THE BRITISH CYNIPOIDEA

(HYMENOPTERA)
DESCRIBED BY P. CAMERON

J. QUINLAN

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 31 No. i

LONDON : 1974

14 AUG1974

THE BRITISH CYNIPOIDEA (HYMENOPTERA)^ 4t ^
DESCRIBED BY P. CAMERON

BY

JOHN QUINLAN

Pp. 1-2 1

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 31 No. i

LONDON : 1974

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 Vol. 31 No. i 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.)

Trustees of the British Museum (Natural History), 1974

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued i August, 1974 Price 1.15

THE BRITISH CYNIPOIDEA (HYMENOPTERA)
' DESCRIBED BY P. CAMERON

By J. QUINLAN

CONTENTS

Page

SYNOPSIS ........... 3

INTRODUCTION .......... 3

ACKNOWLEDGEMENTS ......... 6

CYNIPIDAE ........... 6

Charipinae .......... 6

Cynipinae .......... 9

EUCOILIDAE ........... 10

FlGITIDAE ........... 16

Anacharitinae .......... 16

Aspicerinae .......... 17

Figitinae ........... 17

SUMMARY OF THE PRESENT NAMES DISCUSSED ..... 18

SUMMARY OF THE LECTOTYPES AND NEOTYPES DESIGNATED . . . 18

REFERENCES ........... 19

INDEX ............ 20

SYNOPSIS

The British species of Cynipoidea described by P. Cameron are re-classified. Five new
combinations and seven new specific synonyms are established. Twelve lectotypes and five
neotypes are designated. Nine specific names are at present nomina dubia.

INTRODUCTION

CAMERON described 50 British species of Cynipoidea during the period 1875-89.
Most of these species have been variously referred to by European workers but some
have remained unrecognized. Before further progress could be made with the
preparation of keys to the British Cynipoidea in the Handbooks for the Identification
of British Insects series it became necessary to study the type-material of Cameron's
species and to correlate it with the original descriptions. The type-specimens of
36 of these species have been located in the British Museum (Natural History)
(hereafter abbreviated to BMNH) ; of the remaining 14 species, neotypes have been
designated for five and nine remain nomina dubia.

Three Cameron collections are to be found in the BMNH. One bears the
registration label '1886-3' an d is a collection of galls and gallflies purchased from
Cameron and collected mainly in Europe. The second collection bears the regi-
stration label 'Cameron 1896-76' and was presented to the BMNH by F. D. Godman;
the Museum Register states that this collection consisted of Tenthredinoidea and

4 J. QUINLAN

Cynipoidea, '618 sps., 141 types, 391 microscope slides, 103 larvae and 242 drawings'.
The third collection has the registration label '1914-110'.

In preparing this paper I have contacted the following British and European
Museums in order to ensure, as far as possible, that no type-material of Cameron's
British Cynipoidea has been overlooked: Leicester Museum; Royal Scottish Museum,
Edinburgh; University Museum, Oxford; Termeszettudomanyi Muzeum, Budapest.
Most of these museums have kindly loaned me material that might have contained
the Cameron material for which I was looking. During the course of this study I
have traced where possible the collections of Thomas Richard Billups, the Rev.
Thomas Anself Marshall and Eduard Arthur Fitch, who were contemporaries of
Cameron and in a few instances the actual collectors of species described by him.

Cameron's British Cynipoidea are all mounted on rectangular cards. Many of
them have been remounted by various people on similar rectangular cards, but in
most such cases the original card mount has been left attached to the pin. Most
type-material has been found in that collection bearing the accession label 'Cameron
1896-76', and in many instances the locality has been written on this label.
Cameron's original specific labels are sometimes attached to the specimens. Some
of the rectangular cards on which Cameron mounted his material have a number
in the bottom left-hand corner. In a few cases this number has been crossed out
and another number has been written in the right-hand corner. The number in
the left-hand corner can usually be correlated with the figure number on the plate
in Cameron (1890); occasionally this number seems to be the paragraph number
of the species description in Cameron's monograph (1890). Only when the descrip-
tion, the locality and other data such as Cameron's original determination labels
are present and agree with that published, has credence been given to this number.
Cameron did not indicate which specimen of a series was to be regarded as the type.
Unfortunately, prior to my examination of the Cameron material, the specimens
had been moved in the collection from species to species and had not been given
labels to indicate their original placements. There is usually no way of telling
from Cameron's descriptions how many specimens he had before him when describing
a new species; only very rarely did he state the number of specimens upon which
he based a description. Where only one type-specimen has been found and there
is no evidence that more than one specimen ever existed, I have accepted and
labelled that specimen as the holotype, provided that it agrees with the original
description and bears the same data as that published. Cameron was not always
the collector of the specimens he described and in those cases where another collector
is cited in the descriptions this is indicated in the type-data. All nominal species are
arranged alphabetically under the genus in which they were described and within
their respective subfamilies, together with references to the original description
and other relevant papers in which they have been referred to or discussed, in the
following sequence.

Name; author, date and page reference of the original publication; status and
sex of primary type where known; present lectotype designation (if necessary);
locality of primary type ; type-depository.

BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 5

Number and sex of paralectotypes if such exist, with data and depository
information as for primary types.

Statement on the labelling of the type-material and its condition if damaged.

A statement (headed 'Identity.'} on the generic placement and taxonomic
validity of the name, accompanied when known by similar data to that above
for the names of senior and junior synonyms.

Notes on other aspects of the information supplied are given below.

Lost types. In those cases where the type-material is lost and there is no
evidence on the number of original specimens or their sex, the statement 'Type(s)
(? sex)' is used to show this lack of information.

Locality. Cameron did not always write the locality on his labels. In a number
of instances one, two or three letters only were used to indicate the locality; I have
given these localities in full. I have also added 'GREAT BRITAIN' and the county,
although this information was not published in the original descriptions.

Collectors. Unless specifically stated to the contrary Cameron was the collector.
In those cases where another collector was named and the material has not been
found, it is presumed to have been returned to the collector and has had to be
regarded as lost.

Type-depositories. These are given for the primary types of the synonyms of
Cameron's names and for Cameron's nominal species. The following abbreviations
are used to indicate these depositories.

BMNH British Museum (Natural History), London.

IPK Institut fur Pflanzenschutzforschung Kleinmachnow, Eberswalde.

MNHU Museum fur Naturkunde der Humboldt-Universitat, Berlin.

NM Naturhistorisches Museum, Vienna.

NR Naturhistoriska Riksmuseum, Stockholm.

UM University Museum, Oxford.

UZI Universitetets Zoologiska Institution, Lund.

ZSBS Zoologische Sammlung des Bayerischen Staates, Munich.

Taxonomic arrangement. The family and subfamily arrangement is as given by
Eady & Quinlan (1963 : 7). The number of segments in the antennal club has
been important in the identification of many of the species. It is not clear what
Cameron meant by the term 'club'; I have regarded as club segments those antennal
segments that bear rhinaria. as defined by Richards (1956).

Kleidotoma Westwood (1833 : 494). Cameron first used the name Kleditoma
(i888 : 165) when describing the species nigripes. In subsequent papers, including
his monograph (1890 : 216) in which he describes and keys species in the genus
Kleidotoma Westwood, he consistently spells the generic name as Kleditoma
and refers to Kleditoma Westwood (1833 : 494). In interpreting Cameron's spelling
of Kleidotoma it cannot be demonstrated that he intended to correct the original
name and, therefore, the name Kleditoma is an 'incorrect subsequent spelling' as
defined in Article 33 (b) of the International Code of Zoological Nomenclature.

Neotypes. The neotypes here designated accord with the provisions of Article
75 (a) of the current International Code of Zoological Nomenclature, and this paper

6 J. QUINLAN

is considered to be revisionary work in the terms of the Code and of the proposed
amendments to Article 75 (a) (Bull. zool. Nom. 29(2) : 90, 1972).

ACKNOWLEDGEMENTS

I would like to thank those colleagues in other museums and institutions who
have provided me with types on loan and information on the Cameron collection.
For such assistance I warmly thank the following: Dr R. Danielson (Universitetets
Zoologiska Institution, Lund) ; Dr Max Fischer (Naturhistorisches Museum, Vienna) ;
Dr M. W. R. de V. Graham (University Museum, Oxford) , Dr W. Hellen (Zoological
Museum of the University, Helsinki) ; Dr E. Konigsmann (Museum fur Naturkunde
der Humboldt-Universitat, Berlin) ; Dr F. Kiihlhorn (Zoologische Sammlung des
Bayerischen Staates, Munich), Dr G. Morge (Institut fur Pflanzenschutzforschung
Kleinmachnow, Eberswalde) .

CYNIPIDAE

CHARIPINAE

Allotria ancylocera Cameron, 1886 : 85. Holotype $, GREAT BRITAIN: Scotland, West
Lothian, Carruber Glen, 14. viii (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76 Carruber Glen'. On the
underside of the label is the name 'ancylocera'. A further rectangular card mount has the
name 'Carruber' and 'ancylocera' together with the date '14/8'. A separate label with the
word 'Holotype' printed in the centre has been added together with a determination label
'HOLOTYPE of Allotria ancylocera Cam. det. J. Quinlan. 1973'.

Identity. Syn. n. of Alloxysta victrix (Westwood, 1833 : 495), holotype $, GREAT BRITAIN
(UM, Oxford) [examined]. Hellen (1963 : 16) first established the combination Alloxysta
victrix (Westwood) and it is here confirmed.

Allotria basimacula Cameron, 1886:87. LECTOTYPE $, GREAT BRITAIN: Scotland,
Stirlingshire, Mugdock (BMNH, ex coll. Cameron), here designated.

Paralectotype. i <$, same data as the lectotype (BMNH).

The lectotype and the paralectotype have the BMNH accession label 'Cameron 96-76
Mugdock'. Both specimens have the original rectangular card mount with the name
'Mugdock' and an unintelligible date written on the underside. The card mount attached
to the lectotype has the name 'basimacula' written on it. The paralectotype has a separate
label in Cameron's handwriting 'basimacula'. A further purple-edged circular label with
the printed word 'Lectotype' in the centre has been added to the lectotype. A similar
blue-edged paralectotype label has been added to the paralectotype. Both specimens
have a determination label 'LECTOTYPE' and 'PARALECTOTYPE' respectively 'of Allotria
basimacula Cam. det. J. Quinlan. 1973'.

Identity. Valid species of Alloxysta Forster. Dalla Torre & KiefEer (1910 : 258) first
established the combination of Alloxysta basimacula (Cameron) and it is herein confirmed.
Allotria caledonica Cameron, 1886 : 88. Holotype $, GREAT BRITAIN: Scotland, Stirlingshire,
Mugdock, 27. vii (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76 Mugdock'. On the
underside of the label is the name 'caledonica'. The rectangular card on which the holotype
is mounted has the number '2' in the left-hand corner; this is the figure number on the plate
in Cameron (1890). On the underside of the card mount is the name 'caledonica Cam.'

BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 7

and the date '27/7' and the letters 'Mugd'. A further label 'caledonica' in Cameron's
handwriting is attached to the holotype. A red-edged circular label with the word
'Holotype 1 printed in the centre has been added together with a determination label 'HOLOTYPE
of Allotria caledonica Cam. det. J. Quinlan 1973'. The holotype has only 8 segments of the
right antenna and 10 of the left antenna remaining; all the tarsi are damaged and the wings
are damaged by the glue used for mounting the specimen.

Identity. Valid species of Alloxysta Forster, combination first established by Dalla
Torre & Kieffer (1910 : 256) as Alloxysta caledonica (Cameron) and herein confirmed.
Allotria collina Cameron, 1889 157. Type(s) $, GREAT BRITAIN: Scotland, Stirlingshire,
Mugdock (Cameron Coll.) (lost).

Identity. Unknown, the name remains a nomen dubium. Dalla Torre & Kieffer
(1910 : 280) placed collina in Charips Marshall (1870 : 181), attributing the latter
name to 'Haliday in Marshall' (loc. cit.) as does lonescu (1969 : 237). It seems most probable
from the species description that collina is assignable to Alloxysta Forster but in the absence
of type-material the name must remain a nomen dubium.

Allotria crassa Cameron, 1889:59. LECTOTYPE $, GREAT BRITAIN: Scotland,
Sutherland, Bonar Bridge (BMNH, ex coll. Cameron), here designated.

Paralecto types. GREAT BRITAIN: i $, Scotland, Argyll, Cladich; i $, Scotland,
Dumfries-shire, Dumfries (correctly associated) ; i^ (misassociated), Scotland, Dumfries-shire,
Dumfries.

The lectotype and the paralectotypes have the BMNH accession label 'Cameron 96-76'
together with their respective localities as listed above. The lectotype has the original
rectangular card mount on the underside of which is the locality 'Bonar'. A further
handwritten label 'crassa Cam' is attached, on the underside of which is the name 'scipes
Thorn'. This is deduced as being originally 'fuscipes' Thorn' before being used to write
the name 'crassa Cam' on the other side. A purple-edged circular label with the printed
word 'Lectotype' has been added together with a determination label 'LECTOTYPE of
Allotria crassa Cam. det. J. Quinlan. 1973'. The two correctly associated paralectotypes
both have blue-edged circular paralectotype labels and determination labels 'Allotria crassa
Cam. det. J. Quinlan 1973'. The misassociated paralectotype has a blue-edged circular
paralectotype label and a label in Cameron's handwriting 'Allotria fuscipes' . A further
label 'PARALECTOTYPE of Allotria crassa Cam. wrongly associated det. J. Quinlan 1973'
is attached. This wrongly associated paralectotype is referable to the genus Phaenoglyphis
Forster.

Identity. Valid species of Alloxysta Forster, first established by Dalla Torre & Kieffer
(1910 : 261) and herein confirmed.

Allotria dolichocera Cameron, 1889:56. LECTOTYPE <j>, GREAT BRITAIN: Scotland,
Dumfries-shire, Dumfries (BMNH, ex coll. Cameron).

Paralectotypes. GREAT BRITAIN: i $ (misassociated), Scotland, Lanarkshire, Cadder
Wilderness (BMNH); i $ (misassociated), England, London, Peckham (T. R. Billups)
(BMNH).

The lectotype herein designated and the paralectotypes have the BMNH accession label
'Cameron 96-76' together with their respective localities as listed above. The lectotype
has a label in Cameron's handwriting 'dolichocera', on the other side of this label is the
name 'brevis Thorns' but the top of this name has been cut off. A further rectangular card
on which the lectotype was originally mounted has on the underside the locality 'Dumfries'
and the word 'brevis'. The two misassociated paralectotypes clearly belong in the genus
Alloxysta Forster and do not agree with Cameron's descripton of dolichocera on the relative
lengths of the antennal segments. The paralectotype collected by Billups has both antennae
damaged but the critical segments are still present. A purple-edged circular label with
the word 'Lectotype' has been added, together with a determination label 'LECTOTYPE of
Allotria dolichocera Cam. det. J. Quinlan 1973'.

Identity. Valid species of Phaenoglyphis Forster, new combination Phaenoglyphis dolichocera
(Cameron) comb. n. here established.

8 J. QUINLAN

Allotria maculicollis Cameron, 1886 : 87. Type(s) $, GREAT BRITAIN: Scotland, Kirkcud-
brightshire, New Galloway (Cameron Coll.) (lost).

Identiy. Unknown, the name remains a nomen dubium. The identity cannot be deduced
from Cameron's (1890 : 252) key placement, in which Allotria macrophadna is compared
with maculicollis. The name was applied almost certainly to an Alloxysta species.
Allotria megaptera Cameron, 1889 : 54. Holotype $, GREAT BRITAIN : Scotland,
Lanarkshire, Cadder Wilderness (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76 Cadder Wilderness'. On
the underside of this label is the name 'megaptera'. Two further labels in Cameron's
handwriting are attached to the holotype pin: they read 'Allotria' and 'megaptera Cam'.
On the original rectangular card is the number '7'; this is the figure number on the plate
in Cameron (1890). A further red-edged circular label with the word 'Holotype' printed
in the centre has been attached together with a determination label 'HOLOTYPE of Allotria
megaptera Cam. det. J. Quinlan 1973'. The left fore wing and both hind wings are missing.

Identity. Valid species of Alloxysta Forster, new combination Alloxysta megaptera
(Cameron) comb. n. here established. Cameron first listed megaptera in a key to the
British Allotria without designating it 'sp. nov.' as was usually his custom. The description
in the couplet of the key to Allotria satisfies the provisions of Article u of the International
Code of Zoological Nomeclature. Allotria melanogaster Hartig (1840 : 200), misidentified
by Cameron (1886 : 86), was later described as Allotria megaptera Cameron (1889 : 54).
Allotria mullensis Cameron, 1883 : 366. Holotype $, GREAT BRITAIN: Scotland, Argyll,
Mull (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76 Mull'. On the underside
of this label is the name 'mullensis'. The original card mount has the number 'i' in the left-
hand corner; this is the figure number on the plate in Cameron (1890). A further label
'mullensis Cam', in Cameron's handwriting is attached. A red-edged circular label with
the word 'Holotype' printed in the centre has been added together with a determination
label 'HOLOTYPE of Allotria mullensis Cam. det. J. Quinlan 1973.'

Identity. Valid species of Alloxysta Forster, new combination Alloxysta mullensis
(Cameron) comb. n. here established.

Allotria perplexa Cameron, 1889 : 58. Type(s) $, GREAT BRITAIN: Scotland, Sutherland;
Inverness-shire, Kingussie; Lanarkshire, Clydesdale; Kirkcudbrightshire, New Galloway;
Dumfries-shire, Dumfries (Cameron Coll.) (lost).

Identity. Unknown, the name remains a nomen dubium. Hellen (1963 : 12) placed
perplexa in the genus Alloxysta Forster, but without seeing type-material. In the absence
of type-material the name must remain a nomen dubium.

Allotria piceotnaculata Cameron, 1883 : 367. Holotype $, GREAT BRITAIN: Scotland,
Dumfries-shire, Dumfries, vi (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76 Dumfries', on the underside
of this label is the name 'piceomaculata'. The rectangular card on which the holotype was
originally mounted has the number '3'; this is the figure number on the plate in Cameron
(1890). On the underside of the card mount is the locality 'Dumfries' and the name
'piceomaculata'. The rectangular card mount on which the holotype is mounted also has
the number '3' in the left-hand corner. A further label in Cameron's handwriting
'piceomaculata Cameron' is attached to the holotype. A red -edged circular label with
the word 'Holotype' printed in the centre has been added together with a determination
label 'HOLOTYPE of Allotria piceomaculata Cam. det. J. Quinlan 1973'. The holotype has
only one segment remaining of the left antenna.

Identity. Valid species of Alloxysta Forster, combination first established by Cameron
(1886 : 88) and here confirmed.

Allotria pleuralis Cameron, 1879 : 113. LECTOTYPE $, GREAT BRITAIN: Scotland,
Lanarkshire (BMNH, ex coll. Cameron), here designated.

Paralecto types. GREAT BRITAIN: 2 $, Scotland, Lanarkshire (BMNH, ex coll. Cameron).

The lectotype has the BMNH accession label 'Cameron 96-76'. Both the rectangular

BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 9

card on which it is mounted and the original card mount have the number '8' in the bottom
left-hand corner. This number is the reference to the figure number on the plate in
Cameron (1890). A further purple-edged circular lectotype label has been added together
with a determination label 'LECTOTYPE of Allotria pleuralis Cam. det. J. Quinlan 1973'.
The left antenna has the apical segment missing. The two paralectotypes are labelled
'Cameron 96-76' and carry the label 'pleuralis'. Two further specimens have not been
located. (Five specimens were referred to in the original description, three from Clyde near
Newton and two from Fossil Marsh, all taken in July. One paralectotype has the locality
Fossil Marsh.)

Identity. Syn. n. of Xystus testacea Hartig, 1841 : 352, holotype $, AUSTRIA (ZSBS,
Munich) and new combination Alloxysta testacea (Hartig) comb. n. here established.
Allotria ruficeps Cameron, 1883:365. Type(s) $, GREAT BRITAIN: Scotland, Kirkcud-
brightshire, New Galloway, vi (Cameron Coll.) (lost).

Identity. Unknown, the name remains a nomen dubium. Dalla Torre & Kieffer
(1910 : 283) proposed a replacement name Charips (Charips) cameroni for Allotria ruficeps
Cameron, which was preoccupied by Allotria ruficeps (Zetterstedt, 1883 : 410).
Allotria ruficollis Cameron, 1883 : 385. Holotype $, GREAT BRITAIN: Scotland, Argyll,
Mull, vi (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76 Mull'. The original
rectangular card mount has on the underside the words 'Mull ruficollis'. A further hand-
written label in Cameron's handwriting 'ruficollis Cam', is attached. A red-edged circular
label with the word 'Holotype' printed in the centre has been added together with a
determination label 'HOLOTYPE of Allotria ruficollis Cam. det. J. Quinlan 1973'.

Identity. Syn. n. of Xystus erythrothorax Hartig, 1840 : 200, holotype <$, GERMANY
(ZSBS, Munich). Dalla Torre & Kieffer (1910 : 257) first established the combination
Alloxysta erythrothorax (Hartig) and it is here confirmed.

Allotria salicis Cameron, 1883 : 367. Type(s)? $, GREAT BRITAIN: Scotland, Lanarkshire,
Clydesdale, Kilpatrick Hills (Rev. T. A. Marshall) (lost).

Identity. This nominal species clearly belongs in the genus Phaenoglyphis Forster.

Cameron (1890 : 237) first established salicis as being in Phaenoglyphis but in the absence

of the type(s) or reliably determined material the specific name remains a nomen dubium.

Phaenoglyphis forticornis Cameron, 18886:210. Type(s) $, GREAT BRITAIN: England,

Barnstaple, Lastingham (Rev. T. A. Marshall) (lost).

Identity. Unknown, the name remains a nomen dubium. Dalla Torre & Kieffer
(1910 : 294) and lonescu (1969 : 274) placed forticornis in Phaenoglyphis Forster but without
seeing type-material; the name must remain a nomen dubium.

CYNIPINAE

Aulax graminis Cameron, 1875:322. LECTOTYPE ?, GREAT BRITAIN: Scotland,
Lanarkshire, Glasgow, near Partick (BMNH, ex coll. Cameron), here designated.
Paralectotypes. 3 $, same data as the lectotype (BMNH).

The lectotype and the paralectotypes have the BMNH accession label 'Cameron 96-76'.
The lectotype has a label in Cameron's handwriting 'graminis Cam'. A further purple-edged
circular label with the printed word 'Lectotype' has been added, together with a further
determination label 'LECTOTYPE of Aulax graminis Cam. det. J. Quinlan 1973'. A blue-
edged circular label with the printed word 'Paralectotype' has been added to each
paralectotype.

Identity. Junior synonym of Aulacidea hieracii (Bouche, 1834 : I ^4), holotype $,
GERMANY (DEI, Eberswalde). The synonymy of graminis with hieracii was first established
by Cameron (1893 : 50) and recently confirmed by Eady & Quinlan (1963 : 20), who
inadvertently marked the synonymy as new.

Diastrophus (?) aphidivorus Cameron, 1889 167. Holotype <$, GREAT BRITAIN: England,
PBarnstaple (Rev. T. A. Marshall) (BMNH, ex coll. Cameron).

> J. QUINLAN

The holotype has the BMNH accession label 'Cameron 96-76'. The original retangular
card mount has on the underside the words 'Aphis of Nettle N'. On the upperside of the
card mount can be seen the lighter areas where the holotype and the aphis referred to in
Cameron's description had been mounted. The number '7', badly faded, can be seen that
refers to the plate figure reference in Cameron (1893). A further label 'aphidivorus' in
Cameron's handwriting is attached to the pin. A red-edged circular label with the printed
word 'Holotype' in the centre has been added together with the determination label
'HOLOTYPE of Diastrophus aphidivorus Cam. det. J. Quinlan 1973'. The holotype has
the left hind tibia and tarsus missing.

The biological information in Marshall's handwriting 'Aphis of Nettle N' is suspect.
The 'N' on the label gives rise to doubt on the actual type-locality published by Cameron
as 'Barnstaple'. Morley (1915 : 23), who lists the Rev. T. A Marshall's localities, gives the
letter 'N' as Marshall's abbreviation for Nunton, near Salisbury (Wiltshire).

Identity. Syn. n. of Diastrophus rubi (Bouche, 1834 : l6 3)> holotype $, GERMANY (MNHU,
Berlin) [examined].

Dalla Torre & Kieffer (1910 : 49) listed aphidivorus as a species of Trischiza Forster
(Figitinae) .

EUCOILIDAE

Eucoila fortinervis Cameron, 1889 : 66. Holotype $, GREAT BRITAIN: England, Gloucester
(BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76 Gloster'. The original
rectangular card on which it was mounted has the number '9' on the upper surface and the
locality 'Gloster' on the underside. The number '9' is the reference to the figure number
on the plate in Cameron (1890). The holotype has the label 'fortinervis Cam' in Cameron's
handwriting attached to it. Although Cameron did not describe the female of fortinervis,
pi. 9, fig. 9 in Cameron (1890) of a female suggests that the figure was transposed with that
of another species, most probably proxima. A red-edged circular label with the word
'Holotype' printed in the centre has been attached, together with a determination label
'HOLOTYPE of Eucoila fortinervis Cam. det. J. Quinlan 1973.' The holotype has segments
14-15 of the right-hand antenna missing.

Identity. Valid species of Trybliographa Forster, new combination Trybliographa
fortinervis (Cameron) comb. n. here established.

Eucoila gracilicornis Cameron, i888a : 168. LECTOTYPE $, GREAT BRITAIN: Scotland,
Lanarkshire, Banks of Clyde nr Cambuslang.

Paralectotypes. 2 g, same data as the lectotype (BMNH, ex coll. Cameron).

The lectotype herein designated has the BMNH accession label 'Cameron 96-76 S. BK.
Clyde'. A rectangular label with 'gracilicornis CBL' and the name 'similis' crossed out is
attached. On the underside of this label is the number '4' ; this number seems to be the
paragraph number in Cameron (1890 : 199), in which the species is described. Another
label added at a later stage by G. J. Kerrich has the abbreviation 'Pseudeuc' on it. A
circular purple-edged label with the printed word 'Lectotype' has been added together with
a determination label 'LECTOTYPE of Eucoila gracilicornis Cam. det. J. Quinlan 1973'. Both
paralectotypes have had blue-edged circular paralectotype labels added to them, together
with determination labels.

Identity. This valid species was first placed in Trybliographa Forster by Hellen (1960 : 12).
The placing of gracilicornis in Trybliographa by Hellen is herein confirmed.
Eucoila proxima Cameron, 1889 : 67. LECTOTYPE $, GREAT BRITAIN: England, Essex,
Benfleet (BMNH, ex coll. Cameron), here designated.

Paralectotypes. 2 $, same data as the lectotype except that Benfleet is omitted (BMNH,
ex coll. Cameron).

The lectotype has the BMNH accesssion label 'Cameron 96-76 Benfleet'. A further

BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON II

purple-edged circular label with the printed word 'Lectotype' in the centre has been added
together with a determination label 'LECTOTYPE of Eucoila proximo, Cam. det. J. Quinlan
1973'. One paralectotype has the additional label 'proxima Cam' in Cameron's handwriting
and the number '16/83', together with the number '14' in one corner and the number '8'
in the other. The number '8' is the number of the figure on the plate in Cameron (1890).
This plate and figure shows a male ; Cameron did not describe the male of proxima and this
suggests that the figure was transposed with that of fortinervis Cameron. The other
paralectotype has a label 'proxima Cam', in Cameron's handwriting and the number '15'
on the original card mount.

Identity. Syn. n. of Trybliographa glottiana (Cameron 1883 : 368), holotype $, GREAT
BRITAIN (BMNH, ex coll. Cameron) [examined, see p. 14].

Eucoila scotica Cameron, 1889 : 65. LECTOTYPE $, GREAT BRITAIN : Scotland, Dumfries-
shire, Dumfries (BMNH, ex coll. Cameron), here designated.

Paralectotypes. GREAT BRITAIN: i ^, Scotland, Stirlingshire, Mugdock (BMNH, ex coll.
Cameron) ; i , i $, Scotland, Dumfries-shire, Dumfries (BMNH, ex coll. Cameron) ; I $,
Scotland, Dumfries-shire, Dumfries (BMNH, ex coll. Cameron); i $, Scotland, Kirkcud-
brightshire, Colvend (BMNH, ex coll. Cameron); i $, Scotland, no locality; i <J, Scotland,
West Lothian, Carruber Glen (BMNH, ex coll. Cameron) ; 2 $, Scotland, Ayrshire, Dairy
(BMNH, ex coll. Cameron).

The lectotype has the BMNH accession label 'Cameron 96-76. Dumfries', together
with another rectangular label with the locality and name 'Dumfries scotica'. A further
label in Cameron's handwriting is attached and reads 'scotica Cam'. On the reverse side
of this label is the abbreviation 'clia (new sp. ?) $'. A purple-edged circular label with the
printed word 'Lectotype' has been added, together with a determination label 'LECTOTYPE
of Eucoila scotica Cam. det. J. Quinlan 1973'. The paralectotypes all have the BMNH
accession label 'Cameron 96-76' together with their respective locality names and the deter-
mination label 'scotica'. The paralectotype from Colvend has the name labels 'ciliaris'
and 'scotica'; on the underside of this label is the number '3', which is the figure number
on the plate in Cameron (1890).

Identity. Valid species of Trybliographa Forster, new combination Trybliographa scotica
(Cameron) comb. n. here established.

Kleditotna affinis Cameron, 1889 164. Holotype $, GREAT BRITAIN: Scotland, Sutherland,
Bonar Bridge (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76 Bonar Bridge'. A further
label in Cameron's handwriting is attached and reads 'affinis Bonar'. A circular red-edged
label with the printed word 'Holotype' has been added, together with a determination label
'HOLOTYPE of Kleidotoma affinis Cam. det. J. Quinlan. 1973.'.

Identity. This valid nominal species is clearly assignable to Kleidotoma subgen.
Tetrahoptra Forster (1869 : 342), senior synonym of Tetratoma Cameron (1890 : 223).
Kleditotna caledonica Cameron, i888a : 167. Type(s) $, GREAT BRITAIN: Scotland, Argyll,
Claddich, Loch Awe (Cameron) (lost).

NEOTYPE $, GREAT BRITAIN: England, Buckinghamshire, Greenlands, 3i.vii.i936,
'reared from nest of Heron, No. 2.' (BMNH), here designated. I have been unable to
locate the original female type(s). The neotype here designated has, in addition to the
data above, a red-edged circular label with the printed word 'Neotype' in the centre, an
identification label 'Kleidotoma caledonica Cam. J. F. P. 194?.', and the label 'NEOTYPE of
Kleidotoma caledonica Cam. det. J. Quinlan 1973.'.

Identity. This valid species is clearly assignable to Kleidotoma subgen. Kleidotoma
Westwood. Dalla Torre & Kieffer (1910 : 210), Hellen (1960 : 24) and lonescu (1969 : 139)
have referred to this species and have used the characters that Cameron used in his original
description for identifying it. The BMNH collection contains 2 $, identified and labelled as
caledonica by J. F. Perkins, which agree with the original description; one of these has been
designated neotype. Cameron's description isolates caledonica from other related species
on the antennal club being shorter than the flagellum, the scutellum not projecting at the

12 J. QUINLAN

apex, the legs being black, the radial cell elongate and the joints of the antennal club being
attenuate. Cameron did not know the male of the species. Dalla Torre & Kieffer (1910 : 21 1)
and lonescu (1969 : 139) refer in error to both male and female in Cameron's works
(i888a : 166; 1890 : 224).

Kleditoma crassiclava Cameron, i888a : 166. Holotype $, GREAT BRITAIN: Scotland,
Sutherland, Bonar Bridge (BMNH, ex coll. Cameron) .

The holotype has the BMNH accession label 'P. Cameron Coll. B.M. 1914-110'. The original
rectangular card mount has the name 'crassiclava' on the underside ; the same card mount has
the locality 'Bonar' written across the specific name. Both the species name and the locality
are written in Cameron's handwriting. The holotype has been remounted on a card point.
A red-edged circular label with the printed word 'Holotype' in the centre has been attached,
together with a determination label 'HOLOTYPE of Kleidotoma crassiclava Cam. det. J.
Quinlan. 1973.'.

Identity. This valid species is assignable to the genus Rhynchacis Forster. The
combination with Rhynchacis Forster was first established by Cameron (1890 : 217).
Kleditoma elegans Cameron, 1889 : 60. LECTOTYPE <j>, GREAT BRITAIN : Scotland,
Stirlingshire, Mugdock Wood, nr Glasgow (BMNH, ex coll. Cameron), here designated.

Paralectotype. GREAT BRITAIN: i $ (misassociated), Scotland, Lanarkshire, Clober
Glen, nr Glasgow (BMNH, ex coll. Cameron).

The lectotype has the BMNH accession label '96-76 Mugdock Wood'. The specific
name 'elegans Cam.' in Cameron's handwriting is attached to the lectotype. The original
card mount has the number '9' in the bottom right-hand corner; this is the figure number to
the plate in Cameron (1890). The misassociated paralectotype has a specific label 'elegans?'
in Cameron's handwriting; although similar to elegans, it has the antennal segments
differently shaped and of different comparative lengths. It is Kleidotoma pentatoma
Thomson.

Identity. This valid species is assignable to Kleidotoma subgen. Pentakleidota Weld and
is the type-species of this subgenus.

Kleditoma filicornis Cameron, 1889:62. Type(s) $, GREAT BRITAIN: England, Devon,
Bishopsteignton (Rev. T. A. Marshall) (lost).

NEOTYPE $, GREAT BRITAIN: England, Norfolk, Holkham, I7.vii.i968 (M. Crisp)
(BMNH), here designated.

I have been unable to locate the original female type(s). Various authors, Dalla Torre
& Kieffer (1910), Hellen (1960) and lonescu (1969) have all referred to this species in
published works and have used the characters that Cameron used in his original description
for identifying this species. The neotype here designated is one of 8 females with identical
data; a red-edged circular label with the printed word 'Neotype' in the centre has been
attached, together with a further label 'NEOTYPE of Kleidotoma filicornis Cam. det. J.
Quinlan 1973.'.

Identity. This valid species is assignable to Kleidotoma subgen. Kleidotoma Westwood
and is separated from related species on the long thin antennae, the less abrupt club and the
long narrow radius of the forewing.

Dalla Torre & Kieffer (1910) wrongly cite Cameron (1890) as describing the male.
Kleditoma gracilicornis Cameron, 1889 : 63. Type(s) $, GREAT BRITAIN: England, Wiltshire,
Nunton (publ. as Munton] (Rev. T. A. Marshall) (lost).

Identity. This nominal species clearly belongs in the genus Kleidotoma subgen.
Tetrahoptra Forster, but in the absence of the type(s) or reliably determined material the
specific name remains a nomen dubium.

The male of this species was not described.

Kleditoma longicornis Cameron, 1899 : 62. Type(s) $, GREAT BRITAIN: England, Devon,
Barnstaple (Rev. T. A. Marshall) (lost).

Identity. This nominal species belongs in the genus Kleidotoma subgen. Kleidotoma
Westwood, but in the absence of the type(s) or reliably determined material the specific
name remains a nomen dubium.

BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 13

Kleditoma longipennis Cameron, 1889 : 59. Holotype $, GREAT BRITAIN: Scotland,
Lanarkshire, Clober Moor, nr Glasgow (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76 Clober Moor', on the reverse
side of which is the name 'longipennis'. A further label in Cameron's handwriting 'longipennis
Cam.' and a smaller label with the name 'longipennis' is attached. The original card on
which the holotype was mounted has the data 'Clober Moor'. The rectangular card on
which the holotype is mounted has the number '8' in the bottom left-hand corner with a
line crossing the number out; in the right-hand corner of the card mount is the number
'9'. The latter number is the figure number on the plate in Cameron (1890).

Identity. This valid species belongs in the genus Kleidotoma subgen. Pentakleidota Weld.

The male of this species was not described.

Kleditoma marshalli Cameron, 1889 : 61. Type(s) $ <$, GREAT BRITAIN: England, Devon,
Barnstaple (Rev. T. A. Marshall) (lost).

Identity. This nominal species clearly belongs in the genus Kleidotoma subgen.
Kleidotoma Westwood, but in the absence of the type(s) or reliably determined material the
specific name remains a nomen dubium.

Kleditoma melanopoda Cameron, i888a : 167. Holotype $, GREAT BRITAIN: England,
London District (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76' ; no locality data are attached.
The holotype is mounted on a rectangular card in the bottom left-hand corner of which is
the number '10'; this number is the figure number on the plate in Cameron (1890). A label
'melanopoda' in Cameron's handwriting is attached. A red-edged circular label with the
word 'Holotype' printed in the centre has been attached, together with a determination
label 'HOLOTYPE of Kleidotoma melanopoda Cam. det. J. Quinlan. 1973.'. (Although not
stated in the original description, Cameron (1890 : 231) indicates that the Rev. T. A.
Marshall was the collector of this specimen from the same locality 'London District'.)

Identity. This valid species belongs in the genus Kleidotoma subgen. Arhoptra Kieffer.

The male of this species was not described.

Kleditoma nigripes Cameron, i888a : 165. Type(s) $, GREAT BRITAIN: England, London,
Dulwich (T. R. Billups) (lost).

NEOTYPE , GREAT BRITAIN: England, Hertfordshire, Harpenden, Rothamsted, 1951
(B. R. Laurence), B.M. 1951-450 (BMNH), here designated.

No Cameron material of this species stands in the BMNH collection under this name and all
attempts to trace the type(s) in collections that could have contained T. R. Billups material
have been fruitless. The neotype here designated is one of four females with the above
data in the BMNH collection determined as 'Rhynhacis nigripes (Cam)' by G. J. Kerrich,
1951. It has a red-edged circular label with the word 'Neotype' printed in the centre,
together with a determination label 'NEOTYPE of Kleidotoma nigripes Cam. det. J. Quinlan.
1973-'-

Identity. Valid nominal species of Rhynchacis Forster, combination Rhynchacis nigripes
(Cameron) first established by Cameron (1890 : 218) and here confirmed.

Kleditoma picipes Cameron, 1886 : 92. Holotype $, GREAT BRITAIN: Scotland (BMNH,
ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76'. No data label for the
locality is attached. A label in Cameron's handwriting is attached and reads 'picipes'. The
holotype is mounted on a rectangular card; in the bottom left-hand corner is the number
'10', which is the figure number on the plate in Cameron (1890). The holotype bears a
red-edged circular label with the printed word 'Holotype' in the centre, together with a
determination label 'HOLOTYPE of Kleidotoma picipes Cam. det. J. Quinlan. 1973-'-

Cameron did not give a locality in his original description, although the paper was on the
fauna of Scotland with particular reference to Clydesdale. In a later reference to the species,
Cameron (1890 : 226) gave Clydesdale as the locality for the species.

Identity. This valid species of Kleidotoma Westwood is newly assigned to the subgen.

14 J. QUINLAN

Kleidotomidea Rohwer & Fagan (1917 : 369). In placing picipes I have by examination
counted as club segments of the antennae those segments having rhinaria.

The male of picipes was not described by Cameron.

Kleditoma striata Cameron, 1886 :gi. Holotype <, GREAT BRITAIN: Scotland, Lanarkshire,
Lanark, nr Newton on the Clyde (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76. S. Bk. Clyde'. Two determi-
nation labels in Cameron's handwriting are attached, one has the name 'striata' and the
other has the name 'Kleditoma striata Cam 3'. The number '3' is repeated again on the
rectangular card mount and is the figure number on the plate in Cameron (1890). The
holotype bears a red-edged circular label with the printed word 'Holotype' in the centre.
A further determination label 'HOLOTYPE of Kleidotoma striata Cam. det. J. Quinlan. 1973.'.
has been attached.

Identity. This valid species belongs in Kleidotoma subgen. Pentakleidota Weld.

Cameron did not describe the male of this species although one male from his collection
labelled as striata is in the BMNH collection.

Kleditoma striaticollis Cameron, i888a : 167. Holotype $, GREAT BRITAIN: Scotland,
Kirkcudbrightshire, New Galloway (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76 Galloway' together with
a further label with the locality 'Galloway' and the name 'striaticollis' in Cameron's hand-
writing. The holotype is mounted on a rectangular card; in the bottom left-hand corner
is the number '8'. This number is the paragraph number given to the species in Cameron
(1890 : 222). No figure of the species was given by Cameron. A red-edged circular label
with the printed word 'Holotype' in the centre has been attached to the holotype together
with a determination label 'HOLOTYPE of Kleidotoma striaticollis Cam. det. J. Quinlan.

1973-'-

Identity. This valid species belongs in the genus Kleidotoma subgen. Kleidotoma
Westwood.

The male of striaticollis was not described by Cameron.

Kleditoma truncata Cameron, 1889 : 60. Holotype $, GREAT BRITAIN: Scotland, Renfrew-
shire, Bishopton, 31. vii (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76 Bishopton'. A further
label with the locality 'Bishopton 31/7' is attached to the holotype together with a label
'truncata Cam.' in Cameron's handwriting and another label with 'truncata 2' is also attached
to the holotype. The number '2' on the second name label is the number of the figure on
the plate in Cameron (1890). A red-edged circular label with the printed word 'Holotype'
in the centre has been added, together with a determination label 'HOLOTYPE of Kleidotoma
truncata Cam. det. J. Quinlan. 1973.'.

Identity. This valid species of Kleidotoma Westwood is newly assigned to the subgen.
Pentakleidota Weld (1952 : 225).

Psichacra dalei Cameron 1879 : 115. Holotype $, GREAT BRITAIN: England, Dorset,
Wooton (/. C . Dale] (lost).

NEOTYPE ?, GREAT BRITAIN: England, Monmouth, Monks Wood, 5.x.i878 (BMNH),
here designated. I have been unable to locate the original female holotype of this species.
Cameron (1890 : 198) gives an additional locality to that in his first description: Worcester
(Fletcher}. The neotype here designated, in addition to the data above, has the labels
'Cameron 96-76 Monks Wd.' and a handwritten label 'rufula Forst'. The neotype is one
of three specimens with identical data in the BMNH and is distinguished from them by a
red-edged circular label with the printed word 'Neotype' in the centre. A further label
'NEOTYPE of Psichacra dalei Cam. det. J. Quinlan 1973.' has been added.

Identity. Junior synonym of Psichacra rufula (Forster, 1855 : 257), synonymy first
established by Cameron (1890 : 197) and here confirmed.

Psichacra glottiana Cameron, 1883 : 368. Holotype $, GREAT BRITAIN: Scotland, Lanark
shire, Cambuslang (publ. as Cambusland), along the banks of the Clyde, vii (BMNH, ex
coll. Cameron).

BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 15

The holotype has the BMNH accession label 'Cameron 96-76 South Bank of the Clyde'.
A further rectangular card on which the holotype was originally mounted has the number
'7' in the right-hand corner; this is the number of the figure on the plate in Cameron (1890).
On the underside of this label are the initials 'CBL' and the specific name 'glottiana' in
Cameron's handwriting. A further label in Cameron's writing 'glottiana' is also attached
to the holotype. An additional label is attached to indicate the name under which the
holotype stood in the BMNH collection; it reads 'In BM. coll. under Trybliographa
mandibularis (Zett)'. A red-edged circular label with the printed word 'Holotype' in the
centre has been attached, together with a determination label 'HOLOTYPE of Psichacra
glottiana Cam. det. J. Quinlan. 1973'.

Identity. Valid species of Trybliographa Forster, new combination Trybliographa glottiana
(Cameron) comb. n. here established, and senior synonym of proxima Cameron, 1889 (see
p. 10).

Psichacra tnarshalli Cameron, 1883 : 369. Type(s) <$, GREAT BRITAIN: England, Devon,
Barnstaple (Rev. T. A. Marshall) (lost).

NEOTYPE <J, GREAT BRITAIN: England, Surrey, Boxhill (C. G. Champion] (BMNH,
ex coll. Cameron), here designated.

I have been unable to locate the original male type(s). Dalla Torre & Kieffer (1910) and
Hellen (1960) have referred to this species in published works and have used the characters
that Cameron used in the original description for identifying it. The identity of marshalli
is clear from the neotype here designated. It has a label 'Box Hill 67 74', a handwritten
label 'Gronotoma marshalli', the BMNH accession label 'Cameron 96-76 Box Hill', and a
red-edged label with the printed word 'Neotype' in the centre. A determination label
'NEOTYPE of Psichacra marshalli Cam. det. J. Quinlan 1973,' has also been attached to it.
A further label attached to the neotype reads 'New Genus B of Weld 1952, G. J. Kerrich
det. 1960'.

Identity. This valid species is not assignable to a recognized genus at present. It runs
in the keys to genera of Weld (1952 : 108) to his genus 'B'. It is proposed to publish a
description of this new genus in another publication.

Psichacra sirnilis Cameron, 1883:368. LECTOTYPE <j>, GREAT BRITAIN: Scotland,
Lanarkshire, Cambuslang [publ. as Cambusland along the banks of the Clyde] (BMNH,
ex coll. Cameron), here designated.

Paralectotype. i <$, same data as the lectotype except that the BMNH accession number
is 'Cameron 86-3 South Bank Clyde' and a label 'CBL'. A further label in Cameron's
handwriting 'Eucoela mandibularis Zett' is attached.

Non-syntypic specimens. In the BMNH collection are five Cameron specimens that
agree with the original description but they do not have the lectotype data: i without
data; i $, Clober; i ^, Gloucester; i $, labelled 'N'; and i $, Dairy. Cameron referred
only to 'Cambusland' as the type-locality. All five specimens have been labelled 'These
specimens have no type status'.

The lectotype, which has been remounted on a rectangular card, has the BMNH accession
label 'Cameron 96-76'. The original card mount is attached to the pin; on the underside
of this card mount are the specific names 'similis' and 'mandibularis' written in ink over
the pencilled word 'Cambusland 7/9?'. On the upper surface of this card in the bottom
left-hand corner is the number '5'; this is the figure number on the plate in Cameron
(1890 : 220) to the species Trybliographa mandibularis (Zetterstedt) . A circular purple-
edged label with the printed word 'Lectotype' in the centre has been attached together
with a determination label 'LECTOTYPE of Psichacra similis Cam. det. J. Quinlan. 1973.'-

Identity. Junior synonym of Trybliographa mandibularis (Zetterstedt, 1838 : 410)
(Figites), syntypes $ $, SWEDEN (NR, Stockholm) [examined]. The synonymy of similis
with mandibularis was first established by Cameron (1890 : 200) and is here confirmed after
direct comparison of the lectotype and syntypes.

Trybliographa crassicornis Cameron, 1889 : 64. Holotype <j>, GREAT BRITAIN: Scotland,
Lanarkshire, Cambuslang on the Clyde (BMNH, ex coll. Cameron).

16 J. QUINLAN

The holotype has the BMNH accession label 'Cameron 96-76 South Bk Clyde' ; a further
label in Cameron's handwriting has the following abbreviations 'CBL' and 'crassi'. This
same rectangular label has the number '4' in the bottom left-hand corner; this is the figure
number on the plate in Cameron (1890). A red-edged circular label with the printed word
'Holotype' in the centre has been attached to the holotype, together with the determination
label 'HOLOTYPE of Trybliographa crassicornis Cam. det. J. Quinlan. 1973.'.

Identity. This valid species clearly belongs in the genus Trybliographa Forster and this
combination is herein confirmed.

Trybliographa nigricornis Cameron, 1883:369. LECTOTYPE <j>, GREAT BRITAIN:
Scotland, Lanarkshire, Clober Wood (BMNH, ex coll. Cameron), here designated.

Paralectotype. GREAT BRITAIN: i <$, Scotland, Kirkcudbrightshire, Dairy [labelled Daly],
(BMNH, ex coll. Cameron).

The lectotype has the BMNH accession label 'Cameron 96-76 Clober Wood'. A further
label in Cameron's handwriting is attached to the lectotype and reads 'Clober' and 'nigricornis
Cam'. A circular purple-edged label with the printed word 'Lectotype' in the centre has
been added, together with the determination label 'LECTOTYPE of Trybliographa nigricornis
Cam. det. J. Quinlan. 1973.'.

Identity. Syn. n. of Trybliographa atra (Hartig, 1840 : 201) (Cothonaspis) , Holotype
9, GERMANY (ZSBS, Munich) [examined].

Trybliographa testaceipes Cameron, 1883 : 370. Holotype $, GREAT BRITAIN: Scotland,
Ayrshire, Dairy (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76 Dairy'. The original
rectangular card mount has the name 'testaceipes' on the underside and the number '8' on
the upper surface. This number is the figure number on the plate in Cameron (1890). A
further label 'testaceipes' in Cameron's handwriting is attached. A red-edged circular label
with the printed word 'Holotype' has been attached, together with a determination label
'HOLOTYPE of Trybliographa testaceipes Cam. det. J. Quinlan. 1973.'.

Identity. Valid species of Trybliographa Forster.

FIGITIDAE

ANACHARITINAE

Aegilips bicolorata Cameron, 1887 : 194. Holotype $, GREAT BRITAIN: England [publ.
as probably London district], (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76'. On the reverse side of
this label is the name 'bicolorata'. Two further labels with the name 'bicolorata' in Cameron's
handwriting are attached to the holotype. A label with the number '5' is attached to the
holotype; this is the figure number on the plate in Cameron (1890). The holotype has a
label 'B.M. Type Hym. 7. 39'., together with a determination label 'B.M. Type Hym.
Aegilips bicolorata Cameron. 1887.' Both forewings are damaged at the apices.

Identity. Valid species in the genus Aegilips Walker.

Aegilips ruficornis Cameron, 1883 : 372. Holotype $, GREAT BRITAIN: Scotland, Renfrew-
shire, Bishopton (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76 Bishopton'. On the reverse
side of this label is the name 'ruficornis'.

Further labels are attached and read 'Bishop' and 'ruficornis'; both are in Cameron's
handwriting. A label with the number '8' is also attached; this number is the figure number
on the plate in Cameron (1890). The holotype bears a BMNH Type Hym. 7.40. label. A
further label 'B.M. Type. Hym. Aegilips ruficornis Cameron. 1883' is attached to the holotype.

Identity. Valid species in the genus Aegilips Walker.

BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 17

Aegilips scotica Cameron, 1883 : 372. LECTOTYPE g, GREAT BRITAIN: Scotland,
Inverness-shire, Glenmoriston (BMNH, ex coll. Cameron), here designated.

Paralectotype. i , same data as the holotype.

The lectotype and paralectotype have the BMNH accession label 'Cameron 96-76'. The
lectotype has a label 'scotica' in Cameron's handwriting. A purple-edged circular label
with the printed word 'Lectotype' has been added, together with a determination label
'LECTOTYPE of Aegilips scotica Cam. det. J. Quinlan. 1973.'. The left antenna of the lectotype
is damaged.

Identity. Syn. n. of Xyalaspis abietina (Thomson, 1861 : 412), holotype < [publ. as $],
SWEDEN: Ostergothland (Prof. Zetterstedt] (UZI, Lund) [examined]. (One other male, not a
syntype, stands under abietina in the UZI, Lund.)

Aegilips striolata Cameron, 1883 : 373. Holotype^, GREAT BRITAIN: Scotland, Stirlingshire,
Mugdock near Glasgow (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76' ; on the reverse side of this
label is the locality 'Mugdock'. A further label with the number '3' is attached; this number
is the paragraph number on the appropriate page in Cameron (1890 : 181). An additional
label in Cameron's handwriting 'striolata' is attached. A red-edged circular label with the
printed word 'Holotype' has been attached to the holotype, together with a determination
label 'HOLOTYPE of Aegilips striolata Cam. det. J. Quinlan. 1973'.

Identity. Valid species in the genus Aegilips Walker.

ASPICERINAE

Onychia nigripes Cameron, 1879 : 112. Holotype $, GREAT BRITAIN: England, Suffolk,
Norwich (/. B. Bridgman) (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76'; on the reverse side of
this label is the name 'nigripes'. On the reverse side of the original card mount is the
number '198' or '198'. A further label has the number '6', which is the figure number on
the plate in Cameron (1890). A determination label 'nigripes' in Cameron's handwriting is
attached. No locality label is attached but according to the original description Cameron
only saw one specimen collected by Bridgman. A red-edged circular label with the printed
word 'Holotype' in the centre has been attached, together with a determination label
'HOLOTYPE of Onychia nigripes Cam. det. J. Quinlan. 1973.'.

Identity. Junior synonym of Callaspidia defonscolombei Dahlbom, synonymy first
established by Cameron (1890 : 177) and here confirmed by comparison of types [defonsco-
lombei holotype $, SWEDEN: Furillen vid Gottland (UZI, Lund)]. (This species was first
described by Dahlbom (1842 : 13) as Callaspidia De Fonscolombei. Dalla Torre & Kieffer
(1910 : 64) were the first authors to coalesce the name to defonscolombei, which is correct under
the International Code of Zoological Nomenclature.}

FIGITINAE

Melanips femoralis Cameron, 1883 :37i. Holotype $, GREAT BRITAIN: Scotland, Suther-
land, Bonar Bridge, vi (BMNH, ex coll. Cameron).

The holotype has the BMNH accession label 'Cameron 96-76 Bonar Bridge'. A determi-
nation label 'femoralis' and the data label 'Bonar' in Cameron's handwriting is attached
to the holotype. A red-edged circular label with the printed words 'Type H. T.', together
with a label 'B. M. Hym. 7.41' and a determination label 'B. M. Type Hym. Melanips
femoralis Cameron 1883', are attached to the holotype.

Identity. Valid species of Melanips Giraud.

i8 J. QUINLAN

SUMMARY OF THE PRESENT NAMES DISCUSSED

New synonymy

Aegilips scotica Cameron syn. n. of Xyalaspis abietina (Thomson).
Allotria ancylocera Cameron syn. n. of Alloxysta victrix (Westwood).
Allotria pleuralis Cameron syn. n. of Alloxysta testacea (Hartig).
Allotria ruficollis Cameron syn n. of Alloxysta erythrothorax (Hartig).
Diastrophus aphidivorus Cameron syn. n. of Diastrophus rubi (Bouche).
Eucoila proxima Cameron syn. n. of Trybliographa glottiana (Cameron).
Trybliographa nigricornis Cameron syn. n. of Trybliographa atra (Hartig).

Confirmed synonymy

Aulax graminis Cameron, junior synonym of Aulacidea hieracii (Bouche).

Psichacra similis Cameron, junior synonym of Trybliographa mandibularis (Zetterstedt) .

Onychia nigripes Cameron, junior synonym of Callaspidia defonscolombei Dahlbom.

New combinations

Alloxysta testacea (Hartig) comb. n.
Phaenoglyphis dolichocera (Cameron) comb. n.
Trybliographa fortinervis (Cameron) comb. n.
Trybliographa glottiana (Cameron) comb. n.
Trybliographa scotica (Cameron) comb. n.

Confirmed combinations

Alloxysta basimacula (Cameron).
Alloxysta caledonica (Cameron).
Alloxysta erythrothorax (Hartig).
Alloxysta megaptera (Cameron).
Alloxysta mullensis (Cameron).
Alloxysta piceomaculata (Cameron).
Trybliographa gracilicornis (Cameron) .
Rhynchacis crassiclava (Cameron).

Nomina dubia
Allotria collina Cameron.
Allotria maculicornis Cameron.
Allotria perplexa Cameron.
Allotria ruficeps Cameron.
Allotria salicis Cameron.
Phaenoglyphis forticornis Cameron.
Kleditoma gracilicornis Cameron.
Kleditoma marshalli Cameron.
Kleditoma longicornis Cameron.

SUMMARY OF LECTOTYPES AND NEOTYPES DESIGNATED

Lectotypes

Aegilips scotica Cameron.
Allotria basimacula Cameron.

BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 19

Allotria crassa Cameron.
Allotria dolichocera Cameron.
Allotria pleuralis Cameron.
Aulax graminis Cameron.
Eucoila gracilicornis Cameron.
Eucoila proxima Cameron.
Eucoila scotica Cameron.
Kleditoma elegans Cameron.
Psichacra similis Cameron.
Trybliographa nigricornis Cameron.

Neotypes

Kleditoma caledonica Cameron.
Kleditoma fill cornis Cameron.
Kleditoma nigripes Cameron.
Psichacra dalei Cameron.
Psichacra marshalli Cameron.

REFERENCES

BOUCHE, P. F. 1834. Naturgeschichte der Insekten pp. i-v, 1-216. Berlin.

CAMERON, P. 1875. On some new or little known British Hymenoptera. Proc. nat. Hist.

Soc. Glasgow 2 : 304-324.
1879. On some new or little known British Hymenoptera. Trans, ent. Soc. Lond.

12 : 107-119.
1883. Descriptions of sixteen new species of parasitic Cynipidae, chiefly from Scotland.

Trans, ent. Soc. Lond. 16 : 365-374.
1886. The Fauna of Scotland with special reference to Clydesdale and the Western District.

Hymenoptera, part II. Supplement to Tenthredinidae, pp. 53-95. Glasgow.
1887. Hymenopterological notes. I. On some new or little known British Hymenoptera.

Entomologist's mon. Mag. 23 : 193-195.
i888a. Descriptions of twenty three new species of Hymenoptera. Mem. Proc. Manchr

lit. phil. Soc. (4) 1 : 160-163.
18886. On some new or little known British parasitic Cynipidae. Entomologist's mon.

Mag. 24 : 209-211.
1889. On the British species of Allotrinae, with descriptions of other new species of

parasitic Cynipidae. Mem. Proc. Manchr lit. phil. Soc. (4) 2 : 53-69.

1890. Monograph of British phytophagous Hymenoptera 3 (1889): 1-274. London.

1893. Monograph of British phytophagous Hymenoptera 4 (1892): 1-248. London.

DAHLBOM, G. 1842. Onychia och Callaspidia, tvenne for Skandinaviens Fauna nya Insekt-
Sldgten, horande till Galldple-Steklarnes naturliga grupp. 1-16, 9 figs Lund.

DALLA TORRE, K. W. & KIEFFER, J. J. 1910. Cynipidae. Das Tierreich 24 : 1-891. Berlin.

EADY, R. D. & QUINLAN. J. 1963. Hymenoptera Cynipoidea. Key to families and sub-
families, and Cynipinae (including galls). Handbk I dent. Br. Insects 8 (la) : 1-81.

FORSTER; A. 1869. Ueber die Gallwespen. Verh. zool.-bot. Ges. Wien 19 : 327-370.

GIRAUD, J. 1856. Observations sur quelques especes d'Hymenopte'res rares ou peu connues,
trouvees dans les environs de Vienne. Verh. zool.-bot. Ver. Wien 6 : 179-188.

1860. Enumeration des Figitides de 1'Autriche. Verh. zool.-bot. Ges. Wien. 10 : 23-176.

HARTIG, T. 1840. Ueber die Familie der Gallwespen. Z. Ent. (Germar) 2 : 176-209.

1841. Erster Nachtrag zur Naturgeschichte der Gallwespen. Z. Ent. (Germar) 3 : 322-358.

1843. Zweiter Nachtrag zur Naturgeschichte der Gallwespen. Z. Ent. (Germar) 4 : 395~

422.

20 J. QUINLAN

HELLEN, W. 1960. Die Eucoilinen Finnlands (Hym: Cyn). Fauna fenn. 9: 1-29.

1963. Die Alloxystinen Finnlands (Hymenoptera, Cynipidae). Fauna fenn. 15 : 1-23.

IONESCU, M. A. 1969. Hymenoptera Cynipoidea. Fam. Ibaliidae, subfam. Ibaliinae, Fam.

Figitidae Aspicerinae, Anacharitinae. Figitinae. Fam. Cynipidae subfam. Eucoilinae,

Charipinae. [In Romanian.] Fauna Repub. pop. rom. (Insecta) 9 (6) : 1-285, 57 n g s . 22 pis.

8 maps.
KERRICH, G. J. & QUINLAN, J. 1960. Studies on Eucoiline Cynipoidea (Hym). Opusc.

ent. 25 : 179-196.

KLOET, G. S. & HINCKS, W. D. 1945. A check list of British insects pp. lix, 483. Stockport.
MARSHALL, T. A. 1870. On some British Cynipidae. Entomologist's mon. Mag. 6 : 178-181.
MORLEY, C. 1915. The Rev. T. A. Marshall's localities. Entomologist 48 : 23-24.
RICHARDS, O. W. 1956. Hymenoptera, Introduction and Key to Families. Handbk Ident.

Br. Insects 6 (i) : 1-38.
ROHWER, S. A. & FAGAN, M. 1917. The type species of the genera of Cynipoidea, or the gall

wasps and the parasitic Cynipoids. Proc. U. S. natn. Mus. 53 : 357-380.
THOMSON, C. G. 1862. Forsok till uppstallning och beskrifning af Sveriges Figiter. Ofvers.
K. VetenskAkad. Fork. Stockh. 1861 : 395-420.

1877. Ofversigt af Sveriges Oym/>s-Arter. Opusc. ent. 8 : 778-820.

WELD, L. H. 1952. Cynipoidea (Hym.) 1905-7950, pp. 351. Ann Arbor, Michigan.

[Privately published.]
WESTWOOD, J. O. 1833. Notice of the habits of a Cynipideous insect, parasitic upon the

Rose Louse (Aphis rosae), with descriptions of several other parasitic Hymenoptera.

Mag. nat. Hist. 6 : 491-497.
ZETTERSTEDT, J. W. 1838. Insecta Lapponica 1 : 1-1139. Lipsiae.

INDEX

Aegilips, 1 6, 17 forticornis, 9

affinis, ii fortinervis, 10

Allotria, 6, 7, 8, 9

ancylocera, 6 glottiana, 14

aphidivorus, 9 gracilicornis, Eucoila, 10

Aulax, 9 gracilicornis, Kleditoma, 12

graminis, 9
bicolorata, 16

basimacula, 6 Kleditoma, n. 12, 13, 14

caledonica, Allotria, 6

caledonica, Kleditoma, n longicorms, 12

collina, 7 longipenms, 13

crassa, 7

crassiclava, 12 maculicollis, 18

crassicornis, 15 marshalli, Kleditoma, 13

marshalli, Psichacra, 15

dalei, 14 megaptera, 8

dolichocera, 7 Melanips, 17

melanopoda, 13

elegans, 12 mullensis, 8

Eucoila, 10, ii

nigricornis, 16

femoralis, 17 nigripes, Kleditoma, 13

filicornis, 12 nigripes, Onychia, 17

BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 21

Onychia, 17 salicis, 9

scotica, Aegilips, 17

perplexa, 8 scotica, Eucoila, n

piceomaculata, 8 similis, 15

picipes, 13 stria ta, 14

pleuralis, 8 striaticollis, 14

proxima, 10 striolata, 17
Psichacra, 14, 15

ruficeps, 9 testaceipes, 16

ruficollis, 9 truncata, 14

ruficornis, 16 Trybliographa, 15, 1 6

J. QUINLAN

Department of Entomology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON SW7 5BD

ENTOMOLOGY SUPPLEMENTS

3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera) . Pp. 177:
18 plates, 270 text-figures. August, 1965. 4.20.

4. 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.

6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso-
philidae. Pp. 129: 328 text-figures. May, 1966. 3.

7. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967.

3.15.

8. 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.

9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species
(Lepidoptera: Rhopalocera) . Pp. 509. 8.50. Reprinted 1972.

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho-
palocera). Pp. 322: 348 text-figures. August, 1967. 8.

11. MOUND, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 172:
82 text-figures. May, 1968. 4.

12. 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.

13. 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.

14. CROSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and
its Islands. Pp. 198: i plate, 331 text-figures. July, 1969. 4.75.

15. ELIOT, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera:
Nymphalidae). Pp. 155: 3 plates, 101 text-figures. September, 1969. 4.

16. GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe
(Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969.

19-

17. WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198:

68 plates, 15 text-figures. October, 1971. 12.

18. SANDS, W. A. The Soldierless Termites of Africa (Isoptera Termitidae).
Pp. 244: 9 plates, 661 text-figures. July, 1972. 9.90.

19. 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.

20. VON HAYEK, C. M. F. A Reclassification of the Subfamily Agrypninae
(Coleoptera: Elateridae). Pp. 309: 17 text-figures. October, 1973. 12.30.

21. CROSSKEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia,
including keys to the supraspecific taxa and taxonomic and host catalogues.
Pp. 221: 95 text-figures. December, 1973. 9.55.

PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS, oto WOKING, SURREY, ENGLAND

-8 NOV1974

A CATALOGUE
OF THE TYPE-SPECIMENS OF

THE CETONIINAE

(COLEOPTERA : SCARABAEIDAE)

DESCRIBED BY G. J. ARROW

WITH A COMPLETE BIBLIOGRAPHY OF HIS
ENTOMOLOGICAL WORKS

M. E. BACCHUS

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 31 No. 2

LONDON : 1974

\

r<^
-8 NOV1974

A CATALOGUE OF

THE TYPE-SPECIMENS OF THE CETONIINAE

(COLEOPTERA : SCARABAEIDAE)

DESCRIBED BY G. J. ARROW

WITH A COMPLETE BIBLIOGRAPHY OF HIS
ENTOMOLOGICAL WORKS

BY

MICHAEL EDWARD BACCHUS

Pp. 23-44

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 31 No. 2

LONDON : 1974

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 Vol. 31 No. 2 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.)

