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BULLET
THE BRITISH MUSEUM
(NATURAL HISTORY)
ENTOMOLOGY
VOL A
1951-1952
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
LONDON : 1952
DATES OF PUBLICATION OF THE PARTS
No. 1. 12 December 1952
No. 2. 21 December 1952
No. 3. 24 November 1952
No. 4. 6 February 1952
No..5. 21 April 1952
No. 6. 18 December 1952
PRINTED IN
GREAT BRITAIN
AT THE
BARTHOLOMEW PRESS
DORKING
BY
ADLARD AND SON, LTD,
CONTENTS
ENTOMOLOGY VOLUME II
. The early literature on Mallophaga. Part II. 1763-75.
By THERESA CLAY and G. H. E. HOPKINS (Pls. 1-3.)
. A revision of the Australian and Tasmanian Gripopterygidae
and Nemouridae (Plecoptera). By D. E. KIMMINS
. Sphecidae and Pompilidae (Hymenoptera) collected by Mr.
K. M. Guichard in West Africa and Ethiopia. By G. ARNOLD
(Pls. 4-5.)
. A collection of Fleas from the bodies of British Birds. With
notes on their distribution and host preferences. By MIRIAM
ROTHSCHILD
. The Sub-Genus Stegomyia (Diptera: Culicidae) in the Ethio-
pian Region. Part I. A preliminary study of the distribu-
tion of species occurring in the West African Sub-Region with
notes on taxonomy and bionomics. By P. F. MATTINGLY
. A review, and a revision in greater part, of the Cteniscini of
the Old World (Hym., Ichneumonidae). By G. J. KERRICH
mm j . L va
Index to Volume II £ “& uk UY hs Jr ‘|
43
95
185
Pp. 103.
P2035:
P. 225.
CORRIGENDA
The legends to Figs. 5 and 5a (and to Pl. 4, fig. 1) to read
Cerceris longiuscula instead of Cerceris guichardi.
Below last entry under ‘“ Host ’’ add: “ Larus argentatus L.
Herring Gull. BM.— 1 —.”
Line 7 from bottom should read ‘‘ an atypical female, probably
of C. borealis, and one male of C. garet (Rothschild, 1948), etc.”
Sa
17 DEC 1954
THE
ON MALLOPHAGA
(PART II)
THERESA CLAY
AND
G. H. E. HOPKINS
| BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 2 No. 1
LONDON : 1951
THE EARLY LITERATURE ON
MALLOPHAGA
BY
THERESA CLAY
AND ?
| YZ ‘BS
G. H. E. HOPKINS \
PART II. 1763-1775
Pp. 1-36; Pls. 1-3; 45 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 2 No. 1
LONDON : 1951
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is tssued
in five series, corresponding to the Departments of the
Museum.
Parts appear at irregular intervals as they become
veady. Volumes will contain about three or four hundred
pages, and will not necessarily be completed within one
calendar year.
This paper 1s Vol. 2, No. r of the Entomology sertes.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued November 1951 Price Ten Shillings.
THE EARLY LITERATURE ON MALLOPHAGA
By THERESA CLAY and G. H. E. HOPKINS
PART II, 1763-1775
SYNOPSIS
This is the second part of a series of papers in which it is intended to review the species of Mallophaga
described between 1758 and 1818. This part deals with the twenty species described by G. A. Scopoli in
1763 and 1772, the single species described by Pontoppidan in 1763, and the three species described by
J. C. Fabricius in 1775.
IN the first part of this work (Clay and Hopkins, 1950) we remarked on the possibility
that specimens in the Nitzsch collection that are not the types of names given by
Nitzsch might wrongly be considered as types by writers who do not accept the rules
of nomenclature. We were not aware, when we wrote this, that Kéler (1941) had listed
many of the Halle specimens of species described by Nitzsch in 1818 by reference to
older descriptions as the types of the 1818 names. Leaving aside queried instances,
the names affected that concern us in our discussion of the early literature are as
follows:
Trichodectes crassus Nitzsch (p.129). The specimens of Pediculus melis in Fabricius’s
collection are still preserved at Copenhagen and are of necessity the types of
crassus as well as of melis.
Trichodectes scalaris Nitzsch (p. 130). Since scalaris is merely a nomen novum for
Pediculus bovis Linn., the types are the lost specimens of Linné, which we have
replaced (1950: 227). The specimens in Nitzsch’s collection have no special
status.
Trichodectes longicornis (p. 130). Although the specimens in the Nitzsch collection
at Halle are not types, there is no reason why one of them should not be erected
as a neotype, and we intend to take this course in the final instalment of this
work. Until this is done they have no special status.
No host-records are given in Kéler’s list mentioned above, but the hosts can be
ascertained by reference to his two papers ‘Baustoffe zu einer Monographie der
Mallophagen’, Parts I and II (1938 and 1939), and (as it happens) in none of the
above-mentioned cases is the host of Nitzsch’s material definitely different from that
of the earlier author on whose description or figure Nitzsch’s 1818 names rest their
sole claim to validity. We therefore considered whether we could accept Kéler’s
listing of these specimens as types of Nitzsch’s names as constituting them neotypes
of the earlier names, but we have decided against this. In the instances mentioned
above (except in the case of crassus) this course might be possible, but in the case of
crassus the original types are still in existence at Copenhagen and in other cases, also,
it would be quite impossible to accept Kéler’s listing. As an instance, Goniodes dis-
similis was first described by Denny in 1842 and the specimens listed by Kéler (p. 133)
would (if they have any status at all) be types of Gontodes dissimilis Nitzsch 1874 (not
G. dissimilis Denny 1842, nor G. dissimilis Gurlt 1842) if the species were not the
same. We think the principle underlying these statements of fact to be of such
4 THE EARLY LITERATURE ON MALLOPHAGA
importance that we find ourselves unable to stretch a point by regarding Kéler’s
listing of specimens as ‘Typen’ of Nitzsch’s 1818 names as constituting them neo-
types of the older names on which Nitzsch based them in 1818. We are confirmed in
this attitude by the fact that Kéler has evidently not studied the early literature at
all—‘ equi (Lin.)’, for instance, does not exist, no species having been described under
this name prior to 1842, when Denny described it.
We are indebted to the Trustees of the British Museum for permission to publish
Figs. 9, 10, 21, 31, 32, 34, drawn by Mr. A, J. E. Terzi, and to Colonel Meinertzhagen
for permission to publish Figs. 8, 16-18, 23-25, 29-30, 38-40 drawn by Mr. R. S.
Pitcher. Figs. 2, II, 12, 15, 20, 33, 36 were drawn by Miss B. A. Read. The remaining
figures were drawn by Miss T. Clay. We are also indebted to Captain W. H. Pollen
for the photographs on PI. II, figs. 1-2; the other photographs, with the exception of
Pl. I, fig. 2, were taken by Mr. H. M. Malies.
In the measurements given under each species the length of the male genitalia
has almost always been taken from a specimen other than the neotype: it has,
therefore, not been considered necessary to note this fact in individual instances,
as we did in Part I of this work.
ScopoLt 1763 (Entomologia Carniolica. Vindobonae: 381-385)
The names in this work are unquestionably in valid form, and the descriptions,
though very brief, are usually adequate for the recognition of the genus, which is as
much as we can expect from the old descriptions. Scopoli’s collections are stated to
have been destroyed, either by fire or shipwreck, in 1776.1
Pediculus haematopus (p. 381)
The description is undoubtedly that of a Philopterus s.l. and the host-record is
‘Habitat in Falcone Palumbrio, & Sirige Ulula’. The name would, therefore, refer to a
mixture of a Craspedorrhynchus and a Strigi-
philus but for the fact that Scopoli notes that
the specimens from the owl were slightly
different, thus constituting Falco palumba-
vius = Accipiter gentilis (Linn.) the type-
host. J. C. Fabricius (1775: 806) unneces-
sarily renamed the species Pediculus Strigis
(nec P. strigis Pontoppidan, 1763), and
Nitzsch (1818: 290) again renamed it Philo-
pterus (Docophorus) platyrhynchus. Harrison’s
erroneous belief that haematopus is preoccu-
pied by haematopi Linn., 1758 (which is not
a homonym), has caused most later authors
to call the species by the name given to it by
\ Nitzsch.
Fic. 1. Craspedorrhynchus haematopus aoannies benencrts Latreille 2 “E056 "{y0e
(Scopoli), male genitalia. x 351. Linné 1758) is also a synonym of Craspe-
dorrhynchus haematopus (Scopoli). As pointed
1 Horn, W., and Kahle, J., 1936. Uber entomologische Sammlungen. Ent. Betheften 3: 252.
THE EARLY LITERATURE ON MALLOPHAGA 5
out by Hopkins (1949, Ann. Mag. nat. Hist. (12) 2: 48), it is based on a drawing
copied from one copied by Hooke (in Albin’s Natural History of Spiders and Other
Curious Insects, 1736) from one of Redi’s figures of ‘ Pollini dell’ Astore’, so the host
is Accipiter gentilis (Linn.).
Neotype male (Fig. 1) and neallotype female (Fig. 2) of Craspedorrhynchus haema-
topus (Scopoli) from Accipiter g. gentilis (Linn.) from Estonia (Meinertzhagen collec-
tion slide No. 1464); these specimens agree in characters other than those figured
here with the figures published by Merisuo (1945, pl. 2, fig. D; pl. 3, figs. D, d).
Neoparatypes: 14 males and 12 females from the same host-form, Estonia.
Fic. 2. Craspedorrhynchus haematopus (Scopoli), terminal segments of female abdomen. x 102.
The neotypes are automatically also neotypes of Craspedorrhynchus strigis (J. C.
Fabricius) 1775 and of Craspedorrhynchus platyrhynchus (Nitzsch) 1818.
Pediculus maximus (p. 382)
The particulars given, especially the size, can apply only to a Laemobothrion, and
this fact has been recognized ever since the genus was erected. The host is Falco
Buteo = Buteo buteo (Linn.).
J. C. Fabricius (1776: 309) renamed the species Pediculus Buteonts ; Nitzsch (1818:
301) proposed the name Liotheum (Laemobothrion) giganteum for maximus Scopoli,
buteonis Fabricius, and circi Geoffroy, and we shall deal with Nitzsch’s name in such
a way as to make it a synonym of Laemobothrion maximum (Scopoli).
Eichler has figured the species (1941, fig. 28, and 1942, fig. 4) and in the second of
these papers he erects (p. 59) a neotype for the species, the neotype being from a
specimen of Buteo vulgaris collected at Agna Manja, Teneriffe ; Dr. Eichler informs us
(in litt.) that this was probably Buteo buteo insularum Floricke.
Eichler’s neotype of Laemobothrion maximum (Scopoli) is automatically also neo-
type of L. buteonis (J. C. Fabricius), but not of L. giganteum (Nitzsch).
6 THE EARLY LITERATURE ON MALLOPHAGA
Pediculus coarctatus (p. 382)
The host-record is ‘Lanio Collur’. Harrison (1916: 12) considered the name to
apply to a Menopon s.l. and placed M. fuscocinctum Denny and other names in the
synonymy, but this identification of Scopoli’s species is quite certainly incorrect, for
the mention of a white abdomen with seven conical fuscous spots on each side and
with a fuscous apical fascia, coupled with the host-record, is completely diagnostic of
a Philopterus. J. C. Fabricius (1798: 570) described a Pediculus lani, from Lanius
collurio Linn., which we consider to be the same species, and Schrank (1803: 187)
described a Pediculus collurtonis from ‘Dorndreher’ (= L. collurio). Schrank’s
description is independent, but he quotes coarctatus as a synonym and his species is
certainly a Philopterus ; we identify this, also, as a synonym of coarctatus.
The species of Philopterus parasitizing the small European Passeres are very
similar to each other, and in many cases it is difficult (if not impossible) to distinguish
the females of two species. Specific differences in the head may be found in the form
of the hyaline margin and the shape of the dorsal anterior plate (clypeal signature).
The measurements of the head are, in general, unreliable characters ; not only is there
considerable variation within a species but, although species may be distinguished
from each other by the means of the length or breadth, there is always considerable
overlap, making the identification of many specimens by measurements impossible.
Again, in most species the proportions of the head show no reliable specific differences ;
the cephalic index (breadth : length) tends to remain constant and gives less indica-
tion of differences than do the actual measurements of length and breadth. Tables of
the C.I. and breadth measurements for three species illustrate these points (Tables
I-4).
As in most species with heavily sclerotized plates, there is always considerable
variation in the outline of these plates. The exact outline of the prosternal plate, the
abdominal tergites, and the sternites of the male and female genital regions cannot,
therefore, be used as specific characters. The prosternal plate may vary considerably
in outline within one species (Fig. 3), but in some species (P. citrinellae (Schrank), for
instance) the posterior part is more heavily
sclerotized and pigmented. The female genital
plate varies in outline (Fig. 44 shows the out-
line in four specimens of P. fringillae taken
from the same host individual), but such
characters as the curvature of the anterior
margin and the ratio of breadth to length may
Fic. 3. Philopterus coarctatus (Scopoli), be of specific importance. Comparison of the
prosternal plates of 6 females taken from female genital plate of P. coarctatus with that
one host individual. x 175. stig . :
of P. fringillae shows that the anterior margin
always tends to be more rounded in the former than in the latter. The ratio of
breadth to length is given in Table 5; the variation is caused by the variation in
the length of the plate due to the lack of posterior sclerotization in some specimens, the
breadth tends to be fairly constant.
The chaetotaxy of the thorax and abdomen shows much variation, but some
THE EARLY LITERATURE ON MALLOPHAGA
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Fic. 4 Fic. 5
Fias. 4-5. Philopterus coarctatus (Scopoli): 4. Male (setae not shown on legs).
Fic. 6
Fics. 6-7. Philopterus coarctatus (Scopoli), terminal segments of female abdomen: 6. Ventral
view. X152. 7. Dorsal view. X 107.
5. Male genitalia. x 196.
FIG. 7
8 THE EARLY LITERATURE ON MALLOPHAGA
species can be distinguished by the unusually large or small number of setae on cer-
tain segments.
The male genitalia, as is usual throughout the Ischnocera, give the most reliable
specific differences, although in these characters, also, the differences are of small
magnitude.
Philopterus coarctatus (Scopoli) (Figs. 3~7; Pl. I, fig. 1; Tables 1-5) is distinguished
in the male by the characters of the genitalia, and in the female by the form of the
hyaline margin, the prosternal plate, and the proportions of the genital plate.
Measurements in mm.
Male Female
Length Breadth Length Breadth
Head ‘ : : 0°55 0°50 0:60 0°57
Prothorax. ie 0:28 ar 0°32
Pterothorax : a 0°42 ee 0-48
Abdomen . : : 0-78 0-62 I-02 0-80
Total 5 ; 1°55 mA 1°67
Genitalia . ‘ : 0:27 did
Neotype male (Figs. 4-5) and neallotype female (Figs. 3, 6, 7; Pl. I, fig. 1) of
Philopterus coarctatus (Scopoli) from Lantus c. collurio Linn., from Yugoslavia (British
Museum (Natural History) collection, slide No. 485). Neoparatypes: 28 males and
51 females from the same host-form from Yugoslavia, Poland, Estonia, Sweden, the
British Isles, and the Anglo-Egyptian Sudan.
Pediculus ocellatus (p. 382)
The unusually good description places it beyond doubt that this species is a
Philopterus, as has long been accepted. The hosts, however, are Corvus corax and
C. cornix, so that the name originally applied to a mixture of Philopterus corvi (Linn.)
and a second species which had not then been described. It has commonly been ac-
cepted as a principle that the first host mentioned by an author should be regarded
as the type-host, but we can only accept this principle when there is no strong reason
in favour of its rejection. In the present instance the fact that ocellatus has been
accepted since 1818 (at least) as referring to the species found on Corvus corone cornix
would be a very strong argument against accepting Corvus corax as type-host even if
Nitzsch (1818: 290) had not restricted ocellatus to the species found on C. corone and
its sub-species.
J. C. Fabricius (1775: 807) unnecessarily renamed the species as Pediculus Cornicts
and dropped the mention of Corvus corax as a host. Nitzsch (1818: 290) retained
Scopoli’s name, contrary to his usual practice, and the species was subsequently
always known as ocellatus (the authorship usually wrongly ascribed to Nitzsch) until
Thompson (1935: 214) mistakenly replaced this name by Philopterus corvi (Linn.),
with which we have already dealt and which is not the same species.
Philopterus ocellatus (Scopoli) is distinguished from P. corvi (Linn.) by having the
THE EARLY LITERATURE ON MALLOPHAGA 9
anterior plate (= clypeal signature) heavily sclerotized and pigmented posteriorly,
and by the characters of the male genitalia and female genital region.
Fic. 8. Philopterus ocellatus (Scopoli), male. Fic. 9. Philopterus ocellatus (Scopoli),
male genitalia.
Measurements in mm.
Male Female
Length Breadth Length Breadth
Head : A 7 0-75 0:76 0-80 0:80
Prothorax. rae 0°47 fat 0-48
Pterothorax wit 0:68 ae 0:70
Abdomen . : : 1°34 err 1°36 1°32
Total : ‘ ; 2°46 oe 2°50
Genitalia . : . 0:48 i
Neotype male (Figs. 8-9, 11; Pl. I, fig. 2) and neallotype female (Figs. 10, 12)
of Philopterus ocellatus (Scopoli) from Corvus corone sardonius Kleinschmidt from
ENTOM. 2, I B
Io THE EARLY LITERATURE ON MALLOPHAGA
Yugoslavia (British Museum (Natural History) collection, slide No. 484). Neopara-
types: 10 males and g females from the same host-form from Yugoslavia, Greece,
Palestine, and Egypt and 40 males and 30 females from Corvus corone cornix Linn.
from the British Isles, Estonia, and Sweden.
ist Three WAT an
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Fic. 10. Philopterus ocellatus (Scopoli), female.
These neotypes are also neotypes of Philopterus cornicis (J. C. Fabricius). There is
no such species as Philopterus ocellatus (Nitzsch).
Pediculus dolichocephalus (p. 382)
There has never been any doubt about the identity of this species, the very brief
description agreeing with no oriole-parasite except Ricinus. The original host-record .
is ‘Halitat in Coracia Onolo’, which Harrison (1916: 66) misconstrued as meaning
Coracias and Oriolus, evidently not realizing that in Scopoli’s time the oriole was
THE EARLY LITERATURE ON MALLOPHAGA. II
contained in the genus Coracias. Similar errors will be mentioned under Pediculus
ardealis and P. troglodytis.
J. C. Fabricius (1776: 310) renamed the species Pediculus Orioli, and Nitzsch (1818:
302) called it Liotheum (Physostomum) sulphureum, but there is not the slightest
Fic. 12
Fics. 11-12. Philopterus ocellatus (Scopoli), terminal segments of abdomen: 11. Male. x 89.
12. Female. x67.
justification for either of these nomina nova. The description given by Fabricius is an
abridged version of that of Scopoli, and Nitzsch gives no description and no ‘indica-
tion’ except a reference to Scopoli, so both these names derive their validity solely
from Scopoli’s description.
This species is distinguished from other species of Ricinus by the characters of the
12 THE EARLY LITERATURE ON MALLOPHAGA
mandibles (Fig. 13),! by the shape of the head and the characteristic colour-pattern
of the body (PI. I, fig. 3), the characters of the male genitalia (Fig. 14), and the
terminal segments of the female abdomen (Fig. 15).
Fic. 13 Fic. 14
Fics. 13-14. Ricinus dolichocephalus (Scopoli): 13. Left mandible. Xx 191. 14. Male genitalia.
Fic. 15. Ricinus dolichocephalus (Scopoli), terminal segments of female abdomen, ventral. x 75.
Measurements in mm.
Male Female
Length Breadth Length Breadth
Head : : ; 0-76 0:68 0:97 0-78
Prothorax. : oe 0°57 ate 0-70
Pterothorax ; ; ae 0°75 Ke I'03
Abdomen . : 3 ote 0:83 aie I'16
Total : : ‘ 3°50 Ae 4°65
Genitalia . : = 0°46
Neotype male (Figs. 13-14) and neallotype female (Fig. 15; Pl. I, fig. 3) of Ricinus
dolichocephalus (Scopoli) from Oviolus o. oriolus (Linn.) from NE. Poland (Meinertz-
1 These structures show characters of specific value throughout the genus Ricinus.
THE EARLY LITERATURE ON MALLOPHAGA 13
hagen collection, slide No. 4190). Neoparatypes: 3 males and 10 females from same
host-form, NE. Poland, Switzerland, and Cyprus.
These neotypes are, of necessity, also neotypes of Ricinus oriolt (J. C. Fabricius)
and of R. sulphureus (Nitzsch).
Pediculus fasciatus (p. 383)
The host is Cuculus canorus and the description, unmistakably that of the charac-
teristic Cuculiphilus found on this bird, has long caused the name to be correctly
ascribed to this louse.
J. C. Fabricius (1775: 807) renamed the
species Pediculus Cucult, his description be-
ing copied from that of Scopoli, and Nitzsch
(1818: 300) proposed Liotheum (Menopon)
phanerostigmaton as a nomen novum for it.
The species went under this latter name
(and its variant, phanerostigma Giebel)
until Harrison (1916: 47) restored Scopoli’s
name. Uchida (1926: 47) designated Pedi-
culus fasciatus Scopoli as type species of
Cuculiphilus.
Neotype male (Figs. 16-17) and neallotype
female (Fig. 18) of Cuculiphilus fasciatus
(Scopoli) from Cuculus c. canorus Linn.
from NE. Poland (Meinertzhagen collection,
slide No. 4211). Neoparatypes: 3 males and
8 females from same host-form, NE. Poland,
Ushant (France), and Tanganyika Territory.
These neotypes are automatically also
neotypes of Cuculiphilus cuculi (J. C. Fabri-
cius) and of C. phanerostigmaton (Nitzsch).
Neotype of Cuculiphilus phanerostigma
(Giebel), a male from Cuculus c. canorus
Linn. from Ushant, France (Meinertzhagen
collection, slide No. 780), agreeing with the
neotype of Cuculiphilus fasciatus (Scopoli).
Fic. 16. Cuculiphilus fasciatus (Scopoli),
male.
- Measurements in mm.
Male Female
Length Breadth Length Breadth
Head ; ‘ ‘ 0°35 0°58 0°35 0°63
Prothorax. as 0°37 aie 0:39
Pterothorax ‘ a 0°45 ee 0:48
Abdomen . é ‘ 0-90 0°73 1:02 0:82
Total * , é 1°52 be I'79
Genitalia . ‘ . 0-48 3
14 THE EARLY LITERATURE ON MALLOPHAGA
Pediculus auritus (p. 383)
The hosts are Picus major and P. martius, i.e. Dryobates major (Linn.) and Dryo-
copus martius (Linn.), and the description unquestionably refers to a Penenirmus. In
the absence of any indication to the contrary, we regard the first host mentioned by
Scopoli as the type-host. Schrank (1803: 188) described a Pediculus Pict from
‘Schwarzpecht’ (= Dryocopus martius), placing auritus Scopoli as a synonym, but
we cannot accept this as a restriction of auritus to one host, especially as Schrank also
placed Pulex picae Redi (i.e. Myrsidea picae (Linn.), from Pica pica) as a synonym.
