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BULLETIN OF 

THE BRITISH MUSEUM 

NATURAL HISTORY 



ENTOMOLOGY 

Vol. XII 

1962 




PRINTED BY ORDER OF THE TRUSTEES OF 

THE BRITISH MUSEUM (NATURAL HISTORY; 

LONDON : 1963 



DATES OF PUBLICATION OF THE PARTS 

No. I - 17 April 1962 

No. 2-17 April 1962 

No. 3-29 May 1962 

No. 4-17 July 1962 

No. 5-17 July 1962 

No. 6-17 July 1962 

No. 7-10 August 1962 

No. 8-26 October 1962 

Contents & Index — January 1963 



PRINTED IN 

GREAT BRITAIN 

AT THE 

BARTHOLOMEW PRESS 

DORKING 

BY 

ADLARD AND SON, LTD. 




CONTENTS 

ENTOMOLOGY VOLUME XII 

PAGE 

No. I. The British Pseudococcidae (Homoptera : Coccoidea). By D. J. 

Williams i 

No. 2. New African Caddis-flies (Order Trichoptera) . B3' D. E. Kimmins 81 

No. 3. Notes on the EpitoMnae with descriptions of a new genus and new 
species and subspecies (Lepidoptera : Lycaenidae). By T. H. E. 
Jackson 123 

No. 4. A revision of the African Laemophioeinae (Coleoptera : Cucujidae). 

By L. P. Lefkovitch 165 

No. 5. Microlepidoptera from the New Hebrides. Records and descriptions 
of Microlepidoptera collected on the Island of Aneityum by Miss 
Evelyn Cheesman, O.B.E. By J. D. Bradley 247 

No. 6. The Acridoidea (Orthoptera) of Madagascar. I. Acrididae (except 

Acridinae). By V. M. Dirsh 273 

No. 7. Notes on Tunga caecigena (Siphonaptera : Tungidae). B3' Karl 
Jordan. Neotunga euloidea gen. n., sp. n. (Siphonaptera : Pulici- 
dae). By F. G. A. M. Smit 351 

No, 8. Holarctic sawflies (Hymenoptera : Symphyta). By Robert B. 

Benson 379 



THE BRITISH 

PSEUDOCOCCIDAE 

(HOMOPTERA : COCCOIDEA) 



D. J. WILLIAMS 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 12 No. i 

LONDON: 1962 



THE BRITISH PSEUDOCOCCIDAE 
(HOMOPTERA : COCCOIDEA) 



BY 



D. J. WILLIAMS 



(Commonwealth Institute of Entomology) ^J 



}is 




pp. 1-79 ; 29 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol 12 No. i 

LONDON: 1962 



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. 

This paper is Vol. 12, No. i of the Entomological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 



© Trustees of the British Museum, 1962 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued 17 April 1962 Price Twenty-eight Shillings 



THE BRITISH PSEUDOCOCCIDAE 
(HOMOPTERA : COCCOIDEA) 

By D. J. WILLIAMS 

SYNOPSIS 

This study of the British Pseudococcidae has necessitated changes in nomenclature of many of 
the 42 species discussed, resulting in new combinations and new synonymy of names. The species 
are placed in 21 genera, one of which, Balanococcus, is described as new. Twenty-eight species 
are regarded as native and the remainder are found in greenhouses. Four new species are 
described : Atrococcus cracens, Balanococcus boratynskii, Saccharicoccus penium and Trionytnus 
Orestes. Illustrations are given of 29 species, the others having been illustrated elsewhere by 
other authors. 

INTRODUCTION 

The British Pseudococcidae or mealy-bugs discussed here were included in the sub- 
family Dactylopiinae by Newstead (1903) and later by Green in various accounts of 
the British species. The group is equivalent to the tribe Pseudococcini as defined by 
Balachowsky (1948) or to the subfamily Pseudococcinae recognized by Borchsenius 
(1949). In the present work the group is recognized as that defined by Ferris (1950) 
as the family Pseudococcidae. 

HISTORY 

Very few mealy-bugs were known from Britain in the last century and about half 
of these had been described from greenhouses. One in particular, collected in a green- 
house at the Royal Botanic Gardens, Kew, has still not been found in its natural 
habitat. The first native record seems to be that of Coccus halophilus described by 
J. Hardy in 1868. Towards the end of the century J. Douglas added other names to 
the British Hst but it was R. Newstead who first collected this group seriously. In 
a monograph of British coccids, Newstead (1903) gave descriptions of 12 mealy-bugs, 
7 of which were native species. 

During the present century E. E. Green was the one most active in this field adding 
many new records and describing 15 British species as new to science. 

The inadequacy of existing descriptions of British Pseudococcidae makes it impos- 
sible, except in a few instances, to identify species from the literature with any 
degree of certainty. This also applies to many species described from the Continent, 
although in recent years some effort has been made to describe them more fully. 

The purpose of this study is to redescribe and illustrate the British Pseudococcidae 
and to bring together the somewhat scattered records. Particular attention has been 
given to a work by Borchsenius (1949) revising the species from Russia. In this 
work 80 are described as new and many others have since been described from that 
area. Many of the Russian species will doubtless be found in Western Europe and, 

ENTOM. 12, I. I 



4 D. J. WILLIAMS 

taking as a guide the frequency of new species recently discovered in Western Russia, 
it would seem safe to assume that many new ones remain to be discovered in Western 
Europe and even in Britain. 

When some of the continental species are known to be identical with those 
described here then their names have been reduced to synonyms. On the other hand 
a number of the British names may have to be sunk when the continental species are 
redescribed. 

Holotypes of the four new species are deposited in the British Museum (Natural 
History). 

GEOGRAPHICAL DISTRIBUTION 

The species settled on for discussion are given in the distributional list opposite. 

Newstead and Green often recorded species as British merely because they had 
been collected alive on imported plants. In recent years Ferrisiana virgata (Cockerell) 
and Dysmicoccus brevipes (Cockerell) , two important cosmopolitan species, have found 
their way into this country but no records of their establishment are known here. 
A few species, especially some members of the genera Pseudococcus and Rhizoecus, 
have become established in greenhouses throughout the country and for purposes of 
recognition they are included. Some other greenhouse species are discussed where 
changes in nomenclature are due or where a redescription will save the species from 
obscurity. 

No original material seems to exist of Dactylopius theobromae described by Douglas 
(1889) and it is impossible to recognize it from the meagre description : the name, 
therefore, must be regarded as a nomen dubium. 

Although normally regarded as part of the continental fauna, the mealy-bugs of 
the Channel Islands are included because, of the 9 species now recorded, all but 3 are 
known to occur in Britain. The paucity of records from Wales, Scotland and Ireland 
immediately becomes apparent from the list. This is undoubtedly due to a lack of 
collecting and when the total number of British species is finally known, these areas 
will probably have the most interesting and varied fauna. The British records 
included relate only to specimens actually studied during the course of this work. 



MORPHOLOGY 

All morphological terms mentioned are based on those used by Ferris (1950, 1953). 
McKenzie (i960) has given an excellent generalized drawing representing the morpho- 
logical structures of the family Pseudococcidae. Measurements of the labium refer 
to the two fused and sclerotized segments constituting the cone. The small basal 
segment is not included. As pointed out by McKenzie, the presence of a denticle on 
the claw, although a useful character, is now of less significance in placing a species in 
the Phenacoccus series. Some species are now known with a denticle but which are 
not typical of this series and others have been described recently with an obvious 
relationship to the Phenacoccus series, yet lack the denticle. One species is described 
here as Euripersia europaea (Newstead) from specimens where the denticle is present 
or entirely lacking. 



THE BRITISH PSEUDOCOCCID AE 



Distributional List of the British Pseudococcidae 



Antonina crawi* 

Atrococcus cracens sp. n. . 
A. luffi .... 

A. paludinus . 
Balanococcus horatynskii sp. n. 

B. scirpi 

Chnaurococcus subterraneus 
Chorizococcus lonnshuryi* 



Dysmicoccus walkeri 

D. wistariae* . 
Euripersia europaea 

E. tomlinii 
Heliococcus minutus 
Heterococcus pulverarius 
Nipaecoccus nipae* . 
Peliococcus balteatus . 
Phenacoccus aceris . 
P. interruptus 

P. sphagni 
Planococcus citri* 
Pseudococcus adonidum* 
P. fragilis 
P. latipes* 
Rhizoecus albidus 
R. cacticans* . 

R. dianthi* 
R. elongatus* . 
R. falcifer* 
R. halophilus . 
Saccharicoccus penium sp. 
Spilococcus cactearum* 

S. filicicola* . 
Spinococcus calluneti 



Trionymus dactylis 
T. diminutus . 



T. newsteadi . 
T. Orestes sp. n. 
T. perrisii 
T. phalaridis 
T. radicum 
T. thulensis 
T. tomlini 



Id 




Ti 




^^ 




c 




c 


-d 


^ 45 








Q 


§ 


«3 rt 


Other localities 




$ 


^ 


(—1 


f{ In 




X 




X 






Japan, Formosa, U.S.A., Hawaii 
France, Russia. 



X 
X 
X 
X 
X 

X 
X 
X 
X 

X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 

X 
X 
X 
X 
X 
X 

X 
X 



X X 



S. Africa, Kenya, Egypt, Hawaii, 

U.S.A. Holland. 
Spain, France, Germany. 
Japan, U.S.A. 
Holland, Germany. 
Corsica, Germany. 
Spain. 

Probably cosmopolitan. 
Russia, Sweden, Czechoslovakia. 
Palaearctic and Nearctic. 

Holland. 
Cosmopolitan. 
Cosmopolitan. 
Cosmopolitan. 
Probably cosmopolitan. 
France, Germany. 
North and South America, green- 
houses in Germany. 

Holland. 

Probably cosmopolitan. 



U.S.A., Mexico, greenhouses in 
Europe. 

Sweden, Germany, Russia, 

Czechoslovakia . 
France. 
New Zealand, U.S.A., Russia, 

Formosa, Italy. 
Germany, Russia, Czechoslovakia. 

France, Germany , Russia, Holland. 

Russia. 

Holland. 

Iceland, France. 



* Greenhouse species. 



6 D. J. WILLIAMS 

ACKNOWLEDGEMENTS 

The writer is grateful to Dr. Harold Morrison of the United States Department of 
Agriculture, Washington, D.C. for the prompt and generous way he has answered 
queries, studied specimens submitted to him and given helpful advice. His highly 
valued opinions on many of the species herein discussed are accepted and included. 

Professor H. L. McKenzie of the University of California, Davis, California has 
co-operated fully in discussing many points of interest and giving the writer valuable 
information on his latest paper on the Californian mealy-bugs before it was published. 
His kindness in this respect has been most welcome. 

Professor N. S. Borchsenius of the Zoological Institute, Academy of Science of 
the U.S.S.R., Leningrad, has kindly examined a photograph of the illustration of 
Balanococcus scirpi (Green) and given valuable comments on its relationship to the 
genus Kiritshenkella described by him. He also compared specimens of Trionymus 
hibernicus with Russian specimens of T. perrisii for which the writer is deeply 
indebted. 

Dr. A. Reyne of the Zoologisch Museum, Amsterdam, Holland, kindly made 
available Dutch specimens of Trionymus perrisii and T. radicum for which thanks 
are cordially given. 

Particular gratitude is expressed to Dr. K. L. Boratynski of Imperial College, 
London, who has taken a lively interest in this study since its commencement. Dr. 
Boratynski has made available his collection of British mealy-bugs, adding many 
new records and two new species which are described herein. His help and encourage- 
ment have been much appreciated. 

Key to Genera 

1 Legs entirely lacking ....... ANTONINA Signoret 

- Legs present, although at times reduced in size ...... 2 

2 Trilocular pores present somewhere on the body ...... 3 

- TrUocular pores absent, replaced by quinquelocular pores HETEROCOCCUS Ferris 

3 (2) With some crateriform ducts present, each with the orifice at the apex of a 

sclerotized prominence and often with one or more setae attached to the base 

of the duct prominence HELIOCOCCUS SUI9 

- Without this type of duct .......... 4 

4 (3) With some dorsal body setae spine-like and same size or nearly same size as 

those of cerarii ........... 5 

- Dorsal body setae never enlarged and spine-like, either slender or minutely 

lanceolate ............ 6 

5 (4) Quinquelocular disc pores present on venter, claw with a denticle, dorsal cerarii 

present 5P7NOCOCCf/5 Borchsenius 

- Quinquelocular disc pores absent, claw without a denticle, dorsal spine-like 

setae not forming cerarii ..... NIPAECOCCUS SUI9 

6 (4) Dorsal multilocular disc pores present, arranged in clusters of 3-6 and with a 

minute tubular duct at the centre . . . PELIOCOCCUS Borchsenius 

- Dorsal multilocular disc pores, if present, not arranged in clusters . . 7 

7 (6) Oral rim ducts present somewhere on the body ...... 8 

- Oral rim ducts lacking ..........12 

8 (7) With 17 pairs of cerarii, each possessing slender auxiliary setae in addition to 

cerarian setae PSEUDOCOCCUS Westwood 



THE BRITISH PSEUDOCOCCID AE 7 

"With 0-17 pairs of cerarii but only anal lobe pair with auxiliary setae . . 9 

Body content black or blue-black, especially apparent when boiling in potash or 

lacto-phenol ............ u 

Body content never black or blue-black, usually yellow . . . . 10 

With from o to 4 pairs of cerarii present CHORIZOCOCCUS McKenzie 

With from 6 to 17 pairs of cerarii present, part of . SPILOCOCCUS Ferris 

With 1-7 pairs of cerarii, circulus absent . . . ATROCOCCUS Goux 

With at least 9 pairs of cerarii, circulus present, part of .SPILOCOCCUS Ferris 

Circulus present, longer than wide, hour-glass shaped SACCHARICOCCUS Ferris 
Circulus, if present, never longer than wide or hour-glass shaped . . . 13 

With a denticle or tooth, either well developed or small, on the plantar surface of 

the claw ............ 14 

Claw without a denticle .......... 15 

With 9-18 pairs of cerarii .... PHENACOCCUS Cockerell 

With 2-4 pairs of cerarii, part of .... EURIPERSIA Borchsenius 

Anal lobes without recognizable cerarii, bi- or tritubular pores present somewhere 

on the body RHIZOECUS Kiinckel d'Herculais 

Anal lobes with cerarii represented by paired conical setae, bi- or tritubular 

pores lacking ........... 16 

With 18 pairs of cerarii PLANOCOCCUS Ferris 

With 1-17 pairs of cerarii ......... 17 

With 6-17 pairs of cerarii DYSMICOCCUS Ferris 

With 1-5 pairs of cerarii .......... 18 

With some quinquelocular pores on venter, part of EURIPERSIA Borchsenius 

Never with quinquelocular pores on venter . , . . . . . 19 

Antennae 7-8-segmented TRIONYMUS Berg 

Antennae 6-segmented .......... 20 

Multilocular disc pores forming marginal zone around dorsum and venter 

BALANOCOCCUS WUliams 
— Multilocular disc pores on venter of abdomen only, not forming marginal zone 

CHNAUROCOCCUS Ferris 

Notes. Although the key is based mainly on morphological characters some 
difficulty may be encountered with couplet 9 (8) when identifying specimens already 
prepared on slides. This is due to our incomplete knowledge of the genera Chorizo- 
coccus, Spilococcus and Atrococcus and further discussions are given under these 
genera. 

DESCRIPTIONS OF SPECIES 

ANTONINA Signoret 
Antonina Signoret, 1875 : 24. 
Antonina Signoret, Borchsenius, 1949 : 311. 
Antonina Signoret, Ferris, 1953 : 289. 

Type of genus Antonina purpurea Signoret. 

The type of the genus was described from France but the genus is represented 
mainly in the warmer regions. Only Antonina crawi Cockerell is known in this 
country and this is found in greenhouses. It is mentioned here to synonymize the 
name A . socialis Newstead. All members of this genus are devoid of legs and many 
have the posterior segments of the abdomen sclerotized at maturity. They are found 
only on grasses. 



9 


(8) 


10 


(9) 


II 


(10) 


12 


(7) 


13 


(12) 


14 


(13) 


15 


(13) 


16 


(15) 


17 (16) 


18 


(17) 


19 


(18) 


20 


(19) 



8 D. J. WILLIAMS 

Antonina crawi Cockerell 

Anionina crawi Cockerell, 1900 : 70. 
Antonina socialis Newstead, 1901 : 85 (syn. n.). 
Antonina socialis Newstead, Newstead, 1903 : 207. 
Antonina crawii Cockerell, Ferris, 1953 : 292. 

The type of Antonina socialis is identical with A. crawi. Specimens are at hand 
from England, Hertfordshire, Broxbourne, Arundinaria japonica, 26.1.1899 and 
Scotland, East Lothian, Prestonkirk, on bamboo, ii.1905, all taken under glass. 
Ferris has redescribed and illustrated this species. 

ATROCOCCUS Goux 

Atrococcus Goux, 1941 : 69. 

Type of genus Atrococcus melanovirens Goux. 

Goux erected this genus for a few species possessing the following characters : 
cerarii on abdomen only, circulus absent, oral rim ducts on dorsum and usually on 
lateral areas of venter, presence of a group of oral collar ducts on prothorax in front 
of anterior spiracles, these often accompanied by a group of multilocular disc pores, 
body content showing a black pigment after death and especially when placed in 
potash. 

The black or blue-black body content in mealy-bugs is often a preliminary aid 
in identification and certainly the character has some significance in such genera 
as Nipaecoccus §ul9, Naiacoccus Green and Amonostherium Morrison & Morrison and 
others. There seem to be no special external morphological characters associated 
with the internal body colour. Once a specimen is mounted on the slide it is, so far, 
impossible to determine the original body colour. 

Two genera, Spilococcus Ferris and Chorizococcus McKenzie, come very close to 
Atrococcus. The genus Chorizococcus has 0-4 pairs of cerarii and Spilococcus has 
6-17 pairs ; these are the only distinguishing characters. The number of cerarii in 
Atrococcus is 1-7 pairs, combining the characters of the other two genera and, indeed, 
Chorizococcus hrevicruris McKenzie with 2 pairs of cerarii and with a noticeable 
lateral group of tubular ducts and multilocular disc pores on the prothorax could 
easily be placed in Atrococcus except for the absence of the black colour after death. 
For the purposes of this work the three genera are regarded as distinct but the 
discovery or redescription of other species in these genera may clarify the position. 

The following species, including one from the Channel Islands, may be separated 
by the key : 

1 Cerarii on anal lobes only ; oral rim ducts numerous, about 20 on each tergite 

luffi (Newstead) 

- Cerarii 3-7 pairs ; oral rim ducts less numerous, about 6 to each tergite ... 2 

2 Cerarii 3-4 pairs ; noticeable groups of multilocular disc pores in submedian areas of 

dorsum ; number of multilocular disc pores on prothorax equals or exceeds that of 
tubular ducts .......... cracens sp. n. 

- Cerarii 6-7 pairs ; dorsal multilocular disc pores on margins only of posterior abdominal 

segments ; multilocular disc pores on prothorax always less in number than 
tubular ducts ......... paludinus (Green) 



THE BRITISH PSEUDOCOCCID A E 9 

Atrococcus cracens sp. n. 

(Text-fig. i) 

Pseudococcus paludinus Green, Green, 1934 : m- (Misidentification.) 

Habit and Distribution. External appearance not known although Green has 
recorded it as forming ovisacs in the angles of prominent leaf veins. Material has 
been examined from the following localities : England. Surrey : Guildford, 
Centaurea nigra, 21.vii.1922 (E. E. Green). Kent: Bearsted, Chrysanthemum 
leucanthemum, 16. ix. 1932 (E. E. Green). Hampshire: Yateley, Crepis sp., 25. ix. 
1926 (E. E. Green). Berkshire : Silwood Park, Achillea sp., 27. vi. 1949, (Holotype) 
Deschampsia caespitosa, 27.x. 1948, Veronica sp., 31.viii.1948 (K. L. Boratynski). 
Wales. Flintshire: Prestatyn, grass, viii.1918, R. Newstead. 

Recognition characters. An ovoid-elliptical species about 2-5 mm. long. Antennae 
8-segmented, 355-365 microns long. Legs well developed, slender, posterior coxae with a few 
translucent pores. Trochanter + femur 225-240 microns, tibia + tarsus 240-260 microns. 
Labium 95 microns long. Circulus absent. Ostioles present with about 8 trilocular pores and 
I or 2 setae on each lip. Anal ring with 6 setae in length about twice diameter of ring. Cerarii 
on last 3-4 segments of abdomen only ; anal lobe cerarii each with a pair of slender conical setae 
and a loose group of trilocular pores accompanied by 3-4 auxiliary setae and surrounded by a 
small area of sclerotization. Anterior cerarii similar but cerarian .setae smaller and without 
auxiliary setae. 

Dorsal setae not numerous, all short and slender. Multilocular disc pores in small submarginal 
groups on the abdominal segments anterior to anal lobes and in noticeable submedian groups 
on all segments including head except segments IX + X. Tubular ducts with oral collar of two 
sizes, present in small numbers around the abdominal margins and associated with the submar- 
ginal groups of multilocular disc pores. Tubular ducts with oral rim arranged in more or less 
single transverse rows, a common number being about 8. Trilocular pores present. 

Ventral surface with a small elongate area of sclerotization on each anal lobe terminating in an 
apical seta longer than anal ring setae. Other ventral setae slender but longer than those on 
dorsum. Multilocular disc pores numerous in double or triple rows at posterior edges of segments 
V-VIII and at anterior edges of segments VII and VIII, numerous on segments IX -f X and 
in a marginal group containing 15-27 pores opposite each anterior coxa ; others present between 
each pair of coxae. Tubular ducts with oral collar of same two sizes as on dorsum, a larger type, 
numerous in submarginal groups on the posterior abdominal segment, these often extending to 
the submedian areas ; there being also about 10-15 within each group of multilocular disc pores 
on prothorax. A smaller type of duct situated mainly in a transverse row on each of posterior 
abdominal segments. Tubular ducts with oral rim arranged in pairs around the margins of 
prevulvar abdominal segments and in numbers of 1-4 opposite the spiracles. Trilocular pores 
not numerous. 

Notes. This species may be easily distinguished from others in the genus by the 
presence of dorsal groups of multilocular disc pores. It comes closest to A. paludinus 
(Green) from which it differs in possessing fewer cerarii and in the larger proportion 
of multilocular disc pores to tubular ducts opposite the first coxae. 

The accompanying illustration has been prepared from material kindly made 
available by Dr. K. L. Boratynski. 



D. J. WILLIAMS 




Fig. I. Airococcus cracens sp. n. 



THE BRITISH PSEUDOCOCCID AE II 

Atrococcus luffi (Newstead) (comb, n.) 
(Text-fig. 2) 

Dactylopius luffi Newstead, 1901 : 85. 
Pseudococcus luffi (Newstead), Green, 1925 : 520. 

Habit and distribution. The ovisac is, apparently, rather closely felted, long, 
cylindrical, and of equal width throughout ; female remaining uncovered at the 
cephalic extremity. Newstead did not mention the colour but Green described it as 
pale flesh-colour, thinly dusted with white mealy powder. All of this material is now 
black and the specimens become purple-brown or black in potash. Described 
originally from Channel Islands, Guernsey, on lower stems and roots of Lepigonum 
rupestre, now Spergularia rupicola. Other specimens seen, all from Guernsey, are 
on Silene maritima, Petit Bot, Vazon Bay and on Armeria vulgaris, Houmet Benest. 
This is one of the species described from the Channel Islands which so far has not 
been collected in Britain. 

Recognition characters. Shape ovoid, about 2-5 mm. long, anal lobes poorly develop>ed. 
Antennae 8-segmented, about 415 microns long. Labium 1 15-120 microns long. Legs normal, 
posterior coxae with a few translucent pores ; trochanter -\- femur 245-260 microns, tibia -|- 
tarsus 260-285 microns. Circulus absent. Ostioles well developed with inner edges of lips 
sclerotized and a few trilocular pores and an occasional seta on each lip. Anal ring with 6 setae 
twice length of ring at its greatest diameter. Cerarii represented by a single pair on anal lobes 
only, each with a pair of slender conical setae and about 8 trilocular pores. 

Dorsal setae mainly short and slender, not numerous. Oral rim ducts in more or less single 
transverse rows on the abdominal segments, a common number being about 20 on each segment ; 
on the thorax they tend to be in double transverse rows but anteriorly they become scattered. 
Segments III-VIII with small submarginal groups of multilocular disc pores and tubular ducts 
with oral collar of two sizes, a larger type usually mingled with the disc pores or just posterior 
to them and a smaller type just anterior to the groups. Trilocular pores evenly distributed. 
Simple circular pores, slightly larger than a trilocular pore, present in small numbers. 

Ventral surface of each anal lobe with a small area of sclerotization and an apical seta longer 
than anal ring setae. Ventral setae mainly longer and stouter than those on dorsum. Multi- 
locular disc pores rather numerous in the mid-region of the fourth and posterior segments and 
in submarginal groups on all abdominal segments but these becoming less numerous anteriorly ; 
a noticeable submarginal group of about 12 pores also present opposite each anterior coxa. 
Tubular ducts with oral collar of two sizes and similar to those on dorsum ; a smaller type in 
transverse rows in the middle of the posterior abdominal segments and a few in the submarginal 
areas ; a larger type present just anterior to the multilocular disc pores in the mid-region of 
the abdomen and mingled with the submarginal groups, there being also 3-4 associated with 
the submarginal groups of disc pores on the prothorax. Tubular ducts with oral rim in small 
numbers around the submargins of abdomen and on the pro- and mesothorax where they occupy 
the median areas also. Trilocular pores sparse. Simple circular pores present but not numerous. 

Atrococcus paludinus (Green) 
(Text-fig. 3) 

Pseudococcus paludinus Green, 1921 : 191. 
Atrococcus paludinus (Green), Goux, 1941 : 80. 

Habit and distribution. Described from Wicken Fen, Cambridgeshire, Eng- 
land, on the under surface of the foliage of Eupaiorium cannahinum, Symphytum 



D. J. WILLIAMS 




Fig. 2. Atrococcus luffi (Newstead). 



THE BRITISH PSE UDOCOCCI D AE 



13 



officinale, Urtica sp., Lysimachia sp., Convolvulus sp., and Spiraea sp. This is the 
only correct record as other specimens collected or recorded under this name are 
referred to A. cracens. Green described the insect as " Adult female brownish pink, 
masked (on the dorsum) by a thin covering of white pulverulent secretion. Posterior 
extremity with a pair of short, stout, waxy tassels, with a smaller tassel on the sides 
of each of the two preceding segments. Younger examples are of a pale olivaceous 
colour." 

Recognition characters. An ovoid species attaining a length of 2-5 mm., anal lobes 
moderately developed. Antennae 8-segmented, 395-475 microns long. Legs slender, well 
developed, with translucent pores on hind coxae, trochanter + femur 330-370 microns, tibia + 
tarsus 325-380 microns. Labium 130-140 microns long. Ostioles present, each lip with a few 
trilocular pores and i or 2 setae. Anal ring setae about twice length of diameter of ring. Cerarii 
numbering 6-7 pairs on posterior abdominal segments only ; anal lobe cerarii, each with a pair 
of slender conical setae surrounded by a loose group of trilocular pores and a few auxiliary setae. 
Other cerarii similar but cerarian setae becoming more slender anteriorly and without auxiliary 
setae. 

Dorsal setae short and slender, not numerous. Tubular ducts with oral collar and multilocular 
disc pores in submarginal groups up to segment IV where they become less numerous ; always 
with a larger proportion of ducts to pores ; an occasional multilocular disc pore also present in 
median areas of posterior abdominal segments. Tubular ducts with oral rim on all segments 
except IX + X, in single transverse rows, a common number being about 6. Trilocular pores 
evenly distributed. 

Ventral surface with a small elongate area of sclerotization on each anal lobe and with a stout 
apical seta longer than anal ring setae. Ventral setae sparse, all slender but longer than those on 
dorsum. Multilocular disc pores numerous around margins of second and posterior abdominal 
segments and abundant in transverse rows on fifth and posterior segments ; a few also present 
in mid-region of some anterior segments. Another group of 2-8 always present lateral to first 
coxae, these accompanied by a group of 8-45 tubular ducts with oral collar similar to those on 
dorsum, the number of ducts although variable, always greater than number of multilocular 
disc pores. Other tubular ducts numerous in submarginal groups on second and posterior 
segments ; present also in median areas of some anterior abdominal segments. A smaller type 
of duct, with oral collar, sparse in median areas of posterior abdominal segments and occasionally 
near margins. Tubular ducts with oral rim arranged singly or in pairs around the submargins 
as far anterior as metathorax and a group of 5-6 opposite each first spiracle. Trilocular pores not 
numerous. 

BALANOCOCCUS gen. n. 

Type of genus Riper sia scirpi Green. 

Recognition characters. Pseudococcidae with elongate oval body, anal lobes poorly 
developed. Antennae 6-segmented. Legs normal. Anterior and posterior ostioles present, poorly 
developed. Circuli present or absent, up to 3 small oval circuli present in one species. Anal 
ring with a double band of pores and 6 setae. Cerarii numbering i or 2 pairs on abdomen only. 
Tubular ducts with oral collar on dorsum and venter, in zone around body, these of a distinctive 
shape, the collar flange-shaped and occupying either half or nearly half of total length of duct. 
Multilocular disc pores forming zone around entire body on both dorsum and venter, and present 
in transverse rows on abdomen. Trilocular pores evenly distributed. 

Notes. This genus comes closest to Kiritshenkella Borchsenius but differs mainly 
in having the trilocular pores evenly distributed over the body whereas one of the 



14 



D. J, WILLIAMS 




Fig. 3. Atrococcus paludinus (Green). 



THE BRITISH PSEUDOCOCCI D AE 15 

important characters of Kiritshenkella is the concentration of trilocular pores on the 
mid-hne of both surfaces. Furthermore, there are in Kiritshenkella only posterior 
ostioles which are well developed, whereas Balanococcus possesses 2 pairs and these 
are poorly developed. Although the tubular ducts of both genera are similar, those 
of Kiritshenkella seem to be much flatter and are described as disc-like. 

The writer is much indebted to Professor N. S. Borchsenius for kindly examining a 
photograph of the accompanying illustration of B. scirpi and advising on its differences 
with Kiritshenkella. 

The two species of Balanococcus may be separated by the following key : 

Cerarii present on anal lobes only, circuli absent .... scirpi (Green) 

Cerarii present on anal lobes and segment VIII, circuli 3 in number . boratynskii sp. n 



Balanococcus boratynskii sp. n. 

(Text-fig. 4) 

Habit and distribution. External appearance not known. Found under the 
leaf sheaths of Deschampsia caespitosa, England, Berkshire, Silwood Park, 31 . v . 1945 
(K. L. Boratynski). 

Recognition characters. Body elongate with almost parallel sides, length about 3-5 mm., 
anal lobes moderately developed. Antennae 6-segmented, 260-310 microns long. Legs rather 
small and slender, posterior coxae with a few translucent pores ; trochanter -f femur 190-220 
microns, tibia + tarsus 200-240 microns. Labium 70-75 microns long. Circuli 3 in number, 
small and oval, the posterior circulus much smaller than others. Ostioles poorly developed 
with 1-3 trilocular pores on each lip. Cerarii on anal lobes and segment VIII only, anal lobe 
cerarii, each with a pair of slender conical setae accompanied by 5-6 trilocular pores and a 
single auxiliary seta. Penultimate cerarii usually with a single conical seta and i or 2 trilocular 
pores. Anal ring with 2 rows of pores and 6 setae about twice length of diameter of ring. 

Dorsal surface with rather numerous setae, all short and slender. Trilocular pores with an 
even distribution. Multilocular disc pores in a marginal zone around entire body, more numerous 
at anterior and posterior ends ; present also in transverse rows on segments VI-VIII. Tubular 
ducts short with large oral collar occupying slightly less than half total length of duct. They are 
interspersed with the multilocular disc pores around the margins and form transverse rows on 
abdominal segments ; a few scattered on head. 

Ventral surface with each apical seta slightly shorter than anal ring setae. Ventral setae 
similar to those on dorsum but some on head and posterior segments tending to be longer. 
Trilocular pores evenly distributed. Multilocular disc pores present in a transverse row on 
segment V and abundant in transverse rows on posterior abdominal segments. Abundant also 
in a marginal zone around entire body. Tubular ducts of one size only, similar to dorsal ducts, 
situated in transverse rows on segment V and posterior segments. Present also in a marginal 
zone among the multilocular disc pores and particularly numerous in marginal groups on segment 
VI and posterior segments. 

The writer has much pleasure in naming this species after Dr. K. L. Boratynski 
who kindly made available material for study and who has helped in many other 
ways during the preparation of this work. 



I6 



D. J. WILLIAMS 





^ 



b 




Fig. 4. Balanococcus boratynskii sp. n. 



THE BRITISH PSEUDOCOCCI D AE 17 

Balanococcus scirpi (Green) (comb, n.) 
(Text-fig. 5) 
Ripersia scirpi Green, 192 1 : 192. 

Habit and distribution. This species was described by Green as " Concealed 
at base of stems of Scirpus caespitosa [—Trichophorum caespitosum], in boggy 
ground ; Camberley." [Surrey, England.] The adult is apparently pink coloured 
and concealed beneath a coating of white mealy secretion. 

Recognition characters. Adult female elongate, sides subparallel, widest across fourth 
abdominal segment, length approximately 3-5 mm., posterior end rounded. Antennae 6-seg- 
mented, 150-190 microns long. Legs small and slender, posterior coxae with a few translucent 
pores, trochanter -f femur about 130 microns, tibia + tarsus 105-130 microns. Labium 50-60 
microns long. Anterior and posterior ostioles represented by mere slits without trilocular pores 
or setae on the lips. Circulus absent. Anal ring with 2 rows of pores and 6 setae about twice as 
long as diameter of ring. Anal lobe cerarii present only, each with a pair of conical setae set 
close together and surrounded by about 4 trilocular pores. 

Dorsal surface with small pointed setae, not numerous. Trilocular pores sparse but with an 
even distribution. Multilocular disc pores situated mainly in a narrow zone around the margins, 
the heaviest concentrations on the head and posterior abdominal segments. Others extending to 
median areas of segments VI and VII and occasionally in median areas of some anterior segments. 
Tubular ducts short, oral collar large, occupying about half total length of duct. A few present 
on head margins, a few on margins of fourth and posterior segments and small concentrations on 
segments VII-IX where they also occupy the median areas. 

Ventral surface with apical setae shorter than anal ring setae. Body setae small and sparse on 
each segment but tending to be more numerous towards margins. Trilocular pores in transverse 
groups in median areas. Multilocular disc pores numerous in a submarginal zone around body. 
They are also scattered on head and form transverse rows on segments VII-IX. Tubular ducts, 
similar to those on dorsum, situated around body within zone formed by multilocular disc pores. 
A smaller type of duct but otherwise similar in all respects, confined to median areas of segments 
VII, VIII and IX + X. 

Chnaurococcus Ferris 
Chnaurococcus Ferris, 1950 : 40. 

Type of genus Ripersia villosa Ehrhorn. 

This genus was erected for two rotund species each with 6-segmented antennae 
and with a single pair of cerarii on anal lobes only. The following species seems to 
belong to this genus although like C. trifolii (Forbes), it lacks the dorsal tubular 
ducts of the type species. 

Chnaurococcus subterraneus (Newstead) (comb, n.) 

(Text-fig. 6) 

Ripersia subterranea Newstead, 1893 : 79. 

Ripersia tomlinii Newstead, Newstead, 1903 : 187 (in part). 

Ripersia subterranea Newstead, Newstead, 1903 : 189. 

Ripersia formicarii Newstead, In Donisthorpe, 1907 : 5 (syn. n.). 

Ripersia europaea Newstead, Green, 1921 : 191. (Misidentification.) 

Ripersia europaea Newstead, var., Green, 1926 : 183. 

ENTOM. 12, I. 2 



i8 



D. J. WILLIAMS 



^ 




Fig. 5. Balanococcus scirpi (Green). 



THE BRITISH PSEUDOCOCCID AE 19 

Habit and distribution. Newstead described the insect originally as " female 
adult ; dark red-brown, turns dark purple in caustic potash ". Specimens have been 
examined from the following localities : England. Norfolk : Ingoldisthorpe, King's 
Lynn, on roots of Nardus stricta in nests of Formica flava (types). Dorsetshire : 
Portland, nest of Lasius niger. Cornwall : Pudstone, nests of Lasius niger, 8 . vii . 1920 
(H. Donisthorpe) ; Stepper Point, vii. 1920. Somersetshire: Porlock, nests of 
Lasius niger, v. 1915 (W. C. Crawley) ; Minehead, nest of Lasius niger, ix. 1920 (E. E. 
Green). Devonshire: Dartmouth, nest of Lasjws wt^^y (H. Donisthorpe). Gloucester- 
shire: Bristol, in ant's nest, vii. 1924 (H. Donisthorpe). Sussex: Eastbourne, 
nests of Lasius niger, 6.xi.i924 (H. Donisthorpe). Kent : Charing, with Lasius 
niger (type of R. formicarii). Scotland. East Lothian, North Berwick Law (E. E. 
Green) . 

Recognition characters. A broadly oval species measuring approximately 2 mm. long ; 
anal lobes moderately developed. Antennae 6-segmented, 235-260 microns long. Legs small and 
stout, normal, trochanter -\- femur 180-190 microns, tibia + tarsus 190-200 microns. Labium 
105-115 microns long. Circulus absent. Ostioles well developed, with inner edges of lips sclero- 
tized and each lip with 4-5 trilocular pores and an occasional seta. Anal ring with a double 
band of pores and 6 setae only slightly longer than diameter of ring. Cerarii represented by a 
pair on anal lobes only, each with a pair of short conical setae, these usually straight but sometimes 
curved and surrounded by a loose group of 6-7 trilocular pores. 

Dorsal and ventral setae short and mainly rather stout. Apical setae longer than anal ring 
setae. Trilocular pores somewhat numerous and evenly distributed on both surfaces. Ventral 
surface with multilocular disc pores on fifth and posterior segments only, in single or slightly 
irregular transverse rows at posterior edges of segments. Tubular ducts of one size, on venter 
only, situated in single transverse rows at middle of most abdominal segments and in small 
submarginal groups. 

Notes. This species comes closest to C. trijolii (Forbes), another species found in 
ants' nests and known from U.S.A., but differs in possessing only 6-7 trilocular pores 
with each cerarius whereas in C. trifolii the numbers of trilocular pores are much 
more numerous. 

CHORIZOCOCCUS McKenzie 
Chorizococcus McKenzie, i960 : 692. 

Type of genus Chorizococcus wilkeyi McKenzie. 

This genus was erected for species with oral rim ducts, the cerarii numbering 
from 0-4 pairs without auxiliary setae except those on anal lobes. It differs from 
Spilococcus mainly in possessing fewer cerarii but is almost identical with Atrococcus 
discussed earlier. So far as is known, all species of Spilococcus are yellow or pinkish 
but species of Atrococcus are black, especially after death, and turn a distinct black 
or blue-black in potash. For the purposes of this work the genera are regarded as 
distinct. 

Chorizococcus lounsburyi (Brain) 

Pseudococcus lounsburyi Brain, 1912 : 179. 

Pseudococcus [Trionymus] peregrinus Green, 1925a : 40, 41 (syn. n.). 

Trionyntus lounsburyi (Brain), Ferris, In Zimmerman, 1948 : 260, 261. 



D. J. WILLIAMS 



<3. 



^ 




Fig. 6. Chnaurococcus subterraneus (Newstead). 



THE BRITISH PSEUDOCOCCI D AE 21 

Trionymus lounsburyi (Brain), Ferris, 1950 : 271. 
Pseudococcus lounsburyi Brain, De Lotto, 1958 : 96, 97. 
Chorizococcus peregrinus (Green), McKenzie, i960 : 701. 

This is a difficult species to identify satisfactorily. It has never been figured ade- 
quately from the original type material from South Africa and the only illustration 
available is that prepared by Ferris from material collected in Hawaii. McKenzie 
has considered, on information received from Dr. H. Morrison, that this material 
is the same as that described by Green as Pseudococcus (Trionymus) peregrinus and 
differs from C. lounsburyi. This distinction is based mainly on the differences in 
colour and habit of the adult female. The writer has examined many specimens 
from South Africa, Egypt, Hawaii, Holland and England and, although there is 
variation in the numbers of multilocular disc pores and oral rim ducts, there is little 
evidence that the material represents different species. On a request from the author, 
Dr. H. Morrison has kindly re-examined material at Washington and has come to 
similar conclusions, although he has stated that the results must be regarded as 
tentative until more material is studied. As C. peregrinus comes within the known 
range of variation, the name is sunk as a synonym of C. lounsburyi. The variation 
has also been discussed by De Lotto who has examined many specimens from Africa. 
For the time being the illustration given by Ferris is accepted although the numbers 
of multilocular disc pores and oral rim ducts tend to be rather high. As pointed out 
by De Lotto there are often multilocular disc pores on the posterior dorsal abdominal 
segments. 

Although there are exceptions, most of the records are from the plant families 
Amaryllidaceae and Liliaceae. Material is at hand from England, all under glass, 
from Hampshire, Exbury, Nerine roots {peregrinus type) and Surrey, Abinger 
Hammer, on bulbs of Crinum sp., Sprekelia sp. and Hippeastrum sp. Other material 
has also been examined from England, without locality, from the bulbs of Amaryllis 
sp. 

DYSMICOCCUS Ferris 
Dysmicoccus Ferris, 1950 : 53. 

Type of genus Dactylopius hrevipes Cockerell. 

The writer has accepted the interpretation of this genus given by McKenzie (i960) 
and a further discussion is given under Trionymus. Only two species recorded from 
Britain belong to the genus and these may be separated as follows : 

Body elongate, multilocular disc pores numerous in transverse rows on posterior 

abdominal segments of venter ...... walkeri (Newstead) 

Body rotund, multilocular disc pores few, situated around vulva only wistariae (Green) 

Dysmicoccus walkeri (Newstead) (comb, n.) 
(Text- fig. 7) 

Dactylopius walkeri Newstead, 1891 : 164. 
Dactylopius walkeri Newstead, Newstead, 1903 : 169. 
Pseudococcus walkeri (Newstead), Fernald, 1903 : 112. 
Pseudococcus walkeri (Newstead), Green, 191 6 : 31. 
Pseudococcus walkeri (Newstead), Green, 1925 : 517. 



D. J. WILLIAMS 



"^. 




D 



& 



b 



Fig. 7. Dysmicoccus walkeri (Newstead). 



THE BRITISH PSEUDOCOCCID AE 23 

Pseudococcus walkeri (Newstead), Green, 1926 : 182, 183. 
Pseudococcus walkeri (Newstead), Green, 1928 : 30. 

Habit and distribution. Described as " Elongate brown. Antennae and legs 
pale yellow-brown, covered with a dense white powder ; the cottony appendages of 
the margin stout, six of these at the abdominal extremity longer and stouter than the 
others, the second pair longest and very stout, in length equal to width of body. . . . ". 
This is normally a grass-infesting species and specimens have been examined from 
the following localities : England. Cheshire : Manley, near base of Agrostis tenuis, 
ix.1890 (R. Newstead). Surrey: Camberley, vii.1915, 13. ix. 1920, viii.1922, 
5.ix.i922, viii.1926 {Calluna sp. stems), 22.viii.1939 (E. E. Green) ; Ashtead, 
22.viii.1939. Kent: Bearsted, 14. ix. 1926 (E. E. Green). Somersetshire: Cheddar, 
viii.1926 (E. E. Green). Wales. Brecknockshire: Llangammarch, ix.1925 (E. E. 
Green). Montgomeryshire: Nant Cwmdu Aberhosan, 20.vii.1919 (R. Newstead). 
Scotland. East Lothian : Aberlady, viii-1925 (E. E. Green). Aberdeenshire : 
Monjnnusk, viii.1920 (F. Laing). Channel islands. Herm : Petit Bot. 

Recognition characters. An elongate-oval species measuring up to 4-5 mm. long, anal 
lobes poorly developed. Antennae 8-segmented, 410-515 microns long. Legs normal, trochanter 
+ femur 370-395 microns, tibia + tarsus 375-440 microns. Labium 125-130 microns long. 
Circulus not large, rounded. Ostioles with 3-4 trilocular pores and 2-3 setae on each lip. Anal 
ring with 6 setae twice as long as its diameter. Cerarii numbering 15-17 pairs, usually 15 pairs, 
those lacking being on the head ; anal lobe cerarii each with a pair of conical setae and numerous 
trilocular pores accompanied by about 10 auxiliary setae and all borne on a distinct area of 
sclerotization, larger in area than anal ring. Anterior cerarii each with smaller cerarian setae, a 
few trilocular pores and one or two auxiliary setae. 

Dorsal setae of various sizes but mainly short and slender. One or two multilocular disc pores 
usually present on penultimate segment. Tubular ducts with oral collar of 2 sizes, a larger type 
numerous on all segments and a smaller type sparse on some of the posterior abdominal segments. 
Trilocular pores evenly distributed. 

Ventral surface with a small area of sclerotization on each anal lobe terminating at a stout 
apical seta longer than anal ring setae. Ventral setae rather numerous, all slender but mainly 
longer than those on dorsum. Multilocular disc pores numerous on posterior abdominal segments, 
there being double or triple rows on the posterior edges of segments VI-VIII and on the anterior 
edges of segments VII and VIII ; numerous also on segments IX + X, a few also present 
behind first coxae, occasionally at random on thorax. Tubular ducts of same sizes as on dorsum, 
a larger type on fourth and posterior segments and numerous on margins of posterior abdominal 
segments ; smaller marginal groups present on thorax and head. A smaller type of duct in median 
areas of fourth and posterior segments only, sparse. Trilocular pores present. 

Notes. This species comes close to D. timherlakei (Cockerell), another elongate 
species on grasses described from U.S.A. and with 17 pairs of cerarii but in this case 
the circulus is moderately large and divided by a distinct intersegmental furrow. 
In D. walkeri the circulus is oval and not divided. 

Dystnicoccus wistariae (Green) (comb, n.) 

Pseudococcus wistariae Green, 1923 : 218. 
Pseudococcus piricola Siraiwa, 1935 : 69 (syn. n.). 
Pseudococcus cuspidatae Rau, 1937 • ^95 (syn. n.). 
Dysmicoccus cuspidatae (Rau), Ferris, 1950 : 61. 
Dystnicoccus piricola (Siraiwa), Takahashi, 1957 : 3. 



24 D. J. WILLIAMS 

This species was described from material collected on Wistaria sp. at a Japanese 
nursery garden, presumably under glass, at St. Albans, Hertfordshire, England. 
Dr. Harold Morrison has kindly examined a specimen and he agrees that it is identical 
with D. cuspidatae described from U.S.A. The latter name has already been synony- 
mized by Takahashi with D. piricola described from Japan. Ferris has redescribed 
and illustrated this species under D. cuspidatae. 

EURIPERSIA Borchsenius 
Euripersia Borchsenius, 1948 : 955. 

Type of genus Euripersia amnicola Borchsenius. 

Two British species are included in this genus on the basis of the descriptions of 
E. amnicola Borchs. and E. brevispina recently described by Borchsenius & Ter- 
Gregorian (1956). The essential features of the genus are the oval body ; antennae 
6-7-segmented ; claw with a denticle although at times absent ; quinquelocular 
disc pores usually present on venter although in the type species they are present 
only in the first stage larva ; cerarii numbering at most 3 pairs, 2 of which confined 
to segments VHI and IX ; circuli numbering from 1-3 or absent entirely ; multi- 
locular disc pores confined to ventral abdominal segments. 

Although the two following species were described originally in the genus Ripersia, 
they are excluded from this genus despite certain similarities to Ripersia corynephori 
Signoret, the type species, redescribed by Reyne (1951). This species has 8-segmented 
antennae, 2 pairs of cerarii, multilocular disc pores on both dorsum and venter, 
quinquelocular disc pores absent although present in the first stage larva. As very 
little else is known about this species and as the following species have no multilocular 
disc pores on the dorsum, the genus Ripersia is disregarded from the present study. 

The two British species have been mentioned in the literature under a variety of 
names and consequently only the original references can be trusted. 

Euripersia europaea (Newstead) (comb, n.) 
(Text-fig. 8) 

Ripersia europaea Newstead, 1897 : 167. 

Ripersia tomlinii Newstead, Donisthorpe, 1907 : 5. (Misidentification.) 

Ripersia tomlinii Newstead, Green, 1920 : 122. (Misidentification.) 

Ripersia tomlinii Newstead, Green, 1925 : 518 (in part). 

Ripersia wiinni Reyne, 1953 : 235-239 (syn. n.). 

Habit and distribution. Newstead described this species as " densely clothed 
with white mealy wax ; segmentation more or less distinct ". Its most favoured 
positions are apparently under stones in ants' nests and they presumably feed on 
the fine rootlets. Specimens are known from Britain as follows : Channel Islands. 
Guernsey : Watville, ants' nests (B. Tomlin), {europaea types) ; Fort Doyle, under 
stones with ants, 9 . ix . 1924 (E. E. Green) ; 29 . xii . 1926, 8 . vi . 1947 (R. H. Le Pelley) ; 
also material without data from other localities in nests of Lasius niger. England. 
Isle of Wight : Blackgang, with Lasius flavus and L. niger (H. Donisthorpe) ; San- 
down, nest of Lasius niger, 4.V.1910 (H. Donisthorpe) ; Ventnor, 26. ix. 1922 (H. 



THE BRITIH PSEUDOCOCCID AE 



25 



<S 




Fia 8. Euripersia europaea (Newstead). 



26 D. J. WILLIAMS 

Donisthorpe). Kent : Dover, nest of Lasius niger (H. Donisthorpe) . Sussex : 
Eastbourne, with Lasius niger, 6.ix.i924 (H. Donisthorpe). Dorsetshire : Swanage, 
14. ix, 1904 (B. Tomhn). Cornwall: Whitsand Bay, with Lasius niger (H. Donis- 
thorpe). 

Recognition characters. A small broadly oval species approximately 2-5 mm. long, anal 
lobes poorly developed. Antennae 6-segmented, 225-260 microns long. Legs small, claw with 
or without a denticle, in some cases only the faintest sign of a denticle or may be present or 
absent on different claws of same specimen. Trochanter + femur 155-165 microns, tibia + tarsus 
170-205 microns. Labium 90-95 microns long. Circulus absent. Ostioles present, each lip with 
about 4 trilocular pores but apparently without setae. Anal ring with a double band of pores 
and 6 setae about same length as ring at its greatest diameter. Cerarii numbering 2-4 pairs ; 
anal lobe cerarii each with 2 slender conical setae and about 5 trilocular pores, the space between 
the setae lightly sclerotized ; penultimate cerarii similar but cerarian setae slightly smaller. 
The preocular cerarii often present, although difficult to see. Their presence can be detected by 
the paired setae and the small concentration of trilocular pores. Another pair of cerarii sometimes 
present on thorax. 

Dorsal setae sparse, all small and slender. The only pores present on the dorsum are trilocular, 
rather numerous with an even distribution. 

Ventral surface with a stout pair of apical setae longer than anal ring setae. Ventral setae not 
numerous, all slender and short but tending to be longer than dorsal setae especially on posterior 
segments. Multilocular disc pores at anterior and posterior edges of segments VII and VIII and 
present on segments IX + X, there being scarcely more than 50 altogether. Quinquelocular 
disc pores present in groups around the basal antennal segments and each coxa. Tubular ducts 
sparse, of one size only, confined to metathorax and abdomen, forming transverse rows. Trilocular 
pores of similar distribution to those on dorsum. 

Notes. Specimens examined of Ripersia wiinni Reyne are identical with E. euro- 
paea. As pointed out by Reyne, the denticle on the claw is, at times, hardly percepti- 
ble and in some specimens it is absent entirely. 



Euripersia tomlinii (Newstead) (comb, n.) 

(Text-fig. 9) 

Ripersia tomlinii Newstead, 1892 : 146, 147. 

Ripersia tomlinii Newstead, Newstead, 1903 : 186 (in part). 

Ripersia exul Green, 1924 : 46, 47 (syn. n.). 

Ripersia mesnili Balachowsky, 1934 : 67-70 (syn. n.). 

Ripersia exul Green, Green, 1934 : m- 

Habit and distribution. Described by Newstead as " Dull orange-yellow, . . . 
Sac of the female globose or ovate, composed of close, white waxy material, very 
compact on the inside ;" Green described the ovisac as measuring 5 mm. in diameter. 
The insect lives on grass roots in association with ants' nests. Material has been 
examined from the following localities : Channel Islands. Guernsey : Moulin 
Huet, ix.1891 (Miss Tomlin) (Type) ; Chappelle dom Hue, 24.vii.1923 (J. R. le B. 
Tomlin {exul Type) ; viii . 1958 (R. H. Le Pelley) ; Vazon Bay, ix . 1924 (E. E. Green) ; 
Belle Elizabeth; Lihou, ix.1925 (E. E. Green). Herm : ix.1924 (E. E. Green); 
Rat Is., viii. 1923 (J. R. le B. Tomlin). Alderney : viii. 1892 (W. A. Luff). Sark : 
(W. A. Luff). England. Suffolk : ix,.i932 (A. S. Watt). 



THE BRITISH PSEUDOCOCCID AE 



27 




Fig. 9. Euripersia tomlinii (Nevvstead). 



28 D. J. WILLIAMS 

Recognition characters. Adult female broadly oval about 3-5 mm. long, anal lobes poorly 
developed. Antennae 6-7-segmented, 230-260 microns long, the third segment often divided 
into two of equal length and the terminal segment often showing signs of division. Legs small 
and slender, claw with a distinct denticle ; trochanter + femur 145-175 microns, tibia + tarsus 
170-205 microns. Labium 70-80 microns long. Circuli 2 or 3 in number, on the second, third 
and fourth segments, all transversely elliptical and with a fiat protuberance ; the circulus on the 
fourth segment much smaller than the others and sometimes absent. Ostioles with inner edges 
of lips sclerotized and with about 3 trilocular p>ores and an occasional seta on each lip. Anal ring 
with 3 rows of pores and 6 setae over twice length of diameter of ring. Cerarii numbering 2 pairs 
only ; anal lobe cerarii each with a pair of small conical setae and about 5 trilocular pores, the 
space between the setae sclerotized ; a single auxiliary seta present anterior to the cerarius. 
Penultimate cerarii similar but cerarian setae smaller. 

Dorsal setae not numerous, a short slender type interspersed with a minute lanceolate type. 
Tubular ducts present each without noticeable collar, situated more or less in single rows at 
anterior and posterior edges of abdominal segments but scattered on thorax and head ; sparse. 
Trilocular pores not numerous, evenly distributed. 

Ventral surface with apical setae longer than anal ring setae. Ventral setae few, all slender 
but longer than those on dorsum. Multilocular disc pores arranged in single rows at anterior 
and posterior edges of abdominal segments, mainly in median areas ; a few also present on 
thorax. Quinquelocular disc pores present between first coxae and clypeus and labium, usually 
very few, in which case they may number about 6 but sometimes they are more numerous and 
may be present on the mesothorax. Tubular ducts similar to those on dorsum, situated between 
the multilocular disc pores on each of abdominal segments and on the lateral areas ; on the 
thorax and head they become scattered. Trilocular pores sparse. 

Notes. Some confusion seems to have arisen in identifying this species. The third 
antennal segment is often divided into two segments of equal length and the terminal 
segment often shows signs of division. When Green described R. exul he stated that 
it differed from R. tomlinii in having more robust antennae and a broader and 
shorter labium. It now seems evident that Green based this distinction on misidenti- 
fied specimens of Euripersia europaea. The type of R. exul is identical with E. tomlinii. 

Specimens of Ripersia mesnili Balachowsky, described from Corsica, also seem to 
be identical with E. tomlinii. Balachowsky described the tubular ducts as being on 
the venter only but they are present on both surfaces on specimens examined from 
Lac de Nino and Bergeries de Paratella mentioned in his original description. The 
name R. mesnili is, therefore, sunk as a synonym of E. tomlinii. 



HELIOCOCCUS §ul9 
Heliococcus SUI9, 1912 : 39-48. 

Type of genus Heliococcus bohemicus Sul?. 

Since Sulg erected this genus, a few more species were added to it by Goux (1934) 
and a number of new species have been described by Borchsenius (1949). By far the 
greatest number occurs in the Palaearctic Region but four are known from North 
America and one from the Ethiopian Region. 

Belonging to the Phenacoccus group with a denticle on the claw and with quinque- 
locular pores, the genus is easily recognized by the peculiar crateriform ducts, the 
larger ducts often with 1-4 minute setae attached to the base of the duct prominence. 
The following species seems definitely to belong to this genus. 



THE BRITISH PSEUDOCOCCID AE 29 

Heliococcus minutus (Green) (comb, n.) 

(Text-fig. 10) 
Phenacoccus minutus Green, 1925 '. 519. 

Habit and distribution. Described as clustered on the underground stems of 
Erica cinerea, Channel Islands, Guernsey, L'Ancresse, September. The adult 
female is described as dull purplish. Body rather closely dusted with coarse mealy 
secretion. No marginal tassels but with a pair of divaricating liguliform processes 
projecting from the anal area. This insect has so far not been found in Britain. 

Recognition characters. A small oval species measuring approximately 1-5 mm. long, 
anal lobes well developed, with a dorsal area of weak sclerotization either same size or larger in 
area than ring. Antennae 9-segmented, 310-370 microns long. Legs normal, claw with a distinct 
denticle, trochanter + femur 215-220 microns, tibia + tarsus 225-260 microns. Labium 
95-105 microns long. Circulus absent. Ostioles present, each with 4-5 trilocular pores, setae 
often absent, especially on the posterior pair, but usually 2-3 present on each lip of anterior pair. 
Anal ring with 6 setae about twice length of diameter of ring. Cerarii numbering 18 pairs, each 
with 2 lanceolate setae ; anal lobe cerarii with a small concentration of about 6-8 trilocular pores ; 
anterior cerarii with cerarian setae much smaller and accompanied by 3 or 4 trilocular pores. 

Dorsal surface with minute lanceolate setae. Trilocular pores not numerous but evenly 
distributed. Crateriform ducts present, of 3 sizes. The largest type each with 3 minute setae 
attached to the base of the duct prominence, situated mainly on the mid-line and around the 
submargins but their positions not constant. An intermediate type, usually with 2 setae attached 
to the base of the duct prominence, situated mainly in the submedian areas. Both types often 
occupying similar positions in different specimens. A small type of duct usually present on the 
margins and occasionally in the submedian areas, these very sparse, without setae attached to 
the base of the duct prominence but often a single seta in close association with it. 

Ventral surface with a pair of stout apical setae longer than anal ring setae. Anal lobes each 
with a small elongate strip of sclerotization, not attached to the apical seta. Ventral setae 
slender and sparse but a few lanceolate setae, similar to those on dorsum, occupying the lateral 
areas. Multilocular disc pores confined to fifth and posterior segments, very few and present in 
the median areas at posterior edges of segments. Quinquelocular disc pores in median and 
submedian areas, scattered on head and thorax, but lying in transverse rows on abdominal 
segments mainly at anterior edges. Trilocular pores sparse. Tubular ducts with oral collar on 
abdomen and occasionally on metathorax, situated in single transverse rows but absent in 
submarginal areas, there being scarcely more than 20 altogether. A few crateriform ducts of the 
same size as the smallest type on dorsum, present around margins of metathorax and some of 
abdominal segments. 



HETEROCOCCUS Ferris 

Heterococcus Ferris, 191 8 : 65. 
Heterococcus Ferris, Morrison, 1945 : 45. 

Type of genus Heterococcus arenae Ferris. 

The important characters of this genus are the 8-9-segmented antennae, claw 
usually with a denticle, and the presence of quinquelocular disc pores on the dorsum 
and venter which replace the normal trilocular pores. All the species are found at 
the bases of grass stems and only one species is known from Britain. 



30 



D. J. WILLIAMS 




Fig. io. Heliococcus minuius (Green). 



THE BRITISH PSEUDOCOCCI D AE 31 

Heterococcus pulverarius (Newstead) 

Ripersia pulveraria Newstead, 1892 : 145. 
Phenacoccus nudus Green, 1926 : 172. 
Heterococcus nudus (Green), Green, 1928 : 21. 
Heterococcus pulverarius (Newstead), Williams, 1961 : 673. 

The name has been given in error to a few species here discussed under Trionymus. 
A redescription and illustration has been given recently by Williams. 

England. Cheshire : Sandiway, A grostis tenuis, viii.1891 (R. Newstead). Surrey : 
Camberley, vi.1925 (E. E. Green). Berkshire : Silwood Park, Holcus mollis, Festuca 
sp. 8.viii.i949 (K. L. Boratynski). 

NIPAECOCCUS Sulg 
Nipaecoccus SUI9, 1945 : 1-48. 
Nipaecoccus SUI9, Ferris, 1950 : 103. 

Type of genus Dactyhpius nipae Maskell. 

The members of this genus are characterized by a peculiar blue-green or brown 
body content which shows up particularly during the preparation in caustic potash. 
There are always some dorsal setae of the same size or near the same size as cerarian 
setae and the antennae are usually 7-segmented, 



Nipaecoccus nipae (Maskell) 

Dactylopius nipae Maskell, 1893 : 230. 
Pseudococcus nipae (Maskell), Fernald, 1903 : 107. 
Pseudococcus nipae (Maskell), Green, 1917 : 262, 263. 
Pseudococcus nipae (Maskell), Green, 1930 : 10. 
Nipaecoccus nipae (Maskell), Sul^, 1945 : 1-48. 
Nipaecoccus nipae (Maskell), Ferris, 1950 : 109. 

A species recorded from palms in a few greenhouses in Britain. For a description 
and illustration see the work by Ferris. 

England. Surrey : Kew, Royal Botanic Gardens, Cocos, Kentiopsis, Sahal, 
xii.1916. London: Palm (J. C. F. Fryer). Scotland. Midlothian: Edinburgh, 
Royal Botanic Gardens, Calyptrogyne sp. ix.1925 (E. E. Green). 



PELIOCOCCUS Borchsenius 

Peliococctis Borchsenius, 1948 : 954. 

Type of genus Phenacoccus chersonensis Kiritchenko. 

This genus is probably of world wide distribution although the bulk of the known 
species is from the Palaearctic Region. The distinguishing features of the genus are 
clusters of multilocular disc pores, each cluster with one or more slender tubular 
ducts near the centre ; claw with a denticle ; often with quinquelocular disc pores 
on the venter. 



32 D. J. WILLIAMS 

Peliococcus balteatus (Green) 
(Text-fig. ii) 

Phenacoccus balteatus Green, 1928 : 21. 

Peliococcus balteatus (Green), Borchsenius, 1949 : 244. 

Habit. Described by Green as " Colour of living examples pale yellow with a fringe 
of very short tassels, with a pair of slightly longer tassels at the posterior extremity." 
From the under surface of the foliage of various grasses, more particularly on Arrhena- 
therum elatius, Cheddar Gorge, Somerset, England, viii.1926. This is the only 
record for Britain. 

Recognition characters. Adult female elongate oval attaining a length of 3 '5 mm. in the 
available specimens ; anal lobes moderately developed. Antennae g-segmented, 450-490 
microns long. Legs normal, rather large, claw with a denticle, trochanter + femur 315-350 
microns, tibia + tarsus 380-425 microns. Labium about no microns long. Circulus present, 
oval. Anterior and posterior ostioles with 2-4 setae and a few trilocular pores on each lip. Anal 
ring with 6 setae about twice length of its diameter. Cerarii numbering 1 8 pairs. Anal lobe cerarius 
usually with a group of 6 lanceolate setae and a few trilocular pores, the middle setae of the 
group surrounding a small sclerotized area. Anterior cerarii each with 2 smaller cerarian setae 
except those on head where there are up to 4 setae present ; each cerarius accompanied by a 
few trilocular pores. The cerarii often elevated slightly from the surrounding derm, especially 
those at the anterior and posterior ends of the body. 

Dorsal surface of body with minute setae, sparse, some often with one or two trilocular pores 
near the base. Trilocular pores very few. On all segments except IX + X, there are pore clusters 
of 3-6 multilocular disc pores, the commonest number being 4. A single minute tubular duct 
present in the centre of each cluster. Clusters arranged in transverse rows at posterior edges 
of segments except on head where they become scattered. 

Ventral surface of each anal lobe with a small area of sclerotization and an apical seta longer 
than anal ring setae. Ventral body setae around the margins, small and similar to those on 
dorsum ; in the median areas they are slender and much longer. Pore clusters arranged mainly 
on head, in transverse rows on segments III-V and around submargins of third and all posterior 
segments except IX + X. Multilocular disc pores, smaller than those in clusters, abundant at 
posterior edges of sixth and posterior segments. Quinquelocular disc pores few, present in 
median areas only between coxae and at anterior edges of segments II-VII. Tubular ducts with 
oral collar of one size, in transverse rows just anterior to the multilocular disc pores on segments 
VII and VIII and occasionally on margins of these segments. Trilocular pores sparse. 

Notes. This species seems to come close to P. venustus (Green) from Iceland and 
P. saratogensis (Rau) from U.S.A. These two species have clusters of multilocular 
disc pores consisting mainly of 3 pores, whereas in P. balteatus the clusters consist 
mainly of 4 pores. Furthermore P. venustus has 8-segmented antennae and those of 
the other two species are 9-segmented. Both P. balteatus and P. saratogensis have a 
similar distribution of multilocular disc pores in transverse rows on segments VI-VIII 
but P. venustus has an extra row on segment V. 

PHENACOCCUS Cockerell 

Phenacoccus Cockerell, 1893 : 318. 
Phenacoccus Cockerell, Ferris, 1950 : 120. 

Type of genus Pseudococcus aceris Signoret. 

Some of the important characters of this genus are the 8-9-segmented antennae, 
a denticle on the claw, cerarii numbering from 9-18 pairs. Although not always 



THE BRITISH PSEUDOCOCCID AE 




Fig. 11. Pel iococciis balteadts (Green). 



ENTOM. 12, I. 



34 D- J. WILLIAMS 

present, there are often quinquelocular disc pores on the venter and the dorsal setae 
are often minute and lanceolate. Borchsenius (1949) has resurrected the genus 
Paroudahlis Cockerell because the type species Boisduvalia piceae Loew has larger 
tubular ducts on the dorsum than on the venter. No such distinction is made here 
although two British species possess these larger ducts. Only three species are known 
from Britain and these may be separated by the following key : 

1 With 9-14 pairs of cerarii .......... 2 

- With 18 pairs of cerarii ......... aceris (Signoret) 

2 Cerarii numbering 9 pairs, circuli absent ..... interruptus Green 

- Cerarii numbering 14 pairs, circuli numbering 2-3, small and with rounded projections 

sphagni (Green) 

Phenacoccus aceris (Signoret) 

Pseudococcus aceris Signoret, 1875 : 329. 
Pseudococcus ulicis Douglas, 1888 : 88. 
Pseudococcus ulmi Douglas, 1888a : 124. 
Pseudococcus aceris Signoret, Douglas, 1890 : 153. 
Pseudococcus quercus Douglas, 1890 : 154. 
Pseudococcus socius Newstead, 1892 : 144. 
Phenacoccus aceris (Signoret), Fernald, 1903 : 90. 
Phenacoccus aceris (Signoret), Green, 191 5 : 180. 
Phenacoccus aceris (Signoret), Green, 1921 : 151. 
Phenacoccus aceris (Signoret), Green, 1923 : 215. 
Phenacoccus aceris (Signoret), Green, 1926 : 182. 
Phenacoccus aceris (Signoret), Green, 1928 : 30. 
Phenacoccus aceris (Signoret), Ferris, 1950 : 126. 

This is one of the most widely spread mealy-bugs in Britain. It is found on a variety 
of trees and is, apparently, very common on gorse. Ferris has given an excellent 
illustration and in his description he stated that there are 2 circuli present. This 
condition is probably the more common, but specimens at hand collected at Adel, 
Yorkshire, on gorse have only a single circulus. In other respects they are similar 
to the specimens with 2 circuli. 

Borchsenius (1949) has accepted the name Phenacoccus quercus (Douglas) for 
specimens collected in Armenia on oak, stating that the species differs from P. aceris 
in possessing only a single circulus. Two specimens of Douglas' original material 
have been prepared and, although they are in poor condition, there is sufficient 
evidence of the presence of 2 circuli. It seems possible that the Armenian material 
represents a different species. 

The specimens actually seen by the writer are listed below but there are many other 
records in the literature. 

England. Berkshire : Windsor, Quercus sp., 1930 (H. Donisthorpe) ; Silwood 
Park, Quercus sp., 26. v. 1948, Carpinus betulus, 26. iv. 1949, Fagus sp., 3. v. 1955 
(K. L. Boratynski). Buckinghamshire : Latimer, Ulmus sp., 29. v. 1929 (F. Laing). 
Cheshire : Chester, Sorbus aucuparia, Laburnum sp., Quercus sp. (R. Newstead). 
Dorsetshire : Lyme Regis, Ulex europaeus, iv.1920. Essex : Epping Forest, Quercus 
sp., 25. V. 1923 (C. L. Wittycombe) . Hampshire: Bournemouth, Ulex europaeus, 
16.iv.1902 ; Beaulieu, Myrica gale, vi.1919. Kent: Bearsted, Ulex europaeus, 



THE BRITISH PSEUDOCOCCID AE 35 

i.vi.i8g6. London: Lewisham (?^/w/ Type). Norfolk: Ingoldisthorpe, U lex euro- 
paeus, viii.1891 (R. Newstead). Northumberland : Riding Mill, Sycamore, vii.1950 
(D. J. Williams). Surrey : Camberley, U lex minor, ^.xi.igzg (E. E, Green) ; Oxshott, 
Castanea sativa, Fagus sylvatica, 22.iv.1921 (K. G. Blair) ; Kew, Apple, 3. v. 1955 
(K. L. Boratynski) ; Peach Hill, Ulex europaeus, 12.vii.1956 (K. L. Boratynski). 
Warwickshire: Farnborough, Buxus sempervirens, 25.vii.1922 (R. Newstead). 
Wiltshire: Wootton Bassett, peach, 21. iv. 1938. Yorkshire: Wakefield, currant 
(R. Newstead) {socius Type) ; Adel, Ulex europaeus, 24. vi. 1956 (D. J. Williams). 
Scotland. Kincardineshire : Strachan, Ulex europaeus, 30. iv. 1927 (G. D. Morison). 

Phenacoccus interruptus Green 
(Text-fig. 12) 

Phenacoccus interruptus Green, 1923 : 215. 
Phenacoccus interruptus Green, Green, 1925a : 43. 
Phenacoccus interruptus Green, Green, 1928 : 30. 

Habit and distribution. Described originally as " Colour pale purplish, masked 
by a rather close covering of white powdery secretion." Found on grasses from the 
following localities in England. Surrey : Camberley, 28.viii.1922 (Type), I5.viii. 
1923, 3.ix.i934 (E. E. Green). Somersetshire: Cheddar (E. E. Green). 

Recognition characters. Body elongate elliptical, sides subparallel, length about 3'5 mm. 
anal lobes well developed. Antennae g-segmented, 380-405 microns long. Legs well developed, 
normal ; claw with a distinct denticle, trochanter + femur 240-275 microns, tibia + tarsus 
320-340 microns. Labium 90-120 microns long. Circulus absent. Ostioles present ; posterior 
pair with about 6 trilocular pores on each lip and apparently without setae ; anterior pair with 
about 6 trilocular pores and 4 setae on each lip. Anal ring with 2 rows of pores and 6 setae over 
twice length of diameter of ring. Cerarii confined to anterior and posterior ends of body only, 
there being 9 definite pairs present. Anal lobe cerarii, each with a pair of well-developed 
lanceolate setae and a loose group of trilocular pores accompanied by about 2 minute auxiliary 
setae and all borne on a sclerotized plate slightly less in area than anal ring. Cerarii on segments 
VIII and VII on smaller sclerotized plates, each cerarius with a pair of short lanceolate setae 
and about 4 trilocular pores. On segments VI and V, cerarii each with a pair of setae and 3-4 
trilocular pores but not on sclerotized plates. First 3 cerarii on head each with about 3 setae and 
fourth with 2 setae, all on sclerotized plates which become progressively smaller posteriorly. 
Sometimes single setae, a little stouter than dorsal body setae, present in positions of some of 
thoracic cerarii. 

Dorsal surface with numerous minute lanceolate setae. Trilocular pores evenly distributed. 
Tubular ducts present of one size only, these rather large and without an oral collar, scattered on 
head and thorax but forming transverse rows on abdominal segments. 

Ventral surface with a small elongate area of sclerotization on each anal lobe and an apical 
seta longer than anal ring setae. Ventral setae mainly slender and much longer than those on 
dorsum but small lanceolate setae present around margins. Multilocular disc pores on fifth and 
posterior segments only. Segment V with about 4 pores only, VI with 24-34, VII with 53-62, 
VIII with 66-84, IX +X with 36-48. Quinquelocular disc pores in median areas only. Trilocular 
pores on abdominal segments and around margins of thorax and head. Tubular ducts narrower 
than dorsal ducts, without an oral collar, present in transverse rows on all abdominal segments 
and a few in median and marginal areas of thorax. 

Notes. In his original description. Green described this insect as possessing " A 
single rather obscure, circular medio-ventral osteole." Furthermore he shows this 



D. J. WILLIAMS 











•fi^ 



® 




J 



o 



Fig. 12. Phenacoccus interruptus Green. 



THE BRITISH PS E UDOCOCCI D AE 



37 



in a figure. The holotype specimen shows a round hole in the position of the circulus 
but it is evident that it is merely damage, probably caused by a needle as there are 
similar holes near it. Other specimens examined show no sign of a circulus and the 
species must be considered as not possessing one. Borchsenius (1949) has recorded 
this species from Russia as Paroudablis interruptus and possessing a large circulus 
but this is probably a different species. A single distorted specimen is at hand from 
Dorsetshire collected on sedge which shows a rather large circulus and this is possibly 
the same as the Russian species. It has not been possible to describe this species and 
the matter is left in abeyance until more satisfactory material is collected. 

Phenacoccus sphagni (Green) 
(Text-fig. 13) 

Pseudococcus sphagni Green, 1915 : 178, 179. 
Pseudococcus sphagni Green, Green, 1920 : 119, 120. 
Phenacoccus sphagni (Green), Reyne, 1958 : 20. 

Habit and distribution. Colour of adult female pinkish or dull red-brown, 
lightly dusted with wax and with wax projections at anterior and posterior ends of 
body. Described originally in nests of Formica picea amongst sphagnum moss in 
swampy ground. England. Hampshire : New Forest, Matley Bog (H. K. Donis- 
thorpe) and again in the New Forest, July, 1918. This is the only record for Britain 
but Reyne has recently recorded it from Holland. 

Recognition characters. Body elongate oval about 4 mm. long, anal lobes well developed. 
Antennae 8-segmented, total length 400-450 microns. Legs normal, claw with a distinct denticle. 
Trochanter + femur 300-310 microns, tibia -f- tarsus 335 microns. Labium 75-90 microns 
long. Anterior and posterior ostioles present, lips each with 2-3 setae and 6-8 trilocular pores. 
Anal ring with 2 rows of pores and 6 setae about twice length of diameter of ring. Cerarii number- 
ing 14 distinct pairs, those missing being on the meso- and meta thorax and second abdominal 
segment. Each cerarius with 2 lanceolate setae and a group of trilocular pores. Circuli 2-3 in 
number lying on the third, fourth and occasionally fifth abdominal segments. Circulus on segment 

III very small with a rounded projection ; circulus on segment IV slightly larger with projection 
flattened apically ; circulus on segment V, when present, minute. 

Dorsal surface with minute setae. Trilocular pores evenly distributed. Tubular ducts of 
one size only, each with oral collar barely perceptible, arranged in a wide band across the meso- 
and metathorax and each abdominal segment. 

Ventral surface with a stout pair of apical setae longer than anal ring setae and long single 
marginal setae on abdominal segments which become progressively shorter anteriorly. Body 
setae slender but much longer than on dorsum except for a few minute lanceolate setae around 
margins. Multilocular disc pores on sixth and posterior segments only ; numerous on segments 
IX + X and at anterior and posterior edges of VII and VIII ; a transverse row on posterior edge 
of segment VI. Tubular ducts of two sizes. A larger type, similar to those on dorsum, arranged 
singly or in pairs on margins of segments VI-VIII. A smaller type in marginal groups on segment 

IV and posterior segments, becoming fewer anteriorly. Small numbers also around margins of 
anterior abdominal segments and thorax and in median areas of thorax. Quinquelocular disc 
pores very few between anterior coxae, clypeus and labium and an occasional pore on mesothorax. 
Trilocular pores evenly distributed. 

Notes. The distinguishing features of this species are the 8-segmented antennae, 
the reduced number of cerarii and the small pecuHar circuli which are variously 
knobbed. 



38 



D. J. WILLIAMS 




Fig. 13. Phenacocciis sphagni (Green). 



THE BRITISH PSEUDOCOCCID AE 39 

PLANOCOCCUS Ferris 

Planococcus Ferris, 1950 : 164. 

Planococcus Ferris, Ezzat & McConnell, 1956 : 60. 

A genus erected by Ferris for a number of species, the essential characters being 
the presence of an anal bar on the ventral surface of each anal lobe, 18 pairs of 
cerarii, these without auxiliary setae, tubular ducts with oral rim entirely lacking. 



Planococcus citri (Risso) 

Dorthesia citri Risso, 1813 : 416-418. 
Pseudococcus citri (Risso), Fernald, 1903 : 99. 
Planococcus citri (Risso), Ferris, 1950 : 165. 
Planococcus citri (Risso), Ezzat & McConnell, 1956 : 65. 

Newstead (1903) described an insect under the name Dactylopius citri but the 
specimens are undoubtedly Pseudococcus latipes Green. The only specimens seen 
from Britain are from greenhouses in England from various localities and hosts. 
Excellent illustrations are available in the works by Ferris and Ezzat & McConnell. 



PSEUDOCOCCUS Westwood 

Pseudococcus Westwood, 1840 : 118. 
Pseudococcus Westwood, Ferris, 1950 : 170. 

Type of genus Coccus adonidum Linnaeus. 

The limits of this genus were defined by Ferris and the essential features are the 
presence of oral rim ducts, at least on dorsum, and 16 or 17 pairs of cerarii with 
auxiliary setae. 

There are no native British species and, although P.fragilis Brain occurs in the open 
in the southern counties, it has been introduced. The others are all greenhouse 
species. After remounting most of the specimens at hand, it is evident that all those 
identified and recorded as P. comstocki (Kuwana) from imported bananas should be 
referred to Dysmicoccus alazon recently described by Williams (i960). Specimens 
identified as P. maritimus (Ehrhorn) should be referred to P. latipes Green (=P. 
malacearum Ferris) . Evidence in recent years suggests that P. latipes is a widespread 
species and may have been confused with P. maritimus and P. comstocki. A key to 
the British species is given below. 

1 With not more than a single oral rim duct dorsally near each of most of cerarii. Multi- 

locular disc pores in transverse rows on most abdominal segments ... 2 

- With 2-3 oral rim ducts of different sizes dorsally near each of most of cerarii. Multi- 

locular disc pores about vulva only ...... adonidum (Linnaeus) 

2 With a large oral rim duct dorsally, posterior to each frontal cerarius. Anal lobe 

cerarii placed near inner edge of an area of sclerotization . . . latipes Green 

- Without large oral rim duct dorsally posterior to each frontal cerarius. Anal lobe 

cerarii placed near the centre of surrounding sclerotized area . . fragilis Brain 



40 D. J. WILLIAMS 

Pseudococcus adonidum (Linnaeus) 

Coccus adonidum Linnaeus, 1766 : 740. 
Dactylopius longispinus Targioni, 1867 : 1-87. 
Dactylopius longispinus Targioni, Newstead, 1903 : 168. 
Pseudococcus longispinus (Targioni), Fernald, 1903 : 104. 
Pseudococcus adonidum (Linnaeus), Ferris, 1950 : 174. 

In Britain, a greenhouse species only and, apparently, not so common as hitherto 
supposed. England. Cambridgeshire : Cambridge, fern, 17.x. 1935 (H. C. James). 
Berkshire : Reading, Phormium tenax, 24.viii.1948 (M. I. Crichton). Surrey : Kew, 
Royal Botanic Gardens, Stangeria sp., 7 . v . 1896. Scotlanid. Midlothian : Edinburgh, 
Royal Botanic Gardens, Cycas sp. (R. S. MacDougall). 

Pseudococcus fragilis Brain 

Pseudococcus fragilis Brain, 191 2 : 186. 
Pseudococcus gahani Green, 191 5 : 180. 
Pseudococcus gahani Green, Green, 1920 : 120. 
Pseudococcus gahani Green, Green, 1921 : 151. 
Pseudococcus gahani Green, Green, 193 1 : 100. 
Pseudococcus fragilis Brain, Essig, 1942 : 351. 
Pseudococcus gahani Green, Ferris, 1950 : 180. 
Pseudococcus fragilis Brain, De Lotto, 1958 : 96. 

Numerous specimens have been examined from various localities in England, 
Scotland, Wales and Channel Islands, some from greenhouses and others living 
in the open, especially in the southern counties. There is an excellent description 
and illustration under the name P. gahani in the work by Ferris. 

Pseudococcus latipes Green 

Dactylopius citri (Risso), Newstead, 1903 : 164. (Misidentification.) 
Pseudococcus longispinus var. latipes Green, 191 7 : 264. 
Pseudococcus maritimus (Ehrhom), Green, 1920 : 121. (Misidentification.) 
Pseudococcus maritimus (Ehrhorn), Green, 1921 : 151. (Misidentification.) 
Pseudococcus maritimus (Ehrhom), Green, 1928 : 31. (Misidentification.) 
Pseudococcus maritimus (Ehrhorn), Green, 1931 : 99. (Misidentification.) 
Pseudococcus malacearum Ferris, 1950 : 185 (syn. n.). 

This species is probably the commonest of the greenhouse mealy-bugs in Britain. 
It was first described as a variety of P. longispinus and later Green regarded it as 
being identical with P. maritimus but it is quite distinct from both. A detailed 
description has been given by Ferris under the name P. malacearum and there seems 
to be little doubt that it is the same as P. latipes. The most striking feature is the 
anal lobe cerarius with two conical setae surrounded by a crowded mass of pores. 
This cerarius is situated at the inner edge of an oval sclerotized area which extends 
posteriorly to the base of the apical seta. Very often there is a small prolongation 
at the anterior edge of the sclerotized area. 

Specimens are at hand on a wide variety of hosts from many greenhouses in 
England, Wales, Scotland and Channel Islands. 



THE BRITISH PSEUDOCOCCI D A E 41 

RHIZOECUS Kiinckel d'Herculais 

Rhizoecus Kiinckel d'Herculais, 1878 : 163. 

Rhizoecus Kiinckel d'Herculais, Hambleton, 1946 : 50. 

Rhizoecus Kiinckel d'Herculais, Ferris, 1953 : 426. 

Type of genus Rhizoecus falcifer Kiinckel d'Herculais. 

All the hypogeic mealy-bugs in Rhizoecus and related genera have been studied in 
great detail by Hambleton. Differences in number of antennal segments, the lengths 
of the claw digitules and the presence or absence of eyes have led Hambleton to place 
some of the following species in the genera Ripersiella Tinsley and Morrisonella 
{=Coccidella Hambleton). In the present work all species are left in the genus 
Rhizoecus following the definition by Ferris (1953) . Six species are known from Britain 
and may be identified from the key. 

1 Circulus absent ............ 2 

- Circulus present ............ 3 

2 (i) Antennae 5-segmented, abdominal tergites each with 10-16 tritubular pores 

falcifer Kiinckel d'Herculais 
Antennae 6-segmented, abdominal tergites each with 2 or 3 tritubular pores at most 

dianthi Green 

3 (i) Multilocular disc pores present on venter, bitubular pores present halophilus (Hardy) 

- Multilocular disc pores absent on venter, tritubular pores present ... 4 

4 (3) With some tritubular pores on median and submedian areas of dorsum, labium 

60-85 microns long ........... 5 



- With a total of no more than 4 tritubul 

90-105 microns long . 
5 (4) Labium 60-70 microns long . 

- Labium 80-85 microns long . 



:ir pores on dorsal margins only, labium 

elongatus Green 

albidus Goux 

.cacticans (Hambleton) 



Rhizoecus albidus Goux 

(Text-fig. 14) 

Ripersia halophila (Hardy), Green, 1917 : 262. (Misidentification.) 
Rhizoecus (Pararhizoecus) albidus Goux, 1942 : 40. 

Habit and distribution. In life these insects are white and so small as to resemble 
Collembola, for which they may be mistaken. They feed in the soil on the fine rootlets 
of grasses and often do extensive damage. 

The species was first described from the roots of Festuca sp., at Bcssenay, France. 
Material is at hand from the following locahties in Britain : England. Hertford- 
shire : Rothamsted, 1936. Shropshire: Newport, 24. ii. 1939 (H. C. F. Newton). 
Staffordshire : 1947 (H. C. F. Newton) ; Newcastle, ix.1960. Surrey : Camberley 
(E. E. Green); Egham, xii.1959 (K. L. Boratynski). Herefordshire: Hereford, 
i.1932 (H. E. Durham). 

Recognition characters. A small elongate-oval species measuring approximately 1-5 mm. 
long. Posterior end of body rounded. Anal lobes poorly developed, each with i long ventral 
seta and 2 long dorsal setae, appearing as a group of 3. Eyes small. Antennae 6-segmented, 
geniculate, length 165-170 microns. Legs well developed, trochanter -f femur 120-130 microns, 
tibia + tarsus 130-135 microns ; claw long and slender with claw digitules as long as or slightly 
longer than claw, with a very minute apical knob. Labium 60-70 microns long. Ostioles well 
developed, lower lip of each posterior pair without setae or trilocular pores ; other lips each 



42 



D. J. WILLIAMS 




Fig. 14. Rhizoecus albidus Goux. 



THE BRITISH PSEUDOCOCCID AE 43 

with 2-4 setae and 4-7 trilocular pores. Circulus present within borders of segment IV, heavily 
sclerotized and cone shaped with distal circular plate containing about 7 circular areas. Anal 
ring rather large in comparison to size of body ; anal ring setae about twice length of ring at its 
greatest diameter and about same length as apical setae. 

Dorsal surface with numerous short pointed setae and forming distinct groups on thorax 
and head. Trilocular pores following the general distribution of the setae. Tritubular pores not 
numerous, there being rarely more than 3 on any segment of the abdomen or thorax and sometimes 
only a single one on mid-line ; tending to be scattered on head. Some very minute tubular ducts 
present on abdominal segments, these very simple and heavily sclerotized ; some segments 
entirely without ducts and others with at most 6. 

Ventral surface with a more or less quadrate sclerotized area containing 4 marginal setae just 
anterior to clypeus. Body setae short and slender. Multilocular disc pores absent. Trilocular 
pores sparse. The setae and trilocular pores forming groups on thorax and head. Tritubular 
pores very few, there being a single marginal pore on some of posterior abdominal segments, 
occasionally one on thorax and one between antennal bases. Small tubular ducts, similar to 
those on dorsum, arranged 2-6 across some of posterior abdominal segments. 

Notes. Goux placed this species in the subgenus Pararhizoecus because it possesses 
a circulus similar to that of the type species R. [Pararhizoecus) petiti Goux. It is left 
in Rhizoecus following the definition of this genus by Ferris. Dr. Harold Morrison 
has kindly examined specimens of this species and has given valuable comments on 
its differences with R. cacticans and R. elongatus. 

Rhizoecus cacticans (Hambleton) 

Rhizoectis elongatus Green, Green, 1926 : 174 (in part as a misidentitication). 
Ripersiella cacticans Hambleton, 1946 : 64. 
Rhizoecus cacticans (Hambleton), Ferris, 1953 : 432. 

Specimens examined, all from greenhouses, include the following : England. 
Hampshire : Bournemouth, Phyllocactus sp. (as a misidentification of R. elongatus). 
Isle of Wight: Sandown, roots of cactus, ii.1949 (E. Elkan). Essex: Laindon, 
Mammillaria sp., roots, xii.1928 (G. Fox-Wilson). 

This species belongs to a group similar to R. albidus and R. elongatus. All are 
devoid of multilocular disc pores and possess smaU sclerotized tubular ducts. It 
comes closest to R. albidus but differs mainly in the length and shape of the labium 
(see key). Ferris has redescribed and illustrated this species. 

Rhizoecus dianthi Green 

(Text-fig. 15) 

Rhizoecus dianthi Green, 1926 : 175. 

Morrisonella dianthi (Green), Hambleton, 1946 : 23. 

Coccidella dianthi (Green), Hambleton, 1946a : 177. 

Habit and distribution. Appearance in life not known. This species is known 
only from the roots of various plants in greenhouses. It was described from Dianthus 
plumarius and D. harhatus, England, Surrey, Wisley, Royal Horticultural Society's 
Gardens, and from other plants in the London area. Collected since from Wisley 
on other occasions on various plants. Specimens have also been seen from Hampshire: 
Salisbury, Pelargonium sp. roots, viii . 1926 (G. Fox- Wilson). Somersetshire : Taunton, 
Adiantum sp., 7 , iii . 1904. 



44 



I). J. WILLIAMS 




^ 





Vio. 15. Rhizoecus dianthi Cireen. 



THE BRITISH PSEU DOCOCCl D A E 



45 



Recognition characters. Body broadly oval, length about 2 mm. Anal lobes poorly 
developed, each with a long ventral apical seta and 2 slightly shorter dorsal setae. Antennae 
6-segmented about 170-175 microns long. Eyes present, very small. Legs normal for the genus, 
slender ; posterior coxae with a few large oval areas ; trochanter + femur 145-155 microns, 
tibia + tarsus 145-155 microns. Labium 85-95 microns long. Claw slender with very short 
pointed digitules. Ostioles with inner edges of lips sclerotized and with 2-4 setae and 5-8 
trilocular pores on each lip. Circulus absent. Anal ring inclined to ventral surface when mounted 
on the slide. 

Dorsal surface with numerous short slender setae. Trilocular pores evenly distributed. 'Tritu- 
bular pores sparse, there being from 2-4 on some abdominal segments and absent entirely on 
other segments ; occasional pores present on thorax and head. 

Ventral surface with short setae similar to those on dorsum. Multilocular disc pores few on 
posterior abdominal segments only. Segment VII with i or 2, VIII with 4-7, IX + X with 3-6, 
there being scarcely more than 12 altogether. Tubular ducts absent. Trilocular pores distributed 
fairly evenly. Tritubular pores not constant in position although there is often a single marginal 
pore present on one of posterior abdominal segments and a single pore on margins of pro- and 
metathorax. 

Notes. This species comes very close to R. cypercdis (Hambleton) described from 
El Salvador and both may be identical. 

Rhizoecus elongatus Green 

(Text-fig. 16) 

Rhizoecus elongatus Green, 1925a : 174. 

Rhizoecus mesembryanthemi Green, 193 1 : 103 (syn. n.). 

M orrisonella elongcUa (Green), Hambleton, 1946 : 25. 

Morrisonella mesembryanthemi (Green), Hambleton, 1946 : 1,^. 

Coccidella elongata (Green), Hambleton, 1946a : 177. 

Coccidella mesembryanthemi (Green), Hambleton, 1946a : 177. 

Habit and distribution. Appearance in life not known. A greenhouse species 
described from the roots of Phyllocactus sp., England, Hampshire, Bournemouth. 
Recorded also from the roots of Mesembryanthemum sp., Surrey : Oxted (Type of 
R. mesembryanthemi). These are the only records from Britain but specimens are 
also at hand, collected by Dr. A. Reyne from Holland, Zeist Utrecht, on Aloe variegata 
roots. 

Recognition characters. A small elongate species with parallel sides attaining a length of 
2 mm., posterior end rounded, anal lobes with i long ventral seta and 2 slightly shorter dorsal 
setae. Antennae 6-segmented, about 240 microns long. Legs normal for the genus, claw slender 
and elongate with claw digitules either same length or slightly longer than claw, with minute 
apical knob ; trochanter -f femur 165-175 microns, tibia + tarsus 190 microns. Labium 
elongate, about 90-105 microns long. Eyes present, very small. Dorsal ostioles fairly well 
developed ; lower lip of each posterior pair \vith about 4 trilocular pores and without setae ; 
all other lips with 2-4 setae and about 4 trilocular pores. Cerarii absent. Circulus within borders 
of fourth abdominal segment, heavily sclerotized in the form of a truncated cone, wider than 
deep, the distal circular plate with numerous subcircular areas. Anal ring rather large for size 
of body, with 6 setae about twice length of diameter of ring and same length as apical setae. 

Dorsal surface of body with numerous short slender setae and trilocular pores, both evenly 
distributed on abdomen and metathorax but anteriorly forming large groups so that there are 
some areas entirely clear. Minute tubular ducts present on thorax and abdomen ; each in the 
form of a simple sclerotized internal tube ; majority of segments with, at most, about 6-8 



46 



^- 



D. J. WILLIAMS 




O 










i^.cA 




Fig. 1 6. Rhizoecus elongatiis Green. 



THE BRITISH PSEUDOCOCCI D A E 47 

in a transverse row. Tritubular pores sometimes absent entirely or more often with i or 2 on 
margins of each side of body or on one side only, there being usually one on head and others on 
some abdominal segments. 

Ventral surface with a small sclerotized plate anterior to clypeus. Short slender setae and 
trilocular pores numerous on abdomen but anteriorly they become sparse and form definite 
groups. Multilocular disc pores and tritubular pores absent. Minute tubular ducts, similar to 
those on dorsum, very sparse, there being 2-8 in transverse rows on abdominal segments and 
occasional ducts on thorax. 

Notes. This is a distinctive species with reduced numbers of tritubular pores. 
In his original description, Green stated that these pores were absent but in the speci- 
mens available this condition is very rare as there are usually one or two present on 
dorsum. Green's statement of the two different sized limbs in the type slide was due 
to the presence also of specimens of R. cacticans. He also stated, in his description 
of R. mesembry anthem i, that the body was completely devoid of setae. The original 
material has numerous setae and the specimens are identical with R. elongatus. 



Rhizoecus falcifer Kiinckel d'Herculais 

Rhizoecus falcifer Kiinckel d'Herculais, 1878 : 164. 

Riper sia terrestris Newstead, 1895a : 213. 

Ripersia terrestris Newstead, Newstead, 1903 : 190. 

Rhizoecus decor atus Green, 1926 : 177. 

Rhizoecus decoratus Green, Green, 1928 : 31. 

Rhizoecus decoratus Green, Green, 1930 : 10. 

Rhizoecus decoratus Green, Green, 1931 : 102. 

Rhizoecus terrestris Newstead, Green, 1931 : 102. 

Rhizoecus decoratus Green, Green, 1934 ^ m- 

Rhizoecus falcifer Kiinckel d'Herculais, Hambleton, 1946 : 53. 

Rhizoecus falcifer Kiinckel d'Herculais, Ferris, 1953 : 444. 

This species is now known from many parts of the world. In Britain it is found 
only in greenhouses on roots of numerous plants and is, apparently, quite common. 
Formerly known in Britain under the names Rhizoecus terrestris and R. decoratus, 
it was shown by Hambleton that these were the same as R. falcifer. For an illustration, 
see the work by Ferris. British specimens examined include the following : England. 
London : palm roots, 1895 (C. O. Waterhouse) {terrestris type). Yorkshire : Hull, 
Eastington, roots of Abutilon sp., iv.1925 {decoratus type). Lancashire : Fallowfield, 
Carex sp., roots, I9.viii,i926 (J. H. Watson). Surrey : Wisley, Royal Horticultural 
Society's Gardens, Dracaena sp. roots, 6.iii.i928 (G. Fox- Wilson), roots of various 
plants, 5. V. 1953 (D. J. Williams) ; Kew, Royal Botanic Gardens, 2.11.1932 (G. 
Fox-Wilson); Richmond, Gardenia roots, xii.1950. Warwickshire: Studley, lily 
roots, 27. V. 1935 (J. F. Perkins). Ireland. Dublin, Adiantum sp., roots, iii.1901. 

Rhizoecus halophilus (Hardy) 
(Text-fig. 17) 

Coccus halophilus Hardy, 1868 : 136, 137. 

Dactylopius radicum Newstead, 18956 : 235 (in part, misidentification). 

Ripersia halophila (Hardy), Newstead, 1903 : 192. 



48 



D. J. WILMAMS 




Fig. 17. Rhizoecus halophilus (Hardy) . 



THE BRITISH PSE U UOCOCCI D AE 49 

Ripersia halophila (Hardy), Green, 1921 : 191. 
Rhizoecus halophila (Hardy), Green, 1926 : 174. 

Habit and distribution. The reference to this species has long since been for- 
gotten and as it is in a journal which may not be readily available, the full original 
description is given below. 

" 6. Coccus halophilus, J. H. The Cocci offer few tangible specific characters. The 
present species apart from its peculiar habitudes cannot be readily discriminated by 
words. It is scarcely a quarter of a line long, oval, opaque white, without a hard 
scale, abundantly supplied with a white secretion. Found on the steep sea-banks 
near Fastcastle [Scotland : Berwickshire] among the roots of Ligusticum Scoticum, 
and Rhodiola rosea ; and afterwards on the roots of Statice A nneria in the grey wacke 
cliffs near Siccar Point. In both cases it follows the long fibrils minutely interwoven 
through the loose slaty debris." 

Newstead has described the insect as pure white and with two appendages at the 
caudal extremity. It feeds only on the roots and is, apparently, not associated with 
ants. The known distribution is as follows : 

England. Norfolk : Blakeney Point, roots of Armeria maritima, vii.1920 (E. E. 
Green). Isle of Man: Port Erin, roots of grass, 21. ix.1918 (R. Newstead). Scotland. 
Fifeshire : Isle of May, roots of grasses, ix.1913 (W. Evans). Inverness-shire: 
(Outer Hebrides) St. Kilda, grass roots, vi. 1902 (R. Newstead). Ross and Cromarty : 
Swordale, roots of Calluna sp., 21.1.1921 (D. J. Jackson). Wales. Anglesey: 
Puffin Island, on Armeria muritima. Ireland. Donegal: among turf, viii.1942 
(J. Lister). 

Recognition Characters. Adult female ovate, about 1-5 mm. long. Posterior end with 
poorly developed anal lobes, each with i long ventral apical seta and 2 smaller dorsal setae. 
Antennae 6-segmented, total length approximately 155-170 microns. Legs normal, trochanter + 
femur 11 5-1 30 microns, tibia -\- tarsus 130-145 microns, claw with slender digitules about same 
length or longer than claw with very small knob at apex. Labium about 75 microns long. 
Eyes present, minute. Ostioles well developed, inner edges of lips sclerotized and each lip with 
about 2 setae and 2 or 3 trilocular pores ; occasionally without setae. Circulus present within 
borders of fourth abdominal segment, sclerotized and small, in form of truncate cone, wider 
than deep, distal surface with minute circular areas. Anal ring with 6 setae twice length of its 
diameter and about same length as apical setae. 

Dorsal surface with slender setae of various lengths but mostly small. Trilocular pores not 
numerous. Tubular ducts small and slender without oral collar, in transverse rows of up to 8 on 
abdominal segments and becoming scattered on thorax and head. Bitubular pores present, 
very sparse, there being a marginal series of about 8 on each side of body and 2 or 3 on midline. 

Ventral surface with short slender setae but tending to be longer on posterior abdominal 
segments, not numerous. Trilocular pores sparse. Bitubular pores absent. Tubular ducts, 
similar to those on dorsum, very few, situated in transverse rows on abdominal segments and 
becoming scattered on thorax and head. Multilocular disc pores on seventh and posterior 
segments only, there being scarcely more than 50 altogether but some specimens with as few 
as 30. 

Notes. The identification of this species is based on those made by Newstead and 
Green. It comes very close to Rhizoecus {Pararhizoecus) petiti described by Goux 
(1941^) another species with bitubular pores but in this case the multilocular disc 
pores are much more numerous and are situated on the fifth and posterior segments 
whereas in R. halophilus they are present on the seventh and posterior segments. 

ENTOM. 12, 1. 4 



50 D. J. WILLIAMS 

SACCHARICOCCUS Ferris 

Saccharicoccus Ferris, 1950 : 216. 

Type of genus Dactylopius sacchari Cockerell. 

Recognition characters. Pseudococcidae with 7-segmented antennae. Two pairs of dorsal 
ostioles present. Circulus longer than wide, situated across the segmental line between fourth 
and fifth abdominal segments, medially constricted. Legs without a denticle on the claw. Minute 
irregular pores present on the surrounding derm near the attachment of the posterior coxae. 
Tubular ducts either few on ventral side of abdomen or numerous on both surfaces. Multilocular 
disc pores on dorsum and venter. Cerarii i pair, on anal lobes only. 

Notes. This genus was erected for the single species Dactylopius sacchari, having 
many characters similar to those of Trionymus. The description given by Ferris has 
been modified to cater for the species described below as new. One of the striking 
features of the new species is the large number of small irregular-shaped pores near 
the attachment of the posterior coxae. This character is shared also by 5. sacchari 
but has, apparently, been overlooked. An examination of many correctly stained 
specimens of 5. sacchari from various localities, shows that these pores are always 
present. The long stout marginal setae on the abdomen of 5. sacchari are probably 
only of specific value and have been excluded from the generic definition as given 
by Ferris. 

Saccharicoccus penium sp. n. 

(Text-fig. 18) 

Habit and distribution. Description in life not known. Found at the base of 
grasses, England, Hampshire, Yateley, 25. ix. 1926 (E. E. Green). 

Recognition characters. Adult female elongate, with more or less parallel sides, attaining 
a length of 3-5 mm. Anal lobes moderately developed. Antennae 7-segmented, small, about 
240-250 microns long. Legs small and slender, trochanter + femur 150-170 microns, tibia + 
tarsus 165-175 microns. Labium about 70 microns long. Anterior and posterior ostioles poorly 
developed, each lip with 2-3 trilocular pores and lacking setae. Circulus large, longer than wide 
with sides constricted, hour-glass shaped. Cerarii represented by anal lobe pair only, each with 
2 short conical setae set close together, surrounded by a group of about 7 or 8 trilocular pores and 
accompanied by a single auxiliary seta. Anal ring with 6 setae about twice as long as diameter 
. of ring. 

Dorsal surface with short pointed setae, not numerous. Trilocular pores with an even distri- 
bution. Multilocular disc pores forming definite transverse rows at posterior edges of segments 
V-VIII. Elsewhere on the body they tend to be scattered, although there is a noticeable concen- 
tration on the head margin and they are absent or nearly so in the median areas of the thorax. 
Tubular ducts of 2 types. A smaller type, slender, without any appreciable oral collar, in trans- 
verse rows in the middle of metathorax and abdominal segments. A larger type of duct, each 
with a large collar and a larger flange-shaped membranous ring arising from the inner edge of 
the collar, situated in transverse rows on segments VI-VIII and in a small group on anal lobes. 
Others present around head margins and an occasional one present elsewhere on the body. 

Ventral surface with a pair of apical setae only slightly longer than anal ring setae. Body 
setae not numerous, all short and slender but tending to be longer than those on dorsum, especi- 
ally in median areas. Minute simple pores of different shapes and sizes clustered near posterior 
coxae from which they extend laterally and posteriorly to segment III. Trilocular pores evenly 
distributed, sparse. Multilocular disc pores abundant in transverse rows at posterior edges of 
segments V-VIII. They are also numerous on segments IX + X, at anterior edges of segments 



THE JiRlTlSll 1'SEL'DUC(JCC1DAE 



51 



O 




J 



o 



o 



& 






■^ 



:A 



o 



J 



o 



Fig. 18. Saccharicocciis peniiiiu sp. n. 



52 D. J. WILLIAMS 

VI-VIII and in a submarginal zone around body. Elsewhere they become scattered. Tubular 
ducts of same 2 sizes as on dorsum. A smaller type mainly in transverse groups in middle of 
segments V-VIIf. A larger type numerous in marginal groups on abdominal segments, sometimes 
extending to median areas especially on posterior abdominal segments. Present also around the 
submargins of thorax and head in close association with the multilocular disc pores. 

Notes. This species differs from S. sacchari in possessing much more numerous 
multilocular disc pores and tubular ducts. The minute irregular pores tend to be 
concentrated lateral and posterior to the hind coxae whereas in 5. sacchari they are 
mostly situated just anterior. 

SPILOCOCCUS Ferris 

Spilococcus Ferris, 1950 : 219. 
Spilococcus Ferris, McKenzie, i960 : 755. 

Ferris erected this genus for species with oral rim ducts and with the cerarii lacking 
auxiliary setae. As stated in the discussion on Atrococcus, McKenzie has limited 
the genus to those species possessing from 6-17 pairs of cerarii and has erected the 
genus Chorizococcus for species with from 0-4 pairs of cerarii. Although Atrococcus 
is distinct in having species with a black body content and with the cerarii on abdomen 
only, the position is complicated with Spilococcus cactearum McKenzie, a species 
with an obvious relationship to other species of Spilococcus yet possessing a black 
body content. It is left in this genus for the time being pending further research. 

Apart from S. cactearum, the other British species, 5. filicicola (Newstead), is 
placed here as a temporary measure. 



Spilococcus cactearum McKenzie 
(Text-fig. 19) 

Pseudococcus mamillariae Bouch^, Green, 1930 : 9. (Misidentification.) 
Spilococcus cactearum McKenzie, i960 : 757. 

Habit and distribution. According to McKenzie the adult female is covered 
with an even, light grey secretion. When boiled in caustic potash the body content 
turns black or blue-black. So far as is known, the species is confined to the Cactaceae. 
Specimens have been examined as follows : England. Essex : Laindon, vi.1928 
(T. M. Endean). Middlesex : Pinner, v. 1947 (R. E. Elkan). 

Recognition characters. Adult female broadly oval attaining a length of 2-5 mm. Anal 
lobes well developed. Antennae 8 -segmented, 310-335 microns long. Legs normal, posterior 
coxae with a few translucent pores, posterior tibia with a small group of such pores ; claw with 
a denticle ; trochanter + femur 215-240 microns, tibia + tarsus 240-250 microns. Labium 
about 70 microns long. Anterior and posterior ostioles present, inner edges of lips sclerotized, 
each lip with 2 or 3 setae and about 6 trilocular pores. Circulus rather large, oval. Anal ring 
with a double band of pores and 6 setae about twice length of its diameter. Cerarii numbering 
9-14 pairs, those on head and thorax variable in number but there are usually 7 pairs present 
on abdomen. Anal lobe cerarii each with a pair of slender conical setae set close together and 
accompanied by 5 or 6 trilocular pores and 3 or 4 auxiliary setae, the area at base of cerarian 
setae lightly sclerotized. Anterior cerarii each with setae more slender, sometimes wide apart 
but their presence can be detected by a group of 3 or 4 trilocular pores, without auxiliary setae. 



THE BRITISH PSEUDOCOCCI D AE 



53 




Fig. 19. Spilococcus cactearum McKenzie. 



54 D. J. WILLIAMS 

Dorsal surface of body with very small slender setae, not numerous. Trilocular pores evenly 
distributed. Tubular ducts with oral rim on segment VIII and all anterior segments, arranged 
in single transverse rows on abdomen, there being rarely more than 6 on each segment ; on the 
thorax and head they become scattered. 

Ventral surface with a pair of long apical setae longer than anal ring setae. Other setae slender 
but usually longer than those on dorsum especially on median areas, not numerous. Multilocular 
disc pores on abdomen only, confined to segments posterior to circulus. Arranged in transverse 
rows at posterior edges of segments and at anterior edges of segments VII and VIII. Tubular 
ducts with oral collar of two sizes. A small type, very few in transverse rows in middle of some 
of posterior abdominal segments and on anal lobes, an occasional duct sometimes present on 
thorax. A larger type in transverse rows on fourth and posterior segments immediately anterior 
to multilocular disc pores and usually continuous with noticeable marginal groups. One or two 
also present on thorax. Tubular ducts with oral rim arranged singly or in pairs on margins of 
segments IV-VIII but forward to metathorax they are more numerous, often with 2 or 3 on each 
segment. There is usually a pair between antennal bases and often i in median area of thorax. 
Trilocular pores not numerous but with an even distribution. 

Notes. This species has often been misidentified as Pseudococcus mamillariae 
based on the description of Coccus mamillariae Bouche. It has been shown by McKen- 
zie, on the basis of a report by Lindinger (1934), that it cannot be the species described 
by Bouche and consequently McKenzie has described this black cactus-infesting species 
as new, British specimens have been checked against material from France collected 
by A. Balachowsky and from Italy collected by G. Paoli. 

Spilococcus filicicola (Newstead) (comb, n.) 
(Text-fig. 20) 

Ripersia filicicola Newstead, 1898 : 96. 

Ripersia filicicola Newstead, Newstead, 1903 : 184. 

Habit and distribution. Newstead stated that in life this insect was pale ochreous 
yeUow or red-pink ; farinose, with broad irregular wax appendages on the margin 
of the abdominal segments. The insects also secrete slender iridescent filaments 
radiating from the sides of the body. 

Known only from a greenhouse in the Royal Botanic Gardens, Kew, England, 
on the fronds of Trichomanes spicatum, 27 . ii . 1897. It is possible that it will be found 
eventually in Central or South America. 

Recognition characters. A broadly oval species, length of available specimens 1-5 mm., 
anal lobes well developed. Antennae 6-segmented, 240 microns long. Legs normal, stout, 
trochanter -f- femuri 80 microns, tibia -f- tarsus 170 microns. Labium 105 microns long. Circulus 
absent. Ostioles well developed, inner edges of lips sclerotized, each lip with 0-2 setae and 
4-6 trilocular pores. Anal ring with a double band of pores and 6 setae about twice length of ring 
at its greatest diameter. Cerarii numbering 10 pairs, there being 2 pairs on head and 8 pairs on 
abdomen. Anal lobe cerarii each with a pair of stout conical setae and a small group of about 
7 or 8 trilocular pores accompanied by 6 auxiliary setae and a few other trilocular pores all 
on a large oval sclerotized plate the same area as anal ring. Anterior cerarii each with 3-7 trilocular 
pores and a pair of small conical setae except the frontal pair which have 3 conical setae, all 
without auxiliary setae. 

Dorsal surface sparsely beset with short slender setae. Trilocular pores evenly distributed, 
not numerous. Tubular ducts present, of oral rim type, rather large. Each duct often with 1-3 
trilocular pores and i or 2 setae immediately surrounding base of oral rim but not attached to it, 
although some ducts entirely without these. In the 2 specimens at hand there are 7 ducts 



THE BRITISH PSEUDOCOCCID AE 



55 




Fig. 20. Spilococcus fiUcicola (Newstead). 



56 D. J. WILLIAMS 

equispaced on either side of margin, 2 on mid-line of thorax and a pair in submedian areas of 
metathorax. 

Ventral surface witli small triangular sclerotized area on each anal lobe and an apical seta 
shorter than anal ring setae. Body setae few, short and slender but, for the most part, longer 
than dorsal setae. Trilocular pores evenly distributed but sparse. Multilocular disc pores in 
single transverse rows on median area of abdomen only. Tubular ducts with oral collar very 
few, in transverse rows on segments IV-VII. Tubular ducts with oral rim of 2 types. A single 
duct, slightly smaller than those on dorsum, but with rim of larger diameter than a multilocular 
disc pore, present on either side of segment VF. A smaller type of duct, each with a narrow rim 
of smaller diameter tlian a multilocular disc pore, present around the margins where they are 
more numerous on thorax. 

Notes. The placing of this species in Spilococcus is tentative as the dorsal sclero- 
tization on the anal lobes and the very large oral rim ducts are not found on other 
species in the genus. Although some of the oral rim ducts have one or two setae and 
some trilocular pores around the base of the rim, this character is not constant. 

SPINOCOCCUS Borchsenius 

Spinococcus Kiritchenko, 1931 : 314 (worn. nud.). 
Spinococcus Borchsenius, 1949 : 393. 

Type of genus Acanthococcus marrubii Kiritchenko. 

The main characters of this genus are the 17-18 pairs of cerarii with conical setae, 
these elevated slightly from the surrounding derm and with 2-10 trilocular pores 
immediately at the base. Dorsal setae similar in shape and size to cerarian setae, 
there being usually a row of dorsal cerarii on mid-line. Antennae 8-9-segmented. 
Claw with distinct denticle. Quinquelocular disc pores on ventral surface. 



Spinococcus calluneti (Lindinger) 
(Text-fig. 21) 

Pseudococcus calluneti Lindinger, 1912 : 90. 

Parapedronia calluneti (Lindinger), Balachowsky, 1954 [i953] ■ 230. 
Spinococcus calluneti (Lindinger), Zahradnik, 1959 : 537. 
Spinococcus calluneti (Lindinger), Danzig, i960 : 176. 

Habit and distribution. Apparently confined to Calluna spp. and Erica spp. 
where it feeds mainly on the roots. Described originally from Germany and found 
on two occasions only in Britain as follows : England. Cheshire : Chester, roots 
and flowers of Erica cinerea, Calluna sp., viii. 1922 (R. Newstead). Surrey : Oxshott, 
Calluna vulgaris roots, x.1925 (K. Mansour). 

Recognition characters. Adult female ovate about 1-5 mm. long. Anal lobes moderately 
developed, set well apart. Antennae 7-8-segmented, 245-320 microns long. Legs normal, claw 
with a distinct denticle. Trochanter + femur 170-180 microns, tibia + tarsus 190-200 microns. 
Labium 70-90 microns long. Circulus small and round within borders of segment IV. Ostioles 
present, inner edges of lips sclerotized, each lip usually with 3 trilocular pores and an occasional 
seta. Anal ring with a double band of pores and 6 setae about twice length of diameter of ring. 
Marginal cerarii numbering 18 pairs, the cerarian setae on small projections of the derm. Anal 
lobe cerarii each with 2 conical setae on a large sclerotized plate nearly same size as anal ring, 
containing about 10 trilocular pores and i or 2 minute setae. Anterior cerarii each with 2 smaller 



THE BRITISH PSEU DOCOCCI D AE 



57 




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O 



Fig. 21. Spinococcus calluneti (Lindinger). 



58 D. J. WILLIAMS 

conical setae set close together and with 2-4 trilocular pores immediately around the base, the 
base of each seta elevated from the surrounding derm. 

Dorsal surface with setae of various sizes but of similar shape to cerarian setae. Dorsal cerarii 
each consisting of 2 conical setae situated on mid-line of body, these setae of same size as those 
of marginal cerarii. Similar sized setae in transverse rows, each with i or at most 2 trilocular 
pores on or near the basal projection ; occasionally replaced by definite cerarii with 2 setae. 
Other dorsal setae very small. Multilocular disc pores present in transverse rows on thorax and 
abdomen and a few scattered on head. Tubular ducts with a similar distribution to multilocular 
disc pores, a single duct usually situated between 2 multilocular pores. Trilocular pores evenly 
distributed, not numerous. 

Ventral surface with apical setae longer than anal ring setae. Other setae slender, of various 
lengths but usually longer in median areas. Multilocular disc pores in transverse rows on abdomi- 
nal segments but forming groups towards margins. Marginal groups also present on thorax and 
I or 2 pores situated near each coxa. Quinquelocular disc pores present in median areas only on 
head and thorax and all prevulvar abdominal segments. Trilocular pores few, absent in median 
areas of anterior abdominal segments and thorax. Tubular ducts with oral collar of two sizes. 
A larger type, similar to dorsal ducts, around the margin, there being usually a single duct 
between two multilocular disc pores. A smaller type, about half the width, mainly concentrated 
in median areas of abdomen but some extending laterally among the larger type. An occasional 
smaller duct also present in median areas of thorax. 

Notes. The identity of this species is based on specimens from Erlangen, Germany, 
collected by H. Schmutterer. 

TRIONYMUS Berg 

Trionymus Berg, 1899 : 78. 

Trionymus Berg, Ferris, 1950 : 251, 1953 : 482. 

Trionymus Berg, McKenzie, i960 : 764. 

Type of genus Westwoodia perrisii Signoret. 

This genus has been a source of confusion in recent years. It is very close to 
Dysmicoccus and it is doubtful if the latter will be accepted when further species 
are described. Ferris laid particular stress on the shape of the circulus which, in 
Trionymus, should be small and round or oval whereas in Dysmicoccus it should be 
large and divided by an intersegmental fold. McKenzie has disregarded the shape of 
the circulus and species with 6 or more cerarii are placed in Dysmicoccus and those 
with 5 or less are placed in Trionymus. 

McKenzie's interpretation is accepted here although difficulties arise with T. 
newsteadi. This species possesses 4 or 5 pairs of cerarii and has close affinities with 
the genus Dysmicoccus with regard to body form and the shape of the circulus. 
Furthermore, its host is Fagus and, although there are exceptions, most species of 
Trionymus are found on grasses. As it has no obvious relationship to its present 
genus Pseudococcus, it is left in Trionymus for the time being. 

This is by far the largest British genus and the 9 species may be separated by the 
following key : 

1 Anal lobe cerarii borne on a distinct sclerotized plate as large as or larger than area 

of anal ring ............ 2 

- Anal lobe cerarii not borne on a sclerotized plate ...... 5 

2 (i) Circulus large, divided by an intersegmental fold .... tomlini Green 

- Circulus small, round or oval, not divided by an intersegmental fold. . . 3 



THE BRITISH PSEUDOCOCCID AE 



50 



3 (2) Multilocular disc pores absent on dorsum except for total of no more than 4-6 on 

some of last abdominal segments ...... dactylis Green 

- Multilocular disc pores on dorsum in transverse rows ..... 4 

4 (3) With 26-45 trilocular pores on the sclerotized plate on anal lobes ; tarsus about 

half length of tibia perrisii (Signoret) 

With 70-75 trilocular pores on the sclerotized plate on anal lobes ; tarsus about 

one-third as long as tibia phalaridis Green 

5 (i) With 4 or 5 pairs of cerarii ; circulus large and quadrate-shaped newsteadi (Green) 

- With I or 2 pairs of cerarii ; circulus small and round ..... 6 

6 (5) Tubular ducts with the oral collar nearly half total length of duct radicum (Newstead) 

- Tubular ducts with the oral collar very small ...... 7 

7 (6) Multilocular disc pores absent on dorsum ..... thulensis Green 

- Multilocular disc pores present on dorsum ....... 8 

8 (7) With a single pair of cerarii on anal lobes, cerarian conical setae slender orestes sp. n. 

- With 2 pairs of cerarii, cerarian conical setae on anal lobes short and stout 

diminutus (Leonardi) 

Trionymus dactylis Green 
(Text-fig. 22) 

Pseudococcus [Trionymus] dactylis Green, 1925 : 523. 
Trionymus dactylis Green, Green, 1928 : 22. 

Habit and distribution. Appearance in life not recorded. Known only from 
beneath the leaf sheaths of grasses. Channel Islands. Guernsey : St. Sampson, 
Dactylis glomerata, ix . 1924 (E. E. Green) (Type) ; Petit Bot Bay, Deschampsia 
caespitosa, ix.1927 (F. Laing). England. Somersetshire: Cheddar, Dactylis 
glomerata, viii.1926 (E, E. Green). 

Recognition characters. Body elongate elliptical attaining a length of 5 mm. Anal lobes 
moderately developed. Antennae 8-segmented, length 380-425 microns. Legs normal, slender, 
posterior coxae with numerous translucent pores, trochanter + femur 315-340 microns, tibia -f- 
tarsus 335-370 microns. Labium 75-80 microns long. Anterior and posterior ostioles present, 
each lip with 4-6 trilocular pores and usually without setae although at times one may be 
present. Circulus very small, oval. Anal ring with a double band of pores and 6 setae twice 
length of diameter of ring. Cerarii numbering 3 or 4 pairs. Anal lobe cerarii each with 2 conical 
setae on a sclerotized plate, larger in area than anal ring, containing 12 long setae and about 40 
trilocular pores. Penultimate cerarii each with 2 smaller cerarian setae accompanied by about 
12 trilocular pores and 2 or 3 auxiliary setae and surrounded by a small sclerotized area. Cerarii 
on segment VII similar but without sclerotized area. Cerarii on segment VI, when present, 
with I or 2 small cerarian setae and 2 or 3 trilocular pores. 

Dorsal surface with slender setae of various sizes. Multilocular disc pores either absent entirely 
or numbering only 3-4 at most on posterior abdominal segments. Trilocular pores numerous. 
Tubular ducts with oral collar of 2 sizes. A larger type, quite numerous on all segments, tending 
to form transverse rows on abdominal segments but becoming scattered on thorax and head. 
A smaller type present on abdomen, sparse, in transverse rows in middle of segments and occa- 
sionally on thorax. 

Ventral surface of body with apical setae longer than anal ring setae. Other setae of various 
sizes and similar to dorsal setae. Multilocular disc pores on fifth and posterior segments only, 
except for an occasional pore anteriorly. Tubular ducts of same 2 sizes as those on dorsum. 
A larger type in transverse groups or rows on fifth and posterior segments and forming rather 
large groups laterally. Marginal or submarginal groups also present as far forward as head. 
Occasionally single ducts may be present in median areas of thorax. A smaller type of duct 



6o 



D. J. WILLIAMS 




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Fig. 22. Trionymus dactylis Green. 



THE BRITISH PSE UDOCOCCI D AE 61 

mainly present on abdomen across the middle of the segments, much fewer than larger type. 
Trilocular pores with an even distribution. 

Trionymus diminutus (Leonard!) 

Pseudococcus diminutus Leonardi, 191 8 : 198. 
Trionymus diminutus (Leonardi), Ferris, 1950 : 261. 

Specimens are at hand from Channel Islands, Jersey (G. Fox-Wilson) on Phor- 
mium tenax to which plant it seems to be confined. It was described from Italy and 
is now known from New Zealand, U.S.A., Formosa and Russia. Ferris has redescribed 
and illustrated this species. 

Trionymus newsteadi (Green) 
(Text-fig. 23) 

Pseudococcus newsteadi Green, 1917 : 265. 
Trionymus newsteadi (Green), Zahradnik, 1959 : 538. 

Habit and distribution. Described as " Body pale purphsh gre}' ; limbs and 
antennae pale stramineous ; venter thinly, dorsum rather thickly and evenly 
covered with white mealy secretion ; terminal three or four segments of abdomen 
with short, stout, waxy tassels." It is known only from beech. England. Surrey : 
Camberley, vii.1916, v.1917 (Type), 15. vi. 1932 (E. E. Green). Buckinghamshire : 
Chartbridge (E. J. Joseph). Hertfordshire: Berkhamsted, 16.1.1913. Essex: Ep- 
ping Forest, 25. v. 1923 (C. L. Withycombe). Sussex : Midhurst, 18. v. 1946 (W. J. 
Hall). Berkshire : Silwood Park, 31. v. 1949 (K. L. Boratynski). 

Recognition characters. Body broadly oval attaining a length of 3 mm., anal lobes 
moderately developed. Antennae 8-segmented, 340-410 microns long. Legs normal, posterior 
coxae and tibiae with a few translucent pores. Trochanter + femur 260-285 microns, tibia + 
tarsus 275-305 microns. Labium 105-120 microns long. Circulus large, posterior edge usually 
shorter than anterior edge. Posterior ostioles present only, each lip with 3 or 4 trilocular pores 
and apparently without setae. Anal ring with 2 rows of pores and 6 setae, these about twice 
length of diameter of ring. Cerarii numbering 4 or 5 pairs on abdomen. Anal lobe cerarii each 
with 2 slender conical setae and about 6 or 7 trilocular pores accompanied by 3 or 4 auxiliary 
setae, the area around the base of the setae sclerotized. Anterior cerarii on segments VI-VIIl 
similar but cerarian setae smaller, auxiliary setae absent. Cerarii on fifth segment usually with 
setae similar to body setae and accompanied by about 3 trilocular pores. 