Trustees of the British Museum (Natural History), 1974

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 30 October, 1974 Price 1.35

A CATALOGUE OF

THE TYPE-SPECIMENS OF THE CETONIINAE

(COLEOPTERA : SCARABAEIDAE)

DESCRIBED BY G. J. ARROW

By M. E. BACCHUS

CONTENTS

Page

SYNOPSIS ........... 25

INTRODUCTION .......... 25

ABBREVIATIONS .......... 26

ACKNOWLEDGEMENTS ......... 27

CATALOGUE OF THE TYPE-SPECIMENS OF THE CETONIINAE DESCRIBED

BY G. J. ARROW .......... 27

BIBLIOGRAPHY .......... 36

SYNOPSIS

A list is given of all the traceable type-specimens of the species and varieties of Cetoniinae
described by Gilbert J. Arrow. The current valid name or current generic assignment is given
where appropriate. Lectotypes are designated where necessary. A complete bibliography
of Arrow's entomological works is included.

INTRODUCTION

FROM 1896 to 1948 Gilbert J. Arrow was employed at the British Museum (Natural
History) and, during that time, described about 1800 species of Coleoptera in 24
families, mostly from material in the museum collection. He was well aware of the
importance of types and always labelled one specimen (occasionally two) from a
series as 'type'. However, the vague or ambiguous nature of his published type
designations, when any were made at all, renders most of these 'types', except the
uniques, unacceptable as holotypes by today's standards. The remainder of the
series, which he called 'co-types', were rarely all labelled as such. His normal
practice was to label very few, if any, of these additional specimens, whether from
the collection in the British Museum (National History) or borrowed by him from
elsewhere.

In the course of evaluating the type status of Arrow material for routine curation
of the collection and for an increasing number of inquiries during the past few years,

26 M. E. BACCHUS

it was found that the data quoted in the descriptions are rarely comprehensive.
The localities given are often a precis of the data labels of the series. Instances
have been noticed of published data, especially bionomic notes, apparently from
oral sources or contemporary correspondence, not appearing on the data labels.
The name of the 'collector' may be that of the donor of the collection rather than
that of the actual collector(s). The number and sex of the specimens is usually
omitted. These circumstances, together with the fact that a considerable number
of syntypes and paratypes must have been exchanged or given away without
record, make it impossible in many cases to be certain that the whole of the series
is accounted for.

Without access to the collection, museum registers of donations, various journals
and lists available only in the museum libraries, and some knowledge of the history
of the collection and of the vagaries of Arrow's method, accurate and consistent
collation of his type-series is virtually impossible.

This paper is the first of a series which will catalogue all the traceable type-
specimens of species described by Gilbert Arrow. In those families which are
within the specialist field of the cataloguer, lectotypes will be designated where
necessary; in others the syntype series will be listed.

All specimens are in the British Museum (Natural History) except where otherwise
stated. The original genus of every species is given in parenthesis after the specific
name, and the current valid name, when it differs from the original, is given at the
end of the entry. The number and sex of the original series is given, in square
brackets, whenever possible. The data quoted are not verbatim but have been
rationalized for the sake of clarity, brevity and convenience to users. Where
possible the present name of the country of origin is used and all available data
arranged in a standard form. The museum registration numbers have been omitted
as they are of no use without the relevant registers.

The specimens labelled by Arrow as 'type' have, until now, all been regarded as
holotypes and all bore the round, red-bordered type labels in use in this museum.
Where lectotypes are necessary they will, except where otherwise stated, be these
specimens. Normally when both male and female 'types' were so-labelled by
Arrow the male will be designated lectotype.

Every specimen catalogued in this series of papers will have a round label edged
in red (holotype), yellow (paratype), purple (lectotype) or blue (paralectotype or
syntype) as appropriate and a label with the name of the taxon written by the
cataloguer and with his name printed below.

The bibliography of this part of the catalogue is in the form of a complete list of
the entomological works of Gilbert Arrow. Future parts will have a bibliography
pertaining to that part only.

ABBREVIATIONS

BMNH British Museum (Natural History), London.

IRSNB Institut Royal des Sciences Naturelles de Belgique, Brussels.

MCSN Museo Civico di Storia Naturale, Genoa.

CETONIINAE DESCRIBED BY G. J. ARROW 27

MNHN Museum National d'Histoire Naturelle, Paris.

NR Naturhistoriska Riksmuseum, Stockholm.

RHN Rijksmuseum van Natuurlijke Historic, Leiden.

UM University Museum, Oxford.

ZSI Zoological Survey of India, Calcutta.

ACKNOWLEDGEMENTS

I am indebted to Dr R. Damoiseau, Dr J. Krikken, Dr P. I. Persson, Mons. G.
Ruter, Dr T. Sen Gupta, Mr E. Taylor and Dr E. Tortonese for data from their
collections or the loan of specimens. I also thank Mr R. D. Pope for much helpful
advice and Miss V. I. Dick for preparing the index of species described by Gilbert
Arrow on which this catalogue is based.

CATALOGUE OF THE TYPE-SPECIMENS OF THE
CETONIINAE DESCRIBED BY G. J. ARROW

acutangula (Charadronota), 19226 : 529. LECTOTYPE^, CAMEROUN: Dendeng, 16.^.1914;
here designated.

Paralectotypes. UGANDA: i , Mabira Forest, Chagwe, 3500-3800 ft, 1 6-25. vii. 1911
(S. A. Neave); 2 $, Budongo Forest, Unyoro, 3400 ft, 11-15. xii. 1911 ($ A. Neave).

acutipes (Coenochilus), igioc : 210. Holotype $, INDIA: Bombay, Igatpuri, ao.vi.i9O4
(H. M. Lefroy) [Elytra and scutellum on separate card, pygidium detached and mounted on
same card as remainder of specimen.]

angustata (Coilodera), ig^6c : 144 [original series: 2]. LECTOTYPE <$, SUMATRA (/. R.
Grey) ; here designated.

Paralectotype. SUMATRA: I $, Sungei Kumbang, Korinchi, 4500 ft, iv. 1914.

auritus (Mycteristes), igioc : 39. Holotype <$, INDIA: Nilgiri Hills (H. L. Andrewes).
Paratype. INDIA: i <$, same data as holotype.

aurora (Glycyphana), 1941^:84. LECTOTYPE <$, SOLOMON ISLANDS: [Florida Island],
Tulagi, Antigone [sic] flowers, 5.x. 1934 (H- T. Pagderi); here designated.

Paralectotypes. SOLOMON ISLANDS: 5 $, i $, same data as lectotype; 2 <$, same data as
lectotype but i6.ix.i934; 2 , Tulagi, Antigonum [sic], I2.viii.i933 (H. T. Pagderi); 2 <J, I $,
Tulagi, 26.i.i934 (H. T. Pagderi); i <, i $, Tulagi Ridge, 13^.1934 (H. T. Pagderi); I <j>,
Tulagi, 22.vii.i934 (R. A. Lever).

aurocincta (Glycyphana), igioc : 122. Holotype^, BHUTAN: 1898 (L. Durel).

Paratypes. BHUTAN: 2 <$ [not $ as stated by Arrow], i $, same data as holotype (MNHN,
Paris).

mini lent a (Gylcyphana horsfieldi var.^), igioc : 121. LECTOTYPE $, INDIA; here desig-
nated.

[No indication of number of original specimens; one only found.]

bhutanus (Oreoderus), igioc : 225. LECTOTYPE <J, BHUTAN: 1898 (L. Durel); here
designated.

Paralectotypes. BHUTAN: 5 < (BMNH, London), 4 ^ (MNHN, Paris), same data as
lectotype; 2 <J, 2 $ (BMNH, London), 5 <$, 2 $ (MNHN, Paris), Maria Basti (L. Durel).
*

28 M. E. BACCHUS

bidentipes (Protaetia), igoyc : 351 [original series: i $, 2 $]. LECTOTYPE $, NICOBAR Is.
(G. Rogers) ; here designated.

Paralectotypes. i $, same data as lectotype; i <, Rangoon, Andaman Is. [sic] (ZSI,
Calcutta) [Arrow says that this specimen is no doubt wrongly labelled].

binghami (Protaetia), igioc : 156. Holotype $, BURMA: Tenasserim.

Paratype. BURMA: i $, Tenasserim (Captain Bingham) (RNH, Leiden).

biplagiata (Glycosia), igojc : 351. Holotype $, ANDAMAN Is. (Roepstorff).

Paratype. i , Rangoon [sic] [Arrow says that this specimen is 'labelled "Rangoon",
probably by mistake'].

bryanti (Oreovalgus), 1944/1 228. Holotype^, SARAWAK: Quop, 11.^.1914 (G. E. Bryant).
bufo (Anthracophora), igojc : 353. Holotype <$, INDIA: Sylhet (Bowring).

bufo (Charitovalgus), IQ44/ : 231. Holotype $, LAOS: Taevieng, i^.vii.igig (R. Vitalis de

Salvaza) .

bufo (Macronota), igioc : 54. LECTOTYPE $, INDIA: Travancore Tea Co. (G. S. Imray);
here designated.

carbonarius (Dasyvalgus), igioc : 239. LECTOTYPE $, INDIA: Kurseong ( Verschraeghen) ;
here designated.

Paralectotypes. INDIA: i $, same data as lectotype. BURMA: i <$ (BMNH, London),
i <$ (RNH, Leiden), 3 (MNHN, Paris), Ruby Mines, 1200-2300 m, 1890 (Doherty).

[The single female Burma: Ruby Mines (Doherty) described by Arrow is not designated
as a paralectotype because Arrow is indefinite about the association with the males.]

[The collection of Baron P. de Moffarts, which contains more syntypes, has not been
traced.]

cataphracta (Protaetia), 1913^ : 405 [original series: 12]. LECTOTYPE^, JAPAN: Kuri-
gahara, Usui Pass, in flowers, 5.viii.i88i (G. Lewis); here designated.

Paralectotypes. JAPAN: 3 $, Junsai Lake (G. Lewis); 2 $, Kashiwagi, 15-24^1.1881
(G. Lewis); i $, Nikko (G. Lewis); i $, 'Japan'.

catena (Glycyphana), igioc : 122 [original series : 3^]. Holotype^, INDIA: Darjeeling.
Paratype. SIKKIM: i $ (ZSI, Calcutta).

[One other male paratype should be in the Oberthiir Collection (MNHN, Paris) but could
not be found in a recent search by Mons. G. Ruter.]

caudata( Protaetia), igioc : 147. LECTOTYPE , SIKKIM : Kurseong, 1894 ( R - p - Bretandeau) ;
here designated.

Paralectotypes. SIKKIM: 2 $, environs de Kurseong (R. P. Bretandeau) (MNHN, Paris).
[No specimens from Bhutan: Maria Basti, or India: Darjeeling have been found.]

clypealis (Oreoderus), 19447:242. Holotype <$, BURMA: Sadon, 1200 m, 28.vi-5.vii. (R.

Malaise).

cobaltinus (Leucocelis), igoga : 521. LECTOTYPE , RHODESIA: 'Gazaland', Chirinda
Forest, iii.igoy (D. Odendaal); here designated.

Paralectotypes. RHODESIA: 2 $, 'Mashonaland', Chirinda Forest, iii.igoj (G. A. K.
Marshall); i $, 'Gazaland', nr Chirinda Forest, iii.igoj (G. A. K. Marshall).

[By modern definition, Chirinda Forest is not in Gazaland or Mashonaland. It is no miles
south of Umtali.]

coorgica (Macronota), ig^ic : 82. LECTOTYPE <$, INDIA: North Coorg, Santikoppa, 4-
io.v.i9i4 (Fletcher); here designated.

Paralectotypes. INDIA: i <$, North Coorg, Sonawarpet, g.vii.igi^ (L. Newcome); i <$,
Coorg, Sidapur, 14.^.1917 (T. R. N.); i $, Coorg, Sidapur, 3000 ft, 28.iv.i9i7 (T. R. N.).

CETONIINAE DESCRIBED BY G. J. ARROW 29

coriaceus (Dasyvalgus), IQ44/ : 236. Holotype $, MALAYA: Pahang, GunongTahan, Padang,
5500 ft, u.xii.i922 (H. M. Pendlebury).

Paratypes. MALAYA: i <$, Pahang, Gunong Tahan, Seat Point, 5460 ft, 27.xii.ig22 (H. M.
Pendlebury}; i $, as above, except 2o.xii.i922; 3 <, Perak Mts (Doherty).

crassipes (Genuchus), 19266 : 650. Holotype $, KENYA: Maramas District, 14 mis E. of
Mumias, Ilala, 4500 ft, i8-2i.vi.i9ii (S. A. Neave).

crassipes (Oreoderus), 19446 : 243. Holotype $, INDIA: Dehra Dun, Karwapan, I4.xii.i9io
(F. Z.).

croesus (Charitovalgus), 1944 / : 230. Holotype <$, LAOS: Haut Mekong, Muong Sing,
i8.iv.igi8 (R. V. de Salvaza).

dimorpha (Clinteria), igi6c : 4gy [original series: 3 $, 2 $ (see below)]. LECTOTYPE <$,
MOLUCCAS: Larat Island, xii.igoy (F. Muir); here designated.

Paralectotype. MOLUCCAS: I $, Larat (F. Muir).

[The 2 $ specimens mentioned by Arrow are excluded from the paralectotype series because
it is definitely stated that their association with the males is uncertain. There is one female
only, with same data as paralectotype, in BMNH, London.]

dispar (Anagnathocera), ig22b : 525. LECTOTYPE,^, RHODESIA: 'Mashonaland', Chirinda
Forest, x. igo5 (G. A. K. Marshall); here designated.

Paralecto types. RHODESIA: i <$, 'Gazaland', Chirinda Forest, x. igo5 (G. A. K. Marshall);
i $, 'Gazaland' (D. Odendaal).

[By modern definition, Chirinda Forest is not in Gazaland or Mashonaland. It is no miles
south of Umtali.]

dispar (Heterorrhina), igojc : 347 [original series : i $, i $]. LECTOTYPE <$, INDIA : 'N. Ind.' ;
here designated.

Paralectotype. i $, same data as lectotype.

disparilis (Glycyphana) , igi6c : 497 [original series: 6<^, 4$]. LECTOTYPE^, MOLUCCAS:
Larat Island, xii.igo7 (F. Muir); here designated.

Paralectotypes. MOLUCCAS : 2 $, same data as lectotype ; 2 <$, 3 $, same data as lectotype
but no date.

Current valid name: Glycyphana maculiceps Moser (Arrow, ig4i6 : 79).

elgonensis (Endoxazus), ig4ic : 86 [original series: 3]. LECTOTYPE , KENYA: Mt Elgon,
ii 550 ft, ii. ig35 (F. W. Edwards); here designated.
Paralectotypes. KENYA: 2 , same data as lectotype.
Current valid name: Glaphyronyx bayeri Moser (Burgeon, ig46, Revue fr. Ent. 13 : in).

exitnia (Charadronota), 19226:528 [original series: i <, i $]. LECTOTYPE^, UGANDA:
Entebbe, 2 2. ii. 1914; here designated.

Paralectotype. UGANDA: i $, Tero, ig.iv.ign.

exitnia (Macronota), ig^ic : So. LECTOTYPE <, MALAYA: Pahang, Cameron Highlands,
6040 ft, 8.v.ig3g (G. Berumbari); here designated.

Paralectotypes. MALAYA: 2 $, 2 $, same data as lectotype but 6036 ft, 2i.v.ig3g; i <J,
Pahang, Cameron Highlands, Rhododendron Hill, 5200 ft, 2i.vi.ig33.

felix (Clinteria klugi var.), igioc : 187 [original series: 3]. LECTOTYPE < INDIA (Colonel

Buckley) ; here designated.

Paralectotypes. INDIA: 2 $, same data as lectotype.

felix (Genuchus), 19266 : 651. Holotype $, UGANDA: Tero Forest, 4.vii.igi2 (C. C. Gowdey).

fenestrata (Coilodera), ig^6c : 143 [original series: 4]. LECTOTYPE , SUMATRA: Medan;
here designated.

3 o M. E. BACCHUS

Paralecto types. SUMATRA: 2 <$, same data as lectotype. MALAYA: i $, Selangor (H. C.
Robinson) .

festivus (Trichius), igioc : 252. LECTOTYPE $, BURMA: Ruby Mines (Doherty); here
designated. [Although Arrow described the male only and said that the female was unknown,
this specimen bears Arrow's 'Type' label with $ crossed out. It also bears a printed $ label.
No other specimens found.]

festivus Arrow var. funebris (Trichius), see funebris.

flavipennis (Microvalgus), 1944/1 245 [original series: 2 <]. LECTOTYPE <$, AUSTRALIA:
New South Wales, North Sydney, i.xii.igoS (G. E. Bryant); here designated.

Paralecto type. AUSTRALIA: i <$, New South Wales, Jervis Bay, on flowers of Leptosper-
mum scoparium, xi.i924 (F. A. Rodway).

fulvicauda (Dasyvalgus), igioc : 242 [original series: 5 <]. LECTOTYPE ^ BURMA: Karen
Mts (Doherty) ; here designated.

Paralectotypes. BURMA: 4 <$, same data as lectotype.

funebris (Trichius festivus var.J, igioc : 253. No specimens found.

gestroi (Mycteristes), igioc : 38. Holotype <, BURMA: Carin Cheba, 900-1100 m, xii.i888
(L. Fea) (MCSN, Genoa).

Paratype. BURMA: i $, same data as holotype (MCSN, Genoa).

gracilis (Heterorrhina) , 19100 : 96. LECTOTYPE < INDIA: Nilgiri Hills (H. L. Andrewes);
here designated.

Paralectotypes. INDIA: 2 ty, same data as lectotype.

gracilis (Macronota), 1907^:350. LECTOTYPE $, INDIA: Assam, Nagas (Doherty); here
designated.

Paralectotype. INDIA: i $, same data as lectotype.

gravis (Oreoderus), igioc : 228. LECTOTYPE <J, INDIA: Nilgiri Hills (H. L. Andrewes);
here designated.

Paralectotypes. INDIA: 3 <$, i $, same data as lectotype; i <$, Nilgiri Hills, O. Valley,
3500 ft, n.vii. (H. L. Andrewes); i <J, 'S. Ind.'; i . Travancore State, Madras, Pirmaid,
936'N. 77E., 1900-1902 (Mrs Imray) (UM, Oxford).

griseus (Podovalgus), igioc : 230. LECTOTYPE <J, INDIA: Barway (P. Cardon) (IRSNB,
Brussels) ; here designated.

Paralectotypes. INDIA: 4 $, 3 $ (IRSNB, Brussels), i $ (RHN, Leiden), 2 $, i $ (BMNH,
London), same data as lectotype; i $, 3 $, Bengal (IRSNB, Brussels).

[Although Arrow states that the type of this species is in the BMNH, London, the specimens
labelled 'Type ' and 'Type ^' by Arrow are both in IRSNB, Brussels, the male being the
lectotype.]

hondana (Protaetia), igi^d : 404. Holotype ^, JAPAN: Kobe, Harada, 8.vii.i9i3.

Paratypes. Neither the monstrous specimen from Tanegashima (BMNH, London) or
the example from Hiroshima (RHN, Leiden) could be found.

Current valid name: Protaetia lenzi (Harold) (Arrow, 19140 : 79).

horsfieldi (Hope) var. aurulenta (Glycyphana), see aurulenta.

hystrix (Dasyvalgus), igioc : 241. LECTOTYPE ^, INDIA: Assam, Patkai Mts (Doherty);
here designated.

Paralectotypes. INDIA: 4 $, same data as lectotype.

ichthyurus (Leucocelis), igoga : 521. LECTOTYPE^, RHODESIA: 'Mashonaland', Chirinda
Forest, 22-24.11.1907 (C. F. M. Swynnerton); here designated.

CETONIINAE DESCRIBED BY G. J. ARROW 31

Paralectotypes. RHODESIA: 6 <$, 5 $, same data as lectotype; g $, i $, 'Gazaland', Chirinda
Forest, iii.igoy (D. Odendaal}; 2 <$, i $, as previous, but no date; 5 ^, i $, 'Gazaland', nr
Chirinda Forest, iii. 1907 (G.A.K. Marshall) ;2$, i $, as previous, but no date ; 3 , 'Gazaland'
Mt Chirindi, xii.igoi (G. Marshall); i $, 'Gazaland', Mt Chirindi, 1-3.1.1907
(D. Odendaal); i $, 'Mashonaland', Mt Chirinda, xii.igoi (G. A. K. Marshall); i $, Chirinda
(C. F. M. Swynnerton); i $, 'Mashonaland', Chirinda Forest, 3800 ft, 22.11.1907 (C. F. M.
Swynnerton); I <, Mashonaland, Salisbury (G. A. K. Marshall).

[By modern definition, Chirinda Forest is not in Gazaland or Mashonaland. It is 1 10 miles
south of Umtali.]

inscriptus (Trichius), 19380 : 3 [original series: 2 <$, i $]. Holotype $, BURMA: Seinkhu
Valley, Wang, 285'N. 9735' E., 5000 ft, 19^.1926 (F. Kingdon Ward).

Paratypes. BURMA: i , same data as holotype; i $, Kambaiti, 7000 ft, 4-8/^1.1934
(R. Malaise) (NR, Stockholm).

insularis (Dasyvalgus), igioc : 240. LECTOTYPE $, NICOBAR ISLANDS: (Roepstorff);
here designated.

Paralectotype. i $, ANDAMAN ISLANDS (Captain Wimberley).

jansoni (Macronota), igioc : 64. LECTOTYPE $, SIKKIM; here designated.
Paralectotypes. INDIA: 1^,1$, Khasis [Hills], v. 1896 (MNHN, Paris).
[The male specimen in MNHN, Paris has not been designated lectotype because the female
in BMNH, London has been labelled as, and used as, the holotype since 1910.]

jansoni (Platysodes), igioc : 200. Holotype <$, INDIA: Assam, Khasi Hills, 1899 (native
collected) .

kanarensis (Dasyvalgus), igioc : 245. LECTOTYPE^, INDIA: Kanara; here designated.
Paralectotypes. INDIA: 4 <$, same data as lectotype.

kenyensis (Endoxazus), ig^ic : 87 [original series: 4]. Holotype $, KENYA: Mt Kenya,
8500-9700 ft, 23. iv. 1935 (G. L. R. Hancock).

Paratypes. KENYA: 3 $, same data as holotype.

Current valid name: Glaphyronyx kenyensis (Arrow) (Burgeon, 1946, Revue fr. Ent. 13 : in).

kinabaluana (Macronota), 1932** : 127. LECTOTYPE , BORNEO: Mt Kinabalu, Lumu
Lumu, 5500 ft, i6.iv.i929 (H. M. Pendlebury); here designated.

Paralectotypes. BORNEO: i $, same data as lectotype; i $, same data as lectotype, but
9.^.1929; i ^, same data as lectotype, but 7.^.1929.

klugi (Hope) var.felix (Clinteria), see felix (Clinteria).

laetus (Chromovalgus), 1913^:408. LECTOTYPE^, JAPAN: Luchu Islands, Oshima,
17^.1913; here designated.

Paralectotypes. JAPAN: 2 $, same data as lectotype; i , same data as lectotype but

Current valid name: Charitovalgus laetus (Arrow) (Arrow, I944/ : 2 3)-

laevis (Glyptothea), 19320 : 126. LECTOTYPE <, BORNEO: Mt Kinabalu, Lumu Lumu,
5500 ft, 15.^.1929 (H. M. Pendlebury); here designated.

Paralectotypes. BORNEO: i ^, same data as lectotype, but 7.^.1929; i $, same data as
lectotype, but i6.iv.i929; i $, same data as lectotype, but 8.^.1929.

laeviventris (Cetonia), igioc : 134. LECTOTYPE ^, INDIA: Manipur (Doherty); here
designated.

Paralectotypes. INDIA: 2 , 2 $, same data as lectotype.

laticollis (Hybovalgus), 19447:238 [original series: 2]. LECTOTYPE . CHINA: Wei-hai
wei, on Oak, 25.^.1899 (T. B. Fletcher); here designated.
Paralectotype. CHINA: i $.

32 M. E. BACCHUS

latipennis (Myoderma), 19410 : 84. Holotype $, SIERRA LEONE: Njala, 20. v. 1930 (E.

Hat 'greaves) .

latipes (Oreoderus), 1944/1 243. Holotype^, VIETNAM (NORTH): Tonkin, Hoabinh (A. de
Cooman) .

lobata (Macronota), 19320 : 128 [original series: i <J, 2 $]. LECTOTYPE $, BORNEO:
Kinabalu, Lumu Lumu, 5500 ft, i6.iv.ig29 (H. M. Pendlebury) ; here designated.

Paralecto types. BORNEO: i $, same data as lectotype, but 8.iv.i929; i $, Mt Kinabalu,
Kamborangah, 7000 ft, 26.iii.i929 (H. M. Pendlebury).

longicollis (Microvalgus), 19447: 245. Holotype, ZAIRE: Ituri, 1918 (L. Burgeon).

longipennis (Protaetia), igioc : 146 [original series: 2]. Holotype $, BURMA: Monti Cariani,
1892 (D. Tornatore) (MCSN, Genoa).

[The second specimen seen by Arrow could not be found in the O. E. Janson Collection
(RHN, Leiden).]

luridus (Calometopus), 19226 : 532. Holotype <$, MALAWI: Mt Mlange, 30. x. 1913 (S. A.
Neave).

malaisei (Trichius), 19380 : 4. Holotype^, BURMA: Kambaiti, 7000 ft, 22.vi.i934 (R. Malaise)
(NR, Stockholm).

Paratypes. BURMA : i $, same data as holotype; i $, same data as holotype but 4-8. vi. 1934
(NR, Stockholm) ; i $, same data as holotype, but 28^.1934 (NR, Stockholm) ; i <$, same data
as holotype but 2000 m, 28^.1934; 2 <$, 'N. E. Burma' (R. Malaise).

maritimus (Oreoderus), I944/: 241 [original series: 8]. LECTOTYPE <$, MALAYA: Langkawi
Islands, i.v.ig28 (H. M. Pendlebury); here designated.

Paralectotypes. MALAYA: i <$, i $ ,same data as lectotype; 1^,1$, same data as lectotype,
but 22.iv.i928; i $, same data as lectotype, but 26.iv.ig28. BURMA: i <$, Rangoon, v-vi.i887
(Fea).

mashunus (Eccoptocnemis), igoga : 520. LECTOTYPE $, RHODESIA: Mashonaland
(H. B. Dobbie) ; here designated.

Paralectotypes. RHODESIA: i <$, same data as lectotype; i <$, same data as lectotype, but
(F. Brooks); i $, Mashonaland, Salisbury, xi.i897 (G. A. K. Marshall); i $, as previous, but
on Cassia pods, xi. 1899.

tnilitaris (Dasyvalgus), igioc : 237. Holotype [not $ as stated by Arrow], INDIA: Niligiri
Hills (G. F. Hampson).

minimus (Dasyvalgus), igioc : 244. LECTOTYPE <$, BURMA: Karen Mts (Doherty); here
designated.

Paralectotypes. BURMA: 5 <J, same data as lectotype; i $, Ruby Mines (Doherty}; 6 <$,
as previous, but 5500-7500 ft.

momeitensis (Oreoderus), igioc : 224. Holotype <$, BURMA: fitat de Momeit, 600 m, 1890

(Doherty} .

Paratypes. BURMA: 2 <$, same data as holotype (BMNH, London; MNHN, Paris).

monstrosus (Placodidus), 19266 : 648. Holotype $, TANZANIA: Dodoma, 1918 (H. L.

Andrewes).

nanus (Xenoreoderus), 19447: 240 [original series: 6]. LECTOTYPE <, BURMA: Yedashe,
9. in. 1918 (A. G. R.); here designated.

Paralectotypes. BURMA: 5 , same data as lectotype.

nasalis (Genuchus), 19266 : 650. LECTOTYPE $, UGANDA : shores of L. Isolt or Wamala,
3800 ft, 7-8.1.1912 (S. A. Neave); here designated.

Paralecto type. UGANDA: Kampala, I7.X.I9I5 (C. C. Gowdey).

CETONIINAE DESCRIBED BY G. J. ARROW 33

ntgra (Myoderma), 19010 : 258 [original series: 2]. LECTOTYPE $, CAMEROUN: here desig-
nated.

Paralectotype. I $, same data as lectotype.

nigrolineata (Gnathocera), 19226 : 526. Holotype $, GUINEA: Kondia.

nitidicauda (Trichius), 1938^:2. Holotype $, BURMA: Kambaiti, 7000 ft, 3^.1934 (R.

Malaise) (NR, Stockholm).

Paratypes. BURMA: i $>, same data as holotype, but 8.vi.i934 (NR, Stockholm); i <$,
same data as holotype, but 12. iv. 1934; i $, same data as holotype, but 2000 m, 2.vi.i934.

nitidus (Coenochilus), igioc : 210. Holotype^, INDIA: Bombay.
Paratype. INDIA: Kanara (H. E. Andrewes).

oberthuri (Clinteria), igioc : 181. LECTOTYPE <, SIKKIM: Kurseong, 1934 ( R - p -

Bretandeau) ; here designated.

Paralectotypes. SIKKIM : i <J, same data as lectotype (MNHN, Paris) ; i <$, environs de
Kurseong (R. P. Bretandeau) (MNHN, Paris).

occidentalis (Xenoreoderus), igioc : 233. LECTOTYPE $, INDIA: Belgaum, Bombay;
here designated.

Paralectotypes. INDIA: i , same data as lectotype; 5 $, Belgaum.

opacicauda (Coelocorynus), 19266:654 [original series: 3 ]. LECTOTYPE <, KENYA:
Aberdare Mts, 10 ooo ft, vii.igig (H. L. Andrewes); here designated.
Paralectotypes. KENYA: 2 <$, same data as lectotype.

opacipennis (Leucocelis), igoga : 522. Lectotype <$, ZAIRE: 150-200 miles W. of Kambove,
3500-4500 ft, 24. ix. 1907 (S. A . Neave) ; designated by Ruter (1967, Rev . Zool. Bot. afr. 76 : 42).
Paralectotypes. ZAIRE : 6 $, same data as lectotype.

ovaliceps (Macronota), i94ie : 83. Holotype $, BURMA: Sima, i6.viii.i9i8 (P. M. R. Leonard).

ovicollis (Dasyvalgus), igioc : 242. Holotype^, BURMA: Ruby Mines (Doherty).

Paratypes. BURMA: 2 <J (BMNH, London), 3 (MNHN, Paris), same data as holotype
but 12202300 m, 1890.

peninsularis (Acanthovalgus), 19447:232 [original series: i <J, 2 $]. LECTOTYPE <,
MALAYA: Pahang, Cameron Highlands, Gunong Tesar, 5565 ft, 2O.V.I939 (H. M. Pendlebury);
here designated.

Paralectotypes. MALAYA: 2 $, Pahang Cameron Highlands, 5000 ft.

planata (Heterorrhina), igioc : 94. LECTOTYPE <J, INDIA; here designated.

Paralectotypes. INDIA: 3 <, Nilgiri Hills i <J, Kanara (T. R. Bell); i $, Kanara; 3 ^,
Mercara Coorg, 24.vii.i9O5 (H. M. Lefroy); i $, Malabar, Nagdani.

planisslmus (Placodidus), 19266 : 649. Holotype $, RHODESIA: Lukosi, ii. 1905.

polychrous (Dasyvalgus), 19447:235 [original series: 6 <J]. LECTOTYPE <$, BORNEO:
W. Sarawak, Mt Matang, 3000 ft, 2o.xii.i9i3 (G. E. Bryant); here designated.

Paralectotypes. BORNEO: i , same data as lectotype, but 1000 ft, 11.11.1914; i $, same
data as lectotype, but 1000 ft, 12. ii. 1914; i <, Sarawak, Bau Mantang, via Single, 12-20^.1909
(C. J. Brooks); i $, Sarawak, Matang, lo.v.igog (C. J. Brooks). [Arrow gives the date as
'Nov.' despite the fact that the specimen labelled by him as 'type' is clearly '2o.xii.i9i3'.
It appears that he read the 'ii' on the other two specimens collected by Bryant as 'n'.]

prunina (Protaetia), igioc : 147. LECTOTYPE $, BURMA: Moulmein (Weston); here
designated.

Paralectotype. BURMA: i $, same data as lectotype. [Specimen(s) from Yun-za-lin
missing.]

34 M. E. BACCHUS

pubescens (Glyptothea), 19320 : 126 [original series: 8 <]. LECTOTYPE <$, BORNEO: Mt
Kinabalu, Lumu Lumu, 5500 ft, 8.iv.i929 (H. M. Pendlebury) ; here designated.

Paralecto types. BORNEO: i <$, same data as lectotype, but 14. iv. ; I $, same data as
lectotype, but 28. iv. 1929; i <$, same data as lectotype, but 30.^.1929; i <$, same data as
lectotype, but 2^.1929; i $, Mt Kinabalu, Kamborangah, 8000 ft, 3.^.1929; 1$, Mt Kinabalu,
Marei Pare!, 5000 ft, 27.^.1929.

pulchellus (Cymophorus), igioc : 203. LECTOTYPE $, INDIA: Chota Nagpore, Barway
(R. P. Car don) ; here designated.

Paralecto types. INDIA: i $, same data as lectotype (MNHN, Paris); 2 <$, Chota Nagpore,
Nowatoli (R. P. Cardon) (MNHN, Paris) ; I $, Chota Nagpore, Chandernagore (R. P. Cardon)
(RNH, Leiden).

pusilla (Myoderma), 19066 : 135. LECTOTYPE^, ANGOLA: Bihe; here designated.

Paralectotypes. ANGOLA: 5 $, same data as lectotype; 2 , Loanda, Pundo Andongo

(Dr A nsorge) .

pusillus (Callinomenes), igioc : 216 [original series: 3]. LECTOTYPE <J, INDIA: Assam,
North- West Frontier; here designated.

Paralecto type. INDIA: i $, Patkai Mts (Doherty).
[Specimen from Sikkim, Mungphu missing.]

pusillus (Pogonopus), igioc : 117. Holotype g, INDIA: Coimbatore, Podanur.

pygmaea (Heterorrhina (Ptychodesthes)), 19226 : 527. LECTOTYPE g, RHODESIA:
'Gazaland', Chirinda Forest (David Odendaal); here designated.

Paralectotypes. RHODESIA: 4 , same data as lectotype; 4 <$, 'Mashonaland', Chirinda
Forest, 1906 (C. F. M. Swynnertori) ; i , 'Gazaland', nr Chirinda, viii.i9O7 (G. A. K.
Marshall}; i $, 'Gazaland', nr Chirinda, 3800 ft, 5-31.111.1908.

[By modern definition, Chirinda Forest is not in Gazaland or Mashonaland. It is no miles
south of Umtali.]

Current valid name: Ptychodesthes alternata Klug subsp. pygmaea Arrow (Schein, 1941,
Mitt, munch, ent. Ges. 33 : 36).

quadricarinatus (Oreoderus), 19447:242. Holotype $, VIETNAM (NORTH): Hat Thoung
10. xi. 1917.

Paratypes. i g, LAOS: Pak Neun, i.x.igiS. THAILAND: i , Souvannaphoun, 2i.ix.i9i8.

ranu (Protaetia), igioc : 153 [original series: 2 , i $]. Holotype <, INDIA: Assam, Shillong.
Paratypes. INDIA: i g, same data as holotype; i $, Assam, Khasi Hills (RNH, Leiden).

reticulatus (Microvalgus), 19447 : 246 [original series: 3 <$]. LECTOTYPE $, GHANA: Aburi,
1912-13 (W. H. Patterson); here designated.

Paralectotypes. GHANA : 2 $, same data as lectotype.

rotundicollis (Macronota), IQ^IC : 81. Holotype <$, INDIA: Shevaroys, Yercaud, 4500 ft,
2i.iv-4.v.i9i3 (Y. R.).

ruflcauda (Dasyvalgus), I944/: 237 [original series: 2 <$]. LECTOTYPE,^, MALAYA: Kedah
Peak, 3950 ft, 27.iii.i928 (H. M. Pendlebury); here designated.

Paralectotype. MALAYA: i , same data as lectotype, but 3300-3950 ft, iii.i928.

rufipennis (Dasyvalgus), 1944 / : 235 [original series: 7 $}. LECTOTYPE ^, BORNEO: W
Sarawak, Mt Matang, 12.11.1914 (G. E. Bryant}; here designated.

Paralectotypes. BORNEO: i ^, same data as lectotype, but 1000 ft, 9.11.1914; 2 $, same
data as lectotype, but 1000 ft, 10.11.1914; 2 $, same data as lectotype, but 1000 ft, 11.11.1914;
i $, same data as lectotype, but 1000 ft, no date.

CETONIINAE DESCRIBED BY G. J. ARROW 35

[Arrow erroneously records the date of collection as 'Nov.', interpreting 'ii' on the labels
as 'n'. I have confirmed from his collecting journal that Bryant was not at Mt Matang in
November 1914.]

rufiventris (Ligyromorphus), 19010 : 257. LECTOTYPE <$, RHODESIA: Mashonaland,
Salisbury, flying at dusk, xi.igoo (G. A. K. Marshall); here designated.
Paralecto types. RHODESIA: i <, 2 $, same data as lectotype; 1^,1$, same data as
lectotype, but no date.

Current valid name: Diploa rufiventris (Arrow) (Peringuey, 1907, Trans. S. Afr. phil. Soc.
13 : 316).

rufus (Mimovalgus), I944/: 239. Holotype $, MALAYA: Penang (Bowring).

runsorica (Tmesorrhina), igogd : 192 [original series : i <$, 2 $]. LECTOTYPE <$, UGANDA:
Ruwenzori, 7000-8000 ft (Scott Elliot) ; here designated.

Paralectotype. UGANDA: i $, E. Ruwenzori, Old Camp, 6000-7000 ft, 1906 (Hon. G.
Legge &- A.F. R. Wollaston).

scintillans (Trigonophorus), igioc : 105. LECTOTYPE <$, INDIA : Mungphu ; here designated.
Paralecto types. INDIA: i $, same data as lectotype; i $, Darjeeling (Col. Buckley};
3 ? (BMNH, London), i $ 'India' (MNHN, Paris). SIKKIM: 3 $, 5 $, Kurseong, 1898 (R. P.
Decoly) (MNHN, Paris) ; i $ (chasseurs indigenes) (MNHN, Paris) .

Scorpio (Charitovalgus), I944/ : 2 3 I - Holotype $, NEW GUINEA: Kokoda, 1200 ft, vii.i933
(L. E. Cheesman) ['Lower rain forest, 1300 ft' on a separate label.]

sex-dentatus (Hybovalgus), 19447:238 [original series: 3]. LECTOTYPE <$, LAOS: Ban
Naban, i6.iii.i9i5 (R. V. de Salvaza); here designated.

Paralecto types. LAOS: i <, same data as lectotype; i $, Xieng Khouang, 2o.iii.i9i5
(R. V. de Salvaza).

sobrinus (Trichius), 19226 : 529 [original series: 2 ^, 5 $]. LECTOTYPE <J, UGANDA: S. of
L. George, 3200-3400 ft, 17-19. xi. 1911 (S. A. Neave); here designated.

Paralectotypes. UGANDA: i g (BMNH, London), i $ (RNH, Leiden), same data as lecto-
type; 2 , Daro or Durro Forest, Toro, 4000-4500 ft, 25-29.x.i9ii (S. A. Neave); i $, Mabira
Forest, Chagwe, 3500-3800 ft, i6-25.vii.i9ii (S. A. Neave).

Current valid name: Pileotrichius austerus Bourgoin (Arrow, 1941^ : 77).

solidus (Coenochilus), igioc : 209 [original series: i <$, i $]. Holotype $, BHUTAN: British
Bootang, 1898 (L. Durel).

Paratype. BHUTAN: i ^, British Bootang, Padong (MNHN, Paris).

spectralis (Stripsipher), 19266 : 652. Holotype <$, SOUTH AFRICA: Pondoland, Port St John,
ix.i923 (R. E. Turner).

striata (Myoderma), ig^ic : 85. Holotype <$, TANZANIA: 35 miles E. of Singida, Msagaa,
x-xii.i935 (E. Burt).

Paratypes. TANZANIA: 3 $, same data as holotype.

subopaca (Anomalocera), igojc : 348 [original series : 6]. LECTOTYPE <J, INDIA: Manipur
(Doherty) ; here designated.

Paralectotypes. INDIA: 5 $, same data as lectotype.

Current valid name: Rhomborrhina olivacea Janson subsp. subopaca (Arrow) (Miki6, 1967,
Ent. Abh. Mus. Tierk. Dresden 35 : 304).

suco (Oreoderus), 19447 : 2 43- LECTOTYPE^, INDIA: Coorg, Sidapur, i.x.i9i7 (T. R. N.);
here designated.

Paralectotypes. INDIA: i $, same data as lectotype, but 14.^.1917; i c?, same data as
lectotype, but 22. iv. 1917; i ^, same data as lectotype, but 6^.1917; i <$, same data as
lectotype, but 7^.1917.

36 M. E. BACCHUS

tessellatus (Dasyvalgus), 1944/1236. Holotype $, MALAYA: Cameron Highlands, Pahang,
Gimling Kial, 5000 ft, 24^.1939 (H. M. Pendlebury).

tigrina (Glycyphana (Gametis)), igogd : 193. Holotype $, UGANDA: E. Ruwenzori, 6000-
13 ooo ft, 1906 (Hon. G. Legge <& A. F. R. Wollaston).

Current valid name: Phonotaenia tigrina (Arrow) (Schenkling, 1921, Coleoptm Cat. 72 : 311).

tonkinensls (Valgus), 19447: 229. Holotype $, VIETNAM (NORTH): Hoabinh (A. de Cooman).
Paratype. VIETNAM (NORTH) : i <$, same data as holotype.

transparens (Calometopus), 19226:530. LECTOTYPE <, MALAWI: Mlange, 29.xii.igi3

(S. A. Neave) ; here designated.

Par alecto types. MALAWI: i <$, same data as lectotype, but y.xi.i922; i $, same data as
lectotype, but 4.xii.i9i2; i , same data as lectotype, but 22.1.1913; i $, same data as
lectotype, but 4.^.1913; i , same data as lectotype, but 6.xii.i9i3.

tridens (Diploa), 19066 : 136. LECTOTYPE , ANGOLA: Garengaze; here designated.
Paralectotypes. ANGOLA: 2 $, same data as lectotype; 2 <$, Bihe.

tridens (Euryvalgus), 19447 : 233. Holotype <, BORNEO: W. Sarawak, Quop, 8.iv.i9i4
(G. E. Bryant).

truncata (Clinteria), igojc : 352. LECTOTYPE <, INDIA: Nilgiri Hills (G. F. Hampson);
here designated.

Paralectotypes. INDIA: i <^, same data as lectotype; I <$, i $, same data as lectotype, but
6000 ft (H. L. Andrewes); I $, i $, same data as lectotype, but (Captain Downing); 4 <$, 5 $,
Nilgiri Hills; i <j>, Madras, Nilgiris, Naduvatum, 7000 ft, v.igo4 (W. Rowson).

turneri (Stripsipher), 19266 : 652. LECTOTYPE $, SOUTH AFRICA: Pondoland, Port
St John, xi. 1923 (R. E. Turner] ; here designated.

Paralectotypes. SOUTH AFRICA: i g, same data as lectotype; 1^,1$, same data as lecto-
type, but ix.i923; 2 $, same data as lectotype, but x.ig23; 2 $, same data as lectotype, but
1.1924; i $, same data as lectotype, but xii.i924.

variegatus (Parapilinurgus), igioc : 204 [original series: 2]. Holotype , BURMA: Karen
Mts (Doherty).

Paratype. VIETNAM (NORTH): i $>, Dong Van, 1898 (Captain Gadel) (MNHN, Paris).

viduatus (Dasyvalgus), igioc : 236. Holotype $, BURMA.

waterhousei (Macronota), igioc : 56. LECTOTYPE $, INDIA: Nilgiri Hills (H. L.
Andrewes); here designated.

Paralectotypes. INDIA: 5 , 8 <j>, same data as lectotype; i $, same data as lectotype, but
(G. F. Hampson); i $, same data as lectotype, but no collector; 2 $, Nilgiris (H. M. Lefroy)',
2 (ff, 2 $, Nilgiris, Naduvatum, 7000 ft, v.1904 (W. Rowson) ; 2 <J, Nilgiris, Naduvatum, 7000 ft;
4 (, i ?, Anamalai Hills; i $, 'India'.

zulu (Scaptobius), I936c : 260. Holotype [impossible to determine sex without dissection],
SOUTH AFRICA: Zulu, lives in ants' nests only, with the same species of ants as Paussus
cucullatus.

BIBLIOGRAPHY

This bibliography includes all G. J. Arrow's entomological works.

ARROW, G. J. 18990. On sexual dimorphism in beetles of the family Rutelidae. Trans.

ent. Soc. Land. 1899 : 255-269.
18996. Notes on the Rutelid genera Anomala, Mimela, Popillia and Strigoderma. Trans.

ent. Soc. Land. 1899 : 271-276.

CETONIINAE DESCRIBED BY G. J. ARROW 37

ARROW, G. J. iSggc. On the Rutelid beetles of the Transvaal; an enumeration of a collection

made by Mr W. L. Distant. Ann. Mag. nat. Hist. (7) 4 : 118-122.
18990". Notes on the classification of the coleopterous family Rutelidae. Ann. Mag.

nat. Hist. (7) 4 : 363-370.
1899^. On sexual dimorphism in the Rutelid genus Parastasia, with descriptions of new

species. Trans, ent. Soc. Lond. 1899 : 479-499, pi. 17.
iSgg/. Anomala donovani Marsham: synonymical note. Entomologist's mon. Mag.

35 : 269.
19000. In Gahan, C. J. & Arrow, G. J., 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.

5. Coleoptera. Proc. zool. Soc. Lond. 1900 : 21-33, pi- *
19006. On pleurostict Lamellicorns from Grenada and St Vincent (West Indies). Trans.

ent. Soc. Lond. 1900 : 175-182.
igooc. In Andrews, C. W. et al. A monograph of Christmas Island, British Museum

(Natural History), pp. 89-127, pis 10, n.
19010. Remarks upon the genus Rhysodes, with descriptions of some new oriental species.

Ann. Mag. nat. Hist. (7) 7 : 83-89.
19016. Remarks on secondary sexual differences in Rutelid Coleoptera, with descriptions

of some new forms. Ann. Mag. nat. Hist. (7) 7 : 393-401, 2 figs.
igoic. The Rutelid genus Adorodocia and a new allied form. Ann. Mag. nat. Hist.

(7) 8 : 35-38, 4 figs.

igoid. The Rutelid genus Adorodocia. Ann. Mag. nat. Hist. (7) 8 : 193-196, 4 figs.

19010. On a new genus and two new species of African Cetoniidae. Ann. Mag. nat.

Hist. (7) 8 : 257-259.
1 90 if. The Carabid genus Pheropsophus : notes and descriptions of new species. Trans.

ent. Soc. Lond. 1901 : 193-207, pi. 9.
1901^. The genus Hyliota, of the coleopterous family Cucujidae, with descriptions of new

forms and a list of the described species. Trans, ent. Soc. Lond. 1901 : 593-601.
19020. On Rutelid and Melolonthid beetles from Mashonaland and East Africa. Ann.

Mag. nat. Hist. (7) 9 : 89-101.
19026. Notes and descriptions of some Dynastidae from tropical America, chiefly

supplementary to the 'Biologia Centrali- Americana'. Ann. Mag. nat. Hist. (7) 10 : 137

147.
19030. On the affinities and nomenclature of certain genera of Melolonthid and Rutelid

Coleoptera. Ann. Mag. nat. Hist. (7) 11 : 303-306.
19036. On the laparostict Lamellicorn Coleoptera of Grenada and St Vincent (West

Indies). Trans, ent. Soc. Lond. 1903 : 509-520, 4 figs.
19040. Note on two species of Coleoptera introduced into Europe. Entomologist's mon.

Mag. 40 : 35-36.
19046. In Gahan, C. J. & Arrow, G. J. List of the Coleoptera collected by Mr A. Robert

at Chapada, Matto Grosso (Percy Sladen expedition to central Brazil). Proc. zool. Soc.

Lond. 1903 (2) : 244-258, pi. 28.
I904C. On the coleopterous group 'Heptaphyllini' of De Borre. Ann. Mag. nat. Hist.

(7) 14 : 30-33.
19040". Sound-production in the Lamellicorn beetles. Trans, ent. Soc. Lond. 1904 : 709-

750, pi. 36.
1905. On some oriental Aphodiid Coleoptera of the Rhyparus group, with description

of a new genus. Ann. mag. Nat. Hist. (7) 15 : 534-540.
19060. On three remarkable new Melolonthid Coleoptera from Sumatra and Borneo in

the British Museum. Ann. mag. nat. Hist. (7) 18 : 48-50.
19066. On Lamellicorn Coleoptera from Portuguese West Africa, with descriptions of

new species. Ann. Mag. nat. Hist. (7) 18 : 127-136, i fig.
19070. A contribution to the classification of the coleopterous family Passalidae. Trans.

ent. Soc. Lond. 1906 : 441-469.

38 M. E. BACCHUS

ARROW, G. J. 19076. On two new parasitic Coleoptera (Fam. Staphylinidae) from South

America. Ann. Mag. nat. Hist. (7) 19 : 125-127.
19076. Some new species and genera of Lamellicorn Coleoptera from the Indian empire

(part i). Ann. Mag. nat. Hist. (7) 19 : 347-359.

19076?. Some new species and genera of Lamellicorn Coleoptera from the Indian empire
(part 2). Ann. Mag. nat. Hist. (7) 19 : 416-439, 3 figs.

19070. Resultats de 1'expedition scientifique Neerlandaise a la Nouvelle Guinea.

Lucanidae and Scarabaeidae P.P. Nova Guinea 5 : 2728.
19080. Notes on the coleopterous genus Oniticellus and descriptions of some new species

from India. Ann. Mag. nat. Hist. (8) 1 : 178-183.
19086. On some new species of the coleopterous genus Mimela. Ann. Mag. nat. Hist.

(8) 1 : 241-248.
- 19086. A contribution to the classification of the coleopterous family Dynastidae.

Trans, ent. Soc. Lond. 1908 : 321-358.

19080". In Schultze, L., Forschungsreise im westlichen und zentralen Siidafrika 1903-5.
Ruteliden und Melolonthiden. Denks. med. nat. Ges. 13 : 435-438.

19090. On some new species of Coleoptera from Rhodesia and adjacent territories. Ann.

Mag. nat. Hist. (8) 3 : 517-523.

19096. Four new Lamellicorn Coleoptera from the Oriental Region. Ann. Mag. nat.
Hist. (8) 4 : 91-94.

igogc. Systematic notes on Coleoptera of the clavicorn families. Ann. Mag. nat. Hist.

(8) 4 : 190-196.
19090". In Arrow, G. J. et al., Ruwenzori expedition reports- 14. Coleoptera. Trans.

zool. Soc. Lond. 19 : 185-200.
19090. On the characters and relationships of the less-known groups of Lamellicorn

Coleoptera, with descriptions of new species of Hybosorinae etc. Trans, ent. Soc. Lond.

1909 : 479-507.

19100. On the Lamellicorn beetles of the genus Peltonotus with descriptions of four new
species. Ann. Mag. nat. Hist. (8) 5 : 153-157.

19106. On a few new Bornean beetles of the Rutelid genera Mimela and Anomala.

Ann. Mag. nat. Hist. (8) 6 : 64-72.
191 oc. Fauna Br. India. Coleoptera, Lamellicornia (Cetoniinae and Dynastinae).

xiv + 322 pp., 2 pi., 76 figs.
19110. A synoptical revision of the Dynastid genus Lonchotus. Ann. Mag. nat. Hist.

(8) 7 : 84-89.
19116. Notes on the Lamellicorn beetles of the genus Golofa with descriptions of three

new species. Ann. Mag. nat. Hist. (8) 7 : 136-141.
191 ic. On Lamellicorn beetles belonging to the subfamilies Ochodaeinae, Orphninae,

Hybosorinae and Troginae. Ann. Mag. nat. Hist. (8) 7: 390-397.
19110". Upon the Dynamopinae, a new subfamily of Lamellicorn beetles. Ann. Mag.

nat. Hist. (8) 7 : 610612.
19110. Notes on the coleopterous subfamily Dynastinae, with descriptions of new genera

and species. Ann. Mag. nat. Hist. (8) 8 : 151-176, pis 4, 5.
191 1/. A new genus of Ruteline Coleoptera from the Indian region. Ann. Mag. nat.

Hist. (8) 8 : 268-271.
191 ig. On the Ruteline Coleoptera of Ceylon, with descriptions of new species. Ann.

Mag. nat. Hist. (8) 8 : 354-365.
1911/1. Some new species of the coleopterous genus Anomala from southern India. Ann.

Mag. nat. Hist. (8) 8 : 473-488.
19120. Some new species of the Lamellicorn genus Anomala from Sikkim, north India.

Ann. Mag. nat. Hist. (8) 9 : 72-84.
19126. On a new species of Melolonthid beetle (Phytalus smithi) destructive to sugar

cane. Ann. Mag. nat. Hist. (8) 9 : 455-459, 4 figs.

CETONIINAE DESCRIBED BY G. J. ARROW 39

ARROW, G. J. 19120. A synoptical review of the coleopterous genus Hexodon (Dynastinae) .

Ann. Mag. not. Hist. (8) 9 : 594-600.
191 2d. Scarabaeidae : Pachypodinae, Plecominae, Aclopinae, Glaphyrinae, Ochodaeinae,

Orphninae, Idiostominae, Hybosorinae, Dynamopinae, Acanthocerinae, Troginae.

Coleoptm Cat. pars 43, 66 pp.
19120. Descriptions of some new Burmese species of Ruteline Coleoptera belonging to

the genus Anomala. Ann. Mag. nat. Hist. (8) 10 : 327-340, 2 figs.
19130. Some new species of Lamellicorn beetles from Brazil. Ann. Mag. nat. Hist.

Hist. (8) 11 : 456-466.
19136. Notes on the Lamellicorn genus Popillia and descriptions of some new Oriental

species in the British Museum. Ann. Mag. nat. Hist. (8) 12 : 38-54, 15 figs.
1913^. Zoological results of the Abor expedition 1911-12, Coleoptera IV Lamellicornia.

Rec. Indian Mus. 8 : 191-196.
1913^- Notes on the Lamellicorn Coleoptera of Japan and descriptions of a few new

species. Ann. Mag. nat. Hist. (8) 12 : 394-408.
19135. Synopsis of the Melolonthid genus Ancistrosoma, with descriptions of new species

and an allied new genus. Ann. Mag. nat. Hist. (8) 12 : 425432, 4 figs.
19140. On the Ceylonese species of Ruteline Coleoptera belonging to the genus Adoretus.

Ann. Mag. nat. Hist. (8) 13 : 587-594.
19146. On the Burmese species of Ruteline Coleoptera belonging to the genus Adoretus.

Ann. Mag. nat. Hist. (8) 13 : 594-601.
1914^. Some further notes on Lamellicorn beetles of the subfamily Dynastinae. Ann.

Mag. nat. Hist. (8) 14 : 257-276, pi. 13.
I9i5#. Notes on the coleopterous family Dermestidae, and descriptions of some new

forms in the British Museum. Ann. Mag. nat. Hist. (8) 15 : 425-451.
19156. Note on the nomenclature of certain species of Ruteline Coleoptera. Ann.

Mag. nat. Hist. (8) 16 : 231-232.
1915^. Upon a remarkable new genus of Lamellicorn beetles from Borneo. Ann. Mag.

nat. Hist. (8) 16 : 317-319.
1915^. Upon the beetles of the Melolonthid genus Rhopaea found in the Fiji Islands.

Ann. Mag. nat. Hist. (8) 16 : 319-321.

19150. In Arrow, G. J. et al., Report on the Coleoptera collected by the British Orni-
thologists' Union and the Wollaston expedition in Dutch New Guinea. Trans, zool. Soc.

Lond. 20 : 497-540, pi. 39.
19160. A new species of Thaumaglossa bred from the egg clusters of Mantidae. Proc.

ent. Soc. Lond. 1915 : cxii-xciii.

19166. The melolonthine beetles of Ceylon. Ann. Mag. nat. Hist. (8) 18 : 429-444.

igi6c. On the Lamellicorn Coleoptera of Larat Island. Ann. Mag. nat. Hist. (8)

18 : 492-498.

19170. Some systematic notes on Melolonthine Coleoptera. Ann. Mag. nat. Hist. (8)

19 : 59-65.

19176. Fauna Br. India. Coleoptera, Lamellicornia part 2 (Rutelinae, Desmonycinae

and Euchirinae). xii + 387 pp., 5 pi., 77 figs.
igi7c. The Khapra beetle (Trogoderma khapra sp. n.), an Indian grain-pest. Ann.

Mag. nat. Hist. (8) 19 : 481-482.
191 70". A systematic revision of the African species of the coleopterous family Erotylidae.

Ann. Mag. nat. Hist. (8) 20 : 137-156.
191 je. Some insects injurious to cacao plants in the Belgian Congo. Bull. ent. Res.

8 : III-H2.
I9I7/. The life history of Conwentzia psociformis Curt. Entomologist's mon. Mag.

53 : 254-257, i fig.

191 7g. A note on the coleopterous genus Euxestus. Ann. Mag. nat. Hist. (8) 20 : 368.

igiSa. The life history of Scymnus capitatus F. Entomologist's mon. Mag. 54 : 8-9,

i fig.

4 o M. E. BACCHUS

ARROW, G. J. 19186. The larva of Scymnus. Entomologist's mon. Mag. 54 : 39-40.

19190. A remarkable new ball-rolling beetle (family Scarabaeidae) . Ann. Mag. nat.

Hist. (9) 3 : 433-435. i fig-
19196. Systematic notes on a few Melolonthine Coleoptera. Ann. Mag. nat. Hist.

(9) 4 : 21-29, pi. i.
19190. Notes on Ruteline Coleoptera and descriptions of a few new species in the British

Museum. Ann. Mag. nat. Hist. (9) 4 : 379-385, pi. 8.

igzoa. Horned beetles. Proc. ent. Soc. Lond. 1920 : xviii-xx.

10206. A list of the Endomychid Coleoptera of Indo-China with descriptions of new

species. Ann. Mag. nat. Hist. (9) 5 : 321-336.
19200. A contribution to the classification of the coleopterous family Endomychidae.

Trans, ent. Soc. Lond. 1920 : 1-83, pi. i.
19200*. A remarkable new genus of Lamellicorn beetles. Ann. Mag. nat. Hist. (9)

6 : 431-434-

19200. A peculiar new genus of Australian beetles. Ann. Mag. nat. Hist. (9) 6 : 434-437.

I92O/. A new genus of Clavicorn beetles. Ann. Mag. nat. Hist. (9) 6 : 437-440.

1920^. On the Oriental members of the coleopterous group Macrodactylides (Melo-

lonthidae). Ann. Mag. nat. Hist. (9) 6 : 441-455.
1920/1. Some new West Indian species of the Melolonthid genus Lachnosterna. Bull.

ent. Res. 11 : 189-193.
I92ia. Two remarkable new Ruteline beetles from Indo-China. Ann. Mag. nat. Hist.

(9) 7 : 378-381, pi. 10.
19216. A revision of the Melolonthine beetles of the genus Ectinohoplia. Proc. zool.

Soc. Lond. 1921 : 267-276. pi. i.

19210. Oviposition of Aphelinus chaonia Walker. Entomologist's mon. Mag. 57 : 211.

192 id. In Babault, G., Mission dans les provinces centrales de I'Inde et dans la rdgion

occidentale de I'Himalaya. Insectes Coleopterds, Lamellicornes, Melolonthinae, Rutelinae,

Dynastinae. 16 pp., i pi. Paris.
19220. A list of the Erotylid Coleoptera of Indo-China, with descriptions of new species.

Trans, ent. Soc. Lond. 1921 : 285-306.
19226. A few new African Cetoniine beetles. Ann. Mag. nat. Hist. (9) 9 : 525-532,

pi. 8.
19220. Coleoptera, Erotylidae and Endomychidae, from the Seychelles, Chagos and

Amirantes Islands. Ann. Mag. nat. Hist. (9) 10 : 73-83, pi. 3.
19230. Notes on Endomychid Coleoptera and descriptions of new species in the British

Museum. Trans, ent. Soc. Lond. 1922 : 484-500, i fig.
19236. The fauna of an island in the Chilka Lake. Certain Coleoptera of Barkuda.

Rec. Indian Mus. 25 : 259-263.

19230. New Erotylidae. Treubia 3 : 272274.

19230*. Obituary [Canon William Weekes Fowler.] Entomologist 56 : 170.

19240. New Erotylidae. Nova Guinea 15 : 5-8.

19246. On the Stethaspis group of Melolonthid Coleoptera. Ann. Mag. nat. Hist.

(9)13: 546-553-
19240. Vocal organs in the coleopterous families Dytiscidae, Erotylidae and

Endomychidae. Trans, ent. Soc. Lond. 1924 : 134-143, 2 figs.
19250. Notes on the coleopterous families Hybosoridae and Trogidae. Ann. Mag. nat.

Hist. (9) 15 : 328-331.
19256. Fauna Br. India. Coleoptera, Clavicornia (Erotylidae, Languriidae and

Endomychidae). xv + 416 pp., 76 figs, i map.
19250. A few new species of Melolonthine Coleoptera. Ann. Mag. nat. Hist. (9) 16 : 209-

211, 5 figs.

19260. Fauna Buruana. Erotylidae, Languriidae and Endomychidae. Treubia 7 : 122-127.

19266. A few new African species of Cetoniine Coleoptera. Ann. Mag. nat. Hist.

(9) 17 : 648-654, i fig.

CETONIINAE DESCRIBED BY G. J. ARROW 41

ARROW, G. J. 19266. Fauna Sumatrensis. Endomychidae and Erotylidae (Col.). Ent.Mitt.

15 : 248-262.

19260". Notes on Oriental Endomychidae and Erotylidae (Col.). Ent. Mitt. 15 : 354-358.

19260. Mimicry in Coleoptera. Proc. ent. Soc. Lond. 1 : 1819.

1927*3. A note on the coleopterous genus Aserica (Melolonthinae) . Proc. ent. Soc.

Wash. 29 : 69-70. .
19276. Notes on the coleopterous genus Sisyphus. Ann. Mag. nat. Hist. (9) 19 : 456-465,

pi. 12.
19276. A note on the coleopterous genus Trox, with descriptions of a few new Asiatic

species. Ann. Mag. nat. Hist. (9) 19 : 465-468.

19270*. Fauna Sumatrensis. Endomychidae and Erotylidae. Supplta ent. 15 : 112-114.

19270. An entomological mystery solved. Nat. Hist. Mag. 1 : 91-93.

19277. Coleoptera, Clavicornia and Lamellicornia. Insects Samoa. Pt. 4, fasc. i : 35-66,

13 figs.

19280. Some more mimetic beetles. Nat. Hist. Mag. 1 : 244-251.

19286. Polymorphism in horned beetles. Trans, ent. Soc. Lond. 1928 : 73-77, pi. 5.

19286. Coleoptdres Erotylides et Endomychides de 1'Indo-Chine Fran9aise. Faune

Colon, fr. 2 : 329-357, 14 figs.
1929^. A revision of the African Coleoptera belonging to the family Languriidae. Proc.

zool. Soc. Lond. 1929 : 1-15, pi. i.

19296. A new genus of Silphid Coleoptera from Persia. Zool. Anz. 82 : 96-99, 2 figs.

19296. In Schouteden, H. etal., Voyage au Congo de S.A.R. le Prince Leopold de Belgique

1925, Coleoptera 9 Passalidae. Revue Zool. Bot. afr. 17 : 112.
19290*. On the families of Coleoptera related to the Erotylidae, with descriptions of a

new family, two new genera, and a few new species. Ann. Mag. nat. Hist. (10) 4 : 305-

322.
19290. A remarkable new genus of Coccinellid Coleoptera. Ann. Mag. nat. Hist.

(10) 4 : 463-465, i fig.
19300. A new family of Heteromerous Coleoptera (Hemipeplidae), with descriptions of

a new genus and a few new species. Ann. Mag. nat. Hist. (10) 5: 225-231, 2 figs.

19306. An interesting case of mimicry in beetles. Proc. R. ent. Soc. Lond. 4 : 113114.

19306. Two notable new Lamellicorn beetles from Indo-China. Ann. Mag. nat. Hist.

(10) 6 : 541-543. pl- 14-

19300*. Swarming of a Carabid beetle to light. Entomologist's mon. Mag. 66 : 232.

19300. Aberration of instinct in a bark beetle. Entomologist's mon. Mag. 66 : 232.

19310. Coleoptera (Lamellicornia, Clavicornia and Rhyncophora) collected in Korinchi,

West Sumatra by Messrs H. C. Robinson and C. Boden Kloss. /. fed. Malay St. Mus.

8 (3) : 206-209.
19316. The coleopterous genus Trichillum (Copridae), with a key to the species. Ann.

Mag. nat. Hist. (10) 8 : 609-611.
19316. Two new species of Lamellicorn beetles belonging to the genus Peltonotus. Ann.

Mag. nat. Hist. (10) 8 : 611-612.
191310". Fauna BY. India. Coleoptera, Lamellicornia part 3 (Coprinae). xii + 428 pp.,

13 pis, i map, 60 figs.
19320. A few new species of Cetoniine Coleoptera from Mt Kinabalu, British North

Borneo. Ann. Mag. nat. Hist. (10) 9 : 125-128.
19326. A few new species of Melolonthine Coleoptera. Ann. Mag. nat. Hist. (10) 9 : 189-

197-
19326. New species of Lamellicorn beetles (subfamily Coprinae) from South America.