Fic. 17. Cuculiphilus fasciatus (Scopoli), Fic. 18. Cuculiphilus fasciatus (Scopoli),
male genitalia. female.
Burmeister (1838: 427) and Giebel (1874: 94, pl. Io, fig. 3) described as Docophorus
superciliosus an insect that seems to be certainly the same as Penenirmus auritus
(Scopoli) and is from Dryobates major (Linn.). Harrison (1916: 88) listed auritus in
Philopterus with superciliosus as a synonym.
THE EARLY LITERATURE ON MALLOPHAGA 15
Neotype male (Fig. 19; Pl. I, fig. 4) and neallotype female (Fig. 20) of Penenirmus
auritus (Scopoli) from Dryobates major pinetorum (Brehm) from Yugoslavia (British
Museum (Natural History) collection, slide No. 498). Neoparatypes: 12 males and
21 females from same host-form, Jugoslavia and NE. Poland; 23 males and 26
females from Dryobates major major (Linn.), Estonia.
Measurements in mm.
Male Female
Length Breadth Length Breadth
Head . P 5 0°53 0:48 0°57 0:50
Prothorax. . as 0:29 As 0°30
Pterothorax : ‘ 0°40 0:46 0°43 0-51
Abdomen . 4 . 0°93 0-60 I-12 0°65
Total ‘ 2 5 1°72 as I'99 ae
Genitalia . ; ; 0:28 Vs a5
enenne
“0s:
Fic. 19 Fic. 20
Fics. 19-20. Penenirmus auritus (Scopoli): 19. Male genitalia. 20. Terminal segments of female
abdomen. x 87.
Neotype of Penenirmus superciliosus (Burmeister) a male from Dryobates m. major
(Linn.), Estonia (Meinertzhagen collection, slide No. 1568), agreeing with our descrip-
tion and figures of the neotype of Peneniymus auritus (Scopoli).
Pediculus dentatus (p. 383)
The host-record is merely ‘in Anate’, but as the next species described by Scopoli
is from Anas boschas, now known as A. platyrhynchos Linn., we think it only reason-
able to assume that dentatus was from the same host, the specific name of which was
inadvertently omitted. The description is definitely that of an Anatoecus. Eichler
16 THE EARLY LITERATURE ON MALLOPHAGA
(1946: 75) selected Anas platyrhynchos as type-host of this species, but without
erecting neotypes.
We can find nothing in the subsequent literature that adds anything to our know-
ledge of this species ; Harrison (1916: 13, 93) referred it to Philopterus, placing most
Fic. 21 Fic. 22
Fics. 21-22. Anatoecus dentatus (Scopoli), male. 21. Genitalia. 22. Terminal segments of
abdomen. X 97.
of the duck-infesting species as synonyms, but pending redescription of the numerous
species that have been described from ducks we are unable to suggest which names
are synonyms of Anatoecus dentatus (Scopoli).
Cummings (1916: 652) was the first author to draw attention to the two distinct
forms of genitalia found in male Anatoecus—those with and those without the so-called
‘effractor’. Unfortunately his otherwise excellent account of various species found
on the Anatidae is rendered largely useless by his failure to give any indication of the
THE EARLY LITERATURE ON MALLOPHAGA 17
host of the majority of the species that he figured. Cummings did not discuss the
distribution of the two groups of species, but it has been found that on all the European
ducks from which material is available there are two species of Anatoecus, the males
of which are separable by the presence or absence of the effractor in addition to other
less obvious characters. It has not been possible to assign the females to the different
males with certainty, and for this reason no female neallotype nor neoparatypes will
be designated for dentatus or other species of Anatoecus with which we shall deal.
Cummings made Philopterus icterodes Nitzsch the type-species of Anatoecus and
placed it in the group without an effractor; this species will be dealt with later and
Cummings’s interpretation will be followed. Cummings did not mention dentatus
Scopoli, and in order to fix representative species of the two main groups by the
erection of neotypes and publication of figures we have chosen the species from Anas
platyrhynchos in which the effractor is present to bear the name dentatus Scopoli.
Measurements in mm.
Male
Length Breadth
Head : : a 0°47 0°43
Prothorax. ; : se 0:28
Pterothorax ‘ ; Re 0°37
Abdomen . : : 0:72 0:60
Total : . : 1°45
Genitalia . . ; 0-46
Measurements in mm. of heads of specimens examined
Individual No. 7; 2 3 4 5 6 7 8 9 Io
Total length in midline - | 0°47 | 0°50 | 0°47 | 0°48 | 0°45 | 0°45 | 0°50 | 0°46 | 0°47 | 0°48
Length of hyaline margin . | 0-16 j 0-17 | 0-16 | 0-16 | O-15 | O-15 | O-17 | O16 | O-15 | O-T5
Breadth at temples. 0°43 | 0°43 | 0°42 | 0°43 | 0°42 | 0-42 | 0°43 | O-41 | 0-42 | 0°43
Breadth at base of hyaline
margin. : 0°25 | 0:26 | 0°25 | 0°25 | 0°25 | 0:26 | 0:27 | 0°24 | 0:26 | 0:25
Neotype male (Figs. 21-22; Pl. I, fig. 5) of Anatoecus dentatus (Scopoli) from Anas
p. platyrhynchos Linn. from NE. Poland (Meinertzhagen collection, slide No. 4176).
Neoparatypes: g males from same host-form, Poland and England.
Pediculus crassicornis (p. 383)
The host is Anas Boschas and the louse is undoubtedly an Anaticola.
Schrank (1781: 503) described as Pediculus Anatis a form which is certainly partly
crassicornis and which will be discussed later ; his host is ‘ Anas Boschas, varietas fera’,
but there is some evidence of confusion with Anaticola anserts (Linn.). J. C. Fabricius
(1798: 571) also described a Pediculus Anatis from Anas Boschas, his description
being apparently independent. Nivmus crassicornts ‘Olfers’ (see Harrison, 1916: 13)
does not exist, von Olfers correctly attributing the name to Scopoli, though his host-
records indicate the confusion between this species and anseris for which Schrank
appears to be responsible. Nitzsch (1818: 292) proposed Philopterus (Lipeurus)
ENTOM. 2, I Cc
18 THE EARLY LITERATURE ON MALLOPHAGA
squalidus as a nomen novum for Pediculus anatis Fabricius; this name, also, is a
synonym of Anaticola crassicornis (Scopoli).
This species is distinguished from Anaticola anseris (Linn.) (see Clay and Hopkins,
1950: 239) by the characters of the anterior region of the head and the shorter penis
(see measurements).
Fic. 23 Fic. 24
“‘
Fics. 23-24. Anaticola crassicornis (Scopoli): 23. Male. 24. Male genitalia.
x 144; a. Tip of paramere from side. x 437.
Measurements in mm.
Male Female
Length Breadth Length Breadth
Head ° é . 0:67 0°43 0-68 0-46
Prothorax. os 0:27 ie 0°30
Pterothorax . she 0°45 te 0°47
Abdomen . : - I-63 0°50 2:10 0°53
Total ; A 5 2°92 sys 3°40 <
Genitalia . ; : 0-70 a ete
Length of penis (number of specimens in brackets).
Anaticola crassicornis . . 0-13 mm. (1), 0-15 (7), 0-16 (x).
Anaticola anseris . : - 0-27 mm. (3).
THE EARLY LITERATURE ON MALLOPHAGA 19
Neotype male (Figs. 23-24) and neallotype female (Fig. 25) of Anaticola crassicornis
(Scopoli) from Anas p. platyrhynchos Linn., from NE. Poland (Meinertzhagen collec-
tion, slide No. 4242). Neoparatypes: 12 males
and 20 females from same host-form, Poland,
Hungary, and the British Isles.
Pediculus pilosus (p. 384)
The host is Fulica atra Linn., but all the
description given is that the louse has a red
head, a long abdomen, and a particularly
hairy anus with the hairs parallel.
Although this description is extremely un-
satisfactory and Harrison (1916: 17) rejected
pilosus as unrecognizable, we think that the
description of the anus can only apply (among |
coot-parasites) to a Pseudomenopon, and the
other characters are not inconsistent with this
identification. Pseudomenopon could certainly
be described as having a red head and its
abdomen is moderately elongate, much more
so than that of Incidifrons, for instance, which
seems the only alternative.
The species was redescribed as Menopon
tridens by Burmeister (1838: 440) from material
from the same host (Fulica atra) ; Piaget (1880:
480), wrongly taking Gallinula chloropus as
the type-host of tvidens, renamed the form
from Fulica atra as Menopon tridens var.
major (preoccupied by Menopon quadrifascia-
tum var. major Piaget 1880: 441); Eichler
(1937: 97), noting that the name of Piaget’s Fic. 25. Anaticola crassicornis (Scopoli),
variety from the coot was preoccupied, re- soele
named it Pseudomenopon thompsont. Mjéberg (1910: 51) made Menopon tridens ‘N.’
the type-species of his genus Pseudomenopon.
Neotype male and neallotype female of Pseudomenopon pilosum (Scopoli) from
Fulica a. atra Linn. from Ireland (Meinertzhagen collection, slide No. 16388). These
specimens agree with the figures published by Ferris (1924, Parasitology, 16: 64,
fig. 4), although the male drawn by Ferris was from the American coot, Fulica a.
americana Gmelin. The stout spine-like seta and the smaller seta below it on each side
of the dorsal surface of the prothorax are not shown in fig. 4a (Ferris, 1924), nor are
the three setae on each side of the dorsal surface of the metathorax. Neoparatypes:
12 males and 41 females from Fulica a. atra Linn., British Isles.
Neotype of Pseudomenopon tridens (Burmeister) a male (Meinertzhagen collection,
slide No. 10510) from Fulica a. atra Linn. from Ireland. This specimen agrees with
the neotype of P. pilosum (Scopoli).
20 THE EARLY LITERATURE ON MALLOPHAGA
The types of Pseudomenopon thompsoni Eichler (Menopon var. major Piaget 1880:
480 nec 441) are in the British Museum and also agree with the neotypes of P. pilosum
(Scopoli).
Pediculus colymbinus (p. 384)
The host is Colymbus auritus Linn. (= Podiceps auritus (Linn.) of European
authors), and Scopoli states that young specimens have an ovate abdomen with
elliptical black dorsal spots and that the adult becomes rufous. This description is
Fic. 26 Fic. 27.
Fics. 26-27. Aquanirmus colymbinus (Scopoli), male: 26. Genitalia. x 164.
27. Terminal segments of abdomen. xX 223.
Fic. 28
Fic. 28. Aguanirmus colymbinus (Scopoli), terminal
segments of female abdomen, ventral. x 208.
extremely inadequate, but the black dorsal spots (if not gut-contents) confine us to
the Ischnocera and the only genus of Ischnocera reliably reported from the grebes is
Aquanirmus, Denny (1842) described a Nirmus fusco-marginatus (a male Aquanirmus)
from the same host and a Nirmus podicepis (a female of the same genus) from another
THE EARLY LITERATURE ON MALLOPHAGA 21
species of grebe, and later authors have identified what Scopoli considered to be young
specimens with N. podicepis Denny. Although thinking the identification of colym-
binus as an Aquanirmus far from certain, we see nothing in the description that dis-
proves it and there is no object in disturbing the accepted application of Scopoli’s
name. We cannot accept the assumption that Denny’s male and female are conspecific
pending a much more careful examination of the forms occurring on different species
of grebes than has yet been made, but the lectotype of fusco-marginatus Denny
agrees with our male neotype of colymbinus.
Measurements in mm.
Male Female
Length Breadth Length Breadth
Head : : : 0-46 0°35 0°53 0°42
Prothorax. as 0:28 . 0°33
Pterothorax : ce 0°37 oye 0°45
Abdomen . , ss 0:87 0-40 I'47 0°55
Total - 2 : I-65 Ae 2°35 aie
Genitalia . ‘ ; 0°37 5c
Neotype male (Figs. 26-27; Pl. II, fig. 1) and neallotype female (Fig. 28; Pl. IT,
fig. 2) of Aquanirmus colymbinus (Scopoli) from Colymbus auritus Linn. from Devon,
England (Meinertzhagen collection, slide No. 14736). Neoparatypes: I male and
6 females from the same host-form, England.
There are two males and one nymph of Nivmus fusco-marginatus from Podiceps
auritus (= Colymbus auritus) in the Denny collection ; one of these males (slide No.
349) is hereby selected as lectotype of Aquanirmus fusco-marginatus (Denny).
Pediculus ardealis (p. 384)
Harrison (1916: 11) wrongly considered this name to be a synonym of Ardeicola
ardeae (Linn.), doubtless through misreading of the host-record ‘Ardea Ciconia’
(= Ciconia ciconia). Of the species found on Ciconia ciconia (Linn.), Scopoli’s very
poor description could only apply to Neophilopterus incompletus (Denny) or to a
nymph of Ardeicola ciconiae (Linn.). But, in addition to the description, Scopoli
gives two references: to the less hairy figure on Frisch Insect. 5, pl. 4, and to Linné,
1758, p. 613, No. 26. The reference to Frisch is an obvious slip, for plate 4 only con-
tains one insect, but the reference to Linné is to Arvdeicola ciconiae, and under this
species Linné refers to Frisch’s plate 6; the latter plate does show two insects, one
with hairs and the other without, both belonging to the genus Ardezcola. It is, there-
fore, certain that Scopoli’s insect was the Ardeicola and not the Neophilopterus.
Neotype of Ardeicola ardealis (Scopoli) a male from Ciconta c. ciconia (Linn.) from
South Africa (British Museum collection, slide No. 430) that agrees with the neotype
of Ardeicola ciconiae (Linn.) (see Clay and Hopkins, 1950: 253).
Pediculus ovalis (p. 384)
The host is Scolopax arquatus and the description states that the louse is smaller
than Pediculus humanus, has eight abdominal segments, and a depressed ovate body,
22 THE EARLY LITERATURE ON MALLOPHAGA
the antennae are shorter than the head but longer than the thorax, and the colour is
rufous-brown. No subsequent author has anything useful to say about the species,
and Harrison (1916: 17) discards it as unrecognizable, but we claim that it is recogni-
zable with certainty.
Admittedly there is no species on the curlew that agrees perfectly with Scopoli’s
Fic. 29 Fic. 30
Fics. 29-30. Cummingsiella ovalis (Scopoli): 29. Male. 30. Male genitalia.
description, and in particular none with an 8-segmented abdomen, but there are only
three species normally found on this host that could possibly be described as rufous-
brown and oval—a Cummingsiella, an Austromenopon, and perhaps Saemundssonia
humeralis (Denny). Quite apart from the facts that the Cummingsiella fits the colour-
character best and is by far the commonest of the three species (and therefore the
most likely to have been observed by Scopoli), the description of the antenna con-
vinces us that this is the form that Scopoli described. In the Austromenopon the
antennae are concealed and the palps (sometimes mistaken for antennae in early
descriptions) certainly do not project far enough beyond the margin of the head to be
described as being longer than the thorax, but in the male of the Cummingsiella the
antennae are almost as long as the thorax. This identification of Scopoli’s species
THE EARLY LITERATURE ON MALLOPHAGA 23
involves the relegation of Cummingsiella testudinaria (Denny) to synonymy, but this
is in any case inevitable because Docophorus testudinarius Denny and D. biseriatis
Denny (the name used on p. 250 in Denny’s explanation of his plate 1, fig. 6) are not
independent names but merely misdeterminations of Children’s Nirmus testudinarius
Fic. 31. Cummingsiella ovalis (Scopoli), female.
and N. biseriatus and therefore invalid. The only other synonym of Cummingstella
ovalis (Scopoli) is Nirmus pseudonirmus Nitzsch.
Neotype male (Figs. 29-30) and neallotype female (Fig. 31) of Cummingsiella ovalis
(Scopoli) from Numenius a. arquatus (Linn.) from Yugoslavia (British Museum col-
lection, slide No. 522). Neoparatypes: 86 males and gg females from the same host-
form, Yugoslavia, Hungary, and the British Isles.
Neotype of Cummingsiella pseudonirmus (Nitzsch) a male (British Museum (Natural
24 THE EARLY LITERATURE ON MALLOPHAGA
History), slide No. 523) from Numenius a. arquatus (Linn.) from Scotland, which
agrees with the neotype of C. ovalis (Scopoli).
Measurements in mm.
Male Female
Length Breadth Length Breadth
Head . . ‘ 0-70 0-72 0°75 0°73
Prothorax. F : ae 0°42 sts 0°45
Pterothorax : ‘ sis 0°65 ne 0-70
Abdomen . : : I-16 1:03 1°50 I°rl
Total Z : : 2°10 fe 2°56
Genitalia . - ; 0°65 Ste
The Denny collection contains one male and two female Cummuingstella labelled
Docophorus testudinarius but without host-record. These specimens agree with the
neotypes of C. ovalis (Scopoli).
Pediculus junceus (p. 384)
The host is Tvinga vanellus = Vanellus vanellus (Linn.), and the description is
obviously that of a Degeeriella s.l. Denny (1842: 53, 143, pl. 9, fig. 5) made an identifica-
tion of this species with which we are in full agreement ; his specimens (3 females)
came from the same host, although on p. 143 he added two other hosts. We do not
agree with Harrison in considering Pediculus tringae Schrank 1803 (nec O. Fabricius,
1780) to be the present species, and it will be discussed separately. On the other hand,
in spite of Schrank’s definite statement (1803: 190) that his Pediculus Vanelli, from
‘Kybize’ (= Vanellus vanellus) is not Scopoli’s insect we can find nothing in his
description that supports this statement and suspect that the explanation must be
that he misidentified junceus ; our suspicion that this is the case is much strengthened
by the fact that we have been unable to find any nirmoid species except junceus on
Vanellus vanellus.
Measurements in mm.
Male Female
Length Breadth Length Breadth
Head 3 2 8 0°47 0+30 0°47 0°32
Prothorax. ote 0°22 es 0:22
Pterothorax : $< 0-30 ae 0°32
Abdomen . ; : 0:93 O-4I 1:03 0°45
Total : ‘ : 1-64 aie 1-82 ie
Genitalia . 3 : 0:30
Neotype of Quadraceps junceus (Scopoli) a male (Fig. 32; Pl. II, fig.3) and neallo-
type a female (Fig. 33; Pl. II, fig. 4) from Vanellus vanellus (Linn.) from Italy
(British Museum (Natural History), slide No. 524). Neoparatypes: 154 males and
140 females from the same host-form, Italy and the British Isles.
THE EARLY LITERATURE ON MALLOPHAGA 25
Pediculus cuspidatus (p. 385)
Denny (1842: 51, 130, pl. 6, fig. 2) redescribed as Nirmus cuspidatus a species that
he took to be the same as that of Scopoli, though he thought it necessary to query the
Fic. 32 | Fie. 33
Fics. 32-33. Quadvaceps junceus (Scopoli): 32. Male genitalia. 33. Terminal segments of
: female abdomen. xX 125.
determination. Scopoli’s host-record is ‘ Rallo aquat.’, and Denny’s original specimen
(no longer in his collection) evidently also came from Rallus aquaticus Linn., because
this is the only host mentioned on p. 51. Scopoli’s description is not by any means
diagnostic, but so far as it goes it fits Rallicola at least as well as can be expected
from these old descriptions. No subsequent author seems to have seen the species.
Neotype male (Figs. 34-35; Pl. Il, fig. 5) and neallotype female (Fig. 36; Pl. II,
fig. 6) of Rallicola cuspidatus (Scopoli) from Rallus a. aquaticus Linn. from Kent,
ENTOM. 2, I D
26 THE EARLY LITERATURE ON MALLOPHAGA
England (Meinertzhagen collection, slide No. 8332). Neoparatypes: 13 males and
48 females from the same host-form, British Isles.
Fic. 34
Fics. 34-36. Rallicola cuspidatus (Scopoli): 34. Male genitalia. 35. Male antenna. x 173.
36. Terminal segments of female abdomen. X1II5.
Measurements in mm.
Fic. 36
Male Female
Length Breadth Length Breadth
Head 0°37 0°30 0°42 0°35
Prothorax. ws o-I9 0:22
Pterothorax Bi 0:27 -* 0°30
Abdomen . 0-62 0°35 0°85 0°43
Total I'll $f 1°44 ae
Genitalia . 0°25 sid
THE EARLY LITERATURE ON MALLOPHAGA 27
Pediculus bidentatus (p. 385)
The host is Columba palumbus Linn., and the reference to a hemispherical head
rules out all genera of Mallophaga known from European pigeons except Coloceras
and Campanulotes. Pediculus bidentatus was constantly placed as a synonym of
Campanulotes compar (Burmeister), from Columba livia domestica, until Harrison
(1916) reversed this arrangement. Although we find nothing in the original descrip-
tion of bidentatus that enables us to decide which of the two genera Scopoli had before
him, we think it best to accept this long-standing generic determination of his species.
This species is near C. compar (Burmeister) from Columba livia, from which it is
distinguished by its greater size (see measurements below). The chaetotaxy and
Fic. 37. Campanulotes bidentatus (Scopoli), terminal segments of female abdomen. x 121.
general characters are as figured by Kéler (1939: 158-160, figs. 89-91) for C. compar
except that in fig. 89 a median ventral seta has been omitted on segments VI and VII;
in fig. go the latero-dorsal temple spine has been omitted, the dorsal setae on abdominal
segments II-III should be about twice as long as shown, and the median dorsal setae
on segments V—VI have been omitted ; and in fig. 91 the long ventral seta on the last
abdominal segment has been omitted. The genital region of the female of C. bdentatus
is shown in Fig. 37.
Measurements in mm.
Male Female
Length Breadth Length Breadth
Head : , ? 0°34 0°47 0°43 0°53
Prothorax. ae 0:29 ae 0°34
Pterothorax F Ar: 0°37 Si 0°42
Abdomen . ‘ : 0-61 0°57 0-90 0:66
Total ; : i 1°20 at 1°58 te
Neotype of Campanutlotes b. bidentatus (Scopoli) a male (PI. III, fig. 1) and neallotype
a female (Fig. 37) from Columba p. palumbus Linn. from Somerset, England (Meinertz-
hagen collection, slide No. 864). Neoparatypes: 63 males and 58 females from the
British Isles.
ENTOM. 2, I D2
28 THE EARLY LITERATURE ON MALLOPHAGA
Comparison of breadth of heads, in mm.
