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Full text of "Bulletin of the British Museum (Natural History)"

BULLETIN OF 

THE BRITISH MUSEUM 

(NATURAL HISTORY) 




ENTOMOLOGY 
VOL. XXII 

1968 



THE BRITISH MUSEUM (NATURAL HISTORY) 
LONDON: 1969 



DATES OF PUBLICATION OF THE PARTS 

No. i . . . . -23 February 1968 

No. 2 . . . . . ' . 28 May 1968 

No. 3 . . . . . .28 May 1968 

No. 4 . . . . . .5 May 1968 

No. 5 . . . . .13 September 1968 

No. 6 . . . . . .20 September 1968 

No. 7 ...... 4 October 1968 

No. 8 . . . . . .21 December 1968 



PRINTED IN GREAT BRITAIN 

BY ADLARD AND SON LIMITED 

DORKING, SURREY 







CONTENTS 



ENTOMOLOGY VOLUME XXII 

No. I. The evolution and dispersal of the grassland leafhopper genus Exitianus, 
with keys to the old world species (Cicadellidae : Hemiptera). By 
H. H. Ross 



No. 2. A revision of the genus Microgaster Latreille (Hymenoptera : 

Braconidae). By G. E. J. NIXON 31 



No. 3. An index-catalogue of African Phaneropterinae (Orthoptera : Tetti- 

goniidae). By D. R. RAGGE 73 



No. 4. Hymenoptera from Turkey (Symphyta). By R. B. BENSON 



109 



No. 5. The subgeneric divisions of the genus Bombus Latreille (Hymenoptera : 

Apidae). By O. W. RICHARDS 209 

No. 6. A list of the type-specimens of Libellulidae and Corduliidae (Odonata) 

in the British Museum (Natural History). By D. E. KIMMINS 277 

No. 7. A revision of the genus Aiolopus Fieber (Orthoptera : Acridoidea). 

By D. HOLLIS 307 

No. 8. Revision of the genus Catasarcus Schonherr (Coleoptera : Curculion- 

idae). By R. T. THOMPSON 357 



Index to Volume XXII 



455 



THE EVOLUTION AND DISPERSAEW^ 

OF THE GRASSLAND LEAFHOPPER 

GENUS EXITIANUS, WITH KEYS 

TO THE OLD WORLD SPECIES 

(CICADELLIDAE : HEMIPTERA) 



H. H. ROSS 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 22 No. i 

LONDON; 1968 



THE EVOLUTION AND DISPERSAL OF THE 

GRASSLAND LEAFHOPPER GENUS EXITIANUS, 

WITH KEYS TO THE OLD WORLD SPECIES 

(CICADELLIDAE : HEMIPTERA) 



BY 



H. H. ROSS 

>~*\ 

Illinois Natural History Survey, Uft>a 



Pp. 1-30 ; 82 Text-figures 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 22 No. i 

LONDON: 1968 



THE BULLETIN OF THE BRITISH MUSEUM 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

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

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

This paper is Vol. 22, No. i of the Entomological 
series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



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



Trustees of the British Museum (Natural History) 1968 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 23 February, 1968 Price Thirteen Shillings 



THE EVOLUTION AND DISPERSAL OF THE 

GRASSLAND LEAFHOPPER GENUS EXITIANUS, 

WITH KEYS TO THE OLD WORLD SPECIES 

(CICADELLIDAE : HEMIPTERA)* 



By H. H. ROSS 

CONTENTS 

Page 
INTRODUCTION ........... 3 

EXITIANUS BALL 4 

KEY TO GROUPS AND OLD WORLD SPECIES MALES .... 4 

Nanus-Group ........... 7 

Obscurinervis-Group ......... 8 

Distanti-Group .......... 8 

Taeniaticeps-Group .......... 9 

Okahandia-Group .......... 16 

UNPLACED SPECIES .......... 21 

SPECIES TRANSFERRED TO OTHER GENERA . . . . . . 21 

PROBLEMS OF SPECIES RECOGNITION ....... 22 

PHYLOGENY ........... 24 

PALAEODISPERSAL . . . . . . . . . .28 

ACKNOWLEDGMENTS . . . . . . . . . .28 

REFERENCES ........... 29 



SYNOPSIS 

Keys and illustrations are presented for the identification of the five species-groups of 
Exitianus and for the identification of the species occurring in the Old World, the latter including 
seventeen species, nine new to science. Synonymy is included for the described Old World 
species plus a brief synopsis of their distribution. The phylogenetic relationships are adduced 
for the world fauna. When geographic distribution is compared with the postulated family tree 
there are strong indications that the genus evolved primarily in Africa with an ancient dispersal 
to the Americas and more recent dispersals to the Oriental and Australian regions. 

INTRODUCTION 

AMONG the most widespread and frequently the most abundant species of grassland 
leafhoppers are the moderately large tawny forms comprising the genus Exitianus, 
a member of the subfamily Deltocephalinae. This genus differs from related forms 
in having only a small number of apical, stout setae on the male pygofer ; the female 
has a relatively long ovipositor extending conspicuously beyond the last dorsal 
segment. These characters are shared only by the presumably sister genus Nepho- 
tettix, readily differentiated from Exitianus by having the crown sharply ridged 
where it meets the face, and in being opaque green with various black markings. 

* This work was supported in part by a research grant from the National Science Foundation, U.S.A. 

ENTOM. 22, I. 1 



4 H. H. ROSS 

Based on similarities of male and female genitalia, Exitianus and Nephotetiix are 
probably most closely related to Doratum and its allies; the entire complex can be 
segregated as a grass-feeding tribe, the Doraturini. 

On every continent at least one common form of Exitianus has a conspicuous 
black band across the top of the head. So similarly coloured are these forms that 
considerable difference of opinion has existed concerning the number of species 
involved. The same difference has been expressed concerning the number of 
species occurring on the same continent. DeLong & Hershberger (1947) segre- 
gated eleven American species on the basis of colour and the number and position 
of the large pygofer spines. Linnavuori (1959) added a few more segregates from 
South America. No one appears to have undertaken a study of the Old World 
fauna of this genus. An opportunity in 1964 to examine types of the genus at the 
British Museum (Natural History) and Stockholm and to see large series of specimens 
from Africa, India, and Madagascar led me to attempt such a study. As new 
characters and species were found, the possibility became evident of reconstructing 
the phylogeny of the entire genus with a high degree of probability. 

This paper treats the genus Exitianus in the following topical order : (i) a diagnosis 
and description of the world groups and the Eastern Hemisphere species, (2) a 
discussion of the peculiar problems encountered concerning species distinctions in 
the Eastern Hemisphere populations, (3) a derivation of the phylogeny of the genus, 
and (4) an application of these hypotheses to the question of its palaeodispersal. 

This study is based entirely on the male sex. In several species, associated 
females are known but they have proven difficult to diagnose. Sexual dimorphism 
in colour is common, females of certain indubitably different species appear identical, 
and certain females that appear distinctive have as yet not been associated with 
males. In the face of these circumstances it appears that our present knowledge 
of Exitianus females does not lend itself to either keys or diagnoses of worthwhile 
predictive value. 

EXITIANUS Ball 

Exitianus Ball, 1929 : 5. Type-species: Cicadula obscurinervis Still. 
Mimodrylix Zachvatkin, 1935 : 108. Type-species: Athysanus capicola Stal. 

Body usually about 4-5 mm. long. Colour chiefly tawny with various patterns of brown or 
black. Head only moderately produced (Text-figs. 1-14), the crown rounding into the face. 
Wings elongate and having typical leafhopper venation. Male pygofer having from two to 
six large dark or black spines along apical margin and only one or two other conspicuous spines. 
Aedeagus having an articulation between shaft and base, the shaft relatively simple in structure. 
Female ovipositor extending a considerable distance beyond abdominal tergites. 

Distribution. Recorded from all continents except Antarctica. 

KEY TO GROUPS AND OLD WORLD SPECIES MALES 

i Plate short and ovoid, without lateral setae (Text-figs. 54, 55) 

okahandia-group 18 
Plate elongate-triangular, with a row of long, stout lateral setae extending the 

length of the plate (Text-fig. 78) 2 



THE OLD WORLD EXITIANUS 



2 (i) Pygofer having four to six apical brown or black spines (Text-fig. 15) 

nanus-group 
Pygofer having two or three apical brown or black spines (Text-figs. 19, 21) . 

3 (2) Dorsal margin of aedeagus with only a slight, irregular concavity at apex 

(Text-fig. 20) ...... . obscurinervis-group 

Dorsal margin of aedeagus with a large arcuate concavity at apex (Text-fig. 22) 

4 (3) Spine 2 of pygofer as long and slender as spine i (Text-fig. 21) 

dist anti-group 

Spine 2 of pygofer much shorter than and thicker than spine i (Text-fig. 24) 

taeniaticeps-group 

NANUS-GROVP 



7 (4) 



5 (2) Known only from one widespread Old World species 

6 (3) 



minus (p. 7) 



OBSC URINER F/S-GROUP 

Known from several New World species not treated further here. For diag- 
nostic treatments see DeLong & Hershberger (1947) an d Linnavuori (1959). 



DISTANTI-GROVP 

Known only from one African species 





distanti (p. 8) 







FIGS. 1-6. Head, pronotum and mesonotum of Exitianus spp. i, E. nanus (India); 
2, same, dark form (Okahandia, Rhodesia); 3, same, light form (Okahandia, Rhodesia); 
4, E. okahandia (holotype) ; 5. E. turneri (holotype); 6, E, natalensis (holotype). 



H. H. ROSS 



TA EN I A TICEPS- 

8 (4) Socket of spine 2 of pygofer almost contiguous with that of spine i (Text-fig. 

23) ; head with extensive dark markings (Text-fig. 13) . kilitnanus (p. 16) 

Socket of spine 2 a considerable distance from that of spine i (Text-fig. 24) ; 

head variously marked .......... 9 

9 (8) Spine 2 of pygofer only slightly asymmetrical (Text-figs. 64, 65) ; aedeagal shaft 

tapering almost uniformly from base to gonopore (Text-fig. 33) 

attenuatus (p. 10) 

Spine 2 of pygofer markedly twisted or bent at tip (Text-figs. 24, 26) ; aedeagal 
shaft either parallel-sided, or enlarged near gonopore, or markedly con- 
stricted at base of dorsal points . . . . . . . . 10 





8 





10 





12 




13 




14 



FIGS. 7-14. Head, pronotum and mesonotum of Exitianus spp. 7, E. picatus (Mexico; 
8, E. exitiosus (Illinois) ; 9, E. indicus (China) ; 10, E. indicus (holotype) ; n, E. coronatus 
(holotype); 12, E. ootacamundus (holotype); 13, E. kilimanus (holotype); 14, E. greens- 
ladei (paratype). 



10 (9) Shaft of aedeagus as thick as in Text-figs. 25, 28, 30, 31, its paired dorsal 

points usually well separated at base as in Text-fig. 27, its apex long, 
slender and markedly curved ......... 1 1 

Shaft of aedeagus thinner (Text-figs. 34-36, 39-51), or its dorsal points nearly 

contiguous at base .......... 13 

11 (10) Dorsum of head yellow with rigidly defined black spots as in Text-fig, n 

coronatus (p. 14) 
Dorsum of head otherwise (Text-figs. 10, 12) ...... 12 

12 (n) Dark markings on top of head as extensive as in Text-fig. 12 

ootacamundus (p. 14) 
Dark markings on top of head chiefly a single, arcuate, dark line (Text-fig. 10), 

sometimes with additional light brown suffusions . . indicus (p. 12) 

13 (10) Dorsal profile of shaft with a slight notch on which dorsal points are situated 

(Text-figs. 34-36, 39-42, 47) 14 



THE OLD WORLD EXITIANUS 7 

Dorsal profile of shaft with no indication of a notch where dorsal points are 

situated (Text-figs. 43-46, 48-51) ........ 16 

14 (13) Profile of aedeagal shaft markedly angulate below gonopore, gonopore opening 

very long (Text-fig. 34) . . .... pondus (p. n) 

Either profile of aedeagal shaft rounded below gonopore (Text-figs. 39-42) or 

gonopore opening much shorter (Text-figs. 35, 47) ..... 15 

15 (14) Either aedeagal shaft slightly to moderately flared before gonopore (Text-figs. 

39-41) or dorsal points close to base of shaft (Text-fig. 42) taeniaticeps (p. n) 
Aedeagal shaft not flared and dorsal points as far from base of shaft as shown 

in Text-figs. 35, 36, 47 Jrontalis (p. 12) 

16 (13) Head with extensive dark markings (Text-fig. 14) . . greensladei (p. 16) 

Head with only a dark arcuate band, as in Text-fig. 10 . . . . 17 

17 (16) Profile of aedeagal shaft with lateral ridge close to margin and apical excava- 

tion forming an obtuse angle with body of shaft (Text-figs. 48-51) 

plebeius (p. 15) 

Profile of aedeagal shaft with lateral ridge closer to central line of shaft and 
apical excavation forming nearly a right angle with body of shaft (Text- 
figs. 43-45) ......... rnucronatus (p. 14) 

OKA HA NDIA -GROUP 

1 8 (i) Apex of pygofer forming an upturned, fingerlike tip bearing one extremely 

stout, dorsally projecting spine and sometimes another much more slender 
spine (Text-figs. 60, 63) ; apex of style scarcely widened (Text-fig. 58) . . 19 
Apex of pygofer wider, not upturned, and bearing two fairly stout spines 

(Text-figs. 52, 56); apex of style greatly widened (Text-figs. 54, 55) . . 20 

19 (18) Apex of pygofer bearing only one spine, this spine wider than pygofer tip 

(Text-fig. 63) ; profile of aedeagus wider (Text-fig. 62) . zuluensis (p. 18) 

Apex of pygofer bearing a slender seta mesad of large one, the latter narrower 
than pygofer tip (Text-fig. 60); profile of aedeagus narrower (Text-fig. 59) 

natalensis (p. 20) 

20 (18) Style markedly shorter than plate (Text-fig. 55); costal area of front wing not 

forming a conspicuous light stripe ..... okahandia (p. 17) 

Style projecting beyond plate (Text -fig. 54) ; costal area of front wing forming 

a conspicuous light stripe ....... turneri (p. 17) 



NANUS-GROVP 

Exitianus nanus (Distant) comb. n. 
(Text-figs, i, 2, 3, 15, 16, 17, 18, 76) 

Athysanus nanus Distant, 1908 : 345. India. 
Athysanus insularis Distant, 1909 : 47. Amirante Is. syn. n. 
Athysanus simillimus Matsumura, 1914 : 185. Formosa, syn. n. 
Euscelis vulnerans Bergevin, 1925 : 42. Southern Algeria, syn. n. 

This is the most widespread Old World species of the genus, occurring throughout 
Africa, Asia Minor, southern Asia, Formosa, islands in the Indian Ocean, and 
Australia. The dorsal head markings may be faint or form dark spots or crescentic 
bars. Well-marked specimens show considerable geographic variation. In speci- 
mens from Asia, the dark markings usually form a solid curved arc interrupted on 
the meson (Text-fig, i, as in the type of nanus) ; specimens from the southern half 



8 H. H. ROSS 

of Africa invariably have each half of the mark either narrowed in the middle (Text- 
fig. 2) or completely pinched off into a pair of black spots (Text-fig. 3). In specimens 
from Asia Minor and North Africa, the dark marks intergrade between the two 
darker types and occasionally these intergradations are encountered in Asian 
material also. 

The type male of insularis (Text-fig. 15) has pygofer spines 2-4 relatively slender ; 
males from Tanzania have them slightly shorter and more robust (Text-fig. 17); 
specimens from Johannesburg, Republic of South Africa, have a mixture of slender 
and stouter spines on each specimen, and sometimes five or six spines on a side 
(if so, usually not the same number on both sides). The spination of the pygofer 
therefore mirrors the variation found in the colour pattern. 

I have studied the types of nanus Distant and insularis Distant but not those of 
simillimus Matsumura and vulnerans Bergevin. The latter two species I am placing 
on the basis of illustrations by Ishihara (1954, fig. 5) and illustrations in the original 
description, respectively. 

The female seventh sternite is characterized by the bi-emarginate apical margin 
and the slightly bowed mesal area (Text-fig. 76). 

OBSCURINERVIS-GROVP 

(Text-figs. 7, 8, 19, 20) 

The members of this American group of species are remarkably homogeneous in 
shape of aedeagus and pygofer, and in spines i and 2 of the pygofer. In all members 
spine 3 is absent, in most members spine 4 is present, and in occasional individuals 
of the latter species an additional spine 5 may be present on one or both sides of 
the capsule. The females exhibit a considerable number of inter-specific differences 
in the shape of the apical margin of the seventh sternite. 

DeLong & Hershberger (1947) and Linnavuori (1959) deal with eleven and seven 
American species, respectively. I am making no attempt to correlate the two 
accounts pending the availability of longer tropical series needed to establish the 
associations of the two sexes and possible synonymy. 



Exitianus distant! sp. n. 
(Text-figs. 21, 22) 

<J. Length 4 mm. Colour of dorsum pale greenish yellow, face the same but with arcuate 
dark lines, venter of thorax and abdomen almost black, legs with coxae black, the remaining 
segments tawny with dark streaks. Dorsum of head proportioned as in Text-fig. 9, without 
any trace of arcuate banding. General structure typical for genus. Pygofer having only 
setae i and 2, both of them straight and slender (Text-fig. 21). Aedeagus with shaft elongate, 
slender, its apex arcuately incised, as seen from lateral view, and the shaft bearing a pair of 
dorsal projections below middle (Text-fig. 22). 

Holotype <$ (in glycerine). [TANZANIA] Tanganyika: Ilonga, 28. vi. 1962 (L. 
Robertson). In B.M.(N.H.). 



THE OLD WORLD EXITIANUS 



Paratype. [TANZANIA] Tanganyika: Ukirigiru, i 
cerine), 27. iv. 1961, light trap. In B.M.(N.H.). 



(pinned, abdomen in gly- 



The two specimens have been preserved in alcohol, and the greenish cast may be 
due to the action of that preservative. This species forms a unique group all of 
its own, characterized by the possession of only spines I and 2 on the pygofer, and 
the long and slender spine 2 that is much like spine i. In all other known species 
having only spines i and 2, spine 2 is much thicker and more densely pigmented 
than spine i. 




22 



FIGS. 15-22. Male genital parts of Exitianus spp. 15, 16, E. nanus (holotype of insularis] ; 
17, 18, E. nanus (Tanzania); 19, 20, E. exitiosus (Illinois); 21, 22, E. distanti (holotype); 
15, 17, 19, 21, pygofer; 16, 18, 20, 22, aedeagus, lateral aspect. 



TAENIATICEPS-GROUP 

In this group the aedeagus has a well-defined preapical notch, a sharp, curved apex, and a 
pair of small, dorsal, sclerotized points; the pygofer usually possesses only spines i and 2, of 
which spine i is slender and elongate, and spine 2 is much shorter, black, and more or less 
twisted at the tip (Text-figs. 23, 24, 64, 65). In most species the vertex of the head has only 
a single transverse-arcuate black band (Text-fig. 9), often with suffusions of paler brown (Text- 
fig. 10) ; in a few the vertex is darker (Text-fig. 12) or has a different pattern of banding (Text- 
fig, n). 

In those species for which the two sexes are associated (indicus, taeniaticeps, 
frontalis, and plebeius), the female eighth sternite has a narrow, V-shaped apical 
incision bordered with darker colouring (Text-fig. 69). In the case of mucronatus, 



ENTOM. 22, I. 



' 



io H. H. ROSS 

pondus, and several other species, females of the same type were present from the 
same or nearby localities. It seems likely that females of all members of this group 
share this distinctive configuration of the eighth sternite. The unassociated female 
type of capicola (Stal) (Text-fig. 70) can therefore at present be assigned to this 
group but not definitely to any species. As explained below under indicus Distant, 
geographic location gives a clue as to the possible identity of fusconervosus (Mots.). 




FIGS. 23-32. Male genital parts of Exitianus spp. 23, E. kilimanus (holotype); 24, 25, 
E. ootacamundus (holotype) ; 26, 27, 28, E. indicus (type series) ; 29, 30, E. indicus 
(Ceylon); 31, 32, E. coronatus (holotype). 23, 24, 26, 29, 32, pygofer; 25, 28, 30, 31, 
aedeagus, lateral aspect; 27, aedeagus, posteroventral aspect. 



Exitianus attenuatus sp. n. 

(Text-figs. 33, 64, 65, 66) 

<J. Length 4-5 mm. Colour above and below tawny, with a thin arcuate black line across 
dorsum of head (as in Text-fig. 9) and frontal area of head and legs with small dark markings. 
General structure typical for genus. Pygofer with spines i and 2, or i, 2 and 3; spine i is long 
and slender, spine 2 or spines 2 and 3 are shorter, thicker, darker and very slightly asymmetrical 
at apex (Text-figs. 64, 65). Aedeagus with shaft tapering almost uniformly from base to 



THE OLD WORLD EXITIANUS n 

gonopore, the portion beyond that slender and arcuately incised (Text-fig. 33) ; near the base 
of the shaft are a pair of dorsal points or projections that are finger-like, flared laterally, the 
two well separated at the base (Text-fig. 66). 

Holotype $ (pinned, genitalia in glycerine). SOUTH AFRICA: Cape Province, 
Somerset East, xi.i93o (R. E. Turner}. In B.M.(N.H.). 

Paratype. Same data, i <$ (pinned, genitalia in glycerine). In B.M.(N.H.). 

Both type specimens have the pygofer spines somewhat damaged, but the remain- 
ing evidence indicates several interesting items: (i) spine i is long and slender; 
(2) spine 2 is only slightly asymmetrical (Text-figs. 64, 65), not approaching the 
bent condition (Text-figs. 23, 24, 26) of related species; and (3) the genetic control 
for spine 3 has not been completely lost. The latter two points warrant a serious 
consideration of this species as the most primitive of its group. 

Exitianus pondus sp. n. 

(Text-figs. 34, 67) 

cj. Length 5 mm. Colour tawny except for a fine dark arcuate line across anterior portion 
of vertex and a few dark marks on the face and legs. General structure typical for genus. 
Pygofer with spines i and 2 only, spine i long and slender, spine 2 deeper, shorter, almost jet 
black, and markedly asymmetrical (Text-fig. 67), the extreme tip slightly but definitely hooked 
ventrally. Aedeagus (Text-fig. 34) with profile of shaft wide at base and tapering slightly to 
gonopore, the apex with an arcuate incision occupying the apical third of the shaft and forming 
a long narrow apex sharply hooked at tip; near the base of the shaft is a prominent pair of 
dorsal points. 

Holotype <$ (pinned, abdomen in glycerine). S. AFRICA: Pondoland, Port St. 
John, 1-9. vii. 1923 (R. E. Turner). In B.M.(N.H.). 

Paratypes. Same data, 2 $. In B.M.(N.H.). 

Three specimens that resemble the above types very closely but lack dorsal points 
at the base of the shaft are tentatively considered as this species but not included 
in the types, as follows: S. AFRICA: Zululand, Gingindhlovu, 2 <$, 15. vi. 1926 (R. E. 
Turner)', MADAGASCAR: Morondova, forest south of Befasy, i <$, 1.1956 (R.P.). 

Exitianus taeniaticeps (Kirchbaum) 

(Text-figs. 37, 39-42, 71) 
Athysanus taeniaticeps Kirchbaum, 1868 : 87. Messina, Syracuse. 

I have not seen the type of this species, but the description fits perfectly the 
species here considered as taeniaticeps, which is the only species of the group so far 
identified from the circum-Mediterranean countries. Specimens from Iraq, Lebanon, 
Cyprus, France, and Libya are remarkably uniform, in contrast with other popula- 
tions in the species, in having the dorsal points some distance from the base of the 
shaft (Text-figs. 39-41). Males from Rhodesia, considered as this species, usually 



12 H. H. ROSS 

have these points almost at the base of the shaft (Text-fig. 42). Large series of 
males from central Sudan are much more variable in this character, completely 
bridging the morphological gap between the Mediterranean and Rhodesian popu- 
lations. 

This situation suggests that the Mediterranean and South African populations 
represent previously isolated segments of an ancestral population that became 
morphologically different, then were brought together by changing ecological con- 
ditions and have since formed an intermingled, freely-hybridizing population in at 
least the Sudan. 

In a few Rhodesian males the apico-central corner of the pygofer is almost square 
(Text-fig. 37), in contrast to the sinuate condition normally found throughout other 
populations of the species (as indicated by the broken line in Text-fig. 37). 

The illustrations of Ribaut (1952 : 138, figs. 245-249) undoubtedly refer to this 
species, although pygofer spine 2 is shown as straight rather than contorted at the 
tip. It should be noted that in Fig. 248 the shaft is folded into the socket so that 
the latter hides the real base of the shaft and the dorsal points of the shaft therefore 
erroneously appear to be basal. 

Exitianus frontalis (Distant) comb. n. 
(Text-figs. 35, 36, 47) 

Athysanus frontalis Distant, 1917 : 317. Seychelles. 

An extremely close relative of taeniaticeps , this species as here recognized is 
diagnosed by the uniformly thick shaft (Text-figs. 35, 36) in contrast with the slightly 
to markedly clavate shaft of taeniaticeps (Text-figs. 39-42). In frontalis the dorsal 
points of the shaft are some distance from the shaft's base, as sometimes occurs in 
Mediterranean specimens but not in South African specimens of taeniaticeps, sug- 
gesting that in the latter species there has been a displacement of this character 
in the southern part of Africa where both taeniaticeps and frontalis occur in the 
same area. 

To date this species has been found in the Seychelles, Madagascar (Diego-Suarez, 
Montagne des Francais), and South Africa (Pondoland). It is interesting that the 
three specimens from Pondoland exhibit almost exactly the same range of variation 
in the aedeagal shaft as found in the two specimens illustrated from Madagascar. 
The male from the type series (Text-fig. 47) is a slender extreme, matched almost 
perfectly by a specimen from Pondoland in the Turner collection, B.M.(N.H.). 
I have studied the female type, in which the eighth sternite is notched as in other 
species of the group. 

Exitianus indicus (Distant) comb. n. 
(Text-figs. 9, 10, 26-30, 69) 

PAthysanus fusconervosus Motschulsky, 1863 : 97. Ceylon. 
Athysanus indicus Distant, 1908 : 344. India. 
Athysanus atkinsoni Distant, 1908 : 345. India, syn. n. 



THE OLD WORLD EXITIANUS 



33 34 35 




FIGS. 33-51. Male genital parts of Exitianus spp. 33, E. attenuatus (holotype); 34, E. 
pondus (holotype) ; 35, 36, E. frontalis (Madagascar) ; 37, E. taeniaticeps (Rhodesia) ; 
38, E. frontalis (type series) ; 39, 40, 41, E. taeniaticeps (Tripoli) ; 42, the same (Rhodesia) ; 
43, 44, 45, E. mucronatus (holotype and two paratypes, respectively) ; 46, E. greenslactei 
(holotype) ; 47, E. frontalis (type lot) ; 48, E. plebeius (Queensland) ; 49, the same (para- 
type of plebeius}; 50, the same (Samoan Is.); 51, the same (homeotype of norrisi). 37, 
38, pygofer; all others, aedeagus or shaft only, lateral aspect. 



i 4 H. H. ROSS 

This is a pale species having a faint or thin, dark, transverse, arcuate band on the 
vertex. It has been taken at several localities in India (in great abundance at 
Kharagpur), Ceylon, Formosa, and Japan. 

I have studied the female types of both Distant names. These are in B.M.(N.H.), 
bearing identical data 'Calcutta, Atkinson Col. 92.6.' The common type series of 
the two species contains several males that demonstrate the characters of this sex 
(Text-figs. 26-28). 

There is a possibility that fusconervosus Motschulsky, described from the summit 
of Mt. Patannos, Ceylon, may apply to this species, which is the only one of the 
genus thus far recorded for the island, but the unavailability of the type mitigates 
against the use of its carrier name at the present time. Ishihara's illustrations of 
E. capicola (Stal), listed under Mimodrylix Zakhvatkin, undoubtedly apply to this 
species (Ishihara, 1954 : 6). 

Exitianus coronatus (Distant) comb. n. 
(Text-figs, n, 31, 32) 

Phrynomorphus coronatus Distant, 1918 : 52. India. 

This species is remarkably like indicus in morphological details of the male genitalia 
(Text-figs. 31, 32) but differs markedly in the dorsal spotted pattern of the head 
(Text-fig, n). 

The species is known only from the type, which I have studied. It is in B.M. 
(N.H.) and was collected at Kumaon, N.W. India. 

Exitianus ootacamundus (Distant) comb. n. 

(Text-figs. 12, 24, 25) 
Phrynomorphus ootacamundus Distant, 1918 : 51. India. 

As is true for the above-mentioned species, this one also is practically identical 
in details of male genitalia (Text-figs. 24, 25) with indicus, differing primarily in 
having a much darker head (Text-fig. 12). 

To date the only known record is the type, which I have studied. It is in B.M. 
(N.H.) and was collected at Ootacamund, South India. 

Exitianus mucronatus sp. n. 

(Text-figs. 43, 44, 45) 

$. Length 4-5 mm. Colour pale tawny except for a black arcuate band across the vertex, 
much as in Text-fig. 9, and a few pale marks on face and legs. General structure typical 
for genus. Pygofer with spines i and 2 only, spine i very long slender, spine 2 much shorter, 
asymmetrical, and slightly hooked at apex as in Text-fig. 37. Aedeagus with shaft moderately 
long and thick (Text-figs. 43-45), with an apical arcuate incision resulting in a sharp hooked 
tip; the dorsal points are minute and their bases do not interrupt the even contour of the 
dorsal surface. 



THE OLD WORLD EXITIANUS 15 

Holotype < (pinned, abdomen in glycerine). MADAGASCAR: Ft. Dauphin (R.P.). 
In collection of Madagascar Scientific Institute. 

Paratypes. MADAGASCAR: Ft. Dauphin, Tananarive, Bas Mangoky, Nosivola, 
Perinet, Monronova forest south of Befasy, Montagne des Francais and Andova, 
28 $. In collections of Madagascar Scientific Institute, Illinois Natural History 
Survey and B.M.(N.H.). 

This series of specimens appears to represent a distinctive and perhaps an endemic 
Madagascar species differing from plebeius in having a stouter aedeagal shaft whose 
ventral lateral ridges curve dorsally and disappear toward the apex. In Madagascar, 
mucronatus appears to displace taeniaticeps almost completely, although the latter 
species appears to be abundant in Madagascar at Mohali, lac Iconi, and one specimen 
has been taken from Rte de Fianarantsoa, Km. 334. These specimens of taeniaticeps 
are like the more massive Rhodesian type rather than the narrower type found 
around the Mediterranean. 



Exitianus plebeius (Kirkaldy) sp. rev., comb. n. 

(Text-figs. 48-51, 68) 

Nephotettix plebeius Kirkaldy, 1906 : 331. Queensland. 
Eutettix norrisi Evans, 1938 : 14. West Australia, syn. n. 
Euscelis transversus Metcalf, 1946 : 122. Guam. syn. n. 

This species has the same tawny colour and black arcuate stripe on the vertex 
as is found in indicus and several other species of the genus (as in Text-fig. 9). It 
is most closely related to frontalis, from which it differs in lacking a notch on the 
dorsal profile of the shaft where the dorsal spines arise (Text-figs. 48-51), and to 
E. mucronatus, from which it differs in the longer and more slender aedeagal shaft. 

The holotype $ and paratype <$, which I have examined, are from Cairns, Queens- 
land, Australia, deposited in the Bernice P. Bishop Museum, Honolulu. Both 
appear to be the same species and I am herewith designating the ^ paratype as the 
allotype. I have also examined the <$ holotype of transversus, which agrees exactly 
with males from Australia. 

Concerning norrisi, I am following the synonymy of Evans (1966). To date the 
only records of this species I have personally examined are from Australia, Guam, 
and Samoa. The records of capicola cited by Linnavuori (1960) from Fiji are based 
on females, but probably refer to this species. Previously this species has been 
confused with other species in the Old World having the same colour, and the 
Australian records have been labelled as capicola. The distinct, long shaft and the 
inconspicuous dorsal aedeagal spines indicate, however, that this species is different 
from those occurring on other continents, 



[6 H. H. ROSS 

Exitianus greensladei sp. n. 

(Text-figs. 14, 46, 78) 

<J. Length 4 mm. Background colour tawny with many dark areas; vertex with a wide 
black arcuate band covering most of the area, face almost entirely black, the facial marks 
running up slightly over the edge of the vertex; exposed thorax with heavy black spots (Text- 
fig. 14); legs and venter of body with irregular dark marks; membrane of wings almost hyaline, 
the veins bordered by dark brown striping. General structure typical for genus. Pygofer 
with only spines i and 2, spine i long and slender, spine 2 shorter, thicker, darker, and slightly 
hooked at apex, as in Text-rig. 24. Aedeagus with shaft long and slender (Text-fig. 46), 
the dorsal points minute and their bases below the dorsal contours. 

Holotype <$ (pinned, abdomen in glycerine). SOLOMON ISLANDS: Guadalcanal, 
Kau Kan, 17. xi. 1962, no. 2924 (P. Greenslade). In B.M.(N.H.). 

Paratype. Same data, i < (pinned, abdomen not cleared). In B.M.(N.H.). 

This heavily marked species is a close relative of plebeius, which it resembles in 
structure of genitalia and from which it differs primarily in the dark colour pattern. 
The two specimens of the type series are both markedly smaller (4 versus 5 mm.) 
than any specimens of plebeius I have yet seen. 



Exitianus kilimanus(]a.cobi) comb. n. 

(Text-figs. 13, 23) 
Phrynomorphus kilimanus Jacobi, 1910 : 131. Kilimandjaro, Africa. 

This species is readily distinguished by the twisted spine 2 of the pygofer and the 
very close proximity of the bases of spines i and 2 (Text-fig. 23). The head and 
pronotum are heavily marked with black (Text-fig. 13), much as in greensladei. 
The type < (which I have seen) bears the collection data: Meru, 3,000-3,500 m 
Jan. (Sjostedt). Another female from the same locality on Mount Kilimandjaro 
bears a type label but a second female bearing a red 'Allotypus' label is evidently 
another species. I was not in the Stockholm Museum long enough to clear the 
genitalia of the type but the diagnostic position of the pygofer spines was readily 
visible on the dried specimen. 

OKAHANDIA-GROUP 

Among the extensive grass-inhabiting species collected in Africa by R. E. Turner, 
I found four species that form a distinctive group of Exitianus in which the plates 
are short and lack lateral spines, and the styles have the apex truncate and projecting 
nearly posteriorly, rather than being sharp and curved sharply laterally (Text-fig. 
82) as is true of all other members of the genus. 



THE OLD WORLD EXITIANUS 17 

Exitianus okahandia sp. n. 

(Text-figs. 4, 55, 56, 57, 72) 

$. Length 4 mm. Colour tawny with the following black marks: a fairly narrow arcuate 
band on vertex, small black spots on pronotum, several black spots on scutellum (Text-fig. 
4), plus markings on the face, legs, and venter. General structure typical for genus. Plates 
ovate, without lateral spines, and about as long as wide (Text-fig. 55), extending considerably 
beyond apex of styles. Styles with apex broadened and truncate (Text-fig. 55). Pygofer 
of usual shape for genus, with only spines i and 2, both symmetrical, spine 2 shorter, thicker, 
and darker than spine i (Text-fig. 56). Aedeagus moderately long, its apex with only a slight 
and inconspicuous incision (Text-fig. 57). 

$. Length 5-5 mm. Colour and general structure similar to male. Apex of eighth sternite 
slightly produced in middle, the segment often with a small dark spot on each side near the 
middle of the lateral half of the segment (Text-fig. 72). 

Holotype $ (pinned, abdomen in glycerine). SOUTH AFRICA: Natal, Van Reenen, 
Drakensberg, 1-22.1.1927 (R. E. Turner}. In B.M.(N.H.). 

Allotype $. SOUTHWEST AFRICA: Okahandja, 10-16.11.1928 (R. E. Turner). 
In B.M.(N.H.). 

Paratypes. Same data as allotype $, 3 <$, 8 $, with inclusive dates 27.i-i8.iii. 
i (, i $ in Illinois Natural History Survey collection, remainder in B.M.(N.H.). 

Additional females that appear to belong to this species are from SOUTH AFRICA: 
Cape Province, Queenstown ; Transkei, Untata. [TANZANIA] Tanganyika : Ukiriguru. 



Exitianus turneri sp. n. 

(Text-figs. 5, 52, 53, 54, 73) 

cJ. Length 4-5 mm. Colour tawny, head with a black arcuate band slightly interrupted 
at middle (Text-fig. 5), face with a narrow mesal stripe in addition to more lateral markings; 
legs with dark marks; tegmina with all veins pale brown except costal margin which is con- 
trastingly cream coloured. General structure typical for genus. Pygofer moderately narrow 
at apex, which bears spines i and 2, both long and symmetrical, spine 2 thicker and shorter 
than spine i and with a marked crease at its base (Text-fig. 52). Plates short and obliquely 
truncate, without large lateral spines (Text-fig. 54). Style with flared apex, extending slightly 
beyond plate (Text-fig. 54). Aedeagus with shaft short and slightly sinuate (Text-fig. 53). 

$. Length 5-5 mm. Colour and general structure similar to male except for the median 
stripe on the face which is usually paler. Eighth sternite (Text-fig. 73) with a fairly deep 
broad cleft apically, and with a black oblique bar on each side almost at the apical margin. 

Holotype < (pinned, abdomen in glycerine). SOUTH AFRICA: Pondoland, Port 
St. John, i-n.vi.ig23 (R. E. Turner). In B.M.(N.H.). 

Allotype ?. SOUTH AFRICA: Zululand, Gingindhlovu, 15^.1926 (R. E, Turner). 
In B.M.(N.H.). 



i8 



H. H. ROSS 




54 



FIGS. 52-54. Male genitalia of Exitianus turneri. 52, pygofer; 53, aedeagus, lateral aspect; 
54, genital assemblage, ventral aspect (one style omitted), a, aedeagus; c, connective; 
P> pygofer; pl> plate; s, style. 



Paratypes. Same data as allotype, 3 $ (one without abdomen), 4 $, with inclu- 
sive dates 9.vi-5.vii.i926; Natal, Kloof, 1500 ft., i $, viii.ig26 (R. E. Turner). 
In B.M.(N.H.), one pair in Illinois Natural History Survey collection. 

This species differs from the preceding in the shorter plates and other details of 
the genitalia, and in the contrasting pale costal margin of the tegmina. In this 
latter character it resembles the following species, from which it differs in details 
of the male genitalia and the female eighth sternite. 



Exitianus zuluensis sp. n. 

(Text-figs. 61, 62, 63, 74) 

cJ. Length 5 mm. Colour tawny, the head with an arcuate black band on vertex, and 
almost equally dark median stripe on the face. Lateral areas of face and legs with dark mark- 
ings. Tegmina with all veins brown except the costal margin which is cream colour. General 



THE OLD WORLD EXITIANUS 19 

structure typical for genus. Pygofer (Text-fig. 63) with apical half strongly narrowed and 
upturned, ending in a single, dorsally-directed heavy black spine (presumably number 2). 
Plates short, broadly rounded at apex, and without large lateral setae (Text-fig. 61). Style 
with apex obliquely truncate, only moderately wide, extending almost to the apex of the 
plate (Text-fig. 61). Aedeagus with shaft relatively small, slightly angulate near middle, 
and narrowed at extreme tip (Text-fig. 62). 

$. Length 5-5 mm. Colour and general structure same as male. Eighth segment with 
transverse posterior margin and with a pair of dark areas, one on each side, just in front of 
posterior margin (Text-fig. 74). 




FIGS. 55-63. Male genital parts of Exitianus spp. 55, 56, 57, E. okahandia (holotype); 
58, 59, 60, E. natalensis (holotype); 61, 62, 63, E. zuluensis (holotype). 55, 58, 61, 
style, ventral aspect, with outline of plate shown beneath; 56, 60, 63, pygofer; 57, 59, 
62, aedeagus, lateral aspect. 



20 H. H. ROSS 

Holotype $ (pinned, abdomen in glycerine). SOUTH AFRICA: Zululand, Eshowe, 
vi.i926 (R. E. Turner}. In B.M.(N.H.). 

Allotype <j>. Same data as holotype. In B.M.(N.H.). 

Paratypes. Same data as holotype, 3 $, 3 without abdomens, with inclusive 
dates, vi-vii. In B.M.(N.H-), i 9 in Illinois State Natural History Survey collection. 

This species is readily distinguished from the preceding by the peculiar upturned 
pygofer, and from the species that follows by its pale colour pattern. 




FIGS. 64-67. Male genital parts of Exitianus spp. 64, E. attenuatus (holotype), apex 
of right side of pygofer; 65, same (paratype), apex of right and left sides of pygofer, 
respectively; 66, same (paratype), quartering posterior view of aedeagal shaft; 67, 
E. pondus (holotype), apex of pygofer. 



Exitianus natalensis sp. n. 

(Text-figs. 6, 58, 59, 60, 75) 

cj. Length 4-5 mm. Base colour of dorsum chiefly tawny, vertex of head (Text-fig. 6) 
with a very wide black arcuate band, pronotum with lateral dark marks, mesonotum with 
lateral black triangles, face mostly black with tawny enclosed areas, the facial marks running 
up onto vertex; venter and legs mostly black; tegmina with veins brown and costa contrast- 
ingly cream coloured. General structure typical for genus. Pygofer (Text-fig. 60) with apical 
half strongly narrowed and upturned, the tip bearing spines i and 2, their bases contiguous, 
spine i very slender, shorter than 2, spine 2 long and massive. Plates fairly short, and apex 
broadly rounded, the lateral margin without large setae (Text-fig. 58). Style with relatively 
long apex, which is truncate and which almost reaches the margin of the plates (Text-fig. 58). 
Aedeagus with shaft relatively small, curved near middle, and narrowed at extreme tip (Text- 
fig- 59). 

Holotype <$ (pinned, abdomen in glycerine). SOUTH AFRICA: Natal, Van Reenen, 
Drakensberg, 1-22.1.1927 (R. E. Turner). In B.M.(N.H.). 

Three females from Pondoland, Port St. John (B.M.(N.H.)), may belong to this 
species. In general colour pattern they resemble the male closely, but the prono- 
tum and mesonotum are both much darker and therefore there is a possibility that 
they may represent another undescribed species. The eighth sternite has the apical 
margin transverse and uniformly dark coloured (Text-fig. 75). 



THE OLD WORLD EXIT I AN US 



f 



s, 




r ^v \ \ 

1 Q \ ' T* \ 





69 * 73 

\ 



74 l 77 



~\ r TOT \ 

71 I ' T 75 



FIGS. 68-77. Apex of female eighth sternite. 68, E. plebeius (holotype) ; 69, E. indicus 
(holotype); 70, . capicola (holotype); 71, E. taeniaticeps (Libya); 72, . okahandia 
(allotype) ; 73, . turneri (allotype) ; 74, . zuluensis (allotype) ; 75, . natalensis 
(Pondoland) ; 76, . nanus (holotype); 77, E. africanus (holotype). 

UNPLACED SPECIES 

Bythoscopus africanus Walker, 1851 : 873. Type 9 seen m B.M.(N.H.), from 
Sierra Leone (Morgan), 42, 31. The banded head (much as in Text-fig. 10) and the 
elongate ovipositor indicate that this species is a member of Exitianus, but the 
eighth sternite has a straight posterior margin and a pair of large posterior dark 
spots (Text-fig. 77). No males have been associated with this distinctive specimen, 
apparently representing a valid species. 

Athysanus capicola Stal, 1855 : 99. Type $ seen in the Stockholm Museum, from 
Meru, Neider. This is a typical member of the taeniaticeps group, with head banded 
much as in Text-figure 9 and the eighth sternite with a median notch (Text-fig. 70). 
At present it is impossible to associate this specimen with a recognized species. 

Eutettix pallida Evans and selbyi Evans, 1938 : 14. Both of these species were 
described from 9 types collected in Queensland, and might prove to be additional 
junior synonyms of plebeius. The leafhopper fauna of Australia is as yet not well 
enough collected to make undue assumptions concerning the identity of females in 
this group of species. 

SPECIES TRANSFERRED TO OTHER GENERA 

Two species placed in Exitianus by Evans (1966) do not belong in this genus 
Nephotettix contemptus Kirkaldy (1906) and Nephotettix eurytus Kirkaldy (1907), 
both described from Sydney, N.S.W., Australia. I have examined the $ holotypes 
and an associated $ of contemptus; they appear to belong to Hybrasil Kirkaldy 
(1907 : 41) with brani Kirkaldy as type-species. I have not examined the type of 
the latter, but the Australian specimens mentioned agree in all but details of indi- 
vidual parts with Linnavuori's (1960) illustrations and description of brani. 

The types of Kirkaldy's two Australian species appear identical. The description 



22 H. H. ROSS 

of Carvaka fulvida Evans (1966), described from the Bunya Mountains, Queensland, 
Australia, is highly suggestive of the same species. The following synonymy is 
therefore offered: 

Hybrasil contemptus (Kirkaldy) comb. n. 

Nephotettix contemptus Kirkaldy, 1906 : 332. 
Nephotettix eurytus Kirkaldy, 1907 : 54. syn. n. 
PCarvaka fulvida Evans, 1966 : 244. comb. n. 

The taxonomic position of Hybrasil is uncertain. Linnavuori (1960) placed it in 
the Drabescinae and Evans (1966) his C. fulvida in the Selenocephalini (Cicadellinae) . 
Salient features of the genus include the following: Clypellus constricted in middle 
and expanded at apex; front margin of head with a yellow band defined above and 
below by a low but sharp carina, and extending from eye to eye; venation typical 
but with apical two costal crossveins edged with black (as in Fieberiella); plate 
elongate, the apex narrow and partly membranous, and without large lateral setae ; 
aedeagus with shaft articulated with socket; and ovipositor extending well beyond 
pygofer (as in Exitianus). Characters of the aedeagal shaft and ovipositor suggest 
an affinity with the Doraturini. 





78 



FIGS. 78-81. Male genital parts of leafhoppers. 78, Exitianus greensladei (holotype) ; 
79, 80, 81, Nephotettix apicalis (India). 78, 79, plate, ventral aspect; 80, pygofer; 81, 
aedeagus, lateral aspect. 



PROBLEMS OF SPECIES RECOGNITION 

On the basis of material that has been examined in this study, there is little 
problem in establishing satisfactory species-limits in the nanus-gioup (only one 
species), the distanti-group (only one species), and the okahandia-group (four species). 



THE OLD WORLD EXITIANUS 23 

In the obscurinervis and taeniaticeps-groups, however, the situation is far from clear. 

OBSC URINER F/5-GROUP 

DeLong & Hershberger (1947) recognized eleven American species, using the 
number and juxtaposition of the pygofer spines as their primary bases of species 
separation, supplemented with colour differences. Linnavuori (1959) uses the same 
sequence of characters in keying the seven species he recognized from the neotropical 
region. Oman (1949) questions the value of the number of the pygofer spines as 
diagnostic characters. He points out that it is not uncommon to find a specimen 
having unequal numbers of these spines on the left and right sides of the pygofer; 
I have observed this same phenomenon. It is possible, however, that in some 
species the relative position of the spines (if present) may represent diagnostic 
characters. 

The real crux of whether or not different entities should be considered as distinct 
species is whether or not each unit forms a continuing series of populations progress- 
ing through time and evolving independently of other genetic lineages (Simpson, 
1951; Ross, 1962). This is a theoretical concept that can be tested experimentally 
by crossbreeding if adequate material is available and techniques are known. With 
Exitianus this experimental avenue is not yet available. The only other highly 
reliable criterion of the evolutionary distinctness of bisexual phylogenetic lines is 
whether or not the species segregates occur in the same area, yet remain distinct, 
with little or no indication of hybridization. In a number of collections of American 
Exitianus, I have found excellent evidence of different pairs of species occurring 
together with no sign of intergradation, and I infer from this that most of the des- 
cribed American species are indeed independent phylogenetic lines. 

In sorting through these collections, however, another problem has arisen that 
affects the use of existing keys and descriptions. In almost every species the males 
are darker than the females and frequently have colour patterns that are more 
distinctive and more sharply delineated than those of the females. In several 
instances the differences between the females are negligible whereas the colour 
patterns of the males are readily differentiated. This is suggestive that the distinc- 
tive patterns of the males have evolved as differential display stimuli to which only 
the correct female normally responds. 

This sexual dichromatism complicates the matter of identification, because in 
existing keys females of several species would key out to the same place, and would 
run to a different couplet than would the male of the same species. To compound 
this situation, the shape of the female seventh sternite gives a different visual image 
depending on the position in which the female died, and use of this character for 
diagnosis must be attended by extreme caution. Until much more information is 
available concerning these problems, it seems best to treat all the currently des- 
cribed American species as distinct entities. In Text-fig. 82 the species of the 
obscurinervis-group have been arranged phylogenetically on a tentative basis using 
the characters illustrated by DeLong & Hershberger and by Linnavuori. 



24 H. H. ROSS 

TAENIATICEPS-GROUP 

Of the described species, ootacamundus , kilimanus, coronatus, and greensladei are 
the only ones having distinctive colour patterns. The remainder have simply a 
more or less distinct black band across the top of the head. When abdomens of 
this unicolorous group were cleared and examined, the first thing evident was the 
striking similarity in all of them of the two pygof er spines : spine i long and slender, 
somewhat variable in length, and spine 2 short, black, and twisted at the apex. 
Closer inspection revealed small differences in the shape of the aedeagus, especially 
its profile. In large series of indicus, this structure has a distinctive broad shape 
compared with material from islands of the Indian Ocean and from Australia. 
Material from Asia Minor and North Africa has a distinctive clavate profile with a 
pronounced step where the two short processes arise ; in some specimens from South 
Africa the aedeagus has essentially the same shape but is more massive; and in 
collections from the Sudan, unimodal intergrading populations were found that 
bridged the gap between the Asia Minor and South African types. A long series 
of material from Madagascar added two other types, one of them similar to the 
island type and the other differing only slightly from the Asia Minor type. In the 
first sorting it appeared as if the aedeagus varied randomly and locally in a fashion 
that almost defied segregation. 

On further analysis two points of considerable interest emerged. First, in the 
three Indian species, indicus, ootacamundus, and coronatus, the aedeagi were of the 
same type (Text-figs. 25, 28, 30, 31). Furthermore, in a series of several thousand 
specimens from Kharagpur there was no indication of intergradation between the 
plain banding on the head of indicus and the colour patterns distinctive for ootaca- 
mundus and coronatus. These circumstances suggest strongly that a line of the 
taeniaticeps group dispersed into India, evolved the indicus type of aedeagus, and 
subsequently evolved into three species now differentiated primarily on the basis 
of head patterns. Second, in Madagascar two types of aedeagi occurred in some 
of the same localities yet no intergrades were found between them. This suggests 
that these two morphological types represent true genetic isolation and that two 
phylogenetic lines are involved. 

On the basis of these two different cases, there seems to be a good possibility 
that the different morphological types of the taeniaticeps group, though sometimes 
remarkably close, represent distinctive species, and they have been so treated in 
this paper. Unquestionably a tremendous amount of collecting will be necessary 
to establish the true nature of these species but until this additional information 
is forthcoming, the only clear presentation seems to call for a course that may result 
in some splitting rather than taking the chance of obscuring future leads by lumping. 

PHYLOGENY 

After the various species of Exitianus had been segregated, the next problem was 
to decipher the evolutionary sequence and geographic dispersal of the various 
lineages. As a first step in this evolutionary reconstruction, the species most 
similar to each other (essentially the groups expressed in the taxonomic analysis) 



THE OLD WORLD EXITIANUS 25 

were examined for inter-group differences. The various conditions found for any 
one character were then compared with conditions found in related genera in an 
effort to determine which were the ancestral and which the derived conditions. 

As is true in most inducto-deductive investigations, certain theoretical possibilities 
are realized when the study commences, and character differences are viewed in 
their possible evolutionary light from the earliest observations of character differences. 
The present study was no exception. It was first noticed that in a number of Old 
World species, pygofer spine 2 was twisted at the apex (Text-fig. 24) and that the 
profile of the aedeagus was incised at the tip to form a fine curved point (Text- 
fig. 25), whereas in the New World species spine 2 was symmetrical (Text-fig. 19) 
and the profile of the aedeagus was blunt and little incised (Text-fig. 20). Next it 
was seen that (i) in one Old World species the pygofer had four, five, or six spines 
forming an even row (Text-fig. 15), whereas in most New World species there were 
only three spines with a large space between the two more ventral spines (spine 3 
was missing, Text-fig. 19), and (2) in those Old World species with the contorted 
spine 2, there were only two spines. Finally, another group of African species came 
to light in which the plates were short and had only very small lateral setae (Text- 
fig. 54), whereas in all other species the plates were long and each had a lateral 
row of long spines (Text-fig. 78). 

A search for related genera with which to compare conditions found in Exitianus 
soon divulged the information that (i) the genus Nephotettix was the only demon- 
strable close relative of Exitianus, (2) the genera Gillettiella Osborn, Stirellus Osborn 
& Ball, Doratura Sahlberg, and other genera possessing a long ovipositor were 
components of the same tribal unit (Doraturini) as Exitianus, and (3) many genera 
related to Athysanus Burmeister and Recilia Edwards belonged to the same large 
branch of the leaf hoppers. 

In Nephotettix the pygofer has a set of apical dark spines much like those in 
Exitianus. The dorsal spine (Text-fig. 80) seems to correspond with spine i of 
Exitianus ; the lower four spines seem to correspond with spines 3 and 4 of Exitianus 
but are each represented by a pair of spines; spine 2 seems to be missing. The 
plate is long and triangular, bearing lateral long setae (Text-fig. 79), exactly like 
those of Exitianus. The aedeagus has the same parts as in Exitianus, but the shaft 
is highly ornamented (Text-fig. 81). Nephotettix, however, has an unusual opaque 
green ground colour and a sharply carinate forehead, and thus would appear to be 
a separate branch arising before the evolution of Exitianus as it is now represented. 
It seems reasonable to suppose that Nephotettix and Exitianus arose from a common 
ancestor that was remarkably like the nanus group of Exitianus, but probably having 
a more cryptic colour pattern similar to that found in many other grass-inhabiting 
leaf hoppers. 

Comparing the conditions exhibited by Exitianus and Nephotettix with those found 
in other leafhopper genera, it was evident that (i) the long plates with lateral spines 
represented the ancestral condition of this structure within Exitianus, and the 
short plates represented a derived condition ; (2) the larger number of pygofer spines 
was the ancestral condition for the structures, the reduced number the derived 
condition ; (3) the symmetrical pygofer spine 2 was the ancestral condition, the twisted 



26 



H. H. ROSS 



spine 2 the derived condition; and (4) the blunt aedeagus was the ancestral con- 
dition for this structure, the incised and pointed type the derived condition. 

From these considerations of ancestral and derived conditions, it is most likely 
that the ancestor of Exitianus (Ancestor i in Text-fig. 82) possessed triangular setose 
plates, at least four or five pygofer spines, all symmetrical, and a blunt aedeagus. 
These conditions are exemplified by nanus, which would therefore seem to be the 
living representative still bearing this archaic combination of ancestral characters. 
The remainder of the species have lost either spine 3 or spines 3, 4, and 5. It is 
therefore reasonable to suppose that Ancestor i gave rise to a lineage losing spine 
3 and represented by Ancestor 2. Most of the native American species comprising 
the obscurinervis group represent this condition (Text-figs. 19, 20). 




II 



II 

c/> ^ 



3 

^ <fc 

li 



^ i 
s ^ 

<o $ 



< 




CO 

W m 
ui -5 in i" 


CO 

-j - 




Style 



FIG. 82. Proposed family tree of Exitianus. Arabic numerals refer to putative ancestral 
forms explained in text. Roman numerals indicate postulated past dispersals as 
follows: I, Africa to New World; II, Africa to South Pacific; III, Africa to India. 



THE OLD WORLD EXITIANUS 27 

If one adds together the primitive conditions found in the remainder of the species 
(all native to the Old World), the result is a form having triangular setose plates, 
blunt aedeagus, and only two aedeagal spines. This ancestral form (Ancestor 3 
of Text-fig. 82) differs from Ancestor 2 only in lacking pygofer spine 4. Ancestor 3 
apparently gave rise to two lineages. In one of these, represented by Ancestors 
4 and 5, the plates did not change but the aedeagus became incised and pointed at 
the tip ; in the other lineage, represented by Ancestor 6, the aedeagus did not change 
but the plates became short and their setae greatly reduced. In Ancestor 4, pygofer 
spine 2 was probably still symmetrical. Such a stage may be represented by distanti, 
although in this species spine 2 is more elongate and in this respect is specialized 
over the theoretical structure of Ancestor 4. In addition to distanti, Ancestor 4 
gave rise to Ancestor 5 in which pygofer spine 2 became twisted at the apex. 

From Ancestor 5 evolved the species of the taeniaticeps group. The most primitive 
species of the group is attenuatus, in which pygofer spine 2 is only slightly contorted 
(Text-figs. 64, 65) and appears to represent a transitional condition leading to the 
more extreme asymmetry of this spine found in all other members of the taeniaticeps 
group. Nine of the remaining ten species of the group are readily grouped into 
three complexes. In the three species indicus, ootacamundus , and coronatus the 
aedeagal shaft is unusually short and broad (Text-figs. 25, 28, 30, 31). In pondus, 
taeniaticeps, and frontalis the dorsal points of the shaft are situated on a steplike 
sinuation of the profile (Text-figs. 34-36, 39-42, 47). In mucronatus, plebeius, and 
greensladei the basal profile of the shaft has no such basal sinuation (Text-figs. 
43-46, 48-51). At present there seems to be no logical basis for knowing the exact 
affinities of these three complexes, hence in Text-fig. 82 their origin is depicted as 
a trichotomy. The tenth species, kilimanus, can at present be placed in the family 
tree only tentatively because of a lack of knowledge of the internal structures. 

Ancestor 6, the 'short plate' offshoot of Ancestor 3, is represented by the okahandia 
group. In the species okahandia, the wing colour and head proportions are similar 
to those of paler species in the other groups; in turneri, natalensis, and zuluensis 
the tegmen or front wing has a fairly broad whitish costal stripe running the full 
length of the wing. These three apparently represent an offshoot of Ancestor 6 
in which this character became established. In both okahandia and turneri the 
shape of the pygofer and the position of its apical spines is typical of other members 
of the genus, hence this condition must have been present in Ancestor 6 also. From 
this we can infer that the broad, flared styles found in these two species were also 
present in Ancestor 6. From the lineage leading to turneri there arose a branch 
in which the apex of the styles became elongate (Text-figs. 58, 61) and the apex of 
the pygofer became narrowed and upturned, pygofer spine 2 became greatly en- 
larged and spine I reduced (Text-fig. 60), ultimately lost in zuluensis (Text-fig. 63). 
Judging by the character by character changes that appear to characterize much 
of insect evolution, it is a reasonable prediction that additional collecting will bring 
to light an ancestral form having either the elongate style or the highly modified 
pygofer, but not both, now found in natalensis and zuluensis. Such a form would 
give us information as to which of these character changes evolved first. 



28 PALAEODISPERSAL 

When geographic distribution is compared with the postulated family tree of 
Nephotettix and Exitianus (Text-fig. 82), it is apparent that the genus is tropical or 
subtropical (except for the American exitiosus), and therefore has in all likelihood 
been restricted to the tropics and subtropics during almost all of its evolution. 
It is further apparent that the evolution of the entire group occurred in the Old 
World tropics with the exception of the obscurinervis group, which evolved in 
North and South America. Linnavuori recorded several South American specimens 
as taeniaticeps , which I am presuming represent a transport by man. If this is 
indeed the case, then the only natural intercontinental dispersal of which we have 
record was an Old World-to-New World dispersal of Ancestor 2. 

Considering only the Old World records, the simplest explanation is reached by 
assuming that the genus evolved primarily in Africa. The indicus branch of Ancestor 
5 apparently is a moderately old dispersal from Africa to southeastern Asia. The 
plebeius-greensladei branch of Ancestor 5 represents another dispersal eastward to 
Australia and the South Pacific. The present day widespread distribution of 
nanus and frontalis could be the result of natural dispersal. This seems especially 
likely with regard to nanus. 

This probable African evolution of the genus poses a problem with regard to 
probably the oldest intercontinental dispersal of the genus, that of Ancestor 2 to 
the Americas, represented by Ancestor 2A in Text-fig. 82. If this occurred 
sufficiently long ago, possibly in the Eocene or early Oligocene, it would be simple 
to postulate an overland dispersal through subtropical climates across the Bering 
bridge. If this did happen it is puzzling that a branch of the genus comparable to 
the obscurinervis group did not evolve and persist in southeastern Asia. 

A second possible solution is that Ancestor 2 was carried from Africa to South 
America by wind dispersal. This explanation seems more probable because of the 
discovery of long-distance migrations of much smaller leafhoppers in moving masses 
of warm, moist air. If this were the case, it is remarkable that apparently only 
one such dispersal occurred and became established during the entire evolution of 
the genus. Dispersal patterns of grass subgenera would indicate that satisfactory 
grass hosts for Exitianus have been present in the Americas for at least the probable 
period of Exitianus evolution. 

A third explanation is that Ancestor 2 was a species with an unusually wide 
ecological tolerance, much like that of present day Nearctic exitiosus, that the species 
was able to cross into North America through warm temperate rather than tropical 
climates, and that subsequently, for one reason or another, the range of the wide- 
spread species became fragmented into Old and New World segments that eventually 
lost their ability to live in temperate climates. 

A better insight into the circumstances surrounding the American dispersal of 
Exitianus may come when we understand more about the dispersal of grassland 
communities as a whole. 

ACKNOWLEDGMENTS 

I have received much technical and advisory help in these studies from Dr. W. J. 
Knight and other officers of the British Museum, Dr. Alfred Orian, Mauritius Depart- 



THE OLD WORLD EX1TIANUS 29 

ment of Agriculture, and Dr. J. P. Kramer, U.S. Department of Agriculture. To 
Mrs. Nell Miles, Urbana, 111., Dr. F. R. Steggerda, University of Illinois, Dr. H. G. 
Walker, Los Angeles County Museum, Dr. L. W. Quate, University of Hawaii, 
Dr. D. H. Murphy, University of Singapore, Dr. J. W. Evans, Australian Museum, 
Dr. G. C. Sanderson, Illinois Natural History Survey, and Mrs. N. J. Kemp, Australia, 
I am indebted for Old World material collected or loaned, and especially to Dr. 
L. T. Gressitt and his associates, B.P. Bishop Museum, for the loan of type material. 
To the officers of the Stockholm Museum I am much indebted for courtesies accorded 
on a visit to their Museum. 

This study has also been supported by a research grant from the U.S. National 
Science Foundation and in many ways by the Illinois Natural History Survey. 



REFERENCES 

BALL, E. D. 1929. A supplemental revision of the genus Athysanus in North America 

(Homoptera : Cicadellidae) . Trans. Am. ent. Soc. 55 : 1-81. 
BERGEVIN, E. DE. 1925. Description d'une nouvelle espece d'Athysanus suceur de sang 

humain de 1'extreme Sud Algerien (Hemiptere-Homoptere, Jassidae). Archs Inst. Pasteur 

Alger. 3 : 42-44, 5 figs. 
BELONG, D. M. & HERSHBERGER, R. V. 1947. The genus Exitianus in North America 

including Mexico. Ohio J. Sci. 47 : 107-116, 2 pis. 
DISTANT, W. L. 1908. The Fauna of British India including Ceylon and Burma. Rhynchota- 

Homoptera. 4, 501 pp., 282 figs. London. 

1909. 'Sealark' Rhynchota. Trans. Linn. Soc. Lond. Zool. 13 (2) : 29-47, 4 pis. 

1917. Rhynchota. Part II: Suborder Homoptera. The Percy Sladen Trust Expedi- 
tion to the Indian Ocean in 1905, under the leadership of Mr. J. Stanley Gardiner, M.A. 
Trans. Linn. Soc. Lond. Zool. 17 : 273-322, pis. 49-51; text-figs. 1-17. 

1918. The Fauna of British India, including Ceylon and Burma. Rhynchota. Homoptera: 
Appendix. Heteroptera: Addenda. 7 : i vii, 1-210, 90 pis. London. 

EVANS, J. W. 1938. Australian leafhoppers (Homoptera, Jassoidea). Part VIII. Pap. 
Proc. R. Soc. Tasm. 1938 : 1-18, 3 pis. 

1966. The leafhoppers and froghoppers of Australia and New Zealand (Homoptera : 
Cicadelloidea and Cercopoidea) . Mem. Aust. Mus. 12 : 347, 48 figs. 

ISHIHARA, T. 1954. Homopterous notes. Scient. Rep. Matsuyama agric. Coll. 14 : 1-28, 

17 figs. 
JACOBI, A. 1910. Wissenschaftliche Ergebnisse der Schwedischen Zoologischen Expedition nach 

dem Kilimandjaro, dem Meru und den Umgebenden Massaisteppen Deutsch-Ostafrikas 

1905-1906. Unter Leitung von Prof. Dr. Yngve Sjostedt. Herausgegeben mit Unter- 

stiitzung von der konigl. Schwedischen Akad. der Wiss. 12. Hemiptera. 7 Homoptera. 

2: 97-136, pis. 1-2, figs. A-N. Stockholm. 
KIRSCHBAUM, C. L. 1868. Die Cicadinen der Gegend von Wiesbaden und Frankfurt a. M. 

nebst einer Anzahl neuer oder schwer zu unterscheidender Arten aus anderen Gegenden 

Europa's. Jb. nassau ver. Naturk. 21-22 : 1-202. 
KIRKALDY, G. W. 1906. Leafhoppers and their natural enemies. Pt. IX, Leafhoppers. 

Hemiptera. Bull. Div. Ent. Hawaiian Sug. Plrs' Ass. Exp. Stn. I (9) : 271-479, pis. 21-32. 

1907. Leaf hoppers -supplement. (Hemiptera.) Bull. Div. Ent. Hawaiian Sug. Plrs' Ass. 
Exp. Stn. 3 : 1-186, pis. 1-20. 

LINNAVUORI, R. 1959. Revision of the neotropical Deltocephalinae and some related sub- 
families (Homoptera). Suomal. eldin-ja kasvit. Seur. van. eldin. Julk: 1-370, 144 figs. 

1960. Cicadellidae (Homoptera, Auchenorrhyncha) of Fiji. A eta ent. fenn. 15 : 1-71 
22 figs. 



30 H. H. ROSS 

MATSUMURA, S. 1914. Die Jassinen und einige neue Acocephalinen Japans. /. Sapporo 

agric. Coll. 5 : 165-240, figs. 1-12. 
METCALF, Z. P. 1946. Homoptera. Fulgoroidea and Jassoidea of Guam. Bull. Bernice 

P. Bishop Mus. 189 : 105-148, figs. 1-30. 
MOTSCHULSKY, V. I. DE. 1863. Essai d'un catalogue des insectes de Tile Ceylan. Byull. 

mosk. Obshch. Ispyt. Prir. 36 : 1-153, illus. (94-113). 

OMAN, P. W. 1949. The Nearctic leafhoppers (Homoptera : Cicadellidae) . A generic classi- 
fication and check list. Mem. ent. Soc. Wash. 3 : 1-253, pis- I- 44- 
RIBAUT, H. 1952. Faune Fr. Homopteres Auchenorhynques. II (Jassidae). 57 : 1-474, 

figs. I-I2I2. Paris. 
Ross, H. H. 1962. A Synthesis of Evolutionary Theory. 387 pp., 146 figs. Englewood 

Cliffs, New Jersey. 

SIMPSON, G. G. 1951. The Species concept. Evolution 5 : 285-298, 4 figs. 
STAL, C. 1855. Hemiptera fran Kafferlandet. Ofvers. K. VetenskAkad. Forh. 12 : 89-100. 
WALKER, F. 1851. List of the specimens of Homopterous insects in the collection of the British 

Museum 3 : 637-907. London. 
ZACHVATKIN, A. A. 1935. Notes on the Homoptera-Cicadina of Jemen. Uchen. Zap. mosk. 

gos. Univ. 4 : 106-115. 




A LIST OF SUPPLEMENTS 
TO THE ENTOMOLOGICAL SERIES 

OF THE BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 



1. MASNER, L. The types of Proctotrupoidea (Hymenoptera) in the British 
Museum (Natural History) and in the Hope Department of Entomology, Oxford. 
Pp. 143. February, 1965. 5. 

2. NIXON, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera : 
Braconidae). Pp.284; 348 Text-figures. August, 1965. 6. 

3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177 ; 
18 plates, 270 Text-figures. August, 1965. 4 45. 

4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172 ; 500 Text-figures. October, 

1965- 35*. 

5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156 ; 
475 Text-figures. November, 1965. 2 155. 

6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae & Drosophilidae. 
Pp. 129 ; 328 Text-figures. 3. 

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

3&3S- 

8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the 
world species of Cleora (Lepidoptera : Geometridae). Pp. 119; 14 plates, 146 
Text-figures, 9 maps. February, 1967. 3 los. 

9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera : Rhopalocera). Pp. 509. August, 1967. 8 IDS. 

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rhopa- 
locera). Pp.322; Coloured frontispiece, 348 text-figures. August, 1967. 8. 



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



(HYMENOPTERA : BRACONIDAE) 



G. E. J. NIXON 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 22 No. 2 

LONDON: 1968 



A REVISION OF THE GENUS 

MICROGASTER LATREILLE 
(HYMENOPTERA : BRACONIDAE) 



BY 




G. E. J. NIXON 

jr r f*^A 

Commonwealth Institute of EntomologjJ 



Pp. 31-72 ; 33 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 22 No. 2 

LONDON: 1968 



THE BULLETIN OF THE BRITISH MUSEUM 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

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

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

This paper is Vol. 22, No. 2 of the Entomological 
series. The abbreviated titles of the periodicals cited 
follow those of the World List of Scientific Periodicals. 



World List abbreviation : 
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Trustees of the British Museum (Natural History) 1968 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 28 May, 1968 Price Eighteen Shillings 



A REVISION OF THE GENUS MICROGASTER 
LATREILLE (HYMENOPTERA : BRACONIDAE) 

By G. E. J. NIXON 

CONTENTS 

Page 

SYNOPSIS ........... 33 

ACKNOWLEDGMENTS ......... 33 

THE GENUS Microgaster LATREILLE . . . . . . . 33 

Key to species (Females) and description of species . . . . 34 

SPECIES INQUIRENDAE ......... 69 

REFERENCES ........... 70 

INDEX ............ 71 

SYNOPSIS 

The genus Microgaster, as denned by Nixon in 1965, is revised and all species falling within 
the new definition, as far as they have been recognized, are taken into account. Forty-six 
species are keyed and discussed and fourteen of these are brought forward as new. Five 
species are put in synonymy. 

ACKNOWLEDGMENTS 

IN preparing this revision I have been much helped by a very fine collection of 
Microgaster from Ireland, received on loan from Mr. A. W. Stelfox of Newcastle, 
Co. Down, N. Ireland and now the property of the U.S. National Museum. I am 
much indebted to Mr. Stelfox. I am also grateful to the following gentlemen for the 
loan of useful material: Dr. Max Fischer of the Naturhistorisches Museum, Vienna, 
Dr. Wolter Hellen of the Helsinki Museum, Helsinki, Dr. C. F. W. Muesebeck of 
the U.S. National Museum, Dr. Jeno Papp of the Hungarian Natural History 
Museum, Budapest,and Dr. V. I. Tobias of the Zoological Institute, Leningrad. 



IN my revision of the Microgasterini I redefined this genus (1965 : 267 and in key, 
: 13) restricting the use of the name to the group of species showing an obvious and 
natural relationship with the type-species, deprimator Fab. (Muesebeck & Walkley, 
1951 : 135). Many of the better known of the remaining European species were 
transferred by me to the genus Protomicroplitis. 

Microgaster of authors now consists virtually of two genera Microgaster, as 
used here, essentially N. Temperate, and Protomicroplitis Ashmead with a world- 
wide distribution and best represented in tropical regions. 

In 1965 I divided Protomicroplitis into many species-groups, some of which were 
established for species that were traditionally included in European Microgaster, 
such as marginatus Nees, abdominalis Nees and scotica Marshall. 

The species of Microgaster s. str. are remarkably alike in general facies and quite 

ENTOM. 22, 2. 2 



34 G. E. J. NIXON 

impossible to recognize on the characters that have been used in the past. Species 
with red hind femora have usually been called globata L. Indeed, this is the name 
that appears most frequently in the literature on European Microgaster. I do not 
know to what species it should be applied. The Linnean type no longer exists and 
the little that Linne said about it indicates that it is a gregarious parasite living in 
stalks (? reeds) " in Culmis, intra folliculum communem, bombycinum, subrotundum, 
album ". Microgaster globata auctt. is a solitary parasite. The name "globata " 
is not used in this paper. 

Microgaster s. str. was recognized as a distinct segregate within the older concept 
of the genus by earlier workers. Its species have always presented a tough taxo- 
nomic problem, baffling taxonomists by what was thought to be their extreme 
variability. Only Thomson, always a pioneer in the taxonomy of difficult groups 
of insects, succeeded in defining the limits of some of the species. Marshall, who 
lacked neither carefulness nor good judgement, was handicapped by the practice, 
fashionable in his time, of gumming insects on pieces of card, thus obscuring import- 
ant areas of their anatomy. Much more recently Fahringer (1936), Telenga (1955) 
and Papp (1959, 1960) have tried to define what they consider to be species of 
Microgaster but none, unhappily, has discovered any new characters that permit 
even a partial break-down of the genus. 

Having studied Microgaster intermittently for several years, I have been able to 
find a few structural characters that narrow considerably the margin of error in 
naming species. These characters are : the presence of teeth, or even a lobe (deductor) , 
on the claws, the relative proportions of the abscissae of the basal vein of the fore 
wing and the degree of sclerotization of the hypopygium. But even with the help 
of these characters, I cannot claim to have cleared away all the difficulties of species- 
differentiation in Microgaster. The correlation of the North American and Euro- 
pean species is specially in need of investigation. 

When I defined the limits of Microgaster in 1965, I stated that the first abscissa 
of the discoideus is fully equal to the second. This is true of all the species included 
below with the exception of consors sp. n. from England. In all other respects, this 
species is typical of the genus. 

As stated earlier, Microgaster is mainly confined to north temperate regions. 
It does, nevertheless, extend as far as the Oriental region, tjibodas from Java, 
kuchingensis from Borneo and magnifica from Queensland. I include also one 
species from Mexico, nerione. 

In conclusion, it is important to state that this revision is based only on material 
that I have personally examined. For this reason, published host records are 
excluded, since no reliance can be placed on the identification of the parasites. 



KEY TO SPECIES 

FEMALES 

Notaulic courses widened behind and here tending to coalesce (at most separated 
by a weak keel) so that the posterior half of the mesoscutum shows a large, more 
or less sunken area of coarse rugose-punctation or rugose-reticulation. 



REVISION OF MICROGASTER 35 

Gaster always in part red ; tergite 2 extremely coarsely rugose and considerably 
longer than 3; apical tarsal segment of all legs enlarged; claws very large 
hypopygium heavily sclerotized all over and hence without trace of 
lateral creasing; r us satus -group ......... 2 

Notaulic courses, if indicated at all, never reaching beyond the middle of the disc; 
posterior part of mesoscutum thus never with an area of sunken rugosity; 
deprimator-group ............ 4 

Hind coxa red; tergite 2 hardly more than twice as wide as long; hind tarsal segment 
3 fully one and a half times longer than wide; ovipositor sheath distinctly longer 
than the hind basitarsus .......... 3 

Hind coxa blackish ; tergite 2 fully two and a half times wider than long ; hind tarsal 
segment 3 hardly longer than wide; ovipositor sheath distinctly shorter than the 
hind basitarsus 

Gaster short, broad (Text-fig. 31); tergites 1-3 black; rest of gaster red; inner 
spur of the hind tibia not reaching middle of hind basitarsus. 

Europe rugulosus Nees (p. 68) 

Tergites i and 2 and most of 3 red, the rest black ; head behind the ocelli shiny and 
with only weak traces of punctation; preapical segment of the antenna not more 
than twice as long as wide; flagellum yellow. Europe. Japan 

russatus Haliday (p. 67) 

Tergites i and 2 black, the rest broadly banded with black or more or less entirely 
black; head behind the ocelli dull, densely rugose-punctate; preapical segment of 
the antenna fully two and a half times longer than wide; flagellum brownish but 
paler beneath. North America .... melligaster Provancher (p. 68) 

Tergite 2 entirely smooth; costad abscissa of the basalis about one fifth as long as 
the mediad abscissa 

Hind coxa and hind femur black. Europe . . . politus Marshall (p. 48) 

Tergite 2 never as smooth as this ; almost always very strongly, coarsely sculptured ; 
if almost smooth (tjibodas) then the hind coxa in part, and the hind femur entirely, 
yellow; costad abscissa of the basalis at least one quarter as long as the mediad 
abscissa ............. 5 

Thorax entirely bright reddish fulvous; hypopygium without lateral, membranous 
creases and, seen from the side, with its apex produced to form a long, strongly 
sclerotized spine (Text-fig. 15) 

Preapical segment of the antenna tapered distally and about two and a half 
times longer than its basal width; wings strongly, evenly, smoky yellow. Queens- 
land. ......... magnificus Wilkinson (p. 66) 

Thorax black or blackish; hypopygium not thus produced at apex, though some- 
times it may be without lateral creases (grants-subgroup) .... 6 

Face either virtually impunctate or more often with an even punctation, there being 
no indication of transverse or almost transverse vermiculate rugosities towards 
sides. 

Vertex around the ocelli smooth, shining; claws simple .... 7 

Face much less shining and with a predominantly rugose sculpture and almost 
always with a clear indication of vermiculate rugosities towards sides; rarely the 
sculpture reduced to a vague, confused aciculation (tjibodas) or obsolescent 
(nerione) ............. 14 

Face smooth, except for minute setiferous punctures. 

Basal half of ventral surface of gaster bright yellow; flagellum long, thin, with 
somewhat outstanding pubescence; its preapical segment hardly less than twice 
as long as wide. Europe procerus Ruthe (p. 45) 

Face with distinct punctation .......... 

Face, anterior part of mesopleurum, mesosternum, with a heavy discrete puncta- 
tion; preapical segment of antenna about two and a half times longer than wide; 



36 G. E. J. NIXON 

hypopygium, except at extreme tip, heavily sclerotized all over, without trace of 
a fold along the middle line 

Very large species, c. 6 mm., with the whole of the hind leg blackish. E. 

Siberia reticulatus Shestakov (p.65) 

Face and more especially the anterior part of the mesopleurum and mesosternum 
with a much finer, sparser punctation, the surface at first sight sometimes appear- 
ing polished and almost impunctate; preapical segment of the antenna at most 
hardly twice as long as wide; hypopygium, except in one species which has the 
hind femur almost entirely red, tightly folded along the middle line in death and 
with more or less distinct lateral creases ....... 9 

9 Hairy part of ovipositor sheath very distinctly less than half as long as the hind 
tibia ; hind femur red, blackened at tip ; inner spur of the hind tibia not extending 
beyond basal two thirds of hind basitarsus. 

Very small species, c. 2-5 mm. without ovipositor. Europe opheltes sp. n. (p. 48) 

- Hairy part of ovipositor sheath at least about two thirds as long as the hind tibia ; if 
somewhat shorter, then the hind femur entirely dark ; inner spur of the hind tibia 
longer 10 

10 Hind femur entirely red. 

Antennal segments 15-17 somewhat barrel-shaped so that the apical antennal 
segments appear loosely articulated (Text-fig. 5). Europe 

Julvicrus Thomson (p. 45) 
Hind femur at least with a darkened tip . . . . . . . . n 

11 Hind femur entirely black .......... 12 

Hind femur red with a darkened tip ......... 13 

12 Hairy part of ovipositor sheath not more than two thirds as long as the hind tibia; 

posterior tangent to the anterior ocellus not, or hardly, cutting the posterior pair. 
Very dark species with heavily infumated wings; antennal segments 15-17 
cylindrical, with 1 7 varying from one and a half to one and two thirds times longer 
than wide. Europe curvicrus Thomson (p. 46) 

Hairy part of ovipositor sheath about three quarters as long as the hind tibia; 

posterior tangent to the anterior ocellus distinctly cutting the posterior pair 
(Text-fig. 14). Europe erro sp. n. (p. 46) 

13 Tergites with very narrow, pale, apical margin; tergite 3 with a yellowish spot on 

each side ; first abscissa of the discoideus not longer than the second ; setae of the 
median cell as dark towards base of cell as at apex. Europe obsepiens sp. n. (p. 47) 
Tergites entirely black ; no yellow spot on each side of tergite 3 ; first abscissa of the 
discoideus more or less distinctly longer than the second; setae of the median cell 
dark only within about apical quarter; elsewhere colourless and almost absent. 
Ireland ....... fulvicrus Thomson, var. A (p. 46) 

14 Hind coxa entirely, or in large part, yellow; (if the yellow colour is not sharply dis- 

crete, then the apical tergites are marked with yellow) . . . . . 15 
Hind coxa black or dark brown ; at most somewhat paler beneath or at apex (epagoges, 

brittoni) ............. 19 

15 Hind coxa entirely yellow. 

Gaster conspicuously pale-marked or banded with yellow on apical segments . 16 

Hind coxa darkened at base .......... 17 

1 6 Claws long, thin, evenly curved (Text-fig. 12); inner spur of the middle tibia not 

reaching apex of basal segment of tarsus; tergites 4-6 with weakly indicated, 

pale, longitudinal, median band. Europe . . . auriculatus Fab. (p. 55) 

Claws of ordinary form ; inner spur of the middle tibia reaching apex of basal segment 

of tarsus ; tergites 46 with dark, longitudinal, median band, sometimes so narrow 
that segments appear virtually yellow. 

Hind tarsus and apex of hind tibia deeply infuscate. North America 

leechi Walley (p. 65) 



REVISION OF MICROGASTER 



37 



17 Apical tergites conspicuously banded with yellow. 

Ventral surface of gaster entirely yellow; hind leg distal to coxa entirely reddish 
yellow; anterior half of mesoscutum with fine, shrivelled rugosity, stronger along 
the imaginary course of the notaulices; radius leaving stigma far distal to middle. 
North America ... ..... gelechiae Riley (p. 65) 

Apical tergites entirely dark .......... 18 

1 8 Gaster largely dull reddish; tergite 2 almost smooth; mesoscutum, except for its 

anterior declivity, polished, smooth-looking; hind femur entirely yellow; claws 

with minute, basal spine. Java ..... tjibodas Wilkinson (p. 67) 

Gaster virtually black; tergite 2 very coarsely rugose-reticulate; mesoscutum 

markedly dull, rugose almost all over; hind femur weakly infuscate at apex; 
claws with 2-3 teeth. North America. England pantographae Muesebeck (p. 62) 

19 Claws with a conspicuous, basal lobe (Text-fig. 9); head from in front markedly 

triangular (Text-fig. 29). 

Mesoscutum densely, heavily punctate all over and, at the origin of the notaulic 
courses, rugose-punctate. Europe. .... deductor sp. n.(p. 49) 

Claws never lobed, but sometimes toothed or pectinate; head from in front not 

triangular ............. 20 

20 Hind claw with one to two long teeth or two to four fine, close, black spines (Text- 

figs. 8, 6) 21 

Hind claw at most with a single, fine, black spine at base ..... 30 

21 Hind claw bent almost at right angles near base and with one to two strong teeth 

(Text-fig. 8) ; the distal tooth almost as pale and as long as that part of the claw 
distal to it. 

Preapical segment of the antenna more or less square in outline; hairy part of 
the ovipositor sheath fully three quarters as long as the hind tibia . . . 22 

Hind claw less bent and with two to four spine-like teeth that are somewhat darkened 

and do not reach the level of the claw-tip ; the claw, hence, simply pectinate (Text- 
fig. 6) 24 

22 Metacarp short, only about one and one third times longer than its distance from the 

apex of the radial cell; stigma rather short and broad (Text-fig. 25); at least the 
anterior half of the mesoscutum with much fine, shrivelled rugosity, the lines of the 
notaulices more strongly rugose ; tergite 2 very coarsely rugose-reticulate, without 
obvious longitudinal elements in the sculpture; slightly longer than 3, the dividing 
suture deep, somewhat rugose; tergite 3 with distinct traces of rugosity (vague 
rugose-punctation) ; inner spur of the hind tibia longer. North America 

congregatiformis Viereck 

Metacarp longer, at least twice as long as its distance from the apex of the radial cell ; 

stigma longer, narrower; anterior half of the mesoscutum more shiny and smoother; 
tergite 2 with weaker rugosity, in which there are obvious longitudinal elements; 
not longer than 3, the dividing suture smooth; tergite 3 virtually smooth; inner 
spur of the hind tibia shorter, only about three fifths as long as the hind basitarsus 23 

23 Flagellum bristly, the pubescence of segment 17 being equal to about one third the 

width of the segment (Text-fig, i); hind femur entirely red or with faint, dark 
patch above at base; no trace of short, longitudinal keel between the posterior 
ocelli. Europe ....... crassicornis Ruthe (p. 49) 

- Flagellum less bristly, the pubescence of segment 17 not more than one quarter the 
width of the segment; hind femur blackened at base and along upper margin; 
nearly always a very short, longitudinal keel between the posterior ocelli. Europe. 

areolaris Thomson (p. 49) 

24 Hypopygium heavily sclerotized all over and without a trace of lateral creases. 

Hind femur black; hind tibia deeply infuscate but slightly paler along dorsal 
surface; ovipositor sheath about half as long as the hind tibia; ovipositor thick, 
evenly curved. Mexico ....... nerione sp. n. (p. 68) 

ENTOM. 22, 2. 2 



38 G.E.J. NIXON 

Hypopygium more feebly sclerotized laterally thana long the middle line and always 

with some lateral creasing in the dead insect ....... 25 

25 First abscissa of the discoideus distinctly shorter than the second (Text-fig. 23) ; 

costad abscissa of the basalis hardly more than one quarter as long as the mediad 
abscissa ; hind claw with two pale spines ; ovipositor sheath about three quarters as 
long as the hind tibia. England ...... consors sp. n. (p. 50) 

First abscissa of the discoideus as long as, or longer than, the second ; costad abscissa 

of the basalis fully one third as long as the mediad abscissa; hind claw with two 

to four thickened spines .......... 26 

26 Hind coxa somewhat brownish beneath; middle coxa yellowish brown. 

Flagellum fulvous beneath; hind tibia reddish fuscous, becoming faintly darker 
towards apex and with yellowish, basal ring. North America epagoges Gahan (p. 52) 
Hind and middle coxa black . . . . . . . . . .27 

27 Metacarp more sharply defined, short, only a little longer than its distance from the 

apex of the radial cell; hind femur black; hairy part of the ovipositor sheath 
distinctly less than half as long as the hind tibia. 

Mesoscutum conspicuously punctate over anterior half. Europe tibialis Nees (p. 50) 
Metacarp less sharply defined, at least about one and a half times longer than its 
distance from the apex of the radial cell and then, canadensis, mesoscutum not 
obviously punctate in front; hind femur red, with at most the apex darkened; 
hairy part of ovipositor sheath fully two thirds as long as the hind tibia . . 28 

28 Metacarp about one and a half times longer than its distance from the apex of the 

radial cell ; first abscissa of the radius virtually straight ; hind tibia with weak, but 
distinct apical infuscation; hairy part of ovipositor sheath hardly two thirds as 
long as the hind tibia. North America . . . canadensis Muesebeck (p. 64) 
Metacarp about twice as long as its distance from the apex of the radial cell; first 
abscissa of the radius markedly convex on its distal side (Text-fig. 24) ; hind tibia 
with conspicuous, apical infuscation ; hairy part of the ovipositor sheath distinctly 
more than two thirds as long as the hind tibia ...... 29 

29 Scutellum coarsely punctate; the punctures large and separated by one to two 

diameters, except posteriorly; mesopleurum towards front with clearly defined, 

dull surface sculpture between its large, round punctures. Europe eupolis sp. n. (p. 51) 

Scutellum with at most scattered punctures along sides; mesopleurum smooth and 

shining between its punctures 

Species with deeply blackened hind tarsus and black tip to hind tibia, con- 
trasting with deep red hind femur. Europe. Mediterranean Region 

deprirnator Fab. (p. 51) 

30 Costad abscissa of the basalis hardly one quarter as long as the mediad abscissa. 

Gaster yellow on basal half beneath; antenna thin, with segment 16 fully twice 
as long as wide ; flagellum bristly ; hairy part of ovipositor sheath about one third 
as long as the hind tibia. Europe. North America . novicius Marshall (p. 53) 

Costad abscissa of the basalis one third to one half as long as the mediad abscissa . 31 

31 Hind tibia black with white basal ring; fore wing hyaline, sharply infuscate distal to 

middle of radial cell and with dark band enveloping first abscissa of radius 

Hind spurs white and hind tarsus as black as its tibia ; scape somewhat reddish ; 
costad abscissa of the basalis at most about one third as long as the mediad 
abscissa (Philippines) ; much shorter than this (type series, Borneo) . Indo- 
Australian Region ...... kuchingensis Wilkinson (p. 66) 

Hind tibia never black with white, basal ring; hyaline and infuscate areas of fore 

wing never as sharply discrete as this . . . . . . . .32 

32 Eyes strongly convergent below (Text-fig. 32). 

Small species, c. 2-3 mm., with the mesoscutum confusedly rugose-punctate 
almost all over and the hind femur bright reddish yellow with darkened apex. 
Macedonia ......... phryne sp. n. (p. 63) 



REVISION OF MICROGASTER 



39 



Eyes never as strongly convergent below as this . . . . . . 33 

33 Hypopygium very short, heavily sclerotized all over, not tightly folded along the 

middle line and without trace of lateral creases. 

Antenna very long, the preapical segment fully twice as long as wide; surface 
between and around the ocelli with some sort of sculpture ; hairy part of ovipositor 
sheath at most nearly half as long as the hind tibia, its apical edge with a row of 
stiff bristles that contrast with the longer hairs below them (Text-fig, n); this 
tuft poorly defined in the North American brittoni but this species has the dorsum 
of the gaster conspicuously yellow-marked ....... 34 

Hypopygium usually considerably longer and not thus evenly and heavily sclerotized 
so that, in the dead insect, it tends to be tightly folded along the middle line and 
usually shows one or more lateral creases ; (creases hardly evident in fischeri but 
this species is less than 2-5 mm.) ......... 36 

34 At least tergite (2 + 3) heavily marked with yellow ; hairy part of ovipositor sheath 

almost half as long as the hind tibia. North America . brittoni Viereck (p. 54) 

Dorsum of gaster entirely black; hairy part of ovipositor sheath only about one 

third as long as the hind tibia ......... 35 

35 Mesoscutum, at least over anterior two thirds, with large, well separated punctures ; 

hairs of the head and mesoscutum unusually long and conspicuous; hind tibia 

infuscate at apex. Europe grandis Thomson (p. 53) 

Mesoscutum without distinct punctation, but strongly rugulose in front; hairs of 
the head and mesoscutum short, inconspicuous ; hind tibia uniformly red through- 
out. Europe ......... acilius sp. n. (p. 53) 

36 Claws unusually large and conspicuous (Text-fig. 10) ; mesoscutum dull, densely 

rugose-punctate almost everywhere; apart from the medial, polished area, the 
mesopleurum in front and below is densely rugose-punctate; posterior tangent to 
the anterior ocellus not touching the posterior pair; the rugosity of the meso- 
pleurum extends over the area posterior to the sternaulic pit. Europe 

deceptor sp. n. (p. 55) 

Claws normal ; mesoscutum never as dull or as extensively sculptured as this, though 
there may be conspicuous, shrivelled rugosity along the imaginary course of the 
notaulices ; mesopleurum shiny and with at least a somewhat superficial, less close 
punctation ; ocelli in a lower triangle, the posterior tangent to the anterior ocellus 
at least touching the posterior pair; if the mesopleurum shows coarse rugosity, 
then this does not extend over the area posterior to the sternaulic pit . . 37 

37 First abscissa of the discoideus very distinctly longer than the second (Text-fig. 22) 

Mesoscutum rugulose almost all over; hind femur infuscate at tip; hind tibia 
deeply infuscated on about apical third; basal half of ventral surface of gaster 
bright yellow. Austria ....... caris sp. n. (p. 61) 

First abscissa of the discoideus never distinctly longer than the second ... 38 

38 Ovipositor sheath very long, its hairy part as long as the hind tibia. 

Both hind femur and hind tibia with blackened tip ; flagellum very slender, with 
bristly pubescence and the preapical segment at least one and a half times longer 
than wide. Europe ....... subcompletus Nees (p. 56) 

Ovipositor sheath at most three quarters as long as the hind tibia and then the hind 

femur is not tipped with black ......... 39 

39 Distance between the anterior ocellus and a posterior ocellus about equal to the 

diameter of the anterior ocellus. 

Surface between and around the ocelli almost smooth ; hind femur entirely red ; 
hind tarsus almost as red as its tibia; hairy part of ovipositor sheath about two 
thirds as long as the hind tibia. Europe . . laeviscuta Thomson (p. 56) 

This distance usually markedly less than the diameter of the anterior ocellus ; if not, 
then the surface around the ocelli shows some striation radiating outwards from 
the ocellar triangle or the hind femur is blackened at least at apex ... 40 



4 o G. E. J. NIXON 

40 Tergite i tending to become smooth and polished over the greater part of its horizon- 

tal surface; hind tibia tricoloured, deeply infuscate over distal half, red at middle 
and whitish on about basal fifth 

Mesoscutum smooth, shining, even in front; scape bright reddish. S. Europe 

asramenes sp. n. (p. 62) 

Tergite i strongly sculptured everywhere; hind tibia not tricoloured . . " . 41 

41 Hypopygium very strongly developed, long, acute, extending well beyond the apex 

of the gaster (Text-fig. 18). 

Hind femur blackish ; hind tibia becoming progressively more strongly infuscate 

from base to apex; flagellum very bristly, with the preapical segment about one 

and one third times longer than wide; metacarp nearly three times as long as its 

distance from the apex of the radial cell. Finland . . ductilis sp. n. (p. 58) 

Hypopygium rarely approaching this length and then the hind femur is entirely or 

predominantly reddish or yellowish and the flagellum is not noticeably bristly . 42 

42 Hind femur varying from entirely black to red with usually at least a dark 

streak above at base; if entirely red, then wings virtually hyaline. 

Small species, at most about 3 mm., without ovipositor .... 43 

Hind femur rarely blackened and then the species are considerably larger . . 44 

43 Larger, about 3 mm. without ovipositor; hind tibia pale reddish, without trace of 

apical infuscation 

Wings virtually hyaline; antenna short with segments 16-17 and often 15 not or 
only very slightly longer than wide; ovipositor evenly curved throughout (Text- 
fig. 17), thinner than in laeviscuta. Europe. North America. 

hospes Marshall (p. 57) 

Smaller, about 2-3 mm. without ovipositor; hind tibia becoming infuscate in about 

apical third 

Hind femur infuscate almost throughout or entirely ; ovipositor sheath very short, 
hardly more than half as long as the hind tibia; propodeum having a somewhat 
flattened appearance. Europe ...... fischeri Papp (p. 64) 

44 Anterior half of mesoscutum very distinctly punctate, though the punctures tend to 

become crowded and form rugose-punctation at the origin of the imaginary 
notaulic courses; anterior part of the mesopleurum somewhat coarsely rugose. 

Wings strongly infumated; at least the apex of the hind femur strongly infus- 
cated ; surface around the ocelli polished, smooth ; ovipositor sheath with numerous, 
outstanding bristles. Austria ..... famulus sp. n. (p. 59) 

Anterior half of mesoscutum without such distinct punctation though there may be 

considerably rugosity along the course of the notaulices; with punctation feebly 
indicated at front of middle lobe; anterior part of mesopleurum with a prevailing 
sculpture of discrete punctation that may be very weak or superficial; if the 
sculpture here is somewhat coarse and not resolvable as punctation, then the 
wings are virtually hyaline .......... 45 

45 Tergite 2 at most about two and a half times wider than long (Text-fig. 33) and dis- 

tinctly longer than 3. 

Hairy part of the ovipositor sheath a little less than half as long as the hind 
tibia; hind tarsus almost as red as its tibia; head considerably sculptured around 
and between the ocelli; flagellum long, tapering, the preapical segment fully one 
and a half times longer than wide; front part of mesoscutum strongly rugose or 
rugose-punctate. Europe ...... sticticus Ruthe (p. 59) 

Tergite 2 about three times wider than long and not distinctly longer than 3 ; hairy 

part of ovipositor sheath distinctly more than half the length of the hind tibia . 46 

46 Femora predominantly yellowish; smaller, c. 3 mm. without ovipositor. North 

America ......... peroneae Walley (p. 58) 

Femora predominantly reddish; larger, c. 4-2 mm. without ovipositor . . . 47 



REVISION OF MICROGASTER 41 

47 Ovipositor abruptly downcurved in about apical third; no transverse striation to 
the side of the ocelli; spines of the outer side of the hind tibia rather short and 
sparse, especially at about middle ; sculpture at the origin of the notaulic courses 
fine, weak. Europe ...... parvistriga Thomson (p. 61) 







II 13 

FIGS. 1-13. Microgaster, $: Apical flagellar segments of i, crassicornis Ruthe. 2, sticticus 
Ruthe. 3, erro sp. n. 4, curvicrus Thomson. 5, fulvicrus Thomson. 6, deprimator 
Fab., hind claw. 7, laeviscuta Thomson, inner spur of left hind leg. 8, crassicornis 
Ruthe, hind claw. 9, deductor sp. n., hind claw. 10, deceptor sp. n., hind claw, n, 
grandis Thomson, ovipositor sheath (lateral). 12, auriculatus Fab., middle claw. 13, 
procerus Ruthe, ovipositor sheaths (dorsal). 



42 G. E. J. NIXON 

Ovipositor weakly but evenly curved throughout ; considerable striation to the side 
of the ocelli; spines of the outer side of the hind tibia longer, more numerous; 
sculpture at the origin of the notaulic courses very coarse, the sculpture extending 
much further onto the disc than in parvistriga. 

Hind femur red with at most the faintest trace of apical infuscation. Europe 

alebion sp. n. (p. 60) 




FIGS. 14-19. Microgaster $: 14, erro sp. n., ocellar triangle. 15, magnificus Wilkinson, 
apex of gaster (lateral). 16, nerione sp. n., hypopygium and ovipositor (lateral). 17, 
hospes Marshall, apex of gaster (lateral). 18, ductilis sp. n., apex of gaster (lateral). 
19, deceptor sp. n., head from above. 



REVISION OF MICROGASTER 



43 



20 




21 



22 




25 



FIGS. 20-25. Microgaster, $ : Distal half of fore wing of 20, deceptor, sp. n. 21, tibialis 
Nees. 22, caris, sp. n. 23, consors sp. n. 24, deprimator Fab. 25, congregatiformis, 
Viereck. 



44 



G. E. J. NIXON 



26 




31 



FIGS. 26-33. Microgaster, <j>: Distal part of fore wing of 26, politus Marshall. 27, fulvicrus 
Thomson. 28, grandis Thomson. 29, deductor sp. n.; head (from in front). 30, laevi- 
scuta Thomson, distal part of fore wing. 31, rugulosus Nees, gaster (dorsal). 32, phryne 
sp. n., head (from in front). 33, stictious Ruthe, gaster (dorsal). 



REVISION OF MICROGASTER 45 

Microgaster procerus Ruthe 

(Text-fig. 13) 
Microgaster procerus Ruthe, 1860 : 115. 

?. Basal half of ventral surface of gaster yellow. Maxillary palpi pale yellow. Hind 
femur red with darkened tip; hind tibia at apex and hind tarsus, infuscated. Fore wing faintly 
smoky; metacarp paler than the stigma. 

Head above smooth, polished. Flagellum thin, somewhat pale beneath, distinctly bristly; 
the preapical segment almost twice as long as wide. 

Mesoscutum having a highly polished appearance, even in front, where there are traces of 
very superficial punctation; no rugosity at the origin of the notaulic courses. Costad abscissa 
of the basalis fully two-fifths as long as the mediad abscissa; first abscissa of the discoideus 
distinctly a little longer than the second. Inner spur of the hind tibia fully three quarters 
as long as the hind basitarsus. 

Tergite i somewhat narrowly triangular, its sculpture towards sides tending to be predomin- 
antly longitudinal. Hairy part of ovipositor sheath about half as long as the hind tibia; seen 
from above, the hairs of the sheaths are particularly long and outstanding (Text-fig. 13). 

o\ Like the female in colour and sculpture. 

Length: $ $, c. 5 mm. without ovipositor of female. 

Material examined: FINLAND. GERMANY. IRELAND. 3^,4$. 

Distinct among those species with non-rugose face on general reduction of sculpture 
together with colour, especially that of the ventral surface of the gaster. 

The single Irish male I have seen differs from two Finnish males in that the 
middle femur is streaked with black above and the base of the gaster is not bright 
yellow; this may be due to discoloration. 

Microgaster fulvicrus Thomson 
(Text-figs. 5, 27) 

Microgaster fulvicrus Thomson, 1895 : 2240. 

$. Maxillary palpi brownish, the basal segments almost black. Hind femur entirely red; 
hind tibia with darkened tip, sometimes hardly noticeable; hind tarsus deeply infuscated. 
Fore wing somewhat darkened; the radial cell contrastingly paler than the large third cubital 
cell adjacent to it. 

Head above smooth, polished. Apical flagellar segments (Text-fig. 5). 

Front part of mesoscutum with faint to rather sharp, fine punctation. Scutellum virtually 
impunctate. Costad abscissa of the basalis about one third as long as the mediad abscissa; 
stigma rather broad, emitting the radius virtually at middle (Text-fig. 27). 

Tergite 2 as coarsely rugose as tergite i. Hairy part of the ovipositor sheath slightly more 
than half as long as the hind tibia; outstanding hairs fewer than in procerus (cf. Text-fig. 13). 

cJ. Like the female in colour and associated with it on the evidence of the short series bred 
from D. ocellana. Unless careful attention is paid to the sculpture of the face, the male is 
virtually indistinguishable from the males of several other species. 

Distribution: ENGLAND. FINLAND. IRELAND. SWEDEN. 

Host: Depressaria ocellana Fab. (Oecophoridae) , in England, Westmorland, 
Witherslack. Parasites emerge in May from hosts collected previous June. 
Type in the Entomological Institute, Lund, Sweden. Examined. 

ENTOM. 22, 2. 2 



46 G. E. J. NIXON 

Microgaster fulvicrus, var. A. Seven females from counties Wicklow and Dublin 
in Ireland. These differ from typical fulvicrus as follows : the apex of the hind 
femur is conspicuously blackened; the two preapical segments of the antenna are 
slightly more elongate; the anterior part of the mesoscutum is more densely and 
more sharply punctate; the first abscissa of the discoideus is usually distinctly 
longer than the second; the hairs of the median cell are darkened only in about 
apical quarter of cell; elsewhere the cell appears bare, the scattered setae being 
sparse and colourless, whereas in typical fulvicrus, the setae are dark wherever 
they occur. 

Microgaster curvicrus Thomson 

(Text-fig. 4) 
Microgaster curvicrus Thomson, 1895 : 2242. 

$. Middle and hind femur entirely black; front femur yellow on about distal half; hind 
tibia dusky red, darkened at tip but the infuscation not sharply discrete and more extensive 
on inner side. Radial cell more or less as dark as the third cubital cell adjacent to it. 

Flagellum differs from that of fulvicrus in that the two preapical segments are more elongate 
and evenly cylindrical (Text-fig. 4). 

Anterior half of mesoscutum very shiny, tending to be sharply and distinctly punctate. 
Costad abscissa of the basalis a little longer than in fulvicrus so that the discoidal cell is slightly 
more elongate; abscissa i of the radius slightly more obliquely placed than in fulvicrus and 
slightly more curved. 

Ovipositor sheath as in fulvicrus. 

<$. I associate with the female numerous males, caught in certain localities, with females, 
in England (Kent: Wye and Dartford Heath) and showing the same colouration. 

Distribution: ENGLAND. FINLAND. SWEDEN. A common species ; some seventy- 
five specimens examined. 

Host : Depressaria pallor ella Zeller (Oecophoridae) . Host collected July ; parasites 
emerge following May. In southern England, Mr. R. L. E. Ford has swept the 
species commonly from broom (Cytisus] in June. 

Type in the Entomological Institute, Lund, Sweden. Examined. 

Microgaster curvicrus, var. A. Three females from SWITZERLAND: Valais, Les 
Hauderes and Ferpecle, vi and CZECHOSLOVAKIA: Tatra Mts., v. These females 
have the hind tibia blackened throughout; the apical segments of the flagellum 
are slightly more slender than in typical examples and slightly more bristly; the 
hairy part of the ovipositor sheath is slightly less than half as long as the hind tibia. 

Superficially, curvicrus s.l. is much like tibialis but differs strikingly in the sculp- 
ture of the face. 

Microgaster erro sp. n. 

(Text-figs. 3, 14) 

$. Extremely like curvicrus and differing from it by little more than the characters given 
in the key. The most obvious difference is the longer ovipositor of erro but the difference in 
the shape of the ocellar triangle may have equal importance. 



REVISION OF MICROGASTER 47 

Whereas in curvicrus, the apical infuscation of the hind tibia tends to spread over almost a 
third of the tibia, in erro it is restricted to the extreme apex but tends to be faint and in one 
of the three females from Semsjarvi is absent. 

Flagellum not at all tapered apically. (Text-fig. 3). First abscissa of the radius slightly 
more curved than in curvicrus. 

Length: larger than curvicrus, c. 6 mm. without ovipositor. 

Type $. FINLAND: Lemland (Hellen), Helsinki Museum. 

FINLAND: Bergo, i $, 17- 28 . vi . 1946 (Hellen); Carelia, Semsjarvi, 3 $, 6.vi.i943 
(Hellen); Helsingfors, 3 $ (W. Nylander). SWITZERLAND: Valais, Les Hauderes, 
4,000-8,000 ft., 3 $, vi.i935 (/. E. & R. B. Benson). The seven Finnish females 
are paratypes. 

The three females from Switzerland are somewhat doubtful; they do not have 
the flagellar segments so bristly as in the type series. In one of them, the hind 
tibia is infuscate almost throughout ; and in colour is thus exactly like var. A of 
curvicrus. 

In curvicrus the ovipositor itself is strongly curved, rather thick and distinctly 
shorter than the hind tibia; in erro, it is longer than the hind tibia, less thick and 
less curved. 

I have examined a female of what I believe to be this species from Cyprus; it 
differs from the Finnish specimens as follows: hind tibia without a trace of apical 
infuscation. Flagellum somewhat tapered apically, with the two preapical segments 
slightly longer, though this appearance may be deceptive for the segments are 
somewhat collapsed. Ocelli in a higher triangle, the posterior tangent to the an- 
terior ocellus not cutting the posterior pair. Anterior part of the mesososcutum 
more shining and having altogether a much more polished appearance, the punc- 
tation being almost absent. 

The combination of the above differences may constitute specific validity but 
needs to be seen in further specimens before a decision on this point can be reached. 

Microgaster obsepiens sp. n. 

$. Wings strongly darkened; stigma suffused with reddish on about basal third. All the 
femora red, except that the hind femur is faintly tipped with black; hind tibia also faintly 
darkened at tip and yellowish on about basal fifth. Basal half of ventral surface of gaster 
reddish yellow; a reddish yellow spot on each side of tergite 3. 

Flagellum as in curvicrus, the apical segments tightly articulated and having an evenly 
cylindrical appearance; its pubescence so short as to be virtually not outstanding. 

Front part of mesoscutum sharply and quite strongly punctate. Front femur slightly less 
thick and slightly longer than in both curvicrus and fulvicrus; inner spur of the hind tibia 
relatively a little shorter than in both curvicrus and fulvicrus, not more than two thirds as 
long as the hind basitarsus. Radius leaving stigma more obviously distal to middle than in 
both fulvicrus and curvicrus ; first discoidal cell exactly as in curvicrus. 

Ovipositor sheath slightly more than half as long as the hind tibia. Tergites with thin, 
pale apical margin. 

Length: 5 mm. without ovipositor. 

Type $. GERMANY: Berlin, Finkenburg (Hellen), Helsinki Museum. 



48 G. E. J. NIXON 

In colour, this species approaches procerus more closely than it does either fulvicrus 
or curvicrus but differs from Ruthe's species in having very distinct punctation on 
both face and mesoscutum ; the flagellum is also considerably shorter than in procerus. 

Microgaster opheltes sp. n. 

$. Hind femur red with dark tip; hind tibia also red with dark tip; middle femur with 
faint dark streak above. 

Flagellum without bristly pubescence; preapical segment about one and one third times 
longer than wide. 

Mesoscutum very shiny, even in front, and here with only a weak trace of punctation; much 
weaker than in both curvicrus and fulvicrus. Costal abscissa of the basalis between one third 
and one half as long as the mediad abscissa; first discoidal cell elongate, as in curvicrus; first 
abscissa of the radius virtually straight and leaving the rather broad stigma only very slightly 
beyond middle. Hind spurs whitish, the inner one not extending beyond basal two thirds 
of hind basitarsus. 

Tergite 2 weakly sculptured (compared with most species of the genus), distinctly shorter 
than 3. Ovipositor sheath (hairy part) distinctly less than half as long as the hind tibia, ca. 

3 : 7- 

Length: c. 2-5 mm., without ovipositor; a small species. 

Type $. Europe, YUGOSLAVIA: Macedonia, Lake Ochrid, i6.vi.i958 (R. L. 
Coe), B.M. (N.H.). 

Paratype <j>. NORTHERN IRELAND: Co. Down, Cranfield Pt, i.vii.i957 (A. W. 

Stelfox) . 

Size, colour and short ovipositor are the main features of this species. The 
usefulness of the other characters mentioned must be accepted with caution since 
they are based only on two specimens. 

Microgaster politus Marshall 

(Text-fig. 26) 
Microgaster po litus M arshall, 1885 : 260. 

$. Legs predominantly brownish, the hind femur almost black. Wings more or less hyaline. 

Face smooth-looking, almost polished and with only the merest trace of punctation. Pre- 
apical segment of flagellum about one and one third times longer than wide. 

Mesoscutum highly polished. Suture between scutellum and mesoscutum reduced to a very 
narrow, foveolate groove. Hind spurs whitish, the inner one reaching only a little beyond 
the middle of the hind basitarsus. Fore wing (Text-fig. 26). 

Ovipositor sheath nearly three quarters as long as the hind tibia. 

<J. Like the female in colour and sculpture. 

Length: <J <j>, c. 3 mm. without ovipositor of female. 

Distribution: ENGLAND. IRELAND. SWEDEN. Not common. 
Type in the B.M. (N.H.). 

An easily recognized species. Apart from the polished second tergite, the short 
costad abscissa of the basalis is very characteristic. 



REVISION OF MICROGASTER 49 

Microgaster deductor sp. n. 

(Text-figs. 9, 29) 

A highly aberrant species on account of its lobed claws (Text-fig. 9). 

$. Hind femur entirely red; middle femur dark beneath; hind tibia red throughout. Wings 
almost hyaline. 

Head from in front rather narrowly triangular (Text-fig. 29). Pubescence of flagellum 
hardly noticeable; preapical segment about one and a quarter times longer than wide. 

Mesoscutum densely punctate almost all over and coarsely rugose-punctate at the origin 
of the notaulic courses. Scutellum closely punctate along each side. Mesopleurum with large, 
conspicuous punctures almost everywhere. 

Tergite 2 fully as long as 3. Tergites beyond 2 unusually densely and extensively hairy. 
Ovipositor sheath about three quarters as long as the hind tibia. 

<$. Like the female but the palpi almost black and the middle femur more extensively 
blackened at base. The lobe of the claws is less well developed than in the female but will 
nevertheless distinguish this male from all others available for consideration in this synopsis. 

Length: $ $, c. 5-2 mm. without ovipositor of female. 

Type $. FINLAND, Ivalo (Helleri), Helsinki Museum. 

Paratypes. Same data : i <, i $. LAPLAND : Tornekrask, i 9- 

In the Lapland female, the hind femur is darkened at base above and the middle 
and front femora are more extensively blackened at base than in the two Finnish 
females. The colour of the legs is likely to be variable. 

An unmistakable species on account of the structure of the claws. 

Microgaster crassicornis Ruthe 

(Text-figs, i, 8) 
Microgaster crassicornis Ruthe, 1860 : 124. 

This species has been confused in the literature with some of the other species 
with equally red legs but because of its toothed claws crassicornis could actually 
be confused only with areolaris. The essential differences between the two species 
have been given in the key. 

$. The flagellum is more bristly than in any other species known to me and this, combined 
with the shortness of the apical segments, provides one of the most useful features for recognizing 
the species (Text-fig, i). 

Mesoscutum in front without punctation but there is considerable rugosity at the origin 
of the notaulic courses. Inner spur of the hind tibia hardly equal to two thirds the length 
of the hind basitarsus. Ovipositor sheath fully three quarters as long as the hind tibia. 

Distribution: ENGLAND. FINLAND. SWEDEN. GERMANY 4 $. 
Host: Eupithecia denotata Guenee (now pimpinellata Hiibnagel) (Geometridae), 
England, Lyle Coll. in B.M. Bred from same host in Germany, Hinz Coll. 

Microgaster areolaris Thomson 
Microgaster areolaris Thomson, 1895 : 2240. 

?. The legs are altogether darker than in crassicornis, the basal and apical infuscation of 
the hind femur being a conspicuous feature of all the specimens I have examined. 



50 G. E. J. NIXON 

Flagellum considerably longer than in crassicornis. Face somewhat gabled along the middle ; 
towards the antennal sockets, the keel is more prominent than in crassicornis. 

Sculpture of tergite 2 very variable and on the whole very weak; in one of two Scottish 
examples, the surface is almost smooth; when clearly present, the sculpture is predominantly 
longitudinal. 

Metacarp almost three times as long as its distance from the apex of the radial cell, a little 
longer than in crassicornis. Claws slightly weaker than in crassicornis. 

<J. Ten males that I associate with this species (Poland, 8), (Switzerland, 2) have the claws 
toothed like the female ; in all of them the middle femur is blackened and in the two specimens 
from Switzerland (Valais) the hind femur is almost entirely black. 

Material examined: N. ENGLAND. FINLAND. SCOTLAND. SWEDEN. SWITZER- 
LAND. 

This is perhaps a Northern species. 

Microgaster consors sp. n. 

(Text-fig. 23) 

$. Hind femur red with infuscation at base beneath (type) or entirely blackish; hind tibia 
without apical infuscation. Wings only faintly tinted, almost hyaline. 

Flagellum not bristly; preapical segment almost square in outline. 

Mesoscutum on the whole smooth-looking but finely rugose at the origin of the notaulic 
courses. Propodeum unusually finely and evenly rugose and with only a very weak, medial 
keel. Costad abscissa of the basalis short, not much more than one quarter as long as the 
mediad abscissa; stigma somewhat broad; metacarp fully three times as long as its distance 
from the apex of the radial cell (Text-fig. 23). 

Hairy part of the ovipositor sheath fully three quarters as long as the hind tibia; apex of 
the ovipositor with two to three very small serrations, as seen from above. 

Length: c. 3-8 mm. without ovipositor. 

Type <j>. ENGLAND: S. Devon, Kingsteignton, Stark's Pond, 24. .1942 (/. F. 
Perkins), B.M. (N.H.). 

Paratypes. ENGLAND: Surrey, Byfleet, i $, 9-11. .1947 (R. B. Benson); Herts., 
Brickett Wood, i $, 20. .1936 (R.B.B.). 

I associate with the females seven males, on the strength of wing and claw charac- 
ters. These were all captured in May (England, Herts., Brickett Wood ; Bucks., 
Farnham Common; Surrey, Wimbledon Common). With one exception (Brickett 
Wood) all have the hind femur infuscate throughout, that is, very dark brown. 

This species is largely characterized by the shortness of the first abscissa of the 
discoideus, and long ovipositor sheath. 

Microgaster tibialis Nees 

(Text-fig. 21) 

Microgaster tibialis Nees, 1834 : 168. 
Microgaster pluto Morley, 1936 : 211, syn. n. 

$. Rather small, very dark species with dusky reddish hind tibia, infuscated on about 
apical third; both middle and hind femur entirely black or blackish. Wings strongly smoky. 



REVISION OF MICROGASTER 51 

Antenna rather short, not bristly; preapical segment at most about one and a quarter times 
longer than wide. 

Mesoscutum conspicuously punctate, the punctures tending to fade out at about posterior 
third. Scutellum with a few scattered punctures. Costad abscissa of the basalis between 
one third and one half as long as the mediad abscissa; first abscissa of the discoideus fully 
as long as the second; first abscissa of the radius leaving the stigma considerably distal to 
middle and very obliquely placed (Text-fig. 21). 

N. W. EUROPE. Common. 

Host : Gracillaria tringipennella Zeller (Gracillariidae) ; Peronea aspersana (Hiibner) 
(Tortricidae) . 

In size and colour this species is much like curvicrus but easily separated by the 
sculpture of the face, among several other characters. 

Microgaster deprimator Fab. 

(Text-figs. 6, 24) 

Ichneumon deprimator Fab., 1798 : 227. 
Microgaster australis Thomson, 1895 : 2240, syn. n. 

$. This large species is almost recognizable to the naked eye by its deeply infumated wings 
and strongly blackened hind tarsus, contrasting sharply with the intensely red hind femur. 
In one female (Switzerland, Miistertal, 1,400 m.) the hind femur is weakly infuscated at base; 
it is possible that more extensive darkening may occur in specimens taken at high altitudes. 

Whole of median part of face with simple, though rather coarse punctation. Flagellum 
thick, markedly tapered apically; its pubescence so short as to be almost imperceptible; pre- 
apical segment from one and one third to one and a half times longer than wide. 

Mesoscutum much less noticeably punctate than in tibialis. Mesopleurum, except for a 
small posterior area, closely punctate. Lateral keel of the propodeum very strongly raised 
behind so that the surface between it and the medial keel is almost concave. 

o*. Claws pectinate like the female (Text-fig. 6). Colour usually similar but the hind tarsus 
often not contrastingly darkened. Propodeum equally characteristic. 

Length: Q* $, c. 5-5 mm. without ovipositor of female. 

Distribution: CENTRAL EUROPE as far as SPAIN and TURKEY. CYPRUS. PERSIA. 
N. MONGOLIA (i<3). Thomson records australis from N. Italy, not N. Europe, as 
given by Fahringer (1937 : 334). 

Microgaster eupolis sp. n. 

$. Apart from the essential differences given in the key, this species may be compared with 
deprimator as follows: 

Wings much less smoky. Distribution of light and dark leg-colouration much as in depri- 
mator, except that the hind tarsus, especially in the type, is considerably reddened. 

Surface immediately surrounding the ocelli considerably rugose; temples coarsely rugose in 
comparison with deprimator. Apical segment of antenna less narrowly conical but since only 
two females are available, this may have no significance. 

Mesoscutum coarsely punctate; course of notaulices marked by a band of coarse rugose- 
punctation. Propodeum exactly as in deprimator. Hind coxa on lower half very closely, 
strongly punctate. 

Length: c. 5-2 mm. without ovipositor. 



52 G. E. J. NIXON 

Type $. AUSTRIA: Tyrol, Seiseralpe, 1893, Naturhistorisches Museum, Vienna. 

Paratype. AUSTRIA: Innsbruck, Hottinger Berg, 1,700 m., i $, 20.viii.i95o 
(Pechlauer) . 

I associate with the above two females a single male from ITALY (Dolomiti, Passo 
Campolongo, 6.ix.i954 (G. J. Kerrich)}. This has the characteristic mesoscutal 
sculpture of the females and the densely punctate hind coxa, but the punctures 
of the scutellum are smaller and hence more widely spaced; the punctation of the 
mesopleurum is equally strong and conspicuous but the interspaces show hardly 
a trace of surface sculpture. 

Microgaster epagoges Gahan 

Microgaster epagoges Gahan, 1917 : 197. 

$. Flagellum fulvous throughout but darker above. Hind femur red with faint apical 
infuscation. Basal half of ventral surface of gaster yellowish. 

Flagellum not bristly; preapical segment about one and one third times longer than wide. 

Mesoscutum not punctate in front but considerably roughened, the sculpture still stronger 
at the origin of the notaulic courses. Hind claw with three to four spine-like teeth. 

Sculpture of tergites i and 2 particularly coarse; 3 also considerably sculptured. Hypopy- 
gium thin, membranous, yellowish and with numerous lateral creases. 

Length: c. 3-5 mm. without ovipositor. 

Material examined: U.S.A., Tennessee, Nashville, i $ paratype, in B.M. (N.H.). 

Host: Epagoge sulfur eana Clemens (Tortricidae) . 

I cannot be sure that this species, represented by the single paratype in the 
British Museum, is really distinct from Microgaster phthorimaeae, as represented by 
a single female in the B.M.(N.H.), determined by Muesebeck and evidently 
one of three specimens recorded by him (1922 : 41) from California, Pasadena but 
not part of the type series. These three examples of phthorimaeae were bred from 
Phthorimaea operculella Zeller. 

Apart from slight differences in colour the broken flagellum of phthorimaeae is 
pale only beneath I can find no differences of particular significance. Certainly 
there is a short keel beneath the antennal insertions in epagoges, as specially men- 
tioned by Muesebeck and only the merest trace of one in the single specimen of 
phthorimaeae but in my opinion this is a feature of only doubtful specific value. 



Microgaster phthorimaeae Muesebeck 
Microgaster phthorimaeae Muesebeck, 1922 : 40. 

See discussion under epagoges. Not included in key because not sufficiently differ- 
entiated from epagoges on the material available to me in the B.M. (N.H.). 

Distribution: U.S.A., California, Oxnard (type locality); Pasadena, i $ paratype, 
in B.M. (N.H.). 

Host: Phlyctaenia ferrugalis Hiibner (Pyralidae), host of type series; Phthorimaea 
operculella Zeller (Gelechiidae) . 



REVISION OF MICROGASTER 53 

Microgaster novicius Marshall 

Microgaster novicius Marshall, 1885 : 252. 

Microgaster swammerdamiae Muesebeck, 1922 : 37, syn. n. 

$. This is a small species, characterized essentially by three features : long, thin antenna 
with bristly flagellum; very short costad abscissa of the basalis and very short ovipositor. 

The pale parts of the legs are yellow rather than red; hind spurs whitish; hind femur faintly 
darkened at tip above. 

Metacarp almost four times as long as its distance from the apex of the radial cell. 

Ovipositor sheath only about one third as long as the hind tibia. Hypopygium rather 
heavily sclerotized but still tightly folded along the middle line in death and without lateral 
creases. 

Length: c. 3 mm. without ovipositor. 

Distribution : SCOTLAND (Lanark, Cadder) (typeinB.M.(N.H.)). FINLAND, Kempele, 
i $, Flutis, i $ (hind femur of this female entirely infuscate and fine rugosity of 
the notaulic courses extending further backwards than in other females). NORTH 
AMERICA ($, paratype of swammerdamiae, in B.M.(N.H.)). 

Host: Swammerdamia castanea Busk (Hyponomeutidae), host of type series of 
swammerdamiae in North America. 

Microgaster grandis Thomson 

(Text-figs, n, 28) 
Microgaster grandis Thomson, 1895 : 2242. 

$. Hind femur varying from entirely red to entirely black. 

Flagellum long, markedly tapering to apex; preapical segment fully twice as long as wide. 

Hairs of the median cell sparse, becoming still sparser to widely absent along the medius 
side of the cell; radius leaving stigma very obviously beyond middle; stigma rather narrow 
(Text-fig. 28) ; costad abscissa of the basalis fully half as long as the mediad abscissa. 

Hairy part of the ovipositor sheath about one third as long as the hind tibia. 

($. Like the female in hairiness and punctation of mesoscutum. The median cell shows 
the same sparseness of hairs. 

Length: $ $, 4-5 mm. without ovipositor of female. 

Distribution: ENGLAND. FINLAND. IRELAND. SWEDEN. 

Host: Cnephasia chrysanthemana Duponchel (Tortricidae), i $, England, Kent, 
Bexley, collected May, emerged June same year (R. L. E. Ford). Aphelia viburniana 
Fab. (Tortricidae), i $, England, Yorks., Ling Hill, emerged 2.vii.ig65 (/. Bradley}; 
the hind femur of this female is entirely black. 

This is one of the more distinct species, characterized largely by the long hairs 
and very characteristic punctation of the mesoscutum. 

Microgaster acilius sp. n. 

$. Has a hypopygium similar to that of grandis, with which, in addition to the characters 
given in the key, it may be compared as follows: 

Hind femur red, except for faint infuscation at extreme tip. 
Upper surface of the head with more rugosity. 



54 G. E. J. NIXON 

The coarse rugosity at the origin of the notaulic courses extends almost to middle of disc; 
similar rugosity occurs along the middle line in front so that the mesoscutum anteriorly has a 
strongly rugose appearance but along the middle of the lateral lobes the surface tends to 
be much smoother. Costad abscissa of the basalis slightly shorter; first discoidal cell slightly 
less elongate; metacarp very slightly shorter, about one and two thirds times longer than its 
distance from the apex of the radial cell; median cell showing an even distribution of setae. 
Inner spur of the hind tibia slightly shorter, not more than two thirds as long as the hind 
basitarsus. 

Gaster slightly narrower. Tergite 2 a little longer than 3. Seen in profile, the postero- 
dorsal edge of the hypopygium is deeply sinuate; this is more obvious than in grandis. 

Length: c. 3-5 mm. without ovipositor. 

Type ?. ENGLAND: Kent, Wye Downs, 2.vi.i949 (R. L. E. Ford), B.M. (N.H.). 

Paratypes. Same data: i $, 8.vii.ig46 (G. E. J. Nixon). Kent, Dartford 
Heath, 2 ?, i6.ix.i96o (R. L. E. Ford}; Dulwich, 2 ?, 24.viii.i885 (Billups). 

Mr. Ford has taken a series of nine males with the two females from Dartford 
Heath that I think may well belong to this species; they show the same strong 
rugosity of the anterior part of the mesoscutum a similarly short metacarp and 
similarly coloured legs. The association, however, must be regarded as circum- 
stantial for I am unable to define the male clearly in relation to the males of other 
species. 

In general facies, the female bears a striking resemblance to sticticus, differing 
from it by hardly more than the degree of sclerotization of the hypopygium, the 
arrangement of bristles at the apex of the ovipositor sheath and the longer flagellar 
segments. 



Microgaster brittoni Viereck 

Microgaster (Microgaster) brittoni Viereck, 1916 : 202. 
Microgaster brittoni Viereck; Muesebeck, 1922 : 36. 

Through the courtesy of Dr. Muesebeck, I have been able to examine a male and 
two females of this species, all determined by him. 

?. Apart from differing from acilius in its extensively yellow-marked gaster, brittoni is 
extremely like that species. 

The front part of the mesoscutum is more evenly and more evidently rugose than in acilius. 
First discoidal cell more elongate, as in grandis. Hind femur narrow and very evenly widened 
towards apex. Tergite i appears shorter than in acilius and is more abruptly and strongly 
widened towards apex. Ovipositor sheath distinctly longer. 

cJ. In colour and in sculpture of mesoscutum like the female. Beyond this there is nothing 
useful I can say about this specimen. Dr. Muesebeck assures me that it is an excellent match 
of Viereck's type. 

Material examined: U.S.A., Mass., Holliston, i <j>. Michigan, Tuscola, i <$. 
CANADA: Toronto, i $. 

This species, together with grandis and acilius form a small species-group within 
the larger aggregate, characterized by the long, somewhat tapering flagellum, with 
its very elongate preapical segment, the evenly sclerotized hypopygium and the 



REVISION OF MICROGASTER 55 

differentiated tuft of bristles at the apex of the ovipositor sheath. This tuft is 
not too readily seen in the two females of brittoni but is more distinct in the specimen 
from Holliston. 

Microgaster deceptor sp. n. 

(Text-figs. 10, 19, 20) 
An aberrant species on account of large claws and heavily sculptured thorax. 

$. All femora and all tibiae entirely reddish yellow. Basal half of gaster beneath yellowish; 
gaster otherwise, except for the black first and second segments, dark brown. 

Head from above strongly narrowed behind the eyes (Text-fig. 19). Temples almost coarsely 
rugose; traces of rugose-punctation immediately behind the ocelli. Flagellum thin, the pre- 
apical segment about one and a half times longer than wide. Ocelli in a higher triangle than 
in the other species in this synopsis except auriculatus. 

Costad abscissa of the basalis fully two thirds as long as the mediad abscissa; areolet some- 
what characteristic in shape (Text-fig. 20). Apical tarsal segments of all legs somewhat en- 
larged (Text-fig. 10). Propodeum with a longer, more clearly differentiated, dorsal surface 
than in the other species of the genus, somewhat dull looking, densely intricately rugose; the 
posterior corners rather strongly produced backwards. 

Tergite 2 three times as wide as long and markedly longer than 3; tergite 3 with a group 
of large punctures towards sides. Widened, hairy part of ovipositor sheath a little less than 
half as long as the hind tibia. Hypopygium in type shows no lateral creasing but is tightly 
folded along the middle line; in paratype distinct creasing is visible. 

Length : 4 mm. without ovipositor. 

Type $. FINLAND: Juuga (Woldstedt), Helsinki Museum. 

Paratype. Same data: i $. 

Apart from its long claws, this species has the mesoscutum more heavily sculp- 
tured than in any other species, excepting the russatus-group (but cf. auriculatus) 
On structure of propodeum, enlargement of apical tarsal segment and lengthening 
of tergite 2, this species and auriculatus are transitional towards russatus and 
rugulosus, the two species that formerly constituted the genus Hygroplitis, now 
treated as a synonym of Microgaster (Muesebeck & Walkley, 1951 : 135, Nixon, 
1965 : 267). 

Microgaster auriculatus Fab. 

(Text-fig. 12) 

Ichneumon auriculatus Fab., 1804 : 69. 
Microgaster auriculatus Fab.; Spinola, 1808 : 147. 

A most distinctive species, characterized by the pale-marked gaster, short ovi- 
positor and especially by the long claws. On claw-length different from typical 
species of Microgaster with the exception of deceptor to which it appears to be closely 
related and with which it may be compared as follows: 

?. Hind coxa entirely reddish yellow; (hind tibia missing); Scape reddish yellow on basal 
half; flagellum pale. Tergite i suffused with reddish; 2 still paler; 3 reddish yellow; a wedge- 
shaped mark of the same colour extending along the middle of the following three tergites. 



56 G. E. J. NIXON 

Head from above less exaggerated in appearance, less narrowed behind the eyes (cf. Text- 
fig. 19). Ocelli in a high triangle as in deceptor. Flagellum thin as in deceptor but broken in 
the single specimen available; twelve antennal segments present on left side. 

Mesoscutum less rugose. Front part of mesopleurum without the dense punctation charac- 
teristic of deceptor but nevertheless with coarse rugosity within the prepectal area. Claws 
thinner, slightly longer and more evenly curved; apical tarsal segment of all legs less obviously 
enlarged. Areolet of the fore wing not characteristic in shape. 

The pale honey-yellow hypopygium is more evenly sclerotized than in deceptor and is without 
a trace of lateral creasing. Ovipositor sheath as in deceptor. 

Material examined: GERMANY: i $, labelled " Fiirstenberg i.M. Konow ", " auri- 
culatus F. Coll. Marshall " and " Microgaster auriculatus F. det Szepligeti ". This 
specimen received on loan from the Hungarian Natural History Museum. 

M. auriculatus and M. deceptor clearly represent a small species-group within 
Microgaster, characterized essentially by the long claws. 



Microgaster subcompletus Nees 

Microgaster subcompletus Nees, 1934 : J ^5- 

Microgaster carinata Packard, 1880 : 25, Muesebeck, 1922 : 38, syn. n. 

$. Scape and flagellum pale beneath. Pale parts of the legs somewhat yellowish; front 
and middle femora entirely pale. Basal half of ventral surface of gaster yellowish. 

Mesoscutum without punctation in front but rugose at the origin of the notaulic courses; 
the rugosity sometimes extending almost to middle of disc. Mesopleurum in front strongly 
shining and with only minute setiferous punctures. 

Ovipositor evenly but weakly curved and rather thick. 

N.W. EUROPE. NORTH AMERICA, ? Mass, (carinata). According to Muesebeck, 
in litt. , carinata may not be native to North America. 

Host: Pyramis atalanta Linn., (Nymphalidae), in Europe and North America. 
Notarcha ruralis Scopoli (Pyralidae), Europe in B.M. (N.H.). Both hosts live 
concealed in folded or rolled leaves of nettle (Urtica). 

This, the only gregarious species of Microgaster known to me, is easily recognized 
by its long, thick ovipositor, thin, bristly antenna and black-tipped hind femur 
and hind tibia. 

Dr. Muesebeck has confirmed that carinata falls as a synonym of subcompletus. 



Microgaster laeviscuta Thomson 

(Text-figs. 7, 30) 
Microgaster laeviscuta Thomson, 1895 : 2239. 

This is one of a small group of species previously confused by older writers under 
the name " globatus Nees ". 

$. All the femora and tibiae red; hind tarsus almost as red as its tibia; hind spurs as red 
as their tibia. 



REVISION OF MICROGASTER 



57 



Flagellum somewhat thick, not bristly, usually pale beneath and with the two preapical 
segments obviously longer than wide. The two apical segments of the maxillary palpi rather 
short. 

Mesoscutum considerably roughened in front but without obvious punctation. Front part 
of the mesopleurum shiny, at most finely rugose. Spines of the outer side of the hind tibia 
pale and short, the thicker ones along upper edge of tibia clearly a little shorter than the finer 
spines on the lower side; inner spur of the hind tibia rather short in relation to the outer one 
and not reaching beyond basal two thirds of the hind basitarsus (Text-fig 7). First abscissa 
of the radius very obliquely placed on the stigma (Text-fig. 30). 

Ovipositor thick, forming an even curve; the straight line joining its extremities equal to 
the length of the hind tibia. 

Length : c. 4 mm. without ovipositor. 

CZECHOSLOVAKIA. ENGLAND. FINLAND. SWEDEN. 

Type in the University Museum, Lund, Sweden. Examined. 

Host: Argyroploce dimidiana Sodoffsky (Eucosmidae) (in England); Acrobasis 
tumidana Schiffermiiller (Crambidae) (in Czechoslovakia); Tortrix viridana L. 
(Tortricidae) (in Czechoslovakia). 

Very important for the recognition of this species is the shortness of the inner 
spur of the hind tibia (not an easy feature to appreciate!), and the thick, evenly 
curved ovipositor. 

Microgaster hospes Marshall 
(Text-fig. 17) 

Microgaster hospes Marshall, 1885 : 257. 
Microgaster comptanae Viereck, 1911 : 403, syn. n. 

?. Very close to laeviscuta but smaller and differing from it only in a few details; these are 
mainly concerned with colour. 

All the femora clouded with infuscation at least basally; sometimes the hind femur entirely 
blackish; in palest specimens the middle femur always shows a dark streak above. 

Flagellum rather short, a little shorter than in laeviscuta but the preapical segment at least 
very slightly longer than wide. 

Inner spur of the hind tibia slightly longer in relation to the length of the outer one and 
to the length of the hind basitarsus. First abscissa of the radius very slightly less curved 
than in laeviscuta (cf. Text-fig. 30). 

Hairy part of the ovipositor sheath shorter than in laeviscuta, not more than half as long as 
the hind tibia. 

Length: 3-3-2 mm. without ovipositor. 

Distribution: N.W. EUROPE. NORTH AMERICA (comptanae). 

Type in the B.M.(N.H.). 

Host: Peronea hastiana L., Depressaria aspersana Hubner (Tortricidae). Bred 
from both these hosts in England by R. L. E. Ford. Parasite overwinters in cocoon 
and emerges in May and June. Ancylis comptana Frohlich (Tortricidae) in North 
America, host of comptanae. 

I sent specimens of hospes to Dr. Muesebeck for comparison with Viereck's type of 
comptanae and he has confirmed that they are the same species. 



5 G. E. J. NIXON 

The hind tibia of hospes has the same subtly distinctive appearance as that of 
laeviscuta, all the spines being just as short and pale golden in colour. 

I am not satisfied that I have clearly recognized the specific limits of hospes; 
it is a species needing further study. 

Microgaster peroneae Walley 

Microgaster peroneae Walley, 1935 : 56. 
This is a species with only a minute black spine at the base of the claws. 

$. Flagellum a little pale beneath. Hind femur bright yellowish rather than reddish with 
only a faint touch of infuscation at tip; hind tibia with weak infuscation at extreme apex; 
hind tarsus about as dark as the apex of the hind tibia but the hind basitarsus whitish at base. 

Maxillary palpi somewhat short, in comparison with, say, leechi, bright yellow. Temples 
considerably rugose. Antenna rather thin; flagellum slightly bristly, the preapical segment 
fully one and one third times longer than wide; the segment proximal to this still longer. Face 
evenly dull, confusedly rugulose. 

Anterior part of the mesoscutum considerably roughened, the lines of the notaulices markedly 
rugose as far as about middle. Anterior part of mesopleurum shiny and with large, but very 
ill denned punctures. Inner spur of the hind tibia three fifths as long as the hind basitarsus; 
hind coxa on outer side more evenly and more closely punctate than in hospes. Medial groove 
of mesosternum wide enough to show its costae; this groove is narrower in hospes. 

In one female, a paratype, tergite 3 is as rugose as 2; in the other two females, this tergite, 
though still markedly rugose, is much less so than tergite 2. Hypopygium large, tightly folded 
along the middle line in the dead insect and with numerous lateral creases. Hairy part of the 
ovipositor sheath almost two thirds as long as the hind tibia. 

Material examined: CANADA: N.S., Grand River, i $ paratype, received for 
B.M. (N.H.) through the courtesy of W. M. Mason. Ontario, Biscotasing, i $, ex 
P. variana. British Columbia, Creston, i $, ex Acleris variana on Cranberry. 

Host: Peronea variana Fernald (now Acleris v.) (Tortricidae) . 

There is very little of substance to separate this species from hospes, beyond 
colour and the slightly longer ovipositor of peroneae. It is possible that the hypo- 
pygium of peroneae is larger than that of hospes but the difference is difficult to 
assess with only two females of peroneae available. 

Microgaster ductilis sp. n. 

(Text-fig. 18) 

$. Dark species; hind femur blackish; hind tibia showing a dark reddish infuscation that 
deepens distal to middle; hind tarsus infuscate, as dark as the apical part of the hind tibia; 
hind spurs contrasting pale yellow. Wings strongly infumated. 

Flagellum thin, bristly; preapical segment only very slightly longer than wide. 

Anterior part of the mesopleurum very shiny, with very shallow but more or less discrete 
punctation. Metacarp fully three times as long as its distance from the apex of the radial 
cell; costad abscissa of the basalis fully one third as long as the mediad abscissa. Inner spur 
of the hind tibia three quarters as long as the hind basitarsus. 

Ovipositor sheath with unusually long, thin, basal stalk; hairy part of sheath about two 
thirds as long as the hind tibia; ovipositor slightly but evenly curved throughout. 

Length : 4 mm. without ovipositor. 



REVISION OF MICROGASTER 



59 



Type $. FINLAND: Siikajoki (Wuorentaus), Helsinki Museum. 
Paratypes. FINLAND: Aitlolahti, I $, vii. ; Kyolimaj, i 9; Kangasala, i $, vii. 
This dark-legged species is essentially characterized by the bristly flagellum and 
the long hypopygium. 