Dorsal surface with slender setae. Tubular ducts numerous with deep collar occupying nearly 
one-third length of duct ; a flange-shaped membranous ring also arising from the inner end of 
the collar. The ducts and dorsal setae are arranged in definite groups rather than in transverse 
rows leaving bare areas except for trilocular pores which are more evenly distributed, as in the 
accompanying diagram. 

Ventral surface with apical setae longer than anal ring setae. Body setae slender but mainly 
longer than those on dorsum. Multilocular disc pores confined to abdomen except for occasional 
pores near the coxae. Fifth segment with about 10-18 pores and posterior segments with trans- 
verse rows at anterior and posterior edges of segments. Tubular ducts of 2 sizes, a larger type 
similar to those on dorsum, rather numerous in marginal groups on abdomen and less numerous 
in groups around the thorax and head. Present also in transverse rows on fourth and posterior 
segments. A smaller type of duct, similar in all respects to larger type, sparsely distributed in 
median areas of posterior abdominal segments and occasionally on abdominal margins. Trilocular 
pores not numerous. 



62 



D. J. WILLIAMS 




^ 






Fig. 23. Trionymus newsteadi (Green) 



THE BRITISH PSE U DOCOCCID AE 63 

Trionynius orestes sp. n. 

(Text-fig. 24) 

Trionymus pulverarius (Newstead), Green, 1928 : 30. (Misidentification.) 

Habit and distribution. External appearance not known : probably found 
on the leaf sheaths. England. Surrey : Box Hill, Brachypodium sylvaticum, 
26.ix.1921 (E. E. Green) (Holotype). Somersetshire: Cheddar, Brachypodium 
sylvaticum, viii.1926 (E. E. Green). 

Recognition characters. An elongate species attaining a length of 3-5 mm. in available 
specimens. Anal lobes poorly developed. Antennae 7-segmented, 275-310 microns long. Legs 
rather small and slender, posterior coxae with a few translucent pores. Trochanter + femur 
200-230 microns, tibia -f tarsus 230-260 microns. Labium 75-80 microns long. Ostioles small, 
each lip with about 3 trilocular pores and without setae. Circulus present, small and oval. 
Anal ring with a double band of pores and 6 setae, about twice as long as diameter of ring. 
Cerarii on anal lobes only, each with 2 conical setae set close together and about 5 trilocular pores 
accompanied by a longer auxiliary seta. 

Dorsal surface with slender setae of various sizes, not numerous. Multilocuiar disc pores 
arranged more or less at anterior and posterior edges of abdominal segments in single or double 
transverse rows becoming less numerous anteriorly. A few also present in no definite arrangement 
on thorax and head. Trilocular pores evenly distributed. Tubular ducts with oral collar of 2 
types, a larger type in transverse rows on abdomen except on segments IX + X ; present also 
on thorax and head where they have a random distribution. A smaller type, sparse, situated 
across the middle of abdominal segments except the last. 

Ventral surface with apical setae longer than anal ring setae. Other ventral setae similar to 
those on dorsum but tending to be longer in median areas. MultUocular disc pores on posterior 
edges of segments V-VIII and on anterior edges of segments VI-VIII, present also on segments 
IX + X. Small marginal groups extending forward to head. Tubular ducts of two sizes as on 
dorsum. A larger type present mainly in marginal groups around body and a smaller type in 
transverse rows on abdomen and sometimes a few on thorax. Trilocular pores not numerous. 

Notes. This species is very close to 7\ caricis McConnell described from Carex 
tribuloides, Maryland, U.S.A., but differs in the greater number of dorsal multilocuiar 
disc pores which tend to form transverse rows on the abdominal segments. In T. 
caricis they are sparse and scattered. 

Trionymus perrisii (Signoret) 

(Text-fig. 25) 

Westwoodia perrisii Signoret, 1875 : 337. 

Dactylopius hibernicus Newstead, 1895 : 167 (syn. n.). 

Trionymus perrisii (Signoret), Berg, 1899 : 78. 

Dactylopius hibernicus Newstead, Newstead, 1903 : 172 (in part). 

Dactylopius pulverarius (Newstead), Newstead 1903 : 174 (in part). 

Pseudococcus pulverarius (Newstead), Green, 1915 : 178. (Misidentification.) 

Pseudococcus hibernicus (Newstead), Green, 1920 : 120. 

Trionymus pulverarius (Newstead), Green, 1926 : 182. (Misidentification.) 

Trionymus hibernicus (Newstead), Green, 1926 : 183. 

Trionymus pulverarius (Newstead), Green, 1928 : 30. (Misidentification.) 

Habit and distribution. Living at the base of the leaf sheath of grasses. Adult 
dull yellow to pale purplish, dusted with a fine white powder. Short tassels on last 



64 



^ 



D. J. WILLIAMS 



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Fig. 24. Trionynius orestes sp. n. 



THE BRITISH PSEU DOCOCCID AE 65 

two abdominal segments. This is, apparently, a common species in Britain and 
material has been examined from the following localities : 

England. Berkshire: Silwood Park, Deschampsia caespitosa, Holcus sp.. 27.x. 
1948, Festuca rubra, 22. ix. 1948, 13. ix. 1953 (K. L. Boratynski). Cheshire : Helsby 
Hill, Agrostis sp., i.viii.1896 (K. Newstead). Cumberland: Whinlatter Pass, 
Deschampsia caespitosa 10.viii.1960 (D. J. Williams). Devonshire: Budleigh 
Salterton, 10. ix. 1896 (E. E. Green). Durham : Waldridge Fell, 7.viii.i96o (D. J. 
Williams). Kent: Deal, ix. 1899 (B. Tomlin). Norfolk: Ingoldisthorpe, viii . 1892, 
(R. Newstead). Somersetshire: Cheddar, viii. 1926 (E. E. Green). Surrey: Guild- 
ford, 21.vii.1922 (E. E. Green) ; Camberley, ix.1914, 19. ix. 1928, 26. ix. 1930 
(E. E. Green). Yorkshire : Selby, ix . 1918 (E. E. Green). Hawkesworth, 15 . viii . 1958, 
6. viii. i960 (D. J. Williams). Scotland. East Lothian: Gullane, Aberlady, North 
Berwick Law, viii. 1925 (E. E. Green). Ireland. Antrim: BaUintoy, 3.ix.i893 
(L. E. Tomlin) {hibernicus type). 

Recognition characters. Adult female elongate oval, up to 4-5 mm. in length. Anal lobes 
usually well developed. Antennae 8-segmented, 335-340 microns long. Legs normal, posterior 
coxae with translucent pores, trochanter + femur 215-260 microns, tibia + tarsus 250-285 
microns. Labium about 95 microns long. Anterior and posterior ostioles moderately developed, 
each lip with 4-6 trilocular pores and rarely with setae. Circulus present, small and oval between 
fourth and fifth segments. Occasionally a second smaller circulus may be present on the next 
posterior segment and very rarely up to 4 circuli present in which case the extra ones are minute. 
Anal ring with 2 rows of pores and 6 setae twice as long as diameter of ring. Cerarii numbering 
2 or 3 pairs. Anal lobe cerarii each with 2 conical setae on a sclerotized plate, slightly larger in 
area than anal ring. This plate also containing about 26-45 trilocular pores and 9 auxiliary 
setae. Cerarii on segment VIII each with a pair of smaller conical setae and on a small sclerotized 
plate with 8 trilocular pores and about 2 auxiliary setae. On segment VII cerarii often absent 
entirely but when present, with either i or 2 conical setae and 2 or 3 trilocular pores and not 
surrounded by a sclerotized area. 

Dorsal surface with slender setae. Multilocular disc pores variable in number and distribution. 
On the abdomen they form transverse rows at the anterior and jiosterior edges of segments 
VI-VIII. On the anterior abdominal segments they become sparse and may be missing entirely 
on some of the edges. On the thorax they become scattered and sometimes they are present on 
the head. Trilocular pores rather numerous. Tubular ducts with oral collar of 2 sizes. A smaller- 
sized duct sparse, there being a few across the middle of most abdominal and thoracic .segments. 
A larger type of duct, numerous, forming transverse bands on the abdominal segments. In some 
S|>ecimens they reach from the anterior to posterior edges of most segments but in other specimens 
they are concentrated in the middle of the segments or towards the posterior edges. On the 
thorax and head they are less numerous and become scattered. 

Ventral surface with apical setae longer than anal ring setae. Ventral setae slender but 
tending to be longer than dorsal setae. Multilocular disc pores numerous in transverse rows at 
anterior and posterior edges of segments VI-VIII ; present also on segments IX + X and on 
posterior edge of segment V. Elsewhere on the abdomen they are sparse and may be present or 
absent around the margins. On the thorax and head they are scattered on median areas and 
often extend around margins. Trilocular pores evenly distributed. Tubular ducts, of smaller 
type, present in middle of abdominal segments and in median areas of thorax ; not numerous. 
A larger type in transverse rows and marginal groups on abdominal segments and extending 
around body in a marginal or submarginal zone. 

Notes. Dr. A. Reyne has very kindly sent specimens of T. perrisii from Holland 
and these agree with specimens of T. hibernicus described from Britain. Professor 
N. S. Borchsenius has also kindly examined some British specimens of T. hibernicus 

ENTOM. 12, I. 5 



66 



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D. J. WILLIAMS 



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Fig. 25. Trionymus perrisii (Signoret). 



THE BRITISH PSE U DOCOCCI D AE 67 

and it is his opinion that they are the same as T. perrisii. All of these specimens 
agree with the illustration of T. perrisii given by Marchal (1908). 

Continental specimens usuall}^ have 3 distinct pairs of cerarii but on at least i 
specimen from Holland there are only 2 pairs. British specimens usually have 2 
pairs of cerarii but the third pair is often represented by a single conical seta and 
2 or 3 trilocular pores. Although a single circulus seems to be the more normal 
condition, some specimens in Britain have 2 and in rare cases up to 4. 

Trionytnus phalaridis Green 

(Text-fig. 26) 

Pseudococcus {Trionytnus) phalaridis Green, 1925a : 37. 

Habit and distribution. Described as " Colour reddish purple to dark slaty 
grey ; closely dusted with white powdery secretion. Posterior extremity with four 
short waxy tassels." Living beneath the leaf sheath and stem. England. Surrey : 
Frimley, 8.ix.i922 (E. E. Green) (Type) ; Camberley, 30.viii.1921, 30.ix.1921 
(E. E. Green), all on Phalaris arundinacea. Gloucestershire : ix.1922 (K. G. Blair). 

Recognition characters. Body elongate, sides subparallel, attaining a length of 6-5 mm. 
Anal lobes well developed. Antennae 8-segmented, 450-495 microns long. Legs rather long and 
slender, posterior coxae with a few translucent pores, trochanter + femur 380-395 microns, 
tibia about 3 times as long as tarsus, measuring together 405-445 microns. Labium 100 microns 
long. Ostioles present, usually with about 10-15 trilocular pores on each lip of posterior pair and 
about 4-6 on each lip of anterior pair ; without setae. Circulus oval. Anal ring with a double 
band of pores and 6 setae over twice length of ring at its greatest diameter. Cerarii on last 2 
segments only. Anal lobe cerarii each with 2 conical setae on a large sclerotized plate, larger in 
area than anal ring and accompanied by 70-75 trilocular pores and 12 or 13 slender auxiliary 
setae. Penultimate cerarii each with 2 smaller conical setae on a small round sclerotized plate 
containing about 15 trilocular pores and 5 auxiliary setae. 

Dorsal surface of body with numerous slender setae of various sizes. Multilocular disc pores 
arranged in transverse rows at anterior edges of segments V-VIII and with only a few at posterior 
edges of these segments. On the anterior abdominal segments and thorax they are sparse and 
become scattered. Trilocular pores numerous and evenly distributed. Tubular ducts with oral 
collar of 2 sizes. A smaller type, few on abdomen, situated mainly across middle of segments. 
A larger type abundant, occupying most of the area of segments II-VIII except at anterior 
edges. On the thorax they become scattered but are well separated between the segments and 
on the head they lie in definite groups. 

Ventral surface with a pair of apical setae longer than anal ring setae. Slightly shorter setae 
present around margins of abdomen. Other ventral setae of various sizes but all slender. Multi- 
locular disc pores numerous on segments IX + X, in transverse rows at anterior and posterior 
edges of segments VI-VIII, and on posterior edges of segment V. A few also present on segment 
IV, around the margins to head and occasionally in median areas of thorax. Trilocular pores 
numerous. Tubular ducts of same 2 sizes as on dorsum. A smaller type sparse on abdomen and 
a larger type in transverse rows and marginal groups on the abdominal segments. Present also 
in groups around the margins to head and a few in median areas of thorax. 

Notes. This species comes close to T. perrisii but is usually much larger. The 
trilocular pores on the sclerotized plate on anal lobes are much more numerous and 
the tibia is about three times as long as the tarsus whereas in T. perrisii it is only 
twice as long. 



68 



D. J. WILLIAMS 



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l-'iG. 20. Trionymtis phalaridis Green. 



THE BRITISH PSEUDOCOCCID AE 69 

Trionymus radicum (Newstead) (comb, n.) 

(Text-fig. 27) 

Dactylopius radicum Newstead, 18956 : 235. 

Dactylopius hibernicus Newstead, Newstead, 1903 : 172 (in part, misidentification). 

Ripersia donisthorpei Newstead, In Donisthorpe, 1907 : 5 (syn. n.). 

Habit and distribution. Living at the base of grasses. Described originally as 
" pale reddish-pink ... At period of gestation the $ completely envelops herself 
in a very loose cottony material, in which the pale flesh-coloured ova are laid." 
England. Isle of Man: Port Erin, 21.ix.1918. Buckinghamshire: Waddesdon, 
17. V. 1953 (V. F. Eastop). Kent : Charing, in nest of Ponera contracta (H. Donis- 
thorpe) (^om's^^or/)^/ Type) . Wales. Anglesey: Puffin Is., vi. 1895 (ra^?icww Type). 

Recognition characters. Body elongate oval, about 3*5 mm. long. Anal lobes moderately 
developed. Antennae 7-segmented, 305-335 microns long. Legs normal, slender, hind coxae 
with a few translucent pores. Trochanter + femur 215-240 microns, tibia + tarsus 225-245 
microns. Labium 70-75 microns long. Ostioles poorly developed with 4-7 trilocular pores on 
each lip and rarely with setae. Circulus very small, oval. Anal ring with a double band of pores 
and 6 setae longer than ring at its greatest diameter. Cerarii numbering 2 pairs only. Anal lobe 
cerarii each with a pair of conical setae and about 6 or 7 trilocular pores accompanied by a single 
auxiliary seta, the area between the cerarian setae sclerotized. Cerarii on segment VIII each 
with a pair of smaller conical setae and about 4 trilocular pores. 

Dorsal surface of body with short slender setae, not numerous. Multilocular disc pores very 
few, scattered at random on thorax and abdomen. Trilocular pores evenly distributed. Tubular 
ducts arranged in transverse rows on thorax and abdomen and in groups on head, each duct with 
a large flange-shaped collar occupying nearly half total length of duct. 

Ventral surface with apical setae longer than anal ring setae. Ventral setae all slender but 
mainly longer than dorsal setae. Multilocular disc pores numerous on segments IX + X and 
present in transverse rows at posterior edges and usually on anterior edges of segments V-VIII. 
Small numbers also on anterior abdominal segments and around the margins to head and one or 
two in median areas of thorax. Tubular ducts of 2 sizes ; a larger type, similar to dorsal ducts 
situated in groups around the margins and a few present in median areas of segments V-VIII. 
A smaller type of duct, similar to the larger type but with inner end rounded, present on abdomen 
mainly in the median areas. 

Notes. When Newstead collected this species he also found some smaller specimens 
which he thought were the larvae. They were, however, adult females of Rhizoecus 
halophilus and he mentioned this error in 1903. In this same work he sunk the name 
Dactylopius radicum as a synonjmi of D. hibernicus, stating that the " curiously 
shaped pores or glands " were caused by a parasite. The species is quite distinct and 
the tubular ducts, with the collar flange-shaped and nearly half the total length 
of the duct, serve to distinguish this species from all the other British species of 
Trionymus. 

Trionymus thulensis Green 

(Text-fig. 28) 

Trionymus pulverarius (Newstead), Green, 1926 : 182. (Misidentification.) 
Trionymus thulensis Green, 1931a : 267. 

Habit and distribution. Found beneath the leaf sheath and stem of grasses. 
Described originally from Iceland, VarmahliS" on Festuca rubra, 3.vi.i929 (C. H. 
Lindroth). British specimens are at hand from the following localities : England. 



70 



D. J. WILLIAMS 



^ 





.* • * • ■ . • • • • ' . 



'...;\V^-;-.V:V.-V.H 

■ ;i-.V-\- ■/.■.:.■:.•.:• 



•'\ ' 



•V- 



•.v> 



d 

o 



^ 




Fig. 27. THonymus radicum (Newstead). 



THE BRITISH PSEUDOCOCCI D AE 



71 



^ 





'& 



o 



Fig. 28. Trionymus thulensis Green. 



72 D. J. WILLIAMS 

Berkshire : Silwood Park, Holcus mollis, 31.V.1951 (K. L. Boratynski). Durham : 
Low Fell, vi.1943 (R. S. Bagnall). Kent : Thurnham, Trisetum flavescens, I3.vii. 
1921 (E. E. Green). Isle of Wight : Seaview, 3.ix.i92i (E. E. Green). Norfolk: 
Snettisham, 18.vii.1895 (R. Newstead). Northumberland: Alnmouth, 2.iii.i935 
(R. S. Bagnall). Surrey : Camberley, 5.ix.i923, vii.1929, 23.vii.1932 (E. E. Green). 
Yorkshire: Askham Bog, Agrostis tenuis, 22. vi. 1956 (D. J. Williams). Wales. 
Brecknockshire : Llangammarch, vi.1926 (E. Earle). 

Recognition characters. Body elongate elliptical to elongate oval, attaining a length of 
4 mm. Antennae 7- or 8-segmented, length about 380 microns. Labium 80-95 microns long. 
Legs normal, posterior coxae with a few translucent pores ; trochanter + femur 265-285 
microns, tibia + tarsus 285-320 microns. Circulus small and round. Ostioles with about 6 
trilocular pores on each lip and without setae. Anal ring with 6 setae twice as long as diameter 
of ring. Cerarii present on last 2 abdominal segments. Anal lobe cerarii each with a pair of 
conical setae, a small cluster of 7 or 8 trilocular pores, not on a sclerotized plate but small area 
between the setae sclerotized. Accompanied also by a single auxiliary seta immediately above 
cerarian setae. Cerarii on segment VIII with 2 smaller conical setae and about 6 or 7 trilocular 
pores and no auxiliary setae. 

Dorsal surface of body with short slender setae. Multilocular disc pores absent. Trilocular 
pores not numerous, evenly distributed. Tubular ducts with oral collar of one size ; absent on 
segments IX + X but present on all anterior abdominal segments and thorax in transverse bands 
and tending to become scattered on head. In some specimens they are quite sparse on thorax 
and head. 

Ventral surface of body with a stout pair of apical setae longer than anal ring setae. Other 
ventral setae slender but mainly longer than on dorsum ; not numerous. Multilocular disc pores 
present on segments IX -f X and in transverse rows at posterior edges of segments V-VIII in 
median areas. Present also at anterior edges of segments VI-VIII and occasionally in median 
areas of anterior abdominal segments, thorax and head. Tubular ducts of 2 sizes, a larger type, 
similar to dorsal ducts, numerous in groups around margins to head and also in transverse rows 
on abdominal segments. Some also present in median areas of thorax. A smaller type of duct 
less numerous, situated in median areas of abdominal segments and sometimes among the lateral 
groups of the larger ducts. 

Notes. This species is, apparently rather common in Britain, and has been misi- 
dentified in the British literature as T. pulverarius (Newstead) . The illustration has 
been prepared from specimens collected at Askham Bog, Yorkshire, which agree 
well with the type. Some other specimens studied show fewer multilocular disc pores 
and tubular ducts but these represent an extreme in variation. 

Trionymus tomlini Green 
(Text-fig. 29) 

Pseudococcus [Trionymus) calif ornicus Ehrhorn, Green, 1925 : 522. (Misidentification.) 
Pseudococcus (Trionymus) tomlini Green, 1925 : 525. 

Habit and distribution. External appearance not known. Found beneath the 
leaf sheaths of grasses. Channel Islands. Guernsey : Port Grat, Psamma arenaria 
{=Ammophila arenaria), ix. 1925 (J. R. le B. TomHn) (Type) ; Vazon Bay, Agropyron 
repens, ix.ig24. England. Isle of Man : Dalby in sandy shingle above high water 
mark ix.1922 (L. E. Tomlin). 

Recognition characters. An elongate oval species measuring about 3'5 mm. long. Anal 
lobes moderately developed. Antennae 8-segmented, 425 microns long. Legs rather long. 



THE BRITISH PS EU DOCOCCI D A E 



73 



^ 




Fig. 29. Trionymus tomlini Green. 



74 D. J. WILLIAMS 

posterior coxae with a few translucent pores. Trochanter + femur 285-340 microns, tibia + 
tarsus 355-370 microns. Labium about 95 microns long. Anterior and posterior ostioles present, 
each lip with 6-8 trilocular pores, usually without setae but occasionally present. Circulus large 
and oval with a distinct intersegmental fold. Anal ring with 6 setae about twice length of its 
diameter. Cerarii numbering 2 pairs. Anal lobe cerarii each with 2 conical setae on an oval 
sclerotized plate nearly as large as anal ring and containing about 35-42 trilocular pores and 8 or g 
auxiliary setae. Penultimate cerarii not on a sclerotized area, each with 2 smaller conical setae, 
about 15 trilocular pores and 2 or 3 auxiliary setae. 

Dorsal surface with slender setae of various sizes. Multilocular disc pores absent on segments 
IX -}- X and present at anterior and posterior edges of the other abdominal segments in no 
definite pattern ; not numerous and becoming sparse on anterior abdominal segments ; a few 
also present on thorax. Trilocular pores evenly distributed. Tubular ducts with oral collar of 
2 sizes. A smaller type, not numerous, in middle of abdominal segments except IX + X and 
also present on thorax. A larger type of duct more numerous, arranged on segments II-VIII 
mainly in transverse rows on posterior edges and in marginal groups, but a few also present 
towards anterior edge of each segment. On the meta- and mesothorax they become scattered 
and on the prothorax and head form definite groups. 

Ventral surface with apical setae longer than anal ring setae ; each anal lobe also with a small 
sclerotized patch which is not attached to the apical seta. Body setae rather numerous, all 
slender, of various sizes but mainly long. Multilocular disc p>ores numerous on abdomen, present 
on segments IX -f X and in transverse rows at posterior edges of segments IV-VIII. Situated 
also on anterior edges of segments VI-VIII and a few present on anterior abdominal segments 
and in median areas of thorax and head. Tubular ducts of same two types as on dorsum. A 
smaller type on abdomen, mainly across the middle of each segment. A larger type present in 
transverse rows at posterior edges of abdominal segments and in marginal groups which extend 
to head. Groups also present in median areais of thorax and head. 

Notes. In its general distribution of pores and ducts, this species comes very close 
to T. perrisii but differs in the form of the circulus which is quite large and divided 
by an intersegmental fold, whereas the circulus of T. perrisii is small and oval. It 
seems to have close affinities with T. modocencis (Ferris) and T. haancheni McKenzie, 
described from the U.S.A. which also have 2 pairs of cerarii and a large divided 
circulus. In this respect T. tomlini comes closest to T. haancheni but possesses more 
numerous multilocular disc pores on the dorsum, and on the venter they extend to 
the thorax and head whilst in T. haancheni they are present on the abdomen only. 

REFERENCES 

Balachowsky, A. S. 1934. Contribution k l'6tude des Coccides de France ( i je note) . Recherches 

compl6mentaires sur la faune de Corse. Bull. Soc. ent. Ft. 39 : 67-72. 
1948. Les Cochenilles de France, d'Europe, du nord de I'Afrique, et du Bassin M6diter- 

ran6en. IV. — Monographic des Coccoidea ; Classification — Diaspidinae (Premifere Partie). 

AcUialitSs sci. industr. 1054 : 243-394. 
1954 [1953]- Sur un Pseudococcini (Hom. Coccoidea) nouveau du Cap Bon (Tunisie). 

II. Presence de Rhizaspidiotus bivalvatus Goux (Coccoidea, Diaspidinae) en Afrique du 

Nord. Bull. Soc. not. Tunis. 6 : 227-253. 
Berg, C. 1899. Substitution de nombres genericos. III. Comun. Mus. nac. B. Aires, 1 : 77-80. 
BoRCHSENius, N. S. 1948. Toward a revision of the genus Phenacoccus Ckll. (Insecta, Homop- 

tera, Coccoidea). C.R. Acad. Sci. U.R.S.S. 61 : 953-956. 

1949. Fauna U.S.S.R. Pseudococcidae. Zool. Inst. Acad. Sci. (n.s. 38) 7, 383 pp. 

BoRCHSENius, N. S. & Ter-Gregorian, M. a. 1956. Coccoidea injurious to wheat and other 

plants in Armenia. Izv. Akad. Nauk Arm. S.S.R., Erevan, 9 : 17-19. 



THE BRITISH PSEUDOCOCCI D AE 75 

Brain, C. K. 1912. Contribution to the knowledge of mealy bugs, genus Pseudococcus, in 

the vicinity of Cape Town, South Africa. Ann. ent. Soc. Amer. 5 : 177-189. 
CocKERELL, T. D. A. 1893. Notes on the genus Pseudococcus Westwood. Ent. News, 4 : 317, 

318. 

1900. Some Coccidae quarantined at San Francisco. Psyche, 9 : 70-72. 

Danzig, E. M. i960. Some new and little known mealybugs (Homoptera, Coccoidea, Pseudo- 

coccidae) of the Leningrad Region. Rev. Ent. U.R.S.S. 39 : 172-181. 
De Lotto, G. 1958. The Pseudococcidae (Hom. : Coccoidea) described by C. K. Brain from 

South Africa. Bull. Brit. Mus. {nat. Hist.) Ent. 7 : 77-120. 
DoNiSTHORPE, H. St. J. K. 1907. Myrmecophilous notes for 1906. Ent. Rec. 19 : 4-7. 
Douglas, J. W. 1888. Notes on some British and exotic Coccidae (No. 10). Ent. mon. Mag. 

25 : 86-89. 

1888a. Notes on some British and exotic Coccidae (No. 11). Ibid. 25 : 124, 125. 

1889. Notes on some British and exotic Coccidae (No. 14). Ibid. 25 : 314-317. 

1890. Notes on some British and exotic Coccidae (No. 16). Ibid. 26 : 153-155. 

EssiG, E. O. 1942. College Entomology, New York. pp. vii -(- 900. 

Ezzatt, Y. M. & McCoNNELL, H. S. 1956. The Mealybug Tribe Planococcini (Pseudococcidae, 

Homoptera). Bull. Md. agric. Exp. Sta. A-84. 
Fernald, M. E. 1903. A Catalogue of the Coccidae of the World. 360 pp. 
Ferris, G. F. 1918. The California species of mealybugs. Stanf. Univ. Publ. biol. Sci. 78 pp. 

1950, 1953. Atlas of the Scale Insects of North America, 5, 6. 

Goux, L. 1934. Notes sur les Coccides (Hem.) de la France. (9e note). Contribution k I'dtude 
du genre Heliococcus avec description de deux especes nouvelles. Bull. Soc. ent. Fr. 39 : 

164-171. 
194 1. Contribution k I'^tude d'un faisceau d'esp^ces constituent un sous-genre nouveau 

du genre Pseudococcus (Hem. -Coccidae). Bull. Mus. Hist. nat. Marseille, 1 : 66-83. 
19410. Description d'un Rhizoecus nouveau et de sa larve neonate (!). Ibid. 1 : 197- 

203. 
1942. Notes sur les Coccides (Hem. Coccoidea) de la France. (32me note). Description 

d'un Phenacoccus et d'un Rhizoecus nouveaux. Ibid. 2 : 33-45. 
Green, E. E. 1915. Observations on British Coccidae in 1914, with descriptions of new species. 

Ent. mon. Mag. 51 : 175-185. 
191 6. On two new British Coccidae. with notes on some other British species. Ibid. 52 : 

23-31- 
1917. Observations on British Coccidae ; with descriptions of new species. Ibid. 53 : 

201-210, 260-264, 265-269. 

1920. Observations on British Coccidae. No. V. Ibid. 56 : 114-130. 

192 1. Observations on British Coccidae, with descriptions of new species. No. VI. Ibid. 

57 : 146-152, 189-200. 

1923. Observations on British Coccidae. — VIII. Ibid. 59 : 211-218. 

1924. On some new species of Coccidae from various sources. Bull. ent. Res. 15 : 41-48. 

1925. Notes on the Coccidae of Guernsey (Channel Islands) with descriptions of some new 

sp)ecies. Ann. Mag. nat. Hist. (9) 16 : 516-527. 

1925a. Observations on British Coccidae. — IX. Ent. mon. Mag. 61 : 34-44. 

1926. Observations on British Coccidae. — X. Ibid. 62 : 172-183. 

1928. Observations on British Coccidae. — XI. With descriptions of new species. Ibid. 

64 : 20-31. 

1930. Observations on British Coccidae. — XII. Ibid. 66 : 9-17. 

1931. Observations on British Coccidae. — XIII. Ibid. 67 : 99-106. 

1931a- Notes on some Coccidae from Iceland. Ent. Tidskr. 52 : 263-269. 

1934. Observations on British Coccidae. — XIV. Ent. mon. Mag. 70 : 108-114. 

Hambleton, E. J. 1946. Studies of hypogeic mealybugs. Rev. Ent., Rio de J. 17 : 1-77. 

1946a. A new name for a mealybug. Proc. biol. Soc. Wash. 59 : 177. 

Hardy, J. 1868. Notes for 1864. Hist. Berwicksh. Nat. CI. 5 : 136, 137. 



76 D. J. WILLIAMS 

KiRiTCHENKO, A. N. 1 93 1. Second contribution to the Coccid fauna of Ukraine and the 

Crimea. PI. Prot. Leningr. 7 : 307-321. 
KiJNCKEL d'Herculais, J. 1878. Histoire de la cochenille vivant sur las racines des palmiers 

de la section des Seaforthia. Expos6 des caractferes du genre Rhizoecus. Ann. Soc. ent. Fr. 

(5) 8 : 161-164. 
Leonardi, G. 1918. Terza contribuzione alia conoscenza delle Cocciniglie Italiane. Boll. 

Lab. Zool. Portici. 12 : 188-216. 
LiNDiNGER, L. 1912. Die Schildlduse (Coccidae) Enropas, Nordafrikas, und Vorderasiens, 

einschliesslich der Azoren, der Kanuren und Madeiras. Stuttgart. 388 pp. 

1934. Die Schildlaus-Arten P. Fr. Bouch^s und ihre Deutung. Eni. Jb. 43 : 162. 

Linnaeus, C. 1766. Insecta Hemiptera. Coccus. Systema Nat {Ed. 12) 1 : 739-743. 
McKenzie, H. L. i960. Taxonomic Study of California Mealybugs with descriptions of new 

species. (Homoptera : Coccoidea : Pseudococcidae.) Hilgardia, 29 : 681-770. 
Marchal, p. 1908. Notes sur les Cochenilles de I'Europe et du nord de I'Afrique. Ann. Soc. ent. 

Fr. 77 : 223-309. 
Maskell, W. M. 1893. Further Coccid notes, with descriptions of new species from Australia, 

India, Sandwich Islands, Demarara, and South Pacific. Trans. N.Z. Inst. (1892) 25 : 201- 

252. 
Morrison, H. 1945. The mealybug genus Heterococcus Ferris and some of its relatives. 

/. Wash. Acad. Sci. 35 : 38-55. 
Newstead, R. 1891 . On some new or little known Coccidae found in England. Eni. mon. Mag. 

27 : 164-166. 

1892. On new or little known Coccidae, chiefly English (No. 2). Ibid. 28 : 141-147. 

1893. New or little known Coccidae, chiefly English (No. 3). Ibid. 29 : 77-79. 

1895. Observations on Coccidae (No. 11). Ibid. 31 : 165-167. 

1895a. Observations on Coccidae (No. 12). Ibid. 31 : 213, 214. 

18956. Observations on Coccidae (No. 13). Ibid. 31 : 233-236. 

1897. Observations on Coccidae (No. 16). Ibid. 33 : 165-171. 

1898. Observations on Coccidae (No. 17). Ibid. 34 : 92-99. 

1901. Observations on Coccidae (No. 19). Ibid. 37 : 81-86. 

1903. Monograph of the Coccidae of the British Isles, Ray Society, 2 : 162-195. 

Rau, G. J. 1937. Two apparently undescribed mealybugs (Hemiptera Pseudococcidae) 

from New York State. Bull. Brooklyn ent. Soc. 32 : 195-201. 
Reyne, a. 195 1. A re-description of Ripersia corynephori Sign. Tijdschr. Ent. 94 : 189-206. 

1953- Ripersia wiinni n. sp. (Pseudococcidae). Ent. Ber., Amst. 14 : 235-239. 

1958. Phenacoccus sphagni (Green) in Holland. Ibid. 18 : 20-26. 

Risso, A. 1813. M6moire. Sur I'Histoire Naturelle des oranges, bigaradiers, limettiers, 

c^dratiers, limoniers ou citroniers, cultiv6s dans le department des Alpes maritimes. Ann. 

Mus. Hist. not. Paris, 30 : 416-418. 
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SuLg, K. 1912. Heliococcus bohemicus n. gn., n. sp. Acta. Soc. ent. Bohem. (Csl.) 9 : 39-48. 
1945. Zevni morphologie, metamorfosa a beh zivota cervce Nipaecoccits n. gn. nipae 

Maskell. Acta Soc. Sci. nat. Morav. 17 : 1-48. 
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three new genera and two new species. Bull. Univ. Osaka Pref. B, 7 : 1-8. 
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1-87. 
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natural habits and corresponding organization of different families. Synopsis of the British 

Insects. 2 : 158 pp. 
Williams, D. J. i960. A new species of Dysmicoccus Ferris (Pseudococcidae : Homoptera) 

on banana. Bull. ent. Res. 51 : 239-241. 



THE BRITISH PSEUDOCOCCID AE 



77 



Williams, D. J. 1961. Notes on the genus Heterococcus Ferris (Coccoidea, Homoptera) with a 
description of a new species injurious to guineacorn [Sorghum vulgare) in Nigeria. Ibid. 51 : 
671-675. 

ZahradnIk, J. 1959. Klic Zvireny C.S.R. Cervci-Coccinea, Praha, 3 : 527-552. 

Zimmerman, E. C. 1948. Insects of Hawaii, Honolulu, 5 : 132-436. 



INDEX 



aceris Phenacoccus, 34 
aceris Pseudococcus, 34 
adonidum Coccus, 40 
iidonidum Pseudococcus, 40 
alazon Dysmicoccus, 39 
albidus Pararhizoecus, 41 
albidus Rhizoecus, 41 
ANTONINA, 7 

crawi, 8 

socialis, 8 
ATROCOCCUS, 8 

cracens, 9 

luffi, IT 

paludinus, 1 1 
BALANOCOCCUS, 13 

boratynskii, 15 

scirpi, 17 
balteatus PeUococcus, 32 
balteatus Phenacoccus, 32 
boratynskii Balanococcus, 15 
brevipes Dysmicoccus, 4 
cactearum Spilococcus, 52 
cacticans Rhizoecus, 43 
cacticans Ripersiella, 43 
californicus Pseudococcus, 72 
californicus Trionymus, 72 
calluneti Parapedronia, 56 
calluneti Pseudococcus, 56 
calluneti Spinococcus, 56 
CHNAUROCOCCUS, 17 

subterraneus, 17 
CHORIZOCOCCUS, 19 

lounsbur>'i, ig 

peregrinus, 21 
citri Dactylopius, 40 
citri Dorthesia, 39 
citri Planococcus, 39 
citri Pseudococcus, 39 
Coccidella dianthi, 43 
elongata, 45 

mesembryanthemi, 45 
Coccus adonidum, 40 

halophilus, 3, 47 
comstocki Pseudococcus, 39 
cracens Atrococcus, 9 
crawi Antonina, 8 
cuspidatae Dysmicoccus, 23 



cuspidatae Pseudococcus, 23 
dactylis Pseudococcus, 59 
dactylis Trionymus, 59 
Dactylopius citri, 40 

hibernicus, 63, 69 

longispinus, 40 

luffi, II 

nipae, 31 

pulverarius, 63 

radicum, 47, 69 

theobromae, 4 

walkeri, 21 
decoratus Rhizoecus, 47 
dianthi Coccidella, 43 
dianthi Morrisonella, 43 
dianthi Rhizoecus, 43 
diminutus Pseudococcus, Oi 
diminutus Trionymus, 61 
donisthorpei Ripersia, 69 
Dorthesia citri, 39 
DYSMICOCCUS, 21 

alazon, 39 

brevipes, 4 

cuspidatae, 23 

piricola, 23 

walkeri, 21 

wistariae, 23 
elongata Coccidella, 45 
elongata Morrisonella, 45 
clongatus Rhizoecus, 45 
elongatus Rhizoecus, 43 
EURIPERSIA, 24 

europaea, 4, 24 

tomlinii, 26 
europaea Euripersia, 4, 24 
europaea Ripersia, 17, 24 
exul Ripersia, 26 
falcifer Rhizoecus, 47 
Ferrisiana virgata, 4 
filicicola Ripersia, 54 
filicicola Spilococcus, 54 
formicarii Ripersia, 17 
fragilis Pseudococcus, 40 
gahani Pseudococcus, 40 
halophila Ripersia, 41, 47 
halophilus Coccus, 3, 47 
halophilus Rhizoecus, 47, 49 



78 

HELIOCOCCUS, 28 

minutus, 29 
HETEROCOCCUS, 29 

nudus, 31 

pulverarius, 31 
hibernicus Dactylopius, 63, 69 
hibernicus Pseudococcus, 63 
hibernicus Trionymus, 63 
interruptus Phenacoccus, 35 
latipes Pseudococcus, 40 
longispinus Dactylopius, 40 
longispinus Pseudococcus, 40 
lounsburyi Chorizococcus, 19 
lounsburyi Pseudococcus, 19, 21 
lounsburyi Trionymus, 19, 21 
luffi Atrococcus, 1 1 
lujffi Dactylopius, 1 1 
luffi Pseudococcus, 1 1 
malacearum Pseudococcus, 40 
mamillariae Pseudococcus, 52 
maritimus Pseudococcus, 40 
mesembryanthemi Coccidella, 45 
mesembryanthemi Morrisonella, 45 
mesembryanthemi Rhizoecus, 45 
mesnili Ripersia, 26 
minutus Heliococcus, 29 
minutus Phenacoccus, 29 
Morrisonella dianthi, 43 
dongata, 45 

mesembryanthemi, 45 
newsteadi Pseudococcus, 61 
newsteadi Trionymus, 61 
m^oe Dactylopius, 31 
nipae Nipaecoccus, 31 
nipae Pseudococcus, 31 
NIPAECOCCUS, 31 

nipae, 31 
nudus Heterococcus, 31 
nudus Phenacoccus, 31 
orestes Trionymus, 63 
paludinus Atrococcus, 1 1 
paludinus Pseudococcus, 9, 11 
Parapedronia calluneti, 56 
Pararhizoecus albidus, 41 
PELIOCOCCUS, 31 

balteatus, 32 
peniura Saccharicoccus, 50 
peregrinus Chorizococcus, 21 
peregrinus Pseudococcus, 19 
peregrinus Trionymus, 19 
perrisii Trionymus, 63 
perrisii Westwoodia, 63 
phalaridis Pseudococcus, 67 
phalaridis Trionymus, 67 



D. J. WILLIAMS 



PHENACOCCUS. 32 

aceris, 34 

balteatus, 32 

interruptus, 35 

minutus, 29 

nudus, 31 

sphagni, 37 
piricola Dysmicoccus, 23 
piricola Pseudococcus, 23 
PLANOCOCCUS, 39 

citri, 39 
PSEUDOCOCCUS, 39 

aceris, 34 

adonidum, 40 

calif ornicus, 72 

calluneti, 56 

ct/n, 39 

comstocki, 39 

cuspidatae, 23 

dactylis, 59 

diminutus, 61 

fragilis, 40 

gahani, 40 

hibernicus, 63 

latipes, 40 

longispinus, 40 

lounsburyi, 19, 21 

/mJ^, II 

malacearum, 40 

mamillariae, 52 

maritimus, 40 

newsteadi, 61 

nipae, 31 

paludinus, 9, 11 

peregrinus, 19 

phalaridis, 67 

piricola, 23 

pulverarius, 63 

quercus, 34 

socius, 34 

sphagni, 37 

tomlini, 72 

ulicis, 34 

w/wi, 34 

walkeri, 21, 23 

wistariae, 23 
pulveraria Ripersia, 31 
pulverarius Dactylopius, 63 
pulverarius Heterococcus, 31 
pulverarius Pseudococcus, 63 
pulverarius Trionymus, 63, 69 
quercus Pseudococcus, 34 
radicum Dactylopius, 47, 69 
radicum Trionymus, 69 



THE BRITISH PSEU DOCOCCID AE 



79 



RHIZOECUS. 41 

albidus, 41 

cacticans, 43 

decor atus, 47 

dianthi, 43 

elongatus, 45 

elongatus, 43 

falcifer, 47 

halophilus, 47, 49 

mesembryanthemi, 45 

terrestris, 47 
Ripersia donisthorpei, 69 

europaea, 17, 24 

&xul, 26 

filicicola, 54 

formicarii, 17 

hcUophila, 41, 47 

mesnili, 26 

pulveraria, 31 

scirpi, 17 

subterranea, 17 

terrestris, 47 

tonUinii, ij, 24, 26 

«;Mnm, 24 
Ripersiella cacticans, 43 
SACCHARICOCCUS. 50 

penium, 50 
scirpi Balanococcus, 17 
scirpi Ripersia, 1 7 
socialis Antonina, 8 
socius Pseudococcus, 34 
sphagni Phenacoccus, 37 
sphagni Pseudococcus, 37 
SPILOCOCCUS, 52 
cactearum, 52 
filicicola, 54 



SPINOCOCCUS, 56 

calluneti, 56 
subterranea Ripersia, 17 
subterraneus Chnaurococcus, 17 
terrestris Rhizoecus, 47 
terrestris Ripersia, 47 
theobromae Dactylopius, 4 
thulensis Trionymus, 69 
tomlini Pseudococcus, 72 
tomlini Trionymus, 72 
tomlinii Euripersia, 26 
tomlinii Ripersia, 17, 24, 26 
TRIONYMUS, 58 

californicus, 72 

dactylis, 59 

diminutus, 61 

hibernicus, 63 

lounsburyi, 19, 21 

newsteadi, 61 

Orestes, 63 

peregrinus, 19 

perrisii, 63 

phalaridis, 67 

pulverarius, 63, 69 

radicum, 69 

thulensis, 69 

tomlini, 72 
li/ici^ Pseudococcus, 34 
/(/mi Pseudococcus, 34 
virgata Ferrisiana, 4 
walkeri Dactylopius, 21 
walkeri Dysmicoccus, 21 
walkeri Pseudococcus, 21, 23 
PT^s^if ooiia perrisii, 63 
wistariae Dysmicoccus, 23 
wistariae Pseudococcus, 23 
wiinni Ripersia, 24 




PRINTED IN GREAT BRITAIN BY 
ADLARD AND SON, LIMITED 
BARTHOLOMEW PRESS, DORKING 



NEW AFRICAN CADDIS-FLIES 
(Order TRICHOPTERA) 




D. E. KIMMINS 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY VoL 12 No. 2 

LONDON: 1962 



NEW AFRICAN CADDIS-FLIES 
(Order TRICHOPTERA) 



BY 

D. E. KIMMINS 



British Museum (Natural History 






Pp. 81-121 ; 107 Text-figures 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 12 No. 2 

LONDON : 1962 



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. 

This paper is Vol. 12, No. 2 of the Entomological 
series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



Trustees of the British Museum, 1962 



PRINTED BY ORDER OF THE TRUSTEES OF 
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Issued 17 April, 1962 Price Fifteen Shillings 



NEW AFRICAN CADDIS-FLIES 
(Order TRICHOPTERA) 

By D. E. KIMMINS 

SYNOPSIS 

This pap>er deals mainly with species of the subfamilies Oestropsinae and Leptocerinae and 
includes a lectotype designation and redescription of the male genitalia of Phanostoma senegalense 
Brauer and a redescription and figures of the female holotype of Setodellina albopunctata Lestage. 
The synonymy of the genus Phanostoma Brauer with the genus Amphipsyche McLachlan is 
confirmed and four new species of the latter genus are described. The genus Chloropsyche 
McLachlan is placed as a synonym of A ethaloptera Brauer. A revised key to the African genera 
of the Leptocerinae is given and the genus Setodellina is synonymized with Oecetis. 

This paper is based upon material sent for identification by Dr. Marjorie Scott 
(South Africa), Dr. Philip S. Corbet (Uganda) and on British Museum (Nat. Hist.) 
accessions. The author wishes to express his thanks to Drs. Scott and Corbet for 
their generosity in presenting to the British Museum (Nat. Hist.) the types of the 
new species based upon their material, in addition to other duplicate material. 
Paratypes of South African material (where available) have been returned to Dr. 
Marjorie Scott, and will in due course be deposited in the South African Museum. 

The South African material was collected by the following workers : Messrs. 
John Agnew, Brian Allanson, Mark Chutter, Dr. A. D. Harrison and Mr. W. D. Olliff, 
in addition to Dr. Scott herself, but the individual collectors' names are not given on 
the locality labels. 

For the loan of the type series of Phanostoma senegalense Brauer the author is 
indebted to Dr. M. Beier, Naturhistorisches Museum, Vienna, and for the loan of the 
type of Setodellina albopunctata Lestage to Dr. P. Basilewsky, Musee Royale de 
I'Afrique centrale, Tervuren, Belgium. 

Family PSYCHOMYIIDAE 

Paduniella ankya Mosely 

Two males, one from Volksrust and the other from the Natal National Park, are 
referred to this species. The genitalia do not entirely agree with the original descrip- 
tion and figures, but I do not consider the differences to be specific. The shape of the 
clasper of the Natal specimen is intermediate between that of the type specimen and 
that of an example from the Semliki Forest. The locality of the type (Namwamba 
Valley) and the Semliki Forest are both in the Ruwenzori region. 

ENTOM. 12, 2. 6 



84 D. E. KIMMINS 

Family HYDROPSYCHIDAE 

Subfamily OESTROPSINAE 

THE GENERA AMPHIPSYCHE McLACHLAN AND 

PHANOSTOMA BRAUER 

These two genera were originally separated on the difference in the tibial spur 
formula {Amphipsyche 1.4.4, Phanostoma 2.4.2) and on the presence of a weak 
" false discoidal cell " in Phanostoma. As additional species of Amphipsyche were 





Figs. 1-3. Paduniella ankya Mosely, ^ clasper from left, i, Ruwenzori, Semliki Forest 
2, Ruwenzori, Namwamba Valley (Type) ; 3, Natal National Park. 



discovered in Asia, it became evident that the tibial spur formula was not a constant 
feature of the genus, and Martynov (1935 : 201) proposed that the genus Phanostoma 
should be considered a synonym of Amphipsyche. This was not generally accepted 
and Ulmer (1951 : 197) maintained Phanostoma as a distinct genus on the grounds 
of the false discoidal cell in the fore wing and the tibial spur formula of 2.4.2. I 
have been able to study the type-series of Phanostoma senegalense Brauer and find 
that Brauer was incorrect in stating that there were two very small tibial spurs on 
the anterior tibia. Microscopic examination reveals neither spurs nor spur-bases on 
the inner apical angle of the fore tibia. There is however a variable number of micro- 
scopic spines on the outer apical angle, which Brauer may have taken for apical 
spurs. The spur formula of Phanostoma senegalense Brauer is therefore 0.4.2, 



NEW AFRICAN CADDIS-FLIES 85 

which falls within the range admitted by Ulmer for Amphipsyche. It is at times 
difficult to decide whether a false discoidal cell is present and both genera have the 
intermediate tibiae and tarsi of the male moderately flattened. 

I therefore adopt Martynov's proposal and to combine the genera Amphipsyche 
and Phanostoma. The synonymy is as follows : 

AMPHIPSYCHE McLachlan, 1872 

McLachlan, 1872 : 68, pi. 2, figs. 7-7^. 

Type-species (monotypic) Amphipsyche proluta Mch., 1872. 
Phanostoma Brauer, 1875 : 69, pi. 4, figs. 5-56. 

Type-species (monotypic) Phanostoma senegalense Brauer, 1875. 
? Amphipsychella Martynov, 1935 : 201. 

Type-species (monotypic) Amphipsychella extrema Martynov , 1935. 

Spur formula (^, 9. I-4-4, I-4-3. I-4-2, 0.4.3, 0.4.2. Mid-tibia and first three 
basal tarsal segments moderately flattened in (^, strongly so in $. Maxillary palpi 
weak, rather short. Aedeagus with or without parameres, [A. proluta and A. sene- 
galensis are certainly without spine-like parameres, nor does Martynov's figure of 
A. distincta give any indication of them. The four new African species have such 
parameres, as do /I. indica Mart, and A. bengalensis Mart.) 

Key to Akkican Spkcies of Amphipsyche (Males) 

1 Stem of R2+3 in fore wing closely approximated to R^ ; aedeagus without spiniform 

parameres ......... senegalensis (Brauer) 

- Stem of i?2+3 in fore wing widely separated from 7^4 ; aedeagus with two spiniform 

parameres ............. 2 

2 Lobes of tenth segment, in dorsal view, arising from broad, shouldered bases . . 3 

- Lobes of tenth segment, in dorsal view, not arising from broad bases ... 4 

3 Lobes of tenth segment parallel in dorsal view, deep in side view . . corbeti sp. n. 

- Lobes of tenth segment divergent in dorsal view, narrow in side view . berneri sp. n. 

4 Apex of aedeagus, in side view, strongly clavate, parameres angled abruptly upwards. 

Lobes of tenth segment deep in side view ..... ulmeri sp. n. 

- Apex of aedeagus, in side view, produced but not strongly clavate, parameres gently 

curved. Lobes of tenth segment narrow in side view . . . scottae sp. n. 

Amphipsyche senegalensis (Brauer) 

(Text-figs. 4-13) 

Phanostoma senegalense Brauer, 1875 : 71, pi. 4, figs. 5-5C (Senegal) ; Ulmer, 1907 : 26, fig. 12 
(Congo) ; id., igoja : 32-33, fig. 24 ; id., 1907b : 159, pi. 21, fig. 198 ; Mosely, 1932 : 5, pi. i, 
fig. 6 (Mozambique) ; Barnard, 1934 • 3^9 • 

Brauer's description is based upon males only and the account of the general 
appearance is good. His figure of the wing venation is also reasonably good, but he 
shows the veins i?2+3 ^.nd R^ in the fore wing as fused near their bases, whereas 
(as Ulmer has pointed out) they are separate but close together. Cell jRa+s ^^ hind wing 
may be either just sessile or with a short footstalk. His description and figures of 
the male genitalia, being made from the dried material, are rather inadequate by 
modern standards and I am therefore giving a new description and figures. 

ENTOM. 12, 2. 6§ 



86 



d. e. kimmins 
Designation of Lectotypi': 



No lectotype has been selected from Brauer's type series, so I take this opportunity 
of designating the example with the following labels : " Steindachner, 1869, Senegal "; 
" senegalense det. Brauer "; " senegalense det. Ulmer "; " Phanostoma senegalense 
Brauer, <^, Lectotype, D. E. Kimmins det. i960 " (Vienna Museum). The lectotype 
has one pair of wings mounted between sheets of celluloid, the abdomen and one fore 
leg cleared in KOH solution and preserved in a small vial of glycerine, both prepara- 




FiGS. 4-9. Amphipsyche senegalensis {'Bxa.VLex), (S^^ctoty^Q. 4, wings ; 5, fore tibia, with 
apex more enlarged ; 6, genitalia, lateral ; 7, aedeagus, lateral ; 8, genitalia, dorsal ; 
9, ventral and from behind. 



NEW AFRICAN CADDIS-FLIES 



87 



tions being attached to the staging pin. There are also four other males from the 

type-series. 

cJ GENITALIA of lectotypc. Ninth segment with the dorsal apical margin only slightly produced 
at its centre. Side-pieces large and rounded. Tenth segment deeply excised at its centre to form 




Figs. 10-13. Amphipsyche senegalensis (BrsiueT), ^ {Luailaba. River). 10, wings ; 11, genitalia, 
lateral ; 12, ventral ; 13, bursa copulatrix, ventral, more enlarged. 



a pair of narrow ovate lobes, set well below the level of the ninth tergite and linked to the latter 
by membrane. The upper surface of the lobes bears a number of small setae. Lower margins of 
the tenth segment partly enfold the stem of the aedeagus. The latter organ has a wide base, 
constricting to a slender stem and then dilating to form a clavate apex. The apical surface of 
aedeagus bears two small triangular teeth or lobes (shown in Brauer's fig. 5^). There are no 



88 



D. E. KIMMINS 



spiniforni parameres. Claspers long, slender and sinuous, the articulation between the basal 
and terminal segments somewhat indefinite. 

$ GENITALIA (Congo, Lualaba River). Pleurosternuni of eighth segment large, plate-hke, 
its convex lower margin almost meeting that of the opposite side ventrally. Posterior margin 
slightly excised. Eighth sternitc reduced to a narrow, lightly sclerotized tongue, not extending 




Figs. 14-18. Amphipsychc ulmeri sp. n. J. 14, wing.s ; 15, genitalia, lateral; 
dorsal ; 17, apex of aedeagus, dorsal ; 18, genitalia, ventral and behind. 



16, 



as far as apices of the pleurosterna. Ninth tergite forming a hood, clasper groove obscure. 
Ninth sternite narrow, apex slightly excised. Tenth segment with the usual three pairs of 
processes. 

Length of fore wing, cJ. 13-15 mm.; ?, 9-10 mm. 

I have seen no females from the type-locahty, Senegal. I have therefore en- 
deavoured to fix the female of senegalensis by examination of females taken at the 
same time and place as males of senegalensh. Examination of females fulfilling these 
conditions from the Congo (Lualaba R.) and Lake Nyasa suggests that the female 



NEW AFRICAN CADDIS-FLIES 89 

venation differs appreciably from that of the male. In the fore wing, veins /?2+3 
and R^ are not closely approximated and Rs is more or less strongly sinuous, much more 
so than in the male. Examples from Lake Victoria also show this sinuosity of Rs 
in the fore wing, and in this locahty the males would appear to be extremely rare, 
since I have seen only one example, although Dr. Corbet and Dr. Hickin have sent 
me several hundred females. The fact that females with a sinuous Rs in the fore wing 
appear to be associated with male senegalensis suggests that Phanostoma curvinerve 
Navas, based upon females from Egypt, may be a synonym of senegalensis. This 
sinuosity of Rs in the fore wing does not appear to be a constant generic character 
in Amphipsyche since, in the females of A. scottae sp. n. from S. Africa, Rs is almost 
straight. 

Distribution. I have seen males from Senegal, Congo, S. Rhodesia, Nyasa- 
LAXD and Uganda. 

Amphipsyche ultneri sp. n. 

(Text-figs. 14-18) 

Phanostoma senegalense Brauer ; Ulmer, 1923 : 19 {partim, Sennar) ; id., 1924 : 2 (Sennar). 

Egyptian Sudan : Sennar, 18-27. ii.1914, ^bner, 7 (^, two labelled " Phanostoma 
senegalense Brau." in Ulmer 's writing. 

^ HOLOTYPE (pinned). General colour pale tawny, antenna finely annulated with reddish. 
Thorax clothed with short, pale pubescence. Fore wing very pale tawny, with slightly darker 
clouding over the anastomosis. Rs almost straight, i?2+3 separated from i?4 at their bases M' 
about the distance between R^ and R^. In the hind wing, cell R2+3 is sessile. 

cJ GENITALIA foUowing the general pattern of the genus. Aedeagus with two curved spine-like 
parameres, angled abruptly upwards. Stem of aedeagus less slender than in senegalensis, apex 
clavate, its lower margin projecting in two rounded, triangular lobes. 

Length of fore wing, 13-14 mm. 

(^ HOLOTYPE (pinned, with wings between celluloid and abdomen in a small vial 
of glycerine) and paratypes in the Naturhistorisches Museum, Vienna. In the course 
of correspondence with Dr. Ulmer in 1956 concerning Phanostoma, he informed me 
that he had in his collection two females from Sennar and that they had Rs in the 
fore wing curved but less so than in Egyptian examples which he had identified as 
P. curvinerve Navas. This fact may be additional evidence in favour of A. ulmeri 
being distinct from A . curvinervis. 

Amphipsyche corbeti sp. n. 

(Text-figs. 19-23) 

Uganda : Northern Province, Victoria Nile, Karuma Falls, P. S. Corbet, numerous 
males. 

cJ HOLOTVP1-: (in alcohol). General colour very pale tawny. Antenna finely annulated witli 
fuscous. Anterior femur with a longitudinal fuscous streak on ventral surface. Mesonotum 
rather darker tawny. Abdominal segments whitish, faintly marked with very pale purplish 
transverse bands on the tergites and with a similarly coloured median ventral streak. In fore 
wing, /?2+3 widely separated from R^ basally. 



go 



D. E. KIMMINS 



^ GENITALIA. Ninth segment with its dorsal apical margin only slightly produced. Side- 
pieces rounded. Lobes of the tenth segment broad basally, the apical portion about half as 
wide as the basal, which projects laterally as a shoulder. In dorsal view, the lobes are parallel, 
with rounded apices. Aedeagus with a stout stem, in side view somewhat constricted about 
mid- way, lower apical margin produced in a pair of lobes. In ventral view, the inner margins 




Figs. 19-23. Amphipsyche corheti sp. n. ^. 19, wings ; 20, genitalia, lateral ; 21, 
dorsal ; 22, apex of aedeagus, dorsal ; 23, genitalia, ventral and behind. 



of the lobes are parallel, their basal angles hooked. Above these lobes are a pair of stout, spini- 
form parameres, curving upwards in side view. Claspers slender, indistinctly two-segmented. 

$ unknown. 

Length of fore wing, 11 mm. 

^ HOLOTYPE (now mounted as microscope preparations), paratypes in 2% formal- 
dehyde solution, in British Museum (Nat. Hist.). This species differs from A. ulmeri 



NEW AFRICAN CADDIS-FLIES 



91 



in the less constricted stem of the aedeagus, whose apex is more produced, and in 
the stouter parameres. The tenth segment is wider at the base in dorsal view, with 
definite shoulders. 

Amphipsyche berneri sp. n. 

(Text-figs. 24-28) 
Phanostoma senegalense Brauer : Kimmins, 1957 • ^3 (Gold Coaist examples). 
Gold Coast : Volta River, Senchi, i.viii.1950, Lewis Berner, 4 (^. 

(J HOLOTYPE. Specimen originally preserved in alcohol, general colour tawny yellow, antenna 
with fine fuscous annulations, abdomen with faint purplish markings. Fore wing yellowish 




Figs. 24-28. Amphipsyche berneri sp. n. ^. 24, wings ; 25, genitalia, lateral ; 26, dorsal 
27, apex of aedeagus, dorsal ; 28, genitalia, ventral and behind. 



02 D. E. KIMMINS 

brown, with darker shading behind the sinuously curved portions of Sc and Ri. Cell i?2+3 at 
base at least as wide as cell i?4. Rs straight. 

c? GENITALIA resembling A. corbeti in general pattern. Tenth segment also shouldered at its 
base, but in the type the lobes of the tenth tergite are strongly divergent in dorsal view and in 
side view appear as narrow plates. The aedeagus is similar but, in side view, there two pro- 
jections on the dorsal margin. In ventral aspect the apex is more clavate. 

Length of fore wing, 12 mm. 

cJ HOLOTYPE (mounted as microscope preparations) and i paratype in 2% formal- 
dehyde solution, in British Museum (Nat. Hist.), 2 <^ paratypes in University of 
Florida. 




Figs. 29-33. Amphipsyche scottae sp. n. J. 29, wings ; 30, palpi ; 31, genitalia, lateral 
32, dorsal, with ape.x of aedeagus ; 33, ventral and behind. 



NEW AFRICAN CADDIS-FLIES 



93 



Amphipsyche scottae sp. n. 

(Text-figs. 29-37) 

South Africa: Vaal River barrage, 20.iii.1956, 6 c^, 31.iii.1958, i ^ ; below 
Vaal barrage wall, 29 . vi . 1956, 8 ^ ; Vaal River, below barrage, 23 . viii . 1957, i $ 




Figs. 34-37. Amphipsyche scottae sp. n. ?. 34, wings ; 35, genitalia, lateral ; 36, ventral 

37, bursa copulatrix, ventral. 



and pelt ; Waterval River, at Joburg-Durban Road, 16 . x . 1958, 2 c^ ; Wilge River 
at Warden-Reitz Road, 8.xii.i958, 8 ^ ; Wilge River at Frankfort, 8.xii.i958, 
3 (^ ; Waterval River, on National Road between Standerton and Greylingstad, 
12.1.1959, 3 ^ ; Wilge River, 5 mUes below Harrismith, 10. ii. 1959, 3 J; Wilge 
River, between Warden and Reitz, 9 . ii . 1959, 9 3- All specimens in alcohol. 



ENTOM. 12, 2. 



6§§ 



94 D. E. KIMMINS 

^ HOLOTYPE. As the specimen was in alcohol, one can only say of its general appearance that 
it was tawny yellow, antenna finely annulated with reddish at the joints. Abdomen lightly 
tinged with reddish purple. Fore wing faintly clouded with brownish. Rs almost straight, 
R2+3 not closely adjacent to Rt, the space between them not much less than that between R^ 
and Ri. In hind wing, cell i?2+3 generally broadly sessile. 

cJ GENITALIA following the general pattern of A. corbeti and A. berneri. Tenth segment not 
broadly shouldered at base, the lobes with their inner margins concave in dorsal view and in side 
view thin as in berneri. The aedeagus with a pair of spiniform parameres, projecting from beneath 
a pair of rounded dorsal lobes. Apical lobe of aedeagus truncate, not divided, its ventral surface 
bearing a pair of small triangular plates, facing each other. 

? ALLOTYPE. This specimen was bred and is rather teneral. Colours much as in male. In the 
fore wing, Rs runs almost straight to the anastomosis, not sinuous as in senegalensis . Apical 
cellules narrower than in senegalensis. 

? GENITALIA. Pleurostcmum of eighth segment large, plate-like, its curved lower margins 
almost meeting over the eighth sternite. The latter is lightly sclerotized and does not extend 
beyond the pleurosternum. Ninth tergite forming a hood, clasper groove shallow. Ninth sternite 
in the shape of a transverse plate, whose apical margin is shallowly excised at the centre. Tenth 
segment with the usual three pairs of processes. 

Length of fore wing, (^,18 mm. ; ?, 14 mm. 

S HOLOTYPE (Wilge R., 5 miles below Harrismith, 10. ii. 1959), 9 allotype (Vaal 
River, below barrage, 23.viii.1957), both mounted as microscope preparations 
in British Museum (Nat. Hist.) ; paratypes in South African Museum and British 
Museum (Nat. Hist.). This species is related to A. corbeti and A. berneri but may be 
separated by the form of the tenth tergite and of the aedeagus. This species is 
named after Dr. Marjorie Scott, of the University of Cape Town. 

THE GENERA AETHALOPTERA AND CHLOROPSYCHE 

(Text-figs. 38-47) 

The genus Aethaloptera was described by Brauer (1875 : 71), with yl. dispar Brauer 
(W. Africa) as the only included species. McLachlan (1880, Suppl. 2 : Ixix-lxx) 
described the genus Chloropsyche, with type species evanescens McL. (Amurland), 
and among the characters given to separate it from Aethaloptera Brauer are the spur 
formula 0.2.2 [Aethaloptera 0.3.2), and the male intermediate tibiae slender, with 
the tarsi slightly flattened, fringed but not dilated. McLachlan's figure of the male 
wings shows that in the hind wing, fork R^ is sessile as in Aethaloptera. Ulmer 
(1906 : 62) described Aethaloptera maxima from S. Africa, which had a spur formula 
of o. 2 . 2, and fork R^ in the hind wing stalked. This species was transferred by Ulmer, 
(1907^ : 17) to Chloropsyche McLachlan, with which it agreed in spur formula but 
not in the shape of fork R^ in the hind wing. As a result, Chloropsyche then differed 
from Aethaloptera only in possessing two instead of three spurs on the intermediate 
tibiae, the latter being not or less dilated than in Aethaloptera. 

As the male genitaha of Chloropsyche maxima appeared to be almost identical with 
those of A . dispar, it was decided to investigate more thoroughly the type-specimen 
of Chloropsyche evanescens McL. It was at once obvious that McLachlan's statement 
(1880 : Ixix) " All the legs are slender in the male " was not strictly accurate. The 
intermediate tibiae and tarsi are sHghtly but definitely dilated, the latter being nearly 
twice as wide as the posterior tarsi. The intermediate tibiae also showed a slight 



NEW AFRICAN CADDIS-FLIES 



95 



swelling just beyond the base and in this area (in the pinned specimen) near the 
anterior margin there appeared to be a minute pit. Examination of the cleared 
intermediate tibiae of a second specimen under a compound microscope suggested 
that this pit was the rudimentary socket of the third or pre-apical spur. No actual 
spur was seen in the four specimens studied and it would seem that the pre-apical 




Figs. 38-42. Wings oi A eihalopter a spp. 38, A. dispar Brctuer, <S (Uganda) ; 39 A. dispar, 
? (Sierra Leone) ; 40, ^4. maxima Ulmer, ^ (S. Africa) ; 41, A. maxima, ? (SW. Africa) ; 
42, A. evanescens McLachlan, <? (Amur). 



spur of the intermediate tibia is either very easily detached or obsolete in evanescens. 
Specimens of C. maxima have the intermediate tibiae and tarsi about as much dilated 
as in evanescens, but microscopic examination reveals neither a swelling just beyond 
the base nor any evidence of a spur socket. Other cases are known in which the tibial 
spur formula has proved variable and unreliable as a generic character and the genus 
Chloropsyche does not appear to me to be worthy of separation from Aethaloptera 



96 



D. 



KIMMINS 



and I place it in the synonymy of that genus. Paraethahptera Martynov (1935 : 193), 
type species P. gracilis Mart., 1935, also seems doubtfully distinct from Aethaloptera. 
The type of Paraethaloptera punctata Banks has a spur formula of 0.3.2, and should 
be transferred to Aethaloptera. 

AETHALOPTERA Brauer 

1875 : 71. Type-species, A. dispar Brauer, 1875. 
Chloropsyche McLachlan, 1880 : Ixix (syn. nov.). 

Type species, C. evanescens McL., 1880. 
? Paraethaloptera Martynov, 1935 : 193. 

Tjrpe species, P. gracilis Mart., 1935. 

Four species of this genus have been recorded from Africa, A. dispar Brauer, 
A. maxima Ulmer, A. maesi (Navas) and A. marcina (Navas), the two latter species 





Figs. 43-47. Median tibiae, etc., of Aethaloptera spp. 43, A. dispar Brauer, ^ (Gold 
Coast) ; 44, A. evanescens McL., c? (Amur) ; 45, A. maxima Ulmer, ^ (Port. E. Africa) ; 
46, A. dispar, ?, (Sierra Leone) ; 47, A. maxima, ?, (SW. Africa). 



being considered as synonyms of A. dispar. Whilst A. dispar and A. maxima can 
generally be separated on the spur formula of the intermediate tibiae and on the 
stalked or sessile condition of fork R^^ in the hind wing, both Ulmer and Barnard 
have commented on the fact that the male genitalia are apparently identical. I have 
examined numerous specimens and a number of cleared preparations of both species 
and find that the genitalia, though similar, are not uniform in details, but the varia- 
tions do not appear to be correlated with any particular spur formula or arrange- 
ment of wing venation. The females similarly do not show any constant differences 



NEW AFRICAN CADDIS-FLIES 



97 



in genitalia which might be considered as specific. The venation of the hind wing in 
the females also shows some variation, specimens with three tibial spurs sometimes 
having a short footstalk to fork R^ instead of having it sessile. Such variation is not 




Figs. 48-55. A ethaloptera dispar BrSLuer, ,^ genitalia,. 48, lateral ; 49, dorsal ; 50, apex of 
aedeagus, dorsal, more enlarged; 51, claspers and aedeagus, ventral; 52, aedeagus, 
lateral (Gold Coast) ; 53, the same (Sierra Leone) ; 54, the same (Nyasaland) ; 55, 
ninth segment, dorsal (Nyasaland). 



surprising when one considers that the stalked state of fork R^ in this genus is a 
secondary condition. R^ and R^ separate at the anastomosis, rapidly running together 
again, forming a small cell enclosing the corneous point, subsequently separating 
again to form fork R^. Fork M3 in the fore wing of the female may be either sessile 
or stalked in dispar and usually stalked in maxima. 



98 



D. E. KIMMINS 



These two species in Africa possibly represent local forms of one widespread species, 
maxima being centred in S. Africa and spreading northwards into N. Rhodesia and 
Portuguese East Africa. A. dispar has its centre in equatorial Africa, extending 
from west to east and northwards into the Sudan. In Nyasaland one gets a form which 
is intermediate, the venation and spur formula being that of dispar and the male 
genitalia tending towards that of maxima. Ulmer's record of dispar from Livingstone, 
N. Rhodesia suggests that the range of both species overlaps in Northern Rhodesia. 
For the present, however, it is proposed to treat the two as closely allied species. 




Figs. 56-59. A ethaloptera maxima Vlmer, (^ genitalia. 56, lateral ; 57, aedeagus, lateral 
58, genitalia, dorsal ; 59, claspers and aedeagus, ventral. 



Aethaloptera dispar Brauer 
(Text-figs. 38, 39, 43, 46, 48-55, 60-61) 



1875 : 72, pi. 4. figs. 4-4C. 
Prim.erenca maesi Navds, 1914 : 182. 
Primerenca marcina Navas, 1916 : 26, 



NEW AFRICAN CADDIS-FLIES 



99 



From records and specimens the distribution appears to be Senegal, Sierra 
Leone, Gold Coast, S. Nigeria, Belgian Congo, S. Sudan, Uganda, Tanganyika, 
Northern Rhodesia (Livingstone) and Nyasaland. 




Figs. 60-63. Female genitalia of, 60-61, A . dispar Brauer (Uganda), and 62-63, ^ ■ maxima 
Ulmer (S. Africa). 60, lateral ; 61, ventral ; 62, lateral ; 63, ventral. 



Figures are given of the ^ genitalia of an example from Gold Coast, and comparative 
figures of the aedeagus in side view of specimens from Sierra Leone and Nyasaland. 
It will be seen that the shape of the apex is by no means uniform and that there is 
variation in the contour of the lower margin of the stem. The lower surface towards 



loo D. E. KIMMINS 

the apex is often more keeled than in maxima. The apical margin of the ninth segment 
varies in the amount of its production but is generally without a median dorsal 
furrow. The degree to which the lobes of the tenth segment diverge is not constant. 

In the female, the produced setose lobes on the apical margin of the eighth sternite 
are variable in shape. The apex of the ninth sternite is only very sparsely setose. 
Bursa copulatrix rather complex in structure but possibly variable. 

Length of fore wing, (J, 10-16 mm.; $, 7-10 mm. 

The female specimens recorded doubtfully by Ulmer (1912 : 95) as Aethaloptera 
sexpunctaia Kolenati from Kamerun may possibly belong here. Since he recorded 
them as Aethaloptera, one presumes that they had the typical spur formula 0.3.2. 
On the other hand, fork R^ in the hind wing is stalked, which suggests that they 
might be A. maxima, which would be a rather unusual distribution for that species 
as we know it at present. 

Aethaloptera maxima Ulmer 

(Text-figs. 40, 41, 45, 47, 56-59, 62, 63) 

1906 : 62, fig. 66. 

Chloropsyche maxima (Ulmer), 1907 : 17, fig. 2. 

Records and specimens give the distribution as follows : S. Africa, SW. Africa, 
Portuguese E. Africa, NE. Rhodesia. 

In the male genitalia, the apical margin of the ninth segment is generally more 
produced and there is often a longitudinal dorsal furrow. The apex of the aedeagus 
is rather more rounded in side view than in dispar. In the female, the genitalia are 
much as in dispar but the apex of the ninth sternite is densely setose. 

Length of fore wing, (^, 14-18 mm.; $ 9-13 mm. 

Family HYDROPTILIDAE 

Oxyethira velocipes (Barnard) 

(Text-figs. 64-67) 

, After examination of a paratype male and other material of this species, I agree 
with the suggestions of Drs. Ross and Ulmer that Argyrobothrus is not generically 
distinct from Oxyethira. It has the same spur formula and the arrangement of the 
warts on the mesoscutellum and the shape of the metascutellum agree. There is a 
slender spine at the apex of the seventh sternite. The male genitalia of this species 
are not easy to interpret and are rather more complex than Barnard's figures suggest 
(1934 : 392). I am therefore refiguring them on a larger scale from a paratype and 
giving a new description of them. 

(J GENITALIA. Eighth Segment synscleritous, almost completely encasing the ninth segment, 
from the side obliquely truncate, and from beneath with its apex excavate. Ninth segment 
about as long ventrally as eighth, but dorsally narrowed to a transverse band. Apex of ninth 
segment also excavate ventrally. Aedeagus long, slender, with a spiral spine somewhat beyond 
mid-way, and another curved spine shortly before the apex. Claspers reduced to two short, 
wide lobes, projecting a little beyond the margin of the ninth segment in the excavation. Basally 
they are linked to a structure, possibly the basal plate, which takes the form of a quadrate plate 



NEW AFRICAN CADDIS-FLIES lOi 

beneath the aedeagus. Its upper margins extend dorsally at the base and completely encircle 
the aedeagus, this upper part being joined by membrane to the ninth segment. On each side 
of the basal plate is a digitate, transparent process, terminating in a seta. Above the aedeagus 
is a membranous lobe (? tenth segment). The basal plate is the structure referred to by Barnard 
as the tenth segment. 




Figs. 64-67. Oxyethira velocipes (Barnard), genitalia. 64, c?. lateral ; 65, cJ, dorsal ; 66, 

cJ, ventral ; 67, ?, ventral. 



I02 D. E. KIMMINS 

Family LEPTOCERIDAE 

SubfamUy LEPTOCERINAE 

Whilst Kimmins' (1956) paper on the African Leptocerinae was in the press, 
Marlier (1955 : 150) added one new genus, Limnoecetis, and I therefore take this 
opportunity of revising my key to the African genera. 

1 Fork Cui„ in hind wing present ....,..., 4 

- Fork Cuia in hind wing absent .......... 2 

2 Wings reduced, very narrow, no apical forks in hind wing . LIMNOECETIS* 

- Wings not reduced, hind wing with at least fork R^ . . . . . . 3 

3 In fore wing, stem of M (usually) obsolete, thus thyridial cell is absent ; if stem of 

M is present, then stem of epicranial suture is absent . TRIAENODES 

- In fore wing, stem of M present, complete, thyridial cell present ; stem of epicranial 

suture present .......... ADICELLA 



5 
7 
6 

LEPTOCERINA ? 

ATHRIPSODES ? 



Fore wing with forks R2, M^ and Cui^ present 

Fore wing with forks i?2 and Cuia only 

In fore wing, cell R5 with a footstalk 

In fore wing, cell R^ sessile 

In hind wing, Rs is more or less parallel to i?i 

In hind wing, Rs diverges from Ri, R2+3 arching towards i?i 

PSEUDOLEPTOCERUS ? 
Rs in hind wing completely or partially obsolete ...... 8 

Rs in hind wing not obsolete .......... 12 

In hind wing, M also obsolete at base LEPTOCELLA 

In hind wing, M not obsolete at base ........ g 

Spurs 1.2.2, that of anterior tibia small ; fork R^ in hind wing present or absent 

SET ODES {partim) 
Spurs 0.2. 2; fork i? 2 in hind wing present ....... 10 

Hind wing moderately broad (about 2-|-3 times as long as broad), posterior margin 

moderately convex HEMILEPTOCERUS 

Hind wing narrow (about 4-5 times as long as broad), posterior margin only slightly 

convex ............. 11 

Basal segment of antenna (in both sexes) rather short, stout, tapering to apex and 

without a tuft of long hairs at apex in male .... SETODES (partim) 
Basal segment of antenna (in both sexes) slender, cylindrical, in male with a tuft 

of long hairs at apex TRICHO SETODES 



12 M in hind wing unforked 

— M in hind wing forked 

13 Spurs 1.2.2, fork 7? 2 in hind wing present 

— Spurs 2.2.2, fork JF?2 in hind wing absent 

14 In fore wing, M3+4 clearly arises from M 



13 

14 

PARASETODES 
PTOCHOECETIS 

15 

— In fore wing, M3+4 arises from w-cm cross-vein or from Cu^^ 

OECETIS {=SETODELLINA) 

15 In fore wing, M3+4 arises at or before anastomosis . . . . . . 16 

— In fore wing, Ms+4 arises beyond anastomosis . . . . . . . 17 

16 Spurs 0.2.2 LEPTOCERUS 

— Spurs 2.2.2 LEPTOCERINA^ 

17 In hind wing, Rs diverges from 7?2 to enclose a wide radial area 

PSEUDOLEPTOCERUS S 

— In hind wing, Rs runs more or less parallel to 7?^ . . . . . 18 

* In October, 1961, after this paper had gone to press, Marlier {{Inst. R. Sci. nat. Belg., Bull. 38 (28) : 
1-6) transferred his genus Limnoecetis to the family Polycentropodidae, Dipseudopsinac. Couplet 2 in 
the key should therefore be deleted. 



NEW AFRICAN CADDIS-FLIES 

1 8 In fore wing, venation differing in the sexes, fork Mj present in the ? 

— In fore wing, venation aUke in both sexes .... 

19 Fork i?2 present in hind wing ....... 

— Fork Ro absent in hind wing . . ... 



103 

ATHRIPSODES 

19 

. HOMILIA 

LEPTECHO 



THE GENUS TRIAENODES McLACHLAN 

The majority of the species of this genus are easily recognized by the absence of 
the stem of the media in the fore wing and this has been used as the main recognition 




Figs. 68-70. Wings of Triaenodes spp. n. 68, T. uncata, ^ ; 69, T. uncata, ? ; 70, T. moselyi, cJ. 



character for the genus. Barnard (1934 : 357) described as Adicella sicula a species 
which had venation typical of Adicella, that is, it had the media normally formed and 
complete, but he commented on the remarkable similarity of the genital structure of 
the male with that of Triaenodes serrata Ulmer. Mosely (1936 : 434) described 
Triaenodes darfurica, which has the male genital structure of the Triaenodes pattern ; 
he figures the venation but strangely enough makes no comment on the fact that 



I04 D. E. KIMMINS 

the venation is atypical, having the stem of the media complete in the fore wing. 
Since then, several other species of Triaenodes have been described in which the stem 
of the media in the fore wing shows varying degrees of completeness, the extreme 
base being most often obsolete. Thus the absence of the stem of the media in the 
fore wing can no longer be relied upon entirely as a character of the genus Triaenodes. 

On the other hand, judging by the male genital structures, there does not seem 
to be any question that Triaenodes and Adicella are generically distinct. Other possible 
characters were therefore investigated and it seems that a feature mentioned by 
Ross (1944 : 212) may be used to supplement the venational character. This is the 
presence or absence of the stem of the epicranial suture. In Triaenodes it is absent 
in nineteen out of twenty species examined, the exception being the unique type of 
Triaenodes chelifera (Mosely), in which it is present but weakly formed. This species 
is the type-species of Mosely's genus Triaenodella, which was distinguished from 
Triaenodes primarily on the presence in the male of a lobe covering a scent tuft on 
the basal segment of the antennae. Similar lobes occur in other species of Triaenodes 
which do not have the stem of the epicranial suture developed. In Adicella this stem 
is well developed in six out of eight species examined and in Erotesis (closely allied 
to Adicella) it is present in both species examined. The exceptions are A . triaenodi- 
formis Ulmer (known only from the female type) and A. pulcherrima Ulmer. In 
the first of these exceptions, it is perhaps significant that Ulmer has given it a name 
suggesting a resemblance to Triaenodes, and it may be that when the male is recog- 
nized, it will prove to have genitalia of the Triaenodes pattern. In the second case, 
I have seen only two examples and in these, the genitalia are more like those of Adicella 
than Triaenodes. 

As a result of these investigations, I have reversed my opinion (1956 : 146) when 
I transferred Triaenodes darfurica Mosely to Adicella on venational grounds, and I 
am also considering Adicella sicula Barnard to be a Triaenodes. I also propose that 
Adicella triaenodiformis Ulmer be placed provisionally in Triaenodes. 

Key to Males of African Species of Triaenodes 

(Excluding those of the Mediterranean Region). 

The following species are omitted : T. africana Ulmer, figures of male genitalia inadequate ; 
T. palpalis Banks, no description of genitalia, sex not given ; T. triaenodiformis (Ulmer), 
known from female only. 

1 Clasper with a curved spine arising from the base, in addition to a basal branch . 2 

- Clasper without a curved basal spine ........ 6 

2 Clasper slender, basal branch very small ..... elegantula Ulmer 

- Clasper stouter, basal branch conspicuous ....... 3 

3 Clasper, in side view, short, rectangular (known to me from literature only) 

hastata Ulmer 

- Clasper, in side view, not rectangular ........ 4 



Apex of clasper, in side view, obliquely truncate 
Apex, in side view, acutely produced 
Apex of clasper directed caudad in ventral view 
Apex incurved ...... 

Tenth segment with one (rarely two) arched spine, arising from a socket on dorsal 
surface ............. 



uncata sp. n. 

5 
darfurica Mosely 
wambana Mosely 



NEW AFRICAN CADDIS-FLIES 105 

- Tenth segment without such spine ......... 9 

7 Clasper, in side view, short, about as long as deep, basal branch short and broad, aj>ex 

scarcely bifid ......... falculata Kimmins 

- Clasper, in side view, at least twice as long as deep, basal branch slender with bifid 

apex .............. 8 

8 Cerci shorter than tenth segment (excluding spine). Lower fork of basal branch of 

clasper shorter than apical branch ...... sicula (Barnard) 

- Cerci longer than tenth segment. Lower fork of basal branch of clasper longer and 

narrower than apical branch ....... serrata Ulmer 

g Basal branch of clasper slender, terminating in two long, slender forks . . . 10 

- Basal branch stouter, not terminating in two long, slender forks . . . . 12 

10 Inner and outer margins of clasper, in ventral view, straight and parallel, clasper 

abruptly narrowed at apex ........ moselyi sp. n. 

— Inner margins convex or sinuous, clasper not abruptly narrowed at apex . 11 

11 Clasper, in ventral viev\% three times as long as wide, upper fork of basal branch 

clavate ........... difformis Mosely 

— Clasper, in ventral view, twice as long as wide at base, tapering to apex. Upper fork 

spine-like .......... hickini Kimmins 

12 Side lobes of tenth segment fused to form a single spine ; median lobe reduced to two 

small processes, or absent ..........13 

— Side lobes separate, except at base, spiniform ; median lobe well developed or 

reduced ............. 14 

13 Basal branch of clasper simple, clavate, with a ventral spine . . kimila Mosely 

— Basal branch of clasper bifid, both forks arched downward . . . ghana Kimmins 

14 Median lobe of tenth segment reduced to two small, transparent fingers at the bases 

of the cerci ........... legona Mosely 

— Median lobe of tenth segment produced, finger-like, longer than cerci . . . 15 

15 Clasper, from beneath, not longer than wide, basal branch strongly incurved 

polystachya Marlier 

— Clasper, from beneath, distinctly longer than wide . . . . . . 16 

16 Apical part of basal branch of clasper flattened, blade-like, lower edge serrate and 

with a long acute tooth ........ clavata (Mosely) 

— Apical part of basal branch forming a rounded lobe, with a stout, blunt projection 

beneath it ......... chelifera (Mosely) 



Triaenodes moselyi sp. n. 

(Text-figs. 70-75) 
Triaenodella haslata (Ulmer) Mosely, 1939 : 16. 

Uganda : Ruwenzori Range, xii . 1934-i . 1935, /' • W, Edwards, Namwamba Valley, 
6,500 ft., I c^, Fort Portal, 5,000 ft., 5 ?. 

<S HOLOTYPE (pinned). General colour pale tawny, with golden pubescence. Antenna finely 
annulated with fuscous, basal segment long and slender and with a plate covering the dorsal 
surface, although no pencil of hairs is visible beneath it. Palpi long, sparsely pubescent. Fore 
wing with golden pubescence and with indistinct spots of brownish pubescence in the apical 
half. Hind wing with sparse brownish pubescence. Fore wing rather narrow, discoidal cell 
long, almost twice as long as its footstalk. In hind wing, fork R^ has a short footstalk. 

cJ GENITALIA. Ninth segment long ventrally but reduced dorsally to a narrow transverse 
band. Side-pieces large and triangular, with rounded setose apices. Lateral margins at base of 
claspers extended upwards and inwards to support the base of the aedeagus. Median lobe of 
tenth segment inconspicuously trifid, the side branches very short, centre branch slender, 
slightly clavate and setose at apex. Side lobes fused together beneath the median lobe to form 



io6 



D. E. KIMMINS 



a long, blade-like, curved spine, with a short projection about mid-way on the left side. Cerci 
long and slender, of unequal length. Aedeagus arched, with a median dorsal furrow, membranous 
towards apex dorsally and with a membranous upper lobe on each side. Clasper about twice as 
long as wide, quadrate in side and ventral views, the outer apical angle, in ventral view, produced 




Figs. 71-75. Triaenodes moselyi sp. n. Genitalia. 71, c?. lateral ; 72, S, tenth segment, 
dorsal ; 73, cj, claspers and aedeagus, ventral ; 74, ?, lateral ; 75, ?, ventral. 

in a short, blunt, incurved process. Dorsal, apical and inner margins serrate, set with stout 
setae. Inner margins parallel. From the base of the left clasper, the upper margin is produced 
upwards in a long finger, which bifurcates about mid-way into two spines. The outer spine is 
directed tailward and is setose apically, the inner (and longer) spine is bent downwards alongside 
the aedeagus. In the right clasper of the type, this basal branch is either asymmetric or damaged, 
taking a bigger sweep basally and its apex is broken away. 



NEW AFRICAN CADDIS-FLIES 107 

$ ALLOTYPE. General appearance much as in male. Basal segment of antenna similarly 
elongate, but without the dorsal flap. Wings rather denuded but apparently with similar 
markings. 

9 GENITALIA. Eighth stemite with its apical margin sinuous, the margin itself and the area 
anterior to it densely setose. Ninth and tenth tergites fused, forming a short, truncate anal 
tube. From the dorsal surface of the fused segments arise two short, unequal, transparent 
processes, directed caudad. Sides of anal tube raised to form two low, hairy warts. Lateral 
gonapophyses short, foliate, with rounded apices, directed obliquely downward, outer surfaces 
convex and setose. Beneath them is a broad, bifid subgenital plate, deep in side view, the branches 
directed obliquely upwards between the lateral gonapophyses. From beneath, the branches 
are separated by a deep narrow excision. 

Length of fore wing, c?, 8-5 mm.; ?, 9-9-5 mm. 

cJ HOLOTYPE (now mounted as microscope preparations), $ allotype, pinned 
(abdomen mounted as microscope preparation) and $ paratypes in British Museum 
(Natural History). These specimens were originally determined by Mosely as 
Triaenodella hastata (Ulmer), but even allowing for the fact that Ulmer's figure of 
the male genitalia was drawn from a dried example, I do not think that they could 
justifiably be placed as hastata. The latter is shown with definite curved, blade-like 
basal spines arising from the clasper and the basal branch is clavate, not bifid. 
T. moselyi is closely related to T. hickini Kimmins (1957 : 22) from Kenya, both in 
male and in female genital structures. In the male the centre branch of the median 
lobe of the tenth segment is much longer than the side branches and the two side 
lobes are fused into a single spine. In side view, the clasper is more rectangular and 
in ventral view the inner margins are parallel to each other and to the outer margins. 
In the female, there are two dorsal processes on the fused ninth and tenth segments 
and the lower lip of the anal tube is not produced and the lateral gonapophyses are 
rounded (not acute) apically. 

Triaenodes uncata sp. n. 

(Text-figs. 68, 69, 76-81) 
Uganda : Mbanga Forest, Mpigi, P. S. Corbet, i c^, 4 $. 

cj HOLOTYPE (in alcohol). General colour pale tawny yellow. Antenna finely annulated with 
fuscous, basal segment without dorsal flap but with a small, median, longitudinal carina on 
apical half of dorsal surface, which curves over outwards. Fore wing with golden pubescence 
and traces of fuscous pubescence (rather denuded). Apical half of stem of M present. Cui,, 
represented by a weak cross-vein from anastomosis to Cm2 + A. In hind wing, i?4+5 and M1+2 
confluent for a short distance. 

(^ GENITALIA. Resembling T. wambana Mosely (1939 : 15), but differing in detail. Ninth 
segment with prominent, triangular side-pieces and below them there is a triangular excision 
of the lateral margin. Tenth segment comprising a pair of long, slender spines, strongly arched 
in side view, their apices crossing in the type. Above these spines (which are fused basally) 
are two long, slender cerci and two short processes. Aedeagus arched basally, with a median 
dorsal groove, apical part membranous. Clasper stout, about twice as long as wide in side view, 
its upper margin elevated about mid-way, then excised before the acute apex. Lower margin 
sinuous in side view, apex obliquely truncate. In ventral view, the clasper tapers gradually from 
base almost to apex and is then suddenly constricted to a finger, curving inwards from the outer 
angle. Inner margin of the clasper produced upwards in apical half to form a rounded setose 
lobe. At the extreme base of the clasper there arises, from the dorsal margin, a strong, sickle- 
shaped spine, adjacent to the stem of the aedeagus. Just posterior to this spine arises the basal 



io8 



D. E. KIMMINS 



branch, slender bcisally, directed obliquely upward and caudad, dilating apically and setose. 
This branch is very thin and lightly sclerotized and in the course of mounting has twisted 
spirally and changed its direction. 

? ALLOTYPE (in alcohol). General appearance as in male. Antenna without median carina 
on basal segment. In fore wing there is a weak, incomplete vein representing the basal half of 




Figs. 76-81. Triaenodes uncata sp. n. GenitaUa. 76, cj, lateral; 77, S, tenth segment, 
dorsal ; 78, ^, claspers, ventral ; 80, ?, lateral ; 81, ?, ventral. 



the stem of M. Cui,, normal. In hind wing, 7^4+6 and M1+2 touching or hnked with a short 
cross-vein. 

? GENITALIA. Apical margin of eighth sternite straight, setose. Ninth and tenth segments 
fused, forming a short anal tube, ventral margin of which is produced in a short spatulate lobe. 
Sides of anal tube convex, with long setae. Lateral gonapophyses large, ovate, outer surfaces 
convex. Subgenital plate complex ; on each side at the base is a deep, ovate hollow. From 
beneath, the plate is broad, with rounded lateral margins and a raised median carina at base. 



NEW AFRICAN CADDIS-FLIES 109 

The anterior margin has a wide, V-shaped excision, outer angles acute. Beyond the excision is 
another triangular plate, convex and with a deep and narrow apical excision. This plate is 
directed obliquely upwards between the lateral gonapophyses. 
Length of fore wing, ^, ?, 7 mm. 

^ HOLOTYPE, $ ALLOTYPE (mounted as microscope preparations) and $ paratypes 
in 2% formaldehyde solution, in British Museum (Natural History). The male 
genitalia of this species are similar in structure to those of T. wamhana Mosely. 
The chief differences are in the clasper, which in .side view is shorter and obliquely 
truncate at apex ; the inner marginal lobe is smaller ; the sickle-shaped spine has 
no spur. From beneath, the clasper tapers much more abruptly at its apex. 

Parasetodes maguira (Mosely) 

Leptocerus maguirus Mosely, 1948 : 32, figs. 1-4. 
Parasetodes maguira (Mosely) Kimmins, 1956 : 124. 

Mosely describes and figures the upper branch of the clasper as being clavate or 
truncate at the apex. Examples from the Vaal and Wilge Rivers have genitalia 
almost identical with the figure of maguira, apart from the fact that the apex of the 
upper branch of the clasper is produced in a slender finger. Mosely 's description was 
based upon a single example ; I have studied the preparation carefully and, under 
a 1^ in. objective, it is evident that the extreme apex is damaged and it is probable 
that there was originally a similar slender finger in the type of maguira. 

Athripsodes harrisoni (Barnard) 

Leptocerus harrisoni Barnard, 1934 • 337~"338. ^^%^- 25-26. 
Leptocerus calcaratus Barnard, 1941 : 106-07; fig. b (syn. nov.). 

Barnard distinguished his species calcaratus from harrisoni on the presence of an 
acute tooth on each side of the ninth segment (loth tergite of Barnard) and slight 
differences in the shape of the clasper. These differences seem to me to be no more 
than individual variation, especially as a paratype of harrisoni shows traces of the 
lateral acute processes on the ninth segment. 

OECETIS McLachlan 

1877 : 329-330 (type-species Leptocerus ochraceus Curtis, 1825). 

Setodellina Lestage, 191 9 : 202, fig. 5. (type-species Setodellina albopunctata Lestage, 1919) 
(syn. nov.). 

The genus Setodellina was based by Lestage upon a single example, but in neither 
the generic nor the specific description does he give any indication of the sex of this 
example, merely remarking that the genitalia are difficult to describe on account of 
the bad state of the apex of the abdomen. He figures the venation and compares 
it with Pseudosetodes Ulmer. No comparison is made with Oecetis, with which genus 
it agrees in most characters other than the absence of fork R^ in the hind wing, a 
character which has since been found to be variable in the genus Athripsodes. His 
figure shows Sc and R^ to be completely fused in the fore wing but no comment is 



no D. E. KIMMINS 

made on this point ; in the hind wing these veins are shown as separate for most of 
their length but in his description they are said to be almost fused and scarcely 
visible. 

Ulmer (1923 : 15) placed a second species (S, brunnescens) in the genus Setodellina, 
figuring the venation and the male genitalia. The hind wing of this species also lacks 
the fork R^, but the wings are broader than in the figure of 5. alhopunctata and Ulmer 
comments that they are otherwise similar to those of Oecetis. The genitalia of S. 
brunnescens resemble those of Oecetis gradata, both having the cerci more or less 
fused to the tenth segment, forming a broad plate, Martynov (1936 : 271) placed a 
number of Asiatic and African species into Setodellina and gave an extended diagnosis 
of the genus, including genital characters, particularly the fused cerci. As regards 
venation, he remarks that apical fork R^ may be present or absent in the hind wing, 
a fact that lessens its value as a character separating Setodellina from Oecetis. As 
far as I known, Martynov did not have the opportunity of studying the type of 
S. alhopunctata and the genital characters which he gives for Setodellina are based 
upon S. brunnescens Ulmer, upon his own new species and upon various other species 
of Oecetis, which he had transferred to Setodellina. In my own paper (1956 : 135), 
I accepted Martynov's interpretation of the genus Setodellina, but as material of 
further African species has come to hand it has become increasingly difficult to 
decide when the cerci cease to be fused with the tenth segment to form a broad plate 
or anal tube, some species being intermediate in this respect between Setodellina 
and Oecetis, as defined by its type-species, 0. ochracea (Curtis). Both have the inflated, 
somewhat globular aedeagus with a downwardly-hooked apex. 

Thus a conception of the genus Setodellina had grown up which was based largely 
upon species subsequently placed in it and which were not necessarily congeneric 
with it. I have been able to study the holotype of Setodellina albopunctata Lestage. 
The specimen as sent to me lacked the preparation of the hind wing (Lestage had 
removed both left wings for figuring and attached them by their bases to a card with 
gum ; the hind wing has at some time become detached from the gum and was miss- 
ing). For greater safety, I have now mounted the left fore wing and the right hind 
wing (which was somewhat damaged) between two pieces of celluloid. It is clear 
from these wings that Lestage's figure of the venation is incorrect as far as the sub- 
costa and radius of both wings are concerned. I can see no definite evidence of fork 
R2 in the remaining hind wing, although there is a slight thickening of the vein at 
the point where the fork might be expected to occur. A female of albopunctata in the 
British Museum (Nat. Hist.) from Nigeria, Gadau, has a definite fork R^ in the hind 
wing. I am refiguring the wings of the type (with fore wing reversed) and have 
attempted an impression of the pigmentation of the fore wing. The pattern is rather 
more definite than I had expected from Lestage's description and recalls that of 
examples from Uganda determined by myself as 5. maculipennis (Ulmer). Examina- 
tion of the abdomen of the type of albopunctata suggested that it was a female, an 
impression which was confirmed by clearing in KOH solution. 

Comparison of the wing venation of the female holotype of albopunctata with 
Ulmer's figure of maculipennis (1922 : 61) reveals one difference which, though not 
of generic importance, may be considered specific (Text-figs. 8ia-c). In the fore 



NEW AFRICAN CADDIS-FLIES iii 

wing of albopunctata, M forks basad of the r-m cross- vein. M^j^^ simulates a cross- 
vein from M to Cu^^ ^"^^ fuses with the latter vein for a short distance, appearing 
as a short stalk to the false fork formed by the divergence of M3_,_4 and Cu^^- ^^ 
Ulmer's figure (and in the Uganda examples now placed as maculipennis) the fork 
of M is situated beyond the r-m cross-vein and M^_^^ does not fuse with Cwj^ but is 
linked to it by a cross-vein. The use of this character to separate Setodellina from 




81 A 





Figs. 8ia-c. Oecetis spp., wings. 8ia, 0. albopunctata (Lestage), ? holotype ; b, presumed 
male of O. albopunctata ; c, O. maculipennis Ulmer, male. 



112 D. E. KIMMINS 

Oecetis is not supported by other characters in the various species examined and it 
does not seem worth while to maintain any longer the separation of these two genera. 

Oecetis albopunctata (Lestage) 

(Text-figs. 8lA, B, D, E, H-j) 

Setodellina albopunctata Lestage, lyiq : 202-203, fig- 5 (Congo). 

Uganda: Albert Nile, Pakwach, 28-29, iv. 1956, near Laropi, 27-28. iv. 1956, 
P. S. Corbet, S- 
Nigeria : Gadau, ii.1933, Buxton & Lewis, i $. 
S. Africa : Pongola River, Pongola Settlements, 17. ix. 1954, i c^. 

$ GENITALIA OF HOLOTYPE. Eighth stcmite produced at the centre of its apical margin in an 
acutely cuspidate subgenital plate, which is margined at its base by a narrow band of brownish 
pigment, which simulates an excision. Ninth tergite rather short and deep, the centre of its 




Figs. 8id-g. Oecetis spp., female genitalia. 81D, O. albopunctata (Lestage), holotype, 
lateral ; e, the same, ventral ; f, O. macidipennis Ulmer, lateral ; g, the same, ventral. 



NEW AFRICAN CADDIS-FLIES 113 

apical margin acutely produced in side view. Tenth segment and cerci fused to form an anal 
tube with upper and lower margins produced. Lateral gonapophyses large, thin, the ventral 
margin of each turned outwards, slightly hairy, forming a pair of somewhat reniform lobes in 
ventral view. 

Amongst examples from various localities, originally determined as maculipennis Ulmer, a 
short series of males from Albert Nile, Pakwach has the form of M in the fore wing basad of the 
r-tn cross-vein and the pigmentation of the membrane much as in the type of albopunctata. 
Fork i?2 is present in the hind wing (although I do not attach much importance to this), the 
examples are smaller than maculipennis and show certain differences in male genitalia. It is 
proposed to consider them as the probable male of O. albopunctata (Lestage). 

cj GENITALIA. Ninth segment with its dorsal apical margin moderately produced at its centre. 
Side-pieces broadly triangular. Cerci ovate, almost completely fused to the tenth segment, 
which projects somewhat beyond it in a convex plate with a shallowly excised apical margin. 
Aedeagus short, moderately inflated, its apex produced downwards in a short lobe, which is 
rounded in side view (acute in S. African example). Clasper slender in side view, extending 
beyond the tenth segment. It curves slightly upwards, upper margin sinuously curved. The 
apex is not strongly curved upwards beyond mid- way. From beneath, the clasper is moderately 
broad at base, apical half constricted and incurved, inner and outer margins sinuous. 

Oecetis maculipennis Ulmer 

(Text-figs. 81C, F, G, k-m) 

1922 : 61-63, figs. 22-25 (Egyptian Sudan). 

Specimens from Uganda and Ghana determined by myself as maculipennis agree 
fairly well with Ulmer's figures. New figures of the ^ genitalia are given from an 
example from Lake Victoria, for comparison with albopunctata, in which it will be 
seen that the side-pieces of the ninth segment are narrower than in the presumed 
albopunctata, and the claspers are broader basally in side view and are curved sharply 
upwards in the apical half. The chief difference between these figures and those given 
by Ulmer is that in ventral view the claspers are incurved apically, not divergent as 
shown by Ulmer. 

The female genitalia also follow the pattern of albopunctata, but as in the male, 
they differ in detail. The ninth tergite in side view has the apical margin almost 
straight and the lower lateral margin is much wider. The lateral gonapophyses are 
rather shorter and the apical margin in side view is less obliquely truncate. In ventral 
view the apical margin is sinuous. Subgenital plate more abruptly shouldered. 
Cerci less closely fused to the tenth segment. 

Oecetis absimilis sp. n. 

(Text-figs. 82-88) 
Uganda : Mbanga Forest, Mpigi, P. S. Corbet, 2 ^. 

J HOLOTYPK (in alcohol). General colour pale ochraceous. Antennae with fine piceous annula- 
tions, basal segment tinged with pale fuscous. Palpi and anterior legs with sparse fuscous 
pubescence. Anterior wing with sparse pale fuscous pubescence, the membrane marked with 
fuscous as in the figure. 

<J GENITALIA. Ninth segment narrowed dorsally. Tenth segment forming a narrow, hood-like 
plate, dilated basally in dorsal view and with the apex slightly excised. At its base, on the dorsal 
surface, arise two very short, transparent processes, one on each side of a longer, slender finger, 
bearing a few setae. Cerci short and blunt, rather less than half the length of the tenth segment. 



"4 



D. E. KIMMINS 







Figs. 8ih-m. Oecetis spp., male genitalia. 8ih, presumed O. albopunctata (Lestage) 
lateral ; i, the same, dorsal ; j, the same, ventral ; k, O. maculipennis Ulmer, lateral • 
L, the same, dorsal ; m, the same, ventral. 



NEW AFRICAN CADDIS-FLIES 



1^5 



Aedeagus short, strongly arched downwards, its apical part with a median dorsal furrow, in 
which arises a short, slender, arched spine. Clasper large, with a flattened, erect basal lobe, its apex 
broader than its base, slightly concave in side view and armed with strong setae. The main 
part of the left clasper is roughly rectangular in side view, slightly arched and with the upper 
apical angle rounded. From above or beneath, the clasper is subtriangular, upper margin bearing 
a few stout setae, inner surface concave apically. The upper margin of the right clasper is 




Figs. 82-88. Oecetis absimilis sp. n. ,$■ 82, wings ; 83, genitalia, lateral ; 84, aedeagus, 
lateral ; 85, tenth segment, dorsal ; 86, apex of aedeagus, dorsal ; 87, claspers, dorsal ; 
88, the same, ventral. 



produced in an inwardly directed, spatulate lobe, overlapping the left clasper. Internally, near 
the base of each clasper, is a strongly setose lobe. 
Length of fore wing, ^, 6 mm. 

(^ HOLOTYPE (in 2% formaldehyde solution), <^ paratype mounted as microscope 
preparations, in British Museum (Nat. Hist.). This species appears to belong to the 
kathia-angustipennis group (Mosely, 1939 : 17-18), in which there is a basal lobe on 
the clasper. In 0. absimilis, this lobe is more developed than in most of them and it 
also differs from them in having the tenth segment in the form of a hood rather than 
a pair of spines. The specific name refers to the dissimilarity in the form of the right 
and left claspers. 



Ii6 



D. E. KIMMINS 



Setodes njala sp. n. 

(Text-figs. 89-94) 

SiHKKA Leone : Njala, i.iv.1932, E. Hargreaves, i (S, " Setodes sp. n. det. M. E. 
Mosely ", i ?, " Setodes sp. det. M. E. Mosely ". 

^ HOLOTypE (pinned). Head tawny yellow, with golden pubescence and three snuw-white 
longitudinal lines on vertex. Antenna tawny yellow, basal segment with a .snow-white line 




Figs. 89-94. Setodes njala sp. n. 89, wings,- ^ ; 90, S, genitalia, lateral ; gi, ^, dorsal 
92, c?. claspers and aedeagus, ventral ; 93, $, genitalia, lateral ; 94, $, ventral. 



NEW AFRICAN CADDIS-FLIES 117 

externally. Palpi tawny. Thorax tawny, with golden pubescence and three snow-white longi- 
tudinal lines. Legs pale luteous. Fore wing with dense golden brown pubescence, most of the 
cells with a longitudinal white line bordered with fuscous pubescence. Apical cells with a white 
spot bordered with fuscous. Hind wing hyaline. 

cJ GENITALIA. Ninth segment rather short, its dorsal margin short and fused with the tenth 
segment. Cerci about as long as the ninth segment, slender, somewhat flattened dorsoventrally. 
Tenth segment produced in two long, slender, slightly sinuous spines arising from a common 
base. Aedeagus short, stout, apex downcurved, upper surface membranous and enclosing a 
shorter, stout spine. Clasper short and stout, the lower part forming a rounded plate, whose 
inner margin is slightly produced. Upper portion produced in two short processes, the upper 
incurved and acute, the lower digitate and hairy. 

? ALLOTYPE. Resembling the male but rather smaller. $ genitalia. Ninth segment forming 
a complete ring. From its upper margin project two short, digitate lobes, bearing a few setae. 
Centre of the ventral margin with a small rounded depression, giving the appearance of an 
excision. Lateral gonapophyses large, subquadrate, from the side, shallowly excised at the apex. 
From beneath, they are acute apically, lower margin carinate, inner surface convex, outer 
surface concave basally. Tenth tergite reduced to a small, rounded, transparent knob between 
the digitate processes of the ninth segment. Within the ninth segment is a shallow, convex 
subanal plate, with a rounded apex. 

Length of fore wing, cJ, 4 mm.; $, 37 mm. 

^ HOLOTYPE pinned (with one pair of wings and the abdomen mounted as micro- 
scope preparations), $ allotype pinned (abdomen cleared and in glycerine), in 
British Museum (Nat. Hist.). The markings of the fore wing recall those of Hemilep- 
tocerus gregarms Ulmer (1922 : 56) and H. hargreavesi (Ulmer, 1931 : 25), but the 
hind wing is relatively less wide than in these species. The types are smaller than 
any described species of Setodes from Africa known to me, and the ^ genitalia are 
also different. 



Trichosetodes triangularis sp. n. 

(Text-figs. 95-101) 

S. Africa : Orange Free State, Swinburne, 9.xii.i958, i <^. 

(^ HOLOTYPE. The specimen was collected in alcohol and is rather rubbed and 
faded. The wings bore only traces of pubescence and no description of the wing 
pattern (if any) can be given. The hair tufts on the basal segment of the antennae 
are likewise abraded, but these segments are of the typical elongate, cylindrical form 
of Trichosetodes. The wings are more acute apically than in T. anysa Mosely (1948 : 39), 
semibrunnea Ulmer (1923 : 13) or lacustris Kimmins (1953 : 278). 

cJ genitalia. Conforming to the general pattern of the African species. Ninth segment cut 
back dorsally to a narrow, transverse band. Tenth segment produced in a pair of simple, slender 
spines, extending beyond the apices of the claspers. Cerci short, about one-third the length of 
the spines of the tenth segment. Aedeagus broad at its base in side view, arched sharply down- 
ward and tapering to a blunt apex. From its dorsal surface arises a strong, acute paramere, 
arching over the aedeagus and separated from it by a wide gap. Clasper triangular in side view, 
stout at base, apex curving upward and acute. On its upper surface arise two processes, the 
basal one the longer and more slender, its apex triangularly dilated inwardly in dorsal aspect. 
The second one arises about mid-way, is about half as long as basal and is stouter. From beneath, 
the clasper is convex on its outer margin. Inner margin more or less straight, with a small 



xi8 



D. E. KIMMINS 




Figs. 95-101. Trichosetodes spp. S- 95-98, T. triangularis sp. n. 95, wings ; 96, 
genitalia, lateral ; 97, dorsal ; 98, claspers, ventral ; 99, T. lacustris Kimmins, aedeagus 
and claspers, lateral; 100, T. anysa Mosely, the same aspect; 10 1, T. semibrunnea 
Ulmer, same aspect. 



NEW AFRICAN CADDIS-FLIES 119 

excision in its basal half. Ventral apical margin of ninth segment shallowly excised at its 
centre. 

Length of fore wing, 7 mm. 

^ HOLOTYPE mounted as microscope preparations, in British Museum (Nat. Hist.). 
This species differs from the other described African species in its stout, triangular 
clasper and the wide separation of the paramere and aedeagus. This very noticeable 
feature led me to re-examine the preparations of the other species. Indications of a 
slender paramere occur in both semibrunnea and in anysa, but there is no evident 
paramere in lacustris. Figures are given of the lateral aspect of the aedeagus and 
clasper of the other three species for comparison. 



Leptocerus inflatus sp. n. 

(Text-figs. 102-107) 

S. Africa: Transvaal, Vaal River, above Klip confluence, x.1957, 2 J. i ? i 
Vaal Barrage, 1957, 2 c^, 4 ?. 

(J HOLOTYPE (collected in alcohol). Head brown, two basal segments of antenna pale brown, 
remaining segments pale luteous, annulated with piceous. Palpi dull brownish, with fuscous 
pubescence. Thorax brown. Legs very pale brownish. Fore wing with fuscous pubescence, with a 
patch of creamy white, broadened hairs on anterior margin near apex of Sc, extending triangularly 
as far as anterior margin of discoidal cell, and with iridescent scales extending to posterior margin 
of cell. There are traces of creamy white hairs just basad of the discoidal cell and a patch of 
iridescent scales between Rs and Cu^a at the same level. A few scattered iridescent scales near 
base of wing. Wings narrow, acute, hind wing tapering more gradually than in L. neavei (Mosely, 
1932a : 130). 

(J GENITALIA. Ninth segment reduced dorsally to a narrow, transverse band. Ventral margin 
slightly produced, with a small median excision. Tenth segment with an ovate median lobe, 
apex truncate, in side view raised above the side lobes. The latter consist of two asymmetric 
blade-like spines and two slender digitate processes, about equal in length to the longer of the 
spines and terminating in a few setae. Aedeagus strongly arched downward, with a median 
dorsal trough in the apical third, apex narrowed and rounded in ventral view. On each side, at 
about mid-way, the upper margin is expanded in a membranous, bilobed process, covered with 
minute setae. Clasper gently sinuous in side view, without the basal ventral projection present 
in the other described African species. Its dorsal margin towards the base is slightly elevated. 
In ventral view, the clasper is broad at its base, tapering to a narrow, slightly incurved apex. 

9 ALLOTYPE (collected in alcohol). General appearance much as in male, but the iridescent 
scales on the fore wing are few, possibly denuded. 

? GENITALIA. Eighth stemite produced in a more pigmented subgenital plate, its apical margin 
triangular, lateral margins slightly concave, apex rounded. Ninth and tenth tergites fused, 
the tenth more sclerotized and bearing patches of setae, apical margin rounded. Lateral gonapo- 
physes extending beyond the apex of tenth segment, elongate, slightly dilated and obliquely 
truncate. 

Length of fore wing, ,S, ?, 7 mm. 

^ HOLOTYPE (mounted as microscope preparations), $ allotype (in 2% formal- 
dehyde solution) in British Museum (Nat. Hist.), c^, $ paratypes in Dr. Scott's 
collection. This species is perhaps closest to L. neavei (Mosely) in male genital struc- 
ture, but it differs from all the described African species of Leptocerus in the absence 
of the ventral projection at the base of the clasper. 



D. E. KIMMJNS 




Figs. 102-107. Leptocerus inflatus sp. n. 102, cJ, wings ; 103, ^, genitalia, lateral ; 104, 
(J, dorsal ; 105, (J, claspers, ventral ; 106, ?, genitalia, lateral ; 107, ?, ventral. 



BIBLIOGRAPHY 

Barnard, K. H. 1934. South African Caddis-flies. Trans. R. Soc. S. Afr. 21 : 291-394, 52 figs. 
1941- May-flies and Caddis-flies from Natal, Basutoland and Pondoland. Ann. Durban 

Mus. 3 : 105-108, figs. A-c. 
Brauer, F. 1875. Beschreibung neuer und ungenugend bekannter Phryganiden und Oestri- 

den. Verh. zool. bot. Ges., Wien, 25 : 69-74, P^- 4- 
KiMMiNS, D. E. 1953. Trichoptera collected by Miss R. H. Lowe in Uganda, with descriptions 

of three new species of Leptoceridae. Entomologist, 86 : 274-278, 8 figs. 



NEW AFRICAN CADDIS-FLIES 121 

KiMMiNS, D. E. 1956. New and little-known species of the Leptocerinae (Trichoptera) from 

the African mainland (south of the Mediterranean region). Trans. R. ent. Soc. Lond. 

108 : 1 17-146, 18 figs. 
1957- New and little-known species of African Trichoptera. Bull. Brit. Mus. {not. Hist.) 

Ent. 6 : 1-37, 27 figs. 
Lestage, J. -A. 1919. Notes sur quelques N^vropt^res (Eph6meres et Trichopteres) du Congo 

Beige. Rev. Zool. afr. 6 : 193-204, 5 figs. 
Marlier, G. 1955. Un Trichoptere p61agique nouveau du lac Tanganyika. Rev. Zool. Bot. 

afr. 52 : 150-155. figS- i-3- 
Martynov, a. B. 1935. O'^ ^ collection of Trichoptera from the Indian Museum. Part I. 

Annulipalpia. Rec. Ind. Mus. 37 : 93-209, 109 figs. 

1936. On a collection of Trichoptera from the Indian Museum. Part II. Integripalpia. 

Ihid. 38 : 239-306, 74 figs. 

McLachlan, R. 1872. Mat^riaux pour une Faune n^vropterologique de I'Asie Septentrionale. 

Ann. Soc. ent. Belg. 15 : 60-71, pis. 1-2. 
1874-80. A monographic revision and synopsis of the Trichoptera of the European Fauna. 

iv -f- 523 pp. and Supplements, parts I-II, pp. i-lxxxiv, 59 pis. London, John van Voorst. 
MosELY, M. E. 1932. Contributions k I'^tude de la Faune du Mozambique, Voyage de M. P. 

Lesne (1928-29) 12* Note — Trichoptera. Mem. Est. Mus. zool. Univ. Coimbra, (i) 64 : 1-8, 

pi. I. 

1932a. More African Leptoceridae (Trichoptera). Stylops, 1 : 128-134, i^ figs. 

1936. New African Trichoptera. Ann. Mag. n. H. (10) 17 : 429-451, 48 figs. 

1939. Trichoptera. Ruwenzori Exped. 1934-5, 3 (i) : 1-40, 3 pis., 123 figs. 

1948. Trichoptera collected by Miss R. H. Lowe at Lake Nyasa. Ann. Mag. n. H. (12) 1 : 

31-47, 27 figs. 
Navas, L. 1914. Notes sur quelques N^vropteres du Congo Beige. III. Rev. zool. afr. 4 : 

172-182, pi. 5 

1916. Neuropteros nuevos o poco conocidos (Septima serie). Mem. Ac. Ci. Barcelona, 

12 (13) : 1-27. 

Ross, H. H. 1944. The Caddis Flies, or Trichoptera, of Illinois. ///. nat. Hist. Surv. Bull. 

23 (i) : 1-326, 961 figs. 
Ulmer, G. 1906. Neuer Beitrag zur Kenntnis aussereuropaischer Trichopteren. Notes 

Leyden Mus. 28 : 1-116, 114 figs. 
1907. Neue und wenig bekannte Trichopteren der Museen zu Briissel und Paris. Ann. 

Soc. ent. Belg. 49 : 17-42, 31 figs. 
1907a. Trichopteren (Macronematinae). Collections zoologiques du Baron Edm. de Selys 

Longchamps. Catalogue systematique et descriptif Fasc. 6 (2) : 1-121, 6 pis. 119 figs. 

19076. Trichoptera. Wytsman's Genera Insectorum. Fasc. 60a : 1-259, 41 pis. 

1912. Trichopteren von Aquatorial-Afrika. Wiss. Ergeb. D. Zentral-Afrika Exp. 4 (Zool.): 

81-125, figs. 1-50. 
1922-23. Trichopteren aus dem aegyptischen Sudan und aus Kamerun. Mitt. Miinch. 

ent. Ges. 12 : 47-63, figs. 1-26, 13 : 9-20, figs. 26-36. 
1924. Wissenschaftliche Ergebnisse der mit Unterstiitzung der Akademie der Wissens- 

chaften in Wien aus der Erbschaft Treitl von F. Werner unterbommenen zoologischen 

Expedition nach dem Anglo-Aegyptischen Sudan (Kordofan) 1914. XII. Trichopteren und 

Ephemeropteren. Denks. Akad. Wiss. Wien, Math. Naturw. 99 : 1-9, 9 figs. 
1931. Trichoptera von Afrika (hauptsachlich aus dem Britisch Museum). Dtsch. ent. 

Zeitschr. 1931 : 1-29, 30 figs. 
195 1. Kocherfliegen (Trichopteren) von den Sunda-Inseln (Teil I). Arch. Hydrobiol. 

Suppl. Bd. 19 : 1-528, 28 pis. 



ENTOM. 12, 2. 




PRINTED IN GREAT BRITAIN BY 
ADLARD AND SON, LIMITED, 
BARTHOLOMEW PRESS, DORKING 



> # ' / / . l^ i 



NOTES ON THE EPITOLINAE 

WITH DESCRIPTIONS OF A NEW 

GENUS AND NEW SPECIES AND 

SUBSPECIES (LEPIDOPTERA : 

LYCAENIDAE) 



T. H. E. JACKSON 




BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 12 No. 3 

LONDON : 1962 



NOTES ON THE EPITOLINAE 

WITH DESCRIPTIONS OF A NEW GENUS 

AND NEW SPECIES AND SUBSPECIES 

(LEPIDOPTERA : LYCAENIDAE) 



BY 



T. H. E. JACKSON 

Kapretwa, Kitale, Kenya Y' 



i 




pp. 123-163 ; Plates 1-13 ; i Text-figure 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 12 No. 3 

LONDON: 1962 



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 
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within one calendar year. 

This paper is Vol. 12, No. 3 of the Entomological 
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those of the World List of Scientific Periodicals. 



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NOTES ON THE EPITOLINAE 

WITH DESCRIPTIONS OF A NEW GENUS AND NEW SPECIES 
AND SUBSPECIES (LEPIDOPTERA : LYCAENIDAE) 

By T. H. E. JACKSON 



CONTENTS 



Epitola 

The HEWiTSONi group 

The POSTHUMUS group 

The CARCINA and nitide groups 

The PiNODES group 

The suBLUSTRis group 

The ALBA group 
/ Species incertae sedis 

Species incorrectly placed in Epitola 

Discussion 
Stempfferia . 
Neavia 
Epitolina 
Hewitsonia 
Acknowledgements 
References 
Index 



Page 

126 

128 

130 

142 

152 

155 

155 

155 

156 

157 

158 

159 

160 

161 

161 

163 



SYNOPSIS 

The genus Epitola is brought up to date with further association of the sexes. New species and 
subspecies are described, as well as a new genus. 



Genus EPITOLA Westwood ^ 

The genus Epitola Westwood was revised by P. J. L. Roche (1954). In this excellent 
paper will be found a history of the genus, a check list of the species, a working key 
to the males and descriptions of all new species to that date. Since then, however, 
many new species have been discovered and an attempt is made here to bring the 
genus up to date, and further, to integrate the females. The latter is admittedly 
a difficult task ; it was scarcely touched by Aurivillius in Seitz (1920), and not at all 
by Roche, but considerable evidence has accumulated either from species taken 
in cop., bred, or taken together in large numbers to warrant a preliminary arrange- 
ment, and it is felt that this is long overdue. In addition, a careful study of the 
underside markings is sometimes sufficient. The arrangement of the species in this 
paper largely follows that of Roche but, with the exception of the first two groups 
and the last, they have been rearranged according to the colour and markings of the 
females. This produces a large measure of agreement with Roche's keys, but some 

ENTOM. 12, 3. 8 



126 T. H. E. JACKSON 

interesting discrepancies. With the exception of those of a few type specimens, 
which are not available for study, all records mentioned in this paper have been 
personally checked by the present author or by Monsieur H. Stempffer of Paris. 
All types are in British Museum (Nat. Hist.) unless otherwise stated. 

THE HEWITSONI GROUP 

Epitola crippsi Stoneham 

$ Epitola crippsi Stoneham, 1933 : i, pi. 2. 
Type. Kenya : Soy. (Stoneham Mus., Kenya.) 

^ undescribed : the male of this species was taken by the writer some years ago 
at Busia, Uganda and forwarded to Europe. The specimen has unfortunately been 
mislaid and so the description cannot be included here. The male is almost identical 
with the female, but with sex brands. 

Recorded from : Kenya : Soy ; Uganda : Busia. 

Epitola hewitsoni Mabille 

(PI. 13, figs. 124, 130) 

(J Epitola hewitsoni Mabille, 1877 : 221. 

Type. Angola : Cabinda, Landana. (Paris Mus.) 

Epitola hewitsoni Mabille ; Seitz, 1920 : pi. 65, d. 

Recorded from : Cameroons : Bitje ; Angola : Cabinda. 

Epitola hewitsonioides Hawker-Smith 

(J, $ Epitola hewitsonioides Hawker-Smith, 1933 • i^- 
Types. Congo : Middle Lowa Valley, near Walikale. 

Recorded from : Cameroons : Bitje ; Fr. Equat. Africa : Etoumbi, MambiH 
Forest ; Congo : Equator, Eala, Lomami River. 

Epitola miranda miranda Staudinger 
(PI. I, fig. I, PL 2, fig. 10) 

<J Epitola miranda Staudinger, 1889 : 176. 
Type. Sierra Leone. (Staudinger Coll.) 
Epitola miranda Staudinger ; Seitz, 1920 : pi. 64, e. 

?. Legs dark brown with paler spots at the joints ; palps light brown below with black tips, 
dark brown above ; eyes dark brown ; frons dark brown with two pale lines ; antennal shaft 
black above, minutely checkered white below ; club with orange tip. 

Upperside forewing : brown ; a subapical series of elongate yellowish spots in 5, 6 and 7, 4 mm. 
long, and a yellow streak in 8. A faint pale spot astride vein 2. 

Upperside hindwing : brown, immaculate. 

Underside : similar to the male, but paler, the yellow fore wing spots reproduced below, but 
paler. 

Length of fore wing : 27 mm. 

Neallotype $. Sierra Leone. Rothschild bequest. 