Species and No. of specimens Males
Breadth in mm. | 0:38 | 0:39 | 0°40 | 0:44 | 0°45 | 0°46 | 0:47 | 0:48
C. bidentatus compar . : 2 se) 7
C. bidentatus bidentatus .
Breadth in mm. | 0:43 | 0:44 | 0:45 | 0:46 | 0-47 | 0:49 | 0-50 | O-5r | 0-52 | 0-53
C. bidentatus compar . F I 4 5 8
C. bidentatus bidentatus.
Pediculus albiventris (p. 385)
The original host-record is ‘Motacilla Troglodyte’, which Harrison (1916: 87)
wrongly took to mean Motacilla and Troglodytes, whereas only one species, the bird
now known as Tvoglodytes troglodytes (Linn.), is mentioned. Clay and Meinertzhagen
(1938: 73) showed that Pediculus albiventris (Scopoli) is the same as Docophorus
Fic. 38
Fics. 38-39. Penenirmus albiventris (Scopoli): 38. Male. 39. Male genitalia. Xx 600.
Fic. 39
THE EARLY LITERATURE ON MALLOPHAGA 29
troglodytis (Waterston), from Troglodytes troglodytes zetlandicus Hartert, and made it
the type species of Penenirmus. J. C. Fabricius (1776: 310) renamed the species
Pediculus motacillae.
Measurements in mm.
Male . Female
Length Breadth Length Breadth
Head = A “ 0°42 0°37 0:48 0°45
Prothorax. : ; ae 0-20 ie 0°25
Pterothorax 2 ic 0°33 ake 0:40
Abdomen . : ‘ 0°73 0°45 I'I3 0°63
Total ; : : 1°33 oe 1°85 ers
Neotype of Penenirmus albiventris (Scopoli) a
male (Figs. 38-39) and neallotype a female (Fig.
40) from Troglodytes t. troglodytes (Linn.) from
Wiltshire, England (Meinertzhagen collection,
slide No. 15399). Neoparatypes: 44 males and 52
females from various subspecies of Tvoglodytes
troglodytes from the British Isles.
These neotypes are also automatically neo- f
types of Penenirmus motacillae (J. C. Fabricius) f
PONTOPPIDAN, 1763 (Den Danske Atlas. 1
Kidbenhavn: 699)
Only one of the names of Mallophaga published
in this work is new.
Pediculus strigis (p. 699, pl. xxx)
This species is stated to be new and is des-
cribed with a reference to plate xxx 0, a figure of
an obvious Philopterus s.l. There is no host-
record other than that provided by the specific
name. Before we go on to discuss the identity of
the species it will be as well to consider the sub-
sequent history of the name.
Scopoli (1772: 124) gives a completely inde-
pendent description of a Pediculus strigis which
is also clearly a Philopterus s.l.; there is no host-
record, but as he describes a Hippoboscid from
Strix bubo we think it nearly certain that this Fic. 40. Penenirmus albiventris
species was also the host of the louse. J. C. Fabri- Oy od): tea
cius (1775: 806) applied the name Pediculus strigis to Pediculus haematopus Scopoli,
Fabricius’s nomen novum thus being not only unnecessary but twice preoccupied, to
say nothing of the fact that haematopus (and therefore strvigis Fabricius) is not from
an owl but from a hawk. Miiller (1776: 185) mentioned the name with a reference to
30 THE EARLY LITERATURE ON MALLOPHAGA
‘P.D.A.’ (= Pontoppidan’s Danske Atlas). O. Fabricius (1780: 216) gave references
to Pontoppidan and Miiller and also an independent description of the differences
between his material (from Stvix nyctea) and Pontoppidan’s plate. Harrison (1916:
18) did not know stvigis Pontoppidan and rejected all the other uses of the name
as either unrecognizable or preoccupied. The position may be summarized as follows:
P. strigis Pontoppidan is a valid name and easily determinable to the genus.
P. strigis Scopoli may or may not be Pontoppidan’s species but is certainly con-
generic.
P. strigis J. C. Fabricius is an unwanted nomen novum for Pediculus (now Craspe-
dorrhynchus) haematopus Scopoli. It is twice preoccupied and is not congeneric
with the others.
P. strigis ‘Miiller’ does not exist, Miiller merely listing Pontoppidan’s species.
P. strigis O. Fabricius is partly a reference to Pontoppidan and partly a new
species, the name of the latter thrice preoccupied.
In the circumstances it seems to us that much the most satisfactory course is to fix
strigis Pontoppidan in such a way that it is the same as st¢vigis Scopoli and (if possible)
so that its restoration does not upset any well-established name of later date. Not
only does Bubo bubo, which we consider to be the host of Scopoli’s species, occur in
Denmark, but Pontoppidan definitely described another parasite from this host, so
that it is very probable that his louse came from this species of bird. Two species of
Strigiphilus occur on Bubo bubo, one of which has long been known as S. heteroceros
(Nitzsch) whereas the other had not been named until Eichler (1949: 14) named it
‘Neodocophorus’ hopkinst, though it is probably a component of S. cursor (Burmeister)
as described by Giebel (1874: 70). The first species, however, cannot retain the name
Strigiphilus heteroceros (Nitzsch), because this species was not described until 1861
whereas Grube used the same name (Docophorus heterocerus) for a species found on
Strix uralensis liturata Tengmalm ten years earlier (Grube, 1851: 469). Eichler
(1949: 11) has correctly pointed out this fact and renamed the species with sexually
dimorphic antennae found on Bubo bubo as Strigiphilus gontodicerus. Both the
species concerned have, therefore, been named and both the names are of equal
seniority, so it does not matter to which species we apply the name strigis Pontoppidan
(his figure applying fairly well to either), so we have selected hopkinsi to bear
Pontoppidan’s name.
Measurements in mm.
Male
Length Breadth
Head : A ‘ 0:69 0:63
Prothorax. es 0°40
Pterothorax ; ae 0:56
Abdomen . : ; 0:94 0-84
Total : - : I'93 ea
Genitalia . : & 0°47
This species is distinguished from S. goniodicerus Eichler, from the same host, by
the antennae being similar in the two sexes and by the smaller and less complicated
THE EARLY LITERATURE ON MALLOPHAGA 31
male genitalia. Although there are two species of females represented in the available
material it has not been possible to assign them with certainty to the males, so no
neallotype or female neoparatypes have been erected.
Fic. 41 : Fic. 42
Figs. 41-42. Strigiphilus strigis (Pontoppidan), male: 41. Terminal segments of abdomen. x 74.
42. Genitalia. x 294.
Neotype of Strigiphilus strigis (Pontoppidan) a male (Figs. 41-42, Pl. III, fig. 2)
from Bubo bubo (Linn.) from Russia (Meinertzhagen collection, slide No. 10975).
Neoparatypes: g males from the same host-form, Russia and Italy (captive host).
Neotype of Strigiphilus strigis (Scopoli) a male (Meinertzhagen collection, slide No.
109750) from Bubo bubo (Linn.) from Russia, that agrees with the neotype of Sérigi-
philus strigis (Pontoppidan).
SCOPOLI, 1772 (Annus V Historico-Naturalis. [PC.] 5. Observationes Zoologicae.
Lipsiae: 124-125)
The descriptions in this work are very poor. Fortunately they are also very few.
Pediculus hirct jumoris (p. 124)
We only mention this name because we feared at first that it might be the earliest
name for a chewing louse of the goat. But we are certain that the description does
not refer to a member of the Mallophaga and we think it likely that the insect was one
of the Anoplura.
Pediculus strigis (p. 124)
This name has been dealt with under Pediculus strigis Pontoppidan and a neotype
has been erected. Sérigiphilus strigis (Scopoli) is both a homonym and a synonym of
S. strigis (Pontoppidan).
32 THE EARLY LITERATURE ON MALLOPHAGA
Pediculus ralli (p. 125)
There is no host-record other than that provided by the specific name, but the host
must be assumed to have been some bird that occurs in Carniolia and was included
in the genus Rallus in Scopoli’s time.
The entire description is that the head of the insect is bidentate and the abdomen
glabrous, with pilose and crenate sides. Among parasites of the Rallidae this descrip-
tion could only apply to the genus Incidifrons, but as we have not seen this genus
from any bird that complies with the conditions we have mentioned as necessary
assumptions we are unable to erect a neotype for the species. We think it of the first
importance that Incidifrons ralli (Scopoli) should eventually be fixed in such a way
that Incidifrons ralli (Denny) becomes a synonym as well as a homonym, thus avoid-
ing the confusion that would be caused by the transfer of the name valli from one
species to another. We therefore intend to assume in all future work that the host
of Incidifrons ralli (Scopoli), like that of I. ralli (Denny), was Rallus a. aquaticus
Linn., and we most strongly urge other workers to make the same assumption. There
are no specimens of J. valli in the Denny collection.
Pediculus fringillae (p. 125)
The entire description is that the head is bidentate and the sides of the abdomen
are pilose and rugose, but among parasites of the birds included in Fringilla in
Scopoli’s time only Philopterus fits this description. There
al is no host-record other than that provided by the name,
so we have chosen as host of our neotype one of the
commoner birds included by Scopoli in Fringilla, namely,
Fringilla domestica, now known as Passer domesticus
Linn.?
We hope that our action will finally settle the confusion
that has arisen (as shown in Part I of this work (Clay
and Hopkins, 1950: 270)) through the application of
Geoffroy’s invalid appellation ‘ subflavescens’ to the Philop-
terus from this host. Fourcroy (1785: 519) gave the name
Pediculus passeris to the species described by Geoffroy,
and Philopterus passeris (Fourcroy) and P. passeris
(Piaget) (together with subflavescens of authors subsequent
to Geoffroy) become synonyms of Philopterus fringillae
Fic. 43. Philopterus frin- .
gillae (Scopoli), male geni- (Scopoh). : :
talia. x 3109. This species has been discussed above under P. coarctatus
(Scopoli) 1763, from which it differs in having a median
indentation in the hyaline margin of the head, and in the characters of the male
genitalia and female genital plate.
Neotype male (Fig. 43 ; Pl. III, fig. 3 ; Tables r and 3) and neallotype female (Fig. 44;
Tables 2 and 4-5) of Philopterus fringillae (Scopoli) from Passer d. domesticus (Linn.)
‘This host is not given in the publication under discussion, in which no species of Fringilla are men-
tioned, but in Part I of the same work, published in 1769, Scopoli refers to Fringilla domestica on p. 149.
THE EARLY LITERATURE ON MALLOPHAGA 33
from Hungary (Meinertzhagen collection, slide No. 8077a). Neoparatypes: 47 males
and 57 females from the same host-form, Hungary, Estonia, and the British Isles.
Measurements in mm.
Male Female
Length Breadth Length Breadth
Head - 4 ; 0°55 0°50 0:58 0°56
Prothorax. i 0:29 ae 0:32
Pterothorax ar 0°45 aie 0°50
Abdomen . : é 0-70 0:68 I-02 o-g1
Total ° . . 1°45 Ar 1°85
Genitalia . . A 0°31 Be
Fic. 44. Philopterus fringillae (Scopoli), genital plates of 4 females
taken from one host individual. x 102.
Pediculus part palustris (p. 125)
‘Cauda quadriseta, ut in P. Pari majoris Linn.’ As no differences from Linné’s
species are mentioned we consider this to be a nomen nudum, but in any case the
name clearly does not refer to the Mallophaga but to a mite.
J. C. FaBRIcIUS, 1775 (Systema Entomologiae. Flensburgi et Lipsiae: 804-810)
The great majority of the names mentioned in this work have already been dealt
with, being either quoted from Linné or entirely unnecessary renamings of Scopoli’s
species, with descriptions quoted from the latter author. Only the following are
genuinely new:
Pediculus vulturts (p. 806)
The description is quite obviously that of a Laemobothrion, but the host-record is
merely ‘Habitat in Indiae orientalis vulturibus’. In these circumstances the most
reasonable procedure seems to be to attach the name to a Laemobothrion from one of
the Indian vultures. We have chosen Pseudogyps bengalensis (Gmelin).
This species differs from L. tinnunculi (Linn.) (see Clay and Hopkins, 1950: 228)
34 THE EARLY LITERATURE ON MALLOPHAGA
in the larger size, the shape of the head (PI. III, figs. 4, 6), in the greater number of
prosternal setae’ and the shape of the sternal plates (Fig. 45), the presence of a line of
setae on the lateral margins of the sternal plates, and in details of the
— male genitalia. No material of Laemobothrion maximum (Scopoli) has
been seen from Buteo buteo, but from Eichler’s figures (1941: 363, fig. 28 ;
1942: 59, fig. 4), and examination of specimens (possibly not conspecific
with maximum) from other species of Buteo, it seems to differ from
vulturis in the smaller size, the smaller number of prosternal setae,
the shape of the prosternal plate, and possibly in the details of the male
genitalia.
Measurements in mm.
Male Female
Length Breadth Length Breadth
Head* I+52 1°70 1°56 1:80
Abdomen . 5°76 2:48 6:92 3:02
Total 9°24 ks 10°56
Genitalia . 2°90 fc
FIG. 45.
Laemoboth-
* The head is liable to distortion in mounted specimens of Laemobothrion and
rion vulturis throughout the genus these measurements may show considerable variation in one
(J. C. Fabri- species.
cius), female
sternal Neotype male (Pl. III, figs. 4-5) and neallotype female (Fig. 45; Pl.
Plates. x31. ITT, fig. 6) of Laemobothrion vulturis (J. C. Fabricius) from Pseudogyps
bengalensis (Gmelin) from Deccan, India (Meinertzhagen collection, slide No. 8607).
Neoparatypes: 1 male and 2 females from the same host-form, Deccan and Siam.
Pediculus procellariae (p. 808)
An elongate, filiform, fuscous species occurring ‘in Brasiliae procellariis’ can only
be a Halipeurus or a Perineus, the former being the more probable identification
because members of this genus are more elongate than those of Pevineus (Fabricius
seems to have been particularly impressed by this character) and also more fuscous.
But in view of the number of species of petrels which occur in Brazilian waters the
selection of host must be purely arbitrary. Mr. R. L. Edwards informs us that in a
paper he is about to publish he will redescribe Halipeurus procellariae (J. C. Fabricius)
from specimens taken from Pterodroma m. macroptera (A. Smith). Halipeurus con-
strictiventris (Pess6a and Guimaraes) 1935 will thus become a synonym of H.
procellariae.
Pediculus diomedeae (p. 808)
One of us (Clay, 1940: 300-302) has already discussed this species and erected
neotypes from Diomedea m. melanophris Temminck. It is perhaps as well to repeat
that it is a Perineus, not a Harrisoniella, has nothing to do with Harrisontella ferox
(Giebel), and that the type species of Harrisoniella is Esthiopterum diomedeae Harrison
1916 (nec J. C. Fabricius 1775) (= Lipeurus ferox Giebel).
t Examination of further material suggests that this is an unreliable character owing to individual
variation. See Part III (in press) for further notes on Laemobothrion vulturis.
THE EARLY LITERATURE ON MALLOPHAGA 35
Pediculus hirundinis (p. 810)
We have noted above (Clay and Hopkins, 1950: 26) that this name must be ascribed
to Linné and have discussed it under that author.
TABLES 1-5, MEASUREMENTS OF PHILOPTERUS SPECIES
TABLE 1. Breadth (in mm.) of head at temples of males, with number of specimens
O°45— 0°47— 0-49- O*5I- O:53- 0°55=
0-46 0-48 0:50 0°52 0°54 0:56
coarctatus : : : os 6 17 4 Sr ye
fringillae : Z : Af a 3 15 7 2
citrinellae A f : 3 22 2 ae : ;
TABLE 2. Breadth (in mm.) of head at temples of females, with number of specimens
o-47— | o'4g- | o-sz- | 0-53- | o-55- | 0-57- | 0-59- | o-6r-
0:48 0°50 O52 | 0°54 0:56 0°58 0:60 0:62
coarctatus A 7 : ae : 4 18 ate ie
fringillae A ; 6 13 4 5
citrinellae : , : I 4 as ws ois
TABLE 3. Cephalic index of males, with number of specimens
0-90- 0-92-— 0:94- 0:96- 0-98— I-00- I:02-—
o-9r 0:93 0:95 0:97 0-99 I-or I'03
coarctatus ; ‘ 2 3 5 fe) 7 7 oe ue 3
fringillae é : : = 2 5 Io 3 4
citrinellae : : : 4 10 9 3 I
TABLE 4. Cephalic index of females, with number of specimens
0-90— | 0-92— | o-g4- | 0:96— | o-g8— | I:oo—- | T-02- | rIo4-
0-91 0°93 0°95 0:97 0°99 I-or I-03 I-05
coarctatus : 5 2 3 5 II 4 6 I ate as
fringillae ‘ ; ; i I 4 7 7 7 3 I
citrinellae . ; , 2 8 II 7 2 ae o ee
TABLE 5. Ratio of breadth: length of female genital plate, with number of specimens in
parentheses
coarctatus . . i . 1°38 (1), 1°45 (1), 1°53 (2), 1°54 (I), 1°55 (1), 1°58 (x),
1°59 (1), 169 (2), 1°75 (I).
fringillae . - 4 - 1°79 (1), 1°88 (3), I-o1 (1), 194 (2), 2:00 (3), 2°10 (1).
List OF SPECIES
The synonymy of the following names has been established.
Specific name Present status Page
albiventris Scopoli. Penenirmus albiventris (Scopoli). 28
anatis Fabricius. Anaticola crassicornis (Scopoli). 17
36 THE EARLY LITERATURE ON MALLOPHAGA
Specific name
arvdealis Scopoli.
auritus Scopoli.
bidentatus Scopoli.
biseriatis Denny.
buteonis Fabricius.
coarctatus Scopoli.
collurionis Schrank.
colymbinus Scopoli.
constrictiventris Pesséa &
Guimaraes.
cornicis Fabricius.
crassicornis Scopoli.
cucult Fabricius.
cuspidatus Scopoli.
dentatus Scopoli.
diomedeae Fabricius.
dolichocephalus Scopoli.
fasciatus Scopoli.
fringillae Scopoli.
fusco-marginatus Denny.
gigantum Nitzsch.
haematopus Scopoli.
junceus Scopoli.
lanii Fabricius.
maximus Scopoli.
motacillae Fabricius.
ocellatus Scopoli.
oriolt Fabricius.
ovalis Scopoli.
passeris Fourcroy.
passeris Piaget.
phanerostigma Giebel.
phanerostigmaton Nitzsch.
pilosus Scopoli.
platyrhynchus Nitzsch.
procellariae Fabricius.
pseudonirmus Nitzsch.
valli Scopoli.
valli Denny.
squalidus Nitzsch.
strigis Pontoppidan.
strigis Scopoli.
strigis Fabricius.
subflavescens auctorum.
sulphureum Nitzsch.
superciliosus Burmeister.
testudinarius Denny.
thompsoni Eichler.
tinnuncult Latreille.
tvidens Burmeister.
tridens var. major Piaget.
troglodytis Waterston.
vanelli Schrank.
vulturis Fabricius.
Present status
Ardeicola ciconiae (Linn.).
Peneniymus auritus (Scopoli).
Campanulotes bidentatus (Scopoli).
Cummingsiella ovalis (Scopoli).
Laemobothrion maximum (Scopoli).
Philopterus coarctatus (Scopoli).
Philopterus coarctatus (Scopoli).
Aquanirmus colymbinus (Scopoli).
Halipeurus procellariae (Fabricius).
Philopterus ocellatus (Scopoli).
Anaticola crassicornis (Scopoli).
Cuculiphilus fasciatus (Scopoli).
Rallicola cuspidatus (Scopoli).
Anatoecus dentatus (Scopoli).
Perineus diomedeae (Fabricius).
Ricinus dolichocephalus (Scopoli).
Cuculiphilus fasciatus (Scopoli).
Philopterus fringillae (Scopoli).
Aquanirmus colymbinus (Scopoli).
Laemobothrion maximum (Scopoli).
Craspedorrhynchus haematopus (Scopoli).
Quadraceps junceus (Scopoli).
Philopterus coarctatus (Scopoli).
Laemobothrion maximum (Scopoli).
Penenirmus albiventris (Scopoli).
Philopterus ocellatus (Scopoli).
Ricinus dolichocephalus (Scopoli).
Cummingsiella ovalis (Scopoli).
Philopterus fringillae (Scopoli).
Philopterus fringillae (Scopoli).
Cuculiphilus fasciatus (Scopoli).
Cuculiphilus fasciatus (Scopoli).
Pseudomenopon pilosum (Scopoli).
Craspedorrhynchus haematopus (Scopoli).
Halipeurus procellariae (Fabricius).
Cummingsiella ovalis (Scopoli).
Incidifrons valli (Scopoli).
Incidifrons valli (Scopoli).
Anaticola crassicornis (Scopoli).
Strigiphilus strigis (Pontoppidan).
Strigiphilus strigis (Pontoppidan).
Craspedorrhynchus haematopus (Scopoli).
Philopterus fringillae (Scopol).
Ricinus dolichocephalus (Scopoli).
Penenirmus auritus (Scopoli).
Cummingsiella ovalis (Scopoli).
Pseudomenopon pilosum (Scopoli).
Craspedorrhynchus haematopus (Scopoli).
Pseudomenopon pilosum (Scopoli).
Pseudomenopon pilosum (Scopoli).
Penenirmus albiventris (Scopoli).
Quadraceps junceus (Scopoli).
Laemobothrion vulturis (Fabricius).
THE EARLY LITERATURE ON MALLOPHAGA 37
REFERENCES
Only those papers not referred to in Part I are listed here.
Cray, T. 1940. Anoplura. British Graham Land Exped. Sci. Rep. (Brit. Mus. (Nat. Hist.)),
1: 295-318.
—— & Hopkins, G. H. E. 1950. The Early Literature on Mallophaga. Pt. I. Bull. Brit. Mus.
(Nat. Hist.) Ent. 1 (3): 223-272.
EICHLER, W. 1941. Zur Klassifikation der Lauskerfe. Arch. Naturgesch. B (N.F.), 10: 345-398.
1942. Mallophagen-Synopsis.—III. Genus Laemobothrion. Zool. Anz. 187: 52-63.
—— 1946. Mallophagen-Synopsis.—VIII. Genus Anatoecus. Tijdschr. Ent. 87: 74-70.
1949. Die Eulenfederlinge. Beitr. Taxon. Zool. 1: 7-22.
KE&LER, S. 1938. Baustoffe zu einer Monographie der Mallophagen.—I. Teil. Nova Acta Leop.
Carol. (N.F.), 5: 385-467.
1939. Baustoffe zu einer Monographie der Mallophagen.—II. Teil. Nova Acta Leop. Carol.
(N.F.), 8: 1-254.
1941. Systematisches Verzeichnis der von Nitzsch begriindeten und von Giebel und
Taschenberg fortgefiihrten und bereicherten Sammlung von Mallophagen des Zoologischen
Instituts der Universitat Halle. Z. Naturw. 95: 123-136.
Merisuo, A. K. 1945. Notulae Mallophagologicae.—II. Ann. Ent. Fenn. 11: 101-112.
WATERSTON, J. 1915. On some Mallophaga in the Kgl. Zool. Museum, K6nigsberg. Zool. Jb.
(Abt. Syst.), 89: 17-42.
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PLATE 1
Philopterus coarctatus (Scopoli), 2.