Microgaster famulus sp. n. 

9. This species is superficially extremely like tibialis, differing from it as follows: 

In four out of six females, the hind femur is predominantly red with considerable apical 
infuscation. Wings as dark as in tibialis. 

Flagellum slightly longer and slightly bristly. 

Front part of the mesoscutum less clearly punctate. Front part of mesopleurum rather 
coarsely rugose; the same area in tibialis shows well defined, discrete punctures. Metacarp 
longer, about twice as long as its distance from the apex of the radial cell. Claws less bent 
and without spines. 

Ovipositor sheath, seen from above, with many more outstanding bristles. 

Type $. AUSTRIA: Loitsch-Krain, vii-viii (Graeffe Coll.}. Naturhistorisches 
Museum, Vienna. 

Paratypes. AUSTRIA: Steiermark, Tratten b. Murau, 2$, ix.; Feistritz i.d. 
Wochein, i $. 

The face of this species is very shiny and has rather ill denned pits along the 
middle line. This is an observable difference, compared with tibialis, but it may 
have little significance. In spite of the differences given above, the specific validity 
of famulus should be accepted with some caution. 

Microgaster sticticus Ruthe 

(Text-figs. 2, 33) 
Microgaster sticticus Ruthe, 1858 : 5. 

This species closely resembles acilius in general facies but there is a fundamental 
difference between the two species in the sclerotization of the hypopygium; that of 
sticticus, though short, is tightly folded along the middle line and laterally usually 
shows one or more creases in death. 

$. Legs, excluding the coxae, red; rarely the middle femur with a short, dark streak above 
at base. Wings faintly, and virtually evenly, brownish. 

Head rather deeply emarginate behind. Space between the eye-margin and the ocelli trans- 
versely striate. Flagellum not bristly (Text-fig. 2). 

Mesoscutum conspicuously rugose in front, the sculpture coarser along the course of the 
notaulices. Front part of the mesopleurum strongly shining, very superficially punctate. 
Propodeum rather evenly rounded from back to front ; its sculpture fine and close for the genus ; 
medial keel, and lateral keel behind, weak. As in acilius, the radius leaves the stigma markedly 
distal to middle. Inner spur of the hind tibia about three quarters as long as the hind basi- 
tarsus. 

Gaster decidedly elongate (Text-fig. 33). 

Length: c. 3-8 mm. without ovipositor. A medium sized species of slender build. 

N.W. EUROPE. Common. 

This species could be confused with the similarly coloured laeviscuta; the latter, 



60 G. E. J. NIXON 

however, has the head more transverse, the second segment of the gaster more 
transverse, the inner spur of the hind tibia relatively shorter and the ovipositor 
sheaths longer. 

Microgaster alebion sp. n. 

I regard this species, provisionally, as being composed of two generations a 
spring form emerging from larvae of Platyptilia and a summer one parasitizing 
various lepidoptera feeding on nettle (Urtica). These two generations are separable 
on a small difference in the length of the ovipositor but by no other criteria that 
I believe to have specific validity. It is the spring generation from Platyptilia to 
which I give the name " alebion ". 

$. The two basal sternites usually extensively yellow; in such individuals, the femora are 
entirely red ; in one out of two bred females from Boxhill, the middle femur shows faint darken- 
ing at base above; hind tibia without apical infuscation; hind tarsal segments becoming in- 
fuscated towards apex, but generally as reddish as tibia. Fore wing almost hyaline. 

Head above with a considerable amount of rugosity; between the ocelli and the eye-margin 
there is always at least a trace of striation. Distance between a posterior ocellus and the 
front ocellus distinctly less than the diameter of the anterior ocellus. Flagellum not bristly, 
the preapical segment hardly one and one third times longer than wide. 

Mesoscutum very strongly rugose in front, the coarse, wrinkly rugosity extending backwards 
along the course of the notaulices almost to middle; the sculpture much stronger than in the 
very similar laeviscuta. Front part of the mesopleurum quite strongly rugose-punctate. Hind 
tibia slightly less thick than in laeviscuta, the spines of its outer surface having a very prickly 
appearance, being long, sharply pointed and more upstanding than in laeviscuta; inner spur 
of the hind tibia about three quarters as long as the hind basitarsus, much longer in proportion 
to the outer spur than in laeviscuta. Costad abscissa of the basalis rather short, hardly more 
than one third as long as the mediad abscissa. 

Ovipositor sheath about three quarters as long as the hind tibia. 

Length : ca. 4 mm. without ovipositor. 

Type $. ENGLAND: Kent, Gravesend, collected i.vi.igso, emerged i6.vi.i95o, 
ex Platyptilia gonodactyla (R. L. E. Ford], B.M. (N.H.). 

Paratypes (females only). Same data as type, 3 $, 4 $; I ?, i <$, io.vi.i95i. 
England: Kent, Polhill, 3$, I <$, 12. vi. 1952; Surrey, Boxhill, 2$, 3 , v.i95i; 
3 $, 2 <$, i.vi.i959. All the foregoing bred by R. L. E. Ford from P. gonodactyla. 
Surrey, Redhill, I $, 9^.1934, ex P. gonodactyla (W. Rail-Smith]. Herts., Gade 
Valley, I $, 23. vi. 1936 (R. B. Benson). 

Host : Platyptilia gonodactyla Schiffermuller (Pterophoridae) on Tussilago farfara 
L., (Coltsfoot). 

Microgaster alebion, var. A 

$. Differs from the nominate form only in the length of the ovipositor, the sheaths being 
two thirds as long as the hind tibia. 

ENGLAND: Hants, Stockbridge, i $, i ^, vii. ex Vanessa atalanta. Kent, Bexley, 
i $, i 3, collected 6. vii. 1938, emerged i.viii.1938, ex Vanessa atalanta (R. L. E. 
Ford); 8$, 3^, vii. 1943, ex Choreutis myllerana (R.L.E.F.). Surrey, Esher, 2$, 
i <$, collected 12. vi. 1947, emerged 30. vi. 1947, ex Notarcha ruralis (G. E. J. Nixon) ; 



REVISION OF MICROGASTER 61 

Wimbledon Common, i $, 20. .1947, ex N. ruralis; Redhill, I $, I4.viii.i947, ex 
Simaethis fabriciana. Sussex, Hailsham, i $, vii, ex N. ruralis. 

Host: Choreutis myllerana Fabr., Notarcha ruralis Scopoli, Simaethis fabriciana L. 
These three hosts, like Platyptilia gonodactyla, the host of the nominate form, are 
all Pyralidae. Vanessa atalanta L. (Nymphalidae). Since this host, like the others 
of var. A., lives in a rolled or folded leaf of Urtica (Nettle), the possibility of mis- 
identification of host cannot be ruled out here. 

Platyptilia gonodactyla lays its eggs in September on Tussilago so that the early 
stages are available for alebion, var. A. after its cycle on the Urtica-ieeding hosts. 

I have examined a series of seven females from various localities in Finland, that 
resemble var. A. in the shortness of the ovipositor sheaths but show considerable 
leg darkening. In all these specimens both hind femur and hind tibia show well 
defined apical infuscation and in some individuals, the hind femur shows also a 
patch of basal infuscation. 

Microgaster parvistriga Thomson 

Microgaster parvistriga Thomson, 1895 : 2241. 

$. All the femora reddish yellow. Wings almost hyaline. 

Face rather finely rugulose-aciculate, lacking the somewhat coarse punctate element seen in 
the facial sculpture of hospes. Vertex between the ocelli and the eye-margin without trans- 
verse striation; immediately behind and between the ocelli, the surface virtually smooth and 
highly polished. Flagellum rather short, not bristly; the two preapical segments about one 
and a quarter times longer than wide. 

Mesoscutum with a polished, smooth appearance in English specimens, the weak sculpture 
at the origin of the notaulic courses hardly spreading backwards onto the disc; in a large indi- 
vidual from Germany, Berlin (Ruthe Coll. in B.M. (N.H.)), the front part of the mesoscutum shows 
more rugosity than in the English material. Costad abscissa of the basalis not more than 
one third as long as the mediad abscissa, that is, relatively shorter than in hospes. 

Tergites i and 2 rather weakly sculptured compared with hospes and laeviscuta, the surface 
having a smoothed-out appearance. 

ENGLAND. GERMANY. SWEDEN. 

Host: A series in the B.M. (N.H.) was bred in August, 1945, from catkins of 
Betula found in March of the same year. 

The determinative feature for recognizing this species is the shape of the ovipositor. 
When this is concealed within its sheaths, the species could easily be confused with 
palest forms of hospes and smallish examples of laeviscuta. From the latter, parvi- 
striga differs in having the first abscissa of the radius much less obliquely placed 
(cf. Text-fig. 30) and from hospes in having tergites i and 2 more finely sculptured. 

Microgaster caris sp. n. 

(Text-fig. 22) 

$. A brightly coloured species. The pale parts of the legs are more yellowish than reddish; 
the hind spurs are whitish and contrast strongly with the deeply infuscated apex of the hind 
tibia and the equally infuscated hind tarsus. 



62 G. E. J. NIXON 

Maxillary palpus pale yellow. Wings only faintly and evenly darkened. 

Head considerably roughened above; traces of transverse striation between the eye-margin 
and the ocelli and at temples. Flagellum thin, bristly, with the underside markedly paler; 
antennal segment 16 fully one and a half times longer than wide. 

Mesoscutum densely sculptured at least to middle, the sculpture having a somewhat shrivelled 
appearance. 

Tergites i and 2 as densely sculptured as in the majority of the species. Tergite 2 distinctly 
longer than 3; tergite 3 with pale apical margin. Ovipositor sheaths about three fifths as long 
as the hind tibia. 

Length: c. 3-5 mm. without ovipositor. 

Type . AUSTRIA: N. Tyrol, Salvenburg (Kohl), Naturhistorisches Museum, 
Vienna. 

Same data: i $. AUSTRIA: Jitter, i $ (Kohl). 

The single male that I associate with the female has the underside of the flagellum 
almost yellow. The sculpture of the mesoscutum is a little less extensive but the 
venational details and the colour of the legs are the same. 

Microgaster asramenes sp. n. 

$. This species has been largely characterized by the details given in the key; there is little 
to add. 

Wings faintly darkened, the colour deeper apically; a dark band envelopes the first abscissa 
of the radius and this contrasts with a large, pale area occupying almost proximal half of radial 
cell. Scape almost entirely reddish; flagellum brown, paler beneath. Basal half of gaster 
beneath, yellowish. 

Sculpture of face decidedly fine. Flagellum somewhat bristly, tapering apically; the pre- 
apical segment fully one and a half times longer than wide. Distance between a posterior 
ocellus and the anterior ocellus equal to about half the diameter of the anterior ocellus ; posterior 
ocellus separated from eye-margin by about one and two thirds its diameter. Vertex between 
and around the ocelli highly polished. 

Inner spur of the hind tibia fully four fifths as long as the hind basitarsus. Submediellan 
cell of the hind wing entirely free from hairs. 

Tergite 2 as long as 3. Hairy part of the ovipositor sheath fully two thirds as long as the 
hind tibia. 

Length: c. 4-5 mm. without ovipositor. 

<J. Flagellum strongly tapered, the thick, basal segments somewhat flattened; the entire 
flagellum almost yellow beneath. Tergites i and 2 more extensively smooth and polished 
medially than in the female. 

Type ?. TURKEY: Cayeli, 50 ft., 22.viii.i959 (K. M. Guichard),~BM. (N.H.). 
Taken in Alnus plantation near stream. 

Paratypes. ITALY: Napoli, 2 $, I5.viii.i953. 

Microgaster pantographae Muesebeck 

Microgaster pantographae Muesebeck, 1922 : 34. 

$. Front and middle coxae, except at base, pale yellow; trochanters of all legs and front 
and middle tibiae pale yellow; hind femur yellowish, faintly infuscate at extreme apex; the 
blackened hind tarsal segments are faintly annulated with white at base. Scape and flagellum 
brown, both paler beneath. Wings faintly and more less evenly darkened (England) ; virtually 
hyaline in paratype from U.S.A. 



REVISION OF MICROGASTER 63 

Antenna somewhat tapering to apex; with segment 16 fully one and a half times longer than 
wide; flagellum somewhat bristly. Distance between a posterior ocellus and the anterior 
ocellus about half the diameter of the anterior ocellus; posterior ocellus separated from the 
eye-margin by about one and two thirds its own diameter. 

Submedian cell of the hind wing with scattered, colourless hairs. 

Tergite 2 very distinctly longer than 3, very coarsely reticulate-rugose; tergite 3 almost as 
strongly rugose over at least basal half. Hairy part of the ovipositor sheath about two thirds 
as long as the hind tibia; ovipositor itself evenly curved throughout. 

Length: c. 4-5 mm. without ovipositor. 

Distribution: U.S.A., Maine, Bangor (type locality). Various other localities 
recorded by Muesebeck. ENGLAND : Surrey, Horsley, 12 . vi . 1957, i $ (/. F. Perkins], 
in B.M. (N.H.). 

Host: According to Muesebeck, pantographae described from the linden-leaf 
roller, Pantographa lineata Grote and Robinson (Pyralidae). Same author also 
gives Gelechia cercerisella Chambers (Gelechiidae) as another host. 

In the English female the hind tibia is strongly infuscated on about apical third; 
the middle part of the tibia is almost reddish and the basal fifth is yellowish; in 
the American examples (paratype and one other female examined), this contrast in 
colour is much less well marked, the apex of the hind tibia being only weakly 
infuscate. 

This species is readily distinguished from all other N.W. European species by 
its predominantly yellow-marked hind coxa. 



Microgaster phryne sp. n. 

(Text-fig. 32) 

$. Front and middle legs distal to coxae entirely pale reddish yellow; hind tibia with 
darkened tip and hind tarsus infuscate more or less throughout. Wings almost hyaline. 

Face finely rugose and with no obvious punctate element (Text-fig. 32). Temples consider- 
ably roughened. Flagellum thin; preapical segment about one and a third times longer than 
wide. 

Mesoscutum somewhat dull and rugulose almost all over though the sculpture fades out and 
the surface becomes more shiny towards back. Costad abscissa of the basalis from one thrid 
to nearly one half as long as the mediad abscissa; stigma short, somewhat broad. Claws simple. 

Tergite 2 weakly sculptured, sometimes almost smooth. Hypopygium more or less evenly 
sclerotized, without lateral creases. Ovipositor sheaths very short, narrow, about as wide as 
the hind basitarsus ; hairy part of sheath about two fifths as long as the hind tibia. Ovipositor 
rather thin, weakly downcurved at apex. 

cJ. Like the female in colour and sculpture but the eyes slightly less convergent. 

Length: $ $, c. 2-3 mm. without ovipositor of female. 

Type -. EUROPE: Yugoslavia, Macedonia, Prespa Geul, 22. vi. 1958, verge of 
oak-wood (R. L. Coe), B.M. (N.H.). 

Paratypes. Same data : 4 ., 2 <. 

This unusually small species is a typical Microgaster. It differs from all other 
species in having convergent eyes. This character will separate it from opheltes, 
to which it otherwise bears a very close resemblance. 



64 G. E. J. NIXON 

Microgaster fischeri Papp 
Microgaster fischeri Papp, 1960 : 120 

My knowledge of this little species is based on two male paratypes kindly lent 
to me by Dr. Max Fischer of the Naturhistorisches Museum, Vienna. 

The species was described from the neighbourhood of Vienna and, being based 
unfortunately on the male sex only, is difficult to interpret. Nevertheless, there 
are, in the B.M. (N.H.), two females and one male, from England, all of which 
agree with the paratypes in size, colour and sculpture. It is on the characters of 
the two females that I am interpreting the species and finding a place for it in the 
synopsis. 

<J $. Wings considerably, evenly, darkened. Middle and hind femur entirely blackish in 
the three English examples; in one of the paratypes, the middle femur is flushed with reddish 
along outer side in apical half; hind tibia brownish red, gradually becoming darker in about 
apical third. 

$. Face finely rugose, without a punctate element, much as in phryne. Flagellum some- 
what thin, with the preapical segment about one and a third times longer than wide, exactly 
as in phryne. Eyes not at all convergent. 

Mesoscutum considerably rugulose in front half in the two paratypes; less so in the English 
examples; the sculpture virtually not distinguishable from that of phryne. Propodeum with 
hardly an indication of a medial keel; it is sharply defined in phryne. 

In the paratypes, tergite 3 is sculptured over basal half; in the English examples, this segment 
is smooth, though not as polished as in phyrne. Ovipositor sheath as short as in phryne but 
wider; ovipositor slightly thicker, evenly curved throughout. 

Length: <J 9. c - 2 '3 mm. without ovipositor of female. 

AUSTRIA: Vienna (type locality). ENGLAND: Bucks., Princes Risborough, i $, 
2i.vi-4.vii.i942; Aston Clinton, i <, 6.vi.i953; Wendover, i $, 28. vi. 1955. All 
R. B. Benson. 

Small size and very short ovipositor are important features of this species. 



Microgaster canadensis Muesebeck 

Microgaster canadensis Muesebeck, 1922 : 38. 

$. Anterior part of mesoscutum without punctation but markedly rugose at the origin of 
the notaulic courses. Preapical segment of the flagellum almost square in outline ($ from 
Canada) ; flagellum not in the least bristly. 

<J. Flagellum very long, heavy-looking; the basal segments decidedly flattened; the whole 
flagellum markedly pale beneath. Hind claw pectinate like that of female. 

Material examined: CANADA (type locality). Ontario, i $, i <$, ex Compsolechia 
niveopulvdla; these two specimens in B.M. (N.H.) as well as female paratype. 

Host: Compsolechia niveopulvella Chambers (Gelechiidae) . No host known for 
the type series. 

Rather close to crassicornis from which it differs in not having bristly antennae 
and differently armed claws. 



REVISION OF MICROGASTER 65 

Microgaster reticulatus Shestakov 
Microgaster reticulatus Shestakov, 1940 : n. 

I have seen a female of this species from the Riksmuseum, Stockholm, labelled 
" Microgaster reticulatus, sp. n., det Shestakov, Vladivostok, Suchan (Malaise) " . 

$. Flagellum very long, much tapered towards apex, bristly. Costad abscissa of the 
basalis short, only about one quarter as long as the mediad abscissa. Outer side of the hind 
tibia unusually densely beset with short, thick, fire-red spines. Propodeal rugosity coarser 
than in other species known to me; the propodeum itself, especially towards sides, is clothed 
with long, silvery white hairs. 

Gaster somewhat short, tergite (2 + 3) being longer than following tergites together; the suture 
between tergites 2 and 3 is much deeper, broader and more distinctly costate than in any other 
species in this revision. 

Length : $, 6 mm. without ovipositor. 

E. SIBERIA: Vladivostok. 

This is a most distinctive species, aberrant among those species with simple claws 
and probably representing a distinct species-group. 

Microgaster leechi Walley 
Microgaster leechi Walley, 1935 : 57. 

$. This species is richly marked with yellow. Scape predominantly pale; flagellum pale 
almost throughout but the 2-3 basal segments darkened above. Hind coxa yellow throughout ; 
hind femur reddish yellow; hind tibia almost dull reddish, with conspicuous apical infuscation; 
hind tarsus deeply infuscate throughout. The rather long maxillary palpi pale yellow. 

Eyes, in a facial view, markedly convergent and the face with well marked keel on upper 
half; in one of the two specimens available (Ottawa), this keel is distinct as far as the clypeus. 
Flagellum somewhat thin, markedly bristly; penultimate segment one and a half times, and 
antipenultimate segment almost twice, as long as wide. 

Costad abscissa of the basalis fully two fifths as long as the mediad abscissa. Claws simple 
(cf. gelechiae) ; outer side of hind tibia having a prickly appearance, its spines numerous and 
rather long. 

Gaster beyond tergite 2 almost entirely reddish yellow (Clement) or with broad, medial, 
blackish band (Ottawa). Ovipositor sheath fully two thirds as long as the hind tibia; ovipositor 
evenly curved throughout. 

o*. The flagellum is strikingly yellowish throughout, except that the two to three basal 
segments are darkened above; this contrast is more striking than in the female. Apex of hind 
femur faintly darkened above. 

Material examined: CANADA: Quebec, Clement, i $, det. Mason. Ontario, 
Ottawa, i <J, i $, vii and viii.1947 (W. R. M. Mason). U.S.A.: Maine, Dryden, 
3 cJ, 26. vii. 1959 (G. H. Heinrich}; all in B.M. (N.H.). 

Microgaster gelechiae Riley 
Microgaster gelechiae Riley, 1869 : 178. 

Superficially very like leechi with which it may be compared as follows: 

?. Entire hind leg distal to the coxa reddish yellow, the hind tarsus being as brightly coloured 
as its tibia; the hind coxa becomes darkened on about basal half. 



66 G. E. J. NIXON 

Face broader than in leechi, the eyes less convergent and the sculpture more evenly rugose, 
finer. Hairs of the eyes slightly shorter. Flagellum not in the least bristly; the two preapical 
segments about one and one third times longer than wide. Hind claw with three pale spines. 
Ovipositor sheath almost three quarters as long as the hind tibia. Mesosternum with fine, 
transverse striation. 

cj. Flagellum blackish above, faintly paler beneath. 

Material examined: U.S.A., Virginia, Great Falls, i $, i <, det. Gahan, ex Gnori- 
moschema gallae-solidaginis , in B.M. (N.H.). 

Host: Gnorimoschema gallae-solidaginis Riley. 

There is in the B.M. (N.H.) a single female from CANADA (Toronto, Ottawa, 
22. v. 1956, R. B. Benson) that I believe to be this species. The yellow markings 
of the dorsal surface of the gaster are reduced to a yellow spot at each lateral corner 
of tergite 3; these spots are united by the faintly yellow, apical margin of the tergite. 

Microgaster kuchingensis Wilkinson 
Microgaster kuchingensis Wilkinson, 1927 : 176, 1929 : 120. 

A species largely characterized by the heavily blackened hind tibia with its white 
basal ring. 

$. Front and middle tarsi whitish yellow. 

Submedian cell of the hind wing without hairs. Hairy part of ovipositor sheath about half 
as long as the hind tibia; ovipositor thick, evenly downcurved. Hypopygiom short, evenly 
sclerotized and without lateral creases. 

BORNEO: Kuching (type locality). PHILIPPINES: Mt. Macolod, i $, 24.x. 1953 
(H. M. & D. Townes); Mindanao, Surigao, i $ (Baker). 

Type in the B.M. (N.H.). 

I am puzzled by a single female from INDIA: United Provinces, Dehra Dun, ex 
Pyrausia codesalis Walker, defoliating bamboo, (Wilkinson, 1929 : 120). The 
darkened areas of the fore wing are less contrasted than in typical material, the 
hind tibia is not so deeply infuscated and the pale basal ring is less sharply discrete 
and covers almost basal quarter of tibia. The striate element in the striate-punctate 
sculpture of the basal tergites of the far eastern material is absent in this single 
specimen and most of the surface of tergites i and 2 is describable as predominantly 
punctate and shiny. 

Microgaster magnificus Wilkinson 

(Text-fig. 15) 
Microgaster magnified Wilkinson, 1929 : 120. 

$. With its entirely fulvous red thorax, darkened gaster and dark, smoky yellow wings, 
this is a very striking species. The curiously prolonged apex of the hypopygium is distinctive 
and unique among the species dealt with in this paper. 

Hind tibia deeply inf uscate but paler at extreme base ; hind coxa inf uscate but with a yellow- 
ish streak above and below. 



REVISION OF MICROGASTER 67 

Costad abscissa of the basalis fully one third as long as the mediad abscissa. Outer side 
of the hind tibia very densely spinose (cf. tjibodas). 

Tergite 2 almost as smooth and as shining as 3, but with some vague punctation along anterior 
margin. Hairy part of the ovipositor sheath about one third as long as the hind tibia; ovi- 
positor thin, straight basally but sharply downcurved at apex (Text-fig. 15). 

cj. There are two paratypes in the B.M. (N.H.). One has the gaster considerably reddened 
medially. Both have the antenna brown but considerably paler beneath. 

AUSTRALIA: Queensland, 2 $, (type and paratype); 2 <$, (paratypes). 

Type in the B.M. (N.H.). 

In general heaviness of build this species is closer to kuchingensis than to tjibodas 
but for all that the two species are widely different. Whereas kuchingensis is 
striking on colour only and structurally is close to the European species with rugose 
face and simple claws, magnificus is aberrant on the shape of the hypopygium. 
This, of course, may turn out to be only a species-group character. 

Microgaster tjibodas Wilkinson 
Microgaster tjibodas Wilkinson, 1927 : 177. 

$. A brightly coloured species with the legs almost entirely yellow. The thin hind tibia 
is reddish yellow with faint apical darkening; the hind tarsus is weakly infuscated and is much 
the same colour as the darkened apex of the hind tibia. 

Sculpture of the face very fine, feebly rugose and very shiny. Flagellum thin, bristly, with 
the preapical segment fully one and a half times longer than wide. 

First abscissa of the discoideus a little shorter than the second ; costad abscissa of the basalis 
slightly more than one quarter as long as the mediad abscissa. Spines of the outer side of the 
hind tibia very fine and very sparse. 

Horizontal part of tergite i smooth-looking, very shiny and with scattered irregular pits 
and punctures. Tergite 2 almost smooth. Hairy part of the ovipositor sheath about three 
fifths as long as the hind tibia. Hypopygium thin, membranous and with several lateral 
creases. 

JAVA: Tjibodas (type locality). 

Type in the B.M. (N.H.). 

I know this species only from the type. 

Microgaster russatus Haliday 

Microgaster russatus Haliday, 1834 : 237. 

9- Flagellum pale at least beneath. Stigma usually markedly bicoloured but in one speci- 
men in the B.M. (N.H.) entirely yellow. The hind tarsus varies from black to dull reddish 
with the apical segment the darkest. 

Japanese specimens ($) have the pale parts of the body and legs yellowish rather than red 
and the apical infuscation of the hind tibia more extensive. 

EUROPE. JAPAN. 

Host: Orthotelia sparganella Thunberg (Plutellidae) (in the literature). Chilo 
suppressalis Walker; Chilo simplex Butler. Both in Japan. Specimens in B.M. 
(N.H.) C r ambus paludellus Hiibnagel (Crambidae) (in England, bred by R. L. E.Ford). 

I do not think there has ever been any confusion about the identity of this very 
distinctive species. 



68 G. E. J. NIXON 

Microgaster rugulosus Nees 

(Text-fig. 31) 
Microgaster rugulosus Nees, 1834 : 163. 

This species departs from the typical Microgaster pattern in that the apical, polished band 
of the scutellum becomes very narrow at middle owing to the considerable amount of punctation 
at the apex of the scutellar triangle. 

The general surface of the gaster is much duller and more sculptured than russatus and 
tergite (2 -f 3) occupies a much larger area beyond the first tergite (Text-fig. 31). 

Hind coxa densely punctate all over. Hind tibia without apical infuscation; its inner spur 
unusually short, not quite reaching the middle of the hind basitarsus. 

EUROPE. 

Microgaster melligaster Provancher 

Microgaster melligaster Provancher, 1886 : 143. 
Microgaster rubricoxus Provancher, 1888 : 386. 

This species is much more closely related to russatus than to rugulosus and in fact is very 
like the former species from which it differs as follows: 

?. Two basal segments of gaster entirely dark; remaining segments either entirely darkened 
(i $, in B.M., Canada, Ontario) or reddened along sides (2 $, U.S.A., Maine). 

Disc of scutellum more narrowed behind, even slightly constricted. Segments of the hind 
tarsus considerably shorter. Areolet in six out of seven specimens triangular; in russatus, it 
is distinctly four-sided. 

Two basal segments of gaster less transverse; tergite 3 (i.e. tergite (2 + 3) beyond the suture) 
smooth, unsculptured ; in russatus, this segment shows considerable coarse, rugose punctation. 

U.S.A.: Maine, 2 ?, 2 . CANADA: Ontario, 2 <$, i $. All in the B.M. (N.H.). 
Other characters for this species are given in the key. 

Microgaster nerione sp. n. 

(Text-fig. 16) 

?. Middle femur infuscate throughout; front femur somewhat darkened. Fore wing hyaline 
proximal to the areolet and also within the radial cell; otherwise faintly darkened. Gaster 
blackish but becoming dark brown beneath. 

Face strongly shining, its sculpture very weak, obsolescent. Ocelli in a low triangle, the 
posterior tangent to the anterior ocellus cutting the posterior pair. Flagellum long, rather 
thin, slightly bristly; antennal segment sixteen fully twice as long as wide (two apical segments 
missing) . 

Mesoscutum very smooth-looking, virtually without a trace of punctation in front and with 
only very weak rugosity at the origin of the notaulic courses. First abscissa of the discoideus 
distinctly shorter than the second; costad abscissa of the basalis hardly one third as long as 
the mediad abscissa so that the first discoidal cell is somewhat high. Inner spur of the hind 
tibia unusually short, only just reaching beyond middle of hind basitarsus; spines of the outer 
side of the hind tibia somewhat sparse; those along upper edge golden; those below, whitish; 
hind claw with three, close, blackish spines. Anterior part of the mesopleurum strongly 
shining, virtually impunctate. 

Tergite i rather strongly narrowed basally and hence markedly triangular. Tergite 2 about 
three times as wide as long, showing the usual type of rugosity but this tending to fade out 



REVISION OF MICROGASTER 69 

medially; tergite 3 as long as 2. Ovipositor thick, strongly curved. Posterior edge of hypo- 
pygium markedly sinuate (Text-fig. 16). 
Length: 4-3 mm., without ovipositor. 

Type $. MEXICO: Guerrero, Omilteme, 8,000 ft., viii (H. H. Smith], B.M. (N.H.). 

A rather slender species, distinguished from all others with toothed claws by the 
evenly sclerotized hypopygium. This feature, however, does not relate the species, 
in my opinion, to the European grandis-group. 

SPECIES INQUIRENDAE 

1. Microgaster campestris Tobias, 1964 : 210. 

KASAKHSTAN. Tobias states that this species is close to curvicrus Thomson. 

2. Microgasier claritibia Papp, 1959 : 405. 

HUNGARY. Almost certainly a species of Protomicroplitis. 

3. Microgaster fusca Papp, 1959 : 407. 

HUNGARY. Papp states that this species is related to Microgaster postica 
Nees. Probably a species of Protomicroplitis. 

4. Microgaster procris Fischer, 1964 : 42. 

AUSTRIA. Bred from Procris notata Zeller (Zygaenidae) . Probably a species 
of Protomicroplitis. 

5. Microgaster rugosicoxa Papp, 1959 : 408. 

HUNGARY. Papp states that this species is related to Microgaster scotica 
Marshall (now Protomicroplitis scotica (Marshall) Nixon, 1965 : 252. Evi- 
dently a species of Protomicroplitis. 

6. Microgaster tegularius Papp, 1959 : 407. 

HUNGARY. Papp says this species comes close to Microgaster circumvectus 
Lyle (now Protomicroplitis circumvectus (Lyle) Nixon, 1965 : 256). Evidently 
a species of Protomicroplitis. 

7. Microgaster dudichi Papp, 1961 : 154. 

GERMANY. This species certainly belongs to Microgaster s. str. Papp places 
it near tibialis Nees but this does not necessarily relate it to the species for 
which I use the name " tibialis " in this paper. 

8. Protomicroplitis kasachstanica (Tobias) comb. n. 

Hygroplitis kasachstanica Tobias, 1964 : 208. KASAKHSTAN. One female 
paratype in the B.M. (N.H.), presented by Dr. Tobias. Tobias states that 
this species is related to Hygroplitis abdominalis Nees (now Protomicroplitis 
abdominalis (Nees) Nixon, 1965 : 254) ; it is certainly closely related to this 
species and may not really be distinct from it. Nevertheless, without examin- 
ing further specimens I hesitate to sink Tobias' species. 

9. Protomicroplitis stepposa (Tobias) comb. n. 

Hygroplitis stepposa Tobias, 1964 : 209. 

This species seems to be closely related to Protomicroplitis meges Nixon, 
1965 : 251, a species transitional between the abdominalis- and sco^'ca-groups 



yo G. E. J. NIXON 

of Protomicroplitis (Nixon, 1965) ; stepposa is a much more brightly coloured 
species than meges, with the hind tibia entirely reddish yellow and the meso- 
scutum with larger punctures and their interspaces much more polished. 

REFERENCES 

FABRICIUS, J. C. 1798. Entomologia systematica emendata et aucta. Supplementum. pp. 

572. Hafniae. 

1804. Sy sterna Piezatorum. Brunsvigae. 

FAHRINGER, J. 1925-37. Opuscula braconologica. 4. Wien. 

FISCHER, MAX. 1964. Zwei neue geziichtete Braconiden. Entomophaga 9 : 39-44. 

GAHAN, A. B. 1917. Descriptions of some new parasitic Hymenoptera. Proc. U.S. natn. 

Mus. 53 : 195-217. 
HALIDAY, A. H. 1834. Essay on the classification of parasitic Hymenoptera. Ent. Mag. 

2 : 225-259. 

HELLEN, W. 1954. Ubersicht iiber die Microgasterinen Finnlands. Notul.ent. 34 : 106-121. 
MARSHALL, T. A. 1885. Monograph of British Braconidae. I. Trans, ent. Soc. Land., 

1885 : 1-280. 

MORLEY, C. 1936. Notes on Braconidae: XV. Microgasterinae. Entomologist 69 : 209-213. 
MUESEBECK, C. F. W. 1922. A revision of the North American Ichneumon-flies belonging to 

the subfamilies Neoneurinae and Microgasterinae. Proc. U.S. natn. Mus. 61 : 1-76. 
NEES VON ESENBECK, C. G. 1834. Hymenopterorum Ichneumonibus affinium monographiae, 

genera Europaea et species illustrantes. I. 320 pp. Stuttgart et Tubingen. 
NIXON, G. E. J. 1965. A ^classification of the tribe Microgasterini (Hymenoptera, Braconi- 
dae). Bull. Br. Mus. not. Hist. (Ent.). Suppl. 2 : 1-284. 
PACKARD, G. M. 1881 (1880). Descriptions of some New Ichneumon parasites of North 

American butterflies. Proc. Boston Soc. nat. Hist. 21 : 18-38. 
PAPP, J. 1959. The Microgaster Latr., Microplitis Forst., and Hygroplitis Thorns, species of 

the Carpathian Basin (Hymenoptera, Braconidae). Annls hist. natn. Mus. hung. 51 : 

397-4I3- 

1960. Zur Kenntnis der Microgaster Latr., -und Microplitis Forst. Arten Osterreichs 
(Hym., Braconidae). Z. ArbGem. ost. Ent. 12 (3) : 117-214. 

1961. Untersuchungen liber drei Microgaster-Arten. Beitr. Ent. 2 : 154-159. 
PROVANCHER, L. 1885-1889. Faune Entomologique du Canada traitant des Hymenopteres. 

2. Additions et Corrections. 1475. Quebec. 
RILEY, C. V. 1869. First Annual Report on the noxious, beneficial and other insects of the state 

of Missouri, Jefferson City. 180 pp. 

RUTHE, J. F. 1860. Deutsche Braconiden. Berl. ent. Z. 4 : 105-160. 
SHESTAKOV, A. 1940. Zur Kenntnis der Braconiden Ostsibiriens. Ark. Zool. 32 : 1-21. 
SPINOLA, M. 1806-08. Insectorum Liguriae species novae aut rariores. Genuae. 
TELENGA, N. A. 1955. Braconidae: Microgasterinae, Agathinae. Fauna S.S.S.R. 5 (4), 

311 pp. Moscow and Leningrad. 

THOMSON, C. G. 1869-97. Opuscula entomologica 2452 pp. Lund. 
TOBIAS, V. I. 1964. New species and a new genus of Braconidae from Kazakhstan. Trudy 

zool. Inst., Leningr. 34 : 177-234. 
WALLEY, G. STUART. 1935- Five new species of Braconidae with host records of additional 

species. Can. Ent. 67 : 55-61. 

WILKINSON, D. S. 1927. On the Indo-Malayan species of the genus Microgaster (Hymenop- 
tera, Braconidae). Bull. ent. Res. 18 : 171-178. 
1929- A revision of the Indo-Australian and Ethiopian species of the genus Microgaster. 

Trans. R. ent. Soc. Lond. 77 : 99-123. 
MUESEBECK, C. F. W., KROMBEIN, K. V. & TOWNES, H. K. 1951. Hymenoptera of America, 

North of Mexico. Agriculture Monogr. 2 : 1420 pp. 



REVISION OF MICROGASTER 



INDEX 



Numbers in bold type are the most important page references. 



abdominalis-group (of Protomicroplitis), 69 

abdominalis, Hygroplitis, 69 

abdominalis, Microgaster, 33 

acilius, Microgaster, 39, 53, 54 

alebion, Microgaster, 42, 60 

Alnus, 62 

Aphelia, 53 

areolaris, Microgaster, 37, 49 

Argyroploce, 57 

asperdana, Depressaria, 57 

asramenes, Microgaster, 40, 62 

atalanta, Vanessa, 56, 60 

auriculatus , Ichneumon, 55 

auriculatus, Microgaster, 36, 55 

australis, Microgaster 51 

bamboo, 66 

Betula, 6 1 

brittoni, Microgaster, 39, 54 

broom, 46 

campestris, Microgaster, 69 
canadensis, Microgaster, 38, 64 
carinata, Microgaster, 56 
caris, Microgaster, 39, 61 
castanea, Swammerdamia, 53 
cercerisella, Gelechia, 63 
Chilo, 67 
Choreutis, 60 

chrysanthemana, Cnephasia, 53 
circumvectus, Microgaster, 69 
circumvectus, Protomicroplitis, 69 
claritibia, Microgaster, 69 
Cnephasia, 53 
codesalis, Pyrausta, 66 
Coltsfoot, 60 
Compsolechia, 64 
comptana, Ancylis, 57 
comptanae, Microgaster, 57 
congregatiformis, Microgaster, 37 
censors, Microgaster, 34, 38, 50 
Crambidae, 57, 67 
Crambus, 67 

crassicornis, Microgaster, 37, 49, 64 
curvicrus, Microgaster 36, 46, 48, 69 
Cytisus, 46 



denotata, Eupithecia, 49 
Depressaria, 45, 46, 57 
deprimator, Ichneumon, 51 
deprimator, Microgaster, 33, 38, 51 
dimidiana, Argyroploce, 57 
ductilis, Microgaster, 40, 58 
dudichi, Microgaster, 69 

Epagoge, 52 

epagoges, Microgaster, 38, 52 

erro, Microgaster, 36, 46 

Eucosmidae, 57 

Eupithecia, 49 

eupolis, Microgaster, 38, 51 

fabriciana, Simaethis, 61 
famulus, Microgaster, 40, 59 
farfara, Tussilago, 60 
ferrugalis, Phlyctaenia, 52 
fischeri, Microgaster, 40, 64 
fulvicrus, Microgaster, 36, 45, 46 
fusca, Microgaster, 69 

gallae-solidaginis, Gnorimoschema, 66 

Gelechia, 63 

gelechiae, Microgaster, 37, 65 

Gelechiidae, 52, 63, 64 

Geometridae, 49 

globata, Microgaster, 34 

globatus, Microgaster, 56 

Gnorimoschema, 66 

gonodactyla, Platyptilia, 60 

grandis, Microgaster, 39, 53, 54 

hastiana, Peronea, 57 
hospes, Microgaster, 40, 57, 58, 61 
Hygroplitis, 69 
Hyponomeutidae, 53 

kasachstanica, Hygroplitis, 69 
kasachstanica, Protomicroplitis, 69 
kuchingensis, Microgaster, 34, 38, 66 
laeviscuta, Microgaster, 39, 56, 61 

leechi, Microgaster, 36, 65 
lineata, Pantographa, 63 



deceptor, Microgaster, 39, 55 
deductor, Microgaster, 34, 37, 49 



magnificus, Microgaster, 35, 66 
magnifica, Microgaster, 34, 66 



INDEX 



marginatus, Microgaster, 33 
melligaster, Microgaster, 35, 68 
myllerana, Choreutis, 60 

nerione, Microgaster, 34, 37, 68 

nettle, 56, 61 

niveopulvella, Compsolechia, 64 

Notarcha, 60 

notata, Procris, 69 

novicius, Microgaster, 38, 53 

Nymphalidae, 56 

obsepiens, Microgaster, 36, 47 
ocellana, Depressaria, 45 
Oecophoridae, 45 
opheltes, Microgaster, 36, 48 
Orthotelia, 67 

pallorella, Depressaria, 46 
paludellus, Crambus, 67 
Pantographa, 63 

pantographae, Microgaster, 37, 62 
parvistriga, Microgaster, 41, 61 
peroneae, Microgaster, 40, 58 
phryne, Microgaster, 38, 63, 64 
phthorimaeae, Microgaster, 52 
Platyptilia, 60 
Plutellidae, 67 
pluto, Microgaster, 50 
pimpinellata, Eupithecia, 49 
politus, Microgaster, 35, 48 
postica, Microgaster, 69 
procerus, Microgaster, 35, 45, 48 
Procris, 69 

procris, Microgaster, 69 
Protomicroplitis, 33, 69 
Pterophoridae, 60 
Pyralidae, 52, 56, 63 



Pyramis, 56 
Pyrausta, 66 

reticulatus, Microgaster, 36, 65 
rubricoxus, Microgaster, 68 
rugosicoxa, Microgaster, 69 
rugulosus, Microgaster, 35, 68 
ruralis, Notarcha, 56, 60 
russatus, Microgaster, 35, 67 

scotica-group (of Protomicroplitis), 69 
scotica, Microgaster, 33, 69 
scotica, Protomicroplitis, 69 
Simaethis, 61 
simplex, Chilo, 67 
sparganella, Orthotelia, 67 
stepposa, Hygroplitis, 69 
stepposa, Protomicroplitis, 69 
sticticus, Microgaster, 40, 54, 59 
subcompletus, Microgaster, 39, 56 
sulfureana, Epagoge, 52 
suppressalis, Chilo, 67 
swammerdamiae , Microgaster, 53 

tegularius, Microgaster, 69 

tibialis, Microgaster, 38, 46, 50, 59, 69 

tjibodas, Microgaster, 34, 35, 37, 67 

Tortricidae, 52, 53, 57 

tumidana, Acrobasis, 57 

Tussilago, 60, 6 1 

Urtica, 56, 61 

Vanessa, 60 
variana, Acleris, 58 
variana, Peronea, 58 
viburniana, Aphelia, 53 
viridana, Tortrix, 57 

Zygaenidae, 69 




A LIST OF SUPPLEMENTS 
TO THE ENTOMOLOGICAL SERIES 

OF THE BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 



1. MASNER, L. The types of Proctotrupoidea (Hymenoptera) in the British 
Museum (Natural History) and in the Hope Department of Entomology, Oxford. 
Pp. 143. February, 1965. 5. 

2. NIXON, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera : 
Braconidae). Pp.284; 348 Text-figures. August, 1965. 6. 

3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177 ; 
18 plates, 270 Text-figures. August, 1965. 4 45. 

4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172 ; 500 Text-figures. October, 

1965- 35s. 

5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156; 
475 Text-figures. November, 1965. 2 155. 

6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae & Drosophilidae. 
Pp. 129 ; 328 Text-figures. 3. 

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

33s. 

8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the 

world species of Cleora (Lepidoptera : Geometridae). Pp. 119; 14 plates, 146 
Text-figures, 9 maps. February, 1967. 3 los. 

9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera : Rhopalocera). Pp, 509. August 1967. 8 los. 

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rhopa- 
locera). Pp. 322; 233 Text-figures, coloured frontispiece. September 1967. 

11. MOUND, L. A. A review of R. S. Bagnall's Thysanoptera Collection. Pp. 181; 
82 Text-figures. May, 1968. 4. 



PRINTED IN GREAT BRITAIN BY ADLARD & SON LIM ITED, EARTH OLOMEW PRESS, DORKING 



i" V ' 



AN INDEX-CATALOGUE OF 

AFRICAN PHANEROPTERINAE 

(ORTHOPTERA : TETTIGONIIDAE) 



D. R. RAGGE 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 22 No. 3 

LONDON: 1968 



AN INDEX-CATALOGUE OF AFRICAN 

PHANEROPTERINAE 
(ORTHOPTERA : TETTIGONIIDAE) 



BY 

D. R. RAGGE 

British Museum (Natural History 




Pp. 73-108 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 22 No. 3 

LONDON : 1968 



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

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

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

This paper is Vol. 2,2, No. 3 of the Entomological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 



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



Trustees of the British Museum (Natural History) 1968 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 28 M ay, 1968 Price Sixteen Shillings 



AN INDEX-CATALOGUE OF AFRICAN 

PHANEROPTERINAE 
(ORTHOPTERA : TETTIGONIIDAE) 

By D. R. RAGGE 

CONTENTS 

Page 

SYNOPSIS ............ 75 

INTRODUCTION ........... 75 

TREATMENT ........... 76 

THE STATUS OF THE PHANEROPTERINAE ...... 78 

CATALOGUE OF AFRICAN PHANEROPTERINAE ...... 79 

REFERENCES ........... 98 

INDEX ............ 103 

SYNOPSIS 

A catalogue is given of all the genera and species of Phaneropterinae known at present to occur 
in Africa and its off-shore islands (but not those of the Mediterranean Sea or the Malagasy Sub- 
region) , and in the Arabian peninsula. The species are arranged alphabetically under the genera to 
which they have been assigned in the most recent literature, and the arrangement of the genera 
is also alphabetical. Generic and specific synonyms, junior homonyms and deliberate variations 
in spelling are listed chronologically under the valid name. The type-species is cited for all 
generic names, and full information on the type-material is given for all specific names. The 
catalogue includes 115 genus-group and 358 species-group names; of these, eighty-three generic, 
295 specific and seven subspecific names are currently regarded as valid. The status of the 
Phaneropterinae is discussed. 

INTRODUCTION 

THIS catalogue has been compiled as the first stage of a comprehensive revision of the 
African Phaneropterinae. It includes all the species of Phaneropterinae known at 
present to occur in Africa, listed alphabetically under the genera to which they have 
been assigned in the most recent literature. As a deliberate policy, no new combina- 
tions (with the exception of an unavoidable one mentioned below) or synonyms are 
established in this paper; any necessary nomenclatorial changes will appear in the 
systematic revision to be published later. With the exception of groups covered by 
recent revisions (Ragge, 1955, 19560, 19566, 1957, 19600, 19606, 19610, 19616, 19620, 
19626, 1964) the nomenclature will thus be subject to some alteration during the next 
few years. However, the standard of taxonomic work on the Tettigoniidae during 
most of the past century has been remarkably high and there are very few nomen- 
clatorial problems. 

The geographical region covered by this catalogue includes the whole of the African 
mainland and off-shore islands (e.g. Socotra, Zanzibar and the islands of the Gulf of 
Guinea) and the Arabian peninsula, but excludes the Malagasy Sub-region; there is 

ENTOM. 22, 3. 3 



76 D. R. RAGGE 

such a high degree of endemism in the Phaneropterinae of this sub-region that it is 
more appropriate to treat them separately. The Mediterranean islands are not 
included, and the inclusion of North Africa adds only thirteen species, all belonging 
to the genus Odontura Rambur. 

TREATMENT 

The eventual systematic arrangement of the African genera of Phaneropterinae 
will naturally depend on the comprehensive revision to be undertaken during the 
next few years, and it has therefore been thought better at this stage to list the genera 
alphabetically. Generic synonyms, junior homonyms and deliberate variations in 
spelling are listed chronologically under the valid generic name. The entries for 
each genus and generic synonym include a reference to the original description and 
the citation of the type-species and its mode of fixation ; where the type-species was 
fixed by subsequent designation, a bibliographical reference is given to the fixation. 
The type-species is cited in its original combination; where it is now a junior synonym 
the present valid name is given in brackets. Where a type-species does not occur in 
Africa, the country of the type-locality is given in brackets. All generic synonyms 
are cited, including those based on non- African type-species; the type-species are 
cited in the same way as those of valid generic names. 

The African species of each genus are listed alphabetically, together with references 
to their original descriptions and a statement of the nature and data of the type- 
material and the institution in which the type-material is housed. The absence of 
any information on the number and sex of the specimens in the type-series indicates 
that this information was not given with the original description and has not been 
determined subsequently. Where a species is based on a type-series from more than 
one locality, the citation of one depository after the last locality indicates that all the 
type-material is believed to be in that depository. It should be noted that the stated 
numbers of syntypes are not always known for certain, but always with at least a high 
degree of probability. A number of species described by Sjostedt (1909) from 
Kilimanjaro and Meru were based on long series of syntypes, a proportion of which 
were sent out from Stockholm, where Sjostedt worked, to other museums; as the 
exact extent of this distribution of syntypes is unknown, the only depository cited 
in these cases is the Naturhistoriska Riksmuseum, Stockholm. 

Where a species has been transferred from its original genus, the name of the latter 
is given in brackets after the reference to the original description ; a change in status 
is noted in a similar way. A question-mark before the original genus indicates that 
the original generic assignment was doubtful; one before the specific name itself 
indicates that the present generic assignment is doubtful. Where a species is 
currently regarded as polytypic, an entry is given for each subspecies and there is no 
separate entry for the species as a whole. 

Specific synonyms (including those based on non-African type-specimens) and 
deliberate variations in spelling are indented and listed chronologically under each 
species or subspecies; they are otherwise given the same treatment as valid specific 
names. A question-mark before a specific synonym indicates that the synonymy is 
doubtful. 



AFRICAN PHANEROPTERINAE 77 

Territories in which type-localities are situated are given their current political 
names. Where the country is unknown the locality data originally published is 
quoted between inverted commas. Provinces are not cited unless given with the 
published locality data except in the case of South African specimens, for which the 
provinces (if known) are always cited. 

Where a lectotype has been designated a bibliographical reference is given to the 
designation. No lectotypes are designated in this catalogue. 

The museums and other institutions in which the type-material is housed are 
abbreviated in this catalogue as follows. 

ANS Academy of Natural Sciencies of Philadelphia. 

BMNH British Museum (Natural History). 

DEI Deutsches Entomologisches Institut, Eberswalde (East Germany). 

EIETH Entomologisches Institut of the Eidgenossische Technische Hochschule, 

Zurich. 

IEE Institute Espanol de Entomologia, Madrid. 

ILEA Institute e Laboratorio di Entomologia Agraria, Portici. 

IMZU Istituto e Museo di Zoologia of the Universita di Torino, Turin. 

IRSNB Institut Royal des Sciences Naturelles de Belgique, Brussels. 

IZPAN Instytut Zoologiczny of the Polska Akademia Nauk, Warsaw. 

IZU Istituto di Zoologia of the University di Pavia. 

MCSN Museo Civico di Storia Naturale, Genoa. 

MHN Mus6um d'Histoire Naturelle, Geneva. 

MLZA Museu e Laboratorio Zoologico e Antropologico, Lisbon. 

MNHN Museum National d'Histoire Naturelle, Paris. 

MRAC Mus6e Royal de 1'Afrique Centrale, Tervuren. 

NM Naturhistorisches Museum, Vienna. 

NR Naturhistoriska Riksmuseum, Stockholm. 

SAM South African Museum, Cape Town. 

SMN Staatliches Museum fur Naturkunde, Stuttgart. 

TM Terme'szettudomanyi Muzeum, Budapest. 

UM University Museum, Oxford. 

UZI Universitetets Zoologiska Institution, Lund. 

ZM Zoologisches Museum, Hamburg. 

ZMHU Zoologisches Museum of the Humboldt-Universitat, Berlin. 

In the course of compiling this catalogue I have come across one generic and two 
specific names published before Kirby's catalogue of 1906, but not included in it. 
The generic name is Cameronia, which first appeared in the combination Symmetro- 
pleura (Cameronia) africana Brunner in a paper by Karsch (1889 : 450). The name 
was ignored by Brunner (1891) in the supplement to his earlier monograph (Brunner, 
1878) but was quoted, in the same combination, by Bolivar (1906 : 349). As far as 
I know these are the only two occasions on which this name has appeared in print. 
Cameronia is clearly a genus-group name and is available under Article 16 (a) (v) of 
the International Code of Zoological Nomenclature, with Symmetropleura africana 
Brunner as its type-species, by monotypy. It seems highly improbable that Karsch 
intended the name to be regarded as a subgenus of Symmetropleura Brunner ; there is 
no other instance of either him or his contemporary Brunner using a subgeneric name, 
and on the following page of his 1889 paper (op. cit., p. 451) he describes another 



7 8 D. R. RAGGE 

species of Symmetropleura without giving it a subgeneric assignment. It seems more 
likely that Karsch was considering Cameronia as an alternative generic name for 
Symmetropleura africana Brunner, and so cited it in brackets. No subgenera are 
recognized in this catalogue and I have merely listed Cameronia in the synonymy 
of Symmetropleura. If, as seems probable, revisionary work shows that the African 
species at present assigned to the otherwise New World genus Symmetropleura should 
be separated at the generic level, the name Cameronia is available for any resulting 
genus that contains Symmetropleura africana Brunner. 

The two specific names missed by Kirby (1906) are Cestromoecha crassipes Karsch 
(described in a footnote Karsch, 1890 : 364) and Rhegmatopoda brunneri Kirby. 
The latter name, proposed as a nomen novum (Kirby, 1900 : 215), was based on 
Brunner's (1878 : 89) description of a male specimen from Grahamstown in the 
Naturhistorisches Museum, Vienna, then regarded by Brunner as Horatosphaga 
leptocerca Stal, and Kirby's examination of a male specimen from Barberton in the 
British Museum (Natural History). Although these specimens were not described 
by Kirby, the name R. brunneri is available under Article 16 (a) (i) of the International 
Code of Zoological Nomenclature. The name Rhegmatopoda Brunner has since be- 
come a junior synonym of Conchotopoda Karsch (Ragge, 19600 : 325) and brunneri 
is thus listed in this catalogue under Conchotopoda Karsch, forming the new com- 
bination Conchotopoda brunneri (Kirby) comb. n. 

THE STATUS OF THE PH ANEROPTERIN AE 

The inter-relationships and systematic arrangement of the African genera of 
Phaneropterinae will be considered in the course of forthcoming revisionary work, but 
it is appropriate at this stage to consider briefly the status of the group as a whole. 

The Phaneropterinae have been regarded as a subfamily of Tettigoniidae (also 
called at various times, and often in a broader sense, Locustariae, Locustina, Locusti- 
dae, Locustodea and Phasgonuridae) for well over a century : they were given this 
status, together with Mecopoda and Phyllophora (now the type-genera of two other 
subfamilies), by Burmeister (1838 : 684). In 1878 Brunner, in his classic Mono- 
graphie der Phaneropteriden, established the group in its present sense and laid the 
foundation for all subsequent systematic work on the Phaneropterinae. The 
subfamily suffix -inae was not in use at this time, but Brunner referred to the group 
as a " Subfamilie " and used for it the Latin category " tribus ". Brunner reaffirmed 
his classification in his works of 1891 and 1893. The group-name appeared with the 
currently used subfamily suffix in Saussure & Pictet's work of 1897, and was again 
used in this form by Kirby (1906). This usage was adopted almost universally and 
still prevails today (Bei-Bienko, 1954, 1964 ; Blatchley, 1920 ; Brues, Melander & 
Carpenter, 1954; Chopard, 1922, 1943, 1947; Essig, 1947; Grant, 1964; Imms, 1957; 
Zeuner, 19360). 

During this period the general classification of the Ensifera came under the strong 
stabilizing influence of two major works on their morphology and phylogeny, one 
by Ander and the other by Zeuner, published almost simultaneously in 1939. The 
thoroughness and comprehensiveness of these studies clearly makes them the best 
available basis for considering the inter-relationships of the sub-groups of Ensifera, 



AFRICAN PHANEROPTERINAE 79 

and it is significant that the classifications adopted in the two works are almost 
identical. The Phaneropterinae were again regarded as a subfamily of the Tetti- 
goniidae. 

This long period of stability in the status of the Phaneropterinae has been affected 
during the past twenty years by a movement among European orthopterists to raise 
the status of nearly all the groups of Saltatoria, i.e. to elevate most of the families to 
superfamilies or even suborders, and most of the subfamilies to families. This move- 
ment does not stem from new morphological or phylogenetic discoveries but simply 
from the view that these groups are at present ranked at too low a level, or, in one 
case (Dirsh, 1961 : 358), from the belief that family and subfamily rank can be 
objectively defined. The effect of this action on the Phaneropterinae is to raise them 
to family rank, the Tettigoniidae becoming a superfamily more restricted in scope 
than the Tettigonioidea of Ander (op. cit.}, which is left without a name. 

Although there are a few earlier instances of the Phaneropterinae being accorded 
family status (e.g. Burr, 1910 ; Zacher, 1917), the first major recent work in which this 
was done was that of Chopard (1949), and the practice has since been adopted by 
Beier (1955) and in the faunal works of Harz (1957, 1960). Although it seems likely 
that some other modern orthopterists whose specialities lie outside the Ensifera 
would favour family status for the Phaneropterinae, the group is still regarded as a 
subfamily in most recent taxonomic publications in both Europe and America (e.g. 
Bei-Bienko, 1964; Grant, 1964; Karabag, 1964; Ragge, 1964); indeed, some orthop- 
terists (e.g. Uvarov, 1924:492-493; Zeuner, 1936^:107, 19366 : 300-301, 
1939 : 115) have suggested that a number of the other subfamilies of Tettigoniidae 
would be better regarded as tribes. 

There is of course a trend during the earlier stages of the taxonomic study of all 
groups towards the raising in rank of each level of the hierarchy, and it may seem at 
first sight that there is still a need for such a trend in the classification of the Ensifera. 
However, in my view the classification firmly established by Ander (op. cit.) and 
since widely adopted by orthopterists offers an acceptable compromise in the rank 
accorded to each group, and there is no reason why it should not be used, with minor 
modifications, as a framework for taxonomic work on this suborder for many years 
to come. In the case of the Phaneropterinae there is room for only one category 
between the group as a whole and its genera, one that is well provided for by tribes ; 
if the group is regarded as a family, either the subfamily or tribe category will be left 
unemployed. This is not the place for further discussion of the problems of the 
classification of the Ensifera ; it suffices for me to lend my support to the view still 
expressed by the majority of orthopterists that the Phaneropterinae are best 
regarded as a subfamily. 

CATALOGUE OF AFRICAN PHANEROPTERINAE 

PHANEROPTERINAE Burmeister 

Phaneropteridae Burmeister, 1838 : 684. 

Camptoxiphae Serville, [1838] : 399. (Not based on the name of a contained genus and there- 
fore unavailable under Article n (e) of the International Code of Zoological Nomenclature.) 
Phaneropterinae; Saussure & Pictet, 1897 : 310. 



8o D. R. RAGGE 

ANISOTOCHRA Karsch 
Anisotochra Karsch, 1889 : 430. Type-species: Anisotochra gracilipes Karsch, by monotypy. 

gracilipes Karsch, 1889 : 431. Hole-type?, CONGO (KINSHASA): Kuako to Kimpoko (ZMHU, 
Berlin). 

ARANTIA Stal 

Arantia Stal, 1874 : 25. Type-species: Holochlora fatidica Stal, by monotypy. 

accrana Karsch, 1889 : 436. Holotype $, GHANA: Accra (ZMHU, Berlin). 

angustipennis Chopard, 1954 : 3 6 - Holotype <J, GUINEA: Nimba (MNHN, Paris). 

brevipes Chopard, 1954 : 37- J <$ I ? syntypes, GUINEA: Nimba (MNHN, Paris). 

excelsior Karsch, 1889 : 435. Holotype <$, SIERRA LEONE (IZPAN, Warsaw). 

fasciata Walker, 1869 : 378 (Phylloptera) . Holotype $, GAMBIA (BMNH, London). 
atrolineata Brunner, 1891 : 69. Holotype -, "SLAVE COAST" (EIETH, Zurich). 

fatidica Stal, 1873 : 43 (Holochlora). Holotype $, SIERRA LEONE (NR, Stockholm). 

gabunensis Brunner, 1891 : 64. Holotype $, GABON (NM, Vienna). 

gestri Griffini, 1906 : 384. Holotype <J, CONGO (BRAZZAVILLE): N'jole (MCSN, Genoa). 

hydatinoptera Karsch, 1889 : 436. i $, i $ syntypes, NIGERIA (IZPAN, Warsaw). 

latifolia Karsch, iSgoc : 356. i #, i $ syntypes, CAMEROUN : Barombi (ZMHU, Berlin). 

leptocnetnis Karsch, iSgoc : 358. Holotype ., CAMEROUN : Barombi (ZMHU, Berlin). 

matnmisignum Karsch, 1896 : 332. Holotype $, CAMEROUN: Lolodorf (ZMHU, Berlin). 

rnanca Bolivar, 1906 : 332. Holotype <$, CAMEROUN (IEE, Madrid). 

martnorata Karsch, 1889 : 435. Holotype $, "CONGO" (IZPAN, Warsaw). 

melanotus Sjostedt, 1901 : 6. Holotype $, CAMEROUN: Bonge (NR, Stockholm). 

orthocnetnis Karsch, iSgoc : 357. i $ syntype, CAMEROUN; i - syntype, CAMEROUN: Bar- 
ombi (ZMHU, Berlin). 

ovalipennis Chopard, 1954 : 38- Holotype -, GUINEA: Nimba, N'Zo (MNHN, Paris). 

rectifolia Brunner, 1878 : 137. Holotype <J, FERNANDO Poo (NM, Vienna). 

regina Karsch, 1888 : 434. Holotype $, GABON (ZMHU, Berlin). 

retinervis Karsch, 1888 : 437. 2 <$ syntypes, CAMEROUN: Barombi (ZMHU, Berlin); i <J 
syntype, "Chinchoxo" (ZMHU, Berlin); i $ syntype, SIERRA LEONE (IZPAN, Warsaw). 

scurra Karsch, 1896 : 331. Holotype <$, CAMEROUN: Lolodorf (ZMHU, Berlin). 

simplicinervis Karsch, 1889 : 438. i <J syntype, "Chinchoxo"; i - syntype, CONGO (KIN- 
SHASA) : Kuako to Kimpoko (ZMHU, Berlin) . 

spinulosa Brunner, 1878 : 137. i <J, 2 $ syntypes, SOUTH AFRICA: Natal, Durban (NM, 
Vienna). 

tigrina Bolivar, 1906 : 330. Holotype <J, FERNANDO Poo (IEE, Madrid). 

ugandana Rehn, 1914 : 176. Holotype $, UGANDA (DEI, Eberswalde). 

ATLASACRIS Rehn 
Atlasacris Rehn, 1914 : 153. Type-species: Atlasacris peculiaris Rehn, by original designation. 

peculiaris Rehn, 1914 : 154. Holotype <J, "OST-AFRIKA" (ZMHU, Berlin). 

A Z AMI A Bolivar 

Azamia Bolivar, 1906 : 339. Type-species: Azamia biplagiata Bolivar, by monotypy. 

biplagiata Bolivar, 1906 : 341. Holotype <$, CAMEROUN (IEE, Madrid). 

doriae Griffini, 1906 : 395 (Vossia). Holotype <$, FERNANDO Poo: Musola (MCSN, Genoa). 



AFRICAN PHANEROPTERINAE 81 

BONG El A Sjostedt 
Bongeia Sjostedt, 1901 : 9. Type-species: Bongeia puncticollis Sjostedt, by monotypy. 

brevicauda Ebner, 1943 : 265. Holotype ., FERNANDO Poo (DEI, Eberswalde). 
puncticollis Sjostedt, 1901 : 9. Holotype $, CAMEROUN (NR, Stockholm). 

BUEACOLA Sjostedt 

Bueacola Sjostedt, 1912 : 7. Type-species: Bueacola cornigera Sjostedt, by monotypy. 
cornigera Sjostedt, 1912 : 8. Holotype Q*, CAMEROUN: Buea (NR, Stockholm). 

BUETTNERIA Karsch 

Buttneria Karsch, 1889 : 444. Type-species: Buettneria maculiceps Karsch, by monotypy. 
Buettneria Karsch, Brunner, 1891 : 105. 

Stenacropteryx Karsch, 1896 : 339. Type-species: Stenacropteryx eburneiguttata Karsch 
(= Buettneria maculiceps Karsch), by monotypy. 

maculiceps Karsch, 1889 : 445. Holotype ?, CONGO (KINSHASA) : Kuako to Kimpoko (ZMHU, 

Berlin). 

eburneiguttata Karsch, 1896 : 339 (Stenacropteryx). Holotype <$, CAMEROUN : Lolodorf (ZMHU, 
Berlin). 

CATOPTROPTERYX Karsch 

Catoptropteryx Karsch, iSgoc : 361. Type-species: Catoptropteryx guttatipes Karsch, by mono- 
typy (two other species doubtfully included). 

afra Karsch, 1889 : 446 (Caedicia). Holotype ., NIGERIA: Benue (IZPAN, Warsaw). 
apicalis Bolivar, 1893 : 177 (Caedicia). Holotype -, IVORY COAST: Assinie (IEE, Madrid). 
capreola Karsch, 1896 : 332. Holotype <J, CAMEROUN: Lolodorf (ZMHU, Berlin). 
extensipes Karsch, 1896 : 336. Holotype (J, CAMEROUN: Lolodorf (ZMHU, Berlin). 
guttatipes Karsch, iSgoc : 362. Holotype <$, CAMEROUN: Barombi (ZMHU, Berlin). 
irnmaculipennis Karsch, 1896 : 333. Holotype <J, CAMEROUN: Lolodorf (ZMHU, Berlin). 
latipennis Chopard, 1955 : 267. Holotype Q*, SOUTH AFRICA: Cape Province, Tzitzikama 

Forest, Stormsrivierpiek (UZI, Lund). 

maculipennis Karsch, 1896 : 333. Holotype $, CAMEROUN: Lolodorf (ZMHU, Berlin). 
neutralipennis Karsch, 1896 : 334. i $ syntype, TOGO: Misahohe; i - syntype, TOGO: 

Bismarckburg (ZMHU, Berlin). 

nigrospinosa Brunner, 1891 : 97 (Caedicia). Holotype $, CAMEROUN (IZPAN, Warsaw). 
punctulata Karsch, 18906 : 260 (Caedicia). Holotype $, CAMEROUN: Kribi (ZMHU, Berlin). 
signatipennis Karsch, 1896 : 334. i Q* syntype, CAMEROUN: Victoria; i $ syntype, CAMEROUN: 

Lolodorf (ZMHU, Berlin). 

CESTROMOECHA Karsch 

Cestromoecha Karsch, 1893 : 128. Type-species: Poreuomena tenuipes Karsch, by original 
designation. 

crassipes Karsch, iSgoc : 364 (Poreuomena). Holotype $, CAMEROUN (ZMHU, Berlin). 
mundamensis Karsch, 1896 : 338. 1^,1$ syntypes, CAMEROUN: Mundame (ZMHU, Berlin). 
tenuipes Karsch, iSgoc : 363 (Poreuomena). 3- syntypes, CAMEROUN: Barombi (ZMHU, 
Berlin). 



82 D. R. RAGGE 

CONCHOTOPODA Karsch 

Conchotopoda Karsch, 18870. : 44. Type-species: Conchotopoda belcki Karsch, by monotypy. 
Rhegmatopoda Brunner, 1891 : 44. Type-species: Horatosphaga leptocerca Stal, by monotypy. 

belcki Karsch, 1 887*2 : 45. Holotype Q*, SOUTH WEST AFRICA: Damaraland (ZMHU, Berlin). 

brunneri Kirby, 1900 : 215 (Rhegmatopoda) (see p. 78). i $ syntype, SOUTH AFRICA: Trans- 
vaal, Barberton (BMNH, London) ; i $ syntype, SOUTH AFRICA : Cape Province, Grahams- 
town (NM, Vienna). 

crassicauda Ragge, 19600 : 328. Holotype $, SOUTH AFRICA: Transvaal, Pietersburg (SAM, 
Cape Town). 

grallatoria Stal, 1856 : 170 (^Phaneroptera). Holotype $, SOUTH AFRICA: "Cam-aria" (NR, 
Stockholm). 

leptocerca Stal, 1876 : 59 (Horatosphaga). Holotype $, SOUTH WEST AFRICA: Damara (NR, 
Stockholm) . 

parva Ragge, 19600 : 330. Holotype $, SOUTH AFRICA: Orange Free State, Witzieshoek 
(BMNH, London). 

CORYCOMIMA Karsch 

Corycomima Karsch, 1889 : 457 (proposed conditionally, but available from this date under 
Article 17 (8) of the International Code of Zoological Nomenclature). Type-species: Plangia 
earner ata Karsch (= Corycomima flavescens (Walker)), by monotypy. 

Corycomima Karsch, Karsch, 1896 : 343 (proposed unconditionally). 

flavescens Walker, 1869 : 386 (Orophus). Holotype -, GAMBIA (BMNH, London). 

camerata Karsch, 1889 : 457 (Plangia). Holotype $, CAMEROUN: Barombi (ZMHU, Berlin). 

CORY MET A Brunner 

Corymeta Brunner, 1878 : 126. Type-species: Phaneroptera amplectens Schaum, by monotypy. 

amplectens Schaum, 1853 : 777 (Phaneroptera}. Holotype <$, MOZAMBIQUE (ZMHU, Berlin). 

DANNFELTIA Sjostedt 
Dannfeltia Sjostedt, 1901 : 19. Type-species: Dannfeltia nana Sjostedt, by monotypy. 

nana Sjostedt, 1901 : 19. Holotype $, " KONGO " (NR, Stockholm). 

DAPANERA Karsch 

Dapanera Karsch, 1889 : 441. Type-species: Dapanera genuteres Karsch, by monotypy. 

eidrnanni Ebner, 1943 : 263. Holotype <J, FERNANDO Poo (DEI, Eberswalde). 
genuteres Karsch, 1889 : 441. 2 <$ syntypes, GHANA: Accra (ZMHU, Berlin). 
irregularis Karsch, 18906 : 258. i Q*, i ? syntypes, CAMEROUN: Kribi (ZMHU, Berlin). 

DEBRONA Walker 

Debrona Walker, 1870 : 480. Type-species: Debrona cervina Walker, by monotypy. 
Otiaphysa Karsch, 1889 : 453. Type-species: Otiaphysa habetata Karsch ( Debrona cervina 
Walker), by monotypy. 

angustipennis Burr, 1900 : 43 (Otiaphysa). Holotype -, SOMALI REPUBLIC: Whardi Datal 

(UM, Oxford). 
cervina Walker, 1870 : 481. Holotype $, SOUTH AFRICA (BMNH, London). 

habetata Karsch, 1889 : 454 (Otiaphysa). Holotype ., TANZANIA: Usambara (ZMHU, Berlin). 



AFRICAN PHANEROPTERINAE 83 

DIOGENA Brunner 

Diogena Brunner, 1878 : 224. Type-species: Phaneroptera fausta Burmeister, by monotypy. 

denticulata Chopard, in Chopard & Kevan, 1954 : 3 2 &- Holotype $, KENYA: Northern 

Frontier Distr., El Wak (BMNH, London). 
fausta Burmeister, 1838 : 689 (Phaneroptera). Holotype $, SUDAN: Nubia (ZMHU, Berlin). 

DIONCOMENA Brunner 
Dioncomena Brunner, 1878 : 208. Type-species: Dioncomena ornata Brunner, by monotypy. 

ornata Brunner, 1878 : 208. 2 $, i $ syntypes, "ZANZIBAR" (NM, Vienna); i $ syntype, 

"ZANZIBAR" (IEE, Madrid). 
superba Karsch, 1889 : 449. Unknown number of syntypes of both sexes, TANZANIA: Bondei 

and Usambara (ZMHU, Berlin). 

DITHELA Karsch 

Dithela Karsch, iSgoc : 354. Type-species: Dithela rectiloba Karsch, by monotypy. 

acuticercus Sjostedt, 1912 : 5. 2 <J, i $ syntypes, CAMEROUN: Buea (NM, Stockholm). 
rectiloba Karsch, iSgoc : 355. i <$, 2 . syntypes, CAMEROUN : Barombi (ZMHU, Berlin). 

DREPANOPHYLLUM Karsch 

Drepanophyllum Karsch, 18900 : 358. Type-species: Drepanophyllum marmoratum Karsch, 

by monotypy. 
Karschia Brunner, 1891 : 41. Type-species: Karschia corrosa Brunner (= Drepanophyllum 

marmoratum Karsch), by monotypy. 

corrostfoliutn Karsch, 1896 : 326. Unknown number of syntypes of both sexes, CAMEROUN: 

Yaunde and Lolodorf (ZMHU, Berlin). 

furcatum Ragge, 1962^ : 307. Holotype ,$, UGANDA : Entebbe (BMNH, London). 
marmoratum Karsch, iSgoc : 360. Holotype $, CAMEROUN: Barombi (ZMHU, Berlin). 
corrosa Brunner, 1891 : 41 (Karschia). Holotype $, CAMEROUN (IZPAN, Warsaw). 

DUCETIAStal 

Ducetia Stl, 1874 : n. Type-species : Locusta japonica Thunberg, by monotypy (type-locality : 

Japan). 
Paura Karsch, 1889 : 439. Type-species: Paura biramosa Karsch, by subsequent designation 

(Kirby, 1906 : 407). 
Pseudisotima Schulthess, 1898 : 199. Type-species: Pseudisotima punctata Schulthess, by 

monotypy. 
Kuwayamaea Matsumura & Shiraki, 1908 : 7. Type-species: Kuwayamaea sapporensis Mat- 

sumura & Shiraki (= Ducetia chinensis (Brunner)), by original designation (type-locality: 

Japan). 
Telaea Bolivar, 1922 : 201. Type-species: Telaea quadripunctata Bolivar ( Ducetia puncti- 

pennis (Gerstaecker)), by monotypy. 

biramosa Karsch, 1889 : 439 (Paura). Holotype $, TANZANIA: Usambara (ZMHU, Berlin). 
chelocerca Ragge, 1961^ : 197. Holotype <, SOUTH AFRICA: Transvaal, junction of Crocodile 

and Marico Rivers (SAM, Cape Town). 
ENTOM. 22, 3. 3 



84 D. R. RAGGE 

costata Ragge, 19610 : 196. Holotype $, CONGO (KINSHASA): Nyangwe (MRAC, Tervuren). 
crosskeyi Ragge, 19610, : 192. Holotype <$, NIGERIA: Niger Province, Diko (BMNH, London). 
fuscopunctata Chopard, 1954 : 35- Holotype <$, GUINEA: Nimba, Yalanzou (MNHN, Paris). 
loosi Griffini, 1908^ : 204. Holotype -, CONGO (KINSHASA): Popocabacca (IRSNB, Brussels). 
macrocerca Ragge, 19610 : 195. Holotype <$, CONGO (KINSHASA) : Katanga, Kiambi (MRAC, 

Tervuren) . 

parva Ragge, 19610 : 205. Holotype $, SOMALI REPUBLIC: Haud (BMNH, London). 
punctata Schulthess, 1898 : 199 (Pseudisotima). Holotype $, ETHIOPIA: Girma (MCSN, 

Genoa). 
punctipennis Gerstaecker, 1869 : 215 (Phaneroptera) . Holotype <, KENYA: Mt. Ndara 

(ZMHU, Berlin). 

reticulosa Karsch, 1889 : 440 (Paura). Holotype $, KENYA: Mombasa (ZMHU, Berlin). 
quadripunctata Bolivar, 1922 : 201 (Telaea). Holotype <J, KENYA: Simba (lost). 
ramulosa Ragge, 19610: 198. Holotype <$, ZAMBIA: Luano Valley, R. Mulungushi (BMNH, 

London). 

sagitta Ragge, 19610 : 199. Holotype <, ANGOLA: Lepi (BMNH, London). 
vitriala Ragge, 19610 : 204. Holotype $, SOMALI REPUBLIC: Haud (BMNH, London). 

ENOCHLETICA Karsch 
Enochletica Karsch, 1896 : 336. Type-species: Enochletica ostentatrix Karsch, by monotypy. 

ostentatrix Karsch, 1896 : 337. Unknown number of syntypes of both sexes, CAMEROUN: 
Lolodorf (ZMHU, Berlin). 

EPIPHLEBUS Karsch 

Epiphlebus Karsch, 1896 : 325. Type-species: Epiphlebus crypterius Karsch, by monotypy. 

crypterius Karsch, 1896 : 325. Holotype -, SOMALI REPUBLIC (type-depository unknown). 
ruspolii Schulthess, 1898 : 197. Holotype -, SOMALI REPUBLIC: Ganana (MCSN, Genoa). 

EULIOPTERA Ragge 

Eulioptera Ragge, 19560 : 266. Type-species: Phaneroptera reticulata Brunner, by original 
designation. 

breviala Ragge, 19560 : 277. Holotype $, CONGO (KINSHASA): Kivu, Kibumba (MRAC, 

Tervuren) . 

longicerca Ragge, 19560 : 273. Holotype $, CONGO (KINSHASA) : Kinshasa (MRAC, Tervuren). 
reticulata leptomorpha Ragge, 19560 : 271. Holotype , ZAMBIA: Mporokoso distr., Mweru 

Wa Ntipa (BMNH, London). 
reticulata reticulata Brunner, 1878 : 213 (Phaneroptera reticulata}. Holotype -, SOUTH 

AFRICA: Cape Province, Grahamstown (NM, Vienna). 
spinulosa Ragge, 19560 : 274. Holotype $, CONGO (KINSHASA): Kasenyi (BMNH, London). 

EURYCORYPHA Stal 

Eurycorypha Stal, 1873 : 40. Type-species: Phylloptera cereris Stal, by original designation. 
Myrmecophana Brunner, 1883 : 248. Type-species: Myrmecophana fallax Brunner, by mono- 
typy. 

adicra Karsch, 18926 : 75. Holotype ?, CAMEROUN: Buea (ZMHU, Berlin). 

aequatorialis Krauss, 1890 : 663. Holotype <j>, ROLAS I. (Gulf of Guinea) (SMN, Stuttgart). 



AFRICAN PHANEROPTERINAE 85 

arabica arabica Uvarov, 1936 : 534. Holotype <J, SAUDI ARABIA: S. Hedjaz, Taif (BMNH, 

London) . 

arabica media Uvarov, 1936 : 535. 3 <$ syntypes, SAUDI ARABIA: El Kubar (BMNH, London). 
arabica reducta Uvarov, 1936 : 536. Holotype $, SOUTH ARABIA: Hadramaut, Wadi Humum, 

nr. Makalla (ZM, Hamburg). 

brevicollis Stal, 1876 : 61. Holotype -, SOUTH WEST AFRICA: Ovambo (NR, Stockholm). 
canaliculata Karsch, 18906:261. Holotype^, CAMEROUN: Kribi (ZMHU, Berlin). 
cereris Stal, 1856 : 170 (Phylloptera) . Holotype $, SOUTH AFRICA: Natal, Durban (NR, 

Stockholm) . 

compressa Walker, 1869 : 386 (Orophus). Holotype $, SOUTH AFRICA (BMNH, London). 
cuspidata Krauss, 1901 : 292. i $ syntype, SOUTH WEST AFRICA: Okahandja; i $ syntype, 

SOUTH WEST AFRICA: Windhoek (SMN, Stuttgart). 

darlingi Uvarov, 1936 : 536. Holotype <J, SOUTH YEMEN: Makhnung (BMNH, London). 
diminuta Chopard, 1938 : 121. Holotype <J, KENYA: Turkana Desert, Lodwar (MNHN, 

Paris) . 
fallax Brunner, 1883 : 248 (Myrmecophana). Nymphal holotype, sex unknown, SUDAN: 

Ambucarra (IRSNB, Brussels). 

montana Sjostedt, 1901. Holotype ?, CAMEROUN: Mapanja (NR, Stockholm). 
tnutica Karsch, 1892 : 327. Holotype $, CAMEROUN: Barombi (ZMHU, Berlin). 
ornatipes Karsch, 18906 : 260. Holotype <$, CAMEROUN: Kribi (ZMHU, Berlin). 
proserpinae Brunner, 1878 : 274. Holotype -, SOUTH AFRICA: Natal, Durban (NM, Vienna). 
punctipennis Chopard, 1938 : 121. Holotype $, KENYA: Mt. Elgon (MNHN, Paris). 
securifera Brunner, 1878 : 275. Holotype $, SOUTH WEST AFRICA: Herrero-Land (NM, 

Vienna) . 
simillima Chopard, in Chopard & Kevan, 1954 : 33- Holotype , KENYA: N. Turkana, 

Lokitaung (BMNH, London). 
spinulosa Karsch, 1889 : 455. Holotype $, CONGO (KINSHASA): Kuako to Kimpoko (ZMHU, 

Berlin). 
strangulata Walker, 1869 : 385 (Orophus). i $ syntype, SIERRA LEONE; i $ syntype, "WEST 

AFRICA" (BMNH, London). 

stylata Stal, 1873 : 43. Holotype , SIERRA LEONE (NR, Stockholm). 
sudanensis Giglio-Tos, 1907 : 18. Holotype <$, SUDAN: Port Sudan (IMZU, Turin). 
varia Brunner, 1891 : 136. i $, i . syntypes, TANZANIA: Kilimanjaro (NM, Vienna). 
velicauda Karsch, 1893 '.131. i <J, i $, i nymphal syntypes, TOGO: Bismarckburg (ZMHU, 

Berlin). 
zebrata Bruner, 1920 : 135. Holotype $, CAMEROUN: Lolodorf (ANS, Philadelphia). 



GABONELLA Uvarov 

Gabonia Bolivar, 1906 : 327. Type-species: Gabonia cothurnata Bolivar, by monotypy. 

(Homonym of Gabonia Jacoby, 1893 : 101.) 
Gabonella Uvarov, i94oa : 174 (proposed as replacement name for Gabonia Bolivar). 

cothurnata Bolivar, 1906 : 328 (Gabonia). 2 <$, 3 $ syntypes, CAMEROUN (IEE, Madrid). 



GEL A TOPOIA Brunner 

Gelatopo'ia Brunner, 1891 : in. Type-species: Gelatopoia bicolor Brunner, by monotypy. 

Gelatopoia Brunner, Bolivar, 1906 : 349. 

Gelatopoea Kirby, 1906 : 439. (Unjustified emendation.) 

bicolor Brunner, 1891 : 112. Holotype <J, SIERRA LEONE (IZPAN, Warsaw). 



86 D. R. RAGGE 

GOETIA Karsch 

Goetia Karsch, 18920, : 322. Type-species: Goetia galbana Karsch, by monotypy. 
Goetia Karsch, Kirby, 1906 : 428. 

dimidiata Bolivar, 1906 : 335. i <J, i $ syntypes, CAMEROUN; i <J, i $> syntypes, FERNANDO 

Poo; i $, i $ syntypes, no data (IEE, Madrid). 
galbana Karsch, 18920 : 323. Holotype $, CAMEROUN: Barombi (ZMHU, Berlin). 



GONATOXIA Karsch 

Gonatoxia Karsch, 1889 : 441. Type-species: Gonatoxia immaculata Karsch, by subsequent 
designation (Kirby, 1906 : 424). 

immaculata Karsch, 1889 : 442. Holotype $, TANZANIA: Usambara (ZMHU, Berlin). 
maculata Karsch, 1889 : 442. Holotype $, SOMALI REPUBLIC (IZPAN, Warsaw). 



GRAVENREUTHIA Karsch 

Gravenreuthia Karsch, 18926 : 73. Type-species: Gravenreuthia saturata Karsch, by monotypy. 
saturata Karsch, 18926 : 74. i <J, 2 <j> syntypes, CAMEROUN : Buea (ZMHU, Berlin). 

HARPOSCEPA Karsch 

Harposcepa Karsch, 1896 : 328. Type-species: Harposcepa lobulipennis Karsch, by monotypy. 
1896. 

karschiana Schulthess, 1898 : 202. Holotype -, SOMALI REPUBLIC (or ETHIOPIA?): Gubbet 

(IZU, Pavia). 
lobulipennis Karsch, 1896 : 329. Holotype ., SOMALI REPUBLIC (type-depository unknown) . 

HIMERTULA Uvarov 

Himerta Brunner, 1878 : 118. Type-species: Himerta marginata Brunner, by monotypy (type- 
locality: India). (Homonym of Himerta Foerster, 1868 : 200.) 
Himertula Uvarov, 19400 : 174 (proposed as replacement name for Himerta Brunner). 

feana Griffmi, 1906:380 (Himerta). Holotype <J, CONGO (BRAZZAVILLE): Nkogo (MCSN, 
Genoa). 

HORATOSPHAGA Schaum 

Horatosphaga Schaum, 1853 : 778. Type-species: Horatosphaga serrifera Schaum, by monotypy. 

Plegmatoptera Karsch, 1889 : 429. Type-species: Plegmatoptera reticulata Karsch, by monotypy. 
(Homonym of Plegmatoptera Spinola, 1839 : 283.) 

Pantolepta Karsch, 1889 : 429. Type-species: Pantolepta heteromorpha Karsch, by monotypy. 

Pachypyga Karsch, 1893 : 124. Type-species: Pachypyga inclusa Karsch, by monotypy. 
(Homonym of Pachypyga Steinheil, 1873 : 564.) 

Euthyphlebia Schulthess, 1898 : 204. Type-species : Euthyphlebia parallela Schulthess (= Horato- 
sphaga heteromorpha (Karsch)), by monotypy. 



AFRICAN PHANEROPTERINAE 87 

Thaumatoxenia Kirby, 1909 : 65. Type-species: Thaumatoxenia leggei Kir by, by monotypy. 
Keniacola Sjostedt, 1912 : 13. Type-species: Keniacola gracilis Sjostedt, by monotypy. 
Schulthessinia Sjostedt, 1912 : 15. Type-species: Conchotopoda ruspolii Schulthess, by original 

designation. 

Pachypygiana Strand, 1921 : 106 (proposed as replacement name for Pachypyga Karsch). 
Eupantolepta Bolivar, 1922 : 200. Type-species: Eupantolepta regularis Bolivar, by monotypy. 
Plegmatia Uvarov, 19406 : 116 (proposed as replacement name for Plegmatoptera Karsch). 

concava Ragge, 1960 : 308. Holotype <$, UGANDA: Gulu (BMNH, London). 

crosskeyi Ragge, 1960 : 318. Holotype <$, NIGERIA: Niger Province, Diko (BMNH, London). 

diminuta Chopard, in Chopard & Kevan, 1954 : 3 X 9 (Peronura). Holotype <$, KENYA: Moyale 

(BMNH, London). 

elgonis Chopard, 1938 : 119 (Peronura). Holotype <$, KENYA: Mt. Elgon (MNHN, Paris). 
elongata Rehn, 1914 : 163 (Conchotopoda). Holotype $, CONGO (KINSHASA): Usumbura 

(ZMHU, Berlin). 
? kandti Rehn, 1914 : 167 (? Horatosphaga). Holotype ?, CONGO (KINSHASA): Lake Kivu 

(ZMHU, Berlin). 

gracilis Sjostedt, 1912: 14 (Keniacola). Holotype <J, KENYA: Mt. Kenya (NR, Stockholm) 
heterotnorpha Karsch, 1889 : 430 (Pantolepta) . Lectotype , KENYA: Mombasa (ZMHU, 

Berlin); designated by Ragge (1960 : 298). 
hoehneli Brunner, 1891 : 44 (Plegmatoptera). Holotype -, TANZANIA: Kilimanjaro (NM, 

Vienna) . 
parallela Schulthess, 1898 : 205 (Euthyphlebia) . Holotype $, ETHIOPIA: Dolo (MCSN, 

Genoa) . 
kilimandjarica Sjostedt, 1909 : 132 (Pantolepta). Lectotype <$, TANZANIA: Mt. Meru (NR, 

Stockholm); designated by Ragge (1960 : 298). 
gracilis Bolivar, 1922 : 198 (Plegmatoptera). Holotype $, KENYA: Loroghi Mtns. (MNHN, 

Paris). 
inclusa Karsch, 1893 : 125 (Pachypyga). i , 5 - syntypes, TOGO: Bismarckburg (ZMHU, 

Berlin). 
leggei Kirby, 1909 : 65 (Thaumatoxenia). Lectotype <J, UGANDA: E. Ruwenzori, Mubuku 

Valley (BMNH, London); designated by Ragge (1960 : 292). 
robusta Rehn, 1914 : 156. Holotype?, CONGO (KINSHASA): 90 km. W. of Albert-Nyanza 

(ZMHU, Berlin). 
amplipennis Rehn, 1914 : 165 (Conchotopoda). Holotype $, CONGO (KINSHASA): Lake Kivu, 

Idjwi I. (ZMHU, Berlin). 

affinis Bolivar, 1922 : 198 (Peronura). Holotype $, KENYA: Escarpment (MNHN, Paris). 
linearis Rehn, 1914 : 167 (Conchotopoda). Holotype <$, UGANDA: Mpororo (ZMHU, Berlin). 
? karschi Rehn, 1914 : 161 (Pachypyga). Holotype $, CONGO (KINSHASA) : Ruanda, Lake. 

Mohasi (ZMHU, Berlin). 

longipes Bolivar, 1922 : 197 (Peronura). Holotype $, KENYA: Naivasha (MNHN, Paris). 
tnagna Ragge, 1960 : 304. Holotype $, ETHIOPIA: nr. Scillare (BMNH, London). 
media Ragge, 1960 : 299. Holotype <$, ZAMBIA: Lusaka (BMNH, London). 
meruensis Sjostedt, 1909 : 130 (Plegmatoptera). Lectotype <J, TANZANIA: Mt. Meru (NR, 

Stockholm); designated by Ragge (1960 : 311). 
montivaga Sjostedt, 1909 : 129 (Peronura). Lectotype <J, TANZANIA: Kilimanjaro (NR, 

Stockholm); designated by Ragge (1960 : 312). 
nomirna Karsch, 1896 : 327 (Peronura). i 5*, 2 $ syntypes, TANZANIA: Mpwapwa (ZMHU, 

Berlin). 

nuda Ragge, 1960 : 309. Holotype <J, SUDAN: Imatong Mtns., Kippia (BMNH, London). 
regularis Bolivar, 1922 : 200 (Eupantolepta). Holotype <J, KENYA: S. of Lake Rudolph 

(MNHN, Paris). 

? reticulata Karsch, 1889 : 429 (Plegmatoptera). Holotype ., TANZANIA: Lake Tanganyika 
(ZMHU, Berlin). 



88 D. R. RAGGE 

ruspolii Schulthess, 1898 : 204 (Conchotopoda) . Holotype <J, KENYA: Balessa (MCSN, Genoa). 
? rivae Schulthess, 1898 : 201 (Peronura). Holotype ?, ETHIOPIA: Dolo (MCSN, Genoa). 
trochlearis Bolivar, 1922 : 199. Holotype $, KENYA: S. of Lake Rudolph (MNHN, Paris). 
? viridis Chopard, in Chopard & Kevan, 1954: 318 (Peronura). Holotype ., KENYA: Isiolo 

distr., Bambota (BMNH, London). 
samburu Kevan, in Chopard & Kevan, 1954: 320 (Peronura). Holotype <J, KENYA: Samburu 

(BMNH, London). 
serrifera Schaum, 1853 : 778. Holotype $, MOZAMBIQUE (ZMHU, Berlin). 

vosseleri Karny, 1910 : 52 (Pantolepta) . 10 $ syntypes, SOUTH WEST AFRICA: Okahandja; 

i <J syntype, SOUTH WEST AFRICA: Kung-Buschmannland (ZMHU, Berlin). 
multivenosa Chopard, 1935 : 2 (Pantolepta). Holotype $, MOZAMBIQUE: Nova Chupanga, 

nr. Chemba (MNHN, Paris). 
somali Schulthess, 1898 : 200 (Peronura). Holotype ?, SOMALI REPUBLIC: Lugh (MCSN, 

Genoa) . 
peeli Burr, 1900 : 44 (Rhegmatopoda) . Holotype <J, SOMALI REPUBLIC: Whardi Datal (UM, 

Oxford). 
stuhlmanni Karsch, 1896 : 329 (Conchotopoda). Holotype <J, "AFRICA ORIENTALIS": Buginda 

(ZMHU, Berlin). 
kasindina Rehn, 1914:160 (? Horatosphaga). Holotype ?, CONGO (KINSHASA): Kasindi 

(ZMHU, Berlin). 
stylifera Karny, 1910 : 51 (Pantolepta). Holotype $, SOUTH WEST AFRICA: Windhoek 

(ZMHU, Berlin). 

vicina Chopard, in Chopard & Kevan, 1954 : 3 J 8 (Peronura). Holotype $, KENYA: Marsabit, 
Chopa Gof (BMNH, London). 

ITOKIIA Sjostedt 

Itokiia Sjostedt, 1901 : 10. Type-species: Itokiia silvarum Sjostedt, by monotypy. 
silvarum Sjostedt, 1901 : 10. Holotype <$, CAMEROUN : Itoki (NR, Stockholm). 

IVENSIA Bolivar 

Ivensia Bolivar, 1890 : 218. Type-species: Ivensia uncinata Bolivar, by monotypy. 
uncinata Bolivar, 1890 : 218. Holotype <J, "Quango" (MLZA, Lisbon). 

JAPYGOPHANA Carl 

Japygophana Carl, 1921 : 301. Type-species: Japygophana peloti Carl, by monotypy. 
peloti Carl, 1921 : 302. Holotype <J, GABON (MHN, Geneva). 

KEVANIELLA Chopard 

Kevaniella Chopard, in Chopard & Kevan, 1954 : 33 2 - Type-species: Kevaniella bipunctata 
Chopard, by original designation. 

bipunctata Chopard, in Chopard & Kevan, 1954: 333. Holotype <$, KENYA: Mandera distr., 
Rahmu (BMNH, London). 

LAMECOSOMA Ragge 

Lamecosoma Ragge, 1960^ : 331. Type-species: Lamecosoma tenuis Ragge, by original designa- 
tion. 

inermis Ragge, 19616 : 217. Holotype $, KENYA: Nairobi, Karura Forest (BMNH, London). 
tenuis Ragge, 19600 : 332. Holotype <J, ZAMBIA: Kipundu (MRAC, Tervuren). 



AFRICAN PHANEROPTERINAE 89 

LEIODONTOCERCUS Chopard 

Leiodontocercus Chopard, 1954 : 83. Type-species: Leiodontocercus angustipennis Chopard, by 
original designation. 

angustipennis Chopard, 1954 : 84. Holotype $, GUINEA: Nimba (MNHN, Paris). 
condylus Ragge, 19626 : 15. Holotype <$, CONGO (KINSHASA): Kibali-Ituri, Yindi (MRAC, 

Tervuren) . 
malleus Ragge, 19626 : 14. Holotype <?, GHANA: Western Region, nr. Wiawso (BMNH, 

London) . 

LEPTOPHYES Fieber 

Leptophyes Fieber, 1853 : 174. Type-species: Barbitistes albovittatus Kollar, by monotypy 
(type-locality : Austria) . 

antinorii Bormans, 1880 : 217. Holotype $, ETHIOPIA: Shoa (MCSN, Genoa). 

LET AN A Walker 

Letana Walker, 1869 : 277. Type-species: Letana linearis Walker, by monotypy (type-locality: 

India). 
Pyrrhicia Stal, 1873 : 41. Type-species: Phaneroptera rubescens Stal, by original designation 

(type-locality: Hong Kong). 
Pyrrhizia Brunner, 1891 : 55. (Unjustified emendation.) 

conradti Bolivar, 1906 : 329 (Pyrrhicia). Holotype <, CAMEROUN (IEE, Madrid). 
zanzibarica Brunner, 1891 : 55 (Pyrrhizia). Holotype $, "ZANZIBAR" (NM, Vienna). 

MANGOMALOBA Sjostedt 
Mangomaloba Sjostedt, 1902 : 14. Type-species: Mangomaloba monticola Sjostedt, by mono- 

typy- 

angustipennis Chopard, 1958 : 76. Holotype $, PRINCIPE (MNHN, Paris). 
latipennis Chopard, 1954 : 86. Holotype <$, GUINEA: Nimba (MNHN, Paris). 
monticola Sjostedt, 1901 : 15. Holotype <J, CAMEROUN: Mt. Cameroon (NR, Stockholm). 
royi Chopard, 1954 : 85. Holotype <J, GUINEA: Nimba (MNHN, Paris). 

MELIDIA Stal 
Melidia Stal, 1876 : 60. Type-species: Melidia brunneri Stal, by monotypy. 

brunneri Stal, 1876 : 60. Holotype ?, SOUTH WEST AFRICA: Damara (NR, Stockholm). 
kenyensis Chopard, in Chopard & Kevan, 1954:326. Holotype <j>, KENYA: Moyale distr., 

Yasere (BMNH, London). 
laminata Chopard, in Chopard & Kevan, 1954 : 3 2 4- Holotype <J, KENYA: Wajir distr., El 

Katulo (BMNH, London). 

MERUTERRANA Sjostedt 

Meruterrana Sjostedt, 1912 : 10. Type-species: Meruterrana elegans Sjostedt, by monotypy. 
elegans Sjostedt, 1912 : 12. 4 <J syntypes, TANZANIA: Mt. Meru (NR, Stockholm). 



90 D. R. RAGGE 

MILITITSA Burr 

Milititsa Burr, 1900 : 42. Type-species: Milititsa somaliensis Burr, by monotypy. 
somaliensis Burr, 1900 : 43. Holotype ?, SOMALI REPUBLIC: Whardi Datal (UM. Oxford). 

MILTINOBATES Sjostedt 

Miltinobates Sjostedt, 1901 : 15. Type-species: Miltinobates blandus Sjostedt, by monotypy. 
blandus Sjostedt, 1901 : 16. Holotype $, "KONGO" (NR, Stockholm). 

MONTEIROA Karsch 

Monteiroa Karsch, 1889 : 458. Type-species: Monteiroa latifrons Karsch, by monotypy. 
latifrons Karsch, 1889 : 458. i $, i $ syntypes, MOZAMBIQUE: Delagoa Bay (ZMHU, Berlin). 

MONTICOLARIA Sjostedt 

Monticolaria Sjostedt, 1909 : 128. Type-species: Monticolaria meruensis Sjostedt, by PRE- 
SENT DESIGNATION. 

Both the species listed below were originally included in Monticolaria Sjostedt, but neither was 
designated as the type-species. I here designate Monticolaria meruensis Sjostedt as the type- 
species of Monticolaria Sjostedt. 

kilimandjarica Sjostedt, 1909 : 129. 2 $ syntypes, TANZANIA: Kilimanjaro (NR, Stockholm). 
meruensis Sjostedt, 1909 : 128. 10 <, 3 $ syntypes, TANZANIA: Meru (NR, Stockholm). 

MORGENIA Karsch 
Morgenia Karsch, 18906 : 261. Type-species: Morgenia hamuligera Karsch, by monotypy. 

hamuligera Karsch, 18906 : 263. Holotype $, CAMEROUN : Kribi (ZMHU, Berlin). 

melica Karsch, 1893 : 196. Holotype $, CAMEROUN : Victoria (ZMHU, Berlin). 

modulata Karsch, 1896 : 340. i <J, i $ syntypes, CAMEROUN : Lolodorf (ZMHU, Berlin). 

rubricornis Sjostedt, 1913 : 4. Holotype $, CONGO (KINSHASA): Mukimbungu (NR, Stock- 
holm) . 

spathulifera Griffini, 1908 : 209. i $ syntype, " Bussanga " ; i ? syntype, CAMEROUN : Mukonje 
Farm (IRSNB, Brussels). 

MYLLOCENTRUM Ragge 

Myllocentrum Ragge, 19626 : 15. Type-species: Phlaurocentrum stigmosum Karsch, by original 
designation. 

stigmosum Karsch, 1896 : 336 (Phlaurocentrum}. Holotype $, CAMEROUN: Lolodorf (ZMHU, 
Berlin). 

ODONTURA Rambur 

Odontura Rambur, 1838 : 44. Type-species: Barbitistes glabricauda Charpentier, by subsequent 
designation (Kirby, 1906 : 386) (type-locality: Portugal). 

algerica Brunner, 1878 : 75. i $, i <j> syntypes, ALGERIA (NM, Vienna). 
borrei Bolivar, 1878 : 71. Unknown number of $ syntypes, ALGERIA: Constantine Province 
(IRSNB, Brussels). 



AFRICAN PHANEROPTERINAE 91 

brevis Werner, 1932 : 114. Holotype <J, MOROCCO: Ifrane (NM, Vienna). 

capensis Walker, 1869 : 242. Holotype $, SOUTH AFRICA (BMNH, London). 

liouvillei Werner, 1929 : 184. Unknown number of <J syntypes, MOROCCO: nr. Tiflet, Mamora 

Forest; i Q* syntype, MOROCCO: Malay Idris (NM, Vienna). 
maroccana Bolivar, 1908 : 333 (as var. of Odontura spinulicauda; raised to sp. by Bolivar, 

1914 : 237). 7 (J, i $ syntypes, MOROCCO: Marrakesh (IEE, Madrid). 
teknicus Morales, 1947 : 250 (as subsp. of Odontura liouvillei}. Holotype Q*> IFNI: Zoco 

Tlata Sboya (IEE, Madrid). 

microptera Chopard, 1943 : 107. Holotype?, ALGERIA: Ain Mlila (MNHN, Paris). 
moghrebica Morales, 1950 : 163. Holotype <J, MOROCCO: Isaguen (IEE, Madrid). 
plasoni Ebner, 1915 : 419. 3 <J, i $ syntypes, TANZANIA: Tassamaganga (NM, Vienna). 
pulchra Bolivar, 1914 : 237. 3 <J, 3 $ syntypes, MOROCCO: Tangier (IEE, Madrid). 
quadridentata Krauss, 1893 : xcvi. Unknown number of syntypes of both sexes, ALGERIA: 

Tessala (type-depository unknown). 
terniensis Finot, 1893 : xxx. Unknown number of syntypes of both sexes, ALGERIA: Terni, 

Sebdou and Tlemcen (MNHN, Paris). 
spinulicauda Rambur, 1838 : 45. Lectotype $, SPAIN (IEE, Madrid); designated by Uvarov 

(1948 : 380). 

stenoxypha Fieber, 1853 : 187 (Barbitistes). Holotype $, SICILY (type-depository unknown). 
uvarovi Werner, 1929 : 184. Holotype $, MOROCCO: Fez (NM, Vienna). 

OXYECOUS Chopard 
Oxyecous Chopard, 1935 : 4. Type-species: Oxyecous lesnei Chopard, by monotypy. 

apertus Ragge, 19566 : 187. Holotype <J, KENYA: Rabai (BMNH, London). 
lesnei Chopard, 1935 : 4. Holotype <$, MOZAMBIQUE: nr. Vile Pery (MNHN, Paris). 
magnus Ragge, 19566 : 188. Holotype Q*. TANZANIA: Kilosa (BMNH, London). 
undulatus Ragge, 19566 : 190. Holotype Q*. UGANDA: Bugoma Forest (BMNH, London). 

PARAPYRRHICIA Brunner 

Parapyrrhicia Brunner, 1891 : 149. Type-species: Parapyrrhicia zanzibarica Brunner, by 
monotypy. 

zanzibarica Brunner, 1891 : 149. Holotype $, "ZANZIBAR" (NM, Vienna). 

PARDALOTA Brunner 

Pardalota Brunner, 1878 : 133. Type-species: Pardalota versicolor Brunner, by monotypy. 

asymmetrica Karsch, 1896 : 330. 7 syntypes of both sexes, "AFRICA ORIENTALIS CENTRALIS " : 

Kirewia; Itimba; Bundeko; Bugundi; Sossian-Ngoroine, Massai Territory; W. Victoria 

Nyansa, Ussoga-Uganda (ZMHU, Berlin). 
cloetensi Griffini, igoSa : 77. Holotype ., CONGO (KINSHASA) : Beni Bendi, Sankuru (IRSNB, 

Brussels) . 
haasi Griffini, 1908^ : 76. 2 $, i $ syntypes, CONGO (KINSHASA): Luluabourg (IRSNB, 

Brussels) . 
karschiana Enderlein, 1907 : 197. 2 Q* syntypes, TANZANIA: Dar es Salaam (i in ZMHU, 

Berlin; i in IZPAN, Warsaw). 

reimeri La Baume, 1911 : 317. 2 $, 3 nymphal syntypes, TANZANIA: Lindi (ZMHU, Berlin). 
superba Sjostedt, 1913 : 2. Holotype $, TANZANIA: Lukuledi (NR, Stockholm). 
versicolor Brunner, 1878 : 134. Unknown number of syntypes of both sexes, "Chinchoxa in 

Benguela " (i $ in ZMHU, Berlin; i $ in NM, Vienna). 



92 D. R. RAGGE 

PERONURA Karsch 

Peronura Karsch, 1889 : 426. Type-species: Peronura clavigera Karsch, by subsequent desig- 
nation (Kir by, 1906 : 391). 

clavigera Karsch, 1889 : 427. 2$ syntypes, KENYA: Mombasa (ZMHU, Berlin). 

PEROPYRRHICIA Brunner 

Peropyrrhicia Brunner, 1891 : 37. Type-species: Dichopetala massaiae Bormans, by monotypy. 

cooperi Uvarov, 1934 : 59^. Holotype $, ETHIOPIA: Jem-Jem Forest (BMNH, London). 
maculata Schulthess, 1898 : 198. i $, i ? syntypes, ETHIOPIA: Biddwara (MCSN, Genoa). 
massaiae Bormans, 1880 : 218 (Dichopetala). 2 $ syntypes (i nymphal), ETHIOPIA: Let- 

Marefia (MCSN, Genoa). 
scotti Uvarov, 1934 : 597- Holotype <$, ETHIOPIA: between Jem-Jem and Addis Ababa 

(BMNH, London). 

PHANEROPTERA Serville 

Phaneroptera Serville, 1831 : 158. Type-species: Gry llus falcata Poda, by subsequent designation 

(Hemming, 1944 : 211) (type-locality unknown) . 

Anevota Caudell, 1921 : 488. Type-species: Gryllus falcata Poda, by original designation. 
Paranerota Karny, 1926 : 105. Type-species: Phaneroptera gracilis Burmeister, by original 

designation. 
Euanerota Karny, 1927 : 12. Type-species: Phaneroptera brevis Serville, by tentative original 

designation (see Ragge, 1956 : 206) (type-locality: Java). 

acaciae Chopard, in Chopard & Kevan, 1954 : 3 2 3- Holotype <J, KENYA: Mandera distr., 

Rahmu (BMNH, London). 

africana Steinmann, 1966: 411. Holotype <$, GUINEA: Conakry (TM, Budapest). 
albida Walker, 1869 : 335. Holotype <J, no data (BMNH, London). 
fragilis Ragge, 19606 : 245. Holotype <$, ETHIOPIA: Danot (BMNH, London). 
gracilis Burmeister, 1838 : 690. i $, 2 $ syntypes, JAVA (type-depository unknown). 
roseata Walker, 1869 : 343. Holotype ?, INDIA (BMNH, London). 
marginalis Brunner, 1878 : 214. Holotype?, SOUTH AFRICA (ZMHU, Berlin). 
indica Brunner, 1878 : 215. Lectotype <$, INDIA: Himalayas (NM, Vienna); designated by 

Ragge (19606 : 250). 
elongata Brunner, 1878 : 217. Lectotype?, JAVA (ZMHU, Berlin); designated by Ragge 

(19560 : 251). 

subcarinata Bolivar, 1900 : 764. Holotype 3, INDIA: Madura (IEE, Madrid). 
longispina Ragge, 19560 : 244. Holotype (J, UGANDA: Ruwenzori Range, Kilembe (BMNH, 

London) . 

maculosa Ragge, 19560 : 243. Holotype $, CAMEROUN: Lolodorf (ZMHU, Berlin). 
magna Ragge, 19560 : 242. Holotype $, UGANDA: Mabira Forest (BMNH, London). 
minima Brunner, 1878 : 214. Holotype $, EGYPT (ZMHU, Berlin). 
nana nana Fieber, 1853 : 173 (as sp.). Type-locality: PORTUGAL; type-material lost. 

quadripunctata Brunner, 1878 : 212. Lectotype $, ITALY: Trieste (NM, Vienna); designated by 

Ragge (1956 : 233). 

nana sparsaSt&l, 1856 : 170 (assp.). Holotype?, SOUTH AFRICA: "Cam-aria" (NR, Stockholm). 
lurida Walker, 1869 : 339. Holotype <J, "W. AFRICA" (erroneously cited as "Natal" in 

original description) (BMNH, London). 
tetrasticta Gerstaecker, 1869 : 215. Holotype $, TANZANIA: Uru (ZMHU, Berlin). 



AFRICAN PHANEROPTERINAE 



93 



conspersa Still, 1874 : 29 (apparently proposed unnecessarily as replacement name for Phan- 

eroptera sparsa Stal). 

punctulata Burr, 1900 : 41. Holotype <$, SOMALI REPUBLIC: Whardi Datal (UM, Oxford). 
tenuicerca Ramme, 1951 : 348. Holotype $, LEBANON: Djezin (ZMHU, Berlin). 
nigropunctata Chopard, 1955 : 269. Holotype <$, SOUTH AFRICA: Cape Province, Drakens- 

bergen, 8 miles E.N.E. of Rhodes (UZI, Lund). 
parva Ragge, 1956^ : 240. Holotype $, CONGO (KINSHASA): Kasongo (MRAC, Tervuren). 

PHANEROPTILA Uvarov 

Phaneroptila Uvarov, in Uvarov & Popov, 1957 : 363. Type-species: Phaneroptila insularis 
Uvarov, by monotypy. 

insularis Uvarov, in Uvarov & Popov, 1957 : 3^3- Holotype <$, SOCOTRA: Hijama (BMNH, 
London). 

PHAULULA Bolivar 

Phaula Brunner, 1878 : 167. Type-species: Phaula laevis Brunner, by subsequent designation 
(Kirby, 1906 : 425) (type-locality: Philippine Is.). (Homonym of Phaula Thomson, 1857 : 

303-) 
Phaulula Bolivar, 1906 : 347 (proposed as replacement name for Phaula Brunner). 

inconspicua Brunner, 1891 : 83 (Phaula). Holotype -, CAMEROUN (NM, Vienna). 
lenzi Brunner, 1891 : 83 (Phaula). Unknown number of <$ syntypes, CAMEROUN (Liibeck, 
museum unknown) and GABON (NM, Vienna). 

PHLAUROCENTRUM Karsch 

Phlaurocentrum Karsch, 1889 : 445. Type-species: Phlaurocentrum latevittatum Karsch, by 
monotypy. 

latevittatum Karsch, 1889:446. Holotype?, CONGO (KINSHASA): Kuako to Kimpoko 

(ZMHU, Berlin). 

lobatum Ragge, 19626 : 9. Holotype <$, CONGO (KINSHASA): Eala (MRAC, Tervuren). 
maculatum Ragge, 19626 : 10. Holotype <$, CONGO (KINSHASA) : Mawambi-Irumu (NM, 

Vienna) . 

mecopodoides Karsch, 18920! : 321. Holotype $, CAMEROUN: Barombi (ZMHU, Berlin). 
tuberosum Ragge, 19626 : 8. Holotype $, UGANDA: Mabira Forest (BMNH, London). 
turbatum Walker, 1869 : 340 (Phaneroptera) . Holotype <, " CONGO " (BMNH, London). 

PHYSOCORYPHA Karsch 

Physocorypha Karsch, 1896 : 344. Type-species: Physocorypha politurata Karsch, by monotypy. 

politurata Karsch, 1896 : 345. i $ syntype, CAMEROUN: Victoria; i $ syntype, CAMEROUN: 
Mundame (ZMHU, Berlin). 

PLANGIA Stal 
Plangia Stal, 1873 : 40. Type-species: Phylloptera graminea Serville, by original designation. 

graminea Serville, 1839 : 405 (Phylloptera). Unknown number of syntypes of both sexes, 

SOUTH AFRICA: Cape of Good Hope (lost). 

natalensis Walker, 1869 : 379 (Phylloptera). Holotype $>, SOUTH AFRICA: Natal (BMNH, 
London) . 



94 D. R. RAGGE 

karschi Chopard, 1954 : 44- Holotype ?, GUINEA: Nimba, Nion (MNHN, Paris). 
taminifera Karsch, 1896 : 343. Holotype?, CAMEROUN: Lolodorf (ZMHU, Berlin). 
nebulosa Karsch, 18900 : 366. Holotype?, CAMEROUN: Barombi (ZMHU, Berlin). 
unimaculata Chopard, 1955 : 271. Holotype <$, SOUTH AFRICA: Cape Province, Tzitzikama 

Forest, Storms River Mouth (UZI, Lund). 
villiersi Chopard, 1954 : 45- Holotype?, GUINEA: Nimba (MNHN, Paris). 

PLANGIODES Chopard 

Plangiodes Chopard, in Chopard & Kevan, 1954 : 33- Type-species: Plangiodes carinatus 
Chopard, by original designation. 

carinatus Chopard, in Chopard & Kevan, 1954 : 33- Holotype $, KENYA: Garissa distr., 
Afwein (BMNH, London). 

PLANGIOLA Bolivar 

Plangiola Bolivar, 1906 : 336. Type-species: Plangiola herbacea Bolivar, by monotypy. 
herbacea Bolivar, 1906 : 337. 2 <$, i ? syntypes, CAMEROUN (IEE, Madrid). 

PLANGIOPSIS Karsch 
Plangiopsis Karsch, 1889 : 459. Type-species: Plangiopsis semiconchata Karsch, by monotypy. 

adeps Karsch, 1896 : 338. 2 ? syntypes, CAMEROUN: Lolodorf (ZMHU, Berlin). 
foraminata Karsch, 18920, : 324. Holotype <$, CAMEROUN: Barombi (ZMHU, Berlin). 
schoutedeni Griffini, 1908 : 218. Holotype?, CAMEROUN: Mukonje Farm (IRSNB, Brussels). 
semiconchata Karsch, 1889 : 460. Holotype?, CAMEROUN: Barombi (ZMHU, Berlin). 

POECILOGRAMMA Karsch 

Poecilogramma Karsch, 18876 : 52. Type-species: Poecilogramma striatifemur Karsch, by sub- 
sequent designation (Kir by, 1906 : 405). 

annulifemur Karsch, 18876 : 53. 2 <J, 3 ? syntypes, TANZANIA: Kawende (ZMHU, Berlin). 
striatifemur Karsch, 18876 : 53. 2 $ syntypes, KENYA: Mombasa (ZMHU, Berlin). 

POREUOMENA Brunner 
Poreuomena Brunner, 1878 : 187. Type-species: Poreuomena africana Brunner, by monotypy. 

africana Brunner, 1878 : 187. Holotype <J, GABON (MHN, Geneva). 

duponti Griffini, 19080, : 84. i <$, 6 ? syntypes, CAMEROUN: Mukonje Farm (IRSNB, Brussels). 

forcipata Sjostedt, 1901 : 12. Holotype <J, CAMEROUN (NR, Stockholm). 

gladiator Bolivar, 1906 : 337. Holotype g, CAMEROUN (IEE, Madrid). 

lamottei Chopard, 1954 : 4- I 6 A syntype, GUINEA: Nimba, N'Zo; i ? syntype, GUINEA: 

Nimba, Yalanzou (MNHN, Paris). 
wilverthi Griffini, 19080 : 85. Holotype <J, CONGO (KINSHASA): Umangi (IRSNB, Brussels). 

PREUSSIA Karsch 

Preussia Karsch, 18900 : 364. Type-species: Preussia lobatipes Karsch, by monotypy. 
lobatipes Karsch, 18900 : 365. Holotype ?, CAMEROUN: Barombi (ZMHU, Berlin). 



AFRICAN PHANEROPTERINAE 95 

PRONOMAPYGA Rehn 

Pronomapyga Rehn, 1914 : 171. Type-species: Pronomapyga grandis Rehn, by original desig- 
nation. 

grandisRehn, 1914 : 172. Holotype g, " OST-AFRIKA" (ZMHU, Berlin). 
graueri Rehn, 1914 : 171. Holotype $, "OST-AFRIKA" (ZMHU, Berlin). 

PROSPHAGA Ragge 

Prosphaga Ragge, ig6oa : 322. Type-species: Pantolepta calaharica Karny, by original desig- 
nation. 

calaharica Karny, 1910 : 52 (Pantolepta). Holotype $, SOUTH AFRICA: Bechuanaland, Kala- 
hari, Ku Gudie, nr. Pitsani (ZMHU, Berlin). 
splendens Ragge, 19600 : 324. Holotype $, ETHIOPIA: Wardere (BMNH, London). 

PSEUDOPYRRHIZIA Brunner 

Pseudopyrrhizia Brunner, 1891 : no. Type-species: Pseudopyrrhizia punctata Brunner, by 
monotypy. 

punctata Brunner, 1891 : no. Holotype $, "ZANZIBAR" (NM, Vienna). 

SCHUBOTZACRIS Rehn 

Schubotzacris Rehn, 1914 : 169. Type-species: Schubotzacris producta Rehn, by original desig- 
nation. 

producta Rehn, 1914 : 169. Holotype <$, CONGO (KINSHASA): Mboga (ZMHU, Berlin). 

STENAMBLYPHYLLUM Karsch 

Stenamblyphyllum Karsch, 1896 : 326. Type-species: Stenamblyphyllum dilutum Karsch, by 
monotypy. 

dilutum Karsch, 1896 : 327. Lectotype $, CAMEROUN: Victoria (ZMHU, Berlin); designated 
by Ragge (19620 : 309). 

SYMMETROPLEURA Brunner 

Symmetropleura Brunner, 1878 : 245. Type-species: Symmetropleura laevicauda Brunner, by 
subsequent designation (Kirby, 1906 : 446) (type-locality: South America, "Bahia"). 

Cameronia Karsch, 1889 : 450. Type-species: Symmetropleura africana Brunner, by monotypy. 
(See p. 77.) 

africana Brunner, 1878 : 246. Holotype <$, "CONGO" (NM, Vienna). 

plana Walker, 1869 : 339 (Phaneroptera) . Holotype <$, SOUTH AFRICA: Natal (BMNH, 
London) . 

TAPIENA Bolivar 

Tapeina Brunner, 1878 : 163. Type-species: Tapeina acutangula Brunner, by monotypy (type- 
locality : India). (Homonym of Tapeina Le Peletier & Serville, 1828 : 545.) 
Tapiena Bolivar, 1906 : 334 (proposed as replacement name for Tapeina Brunner). 

minor Bolivar, 1906 : 334. Holotype 9, GHANA: "Ashante" (IEE, Madrid). 



96 D. R. RAGGE 

TERPNISTRIA Stal 
Terpnistria Stal, 1873 : 42. Type-species: Phaneroptera zebrata Serville, by original designation. 

lobulata Stal, 1876 : 61. Holotype $, SOUTH WEST AFRICA: Ovambo (NR, Stockholm) (the 

additional citation of " Caffraria " is probably erroneous). 
tuberculata Chopard, in Chopard & Kevan, 1954 : 3 2 ^- Holotype <$, KENYA: Northern 

Frontier Distr., Garissa (BMNH, London). 
zebrata Serville, 1839 : 424 (Phaneroptera). Holotype $, SOUTH AFRICA: Cape of Good Hope 

(lost). 

TETRACONCHA Karsch 

Tetraconcha Karsch, 18900 : 61. Type-species: Tetraconcha fenestrata Karsch, by monotypy. 
Tellidia Bolivar, 1893 : 177. Type-species: Tellidia longipes Bolivar, by monotypy. 

banzyvilleana Griffini, 1909:11. Holotype?, CONGO (KINSHASA): Banzyville, Jusoro 

(MRAC, Tervuren). 

fenestrata Karsch, iSgoa : 62. Holotype $, CAMEROUN (ZMHU, Berlin). 
longipes Bolivar, 1893 : 178 (Tellidia}. Holotype?, IVORY COAST: Assinie (IEE, Madrid). 
smaragdina Brunner, 1891 : 116. Holotype <$, CAMEROUN (Lubeck, museum unknown). 
stichyrata Karsch, iSgoc : 360. Holotype $, CAMEROUN: Barombi (ZMHU, Berlin). 
scalaris Brunner, 1891 : 116. Holotype <$, GABON (NM, Vienna). 

TRIGONOCORYPHA Stal 

Trigonocorypha Stal, 1873 : 39. Type-species: Locusta crenulata Thunberg ( = Trigonocorypha 
unicolor (Stoll)), by original designation (type-locality unknown). 

tihamae Uvarov, 1952 : 177. Holotype <$, SAUDI ARABIA: near Qunfida, Wadi Qanuna 
(BMNH, London). 

TROPIDONOTACRIS Chopard 

Tropidonotacris Chopard, in Chopard & Kevan, 1954 : 3 21 - Type-species: Tropidonotacris 
carinata Chopard, by original designation. 

atnabilis Ragge, 1957 : I21 - Holotype <J, ETHIOPIA: Ual Ual (BMNH, London). 

carinata Chopard, in Chopard & Kevan, 1954 ' 3 22 ( as Tropidonotacris carinatus; corrected by 

Ragge, 1957 : 119). Holotype $, KENYA: Mandera distr., Takabba (BMNH, London). 
grandis Ragge, 1957 : 122. Holotype <$, TANZANIA: Little Mahenge (BMNH, London). 

TR OP ID OPHR YS Karsch 

Tropidophrys Karsch, 1896 : 340. Type-species: Tropidophrys amydra Karsch, by monotypy. 
amydra Karsch, 1896 : 341. Holotype <$, CAMEROUN: Victoria (ZMHU, Berlin). 

TYLOPSIS Fieber 

Centrophorus Fischer, 1846 : 361. Type-species: Centrophorus spinosus Fischer ( Tylopsis 
lilifolia (Fabricius)), by monotypy. (Homonym of Centrophorus Miiller & Henle, 1837 : 115.) 
Tylopsis Fieber, 1853 : 172. Type-species: Locusta lilifolia Fabricius, by monotypy. 



AFRICAN PHANEROPTERINAE 97 

ampla Ragge, 1964 : 315. Holotype <$, ANGOLA: Mexico distr., upper Mu-Simoj R. (BMNH, 

London) . 
bilineolata Serville, [1838]: 419 (Phaneroptera) . Holotype <$, SOUTH AFRICA: Cape of Good 

Hope (lost). 

attenuata Walker, 1869 : 338 (Phaneroptera). Holotype?, SOUTH AFRICA (BMNH, London). 
marginata Brunner, 1891 : 113. Holotype ?, SOUTH AFRICA: Natal, Durban (NM, Vienna). 
brevis Ragge, 1964:314. Holotype cJ, CONGO (KINSHASA): Volcan Nyamlagira (MRAC, 

Tervuren) . 
continua Walker, 1869 : 337 (Phaneroptera). Lectotype <$, SOUTH AFRICA: Natal, Durban 

(BMNH, London); designated by Ragge (1964 : 318). 
vicaria Walker, 1869 : 338 (Phaneroptera). Holotype <$, SOUTH AFRICA: Natal, Durban 

(BMNH, London). 

longipennis Stal, 1876 : 58. Holotype <J, SOUTH WEST AFRICA: Damara (NR, Stockholm). 
vittata Brunner, 1878 : 229. Lectotype $, SOUTH AFRICA: Natal, Durban (NM, Vienna); 

designated by Ragge (1964 : 318). 

inhamata Karsch, 1889 : 453. Lectotype <J, MOZAMBIQUE: Delagoa Bay (ZMHU, Berlin). 
dispar Sjostedt, 1909 : 135. Lectotype <, TANZANIA: Mt. Meru (NR, Stockholm); designated 

by Ragge (1964 : 309). 
dubia Giglio-Tos, 1907 : 3. i 6*. T nymphal syntype, UGANDA: Ibanda; i ? syntype, UGANDA: 

Mitiana (IMZU, Turin). 

fissa Ragge, 1964 : 312. Holotype <$, UGANDA: Entebbe (BMNH, London). 
gracilis Chopard, in Chopard & Kevan, 1954 ' 3 2 ^- Holotype <$, KENYA: Mandera distr., 

Damassa (BMNH, London). 

irregularis Karsch, 1893 : 130. Lectotype <$, TOGO: Bismarckburg (ZMHU, Berlin); desig- 
nated by Ragge (1964 : 307). 

perpulchra Burr, 1900 : 43. Holotype <$, SOMALI REPUBLIC: Whardi Datal (UM, Oxford). 
obscuripes Chopard, 1945 : 166. Lectotype <, CAMEROUN: Bambouto Mtns. (MNHN, Paris); 

designated by Ragge (1964 : 307). 
villiersi Chopard, 1950 : 133. Lectotype <$, NIGER: Agades (MNHN, Paris); designated by 

Ragge (1964 : 307). 
lamottei Chopard, 1954 : 4 2 - Lectotype o*. GUINEA: Nimba, Serengbara (MNHN, Paris); 

designated by Ragge (1964 : 307). 

lilifolia Fabricius, 1793 : 36 (Locusta). Type-locality: ITALY; type-material lost. 
gracilis Germar, 1817 : 251 (Locusta). Holotype?, YUGOSLAVIA: Zadar (lost). 
liliifolia Rambur, 1838 : 44 (Locusta). (Unjustified emendation.) 
praeusta Fischer, 1846 : 142 (Phaneroptera). Holotype?, U.S.S.R. : Azerbaijan, Kirovabad 

(lost). 
spinosus Fischer, 1846 : 362 (Centrophorus). Unknown number of nymphal syntypes of both 

sexes, U.S.S.R.: Crimea (lost). 
margineguttata Serville, 1839 : 442 (Phaneroptera). i syntype, SARDINIA; i $ syntype, 

SICILY (lost). 

? coi Jannone, 1936 : 147. Holotype <$, DODECANESE: Kos (ILEA, Portici). 
rubrescens Kirby, 1900 : 216. Holotype <J, MALAWI: Zomba (BMNH, London). 

punctulata Kirby, 1900 : 216. Lectotype <$, MALAWI: Zomba (BMNH, London); designated 

by Ragge (1964 : 320). 
meruensis Sjostedt, 1909 : 134. Lectotype $, TANZANIA: Mt. Meru (NR, Stockholm); 

designated by Ragge (1964 : 320). 
confluens Karny, 1915 : 124. Holotype?, GUINEA: Upper Niger (NM, Vienna). 

VOSSIA Brunner 

Vossia Brunner, 1891 : 139. Type-species: Vossia obesa Brunner, by monotypy. 
obesa Brunner, 1891 : 140. Holotype?, CAMEROUN (NM, Vienna). 



9 8 D. R. RAGGE 

WEISSENBORNIA Karsch 

Weissenbornia Karsch, 1888 : 65. Type-species: Weissenbornia praestantissima Karsch, by 
monotypy. 

praestantissima Karsch, 1888 : 66. Holotype <$, CAMEROUN: Lowry-Criby-Mundung (ZMHU, 
Berlin). 

ZEUNERIA Karsch 
Zeuneria Karsch, 1889 : 443. Type-species: Zeuneria melanopeza Karsch, by monotypy. 

biramosa Sjostedt, 1929 : 40. Holotype $, CONGO (KINSHASA): Bas-Uele (MRAC, Tervuren). 
centralis Rehn, 1914 : 179. Holotype <$, CONGO (KINSHASA): Ruwenzori (ZMHU, Berlin). 
longicercus Sjostedt, 1929 : 39. Holotype $, CONGO (KINSHASA): Inkisi (MRAC, Tervuren). 
melanopeza Karsch, 1889 : 443. Holotype ., CAMEROUN: Barombi (ZMHU, Berlin). 

REFERENCES 

ANDER, K. 1939. Vergleichend-anatomische und phylogenetische Studien iiber die Ensifera 

(Saltatoria) . Opusc. ent. Suppl. II. viii -f 306 pp., 172 figs. 
BEI-BIENKO, G. YA. 1954. Orthoptera 2 (2). Phaneropterinae. [In Russian.] Fauna 

SSSR 59, 384 pp., 210 figs. 

1964. Key to the insects of the European part of the U.S.S.R. 1. Lower Palaeoptera, with 
imperfect metamorphosis. [In Russian.] 936 pp., 344 figs. Moscow. 

BEIER, M. 1955. Bronn's Kl. Ordn. Tierreichs. Funfter Band: Arthropoda. III. Abteilung: 
Insecta. 6. Buck: Embioidea und Orthopteroidea. i + 304 pp., 197 figs. Leipzig. 

BLATCHLEY, W. S. 1920. Orthoptera of northeastern America with especial reference to the 
faunas of Indiana and Florida. 784 pp., 246 figs., i pi. Indianapolis. 

BoLfvAR, I. 1890. Ortopteros de Africa del Museo de Lisboa (conclusion). Jorn. Sci. math, 
phys. nat. (2) 1 : 211-232, i pi. 

1893. Voyage de M. Ch. Alluaud dans le territoire d'Assinie (Afrique occidentale) en 
juillet et aout 1886. 146 Memoire. Orthopteres. Annls Soc. ent. Fr. 62 : 169-184, i pi. 

1900. Les Orthopteres de St- Joseph's College a Trichinopoly (Sud de 1'Inde). Pt. II. 

Annls Soc. ent. Fr. 68 : 761-812, 2 pis. 
1906. Fasgonuridos de la Guinea espanola. Mems R. Soc. esp. Hist. nat. 1 : 327-377, 

i pi. 
1908. Algunos ort6pteros nuevos de Espafia, Marruecos y Canarias. Boln R. Soc. esp. 

Hist. nat. 8 : 317-334. 
1914. Dermapteros y ortopteros de Marruecos. Mems R. Soc. esp. Hist. nat. 8 : 157-238. 

1922. Voyage de M. le Baron Maurice de Rothschild en Ethiopie et en Afrique Orientale 
Anglaise (1904-1905). Resultats scientifiques. Animaux articules. Premiere Partie. Orthop- 
teres. Pp. 169-219, 4 pis. Paris. 

BORMANS, A. DE. i88o. Spedizione Italiana nell'Africa equatoriale. Risultati zoologici. 

Ortotteri. Annali Mus. civ. Star. nat. Genova 16 : 205-221, 4 figs. 
BRUES, C. T., MELANDER, A. L., & CARPENTER, F. M. 1954. Classification of insects. Bull. 

Mus. comp. Zool. Harv. 108, v + 917 PP-, 1219 figs. 
BRUNER, L. 1920. Orthoptera from Africa, being a report upon some Saltatoria mainly 

from Cameroon contained in the Carnegie Museum. Ann. Carneg. Mus. 13 : 92-142. 
BRUNNER VON WATTENWYL, C. 1878. Monographic der Phaneropteriden. 401 pp., 8 pis. 

Wien. 
1883. Ueber hypertelische Nachahmungen bei Orthopteren. Verh. zool.-bot. Ges. Wien 

33 : 247-249, i pi. 



AFRICAN PHANEROPTERINAE 99 

BRUNNER VON WATTENWYL, C. 1891. Additamenta zur Monographic der Phaneropteriden. 

Verh. zool.-bot. Ges. Wien 41 : 1-196, 2 pis. 
1893. Revision du systeme des Orthopteres et description des especes rapportees par 

M. Leonardo Fea de Birmanie. Annali Mus. civ. Stor. nat. Genova (2) 13 : 5-230, 6 pis. 
BURMEISTER, H. 1834-1839. Handbuch der Entomologie 2, xii + 1050 pp. Berlin. 
BURR, M. 1900. On a collection of insects and Arachnids made in 1895 and 1897, by Mi. 

C. V. A. Peel, F.Z.S., in Somaliland, with descriptions of new species. Pt. 7. Orthoptera. 

Proc. zool. Soc. Lond. 1900 : 35-46, i pi. 

1910. A synopsis of the Orthoptera of western Europe. 160 pp. London. 

CARL, J. 1921. Phasgonurides nouveaux du Museum de Geneve. Rev. suisse Zool. 28 : 301- 

309, 5 figs. 
CAUDELL, A. N. 1921. On the orthopterous group Phaneropterae (= Scudderiae), with 

descriptions of a new genus and species. /. Wash. Acad. Sci. 11 : 487-493, i fig. 
CHOPARD, L. 1922. Orthopteres et Dermapteres. Faune Fr. 3, 22 pp., 466 figs. 

1935. Contributions a 1'etude de la faune du Mozambique. Voyage de M. P. Lesne 
(1928-1929). 19. Orthopteres Ensiferes. Mems Estud. Mus. zool. Univ. Coimbra (I) 
85 : 1-32, 18 figs. 

1938. Mission scientifique de 1'Omo. Tome IV. Fasc. 33. Orthoptera. i. Dictyop- 

tera, Phasmodea, Ensifera. Mem. Mus. natn. Hist, nat., Paris 8 : 89-134, 34 figs. 

1943. Orthopteroides de 1'Afrique du Nord. Faune Emp. fr. 1, 450 pp., 658 figs. 

1945. Orthopteroides recueillis dans les montagnes du Cameroun par la mission Lepesme, 
Paulian, Villiers. Revue fr. Ent. 11 : 156-178, 27 figs. 

1947. Atlas des Apterygotes et Orthopteroides de France, in pp., 12 figs., 12 pis. Paris. 

- 1949. In Grass, P. -P., [Ed.], Traite de Zoologie 9, 1117 pp., 752 figs., 2 pis. Paris. 

- 1950. Contribution a 1'etude de 1'Air (Mission L. Chopard et A. Villiers). Orthopteroides. 
Mem. Inst.fr. Afr. noire 10 : 127-145, 2 figs. 

1954- La reserve naturelle integrate du Mont Nimba. Fasc. II. Pt. III. Orthopteres 

Ensiferes. Mem. Inst. fr. Afr. noire 40 (2) : 25-97, 42 figs. 

1955. In Hanstrom, B., Brinck, P. & Rudebeck, G., [Edd.], South African Animal Life 
2, pp. 266-300 (Orthoptera Ensifera), 26 figs. Stockholm. 

- 1958. Mission du Museum dans les lies du Golfe de Guinee. Entomologie, VI. Orthop- 
teroides. Bull. Soc. ent. Fr. 63 : 73-85, 2 figs. 

CHOPARD, L., & KEVAN, D. K. McE. 1954. Orthoptera-Ensifera from Northern Kenya and 

Jubaland. Trans. R. ent. Soc. Lond. 105 : 315-353, 22 figs. 
DIRSH, V. M. 1961. A preliminary revision of the families and subfamilies of Acridoidea 

(Orthoptera, Insecta). Bull. Br. Mus. nat. Hist. (Ent.) 10 : 351-419, 34 figs. 
EBNER, R. 1915. Neue tropische Odonturen (Orthoptera). Zool. Anz. 45 : 419-422. 

- 1943. Einige Orthoptera Saltatoria von Fernando Poo (Spanisch-Guinea) . 28. Beitrag 
zu den wissenshaftlichen Ergebnissen der Westafrika-Expedition Eidmann 1939/40. 
Zool. Anz. 143 : 259-274, 12 figs. 

ENDERLEIN, G. 1907. Pardalota karschiana, eine neue ost-afrikanische Orthoptere. Zool. 
Jb. (Syst. Geogr. Biol.) 25 : 197-200, i fig., i pi. 

ESSIG, E. O. 1947. College entomology, vii + 900 pp., 308 figs. New York. 

FABRICIUS, J. C. 1793. Entomologia systematica emendata et aucta 2, viii + 519 pp. Copen- 
hagen. 

FIEBER, F. X. 1853. Synopsis der europaischen Orthoptera [fourth part]. Lotos 3 : 168-176. 

FINOT, A. 1893. Seance du 25 Janvier, 1893. Annls Soc. ent. Fr. 62 : xviii-xli. 

FISCHER DE WALDHEIM, G. 1846. Orthoptera ImperiiRossici. iii + 443 pp., 37 P ls - Moscow. 

FOERSTER, A. 1868. Synopsis der Familien und Gattungen der Ichneumonen. Verh. 
naturh. Ver. preuss. Rheinl. 25 : 135-221. 

GERMAR, E. F. 1817. Reise nach Dalmatien und in das Gebiet von Ragusa. xii + 323 pp., 
ii pis. Leipzig & Altenburg. 

GERSTAECKER, A. 1869. Beitrag zur Insekten-Fauna von Zanzibar. II. Orthoptera et 
Neuroptera. Arch. Naturgesch. 35 (i) : 201-223. 



ioo D. R. RAGGE 

GiGLio-Tos, E. 1907. Spedizione al Ruwenzori di S. A. R. Luigi Amedeo di Savoia Duca 
degli Abruzzi. XVI. Ortotteri nuovi (diagnosi preventive). Boll. Musei Zool. Anat. 
comp. R. Univ. Torino 22 (556) : 1-3. 

GRANT, H. J. 1964. A revision of the genera Ceraia and Euceraia, with notes on their re- 
lationship to Scudderia (Orthoptera; Tettigoniidae ; Phaneropterinae) . Proc. Acad. nat. 
Sci. Philad. 116 : 29-117, 228 figs. 

GRIFFINI, A. 1906. Ortotteri raccolti da Leonardo Fea nell'Africa occidentale. I. Hetro- 
didi, Conocephalidi, Meconemidi, Pseudophyllidi, Mecopodidi e Fanerotteridi. Annali 
Mus. civ. Stor. nat. Genova (3) 2 : 358-397. 

I9o8#. Phasgonouridae Africane del R. Museo di Storia Naturale di Bruxelles. 5. 
Phaneropteridae i. Mem. Soc. ent. Belg. 15 : 74-86. 

19086. Phasgonouridae Africane del R. Museo di Storia Naturale in Bruxelles. 6. 
Phaneropteridae 2. Mem. Soc. ent. Belg. 15 : 201-226. 

1909. Note sopra alcune Phasgonouridae del Congo. Annls Soc. ent. Belg. 53 : 9-28, 

i fig. 

HARZ, K. 1957. D Geradfliigler Mitteleuropas. xxiii + 494 pp., 254 figs., 20 pis. Jena. 
1960. Geradfliigler oder Orthoptera (Blattodea, Mantodea, Saltatoria, Dermaptera) . 

Tierwelt Dtl. 46, xii + 232 pp., 566 figs. 
IMMS, A. D. 1957. A general textbook of entomology . gth edn, revised by O. W. Richards and 

R. G. Davies. x + 886 pp., 606 figs. London. 
JACOB Y, M. 1893. Descriptions of some new species of Eumolpidae and Halticidae from 

Africa (Gaboon). Entomologist 26 (Suppl.): 97-102. 
JANNONE, G. 1936. Nuovi contributi alia conoscenza della fauna delle isole italiane dell'Egeo. 

Pt. 5. Studio bio-ecologico e sistematico dell'Ortotterofauna con notizie sui Blattoidei, 

Mantoidei e Fasmoidei. Boll. Lab. Zool. gen. agr. R. Scuola Agric. Portici 29 : 47-248, 

48 figs., i pi. 
KARABAG, T. 1964. Some new species and new records of Tettigoniidae (Orthoptera) from 

Turkey. Communs Fac. Sci. Univ. Ankara (C) 13 : 36-55, 40 figs. 
KARNY, H. H. 1910. Zoologische und anthropologische Ergebnisse einer Forschungsreise im 

westlichen und zentralen Sudafrika ausgefuhrt in den Jahren 1903-1905 mit unterstutzung 

der Kgl. Preussischen Akademie der Wissenschaften zu Berlin von Dr. Leonhard Schultze. 

4. Systematik und Tiergeographie. Orthoptera (s. str.). Denkschr. med.-naturw. Ges. 

Jena 16 : 35-90, i pi. 
1915. Ergebnisse der Forschungsreise des Herrn Dr. Adalbert Klaptocz nach Franzosisch 

Guinea. Orthoptera und Oothecaria. Zool. Jb. (Syst. Geogr. Biol.) 40 : 119-146. 
1926. On Malaysian katydids (Tettigoniidae). Represented in the collections of the 

F.M.S. Museum (Kuala Lumpur) and the Raffles Museum (Singapore). /. fed. Malay St. 