Recorded from : Sierra Leone ; Nigeria : Lagos Dist., Oshodi and Kaba 
Prov. : Cameroons. 



NOTES ON THE EPITOLINAE 127 

Epitola miranda vidua Talbot 

(PI. I, fig. 2, PL 2. fig. II) 

$ Epitola miranda vidua Talbot, 1935 : 75. 

Type. Uganda: Sesse Isles, Bugalla Is., Lutoboka. (Hope Dept., O.xford.) 

This is not a well-differentiated subspecies and the male differs scarcely at all from 
miranda miranda. 

cJ. Legs dark brown, faintly paler at the joints ; palps light brown below with black tips, 
dark brown above ; eyes and Irons brown ; antennal shaft black above, very minutely checkered 
white below ; club with small yellow tip. 

The following small points may be noted : 

On fore wing above the blue spot in space 2 is narrower and the spots in 4, 5 and 6 are longer ; 
in hind wing the distal border is narrower, especially at anal angle. Below the postdiscal pale 
spots on fore wing are longer and more distinct, especially in 2 and 3. Otherwise as in the typical 
race. 

Neallotype ^. Uganda: Bwamba, iv.1942. T. H. E. Jackson. 

Seen flying rapidly across the open, between trees on an open hillside at Katera, 
Buddu about 11 a.m. ; the females laying on lichen-covered, ant-infested tree trunks 
later in the day. 

Recorded from : Uganda : Katera, Kayonza, Kigezi and Bwamba, Sesse Islands, 
Lutoboka, Bugalla Island, Siroko River near Mt. Elgon. 

Epitola decellei Stempffer 

(J Epitola decellei Stempffer, 1956 : 30, fig. 16, genitalia pi. 2, figs. 11, 12. 
Type. Congo : Stanleyville, Yangambi. (M.R.C.B., Tervuren.) 

$. Unknown. 

Recorded only from the type locality. 

Epitola stempfferi sp. n. 

(PI. 13, figs. 122, 123, 128, 129) 

cJ and 9. Closely allied to hewitsoni Mabille and decellei Stempffer, and of the 
same shape. 

Legs dark sepia, unspotted ; palps dark brown above and below ; frons and eyes dark brown ; 
antennal shaft black above, checkered white below ; club black. 

cj. Upper side fore wing : black with brilliant greenish blue basal and discal areas ; costal 
border black up to the upper median and thereafter linear to the apical patch ; apex and margin 
broadly black ; blue area extends from inner margin to upper median, covering i and 2 to within 

4 mm. of distal margin, the bases of 3, 4 and 5 and the whole of the cell ; thereafter three blue 
streaks postdiscal in 7, 8 and 9 and two small blue spots in 5 and 6 ; large V-shaped wedge of 
black ground colour invading the blue area above apex of cell. 

Upperside hind wing : black with blue basal discal patch from veins 1-7 with a blue streak 
in the base of space 8 ; distal margin of this patch irregular leaving a broad black distal border 

5 mm. wide on vein 5 and 3 mm. wide on vein 2 ; space ib black throughout. 

Underside fore wing : smoky brown with lighter patch along inner margin nearly to vein 2 
and a series of small obscure whitish subapical spots placed obliquely in spaces 5-7 ; a slightly 
paler submarginal area between veins 3-5. 



128 T.H.E. JACKSON 

Underside hind wing : clear brownish violet with black or brown spots and markings ; three 
Small black sub-basal spots, the first just below 8, the second in the cell and the third between 
vein 2 and the lower median ; five rounded black discal spots, the first below vein 8, the second 
and third confluent and in the cell, the fourth between vein 2 and the lower median, and the 
fifth between 2 and ib ; a black spot on the lower discocellular ; a small, triangular black spot 
between the lower median and the origin of vein 2 ; a fine, very irregular postdiscal black line 
running between 6 and 2 ; a series of large confluent, submarginal brown spots, forming a broad 
band, nearly continuous from apex to anal angle. 

This species may be distinguished at once, in the male, from its nearest neighbours, 
decellei Stempffer and hewitsoni Mabille by the much broader black border in hind 
wing and by the invasion of the blue band in fore wing by a wedge of the black ground 
colour. 

It is worthy of note that both the above are female characters and this is the only, 
species known with such characters in the male. 

?. Upperside fore wing : brown with blue patch from base to beyond the middle, bounded 
along the costa by the subcostal vein and reaching the inner margin. Distally covering the base 
of spaces 4, 5 and 6, projecting sharply distad in 2 and 3 and again in i and la. A series of small 
blue postdiscal spots in 5, 6 and 7. 

Upperside hind wing : brown with large blue patch reaching to within 2 mm. of the margin 
in I and 3 and gradually receding to 6 mm. on vein 6. Bounded costally by vein 6 but with a few 
blue scales in the base of space 6 above it. The distal margin of the blue area very dislegnic. 

Underside : exactly as in the male but with larger postdiscal spots in fore wing. 

Length of fore wing : cj 24 mm., ? 27 mm. 

HolotypCf^. Gaboon: Douano. 27.ix.1951. P. Rougeot leg. Coll. Stempffer. 

Allotype ?. Fr. Equat. Africa: Ouesso, Ketta Forest, vii.1959. T. H. E. 
Jackson. 

Recorded from : Gaboon : Douano ; Fr. Equat. Africa : Ouesso ; Congo : 
Buta Dist., Yopole. 

This species is named in honour of Monsieur H. Stempffer of Paris. 

THE POSTHUMUS GROUP 

Epitola posthumus (Fabricius) 

J Papilio posthumus Fabricius, 1793 : 149. 

Type. Missing. 

Epitola posthumus (Fabricius) ; Seitz, 1920 : pi. 64, e. 

? Epitola belli Hewitson, 1874 : 382. 

Recorded from : Nigeria ; Br. Cameroons ; Fr. Cameroons ; Ghana ; Fr. 
Equat. Africa ; Congo ; Uganda. 

Epitola urania urania Kirby 
(PI. I, fig. 3, PI. 2, fig. 12, PL 9, fig. 85, PI. 10, fig. 95) 

(? Epitola urania Kirby, 1887 : 441. 
Type. Cameroons. 

$. Legs black with small pale brown lines at joints ; palps black above and below, with a 
small white spot above ; frons black with white edges ; eyes black ; antennal shaft black above 
and below ; club yellow-tipped below. 



NOTES ON THE EPITOLINAE 129 

Upper side fore wing : black with blue basal and discal area bounded by the subcostal and 
reaching the inner margin, distally covering the base of spaces 4, 5 and 6 for 5 mm.; projecting 
distad in 3 and 2 and to within 3 mm. of the margin in i, and 4 mm. in la. Small blue spots 
half-way along the costa in 7 and 8 and streaks in 9 and 10. 

Upperside hind wing : black with large blue discal patch reaching to the base and bounded 
by veins i and 7. A wide black border distally, 7 mm. on vein 6 but becoming narrower towards 
the anal angle, where the blue colour almost reaches the margin. 

Underside : as in the male. 

Length of fore wing : 35 mm. 

Neallotype $. Cameroons : Suffer! bequest. 

Recorded from : Sierra Leone ; Ivory Coast ; Liberia ; Nigeria ; Gaboon ; 
Fr. Equat. Africa. 

Epitola urania tanganikensis Joicey & Talbot 
(PI. I, fig. 4. PI. 2, fig. 13) 

^ Epitola urania tanganikensis Joicey & Talbot, 1921 : 86, pi. 14, fig. 63. 
Type. Belgian Congo : Albertville. 

$. Legs, eyes, etc. as in the typical race, but antennal shaft checkered white 
below. 

Agrees with the description of the male, loc. cit., except for the presence of a costal 
spot, in 7 and the reduction of the blue area in hind wing, as in the typical female. 

Neallotype $. Uganda: Toro, Kibale Forest, ii.1951. T. H. E. Jackson. 

Males observed flying, briefly, high up in Kibale Forest, Toro, and settling on 
creeper tendrils ; females laying on tree trunks among Crematogaster ants. 

Recorded from : Congo : Albertville, Katanga ; Uganda : Bwamba and Toro. 

Epitola crowleyi crowleyi Sharpe 

(J, $ Epitola crowleyi Sharpe, 1890 : 106. 

Types. Sierra Leone. 

(J, $ Epitola crowleyi Sharpe ; Seitz, 1920 : pi. 64, e. 

Recorded from : Sierra Leone ; Nigeria ; Cameroons ; Fr. Equat. Africa ; 
Ghana ; Congo ; Uganda. 

Epitola setnibrunnea Baker 

(J Epitola crowleyi var. semibrunnea Baker, 191 6 : 378. 
Type. Cameroons. 

$ Epitola ammon Joicey & Talbot, 1921 : 83, pi. 14, figs. 60, 61. 
Type. Congo : Penghe, North bank, Ituri River. 

Only the types are known to the author. 

Epitola ceraunia Hewitson 

(J Epitola ceraunia Hewitson, 1879 : 149. 

Type. West Africa (Rutherford). 

Epitola ceraunia Hewitson ; Seitz, 1920 : pi. 64, e. 

$ Epitola dewitzi Kirby, 1887 : 442. 

Type. Cameroons. Bred. 



130 T. H. E. JACKSON 

Recorded from : Sierra Leone ; Ivory Coast ; Liberia ; Togo ; Ghana ; 
Nigeria ; Br. Cameroons ; Fr. Cameroons ; Fr. Equat. Africa ; Congo ; 
Uganda. 

Epitola adolphifriderici Schultze 

<J, $ Epitola adolphi friderici Schultze, 191 1 : 95. 
Epitola adolphifriderici Schultze ; Seitz, 1920 : pi. 65, b. 
Types. South Cameroons : Yukaduma. (Coll. Senckenberg.) 



THE CARCINA AND NITIDE GROUPS 

A, $$. Brown with large white discal patch in fore wing. 
Epitola marginata marginata Kirby 

(J Epitola marginata Kirby, 1887 : 443. 

T^'pe. Cameroons. 

Epitola marginata Kirby ; Grose-Smith & Kirby, i. 1889 : Lye. Afr. pi. 7, figs. 5, 6. 

$ Epitola marginata Kirby ; Joicey & Talbot, 192 1 : 85, pi. 14, figs. 56, 57. 

Type. Congo : Albertville. 

Recorded from : Cameroons : Bitje, Johann Albrechts Hohe ; Nigeria : Pt. 
Harcourt and Bassa Prov. ; Fr. Equat. Africa : Ketta and Mambili ; Congo : 
Albertville ; Uganda : Kigezi, Kayonza. 

Epitola marginata umbratilis Holland 

Epitola umbratilis Holland, 1890 : 425. 

Type. Gabon : " Upper waters of the River Ogove." (Carnegie Mus., Pittsburg.) 

This subspecies is probably a synonym of marginata Kirby. 
Recorded from : Cameroons : Yukaduma, Barombi. 

Epitola moyambina Baker 
(PL I, fig. 5. PI. 2, fig. 14) 

<J Epitola moyambina Baker, 1903 : 330. 
Type. Sierra Leone : Moyamba. 

$ Very similar to the females of cercene Hewitson, convexa Roche and intermedia Roche, and 
about the same size. 

Legs dark brown with white spots at joints ; palps brown, black-tipped with whitish lateral 
spots ; frons brown with white edges ; eyes brown ; antennal shaft black above, checkered 
white below ; club orange-tipped. 

Upperside fore wing : black-brown with white postdiscal band and basal areas blue. Cell, 
half space i and the extreme base of 2 blue. A large triangular spot of the black ground colour 
invading the discal area with apex half-way along the discocellular. Postdiscal white band from 
1-7, its distal edge nearly straight from veins 2-7, and sharply toothed in space 2. A small 
kink basad on vein 7 and narrowing abruptly in space i. The almost straight distal margin 
of the white band distinguishes it from the other members of this group. In specimens from 
Sierra Leone this band is broader. 

Upperside hind wing : dark grey-brown, immaculate. 



NOTES ON THE EPITOLINAE 131 

Underside fore wing : ground colour brown-grey, the white postdiscal band reproduced as 
above but reaching the costa and becoming pale yellow from space 3 to the costa. Grey sub- 
marginal and marginal lines as in the male. 

Underside hind wing : brown-grey with markings as in the male. 

Length of fore wing : 20 mm. 

Neallotype $. Nigeria: Lagos Dist., Oshodi. iv.1955. T. H. E. Jackson. 
Recorded from: Sierra Leone ; Ivory Coast ; Nigeria ; Ghana : Ashanti. 

Epitola cercene Hewitson 

(J Epitola cercene Hewitson, 1873 : 150. 

Type. Angola. 

Epitola cercene Hewitson ; Seitz, 1920 : pi. 65, /. 

9 Epitola versicolor Kirby, 1887 : 444 (syn. n.). 

Type. Cameroons. 

Recorded from : W. Africa to Uganda and Angola. 

Epitola intermedia Roche 
(PI. I, fig. 6, PI. 2, fig. 15) 

cj Epitola intermedia Roche, 1954 • 49^' pl- 20, figs. 11, 12. 
Type. Uganda : Katera. 

9. Similar to moyambina Bethune-Baker and to cercene Hewitson, but distal edge of fore 
wing white patch more or less rounded and underside different. Legs light brown with lighter 
joints ; palps light brown above and below ; frons light brown with white edges ; eyes brown ; 
antennal shaft black above, checkered minutely white below ; club orange-tipped. 

Upperside fore wing : black-brown ; cell and basal third of space i brown with sparse blue 
scales. A broad white postdiscal band with rounded distal edge, just invading the cell proximad. 
A small triangular patch on the discocellular. The white band is slightly dentate in 2 and 3, but 
thereafter rounded and even to vein 9. This band stretches from la-g, as in cercene. 

Upperside hind wing : dark grey-brown ; immaculate. 

Underside fore wing : very pale brownish grey with white band as above, but slightly yellowish 
near the costa markings in apical area faint. 

Underside hind wing : very pale brownish grey with markings as in the male. 

Length of fore wing ; 18 mm. 

Neallotype $. Uganda: Katera. ix.1949. T. H. E. Jackson. 
Recorded from : Uganda : Katera and Budongo. 

Epitola convexa Roche 
(PI. I, fig. 7, PL 2, fig. 16) 

^ Epitola convexa Roche, 1954 : 496, pi. 20, figs. 11, 12. 
Type. Uganda : Bwamba. 

$. Very similar to the females of moyambina Baker, cercene Hewitson and intermedia Roche. 
Legs brown, heavily scaled whitish ; palps light brown above and below ; eyes brown ; frons 
brown edged whitish ; antennal shaft black above, checkered white below ; club yellow- 
tipped. 

Upperside fore wing : black-brown ; basal area of i and cell brown with blue scales, more blue 
than in intermedia but less than in cercene and moyambina. A triangular black spot on the disco- 
cellular, longer and more pointed than in the other species and reaching almost to the lower 



132 T. H. E. JACKSON 

angle of the cell. Broad white postdiscal band, double-toothed in space 2, thereafter narrowing 
in squared steps between 3, 4 and 5, and 6-9, stretching from veins la-g. 

Upperside hind wing : dark grey-brown ; immaculate. 

Underside fore wing : scarcely differs from cercene Hewitson, except for the slightly different 
distal margin of the white band. Submarginal and marginal lines as in male. 

Underside hind wing : grey-brown, with markings as in the male. 

Length of fore wing ; 21 mm. 

Neallotype $. Uganda: Unyoro, Budongo Forest, viii.1957. T. H. E. 
Jackson. 

Recorded from : Fr. Equat. Africa : Etoumbi and Ouesso ; Uganda : Bwamba, 
Budongo and Mpigi. 

Epitola cercenoides Holland 

(J Epitola cercenoides Holland, 1890 : 424. 

Type. Gaboon : " Upper waters of the River Ogove." (Carnegie Mus., Pittsburg.) 

9 Epitola batesi Druce, 1910 : 365, pi. 34, figs. 2, 3. 

Type. Cameroons : Bitje. 

(J, $ Epitola batesi Druce ; Seitz, 1920 : pi. 64, h. 

Recorded from : West Africa to Uganda. 

Epitola uniformis Kirby 
(PI. I, fig. 8, PI. 2, fig. 17) 

{J Epitola uniformis Kirby, 1887 : 445. 

Type. Nigeria : Lagos. 

Epitola uniformis Kirby ; Seitz, 1920 : pi. 64, h. 

? Epitola versicolor Kirby, 1887 : 444. (Described as the male.) 

Epitola leonina Staudinger ; Seitz, 1920 : 355. 

$. There has been considerable confusion in the past over the females of uni- 
formis Kirby, leonina Staudinger and zelza Hewitson. The female described by 
Kirby, loc. cit. and figured in Seitz, 64, h, is the female of zelza Hewitson. Similarly, 
the female ascribed by Aurivillius in Seitz, p. 355, to leonina Staudinger is the 
female of uniformis Kirby. The following evidence is considered conclusive. 

1. An examination of the underside of leonina Staudinger shows that it belongs to 
the group containing zelza Hewitson, tumentia H. Druce, hadura Kirby and nigro- 
venata sp. n. All the females of this group are brown with blue discal areas. It 
seems probable, therefore, that the female of leonina will be similar. Further, in 
the eastern Congo and extreme west of Uganda (Bwamba) leonina flies with a blue 
female and in the west a similar female is found in the same areas as the male. 

2. If the above is correct then the black and white female must belong to uniformis 
and it is significant that this female occurs in all localities cited for the male. The 
only alternative to this is that this black and white female is merely a variety of 
cercenoides HoUand with blue scaling covering the disc of the hind wing, an unlikely 
hypothesis, since it would leave uniformis, a comparatively common species, without 
a known female of any sort. On average; also, the females with the blue discal area 
in hind wing, are smaller than those of cercenoides. 



NOTES ON THE EPITOLINAE 133 

This insect was described by Aurivillius in Seitz, under the name leonina Staudinger, 
but a fuller description is given below : 

9. Nearest to cercenoides Holland, but with blue discal area in hind wing. 

Legs dark brown with paler joints ; palps dark brown above and below, also eyes and frons ; 
antennal shaft black above, checkered below ; club orange-tipped. 

Upperside fore wing : black-brown ; cell and extreme base of i blue ; a large black triangular 
spot on the discocellular ; broad white postdiscal band not invading the cell and with evenly 
curved distal edge, becoming yellowish towards the costa and stretching from veins la to 9. 
Slight indentations in 2 and 3 but less so than in cercenoides. 

Upperside hind wing : dark brown ; a blue patch covering the basal two-thirds of the cell, 
the extreme base of space 3 and the basal third of i . This blue area is variable ; it may extend 
over the whole cell and beyond or be reduced to some scattered blue scales, but is present in all 
specimens before me. 

Underside fore wing : grey-brown ; the postdiscal band strongly yellow from vein i to the 
costa ; apical markings prominent as in male. Basal grey-brown area distally V-shaped in which 
it differs from cercenoides. 

Underside hind wing : grey-brown with prominent lines as in male. 

Length of fore wing : 16 mm. 

Neallotype $. Fr. Equat. Africa : Ouesso, Ketta Forest, ix.1959. T. H. E. 
Jackson. 

Recorded from : Sierra Leone ; Ivory Coast ; Nigeria ; Ghana ; Gaboon ; 
Fr. Equat. Africa ; Congo ; N. Angola. 

Epitola dorothea Baker 
(PL 9, fig. 87, PI. 10. fig. 97) 

(J, $ Epitola dorothea Baker, 1904 : 227. 
Type. Sierra Leone. 

Recorded from : Sierra Leone : Moyamba, Mabangi ; Ivory Coast : Guiglo ; 
Ghana : Kumasi. 

Epitola insulana Aurivillius 
(PI. I, fig. 9, PI. 2, fig. 18. PI. 9, fig. 88, PI. 10, fig. 98) 

(J Epitola insulana Aurivillius, 1923 : 1203. 

Type. Fernado Po : San Carlos. (Riksmuseum, Stockholm.) 

$. Legs and palps below pale brown, heavily dusted with grey scales ; palps above darker 
brown ; frons light brown laterally fringed with white ; eyes dark brown ; antennal shaft dark 
brown above, below minutely checkered white ; club orange-tipped. 

Upperside : nearest to cercene Hewitson and cercenoides Holland but differs from both in the 
bright blue basal area fore wing. 

Upperside fore wing : black ; broad white postdiscal band from 2-7 broadest in space 2, 8 mm., 
where it almost reaches the margin, slightly invading space i at the distal end and narrowest, 
2 mm., in space 6, proximally this band is straight from vein 7 to the origin of vein 3 and then 
bent distad, running almost along vein 2 ; whole basal area bright blue ; extending in space i 
to within 2 mm. of the margin, small triangular black spot, at junction between white and blue 
areas arising from vein 7. 

Upperside hind wing : plain dark brown. 

Underside fore wing : pale brown with white band of upperside showing through, but exten- 
ding to space I, proximal edge, therefore, straight throughout ; a few faint paler markings in 
apex. 



134 T. H. E. JACKSON 

Underside hind wing : pale brown, with very faint traces of submarginal and postdiscal lines. 
Length of fore wing : 17 mm. 

Neallotype 9- Fernando Po : 3,000-4,000 ft., vi. 1926. T. A. Barns. Joicey 
Bequest. 

Known only from Fernando Po. 

An outlying member of this group is Epitola katherinae Poulton, in which the 
female is similar, but the light patch in fore wing is orange. 

Epitola katherinae Poulton 

cj, $ Epitola katherinae Poulton, 1929 : 494, pi. 30, figs. 5, 7. 
Type. Nigeria : Ibadan. (Hope Dept., Oxford.) 

Recorded from : Nigeria : Oshodi and Ibadan. 
The following species probably belongs here : 

Epitola carilla Roche 

cJ Epitola carilla Roche, 1954 • 495. P^- 20, figs. 13, 14. 
Tj^e. Uganda : Entebbe. 

9 unknown. 

Known only from the type. 

B. 9$ brown with large blue discal areas in both wings, but no postdiscal white 
line or blue spots ; occasionally the blue area in fore wing is distally white. 

Epitola staudingeri staudingeri Kirby 
(PI. 3, fig. 19, PI. 4, fig. 29) 

cJ Epitola staudingeri Kirby, 1890 : 271. 
Type. Sierra Leone. (Staudinger Coll.) 
Epitola staudingeri Kirby ; Seitz, 1920 : pi. 65, d. 

$. Legs dark brown, with much lighter scaling at the joints ; palps light brown with whitish 
lateral edges ; antennal shaft, black above, checkered white below ; club tipped bright orange. 

Upperside fore wing : dark brown, with dull blue basal area from inner margin to upper median 
vein covering the cell, but not extending beyond it ; distally very irregular and arranged as 
follows : a long sharply pointed streak in 5, extending half-way along the cellule ; a very small 
spot in the base of 4 ; a streak in 3, 5 mm.; the whole of i and 2 almost to the margin. 

Upperside hind wing : dark brown ; discal and basal dull blue patch from veins 1-6, not 
sharply defined distally ; a broad dark brown border 2-3 mm. but containing scattered blue 
scales ; costal border dark brown. 

Underside : clay coloured, slightly paler along inner margin of fore wing ; faint traces of two 
submarginal crenulate lines on both wings, otherwise unmarked. 

Length of fore wing : 20 mm. 

Neallotype $. Sierra Leone : Baudajuma, No. 1898. G. I. Arnold. 
Recorded from : Sierra Leone and Nigeria. 



NOTES ON THE EPITOLINAE 135 

Epitola staudingeri gordoni Druce 

^ Epitola gordoni Druce, 1903 : 70. 

Type. Nigeria : Bonny. (Hope Dept., Oxford.) 

$ unknown. 

The type has been examined in Hope Department, Oxford. The underside 
markings, mentioned by Druce are present to a greater or lesser degree in the series 
in British Museum (Nat. Hist.) and gordoni should be considered a synonym of 
staudingeri Kirby. 

Epitola staudingeri aequatorialis ssp. n. 

(PI. 13, figs. 120, 121, 126, 127) 

This subspecies occurs in French Equatorial Africa, i.e. in the zone usually referred 
to as Gaboon. It differs from the typical race in the female, which is brown, un- 
marked, thus linking it with concepdon Suffert and elissa Grose-Smith. 

(J. Does not differ either above or below from the typical race. 
$. Upper side : dark brown, unmarked. 

Underside : pale warm brown ; traces of paler postdiscal spots in fore wing and of submarginal 
and postdiscal crenulate lines in hind wing. 
Length of fore wing : 18 mm. 

Holotype $. Fr. Equat. Africa : Moy en Congo, Etoumbi. ix.1960. T. H. E. 
Jackson. 
Allotype c^. Same data as holotype. 
Recorded from : Gaboon and Fr. Equat. Africa. 

Epitola mora Talbot (stat. n.) 
(PI. 3, fig. 20, PI. 4, fig. 30) 

(J Epitola staudingeri mar a Talbot, 1935 : 75. 
Type. Vict. Nyanza : Nr. Kakindu. (Hope Dept., Oxford.) 
^ Epitola jacksoni Roche, 1954 • 49^> P^- 20, figs. 15, 16 (syn. n.). 
Type. Uganda : Katera. 

The type of staudingeri mar a has been examined in Hope Department, Oxford 
and found to be identical with the species described by Roche (1954 : 496) under 
the name of Epitola jacksoni which name sinks as a synonym. The present author 
however, agrees with Roche that mara Talbot is not a ssp. of staudingeri Kirby. 

$. Frons, legs, palps light brown ; eyes black ; antennal shaft brown ; club black above 
with pale brown tip, brown below. 

Upperside fore wing : dark brown ; scattered blue scales throughout cell, extended in a streak 
along vein 5 for about half its length ; half of cellule 2 and the whole of i, up to 2 mm. of the 
margin. 

Upperside hind wing : dark brown with basal blue patch extending from vein i to vein 6, 
covering the whole of the cell, and distad to within 5 mm. of the margin. The distal margin of 
the blue patch, however, fades out into the ground colour, without any clearly defined boundary. 

Underside : pale clay coloured on both wings with traces of the postdiscal silvery spots, 
especially the large one on the inner margin ; no other markings. 

Length of fore wing : 20 mm. 



136 T. H. E. JACKSON 

Neallotype $. Uganda: Masaka, Sango Bay, Katera. xi.1955. T. H. E. 
Jackson. 

This species flies high in and under the canopy and can only be taken by building 
a platform in a suitable place ; the males fly later than most Epitola and are still 
moving up to i p.m. The females come down to lay on dead twigs, etc. 

Recorded only from : Uganda : Masaka Dist., Sango Bay, Katera and Kakindu. 

Epitola leonina Staudinger 
(PI. 3, fig. 21, PI. 4, fig. 31) 

(J Epitola leonina Staudinger, 1888 : 268. 

Type. Sierra Leone. (Staudinger Coll.) 

Epitola leonina Staudinger ; Seitz, 1920 : pi. 65, e. 

Epitola ciconia Grose-Smith & Kirby, 1892 : Afr. Lye. pi. 17, figs., 5, 6. (See Aurivillius, 1898). 

The female described in Seitz (1920 : 355) as E. leonina is the female of uniformis 
Kirby (see explanation under that species) ; the correct female is described below : 

$. Legs dark brown, lighter at the joints ; eyes black ; palps dark brown ; frons grey-brown ; 
antennal shaft brown above, checkered white below ; club black with light brown tip. 

Upperside fore wing : black-brown ; a basal blue patch extending from the inner margin to 
vein 5 and covering the cell except for a streak in the upper angle. The distal edge is broadly 
dentate and includes almost the whole of space i, half 2 and 3 mm. of 3 and 4 ; a fine black line 
along the discocellular as in the male ; veins black. 

Upperside hind wing : black -brown with blue patch from veins 1-6, extending within 3-4 mm. 
of the margin ; veins black. 

Underside : grey-brown ; markings as in the male but more prominent, especially the discal 
and postdiscal lines. 

Length of fore wing 19 mm. 

Neallotype ?. Nigeria: Calabar Prov., Okarara, iii.igsS. T. H. E. Jackson. 

Recorded from : Sierra Leone ; Nigeria : Okarara and Ikom ; Br. Cameroons : 

Mamfe; Fr. Cameroons : Bitje; Fr. Equat. Africa : Etoumbi; Uganda: Bwamba. 

Epitola badura Kirby 

(J Epitola badura Kirby, 1890 : 271. 

Type. Cameroons. (Missing.) 

Epitola badwia [sic] Kirby ; Seitz, 1920 : pi. 64, g. 

$ Epitola badura Kirby ; Aurivillius in Seitz, 1920 : 355. 

Recorded from : Cameroons : Barombi, Bitje, Johann Albrechts Hohe ; Fr. 
Equat. Africa : Etoumbi ; Congo : Irumu, Beni ; Uganda : Bwamba. 

Epitola nigrovenata sp. n. 

(PI. 3, figs. 22, 23, PI. 4, figs. 32, 33) 
This species belongs to the subgroup of leonina Staudinger and badura Kirby. 

(J, ?. Legs dark brown, spotted with whitish yellow at the joints ; palps, eyes and frons 
dark brown. Antennal shaft black above, minutely checkered white below ; club orange-tipped. 

(J. Upperside fore wing : black with blue basal and discal area from inner margin to and 
including the cell ; distal edge of this sharply "and deeply indented and broken into spots by 
black along the veins ; veins 2, 3 and 4, the lower median and discocellular all broadly black ; 
along the inner margin in spaces i and 2 the blue colour reaches within ^ mm. of the margin. 



NOTES ON THE EPITOLINAE 137 

Upperside hind wing : blue from veins 1-5 from base to margin, only the fringes black ; beyond 
this black to costa ; vein 4 and the median blackened. 

Underside fore wing : dark brownish grey ; obscure silvery submarginal lines becoming 
prominent at inner angle ; fine, silvery subapical and postdiscal lines enclosing a darker band 
of the ground colour ; a few silvery spots in cell. 

Underside hind wing : dark brownish grey with traces of five silvery lines. This underside 
pattern is typical of all members of the subgroup. 

$ Upperside fore wing : black with blue area as in male, differing in extent only in the cell, 
where the basal two-thirds is black ; veins blackened as in male, but finer. 

Upperside hind wing : black ; blue basal area restricted, leaving a broad black border i mm. 
wide at anal angle and 4 mm. on vein 5 ; distal edge not sharply delineated ; vein 4 and the 
median blackened. 

Underside : as in the male but all lines and markings more prominent. 

Length of fore wing : ,^ 23 mm., $ 24 mm. 

Holotype cJ. S. Nigeria : Oban, ii.1921. Cator Coll. 

Allotype $. Fr. Equat. Africa: Moyen Congo, Etoumbi, iii.1959. T. H. E. 
Jackson. 

Recorded only from the two localities mentioned above. 

Epitola tutnentia Druce 
(PI. 3, fig. 24, PI. 4, fig. 34) 

^ Epitola tumentia Druce, 1910 : 366, pi. 33, fig. 15. 

Type. Nigeria : Afikpo. 

Epitola tumentia Druce ; Seitz, 1920 : pi. 64, g. 

$. Very similar to the female of leonina Staudinger. Legs dark brown with paler joints ; 
palps dark brown ; eyes black ; frons grey-brown ; antennal shaft black above, finely checkered 
black and white below ; club black with orange tip. 

Upperside fore wing : black-brown with blue basal patch extending almost as in leonina, but 
slightly restricted in space 4 ; veins not or little blackened ; black discocellular streak finer. 

Upperside hind wing : black-brown with blue basal patch from veins 1-6 ; distal edge ill- 
defined ; margin i mm. at vein i and 3 mm. at vein 6 ; veins not blackened. 

Underside : as in the male ; differs from leonina Staudinger in the greyer, less brown, ground 
colour and in the thick silvery conjoined double submarginal lines in both wings. Length of 
fore wing : 18 mm. 

Neallotype $. Fr. Equat. Africa: Moyen Congo, Etoumbi. xii.1958. 
T. H. E. Jackson. 

Recorded from : Nigeria : Mamu, Awka Dist., and Uwet ; Fr. Equat. Africa : 
Ouesso and Etoumbi. 

Epitola zelza Hewitson 
(PI. 9. fig. 84, PI. 10, fig. 94) 

(^ Epitola zelza Hewitson, 1873 : 151. 

Type. Nigeria : Old Calabar. 

Epitola zelza Hewitson ; Seitz, 1920 : pi. 65, e. 

Epitola badia Kirby, 1887 : 444. 

Type. Cameroons. 

$ Epitola uniformis Kirby ; Seitz, 1920 : pi. 64, h. 

$ figured in Seitz, pl. 64, h, as uniformis Kirby and described under that species. 
A redescription is given below of the male ; cf. Aurivillius in Seitz, who states that 



138 T. H. E. JACKSON 

this species was described from a damaged specimen and borne out by the figure, 
which is totally incorrect. This figure is but a copy of Hewitson's figure in ///. 
Diurn. Lep. Suppl. pi. lb, figs. 15, 16. 

(J. Legs brown with paler joints ; palps mouse-brown ; eyes black ; frons black-brown ; 
antennal shaft black above, finely checkered white below ; club black with pale brown tip. 

Upperside fore wing : black with bright blue basal patch from inner margin to vein 7 ; distal 
edge evenly curved but dentate, reaches almost to margin in i ; costa broadly black ; median 
vein swollen at base. 

Upperside hind wing : blue from vein i nearly to 7, with narrow black distal margin. 

Underside fore wing : mouse-grey with faint but distinct dentate lines, two submarginal, one 
postdiscal, one shorter, across end of cell ; some faint spots in cell. 

Underside hind wing : mouse-grey with five pale dentate lines ; two submarginal, one post- 
discal, one across end of cell, one very faint across cell. 

Length of fore wing : 15 mm. 

Recorded from : Nigeria : Mamu, Awka, Udi, Eket, Oban and Uwet ; Fr. 
Equat. Africa : Etoumbi and Ouesso ; Coi;go : Lowa Valley. 

Epitola gerina Hewitson 

^ Epitola gerina Hewitson, 1878 : 19, pi. 16, figs. 13, 14. 

T3rpe. Congo. 

Epitola gerina Hewitson ; Seitz, 1920 : pi. 65, b. 

$ Epitola goodii Holland, 1890 : 24. 

Type. Gaboon : " Upper waters of the River Ogove." (Carnegie Mus., Pittsburg.) 

$ Epitola zelica Kirby, 1890 : 272. 

Type. Cameroons : Barombi. 

Epitola daveyi Roche 

<J, $ Epitola daveyi Roche, 1954 • 499- P^- 20, figs. 19, 29 (J ; figs. 21, 22 $. 
Type. SE. Nigeria : Isoba. 

Known only from the types. 

Epitola albomaculata Baker 

(J Epitola albomaculata Baker, 1903 : 329. 
Type. Sierra Leone. 

Epitola albomaculata Baker ; Seitz, 1920 : pi. 65, e. 
$ Epitola albomaculata Baker, 1904 : 230. 

Recorded from : Sierra Leone : Moyamba ; Nigeria : Warn. 

Epitola liana Roche 

(PI. 13, figs. 119, 125) 

(J Epitola liana Roche, 1954 ■ 5°°' pl- 21, figs. 23, 24. 
Type. Uganda : Bwamba. 

$. Legs white ; palps white with black tips, frons white ; eyes brown ; antennal shaft 
black above ; finely checkered white below. 

Upperside fore wing : black with dark blue basal area from the upper median to inner margin 
and extending distad as follows : upper half of cell and space 4 white to within 3 mm. of margin ; 
base of spaces 2 and 3 blue and a large white spot mid -way in 2 ; space i blue, distally white to 
within 2 mm. of margin. A small black streak on the discocellular. 



NOTES ON THE EPITOLINAE 139 

Upper side hind wing : dark grey ; blue basal patch from veins 1-6, covering half space 2 and 
the bases of 3-6 ; a large whitish spot mid-way in 5 and 6 and space 8 whitish ; a fine black 
streak on the discocellular ; fine black marginal line and fringes checkered. 

Underside : as in the male, but white ground colour more prominent. 

Length of fore wing : 15 mm. 

Neallotype $. Uganda: Masaka, Sango Bay, Katera. viii.1960. T. H. E. 
Jackson, 

Recorded from : Uganda : Budongo, Katera and Bwamba. 

Epitola nitide Druce 

(J Epitola nitide Druce, 1910 : 336, pi. 34, fig. i. 

Type. Cameroons : Bitje, Ja River. 

Epitola nitida [sic] Druce ; Seitz, 1920 : 356. 

$ Epitola subalba Baker, 1915 : 190. 

Type. Cameroons : Bitje. 

Epitola nitide Druce ; Dufrane, 1953 : 50 [Redescription] . 

Recorded from : Ghana : Ho, Kapandu ; Cameroons : Bitje ; Fr. Equat. 
Africa : Kelle ; Uganda : Mukono, Katera. 

Epitola virginea Baker 

$ Epitola virginea Baker, 1904 : 230. 

Type. Sierra Leone. 

^ Epitola virginea Baker ; Roche, 1954 • 5°!. pl- 20, figs. 17, 18. 

Allotype c?- Sierra Leone : Daru. 

Recorded from : Sierra Leone : Moyamba, Daru ; Ghana : Kumasi. 
G. ?$. Plain brown above. 

Epitola concepcion Suffert 

$ Epitola concepcion Suffert, 1904 : 54. 
Type. Cameroons : Barombi (Berlin Mus.). 
Epitola concepcion Suffert ; Seitz, 1920 : pi. 65, a. 
^ Epitola concepcion Suffert ; Aurivillius, 1923 : 1199. 
Allotype, S. Cameroons : Molundu. 

There are no specimens of this species in the British Museum (Nat. Hist.) or in 
the author's collection. 

Epitola elissa Grose-Smith 
(PI. 3. fig. 25; PI. 4. fig. 35) 

cj Epitola elissa Grose-Smith, 1898 : 354. 

Type. Nigeria : Warri. 

Epitola oniensis Baker, 191 3 : 501. 

Type. Nigeria : Oni near Lagos. (Hope Dept., Oxford.) 

9. Legs brown with paler joints, almost checkered ; palps pale brown with black tips ; 
eyes black ; frons dark brown ; antennal shaft dark brown above, checkered below ; club 
yellow-tipped. 

Upperside : plain dark brown in both wings ; no markings. 

ENTOM. 12, 3. 9 



I40 T. H. E. JACKSON 

Underside fore wing : warm brown ; two faint dentate submarginal lines ; a stronger post- 
discal crescentric line from costa to vein 2 ; no other marks ; space i pale grey. 

Underside hind wing : warm brown ; traces of four-dentate lines all distad of the cell and 
broken into spots. Both wings with fine dark margins. 

Length of fore wing : 15 mm. 

Neallotype $. Nigeria : Ogoja Prov., Ikom. ii.1956. T. H. E. Jackson. 
Recorded from : Sierra Leone : Moyamba ; Nigeria : Ikom and Warri ; 
Br. Cameroons : Mamfe. 

Epitola ikoya Roche 

cj, $ Epitola ikoya Roche, 1954 • 497- P^- ^9. figs. 5-6 (J ; 7, 8 $. 
Type. Nigeria : Lagos. 

Recorded from : Nigeria : Oshodi, Mamu Forest, Ubiaja. 

D. $$. Brown with blue discal areas in both wings and a series of white postdiscal 
spots, fore wing. 

Epitola congoana AuriviUius 

(J, 9 Epitola congoana AuriviUius, 1923 : 1201. 

Type. Congo : Kimuenza. (Riksmuseum Stockholm.) 

Recorded from : Nigeria : Oshodi and Ubiaja ; Congo : Kimuenza, 



Epitola flavoantennata Roche 

(J, $ Epitola flavoantennata Roche, 1954 • 495- P^- ^9' figs, i, 2 (J ; 3, 4 ?. 
Type. Cameroons : Bitje, Ja River. Taken in cop. 

Recorded from : Cameroons : Bitje ; Fr, Equat. Africa : Ketta and Mambili ; 
Congo : Beni and LubiUnga Valley, Lubutu. 

Epitola carcina Hewitson 

$ Epitola carcina Hewitson, 1873 : 150. 

Type. Nigeria : Old Calabar. 

(J, $ Epitola kholifa Baker, 1904 : 229. 

Type. Sierra Leone. 

(J, $ Epitola carcina Hewitson ; AuriviUius in Seitz, 1920 : 353, pi, 65, d. 

? Epitola entebbeana Baker, 1926 : 392. 

Allotype. Nigeria : Oban. 

Epitola coerulea sp. n. 

(PI. 3, figs. 26, 27, PI. 4, figs. 36, 37) 
Nearest to carcina Hewitson. 

(J. Legs very dark brown with small pale bands at the joints ; palps dark grey -brown above 
and below ; frons dark brown ; eyes rather browner than in other species ; antennal shaft 
black above, checkered black and white below ; club with orange tip. 

Upper side fore wing : black with shining azure blue basal patch, much brighter and more 
extended than in carcina ; blue colour extends from inner margin to vein 7, filling half spaces 
5 and 6, more than half 3 and 4, and almost the whole of i and la, distally evenly curved, 



NOTES ON THE EPITOLINAE 141 

reaching almost to the margin in i. A basal scent patch in the base of space i, overlapping a 
little into the cell ; in carcina this patch extends also into la and the blue colour is restricted in 
4 and 5, and is often absent in 6. Vein i, the median and subcostal swollen at base. 

Upper side hind wing : much as in carcina ; blue from veins 1-6 with fine black margin ; 
costal margin broadly black. 

Underside : differs from carcina in the paler ground colour, less contrasting pattern and in the 
much smaller silvery area along the inner margin, which in carcina reaches vein 3, and in coerulea 
scarcely to vein 2. 

Underside fore wing : dark grey with pale silvery markings as follows : a crescentric submargi- 
nal line, followed by a broad second line almost a band ; two very faintly defined postdiscal 
lines about 3 mm. apart on vein 4, enclosing a darker patch of the ground colour ; silvery at 
base of costa ; an inner marginal silvery patch in the distal half of space i and la reaching to vein 
2. 

Underside hind wing : dark grey with pale silvery markings ; a submarginal crescentric line, 
followed by a broad area of silvery scales ; two well-defined postdiscal lines divided into spots 
and enclosing a darker band 2 mm. wide ; some scattered silvery spots in cell and basal area. 

Length of fore wing : 20 mm.; on the whole slightly larger than carcina. 

$ differs from carcina Hewitson in the larger size, and paler brighter blue areas. 

Antennae, etc. as in male. 

Upperside fore wing : black ; bright light blue basal patch from vein i to cell and distad only 
to lower angle of space 4, half 2 and 3, and most of i and la. Distal edge dentate, slightly 
extended to i, to within 2 mm. of margin. A black streak on the discocellular more prominent 
than in carcina ; four large white postdiscal spots in 4-7 in an even curve from half-way along 
costa. 

Upperside hind wing : black with restricted light blue patch from veins 1-6 ; black costal 
margin and black border 4 mm. wide throughout ; distal margin of blue area quite even, but not 
sharply defined. 

Underside : greyish brown ; very similar to carcina Hewitson. 

Underside fore wing : two pale submarginal crescentric lines ; a postdiscal series of large 
whitish spots on the veins from 2-7 and a streak in 8. No basal spots. 

Underside hind wing : two crescentric submarginal lines with much silvery, scaling between 
them ; a postdiscal series of small whitish spots ; an obscure double irregular line across end of 
cell, and a few scattered basal spots. Margins fine ; dark brown. 

Length of fore wing : 23 mm. (18 mm. in carcina). 

Holotype ,^. Fr. Equat. Africa : Moyen Congo, Etoumbi. iii.1959. T. H E. 
Jackson. 

Allotype $. Fr. Equat. Africa : Ouesso, Mambili Forest, viii.1959. T. H. E. 
Jackson. 

Flies with carcina in the same area in Fr. Equat. Africa. 

Recorded from : Fr. Equat. Africa : Ouesso and Etoumbi. 

Epitola dunia Kirby 
(PI. 3, fig. 28, PI. 4, fig. 38) 

<J Epitola dunia Kirby, 1887 : 441. 

Type. Cameroons. 

Epitola dunia Kirby ; Grose-Smith & Kirby, i.1889 : Lye. Afr. pi. 7, figs, i, 2. 

This species was not mentioned by Aurivillius in Seitz and it appears that he 
considered it a synonym of carcina Hewitson. 

$. Similar to the females of carcina Hewitson, coerulea sp. n. and flavoantennata Roche, but 
darker with restricted blue areas and a thick black streak on the hind wing discocellular. 



142 T. H. E. JACKSON 

Legs dark brown with some small light spots at the joints ; palps dark grey-brown above and 
below ; eyes brown ; frons dark brown ; antennal shaft black above and minutely checkered 
white below ; club yellow tipped. 

Upperside fore wing : black with blue inner marginal patch covering a little over half la and 

1 and 2 to within 2 mm. of the margin ; occasionally two blue spots in cell divided by black 
ground colour. A series of four postdiscal white spots in 4-7 in an even curve to the middle of 
the costa. 

Upperside hind wing : black with blue area covering the cell, base of 1-3, and half 3 and 4, 
distally variable in extent. A thick black streak on the discocellular occasionally reduced to a 
fine line. Broad dark border ; blue more restricted than in allied species. 

Underside : light brown as in carcina and coerulea. 

Underside fore wing : two faint pale crescentric submarginal lines ; series of prominent post- 
discal white spots from 2-7, but sometimes missing in 3 ; a silvery white patch in i and la about 

2 mm. from margin to half the cellules ; no basal spots. 

Underside hind wing : two crescentric submarginal lines and a postdiscal series of small whitish 
spots ; markings faint or sometimes well defined. 
Length of fore wing : 17 mm. 

Neallotype ?. Nigeria: Ogoja Prov., Ikom. ii.1956. T. H. E. Jackson. 
Recorded from : Sierra Leone : Mano ; Nigeria : Ikom, Benin, Oban, Warri 
and Uwet ; Br. Cameroons : Mamfe, Barombi. 

The following species probably belongs here : 

Epitola iturina Joicey & Talbot 

(PI. 9, fig. 86 ; PI. 10, fig. 96) 

<J Epitola iturina Joicey & Talbot, 1921 : 85, pi. 14, fig. 62. 

Type. Congo : forest between Ituri and Lindi Rivers, SW. of Avakubi. 

Epitola hella Aurivillius, 1923 : 1200, taf. i, lig. 2. 

Type. Cameroons : N'gumesok. (Riksmuseum, Stockholm.) 

$ unknown. 

Recorded from : Cameroons : Bitje, N'gumesok ; Fr. Equat. Africa : 
Etoumbi and Mambili ; Congo : Lusambo, Lowowo Valley, Ituri River. 

THE PINODES GROUP 

A. $$. Plain brown above, sometimes with small obscure pale postdiscal spots 
in fore wing. 

Epitola viridana viridana Joicey & Talbot 
(PI. 5, fig. 39; PI. 6, fig. 49) 

(J Epitola viridana Joicey & Talbot, 192 1 : 84, pi. 14, figs. 58, 59. 
Type. Congo : South side of Ituri River, 5 days west of Irumu. 

$. Legs brown and white checkered ; palps whitish below with black tips, dark brown above ; 
eyes dark brown ; frons dark brown ; antennal shaft black above, checkered below ; club 
minutely yellow tipped, below only. 

Upperside : plain brown ; traces of 3 or 4 very faint postdiscal spots in 4, 5 and 6 fore wing. 

Underside fore wing : as in the male ; brown with broad black inner marginal streak from 
2-5 ; prominent pale postdiscal spots in 4-7, and a pale spot in 2 in the black area ; two pale 
crescentric submarginal lines. Space i pale, whitish. 



NOTES ON THE EPITOLINAE 143 

Underside hind wing : brown ; a series of three obscure parallel crescentric lines in the marginal 
area. 

Length of fore wing : 15 mm. 

Neallotype $. Uganda: Bwamba. ix.1941. T. H. E, Jackson. 

Taken in cop. 

The males of this species fly in clearings or over clumps of bush at the edge of the 
forest from about 9-9.30 a.m. The females fly later laying on dead twigs. 

Recorded from : Fr. Equat. Africa : Etoumbi, Ouesso ; Congo : Epulu, 
Upper Lowa Valley, Yanga ; Uganda : Bwamba, Kigezi, Katera, Budongo, Jinja, 
W. Elgon, etc. 

Epitola viridana radiata Baker 

(J, $ Epitola radiata Baker, 1926 : 393. 
Type. Cameroons : Bitje, Ja River. 

Recorded from : Cameroons : Bitje, Johann Albrechts Hohe. 

Epitola pinodes Druce 

^ Epitola pinodes Druce, 1890 : 24. 

Type. Nigeria : Lagos. 

$ Epitola pinodes Druce ; Seitz, 1920 : pi. 65, a. 

Epitola pinodes Druce ; Grose-Smith & Kirby, iv.1891 : Afr. Lye. pi. 13, figs, i, 2. 

Type missing. 

Recorded from : Sierra Leone : Moyamba, Batkanu ; Ivory Coast ; Nigeria : 
Eket. 

Epitola orientalis Roche 
(PI. 5, fig. 42; P1.6. fig. 52) 

Epitola orientalis Roche, 1954 • 499> P^- 22, figs. 35, 36. 
Type. Uganda : Bwamba. 

Very similar to viridana Joicey & Talbot, but differs below. 

$. Legs brown and white checkered ; palps whitish, streaked black below with black tips, 
above basally whitish with long black tips ; frons dark brown ; antennal shaft black above, 
checkered white and black below ; club orange-tipped. 

Upperside : plain dark brown with four bluish postdiscal spots from 4-7 on fore wing. 

Underside : as in the male, cf. figure in Roche loc. cit.; the dark postdiscal band in hind wing 
below is plainly visible. 

Length of fore wing : 16 mm. 

Neallotype $. Kenya: Kakamega. ix.1937. T. H. E. Jackson. 
Males of this species were taken at Lake Nabugabo flying over clumps of bushes 
early in the morning. 

Recorded from : Uganda : Bwamba and Lake Nabugabo ; Kenya : Kakamega. 

Epitola catuna Kirby 

(PI. 5, fig. 40; PI. 6, fig. 50) 

(^ Epitola catuna Kirby, 1890 : 273. 

Type. Cameroons. (Staudinger Coll.) 

Epitola catuna Kirby ; Grose-Smith & Kirby, iv.1892 : Lye. Afr. pi. 17, figs. 11, 12. 



144 T. H. E. JACKSON 

Epitola catuna Kirby ; Seitz, 1920 : pi. 64, g. 

$ Epitola mus Suffert, 1904 : 53 (syn. n.). 

Type. Cameroons : Barombi Station. (Berlin Mus.) 

Epitola mus Suffert ; Seitz, 1920 : pi. 65, e. 

The type of Epitola mus Suffert has been examined and proves to be a female of 
Epitola catuna Kirb\^ 

$ Legs brown with small pale spots on the joints ; palps dark brown above and below ; frons 
and eyes brown ; antennal shaft black above, checkered below ; club orange-tipped. 

Upperside : plain dark brown ; immaculate. 

Underside fore wing : brownish grey as in the male ; inner margin broadly pale, then base to 
vein 5 very slightly darker than ground colour, a few obscure marks in apical area. 

Underside hind wing : brownish grey ; three very faint crescentric lines in submarginal and 
postdiscal areas. 

Length of fore wing : 15 mm. 

Neallotype $. Uganda: Bwamba, Bundibugyo. v. 1954. T. H. E. Jackson. 

Recorded from : Fernando Po ; Spanish Guinea : Riomuni ; Br. Cameroons : 
Barombi ; Fr. Cameroons : Bitje ; Gaboon : Kuilu ; Fr. Equat. Africa : 
Ouesso and Etoumbi ; Congo : Epulu, Beni, Ituri ; Uganda : Bugoma, Budongo, 
Bwamba, Toro, etc. 

Epitola carpenteri Baker 
(PI. 5, fig. 41, Ph 6. fig. 51. PL 9, fig. 83, PI. 10, fig. 93) 

(J Epitola carpenteri Baker, 1921 : 462. 

Type. Uganda : Bugalla Is. Lutoboka. (Hope Dept., Oxford.) 

It has apparently been overlooked by Roche et al., that Baker described the male 
of the above species in his original description, loc. cit. and designated a holotype. 
This specimen has been examined in the Hope Dept., Oxford and found to belong 
to a species near catuna Kirby. Baker's allotype female belongs to a different species, 
Epitola azurea sp. n., which is described later in this paper. 

To add to the original description of the male, it may be said that carpenteri differs from catuna 
in the more prominent blue streaks on fore wing in spaces 4 and 5, in the wider black border in 
hind wing and in the different underside. In the former, the underside ground colour is warm 
brown instead of dark grey ; the black basal patch, fore wing, is much more prominent, as are 
also the darker submarginal and discal lines in hind wing. Further, it is larger ; length of fore 
wing ; 15 mm.; in catuna 13-14 mm. 

$. Upperside : plain dark brown in both wings, with some very scattered blue scales in the 
discal and postdiscal areas of the fore wing. 

Underside : as in the male, but paler ; the pale markings in the cell of fore wing less distinct. 

Length of fore wing : 14 mm. 

Neallotype?. Uganda: Sesse Isles, Bugalla Is. ix-x.1958. T. H. E. Jackson. 

Recorded only from : Uganda : Sesse Isles, Bugalla and Entebbe. 

This species is very near catuna Kirby, but the differences are deemed sufficient 
to accord it specific rank, especially since catuna is known to occur very near 
Entebbe. 

Both sexes have been taken feeding on low shrubs among Crematogaster ants at 
the edge of the forest on Bugalla. 



NOTES ON THE EPITOLINAE 145 

Epitola ghesquieri Roche 

(PI. 5, fig. 43, PI. 6. fig. 53) 

(J Epitola ghesquieri Roche, 1954 • 49^. pl- 22. figs. 33, 34. 
Type. Congo : Equateur DLst., Eala. (M.R.C.B., Tervuren.) 

$. Legs black with small pale spots at the joints ; palps black above and below ; eyes 
brown ; frons black ; antennal shaft black above and checkered black and white below ; club 
orange tipped. 

Upperside : plain dark brpwn ; no markings. 

Underside fore wing : dove grey ; large black area between veins 2 and 5 from base to within 
I mm. of margin, broadly club-shaped. Two very faint crenulate submarginal lines. 

Underside hind wing : dove grey, markings scarcely visible but faint traces of submarginal 
lines. 

Length of fore wing : 13 mm. 

Taken in cop. 

Neallotype $. Fr. Equat. Africa: Ouesso, Ketta Forest, x.1959. T, H. E. 
Jackson. 

Recorded from : Fr. Equat. Africa : Ouesso Dist. ; Congo : Equateur Dist., 
Eala. 

Epitola nigeriae sp. n. 

(PL 5, figs. 44, 45, PI. 6. figs. 54, 55) 
Above nearest to ghesquieri Roche, but quite different below. 

^. Legs black, spotted with white at the joints ; palps black above, below spotted white and 
with white tips, covered in long hair ; eyes brown ; frons black, hairy ; antennal shaft black 
above, checkered black and white below ; club minutely yellow-tipped. 

Upperside fore wing : black ; some scattered dark blue scales in the upper basal portion of 
cellule I. Vein i for half its length, the median, subcostal and costal swollen. 

Upperside hind wing : black with very dark blue basal and discal patch from veins 1-6 ; 
inner margin black ; border black of even width throughout just over i mm. Median vein and 
extending half-way along vein 4 blackened ; veins 2 and 3 also black. 

Underside fore wing : dark grey ; traces of the crenulate submarginal lines only visible under 
a glass ; a club-shaped black patch from base to within i mm. of the distal margin, rounded 
distally between veins 2-5 ; a large square patch of the ground colour on the distal half of the 
inner margin. 

Underside hind wing : dark grey ; the usual crenulate lines scarcely visible. 

?. Upperside : plain dark brown, fringes light brown. 

Underside : exactly as in the male but ground colour ochreous brown ; the lighter patch on 
the distal margin, fore wing, confined to space la. 

Length of fore wing : (J and $ 16 mm. 

Holotype (^. Nigeria: Ogoja Prov., Ikom. x.1955. T. H. E. Jackson. 
Allotype?. Nigeria: Ogoja Prov., Ikom. i.1956. T. H. E. Jackson. 
Recorded from : Nigeria : Ikom ; Cameroons : Bitje ; Fr. Equat. Africa : 
Ouesso Dist. 

Epitola nigra Baker 

cJ, $ Epitola nigra Baker, 1903 : 331. 
Types. Sierra Leone : Moyamba. 

Recorded from : Sierra Leone : Moyamba ; Nigeria : Eket ; Cameroons : 
Bitje. 



146 T. H. E. JACKSON 

B. $$ brown with 3-4 white (or pale blue) postdiscal spots beyond cell and between 
cell and inner margin of fore wing ; hind wing broadly suffused blue to very pale 
blue, between veins i and 6 from base to margin, or occasionally with only a few 
scattered blue scales. 

Epitola cephena Hewitson 

The female described by Hewitson as cephenu is the female of doleta Kirby, and 
the former having priority the name doleta must sink as a synonym. The male 
placed by Roche (1954 : 501) to cephena is mercedes Suffert, and has a different 
female. 

Epitola cephena cephena Hewitson 

$ Epitola cephena Hewitson, 1873 : 151. 

Type. Gaboon. 

Epitola cephena Hewitson ; Hewitson, 1878 : pi. ib, figs., 9, 10. 

Epitola cephena Hewitson ; Seitz, 1920 : pi. 65, e. 

^ Epitola doleta Kirby, 1890 : 273 (syn. n.). 

Type. Sierra Leone. 

Epitola doleta Kirby ; Grose-Smith & Kirby, iv.1892 : Lye. Afr. pi. 17, figs, i, 2. 

Epitola doleta Kirby ; Seitz, 1920 : pi. 64, g. 

Phytala leonina Baker, 1903 : 328. 

Type. Sierra Leone. 

Epitola leonensis Baker, 1904 : 227. 

Type. Sierra Leone. 

Recorded from : Sierra Leone : Moyamba, Fula Wusu ; Cameroons : Bitje ; 
Nigeria : Eket, Oshodi, Ubiaja ; Ghana : Kumasi ; Fr. Equat. Africa : Ouesso ; 
Gaboon. 

Epitola cephena entebbeana Baker (stat. n.) 
(PI. 5, fig. 46; PI. 6, fig. 56) 

(J Epitola entebbeana Baker, 1926 : 392. 
Type. Uganda : Entebbe. 

$. As pointed out by Roche (1954), the female described by Baker belongs to 
carcina Hewitson and the true female is undescribed. 

$. Legs whitish, unmarked ; palps below whitish brown, tips brown, eyes black ; antennal 
shaft black above, finely checkered white below ; club minutely tipped orange ; frons grey -brown. 

Upperside fore wing : very similar to cephena cephena Hewitson, ground colour black ; three 
white blue-edged postdiscal spots in 5-7, half-way along costa ; an inner marginal bluish-white 
patch covering the basal half of la, and most of i and 2 to within 2 and i mm. of margin, 
respectively ; the spot in 2 white, except in the base. 

Upperside hind wing : brownish black ; scattered blue scales across cell and bases of spaces 

1-5- 

Underside fore wing : whitish grey with jet black patch from base to i mm. of margin between 
veins 2 and 5 ; four white postdiscal spots in 4^7, that in 4 a streak and that in 5 large and trian- 
gular ; a streak and large white spot in distal half of i in the black patch ; space la pale grey. 

Underside hind wing : pale grey unmarked, as in the typical subspecies. 

Length of fore wing : 17 mm. 



NOTES ON THE EPITOLINAE 147 

Neallotype $. Congo Belge : Mambasa, Epulu. x.ig^y. T. H. E. Jackson. 
The males behave as in other species, flying in clearings early in the morning. 
Recorded from : Congo : Beni, Epulu, Middle Lowa Valley ; Uganda : Entebbe, 
Mpigi, W. Elgon. 

Epitola mercedes Suffert 
(PI. 5, fig. 47 ; PI. 6, fig. 57) 

(^ Epitola mercedes Suffert, 1904 : 53. 

Type. Cameroons : Johann Albrechts Hohe. (Berlin Mus.) 

^ Epitola cephena Hewitson ; Roche, 1954 : 501, pi. 22, figs. 31, 32. 

Roche (1954) was well aware that there were two forms of female associated 
with cephena Hewitson, but considered them to be varieties of the same thing. This, 
however, is incorrect ; the heavily marked form as figured in Seitz, belongs to 
cephena and agrees well with the female of the eastern subspecies entehheana Baker 
and the more lightly marked insect is the female of mercedes Suffert. 

The type of Epitola mercedes Suffert has been examined, and there is no doubt 
that it is identical with the male described by Roche (1954) as cephena Hewitson. 

$. Legs brown with paler joints ; eyes black ; frons brown ; palps pale brown below, dark 
brown above ; antennal shaft black above, strongly checkered white below ; club orange- 
tipped. 

Upperside fore wing : greyish black ; blue postdiscal spots in 5 and 6 and a trace of a third in 
7 ; two others, larger, about mid -way in spaces i and 2. No other marks. 

Upperside hind wing : greyish black ; a small rounded blue spot at the end of cell, but some- 
times unmarked, which is also true of the male. 

Underside fore wing : brownish grey ; postdiscal whitish spots in i and 2, as above, and in 
4-6 ; jet black streak between veins 2-5, erupting into space 5, from base to i mm. of margin ; 
obscure crenulated submarginal lines. 

Underside hind wing : brownish grey with much grey scaling ; two obscure crenulated sub- 
marginal lines. 

Length of fore wing : 16 mm. 

Neallotype $. Nigeria: Lagos Dist., Oshodi. iv.1955. T. H. E. Jackson. 
Both sexes were seen at Oshodi flying low dowTi, in deep shape under Crematogaster- 
infested trees. 

Recorded from : Nigeria : Lagos Dist. ; Cameroons : Johann Albrechts Hohe. 

Epitola ouesso sp. n. 

(PI. 7, figs. 59, 60 ; PI. 8, figs. 69, 70) 

Nearest to cephena Hewitson in the male and to pinodes Druce and mercedes 
Suffert in the female. 

cJ. Legs dark brown with small paler joints ; palps dark brown above and below with some 
grey hairs ; frons black ; eyes brown ; antennal shaft black above, finely white checkered 
below ; club yellow-tipped. 

Upperside fore wing : black ; dark blue streak in lower half of cell and dark blue spots in 
extreme base of 2 and basal half of i . 

Upperside hind wing : black ; dark blue patch covering cell and basal half of 4 and 5 ; more 
cattered blue scales in 1-3 ; no postdiscal spots and no other markings. 



148 T. H. E. JACKSON 

Underside fore wing : dark brown, no grey coloration ; black basal streak restricted by ground 
colour in 1-3 ; no postdiscal spots or other markings. 

Underside hind wing : dark brown, usual markings scarcely visible. 

$. Very similar to female mercedes Suffert differing only below, and in the extended blue in 
hind wing above. 

Upperside forewing : brownish black ; small blue postdiscal spots in 5-7, and in base of i 
and 2. 

Upperside hind wing : obscurely scaled blue from cell to mid-way in 4 and 5 and in the base of 
I and 2, otherwise brownish black. 

Underside fore wing : browner than in mercedes ; upper edge of black inner marginal patch 
perfectly straight along vein 5 ; in mercedes there is a broad extension mid-way into space 5 ; 
large postdiscal white spots in black area in i and 2 ; very small white spots in 4-7. 

Underside hind wing : brown, faint traces of submarginal and postdiscal crenulate lines. 

Length of fore wing : <J and $ 16 mm. 

Holotype ^. Fr. Equat. Africa: Ouesso, Ketta Forest, x.1959. T. H. E. 
Jackson. 

Allotype $. Same data as holotype. 

Recorded from : Fr. Equat. Africa : Ouesso Dist. ; only the types. 



Epitola azurea sp. n. 

(PI. 7, fig. 61, PI. 8, fig. 71) 

$ Epitola carpenteri Baker, 1921 : 462, pi. 22, figs. 37, 38. 

Allotype. Uganda : Dama Is., L. Victoria. (Hope Dept., Oxford.) 

The male associated with this female by Roche belongs to mpangensis sp. n. 
See below. 

^. Legs brown with pale spots on joints ; palps pale whitish below with dark tips, black 
above ; frons whitish grey with broad brown median band ; antennal shaft black above, finely 
checkered white below ; club pale-tipped. 

Upperside fore wing : black with pale bright blue spots as follows : a spot in base of cell, 
another linear and irregular in centre, a large spot covering all of i to within 2 mm. of margin, 
another spot in base of 2 and a few blue scales in la ; a black spot of the ground colour usually, 
but not always present in upper centre of blue area in i. 

Upperside hind wing : black ; large pale bright blue discal area from veins 1-6 ; narrow black 
margin and black costal area. 

Underside fore wing : silvery grey ; black inner marginal streak bounded by vein 5 ; two very 
large silvery white spots in distal half of i and 2, almost obliterating the black area distally ; 
silvery crescentric submarginal lines ; three silvery spots in line, horizontally along upper edge 
of black area. 

Underside hind wing : silvery grey ; heavily irrorated with silver scales throughout ; sub- 
marginal and postdiscal silvery crescentric lines and a square silvery patch in the base of 4. 

Length of fore wing : 13 mm. 

This species differs from mpangensis sp. n. in the much paler brighter blue, the 
extended blue area of fore wing and in the smaller expanse. A series of males 
were taken in the type locality, with females and there is no doubt as to its identity. 

Holotype $. That described by Baker as allotype of E. carpenteri Baker. 

Allotype cJ. Uganda: Masaka, Sango Bay, Katera. v. 1956. T. H. E. Jack- 
son. 



NOTES ON THE EPITOLINAE 149 

The males fly under the canopy, never venturing outside ; flight is fast but short 
and continues later than in most species up to about 11 a.m. The females fly later, 
laying on lichen-covered tree trunks. So far known only from this locality. 

Recorded from : Uganda : Sango Bay, Katera, Dama Is., L. Victoria. 

Epitola mpangensis sp. n. 

(PI. 5. fig. 48, PI. 6. fig. 58) 
(J Epitola carpenteri Baker ; Roche, 1954 • 5°°' P^- ^-z, figs. 37, 38. 

c^. Described and figured by Roche, loc. cit., as carpenteri Baker. 

$. Similar to azurea sp. n., but with much darker underside. 

Legs dark brown with pale spots on the joints ; palps dark grey below, pale grey above with 
black tips ; frons and eyes dark brown ; antennal shaft checkered white above and below ; 
club with minute yellow tip. 

Upperside fore wing : black ; four white blue-edged postdiscal spots in 4-7 ; a large white 
spot covering most of 2 ; space i blue, distally bluish white to within 2 mm. of margin, space 
I a blue from base to middle. 

Upperside hind wing : black ; large blue discal area from veins 1-6, distally paler ; border 
of even width about 3 mm. 

Underside fore wing : brownish grey ; postdiscal spots in 4-7 and i and 2 much larger than 
in azurea, the latter filling up most of the black inner marginal streak ; latter as in azurea ; 
submarginal lines merged into wide grey border. 

Underside hind wing : brownish grey with two submarginal and one postdiscal crenulate lines 
and square grey patch in 4 as in azurea ; scattered basal grey spots. 

Length of fore wing : 15 mm. 

This species can be distinguished from azurea in the male by the darker blue and 
more restricted spots on fore wing and by the greater expanse. In the female by 
the darker underside. 

Holotype ^. That cited by Roche, loc. cit., as " Allotype " ^ Epitola carpenteri 
Baker. Uganda: Bwamba. x,i94i. T. H. E. Jackson, 

AUotj^e ?. Uganda: Mpigi, Mpanga Forest, viii.1959. T. H. E. Jackson. 

Recorded from : Congo : Lowa Valley and Katanga ; Uganda : Bwamba and 
Mpigi. 

Epitola katerae sp. n. 

(PI. 7, figs. 62, 63, PI. 8, figs. 72, 73) 

In the male nearest to ghesquieri Roche, nigeriae sp. n. and mercedes Suffert of the 
previous group and in the female to azurea sp. n. and mpangensis sp. n. 

cj. Legs brown with very small pale spots at the joints ; frons and eyes brown ; palps brown 
below with some grey hairs, brown above with black tips ; antennal shaft checkered black and 
white above and below (a very unusual character) ; club black, very minutely orange-tipped. 

Upperside fore wing : black ; immaculate. 

Upperside hind wing : black with dark blue discal basal patch between veins 1-6, covering 
only the basal portions of spaces 1-5 and ending distally in scattered blue scales ; a wide black 
distal border, 3-4 mm. and black costal border ; veins 2-4 black ; distal edge of blue patch 
ill-defined. 

Underside fore wing : dark grey ; black inner-marginal streak between veins 2-5 ; large 
pale grey square spot in distal part of i and la ; postdiscal pale spots in 4-7, conjoined to form 
a line ; another larger silvery spot in 2 ; margin speckled with silvery scales. 



I50 T. H. E. JACKSON 

Underside hind wing : dark grey ; irrorated with silvery scales, the individual lines obscured ; 
a more prominent patch of silver scales in the base of 4 and 5. 

$. Very similar to azurea. 

Legs, eyes, palps and frons much as in male, but paler ; antennal shaft checkered on both 
surfaces as in male (in aztirea jet-black above in both sexes). 

Upper side fore wing : greyish black ; large white, blue-edged postdiscal spots in 4-7, the latter 
a streak ; another, broad, elongate in 2 to i mm. of margin ; the whole of i to 2 mm. of margin 
light bluish ; basal half of la, also, bluish ; a small black streak from mid -way along median 
vein, invading blue area in i. 

Upperside hind wing : paler, blackish grey ; pale bluish basal-discal patch from veins 1-6, 
distal and costal borders of ground colour, former about 2 mm.; the blue area slightly paler 
distally. 

Underside fore wing : differs from azurea in the greater amount of dark ground colour and 
from mpangensis in the absence, as such, of the inner-marginal black streak, which in this species 
merges into the discal ground colour. Ground colour dark grey-brown ; large white patch on 
inner-margin comprising the postdiscal spots in i and 2 and covering the whole distal half of i 
and more than half 2 ; white postdiscal spots in 4-7 above ; a prominent thick white submarginal 
crenulate line ; apex and distal border thickly irrorated with silvery white scales. 

Underside hind wing : brownish grey ; the whole covered with silvery white scales with a 
more prominent silvery white patch in the base of 4 and 5 ; a dark crenulate submarginal line. 

Length of fore wing : ^ and $, 16 mm. 

Holotype ^. Uganda: Masaka, Sango Bay, Katera. viii.1956. T. H. E. 
Jackson. 

Allotype $. Same data as holotype. xi.1954. 

This species is, so far, only known from Katera, Masaka Dist., where the males 
were taken on an open hillside flying over small trees and bushes from about 9-9.30 
a.m. The females were found laying on the lichen-covered trees among Cremato- 
gaster ants. 

Epitola rileyi Audeoud 

cj, $ Epitola rileyi Audeoud ; 1936, 704, pi. 7, figs. 3 $ ; 4 c^. 
Types. Cameroons : Lolodorf. (Audeoud, Coll. Geneva.) 

This species, probably belongs here, although it differs from all others in the 
crenulated margins to the wings. 

There are no specimens of this species in the British Museum (Nat. Hist.). 



Epitola kamengensis sp. n. 

(PI. 7, figs. 64, 65 ; PI. 8, figs. 74, 75) 
Near katerae sp. n., but larger and differing below. 

^. Legs brown, white at the joints ; palps brown, scaled grey below ; brown, black-tipped 
above ; eyes and frons brown ; antennal shaft checkered black and white on both surfaces as in 
katerae ; club black, orange-tipped. Thorax clothed in green hair. 

Upperside fore wing : plain black, unmarked. 

Upperside hind wing : black with large dark, electric blue discal area from veins 1-6, reaching 
nearly to the margin ; black, even, distal border i mm. wide, and black costal border ; space 
lb black. 

Underside fore wing : dark grey with black inner-marginal streak from veins 2-5, broken at 
distal half of inner margin by a dark grey patch ; a light grey spot mid-way in i and a small one 



NOTES ON THE EPITOLINAE 151 

in base of 2 ; small pale streaks, discal, in 4-7 ; prominent white submarginal spots from 3 to 
costa and silvery scales in distal border. 

Underside hind wing : dark grey, heavily coated with silver-grey scales throughout the whole 
of the distal half, arranged roughly into two submarginal bands and a broader band horizontally 
across the wing, distal of the cell, leaving a small dark area at the base of 2 and 3 ; basal area 
with scattered silver-grey spots. 

$. Similar to azurea sp. n., katerae sp. n., etc., but larger. 

Legs pale brown, slightly paler at the joints ; palps pale brown dusted with pale grey scales ; 
frons dark brown ; eyes brown ; antennal shaft black, conspicuously checkered with white, 
above and below ; club broad and flattened, orange-tipped. 

Upper side fore wing : blackish brown ; three large white postdiscal spots in 5-7 with a streak 
in 8 ; other spots in 1-3 inclusive, that in i blue, extending from base to middle ; in 2 from base 
to within 3 mm. of margin and basally blue ; in 3 white to within 2 mm. of margin ; discal 
area and beyond somewhat darker black. 

Upper side hind wing : blackish brown with large blue discal patch extending from veins 1-6, 
and distad, dislegnic, to within 3 mm. of the margin, leaving a black-brown marginal band ; 
vein 6 to costa black-brown ; a paler patch due to admixture of white scales, in the distal half 
of blue area in spaces 5 and 6. Fringes dark brown in fore wing and from vein 3 to anal angle 
hind wing, but white from vein 3 to costa in hind wing. 

Underside : the dark basal area in hind wing, exactly as in the male, leaves no doubt as to the 
identity of this insect. 

Underside fore wing : dark sepia with white postdiscal spots as above, but, in addition, a 
small white spot in space 4, this completing the series ; apex and distal margin broadly greyish 
white, proximal edge indented. 

Underside hind wing : pale brown with four crescentric pale greyish white bands, merging into 
each other and much grey scaling ; the paler patch in the blue area above in 5 and 6 reproduced 
as a square whitish spot ; basal area including most ot cell, much darker than rest of wing and 
strongly contrasting as in the male. 

Length of fore wing : ^J 19 mm.; $ 17 mm. 

Holotype cJ. Uganda: Mawakota, Kamengo. viii.ig53. V. G. L. Van 
Someren. 

Allotype ?. Brit. E. Africa : Nandi Escarpment (Forest), 5,%o ft. 29.V.1911. 
S. A. Neave. 

This species was taken by Van Someren, flying from the forest edge out into a 
coffee plantation at about 9.30 a.m. 

Recorded from : Uganda : Kamengo and Katera ; Kenya : Nandi Escarpment. 

C. The following two species are included in the pinodes group, in spite of the 
convex margin of the fore wing, since in all other characters, including those of the 
females, they are similar 

Epitola conjuncta conjuncta Grose-Smith & Kirby 

c?, $ Epitola conjuncta Grose-Smith & Kirby, i.1893 : Lye. Afr. 86, pi. 20, figs. 3-5. 
Type. Sierra Leone. (Staudinger Coll.) 

Recorded from : Sierra Leone ; Fr. Equat. Africa : Sembe, Etoumbi. 

Epitola conjuncta budduana Talbot 

S, ? Epitola conjuncta budduana Talbot, 1937 • 62, pi. i, figs. 2 ^ ; 3 $. 
Types. Uganda : Katera. 

Recorded from : Uganda : Budongo, Bwamba, Katera, Tero Forest, Buddu. 



152 T. H. E. JACKSON 

Epitola dolorosa Roche 

cj, $ Epitola dolorosa Roche, 1954 • 49^' P^- 21, figs. 27, 28 <J, 29, 30 $. 
Types. <J, Uganda : Bwamba. Allotype ; $, Uganda : Entebbe. 

Recorded from : Uganda : Bwamba, Entebbe, Tero Forest, Buddu. 

The following species also belong to the pinodes group, but I am unable to place 
them in any of the subdivisions defined in this paper, since the females are 
unknown. 

Epitola obscura Hawker-Smith 

(J Epitola obscura Hawker-Smith, 1935 : 11. 
Type. Nigeria : Lagos. 

$ unknown. 

Recorded from : Nigeria : Lagos, Ilesha ; Ghana : Ashanti. 

Epitola maculata Hawker-Smith 

^ Epitola maculata Hawker-Smith, 1926 : 240. 

Type. Congo : Stanleyville, Elisabetha near Basoko. (M.R.C.B., Tervuren.) 

$ unknown. 

Recorded from : Congo : Stanleyville ; Uganda : Bwamba. 

Epitola pinodoides Grose-Smith & Kirby 

<J Epitola pinodoides Grose-Smith & Kirby, i.1893 : Lye. Afr. pi. 20, figs, i, 2. 
Type. Sierra Leone. (Berlin Mus.) 

$ unknown. 

The type of this species is in the Humboldt Universitat, Berlin, who kindly for- 
warded it for examination. It is very similar to Epitola cephena Hewitson, although 
the colour of the blue spots is paler and the insect is smaller. Without more 
material from Sierra Leone it seems inadvisable to sink it, however, to cephena. The 
type was examined by Monsieur H. Stempffer and found to be a true Epitola. 

Epitola lamborni Baker 

$ Epitola lamborni Baker, 192 1 : 461. 
Type. S. Nigeria : Ibadan. 

^ unknown. 

The type of this species is missing and it is, therefore, impossible to be sure of its 
identity. From the description, it is near alhomaculata Baker. 

THE SUBLUSTRIS GROUP 

The four species which follow differ from all known Epitola in the presence of a 
large patch of lustrous scales covering the whole discal area of the fore wing. 



NOTES ON THE EPITOLINAE 153 

Epitola pseudoconjuncta sp. n. 

(PI. 7, figs. 66, 67, PI. 8, figs. 76, 77) 

Very similar to conjunda huddvuina Talbot, but much smaller and with large patch 
of lustrous scales over the whole discal area of fore wing. 

(J and $. Legs dark brown with paler joints ; palps brown with grey scales below, black 
above ; eyes black ; frons black ; antennal shaft black above, finely checkered white below ; 
club broad and flattened, minute orange tip. 

<J. Upperside fore wing : black with dark blue spots as follows : three postdiscal in 4-6 and 
a minute dot in 7 ; blue spot half-way in 2 ; long blue streak in i, covering most of the cellule 
except angle between vein 2 and median and reaching to within 2 mm. of distal margin ; whole 
discal area up to blue postdiscal spots covered in black lustrous scales. 

Upperside hind wing : black ; large blue discal area from vein i to just short of vein 7 ; distal 
border narrow black, ^ mm. costal border black ; veins black, especially median. 

Underside fore wing : brownish grey ; large black inner-marginal patch from veins 2-5 and 
to I mm. of distal margin ; small silver-grey postdiscal spots on vein 2 and in 4 and 5, inner 
margin silver-grey ; faint traces of darker submarginal line. 

Underside hind wing : brownish grey ; faint dark submarginal and marginal lines ; no other 
markings. 

$. Legs dark brown checkered white at the joints ; palps, frons and eyes dark brown ; 
antennal shaft dark brown, boldly checkered with white, club minutely orange-tipped. 

Fore wings strongly convex, as in the male. 

Upperside : plain dark brown, unmarked. 

Underside fore wing : pale straw-coloured ; space i very pale grey, almost white ; thereafter 
a black patch from veins 1-5 ; white postdiscal spots in 2, 5 and 6 ; very obscure submarginal 
line. 

Underside hind wing : pale straw-coloured, with faint, paler, submarginal and postdiscal lines. 

Length of fore wing : 16 mm. ^ ; 15 mm. $. 

Holotype (^. Uganda: Masaka, Sango Bay, Katera. xi.1956. T. H. E. 
Jackson. Note : a beak-mark is plainly visible on left fore wing of holotype. 

Allotype $. Uganda: Masaka, Sango Bay, Katera. xi.1933. T. H. E. 
Jackson. 

This species has only been taken at Katera, where it flies with conjunda hudduana 
Talbot, which it closely resembles. Both species were taken together flying from 
high up to some 10 ft. from the ground early in the day, and the resemblance to 
the larger, more powerful conjunda is obviously protective ; pseudoconjuncta con- 
tinues to fly a little later than conjunda, up to about 11 a.m. 

Epitola tnengoensis Baker 
(PI. 9, fig. 79. PI. 10, fig. 89) 

<J Epitola mengoensis Baker, 1906 : 105. 
Type. Uganda : Mengo. 

?. Upperside : warm brown in both wings ; no markings. 

Underside fore wing : warm brown ; the usual darker basal patch, but not strongly contrasting ; 
a pale spot at end of cell and another in space 2 and the whole of i pale grey ; a series of strongly 
marked, submarginal, crescentric spots. 

Underside hind wing : warm brown ; a darker patch basad ; a well-defined crescentric, 
submarginal line followed by another, similar, but finer. 

Length of fore wing : 17 mm. 



154 T. H. E. JACKSON 

Allotype $. Mengo (Uganda), iii.1900. F. J. Jackson. 
Recorded from : Uganda : Mengo, Mpigi, Entebbe. 

Epitola vinalli Talbot 
(PI. 7, fig. 68; PI. 8, fig. 78) 
(J Epitola vinalli Talbot, 1935 : 75. 

Type. Belgian Congo : Tshuapa, Basankusu. (Hope Dept., Oxford.) 
cj Epitola subcoerulea Roche, 1954 • 49^- P^- 21, figs. 25, 26 (syn. n.). 
Type. Sierra Leone. 

The type of E. vinalli Talbot is identical with that of subcoerulea Roche and the 
latter must sink as a synonym. 

$. Legs dark brown with white spots at the joints ; eyes, palps and frons dark brown ; 
antennal shaft black above, checkered white below ; club orange-tipped. 

Upperside : plain dark brown ; distal margin of fore wing strongly convex as in male ; a 
lighter patch in fore wing between spaces 4-6, where the white postdiscal spots of underside 
show through. 

Underside fore wing : brown ; jet black inner-marginal patch from base to i mm. of margin, 
between veins 2-5 ; inner-marginal border silvery white throughout ; very small pale postdiscal 
spot on vein 2 ; two white postdiscal spots in 4 and 5 and small pale spots in 6 and 7 ; crenulate 
submarginal line. 

Underside hind wing : brown ; faint submarginal and postdiscal lines. 

Length of fore wing : 15 mm. 

Neallotype $. Fr. Equat. Africa : Moyen Congo, Etoumbi. iii . 1959. T. H. E. 
Jackson. 

Recorded from : Gambia ; Sierra Leone ; Fr. Equat. Africa : Etoumbi. 

Epitola sublustris Baker 

^, $ Epitola sublustris Baker, 1904 : 228. 
Type. Sierra Leone. 

In the Lamborn duplicate collection at the Hope Department, Oxford was found 
a female Epitola labelled : " Comp. with Type Epitola sublustris Bethune-Baker ", 
Lagos, Oni, W. A. Lamborn, iii-xii.1911. The type female is in British Museum 
(Nat. Hist.) and the Oxford specimen is a female of Epitola dolorosa Roche. 

Recorded from : Sierra Leone : Moyamba ; Ivory Coast ; Nigeria : Ubiaja, 
Oshodi ; Br. Cameroons : Kumba ; Fr. Cameroons : Johann Albrechts Hohe ; 
Ghana : Aburi, Ashanti. 

The following species differs from all other known Epitola, in the underside mark- 
ings, but, from the convex distal margin of the fore wing, belongs near conjuncta. 
No lustrous scales in fore wing. 

Epitola barombiensis Kirby 

(PI. 9, fig. 80 ; PI. 10, fig. 90) 
(^ Epitola barombiensis Kirby, 1890 : 274. 
Type. Cameroons : Barombi. (Staudinger Coll.) 

$. Upperside : both wings dark brown— unmarked. 
Underside : as in the male, but paler. 

Neallotype $. Uganda: Bwamba. ii-iii.1957. R. Carcasson. 



NOTES ON THE EPITOLINAE 155 

THE ALBA GROUP 

The following new species is quite unlike any known Epitola and must remain 
in a group of its own, 

Epitola alba sp. n. 

(PI. 9, figs. 81, 82, PI. 10, figs. 91, 92) 

(^. Legs brown, faintly paler at the joints ; eyes, palps and frons brown ; antennal shaft 
black above, minutely checkered below ; club minutely yellow -tipped. 

Upperside fore wing : black-brown ; large white discal area from ia-6, scaled blue at the base, 
distally rounded and dentate ; a broad, triangular black spot at end of cell from costa ; costal 
margin narrowly black ; wide black-brown distal border ; median and subcostal veins swollen 
at base, the former black to lower angle of cell. 

Upperside hind wing : black-brown ; white discal patch between veins 1-6, from base over 
cell, bases of 2, 3 and 4 and half space i ; broad black distal and costal borders ; the white discal 
patch is elongated, being broader on vein 6 than on vein i. 

Underside fore wing : dark silver-grey ; white discal patch as above ending at the middle of 
the costa in 10 ; a broad dark grey postdiscal band to vein 4, with fine pale line distad ; small 
rounded basal patch on mid-costa, extending basad to inner margin ; very obscure submarginal 
line ; distal border broadly dark grey. 

Underside hind wing : dark silver-grey ; white discal patch as above ; some obscure dark 
grey costal markings ; an obscure dark postdiscal line ; a crenulated dark grey submarginal 
line followed by a fine pale line ; border dark grey, i mm. 

$. Upperside : similar to male ; basal blue area fore wing smaller and median vein not black ; 
triangular cistal spot larger and darker, hind wing as in the male but basal blue smaller. 

Underside : paler, more brownish but markings as in the male ; broad dark postdiscal band 
extends to vein 3, fore wing ; the postdiscal line on hind wing shifted further distad ; otherwise 
as in the male. 

Length of fore wing : (J 16 mm.; $ 17 mm. 

Holotype cJ. Fr. Equat. Africa : Ouesso, Mambili Forest, vi.1959. T. H. E. 
Jackson. 

Allotype. Same data as holotype. viii.1959. 
Recorded only from the above locality. 

SPECIES INCERTAE SEDIS 

Epitola pulverulenta Dufrane 

Epitola pulverulenta Dufrane, 1953 : 51. 

Type. Monsieur Dufrane states that the type is lost. 

Since many of these small brown females are very much alike, it is not possible 
to be certain of an identity from a description alone. 

SPECIES INCORRECTLY PLACED IN EPITOLA 
Phytala benitensis (Holland) (comb, n.) 

cj Epitola benitensis Holland, 1890 : 425. 

Type. Gaboon : " Upper waters of the River Ogove." (Carnegie Mus., Pittsburgh.) 

This species belongs to Phytala West wood. Mr. Harry K. Clench of the Car- 
negie Museum, Pittsburgh, who has kindly examined the type, writes as follows : 
" I have checked the type for venation and find that it is a Phytala. From the figures 

BNTOM. 12, 3. 10 



156 T. H. E. JACKSON 

in Grose-Smith & Kirby 1893 [Rhop. Exot. 23 ; Lye. Afr, 86) I suspect it to be very 
close to rezia Grose-Smith & Kirby, if not the same ". The figure and description 
of the female in Grose-Smith & Kirby, loc. cit., is that of a male. The authors state, 
in a footnote, that rezia does not have the peculiar falcate fore wings of henitensis 
Holland, and it is considered by the present author that the two species are distinct. 

Liptena langi (Holland) (comb, n.) 

cJ Epitola langi Holland, 1920 : 217, pi. 12, fig. 10. 
Type. Congo : Medje. (American Mus., N. York.) 

Judging by the coloured plate this insect appears to be a male of Liptena despecta 
Holland and Mr. Frederick H. Rindge of the American Museum, New York, who 
has kindly examined the type, writes : "I have examined Holland's type. By 
using the keys and discussion in Seitz, I have concluded that langi Holland should 
definitely be placed in the genus Liptena. The venation is of this genus and not of 
Epitola." There appears to be very little doubt that Epitola langi should sink 
to Liptena despecta Holland. 

Phytala rezia (Grose-Smith & Kirby) (comb, n.) 
$ Epitola rezia Grose-Smith & Kirby, i.1893 : Lye. Afr. 86, pi. 20, fig. 6. 
This species belongs to Phytala West wood. 

Aslauga subfulvida (Holland) (comb, n.) 

$ Epitola subfulvida Holland, 1890 : 423. 

Aslauga leonae Aurivillius in Seitz, 1920 : 343 (syn. n.) 

Aslauga purpurascens (Holland) (comb, n.) 
$ Epitola purpurascens Holland, 1890 : 424. 

Through the kindness of Mr. Harry K. Clench of the Carnegie Museum, Pittsburg, 
I am enabled to state that the above two species belong to Aslauga Kirby, the first 
being synonymous with leonae Aurivillius, and the second with Aslauga purpurascens 
HoUand. 

DISCUSSION 

Comparing this arrangement with the key in Roche (1954), the following, more 
important, points emerge : 

{a) The hew it son i and posthumus groups remain as before. 

(&) Part of the first section of the pinodes group is treated separately and is 
named the sublustris group. This includes four species listed by Roche with 
convex distal margin in fore wing. All four are small and have a large patch of 
lustrous scales in the discal area of the fore wing. 

(c) The remainder of the pinodes group remains as before, but subdivides on the 
pattern of the females into two. The species concerned, however, intergrade and 
are obviously closely allied. 

(d) On the pattern of the females the carcina and nitide groups merge and, 
from a study of the uppersides of the males, and of the pattern of the females, 



NOTES ON THE EPITOLINAE 157 

this is probably correct. On the other hand the four subdivisions are rather further 
from each other than are those of the Pi nodes group. 

[e) Epitola alba sp. n., cannot be united with any of the existing groups and 
Epitola haromhiensis Kirby is separated on its pecuhar underside. 

Genus STEMPFFERIA gen. n. 

Type species: Stempfferia carcassoni sp. n. 

Differs from Deloneura Trimen and Neaveia Druce in the genital armature ; from 
Batelusia Druce in the venation (only ii veins in the latter) ; from Turner epes 
Baker in that the legs are not swollen ; from Epitola Westwood, Hewitsonia Kirby 
and Phytula Westwood in the pattern and in the shape of the wings ; eyes faintly 
hairy ; second segment of palps, laterally compressed ; third segment cylindrical, 
terminating in a point ; antennae half length of costa ; club slightly swollen. 

Wings ; costal border of fore wing arched at base, then straight ; external border strongly 
convex ; hind wing suboval, anal angle obtuse. In both sexes a fringe of long hair along inner- 
margin of fore wings. Venation : fore wing 12 veins, median much swollen at base in male, less 
so in female ; vein 3 from a little before the lower angle of the cell : vein 4 from this angle ; 
vein 5 from much nearer 6 than 4 ; vein 6 from the upper angle of the cell ; vein 7 from well 
before this angle, ending below the apex ; vein 8 branching from 7 and ending on the costal 
border ; vein 9 long, branching from 7 ; in the male vein 10, feebly stalked, arising from 7, not 
far from its origin ; vein 11 free, from same origin as 7 ; in the female ; vein 10 arising from the 
same point as 7 ; vein 1 1 from a little before ; hind wing ; veins 3 and 4 briefly stalked, from 
lower angle of cell ; lower discocellular concave ; vein 5 nearer 6 than 4 ; 7 from a little before 
the lower angle of cell ; genital armature ; of the usual Epitolinae type ; uncus in the form of a 
crescent, edging the subtriangular tegumen ; subunci long, arched near the base ; swollen at the 
level of the bend, ending in a slightly curved fine point (in the figured slide, the subunci have been 
set aside laterally in order to be better visible, but in situ they are folded up below the tegumen). 
Vinculum rather narrow, prolonged by a triangular saccus which is, at its extremity, furnished 
with long scales ; lower fultura very reduced, valvae subrectangular, the upper process ending 
in a slightly curved point, penis short, very stout, the distal extremity obliquely cut and widely 
open on the dorsal side. Uncus furnished with long, fine hair, much fine hair also on the distal 
half of the upper process of valvae (in the figure, the bases of hair only are figured, this for the 
sake of clearness). 

Sexual dimorphism very striking ; the male upperside is blue, that of the female yellowish 
white. 

This genus is named in honour of Monsieur H. Stempffer of Paris, who, more than 
any other person, has succeeded in unravelling the complexities of the African 
Lycaenidae, and achieving an orderly presentation of their systematics. 

Stempfferia carcassoni sp. n. 

(PI. II, figs. 103, 108, PI. 12, figs. 113, 118) 

^ and $. Legs, frons and palps black ; eyes brown, slightly hairy ; antennal shaft black 
above, checkered white below ; wings rounded ; fore wing obtusely angled at apex ; fringes 
white, checkered black at the veins. 

(? Upperside fore wing : black with broad blue discal-basal patch from inner margin to costa, 
extending to within 2 mm. of the margin along inner margin and to within 9 mm, of apex along 
costa ; distal edge evenly rounded ; distal border broadly black. 



158 



T. H. E. JACKSON 



Upper side hind wing : black with blue discal patch from half space ic to vein 6, evenly rounded 
distally, costal and distal margins and space ih to half ic, black ; costal margin 3 mm. wide 
throughout. 

Underside : both wings lustrous white, unmarked. 

$. As in the male, but larger and blue areas replaced by yellowish white. 

Upperside fore wing : black with yellowish white discal area from base and inner margin to 
vein 7, with a small extension in space 7 ; broad black distal border and narrow black costal 
border ; discal patch evenly curved distad. 

Upperside hind wing : black with yellowish white discal patch, extending from base and inner 
margin through to the costa ; brownish black distal border 4 mm. wide. 

Underside : as in the male, but less lustrous. 




Fig. I. Genital armature of male Stempfferia carcassoni gen. et sp. n. 

Holotype (^. Congo: Prov. Orientale, Lomami River, Opala. iii.1959. R. H. 
Carcasson. 

Allotype $. Same data as holotype. 

Recorded from : Fr. Equat. Africa : Ouesso ; Congo : Lomami River. 

This fine new genus and species was first discovered by Mr. R. H. Carcasson of 
the Coryndon Museum, Nairobi in the Congo, and subsequently has been taken 
also in French Equatorial Africa. He has kindly allowed me to describe it in this 
paper and the species is named in his honour. 



Genus NEAVEIA Druce 
Neaveia lamborni orientalis ssp. n. 

(PI. II, figs. loi, 102, PI. 12, figs. Ill, 112) 

The eastern subspecies has not been described. Differs from the typical race in 
narrower apical black patch on fore wing above and in the presence of submargina] 
and postdiscal lines in hind wing beneath. 



NOTES ON THE EPITOLINAE 159 

^ and $. Legs dark brown, paler at the joints ; palps short, dark brown ; frons brown with 
many white scales ; eyes black ; antennal shaft black above, checkered white below ; club 
black. 

cJ. Upperside fore wing : greenish white ; black apical patch to vein 3, then continued as a 
thin marginal line ; in lamborni lamborni it is still i mm. wide on vein i ; costal border only 
dusted with black scales. 

Upperside hind wing : greenish white ; black apical patch to vein 3, then continued as a thin 
marginal line ; in lamborni lamborni it is still i mm. wide on vein i ; costal border only dusted 
with black scales. 

Underside fore wing : white ; black apical patch shows through from above and is narrow 
and, as above, ends at vein 3 ; some small brown striae along the margin from vein 3 to the apex 
and others, submarginal, in 4, 5 and 6 and along the distal half of the costal margin. 

Underside hind wing : strongly green ; marginal border striated with scattered brown scales ; 
a series of submarginal brown spots in spaces 1-8 ; another series postdiscal, arrow-shaped ; 
a few obscure discal markings ; a fine brown line on the discocellular. 

$. Similar to male, but markings below a little fainter. 

Length of fore wing : ^ and ?, 20 mm. 

Holotype (J. Uganda: Masaka, Sango Bay, Katera. xi.1954. T. H, E. 
Jackson. 

Allotype $. Same data as holotype. xii.1954. 

This species was observed flying in the canopy of lichen-covered trees among 
Crematogaster ants at Katera, Sango Bay. 

Recorded from : Fr. Equat. Africa and the eastern Congo ; Uganda : Bwamba, 
Budongo, Katera. 

Genus EPITOLINA Aurivillius 

Epitolina catori ugandae ssp. n. 

(PI. II, figs. 99, 100, PL 12, figs. 109, no) 

Differs from catori catori Baker in the much larger orange postdiscal band in the 
fore wing of the female and in the darker underside in both sexes. 

cj and ?. Legs dark brown covered in yellow hair ; eyes and frons black ; palps black above, 
yellow below ; antennal shaft widely checkered black and white on both surfaces ; club broad, 
flattened, minute pale tip. 

cj. Upperside : does not differ from catori catori. 

Underside : much darker, red markings more prominent ; on both wings a broad black 
postdiscal band, distal of the red spots. 

$ Upperside : the orange patch on fore wing extends from veins 10-5, with a small spot also 
in space 5 ; proximad it runs in a curve skirting the cell to the origin of vein 2, then vertically to 
la ; distally covers the base of spaces 4 and 5, half 2 and 3, and the middle of i and la ; 3 mm. 
broad in i and la, and 5 mm. on vein 2. 

Underside : as in catori catori, but a broad blackish brown postdiscal band in both wings distal 
of the orange spots ; these bands are very prominent. 

Length of fore wing : ^ and $,12 mm. 

Holotype (^. Uganda: Mpigi, Mpanga Forest, viii.1959. T. H. E. Jackson, 
Allotype $. Same data as holotype. iv.1959. 
Recorded also from many other parts of Uganda. 



i6o T. H. E. JACKSON 

Genus HEWITSONIA Kirby 

Hewitsonia boisduvalii (Hewitson) 

Cory don boisduvalii Hewitson, 1869 : i, pi. i. 

Hewitsonia boisduvalii (Hewitson) Hewitson, 1878 : 17, pi. \a, figs, i, 2. 

The type of this species, a female, came from Gaboon and occurs throughout the 
Southern Cameroon and French Equatorial Africa in the same form, changing to 
subspecies congoensis Joicey & Talbot in the eastern Congo. Specimens from Nigeria 
and Northern Cameroons differ considerably and warrant a further subspecific 
name. 



Hewitsonia boisduvalii nigeriensis ssp. n. 

(PI. II, figs. 104, 105, PI. 12, figs. 114, 115) 

cj. Differs from the typical race as follows ; the four subapical spots are small, rounded, 
blue with white centres ; the spot in space 5, as an example, is 2 mm. long, as against 7 mm. 
The blue colour is darker. The small blue spots extending from the blue patch in the anal angle 
fore wing into the base of space 2 are here reduced to one small blue streak and a few scattered 
scales in the base. On hind wing blue patch less extensive, particularly in spaces 5 and 6. 

Below, as in typical boisduvalii. 

$. Subapical yellow band narrower, 4 mm. wide as against 6 mm. in boisduvalii ; yellow 
spot in inner angle fore wing 6 mm. long, as against 8 mm.; otherwise not differing. Below not 
differing. 

This race is on average smaller than the typical race and in this agrees with congoensis Joicey 
& Talbot. 

Holotype c^. Nigeria: Lagos District, Oshodi. iv.1955. T. H. E. Jackson. 
Allotype $. Nigeria: Onitsha Prov., Awka, Mamu. xi.1959. T. H. E. 
Jackson. 

Hewitsonia similis ugandae ssp. n. 

(PI. II, figs. 106, 107, PI. 12, figs. 116, 117) 
This is the eastern subspecies and differs as follows : 

<?. Upperside : differs from similis similis Aurivillius in the reduction of the blue spots in 
fore wing ; the spot in the inner angle is missing or represented by a few blue scales ; no blue 
spots in the cell and those in 2 and 3 much reduced in size ; four white subapical spots in fore 
wing ; hind wing as in the typical form but with a series of triangular black marginal spots, one 
in each cellule. 

?. Upperside : the subapical white or cream spots are larger than in specimens from Nigeria 
and Sierra Leone, but smaller than those from the Cameroons — otherwise not differing. 

Underside : both sexes ; differs in the much paler ground colour and reduction in the black 
streaks on hind wing ; the red-brown patch at the distal end of the hind wing costa is scarcely 
visible, and does not contrast with the rest of the wing. 

Holotype c^. Congo: Ituri, Beni, 4,000 ft. ix.1946. T. H. E. Jackson. 
Allotype $. Uganda: Bwamba. ix.1942. T. H. E. Jackson. 



NOTES ON THE EPITOLINAE l6i 

Hewitsonia intermedia Joicey & Talbot (stat. n.) 

Hewitsonia kirhyi ? f. intermedia Joicey & Talbot, 192 1 : 86. 

This insect differs greatly from kirbyi Hewitson and should be considered as a 
separate species, 

ACKNOWLEDGEMENTS 

My thanks are due to Monsieur H. Stempffer of Paris, for much help and advice 
in writing this paper. To Mr. Harry K. Clench, Carnegie Museum, Pittsburg, and 
Mr. Frederick H. Rindge, American Museum, New York, for information over 
Holland's types. To the Keeper and staff of the Department of Entomology, 
British Museum (Natural History) for all facilities, and particularly to Captain 
N. D. Riley, C.B.E., Messrs. W. H. T. Tams and T. G. Howarth for their advice. 
To the Director, Humboldt Universitat, Berhn, for kindly lending me their types. 



REFERENCES 

AuDEOUD, G. E. 1936. Description de six especes ou sous-especes nouvelles de Rhopaloceres 

africains. Mitt, schweiz. ent. Ges. 16 : 702-705, pi. 7, figs. 3, 4. 
AuRiviLLius, C. 1898. Rhopalocera Aethiopica. 561 pp. Stockholm. 

1920. In Seitz, Macrolepidoptera of the World, 13 : pis. 64, 65. Stuttgart. 

1923. Fortsetzung. Ergbn. zweit. Deutsch. Zentr. Afr. Exped. 1910-11, Zool. 17 : 1195- 

1241. 
Baker, G. T. B. 1903. On new species of Lycaenidae from West Africa. Ann. Mag. nat. Hist. 

(7) 12 : 324-334. 

1904- On new species of Rhopalocera from Sierra Leone. Ibid. (7) 14 : 222-233. 

1906. Description of some new African Butterflies. Ibid. (7) 17 : 104-110. 

1913- Notes on Lycaenidae collected by W. A. Lamborn in the Lagos district of West 

Africa with new species. Trans. R. ent. Soc. Lond. 46 : 499-507. 

1915- Descriptions of new species of Lepidoptera from Africa and the East. Ann. Mag. 

nat. Hist. (8) 16 : 186-203. 

1916. Descriptions of new species of Lepidoptera. Ibid. (8) 17 : 378-385. 

1 92 1. Descriptions of new species of Lepidoptera, chiefly Lycaenidae from Southern 

Nigeria, and one from Damba Island, Victoria Nyanza. Trans. R. ent. Soc. Lond. 54 : 

461-464. 
1926. Descriptions of new species of Rhopalocera from the Ethiopian Region. Ann. 

Mag. nat. Hist. (9) 17 : 384-402. 
Druce, H. H. 1890. Descriptions of twelve new species of Lycaenidae from West Africa 

and one from Solomon Islands, in the Coll. of Herbert Druce. Ibid. (6) 5 : 24-31. 

1903. Descriptions of four new species of West African Lycaenidae. Ibid. (7) 11 : 69- 72. 

1 9 10. Descriptions of new Lycaenidae and Hesperidae from tropical West Africa. Proc. 

zool. Soc. Lond. 1910 : 356-378. 
DuFRANE, A. 1953. L^pidopteres du Kivu. Bull. {Ann.) Soc. ent. Belg. 89 : 41-57. 
Fabricius, J. C. 1793. Ent. Syst. Emend. 3 {i). 349 pp. Stockholm. 
Grose-Smith, H. & Kirby, W. F. 1887-97. Rhopalocera Exotica, 1-2. London. 
1898. Descriptions of new species of African butterflies in the Tring Museum. 

Novit. Zool. 5 : 350-358. 
Hawker-Smith, W. 1926. Some new Lycaenidae in the Congo Museum. Rev. zool. afr. 

14 : 237-241. 
1933- New species and races of Lipteninae (Lepidoptera, Lycaenidae). Stylops, 2 : i-ii. 



Hewitson, W. C. 1869-78. Illustrations of Diurnal Lepidoptera, Suppl. 47 pp. London. 



I62 T. H. E. JACKSON 

Hewitson, W. C. 1873. Descriptions of six new species of Epitola from the west coast 
of Africa. Ent. mon. Mag. 10 : 149-151. 

1874. A list of butterflies taken on the march to Coomassie with descriptions of new 

species. Ann. Mag. nat. Hist. (4) 13 : 380-383. 

Holland, W. J. 1890. Descriptions of new West African Lycaenidae. Psyche, 5 : 423-431. 

1920. Lepidoptera of the Congo, being a systematic list of butterflies and moths collected 

by the American Museum of Natural History Congo Expedition, together with descriptions 
of hitherto undescribed species. Bull. Amer. Mus. nat. Hist. 43 : 109-331, pi. 12, fig. 10. 

JoiCEY, J. J. & Talbot, G. 192 i. New Lepidoptera collected by Mr. T. A. Barnes, in East 

Central Africa. Bull. Hill. Mus. 1 : 40-166, pi. 14. 
KiRBY, W. F. 1887. Descriptions of new species of Epitola from the Cameroons, &c. in the 

collection of Henley Grose-Smith. Ann. Mag. nat. Hist. (5) 19 : 441-445. 
1890. Descriptions of new species of African Lycaeinidae, chiefly from the collections of 

Dr. Staudinger & Mr. Henley Grose-Smith. Ibid. (6) 6 : 261-274. 
Mabille, p. 1877. Catalogue des L6pidopteres du Congo. Bull. Soc. zool. Fr. 2 : 214-221. 
Roche, P. 1954. Revisional notes on the genus Epitola Westwood (Lepidoptera : Lycaenidae). 

Bull. Brit. Mus. (nat. Hist.) Entom. 3 : 489-501, pis. 19-22. 
PouLTON, E. B. 1929. African Rhopalocera : Descriptions and notes. Trans. R. ent. Soc. 

Lond. 77 : 475-504. pi. 30. 
Seitz, a. 1908. Macrolepidoptera of the World, 13 : 1-613, pi. 64, 65. 
Schultze, a. 191 1. Eine neue Epitola aus dem afrikanischen Aequatorial-Urwalde. Iris, 

25 : 95-96. 
Sharpe, E. M. 1890. On some new species of African Lycaenidae in the collection of Philip 

Crowley. Ann. Mag. nat. Hist. (6) 6 : 103-106. 
Staudinger, O. 1888. Exotische Schmetterlinge, 1. 333 pp. Fiirth. 

1889. Zwei neue afrikanische Lycaeniden. Ent. Nachr. 15 : 176-180. 

Stempffer, H. 1956. Contribution a I'^tude des L6pidopteres Lycaenidae de I'Afrique 

^quatoriale. Ann. Mus. Cong. belg. (8) 49 : 7-52. 
Stoneham, H. F. 1933. New species of Lycaeinidae Leach from Kenya Colony, British East 

Africa. Bull. Stoneham Mus. 17 : 1-3, pi. i. 
Suffert, E. 1904. Neue afrikanische Tagfalter. Iris, 17 : 12-107. 
Talbot, G. 1935. New species and forms of African Lycaenidae (Lep. Rhop.). Ent. mon. 

Mag. 71 : 69-78. 
1937- New African Lycaeinidae and Nymphalidae and two new Diestogyna (Lepidoptera). 

Trans. R. ent. Soc. Lond. 86 : 59-72, pis. 1-2. 



INDEX 



163 



adoiphifriderici, 130 
aequatorialis, 135 
alba, 155 

albomaculata, 138 
ammon, 129 
Aslauga, 156 
azurea, 148 

badia, 137 
badura, 136 
badwia, 136 
barombiensis, 154 
batesi, 132 
Batelusia, 157 
bella, 142 
belli, 128 
benitensis, 155 
boisduvalii, 166 
budduana, 151 

carcassoni, 157 
carcina, 140 
carilla, 134 
carpenteri, 144 
catori, 159 
catuna, 143 
cephena, 146 
ceraunia, 129 
cercene, 131 
cercenoides, 132 
ciconia, 136 
coerulea, 140 
concepcion, 139 
congoana, 140 
congoensis, 160 
conjuncta, 151 
convexa, 131 
crippsi, 126 
crowleyi, 129 

daveyi, 138 
decellei, 127 
Deloneura, 157 
despecta, 156 
dewitzi, 129 
doleta, 146 
dolorosa, 152 
dorothea, 133 
dunia, 141 



INDEX 

elissa, 139 
entebbeana, 146 
Epitola, 125 
Epitolina, 159 

flavoantennata, 140 

gerina, 138 
ghesquieri, 145 
goodii, 138 
gordoni, 135 

hewitsoni, 126 
Hewitsonia, 160 
hewitsonioides, 126 

ikoya, 140 
insulana, 133 
intermedia, 131, 161 
iturina, 142 

jacksoni, 135 

kamengensis, 150 
katerae, 149 
katherinae, 134 
kholifa, 140 
kirbyi, 161 

lamborni, 152 
langi, 156 
leonae, 156 
leonensis, 146 
leonina, 136, 146 
liana, 138 

maculata, 152 
mara, 135 
marginata, 130 
mengoensis, 153 
mercedes, 147 
miranda, 126 
moyambina, 130 
mpangensis, 149 
mus, 144 

Neaveia, 158 
nigeriae, 145 
nigeriensis, 160 



nigra, 145 
nigrovenata, 136 
nitida, 139 
nitide, 139 

obscura, 152 
oniensis, 139 
orientalis, 143, 158 
ouesso, 147 

Phytala, 155 
pinodes, 143 
pinodoides, 152 
posthumus, 128 
pulverulenta, 155 
purpurascens, 156 
pseudoconjuncta, 153 

radiata, 143 
rezia, 156 
rileyi, 150 

semibrunnea, 129 
similis, 160 
staudingeri, 134 
stamp fferi, 127 
Stempfiferia, 157 
subalba, 139 
subcoerulea, 154 
subfulvida, 156 
sublustris, 154 

tanganikensis, 129 
tumentia, 137 
Tumerepes, 157 

ugandae, 159, 160 
umbratilis, 130 
uniformis, 132 
urania, 128 

versicolor, 131 
vidua, 127 
vinalli, 154 
virginea, 139 
viridana, 142 

zelica, 138 
zelza, 137 



ENTOM. 12, 3. 



PLATE I 

Figs. 1-9. Uppersides of Epitola : (i) miranda miranda Staudinger neallotype female, 
(2) miranda vidua Talbot neallotype male, (3) urania urania Kirby neallotype female, (4) urania 
tanganikensis Joicey & Talbot neallotype female, (5) moyambina Baker neallotype female, 
(6) intermedia Roche neallotype female, (7) convexa Roche neallotype female, (8) uniformis 
Kirby neallotype female, (9) insulana Aurivillius neallotype female. 



4 JUN'9&2 I 



Bull. B.M. (N.H.) Enlom. 12, 3 



PLATE I 









8 




PLATE 2 

Figs. 10-18. Undersides of Epitola : (10) miranda miranda Staudinger neallotype female, 
(11) miranda vidua Talbot neallotype male, (12) urania urania Kirby neallotype female, (13) 
urania tanganikensis Joicey & Talbot neallotype female, (14) moyambina Baker neallotype 
female, (15) intermedia Roche neallotype female, (16) convexa Roche neallotype female, (17) 
uniformis Kirby neallotype female, (18) insiilana Aurivillius neallotype female. 



Bull. B.M. (N.H.) Entom. 12, 3 



PLATE 2 





14 





y 



15 





17 




18 



PLATE 3 

Figs. 19-28. Uppersides of Epitola : (19) staudingeri Kirby neallotype female, (20) mara 
Talbot neallotype female, (21) leonina Staudinger neallotype female, (22) nigrovenata sp. n. 
holotype male, (23) nigrovenata sp. n. allotype female, (24) tumentia Druce neallotype female, 
{25) elissa Grose-Smith neallotype female, (26) coernlea sp. n. holotype male, (27) coerulea sp. n. 
allotype female, (28) dunia Kirby neallotype female. 



Bull. B.M. (N.H.) Entom. 12, 3 



PLATE 3 











m 1 


1 




^19 


•^^#^ 




y 

20 


^mI 







28 



PLATE 4 

Figs. 29-38. Undersides of Epitola : (29) staudingeri Kirby neallotype female, (30) mara 
Talbot neallotype female, (31) leonina Staudinger neallotype female, (32) nigrovenata sp. n. 
holotype male (33) nigrovenata sp. n. allotype female, (34) tumentia Druce neallotype female, 
(35) elissa Grose-Smith neallotype female, (36) coerulea sp. n. holotype male, (37) coerulea sp. n. 
allotype female, (38) dnnia Kirby neallotype female. 



Bull. B.M. (N.H.) Entom. 12. 3 



PLATE 4 





29 



34 




^^ 



30 



35 




i 



31 



#^ 



36 




32 






33 



38 



PLATE 5 

Figs. 39-48. Uppersides of Epitola : (39) viridana viridana Joicey & Talbot neallotype 
female, (40) catuna Kirby neallotype female, (41) carpenteri Baker neallotype female, (42) 
orientalis Roche neallotype female, (43) ghesquieri Roche neallotype female, (44) nigeriae sp. n. 
holotype male, (45) nigeriae sp. n. allotype female, (46) cephena entebbeana Baker neallotype 
female, (47) mercedes Suffert neallotype female, (48) mpangensis sp. n. allotype female. 



Bull. B.M. (N.H.) Entom. 12, 3 



PLATE 5 





39 




40 




41 




42 






44 




45 




*«s-^;r* 



46 




47 




43 



48 



PLATE 6 



Figs. 49-58. Undersides of Epitola : (49) viridana viridana Joicey & Talbot neallotype 
female, (50) catuna Kirby neallotype female, (51) carpenteri Baker neallotype female, (52) 
orientalis Roche neallotype female, (53) ghesquieri Roche neallotype female, (54) nigeriae sp. n. 
holotype male, (55) nigeriae sp. n. allotype female, (56) cephena entebbeana Baker neallotype 
female, (57) mercedes Suffert neallotype female, (58) mpangensis sp. n. allotype female. 



Hull. B.M. {N.H.) Enfom. 12, 3 



PLATE 6 





49 



54 




50 





55 




51 




56 




52 




57 



'^ir 



53 




PLATE 7 

Figs. 59-68. Uppersides of Epitola : (59) oiiesso sp. n. holotype male, (60) ouesso sp. n. 
allotype female, (61) aziirea sp. n. holotype male, (62) katerae sp. n. holotype male, (63) katerae 
sp. n. allotype female, (64) kamengensis sp. n. holotype male, (65) kaniengensis sp. n. allotype 
female, (66) pseudoconjuncta sp. n. holotype male, (67) pseudoconjimcta sp. n. allotype female, 
(68) vinalH Talbot neallotype female. 



Bull. B.M. (N.H.) Entom. 12, 3 



PLATE 7 





59 



64 



%m 




60 



65 



M 




61 



66 







62 



67 




Vw 



63 



68 



PLATE 8 



Figs. 69-78. Undersides of Epitola : (69J ouesso sp. n. holotype male, (70) ouesso sp. n. 
allotype female, (71) azurea sp. n. holotype male, (72) katerae sp. n. holotype male, (73) katerae 
sp. n. allotype female, (74) kamengensis sp. n. holotype male, (75) kamengensis sp. n. allotype 
female, (76) pseudoconjuncia sp. n. holotype male, (77) pseudoconjuncta sp. n. allotype female, 
(78) vinalli Talbot neallotype female. 



Bull. B.M. (N.H.) Entom. 12, 3 



PLATE 8 



\ / 





69 





/ 



70 



75 



^W^ 




71 



76 





72 



77 







73 



78 



PLATE 9 

Figs. 79-88. Uppersides of Epitola : (79) mengoensis Baker neallotype female, (80) barom- 
biensis Kirby neallotype female, (81) alba sp. n. holotype male, (82) alba sp. n. allotype female, 
(83) carpenteri Baker male, (84) zelza Hewitson male, (85) urania urania Kirby male, (86) 
iturina Joicey & Talbot male, (87) dorothea Baker male, (88) insulana Aurivillius male. 



Bull. B.M. {N.H.) Entom. 12, 1 



PLATE 9 




79 




80 




%ir -m0 



82 




83 




PLATE lo 



Figs. 89-98. Undersides of Epitola : (89) mengoensis Baker neallotype female, (90) barom- 
biensis Kirby neallotype female, (91) alba sp. n. holotype male, (92) alba sp. n. allotype female, 
(93) carpenteri Baker male, (94) zelza Hewitson male, (95) urania urania Kirby male, (96) iturina 
Joicey & Talbot male, (97) dorothea Baker male, (98) insulana Aurivillius male. 



Bull. B.M. (N.H.) Entom. 12, 3 



PLATE 10 




89 




90 





91 





92 



97 




93 




PLATE II 

Figs. 99-108. Uppersides of Epitolina, Neaveia, Hewitsonia and Stempfferia : (99) E. catori 
ugandae ssp. n. holotype male, (100) E. catori ugandae ssp. n. allotype female, (loi) N. lamborni 
orientalis ssp. n. holotype male, (102) N. lamborni orientalis ssp. n. allotype female, (103) Stemp- 
fferia carcassoni gen. et sp. n. holotype male, (104) H. boisduvalii nigeriensis ssp. n. holotype 
male, (105) H. boisduvalii nigeriensis ssp. n. allotype female, (106) H. similis ugandae ssp. n. 
holotype male, (107) H. similis ugandae ssp. n. allotype female, (108) Stempfferia carcassoni 
gen. et sp. n. allotype female. 



Bull. B.M. {N.H.) Entom. 12, 3 



PLATE II 



vfv 



99 



«tf 



100 






101 



f^V^ 



102 





106 



103 




108 



PLATE 12 

Figs. ioq-ii8. Undersides of Epitolina, Neavia, Hewitsonia and Stempfferia : (109) E. catori 
ugandae ssp. n. holotype male, (no) E. catori ugandae ssp. n. allotype female, (in) N. lamborni 
orientalis ssp. n. holotype male, (112) .V. lamborni orientalis ssp. n. allotype female, (113) Stemp- 
fferia carcassoni gen. et sp. n. holotype male, (114) H. boisduvalii nigeriensis ssp. n. holotype 
male, (115) H. boisduvalii nigeriensis ssp. n. allotype female, (116) H. similis ugandae ssp. n. 
holotype male, (117) H. similis ugandae allotype female, (118) Stempfferia carcassoni gen. et 
sp. n. allotype female. 



Bull. B.M. (N.H.) Entom. 12, 3 



PLATE 12 




109 



M 



no 



r^M^^ 



111 



r4 1 



112 




117 




113 



118 



PLATE 13 

Figs, i 19-130. Uppersides and undersides of Epitola : (119, 125) liana Koche neallotype 
female, (120, 126) staudingeri aequatorialis ssp. n. holotype male, (121, 127) staudingeri aequa- 
torialis ssp. n. allotype female, (122, 128) stempfferi sp. n. holotype male, (123, 129) stempfferi 
sp. n. allotype female, (124, 130) hewitsoni Mabille holotype male. 



UitU. B.M. (N.I I.) Enlom. 12, 3 



PLATE 13 




124 




130 



PRINTED IN GREAT BRITAIN BY 
ADLARD AND SON, LIMITED, 
BARTHOLOMEW PRESS, DORKING 



6. 

A REVISION OF AFRICAN 

LAEMOPHLOEINAE 
(COLEOPTERA : CUCUJIDAE) 



L. P. LEFKOVITCH 




BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 12 No. 4 

LONDON: 1962 



A REVISION OF AFRICAN LAEMOPHLOEINAE 
(COLEOPTERA : CUCUJIDAE) 



BY 



L. P. LEFKOVITCH 

Pest Infestation Laboratory, Slough 



Yv^| 



Pp. 165-245 ; 82 Text-figures 




BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 12 No. 4 

LONDON : 1962 



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. 

This paper is Vol. 12, No. 4 of the Entomological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 



© Trustees of the British Museum 1962 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued 17 July 1962 Price Twenty-seven shillings and sixpence 



A REVISION OF AFRICAN LAEMOPHLOEINAE 
(COLEOPTERA : CUCUJIDAE) 

By L. P. LEFKOVITCH 



CONTENTS 

Introduction ...... 

Structure of the Adult .... 

Key to Genera of African Laemophloeinae 
Descriptions of Genera and Species, with Keys 
I. Cannes gen. n. 
II. Placonoius Macleay 

III. Xylolestes gen. n. 

IV. Xylophloeus gen. n. 
V. Blubos gen. n. 

VI. Microlaemus gen. n. 

VII. Magnoleptus gen. n. 

VIII. Leptophloeus Casey 

IX. Narthecius Leconte 

X. Passandrophloeus Kessel 

XI. Mestolaemus gen. n. 

XII. Mariolaemus gen. n. 

XIII. Notolaemus Lefkovitch 

XIV. Planolestes Lefkovitch 
XV. Cucujinus Arrow 

Paracucujinus subgen. n. 
XVI. Cryptolestes Ganglbauer 
Acknowledgements . 
Appendix .... 
References 
Index .... 



Page 
167 
168 
171 
174 
174 
176 
187 
189 
194 
196 
199 
201 
210 
213 

215 
216 
220 
226 
228 
230 

231 
240 
241 

242 
243 



SYNOPSIS 
Keys to, and descriptions of, sixteen genera, one subgenus and eighty-six species of Laemo- 
phloeinae from the Ethiopian region (including Madagascar, Madeira, the Canaries and the 
Salvages) and some North African species which are primarily Palaearctic are provided. Eight 
new genera, one new subgenus and thirty-nine new species are described ; two new generic 
and eleven new specific synonymies are made. 



INTRODUCTION 

In a previous paper (Lefkovitch, 19596), it was pointed out that Laemophloeus 
was a heterogeneous group. Based upon a study of species occurring in Europe, 
the genus was divided into more natural assemblages which were considered to merit 
generic status. Since a number of species of the Laemophloeinae are of economic 
importance, it was considered justifiable to extend the study of the group to those 
species not occurring in Europe. This paper is the first of such studies. 



ENTOM. 12, 



i68 L. P. LEFKOVITCH 

The Species considered here are from the Ethiopian region including Madagascar 
(some species from Madagascar belong to a fauna whose affinities are with the 
Oriental or Pacific regions) and those apparently native to Madeira, the Canaries and 
the Salvages ; those North African species which are primarily European and 
represented in Mediterranean Africa but are here excluded are Lathropus sepicola 
(Miiller), Laemophloeus nigricollis Lucas, L. monilis (Fabricius), L. muticus (Fabri- 
cius), Notolaemus castaneus (Erichson), N. unifascicUus (Latreille), Cryptolestes 
duplicatus (Waltl), C . fractipennis (Motschulsky), C. corticinus (Erichson), Placonotus 
testaceus (Fabricius), Leptophloeus alternans (Erichson), L. juniperi (Grouvelle), 
L. hypohori (Perris) and L. clematidis (Erichson) ; reference should be made to 
Lefkovitch (19596) for a discussion of these. 

There is one African species Laemophloeus notabilis Grouvelle, 1904 : 183, whose 
type specimens I have been unable to see and whose description is inadequate for 
its generic position to be established. It should be regarded as a species incertae 
sedis for the present. So far as I know, all other previously described species have 
been included. 

The most interesting point that has arisen from the study of the African and 
European species of the subfamily (Lefkovitch, 1959&) is the difficulty in finding 
adult characters which would enable a tribal grouping of the genera. Quite clearly, 
Cucujinus Arrow, by its extreme flattening, the grooved prothoracic sublateral lines, 
the large contribution made by the metepisternum to the mesocoxal cavity and the 
curved suture between that sclerite and the metasternum, is distinct from the other 
genera. This degree of difference is such that the lack of apparent comparable 
differences between the other genera suggests that the appropriate phylogenetic 
characters are yet to be discovered. The recognition of these characters may be 
easier after a study of the Laemophloeinae in the remainder of the world. No tribal 
grouping of the genera will therefore be attempted at this stage. 