Philopterus ocellatus (Scopoli), 3.
. Ricinus dolichocephalus (Scopoli), @.
Penenirmus auritus (Scopoli), 3.
Anatoecus dentatus (Scopoli), 3.
Bull. B.M. (N.H.) Ent. II, 1 PLATE 1
Fic.
Fic.
Fic.
Fic.
Fic.
Fic.
Po Se ee
PLATE 2
Aquanirmus. colymbinus (Scopoli),
Aquanirmus colymbinus (Scopoli),
Quadraceps junceus (Scopoli), ¢.
Quadraceps junceus (Scopoli), 2.
Rallicola cuspidatus (Scopoli), 3.
Rallicola cuspidatus (Scopoli), 9.
3.
St
Bull. B.M. (N.H.) Ent. II, 1 ' PLATE 2
ves"
yeOUTT
nN
Fic.
Fic.
Fic.
Fic.
Fic.
Fia.
AAW H
PLATE:3
Campanulotes bidentatus (Scopoli), 3.
Strigiphilus strigis (Pontoppidan), ¢.
Philopterus fringillae (Scopoli), 3.
Laemobothrion vulturis (J. C. Fabricius), 3.
Laemobothrion vulturis (J. C. Fabricius), ¢ genitalia.
Laemobothrion vulturis (J. C. Fabricius), 9.
PLATE 3
Bull. B.M. (N.H.) Ent. II, 1
PRESE NTED
_
4
PRINTED IN _ eis
GREAT BRITAIN © aie E
UNIVERSITY PRESS : | i,
OXFORD \ Me aa
a
CHARLES BATEY
PRINTER
TO THE
UNIVERSITY
17 DEC 02
REVISION OF
THE AUSTRALIAN AND
TASMANIAN
GRIPOPTERYGIDAE
AND NEMOURIDAE
(PLECOPTERA)
D. E. KIMMINS
BULLETIN OF
_ THE BRITISH MUSEUM (NATURAL HISTORY)
_ ENTOMOLOGY Vol. 2 No. 2
| LONDON : 1951
A REVISION OF THE AUSTRALIAN AND
TASMANIAN GRIPOPTERYGIDAE AND
NEMOURIDAE (PLECOPTERA)
BY
D. E. KIMMINS
Tee!
Pp. 43-93; 40 Text-figures
| BULLETIN: OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 2 No. 2
LONDON : 1951
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, 1s issued
in five series, corresponding to the Departments of the
Museum.
Parts appear at irregular intervals as they become
ready. Volumes will contain about three or four hundred
pages, and will not necessarily be completed within
one calendar year.
This paper ts Vol. 2, No. 2, of the Entomology series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued December 1951 Price Ten shillings
A REVISION OF THE AUSTRALIAN AND
TASMANIAN GRIPOPTERYGIDAE AND
NEMOURIDAE (PLECOPTERA)
By D. E. KIMMINS
SYNOPSIS
In this paper sixteen new species of the family Gripopterygidae and four in the family Nemouridae are
described. The Australian species of Nemouridae are transferred to the subfamily Notonemourinae Ricker,
containing nine genera with a distribution ranging over Australia, Tasmania, New Zealand, southern South
America, and Cape Province, South Africa. The type of the genus Leptoperla Newman has been studied
and redescribed, with the result that a later genus described by Enderlein must fall into synonymy.
In addition, the nymphs of three genera of the Gripopterygidae are described and figured.
THE Gripopterygidae are the dominant Plecopterous family in Australia and New
Zealand ; Tillyard gave ten as the number of Australian and Tasmanian species, a
figure which included some undescribed species in his collection, as only eight species
had then been published, contained in five genera. Examination of the British
Museum accessions and the Tillyard collection has enabled this list to be increased by
sixteen new species and the genera to be reduced to four.! In addition, it has been
possible to describe the nymphs of three of the genera, and it is highly probable that
there are still more species to be described, as more than one species of Tvinotoperla
nymph has been recognized from Tasmania, whence no adult has yet been recorded.
Apart from Australia, Tasmania, and New Zealand, this family also occurs in the
Fiji Islands, Auckland Islands, and the colder, mountainous parts of South America.
The Nemouridae are represented in Australia and Tasmania by five species (four
here described for the first time), in two genera. For these insects Dr. Ricker’s sub-
family name Notonemourinae isadopted. This subfamily is not restricted to Australiaand
New Zealand, but occurs also in southern South America and in the mountain ranges
of Cape Province, South Africa. In the latter region they are the dominant Plecoptera,
seventeen species in four genera being listed by Barnard, the only other Plecoptera
recorded being two sub-tropical Perlidae. Further collecting in Australia should pro-
duce more species, their small size and dull colouring no doubt causing them to be
overlooked, but according to Tillyard they are decidedly less common than the
Gripopterygidae.
It is difficult to understand why this nemourid subfamily should have established
itself so securely in the mountains of Cape Province, whereas the Gripopterygidae,
which occur with it in other regions, are apparently absent. This absence would not
seem to be due to lack of collecting, as Tillyard asked Dr. Barnard to make a special
search in this region in the hope of finding examples of Gripopterygidae. Fossil
records may show whether this family did ever exist in the Cape, but if they did, they
appear to have died out. The South African Notonemourinae would appear to have
been isolated for a longer period than the Australian and South American forms, as
* Unless otherwise stated, types of the new species are in the British Museum (Nat. Hist.).
46 A REVISION OF THE AUSTRALIAN AND TASMANIAN
all the four South African genera are absent from the other regions. Udamocercia
occurs in South America and Tasmania, two genera are known only from New Zealand,
and Spaniocerca occurs in Australia, Tasmania, and New Zealand.
GRIPOPTERYGIDAE Enderlein
Gripopterygidae Enderlein, 1909, Zool. Anz. 84: 388.
Leptoperlinit Banks, 1913, Trans. Amer. Ent. Soc. 89: 202.
Leptoperlidae Tillyard, 1921, Canad. Ent. 58: 36; 1921, Trans. Roy. Soc. S. Aust. 45: 270-273;
1923, Tvans. N. Zeal. Inst. §4: 202-203; 1926, Ins. Aust. & N.Z.: 119.
Tillyard has adopted for this family the name Leptoperlidae, but the reasons which
he has set out in its support are, unfortunately, based upon a misunderstanding of an
early paper by Newman. Tillyard (1921) writes: ‘This family was first recognised as
a distinct group by Newman in 1839, and has been raised to tribal rank by Banks
under the name Leptoperlini, these authors not considering the whole of the Perlaria
to be entitled to more than family rank. Now that the ordinal rank of the Perlaria is
well established, the various tribes of Banks take family rank.’
In the first place, Newman merely erected a new genus, Leptoperla, with one species,
beroé, and his.introduction makes it clear that it was a genus of the family Perlidae.
No mention is made that it belonged to a ‘distinct group’. Secondly, Newman did
not consider the whole of the Plecoptera to belong to one family. He heads his paper
‘Class—NEUROPTERA. Natural Order—PERLITES. Family—PERLIDAE’, and in his
first paragraph he states that he uses the family Perlidae in a restricted sense, ‘includ-
ing only those genera which are furnished with caudal setae: Leach and Stephens
incorporate with them the ecaudate Nemourae’. Thus it is evident that he recognized
at least two families in what are now called Plecoptera.
The first worker to give family rank to this section of the Plecoptera was Ender-
lein, who in 1909 proposed the name Gripopterygidae ; his family was a rather com-
posite one, including the genera Paragripopteryx Enderlein, Gripopteryx Pictet,
Eusthenia Gray, Stenoperla McLachlan, Antarctoperla Enderlein, Notoperla Enderlein,
and Paranotoperla Enderlein, but omitting Leptoperla Newman. In 1913, under the
tribal name Leptoperlini (sub-family Pteronarcinae), Banks groups Austroperla
Needham, Gripopteryx Pictet, Leptoperla Newman, Notoperla Enderlein, Auck-
landobius Enderlein, Antarctoperla Enderlein, and Paranotoperla Enderlein. Tillyard
in 1921 raised Bank’s tribe to family status and at the same time separated off
Austroperla to form a distinct family.
Leptoperla Newman undeniably belongs to the same family as Gripopteryx Pictet,
Paranotoperla Enderlein being actually a synonym of Leptoperla, and thus the name
Leptoperlidae must give place to Gripopterygidae on grounds of priority.
Diagnosis of family
Fore wing with Sc terminating at from one-half to two-thirds of the wing length
from the base, thus leaving a long pterostigmatic area, with or without cross-veins.
Cross-veins, more or less numerous, are present in the distal half of the wing, and
there is no transverse cord. Three anal veins present, first long, second and third
shorter and fused at their bases. On posterior margin, between the terminations of
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 47
Cur and Cuz, the marginal fringe of hairs is modified into a row of hooked hairs,
probably a form of wing-coupling. I have not seen this structure in any other family
of Plecoptera, nor have I seen any previous record of it. In the hind wing, M3+4 is
always fused with Cur shortly after it separates from M1-+2; it may remain thus
fused with Cur to the margin of the wing, or it may separate again, thus appearing
like a fork of Cur. Anal fan always forming an angle with the hind margin of the
wing, and containing only six simple veins, including 1A; it is generally without
cross-veins.
The nymphs may be readily distinguished by the tuft or rosette of filiform anal
gills, whitish, pinkish, or pale mauve in colour. According to Tillyard, these nymphs
are sluggish in habits, being found clinging to rocks in the swifter parts of fast-
running streams. In the more mature nymphs the developing wing pads show the
main venation of the adults quite clearly, and by this means it has been possible to
recognize the nymphs of the three genera Tvinotoperla, Dinotoperla, and Leptoperla.
KEY TO GENERA OF GRIPOPTERYGIDAE OF AUSTRALIA, TASMANIA,
NEW ZEALAND, AND AUCKLAND ISLANDS
I. Rs forked in both pairs of wings
Rs simple in both pairs of wings
2. Cur in fore wing forked .
Cur in fore wing simple :
3. A series of cross-veins between Cuz and IA in fore wing
EUNOTOPERLA Tillyard, p. 47
hwWUN
Not more than one cross-vein in this area. . TRINOTOPERLA Tillyard, p. 75
4. Fork of Rslong . ; ; ; : MEGALEPTOPERLA Tillyard
Fork of Rs terminal : 7 ; . ZELANDOBIUS Tillyard
5. M3+4 and Cur in hind wing separate at base and apex, fused in middle of wing
DINOTOPERLA Tillyard, p. 62
M3+4 and Cur acs at base of hind wing, fused from centre of wing to
margin ; : 2.6
6. Generally only one cross-vein between Cur and Cuz i in hind wing
LEPTOPERLA Newman, p. 48
More than two or three cross-veins in this area . ; ; : : rae |
7. A series of cross-veins in the pterostigma in both wings
ZELANDOPERLA Tillyard
No cross-veins in the pterostigma_ . ; é : ; . : : 3
8. Fore wings shorter than the hind wings . AUCKLANDOBIUS Enderlein
Fore wings longer than hind wings . : ‘ NESOPERLA Tillyard
EUNOTOPERLA Tillyard
Eunotoperla Tillyard, 1924, Trans. Roy. Soc. S. Aust. 48: 194.
Type species: E. kershawi Tillyard, 1924 (fixed by Tillyard, 1924).
_ This genus is so far not represented in the British Museum (Nat. Hist.) and I am
therefore quoting Tillyard’s descriptions.
t
48 A REVISION OF THE AUSTRALIAN AND TASMANIAN
‘Fore wing with Rs distally forked or three-branched, M forked from near middle
of wing, Cur three-branched, or more rarely, simply forked ; a thickened cross-vein
between 1A and 24; cells between M and Cu partially double ; cubito-anal space with
cross-veins. Hind wing with Rs terminally forked or simple, M forked, Cur forked or
simple ; apex well rounded; anal fan moderately wide, with a few weak cross-veins
developed in the region of 1A and 2A. Cerci short. Size of species large, expanding
50 mm. or more.
‘This genus shows an approach to the Eustheniidae in the beginning of the develop-
ment of cross-veins on the anal fan, and is in other respects somewhat similar to the
genus Stenoperla of that family ; it can be at once distinguished by the marked angle
between the border of the anal fan and the rest of the hind wing, this angle being
entirely absent in the Eustheniidae.’
Distribution: AUSTRALIA, Victoria.
Eunotoperla kershawi Tillyard
Fic. I
Eunotoperla kershawi Tillyard, 1924, Trans. Roy. Soc. S. Aust. 48: 195, fig. 2; 1926, Ins. Aust. &
N Zot 319; pl 20, fig. 16;
‘g. Length of body (abdomen much shrunken): 11 mm., of fore wing 25 mm.,
antennae 16 mm., cerci 4mm. Body, legs, and antennae blackish, with front border of
foe F
NEES CURT
SET a ao
Fic. 1. Eunotoperla kershawi Tillyard. Right fore wing (after Tillyard).
pronotum brownish. Wings dull brownish with dark brown venation; cross-veins nearly
all enclosed in pale transparent whitish areas; thickened cross-vein between IA and
2A of fore wing dark brown; supra-anal plate with a copulatory process in the form
of a slender, downcurved spine; paraprocts with a pair of shorter, upcurved spines.
‘Q. Length of body (not shrunken) : 25 mm., fore wing 30 mm. ; differs from male in
having abdomen and pronotum brown, wings dark brown, the hind wings somewhat
fuscous.
‘Types: Holotype male and allotype female (Warburton, Victoria, 12.94) in
National Museum, Melbourne; paratype male, from same locality, in Cawthron
Institute collection, Nelson, N.Z., also a male from Thorpdale, Gippsland.’
LEPTOPERLA Newman
Leptoperia Newman, 1839, Mag. Nat. Hist. 8 (n.s.): 89.
Type species: L. beroé Newman 1839 (only species included by Newman).
Walker, 1852, Cat. Ins. B.M.1: 169; Banks, 1913, Trans. Amer. Ent. Soc. 39: 203 ; Tillyard, 1921,
Canad. Ent. 58: 42, fig. 4a; 1921, Trans. Roy. Soc. S. Aust. 45: 273.
Paranotoperla Enderlein, 1909, Zool. Anz. 34: 393, 416 (Syn. nov.)
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 49
Type species: P. australica Enderlein (fixed by Enderlein, 1909).
Enderlein, 1909, Disch. Ent. Z. 1909: 634, fig. 2; Tillyard, 1921, Trans. Roy. Soc. S. Aust. 45: 273.
Newman’s description of the genus is very brief. ‘Proalarum nervurae apicales 6,
nervuris transversis numerosis intersectae: antennae setaeque caudales elongatae ;
pedes elongati.’
The type of L. beroé is gummed on a card and the wings are somewhat crumpled.
Tillyard examined the specimen in 1920 and gave an extended generic diagnosis in
the Canadian Entomologist. For some reason he did not completely spread out the
left fore wing when he softened it forexamination, and consequently the fact that the
right fore wing (with Rs forked almost to its origin) was abnormal escaped his notice.
Through the courtesy of Professor G. Hale Carpenter I have been able to study,
amongst other Australian material, the type of L. beroé. Included in this material
was an example resembling beroé, but without the forked Rs in the fore wing. This
Fic. 2. Leptoperla beroé Newman, g type. a, right fore wing (after Tillyard, showing aberrant
fork of Rs) ; b, left fore wing (original).
aroused my suspicions and I removed and spread out the left fore wing of the type.
In this, Rs was simple and the whole venation was of the pattern of Paranotoperla
Enderlein (Fig. 25). Genitalia and cerci also agree with that genus, and I have no
hesitation in placing Enderlein’s genus in the synonymy of Leptoperla.
The following is my revised diagnosis of the genus. Fore wing with Rs simple,
M forked near the middle of the wing, Cur simple; cubito-anal space with only one
cross-vein: a thickened cross-vein between IA and 2A. Hind wing with Rs simple,
Rs and M generally separating close to the base of the wing ; in L. rugosa, L. reticulata,
and L. nigrifrons they remain fused for a much greater distance ; posterior branch of
M fused with Cur from near the fork of M to the wing margin. Generally only one
cross-vein between the branches of Cu. Anal fan rather narrow. Cerci long.
Distribution: Australia, Tasmania, Fiji Islands.
Leptoperla beroe Newman
FIGS. 2, 3
Leptoperla beroé Newman, 1839, Mag. Nat. Hist. 3 (n.s.): 89-90; Walker, 1852, Cat. Neur. Ins.
B.M.1: 169; Banks, 1913, Trans. Amer. Ent. Soc. 89: 203; Tillyard, 1921, Canad. Ent. 58: 42,
fig. 4a; 1921, Trans. Roy. Soc. S. Aust. 45: 273; 1926, Ins. Aust. & N.Z.: 119.
Newman’s description (given below) is good, but owing to the aberrant fore wing,
his statement that there are s7x parallel veins in the fore wing should be read as five.
‘Sp. 1. Leptoperla Beroé. Fusca, alae opacae, fuscae, versus apicem maculis
50 A REVISION OF THE AUSTRALIAN AND TASMANIAN
albidis notatae ; pro- et mesopedes fusci, tibiis medio testaceis: metafemora testacea,
apice fusca. (Corp. long. -3 unc. ant. -475 unc. set. caud. -475 unc. alar. dilat. -9 unc.)
‘This is a slender and very elegant insect, it differs generically from Isogenus and
Perla in the neuration of the fore wings, the exterior portion of which is occupied by
six strong parallel longitudinal nervures ; of these the fourth is furcate at the extremity
and the fifth unites with the fourth just before its furcation;! these longitudinal
nervures are intersected by several very delicate transverse nervures: the antennae
and caudal setae are very slender, and much longer than the body of the insect ; the
terminal segment of the abdomen is furnished below with two leaf-like processes,
which curve upwards, passing between
the caudal setae and terminating in acute
points: the legs are very long and slen-
der: the insect is of a dark brown colour,
the wings being opaque, dark brown,
and the exterior portion of the fore
wings regularly spotted with dirty
white ; the hind wings are immaculate ;
the pro- and mesofemora having a
bright testaceous ring; the metafemora
are testaceous, with the apex only dark
brown, the ¢biae are rather paler, and
the tarsi nearly black.
‘Inhabits Van Diemens Land. There
is a single specimen in the cabinet of
the Rev. F. W. Hope.’ (Now in the
Hope Department, University Museum,
Oxford.)
To this description may be added the
a following account of the male genitalia.
F =—— —_——_—sqAAApicali tergites pigmented at bases and
Fic. 3. rence ee sale mee SAE erase sides, sternites with a pair of basal spots.
oy at Tenth tergite large, narrowed apically,
apical margin truncate from the side and carrying a short triangular process. There
is a low median dorsal carina, and at the centre of the basal margin is a small quad-
rate, membranous area. Supra-anal lobe slender, not projecting much beyond the
margin of the tergite. From the side there is a dorsal excision before the apex.
giving the appearance of a crochet-hook, and several acute dorsal teeth. Sub-anal
plates large, broad, leaf-like, curving upwards, with somewhat twisted apices ; lower
basal margins heavily fringed with hairs. Cerci long, at least thirty-seven-segmented.
Subgenital plate triangularly produced, pigmented at its centre, giving the im-
pression of being narrower.
The above genitalic description has been made from a second example in the Hope
Department, University Museum, Oxford.
1 Tillyard has shown this to be an aberration; in the left fore wing all these veins run parallel.
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 51
Leptoperla australica (Enderlein)
Fic. 4
Paranotoperla australica Enderlein, 1909, Zool. Anz. 84: 416; 1909, Dtsch. Ent. Z. 1909: 684;
1912, Fauna SW. Aust.: 60; Tillyard, 1921, Trans. Roy. Soc. S. Aust. 45: 270; ? SAmal, 1921,
Casopis, 18: 66-70.
Sdmal’s reference to the occurrence of this species in Tasmania is probably based
on a mis-determination and may possibly refer to L. beroé. I have not seen any
examples of L. australica from the island and I believe it to be restricted to west and
south-west Australia.
— mente / a a =
Se
Fic. 4. Leptoperla australica (Enderlein) 9. a, wings; b, genitalia, ventral.
Head brownish-yellow, darker above. Eyes black. Thorax brown, almost rect-
angular, somewhat longer than broad, front and hind margins straight, lateral margins
straight and diverging slightly from front to back ; no impressed median line, but with
anterior and posterior transverse furrows ; anterior angles rounded. Abdomen light
brown, in the female paler beneath, subgenital plate brown. Cerci dark brown,
longer than abdomen, about thirty segments. Legs brownish-yellow, marked with
dark brown at the knees, apices of tibiae, and on the tarsi. Wings light brown, with
brown venation, distal cross-veins pale and enclosed in whitish spaces; generally
without cross-veins between Cur and wing margin in fore wing.
I have seen no males; Enderlein says that the last tergite in this sex is triangular.
His description of the subgenital plate apparently refers to the female.
?. Subgenital plate large, quadrate, angles rounded, projecting somewhat beyond
the segment, brown, with pale triangular excised areas, a broad shallow area at apex
and a narrow deep one at base. Sub-anal plates triangular, apices curving outward.
Length of fore wing g 9°-5-I0 mm., 2 7-5-9°5 mm.
Length of hind wing ¢ ?, 2 6-5-8-5 mm.
SW. AvusTRALIA: Lunenberg; 22.ix.1905, Serpentine, 23-25.ix.1905, Harvey,
27.vii.1905 (Hamburg SW. Australia Expedition). Wrst AusTRALIA: Bolgonup,
I.x.1922 (R. J. Tillyard).
Type in Stettin Museum.
Leptoperla tasmanica sp.n.
FIG. 5
(In fluid.) Head, thorax, and three terminal segments of abdomen brown, shin-
ing, segments one to seven whitish. Antennae longer than fore wing. Pronotum
ENTOM. 2, 2 F
52 A REVISION OF THE AUSTRALIAN AND TASMANIAN
trapezoidal, anterior margin two-thirds as wide as posterior, a little shorter than width
at base. Legs brownish, distinctly banded with dark brown towards apex of femur and
base of tibia, apex of tibia and tarsus clouded with darker brown. Wings elongate,
fore wing medium fuscous, with pale markings; all cross-veins, and to a lesser degree
Fic. 5. Leptoperla tasmanica sp.n. a, wings, 3; b-d, genitalia 3, left cercus omitted, b, lateral,
c, dorsal, d, ventral; e, genitalia 2, ventral.
also the longitudinal veins in the apical half margined with whitish, so that each
cellule is dark, surrounded by a pale line. In female, cross-veins are more numerous,
and main veins are not bordered with whitish, only the cross-veins. In the basal half
of the wing it is the cells which are pale, bordered with fuscous. Veins pale fuscous,
except apical cross-veins which are whitish (pale purplish in dried examples). Hind
wing uniform pale fuscous, with darker veins.