Stylops 1 : 223-226, 3 figs.
I 933- A few new Melolonthine Coleoptera from Mexico. Ann. Mag. nat. Hist. (10)

11 : 145-151-
19336. The genus Uroxys (Coleoptera, Copridae), with descriptions of some new species.

Ann. Mag. nat. Hist. (10) 11 : 385-399, 9 figs.

42 M. E. BACCHUS

ARROW, G. J. 1933^. A further note on the coleopterous genus Aserica (Melolonthinae) .

Proc. ent. Soc. Wash. 35 : 71-73.
I 933d. Notes on Coprid Coleoptera, with descriptions of a new genus and a few new

species. Ann. Mag. nat. Hist. (10) 12 : 421-430, 4 figs.
i-933 e - I n Sicard, A., Mader, L. & Arrow, G. J., Resultats scientifiques du voyage aux

Indes Orientales Neerlandaises LL. AA. RR. le Prince et la Princesse Leopold de Belgique,

Coccinellidae. Mem. Mus. r. Hist. nat. Belg. (Hors serie) 4 (9) : 51-57.
19340- Schwedische-Chinesische wissenschaftliche Expedition nach des Nordwestlichen

Provinzen Chinas. Coleoptera, Melolonthinae. Ark. Zool. 27(A) 19 : 7-10.

- 19350. A note on the genus Narycius and a few other genera of Cetoniid Coleoptera.
Sty lops 4 : 101.

!9350- A contribution to the classification of the coleopterous family Lucanidae. Trans.

R. ent. Soc. Lond. 83 : 105-125, pi. 4, 5 figs.
I 935 C - The undesirability of establishing genera based on characters not common to

both sexes and of the naming of subgenera. Proc. R. ent. Soc. Lond. 10 : 33-34.

I 935^- A note on the genus Rhyparus (Coleoptera, Aphodiidae), and descriptions of
three new species. Ann. Mag. nat. Hist. (10) 16 : 157-160.

19360. The beetles belong to the Lamellicorn genus Chiron. Ann. Mag. nat. Hist.

(10) 17 : 150-153.

19366. Three genera of Erotylid Coleoptera new to the African fauna. Novit. zoo/.
39 : 253-256.

19360. Notes on some Lamellicorn beetles from South and East Africa, with descriptions

of new species. Novit. zool. 39 : 257-260.

19360*. A few new African species of Endomychidae (Coleoptera) in the British Museum.
Ann. Mag. nat. Hist. (10) 18 : 373-378.

- 19360. A beetle in a bedstead. Entomologist's mon. Mag. 72 : 229.

19370. Systematic notes on beetles of the subfamily Dynastinae, with descrptions of a
few new species in the British Museum collection (Coleoptera). Trans. R. ent. Soc. Lond.

86 : 35-58. Pi- L

19376. Notes on some Clavicorn Coleoptera and descriptions of a few new species and
genera. Ann. Mag. nat. Hist. (10) 20 : 101-113.

19370. Dimorphism in the males of stag-beetles (Coleoptera, Lucanidae). Trans. R.
ent. Soc. Lond. 86 : 239-245, pis 1-3.

19370*. Scarabaeidae : Dynastinae. Coleoptm Cat. pars 156, 124 pp.

19380. Some notes on stag-beetles (Lucanidae) and descriptions of a few new species.
Ann.. Mag. nat. Hist, (n) 2 : 49-63, pi. 4.

19386. The coleopterous genus Anthypna (Glaphyridae) . Ann. Mag. nat. Hist, (n)

2 : 285-290.

- 19386. Entomological results from the Swedish expedition, 1934, to Burma and British
India. Coleoptera, Cetoniidae collected by R. Malaise in Burma. Ark. Zool. 30(B)
14 : 1-4.

19380". Notes on some Melolonthine Coleoptera from the Malay Peninsula, and descrip-
tions of a few new species. /. fed. Malay St. Mus. 18 : 267-278.

- 19380. Some new species of Melolonthine beetles from Borneo. /. fed. Malay St. Mus.
18 : 319-326.

19390. Dr C. J. Gahan - Obituary. Entomologist 72 : 25-26, pi. i.

- 19396. Dr C. J. Gahan - Obituary. Nature 143 : 401.

- 19390. Three new species of Lamellicorn beetles from the Caroline Islands. Entomo-
logist's mon. Mag. 75 : 84-87.

- 19390". Walther Horn - Obituary. Entomologist's mon. Mag. 75 : 204-205.

- 19390. Notes on the coleopterous family Languriidae and descriptions of a few new
African species. Proc. R. ent. Soc. Lond. (B) 8 : 200-203.

I939/. The Lucanid Coleoptera of the Caroline Islands. Ann. Mag. nat. Hist, (n)

4 : 579-582, pi. 4.

CETONIINAE DESCRIBED BY G. J. ARROW 43

ARROW, G. J. 1939^. Dimorphism in the males of Coleoptera. Proc. R. ent. Soc. Lond. (A) 14:

113-114.
19400. A nomenclatural note. Proc. R. ent. Soc. Lond (B) 9 : 16.

- 19406. Entomological results from the Swedish expedition 1934 to Burma and British
India. Coleoptera, Clavicornia. Erotylidae, Languriidae, Endomychidae, Discolomidae
and Georyssidae collected by R. Malaise. Ark. Zool. 31 (A) 17 (1939) : 1-9.

- 1940^. A new genus of blind beetles of the family Lucanidae (Col.). Proc. R. ent. Soc.
Lond. (B) 9 : 93-96, n figs.

19400*. A note on certain genera of Brenthid Coleoptera and description of a new species.

Ann. Mag. nat. Hist, (n) 6 : 318-320.

19410. Some undescribed species of Melolonthid Coleoptera from western New Guinea
and the adjacent islands of Waigeu and Japen. Ann. Mag. nat. Hist, (n) 7 : 448-464.

19416. A few new species of the genus Onthophagus (Coleoptera, Copridae) from New

Guinea and Japen Island. Ann. Mag. nat. Hist, (n) 8 : 48-55.

1 94 ic. Systematic notes on the Cetoniid Coleoptera and descriptions of a few new

species. Ann. Mag. nat. Hist, (n) 8 : 73-88.

1 94 id. Entomological results from the Swedish expedition 1934 to Burma and British

India. Coleoptera, Melolonthidae. Ark. zool. 33 (A) 8 : 1-8.

19410. Systematic notes on beetles of the family Dynastidae and descriptions of a few

new species. Ann. Mag. nat. Hist, (n) 8 : 273-283.
19420. Can indistinguishable specimens be different species? Entomologist 75 : 169-

171.

19426. The origin of stridulation in beetles. Pvoc. R. ent. Soc. Lond. (A) 17 : 83-86.

I942C. The beetle family Rhysodidae, with some new species and a key to those at

present known. Proc. R. ent. Soc. Lond. (B) 11 : 171-183.

- 19420". A few new species of the Scarabaeid subfamily Hybosoridae (Coleoptera), with a
key to the genus Phaeochroops. Ann. Mag. nat. Hist, (n) 9 : 923-928.

- i943a. The Endomychid Coleoptera of New Guinea and the neighbouring islands, with
some new species. Ann. Mag. nat. Hist, (n) 10 : 128-136.

19436. Two remarkable new genera of Erotylid Coleoptera. Ann. Mag. nat. Hist.

(n) 10 : 389-393. ^ figs.

- I943C. A new mimetic genus of Erotylid Coleoptera. Ann. Mag. nat. Hist, (n) 10 : 394-
397, 2 figs.

19430". On the genera and nomenclature of the Lucanid Coleoptera, and descriptions of a

few new species. Proc. R. ent. Soc. Lond. (B) 12 : 133-143.

- 19430. Systematic notes on the Melolonthine beetles belonging to the genus Lepidiota
and some related genera. Ann. Mag. nat. Hist, (n) 10 : 773-785.

- 19440. Polymorphism in giant beetles. Proc. zool. Soc. Lond. (A) 113 (1943) : 113-116,
pi. i.

19446. The Asiatic beetles of the Melolonthine genus Schizonycha. Ann. Mag. nat.

Hist, (n) 11 : 194-200, 6 figs.
I944C. On Erotylid beetles belonging to Spondotriplax and some allied genera, with

descriptions of a few new species (Coleoptera). Proc. R. ent. Soc. Lond. (B) 13 : 5357.

- 19440*. Reduction of segmentation in the Coleoptera. Proc. R. ent. Soc. Lond. (A)
19 : 107-108.

19440. Systematic notes on Melolonthine beetles belonging to Holotrichia and related

genera. Ann. Mag. nat. Hist, (n) 11 : 631-648.
I944/. The beetles of the Lamellicorn subfamily Valginae, with a synopsis of the genera

and descriptions of some new species. Trans. R. ent. Soc. Lond. 94 : 225-246.
19450. On the Melolonthine beetles of the genus Hemiserica, including a few new species.

Ann. Mag. nat. Hist, (u) 12 : 117-121.
19456. Systematic notes on a few genera of Erotylid Coleoptera. Proc. R. ent. Soc.

Lond. (B) 14 : 117-118.

44 M. E. BACCHUS

ARROW, G. J. ig^6a. A new giant species of Ruteline Coleoptera. Proc. R. ent. Soc. Lond.
(B) 15 : 49-5. I fi g-

- 19466. Entomological results from the Swedish expedition 1934 to Burma and British
India. Coleoptera, Melolonthidae collected by Rene Malaise. Ark. Zool. 38 (A) 9 : 1-33,
i fig., 2 pis.

I94&C. Synonymic notes on some Oriental Cetoniine beetles, with descriptions of two new

species. Proc. R. ent. Soc. Lond. (B) 15 : 140-144, 8 figs.

- i947. Notes on Aserica and some related genera of Melolonthine beetles, with des-
criptions of new species and two new genera. Ann. Mag. nat. Hist, (n) 13 : 264-283.

19476. A few notes on West Indian Dynastine beetles and descriptions of two new
species. Ann. Mag. nat. Hist, (n) 14 : 221224.

19480. The Melolonthine beetles of the Island of Mauritius, with a key to the genera
and species. Proc. R. ent. Soc. Lond. (B) 17 : 25-34, 1 1 n g s -

- 19486. Further notes on the beetle genus Lachnosterna (Coleoptera, Melolonthinae),
with descriptions of three new genera. Proc. R. ent. Soc. Lond. (B) 17 : 49-54.

i948c. A note on the Indian beetle genus Narycius. Ann. Mag. nat. Hist. (12) 1 : 71-72.

1948^. A horned Melolonthine beetle from South America. Ann. Mag. nat. Hist.
(12) 1 : 371-375. 2 figs.

19485. A few new African species of Endomychid Coleoptera. Bull. Annls Soc. R. ent.

Belg. 84 : 46-56.
1949. Fauna BY. India. Coleoptera, Lamellicornia part 4 (Lucanidae and Passalidae.

xi -f 274 pp., 23 pi.

1951. Horned beetles, a study of the fantastic in nature. 154 pp., 15 pis. The Hague.

M. E. BACCHUS

Department of Entomology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON, SW7 560

ENTOMOLOGY SUPPLEMENTS

3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177:
18 plates, 270 text-figures. August, 1965. 4.20.

4. 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-

6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso-
philidae. Pp. 129: 328 text-figures. May, 1966. 3.

7. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967.

3.15.

8. 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.

9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species
(Lepidoptera: Rhopalocera). Pp. 509. 8.50. Reprinted 1972.

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho-
palocera). Pp. 322: 348 text-figures. August, 1967. 8.

11. MOUND, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 172:
82 text-figures. May, 1968. 4.

12. 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.

13. 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.

14. CROSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and
its Islands. Pp. 198: I plate, 331 text-figures. July, 1969. 4.75.

15. ELIOT, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera:
Nymphalidae). Pp. 155: 3 plates, 101 text-figures. September, 1969. 4.

16. GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe
(Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969.

17. WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198:
68 plates, 15 text-figures. October, 1971. 12.

18. SANDS, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae).
Pp. 244: 9 plates, 661 text-figures. July, 1972. 9.90.

19. 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.

20. VON HAYEK, C. M. F. A Reclassification of the Subfamily Agrypninae
(Coleoptera : Elateridae). Pp. 309: 17 text- figures. October, 1973. 12.30.

21. CROSSKEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia,
including keys to the supraspecific taxa and taxonomic and host catalogues.
Pp. 221: 95 text-figures. December, 1973. 9.55.

PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS OLD WOKING SURREY ENGLAND

-8 NOV1974

A REVISION OF

ATOPOPOMPILUS ARNOLD, WITH

A NOTE ON THE IDENTITY

OF ANOPLINELLUS BANKS

(HYMENOPTERA : POMPILIDAE)

M. C. DAY

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 31 No. 3

LONDON : 1974

NOVI974

A REVISION OF

ATOPOPOMPILUS ARNOLD, WITH A NOTE

ON THE IDENTITY OF ANOPLINELLUS BANKS

(HYMENOPTERA : POMPILIDAE)

BY

MICHAEL CHARLES DAY

Pp. 45-70; 34 Text-figures; 2 Maps

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 31 No. 3

LONDON : 1974

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 Vol. 31 No. 3 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.)

Trustees of the British Museum (Natural History), 1974

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 7 November, 1974 Price 1.55

A REVISION OF

ATOPOPOMPILUS ARNOLD, WITH A NOTE

ON THE IDENTITY OF ANOPLINELLUS BANKS

(HYMENOPTERA : POMPILIDAE)

By M. C. DAY

CONTENTS

Page
SYNOPSIS .... ...... 47

INTRODUCTION

ACKNOWLEDGEMENTS

THE IDENTITY OF A noplinellus BANKS

GENUS Atopopompilus ARNOLD .
Key to the species
Descriptions of the species

REFERENCES ....

INDEX

47

49

AND ITS TYPE-SPECIES . . 49

51

52

54

68

69

SYNOPSIS

Anoplinellus Banks, a genus based on a misidentified type-species, is disposed of as a synonym
of Arachnophroctonus Howard. Atopopompilus Arnold is recalled from synonymy and redefined
to include Ethiopian, Malagasy and Oriental species; the males of (all species are recognized.
Six species are revised and a seventh described as new; a key and distributional data are given.
Lectotypes are designated for four nominal species, and type-material of a fifth is presumed lost;
eleven new specific synonyms are established.

INTRODUCTION

VARIOUS groups of Pompilidae exhibit quite substantial sexual dimorphism, and
this phenomenon has, on occasion, given rise to considerable taxonomic confusion.
New taxa have been established for species or groups of species recognized only
from one sex, the other remaining undescribed, or being assigned to other taxa.
The genus here revised, Atopopompilus Arnold, 1937, is an example of such confusion,
as will be clear from this introduction.

Haupt (1929), in his treatment of the genus Paracyphononyx Gribodo, 1884,
included species of two distinct groups, as is shown by the immediate dichotomy
in his keys to females and to males. Arnold (1936) revised the Ethiopian species
of Paracyphononyx, including the males of both groups keyed by Haupt, but the
females only of one group, those most closely related to P. melanicrus Gribodo,

48 M. C. DAY

1884, type-species of Paracyphononyx. In 1937, Arnold proposed Atopopompilus
for the second group of females. However, males of one group which character-
istically have a median carina on the face between and below the antennae, had
been associated by Haupt with females of three species, P. nefas Dalla Torre, 1897,
P. coloratus Haupt, 1929, and P. venans Kohl, 1894, the last being type-species of
Atopopompilus. In his revision of Paracyphononyx, Arnold (1936) had recognized
Haupt's carinate-faced male, P. affinis Haupt, 1929, and had placed Haupt's other
carinate-faced species in the synonymy of Pompilus carinatus Radoszkowski, 1881.
In 1937, Arnold stated 'male unknown' after his diagnosis of Atopopompilus.

Atopopompilus as here discussed includes five Ethiopian and one Malagasy
species; a related species occurs in the Oriental and Indonesian regions. Banks
(1934) proposed Anoplinellus as a genus for this Oriental species, known to him only
from females which he misidentified as Pompilus clotho Smith, 1879. Van der
Vecht (unpublished) correctly associated this female with its male in long series of
specimens from Java, identified as the subsequently described Anoplinellus javanus
Haupt, 1935. The males have a carinate face. A female and male, determined by
Van der Vecht, are in the collections of the National Museum of Rhodesia as a result
of exchanges between Arnold and Vecht. However, Arnold did not associate the
female with Atopopompilus, or the male with 'carinate-faced Paracyphononyx'.

Haupt (1950) re-assessed Atopopompilus and gave a key to the females of the
species he recognized. He again referred to a male of A. venans, in cursive text,
but did not give sufficient information to enable other workers readily to identify it.

Evans (1966), in his re visional study of North and Central American Pompilinae,
listed Atopopompilus as a synonym of Paracyphononyx, and suggested that
Atopopompilus might ultimately be used as a subgenus within the latter genus.
However, Evans based his knowledge of Atopopompilus only on female specimens;
his diagnosis of Paracyphononyx does not include carinate-faced males. Neo-
tropical species placed in Atopopompilus by Banks (1947) are here regarded as
members of Paracyphononyx, following Evans (1966).

After study of reasonable series of specimens and their geographical distribution,
and an antero-postero gynandromorph individual, the association of sexes originally
made by Haupt can now be confirmed unreservedly. The carinate-faced males are
here recognized as the true males of Atopopompilus. It is the purpose of this paper
to revise the species of this Palaeotropical genus, here recalled from the synonymy
of Paracyphononyx. The identity of Anoplinellus is examined, and the name
disposed of as a synonym of the genus to which its type-species belongs.

I continue to follow the terminology of Evans (1966) ; abbreviations here employed
are as follows:

LID : lower interocular distance (distance between eyes below antennae)

MID : middle interocular distance (distance between eyes across front)

SMC: submarginal cell

TFD : transfacial distance (width of head)

UID: upper interocular distance (distance between eyes on vertex).

Depositories in which material studied is housed are as follows; abbreviations for
these institutions used in the text are also here listed.

REVISION OF A TOPOPOMPILUS 49

AM Albany Museum, Grahamstown, Cape Province, Republic of South Africa.

BMNH British Museum (Natural History), London.

IRSNB Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium.

ISM Institut Scientifique, Madagascar.

MB Museu Bocage, Lisbon, Portugal.

MCZ Museum of Comparative Zoology, Boston, U.S.A.

MHN Museum d'Histoire Naturelle, Geneva, Switzerland.

MNHN Museum National d'Histoire Naturelle, Paris, France.

MNHU Museum fiir Naturkunde der Humboldt-Universitat, Berlin, East Germany.

MRAC Musee Royal de 1'Afrique Centrale, Tervuren, Belgium.

NM Naturhistorisches Museum, Basle, Switzerland.

NMR National Museum of Rhodesia, Bulawayo, Rhodesia.

NR Naturhistoriska Riksmuseum, Stockholm, Sweden.

TM Transvaal Museum, Pretoria, Transvaal, Republic of South Africa.

UM University Museum, Oxford, United Kingdom.

USNM United States National Museum, Washington, U.S.A.

ZM Phyletisches Museum, Halle an der Saale, East Germany.

coll. Wahis Private collection of Monsieur R. Wahis, Chaudfontaine, Belgium.

ACKNOWLEDGEMENTS

I am indebted to the following staff of other institutions who have arranged loans
of type or other material: Dr C. Baroni Urbani (NM), Dr C. Besuchet (MHN),
Dr J. Decelle (MRAC), Prof. H. E. Evans (MCZ), Dr G. Fagel (IRSNB), Dr J. de A.
Fernandes (MB), Mr F. W. Gess (AM), Prof. J. O. Husing (ZM), Dr S. Kelner-Pillault
(MNHN), Dr E. Konigsmann (MNHU), Dr A. S. Menke (USNM), Mr F. C. de
Moor (NMR), Mr C. O'Toole (UM), Dr P. I. Persson (NR), Mr J. van Reenan (TM).
I am indebted to: Monsieur R. Wahis, both for helpful correspondence and for the
loan of specimens; Prof. H. E. Evans has discussed the identity of P. clotho with
me.

THE IDENTITY OF ANOPLINELLUS BANKS, AND ITS TYPE-SPECIES

Banks (1934) proposed Anoplinellus for females of a species of Pompilidae from
the Philippines and Indonesia which he misidentified as Pompilus clotho Smith,
1879, which species is type by monotypy. Female specimens bearing Banks's
labels have been examined, and are here identified as conspecific with Pompilus
daedalus Bingham, 1896, a species here regarded as congeneric with Pompilus
venans Kohl, 1894, type-species of Atopopompilus Arnold. The material consists
of the following specimens: a female from Iligan, Mindanao (coll. Baker) in the
USNM, bearing a label 'Anoplinellus clotho Sm.'; a female in MCZ from Sandakan,
Borneo (coll. Baker) bearing a label 'Anoplius clotho Sm.'; and two females from
Celebes (Sulawesi) (coll. C. F. Clagg). One of the Celebes females bears a Banks
manuscript name in Anoplinellus; he referred to this undescribed material in 1934.

P. clotho Smith was described from a single female specimen. This holotype is in
the BMNH, and agrees well with the description; it is a species of Anoplius Dufour.
It bears a label in Smith's hand, 'Pompilus clotho type Sm.', and a second small
white circular label has written on one side the figures '54. 76', and 'Sumatra' on the

50 M. C. DAY

reverse. The figures refer to an entry in the accessions register of BMNH dated
1854. The data recorded indicate that the specimen came to the museum as part
of a collection disposed of by the Zoological Society of London through the London
dealer, Stevens. This collection included material gathered from several sources;
some from Sumatra, collected for Sir Stamford Raffles, and some from 'San Domingo'
collected by Hearne; also material from Brazil, Colombia, 'Niger', Trebizond and
Madagascar. The holotype of P. clotho is labelled as part of the material collected
in Sumatra. However, I know of no other Oriental or Indonesian specimens similar
to the holotype. The specimen, which has lost parts of some legs and an antenna,
and has its pubescence much rubbed, nevertheless is immediately noteworthy in
that it presents the characteristic facies of many Nearctic species of Pompilidae.

The type of P. clotho is in my opinion consubgeneric with the type-species of
Arachnophroctonus Howard, a subgenus of Anoplius. The species of Arachnophroc-
tonus known to me from the East Indies are markedly different in coloration, and
readily separated morphologically, from P. clotho. In Evans's (1951; 1966) keys
to species of North and Central American Arachnophroctonus, the specimen keys out
as A. relativus Fox, 1893, recorded by Evans as a 'somewhat variable species'. The
holotype of P. clotho (bearing in mind its poor condition) is closely similar to speci-
mens of A . relativus.

Since other material in the collection obtained by the BMNH from Stevens came
from San Domingo rather than Sumatra, there is a reasonable possibility that this
type-specimen could have been collected in the West Indies or Central America.
San Domingo is normally construed as referring to the present-day Dominican
Republic. It is conceivable but unlikely that some other locality of this name on
the Central American mainland was intended. H. E. Evans (personal communica-
tion) informs me that A. relativus is not yet known to occur in the West Indies.
However, he has seen males of a closely related undescribed species from the
Dominican Republic.

It would seem likely that the holotype could have been mislabelled 'Sumatra'
when incorporated into the BMNH collections; certainly, the type-locality should be
regarded as doubtful. However, three possibilities should be considered:

(a) P. clotho may be an Indonesian species of Anoplius, known only from the
type-specimen;

(b) P. clotho may be a West Indian species of Anoplius;

(c) P. clotho may be a senior synonym of A . relativus.

On the balance of present evidence, the second possibility seems most probable.
It is not feasible here completely to resolve the problem since our knowledge of the
pompilid faunas of East and West Indies is so imperfect.

Haupt (1935) described Anoplinellus javanus from two female specimens collected
in Java. He recognized the genus Anoplinellus as described by Banks, and stated
'male unknown'. No authors have subsequently used the name Anoplinellus, save
in a catalogue of the genera of Pompilidae (Pate, 1946).

Although proposed at a later date than Anoplinellus, Atopopompilus has enjoyed
more frequent use (Arnold, 1937; 1951; Haupt, 1950; de Saeger, 1945) and has had

REVISION OF A TOPOPOMPILUS 51

more nominal species referred to it. The type-species has always been correctly
identified. In view of the more limited use of Anoplinellus, the misidentification of
its type-species, and the doubt surrounding the provenance of the type-specimen of
P. clotho, I propose to place Anoplinellus in the synonymy of Arachnophroctonus,
with the type-species of which P. clotho is consubgeneric. The nomenclatural
changes are formalized below.

Subgenus Arachnophroctonus Howard

Arachnophroctonus Howard, 1901: pi. 7, fig. n. Type-species: Sphex tropicus Linnaeus sensu
Fabricius, 1775 (misidentification) [ Psammochares marginalis Banks, 1910], by subsequent
designation (Pate, 1946 : 129).

Anoplinellus Banks, 1934 : 84. Type-species: Pompilus clotho Smith, 1879, by monotypy.
Syn. n.

Anoplius (Arachnophroctonus} clotho (Smith) comb. n.

Pompilus clotho Smith, 1879 : 146. Holotype $, provenance uncertain: labelled 'Sumatra',
but more probably DOMINICAN REPUBLIC (BMNH) [examined].

Atopopompilus Arnold is thus the only available name for the genus here revised.
Genus ATOPOPOMPILUS Arnold stat. rev.

{Anoplinellus Banks, 1934 : 84. Misidentification, see above]

Atopopompilus Arnold, 1937 : 22. Type-species: Pompilus venans Kohl, 1894 C= Pompilus
carinatus Radoszkowski, 1881], by original designation.

$ ^. Length 7-23 mm. Body colour black; females of some species with more or less
red-brown coloration on head, thorax or legs, more rarely light red-brown ; antennae sometimes
red-brown to orange; palpi frequently light red-brown. Males as above, but frequently with a
yellow line on posterior pronotal margin, often extensively yellow on face, dorsal surface of
hind tibia, and often on tarsi and seventh tergite. Female wings infuscate, darker at margin;
male, hyaline with infuscate margin or fusco-hyaline with darker margin. Both sexes with
more or less grey or white pubescence on face, thorax, coxae and abdomen. Female with
sparse, long, thin erect hairs on head, thorax, coxae and abdomen, each arising from a more or
less distinct small pit; less noticeable on dorsum of thorax and abdomen. Male often less
hirsute, but hairs frequently white, more noticeable. Female post-scutellum and propodaeum
often with numerous short thick, dark erect hairs also, which may appear thickened or adpressed,
and sometimes reflect violaceous. Male post-scutellum and posterior half or third of pro-
podaeum with very dense, long, erect white pubescence, perpendicular to insertion or at an
angle. Posterior tergite and sternites of female with more or less marked strong erect hairs.

Head thin in lateral view (Text-figs i, 3), inserted low on thorax; vertex abrupt, not gently
rounded; labrum exposed, semicircular or truncate; mandibles short, normally not exceeding
opposite edge of labrum when closed, with a single tooth on inner margin. Malar space present,
not exceeding half thickness of first flagellar segment. Female face with a small interantennal
tubercle, sometimes with a faint trace of a carinate edge continuing just below the antennal
insertions. Male face with a more or less strong facial carina, from middle of face passing
between antennal insertions, normally to reach clypeus, where it may divide in two and become
lost (Text-figs 29-34). Female antennae normally elongate, thin, but rarely with shorter,
thicker antennae. Male antennae strongly crenulate. Occiput concave transversely, vertex
well defined, never carinate. Temples almost obsolete. Thorax relatively massive, pronotum
narrowed in front, short, with more or less arcuate posterior margin. Postnotum short, very

52 M. C. DAY

slightly expanded on either side of the median line. Female propodaeum frequently with
more or less well denned sloping posterior declivity, sometimes flattened also on lateral corners ;
rarely, propodaeum rounded. Declivity occasionally concave, sometimes with very fine
aciculae centrally; one species with distinct tubercles produced laterally. Male propodaeum
usually with low lateral profile, rounded. Legs elongate, strongly spinose, but female without
tarsal comb; foremetatarsus with two short lateral spines in addition to terminal spine.
Ultimate tarsal segments lack spines beneath, claws strongly bifid, with a broad, truncate rear
tooth. Apical segment of male foretarsus unmodified with claws asymmetric, inner claw more
curved, tooth stout and blunt. Female abdomen somewhat compressed apically, last sternite
strongly so with well-marked blunt, polished ventral carina at least terminally. Forewings
with SMC2 approximately equal to SMC3, latter often narrowed above. Radial vein beyond
SMC's with characteristic curve (Text-fig. 9). Male genitalia with basal booklets single.
Parameres only just exceeding other appendages, very thin, with more or less long hairs.
Subgenital plate with more or less triangular termination, sometimes truncate at tip, hairy but
without thick spines or long hairs on posterior margin (Text-fig. 23).

Females may readily be recognized by the combination of bifid claws, malar
space, and the compressed posterior abdominal segments with a short ventral carina
terminally. The general habitus, especially the high, broad thorax and thin head,
render the genus particularly recognizable. The males, with the conspicuous facial
carina and crenulate antennae, can be confused only with Batozonellus Arnold or
Paracyphononyx in the Old World. The former may be recognized by the char-
acteristic venation, broad face, and arcuate expansion of the postnotum either side
of the midline. Males of the latter genus lack the carinate face.

DISTRIBUTION. Forest, woodland and savanna areas of the Old World tropics
including Ethiopian and Malagasy, Oriental and Indonesian regions.

BIOLOGY. Unknown. Morphology indicates a non-fossorial habit. This genus
may utilize pre-existing nest-cavities, or leave prey in situ at site of attack.

INCLUDED SPECIES. Five Ethiopian, one Malagasy, and one Oriental species are
here regarded as members of Atopopompilus . The Ethiopian species form a more
closeknit group, the two remaining species being each somewhat distinctive;
however, I do not feel that any infrageneric grouping is necessary.

Although Atopopompilus has much in common with Paracyphononyx, it is a very
distinctive and comparatively homogeneous group of species. It is not possible at
this stage to state with any certainty whether the similarities of the two genera
are due to convergence or to close relationship. It seems most convenient, in the
absense of biological information, to regard Atopopompilus as a genus perhaps
standing in a similar relationship to Paracyphononyx as does Batozonellus to Episyron
Schiodte. Its status may better be reassessed when the more diverse and wide-
spread Paracyphononyx has been adequately revised.

KEY TO THE SPECIES OF ATOPOPOMPILUS

FEMALES

i Propodaeum with well-developed lateral tubercles (Text-fig. 16). Black, with more
or less red-brown anteriorly, on head and thorax. Madagascar.

nefas (Dalla Torre) (p. 63)

REVISION OF A TOPOPOMPILUS

53

FIGS 1-16. 1-8. Female head: i. A. carinatus, side view. 2. Facial view. 3. A.
daedalus, side view. 4. Facial view. 5-8. Facial views. 5. A. nasutus. 6. A. kili-
mandjaroensis. 7. A. nefas. 8. A. jacens. 9. Submarginal cells of fore wing, A.
carinatus, female. 10. Female antenna, A. crassicornis. n 16: Female propodaeum,
lateral profile, n. A. carinatus. 12. A. kilimandjaroensis. 13. A. jacens. 14. A.
daedalus. 15. A. nasutus. 16. A. nefas.

54 M. C. DAY

Propodaeum without lateral tubercles. If with red-brown head and prothorax, then

antennae also so coloured .......... 2

2 Head short, broad, flat in the face (Text-figs 3, 4). Oriental region

daedalus (Bingham) (p. 66)

- Head less short, face more convex (Text-figs i, 2, 5, 6, 8). Ethiopian region . . 3

3 Unicolorous black, or black with very dark brown appendages; often with a little

red-brown ventrally on antennae ......... 4

- Black with antennae substantially red-brown or orange, sometimes legs and/or head

and prothorax red-brown .......... 5

4 Second and subsequent few flagellar segments at least 4-0 x as long as wide.

carinatus (Radoszkowski) (p. 54)

- Second and subsequent few flagellar segments not more than 3-0 X as long as wide

(Text-fig. 10) crassicornis sp. n. (p. 57)

5 Prothorax red-brown or yellow-brown .... jacens (Bingham) (p. 61)

- Prothorax black ............ 6

6 Antennae orange, sometimes darker basally; face almost maroon, area about ocelli

black (Text-fig. 6). Propodaeum massive, with marked declivity (Text-fig. 12)

kilimandjaroensis (Cameron) (p. 59)

- Antennae and legs, clypeus and lower part of face red- or orange-brown (Text-fig. 5).

Propodaeum rounded (Text-fig. 5) nasutus (Haupt) (p. 58)

MALES

Propodaeum excised behind ; wings lightly infuscate, head and anterior part of thorax

red-brown, otherwise black. Madagascar . . . nefas (Dalla Torre) (p. 63)

Propodaeum not excised behind: wings fusco-hyaline, or hyaline with infuscate

outer margins. Body black, with some yellow colour; if with red-brown on

thorax, then legs or antennae so coloured. Oriental and Ethiopian regions . 2

Wings fuscohyaline with tips infuscate, body colour black with some yellow on face,

prothorax, hind tibiae and seventh tergite ....... 3

Wings fuscohyaline, or hyaline with infuscate tips. Body colour black and yellow,

also with some red-brown at least on appendages, often also on head and thorax 5

Facial carina not distinct on clypeus (Text-figs 29, 30). Oriental region. Antennae

as in Text-fig. 24 daedalus (Bingham) (p. 66)

Facial carina distinct on clypeus, forked to give two lateral arms (Text-figs 31-33).

Ethiopian region ........... 4

Antennal crenulations angulate ventrally in profile (Text-fig. 25)

carinatus (Radoszkowski) (p. 54)

Antennal crenulations curved ventrally in profile (Text-fig. 27) crassicornis sp. n. (p. 57)
Antennal crenulations angulate ventrally (Text-figs 25, 28) ..... 6

Antennal crenulations curved ventrally (Text-fig. 27)

kilimandjaroensis (Cameron) (p. 59)
Forewing hyaline with infuscate tip; antennae shorter, stouter, with less curvature

dorsally on each segment (Text-fig. 28) . . . . jacens (Bingham) (p. 61)
Forewing fuscohyaline with infuscate tip ; antennae more elongate, with more distinct

segmental curvature dorsally (Text-fig. 25) . . nasutus (Haupt) (p. 58)

Atopopompilus carinatus (Radoszkowski) comb. n.
(Text-figs i, 2, 9, n, 17, 23, 25, 31, 32)

Pompilus carinatus Radoszkowski, 1881 : 212. Holotype ^, ANGOLA (MB, Lisbon) [examined].

REVISION OF A TOPOPOMPILUS 55

Pompilus venans Kohl, 1894 : 3 X 5- Holotype $, SIERRA LEONE (TM, Pretoria) [examined].

Syn. n.

Paracyphononyx venans (Kohl) Haupt, 1929 : 168; $, $.
Paracyphononyx affinis Haupt, 1929 : 170. Holotype , CAMEROUN (MNHU, Berlin)

[examined]. Syn. n.

Paracyphononyx affinis Haupt; Arnold, 1936 : 445; <J.
Atopopompilus venans (Kohl) Arnold, 1937 : 2 3I ?
Atopopompilus venans (Kohl); Haupt, 1950 : 57; $, <$.

$. Length 13-21 mm. Black; legs and antennae sometimes very dark brown; sensory
areas of antennae often red -brown. Face below antennae with some silvery pubescence,
otherwise pubescence more commonly brownish. Hairs of head and thorax relatively long
(often as long as thickness of scape), dense, often lost in worn specimens.

Antennae most usually elongate, but occasionally less so (second flagellar segment only 4 x
as long as thick). Front narrow, high (Text-fig. 2), proportions variable. Inner orbits converge
strongly above, converging or sub-parallel below. Thorax of form described for genus,
propodaeum with a pronounced declivity which frequently has a central area of very fine
aciculae; with long erect hairs and adpressed short hairs. SMC3 narrowed above, shorter on
radial vein than is SMC2.

(J. Length 7-15 mm. Black; border of inner orbits, tip of facial carina, lateral area of
clypeus, posterior pronotal margin, area on dorsal surface of hind tibia, yellow. At least some
reddish ventrally on basal antennal segments, some populations with much of antennae, labrum,
tip of clypeus, and part of legs dark or light red-brown. Wings fusco-hyaline with darker
margins. Body extensively silvery pubescent, with much erect silvery hair, particularly on
posterior third of propodaeum, less on anterior half of first tergite. Seventh tergite with a
small amount of whitish yellow colour in dark specimens, more extensively so in specimens with
some red-brown coloration.

Antennae (Text-fig. 25) markedly crenulate, each segment with a sharp ventral angle when
viewed in profile. Face with well marked carina extending onto clypeus. Inner orbit of eyes
converging above, sub-parallel, then diverging below (Text-fig. 32). Genitalia as in Text-fig. 17.

The holotype of P. carinatus stands under that name in the collections of MB,
Lisbon. It is in fair condition but bears no original labels. Radoszkowski's
description, though brief, is unmistakeable and applies well to the type-specimen.
However, Arnold applied the name to A. jacens (p. 61), despite the fact that the
description of P. carinatus does not mention any red coloration.

Haupt associated a male from Bonjongo with P. venans. This specimen, in the
collections of MNHU, Berlin, bears Haupt's determination label and a 'type' label
(i.e. so-called 'allotype' of P. venans Kohl). It has no such status. It is clearly
conspecific with the type of P. affinis, though the yellow coloration of the former
specimen is much reduced. It was on a basis of this colour difference that Haupt
separated the two in couplet 12 of his key to males (1929). The confusion amongst
the African species arises mainly from this inadequate key, and from Arnold's
consequent misidentifications.

DISTRIBUTION. The species is characteristic of the dense humid forest regions of
Central and West Africa; summarized on Map i, p. 60.

VARIATION. The antennae of females show much variation in length relative to
thickness. This correlates broadly with the relative proportions of the thorax and
legs, in that females with shorter, thicker antennae tend to have shorter, thicker
legs and a more massive thorax. The face of a female with attenuate antennae is

M. C. DAY

FIGS 17-34. 17-22. Male genitalia, left half, ventral view. 17. A. carinatus. 18. A.
nasutus. 19. A. kilimandjaroensis. 20. A. jacens. 21. A. nefas. 22. A. daedalus.
23. Subgenital plate, A. carinatus, male. 24-28. Male antennal flagellum, i.e. excluding
scape and pedicel. 24. A. daedalus. 25. A. carinatus. 26. A. nefas. 27. A. kiliman-
djaroensis. 28. A. jacens. 29-34. Male head. 29. A. daedalus, side view. 30. Facial
view. 31. A. carinatus, side view. 32. Facial view. 33-34- Facial views. 33. A.
crassicornis. 34. A. nefas.

REVISION OF A TOPOPOMPILUS 57

illustrated in Text-fig. 2; females of the more compact structure tend to have a
slightly broader face; it is difficult to segregate these females from those of A.
crassicornis sp. n. (see below). Males in West Africa have a tendency towards the
development of red-brown coloration on antennae and legs.

MATERIAL EXAMINED.

Pompilus carinatus Radoszkowski, holotype , ANGOLA: no further data
(Welwitsch) (MB, Lisbon). Pompilus venans Kohl, holotype 9 SIERRA LEONE:
25.viii.i892 (H. Brauns) (TM, Pretoria). Paracyphononyx affinis Haupt, holotype
(J, CAMEROUN: 'Joh. Albrechtshohe' (Conradt) (MNHU, Berlin).

ANGOLA: Salazar, field station of I. LA. A., on path through dense humid forest,
9-i5.iii.i972, 2 $ (D. Hollis); same, in Malaise trap, I <$; 7 miles west of Gabela, in
coffee plantation, swept from vegetation beside path, 17.^1.1972, I <$ (M. C. Day)
(BMNH Southern African Expedition, 1972) (BMNH). CAMEROUN: Lolodorf, 2 $
(Conradt); Bonjongo, 17.^.1873, I <$ (so-called 'allotype' of P. venans}; Victoria,
on flowers of Bidens pilosus, s.vii.iSgo, I $ (Preuss) (MNHU); R. Dja, 3i5' N.,
I33o' E., v-vii.J-936, I $ (F. G. Merfield) (BMNH). DAHOMEY: no further data,
1903, i $ (E. Poisson) (MNHN). GABON: Mts de Cristal, i5-3i.x.ig69, i <j> (A.
Villiers) (MNHN). GHANA: Ashanti, Obuasi, 1908, i $ (W. M. Graham); Aburi,
1400', xii. 1941, i c? (K. M. Guichard) (BMNH). NIGERIA: Olokemeji, near Ibadan,
i c? (Bridwell) (USNM); Ibadan, 4^.1922, TL $ (A. W . /. Pomeroy) (BMNH).
SIERRA LEONE: Njala, x. 1935, i $ (E. Hargreaves) (BMNH). TOGO: Bismarck Mts,
1890, i <j> (Buttner) (MNHU). UGANDA: Mabira Forest, Chagwe, 3500 '-3800',
i6-25.vii.i9ii, i $; W. shores of Lake Victoria, Buddu, 3700', 19-25. ix.ign, i <j>;
Entebbe, i-n.xi.ign, 2 $ (5. A. Neave); Mabira Forest, 3.vii.i9i3, i $ (C. C.
Gowdey) (BMNH). RWANDA: Bukavu (Costermansville), 1939, i $, i <$ (Hautmann);
1948, i <$ (P. H. Vercammen); Muhavura, 2100 m, 28.1.1953, i <$ (P. Basilewsky)
(MRAC). ZAIRE: Kibali-Ituri, Epulu, x. 1956, i $ (M. Poll); Kivu, Kavumu, 1951,
I $ (H. Bomans); Paulis, viii. 1947, i ? (P. L. G. Benoit); Bambesa, 20.ix.i933, i $
(H. J. Bredo); 20.X.I933, i $ (/. Leroy); viii-xii. 1937, 4 $ (/. Vrydagh); v. 1938, 3 $
(P. Henrard) (MRAC); Bambesa, ii.ix.ig37, i ? (/. Vrydagh) (IRSNB); Bambesa,
25.ix.i933, i $ (H. J. Bredo); Ubangi, Duma, lo.x.igio, i ? (H. Schubotz) (MNHU);
Bumba, 1940, i $ (H. de Saeger) (MRAC); Coquilatville, io.vii.ig36, i $ (R. P.
Hulstaert) (MNHN); Eala, xi. 1935, I <$ (J. Ghesquiere); Tshuapa, Bamanya, xi.
1964, i $ (R. P. Hulstaert); Sankuru, Komi, ii. 1930, i <$ (J. Ghesquiere} (MRAC);
Lualaba River, 5^.1907, i $ (S. A. Neave} (BMNH); Luluabourg, i $ (MHN);
Kasai, Kondue, i $ (E. Luja); Kasongo, ix. 1959, i $ (P. L. G. Benoit); Mutsors?,
1939, i $ (Hackars) (MRAC).

Atopopompilus crassicornis sp. n.

(Text-figs 10, 27, 33)

$. Length 11-14 mm. Black; morphologically very close to A. carinatus, but of more
compact structure. Face wider, thorax stouter, less hirsute, legs and antennae shorter, thicker

58 M. C. DAY

(antenna as in Text-fig. 10). Antennal segments tend towards oval section, second flagellar
segment not more than 3 X as long as greatest width.

<-. Length n mm. Black with yellow maculation on face and prothorax; morphologically
close to A. carinatus. Head broad, less high than in A. carinatus (Text-fig. 33). Antennal
segments curved beneath in profile, ventral sensory areas as those of A. kilimandjaroensis
(Text-fig. 27). However, dimensions of segments more nearly those of A. carinatus (Text-fig.
25).

Two females of this species from Uganda were collected together with typical
females of A . carinatus, and as a matter of direct comparison are clearly distinct both
in the more compact form of the thorax, and the short, thick antennae. However,
females from other parts of the range of A . carinatus tend to bridge the gap between
the extremes represented by these Ugandan females. On the basis of female
structure alone, I would hesitate to describe this taxon. However, certain char-
acters that I believe are those of the male are exhibited by a gynandromorph
specimen, of which all save the abdomen is male. I am confident that the structure
of the antennae and head are typical of the species it represents, and not modified
as a result of its mosaic genotype. The shape of the head, together with the curved
antennal crenulations and general coloration separate this specimen from all other
males I have seen. In particular, I have found antennal structure to give reliable
identification of males in this genus, and the nature of the antennal differences
observed in this specimen relative to the male of A . carinatus is entirely consistent
with the sorts of differences observed between other species. Geographical distri-
bution further supports the association of this gynandromorph with the females here
described.

DISTRIBUTION. East Africa; highland and lake area of the Great Rift Valley in
Uganda and Eastern Zaire: summarized on Map I, p. 60.

MATERIAL EXAMINED.

Holotype <j>, UGANDA: Entebbe, i-n.ix.ign (S. A. Neave) (BMNH).

Paratypes. UGANDA: west shores of Lake Victoria, Buddu, 3700', ig-as.ix.ign,
i $ (S. A. Neave) (BMNH). ZAIRE: Mt Hoyo, Ituri, 1250 m, 5.x. 1957, gynan-
dromorph (E. S. Ross & R. E. Leach] (coll. Wahis) ; Kibali-Ituri, Kilo, Mongbwalu,
1937, i $ (Harford-Jordens); Kilo, Mines, 1955, I $ (R. Andry) (MRAC).

Atopopompilus nasutus (Haupt) comb. n.
(Text-figs 5, 15, 18, 25)

Paracyphononyx nasutus Haupt, 1929 : 170. Holotype <$, TANZANIA (MNHU, Berlin)

[examined] .

{Atopopompilus venans (Kohl) sensu Arnold, 1937 : 2 4- ? colour variety. Misidentification.]
Atopopompilus bruneipes Haupt, 1950 : 57. Holotype $, ZAIRE (MRAC, Tervuren) [examined].

Syn. n.

$. Length 12-17 mm. Black; face below antennal insertions, clypeus, labrum, mandibles,
antennae, tibiae and tarsi orange-brown; sometimes also part or all of femora. Face below
and beside antennae with silvery pubescence, otherwise pubescence mostly brownish. Hairs
of head relatively long and dense as in A . carinatus.

REVISION OF A TOPOPOMPILUS 59

Antennae elongate. Inner orbits converging above, less strongly so below. SMC3 narrowed
above, half as long on radial vein as is SMCa. Propodaeum sloping gently (Text-fig. 15),
rounded in profile, with incipient adpressed hairs or very short fine erect hairs projecting
backwards, not vertical. Face as in Text-fig. 5.

<$. Length 8-12 mm. Black; clypeus, ventral surface of proximal antennal segments,
tibiae and tarsi red-brown; femora sometimes also so. Border of inner orbits, part of clypeus,
posterior pronotal margin, and dorsal surface of hind tibia, yellow. Seventh tergite whitish
yellow. Wings fusco-hyaline with darker margins. Otherwise morphologically very close to
A. carinatus. Genitalia as in Text-fig. 18.

Arnold placed P. nasutus in the synonomy of P. carinatus, by which name he
misidentified A . jacens.

I have seen only two males, only one of which has part of its antennae. The
proximal six segments are similar in form to those of A . carinatus (Text-fig. 25) .

DISTRIBUTION. East Africa; summarized on Map i, p. 60.

MATERIAL EXAMINED.

Paracyphononyx nasutus Haupt, holotype <^, TANZANIA: Usambara, Nguela,
(Rolle) (MNHU, Berlin). Atopopompilus bruneipes Haupt, holotype $, ZAIRE:
Pare National Albert, Rutshuru, io-24.vi.iQ34 (G. F. de Witte) (MRAC, Tervuren).

UGANDA: Entebbe, viii.iQii, 3 $ (C. C. Gowdey); Entebbe, i-ii.ix.ign, i $
(S. A. Neave); Kampala, I9~28.xii.i9i5, i $ (C. C. Gowdey); Tero Forest, S.E. of
Buddu, 3800', 20-30.ix.i9ii, i $ (5. A. Neave); Mubendi, 4^.1911, i $ (C. C.
Gowdey); 1918, no further data, i $ (C. C. Gowdey) (BMNH). ZAIRE: Rutshuru,
xi. 1937, i <$ (J. Ghesquiere) (MRAC).

Atopopompilus kilimandjaroensis (Cameron) comb. n.
(Text-figs 6, 12, 19, 27)

Pompilus kilimandjaroensis Cameron, 1910 : 252. Holotype $, TANZANIA (NR, Stockholm)

[examined].
Paracyphononyx personatus Haupt, 1929 : 170. Holotype <$, TANZANIA (MNHU, Berlin)

[examined]. Syn. n.

{Paracyphononyx parallelus Haupt sensu Arnold, 1936 : 445; $. Misidentification.]
Atopopompilus venans race mlanjiensis Arnold, 1937 : 24. Holotype $, MALAWI (BMNH)

[examined]. Syn. n.

$. Length 14-23 mm. Black; antennae, often including scapes, yellow-orange to orange-
brown. Head capsule dark red-brown, almost maroon; with varying amounts of black on face,
normally at least a horizontal band through anterior ocellus, sometimes reaching down to
antennal insertions and back to vertex. Pubescence mostly brownish. Erect hairs of head
and thorax shorter than in A. carinatus (mesopleural hairs shorter than thickness of second
flagellar segment) .

Antennal segments long, more than 4 X as long as wide. Inner orbits converging above, less
strongly so below (Text-fig. 6). SMC3 narrowed above, usually shorter on radial vein than is
SMC2. Propodaeum (Text-fig. 12) with abrupt and steep declivity, and posterior corners also
flattened at 45 degrees. Erect hairs long (almost thickness of scape), adpressed hair dense and
extensive.

(J. Length 9-12 mm. Black; mouthparts and clypeus, antennae and legs substantially
red-brown; part of clypeus, facial carina, borders of inner orbits, temples, posterior pronotal

6o

M. C. DAY

margin, spots on forelegs and dorsal surface of hind tibia, yellow. Last tergite white. Wings
hyaline with infuscate margins. Body hair extensive, silvery, pubescence silver or grey.
Propodaeum, save spiracular area, with extensive, dense silvery erect hair.

Morphologically very similar to A . carinatus, but antennal segments in lateral view rounded
beneath (Text-fig. 27). Genitalia as in Text-fig. 19.

Arnold placed P. personatus in the synonymy of his P. carinatus (A. jacens).
He also misidentified a female as P. parallelus, whilst excluding P. venans from
Paracyphononyx.

A. carinatus

A.crassicornis

A.nasutus

A. kilimandjaroensis

A. jacens

A.nefas

MAP i. Distribution of Atopopompilus species in the Ethiopian and Malagasy regions.

REVISION OF ATOPOPOMPILUS 61

DISTRIBUTION. Through much of the woodland and savanna areas of Africa;
summarized on Map i, p. 60.

VARIATION. Two females, from Bambesa and Moto, have a colour pattern very
similar to that of A. nasutus females. The form of the propodaeum, however, is
that of A. kilimandjaroensis. The colour pattern of the face of West African
populations is more extensively dark, closely similar to that of A . nasutus.

I have seen only three males of this species.

MATERIAL EXAMINED.

Pompilus kilimandjaroensis Cameron, holotype $, TANZANIA: Kilimandjaro,
Kibonoto, 1300-1900 m, H.v.i9o6 (Sjostedt) (NR, Stockholm). Paracyphononyx
personatus Haupt, holotype <, TANZANIA: Langenberg, v. 1898 (Fulleborn) (MNHU,
Berlin). Atopopompilus venans mlanjiensis Arnold, holotype $, MALAWI: Mlange,
6.1.1913 (S. A. Neave) (BMNH).

ANGOLA: no further data, i $ (Welwitsch) (BMNH). GHANA: Accra, 13^.1941,
i $ (K. M. Guichard); Tafo, 28.X.I97O, i $ (B. Boltori) (BMNH). IVORY COAST:
Assinie, 1886, i $ (C. Alluaud) (coll. Wahis). KENYA: Kwali Forest, 20 miles west
of Mombasa, 1.^.1948, i $ (M. Steele) (BMNH). MALAWI: Mt Mlange, xi. 1912-
xi. 1913, 24 $, i cJ; near mouth of Lusangazi river, i-3.ix.i9io, i $; valley of N.
Rukuru, Karonga district, 2000-4000', I5~i8.vii. 1910, i $ (S. A. Neave) (BMNH);
Mlanje, 2000', 20.1.1945, i $ (NMR). MOZAMBIQUE: foothills N. of Mt Chiperone,
I9.xi.i9i3, 2 $; Kola valley E. of Mt Chiperone, 2i.xi.i9i3, 2 $ (S. A. Neave)
(BMNH). SIERRA LEONE: no further data, i $ (BMNH). TANZANIA: Langenberg,
20.viii-i.ix.i898, i $ (Fulleborn) (MNHU). ZAIRE: Moto, 1922, i $ (L. Burgeon);
Bambesa, io.iv.i937, i $ (/. Vrydagh); Sankuru, M'Pemba Zeo, I3.xii.i959, i $
(D. R. Marechal); Kabinda, 1935, i $ (P. Henrard); Lulua, Kapanga, x. 1932, i $
(F. G. Overlaet); Katanga, Lubombo, ix. 1928, i $ (C. Seydel)', Kasongo, ix. 1959, i ?
(P. L. G. Benoit) (MRAC); Elizabethville, iv. 1935, i $ (F. de Loose)(NUR).

Atopopompilus jacens (Bingham) comb. n.
(Text-figs 8, 13, 20, 28)

Pompilus jacens Bingham, 1912 : 560. Holotype ^ (not $ as originally stated), RHODESIA

(UM, Oxford) [examined].
Paracyphononyx parallelus Haupt, 1929 : 168. Holotype $, SOUTH AFRICA (MNHU, Berlin)

[examined]. Syn. n.
Paracyphononyx coloratus Haupt, 1929 : 168. LECTOTYPE $, CAMEROUN (MNHU, Berlin),

here designated [examined]. Syn. n.

{Paracyphononyx carinatus (Radoszkowski) sensu Arnold, 1936 : 443; <$. Misidentification.]
Atopopompilus marshalli Arnold, 1937 : 24. Holotype $, RHODESIA (TM, Pretoria) [examined].

Syn. n.

Atopopompilus parallelus (Haupt) Haupt, 1950 : 58; $.
Atopopompilus coloratus (Haupt) Haupt, 1950 : 58; $.

LECTOTYPE DESIGNATION. Paracyphononyx coloratus Haupt. This species was
described in the couplets of keys to females and males of species of Paracyphononyx.

62 M. C. DAY

A female and a male in the collections of the MNHU, Berlin bear Haupt's deter-
mination labels Taracyphononyx coloratus Haupt det. Haupt 1928'. Neither
bears any form of 'type' label. The female is labelled 'Kamerun' and agrees with
the short description. The male agrees with the short description, and with fig. 41
in the text of Haupt's paper, showing the head in facial view. A putative second
male specimen is indicated by the citation of a range of measurements of body
length (10-11 mm.). Records of MNHU indicate that a 'type' specimen of P.
coloratus should be deposited in the collections, but Dr Konigsmann of that institution
is unable to trace other material. I am satisfied that the female and male I have
seen were before Haupt when he described this species; since a male has been
figured, I have labelled, and here designate as lectotype, the male specimen. Should
a further male specimen of this species come to light, bearing Haupt's determination
label, it should be treated as a paralectotype of P. coloratus.

9- Length 9-17 mm. Black; antennae, head, part or all of pronotum, mesonotum,
scutellum, tibiae and tarsi, sometimes also parts of femora, yellow-brown to red-brown, quite
variable; occasionally (northern populations) face and pronotum substantially yellow. Erect
hairs often quite long, intermediate between A . carinatus and A . kilimandjaroensis. Pubescence
mostly brownish, occasionally silvery on face.

Antennal segments long, at least 4 x as long as wide. Head short, broad, inner orbits
converging above and below (Text-fig. 8). SMC3 narrowed above, normally half as long on
radial vein as is SMC2. Propodaeum declivous but not abruptly so, posterior corners flattened
but not strongly so (Text-fig. 13); short erect hairs of propodaeum not markedly adpressed,
but backwardly directed.

<. Length 8-12 mm. Black; with more or less red-brown coloration on head, pronotum,
mesonotum, scutellum, and coxae; antennae and legs always so. Always with some yellow
coloration behind eyes on temples, posterior pronotal margin, and on dorsal surface of hind
tibia. Yellow may replace red-brown extensively on face and clypeus, pronotum, fore legs
and tarsi. Wings largely hyaline with infuscate margins. Pubescence extensive, silvery, also
much erect silvery hair; propodaeum, save spiracular area, largely so covered. Seventh
tergite white.

Antennae (Text-fig. 28) short, stout, each segment with sharp ventral angle when viewed in
profile. Face with well-marked carina extending onto clypeus. Eyes convergent above and
below, but UID clearly exceeds LID. Genitalia as in Text-fig. 20.

All the new taxa described in Bingham's posthumously published paper (1912)
were designated 'form n.'. A. jacens has thus on occasion been recorded as 'P.
festivus f. 'jacens', since P. festivus Klug was the immediately preceding species
dealt with by Bingham. Arnold erroneously determined A. jacens as P. carinatus,
and recorded P. jacens, P. nasutus and P. personatus as synonyms. The types of
P. jacens and A. marshalli are both part of a series of specimens collected by G. A. K.
Marshall in Salisbury and dispersed to Oxford, Bulawayo and London.

DISTRIBUTION. Widely distributed through parts of the woodland and savanna
regions of Africa; summarized on Map i, p. 60.

VARIATION. Males from southern Africa exhibit least red and yellow coloration
on head and thorax. The extent of areas so coloured increases clinally in a north-
wards direction; thus, males from Cameroun and Nigeria are most extensively
marked with red and yellow. Females from the most northern part of the range
(Sudan, Cameroun, Nigeria) also have a tendency towards replacement of red-brown

REVISION OF A TOPOPOMPILUS 63

by yellowish colour, particularly on the head and pronotum. The type-material of
P. coloratus is of this extreme colour form.

MATERIAL EXAMINED.

Pompilus jacens Bingham, holotype <$, RHODESIA: Salisbury, x.iSgg (G. A. K.
Marshall) (UM, Oxford). Paracyphononyx paralldus Haupt, holotype -, SOUTH
AFRICA: Natal, no further data (MNHU, Berlin). Paracyphononyx coloratus
Haupt, lectotype $, CAMEROON: Bosum, 21-31^.1914 (Tessmanri) (MNHU, Berlin).
Atopopompilus marshalli Arnold, holotype $, RHODESIA: Salisbury, iv. 1900 (G. A. K.
Marshall) (TM, Pretoria).

CAMEROUN: Bosum, i-io.vi.i9i4, i $ (Tessmann); ? Odessi, 5^.1904, i $ (para-
lectotype of P. coloratus) (MNHU). KENYA: Mombasa Island, xi. ign-iii. 1912,
I $ (paratype of A. marshalli); Rabai, viii. 1930, 9 $; viii. 1937, I <$ (van Someren);
Diani Beach, ix. 1951, i <$ (N. L. H. Krauss) ; Tanger province, ix. 1950, i $ (R. C. H.
Sweeney); Ukumbani, Nzoi, i-ii. 1889, i $, i <$ (BMNH). MALAWI: Valley of N.
Rukuru, Karonga district, 2000-4000', i5-i8.vii.i9io, i $ (S. A. Neave) (paratype
of A. marshalli) (NMR). NIGERIA: Yola, i $ (L. N. Lee); Kano, Azare, 4.ix.i925,
i $ (L. Lloyd) (BMNH). RHODESIA: Salisbury, i <j> (G. A. K. Marshall) (paratype
of A. marshalli) (NMR); Salisbury, i-iv. 1900, 2 9, I ^; xi. 1903, i <$; i. 1905, i -
(G. A. K. Marshall); Shangani, de Beer's ranch, v. 1932, i (J. Ogilvie) (BMNH);
S. Rhodesia, 9.xi.i935, i <$ (R. H. R. Stevenson) (NMR); Matopos National Park,
in Malaise trap, 1-2.^.1968, 2 <$ (P. J. Spangler) (USNM). SENEGAL: Dakar, 1907,
i $ (Waterlot) (MNHN). SIERRA LEONE: no further data, 2 $ (Moquerys) (MNHN);
Freetown, Tower Hill, iii. 1908, i $ (A. Pearse) (BMNH). SOUTH AFRICA: Algoa
Bay, 31.111.1892, i , i $ (H. Brauns) ($ paratype of A. marshalli) (TM); Grahams-
town, Strowan, 19.11.1967, i ?; 26.11.1967, i $; 21.1.1968, i $, 10. iii. 1968, i $, i <$;
17.111.1968, i $; 31.111.1968, i <$ (C. Jacot-Guillarmod); Grahamstown, Hilton,
9.^.1967, i ? (C. Jacot-Guillarmod); Grahamstown, 17-25.1.1970, i <$ (J. G. H.
Londt); Grahamstown, Belmont Valley, 20.1.1970, i <; 26.1.1970, i <$ (F. W. Gess);
Kenton-on-Sea, in Malaise trap, iii. 1971, i ? (R. A. Jubb); Howison's Poort, in
Malaise trap, 1-7.11.1972, i $ (F. W. Gess) (AM); Transvaal, 5 miles W. of Warmbad,
24-25.11.1968, i <$ (Krombein & Spangler) (USNM). SUDAN: West Darfur, N. Jebel
Murra, Killing, 29^1.1932, 2 $ (M. Steele) (BMNH). TANZANIA: Stigi, x. 1917, i #
(G. D. H. Carpenter) (BMNH). ZAIRE: Elizabethville, 21^.1923, 2 (M. Bequart)
(MRAC); Kabinda, Lomami, i $ (J. Mutter] (IRSNB); Coquilatville, i $ (NMR).
ZAMBIA: Upper Luangwa River, 27.vii.-i3.viii.i9io, i $ (S. A. Neave} (paratype of
A. marshalli}; i6S., 26E., 25^111.1967, i ? (BMNH).

Atopopompilus nefas (Dalla Torre)
(Text-figs 7, 16, 21, 26, 34)

Salius collaris Saussure, 1891 : 263. Holotype (?) $, MADAGASCAR (originally in MHN,
Geneva, but not located and presumed lost).

64 M. C. DAY

Salius (Schistosalius) collaris Saussure; Saussure, 1892 : 322; $.

Pompilus nefas Dalla Torre, 1897 : 304. Replacement name for Salius collaris Saussure, 1891,

junior secondary homonym in Pompilus of Sphex collaris Fabricius, 1775.
Paracyphononyx nefas (Dalla Torre) Haupt, 1929 : 168; ty, <$.
Psammoderes collaris (Saussure) Banks, 1941 : 353; $.
Atopopompilus nefas (Dalla Torre) Haupt, 1950 : 56; $.

TYPE-MATERIAL. Salius collaris Saussure. The original description of the
female clearly applies to this distinctive species, but Saussure did not indicate
whether he saw more than one specimen. He gives a length of 17 mm. Two very
similar females, both more than 20 mm in length, but otherwise agreeing well with
the description, are in the collections of the MHN, Geneva; there is some ambiguity
in the attached labels of one specimen. Both bear blue printed labels, 'NOSSI BE',
and a second label, printed 'Madagascar' above handwritten 'H. de Saussure'.
One female also bears a label in Saussure's hand, 'Salius collaris $ Ss', and a pink
printed label, 'Madagasc. Antananarivo'. In 1892, Saussure gave a more
detailed description of 5. collaris, and recorded the species from Antananarivo.
Saussure wrote 'Capturee dans la region d' Antananarivo', and I believe his use of
the singular denotes a single specimen rather than the species.

Both females are mounted on thick pins. Saussure's handwritten label also has
pinholes smaller than those made by the thick pin of the specimen currently bearing
this label.

I believe it probable that the holotype was badly damaged and discarded at some
time in the past; as a matter of continuity of interpretation, some person perhaps
attached the labels to a more satisfactory conspecific specimen which, however,
probably postdates description. I have glued the pertinent labels to a separate
card, to preserve them in their present state. I presume the holotype to be
permanently lost. However, Saussure's figure (1892: pi. 23, fig. 13) is quite sufficient
to identify the species satisfactorily.

$. Length 11-22 mm. Black; head except mouthparts, most of pronotum, occasionally
mesonotum and scutellum orange-brown to dark red-brown. Erect hairs of face and pronotum
much as in other species of the genus, but substantially reduced on rest of thorax. Pubescence
dark, sometimes reflecting obscurely violaceous or blue; with a little silvery adpressed
pubescence on propodaeum, frequently lost.

Antennal segments fairly thick, second flagellar segment approximately 4 x as long as thick.
Inner orbits converging above and below, but much less strongly so above than in Ethiopian
species (Text-fig. 7). SMC3 narrowed above, but configuration of SMC2 and SMC3 very
variable. Propodaeum distinctively modified (Text-fig. 16), with a pair of bluntly pointed
postero-lateral prolongations and a shallow median longitudinal impression. Sides of pro-
podaeum almost parallel, very slightly constricted in between spiracle and posterior prolongation.

(. Length 11-15 mm. Black; head (except mouthparts and, occasionally, ocellar region),
frequently pronotum, and occasionally mesonotum and scutellum, light or dark red-brown.
Erect hair and pubescence mostly black, but silvery on lower face and coxae, occasionally
silvery or brownish on propodaeum. Wings fuscous with darker margin.

Antennae roundly crenulate below (Text-fig. 26). Facial carina reaches onto clypeus, but
does not fork (Text-fig. 34). Inner orbits not strongly convergent above, sub-parallel below,
UID clearly exceeds LID. Propodaeum with distinct declivity and lateral prolongations as in
female, but less strongly so. Erect hair of propodaeum quite marked, but much less dense
than on Ethiopian species. Genitalia as in Text-fig. 21.