Data relating to specimens consisting of " Miss. H. de Saeger " followed by a 
number are given in the appendix ; further details are given by de Saeger (1956). 
The specimens were collected in Garamba in the Congo. 

GENERAL STRUCTURE OF ADULT LAEMOPHLOEINAE 

Head. The head is perhaps the most obviously variable part of the body. 
The lateral lines, which when present (they are absent in Lathropus) have much 
the same structure as those of the pronotum, may often be joined anteriorly by a 
transverse line. In some genera, this transverse line is in fact a groove, which to- 
gether with the median impressed line suggests the primitive Y-shaped epicranial 
sutures, e.g. Text-fig. 5. These sutures are not homologous with the ecdysial lines 
of immature stages of Coleoptera and the transverse suture is probably between 
the ante- and postclypeus and not between the frons and the clypeus. In some 
other genera the transverse fine is raised and is obviously no more than a transverse 
continuation of the lateral lines. In some of these genera, examination by trans- 
mitted light reveals a transverse suture in the head which does not coincide with 
the transverse line. In the broadest terms, the shape of the head varies from 



A REVISION OF AFRICAN LAEMOPHLOEIN AE 169 

truncated triangular to rectangular and can even be rostrate as in Rhinolaemus, 
Rhinophloeus and Rhinomalus. In general the gular sutures are visible in all the 
genera and are never confluent even in those genera (e.g. Magnoleptus gen. n.) 
which recall the Passandridae in their general appearance. The anterior angles of 
the genae are variable in the degree of development ; in most genera they present no 
unusual features but in those genera in which males have a lateral expansion exter- 
nally upon the mandibles, the genae are often enlarged proportionally. In males of 
Passandrophloeus the degree of development is such that mandibular development 
is quite clearly linked with that of the genae, which appear to prevent the mandibles 
from crossing too far in the median Une (Text-fig. 39). Antennae vary to some 
extent in the subfamily ; the scape, pedicel and three-segmented club may be con- 
siderably more robust than the intermediate segments (e.g. Lathropus) ; the degree 
of size differentiation is almost imperceptible in some genera ; the antennal seg- 
ments may even be quite elongate and a club completely indistinguishable. Sexual 
dimorphism may be apparent in the structure of the antenna. In males of many 
species in many genera, the antennal segments are elongate and there is no apparent 
club (e.g. Text-fig. 54), whereas those of females are shorter and a club is often quite 
distinct (e.g. Text-fig. 64). In addition, the male antennae may be as long as, or 
longer than the rest of the body whereas those of females of the same species are 
often equal to no more than half the body length. In some species the scape of the 
male is developed into an unusual form (Lefkovitch, 19586) ; two species with such 
developments occur in the African region. 

MouTHPARTS. Laemophloeinae may be distinguished from other Cucujidae in 
that the lacinia is completely devoid of setae. As noted above, the mandibles may 
be produced laterally, resembling the condition in males of some species of Uleiota. 

In this paper, " front of the head " refers to the dorsal margin between the 
antennal insertions, " epistome " to that part of the margin above the mouthparts. 

Pronotum. In general, the pronotum is broader than long, although in Lepto- 
phloeus and Narthecius the reverse may hold. Normally the anterior angles are 
distinct but are less prominent than the posterior angles except in Cannes and some 
Placonotus. Each lateral line (stria of earher authors) may consist of a simple 
raised ridge (e.g. Text-fig. 20) or it may be a double structure consisting of 
a raised ridge and an internal groove, the whole appearing as a single structure 
(e.g. Text-fig. 24). In some genera the groove may deepen behind the mid- 
point and form a deep fovea (e.g. Text-fig. 5). A line of punctures may be 
situated adjacent to the ridge externally, their outer margins sometimes running 
together to form what appears to be a second lateral fine (e.g. Text-fig. 40). When 
such a condition holds, it is usually easy to decide which of the two lateral Hnes is 
the primary since the outer margin of the raised part of the primary line is continuous 
with the line adjacent to the anterior and posterior margins of the sclerite. In 
Cucujinus the lateral line is simply an elongate groove without any distinct edges, 
disappearing anteriorly and posteriorly before reaching the edge of the sclerite 
(Text-fig. 55). 

Elytra. Punctured striae are typical and are probably always present. 
However, superimposed upon this puncturation there is in many genera a system 



170 



L. P. LEFKOVITCH 



of longitudinal raised areas. The position of these raised areas is quite constant. 
They occur in the first, third, fifth and seventh intervals of each elytron, but there 
is some variation between genera, and between the species in some genera, in the 
degree of elevation and in the lateral width. The regions enclosed by these raised 
areas, which are continuous both apically and basally, are called a cell for the 
purposes of this paper. The longitudinal raised region between the cells is here 
called a secondary interval. The secondary intervals may be quite narrow as in 
Microlaemus (Text-fig. 24) in which case the primary strial interval may then be 
considered to be carinate or just elevated ; or they may be broad as in M agnoUptus , 
where an accurate brief description of the primary interval may be difficult to give 
(Text-fig. 27) . It is quite clear that each cell contains two rows of punctured striae 
and that any rows of punctures situated on a secondary interval do not represent 
striae (Text-fig. i). Secondary rows of punctures may be present in the cells in 
addition to the striae, e.g. in Cryptolestes capensis which has one additional row and in 
C. ferrugtneus which has two. A brief examination will often suffice to distinguish 
which rows of punctures are striae and which are secondary. 

Ventral side of the thorax. The pro-episternal region is quite distinct and 
the suture between this sclerite and the prosternum often reaches the lateral margin 
of the pro thorax. The pro-epimeron is often exposed and may close the anterior 
coxal cavity in conjunction with the intercoxal process of the prosternum (Text-fig. 
38). The amount of closure of the anterior cavities is variable ; the cavities may 
be open posteriorly (Text-fig. 19), they may be closed by a meeting of the prosternum 
and epimeron either as points (Text-fig. 13) or in a broad front (Text-fig. 38). The 
prosternal process may be either very broad (Text-fig. 56) or very narrow (Text-fig. 
21), It may be straight posteriorly, convex or concave. These features may be of 




Fig. I. Diagrammatic dorsal view of right elytron of typical Laemophloeinae. 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 171 

value in a tribal classification, since they must be of importance in the kind of move- 
ment that the animal is able to make and thus of significance in the evolution of the 
group. There is some variation in the construction of the ventral side of the meso- 
thorax in different genera. In most genera both the mesepisternum and the 
mesepimeron form part of the mesocoxal cavity, or if the mesepimeron does not, 
then it only just fails to do so. In many genera the metepisternum either does not 
contribute to the mesocoxal cavity (Text-fig. 15) or only just does so, but in Cucujinus 
the contribution of this sclerite is such that it forms an important part of the 
posterolateral margin of the cavity (Text-fig. 56). 

The metasternum presents no special features except in the degree of definition 
of the median suture, and in the shape of that region in contact with the intercoxal 
process of the first visible abdominal sternite. 

Ventral side of the abdomen. The intercoxal process of the first visible 
abdominal sternite has been considered by Lefkovitch (19596) ; nothing further 
has been revealed by the African species of the subfamily. There is considerable 
variation in the overall shape of the five sternites which are visible, but it seems 
likely that this variation represents generic and not tribal differences. In general, 
either the first four visible sternites are as broad as each other or they become 
progressively narrower, the fifth sternite always being the narrowest. The first 
sternite is longest although the fifth sternite may be nearly as long as the first in 
some genera. 

Legs. The legs show a certain amount of variation although much of this is 
often specific in character. The trochantero-femoral junctions are usually of the 
normal type, but occasionally an approach to both the heteromeroid and the 
elongate type may be seen. The femora are often dilated, and, in very flattened 
species, the dilation may be considerable in one plane. The tibiae are usually 
slightly sinuate and bear two spines apically. In the peculiar Laemophloeus mirificus 
Grouvelle, these two spines are produced into setae which are equal in length to the 
combined lengths of the tarsal segments. The tarsi are usually five-segmented 
but may be four-segmented in the hind legs only, as in the males of many species. 
The basal four segments together are usually slightly longer than the terminal 
segment, but may be much shorter as in Lathropus Erichson, where the basal 
segments are shortened, or as in Laemophloeus gestroi Grouvelle where the terminal 
segments are considerably elongate in comparison with the usual condition. The 
terminal segment is never short. There is at most the very smallest indication of 
lobing of the segments ventrally. 

(The generic attribution of mirificus and gestroi has not yet been established.) 

Key to Genera 

1 . Frontoclypeal suture situated at the base of a transverse groove .... 2 
- Frontoclypeal suture either not situated at the base of a transverse groove or obsolete 4 

2. Intercoxal process of the first abdominal sternite pointed anteriorly ; anterior 

angles of the pronotum never produced into an acute tooth 

LAEMOPHLOEUS Dejeani 

* This genus is included in the key since although it is not represented in the Ethiopean region proper, 
it is represented in Mediterranean Africa : supra p. 168. 



172 L. P. LEFKOVITCH 

- Intercoxal process of the first abdominal sternite straight or slightly rounded 

anteriorly ; anterior angles of the pronotuna often produced into an acute tooth 3 

3. Elytra always covering the whole abdomen ; last visible abdominal sternite in the 

mid-line at most only slightly longer than the preceding sternite ; pronotum 
1-3 times as broad as long ; right mandible of male produced laterally and 
dorsally I. CANNES gen. n. (p. 174) 

- Elytra leaving at least the tip of the abdomen exposed (Text-fig. 3), often the last 

segment is completely exposed dorsally ; last visible abdominal sternite usually 
distinctly longer than the preceding sternite, often twice as long ; pronotum 
usually quadrate or very slightly transverse ; mandibles of the male without any 
lateral production II. PL>lCOiVOTl75 Macleay (p. 176) 

4. Head strongly produced anteriorly to the antennal insertions forming a distinct 

rostrum, this region being broad and flattened ; pronotal lines strongly carinate ; 
anterior coxal cavities widely open posteriorly ; coxae very close together on 
each segment, separated by a distance equal to less than the width of a coxa ; 
intercoxal process of the first visible abdominal sternite pointed anteriorly ; 
antennae very short, hardly reaching the mid-point of the pronotum 

Laemophloeus raffrayi Grouvelle^ 

- Head only normally developed anteriorly to the antennal insertions or not produced 

at all ; pronotal lines only weakly carinate ; anterior coxal cavities either broadly 
or narrowly open, or closed posteriorly ; coxae separated by a distance equal to 
or greater than the width of a coxa ; intercoxal process of the first visible abdomi- 
nal sternite never pointed anteriorly ; antennae usually reaching at least the 
mid-point of the pronotum .......... 5 

5. Lateral lines on the head joined transversely just behind and very close to but 

distinct from the epistome by an almost straight but sometimes irregular line, 
this line possibly representing the frontoclypeal suture ..... 6 

Lateral lines of head, if joined by a line, the line is not straight but clearly curved 

(or the line is the line of the epistome) ....... 7 

6. Labrum truncate or shallowly emarginate anteriorly ; antennae usually inserted at 

the front corners of the head ; eyes usually occupying slightly more than one-third 
of the lateral margin of the head measured from the front angles to the base 

III. XYLOLESTES gen. n. (p. 187) 

- Labrum evenly rounded anteriorly ; antennae usually inserted laterally on the 

head, behind the front comers ; eyes usually occupying about one-quarter of the 
lateral margin of the head . . . .IV. XYLOPHLOEUS gen. n. (p. 189) 

7. Lateral lines of pronotum never reaching its base (Text-fig. 55) ; head expanding 

laterally from base to just behind eyes, at widest usually broader than the prono- 
tum ; elytra leaving at least the last and often the preceding segments exposed 

XV. CUCUJINUS Arrow (p. 228) 

- Lateral lines of the pronotum always reaching its base ; head parallel-sided from 

base to eyes, at most as broad as the pronotum ; elytra never leaving more than 

the tip of the pygidium exposed ......... 8 

8. Prothorax with suture between pronotum and pro-episternum visible dorsally ; 

about 4 mm. in length and Carabid-like in general appearance ; elytra with very 
distinct punctate striae (Text-fig. 20) . . . V. BLUBOSgen.n. (p. 194) 

Prothorax with suture between pronotum and pro-episternum not visible dorsally 
or if so, beetles cylindrical, resembling some Colydiidae and about 2 mm. in 
length ; elytra with less distinct punctate striae ...... 9 

9. Epistome either rounded, truncate, or with a single medial spine anteriorly or with 

two teeth (Text-figs. 28-30) or, if with one emargination, either the mesepimera 

^ This species does not belong to the Cucujidae and a detailed study has failed to show its proper 
family position. It is included here since it was originally described as a Laemophloeus and could 
possibly be misidentified as a Laemophloeine (see Lefkovitch, 1962). 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 173 

contribute to the mesocoxal cavities or the body is never longer than 3 mm. and 

the clypeus is rarely produced into horns in males . . . . . . 10 

Epistome with three or five emarginations or, if with one emargination, either the 
mesepimera do not contribute to the mesocoxal cavities or the clypeus is always 
produced into horns in the males (Text-fig. 54) and the body is usually 3 mm. or 
more in length ............ 15 

10. Frontoclypeal suture represented by a line strongly curved away from the mouth- 

parts (Text-fig. 24) (in the smallest species, this line is not easily seen) ; a dark 
mark is present anteriorly on the lateral lines of the head at the junction between 
the lines and the frontoclypeal suture ; first visible abdominal sternite 2-5 times 
as long as the second sternite in the mid-line ; last visible abdominal sternite 
1-5 times as long as the preceding sternite . VI. MICROLAEMUS ^en. n. (p. 196) 

— Frontoclypeal suture obsolete ; no dark marks anteriorly on the lateral lines of the 

head ; first visible abdominal sternite usually only twice as long as the second 
sternite in the mid-line ; last visible abdominal sternite only rarely longer than 
the preceding sternite .......... 11 

1 1 . Body almost cylindrical ; antennae less than half the body length in both sexes ; 

head and pronotum never with two carinae on each side . . . . . 12 

— Body more flattened in section ; antennae in female just less than half the body 

length or longer, in male often considerably longer than those of the female, often 
either as long as the body, or head and pronotum with two lines on each side, or 
these characters combined . . . . . . . . . . 14 

12. Head longitudinally grooved on each side of a median epistomal horn 

IX. NARTHECIUS Leconte (p. 210) 

— Head, if with a median epistomal horn, not longitudinally grooved on each side of it 13 

13. Larger beetles, 3'5-5*o mm. in length ; male tarsal formula 5-5-4 ; body usually 

black or reddish black in colour . . . VII. MAGNOLEPTUS gen. n. (p. 199) 

— Smaller beetles, i •8-3-0 mm. in length ; tarsal formula in both sexes 5-5-5 (except 

in two species where it is 5-5-4) ; body usually brown in colour 

VIII. LEPTOPHLOEUS Csisey (p. 201) 

14. Anterior coxal cavities with broad transverse extensions laterally (Text-fig. 38) ; 

head with two lateral lines on each side ; mandibles of male produced laterally 
and ventrally (Text-fig. 39) ; genal horns strongly developed, especially in males 

X. PASSANDROPHLOEUS Kessel (p. 213) 

— Anterior coxal cavities with narrow lateral transverse extensions ; head rarely with 

two lateral lines on each side, usually with one ; mandibles of the males of some 
species produced laterally but never ventrally ; genal horns weakly developed 

XVI. CRyPTOLE5TE5 Ganglbauer (p. 231) 

15. Front of head with a deep emargination just internal to the antennae (Text-fig. 43) ; 

five distinct emarginations exist between the insertions of the antennae 

XI. MESTOLAEMUS gen. n. (p. 215) 

— Front of head only shallowly emarginate just internal to the antennae ; only one 

distinct emargination exists anteriorly on the head, that over the mouthparts ; 
very shallow more lateral emarginations may be present in larger species, but a 
deep emargination just medial to the antennae is never present . . . 16 

16. Intercoxal process of the first visible abdominal sternite evenly rounded anteriorly 

and without a straighter median section ; frons produced anteriorly into two 
horns in males (in one species only the left horn is developed), these horns appar- 
ently being formed by the production of the external angles of the single epistomal 
emargination (Text-fig. 54) ; elytra carinate laterally, with secondary intervals 
well developed and continuing to the apices ; the secondary intervals may be as 
broad as the cells or narrower ; beetles larger, 2-3-3-5 mm. in length (associated 
with Legume pods) .... XIV. PLANOLESTES Lefkovitch (p. 226) 



174 L- P- LEFKOVITCH 

— Intercoxal process of the first visible abdominal sternite rounded and with a straighter 

median section ; frons not produced anteriorly into horns in either sex ; elytra 
carinate or non-carinate laterally, if with raised secondary intervals then either 
the elytra become flat apically or the secondary intervals are always as broad as the 
cells ; beetles usually smaller, i •5-2-5 mm. in length, if larger, then usually 
three emarginations are present anteriorly on the head . . . . . 17 

17. Posterior border of intercoxal process of prosternum concave for reception of 
mesosternum (Text-fig. 44) ; elytra not carinate laterally ; beetles more convex in 
section and body rather short ; elytra with only one raised interval, recurved 
near the humeral angle and with faint punctured striae ; scutellum semi-elliptical ; 
internal angle of metepisternum not contributing to the mesocoxal cavity, i.e. 
mesepimeron and metasternum in contact with each other 

XII. MARIOLAEMUS gen. n. (p. 216) 

— Posterior border of intercoxal process of prosternum evenly rounded (Text-fig. 53) ; 

elytra carinate laterally ; beetles flattened in section and body longer ; elytra 
with at least two raised intervals in addition to the punctured striae ; scutellum 
triangular ; metepisternum contributing to the mesocoxal cavity, i.e. mesepimeron 
separated from metasternum by the anterior angle of the metepisternum 

XIII. NOTOLAEMUS Lefkovitch (p. 220 ) 



DESCRIPTIONS OF GENERA AND SPECIES 

I. CANNES gen. n. 

Type species : Cannes immoderatus sp. n. 

This genus is most closely related to Placonotus Macleay, It is distinguished 
from that genus by the characters given in the key (p. 172). 

About 2 mm.; body flattened. 

Head transverse ; eyes large, usually in contact with prothorax ; frontoclypeal suture distinct 
and situated at the base of a groove ; epistome truncate ; lateral line distinct and forming part 
of the dorsal margin of eye ; antennae about as long as body in male, shorter in female, in female 
there is a loose three-segmented club ; males with right mandibles produced laterally and dorsally. 

Pronotum distinctly transverse, narrowing somewhat basally ; anterior and posterior angles 
acute but hardly, if at all, produced ; lateral line distinctly raised and with an accompanying 
depression, the line not punctured or foveolate along its length. 

Scutellum semicircular ; elytra carinate laterally, with the third cell complete (see p. 170) and 
two other raised intervals. 

Genae normal ; gular sutures obsolete ; anterior coxal cavities closed posteriorly, circular and 
widely separated ; intercoxal process of the prosternum slightly convex posteriorly ; mesepimera 
not contributing to the mesocoxal cavities (Text-fig. 2) ; metasternum equal in length to the 
first two abdominal segments ; intercoxal process of first abdominal segment almost straight 
anteriorly ; first visible abdominal sternite 1-5 times as long as the second sternite, second, 
third and fourth sternites about equal to each other, fifth slightly longer ; legs subequal ; 
trochantero-femoral junctions of normal type ; tarsal formula 5-5-5 in female, 5-5-4 in male. 

Key to Species 

I. Head with punctures very slightly less in diameter than facets of eyes, separated 
by a polygonal micro-reticulation ; discal setae about 2 -5 times as long as diameter 
of a puncture ; fifth antennal segment in males longer than the scape 

2. ambiguus (Grouvelle) (p. 175) 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 175 

- Head with punctures much less than an eye facet in diameter (about 0-25 times), 
separated mainly by smooth cuticle but with small areas of irregular micro- 
reticulation ; setae about 5 times as long as the diameter of a puncture ; fifth 
antennal segment in males shorter than the scape . i. immoderatus sp. n. (p. 175) 

I. Cannes immoderatus sp. n. 

(Text-fig. 3) 

I '9-2 '3 mm.; body reddish yellow to blackish red in colour ; elytra somewhat more yellow 
than the head and pronotum. 

Head across eyes nearly as wide as pronotum at its widest ; frontoclypeal suture evenly 
curved posteriorly ; epistome rounded anteriorly ; antennae as long as body in male, shorter 
in female ; last three segments somewhat enlarged ; head simply punctured, with a micro- 
reticulation, punctures separated by one to two diameters, each subtending a seta equal to 
five punctures in length ; pronotum with puncturation and setation as on the head ; elytra some- 
what oval, broadest about a third of their length from the base and narrowing apically, evenly 
rounded to apico-sutural angle. 

Congo, Uganda. 

HoLOTYPE male " Yangambi 1953. C. Donis z.2634 (= Synsepalum dulciferum 
Sch. and Th, : Sapotaceae) coll. R. Majme. Com. et Bois Congo. R2439 " (Mus. R. 
Congo Beige) ; i paratype with same data as holotype except with z.2992 and R2440 ; 

1 paratype with same data as holotype but with z.2672 (Mus. R. Congo Beige) ; 

2 paratypes "Uganda Kampala 25.vii.1927 H. Hargreaves " (British Museum 
(Natural History)). 

2. Cannes ambiguus (Grouvelle) comb. n. 

Laemophloeus ambiguus Grouvelle, 1923. 

This species differs from G. immoderatus sp. n. in that the setae on the head and 
pronotum are considerably shorter, the elytra are less narrowed apically, and the 
epistome is truncate whereas that of G. immoderatus is rounded anteriorly. 

I •8-2-3 mm.; body yellow to blackish red in colour. 

Head across eyes as wide as or slightly narrower than pronotum ; frontoclypeal suture evenly 
curved posteriorly ; epistome truncate anteriorly ; antennae as long as the body in male and 
the last three segments not forming a club, in female antennae shorter than body and last 
three segments forming a loose club. Head evenly punctured, punctures separated by one 
diameter, micro-reticulated between the punctures ; setae equal in length to 2-5 times diameter 
of one puncture. Pronotum, with puncturation and setation as on head. Elytra less narrowed 
apically than in G. immoderatus sp. n., evenly rounded to apico-sutural angle. 

East Africa, Congo. 

Holotype female " Afrique orient, anglaise Foret de Nairobi Alluaud et Jeannel 
Nov.-Dec. 1911 — 1700 — St II " (Grouvelle coll.). Other specimens seen include : 
I " Benito. Congo Franc, sp. nv." ; i " Ogoove Lambarene R. Ellenberger 1911 " 
I " Af. or. All. Amani/Eichelbaum/La^wo^A/o^MS testaceus F var A ined " (Grouvelle 
collection, Museum National d'Histoire Naturelle, Paris) : i " Congo Beige M2 
Kahuzi S7 iii. 22-viii-i952. Dr. Schedl. s.708 " ; i " Recolte dans I'humus/ 
I.R.S.A.C. — Mus. Congo Kivu : Terr. Uvira, Mulenga 1880-2010 m. (Vest. for. 



176 



L. P. LEFKOVITCH 



ombroph.) V-1951. N. Leleup." ; i " S/L'ecorce de Canarium Schweinfurthij Coll. 
Mus. Congo. Bas-Congo : Sanda. Courtois." ; i " Eala vii-1935. J. Ghesquiere " ; 
I " A la lumiere I.R.S.A.C. — Mus. Congo. Kivu : Terr. Masisi, 720 m Walikale (River 
Loa) xi-53. N. Leleup " ; i " Elisabeth ville -ii-1940 H. J. Brehdo " ; 2 " Kivu : 
Buranga 5-xii-i925 and 9-xii-i925. Dr. H. Schouteden " ; 3 " Yangambi, 1952. 
C. Donis. Z3032 [Synsepalum dulciferum Sch. and Th. : Sapotaceae) " ; Miss. H. 
de Saeger, 3 (3649), i (3519). 2 (3736), i (3738), i (2601). 2 (3379) (see Appendix) 
(Mus. R, Congo Beige) ; i " Uganda. Ruwenzori Range. Mahoma River 6,700 ft. 
13-16. viii. 1952. D. S. Fletcher " ; i " R. E. Dent. Charia R.7800. July, 1929 " ; 
I " Ngong Forest in Boletus. Nairobi B.E. Africa 4.x. 1920. A. F. J. Gedye " ; 
4 " Port St. John. Pondoland Sept. 1923. R. E. Turner " ; 3 " Belgian Congo 18 m. 
sw of Elizabethville 13. xi, 1927. Dr. H. S. Evans"; 2 "Nairobi .xi.1920"; 
(British Museum (Natural History)) ; 2 " Mt. Elgon 0(st) A(frika) 2200 m. iv-25 
leg Alinder" (Zoologische Sammlung des Bayerischen Staates, Munich). 



II. PL/lCONOTt/5 Macleay 

(Text-fig. 5) 
Placonotus Macleay, 1871. 
Silvanophloeus Sharp, 1899. 

TjTpe species : Placonotus longicornis Macleay (by monotypy). 






Figs. 2-5. (2) Front and middle coxal cavities of Cannes immoderatus ; (3) dorsal view 
of G. immoderatus ; (4) front and middle coxal cavities of Placonotus testaceus (F.) ; 
(5) dorsal view of P. testaceus. 

Scale lines : Figs. 2, 4 = 0-25 mm. ; Figs. 3, 5 = i nam. 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 177 

A redescription of this genus is given by Lefkovitch (19596). In addition to the characters 
indicated there, the following should be noted : Lateral line on head formed by raised ridge ; 
that of pronotum by a groove with outer margin slightly raised to form a weak ridge ; genae 
normally developed ; gular sutures obsolete ; hind border of prosternal process convex poster- 
iorly ; anterior coxal cavities circular ; metepisternum not contributing to mesocoxal cavity but 
both mesepislernum and mesepimeron do ; posterior border of mesosternum straight ; trochantero- 
femoral junctions of normal type. 

The structure of the male genitalia presents points of interest in this genus. Two pairs of 
claspers are present which appear to be derived from the sclerites of the tenth abdominal segment. 
The tergite may be partially or completely divided along the median line to form two elongate 
rods ; the sternite is always divided and articulated at its proximal end. Internally, the two 
sclerites may be tuberculate posteriorly. The protrusion of these claspers gives the impression 
that they are ovipositors ; several specimens labelled by their collectors as females were found 
to be males with projecting claspers. 

Key to African Species 

1. Anterior angles of pronotum acute (Text-fig. 6) ...... 2 

- Anterior angles of pronotum rounded (Text-fig. 7) . . . . . . 12 

2. Black or reddish black in colour, without a lighter mark on each elytron ; epistome 

truncate anteriorly ....... i. tastus sp. n. (p. 178) 

- Head and pronotum brown or yellowish in colour, or if black, then each elytron 

with a lighter mark at basal third ; epistome emarginate anteriorly ... 3 

3. Elytra unicolorous ............ 4 

- Elytra with a lighter mark at basal third . . . . . . . 11 

4. Smaller species (about 1-5-2 mm.) ; pronotum distinctly transverse and head punc- 

tate ; elytra 2 •5-3-0 times as long as their combined width .... 5 

- Larger species (about 1-8-2-3 nim.) ; pronotum about as broad as long or if trans- 

verse, then head impunctate ; elytra usually less than 2-5 times as long as their 
combined width ........... 6 



. 2. dolce sp. n. (p. 178) 

3. mestus sp. n. (p. 179) 

13. mossus sp. n. (p. 187) 

7 



5. Epistome concave ..... 

- Epistome convex ..... 
[ 6. Disc of head impunctate but strongly reticulate 

- Disc of head punctured .... 

7. Disc of head polygonally reticulate between punctures 

8. donacioides (Wollaston) (p. 183) 

- Disc of head smooth or with irregular groove-like markings between punctures . 8 

8. Elytra rounded together posteriorly (Text-fig. 8) . . 4. majus sp. n. (p. 179) 

- Elytra separately rounded to apico-sutural angle, semitruncate (Text-fig. 9) . . 9 

9. Ventral claspers of male weakly developed (Text-fig. 10) 6. ealaensis^ sp. n. (p. 182) 

- Ventral claspers of male strongly developed . . . . . . . 10 

10. Ventral claspers straight posteriorly ; dorsal claspers incompletely divided, in shape 

together equivalent to a normal tenth tergite (Text-fig. 11) 

5. subtruncatus^ sp. n. (p. 181) 

— Ventral claspers spatulate posteriorly ; dorsal claspers completely divided except 

for a narrow strut anteriorly, posteriorly no broader than ventral claspers (Text- 
fig. 12) ....... . 7. af r icanus^ sp. n. (p. 182) 

11. Head and pronotum reddish brown . . . .12. exornatus (Grouvelle) (p. 186) 

— Head and pronotum black ..... 11. dccoram* (Grouvelle) (p. 186) 

12. First visible abdominal sternite with femoral lines ; pronotum convex between 

lateral lines ; anterior angles of pronotum not produced (Text-fig. 7) 

9. politissimus (Wollaston) (p. 183) 

^ Good characters for the separation of females have not been discovered. 



178 L. P. LEFKOVITCH 

— First visible abdominal sternite without femoral lines ; pronotum almost flat 
between the lateral lines ; anterior angles of pronotum somewhat produced 

10. bolivari (Grouvelle) (p. 185) 

I. Placonotus tastus sp. n. 

(Text-fig. 6) 

2 mm. ; body reddish black in colour, elytra less black than the head and pronotum ; shining 
but not brilliantly. 

Head somewhat transverse, eyes large slightly in advance of base ; frontoclypeal suture with 
a raised ridge anterior to the shallow groove ; epistome truncate anteriorly ; antennae as long 
as body in male ; disc evenly punctured, punctures separated by a distance equal to two to 
three diameters ; each puncture subtends a seta equal to about one diameter in length ; laterally 
on disc the intervals between punctures micro-reticulate, medially less so ; frons punctured 
just anterior to frontoclypeal suture but only reticulate just behind epistome. 

Pronotum very slightly transverse and slightly narrowed basally, at its broadest slightly 
narrower than head ; anterior angles acute and produced, posterior angles obtuse ; disc with 
a small shallow depression on each side just anterior to the fovea included in lateral line ; disc 
with puncturation and setation as on disc of head ; region between lateral line and lateral margin 
with more distinct micro-reticulation than on disc. 

Elytra slightly broader across shoulders than pronotum at broadest ; 2*5 times as long as 
their combined width ; parallel-sided anteriorly but narrowing somewhat towards apices from 
about mid-point ; third cell developed basally, with part of first cell developed at about mid- 
point ; punctate striae present and complete until just before the apical region where they are 
obsolete ; each elytron subtruncate apically, their combined median regions being straight ; 
surface with micro-reticulation between punctures and lines. 

Female unknown. 

Ethiopia. 

HoLOTYPE male "Abyssinia: Djem-Djem Forest, circa 8000 ft. 25-29. ix. 1926. 
Dr. H. Scott." (British Museum (Natural History)). 

This species is most clearly distinguished from other Placonotus in its colour 
which is a very dark red or a slightly reddish black. In addition, the truncate 
epistome is unique amongst the African species of the genus having acute anterior 
angles to the pronotum. 

2. Placonotus dolce sp. n. 

This species can be distinguished from other Placonotus in that it is quite small, 
the pronotum transverse, the ratio of length of elytra to their combined breadth is 
between 2-5 and 3 and has interesting double puncturing on the discs of both the 
head and the thorax. 

i'5-2*o mm. ; body brownish red in colour. 

Head transverse ; eyes about half a diameter in advance of base of head ; frontoclypeal suture 
with a slight raised ridge anterior to deep groove ; epistome shallowly emarginate anteriorly ; 
antennae about 0-75 times length of the body in m,ale, shorter in female ; antennae oi female 
with last three segments slightly enlarged to form a loose club, in male these segments more 
elongate than those preceding but not broadened laterally ; disc of head with punctures of 
two sizes, the larger with diameter equal to slightly more than that of eye facet, the smaller 
being equal to half or less than this diameter ; the larger punctures separated by a distance 



A REVISION OF AFRICAN L AEMOPHLOEI N AE 



179 



equal to one to two diameters, each subtending a seta equal in length to two diameters ; the 
smaller punctures irregularly and sparsely scattered between larger ; head without micro- 
reticulation between punctures but smooth and shining. 

Pronotum 1-25 times as wide as long, at its broadest slightly broader than head ; anterior 
angles acute and slightly produced, posterior angles acute ; lateral margin with twelve small 
teeth, those anterior being further separated from each other than those posterior ; puncturation 
and setation similar to that of head ; fovea of lateral line deep. 

Elytra at shoulders slightly broader than pronotum ; 2'5-3-o times as long as their combined 
width, hardly narrowing apically ; not or only weakly carinate laterally ; with three cells 
weakly developed but hardly visible apically and basally ; striae distinct ; each elytron separ- 
ately rounded to apico-sutural angle ; without micro-reticulation between punctures. 

Congo. 

HoLOTYPE male " Dans terreau, au Berlese/Kivu : Terr, Lubero, confl. Munobo 
et Lubero, 1880 m. R, P. M. J. Cells ix-1954 " and allotype with the same data 
(Mus. R. Congo Beige). 

3. Placonotus mestus sp. n. 

This species resembles P. dolce sp. n., differing as follows : head and pronotum with punctures 
separated by once to twice their diameter, without micro-puncturation, with micro-reticulation 
between punctures at least peripherally on disc ; epistome convex anteriorly ; eyes situated 
very close to thorax ; elytra no broader across shoulders than pronotum, markedly narrowing 
apically, clearly carinate laterally. 

Sierra Leone, Congo. 

HoLOTYPE male and 15 paratypes " Bas-Uele : Kotell i/2i-i-ig25 Dr. H. 
Schouteden ". Other paratypes : i " Mongende i3-iv-i92i Dr. H. Schouteden " ; 
3 " Eala viii-1935 J. Ghesquiere " ; i " Katanga : Katompe i/i5-vi-i930 Dr. P. 
Gerard " ; i " Recolte sous ecorce d'Hevea Yangambi v-1952 J. DeceUe " ; i 
" Luki 5-xi-i952 R. Mayne R.M.i." ; 3 " Yangambi 1951 C. Donis z.3-17, z.802 
and Z.1368 " (Mus. R. Congo Beige) : 2 " Sierra Leone Njala 4-1-1933 E. Hargreaves. 
Under bark of dead rubber " (British Museum (Natural History)). 

4. Placonotus majus sp. n. 

(Text-fig. 8) 

This species resembles Placonotus tastus sp. n, but differs in that its body colour is 
less dark, in the possession of the small longitudinal impressions between the 




o ^ 



6 7 8 9 -^ 

Figs. 6-g. (6) Pronotum of Placonotus tastus and (7) of P. politissimus ; (8) elytral 
apices of P. majus and (9) of P. subtruncatus. 

Scale line — 0-25 mm. 



l8o L. P. LEFKOVITCH 

punctures of the disc of the head and prothorax, in the greater narrowing of the 
pronotum basally in addition to the emarginate epistome. 

2'(>-2'5 mm. ; body ferrugineous in colour. 

Head transverse ; eyes about half a diameter in advance of base ; frontoclypeal suture with 
an anterior ridge and a shallow groove ; epistome emarginate anteriorly ; antennae about 
0'75 times body length in males, shorter in females ; disc of head punctured, punctures smaller 
than facets of eyes, separated by one to three diameters, each with a short seta equal in length to 
one and a half diameters ; surface between punctures irregular, with a tendency to groove- like 
impressions but otherwise shining. 

Pronotum slightly broader than head, about as broad as long at its broadest, narrowed basally ; 
anterior angles acute and produced, posterior angles obtuse ; puncturation and setation on 
disc similar to that of disc of head ; surface micro-reticulate between lateral line and lateral 
margin ; the fovea included in lateral line shallow but includes a deep puncture. 

Elytra as wide as pronotum at widest, slightly more than twice as long as their combined 
width, carinate laterally, disc concave between carina and suture ; almost parallel-sided until 
just before apices ; apices evenly rounded to apico-sutural angle ; cells weakly developed ; 
striae represented by rows of setae whose subtending punctures are very shallow. 

Rhodesia, Congo, Angola, West Africa. 

HoLOTYPE and 2 paratypes " N.W. Rhodesia Shigariatombwes, 8-vii-i9i3 H. C. 
Dollman " (British Museum (Natural History)) ; paratypes as follows : 3 " N.W. 
Rhodesia Mwengwa 27° 40' E 30° S 26-vi-i9i3 H. C. Dollman " ; i " Sierra Leone " 
(British Museum (Natural History)) ; i " Angola 4194.5 — Alto Chicapa, gal. forest. 
riv. Ngungo, dans, tronc 27-vi-54. Ed. Luna de Carvalho " ; 2 " Angola 4116.20 — 
Alto Cuilo, Poste de Cacolo, rives de la Cavuemba, detr. sol, 26-V-54. A, de Barros 
Machado " ; 3 "Angola 411. 6. 17 — ^Dundo, iii-48, piege lumineux. A. de Barros 
Machado " ; i "Angola 4118.9-Gal. forest, riv. Lunguena, affl. Luangue, Poste 
Xa-Cassau, 7-vi-54. A. de Barros Machado"; 11 "Angola 10656 . 42-Marco de 
Canavezes (Cubal da Ganda), district de Benguela, piege lum ; iii-56. Ed. Luna 
de Carvalho " ; 3 " Angola 3158.24 — Dundo, xii-53, piege lum. Ed. Luna de Car- 
valho " ; 13 " Angola 4895.3 — Cazombo, Alto Zambeze, sous ecorces arbres morts, 
19-1-55. A. de Barros Machado " ; i " Angola 4323.23 — Alto Chicapa, gal. forest. 
riv. Tchirimbo, detr. sol, 9-vii-54. A. de Barros Machado " ; 2 " Angola 4117.15 
— Alto Cuilo, Poste de Cacolo, gal. forest, ruisseau Tcha-Muchito, 2-vi-54. A. de 
Barros Machado"; i "Angola 4742.14 — Rives du Lac Calundo, sous ecorces 
arbres morts, 19-1-55. A. de Barros Machado " ; 3 " Angola 4737.9 as 4742.14 ; 
4 " Angola 3176. II — Dundo, ix-53, piege lum. Ed. Luna de Carvalho " ; 2 " Angola 
4359-3 — -^Ito Chicapa, 7-viii-54. Ed. Luna de Carvalho " ; i " Angola 4370.36 — 
Alto Chicapa, gal. for. riv. Tchimboma, detr. sol, 17-1-55. A. de Barros Machado " ; 
I " Angola 4214.42 — Alto Chicapa, gal. forest, riv. Ngungo, detr. sol, 27. vi.54. A. de 
Barros Machado " ; i " Angola 3115.21 — Dundo, viii-53, piege Lum. Ed. Luna de 
Carvalho " (Museu do Dundo) ; 2 " Yangambi 1951 C. Donis 300, 302 " ; 3 " Luki 
1951 P. Henrard 97 " ; 11 " Eala vii-viii-1935 J. Ghesquiere " ; 24 " Eala iii-1935 
J. Ghesquiere " ; 3 " Eala x-1936 J. Ghesquiere " ; 2 " Kindu L. Burgeon " ; 
I " Stanleyville i9-viii-i928 A. Collart " ; i " Kamajembi (Luebo) 2i-ix-i92i Dr 
H. Schouteden " ; i " Malych 1942 Rev. P. Van Eyen " ; i " Katang : Kanzenge 
12544 N. Leleup i2-ii-i949 " ; i " Tanganyika : Moba, 780 m (a la lumiere) xi-1953 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 



i8i 



H. Bomans " (Mus. R, Congo Beige) ; 23 " IFAN 1953 Bafrechie (Mauritanie) 5-ix 
to lo-x A. Villiers " ; 13 " IFAN 1946 Mr Nimba N.E. 500-700 m. A. Villiers " ; 
2 " Irabellaben Mrs Baguezans 1200-1300 m 28-31-viii IFAN 1947 L. Chopard, 
A. Villiers " (Museum National d'Histoire NatureUe, Paris). 



The subtruncatus Species Group 

There are three species involved in this group, P. subtruncatus sp. n., P. ealaensis 
sp. n. and P. africanus sp. n. It has not been possible to separate these species on 
invariable external characters but there are no difficulties in seeing the differences 
between the male genitalia. Externally, the three species resemble P. tastus sp. n. 
but differ from it in that the body is yellowish brown in colour and in the tendency 
to longitudinal impressions on the surface of the head between the punctures. 

5. Placonotus subtruncatus sp. n. 

(Text-figs. 9, 11) 

2'0-2'5 mm.; body yellowish to reddish brown in colour. 

Head transverse ; eyes a quarter to a half times their diameter in advance of base of head, 
hemispherical in shape ; frontoclypeal sutural region with anterior shallow transverse impression 
consisting of a ridge and a deeper groove ; epistome emarginate anteriorly ; antennae about 
0'75 times body length in males, fifth to eleventh segments about twice as long as broad, in 





II 




Figs. 10-12. Male claspers of (10) Placonotus ealaensis, (11) P. subtruncatus and (12) 
P. africanus. 

Scale line = 0-25 mm. 

ENTOM. 12, 4. 13 



l82 L. P. LEFKOVITCH 

females antennae about 0'6 times body length, fifth and subsequent segments about i'5-2'0 
times as long as broad ; in both sexes, last three segments slightly dilated, more distinctly 
so in female ; disc of head punctured, the punctures slightly smaller than facets of eyes, separated 
by once to three times their diameter, each with a fine seta equal to 1-5 times this diameter in 
length ; surface between punctures polygonally reticulate but generally shining. 

Pronotum slightly broader than head (including the eyes), very slightly broader than long at 
its broadest, narrowed basally ; anterior angles acute and produced, posterior angles obtuse ; 
disc ornamented similarly to disc of head ; fovea included in lateral line encroaches upon disc, 
moderately deep and often includes a deep puncture and blackish mark. 

Elytra slightly broader than pronotum, broadest just anterior to mid-point, about 2-25 times 
as long as their combined width, abruptly deflected laterally but not carinate at fold ; each 
elytron separately and abruptly rounded to apico-sutural angle ; cells weakly developed, 
strial punctures shallow. 

Ventral claspers of male genitalia straight posteriorly with three irregular dorsal rows of small 
tubercles posteriorly, in shape resembling a coal hammer ; dorsal claspers not completely divided 
in mid-line, less strongly chitinized medially than laterally, together of shape appropriate to 
a normal tenth tergite, ventral surface with four irregular rows of tubercles posteriorly ; both 
dorsal and ventral claspers in length equal to fifth externally visible abdominal sternite. 

Congo, West Africa. 

HoLOTYPE male and 8 paratypes : " Nioka xii-53. P. Deville Pap. 49 " ; other 
paratypes as follows : 4 as holotype but dated 26-xii-53 and Pap 47, i with Pap. 46, 
I dated 28-xii-53 and Pap. 68 ; i " Katanga. Katompe i/i5-vi-i930 Dr. P. Gerard " 
(Mus. R. Congo Beige) : 2 " IFAN — 1946 Tonkoui C.I. 900-1200 m. A. Villiers 
For^t Prim. 20/30-ix " (Museum National d'Histoire Naturelle, Paris). 



6. Placonotus ealaensis sp. n. 

(Text-fig. 10) 

This species appears to be identical with P. subtruncatus sp. n. except in the struc- 
ture of the male genitalia. 

Dorsal claspers similar to those of P. subtruncatus but less evenly rounded externally ; ventral 
claspers hardly developed, being a thin rod on each side apparently fused posteriorly with the 
external angles of the dorsal claspers. 

Congo. 

Holotype and i paratype (males) : " Eala viii-1935 J. Ghesquiere " (Mus. R. 
Congo Beige). 

7. Placonotus africanus sp. n. 

As P. subtruncatus sp. n. except that the surface between the punctures is very weakly, if at 
all, polygonally reticulate and also differs in the structure of the male claspers : 

Dorsal claspers completely divided medially except, apparently, for a thin transverse strut 
anteriorly, narrowed posteriorly where no wider than ventral claspers ; tubercles on dorsal 
claspers arranged in two groups, one irregular row running forwards along the lateral margin 
from posterior region until just before anterior margin ; other tubercles about twenty in number 
situated in a group antero-medially ; ventral claspers spatulate posteriorly, situated latero- 
ventrally, bearing about twenty-five long spines internally. 



a revision of african l aemophloein ae 183 

Congo, Angola. 

HoLOTYPE male " Katanga : Kanzenze (a meme I'argile sous bois pourri) 12544 
N. Leleup i2-ii-49 " ; paratypes as follows : i " Katanga : Kansenze 22-xi-i948 
12625 N. Leleup " ; i " Recolte dans I'humus. Distr. Elisabethville Gal. for. de la 
Kisanga 26-ix-5i N. Leleup " ; 4 "Elisabethville 6-X-1912 Miss. Agric." ; i 
" S/Bois divers Kasai : Kakenge Seierie Exforka i-1945. J. Vrydagh " ; 10 " Eala 
iii-1935 J. Ghesquiere 376 " (Mus. R. Congo Beige) ; 18 " N.W. Rhodesia Shigaria- 
tombwes S-vii-igia H. C. DoUman " ; i " Angola 3176. 11 — Dundo, ix-53, piege 
lum. Ed. Luna de Carvalho " ; 2 " Angola 4117.15 — Alto Cuilo, Poste de Cacalo, 
gal. forest ruisseau Tcha-Muchito, 2-vi-54. A. de Barros Machado " ; i " Angola 
41 19. 3 — Alto Cuilo (Cacalo), vi-54. A. de Barros Machado et Ed. Luna de Carvalho "; 
I "Angola 4373.9 — Rives du Lac Calundo, sous ecorces arbres morts, 19-1-55. 
A. de Barros Machado " ; i " Angola 4895.3 — Cazombo, Alto Zambeze, sous ecorces 
arbres morts i2-ii-55, Ed. Luna de Carvalho " ; i " 18 m. S.W. of Ehzabethville 
31-X-1927 Dr. H. S. Evans " (British Museum (Natural History)). 

8. Placonotus donacioides (Wollaston) comb. n. 

Laemophloeus donacioides Wollaston, 1854. 
Laemophloeus granulatus Wollaston, 1854, syn. n. 

Head very slightly transverse in males, as broad as long in females ; epistome shallowly 
emarginate anteriorly ; lateral line represented by raised ridge and an internal groove which 
includes a weakly impressed line ; surface punctured, punctures slightly larger than the facets 
of eyes, separated by once to twice their diameter ; surface between punctures polygonally 
reticulate. 

Pronotum about as broad as long, narrowed both anteriorly and posteriorly ; anterior angles 
acute and produced, posterior angles obtuse ; lateral line and surface ornamentation as that 
of head. 

Elytra with three cells developed, secondary intervals about half the width of cells ; strial 
punctures almost invisible. 

Madeira, Canaries. 

There are 2 specimens without data labelled as types by G. Arrow in the Wollaston 
collection (British Museum). I select the male as lectotype of the species. 

There are also two syntypes without data of Laemophloeus granulatus in the same 
collection. I select a male as lectotype. This species is quite clearly just a small 
form of P. donacioides. 

9. Placonotus politissimus (Wollaston) comb. n. 

Laemophloeus politissimus Wollaston, 1867. 

Laemophloeus (Silvanophloeus) mirus Grouvelle, 1905, syn. n. 

This species, when described by Grouvelle, was considered to be very simila: to 
Laemophloeus nitens Leconte (this American species properly belongs to Placonotus 
and is now placed in that genus comb. n.). The two species were distinguished by 
the presence of femoral Hues on the first visible sternite of the abdomen of P. 
politissimus. In addition it has been found that no other species of the genus from 



i84 L. P. LEFKOVITCH 

Africa possesses femoral lines and thus this feature serves as an adequate distinguish- 
ing character. 

i-5-2*o mm.; body yellow to reddish brown in colour. 

Head transverse ; eyes large and situated close to base ; frontoclypeal suture with shallow 
groove and without raised ridge anteriorly ; epistome emarginate anteriorly ; antennae about 
three-quarters length of body in males, somewhat shorter in females, distal three segments very 
slightly dilated in males, a little more so in females ; disc of head very evenly punctured, punc- 
tures separated by about two to four diameters, each puncture with a seta about i'5 diameters 
in length ; intervals between punctures smooth and shining. 

Pronotum transverse, about 1*25 times as wide as long, slightly broader than head ; broadest 
across anterior third ; anterior angles obtuse and rounded, posterior angles rectangular to 
slightly obtuse ; puncturation and setation as on head ; fovea of the lateral line apparently 
formed by fusion of three large shallow punctures. 

Elytra broader across shoulders than pronotum at its widest, broadening slightly before mid- 
point and then narrowing towards apices, each evenly rounded to apico-sutural angle ; at their 
extreme length, between suture and external angle, elytra about twice as long as their combined 
width ; only the third cell is developed basally together with part of the first near mid-point 
and close to suture ; each elytron with inflected lateral region but not carinate ; striae with 
punctures considerably larger than those of head and pronotum. 

The species exhibits marked heterogony in the adult structure in addition to some variation 
in size and puncturation. 

Atlantic islands off Africa, Congo, Ivory Coast, Madagascar, Seychelles. 

There are eleven specimens in the Wollaston Cape Verde collection in the British 
Museum and two further specimens with a j8 written on the card standing in the 
position indicated by the species name. One of the eleven has a British Museum 
type label and a second label with " politissimus, Woll " in Wollaston's handwriting. 
In his original description Wollaston writes that the species was taken " in pro- 
fusion " ; it would seem likely that these eleven specimens are all, or at least some, 
of the syntypes. I now select the labelled specimen, a male, to be lectotype. Of 
the remaining specimens, ten are the same species ; the specimens labelled /S on 
the card are Cryptolestes capensis (Waltl). Wollaston, in discussing Laemophloeus 
clavicollis Woll. immediately after describing L. politissimus, refers to a " var. ^ 
affmis" and since L. clavicollis is C. capensis (see p. 235), I presume that these two 
specimens are misplaced in the collection and are not to be considered as syntypes of 
L. politissimus. 

Grouvelle, in his original description of Laemophloeus mirus gave the following 
type localities : " Fort-Dauphin (coll. Alluaud) ; localite incertain (coll. A. Grou- 
velle) ; St-Thome (coll. A. Grouvelle) ". In the Grouvelle collection in Paris, 
there are the following specimens of the species; one "Madagascar [mssl/Type 
[black printed on white] /Laemophloeus mirus Grouv. ty [in Grouvelle's mss]/ 
Type [black printed on red] " ; one " Madagascar (Sud) Foret Nord Ft. Dauphin 
Alluaud 1900 [printed] 17 [mss] ". In addition there are six specimens from 
Amani and one from the Cape of Good Hope which are obviously not syntypes. 
I have not located a specimen from St. Thome in the Grouvelle collection either 
named as this species or unnamed but referable to it. I consider that the two 
specimens of the species whose detailed data are given above are syntypes and 
I now select the specimen vhich bears Gruovelle's mss label as lectotype. The 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 185 

other specimen is the same species. It is quite clearly the same species as P. 
politissimus. 

The author has seen specimens of the species from the following places : Yangambi, 
195 1 (C. Donis), Lubilonga, ii.52 (M de Backer), Luki, vii.42 (R. Mayne), Luki, 
1951 (P. Henrard), Elizabeth ville, ii.1940 (H. J. Bredo), Kunungu, 9.iv.i92i (Dr. 
H. Schouteden), Eala, ix.1938 (J, Ghesquiere). Elizabethville — a la lumiere, 
xi.5o/vi.5i (Ch. Seydel), Mobeka, 16.iii.1911 (L. Burgeon) all within the Congo; 
I " Madagascar Namoroka xi.52 R. P(aulian) " ; i " Seychelle Islands Percy Sladen 
Trust Expedition/Mahe 1908-9 " ; i " Madagascar (Sud) Forets Nord Ft. Dauphin 
Alluaud 1900 (17) " ; I " Haut Cote d'l voire, Danaue (Fort Hittos) A, Chevalier 
1 910, Avril " ; i " Madagascar Perrier ". 

10. Placonotus bolivari (Grouvelle) comb. n. 

Laemophloeus bolivari Grouvelle, 1905. 

I -75-2 -25 mm. ; body shining, yellowish red in colour, elytra much more yellow and almost 
transparent. 

Head very transverse, about twice as broad as long ; eyes situated in advance of base ; 
head turning abruptly medially just anterior to antennae and anteriorly again at lateral end 
of frontoclypeal suture ; suture evenly rounded ; epistome truncate ; antennae as long as or 
slightly longer than body in males, about o-j^ times length of body in females ; scape robust and 
about equal in length to second and third segments combined in males, somewhat shorter in 
females ; distal segment slightly curved in male, straight in the female, in female last three segments 
slightly enlarged forming a loose club ; disc of head regularly punctured with irregularly shaped 
small punctures, punctures separated by a distance equal to two eye facets, each with a very 
short seta ; intervals between punctures rather shining and with very fine micro-punctures. 

Pronotum very slightly broader than head at its broadest, 1*2 times as broad as long, broadest 
at about anterior quarter ; anterior angles ventral in p>osition since anterior half of region between 
lateral line and lateral margin inflected ventrally ; angles obtuse and slightly produced ventrally, 
posterior angles rectangular or very slightly produced ; lateral line consisting of a groove, 
expanding from about the mid-point to a maximum at included deep puncture, narrowing 
towards basal fifth ; secondary raised line present basally between primary line and lateral 
margin ; disc with puncturation and setation as on the disc of head. 

Elytra about twice as long as their combined maximum width which is at about their mid- 
point, there broader than pronotum, narrowing slightly until external apical angles, then turning 
more or less abruptly to sutural angle ; thus each elytron subtruncate ; cells not developed ; 
striae distinct ; elytra apparently glabrous except for a few larger setae apically ; abdomen 
only slightly exposed dorsally. 

East and West Africa, Congo. 

HoLOTYPE " Biafra Cabo S. Juan viii-1901 Escalera/Type/15/L. Bolivari Grouv. 
ty." (Grouvelle coll). There are eight other specimens standing next to the type 
of this species all collected in S. Thome, their data labels being written in differently 
coloured inks suggesting that they were acquired by Grouvelle at different times. 
The specimen collected by Escalera is the only one to bear the word type in Grou- 
velle' s handwriting although three of the others have a printed label with the word 
type in addition to the locality label. I do not accept the S. Thome specimens as 
syn types and consider that the Escalera specimen is the holotype. Other specimens 
seen include 2 " Luki 1950 R2306 Com. et Bois Coll. R. Mayne " ; 4 " Yangambi, 



i86 L. P. LEFKOVITCH 

1951 C. Donis Z.304, 321, 323, and 326 Coll. R. Ma5me Com. et Bois Congo R. 2326" ; 
I " Haut-Uele : Moto x-xi-1923 L. Burgeon " ; i " Mayumbe. Pulu-Banzi 17-ii- 
1924 A. Collar! " ; i " Ubangi : Gemena 3-1-1936 C. Leontovitch " ; 2 " Yan- 
gambi 3-vii-i952 Dr. Schedl s.295 " (Mus. R. Congo Beige) ; 2 " St. Thome " ; 
I " Ed. Luga Kondie Congo Beige " (Museum National d'Histoire Naturelle, Paris), 

11. Placonotus decor atus (Grouvelle) comb. n. 
Laemophloeus decor atus Grouvelle, 1908. 

This and the next species are the most distinct of African Laemophloeinae and 
certainly the most distinct Placonotus. P. exornatus (see below) differs in having 
the head, pronotum and the greater part of the elytra brown and not black as P. 
decor atus. 

Since Grouvelle's description is quite adequate, the species will not be fully 
redescribed. 

I' 75-2* 3 mm.; body shining ; mouthparts, basal segments of antennae, tibiae and tarsi 
dark brown ; femora, head, pronotum and the greater portion of each elytron black ; elytra 
otherwise yellow in a region at anterior third from about second or third strial interval to lateral 
margin. Elytra truncate, external apical angle being somewhat more distinct than usual ; 
last segment of abdomen almost entirely exposed dorsal] y ; in male paired accessory structures 
of genitalia often exposed. 

The lectotype, which I now select, is in the Grouvelle collection in Paris and bears 
the following data : " Af. or All Amani/Eichelbaum/Type/La^wo^A/o^ws decor atus 
sp. nv. Grouv," The other syntype in the same collection bears identical data 
except that it does not bear the name of the species in Grouvelle's handwriting. 

South Africa, Congo, Kenya. 

The author has seen specimens as follows : 2 " E. Cape Prov., Katberg 4000 ft. 
xii-1932 R. E. Turner " ; i " E. Cape Prov., Katberg 4000 ft. 13-15^11-1933 R. E. 
Turner " (British Museum (Natural History)) ; 23 " Ruanda : Kayove 2000 m. 
terr. Kisenyi P. Basilewsky 14-11-1953 " ; i " Manyema 1918 Sibatwa Kilengwe 
Dr. Gerard " ; i " Foret de bambous et Podocarpus Kenya : Elgon, Vers. Est. 
2500 m. (dans humus) N. Leleup 4-xii-i953 " ; 2 " Nioka xii - 1953 F. Deville Pap. 
49/Com. et Bois Congo R. 2507 " (Mus. R. Congo Beige) ; i " O A Nandi Reservat 
leg. Ahnder " (Munich). 

12. Placonotus exornatus (Grouvelle) comb. n. 

Laemophloeus (Silvanophloeus) exornatus Grouvelle, 1908. 

It has not been possible to find any external difference between this and the 
previous species except in their colour ; it is therefore possible to confuse immature 
P. decoratus with P. exornatus. This difference in colour may indicate that the 
two species are not really distinct since P. exornatus is only known to the writer 
by two specimens. It may be possible to show that the two species are in fact 
synonymous if P. decoratus is found together with P. exornatus in South Africa 
and the specimens of P. exornatus are kept alive and observed to see whether they 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 187 

change colour with time. A similar colour difference between two species of other- 
wise identical Cryptolestes was found to reflect a specific difference (Lefkovitch, 
1959a). 

South Africa. 

Grouvelle's original description was based upon a single male bearing the following 
data : "C. B. Ep. Donkis/Port Alfred/L. exornatus ty. Grouv." The author has 
seen a second specimen with the following data : " Port St. John Pondoland Nov. 
1923/S. Africa R. E. Turner " (British Museum (Natural History)), 

13. Placonotus mossus sp. n. 

This species most resembles P. donacioides (WoU.) in that the nature of the micro- 
sculpture on the head and pronotum and the consequent dull appearance distinguish 
these two species from other Placonotus. These two species can be distinguished 
easily in that P. donacioides is relatively longer in its build and possesses punctures 
on its head in addition to the reticulation. 

1-8 mm.; head and pronotum reddish brown, rather dull, elytra yellow and transparent. 

Head transverse ; eyes situated about half a diameter in advance of base ; epistome shallowly 
emarginate ; frontoclypeal suture with a groove the anterior margin of which is raised slightly, 
evenly rounded posteriorly ; antennae in male with segments longer than broad, becoming 
progressively longer after third ; scape the most robust segment ; antennae in total length just 
reaching apices of elytra ; disc of head impunctate but strongly reticulate into polygonal areas, 
three of these areas being slightly greater in area than an eye facet ; without setae on disc. 

Pronotum distinctly transverse, about i'25 times as broad as long ; broadest at about mid- 
point ; distinctly broader than head ; distance between anterior angles only just perceptibly 
greater than that between posterior angles ; anterior angles acute and projecting, lateral margin 
just posterior to them evenly sinuate, not abruptly so as in most Placonotus ; posterior angles 
obtuse ; lateral line expanded somewhat just behind mid-point to accommodate the deep 
puncture or fovea ; surface of disc otherwise impunctate but reticulate as on head ; glabrous. 

Elytra broadest at anterior third, there as broad as pronotum ; narrowing from this region to 
apices, each elytron separately rounded to apico-sutural angle ; elytra just twice as long as their 
combined width ; three cells present basally, outer margin of third continues further posteriorly 
than others ; striae more or less visible, apparently without setae. Legs somewhat shorter than 
is usual for the genus. 

Female unknown. 

Congo. 

HoLOTYPE male " Yangambi 1952 C. Donis z.2978 Com. et Bois Congo R.2440 " 
(Mus. R. Congo Beige). 

III. XYLOLESTES gen. n. 

Type species : Laemophloeus unicolor Grouvelle. 

This genus is most closely related to Xyhphloeus gen. n. which is described below. 
It resembles some species of that genus more closely than others, especially Xylo- 
phloeus patens (GrouveUe). The key to genera and the table on p. 190 outline the 
most important differences. 

i'5-2*5 mm.; body moderately flattened. 

Head distinctly transverse ; eyes situated about half their diameter in advance of base, 
occupying slightly more than one-third of lateral margin ; antennae inserted at about front 



i88 L. P. LEFKOVITCH 

corners of head and equal to half body length ; with last three segments more transverse and 
enlarged to form a distinct club ; epistome apparently straight but in fact with three shallow 
emarginations ; lateral line represented by a groove on each side, joined anteriorly by a trans- 
verse groove situated just behind epistome ; frontoclypeal suture obsolete ; labrum parallel- 
sided and truncate to distinctly emarginate anteriorly. 

Pronotum transverse, with rounded anterior and obtuse posterior angles ; lateral line repre- 
sented by a distinct groove ; basally a slight ridge and a shallow groove running laterally from 
lateral line to hind angle ; lateral lines joined anteriorly but not posteriorly by a groove. 

Scutellum triangular with rounded angles ; elytra normally covering whole abdomen ; the 
three cells clearly defined, posterior border of first and borders of third much more clearly 
defined than others which may be obsolescent ; each elytron distinctly carinate laterally, the 
carina lying close to outer margin of third cell after basal third. 

Genae normal ; gular sutures distinct ; anterior coxal cavities narrowly open posteriorly, with 
distinct lateral extension ; intercoxal process of the prosternum slightly concave posteriorly ; 
mesepimeron only just reaching mesocoxal cavities ; posterior margin of mesosternum slightly 
concave ; metasternum about equal in length to first and second visible abdominal sternites 
combined (Text-fig. 13) ; median metasternal line not reaching the anterior margin of sclerite ; 
first visible abdominal sternite about i-5 times as long as second, second to fifth about equal to 
each other ; legs subequal ; trochantero-femoral junctions of normal type ; tarsal formula 
5-5-5 (in both sexes). 

Xylolestes unicolor (Grouvelle) comb. n. 

(Text-fig. 14) 
Laemophloeus unicolor Grouvelle, 1908. 

This species was described by Grouvelle only in his key, to which the above 
reference relates, as " unicolor Grouv. Afrique or. all." Fortunately, it can be 
recognized as a distinct and not uncommon species in Africa. The species is re- 
described below. 

i-5-2'75 mm.; body reddish brown in colour, elytra more yellow than head and pronotum, 
tarsi and antennal clul3 yellow. 

Head transverse, trapezoidal in shape ; antennae just reaching basal quarter of elytra, fourth 
to eighth segments as wide as long, segments one to three more elongate, at most twice as long as 
wide ; last three segments about 1-5 times as broad as eighth and except for last segment, 
I "5 times as long as eighth ; last segment about twice as long as broad and shaped rather like 
an egg ; disc of head irregularly punctured with irregularly shaped punctures, these smallest 
and most nearly circular near mid-line, largest half-way between mid-line and lateral line ; 
the largest punctures somewhat larger in diameter than facets of eyes ; surface of head shining 
between punctures ; with sparse, very fine setae which are often not easily visible. 

Pronotum twice as wide as long, broadest in front of middle where thorax i"i-i-25 times as 
wide as head across and including eyes ; anterior angles rounded, posterior obtuse ; punctura- 
tion on disc finer than on head ; setation as on head. 

Elytra across shoulders as broad or broader than pronotum, slightly more than twice as long 
as their combined width, roughly oval in shape ; broadest at about anterior third, there about 
I -25 times as broad as pronotum ; thinner darker areas of elytra are arranged in rows and prob- 
ably represent striae ; apparently glabrous. 

There are two syntypes of this species in the Grouvelle collection, one with data 
" Zanzibar Raff ray /Laemophloeus unicolor Grouv ty " and the other " Af.or. All. 
Amani/Eichelbaum/Type [mss] " and a Paris museum type label. I now select 
the first of these specimens to be lectotype. 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 



189 



East and West Africa, Congo, Angola. 

I have examined the following specimens of the species : i " Nioka 27-ii-54 
F. Deville Pap, 57 R.2507 " ; 1 " Yangambi, 1951 z.261 R. 2333 " ; 7 " Coquil- 
hatville 27-viii-i930 J. Vrydagh " ; 2 " Eala vii-1935 J. Ghesquiere " ; 2 " Poto- 
pot pres Dima 4-ix-i930 G. F. de Witte " ; 2 " Ituri : La Moto : Madyu L. 
Burgeon " ; i " Equateur : Flandria xi-1929 R. P. Hulstaert " ; i " Haut Uele, 
Walsa, xi-1919 L. Burgeon " ; i " Recolte dans I'humus/Kivu : T. Fizi, Has 
Itombwe 1000 m. (Gal. for. Mukera) N. Leleup ii-52 " ; i " Kwamouth vi-1921 
Dr. H. Schouteden " ; i " Isangi 1929 J. Walkiers " ; i " Lulua ; Kapanga vii- 
1932 G. F. Overlaet " Miss. H. de Saeger : i (2062) ; i (2115) ; i (3563) ; i (3744) 
(see Appendix) (Mus. R. Congo Beige), i " Angola 1253. 11. 2. Dundo, galerie fores- 
tiere Luachimo, sous ecorces arbres tombes, xii-48 A. de Barros Machado " (British 
Museum (Natural History)) ; 22 " IFAN-1946 Tonkui C.I. 900-1200 m. A. Villiers " ; 
I " Amani " ; i " Cote d'lvoire Bimbroko " ; i " S. Thome (Museum National 
d'Histoire Naturelle, Paris). 



IV. XYLOPHLOEUS gen. n. 

Type species : Laemophloeus patens Grouvelle. 

This genus resembles the previous in several ways ; it can be distinguished by 
the characters given in the key to genera on p. 172 but especially by those given in 
the tabulated comparison with Xylolestes (p. 190). 





15 





Figs. 13-18. (13) Front and middle coxal cavities of Xylolestes unicolor ; (14) dorsal view 
of X. unicolor ; (15) front and middle coxal cavities of Xylophloeus patens ; (16) 
pronotum of X. bimaculatus and (17) of X. unifasciatus ; (18) dorsal view of X. patens. 
Scale lines : Figs. 13, 15 = 0-25 mm. ; Figs. 14, 16, 17, 18 =1 mm. 



I go 



L. P. LEFKOVITCH 



2'5 mm.; body moderately flattened. 

Head transverse and rectangular in its general appearance ; lateral lines distinct and forming 
part of dorsal margin of eye, joined anteriorly just behind epistome by an almost straight ridge ; 
epistome with three emarginations ; antennae inserted just behind front corners of head, their 
segments (except for scape and pedicel) slightly transverse, last three forming a loose club ; 
frontoclypeal suture obsolete although depressions corresponding with this and with median 
line can be seen if the incident light is at the correct angle ; eyes situated in advance of base of 
head ; labrum rounded anteriorly. 

Pronotum transverse, somewhat cordiform ; anterior and posterior angles acute ; lateral line 
represented by a ridge raised above the general surface ; internal to lateral line often a row of 
small punctures situated very close to lateral line, somewhat more distinct behind mid-point ; 
externally to lateral line a shallow circular fovea situated at about mid-point, diameter of 
which about equal to distance separating line from margin. 

Scutellum triangular, slightly transverse ; elytra with three cells developed although lateral 
margin of third obsolescent just after base ; no lateral carina present but lateral region evenly 
deflected ventrally from inner border of third cell. 

Genae normal ; gular sutures distinct ; anterior coxal cavities slightly open posteriorly ; 
anterior coxae spherical and well separated ; intercoxal process of prosternum straight posteri- 
orly ; metepimeron not contributing to mesocoxal cavity but mesepisternum and mesepimeron 
contribute equally, forming together about one-third of circumference of cavity (Text-fig. 15) ; 
metasternum equal in length to first two visible abdominal sternites ; first abdominal sternite 
twice as long as second, its intercoxal process straight anteriorly ; second sternite slightly longer 
than either third or fourth, the fifth about equal to or slightly longer than second. 

Legs subequal ; trochantero-femoral junctions of normal type ; tarsal formula 5-5-5 in 
female, 5-5-4 in male. 



Xylolestes gen. n. 

Lateral lines of head represented by groove, 
joined anteriorly by a transverse groove 

Epistome apparently straight but with three 
shallow emarginations 

Antennae inserted at front corners of head 

Labrum parallel-sided, truncate to distinctly 

emarginate anteriorly 
Prothorax with rounded anterior and 

posterior angles 
Lateral line of pronotum represented by a 

groove 
Each elytron carinate laterally 
Intercoxal process of prosternum concave 

posteriorly 
Mesepimeron just reaching middle coxal 

cavities 



Xylophloeus gen. n. 

Lateral lines of head represented by ridges, 

joined anteriorly by a transverse ridge. 
Epistome with three distinct emarginations. 

Antennae inserted behind front corners of 

head. 
Labrum rounded anteriorly. 

Prothorax with acute anterior and posterior 

angles. 
Lateral line of pronotum represented by a 

ridge. 
Elytra not carinate laterally. 
Intercoxal process of prosternum straight 

posteriorly. 
Mesepimeron and mesepisternum contribute 

equally to middle coxal cavities. 



Key to Species 

Pronotum at widest as wide as elytra at shoulders ...... 2 

Pronotum at widest narrower than elytra at shoulders ..... 4 

Pronotum hardly broader than long (i-i times) ; disc of pronotum convex 

2. mimosae sp. n. (p. 192) 
Pronotum transverse (1*3-1 '5 times as broad as long) ; disc of pronotum flat . 3 

Pronotum unicolorous yellow ; elytra coloured as pronotum i. patens (Grouvelle) (p. 191) 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 191 

Pronotum darker in colour on disc between lateral lines than outside ; elytra with dark 
markings 7- dentatus sp. n. (p. 194) 

4. Pronotum entirely black 3. chrysomeloides sp. n. (p. 192) 

- Pronotum red or yellow, at least in part ........ 5 

5. Elytra black but for the humeral angles which are dark red in colour 

4. elgonensis sp. n. (p. 193) 

- Elytra entirely yellow or with a dark maculation on each ..... 6 

6. Hind angles of pronotum slightly acute ; disc of pronotum almost black ; distance 

between hind angles of pronotum equal to or slightly more than that between the 

front angles (Text-fig. 16) 6. bimaculatus sp. n. {p. xg^) 

Hind angles of pronotum obtuse ; disc of pronotum hardly if at all darker than the 
lateral region ; pronotum narrower between the hind angles than between the 
front angles (Text-fig. 17) 5. unifasciatus sp. n. {p. 193) 

I. Xylophloeus patens (Grouvelle) comb. n. 

(Text-fig. 15) 
Laemophloeus patens Grouvelle, 1908. 

2'0-2-25 mm.; yellowish brown in colour, elytra almost transparent. 

Head with front region at slightly lower level than rest (? frons) ; lateral line formed by 
ridge and groove ; secondary line present behind eye ; antennae reaching to basal third of 
elytra, third segment slightly elongate ; disc evenly punctured, punctures equal to or slightly 
smaller than facet of eye, separated by one to three diameters, with fine setae about equal to 
three diameters in length. 

Pronotum 1-7 times as wide as long at maximum width, which varies from about mid-point 
to just behind anterior angles ; at maximum width prothorax very slightly broader than head ; 
lateral lines formed by slightly raised ridge, occasionally not joined by transverse line posteriorly ; 
anterior angles produced further anteriorly than greater part of anterior margin of pronotum, 
acute in shape but extreme apices rounded ; posterior angles sharp and acute-angled ; punctura- 
tion and setation on disc similar to that on disc of head. 

Elytra of maximum width at about basal third, there 1-25 times as broad as pronotum ; 
slightly less than twice as long as their combined width ; punctured striae present but elytra 
apparently glabrous. 

South and East Africa, Congo. 

There are 2 syntypes of the species in the Grouvelle collection ; I select as lecto- 
type a male which bears the following data : " Af. or. All Amani/Eichelbaum/Type/ 
;^g I Laemophloeus patens n.s. ty. Grouv." The data of the paralectotype are " Af. 
or. All. Amani/Eichelbaum/Type [mss] " and a Paris museum type label. 

I have also examined the specimens whose data are given below : 3 " Zululand : 
Eshowe. 6-31-V-1926 S. Africa R. E. Turner " ; 9 " Port St. John, Pondoland. 
Sept 1923 S. Africa R. E. Turner " ; i " Natal, near Durban (Virgin bush), 10- 
18.xii.1904 " ; I " Malvern, Natal G. A. K. Marshall — viii-1897 " (British Museum 
(Natural History)) ; i " Foret de transition/Tanganyika Terr : Uluguru Mts., 
val. Ullulu-Ndogo, 1500 m. 8-V-1957 /Mission Zoolog. I.R.S.A.C. en Afrique orientale 
(P. Basilewsky et N. Leleup) " ; i " Foret de montagne/Tanganyika en Afrique 
orientale (P. Basilewsky et N. Leleup) " ; 2 " Tanganyika Terr : Mt. Oldeani, Versant 
Est, for. bamb. 2350-2500 m. 6/9-vi-57/P. Basilewsky et N. Leleup " ; i " For. 
montange/Kivu : Terr. Kabare, S.E. Kahuzi, 2080/2200 m. N. Leleup viii-1951 " ; 



192 L. P. LEFKOVITCH 

I " Recolte dans Thumus/Kivu : Terr. Uvira, Mulenge 1880-2010 m, (Vest, for^t 
ombroph.) v-1951 N. Leleup " (Mus. R. Congo Beige). 



2. Xylophloeus tnimosae sp. n. 

3'0-4'0 mm.; body rather shining. 

Head 1-5 times as wide across eyes than long ; eyes situated one diameter in advance of base, 
their mid-points equidistant from base and antennal insertions ; lateral line formed by raised 
ridge ; secondary line present posterior to eye ; median line visible as a shadow when viewed 
with lateral illumination ; punctures on disc about equal to i'25 times diameter of eye facet ; 
separated by 2-4 times diameter of puncture, each with seta about equal to two diameters 
in length ; surface of head between punctures with short impressed lines inclined at an angle to 
median line and parallel with each other on each side, about equal to two punctures in length ; 
surface of head between punctures and lines otherwise shining. 

Pronotum i •1-1-2 times as wide as long, slightly broader than head, narrowing basally ; 
front angles acute, hind angles acute and projecting ; lateral line formed by raised ridge ; 
lateral margin weakly toothed at posterior border of fovea ; puncturation, setation and surface 
of disc as on head. 

Elytra broadest at about middle, there nearly twice as wide as pronotum, about 1*75 times as 
long as their combined width ; each elytron with internal margin of first cell visible apically, 
separated by striae from the basally visible internal margin of third cell ; each strial puncture 
with a short seta. 

Ethiopia. 

HoLOTYPE and 4 paratypes (i partially dissected) : " Under bark of decaying 
Mimosa/Djem-Djem Forest nearly 9,000 ft. i-x-1926 Dr. H. Scott " (British Museum 
(Natural History)). 

3. Xylophloeus chrysotneloides sp. n. 

2'5 mm.; trophi, epistomal region of the head, first two and base of third antennal segments 
reddish in colour ; remainder of head, pronotum and antennae black, antennae with yellow 
setae ; elytra yellow but for suture, interstices adjacent to fifth stria and lateral inflected margin 
brown in colour ; striae somewhat darker than interstices ; pygidium brown ; general surface 
rather dull. 

Head i'5 times as wide as long ; lateral line formed by raised ridge with adjacent row of 
closely applied punctures ; secondary line present posterior to eye ; punctures on disc about 
equal to eye facet in size, slightly smaller behind epistomal ridge, separated by one to three 
diameters ; surface of head strongly reticulate between punctures. 

Pronotum i'25 times as wide as head, broadest at about anterior third ; lateral line formed 
by a broad, raised ridge and a closely approximated row of punctures, just turning towards 
centre anteriorly before disappearing and therefore not meeting its fellow behind anterior 
margin of sclerite ; disc punctured as on head and with a similar reticulation ; surface between 
lateral line and lateral margin reticulate and with few punctures ; lateral margin without a 
tooth near the obsolescent fovea ; just before base of disc two very shallow foveae, their centres 
equidistant from lateral line and lateral mairgin. 

Elytra broadest at mid-point, there about i'75 times as broad as pronotum ; about i*8 times 
as long as their combined width ; punctured striae somewhat depressed below general level 
of elytra, each puncture with a seta ; surface with longitudinal reticulations between striae, 
almost micro-rugose ; pygidium coarsely and closely punctured, with long setae. 



a revision of african laemophloein ae 193 

Congo. 

HoLOTYPE female " Recolte dans I'humus/I.R.S.A.C. Kivu : Terr. Unira, Lubuka 
2780 m. (for. bamb.) 4-xi-5i N. Leleup " (Mus. R. Congo Beige). 
Male unkown, 

4. Xylophloeus elgonensis sp. n. 

This sp>ecies resembles X. mimosae sp. n. but differs in the following particulars : head, 
antennae, apical and basal regions of pronotum, scutellum, humeral angles of elytra and legs 
red in colour ; middle region of pronotum, remainder of elytra and pygidium black ; impressed 
lines between the punctures on discs of head and pronotum more dense and slightly deeper ; 
elytra across shoulders 1-3 times as broad as pronotum at its broadest. 

Kenya. 

HoLOTYPE " Dans feuilles mortes de bambous I.R.S.A.C. — Mus. Congo Kenya : 
Elgon, Vers. Est. 300 m. ii-xii-1953 N. Leleup " (Mus. R. Congo Beige). 



5. Xylophloeus unifasciatus sp. n. 

(Text-fig. 17) 

This species resembles X. elgonensis sp. n. very closely in its general shape and differs only in 
its coloration which is as follows : bases of elytra (except along the suture), apices of elytra 
and legs yellowish, body otherwise red. 

East Africa, Congo. 

HoLOTYPE male " Africa or. Katona/ Kilimanjaro x-1904 " (Termeszettudo- 
manyiMuzeum, Budapest) ; paratypes : i " Kivu : Tshibinda xi-1932 L. Burgeon "; 
I " Recolte dans I'humus/Kivu : T. Kalehe, 2200 m. Contref. S.O. Kahuzi (foret 
de bambous) viii-1951. N. Leleup " ; i " Kivu : Terr. Uvira, Lubuka 2180 m. 
(marecage) 3-ix-i95i. N. Leleup " (Mus. R. Congo Beige). 

6. Xylophloeus bimaculatus sp. n. 

(Text-fig. 16) 

3-4 mm.; disc of head and pronotum, middle of each elytron black in colour, remainder of 
body yellow ; shining and almost glabrous. 

Head only slightly broader than long ; antennae with all segments elongate, pedicel shortest ; 
lateral line formed by raised ridge and internal punctured groove ; secondary line present behind 
eye ; disc punctured, punctures somewhat larger than an eye facet, but somewhat smaller 
peripherally ; surface of head otherwise smooth and shining except for a few minute punctures 
between the larger ones. 

Pronotum at widest i'5 times as broad as head, broadest at basal third where lateral margin 
sinuate ; anterior angles produced, acute but apically rounded ; posterior angles slightly acute ; 
lateral line formed by ridge and internal punctured groove ; a deep fovea present in sublateral 
region ; disc with punctures as on head ; punctures of explanate sublateral region much smaller 
and very much sparser than those of disc. 

Elytra at widest 1*5 times as wide as pronotum, broadest at basal third just anterior to darker 
markings ; between humeral teeth elytra i'4 times as wide as pronotum ; elytra about i-5 times 
as long as their combined width ; not carinate laterally but laterally evenly curved ventrally 



194 L. P. LEFKOVITCH 

except for extreme lateral region which is explanate ; third cell well developed basally, outer 
margin disappearing before the maculation ; elytra striate. 

Congo. 

HoLOTYPE female " Ruanda : Yanina, terr. Klibuye, 2,300 m i2-ii-i953. P. 
Basilewsky " and i paratype, female " Recolte dans I'humus/I.R.S.A.C. Kivu : 
Terr. Uvira, Mulenga 1880-2010 m. (Vest. for. ombroph.) v-1951 N. Leleup " (Mus. 
R. Congo Beige). 

7. Xylophloeus dentatus sp. n. 

3'G-3'5 mm.; trophi, anterolateral regions of head, scape, pedicel, lateral regions of pronotum 
and elytra, basal region of elytra adjacent to scutellum and median apices of elytra (other than 
suture) yellow to red in colour, remainder of dorsal surface black ; antennae with yellow setae ; 
moderately shining. 

Head about twice as wide as long, less trapezoidal in shape than usual for the genus ; lateral 
line formed by raised ridge ; a secondary line present behind eye ; disc of head punctured, the 
punctures of diameter about equal to nearly twice that of eye facet, except near and at median 
region where they are smaller ; separated by distance equal to about half their diameter to 
twice their diameter ; disc of the head polygonally reticulate anteriorly but laterally compressed 
elsewhere ; surface of disc otherwise shining. 

Pronotum 1-7 times as wide as long, at widest about 1*3 times as wide as head, broadest at 
about anterior third and equally broad across lateral blunt teeth situated on lateral margin at 
the basal third ; anterior angles produced, acute in shape but rounded at the extreme apices ; 
posterior angles acute and slightly projecting ; lateral line formed by raised ridge and internal 
groove, not joined together transversely behind anterior margin ; sublateral fovea large in area, 
encroaching upon ridge of lateral line ; at region of fovea, lateral margin thickened, forming a 
blunt tooth ; lateral region explanate ; disc with punctures smaller and further distant than 
those of head, separated by a distance equal to twice to six times their diameter. 

Elytra broadest at basal third, 1*7 times as broad as pronotum, slightly broader across shoulders 
than pronotum (i-i times) ; almost twice as long as their combined width ; evenly rounded to 
apico-sutural angle ; in longitudinal section elytra sinuate, being concave basally becoming 
convex at apical third ; carinate laterally, lateral margin narrowly explanate. 

Kenya, East Africa. 

HoLOTYPE " Ravin boise/Tanganyka Terr. : Mt. Meru, Oikokola, versant N.O. 
2750 m. 25-vi-i957/Mission Zoolog. I.R.S.A.C. en Afrique orientale (P. Basilewsky 
et N, Leleup) " (Mus. R. Congo Beige) ; i paratype " Kidumnik (?) Ukunda [Kenya] 
ii-ig47/Com. Inst. Ent. Coll. No. 10840 " (British Museum (Natural History)). 

V. BLUBOS gen. n. 

Type species : Bluhos matris sp. n. 

This genus appears to stand apart from other Laemophloeine genera known to 
the writer and as such it is difficult to find another genus with which it can be 
compared. It resembles, at least superficially, some of the species of Xylophloeus 
gen. n. but differs in many features, the most conspicuous being that the suture 
between the pronotum and pro-episternum is dorsal in position and not lateral as 
in that genus. 



Head parallel at base, turning outwards just behind eyes ; across eyes head as broad as prono- 
tum ; antennae about equal in length to head and pronotum, third segment somewhat longer 



I 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 



195 



than second, distal three segments sHghtly enlarged ; antennae inserted laterally, behind the 
apparent front angles of head, insertions somewhat sunken ; lateral line on head behind eyes 
represented by very fine ridge, anterior to eyes this ridge more completely developed ; lateral 
lines joined anteriorly behind epistome by an evenly curved concave ridge ; that part of frons 
anterior to this ridge evenly emarginate anteriorly, the whole anterior region of head having 
three emarginations ; labrum very slightly emarginate anteriorly, otherwise evenly rounded. 

Pronotum transverse, broadest in front of middle ; anterior angles acute, posterior angles acute 
and produced ; lateral lines represented by groove, its outer margin continuous with anterior 
and posterior discal margins ; secondary line present basally running medially from posterior 
angle and turning anteriorly for short distance just before primary line before disappearing ; 
at about mid-point of lateral region two large, deep punctures or foveae situated between 
lateral line and lateral margin of pronotum ; suture between pronotum and pro-episternum 
visible dorsally, pro-episternum providing outermost lateral region of prothorax. 




Figs. 19-20. (19) Front and middle coxal cavities of 5/m6os ma^m ; (20) donsal view of 
B. matris. 

Scale lines : Fig. 19 = 0-25 mm. ; Fig. 20 = i mm. 



196 L. P. LEFKOVITCH 

Scuiellum triangular, apex sharp ; elytra with distinct, punctate striae, third cell developed 
apically ; not carinate laterally ; somewhat narrowed apically, rounded to apico-sutural angle, 
leaving part of last abdominal segment exposed dorsally. 

Genae normal, gular sutures distinct ; anterior coxal cavities broadly open behind, slightly 
transverse ; hind margin of intercoxal prosternal process concave posteriorly ; intercoxal 
process posteriorly deflected dorsally ; metepisternum not reaching mesocoxal cavity ; mesepi- 
meron and metasternum meeting on a broad front (Text-fig. 19) ; metasternum as long as first and 
second abdominal sternites combined ; intercoxal process of first visible abdominal sternite 
slightly rounded anteriorly ; abdomen narrowing towards apex ; first visible abdominal sternite 
twice as long as second, second to fourth equal to each other, the fifth slightly longer than fourth ; 
legs of normal length for the subfamily ; trochantero-femoral junctions of normal type ; tarsal 
formula 5-5-4 in the male ; female unknown. 



Blubos matris sp. n. 

(Text-fig. 20) 

About 4 mm.; head, pronotum and antennae reddish yellow in colour, elytra more yellow, 
the punctures somewhat red ; each elytron with one dark brownish mark, both together giving 
the impression of the letter " M "; legs yellow with bases of tibiae and tarsi more red in colour ; 
apices of mandibles black. 

Head with large punctures, separated by slightly less than one diameter, surface shining 
between them, each puncture with a short seta. 

Pronotum with puncturation and setation on disc as that of disc of head ; region between 
lateral line and lateral margin with very few, much smaller punctures, each with a seta. 

Elytra with first, third and fifth intervals with a few punctures present basally, these punctures 
further separated than those of striae ; apices of elytra with a few slightly longer setae ; pygidium 
finely punctured, with fine reticulation between punctures. 

Congo. 

HoLOTYPE " Kivu : contr. S. Kahuzi, km. 27, 2200 m., 28-iii-i953 P. Basilewsky " 
(Mus. R. Congo Beige). 

VI. MICROLAEMUS gen. n. 

Type species : Microlaemus turneri sp. n. 

This genus includes species which superficially resemble those of Hypocoprus 
Mots. ; amongst the Laemophloeinae, the genus could be confused with some 
Cryptolestes but can be distinguished from that genus by the possession of a distinctly 
visible frontoclypeal suture, the transverse anterior coxae with their cavities open 
posteriorly (Text-fig. 21), and with the fifth visible abdominal sternite about 
twice as long as the fourth whereas Cryptolestes has closed front coxal cavities, the 
coxae being globular, the frontoclypeal suture obsolete and the last visible abdo- 
minal sternite only as long as the fourth at the most. 

1-2-5 mm. ; body moderately flattened. 

Head somewhat triangular in shape ; eyes moderate in size, situated in advance of base ; 
antennae with scape and pedicel elongate, other segments transverse, distal three segments 
forming a loose club ; epistome shallowly emarginate or truncate anteriorly ; lateral line 
represented by raised ridge and closely approximated internal groove ; frontoclypeal suture 
and median line represented by slight depressions in general surface of head. 



i 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 197 

Pronotum slightly transverse or very occasionally very slightly longer than broad ; anterior 
and posterior angles obtuse to rounded in shape ; lateral line represented by raised ridge and a 
closely situated internal groove. 

Scutelltim triangular in shape ; elytra carinate laterally ; three cells developed, lateral margins 
of first and second indistinct ; secondary intervals narrow. 

Genae moderately well developed lateroventrally to trophi, gular sutures obsolete ; anterior 
coxae transverse, with distinct lateral extensions, their cavities open posteriorly ; intercoxal 
process of prosternum separating coxae to some extent, in the smallest species separation much 
less in proportion to general body structure than in more typical species of genus ; intercoxal 
process convexly rounded posteriorly ; metepisternum just contributing to mesocoxal cavity, 
mesepimeron and mesepisternum contributing about equally ; metasternum equal in length to 
first two visible abdominal sternites combined ; intercoxal process of first visible abdominal 
sternites narrow and rounded anteriorly ; first abdominal sternite 2- 2 5-2' 5 times as long as 
second, second to fourth equal to each other, fifth somewhat longer, legs subequal ; tro- 
chantero-femoral junctions of normal to elongate type ; middle femora less dilated than 
either front or back ; tarsal formula 5-5-5 in female, 5-5-4 in male. 

Key to African Species 

1. Body less than 1-5 mm. in length ; frontoclypeal suture obsolescent 4. slades sp. n. (p. 199) 

- Body more than 1-5 mm. in length ; frontoclypeal suture distinct ... 2 

2. Body very dark brown to black in colour, palpi pale yellow in colour 

3. palpalis (Waterhouse) (p. 198) 
Body brown in colour, palpi almost the same colour as the rest of the body . . 3 

3. Head produced anterior to the insertions of the antennae and rather narrow across 

the epistome (Text-fig. 22) . . . .2. ferrugineus sp. n. (p. 198) 

- Head normally developed anterior to the antennal insertions and of normal width 

across the epistome (Text-fig. 23) . . . . . i. turner i sp. n. (p. 197) 

The following species, hitherto placed in Laemophloeus, belong to Microlaenms : 
M. sylvestris (Grouvelle), M. interceptus (Grouvelle), both from India, M. picipennis 
(Grouvelle) from Ceylon and M. sulcijrons (Grouvelle) from Reunion combb.n.; 
Lathropus hrightensis Blackburn from Australia may belong to this genus although 
further study is necessary before this last opinion can be confirmed. 

I. Microlaemus turneri sp. n. 

(Text-figs. 23, 24) 

i75-2-omm. ; brown in colour ; pubescent. 

Head i -5 times as wide as long ; epistome rounded anteriorly ; frontoclypeal suture evenly 
concave ; distance between anterior margin of eye and apex of epistome equal to that between 
anterior margin of eye and base of head ; antennae just reaching base of pronotum ; mandibles 
expanded laterally so that their posterolateral border is parallel with anterior border of the 
produced genae ; disc of head evenly and shallowly punctured, punctures slightly latter in 
diameter than facet of eye, separated by distance equal to once to twice times their diameter, 
each with seta equal to two diameters in length ; surface between punctures polygonally 
reticulate. 

Pronotum very slightly transverse, widest at anterior quarter, slightly broader than head 
(i* I times), narrowed basally ; anterior and posterior angles slightly obtuse ; disc with punctures, 
setae and surface as on head. 

Elytra twice as long as their combined width, broadest at their mid-point, there i*i times as 
broad as pronotum ; apices of the elytra separately rounded to the apico-sutural angle ; striate, 
setae as long as those of head and pronotum. 

ENTOM. 12, 4. 14 



iqS 



L. P. LEFKOVITCH 



South Africa. 

HoLOTYPE male " Natal : Kloof 1500 ft. Sept. 1926 R. E. Turner " and 8 paratypes 
with same data; other paratypes: 6 " Zululand : Eshowe 1-22. v. 1926 R. E. 
Turner " ; i " Port St. John, Pondoland June i-ii. 1923 R. E. Turner " ; 5 " Stn. 
No. 59 Deepwalls Forest. Knysna, i6-iii-i954, under bark of Yellow-wood, 1725 ft. 
Cape Province J. Balfour Browne " ; 1 " Stn. No. 59 Deepwalls Forest. Knysna, 
i7-iii-i954 small pool in glade, 1725 ft. Cape Province J. Balfour Browne " (British 
Museum (Natural History)). 

2. Microlaetnus ferrugineus sp. n. 

(Text-fig. 22) 

Resembles M. tuvneri sp. n. except as follows : head very much more triangular, epistomal 
region quite narrow ; distance between anterior margin of eye and epistome twice that of 
distance between anterior margin of eye and base of head ; frontoclypeal suture only slightly 
concave in shape. 

HoLOTYPE " Mauritius J. E. M. Brown B.M. 1899-265 " (British Museum (Natural 
History)). 

3. Microlaetnus palpalis (Waterhouse) comb. n. 

Laemophloeus palpalis Waterhouse, 1876. 
Laemophloeus tenebrosus Grouvelle, 1899, syn. n. 

I'S mm.; body reddish black in colour, maxillary palpi and tarsi yellow ; setae on head and 
pronotum white to yellow in colour. 





22 





23 



24 



Figs. 21-24. (21) Front and middle coxal cavities of Microlaetnus slades ; (22) head 
of M. ferrugineus and (23) of M. turneri ; (24) dorsal view of M. turneri. 
Scale lines : Figs. 21-23 = °'^5 "^"^- '> ^^S- 24 = i mm. 



A REVISION OF AFRICAN L AEMOPHLOEIN AE tgg 

Head coarsely punctured, punctures equal in diameter to two eye facets, separated by less 
than half one diameter ; punctures smaller on frons ; surface, including that of internal regions 
of punctures, reticulate. 

Pronotum slightly longer than wide ; disc of pronotum punctured similarly to that of head, 
but punctures further separated ; surface reticulate ; lateral margin with eleven or twelve blunt 
tubercles ; sublateral region with row of large, closely approximated punctures situated close 
to lateral line, their medial margins forming part of lateral line ; external to these punctures a 
second row of deep and smaller punctures alternating with first row but absent at base and apex. 

In its body proportions this species is identical with the generic type. 

HoLOTYPE " Rodriguez G. Gulliver yb-i^lLaemophloeus palpalis [Type) C. 
Waterh." (British Museum (Natural History)). The holotype of L. tenebrosus 
Grouvelle, which bears the data "I. Bourbon [Mauritius] /Type [mss] I Laemophloeus 
tenebrosus Grouv " and a Paris museum type label, was found to be the same species 
as M. palpalis (Waterhouse) . 

4. Microlaetnus slades sp. n. 

i*o mm.; yellowish brown in colour ; pubescent, dull. 

Head 1*3 times as wide as long ; distance between anterior margin of eye and epistome 1*5 
times that between anterior margin of eye and base of head ; disc polygonally reticulate, 
apparently without punctures, covered with long setae, each seta being nearly as long as scape. 

Pronotum i*i times as long as wide, at its widest I'l times as wide as head ; anterior and 
posterior angles obtuse but rounded distally ; surface of disc reticulate and with setae as on head. 

Elytra at their widest as wide as pronotum, twice as long as their combined width, pygidium 
sometimes partially exposed dorsally ; striate and with rows of setae, setae slightly shorter than 
those of head and pronotum. 

Congo. 

Holotype " Bas-Uele : Kotell 1-21.1.1925 Dr H. Schouteden " and 2 para- 
types with the same data (Mus. R. Congo Beige) ; i paratype " Haut-Ovelle Env. 
de Madyu L. Burgeon 1919 " (Museum National d'Histoire Naturelle, Paris). 

Vn. MAGNOLEPTUS gen. n. 

Type species : Magnohptus pugnaceus sp. n. 

This genus differs from the closely related Leptophloeus Casey in several ways, the 
most important being the considerably larger size of the adults, the tarsal formula 
of 5-5-4 in the males as against 5-5-5 in Leptophloeus, the arrangement of the 
intervals on the elytra, the detailed structure and arrangement of the mesocoxal 
cavities and the rather nebulous difference in the surface appearance. The genus 
resembles small Passandridae in its general appearance. 

Body flattened cylindrical or cylindrical in shape ; 3-5 mm. 

Head as long as broad ; eyes small, situated one to two diameters in advance of base ; lateral 
Une represented by raised ridge above and anterior to eyes, posterior to eyes the line obsolescent ; 
frontoclypeal suture obsolete ; epistome truncate anteriorly ; antennae with pedicel shorter 
than scape or third segment, otherwise filiform or almost imperceptibly enlarged distally. 

Pronotum longer than broad, narrowed basally, anterior angles rounded or obtuse, hind angles 
obtuse to acute, sometimes obsolescent ; lateral line represented by a shallow groove closely 
approximated to an external raised ridge. 



200 L. p. LEFKOVITCH 

Scutellum pentagonal ; elytra more than 2*5 tinaes as long as their combined width, sometimes 
more than three times ; carinate laterally, carina formed by outer margin of third cell ; three 
cells present, secondary intervals narrower than cells. 

Genae normal ; gular sutures distinct ; anterior coxal cavities closed posteriorly, the coxae 
spherical, situated fairly close together ; intercoxal process of the prosternum straight or slightly 
convex posteriorly ; metepisternum not contributing to the mesocoxal cavities (Text-fig. 25) ; 
posterior margin of mesosternum straight ; median suture of the metasternum not reaching 
anterior margin of sclerite ; intercoxal process of first visible abdominal sternite narrow and 
rounded anteriorly, this sternite 1*5 times as long as second, second to fifth about equal to each 
other. 

Legs subequal ; trochantero-femoral junctions of normal type ; tarsal formula 5-5-5 in 
female, 5-5-4 in male. 



Key to Species 

Head and pronotum black in colour, rugosely punctured, not shining ; elytra 
parallel-sided and semi-truncate apically ; hind angles of pronotum very obtuse 
(Text-fig. 27) . . . . . . .2. parallelicollis sp. n. (p. 



lOi) 




25 




26 

Figs. 25-27. (25) Front and middle coxal cavities of Magnoleptus parallelicollis ; (26) 
dorsal view of thorax and elytra of M. pugnaceus ; (27) dorsal view of M. paralleli- 
collis. 

Scale lines : Fig. 25 = 0-25 mm. ; Figs. 26, 27 = i mm. 



A REVISION OF AFRICAN LAEMOPHLOEIN AE 201 

Head and pronotum blackish red, simply punctured, shining ; elytra diverging from 
their bases to just behind the mid-point and then narrowing somewhat ; apices 
evenly rounded to the apico-sutural angle ; hind angles of pronotum rectangular 
(Text-fig. 26) ........ I. pugnaceus sp. n. (p. 201) 

I. Magnoleptus pugnaceus sp. n. 

(Text-fig. 26) 

3-0-5-0 mm.; body blackish red in colour except the tarsi which are less black. 

Head as broad as or slightly broader across eyes than pronotum ; disc of head simply punctured, 
punctures somewhat elongate, separated by once to twice their diameter ; surface between 
punctures micro-reticulate ; almost devoid of setae. 

Pronotum somewhat broader than long ; anterior angles rectangular and blunt, posterior 
angles very distinct and projecting ; puncturation and setation as on head. 

Elytra about 2-25 times as long as their combined width, parallel-sided until mid-point and 
then narrowing somewhat apically, evenly rounded to apico-sutural angle ; third cell obsolete 
basally but well developed apically. 

Congo, Rhodesia, Angola. 

HoLOTYPE male " N.W. Rhodesia Kashitu, N. of Broken Hill ii-1915 H. C. 
Dollman " and the allotype with the same data ; paratypes : i " Angola 10656.42 
Marco de Canavezes (Cubal da Ganda), district de Benguela, piege lum., iii-56 
Ed. Luna de Carvalho " (British Museum (Natural History)) ; i " Mulungu, 1951 
P. C. Lefevre 691 Coll. R. Mayne Com. et Bois Congo r.2371 ". The following 
paratypes were collected by Dr. K. Schedl : i Yangambi 15.x. 1952 ; 3 Mulungu 
5.viii.i952; 10 Yangambi 18.vii.1952; i Yangambi 21.x. 1952; 2 Hembe- 
Bitale 18.vii.1952 ; 2 Ruanda Jhembe 29.vii.1952 (Mus. R, Congo Beige). 

2. Magnoleptus parallelicollis sp. n. 

(Text-fig. 27) 

3 "0-3 "5 mm.; head and pronotum black, trophi, antennae, legs and elytra brownish black ; 
body quite cylindrical in section. 

Head rugosely punctured, punctures elongate, intervals between punctures reticulate ; setae 
quite short. 

Pronotum slightly longer than broad ; anterior angles obtuse and sharp, posterior angles 
very obtuse, almost obsolete ; puncturation and setation similar to that on head. 

Elytra three times as long as their combined width, parallel sided, each elytron subtruncate 
apically. 

Congo. 

HoLOTYPE male " Recolte dans cellule de bambous/I.R.S.A.C. — Mus. Congo 
Kivu : Uvira, Hte Luvubu 265 m 27-xii-i95o N. Leleup " and 2 paratypes with 
the same data (Mus. R. Congo Beige). 

Vni. LEPTOPHLOEUS Csisey 

Leptophloeus Casey, 191 6. 
Truncatophloeus Kessel, 192 1, syn, n. 

Type species : Laemophloeus angustuhts Leconte (by original designation). 



202 L. P. LEFKOVITCH 

This genus has been redescribed by Lefkovitch (19596). Additions to that description follow : 
epistome rarely with tooth in mid-line or with two teeth ; lateral line on head formed by raised 
ridge and closely approximated internal groove ; elytra more or less parallel-sided ; genae 
somewhat produced anteriorly ; gular sutures obsolescent or obsolete ; posterior border of 
intercoxal process of prosternum straight or very slightly concave posteriorly ; metepisternum 
not contributing to mesocoxal cavity ; mesepimeron just reaching cavity, mesepisternum contri- 
buting to cavity but limited to a small region laterally ; median line of metasternum not reaching 
anterior third of sclerite ; trochantero-femoral junctions of normal type ; in two species (L. 
problemaiicus sp. n. and L. cornutus sp. n.) the hind tarsi of males four-segmented, otherwise 
5-5-5- 

Two species, Laemophloeus mohilis Grouvelle and L. insularis Grouvelle, are 
included in Kessel's (1921) subgenus Truncatophloeus of Laemophloeus which was 
defined merely by " Der Clypeus ist vorn gerade abgestiitzt ". Examination of the 
tj^e specimens of both species has shown that they have the generic characters of 
Leptophloeus. I now select Laemophloeus mohilis Grouvelle as type species of 
Truncatophloeus Kessel which thereby becomes a junior synonym of Leptophloeus 
Casey ; both of these species are now transferred to Leptophloeus, comb. n. 

The species of this genus fall into three groups which are not sufficiently distinct 
to define as subgenera. The dematidis group includes the darkly coloured, rather 
elongate species with a tendency to rugose sculpturing on the head and prothorax ; 
the alternans group include the brown (but occasionally black) species which are 
less elongate and which have simple puncturation without rugosity on the head and 
prothorax ; the problematicus group is more distinct and includes those species 
having four-segmented hind tarsi in the males and with an unusual epistome. In 
other respects this last resembles the alternans group. 

There remains the possibility that the problematicus species group does not properly 
belong to Leptophloeus ; I am neither able to place them in any other genus nor 
find any difference in the structure of the adults which together with the curious 
epistomal structures and tarsal formula would justify the erection of a new genus 
to contain the species. They perhaps belong to a different subgenus from the other 
species here considered to belong to Leptophloeus but I would prefer to await the 
discovery of a third species, if one exists, before confirming or refuting this suggestion. 

Key to African Species 

1. Epistome produced into a tooth, horn or horns over the labrum ... 2 

- Epistome not produced into a tooth or horns over the labrum but truncate or 

emarginate ............ 4 

2. Epistome with two lateral horns (Text-fig. 28) ; males with four-segmented hind 

tarsi ........ lb. problematicus s^. n. (p. 210) 

- Epistome with one more or less medial tooth ; males with four- or five-segmented 

tarsi ............. 3 

3. Punctures on head separated by a distance less than their diameter ; head with 

medial raised ridge ; males with four-segmented hind tarsi (Text-fig. 29) 

17. cornutus sp. n. (p. 210) 

- Punctures on head separated by a distance greater than their diameter ; head 

without medial raised ridge ; males with five-segmented hind tarsi (Text-fig. 30) 

4. opaculus (Grouvelle) (p. 205) 

4. Epistome truncate or only very slightly emarginate anteriorly (Text-fig. 31) . 5 



A REVISION OF AFRICAN LAEMOPHLOEINAE 



203 



Epistome strongly emarginate anteriorly (Text-figs. 32, 33) .... 10 

5. Body black and shining ........... 6 

- Body brown and dull ........... 7 

6. Posterior angles of pronotum obtuse, clearly defined and somewhat projecting ; 

head simply punctured . . . . . . .1. ater sp. n. (p. 204) 

- Posterior angles of pronotum obsolete ; head longitudinally rugosely sculptured 

6. axillaris (WoUaston) (p. 206) 

7. Head with median line raised above general surface into a ridge 

14. bupleuri (Peyerimhof) (p. 209) 

- Head, if with visible median line, the line is depressed ..... 8 

8. Elytra strongly carinate laterally, more than three times as long as their combined 

width ........ 13. mobilis (Grouvelle) (p. 208) 

- Elytra abruptly deflected laterally but not carinate, about 2-5 times as long as their 

combined width ........... 9 

9. Head and pronotum longitudinally rugosely sculptured ; eyes situated about one 

diameter in advance of the thorax ; elytra at shoulders as broad as the prothorax, 
subtruncate apically . . . . . . .7. parallelus sp. n. (p. 206) 

- Head and pronotum simply punctured ; eyes almost contiguous with the prothorax, 

elytra at shoulders broader than the prothorax, broadly rounded apically 

2. lucidus (Grouvelle) (p. 205) 
[o. Pronotum 1-5 times as long as the breadth between the posterior angles 

8. mucunae sp. n. (p. 206) 
— Pronotum at most 1-25 times as long as broad between the posterior angles . . 11 




28 








31 



Figs. 28-33. Heads of Leptophloeus ; (28) L. problematicus ; (29) L. cornutus 
(30) L. opaculus ; (31) L. ater ; (32) L. janeti ; (33) L. punctatus. 

Scale line = 0-25 mm. 



204 L. P. LEFKOVITCH 

11. Elytra about 2-25 times as long as their combined width . . . . . 12 

— Elytra at most about twice as long as their combined width . . . . 14 

12. Pronotum at most as long as broad . . • 5- stenoides (WoUaston) (p. 206) 

— Pronotum longer than broad . . . . . . . . . .13 

13. Body shining, reddish yellow in colour ; head more or less smooth between the 

punctures ......... 9. cassavae sp. n. (p. 207) 

— Body dull, brown in colour ; head reticulate between the punctures 

10. linearis (Grouvelle) (p. 207) 

14. Third segment of antennae narrower than second ; body dark brown in colour 

II. punctatus sp. n. (p. 208) 

— Third segment of antennae about as broad as second ; body yellow to brown in 

colour ............. 15 

15. Head and pronotum smooth between the elongate punctures 

15. janeti (Grouvelle) (p. 209) 

— Head and pronotum with longitudinal micro-sculpture between the more circular 

punctures . . . . . . . .12. capitus sp. n. (p. 208) 

It has not been possible to include 3. L. anormus (Grouvelle) in this key since it 
has neither been recognized from the description nor has the type been located 
{infra, p. 205). 

The alternans Species Group (see p. 202) 
I. Leptophloeus ater sp. n. 

(Text-fig. 31) 

This species is not unlike the European L. alternans (Erichson) in its general 
appearance but differs in that the colour is black, the eyes are more convex, the 
puncturation is coarser and the elytra are shorter in proportion to the rest of the 
body. 

2-0 mm.; head, pronotum and elytra black, trophi, antennae and legs red. 

Head across eyes slightly broader than long ; epistome truncate anteriorly ; antennae reaching 
basal third of elytra, last three segments enlarged to form a loose club, last segment twice as 
long as wide ; disc of head with large punctures, about one and a half times to twice diameter 
of an eye facet, somewhat elongate in shape ; punctures separated by a distance equal to 
once to twice their diameter basally, by twice to four times their diameter apically ; each 
puncture with a very short seta, or apparently glabrous. 

Pronotum longer than wide (i-i times) ; at its widest narrower tlian head across eyes (0-95 
times) ; anterior angles obtuse and rounded ; posterior angles obtuse, sharp and somewhat 
projecting ; puncturation as on basal region of head, with very short setae or apparently 
glabrous. 

Elytra twice as long as their combined width ; i -i times as broad as pronotum across shoulders, 
very slightly broader at mid-point ; each elytron separately curved to the apico-sutural angle, 
just leaving tip of pygidium exposed dorsally in mid-line ; with punctured striae, each puncture 
subtending a short seta. 

Congo. 

HoLOTYPE and i paratype " Mulungu, 1951 P. C. Lefevre P. C. L. 642 " ; other 
paratypes : i " Mulungu 8-viii-i952 R, Mayne. R.M. 592A " ; 3 " Mulungu 
5-viii-i952 Dr. Schedl Nr. 267 " (Mus. R. Congo Beige). 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 205 

2. Leptophloeus lucidus (Grouvelle) comb. n. 

Laemophloeus lucidus Grouvelle, igo8. 

This species resembles L. ater sp. n. in almost all details and differs primarily from that species 
in the colour of the main part of the body which is brown and not black. Since Grouvelle's 
original description is very detailed, no further redescription will be given here. 

East Africa, Ethiopia. 

HoLOTYPE " Af. or All. Amani/Eichelbaum/Type [mss]/3h /Laemophloeus lucidus 
Grouv ty " and a Paris museum type label. 

In addition to the holotype described from Dar-es-Salaam, the following specimens 
have been examined : i " Abyssinia : Djem-Djem Forest neariy 9,000 ft. i-x-1925 
Dr. H. Scott " ; i " Abyssinia/Schimper 430-50 ". 



3. Leptophloeus anormus (Grouvelle) comb. n. 
Laemophloeus anormus Grouvelle, 1923. 

Although the type specimen of this species has not been located in either the 
Grouvelle or the Alluaud collections, the original description presents sufficient 
information to establish that the species properly belongs to Leptophloeus. Grouvelle 
wrote that the species is related to L. janeti (Grouv.) and L. axillaris (WoU.), being 
distinguished from the former by the truncate epistome and from the latter by the 
more slender antennae, the more elongate body and an emargination at the outside 
of the base of the prothorax. I have not been able to recognize specimens of a 
species resembling L. janeti and L. axillaris, but differing from them as given and 
agreeing with the rest of the description of L. anormus. In my key to species of 
Leptophloeus {supra, p. 202), L. anormus would probably be identified as L. lucidus 
(Grouvelle). 

Kenya. 

According to the original description, the species was found " Riviere Amboni 
dans la zone des prairies inferieures du mont Kenya, alt. 2,000 m." by Alluaud. 

4. Leptophloeus opaculus (Grouvelle) comb. n. 

(Text-fig. 30) 

Laemophloeus opaculus Grouvelle, 1908. 

This species resembles L. lucidus fairly closely but differs in the epistome having a pointed 
tooth in the mid-line. In other respects, this species resembles the species of the alternans grou^. 
The figure of the head (Text-fig. 30) will enable accurate recognition of the species. Since only 
the type is known, there remains the possibility that this specimen is an abnormal individual of 
another species. The shape of the prothorax, however, with its projecting, slightly acute hind 
angles is unlike that of any other African species of the genus. 

East Africa. 

Holotype " Afr. or. All. Amani Eichebaum \sic] Type I Laemophloeus opaculus 
Grouv. ty." 



2o6 L. P. LEFKOVITCH 

5. Leptophloeus stenoides (Wollaston) comb. n. 

Laemophloeus stenoides Wollaston, 1854. 

Resembles L. alternans (Erichson) very closely but differs in the greater emargination of the 
epistome and in having the surface of the head and of the pronotum polygonally reticulate upon 
a moderately rugose sculpturing. 

The surface is so distinct in this species as to distinguish it completely from other Leptophloeus 
occurring in Africa. 

Madeira. 

HOLOTYPE, the only specimen seen, is in the Wollaston collection (British Museum 
(Natural History)). It has a British Museum type label and " Laemophloeus stenoi- 
des, type Woll " in a handwriting I do not recognize but which may be that of Arrow. 

The clematidis Species Group (see p. 202) 
6. Leptophloeus axillaris (Wollaston) comb. n. 

Laemophloeus axillaris Wollaston, 1854. 

This species resembles L. clematidis in its general appearance but is reddish black rather than 
brown in colour, shining rather than dull ; the humeral region of the elytra is slightly redder 
than the remainder. 

Eyes situated very far in advance of base of head, very small ; head large in proportion to 
remainder of body, as long as broad ; surface rugosely punctured, the sculpture tending to run 
into longitudinal ridges ; very slightly pubescent ; median line slightly impressed. 

Pronotum as long as broad, slightly narrowed basally ; anterior angles obtuse, posterior so 
obtuse and rounded as to be obsolete ; surface punctured as on head. 

Madeira. 

HoLOTYPE a single specimen in the Wollaston collection (British Museum (Natural 
History)). It has a British Museum type label and " Laemophloeus axillaris, 
type Woll " in a handwriting I do not recognize but which may be that of Arrow. 
I have not recognized further specimens of this species. 

7. Leptophloeus parallelus sp. n. 

Although belonging to the clematidis group of the genus, this species is most likely to be con- 
fused with L. lucidus (Grouvelle) ; the distinguishing characters presented in the key to species 
should prove adequate to prevent any confusion between these two species. However, L. paral- 
lelus is very closely related to L. axillaris (Wollaston) from which it differs in colour and distri- 
bution ; L. axillaris is reddish black in colour and occurs in Madeira whereas L. parallelus is 
brown (albeit dark) and occurs in the Congo. 

Congo. 

HoLOTYPE and i paratype " Luki 5-vi-i952 Dr. Schedl s.136 " (Mus. R. Congo 
Beige). 

8. Leptophloeus mucunae sp. n. J 

This species can be distinguished from its nearest relatives by the characters 
outlined in the key to species. 

i-5-2'0 mm.; body brownish red in colour, moderately shining. 



J 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 207 

Head transverse, i -6 times as wide as long ; epistome concave anteriorly ; antennae rather 
short, hardly reaching base of pronotum, last three segments enlarged forming a loose club, last 
segment at most 1-5 times as long as broad ; surface of disc punctured, punctures twice as large 
in diameter as facet of an eye, separated by a distance equal to less than one diameter ; each 
puncture subtending a small seta ; surface between punctures somewhat irregular (not reticulate 
but tending towards micro-rugosity) and shining. 

Pronotum considerably narrowed basally, maximum width slightly broader than head, 
1-5 times as long along mid-line as broad between hind angles ; anterior angles obtuse, posterior 
obtuse and somewhat projecting ; surface of pronotum as coarsely and closely punctured as on 
head. 

Elytra 2-3 times as long as their combined width, at shoulders narrower than pronotum at 
broadest ; together evenly rounded apically, not leaving last abdominal segment exposed 
dorsally ; the three cells developed, secondary intervals being very narrow ; with punctured 
striae. 

Uganda, Congo. 

HoLOTYPE and 3 paratypes : " Velvet bean seed/Uganda, Kampala -xi-1932 
H. Hargreaves" (British Museum (Natural History)); i "Miss. H. de Saeger 
3787 " (see Appendix) (Mus. R. Congo Beige). 

9. Leptophloeus cassavae sp. n. 

This is a comparatively small species of the genus ranging from i-6-i-9 mm. It resembles 
L. alternans (Erichson) and L. lucidus (Grouvelle) in many ways but differs in that the body is 
somewhat more slender than in those species, thus resulting in, for instance, the pronotum being 
slightly longer than broad ; the epistome is emarginate anteriorly and not truncate ; the punc- 
turation of the head is coarser than the facets of the eyes and not finer ; the antennae resemble 
those of L. alternans rather than those of L. lucidus although the difference between the antennae 
is very slight ; in other respects the three species appear to be identical externally. 

HoLOTYPE and 3 paratypes " Sierra Leone. Njala Ex Cassava branch ix-1935 
E. Hargreaves " (British Museum (Natural History)). 

10. Leptophloeus linearis (Grouvelle) comb. n. 
Laemophloeus linearis Grouvelle, 1908. 

In the key to African Laemophloeinae presented by Grouvelle (1908), this species 
and L. punctatus sp. n. would be identified as L. hypohori (Perris). Nevertheless, 
the differences between them and L. hypobori are considerable. 

The following differences between L. linearis and L. clematidis will serve as a description of the 

1 species : in L. linearis segments of antennae elongate, eyes moderately convex, pronotum 

I broadest just behind anterior angles, elytra as broad across shoulders as pronotum at widest, 

general surface less pubescent than in L. clematidis where antennal segments transverse, eyes 

hardly convex, pronotum broadest at anterior third and elytra broader across shoulders than 

pronotum at widest. 

Congo, Ethiopia. 

HoLOTYPE " Abyssinie/Type/L. linearis A. Grouv." Other specimens seen 
include 7 " Ituri : Djugu ii-1935. H. J. Bredo " ; i " Kivu : NGweshe v-1938 
J. Ghesquiere " ; i " Kivu : Terr. Kabare Nyakasiba, 1800 m (vestige forfit) 
vi-1951 N. Leleup " ; i " N. Kivu : Loashi viii-1937 J. Ghesquiere " ; i " Uganda : 



2o8 L. P. LEFKOVITCH 

Bugiri, 1400 m savane boisee 5/8-viii-i957 P. Basilewsky et N. Leleup " (Mus. R. 
Congo Beige). 

II. Leptophloeus punctatus sp. n. 

(Text-fig. 33) 

This species resembles the preceding, differing only in the following details : in general shape 
body less flattened, almost cylindrical ; elytra about twice as long as their combined width, 
punctures on head and pronotum simple, surface without any rugosity. 

Congo. 

HoLOTYPE " Galerie forestiere de la Kisanga dans humus. Environ d'Elisabeth- 
ville. Septembre 1948 N. Leleup " ; paratypes : i " Recolte dans Thumus (foret 
de Hagenia)/Kivu : Terr Kabare, contrel S.E. Kahuzi, 2080 m i3-viii-i95i N. 
Leleup " ; 2 " Sur Lobelia IKihall-ltun : Mont Bughera 2200 m xi- 1953 R. R. P. P. 
Bergmans-Celis " ; i " Dans inflorescences de Lohelial'^. Kivu : env, Lubero, 
2300 m 6-viii-i953 R. P. M. J. Cells & Coll." ; i " Kwesi & Kilo io/i9-iv-i9ii Dr. 
Bayer " ; i " Ituri La Moto ; Madyu L. Burgeon " ; i " Miss. H. De Saeger 3787 " ; 
I " Miss. H. de Saeger 1724 " (see Appendix) (Mus. R. Congo Beige). 



12. Leptophloeus capitus sp. n. 

This species resembles L. punctatus sp. n. in most respects but differs as follows : antennal 
segments at most as broad as long, median line on the head visible although not deep, punctures 
on the head separated by one or two diameters whereas L. punctatus has the antennal segments 
longer than broad, the median line of the head obsolete and distance between punctures on head 
equal to half to once their diameter. 

South Africa. 

HoLOTYPE " Port St. John, Pondoland Sept. 1923/S. Africa R. E. Turner " and 
I paratype with the same data (British Museum (Natural History)). 

13, Leptophloeus mobilis (Grouvelle) comb. n. 

Laenwphloeus mobilis Grouvelle, 1908. 

Head as broad as long, eyes somewhat projecting ; antennae reaching basal quarter of elytra, 
terminal three segments at most i -5 times as long as broad ; surface of disc with elongate 
punctures separated by narrow ridges, almost rugose ; with fine, narrow longitudinal reticulation 
between the punctures. 

Pronotum just narrower than head across eyes, slightly longer than broad, weakly narrowed 
basally ; lateral line represented by ridge and internal groove, the latter deeper in mid-region 
than anteriorly or posteriorly ; anterior angles obtuse and blunt, posterior angles obtuse and 
rounded ; surface as on head. 

Elytra at shoulders slightly broader than pronotum at its widest, broadening until basal eighth 
and then parallel-sided until apical third, then evenly rounded together ; 2-5 times as long, as 
their combined width ; the three cells developed, secondary intervals being half width of cells. 

East Africa, Congo. 

The single male in the Grouvelle collection is selected as lectotype and is labelled 
" Zanzih. /Type I Laemophloeus mobilis Grouv. ty." There is another syntype in 



A REVISION OF AFRICAN L AEMOPHLOEI N AE 209 

the Deutsches Entomologisches Institut. I have seen i other specimen " Congo 
Beige Hembe-Bitale i9-viii-i952 Dr. Schedl " (Mus. R. Congo Beige). 



14. Leptophloeus bupleuri (Peyerimhof) emend, comb. n. 

Laemophloeus bupleri I'eyeriinhof, 1948. 

This species was called bupleri in the original description but was referred to as 
bupleuri elsewhere in the paper and was named after Bupleurnnt spinosum L. The 
spelhng bupleri is quite clearly an error and is here corrected. 

L. bupleuri belongs to the dematidis group of Leptophloeus but is unique in having the median 
hne of the head raised into a strong ridge. The head is otherwise rugose, shghtly longer than 
broad ; the antennae do not reach the base of the pronotum ; the front angles of the pronotum 
are obtuse, the hind angles so obtuse as to be obsolescent ; pronotum just broader than the head, 
about 1-25 times as long as broad ; elytra at shoulders slightly broader than the pronotum, 
about 2-25 times as long as their combined width, leaving the tip of the abdomen exposed 
dorsally. 

Algeria. 

There are five syntypes, two mounted on one card with the data " Agouni Boussouil 
Djuidjusa vi-vii-1947/Bupleurum spinosum/Laemophloeus Bupleuri Peyerh. 
Types 1-2 " ; two mounted on separate cards but on one pin have data as above 
but " Types 3-4 " ; the fifth is mounted alone and has the same data except for 
" Type 5 ". The specimen implied by Type i is selected as lectotype. 

15. Leptophloeus janeti (Grouvelle) 

(Text-fig. 32) 
Laemophloeus janeti Grouvelle, 1899. 

L. janeti resembles L. capitus sp. n. very closely, both these species differing from L. punctatus 
sp. n. in being yellow rather than chestnut in colour and in the ratio between the length of the 
elytra to the length of the head and pronotum combined being i -4 or less in contrast with the 
figure of i-6 in L. punctatus. In addition to the characters given in the key, L. janeti differs from 
L. capitus in having the secondary intervals and to a lesser extent the raised regions between 
the strial punctures yellow in colour whereas the elytra of L. capitus are a uniform reddish 
yellow. 

Two specimens in the Grouvelle collection, Paris, bear printed data labels as 
follows : " Museum Paris/Congo/Libreville/H. Lecomte 56-97 ". One of these two 
specimens bears the word " type " in Grouvelle 's later handwriting in his usual 
purple ink ; the other has a ms. label " Laemophloeus Janeti Grouv." in Grouvelle's 
early handwriting in black ink. In his original description Grouvelle gave as 
locahty the following information : " Congo : Libreville. Museum de Paris et 
collection A. Grouvelle. — Espece decouverte a Paris dans une noix de Cacao, 
contentant un nid de Mononiorium floricola, envoyee au Museum par M. Lecomte ". 
Consequently, I feel confident that the two specimens indicated above are syntypes. 
I select the specimen bearing the name " Laemophloeus Janeti Grouv." as lectotype. 
The other specimen is rather smaller than is usual for the species. 



210 L. p. LEFKOVITCH 

It is interesting to note that this species, when correctly recognized in stored 
products, has always been associated with cocoa and coffee beans. 
I have seen specimens from throughout the Congo and from Uganda and Ghana. 

The problematicus Species Group (see p. 202) 

Two species belong here, Leptophloeus problematicus sp. n. and L. cornutus sp. n., 
which appear to be unique in the genus in possessing four-segmented tarsi in the 
hind legs of the male. 

16. Leptophloeus problematicus sp. n. 

(Text-fig. 28) 

The following differences from L. janeti (Grouvelle) will serve as a description of this very 
distinct species : colour brown rather than red, puncturation coarse, close ; epistome produced 
into a plate and two horns over labrum. 