3g. Ninth tergite with a large unpigmented area on dorsum, extending almost to
base. Tenth tergite with a strong longitudinal ridge, and on each side of it a small
rounded depression, base of segment unpigmented ; apical margin triangularly pro-
duced, apex truncate, bent down from the side, and carrying a ridged projection,
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 53
triangular from above, quadrate from the side, its lower apical angle acute. Supra-
anal lobe broad, triangular, curving upward, its upper surface set with teeth. Cerci
brownish, at least as long as abdomen (incomplete in type), more than twenty-three-
segmented, basal segment pale. Sub-anal plates blunt, curving upward, upper margin
curved over outward. Above them is a small setose lobe. Subgenital plate ovate,
projecting about a quarter of its length beyond the sternite, separated from the seg-
ment by an unpigmented band.
2. Dorsum of eighth and ninth segments membranous, tenth pigmented, triangu-
larly produced and deflexed, terminating in a slender blackened spine; dorsally there
are two small rounded pits similar to those in the male. Subgenital plate broad at
base, narrowed to half its width at apex, which is shallowly emarginate or sometimes
merely truncate. It extends nearly to the apex of ninth sternite, and is brownish,
with a narrow triangular whitish area, the base of the triangle adjoining the basal
margin of the plate. Anal plates triangular, apices produced in flattened fingers.
Cerci brownish, about twice as long as abdomen.
Length of fore wing g¢ 12 mm., 9 12 mm.
Length of hind wing g 10 mm., 9 I0 mm.
TASMANIA: Gouldt County, 10.11.1933, I g; Penstock Lagoon, 27—29.1.1933, 3 2
(R. J. Tillyard) ; Great Lake, ii.1934, 2 2 (Critchley Parker).
Type ¢ (from Gouldt County) and a paratype female from Penstock Lagoon are
microscope preparations, the two females from Great Lake are pinned and the
remaining two females are in 2 per cent. formaldehyde solution.
This species has paler wings than L. australica (Enderlein), and the presence of
cross-veins between Cur and the margin in the fore wing will help to separate it from
that species.
Leptoperla exigua sp.n.
Fic. 6
(In fluid.) Head and thorax brownish, abdomen paler, particularly beneath.
Antennae long, brownish. Pronotum quadrate, about one-fifth broader than long,
anterior angles rounded, anterior and posterior transverse furrows shallow. Legs
yellowish-brown, femora with a darker ring just before apex, which is pale, base of
tibiae also darker. Fore wing with apical cross-veins whitish, outer ones arranged in
an almost straight row across the wing. In hind wing Rs and M separate close to base.
g. Tenth tergite with its apical margin quadrately produced and deflexed ; from
a rounded whitish area arises a slender upcurved process, somewhat ridged dorsally.
Supra-anal lobe long and slender, particularly in apical half, apex hooked downward.
Cerci pale brownish, longer than abdomen, about thirty-two-segmented. Sub-anal
plates upcurved, fitting closely on either side of the supra-anal lobe, apices dilated
and terminating in an acute spine. Subgenital plate rather broad-oval, a small
unpigmented area at its apex giving the appearance of being excised.
2. Sides of the eighth and ninth tergites pigmented, dorsum membranous, tenth
pigmented and triangularly produced. Sub-genital plate large, dark brown, apex shal-
lowly excised, sides narrowly unpigmented, giving the appearance of a constriction
54 A REVISION OF THE AUSTRALIAN AND TASMANIAN
before the apex. Basally on each side is a shallow rounded depression. Sub-anal plates
triangular, outer margins with a deep rounded excision. Cerci pale brownish, about
as long as the abdomen.
Length of fore wing, ¢ 6-5 mm., 9 7 mm.
Length of hind wing, ¢ 5-5 mm., 2? 6 mm.
W. AvusTRALIA: Kelmscott, 22.ix.1932, 2g, 102, Bolgonup, I.x.1932, 4 4, 29,
(R. J. Tillyard) ; Chidlow, near Perth, 14.ix.1923, 3 g, 1 9, Mundaring, near Perth,
15.ix.1923, I 2, (G. A. K. Marshall). TasMANIA: 1 g (J. W. Evans).
Fic. 6. Leptoperla exigua sp.n. a, wings 3; b-c, genitalia g, b, dorso-lateral, c, dorsal ;
d, genitalia 9, ventral.
Type male, paratype female from Kelmscott, paratype male from Chidlow as
microscope preparations, others from Kelmscott and Bolgonup in 2 per cent. form-
aldehyde solution, remainder pinned.
In the single male from Tasmania the arrangement of cross-veins is not quite the
same, but such differences as there are in genitalia are so slight that, in the absence
of more material it is proposed to consider it as L. exigua.
Leptoperla varia sp.n.
FIG. 7
(In fluid.) Head, thorax, and apex of abdomen brownish, remainder of abdomen
whitish. Antennae long, brownish. Pronotum a little longer than broad, slightly
narrowed anteriorly, with two broad transverse furrows, median line not well marked.
Legs yellowish-brown, knees strongly banded with dark brown, apices of tibiae and
tarsal segments brownish. Wings pale fuscous with darker venation. Fore wing
largely pale at base, apical half with cross-veins pale, bordered with whitish, and
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 55
with numerous pale speckles. Hind wing uniform pale fuscous, except costal area
and apical cross-veins, which are whitish. Rs and M separating close to base.
Fic. 7. Leptoperla varia sp.n. g. a, wings; genitalia, b, lateral, c, dorsal, d, ventral.
3g. Ninth tergite membranous dorsally, except at base. Tenth with a short median
ridge, apical margin produced in two rounded lobes. The base of the excision
between them is unpigmented and membranous, and from it arises a slender pro-
cess, slightly dilated apically from side. Supra-anal lobe moderately broad at base,
56 A REVISION OF THE AUSTRALIAN AND TASMANIAN
its apex triangularly dilated and hooked, the two side angles each armed with a pair
of hooks, Cerci brownish, at least as long as abdomen, about forty-five-segmented.
Sub-anal plates broad at base, upcurved, hollowed on their lower surfaces, rather
transparent, apices slightly hooked. Subgenital plate triangularly produced, apex
rounded, a pigmented area down its centre making it appear narrower than it
really is; it extends almost to the apex of the tenth sternite.
2 (in the form of a microscope preparation from the Tillyard collection). Too dis-
torted by pressure in mounting for description.
Length of fore wing ¢ 8 mm., 2 9°75 mm.
Length of hind wing ¢ 7 mm., 2 8 mm.
TASMANIA: Lake St. Clair, 6.ii.1933, 1 g, Scottsdale, 2.xi.1927, I g, 23.1.1933,
Ig,19(R. J. Tillyard).
Type male, Lake St. Clair, male and female paratypes in form of microscope
preparations, one male paratype in 2 per cent. formaldehyde solution.
The speckled wings and bilobed tenth tergite of the male separate this species from
any of the genus known to me.
Leptoperla rugosa sp.n.
Fic. 8
(In fluid.) Head, thorax, and two terminal segments of abdomen brown, remainder
of abdomen whitish, with small trans-
verse brown sclerites, four on each ter-
gite, three on each sternite, two anterior
SSS and a larger posterior one. Antennae
slightly longer than fore wing, apices
of segments more dilated than in éas-
manica. Pronotum one-third broader
than long, slightly narrowed anteriorly,
posterior angles square, anterior
rounded. Surface finely rugose or
tuberculate, with deep anterior and
posterior transverse furrows, linked by
a fine median impressed line. Legs
brownish, knees a little darker.
Wings moderately elongate, pale to
medium fuscous, with darker veins.
Both pairs generally with a narrow pale
streak along apical half of costal area.
Fore wing with three cross-veins below
subcosta and four cross-veins in steps
near the apex of the wing, one cross-
A SON alg ie vein between Cul and margin. In hind
Fac 8 Leplopera rugea spn, wings 3: bf. wing, Rsand Mseparateat some distance
e, genitalia Q, ventral. from base, about half-way to fork of M.
Thy
7 1 ve
yayttt /
afl ely
Ney
vigd
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 57
$. Apical margin of tenth tergite deflexed, and bearing a short process, finger-like
from the side, keeled dorsally. Supra-anal lobe upcurved, of medium width at base,
tapering to a slender apex from above, upper lateral margins set with triangular
teeth, a sinuous hooked projection on the ventral surface directed basally. Cerci
brownish, longer than the abdomen, twenty-seven or more segments. Sub-anal plates
broad, upcurving, apices truncate and setose, upper angles curving over outwards.
Subgenital plate ovate, apex slightly flattened, sometimes slightly excised, extend-
ing beyond the bases of the sub-anal plates.
Q. Sides of segments eight and nine brownish, dorsum membranous; tenth tergite
brownish, margin produced and rounded. Subgenital plate lightly chitinized, about
the width of the segment and scarcely produced, its apex shallowly excised. Sub-anal
plates triangular, apices rounded, not produced in long blades. Cerci brownish, about
twice as long as the abdomen.
Length of fore wing, ¢ 7 mm., 2 7°75 mm.
Length of hind wing, ¢ 6 mm., 2 6-5 mm.
AUSTRALIA: Federal Capital Territory, Lee’s Springs, xi.1932, 6 g, 1 2, Murrum-
bidgee River, 1.1922, 1 g (R. J. Tillyard) ; New South Wales, Bathurst, 2,300 ft.,
10.xi.1884, 1 ¢ (from McLachlan collection).
Type male and one paratype male (from Lee’s Springs) as microscope preparations,
Bathurst male pinned, remainder in 2 per cent. formaldehyde solution.
The arrangement of the outer cross-veins of the fore wing in steps, anterior cross-
veins nearer apex, seems reasonably constant, as does the position of separation of
Rs and M in hind wing. The type and female paratype are rather paler than the
others, probably due to the degree of maturity.
Leptoperla reticulata sp.n.
FIG. 9
¢ unknown.
Q (in fluid). Head yellowish-brown, marked with dark brown between the ocelli
and on the occiput. Antennae brown, about 7 mm. long, a little shorter than the
anterior wing. Pronotum dark fuscous, a little broader than long, narrowed anteriorly ;
a narrow, impressed, piceous median line linking similar anterior and posterior
transverse lines. The latter do not extend more than half the width of the pronotum.
Meso- and metanota shining piceous brown. Legs yellowish-brown, marked with
dark brown at apices of femora, extending along posterior surface, bases and apices
of tibiae and on the tarsi. Abdomen whitish above, except the tenth and sides of
seventh to ninth tergites, which are fuscous, and the pale fuscous sternites of the
first seven segments. Subgenital plate and centre of ninth sternite brown. Sub-anal
plates light fuscous, margined exteriorly with dark brown. Wings rather short, pale
greyish, with fuscous venation ; membrane slightly clouded with brownish in costal
area, around the pterostigma and bordering some of the cross-veins. In the fore wing,
the most noticeable feature is the arrangement of the cross-veins in the apical half,
where they tend to form an irregular network between the main veins. Cross-veins
in apical half of hind wing normal.
58 A REVISION OF THE AUSTRALIAN AND TASMANIAN
Subgenital plate broad, only slightly produced; margin with a shallow excision,
strongly pigmented. Ninth sternite with a small dark brown lobe at its centre,
separated from the remainder of the ster-
TS nite by notches in the apical margin. Sub-
peer BE ae Se anal plates broad, apices stout, blunt and
WY, Ty eo Pp ,» ap out, blunt an
Se as ee , divergent. Cerci about 5mm. long, twenty-
six-segmented, yellowish-brown, monili-
form at base, becoming filiform. Tenth
tergite triangularly produced to a small
rounded apex.
Length of fore wing, 8-5 mm., of hind
wing, 7°25 mm.
AUSTRALIA: New South Wales, Mt.
Kosciusko, Spencer’s Creek, xii.1932, I 2
(R. J. Tillyard).
Type in form of microscope preparation.
This species should be easily separable
from other described species by the irregu-
Fic. 9. Leptoperla reticulata sp.n. 9. a, wings, lar reticulation of the fore wing, and by
b, genitalia, ventral. the lobe of the ninth sternite.
Leptoperla nigrifrons sp.n.
Fic. 10
3 (in fluid). Head dark brown, pitchy-black between epicranial suture and labrum ;
antennae brown, basal segment blackish above. Pronotum brown, with blackish
rugosities, median line, anterior and posterior furrows blackish ; quadrate, parallel-
sided, about as long as broad, anterior margin slightly convex, anterior angles
rounded. Meso- and metanota brown. Legs brown, femora blackish. Wings hyaline,
with considerable pale brownish suffusion as in Fig. 10; venation brownish, apical
cross-veins rather weak. In posterior wing the stalk of Rs-+-M is rather long, there
are two or three cross-veins between Cul and Cuz, and the angle between the posterior
margin and the anal fan is rather abrupt. Abdomen and cerci brownish. Ninth
tergite somewhat excised and membranous at the centre of its apical margin. Ninth
sternite moderately produced in a parallel-sided subgenital plate, whose apex in
ventral view is rounded, with a shallow median excision. Tenth tergite small, dorsally
almost divided into two lobes by excisions of the apical and basal margins. Supra-
anal lobe broad at base, in dorsal view with a finger-like process arising from an
elevated trapezoidal base ; below this the lobe is bent downward and tailward. About
midway on the upper surface is a small basally-directed hook, and the lobe terminates
in a downturned hook. Cerci long, slender, with more than thirty segments (incom-
plete), which become progressively longer and more slender towards the apex. The
second segment is obliquely truncate, causing the cerci to be angled in dorsal view.
Sub-anal plates upcurved in side view, gradually dilating to a truncate apex, with
a pointed upper angle. In ventral view they are slender, with out-turned apices.
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 59
Length of fore wing, 8 mm.
TASMANIA (J. W. Evans), 2 ¢.
Type male in form of microscope preparation, paratype in 2 per cent. formaldehyde
solution.
Fic. 10. Leptoperla nigrifrons sp.n. g. a, wings, showing pattern of fore wing ; b, genitalia,
lateral (left cercus omitted), c, genitalia, dorsal, d, the same, ventral.
The venation of the posterior wing suggests that this species be grouped with
L. rugosa and L. reticulata, but it may be separated from them by the pattern of the
fore wing and the shape of the anal fan.
Leptoperla sp. No. I
FIGS. II-13
Nymph (? about half-grown). General colour yellowish-brown, with obscure,
_ slightly darker markings; of rather slender build (about 7 mm. long), legs rather
_ stout. Body sparsely clothed with short hairs, and in some areas very short, stout
spinules. Head rather wider than the pronotum, brownish, smooth, developing
ocelli visible as black spots beneath the integument. Antennae incomplete, fairly
_ stout, tapering towards apex: first segment large, second smaller, but still larger than
any of the next few succeeding ones; a few scattered setae, but no dorsal fringe.
Labrum about twice as broad as long, anterior angles rounded. Labium with
glossae a little shorter than paraglossae, about the same width, with an apical tuft
of setae ; excision between glossae very narrow. Paraglossae pointed, inner margins
straight, outer rounded. Labial palpus three-segmented, terminal segment large,
- ovate, as long as first and second together. Maxilla well developed: lacinia with three
ENTOM. 2, 2 G
‘fl
60 A REVISION OF THE AUSTRALIAN AND TASMANIAN
or four blunt plate-like teeth at its apex and near them, on its inner margin, a row of
strong acute setae. Galea about as long as lacinia, tapering to a setose apex. Palpus
five-segmented, first, second, and fourth short, second half as long again as the first,
third twice as long as the first, fourth equal to the second, and fifth a little longer than
first and second. Mandibles strongly chitinized, with a number of acute teeth
near apex.
Pronotum about as long as broad, slightly narrowed in front, angles rounded,
anterior and posterior margins both some-
what convex; median impressed line about
half as long as pronotum, anterior and
— posterior impressed lines present, angled
N aah Be. towards each other at their centres. Surface
SS GEG of pronotum smooth. Wing pads small,
anterior elongate, posterior triangular ; vena-
tion typical of the genus. Mesonotum with
an irregular row of small spinules along the
he: » anterior margin and over base of wing pad.
( ee. Posterior margins of meso- and metanota
sion) Ye produced backward, plate-like, a narrow
‘i incision separating their lateral angles from
the wing pads.
: Legs short and strong, yellowish-brown,
femora dorsally with short spinules and setae,
rather deep; tibiae a little longer than femora,
with short acute spinules on inner surfaces,
and a sparse row of fine setae exteriorly.
FYI’ First tarsal segment short, second very short,
Fic. 11. Leptoperla sp. No.1. Nymph. _ third about three times as long as first.
Abdomen cylindrical, tergites two to nine
elevated along the median line, apical margin of each produced in a short spur.
This elevated area and the apical margin bears short spinules as well as hairs,
sternites bearing hairs only. Margin of tenth tergite triangularly produced. Anal
gills dull whitish. Cerci incomplete. Sub-anal plates broad, spatulate.
TASMANIA: Scottsdale, Cuckoo Falls Creek, 31.vii.19g31 (R. J. Tillyard). Five
examples, probably all males; it is possible that these nymphs may belong to the
species L. varia Kimmins, which occurs in this locality. The somewhat irregular
arrangement of the cross-veins in the nymphal fore wing suggests this, but further
confirmation is needed.
Leptoperla sp. No. 2
FIGS. 12-13
Full-grown male nymph. General colour fuscous. of slender build, with rather
short legs: length about 6 mm. Body smooth, sparsely clothed with short setae,
whose apices are slender and curled. Head about as wide as pronotum, antennae
incomplete, moderately stout, tapering.
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 61
Labrum about twice as broad as long, anterior angles rounded. Labium with
glossae a little shorter and narrower than paraglossae, with an apical tuft of setae.
Paraglossae less pointed than in sp. No. 1. Labial palpus rather stouter than in
sp. No. 1; maxillae with palpus rather stouter, mandibles similar.
Fic. 13. Leptoperla spp. Nymphal structures. a-g, sp. No. 1; h-k, sp. No. 2. a, mandibles; 8,
labrum ; c, maxilla; d, labium; e, leg; f, dorsal armature of abdominal tergites, lateral ; g, the same,
dorsal; h, mandibles; 7, labrum; 7, maxilla; k, labium.
Pronotum broader than long,’*somewhat narrowed anteriorly, angles rounded,
margins straight. Strong median and transverse impressed lines, anterior shorter than
posterior, both angled towards each other at their centres. Wing pads of medium
size, anterior elongate, overlapping basal two-thirds of posterior; fuscous, with
longitudinal veins slightly darker. Imaginal venation less distinct than in some
species, few cross-veins in apical half of wings. No incision between anal angle of
wings and nota.
Legs paler than body, similar to sp. No. 1, but with rather fewer short setae.
Abdomen long, cylindrical, somewhat ridged along dorsum, apical margins not
62 A REVISION OF THE AUSTRALIAN AND TASMANIAN
produced. Tenth tergite much elevated in a median dorsal ridge, terminating in
an acute finger, projecting beyond the triangularly produced margin. Basal margin
with a pair of acute triangular apodemes. Anal gills pale pinkish, cerci incomplete.
Sub-anal plates about twice as long as broad, parallel-sided for most of their length,
tapering near apex.
TASMANIA: Scottsdale, a number of male nymphs.
Smaller than sp. No. 1, rather more slender; pronotum is definitely broader in
proportion to its length, and tenth tergite is more produced ; sub-anal plates narrower.
DINOTOPERLA Tillyard
Dinotoperla Tillyard, 1921, Canad. Ent. 58: 43, fig. 40.
Type species: Perla opposita Walker, 1852 (fixed by Tillyard, 1921).
Tillyard, 1921, Tvans. Roy. Soc. S. Aust. 45: 270-273, figs. 2-3.
Fore wing with Rs simple, M forked from near the middle of wing, Cur forked; a
thickened cross-vein between IA and 2A; cubito-anal space with a basal cross-vein
only. Hind wing with Rs simple; M and Rs separating some distance from base ;
M forked, posterior branch fused for a space with Cul, separating again before the
margin of the wing; generally only one cross-vein between the branches of cubitus.
Anal fan rather narrow. Cerci shorter than abdomen.
Distribution: Australia, Tasmania.
Dinotoperla serricauda sp.n.
Fic. 14
Head, thorax, and apex of abdomen brown, segments one to eight of abdomen
whitish. Eyes blackish, antennae brown, about as long as fore wing, basal segment
enlarged. Pronotum almost rectangular, broader than long, slightly narrowed
anteriorly. Legs brownish, slightly darker at the knees. Wings pale fuscous, with
darker venation ; in anterior, R, Rs, and most cross-veins broadly shaded with darker
fuscous. Veins in hind wing not shaded.
g. Apical margin of tenth tergite produced at its centre in a pale process, triangular
and acute from the side; from above broad and pigmented at its base, tapering to a
narrow, rounded apex, upper surface closely set with minute setae. Supra-anal lobe
projecting beyond apex of tergite, slender and blade-like from above, apex abruptly
hooked downwards. Cerci brownish, about 1} mm. long, eleven-segmented, tapering
slightly, segments near base shorter than broad, slightly serrately produced on inner
surfaces. Sub-anal plates scimitar-like, apices somewhat incurved; from beneath,
they are seen to arise from broad bases and are contiguous for most of their length.
Margin of ninth sternite produced in a parabolic subgenital plate about as long as its
sternite, from side appearing as a stout, blunt finger.
2. Wings more elongate. Subgenital plate of eighth sternite very short and broad,
scarcely projecting beyond the margin. Sub-anal plates triangular, from beneath
apices produced in slender fingers: near the upper basal angle is a small, rounded,
whitish pit. Cerci less serrate than in male, twelve-segmented. Margin of tergite
produced, rounded and bent down, whitish, with minute setae as in male.
Length of fore wing, g 9°5 mm., 2? II mm.
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 63
Length of hind wing, g 8-5 mm., 2 9°5 mm.
TasMANIA: R. Ouse, 4.ii.1933, 4 3, 2 2 (R. J. Tillyard) ; Deloraine, 20.xii.1884, I 9.
Type male (R. Ouse) in form of microscope preparation, 3 3, 2 2 paratypes in 2 per
cent. formaldehyde solution, 1 9 paratype from Deloraine, pinned.
Fic. 14. Dinotoperla serricauda sp.n. a, wings 3; b-d, genitalia g, b, lateral, c, dorsal, d, ventral ;
é, genitalia 9, ventral.
Compared with D. carpenteri Tillyard, in fore wing the posterior margin between
Cuz and Curd is more bowed, and Curd is longer. In the hind wing, the anal fan is
broader. Tillyard’s figure does not permit of a useful comparison of the male genitalia.
Dinotoperla opposita (Walker)
FIG. 15
Perla (sg. Chloroperla) opposita Walker, 1852, Cat. Neur. Ins. B.M.1: 171.
Leptoperla opposita McLachlan, 1874, Trans. N.Z. Inst. 6 (Append.): xcii; Banks, 1913, Tvans.
Amer. Ent. Soc. 89: 203.
Dinotoperla opposita Tillyard, 1921, Canad. Ent. 58: 43; id., 1923, Trans. N.Z. Inst. 54: 203.
Walker described the species as follows:
“too. PERLA OPPOSITA.