Mus. 13 : 69-156, 48 figs., 2 pis. 
1927. Revision der Gryllacriden des Zoologischen Institutes in Halle a. S., sowie einiger 

Tettigoniiden-Typen von Burmeister und Giebel. Z. Naturw. 88 : 1-14. 
KARSCH, F. 1887^. Verzeichniss der von Herrn Waldemar Belck 1885 im Damaralande gesam- 

melten Orthopteren. Ent. Nachr. 13 : 39-46. 

18876. Zwei neue ostafrikanische Phaneropteriden. Ent. Nachr. 13 : 52-54. 

1888. Weissenbornia, eine neue Orthopteren- (Phaneropteriden-) Gattung aus Deutsch- 

Westafrika. Ent. Nachr. 14 : 65-67. 

1889. Orthopterologische Beitrage III. Berl. ent. Z. 32 : 415-464, i pi. 

iSgoa. Orthopterologische Mittheilungen 4. Ueber Phaneropteriden. Ent. Nachr. 

16 : 57-62. 
18906. Neue westafrikanische, durch Herrn Premierlieutenant Morgen von Kribi 

eingesendete Orthopteren. Ent. Nachr. 16 : 257-276, 4 figs. 
iSgoc. Verzeichniss der von Herrn Dr. Paul Preuss auf der Barombi-Station in Deutsch- 

Westafrika 1890 gesammelten Locustodeen aus den Familien der Phaneropteriden, Meko- 

nemiden und Gryllakriden. Ent. Nachr. 16 : 353-369, 4 figs. 



AFRICAN PHANEROPTERINAE 101 

KARSCH, F. 18920. Uebersicht der von Herrn Dr. Paul Preuss auf der Barombi-Station in 
Kamerun gesammelten Locustodeen. Berl. ent. Z. 36 : 317-346, 7 figs. 

18926. Verzeichniss der von Herrn Dr. Paul Preuss im Kamerungebirge erbeuteten 

Orthopteren. Berl. ent, Z. 37 : 65-78, 3 figs. 

- 1893. Die Insecten der Berglandschaft Adeli im Hinterlande von Togo (Westafrika) 
nach dem von den Herren Hauptmann Eugen Kling (1888 und 1889) und Dr. Richard 
Biittner (1890 und 1891) gesammelten Materiale. i. Apterygota, Odonata, Orthoptera 
Saltatoria, Lepidoptera Rhopalocera. Berl. ent. Z. 38 : 1-266, 35 figs., 6 pis. i map. 

1896. Neue Orthopteren aus dem tropischen Afrika. Stettin, ent. Ztg 57 : 242-359, 
44 fi gs. 

KIRBY, W. F. 1900. Notes on the collection of African Phasgonuridae formed by Mr. W. L. 
Distant in the Transvaal etc., with descriptions of two new species. Ann. Mag. nat. Hist. 
(7) 6 : 211-217. 

1906. A synonymic catalogue of Orthoptera 2, Pt. i, viii + 562 pp. London. 

1909. Ruwenzori Expedition reports. 8. Orthoptera. Trans, zool. Soc. Lond. 19 : 63- 

66. 
KRAUSS, H. 1890. Beitrag zur Kenntniss westafrikanischer Orthopteren. 2. Orthopteren 

der Guinea-Inseln Sao Thome und Rolas, gesammelt von Prof. Dr. Richard Greeff. Zool. 

Jb. (Syst. Geogr. Biol.) 5 : 647-668, i pi. 

1893. Vorlaufige Diagnosen der neuen Arten und Varietaten von Oran. Jh. Ver. vaterl. 
Naturk. Wurtt. 48 : xcvi. 

- 1901. Beitrag zur Kenntniss der Orthopteren Deutsch-Sudwestafrikas. Verh. zool.-bot. 
Ges. Wien 51 : 281-293. 

LA BAUME, W. 1911. Beitrag zur Kenntniss der aethiopischen Orthopteren. Stettin, ent. 

Ztg 72 : 308-326, 19 figs. 
LE PELETIER DE SAINT-FARGEAU, A. L. M., & SERVILLE, J. G. A. 1828. Encyclopedie metho- 

dique. Histoire naturelle. Entomologie, ou histoire naturelle des crustaces, des arachnides 

et des insectes 10, 833 pp. Paris. 
MATSUMURA, S., & SHIRAKI, T. 1908. Locustiden Japans. /. Coll. Agric. imp. Univ. Tokyo 

3 : 1-80, 2 pis. 
MORALES AGACINO, E. 1947. Notas sobre ortopteroides de Ifni y Sahara Espanol. Eos, 

Madr. 23 : 241-283, 4 figs. 

1950. Apuntes sobre los Phasmidae y Tettigoniidae marroquies del Institute Espanol de 
Entomologia. Eos, Madr. 26 : 157-196, 17 figs. 

MULLER, J., & HENLE, J. 1837. Sitzung der physicalisch-mathematischen Klasse (31. Juli). 

Ber. Verh. Akad. Wiss. Berl. 1837 : iii-n8. 
RAGGE, D. R. 19560. A revision of the genera Phaneroptera Serville and Nephoptera Uvarov 

(Orthoptera : Tettigoniidae), with conclusions of zoogeographical and evolutionary 

interest. Proc. zool. Soc. Lond. 127 : 205-283, 131 figs. 

19566. A revision of the genus Oxyecous Chopard, 1935 (Orthoptera : Tettigoniidae). 
Proc. R. ent. Soc. Lond. (B) 25 : 183-190, 23 figs. 

1957- A revision of the genus Tropidonotacris Chopard, 1954 (Orthoptera : Tettigoniidae). 

Proc. R. ent. Soc. Lond. (B) 26 : 119-122, 7 figs. 

19600. The Acrometopae of the Ethiopian Region: a revision, with notes on the sexual 
dimorphism shown by the group (Orthoptera : Tettigoniidae). Bull. Br. Mus. nat. Hist. 
(Ent.) 8 : 269-333. 148 figs. 

19606. Further notes on the genus Phaneroptera Serville, with a revised key to the species 
(Orthoptera : Tettigoniidae). Proc. zool. Soc. Lond. 134 : 237-250, 56 figs. 

- i96ia. A revision of the genus Ducetia Stal (Orthoptera : Tettigoniidae). Bull. Br. Mus. 
nat. Hist. (Ent.) 10 : 171-208, 91 figs. 

- 19616. Further notes on the genus Lamecosoma Ragge, with a description of a new species 
(Orth. Tettigoniidae). Eos, Madr. 37 : 215-219, 5 figs. 

19620. A revision of the genera Drepanophyllum Karsch and Stenamblyphyllum Karsch 
(Orth. Tettigoniidae). Eos, Madr. 38 : 299-309, 21 figs. 



102 D. R. RAGGE 

RAGGE, D. R. 19626. A revision of the genera Phlaurocentrum Karsch, Buettneria Karsch 

and Leiodontocercus Chopard (Orthoptera : Tettigoniidae) . Bull. Br. Mus. nat. Hist. (Ent.) 

13 : 3-17, 32 figs. 
1964. A revision of the genus Tylopsis Fieber (Orthoptera : Tettigoniidae). Butt. Br. 

Mus. nat. Hist. (Ent.) 15 : 297-322, 52 figs. 

RAMBUR, M. P. 1838-1842. Faune entomologique de I'Andalousie 2, 336 pp., 18 pis. Paris. 
RAMME, W. 1951. Zur Systematik Faunistik und Biologic der Orthopteren von Siidost- 

Europa und Vorderasien. Mitt. zool. Mus. Berl. 27 : 3-431, 134 figs., 39 pis., 3 maps. 
REHN, J. A. G. 1914. Wissenschaftliche Ergebnisse der Deutschen Zentral-Afrika-Expedition 

1907-1908 unter Fiihrung Adolf Friedrichs, Herzogs zu Mecklenburg. Band V. Lief. i. 

Orthoptera I. Mantidae, Phasmidae, Acrididae, Tettigoniidae und Gryllidae aus dem 

Zentral-Afrikanischenseengebiet, Uganda und dem Ituri-Becken des Kongo. Pp. 1-223. 

Leipzig. 
SAUSSURE, H. DE, & PICTET, A. 1897-1899. Fam. Locustidae. Biologia cent. -am. (Zool.) 

Insecta. Orthoptera 1 : 285-458, pis. 14-22. 
SCHAUM, H. 1853. Gesammtsitzung der Akademie (15 December). Ber. Verh. Akad. Wiss. 

Berl. 1853 : 747-780. 
SCHULTHESS SCHINDLER, A. DE. 1898. Orthopteres du pays des Somalis, recueillis par L. 

Robecchi-Brichetti en 1891 et par le Prince E. Ruspoli en 1892-93. Annali Mus. civ. 

Stor. nat. Genova (2) 19 : 161-216, i pi. 
SERVILLE, J. G. A. 1831. Revue methodique des insectes de 1'ordre des Orthopteres [second 

part]. Annls Sci. nat. 22 : 134-166. 

1838*. Histoire naturelle des insectes. Orthopteres. xviii + 776 pp., 14 pis. Paris. 

SJOSTEDT, Y. 1901. Beitrage zur Kenntniss der Insektenfauna von Kamerun. 8. Locusto- 

deen aus Kamerun und Kongo. Bih. K. svenska VetenskAkad. Handl. 27 (4, 3) : 1-45, 

4 pis. 
1909. Wissenschaftliche Ergebnisse der schwedischen zoologischen Expedition nach dem 

Kilimandjaro, dem Meru und den umgebenden Massaisteppen deutsch-Ostafrikas 1905-1906 

unter Leitung von Prof. Dr. Yngve Sjostedt. 17. Orthoptera. 6. Locustodea. Pp. 125-148, 

i pi. Stockholm. 
1912. Zur Orthopterenfauna des Kamerungebirges. Ark. Zool. 7 (37) : 1-30, 4 figs., 

3 Pis. 
1913- Neue Orthopteren aus Ost- und Westafrika nebst einigen anderen zugehorigen 

Formen. Ark. Zool. 8 (6) : 1-26, 3 pis. 
1929- Voyage au Congo de S.A.R. le Prince Leopold de Belgique 1925. Orthoptera. 

2. Phasgonurodea. Revue Zool. Bot. afr. 17 : 38-43, 3 figs. 
SPINOLA, M. 1839. Essai sur les Fulgorelles, sous-tribu de la tribu des Cicadaires, ordre des 

Rhyngotes [first part]. Annls Soc. ent. Fr. 8 : 133-337. 
STAL, C. 1856. Orthoptera cursoria och Locustina f ran Caff erlandet. 'Ofvers. K. VetenskAkad 

Forh. 13 : 165-170. 

1873. Orthoptera nova descripsit. Ofvers. K. VetenskAkad. Forh. 30 (4) : 39-53. 

1874. Recensio Orthopterorum. Revue critique des Orthopteres decrits par Linne, De 

Geer et Thunberg 2, 121 pp. Stockholm. 
1876. Bidrag till sodra Afrikas Orthopter-fauna. Ofvers. K. VetenskAkad. Forh. 33 

(3) : 29-76. 
STEINHEIL, E. 1873. Symbolae ad historian! Coleopterorum Argentiniae meridionalis, ossia 

enumerazione dei coleotteri raccolti dal prof. P. Strobel nell'Argentinia meridionale, e 

descrizione delle specie nuove. Centuria II. Atti Soc. ital. Sci. nat. 15 : 554-578. 
STEINMANN, H. 1966. New Phaneroptera Serv. and Nephoptera Uv. species (Orthoptera: 

Tettigoniidae). Acta zool. hung. 12 : 409-417, 29 figs. 
STRAND, E. 1921. Neue Namen dreier Insektengattungen. Ent. Z. Frank/, a. M. 34 : 106. 

* This work is dated " 1839 " and this is often cited as the date of publication. The work was, how- 
ever, published in December, 1838 (teste Sherborn, C. D., 1922, Index Animalium (2) 1 : xxxiii). 



AFRICAN PHANEROPTERINAE 



103 



THOMSON, J. 1857. Description de Cerambycides nouveaux ou peu connus de ma collection. 

Archs ent. Paris 1 : 291-320. 
UVAROV, B. P. 1924. Notes on the Orthoptera in the British Museum. 3. Some less known 

or new genera and species of the subfamilies Tettigoniinae and Decticinae. Trans, ent. 

Soc. Land. 1923 : 492-537, i fig., i pi. 

1934. Entomological expedition to Abyssinia, 1926-7 : Orthoptera of the families Mantidae 
Gryllidae, Tettigoniidae, and Acrididae. /. Linn. Soc. (Zool.) 38 : 591-614, 10 figs. 

1936. Studies in the Arabian Orthoptera. i. Descriptions of new genera, species and 
subspecies. /. Linn. Soc. (Zool.) 39 : 531-554, 5 figs., 2 pis. 

i94O. Twenty-eight new generic names in Orthoptera. Ann. Mag. nat. Hist, (n) 

5 : 173-176. 

19406. Twenty-four new generic names in Orthoptera. Ann. Mag. nat. Hist, (n) 

6 : 112-117. 

- 1952. Studies in the Arabian Orthoptera. III. New genera, species and subspecies 

collected by the Anti-Locust Missions. /. Linn. Soc. (Zool.) 42 : 176-194, 55 figs. 
UVAROV, B. P., & POPOV, G. 1957. The Saltatorial Orthoptera of Socotra. /. Linn. Soc. 

(Zool.) 43 : 359-389, 38 figs., i pi., i map. 
WALKER, F. 1869. Catalogue of the specimens of Dermaptera Saltatoria in the collection of the 

British Museum 2 : 225-423. London. 
1870. Catalogue of the specimens of Dermaptera Saltatoria in the collection of the British 

Museum 3 : 425-604. London. 
WERNER, F. 1929. Wissenschaftliche Ergebnisse einer zoologischen Forschungsreise nach 

Westalgerien und Marokko. II. Dermapteren und Orthopteren aus Westalgerien und 

Marokko. Sber. Akad. Wiss. Wien (I) 138 : 167-188, 7 figs., 4 pis. 

1932. Orthopteren aus Marokko und Westalgerien. Zool. Am. 100 : 113-127, 2 figs. 

ZACHER, F. 1917. Die Geradflugler Deutschlands und ihre Verbreitung. vii + 287 pp., i map. 

Jena. 
ZEUNER, F. E. 19360. The subfamilies of Tettigoniidae (Orthoptera). Proc. R. ent. Soc. 

Lond. (B) 5 : 103-109. 
19366. The Recent and fossil Tympanophorinae (Tettigoniidae, Saltatoria). Trans. R. 

ent. Soc. Lond. 85 : 287-302, 2 pis. 
1939. Fossil Orthoptera Ensifera. xiii + 321 pp., 80 pis. London. 



INDEX 



Valid names are in roman type, invalid names in italics. 



acaciae, 92 

accrana, 80 

acuticercus, 83 

adeps, 94 

adicra, 84 

aequatorialis, 84 

affinis, 87 

afra, 81 

africana, Phaneroptera, 92 

africana, Poreuomena, 94 

africana, Symmetropleura, 77, 78, 95 

albida, 92 

algerica, 90 

amabilis, 96 



ampla, 97 

amplectens, 82 

amplipennis, 87 

amydra, 96 

Anerota, 92 

angustipennis, Arantia, 80 

angustipennis, Debrona, 82 

angustipennis, Leiodontocercus, i 

angustipennis, Mangomaloba, 89 

Anisotochra, 80 

annulifemur, 94 

antinorii, 89 

apertus, 91 

apicalis, 81 



IO4 



INDEX 



arabica, 85 
Arantia, 80 
asymmetrica, 91 
Atlasacris, 80 
atrolineata, 80 
attenuata, 97 
Azamia, 80 

banzyvilleana, 96 

belcki, 82 

bicolor, 85 

bilineolata, 97 

biplagiata, 80 

bipunctata, 88 

biramosa, Ducetia, 83 

biramosa, Zeuneria, 98 

blandus, 90 

Bongeia, 81 

borrei, 90 

breviala, 84 

brevicauda, 81 

brevicollis, 85 

brevipes, 80 

brevis, Odontura, 91 

brevis, Tylopsis, 97 

brunneri, Conchotopoda, 78, 82 

brunneri, Melidia, 89 

Bueacola, 81 

Buettneria, 81 

Buttneria, 81 

calaharica, 95 

earner ata, 82 

Cameronia, 77, 78, 95 

Camptoxiphae, 79 

canaliculata, 85 

capensis, 91 

capreola, 81 

carinata, Tropidonotacris, 96 

carinatus, Plangiodes, 94 

Catoptropteryx, 81 

centralis, 98 

Centrophorus, 96 

cereris, 85 

cervina, 82 

Cestromoecha, 78, 8 1 

chelocerca, 83 

clavigera, 92 

cloetensi, 91 

coi, 97 

compressa, 85 

concava, 87 

Conchotopoda, 82 

condylus, 89 



confluens, 97 

contradti, 89 

conspersa, 93 

continua, 97 

cooperi, 92 

cornigera, 81 

corrosa, 83 

corrosifolium, 83 

Corycomima, 82 

Corymeta, 82 

costata, 84 

cothurnata, 85 

crassicauda, 82 

crassipes, 78, 81 

crosskeyi, Ducetia, 84 

crosskeyi, Horatosphaga, 87 

crypterius, 84 

cuspidata, 85 

Dannfeltia, 82 

Dapanera, 82 

darlingi, 85 

Debrona, 82 

denticulata, 83 

dilutum, 95 

dimidiata, 86 

diminuta, Eurycorypha, 85 

diminuta, Horatosphaga, 87 

Diogena, 83 

Dioncomena, 83 

dispar, 97 

Dithela, 83 

doriae, 80 

Drepanophyllum, 83 

dubia, 97 

Ducetia, 83 

duponti, 94 

eburneiguttata, 81 
eidmanni, 82 
elegans, 89 
elgonis, 87 

elongata, Horatosphaga, 87 
elongata, Phaneroptera, 92 
Enochletica, 84 
Epiphlebus, 84 
Euanerota, 92 
Eulioptera, 84 
Eupantolepta, 87 
Eurycorypha, 84 
Euthyphlebia, 86 
excelsior, 80 
extensipes, 81 



INDEX 



105 



fallax, 85 
fasciata, 80 
fatidica, 80 
fausta, 83 
feana, 86 
fenestrata, 96 
fissa, 97 
flavescens, 82 
foraminata, 94 
forcipata, 94 
fragilis, 92 
furcatum, 83 
fuscopunctata, 84 

Gabonella, 85 

Gabonia, 85 

gabunensis, 80 

galbana, 86 

Gelatopoea, 85 

Gelatopoia, 85 

genuteres, 82 

gestri, 80 

gladiator, 94 

Goetia, 86 

Gonatoxia, 86 

gracilipes, 80 

gracilis, Horatosphaga, 87 

gracilis, Locusta, 97 

gracilis, Phaneroptera, 92 

gracilis, Plegmatoptera, 87 

gracilis, Tylopsis, 97 

grallatoria, 82 

graminea, 93 

grandis, Pronomapyga, 95 

grandis, Tropidonotacris, 96 

graueri, 95 

Gravenreuthia, 86 

guttatipes, 81 

haasi, 91 
habetata, 82 
hamuligera, 90 
Harposcepa, 86 
herbacea, 94 
heteromorpha, 87 
Himerta, 86 
Himertula, 86 
hoehneli, 87 
Horatosphaga, 86 
hydatinoptera, 80 

immaculata, 86 
immaculipennis, 81 
inclusa, 87 



inconspicua, 93 
indica, 92 
inermis, 88 
inhamata, 97 
insularis, 93 

irregularis, Dapanera, 82 
irregularis, Tylopsis, 97 
Itokiia, 88 
Ivensia, 88 

Japygophana, 88 

kandti, 87 

karschi, Pachypyga, 87 

karschi, Plangia, 94 

Karschia, 83 

karschiana, Harposcepa, 86 

karschiana, Pardalota, 91 

kasindina, 88 

Keniacola, 87 

kenyensis, 89 

Kevaniella, 88 

kilimandjarica, Monticolaria, 90 

kilimandjarica, Pantolepta, 87 

Kuwayamaea, 83 

Lamecosoma, 88 

laminata, 89 

laminifera, 94 

lamottei, Poreuomena, 94 

lamottei, Tylopsis, 97 

latevittatum, 93 

latifolia, 80 

latifrons, 90 

latipennis, Catoptropteryx, 81 

latipennis, Mangomaloba, 89 

leggei, 87 

Leiodontocercus, 89 

lenzi, 93 

leptocerca, 82 

leptocnemis, 80 

leptomorpha, 84 

Leptophyes, 89 

lesnei, 91 

Letana, 89 

lilifolia, 97 

liliifolia, 97 

linearis, 87 

liouvillei, 91 

lobatipes, 94 

lobatum, 93 

lobulata, 96 

lobulipennis, 86 

longicerca, Eulioptera, 84 



io6 



INDEX 



longicercus, Zeuneria, 98 

longipennis, 97 

longipes, Horatosphaga, 87 

longipes, Tetraconcha, 96 

longispina, 92 

loosi, 84 

lurida, 92 

macrocerca, 84 

maculata, Gonatoxia, 86 

maculata, Peropyrrhicia, 92 

maculatum, Phlaurocentrum, 93 

maculiceps, 81 

maculipennis, 81 

maculosa, 92 

magna, Horatosphaga, 87 

magna, Phaneroptera, 92 

magnus, Oxyecous, 91 

malleus, 89 

mammisignum, 80 

manca, 80 

Mangomaloba, 89 

marginalis, 92 

marginata, 97 

margineguttata, 97 

marmorata, Arantia, 80 

marmoratum, Drepanophyllum, 83 

maroccana, 91 

massaiae, 92 

mecopodoides, 93 

media, Eurycorypha arabica, 85 

media, Horatosphaga, 87 

melanopeza, 98 

melanotus, 80 

melica, 90 

Melidia, 89 

meruensis, Horatosphaga, 87 

meruensis, Monticolaria, 90 

meruensis, Tylopsis, 97 

Meruterrana, 89 

microptera, 91 

Milititsa, 90 

Miltinobates, 90 

minima, 92 

minor, 95 

modulata, 90 

moghrebica, 91 

montana, 85 

Monteiroa, 90 

monticola, 89 

Monticolaria, 90 

montivaga, 87 

Morgenia, 90 

multivenosa, 88 



mundamensis, 81 
mutica, 85 
Myllocentrum, 90 
Myrmecophana, 84 

nana, Dannfeltia, 82 
nana, Phaneroptera, 92 
natalensis , 93 
nebulosa, 94 
neutralipennis, 81 
nigropunctata, 93 
nigrospinosa, 81 
nomima, 87 
nuda, 87 

obesa, 97 
obscuripes, 97 
Odontura, 90 
ornata, 83 
ornatipes, 85 
orthocnemis, 80 
ostentatrix, 84 
Otiaphysa, 82 
ovalipennis, 80 
Oxyecous, 91 

Pachypyga, 86 
Pachypygiana, 87 
Pantolepta, 86 
parallela, 87 
Paranerota, 92 
Parapyrrhicia, 91 
Pardalota, 91 
parva, Conchotopoda, 82 
parva, Ducetia, 84 
parva, Phaneroptera, 93 
Paura, 83 
peculiaris, 80 
peeli, 88 
peloti, 88 
Peronura, 92 
Peropyrrhicia, 92 
perpulchra, 97 
Phaneroptera, 92 
Phaneroptila, 93 
Phaula, 93 
Phaulula, 93 
Phlaurocentrum, 93 
Physocorypha, 93 
plana, 95 
Plangia, 93 
Plangiodes, 94 
Plangiola, 94 



INDEX 



107 



Plangiopsis, 94 
plasoni, 91 
Plegmatia, 87 
Plegmatoptera, 86 
Poecilogramma, 94 
politurata, 93 
Poreuomena, 94 
praestantissima, 98 
praeusta, 97 
Preussia, 94 
producta, 95 
Pronomapyga, 95 
proserpinae, 85 
Prosphaga, 95 
Pseudisotima, 83 
Pseudopyrrhizia, 95 
pulchra, 91 
punctata, Ducetia, 84 
punctata, Pseudopyrrhizia, 95 
puncticollis, 81 
punctipennis, Ducetia, 84 
punctipennis, Eurycorypha, 85 
punctulata, Catoptropteryx, 81 
punctulata, Phaneroptera, 93 
punctulata, Tylopsis, 97 
Pyrrhicia, 89 
Pyrrhizia, 89 



quadridentata, 91 
quadripunctata, Phaneroptera, 92 
quadripunctata, Telaea, 84 



ramulosa, 84 

rectifolia, 80 

rectiloba, 83 

reducta, 85 

regina, 80 

regularis, 87 

reimeri, 91 

reticulata, Eulioptera, 84 

reticulata, Plegmatoptera, 87 

reticulosa, 84 

retinervis, 80 

Rhegmatopoda, 78, 82 

rivae, 88 

robusta, 87 

roseata, 92 

royi, 89 

rubrescens, 97 

rubricornis, 90 

ruspolii, Epiphlebus, 84 

ruspolii, Horatosphaga, 88 



sagitta, 84 
samburu, 88 
saturata, 86 
scalaris, 96 
schoutedeni, 94 
Schubotzacris, 95 
Schulthessinia, 87 
scotti, 92 
scurra, 80 
securifera, 85 
semiconchata, 94 
serrifera, 88 
signatipennis, 81 
silvarum, 88 
simillima, 85 
simplicinervis, 80 
smaragdina, 96 
somali, 88 
somaliensis, 90 
sparsa, 92 
spathulifera, 90 
spinosus, 97 
spinulicauda, 91 
spinulosa, Arantia, 80 
spinulosa, Eulioptera, 84 
spinulosa, Eurycorypha, 85 
splendens, 95 
Stenacropteryx, 81 
Stenamblyphyllum, 95 
stenoxypha, 91 
stichyrata, 96 
stigmosum, 90 
strangulata, 85 
striatifemur, 94 
stuhlmanni, 88 
stylata, 85 
stylifera, 88 
subcarinata, 92 
sudanensis, 85 
superba, Dioncomena, 83 
superba, Pardalota, 91 
Symmetropleura, 77, 78, 95 

Tapeina, 95 
Tapiena, 95 
teknicus, 91 
Telaea, 83 
Tellidea, 96 
tenuicerca, 93 
tenuipes, 81 
tenuis, 88 
terniensis, 91 
Terpnistria, 96 
Tetraconcha, 96 



io8 



INDEX 



tetrasticta, 92 
Thaumatoxenia, 87 
tigrina, 80 
tihamae, 96 
Trigonocorypha, 96 
trochlearis, 88 
Tropidonotacris, 96 
Tropidophrys, 96 
tuberculata, 96 
tuberosum, 93 
turbatum, 93 
Tylopsis, 96 

ugandana, 80 
uncinata, 88 
undulatus, 91 
unimaculata, 94 
uvarovi, 91 

varia, 85 
velicauda, 85 



versicolor, 91 
vicaria, 97 
vicina, 88 

villiersi, Plangia, 94 
villiersi, Tylopsis, 97 
viridis, 88 
vitriala, 84 
vittata, 97 
vosseleri, 88 
Vossia, 97 



Weissenbornia, 98 
wilverthi, 94 



zanzibarica, Letana, 89 
zanzibarica, Parapyrrhicia, 91 
zebrata, Eurycorypha, 85 
zebrata, Terpnistria, 96 
Zeuneria, 98 




A LIST OF SUPPLEMENTS 
TO THE ENTOMOLOGICAL SERIES 

OF THE BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 



1. MASNER, L. The types of Proctotrupoidea (Hymenoptera) in the British 
Museum (Natural History) and in the Hope Department of Entomology, Oxford. 
Pp. 143. February, 1965. 5. 

2. NIXON, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera: 
Braconidae). Pp. 284; 348 Text-figures. August, 1965. 6. 

3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera) . Pp. 177; 
18 plates, 270 Text-figures. August, 1965. 4 45. 

4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172; 500 Text-figures. October, 

1965- 355. 

5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156; 
475 Text-figures. November, 1965. 2 155. 

6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae & Drosophilidae. 
Pp. 129; 328 Text-figures. 3. 

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

33s. 

8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the 
world species of Cleora (Lepidoptera : Geometridae) . Pp. 119; 14 plates, 146 
Text-figures, 9 maps. February, 1967. 3 los. 

9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera : Rhopalocera) . Pp. 509. August, 1967. 8 los. 

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rhopa- 
locera). Pp. 322; Coloured frontispiece, 348 text figures. August, 1967. 8. 

11. MOUND, L. A. A review of R. S. Bagnall's Thysanoptera Collection. Pp. 181; 
82 Text-figures. May, 1968. 4. 



PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED, B ARTH OLOME W PRE SS, DO RKING 



HYMENOPTERA FROM TURKEY 

SYMPHYTA 




R. B. BENSON 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 22 No. 4 

LONDON: 1968 



HYMENOPTERA FROM TURKEY 
SYMPHYTA 




BY 

R. B. BENSON 



British Museum (Natural History)* 



Pp. 109-207; 42 Text-figs. 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 22 No. 4 

LONDON: 1968 



THE BULLETIN OF THE BRITISH MUSEUM 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

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

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

This paper is Vol. 22, No. 4 of the Entomological 
series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



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



Trustees of the British Museum (Natural History) 1968 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 5 July, 1968 Price 2 



HYMENOPTERA FROM TURKEY 
SYMPHYTA 

By R. B. BENSON* 

SYNOPSIS 

The Sawflies of Asiatic Turkey and neighbouring countries (from Israel and Cyprus to Trans- 
caucasia and Iran) are listed; over 370 from Asiatic Turkey itself, 50 of these for the first time. 
Twenty-three new species or subspecies are described and lectotypes designated for seven 
species. Keys to world species are given for genera or generic-groups centred in this region. 
The account is based mainly on the collections made in Turkey by Messrs. K. M. Guichard and 
D. H. Harvey in 1959, 1960 and 1962. 

CONTENTS 

Page 
INTRODUCTION . . . . . . . . . . . m 

XYELIDAE .......... 112 

PAMPHILIIDAE . . . . . . . . . . i I2 

MEGALODONTIDAE . . . . . . . . . . n^ 

XlPHYDRIIDAE . . . . . . . . . . Il6 

SlRICIDAE ......... Il6 



ORUSSIDAE 
CEPHIDAE 
ARGIDAE . 



117 
117 
124 



ClMBICIDAE ........... 129 

TENTHREDINIDAE .......... 133 

Selandriinae . .... ..... 134 

Blennocampinae ......... 143 

Tenthredininae .......... 154 

Nematinae .......... 198 

REFERENCES ........... 205 

INDEX ............ 206 

INTRODUCTION 

A GENERAL account of the fruitful entomological expeditions of Kenneth M. Guichard 
and David H. Harvey to Turkey in 1959, 1960 and 1962 has already been published 
(Guichard & Harvey, 1967). In the pages that follow, the numbers given (in paren- 
theses) after the names of the Provinces refer to the Guichard & Harvey (1967) 
collecting site-numbers. For dates and ecological details of the sites, reference 
must be made to that paper. The present work on the sawflies, though based 
mainly on the Guichard and Harvey collections, has been broadened to include 
material from other countries bordering the eastern Mediterranean, such as that 
collected by G. A. Mavromoustakis in Cyprus (Benson, 1954), by Dr. Bytinski-Salz 
and others in Israel (Benson, 1955), by Dr. Zhelochovtsev (1941) and Dr. Dadurian 
(1958) in Armenia, by G. Heinrich in the Elburz Mountains, N. Iran (Berlin 

* Since the submission of this paper for publication, we learned with deep regret of the death of the 
author. Certain editorial changes have been made in the paper, which we were unable to submit to him. 
(Editor's Note.) 

ENTOM. 22, 4. 4 



ii2 R. B. BENSON 

Museum) and more recently by D. B. Baker on the S. Caspian Coast and Elburz 
Mountains of N. Iran. 

There is no doubt that in collecting this material, the larger and more highly- 
coloured flower-haunting species have been favoured, whereas most of the small 
dark inconspicuous species on low plants, as well as those associated with forest 
trees, have yet to be explored. The most useful approach therefore seemed to be 
to treat this fauna as though it were an extension of that of C. Europe as described 
by Enslin (1912-18) and Benson (1951-58), and in the families other than the 
Tenthredinidae, the Palaearctic fauna described by Gussakovskii (1935 and 1947). 
References to species given in full by Konow, 1905 or Gussakovskii are not repeated 
here. 

Wherever possible, new keys have been provided to the known world species for 
the genera centred in the E. Mediterranean: i.e., for Corynis, Tenthredopsis, Cuneala, 
Elinora and Sciapteryx. 

In Tenthredo a key has been constructed to world species-groups, and separate 
keys to species have been provided for the following species-groups: bifasciata, 
zonula, and maculata-temula; keys are also provided as follows: Calameuta and 
Trachelus species of the E. Mediterranean; the picipes-ciliatus-group of Dolerus; 
Periclista and the blanda-duodecimpunctata- and the postica-groups of Macrophya 
in the W. Palaearctic. 

I am much indebted to many people who have helped me in this work : especially 
to K. M. Guichard and D. H. Harvey for their careful attention to collecting sawflies 
in Turkey, and to D. B. Baker for the great care he has taken in collecting sawflies 
for me in N. Iran; to Dr. Oehlke of the Deutsches Entomologisches Institut, Ebers- 
wald, Germany for his continued kindness in lending me numerous Konow types 
for study; to Dr. Konigsmann of the Zoologisches Museum der Humboldt-Uni- 
versitat zu Berlin for lending me Dr. Heinrich's material from the Elburz Mountains 
etc. ; to Dr. Zhelochovtsev of Moscow for sending in exchange most valuable repre- 
sentatives of species collected by himself in Armenia ; to Dr. F. Wolf of the Facultes 
des Sciences Agronomiques, Gembloux, Belgium, for lending me material collected 
in Turkey in 1965 by Dr. Demelt. 

Unless otherwise indicated, the new Turkish records given below are of specimens 
collected by Guichard or Harvey in 1959, 1960 or 1962. Those marked with an 
asterisk (*) are new to the recorded fauna of Turkey and a dagger (f) indicates that 
the type has been examined in the course of this work. 

XYELIDAE 

Xyela graeca J. P. E. F. Stein 

Xyela graeca J. P. E. F. Stein; Benson, 1958. 

TURKEY, C. and W. : Ankara (13), 5 $; Izmir (Smyrna). 
ALGIERS, AUSTRIA, GREECE, ROUMANIA, TURKEY and ISRAEL. 



PAMPHILIDAE 

For a key to the palaearctic species see Gussakovskii 1935. 



HYMENOPTERA FROM TURKEY 113 

*Acantholyda hieroglyphica (Christ) 

TURKEY, C. : Ankara (3) i $; Ankara, Cankaya, i $, 13. vi. 1963 (H. Ozeren). 
EUROPE and TURKEY. 

Acantholyda fumata (Enslin) 
TURKEY, S. : Mersin, Toros Dagi [" Cilicischer Taurus "], (type locality). Endemic. 

Cephalcia hartigi (Bremi) 
C. EUROPEAN ALPS and TRANSCAUCASIA. 

*Neurotomafausta (Klug) 

TURKEY, W. : Bursa, Karacabey, iv-v.ig28 (Ajtai). 
C. and S. EUROPE and TURKEY. 

Neurotoma saltuum (L.) 

TURKEY, N. : Sakarya, Adapazari, on Prunus armeniaca L., 2.vi.ig57 (M. 
Akdogan). 

EUROPE, ASIA MINOR, TURKEY and TRANSCAUCASIA. 

*Neurotoma nemoralis (L.) 

TURKEY, C. : Kiitakya, Sureya, 1923, on leaves of Prunus cerasus L. 
EUROPE and TURKEY. 

Celidoptera maculipennis (J. P. E. F. Stein) 

TURKEY, C.: Ankara, Kiire Dag, near Bala, 1500 m., 2 $, 2 ?, lo.v.igsg, flying 
round Prunus spinosa L. (E. S. Brown). 

Previously recorded only from TURKEY, W. and C. : Izmir and Amasya Provinces. 
Endemic. 

Pamphilius caucasicus Gussakovskii, 1935 

TRANSCAUCASIA. Endemic. 

Pamphilius trigarius Konow 
TRANSCAUCASIA. Endemic. 

Pamphilius lethierryi (Konow) 
TRANSCAUCASIA. Endemic. 

Pamphilius aurantiacus (Giraud) 
S. EUROPE and TRANSCAUCASIA. 



n 4 R. B. BENSON 

MEGALODONTIDAE 

The species of this family are found mainly as adults on the flower tables of Umbel- 
liferae, on which plants the larvae feed socially in webs. The species are keyed 
by Gussakovskii, 1935. 

Megalodontes imperialis Konow 
ISRAEL and TURKEY. 

Megalodontes phoenicius (Lepeletier) 
ISRAEL, SYRIA and TURKEY. 

Megalodontes kohli Konow 

TURKEY, C. N. and N.E. : Amasya (5); Kayseri (4); Samsun (3); Erzurum (4). 

4?- 

CRIMEA and TURKEY. 

^Megalodontes multicinctus (Mocsary) 

TURKEY, C.: Ankara, Bala district, Ucern, 3 <j>, 28^.1959 (E. S. Brown). 
TURKEY and TRANSCAUCASIA. 

Megalodontes exornatus (Zaddach) 
HUNGARY, GREECE, TURKEY and TRANSCAUCASIA. 

Megalodontes loewi (Stein) 
TURKEY and TRANSCAUCASIA. 

Megalodontes flabellicornis (Germar) 

TURKEY, C. and N.E. : Nigde (5); Erzurum (i and 4); Artvin (7). 2 <, 8 <f>. 
S. EUROPE and TURKEY. 

Megalodontes escalerai Konow 

LEBANON: Djezzine, i $, 2.vi.i953 (G. A. Mavromoustakis). 
LEBANON, ISRAEL and SYRIA. 

Megalodontes pectinicornis (Klug) 

TURKEY, C. and S. : Ankara (12) ; Amasya (5, 7 and 14) ; Mersin (9). 20 <$, 35 $. 
HUNGARY and TURKEY. 



MESOPOTAMIA. 



HYMENOPTERA FROM TURKEY 115 

Megalodontes olivieri (Brulle) 



Megalodontes medius Konow 

TURKEY, N.W. and C. : Bursa (4); Corum (2). 2 ^, 2 ?. 
S.E. EUROPE, TURKEY and TRANSCAUCASIA. 

^Megalodontes laticeps Konow 

TURKEY, C. : Konya, Aksehir, i $, 1-12. vi. 1955 (Seidenstucker] ; Nigde (i). i ?; 
Kayseri (4), i $. ISRAEL: Carmel, Place of Sacrifice, 2^, i <j>, 26 . iii-g . iv . 1930 
(Tapuchi) . 

S.E. EUROPE, TURKEY and ISRAEL. Not previously recorded outside Europe. 

Megalodontes flavicornis (Klug) 

TURKEY, N.E.: Artvin (7). i ?; IRAN: Azerbaijan, Bazergan, 1300 m., i $, 
25. .1960 (E. S. Brown). 

S.E. EUROPE, TURKEY, TRANSCAUCASIA, N. IRAN and W. TURKMEN. 

Megalodontes aquilus Konow 
TRANSCAUCASIA. 

^Megalodontes klugi (Leach) 

Tarpa spissicornis Klug. 

TURKEY, N.W. : Bursa (9). i ^. 
C. and S. EUROPE and TURKEY. 

Tristactus judaicus (Lepeletier) 

Tarpa judaica Lepeletier. 
T. caesariensis Lepeletier. 
^[Tristactus punctatus Konow, syn. n. 

A series of this species (14 J, n $) from S. TURKEY : Mersin (5) agrees with a series 
given to the B.M. (N.H.) by Enslin from S. TURKEY: Mersin, Tarsus, 23-24.^.1955 
(Seidenstiicker) and S.E. TURKEY: Diyarbakir, Ergani (Osmaniye), 5-6. v. 1955 
(Seidenstiicker}. These all have strongly infuscate wings, the males with a white 
clypeus and mandibles, and the females with dark mandibles and a variable extension 
of black on the clypeus ; it would not be surprising to find individuals with an entirely 
black clypeus and caesariensis would seem to be just such an aberration. A series 
from ISRAEL: Mt. Tiberias, 2$, 20.iii.i939 (Palmoni), i <^, 14.^.1963 (C. H. 
Andrewes); Benkamina, i , 23-25.^.1942 and Elon, i cj, 18^.1946 (H. Bytinski- 



n6 R. B. BENSON 

Salz) differ from the form from Turkey in having subhyaline wings. The punctation 
of the mesonotum is also variable and T. punctatus is not clearly differentiated. 
TURKEY, SYRIA, ISRAEL. 

XIPHYDRIIDAE 

For a key to palaearctic species see Gussakovskii, 1935. 

Xiphydria caucasica Semenov & Gussakovskii 
TRANSCAUCASIA. 

SIRICIDAE 

For keys to species see Benson, 1943. 

SIRICINAE 

Xeris spectrum (L.) 
TURKEY, N.E.: Trabzon (14). 3$. 

EUROPE to CAUCASUS and ASIA MINOR, SIBERIA and mountains to HEPTA- 
POTAMIA, SAKHALIN, boreal N. AMERICA and mountains south to COLORADO. 

Urocerus augur augur (Klug) 

Sir ex cedrorum Smith. 

Urocerus augur bensoni Maa, 1949, syn. n. 

TURKEY, N.E. : Artvin (5). 2 $ in a timber yard. 
Mountains of C. EUROPE, TURKEY and LEBANON. 

Urocerus augur sah Mocsary 

TURKEY, W. : Izmir, Odemis, Boz Sira Daglari, 1300-1900 m., i 9, i6.viii.i95o 
(P. H. Davis). 

Mountains of N. AFRICA, W. TURKEY, TRANSCAUCASIA, N. IRAN, TURKESTAN and 
AFGHANISTAN. 

Urocerus gigas gigas (L.) 
Occasionally introduced from Europe. 

Urocerus gigas argonautarum Semenov 

TURKEY, N. and N.E. : Bolu (3). i $; Trabzon (2). i $ in timber yard. 
TURKEY and TRANSCAUCASIA. 



HYMENOPTERA FROM TURKEY 117 

Sir ex juvencus (L). 
TRANSCAUCASIA (Dadurian, 1958). Holarctic. 

Sirex cyaneus dux Semenov 

This form agrees with 5. cyaneus F. in ovipositor /sawsheath ratio and in all other 
essential characters, but the total ovipositor is longer than in that form and is 
almost as long as a fore wing (see Benson, 1943). It seems reasonable to treat 
forms differing only in ovipositor/fore wing ratio in Sirex as at most subspecies, as 
was done by Benson in Urocerus. TRANSCAUCASIA. 

Sirex noctilio F. 

TURKEY, S.W. : Denizli, ex Pinus bmtia Ten, 3 <$, 2 $, x.ig62 (Dr. Hasan Canak- 
sioglu). 

This series of small dark- winged specimens (15-16 mm. long) and a similar series 
from CYPRUS, x.ig2j (H. M. Morris) in the B.M. (N.H.) may have been imported. 

Temperate EURASIA, introduced into N. AMERICA, NEW ZEALAND and AUSTRALIA. 

TREMECINAE 

Tremex jakovlevi Semenov 
TRANSCAUCASIA. 

ORUSSIDAE 

Orussus abietinus (Scopoli) 
EUROPE, SYRIA and CAUCASUS. 

*Mocsarya syriaca Benson, 1936 

The unique original specimen of this species was collected at Akbes (Meidan 
Ekbes) on the Turkish border of Syria and as it had lost its antennae, its original 
generic assignment was doubtful. Guichard and Harvey collected a second speci- 
men, i $ in N. TURKEY : Zonguldak, near Ulus, c. 100 m., 17^.1962. The posses- 
sion of antennae with a small scape shows that the species had been correctly assigned 
to Mocsarya rather than to Chalinus. The male is only 9 mm., with subhyaline 
wings, and the carinae within the facial field are incompletely developed, not un- 
naturally in a small specimen. 

SYRIA and TURKEY. 

CEPHIDAE 

A classification of the Cephidae was given by Benson (1946) and most of the 
species from Asia Minor can be named in Gussakovskii's work (1935). but keys to 
E. Mediterranean Calameuta and Trachelus are given below. 



n8 R. B. BENSON 

HARTIGIINI 

Hartigia nigra (Harris) 
TURKEY, N.E.: Erzurum (5). i $. 
C. and S. EUROPE and TURKEY. 

Hartigia linearis (Schrank) 
C. and S. EUROPE, TRANSCAUCASIA, S. SIBERIA. 

[Janus femoratus (Curtis) 

TURKEY, N.W. : Istanbul (3). i $. Not recorded yet from Asia Minor. 
This specimen differs from the normal C. European form in having a brown 
instead of yellowish white tegula. 

EUROPE.] 

[Janus compressus (Fabricius) 

TURKEY, N.W. : Istanbul (3). 2 9. Likewise not yet recorded from Asia Minor. 
S. EUROPE to N. CAUCASUS.] 

Syrista parreyssii (Spinola) 

TURKEY, N.W., C. and N.E. : Tekirdeg (i) ; Amasya (3, 4, 5 and 7); Ankara (33); 
Nigde (5); Erzurum (4). 15 <?, 12 $. 

The flight period is from the end of May in Amasya and Nigde, to 23rd July at 
2,300 m., Erzurum. 

S. EUROPE to CAUCASUS, TURKEY, CYPRUS and ISRAEL. 

PAGHYGEPHINI 

Pachycephus smyrnensis J. P. E. F. Stein 

Pachycephus brevis Ghigi, syn. n. 
\Spatulocephus sanctus Pic, syn. n. 

This species is common and very variable in size (6-u mm.) and colour pattern. 

In CYPRUS and ISRAEL it flies in March and April. In TURKEY at Mersin (8), 
at 600 m. and at Amasya (i, 2 and 3) at 400-500 m., it was found in the last week 
of May and first week of June. 39 <$, 50 $. 

BALKANS, TURKEY, CYPRUS, SYRIA, ISRAEL and TRANSCAUCASIA. 

Pachycephus aeneovarius Kohl 

This species is very similar to P. smyrnensis except that the dark colour of the body 
has greenish and bluish metallic reflections, and the head and thorax are even more 
densely punctured. 

Only known from TURKEY: Eskischir, Sebandseln (Kohl, 1905). Endemic. 



HYMENOPTERA FROM TURKEY 119 

Pachycephus persicus Gussakovskii 

This species is based on a unique $ of 6 mm. length, in which the yellowish white 
markings are replaced by white. It would seem probable that this is a form of 
P. smyrnensis. 

S. IRAN: Luristan, in. 1904. 

[Pachycephus cruentatus cruentatus (Eversmann) 

This species is distinguished from the above in: (i) lacking any sign of a genal 
carina on the lower lateral hind margin of the head capsule ; (2) the strongly shining 
surface of head and thorax with obsolescent punctures ; and (3) the deeply infuscate 
wings. 

S. RUSSIA, CRIMEA, N. CAUCASUS]. 

Pachycephus cruentatus konowi Kohl 

This subspecies differs from the typical form in having more extensive red colour 
on the legs. It was described originally from TURKEY : Kayseri, Ereiyas and Dagi, 
and has also been found in ISRAEL, at Urim. 

Characopygus decoratus sp. n. 

$. Black with following parts yellow: palps, mandibles (except teeth), fleck in middle of 
fronto-clypeal area, pronotum, behind, tegula, small fleck on each lateral mesonotal lobe, large 
fleck on scutellum, upper angle of mesepisternum legs (except most of coxae, trochanters and 
bases of femora which are black, and the yellow colour is reddish tinged) tergites 6, 7 and 9 
almost entirely, 3rd laterally and with medial fleck, and 8th with lateral and medial flecks 
broad, apical margin of hypopygium, small fleck on the preceding sternite, and lower edge of 
basal plate flanking the ovipositor. Wings hyaline; stigma, C and Sc yellow; rest of venation 
brown to piceous. 

Length: u mm. (without ovipositor). 

Head normal, subparallel-sided. Distance between antennal sockets compared to distance 
between socket and middle of anterior tentorial pit as i-o : 0-8. Antenna 22-segmented ; 
becoming clavate from about yth segment; loth onwards transverse, and 2oth about two and a 
half times broader than long. POL : OOL : OCL as i-o : 1-3 : 1-9. 

Thorax normal. Legs with hind tarsus about as long as tibia, basitarsus longer than 3 fol- 
lowing tarsal segments together. Spurs normal. Inner hind tibial spur longer than apical 
width of tibia as 1-2 : i-o. Claws with inner tooth about as long as end tooth, parallel with it 
but stouter at base. 

Abdomen with ovipositor about as long as 3 basal hind tarsal segments. Sawsheath at about 
45 out of alignment with basal plate and shorter than basal plate as i-o : 1-2. 

Punctation obsolescent on frons, which is shining; temples shining between shallow punctures 
separated by about two diameters from each other. Mesonotum and mesopleuron with denser 
punctures separated by about one diameter, but on the front lobes the interspaces between the 
punctures are dull, with fine surface sculpture and on the scutellum the punctures are shallow 
and with more widely shining interspaces. Abdomen dull with follicles and fine surface 
striations. 



lao R. B. BENSON 

Pubescence : Head and thorax with dense pubescence about as long as diameter of an ocellus, 
fuscous on the dark surfaces and colourless on the yellow surfaces. Abdomen likewise densely 
clothed all over with very short pubescence. 

(J as $ but lateral lobes of mesonotum entirely black, and abdomen with tergites 4, 6 and 7 
almost entirely, 3 laterally, posterior half of 8 and hypopygium and apical segments entirely 
yellow. 

Antenna with 2oth segment about x 2 broader than long. Sternite of 8th abdominal segment 
with an apical fringe of strap-shaped setae; hypopygium normal, drawn out into a narrow 
tongue-like protuberance, slightly swollen apically, 9 mm. 

Holotype $. ISRAEL: near Jerusalem, Ejn, Karim, 10.111.1959 (H. Bytinski- 
Salz). B.M. (N.H.). 

Paratype. ISRAEL: Holou, i ^, 28.111.1959 (H. Bytinski-Salz). B.M. (N.H.). 

This new species is at once distinguishable from those previously known by its 
rich yellow markings. None of the other species have any tergites banded right 
across with yellow, or mesopleurum or scutellum yellow-flecked. 

In structure the species are very similar, but the 22-segmented antenna may prove 
to be characteristic (C. moricei Konow and C. scythicus Dovnar-Zapolskii have 
20 segments and C. modestus Dovnar-Zapolskii but 18). 

Characopygus scythicus Dovnar-Zapolskii 
TURKEY: Mersin, Gozne, 1,800 m., i $, 2.vi.i96o (Guichard & Harvey). 

Previously known only from S. and S.E. RUSSIA (Askamia-Nova, Sarepta and 
Orenburg). 

CEPHINI 

Attached to wild and cultivated Gramineae. 

Cephus pygmaeus (L.) 

Cephus tanaiticus Dovnar-Zapolskii, syn. n. 
C. notatus Kokujer, syn. n. 

The descriptions of C. tanaiticus Dovnar-Zapolskii from S. RUSSIA and C. notatus 
from Transcaucasia agree with melanic forms of this species that I have seen in 
Britain and elsewhere. Pest of corn and other cultivated grasses. 

This species, apart from its introduction into eastern N. AMERICA, occurs through- 
out EUROPE, west of the Volga and Caspian and south of about 60 lat. and also in 
N. IRAN, TURKEY, SYRIA and ISRAEL. 

Cephus brachycercus C. G. Thomson 
TURKEY: Samsun (5). i <$. 
EUROPE, SIBERIA and TURKEY. 

Cephus nigrinus C. G. Thomson 
SYRIA: Meidan Ekbes (Konow, 1891). 
Throughout EUROPE, SYRIA. 



HYMENOPTERA FROM TURKEY 121 

Cephus berytensis (Pic) 
\PeronistilimoYphus berytensis Pic, 1916. 

Type examined 24.^1.1964. $. Abdomen and antennae beyond I3th segment 
are now missing It would appear to be a true Cephus, because the distance between 
the antennal sockets is approximately the same as the distance from a socket to the 
middle of the anterior tentorial fovea. Antenna swollen from I3th segment. Claws 
with minute inner tooth. Basal antennal segment yellow and legs entirely yellow 
except for trochanters and i coxae. 

SYRIA (Beirut). 

Species incertae sedis 

Cephus nigricarpus Andre (Syria). 

Cephus obscuriventris Pic, 1918 (Lebanon) [Type lost. Pin and labels in Paris 
Mus., seen 24.^1.1964.] 

Cephus politissimus Costa (Armenia). 

CALAMEUTA Konow 

KEY TO E. MEDITERRANEAN MALES AND FEMALES 

1 Either scutellum shining with at most widely spaced punctures, or with one or 

more of the tergites entirely yellow ........ 2 

Scutellum and whole of mesonotum dull with dense microsculpture. No 
tergite entirely yellow. Hind tibia normally with 2 preapical spines. Wings 
subhyaline. No antennal segment transverse. Tergites normally i mar- 
gined with yellow apically but may be entirely black. 9-12 mm. 

filifortnis (Eversmann) 

2 (i) Maxillary palp with 5th segment much shorter than 6th (apical) ... 3 

Maxillary palp with 5th segment almost as long as 6th. 

Very variable in colour but $ always has apical tergite yellow and may 
have other tergites flecked or banded or entirely yellow, but has hind legs 
entirely black ; in $ even the hind legs may be flecked with yellow on tibia 
and femur as also may be the face, mesopleura and pronotum. 7-10 mm. 

haetnorrhoidalis (F.) 

3 (2) Larger species (10-15 mm.) with more than one tergite entirely yellow and 

hind tibia with only one pre-apical spine ...... 4 

- Smaller species (4-10 mm.) with at most only one tergite entirely yellow or hind 

tibia with 2 pre-apical spines ......... 5 

4 (3) Wings strongly infuscate. Head, thorax and legs entirely black. Abdomen 

almost entirely reddish yellow ...... pygmaea (Poda) 

Wings subhyaline. Head, thorax and legs yellow-marked, abdomen banded 

yellow and black idolon (Rossi) 

5 (3) Scutellum impunctate. Abdomen in $ entirely black; in <$ with 4 middle tergites 

yellow-margined. Hind tibia yellow ..... pallipes (Klug) 
Scutellum with definite punctures. Abdomen in $ with at least apical tergite 
yellow, and <$ with middle tergites yellow-margined as in pallipes. Hind 
tibia yellow or infuscate. (S.E. EUROPE and S. SIBERIA) 

pravei (Dovnar-Zapolskii) 



122 R. B. BENSON 

Calameuta filiformis (Eversmann) 

Cephus grombczevskii Jakovlev, syn. n. (Turkestan). 

Cephus infernalis Dovnar-Zapolskii, syn. n. (Caucasus). 

Cephus turanicus Dovnar-Zapolskii, syn. n. (Turkestan bei Taschkent). 

Calameuta amurensis Gussakovskii syn. n. (Amur). 

I have representatives of C. grombczevskii and infernalis named by Gussakovskii 
and they, together with C. amurensis, appear to be melanic forms of C. filiformis, 
C. infernalis retaining a yellow fleck on the side of the 5th tergite and C. grombczevskii 
a yellow fleck on the 5th and 6th tergites; C. infernalis has, on the contrary, paler 
hind tibiae than C. filiformis. C. turanicus appears to be a paler form than C. fili- 
formis. 

In addition to EUROPE, SIBERIA, TRANSCAUCASIA, N. IRAN and SYRIA, the species 
also occurs in LEBANON, where i <$ and i $ of the typical European form were 
collected by G. A. Mavromoustakis at Hammana, i6.v.i953 and i $ at Falouka, 
17. v. 1953. 

Calameuta haemorrhoidalis (F.) 

^Cephus gracilicornis Konow, Caucasus. 

Cephus diver sipes Ghigi, Rhodes. 

Trachelus syriacus Pic, Syria. 

f Calameuta f estiva Benson, 1954, Cyprus, syn. n. 

Guichard and Harvey brought back 10 , 17 $ of this species from GREECE, 
THRACE and various parts of TURKEY, N.W., W., S.W., C., N. and N.E. ; Istanbul (3) ; 
Bursa (i and 2) ; Mugla (7) ; Antalya (13) ; Amasya (i and 7) ; Ankara (15 and 
23) ; Samsun (6 and 7) ; Ersurum (8) . 

Most of them were collected in May and at various altitudes up to 1,800 m., near 
the summit of Elma Dagi, Ankara on 21. v. 1960. The long 5th segment of the 
maxillary palp distinguishes this species from all other Cephinae, and enables a 
vast range of colour-pattern forms to be associated here, including the types of 
C. f estiva and gracilicornis. 

S.E. EUROPE, TURKEY, TRANSCAUCASIA, SYRIA, ISRAEL and W. TURKMEN. 

Calameuta pygmaea (Poda) 

This is mainly a species of S.W. EUROPE and N. AFRICA but it extends through 
EGYPT to ISRAEL, where i $ was taken 20. iv. 1953 at Wadi Ruaz, Beth Hale rem, 
Jerusalem (0. Theodor] and i $ at Oqanim, g.iv.ig6i (Sandier}. 

Calameuta idolon (Rossi) 

\Monoplopus apicicornis Pic, syn. n. 

Another species varying greatly in the amount of yellow colour on the head and 
thorax, so that it is not possible to draw any line of distinction from the form de- 
scribed as apicicornis with the head mainly yellow. Guichard and Harvey collected 



HYMENOPTERA FROM TURKEY 123 

24 , 12 $ in TURKEY, C.: Ankara (4, n, 12 and 14) and Amasya (7), mostly from 
the flowery edges of a lake and streams. 

N. AFRICA, S. EUROPE to CAUCASUS, TURKEY, IRAN, SYRIA, LEBANON and ISRAEL. 

*Calameuta pallipes (King) 

A <$ was collected in TURKEY: Ankara, 800 m., 22. v. 1960 (Guichard & Harvey] 
and i $ on the Black Sea coast at Samsun, nr. Engiz, 17^.1959 (Guichard}. 
Apart from these two records it has not been found outside Europe. 

N. and C. EUROPE and TURKEY. 

TRACHELUS Jurine 
KEY TO MEDITERRANEAN MALES AND FEMALES 

1 Antenna with pre-apical segments at least x 2 as broad as long. 6-9 mm. . 2 
Antenna with no antennal segments x 2 as broad as long. 7-14 mm. . . 3 

2 (i) Hind tibia without pre-apical spine. $ with 3 apical sternites excavated and 

bearing modified setae. Pronotum yellow-flecked. 6-9 mm. libanensis (Andre) 
Hind tibia with i pre-apical spine. <$ without modified apical sternites. Pro- 
notum flecked with yellow ...... judaicus (Konow) 

3 (2) Larger species (10-14 mm.). Tibia pale, abdomen often with some of the 

tergites yellow-margined apically ........ 4 

Smaller species (7-10 mm.). Legs entirely black. Abdomen black, with yellow 
lateral stripe or row of flecks but no tergites yellow-margined apically. 
Usually 2 pre-apical hind tibial spurs ..... tabidus (F.) 

4 (3) Head capsule and mesonotum (apart from scutellum) entirely black. 1-2 pre- 

apical tibial spurs present. Cross vein 3 rm present in fore wing ... 5 

Head capsule, and mesonotum richly marked with yellow as also is abdomen. 
Hind tibiae without pre-apical apurs. Fore wing with vein 3 rm missing. 
(cJ unknown). 12 mm. (ASTRACHAN) ..... tigris (Benson) 

5 (4) Antenna, pronotum, scutellum and abdomen richly marked with yellow. 

10-12 mm. (SPAIN and N. AFRICA) .... flavicornis (Lucas) 

Antenna, pronotum and scutellum entirely black, and abdomen with at most 

apical margins of some of the segments yellow. 10-14 rnm. troglodyta (F.) 

Trachelus judaicus (Konow) 

\Monoplopus judaicus Konow. 

\Monoplopus notaticollis Pic. 

\Micvocephus judaicus (Konow) Benson, 1935. 

This species is very similar to the next following, apart from the characters men- 
tioned in the key, and Benson (1946) concluded that the species belonged to this 
genus despite its loss of the modified apical sternites in the male. 

ISRAEL. 

Trachelus libanensis (Andre) 

\Ateuchopus armenius Konow, syn. n. 

In a series of over 100 specimens of this species from Cyprus, the pronotum varies 
from entirely black to mainly yellow and the development of punctures on the 



124 R- B. BENSON 

mesonotum likewise bridges the gap between the " species " libanensis and armenius. 
In the Transcaucasus the pronotum is said to be entirely black, whereas in Israel 
the pronotum is almost entirely yellow. 

The species was found in TURKEY, C. : Yozgat (i) on a flowery hillside at 1,000 m. 
and at Mersin (14) at 1,600 m., near crops of corn. 

TURKEY, CYPRUS, SYRIA, LEBANON and ISRAEL. 

Trachelus troglodyta (F.) 

^Astatus tenuicornis Konow, 1902, syn. n. 

T. troglodyta has previously been recorded from only N. and C. EUROPE and 
is replaced in Transcaucasia by tenuicornis. The specimens collected from TURKEY, 
N. and N.E. at Rize (2) and Samsun (6 and 28) appear to be normal troglodyta. 
They were collected in swampy woods and the borders of marshes. 

N. and C. EUROPE, to TURKEY and TRANSCAUCASIA. 

Species incertae sedis 

Cephus (Fossulocephus) citriniventris Pic, 1917 (Algeria). 

ARGIDAE 
ARGINAE 

For keys to Palaearctic Arge species see Gussakovskii, 1935. 

Arge ochropus (Gmelin) 
Tenthredo rosae L. ; auctt. nee L. 

TURKEY, N.W., S., C., N., and N.E.: Bursa (2); Bolu (3); Antalya (5 and 6); 
Ankara (3); Sinop (4); Samsun (10) ; Amasya (i, 2, 5, 6 and 7); Tokat (4); Adana 
(3 and 6); Trabzon (3 and 8); Erzurum (6). 12 <$, 22 $, from 4_iv to i6.viii and at 
altitudes up to 1,700 m. 

EUROPE, W. and C. SIBERIA, EGYPT, ISRAEL, LEBANON, SYRIA, TURKEY, CYPRUS, 
TRANSCAUCASIA, N. IRAN and TURKMEN REPUBLIC. 

Arge simulatrix Konow 

TURKEY, C. and E. : Ankara (32); Amasya (i and 2); Nigde (5); Gumusane (i). 

4 < 9 ? 

GREECE, SYRIA, TURKEY and TRANSCAUCASIA. 

Arge frivaldzkyi (Tischbein) 

TURKEY, W. and C. : Bursa (4) ; Ankara (2) ; Amasya (5 and 7). 6 ^, 4 $. 
HUNGARY, GREECE, TURKEY and TRANSCAUCASIA. 



HYMENOPTERA FROM TURKEY 125 

*Arge beckeri (Tournier) 

TURKEY, E.: Erzurum (6). i $, i $. 
S.E. EUROPE and TURKEY. 

Arge pyrenaica (Andre) 
TURKEY, N.E. and E. : Trabzon (16); and Erzurum (4). 1^,1$. 

Mountains of N. AFRICA, PYRENEES, C. and S. EUROPE, SARDINIA, TURKEY, 
TRANSCAUCASIA, N. URAL, C. ASIA, MONGOLIA. 

Arge pagana (Panzer) 

EUROPE, TRANSCAUCASIA, SIBERIA, KAMTCHATKA, MONGOLIA, MANCHURIA, N. 
CHINA and JAPAN. 

Arge carinifrons Enslin 
TRANSCAUCASIA. 

Arge persica Gussakovskii 
N. IRAN. 

Arge impressifrons Konow 
TRANSCAUCASIA and N. IRAN (Talysk). 

Arge cyanocrocea (Forster) 
Hylotoma syriaca Mocsary, syn. n. 

Benson (i958a) treated A. syriaca as a species distinct from cyanocrocea, as he 
found various venational and sculptural differences correlated with the blackened 
legs of syriaca. Much more material shows that these characters are not significantly 
correlated and that numerous intermediate combinations occur. 

In the cooler parts of Europe all the tibiae and basitarsi as well as the hind femur 
are marked with yellow; in the warmer parts the legs from the front become more 
extensively blackened and in various parts of the Mediterranean from Spain to 
Lebanon occur forms with all the legs entirely or almost entirely black. On the 
S. Caspian Coast in N. IRAN, D. B. Baker has collected a series with still more melanic 
tendencies: in 26 $, 12 $, in addition to the legs being black, the wings show a range 
from normal colouring to almost entire infuscation ; and 48 <$, 9 $ have, in addition 
to entirely black wings, a deeply infuscate abdomen at most only obscurely brownish 
on the middle segments. 

TURKEY, N.W., S.W., S., C., N. and N.E.: Istanbul (2 and 3); Mugla (5 and 10); 
Mersin (6) ; Sinop (3) ; Amasya (14) ; Samsun (9 and 10) ; Mersin (6) ; Rize (8) ; Artvin 
(3 and 6). 7 <$, 25$. 

ENTOM. 22, 4. 5 



126 R. B. BENSON 

C. and S. EUROPE, LEBANON, SYRIA, CYPRUS, TURKEY, TRANSCAUCASIA, IRAN 
and TURKMEN. 

Arge melanochroa (Gmelin) 

(Text-fig, i) 
Hylotoma nigritarsis Klug, syn. n. 

In the E. Mediterranean, forms of this species often have the ist tergite i infus- 
cate and have been separated as a distinct species (nigritarsis) but every intergrade 
occurs in our series of over 150 specimens. The species is very similar to the follow- 
ing (scita) but can be distinguished by the black sawsheath in the $ with the large 
inner teeth, and in the <$ by the entirely different form of penis- valve (Text-fig, i) ; 
in both sexes also by absence of a continuous longitudinal glabrous patch on the 
mesosternum. 

TURKEY, N.W., S.W., S., C., N., N.E. and E.: Istanbul (i, 2 and 9); Mugla (5); 
Mersin (6 and 7) ; Ankara (39, 53 and 54) ; Amasya (i, 5, 7 and 14) ; Tokat (3 and 4) ; 
Zongulduk (i); Sinop (3 and 4); Giresun (5); Erzurum (4 and 6). 61 J, 48$. 

C. and S. EUROPE, SYRIA, CYPRUS, TURKEY, TRANSCAUCASIA and IRAN. 

Arge scita (Mocsary) 

(Text-fig. 2) 

Hylotoma proximo, Andr6, syn. n. 
\Avge debilis Konow, syn. n. 
A. zarudnyi Gussakovskii, syn. n. 

As in A. melanochroa, this species varies in the amount of infuscation of the 
ist tergite, and also in whether the anastomosis of M with R in the fore wing is long 
or short, so that it is impossible in our long series from Cyprus and Turkey of over 
120 specimens to segregate proxima and debilis from scita. From melanochroa 
this species is always to be distinguished in the $ by its pale sawsheath with only 
small inner teeth, in the <$ by its very characteristic penis-valve (Text-fig. 2) and 
by the longitudinal glabrous patch on the mesosternum in both sexes. 

TURKEY, W., S.W., S., C. and E. : Bursa (69 and 14); Kutahya (u); Mugla (7); 
Hatay (3); Mersin (6); Nigde (5); Maras (4 and 5); Ankara (3, 4, 17, 53 and 54); 
Amasya (i, 3, 5, 6, 7 and 8); Tokat (3); Erzurum (4). 32 <, 24 $. It has been 
reared from a larva on Prunus amygdalus Batsch (CYPRUS: Paphos, vi.1950 (Th. 

Shiakides) . 

GREECE, ISRAEL, LEBANON, SYRIA, CYPRUS, TURKEY, TRANSCAUCASIA, IRAN and 
TURKMEN. 

Arge cingulata Jakoulev 
Arge turanica Kuznetzov-Ugamskii, syn. n. 

IRAN: Mazanderan, Panjak Rustaq, 860-1,125 m., 7 ^, i $ 23^.1966 and 7^, 
5 ?, 3. v. 1967 (D. B. Baker). 

TURKESTAN and IRAN. 



HYMENOPTERA FROM TURKEY SYMPHYTA 127 

*Arge clavicornis seljuki ssp. n. 

Differs from the closely related A . clavicornis fuscipes Fallen in that the infuscate band below 
the stigma stretches right across the fore wing to the anal margin as in A. dimidiata Fallen. 

Holotype $. TURKEY: Trabzon, Zigana Dagi, 1,400 m., I3.vii.ig6o (Guichard 
& Harvey}. B.M. (N.H.). 

Paratype $. Same data. B.M. (N.H.). 

The species occurs throughout EUROPE, and in TURKEY, TRANSCAUCASIA, SIBERIA 
and N. AMERICA. 

Arge pallidinervis Gussakovskii 
TRANSCAUCASIA. 

Arge aurata (Zaddach) 

This species is very closely related to the European A. ustulata and clavicornis 
but, apart from its entirely dark legs, the pubescence on its head and thorax is 
golden instead of silvery. 

TURKEY, S. and E. : Nigde, Ciftehan, 900 m., 2 <$, 26. v. 1960 (Guichard & Harvey) ; 
and Erzurum (6). I $. 

Arge auripennis Konow 
S.E. EUROPE, SYRIA and TRANSCAUCASIA. 

Arge rustica (L.) 

TURKEY, W., S., C., N., N.E., and E.: Izmit, Alem Deg, 600 m., i $, 30. vi. 1966 
(Demelt); Antalya (13); Nigde (5); Ankara (41); Amasya (6, 12 and 13); Sinop (2); 
Samsun (3); Giresun (2); Gumusane (4); Erzurum (6). 22 $, 18 $. 

EUROPE, ISRAEL, TURKEY and TRANSCAUCASIA. 

Arge pleuritica (Klug) 

TURKEY, C., N., and E. : Aydin, Bozdogan, 2 $, 12 $, iv.igso (Plant Protection 
Institute); Amasya (5), 2 $, I $; Erzurum (5 and 6). 2 <$, 3 $. 
S.E. EUROPE, TURKEY, TRANSCAUCASIA and TURKESTAN. 

Arge berberidis (Schrank) 

TURKEY, S., C., N.E. and E. : Mersin (13); Ankara (37 and 39); Giresun (5); 
Gumusane (i and 5); Erzurum (6). i <$, 9 ?. 
C. and S. EUROPE, TURKEY and TRANSCAUCASIA. 

Arge nigripes (Retzius) 

TURKEY, C., N.E., E.: Ankara (39); Giresun (2); Erzurum (6). 4^, 8$. 
EUROPE, TURKEY, TRANSCAUCASIA and SIBERIA. 



128 R. B. BENSON 

Arge ciliaris (L.) 
EUROPE, TRANSCAUCASIA, SIBERIA, MONGOLIA, MANCHURIA. 

Arge gracilicornis (Klug) 
EUROPE, TRANSCAUCASIA, SIBERIA, JAPAN. 

Arge enodis (L.) 
TURKEY, C. : Ankara (35). i <$. 
C. and S. EUROPE, TURKEY, TRANSCAUCASIA, SIBERIA and JAPAN. 

Kokujewia ectrapela Konow 

Kokujewia dementi Zirngiebl, syn. n. 
^Kokujewia palestina Benson, syn. n. 

The type of palestina was reared from a larva and the supposed differences of 
the sawsheath and tarsi of the adult were probably due to its teneral condition. 
K. dementi is surely a further synonym, as slight differences of colour are only to 
be expected. 

TURKEY, C. : Ankara, Elma Dagi, 1,700 m., i <, 28. vi. 1959 (Guichard); the types 
of dementi were from Konya, Aksehir. 

ISRAEL, TURKEY and TRANSCAUCASIA. 

STERICTIPHORINAE 

Sterictiphora furcata (Villers) 

Hylotoma gastrica Klug, syn. n. 
Schizoceros nigripes Konow, syn. n. 
Schizoceros henschi (Konow), syn. n. 
Schizoceros bleusei Pic, syn. n. 

TURKEY, N.W., C., N. and E. : Istanbul (2 and 3) ; Ankara (39) ; Amasya (3 and 5) ; 
Zonguldak (i); Sinop (4); Erzurum (6). i ^, 9$. 

EUROPE, SYRIA, TURKEY, TRANSCAUCASIA and IRAN. 

Aprosthema tarda (Klug) 

TURKEY: Sinop, i $, i8.vi.i959 (Guichard}. 

Gussakovskii keys 45 palaearctic species of Aprosthema (and Pseudaprosthema] 
but most of these are colour forms of a very few genuine species. In Europe and 
Turkey there are probably only two, tarda and melanura (see Conde, 1934). 

EUROPE, ISRAEL, TURKEY, TRANSCAUCASIA, TURKESTAN, IRAN and SIBERIA. 

Aprosthema melanura (Klug) 
EUROPE to CAUCASUS, TURKESTAN, IRAN and SIBERIA. 



HYMENOPTERA FROM TURKEY j 29 

CIMBICIDAE 
ZARAEINAE 

To distinguish the species see Benson, 1951 : 39. 

Zaraea aenea (Klug) 
TURKEY, E.: Trabzon (3). i $>. 
C.E. EUROPE, TURKEY and TRANSCAUCASIA. 

Abia sericea (L.) 

TURKEY, S. and N. : Mersin (4); Samsun (18). 3 <$, i $. 
EUROPE, TURKEY and TRANSCAUCASIA. 

CIMBICINAE 

To distinguish the species see Gussakovskii, 1947. 

Pseudoclavellaria amerinae (L.) 

TURKEY, E. : Gumusane (5). 5 <, 2 <j>. 
EUROPE, TURKEY, SIBERIA and COREA. 

Palaeocimbex quadrimaculata (Miiller) 
C. and S. EUROPE, TURKEY and ISRAEL. 

CORYNINAE 
COR YNIS Thunberg 

KEY TO MALES AND FEMALES 

1 Abdomen and often head and thorax marked with pale colour (yellowish white 

to orange or red). Claws sub-bifid or with small inner tooth ... 2 

Body entirely dark, brown or black, at most legs sometimes pale. Claws sub- 
bifid ............. 14 

2 (i) Face, scutellum, mesopleura as well as abdomen red-marked. Clypeo-frontal 

area convex above and gibbous in shape . . . (sanguinea-Qroup) 3 

Pale colour yellow or yellowish white and usually less extensive. Clypeo- 
frontal area flattened to form straight line in profile ..... 5 

3 (2) Mesonotum and tergites covered with widely spaced large punctures with the 

interspaces shining though pitted with numerous fine punctures ... 4 

Mesonotum in front, and tergites laterally, densely and coarsely punctured 
without shining interspaces; scutellum and tergites medially with shining 
interspaces scarcely larger than diameter of punctures. 

Colour red with the following parts black: head apart from face below 
antennae, a medial and lateral fleck each side of mesonotum together with 
sunken parts thereof and metanotum, lower mesopleuron and mesosternum, 



130 R. B. BENSON 

coxae and base of femora, ist tergite (except laterally) and fore and hind 
margins of 2nd and 3rd tergites, together with most of sternites. Stigma 
and rest of wing venation brown. Antennae much longer than greatest eye 
measure, with club longer than distance between eyes in front. Hind ocelli 
about as far from eye-margin as from occipital carina (OOL = OOCL). 
Claws with small inner tooth. Inner front tibial spur almost as long as 
basitarsus. Pubescence on parts of head, mesonoton and mesopleurum as 
long as diameter of front ocellus. $ not seen. 6 mm. CANARY ISLANDS, 
MOROCCO and TUNIS .... sanguined (Snellen van Vollenhoven) 

4 (3) Claws sub-bifid. Tibiae and tarsi reddish as femora; tarsi not infuscate apically. 

Hind ocelli one and one-half times further from eye-margin as from occipital 
carina (OOL > OOCL). 

Colour as in sanguined but the 2nd-5th tergites are marked with black 
medially though progressively less. $ not known. ALGERIA and TRIPOLI- 
TANIA ......... f semisanguinea (Pic, 1916) 

Claws with only minute inner tooth. Tibiae and base of tarsi yellowish white 
in contrast to the red apex of the femora, and the apical tarsomeres infuscate. 
Hind ocelli about as far from eye-margin as from occipital carina (OOL 
OOCL). 

Otherwise coloured as in semisanguinea but that the lateral and medial 
dark flecks on the mesonotum are much enlarged and partially joined and 
the scutellum is infuscate laterally. 6mm. $ only. Holotype $. ISRAEL: 
Vadi Ajram, 7. v. 1954 (H- Bytinski-Salz) (in B.M.). ISRAEL haematica sp. n. 

5 (2) Antenna not longer than greatest measure of eye, and club often shorter than 

distance between eyes in front. Face very flat: clypeo-frontal area below 
scarcely raised above level of inner orbits. Frontal area of head and thorax 
almost glabrous ; pubescence on post-ocellar region of head much shorter than 
diameter of front ocellus. Claws sub-bifid. Inner front tibial spur about 
three-fourths as long as basitarsus ........ 6 

Antenna at least about one-fourth longer than greatest measure of eye and club 
often longer than distance between eyes in front. Face with clypeo-frontal 
area with declivous sides raising it above the level of the inner orbits. Head 
and thorax pubescent all over and in part at least almost as long as diameter 
of front ocellus. Claws with small inner tooth or sub-bifid. Inner front tibial 
spur not more than about two-thirds as long as basitarsus (except in citrina) 7 

6 (5) Face below antenna, fleck on mesonotum in addition to scutellum and meso- 

pleuron as well as tegula and pronotum yellow. Thorax with shining 
interspaces between punctures at least as large as punctures on mesopleura 
and sides of mesonotum. Post-ocellar region ? 5-6 mm. S. and E. 
EUROPE (N. to Leningrad) and S.W. SIBERIA ... $ amoena (Klug) 

Head with only clypeus, thorax with only tegula and pronotum, i yellow. 
Thorax very densely punctured, without shining interspaces as large as 
punctures on mesopleura or sides of mesonotum. Post-ocellar region con- 
vexly raised above level of hind ocelli. 6-7 mm. S.E. EUROPE, E. MEDI- 
TERRANEAN and TURKESTAN . . . concinna (J. P. E. F. Stein) 

7 (5) Pronotum and face below antennae i yellow, or malar space at most two- 

thirds as long as diameter of front ocellus in $ and less than one-half in <$ 8 

Pronotum and often face below antennae black. Malar space at least about as 
long as diameter of front ocellus in 9 and two-thirds in <J. 

Claws with only small inner tooth. . . . . . . . n 

8 (7) Claws bifid ............ 9 

Claws with only small inner tooth . . . . . . . . 10 

9 (8) Fleck on mesopleuron, and abdomen with continuous bands on tergites i and 3 

or 4 to 7, as well as all the sternites, yellowish white. Mesonotum with 



HYMENOPTERA FROM TURKEY 131 

shining surface between large widely-spaced punctures. Hind ocelli further 
apart than from nearest eye-margin : POL > OOL. 7-8-5 mm. N. AFRICA 

and ARABIA citrina (Perez) 

Mesopleuron and abdominal sternites all black, and tergites with at most 
continuous yellow band on yth segment. Mesonotum dull with rough surface 
sculpture between the punctures. Hind ocelli as far apart as from nearest 
eye margin : POL = OOL. 8-5-9 mm. PORTUGAL, SPAIN, ALGERIA 

dusmeti (Konow) 

10 (8) Malar space as long as diameter of front ocellus in - and two-thirds as long in 

o*. Pubescence on mesopleura longer than diameter of front ocellus. E. 

MEDITERRANEAN Tforientalis (Konow) 

Malar space only about two-thirds as long as diameter of front ocellus in $ and 
less than half in <J. Pubescence on mesopleura much shorter than diameter 
of front ocellus. SPAIN, ALGERIA ...... jucunda (Klug) 

11 (7) Pubescence on mesonotum and mesopleura only about half as long as diameter 

of an ocellus. Hind ocelli closer to occipital carina than from nearest eye- 
margin . . . . . . . . . . . .12 

Pubescence on mesonotum and mesopleura in part longer than diameter of an 
ocellus. Hind ocelli about as far from occipital carina as from nearest eye- 
margin. 

[POL > OOL. Hind ocelli about twice their own diameter from occipital 
carina.] N. AFRICA, C. and S. EUROPE, TRANSCAUCASUS and UKRAINE 

crassicornis (Rossi) 

12 (n) Clypeo-f rental area entirely black and its medial groove extends only one- 

fourth or one-third of way to antennae. POL = or > OOL. Hypopygium 
($) excised on hind margin each side of middle . . . . . 13 

Clypeo-frontal area marked in front with yellow, and its medial groove extends 
half way to antennae. POL < OOL. Hypopygium ($) only slightly 
emarginate on hind margin each side of middle. 

Hind ocelli about twice their own diameter from occipital carina. E. 
MEDITERRANEAN ........ \jrontina (Konow) 

13 (12) POL > OOL. Hind ocelli twice their own diameter from occipital carina. 

Subcosta of fore wing infuscate except at extreme base. E. MEDITERRANEAN 

lateralis (Brulle) 

POL = OOL. Hind ocelli about one and one-half times their own diameter 
from occipital carina. Subcosta of fore wing brown except at extreme apex. 
SPAIN, N.W. AFRICA ..... mauritanica (Gussakovskii) 

14 (i) Legs partly pale ........... 15 

Legs entirely black ........... 18 

15 (14) Pubescence on mesonotum at least as long as diameter of hind ocellus and up- 

standing. Clypeo-frontal area convex and gibbous above. Malar space 
less than half as long as diameter of front ocellus . . . . . 16 
Pubescence on head and thorax much shorter than diameter of hind ocellus 
and prostrate. Clypeo-frontal area with the middle flattened so that it 
appears as a straight line in profile. Femora mainly black. 7-10 mm. 
Malar space more than half diameter of front ocellus. $ apical tergites 
unmodified. E. MEDITERRANEAN ^similis (Mocsary) 

16 (15) Pubescence on mesonotum and head only about as long as diameter of hind 

ocellus and pubescence on abdomen much shorter than this. Femora black 
only basally. Tergites scarcely arched longitudinally . . 17 

Pubescence on mesonotum, head and abdomen very coarse and about as long 
as 4th antennal segment. Femora mainly black. Tergites strongly arched 
longitudinally. <$ 8th tergite with apical triangle bearing dense fine pubes- 
cence. S.E. EUROPE and E. MEDITERRANEAN . . \kruperi (J. P. E, F. Stein) 



132 R. B. BENSON 

17 (16) Mesopleura very densely punctate without shining interspaces. $ with un- 

modified yth and 8th tergites. E. MEDITERRANEAN . . \reticulata (Benson) 
Mesopleura less densely punctate, so that there are some shining interspaces 
in the middle as large as punctures. $ 7th and 8th tergites with an apical 
medial triangular area bearing dense long fine hairs. ALGERIA \andrei (Konow) 

1 8 (14) Clypeo-frontal area convex above and concave below . . . . . 19 

Clypeo-frontal area flattened medially, slightly concave throughout with raised 
lateral margins. 

$ hypopygium only slightly emarginate laterally. $ 7th tergite with 
medial apical longitudinal line densely pubescent. 5-6 mm. long. S. EUROPE 
and E. MEDITERRANEAN ...... italica (Lepeletier) 

19 (18) Clypeo-frontal area evenly convex above and slightly concave below, showing a 

curved line in profile. Pubescence of head, thorax and abdomen grey. 

(J apical tergites unmodified. 5-5-7 mm. ...... 20 

Clypeo-frontal area strongly convex above, gibbous in shape strongly de- 
pressed below, angular in profile. Pubescence of head, thorax and abdomen 
fuscous, o* 7th tergite with a medial apical triangular area with dense 
pubescence. 4-5-6 mm. long. S.E. EUROPE . . . \atricapilla (Mocsary) 

20 (19) Malar space about as long as diameter of front ocellus. $ hypopygium with 

hind margin excised each side. Punctation on mesonotum coarser and 
more irregular. C. and S. EUROPE and TURKEY . . obscura (Fabricius) 
Malar space not more than one half as long as diameter of front ocellus. $ 
hypopygium only slightly emarginate each side. Punctation on mesonotum 
much finer and more regular. S.E. EUROPE, E. MEDITERRANEAN to N. IRAN 

^caucasica (Mocsary) 

Corynis haematica sp. n. 

Description and comparison with related species is included in the key above. 
ISRAEL. 

\Corynis orientalis (Konow) 
SYRIA, LEBANON, ISRAEL and IRAQ. 

Corynis concinna (]. P. E. F. Stein) 

Amasis sarta Kuznetzov-Ugamskii, syn. n. 
A. bleyli Muche, 1964, syn. n. 

TURKEY, S., C. and E.: Mersin (6). i $. Cankiri, Isik Dag, 1,200 m. i $, 
25. vi. 1966 (Demelt Coll.); Amasya (5, 6 and 7). 17 $; Erzurum (3 and 4). 4 $. 
At flowers of Potentilla hirta. 

S.E. EUROPE, TURKEY and TRANSCAUCASIA. 



Corynis lateralis (Brulle) 

TURKEY, C. and E. : Ankara (41); Amasya (3 and 7); Artvin (2). i <$, 17$. 
S.E. EUROPE, TURKEY, TRANSCAUCASIA and S. URAL. 



HYMENOPTERA FROM TURKEY 133 

^Corynis frontina (Konow) 

TURKEY, S., C., N., N.E. and E. : Mersin (7); Amasya (i, 2, 3, 5 and 7); Ankara 
(41); Samsun (10) ; Artvin (2); Gumusane (14). 39 $, 27 $. 

TURKEY and TRANSCAUCASIA. 

Corynis citrina (Perez) 

ARABIA: Bahrein Island in Persian Gulf, i $, 4.^.1936 (/. Fernandez). 
ALGERIA, TUNISIA, TRIPOLITANIA and ARABIA. 

^Corynis similis (Mocsary) 
IONIAN ISLANDS, CRETE, CYPRUS, SYRIA and ISRAEL. 

^Corynis kruperi (J. P. E. F. Stein) 
Amasis enslini Maidl, syn. n. 

TURKEY, W. and C. : Bursa (3) ; Amasya (14). 3 <$, 3 $. 
BALKANS, TURKEY, CYRENAICA and TRIPOLITANIA. 

*~\Corynis reticulata Benson, 1954 

TURKEY, C. : Nigde, Ulukisla, i $, 16-20. .1955 (Seidenstucker). 
TURKEY and ISRAEL. 

Corynis italica (Lepeletier) 

TURKEY, W. and C.: Bursa (9); Amasya (3, 7 and 9). 8 ^, 8 $. 
S. EUROPE and TURKEY. 

t Corynis caucasica (Mocsary) 

TURKEY, N.W., C. and E. : Edirne (i) ; Amasya (2, 3, 7 and 9) ; Trabzon (3 and 15). 
26 <J, 24 ?. 
S.E. EUROPE, TURKEY, TRANSCAUCASIA and IRAN. 

Corynis obscura (Fabricius) 

TURKEY, N.E., and E.: Trabzon (13); Rize (8); Erzurum (10). i $, i ?. 
EUROPE, TURKEY and TRANSCAUCASIA. 

TENTHREDINIDAE 

The notes on the species of this family are intended for use in comparison with 
the keys given in Benson, 1951-58. 



134 R- B - BENSON 

SELANDRIINAE 

For a recent revision of the Palaearctic species of this subfamily see Zhelochovtsev, 



Thrinax caucasica Schaposchikov 

This species is closely similar to T. mixta Klug, but differs in that the edge of the 
pronotum, tegulae and venation at base of fore wing are red instead of yellowish 
white, that on the legs the coxae and trochanters are black as well as i the femora 
and tibiae, and the frontal area of the head is more transversely wrinkled than in 
mixta. 

TURKEY, N.E.: Trabzon (17). 2 <$, 5 $. 

TURKEY and TRANSCAUCASIA. 

T. macula (Klug) 
C. and N. EUROPE and TRANSCAUCASIA. 

Strongylogaster lineata lineata (Christ) 

TURKEY, N.W., N. and N.E. : Istanbul (2) ; Sinop (2) ; Samsun (19) ; Rize (5). 9 $. 
EUROPE, TURKEY, TRANSCAUCASIA, IRAN, SIBERIA to JAPAN. 

\Strongylogaster lineata cypria Benson, 1954 

This form, only known from Cyprus and Lebanon, has almost entirely yellow 
legs and entirely black antenna and epipygium (the typical subspecies has black- 
based femora, and the two basal antennal segments and the epipygium brown). 

Aneugmenus padi (L.) 
T. coronatus Klug, syn. n. 

The only difference that I can discover between A. padi and A. coronatus is that 
A. coronatus has a small white fleck in the middle of the apical tergites in the -. 
As these two forms occur together over the same range, together with intermediate 
forms which can only be ascribed arbitrarily, it is unlikely that they are specifically 
distinct. 

Only one form of <$ is known and this has a large yellow fleck covering the middle 
of the middle tergites and another covering most of the sternites. The <$ is also 
remarkable in having a sinus sexualis cutting transversely deeply into the 7th tergite. 
The male has occasionally been found in Britain, where however the species is at 
least almost entirely parthenogenetic. In the Mediterranean region, males are 
more numerous than females. 

TURKEY, N.W., S., N. and N.E. : Istanbul (2 and 3) ; Mersin (6) ; Tokat (i) ; Samsun 
(29); Rize (5). I8& 15?- 
LEBANON: Felouka, 5 <$, 17^.1953 (G. A. Mavromoustakis). 

EUROPE, N. AFRICA, TURKEY and SIBERIA (introduced into N. America). 



HYMENOPTERA FROM TURKEY 135 

Aneugmenus oertzeni (Konow) 

This species, which I have never seen, is distinguished from A. padi by its un- 
toothed tarsal claws. From A. fuerstenbergensis it is distinguished in the <$ by 
having the 6th tergite emarginate apically and a transverse groove (sinus sexualis) 
across the yth tergite (as in A. padi) and the tergites yellow from the 3rd segment; 
and in both sexes by the shining surface of the tergites which in A . fuerstenbergensis 
are dull with transverse striations. 

CRETE, TURKEY and S.E. EUROPE. 

Birka cinereipes (Klug) 

Melisandra cinereipes (Klug) Benson. 

TURKEY, N.: Balu (i). i _. 

N. and C. EUROPE, TURKEY and TRANSCAUCASIA. 

Mesoselandria morio (Fabricius) 

Melisandra morio (Fabricius) Benson. 

TURKEY, C., N., and N.E.: Ankara (39); Sinop (4); Trabzon (3, 7 and 9); Rize 

(i and 6). 8$. 

EUROPE, TURKEY, TRANSCAUCASIA and SIBERIA. 

Selandria serva (Fabricius) 
\Selandria serva fuscitar sis Benson, 1954, s y n - n - 

TURKEY, N.W., C., N. and N.E. and E. : Istanbul (3 and 10); Ankara (15, 34, 39 
and 40); Tokat (i and 3); Samsun (5, 7, 10, 20 and 24); Trabzon (8); Gireson (3); 
Erzurum (5). Plentiful, 9.v-6.ix, from altitudes up to 1,300 m. in Karagol lake, 
Ankara. The form with black tarsi does not represent a distinct subspecies as 
numerous intermediates occur. Gramineae and Cyperaceae. 

EUROPE, TURKEY, TRANSCAUCASIA and SIBERIA. 

Brachythops flavens (Klug) 
EUROPE, TRANSCAUCASIA, SIBERIA, N. AMERICA. 

Loderus eversmanni (Kirby) 

TURKEY, N., N.E. and E.: Samsun (5); Trabzon (4); Gumusane (13); Erzurum 
(13). 1,245-2,000 m., 75 $, 15$. Equisetum. 

EUROPE to CAUCASUS and TURKEY, SIBERIA to JAPAN, and N. AMERICA (poly- 
typic species). 



136 R. B. BENSON 

DOLERUS Panzer 
In the list which follows, a key is given to the species of the picipes-ciliatus-group. 

Loderus vestigialis (Klug) 

TURKEY, N.E. and E. : Trabzon (3,4 and 17); Gumusane (10); Rize (2). 500- 
1,800 m., 34 <$, ii $. Equisetum. 
EUROPE to CAUCASUS and SIBERIA to JAPAN and N. AMERICA (polytypic species). 

Dolerus germanicus (Fabricius) 

TURKEY, C., N.E. and E. : Amasya (2) ; Trabzon (7 and 9) ; Rize (5) ; Gumusane (3) ; 
Erzurum (3, 5 and 10). Trabzon and Rize at sea level. 56 <$, 83$, 19. iv.- 
20.iv. 1959; 40^, 24$, 24 . viii . 1959. Erzurum and Gumusane, 1,700-2,000 m. 
19 c, 4$, 3i.v.-i.vi.i962. 

EUROPE and temperate ASIA, (polytypic species). 

The three series from N.E. TURKEY represent the spring and summer flights of 
a double-brooded mountain form. The lowland form would appear to represent 
D. germanicus meridianus Zhelochovtsev, distinguishable from the mountain form, 
the S. European etmscus, through its yellow labrum and, in the summer females, 
through its yellow clypeus also. It is not clear whether the lowland and mountain 
forms are discrete races, kept apart by altitude and differing flying-seasons, or 
whether blending occurs in intermediate altitudes and seasons. Every intergrade 
occurs between the two European forms: germanicus of W. and C. EUROPE with its 
hind legs more extensively black than its front legs and frequently with black marks 
also on front mesonotal lobe and scutellum; and etruscus Klug of C. and S.E. EUROPE 
with its hind legs more extensively yellow than its front legs, and the front mesonotal 
lobe and scutellum usually entirely yellow. 

It would therefore seem very unlikely that the forms D. germanicus germanicus, 
D. germanicus etruscus and D. germanicus meridianus are discrete subspecies. 

\Dolerus melanoptera Konow 

Very closely similar to the preceding but larger (10-11 mm. instead of 7-9 mm.) 
with head more swollen behind eyes, temples more sparsely punctured, with occipital 
groove and carina less developed and hind femur and tibia entirely yellow. The <$ 
in lacking the black flecks on the apical tergites differs from the form of germanicus 
in E. Mediterranean (some <$<$ of germanicus in Italy and Spain lack these 
flecks). 

From D. hispanicus it is distinguished by its partly pale legs and deeply excised 
clypeus (to depth of at least half the total length of clypeus). 

TURKEY, N.W., C. and N. : Edirne (i) ; Ankara (15, 35 and 36) ; Amasya (2 and 7) ; 
Samsun (20). 24 <, 13 $. 

S.E. EUROPE and TURKEY. 



HYMENOPTERA FROM TURKEY 137 

f Dolerus hispanicus Mocsary 

Dolerus geniculatus Lepeletier, 1823, nee Geoffrey, 1785, syn. n. 
\Dolerus nigriceps Konow, 1891, syn. n. 
Dolerus africanus Forsius, 1919, syn. n. 

This species is similar to D. germanicus in general form and sawsheath, but is 
larger, with very sparsely punctured head and mesonotum, with poorly developed 
temporal furrows on head, and the ist tergite and sawsheath in the $ are pale, and 
the legs entirely black. From both germanicus and mdanoptera it is distinguished 
by its less deeply excised clypeus (less than half the total clypeal length). The <, 
like the -, has the mesonotum mainly yellow and the apex of the abdomen un- 
marked with black. The palest forms occur in Algeria, where the whole thorax 
and abdomen above, including ist tergite in both $ and $, can be yellow, and only 
the meso- and metasternum black, but usually the scutellum and the sunken sur- 
rounding parts are i black. 

TURKEY, C.: Amasya, 500 m., i $, 2$, 22.v.-6.vi.i959 (Guichard & Harvey}. 
N.W. AFRICA, SPAIN, TURKEY, TRANSCAUCASIA and N. IRAN. 

Dolerus aericeps C. G. Thomson 
EUROPE and TRANSCAUCASIA. 

*Dolerus anticus seljuki ssp. n. 

In the $, this race is distinguished from the typical race in that the lateral mesonotal lobes 
are entirely reddish yellow except where they meet medially. In colour therefore these speci- 
mens resemble D. madidus f. schulthessi except that the mesopleuron is entirely black. In 
form of saw, sawsheath with curved setae, and antenna with 8th segment only twice as long as 
wide, they are clearly distinguished from D. madidus. The <$ is distinguished from typical 
D. anticus by having the 2nd and yth tergites unmarked with black. 

Holotype $. E. TURKEY: Gumusane, Bayburt, 1,600 m., 26^.1962 (Guichard 6- 
Harvey}; B.M. (N.H.). 

Paratypes. TURKEY: N.W., N., N.E. and E. : Edirne, i $, 6. v. 1960 (Guichard 
6- Harvey); Samsun, Lake Ladig, 800 m., i $, 26.vii.i959 (Guichard); Erzurum, 
Ovacik, 2,000 m., i , 30^.1962 (Guichard & Harvey); Gumusane, Bayburt, 
1,600 m., i $, 26. v. 1962 (Guichard & Harvey). B.M. (N.H.). 

The species not previously recorded outside Europe. 

Dolerus triplicatus triplicatus (Klug) 

This subspecies is distinguished from steini thus: (i) 3 colour pattern is different 
from the $, the thorax and ist tergite being entirely black; and (2) $, though similar 
to steini, in having a reddish yellow thorax with a black fleck on each of the mesonotal 
lobes and a large fleck on the underthorax, covering the mesosternum and much of 
the mesopleura. This fleck does not reach the front edge of the mesepisternum, and 



138 R. B. BENSON 

the depressed lateral parts of the mesonotum are not infuscate. This subspecies 
would appear to be the continental part of an atlantic /continental subspecies pair 
which it forms with steini: these two subspecies overlap in TURKEY. 

TURKEY, N.W., C. and E. : Edirne (i) ; Ankara (13) ; Gumusane (13) ; Erzurum (7) ; 
1,000 to 1,800 m., 43 <$, 7 $. 

SWEDEN, GERMANY, CZECHOSLOVAKIA, AUSTRIA, and N.W., C. and E. TURKEY. 
Dolerus triplicatus steini (Konow, 1885) 

Dolerus triplicatus (Klug) ; Benson, 1952 nee Klug. 
Dolerus triplicatus steini (Konow) ; Benson, 1966. 

This subspecies is the atlantic counterpart of an atlantic-continental subspecies 
pair which it forms with the preceding subspecies. It is distinguished by the <$ 
and $ having the same colour pattern, which differs from that of the triplicatus $ 
in that the black fleck on the underthorax reaches right to the front of the mesepisternum 
and that depressed lateral parts of the mesonotum are i infuscate (see fig. 4 in 
Benson, 1966). 

TURKEY, N.E. : Trabzon, Soganli Gecidi, 2,600 m., i <$, 27. vi. 1962 (Guichard & 
Harvey}; Rize, Sivrikaya, 1,700 m., i $, 3^.1962 (Guichard & Harvey}. 
ENGLAND, SWITZERLAND, ITALY and N.E. TURKEY. 

Dolerus puncticollis C. G. Thomson 

TURKEY, N. andE. : Samsun (12 and 15). 3 $, 31 -, 1,600-2,390 m., 2-11.^.1959. 
Rize (8); Gumusane (3, 10); Erzurum (5, 12 and 14). 26 <$, 25 $, at sea level, 23. v 
to 3.vi.i962. 

EUROPE, ISRAEL and TURKEY. 

[Dolerus gonager (F.) 
(Text-fig. 7) 

TURKEY, N.W. : Istanbul, Belgrat Orman, i , 23.^.1962 (Guichard & Harvey}. 
EUROPE]. 

*Dolerus montivagus sp. n. 

(Text-figs. 4, 9) 

$. Black except for the red knees (the extreme apex of all the femora and extreme base of 
all the tibiae) . Pubescence silvery. Wings subhyaline with black venation. Length : 9-10 mm. 

Head : subparallel-sided behind the eyes, which are separated above from the occipital margin 
by about their own length. Malar space about one-third of inter- an tennal line. Clypeus 
broadly and shallowly emarginate to a depth of only about one-fifth of the clypeal length. 
Antenna with 8th segment about x 5 as long as basal breadth and 3rd segment longer than 
greatest measure of an eye as (i-o : 09). Punctation coarse with several large shining inter- 
spaces on temples and notably a large spot as large as 2 or 3 ocelli adjoining each of the lateral 
ocelli. Occipital furrows not very clearly marked above. 



HYMENOPTERA FROM TURKEY 139 

Thorax: Mesonotum coarsely and rather irregularly punctured but shining between the 
punctures ; large sparsely punctate shining areas occur each side of the middle line of the front 
lobes, most of the front and middle of the lateral lobes and the front and middle of the scutellum ; 
post-tergite of scutellum shining without punctures or microsculpture and longer than the 
shortest measure of a cencher. Mesopleuron coarsely and densely punctured, without inter- 
spaces as large as a puncture. Mesosternum with only very shallow fine scattered punctures. 
Legs normal, with hind tarsus about five-sixths as long as a tibia. Wings normal. 

Abdomen: Normal but with transversely alutaceous surface sculpture obsolescent on the ist 
and middle of the 2nd tergite. Sawsheath in dorsal view long and narrow, but with apical 
setae much as in D. puncticollis (Text-fig. 4). 

Pubescence silvery all over and up to about as long as x 2 diameter of an ocellus on head and 
thorax, obsolete on the 2nd, 3rd and 4th tergites and middle of ist tergite. 

cJ as $ but head contracted slightly behind eyes, which are separated above from the occipital 
margin by little more than their own shortest measure, 8th antennal segment only about 
X 3-5 as long as its basal breadth. Abdomen more pubescent on ist and middle of 2nd and 
3rd tergites, the tergites beyond 4th lacking even the mid-dorsal base line; penis -valve (Text- 
fig. 9)- 

Holotype $. N.E. TURKEY: Trabzon, Soganli Gecidi, 2,600 m., 26. v. 1962 
(Guichard 6- Harvey). B.M. (N.H.). 

Paratypes. N.E. TURKEY: 3 <$ (with same data as holotype); Trabzon, Zigana 
Gecidi, 1,650 m., 13$, 22. v. 1962; Trabzon, Hamsikoy, 1,245 m., i ^, i $, 23- 
24. v. 1962; Rize, Sivrikaya, 1,700 m., i $, i $, 3. 1.1962; and Erzurum, Tortum, 
1,550 m., i $, 10. vi. 1962 (Guichard & Harvey}. Total 4$, 18 <. B.M. (N.H.). 

This species has the sparse punctation of D. gonager on the mesonotum, but the 
narrow sawsheath bears setae more like those of D. puncticollis: from both these 
species though it differs in having a scarcely emarginate clypeus, longer antennae 
in $ (in neither D. gonager nor D. puncticollis $ is the 3rd antennal segment clearly 
longer than an eye), and in <$, absence of bare hairless mid-dorsal line on abdomen 
from 4th tergite, apart from differences in the penis-valve. 



THE PICIPES-CILIATUS-GROUP 

In S.E. EUROPE, ASIA MINOR and TRANSCAUCASIA there occurs a group of species 
related to the two last species (Dolerus gonager F. and puncticollis Thomson) but 
in which the female sawsheath is sharply expanded behind in dorsal view (see 
Benson, 1951-58, fig. 198), the legs are often red about the knees (the joint between 
the femur and the tibia) and the temples have an impunctate spot c. X 3 diameter 
of an ocellus adjoining the postocellar area each side. The males are under 9 mm. 
and have: (i) either the lateral mesonotal lobe punctate at least behind, or infuscate 
pubescence on the head, and (2) either very densely pubescent tergites without a 
bare medial line or a clypeus with a much-reduced medial front marginal excision 
(to less than one-third of total clypeal length). In three of the species the females 
are brachypterous with correlated changes in the mesonotum (flattening and 
shortening of lobes with obsolescence of suture dividing front pair and shortening 
of post-tergite of scutellum). 



140 R. B. BENSON 

KEY TO MALES AND FEMALES 

1 Wings hyaline and fully-developed in $ and $. Clypeus apicMly broadly 

emarginate to a depth of at least one-third of its total length, with the lateral 
lobes narrower than the medial emargination. Pubescence on head and 
thorax silvery ........... 2 

Wings infuscate and often brachypterous in $. Either clypeus with a narrow 
and shallow excision apically (to a depth not exceeding one-third of its total 
length and leaving obtuse or truncate lateral lobes) or pubescence on head 
and mesonotum infuscate ......... 3 

2 (i) Legs not red about the knees (joint between femur and tibia). Head behind the 

eyes slightly swollen, longer than shortest measure of eye and with occipital 
furrows obsolescent. $ abdomen lacks the medial bare line through the 
pubescence on tergites 3-8; penis-valve (Benson, 1951-58, fig. 240). Larger 
species : 7-9 mm. EUROPE ....... picipes Klug 

Legs red about the knees. Head slightly contracted behind eyes, not longer 
than shortest eye-measure and with occipital furrows clearly developed. 
<J abdomen with medial bare line through pubescence on tergites 3-8; penis- 
valve (Text-fig. 5). Smaller species: 7-8 mm. IRAN . hyrcanus sp. n. 

[Dolerus vernalis Ermolenko, 1964 (unique $) (Ukraine) would run here 
and may be synonymous, but has red on front and middle knees limited to 
apex of femur.] 

3 (i) Either clypeus excised medially in front to a depth less the diameter of the 

front ocellus, or pubescence on head and wings silvery. $ i brachypterous 4 

Clypeus broadly emarginate in front to a depth of almost one-third of its total 
length. Upper head and wings with infuscate pubescence. Wings fully- 
formed in both sexes. 

Legs red on the knees. Penis- valve (Text-fig. 6). S.E. EUROPE and 
TRANSCAUCASIA kokujewi Konow 

4 (3) $ wings less than one-half as long as abdomen. At least $ with infuscate 

pubescence on upper head and wings ....... 5 

$ wings much more than one-half as long as abdomen. Pubescence on head 

and wings silvery in $ and -. S.E. EUROPE . . subalatus Kirensky, 1926 

5 (4) Legs entirely black. $ with infuscate pubescence on head and wings. Pubes- 

cence on mesopleura shorter than on head and mesonotum, where it is as 
long as 2nd antennal segment. GREECE (Mt. Parnassus, 2,000 m.) (<$ un- 
known) .......... oertzeni Konow 

Legs with red knees. <J, but not $, with infuscate pubescence on head and 
wings; $ with pubescence on mesopleura as long as that on head and meso- 
notum, where it is about as long as 2nd antennal segment. S.E. EUROPE 
and TRANSCAUCASIA ........ ciliatus Konow 

Dolerus picipes Klug 
EUROPE, TURKEY and TRANSCAUCASIA. 

Dolerus kokujewi Konow 
(Text-fig. 6) 

TURKEY, N.E. : Trabzon, Zigana Gecidi, 1,650 m., i <$, 2. v. 1962 (Guichard & 
Harvey). 

S.E. EUROPE, TURKEY and TRANSCAUCASIA. 



HYMENOPTERA FROM TURKEY 141 

Dolerus hyrcanus sp. n. 

(Text-figs. 3, 5) 

<J. Black with the knees red (the apical half of the femur and basal fourth of the tibia) ; 
pubescence silvery; wings hyaline with black venation. Length 8-9 mm. 

Head: slightly contracted behind eyes which are separated above from the occipital margin 
by about their own shortest measure. Malar space about one-half of inter-antennal line. 
Clypeus broadly emarginate in front to a depth of about one-third of its total length and strongly 
arched transversely. Antenna with 8th segment about x 4 as long as its basal breadth. 
Punctation coarse and without interspaces in front of ocelli but the temples adjoining the post- 
ocellar region each side have an impunctate area as large as 2 to 3 ocelli. Occipital furrows 
clearly marked. 

Thorax : front lobes of mesonotum coarsely punctured laterally without interspaces but with 
the middle half shining with shallow sparse punctures ; middle lobes shining with sparse shallow 
punctures on the front half, widely spaced becoming denser behind; scutellum densely punc- 
tured except in front and along middle line, and with its post-tergite longer than the shortest 
length of a cencher and obliquely striated with rugulae. Mesopleura coarsely punctured but 
with a few impunctate spots in the middle up to about twice the diameter of a puncture. Meso- 
sternum with only very shallow fine scattered punctures. Legs normal with hind tarsus about 
five-sixths as long as tibia. Wings normal. 

Abdomen normal with tergites transversely alutaceous; penis-valve as in Text-fig. 5. 

Pubescence silvery all over ; on head and thorax up to about x 2 diameters of ocellus ; abdomen 
with tergites 1-3 glabrous laterally, rest of tergites densely clothed except for base medial line. 

? as 6* except as follows : 8th antennal segment only x 3 as long as broad at base ; hind tarsus 
only about two-thirds as long as tibia; ovipositor longer than hind femur; sawsheath in dorsal 
view incrassate apically, where it is twice as broad as at base (Text-fig. 3). Saw as in D. picipes. 

Holotype $. IRAN: Mazandaran, Chalus-Chahsavar coast, 23.iii.ig66 (D. B. 
Baker}. B.M. (N.H.). 

Paratypes. Same locality, 19 $, 3 ?, 25. 11-28. iii (D. B. Baker}. B.M. (N.H.). 

The $ is distinguished from other species of the picipes-ciliatus group in the key 
above. The $ penis-valve (Text-fig. 5) is unlike that of any other known species 
but is perhaps most like that of D. puncticollis (cf. Benson, 1951-58, fig. 230). 

Dolerus nigratus (Miiller) 
EUROPE and TURKEY (Bytinski-Salz). 

Dolerus aeneus Hartig 

TURKEY: Trabzon, Soganli Gecidi, 2,600 m., i <$, 27^.1962 (Guichard & Harvey}; 
Trabzon (17). i $. 
EUROPE and ASIA MINOR. 

Dolerus asper Zaddach 
Dolerus asper megapteroides Muche $, ig64a (nee g) syn. n. 

TURKEY, E.: Gumusane (n); Erzurum (7, 12 and 13). 1,800 m., 2,390 m., 5 <J, 
i $, 29 . v . to i . vi . 1962 ; Bolu (Muche) . 

ENTOM. 22, 4. 6 



R. B. BENSON 





FIGS. 1-9. i, 2. Penis-valves: i, Arge melanochroa; and 2, A. scita. 3, 4. Sawsheaths 
from above: 3, Dolerus hyrcanus sp. n.; 4, Z). montivagus sp. n. 5-9. Penis- val ves : 
5, Dolerus hyrcanus sp. n. ; 6, D. kokujewi; 7, Z). gonager; 8, D. nonutimus; and 9, 
>. montivagus sp. n. 

The specimens from Gumusane and Erzerum have more infuscate wings than 
any specimens seen from Europe. 

Species from EUROPE, TURKEY and temperate ASIA to KAMTCHATKA. 

Dolerus haematodes Schrank. 

TURKEY, N., N.E. and E. : Samsun (5, 12 and 15) ; Trabzon (7 and 17) ; Gumusane 
(3). Sea level to 1,600 m., 80 $, 53 ?. 



HYMENOPTERA FROM TURKEY I43 

In the series from sea level at Samsun, 12 out of 23 < have the female colouring, 
with red pronotum and tegulae; in the series from Gumusane, at 1,600 m., only i 
out of 58 <$ have this colouring while 57 J, as in typical W. European form, lack 
any red on pronotum and tegulae. 

EUROPE, TURKEY and TRANSCAUCASIA. 



*Dolerus nonultimus Zhelochovtsev, 1941 (with figs.) 
TURKEY: Trabzon (4). i $. 

This species belongs to the anthracinus-nitens group with sawsheath as in Benson, 
1951-58, fig. 196, but has a distinct penis-valve and saw. The male has infuscate 
pubescence on the upper head but differs from anthracinus in having the head strongly 
contracted behind. The female differs from other known females in the group in 
having infuscate pubescence on its head as in the male, and from anthracinus and 
cor acinus differs also in having unmodified front mesonotal lobes. 

TURKEY and TRANSCAUCASIA. 

*^Dolerus megapterus Cameron 

? Dolerus asper megapteroides Muche, ig6^a, ($ nee $). 

TURKEY: Gumusane (3). 2 <$. G. Ovacik, 2,000 m., 4 ^, i 9, 30^.1962 (Guichard 
& Harvey); Erzurum (5). i $. 

Not previously recorded from outside EUROPE. 

BLENNOCAMPINAE 

ATHALIINI 

Tribe revised by Benson, 1962. 

\Athalia cuspidata Benson, 1954 
ISRAEL, endemic. 

^Athalia dimidiata Konow 
TURKEY, endemic. 

Athalia bicolor Lepeletier 

TURKEY, N.W., W., C., N., N.E., and E.: Edirne (i); Bursa (4 and 9); Usak 
(i) ; Bilecik (2) ; Samsun (5, 6 and 16) ; Trabzon (3 and 4) ; Artvin (4) ; Erzurum 
(7). 29 , 2 ?. 

EUROPE, MEDITERRANEAN, TURKEY, TRANSCAUCASIA to TURKMEN REPUBLIC. 



144 R- B. BENSON 

f Athalia maculata Mocsary 

TURKEY, N. and C. : Bolu (2); Samsun (30); Amasya (9). 3 
S.E. EUROPE, TURKEY and TRANSCAUCASIA. 



\Athalia paveli Mocsary 
TURKEY, endemic. 

^Athalia rufoscutellata Mocsary 

TURKEY, W., C., N.E. and E. : Bursa (3, 9); Ankara (n); Corum (2); Samsun (2); 
Artvin (4); Gumusane (n). 5 ^, 7 $. 

C. and S.E. EUROPE, TURKEY and TRANSCAUCASIA. 



f Athalia glabricollis glabricollis C. G. Thomson 

TURKEY, W., S.W., C., N.E. and E. : Bursa (3); Mugla (7 and 8); Ankara (3, n, 
14 and 63); Amasya (12); Trabzon (9); Erzurum (i); Kars (n). 12 J, 15 $. 
All EUROPE, MEDITERRANEAN and TURKEY. 



f Athalia glabricollis meridiana Benson, 1954 

This subspecies is distinguished from the preceding in having more than half the 
Costa from the base yellow in the fore wing. 

It is known only from ISRAEL and S.W. IRAN (Benson, 1962, was wrong in ascribing 
material from GREECE and TURKEY to this form). 

Athalia ahngeri Kokujev 

TURKEY, C. and E. : Nigde (5); Gumusane (15). 2 $. 

Steppe species. ISRAEL, TURKEY, TRANSCAUCASIA and MESOPOTAMIA to TURK- 
MEN, UZBEK REPUBLICS and AFGHANISTAN. 

Athalia circularis circularis Klug 

TURKEY, N.W., W., S., N., C. and N.E.: Istanbul (3 and 9); Mugla (7); Bolu (3); 
Zonguldak (i) ; Mersin (7) ; Sinop (4) ; Samsun (5, 10, 19 and 21) ; Corum (2) ; Amasya 
(3 and 5) ; Tokat (i) ; Trabzon (14 and 15) ; Artvin (6) ; Rize (6). 21 <$, 2 $. 

Females of the summer flight in the mountains and to a lesser extent those of 
the autumn flight at sea level have the scutellum and sides of the mesonotal lobes 
i yellow (e.g., from Samsun, Kundaz Ovacik, at 1,300 m., 22.vii.i959 and Samsun, 
Engiz at sea level, 22. ix. 1960). 



HYMENOPTERA FROM TURKEY 145 

Typical form from EUROPE and MEDITERRANEAN ; replaced by ssp. melanoptera 
Benson in E. SIBERIA, JAPAN and mountains of C. ASIA (Kopet Dag and 
Afghanistan to the Pamirs, Altai and W. Himalayas). 

Athalia cordata Lepeletier 

TURKEY, W., S.W., N., N.E. and E.: Kutahya (5); Bursa (3); Aydin (2); Mugla 
(i and 9); Bom (3); Samsun (15); Giresun (7); Trabzon (3, 4 and 14); Rize (3); 
Kars (i and 9). 15 $, 15 ?. 

EUROPE, MEDITERRANEAN and TURKEY. 



f Athalia rosae rosae (L.) 

TURKEY, N.W., W., S., N., N.E. and E.: Istanbul (3); Kutahya (3 and 10) ; 
Mersin (2); Adana (3); Antakya (6); Samsun (5, 6, 19, 20 and 21); Trabzon (9); 
Rize (i);Gumusane (4); Kars (i and 7). 8 <$, 12$. 

EUROPE, MEDITERRANEAN, ISRAEL to IRAN; S.W. SIBERIA and N.W. CHINA. 
Replaced by ssp. ruficornis in E. ASIA. 

Athalia liberta liberta Klug 

TURKEY, N.W., S.W., N., C., N.E. and E.: Istanbul (3); Aydin (i) ; Samsun (10); 
Amasya (5 and 7); Trabzon (9 and 14); Gumusane (13); Erzurum (10). n <$, 3 $. 

EUROPE and MEDITERRANEAN to S.W. IRAN, Mountains of C. ASIA to UZBEK, 
KIRGHIZ and the AMUR region of E. ASIA; replaced by ssp. yanoi in Japan. 

*f Athalia cornubiae Benson 

TURKEY, N. and N.E. : Bolu (i) ; Trabzon (7 and 15). Sea level to 2,000 m. 3 $. 
EUROPE and MEDITERRANEAN to IRAN. 

t Athalia paradoxa Konow 

TURKEY, W. and E.: Bursa (3); Gumusane (3); Erzurum (3). 3 $. 
Mountains of C. EUROPE (France, Switzerland and Austria), MACEDONIA and 
TURKEY. 

EMPRIINI 

*Harpiphorus lepidus (Klug) 

TURKEY, E. : Erzurum (5). i $, in Quercus scrub. 
Not previously recorded outside EUROPE. 



146 R. B. BENSON 

Monostegia abdominalis (F.) 

TURKEY, N., C. and N.E.: Samsun (10, n and 21); Ankara (n and 33); Amasya 
(i and 7); Tokat (i); Trabzon (4). 20 $. 

EUROPE to SPAIN and CAUCASUS, ISRAEL, TURKEY, TRANSCAUCASIA, SIBERIA and 
Atlantic coast of N. AMERICA. 

*Monsoma pulverata (Retzius) 

TURKEY, N.E. : Trabzon (4). i 9; Rize (5). i ^; Rize (2). i <j>. 
Not recorded previously outside EUROPE. 

Empria archangelskii Dovnar-Zapolskii 
CYPRUS, TURKEY and TRANSCAUCASIA. 

Empria pravei Dovnar-Zapolskii 

N. IRAN: Mazandaran, Chalus-Shahasavar coast of Caspian, i $, io.iv.ig67 
(D. B. Baker). 

N. CAUCASUS, TRANSCAUCASIA and N. IRAN. 



ia klugii Stephens 

TURKEY, N.E.: Trabzon (17); Rize (2). 39- 
EUROPE, TURKEY and TRANSCAUCASIA. 

^Empria excisa Thomson 

TURKEY, N.: Samsun (2). i <. 
EUROPE and TURKEY. 

Empria baltica Conde 
EUROPE and TRANSCAUCASIA. 

Empria liturata (Gmelin) 

TURKEY, N.E. : Rize (2). i <. 

EUROPE, TURKEY and SIBERIA to Irkutsk. 

Empria tridens (Konow) 
TURKEY, N.E. : Giresun (i); Trabzon (3). 2 <j>. 
EUROPE and TURKEY. 

Empria konowi Dovnar-Zapolskii 
UKRAINE and TRANSCAUCASIA. 



HYMENOPTERA FROM TURKEY 



147 



Ametastegia equiseti (Fallen) 

TURKEY, S., N., C., N.E. and E. : Mersin (7); Samsun (21); Amasya (9); Trabzon 
(3); Rize (i and 6); Erzurum (7). 7 <$, 4 . 

EUROPE, MEDITERRANEAN, TURKEY, SIBERIA to N. AMERICA. 

Protemphytus pallipes (Spinola) 

TURKEY, N. and N.E.: Samsun (6); Trabzon (7). 3 $. 

EUROPE to ICELAND, MEDITERRANEAN, TURKEY to IRAN, SIBERIA, N. AMERICA. 

Protemphytus carpini (Hartig) 
EUROPE, MEDITERRANEAN, TURKEY, TRANSCAUCASIA and SIBERIA. 

Protemphytes tener (Fallen) 
EUROPE, MEDITERRANEAN, TURKEY, TRANSCAUCASIA, SIBERIA to N. AMERICA. 

ALLANTINI 

Taxonus agrorum (Fallen) 

TURKEY, N.E. : Artvin, above Artvin, 700 m., i ^, i $, 2.vi.ig62 (Guichard & 
Harvey}. Not previously recorded outside EUROPE. 

Taxonus sticticus (Klug) 
C. and S.E. EUROPE and TRANSCAUCASIA. 

Allantus togatus Panzer 

TURKEY, E. : Erzurum (10). i <$. 
EUROPE, TURKEY, SIBERIA to JAPAN. 

Allantus viennensis (Schrank) 
C. and S. EUROPE and TRANSCAUCASIA. 

Allantus calceatus (Klug) 
EUROPE and TRANSCAUCASIA. 

Allantus basalts (Klug) 

TURKEY, W. and N. : Bursa (9) ; Samsun (6). i <$, 2 J. 
N. and C. EUROPE, TURKEY and SIBERIA. 



148 R. B. BENSON 

Allantus cinctus (L.) 

TURKEY, C. and N.E. : Amasya (7 and 13) ; Gumusane (n). i <$, 2 $. 
EUROPE, TURKEY, TRANSCAUCASIA, SIBERIA to N. AMERICA. 

Allantus cingulatus (Scopoli) 
EUROPE, TRANSCAUCASIA and SIBERIA. 

Allantus didymus (Klug) 

TURKEY, S., N., C. and E. : Mersin (6); Nigde (5); Bolu (2); Ankara (29); Amasya 
(2, 3, 5, 6 and 7) ; Tokat (i) ; Gumusane (10) ; Erzurum (4 and 6). 19 <$, 12 $. N. 
IRAN : Mesanderan, Chalus-Shahsavar coast of Caspian Sea, alt. 20 m., i $, 19 .iv. 1966 
(D. B. Baker}. 

MEDITERRANEAN, TURKEY and IRAN. 

* Allantus melanarius (Klug) 

N. IRAN: Mesandaran, Chalus-Shahsavar coast of Caspian Sea, alt. 20 m., 2$, 
19. .1966 (D. B. Baker). 

All EUROPE and IRAN. 

CALIROINI 

Endelomyia aethiops (F.) 
TURKEY, N. : Samsun (2 and 25) ; 2 $. 
EUROPE, TURKEY, TRANSCAUCASIA and introduced into N. AMERICA. 

Caliroa cerasi (L.) 

TURKEY, N. : Samsun, Engiz, 9 <$, v.i959 (Guichard). 

HOLARCTIC. Obligatorily parthenogenetic races of this species have been intro- 
duced with rosaceous fruit-trees, from its native country, presumably in EURASIA, 
to N. AMERICA and almost all other temperate regions of the world. The occur- 
rence of males in Turkey may be significant in indicating the native country of this 
species, on the theory that introduced parthenogenetic races are descended from 
normally sexual races, native to some part of the world. 

*Caliroa varipes (Klug) 

TURKEY, N.W., S., N. and C.: Istanbul (i, 3 and 8); Tekirdag (2); Mersin (6); 
Kastamonu (i); Samsun (i, 6, 7 and 13); Amasya (13). 12 $, n $. 
All EUROPE and TURKEY, 



HYMENOPTERA FROM TURKEY 149 

Caliroa annulipes (King) 

TURKEY, N.E. : Trabzon (15 and 16). 5 <J, 3$. 
EUROPE, TURKEY and temperate ASIA to E. SIBERIA. 

HETERARTHRINI 

*Heterarthrus vagans (Fallen) 

TURKEY, N.E. : Rize, at sea level, 2$, 12. iv. 1959 (Guichard). 
All EUROPE and TURKEY. 

FENUSINI 

Metallus beckeri (Konow) 

Very close to European M. pumilus Klug, but the hind legs are almost entirely 
pale (pumilus has hind femur mainly infuscate from base) and the individual mar- 
ginal saw teeth have more-rounded projections. In the saw therefore it resembles 
the nearctic M. rohweri MacGillivray ; M. albipes Cameron and M. gei Brischke 
have saws resembling M . pumilus. The <$ antenna is flattened as in pumilus. 

TURKEY, N. and N.E. : Samsun (9) . i <$, 2 $ ; Trabzon, at sea level, i $, 24 . viii . 1959 
(Guichard} ; Rize (5) ; i $. 

Only known from TURKEY. 

Profenusa thomsoni (Konow) 
EUROPE, TRANSCAUCASIA and N. AMERICA (? introduced). 

Profenusa pygmaea (Klug) 

TURKEY, W. and C. : Bilecik (i); Amasya (13). i <J, i $. 
EUROPE and TURKEY. 

Hinatara recta (C. G. Thomson) 
EUROPE and TRANSCAUCASIA. 

Scolioneura hyrcana sp. n. 

cj. Black except for the following, which are brownish to yellowish white: labrum, palps, 
antennal segments 9 and underside of 5 to 8, tarsi, tibiae and apical one-third to one-half of 
femora. Wings hyaline; stigma, and venation black to piceous. Length 4-5 mm. 

Head with mouthparts normal, antenna about X 1-5 breadth of head; 4th segment = 8th 
+ gth = ist x 0-8. Eyes much enlarged and strongly converging in front where the distance 
between them is much less than the longest measure of the eye (i-o : 1-5). Malar space linear. 

Thorax with tarsal pulvilli of legs developed only on two apical tarsomeres. Claws bent 
medially at right angles and with sharp basal tooth. 

Wings normal. 



150 R. B. BENSON 

Abdomen without any visible sculpture. 

Pubescence dense on upper head and mesopleura; on mesopleura up to about as long as dia- 
meter of an ocellus but on head shorter than this. On mesonotum and abdomen sparse. 

Holotype <$. N. IRAN: Mazandaran, Shalus-Shahsavar coast, i8.iv.i966 (D. B. 
Baker}. B.M. (N.H.). 

This species is distinguished at once from S. betuleti Klug and S. tirolensis Enslin 
by its strongly converging eyes, which are much closer together in front than their 
greatest length (in 5. betuleti and tirolensis they are further apart than their greatest 
length, as 1-2 : i-o) and also by the loss of the tarsal pulvilli on the two basal tarso- 
meres of the legs. 

Messa hortulana (Klug) 
EUROPE and TRANSCAUCASIA. 

Fenusa dohrni (Tischbein) 
TURKEY, E.: Rize (i). i $. 
EUROPE, temperate ASIA and N. AMERICA; introduced to S. AFRICA. 

Kaliofenusa ulmi laevinota ssp. n. 

Differs from the typical north holarctic race in that the coriaceous surface sculpture on the 
front and lateral mesonotal lobes is absent and the lobes are entirely smooth. The frontal 
basin is much shallower and the furrow separating the frontal area from the inner orbits is 
more clearly denned. 

Holotype $. TURKEY: Mugla, Ula (Mezarlik) 700 m., on Ulmus, 17.^.1962 
(Guichard & Harvey}. B.M. (N.H.). 

Paratypes. TURKEY: Mugla, Ula (Mezarlik) 700 m.; on Ulmus, i? and 8 <$, 
17. iv. 1962 (Guichard & Harvey} ; Bursa, Bursa-Mudanya Rd., 50 m., i $, 28 .iv. 1962 
(Guichard & Harvey); Corum, Iskilip, 700 m., i $, 9. .1962 (Guichard & Harvey}. 

Species holarctic in distribution. 

LYCAOTINI 

This tribe was revised by Benson, 1966. 

*-fSeljukia tene.br osa Benson, 1966 

TURKEY, S.: Mersin, Gozne 600 m., 6 <$, n $, 3-5^1.1960 (uichard 6- Harvey). 

The representatives of this species were taken in a rocky broken terrain with 
alder-lined streams, 34 kms north of Mersin. 

PHYMATOCERINI 

Phymatocera aterrima (Klug) 
C. and S. EUROPE and TRANSCAUCASIA. 



HYMENOPTERA FROM TURKEY 151 

TOMOSTETHINI 

Tomostethus nigritus claripennis Enslin 

N. AFRICA and TURKEY. 

Typical subspecies throughout EUROPE and across SIBERIA. 

Eutomostethus gagathinus (Klug) 
E. gagathinus meridionalis Benson, 1954, sy 11 - n - 
TURKEY, E. : Erzurum (13). i $. 
All EUROPE, N. AFRICA, CYPRUS, TURKEY, TRANSCAUCASIA and SIBERIA. 

Eutomostethus luteiventris (Klug) 
All EUROPE, TRANSCAUCASIA and N. AMERICA. 

Eutomostethus ephippium vopiscus (Konow) 
TURKEY, E.: Rize (i) 4 ; (2) i <$; (5) 17 <?, 5 ?. 

This subspecies differs from the typical one in that males predominate and that 
whereas the $ has an entirely black body (as in one of the two $ phases of the typical 
subspecies) the $ is more extensively red than in the red phase of the typical sub- 
species: it has the whole thorax red (in the red phase of the typical subspecies the 
mesoscutellum, with its post-tergite and the mesosternum are black). The legs of 
$ vopiscus differ from those of both $ forms of the typical subspecies in having the 
apical third of the femora, all the tibiae except the extreme apex as well as the basal 
tarsal segment and i the apical one yellowish white ; in the males the pale colour is 
brownish. 

The typical subspecies is now found throughout EUROPE and N. AMERICA (? intro- 
duced) and there is a black-bodied form in the HIMALAYAS: ssp. vopiscus is re- 
corded only from TURKEY and TRANSCAUCASIA. 

Stethomostus fuliginosis (Schrank) 

TURKEY, C. and N.E. : Bolu (3) ; Ankara (4) ; and Rize (i). 4 $, i ?. 
EUROPE, TURKEY, SIBERIA, to JAPAN and N. AMERICA. 

Stethomostus funereus (Klug) 
TURKEY, N.E. : Rize (i). 3^. 
EUROPE, TURKEY and SIBERIA. 

BLENNOCAMPINI 

Monophadnus spinolae (Klug) 
TURKEY, N.W.: Edirne (i). i ?. 
EUROPE and TURKEY. 



152 R. B. BENSON 

Monophadnus fulviventris athalioides Jakovlev stat. n. 

The only known difference between M. athalioides $ and fulviventris Scop. $ is in 
the colour of the thorax; the underthorax is all black in fulviventris, and in athalioides 
with the upper mesopleuron red. The mesonotum, apart from the postergite of 
the scutellum, is all red in fulviventris but is black-flecked in athalioides. They are 
probably best treated as geographical subspecies. M. f. fulviventris is restricted to 
C. and S. EUROPE and M. f. athalioides to TRANSCAUCASIA and CHINA. 

Monophadnus longicornis Hartig 

TURKEY, N.: Samsun (12, 14 and 15). 5 <$, 5 $. 
EUROPE and TURKEY. 

Monophadnus pallescens (Gmelin) 

TURKEY, N.E. : Trabzon (4 and 17); Rize (2 and 5). n $. 
Temperate EURASIA and N. AMERICA. 

PERICLISTA Konow 

Apericlista Enslin 

Cornaria Malaise 1964, syn. n. 

KEY TO EUROPEAN MALES AND FEMALES 

1 Abdomen mainly pale beneath. Hind wing with anal cell sessile or subsessile 

(peduncle not more than one-half as long as width of anal cell). Clypeus 
medially emarginate. Ovipositor not longer than hind tibia ... 2 

Either abdomen mainly dark beneath or anal cell in hind wing with peduncle 
longer than one-half width of anal cell, or clypeus subtruncate, or ovipositor 
longer than hind tibia .......... 4 

2 (i) with abdomen mainly yellowish above and hind wing with marginal vein; 

$ with head black above and hind wing often with enclosed middle cell . 3 

<$ with body entirely black above except for white tegula and apical margins 
of tergites, and hind wing without marginal vein ; $ with head mainly reddish 
yellow above and without enclosed middle cell in hind wing. MEDITER- 
RANEAN (MOROCCO, and PORTUGAL to BALKANS) . . . andrei (Konow) 

3 (2) Antenna about as long as head + thorax; $ with mesonotum mainly red. 

C. EUROPE to SPAIN and CAUCASUS ..... albida (Klug) 

Antenna longer than head + thorax; $ mesonotum all black. (<$ not seen). 

C. EUROPE ......... albiventris (Klug) 

4 (i) Anal cell of hind wing with peduncle longer than one-half of width of cell. 

Hind wing with or without enclosed middle cell in $ or marginal vein in <$. 
Ovipositor longer than hind tibia ........ 5 

Anal cell of hind wing subsessile (peduncle less than one-half of width of cell). 
Hind wing with enclosed middle cell in - and marginal vein in <$. Ovipositor 
shorter than hind tibia. Legs with femora black. SPAIN and PORTUGAL 

dusmeti (Konow) 

5 (4) Clypeus emarginate medially. $ hind wing with enclosed middle cell, and 

$ with marginal vein. Malar space at least two-thirds as long as diameter 

of ocellus in and $ .......... 6 



HYMENOPTERA FROM TURKEY 153 

Clypeus evenly truncate. $ hind wing without enclosed middle cell and <$ 
without marginal vein. Malar space less than one-half of diameter of 
ocellus in $ and linear in <J. Body entirely black except for white-margined 
pronotum and abdominal segments. C. EUROPE . . albipennis (Zaddach) 

6 (5) Head and thorax with only normal pubescence on middle of mesonotum, not 

longer than diameter of ocellus ........ 7 

Head and thorax clothed in woolly pubescence on mesonotum up to x 2 
diameter of ocellus. Abdomen black with pale apical margins to segments and, 
in $, with orange lateral band on down-turned lateral portions of tergites. 
Legs in <$ yellowish with black coxae and bases of femora, but in $ femora 
mainly yellow with only extreme bases infuscate. C. EUROPE 

pubescens (Zaddach) 

7 (6) Legs in both <J and $ mainly yellow except only for coxae and extreme bases 

of femora. Abdomen in $ infuscate below but yellow above; $ mainly 
black with pale apical margins to segments. AUSTRIA and TURKEY lent a Konow 
Legs in both $ and $ with mainly black femora and i pale tibiae. Abdomen 
in both o* and $ mainly black with pale apical margins to segments. C. 
EUROPE .......... lineolata (Klug) 

Periclista lenta Konow 

TURKEY: Mugla, Kestep, 50 m., 3 ^, 12. iv. 1962 (Guichard & Harvey}. 
TURKEY [type locality Brussa]. 

Monardis plana (Klug) 
EUROPE and TRANSCAUCASIA. 

Ardis brunniventris (Hartig) 
EUROPE, TURKEY, SIBERIA to JAPAN and N. AMERICA. 



Cladardis elongatula (Klug) 
EUROPE and TURKEY. 

Claremontia tenuicornis (Klug) Ross, 1951 

Blennocampa tenuicornis (Klug) Konow. 
Pseudoblennocampa tenuicornis (Klug) Malaise, 1935. 
Monophadnoides tenuicornis (Klug) Benson, 1952. 

It seems to me reasonable to separate Claremontia Rohwer, 1909, in which the 
hind-orbits have a deep coriaceous groove and the hind wing has no enclosed middle 
cells, from Monophadnoides, in which the deep coriaceous hind-orbital groove is 
absent and the hind wings have normally an enclosed middle cell M. To Clare- 
montia also belong alternipes Klug, comb. n. puncticeps Konow, comb. n. confusa 
Konow, comb. n. and the following species. 

EUROPE, TURKEY and SIBERIA. 



154 R- B. BENSON 

Claremontia waldheimii (Gimmerthal) comb. n. 
EUROPE and SIBERIA. 

*Monophadnoides ruficruris (Brulle) 

TURKEY, N.E. : Rize (5). i <$. 

Males of this species in Britain have some of the middle tergites i marked with 
yellow. The $ from Turkey has tergites 3-5 and 2 an d 6, and sternites 2-6 
yellow. This may indicate a distinct race but more specimens would be needed to 
demonstrate this. 

Not previously recorded outside EUROPE. 

Halidamia affinis (Fallen) 

TURKEY: Amasya (5). i $. 

EUROPE, TURKEY, TRANSCAUCASIA and N. AMERICA. 

Pareophora putnilio Konow 

N. IRAN: Mazandaran, Chalus-Shahasavar coast of Caspian, i $, 28.iii.i966 
(D. B. Baker}. 

S. and S.W. CASPIAN coast. [Type locality AZERBAYDZHAN, S.S.R., Lenkoran], 

TENTHREDINIDINAE 
ERIOCAMPINI 

*Eriocampa ovata ovata (L.) 

TURKEY, E.: Tokat (i). 3$. 

EUROPE, not previously recorded from TURKEY, introduced into N. AMERICA. 

*Eriocampa ovata nitens ssp. n. 

A series of this species from E. TURKEY lacks the transverse sculpture on the 
abdomen. 

Holotype $. TURKEY: Rize, Cayeli, 15 m., 22.viii.i959 (Guichard). B.M. (N.H.). 

Paratypes. TURKEY: Rize, Cayeli, 15 m., 19$, 22.viii.i959 (Guichard); Rize, 
Rize, 15 m., i $, 2i.viii.i959; Trabzon, Trabzon, at sea level, i $, 24.viii.i959 
(Guichard). B.M. (N.H.). 

The absence of transverse sculpture on the abdomen occurs as a rare aberration 
in the typical race of this species (e.g. SWEDEN: Skane, Kivik, i $, I5.vii.i938 
(D. M. S. 6- /. F. Perkins)), and is also characteristic of certain other species in the 
genus such as E. dorpatica Konow (E. EUROPE) and E. peineae Zirngiebl (!RAN) 
it is possible that the latter is another form of E. ovata. 



IRAN. 



HYMENOPTERA FROM TURKEY 155 

Eriocampa peineae Zirngiebl, 1956 



*Eriocampa umbratica (Klug) 

TURKEY, N.E. : Trabzon (3, 4 and 16) ; Rize (2, 3 and 5) ; Artvin (4). 38 $, 51 ?, 
Not previously recorded from outside EUROPE. 

PERINEURINI 

Aglaostigma langei (Konow) 
C. EUROPE and TRANSCAUCASIA. 

* Aglaostigma aucupariae lacteore ssp. n. 

<J differs from typical race in that the inter-antennal area and the whole face below the 
antennal sockets (except for the anterior tentorial pits) are yellowish white and this colour is 
continued along the edge of the inner orbits. (In the typical race most of the clypeus, the 
inter-antennal area, and the face below the antenna are black.) 

The $ is scarcely distinguishable from that of the typical race except that the costa of the 
fore wing is paler apically. 

Holotype <. TURKEY: Trabzon, Zigana Gecidi, 1,650 m., 22. .1962 (Guichard & 
Harvey}. B.M. (N.H.). 

Paratypes. TURKEY: Bursa, Uludag, 500 m., i <$, i $, 26.^.1962 (Guichard & 
Harvey); Samsun, Samsun, i $, 20.vii.i959 (Guichard); Trabzon, Zigana Gecidi, 
1,650 m., i <$, 22. v. 1962; Artvin, above Artvin, 1,800 m., i $, 6.vi.i962 (Guichard 
6- Harvey). B.M. (N.H.). 

Typical race throughout EUROPE to E. SIBERIA. 