REVISION OF A TOPOPOMPILUS 65

The name Salius collar is is permanently rejected under Article 59(b)(i) of the
International Code of Zoological Nomenclature, as amended on 30 September, 1972.

Haupt (1929) correctly identified the male of A. nefas. However, Arnold did not
include the Madagascan fauna in his works, and was not familiar with this species.

DISTRIBUTION. Madagascar; summarized on Map i, p. 60.

VARIATION. A. nefas is a variable species, with no strong geographic component
in its variation. Specimens with extensive orange-brown coloration on head
and thorax occur sympatrically with others which have the areas of coloration
substantially reduced and the colour deepened to dark red-brown. In addition,
morphological characters such as the shape of the face and the form of the sub-
marginal cells are noted to vary quite widely. I cannot, however, find any basis
for recognition of undescribed taxa in the material available for study; although
selected individuals may superficially appear quite different, intermediates can
always be found.

MATERIAL EXAMINED.

MADAGASCAR: Betsilei, 1880, 3 $ (D. Cowan) (BMNH); Tananarive, 1914, i J;
1916, i 9; 1919, i ? (Waterlot] (MNHN); Tananarive, i <J (Plason) (MRAC);
Tananarive, i <$ (Sikora) (MNHU); Manjakatampo, 3.1.1958, i $ (F. Reiser);
Antisabe, 15.1.1958, i $ (F. Reiser) (NM); Tzimbazaza, ig.xi.i947, i $ (ISM);
Ambalavao, ix-x. 1938, I $ (C. Lamberton) (USNM); Joffreville, S.v.igsS, i

MAP 2. Distribution of Atopopompilus in the Oriental region.

66 M. C. DAY

(Keiser) (NM) ; Antanambe, i $ (USNM) ; Ambohimasoa, Tsarafidy Forest, 1450 m,
xii. 1959-!. 1960, 1 9 (P. Griveaud} (ISM) ; Ambatolampy, 1931, 2 $ (Lasere) (MNHN) ;
Ambatolampy, 1.1.1958, I $ (Keiser) (NM); Fianarantsoa, Anosimparihy, g.vm.ig^S,
I $ (Keiser) (NM); Ranomafana, 21.1.1958, i $; io-n.ix.i958, 2 <J (NM & coll.
Wahis) ; Rogez, v. 1936, i $; iv. 1937, 3 $ (^4 . Seyrig) (MNHN) ; Rogez, Analandraraka,
vi. 1937, i <$ (A. Seyrig) (coll. Wahis); near Rogez, 900 m, 1946, 2 $ (C. Lamberton)
(USNM & coll. Wahis); Perinet, 500 m, 6-12^.1968, i ? (K. M. Guichard) (BMNH);
Perinet, no further data, i $ (ISM); Perinet, xii. i957-x. 1958, 4 <j>, 3 <$ (Keiser}
(NM); Moramanga, 9.X.I958, 2 $ (Keiser} (NM); Ambohimitombo, 1894, i $ (Forsyth-
Major} (BMNH); Inosy, ii. 1933, i < (4. Seyng) (MNHN); Tulear, Sakaraha,
12.111.1958, i $ (Keiser) (NM); Fort Dauphin, 1913, i $ (Gruvel) (MNHN); Nossi
Be, 2 $ (MHN); no further data, 1919, 2 $ (/. de Gaulle) (MNHN & coll. Wahis).

Atopopompilus daedalus (Bingham) comb. n.
(Text-figs 3, 4, 14, 22, 24, 29, 30)

Pompilus daedalus Bingham, 1896 : 429. LECTOTYPE $, BURMA (BMNH), here designated

[examined].

Anoplius styrus Cameron, 1903 : 327. Holotype , INDIA (BMNH) [examined]. Syn. n.
Ceropales pruinosa Cameron, 1905 : 415. LECTOTYPE (not $, as originally stated), INDIA

(UM, Oxford), here designated [examined]. Syn. n.

[Anoplinellus clotho (Smith) sensu Banks, 1934 : 84; $ Misidentification.]
Anoplinellus undescribed sp. ; Banks, 1934 : 84; $.
Anoplinellus javanus Haupt, 1935 : 318. LECTOTYPE $, JAVA (NM, Basle), here designated

[examined]. Syn. n.

LECTOTYPE DESIGNATIONS, (i) Pompilus daedalus Bingham. The description
cites 'Sikkim; Tenasserim', as localities, and gives a range of measurements. Six
conspecific females in the BMNH collections are from three localities in Tenasserim,
Burma; from Sikkim; and from Darjeeling. All are originally from the Bingham
collection. Two bear labels in Bingham 's handwriting, one 'Pompilus daedalus
Bingham $ Type' and the other 'Pompilus daedalus Bingham $'. The specimen
labelled 'type' has 'the antennae, tibiae and tarsi dull piceous red' and thus agrees
best with the description, since the other females are more nearly black. I have
labelled, and here designate as lectotype, the female which bears Bingham's type
label.

(2) Ceropales pruinosa Cameron. A male bearing Cameron's label, 'Ceropales
pruinosa Cam. type' stands next to a second male in the Rothney collection in
UM, Oxford. Although the description states '$', both males fit the description
well, and it can only apply to this sex. I have labelled, and here designate as
lectotype, the male which bears Cameron's type label.

(3) Anoplinellus javanus Haupt. Haupt lists two females from Gedeh after his
description, and states 'i $ in meiner stammlung [sic\\ This specimen, now in the
ZM, Halle, bears Haupt's 'cotype' label, and is conspecific with the female in NM,
Basle, which bears Haupt's holotype label. Both agree well with the description.
I have labelled, and here designate as lectotype, the Basle specimen.

REVISION OF A TOPOPOMPILUS 67

$. Length 11-20 mm. Black; wings infuscate with darker margin; some populations with
small more or less hyaline areas in forewing, more so in hind wing. With extensive silver or
grey pubescence, particularly silver on face, thorax and coxae, more greyish on anterior of
tergites. Otherwise brown pubescent. Erect hair of head and thorax extensive, very long,
exceeding thickness of scape.

Antennae particularly elongate, thin, second segment of flagellum at least 5-0 X as long as
wide. Front very narrow, MID about 0-5 x TFD in smaller specimens, less in larger specimens.
Vertex in facial view below level of eyes in large specimens. Inner orbits strongly convergent
above and below (Text-fig. 4). Thorax typical of genus; propodaeum similar to that of A.
carinatus, but shorter; declivity more concave (Text-fig. 14), with a central area of aciculation;
lateral angles more or less flattened at 45 degrees or slightly protuberant dorsally. Dorsum
and lateral angles with abundant, tangled brown-silver pubescence in addition to long erect
hairs. SMC3 narrowed above, normally much shorter on radial vein than is SMCa, and
approximately equal to or shorter than SMCa on cubital vein.

(J. Length 8-12 mm. Black; ventral surface of scape, most of clypeus, facial carina, lines
bordering inner orbits and temples, posterior pronotal margin, and spot on dorsal surface of
hind tibia, yellow. Antennae sometimes reddish ventrally; seventh tergite sometimes with
white spot. With much silvery erect hair, especially dense on posterior third of propodaeum;
silvery pubescence widely distributed, merging to greyish on anterior halves of tergites. Wings
fuscohyaline with infuscate margins.

Antennae (Text-fig. 24) short, each segment with sharp ventral angle when viewed in profile.
Facial carina just reaches upper margin of clypeus (Text-figs 29, 30), otherwise very similar to
A. carinatus. Genitalia as in Text-fig. 22.

This distinctive species is sole representative of its genus in the Orient. All
the names in the above synonymy were proposed in works of limited scope which
simply described new taxa.

Surprisingly, specimens in collections have often been misdetermined as various
of the nominal species which are synonyms of Anoplius (Orientanoplius) canifrons
Smith, 1855. It would appear that A. daedalus has often been taken together with
this species of Anoplius, but in fewer numbers; cursory examination of series must
have resulted in failure to segregate these females, which have a similar colour
pattern.

Ceropales (and Ceropalidae) was used by Cameron in 1905 to replace Pompilus
(and Pompilidae) which at that time was thought to be preoccupied in the Octopoda.

DISTRIBUTION. Forest areas of the Oriental and Indonesian regions; summarized
on Map 2, p. 65.

VARIATION. Size variation produces minor allometric differences. For example,
larger females have the face and vertex sunk beneath the contour of the eyes, whilst
smaller females have the face more nearly in the same plane as the surface of the
inner orbits; thus, the eyes of large specimens appear to bulge when compared with
those of smaller females.

The Javan population does not have uniformly infuscate wings in the female.

MATERIAL EXAMINED.

Pompilus daedalus Bingham, lectotype $, BURMA: Tenasserim, Thaungyin Valley,
vi. 1894 (C. T. Bingham) (BMNH). Anoplius styrus Cameron, holotype $, INDIA:
Sikkim? (BMNH). Ceropales pruinosa Cameron, lectotype J, INDIA: Khasia Hills

68 M. C. DAY

(Rothney) (UM, Oxford). A noplinellus javanus Haupt, lectotype 9, JAVA: Tjibodas,
Gedeh, 1400-1600111, viii. 1931 (Handschiri) (NM, Basle).

BORNEO: Sandakan, i 9 (C. F. Baker) (MCZ); Sarawak, Mt Dulit, 4000', moss
forest, i6.x.i932, I 9 (B. M. Hobby & A. W. Moore) (misdet. 'Orientanoplius
iliacus, Haupt, 1937'). (BMNH). BURMA: Tenasserim, Tavoy Valley, iii. 1893, i 9
(paralectotype of P. daedalus); Maulmain, v. 1889, i $ (paralectotype of P. daedalus);
Thaugyin Valley, vi. 1894, i (C. T. Bingham) (BMNH). INDIA: S. India, no
further data, i 9 (C. T. Bingham) ; Sikkim, Runjit Valley, iv. 1894, 2 9 (paralectotypes
of P. daedalus); Darjeeling, 7000', 31.111.1894, i 9 (C. T. Bingham) (paralectotype of
P. daedalus); Khasia Hills, 6 9 (BMNH); Khasia Hills, i 9 (Rothney) (UM). JAVA:
Tjibodas, Gedeh, 1400-1600 m, viii. 1931, i 9 (Handschin) (paralectotype of A.
javanus) (ZM); same data, v. 1935, i 9. i $ (V. d. Vecht) (MCZ); same data, i <j>, i $
(Vecht) (NMR); Mt Djampang, Tjigaeha, i-ii. 1938, 6 9; Salatri, xi. 1937, i <j>;
i. 1938, i 9; Gunung Malang, 3000-4000', x. 1937, 2 $; East Java, Tengger High-
lands, noom, v. 1938, 3 9; Soekaboemi, i. 1938, i 9 (K. M. Walsh] (BMNH).
NEPAL: Gum and Lumsa, 7000', 27. viii. 1952, i <$ (BM Expedn to Nepal) (BMNH).
NORTH VIETNAM: Hoa Binh, 1926, i $ (A. de Cooman) (MNHN) ; Hoa Binh, viii. 1918,
i 9 (R. V. de Salvaza) (BMNH). PHILIPPINE ISLANDS: Mindanao, Iligan, i 9
(C. F. Baker) (USNM). SIAM?: Biserat, i $ (BMNH). SOUTH VIETNAM: Tay
Ninh, 2O.xii.i923, i $; io.xi.i924, i $; 29.xi.i924, i 9 (R. V. de Salvaza) (IRSNB).
SULAWESI: Latimodjong Mts, Bontoe Batoe district, 4000', v. 1931, 2 9 (C. F.
Clagg) ('undescribed species' of Banks, 1934) (MCZ). WEST MALAYSIA: Pahang,
Eraser's Hill, 2O.vii.i936, i 9; Pahang, Cameron Highlands, Ginting Rial, 5000',
i8.vii.i938, i <$; 22^.1939, i 9; 27^.1939, i <j>, i (; 47oo'-5ioo', 12^.1939, i 9;
Selangor, Kuala Sleh, 17^.1936, i 9; Bukit Kutu, 3500', 9.ix.i929, i 9 (ex colln Fed.
Malay States Mus.) (BMNH); Cameron Highlands, Rhododendron Hill, i.x.i935,
i 9; Penang, Ayerltam Reservoir, 8.viii.i968, i 9 (H. T. Pagden) (BMNH).

REFERENCES

ARNOLD, G. 1936. The Psammocharidae of the Ethiopian region. Part VI. Subfamily

Psammocharinae continued. Ann. Transv. Mus. 18 : 415-460.

1937- The Psammocharidae of the Ethiopian region. Part VII. Subfamily Psammoch-
arinae continued. Ann. Transv. Mus. 19 : 1-98.
1951. Sphecidae and Pompilidae (Hymenoptera) collected by Mr K. M. Guichard in

West Africa and Ethiopia, 1941-1948. Bull. Br. Mus. nat. Hist. (Ent.) 2 : 97~ l8 3-
BANKS, N. 1934. The Psammocharidae of the Philippines. Proc. Am. Acad. Arts Sci. 69

(i) : 1-117.
1941. Some Psammocharidae from Madagascar (Hymenoptera). Proc. Acad. nat. Sci.

Philad. 92 (1940) : 335-32.
1947. Studies of South American Psammocharidae. Part 2. Bull. Mus. comp. Zoo/.

Harv. 99 (2) : 371-486.
BINGHAM, C. T. 1896. On some exotic fossorial Hymenoptera in the collection of the British

Museum, with descriptions of new species and of a new genus of Pompilidae. /. Linn. Soc.

25 : 422-445.
1912. South African aculeate Hymenoptera in the Oxford Museum. Trans, ent. Soc.

Lond. 44 (1911) : 528-562.
CAMERON, P. 1903. On some new genera and species of parasitic and fossorial Hymenoptera

from the Khasia Hills, Assam. Ann. Mag. nat. Hist. (7) 11 : 3I3-33 1 -

REVISION OF A TOPOPOMPILUS 69

CAMERON, P. 1905. Descriptions of new Species of Sphegidae and Ceropalidae from the

Khasia Hills, Assam (continued). Ann. Mag. nat. Hist. (7) 15 : 415-424.

1910. Fossores. Wiss. Ergebn. schwed. Exped. Kilimanjaro . . . Meru ... 2 (7) : 197-296.

DALLA TORRE, C. G. DE. 1897. Catalogus hymenopterorum hucusque descriptorum systematicus

et synonymicus. 8. i-viii+ 1-749 pp. Leipzig.
EVANS, - H. E. 1951. A taxonomic study of the Nearctic Spider Wasps belonging to the

tribe Pompilini (Hymenoptera : Pompilidae). Part II: Genus Anoplius Dufour. Trans.

Am. ent. Soc. 76 : 207-361.
1966. A revision of the Mexican and Central American Spider Wasps of the sub-family

Pompilinae (Hymenoptera: Pompilidae). Mem. Am. ent. Soc. 20 : 1-442.
Fox, W. J. 1893. New species of fossorial Hymenoptera. Can. ent. 25 : 113-117.
GRIBODO, G. 1884. Spedizione Italiana nell' Africa Equatoriale; Risultati Zoologici;

Imenotteri. Annali Mus. civ. Stor. nat. Giacomo Doria 21 : 277-325.
HAUPT, H. 1929. Weiterer Ausbau meines systems der Psammocharidae. Mit Beschreibung

neuer Gattungen und Arten. Mitt. zool. Mus. Berl. 15 : 109-197.
1935- In von Schulthess Rechberg, Hymenoptera aus den Sundainseln und

Nordaustralien. Revue suisse Zool. 42 (2) : 293-323.
1950. Pompilidae (Hymenoptera Sphecoidea). Explor. Pare natn. Albert Miss. G. F.

de Witte 69 : 1-63.

HOWARD, L. O. 1901. The Insect Book, xxvii + 429 pp. New York.

KOHL, F. F. 1894. Zur Hymenopterenfauna Afrikas. Annln naturh. Mus. Wien 9 : 279-350.
PATE, V. S. L. 1946. The generic names of the Spider Wasps (Psammocharidae olim

Pompilidae) and their type species. Trans. Am. ent. Soc. 72 : 65-130.
RADOSZKOWSKI, M. O. 1881. Hymenopteres. Jorn. Sci. math. phys. nat. 8 : 197-221.
SAEGER, H. DE. 1945. Contribution a 1'etude des Hymenopteres du Congo Beige: Pompilidae.

Revue Zool. Bot. afr. 39 (1946) : 78-114
SAUSSURE, H. DE. 1891. Hymenopteres nouveaux de Madagascar. Mitt, schweiz. ent. Ges.

8 (7) : 253-269-
1892. Histoire physique, naturelle et politique de Madagascar. XX. Histoire naturelles

des Hymenopteres. 177-590 pp., pis 21-27. Paris.
SMITH, F. 1897. Descriptions of new species of Hymenoptera in the collection of the British

Museum, i-xxi + 1-240 pp. London.

INDEX

Synonyms and invalid names are in italics,
affinis, 55 jacens, 61

Anoplinellus, 49 javanus, 66

Anoplius, 51, 66

v iT kihmandiaroensis, 59

Arachnophroctonus, 51

Atopopompilus, 51 marshalli, 61

Batozonellus, 52 melanicrus, 47

bruneipes, 58 mlanjiensis, 59

canifrons, 67 nasutus, 58

carinatus, 51, 54 nefas, 63

Ceropales, 66

, ..f Paracyphononyx, 47, 52

clotho, 49 ,/f , 3

,, . parallelus, 61

collar is, 63

, personatus, 59

color atus, 61 \ .

pruinosa, 66
crassicorms, 57

daedalus, 66 relativus, 49

Episyron, 52 styrus, 66

festivus, 62 venans, 51, 55

M. C. DAY, B.Sc.

Department of Entomology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON SWy 5BD

ENTOMOLOGY SUPPLEMENTS

3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177:
18 plates, 270 text-figures. August, 1965. 4.20.

4. 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.

6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso-
philidae. Pp. 129: 328 text-figures. May, 1966. 3.

7. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967.

&.I5.

8. 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.

9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species
(Lepidoptera: Rhopalocera). Pp. 509. 8.50. Reprinted 1972.

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho-
palocera). Pp. 332: 348 text-figures. August, 1967. 8.

11. MOUND, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 172:
82 text-figures. May, 1968. 4.

12. 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.

13. 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.

14. CROSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and
its Islands. Pp. 198: I plate, 331 text-figures. July, 1969. 4.75.

15. ELIOT, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera:
Nymphalidae). Pp. 155: 3 plates, 101 text-figures. September, 1969. 4.

16. GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe
(Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969.

19-

17. WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198:

68 plates, 15 text-figures. October, 1971. 12.

18. SANDS, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae).
Pp. 244: 9 plates, 661 text-figures. July, 1972. 9.90.

19. CROSSKEY, R. W. A Re visionary Classification of the Rutiliini (Diptera:
Tachinidae), with keys to the described species. Pp. 167: 109 text-figures.
February, 1973. 6.50

20. VON HAYEK, C. M. F. A Reclassification of the Subfamily Agrypninae
(Coleoptera: Elateridae). Pp. 309: 17 text-figures. October, 1973. 12.30.

21. CROSSKEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia,
including keys to the supraspecific taxa and taxonomic and host catalogues.
Pp. 221: 95 text-figures. December, 1973. 9.55.

PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS OLD WOKING SURREY ENGLAND

19

A GUIDE TO THE GENERA W IBR '

N

AND SPECIES OF PARNASSIINAE
(LEPIDOPTERA : PAPILIONIDAE)

P. R. ACKERY

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 31 No. 4

LONDON i 1975

FRONTISPIECE
Representatives of the seven genera comprising the Parnassiinae (natural size)

FIG. i. Parnassius apollo apollo (Linnaeus) $ (Sweden)

FIG. 2. Sericinus montela montela Fruhstorfer <$ (China : Kiangsi)

FIG. 3. Bhutanitis thaidina thaidina (Blanchard) (China: Szechwan)

FIG. 4. Archon apollinus apollinus (Herbst) < (Turkey)

FIG. 5. Hypermnestra helios helios Nickerl g (U.S.S.R.: Turkmenistan)

FIG. 6. Parnalius polyxena polyxena (Denis & Schiffermiiller) (Hungary)

FIG. 7. Luehdorfia puziloi puziloi (Erschoff) (U.S.S.R.: Russia, Primorye)

19

LPR/PV

A GUIDE TO THE GENERA AND SPECIES ^ c
PARNASSIINAE (LEPIDOPTERA : PAPILIONIDAE)

BY

PHILLIP RONALD ACKERY

-***

Pp. 71-105; 16 Plates, 32 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 31 No. 4.

LONDON : 1975

I \J /

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 Vol. 31 No. 4 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.).

Trustees of the British Museum (Natural History), 1975

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 7 February, 1975 Price 4.80

A GUIDE TO THE GENERA AND SPECIES OF
PARNASSIINAE (LEPIDOPTERA : PAPILIONIDAE)

By P. R. ACKERY

CONTENTS

SYNOPSIS ..........

INTRODUCTION .........

ACKNOWLEDGEMENTS ........

KEY TO THE GENERA OF PARNASSIINAE .....

NOTES ON THE GENERA, WITH KEYS TO THE SPECIES WHERE APPLICABLE

Archon Hiibner

Hypermnestra Menetries

Parnassius Latreille

Sericinus Westwood

Parnalius Rafinesque

Luehdorfia Cruger .

Bhutanitis Atkinson.
REFERENCES .
INDEX

Page

73
73
74
74
75
75
75
76
90
90
92
93
94

SYNOPSIS

Illustrated keys are given to the seven genera and 44 species of Parnassiinae here recognized,
together with brief notes on the distribution and larval food plants of each species. For
polytypic species a list is given of the subspecies that differ most markedly from the typical
form. One generic synonym is newly established.

INTRODUCTION

THE inconsistency of the wing pattern within many species of the Parnassiinae has
inspired the description of numerous subspecies and forms, principally in the works
of Fruhstorfer, Bang-Haas, Bryk and Eisner. Thus, although the literature is
extensive, it is concerned mainly with intra-specific variation rather than the
definition and identification of the recognized species. Bryk (1934; 1935) includes
keys to the Parnassiinae in his extensive monograph; however, the lack of availability
of this work, together with the great emphasis placed on variation within species
throughout the study, renders it unsuitable as a ready means of identification to
species. It is hoped that this present work will to some degree fill this apparent
gap in the literature available to lepidopterists interested in the Parnassiinae.

There has been considerable difference of opinion concerning the higher classifica-
tion of this subfamily, the two tribes here recognized, the Parnassiini and Zerynthiini,
being regarded by Bryk (1934; 1935) and Ford (1944) as subfamilies, while Clench

74 P. R- ACKERY

(1955) and Hemming (1960) have suggested that they be accredited with family
status. I have followed Ehrlich (1958) and Munroe (1960) in treating the two
groups as tribes which together form the subfamily Parnassiinae.

At the generic and specific levels there has been more general agreement, except
over the status of the genus Parnassius and its species (see pp. 76-77). Sericinus,
Archon and Hypermnestra are at present monotypic, Bhutanitis contains four
species and the far-eastern genus Luehdorfia consists of two species. Allancastria,
hitherto a valid monotypic genus, is here considered to be a junior synonym of
Parnalius, which now includes three species. I have adopted Munroe's (1960)
broad concept of Parnassius, which is regarded here as comprising 32 species.
Examples of each genus are illustrated in the Frontispiece, figs 1-7.

To include notes on all the described forms is certainly not the purpose of this
work. I have, however, listed under the relevant species some of those subspecies
that seem to differ most markedly from the more typical forms, together with their
localities and points of difference. No attempt has been made to produce a
comprehensive list of all the subspecies described.

ACKNOWLEDGEMENTS

I wish to thank Mr R. I. Vane- Wright for his help in the preparation of this
work together with Messrs C. F. Huggins, R. L. Smiles and M. A. Kirby whose
helpful suggestions led to many improvements, particularly in the key to Parnassius.
I also extend my thanks to Messrs T. G. Howarth, H. K. Clench, H. J. Epstein,
O. Kudrna, A. Tsvetejev, P. M. Hammond, R. B. Grubh, J. C. Eisner, Suguru
Igarashi and Kazuo Saitoh. To Mr P. York and Mr F. Greenaway I am indebted
for the production of the photographs.

KEY TO THE GENERA OF PARNASSIINAE

1 Outer margin of hindwing rounded, without tails . . . (Parnassiini) 2
Tails of various lengths present on the outer margin of hindwing (Zerynthiini) 4

2 (i) Claspers of male narrow (Text-fig, i) ; forewing with five radial veins (Text-fig. 18)

ARCHON Hiibner (p. 75)
Claspers of male various, never narrow ; forewing with four radial veins . . 3

3 (2) Male tarsal claws equal (Text-fig. 29) . . HYPERMNESTRA Menetries (p. 75)

Male tarsal claws sub-equal (Text-fig. 30) . . PARNASSIUS Latreille (p. 76)

4 (i) Vein M 3 of hindwing produced to a short tapering or rounded tooth (Text-figs

17, 19) PARNALIUS Rafinesque (p. 90)

Vein M 8 of hindwing produced to a distinct tail ...... 5

5 (4) Uncus short, bifid (Text-fig. 16); hindwing tail on vein M 3 longer than the

discal cell of hindwing; vein R 3 of forewing usually arising from discal cell

(Text-fig. 26) SERICINUS Westwood (p. 90)

Uncus elongate, bifid ; hindwing tail on vein M s no longer than the discal cell ;

vein R 3 normally stalked with R t + 6 . . . . 6

6 (5) Precostal cell wide; tails well developed on veins Cw ia and Cu^ of hindwing

(Text-fig. 25); internal ventral surface of claspers smooth (Text-figs 14, 15)

BHUTANITIS Atkinson (p. 93)

GENERA AND SPECIES OF PARNASSIINAE 75

Precostal cell narrow; tails rudimentary on veins Cw ia and Cu^ of hindwing
(Text-fig. 23); internal ventral surface of claspers spinose (Text-figs 12, 13)

LUEHDORFIA Criiger (p. 92)

NOTES ON THE GENERA, WITH KEYS TO THE SPECIES
WHERE APPLICABLE

Tribe PARNASSIINI

Parnassinae Swainson, 1840 : 87, 90. Type-genus: Parnassius Latreille.

ARCHON Hiibner

[Doritis Fabricius sensu Hiibner, [1819] : 86.]

Archon Hiibner, [1822]: 2, 6, 8, 9. Type-species: Papilio thia Hiibner, by subsequent

designation (Scudder, 1875 : 117).

Dorarchon Rothschild, 1918 : 219. Type-species: Papilio apollinus Herbst, by monotypy.
Archon Hiibner; Bryk, 1934 : 19.
Archon Hiibner; Munroe, 1960 : 10.

The name Doritis is still commonly used for this taxon, although Papilio apollo
Linnaeus is clearly the type-species of Doritis, not Papilio apollinus Herbst as
was erroneously believed.

Archon apollinus (Herbst)
(PL i, figs i, 2, Text-figs i, 18)

Papilio apollinus Herbst, 1798 : 156.

Papilio thia Hiibner, [1806] : 60.

Archon apollinus (Herbst); Bryk, 1922 : 224; 1934 : 2 3-

Archon apollinus (Herbst); Eisner, 1966 : 89.

DISTRIBUTION. Rumania. Bulgaria. Turkey. U.S.S.R.: Armenia, Turk-
menistan. Greece. Syria. Iraq. Lebanon. Israel.

LARVAL FOOD PLANTS. Aristolochia hastata (Higgins & Riley, 1970). A. bodame
(Bryk, 1934). A. maurorum (Suguru Igarashi, in litt.).

HYPERMNESTRA Menetries

Ismene Nickerl, 1846 : 207. Type-species: Ismene helios Nickerl, by monotypy. [Junior

homonym of both Ismene Savigny, 1816, and Ismene Swainson, 1820.]
Hypermnestra Menetries, 1848 : pi. 6, fig. i. [Replacement name for Ismene Nickerl.]
Hypermnestra Menetries; Bryk, 1935 : 4-
Hypermnestra Menetries; Munroe, 1960 : 10.

Ehrlich (1958) says that there are reports of Parnassius species in which the
males have nearly symmetrical claws as in Hypermnestra Menetries; however, I
have always found the tarsal claws of male Parnassius to be subequal.

76 P. R. ACKERY

Hypermnestra Helios (Nickerl)
(PI. i, figs 3, 4, Text-figs 2, 20, 29)

Ismene helios Nickerl, 1846 : 208, pi. 3, figs a-g.
Hypermnestra helios (Nickerl); Bryk, 1935 : 7-
Hypermnestra helios (Nickerl) ; Eisner, 1966 : 121

DISTRIBUTION. Iran. Afghanistan. U.S.S.R.: Uzbekistan, Kirghizia.

LARVAL FOOD PLANTS. Zygophyllum (Munroe, 1960). Z. turcomanicum (Verity,
1906). Z. atriplicoides (Bryk, 1935). Z. fabago, Z. portulacoides (Eisner, 1968).
Z. gontsharovii (Stschetkin, 1963).

PARNASSIUS Latreille

Parnassius Latreille, 1804 : 185, 199. Type-species: Papilio apollo Linnaeus, by monotypy.
Doritis Fabricius, 1807 : 283. Type-species : Papilio apollo Linnaeus, by subsequent designation

(Dalman, 1816 : 60).
Tadumia Moore, 1902 : 116. Type-species: Parnassius acco Gray, by original designation.

[Synonymized by Munroe, 1960 : n.]
Kailasius Moore, 1902 : 118. Type-species: Parnassius charltonius Gray, by original

designation. [Synonymized by Munroe, 1960 : n.]
Koramius Moore, 1902 : 120. Type-species: Parnassius delphius Eversmann, by original

designation. [Synonymized by Munroe, 1960 : u.]
Lingamius Bryk, 1935 : 538-540. Type-species: Parnassius hardwickii Gray, by original

designation. [Synonymized by Munroe, 1960 : n.]
Eukoramius Bryk, 1935 : 630, 673-674. Type-species: Parnassius imperator Oberthiir, by

monotypy. [Synonymized by Munroe, 1960 : n.]

Bryk (1935) divides Parnassius into five genera on the basis of differences in the
wing venation and relative lengths of the foretibial epiphysis. As shown by
Munroe (1960) the groups so defined do not agree with those indicated by the male
genitalia. I am adopting Munroe 's broad concept of this genus, but not his suggested
division into two subgenera, Parnassius s. str. and Doritis. The latter name
cannot have any valid use as it is a junior objective synonym of Parnassius.

Elwes (1886) attempted to define the limits of the species, focusing attention on
the wide differences in the shape of the sphragis. The comparative morphology
of the species is discussed by Hering (1932) who recognizes only six distinct species,
this arrangement being followed by Fisher (1950). A revision of the mnemosyne
group is presented by Miiller (1973), partly based on new morphological characters
derived from the comparative structure and arrangement of the wing scales.

Valuable information on this genus is to be found in some of the faunistic studies
on butterflies. Kurentsov (1970) illustrates and discusses the Parnassius of the
eastern U.S.S.R. and the distinguishing characters exhibited by the three western
Palaearctic species are given by Higgins & Riley (1970) . Yokohama & Wakabayashi
(1968) illustrate the representatives in the Japanese fauna, whilst five species are
included by Seok (1939) in his check list of Korean butterflies. The Asian species,
nomion Fischer de Waldheim, is doubtfully listed by Wilson (1961) in his identification
key of North American Papilionidae, but excluded by Dos Passes (1964).

GENERA AND SPECIES OF PARNASSIINAE 77

The studies of the Indian fauna provide useful keys, notably in the works of
Evans (1927) and Talbot (1939). A complete key to the 34 species then recognized
is given by Bryk (1935). Munroe (1960) notes 37 species, five more than Eisner
(1966), but the only species included here are those listed by Eisner in his index
to the Tarnassiana' and Tarnassius Nova' series.

The apparent uniformity of structure within the species-groups, as defined by
Munroe (1960), together with the multiplicity of variations in the wing pattern
within most of the species, presents some difficulty in the preparation of a key to
the species of Parnassius. The wing pattern is used as the principal means of
differentiating between the species since the structural characters derived from the
genitalia, wing venation, sphragis and foretibial epiphysis, are generally applicable
at the species-group level only. As the following key is based on wing pattern at
the species level, it is to be expected that atypical specimens, particularly in the
actius-jacquemontii-phoebus subgroup, will not always run out satisfactorily. It
must therefore be emphasized that the present work should be used only as a guide.

This genus is generally distributed throughout the mountainous and northern
areas of the Palaearctic and western Nearctic regions, the inaccessibility of many
likely localities probably accounting for the rarity of some species.

KEY TO THE SPECIES OF PARNASSIUS LATREILLE

1 Uncus normally plainly visible, spatulate, lying between the two elongate

processes on the tenth tergite (Text-fig. 31); sphragis keeled and with a
simple, single, backward-pointing flange (PL 15, fig. 98) [keel lacking in
epaphus Oberthur]; hindwing upperside lacking submarginal series of blue
spots ............. 2

Uncus sometimes enclosed by eighth abdominal tergite and bifid, never
spatulate (Text-fig. 32) [simple and truncate in simo Gray] ; sphragis various,
never keeled and always lacking a single backward-pointing flange (PI. 15,
figs 99-105); hindwing upperside often with submarginal series of blue
centred spots ........... 16

2 (i) Antennae dark, lacking white scales ........ 3

Antennae with some white scales ........ 5

3 (2) Submarginal band of forewing and hindwing broken up into a distinct series

of internervular black spots (PI. i, figs 5, 6) . apollonius (Eversmann) (p. 80)
Submarginal band of hindwing indistinct or absent ..... 4

4 (3) Pubescence of body mostly pale beneath; wing veins above exceptionally

distinct, with a covering of black scales ; forewing above usually lacking red

markings (PI. 2, figs g, 10) bremeri Bremer (p. 81)

Body clothed with thick black pubescence beneath ; wing veins above without
black scales; forewing above usually with red markings (PI. i, figs 7, 8)

honrathi Staudinger (p. 80)

5 (2) Scaling of antennae, excluding the club, completely white above ... 6

Scaling of antennae, excluding the club, dark and white above . . 7

6 (5) Margins distinctly chequered, being black about the wing veins and white in

the internervular areas; upper surface of hindwing often with a distinct
red basal spot (PI. 3, figs 17, 18) . . nomion Fischer de Waldheim (p. 82)
Margins usually white, sometimes indistinctly chequered; upper surface of
hindwing seldom with a red basal spot (PI. 3, figs 23, 24)

apollo (Linnaeus) (p. 83)

78 P. R. ACKERY

7 (5) Males 8

Females ............. 12

8 (7) Eighth abdominal tergite rounded posterio-laterally ..... 9

Eighth abdominal tergite pointed posterio-laterally . . . . . 10

9 (8) Marginal and submarginal forewing bands poorly developed; ground colour

of upperside creamy white; discal spot in cell Cu^ of forewing upperside
absent or rudimentary, seldom distinctly scaled below and never centred with
red (PI. 2, fig. u) . . . . . . phoebus (Fabricius) (p. 81)

Marginal and submarginal bands of forewing well developed; ground colour
of upperside grey- white, often distinctly dusted with black; discal spot in
cell CMjb of forewing upperside usually present, distinctly scaled below and
often centred with red (PL 3, fig. 21) . tianschanicus Oberthiir (p. 82)

10 (9) Basal red spot usually present in hindwing upperside; discal spot in cell Cu^

of forewing underside often heavily scaled, sometimes centred with red

(PL 2, fig. 15) ...... jacquemontii Boisduval (p. 81)

Basal red spot normally absent from the hindwing upperside, if present the
discal spot in cell Cu^, of forewing underside is lightly scaled, not centred
with red ............ n

11 (10) Black basal scaling normally quite extensive; margins of forewings usually

distinctly chequered, being black about the veins and white in the inter-
nervular areas (PL 3, fig. 19) . . . . . epaphus Oberthiir (p. 82)
Black basal scaling usually less extensive; margins of forewing seldom distinctly

chequered (PL 2, fig. 13) . . . . actius (Eversmann) (p. 81)

12 (7) Sphragis lacking a keel (PL 3, fig. 20) ... epaphus Oberthur (p. 82)

Sphragis strongly keeled . . . . . . . . . .13

13 (12) Discal spot in cell Cu^ of forewing underside distinctly scaled; red basal spot

never present in hindwing upperside ; upperside distinctly dusted with black
scales (PL 3, fig. 22) ..... tianschanicus Oberthur (p. 82)
Discal spot in cell Cw x b of forewing underside normally only lightly scaled,
if distinctly scaled then red basal spot normally present in hindwing
upperside; upperside not distinctly dusted with black .... 14

14 (13) Abdomen usually hairy, almost as much so as the male [Asian species only]

(PL 2, fig. 14) ....... actius (Eversmann) (p. 81)

Abdomen not exceptionally hairy [except European phoebus] . . . 15

15 (14) Hindwing upperside normally with a distinct red basal spot [Asian species

only] (PL 2, fig. 16) ..... jacquemontii Boisduval (p. 81)
Hindwing upperside seldom with a distinct red basal spot [N. American,

European & Asian species] (PL 2, fig. 12) . phoebus (Fabricius) (p. 81)

16 (i) Foretibial epiphysis short, not reaching the end of the tibia (Text-fig. 27);

hindwing upperside without a submarginal series of blue spots (except
Orleans Oberthur) .......... 17

Foretibial epiphysis longer, often reaching the end of the tibia (Text-fig. 28) ;

hindwing upperside usually with a submarginal series of blue spots . . 24

17 (16) Hindwing underside lacking red ocelli . . . . . . . 18

Hindwing underside usually with red ocelli ...... 20

18 (17) Outer surface of palpi dark, occasionally with a few light yellow scales; face

lacking white hairs; bifid uncus with paired ventral processes (Text-fig. 6,

PL 4, figs 29, 30) ...... glacialis Butler (p. 84)

Outer surface of palpi with white scales; face with some white hairs; bifid

uncus without ventral processes . . . . . . . . 19

19 (18) Forewing upperside usually with two black spots in the discal cell; clasper

of male weakly produced posteriorly, never ending in a long thin point

(PL 4, figs 25, 26) ..... mnemosyne (Linnaeus) (p. 83)

GENERA AND SPECIES OF PARNASSIINAE 79

Forewing upperside without two black spots in the discal cell ; clasper of male
strongly produced posteriorly, ending in a long thin point (PI. 4, figs 27, 28)

stubbendorfli Menetries (p. 84)

20 (16) Face usually with golden yellow pubescence . . . . . . . 21

- Pubescence of face black, pale yellow or white . . . . . . 22

21 (20) Basal black scaling of hindwing upperside sharply differentiated; margins

of hindwing underside black; wings of male often yellow (PI. 4, figs 31, 32,
PI. 5, figs 33, 34) ..... eversmanni Menetries (p. 84)

Basal dark scaling of hindwing upperside not sharply divided from the white
ground colour; margins of hindwing underside partly white; wings of male
never yellow (PL 5, figs 39, 40) .... clodius Menetries (p. 85)

22 (20) Hindwing upperside and underside with a submarginal series of internervular

blue-centred spots; margins of forewing upperside usually distinctly
chequered, black about the veins and white in the internervular areas (PI. 6,
figs 41, 42) ........ Orleans Oberthiir (p. 85)

Hindwing upperside lacking blue-centred internervular spots ; forewing margins

never divided into distinct black and white areas ..... 23

23 (22) Forewing apex largely hyaline, the submarginal internervular markings

usually absent, if present pale and indistinct; veins of hindwing underside
pale (PI. 5, figs 35, 36) .... nordmanni (Nordmann) (p. 85)

Forewing above with submarginal internervular markings forming a distinct
band in the wing apex; veins of hindwing underside dark (PI. 5, figs 37, 38)

ariadne Lederer (p. 85)

24 (16) Vein R z stalked with vein Ry_ 5 , not arising from discal cell (Text-fig 24) . 25

Vein R z arising from discal cell (Text-figs 21, 22) ..... 28

25 (24) Hindwing upperside usually with large red postdiscal spots .... 26

Red postdiscal spots of hindwing upperside small or absent . . . . 27

26 (25) Hindwing blue-black internervular submarginal spots absent from cell R & ;

sphragis straight (PL 9, figs 67, 68, PL 15, fig. 105) inopinatus Kotzsch (p. 89)
Hindwing blue-black internervular submarginal spot present in cell R 5 ;

sphragis coiled (PL 9, figs 65, 66, PL 15, fig. 104) . charltonius Gray (p. 88)

27 (25) Hindwing upperside without red markings but with an orange postdiscal band,

wide in female, narrow in male (PL 9, figs 71, 72) . autocrator Avinoff (p. 89)
Hindwing upperside with red markings and lacking an orange postdiscal band

(PL 9, figs 69, 70) . . . . . . loxias Piingeler (p. 89)

28 (24) Vein R 2 anastomosing with vein R^ (Text-fig. 2 1) ..... 29

Vein R 2 not anastomosing with vein R t (Text-fig. 22) .... 32

29 (28) Hindwing upperside usually with large blue-centred submarginal spots;

sphragis large, bilobate (PL 8, figs 63, 64, PL 15, fig. 103)

imperator Oberthiir (p. 88)
Hindwing upperside seldom with large blue-centred submarginal spots ; sphragis

never bilobate ........... 30

30 (29) Uncus short, strongly bifid; sphragis normally laterally flattened, forming a

complete ring about the abdomen (PL 7, figs 51, 52, 53, 54) acco Gray (p. 86)

Uncus simple or weakly bifid; sphragis never laterally flattened or forming

a complete ring about the abdomen ...... 31

31 (30) Uncus short, simple (Text-fig. 8); sphragis rudimentary; hindwing underside

with few postdiscal spots (PL 10, figs 73, 74) . . simo Gray (p. 89)

Uncus elongate, weakly bifid (Text-fig. 3); sphragis well formed, distinct;
hindwing underside with a complete row of postdiscal spots (PL 10, figs 75,
76) ........ tenedius Eversmann (p. 89)

32 (28) Clasper broad; internal process half to two-thirds the length of the clasper

(Text-figs 4, 5) ; sphragis simple, usually laterally flattened (PL 15, fig. 100) . 33

So P. R. ACKERY

Clasper narrow, distally pointed; internal process one-third the length of the

clasper (Text-fig. 7); sphragis bilobate (PI. 15, figs 101, 102). 35

33 (3 2 ) Veins of hindwing underside bordered with white scales which cover the veins

(PI. 6, figs 45, 46) ..... szechenyii Frivaldsky (p. 86)

Wing veins clearly visible, not covered by white scales .... 34

34 (33) Male with a distinct row of bristles anterior to the uncus; clasper produced

dorsally to a distinct angle (Text-fig. 4) ; submarginal blue spots of hindwing
upperside usually centred with white (PI. 6, figs 43, 44) hardwickii Gray (p. 86)
Male without a distinct row of bristles anterior to the uncus; clasper large,
rounded; submarginal spots of hindwing upperside, when present, seldom
centred with white (PL 6, figs 47, 48) . . cephalus Grum-Grshimailo (p. 86)

35 (32) Sclerotized area of male eighth abdominal tergite produced posteriorly to

give two finger-like projections; sphragis produced backwards, forming two
lateral points (PI. 15, fig. 101); basal black scaling of hindwing upperside
usually very extensive, often surrounding the red ocelli of hindwing above
(PI. 8, figs 61, 62) .... acdestis Grum-Grshimailo (p. 87)

- Sclerotized area of male eighth abdominal tergite not strongly produced, never

with finger-like projections ; sphragis produced backwards to form two ventral
lobes (PI. 15, fig. 102); basal black scaling of hindwing upperside usually
rather less extensive . . . . . . . . . .36

36 (35) Submarginal spots of hindwing upperside usually distinct and centred with

blue (PI. 8, figs 57, 58, 59, 60) ... delphius (Eversmann) (p. 87)

Submarginal spots of hindwing upperside, when present, paler and not centred

with blue (PI. 7, figs 55, 56) patricius Niepelt (p. 87)

THE APOLLO-GROVP

Parnassius apollonius (Eversmann)

(PI. i, figs 5, 6)

Doritis apollonius Eversmann, 1847 : 71, pi. 3, figs 1,2.
Parnassius apollonius (Eversmann); Bryk, 1935 : 176.
Parnassius apollonius (Eversmann) ; Eisner, 1966 : 89.

DISTRIBUTION. U.S.S.R.: Uzbekistan, Tadzhikistan, Kirghizia. China: western
Sinkiang.

LARVAL FOOD PLANTS. Salsola (Elwes, 1886). Scabiosa (Stichel, 19076). Radiola
semenovi (A. Tsvetajev, pers. com.)

Parnassius honrathi Staudinger
(PI. i, figs 7, 8)

Parnassius honrathi Staudinger, 1882 : 161, pi. i, figs 4, 5a, pi. 2, fig. 5.
Parnassius honrathi Staudinger & Bang-Haas; Bryk, 1935 : 185.
Parnassius honrathi Staudinger & Bang-Haas; Eisner, 1966 : 122.

DISTRIBUTION. Afghanistan. U.S.S.R.: Tadzhikistan.

GENERA AND SPECIES OF PARNASSIINAE 8l

Parnassius bremeri Bremer
(PI. 2, figs 9, 10)

Parnassius bremeri Bremer, 1864 : 6 (Felder in litt.).
Parnassius bremeri Bremer; Felder & Felder, 1865 : 133.
Parnassius bremeri Bremer; Bryk, 1935 : 190.
Parnassius bremeri Bremer; Eisner, 1966 : 94.

DISTRIBUTION. China: Heilunkiang, Shansi, Hopei. U.S.S.R.: Russia (Chita,
Khabarovsk, Sakhalin, Kamchatka). Korea.

LARVAL FOOD PLANTS. Various species of Sedum (Stichel, 19076). 5. aizon,
S. ischida, S. ussuriensis, S. quadriflorum (Kurentsov, 1970).

Parnassius phoebus (Fabricius)
(PI. 2, figs ii, 12, Text-fig. 31)

Papilio phoebus Fabricius, 1793 : 181.
Parnassius phoebus (Fabricius) ; Bryk, 1935 : 206.
Parnassius phoebus (Fabricius); Eisner, 1966 : 157.

DISTRIBUTION. Europe: Italy, Austria and Switzerland. [Maritime Alps and
eastward to Styria and Grossglockner, rare in north, occasional in Allgauer Alps
(Higgins & Riley, 1970).] U.S.A.: Alaska, Washington, Idaho, Montana, Dakota,
Wyoming, California, Nevada, Utah, Colorado, New Mexico. Canada: British
Columbia, Alberta. China: Sinkiang. U.S.S.R.: Russia (Irkutsk, Amur),
Kazakhstan. Mongolia.

LARVAL FOOD PLANTS. Saxifraga aizioides, Sempervivum montanum (Higgins &
Riley, 1970). Sedum stenopetalum, Sempervivum and Saxifraga (Wilson, 1961).
Sedum telephium, S. fabria, S. album, S. roseum and Sempervivum tectorum (Bryk,
I 935)- Saxifraga calycina, S. nivalis (Kurentsov, 1970). Carexfilifolia, Gayophytum
diffusum, Phlox douglasii, Sedum debile, S. obtusatum, S. wrightii (Tietz, 1972).
Sedum lanceolatum (=stenopetalum] (Scott, 1973).

Parnassius actius (Eversmann)

(PI. 2, figS 13, 14)

Doritis actius Eversmann, 1843 : 540, pi. 9, figs 2a, b.
Parnassius actius (Eversmann) ; Bryk, 1935 : 2 49-
Parnassius actius (Eversmann); Eisner, 1966 : 82.

DISTRIBUTION. Afghanistan. U.S.S.R.: Tadzhikistan, Kirghizia, Kazakhstan.
China: Sinkiang, Kansu. Pakistan. Kashmir.

Parnassius jacquemontii Boisduval
(PI. 2, figs 15, 16)

Parnassius jacquemontii Boisduval, 1836 : 400.
Parnassius jacquemontii Boisduval; Bryk, 1935 : 257.

82 P. R. ACKERY

Parnassius jacquemontii Boisduvai; Eisner, 1966 : 128.
Parnassius jacquemontii Boisduvai; Ackery, 1973 : 6.

DISTRIBUTION. Afghanistan. U.S.S.R.: Tadzhikistan, Uzbekistan. China:
Sinkiang, Kansu, Szechwan. N. India. Pakistan. Tibet.

Variable species. Red basal spot sometimes absent. Superficially many forms
resemble epaphus Oberthur, but the sphragis of the female always bears a keel.

Subsp. mercurius Grum-Grshimailo [Tibet, Amdo], subsp. jupiterius Bang-Haas
[Kansu, Pullow mont, Minschan], subsp. tatungi Bryk & Eisner [Kansu, Richthofen
Mts, Nanschan Mts]. Wing margins distinctly chequered.

Subsp. thibetanus Leech [Szechwan, How-Kow, Ta-tsien-lou]. Dusted with
black scales.

Parnassius epaphus Oberthiir
(PI. 3, figs 19, 20)

Parnassius epaphus Oberthur, 1879 : 23.
Parnassius epaphus Oberthur; Bryk, 1935 : 270.
Parnassius epaphus Oberthur; Eisner, 1966 : 106.
Parnassius epaphus Oberthur; Ackery, 1973 : 5.

DISTRIBUTION. Afghanistan. Pakistan. Kashmir. N. India. Nepal. Sikkim.
Tibet. China: Sinkiang, Szechwan, Kansu, Tsinghai.

Variable species. Red basal spot sometimes present causing many specimens
to resemble jacquemontii Boisduvai, but sphragis always without a keel.

Parnassius tianschanicus Oberthur

(PI. 3, figS 21, 22)

Parnassius corybas var. tianschanicus Oberthur, 1879 : 108.
Parnassius tianschanicus Oberthur; Bryk, 1935 : 288.
Parnassius tianschanicus Oberthur; Eisner, 1966 : 184.

DISTRIBUTION. U.S.S.R.: Uzbekistan, Tadzhikistan, Kirghizia. Afghanistan.
Pakistan. Kashmir. China: Sinkiang.

Parnassius nomion Fischer de Waldheim
(PI. 3, figs 17, 18)

Parnassius nomion Fischer de Waldheim, 1823 : 242, pi. 6, figs 3, 4.
Papilio apollo var. nomion; Geyer, [1838] : pi. 207, fig. 1029.
Parnassius nomion Hiibner; Bryk, 1935 : 300.
Parnassius nomion Hiibner; Eisner, 1966 : 149.

DISTRIBUTION. Mongolia. U.S.S.R.: Russia (Irkutsk, Buryat, Amur,
Khabarovsk, Altay). China: Kansu, Tsinghai, Shensi, Heilungkiang, Liaoning.
Korea. U.S.A.: Alaska?

GENERA AND SPECIES OF PARNASSIINAE 83

LARVAL FOOD PLANTS: Yellow-flowered Sedum (Elwes, 1886). S. album (Bryk,

1935).

Red basal spot sometimes absent causing specimens to resemble apollo Linnaeus.

Subsp. epaphoides Bryk & Eisner [Kansu, Richthofen Mts.]. Much smaller
than the typical form.

Parnassius apollo (Linnaeus)
(PI. 3, figs 23, 24, PI. 15, fig. 98)

Papilio apollo Linnaeus, 1758 : 465.
Parnassius apollo (Linnaeus); Latreille, 1804 : 199.
Parnassius apollo (Linnaeus); Bryk, 1935 : 325.
Parnassius apollo (Linnaeus); Eisner, 1966 : 89.

DISTRIBUTION. Sweden. Finland. Poland. Germany. France. Spain.
Switzerland. Austria. Czechoslovakia. Hungary. Rumania. Italy. Bulgaria.
Greece. Albania. Turkey. Syria. U.S.S.R.: Latvia, Lithuania, Ukraine, Armenia,
Caucasus, Russia (Orel, Ural Mountains, Omsk, Altay, Tuva). China: Sinkiang.
Mongolia.

LARVAL FOOD PLANTS: Stonecrop; Sedum, especially S. album, S. telephium, S.
purpurascens, and Sempervivum (Higgins & Riley, 1970). Sedum acre (Rebel &
Rogenhofer, 1893). 5. annuum, S. villosum and S. roseum (Eisner, 1958). S.
maximum (Holik, 1937).

Subsp. geminus Stichel (Switzerland: Grindelwald) . Faintly yellowish, red
spots smaller.

Subsp. bartholomaeus Stichel [Germany: Konigseej. Small, male heavily
marked above.

Subsp. rhodopensis Markovic [Bulgaria: Rila Planina, Rhodopi Planinaj.
Very large.

Subsp. nevadensis Oberthiir [Spain: Sierra Nevada]. Male above with yellow
ocelli.

Subsp. sicilae Oberthiir [Sicily], Very small (Higgins & Riley, 1970).

THE MNEMOSYNE-GROVP

Parnassius mnemosyne (Linnaeus)
(PI. 4, figs 25, 26, PI. 15, fig. 99, Text-fig. 32)

Papilio mnemosyne Linnaeus, 1758 : 465.
Parnassius mnemosyne (Linnaeus) ; Bryk, 1935 : 19.
Parnassius mnemosyne (Linnaeus); Eisner, 1966 : 142.

DISTRIBUTION. Finland. Sweden. Denmark (Baltic Islands). Germany.
France. Spain. Switzerland. Italy. Austria. Poland. Czechoslovakia. Hungary.
Yugoslavia. Rumania. Bulgaria. Albania. Greece. Turkey. Syria. Lebanon.
Iraq. Iran. Afghanistan. U.S.S.R.: Estonia, Latvia, Lithuania, Ukraine,
Uzbekistan, Tadzhikistan, Kirghizia, Russia (Mordov, Ural Mountains).

84 P. R- ACKERY

LARVAL FOOD PLANTS. Corydalis (Higgins & Riley, 1970). C. cava and C.
helleri (Bryk, 1935). C. solida (Rebel & Rogenhofer, 1893). C. parnanica (Kolar,
1937). C. intermedia (Eisner, 1958).

Parnassius stubbendorfii Menetries
(PL 4, figs 27, 28)

Parnassius stubbendorfii M6netries, 1849 : 273, pi. 6, fig. 2.
Parnassius stubbendorfi Menetries; Bryk, 1935 : 107.
Parnassius stubbendorfi Menetries; Eisner, 1966 : 179.

DISTRIBUTION. U.S.S.R.: Russia (Altay, Tuva, Buryat, Chita, Amur,
Khabarovsk, Primorye, Sakhalin, Kurile Islands). Mongolia. Tibet. Korea.
China: Heilungkiang, Kansu, Szechwan, Tsinghai. Japan: Hokkaido.

LARVAL FOOD PLANTS. Corydalis ambigua, C. gigantaea (Kurentsov, 1970).
Aristolochia debilis (Lee, 1958).

Parnassius glacialis Butler
(PL 4, figs 29, 30, Text-fig. 6)

Parnassius glacialis Butler, 1866 : 50.
Parnassius stubbendorfi glacialis Butler; Bryk, 1935 : 128.
Parnassius glacialis Butler; Eisner, 1966 : 117.
Parnassius glacialis Butler; Ackery, 1973 : 7.

DISTRIBUTION. Japan: Hokkaido, Honshu, Shikoku. Korea. China: Hupeh,
Shangtung, Kiangsu, Anhwei, Chekiang.

LARVAL FOOD PLANTS. Corydalis incisa, C. ambigua, C. decumbens and C. remota
(Suguru Igarashi, in litt). Aristolochia debilis (Lee, 1958).

Parnassius eversmanni Me'netries
(PL 4, figs 31, 32, PL 5, figs 33, 34)

Parnassius eversmanni Menetries, 1855 : 73, pi. i, fig. 2.
Parnassius eversmanni Menetries; Hemming, 1934 : 199.
Parnassius eversmanni Menetries; Bryk, 1935 : I 33-
Parnassius eversmanni Menetries; Eisner, 1966 : 108.

DISTRIBUTION. U.S.S.R.: Russia (Irkutsk, Buryat, Primorye, Khabarovsk,
Tuva, Chita, Yakut, Magadan, Kamchatka, Yevrey). Mongolia. Japan: Hokkaido.
U.S.A.: Alaska.

LARVAL FOOD PLANTS. Corydalis gigantea (Fumariaceae) (Wilson, 1961).
Dicentra peregrina (Suguru Igarashi, in litt.).
Subsp. felderi Bremer [U.S.S.R.: Russia (Amur, Khabarovsk, Yevrey)].

GENERA AND SPECIES OF PARNASSIINAE 85

Yellow pigment absent in male. Postdiscal and costal spots often black.
Superficially similar to glacialis Butler.

Parnassius nordmanni (Nordmann)
(PI. 5, figs 35, 36)

Doritis nordmanni Nordmann, 1851 : 423, pi. 13, figs 1-3.
Parnassius nordmanni (Nordmann) ; Hemming, 1934 : J 98.
Parnassius nordmanni Menetries; Bryk, 1935 : I 4^-
Parnassius nordmanni Menetries; Eisner, 1966 : 149.

DISTRIBUTION. U.S.S.R.: Caucasus.

Parnassius ariadne Lederer
(PI. 5, figs 37, 38)

Doritis clarius Eversmann, 1843 : 539, pi. 9, figs lA, B, C (nee clarius Hiibner, [1806] : 61,

nota 6).

Parnassius ariadne Lederer, 1853 : 354.
Parnassius ariadne Lederer; Hemming, 1934 : J 9^'
Parnassius clarius (Eversmann); Bryk, 1935 : 151.
Parnassius clarius (Eversmann) ; Eisner, 1966 : 98.

DISTRIBUTION. U.S.S.R.: Russia (Altay), Tadzhikistan. Western Mongolia.

Parnassius clodius Menetries
(PI. 5, figs 39, 40)

Parnassius clodius Menetri6s, 1855 : 7.
Parnassius clodius Menetries ; Bryk, 1935 : J 56.
Parnassius clodius Menetries; Eisner, 1966 : 98.

DISTRIBUTION. U.S.A.: Alaska, Washington, Idaho, Montana, Wyoming,
Oregon, Nevada. Canada: British Columbia.

LARVAL FOOD PLANTS. Viola, Sedum (Stonecrop), Vaccinium?, Rubus? (Wilson,
1961). Saxifraga sp., Vitis calif ornica (Tietz, 1972).

Parnassius Orleans Oberthiir
(PI. 6, figs 41, 42, Text-fig. 27)

Parnassius Orleans Oberthiir, 1890 : i.
Parnassius Orleans Oberthiir, 1891 : 8, 18, pi. i, fig. 2.
Parnassius Orleans Oberthiir; Bryk, 1935 : 163.
Parnassius Orleans Oberthiir; Eisner, 1966 : 154.

DISTRIBUTION. Tibet. Mongolia. China: Sinkiang, Tsinghai, Kansu, Shensi,
Szechwan, Yunnan.

86 P. R. ACKERY

THE HARDWICKII-GROVP

Parnassius hardwickii Gray
(PI. 6, figs 43, 44, Text-fig. 4)

Parnassius hardwickii Gray, 1831 : 32.
Lingamius hardwickei (Gray); Bryk, 1935 : 54 1 -
Lingamius hardwickei (Gray); Eisner, 1966 : 120.

DISTRIBUTION. Kashmir. N. India. Nepal. Sikkim. Bhutan.

LARVAL FOOD PLANTS. Various species of Saxifrage (Moore, 1902).
Superficially similar to Orleans Oberthiir but normally distinguishable by the
white-centred, blue, submarginal series of spots on the hindwing upperside.

THE SZECHENYII-GROUP

Parnassius szechenyii Frivaldszky
(PI. 6, figs 45, 46, PI. 15, fig. 100, Text-fig. 5)

Parnassius szechenyii Frivaldszky, 1886 : 39, pi. 4, figs i, la.
Koramius szechenyii (Frivaldszky); Bryk, 1935 : 550.
Koramius szechenyi (Frivaldszky); Eisner, 1966 : 181.

DISTRIBUTION. Tibet. China: Tsinghai, Kansu, Szechwan, Yunnan.

Parnassius cephalus Grum-Grshimailo
(PI. 6, figs 47, 48, PI. 7, figs 49, 50, Text-fig. 22, 30)

Parnassius cephalus Grum-Grshimailo, 1891 : 446.
Koramius cephalus (Grum-Grshimailo) ; Bryk, 1935 : 558.
Koramius cephalus (Grum-Grshimailo); Eisner, 1966 : 97.

DISTRIBUTION. Tibet. China: Kansu, Szechwan, Tsinghai. Kashmir.

Subsp. maharaja Avinoff [Kashmir: Ladakh Range]. Markings generally
reduced. Submarginal spots above faint, not centred with blue. Postdiscal and
costal spots absent from hindwing upperside.

This subspecies (PI. 7, figs 49, 50), treated by both Bryk (1935) and Munroe (1960)
as a species, differs from typical cephalus in having veins R^ and R z consistently
anastomosing, causing it to key out as P. acco Gray.

THE ACCO-GROUP
Parnassius acco Gray

(PL 7, figs 51, 52, 53, 54)

Parnassius acco Gray, 1853 : 76, pi. 12, figs 5, 6.
Tadumia acco (Gray); Bryk, 1935 : 631.
Tadumia acco (Gray); Eisner, 1966 : 82.

GENERA AND SPECIES OF PARNASSIINAE 87

DISTRIBUTION. Kashmir. Tibet. Sikkim.

Subsp. liliput Bryk [Tibet: Everest District], subsp. hunningtoni Avinoff
[Tibet: Dochar, Tuna, Tsangpo Valley, Dzara, Kyetrak, Chumbi Valley. Sikkim:
Gangtok], Smaller than the typical form, red markings absent.

THE DELPHIUS-GROUP

Parnassius patricius Niepelt

(PI- 7, figs 55, 56)

Parnassius patricius Niepelt, 1911 : 274.
Koramius patricius (Niepelt); Bryk, 1935 : 568.
Koramius patricius (Niepelt); Eisner, 1966 : 155.

DISTRIBUTION. U.S.S.R.: Kirghizia.

Parnassius acdestis Grum-Grshimailo
(PI. 8, figs 61, 62, PI. 15, fig. 101, Text-fig. 7)

Parnassius delphius var. acdestis Grum-Grshimailo, 1891 : 446.
Koramius acdestis (Grum-Grshimailo) ; Bryk, 1935 : 572.
Koramius acdestis (Grum-Grshimailo) ; Eisner, 1966 : 82.

DISTRIBUTION. U.S.S.R. : Kirghizia. Kashmir. Tibet. Sikkim. Bhutan.
China: Sinkiang, Szechwan.

The arrangement of the forewing radial veins is more variable in this species
than in any other. Although veins R and R z are usually separate, in many
specimens they do appear to touch and in some cases quite definitely anastomose.

Subsp. lucifer Bryk [Sikkim: Gyamtshona]. Postdiscal and costal spots black
in hindwing above.

Subsp. lux Eisner [Tibet: Jung-jung Khola]. Basal black scaling in hindwing
above far less extensive than in the typical form. Postdiscal and costal spots
large.

Parnassius delphius (Eversmann)
(PI. 8, figs 57, 58, 59, 60, PI. 15, fig. 102)

Doritis delphius Eversmann, 1843 : 541, pi. 7, figs ra, b.
Parnassius delphius (Eversmann); Ehves, 1886 : 39.
Koramius delphius (Eversmann); Bryk, 1935 : 583.
Koramius delphius (Eversmann) ; Eisner, 1966 : 102.

DISTRIBUTION. Afghanistan. U.S.S.R.: Tadzhikistan, Kirghizia, Uzbekistan.
Pakistan. Kashmir. N. India. Tibet. China: Sinkiang, Tsinghai.

Highly variable species. Hindwing discal spots often without red scales.

88 P. R. ACKERY

Subsp. pulchra Eisner [Kirghizia: Kungey Alatau Mountains]. Wings exception-
ally dark, semi-transparent.

Although treated by Bryk (1935) and Munroe (1960) as a distinct species,
Parnassius stoliczkanus is here regarded, in accordance with Eisner (1966), as a
subspecies of Parnassius delphius (Eversmann). The following subspecies are
those that Bryk and Munroe would have included in Parnassius stoliczkanus
Felder & Felder.

Subsp. atkinsoni Moore [Kashmir: Pir Pinjal, Sind Valley, Burzil Pass. India:
Himachal Pradesh, Kulu], subsp. beate Eisner [Kashmir: Karakoram, Potu-la
Pass, Chalsi, Leh], subsp. chitralica Verity [Pakistan: Chitral], subsp. florenciae
Tytler [Tibet: Phupes Hundes, Tibu, Churmurti], subsp. gracilis Bryk & Eisner
[India: Himachal Pradesh, Kangra, Rohtang Pass], subsp. imitator Bryk & Eisner
[U.S.S.R.: Tadzhikistan, Pamirs, Beik Pass], subsp. kumaonensis Riley [India.
Uttar Pradesh, Kumaon, Shillung], subsp. nicevillei Avinoff [Kashmir: Pir Pinjal,
Burzil Pass, Sari Sungur Pass, Sapta La], subsp. parangensis Eisner [India: Himachal
Pradesh, Parang Pass, Bara Lacha Pass. Kashmir: Tagalang Pass, Lingti, Ladahk],
subsp. rileyi Tytler [Kashmir: Rupal Valley, Astor], subsp. spitiensis Bang-Haas
[Tibet: Spiti, Tum-Tum-Thang, Churmurti], subsp. stoliczkanus Felder & Felder
[Kashmir: Ladak, Rupshu, Sapta La], subsp. tennis Bryk & Eisner [Kashmir:
Gya-Ladahk, Tagalang Pass], subsp. tytlerianus Bryk & Eisner [Kashmir: Chitral,
Bangol Pass], subsp. zanskarica Bang-Haas [Kashmir: Nira, Zanskar Mts.], subsp.
zogilaica Tytler [Kashmir: Zogila]. Generally smaller than the typical delphius.
Red costal spot of hindwing upperside usually absent.