Congo. 

HoLOTYPE male and 3 paratypes : " Yangambi i8-vii-i952 Dr. Schedl Nr. 468 " ; 
other paratypes : 7 " Yangambi 15-X-1952 Dr. Schedl Nr. 468 " (Mus. R. Congo 
Beige). 

17. Leptophloeus cornutus sp. n. 

(Text-fig. 29) 

This species resembles L. problematicus sp. n. very closely, differing primarily in the structure 
of its epistome which in L. cornutus has a short sharp median horn contrasting with the plate 
and two horns to be found in L. problematicus. 

Congo. 

HoLOTYPE and i paratype " Yangambi i8-vii-i952 Dr. Schedl Nr. 468 " ; other 
paratypes : 4 " Yangambi 15-X-1952 Dr. Schedl Nr. 468 " ; i " Luki 12. vi. 1952 
Dr. Schedl s.i64a " ; 2 " Luki 5-vi-i952 Dr. Schedl s.136 " ; i " Luki ii-vi-1952 
Dr. Schedl s.195 " ; i " Yangambi 9-ix-i952 Dr. Schedl Nr. 802 " ; i " Sankuru : 
Lodja 10-V-1929 J. Ghesquiere " (Mus. R. Congo Beige). 

IX. NARTHECIUS Leconte 

Narthecius Leconte, 1861. 
Paraphloeus Sharp, 1899 syn. n. 

Type species : Narthecius grandiceps Leconte (by monotypy). 

Body cylindrical. 

Head longer than broad ; epistome acuminate apically, epistome symmetrically or asym- 
metrically produced over labrum into narrow horn ; head longitudinally shallowly grooved on 
each side of horn between horn and insertion of antennae, with a raised ridge on outside in some 
species ; eyes situated well in advance of base, flush with the surface of the head or hardly 
convex, with moderately large facets ; antennae inserted just below lateral carinae at base of 
acumination, with terminal three segments broader than remainder, with scape in some species 
an irregular isoseles triangle in shape, pedicel being inserted at external apical angle, in other 



A REVISION OF AFRICAN LAEMOPHLOEUN AE 211 

species scape almost spherical ; frontoclypeal suture obsolete ; lateral carinae represented by 
raised ridge ; an impressed median line usually present ; gular sutures distinct and well separ- 
ated ; genae moderately produced but negligibly so when compared with Passandridae, 
Prostomis or Passandrophloeus ; labrum transverse, shallowly emarginate anteriorly. 

Pronotum longer than broad, as long as or shorter than head ; suture between the pronotum 
and the pleural sclerites not lateral but lateroventral, sometimes partially visible from above ; 
anterior angles rounded, p>osterior obtuse and well defined ; lateral line represented by raised 
ridge and internal groove. 

Scutellum more or less semicircular ; elytra parallel-sided, about twice as long as their combined 
width, about 1-5 times as long as pronotum ; the three cells developed, secondary intervals being 
as broad as cells or narrower. 

Anterior coxal cavities open posteriorly, situated close together ; intercoxal prosternal process 
convexly rounded posteriorly ; mesosternum very small and consisting of a narrow strut posteri- 
orly between middle coxae ; meso- and metepisterna not contributing to mesocoxal cavities, 
these being formed by meso- and metasterna with mesepimeron ; metasternum equal in length to 
first four visible abdominal sternites combined ; first visible abdominal sternite 1-5 times as 
long as second in mid-line, its intercoxal process narrow, sharply rounded anteriorly ; second 
to fourth sternites about equal to each other, fifth longer but not as long as first. 

Legs rather short, almost warranting description as brachypodous ; trochantero-femoral 
junctions of normal type, femora strongly dilated ; tibiae sinuate ; tarsal formula 5-5-5 in 
both sexes ; in some (American) species the paired apical spines on tibia asymmetrically 
developed on the front legs, symmetrically on the others. 

Paraphloeus Sharp was erected to contain two species from Central America ; 
the type species, P. crassiceps Sharp, quite clearly is a Narthecius, probably con- 
specific with N. grandiceps and is now placed in that genus comb. n. The other 
species, P. longicollis Sharp, belongs to a different group within the Laemophloeinae 
meriting generic status. In contrast with Sharp's opinion, Paraphloeus has five 
tarsal segments in each leg and not four (or three, as he claimed to be present in 
the front legs) . The basal segment is almost entirely withdrawn (in dried specimens, 
at least) into the tibia and is hardly, if at all, shorter than the second ; what little 
protrudes from the shaft is normally hidden by the spurs, spines, knobs and setae 
with which the tibia is ornamented in this region. 

I have neither been able to see the type of nor recognize N. haroldi Reitter, 
described from Madagascar. Its description suggests that it is not Narthecius but 
is more related to the Oriental A^. suturalis Grouvelle which, as indicated below, is 
considered to belong to a genus distinct from Narthecius. N. haroldi would appear 
to be distinguished from the species described below as follows : 

Epistome asymmetrically acuminate anteriorly . . N. schedli sp. n. (p. 211) 

Epistome weakly curved anteriorly ..... N. haroldi Reitter 

Narthecius schedli sp. n. 

(Text-figs. 34, 35) 

About 1-8 mm.; body reddish brown in colour, elytra less intensely so. 

Eyes slightly convex, situated 1-5 times their diameter in advance of base ; surface of head 
with slightly longitudinally elongate punctures, not very much smaller to half the diameter of 
facets of eyes, separated by about one diameter ; surface between punctures minutely poly- 
gonally reticulate ; apparently without setae. 



212 L. P. LEFKOVITCH 

Pronottim about 1-25 times as long as broad, slightly longer than head ; surface as on head ; 
elytra about 2-5 times as long as their combined width, parallel-sided until apical third, there 
gently turning inwards and evenly rounded to apico-sutural angles ; strial punctures very minute, 
each with a thin short seta. 

Congo. 

HOLOTYPE " Congo Beige Yangambi 25-ix-i952 Dr. K. Schedl Nr. 961 " ; para- 
types collected in same locality but on different dates : i, 24-vi-i952 (S.215) ; i, 
30-vi-i952 (S.265) ; 2, 24-ix-i952 (R.944) ; 2. 31-X-1952 (R.949) (Mus. R. Congo 
Beige). 

It is rather difficult to know with which species to compare N. schedli. The 
American members of the genus are very different in their bod}' form and can easily 
be distinguished as a group by the following key. Those from the Oriental region 
which are more closely related are as yet undescribed (A^. suturalis Grouvelle is 
probably generically distinct). The following key will indicate the most obvious 
differences : 

1. Eyes situated three diameters in advance of the base, hardly, if at all, convex ; 

scape large and irregularly triangular in shape ; grooves on each side of the frontal 
horn ridged externally ; epistomal horn centrally placed (Text-fig. 36) 

American NARTHECIUS 
Eyes situated 1-5 diameters in advance of the base, slightly convex ; scape globular ; 
grooves on each side of the frontal horn not ridged externally but merging with the 
general surface of the head ; epistomal horn asymmetrically placed ... 2 

2. Epistomal horn with a narrow base, the head otherwise hardly produced anteriorly 

to the antennal insertions (Text-fig. 37) .... Oriental NARTHECIUS 
- Epistomal horn with a broad base, the head being clearly produced anteriorly to the 

antennal insertions (Text-fig. 35) ...... N. schedli sp. n. 





34 

Figs. 34-37. (34) Dorsal view of Narthecius schedli ; (35) head of N. schedli ; (36) of 
American Narthecius and (37) of Oriental Narthecius. 

Scale lines : Fig. 34 = i mm. ; Figs. 35-37 = 0-25 mm. 



A REVISION OF AFRICAN L AEMOPH LOEI N AE 213 

X. PASSANDROPHLOEUS Kessel 
Passandrophloeus Kessel, 1921. 

Type species : Laemophloeus spinosus Grouvelle (present designation). 

This genus could easily be confused with Leptophloeus but can be easily recognized 
by the ventral structure of the head, the anterior coxal cavities (Text-fig. 38), the 
shape of the intercoxal process of the first abdominal sternite in addition to the 
characters given in the key to genera. 

Of the seven species listed by Kessel in Passandrophloeus, I select spinosus Grou- 
velle as the type species. This species is very similar to the single African species 
redescribed below and may prove to be conspecific. 

Body moderately flattened ; 2-4 mm. 

Head in male triangular in general shape (Text-fig. 40) ; eyes situated in advance of base, a 
distinct ridge present behind eye ; width of head across eyes more than that of pronotum at 
widest ; head in female less obviously triangular in shape (Text-fig. 41) ; eyes almost in contact 
with the base, no distinct ridge behind eye ; width across eyes rarely as much as that of prono- 
tum ; in both sexes, frontoclypeal suture and median line obsolete ; epistome truncate anteri- 
orly ; antennae with scape, pedicel and terminal three segments distinctly larger than other six 
segments ; two lateral lines on each side, inner one depressed, outer one ridged and anteriorly 
forming margin to epistome ; external region of mandibles in male produced ventrally and 
laterally, this production often reaching to beneath produced genae ; in female mandibles not 
produced externally lateroventrally. 

Pronotum with indistinct obtuse anterior angles and with distinct acute posterior angles ; 
two lateral lines on each side, inner consisting of internal shallow groove and external ridge, 
outer essentially a groove between ridge of inner line and a secondary ridge ; this outer groove 
may consist of a row of largish punctures behind mid-line ; the outer ridge is recurved externally 
in front, then runs posteriorly close to lateral margin. 

Scutellum triangular ; elytra carinate laterally, the three cells developed, the carinal fold being 
closely approximated to outer margin of third ; apices of elytra usually truncate in male, with 
external apical angles turned outwards ; apices sometimes describable as truncate in females 
but usually evenly rounded to suture. 

Genae normal in female, in male this region developed ventrally and slightly laterally into a 
heavy jowl on each side (Text-fig. 39) ; gular sutures distinct ; anterior coxal cavities closed 
behind, each with a lateral extension ; intercoxal process of prosternum concave posteriorly ; 
coxae well separated ; metepimera not contributing to mesocoxal cavities ; intercoxal process 
of first abdominal sternite almost straight anteriorly ; metasternum about as long as first three 
abdominal sternites ; first abdominal sternite twice as long as second, second to fourth about 
equal in length to each other, fifth somewhat longer ; trochantero-femoral junctions of the 
normal type ; tarsal formula 5-5-5 in female, 5-5-4 in male. 

Passandrophloeus glabriculus (Grouvelle) comb. n. 
(Text-figs. 39-41) 
Laemophloeus glabriculus Grouvelle, 1897. 
Laemophloeus divergens Grouvelle, 1908 syn. n. 

This species is fairly widespread in the Oriental and African regions and appears 
to be associated with trees of the Dipterocarpaceae. 

About 2 mm.; head, antennae, pronotum and legs yellowish red in colour, elytra essentially the 
same colour but since they are transparent or almost so, they appear to be a dirty white over 
the metathoracic wings. 

ENTOM. 12, 4. 15 



214 



L. P. LEFKOVITCH 




40 



Figs. 38-41. Passandrophloeus glabriculus ; (38) Front and middle coxal cavities ; 
(39) lateral view of head of male ; (40) dorsal view of male ; (41) dorsal view of head 
of female. 

Scale lines : Figs. 38, 39 =- 0-25 mm. ; Figs. 40, 41 = i mm. 



I 



A REVISION OF AFRICAN LAEMOPHLOEIN AE 215 

Punctures on head and disc of pronotum slightly smaller in diameter than facet of eyes, 
separated by once to four times their diameter, each with a seta about two diameters in length. 

Pronotum about as long as broad, narrowed basally ; elytra at shoulders as broad as pronotum 
apically, broadening until just posterior to their mid-point where, in male, they are as broad as 
head across eyes and in females, broader than head ; first strial interval of elytra in addition to 
contributing to first cell, possesses a row of punctures situated quite close to suture ; the three 
cells each with row of punctures in addition to two punctured striae ; secondary intervals very 
narrow (these are presumably what Grouvelle in his original description, called the striae since 
he described each elytron as having three striae). 

In the Grouvelle collection there are three males, which I consider to be syntypes 
of this species : two are mounted together and bear the data " Sumatra Deli /Type 
[black printed on white] /Laemophloeus glabriculus Grouv ty ". The lectotype, 
which I now select, is the male on the right and has been indicated by the figure 2 
on the card. The remaining paralectotype bears the data " Sumatra tabac/Type 
[black printed on white] ". 

The holotype of L. divergens bears the following data ; " Af. or. All. Amani/ 
Eichelbaum/T5rpe [mss] /Laemophloeus divergens sp. nv. Grouv." and a Paris 
museum type label. When examined it was found to be the same species as P. 
glabriculus. Lefkovitch (1957a) gave a record of this species arriving at a British 
port on dunnage from West Africa, which suggested that the species, previously 
known only from Sumatra, occurred in Africa. The following data on the speci- 
mens given below confirms that the species does in fact occur there. 

Congo, West Africa. 

I " Yangambi, 1951 C. Donis z.309 Com. et Bois Congo r.2326 " and 3 other speci- 
mens whose data are identical but for z.307, 306, 316 ; i " s/Grumes Boma (quai) 
X-1945 J. Vrydagh " ; 14 specimens " Eala viii-1935 J. Ghesquiere 708 " ; (Mus. 
R. Congo Beige). Five specimens (author's collection) " Palm kernels loaded 
Apapa 23-i-i957 next to damp, moulding logs ss.Martita collected 18. ii. 1957 
R. C. Welch at Bristol " (these last are the specimens referred to in Lefkovitch, 

1957)- 

Certain other species hitherto placed in Laemophloeus properly belong to this 
genus, namely Passandrophloeus ditomoides (Grouvelle) comb, n., P. spinosus 
(Grouvelle) comb. n. and P. falcidens (Grouvelle) comb. n. 

XL MESTOLAEMUS gen. n. 

Type species : Mestolaemus longicornis sp. n. 

The beetles belonging to this curious genus can be distinguished at once from other 
African Laemophloeinae by the very deep outer emargination on the front of the 
head. 

1-5 mm.; body flattened. 

Head broader than long ; eyes moderate in size, situated just more than one diameter in 
advance of base ; frontoclypeal suture obsolete ; epistome emarginate anteriorly, between 
epistome and insertions of antennae two further emarginations on each side, outermost very 
deep ; lateral raised line present and additional short line present behind eye ; antennae with 
all segments elongate, in total longer than body. 



2i6 L. P. LEFKOVITCH 

Pronotum with lateral line carinate, this line without a fovea along its length. 

5a</e//MW flatly triangular ; e/>'^ya with the three cells ; the usual inflected lateral region typical 
of Laemophloeinae hardly deflected but explanate. 

Genae not produced ; gular sutures distinct ; anterior coxal cavities narrowly closed behind, 
coxae spherical and well separated from each other ; mesepimeron contributing appreciably to 
mesocoxal cavity ; metasternum about equal in length to first two abdominal sternites ; inter- 
coxal process of first abdominal sternite straight anteriorly ; first abdominal sternite twice as 
long as second in mid-line, second to fourth subequal, fifth slightly longer ; in overall shape, 
abdominal sternites strongly convex, abdomen consequently semicylindrical in shape ; legs 
subequal in length ; trochantero-femoral junctions of normal type ; hind femora slightly more 
dilated than others ; tarsal formula 5-5-5 in male, female unknown (probably 5-5-5). 

Mestolaemus longicornis sp. n. 

(Text-fig. 43) 

I •5-2-0 mm.; body very flattened ; reddish yellow in colour. 

Head transverse, about twice as broad as long ; labral emargination shallow ; region between 
outer deep emargination and eye strongly produced above antennal insertion ; scape about 
twice as long as broad, other segments three to four times as long as broad ; all segments of 
antennae about as long as each other, antenna about equal to body in length ; last three segments 
hardly if at all broader than those preceding ; surface closely and evenly punctured, diameter 
slightly less than that of an eye facet, separated by half to one and a half times this distance, 
each with a short, stout seta about equal to one and a half diameters in length ; surface between 
punctures smooth and shining. 

Pronotum at the acute, produced anterior angles as broad as head, twice as broad as long ; 
posterior angles slightly obtuse ; lateral margin (including anterior angles) with about six 
rounded teeth, these being more strongly developed anteriorly ; lateral line formed by a raised 
ridge ; sublateral region explanate ; surface with puncturation and setation as on head. 

Elytra oval in shape, broadest at their mid-point, there slightly broader than pronotum, 
slightly longer than their combined width ; cells with two or three rows of punctures additional 
to striae ; each elytron carinate laterally, sublateral region explanate ; setae as long as those on 
head and pronotum. 

West Africa, Angola. 

HoLOTYPE male " Angola 1493, 18. 2. /Rives Chumbe, 40 km. E. de Dundo, galerie 
for., detritus sol. 15-V-1949. A. de Barros Machado " (British Museum (Natural 
History)) ; i paratype male " IFAN-1946 Tonkoui C.I. 900-1200 m. A. Villiers 
Foret Prim. 20-30. ix " (Museum National d'Histoire Naturelle, Paris). 



XII. MARIOLAEMUS gen. n. 

Type species : Laemophloeus misellus Grouvelle. 

I -5-2 -3 mm.; body only slightly flattened and rather broader in general build than is usual 
in the subfamily. 

Head somewhat transverse ; eyes moderate in size, situated in advance of base ; frontoclypeal 
suture obsolete ; epistome trisinuate ; mouthparts projecting ; antennae elongate, not clubbed, 
segments increasing in length from fourth to eleventh ; third segment longer than either second 
or fourth ; lateral line represented by raised ridge ; a short additional line behind eye present. 

Pronotum transverse, narrowed basally, anterior and posterior angles acute ; lateral line 
represented by raised ridge, broadening laterally at base ; just behind mid-point in sublateral 
region a deeply impressed line with posterior obliquity. 



A REVISION OF AFRICAN L AEMOPHLOEI N AE 



217 



Scutellum transversely p>entagonal ; elytra rarely twice as long as their combined width, 
usually only one and a half times or somewhat less ; first cell strongly margined internally at 
base, other cells weakly developed ; greater part of last abdominal tergite often exposed dorsally. 

Genae normal ; gular sutures distinct ; anterior coxal cavities closed posteriorly, circular but 
coxae very clearly transverse internally ; prosternal intercoxal process concave posteriorly ; 
metepisternum not contributing to mesocoxal cavity ; intercoxal process of first visible abdominal 
sternite slightly curved anteriorly ; metasternum about as long as first two abdominal sternites ; 
first abdominal sternite twice as long as second, second to fourth subequal in length, fifth 
somewhat longer, trochantero-femornl junctions of normal type ; legs subequal ; tarsal formula 
5-5-5 in female, 5-5-4 in male. 

This genus is not unlike Phloeipsius Casey in its general appearance, but the two 
are not really closely related. Mariolaemiis can be distinguished from Phloeipsius 
by the strongly formed frontoclypeal suture and the greater degree of flattening in 
the body. Mariolaemus can be distinguished from other African genera by the 
characters given in the key. 







1 



Figs. 42-47. (42) Front and middle coxal cavities of Mestolaemus longicornis ; 
(43) dorsal view of M. longicornis ; (44) front and middle coxal cavities of Mario- 
laemus misellus ; (45) dorsal view of M. misellus ; (46) apices of elytra of M. escalerae 
and (47) of M. misellus. 

Scale lines : Figs. 42, 44, 46, 47 = 0-25 mm. ; Figs. 43, 45 = i mm, 



2i8 L. P. LEFKOVITCH 

Key to Species 

1. Elytra about I '5 or more times as long as their combined width .... 2 

- Elytra less than i'45 times as long as their combined width ..... 3 

2. Head with a striole at the base of the antennae ; body slightly convex 

I. livens (Grouvelle) (p. 218) 

- Head without a striole at the base of the antennae ; body distinctly convex (blackish 

red in colour, with a shining region at the base of the antennae) 

4. eichelbaumi (Grouvelle) (p. 219) 

3. Elytra of greatest length at their apico-sutural angles (Text-fig. 46) ; abdomen 

usually entirely covered by the elytra ; elytra at their broadest wider than the 
pronotum at its maximum width . . . -3- escalerai (Grouvelle) (p. 219) 

- Elytra of greatest length between their external and their apico-sutural angles and 

the base ; tip of abdomen often exposed dorsally ; elytra at their broadest as 
broad as the pronotum at its widest . . .2. misellus (Grouvelle) (p. 218) 

I. Mariolaemus livens (Grouvelle) comb. n. 
Laemophloeus livens Grouvelle, 1923. 

The length of the elytra distinguishes M. eichelbaumi and M. livens from the other 
two species of the genus ; however, the differences that Grouvelle gave to separate 
M. livens from M. eichelbaumi (the lesser convexity, the striole at the bases of the 
antennae and the sublateral impression on the pronotum) are characters difficult 
to appreciate in the absence of the type specimen (s) ; they have not been discernible 
in any specimen examined by me. 

The type of this species has not been located nor have specimens been found to 
correspond with the original description. Grouvelle stated that it was related to 
M. eichelbaumi most closely and also to M. misellus and M. escalerai. 

East Africa. 

2. Mariolaemus misellus (Grouvelle) comb. n. 

(Text-fig. 45) 
Laemophloeus misellus Grouvelle, 1889. 

1 7-2 -o mm. ; body black with a reddish tinge, tarsi yellow to red in colour. 

Head transverse, i -5 times as wide as long ; a short, shallow groove situated on anterior margin 
of impunctate area at base of antennae, antennae about as long as body ; surface evenly 
punctured (except for area at base of antennae), punctures smaller in diameter than eye facets, 
separated by distance equal to two to five times their diameter ; each with a short seta about 
equal to three diameters in length ; surface between punctures with micro-punctures but 
otherwise smooth and shining. 

Pronotum i -5 times as broad as long, as broad as head, narrowing basally ; disc with punctures, 
setae and surface as on head. 

Elytra slightly broader across shoulders than base of pronotum, broadest at about anterior 
third, as broad as pronotum at widest ; elytra 1-2 times as long as their combined width, each 
with punctured striae ; third cell developed internally and anteriorly ; each separately rounded 
to apico-sutural angle ; last segment of abdomen exposed dorsally. 

There are 2 syntypes in the Grouvelle collection ; I select a specimen with the 
following data as lectotype (it appears to be the specimen figured in the original 
description) : " Assinie Cote occid. Afrique Ch. AUuaud 1886 jType I Laemophloeus 
misellus Grouv," The paralectotype, which is the same species as the lectotype, 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 219 

bears the following data ; " Assinie Afrique oc [printed on green paper] /Type [black 
printed on white] /Laemophloeus misellus Grouv [mss]/Type [black printed on red]." 
This species is one of the commonest of the African Laemophloeinae not associated 
with stored products, and is widespread in its distribution. I have seen specimens 
from Abyssinia, the Ivory Coast, the Congo, Uganda, Angola and South Africa. 

3. Mariolaetnus escalerai (Grouvelle)^ emend, comb. n. 

Laemophloeus escalerae Grouvelle, 1905. 
Laemophloeus subniger Grouvelle, 1905, syn, n. 

This species resembles M. misellus (Grouvelle) very closely but differs as follows : head and 
pronotum with very few if any micropunctures between punctures ; pronotum slightly broader 
than head ; elytra across shoulders as broad as greatest breadth of pronotum, at anterior third 
distinctly broader ; elytra evenly rounded together apically. 

There are 3 syntypes of M. escalarai (Grouvelle) in the Grouvelle collection. I 
select a specimen with the data)" Biaffra Escalera " as lectotype. The other two 
S5mtypes, which are mounted together, bear the data ; " Biafra Cabo S. Juan 
VII-1901 Escalera/Type [mss]/L. Escalerae Grouv. ty. [Grouvelle's mss] " and a 
Paris museum type label. They are conspecific with the lectotype. The holotype 
of L. subniger Grouvelle, whose data are " Assinie Cote occid. Afrique Ch. Alluaud 
1886/L. subniger Gr. ty ined.", is the same species as M. escalerai, 

Congo, West Africa. 

The following specimens have also been seen : 2 " Stanleyville : Yangambi 
i/i6-xii-i953. J. Decelle " ; i " Haut-Uele : Moto vi/vii-1923 L. Burgeon " ; i 
" Coquihatville, 100 km en amout de, sur le fieure k la lumi^re 3-V-1949. N. Leleup "; 
I " Yangambi 2o-ix-i952 Dr. K. Schedl " ; i " Congo Beige Leopoldville L, 
Burgeon 1917 " ; 3 "Ogoone Lambarene R. Ellenberger 1911 " (Mus. R. Congo 
Beige) ; 2 " Yaule/Nimba (Guinee) M. Lamotte 2-vi-i942 " (Museum National 
d'Histoire Naturelle, Paris). 

4. Mariolaetnus eichelbaumi (Grouvelle) comb. n. 

Laemophloeus eichelbaumi Grouvelle, 1908. 

This species resembles both M. misellus and M. escalerai but differs in having no groove or 
stria anterior to the impunctate region at the base of the antennae and in the elytra being more 
than I -5 times as long as their combined width ; it resembles M. misellus in having each elytron 
separately rounded to the apico-sutural angle and M. escalerai in the non-exposure of the last 
tergite of the abdomen. 

East Africa, Congo. 

There are 8 syntypes in the Grouvelle collection ; I select the one with the 
following data as lectotype : " Af. or. all. Amsim/ig /Laemophloeus eichelbaumi 
ined. Grouv." All the other S5nitypes bear the following data : " Afr. or. all Amani ", 
and two additionally : " Eichelbaum " on a separate label. 

Grouvelle describes this species as being common at Amani during the months of 
July to November. Two specimens have been examined other than the syntypes, 

^ The emendation accords with the provisions of Article 31 of the International Code of Zoological 
Nomenclature, 1961, 



220 L. P. LEFKOVITCH 

both from the Belgian Congo. One (British Museum (Natural History)), bears 
the following data: " i8 m. S.W. of Elizabethville 1928 Dr. H. S. Evans " ; the 
other (Mus. R. Congo Beige), " Ehzabethville — ii-1940 H. J. Bredo ". 

XIII. NOTOLAEMUS Lefkovitch 
Notolaemus Lefkovitch, 19596. 

Type species : Cucujus unifasciatus LatreiUe (by original designation). 

2-3 mm.; body moderately flattened. 

Head somewhat wider than long ; eyes moderately prominent, situated in advance of base 
and occupying less than one half of lateral margin of head ; front margin of head with a central 
labral, paired mandibular and often small antennal emarginations ; antennae about equal in 
length in both sexes, approaching length of body ; last three segments not forming a distinct 
club ; lateral line represented by raised ridge ; a subsidiary ridge behind eye present in addition. 

Pronotum transverse, narrowed basally ; anterior and posterior angles moderately prominent 
lateral margin sometimes irregularly rounded ; lateral line represented by raised ridge which is 
somewhat broadened basally. 

Scutellum triangular ; elytra often leaving the tip of abdomen exposed dorsally ; the three 
cells present or absent ; outer margin of third cell when present may or may not coincide with 
lateral carina. 

Genae normal ; gular sutures distinct ; anterior coxal cavities widely open posteriorly ; coxae 
globular ; prosternal intercoxal process slightly convex posteriorly ; metepimera not contributing 
to mesocoxal cavities ; posterior border of mesosternum very slightly convex posteriorly ; 
metasternum about as long as first two abdominal sternites, median line broadening towards 
anterior margin ; intercoxal process of first visible abdominal stemite slightly curved anteriorly ; 
this stemite twice as long as second, second to fifth sterites subequal ; trochantero-femoral 
junctions of normal type although approaching heteromeroid type in middle and hind legs ; 
legs subequal in length, middle femora slightly less dilated than front or hind ; tarsal formula 
5-5-5 in female, 5-5-4 in male. 

Key to African Species 

1. Pronotum with two well separated lateral lines on each side 

I. perrieri (Grouvelle) (p. 221) 

- Pronotum with one lateral line on each side ....... 2 

2. Mandibles of the male broadly produced laterally (Text-fig. 48) . 2. elli sp. n. (p. 221) 

- Mandibles not produced laterally but rounded ....... 3 

3. Hind angles of the pronotum acute or rectangular ...... 4 

- Hind angles of the pronotum obtuse ......... 5 

4. Pronotum with a short, longitudinal deep impression on each side external to the 

lateral carina and adjacent to it at its mid-point (Text-fig. 49) 

3. perspicuus (Grouvelle) (p. 222) 

- Pronotum with a transverse, impressed line between the lateral carina and the lateral 

margin at its mid-point (Text-fig. 50) .... 4. riartus sp. n. (p. 223) 

5. Pronotum excavate sublaterally (Text-fig. 51) ; body brown 5. clarus (Grouvelle) (p. 223) 

- Pronotum flattened sublaterally (Text-fig. 52) ; body brown or black ... 6 

6. Body black and rather dull ..... 8. picinus (Grouvelle) (p. 226) 

- Body brown and rather shining ......... 7 

7. Head with a deep transverse groove on each side at the bases of the antennae ; body 

2-5 mm. in length or more . . . . .6. peringueyi (Grouvelle) (p. 224) 

- Head at most with a shallow fovea on each side at the bases of the antennae, usually 

non-foveate and flat ; body rarely longer than 2-3 mm., usually 2-0 mm. or less in 
length ......... 7. liganus sp. n. (p. 225) 



A REVISION OF AFRICAN LAEMOPHLOEIN AE 221 

I. Notolaetnus perrieri (Grouvelle) comb. n. 

Laemophloeus perrieri Grouvelle, 1906. 

This species differs from other Notolaemus in possessing two lateral lines on each 
side on the head and pronotum, and in the broadness of the head. 

2 mm.; brown in colour ; head and pronotum covered with short, fine setae ; punctures 
almost invisible. 

Head transverse, broader across eyes than pronotum at widest, with two lateral lines on each 
side ; eyes one diameter in advance of base ; antennae about as long as body. 

Pronotum 1-3 times as broad as long, broadest just behind anterior angles ; two lateral lines 
on each side, distance between inner and outer equal to that between outer and lateral margin ; 
anterior angles acute, posterior angles obtuse. 

Elytra 1-5 times as long as their combined width, evenly rounded to apico-sutural angle. 

Female unknown. 

HoLOTYPE male " Madagascar /Perrier /Type /L^^wo^/f/o^Ms perrieri Grouv." 
(Grouvelle collection). A second male, with the following data, has been seen : 
" Zamscolla ix-1952 R.P. Institut Scientifique Madagascar " (Museum National 
d'Histoire Naturelle, Paris). 

2. Notolaemus elli sp. n. 

(Text-fig. 48) 

This species is distinct from other Notolaemus in that the mandibles of the male 
are strongly produced laterally and the external angles ot the central epistomal 









50 



51 



52 



Figs. 48-52. (48) Head of Notolaemus elli ; (49) sublateral region of pronotum of 
A'^. perspicHus and (50) of N. riartus ; (51) transverse section of pronotum oi N. clarus 
and (52) of N. peringueyi. 

Scale line =0-25 mm. 



222 L. P. LEFKOVITCH 

emargination are produced anteriorly, recalling the condition in Planolestes ; how- 
ever, the characters included in the key to genera serve to distinguish this species 
from those of that genus. 

2 -1-2 '5 mm.; head and pronotum dark red, elytra more yellow in colour. 

Head widest across eyes, broader than pronotum ; labral emargination of epistome with 
external angle somewhat produced anteriorly, recalling those of a small male Planolestes ; 
antennae as long as body in males, shorter in females ; surface of disc punctured, punctures 
small, circular and separated by once to three times their diameter, each subtending a seta ; 
mandibles of males often strongly produced laterally resembling a claw-hammer in general 
appearance. 

Pronotum broader anteriorly than posteriorly in males, about same width in females ; surface 
with puncturation and setae as on head. 

Elytra evenly rounded to apico-sutural angles. 

Congo. 

HoLOTYPE male " Maceo (Luebo) 23-ix-i92i Dr. H. Schouteden " ; paratypes : 
I with the same data as the holotype (female) ; 4 " Kamalembi (Luebo) 2i-ix-i92i 
Dr. H. Schouteden " (2 males and 2 females) (Mus. R. Congo Beige). 



3. Notolaetnus perspicuus (Grouvelle) comb. n. 

(Text-fig. 49) 

Laemophloeus perspicuus Grouvelle, 1899. 

This species is most closely related to N. elli sp. n. but can be distinguished from 
it in that the mandibles of the male are not produced laterally and that the produc- 
tion of the external angles of the median epistomal emargination is much less. 

2 -0-2 -5 mm.; shining ; head and pronotum reddish brown in colour, elytra yellower. 

Head in males about as broad as pronotum, somewhat narrower in females ; epistome with 
outer pair of emarginations not developed ; outer margins of the central emargination somewhat 
produced anteriorly ; antennae in males about three-quarters length of body, in females about 
half length of body ; surface of head with small, circular punctures separated by once to three 
times their diameter, this surface smooth. 

Pronotum broadest between anterior angles in males, somewhat behind them in females ; 
anterior angles obtuse and rounded, posterior angles rectangular or just acute and sharp ; 
puncturation and setation as on head. 

Elytra about 1-75 times as long as their combined width. 

Rhodesia, Congo, South Africa. 

There are 4 syntypes in the Grouvelle collection : I select as lectotype a male 
" C. B. Esp. PenngueylTyTpe I Laemophloeus perspicuus Grouv. ty ". The three 
other syntypes have the following data : two " C. B. Esp " ; one " C. B .Esp. /Type 
[black printed on white] /i 04 ". Other specimens seen include 2 " Kunungi 9-iv- 
1921 Dr. H. Schouteden " ; 2 " Ubangi i-ii-1931 M. Leontovitch " ; i " Arebi 
vii-1925 Dr. H. Schouteden " ; i (female) " Haut-Uele : Adra 23-iv-i925 Dr. H. 
Schouteden " (Mus. R. Congo Beige) ; 3 " N.W. Rhodesia Shigariatombwes, 
30-vi-i9i3 H. C. DoUman " (British Museum (Natural History)). 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 223 

4. Notolaemus riartus sp. n. 

(Text-fig. 50) 

Closest to N. perspicuus (Grouvelle) with which it could easily be confused. In 
addition to the transverse line between the lateral carina and lateral margin, A^. 
riartus is a dirty yellow in colour rather than reddish brown, is less convex than N. 
perspicuus and has the sides of pronotum more smoothly rounded. 

i'8-2'3 mm.; moderately shining ; body fuscous yellow in colour, elytra almost transparent. 

Head in males as broad as or broader than pronotum, somewhat narrower in females ; epistome 
with outer pair of emarginations weakly developed, outer margins of the labral emargination 
somewhat produced anteriorly ; antennae almost as long as body in males, shorter in females ; 
surface of head punctured, surface micro-reticulate between them. 

Pronotum broadest behind anterior angles in both sexes ; anterior angles obtuse and rounded, 
posterior angles slightly acute and moderately sharp ; surface ornamented as on head. 

Elytra i- 75-2-0 times as long as their combined width, sometimes leaving tip of abdomen 
exposed dorsally. 

Rhodesia, Congo, Angola. 

HoLOTYPE and 2 paratypes "Angola 4117.15 — Alto Cuilo, Poste de Cacolo, 
gal. forest, missean Tcha-Muchito, 2-vi-54. A. de Barros Machado " ; other para- 
types : 3 "Angola 4119.3 Alto Cuilo (Cacolo) vi-54. A. de Barros Machado"; 
I " Angola 4072.6 — ^Alto Cuilo, Poste de Catolo, rives de la Cavuemba, detr. sol. 
26-V-54. A. de Barros Machado " ; i " Angola 4116.20 (as 4072.6) " ; i " N.W. 
Rhodesia Shigariatombwes 8-vii-i9i3 H. C. Dollman " (British Museum (Natural 
History)) ; 3 " Miss. H. de Saeger 3384 " (see Appendix) ; 2 " Ubangi i-ii-1931 " ; 
I " Arebi vii-1925 Dr. H. Schouteden " (Mus. R. Congo Beige). 

5. Notolaemus clarus (Grouvelle) comb. n. 
(Text-fig. 51) 

Laemophloeus clarus Grouvelle, 1899. 

This species is closest to the European N. castaneus (Erichson). The two species 
differ in many ways and can be most easily distinguished in the region of the 
scutellum ; in N. castaneus the scutellum is darker in colour than the adjacent 
parts of the elytra, whereas in N. clarus it is lighter. 

3 mm.; dark brown in colour ; surface somewhat dull. 

Front of head with five emarginations ; slightly narrower across eyes than pronotum at its 
widest ; surface of head closely punctured, punctures circular, separated by one half of their 
diameter, surface between them with a fine micro-sculpture ; each puncture with a seta about 
equal to one to one and one half times their diameter in length ; antennae as long as body in 
male {female unknown). 

Pronotum somewhat irregularly rounded laterally ; front angles acute, posterior angles 
obtuse ; sublateral region somewhat explanate ; puncturation, setation and micro-sculpture 
as on head. 

Elytra twice as long as their combined width. 

South Africa, East Africa. 

HoLOTYPE " C.B. Esp. Perinta [= VeTm^uey}]ITypelio^ I Laemophloeus clarus 



224 L. p. LEFKOVITCH 

Grouv. ty." One other specimen has been seen, a male " Africa or. Katona/Kili- 
mandjaro x-1904 " (Magyar Nemzeti Museum, Budapest (ex Reitter coll.)). 



6. Notolaemus peringueyi (Grouvelle) comb. n. 
(Text-fig. 52) 
Laemophloeus peringueyi Grouvelle, 1899. 

A very variable species. One of the most characteristic and constant features is 
the diagonal grooving of the disc of the head just anterior to the antennae. These 
grooves appear to be indicative of the frontoclypeal suture, but since the impressed 
median line ends anteriorly in advance of the median junction of these grooves, 
the latter cannot be considered to be homologous with those of Placonotus. 

2"5-3"0 mm.; reddish brown in colour. 

Head as broad across eyes as pronotum at widest ; antennae as long as body in the male, all 
segments elongate ; in female, antennae about three-quarters body length, segments less elongate 
and with last three segments forming a distinct club ; surface of head simply punctured, 
punctures separated by two to three times their diameter, each with a seta about two diameters 
in length ; surface between punctures with micro-reticulation. 

Pronotum, transverse, at its widest 1-3-1 -6 times as wide as long, broadest just behind anterior 
angles, narrowing basally ; anterior angles acute and rounded, posterior obtuse but sharper ; 
surface ornamented as head. 

Elytra twice as long as their combined width, evenly rounded apically, giving elytra a truncated 
elliptical shape. 

Additional characters are given by Grouvelle in his original description of the species. 

South Africa. 

Grouvelle, in his original description, indicated that the syntypes of this species 
were in the Cape Museum and in his own collection. In the South African Museum, 
there is a single specimen (male) with the following data ; " CT 5.86/CT 9.86 Under 
bark/102 [black printed on white] /Laemophloeus Peringueyi Grouv. ty [Grouvelle's 
mss] ". In the Grouvelle collection, there are two specimens which I consider to 
be syntj^es of this species, a male " C.B.Es. Raffray/Type [black printed on white]/ 
Laemophloeus Peringueyi Grouv ty." and a Paris museum type label, and a female 
" C.B. Esp. Peringuey/Type [black printed on white] 105 [black printed on white] " 
and a Paris museum type label. Another specimen in the Grouvelle collection 
which may be a syntype bears the data " C.B. Esp. Rafl[ray " written with a different 
colour ink on differently coloured paper from the other Raffray specimen. There 
is yet another specimen of this species which should be mentioned, which although 
bearing a type label, is not a syntype of any properly described species. Its data 
are " Le Cap Chevrolet [sic] /Type [black printed on white] /Laemophloeus capensis 
Chevr [and an illegible word which might be D'tege] ". This species has never been 
validly published, so far as it has been possible to discover, and is not listed by 
Hetschko (1930). I now select the male belonging to the South African Museum 
to be lectotype. 

I have examined the following specimens of the species : 24 " Port St. John 
Pondoland Sept. 1923/R. E. Turner " ; i " Natal ; Van Reenan, Drakensberg 



J 



A REVISION OF AFRICAN L A EiMOPH LOE I N AE 225 

55-6500 ft. X-1926/R. E. Turner " ; i " Natal : Kloof 1500 ft. Sept. 1926/R. E. 
Turner " ; i " E. Cope Prov. Hogsback 4000 ft. io-x-1932/R. E. Turner " ; 4 " E. 
Cape Prov. Katberg i/i3-xi-i932/R. E. Turner"; 3 " E. Cape Prov. Katberg 
4000 ft i4/26-xi-i932/R. E. Turner " ; 2 " E. Cape Prov. Katberg 4000 ft. xii- 
1932/R. E. Turner " ; 6 " Zululand : Eshowe i-22-iv-926/R. E. Turner " ; i 
" George, Cape Prov. 27-vi to i-vii-1920/ R. E. Turner " i " Orange Free 
State : Witziesnoek. 6100 ft. 24-ii-i929/Dr. Hugh Scott " ; i "Estcourt, Natal. 
G. A. K. Marshall " ; 16 " Stn. No. 59, Deepwalls Forest. Knysna 16-111-1954 under 
bark of Yellow-wood, 1725 ft. Cape Province J. Balfour-Browne, B.M. 1954-797 " ; 
2, as previous 16 but dated 17.iii.1954 and " small pool in glade, 1725 ft." (British 
Museum (Natural History)). 



7. Notolaemus liganus sp. n. 

This species resembles N. peringueyi (Grouvelle) but differs in that it is smaller, 
is a dirty yellow in colour and does not have the diagonal grooving anteriorly on the 
head. 

I •5-2-3 mm.; blackish yellow in colour, moderately shining. 

Head slightly narrower across eyes than pronotum at broadest ; antennae almost as long as 
body in males, segments elongate, ninth segment slightly longer (c. 1-25 times) than tenth; 
in females, terminal three segments slightly more thickened than third to eighth ; surface of head 
with punctures equal in diameter to an eye facet, separated usually by once but sometimes as 
much as twice times their diameter, each with a seta equal to two diameters in length ; surface 
between punctures smooth and shining. 

Pronotum transverse, about i -25 times as broad as long ; moderately convex, broadest at 
about anterior third ; anterior angles rectangular but not sharp, posterior angles obtuse ; 
surface ornamented as on head. 

Elytra 2-0-2-2 times as long as their combined width, together evenly rounded apically, 
broadest at about mid-point. 

Congo, Angola. 

HoLOTYPE and i paratype " Miss. H. de Saeger 4007 " ; other paratypes i " Miss. 
H. de Saeger 3186 " ; 2 " Miss. H. de Saeger 3235 " ; 5 " Miss. H. de Saeger 3296 " ; 
3 " Miss. H. de Saeger 3384 " ; 2 " Miss. H. de Saeger 3519 " (see Appendix) (Mus. R. 
Congo Beige) ; i " Angola 4076.1.5 — Gal. forest, riv. Tcha-Pemba, afil. Luangue, 
Poste de Xa-Cassan, 26-V-54. A. de Barros Machado " ; 4 " Angola 4072. 6- Alto 
Cuilo, Poste de Cacalo, rives de la Caouemba, detr. sol. 26-V-54. A. de Barros 
Machado"; 2 "Angola 4116.20 (as for 4072.6)"; 3 "Angola 4118.9 — Gal. 
Forest, riv. Lunguena, affl. Luangue, Poste Xa-Cassan, 7-vi-54. A. de Barros 
Machado " ; i "Angola 4168.12 Alto Chicapot, dans les mousses d'une cascade, 
24-vi-54. A. de Barros Machado " ; i " Angola 4276.16 — Alto Chicapa, gal. forest ; 
riv. Tchirimbo, detr. sol. 9-vii-54. A. de Barros Machado " ; i " Angola 4125.35 — 
Alto Cuilo, Cacolo, rives ruisseau Na-Ipanha, detr. sol. io-vi-54. A. de Barros 
Machado " ; i " Angola 4370.36 Alto Chicapa, gal. for. riv. Tchimboma, detr. sol, 
i-viii-54. A. de Barros Machado " ; 6 " Angola 4082.15 (as for 4072.6) " (British 
Museum (Natural History)). 



226 L. p. LEFKOVITCH 

8. Notolaemus picinus (Grouvelle) comb. n. 

Laemophloeus picinus Grouvelle, 1923. 

This is a very distinct species, being the only black one of its genus ; it most closely resembles 
the European N. unifasciahts (Latreille) in shape and general body proportions but differs in 
that it is black ; the punctures on the head are about twice the diameter of an eye facet and are 
situated close together. 

N. picinus is the only species of the unifasciatus group known to occur in Africa, 
all the other species of the genus in Africa being closer to N. castaneus (Erichson). 

East Africa, Congo. 

In addition to the holotype, which bears the data " Afr. or Ang. (Kikuyu- 
Esct) Kijabe AUuaud et Jeannel Dec. 1911-2100 m-St. 27 [printed] /Type [black 
printed on pink] /Laemophloeus picinus Grouv." and a Paris museum type label, 
I have seen the following specimens : i " Miss. H. de Saeger 3488 " ; i " Miss. H. 
de Saeger, 4007 " (see Appendix) (Mus. R. Congo Beige). 



XIV. PLANOLESTES Lefkovitch 

Planolestes Lefkovitch, 1958a. 

Type species : Planolestes laevicornis Lefkovitch (by original designation). 

In addition to the characters given in the original description, the following points should be 
noted : head with lateral line represented by raised ridge, median line by shallow groove ; 
pronotum with lateral line represented by a raised ridge ; at about middle third a deep groove 
just internal to ridge ; basally a short, raised secondary ridge internal to primary ridge ; lateral 
line appears to be a double structure in basal third, dividing into two very close to base, one 
part turning abruptly to meet hind angles, other continuing to base ; gular sutures obsolete ; 
anterior coxal cavities circular ; prosternal process convex posteriorly ; mesepimeron and 
mesepisternum contributing to the mesocoxal cavities ; posterior margin of the mesosternum 
straight ; median line of metasternum not quite reaching anterior margin of sclerite ; trochantero- 
femoral junctions of normal type. 

I am indebted to Mr. J. C. Davies for collecting larvae of Planolestes ; these 
have emphasized the generic separation of Planolestes from Cryptolestes. A full 
study of these and other Cucujid larvae wiU be published elsewhere. 

No redescriptions of the species belonging to this genus will be given. 

Key to Species 

1. Pro thorax 1-5 times as broad as long; head of male with clypeal horn strongly 

developed on the left side only, vestigial on the right laevicornis Lefkovitch (p. 228) 

- Prothorax less than 1-5 times as wide as long, rarely exceeding 1-3 times ; head of 

male with clypeal horns strongly developed on both sides . . . . 2, 

2. Head coarsely punctured near the median line, punctures longitudinally elongate, 

separated by about one diameter ; thoracic punctures as those on the head ; setae on 
disc of thorax about equal in length to that of the punctures 

brunneus (Grouvelle) (p. 228) 

- Head with very few punctures near the median line, those present being circular, 

two to four diameters apart ; thoracic punctures as those on the head ; no apparent 
setae on the disc of the thorax .... cornutus (Grouvelle) (p. 228) 



J 



A REVISION OF AFRICAN LAEMOPHLOEIN AE 



227 



Four species were considered by Lefkovitch (1958a) to belong to this genus. The 
placing of one of these, described originally as Laemophloeus hiskrensis Grouvelle 
was based upon a specimen labelled " type " in the Grouvelle collection. It has 
since been compared with the true type in the AUuaud collection and found not to 
be conspecific. In fact L. hiskrensis is a Cryptolestes and is considered in this paper 
under that genus (p. 233). The incorrectly named specimen in the GrouveUe collec- 




FiGs. 53-54. (53) Front and middle coxal cavities of Planolestes cornutus ; (54) dorsal 
view of male P. cornutus. 

Scale lines : Fig. 53 = 0-25 mm. ; Fig. 54 = i mm. 



228 L. p. LEFKOVITCII 

tion has proved to be a rather aberrent Planolestes cornutus (Grouvelle), agreeing 
reasonably with the original description of the species with which it had been 
confused. 

Planolestes laevicornis Lefkovitch 

Planolestes laevicornis Lefkovitch, ig58a. 

South Africa. 

No specimens additional to the type and para types have been located. 

HoLOTYPE and 3 paratypes " Fort Beaufort. C[ape] P[rovince] June 1941. J. W. 
Geyer. With scale Lecaniodiaspis mimosae [(Maskell)] " ; other paratypes as 
follows : 4 " Grahamstown C[ape] P[rovince.] June 1908. C. W. Mally. Emerged 
from Mimosa thorn twig " ; 3 " Okahandja i9-29-xii-i927. S.W. Africa. R. E 
Turner " (British Museum (Natural History)). 

Planolestes brunneus (Grouvelle) 

Laemophloeus brunneus Grouvelle, 1877. 

Congo, East Africa. 

HoLOTYPE " Zanzibar Raffray/Type [mss]/L. brunneus Grouv. ty " and a Paris 
museum type label. 

Grouvelle (1908) expressed the opinion that this and the next species were 
synonymous. Lefkovitch (1958a) showed that they could be distinguished by the 
female genitalia and that both sexes differed slightly externally. The following 
specimens, in additon to the type (which bears no data) have been seen : i " Dimbrey 
C.B. Esp." in the Grouvelle collection ; i " Bushurn iii-1937 J. Ghesquiere ", 7 
" Mongbwalu -vii-1938 Mme. Scheitz " (Mus. R. Congo Beige). 

Planolestes cornutus (Grouvelle) 

(Text-fig. 54) 
Laemophloeus cornutus Grouvelle, 1899. 

This species was described from a single male from Madagascar. The species 
appears to be widespread ; I have seen specimens from Madagascar, from various 
parts of the Congo, Ghana, Angola, Uganda, Nigeria, Bechuanaland, Cape 
Province, Pondoland, Southern Rhodesia, Kenya. 

There is a quite definite association with legume pods. Both Mr. F. Ashman of 
the Department of Agriculture, Kenya, and Mr. J. C. Davies of the Department of 
Agriculture, Uganda, formed the opinion that this species, which they sent to me for 
identification, was attacking legume pods. 

XV. CUCUJINUS Arrow 

Cucujinus Arrow, 1920. 

Type species : Cucujinus micromma Arrow (by monotypy). 

This genus is being revised by Mr. W. O. Steel and therefore I include only a 
generic redescription designed to show its relationship with other African Laemo- 
phloeinae. 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 



229 



Head transverse, with distinct rounded temples, expanding from base ; eyes about or more 
than one diameter in advance of base ; frontoclypeal suture obsolete ; front of head between 
antennae with five emarginations, a shallow labral, paired deep mandibular and a shallow pair 
between the mandibular and insertions of antennae ; antennae about half body length, with a 
robust scape, smaU pedicel, slightly elongate third segment, segments increasing in size from 
fourth to terminal, last three or four segments giving the impression of forming a club but in 
fact not so ; lateral line represented by a distinct groove, these grooves not joined anteriorly 
but basally ; basally at junction of lateral and transverse grooves a diagonal groove running 
from junction to base. 

Pronotum very transverse, usually more than twice as broad as long ; anterior angles distinct 
and acute, hind angles rounded ; lateral margin somewhat irregular ; lateral line represented by 
a distinct groove which does not reach anterior margin, basally curved medially but does not 
continue across to join the other. 








Figs. 55-56. Cucujinus (55) dorsal view ; (56) front and middle coxal cavities. 

Scale lines : Fig. 55 == 2 mm. ; Fig. 56 = 025 mm. 
ENTOM. 12, 4. 16 



230 L. P. LEFKOVITCH 

Scntellum semicircular ; elytra leaving at least last abdominal segment and often one or two 
more exposed dorsally ; each elytron subtruncate, each separately rounded to apico-sutural 
angle ; elytra not carinate laterally and with only a sutural line distinct, other regions of cells 
obsolescent or obsolete. 

Gewfle normal; ^w/ay sutures distinct ; anterior co;rae spherical, their cavities closed posteriorly, 
closure dorsal in relation to rest of cavity ; coxae very widely separated ; intercoxal process of 
prosternum slightly concave posteriorly ; mesocoxal cavities of rather irregular shape, not 
circular ; mesepimeron and metepimeron contributing to mesocoxal cavity ; hind margin of 
mesosternum convex posteriorly ; suture between metasternum and metepisternum curving 
outwards posteriorly, not straight ; median line of metasternum not reaching anterior margin ; 
metasternum about as long as first three abdominal sternites combined ; intercoxal process of 
first abdominal sternite very slightly convex anteriorly, this sternite twice as long as second, 
second to fourth subequal, fifth about as long as the first ; trochantero-femoral junctions of 
normal type ; legs subequal ; femora flattened and dilated in one plane ; tarsal formula 5-5-5 
in female, either 5-5-4 or rarely 5-5-5 in male. 

A species described from Madagascar probably belongs to this genus but differs 
from Cucujinus s. s. especially in the structure of the prothorax. This species, for 
which I propose the subgenus Paracucujinus subgen. n., was described as Laemo- 
phloeus hrevipennis Grouvelle. Since only the type has been seen and since L. coquereli 
Grouvelle from Reunion and one other undescribed species from that island are all 
the specimens I have seen belonging to this subgenus, a full study must wait until 
further material from these islands is available. The following key will serve to 
indicate the principal differences between the two subgenera. 

Key to Subgenera of Cucujinus Arrow 

I. Front of head with the outer emargination on each side shallower than the adjacent ; 
labrum almost triangular in shape ; antennae with terminal segments much thicker 
than the intermediate ; pronotum with rounded anterior and obsolete posterior 
angles ; scutellum rounded posteriorly .... CUCUJINUS Arrow, s. s. 

- Front of head with the outer emargination on each side deeper than the adjacent ; 
labrum almost semicircular in shape ; antennae with the terminal segments hardly, 
if at all, thicker than the intermediate, pronotum with acute, somewhat denticulate 
anterior angles and sharp, obtuse posterior angles ; scutellum angled posteriorly 

PARACUCUJINUS subgen. n. 

The type species of Paracucujinus is Laemophloeus hrevipennis Grouvelle, 1906. 
The following key will separate the two described species belonging to the new 
subgenus, each species of which is known only from the type specimen. 



Key to Species of Paracucujinus subgen. n 

1. Eyes situated twice their diameter in advance of the posterior angles of the head 

coquereli (Grouvelle) comb. n. 
- Eyes situated their diameter in advance of the posterior angles of the head 

hrevipennis (Grouvelle), comb. n. 

Of previously described species hitherto incorrectly placed, Laemophloeus curti- 
pennis Grouvelle and L. nehulosus Grouvelle properly belong to Cucujinus s.s. and 
are now placed in that genus comb. n. 



J 



A REVISION OF AFRICAN L A EMOPH LOE I N A E 231 

XVI. CRYPTOLESTES GaLUglhsLuer 

Leptus Thomson, 1863, nee Latreille, 1796, nee Duftschmid, 1825. 
Cryptolestes Ganglbauer, 1899. 
Fractophloeus Kessel, 1921. 

Type species : Cucujus ferrugineus Stephens (selected by Casey, 1916). 
This comparatively large genus includes some species of economic importance and 
of wide distribution in the world. 

I •5-2-5 mm.; body moderately flattened. 

Head about as broad as long ; eyes of moderate size and situated in advance of base ; epistome 
slightly convex, straight or slightly concave anteriorly ; antennae when about half body length, 
distinctly clubbed, when about as long as body {males of some species) not distinctly clubbed ; 
lateral line represented by a ridge. 

Pronotum very slightly to distinctly transverse, rarely exceeding 1-5 times, usually somewhat 
narrowed basally ; anterior angles distinct and usually obtuse ; posterior angles obtuse to acute, 
usually obtuse, sometimes obsolescent ; lateral line represented by a raised ridge and a closely 
approximated internal shallow groove. 

Scuiellum flatly triangular, approximating to pentagonal in shape ; elytra with the three cells 
developed, secondary intervals being quite narrow ; rows of punctures additional to striae may 
be present in cells. 

Genae normal ; gular sutures obsolescent ; anterior coxal cavities closed behind ; anterior 
coxae spherical ; intercoxal process of prosternum straight or very slightly concave posteriorly ; 
metepimeron just not contributing to mesocoxal cavity ; hind margin of mesosternum straight ; 
metasternum almost as long as first two abdominal sternites combined, median line obsolete 
anteriorly ; first visible abdominal sternite twice as long as second, second to fourth subequal, 
fifth somewhat longer ; trochantero-femoral junctions of normal type (front and middle femora 
grooved ventrally for the reception of the tibiae) ; tarsal formula 5-5-5 in femile, 5-5-4 in 
male. 

A discussion of the problems in correctly identifying some species of this genus is 
given by Lefkovitch (19596). Briefly, the adults show heterogony to such a subtle 
but confusing degree that detailed external description is almost valueless. The 
crucial points required for the identification of the species when using external 
characters alone are given in the key. As will be seen, no attempt is made to 
separate those species referred to as the " pusilloides " group, since it was found that 
constant external characters did not exist. Consequently, identifications of this group 
of species must be made by examination of the genitaha. 

Key to African Species Based on External Characters 

Body black in colour ........... 2 

Body brown in colour ........... 6 

Scape of male equal in length to segments two to four combined and toothed dorsally 

apically 8. planulatus (Grouvelle) (p. 236) 

Scape of male much shorter than segments two to four combined and without any 

dorsal teeth ............ 3 

Hind angles of the pronotum obsolescent (Text-fig. 57) . i. spartii (Curtis) (p. 232) 

Hind angles of the pronotum well developed (Text-fig. 58) .... . 4 

Body more than 2-1 mm. in length, well covered with setae ; males with antennae 

hardly longer than those of the female, reaching the basal third of the elytra 

6. lepesmei (Villiers) (p. 236) 



232 L. p. LEFKOVITCH 

- Body 2 mm. or less in length, with or without setae ; males with antennae much longer 

than those of the female, nearly as long as body ...... 5 

5. Body shining, only sparsely covered with setae . . . 10. atulus sp. n. (p. 238) 

- Body rather dull, thickly covered with setae . .11. atuloides sp. n. (p. 238) 

6. Antennae of males considerably longer than those of the females, often reaching to 

the apices of the elytra or beyond ..." pusilloides " group (pp. 236-240) 
7. pusilloides (Steel & Howe), 9. turcicus (Grouv.), 12. pusillus (Schon.), 
13. ugandae Steel & Howe, 14. evansi sp. n., 15. minimus sp. n. 

- Antennae of males hardly longer than those of the females, reaching to about the 

basal third of the elytra .......... 7 

7. Mandibles of male without tooth on their external margin ; elytra nearly twice as 

long as head and prothorax combined . . . . . . . .8 

- Mandibles of male with a tooth on their external margin ; elytra about 1-5 times as 

long as head and prothorax combined ........ q 

8. Antennal segments hardly longer than broad, last three segments cylindrical 

3. brunneus sp. n. (p. 234) 

- Antennal segments at least twice as long as broad, last three segments flattened 

2. biskrensis (Grouvelle) (p. 233) 

9. Hind angles of pronotum obsolescent . . . .5- capensis (Waltl) (p. 235) 

- Hind angles of pronotum distinct . . . 4. /crrMgincu* (Stephens) (p. 234) 

I. Cryptolestes spar Hi (Curtis) 
(Text-figs. 59, 60) 

Cucujus ater Olivier, 1795, nee Fourcroy, 1785. 
Cucujus piceus Stephens, 1831, nee Olivier, 1795. 
Cucujus Sparta Curtis, 1834. 
Laemophloeus rufipes Lucas, 1849. 
Laemophloeus suffusus Wollaston, 1871, syn. n. 

So far as at present known, this species is confined to Mediterranean Africa, 
Madeira and Europe ; it is included in this study since it seems Hkely to be recorded 
in the future. 

I •8-2-5 mrn.; antennae, mouthparts and legs brownish red in colour, remainder of body 
black ; setae yellowish in colour ; in very immature specimens adults entirely brownish red in 
colour and completely indistinguishable morphologically from C. capensis (Waltl). 

Head in males often as broad as or broader than prothorax, truncated triangular in shape, 
narrower in some males and in all females ; eyes situated in advance of base ; median line present 
basally as an impression ; epistome very slightly concave anteriorly ; antennae reaching to 
just beyond base of elytra in both sexes ; mandibles of males with a tooth on outer margin ; 
surface evenly punctured, punctures about as large as an eye facet, separated by about one to 
three diameters, each with a short seta about two diameters in length ; surface between punc- 
tures smooth and shining at vertex, polygonally reticulate elsewhere. 

Pronotum very slightly transverse, strongly narrowed basally, narrower than head in some 
males but usually broader ; anterior angles slightly acute in broad-headed males, otherwise 
obtuse ; hind angles obsolescent or very obtuse, often indistinguishable from slightly denticulate 
lateral margin of prothorax ; surface as on head. 

Elytra about 2 25 times as long as their combined width ; the three cells developed, secondary 
intervals less than half as broad as cells ; each cell with one row of punctures additional to 
striae ; surface with longitudinal reticulation. 

North Africa, Madeira. 

For a discussion of the synonymy of this species, see Lefkovitch (1959&). It is 
now possible to add Laemophloeus rufipes Lucas to this synonymy. There are two 



A REVISION OF AFRICAN L A EMOPHLOEIN A E 



233 



syntypes in the Lucas collection. An examination of these has enabled me to 
confirm Grouvelle's placing of this species (Grouvelle, 1874). In the last drawer of 
the WoUaston Madeira collection in the British Museum (Natural History), there is a 
single specimen agreeing with WoUaston's description of L. suffusus, which was based 
on a single individual collected by Bewicke. The data of this specimen, which I 
accept as the holotype of L. suffusus, are as follows : on the lower side of the card 
on which the specimen is mounted " Bewicke ", an independent label " Laemophloeus 
suffusus, Woll." in WoUaston's handwriting. It is quite clearly the same species as 
C. spartii (Curtis). 



2. Cryptolestes biskrensis (Grouvelle) comb. n. 

(Text-figs. 61, 62) 

Laemophloeus biskrensis Grouvelle, 1899. 

This large Cryptolestes (2-3 mm.) resembles C. pusilloides in its thoracic shape and in the length 
of the elytra but differs in several other important features. The head is rather foreshortened 
anterior to the antennal insertions and the truncate epistome is shorter in proportion than in 
pusilloides. There is a strong sexual dimorphism in the antennae, mainly connected with the 
fourth to the seventh and ninth to eleventh segments of the antennae which in males increase in 
length with hardly if any increase in width and the distal segment is about four times as 
long as broad ; the last three segments in the female are distinctly broader thim those preceding. 
In their total length the antennae of the male reach to about the basal third of the elytra ; those 
of the female are a little shorter. 



57 





61 




58 






63 

Figs. 57-63. Cryptolestes. (57) Hind angle of pronotum of C. spartii and C. capensis ; 
(58) of C. lepesmei and C. ferrugineus ; (59) sclerites associated with male genitalia of 
C. spartii and C. capensis ; (61) of C. biskrensis (63) of C. brunneus ; (60) sclerotization 
on wall of bursa copulatrix of C. spartii and C. capensis ; (62) of C. biskrensis. 
Scale lines : Figs. 57, 58 = 0-25 mm. ; Figs. 59-63 = o-i mm. 



234 L. P. LEFKOVITCH 

This species occurs north of the Sahara and in Central Africa. It was not possible 
to discover any difference in the specimens from the two localities in spite of the 
improbability of their being conspecific. 

North Africa, Uganda, Congo. 

I have examined the following specimens : northern form : i " Algerie Chellala " ; 
I " Environs de Biskra De Vauloger " ; 3 " Moyen Chari/Fort Archambault/ 
Bongoul (Ba-Kare) /Mission Chari -Tchad /Dr. J. Decorse 1904 Mai ent April" 
I " Biskra Vauloger " ; i " Franchetti /Algerie " (Museum National d'Histoire 
Naturelle, Paris) ; southern form : i " Uganda Katona/Mujenje ix-1913 " ; 
9 " Miss. H. de Saeger 63, 578 1803, 1157, 2172, 3012, 3519, 3567, 4057 " (see 
Appendix) (Mus. R. Congo Beige). 

3. Cryptolestes brunneus sp. n. 

(Text-fig. 63) 

I-8-2-5 mm. Resembling C. biskrensis but distinguished easily by its darker colour, being a 
blackish brown, and its almost quadrate prothorax, and the antennal segments which are 
hardly longer than broad. 

Congo. 

HoLOTYPE and i paratype (males) " Miss. H. de Saeger, 3341 " (see Appendix) 
(Mus. R. Congo Beige). 

4. Cryptolestes ferrugineus (Stephens) 
(Text-figs. 64, 65, 66). 

Cucujus tesiaceus PaykuU, 1799, nee Fabricius, 1787. 
Cucujus ferrugineus Stephens, 1831. 
Cucujus monilicornis Stephens, 1831. 
Laemophloeus concolor Smith, 1851. 
Laemophloeus obsoletus Smith, 1851. 
Laemophloeus carinulatus WoUaston, 1877, syn. n. 
Laemophloeus emgei Reitter, 1887. 
Laemophloeus alluaudi Grouvelle, 1906, syn. n. 

This species resembles C. spartii (Curtis) very closely, differing externally in that the body is 
brown in colour, in the hind angles of the pronotum being strongly developed, in the elytra 
possessing two rows of punctures in each cell in addition to the striae and in the epistome being 
ver>' slightly convex. 

The holotype of Laemophloeus alluaudi Grouvelle in the Alluaud collection has 
been examined and found to belong to this species. It is somewhat more pubescent 
than is typical of the species but not unusually so. There are five specimens of 
Laemophloeus carinulatus Wollaston in his collection from St. Helena in the British 
Museum (Natural History). One, a male, has the data " Laemophloeus carinulatus, 
type Woll." and a British Museum type label ; the others, which are females, have 
no data. Wollaston noted that he originally had seven specimens of the species ; 
I have been unable to locate the remaining pair. The five, which I consider to be 
syntypes, are the same species and I now select the male to be lectotype. I have no 
hesitation in synonymising this species with C. ferrugineus, of which WoUaston's 



A REVISION OF AFRICAN L AEMOPHLOEIN A E 



235 



specimens are normal examples ; WoUaston, commenting on his species, wrote that 
it was probably introduced into St. Helena. 

C. ferrugineus is cosmopolitan on stored foods and has also been found under 
bark, in soil and in most habitats where moderately dry plant material occurs. It 
can live entirely upon plant material but is facultatively predatory and cannibalistic. 



5. Cryptolestes capensis (Waltl) 

(Text-figs. 59, 60) 

Cucujus capensis Waltl, 1834. 
Laemophloeus elongatulus Lucas, 1849, syn. n. 
Laemophloeus vermiculatus WoUaston, 1854, syn. n. 
Laemophloeus clavicollis WoUaston, 1854, syn. n. 

This species differs from C. spartii (Curtis) morphologically in one point only ; it is brown in 
colour in the mature adult and not black (see Lefkovitch, 1959a for a biological study of the 
relationship between these two species). 

Europe, South Africa (associated with stored food). 

Previously (Lefkovitch, 19596), I stated that the specimens called Laemophloeus 
elongatulus Lucas that I had seen in European collections of Coleoptera were equally 
divided between large C. ferrugineus and C. capensis. Since then I have examined 
the fifteen syntypes. I select the specimen with a blue label and the words " Laemo- 
phloeus elongatulus Lucas " as lectotype. The species is morphologically identical 
with C. capensis. 



t 





65 





Figs. 64-67. Cryptolestes. (64) Dorsal view of C. ferrugineus ; (65) sclerites associated 
with male genitalia of C. ferrugineus ; (66) sclerotization on wall of bursa copulatrix 
of C. ferrugineus ; (67) of C. lepesmei. 

Scale lines : Fig. 64 = i mm. ; Figs. 65-67 = o- 1 mm. 



236 L. P. LEFKOVITCH 

I have also examined the syntypes of Laemophloeus clavicoUis Wollaston and 
L. vermiculatus Wollaston in the British Museum (Natural History) collection and 
found them to be identical with C. capensis. 

6. Cryptolestes lepesmei (ViUiers) comb. n. 

(Text-fig. 67) 
Laemophloeus lepesmei ViUiers, 1940. 

This species resembles C. spartii very closely but differs externally in that the hind angles of 
the pronotum are as well developed as in C. ferrugineus. In size C. lepesm,ei is larger than C. 
spartii, varying between 2-i-2--j mm. 

In his original description, ViUiers wrote that there was a type and six co types. 
On examining the specimens, I found that no type had been labelled as such nor 
was any indication given in the description of any particular specimen as type. 
In discussion, Mons. Villiers informed me that he had no particular specimen in 
mind to be holotype. I select a male as lectotype, but not that specimen illustrated 
by ViUiers since it is distorted on the left side. None of the specimens bear data. 

Cameroons, Ethiopia. 

In addition to the 7 syntypes, I have recognized 3 specimens of this species in 
the British Museum (Natural History) collection. They bear the following data : 
" Under bark of decaying Mimosa. Abyssinia : Djem-Djem forest. Nearly 9,000 ft. 
1-X-1926. Dr. H. Scott ". 

7. Cryptolestes pusilloides (Steel & Howe) 
(Text-figs. 68, 69) 

Laemophloeus pusilloides Steel & Howe, 1952. 

i-8-2'2 mm.; body reddish brown in colour, the elytra in immature specimens being somewhat 
paler than the rest of the body. 

Head about as broad as or slightly narrower than pronotum, slightly transverse ; epistome 
very shallowly concave anteriorly ; antennae as long as body in male, reaching the basal third 
of elytra in female ; surface punctured, punctures equal in diameter to facet of eye, separated 
by one to three times this distance, each with a seta equal to three to five times this diameter. 

Pronotum transverse, about i- 1-1-2 times as broad as long, narrowing basally ; anterior angles 
obtuse and rounded, posterior rectangular and sharp ; puncturation and vestiture as on head. 

Elytra hardly broader than pronotum, about twice as long as their combined width ; the 
three ceHs developed, each with one row of punctures in addition to two striae. 

The geographical distribution of this species is discussed by Howe & Lefkovitch 
(1957). Briefly, C. pusilloides occurs in southern and eastern Africa, Australia 
and South America and is associated with stored food. 

8. Cryptolestes planulatus (Grouvelle) comb. n. 
Laemophloeus planulatus Grouvelle, 1896. 

This species resembles C. pusilloides in its general shape but differs as follows : pronotum 
somewhat more narrowed basally ; anterior and posterior angles acute ; scape in male equal in 



A REVISION OF AFRICAN L AEMOPH LO E I N A E 



237 



length to segments two, three and four combined, thickened distally, and dorsally produced into 
a tooth above the thickening. The scapes of Laemophloeinae are discussed by Lefkovitch 

(19586). 

Congo. 

HoLOTYPE female " i. Bourbon Fairm./type/L. planulatus Grouv ty ". 

Two colour forms exist, a brown (the typical form) and a black. The following 
specimens in addition to the holotype have been seen : black form : " Yangambi 
1951 C. Donis z.ii77/Com. Et. Bois Congo R.2394 " ; " Yangambi 1951 C. Donis 
z.3o6/Com. Et. Bois Congo R. 2326 " ; brown form : 28 " Miss. H. de Saeger, 928, 
1214. 2007, 3186, 3384. 3519, 3649, 3738, 3744, 4007 " (see Appendix) (Mus. R. Congo 
Beige). 

9. Cryptolestes turcicus (Grouvelle) 

(Text-figs. 70, 71) 

Laemophloeus turcicus Grouvelle, 1876. 

This species differs externally from C. pusilloides principally in the shape of the pronotum, 
which in C. turcicus is almost quadrate and has projecting acute posterior angles. The pronotum 
may be narrowed slightly basally in large specimens somewhat as in C. pusilloides but the 
epistome is slightly convex and never concave anteriorly. 

Temperate regions of the world except Australia and New Zealand ; associated 
with stored food. 




\r-\ 



68 





70 



72 




71 




73 



Figs. 68-73. Cryptolestes. (68) Sclerites associated with male genitalia of C. ^M5?7/oirfe5 ; 
(70) of C. turcicus ; (72) of C. atulus ; (69) sclerotization on wall of bursa copulatrix 
of C. pusilloides ; (71) of C. turcicus ; (73) of C. atulus. 

Scale line = o-i mm. 



238 L. P. LEFKOVITCH 

C. kircicus is not known to breed in Central Africa but has been recorded from the 
Belgian Congo and Uganda (Howe & Lefkovitch, 1957). Additional records : 
3 " C.T. 4.86/C.B. Esp. Peringuey/in a packet of Cocoa " in the Grouvelle collection, 
Paris ; 3 specimens " Flour mill. Cape Town M. Howroyd 1959 " (author's collection). 

10. Cryptolestes atulus sp. n. 

(Text-figs. 72. 73) 

I-8-2-2 mm. This species resembles C. iurcicus but is black and much more shining than that 
species ; the legs, antennae, mouthparts and the impunctate area at the base of each antenna 
are reddish in colour. There are only a few short setae on the head and pronotum. 

S. Thome, Congo. 

HoLOTYPE male " Tshuapa : Bokuma ii/iii-1954 R. P. Lootens " ; paratypes 
as follows : i " Ubaugi : Gemena -xi-1935 P. Henrard " ; 1 " Barumbi -vii-1925. 
fruit ficus pressee. Lt. J, Ghesquiere " ; i " Haut-Uele : Mauda -iii-1925. Dr. H. 
Schouteden " ; i " Luebo -viii-1921. coton. Lt. Ghesquiere " ; i " Luebo 25-viii- 
1921. Dr. H. Schouteden " ; 2 " Eala -iii-1935 J. Ghesquiere " ; i " I.R.S.A.C. 
Tshuapa ; Terr. Bikoro, 350 m. Lac Tumba N. Leleup -x-1955 " ; i " Ruanda : 
Kisenyi 1500 m. 2i-viii-i953 A. E. Bertrand " ; i " N'Gazi i4-vii-i952 " ; i 
" Yangambi 5-viii-i952 " (Mus. R. Congo Beige) ; 3 " S. Thome " (Museum 
National d'Histoire Naturelle, Paris). 

II. Cryptolestes atuloides sp. n. 

(Text-figs. 74, 75) 

I •8-2-3 nim. This species is very closely related to C. atulus sp. n. but differs in that it has longer, 
more abundant pubescence on the head and pronotum. In shape, the pronotum is closer to 
that of C. ferrugineus rather than C. iurcicus, being rather more narrowed basally and without 
such strongly formed hind angles. The setae are about twice as long as the intervals between 
the punctures. 

Congo. 

HoLOTYPE male " Luebo -viii-1921 Lt. Ghesquiere. Bois mort " ; allotype and 
3 other paratypes with the same data as the holotype. Other paratypes : 6 " Eala 
-vii-1935 J. Ghesquiere " ; 2 " Luebo i8-viii-i92i Dr. H. Schouteden " ; 3 " Haut 
Uele : Moto 1920 L. Burgeon " ; i " Yangambi 23-vi-i952 Dr. Schedl s.2i3a " ; 
I " Yangambi 3-vii-i952 s.296 " ; brown specimens externally resembling C. evansi 
sp. n. ; " Kamalembi (Luebo) 2i-ix-i92i. Dr. H. Schouteden " ; " Kasal : Mukumbi 
18-X-921. Dr. H. Schouteden " (Mus. R. Congo Beige) ; i " Belgian Congo 18 m(iles) 
S.W. of Ehzabethville 1927. Dr. H. S. Evans " (British Museum (Natural History)). 

12. Cryptolestes pusillus (Schonherr) 
(Text-figs. 76, 77) 

Cucujus minutus Olivier, 1791, nee Fourcroy, 1785. 
Cucujus pusillus Schonherr, 181 7. 
Cucujus testaceus Stephens, 1831. 



A REVISION OF AFRICAN L AEMOPHLOEI N AE 



239 



Cucujus crassicornis Waltl, 1839. 
Laemophloeus longicornis Mannerheim, 1843. 
Losmophloeiis (sic) brevis Fairmaire, 1850. 
Laemophloeus paralleliis Smith, 1851. 
Laemophloeus pauper Sharp, 1899. 

This species resembles C. pusilloides in many ways and since they both occur in stored products, 
they have often been confused with each other. C. pusillus is much more pubescent, the pronotum 
more transverse (often as much as 1-4 times), the elytra are comparatively shorter in relation to 
the rest of the body, and possess two rows of punctures in the cells in addition to the striae. 
Small specimens of the two species and of C. turcicus and C. ugandae are virtually indistinguish- 
able from each other externally. 

I have seen specimens of this species from almost all parts of Africa ; associated 
with stored food. 

13. Cryptolestes ugandae Steel & Howe 
(Text-figs. 78, 79) 
Cryptolestes ugandae Steel & Howe, 1955. 

Externally, this species is somewhat intermediate between C. pusillus and C. pusilloides. It 
has been impossible to find external characters which are really reliable. The epistome is slightly 
more emarginate than in those species, the anterior margin of the pronotum is just sinuate 
before the anterior angles and the puncturation on the head and pronotum is finer than is usual 
in those two species. But all these species vary so much externally that no real reliance can be 
placed upon these characters. 

The distribution of this species is given by Lefkovitch (1957&) and by Howe & 
Lefkovitch (1957). It appears to be confined to Central Africa, associated with 
stored food. 







74 



76 



78 





Figs. 74-79. Cryptolestes (74) Sclerites associated with male genitalia of C. atuloides ; 
(76) of C. pusillus ; (78) of C. ugandae ; (75) sclerotization on wall of bursa copu- 
latrix of C. atuloides ; (77) of C. pusillus (spermatheca also illustrated) ; (79) of 
C. ugandae. 

Scale line = o-i mm. 



240 



L. ]' 



L E F K O V I r C H 



14. Cryptolestes evansi sp. n. 

(Text-fig. 80) 

i'8-2'2 mm. This species differs from C pusilloides in that the pronotum is less transverse 
being scarcely broader than long and of shape similar to that of a small female C. iurcicus. In 
the length of the body and in its other proportions the species is identical with C. pusilloides. 

Congo. 

HoLOTYPE male " Belgian Congo 18 m(iles) S.W. of Elizabethville 1928 Dr. H. S. 
Evans " ; 2 paratypes with the same data except as follows : male dated 13. xi. 1927 
and female dated 13. ii. 1928 (British Museum (Natural History)). 

15. Cryptolestes minimus sp. n. 

(Text-figs. 81, 82) 

I -8-2 -3 mm. Resembling C. pjtsilloides very closely but differing in the more sharply defined 
posterior angles to the prothorax, in the relative broadness of the elytra in comparison with the 
prothorax (it is difficult to give a value to this difference) and in the yellowish rather than reddish 
brown colour of the body. 

Angola. 

HoLOTYPE male "Angola 4211.9 Alto Chicapa, sous ecorce arbre tombe 27-vi- 
1954. Ed. Luna de Carvalho " ; 4 paratypes (2 males and 2 females) with the same 
data as the holotype ; 3 paratypes " Angola 1824.4.3 Serra do Moco, Luimbale 
(Huambo), sous ecorce arbre tombe, io-ix-1949. A. de Barros Machado " ; i 
" Angola 4194.5 Alto chicapa, gal. forest, riv. Ngungo, dans tronc pourri 27-1-1954. 
Ed. Luna de Carvalho " (British Museum (Natural History)). 






80 



8 



82 



Figs. 80-82. Cryptolestes (80) Sclerotization on wall of bursa copulatrix of C. evansi 
(82) of C. minimus ; (81) sclerites associated with male genitalia of C. minimus. 

Scale line = o-i mm. 



ACKNOWLEDGEMENTS 

I am grateful to the authorities and staffs of the many museums and institutions 
who have lent me specimens or given me hospitality for the furtherance of this 
study. I wish, in particular, to mention the British Museum (Natural History) 
whose staff have given considerable assistance, almost weekly, during the last four 
years. 



A REVISION OF AFRICAN L AEMOPHLOEIN AE 



241 



I am indebted to Miss C. M. F. von Hayek, Dr. E. B. Britton and Mr. R. D. Pope 
for the time they have given in the discussion of various points that have arisen 
and especially to Mr. J. A. Balfour-Browne for the care with which he has read the 
manuscript. 

APPENDIX 

Data relating to specimens collected in the Pare National de la Garamba in the Congo by the 
Mission H. de Saeger and referred to in the text by a number (see also de Saeger, 1956). 

Reference 

Data 
Submerged vegetation. 
Savanna with trees. 
Alighting on fruits of Ficus 

capensis. 
At edge of river in high grass. 
Between burned savanna and 

river in short grass. 
Collected in leaves of Caloncoba 

welwiischii. 
Shaded pool on Canthium cf. 

hispidum. 
Under bark of Cassia siamea 

imported for building. 
Under bark of dead tree, wood 

moist and partially decayed. 
Very moist decomposing wood. 
From grass in savanna. 
Dead trees and branches on 

ground in degenerate forest. 
Savanna, on left bank of Nam- 

bira. 
In fallen trees, decay hardly 

begun. 
In dead trunks of Neoboutonia. 
Savanna, in dry stems of Nauclea 

latifolia killed by fire in Janu- 
ary. 
In stems and bracts in a mass of 

Oxytenanthera abyssinica. 
Under trunk, in spaces between 

the bark of Anogeissus schim- 

peri. 
Savanna with occasional trees ; 

in fallen dead branches. 
Savanna with Lophira trees. 
Savanna with shrubs ; fallen 

dead branches. 
Thin forest ; fallen dead branches. 
Savanna ; in flowers, mainly 

Setaria sphacelata, Sporobolus 

pyramidalis and various Cyper- 

aceae. 



Number 


Collector 


Date 


63 
578 
928 


H. de Saeger 
G. Demoulin 
H. de Saeger 


21.xii.1949 
5.vi.i95o 
2.xi.i950 


1157 
1214 


J. Verschuren 


25.1.1951 
3.11.1951 


1724 


H. de Saeger 


14.V.1951 


1803 


„ 


25.V.1951 


2007 


J. Verschuren 


29. vi. 195 1 


2062 


H. de Saeger 


13.vii.1951 


2115 
2172 
2601 


" 


19.vii.1951 
30.vii.1951 
15.X.1951 


3012 


J. Verschuren 


16.1.1952 


3186 


H. de Saeger 


12.iii.1952 


3235 
3296 


.. 


13.iii.1952 
7.iv.i952 


3341 


>> 


19. iv. 1952 


3379 


»i • 


20. iv. 1952 



3384 

3488 
3519 

3563 
3567 



28. IV. 1952 

20. V. 1952 
26. V. 1952 

3.vi.i952 
30. V. 1952 



242 



L. p. LEFKOVITCH 
A P P E N D I X—cont. 



Reference 






Number 


Collector 


Date 


3649 


H. de Saeger 


18. vi. 1952 


3736 


" 


4.VI1.1952 


3738 


11 


4.vii.i952 


3744 


" 


7.VI1.1952 


3787 


.. 


15.vii.1952 


4007 


.. 


i.ix.1952 


4057 


,, 


1 6. ix. 1952 



Data 

Fallen dead tree on stony rubbish. 

Temporary pool edges, in decay- 
ing vegetation. 

Dead tree of Irvingia smithii 

Dead branches of Erythrophloeum 
Chlorophora. 

In dead, fallen trees of Voacanga 
obtusa. 

Savanna, in dead branches of 
woody shrubs. 

Marsh, in plant layer [Jussiaea). 



REFERENCES 

References to species and genera mentioned in the text but not included in this bibliography are 
given by Hetschko (1930). 

Casey, T. L. 1884. Revision of the Cucujidae of America North of Mexico. Trans. Amer. 

ent. Soc. 11 : 66-112. 
1916. Some random studies among the Clavicornia. In Memoirs on the Coleoptera 

7 : 35-300. 
Grouvelle, a. 1874. Ann. Soc. ent. Fr. (5) 4 : xxix. 
1908. Coleopteres clavicornes de I'Afrique Australe et Oriental. Rev. ent. {Caen) 27 : 127- 

192. 
Hetschko, A. 1930. Cucujidae in Junk, W., Coleopterorum Catalogus 109 : 1-122. 
Howe, R. W. & Lefkovitch, L. P. 1957. The distribution of the storage species of Crypto- 

testes (Col. Cucujidae). Bull. ent. Res. 48 : 795-809. 
Kessel, F. 1 92 1. Neue Monotomiden, Cucujiden u. Passandriden aus der Sammlung des 

Deutschen Entomologischen Museums in Berlin. Arch. Naturgesch. 87 : 25-33. 
Lefkovitch, L. P. 1957a. Further records of Laemophloeinae (Col. Cucujidae) in stored 

products. Ent. mon. Mag. 93 : 239. 
ig^yb. The biology of Cryptolestes ugandae Steel and Howe (Coleoptera, Cucujidae), a 

pest of stored products in Africa. Proc. zool. Soc. Lond. 128 : 419-429. 
1958a. A new genus and species of Laemophloeinae (Col. Cucujidae) from Africa. Ibid. 

93 : 271-273. 
19586. Unusual antennal characters in some Laemophloeinae (Coleoptera : Cucujidae) 

and their taxonomic importance. Proc. R. ent. Soc. Lond. (B) 27 : 93-100. 
1959a. Biological evidence for the specific separation of Cryptolestes capensis (Waltl) 

from C. spartii (Curtis) (Coleoptera : Cucujidae). Proc. R. ent. Soc. Lond. (A) 34 : 44-48. 
1959^- A revision of the European Laemophloeinae (Coleoptera : Cucujidae). Trans. R. 

ent. Soc. Lond. Ill : 95-118. 
1962. The taxonomic position of Laemophloeus raffrayi Grouvelle. Verhand. XI. Int. 

Kongr. Ent. 1 : 78-80. 
Peyerimhof, p. de. 1948. Description d'un Laemophloeus [Col. Cucujidae] algerien 

commensal de Triotemnus Grangeri Peyerh. [Scolytidae] . Bull. Soc. ent. Fr. 53 : 97-99. 
Saeger, H. de. 1956. Entomologie Renseignements Eco-biologiques. Explor. Pare. Nat. 

Garamba, 5 : 1-555. 
Steel, W. O. (in preparation) (A revision of the genus Cucujinus Arrow sensu stricto). 
Steel, W. O. & Howe, R. W. 1952. A new species of Laemophloeus (Col. : Cucujidae) 

associated with stored products. Proc. R. ent. Soc. Lond. (B) 21 : 86-88. 
1955- A new sp>ecies of Cryptolestes (Col. : Cucujidae) associated with stored 

products. Ibid. 24 : 107-109. 
ViLLiERS, A. 1940. In Paulian, R. & Villiers, A., Les coleopteres des Lobelias de montagnes. 

Rev. franc. Ent. 7 : 72-83. 



I 



INDEX 



243 



INDEX OF LAEMOPHLOEINAE 
(Synonyms in italics ; bold page numbers refer to descriptions, tliose in italics to a mention in a key) 



africanus sp. n., Placonotus, lyj, 181, 182 
alluaudi Grouvelle, Laemophloeus, 234 
alternans (Erichson), Leptophloeus, 168, 202, 

204, 206, 207 
ambiguus (Grouvelle), Gannes, 174, 175 
ambiguus Grouvelle, Laemophloeus, 175 
angustulus Leconte, Laemophloeus, 201 
anormus Grouvelle, Laemophloeus, 205 
anormus (Grouvelle), Leptophloeus, 205 
ater Olivier, Cucujus, 232 
ater sp. n., Leptophloeus, 203, 204, 205 
atuloides sp. n., Cryptolestes, 232, 238, 239 
atulus sp. n., Cryptolestes, 232, 237, 238 
axillaris WoUaston, Laemophloeus, 206 
axillaris (WoUaston), Leptophloeus, 203, 205, 

206 



bimaculatus sp, n., Xylophloeus, 189, igi, 

193 
biskrensis (Grouvelle), Cryptolestes, 232, 233, 

234 
biskrensis Grouvelle, Laemophloeus, 227, 233, 
Blubos gen. n., 772, 194 
bolivari Grouvelle, Laemophloeus, 185 
bolivari (Grouvelle), Placonotus, 178, 185 
brevipennis (Grouvelle), Cucujinus (Para- 

cucujinus), 230 
brevipennis Grouvelle, Laemophloeus, 230 
brevis Fairmaire, Loemophloeus (sic), 239 
brightensis Blackburn, Lathropus, 197 
brunneus sp. n., Cryptolestes, 232, 233, 234 
brunneus Grouvelle, Laemophloeus, 228 
brunneus (Grouvelle), Planolestes, 226, 228 
bupleri (sic) Peyerimhof, Laemophloeus, 209 
bupleuri (Peyerimhof), Leptophloeus, 203, 209 

capensis (VValtl), Cryptolestes, 170, 184, 232, 

233. 235 
capensis Waltl, Cucujus, 235 
capensis Chevr. {sic), Laemophloeus, 224 
capitus, sp. n., Leptophloeus, 204, 208, 209 
carinulatus WoUaston, Laemophloeus, 234 
cassavae sp. n., Leptophloeus, 204, 207 
castaneus (Erichson), Notolaemus, 168, 223, 

226 
chrysomeloides sp. n., Xylophloeus, igi, 192 
clams Grouvelle, Laemophloeus, 223 
clarus (Grouvelle), Notolaemus, 220, 2 21,223 



clavicollis WollsiSton, Laemophloeus, 184, 235, 

236 
clematidis (Erichson), Leptophloeus, 168, 

202, 206, 207 209 
concolor Smith, Laemophloeus, 234 
coquereli (Grouvelle), Cucujinus (Paracucu- 

jinus, 230 
coquereli Grouvelle, Laemophloeus, 230 
cornutus Grouvelle, Laemophloeus, 228 
cornutus sp. n., Leptophloeus, 202, 203, 210 
cornutus (Grouvelle), Planolestes, 226 
corticinus (Erichson), Cryptolestes, 168 
crassiceps (Sharp), Narthecius, 211 
crassiceps Sharp, Paraphloeus, 211 
crassicornis Waltl, Cucujus, 239 
Cryptolestes Ganglbauer, 168, 170, iy3, 184, 

187, ig6, 226, 227, 231 
Cucujus (S.I.), 220, 231, 232, 234, 235, 238 
Cucujinus Arrow, 168, 169, 172, 228, 230 
curtipennis (Grouvelle), Cucujinus, 230 
curtipennis Grouvelle, Laemophloeus, 230 

decoralus Grouvelle, Laemophloeus, 186 
decoratus (Grouvelle), Placonotus, J77, 186 
dentatus sp. n., Xylophloeus, igi, 194 
ditomoides (Grouvelle), Passandrophloeus, 

215 
divergens Grouvelle, Laemophloeus, 213 
dolce sp. n., Placonotus, lyj, 178, 179 
donacioides WoUaston, Laemophloeus, 183 
donacioides (WoUaston), Placonotus, 177, 183, 

187 
duplicatus (Waltl), Cryptolestes, 168 

ealaensis sp. n., Placonotus, lyy, 181, 182 
eichelbaumi Grouvelle, Laemophloeus, 219 
eichelbaumi (Grouvelle), Mariolaemus, 218, 

219 
elgonesis sp. n., Xylophloeus, igi, 193 
elli sp. n., Notolaemus, 220, 221, 222 
elongatulus Lucas, Laemophloeus, 235 
emgei Reitter, Laemophloeus, 234 
escalerae Grouvelle, Laemophloeus, 219 
cscalerai (Grouvelle), Mariolaemus, 218, 219 
evansi sp. n., Cryptolestes, 232, 240 
exornatus Grouvelle, Laemophloeus, 186, 187 
exornatus (Grouvelle), Placonotus, lyy, 186 

falcidens (Grouvelle), Passandrophloeus, 215 



244 



INDEX 



ferrugineus( Stephens), Cryptolestes, 170,252, 

■233. 235, 236, 238 
ferrugineiis Stephens, Cucujus, 231, 234 
ferrugineus sp. n., Microlaemus, igy, 198 
fractipennis (Motschuisky), Cryptolestes, 168 
Fractophloeus Kessel, 231 

Cannes gen. n., 169, 172, 174, 176 
gestroi Grouvelle, Laemophloeus (s.l.), 171 
glabriculus (Grouvelle, Laemophloeus, 213 
glabriculus (Grouvelle), Passandrophloeus, 

213 
grandiceps Leconte, Narthecius, 210 
granulatus Wollaston, Laemophloeus, 183 

haroldi Reitter, Narthecius, 211 
hypobori (Perris), Leptophloeus, 168, 207 

immoderatus sp. n., Gannes, 174, 175 
insularis Grouvelle, Laemophloeus, 202 
insularis (Grouvelle), Leptophloeus, 202 
interceptus (Grouvelle), Microlaemus, 197 

janeti Grouvelle, Laemophloeus, 209 

janeti (Grouvelle), Leptophloeus, 203, 204, 

205, 209, 210 
juniperi (Grouvelle), Leptophloeus, 168 

Laemophloeus Dejean (s.s.), 168, 171 
Laemophloeus (s.l.), 167, 168, 171, 202 
laevicornis Lefkovitch, Planolestes, 226, 228 
Lathropus Erichson, 168, 169, 171 
lepesmei (Villiers), Cryptolestes, 231, 233, 

235, 236 
lepesmei Villiers, Laemophloeus, 236 
Leptophloeus Casey, 168, 169, lyj, 199, 201, 

213 
Leptus Thomson, 231 
liganus sp. n., Notolaemus, 220, 225 
linearis Grouvelle, Laemophloeus, 207 
linearis (Grouvelle), Leptophloeus, 204, 207 
livens Grouvelle, Laemophloeus, 218 
livens (Grouvelle), Mariolaemus, 218 
longicollis Sharp, Paraphloeus, 211 
longicornis Mannerheim, Laemophloeus, 239 
longicornis sp. n., Mestolaemus, 215, 216 
longicornis Macleay, Placonotus, 176 
lucidus Grouvelle, Laemophloeus , 205 
lucidus (Grouvelle), Leptophloeus, 203, 205, 

207 

Magnoleptus gen n., 169, 170, iy3, 199 
majus, sp. n., Placonotus, J77, 179 
Mariolaemus gen. n., 174, 216, 217 
matris sp. n., Blubos, 194, 196 



Mestolaemus gen. n., iy3, 215 
mestus sp. n., Placonotus, J77, 179 
Microlaemus gen. n., 170, j/j, 196 
micromma Arrow, Cucujinus, 228 
mimosae sp. n., Xylophloeus, igo, 192 
minimus sp. n., Cryptolestes, 232, 240 
minutus Olivier, Cucujus, 238 
mirificus Grouvelle, Laemophloeus (s.l.), 171 
m,irus Grouvelle, Laemophloeus, 183, 184 
misellus Grouvelle, Laemophloeus, 216, 218 
misellus (Grouvelle), Mariolaemus, 218, 219 
mobilis Grouvelle, Laemophloeus, 202, 208 
mobilis (Grouvelle), Leptophloeus, 203, 208 
monilicornis Stephens, Cucujus, 234 
monilis (Fabricius), Laemophloeus, 168 
mossus sp. n., Placonotus, J77, 187 
mucunae sp. n., Leptophloeus, 203, 206 
muticus (Fabricius), Laemophloeus, 168 

Narthecius Leconte, 169, iy3, 210, 212 
nebulosus (Grouvelle), Cucujinus, 230 
nebulosus Grouvelle, Laemophloeus, 230 
nigricollis Lucas, Laemophloeus, 168 
nitens Leconte, Laemophloeus, 183 
nitens (Leconte), Placonotus, 183 
notabilis Grouvelle, Laemophloeus, 168 
Notolaemus Lefkovitch, 168, iy4, 220 

obsoletus Smith, Laemophloeus, 234 
opaculus Grouvelle, Laemophloeus, 205 
opaculus (Grouvelle), Leptophloeus, 202, 203, 
205 

palpalis Waterhouse, Laemophloeus, ig8, 199 
palpalis (Waterhouse), Microlaemus, igy, 198, 

199 
Paracucujinus subgen. n., 230 
parallelicollis sp. n., Magnoleptus, 200, 201 
parallelus Smith, Laemophloeus, 239 
parallelus sp. n., Leptophloeus, 203, 206 
Paraphloeus Sharp, 210, 211 
Passandrophloeus Kessel, 169, iy3, 211, 213 
patens Grouvelle, Laemophloeus, 189, 191 
patens (Grouvelle), Xylophloeus, 187, 189, 

igo, 191 
pauper Sharp, Laemophloeus , 239 
peringueyi Grouvelle, Laemophloeus, 224 
peringueyi (Grouvelle), Notolaemus, 220, 221, 

224, 225 
perrieri Grouvelle, Laemophloeus, 221 
perrieri (Grouvelle), Notolaemus, 220, 221 
perspicuus Grouvelle, Laemophloeus, 222 
perspicuus (Grouvelle), Notolaemus, 220, 221, 

222, 223 
Phloeipsius Casey, 217 



INDEX 



245 



piceus Stephens, Cucujus, 232 
picinus Grouvelle, Laemophloeus, 226 
picinus (Grouvelle), Notolaemus, 220, 226 
picipennis (Grouvelle), Microlaemus, 197 
Placonotus Macleay, 168, 169, iy2, 174, 176, 

224 
Planolestes Lefkovitch, iy3, 222, 226 
planulatus (Grouvelle), Cryptolestes, 231, 236 
planulatus Grouvelle, Laemophloeus, 236 
politissimus Wollastoii, Laemophloeus, 183, 

184 
politissimus (WoUaston), Placonotus, lyy, 

179, 183, 185 
problematicus sp. n., Leptophloeus, 202, 203, 

210 
pugnaceus sp. n., Magnoleptus, 199, 200, 201 
punctatus sp. n., Leptophloeus, 203, 204, 

207, 208, 209 
pusilloides (Steel & Howe), Cryptolestes, 231, 

232, 233, 236, 237, 239, 240 
pusilloides Steel & Howe, Laemophloeus, 236 
pusillus (Schonherr), Cryptolestes, 232, 238, 

239 
pusillus Schonherr, Cucujus, 238 

raffrayi Grouvelle, Laemophloeus (s.l.), J72 

Rhinolaemus Steel, 169 

Rhinomalus Gemminger & Harold, 169 

Rhinophloeus Sharp, 169 

riartus sp. n., Notolaemus, 220, 223 

rufipes Lucas, Laemophloeus, 232 

schedli sp. n., Narthecius, 211, 212 
sepicola (Miiller), Lathropus, 168 
Silvanophloeus Sharp, 176, 183, 186 
slades sp. n., Microlaemus, igy, 198, 199 
spartii (Curtis), Cryptolestes, 231, 232, 233, 
234. 235, 236 



spartii Curtis, Cucujus, 232 
spinosus Grouvelle, Laemophloeus, 213 
spinosus (Grouvelle), Passandrophloeus, 215 
stenoides WoUaston, Laemophloeus, 206 
stenoides (WoUaston), Leptophloeus, 204, 206 
subniger Grouvelle, Laemophloeus, 219 
subtruncatus sp. n., Placonotus, J77, 179, 

181, 182 
suffusus WoUaston, Laemophloeus 232, 233 
sulcifrons (Grouvelle), Microlaemus, 197 
suturalis Grouvelle, Narthecius, 211, 212 
sylvestris (Grouvelle), Microlaemus, 197 

tastus sp. n., Placonotus, lyy, 178, 179 

tenebrosus Grouvelle, Laemophloeus, 198 

testaceus PaykuU, Cucujus, 234 

testaceus Stephens, Cuctijus, 238 

testaceus (Fabricius), Placonotus, 168, 176 

Truncatophloeus Kessel, 201, 202 

turcicus (Grouvelle), Cryptolestes, 232, 237, 

238, 239 
turcicus Grouvelle, Laemophloeus, 237 
tumeri sp. n., Microlaemus, 196, 197, 198 

ugandae Steel & Howe, Cryptolestes, 232, 239 
unicolor Grouvelle, Laemophloeus, 187, 188 
unicolor (Grouvelle), Xylolestes, 188, 189 
unifasciatus Latreille, Cucujus, 220 
unifasciatus (Latreille), Notolaemus, 226 
unifasciatus sp. n., Xylophloeus, 189, igi 
193 

vermiculatus WoUaston, Laemophloeus, 235, 
236 

Xylolestes gen. n., iy2, 187, 190 
Xylophloeus gen. n., iy2, 189 




ENTOM. 12, 4. 



17 



PRINTED IN GREAT D RITAIN BY 
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BARTHOLOMEW PRESS, DORKING 



i 



A. 

S. 



MICROLEPIDOPTERA FROM THE 
NEW HEBRIDES 

RECORDS AND DESCRIPTIONS OF 

MICROLEPIDOPTERA COLLECTED ON THE 

ISLAND OF ANEITYUM BY 

MISS EVELYN CHEESMAN, O.B.E., 




J. D. BRADLEY 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 12 No. 5 

LONDON: 1962 



I 



MICROLEPIDOPTERA 

FROM 

THE NEW HEBRIDES 

RECORDS AND DESCRIPTIONS OF MICROLEPIDOPTERA 
COLLECTED ON THE ISLAND OF ANEITYUM BY 
MISS EVELYN CHEESMAN, O.B.E. 



BY 



J. D. BMPLEY _Y I 

British Museum (Natural History) \ 



Pp. 247-271 ; Plates 14-27 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 12 No. 5 

LONDON: 1962 



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 
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This paper is Vol. 12, No. 5 of the Entomological 
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PRINTED BY ORDER OF THE TRUSTEES OF 
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MICROLEPIDOPTERA FROM THE NEW 

HEBRIDES 

RECORDS AND DESCRIPTIONS OF MICROLEPIDOPTERA 

COLLECTED ON THE ISLAND OF ANEITYUM BY 

MISS EVELYN CHEESMAN, O.B.E. 

By J. D. BRADLEY 

SYNOPSIS 
A little over 700 specimens of Microlepidoptera collected by Miss Evelyn Cheesman on the 
island of Aneityum in 1955 are studied. Previous records from Aneityum are unknown, and the 
material provides new data on the geographical distribution of many species not previously 
recorded from the New Hebrides. Of the sixty-nine species represented, eighteen species and one 
subspecies are described as new to science. No new genera are described. 

INTRODUCTION 

Aneityum is the southernmost island of the New Hebrides Archipelago, and its 
position in the SW. Pacific, shown in the Map on p. 250, is important biogeographic- 
ally (Cheesman, 1957, Nature, 180 : 903-904). The Microlepidoptera collected by 
Miss Cheesman during her expedition to the island in 1955 have therefore proved 
exceptionally interesting, particularly as no previous records of this group are known 
from Aneityum. 

The collection has greatly increased our knowledge of the distribution of many of 
the species and genera recorded below. A little over 700 specimens representing 
sixty-nine species have been examined and determined to species or genus. 

All the material dealt with in the present paper was collected by Miss Cheesman 
during the period from March to June, 1955, both months inclusive, in the same 
locality, at Red Crest, 1,200 ft., 3 miles north-east of Anelgauhat. This locahty data 
has been slightly abbreviated in the records of the species now listed. A systematic 
arrangement is followed ; and the geographical distribution, so far as known, is given 
for each previously described species. 

The type specimens of the new species described below are in the British Museum 
(Natural History). The insects are described as seen under a low-power lens of up 
to X 15 magnification, and the colour terms used are principally from Ridgway's 
Color Standards and Color Nomenclature. 

The illustrations of the wings, PI. 14, are reproduced from photographs taken by 
Mr. N. Tanti of the photographic section of this museum. The magnification varies 
in the photographs and has not been indicated ; instead the measurements across 
the fully-spread fore wings of the actual specimens studied are given under the figure 

PNTOM. 12, 5. ' j8 



250 



J. D. BRADLEY 




t 



MICROLEPIDOPTERA FROM THE NEW HEBRIDES 251 

captions. The genitalia illustrations on Pis. 15-27 are from drawings by Mr. Arthur 
Smith. 

TORTRICIDAE 

TORTRICINAE 

Tortrix sp. {sensu Meyrick) 

Aneityum : Red Crest, 1,200 ft., iii.1955, 2 $. 

These two specimens are superficially reminiscent of the New Zealand species 
Tortrix excessana (Walker), and belong to a species in the same Australian and New 
Zealand species-group at present accommodated in the composite genus Tortrix 
Linnaeus. There are a number of Austrahan species of this group described by 
Turner which are not known to me, and the New Hebridean specimens perhaps 
belong to one of these. 

Nesoscopa psarodes sp. n. 

(PI. 14, lig. I) 

(^$, 11-13 mm. Labial palpus white, suffused and irrorate with mouse grey exteriorly- Head, 
thorax and tegula whitish diffusely irrorate with mouse grey except front of head (face). Antenna 
greyish fuscous, upper side thinly clothed with white scales near base ; scape very short, whitish 
mixed with mouse grey. Fore wing whitish, usually mixed with grey, the grey coloration tending 
to form transverse striae and strigulae ; a narrow slightly oblique diffuse mouse grey basal fascia 
mixed with blackish on costa and containing a jet black dash below costa and a second similar 
dash near middle, both dashes encircled with ochraceous-orange scales ; a narrow curved mouse 
grey fascia at i /4, variable and often incomplete, usually heavily overlaid with blackish on costa 
forming a quadrate well-defined blackish blotch ; in the type specimen the edges of the fascia 
are moderately well defined and sprinkled with black scales, with in some examples a few 
ochraceous-orange scales at middle of outer margin ; a similar straight slightly outwardly- 
oblique fascia at middle, usually with an admixture of blackish towards inner margin (dorsum) 
and on costa, proximal margin of fascia edged with ochraceous-orange at middle ; distal part of 
wing traversed by several irregular curved oblique blackish striae ; cilia whitish, a greyish 
sub-basal line. Hind wing and cilia smoke grey. 

Male genitalia : PI. 15, figs. 2 and 3. 

Female genitalia : PI. 15, fig. i. 

Holotype ^, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of Anel- 
gauhat, vi.1955 (L. E. Cheesman). Genitalia slide 4911. 

Allotype $, same data. Genitalia slide 5526. 

Paratypes 15 c^, 5 9. same locality data, dated from iii to vi.1955. Genitaha 
shde 5515 (male). 

This species superficially resembles N. exors Meyrick, the only other known species 
in the genus and described from Rapa I. The males of these two species can be 
readily separated by differences in the uncus which can be seen without dissection 
by brushing away a few of the covering scales and examining with a strong lens 
( X 20) . In exors the uncus is broad and obtuse and has two prominent sublateral 
barbs beneath the tip ; in psarodes the uncus is slender and finger-like and is not 
barbed at the tip. 



252 J- D. BRADLEY 

OLETHREUTINAE 
Spilonota cryptogramma Meyrick 
Spilonota cryptogramma Meyrick, 1922, Exot. Microlep. 2 : 520. 

Aneityum : Red Crest, 1,200 ft., iii-vi.1955, 23 (J$. 

Distribution. Fiji. 

Compared with examples from Fiji, the New Hebridean specimens tend to be 
shghtly smaller and have the blackish dashes in the discal and apical areas of the 
fore wing much heavier and darker, forming in some examples a continuous black 
stripe from middle to near apex. They possibly represent a distinct race, and 
when additional material from other localities becomes available their specific 
status will need to be reconsidered together with the species S. infensa Meyrick, 
described from Queensland, of which cryptogramma may prove to be no more than 
a subspecies. 

Herpystis jejuna Meyrick 

Herpystis jejuna Meyrick, 1916, Exot. Microlep. 3 : 16. 

Aneityum : Red Crest, 1,200 ft., iv and v. 1955, 3 (^, i $. 
Distribution. India, Java and Solomon Is. (Rennell I.). 

Eucosma euryochra sp. n. 

(PL 14, fig. 2) 

<J$, 15-19 mm. Labial palpus cartridge buff suffused with cream, basal and second segments 
overlaid with ochreous-tawny or cinnamon-brown exteriorly. Head, thorax and tegula cartridge 
buff, side of crown adjacent to eye, patagia and base of tegula cinnamon-brown. Antenna and 
scape cartridge buff, fiagellum diffusely annulate with fuscous except base. Fore wing cartridge 
buff or whitish, some scattered grey and cinnamon-brown scales, heaviest in female ; a narrow 
cinnamon-brown transverse fascia at 1/3, diffuse and Aveakly defined from costa to middle, 
thence becoming gradually more solid and darkened by an admixture of black scales, almost 
entirely black at inner margin (dorsum) ; basal area of wing between costa and plical fold and 
including upper (costad) half of fascia suffused with light violet-grey, in some examples this 
suffusion extends along the costal area beyond middle of wing but is usually less pronounced ; 
a subtriangular cinnamon-brown mixed with black blotch on inner margin beyond middle, base 
of blotch greater than width of fascia ; discal area without definable ocellus, sometimes shaded 
with pale violet-grey ; costa marked with short cinnamon-brown and blackish strigulae, beyond 
middle of wing these are sometimes linked with rippled strigae of cinnamon-brown scales 
traversing the wing, one somewhat heavier chain intermixed with black scales from termen 
below middle curving inwards towards costa ; a small round cinnamon-brown patch at apex 
centred with black and emitting a strigulation along termen ; cilia cinnamon-brown mixed with 
dark plumbeous. Hind wing with veins 3 and 4 stalked ; cinnamon-brown densely irrorate with 
fuscous scales ; cilia greyish, a slightly darker sub-basal line. Posterior tibiae in both sexes 
rough-scaled above to about middle. 

Male genitalia : PI. 15, fig. 4. 

Female genitalia : PI. 16, figs. 1-3. 

Holotype c?. New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m, NE. of Anel- 
gauhat, iii.1955 {L. E. Cheesman). Genitaha slide 4846. 



1 



MICROLEPIDOPTERA FROM THE NEW HEBRIDES 253 

Allotype $, same locality data, dated iv.1955. Genitalia slide 5525. 

Paratypes 10 cJ, 7 $, same locality data, dated from iii to iv.1955. 

Provisionally placed near a species described from India, E. legitima Meyrick, 
which it superficially resembles in coloration and pattern, but it can be readily 
distinguished by the absence of the large blackish subapical terminal blotch present 
in the fore wing of legitima. 

Bactra blepharopis Meyrick 

Bactra blepharopis Meyrick, 191 1, Proc. Linn. Soc. N.S.W. 36 : 255. 

Aneityum : Red Crest, 1,200 ft., iii and vi.1955, 3 $. 

Distribution. Australia (Queensland), Solomon Is., Fiji and New Hebrides. 

Polychrosis orthomorpha Me3Tick 

Polychrosis orthomorpha Meyrick, 1928, Exot. Microlep. 3 : 443. 

Aneityum : Red Crest, 1,200 ft., iv.1955, i $. 
Distribution. New Hebrides. 

Crusimetra anastrepta Meyrick 

Crusimetra anastrepta Meyrick, 1927, Insects of Samoa, 3(2) : 71. 

Aneityum : Red Crest, 1,200 ft., iv.1955, i ^. 
Distribution. Ceylon, Sumatra, Samoa and Solomon Is. 

Strepsicrates glaucothoe (Meyrick) comb. n. 

Spiloticta glaucothoe Meyrick, 1927, Insects of Samoa, 3(2) : 70. 
Eucosma baryphragma Meyrick, 1937, Exot. Microlep. 5 : 159. Syn. n. 

Aneityum : Red Crest, 1,200 ft., iii and vi.1955, 3 (^, i $. 

The new synonymy above has been confirmed after examination of the types in 
the British Museum (Natural History). 
Distribution. Samoa and Fiji. 

Strepsicrates poliophora sp. n. 

(PL 14, fig. 3) 

(^,18 mm. Labial palpus white interiorly, pale gull grey exteriorly, second segment with 
well-defined dark mouse grey sub-basal and postmedial bands exteriorly and with apex slightly 
projected and suffused deep mouse grey, terminal segment indistinctly ringed with deep mouse 
grey at apex and suffused with similar coloration exteriorly. Head, thorax and tegula fuscous- 
black mixed and irrorate with pale gull grey, front of head white, a light grey patch on posterior 
of thorax. Antenna with notch at about 1/7, mouse grey, scape irrorate with light grey dorsally. 
Fore wing with broad costal fold, enclosing whitish hairs, from base to middle ; pale gull grey, 
costal and distal areas heavily and dorsal (inner margin) area lightly suffused and irrorate with 
an irregular mixture of deep mouse grey and fuscous ; an extensive almost blackish area between 
costa and plical fold reaching beyond middle to discal area ; a prominent tuft of raised light 



254 J- D. BRADLEY 

grey scales barred with deep mouse grey at middle of plical fold ; costa marked from before 
middle to apex with short blackish strigulae ; a large circular blackish pretornal patch, nearly 
coalescent with blackish central area, a similar larger somewhat elongate patch in terminal 
margin, an admixture of sayal brown scales between this and the blackish central suffusion, a 
light sprinkling of sayal brown scales in area of wing having darker coloration ; cilia pale gull 
grey irrorate with deep mouse grey, a broad deep mouse grey basal line along termen. Hind 
wing dark fuscous ; cilia greyish, a mouse grey sub-basal line. 
Male genitalia : PL 17, fig. i. 

Holotype cJ, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, iii.1955 {L. E. Cheesman). Unique. Genitalia slide 4883. 

Very closely related to S. glaucothoe (Meyrick), S. rhothia (Meyrick), and 5. dilacerata 
(Meyrick) ; all four species no doubt originating from the same stock. S. poliophora 
may be readily distinguished from the other three species mentioned by the male 
genitaUa which have a well-developed digitate projection on the ventral margin of 
the cucullus. 

Mesocallyntera dascia sp. n. 

(PI. 14. fig. 4) 

cJ, 24-27 mm. Labial palpus tawny, upper edge of second segment rough-scaled towards apex 
and clothed with plumbeous-black scales tipped with light grey ; terminal segment very short, 
apex suffused plumbeous-black. Head fuscous, with an admixture of plumbeous-black strongest 
on frons and fore part of vertex between antennae, scales at middle and posterior of crown 
tipped with whitish buff. Thorax fuscous overlaid with loosely appressed leaden-metallic plumb- 
eous scales, posterior crest mixed with tawny. Tegula tawny, shaded with dark fuscous at tip, 
overlaid with loosely appressed leaden-metallic plumbeous scales at base matching thorax. 
Antenna and scape fuscous. Fore wing brownish tawny, densely suffused with leaden-metallic 
plumbeous scales forming a nebulous irregular pattern over most of the wing ; markings blackish 
brown, distal margin of basal patch well defined, irregularly sinuous and slightly outwardly- 
oblique from costa at about 1/4 ; a heavy but poorly-defined triangular marking from costa 
beyond middle, apex of this marking confluent in discal area with a tawny dash ; a moderately 
well-defined inwardly-oblique slightly undulate postmedial line, parallel with termen, from 
dorsum near tornal angle to near costa where it merges with a somewhat diffuse subtriangular 
blackish brown patch a little below costal margin, this patch extending obliquely to costa and 
edged outwardly with a fine leaden-metallic plumbeous line from costa to termen ; a small 
wedge-like tawny tuft on inner margin a little proximad of postmedial line ; a similar elongate 
tuft on inner margin before middle ; small interneural tawny dots along termen ; a small tawny 
mixed with blackish subterminal dashes on veins 3 and 4, and similar blackish dashes on veins 5, 
6 and 7, all forming a chain ; costa marked with short pale tawny and blackish strigulae ; 
cilia plumbeous, a diffuse plumbeous-black basal line not evident at tornus where cilia are paler. 
Hing wing and cilia light fuscous ; a slender warm buff hair pencil from mesopleura sometimes 
concealed in vannal fold. Abdomen fuscous with an admixture of grey, a dense patch of 
specialized warm buff lanceolate scales on sternum of second segment. Legs fuscous, tarsi 
banded or suffused warm buff, tufts on fore and middle tibiae purplish black, inner side of 
hind tibia cartridge buff and with a long dense pencil of warm buff hairs from base. 

Male genitalia : PI. 18, fig. i. 

Holotype c^, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, vi.1955 {L. E. Cheesman). Genitalia shde 6020. 
Paratypes 7 3", same data. Genitalia slide 4866. 



MICROLEPIDOPTERA FROM THE NEW HEBRIDES 255 

Allied and similar in appearance to M. squamosa Diakonoff, found in the Celebes 
and, as a subspecies, in New Guinea. Differences in the valva of the male genitalia 
distinguish the two species, that of dasciaheing simple and not furcate as in sqimmosa. 

Olethreutes ancosema ancosema (Meyrick) 

Argyroploce ancosema Meyrick, 1932, Exot. Microlep. 4 : 310. 

Aneityum : Red Crest, 1,200 ft., iv.ig55, 2 S- 

Both specimens resemble the nominate form from Fiji, having the black medial 
marking on the fore wing unbroken, and are at least for the present referred to this 
subspecies. In the Solomons race, 0. ancosema solomonensis Bradley, the medial 
fascia is interrupted at the middle. 

Distribution. Fiji. 

Olethreutes anaprobola (Bradley) 

Argyroploce anaprobola Bradley, 1953, Proc. Hawaii, ent. Soc. 15 : 109. 

Aneityum : Red Crest, 1,200 ft., iii-vi.1955, 11 (^, 1 $, 
Distribution. Fiji and Solomon Is. 

Cryptophlebia ombrodelta (Lower) 

Aroirophora (?) ombrodelta Lower, 1898, Proc. Linn. Soc. N.S.W. 23 : 48. 
Cryptophlebia carpophaga Walsingham, 1899, Indian Mus. Notes, 4 : 106. 

Aneityum : Red Crest 1,200 ft., iv and v. 1955, 2 c^, i ?. 

Distribution. Widespread in the Indo-Australian region, the larva being 
polyphagous on the leaves, pods, seeds and fruits of various tropical trees and 
shrubs. 

Cryptophlebia iridoschema sp. n. 

(PI. 14. fig. 5) 

$,15 mm. Labial palpus buff-yellow, suffused exteriorly with amber brown. Head buff-yellow, 
some amber brown scales at side of face adjacent to compound eye, an orbit of rather slender 
scales on posterior of crown behind ocellus and adjacent to compound eye. Thorax buff-yellow 
suffused with amber brown, most strongly anteriorly. Tegula amber brown, paler at tip. 
(Antennae missing.) Fore wing with basal area to middle of costa and 1/3 inner margin (dorsum) 
amber brown, outer edge of this area sharply defined, almost straight, inwardly-oblique from 
costa to inner margin, edged with whitish bej^ond giving way to mars yellow in distal half of 
wing ; a small triangular amber brown pretornal marking on inner margin ; costa marked with 
very short oblique amber brown strigulae in apical half, those nearest the apex heaviest, terminat- 
ing as thin lines of amber brown scales ; a sprinkling of amber brown scales, sometimes arranged 
in short transverse chains, in distal area ; interspersed between these are very small inconspicuous 
patches of plumbago-grey scales ; cilia pinkish buff, a rather indistinct amber brown sub-basal 
line, shaded with sayal brown beyond. Hind wing mars yellow except basal 3/4 of costal area 
which is whitish ; cilia pinkish buff, amber brown sub-basal line, shaded with sayal brown 
beyond. 

ENTOM. 12, 5. 19 



256 J. D. BRADLEY 

Female genitalia : PI. 16, lig. 4. Bursa copulatrix spherical, surface ornamented with minute 
rhomboidal scutullates, on one side a pair of weakly sclerotized granulose signa are present. 

Holotype $, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, v. 1955 (L. E. Cheesman). Unique. Genitalia slide 6051. 

A species of distinctive appearance, having brighter coloration than is usual for 
the genus, and also an unusual wing pattern. In general coloration nearest the 
two closely related species C. ombodelta (Lower) and C. vitiensis Bradley, but readily 
distinguished from these and other species of the genus by the transverse division 
of the fore wing into two distinct colour areas. 

Cryptophlebia rhynchias (Meyrick) 

Platypeplus rhynchias Meyrick, 1905, /. Bombay nat. Hist. Soc. 16 : 586. 

Aneityum : Red Crest, 1,200 ft., vi.1955, i $. 

Distribution. S. India, Ceylon, Austraha (Queensland), New Hebrides and 
Sudest I. 

SCHOENOTENINAE 
Schoenotenes elaphrodes sp. n. 

(PI. 14, fig. 6) 

(J, 15 mm. Labial palpus white, terminal segment wholly or partially suffused with greyish, 
second segment suffused with greyish exteriorly and with an admixture of blackish grey and 
ochreous in upper and lower margins. Head, thorax, patagia and tegula whitish grey, tinged and 
partially slightly suffused with ochraceous-buff. Fore wing whitish, suffused smoke grey except 
distal area ; scales roughened and raised more especially in discal area ; some vertical diffuse 
ochraceous-orange strigulation superimposed on the greyish suffusion ; a broad black line along 
plical fold from near base to middle of wing, terminating a little above inner margin (dorsum) ; 
a blackish dot at middle joining distally a broad blackish suffusion extending obliquely to costa a 
little before apex ; a moderately heavy diffuse ferruginous mixed with blackish dorsal blotch at 
or just beyond middle ; a few faint irregular transverse ochreous mixed with blackish strigulations 
in whitish terminal area ; several scattered rather inconspicuous black specks along edge of 
inner margin ; cilia whitish, basal half shaded ochraceous-buff, a faint blackish medial line, 
some faint blackish dots along termen at ends of veins. Hind wing greyish fuscous, darker 
distally ; cilia concolorous with wing basally, white apically. 

Male genitalia : PI. 18, figs. 2 and 3. 

Holotype (^, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, v. 1955 (L. E. Cheeseman). Genitalia shde 5516. 

Paratype i ^, same locality data, dated iii.1955. Genitalia slide 6034. 

This species is the only representative of the genus Schoenotenes Meyrick so far 
recorded from the New Hebrides. The centre of development of this genus is New 
Guinea, where the species are numerous, and it is but poorly represented elsewhere 
in the Indo-Australian region. The genus contains species arrangeable in a natural 
sequence according to fascia and genitalia development, and elaphrodes belongs in 
the group with subrectangular or subtriangular fore wings in which rough-scaling 
is reduced and may be present onty in plical and discal scale tufts. The male 
genitalia are without hami, with pendulous .socii, hooked gnathus and simple valva. 



I 



MICROLEPIDOPTERA FROM THE NEW HEBRIDES 257 

CHLIDANOTINAE 

Caenognosis incisa Walsingham 

Caenognosis incisa Walsingham, 1900, A Monograph of Christmas Island, p. 79. 

Aneityum : Red Crest, 1,200 ft., iii and iv.1955, 12 ex. 

Distribution. Australia (Queensland), New Guinea, Philippines and Christmas I. 

Trymalitis cataracta Meyrick 

Trymalitis cataracta Meyrick, 1907, /. Bombay nat. Hist. Soc. 18 : 153. 
Trymalitis optima Meyrick, 191 1, Proc. Linn. Soc. N.S.W. 36 : 294. 
Trymalitis macaiista Meyrick, 1934, Exot. Microlep. 4 : 489. 

Aneityum : Red Crest, 1,200 ft., iii, iv and v,i955, 2 (^, i ?. 
Distribution. E. Austraha, New Guinea, Solomon Is., Bismarck Archipelago, 
Fiji, Siam, Andaman Is., Ceylon and Africa. 

GELECHIIDAE 

Stegasta variana Meyrick 

Stegasta variana Meyrick, 1904, Proc. Linn. Soc. N.S.W. 29 : 394. 

Aneityum : Red Crest, 1,200 ft., iv.1955, i (^. 

Distribution. Widely distributed in the Ethiopian and Indo- Australian regions. 

Thiotricha oxyopis MejTick 

Thiotricha oxyopis Meyrick, 1927, Insects of Samoa, 3(2) : 80. 

Aneityum : Red Crest, 1,200 ft., iii. 1955, i !^. 
Distribution. Samoa and Solomon Is. 

Thiotricha sp. 

Aneityum : Red Crest, 1,200 ft., iii. 1955, i (^. 