‘Nigra, nitens, ferrugineo varia; caput antice testaceum, thorace vix latius ; pro-
_ thorace antice non latius, disco ruguloso, lateribus marginatis rectis, angulis sat
acutis; scutellum flavo maculatum; alae subcinereae, ad venas transversus ob-
scuriores.
64 A REVISION OF THE AUSTRALIAN AND TASMANIAN
‘Black, shining, partly ferruginous: head testaceous in front, hardly broader than
the thorax: antennae very minutely pubescent: prothorax minutely punctured,
rugulose on the disc, not broader in front, with a rim on each side and along the fore-
border; sides straight, angles rather sharp: scutellum with a yellow spot in front:
wings very slightly gray, darker about the transverse veins: veins black. Length of
the body 44 lines, of the wings 13 lines.!
‘a, 6. Van Diemen’s Land. From Dr.
Hooker’s collection.
‘c. Van Diemen’s Land. From Mr. Smith’s
collection.
‘d. New Zealand. Presented by Capt. J. C.
Ross.’
The New Zealand specimen and one of Dr.
Hooker’s examples can no longer be traced in
the British Museum collection. McLachlan
and Tillyard, however, were both of the
opinion that Walker’s opposita does not occur
in New Zealand, and that the record was
based on a mis-identification. This is very
probable, as there is a strong superficial re-
semblance between species of different genera
in the Gripopterygidae.
Fic. 15. Dinotoperla opposita (Walker) 9. To Walker's description may be added that
a, wings; b, genitalia, ventral. the two remaining examples are females, one
with the abdomen damaged by insects. The
pronotum is broader than long, anterior margin rounded, making anterior angles
obtuse. The subgenital plate is scarcely produced, broad, slightly emarginate at
its centre, not or but slightly pigmented. Sub-anal plates broad, triangular, outer
margins obtusely excised, apices broader and less divergent than in D. serricauda
Kimmins. Cerci short, thirteen-segmented (damaged and possibly incomplete in
type). Tenth tergite produced in a rounded, deflexed lobe.
Wings a little more pointed than in serricauda Kimmins, and with more numerous
cross-veins in the apical half.
I have selected as type the female from Mr. Smith’s collection, labelled V.D.L.,
51.153; the abdomen has been made into a microscope preparation and one pair of
wings has been mounted dry on a microscope slide.
I have seen no more examples of this species, which is seemingly restricted to
Tasmania. Tillyard records it from Mt. Wellington, Hobart.
EER HESOC Rome -
e* 1.
RAED
eee
Pedanys
AD
OO
f
RPO MAH
Ura Pag
Wate
Dinotoperla fontana sp.n.
Fic. 16
(In fluid.) Head, thorax, and abdomen brownish, the latter with apices of seg-
ments whitish, whitish areas becoming progressively narrower from base to apex.
1 The wing measurement is obviously the expanse, not the length.
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 65
Eyes blackish, antennae brown, about as long as fore wing. Pronotum about three-
fifths as long as broad, narrowed anteriorly, anterior and posterior transverse furrows
strong, linked by a narrow median line and somewhat shallow lateral troughs. Legs
yellowish-brown, knees and tarsi somewhat darker ; ventral surface of femora with
a pair of dark brown lines.
Wings yellowish fuscous, veins slightly
darker. In fore wing, cross-veins in apical ee ee
half numerous (more than twenty), not very = ETF
strong, in dark specimens faintly bordered S
with brownish. In hind wing, Rs and M
separate at some distance from the base.
$. Apical margin of tenth tergite produced
atits centre in a pale process, cylindrical, and
tapering to an acute apex, which is more
slender than in serricauda; its upper surface
clothed with minute setae. At its base are
two chitinous ribs supporting the supra-anal
lobe. The latter is slender, its apex laterally
compressed, from the side dilated, upper
angle rounded, lower hooked. Cerci brownish,
bent downwards, a little over I mm. in
length, ten-segmented, segments becoming
narrower and longer towards the apex. Sub-
anal plates blade-like, upcurved, outer sur-
face with two or three weak, transverse
corrugations. Subgenital plate dark brown,
parabolic, extending to bases of sub-anal
plates.
Q. Pronotum more rugose than in male.
Abdomen almost wholly pale except tenth, : es
part of ninth segments, and the subgenital Fic. 16. eae hagas roe
plate. The latter is produced in a broad - piece Bay pee hat ee pear
parabola, about half the width of the sternite ; d, dorsal; e, genitalia 9, ventral.
the plate is strongly pigmented, extending to
base of segment, the pigmentation constricted about midway. Sub-anal plates
broad, apices acute and divergent. Cerci brownish, twelve-segmented. Tenth ter-
gite pigmented basally, apical margin pale, the basal pigmentation extending in a
narrow finger into the produced part.
Length of fore wing, gf 10-5 mm., 9 9 mm.
Length of hind wing, ¢ 9:5 mm., 2 8 mm.
AUSTRALIA: Federal Capital Territory, Lee’s Springs, xi.1932, 3 J, 1 9; New South
Wales, Mt. Kosciusko, 10-15.xii.1934, I 2 (R. J. Tillyard).
Type male, paratype female (Lee’s Springs) in microscope preparations, remainder
in 2 per cent. formaldehyde solution.
The form of the supra-anal lobe and the more produced tenth tergite in the male,
66 A REVISION OF THE AUSTRALIAN AND TASMANIAN
and the more produced and pigmented subgenital plate of the female will separate
this species from D. serricauda Kimmins and D. carpenteri Tillyard.
Dinotoperla carpenteri Tillyard
FIG. 17
Dinotoperla carpenteri Tillyard, 1921, Trans. Roy. Soc. S. Aust. 45: 273-274, figs. 1-4.
I have been unable to recognize this species with certainty amongst the British
Museum material, and as I am unable at present to study the type, I am repeating
Tillyard’s description and figures. His illustration of the male genitalia is difficult to
Fic. 17. Dinotoperla carpenteri Tillyard g. Wings and genitalia (after Tillyard).
c, cerci, p, ‘penis’, sa, ‘superior appendages’, probably in dorsal view.
understand ; it was made from a microscope preparation and one which appears to
have been much flattened in mounting. Tillyard says it is a ventral view, but the
median lobe bears considerable resemblance to the supra-anal lobe of Dinotoperla
and one of the sub-anal plates (superior appendages) passes beneath it. The other one
is drawn above the cercus, thus apparently being in ventral aspect. It is probable
that in drawing from a flattened mount, Tillyard was mistaken in the relative position
of the various parts. Under the circumstances I have considered it wiser not to attempt
to re-figure the species from an example which might not be correctly determined,
but to leave the elucidation of this species to an Australian or New Zealand ento-
mologist who has access to the type.
‘Forewing, 10 mm. Expanse, 21-5 mm.
‘Head, thorax and abdomen dull blackish, touched with dark brown behind the
eyes and on notum. Eyes brownish-black. Antennae about as long as the forewing,
very slender, about 50 segments, the basal one slightly enlarged. Pronotum rect-
angular, broader than long. Legs dull brownish, the apices of the femora and tibiae
darkened ; the tarsi darker, except at base of first segment. Cerci rather short, I0- to
11-segmented, tapering, delicately haired, the basal segment thickest and much
longer than any of the next few succeeding segments. In the male, the superior
appendages are slender, elongated, curved processes, projecting on either side of the
penis, the basal portion of which is a broad plate, the distal portion slender and
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 67
upcurved. In the female, the ventral plate is deeply bifid, a deep triangular notch
dividing it into two triangular pieces.
‘Wings: Forewing medium fuscous, with irregular paler sub-hyaline spaces between
the cross-veins; i.e., each cross-vein is surrounded by a rectangle of the fuscous
ground-colour, and the spaces between these rectangles are paler. As the positions
and number of these cross-veins is inconstant, varying for each individual, the
pattern thus produced is very irregular, and is never very conspicuous. Hindwing a
uniform medium fuscous. When at rest, the dark and pale areas of the two wings
appear to re-inforce each other, giving the insect a distinctly banded appearance.
‘Locality: Hornsby, near Sydney, N.S.W. Bred from larvae found on rocks in a
small stream in Old Man’s Valley; also found sitting about on the stems of reeds and
grasses near the stream. July to October.
‘Types: Holotype (Hornsby, Sept. 14, 1917, R. J. T.) and series of paratypes in
Tillyard collection, Cawthron Institute, Nelson, N.Z.
“The set specimens have the abdomens so shrivelled that it is impossible to deter-
mine the sex or study the sexual appendages. The descriptions of these organs given
above were made from slides prepared from specimens of the paratype material.’
Dinotoperla brevipennis sp.n.
Fic. 18
Head and pronotum light yellowish-
brown, with considerable darker brown
markings ; remainder of thorax brown.
Antennae brown, a little shorter than
fore wings. Pronotum quadrate, two-
thirds as long as broad, surface finely
rugulose or granulose, posterior angles
obtuse. Legs brownish, knees and two
lines on lower surface of femora darker.
Abdomen largely brownish above, paler
beneath. Wings short, extending only to
apex of abdomen, light fuscous (hind
wing darker), with darker venation, In
fore wing, apical cross-veins pale, all
cross-veins bordered with brownish.
g. Apical margin of tenth tergite
produced at its centre in a pale process,
shorter than in serricauda, apex rounded,
not acute; upper surface finely setose.
Supra-anal lobe short, laterally com-
pressed, and blade-like, its apex from
the side expanded and produced down- ee ee ee : *
ward in a hook. Cerci brownish, bent 3.'5"7" sential & bite bniare ea tab
downward, about xr mm. in length, genital plate, ventral; ¢, genitalia 9, ventral.
ENTOM. 2, 2 H
d
68 A REVISION OF THE AUSTRALIAN AND TASMANIAN
nine- or ten-segmented, those beyond the fifth of about equal length. Sub-anal plates
blade-like, upcurved, with acute apices. Subgenital plate dark brown, parabolic,
extending to bases of anal plates.
9. Abdomen whitish, except tenth segment, subgenital plate and ninth tergite,
which are brown. Sternites two to seven with four tiny brownish plates. Subgenital
plate rather less produced than in D. fontana Kimmins, about half the width of the
segment. It is strongly pigmented, rather broader at base than apex, and with a
large hyaline area at the base, extending in a narrow finger almost to apex. Sub-
anal plates broad, apices divergent, rather stouter than in fontana. Cerci brownish,
about nine-segmented. Tenth tergite roundly produced at its centre, apical margin
narrowly whitish.
Length of fore wing, ¢ 5 mm., 9 5:5 mm
Length of hind wing, ¢ 4:2 mm., 2 4-6 mm.
AUSTRALIA: New South Wales, Bolaro, 22.xii.1935, 3 g, 69; Rule’s Point, 4,450 ft.,
30.x11.1934, 2¢ (R. J. Tillyard).
Type male, paratype female from Bolaro, in form of microscope preparations,
remainder in 2 per cent. formaldehyde solution.
Dinotoperla thwaitesi sp.n.
FIG. 19 a-c
3 (dried). Resembling D. brevipennis Kimmins but with longer wings and differences
in genital structure. Head and pronotum light brownish, with considerable darker
markings. Antennae brown, incomplete. Pronotum quadrate, about three-fourths
as long as broad, median line short, angles obtusely rounded. Meso- and metanota
dark brown. Legs brownish, darker at knees and with darker lines beneath femora.
Abdomen brownish above, paler beneath, particularly towards base. Wings more
elongate than in brevipennis, and with more numerous apical cross-veins. In hind
wing, anal fan is flatter and less evenly rounded; point of separation of M3+4 and
Cur before intercubital cross-vein. Wings brownish (hind darker), venation brown
and most of cross-veins bordered with brownish.
Apical margin of tenth tergite scarcely produced, membranous at its centre.
Supra-anal lobe slender from above, rather longer than in brevipennis, from side
sinuous and terminating in an acute down-turned hook. Cerci brownish, bent down-
wards, about 1:5 mm. long, twelve-segmented, more finely haired than in brevipennis.
Sub-anal plates blade-like, acute, upcurved, rather broader about midway. Sub-
genital plate dark brown, narrower and more pointed than in brevipenmis.
Length of fore wing 9 mm., of hind wing 8 mm.
AUSTRALIA: Victoria, Melbourne (Thwaites), 1 3.
Type (with abdomen mounted in balsam) in collection of Hope Department,
University Museum, Oxford.
Dinotoperla uniformis sp.n.
FIG. 19 d-g
3 (in fluid). Head, thorax, antennae, and legs pale brownish, head and thorax
with obscure, slightly paler markings, abdomen whitish, ninth and tenth segments
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 69
pale brownish. Antennae about as long as fore wing. Pronotum a little broader than
long, narrowed anteriorly, transverse furrows distinct, lateral margins slightly
raised, a narrow median impressed line. Legs practically without markings, either at
knees or beneath femora. Wings pale fuscous, veins darker, a few paler areas in fore
wing. Ten to twelve rather weak cross-veins in apical half of fore wing. In hind wing,
Rs and M separate some distance from base.
Fic. 19. Dinotoperla spp.n. 3. a-—c, D. thwaitesi; d—g, D. uniformis. Genitalia,
a, lateral; b, dorsal; c, subgenital plate, ventral; d, lateral; e, dorsal; f, sub-
genital plate, ventral; g, wings.
Tenth tergite from above triangularly produced at its centre, pale brownish, only
its extreme apex and a narrow median area whitish. From the side it is much less
produced than in the other species. Supra-anal lobe slender, down-curved towards
its apex, which is acute, and excised on its lower surface. Cerci pale brownish, bent
downwards, about I mm. in length, ten-segmented. Sub-anal plates slender, up-
curved, apices acute and divergent. Subgenital plate produced and rounded, an
ovate pigmented patch at its centre.
2 unknown.
Length of fore wing 9 mm., of hind wing 8 mm.
AUSTRALIA: New South Wales, Rule’s Point, 4,450 ft., 30.xii.1934, 1g (R. J.
Tillyard).
Type in form of microscope preparation.
70 A REVISION OF THE AUSTRALIAN AND TASMANIAN
Dinotoperla evansi sp.n.
FIGS. 20 a-c, 21a
3 (in fluid). Head and thorax light reddish-brown. Head practically without
markings, antennae long, reddish-brown, basal segment dark brown. Pronotum
broader than long, slightly narrowed anteriorly, sides straight, anterior margin
slightly produced and rounded; anterior and posterior transverse furrows distinct.
Colour reddish-brown, with a median longitudinal streak of yellowish-brown. Meso-
and metanota reddish-brown. Legs light reddish-brown, knees darker and two dark
lines on ventral surface of each femur and one on each tibia. Abdomen with segments
Fic. 20. Dinotoperla spp.n. a-c, D. evansi; d, D. nigricoxa. a, g lateral; b, 2 subgenital plate,
ventral; c, 2? sub-anal plate, ventral; d, genitalia 9, ventral.
one to five whitish, remainder largely dark brown. Wings smoky-brownish (hind
wings the darker) with darker venation, costal and stigmal areas darker; apical
cross-veins pale, in fore wings bordered with brownish.
Apical margin of tenth tergite elevated, but without a pale, produced process, its
place being taken by a flat membranous area. Supra-anal lobe rather long, com-
pressed laterally, apex curved downward, rounded, and carrying a small hook. Cerci
brownish, bent downward, rather more than I mm. in length, about ten-segmented.
Sub-anal plates blade-like, upcurved, with rounded apices. Subgenital plate dark
brown, rather narrow, with a very flatly rounded apex.
2. General appearance similar, wings rather darker ; abdomen with only segments
nine and ten, subgenital plate, sub-anal plates, and cerci dark brown. Subgenital
plate broad, not produced, pigmented area with a broad triangular excision at base.
Sub-anal plates with slender produced apices. Cerci slender, about thirteen segments.
Length of fore wing, ¢ I0-5 mm., 9 10:5 mm.
Length of hind wing, ¢ 9 mm., 9 9 mm.
S. AUSTRALIA: Mt. Lofty, x.1931, 1 § (J. Evans) ; Cudlee Creek, 20.xi.1934, 6 2
(R. J. Tillyard) ; Adelaide, 2 2 (Hope Dept., Univ. Mus., Oxford).
Type male, paratype female in form of microscope preparations, others in 2 per
cent. formaldehyde solution, females from Adelaide pinned.
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) aI
This species resembles D. uniformis, but may be distinguished by the form of the
supra-anal lobe and subgenital plate in the male, and by the venation. One of the
females from Adelaide is larger than those from Cudlee Creek (fore wing, 11:5 mm.)
and has more numerous apical cross-veins, but the form and pigmentation is similar.
Dinotoperla fusca sp.n.
Fic. 21 5, c
$ unknown.
@ (in fluid). Head fuscous, with darker markings near antennal bases and behind
epicranial sutures. Antennae fuscous, with blackish basal segment. Pronotum
fuscous, anterior and posterior margins finely bordered with blackish-brown, and
Fic. 21. Dinotoperla spp.n. 9. a, D. evansi wings; b, D. fusca wings; c, D.
fusca genitalia, ventral.
with blackish rugosities on the disk; quadrate, broader than long, anterior and
posterior transverse furrows rather broad. Meso- and metanota dark brown. Femora
_yellowish-brown, ventral carinae and apical third of femur dark brown, tibiae and
tarsi dark brown. Wings brownish hyaline, anterior with whitish areas in most of the
apical, medio-cubital, and cubital cellules. Venation brownish, many of the apical
cross-veins weak and whitish, those in the anterior wing numerous. Abdomen
whitish, pattern of subgenital plate and parts of the ninth and tenth segments, and
cerci brownish-black. Eighth sternite forming a subgenital plate whose margin is
only very slightly produced. There is a strongly pigmented pattern resembling that
in D. evansi, but the pigmented area is much longer than broad, the lateral margins
72 A REVISION OF THE AUSTRALIAN AND TASMANIAN
are more concave and the pale median area is narrower and extends to the apex.
Pigmentation of the ninth sternite interrupted centrally, of tenth complete. Sub-
anal plates broad, subtriangular, outer margins concave, inner straight. Cerci short,
about eleven-segmented.
Length of fore wing 10:5 mm., of hind wing 9:5 mm.
TASMANIA (J. W. Evans), 2 9. ;
Type in form of microscope preparation, paratype in 2 per cent. formaldehyde
solution.
This species resembles D. evansi and D. nigricoxa in the pattern of the female sub- —
genital plate. From the former it may be separated by the rather more pointed wings
and different pattern. In migricoxa the wings are almost without pattern and there
are far fewer apical cross-veins.
Dinotoperla nigricoxa sp.n.
Fic. 20d
¢ unknown.
2 (in fluid). Head yellowish-brown, chestnut-brown between ocelli, occiput —
mottled with darker yellowish-brown. Antennae dark brown, about as long as fore
wing. Pronotum transverse, slightly narrowed anteriorly, anterior margin convex;
transverse furrows broad and distinct, a short median impressed line ; disk on either
side with obscure brownish markings, surface finely rugulose. Legs dull brownish,
darker on knees and tarsi: each femur below with two narrow black streaks, posterior
streak obsolete at base. Meso- and metacoxae on upper surface at base with a distinct
black streak. Mesosternum on each side with a dark brown streak.
Wings brownish, with brown venation ; apices slightly more acute than in uniformis ;
in three out of four examples, Cur in fore wing forks some way beyond the fork of M
and termination of Cuz. (Lines drawn from these points to the fork of Cur form
approximately a right angle; in wniformis this angle is decidedly obtuse.) In hind
wing Rs and M separate some distance from base.
Abdomen whitish, except eighth sternite and parts of ninth and tenth segments,
which are brownish. Eighth sternite more or less produced at its centre in a sub-
genital plate, pigmented as in Fig. 20d. Sub-anal plates broad, apices triangular and
divergent. Cerci short, nine- or ten-segmented.
Length of fore wing Io mm., of hind wing 9 mm.
AUSTRALIA: New South Wales, Mt. Kosciusko, 10-15.xii.1934, 4 2 (R. J. Tillyard). ©
Type female in form of a microscope preparation, paratypes in 2 per cent. formal-
dehyde solution.
This species may possibly be the female of D. uniformis, but as there are a number ~
of minor differences, it has been considered preferable to regard it as distinct. The
chief differences are in the shape of the pronotum, which is less narrowed anteriorly,
the black streaks on the femora and the black markings on the meso- and metacoxae.
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 73
Dinotoperla fasciata Tillyard
FIG. 22
Dinotoperia fasciata Tillyard, 1924, Trans. Roy. Soc. S. Aust. 48: 193; 1926, Ins. Aust. & N.Z.:
119, pl. 10, fig. 13.
The female holotype has been severely damaged by Anthrenus, and little but the
wings remain. Tillyard describes it as follows:
‘Q. Length of body 6 mm., of forewing 10 mm., antenna 7 mm., cerciI mm. Head
black, antennae dark brown ; pronotum pale yellowish ; rest of thorax black, abdomen
dark brown. Forewings suffused with pale fuscous, venation darker fuscous ; distal
cross-veins surrounded by darker fuscous areas
and so arranged that from three to four irregu-
lar transverse fasciae cross that part of the
wing. Hindwings darker fuscous with a few
darkened cross-veins placed distally between
C, Rr, and Rs.’
To this may be added that there are two or
three cross-veins in the pterostigmatic area in
both wings, and that there are distal cross-
veins in the hind wing other than those
specially mentioned by Tillyard.
Holotype female (National Park, S. Queens-
land, 1,500-2,000 ft., Dr. A. J. Turner), in the British Museum (Nat. Hist.), from the
Tillyard collection. Paratype female, slightly smaller in size and with less distinct
fasciae (same locality, March, 1921, G. H. Hardy) in Queensland Museum, Brisbane.
Fic. 22. Dinotoperla fasciata Tillyard.
Q Type. Wings.
Dinotoperla sp. nymph
FIGS. 23-24
General colour yellow-brown, with scanty darker markings ; of medium build, not
very slender, about 7 mm. long; sparsely clothed with small hairs, expanded at their
_ bases, apices slender, generally curved or hooked.
Head a little wider than pronotum, widest at compound eyes, rounded behind ;
stem of epicranial suture about two-sevenths as long as pronotum. Antennae
moderately long, extending beyond the apex of the posterior wing pads; first seg-
‘ment large, flattened dorso-ventrally, slightly tapering, second segment short, third
about as long as second, remainder short, tapering to apex; each segment almost
bare, with only a few short bristles at apex.
Labrum short, broad, anterior angles rounded. Labium with the glossae and para-
glossae of about equal length, apices ciliate. Incision between glossae deeper than
_ that cutting off paraglossae, both very narrow; paraglossae rather broader than
glossae, outer margins strongly convex. Labial palpus three-segmented, segments
increasing in length from base to apex in ratio I: 1-5: 2; second segment dilated
towards its apex, third ovate. Maxillae well developed: lacinia with two acute teeth
at its apex, beneath which, on inner surface, is a short row of stiff setae ; galea scarcely
extending beyond lacinia, curved, tapering towards its apex, which is obliquely
74 A REVISION OF THE AUSTRALIAN AND TASMANIAN
truncate and set with minute setae; palpifer indistinct, membranous; palpus five-
segmented, one and two short, subequal, three as long as one and two combined, four
Fic. 2 3. Dinotoperla sp. Nymph and nymphal venation.
rather longer than two, and five as long as three and four together. Mandibles heavily
chitinized, with a number of strong teeth near the apex, beneath which is a well-
developed mola with a row of short setae.