THE IMPERA TOR-GRoup

Parnassius imperator Oberthiir

(PI. 8, figs 63, 64, PL 15, fig. 103, Text-fig. 21)

Parnassius imperator Oberthiir, 1883 : 77.
Tadumia imperator (Oberthiir); Bryk, 1935 : 675.
Eukoramius imperator (Oberthiir); Eisner, 1966 : 123.

DISTRIBUTION. Tibet. China: Tsinghai, Kansu, Szechwan, Yunnan.
LARVAL FOOD PLANT. Corydalis (Verity, 1907).

THE CHARLTONIUS-GROVP

Parnassius charltonius Gray
(PI. 9, figs 65, 66, PI. 15, fig. 104, Text-fig. 24)

Parnassius charltonius Gray, 1853 : 77, pi. 12, fig. 7.
Koramius charltonius (Gray) ; Bryk, 1935 : 694.
Koramius charltonius (Gray); Eisner, 1966 : 97.

DISTRIBUTION. Afghanistan. U.S.S.R.: Kirghizia, Tadzhikistan. Pakistan.
Kashmir. N. India. Tibet.
LARVAL FOOD PLANT. Corydalis gortschakovi (A. Tsvetajev, pers. com.)

GENERA AND SPECIES OF PARNASSIINAE 89

Parnassius inopinatus Kotzsch
(PI. 9, figs 67, 68, PI. 15, fig. 105)

Parnassius inopinatus Kotzsch, 1940 : 17.
Kailasius inopinatus (Kotzsch); Eisner, 1966 : 123.

DISTRIBUTION. Afghanistan: Firus-Kuhi Range, Koh-i-Baba Range.

Parnassius loxias Piingeler
(PI. 9, figs 69, 70)

Parnassius loxias Piingeler, 1901 : 178, pi. i, figs 5, 6.
Koramius loxias (Piingeler); Bryk, 1935 : 717.
Eitkoramius loxias (Piingeler); Eisner, 1966 : 132.

DISTRIBUTION. U.S.S.R.: Kirghizia. China: Sinkiang.

Parnassius autocrator Avinoff
(PI. 9, figs 71, 72, Text-fig. 28)

Parnassius charltonius autocrator Avinoff, 1913 : 16, pi. 2, fig. 2.
Koramius charltonius autocrator (Avinoff); Bryk, 1935 : 7*6.
Eukoramius autocrator (Avinoff); Eisner, 1966 : 91.

DISTRIBUTION. Afghanistan. U.S.S.R.: Tadzhikistan.

LARVAL FOOD PLANT. Corydalis adiantifolia (Wyatt & Omoto, 1963).

THE TENEDIUS-GROVP

Parnassius tenedius Eversmann
(PI. 10, figs 75, 76, Text-fig. 3)

Parnassius tenedius Eversmann, 1851 : 621.
Tadumia tenedius (Eversmann) ; Bryk, 1935 : 647.
Tadumia tenedius (Eversmann); Eisner, 1966 : 181.

DISTRIBUTION. Mongolia. U.S.S.R.: Russia (Yakut, Tuva, Chita). China:
Inner Mongolia.
LARVAL FOOD PLANT. Corydalis sp. (bracteatal}( A. Tsvetajev, pers. com.).

THE 5/MO-GROUP

Parnassius simo Gray

(PI. 10, figs 73, 74, Text-fig. 8)

Parnassius simo Gray, 1853 : 76.
Tadumia simo (Gray) ; Bryk, 1935 : 654.
Tadumia simo (Gray); Eisner, 1966 : 178,
B*

go P. R. ACKERY

DISTRIBUTION. U.S.S.R.: Kirghizia, Tadzhikistan. Kashmir. N. India.
Mongolia. Tibet. China: Sinkiang, Kansu.

Tribe ZERYNTHIINI

Zerynthianae Grote, 1899 : 17. Type-genus: Zerynthia Ochsenheimer.

SERICINUS Westwood

Sericinus Westwood, 1851 : 173. Type-species: Papilio telamon Donovan, by original

designation.

Sericinus Westwood; Bryk, 1934 : 77-
Sericinus Westwood; Munroe, 1960 : 13.

Sericinus montela Gray
(PI. 10, figs 77, 78, Text-figs 16, 26)

Papilio telamon Donovan, 1798 : pi. 27, fig. i. [Junior homonym of Papilio telamon Linnaeus,

1758 : 486.]

Sericinus montela Gray, 1853 : 78, pi. 13, figs 1,2.
Sericinus telamon (Donovan) ; Bryk, 1934 : 80.
Sericinus telamon montela Gray; Bryk, 1934 : 89.
Sericinus montela Gray; Eisner, 1966 : 142.
Sericinus telamon (Donovan); Eisner, 1966 : 181.
Sericinus montela Gray; Hemming, 1967 : 409.

DISTRIBUTION. China: Heilungkiang, Kirin, Liaoning, Hopei, Shangtung,
Anhwei, Kiangsu, Hunan, Hupeh, Kiangsi, Kansu. Korea.

LARVAL FOOD PLANTS. Aristolochia (Leech, 1893 : 488). A. contorta (Kurentsov,
1970).

PARNAUUS Rafinesque

Thais Fabricius, 1807 : 283. Type-species: Papilio hypsipyle Fabricius, by monotypy. [Junior

homonym of Thais Roding, 1789.]

Parnalius Rafinesque, 1815 : 128. [Replacement name for Thais Fabricius.]
Zerynthia Ochsenheimer, 1816 : 29. [Replacement name for Thais Fabricius.]
Eugraphis Billberg, 1820 : 75. Type-species: Papilio hypsipyle Fabricius, by monotypy.
Parnalius Rafinesque; Sherborn, 1929 : 4765.
Zerynthia Ochsenheimer; Sherborn, 1932 : 7041.
Allancastria Bryk, 1934 : 19, 61-62. Type-species : Thais cerisy Godart, by original designation.

Syn.n.

Zerynthia Ochsenheimer; Bryk, 1934 : 3 1 -
Parnalius Rafinesque; Neave, 19400 : 614.
Zerynthia Ochsenheimer; Neave, 19406 : 689.
Allancastria Bryk; Munroe, 1960 : 10.
Zerynthia Ochsenheimer; Munroe, 1960 : 13.
Zerynthia Ochsenheimer; Hemming, 1967 : 464.
Parnalius Rafinesque; Cowan, 1970 : n. [Zerynthia Ochsenheimer cited as synonym.]

GENERA AND SPECIES OF PARNASSIINAE 91

According to Cowan (1970) Rafinesque introduced the name Parnalius for Thais
Fabricius, which was invalid as a junior homonym. The name is available and
valid, and is a senior objective synonym of Zerynthia Ochsenheimer, 1816. It
has been correctly listed by both Sherborn (1929) and Neave (1940).

The genus Allancastria Bryk is here treated as a synonym of Parnalius Rafinesque.
The differences in venation, as figured by Bryk (1934), do not appear to be consistent
although the genitalia are certainly distinct. If Allancastria Bryk is to be recognized
as a valid genus it would seem to me that there is equal justification for raising the
status of the species groups of Parnassius to genera. In order to maintain con-
sistency in approach I am regarding Allancastria Bryk and Parnalius Rafinesque
as being subjectively synonymous.

KEY TO THE SPECIES OF PARNALIUS RAFINESQUE

1 Cell zA of forewing underside uniformly scaled, without distinct red or black

discal spot; uncus long, bifid; clasper narrow (Text-fig, n, PI. 10, figs 79, 80)

cerisy (Godart) (p. 91)
Cell 2 A of forewing underside with a distinct red or black discal spot; uncus

short, bifid; clasper broad (Text-figs 9, 10) ...... 2

2 (i) Forewing upperside usually with distinct red spots in the discal cell and with

a vitreous spot near the wing apex ; uncus in dorsal aspect narrowing towards
the base; clasper distinctly produced dorso-posteriorly (Text-fig. 10, PI. 1 1, figs
83, 84) ....... rumina (Linnaeus) (p. 92)

Forewing upperside usually with black spots in the discal cell, seldom red, and
vitreous spot absent from the wing apex; uncus in dorsal aspect narrowing
towards the apex; clasper slightly produced dorso-posteriorly (Text-fig. 9,
PI. n, figs 81, 82) . . . polyxena (Denis & Schiffermuller) (p. 91)

Parnalius cerisy (Godart) comb. n.
(PI. 10, figs 79, 80, Text-figs n, 17)

Thais cerisy Godart, [1824] : 812.
Zerynthia cerisyi (Godart) ; Stichel, 19070 : 82.
Allancastria cerisyi (Godart); Bryk, 1934 : 63.
Allancastria cerisy (Godart); Cowan, 1970 : 17, 41.

DISTRIBUTION. Cyprus. Crete. Greece. Yugoslavia. Bulgaria. Rumania.
Albania. Turkey. U.S.S.R.: Armenia. Iran. Iraq. Syria. Israel. Lebanon.

LARVAL FOOD PLANTS: Aristolochia clematis and A. hastata (Bryk, 1934).
A. maurorum (Suguru Igarashi, in litt.).

Parnalius polyxena (Denis & Schiffermuller) comb. n.
(PI. n, figs 81, 82, Text-figs 9, 19)

Papilio hypermnestra Scopoli, 1763 : 149, pi. [17], fig. 425. [Junior homonym of Papilio

hypermnestra Linnaeus, 1763 : 40.]
Papilio polyxena Denis & Schiffermuller, 1775 : 162.

92 P. R. ACKERY

Papilio hypsipyle Fabricius, 1777 : 265.

Zerynthia hypermnestra (Scopoli); Bryk, 1934 : 34.

Zerynthia hypermnestra (Scopoli); Eisner, 1966 : 123.

Zerynthia polyxena (Denis & Schiffermiiller) ; Hemming, 1967 : 436.

Hemming (1967) gives a detailed explanation of the nomenclatorial history of
this species and how polyxena was finally established as the valid name.

DISTRIBUTION. S. France. Austria. Italy. Sicily. Yugoslavia. Hungary.
Rumania. Albania. Greece. Czechoslovakia.

LARVAL FOOD PLANTS. Aristolochia pistolochia. A. rotunda and A. clematis
(Higgins & Riley, 1970). A. sicula (Bryk, 1934).

Parnalius rumina (Linnaeus) comb. n.
(PI. n, figs 83, 84, Text-fig. 10)

Papilio rumina Linnaeus, 1758 : 480.

Thais maturna Butler, 1870 : 232.

Zerynthia rumina (Linnaeus) ; Bryk, 1934 : 5-

DISTRIBUTION. S. France. Spain. Portugal. Algeria. Morocco. Tunisia.
LARVAL FOOD PLANTS. Various kinds of Aristolochia (Higgins & Riley, 1970).
A . pistolochia and A . fontanesi (Bryk, 1934) .

LUEHDORFIA Criiger

Luehdorfia Criiger, 1878 : 128. Type-species: Luehdorfia eximia Criiger, by monotypy.
Luehdorfia Criiger; Bryk, 1934 : 99.
Luehdorfia Criiger; Munroe, 1960 : 13.

KEY TO THE SPECIES OF LUEHDORFIA CRUGER

i Claspers of male covered with thick golden brown pubescence ; ventral spinose setae,
placed internally on claspers, elongate (Text-fig. 12); sphragis of female keeled
(PI. 1 1, figs 85, 86) puziloi (Erschoff) (p. 92)

- Claspers of male covered with thick black pubescence ; ventral setae, placed internally
on claspers, shorter (Text-fig. 13); sphragis of female without keel (PI. n, figs 87,
88, 89, 90) japonica Leech (p. 93)

Luehdorfia puziloi (Erschoff)
(PI. ii, figs 85, 86, Text-fig. 12)

Thais puziloi Erschoff, 1872 : 315.
Luehdorfia eximia Criiger, 1878 : 128.
Luehdorfia puziloi (Erschoff); Bryk, 1934 : IO2>

DISTRIBUTION. U.S.S.R.: Russia (Primorye). Korea. Japan: Hokkaido,
Honshu.
LARVAL FOOD PLANTS. Asarum (Graeser, 1888). A. sieboldi (Kurentsov, 1970).

GENERA AND SPECIES OF PARNASSIINAE 93

Luehdorfia japonica Leech
(PI. n, figs 87, 88, 89, 90, Text-figs 13, 23)

Luehdorfia japonica Leech, 1889 : 25, pi. i, figs i, ib, 1C.
Luehdorfia japonica japonica Leech; Bryk, 1934 : IO2 -

DISTRIBUTION. Japan: Honshu. Taiwan. China: Liaoning, Kirin, Hupeh,
Anhwei, Kiangsu, Kiangsi.

LARVAL FOOD PLANTS. Asarum nipponicum, A . tamaense, A . blumei, A . caulescens
and A. sieboldi (Suguru Igarashi, in litt.).

Subsp. chinensis Leech (China: Hupeh, Anhwei, Kiangsu, Kiangsi). Hindwing
upperside with red submarginal band. Internervular marginal spots of hindwing
yellow.

The latter taxon, variously treated by authors, was regarded by Bryk (1934)
as a subspecies of puziloi Erschoff. The male genitalia, however, show a close
resemblance to those of japonica Leech both in the shape of the claspers and in
the length of the setae placed internally thereon. These characters, together with
the unkeeled sphragis of the female, seem to indicate that this taxon was correctly
placed by Rothschild (1918) as a subspecies of japonica Leech, and it is here so
treated.

BHUTANITIS Atkinson

Armandia Blanchard, 1871 : 809, nota 3. Type-species: Armandia thaidina Blanchard, by

monotypy. [Homonym of Armandia Filippi, 1862.]

Bhutanitis Atkinson, 1873 : 570. Type-species: Bhutanitis lidderdalii Atkinson, by monotypy.
Bhutanitis Atkinson; Bryk, 1934 : IX 3-
Bhutanitis Atkinson; Munroe, 1960 : 13.

KEY TO THE SPECIES OF BH UTANITIS ATKINSON

1 Hindwing upperside with a series of orange marginal internervular markings . 2
- Hindwing upperside without orange marginal internervular markings, being

yellow or grey in these areas ......... 3

2 (i) Vein M 3 of hindwing produced to broad spatulate tail; clasper of male bluntly

produced posteriorly and bearing a tuft of thick black pubescence (Text-fig. 14,

PI. 12, figs 92, 93) thaidina (Blanchard) (p. 93)

Vein M a of hindwing produced to a narrow tail; clasper of male pointed and
bearing sparse pubescence only (Text-fig. 15, PI. 13, figs 94, 95)

lidderdalii Boisduval (p. 94)

3 (i) Vein C lb of hindwing produced to a round lobe; pale bands of wings broad,

resembling Luehdorfia; female bearing a sphragis (PL 12, fig. 91)

mansfieldi (Riley) (p. 94)
Vein C^b of hindwing produced to a distinct tail; pale bands of wings narrow;

female without sphragis (PI. 14, figs 96, 97) . . ludlowi Gabriel (p. 94)

Bhutanitis thaidina (Blanchard)
(PI. 12, figs 92, 93, Text-figs 14, 25)

Armandia thaidina Blanchard, 1871 : 809.
Bhutanitis thaidina (Blanchard); Bryk, 1934 : IX ^-

94 P. R- ACKERY

DISTRIBUTION. China: Shensi, Szechwan, Yunnan.
LARVAL FOOD PLANT. Aristolochia sp. (Bryk, 1934).

Bhutanitis Udder dalii Atkinson
(PI. 13, figs 94, 95, Text-fig. 15)

Bhutanitis lidderdalii Atkinson, 1873 : 570, pi. 50.
Bhutanitis lidderdalii Atkinson; Bryk, 1934 : 118.

DISTRIBUTION. Bhutan. Sikkira. N. India: Assam, Nagaland, Manipur.
N. Burma. China: Szechwan, Yunnan.

Bhutanitis ludlowi Gabriel

(PI. 14, figs 96, 97)
Bhutanitis ludlowi Gabriel, 1942 : 189.

DISTRIBUTION. Bhutan: Trashiyangsi Valley.

As far as I am aware, the type-series of ludlowi Gabriel is unique, no other
representatives of this species being known to me.

Bhutanitis mansfieldi (Riley)

(PI. 12, fig. 9 I)

Armandia mansfieldi Riley, 1939^ : 207, pi. 4.
Bhutanitis mansfieldi (Riley); Riley, 19396 : 267.

DISTRIBUTION. China: Yunnan.

This species, known to me from the female holotype only, bears a curious
resemblance to Luehdorfia Criiger in both pattern and wing shape; furthermore
it is the only Bhutanitis species in which the female bears a sphragis. When more
material becomes available, examination of the male genitalia may show whether
it has been correctly placed here.

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GENERA AND SPECIES OF PARNASSIINAE 97

LINNAEUS, C. 1758. Systema Naturae ed. 10 : 824 pp. Holmiae.

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9 8 P. R. ACKERY

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8

FIGS 1-8. Male genitalia, left clasper removed. I, Archon apollinus amasinus (Staudinger
& Rebel). 2, Hypermnestra helios maxima Grum-Grshimailo. 3, Parnassius tenedius
tenedius Eversmann. 4, P. hardwickii hardwickii Gray. 5, P. szechenyii szechenyii
Frivaldsky. 6, P. glacialis glacialis Butler. 7, P. acdestis lathonius Bryk. 8, P. simo
simonius Staudinger.

14

12

16

FIGS 9-16. Male genitalia, left clasper removed. 9, Parnalius polyxena polyxena (Denis
& Schiffermiiller). 10, P. rumina australis (Esper). 11, P. cerisy speciosa (Stichel).
12, Luehdorfia puziloi puziloi (Erschoflf). 13, L.japonicajaponica Leech. 14, Bhutanitis
thaidina thaidina (Blanchard). 15, B. lidderdalii lidderdalii Atkinson. 16, Sericinus
montela magnus Fruhstorfer.

GENERA AND SPECIES OF PARNASSIINAE

FIGS 1 7-20. Venation of right fore- and hindwings. 1 7, Parnalius cerisy speciosa (Stichel) .
1 8, Archon apollinus amasina (Staudinger & Rebel). 19, Parnalius polyxena polyxena
(Denis & Schiflfermiiller) . 20, Hypermnestra helios maxima Grum-Grshimailo.

102

P. R. ACKERY

FIGS 21-24. Venation of right fore- and hindwings. 21, Parnassius imperator imperator
Oberthiir. 22, P. cephalus ares Bryk & Eisner [forewing only]. 23, Luehd orfia japonica
japonica Leech. 24, Parnassius charltonius deckerti Verity [forewing only].

GENERA AND SPECIES OF PARNASSIINAE

103

25

FIGS 25-26. Venation of right fore and hindwmgs. 25, Bhutanitis thaidina thaidina
(Blanchard). 26, Sericinus montela magnus Fruhstorfer.

io 4

P. R. ACKERY

31

32

FIGS 27-32. 27-28, forelegs, showing foretibial epiphysis of (27) Parnassius Orleans
Oberthur, (28) P. autocrator Avinoff. 29-30. Male tarsal claws of forelegs of (29)
Hypermnestra helios (Nickerl), (30) Parnassius cephalus Grum-Grshimailo. 3132.
Dorsal view of uncus of (31) P. phoebus (Fabricius), (32) P. mnemosyne (Linnaeus).

GENERA AND SPECIES OF PARNASSIINAE

105

INDEX
Synonyms are in italics.

acco, 86
acdestis, 87
actius, 8 1
Allancastria, 90
apollinus, 75
apollo, 83
apollonius, 80
Archon, 75
ariadne, 85
Armandia, 93
autocrator, 89

Bhutanitis, 93
bremeri, 81

cephalus, 86
cerisy, 91
charltonius, 88
clarius, 85
clodius, 85

delphius, 87
Dorarchon, 75
Doritis, 75, 76

epaphus, 82
Eugraphis, 90
Eukoramius, 76
eversmanni, 84
eximia, 92

glacialis, 84

hardwickii, 86
helios, 76
honrathi, 80
Hypermnestra, 75
hypermnestra, 91
hypsipyle, 92

imperator, 88
inopinatus, 89
Ismene, 75

P. R. ACKERY

Department of Entomology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON SW7 5BD

jacquemontii, 81
japonica, 93

Kailasius, 76
Koramius, 76

lidderdalii, 94
Lingamius, 76
loxias, 89
ludlowi, 94
Luehdorfia, 92

mansfieldi, 94
maturna, 92
mnemosyne, 83
montela, 90

nomion, 82
nordmanni, 85

Orleans, 85

Parnalius, 90
Parnassius, 76
patricius, 87
phoebus, 8 1
polyxena, 91
puziloi, 92

rumina, 92

Sericinus, 90
simo, 89

stubbendorfii, 84
szechenyii, 86

Tadumia, 76
telemon, 90
tenedius, 89
thaidina, 93
Thais, 90
tianschanicus, 82
thia, 75

Zerynthia, 90

PLATE i

Upper- and undersides (natural size)
Archon Htibner

FIG. i. apollinus apollinus (Herbst) < (Turkey: Taurus Mts)
FIG. 2. apollinus apollinus (Herbst) $ (Turkey: Taurus Mts)

Hypermnestra Menetries

FIG. 3. helios maxima Grum-Grshimailo (U.S.S.R.: Uzbekistan, Buchara)
FIG. 4. helios maxima Grum-Grshimailo $ (U.S.S.R. : Uzbekistan, Buchara)

Parnassius Latreille

FIG. 5. apollonius narynus Fruhstorfer g (U.S.S.R.: Kirghizia, Fort Naryn)
FIG. 6. apollonius narynus Fruhstorfer $ (U.S.S.R.: Kirghizia, Fort Naryn)
FIG. 7. honrathi honrathi Staudinger (U.S.S.R.: Tadzhikistan, Hasret Sultan mont)
FIG. 8. honrathi honrathi Staudinger $ (U.S.S.R.: Tadzhikistan, Hasret Sultan mont)

Bull. BY. Mus. nat. Hist. (Ent.) 31, 4

PLATE i

.

. < '

..

t

i

PLATE 2

Upper- and undersides (natural size)
Parnassius Latreille

FIG. 9. bremeri bremeri Bremer (China-U.S.S.R. : Amur)

FIG. 10. bremeri bremeri Bremer $ (China-U.S.S.R.: Amur)

FIG. 1 1 . phoebus sacerdos Vorbrodt <$ (Switzerland : St Moritz)

FIG. 12. phoebus sacerdos Vorbrodt $ (Switzerland: San Bernhardino)

FIG. 13. actius actius (Eversmann) <$ (China-U.S.S.R.: Borokhoro Mts)

FIG. 14. actius actius (Eversmann) $ (China-U.S.S.R. : Tien Shan)

FIG. 15. jacquemontii baroghila Tytler g (Pakistan: Chitral)

FIG. 16. jacquemontii baroghila Tytler $ (Pakistan: Chitral)

Bull. Br. Mus. nat. Hist. (Ent.) 31, 4

PLATE 2

PLATE 3

Upper- and undersides (natural size)
Parnassius Latreille

FIG. 17. nomion dis Bryk & Eisner,^ (U.S.S.R.: Irkutsk)

FIG. 18. nomion dis Bryk & Eisner^ (U.S.S.R. : Irkutsk)

FIG. 19. epaphus sikkimensis Elwes $ (S. Tibet: Mago)

FIG. 20. epaphus sikkimensis Elwes $ (S. Tibet)

FIG. 21. tianschanicus tianschanicus Oberthur (U.S.S.R.: Turkestan)

FIG. 22. tianschanicus tianschanicus Oberthur ^ (U.S.S.R.: Turkestan)

FIG. 23. apollo apollo (Linnaeus) ^ (Sweden)

FIG. 24. apollo apollo (Linnaeus) $ (Sweden)

Bull. Br. Mus. nat. Hist. (Ent.) 31, 4

PLATE 3

24

PLATE 4

Upper- and undersides (natural size)
Parnassius Latreille

FIG. 25. mnemosyne similis Bryk & Eisner <$ (Hungary)

FIG. 26. mnemosyne similis Bryk & Eisner $ (Hungary)

FIG. 27. stubbendorfii stubbendorfii Menetries ^ (U.S.S.R.: Russia, Buryat, Sajan)

FIG. 28. stubbendorfii stubbendorfii Menetries $ (U.S.S.R.: Russia, Buryat, Sajan)

FIG. 29. glacialis glacialis Butler ^ (Japan: Hokkaido, Sapporo)

FIG. 30. glacialis glacialis Butler ^ (Japan: Hokkaido, Yesso)

FIG. 31. eversmanni felderi Brerner ^ (U.S.S.R.: Russia, Yevrey, Pompejefka)

FIG. 32. eversmanni felderi Bremer ^ (U.S.S.R.: Russia, Yevrey, Radde)

Bull. Br. Mus. nat. Hist. (Ent.) 31, 4

PLATE 4

28

PLATE 5

Upper- and undersides (natural size)
Parnassius Latreille

FIG. 33. eversmanni eversmanni Menetries <$ (U.S.S.R. : Russia, Buryat, Sajan)

FIG. 34. eversmanni eversmanni Menetries $ (U.S.S.R.: Russia, Buryat, Sajan)

FIG. 35. nordmanni nordmanni (Nordmann) <$ (Caucasus)

FIG. 36. nordmanni nordmanni (Nordmann) $ (Caucasus, Adshara Mts)

FIG. 37. ariadne ariadne Lederer (U.S.S.R. -Mongolia: Altai)

FIG. 38. ariadne ariadne Lederer ^ (U.S.S.R. : Turkestan)

FIG. 39. clodius clodius Menetries <$ (U.S.A. : West Washington)

FIG. 40. clodius clodius Menetries $ (U.S.A. : Oregon)

Bull. Br. A/MS. nat. Hist. (Ent.) 31,4

PLATE 5

PLATE 6

Upper- and undersides (natural size)
Parnassius Latreille

FIG. 41. Orleans Orleans Oberthiir g (China: Szechwan, How-Kow)

FIG. 42. Orleans Orleans Oberthiir $ (China: Szechwan, How-Kow)

FIG. 43. hardwickii hardwickii Gray (N. India: Almora)

FIG. 44. hardwickii hardwickii Gray $ (Nepal: Jargeng Khola)

FIG. 45. szechenyii szechenyii Frivaldsky < (China: Tsinghai, Kuku-Nor)

FIG. 46. szechenyii szechenyii Frivaldsky 5 (Tibet: Arndo)

FIG. 47. cephalus ares Bryk & Eisner 3* (China: Kansu)

FIG. 48. cephalus ares Bryk & Eisner $ (China: Kansu)

Bull. Br. Mus. nat. Hist. (Ent.) 31, 4

PLATE 6

PLATE 7

Upper- and undersides (natural size)
Parnassius Latreille

FIG. 49. cephalus maharaja Avinoff ^ (Kashmir: Ladakh)

FIG. 50. cephalus maharaja Avinoff $ (Kashmir: Ladakh)

FIG. 51. acco tagalangi Bang-Haas <$ (Kashmir: Ladakh)

FIG. 52. acco tagalangi Bang-Haas $ (Kashmir: Ladakh)

FIG. 53. acco hunnyngtoni Avinoff^ (Tibet: Kyetrak)

FIG. 54. acco hunnyngtoni Avinoff $ (Tibet: Chumbi Valley)

FIG. 55. patricius kardakoffi Bryk & Eisner^ (U.S.S.R.: Kirghizia, Kungei-Ala-Tau)

FIG. 56. patricius kardakoffi Bryk & Eisner $ (U.S.S.R. : Kirghizia, Kungei-Ala-Tau)

Bull. Br. Mus. nat. Hist. (Ent.) 31, 4

PLATE 7

PLATE 8

Upper- and undersides (natural size)
Parnassius Latreille

FIG. 57. delphius albulus Honrath (China: Sinkiang, Tien Shan)

FIG. 58. delphius albulus Honrath ^ (China: Sinkiang, Tien Shan)

FIG. 59. delphius beate Eisner,^ (Kashmir: Potu-la Pass)

FIG. 60. delphius beate Eisner ^ (Kashmir: Chalsi)

FIG. 61. acdestis acdestis Grum-Grshimailo <$ (Tibet: Amdo)

FIG. 62. acdestis acdestis Grum-Grshimailo $ (Tibet: Amdo)

FIG. 63. imperator imperator Oberthiir (China: Szechwan, Ta-tsien-lou)

FIG. 64. imperator imperator Oberthiir $ (China: Szechwan, Ta-tsien-lou)

Bull. Br. Mus. nat. Hist. (Ent.) 31, 4

PLATE 8

PLATE 9

Upper- and undersides (natural size)
Parnassius Latreille

FIG. 65. charltonius serenissimus (Bryk)

FIG. 66. charltonius seyenissimus (Bryk)

FIG. 67. inopinatus inopinatus Kotzsch

FIG. 68. inopinatus inopinatus Kotzsch

FIG. 69. loxias loxias Piingeler (China

FIG. 70. loxias loxias Piingeler $ (China

FIG. 71. autocrator autocrator Avinoff
FIG. 72.

(Kashmir: Kutie Pass)
(Kashmir; Dugi Pass & Reylinj;
(Afghanistan : Marak)
(Afghanistan: Marak)
Sinkiang, Aksu)
: Sinkiang, Tien Shan)
(Afghanistan: Hindu Kush)
autocrator autocrator Avinoff $ (Afghanistan: Hindu Kush)

Bull. Br. Mus. nat. Hist. (Ent.) 31, 4

PLATE 9

72

D

PLATE 10

Upper- and undersides (natural size)

Parnassius Latreille

FIG. 73. simo simonius Staudinger (U.S. S.R.: Turkestan)

FIG. 74. simo simonius Staudinger $ (U.S.S.R. : Kirghizia, Transalai)

FIG. 75. tenedius tenedius Eversmann g (U.S.S.R.: Sajan Mts, Arasagun-gol)

FIG. 76. tenedius tenedius Eversmann $ (U.S.S.R.: Sajan Mts, Arasagun-gol)

Sericinus Westwood

FIG. 77. montela shantungensis Hering $ (China: Shantung)
FIG. 78. montela montela Gray $ (China: Hopei, Peking)

Parnalius Rafinesque

FIG. 79. cerisy speciosa (Stichel) <$ (Lebanon: Beirut)
FIG. 80. cerisy speciosa (Stichel) $ (Lebanon: Beirut)

Bull. Br. Mus. nat. Hist. (Ent.) 31, 4

PLATE 10

PLATE ii

Upper- and undersides (natural size)

Parnalius Rafinesque

FIG. 8 1. polyxena polyxena (Denis & Schiffermuller)

FIG. 82. polyxena polyxena (Denis & Schiffermuller)

FIG. 83. rumina rumina (Linnaeus) <$ (Spain)

FIG. 84. rumina rumina (Linnaeus) (Spain)

Luehdorfia Criiger

FIG. 85. puziloi puziloi (Erschoff) (U.S.S.R. -China: Amur)

FIG. 86. puziloi puziloi (Erschoff) $ (U.S.S.R. -China: Amur)

FIG. 87. japonica japonica Leech g (Japan: Honshu, Oyama)

FIG. 88. japonica japonica Leech $ (Japan: Honshu, Oyama)

FIG. 89. japonica chinensis Leech ^ (China: Kiangsu, Nanking)

FIG. 90. japonica chinensis Leech $ (China: Hupeh, Chang- Yang)

(Yugoslavia: Kragouyevatz)
(Hungary)

Bull. Br. Mus. nat. Hist. (Ent.) 31, 4

PLATE ii

PLATE 12

Upper- and undersides (natural size)
Bhutanitis Atkinson

FIG. 91. mansfieldi (Riley) $ holotype (China: Yunnan)

FIG. 92. thaidina thaidina (Blanchard) g (China: Yunnan, Tsekou)

FIG. 93. thaidina thaidina (Blanchard) $ (China: Szechwan, Ta-tsien-lou)

Bull. Br. Mus. nat. Hist. (Ent.) 31, 4

PLATE 12

PLATE 13

Upper- and undersides (natural size)
Bhutanitis Atkinson

FIG. 94. lidderdalii lidderdalii Atkinson g (India: Naga Hills, Kohima)
FIG. 95. lidderdalii, lidderdalii Atkinson $ (Bhutan)

Bull. Br. Mus. nat. Hist. (Ent.) 31, 4

PLATE 13

PLATE 15

Lateral view of sphragis
Parnassius Latreille

FIG. 98. apollo (Linnaeus)

FIG. 99. mnemosyne (Linnaeus)

FIG. 100. szechenyii Frivaldsky

FIG. 101. acdestis Grum-Grshimailo

FIG. 102. delphius (Eversmann)

FIG. 103. imperator Oberthiir

FIG. 104. charltonius Gray

FIG. 105. inopinatus Kotzsch

Bull. Br. A/MS. nat. Hist. (Ent.) 31, 4

PLATE 15

100

102

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19-

17. WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 1981

68 plates, 15 text-figures. October, 1971. 12.

18. SANDS, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae).
Pp. 244: 9 plates, 661 text-figures. July, 1972. 9.90.

19. CROSSKEY, R. W. A Revisionary Classification of the Rutiliini (Dipteral
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20. VON HAYEK, C. M. F. A Reclassification of the Subfamily Agrypninae
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21. CROSSKEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia,
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Pp. 221: 95 text-figures. December, 1973. 9.55.

PRINTED BY Unwin Brothers Limited THE GRBSHAM PRESS OLD WOKING SURREY ENGLAND

23DECi97<

THE NESOTHRIPS COMPLEX ^~ --*'
OF SPORE-FEEDING THYSANOPTERA
(PHLAEOTHRIPIDAE : IDOLOTHRIPINAE)

L. A. MOUND

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 31 No. 5

LONDON : 1974

*

23 DEC

V

THE NESOTHRIPS COMPLEX

OF SPORE-FEEDING THYSANOPTERA

(PHLAEOTHRIPIDAE : IDOLOTHRIPINAE)

BY

LAURENCE ALFRED MOUND

k

Pp 107-188; 74 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 31 No. 5

LONDON : 1974

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 Vol. 31 No. 5 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.)

Trustees of the British Museum (Natural History), 1974

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 12 December, 1974 Price 4-80

THE NESOTHRIPS COMPLEX

OF SPORE-FEEDING THYSANOPTERA

(PHLAEOTHRIPIDAE : IDOLOTHRIPINAE)

By L. A. MOUND

CONTENTS

Page

SYNOPSIS ........... 109

INTRODUCTION .......... 109

RECOGNITION OF GENERA . . . . . . . . . no

DISTRIBUTION OF GENERA . . . . . . . . .no

SPECIATION IN LITTER THRIPS . . . . . . . . in

THRIPS DISPERSION BY MAN . . . . . . . . Ill

ACKNOWLEDGEMENTS . . . . . . . . . 112

ABBREVIATIONS .......... 112

CHECK-LIST OF THE SPECIES DISCUSSED IN THIS PAPER . . . . 113

KEY TO THE GENERA OF THE Nesothrips COMPLEX . . . . 115

GENERA AND SPECIES DISCUSSED ALPHABETICALLY . . . . 1 17

SPECIES REMOVED FROM THE NeSOthfipS COMPLEX. .... l8l

REFERENCES ........... 182

INDEX 186

SYNOPSIS

Difficulties in the definition of genera among the Cryptothripini are correlated with the
biology of these spore-feeding thrips which live in leaf litter and on freshly dead wood. Neso-
thrips is divided into five genera, each with a restricted geographical distribution, but the
structural characteristics on which these genera are based are weak and overlapping. Keys
are provided to twelve genera and one hundred and one species, and three species are transferred
to the subfamily Phlaeothripinae. Two new genera and seven new species are described and
sixteen new synonymies are established. These synonymies suggest that some species have
been transported between various tropical countries by human commerce.

INTRODUCTION

THE GENUS Nesothrips has been interpreted by recent authors in different ways.
Stannard (1957) used the name for a large group of species with a wide range of
morphological characteristics and placed eight generic names in synonymy. Priesner
(1961), however, retained a large element of the traditional classification and recog-
nised five genera in the group when proposing a classification of the suborder Tubuli-
fera. Unfortunately neither of these authors was able to examine the full range
of forms in the Nesothrips-complex because the collections of J. D. Hood and R. S.

no L. A. MOUND

Bagnall were not readily available at the time. In the present study one hundred
species are recognized in the complex and through the courtesy of many colleagues
the author has been able to study type-material or authentic specimens in all
except two instances. As a result a more traditional view has been adopted using
relatively small genera. The generic classification of the whole tribe Cryptothripini
is in need of revision, but the present study is concerned solely with those species
which have been associated with Nesothrips or the closely related genus Rhaebothrips.

RECOGNITION OF GENERA

In the Cryptothripini the individual genera intergrade in a confusing pattern
and it was for this reason that Stannard (1957) used Nesothrips in a broad sense.
However, this has several disadvantages; it obscures the fact that certain species
of Nesothrips are morphologically more similar to species in other genera than they
are to other species of Nesothrips; it conceals the fact that closely related groups
of species tend to occur nearer to each other geographically than do less related
species; it conceals the structural diversity of the Nesothrips group. If the genus
is used in Stannard's sense then Rhaebothrips and Acallurothrips must be included.
Moreover some species of Dichaetothrips and Scotothrips then become difficult to
distinguish from the enlarged genus. In this way the concept of a genus approaches
Priesner's (1961) concept of a tribe or subtribe, and the name Nesothrips no longer
calls to mind any concise set of structural, biological or geographical attributes.
In order that genera reflect evolutionary relationships between groups of species
it is sometimes necessary to admit that there are intermediate species. The use
of subgenera may be helpful at times in these circumstances, but this would not be
practical in the Cryptothripini in view of the complexity of the relationships between
the various forms. The origin of these complex relationships probably lies in the
lack of host or habitat specificity in many of these fungal-spore-feeding thrips
(Mound & O'Neill, 1974 and also below).

DISTRIBUTION OF GENERA

The generic analysis in this paper has been facilitated partly by establishing the
synonymy of several tropical tramp species, and partly by transferring to Neosmerin-
thothrips several South American species described in Gastrothrips. The result is
not completely satisfactory, but in that it establishes a correlation between structural
similarity and geographical distribution this system probably reflects evolutionary
relationships. The genus Nesothrips sensu stricto is based on a species from Hawaii
and is used here for a group of species from the Pacific, Australia and New Zealand.
This is itself a difficult group of species which are not all closely related, and some of
which are very closely related to Rhaebothrips species from the Pacific and New
Zealand. Carientothrips is another related genus but this has its centre of diversity
in Australia rather than the Pacific Islands. In contrast Bolothrips is a holarctic
genus usually associated with grass tussocks, but with several species in South
Africa. Gastrothrips is a Neotropical genus with a few species entering North

NESOTHRIPS COMPLEX in

America and one species-group widespread (? introduced) in the old world. The
remaining large genus, Neosmerinthothrips, is more confusing with species apparently
native to various parts of the tropics. The characters on which these genera are
separated are summarised in the key below, although few of the genera can be
distinguished on single characters.

SPECIATION IN LITTER THRIPS

The species of the subfamily Idolothripinae (= Megathripinae) , to which the
Nesothrips complex belongs, all feed on fungal spores, judging from the gut contents
of the specimens which have been studied. They usually live in leaf litter or on
freshly dead wood and they are found mainly in the tropics. This habitat is wide-
spread and readily available across large parts of the earth's surface, and there is
no evidence that the fungi on which the thrips feed are restricted in their distribution.
This has important effects on the biology of the thrips, their dispersive activity
and resultant speciation, as well as the ease with which they are transported by
man. Although the habitat is widespread the nutritional status of the fungi at
any one site varies with the state of decay of the substratum. This fluctuation in
suitability of the habitat has probably stimulated the evolution of winged and
wingless morphs in many species, as well as the production of major and minor
(oedymerous and gynaecoid) individuals. It is assumed that this structural diversity
within individual species is controlled largely by the environment and that the
flexibility of the morphogenetic process is itself inherited. The evolution of such a
group, within which each species is structurally diverse, can be expected to produce
a complicated series of forms difficult to classify into genera. Moreover the lack
of host specificity and the irregular production of winged individuals is likely to
facilitate gene flow between populations of incipient species. This could have
caused the complicated patterns of variation between populations that are found
apparently irregularly in Carientothrips mjobergi (Mound, 1974) and Biconothrips
species (Mound, 19726) or in clines as in Bolothrips species (q.v.) and Allothrips
megacephalus (Mound, 19720) .

THRIPS DISPERSION BY MAN

Several species of fungus-feeding thrips from leaf litter and dead wood appear to
have been distributed around the world by human commerce. Some of these have
been referred to elsewhere (Mound, 1970 : 87, Nesothrips propinquus and Hoplandro-
thrips flavipes), and the establishment of new synonymy in the present study has
revealed further examples e.g. Scotothrips claripennis, Neosmerinthothrips collaris
and Rhaebothrips lativentris. Such species have probably been transported in
sailing ships: in dry grasses used as fodder and bedding; in seed pods and coconut
husks; in baskets or tubs of plants; on dead wood, or in ballast. Different species
seem to be associated with different shipping routes. Thus N. propinquus is found
along the route from Europe to New Zealand via South Africa. N. collaris is
found in West Africa and the West Indies, the two ends of one of the major slave

ii2 L. A. MOUND

trading routes, and this lends support to the suggestion made in this paper that
Neosmerinthothrips hilaris from West Africa and N. diversicolor from Brazil may
be variants of one species. A similar situation is found in the Phlaeothripine
genus Rhinoceps. The first species in this genus, R. jansei Faure, 1949, was described
from south-east Africa, the second species, R. cornutus zur Strassen, 1972, from
Colombia, and there is a third (?) species in the BMNH collection from Ghana.
However, in view of the very small differences between these 'species' it seems more
likely that they are merely local variants which have been distributed by man.
A distribution involving south-east Africa and the West Indies is also found in
Scotothrips claripennis and Neosmerinthothrips collaris, and again this relationship
calls to mind the slave trading pattern of the seventeenth and eighteenth centuries.
It would not have been possible to transport more than a million men and women
in conditions of appalling squalor without transporting fungus-feeding and detritus-
living insects. The fact that S. claripennis is also found in India may reflect even
earlier patterns of human trading. Ships have crossed the Indian ocean for several
thousand years between such distant points as China and East Africa, and this
could well explain much of the present-day distribution of such species as Nesothrips
brevicollis.

ACKNOWLEDGEMENTS

This study would not have been possible without the active support of many
colleagues. Dr Jacot-Guillarmod of Grahamstown, South Africa and the late Professor
Priesner of Linz, Austria not only loaned many valuable specimens from their
collections, but also gave the author the benefit of their experience with the Crypto-
thripini in the initial stages of the project. Kellie O'Neill of Washington, Professor
Ananthakrishnan of Madras and Dr zur Strassen of Frankfurt also loaned numerous
specimens. To all of these the author is deeply grateful. Figures 1-58 in this
paper were all drawn to the same magnification by my colleague Mrs Jenny Palmer,
and figures 59-74 were all drawn by the author to a different magnification.

ABBREVIATIONS

The following abbreviations are used for depositories.

AMG Albany Museum, Grahamstown, South Africa

ANIC Australian National Insect Collection, C.S.I.R.O., Canberra

BMNH British Museum (Natural History)

BPBM Bernice P. Bishop Museum, Hawaii

CAS California Academy of Sciences, San Francisco

DSIR Department of Scientific & Industrial Research, Auckland, New Zealand

FDA Florida Department of Agriculture, Gainsville, U.S.A.

HP Dr H. Priesner collection, Linz, Austria

INKS Illinois Natural History Survey, Urbana, U.S.A.

NR Naturhistoriska Riksmuseum, Stockholm

Pretoria Plant Protection Research Institute, Pretoria, South Africa

NESOTHRIPS COMPLEX

NZFI New Zealand Forest Research Institute, Rotorua

QM Queensland Museum, Brisbane, Australia

SMF Senckenberg Museum, Frankfurt

TM Termeszettudomanyi Muzeum, Budapest

TNA Professor Ananthakrishnan collection, Loyola College, Madras, India

USNM United States National Museum (National Museum of Natural History),
Washington B.C.

The following abbreviations are used in the descriptions of species for the major
setae of the pronotum.

am anteromarginal setae epim
aa anteroangular setae pa
ml midlateral setae

epimeral setae
posteroangular setae

CHECK-LIST OF THE SPECIES DISCUSSED IN THIS PAPER

Family PHLAEOTHRIPIDAE Uzel

Phloeothripidae Uzel, 1895

Subfamily IDOLOTHRIPINAE Bagnall

Idolothripidae Bagnall, 1908
Megathripidae Bagnall, 1914
Megathripinae Bagnall; Karny,

1919

BOLOTHRIPS Priesner, 1926 gen. rev.

Bolothrips (Botanothrips) Hood, 1939
bicolor (Heeger, 1852)

bicolor brevicornis Priesner, 1928

andrei Watson, 1933
cinctus Faure, 1943
cingulatus (Karny, 1916)
dentipes (Reuter, 1880)

bagnalli (Karny, 1916)
dentis Faure, 1954
embotyi Faure, 1943
gilvipes Hood, 1914

litoreus Hood, 1939 syn. n.
icarus (Uzel, 1895)

icarus pallipes (Uzel, 1895)
insularis (Bagnall, 1914)

icarus tuberculatus (Priesner, 1922)

brachyurus (Bagnall, 1927)

arenarius Priesner, 1950 syn. n.
italicus sp. n.
pratensis Hood, 1939
rachiphilus Cott, 1956
schafferi (Thomasson & Post, 1966)

comb. n.
varius Hartwig, 1948

CARIENTOTHRIPSMoulton,i944Stat.n.
acti Mound, 1974
badius (Hood, 1918) comb. n.

apterus (Girault, 1928)
biformis (Moulton, 1939) comb. n.
capricornis (Mound, 1974) comb. n.
casuarinae Mound, 1974
denticulatus (De Santis, 1963) comb. n.
fijiensis Moulton, 1944
grayi sp. n.

japonicus (Bagnall, 1921) comb. n.
loisthus Mound, 1974
magnetis Mound, 1974
miskoi Mound, 1974
mjobergi (Karny, 1920)

australicus (Priesner, 1928)

incisus (Girault, 1927)

flavitibia (Moulton, 1968)
pedicillus Mound, 1974
pictilis Mound, 1974
reedi Mound, 1974
semirufus (Girault, 1928)
vesper Mound, 1974
DICERA TO THRIPS Bagnall, 1908
validipennis (Hood, 1938) comb. n.

L. A. MOUND

DICHAETOTHRIPS Hood, 1914

glover i (Ramakrishna Ayyar & Marga-

bandhu, 1939) comb. rev.
GASTROTHRIPS Hood, 1912 gen. rev.

Probolothrips Moulton, 1941
abditus Hood, 1935

brasiliensis (Moulton, 1938)
hambletoni (Moulton, 1941)
acuticornis (Hood, 1925)
cybele (Girault, 1927)
noumeae Bianchi, 1945
alticola Hood, 1942
anolis Morgan, 1925
proteus Hood, 1933
callipus Hood, 1935
corvus Priesner, 1933

capitalis Hood, 1935 svn - n -
falcatus Ananthakrishnan, 1968 comb. n.
fulvicauda Hood, 1937
fulviceps Hood, 1937
fumipennis Hood, 1952
intonsus Hood, 1941
mandiocae (Moulton, 1941)
oeceticola De Santis, 1943
tnongolicus Pelikan, 1965 comb. n.
monticola Hood, 1942
procerus Hood, 1956
ruflcauda Hood, 1912
stygicus Hood, 1935
subulatus (Hartwig, 1948) comb. n.
texanus Hood, 1912

NEOSMERINTHOTHRIPS Schmutz,
1913 gen. rev.

Coenurothrips Bagnall, 1921
Galactothrips Moulton, 1933 syn. n.
affinis (Bagnall, 1921) comb. n.
annulipes (Hood, 1950) comb. n.

milleforme (De Santis, 1963) syn. n.
brevicotlis (Bagnall, 1921) comb, n.,

stat. rev.

co liar is (Bagnall, 1917) comb. n.
fuscicauda (Morgan, 1925) syn. n.
marshalli (Priesner, 1934) s Y n ' n *
dominicanus (Hood, 1925) syn. n.
diversicolor (Moulton, 1933) comb. n.
fljiensis (Moulton, 1944) comb. n.
fructuum Schmutz, 1913

ceylonicus (Karny, 1925) syn. n.
hilaris (Priesner, 1937) comb. n.
hoodi (Faure, 1954) comb. n.
inquilinus Ananthakrishnan, 1960
nigrisetis (Hood, 1935) comb. n.
parvidens (Hood, 1935) comb. n.
paulistarum (Hood, 1950) comb. n.

picticornis (Hood, 1936) comb. n.
plaumanni (Hood, 1950) comb. n.
robustus (Ananthakrishnan, 1964)

comb. n.

variipes (Hood, 1950) comb. n.
xylebori Priesner, 1935
NESIDIOTHRIPS gen. n.

alius (Ananthakrishnan, 1970) comb. n.
validus (Bagnall, 1921) comb. n.
NESOTHRIPS Kirkaldy, 1907
Oedemothrips Bagnall, 1910
aoristus Mound, 1974
artocarpi (Moulton, 1942)
brevicollis (Bagnall, 1914)
minor (Bagnall, 1921) syn. n.
formosensis (Priesner, 1935) syn. n.
formosensis karnyi (Priesner, 1935)

syn. n.

carverae Mound, 1974
fodinae sp. n.
hemidiscus Mound, 1974
malaccae sp. n.
melinus Mound, 1974
niger (Moulton & Steinweden, 1932)
oahuensis Kirkaldy, 1907

laticeps (Bagnall, 1910)
propinquus (Bagnall, 1916)
dimidiatus (Hood, 1918)
propinquus breviceps (Bagnall, 1924)
propinquus obscuricornis (Bagnall, 1924)
cestosa (Karny, 1920)
oleriae (Moulton, 1949)
similis (Hartwig, 1948) syn. n.
rhizophorae (Girault, 1927)
semiflavus (Moulton, 1939)
yanchepi Mound, 1974
PHACOTHRIPS gen. n.

ocelloides (Hood, 1950) comb. n.
RHAEBOTHRIPS Karny, 1913
doulli sp. n.
c as topi sp. n.
lativentris Karny, 1913
claripennis (Hood, 1919)
seychellensis (Bagnall, 1921) syn. n.
difficilis (Bagnall, 1921) syn. n.
ipomoeae (Ishida, 1932)
magnus (Moulton, 1928)
yuasai (Moulton, 1928)
fuscus Moulton, 1942
australiensis (Moulton, 1968)
leveri sp. n.
major Bagnall, 1928
nigrisetis Sakimura, 1972
zondagi sp. n.

NESOTHRIPS COMPLEX 115

SCOTOTHRIPS Priesner, 1939 diversus (Ananthakrishnan, 1972)

claripennis (Moulton, 1934) comb. n. syn. n.

trinidadensis (Hood, 1935) syn. n. firtnus (Hood, 1952) comb. n.

indicus (Ananthakrishnan, 1968) SYNCEROTHRIPS Hood, 1935 gen. rev.

syn. n. harti Hood, 1935

Species removed from Nesothrips complex

Subfamily PHLAEOTHRIPINAE Uzel, 1895

Adelothrips lativerticis (Post, 1961) Adelothrips speciosissimus (Karny, 1920)

comb. n. Liothrips debilis (Hood, 1936) comb. n.

KEY TO THE GENERA OF THE NESOTHRIPS COMPLEX

Antennal segment IV with two sense cones (Text-fig. 59) ..... 2

Antennal segment IV with three or more sense cones (Text-figs 61-74) . . 4

Antennal segment VIII less than twice as long as wide, broadly joined to VII ; head
not projecting in front of eyes; interocellar setae twice as long as distance between
two ocelli; female with a fore tarsal tooth [South Africa] Bolothrips dentis (p. 120)

Antennal segment VIII slender, twice as long as wide; head strongly projecting in
front of eyes; no elongate setae near the ocelli (Text-fig. i) ; female without a fore
tarsal tooth ............ 3

Antennal segment III with two sense cones; prothorax and all legs yellow in both
sexes, rest of body brown [Europe and eastern North America]

Bolothrips bicolor (p. 118)

Antennal segment III with one sense cone; prothorax brown in $, legs brown with
yellow markings on the femora, <J with pronotum and pterothorax yellow with
brown shadings laterally [Italy] Bolothrips it aliens (p. 122)

Antennal segment IV with three sense cones (Text-figs 64, 65) .... 5

Antennal segment IV with four or rarely five sense cones (Text-figs 66-74) . . 7

Eyes usually longer ventrally than dorsally, if eyes not prolonged ventrally then
metanotal setae weak and pelta broadly rounded ; tube always black with straight
sides (Text-fig. 13); and $ usually apterous, $ rarely macropterous [mainly
Holarctic] . . . BOLOTHRIPS (p. 117)

Eyes rarely prolonged on ventral surface of head; one pair of metanotal setae
frequently stout; pelta usually triangular and never broadly rounded; tube
variable, frequently yellow, or with apex sharply constricted (Text-figs 14-19) ; $
usually macropterous, rarely micropterous or apterous; <$ macropterous or
apterous [mainly Neotropical] ......... 6

Antennal segment VIII distinct from VII, usually long and slender, rarely less than
o 75 times as long as VII GASTROTHRIPS (p. 134)

Antennal segment VIII broadly fused to VII, the suture incomplete dorsally

SYNCEROTHRIPS (p. 181)

Head with an ommatidia-like papilla on cheek, situated midway between posterior
margin of eye and posterior margin of head (Text-fig. 47) ; tube margins strongly
convex, maximum diameter about five times apical diameter (Text-fig. 51) ; fore
wing without duplicated cilia [Brazil] .... PHACOTHRIPS (p. 170)

Head with normal cheeks, rarely with one isolated ommatidium close behind com-
pound eye; tube frequently with straight sides, if convex then maximum diameter

n6 L. A. MOUND

less than three times apical diameter; fore wing when present rarely lacking
duplicated cilia ............ 8

8 Head with a pair of stout setae anterolateral to the fore ocellus but with no other long

setae near the ocelli (rarely with these setae shorter than one side of ocellar triangle)
(Text-fig. 9) ; antennal segment III more than i 3 times as long as IV, segments

VII and VIII with a complete suture but forming a single unit with an almost
smooth outline (Text-fig. 66); pronotum broad and weakly reticulate, with
anterior and median ridge scarcely thickened even in major males, and mid-
lateral setae usually small ; fore femora of males frequently with a series of stout
spines on inner margin [Neotropics] . . . DICERATOTHRIPS (p. 133)

- Head with a pair of stout setae between or behind the ocelli, or with no long setae

near the ocelli; antennal segment III less than i -2 times as long as IV, segment

VIII usually smaller at base than VII at apex; pronotum frequently with a stout
median thickening, anterior margin often thickened in males, and midlateral
setae rarely small; fore femur of <J without spine-like setae .... 9

9 Head with a pair of long setae arising within the ocellar triangle, midway between

anterior and posterior ocelli (Text-fig. 46) ; fore tarsus of $ with a stout tooth

NESIDIOTHRIPS (p. 156)
Head with a pair of long setae arising between or behind the posterior ocelli, or

ocellar setae all short ; fore tarsus of $ frequently without a tooth 10

10 Fore tarsal tooth well developed in $; head always longer than wide, eyes never

prolonged on ventral surface of head; ocellar setae arising behind posterior
ocelli ; cheeks frequently with several stout setae ; tube slender with straight sides,
evenly narrowing from base to apex . . . . . . . . n

- Fore tarsal tooth usually absent in ?; if present then eyes prolonged on ventral

surface of head, or ocellar setae arising between posterior ocelli, or head as wide as
long and constricted to base without cheek setae, or tube heavy with stout setal
tubercles or even with convex sides . ....... 12

11 Postocellar setae longer than one side of ocellar triangle; cheeks with fine setae

(Text-fig. 8) DICHAETOTHRIPS (p. 134)

- Postocellar setae shorter than one side of ocellar triangle; cheeks frequently with

stout setae SCOTOTHRIPS (p. 177)

12 Maxillary stylets retracted into head as far as postocular setae, parallel in middle of

head and about one third of head width apart (Text-fig. 2) ; pelta of macropterae
usually with very slender lateral wings [Pacific, Australia, Falkland Is]

CARIENTOTHRIPS (p. 125)

Maxillary stylets wide apart, arranged in a V-shape and usually low in head (Text-
figs 40-42) ; pelta of macropterae without slender lateral wings . . . 13

13 Tube heavy with sides convex (Text-figs 35-37) and head with no elongate ocellar

setae ; or fore tarsal tooth present in $, and tube heavy with one or more pairs of
lateral setal bases, ocellar setae rarely elongate [Oriental, Ethiopian and Neo-
tropical regions] NEOSMERINTHOTHRIPS (p. 148)

Tube with straight sides, sometimes slightly constricted at apex (Text-figs 44, 56) ;
fore tarsal tooth rarely present in $ ; postocellar setae usually longer than diameter
of one ocellus, frequently arising between the posterior ocelli [Pacific and
Australian regions, introduced elsewhere] . . . . . . . 14

14 Postocellar setae close together arising between the posterior ocelli (Text-figs 52-

54) ; tube rarely less than three times as long as maximum width ; $ rarely with a

fore tarsal tooth RHAEBOTHRIPS (p. 171)

Postocellar setae frequently arising behind posterior ocelli, if arising between the
ocelli than the tube is shorter, less than 2 5 times as long as maximum width ; $
never with a fore tarsal tooth .... ..... 15

15 Head more than 1-3 times as long as wide . CARIENTOTHRIPS (part) (p. 125)
Head less than 1-2 times as long as wide .... NESOTHRIPS (p. 158)

NESOTHRIPS COMPLEX H 7

GENERA AND SPECIES DISCUSSED ALPHABETICALLY

BOLOTHRIPS Priesner gen. rev.

Bolothrips Priesner, 1926 : 90. Type-species: Phloeothrips bicolor Heeger, by original designa-
tion

Bolothrips (Botanothrips) Hood, 1939 : 605-606. Type-species: Bolothrips pratensis Hood,
by original designation.

The genus Bolothrips is used here for a group of fourteen Holarctic and Ethiopian
species which live at the base of grass tussocks and only rarely produce macropterae.
The more common species appear to exist as irregular clines, for example the gilvipes,
litoreus, rhachiphilus complex in North America, and the insularis, icarus, varius
complex in Europe and Africa. The interpretation of these clines nomenclatorially
must be rather arbitrary.

Most species of Bolothrips have three sense cones on the fourth antennal segment,
although the ventral sense cone is missing in the type-species bicolor as well as in
dentis Faure. Gastrothrips species also have three sense cones on the fourth antennal
segment although these are usually rather stouter than in Bolothrips species. More-
over most species of Gastrothrips have a pair of stout metanotal setae, a triangular
pelta, and well developed wings. Botanothrips has been used for a group of species
in which the eyes are not prolonged ventrally, but this is not accepted here in view
of the variation in insularis and varius. One species, dentis Faure, is retained in
Bolothrips for the present although it combines the characters of several genera.

GENERIC DEFINITION. Medium sized, black or bicoloured, rarely pale species of Cryptothri-
pini. Head usually longer than wide, projecting in front of eyes, with eyes prolonged or at
least narrowed ventrally; maxillary stylets V-shaped and wide apart. Antennae eight segmen-
ted, VIII more than 0-5 times as long as VII; segment III with two (rarely one) sense cone,
IV with three (rarely two) sense cones. Pronotum with epimeral sutures complete; $ with
fore tarsal tooth and frequently with fore tibial apical tubercle; $ without tooth or tubercle.
Usually apterous, fore wing with duplicated cilia in bicolor and icarus. Metanotal median
setae small and slender. Pelta usually broadly rounded; tube with straight sides.

KEY TO THE SPECIES OF BOLOTHRIPS

Body colour light yellowish brown, abdominal segments VIII-X lightest [South

Africa] *embyoti (p. 120)

Body colour dark brown, thorax sometimes yellow, abdominal segments VIII-IX

darker than rest of body .......... 2

Antennal segment IV with two sense cones, the ventral one missing ... 3

Antennal segment IV with three sense cones ....... 5

Fore tarsal tooth present in $ ; interocellar setae longer than distance between ocelli;

tube paler than abdomen, constricted at apex [South Africa] . . dentis (p. 120)
Fore tarsal tooth not present in $; interocellar setae not elongate; tube with sides

straight ............. 4

Prothorax and all legs yellow in both sexes, rest of body brown ; antennal segment

III with two sense cones [Europe, eastern North America] . . bicolor (p. 118)
Prothorax brown in $, legs brown with yellow markings; Q* with pro thorax and

pterothorax yellow but shaded brown laterally; antennal segment III with one

sense cone ; Italy ......... italicus (p. 122)

n8 L. A. MOUND

5 Ventral length of eyes more than i 6 times the dorsal length .... 6

- Ventral length of eyes less than i 3 times the dorsal length . . . . 1 1

6 Pronotum, pterothorax and abdominal segment I yellow ..... 7
Pronotum sometimes reddish yellow, but pterothorax brown .... 8

7 Abdominal segment II yellow; antennal segments II and IV largely brown [Europe]

cingulatus (p. 119)

Abdominal segment II brown; antennal segment II brownish yellow, IV and V

largely yellow [South Africa] ....... cinctus (p. 119)

8 Tube relatively long, at least 2 3 times as long as maximum width ; antennal segment

III largely yellow ........... 9

Tube relatively short, not more than 2 i times as long as maximum width ; antennal

segment III brown with pedicel light . . . . . . . . 10

9 Legs uniform dark yellow; antennal segments III, IV and basal two-thirds of V

yellow [western North America] ....... schqfferi (p. 123)

Legs largely brown ; antennal segments IV and V brown, III yellow with brown

shading at apex [Europe and North America] .... dentipes (p. 119)

10 Head of apterae with reduced ocelli; pronotum reddish yellow [south-eastern North

America] ........... gilvipes (p. 120)

- Head of apterae without ocelli; pronotum black [California] . * rhachiphilus (p. 123)

1 1 Body bicoloured, head and thorax much paler than abdomen [Florida, Texas]

pratensis (p. 122)

- Head and thorax as brown as abdomen . . . . . . . .12

12 Dorsal setae on tergite IX less than 0-8 times as long as tube [South Africa]

varius (p. 123)
Dorsal setae on tergite IX more than 0-9 times as long as tube [Mediterranean to

Latvia] ............. 13

13 Antennal segment IV yellow at base; dorsal setae on tergite IX generally shorter

than tube .......... icarus (p. 121)

- Antennal segment IV brown ; dorsal setae on tergite IX generally a little longer than

tube ............ insularis (p. 121)

* Not studied

Bolothrips bicolor (Heeger)

Phloeothrips bicolor Heeger, 1852 : 477-478. Syntypes ?sex, AUSTRIA (? lost).

Bolothrips bicolor (Heeger) Priesner, 1926 : 90.

Bolothrips bicolor f. brevicornis Priesner, 192801 : 687. Holotype $, HUNGARY (HP) [not

examined].
Oedaleothrips andrei Watson, 1933 : 49-50. Syntypes 4 $, 4 , U.S.A.: Iowa (FDA) [not

examined]. [Synonymized by Stannard, 1957 : 106.]

This species is widespread in central and south-eastern Europe, and according to
Stannard (1957) it is distributed through the northern part of North America.
The present author has not studied specimens from west of North Dakota or Utah.
One macroptera is listed below from Minnesota, and Priesner (1928 : 687) refers to
macropterae from Austria found in July and August. The most closely related
species are dentipes and italicus, whereas cingulatus has much shorter antennal
segments with three sense cones on the fourth segment.

SPECIMENS STUDIED.
HUNGARY: Simontornia, i <j>, i $, 5.V.I924 (F. Pillich). CZECHOSLOVAKIA: i $

NESOTHRIPS COMPLEX 119

ex Uzel Collection. FRANCE: Villefranche sur Saone, 3 <$, 2 $, ix-x.1927 (0. John);
Hyeres Plage, i & ix.igay (R. 5. Bagnall) (BMNH).

U.S.A.: Illinois, Grundy County, i <, x.igyo (Z,. ^4. Mound); Minnesota, Mao
Gregor, 7 $, 3 ^ apterae, i $ macroptera, iv.iQ36 (F. /Iw^re); M., Hay Creek, 2 $,
i <$, xi.i936 (G. Decker); Iowa, Ottumwa, 27 <, 22 $, in Moss, i.1933; I., Boone,
7 , 7 c, from Poa, vii-viii.ig34 (F. ^4mfr0); North Dakota, Northwood, i $, iv.igGi
(#. /. Pos); Utah, Blacksmith Fork Canyon, i $, iv.igyi (G. F. Knowlton) (BMNH).

Bolothrips cinctus Faure

Bolothrips (Bolothrips) cinctus Faure, 1943 : 86-87. Syntypes 2 $, SOUTH AFRICA: Transvaal
(Pretoria) [not examined],

This species has a shorter tube than any other species of Bolothrips, and it can
be distinguished moreover by its colour pattern. The two syntype apterae were
collected from Themeda triandra near Lydenburg, Transvaal.

SPECIMEN STUDIED.

SOUTH AFRICA: Transvaal, Ermelo, i $ aptera on Hyparrhenia, 21^.1949 (J. C.
Faure) (HP).