The wings of the specimen are rubbed and the species cannot be properly 
identified, but the genitalia indicate affinity with T. oxyopis Meyrick. 

Thiotricha sp. 

Aneityum : Red Crest, 1,200 ft., iv.1955, i (^, damaged. 

Superficially similar to T. tethela Bradley which occurs in the Solomon Is. 

Idiophantis lomatographa sp. n. 

(PI. 14, fig. 7) 

$,14 mm. Labial palpus whitish interiorly, drab exteriorly, terminal segment with a deep 
violaceous iridescence exteriorly. Head with face warm buff, crown ochraceous-buff, suffused 
with glossy dark violet-grey mediallv. Antenna and scape whitish, suffused with dark vjolet- 



258 J. D. BRADLEY 

grey anteriorly. Thorax and tegula purplish hair-brown. Fore wing with termen deeply excav- 
ated beneath linear upturned apical prominence ; drab, costal edge paler, markings confined to 
distal area ; an obtusely bent greyish white (plumbeous) blackish-edged subterminal streak, 
thickened and pure white at costa, preceded by an orange streak thinly edged blackish ; area 
distad of subterminal streak orange on upper half, pale apricot yellow on lower half ; a thick 
plumbeous streak along costa towards apex ; a large mixed drab and black circular marking at 
middle of termen, a similar dash above in deep excavation beneath apical prominence, a third 
dash below a little before mid-way to tornus reaching inward from termen to middle of yellow 
blotch ; cilia concolorous, tufted and white tipped with dark grey at extremity of apical projec- 
tion, gold and iridescent reflections opposite mid-termen marking. Hind wing greyish, whitish 
around apex ; cilia concolorous, a black basal line at apex. Abdomen purplish hair-brown above, 
cartridge buff below. Legs cartridge buff, suffused purplish hair-brown exteriorly. 

Female genitalia : PI. 19, figs, i and 2. Ostium very small, circular, situated centrally in a 
slight depression proximad to a deep U-shaped excavation in caudal margin of ostial plate. 
Ductus bursae filamentous to middle thence broadening considerably to bursa copulatrix, which 
is spherical and has a very long convoluted filamentous tube (possibly ductus seminalis) arising 
from proximal half. 

Holotype $, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, iv.1955 {L. E. Cheesman). Unique. Genitalia slide 6580. 

Related to /. discura Meyrick, an Asiatic species, and to /. callicarpa Meyrick, 
a Samoan species ; and distinguished by the blackish dash on the termen of the 
fore wing toward the tornus, not present in other species of the genus. 

Idiophantis soreuta Meyrick 

Idiophantis soreuta Meyrick, 1906, /. Bombay nat. Hist. Soc. 17 : 139. 

Aneityum : Red Crest, 1,200 ft., iv.1955, i $. 
Distribution. Ceylon and St. Matthias I. 

Autosticha silacea sp. n. 

(PI. 14, fig. 8) 

(J$, 12-14 mm. Labial palpus warm buff, diffusedly irrorate with fuscous exteriorly. Head, 
thorax, tegula, antenna and scape warm buff ; thorax and tegula diffusedly irrorate with dark 
brown and fuscous scales with a weak sheen ; antenna weakly marked with fuscous segmental 
bars anteriorly, diffuse and less conspicuous in male. Fore wing warm buff, whole wing diffusedly 
irrorate with dark brown and fuscous scales with a weak purplish sheen ; stigmata weak and 
obscure, plical slightly basad of discal, second discal hardly discernible at end of cell in middle; 
cilia warm buff, suffused greyish ; a diffuse dark grey sub-basal line. Hind wing uniformly 
whitish cream-buff ; cilia concolorous. Legs warm buff, weakly irrorate with fuscous exteriorly. 

Male genitalia : PI. 17, figs. 2 and 3. Tegumen margin broadly produced ventrally. Valva 
with weakly sclerotized glabrous area at middle of costa. Uncus moderately stout, obtuse. 
Gnathus produced medially to a long, slender, digitate point. 

Female genitalia : PI. 20, figs, i and 2. 

Holotype ^, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, iii.1955 [L. E. Cheesman). Genitalia slide 6038. 

Allotype $, same locality data, dated iv.1955. GenitaHa slide 6577. 
Paratype i ^, same data as holotype. Genitalia slide 6527. 



I 



MICROLEPIDOPTERA FROM THE NEW HEBRIDES 259 

Superficially similar to A. brunnea Bradley, from the Solomon Is., but distin- 
guished by the lighter coloration. The structure of the male genitalia of silacea 
indicates a close relationship to A. solita (Meyrick), a Fijian species, and ^. banausopa 
(Meyrick), comb, n., from the New Hebrides, and also to certain other species at 
present in the composite genus Autosticha Meyrick. The present generic placing is 
tentative. A. banausopa was originally described in the genus Pachnistis Meyrick 
and has now been transferred because of its close relationship, evident from the 
morphology of the genitalia, with A. solita. The latter species was originally 
described in Pachnistis but was later transferred by Mej^rick (Mej^rick in Wytsman, 
1925, Genera Insectorum, fasc. 184 : 256). 

A. silacea is larger than banausopa and solita, both these species having a wing 
expanse of lo-ii mm., and is further distinguished by its lighter ochreous coloration, 
which in those two species approaches dark brown. The venation of the fore wing 
also differs ; in silacea vein 7 is present, and veins 8 and 9 arise from vein 7 ; in 
banausopa and solita vein 7 of the fore wing is absent (possibly coincident with vein 8), 
and veins 8 and 9 are .stalked or connate. The generic value of the stalking of 
vein 9 with vein 7 in the fore wing in the Autosticha group of genera seems doubtful 
as the venation appears to vary between closely related species, but it may be reliable 
specifically. The marginal lobe developed ventrally on the tegumen in the male 
genitaha may be a character of generic importance, and is present in silacea, banausopa 
and solita. 



SCAEOSOPHIDAE 
Bathraula sitnulatella (Walker) 

Cryptolechia simulatella Walker, 1864, Cat. Lep. Ins. B.M. 29 : 747. 
Cryptolechia niveosella Walker, 1864, Ibid. 29 : 747. 

Aneityum : Red Crest, 1,200 ft., v. 1955, i (^. 

Distribution. Sarawak, Borneo, Phihppines, Bismarck Archipelago and 
Society I. 

COSMOPTERYGIDAE 

Labdia clodiana Meyrick 
Labdia clodiana Meyrick, 1927, Exoi. Microlep. 3 : 384. 

Aneityum : Red Crest, 1,200 ft., v. 1955, i (^. 
Distribution. New Hebrides. 

Labdia saliens Meyrick 
Labdia saliens Meyrick, Exot. Microlep. 3 : 385. 

Aneityum : Red Crest, 1,200 ft., iii, iv and v. 1955, 14 ^^. 
Distribution. New Hebrides and Solomon Is. 



26o J.D.BR A D L E Y 

Labdia calida Meyrick 

Labdia calida Meyrick, 192 1, Exot. Microlep. 2 : 411. 

Aneityum : Red Crest, 1,200 ft., iv.1955, i $. 
Distribution. Fiji. 

Labdia cedrinopa Meyrick 

Labdia cedrinopa Meyrick, 1928, Exot. Microlep. 3 : 386. 

Aneityum : Red Crest, 1,200 ft., iii.1955, 2 $. 
Distribution. New Hebrides (Efate I.). 

Proterocostna triplanetis Meyrick 

Proterocosma triplanetis Meyrick, 1886, Trans, ent. Soc. Lond. 1886 : 293. 

Aneityum : Red Crest, 1,200 ft., iv and vi.1955, 2 $. 
Distribution. Tonga, Fiji, Samoa, Solomon Is. and New Hebrides. 

Litnnaecia microglypta Meyrick 

Limnoecia [sicl microglypta Meyrick, 1928, Exot. Microlep. 3 : 393. 

Aneityum : Red Crest, 1,200 ft., vi.1955, i <^. 
Distribution. New Hebrides (Efate I.). 

Litnnaecia arsitricha Meyrick 

Limnoecia [sic'] arsitricha Meyrick, 1927, Insects of Samoa, 3(2) : 93. 

Aneityum : Red Crest, 1,200 ft., iv. and v. 1955, i (J, i $. 
Distribution. Samoa, Solomon Is. and New Hebrides. 

Ascalenia sp. 

Aneityum : Red Crest, 1,200 ft., iv and v. 1955, 5 $. 

Representing a species superficially approaching A. armigera Meyrick but with 
general coloration considerably darker. 

A. armigera is known only from two male specimens ; one the type from Fiji, 
and the other a specimen from the Solomon Is. As the specimens from the New 
Hebrides are females it has not been possible to make a genitalic comparison with 
armigera. It would be particularly interesting if this could be done, since the male 
genitaha of armigera have specialized setae on the valva which may be homologous 
with those found in the highly specialized genus Hyposmochoma Butler, endemic in 
the Hawaiian Islands, 



MICROLEPIDOPTERA FROM THE NEW HEBRIDES 261 

CARPOSINIDAE 

Peragrarchis rodea Diakonoff 

Peragrarchis rodea Diakonoff, 1950, Bull. Brit. Mus. [nat. Hist.), Ent. 1 : 299. 

AneityUxM : Red Crest, 1,200 ft., iii.1955, i (^, 
Distribution. New Guinea. 

Peragrarchis pelograpta (Meyrick), comb. n. 

Meridarchis pelograpta Meyrick, 1929, Trans, ent. Soc. Lond. 76 : 502. 

Aneityum : Red Crest, 1,200 ft., iii.1955, 1(^,1$. 

The New Hebridean specimens are smaller than specimens from the Austral Is., 
the only other known locahty for this species, and shght genitalic differences in the 
male, particularly in the aedeagus, indicate some degree of subspeciation. 

Distribution. Austral Is. 

Peragrarchis minima sp. n. 

(PI- 14. fig- 9) 

c^, 14 mm.; 9. ^5 "ii^i. Labial palpus in male about li width of eye, subascending ; in female 
about 3 times width of eye, second segment being greatly extended, porrect ; cartridge buff, 
second segment fuscous below and exteriorly to near dorsal margin, terminal segment with dark 
fuscous medial ring. Antennal ciliations in male 2-3 times width of shaft basally to near middle, 
gradually shortening towards apex to less than width of shaft, which is clothed dorsally with 
cartridge buff scales. Head, thorax, tegula and antennal scape cartridge buff. Fore wing 
cartridge buff irregularly suffused with tawny and irrorate with fuscous and fuscous-black, some 
scales grouped as raised tufts ; 7 or 8 irregular fuscous-black elongate spots on costa ; and ill- 
defined diffuse triangular patch resting on costal spots from middle to 4/5, extending across 
wing beyond middle to fold, its apex truncate, containing a blackish tuft or tufts in disc ; 
termen outlined with tawny ; a diffuse poorly-delined tawny mixed with blackish subterminal 
fascia ; cilia grey, basal half darker, minutely irrorate with black and whitish specks. Hind 
wing light grey ; cilia matching. 

Male genitalia : PI. 22, figs, i and 2. Valva fused along basal portion of sacculus ; cucullus 
weakly sclerotized, bipartite to near base — a characteristic of the genus — halves linear ; 
sacculus with sclerotized denticulate subventral ridge in basal half, a strong curved tooth at 
base, a heavy denticulate apical patch preceded by a solitary, longer, aculiform tooth curved 
cephalad to nearly meet the subventral ridge. Aedeagus about as long as distance from apex 
of uncus to end of saccus, moderately sclerotized, dilated before middle, apex asymmetrical 
and produced at one side to a long slender point ; two short, longitudinal, external patches of 
coarse close-set spine-like setae at about middle, approximate and to one side, one patch slightly 
larger ; a densely compact subapical patch of hair-like setae internally. 

Female genitalia : PL 21, figs. 1-3. 

Holotype ^, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, iii.1955 (L. E. Cheesman). Genitaha slide 6029. 

Allotype 9, same data. Genitalia slide 6055. 

Related to the two closely allied species Peragrarchis pelograpta (Meyrick), 
mentioned above, and Peragrarchis syncolleta (Meyrick), comb, n., which occurs in 
the Solomon Is. and the Andamans. Superficially most similar to pelograpta but 
smaller in size. 



262 J. D. BRADLEY 

Meridarchis celidophora sp. n. 

(PI. 14, fig. 10) 

$, 18-22 mm. Labial palpus porrect, about 3 times as long as width of head ; white, basal 
segment suffused fuscous exteriorly, second segment strongly irrorate with fuscous below and 
exteriorly except at apex and upper margin of apical 2/3, terminal segment with diffuse fuscous 
sub-basal ring. Head, thorax and tegula white, head weakly irrorate with dark fuscous on crown 
and laterally, anterior margin of thorax and basal half of tegula strongly suffused fuscous, pos- 
terior half of thorax thinly irrorate with fuscous-black. Antenna sordid white, weakly annulate 
with fuscous towards base ; scape white. Fore wing white, dusted with fuscous, markings fuscous 
and fuscous-black ; an inwardly-oblique basal patch, darker on costa, outer margin straight 
and edged with raised scales ; an elongate solid fuscous-black costal marking before middle, 
succeeded by three smaller spots equidistant apart to 3/4 ; a similar smaller spot at 4/5 ; an 
ill-defined diffuse triangular fuscous mixed with fuscous-black fascia resting on costal spots from 
before middle to 3/4, extending across wing to inner margin (dorsum), strongest at middle and 
distally, most diffuse inwardly towards inner margin, accompanied by tufts of raised scales in 
discal area ; a series of diffuse confluent spots along termen ; a weak greyish fuscous sub-basal 
shade line parallel with outer edge of medial fascia (or patch) nearer to termen than to fascia ; 
cilia whitish grey, white at tornus, a greyish fuscous medial line along termen. Hind wing 
whitish grey ; cilia matching, whitish basally and apically. Abdomen cartridge buff or whitish, 
suffused fuscous laterally. Legs whitish, marked with fuscous-black exteriorly. 

Female genitalia : PI. 19, fig. 3. Ostium concealed, limen weakly sclerotized, curved ventrally, 
dorsal wall produced as a thin, nearly quadrate plate with blunt digitate medial process. Ductus 
bursae dilated and weakly sclerotized for short distance beyond ostium, weakly scobinate. 
Bursa copulatrix membranous, pyriform, without signum. 

Holotype $, New Hebrides : Aneityum, Red Crest, 1,200 ft,, 3 m, NE. of 
Anelgauhat, v. 1955 {L. E. Cheesman). Genitalia slide 6054. 

Paratypes 7 $, same locality data, dated iii, v and vi.1955. 

Near the Japanese M. excisa Walsingham, and closely resembling it superficially 
except that the medial fascia or patch on the fore wing is more extensive and 
pronounced, but differing in coloration which in excisa is more cinereous or grey. 

Carposina stationaria Meyrick 

Carposina stationaria Meyrick, 1928, Exot. Microlep. 3 : 402. 

Aneityum : Red Crest, 1,200 ft., iv., v and vi.1955, 15 ^$. 
Distribution. New Hebrides. 

Heterogymna cheesmanae sp. n. 

(PI. 14, fig. II) 

6> 30-32 mm.; $, 36-40 mm. Labial palpus in male about li times width of eye, slightly 
recurved ; in female about 2J times width of eye, porrect ; cartridge buff, basal and second 
segments overlaid below and exteriorly, except apex of second segment, with mars brown, 
terminal segment with a broad moderately distinct medial annulus interrupted in male interiorly. 
Head cartridge buff, upper part of front and crown mixed with civette green laterally, an admixture 
of mars brown medially. Antenna strongly ciliate in male, filiform in female ; cartridge buff 
suffused with mars brown, darker basally ; scape civette green. Thorax civette green, a broad 
transverse mars brown patch anteriorly, a similar small patch laterally slightly posterior of 



MICROLEPIDOPTERA FROM THE NEW HEBRIDES 263 

middle. Tegula civette green, usually with strong, mars brown irrorate with mummy brown 
markings at base and on apical half ; in some examples a variable admixture of cartridge buff 
towards apex. Fore wing cartridge buff solidly overlaid with light elm green or bice green (in 
worn specimens the green coloration gives way to cartridge buff) lightly irrorate with mars brown ; 
markings mars brown varying to very dark mummy brown ; a small inwardly-oblique basal 
patch with an admixture of green near costa ; male with costal fold reaching to beyond middle, 
heavily marked with mars brown in basal half and beyond middle ; female with an irregular 
mars brown patch on costa at 1/4, a narrow mars brown marking from before middle to 3/5 of 
costa ; in male this marking is shortened by the costal fold ; in both sexes three small angular 
mars brown costal spots converging and separated by white flecks on costal margin ; a narrow, 
inwardly-oblique irregular transverse band at i /4 interrupted before middle ; a broad blotch- 
like incomplete fascia at 2/3 interrupted and mixed with green, edges irregular and diffuse, 
extending to a little beyond middle of wing and disappearing towards inner margin : a narrow 
irregular subterminal band indented at middle ; termen narrowly edged with small intercon- 
nected irregular wedge-shaped spots not reaching apex ; inner margin (dorsum) strongly irrorate 
or overlaid with mars brown ; cilia brownish, a darker basal line. Hind wing greyish drab, male 
with weak cubital pecten ; basal 2/3 of inner margin broadly whitish, in male including basal 
area of wing ; cilia whitish, shaded with pale drab basally. Abdomen cartridge buff strongly 
overlaid with fuscous ; in male a cartridge buff anal tuft. 

Male genitalia : PI. 22, figs. 3 and 4. Uncus curved, slender, digitate, acutely pointed ; 
cucuUus very narrow, slightly tapered, apex rounded ; sacculus moderately broad, a small 
apical tooth ; basal section of valva with short stout tooth near base of cucuUus ; vinculum 
evenly rounded, produced proximally to an obtuse point. Aedeagus extremely large and stout, 
containing about six large blade-like cornuti. 

Female genitalia : PI. 20, figs. 3 and 4. 

Holotype ^, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, vi.1955 {L. E. Cheesman). Genitalia slide 4890. 

Allotype $, same locality data, dated v. 1955. 

Paratypes 5 (J, i $, same locality data, dated iv and vi.1955. Genitalia slides 
4828 (female) and 4889 (male). 

A species of striking appearance and readily distinguished by the green coloration 
from other species of the genus. The male genitalia are similar to those of H. ochro- 
gramma Meyrick. 

I name this remarkable and singularly distinguished species after the collector, 
Miss Evelyn Cheesman. 

Coscinoptycha improbana Mej^ick 

Coscinoptycha improbana Meyrick, 1881, Proc. Linn. Soc. N.S.W. 6 : 701. 

Aneityum : Red Crest, 1,200 ft., iii-vi.1955, 7 ^, 43 $. 
Distribution. Australia (Queensland and N.S. Wales). 

heliodinidae 

Hieromantis munerata Meyrick 
Hieromantis munerata Meyrick, 1924, Exot. Microlep. 3 : 66. 

Aneityum : Red Crest, 1,200 ft., iii-vi.1955, 10 cJ$. 
Distribution. Fiji, Samoa, New Hebrides. 

KNTOM. 12, 5. 20 



264 J. D. BRADLEY 

Stathmopoda periclina Meyrick 
Stathmopoda periclina Meyrick, 1938, Trans. R. ent. Soc. Lond. 89 : 520. 

Aneityum : Red Crest, 1,200 ft., iii and vi.1955, z ^, 1^. 

Distribution. New Guinea and Solomon Is. 

The type female is the only example known from Papua and has the abdomen 
missing, so that the identity of neither the New Hebridean nor the Solomon Islands 
specimens can be confirmed by examination of the genitalia until more topotypical 
material becomes available. Slight superficial differences are evident in coloration 
and markings of the fore wings of the examples examined from the three locahties. 

Stathmopoda imperator Bradley 

Stathmopoda imperator Bradley, 1957, Nat. Hist. Rennell I., Brit. Solomon Is. 2 (19) : 103. 

Aneityum : Red Crest, 1,200 ft., iii. 1955, 2 $. 
Distribution. Solomon Is. 

Stathmopoda electrantha Meyrick 

Stathmopoda electrantha Meyrick, 1927, Exot. Microlep. 3 : 377. 
Stathmopoda transvecta Meyrick, 1927, loc. cit. 

Aneityum : Red Crest, 1,200 ft., iii-vi.1955, 16 c^$. 
Distribution. New Hebrides and Solomon Is. 

Stathmopoda sp. 

Aneityum : Red Crest, 1,200 ft., iii, v and vi.1955, 5 (^. 

The specimens are in poor condition but apparently belong to a species having 
uniform yellowish-buff coloration on the fore wing with no heavy or extensive 
markings and with an orange or ferruginous patagia (collar). 

GLYPHIPTERYIGIDAE 
lmm.a ochrophara sp. n. 

(PI. 14, fig. 12) 

^J, 19 mm. Labial palpus light orange-yellow, second segment suffused drab below and at base 
exteriorly, paler interiorly ; terminal segment similarly suffused exteriorly. Head pale orange- 
yellow, crown suffused with drab. Antenna and scape light orange-yellow. Patagia pale orange- 
yellow suffused drab. Thorax and tegula drab, weakly violaceous, thorax with light orange- 
yellow posterior crest. Fore wing light orange-yellow, basal half except edge of costa wholly 
overlaid with drab, with a weak violaceous iridescence, distal edge of this area diffuse, strongly 
and evenly convex from before middle of costa to a point opposite on inner margin ; a large 
similarly coloured patch beyond middle in upper discal area, resting on costa, extending to or 
a little beyond middle, outer (distal) margin convex, inclined towards costa, followed by a line 
of darker interneural spots, inner (proximal) edge slightly outwardly-oblique, meeting lower 
(dorsal) edge at end of cell, a small darker spot in angle thus formed, a similar spot obliquely 
above in cell and set in margin of marking ; a blackish somewhat diffuse thin subterminal line ; 



MICROLEPIDOPTERA FROM THE NEW HEBRIDES 265 

cilia drab, darker basally. Hind wing blackish hair-brown, thinly scaled from beyond middle to 
base, semitransparent basally ; a yellowish hair-pencil in slight costal fold on upper surface near 
base ; cilia drab, darker basally. Legs light orange-yellow exteriorly, whitish interiorly ; 
middle and posterior tibiae marked with drab dorsally ; posterior tarsi wholly whitish. 
Male genitalia : PI. 23, fig. i. 

Holotype (J, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of Anel- 
gauhat, iv.1955 (L. E. Cheesman). Unique. Genitalia slide 6040. 

A distinctive species perhaps allied to the Solomon Islands species /, thyriditis 
Meyrick, from which it may be distinguished by the basal marking of the fore wing 
which in thyriditis is with the outer edge straight and outwardly-obHque from costa. 

Imtna oxystoma sp. n. 

(PI. 14, fig. 13) 

(J$, 15-17 mm. Labial palpus cartridge buff in male, cream-buff in female ; a weak admixture 
of fuscous-black interiorly ; male with second segment compressed laterally and proportionately 
widened, strongly suffused with fuscous-black exteriorly ; terminal segment modified into a 
line hair-like curved dark spine ; female second segment cylindrical, with a broad fuscous medial 
band ; terminal segment normal, cylindrical, obtuse, a small fuscous-black marking near base 
exteriorly. Head Front's brown ; anterior quarter of thorax and whole of tegula chestnut- 
brown, tegula somewhat irrorate apically with mummy brown mixed with a trace of whitish and 
elongated with hair-like mummy brown scales, middle of thorax whitish suffused with a mixture 
of chestnut-brown and grey, posterior of thorax fuscous-black, with a trace of whitish irroration 
in caudal margin. Fore wing chestnut-brown mixed with mummy brown, basal third overlaid 
with blackish, edged distally with scattered iridescent violet-plumbeous scales ; three moderately 
large well-defined mustard yellow elliptical spots with suffused fuscous centres on costa, first 
beyond blackish basal area, second a little beyond middle, third at about 3/4 ; an elongate 
transverse blackish bar at end of cell (along discocellulars), a sprinkling of iridescent violaceous 
scales beyond ; a small blackish dot contiguous with inner margin of third costal marking ; 
a similar pre-apical dash on costa ; termen outlined with a thin blackish line in which are set 
minute pale mustard yellow dots at end of radial veins ; cilia fuscous. Hind wing light fuscous, 
basal half thinly scaled, subhyaline ; cilia paler, with a fuscous basal line. 

Male genitalia : PI. 23, figs. 2 and 3. 

Female genitalia : PI. 24, figs, i and 2. 

Holotype ^, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, iii.1955 [L. E. Cheeseman). Genitaha sHde 4884. 

Allotype 9, same data. Genitalia slide 4875. 

Paratype i ^, same data. 

This species shares the peculiar modified labial palpus in the male with /. chelacma 
Meyrick, from Samoa, and also has veins 7 and 8 of the fore wing stalked, and vein 8 
to the costa. 

BLASTOBASIDAE 

Blastobasis anthoptera Lower 

Blastobasis anthoptera Lower, 1907, Trans. R. Soc. S. Austr. 31 : 118. 

Aneityum : Red Crest, 1,200 ft., iii-vi.1955, 2 (^, 9 ?. 

The identification of the above specimens is based on two examples of this species 
in the British Museum (Natural History) collection from the New Hebrides. Both 



266 J. D. BRADLEY 

examples have data labels " Efate I., Vila, vii.1925, P. A. Buxton" and were 
determined by Meyrick. 

Distribution. Australia (Queensland). 

Blastobasis sp. 

Aneityum : Red Crest, vi.1955, i ?. 

Superficially reminiscent of B. transcripta Meyrick, which occurs in China and 
India. 

YPONOMEUTIDAE 
Yponomeuta meracula sp. n. 

(PL 14, fig. 14) 

(J$, 21-24 n^"^- Labial palpus white, upper side of second segment and whole of basal 2/3 
of terminal segment overlaid with fuscous-black. Head, thorax and tegula wholly white. Antenna 
dark grey, whitish near base ; scape pure white. Fore wing smooth-scaled, slightly glossy, snow 
white ; anterior margin of costa strongly marked with fuscous-black basally, rather incon- 
spicuous viewed directly from above ; cilia matching. Hind wing mouse grey, lighter anteriorly 
and becoming white around apex and part of way along terminal margin, white or whitish in 
basal area and part of way along inner margin. Abdomen white. 

Male genitalia : PI. 25, figs, i and 2. 

Female genitalia : PI. 24, figs. 3-5. 

Holotype S, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, v. 1955 (L. E. Cheesman). 

Allotype $, same locality data, dated iii.1955. Genitalia slide 4873. 

Paratypes 6 (^, 13 $, same locality data, dated iii, v and vi.1955. Genitaha 
slide 4905 (male). 

This species is atypical of the genus in having the fore wing wholly white and 
unspotted and without markings except at base of anterior edge of costa. It is the 
sole representative of this genus known from the New Hebrides, and may be tenta- 
tively placed next to the Australian species H. paurodes Meyrick which is typically 
spotted. 

Anticrates dijflua hebridensis ssp. n. 

(PL 14, fig. 15) 

Anticrates difflua Meyrick, 1927, Insecta of Samoa, 3 (2) : 106. 

The examples of this species from the New Hebrides are intermediate super- 
ficially as well as geographically between the nominate race from Samoa and A . difflua 
rennellensis from the Solomon Is., and apparently represent a distinct subspecies. 
Their general coloration is brighter and stronger than in difflua difflua, but the bold 
reticulate markings of the fore wing do not attain the deep coral red coloration as in 
difflua rennellensis. The markings on thorax and head are likewise stronger than 
in the nominate form but are not the deep reddish colour as in rennellensis. The 
New Hebridean specimens are slightly larger — by 1-2 mm.— than specimens from 
Samoa and the Solomon Is. 



MICROLEPIDOPTERA FROM THE NEW HEBRIDES 267 

Male genitalia : similar to those of difflua difflua and d. rennellensts. The genitalia 
of both sexes of this species are illustrated in a previous paper (Bradley, 1957, Nat. 
Hist. Rennell I., British Solomon Is. 2 (19) : figs. 65-69). 

Holotype ^, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, v. 1955 {L. E. Cheesman). GenitaHa slide 4916. 

Paratype i <^, same data. 

PLUTELLIDAE 

Plutella maculipennis (Curtis) 

Cerostoma maculipennis Curtis, 1832, Brit. Entom. 9 (105), No. 420. 

Aneityum : Red Crest, 1,200 ft., v. 1955, i (^, i 9. 
Distribution. Cosmopohtan. 

Orthenches delosticha sp. n. 

(PI. 14, fig. 16) 

{J, II mm. Labial palpus cartridge buff, second segment suffused with fuscous exteriorly 
except at apex, an admixture of warm buff interiorly ; terminal segment tipped with fuscous- 
black. Head, thorax and tegula cartridge buff, lightly shaded with cream-buff, more especially 
at side of head ; lower part of front of head cartridge buff ; a narrow moderately distinct fuscous 
medial line on thorax. Fore wing cartridge buff thinly irrorate with blackish, more concentrated 
towards base ; veins diffusely etched with ochraceous-buff with variable admixture of drab ; 
a small blackish plical dash ; a similar round discal spot at 2/3 with a minute blackish speck 
immediately costad ; a pair of rather thick blackish dashes close together near middle of termen ; 
a single similar pre-apical dash on costa ; cilia warm buff, a cartridge buff medial line from apex 
to near tomus broadly edged with drab. Hind wing smoke grey ; cilia matching, cream-buff 
basally. Abdomen and legs light buff ; legs suffused greyish exteriorly. 

Male genitalia : PI. 25, figs. 3 and 4. 

Holotype (^, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, v. 1955 {L. E. Cheesman). Genitalia slide 6534. 

Paratypes 4 (^, .same locality data, dated iii and iv.1955. 

The geographical distribution of the species in the genus Orthenches Meyrick is of 
considerable interest. The genus is strongly developed in New Zealand, but is well 
represented in Australia and also by several species in South America (Chile and 
Argentina), and one species is known from Assam. 0. delosticha is superficially 
nearest the New Zealand .species 0. chlorocoma Meyrick, but is readily distinguished 
by the radiating nervure lines on the fore wing. The wing venation is slightly 
unstable, vein 2 of the fore wing being approximate, connate or very shortly stalked 
with vein 3 at the end of the cell. 

EPERMENIIDAE 

Epermenia symmorias Meyrick 
Epermenia symmorias Meyrick, 1923, Exot. Microlep. 3 : 52. 

Aneityum : Red Crest, 1,200 ft., vi.1955, i ^. 
Distribution. Fiji. 



268 J. D. BRADLEY 

LITHOCOLLETIDAE 
Acrocercops euthycolona Meyrick 

Acrocercops euthycolona Meyrick, 193 1, Exot. Microlep. 4 : 46. 

Aneityum : Red Crest, 1,200 ft., iv.1955, i $. 
Distribution. Java, India, Malaya and Solomon Is. 

Acrocercops cramerella (Snellen) 

Gracilaria \sic\ cramerella Snellen, 1904, Tidschr. Ent. 46 : 84. 

Aneityum : Red Crest, 1,200 ft., iii-vi.1955, 8 (^$. 
Distribution. Widespread in the Indo-Australian region. 

Caloptilia xanthopharella (Meyrick) 

Gracilaria [sic] xanthopharella Meyrick, 1880, Proc. Linn. Soc. N^.S.W. 5 : 141. 

Aneityum : Red Crest, 1,200 ft., iv.1955, i 9. 
Distribution. Fiji and E. Australia. 

lyonetiidae 

Opogona mendanai Bradley 

Opogona mendanai Bradley, 1961, Bull. Brit. Mus. {nat. Hist.), Ent. 10 : 161. 

Aneityum : Red Crest, 1,200 ft., iii.1955, i $. 
Distribution. Solomon Is. 

Anastathma callichrysa Meyrick 

Anastathma callichrysa Meyrick, 1886, Trans, ent. Soc. Lond. 1886 : 290. 

Aneityum : Red Crest, 1,200 ft., vi.1955, 4 ex. 
Distribution. Fiji. 

Ereunetis capnosticta sp. n. 

(PI. 14, fig. 17) 

(J$, lo-ii mm. Labial palpus in male loosely scaled, apical segment tufted with long fine 
scales ; in female normal, scales closely appressed, slightly roughened beneath, apex of terminal 
segment obtuse and roughened ; in both sexes cream-buff, terminal segment in female strongly 
suffused with fuscous ; terminal segment in male with a small fuscous patch on upper margin at 
base. Head cream-buff, crown sometimes mixed with greyish. Antennal scape cream-buff ; 
flagellum greyish fuscous. Thorax cream-buff, a diffuse hair-brown or fuscous medial line and a 
similar weak suffusion laterally. Tegula cream-buff strongly suffused with hair-brown or 
greyish fuscous. F'ore wing cream-buff sparsely and irregularly irrorate with hair-brown or 
greyish fuscous scales ; markings consisting mainly of diffuse, somewhat irrorate, small hair- 
brown or greyish fuscous subquadrate or rounded patches scattered irregularly over the wing, 
usually larger and often darker near the base, in strongly marked specimens these markings are 



MICROLEPIDOPTERA FROM THE NEW HEBRIDES 269 

more extensive and occasionally coalesce ; an elongate patch at base of costa ; a blackish apical 
dot ; cilia cream-buff suffused with hair-brown except at apices, a faint sub-basal line. Hind 
wing somewhat shining light drab, darker and weakly infuscate along inner margin distally ; 
cilia hair-brown ; male with a specialized pencil of fine cream-buff hairs arising near base a 
little below costa and reaching beyond middle. Legs cream-buff suffused with hair-brown 
exteriorly. Abdomen greyish hair-brown, paler beneath. 

Male genitalia : PI. 26, figs, i and 2. Aedeagus with a small internal sheaf of about 5 or 6 
minute cornuti. 

Female genitalia : PI. 26, fig. 3. 

Holotype rS, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, iv.1955 {L. E. Cheesman). Genitalia slide 6537. 

Allotype $, same data. Genitalia slide 6582. 

Paratypes 21 <^, 7 ?, same locality data, dated iii, iv and vi.1955. 

Additional material : 285 cJ?, same data as type material. These are mostly in 
damaged condition and have not been used for descriptive purposes. 

This species is near E. intertexta Bradley, which occurs in the Solomon Is. It may 
be distinguished from that species by the absence of the light quaker drab and yellow 
coloration present in intertexta in the distal area of the fore wing. The specialized 
hair-pencil on the upper surface of the hind wing in capnosticta is not present in 
intertexta. 

Decadarchis eustropha sp. n. 

(PI. 14. fig. 18) 

cJ$, lo-ii mm. Labial palpus white, second segment marked with fuscous exteriorly to near 
apex, suffused greyish interiorly, bristles dark brown. Head white or sordid white. Antennal 
scape white, sometimes an admixture of fuscous below and anteriorly ; flagellum fuscous-black, 
white-scaled ventrally ; a thin conspicuous white anterior line composed of a chain of very 
slender white scales. Thorax and tegula white overlaid with a mixture of mummy brown and 
fuscous, darker towards posterior of thorax and base of tegula. Fore wing with apex upturned ; 
white ; markings blackish mummy brown ; a small dense patch at base of costa extending along 
anterior edge to a broad, well-defined, solid, transverse fascia from between i /4 to near middle ; 
dorsal half of wing proximad to this fascia diffusely irrorate with fuscous ; an inwardly-oblique, 
diffuse, ill-defined fascia at about 2/3 dilated distally and occupying most of apical half, inner 
edge diffusely irrorate more or less straight from costa at about 3/4 to near middle of inner margin 
(dorsum) where it converges with the prominent submedial fascia ; an intense subapical spot ; 
cilia white irregularly irrorate with fuscous, a blackish mummy brown basal line around apex. 
Hind wing whitish grey ; cilia paler. Legs whitish, tarsi marked with blackish fuscous exteriorly. 

Male genitalia : PI. 27, figs, i and 2. 

Female genitalia : PI. 26, figs. 4 and 5. 

Holotype (^, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, v. 1955 (L. E. Cheesman). Genitaha slide 6535. 

Allot\^pe $, same locahty data, dated vi. 1955. Genitaha shde 6581. 

Paratypes 3 9, same locality data, dated iv, v and vi.1955. 

Near D. discreta Meyrick, which occurs from the Solomon Is. south-west to New 
Guinea, and distinguished by its smaller size, discreta having a wing expanse of 
16-21 mm., and also by the well-defined directly transverse fascia on the fore wing 
^t 1/3, which in discreta is slightly outwardly-oblique, dilated at inner margin of 
wing, and connected medially to the distal fascia. 



270 J. D. BRADLEY 

Decadarchis heterogramma Meyrick 

Decadarchis heterogramma Meyrick, 192 1, Exot. Microlep. 2 : 459. 

Aneityum : Red Crest, 1,200 ft., iv and vi.1955, i c^, 3 $. 
Distribution. Fiji. 

Decadarchis flavistriata (Walsingham) 

Ereunetis flavistriata Walsingham, 1907, Fauna Hawaiiensis, 1 pt. 5 : 716. 
Decadarchis euophthalma Meyrick, 1924, Exot. Microlep. 3 : 83. 

Aneityum : Red Crest, 1,200 ft., iv.1955, i ^. 

Distribution. Hawaiian Is., Solomon Is., Java, Malaya, Fiji, Marquesas, New 
Hebrides and Kermadec Is. 

Decadarchis synclera sp. n. 

(PI. 14, fig. 19) 

cj, 16 mm. Labial palpus whitish, irrorated fuscous or fuscous-black exteriorly, beneath with 
long rough whitish hair-scales throughout, bristles dark brown. Head, thorax and tegula white, 
crown of head tinged light buff, base of thorax overlaid with fuscous or fuscous-black. Antenna 
whitish ; scape diffusely irrorate fuscous ; flagellum with weak greyish suffusion. Fore wing 
ochreous-white ; a thick irregular diffuse fuscous-black streak from base of costa above middle 
to 2/3, lower edge biundulate ; costal area above streak suffused fuscous ; an elongate fuscous- 
black spot on costa at 3/4 ; a diffuse fuscous-black medial streak from end of cell to apex ; 
some light fuscous speckling on dorsal and distal areas ; a conspicuous small black apical spot ; 
a well-defined fuscous-black marginal line from costa well before apex skirting apical spot and 
along termen ; cilia ochreous-white, fuscous opposite apical spot. Hind wing greyish white ; 
cilia concolorous. 

Male genitalia : PI. 27, figs. 3 and 4. 

Holotype cJ, New Hebrides : Aneityum, Red Crest, 1,200 ft., 3 m. NE. of 
Anelgauhat, iv.1955 {L. E. Cheesman). Genitalia slide 6525. 

Paratype i (^, same locality data, dated iii.1955. 

Very closely related to D. ancostyla Meyrick, which occurs in Samoa. The super- 
ficial differences between these two species are slight and difficult to elucidate 
because of the poor condition of the material examined, but the fore wing markings 
of the Samoan species are heavier. The differences in the male genitalia are also 
small, in ancostyla the ventral part of the valva is pointed at apex not rounded as in 
synclera, and the saccus is similarly tapered to a point and not rounded ; and the 
dense, compact, cylindrical patch of cornuti are weaker and less sclerotized in 
synclera. 

Decadarchis capnographa Meyrick 
Decadarchis capnographa Meyrick, 1928, Exot. Microlep. 3 : 402. 

Aneityum : Red Crest, 1,200 ft., iii-vi.1955, 2 cJ, 9 $. 
Distribution. New Hebrides. 



MICROLEPIDOPTERA FROM THE NEW HEBRIDES 271 

Decadarchis semifusca Bradley 

Decadarchis semifusca Bradley, 1961, Bull. Brit. Mus. {nat. Hist.), Ent. 10 : 165. 

Aneityum : Red Crest, 1,200 ft., iii-vi.1955, i 3", 10 $. 
Distribution. Ontong Java Atoll (Solomons Group). 

TINEIDAE 
Gerontha captiosella Walker 

Gerontha captiosella Walker, 1864, List Lep. Ins. B.M. 29 : 782. 

Aneityum : Red Crest, 1,200 ft., iii-vi.1955, 6 $. 

Distribution. Occurring throughout the I ndo- Australian region. 

Tinea sp. 

Aneityum : Red Crest, 1,200 ft., iii and vi.1955, 2 (^, i $. 

These three specimens represent a species of unusual appearance but with typical 
male genitaUa of the genus. All are in poor condition and have labial palpi and 
antennae damaged, and are therefore not considered suitable for descriptive 
purposes. 




ENTOM. 12, 5. 21 



PLATE 14 



Fig. I. Nesoscopa psarodes sp. n., holotype ^ 

Fig. 2. Eucosma euryochra sp. n., holotype (J 

Fig. 3. Strepsicrates poliophora sp. n., holotype cj . 

Fig. 4. Mesocallyntera dascia sp. n., holotype ^ 

Fig. 5. Cryptophlebia iridoschema sp. n., holotype $ 

Fig. 6. Schoenotenes elaphrodes sp. n., holotype (J . 

Fig. 7. Idiophantis lomatographa sp. n., holotype $ 

Fig. 8. Autosticha silacea sp. n., allotype $ 

Fig. 9. Peragrarchis minima sp. n., allotype $. 
Fig. 10. Meridarchis celidophora sp. n., holotype $ . 

Fig. II. Heterogymna cheesmanae sp. n., holotype,^. 

Fig. 12. Imma ochrophara sp. n., holotype cj 

Fig. 13. Imma oxystoma sp. n., holotype ^ 

Fig. 14. Yponomeuta meracula sp. n., holotype cj 

Fig. 15. Anticrates difflua hebridensis ssp. n., holotype (J 

Fig. 16. Orthenches delosticha sp. n., holotype (J 

Fig. 17. Ereunetis capnosticta sp. n., holotype (^ 

Fig. 18. Decadarchis eustropha sp. n., allotype ? 

Fig. 19. Decadarchis synclera sp. n., holotype ^ 



Wing 
expanse 

of 
species 

(mm.) 

11-13 

15-19 

18 
24-27 

15 

15 

14 
12-14 

14-15 
18-22 

/ c?30-32 
\ $36-40 
19 
15-17 
21-24 
15-X6 
II 

lO-II 
lO-II 

16 



Bull. B.M. (N.H.) Entom. 12, 5 




16 

ENTOM. 12, 5. 



X 17 



PLATE 15 

Fig. I. Neoscopa psarodes sp. n., $ genitalia, ventral view (Slide 5526). 

Fig. 2. Ibidem, (^ genitalia with aedeagus removed, ventral view (Slide 5515). 

Fig. 3. Ibidem, aedeagus, lateral view. 

Fig. 4. Eucosma euryochra sp. n., ^ genitalia with aedeagus in situ, ventral view (Slide 
4846). 



Bull. B.M. (N.H.) Entom. 12. 5 



PLATE 15 




PLATE i6 



Fig. I. Eucosma euryochra sp. n., $ genitalia, ventral view (Slide 5525). 

Fig. 2. Ibidem, ostium. 

Fig. 3. Ibidem, enlargement showing detail of one of the pair of signa. 

Fig. 4. Cryptophlebia iridoschema sp. n., $ genitalia, ventral view (Slide 6051). 



Bull. B.M. (N.H.) Entom. 12, 5 



PLATE 16 




-\S 




PLATE 17 

Fig. I. Strepsicrates poliophora sp. n., (^ genitalia with aedeagus in situ, ventral view (Slide 
4883). 

Fig. 2. Autosticha silacea sp. n., ^ genitalia with aedeagus removed, ventral view (Slide 
6038). 

Fig. 3. Ibidem, aedeagus, lateral view. 



Bull. B.M. (N.H.) Eniom. 12, 5 



PLATE 17 






PLATE i8 

Fig. I. Mesocallyntera dascia sp. n., ^ genitalia with acdeagus in situ, ventral view (Slide 
6020). 

Fig. 2. Schoenotenes elaphrodes sp. n., ,^ genitalia with aedeagus removed, ventral view 
(Slide 5516). 

Fig. 3. Ibidem, aedeagus, lateral view. 



I 



Bull. B.M. {N.H.) Entoni. 12. 5 



PLATE I 




ENTOM. 12, 5 



23 



PLATE 19 

Fig. I. Idiophantis lomatographa sp. n., $ genitalia, ventral view (Slide 6580). 

Fig. 2. Ibidem, ostium. 

Fig. 3. Meridarchis celidophora sp. n., § genitalia, ventral view (Slide 6054). 



i 



Bull. B.M. {N.H.) Entom 12, 5. 



PLATE 19 




PLATE 



Fig. I. Autosticha silacea sp. n., $ genitalia, ventral view (Slide 6577). 

Fig. 2. Ibidem, enlargement showing detail of one of the pair of signa. 

Fig. 3. Heterogymna cheesmanae sp. n., $ genitalia, ventral view (Slide 482! 

Fig. 4. Ibidem, enlargement showing detail of one of the pair of signa. 



Bull. B.M. (N.H.) Entoni. 12, 5 



PLATE 20 




PLATE 21 

Fig. I. Peragrarchis minima sp. n., $ genitalia, ventral view (Slide 6055). 
Fig. 2. Ibidem., ostium. 
Fig. 3. Ibidem, signum. 



I 



Bull. B.M. [N.H.) Entom. 12, 5 



V L A T E 2 I 




PLATE 22 

Fig. I. Peragrarchis minima sp. n., ^ genitalia with aedeagus removed, ventral view (Slide 
6029). 

Fig. 2. Ibidem, aedeagus, lateral view. 

Fig. 3. Heterogymna cheesmanae sp. n., (^ genitalia with aedeagus removed, ventral view 
(Slide 4890). 

Fig. 4. Ibidem, aedeagus, lateral view. 



I 



Bull. B.M. (N.H.) Entoni. 12, 5. 



PLATE 22. 




ENTOM. 12. 5. 



24 



PLATE 23 

Fig. I. Imma ochrophara sp. n., ^ genitalia with aedeagus in situ, ventral view (Slide 6040). 
Fig. 2. Imma oxystoma sp. n., ^J genitalia with aedeagus removed, ventral view (Slide 4884). 
Fig. 3. Ibidem, aedeagus, lateral view. 



Bull. B.M. (N.H.) Entom. 12, 5 



PLATE 23 




ENTOM. 12, 5. 



24§ 



PLATE 24 

Fig. I. Imma oxy stoma sp. n., § genitalia, ventral view (Slide 4875). 

Fig. 2. Ibidem, signum. 

Fig. 3. Yponomeuta meracula sp. n., $ genitalia, ventral view (Slide 4873). 

Fig. 4. Ibidem, signum. 

Fig. 5. Ibidem, ostium. 



Bull. B.M. (N.H.) Eniom. 12, 5 



PLATE 24 




PLATE 25 

Fig. I. Yponomeuia meracula sp. n., (^ genitalia with aedeagus removed, ventral view 
(Slide 4905). 

Fig. 2. Ibidem, aedeagus, lateral view. 

Fig. 3. Orthenches delosticha sp. n., ^ genitalia with aedeagus removed, ventral view (Slide 

6534)- 

Fig. 4. Ibidem, aedeagus, lateral view. 



Bull. B.M. (N.H.) Entom. 12, 5 



PLATE 25 




PLATE 26 

Fig. I. Ereunetis capnosticta sp. n., ^ genitalia with aedeagus removed, ventral view (Slide 

6537)- 

Fig. 2. Ibidem, aedeagus, lateral view. 

Fig. 3. Ibidem, $ genitalia, ventral view (Slide 6582). 

Fig. 4. Decadarchis eustropha sp. n., 9 genitalia, excluding bursa copulatrix and portion of 
ductus bursae, ventral view (Slide 6581). 

Fig. 5. Ibidem,, bursa copulatrix and portion of ductus bursae. 



i 



Bull. B.M. (N.H.) Entom. 12, 5 



PLATE 26 




PLATE 27 

Fig. I. Decadarchis eustropha sp. n., (^ genitalia with aedeagus removed, ventral view 
(Slide 6535). 

Fig. 2. Ibidem, aedeagus, lateral view. 

Fig. 3. Decadarchis synclera sp. n., (J genitalia with aedeagus removed, ventral view (Slide 

6525)- 

Fig. 4. Ibidem, aedeagus, lateral view. 



I 



Bull. B.M. (N.H.) Entom. 12, 5. 



PLATE 27. 




PRINTED IN GREAT BRITAIN BY 
ADLARD AND SON, LIMITED 

BARTHOLOMEW PRESS, DORKING 



( 



THE ACRIDOIDEA (ORTHOPTERA) 

OF MADAGASCAR 
I. ACRIDIDAE (EXCEPT ACRIDINAE) 




V. M. DIRSH 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 12 No. 6 

LONDON: 1962 



THE ACRIDOIDEA (ORTHOPTERA) OF 

MADAGASCAR 

I. ACRIDIDAE (EXCEPT ACRIDINAE) 



BY 

V. M. DIRSH 



V- 

Anti-Locust Research Centre, London 



H 



Pp. 273-350 ; 40 Text-figures 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 12 No. 6 

LONDON: 1962 



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 
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within one calendar year. 

This paper is Vol. 12, No. 6 of the Entomological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 



Tiustees of the British Museum 1962 



PRINTED BY ORDER OF THE TRUSTEES OF 
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Issued 17 July 1962 Price Twenty-six shillings 



THE ACRIDOIDEA (ORTHOPTERA) OF 

MADAGASCAR 

I. ACRIDIDAE (EXCEPT ACRIDINAE) 

By V. M. DIRSH 

SYNOPSIS 

The subfamilies Hemiacridinae, Oxyinae, Coptacridinae, Calliptaminae, Eyprepocnemidinae, 
Catantopinae and Cyrtacanthacridinae of the family Acrididae (Acridoidea, Orthoptera) of 
Madagascar are revised. Eleven new genera and nineteen new species are described. Some 
inadequately described genera and species are redescribed and synonymy is checked. Inter- 
relations between species and genera are briefly discussed. 

INTRODUCTION 

In 1956 Dr. R. Paulian, Directeur-adjoint de I'lnstitut de Recherche Scientifique 
de Madagascar (at present Directeur de I'lnstitut d'Etudes Centre-africaines) 
invited me to study the Acridoidea of Madagascar. This offer was gratefully 
accepted and the present paper is the first of the preliminary studies which are 
essential before a full review of the Madagascar fauna can be undertaken. 

This part deals with representatives of subfamilies of the Acrididae, other than 
Acridinae, so far found in Madagascar. The Acridinae and other families of 
Acridoidea will form the subject of further studies when more material has been 
accumulated. 

The material so far studied suggests that probably less than half the genera and 
species of Madagascar Acridoidea are known. The fauna of the island is extremely 
peculiar. It contains some African elements, which probably reached Madagascar 
relatively recently, but most of it consists of purely endemic genera and species. 
From the taxonomic point of view many of the genera are so pecuhar that they are 
difficult to place definitely into any of the known subfamilies. From the zoogeo- 
graphical point of view this endemic fauna suggests affinites with the Acridoidea of 
the Oriental rather than the Ethiopian Region. However, the discussion of this 
question will have to be postponed for the time being. 

Endemic genera and species are redescribed here and many illustrated for the 
first time. 

I wish to express my sincere gratitude to the following persons : Dr. L. Chopard 
(Paris Museum), Mr. E. Morales- Agacino (Madrid Museum), and Dr. Ch. Ferriere 
(Geneva Museum), for providing the types, which were essential for the studies in 
this paper. I am also grateful to Dr. B. P. Uvarov for his valuable advice in the 
course of preparation of this paper, Dr. T. H. C. Taylor, Director of the Anti-Locust 
Research Centre, and Mr. J. P. Doncaster, Keeper of the Department of Entomology 
of the British Museum (Natural History) for the editorial work on the manuscript. 

ENTOM. 12, 6. 25 



276 V. M. DIRSH 

Subfamily HEMIACRIDINAE 

The Madagascar representatives of this subfamily are of peculiar appearance, 
sometimes strikingly so. They also exhibit a tendency towards the reduction and 
even loss of the stridulatory veinlets of the elytron characteristic for the subfamily. 
Sometimes some of the specimens of the same species have clearly detectable, though 
reduced, stridulatory veinlets, while others have lost them completely or almost 
completely. 

The same instability of subfamily characters was observed in the structure of 
the penis. While some of the species have clearly disconnected basal and apical 
penis valves, typical for the subfamily, others have a tendency to form a thin 
flexure, which sometimes, although thin, is rather strongly sclerotized, and in one 
case is rather thick and robust. 

This makes the Hemiacridinae of Madagascar somewhat isolated from other 
members of the subfamily. It is not clear whether they are losing the characteristic 
features of the subfamily or gaining them. 

Although the majority of genera in the subfamily are apparently not closely 
related, some of them show affinities definite enough to allow their formation into 
groups (e.g. Euthymia Group, Gergis Group). Much further study is needed to 
establish the taxonomic value of these groups and their interrelationship with other 
genera. 

Key to Genera 

Antenna compressed, slightly ensiform. Male subgenital plate as long or longer 

than pronotum (Text-fig. i) LEPTACRIS (p. 277) 

Antenna filiform or widened in basal part, but not compressed, male subgenital 
plate much shorter than pronotum. 

Elytra lobiform, lateral, vestigial or absent. 

Elytra lobiform, lateral. 

Dorsum of pronotum smooth, without convexities. 

Posterior margin of pronotum excurved (Text-fig. 2) SPATHOSTERNUM (p. 278) 

Posterior margin of pronotum incurved (Text-fig. 3) 

PARASPATHOSTERNUM (p. 279) 

Dorsum of pronotum rugose with large convexities (Text-fig. 4) 

PSEUDOSERPUSIA (p. 281) 

Elytra vestigial or absent. 

Fully apterous. Antenna filiform (Text-fig. 6) . . . HYSIELLA (p. 284) 

Male with vestigial elytra, female apterous. Antenna widened in basal part 

(Text-fig. 13) XENIPPOIDES (p. 298) 

Elytra fully developed or shortened, but not lobiform. 

Cubital vein of hind wing not connected with medial. 

Head strongly prognathous. Whole body covered with long hair, its lower 
surface concave. Prosternum, without process, with collar-like projecting 
anterior margin (Text-fig. 5) PAULIANACRIS {p. 282) 

15 (14) Head orthognathous or opisthognathous or slightly prognathous. Body only 

slightly hairy or not hairy, its lower surface not concave. Prosternum with 
process, without collar. 

16 (17) Body strongly elongated, slender. Head acutely conical, opisthognathous ; 

fastigium of vertex strongly elongated, narrow (Text-fig. 12) GALIDEUS (p. 296) 



I 


(2) 


2 


(I) 


3 


(12) 


4 


(9) 


5 


(8) 


6 


(7) 


7 


(6) 


8 


(5) 


9 


(4) 


10 


(II) 


II 


(10) 


12 


(3) 


13 


(24) 


14 


(15) 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



277 



17 (16) 



18 


(23) 


19 


(20) 


20 


(19) 


21 


(22) 


22 


(21) 


23 


(18) 


24 


(13) 


25 


(26) 


26 


(25) 


27 


(30) 


28 


(29) 


29 


(28) 


30 


(27) 



Body moderately elongated and not slender. Head subconical to subglobular, 

orthognathous or slightly prognathous. Fastigium of vertex short, wide. 
Pronotum cylindrical or subcylindrical without convexities. 
Pronotum subcylindrical ; transverse sulci comparatively wide ; integument 

slightly rugose (Text-figs. 7, 8) ONETES (p. 284) 

Pronotum cylindrical ; transverse sulci fine ; integument smooth. 
Male supra-anal plate elongate, with acutangular apex (Text-fig. 9) 

EUTHYMIA (p. 290) 
Male supra-anal plate short, wide, with obtusangular apex (Text-fig. 10) 

PROEUTHYMIA (p. 293) 
Pronotum tectiform, with tubercle-like projections (Text-fig. 11) 

LOPHEUTHYMIA (p. 294) 
Cubital vein of hind wing connected with medial (Text-fig. 18). 
Antenna short, widened in basal part and slightly club-like, thickened at apex 

(Text-fig. 17) PACHYCERACRIS (p. 305) 

Antenna long, filiform on whole length. 

Precostal and vannal areas of elytra without regular, parallel, transverse, 

oblique veinlets. 
External apical spine of hind tibia present. Stridulatory veinlets of radial 

area of elytra reduced (Text-fig. 14) . . MALAGAS ACRIS (p. 300) 

External apical spine of hind tibia absent. Stridulatory veinlets of radial area 

of elytra greatly reduced or absent (Text-figs. 15, 16) . GERGIS (p. 302) 

Precostal and vannal areas of elytra with regular, parallel, transverse, oblique 

veinlets (Text-fig. 18) MORONDAVIA (p. 306) 



LEFT ACRIS Walker. 1870 

Leptacris Walker, 1870 : 676. 
I schinacrida St&l, 1873 : 53 ; Uvarov, 1944 : 18. 
Metapa St&l, 1878 : 51, 97 ; Dirsh, 1958 : 239. 
Rhamphacrida Karsch, 1893 : 112 ; Dirsh, 1958 : 239. 
Pretoriana Uvarov, 1922 : 99 ; Uvarov, 1944 • ^^• 



Leptacris hova (Karsch, 1896) 
(Text-fig. i) 

I schinacrida hova Karsch, 1896 : 289. 

I schinacrida pulchra Karny, 1907 : 300, syn. n. 

Leptacris hova (Karsch, 1896) ; Uvarov, 1944 : 18. 

Both species named above were inadequately described. The only difference 
which could be found in the descriptions is in the coloration of internal side of hind 
femur, in /. hova being " infuscatis " and in I. pulchra " luteo rubra, concolora 
(cJ) vel viridi-coerulea, pallidipunctata (?) ". In the series of specimens studied 
the colour of the internal side of hind femur varies from reddish, purplish, bluish 
and brownish, to brown (which is possibly a post mortem change). Small spots, 
whitish or pale but of the same colour as background, may be present or absent ; 
the same colour variability exists in L. violacea (Karny, 1907) and L. monteiroi 
(I. Bolivar, 1820). No structural differences between the two synonymized species 
exist and it may be safely assumed that both belong to the same species. 



278 



V. M. DIRSH 



It should be noted that L. hova is very near to the Airican L. monteiroi, the only 
difference between them being that in L. monteiroi the head is sHghtly shorter than 
in L. hova and L. hova has slightly wider antennae. The phallic complex is of exactly 
the same structure in both of them. It is possible that they represent only subspecies. 

Madagascar Centre: Tananarive Tsimbazaza, 17.viii.1947, i (J (R. Paulian) ; 
Miarinarivo, viii.1958, i (^ nymph (A. Robinson) ; Tsiraonomandidy, sommet de 
I'Ambohiby, 26. v, 1948, i $, nymph (R. Paulian) ; Ambatofinandrahana, 1,180 m., 
26.vii.1957, I ^ (P. Griveaud). 

Madagascar Quest : VHP Reserve Naturelle, Soalala, viii, i $ (R. Paulian) ; 
Namoroka, i (^ (Rakotovao) ; foret de I'Ankarafantsika, Ampijoroa, xii.1951, 

I ? (P. Viette). 

Madagascar Est : Manakara, 15 km. vers Vohipeno, xi.1957, i $ (Jean Elie). 



SPATHOSTERNUM Krauss, 1877 
Spathosternutn malagassum sp. n. 

(Text-fig. 2) 

(^. Type. This species is very near to Spathosternum curium Uvarov, 1953 from Angola, 
from which it differs by sUghtly shorter, lobiform elytra, not overlapping on dorsal side and 
with more reduced and rougher venation, and by better developed and regularly incurved 
lateral carinae of pronotum. The phallic complex is very similar and differs only in small 
details, such as the relative size of the valves of penis. Epiphallus rather similar. 

$. Unknown. 

Length of body 12-2 ; pronotum 2-3 ; elytron 2-6 ; hind femur 6-2 mm. 

Madagascar Centre : Ankazobe, foret d'Ambohitantely, 26. ii. 1948, i ^ type, 2,^ 
paratypes (R. Pauhan). 

Type in Paris Museum. One paratype in the British Museum (Natural History). 




Fig. I. Leptacris hova (Karsch, 1896). Male, 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 279 

PARASPATHOSTERNUM Ramme, 1929 
Paraspathosternum andringitra sp. n. 

(Text-fig. 3) 

cj. Type. Body of medium size, cylindrical, in female slightly widened in middle. Integu- 
ment finely rugose and shiny. Antenna twenty-segmented, thick, filiform, shorter than head 
and pronotum together. Head conical ; fastigium of vertex widely angular, short, slightly 
concave, with obtuse marginal carinulae ; interocular distance wider than basal antennal 
segment ; along whole head there is an obtuse median cannula ; frons strongly oblique, frontal 
ridge deeply sulcate, with thick, obtuse lateral carinulae, almost parallel on whole length and 
at apex fused into narrow ridge. Pronotum sybcylindrical ; dorsum flattened, crossed by 
three sulci ; median and lateral carinae poorly developed, obtuse, almost parallel ; metazona 
less than half length of prozona, its posterior margin obtusangularly incurved. Prosternal 
process spathulate, with deeply bilobate apex ; mesosternal interspace longer than its width, 
slightly constricted in the middle. Elytra lobiform, lateral, with convex surface, reaching middle 
of second or third abdominal tergite ; venation and reticulation thick and rough ; apex subacute. 
Hind femur moderately slender ; lower lobe of hind knee rounded ; external apical spine of 
hind tibia present. Arolium large. Male supra-anal plate short, angular. Cercus straight, 
narrow-conical. Subgenital plate short, subconical. Phallic complex slender. Apodeme 
horseshoe-shaped ; rami narrow ; sheath of penis present ; basal and apical valves of penis 
connected by flexure-like thin connection ; gonopore processes large ; apical valves of j>enis 
small ; valves of cingulum large, with apical extension ; arch of cingulum large. Epiphallus 




Fig. 2. Spathosiernum malagassum sp. n. i, male. 2, prosternal process. 3, phallic 
complex, with ectophallic membrane and epiphallus removed. 4, the same with most 
of cingulum removed. 5, epiphallus. 



28o 



V. M. DIRSH 



bridge-shaped, with wide bridge, small angular ancorae and moderately large lobiform lophi. 
General coloration dark brown. Gena behind eye with ochraceous stripe ; lower part of gena 
with another less developed ochraceous stripe ; lateral carinae of pronotum and lower part of 
lateral lobes light brownish ; pleura with longitudinal, dirty ochraceous stripe ; elytra brown, 
in basal part light, in apical part dark brown ; lower lobe of hind knee ochraceous ; hind tibia 
reddish, with wliitish spines, brown at apices. 

$. Paratype, as male, but larger, with more widened middle part of body. Subgenital plate 
slightly trilobate, with angular middle lobe. Valves of ovipositor slender, with slightly curved 
apices. 

Length of body ^ I7'2, $ 26 ; pronotum ^ 3-2, $ 5-5 ; elytron cj 4, $ 6-6 ; hind femur (J 8-8, 
$ 13 mm. 

Madagascar Centre: Ambalavao, Cirque Boby, 2,500 m., Andringitra, i.1958, 
I (^ type, 2 $ paratypes (R. Paulian). Type and paratype in Paris Museum. One 
female paratype in the British Museum (Natural History). 




Fig. 3. Paraspathosternum andringitra sp. n. i, male, type. 2, prostemal process. 
3, phallic complex, with ectophallic membrane and epiphallus removed. 4, the same, 
with most part of cingulum removed. 5, epiphallus. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



281 



The new species externally is very similar to the Airican Paraspathosternum 
pedestris (Miller, 1929). The phallic complex however is rather different. Flexure- 
like connection between basal and apical valves of penis, which is present in andringi- 
tra is absent in pedestris. Shape of apical valves of penis is different, and valves of 
cingulum are highly complicated in andringitra, but small and simple in pedestris. 
It is possible that they belong to different genera. 

PSEUDOSERPUSIA gen. n. 

Of medium size. Integument rugose and shiny. Antenna filiform, longer than head and 
pronotum together. Head slightly inflated, above subglobular ; fastigiuni of vertex strongly 
sloping forwards, angular, with obtuse, almost truncate apex, longitudinally concave ; inter- 
ocular distance as wide as second basal antennal segment ; frons oblique and slightly incurved ; 
frontal ridge low, flat, in lower half almost obliterated. Eyes large, strongly convex. Pronotum 
subcylindrical, strongly rugose ; dorsum crossed by four deep, wide, transverse sulci ; median 
carina obtuse, forming between first and second, and between third and fourth sulci tubercle- 
like convexities, second being higher than first ; metazona about half length of prozona, its 
posterior margin slightly excurved, almost straight. Prosternal process widely conical, with 
acute apex. Mesostemal interspace wider than its length. Elytra lobiform, lateral, with 
rounded ai>ex, reaching middle of second abdominal tergite. Tympanum vestigial. Hind 




Ejs Gpr 



Fig. 4. Pseudoserpusia polychroma sp. n. i, male, type. 2, end of male abdomen 
from above. 3, phallic complex, ectophallic membrane and epiphallus removed, 
lateral view. 4, the same, with most part of cingulum and sheath of penis removed. 
5, epiphallus. 



282 V. M. DIRSH 

femur slender, with upper carina shallowly serrated ; external apical spine of hind tibia absent ; 
tarsus slightly elongated. Arolium large. Last abdominal tergite of male with pair of small 
acute projections ; supra-anal plate angular, with three pairs of lateral and one pair of medial 
teeth ; cercus acutely conical, straight ; subgenital plate short, obtusely conical. Valves of 
penis paired, divided, not forming flexure ; apical valves of penis and valves of cingulum large 
and strong ; sheath of penis large, wide, poorly sclerotized ; rami of cingulum narrow, strong, 
not connected on ventral side. Epiphallus with wide bridge ; small, wide ancorae and strong, 
lobiform lophi with subacute apices. Valves of ovipositor moderately slender, curved at 
apices. Subgenital plate trilobate. 

Type species : Pseudoserpusia polychroma sp. n. 

The new genus superficially resembles representatives of Serpusia group, but 
according to its genitalia, with disconnected valves of penis, belongs in the subfamily 
Hemiacridinae. It shows no close affinity with any known genus or group of genera. 
In the structure of pronotum it resembles the genus Perineta (see p. 342), but the 
similarity is only superficial, the genitalia being entirely different. 



Pseudoserpusia polychroma sp. n. 

(Text-fig. 4) 

(J. Type. Head strongly rugose ; weak occipital carinula present. Sides of mesosternal 
interspace roundly incurved ; metasternal interspace slightly longer than its width. Lower 
lobes of hind knee angular. General coloration ochraceous. Antenna black, with two basal 
segments ochraceous and two apical segments greenish. On sides of head and pronotum with 
wide, blackish longitudinal stripe, gradually fading towards base of hind femur. Elytron 
ochraceous, with brown anterior and posterior margins. Hind femur dark olive-green on both 
sides ; hind knee, tibia and tarsus bright red. Teeth on supra-anal plate black. 

$. Paratype. As the male, but larger. 

Length of body cj 19, $ 25-5 ; pronotum ^ 5-7, $ 7-5 ; elytron ^ 3-2, 9 5 ; hind femur ^ 13-6, 
$ 17 mm. 

Madagascar Est : Maroantsetra, Ambohitsitondroina, 600 m., iii.1958, i J type 
(P. Soga, E. Raharizonina) , in Paris Museum. 
Madagascar Centre : Tananarive, ii-iii.1950, 2 c?, i ?• 

PAULIANACRIS gen. n. 

Large and slender. Integument finely rugose, extremely hairy. Antenna thin, filiform. 
Head strongly prognathous, subglobular above ; fastigium of vertex wide, roundly merging 
with frons, which is very short ; frontal ridge low, almost obliterated ; ocelli large, lateral ones 
located above bases of antennae ; eyes small, oval, flattened. Pronotum saddle-shaped ; 
dorsum crossed by three sulci, posterior one deep and strong ; carinae absent ; metazona 
slightly longer than prozona, its posterior margin obtusangular, with obtuse apex. Prosternum 
comparatively long, with projecting collar-like anterior margin and deep transverse furrow in 
middle ; its posterior half with longitudinal, irregular, ridge-like convexity and, in the middle 
of it, with deep longitudinal sulcus. Prosternum large. Meso-metasternum longitudinally 
concave ; mesosternal interspace slightly longer than its width. Elytra and wings fully 
developed, well exceeding end of abdomen ; anterior margin of elytron almost straight, apex 
obliquely truncate ; membrane thin, transparent, reticulation sparse ; stridulatory veinlets 
strongly reduced. Wing of normal shape. Tympanum present. All femora compressed ; 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



283 



hind femur slender, not reaching end of abdomen. Hind tibia extremely hairy, external apical 
spine present. Tarsi of normal length. Ovipositor with strongly curved acute valves ; upper 
external margin of upper valve irregularly toothed. 
Male unknown. 

Type species : PauUanacris hirsuta sp, n. 

The new genus is of unusual appearance, with extremely prognathous head, 
elongated prosternum, peculiar form of elytron and exceptional hairiness of body. 
These characters isolate it from all known genera. However, it probably belongs 
to the same group as Hemiacris, Phcdinus and Clonacris which are arboreal in habits. 

The genus is named after Dr. R. Paulian whose exploration and study of Mada- 
gascar contributed very much to our knowledge of this geographical region. 



PauUanacris hirsuta sp. n. 

(Text-fig. 5) 

$. Type. General coloration light grey, with small blackish and brownish dots scattered 
all over pronotum and elytra. Hairs long, whitish. Prosternum in middle, mesosternum above 
furcal suture and metasternum in region of metasternal interspace with black spots. Internal 
and lower sides of hind femur bluish black, internal side with whitish pre-apical and incomplete 
whitish medial fascia. Internal side of hind tibia dark purple. 

Length of body 43 ; pronotum 57 ; elytron 40 ; hind femur 20-5 mm. 

Madagascar SW,, Sakaraha : Lambomakandro, 500 m., vii.1957, i $ type (Andria 
Robinson), in Paris Museum. 




Fig. 5. PauUanacris hirsuta sp. n. i, female, type. 2, sternum, c, collar ; F, transverse 
and longitudinal furrows of prosternum ; pr, presternum ; ms, mesosternal interspace ; 
MT, metasternal interspace. 3, end of abdomen, lateral view, 



284 V. M. DIRSH 

HYSIELLA I. Bolivar, 1906 

Hysia St&l, 1875 : 30. 

Hy Stella I. Bolivar, 1906 : 395. 

Of medium size. Body slender, narrowly cylindrical. Integument finely rugose. Antenna 
filiform, longer than head and pronotum together. Head conical, globular above ; fastigium 
of vertex angular, sloping forwards, separated from vertex by depression ; frons strongly oblique, 
straight ; frontal ridge low, sulcate on whole length, with obtuse carinulae ; interocular distance 
slightly wider than basal antennal segment. Pronotum cylindrical ; carinae absent ; four 
wide sulci crossing dorsum ; metazona one-third the length of prozona, its posterior margin 
slightly incurved. Prosternal process short, antero-posteriorly compressed, widening towards 
apex, which is slightly concave in middle. Mesosternal interspace longer than its width, with 
incurved sides. Elytra, wings and tympanum absent. Hind femur slender, reaching end of 
abdomen. External apical spine of hind tibia present. Tarsus elongated. Arolium large. 
Last abdominal tergite of male with pair of acute projections. Supra-anal plate angular, with 
strong transverse furrow and pair of lateral projections. Cercus straight, acutely conical. 
Subgenital plate short, conical. Female subgenital plate obtusangular. Ovipositor robust, 
with short valves, slightly curved at apices. 

Type species : Hysia nigricornis Stal, 1875. 

This genus, originally described as Hysia Stal, 1875 {nofn. preoc), belongs to the 
Tarbaleus group with which it shares the shape of head, pronotum and general 
appearance. However, its relationships cannot be finally established until the male 
genitalia have been examined. 



Hysiella nigricornis (Stal, 1875) 
(Text-fig. 6) 

Hysia nigricornis St&l, 1875 : 30. 

Hysiella nigricornis (St&l, 1875) ; I. Bolivar, 1906 : 395. 

(J. General coloration brownish olive-green. Antenna black, with ochraceous apex. Along 
head, lateral lobe of pronotum, to base of hind femur a narrow black stripe, and below it a wide 
yellowish stripe. External side of hind femur yellowish along upper part, with black stripe, 
serrated at lower margin ; internal and lower side orange-red ; internal side of hind knee black ; 
hind tibia dark olive-green, lower side almost black ; tarsus olive-green. 

?. As the male, but larger. 

Length of body ^ 19, $ 27-5-36-5 ; pronotum (^ 3-4, $ 5"4-6 ; hind femur ^ 10, 9 I5'2- 
17-6 mm. 

Madagascar Est : Ambila-Lemaitso, i 9 (R- Paulian) ; Mananara, Antanambe, 
5 m., iv.1958, I 9 (P. Soga, E. Raharizonina) ; foret au Nord d'Anosibe, i.1951, 
I c^ (R. Pauhan). 

ONETES Rehn, 1944 

Of medium size. Integument finely rugose. Head slightly inflated. Antenna filiform, 
shorter than head and pronotum together. Fastigium of vertex short, with obtuse almost 
truncate apex, roundly merging with frons ; face short and wide ; frons almost vertical, slightly 
convex ; frontal ridge low. Eyes large, interocular distance wide. Pronotum subcylindrical, 
sometimes slightly constricted, with rounded posterior margin ; carinae absent ; three sulci 
crossing dorsum. Prosternal process low, almost square, with flattened apical surface ; meso- 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 285 




Fig. 6. Hysiella nigricornis {Stkl, 1875). i, male, type, lateral view. 2, the same, 
from above. 3, meso- and metasternum. 4, end of abdomen, from above. 5, the 
same, lateral view. 



286 V. M. DIRSH 

sternal interspace constricted in middle and widened in front. Elytra and wings fully developed 
or slightly shortened ; stridulatory veinlets in radial area of elytron reduced ; wing compara- 
tively wide. Hind femur compressed. External apical spine of hind tibia present. Tarsi of 
normal length or slightly elongated. Ovipositor moderately short with valves curved at apices. 

Type species : Onetes sanguinolentus Rehn, 1944. 

The three known species of the genus have all been described from females. The 
male of a species which is tentatively put into the genus Onetes is described below, 
but since I am not sure about its generic status, the male characters are not included 
in the generic diagnosis but only in the specific description. 

Onetes scudderi (Finot, 1903) 
(Text-fig. 7) 

Euthymia scudderi Finot, 1903 : 630. 

Onetes scudderi (Finot, 1903) Rehn 1944 : 51. 

9. Of medium size. Integument finely rugose and slightly hairy. Antenna thin filiform, 
shorter than head and pronotum together. Head slightly prognathous, globular above ; 
fastigium of vertex wide, sloping forwards, roundly merging with frontal ridge ; frons short, 
convex ; frontal ridge wide, low, with flat, rugose surface, widening between antennae. Ocelli 
large. Pronotum cylindrical, without carinae, crossed by three sulci ; metazona shorter than 
prozona, its posterior margin rounded. Prosternal process low, pyramidal, with flattened and 
concave apical surface, in front view slightly bilobate. Meso-metasternum longitudinally 
concave ; mesosternal interspace elongate, narrow, constricted in middle. Elytra and wings 
fully developed, exceeding end of abdomen ; row of stridulatory veinlets of radial area of 
elytron short, containing seven to nine veinlets ; wing slightly widened, with external margin 
strongly excurved. Middle femur compressed. Hind femur strongly compressed ; hind tibia 
hairy, with external apical spine present ; hind tarsus elongated. Arolium large. Female 
subgenital plate with trilobate apex ; ovipositor short, robust, with obtuse, apically slightly 
curved valves. General coloration pale brownish ; elytra without pattern ; wings infumate ; 
hind femur on external side with few irregularly placed dark brown dots ; internal side red 
with two blackish fasciae and blackish internal side of knee ; hind tibia red. 

Length of body 40 ; pronotum 7-6 ; elytron 32 ; hind femur 21 mm. 

Only female type known, from Maroancetra, Antougi Bay. 



Onetes sanguinolentus Rehn, 1944 

This species is known to me from description only. It differs from 0. scudderi 
mainly by slightly constricted pronotum and by spotted elytra. 
Only female type known. 

Length of body 36-5 ; pronotum 6-6 ; elytron 20-2 ; hind femur 16 mm. 
Type locality " Madagascar ". 

Onetes kraussi (Finot, 1903) 

Euthymia kraussi Finot, 1903 : 632. 

Onetes kraussi (Finot, 1903) Rehn, 1944 : 51. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



287 



This species is known to me from description only. It differs from 0. scudderi 
by smaller size of body ; by elytra hardly reaching end of abdomen and by the 
presence of the spots on elytra. 

According to descriptions it is rather similar to 0. sanguinolentus Rehn. 

Only female type known. Length of body 29 ; pronotum 6-4 ; elytron 18 ; 
hind femur 15-5 mm. 

Type locality " Madagascar ". 




Fig. 7. Oneles scudderi (Finot, 1903). Female, type. 



288 V. M. DIRSH 

Onetes fasciatus sp. n. 

(Text-fig. 8) 

(J. Type. Of medium size. Integument rugose and hairy. Antenna filiform, much longer 
than head and pronotum together. Head slightly inflated, globular above ; fastigium of vertex 
strongly sloping forwards, widening towards apex, shallowly depressed in middle and roundly 
merging with frontal ridge ; carinula of vertex present ; frons short, vertical ; frontal ridge 
low, strongly widened between antennae and constricted below ocellus ; concave in upper part ; 
ocelli large. Pronotum cylindrical, dorsum crossed by three deep sulci ; carinae absent ; 
metazona shorter than prozona, its posterior margin obtusangular, almost rounded. Prosternal 
process very low with flat, trapezoidal apical surface. Meso-metasternum longitudinally 
concave ; mesosternal interspace wide, constricted in middle. Elytra and wings shortened, not 
reaching eighth abdominal tergite ; anterior margin of elytron regularly excurved, apex rounded; 
stridulatory veinlets of radial area strongly reduced ; wing widened, with strongly excurved 
external margin. Middle femur compressed. Hind femur compressed, almost reaching end of 
abdomen ; hind tibia hairy ; external apical spine present ; hind tarsus elongated ; arolium 
large. Male supra-anal plate elongate angular, with slightly attenuate apex. Cercus narrow, 
conical, straight. Subgenital plate short, conical, with slightly attenuate apex. Epiphallus 
with short, angular ancorae and wide lobiform lophi with angular apices. Penis strongly pro- 
truding vertically from upper part of subgenital plate ; basal and apical valves of penis not 
connected ; apical valves of penis strongly elongated. Valves of cingulum strongly shortened ; 
sheath of penis short ; arch of cingulum robust. 

General coloration pale brownish, with yellowish irregular spots. Membrane of elytron 
olive-green ; venation pale ochraceous ; wing infumate. Both sides of hind femur greenish 
black, with three pale ochraceous transverse fasciae ; hind tibia brownish. 

Length of body 27-5 ; pronotum 4-2 ; elytron 15-7 ; hind femur 14-5 mm. 

Ampijoroa, 170 m. Foret I'Ankarafantsika, i.1957, i c^ type, (R.E.), in Paris 
Museum. 

Group EUTHYMIAE 

The group name Euthymiae was used first by Brunner v. Wattenwyl in 1893, to 
include Euthymia Stal, 1875 ; Hysia Stal, 1875 ; Lentula Stal, 1878 and, with 
question mark, Gergis Stal, 1875. In 1929 discussing the group Ramme placed in 
it 12 genera and suggested the possibility of including five more. Uvarov (1943) 
placed eight genera in the group. All these authors were concerned onl}^ with 
Old World genera. In 1938 Rehn revised Neotropical Euthymiae, and in 1944 the 
Old World Euthymiae, and included in this group a large number of genera which 
by their peculiar appearance differed from the other genera of the Catantopinae, 
in which they were formerly included. As a result the group became a highly 
artificial, heterogeneous assemblage. 

Recently, according to the structure of the phallic complex and other characters, 
a number of genera were removed to the family Lentulidae and to the subfamily 
Hemiacridinae of Acrididae (Dirsh, 1956). The remaining genera are still formally 
in the group Euthymiae, confusing the interrelations of Euthymia and allied genera 
with other Acrididae. 

According to the structure of the penis, the basal and apical valves of which are 
near but disconnected, Euthymia and allied genera belong in the subfamily Hemia- 
cridinae. The second external character of the subfamily, namely the presence of 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



289 




1 Gpe 
Ejd Ejs Sps 



Qar Sps 



Fig. 8. Onetes fasciatus sp. n. i, male, type. 2, head and pronotum from above. 3, 
face. 4, sternum. 5, end of abdomen, from above. 6, the same, lateral view. 

7, phallic complex, with most part of ectophallic membrane and epiphallus removed. 

8, the same, with part of cingulum removed. 9, epiphallus. 



ENTOM. 12, 6. 



26 



290 V. M. DIRSH 

the stridulatory veinlets in the radial area of the elytron, however, is not so clear. 
In some species of the genus Euthymia (e.g. E. fasciata) these veinlets are reduced to 
such an extent that they are indistinguishable from ordinary veinlets of the reticula- 
tion. In E. holivari, however, the stridulatory veinlets in males are distinct. 

It is proposed here that the group Euthymiae be restricted to the following genera : 
Euthymia Stal, 1875 ; Lopheuthymia Uvarov, 1943 and Proeuthymia Rehn, 1944, all 
of them from Madagascar. Other genera formerly referred to the group need further 
study before their position can be determined. 

The group Euthymiae can be characterized by the structure of the phallic complex, 
with basal and apical valves very near but disconnected, large sheath covering 
whole apical valves of penis except the tips ; well developed apodemes of cingulum, 
and strong, bridge-shaped epiphallus, with short, wide ancorae and large lobiform 
lophi. Amongst the external characters may be mentioned inflated head, compressed 
hind femur and elongated hind tarsus. 

EUTHYMIA Stal, 1875 

Of medium size. Antenna filiform, longer than head and pronotum together. Head sub- 
globular, slightly inflated. Fastigium of vertex short, wide, with trapezoidal apex, fiat, sloping 
forwards and roundly merging with frons, which is straight and slightly oblique or vertical ; 
frontal ridge wide, low, fiat. Pronotum cylindrical, without carinae ; three narrow sulci 
crossing dorsum ; metazona shorter than prozona, its posterior margin obtusangular. Pros- 
ternal process low, obtusely conical ; mesosternal interspace longer than its width, constricted 
in middle. Elytra and wings fully developed or slightly shortened, not reaching end of abdomen ; 
in males with excurved anterior margin, in females with widened and protruding costal area in 
basal half, stridulatory veinlets of radial area of elytron reduced, sometimes almost indis- 
tinguishable from ordinary ones. Wings wide, with strongly excurved posterior margin. 
Hind femur compressed. External apical spine of hind tibia present. Hind tarsus elongated. 
Arohum large. Male supra-anal plate elongate-angular. Cercus narrow, conical, straight or 
curved. Male subgenital plate short, subconical or acutely conical. Basal and apical valves 
of penis very close, but disconnected ; sheath of penis large ; arch of cingulum present ; apodeme 
strong ; epiphallus with narrow bridge, short wide ancorae and wide, lobiform lophi. Ovi- 
positor with moderately short valves, curved at apices. 

Type species : Euthymia melanocerca Stal, 1875. 

Key to Species 

1 (4) Female elytron with regularly excurved basal half. Male subgenital plate 

obtusely conical ; supra-anal plate with regularly angular apex. Lower margin 
of the lower lobe of hind knee roundly excurved. 

2 (3) Fastigium of vertex protruding in front of eyes ; head relatively narrower and 

less inflated. Basal half of anterior margin of female elytron more excurved. 

Male supra-anal plate relatively wider (Text-figs. 9, 1-9) . . . fasciata 

3 (2) Fastigium of vertex scarcely protruding in front of eyes ; head relatively wider 

and more inflated. Basal half of anterior margin of female elytron less 
excurved. Male supra-anal plate relatively narrower (Text-figs. 9, 10-12) 

polychroma 

4 (i) Basal half of female elytron step-like, separated from apical half . Male subgenital 

plate acutely conical ; supra-anal plate with attenuate apex. Lower margin 

of the lower lobe of hind knee with tooth-like projection (Text-figs, g, 13-15) bolivari 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 291 

Euthymia fasciata (Walker, 1870) 
(Text-fig. 9) 

Oxy a fasciata Walker, 1870 : 646. 

Euthymia fasciata (Walker, 1870) Kirby, 1910 : 384. 

Euthymia melanocerca Stal, 1875 : 29 ; Uvarov, 1943 : 387. 

(J. Of medium size, robust. Integument dotted, hairy. Fastigium of vertex protruding in 
front of eyes ; head moderately inflated. Elytra reaching base of supra-anal plate, with 
regularly excurved anterior margin. Lower margin of lower external lobe of hind knee roundly 
excurved ; hind tarsus about half as long as tibia. Supra-anal plate elongate-angular, with 
obtuse apex and longitudinal convexity in middle. Cercus straight, narrow, acutely conical. 
General coloration olive-green ; antenna blackish ; lateral lobe of pronotum with indefinite 
pinkish spot ; base of elytron with black spot ; hind wing blackish-infumate ; hind femur on 
all sides orange-red, but external side slightly olive-greenish ; hind knee black ; hind tibia 
blackish olive-green ; abdomen with alternating black and ochraceous transverse stripes. 

$. As the male, but larger and more robust. Basal half of external margin of elytron 
strongly, regularly excurved. 

Length of body ^ 22-24, ? 33-38 ; pronotum ^ 4-8-5, $ 7-5-8 ; elytron ^ 14-15, $ 17-21 ; 
hind femur ^ 11 -6-1 2, $ 16-17-5 mm. 

Madagascar Est : Manambato sur I'Anove, i $ ; He Sainte Marie, Ambohidema, 
V.1959, 6 $, I (^ ; Ambatoroa, v. 1909, i $ (E. Razafimandimby) ; Tamatave 
Ranomafana, 3 ? ; Ambila-Lemaitso, 3 $, i (^ ; Rogez, vi.1957, i $ ; Midongy du 
Sud, mont Papango, 1,200 m., iii.1959, i $ (A. Robinson). 

Madagascar Quest : foret de I'Ankarafantsika, 120 m. xii. 1959, 2 $ (E. Rahari- 
zonina). 

Euthymia polychroma Brancsik, 1893 

(Text-fig. 9) 

Euthymia polychroma Brancsik, 1893 • i90- 
Euthymia viridescens Sjostedt, 191 8 : 12, syn. n. 

(J. Differs from E. fasciata (Walk.) by the following characters : 

Size of body smaller. Fastigium of vertex scarcely protruding in front of eyes ; head rela- 
tively wider and more inflated. Anterior margin of elytron slightly, regularly excurved. 
Supra-anal plate narrower, with less obtuse apex. General coloration dark olive-green or dark 
ochraceous ; posterior margin of pronotum, near the basal black spot of elytron, with black 
edge ; internal side of hind femur orange, external side purplish (in ochraceous coloured speci- 
mens external side ochraceous). 

$. As the male, but larger. Anterior margin of elytron, in basal half, regularly excurved. 

Length of body c? 21-22, § 31-33 ; pronotum (J 5-3, $ 7-8 ; elytron ^ I3-I3-5, ? 17^20 ; 
hind femur ^J 12, $ 15-5-18 mm. 

Madagascar Sambirano : Nosy Be, r^erve de Lokobe, 4.ix.i947, i ^J, 2 $ (R. 
Paulian) . 

Madagascar Est : Tamatave, Amboditafana, vi.1958, 2 ^ (E. Razafimandimby). 

Euthymia polychroma Brancsik was synonymized by Uvarov, 1943 with E. fasciata 
(Walker) . However, a comparison of a series of specimens with the description and 
figure by Brancsik showed that E. polychroma should be regarded as a valid species, 
which differs from E. fasciata by the characters indicated in the key. Since the 



2g2 



V. M. DIRSH 




^ 



w 





13 




THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 293 

type of E. polychroma is lost, I designate as neotype the female from Lokobe, Mada- 
gascar, which corresponds best with Brancsik's description and figure. The neotype 
is in the Paris Museum. 

The species varies in general coloration from dark olive-green to ochraceous. 

The type of E. viridescens Sjostedt was examined and proved to be conspecific 
with E. polychroma Brancsik. 

Euthymia bolivari Finot, 1903 
(Text-fig. 9) 

(J. Of medium size. Integument finely dotted and moderately hairy. Head moderately 
inflated ; fastigium of vertex well protruding in front of eyes. Posterior margin of pronotum 
obtusangular, almost rounded. Elytron reaching or exceeding end of abdomen, its anterior 
margin slightly and regularly excurved ; stridulatory veinlets mostly distinct. Lower external 
margin of lower lobe of hind knee with tooth-like projection. Supra-anal plate elongate- 
angular, with attenuated apex and longitudinal convexity in middle. Cercus narrow, incurved 
and downcurved, with acute apex. Subgenital plate acutely conical. General coloration from 
light olive-green to ochraceous ; antenna black ; base of elytron with small black spot ; hind 
wing blackish infumate ; hind femur light orange ; hind knee black ; hind tibia blackish, 
sometimes with orange sub-basal ring ; abdomen with alternating black and ochraceous-orange 
transverse stripes. 

$. As the male, but larger. Basal half of anterior margin of elytron strongly projecting 
and step-like, separated from apical half. Ovipositor slender, with slightly curved valves. 

Length of body ^ 21-24, ? 27-35 ; pronotum <J 4-4*6, ? 5-8-6 ; elytron ^ 17-3-20, $ 23- 
24-5 ; hind femur $ 11-12-7, ? 14-16 mm. 

Madagascar Quest : for^t de I'Ankarafantsika, Ampijoroa, 2 ^. 
Madagascar Est : reserve de Zakamena, i (^, i $ ; Perinet, i $, i cJ. 

PROEUTHYMIA Rehn, 1944, stat. n. 

Euthymia {Proeuthymia) Rehn, 1944 : 31. 

Proeuthymia, which was described by Rehn as a subgenus of Euthymia, is here 
considered as a genus. It differs from Euthymia by the following characters : 

Male supra-anal plate wide, its sides obtusangularly incurved, apex obtusangular with lateral 
angles slightly attenuate ; a middle transverse suture divides it into a basal part with median 
longitudinal concavity, and an apical half, with a pair of lateral tubercles. Lophi of epiphallus 
narrow, acutely angular. Gonopore process of phallus large, wide and very robust. 

Type species : Euthymia saussurei Finot, 1903. 



Fig. 9. 1-9, Euthymia fasciata (Walker, 1870). i, male. 2, phallic complex, with 
ectophallic membrane and epiphaUus removed. 3, the same, with most part of 
cingulum and sheath of penis removed. 4, epiphallus, lophi in horizontal position. 
5, the same, lophi in vertical position. 6, right male elytron. 7, the same, female 
elytron. 8, hind knee, external side. 9, male supra-anal plate. 10-12, Euthymia 
polychroma Brancsik, 1893. 10, female right elytron. 11, hind knee, external side. 
12, male supra-anal plate. 13-15, Euthymia bolivari Finot, 1903. 13, right female 
elytron. 14, hind knee, external side. 15, male supra-anal plate, 



294 



V. M. DIRSH 



Proeuthymia saussurei (Finot, 1903), stat. n. 
(Text-fig. 10) 

Euthymia saussurei Finot, 1903 : 625. 

Euthymia [Proeuthymia) saussurei (Finot, 1903) ; Rehn, 1944 : 32. 

(J. Of medium size. Integument almost smooth. Antenna filiform, longer than head and 
pronotum together. Head slightly inflated ; fastigium of vertex trapezoidal, well protruding 
in front of eyes, with truncate apex ; frons oblique ; frontal ridge low, wide, above ocellus 
flat, below it constricted and concave. Pronotum subcylindrical, without carinae ; three deep, 
wide sulci crossing dorsum ; metazona slightly shorter than prozona, its posterior margin 
obtusangular. Prosternal process low, subconical, with obtuse apex. Mesosternal interspace 
longer than its width, narrowing in front. Elytra slightly exceeding end of abdomen, anterior 
margin in basal half slightly excurved ; stridulatory veinlets strongly reduced, indistinguishable 
from ordinary veinlets. Hind wing relatively narrow. Hind femur moderately compressed ; 
lower margin of lower external lobe of hind knee with tooth-like projection. Hind tarsus less 
than half the length of tibia. Arolium large. Supra-anal plate almost as wide as its length. 
Cercus compressed, narrow, angular, with acute apex. Subgenital plate short, obtusely sub- 
conical. General coloration olive-green ; antenna black ; transverse sulci of pronotum black ; 
hind wing pink, with infumate external margin ; hind femur light green on all sides ; hind knee 
black ; tibia with black base, light green in middle and blackish at apex ; spines black ; hind 
tarsus red. 

$. As the male, but larger. Anterior margin of elytron, in basal half, regularly excurved. 
Ovipositor short, with slender valves, slightly curved at apices. 

Length of body ^ 19-22, $ 23-27 ; pronotum ^ 4-47, ? 6-6-5 '> elytron ^ i2-5-i6-6, $ 19-21 ; 
hind femur (J 11-4-12, $ 13-5-16 mm. 

Madagascar Sud Quest : Sakaraha, Lambomakandro, xii.1956, i cJ, 3 $. 
Madagascar Quest : for6t de I'Ankarafantsika, 20.1.1950, i (^. 



LOPHEUTHYMIA Uvarov, 1943 

Of medium or large size. Integument rugose and hairy. Antenna filiform, longer than 
head and pronotum together. Head slightly prognathous ; fastigium of vertex strongly sloping 
forwards, roundly merging with frons and not protruding in front of eyes ; frons depressed. 





Fig. 10. Proeuthymia saussurei (Finot, 1903). i, female right elytron, 
external side. 3, male supra-anal plate. 4, epiphallus. 



2, hind knee, 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 295 




Fig. II. Lopheuthymia brunneri (Finot, 1903). i, female. 2, male. 3, phallic 
complex, with most part of ectophallic membrane and epiphallus removed. 4, the 
same, with most of cingulum removed. 5, epiphallus, 



296 V. M. DIRSH 

frontal ridge low, flat, widening from base of antennae upwards. Pronotum tectiform, in 
prozona with two high tooth- or tubercle-like projections ; three deep sulci crossing dorsum ; 
metazona shorter than prozona, its posterior margin acutangular. Prosternal process low, 
almost square, with slightly concave apical surface. Mesosternal interspace slightly wider than 
its length and slightly constricted in middle. Elytra and wings not reaching end of abdomen ; 
stridulatory veinlets of radial area of elytron reduced. Hind wing widened, with expanded 
posterior margin. Hind femur compressed ; hind tarsus elongated. Arolium large. Last 
abdominal tergite of male with pair of angular projections ; supra-anal plate angular, with 
trilobate apex, lateral lobes small, and with two lateral tubercles behind median transverse 
suture ; cercus narrow, incurved, with acute apex ; subgenital plate short, subconical. Basal 
and apical valves of penis very close, but clearly divided ; sheath of penis large. Epiphallus 
bridge-shaped, with short, wide ancorae and large lobiform lophi. Female subgenital plate 
deeply trilobate. Ovipositor moderately short, with curved valves. 

Type species : Euthymia brunneri Finot, 1903. 

Lopheuthymia brunneri (Finot, 1903) 
(Text-fig. 11) 

Euthymia brunneri Finot, 1903 : 633. 

Lopheuthymia brunneri (Finot, 1903) Uvarov, 1943 : 388. 

(J. Of medium size. Head slightly prognathous. General coloration brown ; elytra and 
pronotum dirty ochraceous, with small dark brown spots and stripes ; elytra sometimes with 
few small reddish spots ; hind wing blackish-infumate ; external side of hind femur ochraceous, 
apical half with blackish, irregular transverse fascia ; internal side black, with pre-apical 
ochraceous fascia ; hind knee black ; hind tibia reddish, with basal part and part of lower side 
blackened ; hind tarsus red, except blackish tip. 

$. As the male, but larger, with more prognathous head and relatively shorter elytra. 

Length of body ^ 22-25, ? 30-35 ; pronotum (^ 5-5-6, $ 11-12 ; elytron ^ 14-17-5, $ 16-18 ; 
hind femur ,$ 12-15-5, $ 17-18 mm. 

Madagascar Est : Maroantsetra, Fampanambo, i $, i c^ (J. Vadon) ; Ambodita- 
volo, I $ (Capuron) ; Perinet, i (^. 

GALIDEUS Finot, 1908 

Of medium size ; body very strongly elongate and narrow. Integument finely rugose and 
sparsely hairy. Antenna filiform, in male longer, in female shorter than head and pronotum 
together. Head strongly elongate, acutely conical ; fastigium of vertex narrow, acutely 
angular, much longer than longest diameter of eye ; above convex ; at base, in front of eyes, 
with transverse depression and strong transverse furrow ; in profile, with acutely pointed apex ; 
frons slightly incurved ; frontal ridge narrow, low, with low, obtuse parallel lateral carinulae 
and shallow depression between them ; at apex lamelliformly compressed. Interocular distance 
about as wide as basal antennal segment. Ocelli vestigial. Pronotum cylindrical, with four 
shallow sulci, first and second sometimes indistinct ; metazona about one-fourth of length of 
prozona. Prosternal process widening towards apex, which is flattened, widened and slightly 
concave. Mesosternal interspace small, inverse triangular, closed. Elytra and wings shortened, 
slightly exceeding middle of abdomen ; elytron with rounded apex and strongly reduced stridu- 
latory veinlets ; wing wide, with strongly excurved external margin. Hind femur narrow, 
slender, not reaching end of abdomen ; lower external lobe of hind knee angular ; external 
apical spine of hind tibia present ; tarsus slender, elongated, about half as long as tibia. Arolium 
large. Male supra-anal plate elongate angular, with attenuate apex ; cercus long, straight, 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



297 




Fig. 12. 1-7, Galideus mocquerysi Finot, 1908. i, male, type. 2, head, from above. 
3, the same, lateral view. 4, end of male abdomen, from above. 5, the same, lateral 
view. 6, end of female abdomen, lateral view. 7, epiphallus. 8-12, Galideus elegans 
sp. n. 8, head from above. 9, the same, lateral view. 10, phallic complex, with 
most part of ectophallic membrane and epiphallus removed. 11, the same, with most 
part of cingulum removed, 12, epiphallus. 



298 V. M. DIRSH 

almost cylindrical, with subacute apex ; subgenital plate short, acutely conical. Phallic organ 
rather slender. Basal and apical valves of penis connected by thin but strong flexure. Valves 
of cingulum large ; arch of cingulum wide and strong. Epiphallus bridge-shaped, without 
ancorae and with large, hook-shaped lophi. Ovipositor long, narrow, with straight valves. 

Type species : Galideus mocquerysi Finot, 1908. 

Galideus mocquerysi Finot, 1908 
(Text-fig. 12) 

(J. Antenna filiform on whole length. Fastigium of vertex slightly shorter than twice longest 
diameter of eye, with strongly acute, almost pointed apex. General coloration yellowish ; 
antenna black ; on sides, beginning at apex of fastigium of vertex, along whole head, pro- 
notum and elytra there is wide, blackish stripe. Hind femur yellowish ; hind knee reddish ; 
hind tibia blackish purple ; tarsus reddish. 

?. As the male, but larger. 

Length of body cj 31*5-34 ; ? 49 mm. 

The cJ type and $ paratype were studied. 
Madagasc< — - - 
ic? (P. Soga) 



The ^ type and $ paratype were studied. 

Madagascar Est : Sambava, reserve naturelle XII, Marojejy, Andrakato, ii.1959, 



Galideus elegans sp. n. 

(Text-fig. 12) 

(J. Type. Differs from G. mocquerysi Finot by slightly smaller size ; compressed basal part 
of antenna by fastigium of vertex being more than twice as long as longest diameter of eye, 
with apex parabolic. General coloration yellowish orange. Antenna light brownish ; lateral, 
longitudinal stripes of body brownish ; hind knee yellowish orange ; hind tibia bluish-black ; 
tarsus yellowish orange. 

Female unknown. 

Length of body 31 ; pronotum 4 ; elytron 11-3 ; hind femur 14 mm. 

Madagascar Est : Ambodirafia, 7.ii.i934, i (^ (type), (A. Seyrig) in Paris Museum. 
Madagascar Centre: Ambohimitombo Forest, 26. i. 1895, (^ paratype in British 
Museum (Natural History). 

XENIPPOIDES Chopard. 1952 

Small ; body elongate cylindrical, stick-like. Antenna as long as head and pronotum to- 
gether, ensiform, basal half triangular in cross-section. Head elongated, acutely conical ; 
fastigium of vertex narrow angular, strongly protruding in front of eyes, slightly convex, 
separated from vertex by fine furrow, with subacute apex ; frons strongly oblique and incurved ; 
frontal ridge narrow, strongly compressed above base of antennae, with narrow sulcus and 
obtuse lateral carinulae. Eyes elongate oval. Ocelli absent. Pronotum short, cylindrical ; 
dorsum crossed by three fine sulci ; metazona one-quarter the length of prozona, its posterior 
margin shallowly incurved. Prosternal process slightly spathulate, antero-posteriorly com- 
pressed, with slightly bilobate and slightly inflated apex. Mesosternal interspace inverse tri- 
angular, closed. In male, vestigial elytra present, wings and tympanum absent. Anterior 
and middle legs short. Hind femur slender, not reaching end of abdomen. External apical 
spine of hind tibia present. Hind tarsus slightly elongated. Arolium moderately large. Last 
abdominal tergite of male with small, widely rounded projections. Supra-anal plate elongate- 
angular. Cercus long, narrow, conical. Subgenital plate short, conical. Phallic complex 
slender. Sheath of penis absent ; rami large ; basal and apical valves of penis connected by 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



299 



flexure ; apical valves of penis large, lobiform ; valves of cingulum small ; arch of cingulum 
present ; gonopore processes long. Epiphallus bridge-shaped, with divided bridge, small 
obtuse ancorae and large hook-like, incurved lophi. Ovipositor long, slender, with valves 
straight, serrated on external margins and obtuse at apices. Supra-anal plate elongate angular. 
Cerci straight, narrow, stick-like, with obtuse apices. Subgenital plate with obtusely angular 
apex. 

This peculiar genus, according to the shape of head, ovipositor and cercus, is 
probably related to Galideus, from which, however, it differs very strongly by the 
wings being vestigial in the male and absent in the female, by less elongated and 
less acute head, and by shorter tarsi and other characters. 

Type species : Xenippoides elongatus Chopard, 1952. 






Fig. 13. Xenippoides elongatus Chopard, 1952. i, male, type. 2, female, para type. 
3, head from above. 4, end of male abdomen, from above. 5, end of female abdomen, 
lateral view. 6, phallic complex, with ectophallic membrane and epiphallus removed. 
7, the same, with most part of cingulum removed. 8, epiphallus. 



300 V. M. DIRSH 

Xenippoides elongatus Chopard, 1952 
(Text-fig. 13) 

(J. Integument finely rugose and sparsely hairy. Head smooth ; fastigium of vertex slightly 
upcurved and slightly dotted above. Lower margin of lateral lobes of pronotum straight. 
Lower lobe of hind knee angular, with obtuse apex. General coloration ochraceous, with 
indefinite darker stripes along sides of head and part of pronotum. At posterior margin of 
pronotum, on pleura, a small, elongate black spot. Hind tibia reddish. 

?. As the male, but larger. 

Length of body, (J 19-5, ? 23-5 ; pronotum ^ 2-5, $ 3 ; hind femur (J 9-2, $ 9-8 mm. 

Madagascar Sambirano : Mont, Tsaratanana, foret de mousses, i (^ type, i $ 
paratype in Paris Museum. 

Group GERGISINI 

This group contains the following genera : Gergis Stal, 1875 ; Malagasacris Rehn, 
1944 ; Pachyceracris gen. n. and Morondavia gen. n. The most remarkable feature 
of the group is the cubital vein of the hind wing connected with the medial (see 
Text-fig. 18). This may occur in the middle or in the more distal part of the cubitus, 
and is normally present only in this group of genera. Occasionally it has been 
observed also in the genera Pareuthymia Willemse, 1930, Phalinus Rehn, 1944 and 
Enoplotettix I. Bolivar, 1913. 

The phallic complex was studied in one species of Gergis, and in Malagasacris 
and Morondavia. In the first two genera it agrees reasonably well with that of the 
subfamily Hemiacridinae (see Text-figs. 14-16). In Morondavia it possesses a 
rather thick flexure between basal and apical valves of penis which does not as a 
rule occur in Hemiacridinae, but the penis sheath is of Hemiacridinae type. 

The characteristic stridulatory veinlets of Hemiacridinae, between radial and 
medial veins of elytron, are clearly distinct only in Malagasacris, while in other 
genera of the group they are greatly reduced or obliterated. 

Since some of the essential characters of the group are rather obscure, I am 
tentatively placing it in the subfamily Hemiacridinae in the hope that further study 
of more material will clarify the question. 



MALAGASACRIS Rehn, 1944 

Small. Integument rugose. Antenna filiform, longer than head and pronotum together. 
Head inflated, globular above ; fastigium of vertex wide, strongly sloping forwards, with 
truncate apex ; frons short, slightly oblique, strongly rugose, with deep foveolae above antennae; 
frontal ridge wide, low, with flat, rugose surface, widest between and narrowed above and 
below antennae. Eyes and ocelli large, strongly convex ; interocular distance about twice as 
wide as basal antennal segments. Pronotum cylindrical ; dorsum crossed by three or four 
deep sulci ; metazona shorter than prozona, its posterior margin obtusangular, almost rounded. 
Epistemum large, with attenuate anterior angle. Prosternal process low, with rounded apex. 
Mesostemal interspace wide, with incurved sides. Elytra and wings slightly shortened, reaching 
or almost reaching end of abdomen ; stridulatory veinlets of radial area of elytron reduced ; 
hind wing with roundly excurved external margin, cubital vein in apical quarter connected with 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



301 



medial vein. Hind femur slender ; external apical spine of hind tibia present ; hind tarsus 
elongated, exceeding half length of tibia. Arolium large. Male supra-anal plate angular. Cercus 
straight, narrow, conical. Subgenital plate short, subconical. Ectophallus membraneous ; 
rami narrow ; sheath of penis covering whole apical valves of penis ; valves of cingulum well 
developed. Endophallus strongly sclerotized ; basal valves of penis narrow, with very large 
gonopore processes ; apical valves of penis long, narrow. Epiphallus bridge-shaped, with 
strong ancorae and short, wide, lobiform lophi. 

Type species : Malagasacris strateia Rehn, 1944. 

In Malagasacris the connection of the cubitus with the media in the hind wing 
occurs in the distal part of the veins and is less clear than in other genera of the 
group. 




Fig. 14. Malagasacris strateia Rehn, 1944. i, male, paratype. 2, male supra-anal 
plate. 3, phallic complex, with ectophallic membrane and epiphallus removed. 4, 
the same, with most part of cingulum removed. 5, epiphallus. 6, male supra-anal 
plate of Malagasacris rugosa sp. n. 



302 V. M. DIRSH 

Rehn (1944), describing Mcdagasacris strateia, designated the male as the type and 
a female as the allotype ; however, it is doubtful whether the female belongs to the 
same genus as the male. 

Malagasacris strateia Rehn, 1944 
(Text-fig. 14) 

(J. Integument rugose and shiny. Fastigium of vertex moderately wide. Dorsum of 
pronotum crossed by four sulci. Prosternal process tubercle-like. Last abdominal tergite 
without projections. Supra-anal plate angular. General coloration brownish. Lateral lobe 
of pronotum with wide, bright yellow stripe along posterior margin. Membrane of elytron 
bluish, with light, brownish venation and reticulation ; hind wing bluish, with infumate external 
margin. Hind femur on both sides bright yellow, on external side with two, on internal side 
with one blackish transverse fasciae ; knee brown ; hind tibia reddish brown ; tarsus reddish. 

Length of body ^ ly^-i'j-z; pronotum 3-3-3-6 ; elytron io-7-ii-2; hind femur 10-2- 
11-2 mm. 

Only the originally described species from Rogez Distr. is known. 

Malagasacris rugosa sp. n. 

(Text-fig. 14) 

cj. Type. Integument strongly rugose, matt. Fastigium of vertex wide. Dorsum of 
pronotum crossed by three sulci. Prosternal process compressed with subacute apex. Last 
abdominal tergite with a pair of acute projections. Supra-anal plate angular with attenuate 
apex. General coloration brownish. Membrane of elytron bluish, with light brownish venation 
and reticulation ; hind wing bluish, with widely infumate external margin. External side of 
hind femur pale ochraceous, with basal margin black and two wide black transverse fasciae, 
knee brown, with yellowish lower lobe ; internal and lower side blue, with two not very clear 
whitish transverse fasciae ; hind tibia blue ; tarsus brownish. 

Length of body ^ i6-6 ; pronotum 3-7 ; elytron 12 ; hind femur 10 mm. 

Madagascar Est : Moramanga, Ankasoka, 1,130 m., i.1959, i ^ type (P. Griveaud). 
In Paris Museum. 

The two species of the genus are easily distinguishable by the following main 
characters : 

1 (2) Prosternal process tubercle-like, with rounded apex. Last abdominal tergite of 

male without projections. Supra-anal plate simple, angular. Internal and 
lower sides of hind femur yellow ; tibia reddish brown .... strateia 

2 (i) Prosternal process compressed, with subacute apex. Last abdominal tergite of 

male with a pair of acute projections. Supra-anal plate angular, with attenu- 
ated apex. Internal and lower sides of hind femur blue ; tibia blue . rugosa 

GERGIS Stal, 1875 

Size medium or small. Integument finely rugose and sparsely hairy. Antenna filiform, 
longer than head and pronotum together. Head inflated, subglobular above ; fastigium of 
vertex sloping forwards, obtusangular, separated from vertex by depression ; interocular 
distance wider than basal antennal segment ; frons oblique, straight, short ; frontal ridge low, 
wide, constricted below ocellus and almost obliterated below it ; eyes large, strongly convex ; 
ocelli large. Pronotum cylindrical, without carinae ; dorsum crossed by three deep sulci ; 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 303 

metazona half of the length of prozona, its posterior margin rounded. Prostemal process 
obtusely conical. Mesosternal interspace widened at apex and strongly constricted in middle ; 
metasternal interspace inverse-triangular, closed. Elytra and wings shortened, reaching last 
abdominal tergite ; anterior margin of elytron slightly excurved, apex rounded, venation 
rough, reticulation rough and sparse ; stridulatory specialization of elytron strongly reduced, 
sometimes indistinguishable. Hind wing widened with strongly excurved external margin ; 
cubital vein connected with medial one. Hind femur slender. Hind tibia slightly expanded 
in apical part ; external apical spine absent. Hind tarsus more than half the length of tibia. 
Arolium large. Last abdominal tergite of male with a pair of large, lobiform posterior projec- 
tions. Supra-anal plate elongate. Cercus long, narrow, acutely conical, slightly incurved. 
Subgenital plate elongate, acutely conical. Phallic organ robust. Basal and apical valves of 
penis disconnected, but very close. Arch of cingulum and valves of cingulum well developed. 
Sheath of penis large, completely covering apical valves and valves of cingulum. Epiphallus 
bridge-shaped, with small, angular, incurved ancorae and small lobiform lophi. 

Type species : Gergis paLlidinervis Stal, 1875. 

Gergis pallidinervis Stal, 1875 
(Text-fig. 15) 

^. Type (redescription). Of medium size. Apex of fastigium of vertex obtuse. Male 
supra-anal plate elongate-angular, with apex acutely angular and without lateral and median 
carinae. Cercus conical, slightly, irregularly curved. Subgenital plate conical, in profile with 
obtuse apex. General coloration olive-brown. Face brownish with few black dots. Antenna 
brownish. Pronotum with small, irregular, blackish spots. Elytra olive-brownish, with main 
veins ochraceous. Hind wings infumate. Hind femur on external side dirty ochraceous ; 
lower and internal side slightly orange-red. Hind knee blackish. Hind tibia on basal half 
slightly orange above, the rest blackish. Tarsi reddish. 

$. As the male, but larger. Ovipositor long and slender, with valves slightly curved at 
apices. Subgenital plate with angular apex. 

Length of body <J 22, $ 31-2 ; pronotum ^ 4-3, $ 5-2 ; elytron (J 13-5, $ 14 ; hind femur 
(J 12, $ 14 mm. 

Madagascar Est: i cJ type. Ambila-Lemaitso, i.1957, i ? (A. R.). 



Gergis minor sp. n. 

(Text-fig. 16) 

(J. Type. Small. Fastigium of vertex with angular apex. Male supra-anal plate elongate, 
with curved sides, with deep median sulcus, strong lateral carinulae and strong transverse suture ; 
apex angular and attenuate. Cercus conical, slightly incurved. Subgenital plate elongate, 
conical in profile, with acute apex. General coloration brownish. Face ochraceous, with two 
transverse, brown fasciae. Antenna brownish. On head and pronotum a pair of irregular 
ochraceous stripes. Pleura ochraceous. Elytra brown, with ochraceous main longitudinal 
veins. Hind wing infumate. Anterior and middle legs ochraceous. Hind femur on external 
side ochraceous, internal and lower side orange-red ; hind knee bluish-black ; hind tibia orange- 
reddish on basal and brown on apical half ; tarsus red. Lower side of body brown. 

? paratype as male, but larger and less brightly coloured. (End of abdomen broken.) 



304 



V. M. DIRSH 



Length of body (J 17-5-18 ; pronotum (J 3-3-2, $ 4-2 ; elytra ^ 8-5-9, 9 12 ; hind femur 
(J 10-10-3, $ 12 mm. 

The new species differs from Gergis pallidinervis in the following characters : 
minor pallidinervis 



Size smaller 

Fastigium of vertex with angular apex 

Male supra-anal plate with attenuate apex 

and strong lateral carinulae 
Male subgenital plate elongate conical in 

profile with acute apex 



Size larger. 

Fastigium of vertex with obtuse apex. 

Male supra-anal plate with angular apex 

and without lateral carinulae. 
Male subgenital plate conical in profile with 

obtuse apex. 



Madagascar Est : Perinet, 4 c^ (including type), i $. Foret Nord d'Anosibe, 
i,i95i, I ^, (R. Paulian). 

Type and paratypes in Paris Museum. Two (^ paratypes in the British Museum 
(Natural History). 




Fig. 15. Gergis pallidinervis St&l, 1875. i, male type. 2, female. 3, sternum. 
4, end of male abdomen, lateral view. 5, the same, from above. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 

PACHYCERACRIS gen. n. 



305 



Of medium size. Integument rugose. Antenna rod-like, thickened in basal part, much 
shorter than head and pronotum together. Head widely conical, slightly inflated, subglobular 
above ; fastigium of vertex horizontal, angular, convex above, with marginal carinulae, 
separated from vertex by depression ; inter-ocular distance wider than basal antennal segment ; 
frons strongly oblique, short, straight ; frontal ridge wide, convex, without lateral carinulae, 
distinct and tubercle-like above ocellus, obliterated below. An obtuse tubercle between base 
of antenna and eye. Pronotum cylindrical ; dorsum crossed by three deep sulci, metazona 
shorter than prozona, its posterior margin rounded ; trace of median carina noticeable in meta- 
zona only ; lateral carinae absent. Prosternal process short, tubercle-like, with wide, rounded 
apex ; prosternum wide ; mesosternal interspace widened at apex and strongly constricted in 
middle. Elytra and wings shortened, reaching eighth abdominal tergite ; transverse veinlets 
of radial area of elytron not specialized ; hind wing widened, with strongly excurved external 
margin ; cubital vein connected with media at distal end. Hind femur moderately slender. 
External apical spine of hind tibia absent. Hind tarsus half length of tibia. Arolium large. 
Ovipositor elongated, slender, with valves slightly curved at apices. (Only female known.) 

Type species : Pachyceracris fusca sp. n. 

The new genus is strikingly different from all known genera of the subfamily. 
A study of a male, which is unknown, may help to establish more definitely the 
position of the genus. Its most remarkable features are strongly shortened antenna, 
form of head, with tubercle-like upper part of frontal ridge, and pecuHar connection 
of cubitus with media in hind wing. 




Bp-4- 



Ejs Gpr Sps Ap 

4 

Fig. 16. Gergis minor sp. n. i, male, type. 2, end of abdomen, from above. 3, 
phallic complex, with ectophallic membrane and epiphallus removed. 4, the same, 
with most of cingulum removed. 5, epiphallus. 

ENTOM. 12, 6. 27 



3o6 



V. M. DIRSH 



Pachyceracris fusca sp. n. 

(Text-fig. 17) 

$. Type. Integument strongly rugose, but shiny. Antenna eigh teen-segmented. Meta- 
sternal interspace irregularly rounded, closed. Anterior margin of elytron slightly excurved, 
apex rounded ; membrane semi-transparent ; venation and reticulation rough. Subgenital 
plate with angular apex and a pair of pre-apical, small, round, low convexities. General 
coloration brownish. Hind femur ochraceous, on external side with indistinct, brownish 
longitudinal stripe ; hind knee brown on both sides ; hind tibia and tarsus red. Hind wing 
infumate. 

Length of body 32 ; pronotum 5-5 ; elytron 16 ; hind femur 13 mm. 

Madagascar Centre : Andranotobaka, 1,400 m., Ambatolampy, iv.1957, i $, type 
(P. Griveaud) in Paris Museum, 

MORONDAVIA gen. n. 

Small. Integument finely rugose. Antenna filiform, shorter than head and pronotum 
together. Head inflated, globular above ; fastigium of vertex sloping and strongly protruding 
forwards, angular, with obtuse apex ; interocular distance slightly narrower than basal antennal 
segment ; eyes large, moderately convex ; frons strongly oblique, very short, about as long as 
length of clypeus and labrum ; frontal ridge wide, fiat, obliterated below ocellus, roundly 
merging with fastigium of vertex. Pronotum subcylindrical, without carinae ; three weak sulci 
crossing dorsum ; metazona much shorter than prozona, its posterior margin widely rounded. 




Fig. 17. Pachyceracris fusca sp. n. i, female, type. 2, antenna. 3, sternum. 4, end 

of abdomen, lateral view. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 307 

Prostemal process short, tubercle-like, with rounded, inflated apex. Meso- and metastemum 
longitudinally concave ; mesosternal interspace widening at apex and strongly constricted in 
basal part. Elytra and wings not reaching end of abdomen ; venation of elytron rough and 
reduced, but in precostal and vannal areas there are rows of regular, parallel, thickened, oblique 
veinlets ; hind wing slightly widened, with strongly excurved posterior margin ; cubital vein 
connected and fused with medial vein. Hind femur moderately slender ; external apical spine 
of hind tibia absent ; tarsus elongated. Arolium large. Last abdominal tergite of male with 
a pair of small projections ; supra-anal plate elongate-angular ; cercus slender, elongated, with 
acute apex ; subgenital plate narrow, subconical. Phallic complex with well-developed apodeme 
of cingulum, with large sheath, completely covering apical valves of penis and large arch of 
cingulum. Basal valves of penis large ; connected with apical valves by thick flexure-like 
connection. Valves of cingulum small, acute. Epiphallus bridge-shaped, with narrow bridge, 
small ancorae and large, lobiform lophi. Female subgenital plate with acutangular apex and 
pair of pre-apical small, round convexities. Ovipositor elongated, with narrow valves, slightly 
curved at apices ; upper margin of upper valve roughly serrated. 

Type species : Morondavia cephalica sp. n. 

This genus differs strongly from other genera of the group by speciahzation of 
the elytra (possibly stridulatory), represented by rows of regular thickened veinlets 
in precostal and vannal areas, and by inflated head with strongly shortened frons. 

Morondavia cephalica sp. n. 

(Text-fig. 18) 

c^. Type. Head strongly inflated ; fastigium of vertex broadly angular. Cercus strongly 
incurved ; supra-anal plate with parabolic apex. General coloration ochraceous-brownish, 
with ochraceous spots on lateral lobes of pronotum and pleura. External side of hind femur, 
in male, ochraceous, with two infumate transverse fasciae ; internal and lower side red with 
blackish pre-apical fascia ; hind tibia on lower side of basal part reddish, the rest of it on all 
sides blackish ; hind tarsus red. 

?. Paratype Larger than male, with slightly less inflated head. General coloration a.s in 
male, but ochraceous pattern on lateral lobes of pronotum more definite. External side of 
hind femur without fasciae ; internal side with only trace of pre-apical fascia. Hind tibia red. 

Length of body cj 20, $ 24 ; pronotum (J 3-5, ? 37 ; elytron ^ 12, ? 11-4, hind femur (J 10, 
$ 10-5 mm. 

I Madagascar Quest : Antsingy N., 63 km. East Maintirano Forest, vii.1949, i c^ 
type (R. Pauhan) ; for^t de I'Ankaraiantsika, Ampijoroa, xii.1951, i ? paratype. 
Type and parat5^e in Paris Museum. 

Morondavia minor sp. n. 

(Text-fig. 18) 

(J. Type. Differs from M. cephalica by smaller size, slightly less inflated head ; narrow 
fastigium of vertex ; straight male cercus, supra-anal plate of male with angular and slightly 
attenuate apex and by tibia red in both sexes. 

Length of body (J 10-12, ? 22 mm. 

Madagascar Quest : Morondava, foret S. de Befasy, i.1956, i c? type, 2 $ (R. 
Paulian) . 

Madagascar Nord : Montague d'Ambre, Joffreville, xii.1947, i J (P. Cachan). 




Fig. i8. 1-4, Morondavia cephalica sp. n. i, male, type. 2, elytron. 3, hind wing. 
4, end of male abdomen, from above. 5-8, Morondavia minor sp. n. 5, end of male 
abdomen, from above. 6, phallic complex, with ectophallic membrane and epiphallus 
removed. 7, the same, with most part of cingulum removed. 8, epiphallus. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 309 

Type and paratype in Paris Museum, ic^, i? paratypes in the British Museum 
(Natural History). 

Subfamily OXYINAE 

OXYA ServiUe, 1831 

Oxya hutneralis (Walker, 1870) 
(Text-fig. 19) 

Heteracris humeralis Walker, 1870 : 662. 

Oxya humeralis (Walker, 1870) Willemse, 1925 : 39. 

This species was described as Heteracris humeralis Walker, 1870 from a single 
female type from Madagascar preserved in the British Museum (Natural History) 
and still in reasonably good condition. 

Willemse in 1925 synon5mriized this species with Oxya hyla ServiUe, 1831 on the 
basis of the structure of the female subgenital plate and the general pattern of 
coloration, which is, in any case, almost the same in all species of the genus. 

Oxya hyla Serville, the type of which is lost and the type locality Senegal, has 
been recorded from almost the whole of tropical Africa, but a preliminary study of 
African material suggests that several different species may be involved. Therefore, 
until Oxya hyla Serville is redefined and a neotype designated, Walker's species 
must be regarded as valid. 

(J. Supra-anal plate trilobate, with lateral lobes short and small, and the apical lobe large, 
angular. Cercus slightly compressed, in lateral view with almost straight lower and oblique 
and slightly excurved upper margin, apical part of which is slightly oblique, apex subacute. 
Subgenital plate short, narrow, subconical. Phallic complex rather slender. Sheath of penis 
absent ; basal and apical valves of penis connected by thin strongly sclerotized flexure ; valves 
of cingulum large ; arch of cingulum low, robust. Epiphallus bridge-shaped, with bridge 
divided ; ancorae small, lophi Ipbiform or tooth-like. General coloration green or greenish, 
with brown lateral stripe running from eyes, through lateral lobes of pronotum and along 
elytron. Hind femur on all sides green or greenish ; hind knee brown or brownish ; hind tibia 
blue. 

?. As male, but larger. Subgenital plate at apex with two acute teeth and a pair of strongly 
toothed, slightly incurved lateral ridges. Ovipositor slender ; upper valves on external side 
slightly serrated ; lower valves on external sides with acute large teeth. Pattern and coloration 
as in male. 