Fic. 24. Dinotoperla sp. Nymphal struc-
tures. a, labrum; b, labium; c, maxilla;
d, mandibles; e, leg; f, vestiture.
Pronotum with a number of straggling
brownish marks on disk, median suture pale;
broader than long, slightly narrowed anterior-
ly, angles rounded. Wing pads extending
backwards and almost horizontal. Fore wing
pads elongate, overlapping more than half of
the posterior pads; latter somewhat broader,
anal fan not very pronounced. Main venation
visible as lines of pigmentation bearing short
hairs. If viewed by transmitted light, the
veins and cross-veins appear as clearer spaces
in the tissue of the wing. Under these condi-
tions, the cross-vein designated M5 by Tillyard
is very evident and is generally thicker than
the succeeding medio-cubital veins.
Legs short and strong, prothoracic the
shortest, increasing in length to the meta-
thoracic, sparsely clothed with short dilated
hairs, outer surface of tibiae with a scanty
row of longer, very fine setae. Femora shorter and broader than tibiae, anterior
surfaces with a brownish line, not reaching the apex. Tibia with a slender band
A A Rg it tt ae my vam
© ON ae ee eee, ~
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 75
below the knee and a narrow longitudinal streak on anterior surface. Tarsi three-
segmented, second very short, third about four times as long as first, and provided
with a pair of simple claws at apex.
Abdomen cylindrical, segments one to nine with straight apical margins, and
dorsally with a double row of spots, fairly close together. Tenth tergite as long as
five preceding tergites, apical margin produced and somewhat elevated in male,
rounded in female. Sub-anal plates elongate, triangular. Projecting beyond them is
a dense tuft or rosette of mauve anal gill-filaments. Cerci short, tapering to apex,
about twenty segments, basal segments shorter than broad, apical segments longer
than broad, segments with a few short apical setae, no dorsal fringe.
AusTRALIA: F.C.T., Cottar River, 14.ix.1933 (W. L. Raitt) ; other examples, rather
darker, from New South Wales, Rule’s Point, 4,450 ft., 30.xii.1934, and TASMANIA,
R. Ouse, 4.ii.1933 (R. J. Tillyard).
TRINOTOPERLA Tillyard
Trinotoperla Tillyard, 1924, Trans. Roy. Soc. S. Aust. 48: 193.
Type species: T. ivrorata Tillyard, 1924 (fixed by Tillyard, 1924).
Fore wings with Rs forked, sometimes three-branched, M forked near middle of
wing. Cur two- or three-branched (sometimes in males the lowest branch is very
short and forms a closed cell with the one above it) ; cubito-anal space with only a
basal cross-vein ; a thickened cross-vein between IA and 2A. Hind wing with Rs
forked, M forked, posterior branch soon fusing with Cur, and separating again,
generally some distance from the wing margin; apex narrowly rounded; anal fan
moderately or rather narrow, generally without any cross-veins. Cerci short. Size of
species medium to large, expanse from about 25 mm. to over 50 mm.
Distribution: Australia; ? Tasmania, known from nymphs only.
Trinotoperla irrorata Tillyard
FIG. 25
Trinotoperla irrorata Tillyard, 1924, Trans. Roy. Soc. S. Aust. 48: 194, fig. 1; x90, Ins. Aust. &
N.Z.: 119, pl. 10, fig. 14.
The type and allotype have suffered severely from the depredations of Anthrenus,
little remaining apart from the wings. I am therefore quoting the original description
and adding extra notes where possible.
“2 Holotype: Length of body 20 mm., of forewing 25 mm., antennae 17 mm.,
cerci barely 3 mm. Body dark fuscous, marked with dull brown; antennae dull
brownish with slight fuscous annulations; legs dull brownish varied with dark
fuscous. Wings pale greyish with dark fuscous veins, those of the basal half of the
hind wing inclining to brownish’ (the whole of the fore wing and to a lesser degree,
the distal part of the hind wing, speckled with dark brown spots, and most of the
cross-veins margined with darker brown. Posterior margin of fore wing with a small
distinct angular lobe between Cur and Cuz, rather more pronounced than in Till-
yard’s figure). ‘Forewing with a complete series of costal veinlets ; hindwings with
hm and three or four distal costal veinlets only.’ (Costal margin of hind wing very
ENTOM. 2, 2 I
76 A REVISION OF THE AUSTRALIAN AND TASMANIAN
slightly scalloped.) ‘Thickened cross-vein between IA and 2A in forewing blackish,
very prominent.’
Fic. 25. Trinotoperla irrorata Tillyard.
Q Type. Wings.
‘3g Allotype. Smaller and slightly paler
and less strongly irrorated than female.
Length of body 17 mm., forewing 21 mm.
Supra-anal plate with a slender copulatory
process directed upwards and ending in a
small hook directed posteriad; paraprocts
upcurved, forming two flatly rounded lobes
directed forward and upward. Tenth tergite
with a raised flap distally.’
AUSTRALIA: New South Wales, Mt. Kos-
ciusko, 5,000-5,500 ft., 24.xi.1g2I (R. J.
Tillyard). Holotype female, allotype male in
British Museum (Nat. Hist.), from the Till-
yard collection.
Trinotoperla australis Tillyard
Fics. 26-28
Trinotoperla australis Tillyard, 1924, Trans. Roy. Soc. S. Aust. 48: 194; 1926, Ins. Aust. &
N.Z.: 119, pl. Io, fig. 15.
Female holotype: Head, thorax, and abdomen fuscous; antennae fuscous, three-
quarters as long as fore wing. Pronotum a little broader than long, quadrate, anterior
|
Fic. 26. Trinotoperla australis Tillyard. 3 genitalia. Fic. 27. Trinotoperla spp. 2 Wings.
a, lateral, b, apex of supra-anal lobe, lateral, a, T. australis Tillyard; b, T. nivata sp.n.
c, dorsal, d, subgenital plate, ventral.
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 717
and posterior margins slightly rounded, a median impressed line, not reaching
anterior margin, and posteriorly ending in a transverse groove. Legs fuscous. Wings
pale brownish-fuscous, venation darker, cross-veins narrowly bordered. Very few
costal cross-veins, no pronounced angular lobe on posterior margin of fore wing.
Costal margin of hind wing evenly arched. Thickened cross-vein between IA and 2A
in fore wing brownish, less conspicuous than in zrrorata.
Subgenital plate of the female holotype is creased and probably distorted. The
apical margin is slightly produced and rounded, and appears to have a small rounded
excision at its centre. I strongly suspect that this excision is accidental, as I have
examined a number of examples from South Australia, agreeing in other respects, in
which the subgenital plate is evenly rounded and not excised. I am figuring the sub-
genital plate of one of these for comparison. In most of these there is a strongly pig-
mented bar near each lateral margin.
Sub-anal plates broad at their bases,
with rather broad rounded apices,
which are divergent. Cerci filiform, >
brownish, incomplete in the type, in
other examples about fifteen-segmen-
ted. Apical margin of tenth tergite
deflexed, rounded, with a slight pro-
jection at its centre.
Male. Tenth tergite produced in a
pale triangular process, rather flattened
and acute from the side. Supra-anal
lobe long, slender from above; from
the side rather deep and somewhat ,
dilated at its apex. Just before the
apex on the ventral surface is a small
acute excision, giving the apex the ye. 28. Trinotoperla spp. 9. a-b, T. australis
appearance of a blunt hook. Basad of Tillyard; c, T. nivata sp.n. a, genitalia ventral of
this excision are a few minute teeth, bolotype; 5, subgenital plate of another example ;
c, genitalia ventral.
Sub-anal plates long, slender, and a
little upcurved from the side, apices incurving and acute. From beneath they are
broad at their bases, inner lower surfaces membranous. Cerci short, brownish, curv-
ing downwards, basal segment rather constricted from the side, about seventeen
segments. Subgenital plate broad, apical margin rounded, and moderately produced.
The basal margins are more heavily pigmented than the remainder of the tergite.
Length of fore wing, ¢ 18 mm., ? 18-20 mm.
Length of hind wing, ¢ 16 mm., 2 17-18 mm.
AusTrRALi<: The locality of the holotype is New South Wales, Towac, near Mount
Canoblas, 7.x.1916 (R. J. Tillyard). I have seen other specimens from Mt. Kosciusko,
Spencer’s Creek, xii.1932, and Bolaro, 22.xii.1935, and from the Murrumbidgee
River, F.C.T., i.1922, all collected by R. J. Tillyard, and other pinned examples from
the McLachlan Collection, labelled Melbourne, S. Australia, or merely Australia.
This species bears a superficial resemblance to the new species nivata and minor, but
78 A REVISION OF THE AUSTRALIAN AND TASMANIAN
may be distinguished by the male genitalia, and by the rounded sub-anal plates of
the female. The wings are rather narrower.
Trinotoperla nivata sp.n.
Fics. 27, 28
¢ unknown.
(in fluid). Head reddish-brown, with a small area on the clypeus, and the post-
ocellar sutures pale. Antennal brown, long (incomplete in the type). Thorax fuscous,
pronotum a little broader than long, slightly narrowed anteriorly, angles not pro-
duced ; a short, median, impressed line, all margins somewhat elevated. Legs medium
fuscous, knees darker, a pale spot in centre of anterior surface of each femur. Lower
anterior apical angle of each femur produced in a small tooth. Abdomen whitish,
except tenth tergite, eighth to tenth sternites, sub-anal plates and cerci, which are
brownish. Wings greyish-brown, not irrorated, anterior darker than posterior ; veins
darker, faintly shaded. A number of costal cross-veins in fore wing ; posterior margin
of this wing scarcely produced in a lobe about the end of Cur. Angle between anal
fan and posterior margin of hind wing clearly greater than a right angle.
Subgenital plate about twice as broad as long, apical margin not very produced,
broadly excised at its centre ; brown, with a quadrate area at the centre of the apical
margin, and the lateral margins pale. Sub-anal plates broad at bases, apices produced
in acute fingers, curving upward and slightly inward. Cerci short, about 1-7 mm.,
sixteen or seventeen segments, basal segments much broader than long.
Length of fore wing, 16 mm., of hind wing, 15 mm.
AUSTRALIA: Victoria, Snowy River, 3.1.1933, I 9 (R. J. Tillyard).
Type is a microscope preparation.
T. irrorata Tillyard, which this species somewhat resembles, may be separated by
its larger size, distinctly mottled fore wing, a small but distinct rounded lobe on the
posterior margin of fore wing, and by the angle between the anal fan and the posterior
margin of the hind wing being only a little greater than a right angle. The female
holotype of ivrorata suffered greatly from the ravages of Anthrenus whilst in Till-
yard’s possession, and little now remains of the body. The sub-anal plate appears to
be of the pattern of mivata, but the apex is less acute.
Trinotoperla minor sp.n.
FIG. 29
(In fluid.) General appearance much as in T. nivata. Legs with the pale area of the
femur less pronounced, upper surface darker, lower apical angle toothed. Abdomen
whitish, except eighth to tenth segments in male, and tenth tergite, eighth to tenth
sternites in female, which are brownish. Wings rather more elongate than in nivata,
a little paler, and without costal cross-veins, apart from the humeral. In the hind
wing there may be a cross-vein between IA and 2A near their apices. The angle formed
by the posterior margin and the anal fan is more obtuse.
g. Tenth tergite from above produced in a rather narrow, membranous lobe or
flap, rather flattened from the side. Supra-anal lobe slender, somewhat dilated before
ee
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 79
its apex, which curves downward in an acute spine. Cerci pale brownish, curving
gently downward, seventeen-segmented, segments at the base very short and broad,
gradually increasing in length and becoming narrower towards the apex. Sub-anal
plates rather broad, especially from beneath, curving upward on either side of the
supra-anal lobe, and with rounded apices. Subgenital plate strongly produced,
triangular, with a truncate apex.
Q. Very like T. nivata. Subgenital plate without a pale quadrate area at the centre
of the apical margin. In nivata this pale area seems to be more closely set with fine
Fic. 29. Trinotoperla minor sp.n. a, wings 3; b-d, genitalia g, b, dorso-lateral, c, dorsal,
d, ventral; e, genitalia 2, ventral.
hairs than its surroundings, but in minor the clothing is uniform. Apices of the sub-
anal plates a little more incurved, and slightly longer in comparison with their bases.
Cerci similar, seventeen-segmented, about 1-6 mm. long.
Length of fore wing, g 11 mm., 9 12 mm.
Length of hind wing, g 10 mm., 9 II mm.
AUSTRALIA: New South Wales, Bolaro, xii.1935, 2 6, 32 (R. J. Tillyard).
Type male, paratype female as microscope preparations, remainder in 2 per cent.
_ formaldehyde solution. |
Trinotoperla minor nymph
FIGs. 30-32
(Mature nymph, about to emerge.) General colour brownish, pronotum and legs
paler; medium build, legs longer than in Dinotoperla, length of nymph about 9 mm.
A conspicuous fringe or crest of fine setae along the median dorsal line of body, and
along legs, cerci, and basal segments of antennae. Body clothed with a sparse mixture
of fine hairs and short stout ovate setae. Head dark brown, broader than pronotum,
80 A REVISION OF THE AUSTRALIAN AND TASMANIAN
widest at compound eyes, slightly rounded behind ; stem of epicranial suture about
one-third as long as pronotum. Antennae about as long as body, filiform, tapering ;
first segment large, with a dorsal crest of fine setae, second smaller, but longer than
any of the next few succeeding segments, which become progressively longer and
narrower from base to apex.
Labrum short and broad, anterior angles rounded. Labium with glossae slightly
shorter than paraglossae, but of about equal width, apices with a dense tuft of fine
setae. A narrow incision separating the
glossae to their bases, and a wider one divid-
5 ing them from the paraglossae, whose outer
margins are convex. Palpus three-segmented,
segments of roughly equal width, increasing
in length from base to apex in ratio I : 1°5 : 2.
Maxillae well developed, lacinia with a pair
of broad truncate teeth at its apex, beneath
FI Ne which on its inner surface is a row of stout
ie, OARS ‘ setae. Maxillary palpus five-segmented, aris-
iz “ ing from a membranous palpifer; first and
\\ \ ‘ : second segments subequal, third somewhat
. shorter than the first and second together,
= me fourth a shade longer than the second, and
the fifth as long as the third.
ae XE 4 Pronotum pale yellowish-brown, with a
= es short, impressed, dark brown median line,
a and on either side of it a curved brownish
cz band, its convex surface inwards, and a
smaller streak near the posterior angles.
A R Pronotum quadrate, slightly broader than
Fic. 30. Trinotoperla minor spn. Nymph. long, angles rounded; posterior transverse
furrow more noticeable than anterior. Wing
pads extending backwards, almost horizontal. Fore wing pad elongate, cover-
ing about half of posterior pad; the latter is broader, the anal fan extending a full
half of the posterior margin. Enclosed wings folded ready for emergence, venation
appearing as pale lines on brownish ground of wing cover. (The venation figure has
been drawn from a younger nymph, in which the pads are naturally smaller and
relatively broader.)
Legs moderately short, prothoracic the shortest, each with a dense crest of fine long
setae situated on the dorsum of each coxa, trochanter and tarsus, anterior dorsal
margin of femur and anterior surface of tibia. Femur shorter than tibia, with a broad,
light brown band before the knee, and a faint stripe of the same colour near the fringe,
lower apical angle slightly produced. Tibia with a brownish band at the knee and the
outer surface narrowly brown. Tarsus brownish, second segment shorter than the
first, third about three times as long as first and second together, claws simple.
Abdomen cylindrical, brownish, segments one to nine with whitish apical margins, —
tenth tergite rather longer than eighth and ninth together, the centre of its apical
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 81
margin produced ina rounded lobe, which is itself further produced in a slender finger.
A dense tuft of whitish anal gill-filaments. Sub-anal plates brown, roughly triangular,
apices divergent, rounded, whitish, outer margins broadly excised. Cerci long, with
more than forty segments, slender, with a dense dorsal crest of fine, long setae ; each
segment ventrally at its apex with a few short setae.
AUSTRALIA: New South Wales, Bolaro, 22.xii.1935 (R. J. Tillyard).
The material furnishing this description was collected at the same time and place
as the adults of T. minor. It consists chiefly of fully matured nymphs, so mature that
the wings within the pads are folded ready for emergence. There are also from the
same locality two or three smaller nymphs, probably about half-grown, in which the
imaginal venation can be discerned, and this venation closely resembles that of
T. minor adults. I think there can be little doubt that these are the nymphs of
T. minor.
Trinotoperla sp. No. 1
FIGS. 31-32
Last instar nymph: Related to T. minor but differing in details, notably venation
of hind wing, form of mouth-parts and vestiture. General colour light brownish, with
darker markings, length about 13 mm. Median dorsal fringe less prominent ; body
sparsely clothed with a mixture of short curved setae and short dilated hairs. The
3A
Fic. 31. Trinotoperla spp. Nymphal wing pads. a, T. minor sp.n.; b, T. sp. No. 1.
latter are somewhat excised at their apices and trough-shaped in cross-section ; those
of T. minor are relatively larger and cylindrical in section. Head dark brown, paler
at centre of vertex, broader than pronotum, stem of epicranial suture between one-
third and one-quarter as long as pronotum. Antennae longer than body (about
16 mm.), filiform, tapering, first segment large, with a crest of fine setae, second
smaller, but broader and longer than any of the next few succeeding segments, which
also carry a dorsal fringe. Terminal segments longer than broad.
Labrum short and broad, narrowing to a broadly and shallowly excised apex.
Labium with the glossae about the same length but somewhat narrower than the
paraglossae, apices more pointed than in T. minor, and armed with a tuft of setae.
Outer margin of paraglossae slightly sinuous, so that the apex is more acute than in
IT. minor. Palpus three-segmented, first about half as long as second, which is a little
82 A REVISION OF THE AUSTRALIAN AND TASMANIAN
shorter than third. Maxillae well developed, lacina terminating in a single truncate
tooth, and a number of stout setae; galea much as in T. minor. Maxillary palpus
five-segmented, extending about as far as apex of lacinia; first and second subequal,
third twice as long, fourth a little longer than second, fifth about as long as third.
Mandibles with a few rather blunt teeth.
Pronotum similar to T. minor, but more heavily marked. Lower anterior apical
angle of femur not produced. Tibia with an additional narrow brown band on inner
anterior surface, parallel to fringe. Abdomen cylindrical, brownish above, tergites
Fic. 32. Tvinotoperla spp. Nymphal structure. a-g, T. minor; h-m, T. sp. No. 1. a, labrum;
b, labium ; c, maxilla; d, mandibles; e, leg; f, portion of cercus; g, vestiture ; h, labrum ; 7, labium ;
7, maxilla; k, mandibles; m, vestiture.
one to nine with a whitish apical margin ; sternites light yellowish-brown, with golden
pubescence. Tenth tergite longer than eight and nine together, centre of its apical
margin produced, obtuse, not produced in a finger as in T. minor. Anal gills whitish. —
Sub-anal plates much as in minor. Cerci long (Io mm)., slender, about sixty segments,
carrying a dorsal fringe as in minor.
TASMANIA: Scottsdale, Cuckoo Falls Creek, 31.vii.1931 (R. J. Tillyard).
Four male nymphs, two apparently last instar, and two with very rudimentary
wing pads. Up to the time of writing I have not seen any adult Trinotoperla from
Tasmania, but from wing-venation these nymphs undoubtedly belong to that genus.
The presence of cross-veins in the anal fan suggests Eunotoperla, but the venation
of the fore wing is*different. The practice of describing new species of Plecoptera
from the nymphs alone is undesirable, and in consequence I am giving these nymphs
numbers instead of names. I have also seen single examples of two more species of
Trinotoperla from the same locality, but as material is so scanty I propose to do
nothing further with them at present.
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 83
Trinotoperla sp. No. 2
Fics. 33, 34
Mature nymph: General colour light yellowish-brown, male about 9 mm., female
about 12 mm. long; medium to rather stout build, legs long and sprawling ; clothed
throughout with short curved setae, which arise from globular tubercles: setae
widened at their bases, apices very slender. Head flattened, only slightly wider than
Fic. 33. Trinotoperla sp. No. 2. a, nymphal wing pads; b, tracheal venation.
pronotum ; antennae about as long, or a little longer than body, filiform, tapering,
a small dorsal fringe at base.
Labrum bent downwards, concealed under the head, apex directed caudad ; short
and broad, apical margin slightly emarginate, angles rounded. Labium with the
glossae slightly shorter and narrower than the
paraglossae. Palpus short and stout, not reaching
apex of paraglossae, apical segment about twice
as long as basal, second rather shorter than third.
_ Maxilla strong, lacinia with about three acute
teeth at its apex, and a number of bristles. Galea
tapering, rather hairy. Maxillary palpus rather
short and stout, first segment half as long as
second, which is two-thirds as long as third.
Fourth as long as second, fifth as long as third.
Mandibles strong, with numerous acute teeth.
Pronotum quadrate, side margins parallel or
slightly convergent anteriorly, anterior and
posterior margins gently rounded, angles pro-
nounced; a slight median line and a strong
transverse posterior furrow. Wing pads large,
anterior elongate, posterior broader, yellowish-
brown, venation indicated by rows of brownish
setae, arising from globular bases. In the only
large male nymph the tracheal venation is
Fic. 34. Trinotoperla sp. No. 2.
Nymphal structures. a, labrum;
b, labium; c, maxilla; d, mandibles;
e, leg; f, vestiture.
quite evident as well as the imaginal venation, and in the hind wing the fusions of
Rs and M, and M3+-4 and Cur are clearly shown, confirming Tillyard’s study of
ENTOM. 2, 2 K
84 A REVISION OF THE AUSTRALIAN AND TASMANIAN
Dinotoperla. Cross-veins in the apical half of the wing dense, including several in
the pterostigmatic area.
Legs rather long and sprawling, in the male giving a somewhat Ecdyonurid
appearance. Trochanters, particularly of anterior legs, with an acute spur at lower
apical angle. Femora strongly carinate dorsally, all legs with a dense dorsal fringe.
Abdomen cylindrical, in male shorter than hind femur, in female longer. In male,
segments one to nine very short, tenth tergite as long as tergites three to nine, apical
margin produced in an acute, somewhat raised point. Basal margins of tergites nine
and ten each with a pair of stout finger-like apodemes. Cerci long, tapering, with at
least thirty-six segments, bearing a dorsal fringe. Sub-anal plates moderately long,
broad at base, tapering near apex, which is rounded. Anal gills pale mauve. In
female, segments one to nine longer than in male, tenth tergite about as long as
tergites seven to nine, apical margin evenly rounded. Apodemes of tergites nine and
ten more noticeable. Cerci similar to male, sub-anal plates rather shorter and
blunter.