Bolothrips cingulatus (Karny)

Cryptothrips cingulatus Karny, 1916 : 92. Syntypes ? sex, AUSTRIA (HP) [not examined].
Bolothrips cingulatus (Karny) Priesner, 1926 : 90.

Priesner (1964 : 143) refers to this as a south-eastern European species and states
that the macropterae have only the first and second abdominal segments light.
The antennae of cingulatus are shorter than those of bicolor and dentipes.

SPECIMENS STUDIED.

HUNGARY: Simontornyia, i $, 3 $ apterae, vi & ix. 1924 (F. Pillich). FRANCE:
Rhone, Curis, 3 <$, i $ apterae, viii. 1927 (0. John) ; Hyeres Plage, i $ aptera, ix. 1927
(R. 5. Bagnall) (BMNH).

Bolothrips dentipes (Reuter)

Phloeothrips dentipes Reuter, 1880 : 12-14. 2 Syntypes, ? sex, FINLAND (? lost).
Bolothrips dentipes (Reuter) Priesner, 1928^ : 689692.

Cryptothrips bagnalli Karny, 1916 : 94. Syntypes, ? sex, SARDINIA (? lost).
Cryptothrips dentipes var. bagnalli Karny; Priesner, 1925 : 154.

This species is widespread in Europe between Ireland, Poland and Hungary.
There are numerous specimens in the BMNH collections from southern England,
and also one female without data but bearing a determination label 'Reuter det.
3.84'. Stannard (1968) records the species from Illinois, and one specimen has
been studied from Massachusetts.

izo L. A. MOUND

Bolothrips dentis Faure

Bolothrips dentis Faure, 19546 : 155-159. Holotype $, TRANSVAAL (Pretoria) [not examined] .

This species was described from 14 males and 26 females, all apterae, collected
at Mariepskop in Transvaal, mainly on dead branches covered in lichen. The
combination of morphological characters is most unusual. The head is truncate
at the anterior and bears a pair of stout interocellar setae, but the eyes are prolonged
ventrally. The fore tarsus of the female bears a tooth but the fourth antenna!
segment has two sense cones. The tube is pale in colour and constricted at the
apex. This one species thus shows characteristics from each of the genera of the
Nesothrips complex. It is not a true Bolothrips species but is retained in this genus
until the South African fauna is better known.

SPECIMENS STUDIED.

TRANSVAAL: Mariepskop, 2 $ paratypes on dead branches of Syzygium cor datum,
iv. 1951 (]. C. Faure) (HP, AMG).

Bolothrips embotyi Faure

Bolothrips (Bolothrips) embotyi Faure, 1943 : 87-89. Syntypes 5 $, 7 $, SOUTH AFRICA: Pondo-
land (Pretoria) [not examined].

No specimens of this species have been studied. Judging from the description
it is a true Bolothrips with the eyes prolonged ventrally, the head projecting in
front of the eyes, the fourth antennal segment with three sense cones, and the
female without a fore tarsal tooth.

Bolothrips gilvipes (Hood)

Cryptothrips gilvipes Hood, 1914 : 169-170. Holotype $, U.S.A.: Maryland (USNM) [not

examined].
Bolothrips (Bolothrips) litoreus Hood, 1939 : 609-612. Holotype $, U.S.A. : Texas (USNM)

[not examined]. Syn. n.

This species appears to show a gradient in size and colour with the darkest speci-
mens in the southern and western part of its range. The specimens from Florida
listed below have the thorax varying in colour from yellow to light brown even at
one site. In view of this variation there seems to be little point in distinguishing
the larger and darker specimens as litoreus. The Californian species rhachiphilus
is possibly another element in this cline.

SPECIMENS STUDIED.

U.S.A.: Alabama, Mobile, i ^ on dead grass (det. gilvipes by J. D. Hood),
I7.xii.i938 (/. D. Hood); Florida: Cedar Keys, 17 $, I <$, on salt marsh grass,
I2.vii.i939 (P. Oman); F., Lake Wauberg, I $, iv. 1938; F., Hudson, i <? on grass,
vii. 1939; F., Deland, i <j>, i <$ on grass, vii. 1939 (BMNH, USNM).

NESOTHRIPS COMPLEX 121

Bolothrips icarus (Uzel)

Cryptothrips Icarus Uzel, 1895 : 232-233. Syntypes <J ^, CZECHOSLOVAKIA: Bohemia (? lost).
Cryptothrips icarus var. pallipes Uzel, 1895 : 233.
Bolothrips icarus (Uzel) Priesner, 19280 : 692-695.

All of the specimens listed below have the legs more or less yellow with a variable
amount of brown shading. The species is widespread from southern Europe to
Latvia and is very closely related to the Mediterranean insularis and the African
varius, but has paler legs and the eyes usually less prolonged ventrally. Three
macropterous females have been studied from France.

SPECIMENS STUDIED.

FRANCE: Villefranche sur Saone, 3 <$, i $, i $ macroptera, vii. 1927; Rhone,
Curis, i (, 2 $ macropterae, viii. 1927 (0. John) ; La Trinite sur Mer, i <, viii. 1968
(V. F. Eastop). AUSTRIA: Innsbruck, i <$, ix. 1929 (R. S. B agnail). YUGOSLAVIA:
Subotica, i <, vi. 1963. CZECHOSLOVAKIA: Ceje, i <j>, vi. 1950 (/. Pelikan); Bzendei,
3 <?, v. 1964 (V. F. Eastop}. HUNGARY: Simontornia, 3 <j>, i <, vi. 1924 (Pillich}.
U.S.S.R.: Latvia, 4 $, i $, vi. 1928; 2 <J, 3 $, viii. 1929 (0. John) (BMNH).

Bolothrips insularis (Bagnall)

Cryptothrips insularis Bagnall, 1914 : 295. Holotype $, CANARY ISLANDS (BMNH) [examined].
Cryptothrips icarus var. tuberculatus Priesner, 1922 : 105. Holotype $, YUGOSLAVIA (HP)

[examined]. [Synonymized by Mound, 1968 : 141.]
Cryptothrips brachyurus Bagnall, 1927 : 573-574. Lectotype <, FRANCE (BMNH) [examined].

[Synonymized by Mound, 1968 : 141.]
Bolothrips arenarius Priesner, 1950 : 36-37. Syntypes $ $, EGYPT & SYRIA (HP) [examined].

Syn. n.

Priesner (1928 : 693) suggested that this species (under the name tuberculatus)
might be a geographical race of icarus. Whereas icarus is found in continental
Europe, insularis appears to be restricted to the Mediterranean area between the
Canary Islands and Syria. Both species are variable, but they can apparently
be distinguished by the characters given in the key above. Specimens from the
eastern part of the range of insularis, Cyprus and Egypt, generally have the tube
shorter and stouter than specimens from the western part of the range, Spain and
the Canary Islands. The specimens from Cyprus appear to be intermediate be-
tween insularis and typical arenarius from Egypt.

SPECIMENS STUDIED.

Holotype $ of insularis, CANARY ISLANDS (T. V. Wollaston) (BMNH). Lectotype
cJ with i $, i <j> paralectotypes of brachyurus, FRANCE: Tamaris, iii. 1927 (R. S.
Bagnall) (BMNH). Holotype <j>, allotype $ [sic] of tuberculatus, YUGOSLAVIA:
Dalmatia, Ragusa, from grass, 3Xjc,xgx8 (HP).

EGYPT: 40 km east of Meadi, i $, i $ paratypes of arenarius, on Retama, 9.^.1934;
Ismailia, i $, 23.^.1935; Bingash, <$ allotype, 17.1.1935 (HP). CANARY ISLANDS:
Lancerota, i 9 on grass, iii. 1963, i $, ii. 1968 (zur Strassen). SPAIN: Chiclana,

122 L. A. MOUND

2 <j>. FRANCE: St Cyr sur Mer, i <J>, 2 $, ix. 1927 (/?. S. Bagnall). CYPRUS: Akrotiri
Bay, 8 $, 4 $ on Juncus, vii. 1945 (Mavroumoustakis) (BMNH).

Bolothrips it aliens sp. n.

aptera. Colour brown, head and posterior abdominal segments darker than pterothorax;
femora variable in colour, mid and hind tibiae and tarsi dark brown, fore tarsi and inner surface
of fore tibiae lighter; antennal segment III yellow, IV light brown, V slightly paler at extreme
base; terminal setae of abdomen light brown.

Structure of body and antennae very similar to dentipes and bicolor, but head more slender,
pterothorax distinctly alatiform, metanotum not sculptured; antennal segment III with one
sense cone, IV with two sense cones.

MEASUREMENTS (holotype $ in pm). Body length 2900. Head, length 330; median width
230; postocular setae 100. Pronotum, length 175; width across fore coxae 320. Metathorax
total width 315. Tergite IX setae Bj 170. Tube, length 210; basal width 100. Antennal
segments length III-VIII length, 97; 97; 97; 80; 65; 45.

aptera. Thorax paler than $, pronotum yellow but shaded brown laterally, pterothorax
and abdominal segment I almost clear yellow; head and posterior abdominal segments dark
brown; all femora extensively yellow on inner surface, mid and hind tibiae and tarsi brown;
antennal segments IV and V paler than in $. Similar in structure to $ but fore tarsus with a
slender tooth and fore tibia with an apical tubercle.

MEASUREMENTS (paratype in pm). Body length 1900 (body contracted). Head length
290. Tube, length 160; basal width 90.

SPECIMENS STUDIED.

Holotype $, ITALY (taken in quarantine at Detroit, U.S.A.): Stipa tenacissima,
27.xi.i96i (/. Fergusson 4087) (USNM).

Paratypes. 5 $, 6 <? collected with the holotype (USNM, BMNH).

COMMENTS . Unfortunately this species is known only from rather poorly preserved
specimens taken in quarantine. It is described here because of its interesting
relationship to bicolor and dentipes, and because it is the only member of the genus
with one sense cone on the third antennal segment. The females are very similar
to dentipes but have a shorter tube, and some of the specimens have the pterothorax
and femora pale in colour.

Bolothrips pratensis Hood

Bolothrips (Botanothrips] pratensis Hood, 1939 : 606-609. Holotype $, U.S.A.: Texas (USNM)
[not examined].

Despite the fact that the eyes are not prolonged on the ventral surface of the head,
the present author does not agree with Hood that pratensis is closely related to the
European and African icarus-group of species. The tube is particularly stout and
the mesothoracic spiracles project as in gilvipes, and it is possible that pratensis is
derived from forms with ventrally prolonged eyes. Most of the specimens listed
below were collected on dead wood, in contrast to the type-series which came from
grasses as is usual in Bolothrips.

NESOTHRIPS COMPLEX 123

SPECIMENS STUDIED.

U.S.A.; Texas, Palacios (type-locality), i $, i < paratypes, 28.iii. & 23.^.1939
(/. D. Hood); Florida, Upper Matecumbe Key, 5 <$, 10 $ from dead branches,
xii. 1958 (Andre); F., Grassy Key, I ^, I $ on dead branches, xii. 1958 (Andre);
F., Cedar Keys, I , i $, vii. 1939 (P. Oman) (BMNH).

Bolothrips rhachiphilus Cott

Bolothrips (Bolothrips) rhachiphilus Cott, 1956 : 181-182. Holotype $, U.S.A.: California
(not known).

Type-material of this species has not been deposited in the California Academy
of Sciences (Arnaud & Lee, 1973) as was stated in the original description. The
species appears to be related to the dark forms of gilvipes.

Bolothrips schafferi (Thomasson & Post) comb. n.

Nesothrips schafferi Thomasson & Post, 1966 : 31-32. Holotype $, U.S.A.: North Dakota
(INHS) [not examined].

This species is closely related to dentipes in the form of the antennae, head and
tube, but differs in the colour of the legs and antennae. It is known only from
western North America.

SPECIMENS STUDIED.

U.S.A.: North Dakota, Benson County, Wood Lake, 3 $ paratypes on Spartina
pectinata, 2y.viii.i964 (Thomasson < McBride) (BMNH & USNM); Oregon, 2 $,
i , 15.^.1937 (?^. Andre) (BMNH).

Bolothrips varius Hartwig

Bolothrips varius Hartwig, 1948 : 110-112. Holotype $, SOUTH AFRICA: Transvaal (Pretoria)
[not examined].

The original description of this species noted that the ventral length of the eyes
varied from 'about equal to the dorsal length to about 1-4 times the dorsal length'.
The eyes of insularis from the Mediterranean region also vary considerably, and
these two species are closely related. Hartwig lists material of varius from Cape
Province, Basutoland, Natal and Transvaal. The specimen listed below from Kenya
has a slender head as in varius and insularis, but the legs and antennal segments
are pale as in icarus. It is possible that further collecting will produce intermediates
between these named forms.

SPECIMENS STUDIED.

SOUTH AFRICA: Transvaal, Pretoria, i <$ paratype on Panicum maximum, 13.^.1946
(E. K. Hartwig [data as for holotype]; 3 $, i <$ on P. maximum, 8^.1948 (E. K.
Hartwig) (BMNH). KENYA: i $ on Pinus, vii. 1963 (BMNH).

124

L. A. MOUND

FIGS 1-7. i, Bolothrips italicus. 2, Carientothrips fijiensis. 3, C. mjobergi. 4, C
grayi. 5-7. Pelta (first abdominal tergite) : 5, B. italicus. 6, C. fijiensis. 7, C.

NESOTHRIPS COMPLEX 125

CARIENTOTHRIPS Moulton stat. n.

Bolothrips (Carientothrips) Moulton, 1944 : 306. Type-species: Bolothrips (Carientothrips)
fijiensis Moulton, by monotypy.

This genus was placed as a synonym of Nesothrips by Stannard (1957) but is used
here for a group of seventeen species from Australia and the Pacific, with one
possibly unrelated form from South America. The macropterae of the type-species
fijiensis are similar to a new species grayi described below from New Guinea, and
also to the macropterae of the mjobergi complex from Australia. The micropterae
of fijiensis are very similar to the more slender forms loisthus and magnetis from
Australia and New Zealand, but the micropterae of mjobergi are much shorter and
broader and at first sight look very different. These are related to other species
in Australia with short broad heads, miskoi and pedicillus. Within mjobergi there
are forms with the eyes prolonged ventrally or with the eyes equal in size on the
dorsal and ventral surfaces, moreover the legs also vary from yellow to brown,
and there is no clear evidence that these different forms represent distinct species.
The pelta of the macropterae is frequently characteristic with very slender lateral
wings, but the pelta of micropterae and apterae is reduced and such specimens are
very similar to Nesothrips species. These two genera intergrade in Australia, but
Carientothrips is used for a group of species with the stylets relatively close together
in the head. Two species with the stylets wide apart are also included from
Australia, capricornis which seems to be related to semirufus, and badius which is
not closely related to any other species.

GENERIC DEFINITION. Medium sized, dark or pale species of Cryptothripini. Head longer
than wide, or wider than long; eyes normal or reduced, sometimes longer on ventral than on
dorsal surface ; maxillary stylets usually parallel in middle of head and about one third of head
width apart; ocellar setae usually well developed. Antennae eight-segmented, two sense cones
on segment III, four sense cones on IV; segment VIII usually elongate. Fore tarsal tooth
absent in $ (except denticulatus) , present in ^. Pronotum transverse, oedymerous individuals
rare; epimeral sutures usually complete. Wings frequently absent. Pelta frequently with
slender lateral wings; tube with straight sides.

KEY TO THE SPECIES OF CARIENTOTHRIPS

1 Antennal segment III with a shelf-like ring near the base (Text-fig. 60) . . 2

- Pedicel of antennal segment III simple ........ 3

2 Thorax and legs yellow, rest of body and head brown; antennal segments II-IV

largely yellow [Australia] ........ pictilis (p. 132)

- Head light brown, thorax medium brown, abdomen dark, legs yellowish brown;

antennal segments II-IV largely brown [Australia] .... acti (p. 126)

3 Metanotum with four to six pairs of setae medially in apterae, two pairs in

macropterae; epimeral sutures incomplete ....... 4

- Metanotum with one pair of median setae ; epimeral sutures complete ... 6

4 Antennal segment III 2-5 times as long as wide, pedical stout bearing several

transverse ridges (Text-fig 62) [Japan] .... japonicus (p. 130)

- Antennal segment III either longer or shorter, pedicel slender .... 5

5 Antennal segment III 2 i times as long as wide [Australia] . . . reedi (p. 132)

- Antennal segment III 2-8 times as long as wide [Australia] . . vesper (p. 133)

6 Head broad, less than i 2 times as long as wide ...... 7

- Head slender, more than i -3 times as long as wide . . . . . . 12

126 L. A. MOUND

7 Body, legs and antennae dark brown except pedicel of III which is paler; pelta of

macropterae and apterae rounded ; eyes not extended on ventral surface of head
[Australia] .......... casuarinae (p. 128)

- At least antennal segments III-V with yellow pedicels ; frequently with legs, several

antennal segments and anterior half of body pale ...... 8

8 Female with very small fore tarsal tooth at base of pretarsus; ocellar setae less than

20 pm long; antennal segment III stout but sharply constricted at pedicel [Argen-
tina and Falkland Islands] ....... denticulatus (p. 128)

- Female without a fore tarsal tooth ; ocellar setae more than 30 pm ; antennal segment

III tapering to base ........... 9

9 Setae B a on tergite IX about 0-5 times as long as B^ [Society Islands] biformis (p. 127)

- Setae B z on tergite IX as long as or longer than B t [Australia] . . . . 10

10 Antennal segments VI and VII broadly rounded with a sharply slender yellow

pedicel . . . . . . . . . . pedicillus (p. 132)

Pedicel of antennal segments VI and VII less slender, VI frequently yellow in basal

half ii

11 Apical third of tube sharply yellow in contrast to black basal two-thirds; femora

largely yellow .......... tniskoi (p. 131)

- Apical third of tube paler than base but not sharply yellow; femora brown but

tibiae yellow or brown ....... .mjobergi (p. 131)

1 2 Body and appendages uniformly dark brown except for pale base of antennal segment

III; eyes reduced but rounded and protruding [Australia] . . bndius (p. 127)

Head and pronotum sometimes paler than abdomen, antennae or legs with extensive

pale areas; eyes large, if reduced then not round and protruding . . . 13

13 Antennal segment IV shorter than segment III [Australia] . . capricornis (p. 127)

- Antennal segment IV longer than segment III . . . . . . 14

14 Setae B and B 2 on tergite IX less than 0-5 times as long as the tube, with apices

clearly expanded ; head and thorax yellow, abdomen brown, antennal segments I

and II yellow [Australia] ....... setnirufus (p. 132)

- Setae B^ and B z on tergite IX more than 0-5 times as long as tube, apices acute or

blunt; head and thorax usually as brown as abdomen, if paler then antennal
segment II brown ........... 15

15 Antennal segments III and IV almost clear yellow, also basal half of V; segment IV

about 3 5 times as long as wide ; body colour brown, legs yellow [Australia]

magnetis (p. 131)

- Antennal segments III and IV variously shaded with brown ; if pale then segment IV

less than 3-0 times as long as wide; body and femora usually dark . . . 16

1 6 Eyes prolonged on ventral surface of head with one large isolated facet; dorsal

surface of head and pronotum conspicuously reticulate (Text-fig. 4) ; antennal
segments IV and V with elongate slender pedicels [New Guinea] . grayi (p. 129)

- Eyes not longer ventrally than dorsally (Text-fig. 2) ; dorsal surface of head weakly

sculptured; antennal segments IV and V with short pedicels (Text-fig. 61) . 17

17 Eyes reduced to about five facets ventrally; antennal segment III yellow in basal

half, sometimes brown in apical half [Australia & New Zealand] . loisthus (p. 130)

- Eyes with at least ten facets on ventral surface; antennal segment III pale at base

and apex, but with a transverse brown band at basal third [Fiji] . fijiensis (p. 128)

S

Carientothrips acti Mound

(Text-fig. 60)
Carientothrips acti Mound, 1974 : 25-26. Holotype $, AUSTRALIA (ANIC) [examined].

The acti group of species includes japonicus (Bagnall), pictilis Mound, reedi

NESOTHRIPS COMPLEX 127

Mound and vesper Mound. Both acti and pictilis are unusual in having the terminal
setae on the tube flattened at the apex instead of acute, and in having the pedicel
of antennal segment three flared into a ring. However, japonicus has the pedicel
of segment three slightly thickened, and this probably represents an intermediate
condition. These species, apart from pictilis, have the epimeral sutures incomplete,
and the metanotum bears more than one pair of median setae.

SPECIMENS STUDIED.

Holotype $, AUSTRALIA: Australian Capital Territory, Canberra, Black Mt, on
dead leafy Eucalyptus branches, 3.1.1961 (E. M. Reed] (ANIC). Paratypes from
New South Wales, A.C.T. and northern Victoria from leaf litter and grass tussocks
are listed in detail in Mound (1974).

Carientothrips badius (Hood) comb. n.

Cryptothrips badius Hood, 1918 : 143-144. Holotype $, AUSTRALIA: Queensland (USNM)

[examined] .
Elaphrothrips apterus Girault, 1928 (42) : 2. Holotype , AUSTRALIA: Queensland (QM)

[examined]. [Synonymized by Mound, 1974 : 2 3l-
Bolothrips badius (Hood) Hartwig, 1948 : 117.

This species was treated in the genus Bolothrips by Mound (1974), but is now
removed from that genus because the fourth antennal segment bears four sense
cones. However, it is an unusual species with a relatively long head and small
rounded eyes which is not closely related to any other known species. C. badius
is widespread in Australia in grass tussocks, and a detailed list of specimens from
Queensland, New South Wales, and Western Australia is given in Mound (1974).

Carientothrips biformis (Moulton) comb. n.
Bolothrips biformis Moulton, 1939 : 146-147. Holotype $, TAHITI (BPBM) [not examined].

This species is removed from Bolothrips because it has four sense cones on the
fourth antennal segment. It is not closely related to the other species of Cariento-
thrips but has the maxillary stylets closer together than the species of Nesothrips.

SPECIMEN STUDIED.

SOCIETY ISLANDS: Rapa, Mangaoa Pk, 1000-1200 ft, 25.vii.i934 (E. C.
Zimmermann), I $ microptera paratype (BMNH).

Carientothrips capricornis (Mound) comb. n.

Bolothrips capricornis Mound, 1974 : 2 3~ 2 4- Holotype $, AUSTRALIA: Queensland (ANIC)
[examined] .

This species was described in Bolothrips because of its resemblance to the American
species B. pratensis. However, the fourth antennal segment bears four sense

128 L. A. MOUND

cones unlike any species of Bolothrips, and capricornis is probably most closely
related to Carientothrips semirufus.

SPECIMENS STUDIED.

Holotype $ microptera, AUSTRALIA: Queensland, Townsville, 2 miles west of
Saunders Beach, in leaf litter, 2i.vii.i968 (L. A. Mound] (ANIC). Paratype $
collected with the holotype (BMNH).

Carientothrips casuarinae Mound

Carientothrips casuarinae Mound, 1974 : 26-29. Holotype AUSTRALIA : New South Wales
(ANIC) [examined].

This species, which is known from macropterae as well as apterae, is dark brown
but is otherwise similar to semirufus. It has been taken on dead Casuarina branches
at several sites in eastern Australia.

SPECIMENS STUDIED.

Holotype $ aptera, AUSTRALIA: New South Wales, Normanhurst (15 miles north
of Sydney), on dead Casuarina foliage, 1.11.1959 (E. M. Reed) (ANIC). Paratypes
<$ and $ macropterae from New South Wales listed in detail in Mound (1974).

Carientothrips denticulatus (De Santis) comb. n.

Nesothrips (Bolothrips) denticulatus De Santis, 19636 : 66. Holotype $, ARGENTINA: Tierra
del Fuego (Museos de La Plata) [not examined].

This species is removed from Bolothrips because it has four sense cones on the
fourth antennal segment. It is not close to Nesothrips species because the stylets
are parallel in the middle of the head, but it seems unlikely to be closely related
to the other species of Carientothrips. Some of the specimens listed below were
identified by Prof. Luis de Santis as denticulatus. The fore tarsal tooth of the
female is less than 10 /xm long, although that of the male is about 30 /Am long.

SPECIMENS STUDIED.

EAST FALKLAND ISLANDS: Kidney Is., 55 $, 24 <$ in Poa flabellatum tussocks,
29.xi.i967 (E. Holdgate) (BMNH).

Carientothrips fijiensis (Moulton) comb. n.

Bolothrips (Carientothrips) fijiensis Moulton, 1944 : 306-307. Holotype $, FIJI (BPBM)
[not examined].

The original description refers to an apterous paratype female, although this
specimen is actually micropterous with wing lobes 160 /mi long. The eyes of the
microptera are reduced and flattened externally as in loisthus, although the eyes
of the macropterae are large and rounded as in grayi described below. The
Australian species mjobergi is similar to the macropterae oi fijiensis although with a

NESOTHRIPS COMPLEX 129

shorter head, but the apterae and micropterae of mjobergi are much shorter and
broader. The pelta of fijiensis is similar to that of grayi, and the male has a small
fore tarsal tooth.

SPECIMENS STUDIED.

FIJI: Vitilevu, Nandarivatu, 2 <j>, I macropterae, I $ microptera (all paratypes),
9.ix.i938, I $ macroptera paratype i.ix.1938; Mt Korobamba, I $ macroptera
paratype i.viii.i938 (E. C. Zimmermann) (CAS).

Carientothrips grayi sp. n.

(Text-figs 4, 7, 63)

$ macroptera. Colour dark brown with red internal pigment; tarsi light brown; antennal
segments IV and V yellow in basal third, III with yellow pedicel and yellowish brown in apical
half otherwise dark brown; fore wings deeply shaded in third and fourth fifth, but clear in
apical fifth, basal two-fifths clear with dark longitudinal stripe; major setae almost hyaline,
bases of setae at apex of tube dark.

Head longer than wide, projecting in front of large compound eyes, cheeks rounded to base
with about four small setae ; dorsal surface strongly sculptured ; postocellar setae well behind
ocelli; maxillary stylets retracted to postocular setae; eyes with one large ventral facet dis-
placed posterior to normal hind margin of eye. Antennae with segment III much shorter than
IV; segment VII with a distinct pedicel, VIII slender and slightly constricted at base. Prono-
tum similarly sculptured to head, with weak median thickening; epimeral sutures complete,
setae with blunt apices. Metanotum reticulate, median setae slender, 30 ju.m long. Fore
wing slightly constricted medially; subbasal setae short, duplicated cilia variable. Fore
tarsus with no tooth. Pelta reticulate with pronounced lateral wings; setae on tergite IX
softly rounded at apex ; tube slender with straight sides bearing fine setae.

MEASUREMENTS (holotype $ in ^tm). Body length 3250. Head, length 350; median width
260; postocular setae 65; ocellar setae 35. Pronotum, length 195; median width 320; major
setae - am 30, aa ?30, ml ?50, epim 100, pa 70. Fore wing, length 1400; distal width too;
subbasal setae 40, 45, 65; number of duplicated cilia 4-7 (3-6 in one paratype). Tergite VII
setae, B 70; B 2 100. Tergite IX setae, B l 160; B 2 190; B 3 155. Tube, length 290; basal
and apical width 90, 45; terminal setae 175. Antennal segments length, 55; 65; 95; 125;
120; 77; 58; 35; sense cones on III 75.

SPECIMENS STUDIED.

Holotype $, NEW GUINEA: Wau, under bark of Araucaria cunninghamiana,
22.ix.i970 (B. Gray) (BMNH).

Paratypes, NEW GUINEA: Wau; i <j>, 9.1.1966 (/. Sedlacek); i <j>, 6.X.I966
(G.Samuelsori) (BPBM).

COMMENTS. This species is very similar to the macropterae of fijiensis, although
larger and with the eyes prolonged ventrally. In the BPBM collection there is
a female from north-western New Guinea, Lake Wissel, with almost uniformly
brown antennae and no duplicated cilia on the fore wings, but otherwise very
similar to grayi. This may represent a further species. The micropterae of
fijiensis are more slender and very similar to loisthus, although a direct comparison
of loisthus and grayi would not suggest any close relationship between these
two species.

I 3 o L. A. MOUND

Carientothrips japonicus (Bagnall) comb. n.
(Text-fig. 62)

Cry ptothrips japonicus Bagnall, 1921 : 355-356. Holotype $, JAPAN (BMNH) [examined].
My strothrips japonicus (Bagnall) Mound, 1968 : 138.

The unique holotype of this species is rather damaged but is difficult to distinguish
from the eastern Australian species reedi Mound. The third antennal segment is
thicker at the base, suggesting the origin of the condition found in acti Mound.
The apices of the terminal setae on the tube are flattened in acti and pictilis Mound,
but are acute as in most other thrips in japonicus, reedi and vesper.

SPECIMEN STUDIED.
Holotype $, JAPAN: Kobe, 23.viii.i9i6 (Lewis) (BMNH).

Carientothrips loisthus Mound

Carientothrips loisthus Mound, 1974 : 29-30. Holotype $, AUSTRALIA: South Australia (ANIC)
[examined].

This species was based on a single female aptera collected in South Australia,
and at that time the holotype was regarded as a distinct species from specimens
collected in New Zealand. However there is more difference between the specimens
listed below from various localities within New Zealand than there is between some
of these specimens and the Australian holotype. For this reason they are all
regarded as one species. Large specimens from both North and South Island,
New Zealand have the femora brown, whereas the females from Waiwera have the
femora almost as yellow as the tibiae. Moreover these small specimens have the
head and thorax much paler than the abdomen. The large specimens have very
long pronotal setae with acute apices, whereas the small specimens have the epimeral
and midlateral setae bluntly rounded at the apex. The holotype is completely
apterous but has three ocelli, whereas the New Zealand specimens have a small
lobe (15 fim) attached to the lateral corners of the mesonotum, and the lateral
ocelli are scarcely indicated. The males are smaller than the females with almost
clear yellow legs and a large fore tarsal tooth.

MEASUREMENTS (large New Zealand $ in fj,m). Body length 2700. Head, length 310;
median width 225; ocellar setae 85; postocular setae 135; mid-dorsal setae 90-120. Pronotum,
length 175; median width 290; major setae -am 80, aa 100, ml 120, epim 130, pa 135. Tube
length 195.

SPECIMENS STUDIED.

Holotype $, AUSTRALIA: South Australia, Adelaide, Waterfall Gulley, grass by
stream, 14.1.1968 (L. A. Mound) (ANIC).

NEW ZEALAND: South Island, Greymouth, i $ on Juncus, 12. ix. 1972 (V. F.
Eastop); North Island, Levin, 2 <j>, ix. 1966, I <$, v. 1966, i $, iv. 1967 (Manson &
Pritchard); N.I., near Rotorua, 2 9, ix. 1972 (V. F. Eastop); N.I., Waiwera, 4 $,

NESOTHRIPS COMPLEX 131

i ( at base of grasses, viii. 1968 (L. A. Mound}; N.I., Helensville, i <$, viii. 1968
(L. A. Mound) (BMNH).

Carientothrips magnetis Mound

Carientothrips magnetis Mound, 1974 : 30-31. Holotype $, AUSTRALIA: Queensland (ANIC)
[examined] .

This species has the eyes well developed ventrally, but is intermediate between
loisthus and the acti-group of species in the form of the head and epimeral sutures.

SPECIMEN STUDIED.

Holotype $, AUSTRALIA: Queensland, 10 miles north of Townsville, on grasses,
20.vii.i968 (L. A. Mound) (ANIC).

Carientothrips miskoi Mound

Carientothrips miskoi Mound, 1974 : 3 1 - Holotype $, AUSTRALIA: New South Wales (ANIC)
[examined].

This species is known only from three females which were collected on two separate
occasions at the same site. It differs from mjobergi mainly in having the tube
brightly bicoloured.

SPECIMENS STUDIED.

Holotype $ aptera, AUSTRALIA: New South Wales, Cabbage Tree Creek, 7 miles
north of Nelligen, near Bateman's Bay, in leaf litter, 2.11.1969 (5. Misko) (ANIC).

Carientothrips mjobergi (Karny)

Cryptothrips mjobergi Karny, 1920 : 42. Holotype $, AUSTRALIA: Queensland (NR) [examined].

Cryptothrips (?) australicus Priesner, 19286 : 649-651. Holotype $, AUSTRALIA: Queensland
(HP) [examined]. [Synonymized by Mound, 1974 : 31.]

Cryptothrips incisus Girault, 1927 (37) : i. Syntypes 2 ty, AUSTRALIA: Queensland (QM)
[examined]. [Synonymized by Mound, 1974 : 31.]

Bolothrips (Bolothrips) flavitibia Moulton, 1968 : 117-118. Holotype $, AUSTRALIA: Queens-
land (CAS) [examined]. [Synonymized by Mound, 1974 : 31.]

This species is widespread in Australia and highly variable. The eyes are
frequently prolonged ventrally, although this is not true of all populations. The
tibiae are usually brown but sometimes yellow, and similarly the colour of the
fore wings and antennae apparently varies. The number of duplicated cilia on the
fore wing has been found to vary from two to eight even within a single population.
The pelta of the macropterae is very similar to that of macropterae of fijiensis
and grayi, but the distinction between a central body and lateral wings is less
evident in the pelta of micropterae and apterae. Numerous specimens of this
species, or species complex, have been studied from all the States of Australia
except Tasmania and the Northern Territories, mainly from Eucalyptus dead
leaves and branches.

132 L. A. MOUND

Carientothrips pedicillus Mound

Carientothrips pedicillus Mound, 1974 : 32-33. Holotype $, AUSTRALIA (ANIC) [examined].

This species is known only from the holotype. It is very similar to mjobergi
but has the antennal segments six and seven sharply constricted to a narrow pedicel.

SPECIMEN STUDIED.

Holotype $ aptera, AUSTRALIA: Australian Capital Territory, Mt Gingera, in
leaf litter, 13.^.1967 (E. B. Brittori) (ANIC).

Carientothrips pictilis Mound

Carientothrips pictilis Mound, 1974 : 33~34- Holotype $ AUSTRALIA : New South Wales (ANIC)
[examined] .

The unique holotype of this species is bicoloured as in some species of Bolothrips.
However, the fourth antennal segment has four sense cones and the maxillary
stylets are parallel in the middle of the head.

SPECIMEN STUDIED.

Holotype $, AUSTRALIA: New South Wales, Parkes, from Eucalyptus blakelyi,
7.viii.i959 (E. M. Reed] (ANIC).

Carientothrips reedi Mound

Carientothrips reedi Mound, 1974 : 34-35. Holotype $, AUSTRALIA (ANIC) [examined].

This species was described from apterae and one macroptera collected in New
South Wales and Australian Capital Territory, as well as one macroptera from
southern Queensland. This may be a widespread species because the unique
holotype of japonicus Bagnall only differs in the detailed structure of the antennae.
These two species are related to acti Mound in the form of the head and thorax.

SPECIMENS STUDIED.

Holotype $ aptera, AUSTRALIA: Australian Capital Territory, Canberra, Black
Mt, in grass tussock, n.xii.i96o (E. M. Reed] (ANIC). Paratypes from New South
Wales, A.C.T., and Queensland are listed in Mound (1974).

Carientothrips semirufus (Girault)

Elaphrothrips semirufus Girault, 1928(42) : 4. Holotype $, AUSTRALIA : Victoria (QM)

[examined].
Carientothrips semirufus (Girault) Mound, 1974 : 35-

This apterous species is very similar to the bicoloured specimens of loisthus
from New Zealand, but has short expanded major setae and the mesonotum is
closely transversely striate.

NESOTHRIPS COMPLEX 133

SPECIMENS STUDIED.

Holotype <J>, AUSTRALIA: Victoria, Melton, in tussocks, v. 1928 (F. E. Wilson)
(QM).

AUSTRALIA: New South Wales, 13 $, 10 <$ all collected south-east from the Blue
Mountains and detailed in Mound (1974) (BMNH, ANIC).

Carientothrips vesper Mound

Carientothrips vesper Mound, 1974 : 35-36. Holotype $, AUSTRALIA : Western Australia (ANIC)
[examined].

The unique holotype of vesper is very similar to the apterae of reedi but with
longer antennae and shorter expanded setae on tergite nine.

SPECIMEN STUDIED.

Holotype <j>, AUSTRALIA: Western Australia, Manjimup, in grass clump, 28.ix.i967
(L. A. Mound] (ANIC).

DICERATOTHRIPS Bagnall
Diceratothrips Bagnall, 1908 : 193. Type-species: D. bicornis Bagnall, by monotypy.

This genus has been confused with Dichaetothrips (Mound, 1968 : 78), but is
best restricted to a group of Neotropical species. Most of these species have a pair
of elongate anteocellar setae, although in some of the smaller species such as harti
Hood, delicatulus Hood, and setigenis Hood these setae are reduced. It is to this
group of species that Gastrothrips validipennis Hood is related.

Large species of Diceratothrips have a series of stout spines on the inner surface
on the fore femora, at least in the males. The antennae are frequently distinctive,
segment three is relatively long, the sense cones on segments three and four unusually
short, and segments seven and eight frequently form a single elongate unit. The
pronotum is also unusual with no thickening at the anterior margin and very little
medially even in large individuals, moreover the anterior third is reticulate and
the anterior setae short.

Diceratothrips validipennis (Hood) comb, n
(Text-figs 9, 66)

Gastrothrips validipennis Hood, 1938 : 403-406. Holotype $, U.S.A.: Florida (USNM) [not
examined] .

This is a small species of Diceratothrips with the anteocellar setae exceptionally
reduced.

SPECIMEN STUDIED.

U.S.A.: Florida, Homestead, I paratype $, on dead branches, 28.xii.i937 (J. C.

Bradley) (AMG).

134 L. A. MOUND

DICHAETOTHRIPS Hood
Dichaetothrips Hood, 1914 : 164-165. Type-species: D. brevicollis Hood, by monotypy.

The synonymy of this genus and its relationship to Scotothrips have been discussed
recently by Mound (1974) . The Indian species referred to below can be distinguished
from Nesothrips by the presence of a large fore tarsal tooth in the female as well
as the more elongate tube. It is returned to the genus Dichaetothrips on account
of the elongate postocellar setae.

Dichaetothrips gloveri (Ramakrishna & Margabandhu) comb. rev.

(Text-fig. 8)

Neosmerinthothrips gloveri Ramakrishna & Margabandhu, 1939 : 31-32. LECTOTYPE

INDIA (TNA), here designated [examined].

Dichaetothrips gloveri (Ramakrishna & Margabandhu) Shumsher, 1947 : 204-205.
Nesothrips gloveri (Ramakrishna & Margabandhu) Ananthakrishnan, 1960 : 32-33.

The original description of this species does not designate a holotype, although
Shumsher (1947) refers to a holotype in the Zoological Survey of India. The
specimen here designated as lectotype bears the original data and is the only
member of the type-series still available for study.

The postocellar setae of the lectotype are 55 jum long, rather longer than the
distance between the anterior and posterior ocelli. The fore tarsal tooth is slender
and almost as long as the width of the fore tarsus. Unfortunately the specimen
is crushed, but in the opinion of the present author further collecting is likely to
show that this individual is a small specimen of the species known as Dichaetothrips
usitatus Ananthakrishnan & Jagadish, 1970 (= Dichaetothrips indicus Ananthakrish-
nan, 1961).

SPECIMEN STUDIED.

Lectotype $, INDIA, Numkum, Ranchi, on Zizyphus jujuba, 1933 (TVR^o)

(TNA).

GASTROTHRIPS Hood gen. rev.

Gastrothrips Hood, 1912 : 156. Type-species: G. ruficauda Hood, by original designation.
Probolothrips Moulton, 1941 : 319. Type-species: P. hambletoni, by monotypy; treated here
as a synonym of abditus. [Synonymized by Hood, 1952 : 163.]

The genus Gastrothrips is used here for a group of nineteen species, mainly from
the Neotropics, which have only three sense cones on the fourth antennal segment.
The tube is variable in shape, either with straight sides or constricted apically,
and frequently much paler than the rest of the abdomen. In contrast the Neo-
tropical species related to nigrisetis Hood have four sense cones on the fourth
antennal segment (with the exception oipaulistarum), and the tube is black, frequently
with convex sides. These species are treated here under Neosmerinthothrips.

The Gastrothrips species which have a straight-sided tube comprise a single

NESOTHRIPS COMPLEX

135

FIGS 8-12. 8, Dichaetothrips gloveri. 9, Diceratothrips validipennis. 10-12.
Gastrothrips anolis : 10, large male, n, female. 12, pelta of female.

136 L. A. MOUND

species from India, falcatus Ananthakrishnan, and a complicated species-group
with named forms distributed throughout the tropics, the acuticornis-group. All
the other Gastrothrips species are known only from the Neotropics with a few species
extending into temperate latitudes. A major problem with species recognition
is the few species that are known from a series of specimens at more than one site.
Moreover several species are known from only one morph, aptera, microptera or
macroptera, and are therefore difficult to compare with each other. Cryptothrips
citriceps Priesner, 1921 may also belong in this genus but has not been studied.

GENERIC DEFINITION. Medium sized, usually dark species of Cryptothripini. Head usually
rectangular, longer than wide and not projecting in front of eyes; eyes large and usually equally
developed on dorsal and ventral surfaces; head rarely wider than long, rarely with cheeks
bearing stout setae. Antennae eight-segmented, VIII usually elongate and slender, rarely
short and broad ; segment IV with three sense cones, III with one or two sense cones. Pronotum
transverse, sometimes enlarged in major males, anterior setae usually small, epimeral sutures
frequently not complete. Fore tarsal tooth present in , present or absent in $. Wings
present or absent, duplicated cilia present or absent. Mesonotal anterior angles frequently
projecting in major males; metanotal median setae usually well developed. Pelta usually
well defined, triangular, with small lateral wings. Tube with straight sides evenly narrowing
to apex, or with apex sharply constricted ; never with sides strongly convex and not constricted
at apex; frequently yellow or paler than rest of abdomen. Males commonly oedymerous.

KEY TO THE SPECIES OF GASTROTHRIPS

1 Tube with straight sides, not sculptured nor sharply constricted close to terminal

setae (Text-fig. 16); tube dark at least in basal half. ..... 2

Apex of tube sharply constricted in front of terminal setae, tube often sculptured

(Text-figs 17-19) ; basal half of tube frequently yellow or paler than tergite IX . 3

2 Antennal segment VIII broad at base, scarcely o 5 times as long as VII (Text-fig. 64) ;

pelta with a small central body and slender lateral wings; fore wings without
duplicated cilia .......... falcatus (p. 143)

Antennal segment VIII slender, more than 0-7 times as long as VII; pelta without

slender lateral wings; fore wings with duplicated cilia acufico mis-group

(see acuticornis (p. 139), mongolicus (p. 145), procerus (p. 147), & subulatus (p. 148))

3 Basal half of tube yellow or reddish brown, paler than abdominal segment IX . 4

- Tube black or dark brown throughout, not paler than tergite IX ... 9

4 Antennal segments I-III largely yellow, paler than the head; tube bright yellow . 5

Antennal segments I-III brown, as dark as head; tube reddish yellow to reddish

brown ............. 7

5 Pelta small and semicircular, occupying less than o 5 of anterior margin of tergite

II (Text-fig. 12) ; oedymerous $ with pair of elongate tubercles on posterior margin

of pronotum .......... anolis (p. 142)

- Pelta broadly triangular, occupying at least two-thirds of anterior margin of tergite

II (Text-fig. 20) ............ 6

6 Antennal segments I and II yellow in sharp contrast to brown segments IV-VIII;

head of $ with cheeks evenly narrowing to base (Text-fig. 25) ; $ with a fore tarsal
tooth ........... intonsus (p. 145)

- Antennal segments grade evenly from yellow I to brown VIII, even VII has a pale

pedicel; head of $ with cheeks parallel except for slight constriction near base; $
without a fore tarsal tooth fulvicauda (p. 143)

7 Head slightly wider than long (2iOju,m 195 fim); mid and hind femora brown but

with a yellow area on distal posterior margin .... ruflcauda (p. 147)

NESOTHRIPS COMPLEX

14

137

FIGS 13-22. 13-19. Tube (last abdominal segment) : 13, Bolothrips italicus. 14,

Gastrothrips intonsus. 15, G. mandiocae. 16, G. falcatus. 17, G. callipus. 18, G.

abditus. 19, G. ruficauda. 20-22. Pelta: 20, G. intonsus. 21, G. mandiocae. 22, G.
ruficauda.

138 L. A. MOUND

- Head longer than wide ; mid and hind femora uniformly brown .... 8

8 Head less than 1-15 times as long as wide ; tube more than 0-9 times as long as head

corvus (p. 142)
Head more than 1-25 times as long as wide ; tube less than o 75 times as long as head

alticola (p. 140)

9 Legs and antennal segments I-VI largely yellow; tube 0-7 times as long as head;

apterous with metanotal setae fine and less than 15 ^tm long . fulviceps (p. 144)

- Legs and antennal segments largely dark ; tube o 8 times as long as head or longer ;

macropterous or micropterous with metanotal setae more than 30 pm long and

usually stout ............ 10

10 Fore wing with duplicated cilia . . . . \ . . n

- Fore wing without duplicated cilia, or micropterous . . . . . . 13

1 1 Female without a fore tarsal tooth ; postocellar setae less than the diameter of one

ocellus ........... abditus (p. 138)

- Female with a fore tarsal tooth ; postocellar setae usually at least 40 ^im long, usually

twice as long as the diameter of one ocellus . . . . . . . 12

12 Antennal segment III brown with yellow pedicel; sides of tube weakly convex,

curving from base to apex ; pronotal anteromarginal setae less than half as long as
anteroangulars ; median part of pelta narrowly triangular . . stygicus (p. 147)

Antennal segment III largely yellow, shaded brown at apex; sides of tube parallel
medially or even weakly constricted in anterior third; pronotal anteromarginal
setae as long as anteroangulars ; median part of pelta broadly triangular

mandiocae (p. 145)

13 Fore tarsal tooth of $ acute, almost as long as width of fore tarsus . tcxanus (p. 148)
Female without a fore tarsal tooth . ... . .... 14

14 Femora brown with yellow area distally on inner margin; sides of tube weakly

convex, curving from base to apex ...... callipus (p. 142)

Femora uniformly brown without a clear yellow area; tube parallel to weakly
constricted medially, or evenly tapering with sides not rounded ... 15

15 Micropterous; tube tapering from base to near apex . . . .monticola (p. 146)

- Macropterous; tube parallel or weakly constricted medially . fumipennis (p. 144)

Gastrothrips abditus Hood
(Text-fig. 28)

Gastrothrips abditus Hood, 1935 : 177-182. Holotype $, PANAMA: Barro Colorado (USNM)

[not examined].
Hoplothrips brasiliensis Moulton, 1938 : 378-379. Holotype $, BRAZIL (CAS) [not examined].

[Synonymized by Hood, 1952 : 163.]
Probolothrips hambletoni Moulton, 1941 : 320-321. Holotype $, BRAZIL (CAS) [not examined].

[Synonymized by Hood, 1952 : 163.]

Hood described this species from six female and one male macropterae collected
from burrows in dead branches. None of these have been studied by the present
author, but specimens from the type-series of Moulton's two species have been
seen. Apparently abditus is widespread in eastern South America, and it is closely
related to mandiocae and texanus. There are two undescribed species in the USNM
from Mexico which are also related to abditus. One of these is macropterous with
a heavy tube and dark brown third antennal segment. The other species includes
the only hemimacropterous Gastrothrips known to the author, and these may

NESOTHRIPS COMPLEX 139

represent abditus despite the fact that the wing lobes are shaded. The macropterae
of abditus have the fore wings pale or only weakly shaded.

SPECIMENS STUDIED.

BRAZIL: Minas Gerais, on dead Manioc stems, 3 $, i $, 2i.vii.i933, 2 <$, io.iv.1933
(Hambletori) (BMNH) [type data of brasiliensis and hambletoni], PARAGUAY:
Caaguazu, i <$ swept, 1.11.1964 (F. Andre). TRINIDAD: St Augustine, I $ in empty
Psychidae case, i.xi.i97o (L. A. Mound] (BMNH).

Gastrothrips acuticornis (Hood)

Cryptothrips acuticornis Hood, 1925 : 65. Holotype $, WEST INDIES: St Croix (USNM)

[examined] .
Cryptothrips cybele Girault, 1927(38) : i . Holotype <$, AUSTRALIA : Queensland (QM) [examined].

[Synonymized by Mound, 1974 : 53 ]
Gastrothrips noumeae Bianchi, 1945 : 251-254. Holotype $, NEW CALEDONIA (BPBM) [not

examined]. [Synonymized by Mound, 1974 : 53-1
Nesothrips acuticornis (Hood); Ananthakrishnan, 1969 : 181-182.

This species has a remarkable distribution and, moreover, it is difficult to distin-
guish from three other described species: mongolicus from Mongolia, procerus from
Brazil and subulatus from Transvaal. Hood described acuticornis from several
specimens collected on St Croix and Barbados, and in the description refers to
small pale males. Ananthakrishnan (1969) recorded both sexes from southern
India, and Mound (1974) compared Indian specimens with the West Indian holotype
and also specimens from New Caledonia and Australia. The Australian specimens
have the eighth antennal segment slightly shorter than the seventh in contrast
to the other material studied, and moreover the third antennal segment of most
of the Australian specimens is paler than that of either the holotype or the Indian
material. However, one female from Kuala Lumpur in Malaya has the third antennal
segment dark except for the pedicel as in the holotype, but the eighth segment Is
short as in the Australian specimens. For these reasons cybele and noumeae are
regarded as the same species as acuticornis, despite the fact that the males which
have been studied from Australia and New Caledonia are macropterous and brown
like the females, whereas the males from India and the West Indies are yellow
and micropterous. Large males from India have the posterior margin of the meta-
notum produced into a dentate fringe which is excavate medially.

The females of acuticornis are macropterous with short postocellar setae, and
the head of each of the specimens which have been studied is about i-i times as
long as the tube. However, there are two micropterous females from Basutoland
in the Albany Museum, Grahamstown which may represent this species but one
has both postocellar setae 30 /mi long and the other has these setae 12 /xm and
30 jitrn long. The single male collected with these specimens is also micropterous
and slightly oedymerous but has a simple posterior border to the metanotum unlike
the small yellow males from India.

Whilst considering the variation in acuticornis it is necessary to refer to three
other similar species. The South American species procerus has a relatively short

c

146 L. A. MOUND

tube, the head being 1*5 times as long as the tube in all three of the available speci-
mens. The holotype and one male from southern Brazil have a more slender
head than a female from Paraguay, but all three specimens have the third antenna!
segment light brown with a yellow pedicel, and none of them have elongate posto-
cellar setae. The South African species subulatus has the head about 1-4 times as
Iqng as the tube and the postocellar setae less than 15 /z,m long, but the third antenna!
segment is yellow and the fourth variable yellowish brown. Finally, mongolicus
has the head 1*2 times as long as the tube, the third antennal segment is yellow in
contrast to the dark brown fourth segment, but the postocellar setae are 50 /mi
long. These three species are known from too few specimens to assess the signifi-
cance of these differences at present. The form of the pelta differs between them
but this may be a function of body size. It is possible that all three are only local
.populations of acuticornis.

SPECIMENS STUDIED.

Holotype $ macroptera of acuticornis, WEST INDIES: St Croix, swept from grass,
9.iii.i_9i5 (C. B. Williams 553) (USNM 71992). Holotype <$ macroptera of cybele,
AUSTRALIA: Queensland, Gympie, in forest, 29^.1924 (A. A. Girault}.

AUSTRALIA: Queensland, Laidley, i <J, i <j> in forest, 31^.1923 (A. A. Girault};
Q., Grandchester, i (labelled as $} in forest, i.viii.i924 (^4. A. Girault} (QM);
Q., Moura, 2 $ on Rhodes grass, io.x.1967 (Page & Rigby); Q., 100 miles north of
Rockhampton, I ^ in grass, i8.vii.i968 (L. A. Mound) (BMNH).

NEW CALEDONIA: near Noumea, 2 $, i < paratypes of noumeae, beaten from grass,
24.ix.iQ40 (F. X. Williams) (BMNH, BPBM, HP).

WEST MALAYSIA: Kuala Lumpur, i $ from live tree, 26.xii.i969 (F. Andre]
(BMNH). INDIA: Madras, 4 <$ on Barteria sp. twigs, i8.vii.i969, 3 $ on dry Jasmine
twig, 25.vii.i966, i $ from grass, 23.^1.1968; Palghat, i $, i <$ on dry twig, 12^.1970;
Kumili, I $ on dry twig, 22.vii.i969; Ronningtonn, i $ on dry twig, 8.11.1969 (T. N.
A nanthakrishnan) (BMNH) .

Gastrothrips alticola Hood

Gastrothrips alticola Hood, 1942 : 570-573. Holotype $, PERU (USNM) [not examined].

This species was described from thirty-five apterous females and ten apterous
males. Hood states in the description 'basal three fourths of tube blackish brown',
whereas the paratypes which have been studied here have the basal three-fourths of
the tube paler than tergite nine. This has made comparison with other species
particularly difficult. In the key to species above alticola will run to monticola
if the tube is considered to be black. However, monticola is known only from
micropterae and alticola only from apterae. However, both forms were collected
in the Department of Huanuco, Peru, at 3000 metres. Moreover, in the BMNH
collections there is a single macropterous female which was collected within 100
miles of Huanuco. This female has the basal three fourths of the tube dark reddish
brown and paler than tergite nine, the fore wing bears six duplicated cilia, and the

NESOTHRIPS COMPLEX

141

fore tarsus does not bear a tooth. This specimen probably represents alticola.
and monticola may also be the same species.

SPECIMENS STUDIED.

PERU: Dept. Huanuco, Shishmay, 2 $ paratypes from shrub with dry branches,
v at 3000 m, i8.ix.ig37 (F. Woytkowski); Panao, i $, i < from dead branches and
leaves at 3000 m, io.ix.i937 (F. Woytkowski) (AMG).

23 vcK 24 25

27

FIGS 23-28. 23-26. Gastrothrips intonsus : 23, fore tarsus of small male. 24, small
male. 25, female. 26, fore tarsus of female. 27, G. ruficauda. 28, G. abditus.

142 L. A. MOUND

Gastrothrips anolis Morgan
(Text-figs 10-12)

Gastrothrips anolis Morgan, 1925 : 7-8. Holotype $, PUERTO Rico (USNM) [not examined].
Gastrothrips proteus Hood, 1933 : 417-419. Holotype $, PANAMA: Barro Colorado (USNM)
[not examined]. [Synonymized by Hood, 1935 : 186.]

Morgan described this species from one female, and Hood described proteus
from one macropterous and seventeen apterous females with seven apterous males.
The tube is bright yellow as in fulvicauda, intonsus and ruficauda, and the head of
the major males has spiny cheeks as in intonsus. However, the females and the
smallest males have a quadrate head similar to several other species of Gastrothrips.
The major males bear two elongate tubercles on the posterior margin of the pronotum
which overhang the metanotum, and Hood (1935 : 185) has published drawings
of the variation of this species.

SPECIMENS STUDIED.

PANAMA: Barro Colorado Is., Gatun Lake, paratypes of proteus; I $, i <$ on dead
branches, 27.vii.i933, I $ on dead grass and leaves, i.vii.1933, I <$ on grass,
I4.viii.i933 (/. D. Hood] (AMG). CAYMAN ISLANDS: 2 $, 2 < apterae in
Nightingale's nest, i6.vii.i962 (A. Ventura] (BMNH).

Gastrothrips callipus Hood

(Text-fig. 17)
Gastrothrips callipus Hood, 1935 : 182-186. Holotype $, U.S.A., Texas (USNM) [not examined].

Hood described this species from three female and one male macropterae from
Texas, but the species is apparently widespread in the south-eastern part of North
America. The femora are bicoloured as in the North American species ruficauda
and the Central American species anolis; however, the tube is black with weakly
convex sides. The male listed below from Mexico is identified as callipus with
some doubt because it is micropterous with the basal three antennal segments
yellowish brown.

SPECIMENS STUDIED.

U.S.A.: Texas, Victoria, paratype $ on Chenopodium, iv. 1908 (/. D. Mitchell)
(USNM); T., Sinton, i $ on dead branches, 20.xii.i967; T., San Antonio, i $ on
dead branches, 23.xii.i967 (F. Andre); Oregon, Ashland, I $ in berlese funnel,
6.xii.i937 (BMNH) ; South Carolina, Charleston, 3 $ on Solidago, 15^.1943 (W. S.
Fields); Florida, Hialiah, I $ on grass and weeds, 15.^.1969 (C. E. Stegmaier).
BAHAMA Is.: West Palm Beach, i $, 13^.1966 (W. E. Browning. MEXICO: in
quarantine at Nogales, i <$ on tomato fruit, 15.^.1941 (USNM).

Gastrothrips corvus Priesner
Gastrothrips (?) corvus Priesner, 1933 : 55-57- Holotype $, MEXICO (HP) [examined].

NESOTHRIPS COMPLEX 143

Gastrothrips capitalis Hood, 1935 : 174-177. Holotype $, U.S.A.: Texas (USNM) [examined].
Syn. n.

The two unique holotypes of corvus and capitalis have been compared with each
other and capitalis is here regarded as a small macroptera whereas the corvus holotype
is a large microptera. The other differences noted by Hood follow from this morph
difference, but it should be noted that in both specimens the basal two-thirds of
the tube is dark reddish brown in contrast to the black colour of segment nine.
The most closely related species appears to be monticola from Peru which has a
dark tube.

SPECIMENS STUDIED.

Holotype $ microptera of corvus, MEXICO: Stadt, im Zimmer, 20.ii.iQ24 (A . Dampf)
(HP). Holotype $ macroptera of capitalis, U.S.A.: Texas, Brownsville, Palm
Jungle, in beatings, 20.xi.i9ii (C. A. Hart] (USNM 71986).

Gastrothrips falcatus (Ananthakrishnan) comb. n.
(Text-figs 16, 64)

Nesothrips falcatus Ananthakrishnan, 1968 : 969-971. Syntypes n $, 5 <, INDIA (TNA)
[not examined].

The males of this species show extreme allometry. Major males bear several
very stout recurved setae on the posterior margin of the fore femora and fore coxae,
and the metanotum bears a median process projecting backwards over the pelta.
These features are not developed in minor males. Allometry is also known in
four other species of Gastrothrips, acuticornis, alticola, anolis and monticola, although
different parts of the body are affected in each species. In most species of Thysan-
optera which show allometry the major males are much larger than the minor
males, but in anolis and apparently falcatus the differences in body size are not very
great. Unfortunately the measurements published by Ananthakrishnan (1970;
1971) are given as ranges for each character and hence cannot be used to establish
correlations.

G. falcatus appears to be related to the acuticornis-group in the form of the head
and tube, but differs from other members of the genus in having antennal segment
eight short and broad. There is one specimen in the BMNH collections from Valparai
in southern India which is very similar to falcatus in colour and the form of the
wings and antennae, but has the tube constricted at the apex, the pelta broadly
triangular and the postocellar setae close together.

SPECIMENS STUDIED.

INDIA: from dead twigs; Madras, i <j>, ii. 1968, 3 $, xi. 1968, 2 ?, vii. 1969, i $,
xii. 1970; Devikula, I <J, ix. 1970; Goa, 2 $, x. 1968 (BMNH).

Gastrothrips fulvicauda Hood

Gastrothrips fulvicauda Hood, 1937 : 277-280. Holotype $, PERU (USNM) [not examined].
This species was based on three apterous females, and unlike ruficauda the head

I 4 4 L - A - MOUND

is rectangular without any ocelli (Hood, 1937: fig. 40). The tube is yellowish orange
and the distal half of the femora considerably paler than the tibiae. The most
closely related species seems to be alticola which is also based on apterae from Peru,
but these have the first two antennal segments brown and the tube more darkly
coloured.

SPECIMENS STUDIED.

PERU: Cajamarca Dept., near Celendin, from bush containing dry branches,
i paratype $ aptera, i.vi.i936 (F. Woytkowski) (USNM).

Gastrothrips fulviceps Hood

Gastrothrips fulviceps Hood, 1937 : 2 74~ 2 77' Holotype $, PERU (USNM) [not examined].

This species was described from seven apterous females from Celendin, Peru,
and the holotype was collected from a bush containing dry branches and moss.
No other member of Gastrothrips has a similar colour combination of dark tube
but yellowish legs, antennal segments and head. In colour and general shape
fulviceps is very similar to Nesothrips propinquus (Bagnall), but that species has
a more transverse head, four sense cones on the fourth antennal segment, a shorter
and broader eighth antennal segment, and a differently shaped pelta without a
strong subbasal line of sculpture. The head of fulviceps is as broad as long, inter-
mediate in shape between that of ruficauda and the majority of Gastrothrips species.

SPECIMENS STUDIED.

PERU: Cajamarca Dept., Celendin, i $ paratype on Alansoa acutifolia flowers,
2.vi.i936 (F. Woytkowski} (AMG). GUATEMALA: Guatemala City, i $ on Brunella
vulgaris, 7^.1945 (E. J. Hambleton] (USNM) . MEXICO : in quarantine at Brownsville,
Texas, I $ on rose, 6.x. 1939; in quarantine at San Francisco, California, i 9 on
Sedum, 20.iv.i938 (USNM).

Gastrothrips fumipennis Hood

Gastrothrips fumipennis Hood, 1952 : 163. Holotype $, BRAZIL (USNM) [not examined].

Hood described this species from three female and seven male macropterae.
According to the description large males bear a pair of lateral processes on the
mesothorax as in abditus, falcatus and acuticornis. Despite the absence of duplicated
wing cilia fumipennis is very closely related to abditus, although it is rather darker
with shaded wings and brown tarsi and third antennal segment.

SPECIMENS STUDIED.

BRAZIL: Santa Catarina, 2 < paratypes on dry branches, 16^.1949 (F. Plaumann]
(AMG).

NESOTHRIPS COMPLEX 145

Gastrothrips intonsus Hood

. \
(Text-figs 23-26)

Gastrothrips intonsus Hood, 19410, : 180-183. Holotype $ microptera, PERU (USNM) [Hot
examined].

The unique holotype of this species was related by Hood to picticornis on accotmt
of the shape of the head. However, picticornis is treated here in the genus Neo-
smerinthothrips because of the shape of the tube. Moreover the males of intonsus,
particularly large specimens, have the head with convex spiny cheeks as in anolis.
These major males, however, do not have the posterior margin of the pronotum
produced into tubercles. The females have the head broadest just behind the eyes
and strongly tapering to the base, and the pelta is broader than in the male and
similar to fulvicauda and alticola. The bright yellow tube is similar to anolis.

SPECIMENS STUDIED.

PERU: Piedras Grandes, at 3000 m, I major $ (det J. D. Hood) in flowers, 2.xi.iQ37
(F. Woytkowski) (USNM); Huacapistana, I $ on dry leaves, 12.^.1940 (F< Woytkow-
ski); San Felix, I minor from bush with dry leaves, 25.iii.i940 (F. Woytkowski)
(BMNH).

Gastrothrips mandiocae (Moulton)
(Text-fig. 21)

Dichaetothrips mandiocae Moulton, 1941 : 321-322. Holotype $, BRAZIL (CAS) [examined].
Gastrothrips oeceticola De Santis, 1943 : 92-96. Holotype $, ARGENTINA (Museos de la Plata)
[not examined]. [Synonymized by Hood, 1956 : 99.]

Hood (1950 : 22-25) redescribed and figured oeceticola from southern Brazil,
and later synonymized it with mandiocae after a comparison of the type-specimens.
The postocellar setae are frequently more than 100 /xm long, but the female listed
below from Paraguay has one of these setae less than 20 pm long. Because of this
variation the female listed below from Ghana is identified provisionally as mandiocae
despite the fact that the postocellar setae are short. The most closely related
species are abditus and texanus.

SPECIMENS STUDIED.

BRAZIL: Minas Gerais, I $, I < in dead Manioc stems, I5.ix.i933 (Hambleton)
[type data of mandiocae]; Vicosa, 3 $, 2 ^ on dead branches, S.vii & 20^.1964
(F. Andre] (BMNH). PARAGUAY: Caaguazu, i $ swept, 2.^.1964 (F. Andre]
(BMNH).

GHANA: Accra, Legon, I $ from Tapinanthus on Cacao, 1969 (P. Room) (BMNH).

Gastrothrips mongolicus (Pelikan) comb. n.

Nesothrips mongolicus Pelikan, 1965 : 231-233. Holotype $, MONGOLIA (TM) [not examined].
This species was described from two females and is discussed under acuticornis.

14$

L. A. MOUND

Contrary to the original description the ventral setae on the head are not particu-
larly long in comparison to other members of the acuticornis group.

SPECIMEN STUDIED.

MONGOLIA: Koschoo Zaidam, in Milvus nest at 1490 m, I paratype ?, 1.^.1964
(Kaszab) (J. Pelikan collection).

Gastrothrips monticola Hood
Gastrothrips monticola Hood, 1942 : 573-576. Holotype $, PERU (USNM) [not examined].

29

FIGS 29-30. Gastrothrips intonsus, large male : 29, fore tarsus. 30, head, pronotum and

mesonotum with short wings.

NESOTHRIPS COMPLEX 14?

This species was described from two female and one male micropterae, and its
possible synonymy with alticola is discussed under that species.

SPECIMEN STUDIED.

PERU: Huanuco Dept., Piedras Grandes, I $ paratype from a bush at 3000 m,
I3.xi.ig37 (F. Woytkowski) (AMG).

Gastrothrips procerus Hood

Gastrothrips procerus Hood, 1956 : 99-100. Holotype $, BRAZIL (USNM) [examined].