Length of body ^ 21-25-5, $ 26-31 ; pronotum ^ 4-4-5-6, $ 5-6-8 ; elytron ^ 15-23, ? 20-5- 
28 ; hind femur ^ 12-3-14-8 mm. 

This species is highly variable in body size and relative length of elytra, which 
may exceed end of abdomen by about one to one and a half times the length of 
pronotum. 

Madagascar Centre : Tananarive Tsimbazaza, ii.1949, 2 $ (R. Paulian). 

Madagascar Est : Sambava, Marojejy, Ambinanitelo, 500 m., xii.1958, i (J, 2 $ 
(E. Raharizonina) ; foret de Tampolo, vii.1949, 2 (^ (J. Millot) ; Tamatave, Fanan- 
drana, iii.1959, i (^ (E. Razafimandimby) ; station agricole de ITvoloina, xi.1958, 
I ? (J. Ehe) ; foret Nord d'Anosibe, 1.1957, i ? (R. Paulian) ; Brickaville, station 
agricole, i 9. 



3IO 



V. M. DIRSH 




Fig. 19. Oxya humeralis (Walker, 1870). i, male. 2, phallic complex, with ectophallic 
membrane and epiphallus removed. 3, the same, with most part of cingulum removed. 
4, epiphallus. 5, end of female abdomen, lateral view. 6, the same, from below. 7, 
end of male abdomen, from above. 8, male cercus. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 311 

Madagascar Quest: Ambilobe, iv.1957, i (^ (R. Paulian) ; foret de I'Ankara- 
fantsika, Ampijoroa, i $; Antsalova, foret de I'Antsingy, Andobo, ii.1957 ^ c? 
(P. Griveaud). 

Madagascar Sud Quest : station agricole du Bas Mangoky, i $ ; Tulear Saint 
Augustin, ii.1956, I $ (A. Robinson). 

Subfamily COPTACRIDINAE 

PAMPHAGELLA Bruner, 1910 

Of medium size. Integument granose. Antenna compressed, slightly widened on basal and 
apical part, shorter than head and pronotum together. Fastigium of vertex narrow, strongly 
sloping forwards and obtusangularly merging with frontal ridge. Interocular distance wider 
than basal antennal segment ; frons oblique, straight, frontal ridge narrow, between antennae 
slightly widened ; slightly concave, with sharp lateral carinulae, vertex in profile angular. 
Pronotum strongly tectiform, in female almost crest-shaped, crossed by three sulci ; metazona 
much shorter than prozona, its posterior margin acutangular. Prosternal process low, trans- 
verse, with widely rounded apex. Mesostemal interspace short, much wider than its length. 
Elytra lobiform, lateral, covering large tympanum ; precostal area expanded ; apex rounded ; 
venation and reticulation rough. Hind femur short and wide ; upper carina strongly serrated ; 
lower external area located on ventral side ; lower lobe of hind knee rounded. External apical 
spine of hind tibia absent. Arolium of moderate size. Male supra-anal plate elongate-angular. 
Cercus compressed, angular, upcurved, with irregularly serrated upper margin. Subgenital 
plate short, subconical. Female subgenital plate slightly trilobate. Ovipositor short, with 
robust valves, slightly curved at apices. 

Type species : Pamphagella comoroensis Bruner, 1910. 

This genus was described originally as a member of the family Pamphagidae, but 
it was proved (Dirsh, 1958) to belong to the family Acrididae and most probably to 
the subfamily Coptacridinae. Since, however, the only known male type is in poor 
condition, not allowing a study of its phallic organ, the subfamily relationship of the 
genus is doubtful. 

Pamphagella comoroensis Bruner, 1910 
(Text-fig. 20) 

Antenna fourteen-segmented. Pronotum of male less tectiform than in female. General 
coloration brown. Lower and internal side of hind femur reddish. 

Length of body ^ 13-5, $ 24-5-25 ; pronotum <^ 3-4, $ 7-8 ; elytron ^ 2-5, $ 4-5-5 : hind 
femur ^ 7-2, $ lo-ii mm. 

Madagascar Sud-Quest, Sept Lacs 50 m. Tulear, vi-vii.1958, i ?, (A. R.). 
Type locahty : Comoro Is., Circle de Bombao, i c?, I ?• ^ type in Berhn Museum. 

SubfamUy GALLIPTAMINAE 

CALOPTENOPSIS I. Bolivar, 1889 

Caloptenopsis decisa (Walker, 1870) 

(Text-fig. 21) 
Caloptenus decisus "Walker, 1870 : 700. 
Caloptenus baliensis Brancsik, 1893 : 195, syn. n. 
Caloptenopsis madagascariensis Sjostedt, 191 8 : 16, syn, n. 




Fig. 20. Pamphagella comoroensis Bruner, 1910. i, female, paratype. 2, male, type. 
3, face. 4, male antenna. 5, male sternum. 6, the same, female. 7, male cercus. 
8, end of female abdomen, from above. 9, the same, from below. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



313 



The types of C. decisus Walk, and C. madagascariensis Sjos. were compared and 
found identical. The type of C. haliniensis Branc. is lost, but the description fully 
agrees with the types of the above-mentioned species. 

(J. Of medium size and comparatively slender for the genus. Lateral carinae of pronotum 
slightly regularly excurved ; posterior margin of pronotum obtusangular, with slightly incurved 
sides of the angle. Prostemal process slightly flattened in front. Elytra exceed end of 
abdomen, comparatively narrow. Hind femur widened, with moderately expanded lower 
margin. Internal spur of internal side of hind tibia elongated, with moderately curved apex 
and dense hairs. Male supra-anal plate elongate, acutely angular, with median longitudinal 
sulcus and slightly upcurved sides. Cercus comparatively long and narrow, slightly incurved, 




Fig. 21. Caloptenopsis decisa (Walker, 1870). i, male. 2, male cercus, lateral view. 
3, phallic complex, with ectophallic membrane and epiphallus removed. 4, the same, 
with most part of cingulum removed. 5, epiphallus. 



314 V. M. DIRSH 

with long, comparatively narrow upper and long narrow, subacute lower lobe. Subgenital plate 
short, subconical. Phallic complex robust. Apodemes long, narrow ; valves of penis robust, 
with thick flexure ; gonopore process short ; arch of cingulum and valve of cingulum large, 
robust. Epiphallus trapezoidal ; ancorae short, robust. 

General coloration brown. Elytron with few ochraceous spots in medial area and on apical 
part. Internal side of hind femur dark brown, with pre-apical ochraceous fascia, above with 
two faint fasciae ; upper lobe of hind knee blackish or brown with blackish prebasal spot, 
lower lobe ochraceous. Hind tibia greyish or brownish ; with slight purplish or bluish tinge. 

$. As the male, but larger. Ovipositor short, with robust valves, curved at apices. 

Length of body ^ 21-22, $ 27-32 ; pronotum (J /\^-^-^-6, $ 5-5-2 ; elytron (J 17-2-18, $ 22-5- 
26 ; hind femur ^ 13-5-14, $ i7-9-i9-2 mm. 

Madagascar Centre: Tananarive Tsimbazaza, vii.1948, i (^, 3 $ (P. Cachan) ; 
Arivonimamo, x.1948, i $ (E. Razafimandimby) ; Ambatofinandrahana, vii.1957, 
2 (^, 2 $ (P. Griveaud). 

Madagascar Sud : Morombe, lac lotry, 40 m., vii.1957, i 9 (P- Griveaud); 
Beloha, 100 m., vi.1957, i $ ; La Sakoa, i $ (R, P. Cattala). 



Subfamily EYPREPOCNEMIDINAE 

The subfamily Eyprepocnemidinae is abundantly represented in Madagascar, and 
several genera and species described below are endemic in the island. Another 
interesting point is that all Madagascar species of this subfamily manifest a wide 
range of variabihty in pattern and coloration, suggesting that they probably are in 
a progressive stage of evolution on this island. 

Key to Genera 

1 (2) Male cercus with acute apex (Text-fig. 22) . . EYPREPOCNEMIS (p. 314) 

2 (i) Male cercus with widened rounded apex. 

3 (4) Elytra lobiform, lateral (Text-fig. 23) . . . MALAGACETRUS (p. 316) 

4 (3) Elytra fully developed or shortened but not lobiform. 

5 (8) Antenna filiform or almost filiform with slightly widened medial part. 

6 (7) Lateral carinae of pronotum obliterated. General coloration black with red 

pattern (Text-fig. 24) TENEBRACRIS (p. 318) 

7 (6) Lateral carinae of pronotum distinct. General coloration not black, without red 

pattern (Text-figs. 25-28) HETERACRIS (p. 319) 

8 (5) Antenna strongly ensiform (Text-fig. 29) ... HORAEOCERUS (p. 324) 



EYPREPOCNEMIS Fieber, 1853 

Eyprepocnemis brachyptera Bruner, 1910 

(Text-fig. 22) 

(?. Of medium size. Antenna filiform, shorter than head and pronotum together. Fasti- 
gium of vertex parabolic, concave ; vertex with median carinula ; frontal ridge very slightly 
narrowing towards apex. Lateral carinae of pronotum straight, diverging backwards, in 
metazona almost obliterated. Prosternal process subcylindrical, with apex oblique towards 
front, Mesosternal interspace narrow, twice as long as its width. Elytra shortened, reaching 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



315 



sixth abdominal tergite. Cercus narrow, with acute, downcurved, slightly compressed apex. 
General coloration brownish ; longitudinal spot on dorsum of pronotum narrow, dark brown ; 
lateral lobe, in middle, with blackish irregular spot, in middle of which there is ochraceous callous 
convexity ; lower and lower internal part of hind femur orange-red ; base of hind knee with 
black transverse fascia, upper lobe black, apical part of lower lobe light ochraceous ; basal half 
of hind tibia blue with two whitish rings, apical half red ; spines of the same colour as tibia, 
with blackish apices. 

$. As the male, but larger. Elytra shortened or reaching end of abdomen. Subgenital 
plate trilobate. Ovipositor robust, with short, curved valves. 

Length of body (J 17-20, $ 28-33 ; pronotum ^ 3-6-47, ? 5-4-6-5 ; elytron ^ 9-14, $ 13-20 ; 
hind femur <J 11-14, $ 12-5-21 mm. 

E. brachyptera varies in body size and relative length of elytra, which in some 
specimens reach the sixth abdominal tergite, in others exceed end of abdomen. 

The species is quite distinct from any African short-winged species. 

Madagascar Centre : Ambositra, foret d'Ankazomivady, 1,640 m., vii.1957 5 ^, 
6 ? (A, Robinson) ; Ambalavao, Andringitra, plateau Soaindrano, 2,090 m., 
15.1.1958, I $ (R. Paulian) ; Tsarazafy, 15.vii.1913, 2 ? (R. Beck). 

Eyprepocnemis smaragdipes Bruner, 1910 

Euprepocnemis smaragdipes Bruner, 1910 : 643. 

Euprepocnemis malagassus I. Bolivar, 1914 : 11 ; Dirsh, 1958 : 41, 

cJ. Of medium size. Antenna filiform, about as long as head and pronotum together. 
Fastigium of vertex angular, almost parabolic, with obtuse apex, above slightly concave, with 
strong median cannula ; frontal ridge slightly narrowing towards apex. Dorsum of pronotum 




Fig. 22. Eyprepocnemis brachyptera Bruner, 1910. i, male, type. 2, head and pronotum 
from above. 3, prosternal process, frontal view. 4, epiphallus, 



3i6 V. M. DIRSH 

slightly raised in prozona ; median carina sharp ; lateral carinae obtuse, straight, diverging 
backwards and almost obliterated in metazona ; prozona longer than metazona. Prosternal 
process subcylindrical, with rounded apex. Mesostemal interspace about as long as wide. 
Elytra and wings fully developed, exceeding end of abdomen. Cercus narrow, in apical part 
compressed and slightly downcurved. General coloration brownish ; longitudinal spot on 
dorsum of pronotum narrow, dark brown ; lateral lobe of pronotum with brownish, oblique, 
longitudinal stripe on brownish ochraceous background ; external side of hind femur from 
brownish to ochraceous-brown, internal side dark brown, lower surface ochraceous ; hind tibia 
greenish, bluish, greenish grey, sometimes slightly reddish on apical part ; spines whitish, with 
black tips. Epiphallus as in Text-fig. 22. 

$. As the male, but larger. Elytra reach or exceed end of abdomen. Subgenital plate 
trilobate. Ovipositor robust, with curved valves. 

Length of body ^ 22-5-25-5, $ 27-37 ; pronotum ^ 4-8-5-2, ? 5-2-7 ; elytron ^ 19-19-6, 
$ 2i-5-26*2 ; hind femur ^ i4*3-i5-5, $ 17-23. 

E. smaragdipes varies in body .size, relative length of elytra and coloration of 
hind tibia, which is bluish grey, greenish grey and sometimes bluish grey at the 
basal part and reddish in apical part ; there are all intermediate forms between 
them. 

E. smaragdipes is nearest to E. kalkudensis Henry, 1937 from Ceylon. 

It is probable that records of E. plorans from Madagascar by Saussure, 1899 and 
Brancsik, 1893 refer to E. smaragdipes, since plorans was never again recorded from 
the island. 

Madagascar Centre: Tananarive Tsimbazaza, 13. xi. 1947, i $ (R. Pauhan) ; 
Manjakandriana, Ankadimanga, xii.1957, 2 $ (J. Elie). 

Madagascar Est: He Sainte Marie, Ambatoroa, v. 1959, 3 (^, i $. (E. Razafi- 
mandimby) ; Tamatave, iv.1959, 2 (^, i $ (E. Razafimandimby) ; Perinet, 2 ?; 
Ifanadiana, Ranomafand, i (^, i $ (R. Pauhan). 

Madagascar Sud : lac Tsimanampetsotsa, v. 1959, i 9 (R- Pauhan). lies Comores : 
Moroni, 2.iv.i948, i $ (P. Cachan). 



MALAGACETRUS gen. n. 

Of medium size. Integument rugose. Antenna filiform, half again as long as head and 
pronotum together. Fastigium of vertex narrpw, angular, with obtuse apex, roundly merging 
with frontal ridge, obtuse lateral and median carinulae present ; frons oblique ; frontal ridge 
fiat, without lateral carinulae, narrowed at apex. Pronotum slightly tectiform ; median and 
lateral carinae obtuse, lateral carinae irregular and slightly diverging backwards ; three deep 
sulci crossing dorsum ; metazona much shorter than prozona, its posterior margin obtusangular. 
Prosternal process obtusely conical. Mesosternal interspace wider than its length. Elytra 
lobiform, lateral, reaching second abdominal tergite and covering tympanum, its anterior margin 
strongly excurved and apex subacute. Hind femur moderately robust. Aroliurh large. Last 
abdominal tergite with a pair of acute projections. Supra-anal plate elongate, with obtusely 
angular apex. Cercus exceeds end of subgenital plate, widened, in apical half strongly lamelli- 
formly compressed and downcurved, with apex rounded. Paraproct with rounded lower apical 
projection. Subgenital plate short, in profile rounded. 

Only male'known. 

Type species : Paracaloptenus rubripes Chopard, 1919. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 317 

Malagacetrus rubripes (Chopard, 1919), comb. n. 

(Text-fig. 23) 
Paracaloptenus rubripes Chopard, 1919 : 154. 

General coloration dark brown. Antenna very long, with alternate brown and yellowish 
rings. In front and behind eyes yellow vertical stripes present. Pronotum uniformly blackish 
brown. All legs bright cinnabar-red ; hind femur on upper side with three slight, brownish 
fasciae ; hind knee black ; hind tibia at base black, the rest with spines and tarsus cinnabar-red. 
Apical half of cercus reddish. 

Betsimisaraka du Centre, i cJ type. 

The species was described originally as Paracaloptenus, but study of the type 
showed that the shape of cercus, end of abdomen, pronotum and head are as found 
in Eyprepocnemidinae, and not in Paracaloptenus. In Eyprepocnemidinae however 
there is no genus into which rubripes could be included and a new genus had to be 
erected. It can be placed near Heteracris, from which it differs strongly by micro- 
pterism and general shape of body. 




Fig. 23. Malagacetrus rubripes (Chopard, 1919). i, male, type. 2, end of male abdomen, 

lateral view. 3, male cercus. 



3i8 



V. M. DIRSH 



TENEBRACRIS gen. n. 

Of medium size. Integument rugose. Antenna filiform, longer than head and pronotum 
together. Fastigium of vertex widely angular with truncate apex, edged by low lateral 
carinulae, median carinula continuous with occipital one. Frons in profile oblique ; frontal 
ridge low, flat, with small depression under ocellus and rugulose surface ; gradually narrowing 
at apex. Pronotum weakly tectiform with distinct median and fully obliterated lateral carinae. 
Three deep transverse sulci crossing dorsum ; metazona shorter than prozona, with obtus- 
angular posterior margin. Prosternal process subconical, with obtuse, rounded apex. Elytra 
and wings fully developed, exceeding end of abdomen ; membrane thick semi-transparent, 
reticulation dense. Hind femur comparatively robust. Spines of hind tibia sparse. Last 
abdominal tergite of male with a pair of angular projections. Supra-anal plate elongate angular. 
Cercus strongly compressed and widened in apical half, with in- and down-curved, rounded 
apex. Subgenital plate short, rounded. Epiphallus with moderately wide bridge, small 
incurved ancorae and large multi-lobate lophi. 

Type species : Tenebracris splendens sp. n. 

The new genus has no close affinity with any known genera of Eyprepocnemidinae, 
but may be placed near Heteracris, with which it shares filiform antenna and shape 
of cercus, but from which it differs strongly by the obUterated lateral carinae of 
pronotum and by shape of fastigium of vertex. 







Fig. 24. 



Tenebracris splendens sp. n. i, male, type. 2, end of male abdomen, lateral 
view. 3, the same, from above. 4, epiphallus. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 319 

Tenebracris splendens sp. n. 

(Text-fig. 24) 

<^. Type. Black, with faintly marked pair of dark brown lateral stripes on dorsum of 
pronotum. Palpi whitish. Wing deep blue, with apex of remigium infumate. Anterior and 
middle femur with cinnabar-red rings near base. Both sides of hind femur cinnabar-red, with 
three black fasciae and black knee ; hind tibia black with cinnabar-red ring at post-basal part. 

Female unknown. 

Length of body 27 ; pronotum 6-2 ; elytron 19 ; hind femur 17 mm. 

Madagascar Centre: Ambatolampy, Andranotobaka, 1,400 m., vi.1957, i <^, 
paratype (P. Griveaud). 

Madagascar Ouest : Antsalova, foret de I'Antsingy, Andobo, 190 m., ii.1957, i c^, 
type (P. Griveaud). 

Type in Paris Museum, paratype in the British Museum (Natural History). 



HETERACRIS Walker, 1870 

Heteracris Walker, 1870 : 655. 

Thisoicetrus Brunner, 1893 : 150 ; Dirsh, 1958 : 53. 

Bibulus I. Bolivar, 1914 : 31 ; Popov, 1950 : 133. 

Key to Madagascar Species 

1 (8) Elytra and wings fully developed, exceeding end of abdomen. 

2 (7) Dorsum of pronotum with dark longitudinal stripe. 

3 (4) External side of hind femur with two dark, wide, transverse fasciae (Text-fig. 25) 

finoti I. Bolivar 

4 (3) External side of hind femur on basal half with narrow longitudinal dark stripe 

near upper cannula and small spot on apical half (Text-fig. 26), or with numerous 
small spots (Text-fig. 27). 

5 (6) External side of hind femur on upper basal half with dark, narrow stripe and 

small spot on apical half. Lower angle of paraproct in male narrow (Text-fig. 26) 

sikorai I. Bolivar 

6 (5) Pattern on external side of hind femur disintegrated into numerous dark, small 

spots (Text-fig. 27). Lower angle of paraproct in male wide (Text-fig. 27) 

zolotarevskyi sp. n. 

7 (2) Dorsum of pronotum without dark longitudinal stripe . . nobilis Brancsik 

8 (i) Elytra and wings short, reaching base of supra-anal plate (Text-fig. 28) reducta sp. n. 



Heteracris finoti (I. Bolivar, 1914), comb. n. 

(Text-fig. 25) 

Thisoicetrus finoti 1. Bolivar, 1914 : 2. 
Thisoicetrus praestans Carl, 191 6 : 500, syn. n. 

The two species, finoti and praestans, were described from males and females. I 
am designating here the males as types for them both. These were compared and 



320 



V. M. DIRSH 



found identical, a slight difference in pattern on the external side of the hind femur 
not exceeding the range of individual variability. 



(J. Large. Antenna longer than head and pronotum together, shghtly thickened in middle 
part. Fastigium of vertex narrowing towards apex and merging roundly with frontal ridge, 
convex above, with strong median carinula ; frontal ridge narrowed above antennae. Median 
carina of pronotum high and sharp ; lateral carinae straight, almost parallel, only slightly 
diverging backwards ; dorsum slightly tectiform, more so in prozona than in metazona. 
Prosternal process subconical, slightly antero-posteriorly compressed with obtuse apex. Meso- 
sternal interspace longer than its width. Elytra and wings fully developed. Cercus with 
strongly compressed, moderately widened, in- and down-curved apical half, with apex rounded. 
Lower angle of paraproct narrowed and protruding backwards. Subgenital plate very short, 
rounded in profile. General coloration brownish olive-green. Apical part of antenna ochra- 
ceous. Head ochraceous above and with blackish stripes on sides below eyes ; labrum with a 
pair of red spots on sides. Dorsum of pronotum with narrow, longitudinal, parallel-sided, 
blackish stripe. Elytra uniformly olive-green, with dorsal part lighter than lateral. Wings 
blue, with infumate apex. Hind femur ochraceous with large longitudinal basal blackish spot, 
transverse blackish fascia in apical half and blackish knee on both sides ; above with two 
blackish fasciae. Hind tibia with base black, followed by yellow and further by black rings, 
the rest and tarsus bright cinnabar-red. Cercus reddish. 

$. Much larger than male. Antenna not at all thickened. 

Length of body (J 32, $ 47 ; pronotum cj 7, ? 10-5 ; elytron ^ 26, $ 40 ; hind femur ^ 20-5, 
$ 27-5 mm. 



Madagascar Est, Sambava, Marojejy, Ambinanitelo, 500 m., xii.1958, i 
(E. Raharizonina). 



(^ 






Fig. 25. Heteracvis finoti I. Bolivar, 1914. i, male antenna. 2, dorsum of pronotum. 
3, hind femur, external side. 4, end of male abdomen, lateral view. 5, epiphallus. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



321 



Heteracris sikorai (I. Bolivar, 1914), 

(Text-fig. 26) 
Thisoicetrus sikorai I. Bolivar, 1914 : 25. 
Thisoicetrus brevicornis Carl, 191 6 : 498, syn, n. 

The female type of Th. sikorai I. Bol. was compared with the original series of 
males and females of Th. brevicornis Carl, and the two species were found identical. 
One of the males of Th. brevicornis is designated here as the specific type. 

(^. Antenna slightly shorter than head and pronotum together, slightly thickened in middle 
part. Fastigium of vertex angular, with obtuse apex, above with wide, obtuse median carinula ; 
frontal ridge flat, in upper half high, narrowed above antennae. Dorsum of pronotum slightly 
tectiform ; median carina strong but obtuse ; lateral carinae obtuse, slightly diverging back- 
wards ; metazona shorter than prozona, its posterior margin roundly excurved, almost truncate. 
Prostemal process cylindrical, with rounded apex. Mesosternal interspace longer than its 
width. Elytra and wings fully developed, exceeding end of abdomen. Cercus wide, with 
apical part incurved and strongly downcurved, apex rounded. Lower angle of paraproct narrowed 
and protruding backwards. General coloration brown. Head brownish ochraceous, with 
blackish vertical stripe below eyes ; above with brownish longitudinal stripe and brown stripe 
behind eye. Dorsum of pronotum dirty ochraceous or greenish, with narrow, brown longi- 
tudinal stripe with blackish margins, which include narrow, light stripe along median carina. 
Elytra uniformly brown or with indefinite dark brown spots, dorsal part greenish or ochraceous. 
Wing bluish, with slightly infumate apex. Hind femur brownish ochraceous, pre-apical part 
orange-yellowish ; upper part of external side with indefinite longitudinal brownish stripe ; 
similar, but darker stripe on internal side ; lower side orange-red ; knee blackish. Hind tibia 
dark bluish or purplish. 

$. As male, but much larger. Antenna fiUform, not thickened. 

Length of body ^ 25-26, ? 41-48 ; pronotum ^ 5-5-5, § 8-8-6 ; elytron <S 20-22, ? 33-5-35 ; 
hind femur ^ 15-5-18, $ 26-5-29 mm. 

Madagascar Centre : reserve naturelle de Zakamena, Nosivola, i c^ ; Manjakan- 
driana, Ambatoloana, xi.1950, i $ ; Ambositra, mts. Antety, xii.1894, i $ ; foret 
d'Ambohimitombo, i.1895, i $ ; Ambalavao, Andringitra, cirque Boby, 2,600 m., 
i.1958, 5 c?, 4 ? (R. Paulian) ; id., plateau Soaindrano, 2,090 m., i.1958, i $ (R. 
Paulian) ; id., foret Vakoana, 1,530 m., i.1958. 

Madagascar Est : Perinet, Sahamaloto, i.1949, i ? (P. Cachan). 




m 



m 1 



1 

Fig. 26. Heteracris sikorai I. Bolivar, 1914. i, dorsum of pronotum. 2, hind femur, 

external side. 3, end of male abdomen, lateral view. 4, epiphallus. 
ENTOM. 12, 6. 2^ 



322 



V. M. DIRSH 



Heteracris zolotarevskyi sp. n. 

(Text-fig. 27) 

cj. Type. Of medium size. Antenna filiform, longer than head and pronotum together. 
Fastigium of vertex narrowing forwards, with truncate apex ; above with low median carinula 
which is continuous with carinula of vertex and occipital one ; frontal ridge comparatively 
low, flat, slightly narrowed at apex. Dorsum of pronotum slightly tectiform ; median carina 
strong and comparatively sharp ; lateral carinae obtuse, diverging backwards ; metazona 
slightly shorter than prozona, its posterior margin obtusangularly excurved. Prosternal process 
antero-posteriorly compressed, almost transverse, with rounded apex. Mesosternal interspace 
about as long as its width or slightly longer. Hind femur moderately slender. Elytra and 
wings fully developed, exceeding end of abdomen. Cercus with strongly incurved and down- 
curved apical part and obtuse, almost rounded apex. Paraproct large, with lobiform projection 
on lower end. Subgenital plate short, rounded. General coloration brownish. Head ochra- 
ceous-brown, with dark brown stripe below eye, above and behind eye with dark brown longi- 
tudinal stripe. Dorsum with wide, widening backwards, longitudinal, brown stripe. Elytra 
brown with ochraceous spots ; wings blue, with infumate apex. Hind femur ochraceous ; 
external side with brownish fasciae, disintegrated into spots ; knee dark brown. Base of hind 
tibia blackish, followed by light whitish ring, and again by brownish ring ; rest of tibia and 
tarsus blue. 

$. Paratype. As male, but larger ; no other differences, except sexual. 

Length of body $ 24-5, $ 38-5 ; pronotum cJ 5, ? 7-6 ; elytron ^ 20-5, § 34-5 ; hind femur 
(J 15-6, ? 23-3 mm. 

Madagascar Sud : Ejeda, ii.1928, 4 ^ (including type), 6 $ (paratypes). B. N, 
Zolotarevsky. Type in the British Museum (Natural History). 

The new species belongs to the same group as H. guineensis Krauss and H. 
pulchripes jeanneli I. Bol. It differs from them by the fastigium of vertex narrowed 
in apical part, male cercus, pattern and coloration. 

The species is named after the eminent locust expert, Dr. B. N. Zolotarevsky, who 
first collected it. 

Heteracris nobilis (Brancsik, 1893), comb. n. 
Euprepocnemis nobilis Brancsik, 1893 : 195. 

The single female type of the species is lost, but its description and a reasonably 
good coloured figure suggest that the species is distinct. It differs from H. finoti by 




4 ^ -^ -^ 1 

Fig. 27. Heteracris zolotarevskyi sp. n. i, dorsum of pronotum. 2, hind femur, external 
side. 3, end of male abdomen, lateriil view. 4, epiphallus. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



323 



the absence of the dark longitudinal stripe on the dorsum of pronotum and by the 
single transverse stripe on the external side of hind femur. 

Length of body 50 ; pronotum 11 ; elytron 41 ; hind femur 28 mm. 

Type locality : " Madagascar, Ins Nossibe ". 



Meter acris reducta sp. n. 

(Text-fig. 28) 

(J. Type. Of medium size. Antenna shorter than head and pronotum together, almost 
fihform, slightly thickened in middle part. Fastigium of vertex angular with obtuse apex ; 
above convex, with slight median carinula, continuous with carinula of vertex ; frontal ridge 
flat, narrowed at apex. Dorsum of pronotum slightly tectiform ; median carina strong, obtuse ; 
lateral carinae obtuse, slightly excurved and slightly diverging backwards ; metazona shorter 
than prozona, its posterior margin obtusangularly excurved. Prosternal process subcylindrical, 
slightly antero-posteriorly compressed with rounded apex. Mesosternal interspace slightly 
wider than its length. Elytra and wings shortened, reaching base of supra-anal plate. Hind 
femur comparatively slender. Cercus strongly downcurved and moderately incurved, narrow- 
ing to round apex. Paraproct with narrowed and slightly upcurved posterior angle. Subgenital 
plate short, in profile rounded. General coloration brown. Head brownish ochraceous, with 
black stripe under eye ; above with longitudinal brown stripe and black stripe behind eye. 
Dorsum of pronotum with dark brown longitudinal stripe, moderately wide, with parallel sides, 
including light narrow stripe along median carina. Elytra brown without pattern, with greenish 
dorsal part. Wings blue, with slightly infumate apex. Hind femur dirty ochraceous ; on 
external side, in basal upper part, with indefinite brown longitudinal stripe, and small brown 
spot in apical half ; pre-apical part bright ochraceous ; knee blackish ; internal side more 
lightly coloured, with the same pattern as external. Base of hind tibia blackish, followed by 
yellow ring and afterwards with narrow brownish ring ; rest of tibia and tarsus bright red. 

$. Paratype. As male, but larger. Antenna only very slightly thickened in middle part. 
Hind tibia of variable shade of purple. No other differences, except sexual, from the male. 

Length of body (J 24, $ 38-5 ; pronotum ^ 5-4, $ 7-3 ; elytron ^ 12-5, $ 17*3 ; hind femur 
(J i6'5, ? 22 mm. 

Madagascar Sud : reserve naturelle XI, mont Andohahelo, 1,800 m. i (^ (type), 
4 $ (paratypes) (R. PauUan). Type and paratypes in Paris Museum, one paratype 
in the British Museum (Natural History). 




[m 



I I 



LHJ 



4 — U^ J 1 

Fig. 28. Heteracris reducta sp. n. i, dorsum of pronotum. 2, hind femur, external side. 
3, end of male abdomen, lateral view. 4, epiphallus. 



324 V. M. DIRSH 

The new species is near to H. sikorai but differs by reduced elytra and wings, by 
shape of male cercus and less developed median carinula of fastigium of vertex. 

HORAEOCERUS Saussure, 1899 

Of medium size. Integument slightly rugose. Antenna of male from strongly to moderately 
ensiform, about as long as head and pronotum together ; in female weakly widened in median 
part. Fastigium of vertex elongate angular, truncate at apex and tectiformly convex, with 
medium carinula. Frons oblique, frontal ridge in upper half high, with slightly expanded 
lateral margins, flat, weakly depressed under ocellus. Pronotum slightly tectiform with strong 
median and moderately strong, slightly diverging backwards, lateral carinae ; metazona shorter 
than prozona, its posterior margin widely obtusangular. Prosternal process cylindrical, slightly 
antero-posteriorly compressed, with rounded apex. Elytra fully developed, with semi-trans- 
parent membrane and rather sparse reticulation. Hind femur slender. Last abdominal tergite 
of male with a pair of small acutangular projections. Supra-anal plate in both sexes elongate 
angular, in male with slightly attenuate apex. Male cercus widened in apical two-thirds, 
strongly compressed, with downcurved, rounded apex ; in female obtusely conical. Male 
subgenital plate short, rounded ; in female with slightly angular apex. Phallic complex large, 
with U-shaped apodeme, very large rami and expanded distal part of ectophallic membrane, 
which is sclerotized and covering almost completely apical valves of penis and valves of cingulum. 
Basal valves of penis large, connected with apical ones by thick flexure. Epiphallus with large, 
curved ancorae and large lobiform lophi. Valves of ovipositor short, robust, with curved apices. 

Type species : Horaeocerus nigricornis Saussure, 1899. 

Horaeocerus nigricornis Saussure, 1899 
(Text-fig. 29) 

Antenna in male ensiform, in female slightly ensiform. Fastigium of vertex narrow, angular, 
roundly merging with frontal ridge. General coloration brownish or olive-green ; antenna 
blackish, with ochraceous apical part ; gena yellow, under eye black vertical strip>e ; head above 
with brownish, wide, longitudinal stripe, which in basal part is continuous with dark brown 
longitudinal stripe of dorsum of pronotum ; elytron brownish or olive-green, without pattern ; 
hind wing blue, bluish or infumate, with darkened apex ; hind femur on both sides yellowish, 
with three black oblique, incomplete, transverse fasciae ; knee brown ; base of hind tibia dark 
brown, followed by lemon-yellow ring and by wide dark brown ring, the rest pale purplish ; 
spines whitish, with brown apices. 

The species was originally described from male and female. The male is desig- 
nated here as the type. 

A series of specimens shows that the size of body is variable : for male 28-32, for 
female 40-47 mm. (in original description the male is 21, the female 36 mm.). Wing 
in original description : "... sordidae vitreae, venis fuscis ". In the series 
studied wing coloration varies from weakly infumate to bluish, and in dark coloured 
specimens to deep blue. General coloration varies from brown to dark olive-green. 

Madagascar Centre : Tananarive, xi.1952, i 3" (E. S. Brown) ; id., Tsimbazaza, 
2 (^, 3 $ ; Manjakandriana, Ankadimanga, xii.1957, i $ (J. Elie) ; Ankazobe, foret 
d'Ambohitantely, xii.1954, 2 (J. 

Madagascar Est : Maroantsetra, Ivontaka, 8 m., (P. Soga et E. Raharizonina) ; 
Tamatave, 27.xii.1947, i $ ; Perinet, i ^J, i $. 




k 



Fig. 29. 1-7, Horaeocerus nigricornis Saussure, 1899. i, male. 2, male antenna. 3, 
dorsum of pronotum. 4, end of male abdomen. 5, phallic complex, with ectophallic 
membrane and epiphallus removed. 6, the same, with most part of cingulum removed. 
7, epiphallus. 8-10, Horaeocerus antennatus I. Bolivar, 1914, type. 8, male antenna. 
9, hind femur, external side. 10, end of male abdomen, lateral view. 



326 V. M. DIRSH 

Madagascar Quest : foret de I'Ankaraiantsika, Ampijoroa, 170 m., i.1957, i $ ; 
Antsalova, foret de TAntsingy, Andobo, ii.1957, i (^ (P. Griveaud) ; Sakaraha, 
Lambomakandro, xii.1956, i $. 

Horaeocerus antennatus I. Bolivar, 1914 
(Text-fig. 29) 

The species was described from the male and the female. I am designating here 
the male as the type. 

The type and the female paratype are discoloured and damaged by previous 
preservation in spirit. The differences from H. nigricornis Sauss. are the shape of 
male antenna, which in H. antennatus is about twice as wide as in H. nigricornis, 
and the cercus which is less curved and wider than in nigricornis. The females do 
not differ at all. 

Type locaUty : " Madagascar ". 

Subfamily CATANTOPINAE 

Key to Genera 
Fully winged. 
External apical spine of hind femur present. Tarsus longer than half length 

of tibia (Text-fig. 30) SEYRIGACRIS (p. 327) 

External apical spine of hind femur absent. Tarsus shorter than half length 

of tibia CATANTOPS (p. 326) 

Elytra and wings shortened, lobiform or absent. 

Frons, in profile straight. 

Elytra lobiform, lateral. 

Dorsum of pronotum with pair of deep hollows on sides of median carina at 

posterior transverse sulcus (Text-fig. 32) . . . CALDERONIA (p. 331) 

Dorsum of pronotum without hollows (Text-figs. 34, 35) SERPUSILLA (p. 334) 

Elytra strongly shortened, but not lateral, touching one another on dorsal side. 
Male cercus with acute apex (Text-fig. 33) .... AMBREA (p. 331) 

Male cercus with bifurcate apex (Text-fig. 36) . PARASERPUSILLA (p. 338) 
Frons in profile incurved. 
Elytra shortened or lobiform. 
Elytra shortened. Frons in profile strongly incurved. Male cerci straight 

(Text-fig. 37) MANANARA (p. 340) 

Elytra lobiform lateral. Frons in profile slightly incurved. Male cercus 

incurved (Text-fig. 38) PERINETA (p. 342) 

Wings completely absent (Text-fig. 31) . . PSEUDOHYSIELLA (p. 328) 

CATANTOPS Schaum, 1853 
Key to the Madagascar Species 

1 (4) Male cercus with apex simple, rounded. 

2 (3) Larger and slender. Transverse fasciae on external side of hind femur com- 

paratively short and less oblique ....... sacalava 

3 (2) Smaller and less slender. Transverse fasciae on external side of hind femur 

comparatively long and more oblique ..... stenocrobyloides 

4 (i) Male cercus with apex rounded and slightly excised on lower edge , malagassus 



I 


(4) 


2 


(3) 


3 


(2) 


4 


(I) 


5 


(12) 


6 


(9) 


7 


(8) 


8 


(7) 


9 


(6) 


10 


(") 


II 


(10) 


12 


(5) 


13 


(16) 


14 


(15) 


15 


(14) 


16 


(13) 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 327 

Catantops sacalava Brancsik, 1893 

Cantantops sacalava Brancsik, 1893 : 193. 

Catantops acuticercus I. Bolivar, 1902 : 626 ; Dirsh & Uvarov, 1953 : 233. 

Madagascar Sambirano : Nosy Be, Lokobe, 10. ix. 1947, i $ (R. Paulian) ; Nosy 
Komba, crete Nord, v. 1956, i (^ (A. Robinson) ; Nosy Mitsio, 13, 14.1.1960, 2 (^ 
(R. Paulian). 

Madagascar Est: Sambava, Marojejy, Ambinanitelo, 500 m., xii.1958, 2 ^ 
(E. Raharizonina) ; He Sainte Marie, xi.1959, i ^ (E. Razafimandimby) ; Station 
agricole de Brickaville, 2 $ ; Perinet ; Ifanadiana, Ranomafana, 2 (J, i $. 

Madagascar Centre: Manjakandriana, Ankadimanga, xii.1957, i (^ (J. Elie) ; 
Ambatolinandrahana, vii . 1957, 2 J, i $. 

Madagascar Quest : foret de I'Ankarafantiska, Ampijoroa, 2 $, i (^, id., Bevazaha, 
20.1.1950, I cJ ; Antsingy de Bekopaka, vii. 1949, i ^ (R. Paulian) ; Morondava, 
i.1956, I ^ (R. Paulian) ; Sakaraha, Lambomakandro, 4.11.1958, i $ (P. Griveaud). 

Madagascar Sud : Morombe, lac lotry, 8 . vli-1957, i 3* (P- Griveaud) ; Androy, 
Tranomaro, 11. 1954, i $ (J. Elie). 

Catantops stenocrobyloides Karny, 1907 

Madagascar Centre: Manjakandriana, La Mandraka, xll.1952, i ? (N. L. H. 
Krauss) ; id., Ankadimanga, xii.1957, 7 $, 2 cJ (J- Elie) ; Ambatolampy, Andra- 
masina, xi.1956, i $, 2 (^ (R. Paulian) ; Tsiraonomandidy, mont Ambohiby, 
26. V. 1948, 2 $ (R. Paulian) ; Ambositra, foret d'Ankazomlvady, 1,640 m., 23. vii. 
I957» 6 (5^, 6 ? (P. Griveaud) ; Tananarive, Tsimbazaza, 17. iv. 1948, i (^ (R. Paulian). 

Madagascar Est : lie Sainte Marie, Ambatoroa, v. 1959, i (^ (E. Razafimandimby). 

Catantops malagassus Karny, 1907 

Madagascar Centre : Manjakandriana, Ankadimanga, xii.1957, 2 c^ (J. Elie). 
Madagascar Est: Tamatave, vii. 1958, 4 $, i (^ (J. Elie) ; He Sainte Marie, 
Ambohldena, v. 1959, i $ ; id., Ambatoroa, v. 1959, i $ (E. Razafimandimby). 
Madagascar Quest : Morondava, for^t Sud de Befasy, I.1956, i $ (R. Paulian). 
Hes Glorleuses, 5.iv.i948, i ? (P. Cachan). 

SEYRIGACRIS C. BoUvar, 1932 

Of medium size. Integument finely rugose. Antenna filiform, longer than head and 
pronotum together. Head conical ; fastigium of vertex angular, with slight longitudinal 
depression ; interocular distance narrower than basal antennal segment ; frons strongly 
oblique, straight ; frontal ridge wide, with well developed, parallel lateral carinulae. Pronotum 
cylindrical ; dorsum crossed by three sulci ; metazona much shorter than prozona, its posterior 
margin widely obtusangular, almost rounded ; carinae absent. Prosternal process obtusely 
conical. Mesosternal interspace slightly wider than its length, with incurved sides. Elytra 
and wings reaching end of abdomen. Elytron parchment-like with reduced venation and 
reticulation ; without stridulatory specialization. Hind wing widened, with strongly excurved 
external margin ; tympanum present. Hind femur moderately slender. External apical spine 
of hind tibia present. Hind tarsus more than half length of hind tibia, Arolium large. Last 



k 



328 



V. M. DIRSH 



abdominal tergite of male with a pair of large, rounded projections. Supra-anal plate elongate- 
angular. Cercus long, subconical, slightly incurved, with obtuse apex. Subgenital plate short, 
with rounded apex. 
Only male known. 

Type species : Seyrigacris nigrofasciatus C. Bolivar, 1932. 

Seyrigacris nigrofasciatus C. Bolivar, 1932 
(Text-fig. 30) 

(J. Type. General coloration blackish, with yellowish stripes and spots. Antenna black 
with yellowish rings. Face yellowish with black transverse stripe above clypeus and round 
yellowish spot in middle of it. Head and pronotum above with four continuous black stripes 
and yellowish background. Anterior part of elytron, up to radial vein, blackish, rest brownish. 
Hind wing infumate. Hind femur on external side brownish, with a pair of yellowish spots. 
Hind knee black, with lower lobe yellowish. Hind tibia and spines black. Tarsus black with 
yellowish spot on metatarsus and arolium. 

Length of body 24 ; pronotum 5-3 ; elytron 17 ; hind femur 13 mm. 

Only male type, from Rogez, is known. Preserved in Madrid Museum. 



PSEUDOHYSIELLA gen. n. 

Small. Body narrow-cylindrical. Integument rugose. Antenna rod-like, shorter than head 
and pronotum together. Head conical ; fastigium of vertex angular, separated from vertex 
by depression ; interocular distance as wide as basal antennal segment ; frons strongly oblique 
and incurved ; frontal ridge protruding above ocellus, slightly concave in middle, with weak 
lateral carinulae. Pronotum cylindrical ; carinae absent ; three sulci crossing dorsum, first 
being incomplete ; metazona about one-third length of prozona, its posterior margin straight. 
Prostemal process short, widening towards rounded apex. Mesosternal interspace strongly 
constricted in middle. Elytra, wings and tympanum absent. Hind femur slender, not reaching 
end of abdomen. External apical spine of hind tibia absent. Tarsus slightly elongated. 
Arolium large. Last abdominal tergite of male with pair of acutangular projections. Supra- 
anal plate angular, with strong transverse furrow and acutely attenuate apex. Cercus narrow 
conical, straight, slightly upcurved at apex. Subgenital plate acutely conical. Phallic organ 
robust. Basal and apical valves of penis connected by thick, strong flexure. Apical valves of 
penis and valves of cingulum not covered by sheath. Arch of cingulum large, valves of 
cingulum short. Epiphallus bridge-shaped, with short angular, incurved ancorae and small 
lobiform lophi. 

Tj^e species : Hysia inermis Karsch, 1896. 

The new genus superficially resembles Hysiella, but differs from it as follows : 



Pseudohysiella 

Frons incurved ; upper part of frontal ridge 

strongly protruding 
Antenna rod-like, shorter than head and 

pronotum together 
Three sulci crossing dorsum of pronotum 
Mesosternal interspace strongly constricted 

in middle 
Male subgenital plate acutely conical 
External apical spine of hind tibia absent 



Hysiella 

Frons straight ; upper part of frontal ridge 

not protruding. 
Antenna filiform, much longer than head and 

pronotum together. 
Four sulci crossing dorsum of pronotum. 
Mesosternal interspace not constricted, with 

regularly incurved sides. 
Male subgenital plate obtusely conical. 
External apical spine of hind tibia present. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 329 




k 



Fig. 30. Seyrigacris nigrofasciatus C. Bolivar, 1932, type, i, male. 2, head and 
pronotum, from above. 3, face. 4, sternum. 5, end of male abdomen, lateral view. 
6, the same, from above. 7, epiphallus. 



33° 



V. M. DIRSH 

Pseudohy Stella inermis (Karsch, 1896), comb. n. 



(Text-fig. 31) 



Hysia inermis Karsch, 1896 : 278. 
Hy Stella inermis I. Bolivar, 1906 : 395. 



General coloration brownish ochraceous. Sides and lower surface of body reddish. Head 
yellowish, with blackish, irregular spots. Behind eyes and on lateral lobe of pronotum, in- 
definite blackish stripe ; lower margin of lateral lobes yellowish. External side of hind femur 
ochraceous with blackish longitudinal stripe on upper part, interrupted by ochraceous fish-bone 
pattern ; internal and lower internal side bright red ; lower external side blackish ; hind tibia 
above light olive-greenish above, blackish brown below ; tarsus light olive-greenish. 

Female unknown. 

Length of body (J 15-2 ; pronotum 2-3 ; hind femur 8 mm. 

Madagascar Centre : Route de Tamatave, km. 22, 10. x. 1948, i ^, (P. Cachan). 
The type (in Berlin Museum) was described from " Madagascar Australis ". 




Fig. 31. Pseudohy siella inermis (Karsch, 1896). i, male. 2, end of male abdomen from 
above. 3, phallic complex, with ectophallic membrane and epiphallus removed. 
4, the same, with most part of cingulum removed. 5, epiphallus, 



i 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 331 

CALDERONIA I. Bolivar, 1908 

Of medium size ; body robust, fusiform. Integument rugose. Antenna filiform, longer than 
head and pronotum together. Fastigium of vertex in male with angular base and protruding, 
square apical part and excised apex, in female wide trapezoidal, with almost truncate apex ; in 
male with longitudinal concavity, in female fiat, sloping ; interocular distance in male as wide 
as basal antennal segment, in female slightly wider ; frons slightly oblique ; frontal ridge 
between antennae roundly protruding forwards, in male with deep sulcus ; almost obliterated 
below ocellus. Pronotum subcylindrical, slightly constricted in prozona ; dorsum slightly 
inflated, crossed by two sulci ; median carina linear, lateral carinae absent, or discernible in 
frontal part of prozona ; on sides of median carina at posterior sulcus there is pair of very deep, 
oval cavities ; metazona about half length of prozona, its posterior margin straight ; lateral 
lobes of pronotum concave. Prostemal process conical, with wide base and acute apex. 
Mesostemal interspace wider than its length. Elytra lobiform, lateral, covering large tympanum, 
widening towards apex, excised in upper apical part. Hind femur moderately slender. 
External apical spine of hind tibia absent. Arolium moderately large. Last abdominal tergite 
of male with a pair of long projections. Supra-anal plate elongate-angular, with slightly attenuate, 
angular apex. Cercus short, incurved, with obtuse apex. Subgenital plate short subconical, 
with obtuse apex. Phallic complex robust. Sheath of penis absent ; rami large ; basal and 
apical valves of penis connected by long, moderately thin flexure ; apical valves of penis short, 
robust ; valves of cingulum robust ; arch of cingulum large. Epiphallus bridge-shaped with 
large angular ancorae and large, upcurved, wide, lobiform lophi. Female subgenital plate with 
shallowly bilobate apex. Ovipositor short, robust, with valves curved at apices. 

Type species : Colder onia biplagiata I. Bolivar, 1908. 

This genus probably belongs to the group Serpusiae, which is well represented in 
Madagascar, but the phallic complex deviates greatly from any found in the group. 

Calderonia biplagiata I. Bolivar, 1908 
(Text-fig. 32) 

(J. Antenna twenty- three-segmented. Integument of dorsum of pronotum strongly rugose 
and granulose. Lower margin of lateral lobe of pronotum excurved. Elytra with rough net 
of venation, main veins traceable only in base. General coloration brown. First and second 
basal segments of antenna blackish below. Gena and lower part of lateral lobes of pronotum 
dirty ochraceous, upper part of lateral lobe with curved dark brown stripe ; cavities on dorsum 
of pronotum black. External side of hind femur brown, with ochraceous short stripe on basal 
upper part ; internal side black. Hind tibia brownish. 

?. As male, but larger. Pattern and coloration as in male. 

Length of body ^ 15-3, ? 29 ; pronotum cj 5, ? 8 ; elytron cj 3, $ 4 ; hind femur (J 12, $ 16 mm. 

The female type was studied. The male is described here for the first time. 
Madagascar Sambirano : Nosy-Komba, Flanc Nord, v. 1956, i cJ, 2 $ (A. R.). 
Type locality Diego-Suarez. 

AMBREA gen. n. 

Small. Integument moderately rugose. Antenna filiform, slightly widening towards apical 
part, shorter than head and pronotum together. Fastigium of vertex narrow, angular, with 
sUghtly truncate apex ; sides with marginal carinulae and longitudinal concavity between them ; 
interocular distance about as wide as basal antennal segment ; vertex-occipital cannula present. 
Frons oblique, almost straight, frontal ridge narrow, slightly roundly protruding between 
antennae ; fiat, slightly constricted at ocellus ; with lateral carinulae almost obliterated. 



332 



V. M. DIRSH 




Fig. 32. Calderonia biplagiata I. Bolivar, 1908. i, female. 2, male. 3, left elytron. 
4, end of male abdomen, from above. 5, phallic complex, with ectophallic membrane 
and epiphallus removed. 6, the same, with most part of cingulum removed. 7. 
epiphallus, lophi in horizontal position. 8, the same, lophi in vertical position. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



333 



Eyes and ocelli of moderate size. Pronotum subcylindrical, with weakly tectiform dorsum, 
with linear median but no lateral carinae, and three narrow sulci ; metazona much shorter than 
prozona, its posterior margin obtusangular. Prostemal process transverse, spathulate, with 
excised apex. Mesostemal interspace in male as long as its width, in female slightly wider. 
Elytra strongly shortened reaching end of second abdominal tergite and touching one another 
on dorsal side ; precostal area expanded, apex rounded ; venation and reticulation dense. 
Hind femur moderately slender, slightly exceeding end of abdomen ; lower lobes of hind knee 
rounded. Arolium large. Last abdominal tergite of male with pair of moderate-sized projec- 
tions. Supra-anal plate elongate angular, with slightly attenuate apex and upcurved sides ; in 
middle with shallow longitudinal sulcus. Cercus slender, incurved, strongly narrowed in apical 
part and with acute apex. Subgenital plate short, obtusely conical. Phallic complex relatively 
large, with large apodemes and rami. Sheath of penis absent. Basal and apical valves of 
penis connected by thick flexure ; apical valves very large, robust, lobiform. Arch of cingulum 
large ; valves of cingulum small, reduced. Gonopore process long. Epiphallus bridge-shaped, 
with small angular ancorae and large, upcurved, lobiform lophi. Female subgenital plate with 
angular apex. Valves of ovipositor moderately short and slender, with slightly curved apices. 




Fig. 33. Anti)rea acuticerca sp. n. i, male, type. 2, end of male abdomen. 3, male 
left cercus. 4, phaUic complex, with ectophallic membrane and epiphallus removed. 
5, the same, with most part of cingulum removed. 6, epiphallus. 



334 V. M. DIRSH 

Type species : Ambrea acuticerca sp, n. 

This new genus, resembles Paraserpusilla in general appearance. It differs in the 
structure of the male cercus, which in Ambrea is acute and in Paraserpusilla bifur- 
cate ; in the elytra, which in Paraserpusilla reach the third abdominal tergite and 
touch one another at the dorsum, but in Ambrea are lobiform. The two genera 
differ particularly in the structure of the phallic complex, especially in the shape of 
the apical valves of the penis and epiphallus (see Text-figs. 33, 36). 

Ambrea acuticerca sp. n. 

(Text-fig. 33) 

cj. Type. Antenna twenty-segmented. Dorsum of pronotum moderately rugose ; lateral 
lobes of pronotum slightly concave. General coloration brownish. Lateral lobe of pronotum 
with brown, shiny, curved stripe, and diffusing ochraceous spot on lower margin and smaller 
stripe-like spot in upper anterior part ; pleura ochraceous ; elytron brown, with basal two- 
thirds of precostal area blackish. Hind femur brown, internal side dark brown ; hind tibia 
brownish. 

$. Paratype. As male, but larger. Coloration as in male, but duller and without blackish 
spot on elytron. 

Length of body J 15-5, ? 2i'6 ; pronotum (J 4, $ 5 ; elytron ^ 4-2, $ 5 ; hind femur ^ 9-6, 
$ 12 mm. 

Madagascar Nord : Montagne d'Ambre, Les Roussettes 1,100 m., xi-xii.1958, 4 ^ 
(including type) 6 ? (A. Robinson). Type and paratypes in Paris Museum. One 
male, one female paratype in the British Museum (Natural History). 

SERPUSILLA Ramme, 1931 

Small. Integument rugose. Antenna filiform, longer or shorter than head and pronotum 
together. Fastigium of vertex sloping forwards, angular, with obtuse, slightly excised apex 
and longitudinal depression in middle ; interocular distance slightly wider than basal antennal 
segment ; frons oblique, almost straight, with slightly protruding upper part ; frontal ridge 
narrow, with shallow sulcus and obtuse lateral carinulae, gradually diverging downwards ; 
vertex and occiput with strong median cannula. Pronotum strongly or slightly rugose, dorsum 
subcylindrical or slightly tectiform, crossed by three deep sulci, median carina weak, lateral 
carinae replaced by callosities ; metazona about one-third of length of prozona, its posterior 
margin incurved. Prosternal process thick-spathulate, with slightly bilobate apex. Meso- 
sternal interspace slightly longer than its width. Elytra narrow, lobiform, lateral, slightly 
exceeding first abdominal tergite and covering tympanum. Hind femur slender. External 
apical spine of hind tibia absent. Arolium large. Last abdominal tergite of male with a pair 
of small, rounded projections. Supra-anal plate elongate-angular. Cercus narrow, acutely 
conical, straight. Subgenital plate short, conical. Phallic organ robust, with comparatively 
short apodemes and large rami. Basal and apical valves of penis connected by thick flexure. 
Valves of cingulum and apical valves of penis large, not covered by sheath. Arch of cingulum 
large. Epiphallus bridge-shaped, with small, angular ancorae and lobiform lophi. Valves of 
ovipositor slender, with curved apices. 

Type species : Bibracte {?) malagassa Bruner, 1910. 

This genus is related to Auloserpusia Rehn, 1914, but differs in the shape of the 
spathulate prosternal process, which in Auloserpusia is acutely conical, in the less 
rugose pronotum and in the longer elytra. 



J 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 335 

Key to Species 

Integument, especially pronotum, strongly rugose ; median carina almost 
obliterated. 

Pronotum subcylindrical, with comparatively weak lateral callosities. End of 

male abdomen bright red ....... erythropyga Chopard 

Pronotum subcylindrical, but with slightly flattened dorsum and strong lateral 

callosities. End of male abdomen dirty ochraceous . . ochreopyga sp. n. 

Integument moderately rugose ; dorsum of pronotum moderately or slightly 
rugose, lateral lobes shiny ; median carina distinct. 

Fastigium of vertex with deep concavity above. Dorsum of pronotum moder- 
ately rugose. Hind tibia with reddish tinge . . . malagassa Bruner 

Fastigium of vertex only with trace of shallow concavity above. Dorsum of 
pronotum slightly rugose. Hind tibia brownish with slight greenish tinge 

glabra sp. n. 



Serpusilla malagassa (Bruner, 1910) 
(Text-fig. 34) 

Bibracte {?) malagassa Bruner, 1910 : 640. 

Serpusilla malagassa (Bruner, 1910) Ramme, 193 1 : 938. 

cJ. Integument moderately rugose. Fastigium of vertex comparatively wide with deep 
concavity. Pronotum slightly tectiform, with median carina comparatively well developed. 
General coloration brownish. Frons and genae ochraceous ; first and second basal antennal 
segments with dark brown sp>ots. On lateral lobe of pronotum a dark brown longitudinal, 
crescentic curved stripe. Anterior half of elytron shiny, dark brown. Hind femur testa- 
ceous, with brownish longitudinal stripe on both sides, that on external side on lower half and 
not reaching base of femur ; lower side of femur and tibia with reddish tinge. 

$. As the male, but larger. Subgenital plate with angular apex. Valves of ovipositor 
narrow, slender, with curved apices. Brownish stripe on sides of hind femur sometimes dis- 
integrating into several spOts. 

Length of body ^ i2-5-i3-5, $ 17-20 ; pronotum ^ 3-3-5, ? 4-4-5 ; elytron ^ 2-2-4. ? ^"5-3 ." 
hind femur ^ 7-5-9, ? 11 -5-1 2 mm. 

The original material on which Bruner based his description was studied. Here 
the male is designated as the type. 

Type locaHty : " Fianarantsoa, Madagascar ". 
Madagascar Est : Perinet, iv.1951, 3 (^, 5 ? (A. R.). 
Madagascar Centre : Tananarive, ii-iii.1950, 32 (^, 19 $. 

Serpusilla glabra sp. n. 

(Text-fig. 34) 

(J. Type. Integument slightly rugose. Fastigium of vertex narrow, with trace of shallow 
concavity. Pronotum slightly tectiform, with median carina distinct, dorsum only slightly 
rugose. 

General coloration brownish. First and second basal antennal segments with brown spots. 
Frons, clypeus and genae dirty ochraceous. Lateral lobe of pronotum with dark brown longi- 
tudinal crescent-shaped stripe. Costal area of elytron with shiny dark brown longitudinal spot. 
External side of hind femur dirty ochraceous ; on lower part, along lower carinula, a dark 
brown longitudinal stripe with irregular edges, continuous with small basal spot of the same 



336 



M. DIRSH 



colour ; lower and internal side of hind femur brown ; hind tibia brownish, with olive-green 
tinge. 

$. Paratype. As the male, but larger. Apex of subgenital plate obtusangular, almost 
truncate. Ovipositor short, with moderately slender valves, curved at apices. 

Length of body ^ 12-15, ? i9'5-22 ; pronotum ^ 3-2-3-8, ? 5'2-5-3 ; elytron ^ 2-2-3-6, 
? 3-6-4 ; hind femur (J 7-9-2, $ 12-8-13 mm. 






Fig. 34. 1-5, Serpusilla malagassa (Bruner, 1910). i, male. 2, phallic complex, with 
ectophallic membrane and epiphallus removed. 3, the same, with most part of 
cingulum removed. 4, epiphallus. 5, head, from above. G-j, Serpusilla glabra s^p. n. 
6, head, from above. 7, epiphallus. 8, Serpusilla ochreopyga sp. n., epiphallus. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



337 



Madagascar Centre : Manjakandriana, Ambatoloana, vi.1957, i (^ (A. Robinson) ; 
id., La Mandraka, iv.1950, i (^ (A. Robinson). 

Madagascar Quest ; foret de I'Ankarafantsika, Ampijoroa, i $ ; Morafenobe, 
foret Mahajeby, v. 1952, i (^ (type), 4 $ (R. Paulian). 

Type in Paris Museum. One (^ and one $ paratype in the British Museum 
(Natural History). 

Serpusilla ochreopyga sp. n. 

(Text-fig. 34) 

cJ. Type. Body slightly fusiform. Integument strongly rugose. Fastigium of vertex 
narrow, angular, with narrow concavity. Pronotum with slighth' flattened dorsum and strong 
lateral callosities. General coloration blackish brown. Antenna brownish with lighter apical 




Fig. 35. Serpusilla erythropyga Chopard, 1952. i, male, type. 2, head, from above. 
3, end of abdomen, lateral view. 4, phallic complex, with ectophallic membrane and 
epiphallus removed. 5, the same, with most part of cingulum removed. 6, 
epiphallus. 

ENTOM. 12, 6. 29 



338 V. M. DIRSH 

part. Fastigium of vertex, frons, clypeus and genae yellow ; frontal ridge, at ocellus, with 
blackish transverse fascia, which is continuous with large blackish spots at bases of antennae. 
Pronotum blackish, with yellow spot in upper anterior part of lateral lobe ; abdomen brownish, 
with not very distinct yellowish spots ; end of abdomen from sixth segment dirty ochraceous ; 
pleura, in corner between elytron and pronotum, with bright red spot ; elytron black. Anterior 
and middle legs light green. External side of hind femur black, in upper basal half with bright 
red longitudinal stripe, interrupted in posterior part by black spot ; external side of upper 
femoral carina with narrow yellow stripe ; internal side as external, but with less developed red 
pattern. Hind tibia dark green. 

$. Paratype. As male, but larger. Apex of subgenital plate slightly angular. Valves of 
ovipositor slender, with slightly curved apcies. General coloration uniformly brownish, with- 
out pattern. External side of hind femur brown, ochraceous in basal upper part ; lower and 
base of internal side dark red ; hind knee dark red ; hind tibia red. 

Length of body ^ 14-5-15. ? 22-22-5 ; pronotum d* 4, ? 5-5-2 ; elytron (J 3, $ 4 ; hind femur 
(J II, $ 14-6-15 mm. 

Madagascar Est : Andapa, mont Anjanaharibe, 1,600 m., iv.1960, 4 3" (including 
type), 5 $ (P. Soga) ; Sambava, Marojejy, 1,140 m., xi.1959, 3 ^ (P. Soga). Type 
in Paris Museum. One ^ and one $ paratype in the British Museum (Natural 
History). 

Serpusilla erythropyga Chopard, 1952 
(Text-fig. 35) 

Redescription of (^ type. Body cylindrical. Integument strongly rugose and sparsely hairy. 
Fastigium of vertex widely obtusangular. Pronotum subcylindrical, with median carina 
obliterated, lateral carinae represented by moderately developed callosities. General coloration 
blackish brown, with yellowish and red spots. Fastigium of vertex above cheek, below eye, 
frons below ocellus, clypeus and labrum, with large ochraceous spots. Pronotum rust-brownish, 
with brown-blackish spot on dorsum and lateral lobes. All femora yellowish brown ; tibiae and 
tarsi green ; hind knee red. Five basal abdominal tergites dark blackish brown with yellowish 
brown transverse stripes. End of abdomen bright cinnabar-red. 

Length of body ^ 15-5, $ 24 ; pronotum ^ 3-7, $ 3-8 ; elytra cj 3, $ 4 ; hind femur ^ 10, 
$ 13-8 mm. 

Madagascar Sambirano : Mont Tsaratanana, foret de mousses, 1,500 m., i ^, 
type, in Paris Museum. 



PARASERPUSILLA gen. n. 

Of medium size. Integument moderately rugose. Antenna filiform, slightly thickened at 
apex, shorter than head and pronotum together. Fastigium of vertex narrow, angular, with 
incurved sides, slight marginal carinulae extending to vertex, and with sulcus-like concavity ; 
interocular distance narrower than basal antennal segment ; vertex-occipital cannula present. 
Frons oblique ; frontal ridge narrow, slightly roundly protruding between antennae ; at ocellus 
slightly constricted, below it shallowly concave. Eyes and ocelli large. Pronotum subcylin- 
drical with sharp, linear median but no lateral carinae ; three narrow sulci crossing dorsum, 
metazona much shorter than prozona, its posterior margin angular. Prosternal process trans- 
verse, thick, spathulate, with straight or slightly incurved apical margin. Mesosternal interspace 
about as long as wide or, in female, wider than its length. Elytra and wings shortened, reaching 
third abdominal tergite and touching one another at dorsum ; precostal area slightly expanded ; 
apex rounded ; venation and reticulation dense. Wing shorter than elytron. Hind femur 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



339 



moderately robust, reaching or slightly exceeding end of abdomen ; lower lobes of hind knee 
widely rounded. Arolium large. Last abdominal tergite of male with pair of long acutangular 
projections. Supra-anal plate elongate-angular, with upturned sides and narrow median, 
longitudinal sulcus. Cercus incurved, with deeply bifurcate apex. Subgenital plate short, 
subconical. Phallic complex robust. Apodemes large, elongate, rami very large, with expanded 
upper part. Sheath of penis absent ; flexure between basal and apical valves of penis rather 
thick ; arch of cingulum robust ; gonopore process long. Epiphallus bridge-shaped ; ancorae 
small, angular ; lophi large, lobiform. Female subgenital plate with angular apex. Ovipositor 
short, with valves slightly curved at apices. 

Type species : Paraserpusilla furcata sp. n. 

This new genus differs from Serpusilla Ramme, 1931 in the elytra, which touch 
one another on the dorsal side, in the angular posterior margin of the pronotum, 




Fig. 36. Paraserpusilla furcata sp. n. i, male, type. 2, end of abdomen, from above. 
3, male cercus, lateral view. 4, phallic complex, with ectophallic membrane and 
epiphallus removed. 5, the same, with most part of cingulum removed. 6, epiphallus. 



L 



34© V. M. DIRSH 

which in Serpusilla is always incurved and in the bifurcate male cercus, which in 
Serpusilla is conical. The structure of the phallic complex undoubtedly places the 
new genus in the Serpusiae group, but the epiphallus is rather unusual in its strongly 
developed lobiform lophi. 

Paraserpusilla furcata sp. n. 

(Text-fig. 36) 

(J. Type. Antenna twenty-segmented. Dorsum of pronotum regularly, reticulately 
rugose ; a pair of concave callosities on sides ; lateral lobes slightly concave. General coloration 
brown. Scape and pedicel of antenna with dark brown spots. Lateral lobes of pronotum 
brown, shiny ; lower part ochraceous, and with small ochraceous spot on upper part of anterior 
margin. Elytron brown, with shiny dark brown spot, covering basal part of precostal area. 
External side of hind femur brownish ; lower and internal sides reddish, passing to brown on 
apical part. Hind tibia brownish. 

Length of body <S 20, $ 31 ; pronotum <? 5, $ 7 ; elytron <^ 6-8, $ 9-2 ; hind femur S 12, 
? i6"3 mm. 

Madagascar Nord : Montagne des Fran9ais, det. Diego Suarez, ii.igsg, 2 cJ 
(including type), i $ (A. Robinson). Type in Paris Museum. One male paratype 
in the British Museum (Natural History). 

MANANARA gen. n. 

Of submedium size. Integument finely rugose, shiny and sparsely hairy. Antenna thin, 
filiform, much longer than head and pronotum together. Fastigium of vertex strongly sloping 
forwards, narrowing towards apex, and slightly longitudinally depressed in middle ; roundly 
merging with frontal ridge ; interocular distance one-third of the width of antennal scape ; 
vertex and occiput subglobular ; frons slightly oblique and incurved, with upper part protruding 
forwards ; frontal ridge low, roundly protruding above and constricted below ocellus. Prono- 
tum slightly constricted in middle, widening backwards ; weak median carina present ; lateral 
carinae absent ; three sulci crossing dorsum ; metazona much shorter than prozona, its posterior 
margin obtusangular. Prosternal process short, acutely conical. Mesosternal interspace wider 
than its length. Elytra and wings shortened, reaching seventh abdominal tergite ; elytron with 
parchment-like, shiny membrane and thick, reduced venation and reticulation, its anterior 
margin regularly excurved, apex rounded ; wing widened, with strongly excurved external 
margin. Tympanum present. Hind femur exceeding end of abdomen ; external apical spine 
of hind tibia absent. Arolium large. Last abdominal tergite of male with pair of small, 
angular projections ; supra-anal plate angular, with obtuse apex ; cercus straight, acutely 
conical ; subgenital plate short, subconical, with obtuse apex. Basal valves of penis large, 
with very large gonopore processes ; apical valves large, connected with basal ones by thick 
flexure and forming strongly sclerotized proximal projections ; arch of cingulum and valves of 
cingulum large. Epiphallus with small angular ancorae and wide, lobiform lophi. Valves of 
ovipositor moderately long, narrow, slightly curved at apices. 

Type species : Mananara fasciata sp. n. 

The new genus in its pattern and coloration superficially resembles Seyrigacris, 
but differs strongly from it in the pronotum, which is widened towards the posterior 
end, in the presence of a median carina, shortened elytra and wings, in the absence 
of the external apical spine on the hind tibia and in the less elongated hind tarsi and 
straight, acutely conical male cercus. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 341 

Mananara fasciata sp. n. 

(Text-fig. 37) 

(J. Type. General coloration brownish black, with ochraceous, narrow stripes on sides of 
head, pronotum and elytra, and above, along head and median carina of pronotum. Hind femur 
yellow on external side with two black transverse fasciae and small black spot at upper basal 
lobe ; on internal side with similar pre-apical and incomplete median fasciae ; knee blackish on 
both sides. Hind tibia blackish, with light greyish basal ring. 



I 




Fig. 37. Mananara fasciata sp. n. i, male, type. 2, sternum. 3, end of male 
abdomen, lateral view. 4, the same, from above. 5, end of female abdomen, lateral 
view. 6, phallic complex, with ectophallic membrane and epiphallus removed. 7, the 
same, with most part of cingulum removed. 8, epiphallus. 



I 



342 V. M. DIRSH 

$. Paratype. As the male, but larger. 

Length of body (J i8, $ 26 ; pronotum ^ 4-5, $ 6 ; elytron ^8, $ 11-2 ; hind femur (^ 12, 
$ 14 mm. 

Madagascar Est : Mananara, Nosy Mangabe, iv. 1958, i (^ type (E. Raharizonina) ; 
Ambohimakana, Manambato sur I'Anove, i $ (A. Robinson) ; He Sainte Marie, 
foret de Kalalao, iii.1960, i (^, 2 $ (A. Robinson). 

Type and para types in Paris Museum, i (^ and i $ paratypes in the British 
Museum (Natural History). 

PERINETA gen. n. 

Small or medium size and moderately robust. Integument rugose, shiny and sparsely hairy. 
Antenna filiform. Head slightly inflated, subconical, subglobular above ; fastigium of vertex 
narrowing forwards, with truncate apex, concave above ; frons oblique and slightly incurved ; 
frontal ridge low, slightly roundly protruding between antennae ; above and below ocellus 
sometimes with shallow depression. Eyes large, strongly convex ; interocular distance very 
narrow. Pronotum subcylindrical, widening backwards, without lateral carinae ; median 
carina forming two large, tubercle-like projections on prozona, obtuse on metazona ; three wide 
sulci crossing dorsum ; metazona less than half length of prozona, its posterior margin widely 
rounded, almost straight. Prosternal process widely conical. Mesosternal interspace wider 
than its length. Elytra lobiform, lateral, reaching third abdominal tergite. Tympanum 
present. Hind femur slender. External apical spine of hind tibia absent. Hind tarsus more 
than half length of tibia. Arolium large. Last abdominal tergite with slightly upcurved 
posterior margin. Supra-anal plate as wide as long, with excurved sides and slightly attenuate 
angular apex ; middle of apical half with pair of tubercles. Cercus compressed, narrowing 
towards strongly incurved apex. Subgenital plate, in profile, acutely conical ; from above, 
with squarely widened apex. Phallic organ robust. Basal and apical valves of penis connected 
by thin flexure. Apical valves of penis and valves of cingulum distally not covered by sheath. 
Arch of cingulum present, forming long posterior projections. Epiphallus bridge-shaped, with 
small angular ancorae and lobiform upcurved lophi. 

Type species : Perineta bicolor sp. n. 



Perineta bicolor sp. n. 

(Text-fig. 38) 

(J. Type. Interocular distance half width of antennae in middle. Occipital cannula 
present. Pronotum with low convexities on prozona ; lower margin of lateral lobes excurved 
in middle. Prosternal process with obtuse apex. Mesosternal interspace narrowing towards 
apex. Metasternal interspace inversely triangular. Apex of elytron rounded ; membrane 
thin, with reduced venation and reticulation. Supra-anal plate of male with shallow longi- 
tudinal sulcus in middle and upcurved sides. General coloration dark brown. Fastigium of 
vertex, frons in middle, gena under eye and labrum, with ochraceous spots. Pronotum with 
two pairs of wide ochraceous, longitudinal stripes interrupted in middle. Elytra pale ochraceous. 
Hind femur ochraceous, with three blackish, transverse fasciae on both sides ; hind knee brown, 
with ochraceous spot in middle of both sides. Hind tibia brown, with ochraceous ring on base 
and ochraceous spots on external side. Hind tarsus red. 

Length of body 14, pronotum 3-5, elytron 3-7, hind femur 9-2 mm. 

Madagascar Centre : Perinet, i (^ type (in Paris Museum). 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



343 



Subfamily CYRTACANTHACRIDINAE 

Key to Genera 

1 (2) Prosternal process slightly inclined backwards. Body comparatively slender 

(Text-fig. 39) RHADINACRIS (p. 344) 

2 (i) Prosternal process strongly curved backwards, almost touching mesosternum. 

Body comparatively robust. 

3 (4) Costal area of elytra with fine and dense, wavy, elongated reticulation 

NOMADACRIS (p. 348) 

4 (3) Costal area of elytra with usual net of reticulation. 

5 (6) Dorsum of pronotum puncture-like sculptured. Membrane of elytra transparent 

(Text-fig. 40) FINOTINA (p. 344) 

6 (5) Dorsum of pronotum smooth, with velvet-like texture. Membrane of elytra 

parchment-like CYRTACANTHACRIS (p. 346) 







Fig. 38. Perineta bicolor sp. n. i, male, type. 2, end of male abdomen, from above. 
3, phallic complex, with part of ectophallic membrane and epiphallus removed. 4, 
the same with most part of cingulum removed. 5, epiphallus. 



344 V. M. DIRSH 

RHADINACRIS Uvarov, 1923 

Large and slender. Integument moderately rugose and dotted. Antenna filiform, longer 
than head and pronotum together. Fastigium of vertex angular, with truncate apex, concave, 
with distinct marginal carinulae ; frons oblique, slightly excurved ; frontal ridge narrow, with 
parallel carinulae and shallow sulcus. Pronotum slightly saddle-shaped, constricted in middle ; 
three sulci crossing dorsum ; median carina linear obtuse, lateral carinae absent ; metazona 
about as long as prozona, its posterior margin widely obtusangular, almost rounded. Prosternal 
process subconical, slightly inclined backwards, with obtuse apex. Mesosternal interspace 
elongated, narrowing behind. Elytra long, narrow, much exceeding end of abdomen. Hind 
femur slender. Lower lobes of hind knee rounded. Arolium large. Last abdominal tergite of 
male with pair of small, widely rounded projections. Supra-anal plate elongate-angular, with 
attenuate apex. Cercus compressed, widely angular with acute apex. Subgenital plate 
elongate, acutely conical. Phallic complex robust ; apodemes short ; rami large ; basal and 
apical valves of penis connected by thick flexure ; apical valves of penis simple, narrow, 
acutangular ; valves of cingulum narrow ; arch of cingulum present ; gonopore process long. 
Epiphallus bridge-shaped, with small, rounded ancorae and large, lobiform, strongly upcurved 
lophi. Ovipositor short, with robust valves, slightly curved at apices. Subgenital plate with 
angular apex. 

Type species : Acridium schistocercoides Brancsik, 1893. 

Rhadinacris schistocercoides (Brancsik, 1893) 
(Text-fig. 39) 

Acridium schistocercoides Brancsik, 1893 : 192. 

Rhadinacris schistocercoides (Brancsik, 1893) Uvarov, 1923 : 485. 

cJ. Large and very slender. Antenna twenty-five-segmented. General coloration brownish. 
Head above with ochraceous longitudinal stripe. Median carina of pronotum ochraceous ; 
lateral lobe of pronotum with olive-green curved, longitudinal stripe, lower part with oval 
ochraceous spot. Hind wing light blue. Hind femur light brownish ; upper side with two 
transverse fasciae ; external carinulae with small brownish spots ; hind tibia slightly purplish ; 
spines yellowish with dark brown apices. 

$. As male, but larger. 

Length of body ^ 33-5-39. ? 46-48 ; pronotum ^ 6-6-3, ? 8-8-9 ; elytron ^ 32-5-38, ? 44" 
52-5 ; hind femur ^ 18-19, $ 24-27-5 mm. 

Madagascar Centre : reserve naturelle de Zakamena, i $ ; Tananarive, Tsimba- 
zaza, xii.1947, I $ (R. Paulian) ; Moramanga, Anjiro, v. 1948, i $ (A. Robinson) ; 
Fianarantsoa, xi.1957, i $. 

Madagascar Sud : lac Tsimanampetsotsa, v. 1957, i ^ (R. Paulian) ; Itampolo, 
V.1957, 2 <3 (R. Paulian) ; Tranoroa, xii.1951 (R. Paulian) ; Beloha, 100 m., 
vi . 1957 (A. Robinson) ; Ambovombe, Amboasary, iii . i960, i c^ ; Betroka, i c^ 
(R. Paulian) ; La Sakoa (R. P. Cattala). 

FINOTINA Uvarov, 1924 

Large, robust. Integument dotted. Antenna about as long as head and pronotum together. 
Fastigium of vertex trapezoidal, depressed in middle ; frontal ridge narrow, depressed at and 
below ocellus ; interocular distance wider than frontal ridge. Pronotum tectiform, with obtuse 
median carina crossed by three deep sulci. Metazona shorter than prozona, with angular 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



345 



posterior margin. Prosternal process strongly curved backwards, almost touching mesosternum, 
compressed at base, inflated in middle and obtusely conical at apex. Elytra exceeding end of 
abdomen, with transparent membrane and comparatively sparse reticulation, apex rounded. 
Male supra-anal plate trilobate, with small lateral and large angular apical lobe. Cercus 



I 




Fig. 39. RhadinacHs schistocercoides (Brancsik, 1893). i, male. 2, phallic complex, 
with endophallic membrane and epiphallus removed. 3, the same, with most part of 
cingulum removed. 4, epiphallus, lophi in vertical position. 5, the same, lophi in 
horizontal position. 6, end of male abdomen, lateral view. 



346 V. M. DIRSH 

strongly compressed, widely angular, with acute apex. Subgenital plate elongated, acutely 
conical. Phallic complex robust. Rami of cingulum large. Sheath of penis absent. Basal 
and apical valves of penis connected by thick flexure. Apical valves of penis strongly elongated, 
with bilobate apex. Valves of cingulum absent. Arch of cingulum large. Epiphallus robust, 
with small ancorae and large elongate angular lophi. Ovipositor short, robust, with valves 
curved at apices. Subgenital plate with angular apex. 

Type species : Acridium ranavaloae Finot, 1907. 

Finotina radatna (Brancsik, 1893) 
(Text-fig. 40) 

Acridium radama Brancsik, 1893 : 191. 

Finotina radama (Brancsik, 1893) Uvarov, 1924 : 104. 

(J. Large. Integument finely rugose and dotted. Median carina of pronotum sharp, 
linear. General coloration light green, without pattern. Upper external cannula of hind 
femur sometimes reddish. Hind knee sometimes with reddish crescentic spot. Hind tibia 
green, with spines reddish at apices. 

$. As the male, but larger. 

Length of body ^ 38-42, $ 43-57 ; pronotum ^ 9-10, $ 11-15 ; elytron ^ 35"5-36, ? 41- 
56-5 mm. 

Madagascar Nord : Montagne d'Ambre, Les Roussettes, xii. 1948, i $ (R. Paulian). 

Madagascar Centre: Tananarive, Tsimbazaza, ii.1948, i $; iv.1948, 4 $ (R. 
Paulian) ; foret de Didy, Andranomandevy, 1,039 ^^•> x.1936, i $ (P. Griveaud). 

Madagascar Est : Perinet iv.1948, i (^, i $. 

Madagascar Quest : foret de I'Ankarafantsika, Ampijoroa, i $. Type locality : 
Nosy Be. 

Finotina ranavaloae (Finot, 1907) 

Acridium ranavaloae Finot, 1907 : 257. 

Finotina ranavaloae (Finot, 1907) Uvarov, 1924 : 105. 

Differs from Finotina radama by less sharp median carina of pronotum, more 
slender body and general coloration, which is brownish with green stripe along 
head, pronotum and dorsal part of folded elytra. Lower lobe of hind knee greenish ; 
crescentic spot blackish ; hind tibia reddish, tibial spines whitish, brown at apices. 

Length of body (^ 35-36, $ 46-52 ; pronotum ^ 9--9.5, ? 12-13 ; elytron (^ 31- 
38, $ 43-48 ; hind femur ^ 20-24, $ 26-27 . 5 mm. 

Madagascar Est: Sambava, Marojejy, Ambinanitelo, 500 m., xii. 1958, i $ 
(E. Raharizonina) . 

Madagascar Quest : foret de I'Ankarafantsika, i.1957, i ? (E. Razafimandimby). 

Madagascar Sud Quest: Sakaraha, Lambomakandro, i v. i960, i $. Type 
locality : He Sainte Marie. 

CYRTACANTHACRIS Walker, 1870 
Cyrtacanthacris tatarica tatarica (Linnaeus, 1758) 

Gryllus Locusia tataricus Linnaeus, 1758 : 432. 

Gryllus Locusta ranaceus Stoll, 1813 : 30 ; Uvarov, 1923 : 39. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 



347 



Cyrtacanthacris concisa Walker, 1870 : 560 ; Uvarov, 1924 : 97. 
Cyrtacanthacris subliturata Walker, 1870 : 565 ; Uvarov, 1924 : 97. 
Cyrtacanthacris internexa Walker, 1870 : 613 ; Uvarov, 1924 : 97. 
Cyrtacanthacris compta Walker, 1870 : 613 ; Uvarov, 1924 : 97. 

Madagascar Est : Maroantsetre, Ambodivoangy, i $ (J. Vadon) ; Mahanoro, 
I ? (A. Molet). 

Madagascar Quest : ForM de I'Ankarafantsika, Ampijoroa, i $ ; reserve naturelle 
de Namoroka, i $ (R, Paulian). 

Madagascar Sud et Sud Quest: Ankazoabo, le Banian, 70 m., vii.1957, i (^ 
(P. Griveaud) ; Morombe, lac lotry, 40 m., 8. vii.1957 (P. Griveaud) ; La Sakoa 




Apd — V- 





Ejj Cer 



Fig. 40. Finotina radama (Brancsik, 1893). i, female. 2, phallic complex, with 
ectophallic membrane and epiphallus removed. 3, the same, with lateral sclerotization 
of ectophallic membrane removed. 4, the same, with most part of cingulum removed. 
5, epiphallus. 



348 V. M. DIRSH 

I $; Beloha, lOO m., 27. vi. 1957, i 9 (P- Griveaud) ; Tsivory, 15.viii.1948, i $ 
(R. Paulin) ; lac Tsimanampetsotsa, v.1951, i $ (R. Paulian). He Europa, i.ii. 
1950, I 9 (R. Paulian). 

NOMADACRIS Uvarov, 1923 
Nomadacris septemfasciata (Serville, 1838) 

Acridium septemfasciatum Serville, 1838 : 661. 
Acridium coangustatum Lucas, 1862 : 24 ; Uvarov, 1923 : 37. 
Cyrtacanthacris fascifera Walker, 1870 : 558 ; Uvarov, 1923 : 37. 
Cyriacanthacris purpurifera Walker, 1870 : 561 ; Kirby, 1910 : 448. 
Cyrtacanthacris subsellata Walker, 1870 : 559 ; Uvarov, 1923 : 37. 
Acridium sanctae-mariae Finot, 1907 : 349 ; Uvarov, 1923 : 37. 
Nomadacris septemfasciata (Serville, 1838) ; Uvarov, 1924 : 7. 

Fort-Dauphin, Post Adm. Tsivory, Andobolava, xi.1959, i (^, 4 $ (Randrimasy). 
Montague d'Ambre, Lac Mandrit, Diego-Suarez, ix,i957, i (^ (J. Elie). Tananarive, 
Tsimbazaza, 7.1.1948, i (^. Manandrotsy, Betroka, i $ (J. Elie). 



REFERENCES 

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1902. Les Orthopteres de St. Joseph's College k Trichinopoly (Sud de I'lnde). 3me partie. 

Ann. Soc. ent. Fr. 70 : 580-635, pl. 9. 
1906. Rectificaciones y observaciones ortopterologicas. Bol. Soc. esp. Hist. nat. 6 : 390- 

393- 

1908. £tude sur quelques Acridiens d'Afrique. Bull. Soc. ent. Fr. pp. 242-248. 

1913. Rectificacion sinonimica. Bol. Soc. esp. Hist. nat. 13 : 312. 

1914. Estudios entomogicos, Segunda parte. Trab. Mus. Cienc. nat., Madr. (Ser. zool.), 

No. 20, no pp. 
Bolivar, C. 1932. Estudio de un nuevo Acridido de Madagascar del grupo Cranae. Eos, 

Madr. 8 : 391-396, 3 figs. 
Brancsik, K. 1893. Orthoptera nova africana. Jh. naturw. Ver. (Mus.Ver.) Trencsin, 15- 

16 : 175-200, pis. 4, 9, 10, II, 12. 
Bruner, L. 1910. Acridoidea from Madagascar, Comoro Islands and eastern Africa. In 

Voeltzkow, A., Raise in Ostafrika in den Jahren 1903-1905, 2 : 623-644, pl. 27. Stuttgart. 
Brunner von Wattenwyl, C. 1893. Revision du systeme des Orthopteres et description 

des esp^ces rapport^es par M. Leonardo Fea de Birmanie. Ann. Mus. Stor. nat. Genova, 

(2), 13 : 1-230, pis. 1-6. 
Carl, J. 1916. Acridides nouveaux ou peu connus du Museum de Geneve. Rev. suisse Zool. 

24 (6) : 461-518, pl. 2. 
Chopard, L. 1919. Diagnoses d'especes nouvelles d'Orthopteres. Bull. Soc. ent. Fr. 1919 : 

153-154- 
1952. Notes sur les Orthopteroides de Madagascar. IV. Faune de la foret de mouses du 

Tsaratanana. Mem. Inst. sci. Madagascar, E, 1 : 463-516, 34 figs. 
DiRSH, V. M. 1956. The phallic complex in Acridoidea (Orthoptera) in relation to taxonomy. 

Trans. R. ent. Soc. Lond. 108 : 223-356, 66 pl. 

1958. Acridological notes. Tijdschr. v. Entom. 101, i : 51-63, f. 1-34. 

1958. Synonymic and systematic notes on African Acridoidea. Rev. Ent. Mozambique, 

1 (2) : 239-244. 



THE ACRIDOIDEA (ORTHOPTERA) OF MADAGASCAR 349 

DiRSH, V. M. & UvAROv, B. P. 1953- Preliminary diagnoses of new genera and new synonymy 

in Acrididae. Tijdschr. Ent. 96 (3) : 231-237. 
FiEBER, F. X. 1853. Synopsis der europaischen Orthopteren. Lotos, 3 : 90-104, 115-129, 

138-154, 168-176, 184-188, 201-207, 232-238, 252-258. 
FiNOT, A. 1903. Sur le genre Enthymia, de la tribu des Acrididae, famille des Acridoidea, 

ordre des insectes Orthopteres. Ann. Sac. ent. Fr. 71 : 620-636. 
— — 1907. Sur le genre Acridium. Ibid. 76 : 247-354, 27 figs. 

1908. Acridien nouveau de Madagascar. Ibid. 77 : 1-6, pi. i. 

Johnston, H. B. 1956. Annotated catalogue of African grasshoppers. Cambridge Univ. 

Press. : 1-833. 
Karny, H. 1907. Ergebnisse der mit Subvention aus der Erbschaft Treitl unternommenen 

zoologischen Forschungreise Dr. Franz Werner's nach dem aegyptischen Sudan und Nord- 

Uganda, IX. Die Orthopterenfauna des Aegyptischen Sudans und von Nord-Uganda 

(Saltatoria, Gressoria, Dermaptera) mit besonderen Berucksichtigung der Acridoideen- 

gattung Catantops. S.B. Akad. Wiss. Wien, 116 : 267-378, 3 pis. 
Karsch, F. 1893. Die Insekten der Berglandschaft Adeli in Hinterlande von Togo. Berl. 

ent. Z. 38 : 1-266, pis. 1-6. 

1896. Neue Orthopteren aus dem tropischen Africa. Stettin, ent. Ztg. 57 : 242-359. 

KiRBV, W. F. 1910. A Synonymic Catalogue of the Orthoptera. Vol. 3. Orthoptera Saltatoria. 

Part II. Locustidae vel Acridiidae. London. 
Krauss, H. a. 1877. Orthopteren vom Senegal gesammelt von Dr. Franz Steindachner. S.B. 

Akad. Wiss. Wien, 76 (i) : 29-63, pis. i, 2. 
Linnaeus, C. 1758. Sy sterna Naturae, loth ed. 
Lucas, H. 1862. Orthopteres. In Maillard, L., Notes sur I'tle de la Reunion, Annexe i, 

pp. 22-25, pl- 21. Paris. 
Popov, G. 1950. Note on the genus Bibulus I. Bolivar, 1914 (Orthoptera, Acrididae). Proc. 

R. ent. Soc. Lond. B, 19 : 133-135, 15 figs. 
Ramme, W. 1929. Afrikanische Acrididae. Revisionen und Beschreibungen wenig bekannter 

und neuer Gattungen und Arten. Mitt. zool. Mus. Berl. 15 : 247-492, 16 pis., 106 figs. 

1931. Erganzungen und Berichtigungen zu meiner Arbeit " Afrikanische Acrididae ". 

Ibid. 16 (6) : 918-945, pi. xi, 17 figs. 

Rehn, J. A. G. 1938. A revision of the neotropical Euthymiae. Proc. Acad. nat. Sci. Philad. 

90 : 41-102, 7 pis., 43 figs. 
1944- A review of Old World Euthymiae (Orthoptera, Acrididae, Cyrtacanthacridinae). 

Proc. Acad. nat. Sci. Philad. 96 : 1-135, 8 pis., 91 figs. 
Saussure, H. de. 1899. Orthoptera. In Wissenschaftliche Ergebnisse der Reisen in 

Madagascar und Ostafrika in den Jahren 1889-95 von Dr. A. Voeltzkow. Abh. senckenb. 

naturf. Ges. 21 : 567-664, pis. 37, 38. 
Serville, J. G. a. 1 83 1. Revue m^thodique des insectes de I'ordre des Orthopteres. Ann. 

Sci. nat. (Zool.) 22 : 28-65, 134-162, 262-292. Separate loi pp. 
Dec. 1838. Histoire naturelle des Insectes. In Roret, Collection des Suites a Buff on. 

Orthopteres, 776 pp., 14 pis. Paris. 
SjosTEDT, Y. 1909. Wissenschaftliche Ergebnisse der schwedischen Zool. Exped. nach dem 

Kilimandjaro, dem Meru und den umgebenden Masaisteppen deutsch Ostafrikas, 1905-1906. 

17. Orthoptera. 7. Acridoidea: pp. 149-199, i pi., 2 figs. 

1918. Neue Orthopteren aus Afrika und Madagascar. Ark. Zool. 12 (i) : 18 pp., 5 figs. 

Stal, C. 1873. Orthoptera nova descripsit C. Stal. Ofvers. Vetensk Akad. Forh., Stockh. 

30 (4) : 39-53- 
1875. Observations Orthopt^rologiques. (i) K. Svenska Vetensk Akad. Handl. 3(14) : i- 

43- 

1878. Systema Acridiodeorum. Ibid. 5 (4) : i-ioo. 

Stoll, C. 1813. Representation exactement colorees d'apres nature des Spectres ou Phasmes, des 

Mantes, des Sauterelles, des Grillons, des Criquets et des Blattes qui se trouvent dans les quatre 

parties du monde. Acrididae, 42 pp., 23 pis. Amsterdam. 



350 V. M. DIRSH 

UvAROV, B. p. 1922. On some new or little known South African grasshoppers. Ann. Mag. 

nat. Hist. (9) 9 : 99-113, 4 figs. 
1923. Notes on locusts of economic importance with some new data on the periodicity of 

locust invasion. Bull. ent. Res. 14 : 31-39, i fig. 
1923. A revision of the Old World Cyrtacanthacrini. I. Ann. Mag. nat. Hist. (9) 11 : 130- 

144, figs. 1-7. II. (9) 11 : 473-490. III. (9) 12 : 345-366, fig. 8. 

1924. A revision of the Old World Cyrtacanthacrini. IV. Ibid. (9) 13 : 1-19, figs, g, 10. 

V. (9) 14 : 96-113, figs. II, 12. 

1943- Notes on the Old World genera of the group Euthymiae. Ibid. (11) 10 : 384-389. 

1944. The African genera allied to Leptacris Walker, 1870. Ibid. (11) 11 : 13-21. 

Walker, F. 1870. Catalogue of the Specimens of Dermaptera Saltatoria in the Collection of the 

British Museum, Part III, pp. 485-594 ; Part IV, pp. 605-801. 
WiLLEMSE, C. 1925. Revision der Gattung Oxya Serville. Tijdschr. Ent. 68 : 1-60, 65 figs. 

1928. Revision des Acridiodea d^scrites par De Haan, avec descriptions de nouvelles 

especes. Zool. Meded. 11 (i) : 1-27, 6 pis. 

1930. Fauna Sumatrensis. Preliminary revision of the Acrididae. Tijdschr. v. Ent. 

73 : 1-2 10, loi figs. 




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r I' fc • r ' * \y , 



NOTES ON TUNGA CAECIGENA 
(SIPHONAPTERA : TUNGIDAE) 

KARL JORDAN 

NEOTUNGA EULOIDEA GEN. N., SP. N. 
(SIPHONAPTERA : PULICIDAE) 

F. G. A. M. SMIT 




BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 12 No. 7 

LONDON : 1962 



NOTES ON TUNGA CAECIGENA 
(SIPHONAPTERA : TUNGIDAE) 



BY 



KARL JORDAN 



"H 



NEOTUNGA EULOIDEA GEN. N., SP. N. 
(SIPHONAPTERA : PULICIDAE) 



BY 



F. G. A. M. SMIT 

British Museum (Natural History) 



■"H. (f 




pp. 351-377 ; I Plate ; 30 Text-figures 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 12 No. 7 

LONDON: 1962 



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NOTES ON TUNGA CAECIGENA 
(SIPHONAPTERA : TUNGIDAE) 

By KARL JORDAN 

EDITOR'S NOTE 

A FEW years before his death Karl Jordan prepared the manuscript of a paper on a 
Chinese Jigger Flea, Tunga caecigena J. & R., the female of which lives embedded in 
the skin of its host. After describing the extreme specialization of this species to its 
environment and contrasting it with free-living fleas, Jordan postulated the existence 
of a species in which the degree of specialization was intermediate between the two 
extremes. Recently material of a jigger-like flea, a parasite of pangolins in Southern 
Rhodesia, was received by Mr. F. G. A. M. Smit, who realised that in its specialization 
to a sedentary mode of life it accords very closely with the condition postulated by 
Jordan. This species is described for the first time by Smit on p. 365 below. 

This striking fulfilment of Jordan's forecast is believed to justify the publication in 
this context of his manuscript which, though finished in draft, together with the 
relevant drawings, he never finally prepared for publication. Most of it, indeed, was 
left in his minute and barely legible handwriting, and but for Mr. Smit's skill and 
diligence in deciphering it, might never have seen the light of day. As it stands, the 
manuscript contains indications that Jordan would have wished to modify it, perhaps 
substantially, before pubUcation. There are, for example, passages in which the 
author discusses the possible causes of the adaptations he describes, speculative 
essays of a kind which rarely, if ever, occurred in his published work. While in fairness 
to the author such passages have been omitted, and some necessary alterations have 
been made to clarify the original text, everything possible has been done to preserve 
its factual content and the author's characteristic style. 

Mr. Smit has appended references to recent papers relating to the subject and has 
also made corrections to the figures where these were indicated in pencil by Jordan. 
The original manuscript and a transcript are in the Entomological Library, British 
Museum (Natural History). 

SYNOPSIS 

The modifications of intracutaneous fleas, as exemplified by a Chinese jigger, Tunga caecigena, 
are described. The existence of a flea showing a degree of speciaUzation intermediate between 
that of the Pulicidae and that of the Tungidae is postulated. 

Though the public is in general not very observant about the habits of such small 
creatures as insects, the flea has drawn attention to itself not only by the prick of its 
mouthparts, which is a mere fleabite, but particularly by the great nimbleness with 
which the tormentor evades capture. Agility varying in degree in different genera is 

ENTOM. 12, 7. 30§ 



354 KARL JORDAN 

indeed an almost general characteristic of the Siphonaptera, and whoever has 
tried to catch Pulex irritans will agree that this agility is a real advantage to the 
individual flea and therefore to the species. Nevertheless, in some 40 species not far 
removed in relationship from the human flea the female has given up this advantage 
and taken to a sedentary life, attaching herself to the host in a selected place until 
the end of her egg-producing existence. It is obvious that during the period of 
transition in the evolution from an agile flea to an immobile one the species had a 
bad time until new adaptions were perfected by natural selection, such as the 
strengthening of the exoskeleton or the evolution of the habit to settle in a place on 
the host where the parasite is comparatively free from destruction by tooth, beak or 
claw. New adaptations have more than made good the loss of the advantage of 
mobility, the sedentary fleas being so flourishing that at least some of them have 
become pests. 

It is to be expected that the great step from agility of the female to immobility is 
associated with modifications in the morphology. These modifications are consider- 
able, but in observing them we must bear in mind that the question of cause and 
effect implies opposite possibilities, either the change of habit in these fleas having 
been followed by morphological modifications or some profound morphological 
variation having caused a change of habit. 

There are two kinds of sedentary fleas : the extracutaneous stick-fast fleas which 
fasten themselves to the host by the laciniae and epipharynx (which together form 
the piercing-sucking tube), and the intracutaneous jiggers or sand-fleas which live in 
a cyst within the skin of the host. In both kinds the males retain mobility, but are 
also morphologically greatly modified. The jiggers may be called either the most 
advanced or the most degenerate of all fleas both in body and habits. Extremes far 
transcending the average are always instructive and for that reason I have selected 
a Chinese jigger as an illustration of the changes to which I have alluded. The exis- 
tence of this species was first mentioned by W. F. H. Blandford who, in 1894, gave 
an account of specimens sent to the Entomological Society of London from Ningpo 
(90 miles south of Shanghai). He rather doubted the flea to be identical with the 
common South American sand-flea {Tunga penetrans), described the expanded 
specimens and their situation in the ear of sewer-rats {Rattus norvegicus) and men- 
tioned some differences from Tunga penetrans. When we wrote a monograph of the 
Sarcopsyllidae (Jordan & Rothschild, 1906) we asked Blandford for the loan of the 
Chinese specimens, but they could not be found. However, in 1920 he came across 
a slide, which he presented to N. Charles Rothschild and from which we described 
the species Tunga caecigena (Jordan & Rothschild, 1921 : 131). The species was 
redescribed as Dermatophilus lagrangei Roubaud in 1925^ and in 1930 Wu described 
and figured the ear nodules and the flea contained therein. Of 250 rats obtained by 
Wu at Soochow 55 Rattus rattus and 13 Rattus norvegicus were infested ; the flea was 
identified as Tunga caecigena by Riley (1932). A number of years ago I had the 
pleasure of receiving from Dr. Teng Pin Hui, Director of the South Fukien Plague 
Station, the ears of 13 rats {Rattus rattus alexandrinus and R. norvegicus) preserved 
in formahn and aU containing T. caecigena ; the rats were collected in Futsing, South 

^ Referred to as a synonym of T. caecigena by Sharif (1930 : 32). — F. S. 



NOTES ON TUNGA CAECIGENA 355 

Fukien^. The consignment was most welcome, and I express here my sincere thanks 
to Dr. Hui and likewise to Dr. E. Landauer who had asked Dr. Hui to send the fleas 
to me. The total number of fleas in the 13 pairs of ears is nearly 190 (in various stages 
of postmetamorphic development), the highest number in one pair of ears being 31. 

This series of specimens in situ enables me to supplement the previous accounts by 
describing in detail the growth and morphology of the adult female^. The internal 
morphology has only partially been studied ; that task is easier to accomplish with 
fresh specimens. . . .* All 190 specimens had entered from the ear's edge (Fig. i). 
This restricted localization is interesting.^ It is known of some agile species that they 
are found most frequently on definite parts of the body, one species for example on 
the neck and head, another on the rump ; the South American Tunga caccata 
(Enderlein, 1901), like the Chinese jigger confined to the ear of rats, enters the hind 
surface and not the edge. The female of Tunga caecigena arrives at the edge of the 
ear where it burrows into the skin and an environment is created in which it can 
safely accomplish the purpose of its life, the production of hundreds of eggs, each 
one of which is almost exactly as long as an unexpanded flea. The flea has only the 
laciniae and the epipharynx as implements for digging a hole. As the cuticle at the 
edge of a rat's ear is fairly thick, the flea — being only one millimetre long (Text-fig. 3) 
— ^will have disappeared inside before the sensitive portion of the skin is reached 
(Text-fig. 2a]. ... ^ In the large cysts containing specimens with the frontal end 
of the swollen abdomen projecting forward as four large humps, the bottom of the 
cyst shows the impression of the four humps and of the hole into which the mouth- 
parts were inserted : a mould of the anterior end of the flea. The only connection 
with the outside is the hole by which the flea originally entered, the aperture growing 
larger with the flea, and the only part of the flea in contact with the atmosphere is 
the tail end which is but slightly below the surface of the skin of the host. 

Among the fleas found in ears sent by Dr. Hui are a number of specimens of 
different sizes and appearance. In many of the rats' ears the cysts are so closely 
packed that they interfere with each other, thus apparently affecting the growth of 

* No dates are mentioned on the labels. Tunga caecigena is known from the following localities in 
China : Ningpo (Chekiang) (Blandford, 1894 : 229), Shanghai (Roubaud, 1925 : 401), Dong Chia Hong, 
nr. Soochow (Kiangsu) (Wu, 1930 : 59), Wu Tung Chiao (Szechuan) (Li & Chin, 1957 • 120), Foochow 
(Fukien) (Yang, 1955 : 287), Futsing (Fukien). Dr. K. Sakaguti informs me that the species was intro- 
duced into Japan probably about 1940 and that it has been reported from Osaka (Morishita & Arai, 1949) 
and Nishinomiya, nr. K6be (Tanaka & Ikuzawa, 1951). Yang (1955 : 293) records Rattus norvegicus 
and Rattus rattus as the most important hosts, and also found Tunga caecigena on Mus musculus, Suncus 
murinus and an undetermined wild rat. He states that adult rats and shrews are more heavily infested 
than the young, and the difference in the infestation rate of the two sexes is statistically insignificant. 
The fleas are probably univoltine and are usually found in the cold season, the optimum temperature 
zone being between 50° and 60° F. The closely related Tunga callida Li & Chin, 1957, also, has been 
collected only during the period November to March. — F. S. 

* The male Tunga caecigena has only recently been described (Chen & Ku, 1958). — F. S. 

* In the passage here omitted the author suggests that the mode of copulation in T. caecigena is probably 
similar to that of T. penetrans, which has since been described by Geigy & Suter (i960 ; 207, 208). — Ed. 

* Yang (1955 • 293) states : " The parasites are generally found embedded at the edge of the rats' ears, 
but they may attack the dorsal surface of the hosts' ears." Yang's reference that " Once it was also found 
at the base of the tail of a specimen of Rattus rattus " may refer to the closely related Tunga callida Li & 
Chin, 1957 (from Yunnan), the females of which bury themselves at the rear end of the host, especially 
around the anus. — F. S. 

* The author discusses the formation of the cyst (Text-figs. 26, c) and surmises that " the expanding 
fiea moulds the cyst and not the cyst the flea ",— Ed, 



356 



KARL JORDAN 






Figs. 1-3. Tunga caecigena J. & R. i. Part of ear of rat with four embedded fleas in 
various stages of postmetamorphic growth. 2. Cross-section of skin of rat's ear, showing 
embedded fleas in (a) the first stage of postmetamorphic growth, in {b) an intermediate 
stage and in (c) the final stage. 3. Whole female, of. Fig. 2a. 



NOTES ON TUNGA CAECIGENA 357 

some specimens. Though we have no proof that each individual of this flea goes 
through all the stages observed before it reaches its final shape and size, the evidence 
points in that direction. Text-figs. 4-10 represent these stages. The exoskeleton is 
almost intact in Text-fig. 4 ; in Text-fig. 5 the expansion of the abdomen has begun ; 
in Text-fig. 6 the membrane between terga I and IV is greatly extended and at 
segments II and III there is a constriction ; in Text-fig. 7 the anterior swelling 
between segments I and II already conceals part of the thorax and the second con- 
striction has shifted backwards ;' in Text-fig. 8 a small swelling appears all round the 
thorax, a sort of cushion on which lie the thorax and head ; Text-fig. 9 shows a fully 
distended female in which the anterior swellings, two each side, completely conceal 
the head and thorax in lateral aspect^ but they are visible in a view from the front 
or slightly askew (Text-lig. 10).^ 

I have taken about half the number of specimens out of their cysts, and left the 
others in situ. Most specimens have attained full size or nearly, but about 20 are 
smaU and look to me as if they represent the first stages in the individual growth from 
the non-expanded female to the fully expanded one. The smallish specimens vary 
a good deal in shape and this may partially be due to overcrowding. 

The exoskeleton of an adult insect, being a finished product, does not grow any 
more and is not affected, apart from degeneration, by the expansion of the volume 
of the body necessitated in many female insects by the development of the great 
number of comparatively large eggs in the abdomen. Head, thorax and their append- 
ages are the same size in the unexpanded jigger female (Text-figs. 2a, 3) as in the fully 
expanded specimen of over 8 mm. (Text-fig. 2c). However, the hfe in the cyst has a 
remarkable effect on the legs in jigger fleas, the tarsi, tibiae and femora disappearing 
partially or completely, the process beginning with the hind tarsi (cf. Wagner, 
1932 : 249). In the series of some 90 specimens of the Chinese jigger only a few have 
one complete tarsus, and not a single one has a complete hind tarsus. ^° Only the first 
segment is preserved in the hind leg of one specimen and this exhibits strong signs of 
degeneration (Text-fig. 3).^^ . . . ^^ This process seems to set in soon after the skin of 
the host is entered, for the unexpanded and only slightly expanded specimens 
(Text-figs. 4-6) have already lost all tarsi. The small degree of expansion in these 
females, however, may be due at least partially to overcrowding. 

The enlargement of the abdominal cavity is made possible by the postmetamorphic 
growth of the intersegmental membranes as in all adult insects with secondarily 
enlarged abdomen, such as the queen termite, the oil-beetle {Meloe), and repletes of 
honey ants. But the growth of the jigger female differs essentially in the abdomen 
being transformed into a different shape, affecting the function of the segments, 

' This figure closely resembles that given by Yang (1955 : fig. i). — ^F. S. 

* A slightly different version of this drawing was published by Jordan (1948 : fig. 1290). — F. S. 

* In the closely related Tunga callida the gravid females are spherical, but they expand much more on 
the dorsal side than on the ventral, while the segments posterior to the fifth are elevated to form a 
characteristic button-like anal projection (Li & Chin, 1957 • 120). — F. S. 

1" The hind tarsus of male Tunga caecigena is figured by Chen & Ku (1958 : figs, i, 8). — F. S. 

^^ Some species lose parts of their legs more readily than others, e.g. in embedded females of Tunga 
penetrans the legs are usually all intact, even in the final stage of expansion. Are parts of the legs lost in 
certain species during the act of burrowing into the skin ? If those parts are shed after this process one 
would expect to find remnants in the cyst. — F. S. 

12 The author suggests possible reasons to account for the loss of parts of the legs. — Ed. 

ENTOM. 12, 7. 3o§§ 



358 



KARL JORDAN 






Figs. 4-10. Tunga caecigena J. & R. in various stages of postmetamorphic development; 
explanations in text. (Actual size of specimen shown in Fig. 4 : i mm.; in Fig. 5 : 
2 mm.; in Fig. 6 : 2-6 mm.; in Fig. 7 : 3-4 mm.; in Fig. 8 : 5 mm. Figure 8 drawn 
to a somewhat smaller scale than Figs. 4-7 ; Figures 9 and 10 drawn to a much smaller 
scale.) 



NOTES ON TUNGA CAECIGENA 359 

segments I, II and III becoming merely an expanded container and IV-X being 
consolidated into a tough truncate cone and serving as the only connection of the 
parasite with the outside world. It is a teleomorphosis of form affecting function. 

As will be evident from a comparison of Text-figs. 5-7, the intersegmental membrane 
between tergites I to III and between thorax and sternites II and III (there is no 
stemite I in fleas) are the first to grow longer and expand all round, tergite I remaining 
attached to the thorax. In Text-fig. 7 tergites II and III have travelled backwards, 
and a tough belt is formed in front of the two sclerites of segment IV, the specimen 
being here constricted and remaining so ; the anterior part of the abdomen has 
swollen forward. In Text-fig. 8 the head and thorax are on a cushion-like swelling, 
which later on becomes smaller — the specimen is still constricted at segment II. 
Further expansion breaks the constriction at segment II and the swelling extends far 
beyond the head, forming anteriorly on each side two dome-shaped projections, one 
tergopleural, the other sterno-pleural (Text-fig. 9) ; the slits between these four 
humps form an erect cross (Text-fig. 10) ; in a frontal view the head and thorax 
are seen deep down in the centre of the cross. The four humps are not always of 
equal width and length, but are always present in expanded specimens, being a 
specific distinction of the Chinese jigger.^* Among the American species, Tunga 
ierasma Jordan, 1937, likewise has four humps, which, however, are directed side- 
ways, somewhat in the manner of the arms of an X, the head and thorax not being 
concealed. 

After the formation of the belt (Text-fig. 7) in front of the sclerites of segment IV, 
the anal cone undergoes very little change. The tergal plates are well separated from 
the sternal ones and their dorsal and ventral parts, respectively, often show stronger 
chitinization. The cone is firmer than the enlarged part of the abdomen, which is 
chiefly due to the large number of muscles which extend from the segments of the 
cone forward to the thorax. The cone varies slightly in length, being shorter in some 
specimens than in others. In a view from behind the cone is occasionally surrounded 
by a brown ring formed by the faeces (remnants of the blood of the host) as in females 
of other species of Tunga. The exposed area encircled by this brown ring appears 
often to be lower than the margin of the aperture in the host's skin and has erroneously 
been termed a cloaca by various authors. As in all fleas, the rectum and the oviduct 
have separate orifices, the anus being situated between the tergite and the stemite of 
segment X and the oviduct opening between sternites VIII and IX. 

The majority of female Tunginae have no distinct line of demarcation between 
tergites IX and X, the two segments being fused. In a lateral view (Text-fig. 12) 
the triangular sclerite projecting backwards from the sensilium represents the tergum 
of segment X and the lateral short projection with a long apical bristle the tergo- 
pleurite homologous to the stylet present in the large majority of fleas ; below this 
pleurite there is a separate narrow sclerite bearing some bristles which is stemite X. 
In a view vertically on to the terminal surface of the cone (Text-fig. 13) sternum X 
appears as a narrow transverse bar bearing right and left a pair of bristles ; the apex 
of the combined tergites IX and X is deeply sinuate and the backward projection 
each side of the sinus is much foreshortened ; the anus is situated above sternum X. 

*' See footnote 9. 



36o KARL JORDAN 

The midgut of swollen and large specimens is folded twice, thus being divided into 
three parts as shown in Text-fig. ii.^* 

A problem which has remained obscure to me concerns the means of transporting 
the eggs from all parts of the large abdomen to the oviduct. Schimkewitsch (1884 : 
673) found only two longitudinal muscles in Tunga penetrans. ^^ Though the number 
of muscles is much larger in the Chinese jigger, I do not see how, in fleas which are 
often placed in the ear with the head downwards, the longitudinal muscles can guide 
the eggs from the anterior humps upward to the anal cone without the co-ordinated 
help of the transverse or ring muscles. ^^ However it may be accompUshed, eggs do 
arrive in the oviduct and are ejected. As the transverse diameter of the ripe egg is 
much greater than the diameter of the oviduct, the passage of the eggs is only possible 
because the structure of the anal cone admits considerable expansion in width. The 
two segments chiefly concerned in the process of egg-ejection are the tergite and ster- 
nite of segment VIII and the stemite of segment IX. Tergite VIII is divided dorsally 
in the median line into a right and left sclerite, which extend downward to or near 
to the ventral surface of the abdomen. Other tergites are not halved (the apical 
margin of tergite VII is in some specimens shghtly or more deeply incised medially). 
From the spiracle downwards tergite VIII widens considerably and in unexpanded 
specimens the lower areas of the right and left halves approach one another ; on 
the outer surface of the lower area there are three or four bristles and on the inner 
surface about half a dozen directed inward-distad (Text-fig. 13 ; the two halves are 
drawn as they appear in the specimen, the left half being more flattened and expanded 
in the preparation than the right half ; Text-fig. 12, lateral aspect). Sternite VIII 
lies in between the two halves of tergite VIII ; it differs much from the corresponding 
segment of the American jiggers^'' by being soft and tough, and ends in a pointed 
sclerotized cone. Below and in front of it hes stemite VII (Text-figs. 12, 13), which 
is divided medially and bears right and left two bristles, sometimes one. At the side 
of stemite VIII are two humps, flexible like the main body of st. VIII, and above the 
sternum is the orifice of the oviduct. The upper wall of the terminal part of the oviduct 
is formed by a large sclerite homologous to sternum IX and this is a special feature of 
the species of Tunginae. This sclerite extends from the ventral margin of tergite IX 
downward, forming a kind of half cyhnder resembling an elastic plate or a piece of 
paper of which two opposite margins are bent towards each other. The cavity thus 
formed is closed on the frontal side by the intersegmental membrane connecting 
sternum IX with X. The pressure exercised on the egg by the muscles of the abdomen 
will push the egg to the end of the oviduct where it is held between sterna IX and 
VIII and the stiff bristles on the inside of the right and left halves of tergite VIII. 
The egg pressing against the convex surface of the elastic sternum IX, bends this 

1* In fully distended specimens of Tunga penetrans the midgut is folded four times (Geigy & Herbig, 
1949) —F. S. 

1* Geigy & Herbig (1949) found, apart from the hemispherical muscles, only one central muscle cord in 
the abdomen of inflated specimens of Tunga penetrans. — F. S. 

1^ It would seem that the pressure inside an ovariole, caused by the process of egg-growth, is sufficient 
to push the eggs forward till they reach the oviduct, whence the exertion of muscular power will be 
necessary only for the ejection of the eggs. — F. S. 

^' In female Tunga penetrans sternum VIII appears to be entirely membranous ; I have not been able 
to detect it in any mounted specimens. — F. S. 



NOTES ON TUNGA CAECIGENA 



36 r 






12 

Figs. 11-13. Tunga caecigena J. & R. 11. Flea in final stage of expansion, showing 
alimentary canal and a full-sized egg in upper left part of abdomen. 12. Terminal abdo- 
minal segments of female in lateral view. 13. Same segments as viewed from behind. 



362 



KARL JORDAN 



surface upward (there is an egg in this position in one of the specimens of this flea) 
and now a further effort of the muscles would be assisted by the elastic force of 
sternum IX and of the bristles of tergum VIII and achieve the ejection of the egg. 

The third function of the anal cone is the provision of air for the greatly enlarged 
abdominal tracheal system by means of enlarged spiracles. Spiracles, as part of the 
exoskeleton, cannot be enlarged by postmetamorphic growth of the individual ; 
their enlargement can only be due to the result of the evolution of the species. In the 
process of postmetamorphic growth, however, shifting of the segments bearing the 
enlarged spiracles has kept the posterior four of them (on segments V to VIII) in 
contact with the atmosphere, whereas the anterior series are in the cavity and there- 



o o o o o 

o o o o o 

o o o o o 

o o o o o 



PULEX IRRITANS cf 

echidnophaga lar i na (f 
Hectopsylla PSITTACI cT 
TUNGA PENETRANS cf 



#00000 
^00000 

(}( J \ ) \ ) ( } ^^ Hectopsylla psittaci $ 



PULEX IRRITANS ^ 
ECHIDNOPHAGA LARINA § 





Fig. 14. 



TUNGA PENETRANS 



Abdominal spiracular fossae of metepimeron and terga I-VII of four species of 

Pulicid fieas. 



fore shut off from the atmosphere. The three spiracles of segments II to IV have 
hitherto been considered absent. If they were as minute in the Chinese jigger as they 
are in Tunga penetrans^^ I should very Hkely not have discovered them. Nature has 
provided two guides with the help of which the vestiges of these spiracles can be found, 
provided the process of clearing the specimens for the balsam preparation has not 
altogether destroyed the trachea of which the spiracle is the aperture. The guides 
alluded to are a bristle and a pair of minute dots [leucodiscs] . The abdominal terga 
I-VII I of the jiggers, inclusive of the male of T. penetrans, each has one subdorsal 
bristle. That this bristle is present on segment VIII is contrary to what one would 
have expected, there being no bristle above spiracle VIII in any of the fairly numerous 

1* I have failed to detect vestiges of spiracular fossae on terga II-IV in mounted female specimens of 
Tunga penetrans. — F. S. 



NOTES ON TUNGA CAECIGENA 363 

Species of the group of Siphonaptera to which Tunga must be assigned, whereas one 
or more bristles are present above the stigma of VIII in many species of other groups 
of fleas with which Tunga is much more distantly related than for instance with 
Hectopsylla. Nor can I look upon that bristle as a character which was lost in the 
ancestors of the Tunginae and has reappeared. The development of the bristle on 
tergite VIII is most probably due to isomerism, a detail repeated on every segment. 
However that may be, a little more downward and forward from the bristle there are 
two transparent dots, sensory cups, and below these we find the vestigial spiracle 
(Text-fig. 3). The somewhat larger size of the remnants in the Chinese jigger may be 
taken as evidence that there have existed — and possibly there still exist — species in 
which these anterior abdominal spiracles approached the size of the first abdominal 
one (on the metepimeron).^^ That is to say the reduction of spiracles II-IV and the 
compensating enlargement of spiracles V-VII took place gradually or step by step. 
Is there any other evidence bearing on this suggestion ? 

A comparison of the male spiracles of T. penetrans proves that this process in their 
evolution was confined to the female (Text-fig. 14) ; the series of rings, drawn to 
scale (spiracle of tergum VIII omitted), shows neither reduction in II-IV nor enlarge- 
ment in V-VII ; the first spiracle, on the metepimeron, is the same size as the 
following ones. 

In the male of Hectopsyllinae (Text-fig. 14) the proportional size of the spiracles is 
again the same as in male T. penetrans ; in all females, however, II-VII are larger 
than I and increase in size from II to VII. The Hectopsyllinae and the Tunginae are 
the only fleas in which the abdominal spiracle I of the female is much smaller than 
the spiracles of V-VII ; in aU other fleas spiracle I is as large as the following ones or 
larger. This evidence of the existence, in the female of the Tungidae, of a disposition 
toward the evolution of enlarged abdominal spiracles explains the exceptionally 
great size of spiracles V-VII in the jigger females. No other fleas show a correspond- 
ing trend towards the eUmination of spiracles II-IV. The degeneration of these 
spiracles, therefore, must be attributed to the loss of function. 

The Chinese jigger is in some respects of a more generaUzed type than the five 
American species of Tunga. It is therefore not far fetched to assume the existence of 
a jigger which only partially enters the skin of the host. We should expect to find 
in such a species the abdominal spiracle II in the process of degenerating. III and IV 
to be perhaps of the same size as in the male and V-VII more or less enlarged : a 
first step in the modification of the spiracles.^*' It may be left to the imagination 
of the reader to reconstruct the further steps to be passed before the evolution of the 
jigger females as we now know them. 

^* There is now one such species known ; see below, p. 375. — F. S. 
*• See footnote 19. 



364 KARL JORDAN 

REFERENCES 
Chen, C. & Ku, H., 1958, The discovery of a male specimen of the sand-flea, Tunga caecigena 

Jordan et Rothschild, 192 1, with a morphological description. Acta ent. sinica, 8 (2) : 

179-182, pis. I, II. 
Geigy, R. & Herbig, a., 1949, Die Hypertrophic der Organe beim Weibchen von Tunga penetrans . 

Acta. Trop. 6 (3) : 246-262, figs. 1-6. 

& SuTER, P., i960, Zur Copulation der Flohe. Rev. suisse Zool. 67 (2) : 206-210, i fig. 

Jordan, K., 1948, Suctoria in J. Smart — A Handbook for the identification of insects of medical 

importance. London, 2nd Ed.: 211-245, figs. 128-143. 
& Rothschild, N. C, 1906, A revision of the Sarcopsyllidae, a Family of Siphonaptera. 

Thomp.-Yates Lab. Rep. 7 (i) : 15-72, figs. A-G, pis. 1-4. 

& 1921, A new species of Sarcopsyllidae. Ectoparasites, 1 (3): 131-132, figs. 105-106. 

Li, K.-C. & Chin, T.-H., 1957, Tunga callida sp. nov., a new species of sand-flea from Yunnan. 

Acta ent. sinica, 7 (i) : 1 13-120, figs. 1-18. 
MoRiSHiTA, K. & Arai, T., 1949, On the rat chigoe hitherto unrecorded in Japan. Jap. J. sanit. 

Zool. 1 (2) : 37-40. 
Riley, W. A., 1932,^ The ear chigoe of rats in China, Tunga caecigena J. & R. Lingnan Sci. J., 

11 : 285-287. 
RouBAUD, E., 1925, Une nouvelle espece de puce-chique p6n6trante, parasite des rats en Chine : 

Dermatophilus lagrangei n. sp. Bull. Soc. Path, exot., 18 : 399-405, figs. 1-6, pi. i. 
ScHiMKEWiTSCH, W. 1 884. Zur Frage nach der Veranderung der Sarcopsylla penetrans unter 

dem Einfluss der Parasitismus. Zool. Am. 7 (183) : 673-676. 
Sharif, M., 1930, A revision of the Indian Siphonaptera, Part I : Pulicidae. Rec. Ind. Mus. 

32 (i) : 29-62, figs. 1-13. 
Tanaka, H. & Ikuzawa, M., 1951, Ecology study of the rat chigoe, I. Jap. J. sanit. Zool. 

2 (2-3) : 52-55. 
Wagner, J., 1932, Tunga bondari, eine neue Art der Sandflohe. Novit. zool., 38 : 248-249, figs. 

6-9- 
Wu, K., 1930, A study of the common rat and its parasites. Lingnan Sci. J., 9 (1-2) : 51-64, 

pl. 5. 
Yang, H.-S., 1955, Notes on the sandflea, Tunga caecigena Jordan & Rothschild, in Foochow. 
Acta ent. sinica, 5 (3) : 287-293, figs. 1-3. 



NEOTUNGA EULOIDEA GEN.N., SP. N. 
(SIPHONAPTERA : PULICIDAE) 

By F. G. A. M. SMIT 

SYNOPSIS 

A new species of flea, parasitic on pangolins, and showing both pulicid and tungid characters, 
is described. 