TENTHREDOPSIS A. Costa 

KEY TO SPECIES-GROUPS OF THE WORLD 

1 Clypeus subtruncate in front ......... 2 

Clypeus acutely or semicircularly excised in front medially to a depth of more 

than one-fourth of its maximal length. Penis valve without a spine. (Text- 
figs. 10 and 19). EUROPE and E. MEDITERRANEAN . . stigma-roup (p. 157) 

2 (i) Postocellar region slightly convex medially where it is about level with upper 

edge of the occipital carina and the temples. Antennal crests much smaller 
than ist antennal segment, and joined behind to the edge of the frontal area 
without any deep excision ......... 3 

Postocellar region excavated so that medially it is much below the level of the 
sharp edge of the occipital carina and the temples. Antennal crests often 
larger than ist antennal segment and separated from edge of frontal area by 
a deep excision. Penis valve as in stigma-group. N. EURASIA 

auriculata-Qroup (p. 157) 

3 (2) Antenna not white-ringed, nor abdomen with ist tergite black and flecked 

with opaque white, nor mesopleuron dull with dense sculpture ... 5 



156 



R. B. BENSON 




FIGS. 10-19. 1014. Tenthredopsis penis-valves in lateral view: 10, tesselata; n, sordida; 
12, soraria; 13, nassata; and 14, litterata. 15-19. Tenthredopsis penis- valves in dorsal 
view: 15, soraria; 16, sordida; 17, nassata; 18, litterata; and 19, tesselata. 



HYMENOPTERA FROM TURKEY 157 

Either antenna white-ringed, or abdomen with ist tergite black and flecked 

with opaque white, or mesopleuron dull with dense sculpture ... 4 

4 (3) Antenna longer than costa and stigma of fore wing. Clypeus not entirely black. 

Abdomen never with flecks of opaque white, but in $ usually with a pair of 
ventral lateral black stripes. Penis valve as in stigma-group. C. EUROPE 
and C. to E. ASIA ...... tarsata-Qroup (p. 159) 

Antenna not as long as costa + stigma of fore wing. Clypeus often entirely 
black. Abdomen sometimes with flecks of opaque white. Penis valve as in 
stigma-group. S.E. EUROPE and E. MEDITERRANEAN, albonotata- group (p. 159) 

5 (3) 6* with apical tergite carinate medially between two glabrous depressions; 

penis valve with spine (Text-figs. 11-18). - either has hypopygium with the 
posterior medial projection enlarged and strongly excised apically (Text-figs. 
21-23), or has malar space i equal to inter-antennal line (i : i to 1-2) and 
c. x 1-5 diameter of front ocellus. Occipital carina well developed and 
continuous. $ abdomen variously coloured but not with a pair of black 
ventral lateral stripes on a paler background except sometimes in sordida. 
PALAEARCTIC ....... nassata-Qroup (p. 161) 

cJ with apical tergite not modified thus ; penis valve as in stigma-group, without 
a spine. $ with medial posterior projection of hypopygium not enlarged 
nor excised apically. Malar space much shorter than inter-antennal line 
(i : i -3 to 1-5) and at most as long as front ocellus, or occipital carina obso- 
lescent laterally behind the orbits. $ abdomen usually with a pair of black 
ventral lateral stripes. E. MEDITERRANEAN . . /esf/va- group (p. 160) 



KEY TO A URICULATA-GROUP MALES AND FEMALES 

Antenna longer than costa + stigma of fore wing. $ black with following 
parts white: labrum, clypeus, face, antennal crests, orbits, temples, pro- 
notum, tegula, scutellum, flecks on mesonotal lobes and pleura, broad ventral 
and lateral strips on abdomen which dorsally is brown, medially, legs; legs 
brown to piceous with coxae, trochanters and tarsi white; <$ differs in 
that the underside of the thorax is i entirely white, and the head is almost 
entirely white except for frontal area. 9-10 mm. 

The Japanese race, ssp. japonica Takeuchi, is often larger (10-12 mm.) 
and the black is more extensive: the - is entirely black except for the white 
labrum, clypeus, lateral and ventral stripe of the abdomen, and the legs 
are black except for the yellowish white trochanters, front and middle tibiae, 
and tarsi. N. PALAEARCTIC (N. EUROPE to E. SIBERIA and JAPAN) 

carinata Malaise 

Antenna shorter than costa i stigma of fore wing. $ mostly yellowish brown 
with white markings; mostly black above and i white below. 9-10 mm. 
N. PALAEARCTIC (N. EUROPE to E. SIBERIA) . . \auriculata C. G. Thomson 1 



KEY TO STIGMA -GROUP MALES AND FEMALES 

i Mesopleura with mesepisterum shining, scarcely punctured .... 2 

Mesopleura dull with dense surface sculpture on mesepisterum between the 
punctures. Black with 4 middle abdominal segments red : white on clypeus, 
each side of labrum, inner orbits, temples, pronotum, tegula, scutellum and 

1 The following are synonyms of T. auriculata: Tenthredo sachalinensis Matsumura, 1911, syn. n. 
Tenthredopsis camtschatcali Enslin, 1927, syn. n. 

ENTOM. 22, 4. 7 



j 5 8 R. B. BENSON 

metanotum. Wings sybhy aline; venation black except for base of stigma, 
C and Sc of fore wing, which are white. 9-11 mm. S. and S. E. EUROPE 

floricola Costa 

2 (i) Head with occipital carina well developed from gena to vertex ... 3 

Head with occipital carina obsolescent laterally behind eyes. 

Head and thorax black with white on labrum, inner orbits, temples, prono- 
tum, tegula, scutellum; abdomen fulvous except for ist tergite and sawsheath. 
Wings subhyaline and venation black except for base of stigma, C and Sc of 
fore wing. 9-1 1 mm. TURKEY ..... guichardi sp. n. 

3 (2) Claws sub-bifid. Anal cell of fore wing normal ...... 4 

Inner tooth of claws smaller than end tooth and very short on front and middle 
legs. Anal cell of fore wing with cross-vein missing (? abnormal specimen). 
Black species with the following parts white: labrum, sides of clypeus, inner 
orbits, temples, edge of pronotum, tegulae, femora (except for inner stripe) 
and tibiae (except for inner stripe of hind pair) and yellowish white lateral 
stripe from 2nd to yth tergites of abdomen. 8 mm. TURKEY. $ only 

Harvey i sp. n. 

4 (3) Antenna shorter than costa of fore wing in $ and about as long as costa in <J. 

With 8th segment x 2-2-5 times as long as broad. $ wings slightly flavescent 
with venation yellowish white to brown. Black with the following parts 
yellow : labrum, sides of clypeus, fleck on inner orbits, fleck on temple, under- 
side of antenna, edge of pronotum, tegula, scutellum, tergites 3-6 of abdomen 
(except for mid -dorsal stripe) and most of legs. <J as $ except that the 
middle segments of the abdomen are buff with a black mid-dorsal band and 
a ventral lateral band each side. 9-11 mm. C. and N. EUROPE stigma (Fallen) 
Antenna as long as costa in $ and costa + stigma in <$, with 8th segment x 2-5-3 
times as long as broad. Wings subhyaline or slightly infuscate, with venation 
mainly black or brown. 6-5-11 mm. 

This complex of colour-forms more or less segregated into local races but 
with every intergradation have heretofore been treated as divisible into 
numerous species. Basically the $ colour pattern is black with the following 
parts white: labrum, i clypeus, fleck on inner orbits and temples, underside 
of antenna, edge of pronotum, tegula and middle of scutellum. Abdominal 
segments 3-6 are reddish yellow apart from a i developed mid-dorsal line 
and a longitudinal lateral ventral stripe each side. The femora, tibiae and 
tarsi are also mainly reddish yellow. 

In the paler forms the extent of white increases and the head can be almost 
entirely white apart from the frontal area, the posterior orbits, the post- 
genae and occiput; on the thorax the pronotum can become entirely white 
and white flecks can develop on the mesonotal lobes and mesopleura; on the 
abdomen yellowish white and then white flecks appear first on the sides of 
the first tergite and then yellow progressively entirely replaces the reddish 
yellow colour of the abdomen (f. albata Konow, Caucasus). In some forms 
the basic reddish brown is replaced by white and the black mid-dorsal line 
extends along the apical margins of the tergite (f. hungarica Klug and lacti- 
flua Klug, C. and S.E. EUROPE). In f. corcyrensis Andre S. and S.E. EUROPE, 
the basic pattern is altered by the labrum, clypeus, scutellum and legs 
becoming entirely black, with, at the same time, the disappearance of the 
mid-dorsal and lateral ventral abdominal lines. The basic pattern is as 
that described above for stigma. EUROPE and E. MEDITERRANEAN 

tesselata (Klug) 2 

2 Synonyms of T. tessalata : Tenthredo hungarica Klug, syn. n. T. lactiflua Klug, syn. n. Perineura 
excisa Thomson, syn. n. Tenthredopsis corcyrensis Andre, syn. n. T. albata Konow, syn. n. 



HYMENOPTERA FROM TURKEY 159 

KEY TO ALBONOTATA-GROUP MALES AND FEMALES 

1 Eyes only slightly converge in front where they are much further apart than the 

length of an eye. Malar space at least about as long as the diameter of front 
ocellus ............. 2 

Eyes so strongly converging in front that they are closer together here than the 
length of an eye. Malar space correspondingly short, about one-third as 
long as diameter of front ocellus. 

Only (J is known and this is a mainly black species with white labrum, 
mandibles and reddish legs, and antenna white-ringed from jth segment 
(8th and gth segments missing in unique type). Head strongly narrowed 
behind the eyes and mesopleura dull with dense surface sculpture. 9-5 mm. 
ISRAEL ......... convergent Benson 

2 (i) Mesopleura dull with dense surface sculpture. Hind tarsus not white on apical 

segments. Abdomen with or without white flecks on ist tergite ... 3 

Mesopleura shining and sparsely punctured. Hind tarsus with 3rd to 5th 
segments white. Antenna with or without a white ring covering i 6th 
and following segments. 

Black with red banded abdomen and white labrum, scutellum and fleck 
each side of ist tergite. 9-11 mm. S.E. EUROPE and E. MEDITERRANEAN 

albopunctata (Tischbein) 3 

3 (2) Antenna ringed with white or yellowish white on 6th and some of the follow- 

ing segments: Pronotum, tegula and ist tergite entirely black. Otherwise 
black with red-banded abdomen and white labrum and scutellum. 9-11 mm. 
C., S. and S.E. EUROPE and E. MEDITERRANEAN . annuligera (Tischbein) 4 

Antenna brown or black, pale below, but not ringed with white or yellowish 
white. Edge of pronotum and often tegula entirely white, and there is an 
opaque white fleck each side of the ist tergite. Otherwise coloured as in 
the preceding species except that in the red band of the abdomen is suffused 
over or obsolete. 9-10 mm. S.E. EUROPE and E. MEDITERRANEAN 

cJ? albonotata (Brulle) 



KEY TO TARS AT A -GROUP MALES AND FEMALES 

1 Smaller (8-9 mm.) with head strongly contracted behind eyes in dorsal view. 

Mesopleura scarcely punctate but with fine surface sculpture. S.E. ASIA 
(BURMA and FORMOSA) .......... 2 

Larger (10-12 mm.) with head parallel-sided in dorsal view. Mesopleura dull 
with dense surface sculpture between coarse punctures. (Black with 4 middle 
abdominal segments red in $, and scutellum white-marked; $ with the middle 
abdominal segments i infuscate as in scutellum.) C. and S.E. EUROPE 

3 1 ? tarsata (Konow) 

2 (i) Lateral ridge to frontal area of head continuous with antennal crests. Antenna 

not white-ringed and scutellum not marked with white. Two colour phases 
described originally as subspecies : mainly brown and mainly black. BURMA- 
YUNNAN frontier ....... birmanica (Malaise) 5 

3 Synonyms of T. albopunctata : Tenthredo benthini Rudow, syn. n. Tenthredo balcana Mocsary, 
syn. n. Tenthredopsis casia Konow, syn. n. T. quadrannulata Konow, syn. n. T. triforis Konow, 
syn. n. 

4 Synonyms of T. annuligera: Tenthredo tischbeini Frivaldskii, syn. n. *Tenthredopsis andrei Konow, 
syn. n. *T. pisinna Konow, syn. n. 

5 Synonym of T. birmanica: Thomsonia birmanica nigrorufa Malaise, syn. n. 



160 R. B. BENSON 

Lateral ridge to frontal area of head and antennal crests notched medially. 
Antenna often white-ringed from 6th and following segments, and scutel- 
lum usually white-marked. S.E. ASIA (BURMA and FORMOSA) 

insularis Takeuchi* 

KEY TO FESTI VA -GROUP MALES AND FEMALES 

In the B.M. (N.H.) there is a $ unnamed N. American species of this group of Tenthredopsis- 
It was sent to the Museum by Norton in 1865 as a representative of his recently described 
Allantus piceocinctus but it is not even congeneric with the type of that species. It was collected 
at Farrington, Connecticut. The genus is not now recognized in N. America, though larvae 
are recorded by Maxwell, 1955 : 93. 

1 Green species with black markings, and either occipital carina obsolescent 

laterally, or inner tooth of claw diverging from end tooth and slimmer than it 2 

Brown or red and black species. Occipital carina well-developed along whole 
hind margin of head, and inner tooth of claw parallel to end tooth and equal 
to it in girth ........... 3 

2 (i) Occipital carina obsolescent behind eyes. Malar space ($) x 2 diameter of 

front ocellus. Claws with inner tooth parallel to end tooth and equal to it in 
girth. Pulvillus on hind basitarsus only about half as long as apical breadth 
of basitarsus. Green with black fleck on frontal and postocellar regions of 
head, on each of the mesonotal lobes and in the middle of each tergite so as 
to form a medial abdominal row. Stigma entirely green. 8-5 mm. TRANS- 
CAUCASIA ........ viridis Zhelochovtsev 

Occipital carina well developed along whole hind margin of head. Malar space 
($) about x 1-5 diameter of front ocellus. Claws with inner tooth diverging 
from end tooth and slimmer ($) or shorter (<J) than it. Pulvillus on hind basi- 
tarsus about as long as apical breadth of hind basitarsus. Green with black 
markings on head, mesonotum, mesosternum, middle dorsal line and two 
lateral ventral lines on abdomen. Stigma dark with green base. 9-10 mm. 
TRANSCAUCASIA ........ ornatrix Konow 

3 (i) Anal cell of fore wing with cross- vein suberect and nearer base than apex of cell. 

Claws sub-bifid. Abdomen banded yellow or red, and with a pair of ventral 

dark stripes ............ 4 

Anal cell of fore wing with cross- vein oblique and near middle of cell. Claws 
with inner tooth shorter than end tooth. Black with white markings on 
head, thorax and margins of abdominal segments. 

Unique $. 7 mm. (? abnormal dwarf) has asymmetric supernumerary 
cross-veins. TRANSCAUCASIA ...... nigrescens Konow 

4 (3) Head with clypeus, inner orbits, temporal spot and thorax with edge of prono- 

tum and fleck on mesonotum white. Inner hind tibial spur about half as long 

as basitarsus ........... 5 

Head with clypeus black and thorax (apart from tegula and i mesoscutellum) 
entirely black. Inner hind tibial spur only about two-fifths as long as basi- 
tarsus. 

Abdomen with yellow band covering segments 3-9. Legs yellow with 
infuscate coxae and hind tarsus. <J not seen. 7-5 mm. TRANSCAUCASIA 
and TURKEY ......... nigella Konow 

6 Various colour forms of this species were originally described as sympatric subspecies on the Burma- 
Yunnan frontier and together with another from Formosa are synonyms of insularis Takeuchi : Thom- 
sonia insularis brunnescens Malaise, syn. n. T. insularis continentalis Malaise, syn. n. T. insularis 
deannulata Malaise, syn. n. T. insularis fuscicornis Malaise, syn. n. T. insularis ruficornis Malaise, 
syn. n. (Formosa). 



HYMENOPTERA FROM TURKEY 161 

5 (4) Abdomen with segments 2-6 pale yellow. Hind coxae entirely black. <$ not 

seen. 8-10 mm. TURKEY ...... straminata Konow 

Abdomen with segments 2-8 reddish yellow. Hind coxae black with broad 
lateral and ventral yellowish white stripes. 10 mm. TRANSCAUCASIA 

Jestiva Konow 

KEY TO NA SSA TA -GROUP 

1 c?c? 2 

?? 5 

2 (i) Spine on penis-valve shorter than inner hind tibial spur (Text-figs. 11-13 an d 

15-17). Hypopygium with at most a small emargination, not as wide as 
the length of this spur (Text-figs. 20, 22 and 23). Colour variable but 
antennae are always fuscous above and pale brown below .... 3 

Spine on penis-valve longer than inner hind tibial spur (Text-figs. 14 and 18). 
Hypopygium with a wide apical emargination about as wide as the length 
of this spur (Text-fig. 21). Head white below and on orbits, black above; 
antennae reddish yellow below and brown above; thorax yellowish white 
with black and yellow markings above; abdomen mainly reddish yellow 
marked with black on basal tergites. Large species 10-12 mm. EUROPE 
and TURKEY ........ <$ litterata (Geoffrey) 

3 (2) Penis-valve with spine attached before apex (Text-figs, n, 12, 15 and 16) . . 4 

Penis-valve with spine at apex (Text-figs. 13 and 17). 

<$ without external vein to hind wing is thought by some to represent 
the $ of another species known as coquebertii but their exact significance is 
unknown. 7-11 mm. EUROPE, TURKEY and SIBERIA to JAPAN nassata (L.) 

4 (3) Spines set close together on penis-valve and much shorter than apical breadth 

of hind tibia (Text-figs, n and 16). Straw-coloured species with black 
markings on head, thorax and abdomen. 8-10 mm. EUROPE $ sordida (Kl.) 
Spines set about half their own length apart on penis -valve and about as long 
as apical breadth of hind tibia (Text-figs. 12 and 15). Black species with 
yellowish white markings on face, orbits and thorax, n mm. TRANS- 
CAUCASIA ......... <J sororia Konow 

5 (i) Medial projection to hypopygium greatly enlarged so it is longer and broader 

at base than length of an inner middle tibial spur (Text-figs. 21 and 22) . 6 

Medial projection to hypopygium much shorter and narrower than length of 

inner middle tibial spur .......... 7 

6 (5) Projection to hypopygium narrowed behind, where it is narrower than the 

length of an inner middle hind tibial spur, and at its apex it is sharply excised 
(Text-fig. 21). Malar space almost equal to inter-antennal line and diameter 
of front ocellus. Pulvillus on basitarsus about as long as basal breadth of 
basitarsus and almost as long as pulvillus on succeeding tarsomere. 11-13 
mm. .......... $ litterata (Geoffrey) 

Projection to hypopygium scarcely narrowed behind where it is as wide as the 
length of an inner middle hind tibial spur, and at its apex it is broadly and 
slightly emarginate (Text-fig. 22). Malar space only about two-thirds of 
inter-antennal line and equal to diameter of front ocellus. Pulvillus on hind 
basitarsus about two-thirds of basal breadth of basitarsus and two-thirds of 
length of pulvillus on succeeding tarsomere. 12 mm. . $ sororia Konow 

7 (5) Straw-coloured species marked with brown and black. Projection to hypo- 

pygium deeply excised apically (Text-fig. 23). Q-II mm. . $ sordida (Klug) 
Very variable in colour from yellowish white, to brown, or red and more or less 
marked with black and white, to almost entirely black. Projection to 
hypopygium at most emarginate at an extreme apex (Text-fig. 20). 9-12 mm. 

9 nassata (L.) 



1 62 



R. B. BENSON 





sord. 

23 

FIGS. 20-23. Tenthredopsis hypopygia: 20, nassata; 21, litterata; 22, soraria; and 23, sordida. 




[Tenthredopsis floricola A. Costa 
TURKEY: Istanbul (2-3). 2$. 
S. and S.E. EUROPE]. 

* Tenthredopsis guichardi sp. n. 

$. Head and thorax black with the following parts creamy white: labrum, i clypeus, 
inner orbits, fleck behind eyes on lateral hind margin of head above, underside of flagellum, 
hind margin of pronotum, tegula, scutellum and fleck in middle of its post-tergite, metascutel- 
lum, small fleck on front and middle coxae and large fleck on side of hind coxa. Legs yellow 
except for most of coxae, trochanters and extreme base of front and middle and most of hind 
femora, which are black. Wings subhyaline; stigma with basal half white and apical black; 
costa and subcosta yellow; rest of venation piceous except at extreme base of wing. Abdomen 
yellowish brown, except for the mainly black ist, middle of 2nd and gth tergites, and saw- 
sheath. Length: 10-11 mm. 

Head shining, with obsolescent punctation. Clypeus medially emarginate in front to depth 
of nearly one-third of its total length. Malar space nearly as long as diameter of front ocellus. 
Antenna about x 2 as long as width of head behind the eyes. Frontal area with raised lateral 
margins fused with antennal crests and with a deep median groove from the front ocellus to 
the interantennal region. Carina on hind margin of head obsolete laterally behind the eyes so 
that genal carina below is separated from the occipital carina above. Distance between hind 
ocelli less than distance of an ocellus from occipital carina as i-o : 1-7, and less than distance 
of an ocellus to margin of eye as i-o : 1.9. 

Thorax normal with underthorax and mesonotum, apart from front half of front lobes, almost 
impunctate except in the sunken lateral areas of the latter which are rugose and the 



HYMENOPTERA FROM TURKEY 163 

scutellum which is opaque with a line of regular punctures separating the post-tergite which is 
i alutaceous. 

Legs normal; inner hind tibial spur about half as long as basitarsus. 

Wings normal. 

Abdomen with fine surface alutaceous sculpture above. Hypopygium and saw as in T. 
stigma. 

Pubescence over head and thorax pale and shorter than diameter of ocellus. 

6* as in $ but thorax including tegular entirely black and white on inner orbits and temples 
is much reduced. Pubescence on head infuscate. Distance between hind ocelli less than 
distance between ocellus and occipital carina as i-o : 1-3, and as distance between ocellus and 
eye margin as i-o : 1-7. Hind wings with marginal vein. Genitalia much as in T. stigma 
Klug (Text-figs. 10 and 19). 

Length: 9-10-5 mm. 

Holotype $. TURKEY: Ankara, Kubuk, 830 m., 22. v. 1960 (Guichard 6- Harvey). 
B.M. (N.H.). 

Paratypes. Same locality, 3 <$, 5 $, 21-22 . v . 1960 (Guichard & Harvey) ; TURKEY : 
Erzurum, 20 m. from Ispir on Ikizdere Road, 1,700 m., i $, 2.vi.i962 (Guichard 
6- Harvey). B.M. (N.H.). 



*Tenthredopsis harveyi sp. n. 

$. Black with the following parts white to yellowish white: palps, mandible bases, labrum, 
clypeus (except middle third), underside of antenna, spot on gena, inner orbits and spot behind 
temple, edge of pronotum, tegula, legs (except coxae, trochanters, spot on inner base of front 
femur, line on inner side of middle femur, and inner side of hind femur and tibia, most of hind 
tarsus), lateral each side of abdomen covering one-fourth of the width of tergites 2-7, a lateral 
fleck each side of middle of ist tergite and narrow apical margins of succeeding tergites and 
sternites. 

Wings hyaline with costa and subcosta and base of stigma white, rest of venation piceous. 
Length 8-5 mm. 

Head shining, with obsolescent punctation. Clypeus medially emarginate in front to depth 
of nearly one-third of its total length. Malar space slightly longer than diameter of front ocellus. 
Antenna more than x 2 as long as width of head behind eyes. Frontal area slightly convex 
above but with a deep round concavity above the interantennal area. Occipital carina clearly 
defined throughout. Hind ocelli further apart than from occipital carina as i-o : 1-4 and from 
margin of eye as i-o : 1-9. 

Thorax shining but with definite punctures dense on front of mesonotum, sparse on lateral 
lobes, scutellum, episternum and sternum of thorax, but the depressed lateral areas of the 
mesonotum as well as the mesepimeron and metapleura are dull and regulose. Legs normal 
with inner spur of hind tibia about half as long as basitarsus, but the inner tooth of the claw 
is not as long as the end tooth on the hind legs and on the front and middle legs is only about 
half as long as end tooth. 

Wings with the cross-vein to the anal cell missing in the fore wings but this may be an indi- 
vidual abnormality. 

Abdomen finely alutaceous above, hypopygium and saw normal (as in T. stigma). 

Pubescence pale on head and thorax and up to as long as diameter of an ocellus. 

Holotype $. TURKEY: Bolu, Ala Dagi, 2,000 m., Kartal Kaya Tepe, I5.vii.i962 
(Guichard & Harvey). B.M. (N.H.). 



164 R. B. BENSON 

Tenthredopsis tesselata (Klug) 

TURKEY, N.W., W., N. to E. : Widespread from Istanbul (3); Bursa (4); Samsun 
(5 and 6); Erzurum (6). Up to 1,700 m. 52 $, 59 9, iv-vi, in many colour forms 
including f. albata, Konow, f. excisa Thomson and f. corcyrensis Andre. 

N. IRAN: Elburz Mts., 2,000 m. (G. Heinricti). 

All EUROPE and TURKEY to N. IRAN. 

f Tenthredopsis convergens Benson, 1954 
ISRAEL. 

Tenthredopsis albopunctata (Tischbein) 

TURKEY, N.W., W., S.W., S., C., N.: Istanbul (2); Bursa (4 and 9); Mugla (10); 
Antalya (10); Samsun (24); Amasya (2, 5, 6, 7 and 13); Corum (2); up to 1,000 m. 
(near Merzifon in Amasya on 3.vi.59), 17. iv to 6. vi. 1959-62 and 20.vii.59 ( near 
sea level at Samsun ? 2nd brood). Forms with and without white-ringed antennae 
or tarsi occurred together. 20 <$, 2 . 

S.E. EUROPE (Yugoslavia, Bulgaria, Greece and Turkey) and ASIA MINOR. 

Tenthredopsis annuligera (Tischbein) 
TURKEY, C. : Amasya (i and 6). i <, 6 ?. 

C., S. and S.E. EUROPE (France, Germany, Czechoslovakia, Yugoslavia, Corfu, 
Russia) and TURKEY. 

Tenthredopsis albonata (Brulle) 

S.E. EUROPE (Greece and Turkey) and E. MEDITERRANEAN (Syria, Lebanon and 
Israel). 

Tenthredopsis viridis Zhelochovtsev, 1941 
TRANSCAUCASIA. 

^Tenthredopsis ornatrix Konow 
TRANSCAUCASIA. 

t Tenthredopsis nigrescens Konow 

TURKEY (W.): Usak, Bulgaz Dag, Namrun, 1,400-1,600 m., i <j>, 9-16. vi. 1965 
(Gembloux Coll. Belgium), and TRANSCAUCASIA. 

f Tenthredopsis nigella Konow 
TRANSCAUCASIA, TURKEY. 



HYMENOPTERA FROM TURKEY 165 

^Tenthredopsis straminata Konow 
S. TURKEY (Taurus Mountains). 

fTenthredopsis f estiva Konow 
TRANSCAUCASIA. 

Tenthredopsis litterata (Geoffroy) 

TURKEY, N.W., C. and E. : Istanbul (3); Amasya (3 and 7) and Erzurum (6). 
4 & 2 ?. 
All EUROPE and TURKEY. 

^Tenthredopsis sororia Konow 
TRANSCAUCASIA. 

Tenthredopsis nassata (L.) 

TURKEY (C. and N.E.) : Ankara (39); Cankiri, Isik Dag, 1,200 m., i $, 25. vi. 1966 
(Demelt Coll.); Amasya (3, 13); Trabzon (15). 

EUROPE, ASIA MINOR, SIBERIA to JAPAN. 

TENTHREDINIDINI 
RHOGOGASTER Konow 
Rhogogaster was revised by Benson, 1965. 

*1[ Rhogogaster genistae Benson 

TURKEY, C.: Amasya (7). i $, 2 $. 
Also EUROPE, N. AFRICA. 

Rhogogaster picta (Klug) 

TURKEY, N.W., W., C., N.E. and E.: Istanbul (3); Bilecik (i and 2); Konya (3); 
Amasya (7 and 13); Giresun (3); Erzurum (4). 10 <$, 14$. 

Also EUROPE and TURKEY. 

*t Rhogogaster dryas (Benson) 

TURKEY, C. : Amasya (5 and 18). 4$. 
EUROPE and TURKEY. 



166 R. B. BENSON 

Rhogogaster punctulata (Klug) 

TURKEY, N.E. : Trabzon (4 and 15); Artvin (4). 1,246-1,800 m. 5 $. 

These specimens seem to be racially distinct from the normal European form in 
the complete lack of the lateral row of black spots each side of the abdomen. This 
form has also occurred as a rare aberration in Finland. 

EUROPE and TURKEY. 

*t Rhogogaster naias Benson, 1965 

TURKEY, E. : Gumusane (5). n $, 12$. 
Type series. 

Rhogogaster viridis (L.) 

TURKEY, C., N.E. and E.: Ankara (12); Amasya (13); Trabzon (3); Rize (8); 
Artvin (3) ; Erzurum (10). From sea level up to 2,600 m., 3 $, 7 -. Holarctic species. 

*f Rhogogaster chlorosoma (Benson) 

TURKEY, N.W., C., N.E. and E.: Istanbul (9); Ankara (15, 35 and 41); Tokat (i); 
Konya (3); Giresun (3); Erzurum (6). From sea level up to 1,400 m., i <$, 23 $. 

EUROPE, TURKEY and SIBERIA. 

Rhogogaster auctor Weiffenbach, 1967 

TURKEY: Nigde, Ciftehan, vi. 1965 and Ankara, Kizilkahaman, v. 1965 (Demelt 
Coll.). 

TENTHREDO L. 

Keys to the palaearctic species of the bifasciata-, zonula-, scrophulariae- and 
arcuata-groups, (as well as the species now segregated as Elinora and Cuneala] 
were given by Enslin, 1910, and to the olivacea-mesomelas-, cyanatra-atra- and 
maculata-temula-groups by Enslin, 1920. 

KEY TO SPECIES-GROUPS 

1 Antennal crests, over antennal sockets, reach further forward (in dorsal view) 

than the level of the front of the eyes, and in the olivacea-mesomelas group, 
the crests are separated from the sides of the frontal area by a deep excision ; 
the area between the crests is more than half as deep as the distance between 
them. Flagellum of antenna more than x i the width of the head behind 
the eyes ............ 2 

Antennal crests confluent with the sides of frontal area, and not reaching 
forward further than the level of the front of the eyes and/or the area between 
them is not half as deep as the width between them .... 3 

2 (i) Mostly green or green and black species but some partly yellow or brown, with 

antennal crests separated behind from the sides of the frontal area by a deep 
excision ...... olivacea-mesomelas- group (p. 167) 



HYMENOPTERA FROM TURKEY 167 

Species variously coloured, often with black, red, brown or yellow but some 
with green, antennal crests behind confluent with sides of frontal area 

cyanata-atra- group (p. 167) 

3 (i) Head strongly swollen behind eyes and, together with thorax, entirely black 

(except at most for small yellow flecks on edge of pronotum). Wings infus- 
cate on at least apex of front pair (W. PALAEARCTIC) bifasciata-Qroup (p. 169) 
Head not strongly swollen behind eyes, or head and thorax more richly marked 

with yellow. Wings i hyaline or with infuscate patches .... 4 

4 (3) Head with temples shining and at most sparsely punctured, and occipital carina 

obsolete behind post-ocellar region. Antenna less than twice as long as 
breadth of head behind eyes. Male hind tarsal claws without acute basal 
lobe ......... zonula- group (p. 170) 

Head dull with strong punctation above or occipital carina well developed 
along whole of hind margin of head or including the post-ocellar region. 
Antenna long or short. Male hind tarsal claw with or without acute basal 
lobe ............. 5 

5 (4) Wings hyaline in fresh specimens and antenna short (penultimate segment less 

than x i as long as broad). Stigma mostly pale. Claws usually with a 
basal lobe developed. (HOLARCTIC) . . . arcuata-Qroup (p. 176) 

Wings strongly flavescent, or infuscate, at least partly. Antenna often 
elongate (penultimate segment at least x i as long as broad). Stigma 
often dark. Claws often without basal lobe ...... 6 

6 (5) Fore wings evenly i infuscate. Stigma black. Abdomen with metallic 

lustre. Mesosternum each side and mesoscutellum with tooth-like projection. 
Antenna often with penultimate segment less than x i as long as broad. 

(ORIENTAL) opposita- group 

Fore wings subhyaline, flavescent or with infuscate patches. Stigma often pale. 
Abdomen not metallic. Mesosternum and often mesoscutellum without 
tooth-like projection. Antenna with penultimate segment at least x i as 
long as broad ........... 7 

7 (6) Wings flavescent, or fore wing with infuscate apex or front margin. Stigma 

pale. (HOLARCTIC) . . . vespiformis-scrophulariae-&roup (p. 175) 

Wings hyaline or stigma dark. (PALAEARCTIC) maculata-temula-&roup (p. 173) 

flnschi and japonica 

The OLIVACEA-MESOMELAS-group 
Tenthredo mesomelas L. 

TURKEY, N., N.E. and E. : Bolu (i); Zonguldak (i) ; Trabzon (3, 4, 15 and 16); 
Gumusane (7 and 8). 700-2,600 m. 13 $, 28 $. 

EUROPE, TURKEY, TRANSCAUCASIA and SIBERIA. 

The CYANATA-ATRA-group 
Tenthredo atra duplicate, (Enslin) 

This race differs from the typical race in always having white on the genae and 
each side of the ist tergite. 

TURKEY, E.: Trabzon (3). 2 ^, 2 $; Trabzon (15). 5 $; Artvin (4). i ?. 

EUROPE and TRANSCAUCASIA. 

The species occurs throughout EUROPE to E. SIBERIA. 



168 R. B. BENSON 

Tenthredo mandibularis F. 

C. EUROPE and TRANSCAUCASIA. 

Tenthredo araxana Mocsary 
TRANSCAUCASIA. 

Tenthredo caligator Eversmann 

TURKEY, N.E. : Trabzon (15-16). 40 <$, 59 $. 
TURKEY and TRANSCAUCASIA. 

Tenthredo albopicta Puls 

TURKEY, N.E.: Trabzon (15-16). i <$. 
TURKEY and TRANSCAUCASIA. 

Tenthredo discophora Konow 

TURKEY, N., N.E. and E. : Bolu (i) ; Gumusane (7 and n); Trabzon (14, 15, 16 
and 17); Rize (8). 1,400 to 2,500 m. 22 <$, 51 $, 27.v-i4.vii. 

TURKEY and TRANSCAUCASIA. 

Tenthredo purpurea Puls 

TURKEY, N.E. : Trabzon (15-16). 3 <$, 4 $. 
TURKEY and TRANSCAUCASIA. 

Tenthredo livid a L. 

TURKEY, N. : Zonguldak (i). i $. 

EUROPE, TURKEY, TRANSCAUCASIA and SIBERIA. 

Tenthredo colon Klug 

TURKEY, W. and N.E. : Kutahya (9) ; Trabzon (14 and 15). 3 <$, 3 -. 
EUROPE, TURKEY, TRANSCAUCASIA and N. ASIA to KAMTCHATKA. 

Tenthredo balteata Klug 

TURKEY, N.E. : Trabzon (15); Rize (8). 2$. Not previously recorded outside 
EUROPE. 

Tenthredo solitaria Scopoli 
C. and S. EUROPE and TRANSCAUCASIA. 



HYMENOPTERA FROM TURKEY 169 

Tenthredo ferruginea Schrank 

TURKEY, N.E.: Trabzon (15); Rize (8). i <$, i ?. 
EUROPE, TURKEY, TRANSCAUCASIA, SIBERIA to JAPAN. 

Tenthredo luteipennis Eversmann 

TURKEY: Trabzon (13). i $; (14, 15). 17 <, 26 $. 
TURKEY, and CAUCASUS and PAMIRS up to 3,400 m. 

The BIFASCIATA-group 

This group consists of those species with i infuscate wings, and head, greatly 
swollen behind the eyes, together with the thorax entirely black (except at most 
for small lateral yellow fleck on pronotum). Abdomen black with sometimes one 
or more tergites flecked or banded with yellow but very variable in colour pattern 
as are the wings. 

KEY TO MALES AND FEMALES 

1 Mesoscutellum shining, almost impunctate ....... 2 

Mesoscutellum at least partly dull with dense punctures or surface sculpture . 3 

2 (i) Pubescence on head and thorax silvery, not infuscate. 11-13 mm. 6* not 

seen. S.E. EUROPE and E. MEDITERRANEAN . . . kiefferi (Konow) 

Pubescence on head and mesonotum infuscate. 11-13 mm. C. and S. EUROPE, 

E. MEDITERRANEAN to IRAN ...... costata Klug 

3 (i) Mesoscutellum entirely dull with dense surface sculpture between punctures. 

Pubescence on head and mesonotum fuscous ..... 4 

Mesoscutellum with some shining areas anteriorly between the punctures . 5 

4 (3) Abdomen black except at most for i yellow tergite, and occasionally also with 

yellow flecks on 2nd and 4th, but $ often entirely black. Legs entirely black. 
Inner hind tibial spur more than half length of basitarsus. Wings usually 
entirely fuscous in <J, but in $ often flavous basally. 11-15 mm. C. EURO- 
PEAN ALPS and E. MEDITERRANEAN .... caucasica Eversmann 
Abdomen with 3rd to 7th tergites (?) or 3rd and 4th ($) margined apically 
with yellow and legs with yellow tibiae and tarsomeres, necked with brown 
at their apices. Inner hind tibia scarcely half as long as basitarsus. Wings 
of $ flavous infuscate apically. 12-14 mm. C. EUROPEAN ALPS 

stecki (Konow) 

5 (3) Abdomen or legs marked with yellow. Inner hind tibial spur longer than 

apical width of tibia. Wings sometimes flavous ..... 6 

Entirely black species with strongly infuscate wings. Inner hind tibial spur 
about as long as apical width of tibia. 13-14 mm. E. MEDITERRANEAN 

violascens (Konow) 

6 (5) Abdomen with ist tergite black. Scutellum strongly convex, gibbous . . 7 

Abdomen with ist, 3rd, 4th and sometimes 2nd, 5th and 6th tergites flecked 
or margined apically with yellow, as also is often the pronotum; otherwise 
black, except for paler tibiae and tarsi. Mesoscutellum only slightly convex 
in middle. 12-15 mm. <$ not seen. TURKESTAN . . . bractea Enslin 

7 (6) Fore wings fuscous throughout and either stigma dark brown or pubscence on 

head and thorax partly fuscous ........ 8 



170 R. B. BENSON 

Fore wings flavous basally with infuscate apices and yellow stigma. Pubescence 

on head and thorax silvery. 10-12 mm. EUROPE and SIBERIA rossii (Panzer) 
8 (7) Stigma and costa of fore wing yellow or brown. 12-14 mm. C. and S. EUROPE 

bifasciata Miiller 
Sigma and costa of fore wing infuscate. (? subspecies of preceding) . 11-13 mm. 

cJ not seen. E. MEDITERRANEAN ..... diversipes Mocsary 

Tenthredo kiefferi (Konow) 

TURKEY, S.W., S. and C.: Aydin (2); Antalya (5 and 14); Mersin (7, 12 and 14); 
Ankara (12, 14, 15, 21, 35 and 37); Corum (2); Amasya (2, 3 and 7); Kayseri 
(3 and 4); Sivas (3). Up to 1,600 m. in Mersin. 69 <, 93 $. 

S.E. EUROPE, LEBANON, ISRAEL and TURKEY. 

Tenthredo costata Klug 

TURKEY, C., N.E. and E. : Nigde (5); Ankara (15); Amasya (7); Sivas (3); 
Giresun (3); Artvin (2); Erzurum (4 and 6). 700-1,700 m., 2 <$, n $. 

C. and S. EUROPE, TURKEY and IRAN. 

Tenthredo caucasica Eversmann 

TURKEY, N.E. and E. : Trabzon (3, 14, 15 and 16); Gumusane (10); Erzurum (6). 
1,400-2,000 m., 33 $, 60 ?. 

EUROPEAN ALPS (France and Switzerland), TURKEY and TRANSCAUCASIA. 

Tenthredo violascens (Konow) 

TURKEY, E.: Erzurum (6). 13$. 
Also TRANSCAUCASIA. 

Tenthredo diversipes Mocsary 
SYRIA, LEBANON and ISRAEL. 

The ZONULA-group 

KEY TO MALES AND FEMALES 

1 Flagellum of antenna mainly black. Head scarcely swollen behind eyes or 

punctate above. Wings subhyaline; stigma infuscate apically . . 2 

Flagellum of antenna mainly yellow. Head strongly swollen behind eyes and 
impunctate above. Wings yellowish throughout; stigma yellow with brown 
apex. 

Occipital carina continues only up to level with top of eyes, being obsolete 
behind temples. S.E. EUROPE and E. MEDITERRANEAN . flavipennis Brulle 

2 (i) Tongue long (prelabium longer than greatest measure of eye). Occipital carina 

obsolete behind upper half of eyes, temples and postocellar region. Frontal 
area densely punctate. Clypeus excised to only about one-third of its total 
length. Abdomen with ist tergite black, but with whole of 4th and 5th 
segments yellow above and often 3rd and 6th laterally .... 3 



HYMENOPTERA FROM TURKEY 171 

Tongue normal (prelabium shorter than greatest measure of eye). Occipital 
carina denned along whole hind margin of head except behind postocellar 
region. Frontal area sometimes shining with sparse punctures. Clypeus 
excised to more than one-half its total length. Abdomen with i ist, whole 
of 5th, sometimes 4th and apices of some of the following tergites yellow . 4 

3 (2) Frontal area deeply concave in the middle to a depth and size much greater 

than that of an inverted ocellus. Antenna with ist segment yellow. S.E. 
EUROPE and E. MEDITERRANEAN ..... dialeuca (Konow) 

Frontal area not concave in the middle to a depth as great as an inverted 

ocellus. Antenna entirely black. C. and S. EUROPE and E. MEDITERRANEAN 
to IRAN ......... frauenfeldi Giraud 

4 (2) Mesopleura above dull with dense surface sculpture ..... 5 

Mesopleura above shining between the follicles of dense pubescence . . 6 

5 (4) ist antennal segment and tegula mainly yellow. Frons densely punctured and 

lower inner orbits dull with dense surface sculpture. C. and S. EUROPE and 

N. AFRICA ........ distinguenda (R. v. Stein) 

ist antennal segment entirely, and tegula mainly, black. Frons sparsely 
punctured and lower inner orbits shining between surface sculpture. E. 
MEDITERRANEAN ....... hyrcana sp. n. (p. 172) 

6 (4) Hind femur tipped with black at apex only. Antenna with one or two basal 

segments mostly yellow. Hind tibia and tarsus of $ marked with black or 
dark brown. C. and S. EUROPE, N. AFRICA and E. MEDITERRANEAN zonula Klug 
Hind femur with most of inner side black. Antenna entirely black. Hind 

tibia and tarsus of $ marked with reddish brown. EUROPE amoena Gravenhurst 

Tenthredo flavipennis Brulle 

Allantus lautus Konow. 

A. luminosus Konow, syn. n. 

TURKEY, E.: Gumusane (n); Erzurum (6). 1,700-1,800111. 
S.E. EUROPE, TURKEY and TRANSCAUCASIA. 

Tenthredo dialeuca Konow 
A llantus jugalis Konow, syn. n. 

TURKEY, N. and E. : Samsun (8) ; Gumusane (10) ; Erzurum (15). Up to 2,300 m., 
i & 4 ? 
S.E. EUROPE, TURKEY and TRANSCAUCASIA. 

Tenthredo frauenfeldi Giraud 

Allantus helveticus Konow, syn. n. 
A . mevceti Konow, syn. n. 
A. montanus Enslin, 1910. 

TURKEY, N. and E. : Samsun (5); Gumusane (10). Up to 1,800 m., i <$, 2$. 
IRAN : Mazandaran, Kalar Dasht, Sardab, Rud Valley, 3 km. above Rudbarak, 
1,500 m., i$, 17. iv. 1966 (D. B. Baker). 

C. and S.E. EUROPE, TURKEY and IRAN. 



i 7 2 R. B. BENSON 

Tenthredo zonula Klug 

Allantus similis Mocsary. 

A. nazareensis Andre, syn. n. 

A. serena Konow, syn. n. 

TURKEY, N.W., W., S.W., S., C., N.E. and E.: Istanbul (3); Bursa (2 and 14); 
Aydin (2) ; Mugla (9 and 10) ; Antalya (9 and 10) ; Nigde (5) ; Ankara (12) ; Corum (2) ; 
Amasya (2, 3 and 7); Artvin (2); Erzincan (i) ; Erzurum (5, 6 and 14). Up to 
2,250 m. in the east, 50 <$, 35 $. 

C. and S. EUROPE, ISRAEL, SYRIA, LEBANON and TURKEY. 

* Tenthredo hyrcana sp. n. 

(Text-figs. 26, 26a) 

$. Black with the following parts yellow: mandibles, labrum, clypeus, hind margin of pro- 
notum, outer margin of tegula, trochanters extreme base, and apex of femora, tibiae except 
apices of middle and hind pair and apices of middle and hind tarsomere, apical half of isttergite, 
5th abdominal segment and apex of yth, 8th and gth tergites. Wings subhyaline; base of 
stigma and costa yellow, apex of stigma and rest of venation piceous. Length 8-9 mm. 

Head slightly contracted behind and with clearly marked occipital carina, though this is 
obsolete behind postocellar region; labrum acute in front; clypeus deeply excised in front to 
depth of more than half total length of clypeus, slightly convex above; flagellum of antenna 
longer than width of head behind (as 1-3 : i-o) . Frontal area with an 8-shaped medial concavity. 
Punctation sparse and very fine on ridges of frontal area and lower inner orbits but gena dull 
with dense surface sculpture. 

Thorax normal and shining between fine widely-separated punctures on mesonotum. Meso- 
thorax below dull with very dense surface sculpture. Legs normal; inner hind tibial spur 
scarcely half as long as basitarsus. Abdomen dull above with dense transverse alutaceous 
sculpture. Saw. (Text-figs. 26 and 26a.) 

Pubescence white and well-developed on head and thorax and in length up to about x ij 
diameter of an ocellus. 

(J as $ but that the legs are much paler (the whole of the front legs are yellow except for the 
spurs, the claw bearing segment and extreme bases of other segments; the middle legs except 
for the tarsus and an outer black line on the tibia and the hind legs except for the base of the 
coxae, the apical half of the femur together with the whole tibia and tarsus which are black), 
the costa is darker, the ist tergite is pale only on the apical margin and the 3rd-5th sternites 
of the abdomen. 

Holotype $. TRANSCAUCASIA: Armenia, Delizhan, 1,500-2,200 m., i6.vi.i934 
(A. N. Zhelochovtsev). B.M. (N.H.). 

Paratypes. TRANSCAUCASIA: i $ (same data). TURKEY: Ankara, Idris Dagi, 
1,300 m., i $, 30. vi. 1962 (Guichard & Harvey); Amasya, 500 m., 12 <$, 22-23. v. 1959 
(Guichard), 460 m., 2 ^, i 9, 2-6. vi. 1959; Erzurum, Ispir, 20 kms on Ikisdere Rd., 
700 m., i <$, 2.vi.ig6z (Guichard < Harvey). B.M. (N.H.). 

This species is differentiated in the key above from T. distinguenda Stein, to which 
it is most closely related. According to Dr. Zhelochovtsev this species was recognized 
and named " Allantus hyrcanus " by Gussakovskii in his MSS on the Tenthredininae 
in the Faune de 1'URSS before the war, but was never published. I have therefore 



HYMENOPTERA FROM TURKEY 





euph. 25 




hyrc. 





r hyrc. 26a 



FIGS. 24-26. 24. Tenthredo euphorbiae sp. n. penis-valve. 25-26. Tenthredo saws: 
25, euphorbiae sp. n.; and 26, hyrcana sp. n. 25a-26a. Enlarged loth and nth teeth 
from apex of saws: 25a, euphorbiae sp. n.; 26a, hyrcana sp. n. 

selected as the holotype the specimen sent to me by Zhelochovtsev, bearing the 
name given by Gussakovskii. 



The MACULATA-TEMULA -groups 

KEY TO MALES AND FEMALES 

Abdomen only one tergite (3rd) or two (3rd and 4th) entirely yellow; 5th 
tergite at least partly black; <$ with underthorax pale. Mesopleura with 
angular projection. Hind claws with acute basal lobe. 10-12 mm. 

(temula sspp.) 



ENTOM. 22, 4. 



174 R. B. BENSON 

9- Abdomen with two tergites (4th and 5th) entirely yellow or orange; 3rd 
tergite at least mainly black; $ with underthorax black. Mesopleura without 
angular projection. Hind claws without basal lobe. 12-16 mm. 

(maculata complex) 4 

2 (i) o* has 8th tergite with at most apical margin yellow; $ has apex of abdomen 

with at most 8th and gth tergites broadly yellow and a small fleck in the 
middle of the yth ........... 3 

cJ has 8th tergite with at least apical half yellow and $ apex of abdomen has 
at least a large fleck of yellow on the yth tergite and mostly has yellow on 
6th-gth tergites. 

Scutellum sharply raised so as to form right angle in profile. $ 3rd tergite 
entirely yellow and $ 4th tergite black-flecked medially, scutellum all black. 7 

BRITAIN, S.W. FRANCE, N. SPAIN and ITALY . . temula celtica Benson 

3 ( 2 ) 6* 3 r d tergite entirely yellow; $ 4th tergite with medial fleck of black. Scutel- 

lum flatter, with front face more shining and with no yellow fleck. C. 
EUROPE (excluding SPAIN and ITALY) . . . temula temula Scopoli 

cj 3rd tergite with medial black fleck; $ with 4th as well as 3rd tergite entirely 
yellow. Scutellum forming almost a right angle in profile and often with 
yellow fleck. E. MEDITERRANEAN .... temula xanthaspia Enslin 

4 (i) Middle and hind coxae and trochanters mainly, and hind femur entirely, black 

in both sexes. $ abdomen with 4th and 5th segments pale yellow ; <$ abdomen 
pale yellow except for the i infuscate ist and 2nd tergites and two apical 
segments. $ tarsus strongly swollen so that in ventral view the 2nd and 3rd 
tarsomeres are less than twice as long as broad (Text-fig. 29) (maculata sspp.) 5 

Middle and hind coxae and trochanters mainly pale and hind femur pale at base 
in both sexes. - abdomen with 4th and 5th segments orange above and pale 
yellow below; <$ abdomen orange above and pale yellow below except for 
infuscate first 2 tergites. $ tarsus less strongly swollen so that in ventral 
view the 2nd and 3rd tarsomeres are more than twice as long as broad (Text- 
fig. 30). E. MEDITERRANEAN ...... vestita Andre 

5 (4) Hind tibia infuscate apically. $ with 2nd and 6th-8th tergites not marked 

with yellow. ............ 6 

Tibiae not infuscate apically. $ with 2nd, 3rd, 6th and yth tergites flecked 
with yellow laterally and 6th-gth tergites with medial apical margins yellow. 
ITALY, N. Appenine Mountains . . maculata diana ssp. n. (p. 174) 

6 (5) Scutellum, tengula and hind edge of pronotum marked with yellow in $. 

W. and C. EUROPE and ITALY (except N. Appenines) maculata maculata Geoffroy 
Thorax entirely black except for narrow outer edge of tegula. Hungarian 

Plain (CZECHOSLOVAKIA, HUNGARY and AUSTRIA) . maculata semseyiMocsary 

[Tenthredo maculata diana ssp. n. 

$. Black with the following parts yellow: labrum, clypeus, posterior half of pronotum, 
tegula, scutellum, fleck on metapleura and on fore and middle coxae, trochanters and front 
of fore and middle femora, all tibiae and tarsi, 4th and 5th segments of abdomen together with 
a lateral fleck each side of 2nd, 3rd and 6th tergites and medial hind margin of 6th-gth tergites. 

Wings subhyaline, with costa, front of subcosta and veins in anal regions, yellow ; stigma and 
rest of venation black. 

Structure as in T. maculata but mesoscutellum is more convex medially. 13-14 mm. long. 

Holotype $. ITALY: Emilia, Mt. Bretra, i.v.igia (A. Fiori] (Geneva Mus.). 

Paratypes. ITALY, Emilia, La Lama, i 9, 10 . vi . 1962 (A . Servadei] (Padova 
Mus.); Marches, Mt. Catria, i $, v.ig33 (Alzona). B.M. (N.H.)]. 



HYMENOPTERA FROM TURKEY J 75 

Tenthredo vestita Andre 

TURKEY, C., N., N.E., E. : Amasya (3, 6 and 13); Samsun (17); Gumusane (14); 
Trabzon (3, 4 and 15); Erzurum (6). 500-1,400 m., 23^, 24$. 

TURKEY and TRANSCAUCASIA. 

Tenthredo temula xanthaspis Enslin 

TURKEY, N. and N.E. : Sinop (2); Trabzon (3 and 4); Rize (5); Artvin (3). 900- 
1,400 m., 3 & 7 ?. 

Subspecies only in TURKEY and TRANSCAUCASIA. Species otherwise confined to 
EUROPE. 

The VESPIFORMIS-SCROPHULARIAE-group 

These species are coarsely punctured on the head, which is often swollen behind 
the eyes; the antennae often have the flagellum yellow and the wings often have 
infuscate patches. 

Tenthredo marginella Fabricius 

TURKEY, W., N.E. and E. : Bursa (13) ; Trabzon (14) ; Gumusane (4). i <$, 3 $. 
EUROPE, TURKEY and TRANSCAUCASIA. 

* Tenthredo zona Klug 

TURKEY, S.W.: Mugla (10). i ?. 

Not previously recorded outside EUROPE. 

Tenthredo vespa Retzius 

TURKEY, N.E.: Trabzon (15). i $. 

EUROPE, TURKEY, TRANSCAUCASIA and SIBERIA. 

Tenthredo scrophulariae L. 

TURKEY, C. : Tokat (i); Sivas (2). 3 $, 2 $. 
EUROPE, TURKEY and TRANSCAUCASIA. 

Tenthredo excellens (Konow) 

\Allantus persa Konow. 

TURKEY, S., C. and E. : Mersin (12 and 15) ; Ankara (9, 12, 31, 37 and 39) ; Amasya 
(i and 7) ; Kayseri (3) ; Sivas (3) ; Erzurum (7, 8 and 9). 5,000-2,300 m., 21 , 24 $. 

N. IRAN: Elburz Mountains, 1966 (D. B. Baker}. 
S.E. EUROPE, TURKEY, TRANSCAUCASIA and IRAN. 



176 R. B. BENSON 

Tenthredo cinctipleuris (Enslin) 

TURKEY, S. and E. : Mersin (7); Gumusane (5); Erzurum (6). 14 , 12$. 

IRAN: Elburz Mountains, N. side of Kandara Pass, 2,360 m., i ^, i $, 9^1.1965 
(D. B. Baker}. 

EUROPE, TURKEY, TRANSCAUCASIA and IRAN. 

* Tenthredo propinqua Klug 

TURKEY, N.: Kastamonu-Cankiri Border (2). 2,000 m., i <J. 
S.E. EUROPE and TURKEY. 

Tenthredo luteocincta Eversmann 

TURKEY, N.E. : Trabzon (14 and 15). 3 ?. 
TURKEY, TRANSCAUCASIA and S.E. RUSSIA. 

Tenthredo reitteri (Enslin) 
Allantus lituratus Mocselry. 

TURKEY, W., C., N.E. and E. : Kutahya (9); Cankiri (4); Trabzon (4, 14, 15 and 
16); Gumusane (7). 1,400-2,500 m., 42^, 23$. 

TURKEY and TRANSCAUCASIA. 

Tenthredo I ant a (Konow) 
TRANSCAUCASIA. 

Tenthredo luminosa (Konow) 
TURKEY. 

The ARCUATA -group 
The species of this group were revised by Benson (1959). 

Tenthredo schaefferi Klug 

Outside C. and S.E. EUROPE, a few specimens of f. perkinsi of this species were 
found in N. IRAN: Elburz Mountains, 2,500-3,200 m. (see Benson, 1959). 

*^Tenthredo acerrima Benson 

TURKEY, N.E. and E. : Trabzon (14 and 15); Artvin (4); Gumusane (8). 1,400- 
2,450 m., vi-viii, many <$ and $. 

EUROPE and TURKEY. 



HYMENOPTERA FROM TURKEY 177 

*-\Tenthredo titania Benson 

TURKEY: Trabzon, Zigana Dagi, 1,400 m., plentiful vii-viii, 1959-60, and Soganli 
Gecidi, 2,600 m., 40 <$, 84 $, v-vii. 1960-62 (Guichard & Harvey}. 

TRANSCAUCASIA and TURKEY. 

Tenthredo trivittata (Andre) 
\Allantus kussariensis Konow, syn. n. 

TRANSCAUCASIA. 

* Tenthredo euphorbiae sp. n. 

(Text-figs. 24, 25, 25a) 

$. Green with the following parts black: head above the clypeus (except for gena and lower 
outer orbits), with antenna (except underside of flagellum), prothorax (except pronotum behind), 
mesothorax (except tegula, scutellum and its post-tergite, the posterior half of the episternum, 
the epimeron and the sternum), a posterior line on the hind and middle coxae and femora, on 
all the tibiae and tarsi, and on the abdomen a transverse fleck on the middle basal part of each 
tergite. Wings hyaline; stigma, costa, subcosta and anal cells green, rest of venation piceous. 

Length 8-9 mm. 

Head dull with coriaceous sculpture, slightly contracted behind with occipital carina well 
developed from gena to post-ocellar region. Labrum rounded in front. Clypeus excised in 
front of a depth less than half the total length of clypeus (i-o : 2-5), scarcely convex, shining 
with only obsolescent punctures and surface sculpture. Eyes closer together in front than the 
length of an eye (i-o : 1-4). Malar space about half diameter of front ocellus. Antenna more 
than x i \ as long as width of head behind eyes (1-6 : i-o) ; 3rd segment = 4th + 5th. Distance 
between antennal sockets about the same as the diameter of a socket. Antennal crests project 
less than half the distance between them; crests continuous with lateral walls of frontal area. 
Posterior ocelli closer together than distance from occipital carina (i-o : 1-2) and half the distance 
from nearest eye margin (i-o : 2-0) ; postocellar region clearly defined laterally by deep furrows, 
and almost x 2 as broad as long (1-9 : i-o). 

Thorax normal; dull with dense coriaceous sculpture; mesoscutellum slightly roundedly 
convex; mesopleura without any raised protuberance. Legs with hind femur almost as long 
as tibia (i-o : 1-3); hind tarsus longer than tibia (i-i : i-o); inner hind tibial spur about half 
the length of basitarsus; tarsal pulvillus on hind basitarsus about as long as apical width of 
basitarsus. Claws sub-bifid without basal lobe. 

Abdomen with typical transverse alutaceous sculpture above. Hypopygium not emarginate 
laterally. Sawsheath slender and about two-thirds the length of hind tibia. Saw (Text-figs 
25 and 25a). 

Pubescence fuscous on upper head where it is up to a length greater than the diameter of an 
ocellus; colourless on thorax and shorter than diameter of an ocellus. 

cJ as $ but the underthorax and abdomen are almost entirely green except for the black 
sutures between the sclerites, and the medial basal halves of the ist and 2nd tergites. The 
eyes converge more strongly in front where the distance between them to the length of an eye 
is as i-o : 1-7. The malar space is but one-quarter as long as the diameter of the front ocellus. 
The antenna is almost twice as long as the width of the head behind the eyes. 

Hind ocelli closer together than from occipital carina as i.o : 1.4 and then from nearest eye 
margin as i-o : 2-5. Postocellar region wider than long as 2-5 : i-o. Tarsal pulvillus on hind 
basitarsus only about one-third the width of basitarsus, though one-half the width on front legs. 
Penis-valve as in Text-fig. 24. 



178 R. B. BENSON 

Holotype $. TURKEY: Trabzon, Soganli Gecidi, 2,600 m., on flowers of Euphorbia, 
27. v. 1962 (Guichard & Harvey). B.M. (N.H.). 

Paratypes. Same data, 6^, 47$. B.M. (N.H.). 

This remarkable species is to be distinguished at once from all others in the 
arcuata-group by the green ground colour and by the punctation of the head and 
thorax, which are dull all over, with dense even coriaceous surface sculpture without 
coarser punctures: in all the other species known to me the head and thorax are 
coarsely punctured, i shining in places between the punctures especially on the 
temples of the head and mesoscutellum, but the underthorax and frons of head are 
usually dull with coarse surface sculpture. 

CUNEALA Zirngiebl, 1956 

This group of species was previously included in Tenthredo but is readily recognized 
by the long mouthparts (prelabium longer than greatest eye-measure) and the 
subtruncate apex of the labrum. 

Whether this genus is the true Cuneala is not entirely certain, as Zirngiebl gave 
no full description of either the genus or type-species C. tricolor. He says merely 
that the mouthparts are elongate as in Amauronematus , 7 the clypeus prism-shaped 
and the colour as in Tenthredo parviceps (Konow). He also says that the type is 
deposited in the Staatliches Naturkunde Museum, Stuttgart (where unfortunately 
it cannot now be found), and in correspondence Dr. Zirngiebl tells me he has no 
other representative of this species in his own collection. 

KEY TO MALES AND FEMALES 

1 Face and tegula entirely black. Hind tibia and tarsus yellow, with orange 

apices or almost entirely infuscate. Clypeus with coarse irregular punctures 
obscuring the obsolescent reticulate surface sculpture. Hind tarsus of <$ 
longer, but of $ shorter than hind tibia. $ either with densely pubescent 
basal tergites or with shining interspaces between punctures on scutellum . 2 

Face or tegula ^ marked with yellowish white. Hind tibia and tarsus yellow 
with infuscate apices. Clypeus with evident reticulate surface sculpture 
all over between scattered shallow punctures. Hind tarsus of both $ and $ 
longer than hind tibia. $ with 3 basal tergites shining and with only very 
sparse pubescence, but scutellum and mesopleura dull with dense microsculp- 
ture in the interspaces between punctures ...... 3 

2 (i) $ with basal tergites dull all over with dense pubescence; middle tergites 

without red markings. Interspaces between punctures on mesoscutellum 
dull with dense surface sculpture. 

$ body black with yellowish white apical margins to 4th, 5th and 6th 
tergites and sometimes ist and yth also. Legs black with tibia and tarsus 
yellowish white with orange apices; <J as $ but apical margins usually re- 
stricted to 4th and 5th tergites. 10-12 mm. Mountains of C. EUROPE. 

koehleri (Klug) 

7 This is not strictly true of what I here call Cuneala, because in Amauronematus it is the prementum 
and cardo that are elongate, while the glossa, paraglossa, galea and palps are shorter than in the species 
of the related genus Nematus, which has not got elongate mouthparts (c.f. Benson, 1958, figs. 388 and 
389). In the group called here Cuneala all the mouthparts are lengthened, especially the glossa, para- 
glossa and palps. 



HYMENOPTERA FROM TURKEY 

$ with ist-3rd tergites shining medially with only very short and sparse pubes- 
cence; o* middle tergites usually marked with red. Interspaces between 
punctures on mesoscutellum mostly shining with only obsolescent surface 
sculpture. 

Colour extremely variable : <$ and $ can be as in koehleri, but more commonly 
the 3rd and often 4th and 5th tergites in $ are marked with orange or more 
extensively yellow (of 140 $ in B.M. only 18 are without some orange on 
abdomen) ; and in the commonest form of <$ the whitish apical margins to 
the tergites are missing and the 3rd and 4th segments are entirely covered 
by a red band, while the legs are entirely black (85 out of 96 $ in B.M. are 
coloured like this); in some forms the body and legs are entirely black 
(f. atrata Andre). 9-5-12 mm. Mountains of TURKEY and eastwards to N. 
IRAN (1,000-2,400 m.) ...... ^conftnis (Konow, 

3 (i) Larger species, 11-14 mm. long. Face with clypeus, labrum and base of 

mandibles yellowish white. Clypeus with apical excision about as deep as 
half its total length. Scutellum flattened medially. 

Abdomen black with ist tergite i white, and 3rd-6th with entire apical 
margins and 2nd and yth with apical margins laterally white ... 4 

Smaller species, 8-10 mm. long. Face with clypeus and labrum black in $ but 
i white in $. Clypeus with apical excision usually about one-third and at 
most less than half its total length. Scutellum evenly convex, not flattened 
medially. 

Tegula black with pale apical margin. $ abdomen black with ist and 
3rd-yth tergites i margins apically with white and 3rd and 4th may be 
orange ; $ black but 3rd or 3rd and 4th tergites may be yellow. Pubes- 
cence on head and thorax long, so that the longest hairs are about as long as 
diameter of an ocellus. Mesonotum with interspaces between the punctures 
dull with surface sculpture. TRANSCAUCASIA and N. IRAN . ^longipes (Konow) 

4 (3) Tegula mainly yellow with a small black basal spot. Pubescence on head, 

mesonotum and mesopleura shorter (longest hairs up to three-quarters of 
diameter of lateral ocellus). $ and $ with ist tergite enitrely white. Meso- 
notum with interspaces between punctures dull with surface sculpture. 
TURKEY ...... f* atnasiensis (Kriechbaumer) sp. rev. 

Tegula mainly black with a yellow apical margin. Pubescence on head and 
thorax longer (longest hairs about as long as diameter of lateral ocellus), 
ist tergite in 9 entirely white but in <$ black with only apical margin white. 
Mesonotum with interspaces between punctures shining. S.E. EUROPE 
(HUNGARY, JUGOSLAVIA, ALBANIA, CORFU, GREECE) . . dahli Klug 8 

\Cuneala confinis (Konow, 1886) 

\Allantus parviceps Konow, 1898. 

TURKEY, C., N., N.E. and E. : Kayseri (i) ; Bolu (i and 3) ; Cankiri and Kasta- 
monu (2); Trabzon (3, 4, 14, 15 and 16); Gumusane (7 and 8); Rize (8). 1,000- 
2,500 m., 96 (?, 135$, I2.v-9.viii. Many on flowers of Geranium psilostemum 
Ledeb. 

IRAN: C. Elburz, Kandavan Pass, 9 km. above Siahbishe, 2,350 m., i $, 7.vi.i966 
(D. B. Baker). Taken in " Lush vegetation (? former cultivation) bordering stream 
in semi-desert (thorn-cushion) zone ". 

TURKEY, TRANSCAUCASIA and IRAN. 

8 Synonym of C. dahli: Allantus xanthorius Kriechbaumer, syn. n. 



i8o R. B. BENSON 

Cuneala longipes (Konow) 

IRAN: Mazandaran, Chalus-Shahsavar Coast of Caspian Sea, 34^, 219, 24.!!! 
to 6 . vi . 1966, and 9 <, 5 9, 20 . iv to n . v . 1967 (D. B. Baker] . Mainly collected from 
flowers of Ranunculus in woodland and in forest clearings. 

IRAN and TRANSCAUCASIA. 

Cuneala amasiensis (Kriechbaumer) 

TURKEY, W., S.W. and C. : Bursa (2) ; Aydin (2) ; Ankara (14) ; Corum (2) ; Amasya 
(i, 2, 3, 5, 7, 9 and 14). 50 to 560 m., 25 <, 28 $. 

Not known outside TURKEY. 

TENTHREDININAE 
SCIAPTERYGINI 
ELINORA Benson 

The following syntypes of Allantus found to belong to this genus in Konow's 
collection have been examined and the lectotypes labelled. The species are differen- 
tiated in the keys that follow. 

Allantus andrei Konow. LECTOTYPE 9 selected from i $ and i 9- 
A. antigae Konow. LECTOTYPE <j>, with abdomen still intact, labelled " Bar- 
celona " (i.e. " Hisp. prov. Catalonia " of original description), selected from 2 <$ 
and 2 9 one of which had its saw mounted but no abdomen intact. 

A. contiguus Konow. LECTOTYPE 9 with mesopleuron bearing only a yellow 
fleck as in original description, selected from i ^ and 4 9, three of which have 
mesopleura almost entirely yellow. 

A. dusmeti Konow. LECTOTYPE $, selected from i <$ and i 9- 
A. nigritarsis Konow. LECTOTYPE 9, selected from i <$ and i 9- 
A. obscuratus Konow. LECTOTYPE 9, with yellow hind tibia and tarsus as in 
original description, selected from 2 $ and 4 9, three of which have hind tibia 
infuscate behind. Elinora obscurata (Konow) comb. n. 

A. striatipes Konow. LECTOTYPE 9 selected from i ^ and i 9- 
In the keys the following species, described from S.E. EUROPE or TRANSCAUCASIA, 
and thought also to belong to this genus, had to be omitted as no specimens have 
been available for study: Allantus frivalskyi Mocsary, A. limbifer Mocsary and 
A. pubescens Andre. 

The key to the males will be found to be more unsatisfactory than that to the 
females, on which most of the species are based, as many of the females have not 
yet had males correlated with them. 

KEY TO FEMALES 

i Orbits extensively yellow (at least hind orbits lined with yellow for their whole 

length) ............ 2 

At least hind orbits mainly black (at most lower gena yellow) ... 6 



HYMENOPTERA FROM TURKEY 181 

2 (i) Clypeus with front lobes flat and rounded or truncate apically ... 3 
- Clypeus with front lobes raised and acute apically. E. MEDITERRANEAN 

\maculata (Kriechbaumer) 

3 (2) ist tergite black with at most separated pale flecks. Clypeus with surface flat 

and punctate ........... 4 

ist tergite with a broad yellowish white apical band. Clypeus convex and 

shining between sparse punctures. CAUCASUS. 7-5-9 mm. . ornata (Andr6) 

4 (3) More punctate species. Mesopleura, scutellum and ist tergite with evident 

punctation and sculpture. Abdomen with tergites black and i laterally 
pale-margined apically .......... 5 

Sparsely punctate species. Mesopleura, scutellum and ist tergite shining and 
almost impunctate. Abdomen with medial orange band covering tergites 
2-3 and all tergites with a yellowish white apical lateral fleck each side. 

Stigma infuscate apically. Hind tibia with black-tipped apex; inner spur 
about as long as tibial breadth. 11-12 mm. MOROCCO dulcis sp. n. (p. 184) 

5 (4) Clypeus dull with fine surface sculpture. Stigma infuscate apically. ISRAEL. 

10-5 mm. ......... stolida sp. n. (p. 185) 

Clypeus shining between coarse punctures. Stigma unicolorous. 

This species varies greatly in the amount of yellow on the orbits, antennal 
segments and tergites. In S.W. EUROPE the coxae are mostly black and 
the femora, tibiae and tarsi i black-lined; in some of the N. African forms 
the legs are almost entirely pale. S.W. EUROPE and N. AFRICA (PORTUGAL, 
SPAIN, MOROCCO, ALGERIA (including Ahagger Mountains in Sahara Desert) , 
TRIPOLITANIA and CYRENAICA) ..... xanthopus (Spinola) 9 

6 (i) Tegula at least partly black ......... 7 

Tegula entirely pale ......... . . 10 

7 (6) Inner hind tibial spur longer than apical width of tibia .... 8 

Inner hind tibial spur shorter than apical width of tibia .... 9 

8 (7) Antennae and mesopleura entirely black. Head and thorax dull with surface 

sculpture between dense punctures. 8-5-10-5 mm. SPAIN and N. AFRICA 

limbalis (Spinola) 10 

Antenna with basal segment and mesopleura with yellow fleck. Head and 
mesonotum shining between fine punctures, though on mesopleura surface is 
dull between the punctures. 10-11 mm. SPAIN and N. AFRICA (MOROCCO 
and ALGERIA) ........ vilarrubiai (Conde) 

9 (7) Gena pale below. Abdomen black apart from apical lateral margins of tergites. 

8-1 1 mm. N. AFRICA ....... deserta (Enslin) 11 

Gena entirely dark. Abdomen with red girdle covering segments 3-5 and 
segments from 6 and following with increasing white apical margins so that 
9 is entirely white. 9 mm. E. MEDITERRANEAN . . . asiatica (Enslin) 
10 (6) 2nd and following tergites black with yellowish white lateral apical marginal 
flecks which, on following tergites, become more extensive till they form con- 
tinuous apical bands, and on posterior segments cover the whole tergites . 10 
2nd or 3rd and following tergites mainly orange so that the lateral apical yel- 
lowish white flecks, which become continuous apical bands on posterior 
segments, are inconspicuous. 

Stigma scarcely darker apically than basally. 8-5-10-5 mm. E. MEDI- 
TERRANEAN to N. IRAN ....... coniensis (Enslin) 

9 Synonyms of E. xanthopus: \Allantus pectoralis Kriochbaumer, syn. n. \Macrophya cognata 
Kirby, syn. n. ^M. corynetes Kirby, syn. n. fM. jugurtha Kirby, syn. n. \Allantus striatipes Konow, 
syn. n. \A. andrei Konow, syn. n. ^A. tunetensis Konow, syn. n. Tenthredo adequata Enslin, 1910, 
syn. n. Allantus diversipes Pic, 1925, syn. n. Tenthredo limbergorum Forsius, 1930, syn. n. T. afra 
Benson, 1930, syn. n. fT. sahariensis Benson, 1954, syn. n. 

10 Synonyms of E. limbalis: ^Allantus balteatus Kriechbaumer, syn. n. A. gribodoi Konow. 

11 Synonym of E. deserta: \Elinora guichardi Benson, syn. n. 



182 R. B. BENSON 

11 (10) ist tergite almost entirely pale and stigma infuscate apically (pallipes complex) n 

ist tergite with at least broad base black up to one-third of its medial length, 

or stigma entirely pale .......... 14 

12 (n) Abdomen less extensively pale: 3rd and often following tergites without con- 

tinuous pale apical band. Front mesonotal lobe entirely black or with at 

most obsolescent pale flecks. . . . . .. . . 13 

Abdomen more extensively yellowish white: 3rd tergite with continuous pale 
apical band and 4th onwards mainly pale. Front mesonotal lobe with con- 
spicuous white lateral borders. 9-10 mm. TURKESTAN pallipes (Freymonth) 

13 (12) Larger species (n mm.). Hind femur often with apical half black. Scutel- 

lum dull and densely punctate, scarcely convex. 11-12 mm. TURKESTAN 

^dissidua (Konow) 

Smaller species (8-iomm.). Hind femur entirely pale. Scutellum shining 
and sparsely punctate, strongly convex. 9-11 mm. S. and W. CASPIAN 
Coast . caspia (Andre) 

14 (10) All femora and tibiae infuscate behind . . . . . . . 15 

Femora at most infuscate only at base, and tibiae at extreme apex . . 17 

15 (14) Gena black ............ 16 

Gena pale below. N. AFRICA. 9-10 mm. .... \obscuvata (Konow) 

16 (15) Scutellum all black. Pronotum and mesopleura almost entirely yellow. Stigma 

scarcely darker apically. E. MEDITERRANEAN. 7-5-8 mm. barbalis (Enslin) 

Scutellum with pale spot. Pronotum with only hind margin and mesopleura 
with only a small spot pale. Stigma conspicuously bicoloured with apical 
half black. N. AFRICA. 10 mm. ..... rufonigra (Andre) 

17 (14) Wings subhyaline. Prelabium shorter than head capsule. Abdomen with at 

least posterior tergites laterally pale-margined apically. ist tergite often 
i punctate and with surface sculpture between punctures. Inner hind 
tibial spur shorter than apical width of tibia . . . . . . 18 

Wings infuscate. Prelabium longer than head capsule. Abdomen black with 
middle tergites (3-5) red but other not pale-margined apically even laterally 
shining and very sparsely punctured species: ist tergite impunctate and 
without surface sculpture. Inner hind tibial spur longer than apical width of 
hind tibia. 10-11 mm. S.E. EUROPE . . . ^sabariensis (Mocsary) 

1 8 (17) Abdomen with unbroken lateral yellow stripe (tergites are entirely yellow 

laterally), or more extensively yellow. Antenna usually 8-segmented. 
Clypeus i shining between coarse punctures with front lobes acute or 
truncate, ist tergite often without pale apical stripe. W. EUROPE and 
N. AFRICA ............ 19 

Abdomen without lateral yellow stripe (tergites are black basally right up to 
lateral margin). Antenna g-segmented. Clypeus dull with dense surface 
sculpture and with front lobes pressed flat, ist tergite with continuous 
pale apical stripe. 10-12 mm. C. EUROPE to E. MEDITERRANEAN 

flaveola (Gmelin) 

19 (i 8) ist tergite without pale apical band dorsally ...... 20 

ist tergite with pale apical band developed dorsally. 4th and following 

tergites with continuous pale apical band. 10-12 mm. ATLANTIC EUROPE 

(ENGLAND, FRANCE, BELGIUM, PORTUGAL, SPAIN) . ^dominiquei (Konow) 

20 (19) Stigma unicolorous yellow scarcely darker apically 

Pale lateral stripe of abdomen with straight upper margin . . . 21 
Stigma at least partly infuscate . . . . . . . . .22 

21 (20) Clypeus slightly convex and shining between scattered punctures. 7-5-10 mm. 

SPAIN and N. AFRICA . . .. . . . algeriensis (Magretti) 12 

12 Synonyms of E. algeriensis: \Allantus tricolor Kriechbaumer, syn. n. \A. contiguus Konow, 
syn. n. 



HYMENOPTERA FROM TURKEY 183 

Clypeus pressed flat against labrum and with dense surface sculpture between 

punctures. E. MEDITERRANEAN. 7-5 mm. . . \parvula (Kriechbaumer) 

22 Smaller species (under 8mm.). Stigma with at least lower half infuscate. 

Abdomen yellow-brown (? greenish in life) with a continuous black medial- 
dorsal vitta. E. MEDITERRANEAN . . . \vittata (Kriechbaumer) 
Larger species (10-12 mm.). Stigma with apex infuscate. Abdomen with 
the basal tergites black and only the lateral portions yellow, but the yellow 
is progressively more extensive on the apical tergites which beyond the 4th 
segment have also yellow apical margins. SPAIN and N. AFRICA 

h (ic tic (i (Spinola) 13 

KEY TO MALES 

1 Tegulae marked with black ......... 2 

Tegulae entirely pale .......... 8 

2 (i) Inner hind tibial spur longer than apical width of tibia. Stigma strongly 

infuscate apically ........... 3 

Inner hind tibial spur much shorter than apical width of tibia. Stigma either 

unicolorous or infuscate apically ........ 4 

3 (2) Head above entirely dull with dense surface sculpture. N. AFRICA (MOROCCO 

and ALGERIA). 8-10 mm. ...... limbalis (Spinola) 

Head with shining unsculptured patches on temples. SPAIN and N. AFRICA 

(MOROCCO and ALGERIA). 9-10 mm. .... vilarrubiai (Conde) 

4 (2) Stigma yellow often i brownish apically. 8-10 mm. ..... 5 

Stigma black apically with a pale base. 9-10 mm. N. AFRICA rufonigra (F. Andre) 

5 (4) Hind femur at least pale-lined on the outer side ...... 6 

Hind femur almost entirely black. Stigma brownish. 8-10 mm. N. AFRICA 

obscurata (Konow.) comb. n. 

6 (5) Clypeus with front lobes rounded or truncate apically and flattened . . 7 

Clypeus with front lobes acute and convex. 9-10 mm. E. MEDITERRANEAN 

maculata (Kriechbaumer) 

7 (6) Head above antennae, mesonotum and mesopleura i punctate and with fine 

surface sculpture. 8-10 mm. W. MEDITERRANEAN (SPAIN, PORTUGAL 

and N. AFRICA) ........ xanthopus (Spinola) 

Head above antennae, mesonotum and mesopleura smooth and shining with 

at most, sparse obsolescent punctures. 8-9 mm. N. AFRICA deserta (Enslin) 

8 (i) Abdomen with at least one medial or apical tergite entirely pale. Often over 

8 mm. ............ 9 

Abdomen yellow-brown with a continuous black medial dorsal vitta and no 
entirely pale tergites. Hind legs mainly pale yellow except for extreme 
base of femur, apex of tibia and most of tarsi. Mesopleura yellow above and 
black below together with mesosternum. Under 8 mm. E. MEDITERRANEAN 

vittata (Kriechbaumer) 

9 (8) Abdominal tergites 1-7 each at least marked with black at the base medially 

but with the two apical tergites and hypopygium mainly yellowish white. 
Hind femur marked with black. Wings flavescent with unicolorous yellow 
stigma often i brownish below. Large species, 1012 mm. 10 

Abdomen with one or more of the middle tergites entirely reddish brown. If 
the apical tergites and hypopygium are mainly pale then the hind femur is 
not marked with black. Wings often hyaline and stigma usually infuscate 
apically or below. Often smaller species . . . . . . . n 

10 (9) Clypeus with front lobes truncate and flattened. Antenna 9-segmented. 

10-11 mm. C. EUROPE to E. MEDITERRANEAN . . flaveola (Gmelin) 

13 Synonyms of E. baetica: \Allanlus antigae Konow, syn. n. \A. dusmeti Konow, syn. n. 



184 R. B. BENSON 

Clypeus with front lobes acute and convex. Antenna 8-segmented. 10-13 

mm. E. EUROPE . . . . . . . dominiquei (Konow) 

11 (9) Crests above antennal sockets scarcely project beyond level of middle of the 

inter-antennal area .......... 12 

Antennal crests project strongly beyond level of middle of inter-antennal area . 13 

12 (11) Clypeus flat and densely punctate. 7-10 mm. 

algeriensis Magretti (N. AFRICA); boetica (Spinola) (SPAIN); dissidua (Konow), 
fulveota (Zhelochovstev) (TURKESTAN) and parvula Kriechbaumer 

(TURKEY) 
Clypeus strongly convex and shining between sparse punctures. 8-10 mm. 

E. MEDITERRANEAN ........ ornata (Andre) 

13 (n) Three apical tergites black, each with a yellowish white fleck each side . . 14 

Three apical tergites mainly yellow, each with yellowish white fleck each side 
and a. medial black fleck. Mesonotum black except for white fleck on 
scutellum. 9-11 mm. E. MEDITERRANEAN . . . coniensis (Konow) 

14 (13) Front lobe of mesonotum with lateral margins white, as is scutellum. Pale 

lateral flecks on apical tergites reach more than half way to middle of segment. 

8-9 mm. TURKESTAN ....... pallipes (Freymonth) 

Mesonotum usually entirely black. Pale flecks on apical tergites mostly 

confined to lateral ventral portions of tergites . . . . . . 15 

15 (14) Wings infuscate, with stigma pale above and infuscate below. Clypeus and 

mesopleura shining between scattered punctures. Inner hind tibial spur 
longer than apical breadth of tibia. S.E. EUROPE . . sabariensis (Mocsary) 

Wings hyaline; stigma pale basally and infuscate apically. Clypeus and meso- 
pleura above dull with fine microsculpture between punctures. Inner hind 
tibial spur shorter than apical breadth of tibia. S. and W. CASPIAN Coast 

caspia (Andre) 

Elinora maculata (Kriechbaumer, 1869) 

Allantus syriacus Andre. 
A. nigritarsus Konow. 
Tenthredo aulica Enslin, 1912. 

SYRIA, ISRAEL, TURKEY and TRANSCAUCASIA. 

Elinora ornata (Andre, 1881) 
Allantus ^discolor Konow. 

TRANSCAUCASIA. 

Elinora dulcis sp. n. 

$. Black with the following parts yellowish white: mandible base, labrum, clypeus, outer 
orbits to level of top of eyes, basal segment of antenna (rest of antennae missing in unique type), 
posterior half of pronotum, tegula, meso- and meta-scutella, fleck in middle of mesepisternum, 
fleck on metapleura, coxae (except their bases), femora, tibiae (except their extreme apices) 
middle of basitarsi, lateral fleck each side of ist and 2nd tergites, the 3rd, 4th and 5th tergites 
except for a lateral black fleck on each and that dorsally these three tergites are fulvous, and 
there is a lateral yellow fleck on the hind margin of tergites, 6, 7, and continuously on hind 
margin of 8th and gth. 

Wings slightly brownish subhyaline ; basal half of stigma, extreme apex of subcosta and basal 
two-thirds of costa yellow; rest of venation piceous. Length 12 mm. 



HYMENOPTERA FROM TURKEY 185 

Head slightly swollen behind the eyes and shining with sparse punctation. Clypeus slightly 
convex but with the front lobes pressed flat and with a few scattered punctures. Frons with 
a medial groove widening behind. Hind ocelli further apart than from hind margin of head as 
i-o : 0-9, but further from eye margin than from each other (POL : OOL as i-o : 1-6). 

Thorax strongly shining between widely-spaced obsolescent punctures ; mesoscutellum evenly 
convex; post-tergite shorter than shortest measure of a cencher. 

Legs with inner hind tibial spur about as long as apical breadth of tibia. 

Abdomen normal. 

Pubescence on head and thorax pale and up to a length as long as diameter of an ocellus. 

Holotype $. MOROCCO; Grand Atlas, Idni, 8. .1941 (K. M. Guichard}. B.M. 

(N.H.). 

Elinora stolida sp. n. 

$. Black with the following parts yellowish white: mandible base, labrum, clypeus, outer 
orbits to level of top of eyes, ist and upper side of 2nd antennal segments, pronotum, tegula, 
V-shaped outer margin of front mesonotal lobe and fleck on inner side of each of the lateral 
mesonotal lobes, anterior three-quarters of mesoscutellum, upper half of mesepisternum, small 
fleck on mesepimeron and on metapleura, coxae (except extreme bases), femora, tibiae (except 
extreme apices) basitarsus except for a fleck at extreme base and extreme apex, fleck on lateral 
hind margin of ist and following tergites progressively longer from the ist tergite backwards, 
so that on the 5th and following tergites the hind margin is continuously pale. 

Wings slightly brownish subhyaline; front half of stigma, costa, subcosta (front half of 
Sc + R) and veins at extreme base of wings yellow; rest of venation piceous. Length 10 mm. 

Head shining with very fine surface sculptures. Clypeus slightly convex with front lobes 
pressed flat and the whole surface dull with dense surface sculpture. Frons with a L-shaped 
medial groove. Hind ocelli further apart than from hind margin of head as i-o : 0-7, but 
further from the eye margin than from each other (POL : OOL as i-o : 1-7). 

Thorax shining between sparse punctures, which become denser at front of mesonotum and 
in middle of mesonotum, where they are scarcely further apart than the diameter of a puncture. 
Mesoscutellum slightly convex and not separated medially from the raised centre of the post- 
tergite which is about as long as the shortest measure of a cencher. 

Legs with inner hind tibial spur scarcely three-quarters as long as apical width of tibia. 

Abdomen normal. 

Pubescence on head and thorax pale and up to a length as long as the diameter of an ocellus. 

Holotype $. ISRAEL: Jerusalem, 15.111.1923 (P. A. Buxton). B.M. (N.H.). 

Elinora asiatica (Enslin) 
TURKEY. 

Elinora coniensis (Enslin) 
Allantus (Tenthredo) kareli Muche, 1962, syn. n. 

TURKEY: Ankara (57). I <$ and I $; Kayseri (on flowers of Lepidium, Muche, 1962). 
N. IRAN: W. Elburz, Qazvin, c. 760 m., 2 $, i $, 25.^.1967 (D. B. Baker). 
TURKEY, TRANSCAUCASIA and N. IRAN. 

Elinora caspia (Andre) 
S.E. RUSSIA, TRANSCAUCASIA and IRAN. 



186 R. B. BENSON 

Elinora barbalis (Enslin) 
SYRIA. 

Elinora flaveola (Gmelin) 
\Allantus orientalis Kriechbaumer. 

C. and E. EUROPE and TURKEY. 

f Elinora parvula (Kriechbaumer) comb. n. 
TURKEY. 

^Elinora vittata (Kriechbaumer) comb. n. 
ISRAEL and IRAQ. 

TENTHREDININAE 

SGIAPTERYGINI 
SCIAPTERYX Stephens 

Previous keys to world Sciapteryx were compiled by Konow, 1908, Kuznetzov- 
Ugamskii, 1929 and the Caucasian species by Dovnar-Zapolskii, 1930. 

The following species, from studies of the types, are now placed in other genera 
as shown: 

Sciapteryx galerita Konow (Sikkim) = Tenthredo galerita (Konow) comb. n. 

Sciapteryx kozlovi Konow (Tibet) = Tenthredo kozlovi (Konow) comb. n. 

Sciapteryx gilva Konow (Tibet) = Rhogogaster gilva (Konow) comb. n. 

Sciapteryx virescens Konow (Tibet) = Rhogogaster virescens (Konow) comb. n. 

Sciapteryx caucasica Dovnar-Zapolski (Caucasus) and S. montana Dovnar-Zapolski 
(Caucasus) cannot be placed because they are based on inadequately described 
males and are excluded from the key below. 

All the species of this genus have size 7-10 mm. 

KEY TO SPECIES MALES AND FEMALES 

1 Gena dull with dense surface sculpture between punctures or with punctures 

closer together than their diameters. $ hypopygium evenly or acutely 
emarginate each side of middle ........ 2 

Gena behind eyes, shining, with punctures obsolescent or more widely spaced 

than their diameters. $ hypopygium not emarginate each side of middle . 10 

2 (i) Fore wings infuscate or milky and Sc -\- R not longitudinally bicoloured. 

Inner hind tibial spur not longer than apical breadth of tibia and only 4-5 
times as long as broad. 

Hypopygium in $ slightly emarginate each side of middle ... 3 



HYMENOPTERA FROM TURKEY 187 

Fore wings subhyaline and the fused Sc + R paler in front (Sc) than behind (R) . 

Inner hind tibial spur longer than apical breadth of tibia and 7 or 8 times 
as long as broad ........... 7 

3 (2) Sc + R and also usually C in fore wing pale yellow or brown Wings milky 

or with the fore wings i infuscate. Stigma pale with or without dark apex 4 

Sc + R and C in fore wing mainly black. Wings almost uniformly infuscate. 
Stigma pale with dark apex. ATLANTIC EUROPE and MEDITERRANEAN 

jsoror Konow 

4 (3) Stigma of fore wing bicoloured with dark apex and pale base. C of fore wing 

pale as in Sc + R. Fore wings infuscate ...... 5 

Stigma pale without dark apex and C infuscate apically. Wings uniformly 

milky. E. MEDITERRANEAN ...... ^lactipennis Konow 

5 (4) Tegula entirely yellowish white ...... (costalis) 6 

Tegula with most of basal half black. E. MEDITERRANEAN . levantina Andr6 

6 (5) Inner orbits and lower half of outer orbits lined with white in <J and $; <J with 

face below antennae entirely white; with interantennal area pale marked 
CORSICA ....... ^costalis corcyrensis Benson 

Only the inner orbits above pale-lined in $ and interantennal area entirely 
black; in (J whole inner orbits and lower outer orbits pale-lined but clypeus 
marked with black. C. and E. EUROPE . . costalis costalis (Fab.) 

7 (2) Darker species, flagellum of antenna and, even in $, base of clypeus, most of 

face between antennae and clypeus, mesopleura, stigma black. Abdomen 
without a continuous pale lateral band and with no entirely pale apical 
sternites. Antenna with flagellar segments all longer than broad . . 8 

Paler species, underside of flagellum, most of clypeus and face below antennae 
even in $, fleck on mesopleura, most of stigma, a broad lateral band on 
the abdomen and at least i or 2 entire apical sternites yellowish white. 
Antenna with 7th and 8th segments as broad as long ($) or broader than 
long ($). Head and thorax with aeneus reflections. E. MEDITERRANEAN 

]laeta Konow 

8 (7) Inner orbits in $ and $ pale-lined to top of eyes. Head and thorax often with 

cupreous or aeneus metallic reflections. Stigma yellowish white infuscate 
below or at apex ........... 9 

Inner orbits in $ and $ at most only pale-lined on lower half, with a small fleck 
at top of eye. Head and thorax carbonarius or with slight bluish, purplish 
or greenish reflections. Stigma piceous. $ hypopygium slightly and evenly 
emarginate each side of middle. W. and C. EUROPE . consobrina Klug 

9 (8) Stigma pale infuscate below. Labrum and base of clypeus pale. $ hypo- 

pygium sharply excised each side of middle (as in laeta). E. MEDITER- 
RANEAN ....... . circassica Dovnar-Zapolski 

Stigma pale at base and infuscate at apex. Labrum and clypeus black except 
at most for a lateral pale fleck each side of clypeus. $ hypopygium evenly 
emarginate each side of middle (as in consobrina). S.E. EUROPE (THRACE) 

byzantina sp. n. (p. 188) 

10 (i) Stigma bicoloured with infuscate apex or mainly infuscate . . . . u 

Stigma mainly yellowish white .... ..... 12 

11 (10) Stigma bicoloured with infuscate apex. Veins C and Sc -f M in fore wing 

entirely pale. Tegula black with narrow white apex. All legs black with a 
white line above or on outside. S.E. MEDITERRANEAN \cleopatra Benson 

Stigma mainly infuscate. Only base of C, and Sc pale. Tegula black. Legs 

mainly black. CRIMEA ...... semenowi Jakovlev 

12 (10) Tegula entirely yellowish white. TURKESTAN . . . nigriventris Andre 14 

Tegula black with only the apex yellowish white. TURKESTAN hauseri Forsius 

14 Synonym of S. nigriventris: Sciapteryx costalis vernalis Kuznetzov-Ugamskil, syn. n. 



i88 R. B. BENSON 

^Sciapteryx soror Konow 

TURKEY, E. : Gumusane (n). i <$, i $. 

Elsewhere only in ATLANTIC and MEDITERRANEAN EUROPE (Britain, W. France, 
Switzerland, Spain and Italy). This species forms an Atlantic /Continental species 
pair with 5. costalis F. (Benson, 1952). 

\Sciapteryx lactipennis Konow 
TRANSCASPIA, IRAN, TRANSCAUCASIA and ISRAEL. 

Sciapteryx levantina Andre 
SYRIA and LEBANON. 

\Sciapteryx laeta Konow 

TURKEY, E. : Gumusane (n). 4^. 8$. 
TURKEY and TRANSCAUCASIA. 

* Sciapteryx circassica Dovnar-Zaposkii 

TURKEY, E. : Gumusane (n); Trabzon (3 and 17). 31 <$, 28 $. 
TURKEY and TRANSCAUCASIA. 

Sciapteryx cleopatra Benson, 1954 
EGYPT and ISRAEL. 

^Sciapteryx byzantina sp. n. 

$. Black with the following parts yellowish white: labial palps, inner orbits above level of 
antennae, hind edge of pronotum, front half of tegula, extreme apex of femora and i outer 
edge of tibiae, i base of basitarsus and apical margin of all tergites. Wings hyaline: basal 
half of stigma, extreme apex and base of costa and subcosta (front half of Sc + R) yellowish 
white ; rest of venation piceous. Length 8-9 mm. 

Head and thorax normal, dull with dense punctures closer together on gena than their diameters 
and the interspaces i with fine surface sculpture. Antenna with all flagellar segments longer 
than broad. Inner hind tibial spur longer than apical width of tibia and about x 7 or x 8 
times longer than its own basal width. Abdomen dull with dense transverse surface sculpture. 

Hypopygium emarginate each side of middle saw. $ as in $ apart from sexual characters 
but that the clypeus has a small whitish fleck each side and the pale margin of the inner orbits 
reaches to bottom of eyes. 

Holotype $. TURKEY, N.W. : Istanbul, Belgrat Orman, at sea level, 25.iii.62 
(Guichard & Harvey}. B.M. (N.H.). 

Paratypes. Same data, 15^, i $. TURKEY, N.E.: Rize at sea level, i 9, 
22.iv. 1959 (Guichard}. B.M. (N.H.). 



HYMENOPTERA FROM TURKEY 189 

TENTHREDINAE 
MACROPHYINI 

P achy protasis rapae (L.) 

TURKEY, N.E. : Giresun (7); Trabzon (3, 4, 15 and 17). 20 $, 20 $. 
N. temperate HOL ARCTIC species. 

MACROPHYA Dahlbohm 

Two major sections of the genus are defined as species-groups in the present paper 
and keys are given for the species of these groups from EUROPE and S.W. ASIA. 

Macrophya punctumalbum (L.) 
TURKEY, N. and C. : Amasya (6); Samsun (8). 10 $. 
N. IRAN: Mazandaran, Panjak Rostaq, 860-900 m., i $, 29.^.1967 (D. B. Baker). 

Macrophya albicincta (Schrank) 

TURKEY, W., N. and N.E. : Bursa (3, 9); Bolu (i); Samsun (9, 15, etc.); Giresun 
(7); Trabzon (4, 7, 17); Rize (2). 36 $, 39 ?. 

EUROPE, to TRANSCAUCASIA, N. IRAN, W. SIBERIA. 

Macrophya crassula (Klug) 

TURKEY, W., S., N., and N.E. : Bursa (3 and 9); Mersin (6 and 7); Sinop (i); 
Samsun (9, 10, 21, etc.); Trabzon (17, etc.); Artvin (2). 33 , 39 $. 

C. and S. EUROPE to TRANSCAUCASIA. 

Macrophya consobrina Mocsary 
TURKEY and TRANSCAUCASIA. 

Macrophya pallidilabris A. Costa 

TURKEY, W. and N.E. : Bursa (4, 5 and 9); Gumusane-Trabzon (n). 6 $, 6$. 
C. EUROPE to TRANSCAUCASIA. 

Macrophya erythrocnema A. Costa 

TURKEY, N.E.: Rize (8); Artvin (3). 2 $. 
S. EUROPE to TRANSCAUCASIA. 

^Macrophya rufipes orientalis Moscary 

For nee M. orientalis Mocsary; Benson, 1954 vide M. diaphenia sp. n. 
TURKEY, C. and E. : Ankara (35); Gumusane (7). 3$. 

ENTOM. 22, 4. 9 



igo R. B. BENSON 

In this race the red markings on the middle tergites (3 and 4) are entirely absent. 

In N. Africa the species is represented by M. rufipes ruficincta Konow, stat. n., 
which differs from the typical race in having a black clypeus and black hind tibia 
and tarsus in both sexes. 

Macrophya sanguinolenta (Gmelin) 

TURKEY, N.E. and E. : Trabzon (3); Rize (8); Erzurum (9). 2 <, i <j>. 
EUROPE, TURKEY and TRANSCAUCASIA. 

The BLANDA-DUODECIMPUNCTATA-group 

Representatives of this species-group are recognized by the presence of a small 
appendage behind the metepisternum as a separate sclerite between the hind coxa 
and the ist tergite. 

In addition to the species from EUROPE and S.W. ASIA keyed below, M. apicalis 
Smith (JAPAN), M. ignava Smith QAPAN), M. infumata Rohwer (E. SIBERIA) as well 
as M . fumator Norton (N. AMERICA) belong to this group. 

KEY TO MALES AND FEMALES OF EUROPE & S.W. ASIA 

1 Appendage to mesepisternum (between hind coxa and ist tergite) always 

infuscate. Abdomen black with lateral or ventral white flecks. Either 
hind legs red, or hind tibia or tarsus white-flecked. Wings yellowsh or 
subhyaline ............ 2 

Appendage to mesepisternum often mainly white. Abdomen red-banded, 
without white flecks, or hind tibia and tarsus entirely black. Wings often 
infuscate ............ 4 

2 (i) Hind legs not marked with red ......... 3 

Hind legs partly red ........... 7 

3 (2) Wing membrane yellowish; stigma and venation yellowish brown. Hind tarsus 

black. EUROPE, ASIA MINOR, and SIBERIA to JAPAN duodecimpunctata (L.) 
Wing membrane subhyaline. Stigma and venation infuscate. Hind tarsus 

with apical segments white. TRANSCAUCASIA and IRAN . longitarsus Konow 

4 (3) Hind coxa with a white fleck laterally or beneath ..... 5 

Hind coxa entirely black. EUROPE, S.W. ASIA to IRAN and SIBERIA 

annulata (Geoffrey) 

5 (4) (J with hind tarsus normal (4th segment much longer than broad in ventral 

view (Text-fig. 27). $ hypopygium only slightly emarginate apically each 

side ............. 6 

$ with hind tarsus swollen (4th segment scarcely longer than broad in ventral 
view) (Text-fig. 28). $ hypopygium excised apically each side of middle. 
C. TURKEY oedipus sp. n. (p. 194) 

6 (5) (J abdomen mainly black with white flecks below. Pubescence on head and 

thorax up to as long as diameter of an ocellus. Penis-valve Text-fig. 33. 
(9 unknown). Mountains of E. TURKEY .... hatnata sp. n. (p. 194) 
(J abdomen red-banded and without white flecks below. Pubescence on 
head and thorax not longer than half diameter of an ocellus. Penis-valve 
as in Text-fig. 32. EUROPE, ASIA MINOR, to IRAN and SIBERIA blanda (Fabricius) 



HYMENOPTERA FROM TURKEY 



191 



7 (2) Metepisternum with lower two-thirds shining and without surface sculpture 
apart from hair follicles. Frons shining between punctures and without a 
continuous medial groove. Wings subhyaline. $ with pale yellow front 
and middle femora and tibia strongly contrasting with its hind legs and 
with the red front and middle legs of the . EUROPE, ASIA MINOR to 
N. IRAN ......... diversipes (Schrank) 

Metepisternum dull with dense surface sculpture all over. Frons dull with 
dense surface sculpture and with a deep medial groove reaching from front 
ocellus to interantennal area. 

Wings uniformly slightly infuscate. <J with front and middle femora and 
middle tibia reddish as in the hind legs and as in the $. IRAN 

diaphenia sp. n. (p. 195) 



ham. 




oed. 




mac 




vest. 




FIGS. 27-30. 

ENTOM 22. 4. 



hind tarsus: 27, Macrophya hamata sp. n. ; 28, M. oedipus sp. n. 
29, Tenthredo maculata; and 30, T. vestita. 



iga 



R. B. BENSON 



Macrophya duodecimpunctata (L.) 

TURKEY, N.: Samsun (2 and 6). 
EUROPE, TURKEY and SIBERIA to JAPAN. 



32 



bland. 





FIGS. 31-34. Macrophya penis-valves in lateral view: 31, diaphenia sp. n. ; 32, blanda; 
33, hamata sp. n. ; and 34, oedipus sp. n. 



HYMENOPTERA FROM TURKEY 193 

Macrophya longitarsus Konow 

IRAN: Mazandaran, Chalus-Shahsavar coast of Caspian Sea. v-vi.i966 (D. B. 
Baker) . 
TRANSCAUCASIA and IRAN. 

Macrophya annulata (Geoffrey) 

TURKEY, C. and N.: Ankara (15); Samsun (9). 4^, 2$. 

IRAN: Mazandaran, Chalus-Shahsavar coast of Caspian Sea, .1966 (D. B. Baker). 

EUROPE, S.W. ASIA to IRAN and SIBERIA. 

Macrophya blanda (Fabricius) 

TURKEY, W., S., C., N.E. and E. : Usak, Bulgaz Dag Mts., Kizilcahaman, 1,000 m., 
i <$, 2 $, 26.v-i6.vi.i965 (Demelt Coll., Gembloux); Mersin (6, 7 and 14); Ankara 
(31 and 39) ; Amasya (3, 7, 9 and 13) ; Konya (3) ; Samsun (10) ; Giresun (2) ; Trabzon 
(3 and 15); Artvin (3); Erzurum (5, 6). 18 <$, 48 $. 

IRAN: Mazandaran, Chalus-Shahsavar Coast of Caspian Sea, v.1966 (D. B. Baker}. 
EUROPE, TURKEY, TRANSCAUCASIA, IRAN and SIBERIA. 




35a 





36a 




FIGS. 35-36. Macrophya saws: 35, prasinipes; 36, minerva sp. n. 35a~36a. Enlarged 
loth and nth teeth from apex of saws: 35a, prasinipes; 36a, minerva sp. n. 



194 R - B - BENSON 

*Macrophya oedipus sp. n. 

(Text-figs, 28, 34) 

<$. Black with tergites 3 and 4 and sometimes also 5, 6 and 7 +_ red laterally and the follow- 
ing parts white: labrum, +_ front of clypeus, posterior extra sclerite on metepisternum ; front 
face of all coxae and +_ lateral fleck on hind coxae, front face of fore and middle femora and 
tibiae, front tarsus, most of middle tarsus (except line on back of basitarsus, and the claw- 
bearing segment) and lower edge of hind femur. Wings hyaline ; stigma and venation black. 

Length 9-11 mm. 

Head with mouthparts normal; malar space about one-fourth of distance between antennal 
sockets; 3rd antennal segment about as long as 7 + 8 + 9; frontal area not clearly denned; 
frontal groove running from front ocellus to interantennal area; hind ocelli closer together 
than from hind margin of head as i-o : 1-4; and from eye margin as i-o : 2-3. 

Thorax with rounded convex mesoscutellum, medially carinate over posterior half and with 
minute extra appendage attached to posterior margin of metepisternum, which is dull medially 
with dense surface sculpture. Legs with swollen hind tarsus so that the 3rd tarsal segment in 
ventral view is almost as broad as long (Text-fig. 28). Claws bifid with the inner tooth longer 
and stouter than the end tooth. Abdomen normal with penis- valve as in Text-fig. 34. 

Punctation: Head very densely and coarsely punctured on the frontal area and though the 
punctures are sparse on the temples, the surface between the punctures is dull with dense 
coriaceous sculpture. On the thorax the punctures are smaller and more evenly spaced but the 
interspaces are likewise dull with dense coriaceous sculpture. Abdomen dull with dense trans- 
verse alutaceous sculpture. 

Pubescence clothing whole of insect very short, nowhere exceeding about half diameter of an 
ocellus. 

$ as $ in colour but labrum and clypeus + infuscate, the front half of the mesoscutellum is 
white, the legs are black except for white as follows: a fleck on the underside of the fore and 
middle coxae and the side of the hind coxae, a line on the front of the fore femur, tibia and 
basitarsus and +_ on the apex of the middle femur. In structure as ^ except for sexual segments; 
hypopygium emarginate behind each side of middle. Length 12-13 mm. 

Holotype $. TURKEY, C. : Amasya, 500 m., 31^.1959 (K. M. Guichard}. B.M. 

(N.H.). 

Paratypes. TURKEY, C. : Amasya, 500 m., 5 J, i $, 22-24. .1959, 2 , 2$, 
29 . v . 1959, 4 <$, 1-2 . vi . 1959, i $, 6 . vi . 1959, 2 <J, i $, 9 . vi . 1959, and 400 m., 6 <$, 
4$, 30. v. 1959 (K. M. Guichard). B.M. (N.H.). 

*Macrophya hamata sp. n. 

(Text-figs. 27, 33) 

cJ. Colour as in M. oedipus but that the clypeus is black except for its front margin, the fore 
and middle femora and tibiae are only lined with white in front, the hind coxae, though white- 
flecked below, have no lateral white fleck, the abdomen has the red reduced to at most a lateral 
fleck each side of the 2nd tergite, but has a white fleck on the hind lateral corners of each of the 
3rd to 6th tergites, and white apical margins to the sternites. The pubescence is much longer 
than in M. oedipus, blanda and annulata; on head, mesonotum and mesopleura the hairs are up 
to as long as the diameter of an ocellus. Penis-valve as in Text-fig. 33. Otherwise as in 
M. blanda. $ unknown. 

Holotype <$. TURKEY: Artvin, above Artvin, 1,800 m., 6.vi.ig6z (K. M. 
Guichard 6- D. Harvey). B.M. (N.H.). 



HYMENOPTERA FROM TURKEY 195 

Paratypes. i $ (same data), i $ (likewise but at 900 m.) and i $, TURKEY: 
Trabzon, Hamsikoy, 1,24501., 22. v. 1962 (Guichard & Harvey). B.M. (N.H.). 



Macrophya diversipes Schrank 

TURKEY, W., C., N.E. and E. : Mugla (8); Mersin (8); Ankara (9, 37 and 39); 
Corum (2) ; Amasya (i, 2, 3, 5 and 7) ; Artvin (2 and 6) ; Gumusane (i and 10) ; 
Erzurum (i, 4, 5, 8, 10 and n). 26 <$, 41 $. 

C. and S.E. EUROPE, TURKEY, TRANSCAUCASIA and N. IRAN. 



Macrophya diaphenia sp. n. 

(Text-fig. 31) 
Macrophya orientalis Mocsdry; Benson, 1954, nec Mocsdry. 

cJ. Black with the following parts yellowish white: labrum, front half of clypeus, fore tibia 
and tarsus, apex and inner side of middle tibia, and middle tarsus (except apical segment) ; 
trochanters brown; red are the fore femur, middle femur and basal outer side of middle tibia, 
hind femur and tibia (except its extreme apex). Length 11-12 mm. 

Wings brownish-infuscate, paler towards base. Basal half of stigma and basal three-fourths 
of costa brown; rest of venation black. 

Head contracted behind eyes. Malar space one-third as long as distance between antenna 
or diameter of front ocellus. Antenna with 3rd segment about as long as 4th and 5th. Frons 
with a deep medial groove reaching from front ocellus to interantennal area. Hind ocelli 
closer together than from hind margin of head as i-o : 1-7, and from eye margin as i-o : 2-0. 

Thorax with slightly convex mesoscutellum, carinate medially for posterior half; metepister- 
num dull medially with dense surface sculpture and with a small posterior appendage. 

Legs with hind tarsus swollen so that the 4th segment in ventral view is almost as long as 
broad. 

Abdomen normal. penis (Text-fig. 31) not distinguished from that of M. diversipes. 

Punctation dense on head and thorax; interspaces shining on head, but on mesonotum and 
mesopleura the interspaces are dull with dense surface sculpture. 

Pubescence dense all over insect and very short so that nowhere on head and mesonotum are 
the individual hairs as long as the diameter of an ocellus. 

? as (J but the stigma of the fore wing is entirely yellowish brown and all the legs have the 
femur and tibia red and the trochanters and tarsus infuscate, and the hind tarsal segments are 
not so swollen, the 4th being x ij as long as wide in ventral view. Hypopygium slightly 
emarginate each side of middle. Saw not distinguished from that of M. diversipes. 

Holotype. <?. S.W. IRAN: Kuh Send, nr. Bazuft, (Escalem Coll.}. B.M. (N.H.). 

Paratypes. Same data, 3 <$, 2 $ ; N. IRAN: Elburz Mountains, Mt. Demavend, 
2,800 m., i $, vii.i966 (L. G. Higgins). B.M. (N.H.). 

The POST 1C A -group 

Species with eyes converging in front and abdomen with at least a white band 
on the ist tergite and lateral flecks on some of the others ; at least front and middle 
legs, hind coxa and base of hind femora mainly yellow or orange. 



ig6 R.B. BENSON 

KEY TO MALES AND FEMALES 

1 Either mesonotum entirely dull with surface sculpture between fine punctures 

and front lobe usually with dark adpressed pubescence shorter than diameter 
of front ocellus or hind legs with tibia and tarsus mainly black. ($ hypo- 
pygium strongly emarginate) ......... 2 

Mesonotum with the interspaces between the punctures shining and the 
pubescence on the front lobes pale, and usually outstanding and longer than 
diameter of front ocellus ; hind legs with tibia mainly yellow or orange . . 6 

2 (i) Stigma of fore wing and hind tibia and tarsus mainly orange or yellow (except 

in (J aphrodite). Pubescence short, dark and adpressed on front lobe of 
mesonotum ............ 3 

Stigma of fore wing and hind legs mainly black (subsp. arpaklena Ushinskii, 
from TURKESTAN and N. IRAN has $ hind tarsus with segments 2-4 white 
as in (J, and abdomen with pale flecks reduced). EUROPE, N. AFRICA, ASIA 
MINOR to TURKESTAN ....... montana (Scopoli) 

3 (2) Head above dull with dense surface sculpture between punctures. Post-tergite 

of scutellum longer medially than a cencher ...... 4 

Head above shining between punctures, which are very sparse and obsolescent on 

middle of temples. ASIA MINOR to S.W. IRAN . . . ^cyrus Benson 

4 (3) Costa of fore wing infuscate, darker than the yellow stigma. Scutellum with 

depressed posterior third dull with dense sculpture and with anterior two- 
thirds shining between sparse punctures ....... 5 

Costa of fore wing coloured the same as the yellow stigma. Scutellum with the 
posterior half dull with dense sculpture and the anterior half shining between 
scattered punctures. LEBANON, ISRAEL, TRANSCAUCASIA and IRAN 

ottomana Mocsdry 

5 (4) Temples between postocellar region and eyes dull, with surface sculpture 

between the dense punctures: <$ hind tibia mainly orange and brown. C. and 

S.E. EUROPE and TURKEY ....... postica (Brulle) 

Temples between postocellar region and eyes with shining interspaces between 

scattered punctures; $ hind tibia mainly black. CYPRUS . f aphrodite Benson 

6 (i) Stigma of fore wing darker apically than basally. Hind tarsus black. $ 

hypopygium not emarginate each side ....... 7 

Stigma of fore wing unicolorous brown. Hind tarsus reddish brown. $ 
hypopygium strongly emarginate behind each side of middle. S.E. EUROPE 
and TURKEY ........ superba Tischbein 

7 (6) Abdominal segments of $ ringed with pale apical margins to tergites and sternites ; 

$ abdomen yellow above, except for the bases of the tergites medially, and 
entirely yellow laterally and ventrally. Saw with shallow teeth (Text-fig. 
35 and 35a). TRANSCAUCASIA ..... prasinipes Konow 

Abdomen in $ black except for the yellow apical margins of the tergites medially ; 
( unknown). Saw with prominent teeth (Text-figs. 36 and 36a). GREECE 

minerva sp. n. (p. 197) 



Macrophya montana montana (Scopoli) 

TURKEY: Amasya (2, 5 and 7); Samsun (10) ; Sivas (3); Rize (8). 12 <$, 14$. 
(Ssp. arpaklena Ushinskij in TURKESTAN and N. IRAN, Mazandaran on the Caspian 
Sea Coast, iv-v . 1965-1966 (D. B. Baker)). 

EUROPE, N. AFRICA, TURKEY, TRANSCAUCASIA, IRAN and TURKESTAN. 



HYMENOPTERA FROM TURKEY 197 

*-\ Macrophya cyrus Benson, 1954 

TURKEY, C. and E. : Amasya (i, 2, 3 and 7); Nigde (4); Gumusane (13); Erzurum 
(i, 4, 5, 6 and 8). 51^,49?. 

TURKEY and S.W. IRAN. 

Macrophya ottomana Mocsary 
LEBANON, ISRAEL, TRANSCAUCASIA and IRAN. 

Macrophya postica (Brulle) 

TURKEY, S.W., S.C., N., N.E. and E. : Aydin (i and 2) ; Mugla (2, 5 and 7) ; Antalya 
(5 and 89); Mersin (i, 2 and 4); Ankara (12); Amasya (i, 2, 5, 7 and 14); Sinop 
(3 and 4); Samsun (10); Artvin (6); Gumusane (13); Erzurum (4). Many J and $. 

f Macrophya aphrodite Benson, 1954 
CYPRUS. 

Macrophya superba Tischbein 

TURKEY, N., C., N.E. and E.: Izmit, Alem Dag, 600 m., i $, 26-30 . vi . 1966 
(Demelt Coll. Gembloux); Ankara (21, 35 and 39); Cankiri, Isik Dag, 1,200 m., 2 $, 
25. vi. 1966 (Demelt Coll. Gembloux); Amasya (i, 2, 3 and 7); Nigde (5); Artvin (2) 
Gumusane (i); Erzurum (4). 35 J, 42$. 

S.E. EUROPE and TURKEY. 

Macrophya prasinipes Konow 
TRANSCAUCASIA. 

Macrophya minerva sp. n. 

(Text-figs. 36, 36a) 

$. Black with the following parts yellowish white: palps of mouthparts, labrum, clypeus, 
hind margin of pronotum, tegula (except at base), apices of coxae, trochanter, upper outer 
face of hind coxa, femora (except small inner basal fleck) and tibiae of all legs (except apex of 
hind pair) and fore and middle tarsus (except for broken black line on inner side), apical margin 
of ist tergite and lateral dorsal margin of 3rd and following tergites. 

Wings subhyaline ; base of stigma, base of C, and Sc (front half of Sc + R) ; rest of venation 
piceous becoming paler at extreme base. Length: n mm. 

Head normal, slightly contracted behind eyes. Malar space about as long as 2 facets of com- 
pound eye. Flagellar segments of antenna all longer than broad. Occipital carina complete. 
Posterior ocelli about as far apart as from occipital carina (POL = OOCL) ; and closer together 
than from nearest compound eye margin (POL: OOL i-o : 1-5). 

Thorax normal, without the extra metepisternal sclerite between the hind coxa and ist tergite. 
Mesoscutellum slightly convex. 

Abdomen normal; hypopygium only slightly emarginate each side. Saw (Tet-xfigs. 36 
and 36a). 



ig8 R. B. BENSON 

Punctation. Head shining with dense punctures on frons, inner orbits and postocellar region 
but becoming obsolescent on temples and finer and denser on outer orbits. Thorax shining 
between punctures as far apart as their diameters, though more widely spaced on the middle 
of the mesoscutellum and much denser on the underthorax. Abdomen densely alutaceous 
transversely. 

Pubescence on head and thorax dense and pale and up to about as long as twice the diameter 
of a hind ocellus; on abdomen about half this length and pale and dense and recumbent. 

Holotype $. GREECE: Soufli, i $, 5.v.T_96o (Guichard & Harvey}. B.M. (N.H.). 
Distinguished from other members of the postica-group in the key above. 



NEMATINAE 
CLADIINI 

Cladius pectinicornis (Geoffrey) 

TURKEY, N. and C.: Samsun (6 and 21); Amasya (2, 3 and 5); Tokat (4). 4 <$, 

4$. 

EUROPE, N. AFRICA, SIBERIA to JAPAN, TRANSCAUCASIA, N. IRAN and HIMALAYAS. 

Cladius ordubadensis Konow 

GREECE: Samothrace, Therme, 3^, r $, i6.viii.i962 (Guichard & Harvey}. 
CRIMEA, CYPRUS, LEBANON and TRANSCAUCASIA. 

Priophorus morio Lepeletier 
EUROPE, TRANSCAUCASIA, IRAN, SIBERIA, to JAPAN and N. AMERICA. 

Priophorus pallipes Lepeletier 
EUROPE, TRANSCAUCASIA, SIBERIA to JAPAN and N. AMERICA. 

Priophorus rufipes Lepeletier 
Priophorus ulmi (L.) Konow, nee L. 

EUROPE, TURKEY, TRANSCAUCASIA. 

Trichiocampus viminalis (Fallen) 
EUROPE, TRANSCAUCASIA, SIBERIA to JAPAN and N. AMERICA. 

PSEUDODINEURINI 

Pseudodineura fuscula (Klug) 
EUROPE, TRANSCAUCASIA and SIBERIA. 



HYMENOPTERA FROM TURKEY 199 

HOPLOCAMPINI 
HOPLOCAMPA Hartig 

The species of this genus segregate naturally into 3 distinct groups, all feeding as 
larvae in the fruits of Rosaceae: two groups attached to the Prunoidea and one to 
the Pomoidea. Unfortunately the major group characters, the shape of the clypeus, 
the direction of the inner eye-margins and the pubescent clothing of the under- 
thorax, have not been mentioned in previous descriptions, and it is not therefore 
always possible to place species I have not examined. 

KEY TO SPECIES-GROUPS 

1 Clypeus broadly and shallowly emarginate in front to a depth less than half 

total length of clypeus, or (i.e. flava) mesepisternum densely pubescent all 
over and eyes large so that they are closer together in front than ij times 
their own greatest length. Attached to Prunus (Prunoidea) ... 2 

Clypeus deeply excised in front to a depth of at least half the total clypeal 
length. Lower mesepisternum and mesosternum at least partly glabrous 
(with areas devoid of hair follicles). Eyes always further apart in front than 
ij times their length. Attached to Pomoidea : Eurasian species : alpina 
Zetterstedt (Sorbus) ; aviae Benson (Sorbus) ; brevis Klug (Pyrus) ; crataegi 
Klug (Crataegus) ; ephippiata Konow (Malus), pectoralis Thomson (Crataegus) ; 
plagiata Klug (Amelanchier) ; pyricola Rohwer (Pyrus} ; ? sino-birmana Malais; 
testudinea Klug (Malus} (Text-fig. 39). 

Nearctic species (apart from introductions) : alpestris Rohwer (A melan- 
chier); bioculata Rohwer (Amelanchier}; halcyon Norton (Amelanchier}; 
montanicola Rohwer (Amelanchier}; oskina Ross (Crataegus); pallipes Mac- 
gillivray (Amelanchier}; ritcheri Ross (Crataegus); and texas Ross (Crataegus} 

testudinea - group 

2 (i) Eyes strongly diverging in front (where they are further apart than i times 

their length). Mesepisternum below and mesosternum mainly glabrous. 
Nearctic species: idaho Ross; lacteipennis Rohwer (Text-fig. 38); marlatti 
Rohwer; nalema Ross; ? Ashmead ..... oregonensis- group 
Eyes subparallel in front (where they are closer together than i times their 
length). Mesepisternum and mesosternum densely pubescent. 

Eurasian species : chrysorrhoea Klug ; flava L. ; ? formosana Malaise ; minuta 
Klug; prunicola sp. n. ; rutilicornis Klug (Text-fig. 37). 

Nearctic species: cookei Clarke; oskima Ross .... ^fava- group 

Hoplocampa testudinea (Klug) 
EUROPE, TURKEY and TRANSCAUCASIA. 

Hoplocampa brevis (Klug) 
EUROPE and TRANSCAUCASIA. 

^Hoplocampa crataegi (Klug) 

TURKEY, W. : Bursa (i and 3). 8 $, 9 $ from Crataegus blossom. 

A very dark form in which in the $ the antenna, postocellar region and occiput 



200 



R. B. BENSON 




37 rut. 




38 



lact, 




test. 




flav. 




41 




FIGS. 37-42. 37-40. Hoplocampa faces: 37, rutilicornis; 38, lacteipennis ; 39, testudinea; 
and ^o,flava. 41-42. Hoplocampa prunicola sp. n. : 41, penis-valve; and 42, saw. 



HYMENOPTERA FROM TURKEY 201 

of head, almost the entire mesonotum and the hind tibia are black and the stigma 
of the fore wing is slightly infuscate basally. 

C. and S. EUROPE and TURKEY. 

Hoplocampa pectoralis Thomson 

N. IRAN: Mazandaran, Pankaj Rostaq, i 9, 29.^.1967 (D. B. Baker). 
EUROPE, TRANSCAUCASIA and N. IRAN. 

Hoplocampa flava (L.) 

TURKEY, S.: Antalya (n). i <$ at flowers of Prunus insititia L. 
EUROPE, ISRAEL, TURKEY and TRANSCAUCASIA. 

*Hoplocampa prunicola sp. n. 

(Text-figs. 41, 42) 

?. Reddish yellow with the following parts black or piceous: ocellar region, front edge and 
hind margin of pronotum, middle line of front mesonotal lobe, most of lateral mesonotal lobes, 
middle line and hind edge of scutellum and its post-tergite, mesepimeron and metepimeron 
above, sclerotised parts of ist-3rd tergites, and following tergites above. Wings hyaline; 
stigma and venation very pale brown. Length 4 mm. 

Head with clypeus evenly emarginate in front to depth of about one-third of its total length. 
Eyes subparallel in front and closer than i| times their own greatest measure apart in front 
(1-4 : i-o). Hind ocelli further apart than from hind margin of head as i-o : 0-6, and from eye 
margin as i-o : 0-8 (= POL : OOL). 

Thorax normal but with medial suture of front mesonotal lobes obsolescent. 

Abdomen with hypopygium but slightly emarginate each side and ovipositor scarcely as 
long as hind-femur, without and trochanter, and much shorter than hind tibia. Saw as in 
Text-fig. 42. Pubescence pale and evenly spread over head and thorax, much shorter than 
diameter of front ocellus. 

Punctation: surface of head and thorax shining between shallow follicles; abdomen shining 
with faint alutaceous surface sculpture. 

o* as $ but with black more extensive to cover postocellar region and edges of frontal area 
and occiput of head, the whole of the thorax except the pronotum and tegula, and the whole of 
the abdomen except the hypopygium; penis- valve as in Text-fig. 41. 

Holotype $. TURKEY: Izmit, Karamursel, i $, 23.111.1961 on Prunus (H. 
Birkadesler}. B.M. (N.H.). 

Paratypes. Same data, 3 <, 2 ?. B.M. (N.H.). 

This species is very closely related to H. chrysorrhoea Klug, from which it is 
distinguished by the almost entirely yellow head in the female and beaked penis- 
valve in the male (c.f. Text-fig. 41 with Benson, 1958, fig. 398 etc.); in the form 
of its penis-valve it would appear more to resemble H. minuta Klug. 

The type of the Siberian species, H. ephippiata Konow (<), was borrowed for 
comparison but this species was found to be scarcely separable structurally, including 
the penis- valve, from H. pectorialis Thomson. 



202 R. B. BENSON 

DINEURINI 

Caulocampus necopinus Zhelochovtsev 
TRANSCAUCASIA. 

NEMATINI 

* Stauronematus compressicornis (Fabricius) 

TURKEY, C., N.E. and E.: Corum (2); Amasya (13); Rize (i); Trabzon (6 and 9); 
Gumusane (2). 4 ^, 3 $. 

HOL ARCTIC species. 

Pristiphora abbreviata (Hartig) 

CYPRUS: Pera Pedi, 700 m., i $, 4.iv.i952 (G. A. Mavromoustakis). 
C. and S. EUROPE and TRANSCAUCASIA, introduced into N. AMERICA. 

Pristiphora subbifida (C. G. Thomson) 

TURKEY, N.: Samsun, i $, 20. 11.1959 (Guichard). 
EUROPE and TURKEY. 

Pristiphora conjugata (Dahlbom) 

TURKEY, C. : Kula, 2 $, 1.1952 (Plant. Protection Institute), " Larva on poplar". 
EUROPE, ASIA MINOR, SIBERIA to JAPAN. 

Pristiphora fulvipes (Fallen) 

TURKEY, N. and N.E. : Samsun (8); Trabzon (4). 2$. 
EUROPE, ASIA MINOR, TRANSCAUCASIA and SIBERIA. 

Pristiphora sp. near fulvipes 

TURKEY: Trabzon (12). 

Pristiphora crassicornis (Hartig) 

TURKEY: Zonguldak (i); Artvin (3). 2 $. 
EUROPE and TURKEY. 

^Pristiphora rufipes Lepeletier 
TURKEY: Tokat (i). i $. 

HOLARCTIC. 



HYMENOPTERA FROM TURKEY 203 

*Pristiphora ambigua (Fallen) 

TURKEY: Giresun (7). 2 <$, 20 $. 
N. and C. EUROPE and ASIA MINOR. 

* Amauronematus sternalis Enslin 

TURKEY, C. : Ankara (13) (Beynam Forest, 1,000 m.). 2 $. 
EUROPE, TURKEY and SIBERIA. 

*Nematinus luteus (Panzer) 

TURKEY, C. and N.E. : Tokat (3); Rize (3). i <$, 2 $. 
EUROPE and TURKEY. 

Euura mucronata (Hartig) 
EUROPE, TRANSCAUCASIA, SIBERIA, HIMALAYAS and N. AMERICA. 

*Phyllocolpa puella (C. G. Thomson) 

TURKEY, N.E. : Giresun (2). i $. 
EUROPE and TURKEY. 

Phyllocolpa leucosticta (Hartig) 

TURKEY, E.: Gumusane (Soganli Gecidi at 1,800 m.). 2 $, 3 $. 
EUROPE, TURKEY, TRANSCAUCASIA and SIBERIA. 

Pontania vesicator (Bremi-Wolf) 
EUROPE and TRANSCAUCASIA. 

Pontania bridgmanii (Cameron) 
EUROPE and TRANSCAUCASIA. 

Pontania viminalis (L.) 
EUROPE and TURKEY. 

Pontania dolichura (C. G. Thomson) 
Subarctic-Subalpine EUROPE, TRANSCAUCASIA, SIBERIA, N. AMERICA. 

Croesus septentrionalis (L.) 

TURKEY, N.E. : Trabzon (14). i $. 
EUROPE and TURKEY. 



204 R. B. BENSON 

* Croesus varus (Villaret) 
TURKEY, N.E. : Rize (i). 2 ?. 

HOLARCTIC. 

*Nematus lucidus (Panzer) 

TURKEY, N.E. : Artvin (3). i $. 
EUROPE, CYPRUS and TURKEY. 

Netnatus salicis (L.) 
EUROPE and TRANSCAUCASIA. 

Netnatus ribesii Scopoli 
EUROPE and TRANSCAUCASIA, introduced into N. AMERICA. 

Netnatus myosotidis (Fallen) 

TURKEY, C., N., N.E. and E. : Amasya (13); Samsun (6 and n); Giresun(7); 
Trabzon (7, 9, 15 and 16); Gumusane (3). 7$, 9$. 

EUROPE, ASIA MINOR, W. SIBERIA. 

*Nematus oligospilus (Forster) 

TURKEY, C. and N.E. : Ankara (32); Trabzon (9); Rize (i). i <$, 2 <j>. 
HOLARCTIC to HIMALAYAS. 

* Netnatus viridis (Stephens) 

TURKEY (N.E.) : Trabzon (14 and 15) ; Rize (5). i J, 2 -. 
EUROPE and ASIA MINOR. 

Pachynematus rumicis (L.) 
EUROPE, IRELAND, TRANSCAUCASIA, SIBERIA and ALASKA. 

*Pachynematus obductus (Hartig) 

TURKEY, N.E.: Trabzon (18). i . 

HOLARCTIC species reaching ICELAND and GREENLAND. 

Pachynematus vagus (Fabricius) 

TURKEY, C. and N.E.: Tokat (i); Trabzon (4). 2 $. 
HOLARCTIC species. 



HYMENOPTERA FROM TURKEY SYMPHYTA 205 

*Pachynematus ? clitellatus (Lepeletier) 

TURKEY (E.): Erzurum, Cakmak mountains, 2,200 m., i $, 24. v. 1960 (E. S. 
Brown). 
EUROPE and ASIA MINOR. 

REFERENCES 
BENSON, R. B. 1930. Nine Sawflies requiring new names. Entomologist 63 : 107. 

1936. A new species of Mocsarya Konow from Syria (Hymenoptera Symphyta). Proc. 
R. ent. Soc. Lond. (B) 5 : 2-3. 

1938. European Sawflies of the genus Xyela Dalman (sens, lat.) (Hymenoptera Symphyta). 
Proc. R. ent. Soc. Lond. (B) 7 : 32-36, figs. 1-5. 

1943. Studies in Siricidae. . . . Bull. ent. Res. 34 : 27-51, 17 figs., 3 tables. 

1946. Classification of the Cephidae. Trans. R. ent. Soc. Lond. 96 : 89-108, 39 figs. 

1951-58. Hymenoptera Symphyta. Handbk. Ident. Br. Insects 6 (2a-c) : 1-252, 815 figs. 

1954. Some Sawflies of the European Alps and the Mediterranean Region (Hymenoptera : 
Symphyta). Butt. Br. Mus. nat. Hist. (Ent.) 3 (7) : 267-296, 31 figs. 

1955. Sawflies (Hymen. Symphyta) of Israel. Bull. Res. Coun. Israel B, 4 : 451-456. 

1956. Studies in Dolerini (Hymenoptera : Symphyta). Proc. R. ent. Soc. Lond. (B) 
25 : 55-63, 16 figs. 

i958a. Sawflies (Hymenoptera Symphyta) of the Apennine Mountains of Italy. 
Memorie Mus. civ. Stor. nat. Verona 6 : 321-325. 

1959. Revision of the European Sawflies of the Tenthredo arcuata-complex (Hymenoptera 
Tenthredinidae) . Proc. R. ent. Soc. Lond. (B) 28 : 93-102, figs. i-n. 

1962. A revision of the Athaliini (Hymenoptera : Tenthredinidae). Bull. Br. Mus. nat. 
Hist. (Ent.) 11 (7) : 333-382, 58 text-figs. 

1965. The Classification of Rhogogaster Konow (Hymenoptera : Tenthredinidae). Proc. 

R. ent. Soc. Lond. (B) 34 : 105-112, figs. 1-21. 

1966. A new genus of the Lycaotini (Blennocampinae) in Turkey (Hymenoptera Tenthre- 
dinidae). Proc. R. ent. Soc. Lond. (B) 35 : 75-77, 6 figs. 
BYTINSKI-SALZ, H. 1956. Coleoptera and Hymenoptera from a journey through Asia Minor, i. 

Istanb. Oniv. Fen. Fak. Mecm. (B) 21 (4) : 211-229 ( 221 )- 
CONDE, O. 1934. Versuch einer Revision einiger mitteleuropaischen Aprosthema-Arten 

(Hymenoptera, Tent.). Folia zool. hydrobiol. 7 : 20-30. 
DADURIAN, A. B. 1958. [On the horntail and sawfly fauna of Armenia.] Izv. Akad. Nauk 

armyan. SSR 11 : 91-101. 

DALLA TORRE, K. W. VON. 1894. Catalogus Hymenopterorum 1. Lipsiae. 
DovNAR-ZAPOLSKif, D. P. i93o. Neue oder wenig bekannte Chalastogastra. Russk. ent. 

Obozr. 24 : 86-94. 
1931. Ein Uebersicht ueber die Blattwespen des Nord-Kaukasischen Gebietes (Hym. 

Symphyta). Izv. sev.-Kavkaz. Kraev. Sta. Zasch. Rast. 6-7 : 33-62, 10 figs., 2 tables. 
ENSLIN, E. 1910. Das Tenthrediniden-Genus Allantus Jur. Russk. ent. Obozr. 10 : 335-372. 

1912-18. Die Tenthredinoidea Mitteleuropas, 1-7. Dt. ent. Z., Beihefte, 1912-1917. 

1920. Die Blattwespengattung Tenthredo L. (Tenthredella Rohwer). Abh. zool.-bot. Ges. 
Wien 11 : 1-96. 

1927. Die Tenthrediniden (Hymenoptera) der Kamtschatka-Expedition, 1908-1909. 
Ezheg. zool. Muz. 37 (1926) : 363-381. 

ERMOLENKO, V. M. 1964. [A Study of the Hymenoptera Symphyta of Ukrainian Woodlands]. 

Trudy Inst. Zool., Kyyiv 20 : 98-119. 
FORSIUS, R. 1919. tiber einige von Bequaert in Nord Afrika gesammelte Tenthredinoidea. 

Ofvers finska VetenskSoc. Fork. 60 A (13) : i-n (2). 

1930. Uber die von Mag. phil. Hakan Lindberg in Spanien gesammelten Tenthredinoiden. 
Commentat. biol. 3 (15) : 1-6. 



206 R. B. BENSON 

GUICHARD, K. M. & HARVEY, D. H. 1967. Collecting in Turkey 1959, 1960 and 1962. Bull. 

BY. Mus. nat. Hist. (Ent.) 19 (4) : 223-250. 
GUSSAKOVSKII, V. V. 1935-1947. Fauna SSSR. Insectes Hymenopteres. Chalastogastra, 

1 : 1-452, 82 figs. 2 : 1-234, I21 fig 8 - 
KERENSKII, H. N. 1926. [Sawflies and Horntails of Rostov-on-Don. Proc. N. Cauc. Peoples 

University 9]; [not seen] vide Dovnar-Zapolskil, 1931. 
KONOW, F. W. 1899. Chalastogastrorum novae species et varietates, quas D. Escalera ex 

Asia Minor reportavit. Actas Soc. esp. Hist. nat. (2) 8 : 203-207. 

1905. Lydidae, Siricidae and Tenthredinidae. Genera Insect. 27-29. 

1908. Lydidae. Blattwespen aus Tibet. Ezheg. Zool. Muz. 13 : 9-25. 

KUZNETZOV-UGAMSKII, N. N. 1929. Tenthredinologische Notizen (Hymenoptera Tenthre- 

dinoidea). Zool. Anz. 80 : 49-55. 

MALAISE, R. 1935- New Genera of Tenthredinoidea and their Genotypes (Hymen.). Ent. 
Tidsk. 56 : 160-178. 

1945. Tenthredinoidea of South-East Asia, with a general zoogeographical review. 
Subfamily Tenthredininae. Opusc ent. Supplementum 4 : 1-288, plates I-XX, figs. 1-57. 

1964. New Genera and Species of the Subfamily Blennocampinae (Hym. Tenthred.). 
Ent. Tidsk. 85 : 20-39, figs. 1-3. 

MATSUMURA, F. 1911. Erster Beitrag zur Insekten-Fauna Sachalin. /. Coll. Agric. Hok- 
kaido imp. Univ. 4 : 1-145, 2 plates. 

MAXWELL, D. E. 1955. Comparative Internal Larval Anatomy of Sawflies (Hymenoptera 
Symphyta). Can. ent., Supplement 1 : 1-132, figs. 1-153. 

MUCHE, W. H. 1962. Die Tenthredinidae (Hym.), meiner Anatolienausbeute ii. Reichen- 
bachia 1 : 17-20. 

1964. Beitrag zur Kenntnis der Blattwespen Bulgariens mit Beschreibung einer neuen 
Amasis-Krt. Ent. Z. Frankf. a.M. 74 : 17-24. 

i964a. Dolerus asper megapteroides nov. ssp. (Hymenoptera Symphyta). Reichenbachia 
4 : 31-33- 

Pic, M. 1916. Hymenopteres nouveaux d'Orient et du nord de 1'Afrique. Echange 32 : 23-24. 

1917. Notes Hymenopterologiques. Echange 33 hors texte : 1-4. 

1918. Description diverses de Tenthredinides et notes. Echange 34, hors texte : 1-4. 

1925. Hymenopteres nouveaux. Echange 41 : 12-15. 

Ross, H. H. 1951. In MUESEBECK, C. F. W., KROMBEIN, K. V. & TOWNES, H. K. Hymen- 
optera of N. America north of Mexico, Synoptic Catalog. Agriculture Monogr. 2, 1420 pp. 

WEIFFENBACH, H. 1967. Eine neue Rhogogaster aus Kleinasien. NachrBl. Bayer. Ent. 
[In press]. 

ZHELOCHOVTSEV, A. 1928. Uber palaarktische Dolerinae. Zool. Anz. 79 : 105-112. 

1941. On the Sawflies of Armenia. Sb. Trud. gos. zool. Muz. 7 : 123-153. 
ZIRNGIEBL, L. 1956. Blattwespen aus Iran. Mitt, munch, ent. Ges. 46 : 322-326. 

INDEX TO NEW SPECIFIC NAMES AND TO NEW SYNONYMY 
Synonyms in italics 

adequata, Tenthredo, 181 (footnote) balcana, Tenthredo, 159 (footnote) 

afra, Tenthredo, 181 (footnote) balteatus, Allantus, 181 (footnote) 

africanus, Dolerus, 137 bensoni, Urocerus augur, 116 

albata, Tenthredopsis, 158 (footnote) benthini, Tenthredo, 159 (footnote) 

amurensis, Calameuta, 122 , , . c , . 