This species was described from one female macroptera, and is discussed under
acuticornis.

SPECIMENS STUDIED.

Holotype $, BRAZIL: Santa Catarina, on dead branches, v. 1949 (F. Plaumann)
(USNM 71989). BRAZIL: Parana, Ponta Grossa, I $ in grass tufts, I4.X.I972
(F. F. Eastop). PARAGUAY: Caaguazu, i $ sweeping, 2.11.1964 (F. Andre) (BMNH).

Gastrothrips ruficauda Hood
(Text-figs 19, 22, 65)

Gastrothrips ruficauda Hood, 1912 : 156-157. Syntypes 3 $, U.S.A.: Illinois (USNM) [not
examined].

Stannard (1957 : 104-106) records this species from eastern North America.
The males have a pointed process in front of the mesothoracic spiracle as in several
other species of Gastrothrips. The legs are bicoloured as in callipus, and the tube
is bright yellow as in anolis, fulvicauda and intonsus. This is the type-species of
Gastrothrips but the head of the female is relatively small with more rounded cheeks
than other members of the genus. The only specimen studied was an apterous
female with reduced wing-retaining setae but with well developed ocelli.

SPECIMEN STUDIED.

U.S.A.: Virginia, Rosslyn, i $ from dead willow branch, I.xi.i9i4 (C. B. Williams
&J. D. Hood] (BMNH).

Gastrothrips stygicus Hood

Gastrothrips stygicus Hood, 1935 : 186-191. Holotype $, PANAMA: Barro Colorado (USNM)
[not examined].

Hood described this species from five female and one male macropterae. The
tube is stouter than in abditus or mandiocae with weakly convex margins; however,
these three species, together with texanus, are closely related. The fore tarsal
tooth of stygicus is very short and broad, and the postocellar setae are intermediate
in length between those of abditus and mandiocae.

148 L. A. MOUND

SPECIMEN STUDIED.

PANAMA: Porto Bello, I $ paratype on dead vegetation, io.vii.ig33 (J- D. Hood)
(AMG).

Gastrothrips subulatus (Hartwig) comb. n.

Bolothrips subulatus Hartwig, 1948 : 113-115. Holotype $, SOUTH AFRICA : Transvaal (Pretoria)
[not examined].

This species, which was described from 45 and 5 <$ collected in Pretoria and one
female collected in Lourenco Marques, is discussed under acuticornis. The original
description refers to macropterous females and 'dealate' females and males. How-
ever, the paratypes listed below are micropterae with well developed ocelli and
wing retaining setae.

SPECIMENS STUDIED.

SOUTH AFRICA: Transvaal, Pretoria, paratypes; 4 , i <$ on Bignonia tweediana,
1.1.1946 (E. K. Hartwig) [same data as holotype]; I $ on B. tweediana, 16.^.1941;
i ? on bean leaves, 6.1.1940 (E. K. Hartwig} (BMNH, AMG, HP).

Gastrothrips texanus Hood

Gastrothrips texanus Hood, 1912 : 157159. Holotype $, U.S.A.: Texas (USNM) [examined].

This species is known only from the unique holotype macroptera and one damaged
female from the same locality. The pelta is similar to abditus, but the fore tarsus
bears a tooth and the fore wings do not have any duplicated cilia. Despite this
unusual combination of characters the species is closely related to abditus and
mandiocae.

SPECIMENS STUDIED.

Holotype $, U.S.A.: Texas, Brownsville, on Acacia farnesiana, 29^1.1908 (C. A.
Hart] (USNM 71990). I ?, 17.11.1938 (Anderson) (USNM).

NEOSMERINTHOTHRIPS Schmutz gen. rev.

Neosmerinthothrips Schmutz 1913 : 1051. Type-species: N. fructuum Schmutz, by monotypy.
Coenurothrips Bagnall, 192 la : 271. Type-species : C. brevicollis Bagnall, by original designation.
Galactothrips Moulton, 1933 : 404. Type-species: G. diversicolor Moulton, by monotypy.
Syn. n.

Stannard (1957) placed this genus in synonymy with Nesothrips on the grounds
that fructuum was closely related to those species of the Nesothrips complex 'formerly
called Gastrothrips'. Unfortunately, Stannard did not indicate in what sense he
was using the name Gastrothrips, and the present author has removed from Gastro-
thrips to Neosmerinthothrips those species which have four sense cones on the fourth
antennal segment together with paulistarum Hood. The genus Gastrothrips is
limited in this paper to a group of species of the Nesothrips complex with only three

NESOTHRIPS COMPLEX 149

sense cones on the fourth antennal segment. As a result Neosmerinthothrips includes
eighteen named species which are found in the Oriental, Ethiopian and Neotropical
regions. The Oriental and Ethiopian species are more or less closely related,
although it is interesting that the fore tarsal tooth is either present or absent.
One species from Brazil, diver sicolor, is probably introduced; it is so similar to the
West African hilaris. Similarly the West Indian and African species collaris
is very closely related to the Indian species, robustus. However, the annulipes
group of species, including picticornis, plaumanni, paulistarum and variipes, is almost
certainly native to Brazil. This is the group of species, together with two species
from Panama and one from Fiji, which are here removed from Gastrothrips. The
genus Neosmerinthothrips cannot be denned on any single character, but it includes
those members of the Nesothrips complex which have a heavy tube with somewhat
convex margins and the setae on tergite nine relatively elongate.

GENERIC DEFINITION. Medium-sized black or dark brown species of Cryptothripini. Head
variable, longer than wide, or slightly wider than long, stylets V-shaped and wide apart; post-
ocellar setae small (except nigrisetis), eyes not prolonged ventrally. Antennae eight-segmented ;
segment VIII either broad or narrowed at base, but never elongate and sharply constricted;
segment III with two sense cones, IV with four sense cones (three in paulistarum). Pronotum
transverse, epimeral sutures complete; oedymerous forms rare; fore tarsal tooth present in
but large, small or absent in $. Metanotal median setae usually small and fine; fore wing with
duplicated cilia. Pelta broad with lateral wings, lateral setae of tergites VI, VII and IX usually
elongate, tube heavy with convex margins or at least with small setal bearing tubercles laterally,
rarely almost straight with longitudal ridges in basal third.

KEY TO THE SPECIES OF NEOSMERINTHOTHRIPS
(excluding inquilinus, q.v.)

Antennal segment I yellow, if shaded with brown then distinctly paler than the head 2

Antennal segment I dark brown, as dark as the head ...... 8

Fore tarsal tooth absent in $ . . . . . . . . . . 3

Fore tarsal tooth present in $ . . . . . . . . . . 6

Head at least i 2 times as long as maximum width (Text-fig. 33) ; head largely yellow 4
Head less than o 95 times as long as maximum width (Text-fig. 32) ; head dark brown 5

Pronotal anteroangular and midlateral setae 15 /*m long [West Africa] . hilaris (p. 153)
Pronotal anteroangular and midlateral setae more than 35/xm long [Brazil]

diversicolor (p. 152)

Antennal segments I-III largely yellow; tube more than i -8 times as long as maxi-
mum width [India and Ceylon] ....... fructuutn (p. 152)

Antennal segment II brown in basal half, III brown with a yellow pedicel; tube

about i -5 times as long as wide [Brazil] .... picticornis (p. 155)

Antennal segments IV and V yellow, not darker than III and IV [South Africa]

hoodi (p. 153)

Antennal segments IV and V dark brown in contrast to yellow-brown of segment III 7

Fore tarsal tooth of $ less than half as long as width of foretarsus ; tube more than
i -7 times as long as maximum width; large species, antennal segment III about
80 jum long [Seychelles] ....... brevicollis (p. 151)

Fore tarsal tooth of $ about as long as width of fore tarsus ; tube less than i 6 times
as long as maximum width (Text-fig. 36) ; small species, antennal segment III less
than 60 nm long [Java] ........ xylebori (p. 155)

150 L. A. MOUND

8 Fore tarsal tooth absent in $ . . . . . . . . . . 9

- Fore tarsal tooth present in $ but sometimes very small . . . . . 1 1

9 Antennal segment III brown with pedicel yellow .... affinis (p. 150)

- Antennal segment III yellow, shaded light brown in distal half . . . . 10

10 Fore wing shaded only close to bases of subbasal setae ; fore femora largely yellowish

brown, darkly shaded along external margin near base [India] . . robustus (p. 155)

Fore wing shaded completely across base ; fore femora largely dark brown, paler near

apex [Africa and West Indies] ....... collaris (p. 151)

11 Fore wings uniformly shaded light or dark brown . . . . . . 12

- Fore wings not shaded, or shaded at base with median area much paler . . 13

12 Fore wing dark brown; major setae in tergite IX exceptionally dark and stout with

pale slender apices, B v longer than tube; pronotal anteromarginal and antero-
angular setae scarcely longer than posterior discal setae [Brazil] plaumanni (p. 155)

- Fore wing light brown; major setae on tergite IX not exceptionally dark and stout,

B l about half as long as tube; pronotal anteromarginal and anteroangular setae
twice as long as posterior discal setae [Fiji] ..... fijiensis (p. 152)

13 Antennal segment IV with three sense cones [Brazil] . . paulistarum (p. 154)

- Antennal segment IV with four sense cones . . . . . . . 14

14 Fore tarsal tooth well developed, length more than 0-5 of fore tarsal width; head

broadest across posterior part of eyes [Brazil] . . . . . . 15

Fore tarsal tooth small, length less than 0-25 of tarsal width; head broadest across

cheeks [Panama] ........... 16

15 Second antennal segment brown at base, yellow at apex; femora brown, pale at

extreme apices; tube about 2-1 times as long as maximum width annulipes (p. 150)

- Second antennal segment largely yellow ; femora yellow in distal third ; tube less than

2-0 times as long as wide (Text-fig. 37) ...... variipes (p. 155)

1 6 Postocellar setae about 2 . o times as long as diameter of one ocellus ; apices of femora

yellow; major setae on abdomen black ..... .nigrisetis (p. 154)

- Postocellar setae about 0.5 times as long as diameter of one ocellus; femora dark

brown; major setae on abdomen pale ..... parvidens (p. 154)

Neos merint hot hrips affinis (Bagnall) comb. n.

Coenurothrips affinis Bagnall, 19216 : 361-362. Holotype $, CEYLON (BMNH) [examined].

This species is known from a single crushed specimen which is similar to collaris
and robustus in structure but with the third antennal segment largely dark brown.
The fore femur is dark brown along the external margin and yellow along the internal
margin. Unlike brevicollis the fore tarsus has no tooth.

SPECIMEN STUDIED.

Holotype $, CEYLON: among cotton from Hettipold, Ex. Gd., 2O.vi.i9i3
(A. Rutherford) (BMNH).

Neosmerinthothrips annulipes (Hood) comb. n.

Gastrothrips annulipes Hood, 1950 : 13-16. Holotype $, BRAZIL (USNM) [not examined].
Nesothrips (Gastrothrips) milleforme De Santis, 19630 : 12-14. Holotype $, ARGENTINA (Museos
de la Plata) [examined]. Syn. n.

The head of this species is similar to that of the Brazilian species variipes and
picticornis, but the tube is more slender with nearly straight sides and slightly

NESOTHRIPS COMPLEX 151

constricted near the apex. The other species described from Brazil, plaumanni,
has the head wider across the cheeks than across the eyes but is otherwise similar,
and these four species are all closely related. The holotype of milleforme has the
distal parts of the major abdominal setae pale not dark, but cannot otherwise be
distinguished satisfactorily from the paratype of annulipes listed below.

SPECIMENS STUDIED.

Holotype $ of milleforme, ARGENTINA: Tezanosa Pinto (Prov. Entre Rios), 1957
(Rosillo) (Museos de la Plata).

BRAZIL: Rio de Janeiro, Jacarepagua, I paratype of annulipes, on dead branches,
2i.vi.i948 (J. D. Hood] (USNM).

Neosmerinthothrips brevicollis (Bagnall) comb, nov., stat. rev.

Coenurothrips brevicollis Bagnall, 19210 : 271-272. Syntypes g, $, SEYCHELLES (BMNH)
[examined].

This species was synonymized in error with Coenurothrips validus Bagnall in
Mound (1968 : 142), and placed in the genus Nesothrips sensu lato. However,
validus, which is known from a single badly damaged specimen, has a pair of stout
interocellar setae and is here placed in the genus Nesidiothrips. N. brevicollis
is very similar to collaris and robustus although rather smaller. However, it differs
not only in having paler basal antennal segments but in having a small slender fore
tarsal tooth in the female. In contrast to hoodi these three species all have yellowish
brown setae in tergite nine.

SPECIMENS STUDIED.

Syntypes. SEYCHELLES: Mahe, i $, Cascade Estate, top of Mt Serbert, x. 1908-
i. 1909 (H. Scott 104); i $, 1908-09 (3. No. 71); 2 $, 1908-09 (15 No. 71) (BMNH).

Neosmerinthothrips collaris (Bagnall) comb. n.

Cryptothrips collaris Bagnall, 1917 : 26-27. Lectotype <j>, ST. VINCENT (BMNH) [examined].
Gastrothrips fuscicauda Morgan, 1925 : 6-7. Holotype <$, PUERTO Rico (USNM) [examined].

Syn. n.
Bolothrips marshalli Priesner, 1934 : 58-60. LECTOTYPE <j>, SIERRA LEONE (BMNH),

here designated [examined]. Syn. n.
Gastrothrips dominicanus Hood, 1935 : 170174. Holotype <, DOMINICA (USNM) [not

examined]. Syn. n.

Bagnall described collaris from two females and Hood described dominicanus
from two males but there appear to be no significant differences between the speci-
mens. The damaged unique holotype of fuscicauda has been compared with the
paratype of dominicanus as well as the other material listed below. More surpris-
ingly the African specimens described as marshalli apparently represent the same
species together with the specimen from Mozambique, and moreover Indian speci-
mens referred to as robustus Ananthakrishnan can only be distinguished on very

152 L. A. MOUND

trivial characters. Thus collaris appears to be a tramp species which has been
distributed by man, possibly during the slave trade.

SPECIMENS STUDIED.

Lectotype $, with I $ paralectotype of collaris, ST VINCENT: ?i8g5 (H. H. Smith)
(BMNH). Holotype <$ of fuscicauda, PUERTO Rico: from stomach of lizard (G. N.
Wolcott) (USNM type No. 71987) (labelled ?$). Lectotype $ of marshalli, SIERRA
LEONE: Njala, inside Cassava seedpod, 4.xi.ig32 (E. Hargreaves) , with I <$ para-
lectotype bearing identical data (BMNH).

DOMINICA: paratype $ of dominicanus, swept from grass, 13.111.1915 (C. B. Williams
571) (AMG). MOZAMBIQUE: Lourenco Marques, 2 $>, 4 <$ in dry Cassia pods,
vii. 1936 (/. C. Faure) (AMG).

Neosmerinthothrips diversicolor (Moulton) comb. n.
Galactothrips diversicolor Moulton, 1933 : 404-406. Holotype $, BRAZIL (CAS) [not examined].

This is probably only a local and introduced variety of hilaris Priesner from
West Africa. The only apparent differences are the slightly longer setae on the
pronotum.

SPECIMEN STUDIED.
BRAZIL: Bahia, on Galactia sp., i paratype - (Bondar) (BMNH1.

Neosmerinthothrips fijiensis (Moulton) comb. n.

Gastrothrips fijiensis Moulton, 1944 : 286-287. Holotype $, FIJI (BPBM) [examined].

The single specimen from which this species is known has the head rather similar
to the annulipes group from Brazil and the tube is also similar to annulipes. How-
ever, the abdominal setae are rather short, the dorsal setae on tergite nine being
145 jam and the tube 260 /mi. At present fijiensis cannot be placed satisfactorily
in any species group. The median part of the pelta is triangular although the
lateral wings are well developed.

SPECIMEN STUDIED.

Holotype $>, FIJI; Navai Mill, Viti Levu at 2500 ft, i6.ix.i938 (E. C. Zimmerman)
(BPBM).

Neosmerinthothrips fructuum Schmutz
(Text-figs 32, 34, 35, 70)

Neosmerinthothrips fructuum Schmutz, 1913 : 1052-1053. LECTOTYPE $, CEYLON (HP),

here designated [examined].
Oedemothrips ceylonicus Karny, 1925 : 137-139. Holotype $, CEYLON (BMNH) [examined].

Syn. n.

NESOTHRIPS COMPLEX 153

The specimen here designated as lectotype was labelled by Prof. H. Priesner
'Typen material, ex coll. Karny', although the original description does not indicate
if more than one specimen existed. This specimen is very faded and partly crushed,
but otherwise cannot be distinguished from specimens labelled as 'formosensis
var. Kami' by Prof. Ananthakrishnan collected in southern India. The female
of the species lacks a fore tarsal tooth. The true formosensis is treated here under
Nesothrips, and has a tube with straight not weakly convex margins.

SPECIMENS STUDIED.

Lectotype $ offructuum, CEYLON: Peradenyia, in Schoten (Uzel 55) (HP). Holo-
type $ of ceylonicus, CEYLON; Peradenyia, in empty galleries in Cassia multijuga
with 3 immatures, 1^.1924 (F. T. Jepson) (BMNH).

INDIA: Kerala, Kudal, I $ in dry twigs, 6.X.I969 (TNA 158); Vyithri, 3 $,
I5.viii.i969; Ranni, 2 $, 5.x.ig6g-, Kiruatti, i , ig.ix.ig6g (BMNH); Sagar, i$
20.x. 1966 (HP).

Neosmerinthothrips hilaris (Priesner) comb. n.

(Text-fig. 33)
Bolothrips hilaris Priesner, 1937 : 624-626. Holotype <$, SIERRA LEONE (BMNH) [examined].

The tube of this species is heavy with slightly convex sides bearing distinct
setal bases, as in other species of Neosmerinthothrips. There are four sense cones
on the fourth antennal segment, and segment eight is broad at the base. The eyes
are smaller on the ventral than on the dorsal surface, the head does not project
in front of the eyes and the ocellar setae are minute. The epimeral sutures are
sometimes not quite complete, and the setae on tergite nine are elongate. Despite
the colour difference hilaris is related to collaris and fructuum in structure, and
the Brazilian species diversicolor is probably only a local variant.

SPECIMENS STUDIED.

Holotype <, with allotype $, SIERRA LEONE: Njala, in twigs of Bauhinia tomen-
tosa, 1936 (E. Hargreaves); I - paratype at same locality on dead branch of Ficus
(BMNH).

GHANA: near Legon, II $, 6 < from Tapinanthus banguensis on Cacao branches,
1969 and 1970 (P. Room) (BMNH).

Neosmerinthothrips hoodi (Faure) comb. n.

Gastrothrips hoodi Faure, iQ54a : 9-13. Holotype $ macroptera dealate, SOUTH AFRICA
(Pretoria) [not examined].

This species is very similar to fructuum in the shape of the head and tube, and
differs mainly in colour and the presence of a pretarsal tooth. Although it was
described in Gastrothrips, Faure specifically compared it to fructuum (as ceylonicus
Karny) and moreover listed those characters which are here regarded as excluding
species from Gastrothrips sensu stricto.

154 L - A - MOUND

SPECIMENS STUDIED.

SOUTH AFRICA: Zululand, St Richards Bay, I $ mac. paratype on dead branches,
17.11.1946; Kluhluwe, 4 $ dealate paratypes on dead branches of Syzygium cor datum,
I3.V.IQ49 (J. C. Faure) (AMG and BMNH).

Neosmerinthothrips inquilinus Ananthakrishnan

Neosmerinthothrips inquilinus Ananthakrishnan, 1960 : 32-33. Holotype $, INDIA (?TNA)
[not examined].

The holotype female and two male paratypes of inquilinus cannot be found
at present in Prof. Ananthakrishnan 's collection (teste T.N.A. in litt.). However,
judging from the description this is likely to prove to be a senior synonym of robustus
q.v.

Neosmerinthothrips nigrisetis (Hood) comb. n.
Gastrothrips nigrisetis Hood, 1935 : 161-165. Holotype $, PANAMA (USNM) [not examined].

The ocellar setae of nigrisetis are very well developed, about 40 /mi long and
arising well behind a line joining the posterior ocelli. As a result the head and
pronotum look very like a species of Dichaetothrips , but the tube is unusually
heavy with small setal tubercles laterally and the abdominal setae are elongate.
This species and parvidens are closely related and are probably derived from the
annulipes group. They are only distantly related to the other species of Neo-
smerinthothrips, although Hood compared them to collaris (as dominicanus] in
the original description.

SPECIMENS STUDIED.

PANAMA: Barro Colorado Island, on dead branches, 3 $, I <$ paratypes, 30.vii.i933
(/. D. Hood) (AMG).

Neosmerinthothrips parvidens (Hood) comb. n.

Gastrothrips parvidens Hood, 1935 : 165-168. Holotype $, PANAMA (USNM) [not examined].

This species is closely related to nigrisetis but has very small ocellar setae, and
the tube is more slender than any other species of Neosmerinthothrips.

SPECIMENS STUDIED.

PANAMA: Barro Colorado Island, on dead branches, I paratype $, io.viii.i933
(J. D. Hood) (USNM); Canal Zone, Barro Colorado Is., I $ on Swartzia danienensis
fls, io.v.i939 (Zetek) (USNM).

Neosmerinthothrips paulistarum (Hood) comb. n.

Gastrothrips paulistarum Hood, 1950 : 25-27. Holotype $, BRAZIL (USNM) [examined],
Because of the antenna! sense cone formula this species will come out to Gastro-

NESOTHRIPS COMPLEX 155

thrips in the key above. However, it is based on a single specimen which may
be aberrant, and it is closely related to the annulipes group of species. The tube
is dark and heavy with slightly convex margins, and the head shape is intermediate
between that of annulipes and parvidens.

SPECIMEN STUDIED.

Holotype <j>, BRAZIL: Estado de Sao Paulo, Sao Carlos, from dead branches,
I3.vi.i948 (J. D. Hood 6- D. P. de Souza Dias) (USNM).

Neosmerinthothrips picticornis (Hood) comb. n.

Gastrothrips picticornis Hood, 1936 : 272-275. Holotype $, BRAZIL (USNM) [not examined].

The tube of this species is very broad, rather similar to that of xylebori from
Java. However, picticornis is probably closely related to plaumanni and variipes
which also come from Brazil, but which have a fore tarsal tooth in the female.

SPECIMENS STUDIED.

BRAZIL: Rio de Janeiro, I $ paratype on dead twigs of Annonasquamosa, io.viii.i934
(D. Mendes) [holotype in USNM bears identical data] ; State of Sao Paulo, Itanhaen,
i $ on dead branches, 17^.1948 (J. D. Hood & J. Lane) (AMG).

Neosmerinthothrips plaumanni (Hood) comb. n.

Gastrothrips plaumanni Hood, 1950 : 20-22. Holotype $, BRAZIL (USNM) [examined],

This is the only South American member of this genus with dark brown fore
wings. The abdominal setae are long and dark as in picticornis and variipes, and
the tube is moderately heavy.

SPECIMENS STUDIED.

Holotype $, BRAZIL: Sta Catarina, on dry branches, 5.1.1949 (F. Plaumann)
(USNM Type No. 71988).
BRAZIL: Vicosa, 2 $ on dead braches, 20.xi.i964 (F. Andre) (BMNH).

Neosmerinthothrips robustus (Ananthakrishnan) comb. n.

Nesothrips robustus Ananthakrishnan, 1964 : 102-103. Syntypes $, $, INDIA (TNA) [not
examined) .

Contrary to the original description the female of this species does not have a
fore tarsal tooth (Ananthakrishnan, 1968 : 972). It is placed in the genus Neo-
smerinthothrips on account of the heavy tube with weakly convex margins bearing
small setal bases, and the elongate setae on tergite nine. It is very similar to the
widespread species collar is, being rather smaller and paler, and may eventually
prove to be merely a local variety.

D

156 L. A. MOUND

SPECIMENS STUDIED.

INDIA: Madras, 2 ^, 2 ?; Goa, i $; Adar, i <$, i <j>; Thenmatai, i <j>, all determined
by T. N. Ananthakrishnan (BMNH).

Neosmerinthothrips variipes (Hood) comb. n.

(Text-figs 37, 73)
Gastrothrips variipes Hood, 1950 : 16-20. Holotype $, BRAZIL (USNM) [not examined].

This species is very close to plaumanni but the fact that it has rather paler wings
and more yellowish basal antennal segments and femora may not be significant
when more specimens are collected.

SPECIMENS STUDIED.

BRAZIL: Rio de Janeiro, Jacarepagua, i $ paratype on dead branches, 22^.1948
(/. D. Hood 6- T. Borgmeir); State of Sao Paulo, Sao Carlos, i < paratype, 15^.1948
(J. D. Hood & D. P. de Souza Diaz) (AMG).

Neosmerinthothrips xylebori Priesner
(Text-fig. 36)

Neosmerinthothrips xylebori Priesner, 1935 : 370. LECTOTYPE $, JAVA (SMF), here designa-
ted [examined].

5 macroptera. Colour brown, tube blackish; mid and hind femora dark at base and along
anterior margin but yellow at apex and posterior margin; fore femora yellow in apical half;
antennal segments I and II yellow, III light brown, succeeding segments progressively darker;
wings shaded; major setae light brown.

MEASUREMENTS (Lectotype $ in pm). Body length 1550. Head, length 175; maximum
width 190. Fore wing, length 700; subbasal setae 50, 50, 58; number of duplicated cilia 6-8.
Tergite IX setae, fi x 145, B 2 130, f? 8 125. Tube, length 190; basal width 115; apical width
35. Antennal segments length 26; 50; 55; 50; 50; 50; 38; 32.

SPECIMENS STUDIED.

Lectotype $>, JAVA: Tegallega near Tjibadak, 500-600 m, in tunnel of Xyleboms
coffeae (on Cacao?), 30.^.1925 (Menzel), with I <$ paralectotype (SMF); 2 $ para-
lectotypes with identical data (HP).

COMMENTS. This species was described in a key, complete with collection data
but without reference to the number of individuals. The lectotype selected here
is the female labelled as holotype by Prof. Priesner which is mounted on a slide
with one larva. Despite the long slender fore tarsal tooth of the female it is similar
tofructuum, although the only available male is hemimacropterous and oedymerous.

NESIDIO THRIPS gen. n.

Type-species: Nesothrips alius Ananthakrishnan.

Medium-sized dark brown species of Cryptothripini with males frequently oedymerous.

NESOTHRIPS COMPLEX 157

Head longer than wide or about as wide as long; ocellar setae elongate, arising within the
ocellar triangle; postocular setae elongate, major males with two pairs of postocular setae;
maxillary stylets wide apart, V-shaped and retracted to postocular setae. Antennae eight-
segmented, VIII fairly broad at base; two sense cones on segment III, four sense cones on IV.
Pronotum with median thickening, anterior margin thickened in males ; fore femora sometimes
enlarged, fore tarsal tooth well developed in both sexes. Mesonotal lateral setae small; meta-
notal median setae frequently stout; fore wing with duplicated cilia; mid and hind femora
each with one stout dorsal seta. Pelta with small lateral wings; sigmoid wing retaining setae
well developed on tergites III VI, nearly straight on II and VII; lateral setae on tergites VI
and VII elongate, on IX shorter than tube; tube heavy, ridged basally and slightly constricted
near apex.

This genus is erected for two species, one from India and the other from the
Seychelles and the Solomon Islands. These species are closely related to Nesothrips
and Rhaebothrips species, and the mid and hind femora bear a single stout dorsal
seta as in those genera. However, the ocellar setae are elongate and arise within
the ocellar triangle, and the fore tarsus bears a large tooth in the female unlike
both Nesothrips and Rhaebothrips.

KEY TO THE SPECIES OF NESIDIOTHRIPS

I Setae on tergite IX yellowish brown; midlateral pronotal setae more than 80 ^tm

long in $; head slightly longer than tube [India] .... cilius (p. 157)
- Setae on tergite IX dark brown ; midlateral pronotal setae less than 60 pm long in $ ;

head slightly shorter than tube [Seychelles] ..... validus (p. 157)

Nesidiothrips alius (Ananthakrishnan) comb. n.

(Text-figs 46, 48, 50, 69)
Nesothrips alius Ananthakrishnan, 1970 : 52-55. Holotype $, INDIA (TNA) [not examined].

The original description of this species includes illustrations of the dimorphism.
The oedymerous males have a second pair of long postocular setae medially on the
vertex which are not developed on females or minor males. The median metanotal
setae and the pronotal anteroangulars and midlaterals are also elongate in large
males. This species is closely related to validus and further collecting may indicate
that only one widespread species is involved.

SPECIMENS STUDIED.

INDIA: Aryankavu, I $, I $ on dry twiner, ^.x.^g6g (T. N. Ananthakrishnan)
(TNA) [these data are similar to those published for the type-series - Kerala,
Aryankavu, dry twigs, 8.x. 1969].

Nesidiothrips validus (Bagnall) comb. n.
Coenurothrips validus Bagnall, 19210 : 272-273. Holotype $, SEYCHELLES (BMNH) [examined].

The holotype of this species is very badly crushed and the interocellar setae
are not readily visible. The species C. brevicollis Bagnall is not a synonym of
validus contrary to Mound (1968 : 142). The specimens listed below from the Solomon

I 5 8 L. A. MOUND

Islands are identified provisionally as validus, the males show a similar dimorphism
to alius but the terminal abdominal setae are dark brown.

SPECIMENS STUDIED.

Holotype $, SEYCHELLES: Silhouette [Mare aux Cochons, ix. 1908] (BMNH).

SOLOMON ISLANDS: Guadalcanal, Mt Austen, 2 <$, I $ in litter, I9.xi.i963
(P. Greenslade) (BMNH).

NESOTHRIPS Kirkaldy

Nesothrips Kirkaldy, 1907 : 103. Type-species: N. oahuensis Kirkaldy, by monotypy.
Oedemothrips Bagnall, 1910 : 680. Type-species: O. laticeps Bagnall, by monotypy. [Synony-
mized by Bianchi, 1944.]

31

FIG. 31. Nesothrips oahuensis, large male.

NESOTHRIPS COMPLEX 159

Oedemothrips is a synonym of Nesothrips because the names laticeps and oahuensis
have been shown to refer to the same species. However, the other genera which
were placed as synonyms of Nesothrips by Stannard (1957) are here withdrawn
from synonymy for the reasons given in the introduction and key. Unfortunately
this still leaves Nesothrips as an unsatisfactory group of fourteen more or less
related species from the Pacific, Australia and Indonesia, with two species which have
become widespread on trade routes. This group of species is closely related to
Rhaebothrips through aoristus and oahuensis, and it is also closely related to Cariento-
thrips through melinus and propinquus. N eosmerinthothrips fructuum is sometimes
misidentified as Nesothrips brevicollis (or one of its synonyms), but the species
of N eosmerinthothrips have a stout heavy tube with weakly convex instead of
straight margins. Similarly, small species of Scotothrips and Dichaetothrips have
been placed in Nesothrips despite the fact that they have a more elongate tube.

In the present interpretation Nesothrips includes both short broad species such
as oahuensis and propinquus, as well as more elongate species such as niger and
yanchepi. The eyes are prolonged on the ventral surface of the head in some
species and in propinquus the extent of this prolongation is variable. Duplicated
cilia are present on the fore wings of those species in which macropterae are known
with the exception of propinquus which does not have duplicated cilia. The syste-
matics of this group is not likely to be stabilized without further collecting in the
New Guinea and Pacific areas.

GENERIC DEFINITION. Small to medium sized dark species of Cryptothripini. Head fre-
quently wider than long but sometimes longer than wide, usually only weakly extended in
front of eyes; eyes variably prolonged on ventral surface; maxillary stylets wide apart and
V-shaped. Antennae eight-segmented, two sense cones on segment III, four sense cones on
IV; segment VIII short and broad. Pronotum transverse, sometimes enlarged in major
males; epimeral sutures complete. Fore tarsal tooth present in <$, absent in $. Wings, when
present, usually with duplicated cilia. Metanotal median setae usually small. Pelta with
lateral lobes in most species. Tube relatively short, with straight sides.

KEY TO THE SPECIES OF NESOTHRIPS

Antennal segment I yellow or brownish yellow, much paler than VII and VIII and

usually paler than the head ......... 2

Antennal segment I brown, as dark as the head and distal antennal segments . 8

Setae B! on tergite IX more than o 8 times as long as the tube ; major setae unusually
slender; dorsal surface of head with two pairs of elongate postocular setae ; mid and
hind legs dark brown except for the pale hind margin of femora at extreme apex
[Rapa Is.] semiflavus (p. 168)

Setae B t on tergite IX less than 0-7 times as long as tube; major setae stout and
dark; head with only one pair of postocular setae; mid and hind legs sometimes
yellow, or with apex of femora more or less yellow ..... 3

Head more than i-i times as long as wide (Text-fig. 38) [Guam] . artocarpi (p. 161)

Head less than i o times as long as wide, usually distinctly wider than long (Text-
figs 40-42) ... 4

Posterior margin of hind femora yellow, anterior margin dark brown, fore femora
yellow in distal half ; eyes not prolonged on ventral surface of head .
Micropterae with anterior margin of head and antennal segments I-V

160 L. A. MOUND

yellow or brownish yellow ; macropterae and hemimacropterae with head and most
of antennal segments IV-V brown [Mascarenes, Indonesia, Japan, Hawaii] (see
also rhizophorae) ......... brevicollis (p. 162)

Femora coloured differently from above, or with eyes prolonged on ventral surface of

head ............. 5

5 Tube as long as or a little longer than the head ; hind femora dark brown but yellow

in distal third; antennal segments I and II yellow, III yellow-brown, IV scarcely
paler than V-VIII which are dark brown [Malaya, Sumatra] . tnalaccae (p. 164)

- Tube 0-8 times as long as head or shorter; hind femora largely yellow ... 6

6 Lateral lobes of pelta separate from the median lobe or with only a slender connec-

tion, posterior border of pelta eroded medially and separate from anterior border

of tergite II [Australia, New Zealand, South Africa] . . propinquus (p. 167)

Lateral lobes of pelta broadly joined to median lobe, posterior border close to anterior

border of tergite II (Text-fig. 45). . . . . . . . . 7

7 Body colour dark brown; postocellar setae finely pointed [Fiji] . . fodinae (p. 163)

Body colour largely yellow; ocellar setae blunt at apex [Australia] . melinus (p. 166)

8 Ventral length of eyes at least i 5 times as long as dorsal length [Pacific Is.] . 9

Ventral length of eyes less than i -2 times dorsal length [Australia] . . . 10

9 Ocellar setae 60-80^ m long, 40-60 jam apart at base (Text-fig. 31); setae B^ on

tergite IX go-ioo^im [Hawaii] ...... oahuensis (p. 167)

- Ocellar setae 35 /un long, Sojum apart at base (Text-fig. 39) ; setae B t on tergite IX

200 jum [Marquesas Is.] ........ niger (p. 166)

10 Pelta semicircular, lateral lobes not developed; head almost transversely oval,

broadest across cheeks; pronotum very short, more than three times as wide as

long ........... hemidiscus (p. 164)

- Pelta with lateral lobes well developed; head not transversely oval; pronotum

scarcely twice as wide as long ......... 1 1

1 1 Setae B x on tergite I X o 6-0 75 times as long as B 2 ; head less than o 9 times as long

as wide ; mid and hind femora yellow at apex and along posterior margin in distal
third (see also brevicollis) ....... rhizophorae (p. 168)

- Setae B t on tergite IX i -0-1-2 times as long as B 2 ; head longer than wide; mid and

hind femora brown ........... 12

12 Setae B 1 on tergite IX i 2 times as long as B z , but less than o 5 times as long as the

tube; major setae including ocellar and metanotal setae exceptionally stout and
dark aoristus (p. 160)

- Setae B^ on tergite IX about as long as B 2 , and more than 0-5 times as long as the

tube; major setae slender .......... 13

13 Antennal segment III brown with pedicel yellow, IV and V dark brown

carverae (p. 163)

- Antennal segment III largely yellow, shaded near apex, IV and V yellow at base

yanchepi (p. 170)

Nesothrips aoristus Mound

Nesothrips aoristus Mound, 1974 : 68. Holotype $, AUSTRALIA: South Australia (ANIC)
[examined].

The oedymerous male of this species has enlarged fore femora similar to oahuensis,
but the eyes are not prolonged ventrally as in that species. Although the head
and fore legs are similar to Rhaebothrips species the tube and antennae are relatively
short.

NESOTHRIPS COMPLEX

161

SPECIMENS STUDIED.

Holotype $ with 2 $>, 2 <$ paratypes, AUSTRALIA: South Australia, south of
Adelaide, on Olearia ramalosa, 4.xii.ig67 (L. A. Mound] (ANIC, BMNH).

Nesothrips artocarpi (Moulton)

(Text-fig. 38)
Bolothrips artocarpi Moulton, 1942 : 14-15. Holotype $, GUAM (BPBM) [examined].

35

37

FIGS 32-37. Neosmerinthothrips species : 32, N. fructuum. 33, N. hilaris. 34, N.
fructuum, pelta. 35, N. fructuum, tube. 36, N. xylebori, tube. 37, N. variipes, tube.

162 L. A. MOUND

This species, described from two females, is similar to semiflavus in the shape
of the head. The maxillary stylets are retracted into the head as far as the posto-
cular setae, and the postocellar setae are 30 /mi long. The metanotum is almost
unsculptured around the bases of the median setae, and these setae are slender,
40 /mi long and about 45 /mi apart. The pelta has well developed lateral wings.
The tube is relatively long, 195 /mi, with straight sides, and the setae on tergite
nine are dark brown with pale acute apices.

SPECIMEN STUDIED.

Holotype $, GUAM: on Artocarpus communis, 21^.1936 (Sweezey) (BPBM).

Nesothrips brevicoltis (Bagnall)
(Text-figs 40, 43, 44)|

Oedemothrips (?) brevicollis Bagnall, 1914 : 29-30. Holotype $, JAPAN (BMNH) [examined].
Coenurothrips minor Bagnall, 19210 : 287-288. Holotype $, RODRIGUES (BMNH) [examined].

Syn. n.
Neosmerinthothrips formosensis Priesner, 1935 : 368-370. LECTOTYPE $, TAIWAN (HP),

here designated [examined]. Syn. n.
Neosmerinthothrips formosensis var. karnyi Priesner, 1935 : 369-370. LECTOTYPE $, JAVA

(SMF), here designated [examined]. Syn. n.

The name formosensis var. karnyi has been used by Ananthakrishnan in several
publications from Madras (e.g. 1973), but most of the specimens bearing that name
from southern India are referred to here as Neosmerinthothrips fructuum Schmutz.
These specimens have short postocellar setae, the mid and hind femora pale distally,
and the tube broad at the base with convex sides. The single specimen listed
below from India and bearing Ananthakrishnan's identification as formosensis
karnyi has a pair of setae between the posterior ocelli 35 /mi long, the hind femora
yellow along the posterior border, and the tube relatively slender with straight
sides.

Bagnall described this species on a single micropterous female from Japan,
and Priesner described formosensis on two micropterous females from Taiwan (the
only whole specimen is here designated lectotype). Bagnall's specimen is slightly
larger and crushed but cannot otherwise be distinguished. The variety karnyi
and the species minor were both described from macropterous females, which as is
indicated in the key to species have a darker coloured head and median antennal
segments. The males listed below from Hawaii are hemimacropterous, wing
length 500 /mi, and have a similar colour pattern to the macropterae. This also
applies to the micropterous female from Mauritius listed below, although this
specimen has rather long wing lobes, 125 /mi. The Australian species rhizophorae
is possibly only a local colour variant of brevicollis.

In the USNM collections there are three specimens taken in quarentine, apparently
from Guam, which have the ocellar setae scarcely 15 /mi long and arising just behind
a line joining the hind margins of the posterior ocelli. Usually in brevicollis the
ocellar setae are about 50 /mi long and just anterior of a line joining the hind margins

NESOTHRIPS COMPLEX 163

of the posterior ocelli. Mound (1974) refers to similar variation in the position of
the ocellar setae in rhizophorae.

SPECIMENS STUDIED.

Holotype $ of brevicollis, JAPAN: Okinawa, Luchu Is., v. 1913 (J. E. A. Lewis)
(BMNH). Lectotype $ of formosensis, TAIWAN: Hookotoo, Makoo, 5.vi.i93O
(S. Minowa) (HP), with a damaged $ paralectotype on the same slide. Lectotype
$ (labelled holotype) of formosensis var. karnyi, JAVA: Tjibodas, 1400 m, 1923
(Karny) (SMF). Holotype 9 of minor, RODRIGUES: vii-xi. 1918 (H. J. Snell &
H. P. Thomasset) (BMNH).

MAURITIUS: De Pouce Mt, i $, i <$ micropterae on leaves, 2.xi.i939 (R. Mamet)
(HP). REUNION: St Louis, i 9 macroptera on tobacco, I4.vii.i95i (/. R. Williams)
(BMNH).

INDIA: Aryankavu, I $ macroptera on dry twig, 19^.1969 (Ananthakrishnan)
(BMNH); Pachmarki, i $, 20. xi. 1966 (Ananthakrishnan) (HP).

HAWAII: Pupukea, 8 $, 2 ^ on Leucaena, i6.xii.i969; Barbers Point, 2 $, 6 <,
sweeping and on Desmanthus, i6.xii.i969 (F. Andre) (BMNH).

Nesothrips carver ae Mound

Nesothrips carveri Mound, 1974 : 71. Holotype $, AUSTRALIA: South Australia (ANIC)
[examined].

The holotype of this species was collected at Adelaide, but the original description
lists specimens from Western Australia, Victoria and the Australian Capital Terri-
tory. The nomen nudum 'Oedemothrips nigricans BagnalP in Kelly & Mayne
(1934 : 51) refers to this species. The most closely related species is yanchepi with
which carverae has been collected in Western Australia near Perth.

SPECIMENS STUDIED.

AUSTRALIA: South Australia, Adelaide, holotype $ on washing, I5.
(M. Carver) (ANIC). A complete list of specimens is given in Mound (1974).

Nesothrips fodinae sp. n.

$ macroptera. Head, thorax and abdomen dark brown, legs brownish yellow with the
morphological upper surface of all femora deeply shaded at the base, coxae dark brown, tarsi
yellow; antennal segments I-VI yellow, VII and VIII light brown; wings strongly shaded,
particularly at base; major setae dark brown.

Head broader than long, weakly projecting in front of eyes, postocular setae acute; posto-
cellar setae close to inner margin of each posterior ocellus; eyes prolonged on ventral surface.
Antennal sensoria rather slender, two sense cones on III, four sense cones on IV; basal pedicel
of segment VII broad. Pronotum with a weak median thickening; anterior setae reduced,
posterior setae acute; fore femora moderately expanded. Metanotum apparently not sculp-
tured medially, median setae slender and acute, 20 ^m long. Pelta with broad lateral wings;
tergal wing retaining setae reduced on II, sigmoid on 1 1 I-VI I; dorsal and lateral setae on
tergite IX short; sides of tube almost straight.

164 L. A. MOUND

MEASUREMENTS (holotype $ in pm). Body length 1850. Head, length 175; maximum
width 210; basal width 175; postocular setae 50; postocellar setae 23. Pronotum, length
115; median width 230; major setae - aa, am, ml, 16, epim 58, pa 30-35. Fore wing, length
750; distal width 60; subbasal setae 35, 75, 90; number of duplicated cilia 9. Tergite VII
setae, B l 130; B 2 95. Tergite IX setae, B^ 55; B 2 52; B a 70. Tube, length 135; basal width
70; apical width 32; terminal setae no. Antennal segments length (paratype $), 27; 50; 67;
58; 55; 52; 36; 27.

(J macroptera. Head and thorax brown, abdomen dark brown; legs yellow with small
brown markings particularly at base of femora; otherwise similar to $. Fore femora strongly
swollen, fore tibiae stout, fore tarsi with a stout tooth. Pronotum enlarged, anterior margin
and median line strongly thickened; midlateral setae elongate. Metanotal setae 70 yum long.

MEASUREMENTS (paratype in pm). Body length 1800. Head, length 170; maximum
width 187; postocular setae ?go. Pronotum, length 195; median width 250; major setae -
am 20, aa 25, ml 100, epim ?8o, pa ?6o. Fore wing, length 650; subbasal setae 55, 100, 160;
number of duplicated cilia 7. Tergite IX setae, B t 70; B z 60; B 3 120. Tube length 135.

SPECIMENS STUDIED.

Holotype $, FIJI: Lau, in [Ppalm leaf] mines of Promecotheca reichei (Hispidae),
ii. 1932 (R. W. Paine] (BMNH).

Paratypes, 2 $, 4 <$ collected with the holotype, also several immatures of first,
second and fifth instar.

COMMENTS. This species is readily distinguished by the dark body with pale
legs and antennae. N. propinquus is similar in head shape and form of the eyes,
but has a different pelta and usually has a paler head and thorax. The males of
propinquus are apterous. The males of fodinae are interesting not only because
they are macropterous, but also because of the sexual dimorphism in the lengths
of the setae on the head, thorax and wings. The postocellar setae vary from 23-
30 pm but the series is too short to establish whether these also are subject to
sexual dimorphism.

Nesothrips hemidiscus Mound

Nesothrips hemidiscus Mound, 1974 : 71-72. Holotype $, AUSTRALIA: Queensland (ANIC)
[examined].

The D-shaped pelta of this species is unique in the genus.

SPECIMENS STUDIED.

Holotype $ with 7 $, 2 <$ paratypes, AUSTRALIA: Queensland, near Mareeba,
on dead Casuarina twigs, 23.vii.ig68 (L. A. Mound) (ANIC, BMNH).

Nesothrips malaccae sp. n.

(Text-fig. 42)

$ macroptera. Body and legs dark brown, head and abdomen blackish, all femora yellow
in distal third; antennal segments I and II yellow, III light brown with yellow pedicel, IV-
VIII dark brown; major setae black; fore wings deeply shaded, particularly at base, with one
longitudinal dark stripe.

NESOTHRIPS COMPLEX

165

39

40

FIGS 38-45. Nesothrips species: 38, N. artocarpi. 39, N. niger. 40, N. brevicollis.
41, N. fodinae. 42, N. malaccae. 43, N. brevicollis, pelta. 44, N. brevicollis, tube.
45, N. fodinae, pelta.

166 L. A. MOUND

Head broadest behind eyes, cheeks rounded and narrowed to base with two pairs of fine
setae; dorsal surface not sculptured; postocellar setae arise behind the posterior ocelli; posto-
cular setae almost acute ; maxillary stylets wide apart, retracted to postocular setae ; eyes not
prolonged on ventral surface. Antennae typical of genus, two sense cones on segment III,
and four sense cones on IV; VIII not constricted at base. Pronotum transverse, epimeral
sutures complete; anteromarginal setae slender and acute, remaining setae stout and bluntly
acute. Metanotum with a pair of slender setae medially. Pelta with broad lateral wings;
tergal wing retaining setae well developed on II-VII; lateral setae exceptionally long on VII;
tube with straight sides.

MEASUREMENTS (holotype $ in pm). Body length 1900. Head, length 210; maximum
width 215; basal width 150; postocellar setae ?2o; postocular setae 80. Pronotum, length
105; median width 225; major setae - am 35, aa ?2O, ml 35, epim 80, pa 50. Fore wing, length
760; distal width 70; subbasal setae 50, 80, 100; number of duplicated cilia 9. Tergite IV
setae, B^ 100; B 2 55. Tergite VII setae B l 175; B 2 210. Tergite IX setae, B t 160; B 2 145;
B 3 160. Tube, length 210; basal width 95; apical width 42; terminal setae 130. Antennal
segments length, 30; 50; 68; 64; 58; 55; 39; 26.

SPECIMENS STUDIED.

Holotype $, WEST MALAYSIA: Kuala Lumpur, on dead branch, 29. xi. 1969 (R. G.
6- F. Andre) (BMNH).

Paratypes. WEST MALAYSIA: i $ with data similar to holotype except 24.xii.i969
(BMNH). SUMATRA: Fort de Kock, 920 m, xi. 1920 (Jacobson) (SMF) [this paratype
bears a manuscript name].

COMMENTS. This species appears to be related to brevicollis and rhizophorae
but has the dorsal setae (B^ on tergite nine elongate, and all the femora dark brown
with a sharply yellow apex. The paratype female from Sumatra is larger than the
two specimens from Malaya (body length 2400 /xm), and was designated holotype
of an unpublished name by Dr H. Priesner.

Nesothrips melinus Mound

Nesothrips melinus Mound, 1974 : 72-73. Holotype <, AUSTRALIA : Queensland (ANIC) [examined].

This species was described from four males collected in northern Queensland,
Australia, although a fifth male from southern New South Wales may represent
the same species. These males are largely yellow, and in structure appear to be
intermediate between Nesothrips and the Carientothrips species of the mjobergi
complex.

SPECIMENS STUDIED.

Holotype $ with 3 $ paratypes, AUSTRALIA: Queensland, near Ingham, on dead
twigs, 2i.vii.i968 (L. A. Mound] (ANIC, BMNH).

Nesothrips niger (Moulton & Stein weden)
(Text-fig 39)

Bolothrips nigra Moulton & Steinweden, 1932 : 167-168. Holotype $, MARQUESAS Is. (BPBM)
[examined] .

NESOTHRIPS COMPLEX 167

This species is known only from the unique holotype which is mounted under a
broken cover glass. Because of this its relationships cannot be determined at
present. Contrary to the original description the lengths of the tube and B 1 setae
on tergite nine are 260 /urn and 210 /mi.

SPECIMEN STUDIED.

Holotype $, MARQUESAS Is.: Hiva Oa, Mt Temetiu, 24.vii.i929 (Adamson &
Mumford) (BPBM).

Nesothrips oahuensis Kirkaldy
(Text-fig. 31)

Nesothrips oahuensis Kirkaldy, 1907 : 103. Syntype $, HAWAIIAN Is.: Oahu (PBPBM) [not

examined] .
Oedemothrips laticeps Bagnall, 1910 : 680-681. Syntypes , $, HAWAIIAN Is.: Oahu, Lanai

(BMNH) [examined]. [Synonymized by Bianchi, 1944.]
Nesothrips hawaiiensis, lapsus for oahuensis Kirkaldy, Bianchi, 1944 : 31-38.

Bianchi studied a male and female specimen from Kirkaldy's original material
and illustrated these. The present author has examined only two female and one
oedymerous male syntypes of laticeps. The illustration given by Bianchi clearly
refers to a rather small male with the fore femora weakly swollen. The male
illustrated here has a greatly enlarged fore femur which is L-shaped as in Rhaebo-
thrips species. Moreover the head is prolonged in front of the eyes and the ocellar
setae are rather close together. The pelta of this specimen is very broad and
slender.

SPECIMENS STUDIED.

i $ syntype of laticeps, HAWAIIAN ISLANDS: Oahu, near Honolulu, 2000-3000 ft,
vii. 1900 (Perkins 667); I $ syntype, HAWAIIAN ISLANDS: Lanai, 2000 ft, i. 1894
(Perkins 91); i $ syntype without data (Perkins 489) (BMNH).

Nesothrips propinquus (Bagnall)

Oedemothrips propinquus Bagnall, 1916 : 408-409. Holotype $>, AUSTRALIA: Victoria (BMNH)

[examined].
Cryptothrips dimidiatus Hood, 1918 : 145-146. Holotype $, AUSTRALIA: Queensland (USNM)

[examined]. [Synonymized by Mound, 1968 : 141.]
Oedemothrips propinquus var. breviceps Bagnall, 1924 : 634-635. Syntype $, NEW ZEALAND

(BMNH) [examined]. [Synonymized by Mound, 1968 : 141.]
Oedemothrips propinquus form obscuricornis Bagnall, 1924 : 635.
Bagnalliella cestosa Karny, 1920 : 41. Holotype $, AUSTRALIA: Queensland (NR) [examined].

[Synonymized by Mound, 1974 : 73-1
Neosmerinthothrips oleriae Moulton, 1949 : 492-494. Holotype $, SOUTH AFRICA (CAS) [not

examined]. [Synonymized by Mound, 1968 : 141.]
Bolothrips similis Hartwig, 1948 : 103-108. Holotype $, SOUTH AFRICA: Pretoria [not

examined]. Syn. n.

168 L. A. MOUND

In addition to the above synonymy the nomen nudum 'Oedemothrips australis
Bagnall' in Kelly & Mayne (1934 : 51) also refers to this species. The species
is widespread and common in Australia and New Zealand, as well as in South
Africa, and has been recorded from St Helena and New Caledonia. Material from
all these localities is available in the BMNH collections, together with paratypes
of oleriae and similis. In the USNM collections there are three females taken in
quarantine at Boston apparently from the Azores and Portugal. Moreover in the
BMNH collections there is one female apparently collected at Curitiba in southern
Brazil, but this is the only Neotropical record and needs further confirmation.

Males and most females of propinquus are apterous but female macropterae are
collected occasionally. The variation in head shape in Australia includes the form
shown by similis, and the ventral prolongation of the eyes is not constant. In
a few gynaecoid males the eyes are not elongate ventrally. The colour of the legs
and antennae is also variable, and the form obscuricornis has the legs and antennae
largely brown. The anterior setae of the pronotum also vary in length from 15 to
60 jum, but there is no evidence from all this variation that more than one species
is involved. Populations from New Zealand appear to show the greatest degree
of variation.

Nesothrips rhizophorae (Girault)

Cryptothrips rhizophorae Girault, 1927(38) : 2. Syntype $ $, AUSTRALIA: Queensland (QM)

[examined].
Nesothrips rhizophorae (Girault) Mound, 1974 : 74~75-

This species is very similar to brevicollis (Bagnall) and may be merely a local
colour variant with the basal antennal segments brownish instead of yellow.

SPECIMENS STUDIED.

AUSTRALIA: Queensland, Brisbane, 3 <j>, i $ syntypes from galls on mangrove,
28.vi.ig27 (L. Franzen) (QM); Brisbane, Mt Nebo, i $ from shrubs, 27.vii.ig68
(L. A. Mound) (BMNH).

Nesothrips semiftavus (Moulton)
Bolothrips semiflavus Moulton, 1939 : 147-148. Holotype Q, RAPA Is. (BPBM) [examined].

This species is known only from the unique holotype. It appears to be similar
to artocarpi in having the head slightly longer than wide, but there are two pairs
of long setae behind the eyes, the postoculars no /mi and the mid-dorsals 70 /mi
long. The postocellar setae are also 70 /Am long. The median setae on tergite
nine are 155 jam and the tube 175 /mi long. The major setae appear to be paler
and more slender than in other species of Nesothrips.

SPECIMEN STUDIED.
Holotype $, RAPA ISLAND: Mt Tevaitahu at 700 ft, 8.vii.i935 (BPBM).

NESOTHRIPS COMPLEX

47

169

FIGS 46-5 1. 46, Nesidiothrips alius. 47, Phacothrips ocelloides. 48, N. alius, pelta.
49, P. ocelloides, pelta. 50, AT. a/iws, tube. 51, P. ocelloides, tube.

170 L. A. MOUND

Nesothrips yanchepi Mound

Nesothrips yanchepi Mound, 1974 : 75- Holotype $, AUSTRALIA: Western Australia (ANIC)
[examined].

This species is similar to niger (Moulton & Steinweden) but has the eyes less
prolonged ventrally and the setae on tergite nine shorter. It is also closely related
to carveri.

SPECIMENS STUDIED.

Holotype $ with i $, i ^ paratypes, AUSTRALIA: Western Australia, Yanchep
near Perth, from sedges at lakeside, 2Q.ix.i967 (L. A. Mound] (ANIC, BMNH).

PHACOTHRIPS gen. n.

Type-species: Gastrothrips ocelloides Hood.

Medium sized, dark brown species of Idolothripinae, Cryptothripini. Head as in Gastro-
thrips, not projecting at anterior, eyes large, stylets wide apart, but with one large isolated
ommatidium on each cheek midway between posterior margin of head and posterior margin
of eye. Antennae with two sense cones on segment III, four sense cones on segment IV;
segment VIII more than 0-6 times as long as VII but barely constricted at base. Fore tarsal
tooth well developed in both sexes. Median metanotal setae small. Tube very stout, sides
strongly convex, maximum width more than 0-5 times the total length, and about 5-0 times
the minimum width, terminal setae very weak. Macropterous $ with no duplicated cilia;
pelta triangular with slender lateral wings. Micropterous <$ sometimes oedymerous with
enlarged pronotum, pelta broadly triangular, metanotum and tergites with supernumerary
setae.

This genus is possibly related to the nigrisetis-group of species from South America
which is treated here under Neosmerinthothrips. It is distinguished from other
Thysanoptera by the ommatidia on the cheeks and the heavy tube which resembles
that of some Pygothrips species.

Phacothrips ocelloides (Hood) comb. n.
(Text-figs 47, 49, 51, 68)

Gastrothrips ocelloides Hood, 1950 : 9-12. Holotype $, BRAZIL (USNM) [not examined].

This species is here removed from Gastrothrips on account of the four sense cones
on the fourth antennal segment, as well as the structure of the head and tube.

SPECIMENS STUDIED.

BRAZIL: Distrito Federal, Jacarepagua, i $, i <$ paratype on dead branches,
14^.1948 (/. D. Hood); Sao Paulo, Itanhaen, i $ paratype (sic) on dead branches,
17^1.1948 (/. D. Hood & J. Lane) (AMG); Vicosa, on dead branches, 2 $, i <$,
vii. 1964, 5 ?, i cJ, xi. 1964 (F. Andre) (BMNH).

NESOTHRIPS COMPLEX 171

RHAEBOTHRIPS Karny

Rhaebothrips Karny, 1913 : 128. Type-species: R. lativentris Karny, by monotypy.

A revision and diagnosis of this genus was published recently by Sakimura (1971).
However, considerably more material is now available and Rhaebothrips is used
here in a broader sense. Two species are described below which have a small
fore tarsal tooth in the female and one of these, leveri, is unusually small with a
short tube, similar to some Nesothrips species. The other species, doulli, is interest-
ing because it is closely related to two further new species from New Zealand which
do not have a fore tarsal tooth. One of these species is unique in the genus in
having the eyes prolonged on the ventral surface of the head as in some Nesothrips
species. The distinction between these two genera is tenuous and likely to become
more so with further collecting in the Pacific area. Rhaebothrips species always
have a stout pair of setae relatively close together between the posterior ocelli.
They are usually larger than Nesothrips species with relatively longer antennae and
tube.

KEY TO THE SPECIES OF RHAEBOTHRIPS

1 Antennal segment III largely yellow or yellowish brown; V paler at base . . 2
Antennal segment III brown with pedicel pale; V uniformly dark brown . . 4

2 Fore tarsal tooth present in $; antennal segment III less than 2 -o times as long as

wide (Text-fig. 74) [Fiji] leveri (p. 175)

- Fore tarsal tooth absent in $; antennal segment III more than 3-5 times as long as

wide .............. 3

3 Lateral setae in abdominal segments III-VIII light brown to yellowish

lativentris (p. 174)

- Lateral setae on abdominal segments III-VIII dark brown to black .nigrisetis (p. 175)

4 Setae J3 t on tergite IX more than 0-8 times as long as tube, sometimes longer than

tube [Pacific Is.] major (p. 175)

Setae B t on tergite IX less than 0-6 times as long as tube [New Zealand] . . 5

5 Head o- 9-1-0 times as long as tube (Text-fig. 58); antennal segment III 2 -1-2 -2

times as long as wide (Text-fig. 72) ...... zondagi (p. 176)

Head i -2 times as long as tube, antennal segment III 2-4-2-5 times as long as wide

(Text-figs 67, 71) 6

6 Fore tarsal tooth present in $ (only micropterae and hemimacropterae known) ; eyes

not prolonged on ventral surface of head (Text-fig. 52) . . doulli (p. 171)

- Fore tarsal tooth not present in $ (only macropterae known) ; eyes i 4 times as long

on ventral surface of head as on dorsal surface (Text-fig. 53) . . eastopi (p. 173)

Rhaebothrips doulli sp. n.

(Text-figs 52, 55, 67)

$ microptera. Colour dark brown with red internal pigment; tarsi and fore tibiae a little
paler; antennae dark brown except for yellow pedicel of segment III; fore wing rudiment
pale; major setae not very dark, with finely acute apices.

Head longer than wide, cheeks rounded, eyes rather small; ocellar setae arising between
posterior ocelli, median setae on vertex sometimes elongate; maxillary stylets retracted to
postocular setae; antennae similar in structure to eastopi. Pronotum with median thickening

172

L. A. MOUND

FIGS 52-57. Rhaeobothrips species: 52, R. doulli, male. 53, R. eastopi. 54, R.
leveri. 55, 7?. dow//z, fore leg of male. 56, R. eastopi, tube. 57, R. leveri, pelta.

NESOTHRIPS COMPLEX 173

developed, epimeral sutures complete; metanotum not sculptured, median setae 50 fj,m long
and 1 20 p.m apart. Fore femur simple, fore tarsus with a small pointed tooth at apex of inner
margin about one-third as long as tarsal width. Fore wing lobes with a few cilia. Pelta
broad with small lateral wings; tube with straight sides but slightly constricted near apex.

MEASUREMENTS (holotype $ in ^m). Body length 2900. Head length 310; median width
260, ocellar setae 100 long, bases 55 apart; post ocular setae 145; mid dorsal setae 85 (35 in
$ paratype). Pronotum, length 175; median width 370; major setae -am 50, aa 50, ml 8085,
epim 120, pa 100120. Fore wing, length 450; subbasal seta 140. Tergite IV setae, B l
175, B a 100. Tergite VII setae, ^ 240; B 2 280. Tergite IX setae, B 1 145, B 2 150, B 3 240.
Tube, length 260; basal and apical width, 100, 58; terminal setae 195. Antennal segments
length, 50, 70, 105; 100; 85; 77; 55; 40.

<$ microptera. Colour similar to $ (2 <$ paratypes teneral). Pronotum strongly thickened
medially and along anterior border; fore femur enlarged L-shaped; fore tarsal tooth curved,
almost as long as tarsal width ; major setae of head and pronotum longer than in $.

MEASUREMENTS (paratype < in jam). Body length 2400. Head length 290; ocellar setae
100; postocular setae 160, mid-dorsal seta 50. Pronotum, length 225; median width 350;
major setae - am 50, aa 65, ml 145, epim 160, pa 145. Fore wing, length 400; subbasal setae
145, 230. Tube length 225.

SPECIMENS STUDIED.

Holotype $, NEW ZEALAND: Christchurch, on dead walnut twig, 1951 (K. M.
Doull] (BMNH).

Paratypes. NEW ZEALAND: i <j>, 4 < collected with holotype (BMNH) (i $
deposited in DSIR) ; I $, I <$ on Cytisus bark, Cave [100 miles south-west of Christ-
church], g.vii.igGg (R. Zondag) (NZFI).

COMMENTS. This species is closely related to eastopi described below which
unfortunately is known only from a different morph. The third antennal segment
of these two species differs from lativentris and major in being slightly swollen in
the basal third. The female paratype from Cave has fore wings 950 pm long and
a rather stouter fore tarsal tooth than the micropterae, but the male with which
it was collected cannot be distinguished from the other male paratypes.

Rhaebothrips eastopi sp. 11.

(Text-figs 53, 56, 71)

$ macroptera. Colour dark brown, fore tarsi yellowish, fore tibiae light brown; antennae
brown to dark brown, III slightly lighter with pedical yellow; fore wings very weakly shaded;
major setae light brown and finely acute.

Head longer than wide; ocellar setae arising between posterior ocelli, maxillary stylets not
retracted to postocular setae, eyes prolonged and narrowed on ventral surface; antennal seg-
ment III slightly swollen in basal third, VII with a distinct pedicel, two sense cones on III,
four on IV. Pronotum smooth, median thickening not developed, epimeral sutures complete;
metanotum weakly reticulate medially, setae 65 jj,m long. Fore tarsus angulate on inner
surface but without a tooth; fore wing narrow (? partially reduced). Pelta with lateral wings
extending almost to lateral margins of tergite II; tube with almost straight margins.

MEASUREMENTS (holotype $ in //m). Body length 2800. Head length 320; median width
255; ocellar setae 65 long, bases 62 apart; postocular setae 135. Pronotum, length 195;
median width 310; major setae - am 35, aa 25, ml 50, epim 100-110; pa 95-100. Fore wing,
length 950; distal width 80; subbasal setae 50, 50; number of duplicated cilia 3-4. Tergite

E*

I 7 4 L - A - MOUND

IV setae, -B x 145; B 2 80. Tergite VII setae, B t 245; B z 225. Tergite IX setae, B^ 145; B t
130; B 3 225. Tube, length 260; basal and apical width 100, 52; terminal setae 195. Antennal
segments length, 50, 65, 100, 115, 100, 70, 56, 45.

SPECIMEN STUDIED.

Holotype <j>, NEW ZEALAND: South Island, 15 miles S.E. of Greymouth, Moana
Kotula, from Uncinia andjuncus, I2.ix.igj2 (V. F. Eastop) (BMNH).

COMMENTS. The unique holotype is possibly not fully macropterous. The
antennae, particularly segment three, are similar to doulli described above, but
that species has a distinct fore tarsal tooth in the female and the eyes are not
prolonged ventrally.