The pangolin Manis temmincki Smuts is widespread in southern and eastern Africa, 
but never common nor very easy to collect. In 1956, a female pangolin, captured in 
the Sinoia District of Southern Rhodesia, was brought to the University College of 
Rhodesia and Nyasaland at Salisbury, where it was kept alive for two years. After 
it had died, in August 1958, Dr. J. S. Weir of the Zoology Department examined the 
body and found a number of jigger-like fleas embedded in the skin of the underside 
of the animal. An appeal made by Dr. Weir to some members of the staff of the 
National Museum of Southern Rhodesia, at Bulawayo, for more fleas and other 
parasites of pangolins resulted in six additional specimens of the same flea collected 
by Mr. M. P. Stuart Irwin from a pangolin caught near Lonely Mine, Southern 
Rhodesia, in June i960. Dr. Weir sent three of these specimens to Dr. L. H. Finlay- 
son of the Department of Zoology, University of Birmingham, with the request to 
have this material identified. I am deeply indebted to Dr. Finlayson for having sent 
the specimens to the British Museum (Natural History), for these fleas have proved 
to be the most interesting and instructive new species which has ever come my way. 
According to recent classifications this flea belongs to one Family (or Subfamily) on 
the structure of head and thorax, and to another Family (or Subfamily) on most of 
the characteristics of the abdomen. 

I am most grateful to the gentlemen mentioned above for having made an impor- 
tant discovery and for the generosity with which they have placed all specimens 
at my disposal. 

Meanwhile Dr. Weir collected twelve more females from a pangolin captured near 
Que Que, also in Southern Rhodesia. All the fleas so far collected are females. The 
males, which are very likely minute and free-Uving, should be searched for in the 
nesting places of pangolins, which might be difficult to detect. I sincerely hope that 
Dr. Weir and Mr. Stuart Irwin will eventually discover the male specimens which 
they so kindly offered to search for, as the structure of the genitalia of the male would 
give valuable supporting evidence of the origin of sand-fleas. 

NEOTUNGA gen. n. 

Chaetotaxy reduced. Frontal margin of head angulate, with an upward pointing tubercle. 
Eye developed, sometimes with an internal sinus. First two segments of antennal clava lamellate, 
the other segments fused. Epipharynx with a number of minute tubercles along anterior margin ; 



366 



G. A. M. SMIT 



laciniae strongly serrated ; labial palp membranous. Maxillary stipes of median size, extending 
below genal margin. Thorax dorsally narrower than length of terguml. Prosternosome with 
epimeral flange ventrally drawn out into a triangular lobe ; mesosternal furca long and narrow. 
Metasternal ridge with a semi-distinct forward branch. Vincula absent. Oblique hyaline break 
of mid coxa complete. Hind coxa with a patch of spiniforms on inner side and with a ventro- 
anterior spur. Female : spiracular fossa of posterior terga larger than those of anterior terga ; 
two leucodiscs above spiracular fossae. Antesensilial setae absent. No seta above the large 
spiracular fossa of tergum VIII. Genital setae of tergum VIII well developed. Sternum IX 
membranous. Sensilium not divided dorsally, with eight trichobothria each side. Anal tergum 
not much reduced, with a short but stout stylet. Spermatheca rather large, with a dorsal orifice ; 
without a line of demarcation between bulga and hilla. 

Type of genus: Neoiunga euloidea sp. n. 

Neotunga euloidea sp. n. 

(Text-figs. 1-3, 6-12, 15, 16. PI. i) 

Holotype female and 10 female paratypes from Southern Rhodesia, Sinoia 
District : on Manis temmincki (adult $), viii.1958 (/. S. Weir) ; 6 female paratypes 
from Southern Rhodesia, Lonely Mine, 19° 30' S., 28° 42' E. : on Manis temmincki. 





Figs. 1-2. Neotunga euloidea sp. nov. i. Embedded female under the skin of Manis 
temmincki. 2. Expanded female removed from under the skin (the midgut is shown in 
dotted outline). 



NEOTUNGA EULOIDEA GEN. N.. SP. N. 367 

30. vi. i960, (M. P. Sttuirt Irwin) ; 12 female paratypes from Southern Rhodesia, 
15 miles S.E. of Que Que : on Manis temmincki (adult $), 21.x. i960 (/. S. Weir). 

Holotype and a number of paratypes in the British Museum (Tring), other para- 
types in the collection of Lt.-Col. R. Traub, Washington, D.C.; the Canadian National 
Collection of Insects, Ottawa ; the University CoUege of Rhodesia and Nyasaland, 
Salisbury ; the National Museum of Rhodesia and Nyasaland, Bulawayo ; the 
Antiplague Research Institute, Stavropol and the Schweizerisches Tropeninstitut, 
Basel. 

As no other species of Neotunga are known, the generic characters may be taken 
as diagnostic for the species. 

Head (Text-fig. 3). Frons angulate, with a distinct vertical tubercle ; dorsal margin of head 
about twice as long as the frontal margin below the tubercle. Wall of cranium fairly thick. Ventral 
margin of head with a broad rounded genal lobe. Preoral tuber not excessively large. Ocular 
seta very small, placed below level of eye ; genal seta slightly larger than the ocular one. Occipital 
region with two setae. A number of microsetae, most of which are pointing upwards, on frontal 
and dorso-occipital regions. Eye well developed, subglobular, but not very large, with or without 
an internal sinus. Antennal scape with one minute ventroposterior seta ; pedicel with about 
half a dozen longish setae which do not reach to the apex of the clava ; first two segments of 
clava lamellate, the other segments completely fused ; on outer surface of clava only a few 
minute setae. Anterior longitudinal half of antennal fossa covered by the flange-like extension 
of the postocular genal margin which thus covers the anterior half of the antenna. No setae 
bordering the antennal fossa dorso-posteriorly. One leucodisc, representing a placoid, on the 
preantennal region of head, three on the postantennal region. First segment of maxillary palp 
nearly twice as long as the second segment, third segment as long as second, fourth segment a 
little longer than the third ; the palp reaches nearly to the level of the apex of the fore coxa : 
stipes short, but still protruding well below the genal margin. Epipharynx fairly broad, with 
small projections over most of the length of the anterior margin ; lacinia very broad and with 
strong anterior and posterior serrations ; the two-segmented labial palp is membranous — these 
three mouth-parts are about i ^ times as long as the fore coxa. 

Thorax (Text-fig. 3). Thorax dorsally only about half as long as tergum I ; pronotum and 
mesonotum dorsally of the same length, metanotum nearly twice the length of either of the first 
two nota ; there is no fusion between the nota. Pronotum with four slender setae each side, 
mesonotum with two setae and metanotum with one subdorsal seta each side. Flange of proster- 
nosome with a triangular lobe pointing backward, mesosternosome with two setae. Metasternum 
without setae, metepimeron with a row of three slender setae. Pleural arch-joint of metathorax 
and the associated metanotal and pleural ridge well developed. 

Legs (Text-figs. 3, 6-8). Setae on outer surface of fore coxa not very numerous and most of 
them are small. Outer surface of mid coxa with an oblique hyaline break and a relatively small 
number of very short setae at and near the anterior margin, on inner side a few similar setae. 
Hind coxa with short setae scattered over the outer surface ; on the inner surface, apart from 
some thin setae, an oblique dense patch of short spiniforms. Ventro-anterior angle of hind coxa 
drawn out into a pointed triangular tooth. Outer internal rod and inner internal ridge of hind 
coxa not strongly developed. Femora with a number of small setae along the dorsal margin ; 
on inner surface with one lateral seta on proximal part of fore femur with two such setae on mid 
and hind femur ; all femora with one preapical lateral seta on inner surface. Tibiae with two 
notches in posterior margin each of which bearing two short setae the outer one of which is the 
stoutest ; without lateral setae ; ventro-anterior angle of tibiae with two setae, ventro-posterior 
angle with two setae on fore and mid tibia, three on hind tibia. 

Tarsi as shown in Text-figs. 6-8 ; note the reduced chaetotaxy, especially of the fifth tarsal 
segments which only bear one pair of thin sub-basal plantar setae, one pair of preapical plantar 
setae and the usual pair of preapical lateral setae ; claws basally with only a slight swelling. 



368 



F. G. 



M. SMIT 




$ 



Fig. 3. Neotunga euloidea sp. nov. Head, thorax, basal segments of legs, tergum I and 

sternum II of holotype. 



NEOTUNGA EULOIDEA GEN. N.. SP. N. 



369 





Figs. 4-5. Anterior part of females of : 4. Tunga penetrans (L.), 5. Echidnophaga 

gallinacea (Westwood). 



370 



F. G. A. M. SMIT 




6 7 8 

Figs. 6-8. Neotunga euloidea sp. nov. 6. Fore leg. 7. Mid leg. 8. Hind leg (all of paratypes). 



NEOTUNGA EULOIDEA GEN. N., SP. N. 371 

Abdomen (Text-figs. 2, 9-12, 15, 16). Terga I-VII each with a row of two medium-sized setae 
per side. Spiracular fossae of terga II-VII circular and large, increasing in size from tergum II 
onwards (the spiracular fossa on the metepimeron is somewhat larger than that of tergum II). 
Posterior margin of tergum VII dorsally sinuate. Sterna II-VI without setae, sternum VII with 
two setae near the ventral margin. Tergum VIII with about half a dozen setae on the ventral 
half and on the inner side with seven or eight strong and straight setae which are placed on a 
thickening of the wall of the sclerite ; no setae above the large and broad spiracular fossa. Lower 
half of posterior margin of tergum VIII with a small fold. Sternum VIII short and broad, without 
setae. Sensilium with eight trichobothria each side. Anal tergum with some stout dorsoapical 
setae below which a number of slender ones. Anal stylet nearly twice as long as broad in the middle 
with a long apical seta, two preapical ventral setae and two microsetae. Anal sternum short, 
with about half a dozen slender setae per side. 

Bursa copulatrix a simple elongate tube ; ductus obturatus present. Spermatheca with a 
rather thinly walled pyriform bulga, which is internally rather coarsely striated, and a rather 
short and narrow hilla ; orifice of spermatheca dorsal. As in other fleas the spermatheca is 
situated in the right hand side of the abdomen. 

Length : Distended females measure from 4 to 7 mm. in length, while the diameter of the 
abdomen is about 2 mm.; undistended females may be about i'5 mm. long. Eggs are o-y-o-S mm. 
long, 0'35-o*4 mm. broad. 

Embedded females of Neotunga euloidea were found in the soft skin, which is 
abnost completely hairless, on all ventral regions of the pangolin Manis temmincki ; 
in the pangolin from the Sinoia district there were sUght perianal and inguinal con- 
centrations but no such concentrations were observed in the specimen from Que Que. 
There is a tendency for the nodules containing the engorged fleas to be near the lateral 
margin of the venter at the edge of the scales ; the fleas are not found under the 
scales. The nodules are not orientated in any particular way and in one instance there 
were two engorged fleas exactly alongside each other, facing the same direction. 
Unlike the females of Tunga, the embedded flea Ues with its elongate body parallel 
with the skin's surface (Text-fig. i). Dr. Weir informs me that nodules are often 
difficult to see and can best be found by examining the skin for lumps, using gentle 
finger pressure. 

The degree of infestation was rather slight in the pangolins examined : 6, ii and 12 
fleas per host individual. Whether heavier infestations occur in nature remains to be 
discovered. It is interesting to note that the pangolin which was kept alive for two 
years was, on its death, found to be infested with 11 specimens of Neotunga euloidea. 
This might imply that the flea had been breeding in the cage where the pangolin was 
kept, for it seems unlikely that these fleas would remain alive and produce eggs for 
over two years. The pangolin from Que Que showed on the venter a small region of 
scar tissue with indications of former nodules. As considerable quantities of eggs are 
produced — the average number of ripe eggs in three specimens was 30 — it seems 
strange that the captive pangolin was not more heavily infested, but the conditions 
under which it was kept may not have been wholly favourable for the development 
of the larvae. To keep an infested pangolin in a flea-proof cage would seem the ideal 
method of eventually obtaining male and unengorged female fleas. 

Dr. Weir informed me of two negative records : one adult female Manis temmincki 
collected in December, 1959, in the Melsetter District of Southern Rhodesia (near the 
border of Mozambique), and one female collected on 19.xi.1960 between Salisbury 
and Norton. 



372 



F. G. A. M. SMIT 




Figs. 9-12. Neotunga eiiloidea sp. nov. (paratypes). 9. Anal tergum (stylet in dotted out- 
line). ID. Anal stylet. 11. Spermatheca. 12. Terminal abdominal segments [i.e. segments 
VI-X). 



NEOTUNGA EULOIDEA GEN. N., SP. N. 



373 



Nearly all fleas were cut out with surrounding skin and flesh and preserved in 
alcohol. After two or three hours in io% solution of potassium hydroxide the 
embedded fleas could be easily removed, with the aid of two minute mounted needles, 
from the macerated flesh around them, but one has to break the opaque thin silvery 
" skin " which fits tightly over part of the abdomen. 

An astonishing observation was made by Dr. Weir who found that " in some cases 
the flea seemed to be hanging free from the skin and was only attached by its head." 
Indeed, among specimens sent by Dr. Weir are a few without surrounding skin or 
flesh and without the silvery "skin" which is so characteristic of embedded specimens. 
Such specimens, which had no need to be treated with potash in order to free them, 
have been used for the photographs reproduced on PI. i. Mechanically it would 
seem to be impossible for an engorged specimen to penetrate the skin of the host. It 





14 

Figs. 13-14. Terminal abdominal segments of females of : 13. Echidnophaga gallinacea 
(Westwood). 14. Tunga penetrans (L.). 



374 



G. A. M. SMIT 



is more likely that these extracutaneous specimens failed to enter the skin and remained 
attached to it by the strongly developed mouthparts, as for example do females of 
Echidnophaga. If this is indeed the case it is interesting to note that these extra- 
cutaneous specimens become engorged to the same extent as embedded ones. This 
would indicate that engorgement is not correlated with embedding. The opposite 
appears to be the case in Tunga, for Geigy (1953 : 41) found that freshly emerged 
females of Tunga penetrans, when removed from the skin after feeding, without 
giving them a chance to burrow, soon perished without having shown the slightest 
indications of hypertrophy. 





m 






w 



m 




16 

Figs. 15-16. Neotunga euloidea sp. nov. (Paratypes) 15. Dorsal view of segments VII-X. 
16. Spiracular fossae of terga II-VII, 



NEOTUNGA EULOIDEA GEN. N., SP. N. 375 

It would seem a considerable disadvantage to the flea not to be able to enter the 
skin of the pangolin's venter, for the chance may be great that the relatively large 
and heavy flea, hanging down, will be brushed off since the pangolin his short legs 
and the venter may sometimes touch the ground. Moreover, the fact that pangolins 
roll themselves up in a tight spiral when at rest may explain why most of the 
embedded fleas are found on the sides of the venter, close to the scales, for these sites 
will very likely be the easiest for the fleas to reach and there they will presumably be 
least hable to squashing. 

Geigy (1953) and Geigy & Suter (i960) have shown that extracutaneous as well as 
intracutaneous fleas {Echidnophaga gallinacea and Tunga penetrans respectively were 
used for their observations) copulate after the female has fixed herself to the host. 
During and after copulation the female of Echidnophaga remains attached to the host 
by the proboscis, and in the case of Tunga the females are embedded in the skin. 
Since Neotunga euloidea is an Echidnophagoid flea with the habits of a Tungoid, one 
may surmise that here too copulation will take place after the female has entered 
the skin. The female lying in a plane parallel with that of the surface of the skin, 
contact between male and female Neotunga can presumably be closer than that 
between the two sexes of Tunga and less close than between those of Echidnophaga, 
and the aedeagus of Neotunga might therefore be expected to be of a structure inter- 
mediate between that of Echidnophaga and Tunga. 

Of special interest are the size and structure of the abdominal spiracular fossae 
(Text-fig. 16). The metepimeral spiracular fossa is somewhat larger than those of 
terga II and III which are of subequal size, while that of tergum IV is intermediate 
between those two and the large ones of terga V-VII. As Jordan has shown (see 
p. 363 above), in the female Echidnophaga the abdominal spiracular fossae are of 
subequal size and somewhat smaller than the metepimeral fossa, while in female 
Tunga the fossae of terga II-IV are much smaller than the metepimeral one and have 
actually vanished in T. penetrans ; however, in this latter genus the fossae of terga 
V-VII I are greatly enlarged. Jordan postulated the existence of a Tungoid flea in 
which the anterior abdominal fossae would approach the size of the metepimeral 
fossa but with those of terga V-VII I enlarged. As can be seen from Text-fig. i6, this 
is exactly the case in Neotunga euloidea. It is tempting to suppose that the differenti- 
ation in size of abdominal spiracular fossae in this species followed the apparently 
fairly recently acquired Tungoid habit of the female to burrow under the skin of the 
host and is one of several morphological modifications in a basically Echidnophagoid 
structure pointing towards the great morphological adaptations to be observed in 
Tunga. 

In embedded females of Tunga the abdominal organs undergo radical changes 
during the period of hypertrophy. In the hypertrophic females of Neotunga similar 
changes must have taken place, since I found for example the tracheae greatly 
enlarged and very numerous and of the same structure as in Tunga ; the midgut, 
although very long and wide (see Text-fig. 2, in dotted outline), retains its straight 
shape instead of attaining an S-shape or double S-shape as is the case in Tunga, but 
the relative narrowness of the abdomen in Neotunga does not allow a folding of the 
midgut. 



376 f. g. a. m. smit 

Systematic Position of Genus Neotunga 

As has been remarked above, Neotunga euloidea is basically an Echinophagoid flea 
with Tungoid habits. The genus Neotunga has in the female sex the following more 
important characters in common with Echidnophaga : 

Head (Text-fig. 3, cf. Text-fig. 5) : 

(a) frontal margin below the tubercle long ; 
(6) genal seta present, though much reduced ; 

(c) antennal scape and pedicel not set at right angles to the clava ; 

(d) first two claval segments not fused with each other nor with the rest of the 
clava ; 

(e) an indication of an internal supra-antennal ridge ; 

(/) epipharynx with numerous small projections along anterior margin. 

Thorax (Text-fig. 3, cf. Text-fig. 5) : 

[g] prosternosome dorsally tapering to a point which is situated opposite to the 
apex of the antennal clava ; 

(h) prosternosome with a well-developed lateral triangular flange ; 
(i) mesosternal furca long and narrow. 

Legs (Text-fig. 3) : 

(;) hind coxa with a patch of small spiniform setae on the lower part of the inner 
side. 

Abdomen (Text-fig. 12, cf. Text-fig. 13) : 

(k) tergum VIII without a seta above the spiracular fossa ; 
(/) anal tergum with a pair of well-developed anal stylets. 

The following features of female Neotunga are characteristic of members of Tunga : 

Head (Text-fig. 3, cf. Text-fig. 4) : 

[a] frontal tubercle pointing upward (in some members of Echinophaga a frontal 
tubercle may be present, but then it is not so well developed as in A^. euloidea) ; 

(6) microsetae of head pointing upwards ; 

(c) eye somewhat sinuous internally (in some specimens) ; 

[d) claval segments of antenna fused, apart from first two segments ; 
{e) genal lobe much reduced. 

Legs (Text-figs. 6-8) : 

(/) tibiae with only two lateral notches bearing stout setae. 



NEOTUNGA EULOIDEA GEN. N., SP. N. 377 

Abdomen (Text-fig. 12, cf. Text-fig. 14) : 

{g) spiracular fossae, and their tracheae, of terga V-VIII much enlarged (although 
smaller than those in Tunga) ; 
[h) antesensiUal setae absent ; 

{i) genital setae on inner side of tergum VIII much enlarged ; 
(_;) sensilium with eight trichobothria each side. /^ 

General : 

(k) chaetotaxy much reduced. 

In other structural characteristics the genera Tunga, Neottmga and Echidnophaga 
are much ahke, for example aU have a strong hook-like tooth at the ventro-anterior 
angle of the hind coxa which is absent in aU other fleas (there is a small tooth in 
Delopsylla but this monotypic genus is very closely akin to Echidnophaga). 

On the whole the head and thorax and their appendages show a greater afiinity to 
those of Echidnophaga than to those of Tunga, but the abdomen resembles that of the 
latter genus more than that of the former. However, the abdominal modifications are 
a result of adaptation to an ecological niche which is shared only by Tunga and they 
need not necessarily indicate a close phylogenetic relationship. In a system of 
classification Neotunga should be placed somewhere between Echidnophaga and 
Tunga, but nearer to the former than to the latter genus. 

It is possible that Neotunga represents an early step in the evolution from an 
Echidnophagoid ancestor to a form resembling Tunga, and that the ancestors of the 
extant species of Tunga have passed through a stage similar to that exhibited by 
Neotunga. 

To put Echidnophaga in the same supra-generic group as Tunga might seem far- 
fetched as recent classifications have included these genera in two different subfamilies 
or even famihes. However, it should be noted that relationships between groups of 
species tend to be obscured when members of a certain group take to an ecological 
niche or a mode of life which is very different from that occupied by members of 
related groups, and as a result undergo more or less profound morphological changes. 
Thus the intromittent organ of species of Tunga is highly specialized and superficially 
unUke that of any other flea, but in no other flea is the mode of copulation so extreme- 
ly unusual. It follows that the genitalia may not be a reliable guide to phylogenetic 
relationships of fleas which have exchanged the normal ectoparasitic way of liie for 
an endoparasitic one. 

The discovery of the male of Neotunga euloidea will very likely enable us eventually 
to assess the true relationships of Neotunga, but in compliance with General Recom- 
mendation No. 6 of the International Code of Zoological Nomenclature I tentatively 
place this genus, along with Echidnophaga and Delopsylla, in the Family Pulicidae. 

REFERENCES 

Geigy, R., 1953, Sandfloh-Probleme. Naturwissenschaften, 40 : 40-42. 

Geigy, R & SuTER, P., i960, Zur Copulation der Flohe. Rev. suisse Zool. 67 (2) : 206-210, i fig. 



PLATE I 

Neotunga euloidea sp. nov. A. Dorsal view. B. Lateral view. C. Ventral view. 

(Para types). 



Bull. B.M. {N.H.) Entom. 12.7 



PLATE I 



A. 



B. 



C. 




PRINTED IN GREAT BRITAIN BY 
ADLARD AND SON, LIMITED 
BARTHOLOMEW PRESS, DORKING 



HOLARCTIC SAWFLIES 
(HYMENOPTERA : SYMPHYTA) 




ROBERT B. BENSON 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 12 No. 8 

LONDON: 1962 



HOLARCTIC SAWFLIES 
(HYMENOPTERA : SYMPHYTA) 



BY 

ROBERT B. BE NSON 

British Museum (Natural History) \ 




pp. 379-409 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 12 No 8 

LONDON: 1962 



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. 

This paper is Vol. 12, No. 8 of the Entomological 
series. The abbreviated titles of the periodicals cited 
follow those of the World List of Scientific Periodicals. 



Trustees of the British Museum, 1902 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued October, 1962 Price Ten Shillings 



HOL ARCTIC SAWFLIES 
(HYMENOPTERA: SYMPHYTA) 

By ROBERT B. BENSON 

CONTENTS 



1. Introduction. ..... 

2. BlOGEOGRAPHICAL TeRMS 

3. Distribution Patterns of Holarctic Sawflies: 

(a) Circumpolar species 

(6) Circumboreal species 

(c) Incomplete holarctic distributions . 

{d) Introduced species 

4. Note on References and Symbols 

5. List of Holarctic Species 

6. References ...... 



Page 

381 
382 

382 
382 
383 
383 
384 
384 
407 



SYNOPSIS 
A comparative study of the boreal sawflies of the old and new worlds reveals that 134 species 
can now be recorded from both : 34 of these are additions to the N. American fauna ; and 33 more 
were previously treated as different species in the two worlds. Furthermore 30 are presumed to 
have been introduced into N. America from Europe and i into Europe from N. America. These 
holarctic species show several different biogeographical patterns. 

I. INTRODUCTION 

In 1956 a Leverhulme Research Fellowship enabled me to spend five months in 
North America. One of my objects was to correlate the boreal and arctic sawflies of 
North America with those of Eurasia. 

My grateful thanks are due to Mr. G. P. Holland, Head of the Section on Insect 
Taxonomy, of the Entomological Research Institute, Ottawa, for allowing me access 
to the named collections there, including the types, and to the unworked material. 
This latter consisted mainly of the rich collections brought back by the Institute's 
staff since 1947 from many parts of the North American Arctic in the course of their 
Northern Insect Survey (Freeman, 1952). 

Mr. Holland also kindly arranged for me to stay at the Defence Research Northern 
Laboratory, Fort Churchill, Manitoba, on the west coast of Hudson Bay at about 
58° 44' N. lat. for three weeks in June and July. There I was able to make collections 
at the edge of the tundra to compare with similar material I had already collected at 
Abisko in Swedish Lapland in 1948 and 1954. Apart from this material, which is in 
the British Museum, the North American material dealt with in this paper is deposited 
(unless otherwise stated) in the Entomology Research Institute, Ottawa, Canada. 
I am also indebted to Dr. J. Gates Clarke for letting me examine the types of Kincaid, 
Marlatt and Rohwer in the U.S. National Museum, Washington, Dr. Herbert H. Ross 
the types of MacGillivray at Urbana, lUinois, Dr. Rene Malaise (Stockholm) for 



ENTOM. 12, 8 



31 



382 ROBERT B. BENSON 

freely letting me borrow types and other specimens from his own Lapland and Kam- 
tchatka collections, and Mr. E. Lindqvist (Helsingfors) for many loans and gifts of 
representatives of species he has described from North Europe. The Greenland material 
referred to is at Copenhagen and was kindly lent to me by Dr. Borge Petersen in 1959. 

2. BIOGEOGRAPHICAL TERMS 

Owing to the lack of agreement as to how different biogeographical terms should 
be applied it is necessary to define how I am using them here. 

Holarctic: occurring in at least some part of both nearctic and palaearctic 
regions. 

Arctic : associated with tundra beyond or above the tree-line. 

Alpine : similar to the arctic type of fauna, but above the tree-Hne on moun- 
tains distant from regions of northern arctic tundra. 

Montane : other types of mountain fauna above the tree-Hne. 

Subarctic : associated with the boreal coniferous forest belt (taiga). 

Suhalpine : associated with subalpine coniferous forest belt. 

Circumpolar : occurring in arctic tundra not only on the mainland of Fenno- 
scandia but also eastwards across Siberia and the Bering Straits to East Canada. 

Circumboreal : as the preceding (circumpolar) but associated not with tundra 
but mainly with the boreal forest belts. 

The above terms have been applied only in the most general way as the distribution 
of arctic and subarctic sawflies is still very Httle known. In some species almost the 
only information available is from my own collecting in Lapland, Scotland, Switzer- 
land and Canada. It is not therefore reasonable to apply such refinements of distribu- 
tional classification as has been applied for example to the flowering plants by Hulten 
(1950) or Porsild (195 1). 

3. DISTRIBUTION PATTERNS OF HOLARCTIC SAWFLIES 

(a) Circumpolar species 

Species with holarctic distributions are diverse ecologically, but arctic species 
predominate and many of them are circumpolar. Benson (1961) found no less than 
14 circumpolar sawflies in high alpine regions of Switzerland and this number 
included such high arctic species as Amauronematus abnormis and A. arcticola. 
Freeman (1958) estimated that 60 per cent of the Canadian arctic butterflies are 
holarctic in distribution and similar percentages have been estimated for many other 
insect groups. A specimen of Pachynematus parvilabris was collected from Ward 
Hunt Island off the north coast of EUesmere Island, Lat. 83° 05' N. and Long. 
74° 30' W. (less than 500 miles from the North Pole). This is even further north than 
Brenland Fjord, off Independence Fjord, North Greenland, about Lat. 82° N. and 
Long. 31° W., where Amauronematus amentorum (in Copenhagen Mus.) was found. 

(b) Circumboreal species 

These are of various types. Rhogogaster viridis for example shows no subspeciation 
throughout its range, but Pachyprotasis rapae occurs in a shghtly different form in 



HOLARCTIC SAWFLIES (HYMENOPTERA : SYMPHYTA) 383 

North America than in Eurasia though the two forms are assumed to be interfertile 
and conspecific. Other examples are Arge clavicornis and Monophadnoides geniculatus, 
in both of which brown-bodied forms occur sporadically as part of N. American 
though never in Eurasian populations. This increase in brown or red in North Ameri- 
can races of holarctic species also occurs in Loderus, Dolerus and Tenthredo. 

Another kind of circumboreal distribution is shown by Xyela alpigena. This species 
is subalpine in Europe, associated with Pinus cembra. It has also been found in East 
Siberia and Japan where the same Pinus occurs, and presumably the Xyela is spread 
along the intervening mountains in Central Asia on the same host. In eastern North 
America it is associated with the closely related P. strobus. X. obscura has a similar 
distribution. 

(c) Incomplete holarctic distributions 

Many holarctic species are not completely circumpolar or circumboreal. The range 
of some transcontinental Eurasian species such as Heptamelus ochroleuchus , Dolerus 
gessneri, Tenthredo devia or Amauronematus pravus crosses the Bering Straits but 
reaches no further than the Pacific coastal regions of North America. Conversely 
some transcontinental North American species occur in East Asia but do not extend 
beyond the Pacific coastal region : Amauronematus hulteni for example reaches 
Kamtchatka and Dolerus subfasciatus Japan. Some forms such as D. gessneri konowi 
are restricted to the Pacific region of the two continents. 

(d) Introduced species 

It would seem that several species are holarctic now only because they were intro- 
duced accidentally by man. The species marked with an asterisk in the general 
account below are thought, for one or more of the following reasons, to have been 
introduced from Europe to America or, in one instance, from America to Europe : 
(i) that though widespread in Europe or Eurasia as a whole their distribution in 
North America is discontinuous ; they occur only sporadically in the more densely 
populated parts {Macrophya punctum-album and Eriocampa ovata) ; (ii) that their 
discovery in North America has been only recent and they have occurred suddenly 
in epidemic numbers {Gilpinia hercyniae on Picea, G.frutetorum and Diprion similis 
on Pinus, Heterarthrus nemoratus, Profenusa thomsoni and three other Fenusine 
leaf-miners in shade trees, Dolerus nitens and Aneugmenus padi) ; (iii) that they are 
attached to plants that have frequently been imported into North America or are 
not native there [Ardis sulcata on Rosa, Pontania proxima on Salix fragilis, Hoplo- 
campa testudinea on Malus pumila, Pristiphora abbreviata on Pyrus communis, etc.) 
or to a plant introduced from North America to Britain {Nematus tibialis on Robinia 
pseudacacia) . 

It is significant that all except three of the examples of introduced species men- 
tioned above are obhgatory parthenogenetic species, at least in the countries where 
they have become established by introduction. 

In some of them (e.g. Gilpinia hercyniae, and Dolerus nitens) the existence of 
obhgatory parthenogenetic races in Europe was not even suspected until the discovery 
of their introduction into North America was made known. 



384 ROBERT B. BENSON 

It would seem that, other factors being equal, obligatory parthenogenetic races 
have a far greater chance of establishing themselves on introduction, as a single 
individual could give rise to a colony ; but it would also seem that such a colony 
might suffer in its inability to adapt itself to new conditions through lacking the inter- 
play of selection in the recombinations of genes. (Benson, 1950 : 124-8.) 

That nearly all the British Fenusini leaf-mining forest and shade trees have become 
established in North America may have been through the use of forest litter in 
packing goods. 

Some of the introduced species are not associated with plants Ukely to have been 
imported whole and were probably not conveyed in htter used for packing. Species 
such as Dolerus asper, D. nitens and Pachynematus ditellatus were probably introduced 
into North America in ballast through the Atlantic ports. Their larvae feed on Grami- 
neae and Cyperaceae, and they occur in just such open places on the coast as those 
from which the ballast was shipped from England (Lindroth, 1957). 

4. NOTE ON REFERENCES AND SYMBOLS 

References to species already given in Dalla Torre, 1894, Konow, 1905, the North 
American Catalogue of Muesebeck, Krombein & Townes, 1951, or its first supplement 
by Krombein, 1958, are not repeated here. It is assumed that most of the Central 
European species can be identified with the help of Enslin (1912-18), or Benson 
(1952-58) ; exact references to these works are not given unless there is a special 
reason such as the existence of a figure of the saw or penis valve in Benson's work. 
The species marked % are additions to the Hst of North American species and those 
marked f were previously known as N. American but not as holarctic and usually 
by a different name. Species marked with an asterisk are assumed to be holarctic 
only through introduction from Europe or Asia to N. America, or in one instance 
{Nematus tibialis) from N. America to Europe. Species marked $ are obligatorily 
parthenogenetic, normally without functional males, in the races that are holarctic. 



5. LIST OF HOLARCTIC SPECIES 

Arctic and boreal sawfly species are often more variable in colour and form than 
those of more temperate regions, and have more abnormalities. For this reason 
some of the more common species such as Amauronematus fallax, A. polaris, Nematus 
reticulatus and Pontania crassipes have been treated by previous authors as though 
they were groups of species and have been divided accordingly ; and some of the 
more aberrant specimens have even been described as distinct species on the basis of 
single examples. As a result of this a false picture of the Nematinae of boreal and 
arctic Europe has been presented, heading towards chaos. So I have found it is 
necessary to treat all such European species that are based on single specimens as 
species inquirendae until more specimens can be found or their probable identity 
determined. 

Furthermore, if for biological or other reasons it is known that two similar forms 
belong to separate species, it may be necessary to use characters for distinguishing 



HOLARCTIC SAWFLIES (HYMENOPTERA : SYMPHYTA) 385 

them that are extremely difficult to appreciate. But the reverse of this is not true : 
discrete morphological differences often occur between individuals of the same inter- 
breeding populations and their discovery is no proof that more than one species is 
involved. 

XYELIDAE 

t Xyela alpigena (Strobl) 

Pinicola alpigena Strobl, 1895 : 277. 

Xyela brunneiceps Rohwer, syn. n. 

Xyela kamtshatica Gussakovskij, 1935 : 363. 

Xyela alpigena (Strobl) ; Benson, 1938 : 35 and fig. 4. 

Xyela middlekauffi Burdick, 1961 : 343 and figs. 39 and 55. 

Dr. Burdick, to whom I sent Swiss specimens of X. alpigena in exchange for 
paratypes of X. middlekauffi, agrees that the two are synonymous and therefore that 
X. brunneiceps is also. In Europe this species is subalpine with its host Pinus cembra 
L., and occurs presumably on the same plant across Asia to Japan. In eastern N. 
America Burdick gives the hostplant as the closely related P. strobus L. 

f Xyela obscura (Strobl) 

Pinicola julii var. obscura Strobl. 

Xyela japonica Rohwer, 1910 : 99. 

Xyela obscura (Strobl) ; Benson, i960 (c) : iio-iii, figs. 2 and 3. 

Xyela pini Rohwer ; Burdick, 1961 : 327, figs. 45 and 62. 

This species, Uke X. alpigena, is also subalpine in Europe and extends eastwards 
to Japan and N. America. In Europe it is attached to Pinus mugo Terra, but in N. 
America it is known from Pinus banksiana Lamb, P. palustris Mill., P. ponderosa 
Laws, and P. virginiana Mill. The synonymy of X. pini with X. obscura was suggested 
to me by Dr. Burdick. 

PAMPHILIIDAE 

* Acantholyda erythrocephala (L.) 

Europe and North Asia to Japan, introduced into North America. 
Pinus. 

SIRICIDAE 

Sirex cyaneus Fabricius 

Circumboreal species. Sirex varipes Walker is not a synonym of this species (Benson, 

1943)- 
Coniferae. 

f * Sirex noctilio Fabricius 
Sirex noctilio Fabricius ; Benson, 1943 : 37 and fig. la. 
Europe, North Asia and Manitoba (? introduced). 
Coniferae. 



386 ROBERT B. BENSON 

f* Sirex juvencus L. 

Sirex juvencus L.; Benson, 1943 : 37 and fig. 16. 
Europe, North Asia, also Labrador and Newfoundland (? introduced). 
Coniferae. 

f Urocerus gigas flavicornis (Fabricius) 

Urocerus gigas flavicornis (Fabricius) ; Benson, 1943 : 39. 
North American subspecies of Eurasian species, 
Coniferae. 



Xeris spectrum (L.) 



Circumboreal species. 
Coniferae. 



CEPHIDAE 

*Cephus pygmaeus (L.) 

Europe, Asia Minor, Caucasus to Turkestan, introduced into North America. 
Gramineae especially cultivated wheat, barley and rye. 

* Trachelus tahidus (Fabricius) 

Europe, North Africa, Asia Minor, Caucasus and south-west Asia, introduced into 
North America. 

Gramineae especially cultivated wheat and rye. 



DIPRIONIDAE 

* Neodiprion sertifer (Geoffroy) 

North and Central Europe to Caucasus, Corea and Japan. Introduced into north- 
east America, westwards to Ohio. 
Pinus. 

* Diprion similis (Hartig) 

North and Central Europe. Introduced into north-east America. 
Pinus. 

* Gilpinia frutetorutn (Fabricius) 

North and Central Europe. Introduced into north-east America. 
Pinus. 



* Gilpinia hercyniae (Hartig) ? 

North and Central Europe. Introduced into East Canada. 
Picea 



HOLARCTIC SAWFLIES (HYMENOPTERA : SYMPHYTA) 387 

ARGIDAE 

* Arge ochropus (Gmelin) 

Europe, West and Central Siberia, Asia Minor, North Persia, Caucasus and Trans- 
caspia, introduced into Ontario. 
Rosa. 

f Arge clavicornis (Fabricius) 
A . fuscipes (Fall6n) syn. n. 

The North American form is extremely variable in colour and may have its abdo- 
men entirely metallic blue, more or less marked with yellow or almost entirely yellow. 
The European forms are A. clavicornis expansa (Klug) comb. n. of subarctic- 
subalpine regions (Benson, 1951 : 34) and A. clavicornis fuscipes (Fallen) comb, n. 
of temperate parts of Central and Western Europe and are without yeUow markings. 



TENTHREDINIDAE 

Selandriinae 

Selandriini 

Brachythops flavens (Klug) 

Selandridea vanduzeei Rohwer, 191 1, syn. n. 

Circumboreal. 
Carex. 

I Brachythops wiistnei (Konow) 

Central and Northern Europe. Canada, Manitoba, Churchill, 5 ^, 12-13 . vi . 1952 
(J. G. Chillcott). I ^ 29. vi. 1956 {R. B. Benson). 
? Circumboreal. 
Carex. 

Aneugmenini 
* Aneugmenus padi (L.), $ 

Selandria urbis Ross. 

Europe, North Africa, Asia Minor and Transcaucasia, introduced and widespread 
in Canada. 

Pteridophyta. 

Heptamelini 

J Heptamelus ochroleucus (Stephens) 

North Eurasia and British Columbia, Squamish, Diamond Head Trail, 3,300 ft., 
I $, 5.viii.i953 {Edith Mason) (Ottawa). 
Pteridophyta. 

ENTOM. 12, 8 3I§ 



388 ROBERT B. BENSON 

Dolerini 

Loderus eversmanni (Kirby) 

Represented by subsp. acidus MacGillivray in North America, by subsp. obscurus 
Marlatt in Japan, and typical subspecies in Europe. 
Circumboreal. 
Equisetum. 

Loderus pratorum (Fallen) 

Represented by subsp. albifrons Norton in North America and East Asia, and 
typical subspecies in Europe. 
Circumboreal. 
Equisetum. 

Loderus vestigialis (Klug) 

Represented by subsp. apHcus Norton in North America, subsp, insulicula Rohwer 
in Japan, and typical subspecies in Europe. 
Circumboreal. 

Equisetum. 

Loderus genucinctus (Zaddach) 

Represented by subsp. napaeus MacGillivray in North America and typical sub- 
species in Europe. 
Circumboreal. 
Equisetum. 

Dolerus yukonensis Norton 

Full synonymy in Benson, 1956 : 58. 

Circumboreal. 

Equisetum. 

Dolerus gessneri Andre 

Represented by subsp. konowi MacGillivray in Pacific North America and Siberia, 
subsp. labiosus Konow in North Europe and Siberia, and the typical subsp. in Western 
Europe. 

FuU synonymy in Benson, 1956 : 59. 

Equisetum. 

Dolerus subfasciatus Smith 

Dolerus picinus rhodogaster Zhelochovtsev 1935 : 79, syn. n. 
Dolerus pseudoanticus Malaise 1931 : 14-15, syn. n. 

This species is represented by subsp. neoaprilis MacGillivray in N. America and the 
typical subspecies in Japan. 
Equisetum. 



HOLARCTIC SAWFLIES (HYMENOPTERA : SYMPHYTA) 389 

Dolerus pratensis (L.) 

Dolerus frisoni Ross. 

North and Central Europe to East Siberia. Alberta, Manitoba and Illinois in North 
America. 

? Circumboreal. 
Equisetum. 

Dolerus elderi Kincaid 

Circumboreal : North Europe to East Siberia and mountains of Central Asia from 
Mongolia to Turkestan, North America from Alaska and the Cordilleras to the Atlantic 
coast. 

Equisetum. 

Dolerus similis Norton 

Transcontinental boreal North American with melanic races (subsp, nicaeus 
MacGiUivray) in the Pacific area, extends into East Asia to Japan (subsp. japonicus 
Kirby with entirely black males, but normal red-banded females). This species is 
very closely similar to the widespread palaearctic D. germanicus (Fabricius) (Ben- 
son, 1952 : 67, 73-74, figs. 185 and 220) which, however, has a deeply excised clypeus 
in front. The two are nowhere sympatric as D. germanicus does not reach Japan, and 
they may be conspecific. 

Equisetum. 

f Dolerus asper Zaddach 

Dolerus asper Zaddach ; Benson, 1952 : 70, 77, figs. 175, 194, 207, 210 and 235. 
Dolerus tectus MacGiUivray syn. n. 

Europe to East Siberia and, in America, recorded from Connecticut, Massachusetts, 
Michigan and South Dakota. 
Cyperaceae and Gramineae. 

Dolerus nitens Zaddach, $ 

Dolerus wanda Ross $. 

Dolerus nitens Zaddach ; Benson, 1952 : 70, 76, figs. 174, 196, 205, 209 and 237. 

North and Central Europe, New York, Ohio and lUinois in North America. The 
American race is obligatorily parthenogenetic like the high-alpine Swiss race and may 
prove to represent a different species for which the name wanda is available. 

Tenthredininae 

Heterarthrini 

* Heterarthrus nemoratus (Fallen) 

Europe, introduced to Nova Scotia about 1908 and now spread over most of 

eastern North America. 
Betula. 



390 ROBERT B. BENSON 

Empriini 

* Monostegia abdominalis (Fabricius) $ 

Europe to Asia Minor, Caucasus and Siberia, also eastern North America where it 
was probably introduced. 

Primulaceae [Anagallis, Lysimachia and Glaux). 

X Empria candidata (Fallen) 

Central and North Europe to East Siberia and across North Canada from Yukon 
to Quebec. Yukon, Dawson, i $, 28. vi. 1949 {W. W. Judd) ; North West Territory, 
Mackenzie Delta, Reindeer Depot, 3 $, 28 . vi . 1948 (J. R. Vockeroth) ; Salmita Mines, 
64° 06' N., 111° 15' W. 1$, 4.vii.i953 (/. G. Chillcott) ; Alberta, Banff, i $, 
25. V. 1922 (C. B. D. Garrett) ; Manitoba, Gilan, i $, 18. vi. 1950 {W. J. Brown) ; 
Quebec, Pt. Chimo, i ?, 16. vi. 1948 [H. M. Smith). 

Betula. 

Ametastegia equiseti (Fallen) 
Circumboreal. 
Polygonaceae [Rumex spp.), Chenopodiaceae [Chenopodium) and Lythraceae. 

Ametastegia glabrata (Fallen) 
Circumboreal. 
Polygonaceae and Chenopodiaceae. 



Circumboreal. 

Viola. 

Circumboreal. 
Rumex. 



Circumboreal. 
Rosa. 



Ametastegia pallipes (Spinola) $ 
Ametastegia tetter (Fallen) 



AUantini 
Allantus cinctus (L.) 



* Allantus basalis (Klug) 

Europe and Siberia, introduced into eastern North America. 
Rosa. 

Blennocampini 

* Ardis sulcata (Cameron) 

Europe, introduced into Washington. 
Rosa. 



HOLARCTIC SAWFLIES (HYMENOPTERA : SYMPHYTA) 391 

Monophadnoides geniculatus (Hartig) 
Circumboreal. 
Rubus. 

* Halidamia affinis (Fallen) $ 

Europe, introduced into New York. 
Galium aparine L. and G. mollugo L. 

* Stethomostus fuliginosus (Schrank) 

Europe to Caucasus and also Japan, introduced into eastern North America (Massa- 
chusetts and New York). 
Ranunculus. 

Eutomostethus ephippium (Panzer) $ 

Europe, North Africa, Asia Minor, Caucasus, Transcaucasia, Himalayas, Siberia 
and North America. 
Gramineae. 

Eutomostethus luteiventris (Klug) $ 

Europe, Transcaucasia and North America. 
Juncus and Gramineae. 

f Monophadnus pallescens (Gmelin) 

Monophadnus aequalis MacGillivray syn. n, 

Europe, Siberia and eastern North America. 
Ranunculus. 

Galiroini 

* Endelomyia aethiops (Fabricius) 

Europe to Caucasus, introduced into North America where it is now widespread. 
Rosa. 

* Caliroa cerasi (L.), ? 

Throughout temperate Eurasia, probably introduced into North America now 
estabUshed also in South America, South Africa, Tasmania and New Zealand. 
Rosaceous trees, especially cultivated Pyrus and Prunus. 

Fenusini 
I Messa wustnei (Konow) 

Messa alaskana (Kincaid). 

Messa wustnei (Konow) ; Benson, 1959 : 90. 

Boreal and arctic Eurasia and North America. 
Salix. 



392 ROBERT B. BENSON 

f * Profenusa thomsoni (Konow), $ 

Profenusa alumna MacGillivray. 

Profenusa thomsoni (Konow) ; Benson, 1959 : 91. 

Temperate Europe and Japan, introduced into North America where it is now 
widespread. 
Betula. 

* Fenusa dohrnii (Tischbein), $ 

Eurasia and North America, introduced into South Africa. 
Alnus. 

* F. pusilla (Lepeletier) 

Eurasia and North America whither it has probably been introduced. 
Betula. 

* Kaliofenusa ultni (Sundewall), $ 

Europe, introduced into eastern North America. 

Ulmus. 

Eriocampini 

* Eriocampa ovata (L.), $ 

Europe, introduced into British Columbia. 
Alnus. 



Circumboreal. 
Alnus, Betula etc. 



Tenthredinini 
Rhogogaster viridis (L.) 



Tenthredo olivacea Klug. 
Boreal Eurasia and Alaska. 

t Tenthredo colon Klug. 

Tenthredo eburneifrons Kirby syn. n. 

Tenthredo nigricollis Kirby syn. n. 

Tenthredo jocosa Provancher syn. n. 

Tenthredo livida L.; Ross in Muesebeck, Krombein & Townes. nee L. 

Europe, Asia Minor, Armenia, Siberia ; and Ontario, Quebec and Labrador in North 
America. 

Polyphagous. 

f Tenthredo atra L. 

It would seem that T. leucostoma Kirby of eastern North America and other forms 
in the same species-group are geographical races of T. atra, which is widespread in 
Eurasia from north-western Europe to East Siberia. 

Circumboreal. 



HOLARCTIC SAWFLIES (HYMENOPTERA : SYMPHYTA) 393 

f Tenthredo devia (Konow) 

Tenthredo heraclei Kincaid. 
Tenthredo alaskana Enslin. 
Tenthredo devia (Konow) ; Benson, 1959 : 94, 100 and fig. 8. 

Lapland, North Russia, North Siberia to Kamtchatka and Japan, Aleutian Islands 
to Popof Island, Alaska. 

Macrophyini 

Pachyprotasis rapae (L.) 

Europe to Caucasus, Siberia, North China and transcontinental Canada and north 
United States. 

Antirrhinum, Betonica, Scrophularia, Solidago and Fraxinus. 

* Macrophya punctumalbum (L.) $ 

M acrophya erythropa Schrank. 

Europe to Caucasus, introduced into Ontario. 
Fraxinus and Ligustrum. 

Nematinae 
Gladiini 
Priophorus morio (Lepeletier) 
Priophorus hrullei Dahlbom ; Benson, 1958 : 142, figs. 352 and 358. 

Circumboreal. Europe to Caucasus, Japan and North America, introduced into 
New Zealand and Tasmania. I 

Ruhus. 

Priophorus pallipes (Lepeletier) 
Priophorus pallipes (Lepeletier) ; Benson, 1958 : 143, figs. 351, 359 and 360. 

? Circumboreal. Europe to Caucasus and Transcaucasia, Siberia, Japan and North 
America. 

Rosaceae (such as Crataegus, Fragaria, Laurus, Prunus and Pyrus) and also Betula. 

* Trichiocampus viminalis (Fallen) 
Trichiocampus viminalis (Fallen) ; Benson, 1958 : 143, fig. 355. 

Europe and North Asia, introduced to both west and east coasts of North America. 
Populus and sometimes Salix. 

Cladius difformis (Panzer) 

Circumboreal. 
Rosaceae. 



394 ROBERT B. BENSON 

Nematini 

* Hoplocatnpa testudinea (Klug) 

Europe to Caucasus, introduced into Western Canada and New York. 
Malus pumila Mill. 

Hemichroa crocea (Geoffroy) 

Circumboreal. 
Alnus. 

* Pristiphora abbreviata (Hartig) ? 

Pristiphora californica Marlatt. 
Central and southern Europe, introduced into California. 
Pyrus communis L. 

* Pristiphora geniculata (Hartig) 

Europe to Eastern Siberia, introduced into east of North America. 
Sorbus. 

Pristiphora pallipes Lepeletier $ 

Pristiphora rufipes Lepeletier. 
Circumboreal. 
Ribes rubrum L., R. uva-crispa L., etc. 

Pristiphora quercus (Hartig) 
Pristiphora idiota Norton. 
Circumboreal. 
Vaccinium and Betula. 

I Pristiphora staudingeri (Ruthe) 

Pristiphora staudingeri (Ruthe) ; Lindqvist, 1953 : 220-222, figs. 2 and 6 ; Benson 1958 : 167, 
figs. 461, 467. 

Circumpolar : arctic-alpine in Europe, northern Siberia to Kamtchatka and Canada 
Manitoba, Churchill, vi.1956, {R. B. B.). 
Salix. 

X Pristiphora leucostoma (Lindqvist) comb. nov. 
Lygaeophora leucostoma Lindqvist, 1952 : 108-109, figs. 42 and 61. 

? Circumpolar. 

I collected samples of this species in Swedish Lapland in 1954 and in Canada, 
Manitoba, Churchill, vi-vii,i956. It was also found in the North-west Territory, 
Mackenzie Delta, Reindeer Depot, 6.vii.i948 (/. R. Vockeroth). 

Salix. 



HOLARCTIC SAWFLIES (HYMENOPTERA : SYMPHYTA) 395 

I Pristiphora reuteri (Lindqvist) comb. nov. 

Lygaeophora arcticola (Enslin) ; Lindqvist, 1952 : iio-iii, figs. 23, 44, and 62. nee Enslin. 
Lygaeonematus {Lygaeophora) reuteri Lindqvist, i960. 

This arctic Eurasian species (N. Europe to Kamtchatka) is probably circumpolar 
as I found it in Canada, Manitoba, Churchill, vi-vii.1956 [R. B. B). 
Salix. 

X Pristiphora pseudocoactula Lindqvist 
Lygaeonematus pseudocoactulus Lindqvist, 1952 : 115, figs. 29 and 50. 

Probably circumpolar : Canada, Manitoba, Churchill, i$, 3.vii.i956 {R. B. B). 

f Pristiphora coactula (Ruthe) 
Pristiphora Winnipeg (Norton), syn. n. 
Lygaeotus coactulus (Ruthe) ; Lindqvist 1952 : 84-87, figs, i, 31 and 52. 

Vaccinium. 

Circumpolar and arctic-alpine in Europe. This and the two following species are 
kept distinct with some diffidence. The slight differences in the genitalia given by 
Lindqvist are not reliable. The ranges are identical. They are probably forms of one 
species. 

I Pristiphora borea (Konow) 

Lygaeonematus boreus Konow, 1904 : 253. 

Lygaeotus boreus (Konow) ; Lindqvist 1952 : 93-96, figs. 6, 33 and 54. 

Pristiphora borea (Konow) ; Benson, 1958 : 171, figs. 459, 464 and 473. 

Likewise circumpolar and arctic-alpine in Europe. In Canada, Manitoba, Churchill, 
it was common in June- July 1956 with the preceding and succeeding species. It was 
also found in North-west Territory, Mackenzie River, Reindeer Depot, 1948 (J. R. 
Vockeroth), Salmita Mines 64° 00' N., 11° 15' W., 1953 (/. G. ChillcoU), and Padlei, 
1950 {R, A. Henniger) ; Quebec at Bradore Bay, 1929 {W. J. Brown), and Great 
Whale River, 1949 (/. R. Vockeroth) ; and Labrador, Goss Bay, 1948 {W. E. Becket). 

f Pristiphora lativentris (Thomson) 

Pristiphora bucoda Kincaid. 

Lygaeotus lativentris (Thomson) ; Lindqvist, 1952 : 88-90, figs. 2-4, 32 and 53. 

Pristiphora lativentris (Thomson) ; Benson, 1958 : 169, figs. 465 and 472. 

Likewise circumpolar and arctic-alpine in Europe. In Canada, Manitoba, Churchill, 
it occurred with the two preceding species in June-July 1956 and was also found in 
North-west Territory, Mackenzie River, Reindeer Depot, 1948 {J. R. Vockeroth). 

Pristiphora mollis (Hartig) 
Pristiphora mollis (Hartig) ; Benson, 1958 : 169, figs. 457, 462 and 468. 
Circumboreal. 
Vaccinium. 



396 ROBERT B. BENSON 

Pristiphora erichsonii (Hartig) 

Circumboreal. 
Larix. 

Amauronematus amentorum (Forster) 

Amauronematus suavis (Ruthe). 

Pontopristia kamtchatica Malaise, 1931 : 53-54, fig. 24a. 

Amauronematus amentorum (Forster) ; Benson, 1958 : 182, figs. 514, 515 and 533 ; Benson, 1959 
(b) : 102-3, fig. I. 

Circumboreal including North Greenland, Bronlund Fjord, Independence Fjord, 
Klares0, 82° N., 31° E., i ^, 24. vi. 1949 Danish Pearyland Exp., (P. Johnson) ; East 
Greenland, c. 76° 46' N., i $, beginning of vii.1908 ; West Greenland, Jakobshavn 
c. 69° N., I ?, 28. vi. 1951 (C. Vibe). 

I Amauronematus itelmena (Malaise) 

Pontopristia itelmena, Malaise, 193 1 : 54, fig. 246. 

Besides Kamtchatka this species also occurs in Canada, Manitoba, Churchill, where 
I collected i $, 24 . vi . 1956, on Salix alexensis (Anderss) Cov. 

I Amauronematus microphyes (Forster) 

Pontopristia romani Malaise, 1921 : 14, fig. 46. 
Pontopristia nana Lindqvist, 1949 : 68-69, fig- 8. 

Besides Europe this species also occurs in Canada, Ontario, Ottawa, where I caught 
1$, 7.V.1956. 
Salix. 

% Amauronematus groenlandicus Malaise 

Amauronematus groenlandicus Malaise, 1933 : 3-4, figs. la-c. 

A. carbonarius Hell6n, 195 1 : 71-2, syn. n. 

A. carbonarius Hell6n ; Lindqvist, 1959 : 7-8 figs. 13 and 14. 

This high-arctic species is circumboreal. It occurs on mountains in Norway and 
Swedish Lapland (I found it between 2,000 and 3,000 ft. near Laktatjakko and Bjork- 
liden in vi.1954). Lindqvist records it also from Finland and Novaja Zemlja. I am 
indebted to Dr. B. Petersen (Copenhagen) for lending me 2 $ (from Greenland) that 
he had compared with the type of groenlandicus. The Greenland localities are as 
foUows : Scoresby Sund, 19.vii.1956 and Mestersvig, King Oscars Fjord, 27. v. 1953 
(C. Vihe), and the type locality was Wollaston Foreland. These places are all near 
together on the mid-east coast of Greenland between 70° and 75° N. 

In the Canadian national collections at Ottawa there are 6 3* 11 $ from : N.W. 
Territory : Spence Bay, Coral Harbour in Southampton Island and Chesterfield in 
the North Hudson Bay region and also from Cambridge Bay (Victoria Island), 
Coppermine and Norman WeUs ; and from Alaska, Sleese Bay. AU these specimens 
were collected by the staff from Ottawa in 1950, 1951 and 1952. 

? Salix. 

This species is very closely related to A . hyperboreus (Thomson, 1871) and is prob- 
ably only a high-arctic form of this species, but in all the specimens before me the 



HOLARCTIC SAWFLIES (HYMENOPTERA : SYMPHYTA) 397 

third antennal segment in $ groenlandicus is about as long as the shortest eye measure 
and in $ hyperboreus it is about xi^ this length. Unfortunately the comparative 
measurements of antennal segments are not altogether reliable, nor are minute 
differences in saw tooth and penis valves in arctic insects as used by Lindqvist (1959).* 

I Amauronematus hebes Konow 

Amauronematus hebes Konow, 1907 : 17-18 (footnote). 
Amauronematus pristiphorinus Malaise, 1931 : 2 and fig. 29. 
Amauronematus hebes Konow ; Lindqvist, 1941 : 71. 

This species is probably circumpolar as it was found in Canada, North-west 
Territory; Chesterfield, 15.vii.1950 (/. G. Chillcott) ; and Mackenzie Delta, Rein- 
deer Depot, 6.vi.i948 (/. R. Vockeroth) ; and Coppermine, vi-vii.1951 (S. D. Hicks). 

X Amauronematus hulteni Lindqvist 

Amauronematus variabilis var. hulteni Malaise, 1931 : 43. 
Amauronematus hulteni Lindqvist, 1941 (i) : 62-63. 

This species was originally described as one of several varieties of A. variabilis 
Malaise in Kamtchatka and has not been recorded elsewhere. In Canada it has been 
found in : Yukon Territory, Rampart House, 28.V.1951 (/. E. H. Martin) ; North- 
west Territory, Norman Wells, 16-18. v. 1953 (C D. Bird) and Chesterfield I5.vii. 
1950 (/. G. Chillcott) ; and Manitoba, Churchill, in vii.1956 {R. B. B.). The species 
is very close to A . helleni Lindqvist but has a smooth unpunctured scutellum. 

I Amauronem.atus helleni Lindqvist 

Amauronematus helleni Lindqvist, 1941 (2) : 65-66. 

This species was previously known only from Finland and North Sweden. I have 
seen it from Canada as follows : North-west Territory : Mackenzie River, Reindeer 
Depot, 25. vi. 1948 (W. J. Brown), 6.vii.i948 (/. R. Vockeroth) ; Norman Wells, 
3.vii.i949 {W . R. M. Mason) ; Salmita Mines, 64° 5' N., 111° 15' W., 22. vi. 1953 
(/. G. Chillcott) ; Bathurst Inlet, 30.vi.1951 {W. I. Campbell) ; Victoria Island, 
Cambridge Bay, i.vi.1950 {E. H. N. Smith) ; Muskox Lake, 64° 45' N., 108° 10', 
12.vii.1953 (J. G. Chilcott) ; Repulse Bay, 26. vi. 1950 (/. E. H. Martin) ; Baffin 
Island, Frobisher Bay, i $, 24. vi. 1948 {T. N. Freeman) ; Chesterfield, 7-12 . vii . 1950 
(J. G. Chillcott and /. R. Vockeroth) ; Padlei, 30 . vi-19 . vii . 1950 {R. A. Henniger) ; 
Quoich River, 65° N., 94° 30' W., 22. vii. 1950 (/. G. Chillcott). Manitoba : Churchill, 
12. vi. 1950 {H. J. Teskey), 3.vi.i952 (/. G. Chillcott), 22 . vi-4 . vii . 1956 {R. B. B.). 
Quebec : Great Whale River, 20. vi. 1949 (/. R. Vockeroth). 

This species is very close to A . arcticola (Dalla Torre) from which it differs in that 
the saw teeth are not produced at their bases. Furthermore the head and thorax are 
usually far more extensively coloured with yellow. 

* The synonymy of Amauronematus hyperboreus (Thomson, 1871) includes at least : Amauronematus 
poppa Konow, 1904, syn. n. ; A. coracinus Lindqvist, 1939 (i) : 6-7, figs. 9 and 10, syn. n. and ^4. enslini 
Lindqvist, 1959 (i) : 8-9, figs. 13 and 14, syn. n., and probably also A. anihracinus Lindqvist, 1959 (i) : 
II, figs. 17 and 18, A. macrophthalmus Lindqvist, 1959 (i) : 13-14, figs- 21 and 22, and ^. obesus Lindqvist, 
1959(1) : 11-12, figs. 19 and 20. 



398 ROBERT B. BENSON 

I Amauronetnatus arcticola (Dalla Torre) 

Nematus arcticus Thomson, 1871, nee Holmgren, 1869. 

Amauronematus arcticola Enslin, 1915 : 389 nee Dalla Torre, 1894 ; Benson, 1958: 185, 189; 
figs. 516, 551, 558 and 577. 

Previously recorded from arctic and alpine Europe and Kamtchatka ; was found 
in Alaska: Nome, vi.1951 {D. P. Williams) and Canada: North-west Territory, 
Coppermine, 12.vi-10.vii.1951 (S. D. Hicks) and Southampton Island, Coral 
Harbour, 9.vii.i948 {G. E. Shewell)). 

Salix. 

I Amauronematus tillbergi Malaise 

Amauronematus tillbergi Malaise, 1920 : 125, fig. 26. 

A. tillbergi Malaise ; Benson, 1958 : 184, 187 and fig. 551. 

Previously recorded from northern and arctic Europe and Kamtchatka in East 
Asia ; was found in Alaska : Nome : 16. vi. 1951 (D. P. Williams) and Canada, North- 
west Territory, Chesterfield, 25.vii.1950 (J. R. Vockeroth). 

Salix. 

I Amauronematus polaris (Holmgren) 

Amauronematus polaris (Holmgren) ; Lindqvist, 1944 : 14. 

For a full synonymy of this very variable species see Benson, 1961. 

Circumboreal arctic species, also alpine in Europe and West Canada. Holmgren's 
type was from Novaja Zemlja. North American records : Alaska, Steese Highway, 
Mile no, I ?, 31 . vi . 1951 [Mason and McGillis) ; Alberta, near Lake Louise, c. 7,000 ft., 
2 $, 22-25 . vi . 1956 {Joyce Benson). North-west Territory, Salmita Mines, 64° 50' N., 
11° 15' W., I $, 22. vi. 1953 (/. G. Chillcott) ; Reindeer Depot, Mackenzie River, i $, 
28. vi. 1948 (/. R. Vockeroth) ; Southampton Island, Coral Harbour, i $, 26. vi. 1952 
(P. Ehrlich). 

Salix. 

X Amauronematus semilacteus (Zaddach) 
Amauronematus semilacteus (Zaddach) ; Benson, 1958 : 187. 

In addition to its occurrence in North and Central Europe, i $ was collected in 
Canada, British Columbia, Vernon, 24.V.1918 {W. Downes). 

Salix. 

X Amauronematus variator (Ruthe) ? 

Am^auronematus variator (Ruthe) ; Lindqvist, 1959 : 2, figs. 5 and 6. 

This arctic-alpine species of Eurasia is evidently circumboreal with the following 
nearctic records. Alaska: Nome vi.1951 [D. P. Williams); Richard Highway, 
Mile 213, vi.1951 (/. R. McGillis) ; Yukon Territory, Aklavik, vi.1953 (C. B. Bird) ; 
North-west Territory : Norman Wells, vi.1949 (W. R. M. Mason) ; Southampton 
Isle, Coral Harbour, vii . 1948. (G. E. Shewell) ; Padlei, vi-vii . 1950 {R. A . Henniger) . 
Manitoba, Churchill, vii. 1950 {H. J. Teakey) and vi.1952 {J. G. Chillcott). 



HOLARCTIC SAWFLIES (HYMENOPTERA : SYMPHYTA) 399 

It is curious that this widespreading species should be apparently entirely partheno- 
genetic, while its close relative A. godmani Benson (1955 : 104-105, figs. 5-9) with 
normal males, is restricted, so far as is known, to Switzerland and north-west 
Scotland. 

Salix. 

X Amauronematus sagmarius Konow, $ 

Amauronematus viduatinus Malaise, 1931 : 40-41, fig. iic. 
Amauronematus sagmarius Konow ; Benson, 1958 : 179, figs. 536 and 540. 

Circumboreal species. Normally yeUow with only the following parts black : 
antennae above, anchor on mesoscuteUum, metascuteUum and dorsum of abdomen. 

North American specimens as foUows : North-west Territory : Mackenzie River, 
Reindeer Forest, i $, 18.vi.1951 {D. P. Williams) ; Rampart House, 3 $, v-vi.1951 
(J. E. H. Martin) ; Norman Wells, v. 1953 (C. B. Bird) ; Southampton Isle, Coral 
Harbour, vi.1952 (P. B. Ehrlich). Alaska : Nome, i $, 16.vi.1951 [D. P. Williams). 
Alberta ; Lake Louise, 2 $, 25-27 . vi . 1956 {Joyce Benson) . Manitoba, Churchill, 
5 $, 24.vi-4.vii.1956 {R. B. B.). I $, 10. vi.1952 (/. G. Chillcott). Ontario, Ottawa, 
II $, 1.V-22.V.1956 {R. B. B.). Most of these specimens are much darker than in 
the typical form and some even have the thorax entirely black and the head with 
only the temples obscurely brown. 

Salix. 

I Amauronematus pravus Konovf 

Of this species, previously recorded only from North Eurasia, one single example 
has been found in North America : Alaska, Donnelly Dome, Richard Highway, 
Mile 719, I $, 7.vi.i95i [W. R. M. Mason). 

Salix. 

X Amauronematus leptocephalus (Thomson) 

Amauronematus leptocephalus (Thomson) ; Malaise, 1920 : 121-122, fig. 21. 

This arctic Eurasian species has been found in Canada, Manitoba : ChurchiU, i $, 
17. vi.1952 (/. G. Chillcott) and Mile 505, Hudson Bay Railway, 1$, 3. vi.1952 
(J. G. Chillcott). 

f Amauronematus fallax (Lepeletier) 

Nematus trifurcus Kirby, syn. n. 
Pontania quadrifasciata MacGillivray, syn. n. 

Amauronematus nuorbinjargi Saarinen, 1949 (2) : 57-62, figs. 1-5, syn. n. 
Amauronematus hartigi Saarinen, 1950 (i) : 20-22, figs. 4-6. 
Amauronematus striatus Hartig ; Saarinen, 1950 : 22-24, figs. 1-3. 
Amauronematus glacialis Saarinen, 1950 (o) : 45, figs, i, 7 and 13, syn. n. 
Amauronematus amicula Saarinen, 1950 {a) : 47, figs. 2, 8, and 14, syn. n. 
Amauronematus suhnitens Saarinen, 1950 (a) : 49, figs. 3, 9 and 15, syn. n. 
Amauronematus septentrionalis Saarinen, 1950 (a) : 54, figs. 4, 10 and 16, syn. n. 
Amauronematus festivus Saarinen, 1950 (a) : 58, figs. 5, 11 and 17, syn. n. 
Amauronematus propinquus Saarinen, 1950 {a) : 60, figs. 6, 12 and 18, syn. n. 
Amauronematus fallax (Lepeletier) ; Benson, 1958 : 182, 189, fig. 503. 

A common and very variable circumboreal sawfly, subarctic-subalpine in Europe. 
I have collected it in Ontario and Manitoba, Canada and have seen good series of it 



400 ROBERT B. BENSON 

at Urbana, Illinois, and Ottawa, Ontario, from the following North American 
localities : Alaska (Nome and Skagway), North-west Territory (Chesterfield and 
Eskimo Point), British Columbia (Trinity Valley), Alberta (Wabrunum), Yukon 
Territory (Little Salmon and Whitehorse), Manitoba (Avenue, Birtle and Churchill) 
Ontario (Ottawa and St. Martin's Falls, Albany River), Nova Scotia (Kentville). 

Structurally A . fallax has not been distinguished from A . histrio (for differences 
between the two species see Benson, 1958 : figs. 502-503). Furthermore, it is not 
always easy to distinguish between A. fallax (on Salix) and A. schlueteri Enslin, 
1915 : 405 [=A. uUginosae Malaise, 1920 : 122-123, fig. 29, A. lundbohmi Malaise, 
1920 : 119, fig. 19, and ^. squamosus Lindqvist, 1959 (i) : 14, syn. n.) (on Vaccinium) : 
A . schlueteri is usually smaller and the sawsheath is shorter than the basal plate, and 
in dorsal view is only about as long as its greatest breadth. In fallax on the other 
hand the sawsheath is longer than the basal plate and in dorsal view more than x t\ 
its greatest width. 

Of the North- American arctic A. stordalensis (Strand, 1905 17) {= A. marginifer 
(Strand, 1905 : 8) syn. n., A. cogitatus MacGiUivray syn. n., and A. varianus MacGil- 
hvray syn. n.) I have seen Strand's types from S. EUesmere Isle (through the kind- 
ness of Dr. Natvig of Oslo) and specimens from Baffin Land (F. C. Wynne Edwards) 
and, at Ottawa, from Alaska (MacGiUivray t5rpes). Repulse and Cambridge Bay 
(North-west Territory). It differs from A. fallax in its ovipositor being shorter than 
the hind femur [in fallax at least as long as femur and 2nd trochanter together), in its 
malar space (c^ and $) being about xi|^ as long as distance between the antennal 
sockets (in fallax the malar space is only about equal to the distance between the 
antennal sockets) and in its almost entirely black colour [fallax often i marked with 
pale colour). 

The European alpine A. opacipleuris Konow is closely related to A. stordalensis, 
differing in that its head is more strongly contracted behind the eyes. 

f Amauronematus histrio (Lepeletier) 

Atnauronematus luteotergum (Norton), syn. n. 

Amauronematus histrio (Lepeletier) ; Benson, 1958 : 178, 182, 189, figs. 502, 505, 524, 563 and 566 

Another circumboreal species but of more temperate regions giving place north- 
wards to A . fallax. According to Ross, 1951 : 52, it is an eastern species in North 
America, occurring westwards only as far as Illinois. I have, however, collected it at 
Churchill, Manitoba, and in the British Museum is a specimen from Colorado [T. D. A. 
Cocker ell). Ross also gives Alnus as the foodplant of this species on the authority of 
Dyar. The specimens I collected at Ottawa and Churchill were definitely associated 
with Salix and far away from any Alnus. 

f Amauronematus neglectus (W. F. Kirby) 

Amauronematus intermedius Malaise, 1931 : 39-40, syn. n. ; Saarinen, 1949 (i) : 78-81, figs. 
2, 5, 8-10, 16-18 and 24. 

Circumboreal species : in Europe only known in Lapland. East Siberia [Malaise, 
1931) ; British Columbia, Salmon Arm, 1923 [C. R. Buckell) ; Colorado, 1891 [T. D. A. 



HOLARCTIC SAWFLIES (HYMENOPTERA : SYMPHYTA) 401 

Cockerell) ; Yukon, Rampart House, vi. 1951 (/. E. H. Martin) ; North-west Territory, 
Norman Wells, v. 1953, {D. Bird) ; Manitoba, Churchill, vi.1952-1956 (/. G. Chillcott 
and R. B. B.) ; Ontario, St. Martin's Falls, Albany River, before 1844, types of 
Nematus negledus ; and Ottawa v. 1956, {R. B. B.); Quebec, Fort Chimo, vii.1948, 
{R. H. MacLeod); Illinois, Zyon, v. 1956 (/. E. and R. B. B.); Havana, vi.1912 
{A . W. J. Pomeroy) . 

This species is closely related to the European A . mundus Konow, but in the $ shows 
a greater range of colour pattern ; abdomen above entirely yeUow to entirely black 
except for the apex, mesonotum yellow with ± black flecks on the lobes and meso- 
pleura yellow with or without mesosternal fleck to entirely black. {A . mundus always 
has at least three or four basal tergites marked with black.) The differences between 
the saws are figured by Saarinen, 1949 : figs. 8-1 1 and 16-19. 

Salix. 



I Atnauronematus longicauda Hellen 

Nematus (Amauronematus) longicauda Hellin, 1948 : 113 ; Saarinen, 1949 (i) : 81-82, figs. 12, 22 
and 26. 

Saarinen recorded this species only from Lapland. I have seen North American 
specimens as follows : British Columbia, Vancouver, i $, iv.1931 {H. H. Ross) and 
Agassiz, 1$, iv,i932 {R. Glendenning) ; Manitoba, Churchill, 1$, vi.1952 (/. G. 
Chillcott). 

t Amauronematus abnormis (Holmgren) 

Amauronentatus tolli Konow, 1907 : 20-21. 
Amauronematus aulatus MacGillivray, syn. n. 

Amauronematus abnormis (Holmgren) ; Benson, 1958 : 179, figs. 500, 501, 509, 527, 532, 557 and 
569- 

Circumpolar, high arctic and alpine. Lapland, Scotland, Switzerland, Novaja 
Zemlja, Lena River, New Siberian Islands, North Behring Sound, North Alaska 
(Barter and St. Paul Islands), Baffin Island (Marble Canyon) and North-west Terri- 
tory (Chesterfield). 

This species is apparently closely related to the preceding A. longicauda HeUen. 
The curved spine on the penis valve (Benson, 1958, fig. 569) , however, distinguishes the 
male from that of any other species. In the female (Benson, 1958, fig. 501) the short 
wings (shorter than abdomen) with reduced venation and correlated flattening of 
mesonotal lobes should distinguish the species apart from the differences in the shape of 
the saw tooth. The short ovipositor (shorter than hind tibia) will also distinguish it 
from A. longicauda and aU other members of the longiserra-group (in aU others of 
which the ovipositor is as long as or longer than the hind tibia). 

The development of the wings in the female varies greatly between individuals and 
correlated with this the development of the flight muscles in the mesonotum. The 
specimen I found in Switzerland in 1959 had almost normal wings and mesonotum ; 
and it seems probable that females with quite normal wings might occur. 



402 ROBERT B. BENSON 

f Amauronematus leucolaenus (Zaddach) 

Pontania unga Kincaid, syn. n. 

Amauronematus saarineni Lindqvist, 1945 : 106, fig. 5, syn. n. 

This common circumboreal species I have seen from the following localities in 
North America : Alaska, Popof Island, vii . 1899 {T. Kincaid) ; Yukon Territory, 
Rampart House, v.1951 {J. E. H. Martin). North-west Territory, Coppermine, 
vi.1951 (S. D. Hicks) ; Norman Wells, v. 1953 (C. D. Bird) ; Spence Bay, vi.1951 
{A. E. R. Downes) ; Alberta, Lake Agnes near Lake Louise, 6-700 ft., vi.1956 (J. E. 
Benson). Saskatchewan, Saskatoon, v. 1939 {A. R. Brooks). Manitoba, ChurchiU, 
vi.1952 (J. G. Chillcott) and iv.1941 (0. Peck) and Ottawa, v. 1956 {R. B. B.). 

f Amauronematus viduatus (Zetterstedt) 

Amauronematus orhitalis Marlatt, syn. n. 

Amauronematus viduatoides Lindqvist, 1959 [b) : 127-128, fig. i, syn. n. 

This very common European species is also circumboreal but is scarce in North 
America whence I have seen specimens from the following localities : Oregon (type 
of orhitalis). Saskatchewan, Saskatoon, v. 1940 {A. R. Brooks) ; Ontario, Ottawa, 
V.1956 {R. B. B.y. 

I Amauronematus nitidipleuris Malaise 

Amauronematus nitidipleuris Malaise, 1931 : 45 ; Lindqvist, 1945 : 105-106, fig. 4. 

This high arctic circumpolar species described originally from Kamtchatka was 
later recorded from Europe by Lindqvist (1945 : 105, saw fig. 4). I have examined 
the following from Greenland, thanks to Dr. Barge Petersen (Copenhagen) : East 
Greenland : Heklahavn, c, 70° N., i $, vi.1892 [Deichmann) ; Mestersvig, c. 72° N., 
1$; 17. V. 1953, 1$, 22. V. 1953 (C. Vibe); North-west Greenland, Inuarfigssuak, 
Inglefieldland, c. 79° N., i $, 24. vi. 1941 (C. Vibe). In Canada it has so far only been 
found in Manitoba, Churchill, vi.1952 (/. G. Chillcott and P. R. Ehrlich) and vi.1956 
{R.B.B.). 

f Euura mucronata (Hartig) 

Euura insularis Kincaid, syn. n. 

Circumboreal and one of the commonest sawflies in North-west Europe, but seems 
subarctic in North America : Alaska (Kincaid) ; Manitoba, Churchill {R. B. B.). 
This and the following species, which feed as larvae inside Salix buds, have saws 
with projecting marginal teeth (Benson, 1958, fig. 593). E. mucronata feeds on a great 
range of Salix species. Species whose larvae feed in petioles or stems have saws 
without projecting marginal teeth (Benson, 1958, fig. 592). Euura orhitalis Norton, 
1862 from East Canada differs from E. mucronata in having pale hind orbits and 
temples, but may be no more than a geographical race. 



HOLARCTIC SAWFLIES (HYMENOPTERA : SYMPHYTA) 403 

I Euura lanatae Malaise 
Euura lanatae Malaise, 1920 : 105, fig. 4. 

This species is abundant in Lapland where, however, it is attached to Salix lanata L. 
(see preceeding species). At Churchill, Manitoba, I found the species on the closely 
related Salix calcicola Fern, and Wieg in vi-vii.1956, and my wife collected it in 
Alberta, Lake Louise, at about the same time. 

t Euura atra (Jurine) 

Euura nigra Provancher, syn. n. 

This species was not uncommon about Ottawa, Ontario, in vi.1956, and there is a 
bred series in the Museum at Ottawa from Alberta, Brooks, v. 1952. The type of 
E. nigra was from Quebec. The larvae of this species feed in stems of many of the 
narrow-leaved Salix, such as repens L., viminalis 'L.,fragilis L. Sind purpurea L. For 
the saw see note under E. mucronata above. 

I Phyllocolpa coriacea (Benson) 

Phyllocolpa coriacea (Benson) ; Benson, 1960 [a) : 60. 

This species, which is boreo-subalpine in Europe, I have found in north Scandinavia, 
north Britain and Switzerland, and my wife found it near Lake Louise, Alberta, in 
vi.1956. Salix ? spp. This genus Phyllocolpa (Benson, i960 {a)) and the following 
genus Pontania were studied comprehensively by Benson, i960. 

t Phyllocolpa excavata (Marlatt) 

Phyllocolpa excavata (Marlatt) ; Benson, i960 (a) : 60. 

For full synonymy of this species see Benson, i960 : 380. 

Another boreo-subalpine species, circumboreal in distribution, forming leaf edge 
rolls on Salix pentan4ra L. in Europe and presumably on this also where introduced 
into North America as well no doubt as on related species. 

J Phyllocolpa acutiserra (Lindqvist) 

Pontania acutiserra Lindqvist, 1949 : 66-68. 
Phyllocolpa acutiserra (Lindqvist) ; Benson, i960 (a) : 60. 

This species, previously known from Lapland and Scotland, I found at Manitoba, 
Churchill, vi.1956. 

f Pontania crassipes (Thomson) 

The definition and synonymy of this species were discussed by Benson, i960 and 
later in i960 [h) : 179-180. It is a circumpolar arctic-alpine species very variable in 
structure and colour, and forming pea-shaped galls on the main vein usually mainly 
on the underside of a leaf of the following willows : Salix reticulata L., 5. herhacea L., 
5. Polaris Wheb., S. arctica Pall., S. myrsinites L,, S. lapponum L., and 5. arbuscula L. 



404 ROBERT B. BENSON 

I Pontania dolichura (Thomson) 

The synonymy of this species was given by Benson, i960. In Europe it is boreo- 
alpine and in Canada I found it at Manitoba, Churchill, in vi. 1956. The known host 
range is : Salix phylicifolia L., S. nigricans Smith, S. purpurea L., and occasionally 
S. arhuscula L., S. lapponum L., S. myrsinites L., S. lanata L., 5. glabra Scop., 
5. incana Schrank and S. retusa L. in Europe ; 5. sachalinensis R. Schmidt, in Kam- 
tchatka ; and 5. planifolia Pursh. in Canada. 

f* Pontania proxima (Lepeletier) 

Messa hyalina Norton. 

This species is common on Salix fragilis L. and S. alba L. in Europe and where these 
trees have been introduced into Canada. 

Croesus varus (ViUaret) 

Alnus. 

I Nematus crassus (Fallen) 

This widespread Eurasian species was found in North-west Territory of Canada, 
Mackenzie River, Reindeer Depot, i ?, 12.vii.1948. 
Betula, Populus, Salix and Rumex. 

I Nematus villosus Thomson 

This arctic species previously known only from Spitzbergen and Lapland was found 
in North-west Territory of Canada, Chesterfield, i (J, 15.vii.1950 (/. G. Chillcott) and 
Cambridge Bay, i ^, 11.vii.1950 {G. K. Sweetman). 

t Nematus reticulatus Holmgren 

Pontania popofiana Kincaid, and P. glinka Kincaid. 

This circumpolar arctic-alpine is extremely variable in structure and colour, and 
for full synonymy see Benson, 1961. 

In addition to Kincaid's types I have also seen North American material from 
the following localities : Alaska, Pt. Barrow and Upper Colville River, vi-viii.1950 
{N. A. Weber) ; Nome, 16.vi.1951 {D. R. Whillans) ; King Salmon, Naknek River, 
10.viii.1952 (j. B. Hartley) ; North-west Territory, Salmita Mines, 64° 05' N., 111° 
15' W., 22.vi-8.vii.1953 (/. G. Chillcott) ; Muskox Lake, 64° 45' N., 108° 10' W., 
10-11.vii.1953 (/. G. Chillcott) ; Southampton Isle, Coral Harbour, 9.vii.i948 
(G. E. Shewell) ; Manitoba, Churchill, 20. vi. 1952 (C. D. Bird), and 7-11 . vii . 1956 
{R. B. B.) ; Labrador, Hebron, 9 . vii . 1954 (/. F. Mc Alpine) ; New Hampshire, Mount 
Washington, Lake of the Clouds, 29.viii.1954 {Becker, Munroe and Mason). 

Vaccinium. 

Nematus ribesii (Scopoh) 
Ribes. 



HOLARCTIC SAWFLIES (HYMENOPTERA : SYMPHYTA) 405 

* Nematus tibialis (Newman) 

This species, native on Robinia pseudacacia L. in North America, was introduced 
with the tree to Europe. 

I Nematus jugicola Thomson 

I collected a series of this European boreo-subalpine species at Manitoba, Churchill, 
4-1l.vii.1956. The females have an almost entirely black abdomen as in the high- 
arctic forms of Europe. 

Nematus oligospilus Forster 
Nematus mendicus Walsh, syn. n. 

I Pachynematus parvilabris (Thomson) 

Pachynematus inopinatus Lindqvist, 1949 : 82-83, figs. 39-40, syn. n. 
Pachynematus abstrusus Lindqvist, 1949 : 83-84, figs. 43-44, syn. n. 

The two species described by Lindqvist on very shght characters were based on 
three specimens only, and are undoubtedly only forms of P. parvilabris. 

A female collected on Ward Hunt Island (off the North coast of Grant Land, North- 
west Territory) 8^° 05' N., 74° 30' W., at the end of July 1954 [G. H alter sley-Smith) 
must be the most northerly sawfly ever found (cf. Amauronematus amentorum at 
c. 82° 31' N. in North-west Greenland). The specimen is not quite typical, the number 
of marginal teeth to the saw being reduced from 14 to 13, vein 2 rm in the forewing 
is missing and the front lobe of the mesonotum has lost its medial groove. Other 
Canadian specimens are as follows : North-west Territory ; Southampton Island, 
Coral Harbour, 6.vii.i948 [G. E. Shewell) ; Chesterfield, 13 . vii-3 . viii . 1950 (/. R. 
Vockerolh) ; Repulse Bay, 29. vi. 1950 (/. E. H. Martin). 

In Greenland the species has been found at Marrait, Nugssuak, i $, 18. vi. 1949 
(C Vibe). This locality is 70°-7i° N., near the middle of the west coast. 

The species is probably circumpolar. 

- j Pachynematus obductus (Hartig) $ 

Nematus palliventris (Cresson). 
Gramineae. 

t Pachynematus vagus (Fabricius) $ 

Nematus inconspicuus Kirby. 
Pachynematus corticosus MacGillivray. 

Carex and Salix. 

I Pachynematus rumicis (L.) 

Pachynematus rumicis (L.) ; Benson, 1958 : 234, figs. 775 and 798. 

Circumboreal subarctic species reaching Iceland and in North America : Alaska, 
Cold Bay, 163° W., 1$, vii.1952 {W. R. Mason), and Manitoba, Churchill, i ?, 
8.vii.i956 (R. B. B.). 

Rumex. 

This species is closely related to the following species : 



4o6 ROBERT B. BENSON 

I Pachynematus freyi Lindqvist 
Pachynematus freyi Lindqvist, 1949 : 81-82, figs. 37-38. 

This arctic species of Europe and Siberia reaches Canada : North-west Territory, 
Mackenzie, Reindeer Depot, 26-30 . vi . 1948 {W. J. Brown and /. R. Vockeroth), and 
Yukon Territory, Rampart House, 12.vii.1951 (/. E. H. Martin). This is closely 
related to the preceding species P. rumicis but is almost entirely black in colour and 
the saw has only 10 denticulated bands and 5-6 marginal teeth (11 and 9 in rumicis). 

X Pachynematus excisus (C. G. Thomson) 

This arctic European species has been found on the west coast of Greenland at 
Equilet-landet, at about 61° N., and at Gr0nne dal, Godthab, and also at Sarqaq, 
Marrait, and Augpilagtoq between 70° and 73° N., 9-17. vii. 1948-51 (C. Vihe). 
I am indebted to Dr. B0rge Petersen (Copenhagen) for letting me see these specimens. 

Pachynematus moerens (Foerster) 

Pachynematus falonus Ross. 
Gramineae. 



Carex. 



Pachynematus clibrichellus (Cameron) 



f Pachynematus extensicornis (Norton) 

Pachynematus foveolatus Konow, syn. n. 

Pachynematus truncatus (Benson, 1948) ; Benson, 1958 : 237, 241, figs. 788 and 806. 

Gramineae. 

Pachynematus kirbyi (Dahlbom) 

Pachynematus diaphanus Eversmann. 

Pachynematus kirbyi (Dahlbom) ; Lindqvist, 1956 : i ; Benson, 1958 ; 237, 241, figs. 787 and 
805. 

Carex. 

Pachynematus smithiae Ross 

Probably circumboreal arctic-alpine. It was originally described from Mount 
Washington, New Hampshire, but I have collected specimens in Lapland, in the 
Scottish mountains and in the high Swiss Alps. 

I* Pachynematus clitellatus (Lepeletier) 

Pachynematus clitellatus (Lepeletier) ; Benson, 1958 : 237 and 241, figs. 794 and 813. 

I have seen three (^ of this species from East Canada : Labrador, Cartwright, 
14. vii. 1935 {W. J. Brown) and Hebron, 15. vii. 1954 (/. F. McAlpine) ; and Quebec, 
Bonne-Esperance, 14 . vii . 1929 {W. J. Brown) . 

Gramineae. 

Probably introduced into Canada from Europe. 



HOLARCTIC SAWFLIES (HYMENOPTERA : SYMPHYTA) 407 

6. REFERENCES 

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Symphyta). Proc. R. ent. Soc. Lond. (B) 7 : 32-36. 
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Bull. ent. Res., London, 34 : 27-51. 17 figs., 3 tables. 
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Ent. mon. Mag. 84 : 58-65, 36 figs. 
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4o8 ROBERT B. BENSON 

KoNOW, F. W. 1907. Ueber die Ausbeute der Russischen Polar Expedition an Blattwespen im 
arctischen Sibirien. Res. Sci. Exp. Polaire Russe en 1900-1903 sous la direction du baron 
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1941(a). Einige neue Nematinen-Arten und -Aberrationen (Hym. Tenthredinidae). Notul. 

ent., Helsingf. 21 : 64-70. 

1944- t)ber die von Holmgren beschreibenen Nematinae (Hymenoptera Tenthredinidae). 

Notul. ent., Helsingf. 24 : 13-18. 

1944(a). t)ber einige hochnordische Nematinen (Hymenoptera, Tenthredinidae). Notul. 

ent., Helsingf. 24 : 18-24. 

1945, t)ber Blattwespen der Amauronematus-Grxy^^e. Notul. ent. Helsingf., 25 : 102-8, 

7 figs. 

1949. Neue nordische Blattwespen. Notul. ent., Helsingf., 28 : 65-86, 51 figs. 

1952. t)ber alte und neue Lygaeonematus Arten (Hymenoptera, Tenthredinidae). Notul. 

ent., Helsingf. 32 : 80-119, 66 figs. 

1953- tJber Pristiphora staudingeri Ruthe und nahverwandte Arten (Hymenoptera, Ten- 
thredinidae). Opusc. ent. 18 : 220-224, 9 figs. 

1956. Revision einiger von schwedischen Entomologen beschriebenen Nematinen (Hymen- 
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ent., Helsingf., 37 : 92-96 (1957), 97-ii7 (1958), 48 figs. 

1959- t)ber einige neue oder wenig bekannte Amauronematus-Arten (Hymenoptera, Ten- 
thredinidae). Notul. ent. Helsingf. 39 : 1-15, 32 figs. 

1959(a). Neue schwarze Pteronidea- hxten (Hymenoptera, Tenthredinidae). Notul. ent. 

Helsingf. 39 : 54-60, 15 figs. 

1959(6). Drei neue Amauronematus-Kxten (Hymenoptera, Tenthredinidae). Notul. ent., 

Helsingf. 39 : 127-130, 12 figs. 
i960. Zur Kenntnis zweier Nematinen (Hym. Tenthr.). Notul. ent., Helsingf. 40 (i) : 33-34. 



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1931. Entomologische Ergebnisse der Schwedischen Kamtchatka-Expedition 1920-1922. 

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1933- Eine neue Blattwespe aus Ost-Gronland. Skr. Svalb. og Ischavet, 53 : 3-4, fig. la-c. 

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optera Symphyta) aus Finnisch-Lappland. Ann. ent. fenn. 15 : 55-62, 5 figs. 
1950. Amauronematus hartigi n. sp., eine neue Blattwespenart (Hymenoptera Symphyta) 

aus der Sammlung Th. Hartigs in Miinchen. Ann. ent. fenn., 16 : 18-24. 
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Symphyta). Ann. ent. fenn. 16 : 44-63, 24 figs. 
Strand, F. 1905. Coleoptera, Hymenoptera, Lepidoptera and Araneae in Rep. 2nd Norwegian 

Arct. Exped. Fram. 1898-1902, 1 (3) : 7-8. 



HOLARCTIC SAWFLIES (HYMENOPTERA : SYMPHYTA] 



409 



Strobl, G. 1895. Beitrage geographischen Verbreitung Tenthrediniden, V. Theil. Wien. ent. Ztg., 

14 : 277-279. 
Zhelochovtsev, a. N. 1935. Notes sur les Dolerinae (Hymenoptera) pal^arctiques. Arch. Mus. 

Zool. Univ. Moscou, 2 : 79-83, 6 figs. 




INDEX TO VOLUME XII 

New taxonomic names are in bold type 



abbreviata, Pristiphora . 


• 394 


balteatus, Peliococcus 


32. 33 (fig) 


abdominalis, Monostegia 


■ 390 


barombiensis, Epitola 


154, PI. 9, 10 


abnormis, Amauronematus 


401 


baryphragma, Eucosma . 


• 253 


absimllis, Oecetis 


1 1 3-1 15 (fig.) 


basalis, Allantus 


■ 390 


abstrusus, Pachynematus 


. 405 


benitensis, Phytala . 


155-156 


aceris, Phenacoccus 


34-35 


berneri, Amphipsyche 


91 (fig.), 92 


acuticerca, Ambrea 


333 (fig). 334 


bicolor, Perineta 


342, 343 (fig.) 


acutiserra, Phyllocolpa . 


. 403 


bimaculatus, Xylophloeus 


189 (fig.), 193-194 


adolphifriderici, Epitola . 


. 130 


biplagiata, Calderonia 


331, 332 (fig.) 


adonidum. Coccus . 


40 


bisltrensis, Cryptolestes 


• 233 (fig.) 


adonidum, Pseudococcus 


. 40 


blepharopis, Bactra 


• 253 


aequalis, Monophadnus . 


• 391 


Blubos .... 


194-196 


Aethaloptera 


96-100 


boisduvalii, Hewitsonia . 


. 160 


aethiops, Endelomyia 


• 391 


boisduvaiii nigeriensis, Hewitsonia 


affinis, Halidamia . 


• 391 




160, PI. II, 12 


africanus, Placonotus 


182-183 


bolivari, Euthymia 


292 (fig.), 293 


alba, Epitola . 


155. PI- 9. 10 


bolivari, Placonotus 


185-186 


albidus, Rhizoecus 


■ 41. 42 (fig.). 43 


boratynskii, Balanococcus 


15, 16 (fig.) 


albomaculata, Epitola 


. 138 


borea, Pristiphora . 


• 395 


albopunctata, Oecetis iii 


(fig.). 112 (fig.). 113 


brachyptera, Eyprepocnemis 


314, 315 (fig.) 


alluaudi, Laemophloeus . 


• 234 


brevicornis, Thisoicetrus . 


• 321 


alpigena, Xyela 


• 385 


brunneiceps, Xyela 


. 385 


ambiguus, Cannes . 


• 175 


brunneri, Lopheuthymia 


295 (fig.), 296 


Ambrea 


• 331. 333-4 


brunneus, Cryptolestes 


233 (fig-). 234 


amentorum, Amauronematus 


• 396 


brunneus, Planolestes 


. 228 


amicula, Amauronematus 


• 399 


bupleuri, Leptophloeus 


209 


Amphipsyche 


85-94 






anaprobola, Olethreutes . 


• 255 


cactearum, Spilococcus . 


52, 53 (fig-). 54 


anastrepta, Crusimetra . 


• 253 


cacticans, Rhizoecus 


• 43 


ancosema ancosema, Olethrei. 


ites . . 255 


cacticans, Ripersiella 


43 


andringitra, Paraspathosternu 


m 279, 280 (fig.) 


caecigena, Tunga . 


353-364 


ankya, Paduniella . 


. 83 


calcaratus, Leptocerus 


109 


anormus, Leptophloeus . 


205 


Calderonia 


• 331 


antennatus, Horaeocerus 


325 (fig). 326 


calida, Labdia 


. 260 


anthoptera, Blastobasis . 


265-266 


californicus, Pseudococcus 


72 


Antonina 


. 7-8 


callichrysa, Anastathma . 


. 268 


arcticola, Amauronematus 


. 398 


calluneti, Parapedronia . 


. 56 


arsitricha, Limnaecia 


260 


calluneti, Pseudococcus . 


• 57 


asper, Dolerus 


. 389 


calluneti, Spinococcus 


56. 57 (fig). 58 


ater, Leptophloeus . 


203 (fig.), 204 


candidata, Empria 


• 390 


atra, Euura . 


• 403 


capensis, Cryptolestes 


233 (figs.), 235 


atra, Tenthredo 


• 392 


capitus, Leptophloeus 


. 208 


Atrococcus . 


. 8-13 


capnographa, Decadarchis 


270 


atuloldes, Cryptolestes 


• 237 (figs.), 238 


capnosticta, Ereunetis 


. 268, PI. 14 


atulus, Cryptolestes 


• 237 (figs.), 238 


captiosella, Gerontha 


. 271 


aulatus, Amauronematus 


401 


carbonarius, Amauronematus . 


• 393 


axillaris, Leptophloeus 


. 206 


carcassoni, Stempfferia 157-15 


8 (fig.), PI. II, 12 


azurea, Epitola 


148. PI. 7, 8 


carcina, Epitola 


140 






carilla, Epitola 


- 134 


badura, Epitola 


. 136 


carinulatus, Laemophloeus 


• 234 


Balanococcus 


. 13. 15. 17 


carpenteri, Epitola . i 


44, PI. 5, 6, 9, 10 


baliensis, Caloptenus 


. 311 


cassavae, Leptophloeus , 


. 207 



412 

Catantopinae 
Catantops 

cataracta, Trymalitis 
catori ugandae, Epitolina 
catuna, Epitola 
cedrinopa, Labdia . 
celidophora, Meridarchis . 
cephalica, Morondavia 
cephena, Epitola 
cephena entebbeana, Epitola 
cerasi, Caliroa 
ceraunia, Epitola . 
cercene, Epitola 
cercenoides, Epitola 
cheesmanae, Heterogymna 
Chloropsyche 
Chnaurococcus 
Chorizococcus 

chrysomeloides, Xylophloeus 
cinctus, Allantus 
citri, Dactylopius . 
citri, Dorthesia 
citri, Planococcus . 
citri, Pseudococcus 
clarus, Notolaemus 
clavicollis, Laemophloeus 
clavicomis, Arge 
clibrichellus, Pachynematus 
clitellatus, Pachynematus 
clodiana, Labdia 
coactula, Pristiphora 
eoerulea, Epitola 
colon, Tenthredo 
comoroensis, Pamphagella 
concepcion, Epitola 
congoana, Epitola . 
conjuncta badduana, Epitola 
conjuncta conjuncta, Epitola 
convexa, Epitola . 
corbeti, Amphipsyche 
coriacea, Phyllocalpa 
cornutus, Leptophloeus . 
comutus, Planolestes 
cracens, Atrococcus 
cramerella, Acrocercops . 
crassipes, Pontania 
crassus, Nematus 
crawi, Antonina 
crippsi, Epitola 
crocea, Hemichroa 
crowleyi crowleyi, Epitola 
cryptogramma, Spilonota 
Cryptolestes . 
Cucujinus 

cnspidatae, Pseudococcus 
cyaneus, Sirex 

dactylis, Trionymus 
dascia, Mesocallyntera 

daveyi, Epitola 



INDEX 


• 326 


decellei, Epitola 


• 326 


decisa, Caloptenopsis 


• 257 


decoratus, Placonotus 


159, PI. 11, 12 


decoratus, Rhizoecus 


143-144, PI. 5. 6 


delosticha, Orthenches 


260 


dentatus, Xylophloeus 


. 262, PI. 14 


devia, Tenthredo 


307, 308 (fig.) 


dianthi, Coccidella . 


146 


dianthi, Morrisonella 


146, PI. 5, 6 


dianthi, Rhizoecus 


■ 391 


difllua hebridensis, Anticrates 


129 


difformis, Cladius . 


• 131 


diminutus, Pseudococcus 


• 132 


diminutus, Trionymus 


. 262, PI. 14 


dispar, Aethaloptera 95 (fig. 


. 96 




17, 19 


dohrnii, Fenusa 


IQ-21 


dolce, Placonotus . 


192-193 


dolichura, Pontania 


• 390 


dolorosa, Epitola 


40 


donacioides, Placonotus . 


39 


dorothea, Epitola . 


39 


dunia, Epitola 


39 


Dysmicoccus 


221 (fig.), 223 




• 235 


ealaensis, Placonotus 


• 387 


eburneifrons, Tenthredo . 


406 


eichelbaumi, Mariolaemus 


406 


elaphrodes, Schoenotenes 


• 259 


elderi, Dolerus 


• 395 


electrantha, Stathmopoda 


140-141, PI. 3, 4 


elegans, Galideus 


• 392 


elgonensis, Xylophloeus . 


311, 312 (fig.) 


elissa, Epitola 


• 139 


elli, Notolaemus 


140 


elongata, Coccidella 


151-152 


elongata, Morrisonella 


• '51 


elongatulus, Laemophloeus 


131, PI. I, 2 


elongatus, Rhizoecus 


89, 90 (fig.), 91 


elongatus, Xenippoides . 


• 403 


ephippium, Eutomostethus 


203 (fig.), 210 


Epitola 


227 (fig.), 228 


equiseti, Ametastegia 


. 9. 10 (fig.) 


erichsonii, Pristiphora 


268 


erythrocephala, Acantholyda 


• 403 


erythropyga. Serpusilla . 


. 404 


escalerai, Mariolaemus 


8 


euloidea, Neotunga 


126 


Euripersia 


• 394 


europaea, Euripersia 


. 129 


europaea, Ripersia 


• 252 


euryochra, Eucosma 


231-240 


eustropha, Decadarchis 


228-230 


euthycolona, Acrocercops 


23 


Euthymia 


■ 385 


evansi, Cryptolestes 




eversmanni, Loderus 


. 59, 60 (fig.), 61 


excavata, Phyllocolpa 


. 254- PI. 14 


excisus, Pachynematus 


• i3« 


exornatus, Placonotus 



. 127 
311, 313 (fig.), 314 
186 

47 

267 

194 

393 

43 

43 

43- 44 (fig-). 45 

266, PI. 14 

• 393 
61 
61 

96 (fig.), 97 (fig.), 
98, 99 (fig.). 100 

• 392 
178-179 

• 404 
. 152 
. 183 

133. PI- 9. 10 

141-142, PI. 3, 4 

21-24 

181 (fig.), 182 

- 392 
219 

256, PI. 14 

- 389 
264 

297 (fig-). 298 

• 193 
139, PI- 3. 4 

221 (fig.), 222 

- 45 
45 

• 235 

43. 45. 47 (fig). 48 
299 (fig.), 300 

391 

125-157 
390 

- 396 
■ 385 

337 (fig-). 338 

219 

366-377 

24-28 

24, 25 (fig.), 26 

17, 24 

PI. 14 

269, PI. 14 

. 268 

290 

. 240 (fig.) 

- 388 

• 403 
. 406 

186-187 



INDEX 



413 



extensicomis, Pachynematus . . . 406 
exul, Ripersia 26 

falcifer, Rhizoecus . . . -47 
fallax, Amauronematus .... 399 
fasciata, Euthymia . . 291, 292 (fig.) 
fasciata, Mananara 341 (fig.), 342 
fasciatus, Onetes . . . 288, 289 (fig.) 
ferrugineus, Cryptolestes . 234, 235 (fig.) 
ferruginous, Microlaemus 198 (fig.) 
festivus, Amauronematus . 399 
fliicicola, Spilococcus . 54, 55 (fig.), 56 
finoti, Heteracris 319, 320 (fig.) 
Finotina ..... 344-346 
flavens, Brachythops .... 387 
flavistriata, Decadarchis .270 
flavoantennata, Epitola . . . .140 
formicarii, Ripersia 17 

foveolatus, Pacliynematus . 406 

fragilis, Pseudococcus . .40 
freyi, Pachynematus .... 406 
frutetorum, Gilpinia .... 386 
fuliginosus, Stethomostus . -391 
furcata, Paraserpusilla 339 (fig.), 340 
fusca, Pacliyceracris 306 (fig.) 
fuscipes, Arge 387 

gahani, Pseudococcus . . .40 

Galideus ..... 296, 298 
Cannes ..... 174-176 

geniculata, Pristiphora .... 394 
geniculatus, Monophadnoides . .391 

genucinctus, Loderus .... 388 

Gergis 302-303 

Gergisini ...... 300 

gerina, Epitola . . . . .138 

gessneri, Dolerus ..... 388 

ghesquieti, Epitola . . 145, PI. 5, 6 

gigas flavicornis, Urocerus . . 386 

glabra, Serpusilla . 335, 336 (fig.), 337 

glabrata, Ametastegia .... 390 

glacialis, Amauronematus 399 

glabriculus, Passandrophloeus 213, 214 (figs.), 215 
glaucothoe, Strepsicrates .... 253 

granulatus, Laemophloeus .183 

groenlandicus, Amauronematus . 396 

haophila, Ripersia . . . . 41, 47 

halophilus, Coccus ..... 47 
halophilus, Rhizoecus 47, 48 (fig.), 49 

harrisoni, Athripsodes . . . .109 

Iiarrisoni, Leptocerus .109 

hastata, Triaendella . . . .105 

hebes, Amauronematus .... 397 
Heliococcus ..... 28-29 

helleni, Amauronematus . . . 397 

hercyniae, Gilpinia .... 386 

Heteracris . . . . 319 

Heterococcus .... 29-30 

hcterogramma, Decadarchis , , .270 



hewitsoni, Epitola . 
hewitsonioides, Epitola 
hibernicus, Dactylopius 
hibemicus, Pseudococcus 
hibernicus, Trionymus 
Iiirsuta, Paulianacris 
histrio, Amauronematus 
Horaeocerus . 
hova, Leptacris, 
hulteni, Amauronematus 
humeralis, Oxya 
Hvsiella 



ikoya, Epitola 
immoderatus, Gannes 
imperator, Stathmopoda 
improbana, Coscinoptycha 
incisa, Caenognosis 
inermis, Pseudohysiella . 
inflatus, Leptocerus 
inopinatus, Pachynematus 
insulana, Epitola . 
insularis, Euura 
intermedia, Epitola 
intermedia, Hewitsonia 
intermedius, Amauronematus 
interruptus, Phenacoccus 
iridoscliema, Cryptophlebia 
itelmena, Amauronematus 
iturina, Epitola 

janeti, Leptophloeus 
jejuna, Herpystis 
jocosa, Tenthredo . 
juvencus, Sirex 

kamengensis, Epitola 
katerae, Epitola 

katherinae, Epitola 
kirbyi, Pachynematus 
kraussi, Onetes 

laevicornis, Planolestes . 

lamborni, Epitola . 

lamborni orientalis, Neaveia 

lanatae, Euura 

langi, Liptena 

latipes, Pseudococcus 

lativentris, Pristiphora . 

leonina, Epitola 

lepesmei, Cryptolestes 

Leptacris 

leptocephalus, Amauronematus 

Leptophloeus 

leucolaenus, Amauronematus 

leucostoma, Pristiphora . 

liana, Epitola 

llganus, Notolaemus 

linearis, LeptopIiloeu$ 



35 



126, PI. 13 

126 

. 63 

. 63 

. 63 

• 283 (figs.) 

400 

• 324 
277, 278 (fig.) 

■ 397 
309, 310 (fig.) 

284 

140 

175, 176 (fig.) 

264 

■ 263 

• 257 
330 (fig.) 

119, 120 (fig.) 

405 

33, PI. I, 2, 9, 10 
402 
131, PI. I, 2 
. 161 
400 
36 (fig.), 37 
255, PI. 14 

• 396 

142, PI. 9, ID 

203 (fig.), 209 

. 252 

. 392 

. 386 

150-151. PI. 7, 8 
149-150, PI. 7, 8 

• 134 
406 

286-287 

. 228 

. 152 

158, PI. II, 12 

• 403 
. 156 

40 
395 
3. 4 
236 

• 277 
. 399 

201-210 
402 

• 394 
138. PI. 13 

225 
207 



136. PI 
235 (fig) 



414 



INDEX 



livens, Mariolaemus 
lomatographa, Idiophantis 

longicauda, Amauronematus 
longicornis, Mestolaemus 
longispinus, Dactylopius 
longispinus, Pseudococcus 
longispinus var. latipes, Pseudococcus 
Lopheuthymia .... 
lounsburyi, Chorizococcus 
lounsburyi, Pseudococcus 
lounsburyi, Trionymus 
lucidus, Leptophloeus 
luffl, Atrococcus 
luffi, Dactylopius . 
luffi, Pseudococcus 
luteiventris, Eutomostethus 

maculata, Epitola . 
maculipennis, Oecetis in (fig.), 
maculipennis, Plutella 
madagascariensis, Caloptenopsis 
maesi, Primerenca . 
Magnoleptus .... 
maguira, Parasetodes 
maguirus, Leptocerus 
majus, Placonotus . 179 

malacearum, Pseudococcus 
Malagacetrus 

Malagasacris .... 
malagassa, Serpusilla 
malagassum, Spathosternum 
malagassus, Catantops 
mamillariae, Pseudococcus 
Mananara .... 
mara, Epitola 
marcina, Primerenca 
marginata, marginata, Epitola 
marginata umbratilis, Epitola 
Mariolaemus .... 
maritimus, Pseudococcus 
matris, Blubos 
maxima, Aethaloptera 
mendanai, Opogona 
mengoensis, Epitola 153, 

meracula, Yponomeuta 
mercedes, Epitola . 
mesembryanthemi, Coccidella . 
mesembryanthemi, Morrisonella 
mesembryantliemi, Rhizoecus . 
mesnili, Ripersia 
Mestolaemus .... 
mestus, Placonotus 
microglypta, Limnaecia . 
Microlaemus .... 
microphyes, Amauronematus . 
mimosae, Xylophloeus 
minima, Peragrarchls 
minimus, Cryptolestes 
minor, Gergis 303- 

minor, Morondavia 



40 
40 

• 294 
19, 21 

19 
19 

• 205 

II 



391 



218 minutus, Heliococous 29, 30 (fig.) 

. 257, PI. 14 miranda miranda, Epitola . 126, PI. i, 2 

401 miranda vidua, Epitola . . 127, PI. i, 2 

214 (fig.), 2x6 mirus, Laemophloeus (Silvanophloeus) 183 

40 misellus, Mariolaemus 217 (fig.), 218 

mobilis, Leptophloeus .... 208 

mocquerysi, Galideus . . 297 (fig.), 298 

moerens, Pachynematus .... 406 

mollis, Pristiphora ..... 395 

morio, Priophorus ..... 393 

Morondavia ..... 306-307 

moselyi, Triaenodes 103 (fig.), 105, 106 (fig.), 107 
mossus, Placonotus .187 

moyambina, Epitola . . . .130 

mpangensis, Epitola . . 149, PI. 5, 6 

mucronata, Euura ..... 402 

mucunae, Leptophloeus . 206-207 

152 munerata, Hieromantis . . . .263 

112 (fig.), 113 mus, Epitola ...... 144 

267 

311 Narthecius ..... 210-21 1 

98 neglectus, Amauronematus . 400-401 

199-201 nemoratus, Heterarthrus . . 389 

109 Neotunga 365-377 

109 newsteadi, Trionymus . . 61, 62 (fig.) 

(fig.), 1 80-1 8 1 nigra, Epitola . . . 145 

40 nigra, Euura ...... 403 

316 nigeriae, Epitola 145, PI. 5, 6 

300-302 nigricollis, Tenthredo .... 392 

335. 336 (fig.) nigricornis, Horaeocerus . . 324, 325 (fig.) 

278, 279 (fig.) nigricornis, Hysiella . . 284, 285 (fig.) 

327 nigrofasciatus, Seyrigacris . 328, 329 (fig.) 

52 nigrovenata, Epitola 136-137, PI. 3, 4 

340 nipae, Dactylopius . . . -31 

135, PI. 3, 4 nipae, Nipaecoccus . . . .31 

98 nipae, Pseudococcus . . . -31 

130 Nipaecoccus . . . . .31 

130 nitens, Dolerus ..... 389 

216-220 nitide, Epitola ..... 139 

40 nitidipleuris, Amauronematus . . 402 

195 (fig.), 196 njala, Setodes . 116 (fig.), 117 

100 noctilio, Sirex ..... 385 

268 Notolaemus ..... 220-226 

154, PI. 9, 10 nudus, Heterococcus . . . .31 

266, PI. 14 nudus, Phenacoccus .... 31 

147, PI. 5, 6 nuorbinjargi, Amauronematus . 399 

45 

45 obductus, Pachynematus . . . 405 

45 obscura, Epitola . . . . .152 

26 obscura, Xyela ..... 385 

215-216 ochreopyga, Serpusilla 336 (fig.), 337 

179 ochroleucus, Heptamelus . . . 387 

260 ochrophara, Imma . 264, PI. 14 

196-199 ochropus, Arge ..... 387 

396 Oecetis ..... 1 09-11 7 

192 oligospilus, Nematus .... 405 

261, PI. 14 olivacea, Tenthredo .... 392 

240 (figs.) ombrodelta, Cryptophlebia . . . 255 

•304, 305 (fig.) Onetes ...... 284, 288 

307, 308 (fig.) opaculus, Leptophloeus , . 203 (fig.), 205 



INDEX 



415 



orbitalis, Amauronematus 
orestes, Trionymus 

orientalis, Epitola . 
orthomorpha, Polychrosis 
ouesso, Epitola 
ovata, Eriocampa . 
oxyopis, Thiotricha 
oxystoma, Imma 



Pachyceracris 

padi, Aneugmenus . 
pallescens, Monophadnus 
pallidinervis, Gergis 
pallipes, Ametastegia 
pallipes, Priophorus 
pallipes, Pristiphora 
palpalis, Microlaemus 
paludinus, Atrococcus 
paludinus, Pseudococcus 
Pamphagella 

parallelicoUis, Magnoleptus 
parallelus, Leptophloeus 
Paraserpusilla 
parvilabris, Pachyneinatus 
Passandrophloeus . 
patens, Xylophloeus 
Paulianacris . 
Peliococcus 

pelograpta, Peragrarchis . 
penium, Saccharicoccus . 
peregrinus, Pseudococcus 
periclina, Stathmopoda . 
Perineta 

peringueyi, Notolaemus . 
perrieri, Notolaemus 
perrisii, Trionymus 
perrisii, Westwoodia 
perspicuus, Notolaemus . 
phalaridis, Trionymus 
Phenacoccus . 
picinus, Notolaemus 
picinus rhodogaster, Dolerus 
pinodes, Epitola 
pinodoides, Epitola 
piricola, Dysmicoccus 
piricola, Pseudococcus 
Placonotus 
Planococcus . 
Planolestes 

planulatus, Cryptolestes . 
polaris, Amauronematus 
poliophora, Strepsicrates . 
politissimus, Placonotus . 
polychroma, Euthymia . 
polychroma, Pseudoserpusia 
posthumus, Epitola 
praestans, Thisoicetrus 
pratensis, Dolerus . 
pratorum, Loderus 



(Trionymus) 



402 
63, 64 (fig.) 
143. PI. 5, 6 

253 
147, PI. 7, 8 

• 392 

• 257 
. 265, PI. 14 



• 305 

• 387 

• 391 
303, 304 (fig.) 

• 390 
■ 393 

• 394 
198-199 

II, 12 (fig.), 13 
II 

• 311 
200 (fig.), 201 

206 
338-340 

• 405 
213-215 

189 (fig.), 191 
282-283 

31 

261 

50, 51 (fig.), 52 

19 
264 

• 342 

221 (fig.), 224 



63 



, 65. 66 (fig.), 67 

. 63 

222 

67, 68 (fig.) 

32, 39 

226 

388 

143 

152 

23 

23 

176-187 

39 

226-228 

236-237 

. 398 

. 253, PI. 14 

. 183-185 

291, 292 (fig.) 

281 (fig.), 282 

. 128 

• 319 

. 389 

. 388 



pravus, Amauronematus 
problematicus, Leptophloeus 
Proeuthymia . 

propinquus, Amauronematus 
proxima, Pontania 
psarodes, Nesoscopa 
pseudoanticus, Dolerus 
pseudocoactula, Pristiphora 
Pseudococcus 
pseudoconjuncta, Epitola 
Pseudohysiella 
Pseudoserpusia 
pugnaceus, Magnoleptus . 
pulchra, Ischinacrida 
pulveraria, Ripersia 
pulverarius, Heterococcus 
pulverarius, Pseudococcus 
pulverarius, Trionymus . 
pulverulenta, Epitola 
punctatus, Leptophloeus 
punctumalbum, Macrophya 
purpurascens, Aslauga 
pusilla, Fenusa 
pusilloides, Cryptolestes 
pusillus, Cryptolestes 
pygmaeus, Cephus . 



quadrifasciata, Pontania 

quercus, Pristiphora 
quercus, Pseudococcus 

radama, Finotina . 
radicum, Dactylopius 
radicum, Trionymus 
ranavaloae, Finotina 
rapae, Pachyprotasis 
reducta, Heteracris 
reticulatus, Nematus 
reuteri, Pristiphora 
rezia, Phytala 
Rhadinacris . 
Rhizoecus 

rhynchias, Cryptophlebia 
riartus, Notolaemus 
rileyi, Epitola 
rodea, Peragrarchis 
rubripes, Malagacetrus 
rugosa, Malagasacris 
rumicis, Pachynematus 



saarineni, Amauronematus 

sacalava, Catantops 
Saccharicoccus 
sagmarius, Amauronematus 
saUens, Labdia 
sanguinolentus, Onetes 
saussurei, Proeuthymia 
schedli, Narthecius 



• 399 
203 (fig.), 210 

293 

• 399 

• 404 
251, PI. 14 

. 388 

• 395 
39-40 

153. PI- 7. 8 
. 328 
281-282 
ioo (fig.), 201 

• 277 
31 
31 

• 63 

• 63 

• 155 
203 (fig.), 208 

• 393 
. 156 

• 392 
236, 237 (fig.) 

238, 239 (figs.) 
. 386 



• 399 

• 394 

34 

346, 347 (fig.) 

• 47 
69, 70 (fig.), 72 

• 346 

• 393 
323 (fig-) 

• 404 

• 395 
. 156 

• 344 
40-49 

• 256 
221 (fig.), 223 

150 
261 

317 (fig-) 
301 (fig.), 302 

- 405 



402 

• 327 
50-52 

• 399 

• 259 
. 286 

294 (fig-) 
211, 212 (fig.) 



4i6 



INDEX 



schistocercoides, Rhadinacris . 344, 345 (fig.) 
scirpi, Balanococcus 17 

scirpi, Ripersia . 17, 18 (fig.) 

scottae, Amphlpsyche 92 (fig.), 93 (fig.), 94 

scudderi, Onetes . 286, 287 (fig.) 

semibrunnea, Epitola . .129 

semifusca, Decadarchis . . .271 

semilacteus, Amauronematus . 398 

senegalense, Phanostoma 89, 91 

senegalensis, Amphipsyche 85, 86 (fig.), 87 (fig.), 

89 
septemfasciata, Nomadacris . . . 348 

septentrionalis, Amauronematus 
Serpusilla 

sertifer, Neodiprion 
Setodelllna 
Seyrigacris 



sikorai, Heteracris 
sUacea, Autosticha 
similis, Diprion 
similis, Dolerus 
similis ugandae, Hewitsonia 
simulatella, Bathraula 
slades, Microlaemus 
smaragdipes, Eyprepocnemis 
smithiae, Pachynematus 
socialis, Antonina . 
socius, Pseudococcus 
soreuta, Idiophantis 
spartii, Cryptolestes 
spectrum, Xeris 
sphagni, Phenacoccus 
sphagni, Pseudococcus 
Spilococcus 
Spinococcus . 
splendens, Tenebracris 
stationaria, Carposina 
staudingeri, Pristiphora 
staudingeri aequatorialis, Epitola 
staudingeri gordoni, Epitola 
staudingeri staudingeri, Epitola 
stempfleri, Epitola . 
Stempfferia] . 
stenocrobyloides, Catantops 
stenoides, Leptophloeus . 
strateia, Malagasacris 
subcoerulea, Epitola 
subfasciatus, Dolerus 
subfulvida, Aslauga 
sublustris, Epitola . 
subniger, Laemopliloeus . 
subnitens, Amauronematus 
subterranea, Ripersia 
subterraneus, Clinaurococcus 
subtruncatus, Placonotus 

suffusus, Laemopliloeus . 

sulcata, Ardis 
symmorias, Epermenia . 
synclera, Decadarchis 



• 399 
334-335 

. 386 
109 
327-328 
• 321 (fig.) 
. 258, PI. 14 

■ 386 

• 389 
160, PI. II, 12 

■ 259 

• 199 

• 315 
406 

8 

34 

. 258 

232, 233 (fig.) 

. 386 

37, 38 (fig.) 

37 

52-56 

56-58 

318 (fig.), 319 

262 

• 394 
. 135, PI. 13 

• 135 




tabidus, Trachelus . 

tastus, Placonotus . 

tatarica tatarica, Cyrtacanthacris 

tectus, Dolerus 

Tenebracris . 

tenebrosus, Laemophloeus 

tener, Ametastegia 

terrestris, Rhizoecus 

terrestris, Ripersia 

testudinea, Haplocampa 

thomsoni, Profenusa 

thulensis, Trionymus 

tibialis, Nematus . 

tillbergi, Amauronematus 

tomlini, Trionymus 

tomlinii, Euripersia 

tomlinii, Ripersa 

Triaenodes 

triangularis, Tricbiosetodes 

trifurcus, Nematus . 

Trionymus 

triplanetis, Proterocosma 

Truncatophloeus 

tumentia, Epitola . 

turcicus, Cryptolestes 

turneri, Microlaemus 



. 386 

178, 179 (fig.) 

346-348 

389 

318 

198 

390 

47 

47 

394 

392 

69 

405 

398 

72, 73 (fig.), 74 

26, 27 (fig.), 28 

. 17, 24, 26 

103-109 

17, 118 (fig.), H9 

. 399 

58-59 

260 

201 

137, PI. 3, 4 

237 (figs.). 238 

197, 198 (figs.) 



ugandae, Cryptolestes 

ulicis, Pseudococcus 

ulmeri, Ampliipsyclie 

ulmi, Kaliofenusa . 

ulmi, Pseudococcus 

uncata, Triaenodes 

unga, Pontania 

unifasciatus, Xylophloeus 

uniformis, Epitola . 

urania tanganikensis, Epitola 

urania urania Epitola 128- 



vagus, Pachynematus 
vanduzeei, Selandridea 
variana, Stegasta . 
variator, Amauronematus 
varus, Croesus 
velocipes, Oxyethira 
vermiculatus, Laemophloeus 
versicolor, Epitola . 
vestigialis, Loderus 
viduatoides, Amauronematus 
viduatus, Amauronematus 
villosus, Nematus . 
viminalis, Trichiocampus 
vinalli, Epitola 
virginea, Epitola 
viridana radiata, Epitola 
x'iridana viridana, Epitola 
viridescens, Euthymia 
viridis, Rhogogaster 



239 (figs.), 240 

• 34 

88 (fig.). 89 

• 392 

34 

107-109 

402 

189 (fig.), 193 

. 132 

129, PI. I, 2 

129, PI. I, 2, 9, ID 



■ 405 

• 387 
. 257 

• 398 
. 404 

100, lOI (fig.) 

• 235 
. 131 
. 388 

402 
402 

• 404 

• 393 
154. PI- 7. 8 

• 139 
. 143 

142, PI. 5, 6 
291 

■ 392 



walkeri, Dactylopius 
walkeri, Dysmicoccus 
walkeri, Pseudococcus 
Winnipeg, Pristipliora 
wistariae, Dysmicoccus 
wistariae, Pseudococcus 
wiinni, Ripersia 
wiistnei, Brachythops 
wiistnei, Messa 







INDEX 






417 




21 

21, 22 (fig.), 23 

21 


xanthopharella, Caloptilia 
Xenippoides . 
Xylolestes 






. 268 
298-300 
187-189 






• 395 














23-24 
23 


yukonensis, Dolerus 






• 388 






24 

■ 387 

■ 391 


zelza, Epitola 
zolotarevslcyi, Heteracris 


137 


-138 
322 


PI. 9, 10 
(fig-). 323 



"b 



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ADLARD AND SON, LIMITED, 
BARTHOLOMEW PRESS, DORKING