, . . ' a it \ bleusei, Schizoceros, 128 

andrei, Allantus, 181 (footnote) 

andrei, Tenthredopsis, 158 (footnote) yt> Amasis ' J 3 2 

antigae, Allantus, 181 (footnote) brevis > Pachycephus, 118 

apicicornis, Monoplopus, 122 brunnescens, Thomsonia, 160 (footnote) 

armenius, Ateuchopus, 123 byzantina sp. n., Sciapteryx, 188 



INDEX 



207 



camtschatkali, Tenthredopsis, 157 (footnote) 

casia, Tenthredopsis, 159 (footnote) 

dementi, Kokujewia, 128 

cognata, Macrophya, 181 (footnote) 

contiguus, Allantus, 182 (footnote) 

continentalis , Thomsonia, 160 (footnote) 

corcyrensis, Tenthredopsis, 158 (footnote) 

CORN ARIA, 152 

coronatus, Thrinax, 134 

corynetes, Macrophya, 181 (footnote) 

deannulata, Thomsonia, 160 (footnote) 
debilis, Arge, 126 

decoratus sp. n., Characopygus, 119 
diana ssp. n., Tenthredo maculata, 174 
diaphenia sp. n., Macrophya, 195 
diversipes, Allantus, 181 (footnote) 
dulcis sp. n., Elinora, 184 
dusmeti, Allantus, 183 (footnote) 

excisa, Perineura, 158 (footnote) 
euphorbiae sp. n., Tenthredo, 177 

f estiva, Calameuta, 122 

fuscicornis, Thomsonia, 160 (footnote) 

fuscitarsis, Selandria, 135 

gastrica, Hylotoma, 128 
geniculatus Lepeletier, Dolerus, 137 
gribodoi, Allantus, 181 (footnote) 
grombczevskii, Cephus, 122 
guichardi, Elinora, 181 (footnote) 
guichardi sp. n., Tenthredopsis, 162 

haematica sp. n., Corynis, 130 
hamata sp. n., Macrophya, 194 
harveyi sp. n., Tenthredopsis, 163 
helveticus, Allantus, 171 
henschi, Schizoceros, 128 
hungarica, Tenthredo, 158 (footnote) 
hyrcanus sp. n., Dolerus, 141 
hyrcana sp. n., Scolioneura, 149 
hyrcana sp. n., Tenthredo, 172 

infernalis, Cephus, 122 

jugalis, Allantus, 171 

juguvtha, Macrophya, 181 (footnote) 

kareli, Allantus, 185 
kussariensis, Allantus, 177 

lacteore ssp. n., Aglaostigma aucupariae, 155 
lactiflua, Tenthredo, 158 (footnote) 



laevinota ssp. n., Kaliofenusa ulmi, 150 
limbergorum, Tenthredo, 181 (footnote) 
luminosus, Allantus, 171 

megapteroides $, Dolerus, 141 
merceti, Allantus, 171 
meridionalis , Euromostethus, 151 
minerva sp. n., Macrophya, 197 
montivagus sp. n., Dolerus, 138 

nazareensis, Allantus, 172 
nigriceps, Dolerus, 137 
nigripes, Schizoceros, 128 
nigritarsis, Hylotoma, 126 
nigrorufa, Thomsonia, 159 (footnote) 
nitens ssp. n., Eriocampa ovata, 154 
notatus, Cephus, 120 

oedipus sp. n., Macrophya, 194 
quadrannulata, Tenthredopsis, 159 (footnote) 

palestina, Kokujewia, 128 
pectoralis, Allantus, 181 (footnote) 
pisinna, Tenthredopsis, 159 (footnote) 
proximo,, Hylotoma, 126 
prunicola sp. n., Hoplocampa, 201 
punctatus, Tristactus, 115 

ruficornis, Thomsonia, 160 (footnote) 

sachalinensis , Tenthredo, 157 (footnote) 
sahariensis, Tenthredo, 181 (footnote) 
sanctus, Spatulocephalus, 118 
sarta, Amasis, 132 
seljuki ssp. n., Arge clavicornis, 127 
seljuki ssp. n., Dolerus anticus, 137 
serena, Allantus, 172 
stolida sp. n., Elinora, 185 
striatipes, Allantus, 181 (footnote) 
syriaca, Hylotoma, 125 

tanaiticus, Cephus, 120 
tenuicornis, Astatus, 124 
tischbeini, Tenthredo, 159 (footnote) 
tricolor, Allantus, 182 (footnote) 
triforis, Tenthredopsis, 159 (footnote) 
tunetensis, Allantus, 181 (footnote) 
turanica, Arge, 126 
turanicus, Cephus, 122 



vernalis, Sciapteryx, 187 (footnote) 
xanthorius, Allantus, 179 (footnote) 
zarudnyi, Arge, 126 




A LIST OF SUPPLEMENTS 

TO THE ENTOMOLOGICAL SERIES 

OF THE BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
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1. MASNER, L. The types of Proctotrapoidea (Hymenoptera) in the British 
Museum (Natural History) and in the Hope Department of Entomology, Oxford. 
Pp. 143. February, 1965. 5. 

2. NIXON, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera: 
Braconidae). Pp. 284; 348 Text-figures. August, 1965. 6. 

3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177; 
18 plates, 270 Text-figures. August, 1965. 4 45. 

4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172 ; 500 Text-figures. October, 

1965- 35s. 

5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156; 
475 Text-figures. November, 1965. 2 155. 

6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae & Drosophilidae. 
Pp. 129; 328 Text-figures. 3. 

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

335. 

8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the 
world species of Cleora (Lepidoptera : Geometridae). Pp. 119; 14 plates, 146 
Text-figures, 9 maps. February, 1967. 3 IDS. 

9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera : Rhopalocera). Pp. 509. August, 1967. 8 los. 

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rhopa- 
locera). Pp. 322 ; 233 Text figures, coloured frontispiece. September, 1967. 8. 

11. MOUND, L. A. A review of R. S. BagnalTs Thysanoptera Collections. Pp. 184; 
82 Text-figures. May, 1968. 4. 



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THE SUBGENERIC DIVISIONS OF 
THE GENUS BOMBUS LATREILLE ^ 
(HYMENOPTERA : APIDAE) 




O. W. RICHARDS 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 22 No. 5 

LONDON: 1968 



THE SUBGENERIC DIVISIONS OF THE 
GENUS BOMBUS LATREILL 
(HYMENOPTERA : APIDAE) 



GENUS BOMBUS LATREILLE & ^ 



BY 

O. W. RICHARDS 

Imperial College of Science and Technology, London 



Pp. 209-276, 39 Text-figs. 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 22 No. 5 

LONDON: 1968 



THE BULLETIN OF THE BRITISH MUSEUM 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

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

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

This paper is Vol. 22, No. 5 of the Entomological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 



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



Trustees of the British Museum (Natural History) 1968 



TRUSTEES OF 
THE BRITISH MUSEUM (N ATU R AL H ISTOR Y ) 

Issued 13 September, 1968 Price r los 



THE SUBGENERIC DIVISION OF THE 

GENUS BOMBUS LATREILLE 

(HYMENOPTERA : APIDAE) 

By O. W. RICHARDS 

CONTENTS 

Page 
INTRODUCTION . . . . . . . . . . . 211 

ALPHABETICAL LIST OF NAMES AVAILABLE FOR THE SUBGENERA OF Bombus 213 

NOTE ON GROUP-NAMES . . . . . . . . . 2l8 

CHARACTERS OF THE SUBGENERA OF Bombus ..... 219 

DESCRIPTIONS OF THE SUBGENERA . . . . . . .221 

KEYS TO SUBGENERA OF Bombus LATREILLE ..... 260 

Males ........... 260 

Females ........... 265 

REFERENCES ........... 273 

SYNOPSIS 

The described subgenera of Bombus Latreille are revised and listed and their characters 
discussed ; thirty-five subgenera are recognized as valid. Full synonymy is given and keys 
to males and females only are provided. 

THE current system of named subgenera for the large genus Bombus Latreille was 
first started by Vogt (1911) and Skorikov (1914, 1922). Originally, the subgenera 
were merely groups whose males possessed more or less closely similar genitalia 
and the names proposed for them were derived from the specific name of one of the 
included species. The system has gradually been elaborated and more names 
have been added; some of the old ones have tended to drop out because earlier 
names of von Dalla Torre (1880) were found to be available. Very few attempts 
have been made to provide a key to the sub-genera, many of which have never 
been described in detail. The females have been even more neglected than the 
males and though all sorts of characters have been indicated in the very scattered 
literature as helpful in recognizing the females of certain groups, no systematic 
account of the subgeneric classification of this sex is available. Some features 
of the European species were indicated by Kriiger (1917, 1920) and Pittioni (i939a) 
even gave a key of a somewhat artificial type to the East European subgenera. 
Neither of these authors, however, referred to the important characters which can 
be found in the sting. Some of these had been noted by Richards (1927) and the 
North American species have recently been fully illustrated by Hazeltine & Chandler 
(1964). There is clearly a need for proper descriptions of the subgenera and for 
a key to them which includes all of them and not merely those of a small region. 

Meanwhile, Milliron (1961) produced a new classification of Bombus which he 
divided into three genera, two each with two subgenera. He recognized no other 

ENTOM. 22, 5. 10 



212 O. W. RICHARDS 

subgenera though he mentions a number of " species groups ". I find Milliron's 
system unsatisfactory partly for fundamental reasons, partly because there are a 
number of errors in detail ; some of these may be misprints but others are in- 
comprehensible . 

The key to the genera and subgenera (pp. 54-55) depends largely on venational 
characters, which prove to be quite unworkable in a number of species on which 
they have been tested. Moreover, two of his genera, Pyrobombus von Dalla Torre 
and Megabombus von Dalla Torre, each fall in two halves of his key, so that their 
reintegration is essentially arbitrary. A number of exceptions have to be made 
for particular species suggesting that perhaps his genera would really be better 
split. Other points, more of detail, in Milliron's system which are very difficult 
to defend include, (i) putting Obertobombus Reinig in Pyrobombus sgen. Pyrobombus 
von Dalla Torre while Sibiricobombus Vogt (of which it is a synonym) is put in 
Pyrobombus sgen. Cullumanobombus Vogt, (2) putting Thoracobombus von Dalla 
Torre in Pyrobombus and not near its close allies in Megabombus von Dalla Torre, 
(3) separating B. dentatus Handlirsch and its subspecies orichalceus Friese from 
Alpigenobombus Skorikov (to which they clearly belong) and placing them in Bombias 
Robertson, with which they have little in common and further in putting with them 
B. haueri Handlirsch, which is really a member of Crotchiibombus Franklin which 
he places in Pyrobombus von Dalla Torre, (4) placing Kallobombus von Dalla Torre, 
Alpinobombus Skorikov and Orientalibombus Richards in Megabombus von Dalla 
Torre all the other groups of which have spinose mid basitarsi in the female. 

Medler (i962a, 1962^ has made a number of studies of the wings and mouthparts 
of humblebees, measuring in particular the length of the marginal cell, the length 
of the glossa and the ratio of the length of the first segment of the labial palp to 
the length of the prementum (= labial index). He suggests (iQ62a : 217) that the 
group of species allied to B. pratorum (Linnaeus) (subgenus Pyrobombus von Dalla 
Torre) is an unnatural one because of discrepancies in the value of these indices. 
While it has long been known that the very large subgenus Pyrobombus includes 
species with a considerable range of structure I cannot agree that the group is 
unnatural. The subgenera are really defined on a number of characters of both 
sexes though the male genitalia are often the simplest means of reaching a quick 
identification. Indices are often very valuable for distinguishing species but must 
be used with care as indicators of relationships. The palps and mouthparts are 
almost certainly liable to rapid adaptation and may well be imperfect indicators 
of phylogenetic relationships. 

The classification of Bombus has never been very closely set out but most specialists 
in the taxonomy of the genus are reasonably satisfied with something approaching 
the schemes of Vogt & Skorikov. The scheme set out below is far from original and 
cannot be regarded as final. Although the author is responsible for any errors, he 
is very grateful for much help and advice from Drs J. F. Perkins, B. Tkalcu and I. 
H. H. Yarrow. 

Alphabetical list of names available for the subgenera of Bombus. 

Numbers are attached to the valid groups or to groups later described, and show 
the order in which they are dealt with in the descriptions. 



SUBGENERIC DIVISIONS OF BOMBUS 213 

30. ADVENT 'ORIBOMBUS Skorikov, 1922 : 150. 

Proposed as subgenus of Agrobombus Vogt. Type-species Agrabombus [sic] adventor 
Skorikov, 1914 = Bombus adventor (Skorikov, 1914), by designation of Sandhouse, 1943 : 522. 

(P- 254)- 

AGROBOMBUS Vogt, 1911 : 52. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus agrorum (Fabricius) 
= Apis agrorum Fabricius, 1787 nee Schrank, 1781 [ = Apis [Bombus] pascuorum (Scopoli, 
I 736)], by designation of Sandhouse, 1943 : 523. 

Synonym of Thoracobombus von Dalla Torre. Syn. n. 
AGRIBOMBUS Skorikov, 1938 : 145. [Emend.] 

i. ALPIGENOBOMBUS Skorikov, 1914 : 128. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus lefebvrei Lepeletier, 
1836. [ = B. mastrucatus Gerstacker, 1869], by original designation, (p. 221). 

ALPIGENIBOMBUS Skorikov, 1938 : 122. [Emend.] 

18. ALPINOBOMBUS Skorikov, 1914 : 122. 

Proposed as a subgenus of Bombus Latreille. Type-species Bremus alpinus (Linnaeus) 
= Bombus alpinus (Linnaeus) = Apis alpinus Linnaeus, 1758, by designation of Frison, 
1927 : 66. (p. 239). 

ALPINIBOMBUS Skorikov, 1937 : 53- [Emend.] 
ANODONTOMBUSKruger, 1917 : 61,65. 

Proposed as a sectional name but according to Milliron, 1961 : 53 it is synonymous with 
Pyrobombus von Dalla Torre, 1880. 

ATROCINTOB. [OMBUS] Skorikov, 19336 : 244, Table i. 

Not described and no type-species designated, but two species, Bombus atrocinctus Smith, 
1872 and Bombus terminalis Smith, 1872, were included. This must be treated as a nomen 
nudum. 

BOMBELLUS, Zool. Record, Insecta, 1931 : 248, nomen nudum. 
Not in Wild, 1931, Scott. Nat. 190 : 118, as claimed. 

9. BOMBIAS Robertson, 1903 : 176. 

Proposed as a genus. Type-species Bombias auricomus Robertson, 1903 [ = B. nevadensis 
Cresson, 1874 ssp.], by original designation (p. 230). 

5. BOMBUS Latrielle, i8o2a : 385, i8o2b : 437. 

Proposed as a genus. Type-species Apis terrestris Linnaeus, 1758, monobasic, (p. 266) 

BOOPOBOMBUS Frison, 1927 : 62. 
Sectional name. 

BREMUS J urine, 1801 : 164. 

Proposed as a genus. Type-species Bremus terrestris (Linnaeus) = Bombus terrestris 
(Linnaeus) Apis terrestris Linnaeus, 1758 by designation of Morice & Durrant, 1915 : 428. 
Invalidated, Hemming, 1939. 
CALLOBOMBUS von Dalla Torre, see Kallobombus von Dalla Torre. 

CHROMOBOMBUS von Dalla Torre, 1880 : 40. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus muscorum (Linnaeus) 
= Apis muscorum Linnaeus, 1758, by designation of Sandhouse, 1943 : 538. Synonym of 
Thoracobombus von Dalla Torre. Syn. n. 



214 O. W. RICHARDS 

21. COCCINEOBOMBUS Skorikov, 1922 : 157. 

Proposed as a subgenus of Alpigenobombus Skorikov. Type-species Alpigenobombus 
coccineus (Friese) = Bombus coccineus Friese, 1903 : 254 by designation of Sandhouse, 
1943 : 539- (p- 243). 

7. CONFUSIBOMBUS Ball, 1914 : 78. 

Proposed as a subgenus of Bombus latreille. Type-species Bombus confusus Schenck, 
1859. Monobasic. 

CONFUSOBOMBUS Skorikov, 1922 : 149 [apparently independently proposed as a sub- 
genus of Bombus]. Bombus confusus Schenck, 1859 = Confusobombus confusus (Schenck) 
here designated as TYPE-SPECIES (p. 228). Synonym of Confusibombus Ball. 

ii. CROTCHIIBOMBUS Franklin, 1954 : 51. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus crotchii Cresson, 1878 
by original designation, (p. 231). 

13. CULLUMANOBOMBUS Vogt, 1911 : 57. 

Proposed as a subgenus of Bombus Latreille. Type-species Bremus cullumanus (Kirby) = 
Bombus cullumanus (Kirby) = Apis cullumana Kirby, 1802, by designation of Prison, 
1927 : 66. (p. 233) 

CULLUMANIBOMBUS Skorikov, 1938 : 145-146. [Emend.] 

24. DIVERSOBOMBUS Skorikov, 1914 : 406. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus diversus Smith, 1869 
by designation of Sandhouse, 1943 : 546. (p. 247). 

28. EVERSMANNIBOMBUS Skorikov, 1938 : 145. 

Proposed as subgenus of Agribombus = Agrobombus Vogt. Included only A. (E.) evers- 
manniellus (Skorikov) = Mucidobombus eversmanniellus Skorikov, 1922 = B. eversmanni 
Friese, 1911 nee B. modestus Eversmann ssp. eversmanni Skorikov, 1910, which is here desig- 
nated as the TYPE-SPECIES by monotypy and by virtual tautonymy. (p. 252). 

29. EXILOBOMBUS Skorikov, 1922 : 150. 

Proposed as a subgenus of Mucidobombus Skorikov. Type-species Mucidobombus exil 
[misprinted exiln. nov] Skorikov, 1922 = new name for Bombus nymphae Skorikov, 1909 $ 
only, not <J $. Monobasic, (p. 253). 

35. FERVIDOBOMBUS Skorikov, 1922 : 153. 

Proposed as a genus. Type-species Bremus fervidus (Fabricius) = Fervidobombus fervidus 
(Fabricius) Bombus fervidus (Fabricius) = Apis fervida Fabricius, 1798, by designation of 
Frison, 1927 : 69. (p. 259). 

6. FRATERNOBOMBUS Skorikov, 1922 : 156. 

Proposed as a subgenus of Alpigenobombus Skorikov. Type-species Alpigenobombus 
fraternus (Smith) = Bremus fraternus (Smith) = Bombus fraternus (Smith) = Apathus 
fraternus Smith, 1854 by designation of Frison, 1927 : 63. (p. 227). 

22. FUNEBRIBOMBUS Skorikov, 1922 : 157. 

Proposed as a subgenus of Alpigenobombus Skorikov. Type-species Alpigenobombus 
funebris (Smith) = Bombus funebris Smith, 1854. Monobasic, (p. 244). 



SUBGENERIC DIVISIONS OF BOMBUS 215 

HORTOMBUS Vogt, 1911 : 56. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus hortorum (Linnaeus) 
Apis hortorum Linnaeus, 1761 by designation of Sandhouse, 1943 : 559. 

Synonym of Megabombus von Dalla Torre. 
HORTIBOMBUS Skorikov, 1938 : 146. [Emend.] 

HYPNOROBOMBUS Quilis Perez, 1927 : 97. 

HYPNORUBOMBUS Quilis Perez, 1927 : 19. [Error.] 

Proposed as a subgenus of Bombus. Type-species Bombus hypnorum (Linnaeus) = Apis 
hypnorum Linnaeus, 1758. Monobasic. 
Synonym of Pyrobombus von Dalla Torre. 

4. KALLOBOMBUS von Dalla Torre, 1880 : 40. 

CALLOBOMBUS von Dalla Torre, 1896 : 503. Invalid [emendation]. 
Proposed as a subgenus of Bombus Latreille. Type-species Bombus soroeensis (Fabricius) 
= Apis soroeensis Fabricius, 1777 by designation of Sandhouse, 1943 : 561. (p. 225). 

KOZLOVIBOMBUS Skorikov, 1922 : 152. 

Proposed as a genus. Type-species Kozlovibombus kozlovi (Skorikov) = Bombus kozlovi 
Skorikov, 1909 = Bombus kohli Vogt, 1909 nee Cockerell, 1906 by designation of Sandhouse, 
1943 : 561. 

Synonym of Melanobombus von Dalla Torre. Syn. n. 

27. LAESOBOMBUS Skorikov, 1922 : 150. 

Proposed as a subgenus of Agrobombus Vogt. Type-species Agrobombus laesus (Morawitz) 
= Bombus laesus Morawitz, 1875. Monobasic, (p. 251). 

LAESIBOMBUS Skorikov, 1938 : 145. [Emend]. 

LAPIDARIOBOMBUS Vogt, 1911 : 58. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus lapidarius (Linnaeus) 
= Apis lapidaria Linnaeus, 1758 by designation of Sandhouse, 1943 : 562. 

Synonym of Melanobombus von Dalla Torre. 
LAPIDARIIBOMBUS Skorikov, 1938 : 145-146. [Emend.] 

LAPPONICOBOMBUS Quilis Perez, 1927 : 19, 22, 63. 

LAPONICOBOMBUS Neave, 1940 : 143. [Error.] 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus lapponicus (Fabricius) 
= Apis lapponica Fabricius, 1793 by designation of Milliron, 1961 : 58. 
Synonym of Pyrobombus von Dalla Torre. 

LEUCOBOMBUS von Dalla Torre, 1880 : 40. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus terrestris (Linnaeus) 
= Apis terrestris Linnaeus, 1758 by designation of Sandhouse, 1943 : 564. 

Synonym of Bombus Latreille s.s. 

MASTRUCATOBOMBUS Kriiger, 1917 : 66. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus mastrucatus Gerstaker, 
1869 [ = Bombus lefebvrei Lepeletier, 1836). Monobasic. 

Synonym of Alpigenobombus Skorikov. 

23. MEGABOMBUS von Dalla Torre, 1880 : 40. 

[MEGALOBOMBUS Schulz, 1906 : 267. Invalid emendation]. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus ligusticus Spinola, 
1805 = Apis argillacea Scopoli, 1763. Monobasic, (p. 246). 



2i6 O. W. RICHARDS 

15. MELANOBOMBUS von Dalla Torre, 1880 : 40. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus lapidarius (Linnaeus) 
= Apis lapidaria Linnaeus, 1758 by designation of Sandhouse, 1943 : 569. (p. 236). 

8. MENDACIBOMBUS Skorikov, 1914 : 125. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus mendax Gerstaker, 
1869 by designation of Sandhouse, 1943 : 572. (p. 229). 

32. MUCIDOBOMBUS Skorikov, 1922 : 149. 

Proposed as a genus. Type-species Bombus mucidus Gerstaker, 1869 by designation of 
Sandhouse, 1943 : 574. (p. 256). 

NEVADENSIBOMBUS Skorikov, 1922 : 149. 

Proposed as a genus. Type-species Bremus nevadensis (Cresson) = Bombus nevadensis 
Cresson, 1874 by designation of Prison, 1927 : 64. 

Synonym of Bombias Robertson, 1903. 

2. NOBILIBOMBUS Skorikov, i933a : 62. 

Proposed as a genus but no type-species was designated. Listed by Sandhouse (1943 : 
577) but without type-designation. Type-species cited as Bombus nobilis Skorikov, 1904 
by Milliron, 1961 : 54 but there is no such species. TYPE-SPECIES of the subgenus is here 
fixed as B. nobilis Friese (1904) 1905 by virtual tautonymy. The nature of this species is 
unfortunately controversial (see p. 222) but I believe this arrangement to be the best at the 
moment. 

OBERTOBOMBUS Reinig, 1930 : 107. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus oberti Morawitz, 1883. 
Monobasic. 

Synonym of Sibiricobombus Vogt. Syn. n. [but see Sandhouse, 1943 : 579]. 

ODONTOBOMBUS Kriiger, 1917 : 61, 65. 

Proposed as a sectional name but stated by Milliron, 1961 : 57 to be equivalent to Mega- 
bombus von Dalla Torre. 

3. ORIENTALIBOMBUS Richards, 1929 : 378. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus orientalis Smith, 1854 
by original designation, (p. 224). 

POECILOBOMBUS von Dalla Torre, 1882 : 23. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus sitkensis Nylander, 
1848 by designation of Sandhouse, 1943 : 589. Synonym of Pyrobombus von Dalla Torre. 

POMOBOMBUS Kriiger, 1917 : 65. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus pomorum (Panzer) = 
Bremus pomorum Panzer, 1804 by designation of Sandhouse, 1943 : 589. 

Synonym of Rhodobombus von Dalla Torre. 
POMIBOMB US Skorikov, 1938 : 145-146. [Emend.] 

PRATOBOMBUS Vogt, 1911 : 49. 

Proposed as a subgenus of Bombus Latreille. Type-species Bremus pratorum (Linnaeus) 
Bombus pratorum (Linnaeus) = Apis pratorum Linnaeus, 1761 by designation of Prison, 
1927 : 67. 

Synonym of Pyrobombus von Dalla Torre. 

PRATIBOMBUS Skorikov, 1937 : 59: ^S 8 : 146- [Emend.] 



SUBGENERIC DIVISIONS OF BOMBUS 217 

1 6. PRESSIBOMBUS Prison, 1935 : 342. 

Proposed as a subgenus of Bremus Panzer. Type-species, monobasic and by original 
designation, Bremus pressus Prison, 1935 = Bombus pressus (Prison), (p. 237). 

14. PYROBOMBUS von Dalla Torre, 1880 : 40. 

PYRRHOBOMBUS von Dalla Torre, 1882 : 28. [Invalid emendation.] 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus hypnorum (Linnaeus) 
= Apishypnorum Linnaeus, 1758. Monobasic, (p. 234). 

34. RHODOBOMBUS von Dalla Torre, 1880 : 40. 

Proposed as a subgenus of Bombus Latreille. Type-species Bremus pomorum Panzer, 1804 
= Bombus pomorum (Panzer) by designation of Sandhouse, 1943 : 596. (p. 258). 

19. ROBUSTOBOMBUS Skorikov, 1922 : 157. 

Proposed as a subgenus of Alpigenobombus Skorikov. Type-species Alpigenobombus 
robustus (Smith) = Bombus robustus Smith, 1854 by designation of Sandhouse, 1943 : 597. 
(p. 240). 

20. RUBICUNDOBOMBUS Skorikov, 1922 : 154. 

Proposed as a subgenus of Fervidobombus Skorikov. Type-species Fervidobombus rubi- 
cundus (Smith) Bombus rubicundus Smith, 1854 by designation of Sandhouse, 1943 : 597. 
(p. 241). 

17. RUFIPEDIBOMBUS Skorikov, 1922 : 156. 

RUFIPEDOBOMBUS Milliron, 1961 : 54. [Error]. 

Proposed as a genus. Type-species Rufipedibombus rufipes (Lepeletier) = Bombus rufipes 
Lepeletier, 1836. Monobasic, (p. 238). 

RUFOCINCTOBOMBUS Prison, 1927 : [78] pi. 17, fig. 9. 

Type-species Bremus rufocinctus (Cresson) = Bombus rufocinctus Cresson, 1863, monobasic. 
Presumably an error since on p. 65 Prison uses the name Cullumanobombus. 

Synonym of Cullumanobombus Vogt. 

25. SENEXIBOMBUS Prison, 1930 : 3. 

Proposed as a subgenus of Bremus Panzer. Type-species Bremus senex (Vollenhoven) = 
Bombus senex Vollenhoven, 1873 by original designation, (p. 248). 

10. SEPARATOBOMBUS Prison, 1927 : 64. 

Proposed as a subgenus of Bremus Panzer. Type-Species Bremus separatus (Cresson) = 
Bombus separatus Cresson, 1863 = Apis griseocollis Degeer, 1773 by original designation. 
(P- 231). 

12. SIBIRICOBOMB US Vogt, 1911 : 60. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus sibiricus (Fabricius) = 
Apis sibirica Fabricius, 1781 by designation of Sandhouse, 1943 : 599. (p. 232). 

SIBIRICIBOMBUS Skorikov, 1938 : 145-146. [Emend.] 
SOROEENSIBOMBUS Vogt, 1911 : 63. 

SORO&NSIBOMBUS Skorikov, 1923 : 151. [Error.] 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus soroeensis (Fabricius) 
= Apis soroeensis Fabricius, 1777. Monobasic. 
Synonym of Kallobombus von Dalla Torre. 



2i8 O. W. RICHARDS 

33. SUBTERRANEOBOMBUS Vogt, 1911 : 62. 

Proposed as a subgenus of Bombus Latreille. Type-species Bremus subterraneus (Linnaeus) 
Bombus subterraneus (Linnaeus) = Apis subterranea Linnaeus, 1758 by designation of 
Prison, 1927 : 68. (p. 257). 

SUBTERRANEIBOMBUS Skorikov, 1938 : 145-146. [Emend]. 
SULCOBOMBUS (Vogt) Kriiger, 1917 : 65. 

Proposed as a group name to include Mendacibombus and Confusobombus, but type-species 
designated by Sandhouse, 1943 : 502 as Bombus confusus Schenck, 1859. 

Synonym of Confusibombus Ball. 

TANGUTICOBOMBUS Pittioni, 19396 : 201. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus tanguticus Morawitz, 
1886. Monobasic and by original designation. 

Synonym of Melanobombus von Dalla Torre. Syn. n. 

TERRESTRIBOMBUS Vogt, 1911 : 55. 

Proposed as a subgenus of Bombus Latreille. Type-species Bremus terrestris (Linnaeus) 
= Bombus terrestris (Linnaeus) = Apis terrestris Linnaeus, 1758 by designation of Prison, 
1927 : 67. 

Synonym of Bombus Latreille. 

31. THORACOBOMBUS von Dalla Torre, 1880 : 40. 

Proposed as a subgenus of Bombus Latreille. Type-species Bombus sylvarum (Linnaeus) 
= Apis sylvarum Linnaeus, 1761 by designation of Sandhouse, 1943 : 604. (p. 255). 

26. TRICORNIBOMBUS Skorikov, 1922 : 1515. 

Proposed as a subgenus of Agrobombus Vogt. Type-species Agrobombus tricornis (Radosz- 
kowsky) = Bombus tricornis Radoszkowsky, 1888. Monobasic, (p. 249). 

UNCOBOMBUS Kriiger, 1919 : 65. 

Proposed as a group name to include Lapidariobombus, Pratobombus, Cullumanobombus and 
Mastrucatobombus . 

VOLUCELLOBOMBUS Skorikov, 1922 : 149. 

Proposed as a genus. Type-species Volucellobombus volucelloides (Gribodo) = Bombus 
volucelloides Gribodo, 1891. Monobasic. 

Synonym of Robustobombus Skorikov. Syn. n. The page priority of Volucellobombus is 
ignored because the male of B. volucelloides is unknown whereas both sexes are known in 
B. robustus Smith. 

NOTE ON GROUP-NAMES 

A few of the names in the above list seem certainly to have been intended for 
groups of subgenera. Where a type has been fixed they would be available as 
subgeneric names though as it happens none has priority. Where a type has not 
already been fixed I have not done so. The concepts indicated by these names 
(sometimes emended during the present study) seem to be as follows: 

Anodontobombus Kriiger = Subgenera 1-22 
Boopobombus Prison = Subgenera 6-9 
Odontobombus Kriiger = Subgenera 23-35 
Sulcobombus Kriiger = Subgenera 7-8 

Uncobombus Kriiger = Subgenera 10-14, possibly also 15-17. Originally sub- 
genera i and 13-15. 



SUBGENERIC DIVISIONS OF BOMB US 219 

CHARACTERS OF THE SUBGENERA OF BOMBUS 

The order in which the subgenera are arranged below will perhaps be found 
convenient but it is not intended to have a very deep phylogenetic significance. 
It is very difficult to form a theory of the evolution of Bombus. The suggestion of 
Milliron (1961 : 50) that B. mendax Gerstacker has Anthophorine affinities, whereas 
B. fraternus (Smith) arose from some Xylocopine stock seems to me fantastic. 
Euglossa and its allies are perhaps the non-social bees most similar to Bombus. 
If one regards short tongues and malar spaces as primitive, then the serrate mandible 
of Alpigenobombus which accompanies them is surely highly specialized; the habit 
of biting through the base of the corolla can scarcely be primitive. Nevertheless 
a long malar space is practically unknown in any other bee-genus (Michener, 1944 : 
205 otherwise only in Apis, some Trigona, Thrinchostoma, and some Colletes). 

On the grounds of nest-behaviour and brood-rearing, the section Odontobombus 
has been regarded as primitive (Sladen, 1912 : 43 ; Plath, 1934 : 119) but it is 
not certain that structural evolution has necessarily kept pace with evolution of 
social behaviour. The data recorded by Hobbs (1964) make it difficult to construct 
a combined phylogeny. Certain groups of subgenera seem clearly closer to one 
another structurally than to the others ; in the present arrangement such groups 
are: 1-2, 6-9, 13-16, 19-20, 23-25, 26-32, 34~35- 

There are certain characters of subgeneric importance which have not been 
previously fully described or which require a scheme of nomenclature. Some terms 
have already been suggested in Richards (1927), others are noted below. 

Measurements were made with a micrometer scale in the microscope eyepiece. 
Since in general only proportions are important the measurements are expressed 
in micrometer units which are equal to 0-073 mm. The following structures have 
been measured, $ length of malar space, proportions of antennal segments 3:4:5; 
cJ length of malar space, proportions of antennal segments 3:4:5:8, length x 
greatest width of mid and hind basitarsus. 

While the absolute length of the malar space is given, it is usually best to measure 
it against that of the third antennal segment. I have found it more satisfactory 
to measure its length as in Text-fig. I (cd) rather than obliquely (bd) as proposed 
by Kriiger (1920 : 316, fig. B). 

Frons. In the female, on each side of the frons, beside the lateral ocelli, are 
roundish, unpunctured areas (Text-fig. 2). Between these unpunctured areas and 
the eyes, lying along the inner orbit, is often a band of special sculpture. Kriiger 
(1951 : 144) first showed the importance of these areas and it is illustrated by 
Moure & Sakagami (1962 : 85, fig. i) in several species of Fervidobombus ; the sculp- 
ture is less specific in the males. In some subgenera a furrow, sometimes weak, 
transverses the top of the gena (temples) a little below the dorsal limit of the eyes. 
This will be termed the dorsal furrow of the gena. It is of minor subgeneric importance 
but in some groups it is a specific character of some value. It is best developed 
in Pyrobombus and absent in most Odontobombus. It is weaker in the males. 

The sting. Various authors have pointed out the significance of the inner thick- 
enings of the sting-sheath ( = incrustaciones vestibuli aculei) . The outer thickenings 
should also be examined, though less significant. The membrane between the 



220 O. W. RICHARDS 

inner thickenings (presumably in the region of the genital orifice) is often thrown 
into characteristic folds ; there are sometimes blackened patches in the membrane 
and, at least in Pyrobombus, these are not present in virgins and are scars produced 
during copulation by the male (probably by the sagittae) (Cumber, 1949 : 24). 
If the sting is fully extruded over the dorsal side of the abdomen, the sting-sheath 
forms a V of which the point is dorsal. In descriptions, the sting is described in 
this orientation and the essential parts are lettered in Text-fig. 3 (p. 223). 

The male genitalia. The nomenclature of the male genitalia in Hymenoptera 
has been the subject of much controversy and I have earlier indicated a number of 
the alternatives (Richards, 1956). In the present work it has proved most con- 
venient to use the old-fashioned nomenclature. The chief reason for this is that 
Snodgrass (1940 : 59-60) quoting Zander (1900 : fig. 5) gives evidence for believing 
that the lacinia and the squama are parts of one structure, the parameral spine. 
Nevertheless, it is essential for taxonomic purposes to distinguish them and to do 
this the old-fashioned terms seem the best ; once using part of the system it then 
seems best to use the same nomenclatorial scheme throughout (Text-fig. 4). 




FIG. i. Method of measuring length of malar space. 




FIG. 2. Unpunctured areas (X) of frons of Bombus terrestris (Linnaeus), 



SUBGENERIC DIVISIONS OF BOMBUS 221 

I have only illustrated stings and male sternites and genitalia which have not 
been illustrated before. The scale lines are equal to i-o mm. for stings and to 
2-5 mm. for the male sternites and genitalia. 

DESCRIPTIONS OF THE SUBGENERA 
i. ALPIGENOBOMBUS Skorikov, 1914 
Maslrucatobombus Kriiger, 1917 

$. Ocelli separated by three diameters from eyes, lying just in front of postocular line, 
median ocellus almost in a straight line with lateral ones. Frons moderately closely and 
strongly punctured, unpunctured areas large but ill-defined, with some large punctures on 
their outer part, a feebly developed band of fine punctures along the inner margin of eyes. 
Dorsal furrow of gena usually present. Antennal segments 3 : 4 : 5 as j\ : 4^ : 6J, segment 
3 about twice to nearly three times as long as broad. Clypeus very transverse, swollen, closely 
but not very coarsely punctured throughout, apical impression very feeble, with rather dense 
punctures. Labral tubercles flattened, sharply truncate at inner end, furrow wide and deep, 
wider than length of antennal segment 3, lamella wide almost straight. Mandibles strongly 
curved, spoon-shaped, apex with six marginal teeth, incisura deep, no sulcus obliquus. Malar 
space short, very transverse, hardly longer than antennal segment 5 (= 7-7 ), sometimes 
with some fine punctures. Mid basitarsus with posterior apical angle more or less distinctly 
acute. Hind tibia with corbicular surface bare, feebly reticulate, corbicular bristles normal, 
dorsal inner apical angle rather distinctly spinose, hind basitarsus with posterior dorsal angle 
obtusely spinose, surface not densely pubescent but in most species with some long bristles 
arising from lower edge and adjacent disk, especially proximally. Gastral tergite 6 reticulate 
or subrugose, sometimes with a feeble short furrow or very small posterior boss, short hairs 
not conspicuous. Sting with outer thickenings of sheath narrow and hardly widened above, 
inner thickenings a little widened below and gradually narrowed upwards, the degree of widen- 
ing varying, centre or sometimes more of wide part blackened and membrane blackened on 
a lateral lobe projecting down to about mid point. Wings hyaline or in eastern species blackish. 

o*. Eyes not swollen, ocelli separated by three diameters from eyes, lying almost on the 
postocular line. Antennae not very long, penultimate segments not curved, segments 3 14 : 5 :8 
= 7 : 4J : 6J : 6. Mandibles with a broad ventral and two small dorsal teeth, beard long 
and dense. Malar space transverse, shorter than antennal segment 5 (=6). Mid basitarsus 
moderately long and parallel-sided (40 x 9), posterior edge with quite numerous long bristles, 
anterior edge also with a few. Hind tibia considerably widened distally, moderately convex, 
shining with large bare distal disk, both edges with very long hairs. Hind basitarsus 40 x 12, 
dorsal apical angle subacute, dorsal edge with a few long bristles. Gastral sternite 6 scarcely 
modified, a little truncate. Gastral sternite 7 wide, widely produced, end somewhat truncate, 
no fenestrae, bristles short and inconspicuous. Gastral sternite 8 strongly produced, produced 
part with sides somewhat convergent, end obtusely angulated, one central line, bristles short 
and dense. Genitalia rather long and narrow, stipes with a deep, wide inner impression; 
sagittae narrow, much wider in side view, especially towards base which is followed by a ventral 
emargination forming a blunt tooth, apex bent down and hooked inwards, hook rather wide, 
tip rounded, outer edge with a narrow nearly straight flange, proximal end of flange projecting 
a little ; spatha with sides strongly converging, at mid-point two and a half to three times as 
wide as sagitta ; squama rounded quadrate or pointed inwards, proximal inner lobe narrow 
and separated from main lobe by a narrow emargination ; lacinia projecting well beyond 
squama, curved, parallel-sided, end emarginate with inner angle acute, with a small bristle- 
tuft ; volsella region with or without dense bristles. 

Illustrations. $ sting Hazeltine & Chandler 1964 : pi. 6. 6* Gastral sternite 7 and 8 
Pittioni, i939a : pi. 3, fig. 5 ; Prison, 1935 : figs. 5d, e, 6e, f. Genitalia Kriiger, 1920 : pi. 6, 
fig. q ; Pittioni, i93ga : pi. 3, fig. 5 ; Prison, 1935 : figs. 5b, 6c. 

N.W. SPAIN, PYRENEES, ALPS and NORWAY to CHINA and HIMALAYAS. 



222 O. W. RICHARDS 

2. NOBILIBOMBUS Skorikov, 1933 
(Text-figs. 3-6) 

Skorikov (i933a : 62) published a new subgenus, Nobilibombus, including three 
species, B. nobilis Friese, B. validus Friese, and the later described B. morawit- 
ziides Skorikov. No male was known. No type-species of the subgenus was fixed 
but under the International Code of Zoological Nomenclature, Art. 68 (c), B. nobilis 
Friese can be regarded as the type-species by virtual tautonymy. 

Friese's description ( (1904) 1905 : 513) mentions that the mandibles have four 
to five teeth and the species is recorded from several localities in W. China (8 $, 
6 $). On p. 510, B. validus is stated to have untoothed mandibles. Nevertheless, 
in the review of palearctic species (pp. 516-523), he places (p. 517) B. validus in 
a group with B. mastrucatus Gerstacker which he says has a short malar space and 
toothed mandibles. On the other hand, B. nobilis (p. 519) is placed in a group 
with B. lapidarius (Linnaeus) among the species in which the mandibles are un- 
toothed. Dr. B. Tkalcu has seen a " type $ " of B. nobilis from Friese's collec- 
tion and it is a species of subgenus Melanobombus von Dalla Torre [= B. Pyrosoma 
Morawitz, 1890 ssp. honei Bischoff, 1936]. It seems quite likely that Friese's 
series was mixed and that this led to the confusion already noted in the description. 
It seems difficult to believe that Skorikov could have erected the subgenus Nobili- 
bombus for species with toothed mandibles and with malar space longer than in 
Alpigenobombus, unless most of Friese's specimens (now at Leningrad) had these 
characters. Unfortunately it has not proved possible to examine the whole series 
or indeed any specimen other than the one retained by Friese. 

The characters of the subgenus given below are taken from B. sikkimi Friese 
and its differentiation from other subgenera will be seen in the keys (pp. 260, 265). 

$. Ocelli separated by about three diameters from eyes, lying well in front of the postocular 
line, median ocellus almost in line with the lateral ones. Frons with fine and indistinct though 
fairly numerous punctures, especially along inner margins of eyes, unpunctured areas small 
and ill-defined. Dorsal furrow of gena present but usually very weak. Antennal segments 
3:4:5 = loj : 5 : 7, segments 3 about three times as long as wide in centre. Clypeus about 
quadrate, rather swollen, with scattered sparse punctures of variable size, apical impressions 
moderate, with rather close, large punctures. Labral tubercles considerably raised, angulated 
at inner end, shining and coarsely punctured, furrow rather narrow, less wide than length of 
antennal segment 3, lamella nearly straight, almost as wide as a tubercle. Mandibles with 
six apical teeth, the ventral one at bottom of the incision large and directed downwards, almost 
at right angles to the others, no sulcus obliquus. Malar space quadrate, longer than antennal 
segment 3 (= 15). Mid basitarsus with posterior apical angle spinosely produced, whole surface 
with relatively long bristles and on basal half with very long, obliquely erect ones. Hind 
tibia with corbicular surface bare, very feebly reticulate, corbicular bristles normal, dorsal 
inner apical angle scarcely produced. Hind basitarsus posterior dorsal angle acute, surface 
rather densely pubescent, dorsal edge with moderately short dense bristles, ventral part of 
disk and lower edge with long bristles, especially on basal half, almost like those of corbicle. 
Gastral tergite 6 somewhat truncate. Characters of sting (Text-fig. 3) essentially similar to 
Alpigenobombus. Wings subhy aline. 

(J. Eyes not swollen, ocelli separated by three diameters from eyes, lying about on the 
postocular line. Antennae not very long, penultimate segments not curved, segments 3:4:5 
: 8 = 7i : 5i : 7 17. Mandibles with a distinct ventral and two small dorsal teeth, the most 



SUBGENERIC DIVISIONS OF BOMBUS 



223 



dorsal one not very distinct, beard long and dense. Malar space distinctly longer than broad, 
longer than antennal segments 2 + 3 but shorter than 3 + 4 (=12). Mid basitarsus long 
and parallel-sided (45 x n), margins especially posterior one with many long hairs. Hind 
tibia distally a little concave with a large bare disk, both margins and proximal half of disk 
with very long bristles, at least twice as lorg as its greatest width. Hind basitarsus 50 x 13, 
dorsal apical angle just acute, surface with dense, tomentum-like hairs and on disk many very 
long bristles especially on margins. Gastral sternite 6 with apical margin rounded and a little 
thickened with a dense short postapical fringe. Gastral sternite 7 (Text-fig. 4) very widely 
crescentic (i.e. long in longitudinal direction), no lines or fenestrae, one long bristle on each side. 
Gastral sternite 8 (Text-fig. 5b) strongly produced, sides feebly convergent, distal angles almost 
rounded, apex truncate or feebly curved, no fenestrae but distal half with two lines enclosing 
a very narrowly pyriform area, surface with dense short bristles. Genitalia (Text-fig. 6) large 
and rather long, stipes with a wide sharply defined impression, sagittae narrow, much wider 




FIGS. 3-6. Bombus sikkimi Friese. 3, sting : a, dorsal end of sting ; b, outer thickening 
of sheath, c ; d, inner thickening ; e, folds in membrane. 4, gastral sternite 7 of $ ; 
5, gastral sternite 8 of $ ; 6, <$ genitalia, the essential parts labelled. 



224 



O. W. RICHARDS 



in side view with a small ventral tooth at end of postbasal emargination, apex bent downwards 
and hooked inwards, apex of hook just blunt, outer side with a moderate flange which widens 
anteriorly but does not form a tooth ; spatha wide, sides strongly convergent, at centre 2-3 
times as wide as adjacent sagitta ; squama very transverse, inner side much wider than outer 
side, an inner projection forming a rounded-squarish lobe, extending some way beyond the 
stipes, and separated from this a narrow emargination a small proximal lobe ; lacinia extending 
some way beyond the squama, wide, feebly curved and parallel-sided, outer edge raised into 
a keel just inside the edge, apex feebly emarginate, outer corner rounded, inner one forming 
a feeble tooth ; volsellar region with dense, moderately long bristles. 

Illustrations. Sting Text-fig. 3. Gastral sternites 7 and 8 Text-figs 4 and 5. Genitalia 
Text-fig. 6. 

HIMALAYAS (s.l.) (W. INDIA to N. CHINA). 

3. ORIENT ALIBOMBUS Richards, 1929 

$. Ocelli separated by nearly three diameters from eyes, lying well in front of postocular 
line. Frons with a large unpunctured area around the ocelli, only a few fine punctures in 
front of each lateral ocellus and some rather fine punctures along inner margins of eyes. Dorsal 
furrow of gena sometimes feebly developed posteriorly. Antennal segments 3 : 4 : 5 = 10 : 
6 : i\, segment 3 about two and a half times as long as broad. Clypeus elongate, swollen, 
little punctured, apical impressions feeble, little punctured. Labral tubercles angularly con- 
vex but not specially so at inner angles, furrow wide, wider than length of antennal segments 
2 + 3, lamella wide, nearly straight. Mandibles with no incisura, salcus obliquus well-marked 





10 




11 




12 




FIGS. 7-12. Stings of 7, Bombus funerarius Smith ; 8, B. mendax Gerstacker ; 9, B. vorticosus 
Gerstacker ; 10, B. pressus (Frison) ; u, B. eximius Smith ; 12, B. robustus Smith. 



SUBGENERIC DIVISIONS OF BOMBUS 225 

(weaker in B. funerarius Smith). Malar space very long, longer than antennal segments 2 + 3 
(= 19). Mid basitarsus with posterior apical angle obtuse. Hind tibia with corbicular surface 
bare, surface not reticulate, corbicular bristles normal, dorsal inner apical angle rather obtusely 
produced. Hind basitarsus broad, slightly narrowed to the apex, dorsal apical angle just 
acute, surface with fine bristles not dense, marginal bristles short and dense. Gastral tergite 
6 sometimes with a feeble posterior furrow, its short hairs directed inwards. Sting with outer 
thickenings of sheath of moderate, even width, inner thickening moderately widened below 
and narrowing gradually above, lower part of inner edge blackened for a considerable distance 
and the adjacent membrane also blackened. Wings dark. 

<J. Eyes not swollen, ocelli separated by three diameters from eyes, lying just in front of 
postocular line. Antennae very long, penultimate segments each a little curved, segments 
3:4:5:8 = 7^: 8 : n : 13. Mandibles with a broad ventral and one small dorsal tooth, 
beard long and dense. Malar space much longer than broad, a little longer than antennal 
segments 2 + 3 but shorter than 3 + 4 (= 13^), or in -B. funerarius Smith a little longer than 
3 + 4 ( 14), not punctured. Mid basitarsus long and broad (48 x 13), fringes short. Hind 
tibia considerably widened and flattened distally, disk bare, fringes moderately dense but only 
a few bristles as long as maximal tibial width (fringes longer in B. funerarius Smith). Hind 
basitarsus 49 x 15, apical angle just acute, hairs of surface not dense, fringes short. Gastral 
sternite 6 with end thickened and recurved, especially on central quarter, followed by a mem- 
brane and a long, rather dense fringe, still longer at each side. Gastral sternite 7 forming a 
wide transverse band with anterior and posterior margins subparallel, latter feebly bisinuate, 
surface with sparse, scattered, short bristles. Gastral sternite 8 in the form of narrow band 
produced into a very wide tongue-shaped process, sides slightly rounded and angles just rounded, 
apex very feebly emarginate, surface densely covered with short bristles, three small fenestrae 
on mid-line (except in B. funerarius Smith). Genitalia very large and rather elongate, stipes 
broad with a wide, fairly well-defined impression ; sagittae narrow and very little wider in 
side view, with a strong recurved tooth near mid-point beneath, apex somewhat downcurved, 
ending in a long point, with a very slight flange on outer edge ; spatha narrow, hardly wider 
than sagitta ; squama large, transverse, somewhat wider on inner than outer-side, projecting 
in a little past the stipes, all angles rounded ; lacinia mostly narrow, parallel-sided, curving 
inwards, projecting well beyond squama, apex curved over inwards and truncate, dorsal 
corner acute, from base of lacinia and hidden by the squama arises a long, complex, pointed 
process ; or, in B. funerarius Smith, lacinia forms a long acute process and beneath the 
squama bears a small equilateral triangular process ; volsella region with dense, moderately 
long hairs. 

Illustrations. Sting Richards, 1929 : fig. 3 ; Text-fig. 7. Gastral sternites 7 and 8 
Prison, 1935 : n g s - 4-d. e, b, 7d, e ; Richards, 1929 : fig. i. Genitalia Richards, 1929 : fig. 
3 ; Prison, 1935 : fig. 70. 

HIMALAYAS, S.W. CHINA. Bombus funerarius Smith (^ = Bremus priscus Prison, 
J 935) is unusually distinct but probably best included. 

4. KALLOBOMBUS von Dalla Torre, 1880 
Soroeensibombus Vogt, 1911 

?. Ocelli separated by rather more than three diameters from eyes, lying almost behind 
the postocular line. Frons rather closely but not very strongly punctured, unpunctured areas 
small but well-defined, a wide band of fine punctures along inner margins of eyes. Dorsal 
furrow of gena just indicated. Antennal segments 3 : 4 : 5 = 7 : 5 : 5^, segment 3 about 
twice as long as broad. Clypeus rather short and swollen, rather strongly and evenly punctured, 
apical impressions strongly but not more closely punctured. Labral tubercles not much 
raised, moderately angled at inner end, furrow narrow, not as broad as length of antennal 
segment 3, lamella short and curved, inconspicuous. Mandibles with no incisura or salcus 

ENTOM. 22, 5. II 



226 O. W. RICHARDS 

obliquus. Malar space just transverse, a little longer than antennal segment 3 (=8). Mid 
basitarsus with posterior apical angle obtuse. Hind tibia with corbicular surface bare, surface 
moderately reticulate, dorsal inner apical angle not produced. Hind basitarsus with dorsal 
angle moderately acute, surface not densely haired, bristles rather short. Gastral tergite 6 
feebly sculptured, rather shining. Sting with outer thickening of sheath moderately wide, 
especially above, inner thickenings widening rather strongly upwards to above middle, then 
suddenly narrowed, wide part a little blackened at its edge, no black spots in membrane but 
a slight dorsal fold. Wings evenly but not strongly infuscate. 

(J. Eyes not swollen, ocelli separated by fully three diameters from eyes, lying just behind 
the postocular line. Antennae, long, penultimate segments each a little curved, segments 
3 : 4 : 5 : 8 = 4J : 5j : 6J : 7. Mandibles with a broad ventral and small dorsal tooth, beard 
long and dense. Malar space quadrate, longer than antennal segments 2 + 3, shorter than 
3 + 4 (= 7), not punctured. Mid basitarsus not very long, nearly parallel-sided but a little 
narrowed at each end (37 x 8), fringes short. Hind tibia moderately convex, with a consider- 
able shining, nearly bare disk, fringes long, especially the dorsal one. Hind basitarsus 34 x 10, 
considerably narrowed at base, apical angle just acute, fringes short. Gastral sternite 6 a 
little produced on central quarter, not recurved, scarcely thickened, with a short postapical 
fringe. Gastral sternite 7 strongly produced, sides of produced part with a large submem- 
branous, transparent area, apex shallowly emarginate, with a narrow band of dense bristles 
of which some on each side are longer. Gastral sternite 8 produced, production relatively 
narrow and parallel-sided, angles rounded, apex scarcely emarginate, a considerable area of 
dense short bristles, no fenestra but two parallel lines meeting before apex. Genitalia markedly 
elongate, stipes with no real inner impression ; sagittae narrow, bent down at apex, hooked 
outwards, hook forming a sort of half arrowhead, tip rounded (proteus Gerstacker) or pointed 
(soroeensis (Fabricius)), tip of hook pointed, sagitta also narrow in side view but with a large 
isolated pointed tooth beneath centre ; spatha wide, sides strongly converging, at centre three 
times as wide as sagitta ; squama elongate, almost twice as long as broad, inner edge with a 
shallow emargination, all corners rounded ; lacinia long, curved inwards, projecting well 
beyond squama, apex emarginate, outer corner sharp, inner one forming a curved, feebly 
serrate hook, between them a small tuft of bristles ; volsella regions with dense short hairs. 

Illustrations. Sting Richards, 1927 : fig. 54 ; Hazeltine & Chandler, 1964 : pi. 6. 
Gastral sternites 7 and 8 Radoszkowsky, 1884 : fig. 34C ; Pittioni, I939a : pi. 2, fig. 6. 
Genitalia Kriiger, 1920 : pi. 7, fig. u ; Richards, 1927 : fig. 54 ; Pittioni, i93ga : pi. 2, fig. 6. 

EUROPE and adjacent parts of ASIA. 

5. BOMBUS Latreille, i8o2a (s.s.) 

Leucobombus von Dalla Torre, 1880 
Terrestribombus Vogt, 1911 

$. Ocelli separated by nearly three diameters from eyes, lying just in front of postocular 
line. Frons with numerous punctures, mostly fine and coarse mixed, with well-defined un- 
punctured areas and a band of close, mostly fine punctures along inner margins of eyes. Dorsal 
furrow of gena absent or very weak ; a little stronger in B. ignitus Smith. Antennal segments 
3:4:5 = 9:6:7, segment 3 twice as long as broad. Clypeus short, convex, closely and 
rather strongly punctured throughout and especially closely in the well-marked apical impres- 
sions. Labral tubercles flattened, angled at inner end, furrow narrow, less wide than length 
of antennal segments, lamella short, somewhat curved. Mandibles with a strong incisura 
and moderately distinct sulcus obliquus. Malar space transverse, shorter than antennal seg- 
ments 2 + 3 (=9). Mid basitarsus with posterior apical angle obtuse. Hind tibia with 
corbicular surface bare, very weakly reticulate, dorsal inner apical angle distinctly produced. 
Hind basitarsus with dorsal angle just acute, surface densely pubescent, bristles short. Gastral 
tergite 6 feebly sculptured, rather shining. Sting with outer thickenings of sheath narrow but 



SUBGENERIC DIVISIONS OF BOMBUS 227 

broadened dorsally, inner thickenings rather wide and twice emarginate, membrane between 
thrown into central and lateral folds. Wings moderately or in some species strongly infuscate. 

<J. Eyes not swollen, ocelli separated by three diameters from eyes, lying on the postocular 
line. Antennae of moderate length, segments 3:4:5:8 = 7:5:6: 6|. Mandibles with 
a broad ventral and small dorsal tooth, beard long and dense. Malar space almost transverse, 
longer than antennal segment 3 but shorter than 2 + 3 (= 9), with some fine punctures. Mid 
basitarsus relatively broad and parallel-sided (46 x 12), fringes short. Hind tibia slightly 
convex but somewhat hollowed out towards dorsal margin, bare after proximal quarter, fringes, 
especially the dorsal one, long. Hind basitarsus 45 x 15, distal angle just acute, fringes short. 
Gastral sternite 6 with margin considerably recurved and thickened and with a dense short 
postapical fringe. Gastral sternite 7 very transverse, posterior margin rounded with a central 
emargination, a group of 3-5 long bristles on each side, no lines or fenestrae. Gastral sternite 
8 broad, produced, trapeziform, sides strongly converging, apex weakly emarginate with corners 
angled, tufts of long bristles arising near these angles, no lines or fenestrae. Genitalia with 
very slight inner impressions on stipes ; sagittae narrow in dorsal view but undulating and 
widely flaring at apex, in side view very wide, deeply emarginate beneath just after base, apex 
widened and somewhat curved outwards, spatha wide, at least four times as wide as dorsal 
width of sagitta ; squama very transverse with an anterior submembranous inner projection 
separated from the main lobe by a deep emargination, main lobe expanded inwards in a more or 
less trumpet-shaped projection ; lacinia projecting very little beyond squama, end part transverse, 
strongly produced inwards into a process ending in a slight upturned hook, hind margin some- 
what roughened and with dense pubescence ; volsella region with short, rather sparse hairs. 

Illustrations. Sting Franklin, 1913 : figs. 149, 169, 176 ; Richards, 1927 : fig. 55 ; Tkalcu, 
1963 : figs. 13-15 ; Hazeltine & Chandler, 1964 : pi. 3. Gastral sternites 7-8 Franklin, 
1913 : figs, in, 113, 128, 135 ; Richards, 1927 : figs. 21, 23 ; Pittioni, i93ga : pi. 3, fig. i. 
Genitalia Franklin, 1913 : figs. 119, 120 ; Richards, 1927 : figs. 15-27 ; Kriiger, 1920 : 
pi. vii t ; Pittioni, i93ga : pi. 3, fig. i. 

HOLARCTIC and parts of S.E. and S.W. ASIA. 

6. FRATERNOBOMBUS Skorikov, 1922 

$. Ocelli separated by two diameters from eyes, lying well in front of postocular line. Frons 
shining, mostly with rather fine not very close punctures, fine and very close in an area behind 
the ocelli and coarser and less close in front of them, unpunctured areas large and ill-defined, 
no distinct band of denser fine punctures along inner margin of eyes. Dorsal furrow of gena 
absent. Antennal segments 3 : 4 : 5 = 10 : 5 : 6J, segment 3 twice as long as broad. Clypeus 
short, transverse, swollen but rather flattened on ventral third, with large but rather sparse 
and shallow punctures except on disk below, punctures close and deeper in a large region sur- 
rounding the apical impressions which are themselves feeble and ill-defined. Labrum with 
strongly convex tubercles forming smooth rounded knobs at inner end, furrow deep, a little 
wider than length of antennal segment 3, lamella strongly curved, half as wide as labrum. 
Mandibles with no incisura and a strong sulcus obliquus. Malar space very transverse, about 
as long as antennal segment 5 (=7). Mid basitarsus with posterior apical angle rounded. 
Hind tibia with corbicular surface bare, not reticulate, dorsal fringe unusually short and dense, 
mostly shorter than half apical width of tibia, dorsal inner apical angle long and acute. Gastral 
tergite 6 granulate with a subapical somewhat flattened shining triangle. Sting with outer 
thickenings of the sheath narrow and of even width, inner thickenings considerably widened 
and blackened on lower third, then very narrow, membrane not differentiated. Wings evenly 
dark fuscous. 

cJ. Eyes considerably swollen and reaching nearer to occipital margin than usual, ocelli 
separated by less than \ diameter from eyes, very far in front of postocular line. Antennae 
rather long, segments 3:4:5:8 = 9:5:7:8. Mandibles with one broad ventral and one 
smaller dorsal tooth, beard feeble, very short. Malar space absolutely linear, shorter than 



228 O. W. RICHARDS 

antennal segment 2 (= 2^). Mid tibia wide, flattened, longitudinally hollowed out beneath. 
Mid basitarsus long, parallel-sided (66 x 15), posterior fringe short and dense. Hind tibia 
convex, mostly densely hairy but with a more posterior streak which is shining, with sparse 
short hairs, both fringes very short. Hind basitarsus 65 x 17, long, narrow, parallel-sided, 
distal angle a right angle, fringes short. Gastral sternite 6 flattened with a feeble preapical 
transverse thickening. Gastral sternite 7 very large, trapeziform, almost quadrate, apex with a 
deep semicircular emargination covering more than half its width, a wide band of very short 
dense bristles on each side, no lines or fenestrae. Gastral sternite 8 large, strongly produced, 
apex with recurved corners and margin weakly trisinuate, a moderate patch of short bristles 
on each side, two parallel lines on disk posteriorly. Genitalia large and elongate, inner im- 
pressions of stipes distinct but rather narrow ; sagittae narrow and not much wider in side 
view, no tooth beneath, apex hooked inwards and outer side of hook considerably expanded 
by a flange which is hardly perceptibly serrate ; spatha narrow, hardly wider than sagitta 
except right at base ; squama rather long, outer and posterior margins forming a continuous 
curve, inner margin with a strong emargination forming a rather blunt projecting hook at 
anterior inner end ; lacinia elongate, projecting some way beyond squama, pointed, outer edge 
raised and thickened, inner edge flattened, thin and approximately straight, irregularly crenate, 
short haired. Volsella region bare. 

Illustrations. Sting Hazeltine & Chandler, 1964 : pi. 2. Gastral sternites 7 and 8 
Franklin, 1913 : figs, i, 22. Genitalia Franklin, 1913 : figs. 71, 87. 

NORTH and CENTRAL AMERICA. 

7. CONFUSIBOMBUS Ball, 1914 

Sulcobombus Kriiger, 1917 
Confusobombus Kriiger, 1917 

$. Ocelli separated by about two and a half diameters from eyes, lying well in front of 
postocular line. Frons mostly closely and rather finely punctured, unpunctured areas well- 
defined, a wide band of close fine punctures along inner margin of eyes. Dorsal furrow of 
gena absent. Antennal segments 3:4:5 = 11:5^: 5^, segment 3 between three and four 
times as long as broad. Clypeus moderately elongate, strongly swollen but with a wide flattened 
disk, closely and finely punctured, even more closely on lower third, apical impressions ill- 
defined but closely punctured. Labrum with tubercles little raised, much rounded, furrow 
wide and shallow though not quite so wide as length of antennal segment 3, lamella small, 
inconspicuous, curved. Mandibles with no incisura, sulcus obliquus moderately strong. Malar 
space a little longer than broad (= n), a little longer than antennal segment 3. Mid basitarsus 
with posterior apical angle rounded. Hind tibia with corbicular surface bare, moderately 
reticulate, inner dorsal apical angle short but pointed. Gastral tergite 6 moderately shining, 
finely granulate. Sting with outer thickenings of the sheath rather narrow but a little broader 
above than below, inner thickenings very narrow, intervening membrane convex above and 
sending to centre a large blackened lobe on each side. Wings evenly but not strongly infuscate. 

<$. Eyes swollen, temples narrow, ocelli separated by rather more than one diameter from 
eye, far in front of postocular line. Antennae short, segments 3:4:5:8 = 7: 3^ : 4 : 6. 
Mandibles with a broad ventral and a small dorsal tooth, beard moderately long and dense. 
Malar space strongly transverse, shorter than antennal segment 3 (= 5^), almost unpunctured. 
Mid basitarsus moderately long and parallel-sided (=42 x 10) posterior fringe about as wide 
as tarsus. Hind tibia thick and rather clavate, convex, surface punctured and cross-striate 
with short hairs on its whole surface, fringes long dorsally, mostly short ventrally. Hind 
basitarsus 37 x 12, moderately broad and narrowed to base, apical dorsal angle rounded, 
dorsal fringe long, ventral one short. Gastral sternite 6 strongly recurved, a little thickened 
and emarginate. Gastral sternite 7 crescentic but considerably produced into a rounded 
triangle, with a band of scattered short bristles well before apex, no lines or fenestrae. Gastral 
sternite 8 generally crescentic, little produced, feebly truncate, angles rounded, one long black 



SUBGENERIC DIVISIONS OF BOMBUS 229 

bristle only from a black spot on each side, no lines, two minute oval fenestrae on disk. Geni- 
talia small, pyriform ; stipes very broad with deep sharp-edged impressions covering less than 
half their width ; sagittae in dorsal view moderately narrow, dorsal half pale on apical half, 
at mid point with a lateral, somewhat ventral, tooth, beyond this finger-shaped, hardly nar- 
rowed, feebly serrate externally, considerably bent down, in side view at least twice as wide 
as in dorsal view, somewhat narrowing distally ; spatha very wide, at centre five times as 
wide as sagitta, but strongly narrowing distally ; squama elongate, bent into a V-shaped plate, 
the spine of the V forming the outer edge, the inner edge having dorsal and ventral plates, spine 
subacute distally, lower plate with posterior and inner edge rounded together, upper plate 
with a feeble bilobed process adjacent to stipes ; lacinia long, pointed, inner edge opposite 
end of squama produced into a stout, backward-curving tooth, outer edge with short stout 
dense bristles, inner edge with similar but less dense and stout bristles ; volsella region with 
a moderate number of not very long and stout bristles. 

Illustrations. Sting Hazeltine & Chandler, 1964 : pi. 5. Gastral sternites 7 and 8 
Radoszkowsky, 1884 : 42c ; Pittioni, iQ39a : pi. 4, fig. 2. Genitalia Kriiger, 1920 : pi. 
7, fig. r ; Pittioni, igsga : pi. 4, fig. 2. 

EUROPE. 

8. MENDACIBOMBUS Skorikov, 1914 

$. Ocelli separated by two and a half diameters from eyes and lying just in front of post- 
ocular line. Frons finely and rather closely punctured, unpunctured areas well-defined, of 
moderate size, a band of fine punctures along inner margin of eyes. Dorsal furrow of gena 
sometimes feebly indicated. Antennal segments 3:4:5 = 11:4:5, segment 3 nearly four 
times as long as broad. Clypeus very elongate, swollen, feebly punctured, apical impres- 
sions feeble and little punctured. Labrum with tubercles little raised, smooth and rounded, 
gradually narrowed and depressed inwards, furrow very wide and ill-defined, perhaps better 
described as a transverse furrow both above and below the little raised tubercles, lamella wide, 
straight. Mandibles with no incisura or sulcus obliquus. Malar space very elongate (= 14), 
longer than antennal segments 2 + 3. Mid basitarsus with posterior apical angle rounded. 
Hind tibia with basal half of corbicular surface covered with bristles, bare part densely reticulate, 
dull, inner dorsal apical angle not produced, hind basitarsus with long bristles on dorsal edge. 
Gastral tergite 6 shining, moderately closely and strongly punctured. Sting with outer thicken- 
ings of sheath very narrow, inner thickening also very narrow, not blackened, membrane 
undifferentiated. Wings evenly but very little infuscate. 

c. Eyes considerably swollen and reaching back to near the occipital margin, ocelli separated 
by one diameter from eyes, far in front of postocular line. Antennae rather long, segments 
3:4:5:8=1 2^:4^:6: 6J, segment 3 at least four times as long as broad. Mandibles 
rather wider than usual, ending in a single wide tooth only in B. mendax Gerstacker but with 
a smaller, more dorsal tooth in some Asiatic species, beard long and dense. Malar space very 
elongate, just shorter than antennal segment 3 (= u), practically unpunctured. Mid basi- 
tarsus moderately long and broad (=41 X 10), fringes mostly short but a few long bristles 
near base. Hind tibia slightly convex, rather dull, whole surface covered with not very close 
short feathered bristles, fringes very long. Hind basitarsus 34 x 15, rather short and broad, 
dorsal margin with many long hairs, apical dorsal angle rounded-acute. Gastral sternite 6 a 
little truncate with dense short pale pubescence along apex. Gastral sternite 7 crescentic 
with a wide produced piece which is about two and a half times as wide at apex as long, sides 
considerably converging, margin widely membranous, whole surface with scattered, mostly 
short bristles, no lines or fenestrae. Gastral sternite 8 produced in centre into a large sub- 
quadrate piece of which the sides are feebly rounded, apex rather deeply emarginate, surface 
with dense short bristles, no lines or fenestrae. Genitalia with stipes short and stout with 
very slight inner impression ; sagittae rather narrow on proximal half, still more so distally 
where eventually pointed, in side view bisinuate and very little wider, margin weakly crenate 
a little before apex, apical half on inner side with distinct hairs ; spatha moderately wide, 



230 O. W. RICHARDS 

rather more than twice as wide. as basal part of sagitta ; squama stout and dark, not produced 
inwards, in half side view quadrate with rounded angles, a little longer than broad ; lacinia 
elongate, projecting some way beyond squama, rounded-pointed with inner edge more truncate 
or a little concave, with stout, curved, rather short bristles on its outer edge ; volsella region 
bare. 

Illustrations. Sting Text-fig. 8. Gastral sternites 7 and 8 Pittioni, i939a : pi. 4, fig. 3. 
Genitalia Kriiger, 1920 : pi. 7, fig. s ; Pittioni, 1939 : pi. 4, fig. 3. 

High mountains of EUROPE and ASIA. 

9. BOMBIAS Robertson, 1903 
Nevadensibombus Skorikov, 1922 

$. Ocelli separated by rather more than two diameters from eyes and lying far in front 
of postocular line. Frons moderately finely and closely punctured, unpunctured areas large 
and ill-defined, a band of fine punctures along inner margin of eyes. Dorsal furrow of gena 
absent. Antennal segments 3:4:5 = 11:5:6, segment 3 three times as long as broad 
or rather less. Clypeus elongate, strongly swollen, especially dorsally, finely and evenly punc- 
tured, apical impressions feeble and scarcely more punctured. Labrum with tubercles wide 
and much flattened, furrow ill-defined and shallow but essentially narrow, lamella very wide 
with a shining thickened edge, surmounted by dense pubescence. Mandibles with no incisura 
and a feeble sulcus obliquus. Malar space about quadrate (= 12-15), a little longer than antennal 
segment 3. Mid basitarsus with posterior apical angle rounded. Hind tibia with corbicular 
surface bare, weakly reticulate, inner dorsal apical angle feebly produced, hind basitarsus with 
only short pubescence. Gastral tergite 6 finely granulate, rather dull. Sting with outer 
thickenings of sheath a little wider above than below, inner thickenings rather narrow, of almost 
even width throughout, edge recurved especially below, membrane undifferentiated except 
for some lateral blackening below. Wings infuscated. 

<. Rather large, eyes strongly swollen, temples narrow, ocelli separated by one quarter 
their diameter from eye, very far in front of postocular line. Antennae rather long, segments 
3:4:5: 8 = 8^:4^:5:5, penultimate segments not curved. Mandibles with a broad 
ventral and a small dorsal tooth, beard very sparse. Malar space very transverse (=5), about 
as long as antennal segment 5, with a few microscopic punctures. Mid basitarsus long and 
broad (48 x 13), posterior apical angle rounded, dorsal fringe long. Hind tibia considerably 
clavate, somewhat concave, moderately shining but strongly cross-striate, whole surface with 
moderately numerous, very short hairs, fringes short. Hind basitarsus (47 x 17), dorsal 
fringe long, apical dorsal angle rounded. Gastral sternite 6 somewhat depressed with apex 
recurved, with a dense short fringe. Gastral sternite 7 broadly produced, sides anteriorly 
diverging, apex truncate, feebly emarginate, angles moderately distinct, a distal patch of very 
short bristles, no lines or fenestrae. Gastral sternite 8 broadly triangular (no tongue-like 
projection), apex deeply emarginate and with a U-shaped line defining a still deeper mem- 
branous area which is not actually emarginate, two apical patches of short bristles, no other 
lines or fenestrae. Genitalia with stipes broad at apex, inner impressions hardly developed ; 
sagittae in dorsal view narrow, a little downcurved, posterior third outwardly feebly serrate, 
apex more or less pointed (no hooks or flanges), in side view hardly wider, no ventral tooth or 
emargination ; spatha very wide, five times as wide as sagitta, only narrowing just before 
its end ; squama large, subquadrate, sloping obliquely downwards laterally, all angles rounded ; 
lacinia relatively narrow, projecting far beyond the squama, apex subtruncate with a slight 
hook at inner end, whole surface and outer and inner margin with short, rather dense hairs ; 
no long bristles in volsellar regions. 

Illustrations. Sting Hazeltine & Chandler, 1964 : pi. 2 ; Franklin, 1913 : fig. 151. 
Gastral sternites 7 and 8 Franklin, 1913 : figs. 100, 124, 136, 139. Genitalia Franklin, 
1913 : figs. 73, 82, 83. 

N. AMERICA. 



SUBGENERIC DIVISIONS OF BOMBUS 231 

10. SEPAR A TOBOMBUS Prison, 1927 

$. Ocelli separated by two diameters or rather less from eyes and lying well in front of 
postocular line. Frons rather strongly and closely punctured, unpunctured areas small and 
well-defined, a band of fine punctures along inner margin of eyes. Dorsal furrow of gena 
absent. Antennal segments 3:4:5 = 9:4^: 5^, segment 3 twice as long as broad. Clypeus 
rather long and swollen, finely and moderately closely punctured, apical impressions feeble, 
more closely punctured. Labrum with tubercles, flattened, angled at inner end, furrow wide 
and deep, nearly as wide as length of antennal segment 3, lamella wide, curved. Mandibles 
with no incisura but with a strong sulcus obliquus. Malar space short and transverse (=8), 
shorter than antennal segment 3. Mid basitarsus with posterior apical angle rounded. Hind 
tibia with corbicular surface bare, distinctly reticulate, inner dorsal apical angle distinctly 
but not strongly produced, hind basitarsus with only short pubescence. Gastral tergite 6 
shining, feebly granulate. Sting with outer thickenings of sheath moderately wide both above 
and below, inner thickenings rather strongly widened on lower third, then narrowed in a smooth 
curve and thence upwards of an even width, a little blackened where it narrows, membrane 
projecting above and produced downwards on each side into an elongate-oval lobe which is 
blackened at the end. Wings infuscate at tip or throughout. 

<J. Rather large, eyes swollen, temples rather narrow, ocelli separated by two thirds their 
diameter from eyes, far in front of the postocular line. Antennae rather long, segments 
3 :4 : 5 : 8 = 7^ : 5^ : 7 :y, penultimate segments scarcely curved. Mandibles with a 
broad ventral and small dorsal tooth, beard long and dense. Malar space very short and 
transverse (= 3^), hardly longer than antennal segment 2, shorter than 4, hardly punctured. 
Mid basitarsus long and parallel-sided (45 x 9^), posterior apical angle rounded, fringes short. 
Hind tibia not very convex, dull and alutaceous, large disk almost completely bare, dorsal 
fringe long. Hind basitarsus (42 x 10) dorsal fringe long, fully twice as long as tibial width, 
apical dorsal angle a rounded right angle. Gastral sternite 6 with apex a little thickened, 
not recurved, with short dense postapical fringe. Gastral sternite 7 trapeziform, apex very 
feebly emarginate, corners rounded but slightly projecting, a large area of short dense bristles, 
no lines or fenestrae. Gastral sternite 8 subtriangularly produced, sides strongly converging, 
apex a little rounded, almost pointed, a large area of dense, short bristles, with two parallel 
lines which meet at centre and then proceed back as one line, no fenestrae. Genitalia rather 
broad, stipes with wide but very poorly defined impressions ; sagittae in dorsal view narrow, 
apex curved downwards, widely hooked inwards, tip of hook not really acute and its outer 
edge with a slight flange, in side view about twice as wide, with no ventral tooth or emargina- 
tion, spatha small, sides strongly converging, at its centre two and a half times as wide as 
sagitta ; squama small, transverse, rounded-triangular, with a small inner process separated 
by a shallow emargination where it joins the stipes ; lacinia short, only just projecting beyond 
the squama, with an acute outer angle and a moderate inner curved hook ; volsella region 
with dense, short, bristles. 

Illustrations. Sting Hazeltine & Chandler, 1964 : pi. 2. Gastral sternites 7 and 8 
Franklin, 1913 : figs. 31, 127, 129, 134. Genitalia Franklin, 1913 : figs. 85, 86, 89, 90. 



N. AMERICA. 



ii. CROTCHIIBOMB US Franklin, 1954 



$. Ocelli separated by somewhat less than two diameters from eyes, lying well in front 
of postocular line. Frons rather sparsely punctured in region of ocelli except for a densely 
punctured quadrate area immediately behind them, unpunctured areas very ill-defined, a 
number of very fine but not very close punctures between them and the eyes, nearer the latter. 
Dorsal furrow of gena absent. Antennal segments 3 : 4 : 5 = 10 : 6 : 6, segment 3 just 
longer than twice as long as broad. Clypeus short, rather strongly swollen, with not very close 
but general punctures, mostly rather small but some large, apical impressions strong, very 



232 O. W. RICHARDS 

closely punctured. Labrum with tubercles moderately convex, laterally flattened, inner end 
angled, furrow deep but narrower than length of antennal segment 3, lamella curved, moderately 
broad. Mandibles with a feeble incisura and a rather weak sulcus obliquus. Malar space 
transverse (9), a little shorter than antennal segment 3. Mid basitarsus with posterior 
apical angle feebly angled. Hind tibia with corbicular surface bare, feebly reticulate, corbi- 
cular hairs not short and dense as in B. fraternus (Smith), inner dorsal apical angle strongly 
produced, hind basitarsus with short pubescence. Gaster with pubescence short but not 
closely appressed as in Fraternobombus, tergite 6 moderately shining and coarsely rugose. 
Sting with outer thickenings rather wide but somewhat narrow in centre, inner thickenings 
widening for a short distance below, but then rather suddenly narrowing again and remaining 
of even width to dorsal end, lower part considerably darkened, intervening membrane apparently 
not differentiated but a blackened convex area near the narrowing of the inner thickenings. 
Wings evenly infuscate. 

o*. Rather large, eyes large and swollen, ocelli separated by half a diameter from eyes, 
lying far in front of postocular line. Antennae long, segments 3 : 4 : 5 : 8 = 8 : 6 : g : 9. 
Mandibles with a broad ventral and small dorsal tooth, beard long and dense. Malar space 
very short and transverse (=5), shorter than antennal segment 4, not punctured. Mid basi- 
tarsus very long and narrow (52 x 9), posterior apical angle rounded, fringes short. Hind 
tibia narrow, especially at base, gently convex, shining, with sparse short hairs throughout, 
dorsal fringe rather sparse but nearly as long as apical width of tibia. Hind basitarsus (52 x 
13), dorsal apical angle a rounded right angle, dorsal fringe not dense but nearly twice as long 
as its width. Gastral sternite 6 a little thickened but not reflexed subapically, thickening 
truncate in centre, a short dense subapical fringe. Gastral sternite 7 in the form of a wide, 
curved, transverse band, fore and hind margins nearly parallel, a large patch of short dense 
bristles on each side, no lines or fenestrae. Gastral sternite 8 a narrow band, strongly pro- 
duced into a tongue-shaped process, sides slightly convergent, angles rounded, apex approxi- 
mately straight, most of surface with dense, short bristles, two parallel lines and a small, sub- 
apical fenestra. Genitalia large, stipes rather long and broad with a wide, well-defined, inner 
impression. Sagitta narrow, slightly less so in side view, no ventral tooth, end hooked inwards, 
end of hook narrow and pointed, whole hook elongate, outer side with a slight flange which 
does not end anteriorly in a tooth. Spatha broad, sides very rapidly converging, at mid point 
three times as wide as sagitta. Squama rounded triangular, moderately large, with a small 
antero-internal process separated from the rest by a narrow emargination, process wide in the 
transversely dorso-ventral plane. Lacinia not much projecting beyond the squama, curved, 
parallel-sided, end emarginate, outer angle produced into a small acute point, inner angle 
forming a small hook, somewhat widened in dorso-ventral plane. No dense bristles on lacinia 
or volsellar region. 

Illustrations. Sting Hazeltine & Chandler, 1964 : pi. 2. Gastral sternites 7 and 8 
Franklin, 1913 : figs. 117, 130. Genitalia Franklin, 1913 : figs. 81, 88. 

WESTERN U.S.A., MEXICO. 

12. SIBIRICOBOMBUS Vogt, 1911 

Oberiobombus Reinig, 1930 

$. Ocelli separated by two diameters from eyes or rarely fully two and a half diameters, 
lying well in front of postocular line. Frons closely and rather finely punctured, unpunctured 
areas small and well-defined, a wide band of fine sculpture along inner margin of eyes. Dorsal 
furrow of gena sometimes weakly developed. Antennal segments 3 : 4 : 5 = io : 4^ : 5, 
segment 3 about four times as long as broad. Clypeus elongate, strongly swollen, rather closely 
punctured, even more so in the well-marked apical impressions. Labrum with tubercles rather 
small and convex, furrow not wide, much narrower than length of antennal segment 3, lamella 
nearly straight. Mandibles with no incisura and a well-marked sulcus obliquus. Malar space 
very elongate (= 17), longer than antennal segments 2 + 3, with many fine punctures. Mid 



SUBGENERIC DIVISIONS OF BOMBUS 233 

basitarsus with posterior apical angle somewhat produced. Hind tibia with corbicular surface 
bare, feebly reticulate, inner dorsal apical angle rather weakly produced, hind basitarsus with 
short pubescence, dorsal apical angle almost spinose, auricle covered with dense short hairs. 
Gastral tergite 6 rather strongly granulate, usually dull. Sting with outer thickenings of the 
sheath narrow, rarely a little widened above, inner thickenings very little widened below, a 
little blackened near centre in an adjacent membranous lobe, membrane also brown and con- 
vex in a small dorsal area. Wings evenly but not strongly infuscate. 

(J. Eyes normally large and swollen with temples narrow, but unmodified in B. sibiricus 
(Fabricius), when enlarged ocelli lying far in front of postocular line ; ocelli separated normally 
by one diameter from eyes, but by two and a half diameters in B. sibiricus (Fabricius). An- 
tennae very long, 3:4:5:8 = about 9 : 5^ : 8 : 9 or in B. sibiricus (Fabricius) 7 : 5 : g : 9^, 
penultimate segments nearly straight. Mandibles with a broad ventral and a small dorsal 
tooth, beard pale, dense but rather short. Malar space much longer than broad (= c. g), 
longer than antennal segments 2-1-3, with many fine punctures. Mid basitarsus very long 
and parallel-sided (38 x 8), posterior apical angle acute, fringes very short. Hind tibia not 
very elongate, strongly convex (less so in B. sibiricus (Fabricius) ), hairy, the hairs arising 
from granulate punctures, dorsal fringe very long. Hind basitarsus (40 x 12) dorsal apical 
angle acute, dorsal fringe long. Gaster with short not very close hairs, not very closely punc- 
tured or reticulate. Gastral sternite 6 little modified but usually distinctly emarginate (not 
in B. obtusus Richards). Gastral sternite 7 rounded rectangular with apex widely straight 
truncate, with considerable area of fine short hairs, no lines or fenestrae. Gastral sternite 8 
produced but sides of production strongly converging posteriorly, with a small truncate apex 
and an apical tuft of longish bristles, no fenestrae, lines absent or feebly developed at base. 
Genitalia elongate, rather large ; stipes with a strong (in B. sibiricus (Fabricius) weaker) 
sharp-edged inner impression, half its width ; sagitta narrow, dorsal half submembranous, 
strongly bent down at apex where hooked inwards, hook very long and pointed, no real outer 
flange, sagitta in side view much wider but gradually narrowing to apex, no ventral tooth or 
emargination ; spatha fully twice as wide as sagitta, almost parallel-sided ; squama rather 
long, outer and posterior edges forming a continuous curve, inner edge with a deep oval emar- 
gination leaving a wide, rounded, posterior, inner process and a narrow, acute, anterior one ; 
lacinia projecting some way beyond squama, considerably narrowed and curved inwards 
towards apex which is truncate with inner corner forming a slight hook, apex and outer edge 
with dense short bristles ; volsellar region with very dense, moderately long, stout bristles. 

Illustrations. Sting Text-fig. 9. Gastral sternites 7 and 8 Radoszkowski, 1884 : figs. 44C, 
44d ; Pittioni, I939a : pi. 4, fig. i. Genitalia Pittioni, i939a : pi. 4, fig. i. 

EASTERN EUROPE to northern ASIA and mountains of S. ASIA. 

13. CULLUMANOBOMBUS Vogt, 1911 

Rufocinctobombus Frison, 1927 

?. Ocelli separated by rather less than three or (B. rufocinctus Cresson) rather more than two 
diameters from eyes, lying well in front of postocular line. Frons closely and rather finely 
punctured, unpunctured areas small and well-defined, a wide band of fine punctures along 
inner margin of eyes. Dorsal furrow of gena sometimes developed. Antennal segments 
3:4:5 = 8:5:5^ (or B. rufocinctus Cresson 8 : 4^ : 4^), segment 3 about twice as long 
as broad. Clypeus rather swollen, rather closely and evenly punctured or (B. rufocinctus 
Cresson) shining and sparsely and more coarsely punctured, apical impressions rather deep 
or (rufocinctus) small and shallow. Labrum with tubercles rather small and flattened, some- 
what angled at inner end, furrow rather narrower than length of antennal segment 3, lamella 
short and curved. Mandibles with no incisura but with a more or less well-marked sulcus 
obliquus. Malar space just transverse (=9), a little longer than antennal segment 3 or (B. 
rufocinctus Cresson) distinctly transverse and clearly shorter than segment 3. Mid basitarsus 
with posterior apical angle rounded. Hind tibia with corbicular surface bare, finely or (B. 



234 O. W. RICHARDS 

rufocinctus Cresson) more strongly reticulate, inner dorsal apical angle produced but not strongly 
so, hind basitarsus with short, not very dense pubescence, dorsal edge strongly curved (except 
B. rufocinctus Cresson). Gastral tergite 6 dull, granulate and sparsely punctured with traces 
of a longitudinal furrow. Sting with outer thickenings of sheath considerably widened above, 
inner thickenings rather strongly widened and somewhat blackened below, two small blackened 
areas in membrane just above the widening, no other folds in membrane. Wings with apex 
darkened. 

cj. Eyes scarcely or (B. rufocinctus Cresson) more distinctly enlarged, ocelli separated by 
two and a quarter diameters from eyes or (B. rufocinctus Cresson) rather less than two diameters, 
just in front or well in front of postocular line. Antennae moderately long, segments 3:4: 
5 : 8 = 6 : 4^ : y : y (in B. rufocinctus Cresson segment 3 nearly as long as 5), penultimate 
segments straight. Mandibles with a broad ventral and small dorsal tooth, beard dense but 
not very long. Malar space just transverse (= 5), about as long as antennal segment 3, with 
numerous fine punctures. Mid basitarsus moderately long and parallel-sided (= 38 x 9), 
posterior apical angle a rounded right angle, fringes short. Hind tibia moderately clavate, 
disk convex, shining, mostly bare, with small coarse, scale-like cross-striation, dorsal fringe 
long but not dense. Hind basitarsus (38 J x n) with dorsal apical angle rounded but rarely 
acute, fringes short or (B. rufocinctus Cresson) dorsal fringe long. Gaster with dense, relatively 
short hairs. Gastral sternite 6 with its apex somewhat truncate and thickened at the angles. 
Gastral sternite 7 widely crescentic, apical margin rounded with scattered short, coarse bristles, 
apical disk with an area of close fine ones, no lines or fenestrae. Gastral sternite 8 triangularly 
produced, sides not at all parallel, apex somewhat pointed, with a small area of dense, short 
hairs, no fenestra, two parallel lines on disk. Genitalia rather long but not wide ; stipes with 
inner impressions feebly indicated ; sagittae narrow, curved down at end where hooked inwards, 
actual tip of hook a little pointed, no real flange, in side view equally narrow, not widening 
even at base, no ventral tooth ; spatha narrow, hardly widened at base, at centre hardly wider 
than sagitta ; squama transverse though not strongly so, externally more or less rounded, 
distal end feebly emarginate, inner side produced into a strong but not very acute posterior 
point and a very short anterior tubercle, inner margin generally straight-oblique ; lacinia 
projecting far beyond squama, narrow and parallel-sided, curving a little inwards, apex pointed, 
obliquely truncate, a little before apex with close short bristles on both inner and outer side ; 
volsellar region with a group of dense, long bristles. 

Illustrations. Sting Richards, 1927 : fig. 57 ; Hazeltine and Chandler, 1964 : pi. 2. 
Gastral sternites 7 and 8 Franklin, 1913 : 55, 122 ; Pittioni, I939a : pi. 3, fig. 6. Genitalia 
Kriiger, 1920 : fig. 60 ; Franklin, 1913 : figs. 79, 84 ; Richards, 1927 : fig. 36 ; Pittioni, 
I939a : pi. 3, fig. 6. 

EUROPE, N. ASIA, N. AMERICA, northern CENTRAL AMERICA. 

14. PYROBOMBUS von Dalla Torre, 1880 

Pyrrhobombus von Dalla Torre, 1882 
Poecilobombus von Dalla Torre, 1882 
Pratobombus Vogt, 1911 
Hypnorobombus Q. Perez, 1927 
Lapponicobombus Q. Perez, 1927 

This large subgenus includes species which vary much more than usual in the 
length of the malar space ; there is also some variation in the structure of the 
male genitalia. Nevertheless it scarcely seems practical at present to subdivide 
the group. 

$. Ocelli separated by nearly three diameters from eyes, lying just in front of postocular 
line. Frons mostly shining and rather sparsely punctured, in species such as B. atrocinctus 



SUBGENERIC DIVISIONS OF BOMBUS 235 

Smith with much dense, short, velvety pile, unpunctured areas large and ill-defined, no band 
of fine punctures along inner margin of eyes. Dorsal furrow of gena always indicated, some- 
times quite strong. Antennal segments 3:4:5 about = 7:4: 5^, or, B. atrocinctus Smith, 
7^ : 6 : 6, segment 3 hardly twice as long as broad. Clypeus typically moderately long and 
swollen, largely unpunctured, with characteristic deep, well-marked apical impressions con- 
taining many large punctures, in B. atrocinctus Smith these impressions weak and hardly 
punctured. Labrum with tubercles rather flattened, furrow very narrow, narrower than 
length of antennal segment 3, lamella usually short and strongly curved, wider and less curved 
in B. impatiens Cresson. Mandibles usually with a well-marked incisura, no sulcus obliquus. 
Malar space variable, from just longer than broad and a little longer than antennal segment 3, 
to clearly transverse and hardly as long as 3, length varying between 8 and 12 units, not punc- 
tured. Mid basitarsus with posterior apical angle rounded. Hind tibia with corbicular surface 
bare, shining or moderately reticulate, inner dorsal apical angle feebly or not produced (except 
B. atrocinctus Smith), hind basitarsus with short pubescence. Gastral tergite 6 shining, finely 
rather sparsely punctured or granulate. Sting with outer thickenings of the sheath narrow 
but considerably widened for a short distance dorsally, inner thickenings narrow and not 
much widened below, membrane undifferentiated except for two black central spots (copulation 
scars) in copulated specimens. Wings rather evenly but usually not much darkened. 

cJ. Typically rather small. Eyes not at all swollen, ocelli separated by at least three dia- 
meters from eyes, lying about on the postocular line. Antennae not very long, segments 
3:4:5:8 = 6:5: 6: 6, or (B. atrocinctus Smith) segment 5 a little longer (7^), penultimate 
segments straight. Mandibles with a large ventral and small dorsal tooth, beard long and 
dense. Malar space variable as in the $, from as long as antennal segments 2 + 3 (or shorter) 
to as long as segments 3 + 4, usually with scattered fine punctures. Mid basitarsus long and 
parallel-sided (32 x 7), still longer in B. atrocinctus Smith (= 46 x 9), fringes short. Hind 
tibia moderately convex, surface moderately shining, with long hairs except on distal third 
where they are short, fringes long. Hind basitarsus (=33 X 9) apical angle subacute, fringes 
short. Gastral sternite 6 not recurved and hardly thickened at apex, postapical fringe short 
and inconspicuous. Gastral sternite 7 crescentic, not at all produced, with two almost joined 
groups of bristles which are longer on each side, no lines or fenestrae. Gastral sternite 8 strongly 
produced into a subquadrate plate, sides parallel, apex feebly emarginate, angles rounded, 
most of surface covered with short bristles, no lines or fenestrae ; in B. atrocinctus Smith the 
produced part is narrower, with its sides more converging posteriorly, the end forming a rounded 
point. Genitalia typically rather short, not much longer than broad (relatively larger and 
longer in B. atrocinctus Smith) ; stipes rather narrow with effectively no inner impressions ; 
sagittae narrow, bent downwards apically in a rounded right-angle, hooked inwards without 
broad flanges, in side view somewhat wider, emarginate beneath near base, with a small tooth 
at the end of emargination ; in B. atrocinctus Smith the hook is longer with its outer edge 
straighter ; squama very small and subtriangular, mostly in a transverse dorso-ventral plane, 
with a distinct anterior inner process in B. atrocinctus Smith. Lacinia typically very small 
and entirely hidden by the squama but much longer and projecting well beyond it in B. atrocinctus 
Smith, parallel-sided, apex slightly emarginate or with a strong, curved, inner hook in B. 
atrocinctus Smith ; volsellar region normally almost bare but with long dense bristles in B. 
atrocinctus Smith. 

Illustrations. Sting Franklin, 1913 : fig. 172 ; Richards, 1927 : fig. 58 ; Hazeltine & 
Chandler, 1964 : pi. 4. Gastral sternites 7 and 8 Radoszkowski, 1884 : fig. 2c ; Franklin, 
1913 : figs. 105, 125, 131, 133, 137, 140, 142, 144, 145 ; Frison, 1925 : pi. i, figs. 2, 3 ; Prison, 
1934 : n 8 s - 5 C . d, 6c, d ; Pittioni, i939a : pi. 3, fig. 4. Genitalia Radoszkowsky, 1884 : 
figs. 2a, b ; Franklin, 1913 : figs. 64, 65, 67, 190 ; Frison, 1925 : pi. i, fig. i ; Richards, 
1927 : fig. 37 ; Frison, 1934 : n 8 s - 5 a > 6a ; Kriiger, 1920 : pi. 6, fig. p ; Pittioni, i939a : pi. 3, 
fig. 4. 

EUROPE, ASIA (including Burma and " Indo-China " and East Indies), N. 
AMERICA, C. AMERICA. 



236 O. W. RICHARDS 

15. MELANOBOMBUS von Dalla Torre, 1880 

Lapidariobombus Vogt, 1911 
Kozlovibombus Skorikov, 1922 
Tanguticobombus Pittioni, I939b 

This is another large and varied subgenus. The tip of the gaster almost always 
has red hairs (except most specimens of B. flavothoracicus Bingham). The malar 
space is normally a little longer than broad but markedly so in B. simillimus Smith 
of which the male is unknown. The labrum and other characters of B. tanguticus 
Morawitz are somewhat abnormal but there seems no reason to place it in a separate 
subgenus, at least until the male is known. B. rufofasciatus Smith and its allies 
have the eyes more or less enlarged with the ocelli approximated to them and the 
inner side of the squama more strongly emarginate than usual ; nevertheless it 
does not seem that the group is well enough marked to be treated as a distinct 
subgenus. 

$. Ocelli normally separated by nearly three diameters from eye and lying just in front 
of postocular line but in some species separated by not much more than two diameters and 
lying well in front of postocular line. Frons with some large and many fine punctures, un- 
punctured areas rather small and well-defined (larger and ill-defined in B. tanguticus Morowitz), 
a well-marked wide band of fine punctures along inner margin of eye, this band almost ceasing 
behind the level of the mid-point of the ocelli in B. rufofasciatus etc. Dorsal furrow of gena 
usually more or less indicated, absent only in B. incertus Morawitz. Antennal segments 3:4: 
5 = 8 : 5| : 6, segment 3 twice as long as broad. Clypeus long (shorter in B. tanguticus}, 
rather flattened (more convex in B. rufofasciaticus) , rather finely punctured (more punctured 
in B. tanguticus and B. rufofasciatus, and in the first-named the punctures coarser), apical 
impressions feeble and not strongly punctured except in B. tanguticus. Labrum with 
tubercles not much flattened and rather dull, more angular inwardly in B. tanguticus, furrow 
broad, wider than length of antennal segment 5, or deep and narrower and hardly as wide as 
antennal segments in B. tanguticus, lamella moderately wide, little curved. Mandibles typically 
with no incisura and a well-marked sulcus obliquus weak or absent in B. tanguticus, 
B. rufofasciatus and B. keriensis tenellus Friese. Malar space a little longer than broad 
(= 10), a little longer than antennal segment 3, but somewhat longer in B. rufofasciatus 
(as long as antennal segments 2 + 3), or considerably more elongate in B. simillimus and B. 
flavofasciatus Bingham. Mid basitarsus with posterior apical angle rounded or in B. tanguticus 
distinctly angled. Hind tibia with corbicular surface usually bare exceptionally with some 
short hairs (B. simillimus, B. tanguticus), finely reticulate inner dorsal apical angle not at all 
or very little produced. Hind basitarsus with short pubescence, on surface usually dense 
except in B. tanguticus. Gastral tergite 6 with a raised, more or less rounded, sometimes 
furrowed, largely bare boss, apex in B. rufofasciatus somewhat truncate and emarginate, or in 
B. tanguticus distinctly emarginate. Sting with outer thickenings of the sheath narrow except 
at extreme dorsal end, inner thickenings narrow, a little widened on lower third and then 
narrowed again, not blackened except rather extensively in B. simillimus, or in B. tanguticus 
with whole inner thickenings wider and margin medially quite widely reflexed and dorsally 
shortly blackened, intervening membrane very little blackened laterally or more extensively 
so in B. simillimus. Wings typically not at all darkened, even at the tip, but dark throughout 
in B. simillimus. 

c. Typically a good deal smaller than $. Eyes not enlarged and ocelli separated by three 
diameters from eyes and lying just in front of postocular line or (B. rufofasciatus, etc.) eyes 
more or less distinctly swollen (most in B. oculatus Frison) ocelli separated by about two dia- 
meters or less from eyes and lying more or less strongly in front of postocular line. Antennae 
moderately long, segments 3:4:5:8 = about 7:4:6: 5^, penultimate segments straight. 



SUBGENERIC DIVISIONS OF BOMBUS 237 

Mandibles with a large ventral and small dorsal tooth, beard long and dense. Malar space 
about quadrate (=7), not quite as long as antennal segments 2 + 3, with some fine, scattered 
punctures. Mid basitarsus rather long, a little convergent at each end (40 x 9, or B. rufofas- 
ciatus 51 x n), with long hairs on proximal half of dorsal edge and some on disk. Hind 
tibia distinctly convex, dull and reticulate, most of surface with rather sparse, moderately 
long hairs but a considerable bare disk on apical half (absent in B. rufofasciatus} fringes very 
long. Hind basitarsus (=41 X n or B. rufofasciatus 50 x 13), apical angle a rounded right 
angle or B. rufofasciatus, subacute, dorsal margin with some long hairs. Gastral sternite 6 
a little produced, thickened and recurved on central half and here a little emarginate. Gastral 
sternite 7 crescentic with centre of hind margin truncate or emarginate (rather more produced 
in B. rufofasciatus} with a large transverse patch of short bristles, no lines or fenestrae. Gas- 
tral sternite 8 not much produced posteriorly, broadly subtriangular, somewhat truncate or 
emarginate at apex, angles of truncation rounded, two small patches of bristles on each side, 
a few of the bristles long, no fenestrae but one central line. Genitalia relatively long and narrow ; 
stipes narrow with no or with weak inner impressions ; sagittae narrow in both dorsal and 
lateral views, no tooth or emargination beneath, end bent downwards and hooked inwards, 
but the hook angular and pointed like a half arrowhead ; spatha long and narrow, sides feebly 
converging, in centre twice as wide as sagitta ; squama small, not quite quadrate, rounded- 
transverse, with small rounded proximal inner projection, or (B. rufofasciatus} squama smaller 
with posterior vertically truncate and smaller broadly triangular proximal inner lobes ; 
lacinia long, projecting some way beyond squama, curved, parallel-sided, and truncate or 
emarginate so that there are two weak angles, in B. rufofasciatus the inner one forming a curved 
hook ; volsellar region with no conspicuous bristles. 

Illustrations. Sting Richards, 1927 : fig. 36 [figure on pi. 6 of Hazeltine & Chandler, 
1964 seems to be some wrongly determined species]. Gastral sternites 7 and 8 Pittioni, 
I939a : pi. 3, fig. 3 ; Prison, 1933 : n g s - 3 e f ' Prison, 1934 : n 8 s - 4 e > d. Genitalia Kriiger, 
1920 : pi. 6, fig. n ; Richards, 1927 : fig. 35 ; Prison, 1933 : fig. 3c ; Prison, 1934 : n - 4 a - 



EURASIA. 



16. PRESSIBOMBUS Prison, 1935 



$. Ocelli separated by nearly three diameters from eyes, lying just in front of postocular 
line. Frons not very closely or coarsely punctured, part behind ocelli at sides with very close 
fine punctures, temples also with very close, fine punctures, unpunctured areas ill-defined and 
not large, no band of fine sculpture along inner margin of eyes, much of head with dense, 
feathery hairs. Dorsal furrow of gena very weak. Antennal segments 3 : 4 : 5 = 8 : 5 : 6, 
segment 3 about two and a half times as long as broad. Clypeus moderately elongate, little 
swollen, with fairly numerous scattered punctures, mostly small but a few large, apical impres- 
sions well-defined with a few large punctures. Labrum with tubercles angularly raised towards 
inner end where they fall sharply on lower half (less so near clypeus), surface shining with large 
shallow punctures, furrow rather wide and shallow, at least as wide as length of antennal seg- 
ment 3, lamella very inconspicuous, rather short, feebly curved. Mandibles with a feeble 
incisura and no sulcus obliquus. Malar space about quadrate ( 13), about as long as antennal 
segments 2 + 3, not punctured. Mid basitarsus with posterior apical angle obtuse. Hind 
tibia with corbicular surface bare, rather weakly reticulate, inner dorsal apical angle moderately 
produced. Hind basitarsus densely pubescent, lower edge for its whole length and part of 
adjacent disk with long bristles. Gastral tergite 6 slight truncate, with raised boss. Sting 
with outer thickenings of the sheath not wide, but wider above than below, inner thickenings 
wide, widest just above the middle, edge forming a smooth curve, considerably blackened, 
especially near middle, membrane darkened at extreme dorsal end, otherwise undifferentiated. 
Wings moderately, evenly, infuscate. 

cJ. Relatively small. Eyes not swollen, temples wide with dense feathered pubescence, 
ocelli separated by three diameters from eyes, lying on postocular line. Antennae moderately 



238 O. W. RICHARDS 

long, segments 3:4:5:8 = 7:4:5^: 6, penultimate segments straight. Mandibles with 
large ventral and small dorsal apical teeth, beard long, dense, pale. Malar space elongate 
(= 9^), as long as antennal segments 2 + 3, not punctured. Mid basitarsus (35 x 6J) long, 
narrow, parallel-sided, posterior apical angle a rounded right angle, a few long bristles in dorsal 
fringe. Hind tibia feebly convex, strongly shining, disk bare, fringes very long but rather 
sparse. Hind basitarsus (= 37 x 10), rather long and narrowed to base, dorsal apical angle 
subacute, dorsal fringe long but sparse. Gastral sternite 6 with apex a little recurved and 
truncate. Gastral sternite 7 unusually strongly produced, apex feebly curved, a large patch 
of short, black-based bristles and a group of long feathered hairs on each side, no lines or 
fenestra ; gastral sternite 8 considerably produced, sides nearly parallel but somewhat con- 
verging posteriorly, apex truncate, angles quite well-marked, a large patch of short bristles, 
two parallel lines joining at centre and proceeding backwards as one, between them a transparent 
area, almost forming a fenestra. Genitalia relatively large ; stipes with a broad, sharp-edged 
inner impression ; sagitta in dorsal view wide at base, narrow after middle, apex moderately 
bent down, feebly hooked inwards with a small, sharp flange, externally with a much larger 
acute flange, in side view wide at base then somewhat emarginate, then widened again and then 
narrowing to apex though still relatively wide, no ventral tooth ; spatha wide, little narrowed 
posteriorly, nearly twice as wide as sagitta, acutely produced between basal inner processes 
of stipes ; squama very unusual in that it is completely fused to the stipes, represented by a 
long, narrow, acute process, directed obliquely backwards and inwards ; lacinia rather elon- 
gate, twisted so that in dorsal view it appears very narrow, apically, from side apex is obliquely 
truncate with dense rather short bristles at tip and along whole inner edge ; volsellar region 
with short moderately dense hairs. 

Illustrations. Sting Text-fig. 10. Gastral tergites 7 and 8 Prison, 1935 : figs, ic, f. 
Genitalia Prison, 1935 : fig. ib. 



HIMALAYAS. 



17. RUFIPEDIBOMBUS Skonkov, 1922 



Very large species. $. Ocelli separated by three diameters from eyes somewhat in front 
of postocular line. Frons not much punctured, punctures fine, particularly in front of and at 
sides of ocelli, unpunctured area large and ill-defined, some fine punctures along inner margin 
of eyes, much of head with dense velvety hairs as well as bristles. Dorsal furrow of gena 
more or less developed. Antennal segments 3:4:5 = 11:8:9, segment 3 not three times 
but more than twice as long as broad. Clypeus moderately long, swollen, irregularly and 
not very closely or coarsely punctured, apical impressions deep but not closely punctured. 
Labrum with tubercle strongly angled at inner end, furrow moderately distinct, about as wide 
as length of antennal segment 3, lamella wide, little curved. Mandibles with a weak or no 
incisura, no sulcus obliquus. Malar space distinctly longer than broad (= 16), nearly as long 
as antennal segments 4 + 5, feebly punctured ventrally. Mid basitarsus with apical dorsal 
corner not angled. Hind tibia with corbicular surface not reticulate, typically with sparse, 
short feathered hairs, dorsal corbicular fringe very dense, dorsal inner apical angle acutely 
produced. Hind basitarsus not densely pubescent. Gastral tergite 6 variable, finely or 
coarsely granulate, apical half keeled, swollen or hollowed out. Sting with outer thickenings 
of the sheath very narrow, more or less distinctly widened above, inner thickenings considerably 
widened below to just above the middle then narrowed again, edge considerably blackened, 
just inside this on each side a very strong, shining, convex fold of the membrane. Wings 
evenly yellowish or infuscate, or darker at tip. 

c. Large but much smaller than $. Eyes not swollen, ocelli separated by two and a half 
diameters from eyes, just in front of postocular line. Antennae very long, segments 3:4:5:8 
7i : 6J : 10 : 10, penultimate segments straight. Mandibles with a broad ventral and a 
small dorsal tooth, beard long and dense. Malar space distinctly longer than broad (= 12), 
a little longer than 2 + 3 but shorter than 3 + 4, almost unpunctured. Mid basitarsus (= 



SUBGENERIC DIVISIONS OF BOMBUS 239 

48 x n), long and parallel-sided, posterior apical angle rounded, posterior fringe nearly as 
long as its width. Hind tibia not very clavate, flat, shining, with numerous, quite long, feathered 
hairs, fringes long and dense, especially the dorsal one. Hind basitarsus (= 54 x 14) parallel- 
sided, relatively broad, posterior dorsal angle a rounded right-angle, fringes short. Gaster 
with dense, moderately short hairs, tergites closely and rather strongly punctured and strongly 
reticulate. Gastral sternite 6 smooth and shining, central third produced into rounded lobe 
which is a little thickened and reflexed. Gastral sternite 7 widely crescentic, apex shallowly 
curved, with two small groups of short bristles, no lines or fenestrae. Gastral sternite 8 very 
transverse, little produced and produced part wide, feebly emarginate at apex, a small apical 
group of bristles, disk with a bi-emarginate thickening. Genitalia with inner impressions of 
stipes large but ill-defined, inner side with a large, pale membranous band, sagitta in dorsal 
view relatively narrow, dorsal half pale and submembranous, lower half darker sclerotised, 
apex bent down and hooked inwards, hook not very long but moderately sharp with a wide 
outer flange on the bend, in side view it is very wide at base but narrows regularly towards 
the apex, with no ventral tooth or emargination, except a slight projection at the base of the 
flange ; spatha almost quite parallel-sided, a little wider than sagitta ; squama small, almost 
entirely developed on the inner side of stipes in an S-like curve with its inner end forming 
a small backwardly directed hook ; lacinia projecting far beyond the squama, broad at base 
with a central membranous area, narrowing to apex which is roundly pointed but sometimes 
irregular, almost bare, tips of lacinia tending to curve inwards and almost meeting ; volsellar 
region entirely without bristles. 

Illustrations. Sting Text-fig, n. Gastral sternites 7 and 8 Frison, 1930 : figs. 6, 7 ; 
Prison, 1934 : n s - 3 C > d. Genitalia Frison, 1930 : fig. 5 ; Frison, 1934 : n g- 3 a - 

S.W. CHINA, HIMALAYAS, FORMOSA [TAIWAN], EAST INDIES. 

Bombus festivus Smith, of which the male and worker have been unknown, seems 
now almost certainly to belong to Pyrobombus (= B. atrocinctus Smith) von Dalla 
Torre. 

18. ALPINOBOMBUS Skorikov, 1914 

Large species. $. Ocelli separated by three diameters from eyes, lying just in front of or 
on the postocular line. Frons rather feebly punctured, especially behind the level of the 
ocelli laterally, unpunctured areas ill-defined, a rather broad band of very fine sculpture along 
inner margin of eyes. Dorsal furrow of gena distinct. Antennal segments 3:4:5 = 9:6:7 
or 8 : 5 : 6, segment 3 twice as long as broad or slightly less. Clypeus moderately long and 
swollen, generally but not very coarsely punctured, apical impressions rather strong, strongly 
and closely or rather closely punctured. Labrum with tubercles flattened, not much angled at 
inner ends, furrow narrow, narrower than length of antennal segment 3, lamella short, curved, 
inconspicuous. Mandibles with no incisura and a weak sulcus obliquus. Malar space longer 
than broad, longest in B. hyperboreus Schonherr (= 16), as long as antennal segments 3 + 4, 
shortest in B. arcticus Kirby (= 12), as long as antennal segments 2 + 3- Mid basitarsus 
with apical dorsal angle a rounded right angle. Hind tibia with corbicular surface bare, hardly 
reticulate, dorsal inner apical angle shortly and broadly produced. Hind basitarsus not densely 
pubescent. Gastral tergite 6 moderately shining, granulate, sometimes coarsely. Sting with 
outer thicknenings moderately wide, especially above, inner thickenings greatly widened below, 
with two isolated blackened spots (sometimes fused), in the membrane between them. Wings 
if somewhat infuscate, evenly so. 

<J. Relatively large. Eyes not swollen, ocelli separated by three diameters from eyes, 
lying on postocular line. Antennae moderately long, segments 3 : 4 : 5 : 8 = 6J : 4^ : 8 : 8, 
penultimate segments straight. Mandibles with one broad ventral and one small dorsal tooth, 
beard long and dense. Malar space (= c. 9) about quadrate, a little longer than 2 + 3 but 
shorter than 3 + 4, with scattered fine punctures. Mid basitarsus (=38 x n) moderately 
long, rather broad, somewhat narrowed at base, posterior apical angle a rounded right angle, 



240 



O. W. RICHARDS 



a few long hairs in posterior fringe. Hind tibia rather clavate, with large, bare, convex disk, 
both fringes very long. Hind basitarsus (=42 x 13), moderately long, rather broad, posterior 
apical angle a rounded right angle, dorsal fringe very long. Hairs of gaster long and shaggy. 
Gastral tergite 6 with apical margin strongly thickened and somewhat recurved, centre feebly 
truncate, postapical fringe weak. Gastral sternite 7 rather narrowly crescentic with two 
separate groups of long bristles, no lines or fenestrae. Gastral sternite 8 produced into a tongue- 
like process, sides very converging posteriorly, almost pointed, a small posterior area of short 
bristles, no fenestrae, sometimes two weak parallel lines on disk. Genitalia relatively elongate ; 
stipes with no inner impressions ; sagitta in dorsal view rather narrow and sinuate, apex 
not bent down, with a preapical flange forming a small tooth, at apex slightly hooked on both 
sides, in side view narrow at base then considerably widened with a large central tooth, a 
smaller tooth beneath the flange, all these details varying somewhat specifically ; spatha 
wide, sides strongly converging, at centre three times as wide as sagitta ; squama large, longer 
than broad, inner and outer sides parallel and the whole structure somewhat bent inwards, 
the inner edge at a lower level feebly serrate, a slight emargination forming a very small process 
next the end of the stipes, details of squama varying specifically ; lacinia very long, projecting 
some way beyond the squama, narrowed but a little before the apex widened again, apex 
slightly emarginate, inner corner produced inwards into a feeble, rounded, serrate lobe ; bristles 
on volsellar area dense but not long. 

Illustrations. Sting Franklin, 1913 : fig. 173 ; Hazeltine & Chandler, 1964 : pi. 5. Gastral 
sternites 7 and 8 Franklin, 1913 : figs. 115, 123, 165; Richards, 1931 : pi. i ; Pittioni, 
I939a : pi. 3, fig. 3. Genitalia Franklin, 1913 : fig. 173 ; Pittioni, I939a : pi. 3, fig. 3. 

ALPS, ARCTIC EURASIA and AMERICA, Rocky Mts., Calif ornian Sierras. 

19. ROBUSTOBOMBUS Skorikov, 1922 
Volucellobombus Skorikov, 1922 

$. Ocelli separated by two diameters or rather less from eyes, lying well in front of postocular 
line. Frons with large areas both in front and behind the level of the ocelli without punctures, 
unpunctured areas very large and ill-defined, no band of fine sculpture along inner margin of 
eyes. Dorsal furrow of genera absent. Antennal segments 3 : 4 : 5 = 10 : 5^ : 6, segment 
3 rather more than twice as long as broad. Clypeus rather short, strongly swollen, with few 
or only a moderate number of punctures, apical impressions feeble, with a few large punctures 
or with numerous small ones in addition. Labrum with tubercles moderately convex and 
angled at inner end, furrow very wide, about as wide as length of antennal segment 3, lamella 
not wide, curved. Mandibles with a feeble incisura and a strong sulcus obliquus. Malar 
space (= 10) distinctly transverse, shorter than or no longer than antennal segment 3. Mid 
basitarsus with the apical dorsal angle hardly less than a right angle. Hind tibia with corbi- 
cular surface dull, densely reticulate (though much more finely than in Mendacibombus) , 
proximal third of surface with long bristles, dorsal inner apical corner with a short but acute 
process. Hind basitarsus not densely pubescent but whole surface and margins with much 




14 



FIGS. 13-14. Bombus robustus Smith. 13, gastral sternite 7 ; 14, gastral sternite 8. 



SUBGENERIC DIVISIONS OF BOMBUS 241 

coarser bristles than usual. Gastral tergite 6 shining, finely granulate. Sting with outer 
thickenings of the sheath moderately broad and of even width, inner thickenings considerably 
widened below, the wide part ending in an emargination defined by two blackened and some- 
what projecting points, the emargination sometimes bounded externally by a small blackened 
line, intervening membrane undifferentiated or raised dorsally into a horseshoe-like fold, a 
little blackened at each end. Wings infuscate. 

<J. Relatively small. Eyes rather swollen, temples rather narrow dorsally, ocelli separated 
by a little less than one diameter from eyes, far in front of postocular line. Antennae long, 
segments 3:4:5:8 = 7!- 15:8: 7, penultimate segments straight. Mandibles with one 
wide bentral and one small dorsal tooth, beard dense, moderately long. Malar space (= 4) 
transverse, about as long as antennal segment 4, scarcely punctured. Mid basitarsus (= 
42 x 8) very long and narrow, somewhat narrowed at each end, posterior apical angle completely 
rounded, dorsal margin with very long bristles and a few also on the disk. Hind tibia moderately 
clavate, convex, a little dull and reticulate, with moderately long hairs throughout, fringes 
long, especially the dorsal one. Hind basitarsus (43 x n) moderately long and broad, some- 
what narrowed basally, posterior apical angle a rounded right angle, posterior margin with 
numerous long bristles. Pubescence of gaster long and shaggy. Gastral sternite 6 with margin 
rounded, scarcely thickened, with a dense, pale postapical fringe. Gastral sternite 7 widely 
crescentic, apical margin somewhat truncate, almost no bristles, no lines or fenestrae. Gastral 
sternite 8 with a tongue-like projection with its sides converging and only becoming parallel 
near the apex which is straight, angles rounded, a large apical patch of short dense bristles, 
no fenestra, two lines which converge close to one another posteriorly. Genitalia moderately 
large and elongate ; stipes not curved in at apex, with wide, deep, inner impressions ; sagitta 
in dorsal view narrow, bent downwards and hooked inwards at apex, tip of hook with a long, 
relatively narrow point, outer side with a moderately long and broad, weakly serrate flange, 
in side view with sagitta is only about twice as wide, not widened at base and without a tooth ; 
spatha rather narrow, sides not much converging, about one and a half times as wide in centre 
as sagitta ; squama very transverse but considerably widened inwards, outer edge with a 
considerable rounded-angular projection, considerably narrowed to the actual apex and here 
with a small bristle tuft and a little emarginate with the inner corner produced into a long, 
curved hook ; volsella area with short not very dense bristles. 

Illustrations. Sting Text-fig. 12. Gastral sternites 7 and 8 Text-figs. 13, 14. Genitalia 
Franklin, 1913 : figs. 181, 197. 

Southern C. and S. AMERICA. 

20. RUBICUNDOBOMBUS Skorikov, 1922 

$. Ocelli separated by three diameters from eyes, lying a little in front of postocular line. 
Frons rather closely punctured, unpunctured areas large but not well-defined, a well-marked 
band of fine punctures along inner margin of eyes. Dorsal furrow of gena absent. Antennal 
segments 3:4:5 = 8:4: 6, segment 3 slightly more than twice as long as broad. Clypeus 
rather short, basal half much swollen, ventral third flattened, whole with numerous punctures, 
mostly large, apical impressions not defined, with close, rather small punctures. Labrum 
with tubercles convex, rather strongly angled at inner end, furrow very narrow, much narrower 
than the length of antennal segment 3, lamella not wide, moderately curved. Mandibles with 
a feeble incisura and a strong sulcus obliquus. Malar space (= 8) short and transverse, as 
long as antennal segment 3, without any punctures. Mid basitarsus with apical dorsal angle 
almost rounded. Hind tibia with corbicular surface rather dull, finely reticulate, proximal 
two fifths with rather sparse bristles on disk, dorsal inner apical corner distinctly produced. 
Hind basitarsus not densely pubescent but for the bristles on the dorsal edge. Gastral tergite 
6 dull, finely reticulate, finely and sparsely punctured. Sting with outer thickenings of the 
sheath very wide, inner thickenings gradually widening upwards from below but soon ending 
in a right-angled truncation, above this widening again in a smooth curve, margins somewhat 

ENTOM. 22, 5. 12 



242 



O. W. RICHARDS 



blackened, intervening membrane forming on each side a somewhat pyriform, convex, deep 
brown lobe. Wings dark. 

<J. Of moderate size. Eyes hardly enlarged, ocelli separated by one and a half diameters from 
eyes lying far in front of postocular line. Antennae not long, segments 3:4:5:8 = 6:4: 6J 
: 5^, penultimate segments straight. Mandibles with one broad ventral and a narrow dorsal 
tooth, beard long and dense. Malar space transverse (= 5), about as long as antennal segment 3, 
unpunctured. Mid basitarsus (= 39 x 8J) long, nearly parallel-sided, posterior apical angle 
a rounded right-angle, posterior fringe long. Hind tibia rather clavate, convex and with 
numerous rather long bristles on basal two thirds, apical third bare, weakly reticulate, dorsal 
fringe very long, ventral one much shorter. Hind basitarsus (= 38 x 13) relatively short 
and broad, somewhat narrowed to base, dorsal margin somewhat curved, with a very long 
fringe, posterior apical angle rounded. Hairs of gaster dense and rather uneven, tergites 
closely, rather coarsely punctured and reticulate. Gastral sternite 6 somewhat produced, 
especially over central quarter which is truncate and feebly emarginate, with a small depression 
embracing the truncation and extending a short way forwards. Gastral tergite 7 transverse 
with the central quarter moderately produced, shallowly emarginate, corners angled, with 
two dense tufts of bristles, disk with two suboval fenestrae, on each side of produced part 
with a semioval marginal reticulate area. Gastral sternite 8 subtriangular, the produced 
part with strongly converging sides, apex shallowly emarginate, with dense short bristles, 
disk with two parallel lines. Genitalia elongate, rather large ; stipes with a deep, sharp-edged 
inner impression over two-thirds its width ; sagitta in dorsal view narrow, not much bent 
down at tip, apex acutely pointed, behind this on inner side a small flange leading some way 
to a small inner dorsal tooth, in side view sagitta much wider, gradually narrowing to apex, 
no ventral emargination or tooth ; spatha rather narrow, about one and a half times as wide 
as sagitta, sides sub-parallel ; squama quite large, externally rounded, inner side with a large, 
deep, almost circular emargination, producing internally a wide posterior and a much narrower, 
pointed anterior process ; lacinia not projecting beyond squama, rather wide, apex truncate, 
with a dense tuft of relatively long bristles, inner end forming a small hook ; volsellar area 
with dense, short bristles. 

Illustrations. Sting Text-fig. 15. Gastral sternites 7 and 8 Text-figs. 16, 17 ; Franklin 
1913 : fig. 152. Genitalia Franklin, 1913 : figs. 186, 195. 

S. AMERICA. 



16 





FIGS. 15-17. Bombus rubicundus Smith. 15, sting 

17, gastral sternite 8. 



1 6, gastral sternite 7 



SUBGENERIC DIVISIONS OF BOMBUS 

21. COCCINEOBOMBUS Skorikov, 1922 



243 



$. Ocelli separated by two and a half diameters from eyes, lying entirely in front of post- 
ocular line. Frons with rather numerous, not very large punctures, unpunctured areas moder- 
ately large and well-defined but with a wide transverse band of fine punctures across the middle, 
another band of fine and a few large punctures along inner margin of eye, temples with a wide 
unpunctured band next to eyes. Dorsal furrow of gena absent. Antennae with segments 
3:4:5 = io : 6 : 7, segment 3 just over twice as long as broad. Clypeus a little broader 
than long, not much swollen, with moderately numerous, mostly large punctures over whole 
surface, apical impressions fairly distinct, large, with numerous punctures, some large. Labrum 
with tubercles large, strongly punctured but not angular, furrow wide and rather shallow, not 
quite as wide as length of antennal segment 3, ill-defined, lamella much wider than furrow, 
feebly curved. Mandibles with no incisura or sulcus obliquus. Malar space just transverse 
(= 17), about as long as antennal segments 2 + 3- Mid basitarsus with posterior apical angle 
about a right angle. Hind tibia with surface dull, finely reticulate, whole surface with scattered 
but quite numerous short, non-feathered hairs, corbicle moderately dense, inner dorsal apical 
angle long, acute. Hind basitarsus with apical dorsal angle just acute, no bristles on margins 
longer than half its width, surface with dense, feathered, black pubescence. Gastral tergite 6 
dull, granulate, apical quarter with a raised boss (almost as in Melanobombus] but divided 
by a deep, well-defined furrow. Sting with outer thickenings of sheath wide dorsally and 




FIGS. 18-20. Bombus coccineus Friese. 18, sting ; 19, gastral sternite 8 ; 20, (J genitalia. 



244 O. W. RICHARDS 

narrow ventrally, inner thickenings moderately widened on ventral quarter, above this narrow, 
not darkened, intervening membrane dark, rather convex dorsally. Wings subhyaline. 

o*. Relatively small and narrow ; eyes not swollen, ocelli separated by two diameters 
from eyes, lying distinctly in front of postocular line. Antennae moderately long, segments 
3 : 4 : 5 : 8 = yj : 4 : 6 : y, penultimate segments not curved. Mandibles with broad 
ventral and small dorsal tooth, beard long and dense. Malar space about quadrate (=9), 
not quite as long as antennal segments 2 + 3, unpunctured. Mid basitarsus (= 47 x 12) 
long, narrow, somewhat narrowed at each end, posterior apical angle quite rounded off, posterior 
fringe long but diffuse. Hind tibia moderately widened, surface shining but alutaceous, with 
sparse short hairs, fringes, especially dorsal one, long and dense. Hind basitarsus (= 47 x 
13) moderately long and broad, somewhat narrowed at base, posterior apical angle a rounded 
right angle, dorsal edge with a long, rather dense fringe. Hairs of gaster rather shaggy. 
Gastral sternite 6 a little recurved and thickened at apex with a dense apical fringe. Gastral 
sternite 7 transverse, apex widely emarginate with a small tooth in centre of emargination, 
two large reticulate areas on each side, just connected along posterior margin, two small lateral 
areas of short bristles, no fenestrae. Gastral sternite 8 produced in a long, rather narrow, 
mostly parallel-sided tongue, apex truncate, a large apical area of short dense bristles, two 
parallel lines on disk. Genitalia rather short and broad, dull ; stipes wide with a deep wide 
inner impression, basal inner process strong, almost parallel-sided ; sagitta in dorsal view 
narrow, ventral half much more sclerotised than dorsal half, apex bent downwards and hooked 
inwards, hook pointed, inner edge straight (almost a half arrow head) outer side with a narrow, 
very finely serrate edge, sagitta in lateral view twice as wide, no ventral tooth or emargination ; 
spatha about two and a half times as wide in centre as sagitta, sides rather converging ; squama 
short and very transverse, angles rounded, projecting inwards beyond the stipes, a second 
rather thickened inner lobe separated from the posterior one by a narrow emargination ; 
lacinia projecting a short distance beyond squama, generally curved and parallel-sided, a bit 
narrowed at apex but twisting inwards and upwards, actual tip narrow and rounded truncate, 
both inner and outer surfaces with dense short pubescence ; volsellar region with short, not 
very dense pubescence. 

Illustrations. Sting Text-fig. 1 8. Gastral sternites 7 and 8 Text-fig. 19 ; Franklin, 1913 : 
fig. 155. Genitalia Fig. 20 ; Franklin, 1913 : figs. 185, 191, 192. 

Western S. AMERICA. 

22. FUNEBRIBOMBUS Skorikov, 1922 

$. Ocelli separated by two and a half diameters from eyes, lying rather distinctly in front 
of postocular line. Frons quite closely punctured, unpunctured areas large but well-defined, 
a wide band of fine punctures along inner margins of eyes. Dorsal furrow of gena absent. 
Antennae with segments 3:4:5 = 9:5:6, segment 3 nearly two and a half times as long 
as broad. Clypeus short, swollen, with numerous punctures mostly large, apical impressions 
moderately distinct, with close large or moderately large punctures. Labrum with tubercles 
convex, angled at inner end, furrow narrow, clearly narrower than length of antennal segment 
3, lamella little curved, moderately wide. Mandibles with no incisura, ventral corner produced 
into a short, acute process, sulcus obliquus moderately strong. Malar space moderately 
transverse (=9), about as long as antennal segment 3. Mid basitarsus with apical dorsal 
angle almost rounded. Hind tibia with corbicular surface bare, feebly reticulate, inner dorsal 
apical angle almost spinosely produced. Hind basitarsus with apical dorsal angle just less 
than a right angle, surface unusually bare and shining, bristles nowhere dense or long. Gastral 
tergite 6 shining, on posterior half dull, reticulate and finely and rather closely punctured 
with traces of a feeble longitudinal furrow. Sting with outer thickenings of sheath moderately 
broad, inner thickenings gradually but not very strongly widened below, ending above in 
rounded blackened knob, above this slightly widened again and sending down an ill-defined 
process towards the knob, lower part dark and margin blackened, upper process slightly 
blackened, intervening membrane little differentiated. Wings evenly but not greatly infuscate. 



SUBGENERIC DIVISIONS OF BOMBUS 



245 



<J. Relatively small ; eyes a little swollen though temples are quite broad, ocelli separated 
by rather less than two diameters from eyes, lying well in front of postocular line. Antennae 
long, segments 3:4:5:8 = 8:5:7: 7^, penultimate segments hardly curved. Mandibles 
with a broad ventral and a small dorsal tooth, beard long and dense. Malar space just trans- 
verse (=6), a little longer than antennal segment 4 but shorter than 5, with scattered fine 
punctures. Mid basitarsus (40 x 10) moderately long and parallel-sided, posterior apical 
angle a rounded angle, posterior fringe long. Hind tibia clavate, convex, finely reticulate, 
a little shining, hairy throughout, hairs moderately long and not dense, dorsal fringe long 
and moderately dense, ventral fringe shorter and denser. Hind basitarsus ( 37 x n) some- 
what narrowed at base, posterior apical angle acute but not very sharp, dorsal fringe and 
some discal hairs near it very long, at least twice as long as width of tarsus. Hairs of gaster, 
long, shaggy, not very dense. Gastral sternite 6 hardly thickened or reflexed at apex, slightly 
truncate. Gastral sternite 7 widely crescentic, apex weakly emarginate, with a preapical 
submembranous area and two patches of long bristles before apex, no lines (except some trans- 
verse ones) or fenestrae. Gastral sternite 8 widely produced, sides of produced part concave 
and considerably diverging anteriorly, apex a little emarginate, an apical patch of dense, short 
bristles, spreading sparsely down each side, two widely separated feebly curved lines, almost 
parallel. Genitalia relatively long and large ; impressions of stipes small and ill-defined ; 
sagittae in dorsal view narrow, curved down at apex almost at apex, curving outwards, then 
inwards but apex scarcely hooked, a distinct serrate preapical outer flange, in lateral view 
sagitta about twice as wide, no ventral tooth or emargination ; spatha rather wide and strongly 
narrowing posteriorly, at centre two and a half times as wide as sagitta ; squama strongly 
transverse, short and rounded off externally, longer but also rounded off internally, inner 
edge emarginate, with a narrow, subacute anterior lobe, not projecting far inwards beyond 
the stipes ; lacinia projecting a moderate distance beyond squama, rather narrow, moderately 
curved inwards, apex with small, inner hook-like projection, outer edge with rather dense 
hairs ; volsellar region without dense bristles. 

Illustrations. Sting Text-fig. 21. Gastral sternites 7 and 8 Franklin, 1913 : fig. 153 ; 
Text-figs. 22, 23. Genitalia Franklin, 1913 : figs. 178, 180. 

Western S. AMERICA. 



21 




FIGS. 21-23. Bombus funebris Smith. 21, sting 
23, gastral sternite 8. 



22, gastral sternite 7 



246 O. W. RICHARDS 

Section ODONTOBOMBUS Kriiger 

This group of subgenera has always been rather difficult to deal with and its 
members are on the whole more uniform than those of the previous sections. The 
males are often distinct, though it is not always easy to decide to what level of 
divergence subgeneric rank should be given, but the characters of the females are 
much less pronounced and it is difficult to make a useful key to them. The arrange- 
ment adopted below is provisional ; it seemed, at this stage, it might be better to 
define as many groups as possible ; it would not be difficult later to sink some 
of them. 

23. MEGABOMBUS von Dalla Torre, 1880 
Hortobombus Vogt, 1911 

?. Ocelli separated by three diameters from eyes, lying just in front of postocular line. 
Frons finely and rather closely punctured, unpunctured areas small and well-defined, a wide 
band of fine sculpture along inner margins of eyes. Dorsal furrow of gena occasionally deve- 
loped as in B. gerstaeckeri Morawitz. Antennae with segments 3:4:5 = io : 5 : 6, segment 
3 nearly or quite three times as long as broad. Clypeus elongate, swollen dorsally, flattened 
ventrally, general surface little punctured, several rows of minute punctures along dorsal 
third of mid line, apical impressions feeble but more closely punctured. Labrum with tubercles 
rounded, little convex, largely smooth, furrow narrow but shallow, lamella rather broad and 
margin gently curved. Mandibles with no incisura, sulcus obliquus strong or moderate. 
Malar space very elongate (= 21), about as long as antennal segments 2 + 3 + 4. Mid 
basitarsus with apical dorsal angle spinosely produced. Hind tibia with corbicular surface 
bare and shining, inner dorsal apical angle with a short, wide, sharp process. Hind basitarsus 
with apical dorsal angle spinosely produced, surface rather more bristly than usual. Gastral 
tergite 6 shining, finely or coarsely granulate, often with a preapical furrow. Sting with outer 
thickenings of the sheath rather narrow throughout, more or less blackened near centre, inner 
thickenings rather strongly widened below, rather suddenly narrowed just above the middle, 
margins just above and below this extensively blackened, intervening membrane raised into 
a lobe on each side and strongly blackened at end of lobe opposite the point where the inner 
thickenings are narrowed. Wings evenly but only in a few species strongly infuscate. 

cJ. Relatively large. Eyes not at all swollen, ocelli separated by three diameters from 
eyes, lying on the postocular line. Antennae long, segments 3 : 4 : 5 : 8 = 8 : 4^ : y : y, 
penultimate segments each a little curved. Mandibles with one broad ventral and one small 
dorsal tooth, beard long and dense. Malar space very elongate (= 16), about as long as an- 
tennal segment 2 + 3 + 4, unpunctured. Mid basitarsus moderately long (= 38 x 9) and 
slightly narrowed at each end. Posterior apical angle just less than a right angle, fringes 
short. Hind tibia distinctly flattened, disk with rugose margins from which the hairs arise, 
somewhat dull and very finely reticulate, both fringes long. Hind basitarsus rather broad 
(= 36 X 13), apical posterior angle about 80, fringes short. Hairs of gaster rather long and 
dense. Gastral sternite 6 with apical margin a little thickened and recurved, hardly truncate. 
Gastral sternite 7 widely crescentic, central third of hind margin a little more produced, two 
large and narrowly separated patches of long, black bristles, no lines or fenestrae. Gastral 
sternite 8 with a wide base and central quarter with a tongue-like process, sides a little concave, 
apex straight, large apical area of short bristles, two parallel lines, no fenestrae. Genitalia 
rather large ; stipes with inner impressions small and not well-defined ; sagittae in dorsal 
view narrow, distal half serrate on lower outer edge, tip not hooked, sagitta in side view widen- 
ing only at base, no ventral emargination or tooth ; spatha narrow for most of its length, 
hardly wider than sagitta, sides scarcely convergent ; squama large, projecting far beyond 
stipes, on inner side a wide, curved upright lamella which defines and two-thirds encloses an 



SUBGENERIC DIVISIONS OF BOMBUS 



247 



oval area, all corners rounded but anterior end produced upwards into a sharp spike ; lacinia 
rather narrow, extending well beyond squama, curved inwards, tip with a peculiar process 
rather like the " toe of Italy ", volsellar region with stout, not very dense bristles. 

Illustrations. Sting Richards, 1927 : fig. 46 ; Hazeltine & Chandler, 1964 : pi. 3. 
Gastral sternites 7 and 8 Prison, 1935 : figs. 2d, e, 3C, d ; Pittioni, i939a : pi. i, fig. i. 
Genitalia Kriiger, 1920 : pi. 5, fig. a ; Richards, 1927 : fig. 26 ; Prison, 1935 : figs, ac, 3b ; 
Pittioni, i939a : pi. i, fig. i. 

EUROPE to CHINA and JAPAN, apparently not south of China. 

[Bombus melanopoda Cockerell is a true Megabombus and was described from 
Sumatra. The record requires substantiation though the type exists in the British 
Museum and there is no obvious reason to doubt the label.] 

24. DIVERSOBOMBUS Skorikov, 1914 

$. Ocelli separated by three diameters from eyes, lying just in front of postocular line. 
Frons anteriorly shining, coarsely and moderately closely punctured, posteriorly dull, very 
closely and finely punctured, unpunctured areas rather small and well-defined, a moderately 
broad band of fine sculpture along inner margin of eyes. Dorsal furrow of gena occasionally 
indicated. Antennae with segments 3:4:5 = 9:4:7, segment 3 two and a half times as 
long as broad. Clypeus elongate, swollen, finely and generally rather sparsely punctured, 
apical impressions feeble with rather closer punctures. Labrum with tubercles moderately 
convex and angled at inner end, furrow rather deep and narrow, narrower than length of 
antennal segment 3, lamella wide, little curved. Mandibles with no incisura, sulcus obliquus 
strong. Malar space very elongate (= 16), longer than antennal segments 2 + 3, nearly as 
long as 2 + 3 + 4, with a few scattered fine punctures. Mid basitarsus with apical dorsal 
angle spinosely produced. Hind tibia with corbicular surface bare, feebly reticulate, inner 
dorsal angle very feebly produced. Hind basitarsus with dorsal apical angle not very acute, 
bristles all short. Gastral tergite 6 moderately shining, coarsely granulate, apex often with 
a small convex boss. Sting with outer thickenings of sheath moderately widened, especially 
above, inner thickenings strongly, almost angularly, widened, widest well below middle, not 
blackened, intervening membrane hardened and darkened dorsally. Wings evenly infuscate. 

$. Very small compared with $. Eyes not at all swollen, ocelli separated by three diameters 
from eyes, lying just in front of postocular line. Antennae very long, each segment after 
fourth slightly curved, segments 3 : 4 : 5 : 8 = 6 : 4 : 7^ : 7^. Mandibles with one large 
ventral and one small dorsal tooth, beard dense, moderately long. Malar space very elongate 
(= 13), as long as antennal segments 2 + 3, with sparse fine punctures throughout. Mid 
basitarsus (44 X 8) rather long and narrow, slightly narrowed at each end, apical dorsal 




FIGS. 24-26. Stings of, 24, Bombus diversus Smith ; 25, B. senex Vollenhoven 
26, B. tricornis Radoszkowsky. 



248 O. W. RICHARDS 

angle a right angle, fringes short. Hind tibia only slightly convex, shining, most of surface 
with sparse, rather short bristles but a narrow, subapical disk bare, fringes moderately long 
and stout, not very dense. Hind basitarsus (= 43 x 13) moderately broad, considerably 
narrowed at base, apical dorsal angle just less than a right angle, fringes short. Hairs of gaster 
often relatively sparse. Gastral sternite 6 with apex very slightly thickened, not recurved, 
rounded, with a moderately dense postapical fringe. Gastral sternite 7 crescentic ; posterior 
margin rounded, with two rather approximated patches of moderately long bristles. Gastral 
sternite 8 considerably produced but sides of produced part clearly convergent, apex weakly 
angulated, angles almost rounded, a large apical patch of dense, short bristles. Genitalia 
relatively large and elongate ; stipes with a well-marked inner impression for more than half 
its width ; sagittae in dorsal view relatively narrow, somewhat sinuate in then out, apex 
bent down and with a well-marked transverse flange on outer side, top of flange forming a 
small tooth, sagitta in side view much broader, especially at base, emarginate beneath but 
end of emargination hardly forming a tooth ; spatha long and narrow, about as wide as sagitta, 
sides subparallel ; squama mostly narrow and transverse with its posterior margin curved 
parallel with end of stipes, posterior inner end produced into a rounded thumb-shaped process, 
anterior inner end produced into a long curved hook extending as far back as posterior margin ; 
lacinia proximally narrow, curved, parallel-sided, posteriorly with a characteristic spur-like 
process with a posterior spike and anterior crescentic serrate process, another external sub- 
triangular spike preceding the posterior one ; volsella region with long, moderately dense 
bristles. 

Illustrations. Sting Text-fig. 24. Gastral sternites 7 and 8 Prison, 1934 : figs, yc, d. 
Genitalia Prison, 1934 : n g- 7 a - 



ASIA. 