Rhaebothrips lativentris Karny

Rhaebothrips lativentris Karny, 1913 : 129-130. Holotype^, TAIWAN (? lost) [not examined].
Cryptothrips claripennis Hood, 1919 : 90. Holotype $, AUSTRALIA: Queensland (USNM)

[examined]. [Synonymized by Mound, 1974 : 91.]
Cryptothrips seychellensis Bagnall, 1921 : 274-276. Lectotype <$, SEYCHELLES (BMNH)

[examined]. Syn. n.
Cryptothrips difficilis Bagnall, 1921 : 276. Holotype $, SEYCHELLES (BMNH) [examined].

Syn. n.
Machatothrips ipomoeae Ishida, 1932 : 12-14. Holotype $, PONAPE (Hokkaido Univ.) [not

examined]. [Synonymized by Kurosawa, 1968 : 60.]
Cryptothrips magnus Moulton, 19280 : 299. Holotype $, TAIWAN (CAS) [not examined].

[Synonymized by Sakimura, 1972 : 668.]
Gynaikothrips yuasai Moulton, 19286 : 315. Holotype $, TAIWAN (CAS) [not examined]

[Synonymized by Sakimura, 1972 : 668]
Rhaebothrips fuscus Moulton, 1942 : 15-16. Holotype $, GUAM (BPBM) [not examined].

[Synonymized by Sakimura, 1971 : 393-]
Bolothrips australiensis Moulton, 1968 : 118-119. Holotype $, LORD HOWE ISLAND (CAS)

[examined]. [Synonymized by Mound, 1974 : 91.]

This remarkable synonymy is a reflection partly of the variation in structure
of lativentris, but more particularly of the low standard of preparation of most of
the microscope slides on which these nominal species were based. Sakimura (1971)
has given an account of the variation of the species, and moreover distinguished
a new species, nigrisetis from Fiji and New Guinea. However, in the opinion
of the present author, lativentris is native to - and has its centre of diversity in -
the western Pacific. Individuals with black setae which correspond to the descrip-
tion of nigrisetis have been studied from the Solomon Islands and the New Hebrides,
and also from Fiji where they were collected together with normal lativentris. It
seems unlikely that two species are involved in this pattern of variation although
the names are retained separate for the present.

There are specimens of lativentris in the BMNH collection from Samoa, Hawaii,
Guam, Solomon Islands, New Guinea, Queensland, Malaya, Seychelles, Jamaica,
Cayman Islands and Trinidad. Sakimura (1971 ; 1972) also lists southern Japan,
Taiwan, Philippines, Java, Mauritius, Puerto Rico. Cuba, Dominican Republic,
Panama, Bahama and Florida.

NESOTHRIPS COMPLEX 175

Rhaebothrips leveri sp. n.

(Text-figs 54, 57, 74)

$ macroptera. Colour dark brown, tarsi yellow, fore tibiae with yellow markings; antennal
segments I and VI-VIII brown, II yellow at apex, III and IV yellow but light brown distally,
V brown with a yellow pedical ; fore wing shaded ; major setae dark brown.

Head longer than wide; ocellar setae arising on line joining posterior margins of posterior
ocelli; postocular setae acute; maxillary stylets retracted to postocular setae; antennal seg-
ments VI-VIII compact, two sense cones on III, four on IV. Pronotum transverse, median
thickening slender; epimeral sutures just complete; epimeral setae bluntly acute. Metanotum
not sculptured medially, setae 30 ^m. Fore tarsal tooth acute but less than half as long as
tarsal width. Pelta with slender lateral wings ; tergal lateral setae not long, blunt at apex;
sides of tube straight.

MEASUREMENTS (holotype $ in |nm). Body length 1850. Head length 220; width 210;
postocellar setae 60; postocular setae 90. Pronotum, length 105; median width 260; major
setae - am 35, aa 35, ml 58, epim 80-85, pa 50-60. Fore wing, length 750; distal width 65;
subbasal setae 40, 58, 58; number of duplicated cilia 5-6. Tergite IV setae B-^ 80; B z 65.
Tergite VII setae, B^ 135; B z 100. Tergite IX setae, -B x 115; B 2 125, B 3 135. Tube, length
145; width at base 80, at apex 42; terminal setae 135. Antennal segments length, 26, 50,
60; 50; 42; 45; 35; 20.

SPECIMEN STUDIED.
Holotype $, FIJI: Viti Levu, Vatuwaga, I2.vii.i94i (R. A. Lever) (BMNH).

COMMENTS. This species could equally well be placed in Nesothrips. It is
here described in Rhaebothrips partly because it is more closely related to lativentris
than to N. oahuensis, and partly because the only other related species with a fore
tarsal tooth is R. doulli from New Zealand.

Rhaebothrips major Bagnall

Rhaebothrips major Bagnall 1928 : 75-76. PHolotype <$, SAMOA (Plost) [not examined].

The (? unique) holotype male, not female as stated in the original description,
should be in BagnalPs collection at the BMNH but cannot be found (Mound, 1968).
Sakimura (1972) has given a redescription of major from specimens collected in
Samoa, and there is one macropterous male in the BMNH collections from the
Solomon Islands apparently of this species.

Rhaebothrips nigrisetis Sakimura

Rhaebothrips nigrisetis Sakimura, 1972 : 400-402. Holotype^, FIJI (BPBM) [not examined.]

Sakimura described this species from nearly eighty specimens collected in Fiji
and one male from New Guinea. As mentioned above there are specimens appar-
ently of nigrisetis in the BMNH collection from New Hebrides, the Solomon Islands
and New Guinea, as well as some from Fiji which were collected with lativentris,
and it is doubtful if these two species are really distinct. The Rhaebothrips speci-
mens from New Guinea are particularly difficult. Specimens in one series in the

176

L. A. MOUND

BMNH collected on various dates at Bulolo in S.E. New Guinea are similar to
nigrisetis but with the antennae darker. Another series in the BPBM from N.W.
New Guinea (Irian Barat) near Nabire and Lake Wissel have black setae but are
much larger than any other Rhaebothrips which have been studied.

Rhaebothrips zondagi sp. n.

(Text-figs 58, 72)

5 macroptera. Colour brown, head and tube darkest; fore tarsi little paler than femora;
antennae dark brown, pedicel of segment III yellow; fore wing uniformly shaded dark brown;
major setae brown but not very dark, with finely acute apices.

58

FIG. 58. Rhaebothrips zondagi, male.

NESOTHRIPS COMPLEX 177

Head longer than wide, ocellar setae between posterior ocelli ; maxillary stylets not retracted
to postocular setae; antennae relatively short but similar to those of eastopi. Pronotum
with weak median thickening; epimeral sutures complete; median area of metanotum not
sculptured, setae 50-60 pm. long and 50jnm apart. Fore femur simple, fore tarsus with no
tooth; fore wing relatively broad. Lateral wings of pelta not extending fully across anterior
margin of tergite II ; sigmoid setae well developed on II-VII ; tube with straight sides.

MEASUREMENTS (holotype $ inprn). Body length 2450. Head, length 260; median width
225; ocellar setae 70, bases 35 apart; postocular setae 100. Pronotum, length 160; median
width 275; major setae - am 35, aa 20, md 50, epim 90, pa 100. Fore wing, length noo;
distal width 115; subbasal setae 65, 100, 115, number of duplicated cilia 12. Tergite IV
setae, B^ 125, B a 80. Tergite VII setae, B : 220, B z 195. Tergite IX setae, B^ 120; B 2 145;
B 3 195. Tube, length 275; basal and apical width 95, 50; terminal setae 160. Antennal
segments length, 40, 65, 90, 80, 70, 65, 50, 35.

(J macroptera. Colour and structure similar to $; frequently oedymerous with fore femora
enlarged and L-shaped, and fore tarsus with a stout tooth equal in length to tarsal width;
major setae of head and pronotum longer than in $; fore wing relatively small.

MEASUREMENTS (paratype <$ in fj,m). Body length 2300. Head, length 260; median width
210; ocellar setae 90; post ocular setae 135; mid dorsal setae 50-65. Pronotum, length 195;
median width 320; major setae - am 55, aa 30, ml 115, epim 100, pa 135. Fore wing, length
900; distal width 85; number of duplicated cilia 8; sub-basal setae 115, 160. Tergite IX
setae, B 100; B 2 95; B 3 225. Tube length 255.

SPECIMENS STUDIED.

Holotype $, NEW ZEALAND: South Island, Hochstetter State Forest 25 miles
east of Greyinouth, on Phyllocladus alpinus branches, 13.111.1970 (W. A. Holloway)
(DSIR).

Paratypes. NEW ZEALAND: 13 , 8 $ collected with the holotype; 2 ? North
Island, Mangawiri Basin, Whiriaki (Urewera), 40 miles south-west of Rotorua,
on Podocarpus totara twigs, 13.^.1972 (P. J. Alma) (DSIR, NZFI, BMNH).

COMMENTS. This species is related to the other two species described above
from New Zealand in having short setae on tergite nine. The antennae are shorter
than any other species of Rhaebothrips , apart from the aberrant leveri from Fiji,
but the tube is longer than any species of Nesothrips with which it might be confused.

SCOTOTHRIPS Priesner

Scotothrips Priesner, 1939 : 75. Type-species: Adiaphorothrips elephas Karny, by original
designation.

The synonymy of this genus and its relationships to Dichaetothrips have been
discussed recently by Mound (1974). Scotothrips species are found mainly in the
Pacific, Indonesian and Australian regions, and the Brazilian species firmus listed
below is not typical of the group.

Scotothrips claripennis (Moulton) comb. n.

Dichaetothrips claripennis Moulton, 1934 : 503. Holotype $, HAWAII (CAS) [not examined].
Gastrothrips trinidadensis Hood, 1935 : 168-170. Holotype $, TRINIDAD (USNM) [not
examined] . Syn. n.

i 7 8

L. A. MOUND

Nesothrips indicus Ananthakrishnan, 1968 : 967-969. Syntypes <$ ty, INDIA: Madras (TNA)

[not examined]. Syn. n.
Nesothrips diver sus Ananthakrishnan, 1972 : 434-436. Holotype $, INDIA: Madras (TNA)

[not examined]. Syn. n.

This species was based on a single female collected in Honolulu. This specimen
has not been studied, but the above synonymy is based on a female from Honolulu
determined as claripennis by Bianchi. This species does not belong in Nesothrips

FIGS 59-66. Right antennae (III-VIII unless stated otherwise) : 59, Bolothrips italicus.
60, Carientothrips acti, III. 61, C. fijiensis, III-IV. 62, C. japonicus, III lateral.
6^,C.grayi. 64, Gastrothrips falcatus . 65, G. ruficauda. 66, Diceratothrips validipennis .

NESOTHRIPS COMPLEX 179

because of the presence of a fore tarsal tooth in the female, nor in Gastrothrips
because of the four sense cones on the fourth antennal segment. The elongate
straight-sided tube is typical of the Scotothrips I Dichaetothrips complex, and because
the postocellar setae are short claripennis is here placed in Scotothrips.

The vertex of this species is narrowed in dorsal view but is very deep. This
shape is very susceptible to distortion in mounting, and the claripennis and indicus
specimens which have been studied have the head apparently rather broad at the
base. The specimens described as diver sm are here regarded as small pale forms
of the larger species. The most constant characteristic in all this material is the
presence of a pair of pores on the metanotum anterolateral to the metanotal setae.
The colour of antennal segments three and four is variable, although three is usually
yellow with a shaded apex and four is usually light brown. The mid and hind
tarsi are usually yellow but sometimes light brown. In the BMNH collections
there are several specimens of Dichaetothrips usitatus Ananthakrishnan & Jagadish,
1970 (Dichaetothrips indicus Ananthakrishnan, 1961) which had been misidentified
as Nesothrips indicus Ananthakrishnan, 1968 despite the presence of very long
postocellar setae. One of the male paratypes of diversus is also a misidentification
of usitatus. The species claripennis has probably been transported around the
tropics by man.

SPECIMENS STUDIED.

HAWAIIAN ISLANDS: Oahu, Honolulu, Mckinkey High, i $ in Monkey Pod,
iv. 1946 (F. A. Bianchi] (BMNH).

MEXICO: 3 , 2 c? on Cocos nucifera seed, in quarantine at US (USNM). BAHAMAS:
I $ on Cocos nucifera seed, I $ on Poinciana regia pod, in U.S. quarantine (USNM).
JAMAICA: Caenwood, I $, vi. 1970 (K. Heinze) (BMNH); I $, 2 < on Cocos nucifera
fls, i $ on Delonix regia pod, in U.S. quarantine (USNM); TRINIDAD: St Clair, i $
paratype of trinidadensis in hole in dead bamboo, 13.1.1917 (C. B. Williams) (AMG).

MOZAMBIQUE: Lourenco Marques, I $ in Cassia pods, vii. 1936 (/. C. Faure)
(AMG); SOUTH AFRICA: Natal, Winterskloof, 2 $, i $ in Cassia pods, vii. 1942
(F. Casalis) (AMG).

INDIA: Madras, i $, 2 <$, determined as indicus, ii. 1966, i $ paratype of diversus,
xii. 1970 (T. N. Ananthakrishnan) (BMNH).

Scotothrips firmus (Hood) comb. n.

Gastrothrips firmus Hood, 1952 : 162. LECTOTYPE $>, BRAZIL (USNM), here designated
[not examined].

The head of this species is unusual in Scotothrips as the cheeks are constricted
near, but not at, the base, and the eyes are slightly swollen at their posterior margin.
The fore tarsus bears a small tooth in the female, the pronotum is thickened medially
and at the anterior margin, the median metanotal setae are small and slender, the
pelta broadly triangular with the median pores close together, and the tube is long
with straight sides. The specimen here designated as lectotype has not been studied

i8o

L. A. MOUND

but is that specimen which was labelled holotype by Hood but not referred to in
his text. The lectotype bears identical data to the paratype listed below and has
been given the following number: USNM Type Number 71995.

SPECIMEN STUDIED.

BRAZIL: Sao Paulo, Itanhaen, I $ paratype on dead branches, 17^.1948 (J. D.
Hood &J. Lane) (AMG).

67

FIGS 67-74. Right antennae (III-VIII unless stated otherwise) : 67, Rhaebothrips
doulli. 68, Phacothrips ocelloides. 69, Nesidiothrips alius. 70, Neosmerinthothrips
fructuum. 71, Rhaebothrips eastopi, III. 72, R. zondagi. 73, Neosmerinthothrips
variipes. 74, Rhaebothrips leveri.

NESOTHRIPS COMPLEX 181

SYNCEROTHRIPS Hood gen. rev.
Syncerothrips Hood, 1935 : 191-192. Type-species: S. harti Hood, by monotypy.

Stannard (1957) synonymised this genus with Nesothrips on the grounds that the
last two antennal segments of the type-species of Neosmerinthothrips were more
or less closely joined. However, Syncerothrips is closely related to the South
American species of Gastrothrips. There are three sense cones on the fourth antennal
segment and two on the third, the median metanotal setae are very stout, and the
pelta triangular. The only species in the genus differs from Gastrothrips species
in having the last two antennal segments fused with only a partial suture on the
ventral surface. The fore tarsus does not bear a tooth in the female, and the
postocellar setae are long.

Syncerothrips harti Hood

Syncerothrips harti Hood, 1935 : 192-194. Holotype$, U.S.A.: Texas (USNM) [not examined].

This species was based on a single female, but the specimen listed below was
taken at the type-locality and identified by Hood.

SPECIMEN STUDIED.

U.S.A.: Texas, Brownsville, I $ microptera on dead branches, 2.iii.i93Q (J. D.
Hood) (USNM).

SPECIES REMOVED FROM THE NESOTHRIPS COMPLEX

The following three species were described in genera related to Nesothrips, but
they are not even members of the subfamily Idolothripinae in its strict sense.
Species of the genus Adelothrips have moderately broad maxillary stylets and
probably feed on fungus spores, but in view of the structure of the male abdomen
it is likely that this genus is derived from Hoplothrips-like members of the Phlaeo-
thripinae (Mound, 1974).

Adelothrips lativerticis (Post) comb. n.
Bolothrips lativerticis Post, 1961 : 140-143. Holotype $, U.S.A.: Oregon (CAS) [examined].

The original illustration of this species shows five sense cones on the third antennal
segment and six on the fourth, although the description gives the actual number
correctly as three and four. There are two pairs of wing retaining setae on each
tergite, unlike any member of the Cryptothripini discussed in this paper. Moreover
the males of lativerticis, like other members of Adelothrips, have setae B 2 on tergite
nine short and stout, and sternites six and seven have glandular areas. This
species is a typical member of Adelothrips because of the number of antennal sensoria,
the fusion of the last two antennal segments, the position of the maxillary stylets
close together in the middle of the head, the bell-shaped pelta, and the abdominal
structure detailed above.

182 L. A. MOUND

SPECIMENS STUDIED.

Holotype <j>, U.S.A.: Oregon, Hood River County, Herman Creek, in hollow twigs,
31.1.1946 (R. L. Post) (CAS).

U.S.A.: Oregon, Benton County, Corvallis, paratype <j>, 23.^.1946 (Leah & Post)
(R. L. Post coll. North Dakota); Washington, Puyallup, 3 $, 2 <$ in dead willow,
20.iv.i937 (B. Baker) (BMNH).

Adelothrips speciosissimus (Karny)

Nesothrips speciosissimus Karny, 1920 : 42. Holotype <$, AUSTRALIA: Queensland (NR)

[examined].
Adelothrips speciosissimus (Karny) Mound, 1974 : 16.

This species may not be native to Australia as the most closely related species
appears to be skwarrae Priesner from Mexico. The male has setae B 2 on tergite
nine short, and sternites six and seven have reticulate glandular areas.

SPECIMENS STUDIED.

Holotype <, AUSTRALIA: Queensland, Cedar Creek, March (Mjoberg) (NR).

AUSTRALIA: Queensland, Clump Point, Mission Beach, 2 $ on dead branches,
2i.vii.i968 (L. A. Mound 764) (BMNH).

Liothrips debilis (Hood) comb. n.
Neosmerinthothrips (?) debilis Hood, 1936 : 269-271. Holotype^, PANAMA (USNM) [examined].

This species was described from a single male with stout maxillary guides but
with slender maxillary stylets. It is a member of the subfamily Phlaeothripinae
despite the fact that all three pairs of major setae on tergite nine are elongate,
a characteristic shared with several South American species in the genus Liothrips.
The genus is widespread in the tropics and the species feed on the leaves of green
plants.

SPECIMEN STUDIED.

Holotype $, PANAMA: Frijoles, on Panicum maximum, 23.^1.1933 (P. C. Standley)
(USNM).

REFERENCES

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1961. Studies on some Indian Thysanoptera VI. Zool. Anz. 167 : 259-271.

1964. A contribution to our knowledge of the Tubulifera (Thysanoptera) from India.
Opusc. ent. Suppl. 25 : 120 pp.

NESOTHRIPS COMPLEX 183

1968. Two new species of Nesothrips with further remarks on Nesothrips robustus (Thysan.

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1 86

L. A. MOUND

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INDEX
Synonyms and invalid names are in italics.

abditus, 138
Acallurothrips, no
acti, 126
acuticornis, 139
Adelothrips, 181
affinis, 150
alius, 157
alticola, 140
Allothrips, in
andrei, 118
annulipes, 150
anolis, 142
aoristus, 160
apterus, 127
arenarius, 121
artocarpi, 161
australicus, 131
australiensis, 174
australis, 168

badius, 127

bagnalli, 119

bicolor, 118

Biconothrips, in

biformis, 127

Bolothrips, 117

Botanothrips, 117

brachyurus, 121

brasiliensis , 138

breviceps, 167

brevicollis, Neosmerinthothrips, 151

brevicollis, Nesothrips, 162

brevicornis, 118

callipus, 142
capitalis, 143
capricornis, 127
Carientothrips, 125
carverae, 163
casuarinae, 128

cestosa, 167

ceylonicus, 152

cinctus, 119

cingulatus, 119

citriceps, 136

claripennis, Cryptothrips, 174

claripennis, Scotothrips, 177

Coenurothrips, 148

collaris, 151

corvus, 142

cybele, 139

debilis, 182
denticulatus, 128
dentipes, 119
dentis, 120
Diceratothrips, 133
Dichaetothrips, 134
difficilis, 174
dimidiatus, 167
diversicolor, 152
diver sus, 178
dominicanus, 151
doulli, 171

eastopi, 173
embyotyi, 120

falcatus, 143

fijiensis, Carientothrips, 128

fijiensis, Neosmerinthothrips, 152

firmus, 179

flavitibia, 131

fodinae, 163

formosensis, 162

fructuum, 152

fulvicauda, 143

fulviceps, 144

fumipennis, 144

INDEX

187

fuscicauda, 151
fuscus, 174

Galactothrips, 148
Gastrothrips, 134
gilvipes, 1 20
gloveri, 134
grayi, 129

hambletoni, 138
harti, 181
hawaiiensis, 167
hemidiscus, 164
hilaris, 153
hoodi, 153
Hoplandrothrips, in

icarus, 121
incisus, 131
indicus, 178
inquilinus, 154
insularis, 121
intonsus, 145
ipomoeae, 174
italicus, 122

japomcus, 130
karnyi, 162

laticeps, 167
lativentris, 174
lativerticis, 181
leveri, 175
Liothrips, 182
litoreus, 120
loisthus, 130

magnetis, 131
magnus, 174
major, 175
malaccae, 164
mandiocae, 145
marshalli, 151
melinus, 166
milleforme, 150
minor, 162
miskoi, 131
mjobergi, 131
mongolicus, 145
monticola, 146

Neosmerinthothrips, 148

Nesidiothrips, 156

Nesothrips, 158

niger, 166

nigra, 166

nigricans, 163

nigrisetis, Neosmerinthothrips, 154

nigrisetis, Rhaebothrips, 175

noumeae, 139

oahuensis, 167
obscuricornis , 167
ocelloides, 170
oeceticola, 145
Oedemothrips, 158
oleriae, 167

pallipes, 121
parvidens, 154
paulistarum, 154
pedicillus, 132
Phacothrips, 170
picticornis, 155
pictilis, 132
plaumanni, 155
pratensis, 122
Probolothrips, 134
procerus, 147
propinquus, 167
proteus, 142

reedi, 132
rhachiphilus, 123
Rhaebothrips, 171
Rhinoceps, 112
rhizophorae, 168
robustus, 155
ruficauda, 147

schafferi, 123
Scotothrips, 177
semiflavus, 168
semirufus, 132
seychellensis, 174
similis, 167
skwarrae, 182
speciosissimus, 182
stygicus, 147
subulatus, 148
Syncerothrips, 181

!88 L. A. MOUND

texanus, 148 varius, 123

trinidadensis, 177 vesper, 133

tuberculatus, 121 xylebori, 156

validipennis, 133 yanchepi, 170

validus, 157 y uasai > J 74

variipes, 156 zondagi, 176

L. A. MOUND, B.Sc., D.I.C., D.T.A.

Department of Entomology
BRITISH MUSEUM NATURAL HISTORY
CROMWELL ROAD
LONDON SW7 5BD

ENTOMOLOGY SUPPLEMENTS

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3.i5-

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palocera). Pp. 322: 348 text-figures. August, 1967. 8.

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82 text-figures. May, 1968. 4.

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its Islands. Pp. 198: i plate, 331 text-figures. July, 1969. 4.75.

15. ELIOT, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera:
Nymphalidae). Pp. 155: 3 plates, 101 text-figures. September, 1969. 4.

16. GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe
(Hymenoptera: Chalcidoidea) . Pp. 908: 686 text-figures. November, 1969.

19-

17. WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198:

68 plates, 15 text-figures. October, 1971. 12.

18. SANDS, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae).
Pp. 244: 9 plates, 661 text-figures. July, 1972. 9.90.

19. CROSSKEY, R. W. A Revisionary Classification of the Rutiliini (Diptera:
Tachinidae), with keys to the described species. Pp. 167: 109 text-figures.
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20. VON HAYEK, C. M. F. A Reclassification of the Subfamily Agrypninae
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Pp. 221: 95 text-figures. December, 1973. 9-55-

PRINTED BY Un win Brothers Limited THE GRESHAM PRESS OLD WOKING SURREY ENGLAND

LIBRARY

A REVIEW OF THE

OXYINAE
(ORTHOPTERA : ACRIDOIDEA)

D. HOLLIS

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 31 No. 6

LONDON : 1975

A REVIEW OF THE SUBFAMILY OXYINA!

(ORTHOPTERA : ACRIDOIDEA) V ^T"*s^

BY

DAVID HOLLIS
k,

Pp. 189-234; 69 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 31 No. 6

LONDON : 1975

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 Vol. 31 No. 6 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. not. Hist. (Ent.)

Trustees of the British Museum (Natural History), 1975

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 13 January, 1975 Price 2-75

A REVIEW OF THE SUBFAMILY OXYINAE
(ORTHOPTERA : ACRIDOIDEA)

By D. HOLLIS

CONTENTS

Page

SYNOPSIS. ........... 191

INTRODUCTION .......... 191

ACKNOWLEDGEMENTS ......... 192

HISTORICAL ........ 193

EVALUATION OF PRESENT CLASSIFICATION AND CHARACTERS USED . 194

SUBFAMILY OXYINAE ......... 198

Key to the genera of Oxyinae ....... 200

Generic definitions and lists of included species . . . 201

REFERENCES ........... 229

INDEX ............ 232

SYNOPSIS

The Old World Acridid subfamily Oxyinae is reviewed and redefined. A key is given to the
21 genera now included in the subfamily. Each genus is defined and a list is given of its
included species. Of the 169 described species in the subfamily, 125 are regarded as valid.
125 primary types, including those of the genus Oxya, have been examined and as a result n
new generic synonyms, two new specific synonyms and 25 new combinations are proposed.
One new genus is described; one generic name, one specific name and three combinations are
revived.

INTRODUCTION

DURING a recent study on the genus Oxya (Hollis, 1971) it became clear that (i) the
genus Oxya did not represent a monophyletic entity, (ii) the derivations of the
various species-groups involved in Oxya were obscure, and (iii) the current definition
and scope of the subfamily Oxyinae (Dirsh, 1961) needed some form of revision
and rearrangement. At that time it was thought practical to publish the results
of the study on Oxya separately (Hollis, 1971) considering the genus as a convenient
entity but pointing out that some of its species-groups should be transferred to
other genera and that study was continuing on the whole subfamily. The results
of this study are presented below in review form as it was found impractical, both
in terms of time and available information, to make a complete systematic revision
of the group.

This review is based on a study of the extensive collections of Oxyinae housed
in the British Museum (Natural History), London; the Museum fur Naturkunde
der Humboldt-Universitat, Berlin; the Institute Espanol de Entomologia, Madrid;

jg 2 D. HOLLIS

and the collection of the late C. Willemse. Type and other material was borrowed
from institutions in Basle, Eberswalde, Geneva, Leiden, Oxford, Paris and
Stockholm.

Drawings of the parts of the male phallic complex were made after extraction
and maceration in 10 per cent, potassium hydroxide solution, and the abbreviations
used in these figures are based on those used in Hollis (1971). Drawings of other
features were made from dry material.

Type-depositories of species included in the subfamily are given in abbreviated
form as follows.

NM, Basle Naturhistorisches Museum, Basle.

MNHU, Berlin Museum fur Naturkunde der Humboldt-Universitat, Berlin.

BMNH British Museum (Natural History), London.

IRSNB, Brussels Institut Royal des Sciences Naturelles de Belgique, Brussels.

UZM, Copenhagen Universitetets Zoologiske Museum, Copenhagen.

DEI, Eberswalde Deutsches Entomologisches Institute, Eberswalde, D.D.R.

MHN, Geneva Museum d'Histoire Naturelle, Geneva.

MCSN, Genoa Museo Civico di Storia Naturale, Genoa.

RNH, Leiden Rijksmuseum van Natuurlijke Historic, Leiden.

ZI, Leningrad Zoological Institute, Academy of Sciences of U.S.S.R., Leningrad.

IEE, Madrid Institute Espanol de Entomologia, Madrid.

UM, Oxford University Museum, Oxford.

MNHN, Paris Museum National d'Histoire Naturelle, Paris.

ANS, Philadelphia Academy of Natural Sciences, Philadelphia.

ZIHU, Sapporo Zoological Institute, Hokkaido University, Sapporo.

NR, Stockholm Naturhistoriska Rijksmuseum, Stockholm.

MRAC, Tervuren Musee Royal de 1'Afrique Centrale, Tervuren.

ZIUU, Uppsala Zoologiska Institutonen, Uppsala.

NM, Vienna Naturhistorisches Museum, Vienna.

USNM, Washington U.S. National Museum, Washington.

coll. Willemse Dr Fer Willemse, Eygelshoven, Laurastraat 67, Netherlands

ACKNOWLEDGEMENTS

I would like to express my gratitude to the Centre for Overseas Pest Research
for supporting this work and providing funds for visits to museums in Berlin and
Madrid. These visits were made most productive and pleasant by the help and
hospitality of Dr K. K. Giinther, of the Museum fur Naturkunde der Humboldt-
Universitat, Berlin, and Senora Vincente Llorente, of the Institute Espanol de
Entomologia, Madrid. I am indebted to the following colleagues for loans of
type and other material: Dr C. Baroni-Urbani, Basle; Dr M. Descamps and
Dr M. Donskoff, Paris; Dr B. Hauser, Geneva; Dr A. Kaltenbach, Vienna; Dr T.
Kronestedt, Stockholm; Dr B. Petersen, Eberswalde, D.D.R; Dr H. Weidner,
Hamburg; and Dr Fer Willemse, Eygelshoven, Netherlands.

This is an opportune moment for me to thank Dr L. L. Mishchenko, of the
Academy of Sciences, Leningrad, for his help in lending type-material of the genus
Oxya and to apologize for not acknowledging his valuable help towards the
production of my previous paper (Hollis, 1971).

REVIEW OF THE OXYIN^E
HISTORICAL

193

Although first named by Brunner (1893 : 136) as the 'Oxyae', the group was
recognised earlier by Stal (1878 : 45) as 'part i of Division 12' in his key to
Orthopteran genera. I. Bolivar (1918) redefined the group as 'Section Oxyae'
of the family Acrididae; Willemse (1921) listed the genera he considered to belong
to the group Oxyae of the subfamily Cyrtacanthacrinae occurring in the austro-
oriental region; Rehn (1957), in his study of the Australasian Acrididae, gave the

TABLE i

Genera included by various authors in the Oxyinae

Genus Stal

Brunner Bolivar

Willemse Rehn

Dirsh

(1878)

(1893) (1918)

(1921) (195?)

(1961)

Quilta y

y y

y

y

Gesonia

(now Gesonula) y

y y

y y

Oxya y

y y

y y

y

Caryanda y

y y

y

y

Digentia y

y y

y

Bermius -J

y y

y y

Hieroglyphus y

y y

y

Tauchira y

y y

y

Racilia y

y y

y

y

jBadis/ica

y y

y

//z0roc0ry;tr

(now Parahieroglyphus)

y

y

Chitaura

y

y

y

Per

y

y

Genditia

y

y

Carydana

y

Gerista

y

y

Tauchiridea

y

y

Maga

y

y

Cledra

(now Cercma)

y

y

Gemwda

y

y

Pododula

y

Theomolpus

y

y y

Bermiella

y as

Macroquilta y

Beymioides

y

y y

Racilidea

y

y

Austeniella

y

Caledonia

(now Caledonula)

y

Dapperia

y

y

Dibastica

y

y

Lucretilis

y

Pterotiltus

y

Zulua as Oxya

y

Brachycercus

y

Tolgadia

y

194 D. HOLLIS

group tribal status. Dirsh (1956 : 253) defined the group Oxyae of the subfamily
Catantopinae using, in addition to external characters, some male phallic complex
structures, and the same author (Dirsh, 1961 : 400) gave the group its current
definition and status as a sub-family.

The earlier authors, viz. Stal, Brunner and Bolivar, all used a similar assemblage
of characters to define the group, and these in essence were: fastigium of vertex
horizontal or subhorizontal, little produced forward and forming a rounded angle
with the frons; frontal ridge sulcate and continuous to clypeus; prosternal process
conical or transverse but if latter then narrow at base; mesosternal lobes not
contiguous; fore wing without transverse parallel stridulatory veinlets; lower
genicular lobe of hind femur usually produced posteriorly into a spine, sometimes,
angular, rarely rounded; hind tibia with small spines, outer apical spine almost
always present; male subgenital plate conical, produced, with rounded or subacute
apex. Willemse and Rehn added nothing significant to this definition. Dirsh
(1961) used basically this definition adding to it the form of the male epiphallus,
i.e. bridge-shaped with divided or partially divided bridge, with ancorae and lophi;
and the flexured form of the penis valves in the phallic complex. The genera
included in these various authors' groups are given in Table i. It is noticeable
from this table that Dirsh included far fewer genera in the group than did Bolivar,
principally due to the placing of Bermius, Gesonula, Hieroglyphus, Parahieroglyphus
and Tauchira into his subfamily Hemiacridinae, and others presumably into the
subfamily Cantantopinae.

EVALUATION OF PRESENT CLASSIFICATION AND CHARACTERS USED

In his revision of the higher groups of Acridoidea Dirsh (1961 : 358) states
that he regards a subfamily 'as a group of genera with one or more convenient
characters or a combination of characters, which do not normally occur in other
groups of genera of the family, but are not exclusive . . . ', and on this basis divides
the family Acrididae into 17 subfamilies including the Hemiacridinae, Oxyinae
and Catantopinae. It is clear from his key (Dirsh, 1961 : 389) that the
Hemiacridinae are separated from the Oxyinae on the basis of the former having
transverse parallel stridulatory veinlets on the tegmen and divided penis valves,
and the latter having no such stridulatory mechanism, flexured penis valves and
the lower genicular lobe of the hind femur spine-like posteriorly. The Catantopinae
are defined by a series of negatives as a dumping ground for all those genera which
cannot be placed into any of the other 16 subfamilies.

In studying the Oxyinae it is obvious that the limits of the group become very
vague and merge with some genera in the Hemiacridinae. This latter group
is probably polyphyletic and badly needs reviewing. Dirsh's classification of
the Acrididae on an arbitrary basis of convenient characters, although representing
some progress and providing at least a library classification of the group, has no
strong phylogenetic basis, and in terms of predictive value, is often misleading.
Having an existing classification of the acridids one is either forced to fit genera
into its framework or to make a complete revision of the whole family; one cannot,

REVIEW OF THE OXYINAE 195

piecemeal upgrade or downgrade individual groups within the classification. I
regard the erection and definition of acridid subfamilies as highly ambitious given
the present state of our knowledge of the group. All one can say about the Acrididae
at present is that, apart from a few groups including Coptacridinae, Calliptaminae,
Euryphyminae, the Cyrtacanthacridinae and the Acridinae-Truxalinae complex,
it is a large assemblage of genera in which certain groupings are recognizable
but the affinities of a large proportion of the genera are obscure.

Ideally a classification should consist of monophyletic groupings which are
based on uniquely derived characters. To judge whether or not any one character
or group of characters be uniquely derived it is important to understand the develop-
ment and function particularly in relation to behaviour of such characters.

In general apart from a few pest species, acridids are very poorly studied as live
animals and any assessment of the characters used to define the Oxyinae both
previously and in the present paper can only be surmise. Some observations on
these characters are given below.

EPIPHALLUS (Text-figs 13, 14, 20-35, 38-69). Dirsh (1956 : 233) reasonably
concludes that this structure is used to depress and grasp the female subgenital
plate and fix the phallus firmly during copulation; the ancorae, when present,
or the anterior edge of the epiphallus achieve the fixing by either pressing against
the male supra-anal plate or the dorsal surface of the phallus. In the Oxyinae
the epiphallus may or may not have ancorae, also it is normally bridge-shaped
and almost always completely divided. This division is interesting as it is
probably a secondary development and may allow the epiphallus to be used in
more than one plane, perhaps facilitating a method of copulation different from
the normal acridid method. In the genus Stolzia this division appears much more
significant as the two halves of the epiphallus are not symmetrical (Text-figs 31-35)-
In the genus Oxyina the epiphallus is almost shield-like (Text-figs 45-47) and
although the main structure is divided, the epiphallic membrane is sclerotized
above the dorsal margin of the epiphallus, giving the structure rigidity in one plane,
a return to the normal acridid condition but by a derived method. Although
considered here as a feature of the Oxyinae, a divided epiphallus also occurs in
some Hemiacridinae, Coptacridinae, Catantopinae, all the Euryphyminae and in
some of the South American group of genera related to Cornops and Tetrataenia,
the Leptismae.

FLEXURED OR DIVIDED PENIS VALVES (Text-figs 15-19). Dirsh (1961) states
that the South American group Chilacridinae and the Hemiacridinae are the only
subfamilies in the Acrididae in which the genera have divided penis valves. To
these must be added the group Leptismae. In some other groups the flexure is
very weak and difficult to see.

Dirsh (1962) also describes some Hemiacridinae genera from Madagascar with
flexured penis valves. Clearly this character is of doubtful diagnostic value.
Presumably the penis valves are used to compress the spermatophore sac and

1 96

D. HOLLIS

FIGS 1-12. Oxyinae genera, morphological aspects. Cercina obtusa Stal, i, head and
pronotum, lateral view; Cylindrotiltus versicolor Ramme, 2, head and pronotum, lateral
view; Quilta oryzae Uvarov, 3, head, lateral view ; 4, same, dorsal view ; 1 1 , apex of hind femur,
lateral view; Ochlandriphagaxanthelytrana^liller, 5, head, dorsal view; Chitaura brachyptera
I. Bolivar, 6, head, dorsal view; Oxya agavisa Tsai, 7, female ovipositor, lateral view;
Gesonula mundata (Walker), 8, female ovipositor, lateral view; Thanmoia gustavi Ramme,
9, female ovipositor, lateral view; Oxya chinensis (Thunberg), 10, apex of hind femur,
lateral view; Hygracris palustris Uvarov, 12, apex of hind femur, lateral view.

REVIEW OF THE OXYINAE 197

apically as an intromittant organ. The significance of divided or flexured valves
is unknown. In all Oxyinae genera the valves are flexured.

GENICULAR SPINE ON HIND FEMUR (Text-figs 10-12). The function of this
structure is obscure although it forms an interesting continuation of the spines
on the hind tibia and possibly has a defensive function. A similar structure is
common in the Eumastacids and is present in some Hemiacridinae and the South
American group of genera, the Leptismae. The significance of a spined lower
genicular lobe is not understood but it is notable that, in the genera Quilta and
Hygracris, where the upper genicular lobe is extended this is also terminated in a
spine. In other acridid groups where this genicular extension occurs, e.g.
Acanthoxia, Cannula, Amphicremna, the lobes are rounded or angular and not at
all spine-like.

OUTER APICAL SPINE OF HIND TIBIA. This structure seems to be just a
continuation of the normal tibial spines. It is hardly ever present in the supposedly
more highly evolved Acridids and in the Oxyinae its presence is variable. In
some genera, e.g. Oxya, it is always present and obvious, in some it is weak, in
some, e.g. Gerista, it is absent, but in some, e.g. Thanmoia, it may be present or
absent, even to the extent that a specimen may have the spine on one side and
absent on the other. Possibly this structure is a primitive feature which, at the
Oxyinae stage of development of the Acrididae, is in the process of being lost.

EXPANSION AND FLATTENING OF HIND TIBIA. This is a feature obviously
developed as an adaptation to life in marshy habitats. It has arisen several
times in unrelated genera, e.g. Oxya, Paulinia, Jasomenia and Paracinema, and
cannot be regarded as a uniquely derived character. However, it does seem
to be a feature more or less developed in almost all Oxyinae genera, with the
exception of Pterotiltus where, in all probability, it is secondarily lost.

VENTRAL ABDOMINAL HAIR TUFTS. Again this feature seems to be associated
with life in wet habitats and probably provides buoyancy by trapping air when
the grasshopper jumps into water.

OVIPOSITOR VALVES (Text-figs 7-9, 36, 37). Obviously the type of substrate
in which the animal deposits its eggs has had a direct effect, through selection,
on the form of the ovipositor valves. In the Oxyinae the ovipositor valves are
usually slender, somewhat fragile structures with or without teeth or spines. In
Gesonula, Thanmoia and Stolzia they have quite complicated forms and obviously
oviposition sites in these genera are restricted and specialized. It has been found
convenient to exclude genera from the Oxyinae which have the short, curved,
unspined, robust ovipositor valves found in most Acrididae.

PROSTERNAL PROCESS. Uvarov (1966 : 164) in summary suggests that this
structure is an important taxonomic character, has tactile functions and is parti-

198 D. HOLLIS

cularly important to climbing species. In the Oxyinae it is either a simple conical
structure or is transverse with a narrow base and a widened, often trilobate apex.
Both shapes are common in the Acrididae.

TRANSVERSE PARALLEL STRIDULATORY VEINLETS ON TEGMEN. Dirsh (1956 : 255)
used this character to define the subfamily Hemiacridinae on external features.
After examining a number of genera in which this structure occurs I suspect that
it is a stridulatory mechanism which has evolved more than once in related groups
and although generally useful in suggesting affinities with the Hemiacridinae it
should not be used exclusively for that group. This character does occur in a
few genera of Oxyinae, e.g. Tauchira, Gesonula, Per.

In conclusion it is clear that the characters used to define the subfamily Oxyinae
cannot be definitely regarded as uniquely derived, their state of apomorphy or
plesiomorphy is still not clear, and no one character is exclusive to the subfamily.
As the present paper bases the group on slight modifications of these characters
it can be argued that little or no progress has been made and my criticisms above
of the previous classifications are invalid. However, some attempt has been made
to rationalize the characters used and in the taxonomic section below many genera
previously grouped in the Catantopinae are now included in the Oxyinae, and some
genera previously considered as Oxyinae are removed to other subfamilies. If
similar studies of the Hemiacridinae, Tropidopolinae and Catantopinae other than the
Catantops group were made a much clearer idea of the value of characters used to
separate the subfamilies would emerge.

Subfamily OXYINAE

Oxyae Brunrier von Wattenwyl, 1893 : 136. Type-genus: Oxya Audinet-Serville.
Oxyinae Brunner von Wattenwyl; Dirsh, 1961 : 400.

DIAGNOSIS. Head normally subconical in profile, rarely elongate conical; fastigium of
vertex normally short, rounded triangular or pentagonal from above, rarely elongate and then
only when head elongate (Text-figs 1-6) ; frontal ridge sulcate, usually complete to clypeus ;
facial carinae usually present and complete, only incomplete or absent when frontal ridge is
incomplete.

Dorsum of pronotum cylindrical or weakly flattened; median carinae linear, very weak and
often incomplete, lateral carinae absent; prosternal process conical or transverse and if latter
then always with narrow base; mesosternal interspace open and usually longer than wide.
Tegmina and hind wings fully developed, reduced or rarely absent, tegmen sometimes with a
row of transverse parallel stridulatory veinlets in the radial area. Tympanum always present.
Lower genicular lobe of hind femur produced posteriorly into a spine. Hind tibia usually
expanded in apical half or third, with acute upper margins; external apical spine of hind tibia
present or absent.

Distal sternites of abdomen with tufts of short hairs.

Male epiphallus almost always with completely divided bridge, rarely incompletely divided,
bridge usually narrow, ancorae present or absent, at least one pair of lophi present. Cingular

REVIEW OF THE

199

valves fused apically usually forming a curved plate which partially or completely covers
apical penis valves (Text-figs 16-19); cingulum usually produced slightly postero-dorsally
over valvular plate of cingulum; basal and apical penis valves flexured.

Female ovipositor valves almost always slender and fairly long, often with serrate margins
and if curved then only apically so (Text-figs 7-9, 36, 37).

13

01

FIGS 13-19. Oxya species, male genitalia. O. glabra (Ramme), 13, epiphallus; O. hyla
Audinet-Serville, 14, epiphallus; 15, phallic complex, entire, lateral view; 16, same, dorsal
view, epiphallus and epiphallic membrane removed; 17, endophallus, lateral view;
1 8, same, dorsal view; 19, same, ventral view.

200 D. HOLLIS

DISCUSSION. The subfamily is Old World in distribution and includes the
following genera: Oxya, Caryanda, Thanmoia, Bumacris, Stolzia, Badistica, Gerista,
Pterotiltus, Digentia, Cylindrotiltus, Cercina, Per, Gesonula, Quilta, Hygracris,
Oxyina, Lucretilis, Oxytauchira, Tauchira, Chitaura and Ochlandriphaga.

The genera Gerunda and Racilidea do not have spined genicular lobes of the
hind femora or expanded hind tibiae and are not considered as Oxyinae. They
are probably related to Binulacris, Paramaga and Pseudogerunda, but this whole
group needs to be revised as there is probably some synonymy both at generic and
specific levels.

The Australasian genera, viz. Bermius, Bermiella, Bermioides, Caledonulv,
Dapperia, Koscuiscula, Laxabilla, Methiola, M ethiolopsis , Praxibulus, Tauchiridea
and Tolgadia, represent an interesting complex intermediate between the
Hemiacridinae and the Oxyinae. As none of these genera completely fit the
definition of the subfamily given above they are not considered here. I am avoiding
the issue of their exact position within the Acrididae as very little material was
available for study (although Dr Key of C.S.I.R.O., Canberra was kind enough to
send me examples of the type-species of some of the genera concerned) and further
the Australasian fauna in this group is so poorly recorded there may be presently
undescribed but known material to consider. On balance I suspect these genera are
more closely related to Hieroglyphus and Spathosternum. Under the present
classification, therefore, they would be placed in the Hemiacridinae.

The South American group Leptismae present another interesting facet as,
externally, they appear to be very closely related to the Oxyinae. Most have well
developed spines on the lower genicular lobes of the hind femur and expanded
hind tibiae. However, the male cercus is a peculiar shape, very reminiscent of
the African subfamily Eurphyminae, and the penis valves are clearly divided.
The superficial resemblance between the groups is considered here as the result
of convergent evolution in unrelated groups.

KEY TO THE GENERA OF OXYINAE

1 Prosternal process with apex broadened and transverse, often trilobate . . 2

Prosternal process simple, conical with subacute or rounded apex, rarely with
bulbous apex ............ 6

2 Fully winged or brachypterous species, folded tegmina always touching in mid
dorsal line. ............. 3

Micropterous species ........... 5

3 Integument rugulose; hind femora, particularly along carinae, with many tubercles;
frontal ridge evanescing well before clypeus. .... LUCRETILIS (p. 206)

Integument without tubercles; frontal ridge extending down to clypeus. . . 4

4 Tegmen without trace of transverse parallel stridulatory veinlets; external apical

spine of hind tibia present. OXYTAUCHIRA (p. 204)

Tegmen with transverse parallel stridulatory veinlets in radial area; external apical

spine of hind tibia usually absent. ..... TAUCHIRA (p. 201)

5 Fastigium of vertex, from above, rounded triangular, as long as or almost as long as

wide (Text-fig. 6) ; pronotum smooth CHITAURA (p. 204)

- Fastigium of vertex, from above, rounded pentagonal, much wider than long,
without median longitudinal carina (Text-fig. 5) ; pronotum rugulose.

OCHLANDRIPHAGA (p. 207)

REVIEW OF THE OXYIN^ 201

6 Radial area of tegmen with transverse parallel stridulatory veinlets. ... 7

- Tegmen without trace of transverse parallel stridulatory veinlets. ... 8

7 Fastigium of vertex with median longitudinal carinula; male cercus simple, conical;

female ovipositor valves long, slender. ...... FER (p. 208)

- Fastigium of vertex without median longitudinal carinula; male cercus curved

apically; female ovipositor valves short, stubby (Text-fig. 8) . GESONULA (p. 209)

8 Both upper and lower genicular lobes of hind femur spined (Text-figs n, 12) . 9
Only lower genicular lobes of hind femur spined (Text-fig. 10) . . . 10

9 Slender species; head elongate conical, fastigium of vertex almost as long as

longest diameter of eye (Text-figs 3, 4) . . . . . QUILT A (p. 210)

- Robust species ; head subconical, fastigium of vertex wider than long

HYGRACRIS(p. 210)

10 Male epiphallus with ancorae (Text-fig. 20) . . . . . . . n

Male epiphallus without ancorae (Text-fig. 13). . . . . . . . 16

11 Epiphallus strongly asymmetrical (Text-fig. 31) . . . STOLZIA (p. 211)

Epiphallus symmetrical (Text-fig. 20) . . . . . . . . 12

12 Epiphallus plate-like (Text-figs 45-47) OXYINA (p. 228)

Epiphallus normal, bridge-shaped (Text-fig. 20). . . . . . . 13

13 External apical spine of hind tibia present. . . . . . . . 14

External apical spine of hind tibia absent . . . . . . . .15

14 Epiphallus with one pair of outer lophi and two pairs of inner lophi (Text-fig. 40);

fastigium of vertex with median longitudinal carinula, although sometimes weak

and difficult to see GARY AND A (p. 217)

- Epiphallus with one pair of outer lophi and none or only one pair of inner lophi

(Text-figs 48-53) ; fastigium of vertex without median longitudinal carinula.

BADISTICA (p. 220)

15 African species; epiphallus without inner lophi. .... GERISTA (p. 219)
Oriental species; epiphallus with an inner pair of lophi . . THANMOIA (p. 207)

1 6 Fully winged or brachypterous species . . . . . . . 17

- Micropterous species. ........... 18

17 Inner lophi of epiphallus absent, outer pair long and hook-like (Text-fig. 57).

BUMACRIS (p. 222)

Epiphallus with pair of inner lophi, outer pair short, boot-shaped (Text-fig. 14).

OXYA (p. 220)

18 Hind tibia rounded and not expanded apically. . . . PTEROTILTUS (p. 226)

- Hind tibia expanded in apical half or third, with upper outer margins acute. . 19

19 Face and whole pronotum rugulose; antenna with median flagellar segments at

least three times as long as broad; epiphallus incompletely divided (Text-figs

61, 62) DIGENTIA (p. 223)

- Face, pronotum and mesonotum smooth ; antenna with median flagellar segments at

most twice as long as broad ; epiphallus completely divided (Text-fig. 60) . . . 20

20 African species; dorsum of pronotum cylindrical and deeply crossed by transverse

sulci (Text-fig. 2) CYLINDROTILTUS (p. 225)

Ceylonese species; dorsum of pronotum flattened and only shallowly crossed by

transverse sulci (Text-fig, i) CERCINA (p. 224)

GENERIC DEFINITIONS AND LISTS OF INCLUDED SPECIES

Tauchira Stal, 1878

(Text-figs 24, 25)

Tauchira Stal, 1878 : 48. Type-species: Oxya polychroa Stal, designated by Kirby, 1910 : 397.
Bua I. Bolivar, 1918 : 31. Type-species: Tauchira buae I. Bolivar, by monotypy. [Synony-
mised by Willemse, 1930 : 131!.

202 D. HOLLIS

Paratraulia Willemse, 19250 : 355. Type-species: Traulia obliquiannulata Brunner von
Wattenwyl, by original designation. [Synonymised by Ramme, 1941 : 114.]

Servillenia Ramme, 1941 : 121. Type-species: Acridium abbreviatum Audinet-Serville, by
monotypy. Syn. n.

DIAGNOSIS. Head conical; fastigium of vertex parabolic, wider than long, median
longitudinal carinula weakly present; frontal ridge sulcate, complete to clypeus, lateral facial
keels complete. Eyes normal or slightly bulbous. Antenna longer than combined lengths of
head and pronotum. Prosternal process transverse, antero-posteriorly flattened, apex
trilobate. Dorsum of pronotum rugulose or subrugulose, slightly flattened, median carina
weak, lateral carinae absent, crossed by three transverse sulci; relative width of mesosternal
interspace variable. Tegmina and hind wings fully developed or shortened to brachypterous
condition; parallel transverse stridulatory veinlets present, sometimes weakly, in ist radial
or medial area. Lower genicular lobe of hind femur spined ; hind tibia very weakly expanded
apically, with acute upper margins; external apical spine of hind tibia absent. In male loth
abdominal tergite with a weak furcula on postero-dorsal margin; epiphallus with narrow,
divided bridge and well developed ancorae. Female ovipositor valves long, evenly toothed.

DISTRIBUTION. China, W. Malaysia, W. Indonesia.

SPECIES INCLUDED

Tauchira abbreviata (Audinet-Serville) comb. n.

Acridium abbreviatum Audinet-Serville, 1839 : 678. Holotype $, JAVA (lost). Neotype $,
JAVA (MNHM, Berlin), designated by Ramme (1941 : 123) [examined].

T. polychroa (Stal)

Oxya polychroa Stal, 1875 : 32. Holotype $, MALAYA (Malacca) (NR, Stockholm).
Paratraulia bifasciata Willemse, 19250 : 357, fig. Holotype $, MALAYA (BMNH) [examined].
[Synonymized by Willemse, 1930 : 190 (footnote).]

T. bn ac I. Bolivar

Tauchira buae I. Bolivar, 1898 : 92. Syntypes <$ and $, SUMATRA (MCSN, Genoa; IEE,
Madrid) [^ examined].

T. obliquiannulata (Brunner von Wattenwyl)

Traulia obliquiannulata Brunner von Wattenwyl, 1898 : 249. Holotype $, SUMATRA
(? depository).

T. rufotibialis (Willemse)

Paratraulia rufotibialis Willemse, 19250 : 358. Holotype $, SINGAPORE (BMNH) [examined].
Paratraulia elegantula Willemse, 1930 : 190, fig. 98. Syntypes <$ and $, SUMATRA (RNH,
Leiden). [Synonymised by Willemse, 1956 : 202].

T. karnyi (Willemse)
Paratraulia karnyi Willemse, 19250 : 359. Holotype <$, MALAYA (BMNH) [examined].

T. grandiceps (Willemse)
Paratraulia grandiceps Willemse, 1928 : 9. Holotype $, SUMATRA (RNH, Leiden).

T. gressitti Tinkham

Tauchira gressitti Tinkham, 1940 : 304, pi. 10, fig. 4, pi. 13, figs 2, 2a. Holotype <$, CHINA
(Hainan) (? depository).

DISCUSSION. The neotype male of T. abbreviata differs from other species of
Traulia in that the dorsum of the pronotum is slightly less rugulose and flattened

REVIEW OF THE

203

and the parallel stridulatory veinlets of the tegmen are weaker but the latter is
probably due to the shortening of the tegmen. These slight differences are not
considered significant enough to warrant separate generic status for the species
and Servillenia is synonymised with Tauchira.

FIGS 20-30. Oxyinae genera, male epiphalli. 20, Thanmoia olivacea (Willemse); 21,
T. maculata (Willemse); 22, T. gustavi Ramme; 23, Fer coeruleipennis Willemse; 24,
Tauchira polychroa (Stal) ; 25, T. abbreviatum (Audinet-Serville) ; 26, Chitaura mirabilis
(Carl); 27, C. lucida (Krauss); 28, C. indica Uvarov; 29, C. maculata (Willemse); 30,
Oxytauchira aurora (Brunner von Wattenwyl).

204 D. HOLLIS

Oxytauchira Ramme, 1941

(Text-fig. 30)
Oxytauchira Ramme, 1941 : 117. Type-species: Tauchira gracilis Willemse, by monotypy.

DIAGNOSIS. Head conical; fastigium of vertex, from above, parabolic, about as wide as
long, median longitudinal carinula weakly present; frontal ridge sulcate, complete to clypeus;
lateral facial keels complete. Eyes normal. Antenna about as long as combined lengths of
head and pronotum. Prosternal process transverse, antero-posteriorly flattened, trilobate
apically. Dorsum of pronotum weakly flattened, median carina weak, lateral carinae absent,
shallowly crossed by three transverse sulci; mesosternal interspace longer than wide. Tegmen
and hind wing fully developed, no trace of transverse parallel stridulatory veinlets on tegmen.
Lower genicular lobe of hind femur spined ; hind tibia hardly expanded apically but with acute
upper margins, external apical spine of hind tibia present. In male loth abdominal tergite
with a furcula on postero-dorsal margin; epiphallus with narrow divided bridge and developed
ancorae. Female ovipositor valves long, slender, unevenly spined.

DISTRIBUTION. Burma, Sulawesi.

SPECIES INCLUDED

Oxytauchira gracilia (Willemse)

Tauchira gracilis Willemse, 19310 : 242, fig. 99. Syntypes 2 $, SULAWESI (Central) (NM,
Basle) [i $ examined].

O. aurora (Brunner von Wattenwyl) comb. n.

Racilia aurora Brunner von Wattenwyl, 1893 : J 55. pi- 5, fig- 53- Holotype , BURMA (MCSN,
Genoa) .

DISCUSSION. The type of 0. aurora was unavailable for study and the combination
suggested here is based on a study of material in the BMNH collections thought
to be this species. 0. aurora is clearly not congeneric with other species of Racilia
and the genus is returned to the Catantopinae and is thought to be related to
Gerunda.

Chitaura I. Bolivar, 1918
(Text-figs 6, 26-29)

Chitaura I. Bolivar, 1918 : 32. Type-species: Chitaura brachyptera I. Bolivar, designated

by Willemse, 19310 : 244.
Tauracris Willemse, 19310 : 248. Type-species: Tauracris flavolineata Willemse, by monotypy.

[Synonymised by Ramme, 1941 : 135.]
Baliacris Willemse, 1938 : 116. Type-species: Baliacris maculata Willemse, by monotypy.

Syn. n.
Chitaurella Ramme, 1941 : 136. Type-species: Tauchira lucida Krauss, by original designation.

Syn. n.

DIAGNOSIS. Head conical; fastigium of vertex, from above, wider than long, pentagonal
or rounded triangular, median longitudinal carinula weakly present; frontal ridge sulcate,

REVIEW OF THE OXYIN^ 205

complete to clypeus; lateral facial keels complete. Eyes normal; antenna about as long as
or slightly longer than combined lengths of head and pronotum. Prosternal process antero-
posteriorly flattened, trilobate apically. Dorsum of pronotum subcylindrical, median carina
weak, lateral carinae absent, shallowly crossed by three transverse sulci ; mesosternal interspace
as long as or slightly longer than wide. Tegmina and hind wings reduced to micropterous
condition, scale-like, tegmen extending to posterior margin of first abdominal sclerite. Lower
genicular lobe of hind femur spined; hind tibia slightly expanded apically with acute upper
margins; external apical spine of hind tibia small but usually present. In male loth abdominal
tergite with furcula on hind margin; epiphallus with divided bridge and clearly defined
ancorae. Female ovipositor valves long, slender, serrated.

DISTRIBUTION. S. India, W. Indonesia, Sulawesi, Maluku.

SPECIES INCLUDED

Chitaura lucida (Krauss) comb. rev.
Tauchira lucida Krauss, 1903 : 760, pi. 67, fig. 7. Syntypes $ and $, JAVA (? Jena Museum).

C. mirabilis (Carl)

Tauchira mirabilis Carl, 1916 : 472. Syntypes <$ and $, SULAWESI (Central) (MHN, Geneva)
[examined] .

C. satnanga (Carl)

Tauchira samanga Carl, 1916 : 474. Holotype ^, SULAWESI (Central) (MHN, Geneva)
[examined].

C. vidua (Carl)
Tauchira vidua Carl, 1916 : 475. Holotype $, SULAWESI (Central) (MHN, Geneva) [examined].

C. brachyptera I. Bolivar

Chitaura brachyptera I. Bolivar, 1918 : 33. Holotype $, SULAWESI (North) (IEE, Madrid)
[examined].

C. indica Uvarov
Chitaura (?) indica Uvarov, 1929 : 553. Holotype <$, INDIA (South) (BMNH) [examined].

C.flavolineata (Willemse) comb. n.

Tauracris flavolineata Willemse, 19310 : 248, figs 102, 103. Holotype $, SULAWESI (North)
(NM, Basle) [examined].

C. maculata (Willemse) comb. n.
Baliacris maculata Willemse, 1938 : 117. Holotype (j, BALI I. (coll. Willemse) [examined].

C. striata Willemse
Chitaura striata Willemse, 1938 : 117. Holotype $, KEY Is. (RNH, Leiden).

C. elegans Ramme

Chitaura elegans Ramme, 1941 : 128. Holotype $, SULAWESI (South) (MNHU, Berlin)
[examined] .

C. at rat a Ramme

Chitaura atrata Ramme, 1941 : 129. Holotype $, SULAWESI (Central) (MNHU, Berlin)
[examined].

C. mengkoka Ramme

Chitaura mengkoka Ramme, 1941 : 131. Holotype <$, SULAWESI (South) (MNHU, Berlin)
[examined].

C. ochracea Ramme

Chitaura ochracea Ramme, 1941 : 132. Holotype <$, SULAWESI (South) (MNHU, Berlin)
[examined].
**

206 D. HOLLIS

C. moluccensis Ramme
Chitaura moluccensis Ramme, 1941 : 133. Holotype <$, MALUKU (Amboina) (NM, Vienna).

C. poecila Ramme

Chitaura poecila Ramme, 1941 : 131. Holotype <$, SULAWESI (North) (MNHU, Berlin)
[examined] .

C. bivittata Ramme

Chitaura bivittata Ramme, 1941 : 133. Holotype $, MALUKU (Ceram) (MNHU, Berlin)
[examined].

DISCUSSION. Ramme (1941 : 135) first suggested Tauracris may be synonymous
with Chitaura and after examining Willemse's type of T. flavolineata, a female
dried from spirit, this synonymy is confirmed.

The type male of Baliacris maculata lacks the external apical spine on both hind
tibiae but this character appears to be very unstable in Chitaura, often being present
on one hind leg and absent on the other. In general appearance this species is
very similar to C. hicida and, on the evidence from external morphology and the
structure of the male phallic complex, is placed in Chitaura.

C. lucida differs slightly from the bulk of the other species of Chitaura in that
the dorsum of the pronotum is less rounded and the transverse sulci are slightly
deeper, but these features do not warrant separate generic status and it is recombined
with Chitaura.

Lucretilis Stal, 1878

(Text-fig. 55)
Lucretilis Stal, 1878 : 41, 85. Type-species: Lucretilis taeniata Stal, by monotypy.

DIAGNOSIS. Head conical; fastigium forming a stepped join with vertex, pentagonal from
above, wider than long, median longitudinal carinula very weak or absent ; frontal ridge sulcate
but present only down to median ocellus ; lateral facial keels absent or if present then disguised
by rugulosity of facial integument. Eyes bulbous. Antenna very much longer than
combined lengths of head and pronotum. Prosternal process transverse in apical half, trapezoid
in anterior view. Dorsum of pronotum rugulose and coarsely pitted, rounded, shallowly
crossed by three transverse sulci, median carina very weak, lateral carinae absent ; mesosternal
interspace longer than wide. Tegmina and hind wings somewhat reduced, extending to about
half length of abdomen, former without trace of transverse parallel stridulatory veinlets.
Hind femur tuberculate, lower genicular lobe spined; hind tibia very weakly expanded in
apical half, upper margins acute apically; external apical spine of hind tibia present. In
male loth abdominal tergite with or without furcula on postero-dorsal margin; epiphallus
with narrow divided bridge, ancorae and only an outer pair of lophi. Female ovipositor valves
long, slender, completely smooth.

DISTRIBUTION. Malaysia, W. Indonesia.

SPECIES INCLUDED

Lucretilis taeniata Stal
Lucretilis taeniata Stal, 1878 : 85. Holotype $, SUMATRA (NM, Vienna).

L. antennata I. Bolivar
Lucretilis antennata I. Bolivar, 1898 : 88. Holotype , SUMATRA (IEE, Madrid) [examined].

REVIEW OF THE OXYIN^ 207

L. bolivari Miller

Lucretilis bolivari Miller, 1934 : 531, fig. i, pi. 13, fig. 3. Holotype $, MALAYA (BMNH)
[examined] .

L. uvarovi Miller

Lucretilis uvarovi Miller, 1935 : 695, fig. 5, pi. 14, fig. 5. Holotype <$, MALAYA (BMNH)
[examined].

L. maculata Willemse
Lucretilis maculata Willemse, 1936 : 202. Holotype <$, BORNEO (Central) (RNH, Leiden).

L. splendens Willemse

Lucretilis splendens Willemse, 1938 : 118, fig. i. Holotype <$, BORNEO (North) (UM, Oxford)
[topotypic material examined].

L. dohrni Ramme

Lucretilis dohrni Ramme, 1941 : 98, pi. 13, fig. 8. Holotype $, SUMATRA (MNHU, Berlin)
[examined].

L. jucunda Miller
Lucretilis jucunda Miller, 1953 : 142, fig. i. Holotype C J, MALAYA (BMNH) [examined].

Ochlandriphaga Henry, 1933
(Text-figs 5, 58)

Ochlandriphaga Henry, 1933 : T %7- Type-species: Ochlandriphaga xanthelytrana Henry, by
monotypy.

DIAGNOSIS. Head conical; fastigium of vertex forming stepped join with vertex, from
above pentangular, without median longitudinal carinula ; frontal ridge weakly sulcate, complete
to clypeus; lateral facial keels weak and incomplete. Eyes bulbous. Antenna much longer
than combined lengths of head and pronotum. Prosternal process antero-posteriorly flattened,
with weakly trilobate apex. Dorsum of pronotum rounded, coarsely pitted, moderately deeply
crossed by three transverse sulci, median carina very weak, lateral carinae absent; mesosternal
interspace slightly longer than wide. Tegmina and hind wings reduced, scale-like, former
not extending beyond middle of 3rd abdominal tergite. Lower genicular lobe of hind femur
spined; hind tibia hardly expanded apically but dorsal margins acute in apical third; external
apical spine of hind tibia present. In male loth abdominal tergite without furcula; epiphallus
with narrow, divided bridge, without ancorae and only with outer pair of lophi. Female
ovipositor valves long, slender, evenly toothed.

DISTRIBUTION. Ceylon.

SPECIES INCLUDED

Ochlandriphaga xan