25. SENEXIBOMBUS Frison, 1930 



$. Ocelli separated by about three diameters from eyes, lying a little in front of postocular 
line. Frons moderately closely and strongly punctured, very finely and closely behind the 
ocelli, rather less close and more shining at sides, unpunctured areas large, not well-defined, 
no band of fine sculpture along inner margin of eyes. Dorsal furrow of gena absent. Antennae 
with segments 3 : 4 : 5 = 10 : 5 : 5, segment 3 two and a half times as long as broad. Clypeus 
moderately elongate, not much swollen, closely and coarsely punctured on dorsal third, rest 
unpunctured, apical impressions large, deep, moderately coarsely punctured. Labrum with 
tubercles moderately raised and angulated at inner end, shining, coarsely punctured, furrow 
wide and shallow, wider than length of antennal segment 3, lamella straight-edged, very wide. 
Mandibles with a very weak incisura and a strong sulcus obliquus. Malar space about quadrate 
(= 15), longer than antennal segment 3 but about as long as 2 + 3, with numerous fine punc- 
tures, especially on the lower half. Mid basitarsus with apical dorsal angle acutely spinose. 
Hind tibia with corbicular surface bare, very weakly convex, feebly reticulate, inner dorsal 
apical angle strongly produced. Hind basitarsus with apical dorsal angle acute, without 
long bristles. Gastral tergite 5 rather closely and coarsely punctured, tergite 6 feebly granulate, 
moderately shining. Sting with outer thickenings of sheath rather wide, even wider dorsally, 
inner thickenings rather narrow, moderately wide in middle, narrowing above and below, 
upper half blackened, blackening at about the middle, with a small downwardly directed 
process projecting into the membrane which is thrown into two folds. Wings feebly infuscate 
or yellow-brown. 

<J. Relatively large. Eyes not swollen, ocelli separated by three diameters from eyes, 
lying just on the postocular line. Antennae very long, segments 3 : 4 : 5 : 8 = 7^ : 5 : 12 : 
n, penultimate segments each a little curved. Mandibles with a broad ventral and small 
dorsal tooth, beard long and dense. Malar space considerably longer than broad ( 13), 
just longer than antennal segment 5, closely and relatively strongly punctured except for a 
small antero-dorsal patch. Mid basitarsus (= 54 x 14), long, a little narrowed at each end, 



SUBGENERIC DIVISIONS OF BOMBUS 249 

apical dorsal angle acute, fringe shorter than basal width. Hind tibia not very clavate or 
convex, densely hairy with feathered hairs of moderate length, both fringes long and dense. 
Hind basitarsus (= 60 x 18) considerably narrowed to base, apical dorsal angle acute, fringes 
short. Gaster with short rather dense hairs, tergites 1-5 with margins pale, almost white. 
Gastral sternite 6 with apex a little thickened and truncate. Gastral sternite 7 widely cres- 
centic with two closely approximated patches of short bristles, no lines or fenestrae. Gastral 
sternite 8 considerably produced but sides of production diverging anteriorly, apex feebly 
emarginate, its angles rounded, a moderate apical patch of dense short bristles, a short longi- 
tudinal line on disk and sometimes two small fenestrae on each side. Genitalia large ; stipes 
narrow distally, with wide but ill-defined depressions ; sagitta narrow in dorsal view, bent 
down at apex but not curved or widened, outer side serrate on posterior third, first tooth of 
serration much larger, in side view about twice as wide (except quite near base) emarginate 
beneath to form a strong angle near centre ; spatha very narrow hardly wider than sagitta, 
sides scarcely convergent ; squama very large, obliquely transverse, outer end rounded- 
rectangular, inner end produced into a large hook-like lobe extending well anteriorly to end 
of stipes, inside the hook can be seen a downwardly directed bifid process, the outer division 
of which is longer and more acute than the inner one ; lacinia rather narrow, curving inwards, 
apex narrow, curved upwards and outwards into a small hook, inner side with a long dense 
regular fringe continuous with a dense, long, posterior volsellar tuft. 

Illustrations. Sting Text-fig. 25. Gastral sternites 7 and 8 Prison, 1928 : pi. i, figs. 
2, 3 ; Prison, 1930 : figs. 2, 3. Genitalia Prison, 1928 : pi. i, fig. i ; Prison, 1930 : fig. i. 

EAST INDIES. 



26. TRICORNIBOMBUS Skorikov, 1922 

$. Ocelli separated by three diameters from eyes, lying just in front of postocular line. 
Frons anteriorly rather closely and coarsely punctured, behind ocelli (also laterally) very 
closely and rather coarsely punctured, unpunctured areas rather small and well-defined, a 
wide band of close fine punctures along inner margin of eyes. Dorsal furrow of gena absent. 
Antennae with segments 3 : 4 : 5 = y : 4^ : 5^ or (B. atripes Smith, 7^:5^: 6) segment 3 
not quite two and a half times as long as broad. Clypeus elongate, little swollen, closely and 
mostly quite strongly punctured, only a small ventral disk smooth, apical impressions rather 
feeble with close, coarse punctures. Labrum with tubercles only moderately raised, rather 
flattened, not angled, large outer area smooth, furrow very wide and shallow, at least as wide 
as length of antennal segments 2 + 3, lamella very wide, feebly curved. Mandibles with no 
incisura and a strong sulcus obliquus. Malar space, longer than broad (= 12 or B. atripes 
Smith, 14), about as long or nearly as long as antennal segments 3 + 4 with a few fine punctures. 
Mid basitarsus with apical dorsal angle spinosely produced. Hind tibia with corbicular surface 
bare, hardly at all convex, distinctly reticulate, inner dorsal apical angle strongly produced. 
Hind basitarsus with apical dorsal angle distinct but short, bristles short. Gastral tergites 
very closely and coarsely punctured almost throughout except for narrow apical bands, tergite 
6 dull, reticulate and with sparse, coarse granules, dense hairs more confined to sides than usual. 
Sting with outer thickenings only widened dorsally, inner thickenings very wide, bent inwards, 
becoming rather suddenly very narrow dorsally and at this point a bit blackened, membrane 
generally undifferentiated, but a convex central fold at dorsal end. Wings evenly infuscate, 
sometimes strongly so. 

cJ. Relatively small. Pyes not swollen, ocelli separated by three diameters from eyes, 
lying on postocular line. Antennae moderately long, segments 3:4:5:8 = 4:4:7:8 
or 6 : 6 : g% : 12, penultimate segments all distinctly curved. Mandibles with a broad 
ventral and small dorsal tooth, beard rather long. Malar space a little longer than broad 
(= 8 or 9), about as long as antennal segments 3 + 4, scarcely or quite strongly punctured. 
Mid basitarsus (= 39 x 9 or 55 x 13) relatively long and parallel-sided, apical dorsal angle 



250 



O. W. RICHARDS 



rounded or subacute, fringes short. Hind tibia with corbicular surface convex, shining, gran- 
ulate with coarse hairs throughout, fringes short but of very stout bristles. Hind basitarsus 
(B. atripes = 61 x 17) moderately long and parallel-sided, apical dorsal angle acute, fringes 
short. Gaster with short, not very dense hairs, sculpture more or less granulate or confluently 
punctured. Gastral sternite 6 with apex widely rounded, thickened and a bit recurved. 
Gastral sternite 7 widely or long crescentic, apex sometimes a little produced, with a few black- 
based bristles, no or two converging lines, no or two small fenestrae. Gastral sternite 8 strongly 
produced, production almost parallel-sided, apex straight or nearly rounded, with a large 
apical patch of dense short bristles, two parallel lines on disk, sometimes with an oval fenestra 
at base. Genitalia not large ; stipes with distal angle produced inwards, inner impression 
strong and sharp-edged, two-thirds its width ; sagitta in dorsal view rather narrow, distinctly 
so on bent down apical part which is not hooked but has a wide, feebly serrate, outer flange, 
in lateral view sagitta wide with a deep emargination at centre, ending in a tooth, beyond this 
emarginate again before the flange ; spatha wide, at centre one and a half times as wide as 
sagitta, sides strongly converging ; squama with an outer rounded lobe set in an oblique plane, 
considerably or only a little longer than broad, very narrow where it adjoins inner half of stipes, 
at a much lower level produced on inside into two long, very acute lobes, directed obliquely 
forwards and backwards, or one downwards and the other obliquely upwards ; lacinia very 
wide, not extending very far beyond the squama, on inner side proximally with a very long 
acute hook, edge beyond the hook straight truncate, a bit serrate with rather long bristles ; 
volsellar region with long, moderately dense bristles. 

Illustrations. Sting Text-fig. 26. Gastral sternites 7 and 8 Text -figs. 27, 28. Genitalia 
Text-fig. 29. 

East ASIA, mostly northern. 




FIGS. 27-29. Bombus tricornis Radoszkowsky. 27, gastral sternite 7 
28, gastral sternite 8 ; 29, genitalia. 



SUBGENERIC DIVISIONS OF BOMBUS 251 

27. LAESOBOMBUS Skorikov, 1922 

The males of this subgenus are easily recognized but it is not certain that the 
females can be distinguished from all members of subgenus Thoracobombus. 

$. Ocelli separated by about three diameters from eyes, lying just in front of postocular 
line. Frons anteriorly strongly and not very closely punctured, posteriorly much more finely 
and closely, unpunctured areas rather large and well-defined, a narrow band of fine punctures 
along inner margin of eyes. Dorsal furrow of gena absent. Antennal segments 3:4:5 = 
9-4^-5. segment 3 about two and a half times as long as broad. Clypeus about as long 
as broad, rather swollen, with quite numerous punctures, mostly small but a few rather larger, 
apical impressions moderately distinct with close, large punctures. Labrum with tubercles 
flattened and very little raised, not at all angled, furrow wide and shallow, not quite as wide 
as length of antennal segment 3, lamella not strongly marked, half as wide as labrum, little 
curved. Mandibles with a weak incisura and a strong sulcus obliquus. Malar space about 
quadrate (= 10), distinctly shorter than antennal segments 2 + 3, scarcely punctured. Mid 
basitarsus with apical dorsal angle spinosely produced. Hind tibia with corbicular surface 
slightly convex on basal half, shining, not reticulate, inner dorsal apical angle with a strong 
but not very acute projection. Hind basitarsus with dorsal apical angle acutely spinose, 
bristles all short. Gastral tergite 6 moderately shining, coarsely granulate, not at all upturned 
at apex. Sting with outer thickenings of sheath not wide but somewhat more so dorsally, 
inner thickenings moderately widened over most of dorsal half, slightly blackened, membrane 
undifferentiated. Wings slightly and evenly darkened. 

cj. Eyes not swollen, ocelli separated by three diameters from eyes, lying a little in front 
of postocular line. Antennae long, segments 3 : 4 : 5 : 8 = 4^ : 7 : 8 : 9^, individual seg- 
ments a little convex below. Mandibles with broad ventral and small dorsal tooth, beard 
rather long and dense. Malar space just longer than broad (= 6), about as long as antennal 
segment 4, practically unpunctured. Mid basitarsus (= 35 x 8) not long, moderately broad, 
dorsal apical angle moderately acute, fringes short. Hind tibia almost flat, mostly a little 
dull, more shining and concave on distal quarter where it is considerably widened, whole surface 
with sparse short bristles, fringes, especially dorsal one, long. Hind basitarsus (= 37 x n) 
rather short and broad, dorsal edge feebly curved, dorsal apical angle distinct, no long bristles. 
Gaster with hairs short and dense, tergites finely but not rugosely punctate. Gastral sternite 
6 with margin a little thickened, not recurved, with short, dense, pale, postapical fringe. Gastral 
sternite 7 crescentic with central part of margin a little produced and bisinuate, with a large 
continuous area of short bristles, no lines or fenestrae. Gastral sternite 8 narrow with a strongly 
produced central part which is approximately quadrate, with sides, apical margin and angles 
a little rounded, whole produced part with short bristles, two parallel lines on disk, no fenestrae. 
Genitalia rather elongate ; stipes wide, especially distally, inner impressions very weak ; 
sagitta in all views narrow except right at base, a little pointed and downcurved at apex, not 
serrate nor toothed beneath ; spatha broad, short, rapidly narrowing at centre, much wider 
than sagitta ; squama highly differentiated, large outer part mainly pale and submembranous, 
generally transverse, posteriorly produced into a wide rounded lobe, anterior to this the inner 
edge deeply emarginate and produced into an elaborate process which bears a long acute spine 
directed obliquely forwards and above this a large subcircular lobe (mainly in the vertical 
plane) with its dorsal edge serrate and the whole attached to the squama by a narrow stalk ; 
lacinia long and broad, projecting far beyond squama, broadly digitiform with end obliquely 
truncate, inner edge feebly emarginate and weakly serrate, opposite end beneath the tounded 
process of the squama produced inwards into a strong, parallel-sided process with its end 
moderately expanded and with sharp angles ; inner side of lacinia and volsellar region with 
rather dense stout bristles. 

Illustrations. Sting Text-fig. 30. Gastral sternites 7 and 8 Pittioni, i93Qa : pi. 2, 
fig. 4. Genitalia Kriiger, 1920 : pi. 6, fig. i ; Pittioni, i939a : pi. 2, fig. 4. 

SPAIN and N. AFRICA to central U.S.S.R. 



252 



O. W. RICHARDS 

28. EVERSMANNIBOMBUS Skorikov, 1938 



I am uncertain of the status of this group without further study of allied forms. 
Its characters are, however, listed. 

$. Ocelli separated by rather more than three diameters from eyes, lying just in front of 
the postocular line. Frons rather closely and strongly punctured, unpunctured areas small 
and well-defined, the areas narrowed from in front so that they are laterally pointed, two thirds 
of the space to the eye with very dense fine punctures. Dorsal furrow of gena absent. Antennal 
segments 3 : 4 : 5 = 9^ : 5 : 5, segment 3 not quite three times as long as broad. Clypeus 
long, moderately swollen with rather numerous punctures, many large, apical impressions 
small with a narrow band of close punctures. Labrum with tubercles little raised or angled, 
somewhat flattened, furrow moderately wide and deep, not quite as wide as length of antennal 
segment 3, lamella wide, straight, but not prominent. Mandibles with no incisura, sulcus 
obliquus strong. Malar space just longer than quadrate (= 12), just longer than antennal 
segment 3, much of it finely punctured. Mid basitarsus with apical dorsal angle not strongly 
spinose. Hind tibia with corbicular surface not reticulate, inner apical dorsal angle with a 
small acute process. Hind basitarsus with apical dorsal angle moderately acute, no long 
bristles. Gastral tergite 6 moderately shining, moderately coarsely and closely granulate. 
Sting with outer thickenings of the sheath rather narrow, somewhat wider above, inner thicken- 
ings narrow, somewhat widened on central two thirds, and the centre of this part of margin 
somewhat blackened, the adjacent membrane also a bit blackened but otherwise undifferen- 
tiated. Wings moderately infuscate. 

cJ. Relatively large. Eyes not swollen, ocelli separated by three diameters from eyes, 
lying just in front of postocular line. Antennae long, segments 3 : 4 : 5 : 8 = 6 : 5 : 7 : 7^, 



30 




FIGS. 30-34. 30, Sting of Bombus laesus Morawitz ; 31-34, B. eversmanniellus Skorikov, 
31, Sting ; 32, gastral sternite 7 ; 33, gastral sternite 8 ; 34, genitalia. 



SUBGENERIC DIVISIONS OF BOMBUS 253 

penultimate segments hardly curved. Mandibles with a large ventral and a small dorsal 
tooth, beard long and dense. Malar space much longer than broad (= 8J), about as long as 
antennal segments 2 + 3, shorter than 3 + 4, with quite numerous strong punctures. Mid 
basitarsus moderately long nearly parallel-sided (= 38 x 9), apical dorsal angle a right angle, 
fringes short. Hind tibia feebly clavate, rather convex, disk shining with rather sparse coarse, 
bristle-bearing punctures, fringes shorter than the width of tibia. Hind basitarsus moderately 
long and broad (= 40 x 14), apical dorsal angle just acute, all bristles short. Gaster with 
close coarse punctures and some reticulations, hairs rather long and dense. Gastral sternite 6 
flat, apex a little thickened, subtruncate, postapical fringe short, dense, pale. Gastral sternite 
7 widely crescentic, apex rounded, practically whole surface with short bristles which along 
apical margin are longer and feathered, a central thickened line and a darker spot on each 
side of base, no fenestra. Gastral sternite 8 with a strong tongue-shaped projection, sides a 
little concave, apex rounded-truncate, practically the whole process with dense bristles which 
become dense at centre of sides, usually with a very large lannceolate fenestra with thickly 
margined sides, for two-thirds of its length. Genitalia rather long and large ; stipes long and 
wide with inner impressions very feeble ; sagitta in dorsal view narrow, not much curved 
down at end, slightly widened over central fifth and again on apical fifth, this apical part 
forming a slight, very feebly serrate, lateral flange, apex hardly acute, sagitta in side view 
with basal half very wide, suddenly narrowed and after a small emargination produced into 
a wide straight truncate lobe, then another small emargination before the apical flange ; squama 
small, transversely crescentic, sides and posterior margin in one continuous curve, squama 
almost evanescent at inner posterior end of stipes but produced here inwards into a moderately 
acute process which in a more anterior view forms an obliquely-lying plate, the dorsal edge 
of this plate forming a continuous S-shaped curve with posterior margin of squama ; lacinia 
long and broad, thumb-shaped, inner edge nearly straight with very dense curved, feathered 
bristles which are half as long as its width, outer edge more curved, apex narrowly rounded ; 
volsellar region also with dense bristles. 

Illustrations. Sting Text-fig. 31. Gastral sternites 7 and 8 Text-figs. 32, 33 ; Pittioni, 
1937 : n g- 2 - Genitalia Text-fig. 34 ; Pittioni, 1937 : n g- I - 

EASTERN EUROPE. 



29. EXILOBOMBUS Skorikov, 1922 

$. Ocelli separated by three diameters from eyes, lying just in front of postocular line. 
Frons moderately strong but rather sparsely punctured, shining but with a very faint reticu- 
lation, unpunctured areas large, fairly well defined, inner margin of eye with a band of micro- 
scopic punctures which broadens posteriorly and spreads almost half way across unpunctured 
area though the fine punctures here are not very dense. Dorsal furrow of gena absent. An- 
tennal segments 3:4:5 = 8:4:4, segment 3 two and a half times as long as broad. Clypeus 
a little longer than broad, somewhat convex, with numerous punctures, many of them large, 
apical impressions very narrow but rather deep, closely punctured. Labral tubercles little 
raised, smooth and rounded, hardly punctured, furrow broad and shallow, about as broad as 
length ot antennal segment 3, lamella broad and straight, considerably thickened. Mandibles 
with a distinct sulcus, incisura weak, basal area closely, aciculately punctured. Malar space 
a little longer than broad (= 10), longer than antennal segment 3, nearly as long as 2 + 3, un- 
punctured. Mid basitarsus strongly and acutely produced. Hind tibia with corbicular sur- 
face bare, shining, excessively finely reticulate, dorsal inner apical angle strongly produced, 
hind basitarsus with dorsal angle quite strongly produced, surface shining not very densely 
pubescent, margins without long bristles. Gastral tergite 6 dull, very finely and densely 
reticulate, with fine sparse punctures. Sting with outer thickenings of sheath wide above, 
narrow below, inner thickenings moderately widened on lower half, narrow on upper half, 
transition very gradual, thickenings separated from sheath by a narrow paler area, edge of 



254 O. w - RICHARDS 

narrow part of thickening a little darkened near mid point, membrane hardly differentiated, 
with two somewhat darkened lobes above. Wings feebly infuscate. 

<. Not seen. 

Illustrations. Sting Text-fig. 35. 

EASTERN MONGOLIA and USSURI district. 

30. ADVENTORIBOMBUS Skorikov, 1922 

The status of this group is very doubtful. Originally it seems to have been 
intended for all the species of Agrobombus Vogt (= Thoracobombus) which do not 
have a serrate sagitta in the male, that is for all of them except B. pascuorum 
(Scopoli) (= agrorum (Fabricius) ). This is a large and very diverse group of 
species but as it happens B. adventor (Skorikov) is unusually distinct in the female 
(the ( has not been available for study). Whether this species should form a 
separate subgenus or whether it and perhaps some other groups should all be united 
under Thoracobombus will have to be determined in the future. The female is 
described below. 

$. Ocelli separated by three diameters from eyes, lying just in front of postocular line. 
Frons anteriorly with rather sparse, coarse punctures, posteriorly dull, finely and closely 
punctured, unpunctured areas small, moderately well-defined, a wide band of close, fine sculpture 
along inner margin of eyes. Dorsal furrow of gena absent. Antennal segments 3:4:5 = 
7i : 4i : 5, segment 3 about twice as long as broad. Clypeus elongate, swollen, with rather 
sparse, scattered punctures, mostly small, apical impressions very feeble with a row or two 
of moderately close punctures. Labrum with tubercles flattened and rounded, furrow rather 
narrow, clearly narrower than length of antennal segment 3, lamella inconspicuous. Mandibles 
with no incisura and a strong sulcus obliquus. Malar space distinctly longer than broad ( 14), 
at least as long as antennal segments 3 + 4, unpunctured. Mid basitarsus with apical dorsal 
angle long, spinose. Hind tibia with corbicular surface rather convex on basal half, shining, 
not reticulate, inner dorsal apical angle produced into a rather wide process. Hind basitarsus 
with apical dorsal angle quite strong, dorsal margin with some long bristles near base. Gastral 
tergite 6 upturned at apex, shining, with sparse, coarse, granulations. Sting with outer 
thickenings of sheath somewhat wider dorsally than ventrally, inner thickenings widening 
very gradually upwards, widest near top, then rather suddenly narrowing, a slight blackening 
of edge near centre, membrane with small central folds at top. Wings subhyaline. 

Illustration. Sting Text-fig. 36. 

INNER MONGOLIA. 



35 




FIGS. 35-37. Stings of, 35, Bombus exilis Skorikov ; 36, B. adventor Skorikov ; 
37, B. mucidus Gerstacker. 



SUBGENERIC DIVISIONS OF BOMBUS 255 

31. THORACOBOMBUS von Dalla Torre, 1880 

Chromobombus von Dalla Torre, 1880 
Agrobombus Vogt, 1911 

This is a large group with a considerable range in structure. Only some of the 
species have been examined and the characters given here are probably not found 
in all of them. It might have seemed natural to adopt the subgenus Chromobombus 
but the type-species Bombus muscorum Linnaeus raises a nomenclatorial problem 
because the ostensible type in the Linnean collection differs from the usual con- 
ception of the species. There is no doubt about the type-species of Thoracobombus, 
namely Apis sylvarum L., though it is a somewhat peripheral member of the group. 

$. Ocelli separated by three diameters from eyes, lying about on the postocular line. Frons 
shining and somewhat swollen in front with coarse, not very close punctures, posteriorly with 
very close rather fine punctures, unpunctured areas of moderate size, well-defined, a rather 
wide band of close fine punctures along inner margin of eyes. Dorsal furrow of gena absent. 
Antennal segments 3:4:5 about = 7 : 4^ : 5, segment 3 twice as long as broad. Clypeus 
clearly longer than broad, swollen, with more or less fine, scattered punctures, apical impressions 
rather strong with close, moderately coarse punctures. Labrum with tubercles flattened 
and rounded, inner ends rounded, shallow furrow, rather less wide than length of antennal 
segment 3, lamella very wide and straight, sharp-edged or with edge thickened. Mandibles 
with no incisura, sulcus obliquus strong. Malar space distinctly longer than broad (= 12), 
about as long as antennal segments 3 + 4, with scattered fine punctures, especially on distal 
half. Mid basitarsus acutely spinosely produced. Hind tibia a little convex, not reticulate, 
inner dorsal apical angle distinctly produced. Hind basitarsus with short, rather sparse hairs, 
dorsal angle acute. Gastral tergite 6 moderately shining, finely granulate, a little upturned 
at apex. Sting with outer thickenings of sheath moderately wide, especially above, inner 
thickenings strongly widened upwards from below in B. sylvarum but considerably narrower 
in many other species, the middle of the broadest part sometimes projecting as a slight spur 
and margin below this slightly blackened, membrane undifferentiated. Wings evenly but 
usually only slightly infuscate. 

$. Of moderate size. Eyes not swollen, ocelli separated by three diameters from eyes, 
lying on postocular line. Antennae moderately long, segments 3:4:5:8 typically 6 : 4^ : 
8 : 8 but varying specifically, penultimate segments slightly or (B. pascuorum (Scopoli) ) 
strongly thickened beneath. Mandibles with a broad ventral and small dorsal tooth, beard 
long and dense. Malar space distinctly longer than broad (= 8), a little shorter than antennal 
segments 3 + 4, with scattered fine punctures. Mid basitarsus moderately long and parallel- 
sided (40 x 10), dorsal apical angle acute, fringes short. Hind tibia not very clavate, surface 
convex, cross-striate, not very shining, with dense, moderately long hairs throughout, dorsal 
fringe very long. Hind basitarsus moderately long and broad (43 x 13), apical dorsal angle 
subacute, all bristles short. Gaster with dense, moderately long hairs, with granulate punc- 
tures, rather dull. Gastral sternite 6 feebly truncate. Gastral sternite 7 transverse rather 
than crescentic, side apical border with rather long bristles, no fenestrae but sometimes a 
little thinner on oval areas each side of mid-line. Gastral sternite 8 produced into a wide, 
almost parallel-sided process, apex just rounded, angles rounded, with a very large area of 
dense short hairs, no fenestrae but a thickened central line. Genitalia of moderate size ; 
stipes with no definite impressions ; sagitta in dorsal view narrow, not much desclerotized, 
apex hardly bent down, usually with small, outwardly directed, pointed hook and no flange, 
but in B. pascuorum (Scopoli) with a long, narrow, serrate flange, sagitta in side view very 
wide at base, then narrower with an emargination before the central tooth, beyond this narrow 
to apex ; spatha narrowing very soon behind base, mostly about one and a half times as wide 
as sagitta ; squama generally large, subquadrate with outer edge rounded to apex, details 
varying specifically but inner side produced obliquely into a downwardly directed acute plate 



256 O. W. RICHARDS 

or spike ; lacinia wide but not very long, not projecting much beyond squama, inner edge 
at apex produced into a small process and at centre into a spike or a narrow, truncate process, 
bristles short and not dense ; volsellar area with some long bristles posteriorly. 

Illustrations. Sting Richards, 1927 : figs. 50, 51, 52, 53 ; Hazeltine & Chandler, 1964 : 
pi. 5. Gastral sternites 7 and 8 Radoszkowsky, 1884 : figs. 2oc, 2ic, 22c ; Richards, 1927 : 
figs. 22, 24 ; Pittioni, i939a : pi. i, figs. 4, 5, 6, pi. 2, figs, i, 2, 3. Genitalia Radoszkowsky, 
1884 : figs. 20 a, b, 21 a, b, 22 a, b ; Krtiger, 1920 : pi. 5, figs, g, f, h, pi. 6, figs, k, 1 ; Richards, 
1927 : figs. 28, 29, 30, 31, 32 ; Pittioni, i939a : pi. i, figs. 4, 5, 6, pi. 2, figs, i, 2, 3. 

EUROPE to JAPAN, TIBET, but probably not south of the Chinese boundaries. 



32. MUCIDOBOMBUS Skorikov, 1922 

Exilobombus Skorikov, 1922 of which I have not been able to examine the male, 
may prove to be a synonym. As in the other groups allied to Thoracobombus, the 
males of Muddobombus are rather distinctive but the females are difficult to separate 
subgenerically. 

$. Ocelli separated by rather more than three diameters from eyes, lying just in front of 
postocular line. Frons rather closely punctured, more strongly and less closely in front of 
ocelli, unpunctured areas moderately large, well-defined, a wide band of fine sculpture along 
inner margin of eyes. Dorsal furrow of gena absent. Antennal segments 3 : 4 : 5 = 7^ : 3^ : 
4, segment 3 rather more than two and a half times as long as broad. Clypeus elongate, moder- 
ately swollen, with scattered, not at all close, fine punctures, apical impressions feeble but 
with a narrow deeper strip with close, moderately large punctures. Labrum with tubercles 
flattened, inner ends moderately raised, rounded-angular, furrow moderately deep, not broad, 
narrower than length of antennal segment 3, lamella half the width of labrum, feebly curved, 
rather thin. Mandibles with almost no incisura, sulcus obliquus strong. Malar space some- 
what longer than broad (= n), not quite as long as antennal segments 2 + 3, a considerable 
number of scattered, minute punctures. Mid basitarsus strongly spinosely produced. Hind 
tibia with corbicular surface scarcely reticulate, a little convex towards base, inner apical 
dorsal angle acutely produced. Hind basitarsus with dorsal angle acute, all bristles short 
and not numerous. Gastral tergite 6 shining, weakly granulate. Sting with outer thickenings 
of sheath rather narrow, even dorsally, inner thickenings developed for whole length, very 
gradually widening upwards, widest just before the top and for a short distance below this 
somewhat blackened, membrane undifferentiated. Wings lightly infuscate, darker towards tip. 

cJ. Relatively small. Eyes not swollen, ocelli separated by three diameters from eyes, 
lying on postocular line. Antennae with segments 3 : 4 : 5 : 8 = 6 : 3^ : 7 : 7^, penultimate 
segments a little curved. Mandibles with one broad ventral and a small dorsal tooth, beard 
long and dense. Malar space distinctly longer than broad (= 10), nearly as long as antennal 
segments 3 + 4, with scattered fine punctures. Mid basitarsus moderately long and parallel- 
sided (= 31 x 9), apical dorsal angle acute, fringes short. Hind tibia distinctly clavate, 
convex, only a small distal disk bare though the hairs are not long elsewhere, dorsal fringe 
very long. Hind basitarsus short and broad (=35 X 13), dorsal apical angle acute, fringes 
very short. Gaster with long, rather dense hairs, moderately shining with rather close coarse 
punctures and some cross-striation. Gastral sternite 6 with apex rounded, a little thickened. 
Gastral sternite 7 transversely crescentic, apex rounded, with scattered short bristles, no lines 
or fenestrae. Gastral sternite 8 produced into a moderate process which is distally parallel 
sided, apex truncate, angles rounded, a large patch of hairs posteriorly, proximal half with 
two parallel lines on disk with a small fenestra between them. Genitalia relatively long ; 
stipes wide with feeble, ill-defined inner impressions ; sagitta narrow in dorsal view, curved 
down at apex where pointed though not very acutely, no flange or serration, in lateral view 
about twice as wide, no ventral emargination or tooth, but a slight projection at proximal 



SUBGENERIC DIVISIONS OF BOMBUS 257 

end of subapical flange ; spatha small, strongly narrowing posteriorly, at middle about as 
wide as sagitta ; squama large, transverse, outer and posterior margins forming a continuous 
curve, anterior inner end produced forwards and then obliquely downwards and backwards 
in a broad but acute spike ; lacinia very long and broad, generally finger-shaped but in centre 
of inner margin with a wide lobe, denned at each end by a small tooth, the posterior one very 
acute, end of lacinia subacute, inner edge and ventral surface with dense, quite long pubescence ; 
volsella region relatively bare. 

Illustrations. Sting Text-fig. 37. Gastral sternites 7 and 8 Pittioni, i939a : pi. 2, 
fig. 5. Genitalia Kriiger, 1920 : pi. 5, fig. e ; Pittioni, i939a : pi. 2, fig. 5. 

EUROPE, WESTERN ASIA. 



33. SUBTERRANEOBOMBUS Vogt, 1911 

This subgenus has many species, especially in western and central Asia, and there 
is a considerable range in structure ; the malar space in particular varies greatly 
in length. The male genitalia, as far as they are known, though showing good 
specific differences are of the same general type. 

$. Ocelli separated by three diameters from eyes, lying about on the postocular line. Frons 
closely and rather finely punctured, unpunctured areas moderately large, well-defined, a wide 
band of fine sculpture along inner margin of eye. Dorsal furrow of gena absent. Antennal 
segments 3:4:5 about = 9:4:6, segment 3 from rather under to fully three times as long 
as broad. Clypeus elongate, swollen, either little punctured or (B. fragrans (Pallas) ) closely 
and finely punctured throughout, apical impressions feeble, little punctured or (B. difficillimus 
Skorikov) closely and finely punctured. Labrum with tubercles smooth, flattened and con- 
siderably rounded or (B. fragrans} considerably more raised and angled at inner end, furrow 
rather wide, about three-quarters as wide as length of antennal segment 3, lamella moderately 
wide, curved. Mandibles with no incisura, sulcus obliquus strong. Malar space usually 
elongate or even very elongate, longer than antennal segments 2 + 3, about = 3 + 4+1/35 
(= 15), but in B. fragrans hardly more than quadrate and about as long as antennal segments 
2 + 3 (= 13). Mid basitarsus with its dorsal apical angle usually with a relatively short and 
broad spinose production. Hind tibia with corbicular surface shining, inner dorsal apical angle 
acute but relatively short and broad. Hind basitarsus with dorsal apical angle not very acute, 
bristles all short. Gastral tergite 6 not modified, more or less coarsely granulate, sometimes 
with a furrow on the posterior half. Sting with outer thickenings of sheath of even width, 
relatively narrow, inner thickenings rather strongly widened in a very even curve, widest 
at about the middle, a strong black patch just above this, membrane on each side thrown into 
a strong fold which extends down to the middle and almost touches both the inner thickening 
and its fellow from the other side. Wings evenly infuscate, sometimes strongly. 

cj. Relatively small. Eyes not swollen, ocelli separated by three diameters from eyes, 
lying just in front of postocular line. Antennae long or very long, segments 3:4:5:8 = 
7~8 : 5 : 7^8 : 8, penultimate segments somewhat or scarcely curved. Mandibles with a 
broad ventral and small dorsal tooth, beard long and dense. Malar space elongate, as long 
or longer than antennal segments 3 + 4(=nor more), surface unpunctured. Mid basitarsus 
moderately long and parallel-sided (40 x 10 or 46 x 12), dorsal apical angle about a right 
angle, fringes short. Hind tibia with distal half flattened or even concave, covered either 
with short, not very close bristles, or with a considerable bare disk, fringes short or dorsal 
fringe long. Hind basitarsus relatively long and parallel-sided (= 41 x 13 or 47 x 13), 
dorsal apical angle just acute, bristles short. Hairs of gaster long or short, not very dense. 
Gastral sternite 6 with apex gently rounded, not thickened, with a dense, short, postapical 
fringe. Gastral sternite 7 widely and broadly crescentic with scattered bristles and closer 
short hair on disk, no lines or fenestrae. Gastral sternite 8 with a wide tongue-shaped process 

ENTOM. 22, 5. 13 



258 O. W. RICHARDS 

on central third, sides almost parallel, apex just emarginate, with dense short hairs, two weak 
parallel lines but no fenestrae. Genitalia rather large ; stipes with a deep, wide, sharp-edged, 
inner impression ; sagittae in dorsal view generally narrow, apex a little curved downwards, 
a slight broadening at a lower level near centre and apex outwardly triangularly widened or 
with a triangular pointed flange, sagitta in side view broad on the basal half or rather more, 
then narrowed and emitting a process ending in two or three small points ; spatha narrow, 
almost parallel-sided, about as wide as sagitta ; squama transverse or transversely S-shaped, 
all angles, or at least the external ones, rounded, inner side sometimes produced into a vertical 
lamella which looks sharp in dorsal view ; lacinia little or moderately projecting beyond squama, 
rather narrow, a little curved inwards, end pointed or almost rounded, sometimes with a small 
apical tuft ; volsellar region with short dense bristles. 

Illustrations. Sting Franklin, 1913 : fig. 157 ; Richards, 1927 : fig. 49 ; Hazeltine & 
Chandler, 1964 : pi. 6. Gastral sternites 7 and 8 Radoszkowski, 1884 : figs. 26c, 36c ; 
Franklin, 1913 : fig. 138 ; Pittioni, i939a : pi. i, fig. 2. Genitalia Radoszkowski, 1884 : 
figs. 26a, b, 36a, b ; Franklin, 1913 : fig. 157 ; Kriiger, 1920 : pi. 5, figs, c, d ; Richards, 
1927 : fig. 27 ; Pittioni, i939a : pi. i, fig. 2. 

EUROPE, ASIA to the HIMALAYAS, N. AMERICA. 

34. RHODOBOMBUS von Dalla Torre 1880 
Pomobombus Kriiger, 1917 

<. Ocelli separated by three diameters from eyes, lying just in front of postocular line. 
Frons finely, mostly not very closely punctured, a large area in front of median ocellus without 
punctures, unpunctured areas large but well-defined, a narrow band of fine punctures along 
inner margin of eye. Dorsal furrow of gena absent. Antennal segments 3:4:5 = 9:4: 4^, 
segment 3 two and a half times as long as broad. Clypeus elongate, moderately swollen, finely 
and rather sparsely though generally punctured, apical impressions small and narrow but 
rather well-defined, with a few close larger punctures, mostly in transverse rows. Labrum 
with tubercles somewhat flattened and rather rounded, furrow of moderate width, about 
three-quarters as wide as length of antennal segment 3, lamella wide, gently curved. Mandibles 
with no incisura, sulcus obliquus strong. Malar space elongate, about as long as antennal 
segments 2 + 3 (= 13!), with a certain number of scattered fine punctures. Mid basitarsus 
with its dorsal apical angle acutely spinose or in some American species only somewhat angled. 
Hind tibia with corbicular surface somewhat convex on posteroventral part, surface bare, 
weakly reticulate, inner dorsal apical angle not or scarcely produced. Hind basitarsus spinosely 
produced at dorsal apical angle, bristles short. Gastral tergite 6 finely granulate, moderately 
shining. Sting with outer thickenings of sheath rather narrow but somewhat widened above, 
inner thickenings evenly widened for most of their length, edge somewhat emarginate above. 
Wings usually moderately infuscate. 

$. Size moderate. Eyes normal, ocelli separated by three diameters from eyes, lying 
just in front of postocular line. Antennae rather long, segments 3:4:5:8 = 6:4:7: 7^, 
penultimate segments hardly curved. Mandibles with a broad ventral and a small dorsal 
tooth, surface granulate and dull, beard not developed. Malar space elongate, about as long 
as antennal segments 3 + 4 (= 9i), with scattered very fine punctures. Mid basitarsus 
moderately long and parallel-sided (34 x 8), dorsal apical angle a rounded right-angle, fringes 
short. Hind tibia thick and little clavate, surface convex and shining with scattered punctures 
and some cross-striation, with hairs over whole surface, fringes not dense nor longer than its 
width. Hind basitarsus moderately broad and parallel-sided (37 X 13), dorsal apical angle 
acute, fringes short. Gaster with close moderately short hairs. Gastral sternite 6 slightly 
thickened and truncate with a short subapical fringe. Gastral sternite 7 somewhat produced 
in a broad triangle with rounded apex, well behind apex darker transverse band running parallel 
to margin, moderate bristles arising on and before the band, no lines or fenestrae. Gastral 



SUBGENERIC DIVISIONS OF BOMBUS 259 

sternite 8 narrowly transverse with a strongly produced parallel-sided process, angles rounded, 
apex bifid though the emargination partly filled in by transparent membrane, with tufts of 
moderate bristles on each side, a pair of parallel lines on disk. Genitalia of moderate size ; 
stipes with well-defined inner impression of half its width ; sagittae in dorsal view rather 
narrow, not at all desclerotized, apex a little bent down, curved outwards into a truncate 
hook whose anterior end forms a small acute tooth, in side view at least twice as wide with a 
well-marked emargination after base and a strong acute tooth at middle ; spatha moderately 
broad, gradually narrowing posteriorly, at centre one and a half times as wide as sagitta ; 
squama transverse, outer edge curving obliquely inwards and ending in a stout acute process 
at inner posterior dorsal corner, inner edge from anterior inner corner produced obliquely 
downwards and backwards into a twisted plate of the end is hardly acute ; lacinia projecting 
a moderate distance beyond the squama, narrowing to apex and then expanded inwards into 
a small tooth, lacinia almost bare, no dense bristles on volsellar area. 

Illustrations. Sting Richards, 1927 : fig. 48 ; Hazeltine & Chandler, 1964 : pi. 5. Gastral 
sternites 7 and 8 Radoszkowski, 1884 : figs. i8c, igc, 27c ; Pittioni, i93ga : pi. i, fig. 3. 
Genitalia Radoszkowski, 1884 : figs. i7a, b, i8a, b, iga, b, 27a, b ; Kriiger, 1920 : pi. 5, 
fig. b ; Richards, 1927 : fig. 26 ; Pittioni, i939a : pi. i, fig. 3. 

EUROPE to IRAN and CENTRAL ASIA but not beyond the HIMALAYAS or in CHINA. 

35. FERVLDOBOMBUS Skorikov, 1922 

This is a large and varied subgenus with a considerable range of structure amongst 
the species ; moreover several of them are not available in England. 

?. Ocelli separated by three diameters from eyes, lying just in front of postocular line. 
Frons swollen in front and coarsely punctured (closeness varying with the species), very closely 
and finely punctured behind, unpunctured areas rather large and ill-defined, a rather wide 
band of fine punctures along inner margin of eye. Dorsal furrow of gena absent. Antennal 
segments 3 : 4 : 5 = c. g : 4 : 6, segment 3 two to two and a half times as long as broad. 
Clypeus elongate, swollen, rather strongly and closely punctured, especially at sides, disk 
below rather widely unpunctured or with fine punctures, a line of several rows of fine punctures 
on mid line of dorsal third, apical impressions rather distinct with a few but large close punc- 
tures. Labrum with tubercles little raised, flattened and rounded, furrow rather narrow, 
not as wide as length of antennal segment 3, lamella wide, shining, thickened, edge nearly 
straight. Mandibles with no incisura, sulcus obliquus strong. Malar space quadrate to elongate, 
(13-18), from about as long as antennal segment 3 to rather longer than 2 + 3, usually with 
some fine punctures. Mid basitarsus with apical dorsal angle not or very feebly spinose 
produced into a relatively wide angular process. Hind tibia corbicular surface bare, feebly 
reticulate, with posterior ventral part convex, sometimes subangular in proximal part, inner 
dorsal apical angle not or little produced. Hind basitarsus not very acutely produced, bristles 
short. Gastral tergite 6 shining, feebly granulate. Sting variable, outer thickenings of sheath 
usually much wider above than below, inner thickenings strongly widened below then narrowing 
from about the midpoint where there is a large blackened area, membrane above thrown into 
a convex central fold. Wings usually strongly darkened. 

(J. Size moderate. Eyes normal, ocelli separated by three diameters from eyes, lying just 
in front of postocular line. Antennae long, segments 3:4:5:8 = c. 7:4$ : 8 : 8, penul- 
timate segments a little curved. Mandibles with broad ventral and small dorsal tooth, beard 
long, dense and brown. Malar space quadrate to rather elongate (= 11-14), usually a little 
longer than antennal segments 3 + 4, with scattered fine punctures. Mid basitarsus rather 
long and parallel-sided (44 x 9) dorsal apical angle rounded right-angle, fringes short. Hind 
tibia not very long, convex, hairy all over, coarsely cross-striate and obliquely punctured, 
fringes short. Hind basitarsus parallel-sided and rather broad (48 x 16), apical dorsal angle 
a right angle, fringes short. Gaster with hairs rather short and even. Gastral sternite 6 

ENTOM. 22, 5. I3 



260 O. W. RICHARDS 

with end rounded and a little thickened. Gastral sternite 7 very broadly crescentic posteriorly 
rounded, with two distinct discal patches of short bristles, no lines or fenestrae. Gastral 
sternite 8 with central quarter produced into a wide, almost parallel-sided tongue-shaped 
process, apex slightly emarginate, with a large area of dense short bristles, a central line which 
forks a little before the bristle patch. Genitalia of moderate size ; stipes with a moderately 
well-defined impression of half its width ; sagitta seen from above narrow, not bent down 
until just before apex where bent outwards into a hook-like structure of which the outer margin 
is straight-serrate, in side view sagitta considerably broader on basal half up to the mid-ventral 
tooth, then relatively narrow ; spatha narrow, sides not convergent, at centre about twice 
as wide as sagitta ; squama transverse, outer and posterior edges forming a continuous curve 
to a blunt exterior point on outer edge of stipes, inner margin produced into a vertical, plate- 
like anterior lamella and a more dorsal wider rounded process ; lacinia produced a very little 
way beyond squama, strongly widened at apex, angularly produced at outer end and into a 
small curved hook at inner end, volsellar region with dense bristles. 

Illustrations. Sting Moure & Sakagami, 1962 : fig. 3 ; Hazeltine & Chandler, 1964 : 
pi. 3. Gastral sternites 7 and 8 Radoszkowski, 1884 : fig. 28c ; Franklin, 1913 : figs. 32, 
101, 126, 132, 148, 174 ; Moure & Sakagami, 1962 : fig. 4. Genitalia Radoszkowski, 1884 : 
figs. 28a, b ; Franklin, 1913 : figs. 66, 70, 78, 102, 156, 159-164, 166-168, 170-171, 179, 182- 
183, 193, 196 ; Moure & Sakagami, 1962 : figs. 5, 6. 

NORTH and SOUTH AMERICA. 



KEYS TO THE SUBGENERA OF BO MB US LATREILLE 

It is very difficult to make practical keys to the subgenera of Bombus, especially 
in the female sex. Apart from the intrinsic difficulties, few authors have seen more 
than a fraction of the species. There are of course also many nomenclatorial 
problems and points of doubt at the specific and subspecific level, though these 
do not affect a subgeneric key in principle. The keys which I have constructed 
are tentative and I would not claim that an inexperienced entomologist would 
find them very useful ; even the expert will find that some species are troublesome 
and they may also detect errors. Nevertheless, no really comparable attempt 
has been made before and I hope that it may constitute a foundation for more 
successful efforts in the future. It is impossible at the moment to make useful 
keys to the worker caste, which is far too variable and often lacks the diagnostic 
characters of the female. In any district, when a study of the males and females 
has shown which species are present, it is usually possible to identify most of the 
workers. 

MALES 
<J of B. adventor (Skorikoy) and B. exil (Skorikov) were not available. 

i Mandibles distally ending in one large ventral and two small dorsal teeth. Antennal 
segment 4 0'73-o-74, segment 5 0-93 as long as segment 3. Mid and hind basi- 
tarsi with long fringes, hind tibia with a bare disk and very long dorsal and ventral 
fringes. Ocelli separated by three diameters from eyes. Genitalia with spatha 
very wide but strongly narrowed posteriorly ; squama transverse or quadrate, 
inner side proximally with a small process separated by a moderate emargination 
from the posterior lobe ; sagittae hooked inwards, tip blunt, outer side of hook 
with a considerable flange .......... 2 



SUBGENERIC DIVISIONS OF BOMBUS 261 

Mandibles distally ending in one large ventral and one small dorsal tooth (absent 

in some M ' endacibombus) .......... 3 

Malar space a little shorter than antennal segment 3. Squama rounded-quadrate 

or with a long, pointed inner process . . . ALPIGENOBOMBUS (p. 221) 

Malar space half as long as antennal segment 3. Squama very transverse 

NOBILIBOMBUS (p. 222) 

Sagittae narrow and pointed. Antennae very long, antennal segment 4 i* 1-1*4, 
segment 5 1*5-1*7 times as long as segment 3. Malar space nearly twice as long 
as antennal segment 3. Mid and hind basitarsi with short fringes, hind tibia 
bare on disk. Ocelli separated by three to three and a half diameters from eye. 
(Pakistan to W. China) ORIENT ALIBOMBUS (p. 224) 

If the sagittae are of this type, the ocelli are much closer to the eyes or the hind 
tibia is hairy, or if there is a small bare disk, the fourth antennal segment is about 
half as long as the third .......... 4 

Sagittae curved outwards in a rounded hook. Antennal segments 4 and 5 1*2 
and 1*5 times as long as segment 3. Ocelli separated by three diameters from 
eyes. Mid and hind basitarsus without long fringes, hind tibia with a bare disk and 
long fringes. Gastral sternite 7 strongly produced, sternite 8 with a narrow, 
parallel-sided, tongue-shaped projection. Stipes with the inner impression very 
feeble ; squama twice as long as broad. (Europe) . . KALLOBOMBUS (p. 225) 

Sagitta not curved outwards into a recurved hook. Antennal segments 4 and 5 
relatively shorter (4 shorter than 3) except in a few species with short basitarsal 
fringes ............. 5 

Antennal segments 4 and 5 0*36-0-56 and 0-48-0-78 times as long as segment 3. 
Eyes swollen, ocelli separated from them by one diameter or less. Mandibular 
beard usually not very well developed. Sagittae narrow and pointed, or finger- 
shaped, only in one group curved inwards with a flange outside the curve 

(Section BOOPOBOMBUS) 6 

Antennal segments 4 and 5 relatively longer and sagittae usually of a different 
shape, particularly if the antennae are at all similar. Ocelli usually less close 
to the eyes ............. 9 

Sagittae curved inwards with a small flange on the outside of the curve. Squama 
rather elongate, inner edge emarginate. Spatha narrow. Gastral sternite 7 
long trapeziform, apex deeply emarginate. Gastral sternite 8 strongly produced, 
production tongue-shaped. Mid and hind basitarsi and hind tibia with short 
fringes. (S.W. America) FRATERNOBOMBUS (p. 227) 

Sagittae pointed or finger-shaped. Hind basitarsus with a long dorsal fringe . 7 

Squama in section V-shaped, formed of two plates set at an angle and joining at 
their outer edge. Genitalia small, pyriform. Spatha very wide, narrowing 
distally. Gastral sternite 7 crescentic but with rounded-triangular production. 
Gastral sternite 8 crescentic, little produced, subtruncate. Mid basitarsus with 
a fringe as long as its width, fringe of hind tibia long CONFUSIBOMBUS (p. 228) 

Squama subquadrate, all angles rounded, not V-shaped in section ... 8 

Mid basitarsus with fringe mostly short, hind tibia with long fringes. Antennal 
segment 3 unusually long. Gastral sternite 7 crescentic, sternite 8 produced, 
sides a little rounded, apex distinctly emarginate. Spatha about twice as wide 
as sagitta. (Mountains of Eurasia) ... MENDACIBOMBUS (p. 229) 

Mid basitarsus with a long fringe, hind tibia with very short ones. Gastral sternite 
7 broadly triangular, apex feebly emarginate, sternite 8 broadly triangular with 
apex deeply emarginate. Spatha very wide, 5 times as wide as sagitta. (N. 
America) BOMBIAS (p. 230) 

Sagittae in the form of wide sinuate vertical plates. Lacinia projecting very little 
beyond squama which on inner side has a deep anterior emargination of which 
the proximal bounding lobe is linear. Spatha very wide. Mid and hind basitarsi 



262 O. W. RICHARDS 

with short fringes, hind tibiae with a bare disk and both fringes long. Ocelli 
separated by three diameters from eyes. Antennal segments 4 and 5 0-7 and 
0-9 times as long as segment 3. (Eurasia and N. America) . BOMBUS s.s. (p. 226) 
- Sagittae of a quite different form . ........ 10 

10 Sagitta ending in a strong, inwardly directed, recurved hook. Mid basitarsus with 

short fringes. Antennal segment 5 longer or (in Separatobombus) just shorter 

than 3 . . . . . . . . . . . ...n 

Sagitta rarely ending in a strong, inwardly directed, recurved hook and if so the 

mid basitarsus with a long fringe ......... 16 

11 Ocelli separated by less than one diameter from eyes. Malar space shorter than 

antennal segment 4. Hind basitarsus with a long fringe. Sides of spatha 
strongly convergent posteriorly. Squama a small or rather small rounded 
triangle, lacinia hardly projecting beyond it. (America) . . . . . 12 
Ocelli separated by more than one diameter from eyes. Malar space as long or 
longer than antennal segment 3. Sides of spatha subparallel, little convergent 
posteriorly. Sagitta ending in a more or less rounded, blunt-ended hook . . 13 

12 Sagitta ending in a rounded hook, tip not acute. Impression of stipes wide but 

ill-defined. Antennal segment 5 shorter than 3. Gastral sternite 7 trapeziform, 
apex feebly emarginate, sternite 8 subtriangular . SEPARATOBOMBUS (p. 231) 
Sagitta with a somewhat smaller hook, tip acute. Impression of stipes wide and 
well-defined. Antennal segment 5 longer than 3. Gastral sternite 7 subcres- 
centic, sternite 8 with a parallel-sided, tongue-like projection 

CROTCHIIBOMBUS (p. 231) 

13 Squama considerably longer than broad, inner side emarginate, lacinia also elongate 

and projecting well beyond it. Impression of stipes strong, sharp-edged. An- 
tennal segment 5 clearly shorter than 3. Hind basitarsus with a long fringe. 
Ocelli separated by one to two and a half diameters from eyes. Gastral sternite 
7 rounded triangular, widely truncate, sternite 8 rounded triangular but slightly 
truncate or if produced, sides of production strongly divergent. (Eurasia). 

SIBIRICOBOMBUS (p. 232) 

Squama not clearly longer than broad, inner side not emarginate. Impression of 
stipes feebly defined. Antennal segment 5 longer than 3. Gastral sternite 7 
trapeziform or crescentic. 

14 Lacinia scarcely projecting beyond the squama (more so in B. atrocinctus Smith) 

which is small and rounded triangular. Hind basitarsus with a short fringe. 
Ocelli separated by at least three diameters from eye. Gastral sternite 8 with a 
parallel-sided, tongue-like projection. (Eurasia, N. America) PYROBOMBUS (p. 234) 
Lacinia very long, projecting well beyond the squama. Ocelli separated by less 

than three diameters from eyes. Gastral sternite 8 widely subtriangular . . 15 

15 Squama just transverse with inner side emarginate. Hind basitarsus with a short 

or long fringe. Malar space not longer than antennal segment 3. Ocelli separated 
by one and three quarters to two and a quarter diameters from eyes. (Eurasia 

and N. America) CULLUMANOBOMBUS (p. 233) 

Squama S-shaped, inner end pointed, almost the whole of it lying inside the inner 
margin of the stipes. Hind basitarsus with a short fringe. Malar space longer 
than antennal segment 3. Ocelli separated by two and a half diameters from 
eyes. (E. Asia) RUFIPEDIBOMBUS (p. 238) 

1 6 Mid and hind basitarsi with long fringes . . . . . . . . 17 

Mid and hind basitarsi with short fringes. Ocelli separated by three diameters 

from eyes. Malar space more or less elongate (Section ODONTOBOMBUS) 23 

17 Sagitta at end not curved inwards or outwards but with two external teeth, one 

of which is sometimes small. Antennal segment 4 not much shorter than 3, 
5 one and a quarter times longer than 3. Malar space in most species distinctly 
longer than antennal segment 3. Gastral sternite 7 crescentic, 8 subtriangular. 



SUBGENERIC DIVISIONS OF BOMBUS 263 

Spatha wide, sides strongly converging ; stipes with no inner impression ; 
squama longer than broad ; lacinia very long, apex on inner side produced into 
a feeble, rounded, serrate lobe. (Arctic-Alpine in Eurasia & N. America but 
not in mountains of S.E. Asia) .... ALPINOBOMBUS (p. 239) 

Sagitta sometimes curved inwards but never with more than one outer tooth. 

Antennal segment 4 not more than three quarters as long as segment 3, 5 not 
more than one tenth longer than segment 3. Malar space rarely longer than 
antennal segment 3. ........... 18 

1 8 Lacinia projecting far beyond squama. Malar space as long as antennal segment 3 

or a little longer. Sagitta with a sharp projection on inner side of apex. An- 
tennal segment 4 0-57, segment 5 about 0-80 as long as segment 3. (Eurasia) . 19 

Lacinia relatively short, projecting at most a moderate distance beyond the squama 

which has rounded angles and an inner emargination. Malar space usually 
shorter than antennal segment 3. Ocelli separated by two diameters or less 
from eyes. (S. America) .......... 20 

19 Squama completely fused with the stipes, produced into a long acute process directed 

downwards and backwards. Sagitta at apex with a small pointed flange on inside 
and a large pointed one on outside. Gastral tergite 7 widely trapeziform, 8 with a 
parallel-sided projection. Stipes with a broad, sharp-edged impression. Lacinia 
twisted so that its plane is oblique distally, with dense short hairs at apex and 
on inner side. Ocelli separated by three diameters from eyes. (Himalayas) 

PRESSIBOMBUS (p. 237) 

Squama separate from stipes, rounded-transverse with a small, proximal inner 
process. Sagitta with an acute apex and an acute inner flange (half arrowhead). 
Gastral tergite 7 crescentic, 8 broadly triangular. Stipes narrow without an 
inner impression. Lacinia not twisted, with few hairs. Ocelli variable, separated 
by 1-3 diameters from eyes. (Eurasia). . . MELANOBOMBUS (p. 236) 

20 Antennal segment 5 a little longer than 3. Impression of stipes wide and deep. 

Spatha relatively narrow, about one and a half times as wide as sagitta, sides 

not very convergent posteriorly . . . . . . . . .21 

Antennal segment 5 distinctly shorter than 3. Impression of stipes not more than 

half its width. Squama rounded transverse, inner side emarginate . . . 22 

21 Sagitta curved inwards at apex with a serrate flange on the outside. Spatha narrow, 

sides subparallel. Squama very transverse, widening inwards, inner edge emar- 
ginate. Lacinia with a rounded-angular projection on the inner edge, inner corner 
of apex with a long curved hook. Gastral sternite 7 crescentic, 8 with a tongue- 
like projection whose sides converge. Malar space hardly more than half as 
long as antennal segment 3. Ocelli separated by less than one diameter from 

eyes ROBUSTOBOMBUS (p. 240) 

Sagitta slightly curved inwards at apex and with a small inner tooth. Squama 
rounded quadrangular with a large subcircular inner emargination. Lacinia 
wide, posterior end with a short hook, end with a dense tuft of relatively long 
bristles. Gastral sternite 7 transverse with central quarter produced, 8 sub- 
triangular. Malar space a little shorter than antennal segment 3. Ocelli separ- 
ated by one and a half diameters from eyes . RUBICUNDOBOMBUS (p. 241) 

22 Stipes with strong impressions. Sagitta with a somewhat inward-curving pointed 

hook on inside and a narrow serrate flange outside. Lacinia short, apex with 
a small posteriorly directed hook, outer and ventral side with dense short hairs. 
Gastral sternite 7 transverse, apex biemarginate, 8 with a long, parallel-sided 
process. Malar space slightly longer than antennal segment 3. Ocelli separated 
by two diameters from eyes COCCINEOBOMBUS (p. 243) 

Stipes with small ill-defined depressions. Spatha wide with strongly convergent 

sides. Sagitta with apex hooked inwards, externally with a serrate flange. 
Lacinia moderately long, apex with small hook-like projection at inner end. 



264 O. W. RICHARDS 

Gastral sternite 7 crescentic, weakly emarginate, 8 subtriangular. Malar space 
three quarters as long as antennal segment 3. Ocelli separated by less than 
two diameters from eyes FUNEBRIBOMBUS (p. 244) 

23 Lacinia long and narrow, produced inwards at apex into a process like a toe and 

heel. Sagitta narrow, outer side distally strongly serrate (character otherwise 
seen only in B. pascuorum (Scopoli) Thoracobombus), no tooth beneath. Squama 
forming a large elongate, vertical, curved plate on inner side two thirds enclosing 
an oval space, anterior lower corner produced upwards into a sharp spike. Malar 
space as long as antennal segments 2 + 3 + 4- (Eurasia) MEGABOMBUS (p. 246) 

Lacinia long and narrow, produced inwards at apex into a spur-like process with 

several points. Sagitta narrow, sinuate, with an apical outer flange forming a 
small tooth at its proximal end, emarginate beneath but not forming a real tooth. 
Squama narrow and transverse, posterior inner corner produced into a thumb- 
shaped process, anterior inner corner with a long curved hook, extending as far 
back as posterior margin. Malar space as long as antennal segments 2 + 3. 
(E. Asia) DIVERSOBOMBUS (p. 247) 

Lacinia at apex produced at most into a small hook or serrate, rounded lobe. In 

Senexibombus, somewhat similar to Megabombus, but the sagitta is serrate only 

at the recurved distal quarter ......... 24 

24 Antennal segment 5 much (x 1-5-1-9) longer than segment 3 .... 25 

Antennal segment 5 little ( x 1-0-1-3) longer than segment 3 .... 27 

25 Antennal segment 4 less than 0-7 times as long as 3. Spatha very narrow. Squama 

very large, transversely oblique, proximal inner end produced into a large hook, 
beneath this hook can be seen a downwardly directed bifid process, the outer 
division longer and more acute than the inner one. Sagitta narrow, outer side 
serrate on distal quarter, first tooth of serrations large, beneath with a strong 
angle rather than a tooth. Malar space nearly as long as antennal segments 3 + 4. 
(E. Asia) SENEXIBOMBUS (p. 248) 

Antennal segment 4 as long as or longer than segment 3. Spatha very broad with 

convergent sides ............ 26 

26 Lacinia very wide, not extending far beyond outer part of squama, on inner side 

proximally with a very long acute hook, inner edge behind the hook straight 
truncate, somewhat serrate, with long bristles. Squama with elongate outer 
lobe set in an oblique plane, on inner side at a lower level produced into two very 
long acute processes, directed obliquely forwards and backwards respectively. 
Sagitta apically with an outer feebly serrate flange, beneath with a tooth. Malar 
space about as long as antennal segments 3 + 4. Gaster very closely punctured. 
(E. Asia) TRICORNIBOMBUS (p. 249) 

Lacinia very long, broadly digitiform, near centre on inner side produced into a 

strong, parallel-sided process with an expanded end with sharp angles. Squama 
with large, outer part pale and submembranous, generally transverse but posteri- 
orly produced on inside into a rounded lobe, before this deeply emarginate and 
then produced into a large subcircular lobe (mainly in a vertical plane) with its 
dorsal edge serrate and the whole attached to the squama by a narrow stalk. 
Sagitta narrow, simply pointed at apex, not toothed beneath. Malar space as 
long as antennal segment 4. (Eurasia) .... LAESOBOMBUS (p. 251) 

27 Sagitta at end somewhat hooked inwards and on outer side with two large teeth, 

mid-point beneath with bifid or trifid tooth. Stipes with wide, sharp-edged 
impressions. Inner edge of squama not emarginate, anterior inner corner some- 
times produced into a vertical lamella. Hind tibial fringe long or short (Hoi- 
arctic) SUBTERRANEOBOMBUS (p. 257) 

Sagitta curved or hooked outwards, or pointed, or in one species serrate . . 28 

28 Malar space normally a little longer than antennal segments 3 + 4. Sagitta at 

end curved outwards (except B. brevivillus Franklin, Fervidobombus), this end- 



SUBGENERIC DIVISIONS OF BOMBUS 265 

piece serrate or truncate, beneath with a central tooth. Stipes with rather well- 
defined inner impressions. Fringe of hind tibia short. ..... 29 

Malar space a little shorter than antennal segments 3 + 4. Sagitta with a small 
pointed external lobe, or pointed, or serrate. Stipes without any distinct inner 
impressions. Squama produced inside into an obliquely downwardly directed, 
acute plate or spike. (Eurasia) ......... 30 

29 Mandibles without a beard. Squama anteriorly on inside produced obliquely 

downwards and backwards into a twisted plate of which the end is hardly acute. 
Lacinia with apex on inside produced into a small lobe with a small tooth. Trun- 
cate end of sagitta not serrate. Volsellar region with no very conspicuous bristles. 

(Eurasia) RHODOBOMBUS (p. 258) 

Mandibles with a beard. Squama on inside usually produced into a vertical plate- 
like anterior lamella and a more dorsal wider rounded process (details vary con- 
siderably in different species). Truncate end of sagitta sometimes serrate (in 
B. brevivillus Franklin the sagitta is simply pointed). Lacinia with apex on 
inside produced into a small hook or angular process. Volsellar region with dense 
bristles. (N. and S. America) .... FERVIDOBOMBUS (p. 259) 

30 Lacinia long and broad, finger-shaped, centre of inner edge produced into a wide lobe 

defined at each end by a small tooth or else simple, inner and often ventral surface 
with dense, quite long pubescence. Sagitta beneath simple or emarginate but 
with no distinct tooth ........... 31 

Lacinia wide but not very long, apex more pointed, pubescence less dense and wide- 
spread, centre of inner edge produced into a spike or at least a narrow, truncate 
process. Sagitta variable but with a central tooth beneath. Fringe of hind 
tibia long THORACOBOMBUS (p. 255) 

31 Lacinia with centre of inner edge produced into a wide lobe defined at each end by 

a small tooth. Sagitta simply pointed, with no tooth or emargination beneath. 
Hind tibia with a long fringe .... MUCIDOBOMBUS (p. 256) 

Lacinia with inner edge straight or feebly concave, with no lobe or process. Sagitta 
at end hardly acute, with a slight outer, feebly serrate, flange, ventrally bi- 
emarginate but without a tooth. Fringe of hind tibia short 

EVERSMANNIBOMBUS (p. 252) 



FEMALES 

Apex of mandibles with six teeth. Mid basitarsus with posterior apical angle 
more or less clearly acute. Hind basitarsus proximally near ventral margin often 
with a number of bristles almost as long as the corbicular bristles. Sting-sheath 
with the outer thickenings rather narrow, of even width ; inner thickenings 
moderately widened below and gradually narrowed upwards, centre of wide part 
blackened, membrane between blackened on a lateral lobe projecting down to 
mid-point ............. 2 

Apex of mandibles with one small dorsal tooth and, when an incisura is developed, 
the ventral corner may also project to some extent. Hind basitarsus normally 
with no bristles as long as the corbicular bristles [except in Mendacibombus and 
Pressibombus] ......... -3 

Malar space strongly transverse, shorter than antennal segment 3. Ventral man- 
dibular tooth directed inwards. (Pyrenees and Norway to Himalayas and 
China) ALPIGENOBOMBUS (p. 221) 

Malar space about quadrate, longer than antennal segment 3. Ventral mandibular 

tooth directed downwards. (Himalayas s.l.) . . . NOBILIBOMBUS (p. 222) 

Mid basitarsus with posterior apical angle obtuse, more or less rounded. [Except 
for some Sibiricobombus , most of which have the ocelli separated by about two 



266 O. W. RICHARDS 

diameters from eyes ; they also differ from the species of Subterraneobombus 
which they often most resemble in having the whole surface of the auricle of the hind 
basitarsus densely hairy (Text-fig. 38). Bombus tanguticus Morawitz (Melanobombus) 
has the basitarsus produced but has a strong, bare subcircular boss on gastral 
tergite 6 such is not found in Odontobombus'] ....... 4 

- Mid basitarsus with posterior apical angle usually spinose, at least very distinctly 

acute. Malar space usually at least longer than broad, often distinctly elongate. 
Ocelli separated by three diameters from eyes. (Section Odontobombus) . . 22 

4 Outer surface of hind tibia densely reticulate, dull, some long bristles arising from 

disk down to near its mid-point. Malar space very elongate. Antennal segment 
3 fully three times as long as wide in centre. Outer thickenings of sting-sheath 
very narrow, inner ones very narrow, of even width, no blackened areas. (Pyren- 
ees to Himalayas and other asiatic mountains) . MENDACIBOMBUS (p. 229) 

- Outer surface of hind tibia less coarsely reticulate, rarely if ever so dull, long bristles 

confined to margins except very near the base or else the malar space is transverse. 
Antennal segment 3 nearly always shorter ....... 5 

5 Malar space very long, longer than antennal segments 2 + 3. Antennal segment 3 

about four times as long as broad. Lateral ocelli usually separated by not much 
more than two diameters from eyes. Clypeus usually with rather numerous, 
scattered, rather fine punctures, apical impressions with dense fine punctures. 
Mid basitarsus with posterior apical angle somewhat produced. (Some species, 
see couplet 3, are difficult to separate from some of Subterraneobombus, cf. couplet 
29.) Outer thickenings of sting-sheath rather narrow, sometimes a little widened 
above, inner thickenings narrow very little widened below ; membrane brown 
and convex in a small dorsal region and with a somewhat blackened lobe laterally 
near centre. (E. Europe to Asia, where mainly in mountains) 

SIBIRICOBOMBUS (p. 232) 

- Malar space clearly shorter or, if not, antennal segment 3 not so long (about two 

and a half times as long as broad). Mid basitarsus with posterior angle quite 
obtuse ............. 6 

6 Corbicular hairs unusually dense, tibial surface between them covered throughout 

with rather sparse but quite distinct, very short feathered hairs. Very large 
species ; wings usually coloured, dark or yellow brown. Malar space about 
quadrate. Mandibles with weak or no incisura, no sulcus obliquus. Clypeus 
generally little punctured, mid line dorsally with many, usually fine, punctures, 
apical depressions with coarse punctures and sometimes fine ones as well. Punc- 
tures of frons all relatively fine, unpunctured areas large. Outer thickenings 
of sting-sheath narrow but considerably widened above, inner thickenings 
widened from below to just above middle then narrowed again, edge consider- 
ably blackened, adjacent to this on each side a strong, shining, convex fold of 
membrane. (Himalayas to Formosa and Sumatra) RUFIPEDIBOMBUS (p. 238) 

- Corbicular hairs less dense, tibial surface between them without these short feathered 

hairs except in a few Melanobombus (couplet 8). (B. coccineus Friese has short, 
non-feathered hairs, but gastral tergite 6 has a deep furrow and the unpunctured 
area of the frons has a band of microscopic punctures.) ..... 7 

7 Frons with unpunctured areas very large ; most of the area for some distance in 

front of the ocelli unpunctured and a narrow band of punctures between the 
ocelli and eyes ; area immediately behind the ocelli also unpunctured. Large 
species with dark wings. Ocelli separated by three diameters from eyes. Malar 
space considerably longer than broad, clearly longer than antennal segments 2 + 3, 
antennal segment 3 two and a half times as long as wide at apex. Mandibles 
with no incisura. Clypeus with few punctures and these nearly all fine, apical 
impressions sparsely punctured. Labral lamella straight, wide, but not very 
clearly defined. Outer thickenings of sting-sheath of moderate, even width ; 



SUBGENERIC DIVISIONS OF BOMBUS 267 

inner thickenings moderately wide and gradually narrowing upwards, inner 
margin blackened for a considerable distance and adjacent membrane also 
blackened. (Kashmir to S. China) . . . ORIENTALIBOMBUS (p. 224) 

- Frons without this large unpunctured area, especially in front of the ocelli ; or 

else densely punctured right up to them posteriorly and ocelli closer to eyes . 8 

8 Sixth gastral tergite with a bare, convex, more or less rounded boss. Hind tibia 

with dorsal inner corner not or rarely somewhat produced. Hind basitarsus 
as a rule with unusually dense, short, feathery hairs. Wings rarely dark, tip 
not particularly darkened. Malar space about quadrate or, if distinctly longer, 
the ocelli are separated by two to two and a half diameters from eyes, or the 
species is very large and dark winged. Antennal segments 3 : 4 : 5 = 8 : 5^ : 6. 
Frons with quite numerous fine punctures along inner margin of eyes. Outer 
thickenings of sting-sheath generally narrow but much widened for a short 
distance dorsally, inner thickenings widened for a considerable distance then nar- 
rowed again dorsally, edge sometimes blackened and intervening membranous 
folds sometimes also blackened. (Europe to India, China and Formosa) 

MELANOBOMBUS (p. 236) 

- Sixth gastral tergite without a convex rounded boss. Hind tibia with dorsal distal 

inner corner more or less strongly angularly produced (except in Kallobombus 
and a few Pyrobombus B. pratorum (Linnaeus), B. atrocinctus Smith). Hind 
basitarsus rarely so densely haired ........ 9 

9 Whole discal surface of hind tibia with scattered but quite numerous short unbranched 

hairs. Ocelli separated by two and a half diameters from eyes. Frons with 
unpunctured area moderately large crossed by a wide transverse band of micro- 
scopic punctures, inner margin by eye with numerous very microscopic punctures 
and a few rather large ones. Malar space just transverse. Gastral tergite 6 with 
a raised boss (almost as in Melanobombus) but divided by a deep, well-defined 
furrow. Outer thickenings of sting-sheath narrow but considerably widened 
dorsally, inner thickenings narrow but moderately widened, on ventral quarter, 
not darkened ; membrane dark and rather convex dorsally. (Peru) 

COCCINEOBOMBUS (p. 243) 

- Hind tibia at least with a considerable distal discal area bare. Frons without a 

band of microscopic punctures across the unpunctured area. Gastral tergite 

6 never with such a deep, well-defined furrow . . . . . . . 10 

10 Malar space distinctly transverse. Either a definite band of close punctures along 

inner margin of eyes or ocelli separated by about two diameters from eyes, or 

both .............. ii 

Malar space elongate, quadrate or just transverse. In the members of Pyrobombus 

with a distinctly transverse malar space, there are no fine punctures along the inner 
eye-margin and the ocelli are separated by three diameters from eye ; antennal 
segment 3 clearly shorter than 4 + 5.. . . . . . . . 16 

11 Ocelli lying just in front of postocular line, separated by three diameters from 

eyes and corbicular surface of hind tibia entirely bare and shining. Mandibles with 
a strong incisura, sulcus obliquus moderately distinct. Clypeus strongly punc- 
tured on almost its whole surface. Outer thickenings of sting-sheath narrow, 
widened dorsally for a short distance only ; inner thickenings relatively wide, 
twice emarginate, the processes so formed more or less bent inwards, membrane 
undifferentiated. (Eurasia and N. America to Mexico) . BOMBUS s.s. (p. 226) 

Either ocelli not separated by more than two diameters from eyes or the proximal 

half or third of corbicular surface bristly and the whole surface more or less 
strongly reticulate ; or both these characters present. Mandibles with no or 
with a weak incisura, sulcus obliquus strong (except in Crotchiibombus Franklin). 
Clypeus sometimes with some sparse large punctures but if punctures are close 
they are small. ............ 12 



268 O. W. RICHARDS 

1 2 Hind tibia with its corbicular surface bare. Clypeus swollen with fine and rather close 

punctures. Ocelli separated by two diameters or less from eyes, well in front 

of postocular line. (U.S.A. and C. America) . . . . . . . 13 

Hind tibia with proximal half or third of its corbicular surface bristly. (C. and 

S. America) ............. 15 

13 Frons with rather strong, close, punctures along inner margin of eyes. Clypeus 

elongate, impressions feeble, finely and closely punctured. Labral furrow wide 
and deep, nearly as wide as length of antennal segment 3. Mandibles with a 
strong sulcus obliquus. Outer thickenings of sting-sheath moderately wide 
both above and below, inner thickenings rather strongly widened on lower third 
and evenly narrowing and running to top at a constant width ; a little darkened 
where it narrows. Membrane dorsally projecting and produced down each side 
into an elongate-ovate, blackened lobe. (N. America) SEPARATOBOMBUS (p. 231) 

Frons rather sparsely punctured all round the ocelli except for a densely punctured 

area immediately behind them ; unpunctured areas large and ill-defined, without 
a band of fine punctures near the eye though in Crotchiibombus a band of very 
fine punctures may be seen set rather more discally. Clypeus short, impressions 
with coarse punctures. Labral furrow deep and narrower. (Mandibles with a 
weaker sulcus obliquus in Crotchiibombus.) . . . . . . . 14 

14 Malar space clearly shorter than antennal segment 3 which is shorter than twice 

the length of 4. Clypeus more coarsely though shallowly punctured, impressions 
weaker. Labral tubercles more convex but less angular, furrow deep, rather wider 
than length of antennal segment 3. Corbicular hairs shorter than half tibial 
width and dense. Outer thickenings of sting-sheath narrow and of about constant 
width, inner thickenings considerably widened and blackened on lower third, 
above this very narrow ; membrane undifferentiated. (N. America) 

FRATERNOBOMBUS (p. 227) 

Malar space clearly longer than antennal segment 3 which is nearly as long as twice 

the length of 4. Clypeus more finely punctured, impressions stronger. Labral 
tubercles less raised but more angular at inner end, furrow deeper and much 
narrower than the length of antennal segment 3. Corbicular hairs mostly longer 
than half the tibial width and less dense. Outer thickenings of sting-sheath 
rather wide but narrowed in centre, inner thickenings widened for a short distance 
ventrally, then rather suddenly narrowed and of constant width to the top, con- 
siderably blackened on lower half ; membrane with a blackened convex area 
close to the narrowing of the inner thickenings. (N. and C. America) 

CROTCHIIBOMBUS (p. 231) 

15 Frons rather closely punctured, with large but well-defined unpunctured areas and 

a band of fine punctures along inner margin of eyes. Ocelli separated by three 
diameters from eyes and lying a little in front of postocular line. Clypeus with 
numerous punctures, mostly rather large, swollen with ventral third flattened. 
Labral furrow narrow. Hind basitarsus not unusually bristly. Outer thickenings 
of sting-sheath very wide, inner thickenings gradually widening from below but 
soon ending in a right-angled truncation, above this widening again in a regular 
curve, inner margin somewhat blackened, membrane forming two somewhat 
pyriform, convex, deep brown lobes. (C. and western S. America) 

RUBICUNDOBOMBUS (p. 241) 

Frons with large unpunctured or very sparsely punctured areas in front of and 

around the ocelli, no specially defined unpunctured areas, no band of punctures 
along inner margin of eyes. Ocelli separated by two diameters from eyes and 
lying well in front of postocular line. Clypeus strongly swollen, sometimes 
somewhat flattened ventrally, little or moderately punctured, apical impressions 
feeble. Labral furrow wide or very wide. Hind basitarsus with bristles on its 
outer surface longer and more numerous than usual. Outer thickenings of 



SUBGENERIC DIVISIONS OF BOMBUS 269 

sting-sheath of constant width, moderately broad, inner thickenings considerably 
widened below and ending in a finger-shaped blackened process, a somewhat 
similar process projects from above almost to meet it, margin of small area 
enclosed between these processes also sometimes blackened ; intervening mem- 
brane sometimes raised into a horseshoe-shaped fold. (C. and S. America) 

ROBUSTOBOMBUS (p. 240) 

1 6 Malar space much or at least distinctly longer than broad, at least as long as antennal 

segments 2 + 3, often as long as 3 + 4- Clypeus moderately long and swollen, 
generally but not very coarsely punctured, apical impressions strong, strongly 
and more or less closely punctured. Antennal segments 3:4:5 = 9:6:7. 
Frons rather finely punctured, especially behind the level of the ocelli, unpunc- 
tured areas ill-defined, a rather broad band of fine sculpture along inner margin 
of eyes. Ocelli separated by three diameters from eyes, just in front of or almost 
on the postocular line. Inner thickenings of sting-sheath strongly widened below, 
two isolated blackened spots in the membrane between them. (Alps, N. Scan- 
dinavia, Arctic Eurasia, Arctic America, Rockies, Sierras, to Colorado and 
California) ALPINOBOMBUS (p. 239) 

Malar space transverse, quadrate or a little longer than broad, never more than a 

little longer than antennal segment 3.. . . . . . . . 17 

17 Mandibles with no incisura but with ventral apical angle produced into a small 

process, sulcus obliquus rather strong. Ocelli separated by two and a half dia- 
meters from eyes, rather distinctly in front of postocular line. Labral tubercles 
convex, angled at inner end, furrow narrow, narrower than length of antennal 
segment 3. Frons quite closely punctured, unpunctured areas large but well- 
defined, a wide band of fine punctures along the eyes. Outer thickenings of 
sting-sheath moderately wide, inner thickenings gradually but not very strongly 
widened from below, ending in a rounded blackened knob, above this slightly 
widened again and sending down an ill-defined blackened process towards the 
knob ; lower part and adjacent membrane somewhat blackened, membrane 
otherwise undifferentiated. (C. and S. America) . FUNEBRIBOMBUS (p. 244) 

Mandibles sometimes with an incisura but not with the ventral apical angle produced 

into a short process ........... 18 

1 8 Labral tubercles little raised and much rounded, furrow shallow and ill-defined. 

Ocelli separated by two and a half diameters or less from eyes, well in front of 
postocular line. Mandibles with no incisura and a weak sulcus obliquus. An- 
tennal segment 3 as long as or hardly longer than 4 + 5 J 9 

Labral tubercles more or less raised and flattened, inner end more or less angled, 

furrow deeper and well-defined. Ocelli more widely separated from eyes (except 
some Cullumanobombus) . Antennal segment 3 clearly shorter than 4 + 5. 
Malar space quadrate or more or less transverse. ...... 20 

19 Malar space a little longer than broad and a little longer than antennal segment 3. 

Clypeus with a wide flattened disk, closely and finely punctured, especially on 
lower third, impressions ill-defined but closely punctured. Frons mostly closely 
and rather finely punctured, unpunctured areas well-defined, a wide band of 
close fine punctures along inner margin of eyes. Outer thickenings of sting- 
sheath not broad but a little more so above than below, inner thickenings 
very narrow, membrane convex dorsally and sending down to centre two large 
blackened lobes on each side. (C. Europe) . . CONFUSIBOMBUS (p. 228) 

Malar space about quadrate, about as long as antennal segment 3. Clypeus long, 

strongly swollen, closely and finely punctured, impressions very weak and not 
more punctured. Frons moderately strongly and closely punctured, unpunctured 
areas large and ill-defined, a narrow band of rather fine sculpture along inner margin 
of eyes. Outer thickenings of sting-sheath a little wider above than below, inner 
thickenings rather narrow and of almost constant width, edge upturned, especially 



ayo O. W. RICHARDS 

below, membrane undifferentiated except for some lateral blackening. (N. 
America) - . ' . . BOMBIAS (p. 230) 

20 Ocelli separated by somewhat or distinctly less than three diameters from eyes, 

well in front of postocular line. Hind basitarsus rather less pubescent than 
usual. Frons closely and rather finely punctured, unpunctured areas small and 
well-defined, a wide band of fine sculpture along inner margin of eyes. Mandibles 
with no incisura and a rather strong sulcus obliquus. Outer thickenings of sting- 
sheath considerably widened above, inner thickenings rather strongly widened 
and a little blackened below, two small blackened areas in membrane opposite 
centre of widened part. (Eurasia, N. and C. America) 

CULLUMANOBOMBUS (p. 233) 

Ocelli separated by fully three diameters from eyes, hardly in front of postocular 

line .............. 21 

21 Clypeus rather strongly and evenly punctured, impressions strong but not more 

closely punctured. Frons rather closely punctured, unpunctured areas small and 
well-defined, a wide band of fine sculpture along inner margin of eyes. Man- 
dibles with no incisura or sulcus obliquus. Hind basitarsus with sparse pubes- 
cence and no long bristles. Outer thickenings of sting-sheath moderately wide, 
especially above, inner thickenings rather strongly widened upwards to above 
middle then suddenly narrowed, edge of wide part a little blackened, a slight 
dorsal fold but no black spots in the membrane. (Europe) KALLOBOMBUS (p. 225) 

Clypeus, except impressions, largely unpunctured (more strongly and closely in 

B. lapponicus (Fab.). Frons mostly rather sparsely punctured, unpunctured 
areas ill-defined, no fine punctures along inner margin of eyes, the margin being 
largely shining. Mandibles with a well-marked incisura but no sulcus obliquus. 
No long bristles on hind basitarsus which is not usually very densely haired. 
Outer thickenings of sting-sheath narrow but considerably widened for a short 
distance dorsally, inner thickenings narrow and scarcely widened or blackened, 
two large blackened spots in membrane (at least of copulated $) (Europe, Asia, 
including Malayan Archipelago, N. and C. America) PYROBOMBUS (p. 234) 

Clypeus with fairly numerous scattered punctures, mostly small but some large. 

Frons not closely nor coarsely punctured, unpunctured areas not large, ill-defined, 
no fine punctures along inner margin of eyes, the margin being largely shining. 
Mandibles with feeble incisura and no sulcus obliquus. Hind basitarsus densely 
pubescent, lower edge for its whole length and disk in part with long bristles. 
Outer thickenings of sting-sheath not wide but wider above than below, inner 
thickenings very wide, widest just above middle and gradually narrowing above 
and below, considerably blackened, especially near middle ; membrane little 
differentiated except for a darkened dorsal patch. (Himalayas) 

PRESSIBOMBUS (Frison) (p. 237) 

22 Malar space twice as long as antennal segment 3 . . . . . .23 

Malar space less than i -30 times as long as antennal segment 3 . . . .25 

Malar space i -40- 1 -80 times as long as antennal segment 3 ... .27 

23 Antennal segment 3 a little shorter than 4 + 5 (io : n). Mid line of clypeus 

dorsally with a slight furrow or line of close punctures. Furrow between labral 
tubercles narrower. Mid basitarsus acutely spinosely produced. Outer thick- 
enings of sting-sheath rather wide, of fairly constant width, inner thickenings 
considerably widened below to above the mid point, this part with a blackened 
edge and ending in a small blackened projection, above this emarginate and then 
with a rounded projecting lobe of which the base is blackened, membrane between 
thrown into two longitudinal folds. (Eurasia north and west of Himalayas) 

MEGABOMBUS (p. 246) 

Antennal segment 3 clearly shorter than 4 + 5 (at most n-J- : 13). Furrow between 

labral tubercles wider. Mid basitarsus with the production wider, hardly spinose. 24 



SUBGENERIC DIVISIONS OF BOMBUS 271 

24 Inner dorsal angle of hind tibia not or hardly produced apically. Much of clypeus 

rather strongly and closely punctured, mid line on dorsal third with several rows 
of fine punctures (cf. couplet 36). A few species of FERVIDOBOMBUS (p. 259) 
Inner dorsal angle of hind tibia pointed apically though the point is short and broad. 
Clypeus finely or little punctured, no rows of punctures on mid line dorsally 
(cf. couplet 35). . . . Some species of SUBTERRANEOBOMBUS (p. 257) 

25 Frons with large unpunctured areas separated from eyes by a narrow band of close, 

fine punctures. Malar space unpunctured. Mid basitarsus spinosely produced. 
Outer thickenings of sting-sheath not wide, a little wider above, inner thickenings 
moderately widened over most of dorsal half, a little blackened, membrane not 
differentiated. (Eurasia, north and west of the Himalayas) 

LAESOBOMBUS (p. 251) 

Frons with small unpunctured areas, separated from eyes by a wide band of fine 

punctures ............. 26 

26 Mid basitarsus broadly produced. Band of fine punctures along inner edge of 

eye not spreading over the unpunctured area of frons. Labral lamella not 
prominent. Malar space with many fine punctures. Outer thickenings of 
sting-sheath rather narrow but somewhat wider dorsally, inner thickenings 
narrow but somewhat widened on central two thirds and the margin at the 
centre somewhat blackened, adjacent membrane also somewhat blackened but 
otherwise undifferentiated. (E. Europe and W. Asia but not in Himalayas) 

EVERSMANNIBOMBUS (p. 252) 

Mid basitarsus spinosely produced. Band of fine punctures along inner edge of 

eye spreading halfway across unpunctured area of frons. Labral lamella con- 
siderably thickened. Malar space unpunctured. Outer thickenings of sting- 
sheath wide above, narrow below, inner thickenings moderately wide on lower 
half, gradually but much narrowed above, membrane with two indefinite dorsal 
lobes. (E. Mongolia and Ussuri distr.) .... EXILOBOMBUS (p. 253) 

27 Gastral tergites with very close coarse punctures except tergite 6 which has coarse 

granules. Mid basitarsus acutely spinosely produced. Hind tibia with corbi- 
cular surface strongly reticulate. Length malar space : antennal segment 3 = 
i -60. Frons with rather small and well-defined unpunctured areas with a wide 
band of close fine punctures along inner margins of eyes. Outer thickenings of 
sting-sheath not wide except quite dorsally, inner thickenings very wide and 
bent inwards, ending rather suddenly dorsally and here somewhat blackened 
though generally pale brown ; membrane not generally differentiated but with 
a convex central fold at dorsal end. (Asia, especially north-eastern) 

TRICORNIBOMBUS (p. 249) 
Gastral tergites without such close, coarse punctures ...... 28 

28 Frons with no band of close fine punctures along the inner margin of eyes, unpunc- 

tured area large and ill-defined. Clypeus closely and coarsely punctured. 
Length malar space : antennal segment 3 = 1-50. Mid basitarsus acutely 
spinosely produced. Hind tibia with corbicular surface feebly reticulate. Outer 
thickenings of sting-sheath rather wide, even wider dorsally, inner thickenings 
rather narrow with a moderately wide section near centre, above this blackened 
and the black part with a small downward projection into the membrane from 
about the middle, intervening membrane thrown into two big folds. (Malaysia, 
Philippines, Indonesia) SENEXIBOMBUS (p. 248) 

Frons with a band of close fine punctures along the inner margin of eyes. . . 29 

29 Malar space very long and third antennal segment about four times as long as broad. 

Auricle of hind basitarsus with dense brown pile even on surface not apposed to 
hind tibia (Text-fig. 38. Inner thickenings of sting-sheath very little widened even 
below and membrane only with a narrow blackened lateral lobe. 

Some species of SIBIRICOBOMBUS (see couplet 5). 



272 



O. W. RICHARDS 



Malar space and third antennal segment shorter. A few species of Subterraneobombus 

(couplet 34) are not very different but in them the auricle (fig. 39) has dense 
pile only on the surface apposed to the end of the hind tibia and the inner thicken- 
ings of the sting-sheath project strongly, are heavily sclerotized, and the inter- 
vening membrane has two very large lateral folds. . . . . . . 30 

30 Mid basitarsus acutely spinosely produced . . . . . . . .31 

Mid basitarsus acute but not spinose ........ 35 

31 Malar space longer, about as long as antennal segments 2 + 3 + 4. Labral furrow 

narrower and deeper. Inner dorsal angle at apex of hind tibia little produced . 32 

Malar space not longer than antennal segments 3 + 4 ; antennal segment 5 only 

a little longer than 4. Hind basitarsus quite strongly produced 33 

32 Antennal segment 5 clearly longer than 4 which is transverse rather than quadrate 

and shorter than in any other group of Odontobombus. Hind basitarsus little pro- 
duced apically. Outer thickenings of sting-sheath moderately wide, especially 
above, inner thickenings strongly almost angularly widened, widest well below 
middle, not blackened ; membrane hardened and darkened dorsally. (Asia) 

DIVERSOBOMBUS (p. 247) 

Antennal segment 5 very little longer than 4 which is at least quadrate. Hind 

basitarsus distinctly produced apically. Outer thickenings of sting-sheath some- 
what wider dorsally than ventrally, inner thickenings widening very gradually 
upwards, widest near the top then rather suddenly narrowing, a slight blackening 
of the edge near the centre ; two small central folds at the top of the membrane. 
(I have not seen a male of this group and the genitalia do not seem to have been 
illustrated.) (Asia) .... ADVENTORIBOMBUS (p. 254) 

33 A large unpunctured area in front of median ocellus, lateral unpunctured areas large 

but well-defined, band of fine sculpture along inner margins of eye narrow. 
Antennal segment 3 just longer than 4 + 5. Apical impressions of clypeus small 
and narrow with close larger punctures, mostly in rows. Inner dorsal angle at 
apex of hind tibia not or hardly produced. Outer thickenings of sting-sheath 
rather narrow, only a little wider dorsally, inner thickenings moderately and 




FIGS. 38-39. Auricle of right hind basitarsus of, 38, Bombus miniatocaudatus Vogt 

39, B. fragrans Pallas. 



SUBGENERIC DIVISIONS OF BOMBUS 273 

evenly widened over most of their length, thickenings rather dark ; membrane 
thrown into two small dorsal brown folds. (Eurasia north and west of Hima- 
layas), (cf. also couplet 37) .... RHODOBOMBUS (p. 258) 

Area in front of median ocellus coarsely but not closely punctured, lateral un- 

punctured areas of moderate size, fairly well-defined, band of fine sculpture along 
inner margin of eyes wide or rather wide, antennal segment 3 shorter than or as 
long as 4 + 5. Inner dorsal apical angle of hind tibia more acutely produced. 34 

34 Malar space as long as antennal segments 3 + 4- Apical impressions of clypeus 

rather strong with close, moderately coarse punctures. Labral tubercles flat- 
tened, furrow shallow. Outer thickenings of sting-sheath moderately wide, 
especially above, inner thickenings moderately or rather strongly broadened to 
the middle or rather higher, the top of the broad part projecting as a slight spur, 
margin below the broadest part a little blackened, membrane undifferentiated. 
(Eurasia, north and west of Himalayas) . . . THORACOBOMBUS (p. 255) 

Malar space not quite as long as antennal segments 2 + 3- Apical impressions of 

clypeus weak with a narrow deeper strip with close moderately coarse punctures. 
Labral tubercles somewhat raised and angular at the inner end, furrow moderately 
deep. Outer thickenings of sting-sheath rather narrow even dorsally, inner 
thickenings gradually widening upwards for almost whole length, widest just 
before the top and blackened for a short distance below this ; membrane undif- 
ferentiated. (Eurasia, west and north of Himalayas) MUCIDOBOMBUS (p. 256) 

35 Species often large or very large, clypeus swollen with no furrow or lines of punctures 

on dorsal third of mid line, general surface often but not in all species considerably 
punctured. Malar space variable, ratio of its length to that of the third antennal 
segment from 1-43 (B. fragrans Pall.) to over 2-10 (B. difficillimus Skor.). Hind 
tibia with inner dorsal apical angle sharp though production rather wide. Outer 
thickenings of sting-sheath not very wide and of even width, inner thickenings 
rather strongly widened in a regular curve, widest at about the middle, a sharp 
black patch just above this ; membrane dorsally on each side thrown into a 
strong dark fold which touches the inner thickening, about meets its fellow and 
extends down to mid point. (See also couplet 24.) (Eurasia, N. America) 

SUBTERRANEOBOMBUS (p. 257) 

Clypeus with a slight furrow or distinct lines of punctures on mid line of dorsal 

third. Malar space of medium length or, rarely, rather long. Hind tibia with 
inner dorsal apical angle not or scarcely produced ...... 36 

36 Clypeus with widespread but sparse fine punctures. Mid basitarsus more spinosely 

produced. Frons less punctured with a larger area in front of and at sides of 
ocelli unpunctured. (See couplet 32.) . . . RHODOBOMBUS (p. 258) 

Clypeus generally with closer and coarser punctures. Mid basitarsus often not 

very distinctly produced. Frons more punctured, with unpunctured areas 
smaller. Species often large or very large. Outer thickenings of sting-sheath 
much wider above than below, inner thickenings strongly widened below, then 
narrowing from about mid point where there is a large blackened area ; membrane 
dorsally thrown into a convex central fold. (N. and S. America) 

FERVIDOBOMBUS (p. 259) 

REFERENCES 

BALL, F. J. 1914. Les bourdons de la Belgique. Annls Soc. ent. Belg. 58 : 77-108, i plate. 

BISCHOFF, H. 1936. Die Schwedisch-chinesische wissenschaftliche Expedition nach den 
nordwestlichen Provinzen Chinas unter Leitung von Dr. Sven Hedin und Prof. Sxi Ping- 
Chang. 56. Hymenopteren. 10. Bombinae. Ark. Zool. 27A No. 38 : 1-27. 

COCKERELL, T. D. A. 1906. Descriptions and records of bees. XII. Ann. Mag. nat. Hist. 
(i) 18 : 69-75- 



274 O. W. RICHARDS 

CRESSON, E. T. 1863. List of the North American species of Bombus and Apathus. Proc. 

ent. Soc. Philad. 2 : 85-115. 
1874. Descriptions of new Hymenoptera. Trans. Am. ent. Soc. 5 : 99-102. 

1878. Descriptions of new species of North American bees. Proc. Acad. nat. Sci. Philad. 
1878 : 181-221. 

CUMBER, R. A. 1949. The biology of Humble-bees, with special reference to the production 

of the worker caste. Trans. R. ent. Soc. Lond. 100 : 1-45, 10 figs. 
DALLA TORRE, K. W. von. 1880. Unsere Hummel- (Bombus} Arten. Naturhistoriker 2 : 30, 

40-41. 

1882. Bemerkungen zur Gattung Bombus Latr., II. Ber. naturw.-med. Ver. Innsbruck 
12 : 14-31. 

1896. Catalogus Hymenopterorum. 10. Apidae (Anthophila). Leipzig. 
DEGEER, C. 1773. Memoires pour servir a I'histoire des Insectes. Stockholm. 
FABRICIUS, J. C. 1777. Genera Insectorum. Chilonii. 
1781. Species Insectorum. 1. Hamburgi et Kilonii. 

1787. Mantissa Insectorum. 1. Hafniae. 

1793- Entomologia systematica emendata et aucta. 2. Hafniae. 

1798. Entomologiae Systematicae, Supplementum. Hafniae. 

FRANKLIN, H. J. 1912-1913. The Bombidae of the New World. Trans. Am. ent. Soc. 38 
177-486 ; 39 : 73-200, 22 plates. 

1954. The evolution and distribution of American Bumble-bee kinds. Trans. Am. ent. 
Soc. 80 : 43-51. 

FRIESE, H. 1903. Neue Bombus-Arten aus der neotropischen Region (Hym.). Z. Syst. Hym- 
enopt. Dipteral. 3 : 253-255. 

1905. Neue oder wenig bekannte Hummeln des Russischen Reiches. (Hymenoptera). 
Ezheg. zool. Muz. 9 (1904) : 507-525. 

1911. Neue Varietaten von Bombus. III. Dt. ent. Z. 1911 : 571-572. 
1918. tiber Hummelform aus dem Himalaja. Dt. ent. Z. 1918 : 81-86. 



FRISON, T. H. 1925. Contribution to the classification of the Bremidae (Bumble-bees) of 
Central and South America. Trans. Am. ent. Soc. 51 : 137-165, 2 plates. 

1927. A contribution to our knowledge of the relationships of the Bremidae of America 
north of Mexico (Hymenoptera). Trans. Am. ent. Soc. 53 : 51-78, 2 plates. 

1930. The Bumblebees of Java, Sumatra and Borneo. Treubia 12 : 1-22, 8 figs. 

1933. Records and descriptions of Bremus and Psithyrus from India (Bremidae : Hymen- 
optera). Rec. Indian Mus. 35 : 331-342, 5 figs. 

1934- Records and descriptions of Bremus and Psithyrus from Formosa and the Asiatic 

mainland. Trans, nat. Hist. Soc. Formosa 24 : 150-185, 8 figs. 
I935- Records and descriptions of Bremus from Asia (Bremidae : Hymenoptera). Rec. 

Indian Mus. 37 : 339-363, 7 figs. 
GERST ACKER, A. 1869. Beitrage zur naheren Kenntniss einiger Bienen-Gattungen. Stettin. 

ent. Ztg 30 : 315-367- 
GRIBODO, G. 1891. Contribuzioni Imenotterologiche. Sopra alcune species nuove o poco 

conosciute di Imenotteri Antofili. (Generi Ctenoplectra, Xylocopa, Centris, Psithyrus, 

Trigona e Bombus.) Boll. Soc. ent. Ital. 23 : 102-119. 
HAZELTINE, W. E. & Chandler, L. 1964. A preliminary atlas for the identification of female 

Bumble bees (Hymenoptera, Apidae). /. Kans. ent. Soc. 37 : 77-87, 6 plates. 
HEMMING, F. 1939 [Ed.]. Opinion 135. The suppression of the so-called " Erlangen List " 

of 1801. Opin. Decl. int. Commn Zool. Nom. 2 : 912. 
HOBBS, G. A. 1964. Phylogeny of Bumble bees based on brood-rearing behaviour. Can. 

Ent. 96 : 115-116. 
JURINE, L. 1801. In [Panzer, G. W. F.] Nachricht von einem neuen entomologischen Werke, 

des Hrn. Prof. Jurine in Geneve. Intelligenzblatt der Litteratur-Zeitung 1 : 160-165. 

Erlangen. 
KIRBY, W. 1802. Monographia Apum Angliae. Ipswich. 



SUBGENERIC DIVISIONS OF BOMBUS 275 

KRUGER, E. 1917. Zur Systematik der mitteleuropaischen Hummeln (Hym.). Ent. Mitt. 

6 : 55-66, i fig. 
1920. Beitrage zur Systematik und Morphologic der mitteleuropaischen Hummeln. 

Zoo/. Jb. Abt. Syst. 42 : 289-464, 7 plates, figs A-H. 
1951. Phanoanalytische Studien an einigen Arten der Untergattung Terrestribombus 

0. Vogt. (Hymen. Bomb.), i. Teil. Tijdschr. Ent. 93 (1950) : 141-197, 22 figs. 
LATREILLE, P. A. i8o2a. [An. X] [April]. Histoire naturelle des Fourmis. Paris. 

i8o2b. [An. X] [May-Sept.]. Histoire naturelle, generate et particuliere des Crustaces et 

des Insectes. III. Paris. 
LEPELETIER DE SAINT-FARGEAU, A. 1836. Histoire naturelle des Insectes. Hymenopteres. 

1. Paris. 

LINNAEUS, C. VON. 1758. Sy sterna Naturae. 1 Regnum Animale. Holmiae. 

1761. Fauna svecica. Ed. 2. Stockholm. 

MEDLER, J. T. ig62a. Morphometric studies on Bumble bees. Ann. ent. Soc. Am., 55 : 

212-218, i fig. 
ig62b. Morphometric analyses of bumblebee mouthparts. Verh. XI internal. Kongr. 

Ent., Wien 2 : 517-521, i fig. 
I962C. Measurements of the labium and radial cell of Psithyrus (Hymenoptera Apidae). 

Can. Ent. 94 : 444-447, i fig. 
MICHENER, C. D. 1944. Comparative external morphology, phylogeny, and a classification 

of the bees (Hymenoptera). Bull. Am. Mus. nat. Hist. 82 art. 6 : 157-326, 246 figs. 
MILLIRON, J. E. 1961. Revised classification of the bumblebees A synopsis (Hymenoptera : 

Apidae). /. Kans. ent. Soc. 34 : 49-61. 
MITCHELL, T. B. 1962. Bees of the eastern United States. 2. Tech. Bull. N. Carol, agric. 

Exp. Stn. 152 : 557 pp., i plate, 134 figs. 
MORAWITZ, F. 1875. In Fedchenko, Reise in Turkestan. 2 ($&). Hymenoptera Mellifera I. 

Apidae. Moscow. 

1883. Neue russisch-asiatische ow&MS-Arten. Trudy russk. ent. Obshch. 17 : 235-245. 

1886. Insecta in itinere Cl. N. Przewalskii in Asia centrali novissime lecta. I. Apidae. 

Trudy russk. ent. Obshch. 20 : 195-229. 
1890. Insecta a Cl. G. N. Potanin in China et in Mongolia novissime lecta. XIV. 

Hymenoptera Aculeata. II. Trudy russk. ent. Obshch. 24 : 349-3 8 5- 
MORICE, F. D. & DURRANT, J. H. 1915. The authorship and first publication of the " Juri- 

nean " genera of Hymenoptera, etc. Trans, ent. Soc. Lond. 1914 : 339-436. 
MOURE, J. S. & SAKAGAMI, S. F. 1962. As mamangabas sociais do Brasil (Bombus Latr.) 

(Hym. Apoidea). Stud. ent. N.S. 5 : 65-194, 19 figs, 5 maps. 
NEAVE, S. A. ed. 1940. Nomenclator zoologicus, 4 (Suppl.). London. 
NYLANDER, W. 1848. Adnotationes in expositionem monographicam Apum borealium. 

Notiser. Sdllsk. Fauna Fennica Fork. 1 : 165-282, i pi. 
PITTIONI, B. 1937. Eine Hummelausbeute aus dem Elburs-Gebirge (Iran). Konowia 16 

(1937-38): H3-I 2 9, 2 figs. 

- i939a. Die Hummeln und Schmarotzerhummeln der Balkan-Halbinsel. II. Spezialer 
Teil. Izv. tsarsk. prirodonauch. Inst. Sof. 12 : 49-115, 6 plates. 

- i939b. Tanguticobombus subg. nov. (Hymenopt., Apidae). Zoo/. Anz. 126 : 201-205. 
PLATH, O. E. 1934. Bumblebees and their ways, xvi + 201 pp. New York. 

QUILIS PEREZ, M. 1927. Los Apidos de Espafia. Genero Bombus Latr. Trabhs. Lab. Hist. 

nat. Valencia 16 : 1119, 10 plates. 
RADOSZKOWSKY, O. 1884. Revision des armures copulatrices des males du genre Bombus. 

Byull. mosk. Obshch. Ispyt. Prir. 59 : 51-92, 4_figs, 4 plates. 

1888. Etudes Hymenopterologiques. Trudy russk. ent. Obshch. 22 : 3!5-337. 4 plates. 

REINIG, W. F. 1930. Untersuchungen zur Kenntnis der Hummelfauna des Pamir-Hoch- 

landes. Z. Morph. Okol. Tiere 17 : 68-123, 20 figs. 
RICHARDS, O. W. 1927. The specific characters of British Humblebees (Hymenoptera). 

Trans, ent. Soc. Lond. 1927 : 233-268, 4 plates, 5 figs. 



276 O. W. RICHARDS 

RICHARDS, O. W. 1929. (April). A revision of the humble-bees allied to Bombus orientalis 
Smith, with thedescription of a new subgenus. Ann. Mag. nat. Hist. (10) 3 : 378-386, 
7 figs. 

1931. Some notes on the humble-bees allied to Bombus alpinus L. Troms0 Mus. Arsh., 

50 (1927) : 32 pp., 2 plates. 

1956. Hymenoptera. Introduction and keys to families. Handbk Ident. Br. Insects 6 
pt. i : 94 pp., 196 + 22 figs. 

ROBERTSON, C. 1903. Synopsis of Megachilidae and Bombinae. Trans. Am. ent. Soc. 29 : 

163-189. 
SANDHOUSE, G. A. 1943. The type species of the genera and subgenera of bees. Proc. U.S. 

nat. Mus. 92 : 519-619. 

SCHENCK, A. 1859. Die nassauischen Bienen. Jb. Ver. Naturk. Nassau 14 : 1-414. 
SCHRANK, F. DE P. 1781. Enumeratio Insectorum Austriae indigenorum. Augustae Vindeli- 

corum. 

SCHULZ, W. A. 1906. Spolia Hymenopterologica. Paderborn. 
SCOPOLI, J. A. 1763. Entomologia Carniolica. Vindobonae. 
SKORIKOV, A. S. 1909. New forms of Bombus (Hymenoptera, Bombidae). (Preliminary 

diagnoses). III. Russk. ent. Obozr. 9 : 409-413 (in Russian). 

1910. Revision der in der Sammlung des weil. Prof. E. A. Eversmann befindlichen 
Hummeln. Trudy russk. ent. Obshch. 39 (1909) : 570-584. 

1914. Les formes nouvelles des bourdons. Russk. ent. Obozr. 14 : 119-129 (in Russian). 

1922. Les Bourdons de la faune palearctique. Partie i. Biologic generale. Bull. Sta. 
re'g. Prot. Plantes, Petrograd 4 : 102-160, 5 figs., 15 maps. 

i933a. Zur Hummelfauna Japans und seiner Nachbarlandes. Mushi 6 : 53 : 65, 2 figs. 

!933b. Zur Fauna und Zoogeographie der Hummeln des Himalaya. C.R. Acad. Sci. 
U.R.S.S., 2 : 243-248 (in Russian and German). 

1937. Die gronlandischen Hummeln im Aspekte der Zirkumpolarfauna. Ent. Meddr. 20 : 
37-64. 

1938. Zoogeographische Gesetzmassigkeiten der Hummelfauna im Kaukasus, Iran und 
Anatolien. (Hymenoptera, Bombinae). (In Russian). Ent. Obozr. 27, 3-4 : 145-151. 

SLADEN, F. W. L. 1912. The Humble-bee, its life-history and how to domesticate it. xiii -f 283. 

London. 
SMITH, F. 1852. Descriptions of some Hymenopterous insects from northern India. Trans. 

ent. Soc. Lond. (2) 2 : 4548. 

1854. Catalogue of Hymenopterous insects in the collection of the British Museiim. Part 2. 
Apidae. London. 

1869. Descriptions of Hymenoptera from Japan. Entomologist 4 : 205-208. 

1872. Notes on the habits of some Hymenopterous insects from the North-West provinces 
of India. By C. Home. With an appendix containing descriptions of some new species of 
Apidae and Vespidae collected by Mr. Home. By F. Smith. Trans, zool. Soc. Lond. 7 : 
161-196, 4 plates. 

SNODGRASS, R. E. 1941. The male genitalia of the Hymenoptera. Smithson. misc. Collns 

199 (14) : 86 pp., 33 plates, 6 figs. 

SPINOLA, M. 1805. Faunae Liguriae fragmenta. Genuae. [Not seen.] 
TKALCU, B. 1963. Contribution a 1'etude des Bourdons du Japon. Bull. Soc. ent. Mulhouse 

1962 : 81-100, 43 figs. 
VOGT, O. 1909. Studien iiber das Artproblem. Mitt, i : Uber das Variieren der Hummeln. 

Sber. Ges. naturf. Freunde Berl. 1909 : 28-84, J plate. 

1911. Studien iiber das Artproblem. Mitt. 2, Teil 2. Sber. Ges. naturf. Freunde Berl. 
1911 : 31-74 

VOLLENHOVEN, S. C. S. VAN. 1873. Description d'un Bombus nouveau de 1'ile de Sumatra. 

Tijdschr. Ent. 16 : 229-230, i plate. 
ZANDER, E. 1900. Beitrage zur Morphologic der mannlichen Geschlechtsanhange der Hymen- 

opteren. Z. wiss. Zool. 67 : 461-489, i plate. 




A LIST OF SUPPLEMENTS 
TO THE ENTOMOLOGICAL SERIES 

OF THE BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 



1. MASNER, L. The types of Proctotrupoidea (Hymenoptera) in the British 
Museum (Natural History) and in the Hope Department of Entomology, Oxford. 
Pp. 143. February, 1965. 5. 

2. NIXON, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera : 
Braconidae). Pp.284; 348 Text-figures. August, 1965. 6. 

3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177 ; 
18 plates, 270 Text-figures. August, 1965. 4 45. 

4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172 ; 500 Text-figures. October, 

1965- ^355. 

5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156 ; 
475 Text-figures. November, 1965. 2 155. 

6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae & Drosophilidae. 
Pp. 129 ; 328 Text-figures. 3. 

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

33s. 

8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the 

world species of Cleora (Lepidoptera : Geometridae) . Pp. 119; 14 plates, 146 
Text-figures, 9 maps. February, 1967. 3 los. 

9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera : Rhopalocera). Pp. 509. August, 1967. 8 IDS. 

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rhopa- 
locera). Pp.322; Coloured frontispiece, 348 text-figures. August, 1967. 8. 

11. MOUND, L. A. A review of R. S. Bagnall's Thysanoptera Collection's Pp. 184; 
82 Text-figures. May, 1968. 4. 



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






A LIST OF THE TYPE-SPECIMENS 

OF LIBELLULIDAE AND 

CORDULIIDAE (ODONATA) IN 

THE BRITISH MUSEUM 

(NATURAL HISTORY) 



D. E. KIMMINS 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 22 No. 6 

LONDON: 1968 



^ 8H 3 

A LIST OF THE TYPE -SPECIMENS OF 

LIBELLULIDAE AND CORDULIIDAE 

(ODONATA) IN THE BRITISH MUSEUM 

(NATURAL HISTORY) 



BY 

D. E. KIMMINS 

British Museum (Natural History) 



Pp. 277-305 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 22 No. 6 

LONDON: 1968 



THE BULLETIN OF THE BRITISH MUSEUM 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

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

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

This paper is Vol. 22, No. 6 of the Entomological 
series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



World List abbreviation : 
Butt. Br. Mus. nat. Hist. (Ent.). 



Trustees of the British Museum (Natural History) 1968 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 20 September, 1968 Price Twelve Shillings. 



A LIST OF THE TYPE-SPECIMENS OF 

LIBELLULIDAE AND CORDULIIDAE 

(ODONATA) IN THE BRITISH MUSEUM 

(NATURAL HISTORY) 

By D. E. KIMMINS 

SYNOPSIS 

A list of the Libellulid and Corduliid type-specimens in the British Museum (Nat. Hist.) has 
been prepared. Three hundred and forty-three taxa are dealt with and lectotypes are designated 
for ninety-seven of these taxa. 

INTRODUCTION 

IN 1966, following the bequest to the British Museum (Nat. Hist.) of the Fraser 
collection of Odonata, the author published a list of Eraser's Odonata types in the 
BM(NH) collections. This produced valuable information and the present work 
offers a similar list of the type-specimens of Libellulidae and Corduliidae in our 
collection. The publication of such lists is advocated by the International Commis- 
sion on Zoological Nomenclature (Rec. 720(4)) and the preparation of this list has 
proved both useful and interesting. Certain discrepancies in the labelling and 
marking of some of the earlier types have come to light and the opportunity has 
been taken to designate lectotypes in ninety-three cases in which the descrip- 
tions were based upon a series of specimens and a type or holotype was not 
specified by the original author in print. The word LECTOTYPE, in capital letters, 
indicates a designation of lectotype in this paper. 

This list includes not only Eraser's types, dealt with by Kimmins (1966), but also 
the small number of Fabrician Libellulid types from the Sir Joseph Banks collection, 
which were studied by Campion (1917). In both these cases, the original type data 
is not repeated, but reference is made to the appropriate paper in which it may be 
found. 

The general lay-out follows that of Kimmins (1966), each entry beginning with the 
specific name (in alphabetical order), author, the genus in which it was described, 
followed by the date and page and figure references. Full references are given at the 
end of the paper. Next follows the status of the type specimen and the label data, 
that of each label being separated from the next by an oblique stroke. 

Any comments are given in a second paragraph. Square brackets enclose addi- 
tional information, such as the completion of abbreviated locality names; [WFK] 
indicates that a label is in Kirby's writing. Type [McL. label] indicates the small, 
rectangular type-label, usually on mauve or reddish paper, used by McLachlan. 
BM Register numbers are not given. 

In the case of species described by Calvert (1909), it appears to have been his 
practice to label as TYPE the example figured. These are to be considered as 
Holotypes. (Calvert, 1901-08 : xxviii). 

This list of types was completed in February 1968. 

ENTOM. 22, 6 14 



2 8o D. E. KIMMINS 

adolescens Kirby (Beblecia), 1900 : 71-72, fig. 2. LECTOTYPE $ (abdominal segments 

5-10 missing). Free Town, Sierra Leone, I3.ix.99, E. E. Austen /Beblecia adolescens Kb. 

type [WFK]. 

Of the three examples listed by Kirby, only one (the lectotype) bears his determination 

label, as recorded above, but the other two examples, 1^,1$, are recognizable by the locality 

data and are considered to be paralecto types. No type was designated in print, but the 

example bearing his determination label has been selected as Lectotype. This specimen has 

for many years carried a red type label, now replaced by a lectotype label. This taxon is 

currently considered a synonym of Crocothemis divisa Karsch. 
aenea Kirby (Zygonidia), 1905 : 275-277. Holotype <$. Tonkin/Zygonidia aenea Kirby, 

<J Holotype, D. E. Kimmins det. 1961. 

Currently placed as a synonym of Zygonyx iris Selys. 
aequalis Kimmins (ssp. Rhyothemis phyllis), 1936 : 78-80, text-fig. 7 and pi. 3. Holotype <J. 

New Hebrides, Malekula, Ounua, 2i.iii.i929, Miss L. E. CAeesmaw/Rhyothemis phyllis 

aequalis Kimmins <J, det. D. E. Kimmins. 
aethiopica Kimmins (ssp. Atoconeura biordinata), 1958 : 357-358, figs. 7a-c. Holotype <J. 

Ethiopia, Segheria, 25.iii.i948, K. M. Guichard/Atoconeura, biordinata aethiopica Kim., 

(J Type, D. E. Kimmins det. 1958. 
offinis Kirby (Neurothemis), 1889 : 323, pi. 54, fig. 2. Holotype <J. Barang/N. affinis type 

[WFK]. 
This specimen has lost its BM register number, so that it is not possible to check the accuracy 

of the locality label 'Barang', which is in Indochina. It is probably wrongly labelled, since 

the taxon is currently placed as a synonym of Erythrodiplax funerea Hagen, a S. American 

species. 
alcestis Tillyard (Rhyothemis), 1906 : 482-483, pi. 44, fig. i. Holotype ? (lacking head). 

Kuranda, N. Q., F. P. Dodd, xii . 07/Rhyothemis alcestis Till., $ TYPE, R.J.T. 

Described from a single example. Currently placed in the synonymy of Rhyothemis 

braganza Karsch. 
allogenes Tillyard (Agrionoptera), 1908 : 641-643. LECTOTYPE <J. Cairns, N. Q., iv.os, 

E. Allen/AgTionoptera. allogenes Till., TYPE, R.J.T. /Agrionoptera allogenes Tillyard, $ Lecto- 
type, D. E. Kimmins det. 1967. 

Currently placed as a subspecies of A . papuensis Selys. 
amaryllis Selys (Rhyothemis), 1878 : 299. Holotype <J (lacking head). Menado/Type 

[McL. label]/Lib. amaryllis $ Selys. 

The unique holotype was originally in the McLachlan collection. Currently placed as a 

synonym of Rhyothemis phyllis snelleni Selys. 
amazonicaKirby(Ephidatia), 1889 : 331 (pars). LECTOTYPE $. Santarem/i68/Ephidatia 

amazonica type [WFK]/Ephidatia amazonica Kirby, <J Lectotype, D. E. Kimmins det. 1967. 
This species was restricted by Ris, 1913 : 1013 to the <J and $ from Santarem, the # from 

Para becoming a paratype of Ephidatia batesi Ris, 1913. E. amazonica Kirby (restr.) is 

currently considered a synonym of Idiataphe longipes cubensis (Scudder). 
amazonica Kirby (Fylgia), 1889 : 344, pi. 51, figs. 2, 3. LECTOTYPE <J. Para [label 

replaced, DEK]/Fylgia amazonica type $ [WFK]/Fylgia amazonica Kirby, $ Lectotype, 

D. E. Kimmins det. 1967. 

Kirby's label 'Fylgia amazonica type $' had been wrongly attached to an immature <J and 

has been transferred to one of the two $ syntypes, both of which lack heads. 
ambiguus Kirby (Misthotus), igo^a : 193. Lectotype <J (Kimmins, 1957 : 96-97)- C[ape 

of] G[ood] H [ope] /Misthotus ambiguus cotype [WFK] /Misthotus ambiguus Kirby, $ Lectotype, 

D. E. Kimmins det. 1956. 

Currently placed as synonym of Trithemis dorsalis (Rambur). The other syntype of 

ambiguus (Transvaal) was placed by Longfield (1936 : 490) as a synonym of Trithemis risi 

Longfield (1936 : 490). 



LIBELLULID & CORDULIID TYPES IN BM(NH) 281 

anornala Kirby (Anatya), 1889 : 338, pis. 53, fig. 9 and 57, fig. 7. LECTOTYPE <J. Para/3/ 
Anatya anomala type [WFK]/ Anatya anomala Kirby, Lectotype, D. E. Kimmins det. 1967. 
Currently placed as a synonym of A natya guttata (Erichson) . 

anomala Pinhey (Trithemis), 1955 : 38-39, fig- 8. Holotype $. Lake Chila, Abercorn, 
N. R., iv.i954, E- Pinhey /Trithemis anomala Pinh. 1954, Holotype/Male and Female in 
Copula. 

apicalis Fraser (ssp. Hylaeothemis fruhstorferi), 1924. Kimmins, 1966 : 178-179. 

apicalis Kirby (Rhyothetnis), 1889 : 319, pi. 51, fig. 5 Holotype ?. Aneit[yum], Wallace/ 
apicalis type [WFK]. 

This taxon is currently placed as a subspecies of Rhyothemis phyllis (Sulzer) . 

apicalis Kirby ( Untamo), 1889 : 331, pi. 53, fig. 4. Holotype $. Sul[a]/Sula, Wallace /\Jnta.mo 
apicalis type [WFK]. 

Currently placed as a synonym of Neurothemis ramburi (Brauer). 

ardens McLachlan (Thecadiplax), 1894 : 429-430. LECTOTYPE <J. Ta-chien-lu/Type 
[McL. label] /Thecadiplax ardens McL./Thecadiplax ardens McL , $ Lectotype, D. E. Kimmins 
det. 1967. 

Currently placed as a subspecies of Sympetrum eroticum (Selys). 

armstrongi Fraser (ssp. Rhyothetnis regia), 1956. Kimmins, 1966 : 179. 

arsinoe Lieftinck (Diplacina), 1953 : 171-173, fig. 6. Holotype <J. Papua, Kokoda, 1,200 ft., 
viii.1933, L. E. Cheesman/Diplacina, arsinoe sp. n. Holotype, det. M. A. Lieftinck, 1947. 

atra Pinhey (Trithemis), 1961 : 166-167, pi- II > n - J 4- Holotype <$. Uganda, Masindi Port, 
25. iv. 1956, P. S. Corbet/Triihemis atra Pinh., 1956, $ Type. 

attenuata Kirby (Trithemis?), 1889 : 328, pi. 53, fig. 2. LECTOTYPE <J. Amaz[on]/ 
Trithemis attenuata type <J [WFK] /Trithemis? attenuata Kirby, g Lectotype, D. E. Kimmins 
det. 1967. 

Kirby gives as locality 'Santarem', but the locality labels on the specimens marked by him 
as c and $ types are 'Amaz[on]' ( and one of the paratypes is labelled as 'Braz.'. In this case 
I am not interpreting the locality strictly as given in the description but have selected the 
cJ labelled by Kirby as type <J to be the lectotype. The taxon is currently placed in the genus 
Erythrodiplax . 

aurea Fraser (Tetrathemis), 1924. Kimmins, 1966 : 180. 

auricolor Fraser (var. of Notiothemis jonesi), 1944. Kimmins, 1966 : 181. 

austeni Kirby (Perithemis), 1897 : 602, pi. 12, figs. 4, 5. LECTOTYPE <J. Manaos, 

ii.ii.96, (J/Manaos, Brazil, n.ii.96, E. E. Austen /austeni [WFK] /Perithemis austeni Kirby, 

<J Lectotype, D. E. Kimmins det. 1967. 
austeni Kirby (Thermorthemis), 1900 : 72-73, pi. 2, figs, i, xa. LECTOTYPE <$. W. 

Afr[ica]/Thermorthemis austeni <J [WFK]. 

Kirby based his description upon five males, four from W. Africa (without other locality) 

and one male, one female from Sierra Leone. Three of the W. African males have been traced 

and also the male and female from Sierra Leone. This taxon is currently placed as Orthetrum 

austeni (Kirby). 
australis Kirby (Brachydiplax), 1894: 18-19. Holotype <J. Queensland, Turner /australis Kb. 

type [WFK]. 
australis Kirby (Brachymesia), 1889 : 330. Holotype <J. 'Sydney, N.S.W. '/australis type 

[WFK]. 

Locality label believed to be incorrect and the taxon is currently placed in the synonymy 

of Brachymesia furcata Hagen, a South American species. 
azorensis Gardner (ssp. Sympet rum fonscolombei), 1958 : 791-792. Holotype g. Azores, 

Pico 13, i7.viii.52, paired, leading, /. D. Carthy/Sympetrum fonscolombei azorensis Gardner, 

Type $, E. Gardner det. 

The holotype $ and allotype $ were taken paired. 



282 D. E. KIMMINS 

azteca Calvert (Tauriphila), 1906 : 297, 298. Holotype <$ (somewhat teneral). Guadaljaro, 
Jalisco, July, Schumann /Tauriphila azteca Calv. TYPE. P. P. Calvert det. 1906. B.C.A. 
Neur., p. 298. 

basalts Kirby (Micrathyrid), 1897 : 610-612, pi. 12, fig. i. LECTOTYPE <J. W. end of 
Parana de Buyassu, i6.i.96/Parana de Buyassu, Lower Amazon, 16.1.96, E. E. Austen/ 
basalis [WFK]/Micrathyria basalis Kirby, <$ Lectotype, D. E. Kimmins det. 1967. 

Kirby mentions two other localities after his description, Obidos ($) and Breves ($) and I 
consider these three examples to be the syntype series. Three additional males discussed 
later by Kirby are considered as additional non-syntype material, although one of them bears 
a Kirby cotype label. The taxon is currently placed in the genus Erythrodiplax. 

basilinea McLachlan (Libellula), 1894 : 430-431. LECTOTYPE <J. Ta-chien-lu/Type [McL. 

label] /Libellula basilinea McL. /Libellula basilinea McL., $ Lectotype, D. E. Kimmins det. 

1967. 
batesi Kirby (Cannacria), 1889 : 341, pis. 53, fig. i and 57, fig. 9. Holotype <J. Amaz[ons]/ 

Cannacria Batesii type [WFK]. 

The ending of the specific name has been changed to a single 'i' in accordance with Art. 31. 

This taxon is currently placed in the genus Brachymesia. 

bella Kirby (Perithetnis), 1889 : 324, pi. 51, figs. 8, 9. LECTOTYPE <J. ii5/Santarem/bella 
Kb. <J [WFK]/Perithemis bella Kirby, <J Lectotype, D. E. Kimmins det. 1967. 

The example in the BM type collection, though bearing Kirby's determination label, was 
not chosen as lectotype since its locality label is 'Amaz.', whereas the type-locality is specified 
as 'Santarem'. 

biflda Eraser (Tetrathemis), 1941. Kimmins, 1966 : 182. 

bimacula Kimmins (Misagria), 1943 : 156-159, figs. 1-6. Holotype <J. R. Demarara, B. 
Guiana/Misagria bimacula Kim., $ TYPE, det. D. E. Kimmins. 

[biolleyi Calvert (Orthemis), 1906 : 233, 237, pi. 9, figs. 36, 37. 

The example in BM(NH) is one of the syntypes but is not labelled Type by Calvert. In the 
circumstances, Ris (1910 : 286-287) should be considered as having selected the $ in the 
Calvert Collection (Costa Rica, Esparta) as Lectotype, and the <J in BM(NH) from Bugaba 
is therefore a paralectotype.] 

blackburni McLachlan (Lepthemis), 1882 : 229-231. LECTOTYPE <J. Hawaiian Islands/ 
Lepthemis? Blackburni McL./Type [McL. label] /Lepthemis blackburni McLachlan, $ Lecto- 
type, D. E. Kimmins det. 1967. 
Now placed in the genus Nesogonia. 

borneense Kimmins (Orthetrum), 19360 : 70-73, figs. 2, 3. Holotype $. Sarawak, Mt. Dulit, 
4,000 ft., 22.x. 1932. B in cop. A/Oxford Univ. Exped., B. M. Hobby, A. W. Moore &> J. 
Ford/Orthetrum borneense $ sp. n., Holotype, det. D. E. Kimmins. 

braueri Kirby (Lyriothemis), 1889 : 332. Holotype ?. Sula, Wallace, D. E. Kimmins, 
I9&7/W. Borneo [WFK]/L. Braueri type [WFK]. 

Kirby gives the locality as Sula [Moluccas] and this locality is repeated by Ris (1909 : 108, 
in), after his study of the type. When examining this type during the preparation of this 
paper, it was found to be labelled 'W. Borneo' in Kirby's writing. It is very unlikely that Ris 
would not have remarked on the discrepancy in the locality label when he studied the type, 
and one can only assume that at some later date there was been a change of label. I have 
therefore placed a 'Sula' label above the W. Borneo label. This taxon is currently placed as a 
synonym of Lyriothemis cleis Brauer. 

brevistylum Kirby (Orthetrum), 1896 : 521-522. Holotype $. Dobar, Goolis Mts., 4.1!. 
1895, L. L[ort-] P[Ai//t/w]/Orthetrum(?) brevistylum n.s. [WFK]. 
Currently placed as a synonym of Orthetrum taeniolatum (Schneider). 



LIBELLULID & CORDULIID TYPES IN BM(NH) 283 

broadwayi Kirby (Dythemis), 1894^ : 227-228. LECTOTYPE <J. Trinidad/broadwayi/ 
Dythemis broadwayi Kirby, <$ Lectotype, D. E. Kimmins det. 1967. 
Currently placed as a synonym of Dythemis velox Hagen. 

burmeisteri Kirby (Tramea), 1889 : 316. LECTOTYPE <J. N.W. India/Burmeisteri Kb. 
type [WFK]/Tramea burmeisteri Kirby, $ Lectotype, det. D. E. Kimmins, 1967. 

In addition to the allotype $, there are 5 paralectotypes in the BM(NH). The taxon is 
currently placed as a subspecies of Trapezostigma basilaris (Palisot de Beauvois). 

camarense Kirby (Orthetrurri), 1889 : 298. Holotype <J. Cameroons/camarense type 
[WFK]/Orthetrum camarense Kirby, $ Holotype, D. E. Kimmins det. 1967. 

cambridgei Kirby (Micrathyria), 1897 : 608-609, pi. 13, fig. 4. Holotype $. Breves, 
I3.i.96/Breves, Lower Amazon, 13.1.96, E. E. A wstew/cambridgei [WFK]. 

campioni Ris (Allorhizucha), 1915^ : 214-216; Kimmins, 1942 : 47 [selection of holotype]. 
Lectotype <J. KaYima, Sierra Leone, 24. vi. 1912, Jas. J. Simpson/ 'Allorhizucha nova spec. 
C, Det. Dr. F. Ris/Allorhizucha campioni Ris, <J Type, det. D. E. Kimmins. 

cannacrioides Calvert (Dythemis), 1906 : 272, 276, pi. 8, figs. 43, 44. Holotype <J. San 
Isidro, i, 600 ft., C/zaw^ncw/Dythemis cannacrioides Calv. TYPE. P. P. Calvert det. 1906. 
B. C. A. Neur., p. 277. Orig. PI. viii, figs. 43, 44. 

The penis of the lectotype was removed and studied by Dr. K. Buchholz. 

carpenteri Fraser (Oxythemis), 1944. Kimmins, 1966 : 186. 
carpenteri Fraser (Tetrathemis), 1941. Kimmins, 1966 : 186. 

celebensis Kirby (Protorthemis), 1889 : 334-335, pis. 54, fig. 7 and 57, figs. 6, 6a. Holotype <J. 

Mak[assar]* /Celeb [es], WallacejO. celebensis type [WFK]. 

ceylonica Kirby (Zygonidia), 1905 : 273-275 Holotype <$. Ceylon. E. E. Green /Kandy, 
ii.igoi [E. E. Green's writing] /Zygonidia ceylonica Kirby, <$, D. E. Kimmins det. 1961. 
Currently placed as subspecies of Zygonyx iris Selys. 

chirinda Longfield (ssp. Atoconeura biordinata), 1953 : 46, pi. i, figs. 3, B, H. Holotype <J. 
Chirinda For., Melsetter Dist., Dept. Agric. S. Rhodesia, 27.xii.i948, /. A. Whellan/A.toco- 
neura biordinata chirinda, Type <$ subsp. nov., det. Miss C. Longfield. 

chl oe Kirby (Rhyothemis), 1894 : 15-16. LECTOTYPE?. Queensland, Turner/chloe Kirb. 
type [WFK] /Rhyothemis chloe Kirby, ? Lectotype, D. E. Kimmins det. 1967. 

The lectotype $ is the darker of the two syntypes, with (in the hind wing) the 'dark spot on 
the subnodal sector, halfway between the dark blotch on the nodus and the clouded tip of the 
wings' mentioned by Kirby. Currently placed as a subspecies of Rhyothemis phyllis. 

chrysobaphes Ris (Pseudomacromia), 1915^ : 221-223. LECTOTYPE . Sierra Leone, 
Sandea, 14. vi. 1912, /. /. Siw^sow/Pseudomacromia nova spec, d, Type <$, Det. Dr. F. Ris/ 
Pseudomacromia chrysobaphes Ris, $ Lectotype, D. E. Kimmins det. 1967. 
Currently placed in the genus Zygonyx. 

cingulata Kirby (Nesoxenia), 1889 : 336, pi. 53, fig. 8. LECTOTYPE $. [Solomon Islands], 
Alu/Nesoxenia cingulata type [WFK] /Nesoxenia cingulata Kirby, $ Lectotype, D. E. Kimmins 
det. 1967. 

The syntype series also contained i $ (incomplete) and i $. This taxon is currently con- 
sidered a subspecies of Nesoxenia my sis Selys. 

cladophila Tillyard (Tetrathemis), 1908 : 647-648, pi. 14, fig. 5. LECTOTYPE <J. Cook- 
town, N. Q., i.o8, R. J. Tillyard /Tetrathemis cladophila Till. $ TYPE. R. J. T./Tetrathemis 
cladophila Tillyard, <J Lectotype, D. E. Kimmins det. 1967. 

* This label was transcribed by Kirby as Makian, but according to Bradley & Betrem, 1967, 
Bull. Brit. Mus. nat. Hist. (Ent.) 20 (7) : 292, the locality should be Makassar. 



284 D. E. KIMMINS 

corduliformis Longfield (Tetrathemis), 1936 : 484, figs. 6, A-D. Holotype <J. Central 
Africa, Uganda, Bunyoro Distr., Nyamagita Dam, 18.11.1934, C- E. Longfield /Tetrathemis 
corduliformis sp. n. Type $, det. Miss C. Longfield. 

coryndoni Fraser (Lokia), 1952. Kimmins, 1966 : 187. 

cuprina Kirby (Rhyothemis), 1889 : 320, pi. 51, fig. 6. Holotype ?. S. Leone/cuprina Kb. 
type [WFK]. 

Currently placed as a synonym of Rhyothemis fenestrina (Rambur) . 

dalei Tillyard (Nannodythemis), 19080, : 449-450, pi. 6, fig. 2. LECTOTYPE <$. Blue 
Mts., N.S.W., ii.igoS, R. J. Tilly ard/Nannodythemis Dalei Till. ^ TYPE, R. J. T./Nannody- 
themis dalei Tillyard, $ Lectotype, D. E. Kimmins det. 1967. 
Currently placed in the genus Nannophya. 

darwini Kirby (Tramea), 1889 : 315, pi. 51, fig. i. LECTOTYPE $. Galapagos/Galapagos/ 
28/Darwinii Kb. type [WFK]. 

The remaining four female syntypes (in bad condition) are still in the B.M. collection. This 
taxon is currently placed as a synonym of Trapezostigma cophysa (Selys). 

davina Fraser (ssp. Zygonyx iris), 1926. Kimmins, 1966 : 188. 
denticauda Fraser (Tetrathemis), 1954. Kimmins, 1966 : 189. 

diamangae Longfield (Aethiothemis), 1959 : 35, 36, fig. 5A. Holotype <$ (teneral), N. Angola. 
Lunda Prov., Dundo, iii. 1949/Aethio therm's diamangae, Type <$, det. Miss C. Longfield. 

dispar Selys (Trithemis, Paberration), 1883 : 107. Holotype $. Japan/Type [McL. label]/ 
Trithemis phaon, var. $ dispar Selys. 

Currently placed as a synonym of Deielia phaon (Selys) . 

disparilis Kirby (Neurothemis), 1889 : 322-323, pi. 54, fig. 8. LECTOTYPE <J (head 
missing). W. Borneo/N. disparilis ^ [WFK]/Neurothemis disparilis Kirby, <J Lectotype, 
D. E. Kimmins det. 1967. 

dissocians Calvert (Micrathyria), 1906 : 222, 226, pi. 9, figs. 19-21. LECTOTYPE $. 
Atoyac, Vera Cruz, May, H.H.S[mith] /Micrathyria dissocians Calv. Q* TYPE. P. P. Calvert 
det. 1906. B.C.A. Neur., p. 226, Orig. of pi. ix, figs. 19-21 /Micrathyria dissocians Calvert, 
cJ Lectotype, D. E. Kimmins det. 1967. 

Ris (1911 : 440) refers to the Portorico specimen as 'Calvert's Type'. I do not consider 
this to be a designation of Lectotype, and consider that the data on the above mentioned 
example makes it clear that Calvert considered it to be the type, especially as the second Vera 
Cruz example is not marked TYPE. 

distanti Kirby (Stoechia), 1898 : 236-237. LECTOTYPE . Pretoria (W. L. Distant])/ 
distant! Kirby type [Distant's writing] /Stoechia distanti Kirby, $ Lectotype, D. E. Kimmins 
det. 1967. 

Currently placed as synonym of Trithemis dorsalis (Rambur). 

donaldi Lieftinck (Lanthanusa), 1955 : 161-164, n S s - 5~9- Holotype <J. N.E. New Guinea, 
Saiko, 5,500-6,000 ft., Babu River (Upper Waria River), ix-x. 1936, D. Shaw-Meyer /'Lanthan- 
usa, donaldi Lieft. Det. M. A. Lieftinck, 1955. Holotype. 

dorothea Fraser (ssp. Agrionoptera insignis), 1927. Kimmins, 1966 : 189. 
dubia Fraser (Porpacithemis), 1954. Kimmins, 1966 : 190. 

ducalis Kirby (Rhyothemis), 1898 : 230-232. LECTOTYPE <J. Fort Johnston, Nyassaland, 
P. Rendall/Khyoihemis ducalis Kirby, <J Lectotype, D. E. Kimmins det. 1967. 

Two <J, one $ of the four syntypes are from Fort Johnston; the $ from Pretoria, though 
marked 'cotype' by Kirby, was in effect excluded from the syntypes by Kirby's statement 
(231) The specimens from Fort Johnston are to be considered typical.' Currently placed 
as a synonym of R. semihyalina Desjardins. 

eltoni Fraser (Phyllothemis), 1935. Kimmins, 1966 : 191. 



LIBELLULID & CORDULIID TYPES IN BM(NH) 285 

eludens Tillyard (Nannophlebia), 1908 : 645-647, pi. 14, fig. 4. LECTOTYPE <J. Cairns, 
N. Q., v.o5, E. A //ew/Nannophlebia eludens Till. $ TYPE, R.J.T. /Nannophlebia eludens Till- 
yard, (J Lectotype, D. E. Kimmins det. 1967. 

equestris Fabricius (Libelluld), 1781 : 523. Campion, 1917 : 445. 

equivocata Kirby (Thermochoria), 1889 : 339, pi. 52, fig. 8. Holotype <$. W. Afr[ica]/ 
Thermochoria equivocata type [WFK]. 

erichsoni Kirby (Trithemis), 1894 : 263. LECTOTYPE <J. Amaz./erichsoni [WFK]/ 
Trithemis erichsoni Kirby, Lectotype, D. E. Kimmins det. 1967. 

This specimen was the basis of the drawings and notes sent by the present author to D. J. 
Borror in Jan. 1935, and referred to in his monograph on Erythrodiplax. This taxon is 
currently placed in synonymy of Erythrodiplax unimaculata (De Geer) . 

eudoxia Kirby (Accaphila), 1909 : 60. Holotype $. Ruwenzori East, alt. 6,000 ft., ^o.i.ojj 
Ruwenzori Exp./ Accaphila eudoxia Kb. type. 
Currently placed in the genus Atoconeura. 

eurybia Selys (Libellula), 1878 : 293, 298. Holotype <J. Menado/Type [McL. label] /Lib. 
eurybia Selys <J, race de Chinensis? [Selys writing]. 
Currently placed in the genus Trapezostigma. 

exigua Kirby (Fylla), 1889 : 345, pi. 52, fig. 6. LECTOTYPE <J. Gil[olo]/PFa//ac*/Fylla 
exigua Kirby, $ Lectotype, D. E. Kimmins det. 1967. 

The syntype series was found in the museum collection, without type-indication or deter- 
mination labels by Kirby. One can only presume that as Kirby reduced the taxon to syn- 
onymy of Nannophya pygmaea Rambur in his Catalogue of Odonata published the following 
year, he removed his determination labels. The series now contains paralectotypes from 
Borneo, Gilolo and Morty. 

exitnia Kirby (Micrathyria), 1897 : 609, pi. 13, fig. 3. LECTOTYPE <J. Obydos, 2.ii.96/ 
Obydos, Lower Amazon, 2.11.96, E. E. Austenjeximia. [WFK] /Micrathyria eximia Kirby, 
<J Lectotype, D. E. Kimmins det. 1967. 

The lectotype is the specimen which, for many years, has carried a red BM type label. The 
remaining 3 $ syntypes have been labelled paralectotypes. Since Kirby stated that the 
species was described from four males from Obydos, the male mentioned by him from Para is 
excluded from the syntype series. 

extraordinata Fraser (Atoconeura), 1950. Kimmins, 1966 : 191. 

falsum Longfield (ssp. Orthetrum capense), 1955 : 26. Holotype <$. B. E. Africa, Kenya, 
Thoura Forest, Meru, 600 ft., 30.1.1934, far from water, C. E. Longfield/Orthetrum capense 
falsum Longf., Type $, det. Miss C. Longfield. 

Currently placed as subspecies of Orthetrum capicola Kimmins. 

fasciata Kirby (Celithemis), 1889 : 326, pi. 52, fig. 2. LECTOTYPE <J. Georgia/fasciata 
Kb. type [WFK] /Celithemis fasciata Kirby, $ Lectotype, D. E. Kimmins det. 1967. 

fasciata Kirby (Deielia), 1892 : 330, pi. 53, fig. 6. McLachlan, 1892 : 177-178. Holotype $. 
i37/Sandw[ich] Is., Beechey/'Loo Choo Is , Voy. of 'Blossom', See McLachlan, An. Mag. Nat. 
Hist. (6) x. p. 177 (1892) /Deielia fasciata, type [WFK]. 

As explained by McLachlan (1892), the original locality label on this specimen appears to 
have been incorrect, as the insect has not been refound in the Hawaiian Islands. It does 
occur in the Loo Choo (Ryukyu) Islands, which were also visited by the 'Blossom', and it is 
considered most likely that the example came from these islands. The taxon is currently 
considered a synonym of Deielia phaon (Selys). 

ferruginata Fabricius (Libellula), 1781 : 521. Campion, 1917 : 445. 

fitzgeraldi Pinhey in Longfield (Nesciothemis), 1955 : 61-63, n g s - I0 B > D F ; Pinhey, 1955 : 
30-32, fig. 6. Holotype (J. Lake Chila, Abercorn, N. R., 31.1.54, D. Vesey-Fitzgerald/ 
Orthetrum(P) fitzgeraldi Pinh. 1954, Holotype/Nesciothemis fitzgeraldi (Pinhey), det. Miss 
C. Longfield. 
ENTOM. 22, 6. i4 



286 D. E. KIMMINS 

The type of Orthetrum(!) fitzgeraldi Pinhey was presented to the BM(NH) when the manus- 
script was sent to press. Publication was unavoidably delayed, and in the belief that Pinhey's 
paper was due to appear, Miss Longfield included a description of fitzgeraldi in her paper on 
African Orthetrum, placing it in her new genus Nesciothemis, the publication of which un- 
fortunately antedated Pinhey's paper. 

flava Longfield (Monardithemis), 1947 : 23-25, figs. 9-10. Holotype $ (original designation 
as type). Kalukembe, Dec. /Angola, Miss. sc. suisse, 1932-1933/Monardithemis flava genotype 
<$, det. Miss C. Longfield. 

flavescens Kirby (Miathyria), 1897 : 600-601, pi. 13, fig. 2. LECTOTYPE <J. Santarem/ 
113 <J/ Miathyria flavescens Kirby, $ Lectotype, D. E. Kimrnins det. 1967. 

The lectotype $, which has for many years carried a red BM type-label, has no determina- 
tion label by Kirby, but the locality label agrees with the type-data and there is no doubt that 
it is one of the syntypes. On the other hand, an example bearing Kirby's label 'Miathyria 
flavescens K. cotype' is from Para, not Santarem and is from the Saunders collection (not 
Bates) and cannot therefore be a syntype. 

flavescens Kirby (Tetrathemis), 1889 : 343, pi. 52, fig. 4. LECTOTYPE <J. Sar[awak]/ 
Wallace^, flavescens type [WFK] /Tetrathemis flavescens Kirby, <J Lectotype, D. E. Kimmins 
det. 1967. 

flavidulum Kirby (Orthetrum), 1898 : 238. LECTOTYPE $. Pretoria, W. L. D.[istant]/ 
Orthetrum flavidulum Kirby, ? Lectotype, D. E. Kimmins det. 1967. 

Of the four examples (listed by Kirby as females), the example from the Pienaars River, 
iii. 1894 proves to be a <J. Currently placed as synonym of Orthetrum abbotti Calvert. 

flavifrons Kirby (Chalcostephia), 1889 : 337. Holotype <J. Angola/Chalcostephia flavifrons 
type. 

flavopicta Kirby (Orthemis), 1889 : 332-333, pis. 54, fig. i and 57, fig. 5. LECTOTYPE <J. 
Para/43/Orthemis flavopicta type/Orthemis flavopicta Kirby, Holotypus prap. Dr. Buchholz/ 
Orthemis flavopicta Kirby, <J Lectotype, D. E. Kimmins det. 1967. 

From the syntype series of 4 $, i <j>, the $ bearing Kirby's type label has been chosen as 
lectotype. 

foliata Kirby (Belonia), 1889 : 333, pi. 54, fig. 4. Holotype (by original designation). 
[Mexico], Duenas/foliata Kb. type [WFK]. 

Currently placed in the genus Libellula. The $ from Guatemala, doubtfully referred to 
Belonia foliata by Kirby, is now considered to be a synonym of Libellula herculea Karsch. 

jraseri Pinhey (Olpogastra), 1955 : 39-40, pi. 3, fig. i, text-fig. 8. Holotype <J. Aswa R., 
Acholi, Uganda, March 1952, T. H. E. Jackson. Olpogastra fraseri Pinh. 1954, Holotype. 
Currently placed in the genus Zygonoides Pinhey. 

frontalis Kirby (Lyriothemis), 1889 : 332. Holotype $. Sul[a]/Sula, Wallace [L. frontalis 
type [WFK]. 

Currently placed as a synonym of Lyriothemis cleis Brauer. 

fulgens Kirby (Rhyothemis), 1889 : 322. Holotype <J. Sar[awak]/fulgens, Kb. type [WFK] 

gamblesi Longfield (Oxythemis), 1959 : 36-38, figs. 5 B, C. Holotype #. S. Nigeria, Ikorodu, 
Ogun Delta, 26.11. 1956, R. M. Gambles /Oxythemis gamblesi Type <J, det. Miss C. Longfield. 

gardneri Fraser (Hylaeothemis), 1927. Kimmins, 1966 : 194. 

grenadensis Kirby (Brechmorhoga), 18946 : 265. LECTOTYPE <J. Grenada/ io6/grena- 
densis [WFK] /Brechmorhoga grenadensis Kirby, <J Lectotype, D. E. Kimmins det. 1967. 
Currently placed as a subspecies of Brechmorhoga praecox (Hagen). 

[hageni Kirby (Micrathyria), 1890 : 41] 

This name was proposed for Dythemis didyma Hagen, 1861, nee Selys, 1857 and the type 
should therefore be selected from Hagen's material. 



LIBELLULID & CORDULIID TYPES IN BM(NH) 287 

hyalinum Kirby (Orthetrum), 1886 : 326-327, pi. 33, figs. 5, 6. LECTOTYPE <J. N.W. 
Ind[ia]/Campbellpore, i4.xi.85, 7/hyalinum Kirb. (Jtype [WFK] /Orthetrum hyalinum Kirby, 
<$ Lectotype, D. E_. Kimmins det. 1967. 

Currently placed as a synonym of Orthetrum taeniolatum (Schneider). 

hyalina Kirby (Tetrathemis), 1889 : 342, pi. 56, fig. 8. Holotype <J (lacking abdominal 
segments 5-10). Borneo/T. hyalina type [WFK]. 

Currently placed as subspecies of Tetrathemis irregularis Brauer. 

[icteroptera Selys (Libellula), 1857, in Sagra, Hist. Cuba, 7 : 190.] 

This species was briefly differentiated as a race in a note on Libellula metella Selys ; the type 
locality being given as Buenos Ayres. The McLachlan collection included an example 
determined by Selys as Lib. icteroptera Selys $, to which McLachlan had added a type label. 
This cannot however be the type, as its locality label is Minas Geraes. 

immaculata Fraser (Zygonyx), 1933. Kimmins, 1966 : 196. 

imperatrix Selys (Rhyothemis), 1887 : 53-54. Holotype <J. Ryukyu [Loo Choo], vi-viii. 1887 
[Pry /] /Rhyothemis imperatrix Selys, nov. sp.? ou race de splendida, R. <$, Loo Choo [Selys' 
writing]. 

Selys saw originally only this one example, but McLachlan subsequently received three 
females with the same locality data (collected by Pryer), of which one was given to Selys and 
was described by him (1888 : i) ; the other two females are now in BM(NH). 

indica Fabricius (Libellula), 1781 : 521. Campion, 1917 : 444. 
indica Fraser (Hylaeothemis), 1946. Kimmins, 1966 : 196. 

indica Kirby (Brachydiplax), 1889 : 329, pi. 54, fig. 9. LECTOTYPE <J. N. Ind[ia]/B. 
indica $ type [WFK] /Brachydiplax indica Kirby, $ Lectotype, D. E. Kimmins det. 1967. 

The mature ^ has been chosen as lectotype, as this example has been segregated in the type- 
collection since 1940. 

infuscatum Selys (Diplax), 1883 : 90. LECTOTYPE <J. Japan/Type/Diplax infuscata 
Selys, < Japon/Diplax infuscata Selys, <J Lectotype, D. E. Kimmins det. 1967. 
Currently placed in the genus Sympetrum. 

insignis Kirby (Zygonidia), 19000 : 533-534, pi. 12, fig. i. LECTOTYPE <J. 5-fingered Mt., 
Hainan, /. T. Thomasson/Zygonidia insignis, type [WFK] /Zygonidia insignis Kirby, <$ Lecto- 
type, D. E. Kimmins det., 1967. 

Currently placed as a synonym of Zygonyx iris Selys. 

insularis Kirby (Agrionoptera), 1889 : 336-337. LECTOTYPE <J. [Solomon Is.] Santa 
Anna/Agrionoptera insularis $ type [WFK] /Agrionoptera insularis Kirby, <J Lectotype, 
D. E. Kimmins det. 1967. 

Currently placed as a subspecies of Agrionoptera insignis (Rambur). 

insularis Kirby (Cannaphila), 1889 : 341. Lectotype $ (Ris, 1910 : 295). Hayti/Canna- 
phila insularis Kirby, $ Lectotype, F. Ris design. 1910 [D.E.K. hand]. 

I consider that Ris 1 action (1910 : 295) in citing 'British Museum: i $ juv. Hayti (Kirby's 
Type)' is equivalent to selection of a lectotype. There is one other $ in BMNH, which bore 
Kirby's determination Cannaphila insularis, but this has a locality label St. Dom. [Santo 
Domingo], which is in Haiti and was probably a syntype. This is now labelled paralectotype. 
I have been unable to trace the <$ syntype referred to by Kirby. 

Kirby's statement 'Hob. Haiti (types in British Museum); Jamaica (Dublin Museum)' 
eliminated the Dublin Museum example (s) from consideration as lectotype. 

intensa Kirby (Perithemis), 1889 : 326, pi. 51, fig. 7. Holotype $ (abd. segs. 9-10 missing). 
Mex[ico]/intensa Kb. type [WFK]. 

intermedia Fraser (Protorthemis), 1936. Kimmins, 1966 : 198. 



288 D. E. KIMMINS 

internum McLachlan (var. of Orthetrutn japonicum), 1894 : 431-432. LECTOTYPE <$. 
Ta-chien-lu/O. japonicum var. internum McL./Orthetrum japonicum internum McL., <J Lecto- 
type, D. E. Kimmins det. 1967. 

Currently placed as Orthetrum japonicum internum McL. 

iridescens Kirby (Urothemis), 1898 : 235. LECTOTYPE <J (abd. segs. 6-10 missing). Fort 
Johnston, Nyassaland, P. Rendall/ iridescens Kirby type [WFK] /Urothemis iridescens Kirby, 
(J Lectotype, D. E. Kimmins det. 1967. 
Currently placed as synonym of Urothemis edwardsi (Selys). 

isa Fraser (ssp. of Zygonyx iris), 1926. Kimmins, 1966 : 198. 

isis Fraser (Zygonyx), 1924. Kimmins, 1966 : 198. 

iulia Kirby (Orthetrum), 1900 : 75-77, pi. 2, fig. 3. LECTOTYPE <J (abd. segs. 9-10 missing). 
Sierra Leone/Orthetrum julia type [WFK]. 

Kirby writes (: 77) 'The species is described from four specimens from Sierra Leone, two 
of which were collected by Mr. Austen on Aug. 26 and Sept. n (1899).' The example desig- 
nated as Lectotype is the only one bearing Kirby's determination label. The allotype female 
has for many years borne a museum type label but now has no locality label (? Sierra Leone, 
42.31). Of the listed examples collected by Austen, one (without abdomen), labelled 'Free 
Town, Sierra Leone, 26.viii. 1899, E. E. Austen' has been located but the other is missing. 

kalai Longfield (ssp. Orthetrum stemmale), 1936 : 487, 488, fig. 7. Holotype (J. S. Rho- 
desia, Victoria Falls, Zambesi River, Kalai Island, 7.^.1936, C. E. Longfield /OrthetTum 
stemmale kalai sp. nov. Type <J, det. Miss C. Longfield. 

kalula Kirby (Trithemis), 1900 : 69, pi. 2, figs. 2, aa. LECTOTYPE <J. Free Town, Sierra 
Leone, I3.ix.99, E. E. Austen/Trithemis kalula Kb. type [WFK]. 

Kirby lists four males, but only three can now be traced in the collections ; of these, only the 
lectotype bears Kirby's determination label. This specimen has for many years carried a red 
type label and has been considered to be the type. The two remaining males are considered 
paralecto types . 

kenya Longfield (spp. Atoconeura biordinata), 1953 : 46, pi. i, figs. 5, 9, C, G. Holotype <J 
B.E. Africa, Kenya, Meru, 6,000 ft., on swift stream, 29.1.1934, C. E. Longfield. Atoco- 
neura biordinata kenya, Type $ subsp. n., det. Miss C. Longfield. 

kerri Fraser (Amphithemis), 1933. Kimmins, 1966 : 198. 

lacustris Kirby (Trithemis?), 1889 : 329. Holotype <J. [Central Africa] Wadelai, 27.1.1887 
[Emin Pasha] /lacustris Kb. type [WFK]. 

Currently placed in the genus Brachythemis. 

lankana Kirby (Rhyothemis), 1893 : 549-550. Holotype <$. Udagama, 2 6. iv. 92 /Ceylon, 
Yerbury, (on reverse) lankana type/Rhyothemis lankana Kirby, <$ Holotype, D. E. Kimmins 
det. 1967. 

leakeyi Pinhey (Lokithemis), 1955 : 34-35, pi. 3, figs. 3, 4, 6, and text-fig. 7. Holotype <. 
Lake Chila, Abercorn, N. Rhodesia, 31.1.54, D. Vesey Fitzgerald /Lokithemis leakeyi Pinh. 
Holotype. 

[levis Calvert (Orthemis), 1906 : 233, 238, pi. 9, figs. 38, 39.] 

No example in our B.C.A. series was marked TYPE by Calvert and our specimens are there- 
fore considered as syntypes. The true type is probably in some other collection. 

lewisi Selys (Lyriothemis), 1883 : 96-97. LECTOTYPE <J. Japan, I^ww/Lyriothemis 
lewisii Selys $, Japon/Lyriothemis lewisi Selys, $ Lectotype, D. E. Kimmins det. 1967. 

Ris (1909 : 119) in the list of material dealt with cites 'i $ Japon (Pryer, Selys Type der 
L. lewisi)'; this cannot be accepted as a designation of lectotype, since Selys limits his material 
to 'Japon, par M. G. Lewis, communiquee par MacLachlan (coll. MacLachlan, Selys)'. I have 
therefore designated the above specimen as Lectotype. Currently placed as a synonym of 
L. pachygastra (Selys). 



LIBELLULID & CORDULIID TYPES IN BM(NH) 289 

lieftincki Fraser (Rhodothemis), 1954. Kimmins, 1966 : 201. 

lifuana Kimmins (ssp. Agrionoptera insignis), 1953 : 243-244, figs. 1-3. Holotype <J. 
Lifu/Agrionoptera insignis lifuana Kim. $ Holotype, D. E. Kimmins det. 1951. 
Currently placed as a subspecies of Agrionoptera papuensis. 

longipennis Kirby (Belonia), 1889 : 334. Holotype?. Cuensa/longipennis Kb. type [WFK]. 
Currently placed as a synonym of Libellula herculea Karsch. 

lorti Kirby (Orthetrum), 1896 : 522-523. Holotype $ (lacking abd. segs. 6-10). Dobar, 
Goolis Mts., 4.11.1896, L. L[ort-] P[hillips] /Orthetrum lorti. 
Currently placed as synonym of Crocothemis erythraea (Brulle). 

tnachadoi Longfield (Orthetrum), 1955 : 35-39, figs. 5, A, E, J. Holotype <$. N.E. Angola, 
Dundo, Lunda Distr., i3.xii. 1947/Orthetrum machadoi Type <J, in cop. with type $, det. 
Miss C. Longfield. 

macrostigma Longfield (Orthetrum), 1947 : 25-27, fig. n. Holotype <$. Lunda, sept./ 
Angola Miss. sc. suisse, 1932-1933/Orthetrum macrostigma Type $, det. Miss C. Longfield. 

madagascariensis Kirby (Tramea), 1889 : 317. Holotype <$. Betsileo, Madag. [WFK]/ 
madagascariensis Kb. type [WFK]. 

This taxon is currently placed as a synonym of Trapezostigma limbata (Desj .) . 
malabarica Fraser (ssp. Zygonyx iris), 1926. Kimmins, 1966 : 203. 
malcolmi Fraser (Brachydiplax), 1922. Kimmins, 1966 : 203. 
marginata Fabricius (Libellula), 1781 : 523. Campion, 1917 : 445. 

marshalli Kirby (Misthotus), 19050, : 192-193. LECTOTYPE $. Salisbury, Mashonaland, 
Marshall/5. Salisbury, Nov. 1903, in copula. G. A. K. M[arshall] /marshalli Kb. type 
[WFK] /Misthotus marshalli Kirby, <$ Lectotype, D. E. Kimmins det. 1967. 

The 'still more immature male' referred to by Kirby cannot be traced. Currently placed as 
a synonym of Trithemis dorsalis (Rambur). 

maya Calvert (Dythemis), 1906 : 272, 275, pi. 8, fig. 45. Holotype <J. San Geronimo, Vera 
Paz, C&aw^zow/Dythemis maya Calv. TYPE <$. P. P. Calvert det. 1906. B. C. A. Neur., 
p. 275. Original of PI. viii, fig. 45. 

The penis of the lectotype was removed and studied by Dr. Buchholz. 

metallica Fraser (ssp. of Zygonyx iris), 1931. Kimmins, 1966 : 203. 
mildredae Fraser (ssp. of Zygonyx iris), 1920. Kimmins, 1966 : 204. 

monteiroi Kirby (Thermorthemis), 1900 : 73. Holotype $. Angola/monteiroi Kb. type 

[WFK]. 

Currently considered a synonym of Orthetrum austeni (Kirby). 

multipunctata Kirby (Dythemis), 18946 : 265-266. LECTOTYPE $ (left fore wing 
missing). St. Vincent/Pools near Bannonallie, Jan. i2/multipunctata [WFK]/Dythemis 
multipunctata Kirby, <J Lectotype, D. E. Kimmins det. 1967. 

I have chosen as lectotype the $ which has for many years carried a BM red type label and 
of which the penis was removed and studied by Dr. K. Buchholz. 

multipunctata Kirby var. (Dythemis), 18946 : 266. LECTOTYPE <J. Grenada/235/ 
Dythemis multipunctata var. Kirby, J Lectotype, D. E. Kimmins det. 1967/Dythemis sterilis 
Hag., det. D. E. Kimmins. 

Currently placed as a synonym of Dythemis sterilis Hagen. 

nicevillei Kirby (Orthetrum), 18940 : 112-113. LECTOTYPE <J. Burma/nicevillei Kb. 
type $ [WFK] /Orthetrum nicevillei Kirby, <J Lectotype, D. E. Kimmins det. 1967. 

One of Kirby's determination labels had at some time been transferred from a $ syntype to 
a $ which was not received at the BM(NH) until after publication of the description. This 
label has been replaced on its correct specimen. Currently placed as synonym of Orthetrum 
glaucum Brauer. 



2QO D. E. KIMMINS 

nigra Longfield (ssp. of Trithemis donaldsoni), 1936 : 491-493, 496, figs. 9, 10. Holotype $ 
(last 5 abdominal segments missing). W. Africa, Principe I., Ogui Pipi, y.xii. 1932, W. H. T. 
Tams/Trithemis donaldsoni nigra ssp. nov. Type $, det. Miss C. Longfield. 

In the original description, only the above example was mentioned, but in a list (: 496), 
two (J are recorded. The second one (a complete example) had been labelled by Miss Longfield 
as type and the holotype labelled as paratype. This error has now been corrected and the 
second example cannot be considered even as a paratype, as it is not mentioned in the original 
description. 

nigricolor Fraser (Amphithemis), 1922. Kimmins, 1966 : 206. 

nigrifrons Kirby (Orthetrurri), 1894 : 19-20. LECTOTYPE <J. Queensland, Turner I 
Orthetrum nigrifrons Kirby, $ Lectotype, D. E. Kimmins det. 1967. 

The lectotype and paralectotype lack any determination label by Kirby. Currently placed 
in the genus Crocothemis. 

nigrilabris Selys (Lib[ellula]), 1872 : 177; (Urothemis), Selys, 1878 : 304-305. Holotype $. 
Menado/Lib. nigrilabris Selys, $ Type/Macrodiplax lycoris Selys. 

This taxon was first described briefly (1872) in comparison with Libellula lycoris Selys, and 
a full description, with location of type given, in 1878. Currently placed as synonym of 
Macrodiplax cora Brauer. 

notablis Kirby (Onychothemis), 1906 : 277-278. Holotype (J. Tonkin [Fruhstorfer]/ 
Onychothemis notabilis Kirby, Holotype $, det. D. E. Kimmins, 1967. 

No Kirby determination label. Currently placed as synonym of Onychothemis tonkinensis 
Martin, 1904. 

notata Fabricius (Libellula), 1787 : 337. Campion, 1917 : 444. 

novaezealandiae McLachlan (var. of Sympetrutn bipunctatutri), 1894^ : 271-272. LEC- 
TOTYPE $. 29e[Hudson's locality number] /Paikakariki, Wellington, N.Z., Hudson/ 
Sympetrum bipunctatum Br., var. novae-zealandiae McL. Type/Sympetrum bipunctatum var. 
novaezealandiae McL., $ Lectotype, D. E. Kimmins det. 1967. 

The fourth syntype (from Auckland) has not been traced. The name novaezealandiae was 
originally hyphenated and has been emended in accordance with Art. 32(e)(i). Currently 
placed as a synonym of Diplacodes bipunctata (Brauer). 

obsolescens Kirby (Rhyothemis), 1889 : 321. Holotype <J. Borneo/obsolescens type 

[WFK]. 

occidentalis Tillyard (Nannodythemis), igoSa : 450-453, pi. 6, fig. 3. LECTOTYPE <J. 
Bridgetown, W. A[ustralia], 1.07, R. J. Tilly ard/Narmodythemis occidentalis Till., <$ TYPE. 
R.J.T. /Nannodythemis occidentalis Tillyard, $ Lectotype, D. E. Kimmins det. 1967. 
Currently placed in the genus Nannophya. 

octomaculata Fraser (Palpopleura), 1935. Kimmins, 1966 : 207. 
oculata Fabricius (Libellula), 1775 : 421. Campion, 1917 : 444. 
osiris Fraser (ssp. Zygonyx iris), 1936. Kimmins, 1966 : 208. 

Othello Tillyard (Camacinia), 1908 : 639-641, pi. 14, fig. i. Holotype <$. Cooktown, N.Q., 
xii.07, R. J. TillyardlCzmacmia, Othello Till. <J TYPE, R. J. T. 

paciflcus Kirby (Diplax), 1884 : 355. Holotype <$. Tongatabu, with on reverse, Diplax 
pacificus type [WFK]. 

Currently placed as synonym of Diplacodes bipunctatus (Brauer). 

pallidinervis Kirby (Sympetrum), 1889 : 327, pi. 55, fig. 4. LECTOTYPE <J. Ind[ia]/ 
Pseudothemis pallidinervis Kb. type [WFK] /Sympetrum pallidinervis Kirby, <J Lectotype, 
D. E. Kimmins det. 1967. 

Currently placed as Trithemis pallidinervis (Kirby). Although described as a Sympetrum, 
Kirby labelled his specimens as Pseudothemis. 



LIBELLULID & CORDULIID TYPES IN BM(NH) 291 

parana Kirby (Misagria), 1889 : 339, pis. 52, fig. 9 and 57, fig. 8. Holotype <$. Para/Misagria 

parana type (WFK]. 
parasticta Pinhey (Trithemis), 1955 : 35-37, fig. 8. Holotype $. Lake Chila, Abercorn, 

N. R., 31 .1.54, D. Vesey Fitzgerald /Trithemis parasticta Pinh., 1954. 
phaon Selys (Trithemis), 1883 : 106-107. LECTOTYPE <$. Yokohama, Orio/Type [McL. 

label] /Trithemis phaon Selys <$, Japon [Selys label] /Trithemis phaon Selys, $ Lectotype, 

D. E. Kimmins det. 1967. 

Selys gives no indication as to the collection in which the type was deposited. Ris (1911 : 

589) makes no mention of the type in the Selys Collection and therefore the above specimen 

from the McLachlan Collection has been designated as Lectotype. Currently placed in the 

genus Deielia Kirby. 
phillipsi Kirby (Orthetrum), 1896 : 522. Holotype <. Dobar, 2nd Feby. 95, maritime 

plain/Dobar, Goolis Mts , 2.ii. 1895, L. L[ort-] P[hillips] /Orthetrum Phillipsi Kb. type [WFK]. 
It may be pointed out that the locality data given on the second (printed) label and in the 

description differs from that given on the first (mss) label. The date on the printed label has 

been altered to conform with the mss label. Currently placed as a synonym of Orthetrum 

abbotti (Calvert). 

prateri Fraser (Pseudotramea), 1920. Kimmins, 1966 : 210. 
princeps Kirby (Rhyothemis), 1894 : 16-17. LECTOTYPE <$. Queensland, Turner, 

princeps Kb. <$ type/Rhyothemis princeps Kirby, <J Lectotype, D. E. Kimmins det. 1967. 
pruinosa Kirby (Micrathyrial), 18946 : 267-268. Holotype $ (abd. segments 8-10 missing). 

Grenada/233/pruinosa [WFK]/Micrathyria (?) pruinosa Kirby, $ Holotype, D. E. Kimmins 

det. 1967. 

Currently placed as synonym of Micrathyria didyma didyma (Selys). 

pseudeudoxia Longfield (ssp. Atoconeura biordinata), 1953 : 46, pi. i, figs. 2, E, H. Holo- 
type $. Uganda, Ruwenzori Range, Mubuku Valley, Bikoni Peak, 7,000 ft., 3o.xii.ig37, 

C. E. Longfield /Atoconeura. biordinata pseudeudoxia Type <J, ssp. nov. det. Miss C. Longfield. 
pseudodefecta Pinhey (ssp. Hadrothemis defecta), 1961 : 127-128, pi. 9, fig. 2. Holotype <J. 

Entebbe, Uganda, Oct. 1952, E. PzwAey/Hadrothemis defecta pseudodefecta Pinh. Holotype. 
puella Kirby (Aino), 1890 : 113. LECTOTYPE <J (lacking abd. segs. 9-10). N. pygmaea/ 

Aino puella Kirby, $ Lectotype. D. E. Kimmins det. 1967. 

The only example labelled 'puella' and with type label was the female, but the $ syntype 

was discovered labelled solely 'N. pygmaea'. It agrees well with the original description and 

has been designated the lectotype. The locality of the syntypes was unknown. The taxon 

is currently placed as a synonym of Nannothemis bella Uhler, a South American insect. 
pusilla Kirby (Miathyria), 1889 : 318, pi. 52, fig. 3. LECTOTYPE <J. Amaz[ons]/i42/ 

Miathyria pusilla type [WFK] /Miathyria pusilla Kirby, $ Lectotype, det. D. E. Kimmins, 

1967. 

Kirby gives as localities for this species, Tapajos and Santarem; there are three paralecto- 

types from these localities, one of which is labelled 142 ?. In spite of the fact that the speci- 
men bearing Kirby 's type label has only 'Amaz[ons]' as locality, I have accepted it as lectotype. 

This taxon is currently placed as a synonym of Miathyria simplex (Rambur). 
reducta Fraser (Uracis), 1946. Kimmins, 1966 : 211. 
regalis Tillyard (Agrionoptera), 1908 : 643-645, pi. 14, fig. 3. LECTOTYPE <J. Cooktown, 

N.Q., i.o8, R. J. Tillyard /Agrionoptera. regalis Till., <$ TYPE, RJ.T./Agrionoptera regalis 

Tillyard, $ Lectotype, D. E. Kimmins det. 1967. 

Currently placed as synonym of Agrionoptera longitudinalis Selys. 
rendalli Kirby (Urothemis), 1898 : 234. Holotype ?. Fort Johnston, Nyassaland, P. 

Kendall /rendalli Kirb. paratype [Distant's writing] /Urothemis rendalli Kirby $ TYPE, D. E. 

Kimmins det. 1956. 

As U. rendalli was described from one female, Distant's label 'paratype' must be an error. 

Currently placed as synonym of Urothemis edwardsi (Selys). 



292 D. E. KIMMINS 

[resplendens Selys (Rhyothetnis), 1878 : 300-301.] 

The unnamed variety mentioned by Selys (: 301) is in the BM(NH). Although given a type 
label by McLachlan, it is not considered to be a type. 

reticulata Kirby (Crocothemis), 1886 : 328, pi. 33, figs. 8, 9. Holotype <J. Campbellpore, 
2i.xi.85/N.W. Ind[ia] /reticulata Kirb. type. 

Currently placed as synonym of Crocothemis servilia (Drury). 

rezia Kirby (Aethriamantd), 18890, : 298. Holotype <$ (lacking abd. segs. 5-10). Madag- 
[ascar]/rezia Kb. type. 

risi Campion (Oda), 1915 : 489-491, fig. 40. Holotype <J. Utakwa R., Dutch N. Guin., 
2,500-3,000 ft-, "-1913, A. F. R. Wollaston/Oda nov. spec. Det. Dr. F. Ris/Oda risi Cmpn. 
Holotype, Determined by H. Campion. 

Currently placed in the genus Risiophlebia. 

risi Longfield (Trithemis), 1936 : 490, 491, fig. 8A. Holotype <J. Brit. E. Africa, 3i.iii.n, 
S. A. Neave /Trithemis distanti <J, Dr. F. Ris det./Trithemis risi Longfield, <J Holotype, D. E 
Kimmins det. 1967. 

Longfield (: 491) states 'Lectotype <J and $ have been selected from specimens collected by 
Dr. Neave in British E. Africa in 1911.' As she was describing a new species, in effect she 
designated holotype <$ and allotype $. Pinhey (1962, Publfoes cult. Co. Diam. Angola 59 : 271) 
quotes the Transvaal <J of ambiguus Kirby as lectotype of risi Longfield, but although Long- 
field quotes this specimen in synonymy, she designated a holotype for risi Longfield. 

risi Pinhey (ssp. Porpax asperipes), 1958^ : 115-116. Holotype <J. Chibudzana R., Melsetter 
Dist., 26. ii. 1948, Dept. Agric. S. Rhodesia/Porpax asperipes ssp. risi Pinhey, 1956, Type. 
Subsequently raised to specific rank. 

risi Tillyard (Nannophlebia), 19130, : 713-714, pi. 74, figs. 4, 5, 7. LECTOTYPE <J. Bel- 
linger R., N.S.W., 30.xi.ii, R. J. TiJ/yattZ/Nannophlebia risi Till., $ TYPE, R. J. T./Nanno- 
phlebia risi Tillyard, $ Lectotype, D. E. Kimmins det. 1967. 

robertsi Eraser (Notiothemis), 1944. Kimmins, 1966 : 211. 

rubra Kirby (Trithemis), 1889 : 328. LECTOTYPE <J. Australia/T. rubra type [WFK]/ 
Trithemis rubra Kirby, $ Lectotype, D. E. Kimmins det. 1967. 

As Australia is the first locality mentioned, and Kirby's type label was attached to the 
Australian specimen, this has been chosen as the Lectotype. Currently placed as a synonym 
of Diplacodes haematodes Burmeister. 

ruwenzoriensis Fraser (Tetrathemis), 1941. Kimmins, 1966 : 211. 

sagitta Ris (Orthetrum), 19150 : 216-217. LECTOTYPE <J. Sierra Leone, Port Lokko, 
3. v.i 91 2, /. /. Sim^sow/Orthetrum nov. spec. <J (Type) (near africanum) [Ris' writing]/ 
Orthetrum sagitta Ris, $ Lectotype, D. E. Kimmins det. 1967. 

The penis of the lectotype was dissected and studied by Dr. K. Buchholz, 1963. 

schutnanni Calvert (Micrathyria), 1906 : 225, 227, pi. 8, figs. 39, 40, pi. 9, fig. 24. Holo- 
type (J. Guadaljara, Jalisco, July, Sc/mwaww/Micrathyria schumanni Calv., TYPE. P. P. 
Calvert det. 1906. B. C. A. Neur., p. 227, Orig. pi. viii, ff. 39, 40, pi. ix, fig. 24. 
The <J right hamule has been removed and mounted in canada balsam. 

server Kirby (Trithemis), 1900 : 69-70, fig. i. LECTOTYPE <J (abd. segs 7-10 missing). 
Free Town, Sierra Leone, 5.ix.99, E. E. Austen/On reverse, T. serva K[WFK]. 

Kirby is a little obscure in his account of the material upon which this taxon is based. He 
says 'Described from six specimens, two of them taken by Mr. Austen on Sept. 5 and 21 
(1899).' These two specimens are in the collections; one is labelled Trithemis serva Cotype' 
in Kirby's writing, the other (with a red type label) has '1 . serva K' in Kirby's writing on 
underside of locality label. There is also another Sierra Leone specimen, collected by Morgan, 
1835, labelled 'Trith. serva' by Kirby. Three other Free Town examples, collected by Austen, 
bear no determination labels and I consider them to be excluded from the type series by Kirby's 



LIBELLULID & CORDULIID TYPES IN BM(NH) 293 

statement that two were taken by Austen. One must assume that the other three examples 
are missing. I have therefore chosen the specimen which originally bore the red type label 
as Lectotype. 

siamensis Fraser (ssp. of Onychothemis tonkinensis), 1932. Kimmins, 1966 : 213. 
sietnensi Kirby (Uracis), 1897 : 605-606, pi. 12, fig. 2. LECTOTYPE <J. Gurupa, Lower 

Amazon, 22.ii.96, E. E. Austen/siemensi [WFK] /Uracis siemensi holotypus, prap. Dr. Buch- 

holz/Uracis siemensi Kirby, $ Lectotype, D. E. Kimmins det. 1967. 

The penis of the lectotype was removed and studied by Dr. H. Buchholz on one of his visits 

to BM(NH). The 'female from Para' referred to by Kirby is now labelled allotype and the 

second male (marked cotype by Kirby) has been labelled paralectotype. 

simpsoniRis (Cyanothemis), 1915 : 219-221, fig. Holotype $ (original designation). Yana, 
Sierra Leone, i .iv. 1912, Jas. J. Simpson/Nov. gen. nov. spec.c, Det. Dr. F. Ris/Cyanothemis 
simpsoni Ris, $ Holotype, D. E. Kimmins det. 1967. 

Ris (1919 : 219) refers to this Yana specimen in his description of the sub juvenile <J from 
Mongheri as 'Like first (type) specimen,' and also figures the left wings of the Yana specimen 
( : 220) as '(type $}'. I interpret this as a designation of holotype, although the word holotype 
is not used. 

sita Campion (ssp. of Indothemis limbata), 1923 : 29-31, fig. Holotype $. Ceylon, Yerbury/ 
Trincomali, 6 . xii . go/Indothemis limbata sita Campion, $ Holotype. Determined by H. 
Campion. 

srnithi Fraser (Crocorthetrum), 1921. Kimmins, 1966 : 213. 

stnithi Kirby (Cannacria), 18946 : 266-267. LECTOTYPE <J. [St. Vincent], Grand Sable 
(Windward), by pools near sea, Jan. 4/smithii [WFK] /Cannacria smithi Kirby, <$ Lectotype, 
D. E. Kimmins det. 1967. 

Currently placed as synonym of Brachymesia furcata Hagen. 

stigmata Kirby (Bradinopyga), 1893 : 553-554. pi- 41, fig. 3. LECTOTYPE <J. Ceylon, 
Trincomalee, s.viii.go, Yey&wry/stigmaticum [WFK] /Bradinopyga stigmata Kirby, <J Lecto- 
type, D. E. Kimmins det. 1967. 

stigmatizans Fabricius (Libelluld), 1775 : 421. Campion, 1917 : 443. 

strachani Kirby (Apeleutherus), 1900 : 74-75, pi. 2, figs. 4, 4a. LECTOTYPE <. Lagos, 
Dr. H. Strachan/Apeleuiherus strachani <J/App. figd [last two labels in Kirby 's writing]. 

Of the long series recorded by Kirby, only the lectotype <J and allotype $ bear Kirby's 
determination labels. I have been unable to trace the <J from Free Town, 24-viii, listed by 
Kirby. This taxon is currently placed as Bradinopyga strachani (Kirby). 

subpruinosum Kirby (Sytnpetrutn), 1886 : 326, pi. 33, fig. 7. LECTOTYPE $. N.W. 
Ind[ia] [WFK] /subpruinosum Kirb. type [WFK]/Sympetrum subpruinosum Kirby, $ Lecto- 
type, D. E. Kimmins det. 1967. 

Currently placed as synonym of Sympetrum commixtum Selys. 

sylvia Kirby (Nannothemis), 1889 : 343-344. Holotype $. Beara/N. sylvia type [WFK]. 
Currently placed in the genus Oligoclada. 

terminalis Kirby (Lathrecista), 1889 : 336. LECTOTYPE <J. W. Borneo/Lathrecista 
terminalis $ type [WFK] /Lathrecista terminalis Kirby, $ Lectotype, D. E. Kimmins det. 
1967. 

Currently placed as a synonym of Lathrecista asiatica (Fabr.) 

thais Kirby (Perithemis), 1889 : 324-325. LECTOTYPE <J. Para/3 3o/thais Kb. type 
[WFK]/Perithemis thais Kirby, <$ Lectotype, D. E. Kimmins det. 1967. 
The female, without locality, referred to by Kirby, has not been traced. 

thomasi Longfield (Urothemis), 1932 : 34-35. Holotype <J. S. Arabia, Qara Mts., Ain al 
Rizab, 250 ft., 31 .x. 1930, Bertram Thomas /Urothemis thomasi Longfield, $ Holotype. 
Miss Longfield's determination label is missing, and a new one has been provided. 



294 D - E - KIMMINS 

tibialis Kirby (Micrathyria), 1897 : 610, pi. 13, figs. 5-6. LECTOTYPE <J. Santarem/ 
tibialis [WFK] /Micrathyria tibialis Kirby, $ Lectotype, D. E. Kimmins det. 1967. 

tillyardi Fraser (Palaeothemis), 1923. Kimmins, 1966 : 216. 

torrida Kirby (Pseudotnacrotnia), 1889 : 340-341, pi. 52, fig. 7. LECTOTYPE <J. Sierra 
Leone/Pseudomacromia torrida type [WFK]/Pseudomacromia torrida Kirby, $ Lectotype, 
D. E. Kimmins det. 1967. 

The allotype ? (the only one in Kirby 's series) is from Sierra Leone and is very teneral, the 
head now missing. The remaining four syntypes (all males) from Abyssinia, West Africa, Natal 
and Tenerife, have been traced and are now labelled as paralectotypes. Kirby's statement 
(: 341) 'Notwithstanding these differences, I am not at present inclined to treat any of these 
specimens (all of which are males, the only female being described above) as specifically dis- 
tinct' is considered as definite inclusion of these specimens in the type-series. 

torrida Kirby (Tyriobapta), 1889 : 338-339, pi. 54, figs. 5-6. LECTOTYPE <J. W. Borneo/ 
Tyriobapta torrida type [WFK] /Tyriobapta torrida Kirby, $ Lectotype, D. E. Kimmins det. 
1967. 

trameiformis Kirby (Antidythemis), 1889 : 315, pi. 51, fig. 4. LECTOTYPE <J. Para/ 
Carolina Lin. Cab. /Antidythemis trameiformis <J type [WFK] /Antidythemis trameiformis 
Kirby, $ Lectotype, D. E. Kimmins det. 1967. 

The allotype bears a label Dythemis? trameiformis Selys, n. sp. ?, Amazone? 

translucida Kirby (Tratnea), 1889 : 315. Holotype ?. N. India/translucida Kb. type 

[WFK]. 
This taxon is currently placed as a synonym of Trapezostigma limbata (Desj.). 

triangularis Kirby (Rhyothemis), 1889 : 319. LECTOTYPE <J. Sar[awak] 78/Wallace/ 
triangularis type [WFK] /Rhyothemis triangularis Kirby, <J Lectotype, D. E. Kimmins det. 
1967. 

The example in the BM Type collection, although bearing a type-label, proved to be an 
imposter, bearing a label Sandakan, 29.iii.i893 (four years after publication of the descrip- 
tion). The specimen selected above as type was found in the general collection. It is pre- 
sumed that the false type was transferred to the type collection for evacuation in the summer 
of 1940 and has remained undetected until the preparation of this type list. 

tricolor Kirby (Orthetrum), 1893 : 555-556. Holotype <$. Kandy, 30 . vi . 92/Ceylon, 
Yerbury I Orthetrum tricolor [WFK]. 

trifidum Kirby (Acisoma?), 1889 : 341. Holotype <J (lacking head and right hind wing). 
Congo/Acisoma ascalaphoides? [in pencil, writing unknown] /Acisoma trifida, type [WFK]. 
The spelling of the specific name trifida has been corrected to trifidum. 

tristrigata Kirby (Tetrathemis), 1889 : 343. Holotype ?. Gil./Gilolo/T. tristrigata type 

[WFK]. 

Currently placed as a synonym of Tetrathemis irregularis leptoptera Selys. 

turneri Kirby (Rhyothemis), 1894 : 17. LECTOTYPE <J. Queensland, Turner /Turneri 
Kb. (J [WFK] /Rhyothemis turneri Kirby, <$ Lectotype, D. E. Kimmins det. 1967. 
Currently placed as synonym of Rhyothemis resplendens Selys. 

tyleri Kirby (Trithemis), 1899 : 364, pi. 15, fig. i. LECTOTYPE <J. La Chorrera, Panama/ 
T. Tyleri, Named at sight by W. F. K./Trithemis tyleri Kirby, <$ Lectotype, D. E. Kimmins 
det. 1967. 

uniformis Kirby (Belonia), 1889 : 333-334. Holotype $. Mex[ico]/uniformis, type [WFK]. 
Currently placed as a synonym of Libellula croceipennis Selys. 

variegatum Kirby (Acisoma), 1898 : 239-240. LECTOTYPE <J. Fort Johnston, Nyassaland, 
P. Rendalllva.rieg3.tum Kirby paratype [Distant's writing] /Acisoma variegatum Kirby, 
(J Lectotype, D. E. Kimmins det. 1961. 
Currently placed as a synonym of Acisoma ascalaphoides Rambur. 



LIBELLULID & CORDULIID TYPES IN BM(NH) 295 

venusta Kirby (Micrathyria), 1897 ' 612-613, pi- J 3> fig- * Holotype <J. Macapa, 25.ii.g6/ 
Macapa, Lower Amazon, 25.ii.96/venusta [WFK]. 

This taxon is currently placed in the genus Erythrodiplax . 

vibrans Fabricius (Libellula), 1793 : 380. Campion, 1917 : 449-450. 

vittata Kirby (Urothemis), 1893 : 552, pi. 42, fig. 2. LECTOTYPE $. Trincomali, io.ix.gi/ 
Ceylon, Yerbury/vittats. type [WFK] /Urothemis vittata Kirby, ? Lectotype, D. E. Kimmins 
det. 1967. 

vittatus Kirby (Hydrobasileus), 1889 : 314, pi. 51, fig. 10. Holotype <J. Men[ado]/ Wallace/ 
Tramea 4 vittata type [WFK] /Hydrobasileus vittatus Kirby <J, det. D. E. Kimmins, 1967. 
The name Tramea quadrivittata Hag. MS is quoted by Kirby in synonymy of Hydrobasileus 
vittatus Kirby. 

vivax Calvert (Brechmorhoga), 1906 : 279, 280, pi. 8, fig. 46. Holotype <$. Zapote, 
Guatemala, CAaw^now/Brechmorhoga vivax Calvert, TYPE. P. P. Calvert det. 1906. 
B. C. A. Neur. p. 280. Original of PI. viii, f. 46. 

wilsoni Pinhey (Brachythemis), 1952 : 16, fig. $A. Holotype <J. Xambio, S. Sudan, xii. 
1950, C. E. W^scw/Brachythemis ?sp. nov. E. Pinhey det. igsi/Brachythemis wilsoni Pinh. $. 

woodfordi Kirby (Nesocria), 1889 : 335. Holotype <J. [Solomon Islands], Alu/Nesocria 
woodfordi type [WFK]. 

Currently placed in the genus Protorthemis . 

yerburyi Kirby (Tetrathetnis), 1893 : 556-557, pi. 41, fig. 4. LECTOTYPE <J. Kandy, 
30.vi.g2. B. in coitu A/Ceylon, Yerbury/yerburii type [WFK]/Tetrathemis yerburyi Kirby, 
<fj Lectotype, D. E. Kimmins det. 1967. 
The specific name, spelt yerburii by Kirby, has been emended to yerburyi. 

Family CORDULIIDAE 
aculeata Fraser (Macromia), 1927. Kimmins, 1966 : 176. 

aeneothorax Nunney (Ceratogomphus?) , 1895 : 349-351, figs. A, B. Holotype $ (one sup. 
app. and inf. app. glued to card). Ceratogomphus aeneothorax Nunney mss, on reverse 
'Sierra Leone' and TYPE. 

I do not recognise the writing (in red ink) of the locality 'Sierra Leone'. Currently placed 
in the genus Macromia. 

annaimallaiensis Fraser (Macromia), 1931. Kimmins, 1966 : 178. 

atrifrons McLachlan (Syncordulia), 1883 : xc-xci. Holotype <j>. Queensland/Type [McL. 
label] /Syncordulia atrifrons McL. 

Currently placed in the genus Micromidia. 

atuberculata Fraser (Macromia), 1922. Kimmins, 1966 : 180. 

aurolineata Tillyard (ssp. of Metathemis guttata), 1913 : 575. LECTOTYPE <J. Dorrigo, 
N.S.W., R. J. Tillyard, 4. xii. ign/Metathemis guttata aurolineata Tillyard, TYPE <J, R.J.T./ 
Metathemis guttata aurolineata Till., Lectotype, D. E. Kimmins det. 1968. 
Currently placed as Eusynthemis guttata aurolineata (Tillyard). 

bellicosa Fraser (Macromia), 1924. Kimmins, 1966 : 181. 

berthoudi Tillyard (Hesperocordulia), ign : 376-378, pi. 10, figs, i, 3, g. LECTOTYPE <J. 
Waroona, W. Australia], G. F. Berthoud, 27. xi. igog/Hesperocordulia Berthoudi [sic] Till., 
<J TYPE, R.J.T. /Hesperocordulia berthoudi Till., Lectotype, D. E. Kimmins det. ig68. 

binocellata Fraser (Macromia), 1924. Kimmins, ig66 : 183. 
bispina Fraser (Macromia), 1954. Kimmins, 1966 : 183. 

braueri Selys(Epitheca), 1871 : 284-285. Holotype <J. N[ew] Zeal [and] /24/Cordulia Braueri 
de Selys/Type [McL. label] /Epitheca braueri Selys, Holotype, D. E. Kimmins det. ig67. 
Placed by Fraser, ig3g, in his genus Antipodochlora. 



2 g6 D. E. KIMMINS 

burliyarensis Fraser (ssp. of Idionyx corona), 1922. Kimmins, 1966 : 184. 

canis McLachlan (Tetragoneuria), 1886 : 104-105. Holotype <$. [Western North America] 
Washington Territory \K. H. Morrison] /Type [McL. label] /Tetragoneuria canis McL. 

circularis Tillyard (Pseudocordulia), 1908 : 743, pi. 22, fig. i. Holotype <$. Kuranda, 
N.Q., xii.oy, F. P. )o^/Pseudocordulia circularis Till., $ TYPE, R.J.T. 

claviculata Tillyard (Synthetnis), 19086 : 749-750, pi. 22, fig. 5. Holotype <$. Kuranda, 
N. Q., i.o8, E. Allen/Synihemis claviculata Till., TYPE, R.J.T. 

contutnax Selys (Phyllomacromia), 1879 : 103-104. Holotype $ (right fore wing, both hind 
wings somewhat damaged). West Africa, Akele/Type [McL. label] /Phyllomacromia contu- 
max, Selys [McL. hand]. 

Currently placed in the genus Macromia. 

corona Fraser (Idionyx), 1921. Kimmins, 1966 : 187. 

corycia Laidlaw (Macromia), 1922 : 220, 225, fig. 5. Holotype <J. Sarawak, Ulu Baram 
3.xi.i92o, /. C. Moulton/M-acromia. corycia sp. n. <$ Type, det. F. F. Laidlaw. 

The original paper label, in Laidlaw's handwriting, gives the date wrongly as 3.xi.3O. 

costalis Selys (Cordulia), 1871 : 273. Holotype $. Georgia/Cordulia costalis de Selys/C. 
costalis De Selys $. 

Currently placed as a synonym of Tetragoneuria cynosura (Say). 

cupricincta Fraser (Macromia), 1924. Kimmins, 1966 : 187. 
cupricolor Fraser (Hemicordulia), 1927. Kimmins, 1966 : 187. 

cyanitincta Tillyard (Synthemis), 1907 : 724-726, pi. 35, figs. 3-4, pi. 36, fig. i. LECTO- 
TYPE cJ. Margaret R., W. A., 1.07, R. J. Tillyard /Synthemis cyanitincta Till., <$ TYPE, 
R.J.T. /Synthemis cyanitincta Tillyard, $ Lectotype, D. E. Kimmins det. 1967. 

cydippe Laidlaw (Macromia), 1922 : 220-222, 223, figs. 2, 3. Holotype <$. Sarawak, Lio 
Matu, 3. x. 1914, /. C. Moulton/Macromia, cydippe Laidlaw, $ Type. 

cyrene Lieftinck (Synthemis), 19530 : 78-81, figs. 5-9. Holotype <J. N. E. New Guinea, 
Saiko, 5,500-6,000 ft., Babu River (Upper Waria River), ix-x. ig^6/F. Shaw-Mayer/Synthemis 
cyrene sp. n. <$. 

divergens Tillyard (Cordulephya), 1917 : 467-469, pi. 23, figs. 9-10, text-fig. 6. LECTO- 
TYPE $. Hornsby, N.S.W., 8.v.i6, R. J. Tillyard /Cordulephya. divergens Till., ^ TYPE, 
R.J.T. /Cordulephya divergens Tillyard, <$ Lectotype, D. E. Kimmins det. 1967. 

donaldi Fraser (Indomacromia), 1924. Kimmins, 1966 : 189. 

elliptica Tillyard (Pseudocordulia), 19136 : 229-230, pi. 15, figs. 1-2. LECTOTYPE <?. 
Kuranda, N. Queensland], F. P. Dodd, 3.i. igis/Pseudocordulia elliptica Tillyd., TYPE, 
R.J.T. /Pseudocordulia elliptica Till., $ Lectotype, D. E. Kimmins det. 1968. 

ellisoni Fraser (Macromia), 1924. Kimmins, 1966 : 190. 

elongata Campion (Metaphya), 1921 : 64-66, pi. 9, fig. 17. Holotype ?. Loc. Baie Ngo, 
no. 204, 10.11.14. Coll. P. D. Montague, New Caledonia Exped./New Caledonia, P. Z>. 
Montague /Metaphy a. elongata Campion, Holotype, Determined by H. Campion. 

euterpe Laidlaw (Macromia), 1915 : 26-29, figs. 1-2. LECTOTYPE $. Kina Balu, British 
North Borneo, /. C. Moulton/M.. euterpe Type <J, I2.ix.i3, 7 [Laidlaw's writing]/Macromia 
euterpe Laidlaw, 5* Lectotype, D. E. Kimmins det. 1967. 

The example chosen as Lectotype has for many years been labelled 'Type H.T.', but was not 
so designated in print by Laidlaw. 

evelynae Lieftinck (Synthemis), 1953^ : 81-84, fig 3 - io-ii. Holotype <$ (abd. segs. 6-10 
and anal appendages wanting). Papua, Mafulu, 4,000 ft., xii. 1933, L. E. Cheesman/Synihemis 
evelynae sp. n. $. 



LIBELLULID & CORDULIID TYPES IN BM(NH) 297 

fallax McLachlan (Gomphomacromia), 1881 : 141-142. LECTOTYPE <J. Intaj, Ecuador/ 
Gomphomacromia fallax McL./Type [McL. label] /Gomphomacromia fallax McL., Lectotype, 
D. E. Kimmins det. 1967. 

fenella Campion (Synthesis), 1921 : 61-62, pi. 9, fig. 16. text-fig, n. Holotype $. Loc. 
Mt. Mou, No. 725, Date 20.111.14. Coll. P. D. Montague, New Caledonia Exped./Synthemis 
fenella Campion, Holotype. Determined by H. Campion. 

fldelis McLachlan (Hemicordulia), 1886 : 104. Holotype $. [Loyalty Isls.] Uvea/Type 
[McL. label] /Hemicordulia fidelis McLach. 

flavicauda McLachlan (Nesocordulia), 1882 : clxx-clxxii. Holotype $. Madagascar/ 
Nesocordulia flavicauda McL./Type [McL. label]. 

flavicincta Kirby (Phyllomacromia), 1898 : 241-242. LECTOTYPE <J. Rustenberg, 
W. L. D[istant] /flavicincta (type) Kirby [Distant's writing] /Macromia picta Selys H. C[ampion]/ 
Phyllomacromia flavicincta Kirby, <$ Lectotype, D. E. Kimmins det. 1967. 

Currently placed in the genus Macromia, where it is a junior homonym of M. flavicincta 
Selys and a synonym of Macromia picta (Selys) . 

flavicincta Selys (Macromia), 1874 : 25-27. LECTOTYPE <J. North Ind[ia]/Macr. 
flavicincta de Selys/Macromia flavicincta Selys, <J Lectotype, D. E. Kimmins det. 1967. 

The male chosen as Lectotype has for may years been labelled Holotype, but it was not so 
indicated in print by Selys. The original description gives 'Nord du Bengale' as locality, but 
it is only the $ which is so labelled. 

flavocolorata Fraser (Macromia), 1922. Kimmins, 1966 : 192. 
flavovittata Fraser (Macromia), 1935. Kimmins, 1966 : 192. 

flexicauda Campion (Synthemis), 1921 : 57-61, pi. 8, fig. 14, pi. 9, fig. 15, text-fig. 10. Holo- 
type (J. Loc. Mt. Nekando, Date, 24^.14, Coll. P. D. Montague, New Caledonia Exped./ 
New Caledonia, P. D. Montague /Synthemis flexicauda Campion (J. Holotype. Determined 
by H. Campion. 

fulva Laidlaw (Macromidia), 1915 : 29-30, fig. 3. Holotype cJ. Borneo, /. C. Moultonj 
Macromidia fulva, Q* Laidlaw, TYPE. Kina Balu, Borneo, Ii.ix.i9i3, J. C. Moulton. 

fulvia Fraser (ssp. of Idionyx corona), 1924. Kimmins, 1966 : 193. 
galeata Fraser (Idionyx), 1924. Kimmins, 1966 : 193. 

genialis Laidlaw (Macromidia), 1923 : 232. Holotype $. Malay Peninsula, Pahang, 
Gunong Tahan, lo.xii. 1921 /Macromidia genialis $ Type [Laidlaw's writing]. 

gomphomacromioides Tillyard (Synthemiopsis), 1917 : 464-467, pi. 23, figs. 7-8, text-figs 
4-5. LECTOTYPE <J. Cradle Mt., Tas., R. J. Tillyard, i8.i. 1 7/Synthemiopsis gomphoma- 
cromioides Till., o* TYPE, R.J.T. /Synthemiopsis gomphomacromioides Tillyard, $ Lectotype, 
D. E. Kimmins det. 1967. 

gracillima Fraser (Hemicordulia), 1944. Kimmins, 1966 : 194. 

halei Fraser (Macromia), 1928. Kimmins, 1966 : 194. 

ida Fraser (Macromia), 1924. Kimmins, 1966 : 196. 

imbricata Fraser (Idionyx), 1926. Kimmins, 1966 : 196. 

indica Fraser (Macromia), 1924. Kimmins, 1966 : 197. 

insignis Kirby (Pseudogomphus), 18890 2 99- Holotype <J. Cameroons/Pseudogomphus 
insignis [WFK]. 

Currently placed as a synonym of Macromia sophia Selys. The tenth abdominal degment 
and anal appendages are detached. 

insularis Kimmins (Guadalca), 19570 : 318-319, figs. 3-4. LECTOTYPE 6*. Solomon Is., 
Guadalcanal, Tapenanje, 10-23. xii. 1953, /. D. Bradley/Guadalca. insularis Kim., ^ Type, 
D. E. Kimmins det. 1955/Guadalca insularis Kimmins, < Lectotype, D. E. Kimmins det. 1967. 



agS D. E. KIMMINS 

intricata Fraser (Idionyx), 1926. Kimmins, 1966 : 198. 
irata Fraser (Macromia), 1924. Kimmins, 1966 : 198. 

junior Pinhey (ssp. of Macromia kimminsi), 1961 : 116, pi. 8, figs. 10, 15, 16. Holotype <$. 
van Someren, Broderick Falls, Scarp, Kenya, v. 51 /Macromia kimminsi junior Pinh. HOLO- 
TYPE. 

kimminsi Fraser (Macromia), 1954. Kimmins, 1966 : 199. 

kimminsi Lieftinck (Synthemis), 1953^ : 75-78, figs. 1-4. Holotype <$. N.E. New Guinea, 
Saiko, 5,500-6,000 ft., Babu River (Upper Waria River), 24. x. ig^d/F. Shaw Mayer/Synihemis 
kimminsi sp. n. 

laidlawi Fraser (Idionyx), 1936. Kimmins, 1966 : 200. 

In my paper (1966), the reference to this species was omitted. It should be 19360 : 701. 

leachi Selys (Synthemis), iSjia : 561. Holotype <$. N[ew] Holland/Cord. Leachii $ Selys/ 
Synth. Leachii de Selys. 

The left hind wing is missing and the other wings are somewhat battered. The specific 
name has been emended to 'leachi'. 

teoni Fraser (Phyllomacromia), 1928. Kimmins, 1966 : 200. 
lieftincki Fraser (Macromia), 1954. Kimmins, 1966 : 201. 

magniflca Selys (Macromia), 1874 : 22-24. LECTOTYPE ?. N. California, Walsinghamj 
Macromia magnifica $ [Selys' writing] /Macromia magnifica Selys, $ Lectotype, D. E. Kimmins 
det. 1967. 

The <J syntype of this species has not been traced in the McLachlan collection. The $ was 
found (without any type label) and has been designated Lectotype. Selys gives no indication 
that he retained the <$, and Martin (1906 : 67) writes 'Types <? et $ coll. MacLachlan'. 

malabarensis Fraser (Epophthalmia), 1935. Kimmins, 1966 : 203. 

malayana Laidlaw (ssp. of Macromia moorei), 1928 : 133-134. Holotype $. Pahang, 
L. Tamang, 13. vi. 1923, F. N. Chasen/Macromia. moorei malayana n. subsp. [Laidlaw's 
writing] /Macromia moorei malayana <J Holotype, Laidlaw [DEK]. 
Currently placed as synonym of Macromia moorei fumata Kruger. 

marginata Selys (Somatochlora?), 1883 : 109-110. Holotype <$ (head lacking) . Japan/Type 
[McL. label]/Somatochlora? marginata Selys, <$, Japon [Selys' writing]. 
Currently placed in the genus Epitheca. 

[martini Tillyard (Synthemis), 1907 : 726-728, pis. 35, figs. 1-2 and 36, fig. 2.] 

There are no examples of this species in BM(NH) collection. The types may perhaps still 
be in Canberra. 

mascarenica Fraser (Nesocordulia), 1948. Kimmins, 1966 : 203. 

melanosoma Tillyard (ssp. of Metathemis guttata), 19136 : 231-233. LECTOTYPE g. 
Tambourine, Queensland], R. J. Tillyard, 24.xii. 1912/Metathemis guttata melanosoma 
Tillyd., (J TYPE, R.J.T. /Metathemis guttata melanosoma Till., <$ Lectotype, D. E. Kimmins 
det. 1968. 

Currently placed as Eusynthemis guttata melanosoma (Tillyard). 

metallica Tillyard (Lathrocordulia), 1911 : 379-380, pi. 10, figs. 2, 4 and 10. LECTOTYPE 
<. Waroona, W. A[ustralia], G. F. Berthoud, 3-xii. igio/Lathrocordulia metallica Till., 
6* TYPE, R.J.T. /Lathrocordulia metallica Till., ^ Lectotype, D. E. Kimmins det. 1968. 
The lectotype lacks the apex of the left superior anal appendage. 

micans Laidlaw (Metaphya), 1912 : 2-3, pi. i. Holotype <$. Sarawak, igio/Metaphya 
micans Type $ [Laidlaw's writing]. 

The allotype $ was described later (Proc. zool. Soc. Lond. 1913 : 65-67). 

minima Fraser (Idionyx), 1931. Kimmins, 1966 : 204. 



LIBELLULID & CORDULIID TYPES IN BM(NH) 299 

montaguei Campion (Synthemis), 1921 : 55-57, pi. 8, fig. 13, text-fig. 9. Holotype <J. Loc. 
Alt Mou, Date, io.iii.i4 (No. 488), Coll. P. D. Montague, New Caledonia Exped./New Cal- 
edonia, P. D. Montague I Synthemis montaguei Campion, $, Holotype. Determined by H. 
Campion. 

montana Tillyard (Cordulephya), igna : 392-397, pis. 11-12. Hclotype <$. [Medlow], 
Blue Mts., N.S.W., ig.i.io, R. J. Tillyard /Cordulephya, montana Till., ^ TYPE, R.J.T. 

Holotype <J specified by Tillyard (1911 : 397, line 14, 'type-male'). The locality 'Medlow' 
is given in the text in several places but not on the locality label. 

moorei Selys (Macromia), 1874 : 28. Holotype $. Himalaya/Macromia moorei Selys $, 
n. sp. 

mystica Tillyard (Austrophya), 19086 : 739-740, pi. 31, fig. i. Holotype $. Kuranda, 
N. Q., i.o8, F. P. Doddl Austrophya mystica Till., TYPE ?, R.J.T. 

nadganiensis Fraser (Idionyx), 1924. Kimmins, 1966 : 205. 

nasalis Selys (Epitheca), 1874 : 21. Holotype ?. North America/Epith. nasalis de Selys mss., 
var. d'albicincta B.? [Selys' writing]. 

This example has at some time lost its locality label and a new one has been provided. 
Currently placed in the genus Somatochlora. 

nigra Tillyard (Synthemis), 1906 : 489-491. LECTOTYPE <$ (lacking most of left hind wing). 
Kuranda, N. Q., 1.05, R. J. Tillyard /Synthemis nigra Till. ^ TYPE, R.J.T./Synthemis nigra 
Tillyard, <$ Lectotype, D. E. Kimmins det. 1967. 
Currently placed in the genus Eusynthemis. 

nilgiriensis Fraser (ssp. of Idionyx corona), 1922. Kimmins, 1966 : 206. 
nilgiriensis Fraser (Phyllomacromia), 1918. Kimmins, 1966 : 207. 

occidentalis Tillyard (ssp. of Synthemis macrostigma), 1910 : 355. LECTOTYPE <J. 
Bridgetown, W. A., i.oj, R. J. Tillyard /Synthemis macrostigma Selys, subsp. occidentalis 
Till., <J TYPE, R.J.T./Synthemis macrostigma occidentalis Till., $ Lectotype, D. E. Kimmins 
det. 1967. 

olivet Tillyard (Synthemis), 19086 : 747-749, pi. 22, fig. 4. LECTOTYPE <J. Cooktown, 
N. Q., i.o8, R. J. Tillyard /Synthemis olivei Till., ^ TYPE, R.J.T./Synthemis olivei Tillyard, 
(J Lectotype, D. E. Kimmins det. 1967. 
Currently placed in the genus Choristhemis. 

orientalis Tillyard (ssp. of Synthemis macrostigma), 1910 : 355. LECTOTYPE <J. Blue 
Mts., N.S.W., ii.05, R. J. Tillyard /Synthemis macrostigma Selys, subsp. orientalis Till., $ 
TYpE/Synthemis macrostigma orientalis Till., $ Lectotype, D. E. Kimmins det. 1967. 

ornata Fraser (Idionyx), 1921. Kimmins, 1966 : 208. 
pacifica Fraser (Hemicordulia), 1925. Kimmins, 1966 : 208. 
pallida Fraser (Macromia), 1924. Kimmins, 1966 : 208. 
periyashola Fraser (Idionyx), 1939. Kimmins, 1966 : 209. 

piscator Barnard (Presba), 1934 : 168, figs, if-h, 2. LECTOTYPE <J. Groot Drakenstein, 
Cape Province, xii.i932, A. C. Harrison/Presba. piscator Barnard $ TYPE [DEK]/Presba 
piscator Barnard, Lectotype, D. E. Kimmins det. 1967. 

pseudafricana Pinhey (Macromia), 1961 : 109, pi. 8, fig. 5. Holotype <J. Aswe R., Kara- 
moja Suk, Uganda, iv. 1952/Macromia africana Selys, E. Pinhey det. 1952/Macromia africana 
Selys, F. C. Fraser det. 1953/Macromia pseudafricana Pinh. HOLOTYPE. 
Pinhey states that Fraser 's determination was made 'with reservation'. 

pygmaea Selys (Cordulephya), 1871 : 316. Holotype <J. Melbourne, Edwards/ 122 /Cordu- 
lephia pygmaea Selys <J/Type [McL. label]. 



300 D. E. KIMMINS 

refracta Tillyard (Austrocordulia), 19086 : 744-745, pi. 22, fig. 2. Holotype <$. Cooktown, 
N. Q., i.o8, R. J. Tillyard /Austrocordulia refracta Till., <J TYPE, R.J.T. 

regina Selys (Synthemis), 1874 : 31-33. LECTOTYPE <J. N[ew] S[outh] W [ales] /Synth, 
regina de Selys/Synthemis regina Selys, Lectotype, D. E. Kimmins det. 1967. 

Selys gives as locality 'Queensland', but the BM(NH) specimen, which he lists first, is 
labelled New South Wales, the MacLachlan examples being from Queensland. Currently 
placed in the genus Eusynthemis. 

rhinoceroides Eraser (Idionyx), 1934. Kimmins, 1966 : 211. 

rutherfordi Selys (Neophya), 1881 : xvii-xviii, i fig. Holotype <. Old Calabar, R[uther- 
ford]/Type[McL. label] /Neophya Rutherfordi Selys. 

saffronata Eraser (Idionyx), 1924, Kimmins, 1966 : 212. 

selysi Eraser (Idionyx), 1926 : 201-202, pis. 9, fig. 6 and 10, fig. 6. LECTOTYPE ^. I. 
selysi <?. Maymyo, Upper Burma, I5.vi.24. Col. F. Wall coll. [F.C.F.J/Idionyx selysi 
Eraser. < Lectotype, D. E. Kimmins det. 1967. 

Through an unfortunate oversight, this species was omitted from my 1966 paper. The 
above example has been chosen as Lectotype, in preference to the example in the Selys collec- 
tion, which Eraser appears not to have seen. 

selysi Kirby (Macromia), 1900 : 77-78. Holotype <J. Sierra Leone, Free Town, i6.ix.99, 
E. E. Austen/U. Selysi type [WFK]. 

shanensis Fraser (Macromedia), 1927. Kimmins, 1966 : 213. 

smithi White (Cordulia), 1845, pi. 6, fig. 2; Selys, 1871 : 261. Holotype $. New Zealand/ 
Cordulia Smithii White, Zool. Ereb. & Terror, t. . ., f. . . [A. White's writing]. 

The first reference to this species is the plate reference given above. The first description 
is that of Selys, 1871. Currently placed in the genus Procordulia. 

sophia Selys (Macromia), 18710, : 550-551. Holotype $. Cape Coast Castle [Guinea]/i24/ 
Type [McL. label] /Macromia sophia $ Selys [Selys' writing]. 

spiniger Tillyard (Synthemis), 1913 : 572-574, pi. 62, figs. 15-16. Holotype $. Waroona, 
W. Australia], G. F. Berthoud, 27.1. 191 2/Synthemis spinifer [sic] Tillyard, TYPE <J, R.J.T./ 
Synthemis spiniger Till., <J Holotype, D. E. Kimmins det. 1968. 

The specific name was published as 'spiniger' and used consistently five times in the original 
publication. 

stevensi Fraser (Idionyx), 1924. Kimmins, 1966 : 214. 

subjuncta Tillyard (ssp. of Metathemis brevistyla), 1913 : 574, pi. 62, fig. ib. LECTOTYPE 
<. Dorrigo, N.S.W., R. J. Tillyard, 6. xii. 1911 /Metathemis brevistyla subjuncta Tillyard, 
TYPE <J, R.J.T. /Metathemis brevistyla subjuncta Till., $ Lectotype, D. E. Kimmins det. 1968. 
Currently placed as Eusynthemis brevistyla subjuncta (Tillyard). 

superba Tillyard (Hemicordulia) , 1911 : 368-370, pi. 10, figs. 5-8. LECTOTYPE 6*. Fallal, 
N.S.W., R. J. Tillyard, 13. xii. igio/Hemicordulia superba Till., <$ TYPE, R.J.T. /Hemicordulia 
superba Till., $ Lectotype, D. E. Kimmins det. 1968. 

sylvatica Fraser (Macromia), 1954. Kimmins, 1966 : 215. 

tasmanica Tillyard (Synthemis), 1910 : 344-347, pis. 4, fig. 2 and 7, figs. 2, 20. LECTO- 
TYPE <J. St. Patrick's R., Tas., 7-i.og, R. J. Tillyard /Synthemis tasmanica Till., <J TYPE, 
R.J.T. /Synthemis tasmanica Till., <$ Lectotype, D. E. Kimmins det. 1967. 

thalia Lieftinck (Macromia), 1929 : 103-104, fig. 29. Holotype $. Khasia Hills/Macromia 
thalia nov. nom., $ Holotype, vid. Lieftinck, '28 [Lieftinck's writing]. /Macromia thalia 
Lieftinck, <J[DEK]. 

This specimen was originally determined as Macromia Ifraenata Laidlaw, by Laidlaw. 



LIBELLULID & CORDULIID TYPES IN BM(NH) 301 

thetis Ris (Macromia), 1921 : 379-381, figs. 65-66. LECTOTYPE <J. Chirinda For., 
Gazal[and], 3,600 ft., 9. x. 1905, G. A. K. Mar shall /Macromia spec, a, Det. Dr. F. Ris, Phyl- 
lomacromia aequatorialis Martin ?/Macromia thetis Ris <J, det. Miss C. Longfield, 19347 
Macromia thetis Ris, <$ Lectotype, D. E. Kimmins det. 1968. 

With this type and the allotype $ is a mss. note by Miss Longfield. 'These are believed to 
be the types of Macromia thetis Ris, described 1921, in Ann. S. African Mus. Found with all 
labels as at present, except type labels and determination labels of Miss C. Longfield'. 

The female in the South African Museum was excluded from the type series by Ris' reference 
(p. 381) to the 'typical pair in the British Museum'. 

tillyardi Martin (Macromia), 1906 : 72. Holotype $. Kuranda, i.o5/Macromia tillyardi 
Martin, $ TYPE, R.J.T. 

Although Tillyard (1906 : 491) records the capture of three females, only one, 'the type- 
specimen' was sent to Martin and is therefore automatically the holotype 

travencorensis Fraser (Idionyx), 1931. Kimmins, 1966 : 216. 
trituberculata Fraser (Macromia), 1921. Kimmins, 1966 : 216. 
unguiculata Fraser (Idionyx), 1926. Kimmins, 1966 : 217. 

venator Barnard (Presba), 1933 : 167, figs. la-e. LECTOTYPE . French Hoek, C[ape] 
P[rovince], with on reverse, K. H. Barnard, xii. 1932/Presba venator Barnard, <$ TYPE [DEK]/ 
Presba venator Barnard, Lectotype, D. E. Kimmins det. 1967. 
Currently placed as a synonym of Syncordulia gracilis (Burm.). 

virgula Selys (Synthemis), 1874 : 33-34. LECTOTYPE <J. S. Australia], Bake well/Synth. 
virgula de Selys/Synthemis virgula Selys, <$ Lectotype, D. E. Kimmins det. 1967. 
Currently placed in the genus Eusynthemis. 

viridescens Tillyard (Macromia), 1911 : 380-381, pi. 10, fig. n. Holotype <J. Cape York, 
N. Queensland], H. Elgner, 22. xi. igog/Macromia viridescens Till., $ TYPE, R.J.T. /'This is 
said to be the $ of M. terpsichore Foerster. Tillyd., Proc. Linn. Soc. N.S.W., Vol. xxxvii, 
1912, p. 584' [F. C. Eraser's writing]. 
Currently placed as a synonym of Macromia terpsichore Foerster. 



whitei Selys (Macromia), iSjia : 555. LECTOTYPE <J. N. India/Macr. White! <J De Selys 
[Selys' writing] /Macromia whitei Selys, <J Lectotype, D. E. Kimmins det. 1967. 

The BM(NH) collection contains i $, i $ of this species, labelled by Selys in writing very 
similar to that on the label of Synthemis leachi, described in the same publication. Selys quotes 
an unspecified number of examples in 'Coll. Selys et Brit. Mus.' Our examples are labelled 
N. India, which would certainly include part of Bengal. The species is almost certainly named 
after Dr. Adam White, a previous Keeper of the Zoological Department of the British Museum 
and one assumes therefore that our examples would be the types. Martin (1906 : 70-71) 
mentions types of M. cingulata Rambur (of which whitei is a synonym) in the Selys Collection, 
but not of whitei. I do not know of any previous designation of a type for this species and 
have therefore selected our $ (as above) as Lectotype. 

wollastoni Campion (Synthemis), 1915 : 486-488, figs. 38-39. Holotype $. Dutch New 
Guinea, A. F. R. Wollaston/Snow Mts. 4,000 to 6,000 ft., i-ii. I9i3/Synthemis wollastoni 
Cmpn., $ Holotype, Determined by H. Campion. 

The description gives the additional data, Utakwa River, not mentioned on the printed 
labels. 

xanthosticta Tillyard (ssp. of Metathemis nigra), 19136 : 230-231. LECTOTYPE <J. 
Tambourine, Queensland], R. J. Tillyard, 5-i. 1913/Metathemis nigra xanthosticta Till., 
< Lectotype, D. E. Kimmins det. 1968. 

Currently placed as Eusynthemis nigra xanthosticta (Tillyard). 

zeylanica Fraser (Macromia), 1927. Kimmins, 1966 : 219. 



302 



D. E. KIMMINS 



REFERENCES 
BARNARD, K. H. 1933. A new genus of Corduline Dragonfly from South Africa (Odonata). 

Stylops 2 (7) : 165-168, figs. 1-2. 
CALVERT, P. P. 1901-1908. Odonata in Biologia Centrali- Americana: 17-420, pis. 2-10. 

London. 
CAMPION, H. 1915. Report on the Odonata collected by the British Ornithologists' Union 

Expedition and the Wollaston Expedition in Dutch New Guinea. Trans, zool. Soc. Land. 

20 (15) : 485-492, text-figs. 38-40. 
1917. On Fabricius's Types of Odonata in the British Museum (Natural History). Ann. 

Mag. nat. Hist. (8) 19 : 441-450. 
1921. Odonata collected in New Caledonia by the late Mr. Paul D. Montague. Ann. Mag. 

nat. Hist. (9) 8 : 33-67, pis. 8-9, text-figs, i-i i . 

- 1923. A new form of Indothemis limbata from Ceylon (Odonata). Ann. Mag. nat. Hist. 
(9) 11 : 28-31, i text-fig. 

FABRICIUS, J. C. 1775. Systema Entomologiae sistens Insectorum Classes, Ordines, Genera, 

Species, etc. pp. 420-426. Flensburgi et Lipsiae. 
1781. Species Insectorum exhibentes eorum differentias specificas, synonyma auctorum, loca 

natalia, metamorphos, etc., 1 : 519-528. Hamburgi et Kilonii. 

1787. Mantissa Insectorum etc. 1. Hafniae. 

I 793- Entomologia Systematica emendata et aucta, 2 : viii + 519. Hafniae. 
FRASER, F. C. 1926. A revision of the genus Idionyx Selys. Rec. Indian Mus. 28 : 195-207, 

pis. 8-10. 
GARDNER, A. E. 1958. Aspects of the fauna and flora of the Azores, xii(a). Odonata. 

Ann. Mag. nat. Hist. (13) 1 : 791-792. 
KIMMINS, D. E. 1936. Odonata, Ephemeroptera and Neuroptera of the New Hebrides and 

Banks Island. Ann. Mag. nat. Hist. (10) 18 : 68-88, n text-figs., pi. 3. 
1936^. The Odonata of the Oxford University Sarawak Expedition. /. fed. Malay St. 

Mus. 18 (i) : 65-108, 17 text-figs. 
1942. The type of Allorhizucha campioni Ris (Odonata, Libellulinae) . Entomologist 

75 : 47. 
1943- A new South American Dragonfly. Ann. Mag. nat. Hist, (n) 10 : 156-159, figs. 

1-6. 
1953. Miss L. E. Cheesman's Expedition to New Caledonia, 1949 Orders Odonata, 

Ephemeroptera, Neuroptera and Trichoptera. Ann. Mag. nat. Hist. (12) 6 : 241-257, 24 

text-figs. 
1957- Some notes on nomenclature in the family Libellulidae (Odonata). Entomologist 

90 : 95-97- 

- 1958. New species and subspecies of Odonata. Bull. Br. Mus. nat. Hist. (Ent.) 7 (7) : 
347-358, 7 text-figs. 

- 1966. A list of the Odonata Types described by F. C. Fraser, now in the British Museum 
(Natural History). Bull. Br. Mus. nat. Hist. (Ent.) 18 (6) : 173-227. 

KIRBY, W. F. 1884. On the Neuroptera collected during the recent expedition of H.M.S. 

Challenger. Ann. Mag. nat. Hist. (5) 13 : 453-456. 
1886. On a small collection of Dragonflies from Murree and Campbellpore (N.W. India), 

received from Major J. W. Yerbury, R.A. Proc. zool. Soc. Lond. 1886 : 325-329, pi. 33. 
1889. A revision of the subfamily Libellulinae, with descriptions of new genera and 

species. Trans, zool. Soc. Lond. 12 (9) : 249-348, pis. 51-57. 

- i88ga. Descriptions of new genera and species of Odonata in the collection of the British 
Museum, chiefly from Africa. Proc. zool. Soc. Lond. 1889 : 297-303. 

- 1890. Description of a new species of Dragonfly. Ann. Mag. nat. Hist. (6) 5 : 112-113. 
1893. Catalogue of the described Neuroptera Odonata (Dragonflies) of Ceylon, with 

descriptions of new species. /. Linn. Soc., Zool. 24 : 545-566, pis. 41-42. 
1894. On a small collection of Odonata (Dragonflies) from Queensland, with descriptions 



of five new species. Ann. Mag. nat. Hist. (6) 14 : 15-21. 



LIBELLULID & CORDULIID TYPES IN BM(NH) 303 

KIRBY, W. F. 18940. Descriptions of a new species of Dragonfly (Dythemis Broadwayi) from 
Trinidad. Ann. Mag. nat. Hist. (6) 14 : 227-228. 

18946. On some small collections of Odonata (Dragonflies) recently received from the 

West Indies. Ann. Mag. nat. Hist. (6) 14 : 261-269. 

i8g4C. Notes on a small collection of Odonata etc. from Upper Burma, with the descrip- 
tion of a new species. Ann. Mag. nat. Hist. (6) 14 : 111-113. 

1896. On some Dragonflies obtained by Mr. and Mrs. Lort-Phillips in Somaliland. Proc. 
zool. Soc. Land. 1898 : 521-523. 

1897. List of the Neuroptera collected by Mr. E. E. Austen on the Amazons etc. during 
the recent expedition of Messrs. Siemens Bros. Cable S.S. Faraday, with descriptions of 
several new species of Odonata (Dragonflies). Ann. Mag. nat. Hist. (6) 19 : 598-617, pis. 
12-13. 

1898. On a collection of Dragonflies from the Transvaal and Nyasaland. Ann. Mag. 

nat. Hist. (7) 2 : 229-245. 

1899. On a collection of Odonata (Dragonflies) from Panama. Ann. Mag. nat. Hist. 
(7) 3 : 362-371, pi. 15. 

1900. Report on the Neuroptera Odonata collected by Mr. E. E. Austen at Sierra Leone 
during August and September 1899. Ann. Mag. nat. Hist. (7) 6 : 67-79, pi. 2 and 2 text 
figs. 

19000. On a small collection of Odonata (Dragonflies) from Hainan, collected by the late 
John Whitehead. Ann. Mag. nat. Hist. (7) 5 : 530-539, pi. 12. 

1905. List of a small collection of Odonata (Dragonflies) from Ceylon, collected by Mr. E. 
Ernest Green, with notes on the genus Zygonidia and its allies, and descriptions of new 
species of Zygonidia Kirb, and Onychothemis Brauer, from Ceylon and Tonkin. Ann. Mag. 
nat. Hist. (7) 15 : 270-278. 

19050. List of a collection of Neuroptera Odonata (Dragonflies) formed by G. A. K. 
Marshall, Esq., at Salisbury, Mashonaland, with descriptions of a new genus and two new 
species. Ann. Mag. nat. Hist. (7) 15 : 190-193. 

1909. Ruwenzori Expedition Reports. 7. Neuroptera. Trans, zool. Soc. Lond. 19 (i) : 
59-62. 

LAIDLAW, F. F. 1912. On a new genus and species of Odonata from Sarawak. Sarawak Mus. 
J. 1 (2) : 1-3, pi. i. 

1915. Contributions to a study of the Dragonfly fauna of Borneo. Part III. A collection 
made on Mount Kina Balu by Mr. J. C. Moulton in September and October, 1913. Proc. 
zool. Soc. Lond. 1915 : 23-39, text-figs. 1-5. 

1922. Notes on some oriental Dragonflies: the genus Macromia. Journ. Straits Br. R. 
Asiat. Soc. 85 : 218-229, 7 figs. 

1923. A survey of the dragonfly fauna of the Malay Peninsula, with notes on that of 
neighbouring countries. Part I. /. Malayan Br. R. Asiat. Soc. 1 : 319-333, figs. 

1928. Notes on Oriental Dragonflies (Odonata), with descriptions of new species. Proc. 



zool. Soc. Lond. 1928 : 129-138, 3 text-figs. 
LIEFTINCK, M. A. 1929. A revision of the known malaysian Dragonflies of the genus Mac- 
romia Rambur, with comparative notes on species from neighbouring countries and des- 
criptions of new species. Tijdschr. Ent. 72 : 59-108, 22 figs. 

1953. Revisional notes on the genera Diplacina Brauer and Huonia Forster (Odon.). 
Treubia 22 (i) : 153-216, 23 text-figs. 

I953- Revision of the Australasian species of Synthemis Selys (Odon. Corduliidae), with 
descriptions of four new species and a key to their identification. Idea 9 (3-4) : 70-88, 
14 figs. 

1955. Two new species of Lanthanusa Ris, from the high mountains of New Guinea 
(Odonata). Zool. Meded. Leiden 33 (18) : 157-164, 9 text-figs. 

LONGFIELD, C. 1932. A new species of the genus Urothemis from Southern Arabia, and some 
comments on the species of Odonata inhabiting the Cjara Mountains. Stylops 1 : 34-35. 

1936. Studies on African Odonata, with synonymy and descriptions of new species and 

subspecies. Trans. R. ent. Soc. Lond. 85 : 467-498, 10 text-figs. 



304 D. E. KIMMINS 

LONGFIELD, C. 1947. The Odonata of South Angola. Archos Mus. Bocue 16 : 1-31, n figs. 

1953. The Atoconeura problem (Odon., Libellulidae) . Entomologist 86 : 42-49, pi. i. 

1955. The Odonata of North Angola. Part I. Publ foes cult. Co. Diam. Angola 27 (u) : 
13-63, 10 figs. 

1959. The Odonata of North Angola. Part II. Publ coes cult. Co. Diam. Angola 45 : 13-42, 

6 text-figs. 
MCLACHLAN, R. 1 88 1. Description of a new species of Corduliina (Gomphomacromia fallax) 

from Ecuador. Trans, ent. Soc. Lond. 1881 : 141-142. 
1883. Neuroptera of the Hawaiian Islands. Part I. Pseudo-Neuroptera. Ann. Mag. 

nat. Hist. (5) 12 : 226-240. 

18830. Description d'une espece nouvelle de Corduline du sousgenre Syncordulia. C.r. 
Soc. ent. Belg. 27 : xc-xci. 

1886. Two new species of Cordulina. Entomologists' mon. Mag. 23 : 104-105. 

1892. Supplementary note on the Neuroptera of the Hawaiian Islands. Ann. Mag. nat. 

Hist. (6) 10 : 176-179. 

1894. On two small collections of Neuroptera from Ta-chien-lu, in the province of 
Szechuen, Western China, on the frontier of Thibet. Ann. Mag. nat. Hist. (6) 13 : 412-436. 

- 18940. Some additions to the neuropterous fauna of New Zealand, with notes on certain 
described species. Entomologists' mon. Mag. (2) 5 : 238-243, 270-272. 

NUNNEY, W. H. 1895. A new West-African Insect. Ann. Mag. nat. Hist. (6) 16 : 349-351. 

PINHEY, E. C. G. 1952. Three new species of Odonata from Eastern Africa. Occ. Pap. 
Coryndon meml Mus. 3 : 13-16, fig. 5. 

1955- Some dragonflies of East and Central Africa and a rarity from Mauritius. Occ. 

Pap. Coryndon meml Mus. 4 : 17-41, pis 2-3, 9 text-figs. 

1958. Records of Dragonflies from the Zambezi and Rhodesia; a revision of the genus 
Platycypha; a gynandromorph dragonfly from Uganda. Occ. Pap. natn. Mus. Sth Rhodesia 
22B : 97-116, 7 text-figs. 

1961. A Survey of the Dragonflies (Order Odonata} of Eastern Africa, viii + 214 pp., 
ii pis., 2 text-figs. British Museum (Nat. Hist.), London. 

Ris, F. 1909-1919. Libellulinae, Catalogue systematique et descriptif. Coll. zool. Selys- 
Longchamps, Bruxelles, fasc. IX-XVI, 1,278 pp., 8 pis., 692 text-figs. 

19150. New Dragonflies (Odonata) of the subfamily Libellulinae from Sierra Leone, W. 
Africa. Ann. Mag. nat. Hist. (8) 15 : 213-223. 

1921. The Odonata or Dragonflies of South Africa. Ann. S. Afr. Mus. 19 : 245-452, pis. 

5-12, 77 text-figs. 
SELYS-LONGCHAMPS, E. DE. 1871. Synopsis des Cordulines. Bull. Acad. r. Sci. Belg. (2) 

31 : 238-316. 

18710. Synopsis des Cordulines (Suite). Deuxieme Legion. Macromia. Bull. Acad. r. 
Sci. Belg. (2) 31 : 519-565- 

1872. Note sur plusieurs Odonates de Madagascar et des lies mascareignes. Rev. Mag. 

Zool. (2) 23 : 175-183. 

1874. Additions au Synopsis des Cordulines. Bull. Acad. r. Sci. Belg. (2) 37 : 16-34. 
1878. Considerations sur la faune de la Nouvelle Guinee, des Moluques et de la Celebes. 

Mitt. K. zool. mus. Dresden 1878 : 289-322. 

1879. Description of a new species of Phyllomacromia (Corduliina) from West Africa. 
Entomologists' mon. Mag. 16 : 103-104. 

1881. Neophya Selys, nouveau genre de Cordulines. C.r. Soc. ent. Belg. 25 : xv-xviii, 

i fig. 

1883. Les Odonates du Japon. Ann. Soc. ent. Belg. 27 : 82-143. 

1887. Revision des Odonates de 1'Asie septentrionale, du Japon et de 1'Afrique septen- 
trionale. Ann. Soc. ent. Belg. 31 A : 50-68. 

1888. Odonates recueillis aux lies Loo-Choo par feu M. Pryer. C.r. Soc. ent. Belg. 
(3) 101 : xlviii-liii. 

TILLYARD, R. J. 1906. New Australian species of the family Libellulidae. Proc. Linn. Soc. 
N.S.W. 31 : 480-492, pi. 44. 



LIBELLULID & CORDULIID TYPES IN BM(NH) 305 

TILLY ARD, R. J. 1907. The Dragonflies of South-western Australia. Proc. Linn. Soc. N.S.W. 
32 : 719-742, pis. 34-36. 

1908. On some remarkable Australian Libellulinae. Proc. Linn. Soc. N.S.W. 33 : 637- 
649, pi. 14. 

19080. On the genus Nannodythemis, with descriptions of new species. Proc. Linn. Soc. 
N.S.W. 33 : 444-445, pi. 6. 

19086. On some remarkable Australian Corduliinae, with descriptions of new species. 
Proc. Linn. Soc. N.S.W. 33 : 737-751, pis. 21-22. 

1910. Monograph of the genus Synthemis. Proc. Linn. Soc. N.S.W. 35 : 312-377, pis. 4-9. 

1911. Further notes on some rare Australian Corduliinae, with descriptions of new species. 
Proc. Linn. Soc. N.S.W. 36 : 366-387, pi. 10. 

191 la. On the genus Cordulephya. Proc. Linn. Soc. N.S.W. 36 : 388-422, pis. 11-12. 
1913. On some Australian Anisoptera, with descriptions of new species. Proc. Linn. 

Soc. N.S.W. 37 : 572-584, pi. 64. 

1913(2. Description and life history of a new species of Nannophlebia. Proc. Linn. Soc. 
N.S.W. 37 : 712-726, pi. 74. 

I9I3&. Some descriptions of new forms of Australian Odonata. Proc. Linn. Soc. N.S.W. 
38 : 229-241, pi. 15. 

1917. On some new dragonflies from Australia and Tasmania [Order Odonata]. Proc. 
Linn. Soc. N.S.W. 42 : 450-479, pi. 23 and 10 text-figs. 

WHITE, A. 1846-1874. In The Zoology of the Voyage of H. M.S. Erebus and Terror, under the 
command of Captain Sir James Clark Ross, R.N., F.R.S., during the years 1839 to 1843. 
Ed. Richardson, J. & Gray, J. E. 2. Insects, p. 25. London. 




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1. MASNER, L. The types of Proctotrupoidea (Hymenoptera) in the British 
Museum (Natural History) and in the Hope Department of Entomology, Oxford. 
Pp. 143. February, 1965. 5. 

2. NIXON, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera : 
Braconidae). Pp. 284 ; 348 Text-figures. August, 1965. 6. 

3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177 ; 
18 plates, 270 Text-figures. August, 1965. 4 45. 

4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172 ; 500 Text-figures. October, 

1965- 35s. 

5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156 ; 
475 Text-figures. November, 1965. 2 155. 

6. OKADA,T. Diptera from Nepal. Cryptochaetidae, Diastatidae & Drosophilidae. 
Pp. 129 ; 328 Text-figures. May, 1966. 3. 

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

33s. 

8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the 
world species of Cleora (Lepidoptera : Geometridae). Pp. 119; 14 plates, 146 
Text-figures, 9 maps. February, 1967. 3 IDS. 

9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera : Rhopalocera). Pp. 509. August, 1967. 8 los. 

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rhopa- 
locera). Pp. 322; 233 Text-figures, coloured frontispiece. August, 1967. 8. 

11. MOUND, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 184; 
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PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED, BARTHOLOMEW PRESS, DORKING 



A REVISION OF THE GENUS 

AIOLOPUS FIEBER 
(ORTHOPTERA: ACRIDOIDEA) 




D. HOLLIS 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 22 No. 7 

LONDON: 1968 



A REVISION OF THE GENUS AIOLOPUS FIEBER 
(ORTHOPTERA : ACRIDOIDEA) 




BY 

D. HOLLIS 

' ^ ^.'_ "w 

Anti-Locust Research Centre, London 



- 3 7-355 ; 102 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 22 No. 7 

LONDON: 1968 



THE BULLETIN OF THE BRITISH MUSEUM 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

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

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

This paper is Vol. 22, No. 7 of the Entomological 
series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



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



Trustees of the British Museum (Natural History) 1968 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 4 October, 1968 Price 



A REVISION OF THE GENUS AIOLOPUS FIEBER 
(ORTHOPTERA : ACRIDOIDEA) 

By D. HOLLIS 

CONTENTS 

Page 

INTRODUCTION ........... 309 

BIOLOGY ............ 310 

TAXONOMIC TREATMENT ......... 312 

KEY TO SPECIES AND SUBSPECIES . . . . ... 316 

REFERENCES ........... 352 

SYNOPSIS 

The genus Aiolopus is redescribed and defined, and notes are given on its biology. A key is 
given to the seven species and six subspecies included within the genus, and all of these are 
redescribed. 

INTRODUCTION 

FOR the past century the genus Aiolopus has been a source of anxiety to acridid 
taxonomists. It arose in confusion through being described twice in the same year, 
first as Aiolopus (Fieber, 1853, May) and then as Epacromia (Fischer, 1853, Novem- 
ber), and its species have since been continually confused with one another. 

Up to 1938 even the familiar association of Aiolopus was in doubt but firstly 
Zimin (1938 : 27) and later Uvarov (1942 : 336) placed the genus naturally in the 
subfamily Oedipodinae (here considered a part of the Acridinae) close to Encop- 
tolophus and Hilethera. At the same time Uvarov (1942 : 337) erected two genera, 
Platypygius and Epacromius, from species previously included in Aiolopus. 

In recent decades it has become increasingly obvious that the species of Aiolopus 
can and are becoming serious crop pests, being particularly able to adapt from their 
natural grassland savannah habitat to irrigated crop areas. 

In the light of this knowledge and the confused state of the systematics of the 
genus it became clear that some form of revisionary work was necessary. The 
present paper attempts to revise the genus taxonomically and to introduce stability 
of identification of the species concerned. 

Immediately prior to this work the genus was thought to contain seventeen recog- 
nizable species. In a previous paper (Horn's, 1967) the author transferred two of 
these species to other genera. The present paper recognizes seven species, including 
one newly combined species, with four others reduced to subspecific status and the 
remaining five sunk into synonymy. 

All available types have been studied and where a type is lost or is inaccessible 
due to institutional policy this is clearly stated. If a type of a recognized species 
is destroyed a neotype is erected in the interests of stability of the nomenclature and 
where a species is based upon a series of syntypes a lectotype is erected from that 
series. 

ENTOM. 22, 7 15 



3 io D. HOLLIS 

Distribution of the recognized species and subspecies is only considered in the 
light of material studied by the present author. Previous records are not con- 
sidered, unless checked, because of the confusion which would arise from past 
misidentification and misapplication of names. For most species a large amount of 
material was studied and their distribution is presented in map form with notes on 
the countries in which each is found and the months of its known occurrence. 

Type depositories are given in abbreviated form as follows: 

Berlin Zoologisches Museum der Universitat, Berlin. 

BM(NH) British Museum (Natural History), London. 

Copenhagen Universitetets Zoologiske Museum, Copenhagen. 

Leningrad Zoological Institute, Academy of Sciences of USSR, Leningrad. 

Maastricht Natuurhistorisch Museum, Maastricht. 

Madrid Institute Espanol de Entomologia, Madrid. 

Paris Museum National d'Histoire Naturelle, Paris. 

Stockholm Naturhistoriska Riksmuseum, Stockholm. 

Turin Istituto e Museo di Zoologia, Torino. 

Uppsala Zoologiska Institutionen, Uppsala Universitet, Uppsala. 

Measurements and abbreviations of measurements used in this paper follow those 
ofDirsh(i953). 

Abbreviations used in the phallic complex figures are as follows: Ac arch of 
cingulum; Ap apical valve of penis; Apd apodeme of cingulum; Bp basal valve 
of penis; Cv cingular valve; Dp dorsal process of cingulum; Ejd ejaculatory 
duct; Ejs ejaculatory sac; Gpr gonopore process; Rm ramus of cingulum; 
Sps spermatophore sac. 

The opportunity is taken here to thank the Keeper of Entomology of the British 
Museum (Natural History) for allowing me to study the mass of material in his care ; 
Dr. T. H. C. Taylor for editing the manuscript; and the following colleagues for 
loans and information concerning type material; Professor G. Ya. Bei-Bienko, 
Leningrad; Professor J. van Boven, Leuven; Dr. M. Descamps, Paris; Dr. K. K. 
Giinther, Berlin; Dr. L. Hedstrom, Uppsala; Professor J. O. Hiising, Halle (Saale); 
Dr. K. H. L. Key, Canberra; Professor E. Morales Agacino, Madrid; the late Dr. 
B. Hanson, Stockholm; Dr. B. Petersen, Copenhagen; Professor M. Sam, Naples; 
and Dr. F. Willemse, Eygelshoven. 

BIOLOGY 

There is no major study on the biology of the species of Aiolopus available, but 
many authors have studied specific items of the biology of the more common species 
such as A . thalassinus and A . simulatrix (as A . savignyi) . 

Chesler (1938) studied the life history and described the immature stages of A. 
thalassinus in the Transvaal, and Hafez et al. (19620:) studied the same species in 
Egypt, both papers reporting that successful digging and oviposition took place in 
wet soil. Khalifa (1956), studying the egg-pods of A. thalassinus and A. simulatrix, 
mentions that a higher percentage of simulatrix than of thalassinus will oviposit 



REVISION OF GENUS AIOLOPUS 311 

on dry soil and the eggs of both species will develop without contact water as, pro- 
vided they are in an atmosphere of 100 % humidity, they can absorb enough moisture 
from the atmosphere for development. 

Zimin (1938) describes egg-laying and the egg-pods of thalassinus in southern 
Russia; Chapman & Robertson (1958) describe the egg-pods of thalassinus and longi- 
cornis in the Rukwa Valley, and Chapman (1961) describes those of thalassinus from 
Ghana; and Descamps & Wintrebert (1967) describe the egg-pods, eggs, copulation 
and oviposition of A . rodericensis (Butler) and suggest three to four generations per 
year for this species in S.W. Madagascar. 

Joyce (1952) records that A. simulatrix in the Sudan overwinters as adults in 
soil cracks and the present author confirms the presence of this species in soil cracks 
in the Sudan at the onset of winter. Khalifa (1956) states that in Egypt, where the 
winter is more severe, simulatrix overwinters in the egg stage, but thalassinus breeds 
throughout the year. This latter fact is verified for other regions by Nolte (1939) 
and Chapman (1962). Roffey (1965) suggests that, in Thailand, A. thalassinus 
tamulus undergoes more than one generation per year and does not overwinter in 
the egg stage. 

Joyce (1952 and 1954), studying the general biology of simulatrix in the Sudan, 
describes the formation of loose swarms and seasonal migration by this species. 
Khalifa (1956) states that both thalassinus and simulatrix will invade crop areas 
from fallow land and Hafez et al. (1963) describe experiments showing movement of 
thalassinus from Cynodon to irrigated areas for the higher relative humidity rather 
than for food preference, the female showing a better reaction to relative humidity 
than the male. Rungs (1938) describes thalassinus swarming in Morocco. Davey 
et al. (1959) suggest large scale migrations between seasonal habitats for thalassinus 
in Mali. 

Phipps (1959 and 1966) discusses the biology of thalassinus (as Aiolopus sp.) and 
longicornis with special emphasis on egg production, and Robertson & Chapman 
(1962) give short biological notes on longicornis, femoralis and thalassinus in the 
Rukwa Valley. 

Hafez et al. (1962), discussing the general ecology and biology of thalassinus in 
Egypt, showed the ability of both hoppers and adults to mature effectively on a 
variety of food crops and on Cynodon but found both cotton and lentil refused or 
detrimental to growth, and the same authors (Hafez et al., 19630) describe the poss- 
ible humidity and temperature receptors in thalassinus. 

Nolte (1939), discussing the colour forms and chromosome complex of thalassinus 
in Transvaal, considers this species a grassland type with two basic colour forms, 
green and light brown ; the latter was by far the more common and the former was 
not controlled in colour by environment alone since most, but not all, specimens of 
it darkened to brown on maturity. 

Very little is known of the natural enemies of Aiolopus spp., but the literature is 
reviewed by Greathead (1963) and the same author (1966) records Blaesoxipha 
anceps Vill. parasitizing simulatrix (as savignyi) in Eritrea. Chapman (1962) 
records rearing dipterous larvae from Aiolopus sp., Descamps & Wintrebert (1967) 
record Scelio sp. parasitizing eggs of rodericensis in Madagascar; and the present 



312 



D. HOLLIS 



author can record thalassinus killed by Entomophthora sp. (probably grylli) in 
Rhodesia. 

Of the published economic notes on the genus perhaps the most significant are 
those relating to simulatrix. Joyce (1952) gives an account of the damage caused 
by this species (as savignyi) to millet on a mechanized crop scheme in the Sudan. 
Coates (1893) records Epacromia dorsalis (Thunb.) and E. tricoloripes Burm. 
damaging various crops in many parts of India. The record of tricoloripes probably 
refers to A. thalassinus famulus (Fabricius) and that of dorsalis refers to simulatrix 
simulatrix as the figure given, and copied by Fletcher (1914 : 525), is obviously of 
simulatrix. Coates (1893) records simulatrix (as dorsalis) causing serious damage to 
summer rain crops, particularly millet, in the Upper Sind Frontier district; and 
Barlow (1900) records simulatrix (as dorsalis) damaging young wheat seedlings in 
the Ahmednagar district of India. 




AW 



FIG. i. Aiolopus thalassinus thalassinus (Fabr.). Entire insect, $, lateral view. 

(After Dirsh, 1965). 



TAXONOMIC TREATMENT 

Until the present revision, the genus Aiolopus has never been considered in its 
entirety and earlier workers, e.g. Finot (1895 : 421), Brunner von Wattenwyl 
(1882 : 145), I. Bolivar (1898 : 69), Innes (1929 : 30) and Chopard (1943 : 287), 
were only concerned with separating species in particular areas. Consequently no 
set of characters has been tested for the genus as a whole. 

Bei-Bienko (in Bei-Bienko & Mishchenko 1951 : 567) gives the most complete 
key to species previously published; in it five species are separated. He used a fairly 
large combination of characters, the genus being basically divided up on the width 
of the hind femur in relation to the width of the tegmen, coupled with the form of the 
pronotum. The present author has found the femoro-tegminal character unreliable 
as, for any species, the width of the tegmen varies directly with its length but by no 
means always directly with the width of the hind femur. 

The width of the hind femur relative to its length can be used as a rough guide 
to species-groups, although there are overlaps between those with narrow hind femora, 
e.g. A. thalassinus, those with hind femora of medium thickness, e.g. A. longicornis, 



REVISION OF GENUS AIOLOPUS 



- VV o 




Aov 



FIGS. 2-4. Aiolopus thalassinus thalassinus (Fabr.) $ genitalia. 2, ovipositor, ventral 
view; 3, same, lateral view; 4, subgenital plate, dorsal view. Ad apical diverticulum 
of spermatheca; Aov anterior (ventral) ovipositor valve; Avbs anterior ventral 
basivalvular sclerite; Eg egg guide; Jo Jannone's organs; La lateral apodeme; 
Lbs lateral apodeme; Lbs lateral basivalvular sclerite; Lov lateral (dorsal) ovi- 
positor valve; O oviduct; Pd preapical diverticulum of spermatheca; Pov pos- 
terior (inner) ovipositor valve; Pvbs posterior ventral basivalvular sclerite; Sa 
spermathecal aperture; Ss spermathecal sac. 



3H D. HOLLIS 

and the species with broad hind femora, e.g. A. strepens. However, when this 
character is coupled with other characters such as the form of the pronotum (Text- 
figs. 11-20), the frontal ridge (Text-figs. 21-29) and the length of the hind tibia 
relative to the hind femur (Text-figs. 5-8), the species are more clearly defined. 

Identification of subspecies is more difficult as the characters are geographically 
significant trends such as the colouration of the hind tibia and the relative proportions 
of the hind femur and tegmen. The latter character is used to separate A. thalas- 
sinus thalassinus from A. th. rodericensis and A. simulatrix simulatrix from A. s. 
femoralis by comparison of the ratio of the length of the tegmen over the length of 
the pronotum (E/P) with the ratio of the length of the hind femur over its maximum 
width (FL/FW) (Text-figs. 37-38, 90-91). 

The phallic complex appears to be of little use in separating species because, as 
in many other Acridinae genera, it is extremely similar in all the species and can 
only be used as a loose supporting indication of specific identity. 

The female subgenital plate (Text-fig. 4) and ovipositor (Text-figs. 2-3) are very 
uniform within the genus and the spermatheca has a similar range of variation in 
most species. 

AIOLOPUS Fieber, 1853 

Aiolopus Fieber, 1853 : 100. 

Epacromia Fischer, 1853 : 360; Rehn, 1902 : 317. 

Aeolopus [sic] Kirby, 1910 : 190. 

Aeoloptilus Bei-Bienko, 1966 : 1793, syn. n. 

Type-species: Gryllus thalassinus Fabricius, 1781; Kirby, 1910 : 190. 

Medium, size. Integument finely or moderately coarsely pitted. Antenna filiform, slightly 
shorter than, as long as, or longer than combined lengths of head and pronotum. Fastigium 
of vertex pentagonal, slightly longer than wide, moderately concave with well defined margins, 
forward angle acute or broadly rounded, fastigial foveolae trapezoid or rectangular, if former 
then narrowing forwards, shallow, normally with well defined margins; frons oblique; frontal 
ridge in some species slightly convex, in others flat or, sometimes weakly concave at and below 
median ocellus, if latter then marginal carinulae present but poorly developed. Eyes oval, 
long axis always vertical. Pronotum from slightly tectiform to slightly saddle-shaped, 
constricted at junction of prozona and metazona with former sometimes constricted medially; 
median carina linear, crossed only by posterior sulcus; raised lateral carinae absent in most 
species, rarely present in prozona, lateral " shoulders " often present; metazona longer than 
prozona, with obtuseangular posterior margin ; mesosternal interspace as long as wide or slightly 
wider than long, rectangular or trapezoid and widening posteriorly. Tegmen and hind wings 
fully developed ; intercalary vein of medial area of tegmen well developed and serrate at least in 
male, continuing to distal apex of medial area; membrane of tegmen semi-transparent with 
moderately sparse reticulation. Hind femur slender or broad, apical lobes rounded; hind 
tibia as long as or shorter than hind femur, apical spurs not specialized ; arolium of medium size 
or small. Male supra-anal plate rounded-triangular, with moderately elongated, subangular 
posterior apex; cercus narrow-conical with subobtuse apex; subgenital plate short, subconical, 
with rounded apex; epiphallus with moderately narrow bridge, curved ancorae and bilobed 
lophi ; ectophallic membrane sclerotized and forming sheath below apical penis valves ; cingulum 
with horseshoe-shaped arch bearing moderately long apodemes and very weak or more strongly 
developed dorsal processes, from ramus lateral processes extend backwards on each side above 
cingular valves, latter crescent-shaped in profile and slightly expanded medially ; cingular valves 



REVISION OF GENUS AIOLOPUS 315 

less sclerotized than apical penis valves, elipsoid in profile, with acute apices ; basal penis valves 
with moderately or well developed lateral expansions which are rarely recurved ; flexure narrow ; 
apical penis valves very narrow-triangular in profile, with acute apices. Ovipositor of female 
short; valves moderately robust, with curved apices, lower valve with small externo-ventral 
tooth; spermatheca with sac-like preapical and short finger-like apical diverticula. 

When this genus was first erected by Fieber as Aiolopus and later by Fischer as 
Epacromia neither author designated a type-species but both gave a list of contained 
species for their genera. Kirby (1910) designated Gryllus thalassinus Fabricius 
as the type-species for the genus Aiolopus after Rehn's (1902) synonymy of Fieber's 
and Fischer's genera. As G. thalassinus was listed by both Fieber and Fischer in 
their genera, Rehn's synonymy can be accepted. 

Bei-Bienko (1966) erected the genus Aeoloptilus for a new species carinatus Bei- 
Bienko (1966), described from Komodo Island. The present author has been able 
to examine male and female paratypes of this species and a female specimen from 
Sumba Island. Bei-Bienko separates Aeoloptilus carinatus from Aiolopus by the 
presence of lateral pronotal carinae, the weak and elongate-oval fastigial foveolae 
and the shape of the intercalary vein of the medial area of the tegmen. As will be 
seen below, lateral pronotal carinae are not good diagnostic features in Aiolopus, 
as they may or may not be very weakly developed in A. thalassinus famulus 
(Fabricius). The fastigial foveolae in carinatus appear to be of the Aiolopus type 
but weakly developed. The intercalary vein of the medial area of the tegmen in 
carinatus shows no significant difference in shape or position from that of A . thalas- 
sinus (Fabricius). Furthermore the male phallic complex of carinatus is extremely 
similar to that of A. th. famulus (Fabricius). For these reasons the genus Aeolop- 
tilus Bei-Bienko is synonymized with Aiolopus, but the type-species, carinatus, is 
kept distinct for reasons discussed below (p. 334). 

Aiolopus is placed naturally in the oedipodine half of the subfamily Acridinae 
where it may be distinguished by the following combination of characters: filiform 
antenna, trapezoid or rectangular fastigial foveolae, weakly tectiform or saddle- 
shaped dorsum of pronotum, lateral pronotal carinae absent or weakly present only 
in prozona, metazona of pronotum longer than prozona and its posterior margin 
obtuseangular, mesosternal interspace as wide as or wider than long, intercalary 
vein of medial area of tegmen extending to distal apex of medial area, hind femur 
with lower outer area not expanded, hind tibia not expanded and its apical spurs 
not specialized, epiphallus with simple bilobed lophi, male subgenital plate sub- 
conical, and apical penis valves short. 

The genus is widely distributed throughout the Old World south of latitude 50 N. 
and north of latitude 40 S., extending westwards to the Cape Verde Islands and 
eastwards to the Samoan Islands. Its distribution in the Indian Ocean region is of 
particular interest. Two species are present, A. thalassinus and A. simulatrix, both 
on the east coast of Africa and the west coast of India, but thalassinus is not found 
east of the Seychelles Islands and simulatrix is not found west of Coetivy Island. 
This possibly suggests invasion of the region by the genus during different geological 
times. 



316 D. HOLLIS 

KEY TO SPECIES AND SUBSPECIES 

1 Hind tibia considerably shorter than hind femur, with a maximum of nine outer and 

ten inner spines (Text-fig. 8) ; frontal ridge coarsely and densely pitted, without 
lateral carinulae, with margins converging strongly just below fastigium (Text-fig. 
27); prozona of pronotum gradually sloping away on each side of median carina 
and without trace of lateral " shoulders " (Text-figs. 15-16); hind femur broad or 
very broad ............ 2 

- Hind tibia as long as or only slightly shorter than hind femur, with at least nine to 
twelve outer and ten to thirteen inner spines (Text-figs. 5-7) ; frontal ridge more 
sparsely pitted, if coarsely pitted then not narrowing strongly just below fastigium 
(Text-figs. 21-26, 28-29); pronotum almost flat or slightly saddle-shaped, prozona 
either with pattern suggesting presence of lateral carinae or almost flat dorsally 
and without median constriction (Text-figs. 11-14, I 7~' 20 )', hind femur narrow 
or broad ............. 3 

2 Hind femur very broad, ratio of length to width (FL/FW) barely 3-0; tegmen hardly 

surpassing tip of hind femur, ratio of length of tegmen over length of pronotum 
(E/P) normally between 3-7 and 4-5. S.W. Tanzania, Zambia and South Africa 

simulatrix fetnoralis Uvarov (p. 325) 






FIGS. 5-10. Aiolopus spp. 5-8 hind femora, lateral view: 5, A. thalassinus thalassinus 
(Fabr) ; 6, A . longicornis Sjost. ; 7, A . strepens (Latr.) ; 8, A . simulatrix simulatrix (Walker) ; 
9 and 10, pulvillus and claw of hind tarsus, ventral view: 9, A. thalassinus thalassinus 
(Fabr.) ; 10, A . oxianus Uv. 



REVISION OF GENUS AIOLOPUS 



317 



Hind femur slightly less broad, ratio of length to width (FL/FW) about 3-4; tegmen 
well surpassing tip of hind femur, ratio of length of tegmen over length of pronotum 
(E/P) normally between 4-5 and 5-3. Northwards from E. Tanzania to eastern 
Mediterranan region, Arabian peninsula, Middle East, India, Burma, Islands of 
Indian Ocean westwards to Seychelles. . . simulatrix simulatrix (Walker) (p. 320) 

Pronotum almost flat dorsally and without constriction in prozona (Text-figs. 13-14) ; 
hind femur broad, ratio of length to width (FL/FW) about 3-4 (Text-fig. 7); hind 
tibia with ten outer and eleven inner spines, red except for basal quarter ; maximum 
width of face (C) normally less than maximum width of hind femur (FW) ; anten- 
nae shorter than combined lengths of head and pronotum. Southern Palaearctic 
region westwards to Persian Gulf. ..... strepens (Latreille) (p. 327) 

Pronotum slightly saddle-shaped and with median prozonal constriction (Text-figs. 
1 1 -i 2, 17-20); hind femur narrow or broad, if latter then only with nine outer 




FIGS. 11-20. Aiolopus spp., pronota. n, A. thalassinus thalassinus (Fabr.), lateral view; 
12, same, dorsal view; 13, A. strepens (Latr.), lateral view; 14. same, dorsal view; 15, A. 
simulatrix simulatrix (Walker), lateral view; 16, same, dorsal view; 17, A. longicornis 
Sjost., lateral view; 18, same, dorsal view; 19, A. carinatus (Bei-Bienko), dorsal view; 
20, same, lateral view. 



D. HOLLIS 



and ten inner spines (Text-figs. 5-6) ; maximum width of face (C) normally greater 
than maximum width of hind femur (FW) ; antennae at least as long as combined 
lengths of head and pronotum ......... 




25 



28 



FIGS. 21-29. Aiolopus spp., heads, anterior view. 21, A. thalassinus thalassinus (Fabr.), 
$; 22, A. thalassinus tamulus (Fabr.), <j>; 23, A. thalassinus famulus (Fabr.), (J from Christ- 
mas Island; 24, A. thalassinus dubius (Willemse), $; 25, A. oxianus Uv., $; 26, A. longi- 
cornis Sjost., $; 27, A. simulatrix simulatrix (Walker), $; 28, A. strepens (Latr.), $; 
29, ^4. carinatus (Bei-Bienko), ?. 



REVISION OF GENUS AIOLOPUS 319 

Arolium of hind tarsus very small, about one quarter the length of claw (Text-fig. 10) ; 
frontal ridge weakly sulcate and with parallel, obtuse, lateral carinulae which 
extend almost to base of frons (Text-fig. 25). Central Asia oxianus Uvarov (p. 330) 

Arolium of hind tarsus larger, about half length of claw (Text-fig. 9) ; frontal ridge 
normally flat, if sulcate then lateral carinulae gradually converging upwards 
(Text-figs. 21-24, 29) ........... 5 

Fastigial foveolae weak, ovoid, with poorly developed margins (Text-fig. 29) ; 
prozona of pronotum with lateral carinae moderately developed, parallel or 
weakly diverging anteriorly (Text-figs. 19-20); hind tibia with twelve outer and 
twelve or thirteen inner spines. Komodo and Sumba Islands. 

carinatus (Bei-Bienko) (p. 332) 

Fastigial foveolae rectangular or trapezoid, with well developed margins (Text-figs. 
2124) > prozona of pronotum with lateral carinae absent or very weakly developed 
(Text-figs. 11-12) ; hind tibia with nine or ten outer and ten or eleven inner spines 6 

Antennae very long, when extended backwards almost reaching base of hind femur; 
face broad (Text-fig. 26), ratio of length of pronotum over maximum width of 
head (P/C) about 1-05; large species with very prominent criss-cross pattern on 
dorsum of pronotum (Text-figs. 17-18); hind femur of medium thickness. East 
Africa, Lake Chad ...... longicornis Sjostedt (p. 334) 

Antennae not longer than combined lengths of head and pronotum; face narrower, 
ratio of length of pronotum to maximum width of head (P/C) about 1-20 but if 
much less then pronotum short and broad and fastigial foveolae not much longer 
than wide ............. 7 

Fastigial foveolae broadly trapezoid, hardly one and a half times longer than wide; 
hind tibia not black ventrally ; pronotum short and broad ; hind femur broad, ratio 
of length to width (FL/FW) less than 3-7. East and southern Africa 

meruensis Sjostedt (p. 336) 

Fastigial foveolae narrowly trapezoid, about twice as long as broad, but if less then 
hind tibia bluish in apical two thirds or almost completely black ventrally; 
pronotum narrower ; hind femur of medium thickness or narrow, ratio of length to 
width (FL/FW) more than 3 -5 8 

Ventral surface of hind tibia completely black or dark brown in apical three quarters ; 
fastigial foveolae rectangular and about one and a half times longer than wide (Text- 
fig. 24). Polynesia eastwards to Samoa . . thalas sinus dubius (Willemse) (p. 351) 

Ventral surface of hind tibia with dark colouration, if present, broadly interrupted 
with reddish, bluish or ochraceous band; fastigial foveolae narrowly trapezoid, 
about twice as long as wide (Text -figs. 21-23) ....... 9 

Hind femur of medium thickness, ratio of length to width (FL/FW) between 3-4 and 
4-0; tegmen shorter, ratio of length of tegmen over length of pronotum (E/P) 
between 4-1 and 4-9; hind tibia without red colouration Madagascan subregion 
west of Comoro Islands . . . thalassinus rodericensis (Butler) (p. 343) 

Hind femur narrower, ratio of length to width 4-0 or more; tegmen longer, ratio of 
length of tegmen over length of pronotum (E/P) 4 - 7-5'5. Palaearctic, Ethiopian 
and Oriental regions ........... 10 

Frontal ridge narrow, gradually and continuously narrowing upwards (Text-figs. 
22-23) J hind tibia with red colouration, if present, broadly separated from basal 
black band by bluish grey band. East India and Ceylon eastwards to Australia. 

thalassinus tamulus (Fabricius) (p. 347) 

Frontal ridge broad, with lateral margins only slightly convergent upwards to 
fastigium (Text-fig. 21); hind tibia with apical red colouration, if present, narrow- 
ly separated from basal black band by ochraceous band. Ethiopian region, N. 
and W. India westwards to Palaearctic region. 

thalassinus thalassinus (Fabricius) (p. 340) 



320 D. HOLLIS 

Aiolopus simulatrix (Walker, 1870) 
This species is divided into two subspecies which are described and discussed below. 

Aiolopus simulatrix simulatrix (Walker, 1870) stat. n. 
(Text-figs. 8, 15, 16, 27, 30-39) 

Epacromia simulatrix Walker, 1870 : 773. 
Heteropternis (?) savignyi Krauss, 1890 : 262, syn. n. 
Epacromia affinis I. Bolivar, 1902 : 600, syn. n. 
Acrotylus simulatrix (Walker, 1870) Kirby, 1910 : 267. 
Aeolopus laticosta I. Bolivar, 1912 : 270, syn. n. 
Aeolopus strepens deserticola Uvarov, 1922 : 358, syn. n. 

Type locality. South Hindustan; type <$ deposited in the BM(NH). 

Redescription. <$. Integument more strongly rugulose than the other species in the genus. 
Antenna as long as combined lengths of head and pronotum with twenty-two to twenty-four 
segments. Fastigium of vertex pentagonal, slightly longer than wide, moderately concave with 
well defined margins, forward angle narrowly rounded; fastigial foveolae rectangular, shallow, 
coarsely pitted, with moderately well defined margins, lower margin often very weak; frontal 
ridge wide, coarsely and densely pitted (Text-fig. 27), with parallel margins along most of length 
but narrowing strongly just below fastigium. Eye oval, about one and a half times as high as 
wide and almost twice as high as length of subocular groove. Pronotum relatively narrow 
(Text-figs. 15-16); prozona cylindrical above, with very slight median constriction; metazona 
rather flat, with obtuse angular posterior margin ; median longitudinal carina stronger in prozona 
than in metazona; lateral plate of pronotum higher than wide; mesosternal interspace wider 
than long, trapezoid, slightly widening posteriorly. Tegmen relatively long, E/P ratio about 
4-9. Hind femur broad, ratio of length to width about 3-5; hind tibia shorter than hind femur 
(Text-fig. 8), with nine outer and ten inner spines, inner apical spurs slightly less than twice 
as long as outer pair; arolium almost half length of claw. Phallic complex (Text-figs. 30-33) 
with zygoma of cingulum with small dorsal processes and short apodemes ; basal valves of penis 
with small lateral expansions which are not recurved posteriorly. 

General colouration mid-brown with ochraceous (or green) and blackish markings; pronotum 
often unicolorously brown or rarely greenish, sometimes with weak ochraceous X-shaped pattern 
on dorsum; tegmen with two ochraceous transverse fasciae which extend from anterior margin 
to first vannal vein (Snodgrass, 1935), the proximal fascia narrows somewhat posteriorly, apex 
of tegmen mottled; hind wing hyaline, sometimes pale yellow basally, apex slightly darkened; 
hind femur with two dark spots in upper outer area, inner surface ochraceous with two incomplete 
dark fasciae, lower inner area often reddish, hind " knee " blackish; hind tibia narrowly black 
basally, followed by broad ochraceous ring and with broad blackish or grey ring medially, apical 
third reddish or ochraceous. 

$. Larger and more robust than male. Spermatheca as in Text-figs. 34-36. 

Measurements (mm.). Length of body, <J 16-9-26-0, $21-0-30-8. Length of pronotum. 
cJ 3-2-4-5, $ 3-9-5-2. Length of tegmen, <$ 16-1-21-9, ? 18-5-26-6. Length of hind femur, 
c? 9-5-14-5, $11-0-14-6. Maximum width of hind femur, $ 2-7-4-3, $3-3-4-6. Maximum 
width of head (C), < 2-9-4-1, $ 3-6-5-0. 

Ratios (forty males and forty females measured) : 

P/C FL/FW E/P E/F 

r- A * 

Min. Ave. Max. 

i -54 1-69 1-87 
1-54 1-72 1-82 









\ 


f 






^ 


t 






-\ 


Min. 


Ave. 


Max. 


Min. Ave. 


Max. 


Min. 


Ave. 


Max. 


I 'OO 


I'll 


X 


20 


3 


27 3-48 


3 


65 


4' 


75 


4'93 


5'34 


I 'OO 


1-09 


z 


21 


3 


13 3'43 


3 


78 


4 


46 


4'93 


5'3 Z 



REVISION OF GENUS AIOLOPUS 



321 




34 



FIGS. 30-36. A. simulatrix simulatrix (Walker), genitalia. 30, $ epiphallus; 31, $ phallic 
complex, dorsal view, epiphallus and ectophallic membrane removed; 32, same, lateral 
view; 33, $ endophallus, lateral view; 34, $ spermatheca of specimen from Tanzania; 
35, same, from Jordan; 36, same, from S. India. 



322 D. HOLLIS 

Discussion. This subspecies is quite variable in size, general colouration, relative 
width of hind femur and relative length of tegmen. The last two characters show 
a very general tendency to vary according to geographical distribution, the trends 
being towards slightly broader hind femora and shorter tegmina as the species ex- 
tends southwards down the eastern half of the African continent and narrower 
hind femora and longer tegmina as the species extends southwards down the Indian 
peninsula. The former trend reaches its culmination in southern Africa, and the 
forms involved in it are here regarded as a separate subspecies A. simulatrix femoralis 
Uvarov; the latter trend culminates in populations on the Maldive, Chagos and Sey- 
chelles Islands. By comparing the ratios of length to width of the hind femur against 
the ratio of the length of the tegmen over the length of the pronotum these trends 
may be demonstrated, as in Text-figs. 37-38. 

A. simulatrix simulatrix may be readily distinguished from other species in the 
genus by the form of the frontal ridge, pronotum and hind leg. 

Up to the present this species has been recorded in the literature as A. savignyi 
(Krauss), the type of which species does not exist and is known only as a drawing 
in Savigny's " Descriptions de 1'Egypt " (1809-1813 : 182, pi. 6, fig. 16), from which 
a specific description was made by Krauss (1890 : 262) as Heteropternis (?) savignyi. 
The original drawing is not particularly diagnostic and as the specimen from which 
the drawing was made is no longer in existence, the true identity of the name savignyi 
is open to subjective conclusions. Storey (1919 : 55) synonymized H. savignyi 
with Epacromia strepens (Latreille) but in all probability Storey's E. strepens was the 
same species as A . savignyi of other authors. Uvarov (1942 : 337) firmly established 
the synonymy of H. (?) savignyi with Epacromia affinis I. Bolivar (1902) and A. 
strepens affinis Uvarov (1924 : 21). In order not to confuse the literature even 
further the present author accepts Uvarov's interpretation of Savigny's drawing 
as being the species which has, up to the present, been named A. savignyi (Krauss). 

However the type male of Epacromia simulatrix Walker, 1870 represents the same 
species and therefore Krauss's name should fall into synonymy. The use of the 
older Walker name is a strict application of the law of priority but does involve the 
use of a nomen oblitum, as it has not been used as a senior synonym in the primary 
zoological literature since its original publication in 1870; it has been used, in the 
present author's opinion quite wrongly, as a junior synonym by Kirby (1914 : 122) 
and I. Bolivar (1918 : 382). The usage of Walker's name is proposed for the two 
following reasons: firstly, the name in common usage, i.e. savignyi Krauss, is based 
upon the description of a drawing of a specimen which is now lost and which is there- 
fore open to subjective interpretation; and secondly, the older nomen oblitum i.e. 
simulatrix Walker, is based upon the description of an objective type specimen 
which is still in existence. 

Aeolopus strepens deserticola Uvarov is synonymized here, since the male upon 
which Uvarov based the subspecies falls well within the range of individual variation 
of the nominate subspecies. The type male bears the locality label "KAZVIN, 
N.W. Iran " and is deposited in the BM(NH). 

Aeolopus laticosta I. Bolivar, described from the Chagos Islands in the Indian Ocean, 
represents an extreme form of the nominate subspecies which, however, cannot be 



REVISION OF GENUS AIOLOPUS 



323 



38 
3-7 
36 
35 

A 34 
3-3 

3-2 
F'L 
FW 3- I 

30 
2-9i 



x-femoralis 

-simulatrix (Palaearctic) 
-simulatrix (Ethiopian) 
-simulatrix (priental) 
O-simulatrix (Maldive Is!) 



O 

f\ ^L 

fo 



37 38 39 4O 41 42 4-3 44 45 4-6 47 48 4-9 SO 5-1 52 53 54 

37 ' * > 



AS- 



FL 
FW 



38- 
3-7 
36 
35 
3-4 
3 
32 
3-1 
30 
2-9 
2-8 



x-femoralis 

-simulatrix (Palaearctic) 
-simulatrix (Ethiopian) 
-simulatrix (Oriental) 
O-simulatrix (Maldive Is) 



XX X X 













O 



o o 







36 37 38 39 4-O 4-1 42 4-3 44 4-5 4-6 47 4 49 5O 51 5-2 53 5-4 
^ ^ 

FIGS. 37, 38. A. simulatrix subspp., scatter diagrams comparing values of 
FL/FW against E/P. 37, males ; 38, females. 



ENTOM. 22, 7 



16 



324 



D. HOLLIS 



regarded as distinct as it merges very well with the more typical form on the Indian 
mainland through populations on the Maldive and Laccadive Islands. The lecto- 
type male, bearing the locality label " Diego Garcia, Chagos Is. ", was selected by 
Dirsh (1963) and is deposited in BM(NH). 

Epacromia affinis I. Bolivar was described from a male and female from South 
India. The male is here selected as LECTOTYPE and bears the following data: 
" Madure, P. Pantel, E. affinis " ; it is deposited in Madrid. 

Distribution (Text-fig. 39). From the seven hundred and thirty-nine specimens 
of this subspecies examined the following distributional conclusions are made : 

TANZANIA, March to April, July, December to January; KENYA, January to June; 
SOMALIA, June to August, October to November, January; FRENCH SOMALILAND, 
August ; ETHIOPIA, February to April, July, November to December; SUDAN, May 



20 




simulatrix-* 
femoralis -4 



40- 



FIG. 39. A. simulatrix subspp., distribution map. 



REVISION OF GENUS AIOLOPUS 325 

through to January; NIGERIA, May, July to August, October; SENEGAL, August 
to September; MALI, June, August to December, March; UNITED ARAB REPUBLIC, 
May, August to September; SOUTHERN YEMEN, June, August to November; YEMEN, 
March, June to July, December; SAUDI ARABIA, August through to June; ISRAEL, 
June, August, October to November; JORDAN, March, May, June, October; SYRIA, 
August; TURKEY, November; CYPRUS, May, July, September; IRAQ, May to Sept- 
ember, December; IRAN, April to June; WEST PAKISTAN, July, September, November; 
INDIA, June to November, March to April; BURMA, October; LACCADIVE Is., No 
dates; MALDIVE Is., July to August, October, December to February; CHAGOS Is., 
May to December; SEYCHELLES, Coetivy, May to December. 

Aiolopus simulatrix femoralis Uvarov, 1953 stat. n. 

(Text-figs. 37-45) 
Aiolopus femoralis Uvarov, 1953 : 109, figs. 126-128. 

Type locality. Northern Rhodesia : Luano Valley, Chisorwe ; type <$ deposited 
intheBM(NH). 

Differs from the nominate subspecies in the following ways: 

cj. Smaller size; tegmen shorter, hardly reaching or just exceeding tip of hind femur, E/P 
ratio about 4-0; hind femur very broad, about three times longer than maximum width or even 
broader; hind tibia with normally eight or rarely nine inner and normally nine or rarely ten outer 
spines; phallic complex (Text-figs. 40-43) very similar to nominate subspecies, but dorsal pro- 
cesses on zygoma of cingulum a little larger and apodemes a little longer, and lateral expansions 
of basal valves of penis larger and slightly recurved. 

$. Spermatheca as in Text-figs. 44-45. 

Measurements (mm.). Length of body, cJ 16-8-19-9, $21-3-25-5. Length of pronotum, 
cj 3-6-4-1, $ 4-3-5-0. Length of tegmen, $ 14-1-17-0, $ 15-8-20-2. Length of hind femur, 
cj 9-5-11-4, $10-9-13-8. Maximum width of hind femur, <$ 3-1-3-6, $3-8-4-4. Maximum 
width of head, (J 3-1-3-7, $ 4-0-4-6. 

Ratios (twelve males and twelve females measured) : 



FL/FW E/P E/F 




Min. Ave. Max. Min. Ave. Max. Min. Ave. Max. Min. Ave. Max. 
cJ 1-08 1-14 1-18 2-97 3-15 3-28 3-76 4-18 4-43 1-42 1-51 1-58 
? i-oo 1-06 1-14 2-80 2-98 3-19 3-67 3-97 4-39 1-37 1-47 1-58 

Discussion. Only a small number of specimens of this subspecies have been 
examined and variation appears chiefly in size and general colouration. There is a 
general tendency as the subspecies extends southwards for the hind femur to become 
relatively thicker and for the number of spines on the hind tibia to decrease from 
nine outer and ten inner to eight outer and nine inner. 

When first described, femoralis was considered to be a distinct species and by its 
broad hind femora and short tegmina the type series would appear to be fairly well 
distinguished from simulatrix. However, specimens from southern Tanzania are less 
well defined and the specimen examined from South Africa is even more clearly defined 



326 



D. HOLLIS 



than the type series, suggesting a culmination of trends which can be seen in simula- 
trix as it extends southwards down E. Africa. The line of demarkation between 
the two subspecies in Tanzania coincides fairly well with that between the wooded 
steppe of the Sudan and of E. Africa and the savanna of south-eastern Africa (see 
Keay, 1959). 

Distribution (Text-fig. 39) (thirteen males and twelve females examined). TAN- 
ZANIA: Kimamba, iv, i $; Tumba, i, 2 <$, i $; ii, 5 <$; xii, I <; Lake Rukwa, v, i $; 
C. Rukwa, Lundi Mbuga, xii, 4 $; Mshughaa, 35 mis, S.E. of Singida, xii, i <$. 

ZAMBIA: Luano Valley, Chisorwe, ii, 3 $, 3 $; iii, 2 $. 



Ejd Ejs Cpr Sps Km 





Bp 



Apd 



41 



-G 




Ejd Ejs Cpr Sps 



FIGS. 40-45. A. simulatrix femoralis Uv., genitalia. 40, $ epiphallus ; 41, ^phallic 
complex, dorsal view, epiphallus and ectophallic membrane removed; 42, same, lateral 
view; 43, $ endophallus, lateral view; 44, $ spermatheca of specimen from Zambia; 
45, same, from Tanzania. 



REVISION OF GENUS AIOLOPUS 327 

SOUTH AFRICA: Transvaal, Zoutpansberg distr., Limpopo R., nr. Kobeenpan, 
2,200 ft., iv, " along banks ", i $. 



Aiolopus strepens (Latreille, 1804) 
(Text-figs. 7, 13, 14, 28, 46-51) 

Acrydium strepens Latreille, 1804 : 154. 

Gryllus prasinus Thunberg, 1815 : 239, syn. n. 

Acridium vittatum Brulle, 1840 : 78, pi. 5, fig. 7; Finot, 1895 : 422. 

Type locality. "Environs de Bordeaux"; specimen lost. NEOTYPE ($) 
erected bearing the following data: FRANCE: Dordogne, les Eyzies, ix. 1949 (ex. 
Zeuner coll. ), B.M. 1964-194. This specimen was chosen because of its agreement 
with Latreille's original description and the close proximity of its locality to the 
original type locality; it is deposited in the BM(NH). 

Redescription. <$. Robust body-form, integument moderately rugulose. Antenna shorter 
than combined lengths of head and pronotum. Fastigium of vertex pentagonal with forward 
angle broadly rounded, hardly longer than wide, shallowly concave ; fastigial foveolae trapezoid, 
narrowing forwards, very shallow with lower margin somewhat obliterated, about one and three 
quarter times as long as maximum width ; frontal ridge flat or convex, sparsely pitted, gradually 
narrowing upwards (Text-fig. 28). Eye ellipsoid, almost twice as high as maximum width and 
about one and three quarter times as high as length of subocular groove. Dorsum of pronotum 
subtectiform, rather flat, prozona not constricted medially (Text-fig. 13-14); metazona almost 
one and three quarter times as long as length of prozona, with narrowly obtuseangular posterior 
margin ; lateral plate of pronotum higher than wide ; mesosternal interspace rectangular, slightly 
broader than long. Tegmen short and broad, normally only slightly exceeding tip of hind femur. 
Hind femur broad, about 3-3 times as long as maximum width and always broader than maximum 
width of head; hind tibia as long as hind femur (Text-fig. 7), with ten outer and normally eleven 
inner spines; inner apical spurs slightly more than one and a half times as long as outer pair; 
arolium about half length of claw. Supra-anal plate, cercus and subgenital plate normal for 
genus. Phallic complex (Text-figs. 46-49) with zygoma of cingulum without dorsal processes 
but expanded laterally at bases of apodemes; basal valves of penis expanded laterally but not 
recurved. 

General colouration from uniformly brown through brown and ochraceous or green to uniformly 
green; dorsum of pronotum normally unicolourous but sometimes with median longitudinal 
contrasting stripe which may continue forwards along vertex ; tegmen with pale basal transverse 
fascia which extends either as far across as medial vein or as far as ist vannal vein in which case 
it narrows strongly posteriorly, median pale transverse fascia always extending broadly or 
narrowly to ist vannal vein; hind wing hyaline, often bluish basally, apex darkened; hind femur 
apically with inner area black and upper outer area blackish, lower inner area reddish ; hind tibia 
narrowly black basally, basal quarter ochraceous or pinkish, apical three quarters red, the two 
colours separated by a narrow and incomplete black band. 

$. As male but larger and more robust; spermatheca as in Text-fig. 50. 

Measurements (mm.). Length of body, $ 17-7-23-7, $ 22-7-32-0. Length of pronotum, 
<? 3'9~5'> $ 4-6-7-0. Length of tegmen, $ 16-7-23-6, $ 19-6-30-9. Length of hind femur, 
<J 11-3-14-8, $13-4-19-2. Maximum width of hind femur, <J 3-3-4-1, $4-3-5-8. Maximum 
width of head, 3-3-4-1, $ 4-2-5-7. 



328 



D. HOLLIS 



Ratios (twenty males and twenty females measured) : 



P/C 



Min. Ave. Max. 
1-18 1-24 1-29 
1-04 1-24 1-33 



FL/FW 

_A_ 



E/P 
^/^_ 



E/F 



t t , 

Min. Ave. Max. Min. Ave. Max. 

3-19 3'39 3-64 4'6 4'33 4'85 

- ~~ -- ~-.- 3 -60 3 '93 4'S 1 



3-00 



3'45 



Min. Ave. Max. 
1-39 1-52 
1-38 1-51 



1-71 
1-67 



Discussion. There is considerable variation in size and general colouration in 
this species. Specimens from N. Africa are much larger than others and their 
general colouration is very like that of A. simulatrix; they can, if not examined in 
detail, be misidentified as the latter. 

A. strepens differs from other species in the genus in the form of the pronotum; 
also it differs from the thalassinus group in having a broad hind femur and from the 



Bp 



Apd 




50 



FIGS. 46-50. A. strepens (Latr.), genitalia. 46, $ epiphallus; 47, <J phallic complex, 
dorsal view, epiphallus and ectophallic membrane removed; 48, same, lateral view; 
49, (J endophallus, lateral view; 50, spermatheca. 



REVISION OF GENUS AIOLOPUS 



329 




330 D. HOLLIS 

simulatrix group in that although the hind femur is broad the hind tibia is as long 
as the hind femur and has ten outer and eleven inner spines. 

The author has examined the type <$ of Gryllus prasinus Thunberg, deposited in 
Uppsala, and makes the above synonymy although this species has been previously 
considered a junior synonym (Stal, 1873 : 112) of A. thalassinus. 

The type of Acridium vittatum Brulle has not been traced, but from the original 
description and figure it is obvious that Finot's synonymy should be accepted. 

Distribution (Text-fig. 51). Nine hundred and eighty-four specimens of this 
species were examined and the following conclusions on its distribution are made : 

FRANCE, May to October, December; CORSICA, August; SPAIN, April to May, 
August to October; BALEARIC Is., April, September to October; PORTUGAL, August; 
ITALY, July through to May; SICILY, September; SARDINIA, September; MALTA, 
January; GREECE, April to September; YUGOSLAVIA; July to September; ALBANIA, 
August; BULGARIA, August; CYPRUS, April to December; TURKEY, July to Septem- 
ber, November; LEBANON, March to May, July, September, December; SYRIA, 
February to May, November; ISRAEL, April; IRAQ, June; IRAN, March; MADEIRA, 
June to July, November to December; CANARY Is., April, July to September, 
December, January; MOROCCO, June to September, December, ALGERIA, May to 
July, September to October; TUNISIA, January, April; LIBYA, February to May, 
July, September, November; UNITED ARAB REPUBLIC, March, May to August, 
October. 

Aiolopus oxianus Uvarov, 1926 

(Text-figs. 10, 25, 52-56) 
Aiolopus oxianus Uvarov, 1926: 347. 

Type locality. USSR, " Kerki, on River Amu-Darya"; type <$ deposited in 
Leningrad. 

Redescription. $. Slender build, moderately rugulose. Antenna longer than combined 
lengths of head and pronotum, with twenty-two to twenty-four segments, the median segments 
of the flagellum elongate. Fastigium of vertex pentagonal, longer than wide, weakly concave 
with well denned margins ; fastigial foveolae elongate, trapezoid, narrowing forwards, more than 
twice as long as maximum width, with well denned margins ; frontal ridge weakly sulcate along 
most of length (Text-fig. 25), marginal carinulae low but distinct almost to clypeus, parallel 
along most of their length but converging slightly just below fastigium. Eye more rounded 
than in other species, little more than one and a quarter times higher than maximum width but 
twice as high as length of subocular groove. Pronotum narrow, weakly saddle-shaped, prozona 
constricted medially; metazona about one and three quarter times longer than prozona, with 
obtuseangular posterior margin; lateral plate of pronotum higher than wide; mesosternal 
interspace slightly wider than long. Tegmen long and narrow, well exceeding tip of hind femur. 
Hind femur narrow, ratio of length to maximum width about 4-1; hind tibia as long as hind 
femur, with ten outer and eleven inner spines, inner apical spurs slightly longer than outer pair ; 
arolium very small (Text-fig. 10), about one quarter the length of claw. Supra- anal plate, 
cercus and subgenital plate normal for genus. Phallic complex (Text-figs. 52-55) with apodemes 
of zygoma of cingulum short, weakly sinuous and with apices weakly expanded dorsally, dorsal 
processes of zygoma weak or absent ; basal valves of penis expanded apically but not recurved, 
apical penis valves slightly longer than normal. 



REVISION OF GENUS AIOLOPUS 331 

General colouration brown with some ochraceous and black markings; head and pronotum 
with narrow median longitudinal ochraceous stripe sometimes present, ochraceous X-shaped 
pattern on dorsum of pronotum weak or absent; tegmen with broad dark transverse band 
basally and diffuse dark band medially, apical half mottled; hind wing hyaline; upper outer 
area of hind femur with three black spots, outer area with faint subapical black transverse band 
and faint ochraceous apical ring, inner area with three diffuse black spots, lower inner area 
reddish; hind tibia ochraceous in basal half interrupted by broad black ring one third the way 
along, apical one third reddish. 

$. As male but larger; eye a little more ellipsoid than male, about one and a half times as 
high as maximum width and one and a half times as high as length of subocular groove. Sper- 
matheca as in Text-fig. 56. 




54 





Ejs Cpr Sps 



FIGS. 52-56. A. oxianus Uv., genitalia. 52, $ epiphallus ; 53, $ phallic complex, dorsal 
view, epiphallus and ectophallic membrane removed; 54, same, lateral view; 55, <J endo- 
phallus, lateral view; 56, $ spermatheca. 



f 


^ 


i 








"\ 


r 








N 


Min. 


Ave. Max. 


Min. 


Ave. 


Max. 


Min. 


Ave. 


Max. 


I -12 


I -l6 I -21 


3 


97 


4-08 


4' 


18 


4 


82 


4 


99 


5-21 


i -06 


1-13 1-23 


3 


95 


4-07 


4' 


30 


4 


-67 


4 


94 


5-27 



332 D. HOLLIS 

Measurements (mm.). Length of body, $ 18-6-22-0, $ 27-6-29-5. Length of pronotum, 
$3-7-4-4, $4-8-5-8. Length of tegmen, $18-6-21-2, $24-5-27-1. Length of hind femur, 
$ 11-3-12-7, $ 14-2-16-1. Maximum width of hind femur, $ 2-7-3-2, $ 3-4-3-9. Maximum 
width of head, $ 3'3~3'7, $ 4-2-4-8. 

Ratios (eight males and twelve females measured) : 

P/C FL/FW E/P E/F 

Min. Ave. Max. 
1-59 1-67 1-73 
1-64 1-72 1-79 

Discussion. Only a few specimens of this species have been examined and varia- 
tion has been observed in size and general colouration. 

The holotype of A. oxianus is deposited in the Leningrad Academy of Sciences 
and due to institutional policy was not available for study. However, from a study 
of the description and some paratypes in the BM(NH) it is clear that the name 
oxianus should be applied to the taxon described above. 

Superficially A. oxianus strongly resembles Epacromius tergestinus (Charp.) but 
the form of the fastigial foveolae, the position of the intercalary vein of the medial 
area of the tegmen and the form of the male subgenital plate show that the species 
is more naturally placed in the genus Aiolopus. By its slender form and weakly 
saddle-shaped pronotum A . oxianus would appear to be closely related to A . thalas- 
sinus, from which species it may be distinguished by the elongate fastigial foveolae, 
longer antennae, completely sulcate frontal ridge and small arolium. 

Distribution. (Thirteen males and twenty-two females). 

USSR : Kerki, on River Amu-Darya, 4 <$, 3 $, vii ; Tchardzhui, on River Amu- 
Darya, i <$, 2 $, viii; Syr Darya, Nadeshdinsky, 2 $, vi; Samarkand, R. Zaravshan, 
i $, vii; Khiva Town, 7 J, 7 $, " at light ", vii; Khiva distr., Gudzha, i <$, 3 $, vii; 
Karmysh, 2 $, vii; Turkestan, i . 

Aiolopus carinatus (Bei-Bienko, 1966) comb. n. 

(Text-figs. 19, 20, 29, 57-60) 
Aeoloptilus carinatus Bei-Bienko, 1966 : 1793, fig. 7. 

Type locality. Komodo Islands, mountain plateau, 500-600 m. ; type $ deposited 
in Leningrad. 

Redescription. $. Integument moderately rugulose. Antenna as long as combined lengths 
of head and pronotum, with twenty- two segments. Fastigium pentagonal, forward angle 
rounded, weakly concave with margins fairly well developed at least in front of eyes; fastigial 
foveolae subrectangular with rounded angles, shallow, about one and a half times as long as 
maximum width; frontal ridge flat or weakly convex, very slightly narrowing upwards (Text-fig. 
29). Eye ellipsoid, about one and half times as high as maximum width and almost two and a 
half times as high as length of subocular groove. Pronotum with dorsum weakly saddle-shaped 
(Text-figs. 19-20), prozona about three quarters the length of the metazona, latter with obtuse- 
angular posterior margin ; lateral carinae moderately developed in prozona, parallel or very weakly 
divergent in front of first transverse sulcus and weakly divergent back to posterior transverse 



REVISION OF GENUS AIOLOPUS 



333 



sulcus, almost obliterated in metazona; lateral plate of pronotum slightly higher than wide; 
mesosternal interspace slightly wider than long. Tegmen long, well exceeding apex of hind 
femur. Hind femur narrow, about four times as long as maximum width; hind tibia almost as 
long as hind femur, with twelve outer and twelve or thirteen inner spines, apical spurs simple 
with outer pair about one and a half times as long as inner pair; arolium half length of claw. 
Supra-anal plate, cercus and subgenital plate normal for genus. Phallic complex (Text-figs. 
57-60) with zygoma of cingulum without dorsal processes and apodemes laterally flattened; 
basal valves of penis with large lateral expansions which are not recurved. 

General colouration brown with dark brown and ochraceous markings. Lateral carinae of 
pronotum pale brown or ochraceous ; tegmen with costal stripe ochraceous and complete, proximal 
half of tegmen with dark brown area extending from costa to cubital vein, vannal area hyaline, 
distal half of tegmen mottled hyaline and brown ; hind femur brown with pregenicular ochraceous 
ring, upper outer area with two weak triangular brown spots, inner area ochraceous with two 
incomplete dark bands, lower inner area with weak orange tinge; hind tibia narrowly black 
basally, followed by broad ochraceous ring, apical two thirds greyish blue, sometimes apical 
quarter has weak orange tinge ; hind wing hyaline with darker distal area. 

$. As male but slightly larger and more robust; ovipositor normal for genus. 




57 




Apd 




FIGS. 57-60. A. carinatus (Bei-Bienko), male genitalia. 57, epiphallus; 58, phallic com- 
plex, dorsal view, epiphallus and ectophallic membrane removed; 59, same, lateral view 
60, endophallus, lateral view. 



334 D - HOLLIS 

Measurements (mm.). Length of body, $ 16-8, $ 20-6-21-4. Length of pronotum, 3-0, 
$ 3'6-3'8. Length of tegmen, $ 14-8, $ 16-5-17-5. Length of hind femur, $ 10-0, $ 11-4-11-9. 
Maximum width of hind femur, $ 2-3, $ 2-5-2-8. Maximum width of head, <J 2-7, $ 3'2-3'3. 

Ratios (one male and two females measured) : 

FL/FW E/P 





Min. Max. Min. Max. 

<J I'" 4'35 4'93 

$ 1-12 1-15 4-25 4-56 4-58 4-60 

Discussion. When originally described this species was thought to be generically 
distinct from Aiolopus, but for reasons explained above (p. 315) this generic separa- 
tion is not accepted by the present author. 

A. carinatus appears to be very closely allied to A. th. tamulus, from which it 
may be distinguished by the form of the fastigial foveolae, the pronotal carinae and 
the greater number of spines on the hind tibia. 

Distribution. KOMODO Is.: Rintja, i <^, i $, viii (Leningrad). SUMBA Is.: Melolo, 
i $, v (Willemse). 

Aiolopus longicornis Sjostedt, 1909. 

(Text-figs. 6, 17, 18, 26, 61-69) 
Aiolopus longicornis Sjostedt, 1909 : 156, 169. 

Type locality. This species was described from two males and two females from 
Zanzibar and one female from Mombo, Usambara ; one male specimen from Zanzibar 
bears Sjostedt's type label, and this specimen is selected as LECTOTYPE and is 
deposited in Stockholm. 

Redescription. <J. Larger than average size for genus. Integument more strongly rugulose. 
Antenna clearly longer than combined lengths of head and pronotum, with twenty-two to 
twenty-four segments, the median segments elongate and about three times longer than broad. 
Fastigium broadly pentagonal, little longer than wide, moderately concave, strongly sloping 
forwards and roundly merging with frons, marginal carinulae well developed ; fastigial foveolae 
almost rectangular, hardly narrowing forwards, weakly concave with moderately developed 
margins, the lower one often weak; frons oblique; frontal ridge coarsely and densely pitted, 
flat but impressed at median ocellus, merging with genae well before clypeus, lateral margins 
weakly convergent at apex (Text-fig. 26). Eyes oval, slightly less than one and a half times as 
high as maximum width and about twice the length of the subocular groove. Pronotum (Text- 
figs. 17-18) short and wide, hardly more than one and one third times longer than maximum 
width, moderately saddle-shaped, with prozona constricted medially; metazona with obtuse- 
angular posterior margin; lateral plate of pronotum higher than wide; mesosternal interspace 
slightly wider than long, weakly trapezoid, widening posteriorly. Tegmen relatively long (see 
E/P and E/F ratios). Hind femur (Text-fig. 6) moderately broad, FL/FW ratio about 3-7; 
hind tibia almost as long as hind femur, with nine outer and ten inner spines, inner apical spurs 
almost twice as long as outer spurs ; arolium about half length of claw. Phallic complex (Text- 
figs. 61-64) with zygoma of cingulum with well developed dorsal processes and short apodemes; 
basal valves of penis hardly expanded laterally. 

General colouration brown with ochraceous (or rarely green) and blackish markings; median 
longitudinal ochraceous stripe on head and pronotum normally well defined, ochraceous X-shaped 



REVISION OF GENUS AIOLOPUS 



335 




62 



Ejs Cpr Sps 



Ap 





66 





68 



FIGS. 61-68. A. longicornis Sjost., genitalia. 61, <$ epiphallus ; 62, phallic complex, 
dorsal view, epiphallus and ectophallic membrane removed; 63, same, lateral view; 
64, $ endophallus, lateral view; 65, spermatheca of specimen from Chad area; 66, same, 
from Congo; 67, same, from Rukwa, Tanzania; 68, same, from Tingida, Tanzania. 



336 D. HOLLIS 

pattern on dorsum of pronotum clearly defined; tegmen with two V-shaped ochraceous areas 
extending from anterior border to mid-line, apical half mottled, first and second vannal areas 
often ochraceous and when tegmen are folded this colouration forms a continuous stripe with 
that of head and pronotum; hind wing hyaline, sometimes with faintly yellowish base and 
sometimes with darkened apex and posterior border; hind femur with three weak black spots 
on upper outer area, external area mottled ochraceous black with the black colour often weak, 
ochraceous pregenicular ring distinct, apex black ; inner area with complete preapical black band 
and medial black spot in dorsal half, lower inner area reddish ; hind tibia with narrow basal black 
ring followed by broad ochraceous ring and then a broad black ring, ochraceous medially and 
apical third reddish. 

$. Larger and more robust than male. Spermatheca as in Text-figs. 65-68, with apical 
diverticulum varying from a small bubble-like projection to a finger-like tube. 

Measurements (mm.). Length of body, $ 18-8-26-5, ? 2 5'3~3 1 '7- Length of pronotum, 
6* 3'7-5 'Qi ? 4'4-5'5- Length of tegmen, $ 18-6-24-5, $ 23-7-27-8. Length of hind femur, 
<J 11-1-16-0, $14-8-17-6. Maximum width of hind femur, $ 3-3-4-3, $4-0-4-8. Maximum 
width of head, <J 3-6-5-0, $ 4-2-5-8. 

Ratios (fifty males and twenty females measured) : 

P/C FL/FW E/P E/F 



c 






^ 


r~ 








\ 


( 




i 


Min. 


Ave. 


Max. 


Min. 


Ave. 


Max. 


Min. 


Ave. Max. 


0-93 


1-05 


i 


16 


3 


55 


3 


70 


3'97 


4 


'74 


5'3 5 '5 


0-92 


1-04 


i 


09 


3 


53 


3 


68 


3-84 


4 


-87 


5-14 5-48 



Min. Ave. Max. 
1-47 1-58 1-67 
1-56 1-61 1-66 

Discussion. This species is intermediate between the thalassinus group and the 
simulatrix group, having the relatively longer hind femur and tibia and the saddle- 
shaped pronotum of the former and the stouter build and more strongly rugulose 
integument of the latter. A. meruensis is possibly its closest relative, also having a 
relatively broad head and short pronotum, but longicornis may be distinguished 
from meruensis by the broad frontal ridge, the elongate median segments of the 
antennal flagellum, the relatively longer tegmen and the narrower hind femur. 

Distribution (Text-fig. 69). Two hundred and eighty-one specimens of this species 
were examined, indicating the following distribution: 

NIGERIA (Chad area), February; CONGO, (former Belgian), August; ETHIOPIA, 
September, December; SOMALIA, August, October; KENYA, January, March, May, 
October; UGANDA, April to July; TANZANIA, throughout the year; BURUNDI, April; 
ZAMBIA, March, August. 

Aiolopus meruensis Sjostedt, 1909 
(Text-figs. 69-74) 

Aiolopus meruensis Sjostedt, 1909 : 156, 170. 
Aeolopus latus Uvarov, 1922 : 545, syn. n. 

Type locality. Meru (Tanzania) : Flodhastsjoarne; type $ deposited in Stockholm. 

Redescription. <J. Integument moderately rugulose. Antenna at most as long as combined 
lengths of head and pronotum, with twenty-one to twenty- three segments. Fastigium of vertex 
rounded pentagonal, about as long as wide, weakly concave, with well defined margins ; fastigial 
foveolaetrapezoid, broad, barely one and a half times longer than wide, narrowing forwards, margins 



REVISION OF GENUS AIOLOPUS 



337 



well defined but sometimes lower margin weak ; frontal ridge flat or weakly sulcate at and below 
median ocellus, lateral margins gradually and continuously convergent upwards, moderately 
coarsely pitted. Eyes one and a half times as high as maximum width and almost twice as long 
as subocular groove. Pronotum slightly saddle-shaped, broad; metazona about one and two 
thirds the length of prozona, with obtuseangular or almost rounded posterior margin; lateral 
plate of pronotum slightly higher than wide ; mesosternal interspace wider than long, trapezoid, 
widening posteriorly. Tegmen relatively short, E/F ratio about 1-47. Hind femur broad, 
ratio of length to width about 3-39; hind tibia almost as long as hind femur, with nine to ten 
outer and ten to eleven inner spines, inner apical spurs about twice as long as outer spurs; 



^-meruensis 
- long (corn is 




30' 



20 10" 



10* 20 s 30 40" 50 60* 



FIG. 69. Aiolopus spp., distribution map. 



338 D. HOLLIS 

arolium half length of claw. Supra-anal plate, cercus and subgenital plate normal for genus, 
Phallic complex (Text-figs. 70-73) with zygoma of cingulum with well developed dorsal processes 
and lateral expansions at bases of apodemes, latter short; basal valves of penis with rounded 
lateral expansions. 

General colouration brown with ochraceous or greenish markings; head and pronotum with 
median longitudinal pale stripe, pronotum with obvious X-shaped ochraceous pattern; tegmen 
with pale stripe in first vannal area and two transverse pale bands, apical third mottled ; hind wing 
hyaline, often weakly pale yellow basally and with infumate apex; hind femur on outer side 



-Bp 




Apa 



72 





,- r' 

Gpr Sps 



FIGS. 7074. A. mervensis Sjost., genitolia. 70, ^epiphallus ; 71, <Jphallic complex, 
dorsal view, epiphallus; and ectophallic membrane removed; 72, same, lateral view; 
73, (J endophallus, lateral view; 74, ? spermatheca. 



REVISION OF GENUS AIOLOPUS 339 

with weak and diffuse basal, medial and apical obliquely transverse bands, on inner surface with 
two incomplete dark bands, lower inner area reddish ; hind tibia ochraceous in basal third, then 
with incomplete blackish ring, apical one third to two thirds reddish. 

$. As male but larger and more robust. Eyes slightly less than one and a half times higher 
than wide and less than twice as high as length of subocular groove ; spermatheca as in Text-fig. 

74- 

Measurements (mm.). Length of body, 16-0-21-2, $ 21-2-29-7. Length of pronotum, 
cJ 3-2-4-1, $ 3-7-5-4. Length of tegmen, $ 14-3-19-8, $ 14-5-23-1. Length of hind femur, 
,3 9-5-12-8, $11-2-15-9. Maximum width of hind femur, < 2-6-3-8, $3-2-4-8. Maximum 
width of head, ^2-9-4-0, $3-7-5-1. 

Ratios (fourteen males and twenty-nine females measured) : 

P/C FL/FW E/P E/F 



Min. Ave. Max. Min. Ave. Max. Min. Ave. Max. Min. Ave. Max. 

$ 1-02 1-09 I-I7 3-18 3-39 3-66 3-76 4-42 4-89 1-28 1-47 1-64 

$ 0-94 1-04 1-15 3-08 3-37 3-68 3-45 4-35 5-07 1-29 1-47 1-66 

Discussion. In having a short, broad, slightly saddle-shaped pronotum, a broad 
hind femur and a hind tibia almost as long as the hind femur, A . meruensis is probably 
more closely related to A. longicornis Sjostedt than to any other species in the genus; 
it is differentiated from longicornis by the narrower frontal ridge, the much shorter 
antennae, the shorter tegmen and the broader hind femur. 

A. meruensis is variable over its range of distribution but the variations do not 
follow any particular geographical pattern. The form of the frontal ridge may vary 
from flat to weakly sulcate. The posterior margin of the pronotum varies from 
obtuseangular to almost rounded. The relative length of the tegmen is extremely 
varied; normally the tegmen extends to just beyond the tip of the hind femur but 
in the single specimen examined from Kilimanjaro it barely reaches the apex of the 
abdomen, in the specimens examined from Bloemfontein it just reaches the apex 
of the hind femur and in those from Lake Manyara, Tanzania, it extends well beyond 
the apex of the hind femur. 

The original description of this species was not very diagnostic and caused it to 
be previously confused with A . thalassinus thalassinus. Uvarov's type $ of A . latus 
described from Bloemfontein and deposited in the BM(NH), when considered to- 
gether with the other material of meruensis shows no significant difference and is 
therefore synonymized with meruensis. 

Distribution (Text-fig. 69). The forty-five specimens examined were collected 
in the following countries : 

KENYA, August; TANZANIA, January, April; RHODESIA, April; MOZAMBIQUE, 
February, May, July; SOUTH AFRICA, January, April, June, September, November; 
S.W. AFRICA, March. 



Aiolopus thalassinus (Fabricius, 1781) 

This species is divided into four subspecies which are described and discussed 
below. 

ENTOM. 22, 7 17 



340 D. HOLLIS 

Aiolopus thalassinus thalassinus (Fabricius, 1781) stat n. 
(Text-figs. 1-5, n, 12, 21, 75-84. 9. 9 1 ) 

Gryllus thalassinus Fabricius, 1781 : 367. 

Acridium grossum Costa, 1836 : 25, pi. 3, fig. 4, a-d; Fischer, L. H., 1853 : 361. 

Acridiumlaetum Brulld, 1840 : 77, pi. 5, figs, zo-ioa; Finot, 1895 : 423. 

Epacromia angustifemur Ghiliani, 1869 : 179; Kirby, 1910 : 191. 

Ochrophlebia (?) savignyi Krauss, 1890 : 261 (Savigny, Desc. Egypt, pi. 6, fig. 15 (3)); Storey, 

1919 : 54- 

Epacromia lurida Brancsik, 1895 : 2 5 syn. n. 

Aiolopus thalassinus kivuensis Sjostedt, 1923 : 18; Sjostedt, 1929 : 24. 
Aiolopus acutus Uvarov, 1953 : IIX > fi s - ^Q-JS 1 . syn. n. 

Type locality. The type was in Allioni's collection in Turin but has been destroyed 
by fire. NEOTYPE (?) erected bearing the following data : SWITZERLAND : Locarno, 
Maggia Delta, i6.ix.iQ29 (Zeuner); it is deposited in the BM(NH). 

Redescription. <$. Moderately rugulose. Antenna as long as combined lengths of head and 
pronotum, with twenty-two to twenty-four segments. Fastigium pentagonal, forward angle 
rounded, or subangular in specimens from Ethiopian and Oriental regions, moderately concave 
with well defined margins; fastigial foveolae trapezoid, normally slightly narrowing forward, 
about twice as long as maximum width; frontal ridge flat or slightly convex, sparsely pitted, 
lateral margins parallel except at apex where they converge slightly (Text-fig. 21). Eye oval, 
about one and a half times as high as maximum width and almost twice as high as length of 
subocular groove. Pronotum slightly saddle-shaped; metazona almost one and two thirds as 
long as prozona; latter constricted medially and with weak lateral " shoulders "; posterior 
margin of metazona obtuseangular (Text-figs. 11-12); lateral plate of pronotum slightly higher 
than wide; mesosternal interspace almost square. Tegmen long, well exceeding tip of hind 
femur. Hind femur narrow, about four times as long as maximum width (for range see ratios 
below) ; hind tibia as long as or only very slightly shorter than hind femur (Text-fig. 5), with ten 
outer and eleven inner spines, inner apical spurs about one and half times as long as outer spurs ; 
arolium half length of claw. Supra-anal plate, cercus and subgenital plate normal. 

Phallic complex (Text-figs. 75-78) with epiphallus having moderately developed outer lophal 
lobe; zygoma of cingulum with longer apodemes and poorly developed dorsal processes; basal 
valves of penis with lateral expansions strongly developed and recurved. 

General colouration green or brown with ochraceous and black markings. Pronotum with or 
without median longitudinal ochraceous stripe and with or without ochraceous criss-cross pattern 
on dorsum, if without then dorsum unicolorously green or ochraceous and with blackish bordering 
colouration below "shoulders"; costal area of tegmen with interrupted ochraceous stripe, 
generally mottled and without clearly defined transverse bands; hind wing hyaline, sometimes 
weakly yellowish green basally and often with darkened apex; hind femur on upper outer area 
with two triangular black spots which continue onto outer area as incomplete oblique fasciae 
and on inner area as complete fasciae, apex of hind femur blackish with preceding complete 
or incomplete ochraceous ring ; hind tibia narrowly black basally, followed by broader ochraceous 
ring, apical two thirds normally reddish, the red colouration separated from ochraceous ring by 
greyish ring. 

$. As <^ but larger and more robust; frontal ridge often more convex; spermatheca (Text-figs 
79-83) variable, with pre-apical diverticulum of varying lengths; ovipositor as in Text-figs. 2-3. 

Measurements (mm.). Length of body, $ 15-2-21-2, $ 19-8-29-3. Length of pronotum, 
cJ 2-8-4-0, $ 3-8-5-5. Length of tegmen, <J 14-2-20-5, $ 17-0-26-3. Length of hind femur, 
<J 9-5-12-2, $ 10-9-16-5. Maximum width of hind femur, $ 2-3-3-0, $ 2-7-3-8. Maximum 
width of head, $ 2-6-3-4, ? 3 -I -4'5- 



REVISION OF GENUS AIOLOPUS 



Ratios (forty specimens of each sex measured) : 
P/C FL/FW 



E/P 



E/F 



Min. 



Max. 



1-04 
i-io 



{ 










^ 


/" 






^ 


Min. 


Ave. 


Max. 


Min. 


Ave. 


Max. 


4' 


'57 


4 


99 


5 


'44 


1-42 


i -62 


I 


81 


4 


25 


4 


86 


5 


22 


1-53 


i -67 


I 


83 



Ave. Max. Min. Ave. 

1-18 1-27 3-81 4-06 4-33 

1-20 1-37 3 - 8o 4-10 4-50 

Discussion. A. th. thalassinus is widely distributed in the southern palaearctic, 
ethiopian and western oriental regions (see Text-fig. 84), and consequently shows 
great variability. Its general colouration varies from almost completely green, 

Apd 




79 



FIGS. 75-83. A. thalassinus thalassinus (Fabr.), genitalia. 75, $ epiphallus; 76, # phallic 
complex, dorsal view, epiphallus and ectophallic membrane removed; 77, same, lateral 
view; 78, $ endophallus, lateral view; 79, $ spermatheca of specimen from France; 
80, same, of specimen from Morocco; 81, same, of specimen from Kenya; 82, same, of 
specimen from South Africa; 83, same, of specimen from India. 

ENTOM. 22, 7 I7 



342 D. HOLLIS 

through green with brown markings, to brown with ochraceous and black markings. 
The pronotum always has a contrasting pattern dorsally and laterally suggesting 
the presence of lateral carinae. The hind tibia sometimes lacks its reddish coloura- 
tion, cream or grey being the replacement colour. The anterior angle of the fasti- 
gium varies from rounded to quite sharp, specimens with the latter character having 
been described as a distinct species, A. acutus Uvarov (1953). However, as all 
intermediate forms of fastigium can be found and there is no geographical evidence 
to support this separation, Uvarov's species is synonymized. The frontal ridge 
varies from slightly convex to flat with a depression at the median ocellus. 

A small series from Angola, collected from a mangrove swamp 4 mis. S.E. of 
Luanda, which was studied shows rather greater than average variation in the length 
of the pronotum and tegmen and the width of the hind femur. 

Ratios of measurements of Angola specimens : 



P/C 

Min. Ave. Max. 
1-03 i -06 1-09 
i -oo 1-04 i -08 


FL/FW 

Min. Ave. Max. 
3-81 3-86 3-92 
3-90 3-97 4-05 


E/P 

Min. Ave. Max. 
4-28 4-49 4' 8 7 
4'34 4'39 4'43 


E/F 


c i 
Min. Ave. Max. 

1-40 1-46 i -49 
1-40 1-43 1-47 



As only six males and three females were examined and other specimens from other 
parts of Angola appear quite normal, no taxonomic significance is attached to this 
series at present. 

In previous publications the lists of synonymy for this subspecies are at variance 
with the list given in the present paper; the latter is arrived at as follows: 

Gryllus prasinus Thunberg is removed from the synonymy of this subspecies and 
synonymized with A. strepens (Latreille). 

The type of Oedipoda pulverulenta Fischer de Waldheim, 1846 is deposited in 
Leningrad and because of institutional policy was not available for study. Professor 
G. Ya Bei-Bienko has examined the specimen and reports that it does not belong 
to the genus Aiolopus, as thought by Fischer, L. H. (1853 : 36), and is to be referred 
to the genus Epacromius. 

The type of Acridium grossum Costa is destroyed and Fischer's synonymy is 
accepted. Similarly the type of Acridium laetum Brulle is destroyed and Finot's 
synonymy is accepted. 

The type of Epacromia angustifemur Ghiliani, which is deposited in Turin, is also 
unavailable for study, but from the original description it is clear that Kirby's 
synonymy is correct. 

Ochrophlebia (?) savignyi Krauss was described from a drawing by Savigny 
(1809-13 : pi. 6, fig. 15 (3) ) and after examination of Savigny's figure Storey's 
synonymy is accepted. 

The type of Epacromia lurida Brancsik was deposited in Budapest but has been 
destroyed. However, Uvarov (1953 : 109) states that he examined the type and 
found it " appears to be a species of the thalassinus group ". The type locality 
is " Boroma, Zambesi ", and after examination of material from areas close to the 
type locality the present author synonymizes this name with A. thalassinus. 



REVISION OF GENUS AIOLOPUS 343 

A. th. kivuensis Sjostedt was described from a single female specimen from Lake 
Kivu, in the former Belgian Congo, and is deposited in Stockholm. An examination 
of Sjostedt's type has shown that its present synonymy is correct. 

A. acutus Uvarov was described from a series of specimens from Katanga, in the 
former Belgian Congo, and the type $ is deposited in the BM(NH). It is synony- 
mized for the reasons stated above. 

Distribution (Text-fig. 84). From the four thousand specimens examined the 
following general conclusions are drawn : 

FRANCE, June to November; SPAIN, April, September, October; BALAERIC Is., 
September, October; PORTUGAL, August; SWITZERLAND, September; ITALY, Sep- 
tember to November; SARDINIA, September; AUSTRIA, August; CZECHOSLOVAKIA, 
August; HUNGARY, March, July, October; YUGOSLAVIA, May to June, August to 
September; ALBANIA, June, August to September; GREECE, May to August; CYPRUS, 
May, August, October; TURKEY, June to September; USSR, June to September; 
KASHMIR, May to October; NEPAL, July, September to October; INDIA, February 
to December; WEST PAKISTAN, May through to January; AFGHANISTAN, April; 
PERSIA, March to December; IRAQ, May to July, September to December, February; 
SYRIA, April to August; LEBANON, March, May to June; ISRAEL, May to June 
August, October, January; JORDAN, March to April, June; SAUDI ARABIA, All year 
round; YEMEN, June, August to September, December through to April; SOUTHERN 
YEMEN, May through to March ; MUSCAT AND OMAN, August to September, January 
to May; BAHREIN, May; SOCOTRA, August, January to April; EGYPT, May to Nov- 
ember, January to February; LIBYA, September, February to July; TUNISIA, March 
to April; ALGERIA, October, March to July; MOROCCO, June to July, November; 
MADEIRA, July to August, November to December; CANARY Is., July through to 
May; CAPE VERDE Is., No dates; MALI, May through to January; SENEGAL, August 
to September; GAMBIA, April; GUINEA, No dates; SIERRA LEONE, No dates; GHANA, 
All year round; TOGO, February, May; NIGERIA, May to October, December through 
to March; SUDAN, May to August, November to December, February; ETHIOPIA, 
May through to March; SOMALIA, June, August through to March; KENYA, January 
to November; UGANDA, May through to March; RUANDA, August; CONGO, May to 
October, December, February; ANGOLA, March to November; ZAMBIA, October 
through to August; TANZANIA, All year round; MALAWI, June, December; MOZAM- 
BIQUE, June to August, October to November, February, April; COMORO Is., 
September to October; S. W. AFRICA, June, October, December; BOTSWANA, May, 
October; RHODESIA, October through to August; SOUTH AFRICA, August through 
to June. 

Aiolopus thalassinus rodericensis (Butler, 1876) stat. n. 
(Text-figs. 84-91) 

Epacromia rodericensis Butler, 1876 : 410. 

Epacromia famulus [sic] var. pusilla I. Bolivar, 1895 : 378; I. Bolivar, 1912 : 270. 

Chortoicetes rodericensis (Butler, 1876) Kirby, 1910 : 193. 

Aeolopus aldabrensis I. Bolivar, 1912 : 269; Dirsh, 1963 : 264. 



D. HOLLIS 




REVISION OF GENUS AIOLOPUS 345 

Aeolopus dociostauroides I. Bolivar, 1912 : 269; Dirsh, 1963 : 264. 
A eolopus fasciatipes I. Bolivar, 1912 : 270; Dirsh, 1963 : 264. 
A eolopus perpusillus I. Bolivar, 1912 : 270; Dirsh, 1963 : 264. 
Aiolopus rodericensis (Butler, 1876) Uvarov, 1928 : 364. 

Type locality. Rodriguez Is., lectotype <$ designated by Dirsh (1963) and depos- 
ited in the BM(NH). 

Differs from nominate subspecies in the following ways: 

cJ. Fastigial foveolae trapezoid, narrowing forwards, about one and a half times as long as 
maximum width, very shallow; frontal ridge finely and sparsely pitted, gradually and continu- 
ously narrowing upwards, slightly convex, flat or slightly sulcate at and below median ocellus 
and with very weak lateral carinulae; mesosternal interspace slightly wider than long; tegmen 
shorter, E/P ratio averaging 4-60; hind femur wider, ratio of length to maximum width averaging 
3-78. Phallic complex (Text-figs. 85-88) with basal valves of penis hardly expanded and not 
recurved. 

9. E/P ratio averaging 4-58; FL/FW ratio averaging 3-79; spermatheca as in Text-fig. 89. 

General colouration ochraceous and brown or greenish with median longitudinal ochraceous 
stripe which extends forwards weakly onto head ; pronotum normally with cross-shaped ochraceous 
pattern on dorsum or if without then median stripe widens forwards in prozona. Hind femur 
with three incomplete dark fasciae on outer surface which are often weak and irregular; inner 
surface with three black fasciae, apical one always complete, median fascia often weak, and 
basal fascia either occasionally absent or complete and partially fused with median fascia ; hind 
tibia with narrow basal black band follosed by broad ochraceous band, then with narrow 
incomplete black band, then broad bluish band which continues dorsally to apex of tibia but is 
interrupted apico-ventrally by blackish area. Tegmen with three incomplete dark bands, the 
apical one often dispersed, costal area with broken ochraceous or green stripe. 

Measurements (mm.). Length of body, < 13-8-21-5, 9 19-2-28-7. Length of pronotum, 
cJ 2-8-4-2, 9 3-8-5-4. Length of tegmen, <$ 12-4-20-6, 9 16-4-24-9. Length of hind femur, 
cJ 7-8-12-7, 9 11-4-15-6. Maximum width of hind femur, <$ 2-3-3-4, ? 3'-4 -I - Maximum width 
of head, $ 2-5-3-6, 9 3'3~4'6. 

Ratios (twenty specimens of each sex measured) : 

P/C FL/FW E/P E/F 





( 


~\ 


f 




-\ 


1 




> 


( 




i 




Min. Ave. 


Max. 


Min. 


Ave. 


Max. 


Min. 


Ave. 


Max. 


Min. 


Ave. 


Max 


cJ 


i -06 1-15 


1-23 


3-39 


3-78 


4-12 


4-27 


4-60 


4-90 


1-41 


1-51 


i -67 


9 


1-07 1-12 


I -21 


3*64 


3'79 


4-00 


4-09 


4-58 


5-08 


1-41 


1-58 


1-79 



Discussion. This subspecies is quite variable in size, colouration and degree of 
flatness of the frontal ridge. Dirsh (1963 : 267), in discussing the species he syn- 
onymized with A. rodericensis, states " Other characters, such as width of frontal 
ridge which depends upon the stoutness of head do not exceed the range of varia- 
bility; A. laticosta (I. Bol.) being one of the extreme variants in this respect ". In 
fact it is the overall shape of the frontal ridge which is different in laticosta, and 
coupled with the short hind tibia and the form of the pronotum the present author 
disagrees with Dirsh's synonymy and places laticosta into synonymy with A. 
simulatrix (Walker). 

A. th. rodericensis has been previously regarded as a species and Dirsh (1963) 
considered it to belong to the " savignyi group " by virtue of its broad hind femur. 

The present author regards this taxon as a subspecies of A . thalassinus ; the range 



346 



D. HOLLIS 



of stoutness of the hind femur falls between that of A. simulatrix (A. savignyi) 
and that of A . thalassinus, the hind tibia is not as short as in A . simulatrix, and the 
form of the frontal ridge and pronotum are of the same type as A. thalassinus. 

Morphological differences between A . th. rodericensis and the mainland subspecies 
are not very clearly defined; in rodericensis the head tends to be broader and this 
is demonstrated in the P/C ratio, the tegmen is shorter as is demonstrated in the 
E/P ratio, the hind femur is broader as is shown in the FL/FW ratio, the fastigial 
foveolae are slightly more square and there is no red colouration on the hind tibia. 





--Apd 



86 



Cv 




Ejd Ejs Gpr 



FIGS. 85-89. A. thalassinus rodericensis (Butler), genitalia. 85, <$ epiphallus; 86, <$ phallic 
complex, dorsal view, epiphallus and ectophallic membrane removed; 87, same, lateral 
view; 88, <$ endophallus, lateral view; 89, $ spermatheca. 



REVISION OF GENUS AIOLOPUS 347 

A comparison of the E/P and FL/FW ratios for both thalassinus and rodericensis is 
given in Text-figures 90 and 91. 

Certain very general trends are visible in this subspecies across its range from 
Madagascar to the Seychelles Islands; as the populations extend eastwards the 
individuals tend to become smaller, and the tegmen becomes relatively shorter 
and the frontal ridge flatter ; the latter even tends to have very weak lateral carinulae 
at and below the median ocellus. However, these trends are not always complete, 
and specimens showing characters against the general trends are often encountered. 

Type localities of synonymized species. 

Epacromia famulus [sic] var. pusilla I. Bol. Lectotype <$, Mahe (Seychelles), 

designated by Dirsh (1963 : 267) and deposited in Paris. 
Aeolopus aldabrensis I. Bol. Lectotype $, Aldabra, designated by Dirsh (1963 : 267) 

and deposited in BM(NH). 
Aeolopus dociostauroides I. Bol. Lectotype ^, Coetivy (Seychelles), designated by 

Dirsh (1963 : 267) and deposited in BM(NH). 
Aeolopus fasciatipes I. Bol. Lectotype $, Farquar Atoll, designated by Dirsh 

(1963 : 267) and deposited in BM(NH). 

Distribution (Text-fig. 84). Two hundred and thirty-one specimens of this sub- 
species were examined and the following conclusions concerning its distribution 
were made : 

MADAGASCAR, April to June, November through to February; REUNION, June; 
MAURITIUS, January, May; RODRIGUEZ Is., February; ALDABRA GROUP, May to 
December; SEYCHELLES GROUP, February to December. 

Aiolopus thalassinus tamulus (Fabricius, 1798) stat. n. 
(Text-figs. 22, 23, 84, 92-96) 

Gryllus tamulus Fabricius, 1798 : 195. 

Gomphocerus tricoloripes Burmeister, 1838 : 649; Kirby, 1910 : 192. 

Epacromia rufostriata Kirby, 1888 : 550, syn. n. 

Type locality. " Ind: or. et China ", lectotype <$ designated by Key (1967) and 
deposited in the Copenhagen Mus. 

Differs from nominate subspecies in the following ways: 

<J. Fastigium with forward angle more acute (Text-figs. 22-23) ; fastigial foveolae narrowing 
more strongly anteriorly (Text-figs. 22-23), frontal ridge flat and continually narrowing upwards 
(Text-figs. 22-23) I or weakly sulcate and with weak lateral carinulae, and junction with fastigium 
more angular; pronotum with " shoulders" of prozona more parallel in prozona and sometimes 
even with very weak lateral carinae in prozona; posterior margin of pronotum more rounded. 
Phallic complex (Text-figs. 92-95) with basal valves of penis less expanded laterally and not 
recurved. 

$. Spermatheca as in Text-fig. 96. 

General colouration differs in that ochraceous or green stripe along costal area of tegmen is 
usually complete; external area of hind femur without oblique fasciae; hind tibia with red 



348 



D. HOLLIS 



4.4. 

43- 
4-2- 
4- 
4O 
3'9 
38 
A 3-7 

36 

FL 




o 
o 
o 
x o 



*>o 
x o o 



o o o 
x o 

X x X o 

X x 
x x x x 

x x y 


x rodericensis 

thalassinus 
(ethiopiari) 

o thalassinus 
(palaearctic) 


FW 3 . 5 . 
34 


90 

4*5- 
4-4- 

4-3- 
4-2- 

4-1- 
A 4<) . 

3*9- 

FL 3-8- 
FW 

37 


4-3 4 : 4 4^5 4-6 4-7 4 : 8 4-9 5 ; O 5 ; I 5 ; 2 5 ; 3 5 ; 4 5^5 
E ^ 


P 





. 

oo . 

PO o . 



X O O 
00 

o x 2 


* x x rodericensis 
x 
xX x x x 9 thalassinus 
x (ethiopian) 
x xx x 
x o thalassinus 
* (palaearctic) 


91 


4-1 4-2 4 ; 3 4 : 5 4 ; 5 4 ; 6 4 ; 7 4 ; 8 4 ; 9 5O 5-1 5 : 2 5 : 3 5 ; 4 
E ^ 


P ^ 



FIGS. 90-91. A. thalassinus sspp., scatter diagrams comparing values of FL/FW 
against E/P. 90, males; 91, females. 



REVISION OF GENUS AIOLOPUS 



349 



colouration in apical third only or not at all, and broadly separated from median black band by 
broad bluish grey band. 

Measurements (mm.). Length of body, $ 17-9-22-2, $ 21-7-29-5. Length of pronotum, 
<J 3-3-4-3, $ 4'-5'3- Length of tegmen, <J 16-8-21-5, $ 19-8-27-0. Length of hind femur, 
( 10-5-13-1, $11-7-15-9. Maximum width of hind femur, (J 2-5-3-2, $2-9-3-8. Maximum 
width of head, $ 2-7-3-5, $ 3-5-4-5. 

Ratios (twenty specimens of each sex measured) : 



P/C 



FL/FW 



E/P 



f 






A 


1 








^ 


t 






Min. 


Ave. 


Max. 


Min. 


Ave. 


Max. 


Min. 


Ave. 


I'll 


I-2I 


I 


32 


4 


oo 


4-15 


4 


38 


4 


67 


5-0? 


I -12 


1-19 


I 


29 


4 


3 


4-21 


4 


53 


4 


69 


5-03 



Max. 




95 



96 



FIGS. 92-96. A. thalassinus famulus (Fabr.), genitalia. 92, <$ epiphallus ; 93, ^phallic 
complex, dorsal view, epiphallus and ectophallic membrane removed ; 94, same, lateral 
view ; 95, <J endophallus, lateral view ; 96, $ spermatheca. 



350 D. HOLLIS 

Discussion. This subspecies shows considerable variation in size and general 
colouration to the same extent as in the nominate subspecies. In the Australasian 
region the fastigium of the vertex tends to become more rounded, the frontal ridge 
narrows less conspicuously upwards and the hind tibia tends to lose the red coloura- 
tion. As a whole Australasian populations of tamulus show a general transition 
towards the Polynesian subspecies dubius but may be easily separated from the latter 
by the absence of a complete black-brown colouration on the ventral surface of the 
hind tibia. 

A . thalassinus tamulus was previously regarded as a distinct species and identified 
by the form of the frontal ridge and fastigium and the colouration of the hind tibia. 
These characters are fairly distinct in populations in east India and S.E. Asia but 
gradually become less distinct and merge with thalassinus in west India and, as 
mentioned above, with dubius in Polynesia. For this reason tamulus is here con- 
sidered as a subspecies of thalassinus. 

The type of Gomphocerus tricoloripes Burmeister could not be traced in the Halle 
Institute or the Berlin Museum but from the original description it is clear that 
Kirby's syononymy is correct. Gryllus dorsalis Thunberg was removed from the 
synonymy of this taxon and transferred to the genus Anaeolopus (Hollis, 1967). 
The type of Epacromia rufostriata Kirby, deposited in the BM(NH), differs from 
the lectotype of tamulus mainly in the form of the frontal ridge which in the former 
is very acute (Text-fig. 23). As this frontal ridge form is also displayed by spec- 
imens from Java and to a lesser extent by specimens from the Malayan mainland 
Kirby's species is here synonymized. 

A specimen from Timor, identified by C. Willemse as A. timorensis n. sp. (MS), is 
atypical in that the colouration is generally much darker, the pale stripe along the 
costal area of the tegmen is reduced to two pale spots and the hind tibia is without 
the normal bluish band and is orange in the apical half. Morphologically however 
the specimen agrees in detail with tamulus and the present author considers there 
is insufficient evidence at present upon which to base a separate taxon. 

Type localities of synonymized species. 

Gomphocerus tricoloripes Burm. Type not located. 

Epacromia rufostriata Kirby. Holotype <, Christmas Island (Indian Ocean), 
deposited in BM(NH). 

Distribution (Text-fig. 84) . From the five hundred and thirty specimens examined 
of this subspecies the following distributional conclusions were made : 

INDIA, March to December; CEYLON, June, September to November, January; 
EAST PAKISTAN, July to August; NICOBAR Is., March; ANDAMAN Is., February to 
March, June, September; BURMA, May, August, October; THAILAND, February, 
May to August, October; CHINA, June, August to November; HAINAN, April to 
June; HONG KONG, October; TAIWAN, August; MALAYA, June to August, October, 
December through to January; SINGAPORE, January, May; SUMATRA, March, Sep- 
tember, November; JAVA, September; LOMBOK, April; TIMOR, December; CHRISTMAS 
Is., (Indian Ocean), October, January to April; JAPAN, September, December; 



REVISION OF GENUS AIOLOPUS 351 

PHILIPPINE Is., February to May, August, December; MARIANA Is., January, May; 
BORNEO, January; BRUNEI, June; SABAH, February, July, CELEBES, June; NEW 
GUINEA, March, August; PAPUA, March; AUSTRALIA, January to March; LORD 
HOWE Is., November. 

Aiolopus thalassinus dubius Willemse, 1923 stat. n. 

(Text-figs. 24, 84, 97-102) 
Aeolopus dubius Willemse, 1923 : 100. 

Type locality. The species was described from several males and females from 
New Caledonia, Loyalty Islands and the New Hebrides. Efforts have been made 
to trace these specimens but only four females in the British Museum and a male and 
female in Willemse's collection were traced. Of these the following specimen is 
selected as a LECTOTYPE : $, Neukaledonen, Yate (F. Sarasin & J. Roux) ; it is 
deposited in the Maastricht Museum. 

Differs from the nominate subspecies in the following ways : 

$. Fastigial foveolae rectangular and, at most, one and a half times longer than wide ; frontal 
ridge very weakly sulcate (Text-fig. 24) ; prozona of pronotum often with weak lateral carinae 
between second and third transverse sulci. Phallic complex (Text-figs. 99-102) with basal 
valves of penis having rounded lateral expansions which are not recurved. 

$. Spermatheca as in Text-figs. 97-98. 

General colouration dark brown with ochraceous and black markings; hind femur normally 
with inner area black with two incomplete pale bands ; hind tibia black or dark brown on ventral 
and interno-ventral surfaces in apical two thirds; ochraceous costal stripe of tegmen complete. 

Measurements (mm.). Length of body, (J 16-4-20-4, $ 22-2-28-8. Length of pronotum, 
6* 3' 2 -3'9> $ 3 - 8-5-o. Length of tegmen, $ 14-6-19-1, $ 18-7-23-3. Length of hind femur, 
<J 9-7-11-8, $12-2-15-3. Maximum width of hind femur, $2-4-2-9, $3-0-3-6. Maximum 
width of head, <J 2-7-3-2, $ 3-4-4-3. 

Ratios (twenty specimens of each sex measured) : 

P/C FL/FW E/P E/F 



f 






^ 


( 






\ 


f 






i 


Min. 


Ave. 


Max. 


Min. 


Ave. 


Max. 


Min. 


Ave. 


Max. 


I-IO 


1-19 


I 


.32 


3-96 


4-10 


4 


21 


4 


42 


4-78 


5-17 


i -08 


I-I5 


I 


22 


4'3 


4-26 


4 


48 


4 


60 


4-80 


5-10 



Min. Ave. Max. 
i -43 i -54 1-62 
1-41 1-51 1-59 

Discussion. This subspecies appears to be a culmination of a trend in the austro- 
oriental subspecies tamulus as displayed by Australian populations of the latter. 
The structure of the fastigial foveolae and frontal ridge in dubius is similar to that 
of Australian tamulus but the black colouration on the hind tibia in dubius serves for 
separation. To separate a subspecies on the basis of merely a colour difference 
may be regarded with suspicion, but as the separation is backed by the geographical 
distribution of the two forms in that tamulus is not found east of the Australasian 
region and dubius is restricted to the Polynesian region (Text-fig. 84) it seems reason- 
able to regard the two forms as separate taxa of subspecific status. 

Distribution (Text-fig. 84). Two hundred and fifty-six specimens of this sub- 
species were examined and from these the following distributional notes were made : 



352 



D. HOLLIS 



NORFOLK Is., January to February; NEW CALEDONIA, May, July to September, 
November through to January; LOYALTY Is., no dates; NEW HEBRIDES, through- 
out the year; FIJI, April to May, October; FRIENDLY Is., February to March; 
SAVAGE Is., June; SAMOA, February to March, May, August to September. 



Bp 







IOO 



G 



Apd 




-Ap 



X Rm 




Ap 



FIGS. 97-102. A. thalassinus dubius (Willemse), genitalia. 97, $ spermatheca of specimen 
from Samoa; 98, same, of specimen from New Hebrides; 99, <J epiphallus; 100, g phallic 
complex, dorsal view, epiphallus and ectophallic membrane removed; 101, same, lateral 
view; 102, $ endophallus, lateral view. 

REFERENCES 

BARLOW, E. 1900. Notes on insect pests from the entomological section, Indian Museum. 

Indian Mus. Notes 5 : 14-34. 
BEI-BIENKO, G. Ya. 1966. (The orthopteran insects from Komodo and adjacent islands in 

Indonesia). Zool. Zh. 45 : 1779-1795, 7 figs. (In Russian with English summary). 
BEI-BIENKO, G. Ya. & MISHCHENKO, L. L. 1951. [Locusts and grasshoppers of the USSR 

and adjacent countries] 2 : 383-646, figs. 817-1318. Leningrad. (In Russian). 
BOLIVAR, I. 1895. Mission scientifique de M. Ch. Alluaud aux iles Seychelles (Mars-Avril- 

Mai 1892). 6 e mem. Orthopteres. Annls Soc. ent. Fr. 1895 : 369-385. 



REVISION OF GENUS AIOLOPUS 353 

BOLIVAR, I. 1898. Catalogo sin6ptico de los Ortopteros de la Fauna Iberica. Anais sci. nat. 

5 : 54-134- 

1902. Les Orthopteres de St. Joseph's College a Trichinopoly (Sud de 1'Inde). Annls 
Soc. ent. Fr. 1902 : 580-635, i pi. 

1912. The Percy Sladen Trust expedition to the Indian Ocean in 1905 under the leader- 
ship of Mr. J. Stanley Gardiner, M.A. Vol. 4. no. 16. Orthoptera: Acrydiidae, Phasgon- 
uridae, Gryllidae. Trans. Linn. Soc. Lond., Zool. 15 : 26-292, 2 pis. 

1918. Contribuci6n al conocimiento de la fauna indica. Orthoptera (Locustidae vel 

Acridiidae). Revta R. Acad. Cienc. exact, fis. nat. Madr. 16 : 278-289, 374-412. 

BRANCSIK, K. 1895. Orthoptera quaedam nova africana et australica. Jh. naturw. Ver. 
Trencsiner Kom. 17-18 : 243-262, pis. 7-9. 

BRULLE, A. 1840. Orthoptera. In WEBB, P. B. & BERTHELOT, S. Histoire naturelle des 
lies Canaries 2 (2) : 74-78, pi. 5. Paris. 

BRUNNER VON WATTENWYL, C. 1882. Prodromus der europdischen Orthopteren. 446 pp., 
12 pis. Leipzig. 

BURMEISTER, H. 1838. Handbuch der Entomologie 2 : 397-756. Berlin. 

BUTLER, A. G. 1876. Preliminary notice of new species of Orthoptera and Hemiptera col- 
lected in the island of Rodriguez by the naturalists accompanying the Transit of Venus 
Expedition. Ann. Mag. nat. Hist. (4) 17 : 409-412. 

CHAPMAN, R. F. 1961. The egg pods of some tropical African grasshoppers (Orthopt. Acri- 

doidea). /. ent. Soc. sth. Afr. 24 : 259-284, 26 figs. 

1962. The ecology and distribution of grasshoppers in Ghana. Proc. zool. Soc. Lond. 
139 : 1-66, 55 figs., 20 tabs. 

CHAPMAN, R. F. & ROBERTSON, I. A. D. 1958. The egg pods of some tropical African grass- 
hoppers. /. ent. Soc. sth. Afr. 21 : 85-112, 17 figs. 

CHESLER, J. 1938. Observations on the biology of some South African Acrididae. Trans. 
R. ent. Soc. Lond. 87 : 313-351, 27 figs. 

CHOPARD, L. 1943. Faune de V Empire Francais. i. Orthopteroides de 1'Afrique du Nord. 
45 PP- 658 figs. Paris. 

COSTA, O. G. 1836. Fauna del regno di Napoli. Ortotteri. 52 -f 16 pp., 12 pis. Naples. 

COTES, E. C. 1893. A conspectus of the insects which affect crops in India. Indian Mus. 
Notes 2 : 145-176. 

1894. Miscellaneous notes. From the entomological section. Indian Mus. Notes 3 : 39- 

84. 

DAVEY, J. T. DESCAMPS, M. & DEMANGE, R. 1959. Notes on the Acridinae of the French 
Sudan with special reference to the Central Niger Delta, pts. i and 2. Bull. Inst. fr. 
Afr. noire. 21 : 60-112, 565-600, 5 tabs., i map. 

DESCAMPS, M. & WINTREBERT, D. 1966 (1967). Pyrgomorphidac et Acrididae de Madagascar. 
Observations biologiques et diagnoses (Orth. Acridoidea). Eos, Madr. 42 : 41-263, 51 figs. 

DIRSH, V. M. 1953. Morphometrical studies on the phases of the desert locust (Schistocerca 

gregaria Forskal). Anti-Locust Bull. 16 : 34 pp., 31 figs. 

1963. The Acridoidea (Orthoptera) of Madagascar. II. Acrididae, Acridinae. Bull. 
Br. Mus. nat. Hist. (Ent.) 13 : 245-286, 21 figs. 

1965. The African genera of Acridoidea. 579 pp., 452 figs. Cambridge. 
FABRICIUS, J. C. 1781. Species Insectorum 1 : 340-371. Hamburgi et Kilonii. 

1798. Entomologia systematica. Suppl. 572 pp. Hafniae. 

FIEBER, F. X. 1853. Synopsis der europaischen Orthoptera. Lotos 3 : 90-104. 

FINOT, A. 1895. Faune de 1'Algerie et de la Tunisie. Insectes Orthopteres. Annls Soc. ent. 

Fr. 1895 : 401-552, 4 pis. 

FISCHER, L. H. 1853. Orthoptera Europaea. 454 pp., 18 pis. Leipzig. 
FISCHER DE WALDHEIM, G. 1846. Entomographie de la Russie. 4. Orthopteres de la 

Russie. Nouv. mem. soc. Imp. nat. Moscou 8 : 443 pp., 37 pis. 
FLETCHER, T. B. 1914. Some South Indian insects and other animals of importance, considered 

especially from an economic point of view. 565 pp. Madras. 



354 D. HOLLIS 

GHILIANI, V. 1869. Razza o specie nuova di Acridite. Bull. Soc. ent. ital. 1 : 177-180. 
GREATHEAD, D. J. 1963. A review of the insect enemies of Acridoidea (Orthoptera) . Trans. 
R. ent. Soc. Lond. 114 : 437-517, 66 figs. 

1966. Notes on Blaesoxipha spp. (Dipt. Calliphoridae) parasitising Acridoidea in eastern 
Africa. Tech. Bull. Commonw. Inst. Biol. Control 7: 91-100. 

HAFEZ, M. & IBRAHIM, M. M. 1962. On the ecology and biology of the grasshopper Aiolopus 
thalassinus (F.) in Egypt. (Orthoptera: Acrididae). Bull. Soc. ent. Egypte 46 : 189-214, 
3 figs., 5 tabs. 

I962. On the biology of the immature forms of the grasshopper Aiolopus thalassinus 
(F.) in Egypt. (Orthoptera: Acrididae). Bull. Soc. ent. Egypte 46 : 271-282, i fig., 4 tabs. 

1963. Field and laboratory studies on the behaviour of Aiolopus thalassinus (F.) towards 
humidity. (Orthoptera: Acrididae). Bull. Soc. ent. Egypte 47 : 75-96. 

1963^. The temperature reactions of Aiolopus thalassinus (F.). (Orthoptera: Acrididae). 
Bull. Soc. ent. Egypte 47 : 105-116. 

HOLLIS, D. 1967. New combinations affecting the genus Aiolopus (Orthoptera: Acridoidea) 
and a description of a related new genus and species from Australia. /. nat. Hist. 1 : 157- 
162, figs. 1-8. 

IBRAHIM, M. M. 1963. Further investigations into the humidity behaviour of Aiolopus thalas- 
sinus (F.). (Orthoptera: Acrididae). Bull. Soc. ent. gypte 47 : 97-103. 

INNES BEY, W. 1929. Revision des Orthopteres de 1'Egypte. Mem. Soc. ent. Egypte 

3 : 1-176, 7 pis. 

JOYCE, R. J. V. 1952. The ecology of grasshoppers in East Central Sudan. Anti-Locust 
Bull. II : 97PP-. 34 n g s - 

1954. I n SUDAN GOVERNMENT. Ministry of Agriculture. Annual Report of the Re- 
search Division 1951/2. Entomological section, pp. 117, 134-135. 

KEAY, R. W. J. 1959. Vegetation map of Africa south of the Tropic of Cancer. Oxford 
University Press. 

KEY, K. H. L. 1967. The type material of Aiolopus tamulus (F.) (Orthoptera: Acrididae). 
J. Aust. ent. Soc. 6 : 69-70. 

KHALIFA, A. 1956. The egg-pods of some Egyptian grasshoppers and the preference of females 
for soils of different moisture contents. Bull. Soc. ent. Egypte 40 : 175-186, 6 figs., 6 tabs. 

1956^. The incidence of grasshoppers during winter months and the influence of irrigating 
fallow land on grasshopper populations. Bull. Soc. ent. Egypte 40 : 217-229, 2 figs., 2 tabs. 

1957. The development of eggs of some Egyptian species of grasshoppers, with a special 
reference to the incidence of diapause in the eggs of Eyprepocnemis plorans Charp. (Ortho- 
ptera: Acrididae). Bull. Soc. ent. Egypte 41 : 299-330, 7 figs., 8 tabs. 

KIRBY, W. F. 1888. On the insects (exclusive of Coleoptera and Lepidoptera) of Christmas 
Island. In LISTER, J. J. On the natural history of Christmas Island, in the Indian Ocean. 
Proc. zool. Soc. Lond. 1888 : 546-554. 

1910. A synonymic catalogue of the Orthoptera. 3. Orthoptera Saltatoria. II. Locus- 
tidae vel Acridiidae. London. 

1914. The Fauna of British India, including Ceylon and Burma. Orthoptera (Acridiidae). 

276 pp., 140 figs. London. 
KRAUSS, H. 1890. Erklarung der Orthopteren-Tafeln J. C. Savigny's in der " Description 

de 1'Egypte ", Verh. zool.-bot. Ges. Wien 40 : 227-272. 
LATREILLE, P. A. 1804. Histoire naturelle . . . des Crustacds et des Insectes. 12. 424 pp., 

4 pis. Paris. 

NOLTE, D. J. 1939. A comparative study of seven species of Transvaal Acrididae, with 
special reference to the chromosome complex. /. ent. Soc. sth. Afr. 2 : 196-260, 144 figs., 
14 tabs. 

PHIPPS, J. 1959. Studies on East African Acridoidea (Orthoptera), with special reference to 
egg-production, habitats and seasonal cycles. Trans. R. ent. Soc. Lond. Ill : 27-56, i fig. 

1966. The habitat and seasonal distribution of some East African grasshoppers (Orthop- 
tera: Acridoidea). Proc. R. ent. Soc. Lond. (A) 41 : 25-36. 



REVISION OF GENUS AIOLOPUS 355 

REHN, J. A. G. 1902. Notes on some generic names employed by Serville in the Revue 
Methodique, and Fieber in the Synopsis der europdischen Orthopteren. Can. Ent. 34 : 

316-317- 
ROBERTSON, I. A. D. & CHAPMAN, R. F. 1962. Notes on the biology of some grasshoppers 

from the Rukwa Valley, S.W. Tanganyika. Eos, Madr. 38 : 51-114, 2 figs., 31 tabs. 
ROFFEY, J. 1965. Report of the government of Thailand on locust and grasshopper control. 

Food and Agriculture Organization of the United Nations. Report no. 2109. Rome. 
RUNGS, C. 1938. Observation d'unvol en essaimd'Aiolopus thalassinus (Fabr.). (Orthopteres- 

Acrididae). Proc. R. ent. Soc. Land. (A) 13 : 135-136. 

SAVIGNY, J. C. 1809-1813. Description de I'Egypte. Orthoptera. 7 pis. (no text). Paris. 
SJOSTEDT, Y. 1909. Wissenschaftliche Ergebnisse der Schwedischen Zool. Exped. nach devn 

Kilimandjaro, dem Meru und den umgebenden Massaisteppen Deutsch-Ostafrikas, 19051906. 

17. Orthoptera. 7. Acridoidea: 149-199. i pi., 2 figs. Stockholm. 
1923. Zoological results of the Swedish expedition to Central Africa 1921. Insecta. i. 

Acridoidea. Ark. zool. 15 (6) : 39 pp., i pi. 

1929. Orthoptera. i Acridoidea. In Voyage au Congo de S.A.R. le Prince Leopold de 
Belgique (1925). Revue Zool. Bot. afr. 17 (i) : 21-37, 10 figs. 

SNODGRASS, R. E. 1935. Principles of Insect Morphology. McGraw-Hill, New York. 
STAL, C. 1873 Recensio Orthopterorum. Revue critique des Orthopteres decrits par Linne, de 

Geer et Thunberg. 1. 154 pp., Stockholm. 
STOREY, G. 1919. The identification of the Orthoptera figured by Savigny, and other notes 

on Egyptian Orthoptera. Bull. Soc. ent. Egypte 1918 : 49-68. 
THUNBERG, C. P. 1815. Hemipterorum maxillosorum genera illustrata plurimisque novis 

speciebus ditata ac descripta. Mem. Acad. Sci. st.-Petersb. 5 : 211-301, pi. 3, 7 figs. 
UVAROV, B. P. 1922. Records and descriptions of Orthoptera from S.W. Asia. /. Bombay 

nat. hist. Soc. 28 : 351-370, 2 figs. 

ig22a. On some new or little known South African grasshoppers of the subfamily Acrid- 
inae. Ann. Mag. nat. Hist., (9) 9 : 539-551, i fig. 

1924. Some new and interesting Orthoptera in the collection of the Ministry of Agricul- 
ture, Cairo. Bull. Minist. Agric. Egypt tech. scient. Serv. 41 : 1-41, 3 pis. 

1926. New or less known Acrididae from Central Asia. Eos. Madr. 2 : 321-359, i fig. 

1928. The Orthoptera (excluding Blattidae) of Rodriguez Island. Ann. Mag. nat. Hist. 
(10) 2 : 362-364. 

1942. New and less known southern palaearctic Orthoptera. Trans. Am. ent. Soc. 
67 : 303-361, 5 pis. 

1953- Grasshoppers (Orthoptera, Acrididae) of Angola and Northern Rhodesia, collected 
by Dr. Malcolm Burr in 1927-1928. Publcoes cult. Co. Diam. Angola 21 : 1-217, 2 95 n 8 s - 

WILLEMSE, C. 1923. Locustidae (Acrididae a. a.) et Phasgonuridae (Locustidae a. a.) de 

la Nouvelle-Caledonie et des lies Loyalty. Nova Caledonia, Zoologie 3 : 99-112, 10 figs. 

ZIMIN, L. S. 1938. Les Pontes des Acridiens. Morphologic, classification et ecologie. Opred. 
Faune SSSR 23 : 84 pp., 10 pis., 6 figs. 




A LIST OF SUPPLEMENTS 
TO THE ENTOMOLOGICAL SERIES 

OF THE BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 



1. MASNER, L. The types of Proctotrupoidea (Hymenoptera) in the British 
Museum (Natural History) and in the Hope Department of Entomology, Oxford. 
Pp. 143. February, 1965. 5. 

2. NIXON, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera: 
Braconidae). Pp. 284; 348 Text-figures. August, 1965. 6. 

3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177; 
18 plates, 270 Text-figures. August, 1965. 4 45. 

4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172; 500 Text-figures. October, 

1965- 35s. 

5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156; 
475 Text-figures. November, 1965. 2 155. 

6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae & Drosophilidae. 
Pp. 129; 328 Text-figures. May, 1966. 3. 

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

335. 

8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the 
world species of Cleora (Lepidoptera : Geometridae). Pp. 119; 14 plates, 146 
Text-figures, 9 maps. February, 1967. 3 los. 

9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera : Rhopalocera). Pp. 509. August, 1967. 8 los. 

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rhopa- 
locera). Pp. 322; Coloured frontispiece, 348 text figures. August, 1967. 8. 

11. MOUND, L. A. A review of R. S. BagnalTs Thysanoptera Collection. Pp. 184; 
82 Text-figures. May, 1968. 4. 



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



REVISION OF THE GENUS 

CATASARCUS SCHONHERR 

(COLEOPTERA : CURCULIONIDAE) 




R. T. THOMPSON 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 22 No. 8 

LONDON : 1968 



* 

REVISION OF THE GENUS CATASARCUS 



SCHONHERR 



(COLEOPTERA : CURCULIONIDAE) 



BY 



R. T. THOMPSON 

*" v- 

(British Museum (Natural History)) 



Pp. 357-454; i Plate, 64 Text-figures, 4 Maps. 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 22 No. 8 

LONDON: 1968 



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 became 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

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

This paper is Vol. 22, No. 8 of the Entomological 
series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



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



Trustees of the British Museum (Natural History) 1968 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 31 December, 1968 Price 2 



REVISION OF THE GENUS CATASARCUS 

SCHONHERR 
(COLEOPTERA : CURCULIONIDAE) 

By R. T. THOMPSON 

CONTENTS 

Page 

INTRODUCTION ............. 359 

HISTORY OF THE GENUS ........... 359 

DISTRIBUTION ............. 360 

BIOLOGY .............. 361 

SOURCES OF MATERIAL ............ 362 

ACKNOWLEDGEMENTS ............ 362 

Catasarcus SCHONHERR ........... 363 

Characters ............. 363 

Spurious characters used by previous authors ....... 364 

Relationships ............ 365 

Notes on types ............ 366 

Terminology. ............ 366 

Identification ............ 367 

KEY TO SPECIES ............. 368 

DESCRIPTIONS (NOTE ON THE QUADRISPINATE SPECIES, p. 423) .... 423 

CHECK-LIST OF SPECIES (INCLUDING SYNONYMS, ETC.) ...... 452 

REFERENCES ............. 452 

INDEX .............. 453 

SYNOPSIS 

Catasarcus Schonherr is an exclusively Australian genus. As here treated, it comprises 41 
species and one subspecies. The latter, and 19 of the species are described as new. All, with 
one exception, are included in a key. Twenty-one species are figured (19 for the first time) and 
there are photographs of three others. All the types of previously described species (except one) 
have been located and checked and are formally cited. 

Most species have small ranges and these are indicated, where known, by maps and lists of 
localities. Attention is drawn to the importance of accurate locality data in the study of 
flightless insects. 

INTRODUCTION 

THE present work originated in an attempt to complete a revision of Catasarcus begun 
in 1955 by the late Sir Guy Marshall. His revision was far from completion and, as 
much additional material containing several undescribed species became available, 
it was decided to start the revision afresh. 

Four of the species described here bear the manuscript names proposed for them by 
Marshall and this fact is stated in each case. 

HISTORY OF THE GENUS 

Catasarcus was erected by Schonherr (1840) for four species described in this work 
by Fahraeus and two described previously by Boisduval (1835) m the palaearctic 

ENTOM. 22. 8 l8 



360 R. T. THOMPSON 

genus Cneorhinus Schonherr, 1826. Further species were described by Boheman in 
Schonherr, 1845 (i); Germar, 1848 (i); Pascoe, 1870 (34); Blackburn, 1894 (2), 
1896 (i); Lea, 19090; (4) and 1917 (i var.). Numerous unpublished names occur on 
specimens examined by Chevrolat (in his own collection), Pascoe (in his own collec- 
tion) and Marshall (in various museums). None of these names is quoted here. 
Discounting two names published in synonymy by Fahraeus in Schonherr (1840), the 
total of available names is 50. 

By 1931, when the genus appeared in Coleopterorum Catalogue (114: 7), five of these 
names had been placed in synonymy. Of these synonymies, one is here maintained, 
two are altered and two revoked. In the present work, a further 24 names are sunk 
(including the variety, which is also promoted to specific rank) and one is transferred 
to another genus. With the addition of 20 new names, there is a net decrease of three 
in the 1931 total. 

The genus has been redescribed by Labram and Imhoff (1848, No. 27), Lacordaire 
(1863 : 249) and Pascoe, whose revision of the genus appeared in 1870. The species 
were catalogued by Gemminger and Harold (1871 : 2311) and Masters (1872 : 217 
and 1886 : 59 2 )- Lea (1897 : 590-600) published a series of observations, including 
a critical review of Pascoe's treatment of the quadrispinate species. He also included 
species of Catasarcus in various lists (1908 : 129, 19096 : 216, 1911 : 180). The 
genus was mentioned by Froggatt (1907 : 182) and Tillyard (1926 : 242). The name 
occurs in Heyne and Taschenberg (1908 : 226), in a key to the genera of Leptopiini 
by Heller (1923 : 148) and in a list of the Coleoptera described by Blackburn (Lea 
1912 : xvii). As stated above, the genus appeared in Coleopterorum Catalogue in 



DISTRIBUTION 

(Map i) 

The genus Catasarcus is confined to the mainland and certain off-shore islands of 
Australia. It occurs on Kangaroo I. and on several small islands near Perth. No 
species is known to occur in Tasmania, New South Wales or Queensland. There are 
two species and one subspecies in eastern Australia ; the remainder occur in Western 
Australia, mostly south of Geraldton (29 S) but extending along the coast to North 
West Cape (22 S). Most of the species have restricted ranges but one (C. armatus] 
extends from Western Australia into both South Australia and the Northern Territory. 

A glance at the maps showing the ranges of individual species (pp. 403,450) and 
especially that showing the distribution of the two subspecies of C. transversalis (p. 380) 
will show how important precise locality data is in the study of flightless insects. 
The present revision has been severely hampered by the false and inaccurate data 
on many specimens. Lea's comment (19090: : 156) on specimens in European 
museums applies equally to Australian material. Some early specimens have no 
labels at all and the value of data such as 'Interior' is obvious. Apart from patently 
false localities, some recent labels are astonishingly vague, e.g. 'Everard Rgs., S.A. 
to Warburton Rgs., W. A.' (a distance of about 370 miles). In another case 
' Central Austr./26.vi.i927/G. Home ' proper data would have provided a second 
locality for one of the remotest and most interesting species. 



REVISION OF GENUS CATASARCUS 



361 




MAP i. Catasarcus. Range. The Nullarbor Plain should be included (see p. 446). 



BIOLOGY 

Very little has been recorded on the biology of the adults and the immature stages 
are virtually unknown, in spite of the fact that several species are common in well 
populated areas. 

Records of adult host-plants are few and do not indicate which parts of the plant, 
if any, are attacked. A total of five families and nine genera, mostly of xerophytic 
trees and shrubs, is at present implicated, namely: Leptospermum, Melaleuca, 
^Eucalyptus (Myrtaceae) ; Jacksonia, Acacia (Leguminosae) ; Banksia, Hakea 
(Proteaceae) ; Casuarina (Casuarinaceae) and Xanthorrhoea (Xanthorrhoeaceae) . 
This range of families and the fact that three species of Catasarcus have been recorded 
from more than one host-plant family indicate a low degree of host-specificity. 

The habit in many Curculionidae of dropping to the ground when disturbed, seems 
to be especially well marked in Catasarcus, a fact which should be borne in mind by 
collectors and which has been reported for C. transversal-is by Tepper (1887 : 30) and 



362 



R. T. THOMPSON 



by the late F. E. Wilson (personal communication). Wilson states: ' . . . it will 
frequently drop to the ground from its resting place when one gets within a yard or two 
of it. When lying doggo, it is difficult to see on the greyish sand of its habitat '. 

The eggs and young larvae of C. asphaltinus have been seen by Mrs. P. Sundstrom 
and a report of her observations is given under this species (p. 411). 

SOURCES OF MATERIAL 

The following sources of material are indicated in the text by the symbols which 
precede them. 

A Australian Museum, Sydney. 

BM(NH) British Museum (Natural History), London. 

California California Academy of Sciences, San Francisco. 

CSIRO C.S.I.R.O. Division of Entomology, Canberra. 

Dresden Staatliches Museum fur Tierkunde. 

FEW F. E. Wilson, East Malvern, Victoria. (Mr. Wilson's collection is now 

in the National Museum of Victoria.) 

FHUB Dr. F. H. Uther Baker, Applecross, Perth. 

Frey Museum G. Frey, Tiitzing. 

Macleay Macleay Museum, University of Sydney. 

Manchester Manchester Museum, University of Manchester. 

Munich Zoologische Sammlung des Bayerischen Staates. 

New York American Museum of Natural History. 

NSWAg New South Wales Department of Agriculture, Rydalmere. 

Oxford Hope Department of Zoology (Entomology), University Museum, 

Oxford. 

S South Australian Museum, Adelaide. 

Stockholm Naturhistoriska Riksmuseum. 

UW University of Western Australia, Perth. (This material is now in the 

Western Australian Museum.) 

V National Museum of Victoria, Melbourne. 

W Western Australian Museum, Perth. 

Washington United States National Museum. 



ACKNOWLEDGEMENTS 

Among those who have sent me specimens for study I would like to thank 
especially Messrs C. Koch (W), A. Neboiss (V), G. F. Gross (S) and above all Dr. F. H. 
Uther Baker of Perth, who has not only made available to me his private collection 
but has sought diligently for further material, both in collections and in the field and 
has taken endless trouble to trace obscure localities. He has, to his great credit, 
collected no fewer than 23 species in the field, 13 of which are new. 

My thanks are similarly due to Dr. P. B. Carne (CSIRO), A. M. Douglas (Perth), 
Dr. J. W. Evans (A), Dr. H. Freude (Munich), E. Gowing-Scopes (Halstead, England), 
my friend Dr. E. Haaf (formerly of the Frey Museum), Miss E. Hahn and Mrs. J. 



REVISION OF GENUS CATASARCUS 363 

Anderson (Macleay), Dr. R. Hertel (Dresden), the late Dr. W. D. Hincks (Manchester), 
Prof. Dr. J. O. Hiising (Martin Luther Universitat, Halle), the late Dr. E. 
Kjellander (Stockholm), Hugh B. Leech (California), Mrs. P. Sundstrom (Perth), 
E. Taylor (Oxford), Mrs. P. Vaurie (New York), J.-M. Vrydagh (Institut royal des 
Sciences naturelles de Belgique, Brussels), Miss R. E. Warner (Washington) and 
the late F. E. Wilson (East Malvern, Viet.). 

I owe a special debt of gratitude to Dr. G. Kuschel of Nelson, N.Z. (late of Santiago, 
Chile) both for locating and studying the types of Boisduval on my behalf and for 
giving me help and encouragement at a critical stage in this study. 

I am indebted to my senior colleague, Mr. J. Balfour-Browne for checking the 
typescript and making valuable suggestions. I also wish to thank Dr. E. B. Britton 
(now at CSIRO, Canberra) who collected some valuable material during a visit to 
Australia and my present colleagues R. D. Pope, Miss C. M. F. von Hayek, Dr. R. 
Madge and Dr. N. A. Aslam for their interest and advice. 

I am very grateful to M. J. D. Brendell and Miss E. R. Tozer for testing the key. 

The 21 figures of whole insects were executed by Mrs. C. A. O'Brien and Text-figs. 
2-5 by Mr. Arthur Smith. 

CATASARCUS Schonherr 
Catasarcus Schonherr, 1840 : 812. 

Type-species: Catasarcus bilineatus Fahraeus in Schonherr, 1840: 813, by original 
selection. 

Characters 

Apterous Leptopiinae having a dorsal transverse furrow or impressed line near base 
of rostrum ; f rons with median longitudinal sulcus leading off from transverse rostral 
furrow and two or four longitudinal carinae; post-humeral region of elytra with a 
spine, tubercle or bulge (absent in individual specimens of some species). 

Members of this genus also have the following characters in common: 

Mandibles multisetose and usually partly squamose. Rostrum x 1-1-5 as l n g as broad; 
dorsal area flat with margins raised and median carina present, continuous with epistome. 
Antennal funicle with seven segments; scrobes lateral, linear, deep and oblique. Prothorax 
distinctly broader than long, with traces of two dorsal transverse striae, about equidistant from 
each other and the pronotal margins. Scutellum very small or obsolete. Metepisternal suture 
indicated at extreme anterior end only; posterior end dorsally with finger-like process projecting 
posteriad over metathoracic spiracle. Elytra without any swelling at shoulders but often with 
humeral tubercles; costal margin strongly sinuous. Intercoxal process of ventrite i truncate 
and almost twice as broad as a hind coxa ; suture between ventrites i and 2 arcuate, deep at sides, 
becoming fine or obscure in mid-line; lengths of ventrites 3 and 4 subequal and together === 2. 
Legs with femora more or less clavif orm ; tibiae with teeth along ventral (inner) margin, weakly 
mucronate at apex and with corbels enclosed ; tarsi (especially segment 3) all larger in proportion 
to overall size in male than in female; claws free, but only weakly diverging, simple. Ovi- 
positor strongly sclerotized and without styli. 

The transverse furrow is represented by an impressed line in only one species, 
C. memnonius, in which it is also deflected posteriad in the middle and at the sides 
by processes from the rostrum; in all other species it is deeper and straight. 



364 R. T. THOMPSON 

About a third of the species have prominent dorsal spines on the elytra. It is these 
species, with one exception, which have a large spine in the post-humeral region of the 
elytra. The exception is C. albipectus, which has long dorsal spines but only a 
very small tubercle or bulge behind the shoulder. The non-spiny species have either 
a large bulge often surmounted by a blunt tubercle, or a small, sometimes sharp 
tubercle, or a small and often indistinct bulge. In general, this post-humeral 
prominence lies in interstria 10 (between striae 9 and 10) in the non-spiny species 
but in interstria 9 in those with dorsal spines. It is interesting to note that in the 
latter group there is a gap in stria 9 below the spine and the strial punctures on each 
side are drawn upwards towards the base of the spine, as if the spine had emerged in 
interstria 10 and then forced its way through stria 9 into interstria 9. In the non- 
spiny species the tubercle is often closer to stria 9 than 10 and when large causes 
displacement or even interruption of stria 9. In the non-spiny C. memnonius, 
which is believed to have an affinity with the spiny species, the tubercle emerges 
about in the line of stria 9 as it also does in the spiny C, albipectus in which (as stated 
above) it is very small and so does not interrupt the stria. 

The dorsal spines also cause gross distortion of the strial punctures on the disc, so 
that it is not always easy to decide in which interstriae the spines are situated. That 
the spines are not present as such in the pupa is suggested by a specimen of C. 
spinipennis in which, though subsequently fully hardened, the elytra failed to expand 
completely at eclosion; the anterior spines are absent and the distal half of each 
posterior spine is invaginated into the base. It is difficult to see how this condition 
could arise if the spines had been fully formed before eclosion. 

Apart from C. sericeus, of which only five females have been seen, both sexes are 
known to occur in all species. The sex ratio usually approaches parity but males 
predominate in C. albipectus, C. griseus and possibly other species. In addition 
to having larger tarsi, the elytra in the male are usually slightly narrower and more 
evenly rounded, both above and at the sides, than in the female, though in species 
with globose elytra, they are similar or even slightly broader in the male. Ventrite 5 
is usually less strongly convex in the male and weakly truncate and setose at the apex, 
whereas in the female it is entire. In some spiny species, the post-humeral tubercles 
are large in the female but very small or absent in the male. 

Spurious Characters used by previous Authors 

The following characters were used solely, or principally, to distinguish the species 
indicated: 

Bleaching: C. stygmatipennis (Boisd.); C. ceratus Pasc.; C. pollinosus Pasc. ; 

C. albuminosus Pasc. 

This is a curious condition which I have not encountered in any other genus. It is 
found in specimens which have suffered severe abrasion ; not only are the scales and 
clothing setae missing but also the large setae of the legs and the corbellar fringes ; 
in extreme cases, even the tarsal pads may be lost. The cuticle, whether black or red, 
becomes a uniform pale greyish colour with a waxy sheen. This effect is most 
pronounced on the more exposed surfaces; thus the underside, the frontal sulci and 



REVISION OF GENUS CATASARCUS 365 

the strial punctures are often unaffected. The cuticle is not only bleached but 
softened and minor surface irregularities are smoothed out. If the soft layer is 
scraped off, firmer cuticle of the original colour is found beneath. 

This condition is common in some species (e.g. C. impressipennis] but rare or 
unknown in others. Lea (1897 : 593-594) gives a good account of it and suggests 
that it may be caused by the weevils rubbing against comparatively hard leaves 
(Banksia, Acacia, etc.) since he observed that specimens from very soft-leaved 
plants were unaffected. Precisely how the effect is produced is unknown. 

Extraneous Granules : C. capita Pasc. ; C. furfuraceus Pasc. 

Described by Pascoe as ' sand-like exudation ', these granules are resinous in 
nature ; on gentle heating they melt and evaporate, leaving a tarry residue which, on 
stronger heating, leaves a whitish ash. They are insoluble in water, alcohol and 
benzene. 

In the type specimens in question, they are very numerous and hence rather 
conspicuous but I have seen them in smaller numbers on specimens of several other 
species ; they are often attached to the long setae on the tibiae. They are presumably 
derived from the plant on which the weevil lives. 

Cuticle Thickness : C. mollis Lea; C, durus Lea. 

Lea first mentions this character in 1897 (: 599) and comments that it ' has been 
entirely overlooked by Mr. Pascoe '. As indicated in the discussion on p. 420 below, 
I regard this character as having little value. In this instance, Lea merely separated 
teneral and fully hardened specimens of the same species. 

In addition to the above examples, a number of Pascoe's species are based on 
abnormal or defective specimens ; these are discussed under the species concerned. 

Relationships 

Catasarcus has no close relatives. The only form known to me which exhibits any 
of the major distinguishing characters of Catasarcus is an undescribed species and 
genus from the northern part of Western Australia. 

At the same time, it has so far proved impossible to subdivide the genus. Several 
more or less distinct species-groups are apparent but the affinities of many species 
remain in doubt. Pascoe, in his revision, divided the genus according to the number 
of dorsal spines on the elytra. The present study shows that this character can be 
misleading. For example, C. carbo and C. lepidus are shown to be closely related, 
yet the former has four spines and the latter eight. Even as an artificial character 
the spines are not always reliable. A more useful character is afforded by the apex 
of the ovipositor which is laterally compressed or cylindrical in most spineless and 
quadrispinate species but dorso-ventrally flattened and blade-like in some multi- 
spinate species. Serious anomalies occur, however. Thus it is blade-like in the 
quadrispinate C. marginispinis but cylindrical in the apparently closely related 
C. concretus which has six dorsal spines; also, it is cylindrical in C. murex which has 
eight spines and flattened in C. memnonius which has none. Another important 



366 R. T. THOMPSON 

character, used successfully by Marshall in other groups, is the number of setae on the 
mentum. The ten species from C. latheticus to C. longicornis inclusive (except 
C. carinaticeps) have 6 or more setae, while most of the other species have only 4. But 
C. carinaticeps has 4 or 6 and both C. azureipes and C. inaequalis have 6 or more, 
though they are not closely related to the ten species, or to each other. It is worth 
noting, however, that the two species with only 2 setae on the mentum (C. ustulatus 
and C. murex) are each highly distinctive in other respects. 

Notes on Types 

I have cited as holotypes those specimens which I have satisfied myself to be so, 
regardless of the manner in which they have been labelled or previously documented. 
I do not consider it necessary or advisable to designate uncited holotypes as lectotypes 
and I hope that those who do will accept my citations in lieu thereof. It has, however, 
proved necessary to designate one lectotype (for C. humerosus Pasc.) and one neotype 
(for C. hopei Fahrs.). 

The recognition of paratypes has sometimes proved difficult. In the case of 
Hope's material described by Fahraeus, there are additional specimens of some of the 
species concerned both in Stockholm and in the Hope collection at Oxford. Although 
I have listed these specimens, I do not regard them as paratypes since the phrase 
' Dom. Hope. Mus. Schh. ' indicates that the described specimens were retained in 
Stockholm and in any case the descriptions appear to have been based on the 
holotypes alone. 

Pascoe frequently based his descriptions on a series of specimens (indicated by a 
range of lengths) but did not label the paratypes as such, so that they cannot now be 
distinguished from specimens acquired subsequently. Indeed, the presence of a 
determination label on a Pascoe specimen, other than the holotype, is a fair indication 
that the specimen is not a paratype. A further complication exists in Pascoe's case, 
owing to the fact that he examined A. Fry's material of Catasarcus about the time his 
paper was published and many Fry specimens named by Pascoe were labelled 
' TYPE ' by Fry. Most of these specimens were obtained by Fry direct from du 
Boulay and were almost certainly seen by Pascoe after his paper was completed. 
Two of them, however, were obtained by Fry from Pascoe at this time and I have 
accepted these as paratypes (see p. 429). 

Terminology 

The terms herein used are mainly those of Marshall, except that segment replaces 
joint and interstria is preferred to interval. The areas between successive strial 
punctures are gaps ; these and the interstriae taken together are the elytral interspaces. 
The term stria, in relation to the elytra, is purely locational and does not imply the 
presence of an impressed line; where such lines are present, the striae are described as 
impressed. 

To avoid confusion, the fifth tarsal segment is called the claw segment and abdominal 
sternites 3-7 are referred to as ventrites 1-5. In the males of some non-spiny species 
there is a fairly well defined depression on ventrite I behind each hind coxa; these 
are the post-coxal cavities. 



REVISION OF GENUS CATASARCUS 367 

In some species, the apex of the rostrum is abruptly expanded ventrally; this 
expansion is referred to as the chin, from its appearance in profile view (Text-fig. 8) . 
On either side of the epistome, usually near the anterior margin, are its flanking 
setae. Any setae on the corbellar plate are called adventitious setae on account of 
their irregular appearance. 

Scales are sparse when clearly separated from each other, dense when subcontiguous, 
tessellate when pressed together (without overlapping) so as to obscure completely 
the underlying cuticle and imbricate when they overlap strongly; they may be in 
close contact with the cuticle (appressed) or raised up from it (loose). The scales 
collectively, together with the clothing setae, constitute the vestiture. 

The aedeagus is terete when subcircular in transverse section; its subterminal 
orifice is the phallotreme. The halves into which the apex of the ovipositor is divided 
are its valves. 

Carinae, etc. are arched when convex in profile view. The frons is convex when 
the middle is higher than the sides but level longitudinally; when the middle is also 
arched, the frons is said to be dome-shaped. The term triangular denotes an 
equilateral triangle. The prefix micro- is applied to states or structures only clearly 
visible under a magnification of c. X 125. 

Colours are described as they appear under a binocular microscope, using high 
voltage illumination and a bull's-eye condenser. Scales described as bronzy are 
brown with a metallic sheen. 

Identification 

In addition to differences between the sexes (p. 364), many species exhibit great 
variation in size, sculpture and vestiture. Their appearance may also be altered 
by the presence of powdery exudate or as a result of bleaching (p. 364). At the same 
time, the differences between species may be slight or subtle so that correct identifi- 
cation by comparison alone is often impossible. 

The various body proportions given in the key and descriptions were measured 
under a microscope using an eyepiece scale. The proportions of the pro thorax and 
elytra are given with the length first, corrected to 10 in each case. This makes all 
the ratios comparable and avoids the use of figures less than unity for the elytral 
width. As the true elytral length is difficult to measure accurately, the line from 
the scutellum to the apex is used instead (see AC in Text-fig, i). The proportions 
of antennal funicle segments 1-3 are given with 3 (the shortest) corrected to I in each 
case. These proportions vary considerably in each species, and mean figures, not 
ranges, are therefore given. The stated number of specimens from which the means 
are derived should be taken into account when comparing them with fresh data. 
The length of the rostrum was taken from the level of the anterior margins of the 
eyes to the longer of the two genae or the margin of the epistome if this were longer. 

A very useful datum for identifying certain quadrispinate species is what I have 
called the anterior spine index (see Text-fig, i). The measurements must be accurately 
made, using an eyepiece scale and taking care to incline the specimen so as to bring 
the points being measured into the same focal plane, thus ensuring true (maximum) 
readings. As the base of the spine is ill-defined, measurement AB is made to the 



368 R. T. THOMPSON 

centre of the nearest strial puncture; if there is no puncture near the line of 
measurement, the position is obtained by estimation. A table of these indices is 
given on p. 423. 

The aedeagus is sometimes important in identification and should be examined 
wherever possible. Both the aedeagus and the ovipositor can, with care, be drawn 
out of a fully relaxed specimen with watchmaker's forceps without damaging the 
specimen, especially if they are allowed to remain in situ. 




X 100 = anteri r 
AC spine index 



FIG. i. Catasarcus spinipennis Fahraeus $. Elytra, showing how anterior 
spine index is obtained. 

As stated earlier, it is difficult to divide the species of Catasarcus into groups 
morphologically. Nevertheless, in the key to species I have attempted to group the 
non-spiny species according to the condition of the frontal carinae. This may make 
the couplets concerned difficult to interpret but it avoids bringing out all the species 
seriatim, which would make the key very tedious to use. To allow for variation in 
the carinae and other characters, several species have been brought out in more than 
one place. Some indication of the range of each species, where known, is given in the 
key ; this will often serve to confirm determinations. Estimates of frequency would, for 
the most part, be meaningless but I have marked four species as ' Common ' ; this is 
partly to prevent a wrong impression being gained from the fact that one of the four 
is new. 

KEY TO THE SPECIES OF CATASARCUS 
(Except C. albuminosus) 

1 Elytra with dorsal spines .......... 36 

Elytra without dorsal spines ......... 2 

2 (i) Head with 4 distinct longitudinal carinae on frons ..... 4 

Head without distinct frontal carinae ....... 3 

3 (2) Frons convex, with median longitudinal cleft anteriorly; median rostral carina 

very strongly raised and projecting posteriad over the very short but deep 
transverse basal furrow; dorsum densely squamose. W. A., near Busselton 

ustulatus sp. n. (p. 388) 

Frons flat, with shallow median sulcus; transverse basal furrow similarly 
shallow: median rostral carina not strongly raised or projecting posteriad; 
dorsum bare. S . A ., east of Lake Eyre . . memnonius Pascoe (p. 422) 



REVISION OF GENUS CATASARCUS 

4 (2) Frontal carinae all well developed, equidistant from each other and straight (or 
almost so) (Text-fig. 2) ; lateral carinae parallel or weakly converging anteriorly ; 
(admedians often broader than laterals but then well separated by deep 
median sulcus) ; length rarely < 8 mm. W. A ., central and southern parts, 
east of A Ibany ........... 

Frontal carinae either all poorly developed, or one pair differing markedly from 
the other in size, shape, or both (Text-figs. 3-5) (if not markedly different, 
then admedians closer to each other than to laterals) .... 



369 






FIGS. 2-5 . Catasarcus spp. Head in dorsal view to show frontal carinae. 2, C. obesus sp. n. 
3, C. hopei Fahraeus. 4, C. impressipennis (Boisduval). 5, C. opimus Pascoe. 



370 R. T. THOMPSON 

5 (4) Hind femora distinctly curving upwards near base (Text-fig. 10) ; pronotum with 

pair of small pale admedian scale-patches near anterior margin; scales on 
elytra mostly yellowish, on legs and underside of head bluish white with 
coppery reflection; rostrum not, or scarcely, longer than broad, epistome 
large and flat and making an acute angle with mentum. Esperance area 

varus sp. n. (p. 386) 

Hind femora almost straight in profile view (Text-fig, n); pronotum without 
small pale spots; elytra variegated, or with mostly whitish scales; scales on 
underside of head whitish (rarely bright blue), without coppery reflection . 6 

6 (5) Elytra elongate-ovate (10 : < 7); fore femora strongly swollen; legs with 

small vivid metallic blue or green scales, knees black ; underside of head with 
narrow tract of pure white (or vivid blue) scales below eye. Inland, north-east 
of Albany ........ azureipes sp. n. (p. 385) 

Elytra subglobose (10 : > 7) ; fore femora normal; femora and underside of head 

with dense whitish scales throughout ....... 7 

7 (6) Elytra with areas of yellow, white and grey scales forming an irregular pattern ; 

strial punctures very regular, mostly pupillate; prothorax less strongly 
transverse (10 : < 18). Wialki-Nulla Nulla area . aspergetus sp. n. (p. 383) 
Elytra without yellow scales, strial punctures less regular, not pupillate; 
prothorax more strongly transverse (10 : > 18). Southern Cross-Ravens- 
thorpe area ......... obesus sp. n. (p. 382) 

8 (4) Admedian frontal carinae as long as lateral carinae, much closer to each other 

than to laterals and raised above them (if shorter, or not distinctly higher 
than laterals, then knees black and frons without any median elevation); 
humeral tubercle usually sharp, post-humeral tubercle smaller than humeral 
tubercle, or obsolete; interstriae smooth. Esperance-Hopetoun area, inland 
to Widgiemooltha ...... carinaticeps Lea (p. 400) 

Characters not so combined ......... 9 

9 (8) Admedian frontal carinae as long as, and distinctly broader than, the lateral 

carinae (Text-figs. 3, 4). W. A. . . . . . . . 10 

Admedian carinae otherwise ......... 20 

10 (9) Hind femora distinctly curving upwards near base (Text-fig. 10); epistome 

clearly defined, triangular, flat or weakly convex. Esperance area 

varus sp. n. (p. 386) 
Hind femora almost straight in profile view (Text-fig, n); epistome usually 

elongate or with disc depressed . . . . . . . . 1 1 

11 (10) Epistome elongate, merging gradually with median rostral carina in both dorsal 

and profile view; admedian frontal carinae and median rostral carina usually 
strongly raised, bare and shiny ; antennae with f unicle segments 4-7 scarcely 
longer than broad. Albany and extreme south-west . hopei Fahraeus (p. 397) 

Characters not so combined ......... 12 

12 (11) Prothorax twice as broad as long ($); elytra in $ globose (Text-fig. 34), almost 

smooth ; striae not, or weakly, impressed, punctures small (diameter often < 
width of interstriae) ; scales fairly evenly distributed. Tammin 

sericeus Blackburn (p. 396) 

Prothorax less strongly transverse (10 : < 18); elytra less strongly inflated, 
often granulose-rugose ; punctures larger (diameter at least = width of 
interstriae) or striae strongly impressed, or both; scales concentrated in 
depressions ............ 13 

13 (12) Legs with tarsi black above (difficult to see when scales are dense). Mainly 

north of Bunbury ........... 14 

Legs unicolorous, red (sometimes very dark). Mainly south of Bunbury . . 17 



REVISION OF GENUS CATASARCUS 371 

14 (13) Antennae with funicle segments 5-7 not, or scarcely, longer than broad, club 

ovate; mentum with 4 setae; head with no broad scales below eye. Around 
Perth ......... cygnensis sp. n. (p. 412) 

Antennae with funicle segments 5-7 distinctly longer than broad, club fusiform ; 

mentum with at least 6 setae . . . . . . . . 15 

15 (14) Epistome merging with median rostral carina in profile view (ignoring any 

constriction between the two), convex; admedian frontal carinae as long as 
laterals and close together. Hill River area 

pallidiventris sp. n. (p. 406) 
Epistome weakly but abruptly declivous in profile view, flat . . . 16 

16 (15) Epistome with numerous (c. 8-10) flanking setae on each side; scales below 

eye usually narrow and separate; length < 19 mm. Around Perth. Common 

asphaltinus sp. n. (p. 407) 

Epistome with few (c. 4) flanking setae ; scales below eye broader, contiguous or 
imbricate; length probably not > 14 mm. Range uncertain 

longicornis Pascoe (p. 411) 





10 




11 



FIGS. 6-9. Catasarcus spp. Outline of head in profile view (antennae omitted) . 6, C. hopei 
Fahraeus. 7, C. asphaltinus sp. n. 8, C. bilineatus Fahraeus. 9, C. aerosus sp. n. 

FIGS. lo-n. Catasarcus spp. Outline of left femur in profile view. 10, C. varus sp. n. 

n, C. obesus sp. n. 



372 R. T. THOMPSON 

17 (13) Head with patch of scales below eye, admedian frontal carinae straight; dorsal 

area of rostrum with scales concentrated at posterior end. Around Cape 
Naturaliste ........ coruscus sp. n. (p. 414) 

Head without, or with very small filiform scales below eye (not forming a 

patch) ; all frontal carinae usually more or less curved . . . . 18 

18 (17) Elytra with interstria 7 about twice as wide as 6 (Plate i, Figs. 3, 4). Range 

unknown ........ inaequalis sp. n. (p. 421) 

Elytra with interstriae 6 and 7 subequal . . . . . . . 19 

19 (18) Prothorax with narrow tract of imbricate scales at sides, extending from anterior 

constriction to hind angle; elytra less elongate (<$, 10 : 6-5-6-7), interspaces 
convex and smooth. Manjimup ..... laevior sp. n. (p. 417) 
Prothorax without imbricate scales at sides between anterior constriction and 
hind angle; elytra more elongate (<J, 10 : 5-8-6-5), interspaces flat and uneven 
or convex and weakly rugose. Extreme south-west, mainly around Albany. 
Common ...... impressipennis (Boisduval) (p. 417) 

20 (9) Post-humeral tubercle of elytra < segment 3 of antennal funicle, or represented 

by a smooth bulge, or absent ........ 29 

Post-humeral tubercle > segment 3 of funicle ; rostrum with distinct chin (Text- 
figs. 8, 9) . ... 21 

21 (20) Elytra with well marked sutural stripe of pale scales; rostrum with median 

carina raised, arched, bare and shiny; lateral frontal carinae narrow, parallel 
or weakly converging anteriorly; admedian carinae about half as long as 
laterals (often ill -defined). W. A., west coastal region .... 22 
Elytra rarely with any trace of a sutural stripe, lateral frontal carinae distinctly 

converging anteriorly . . . . . . . . . .23 

22 (21) Frons flat, epistome with numerous large flanking setae and smaller setae on disc ; 

form more elongate (elytra in $ 10: 6-4-6-8) ; vestiture of pronotum uniform. 
Wide area around Perth, inland to Beverley. Common 

bilineatus Fahraeus (p-393) 

Frons convex, epistome with 3-6 large adherent flanking setae and a similar 
number of much smaller, separate, setae mesal of these but scarcely any on 
disc; form less elongate (elytra in both sexes c. 10 : 7); pronotum with two 
broad, ill-defined pale stripes or admedian patches. Moore River 

latheticus sp. n. (p. 393) 

23 (21) Post-humeral tubercle of elytra distinct but small (< segment 2 of antennal 

funicle) and sharp. In or near Perth ....... 24 

Post-humeral tubercle large ( > segment 2 of funicle) or represented by a large 

obtuse bulge; length seldom > n mm. . . . . . . .25 

24 (23) Epistome with one tuft of adherent flanking setae (appearing as a single seta) 

on either side; dorsal area of rostrum narrow at base, usually progressively 
widening apically; median rostral carina depressed in middle (Text-fig. 9); 
lateral frontal carinae strongly converging anteriorly; legs slender, tibial 
teeth small. Bejoording-Lancelin area .... aerosus sp. n. (p. 390) 
Epistome with several separate flanking setae; dorsal area of rostrum usually 
distinctly narrower at apex than at base (sometimes lyre-shaped) ; median 
rostral carina usually level (sometimes depressed in middle); lateral frontal 
carinae less strongly converging anteriorly; legs stout, tibial teeth large. 
Perth Gingin area ...... griseus Pascoe (p. 391) 

25 (23) Prothorax very strongly transverse (10 : > 24), sides almost straight, very 

strongly converging anteriorly (Text-fig. 31); elytra globose; antennae with 
segment i of funicle scarcely longer than 2. W. A., around Esperance 

bakeri sp. n. (p. 381) 
Prothorax less strongly transverse; funicle with segment i distinctly longer 

than 2 26 



REVISION OF GENUS CATASARCUS 373 

26 (25) Epistome narrow, flat, weakly arched and continuous with median rostral carina 

in profile view, with several recumbent squamiform or normal setae 
posteriorly. Eastern Australia . transversalis anatolicus ssp. n. (p. 380) 

Epistome broad, triangular, disc more or less depressed, bare ... 27 

27 (26) Scales immediately in front of eye smaller and narrower than those below eye 28 

Scales immediately in front of eye similar or identical to those below eye ; elytra 

with post-humeral tubercle very large and blunt or represented by a broad 
bulge, completely covered with scales; disc of elytra with weak transverse 
folds ; scales mostly pearly but with heavy deposit of yellow-brown powdery 
exudate. W. A., around Albany, inland to Borden . rugulosus Boheman (p. 389) 

28 (27) Form broader (elytra 10 : 7-2-8-1) ; elytra usually with strong continuous sinuous 

transverse folds; median rostral carina usually strongly raised and projecting 
posteriad over transverse furrow; epistome with i or 2 small flanking setae; 
setae on femora small dark and inconspicuous. Eastern Australia 

transversalis Germar (p. 377) 

Form less broad (elytra 10 : 6-8-7-3) ; elytra without, or with weak transverse 

folds; median rostral carina not strongly raised or projecting posteriad; 
epistome with several separate flanking setae ; setae on femora large, pale and 
conspicuous. W. A., Perth-Gingin area .... griseus Pascoe (p. 391) 

29 (20) Mainly west of A Ibany .... 30 

Mainly east of A Ibany ...... 5 

30 (29) Knees black, fore femora strongly swollen ... 31 

Knees red (sometimes infuscate or with black spots but never entirely black), 

fore femora swollen or not ...... .32 

31 (30) Corbel tapering to a point at dorsal end; elytra finely granulate throughout, 

strial punctures ill-defined or irregular; ventrite 5 with transverse carina 
(sometimes obsolete in ?) ; frons densely squamose (scales completely covering 
admedian carinae) and usually with a narrow wedge-shaped median carina 
(Text-fig. 5); pronotum coarsely granulo-rugose ; femora sparsely squamose. 
Between Perth and Geraldton opimus Pascoe (p. 404) 

Corbel not tapering to a point dorsally ; elytra without granules ; strial punctures 

well defined, regular, picked out with white scales; all interspaces equally 
convex, forming a very regular reticulum (mesh pattern) ; frons less densely 
squamose ; femora (at least in part) with vivid metallic blue or green scales. 
Inland, north-east of Albany . . . azureipes sp. n. (p. 385) 

32 (30) Median rostral carina depressed in middle (not at junction with epistome) (Text- 

fig. 9); post-humeral tubercle always present, sharp; lateral frontal carinae 
strongly converging anteriorly; length < 13 mm. Bejoording-Lancelin 
area aerosus sp. n. (p. 390) 

Median rostral carina level or arched; post-humeral tubercle often blunt or 

absent; lateral frontal carinae not, or weakly, converging anteriorly . 33 

33 (32) Epistome with numerous (c. 8-10) flanking setae on either side, poorly defined, 

flat and weakly but abruptly declivous; interstriae of elytra flat or weakly 
convex, rugose; apex of elytra in male finely rugose, appearing shrivelled. 
Wide area around Perth. Common . . . asphaltinus sp. n. (p. 407) 

Epistome with 2-4 flanking setae ........ 34 

34 (33) Frons flat, admedian frontal carinae completely covered with loose imbricate 

scales; centre of frons usually with very narrow cariniform elevation which 
widens posteriorly and merges with vertex. Toodyay-Merredin area 

frontalis sp. n. (p. 402) 

Frons weakly convex, admedian frontal carinae not completely covered with 

scales ; centre of frons without any cariniform elevation . . . . 35 
ENTOM. 22, 8 I9 



374 



R. T. THOMPSON 



35 (34) Epistome convex, merging with median rostral carina in profile view (ignoring 

any constriction between the two) ; scales in front of eye c. x 2-5-5 as l n g as 
broad, < half as wide, on average, as scales below eye ; size and form as in C. 
asphaltinus. Hill River ..... pallidiventris sp. n. (p. 406) 

Epistome flat, weakly but abruptly declivous ; scales in front of eye x i -5-3 as 
long as broad, on average > half as wide as scales below eye ; length probably 
not > 14 mm. Range uncertain .... longicornis Pascoe (p. 411) 

36 (i) Elytra with 6 or more dorsal spines or large tubercles (Text-figs. 18-22) . . 47 

Elytra with fewer than 6 dorsal spines (Text-figs. 13-17, 23). W. A. . . 37 

37 (36) Pronotum dark red, head and elytra black. West coast, north of Geraldton . 38 

Pronotum black or brown, concolorous with head and elytra 39 




16 



FIGS. 1 2-1 8. Catasarcus spp. Outline of left elytron in profile view. 12, C. memnonius 
Pascoe cj. 13, C. intermedius Pascoe ?. 14, C. albipectus sp. n. $. 15, Idem $. 16, 
C. echidna Pascoe <$. 17, C. spinipennis Fahraeus ?. 18, C. concretus Pascoe <$. 



REVISION OF GENUS CATASARCUS 375 

38 (37) Form more elongate (prothorax 10 : 14-7-16-7, elytra 10 : 6-8-7-6); <J with 

posterior dorsal spines very long and cylindrical (Text-fig. 14), post-humeral 
spine represented by a small sharp tubercle, or obsolete ; disc of elytra without 
discrete scale-patches. Murchison River Reserve . albipectus sp. n. (p. 425) 
Form less elongate (prothorax 10 : 16-5-17-3, elytra 10 : 7-4-8) ; both sexes with 
dorsal elytral spines tapering throughout their length; post-humeral spine 
normal; disc of elytra with discrete, but irregular, scale-patches. Between 
Geraldton and Murchison River ..... bicolor sp. n. (p. 427) 

39 (37) Corbels squamose; eyes round ( x 1-2 as long as broad), very strongly convex, 

completely encircled with white scales; disc of pronotum very strongly 
rugose, with impressed median line. Geraldton Shark Bay area 

car bo Pascoe (p. 447) 

Corbels without scales; eyes more elongate, less strongly convex, not, or 
incompletely, encircled with white scales; disc of pronotum less strongly 
rugose than sides ........... 40 

40 (39) Dorsal elytral spines black . . . . . . . . .41 

Dorsal elytral spines red or dark red . . . . . . . .42 

41 (40) Head with lateral frontal carinae absent, admedian carinae very small, tuber- 

culiform ; usually each dorsal elytral spine ascended by a tract of pale scales. 
Bejoording-Lake Grace area .... marginispinis Pascoe (p. 436) 
Head with lateral frontal carinae distinct; dorsal elytral spines bare or with uni- 
form small dark scales. Around Geraldton . . echidna Pascoe (p. 428) 

42 (40) Elytra with interstriae 2 and 3 each with a small sharp shiny forwardly- 

projecting tubercle at base (projecting beyond base), vestiture brown with a 
pale flash at sides; prothorax transverse but barrel-shaped. PGeraldton area 

cicatricosus Pascoe (p. 440) 
Elytra without, or with different tubercles at base ..... 43 

43 (42) Corbels densely setose ( > 20 setae on plate) ; transverse rostral furrow shallow; 

post-ocular lobes of prothorax rather sharply angulate; antennal club short 
and stout (2 : i); dorsal elytral spines small (anterior pair sometimes 
obsolete) and set further back (Text-fig. 13). Between Carnarvon and North 
West Cape ....... intermedius Pascoe (p. 424) 

Corbels with < 20 adventitious setae; transverse rostral furrow deep; post- 
ocular lobes not, or weakly, angulate; antennal club more elongate; dorsal 
spines, on average, larger and set further forward on elytra ... 44 

44 (43) Prothorax subcylindrical (10 : < 16); scales forming a black and white 

pattern (Text-fig. 58) (dorsal elytral spines red). PGeraldton area 

albisparsus Pascoe (p. 436) 
Prothorax much broader at base than at apex (10 : > 16) ; scales, if white, not 

forming a pattern .......... 45 

45 (44) Elytra with anterior dorsal spines evidently nearer base in $ than in $; base of 

interstria 3 with large smooth shiny callus or large granule; head with lateral 
frontal carinae usually extending over top of eye. Around Geraldton 

echidna Pascoe (p. 428) 

Elytra with dorsal spines not evidently nearer base in <J than in $, base of 
interstria 3 simple or, if with large granule, then several other similar but 
smaller granules present in humeral region; head with lateral frontal carinae 
stopping short before eye (and often obscured by scales) .... 46 

46 (45) Aedeagus with transverse wrinkles below phallotreme ; majority of scales on tarsi 

brown (with metallic reflections) ; elytra typically with patches of pearly or 
coppery scales on a dark background and usually without granules. Perth- 
Hill River area nephelodes sp. n. (p. 431) 

Aedeagus without any wrinkles below phallotreme; majority of scales on 
tarsi whitish or pearly; elytra typically with large whitish scales throughout 



376 R. T. THOMPSON 

and often granulose. Perth area, including off-shore islands, north to Yanchep, 
south to Pemberton. Common . . . spinipennis Fahraeus (p. 443) 

47 (36) Head with lateral frontal carinae distinct ; each elytron with i large spine at top 

of declivity and 3 smaller spines (very small in $) on disc (Text-fig. 21); 
humeral tubercle large in $, absent in g. Kalgoorlie, W. A., to Fowler's Bay, 
S. A., inland to Ayer's Rock, N. T. . . . arrnatus Blackburn (p. 443) 

Head with lateral frontal carinae indistinct or absent. W. A. . . . 48 

48 (47) Prothorax dark red; eyes x 1-5 as long as broad, very weakly convex; each 

elytron with 4 large dorsal spines, including one in interstria 4 between middle 
and base (Text-figs. 19, 20). Bridgetown-Lake Grace area rnurex sp. n. (p. 441) 
Prothorax concolorous with head and elytra; eyes < x 1-3 as long as broad, 

moderately to very strongly convex ; elytral spines otherwise ... 49 

49 (48) Corbels squamose; pronotum more strongly rugose on disc than at sides, anterior 

border with bifid median swelling; elytra without a small sharp tubercle at 

base of interstria 3 .......... 50 




19 





21 





FIGS. 19-23. Catasarcus spp. Outline of left elytron in profile view. 19, C. murexsp. n. <. 
20, Idem ?. 21, C. armatus Blackburn $. 22, C. lepidus Pascoe <$. 23, C. carbo Pascoe <$. 



REVISION OF GENUS CATASARCUS 377 

Corbels without scales ; pronotum less strongly rugose or granulate on disc than 
at sides, anterior border not swollen; elytra with a small sharp tubercle at 
base of interstria 3, projecting anteriad beyond base. Hopetoun 

concretus Pascoe (p. 435) 

50 (49) Eyes very strongly convex, broadly encircled with white scales ; each elytron with 
2 large dorsal spines posteriorly (Text-fig. 23). Geraldton-Shark Bay area 

carbo Pascoe (p. 447) 

Eyes less strongly convex, not, or very narrowly, encircled with white scales; 
each elytron with 3 large dorsal spines posteriorly and one near base, in 
interstria 5 (Text-fig. 22). Range uncertain . . lepidus Pascoe (p. 446) 

Catasarcus transversalis Germar sp. rev. 
(Text-figs 24, 30, Map 2) 

Catasarcus transversalis Germar, 1848 : 212. 

Catasarcus transversalis Germar; Taschenberg, 1869 : 31. 

Catasarcus transversalis Germar; Lacordaire, 1863 : 250 (note). 

Catasarcus transversalis Germar; Pascoe, 1870 : 16, 25, 26. 

Catasarcus transversalis Germar; Tepper, 1887 : 30. 

Catasarcus stigmatipennis (Boisduval); Lea, 1918 : 265 [Erroneous synonymy]. 

Length 7-4-11-4 mm. Body black, legs and antennae dark red to black. Scales dense but 
easily abraded, mostly whitish, usually with pink of green reflection (rarely coppery) ; setae 
brown throughout; powdery exudate scanty or absent. Head with frons weakly to distinctly 
convex; frontal carinae very variable, laterals short, straight, subparallel to strongly converging 
anteriorly (rarely almost obsolete) ; admedian carinae straight or weakly curved, parallel with 
laterals or more strongly converging anteriorly, about equidistant from each other and from 
laterals, occasionally irregularly subdivided or with accessory carinulae; frons in mid-line even 
or with smooth elongate tectiform or rounded elevation; scales dorsally fairly dense and 
recumbent, mainly white on frons and olive-green on vertex (dense behind eyes) but white scales 
sometimes restricted to two ill-defined admedian tracts or a pair of dense patches just behind 
level of hind margins of eyes; scales below eyes pearly white, imbricate, becoming less dense 
towards mid-line. Rostrum x 1-1-1-2 as long as broad, distinctly widening apically; epistome 
broad with shallow discal depression accentuated by very low transverse subapical elevation, 
surface pitted and strongly microreticulate anteriorly, flanking setae as in C. rugulosus but 
one or two smaller setae near them and two more in median excision; median carina sharp, 
narrow in front becoming broader and tectiform behind and there moderately to very strongly 
raised (and often arched) and projecting strongly over the very deep transverse furrow with an 
oblique carina supporting the projection on either side; dorsal area rectangular to lyre-shaped, 
lateral sulci sometimes deep; sparsely to densely squamose throughout. Antennae with lengths 
of funicle segments 1-3 in ratio 2:1-5:1 (mean of seven), 4-6 slightly shorter than 3, subequal, 
7 =^= 3 and about x 1-3 as long as broad. Prothorax very strongly transverse (10 : i9'3- 2 3'9), 
broadest in basal half, sides rounded, strongly converging anteriorly; post-ocular lobes fairly 
well developed; upper surface smooth to obscurely granulate, sides distinctly to strongly granu- 
late; transverse striae strongly impressed, usually complete but often irregular; scales below 
and at sides dense, often partly imbricate; scales above less dense (sometimes very sparse) but 
usually with small pale spot on posterior stria at either side and pair of admedian patches, also 
on posterior stria as in C. rugulosus (but less well defined). Scutellum smooth, punctured, with 
variable number of elongate and filiform scales. Elytra shortly ovate-acuminate, globose in some 
females (10 : 7-2-8-1); humeral tubercle basal, well developed, forwardly directed and sharp 
in female, blunt or obsolete in male; interstriae 2, 3 and 5 often slightly raised at base; post- 
humeral tubercle large, conical or subparallel-sided, blunt, strongly reflexed ventrad and 
sometimes posteriad; striae impressed throughout; strial punctures large and deep near sides, 



378 



R. T. THOMPSON 



becoming smaller towards suture; interstria i sometimes flattened and depressed on disc, 
otherwise elytral interspaces strongly raised, forming a reticulate pattern or, more often, a series 
of high sinuous undulating transverse folds, mostly continuous across full width of each elytron ; 
scales very dense on interstriae 9 and 10 (except underside of post-humeral tubercle) and on 
interstria i when depressed, elsewhere less dense and fairly uniform but transverse folds often 
bare or with inconspicuous brown scales which may predominate on declivity. Legs rather 
slender; femora scarcely swollen; fore and middle tibiae very weakly curved, with moderate 
teeth; hind tibiae straight, with unequal teeth, corbels narrow with nought to many adventitious 
setae; femora usually with dense large round scales ventrally and at apex and sparse small 
elongate scales elsewhere (sometimes with large scales throughout) ; tibiae and tarsi with dense, 
mainly brown scales throughout. Underside densely squamose; ventrites i and 2 with small 
scattered granules, strongly raised and bead-like, especially in male; post-coxal cavities linear 
or obsolete; suture between ventrites i and 2 distinct throughout its length. Aedeagus (Text- 




A 




26 







A 

1 / 





29 



FIGS. 24-29. Catasarcus spp. Aedeagus in dorsal and lateral view. 24, C. transversalis 
Germar. 25, C. bakeri sp. n. 26, C. obesus sp. n. (Lake Carmody). 27, C. rugulosus 
Boheman. 28, C. griseus Pascoe. 29, C. varus sp. n. 



REVISION OF GENUS CATASARCUS 



379 



fig. 24) short, depressed, rather strongly curved, terete, smooth, except for numerous fine 
scattered granules on underside of apex; apical region short, tip broadly rounded, not usually 
deflexed. Ovipositor with valves somewhat depressed, together broader than high but each 
smoothly rounded and not at all explanate. 

Holotype <J, Adelaide, [1844-45 (H. H. Behr}}, in Zoologisches Institut, Martin 
Luther Universitat, Halle. Unique (see Taschenberg, 1869 : 31). 

Over 130 specimens seen. 

Localities: SOUTH AUSTRALIA: Kangaroo I. (Kelly's Hill Caves); Yorketown; 
Adelaide; Victor Harbour; Gawler; Nuriootpa; Barossa; Murray Bridge; Tintinara; 
Lucindale. VICTORIA: Yanac; Kiata; Gypsum. See map 2. 

A record for Newcastle, N.S.W. (Frey) is known to be false and a specimen from 
the Pascoe collection (BM (NH)) with ' Champion B. ' must also be wrongly labelled. 
A recent record for Alice Springs, N.T. (xii.1955, W. B. H[itchcock]) (V) must, if 
genuine, almost certainly be a transported specimen. 

Host-plants: Hakea rostrata (Tintinara, 6. i. 1887 (Tepper) (A)). Further host- 
plants are mentioned by Tepper (1887 : 30) with other interesting observations: 
' Catasarcus transversalis, Germar, is one of the commonest beetles in the scrub 
during spring and summer . . . The beetles feed on the leaves of various kinds of 
Leptospermum, Melaleuca (tea-trees) and shrubs. When alarmed they drop down 
at once '. The last observation is also reported by Wilson (p. 363, above). 




30 




FIGS. 30, 31. 30, C. transversalis Germar <$. 31, C. bakeri sp. n. $. 



380 



R. T. THOMPSON 



Lea's erroneous synonymy of this species with C. stygmatipennis is discussed on 
p. 420. In spite of some misgivings, Pascoe identified this species correctly. 

The nominate subspecies, described above, can usually be distinguished from all 
other forms by the strongly raised, sharp, beak-like median rostral carina and the 
very strong transverse folds on the elytra. 

Catasarcus transversalis anatolicus ssp. n. 

(Map 2) 

Length 7-3-10 mm. Distinguished from the nominate subspecies as follows. Head with 
admedian frontal carinae usually closer to each other than to laterals, median sulcus often 
reduced to a short cleft (frons then dome-shaped). Rostrum with epistome usually longer than 
broad, flat, arched, seldom with any subapical elevation, disc with a small number of squamiform 
setae posteriorly and not well denned from median carina which is evenly and less strongly 
raised. Prothorax slightly less transverse (10 : 18-5-20-6); transverse striae less strongly 
impressed (anterior often obsolete) ; scaling more uniform. Elytra with weaker transverse 
folds, often smooth or regularly granulate with strongly impressed striae. Legs almost always 
quite black; femora densely squamose throughout. Aedeagus as in C. t. transversalis except 
that both specimens examined have dorsal margins carinate almost to base. 

Holotype . VICTORIA: Warnambool, Teatree Creek, 9. x. 1964, in the National 
Museum of Victoria, Melbourne. 

Paratypes. 2 <$, Grampians (det. Lea, vii . 1904) (V); i $, same locality, 1934; 
2 <, Hall's Gap (all K. Guichard) (all Manchester) ; i , Kiata, 29 . xii . 1918 (F. E. 



VICTORIA 




C. f. transversa/is O 
C. t. anatolicus + 



MAP 2. Catasarcus t. transversalis Germar and C. t. anatolicus ssp. n. Distribution. 



REVISION OF GENUS CATASARCUS 381 

Wilson) (A); i <j>, ditto but 3i.xii.igi8 (FEW); i <$, ditto but 23. xi. 1952 (BM 
(NH)); i <J, Little Desert, 9 mis. S. of Kiata, x.1948 (A. Musgrave) (A); i $, 2 $, 
Little Desert, 17-25. x. 1952 (. M[atheson\) (2V, iBM(NH)); i ?, same locality, 
23. x. 1946 (4. J3[wms]); i $, Gypsum (C. Oke); i <$, Dimboula (all V); i g, i $, 
without data (V); i <$, Mount Gambier, 29.x. 1957 (W. M. M[oore]); i $, ditto but 
r.xii.i957 (both V); i <, i $, ' S. Australia ' (BM(NH), V). Total: 22 specimens. 

Localities : SOUTH AUSTRALIA : Mount Gambier. VICTORIA : Kiata ; Little Desert ; 
Dimboula; Gypsum; Grampians; Hall's Gap; Teatree Creek (near Warnambool). 
See Map 2. 

In the specimen from Dimboula and one of those from Kiata, the elytra have a high 
proportion of golden scales, especially at the base and along interstria one (the suture). 
In some other specimens, notably those from Mount Gambier, there is a heavy deposit 
of yellow powdery exudate. 

The most easterly representative of the genus. The ranges of the two subspecies 
appear to meet and possibly overlap in the region of the Little Desert. They were 
taken together at Kiata in 1918 and have both been taken at Gypsum (on separate 
occasions). The nominate subspecies was also taken in the Little Desert (without 
precise location) by W. van der Starre in 1964 (FHUB, BM(NH)). 

Catasarcus bakeri sp. n. 

(Text-figs. 25, 31, Map 4) 

Length 6-2-9-1 mm. Body black, legs and antennae dark red to black. Scales dense, mostly 
whitish or pearly; setae brown throughout; no powdery exudate observed. Head with frons 
as in C. rugulosus; scales large and dense throughout, imbricate below eye, mostly white or pearly 
but usually brown on vertex; eyes almost flat, suboblong, x 1-5 as long as broad. Rostrum 
X 1-1-1-3 as l n g as broad, distinctly widening apically; epistome smaller than in C. rugulosus, 
disc flat, more coarsely pitted and often without evident microsculpture ; two (apparently one) 
flanking setae; median carina narrow, often depressed in middle; dorsal area rectangular or 
broadest at junction with upper margin of scrobe; chin fairly well defined, sharp; rather densely 
squamose throughout. Antennae with lengths of funicle segments 1-3 in ratio 1-7 : 1-5 : i 
(mean of five), 3-7 subequal (3 and 7 usually longest). Prothorax very strongly transverse 
(10 : 24-3-26-3), broadest at base, sides almost straight, very strongly converging anteriorly; 
post-ocular lobes poorly developed; upper surface smooth, with a few scattered punctures but 
no granules; sides finely rugose with at most a few obscure granules; transverse striae as in 
C. rugulosus, sometimes strongly impressed, producing between them a well marked transverse 
fold ; scales very dense or imbricate below and at sides, less dense above and there concentrated 
in two very ill-defined longitudinal tracts with, in addition, a small dense spot on posterior 
margin very near sides (best seen without magnification). Scutellum punctate, microrugose, 
sometimes strongly raised, bare or squamose. Mesosternal process broad, apex truncate. 
Elytra globose (10 : 7-5-8-5), apex acuminate, shape similar in both sexes; humeral tubercle 
usually obsolete; post-humeral tubercle very large and broad-based as in C. rugulosus but with 
apex drawn out into an acute but blunt cone with axis deflexed ventrad; striae distinctly 
impressed on declivity only, strial punctures very small, sometimes obscured by scales; inter- 
striae broad, flat, sometimes almost smooth but usually forming a series of low irregular trans- 
verse folds as in C. rugulosus; scales uniformly very dense throughout, whitish but brown 
scales predominate on declivity and form irregular patches on disc or are concentrated on 
interstriae 2, 4 and 6, forming dark stripes (rarely elytra brown throughout, except at sides). 
Legs as in C. rugulosus but hind femora less strongly tapering towards base (viewed from above) 



382 R. T. THOMPSON 

and outer surface near base with large, sometimes confluent, punctures; corbels always with 
many adventitious setae; scales large, round and very dense throughout. Underside as in 
C. rugulosus but granules on ventrite i of male less well developed. Aedeagus (Text-fig. 25) 
very short, very strongly curved, depressed, terete, smooth; apex narrow, tip not deflexed. 
Ovipositor with valves strongly and closely compressed. 

Holotype <j>. WESTERN AUSTRALIA: Esperance, 23.^.1955 (F. H. Uther Baker), 
in the Western Australian Museum, Perth. 

Paratypes. 8 <?, 5 $, same data as holotype (10 FHUB, 2 BM(NH), i W); 3 , 
5 ?, ditto but 22. iv. 1955 (5 FHUB, 2 A, i BM(NH)); 2 <?, 3 ?, ditto but 25.^.1955 
(3 FHUB, 2 BM(NH)); i & i ?, Myrup, 26. iv. 1955 (F. H. Uther Baker) (FHUB); 
2 & i ?, Dalyup, 6. .1960 (F. H. Uther Baker) (2 FHUB, i BM(NH)); i $, i ?, 
Eradu, 21. x. 1914 (/. Clark) (S). Total: 34 specimens. 

Localities: Esperance; Myrup; Dalyup. The record for Eradu must be a mistake. 

The name of this species was proposed by Marshall (i. litt.) as a well deserved 
tribute to Dr. Uther Baker. The general shape and especially that of the prothorax 
will distinguish this species from any other. 

Catasarcus obesus sp. n. 

(Text-figs. 2, n, 26, Map 4) 

Length 7-8-12-6 mm. Body black, legs and antennae very dark red. Scales white (with 
pink and green reflections) and olive-brown, dense throughout body and on legs. Head with 
frons flat or weakly convex; lateral frontal carinae straight (rarely weakly curved), usually 
sharp but not, or weakly, raised, weakly converging anteriorly; admedian carinae as long as 
laterals or nearly so (rarely only half as long), straight or weakly curved, parallel with adjacent 
lateral carinae or with each other and all four carinae usually equidistant from each other 
(Text-fig. 2) ; median frontal sulcus broad, deep and rather short ; centre of frons without any 
median elevation, sides, including admedian carinae and lateral sulci, covered with dense round 
white scales which are continuous with imbricate ovate scales on vertex; latter usually olive- 
brown but often white in continuation of the frontal tracts; centre of frons with scattered or 
dense, mainly olive-brown scales; underside of head throughout with dense ovate scales, often 
becoming imbricate below (and behind) eyes. Rostrum x 1-2-1-4 (o*)> X 1-1-1-2 (?) as long as 
broad, distinctly widening apically; epistome well defined, disc flat or weakly concave, pitted, 
with two (apparently one) small flanking setae on each gena; median carina narrow, usually 
moderately and evenly raised but sometimes strongly raised and arched; dorsal surface densely 
squamose throughout, sides rounded basally, weakly converging apically; chin variable. 
Antennae with lengths of funicle segments 1-3 in ratio 1-9 : 1-5 : i (mean of nine), 4-6 slightly 
shorter than 3, subequal, 7 = 3 and c. x 1-7 as long as broad. Prothorax very strongly trans- 
verse (10 : 18-7-23-8), broadest near base, sides rounded, converging anteriorly; post-ocular 
lobes poorly to fairly well developed ; upper surface smooth, very finely punctured (rarely with 
ill-defined granules at sides) ; transverse striae variable, posterior straight, often extending 
across full width (ill-defined in mid-line), anterior shorter, usually recurved posteriad towards 
sides ; scales imbricate above coxae and along sides, often forming a stripe ; elsewhere less dense 
or mainly olive-brown, sometimes forming two broad white longitudinal tracts on pronotum. 
Scutellum smooth, finely punctured, squamose. Mesosternal process very broad (as broad 
as base of middle femur) and usually abruptly truncate at apex. Elytra globose-acuminate 
(10 : 7-7-7), differing little between the sexes but inflated at base in some females; humeral 
tubercle small (rarely obsolete), usually cariniform with obtuse apex displaced posteriad; 
post-humeral area with a broad bulge, sometimes surmounted by a sharp tubercle; striae 



REVISION OF GENUS CATASARCUS 383 

distinctly impressed on declivity only, elsewhere surface thrown into a series of more or less 
continuous transverse folds; scales dense throughout, imbricate and exclusively white on sides 
around post-humeral tubercle, folds and interstriae bare (? abraded) or with olive-brown scales 
which form an irregular variegated pattern. Legs stout but femora scarcely swollen (Text-fig, 
n); fore tibiae incurved towards apex and with rather large teeth; corbels narrow and usually 
with several adventitious setae; scales very dense throughout, round, white (often with greenish 
reflection) and olive-brown, uniformly mixed; setae slender, brown, inconspicuous. Underside 
very finely rugose; ventrites i and 2 with very small discrete granules; post-coxal cavities 
shallow or absent; densely and evenly squamose throughout, scales white but sometimes each 
ventrite with two olive-brown patches, near sides. Aedeagus (Text-fig. 26) strongly curved, 
smooth; apical half slender, depressed, weakly sulcate dorsally; apex narrow, tip somewhat 
swollen and not, or very weakly, deflexed. Ovipositor with valves strongly compressed. 

Holotype <J. WESTERN AUSTRALIA : Lake Varley, 20 . ix . 1954 (F. H. Uther Baker) , 
in the Western Australian Museum, Perth. 

Paratypes. 7 <$, 2 $, same data as holotype (7 FHUB, 2 BM(NH)); i <j>, Ravens- 
thorpe, 2.ix.i952; i <j>, Lake Carmody, 20. ix. 1954; i <, same locality, 23. ix. 1954 
(aU F. H. Uther Baker) (2 FHUB, i BM(NH)); i ?, Dedari, {.1939 (F. E. Wilson) 
(FEW); i & 4 ?, Southern Cross, viii.1959 (H. Demarz) (4 Frey, i BM(NH)); i <?, 
ditto but 5.ix.i962 (Frey); 10 <$, 6 $, Widgiemooltha, i.x.1962 (A. M. Douglas and 
W. D. Findlay] (12 W, 3 BM(NH), i V); i <J, Kuminin [? = South Kumminin] 
(E. F. du Boulay) (S). Total: 37 specimens. 

Localities : as listed above. 

Host-plants: Jacksonia sp. (series from Widgiemooltha). 

Catasarcus aspergetus sp. n. 

(Text-fig. 32, Map 4) 

<J. Length 12-3-13-7 mm. Body black, legs and antennae dark red. Scaling dense through- 
out, bluish white with black and golden-yellow patches. Head with frons flat or convex; 
lateral frontal carinae strongly raised, fairly sharp, straight or weakly curved, very weakly 
converging anteriorly; admedian carinae narrow, straight, parallel, as long as laterals or shorter; 
lateral sulci deep, filled with dense raised round or ovate yellow scales which extend (less densely) 
over admedian carinae and posteriorly to level of hind margins of eyes; scales behind eyes black, 
elsewhere on vertex and underside of head white, ovate and dense becoming imbricate below 
eyes. Rostrum x 1-1-1-2 ($) as long as broad, widening rather abruptly towards apex; epistome 
flat, triangular, pitted; median carina broad, smooth, slightly or distinctly raised near base 
and projecting over transverse furrow; sides of dorsal area parallel, strongly raised, resulting 
lateral sulci filled with white or yellow scales. Antennae with lengths of funicle segments 
1-3 in ratio 2 : 1-56 : i (mean of two), 4-6 subequal, 7 = 3 and x 1-4-1-6 as long as broad; club 
black, in strong contrast with whitish funicle. Prothorax transverse (10 : 17-7) broadest about 
middle and there angulate, sides posteriorly straight, subparallel or weakly converging, anteriorly 
weakly curved and distinctly converging; post-ocular lobes well developed and with (relatively) 
long vibrissae; pronotum uneven, finely punctured, obscurely granulate at sides; anterior 
transverse stria shallow or obsolete, posterior almost complete but ill-defined in mid-line; 
scales mainly white, very dense and imbricate at sides and above coxae; dorsal surface in 
anterior half with two admedian patches of dense yellow scales which also cover post-ocular lobes 
and surrounding area. Scutellum smooth, with fine punctures and scales. Elytra ovate- 
acuminate (10 : 7-2), strongly and evenly convex; humeral and post-humeral tubercles blunt 



384 



R. T. THOMPSON 



or obsolete; striae impressed only on declivity where punctures are very small; strial punctures 
elsewhere large; interstriae consequently narrow but not deformed; scales dense, whitish, 
yellow and black in patches, forming a complex pattern. Legs with femora squamose through- 
out, scales whitish, round, appressed, mostly contiguous on shaft, tessellate on knee, as also 
on tibia; tibial teeth small, sharp; corbels with several adventitious setae; setae on femora 
dark brown and conspicuous. Underside with dense round mostly imbricate whitish scales 
and long semi-recumbent pale setae which are rather conspicuous, especially on ventrite 5 ; 
post-coxal cavities small or obsolete, cluster of strongly raised granules behind them in paratype ; 
holotype with anterior half of ventrites i and 2 and sides of 3-5 yellow and with some black 
scales among the white elsewhere. Aedeagus as in C. obesus but distinctly tectiform basally, not 
sulcate above and more strongly widening around phallotreme. 

Holotype <J. WESTERN AUSTRALIA: Wialki, ix.i959 (F. H. Uther Baker), in the 
Western Australian Museum, Perth. 

Paratype $. Nulla Nulla, [i9]33~352 (W). 




32 



FIG. 32. Catasarcus aspergetus sp. n. <$ (holotype). 



REVISION OF GENUS CATASARCUS 385 

Localities : as listed above. Of the several localities bearing the name Nulla Nulla, 
that nearest to the type-locality is assumed to be the one at which the paratype was 
taken. 

It will be interesting to see, when further material is available, how the colour- 
pattern in this species varies. Even the present specimens differ ; in the paratype, the 
interstriae and gaps between successive punctures are on the same level and both 
covered with black scales, producing a strongly marked reticulate pattern, while the 
punctures are filled with either white or yellow scales. The latter are in a minority 
and apart from a few on the disc, occur only at the base and in parts of striae 7 and 8. 
Where yellow punctures are adjacent, the intervening scales are also yellow (not 
black) so that continuous yellow areas are formed. This does not, however, disrupt 
the reticulate pattern very much. In the holotype, on the other hand, raised 
transverse folds are present on the disc and black scales occur only on them, so that the 
interstriae are obscured and the scales of both white and yellow punctures are 
contiguous laterally. The yellow punctures are also contiguous longitudinally, as in 
the paratype, but are here more numerous. The general appearance, on the disc, is 
that of a bluish white background with somewhat irregular black transverse lines, 
upon which has been superimposed a number of irregular deep yellow blotches. On 
the declivity, where the trans-strial folds are absent, the interstriae are raised and 
black while the striae are uniformly coloured, yellow dorsally and white at the sides. 
In this species, as in C. azureipes, the centres of the larger strial punctures are bare 
and pupil-like. 

Catasarcus azureipes sp. n. 

(Map 4) 

Length 12-6-17-8 mm. Body black with white and blackish scales; femora red or dark red 
(knees black) with metallic blue or green scales ; tibiae dark red, tarsi black ; antennae dark red ; 
setae brown throughout; red-brown powdery exudate often present. Head with frons flat or 
weakly convex, lateral frontal carinae raised, sharp, weakly curved, subparallel; admedian 
carinae usually as long as laterals (sometimes much shorter), straight, parallel or converging 
anteriorly, separated by a deep, sometimes wide, median furrow; centre of frons posteriorly 
usually with fine longitudinal striations, or rarely with a smooth median elevation; lateral 
sulci densely or sparsely filled with round or ovate white scales which sometimes extend sparsely 
over admedian carinae; vertex with dense ovate olive-brown and metallic blue-green scales; 
underside with a narrow tract of pure white scales below eye. Rostrum x 1-2 (<$), x i-i ($) as 
long as broad, weakly widening apically ; epistome large, triangular, weakly convex, finely pitted, 
clearly defined from median carina which is weakly raised, level, sometimes strongly depressed 
near junction with epistome; sides of dorsal area parallel, strongly raised, the resulting sulci 
filled with sparse or fairly dense ovate-elongate semi-erect white scales; oblique basal sulci 
usually well developed; apex in profile view rounded ventrally. Antennae with lengths of 
funicle segments 1-3 in ratio 2-2 : 1-42 : i (mean of four), 4-7 subequal, slightly shorter than 
3, 7 broader than 6 and x 1-6 as long as broad. Prothorax transverse (10 : 16-7-18-6), broadest 
about middle, sides rather strongly rounded, distinctly constricted behind post-ocular lobes 
which are less well developed than in C. obesus but have longer vibrissae; dorsal surface evenly 
and finely rugose or obscurely granulate, more distinctly granulate at sides ; anterior transverse 
stria obsolete, posterior stria represented by a deeply impressed line on either side ; scales sparse, 
white and olive-brown (latter inconspicuous), white scales somewhat denser at sides and in 
anterior constriction ; underside with irregular patch of very large round imbricate white scales 



386 R. T. THOMPSON 

above coxa. Scutellum microrugose or smooth, punctate, usually with several elongate or 
filiform metallic blue or whitish scales. Elytra ovate-elongate (10 : 6-3-6-8), broadest about 
middle, somewhat flattened above, more steeply declivous posteriorly in female, apex more 
broadly rounded (in dorsal view) in male than in female ; surface smooth and even throughout, 
devoid of granules and with only the finest puncturation and microsculpture ; humeral tubercle 
obsolete in male, cariniform in female ; post-humeral tubercle absent (sometimes represented by 
a low bulge) ; striae strongly impressed on declivity but very weakly on disc, especially in male ; 
strial punctures small near suture, becoming larger towards sides, mostly isodiametric in male, 
strongly transverse in female; interstriae narrow, straight, sometimes partly sinuous; gaps 
between punctures convex, forming a reticulate pattern with interstriae in male, more strongly 
raised and linking together in female, producing a rectangular mesh pattern ; all raised surfaces 
with very small olive-brown scales which are very dense on declivity and along suture ; punctures 
filled with larger round white scales which form continuous tracts towards apex (at least at sides) ; 
centres of punctures bare, hence pupillate. Mesepisternum, mesepimeron and metepisternum 
with dense round white scales ; mesosternum microreticulate and with feather-scales in punctures 
anteri