AUSTRALIA: New South Wales, Bolaro, 22.xii.1935, 3 2; Rule’s Point, 4,450 ft.,
30.x11.1934, I gf, 2 2 (R. J. Tillyard).
The rather sprawling appearance of the nymph, the backwardly deflected labrum,
and the acute spine on the anterior trochanter separate this species from any other
Trinotoperla nymph which I have seen.
Fam. NEMOURIDAE Klapalek 1909
This family, which occurs so abundantly in the Northern Hemisphere, is represented
in the Southern by about eight genera, comprising at present comparatively few
species. They differ to some extent from the typical Northern Nemouridae, particu-
larly in the venation, the fore wing resembling more that of the family Leuctridae.
They are nevertheless true Nemouridae, as in the hind wing the media is clearly
forked and there are five veins on the anal fan. In the Leuctridae the media is ©
apparently simple, Cur being apparently forked, and there are only three veins on —
the anal fan. It is probable that in the latter family the cross-vein from the media
to the cubitus is the branch M3+4, which then fuses with Cur and subsequently
diverges again to form the apparent fork of Cur. In the hind wing, Rs and M are
united at base, whereas in Leuctridae M arises from a cross-vein uniting Rs and Cu.
In view of the differences in venation, it seems desirable to place the Nemouridae
of the Southern Hemisphere in a separate subfamily, characterized as follows:
NOTONEMOURINAE Ricker 1950!
Fore wing with Cuz long, generally extending well into the apical half of the wing;
cross-vein in the pterostigmatic area of both wings either absent or present. Hind
wing with the media forking at or before the radio-medial cross-vein.
The subfamily will include the following genera:
Aphanicerca Tillyard S. Africa.
Aphanicercella Tillyard
Aphamicercopsis Barnard
>”
»?
' See Appendix, p. 92.
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 85
Desmonemoura Tillyard S. Africa
Neonemura Navas Chile,
Notonemoura Tillyard New Zealand.
Spaniocerca Tillyard ys ‘. Australia, Tasmania.
Spaniocercoides Kimmins - os
Udamocercia Enderlein S. America, Tasmania.
Neonemura Navas is probably a synonym of Udamocercia Enderlein. It is described
from a female, which possesses an ovipositor similar to that of Udamocercia. Nemura
pirioni Navas, from Chile, should also be transferred to the Notonemourinae on the
evidence of the long Cu2 figured in the fore wing ; it is probably a Udamocercia.
The venation of the Nemouridae as a whole offers few characters of value for
generic separation. It is comparatively simple, though not primitive in type, and
tends to be variable in the position and number of the cross-veins. The males of the
three species of Spaniocerca from Australia and Tasmania which I have examined,
whilst obviously related to each other, differ somewhat in genitalic pattern from the
New Zealand species; had the females shown similar affinities, it might have been
desirable to separate generically the Australian and Tasmanian species from those
occurring in New Zealand. Unfortunately the females differ in genitalic pattern not
only from each other but also from S. zelandica Tillyard. Pending the acquisition of
further material from these regions, it seems preferable to retain the Australian and
Tasmanian forms in Spantocerca and to extend its generic diagnosis to accommodate
them, rather than to make a new genus based on male genitalic characters only.
The two remaining Nemourid species with which I am dealing in this paper are
placed for convenience into the genus Udamocercia Enderlein. The male genitalia of
one of the species (U. albomacula) are almost identical in pattern with those of two
South American species of this genus. The other species shows a striking develop-
ment of the supra-anal lobe, and the cerci are modified, but the general pattern is
similar. Both species possess a cross-vein in the pterostigma of both wings (contrary
to Enderlein’s diagnosis), but this is probably variable as in both the South American
species which I have examined it is present, although sometimes obscure. In view
of the scanty material available—two males of different species, and a female which
possibly belongs here—it seems unwise to erect a new genus for these Tasmanian
species now.
SPANIOCERCA Tillyard
Spaniocerca Tillyard, 1923, Trans. N.Z. Inst. 54: 216, fig.; Kimmins, 1938, Ann. Mag. Nat. Hist.
(11) 2: 572.
Type species: Spaniocerca zelandica Tillyard, 1923 (Monotypic genus).
Revised generic diagnosis
Sc arching up so as to meet or almost meet costa ; cross-vein M—-Cu in fore wing
ending above on M3+ 4 at an angle to it ; the latter vein generally curved as it leaves
the transverse cord. Pterostigma in both wings without, or with only one cross-vein.
Anal fan moderately narrow, somewhat excised at apex of fourth anal vein.
g. Ninth tergite with apical margin generally produced in a pair of spines or lobes.
ENTOM. 2, 2 K 2
86 A REVISION OF THE AUSTRALIAN AND TASMANIAN
Ninth sternite more or less produced apically as a subgenital plate, with a median
ventrallobe. Titillators of varying length, two-branched (New Zealand) or unbranched
(Australia, Tasmania), arising near the apex of the ninth sternite. Cerci simple,
sometimes angled. Supra-anal lobe in form of upturned hook.
9. Margin of eighth sternite simple, or produced in a subgenital plate. Ninth
sternite sometimes produced.
Spaniocerca tasmanica Tillyard
Fics. 35-36
Spaniocerca tasmanica Tillyard, 1924, Trans. Roy. Soc. S. Aust. 48: 195, fig. 3.
The holotype female has been reduced to two anterior and one posterior wings by
the action of Anthrenus. The following descriptions have been made from a male and
a female in fluid.
Fic. 35. Spaniocerca tasmanica Till- Fic. 36. Spaniocerca tasmanica Tillyard. Genitalia 4, a,
yard, Wings. lateral; b, dorsal; c, ventral; d, genitalia 9, ventral.
$. Head dark brown, occiput orange-brown ; antennae piceous, finely annulated
with yellowish at sutures. Pronotum dark brown, quadrate, a little broader than
long, anterior transverse furrow deep. Meso- and metanota shining piceous. Legs
yellowish-brown to dark brown, femora with a conspicuous median ring of yellowish.
Wings shining smoky-brownish, pterostigma darker, a few inconspicuous paler areas,
especially distad of transverse cord and along cubitus of anterior wing. Venation
brown ; generally no cross-vein in pterostigma of either wing, or if present, indistinct ;
generally two medio-cubital cross-veins between M5 and the transverse cord.
Abdomen whitish except segments nine and ten, which are brown. Ninth tergite
produced on either side of centre in a flattened rounded lobe; between the lobes is
a transparent membranous area. Tenth tergite also with a membranous area at
centre of apical margin, supra-anal lobe in form of a short broad hook, upcurved from
a broad base. Cerci short, stout, lightly pigmented, clothed with short hairs. Ninth
sternite with a strong ventral lobe, from the side dilating triangularly to a truncate
apex ; from beneath quadrate, slightly constricted at its base, apex faintly excised.
pee Pt a Ry
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 87
From the side, the sternite is produced, its apex bent sharply upward and dilated.
From beneath it is slightly narrowed about midway, with an elliptical apex. From
the upper apical margin arises a pair of small, scythe-like hooks, probably reduced
titillators.
Q. Similar, larger. Wings with more definite hyaline areas. Abdomen whitish,
seventh sternite, eighth and ninth segments brownish. Neither seventh nor eighth
sternites produced, margins straight, both with hyaline areas at base. Sub-anal
plates triangular, apices rounded, upper margins shallowly excised. Cerci short and
stout.
Length of fore wing, § 7mm., 2 9:4 mm.
Length of hind wing, $ 5-3 mm., 2 8 mm.
The holotype was taken at Mt. Wellington, TASMANIA, 31.i.1917 (R. J. Tillyard).
Other Tasmanian examples in the British Museum from Sheffield, 8.1.1917 (R. J.
Tillyard), Hobart, 15.x.1916 (C. E. Cole), 4.xi.1916 (G. H. Hardy), Penstock Lagoon,
29.1.1933, and Gouldt County, I0.ii.1933 (R. J. Tillyard). From the mainland of
AusTRALIA I have seen examples from New South Wales, Mt. Kosciusko, 24.xi.1921,
3-9.xii.1921 ; South Australia, Cudlee Creek, 20.xi.1934 (R. J. Tillyard), and Adelaide.
Spaniocerca tillyardi sp.n.
FIG. 37
General appearance similar to S. tasmanica; there appear to be two forms of this
species, one of which is smaller and paler than the other.
g. Head dark brown, occiput a little paler; antennae piceous, with fine pale
annulations at sutures. Pronotum dark brown, quadrate, a little broader than long,
anterior margin somewhat convex, anterior transverse furrow deep. Meso- and
metanota shining dark brown. Legs as in ¢tasmanica. Wings rather narrower, shining
smoky-brownish, with a hyaline area just beyond the transverse cord, and a larger
one in the neighbourhood of Cur. Venation dark brown, no cross-vein in pterostigma
of either wing; one or two medio-cubital cross-veins between M5 and the trans-
verse cord.
Abdominal segments pale or light brownish, apical margins whitish, segments
eight to ten darker. Ninth tergite produced on either side in a flattened lobe, apices
approaching one another calliper-wise. Between them the margin is excised and
membranous. Tenth tergite also with a membranous area at the centre of its apical
margin, giving the appearance of an excision. Supra-anal lobe broad and triangular
at its base, a membranous area at its base separating two rounded hairy elevations.
From the apex arises a slender hook, in side view with a rounded excision on dorsal
surface near base. Cerci longer than in fasmanica, from the side angled upwards in the
_ form of a foot. Ninth sternite long, more slender than in ¢asmanica, not dilated at its
apex, which terminates in a pair of short incurving fingers. From its upper surface
arises a pair of flattened titillators, curving upwards and basally, then downwards
and diverging apically. These titillators are broad at their bases, apical two-thirds
narrow and blade-like. Ventral lobe much narrower in ventral aspect, slightly con-
stricted in its apical half. From the side, it is parallel-sided, apex truncate.
88 A REVISION OF THE AUSTRALIAN AND TASMANIAN
2. Similar, except that abdominal segments one to seven are whitish. Eighth
sternite slightly produced, centre of apical margin still further produced in a small,
triangular lobe; this lobe is more pigmented than the remainder of the margin, and
is connected to the main part of the segment by a narrow band of pigmentation.
Ninth sternite strongly produced apically in an acute triangle, a narrow triangular
membranous area at its centre from base to apex. In side view, this sternite resembles
Fic. 37. Spaniocerca tillyardi sp.n. Genitalia 3, a, lateral; b,
dorsal; c, ventral; d, genitalia 9, lateral; e, ventral.
the valves of a short ovipositor. Cerci short, sub-anal plates subtriangular, apices
rounded.
Length of fore wing, g 5-8-6-6 mm., 2 6-3-7 mm.
Length of hind wing, $ 4-9-5°8 mm., 2 5-2-6 mm.
AUSTRALIA: F.C.T., Lee’s Springs, xi.1932 (R. J. Tillyard).
Type male, paratype female in form of microscope preparations, remainder in
2 per cent. formaldehyde solution.
Spaniocerca bullata sp.n.
Fic. 38
$ (in fluid). Head shining piceous, antennae piceous, finely annulated with yellow- —
ish at the sutures. Pronotum brown, quadrate, a little broader than long, angles
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 89
rounded, anterior transverse furrow deep, its ends curving backward. Meso- and
metanota shining piceous. Legs dark brown, tibiae with a yellowish ring in basal half,
below the knee. Wings pale smoky-brownish (anterior the darker), pterostigma
darker, and a darker area over the transverse cord. Venation brown; no cross-vein
in pterostigma; one to three medio-cubital cross-veins between M5 and trans-
verse cord.
Abdomen pale brownish, segments nine and ten darker. Ninth tergite not pro-
duced. Ninth sternite produced in a large subgenital plate, broad from beneath,
sides sinuous, apex rounded. From the side it is rather narrow, apex roundly trun-
cate, and from the upper margins towards the apex arise a pair of slender arms,
Fic. 38. Spaniocerca bullata sp.n. Genitalia 3, a, lateral; b, dorsal;
c, subgenital plate, ventral; d, apex of the same from behind; e,
genitalia 9, ventral.
curving tailward and inward. From behind, a membranous finger is seen between
the incurving arms. Also towards the apex there arises from the upper surface a pair
of flattened titillators, bent basally about midway, and in the type resting within the
tenth tergite ; in the paratypes these titillators are bent outwards at right angles to
the line of the body. Ventral lobe short and deep from the side, rather narrow from
beneath. Tenth tergite with a V-shaped excision at the centre of its apical margin.
Supra-anal lobe trapezoidal at its base with four elevated knobs on its upper surface ;
the apex is produced upwards at right angles in a broad, rather thin lobe with a
rounded apex. Cerci short, stout, slightly angled near the base in side view, tapering
to a rounded apex. At the base is a small narrow sclerite.
9. Similar to male, larger, wings with a faint indication of a cross-vein at base of
pterostigma. Genitalia following the pattern of S. tillyardi. Eighth sternite more
strongly produced at its centre in a slender, slightly clavate lobe. The whole sternite
evenly pigmented. Ninth sternite triangularly produced, blade-like from the side,
definitely narrower than the eighth sternite. Sub-anal plates broad, from beneath
outer margin concave, apex rounded. Cerci short, somewhat ovate from beneath.
Length of fore wing, ¢ 8 mm., 2 9 mm.
go A REVISION OF THE AUSTRALIAN AND TASMANIAN
TASMANIA: no other data (J. W. Evans), 3 3, 4 9.
The specific name bullata refers to the four knobs on the supra-anal lobe. Type
male, paratype female in form of microscope preparations, remaining paratypes in
2 per cent. formaldehyde solution. The female is associated on general appearance
and on similarity of genital structure with S. ¢illyardi, with which species the male
shows more affinity than with S. tasmanica. The form of the supra-anal lobe and
subgenital plate of the male distinguish it from S. tillyardi.
UDAMOCERCIA Enderlein
Udamocercia Enderlein, 1909, Zool. Anz. 84: 418.
Type species: Leuctra antarctica Enderlein, 1905 (Monotypic genus).
Wings similar to Spaniocerca, but without excision at apex of fourth anal vein in
the hind wing. Anal fan narrower. Cuz in fore wing relatively longer.
36. Ninth tergite not produced ; ninth sternite produced, long, slender ventral lobe
at base. Titillators slender, upcurved, arising from membrane below tenth tergite,
not from apex of the ninth sternite. Supra-anal lobe either in the form of an upwardly
directed hook arising from a broad base, or downwardly directed and bifurcate. Cerci
one-segmented, short, quadrate, or bent inward.
2. In the South American species the eighth sternite is produced in a long sword-
like ovipositor. I have seen but one Tasmanian example which I believe to belong to
this genus. In it the eighth sternite is produced and elevated in a short triangular
subgenital plate, notched at its apex. The ninth sternite is strongly chitinized and its
apical margin triangularly produced. From general appearance of the wings, this
female appears closer to U. albomacula than to U. bifasciata, but as it bears no other
data than Tasmania, I am not assigning it to either of these species.
Enderlein originally asssociated the type species with Leuctra, but it is more
Nemourid than Leuctrid. It has five anal veins in the hind wing, whereas the Leuctri-
dae have only three, and M3+-4 is not partially fused with Cur, as is the case with
Leuctra, which thus has an apparent fork to Cur, and an apparently unforked medius
in the hind wing.
Udamocercia albomacula sp.n.
FIGS. 39, 40 a-c
3 (dried). General colour piceous, shining. Head broader than pronotum ; antennae
slender, shorter than fore wing. Pronotum broader than long, broadest before the
middle, side margins definitely angled at broadest part; a distinct, anterior trans-
verse impressed line. Legs dull yellowish-brown, with darker markings at knees, base
of femur, and apex of tibia. Wings light smoky-brown with paler areas at centre of
pterostigma, distad of transverse cord, between branches of cubitus, and near base of
fore wing. Venation brownish. Abdomen brownish. Ninth tergite not produced,
sternite narrow, apical margin produced in a slender tongue, at least as long as sternite,
curving upward from the side. Ventral process slender, pointed apically. Tenth
tergite with an excision of its basal margin from above, apical margin produced,
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA) 91
rounded, its centre curling downward, and clothed in a dense mat of short black setae.
From beneath it arises the supra-anal lobe, whose base is a broad rounded plate. Its
apical margin is extended upwards as a
slender hook, on each side of which is a small
rounded lobe. Titillators two, slender, up-
curved, membranous on upper surfaces, very
flexibly attached to the membrane below the
tenth tergite. Cerci short, stout, quadrate,
with a chitinous plate at base.
2 unknown.
Length of fore wing 5-8 mm., of hind wing
4°9 mm.
Tasmania: Cradle Mountain, 16.i.1917, Fic. 39. Udamocercia albomacula sp.n.
I g(R. J. Tillyard). Wings.
Type pinned, with the abdomen mounted in balsam.
Udamocercia bifasciata sp.n.
Fic. 40 d—f
3 (in fluid). Head shining piceous, palpi and basal segments of antennae fuscous
(remaining segments lacking in type) ; pronotum fuscous, slightly broader than long,
anterior transverse furrow distinct ; angles rounded, lateral margins slightly convex.
Meso- and metanota fuscous. (Legs lacking in type.) Wings light smoky-brown,
anterior darker than posterior, and with two distinct white transverse fasciae, the
Fic. 40. Udamocercia spp.n. g genitalia. a—c, U. albomacula; d-f, U. bifasciata. a,
lateral; b, dorsal; c, subgenital plate, ventral; d, lateral; e, dorsal; f, subgenital
plate, ventral.
outer straight, at the level of the base of the pterostigma, the inner curved, concave
side towards the apex of the wing, running from apex of Sc basad of transverse cord
nearly to apex of Cuz. Venation brownish, a faint cross-vein at base of pterostigma
in both wings. In posterior wing R4++5 separates from R2+3 beyond the transverse
cord. Abdomen lightly chitinized, segments eight to ten brownish, but membranous
-dorsally. Ninth tergite rather narrow above but longer at its lateral margins. Ninth
92 A REVISION OF THE AUSTRALIAN AND TASMANIAN
sternite produced in a long slender subgenital plate, acute at its apex, and at first
glance resembling an ovipositor. Ventral lobe slender, about as long as lateral
margins of ninth tergite. Attached to the membranous upper surface of the sub-
genital plate towards its base are a pair of elongate titillators, fused throughout most
of their length, apex shallowly emarginate, witha deep narrow excision at its centre.
Tenth tergite small, strongly chitinized at the sides, and bearing a pair of strong,
inwardly hooked cerci whose bases only are visible from the side, appearing as small
lobes. Supra-anal lobe directed obliquely downward, deeply forked in dorsal aspect,
each branch from the side seen as a slender process with a dilated and hooked apex,
placed on either side of the subgenital plate.
Length of fore wing, 5 mm.
TASMANIA: no other data, 1 ¢ (J. W. Evans).
Type in form of microscope preparation. At some future date it may be necessary
to erect a new genus to contain this species, but for the present, in view of the scanty
material available, it is proposed to place it in Udamocercia.
APPENDIX
This paper, though virtually completed in 1941, could not be published under |
war conditions and was revised in 1948, following the receipt of more material.
Further delays in publication ensued, and since it went to press I have received
from Dr. W. E. Ricker (April 1951) a reprint of his paper ‘Some Evolutionary Trends
in Plecoptera’ (1950, Proc. Indiana Acad. Sci. 59: 197-209) in which he proposes
a new subfamily name NOTONEMOURINAE (type genus Notonemoura Tillyard) and in-
cludes also the genera Spaniocerca Tillyard and Spantocercoides Kimmins. My con-
ception of this subfamily is wider, embracing the genera enumerated on pp. 84-85.
Nevertheless I have adopted Dr. Ricker’s subfamily name, but allowed my original
definition to stand. D. Eee
;
_Megaleptoperla .
albomaculata, Udamocercia
antarctica, Leuctra
antarctica, Udamocercia
Antarctoperla
Aphanicerca
Aphanicercella
Aphanicercopsis .
Aucklandobius .
australica, Leptoperla
australica, Pavanotoperla
australis, Trinotoperla
Austroperla
beroé, Leptoperla
bifasciata, Udamocercia
brevipennis, Dinotoperla
bullata, Spaniocerca
carpenteri, Dinotoperla
Desmonemoura
Dinotoperla :
> sp. nymph
Eunotoperla
Eusthenia .
evansi, Dinotoperla
exigua, Leptoperla
fasciata, Dinotoperla .
fontana, Dinotoperla .
fusca, Dinotoperla
_GRIPOPTERYGIDAE
Diagnosis
Key to genera
Gripopteryx
irrorata, Trinotoperla .
_kershawi, Eunotoperla
Leptoperla ,
‘9 spp. nymphs
LEPTOPERLIDAE .
Leptoperlini
-LEUCTRIDAE .
PRESENTED
GRIPOPTERYGIDAE AND NEMOURIDAE (PLECOPTERA)
INDEX
Synonyms printed in italics.
47, 62
minor, Trinotoperla
nymph .
a2? ”?
NEMOURIDAE
Neonemura :
Nesoperla . , :
nigricoxa, Dinotoperla
nigrifrons, Leptoperla .
nivata, Trinotoperla
Notonemoura
Notonemourinae
Notoperla .
Nymphs
Dinotoperla
Leptoperla
Trinotoperla .
opposita, Chloroperla .
opposita, Dinotoperla .
opposita, Leptoperla
opposita, Perla
Paragripopteryx
Paranotoperla
pirioni, Nemura .
reticulata, Leptoperla .
rugosa, Leptoperla
serricauda, Dinotoperla
Spaniocerca
Spaniocercoides .
Stenoperla.
tasmanica, Leptoperla
tasmanica, Spaniocerca
thwaitesi, Dinotoperla
tillyardi, Spaniocerca .
Trinotoperla
9 spp. nymphs
Udamocercia :
uniformis, Dinotoperla
varia, Leptoperla
zelandica, Spaniocerca
Zelandobius
Zelandoperla
93
BY
- CHARLES BATEY ~ .
PRINTER
TO THE
UNIVERSITY
tex =} 17 DEC. 198
SPHECIDAE AND POMPILIDAE
(HYMENOPTERA)
COLLECTED BY MR. K. M. GUICHARD
IN WEST AFRICA AND ETHIOPIA
G. ARNOLD
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 2 No. 3
LONDON : 1951
SPHECIDAE AND POMPILIDAE (HYMENOPTERA)
COLLECTED BY MR. K. M. GUICHARD
IN WEST AFRICA AND ETHIOPIA
1941-1948
BY
G. ARNOLD
National Museum of Southern Rhodesia
Bulawayo
A MON
Pp. 95-183; Pls. 4-5; 65 Text-figures
BULLETIN OF
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LONDON : 1951
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is issued
in five series, corresponding to the Departments of the
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SPHECIDAE AND POMPILIDAE (HYMENOPTERA)
COLLECTED BY MR. K. M. GUICHARD IN WEST
AFRICA AND ETHIOPIA, 1941-1948
By G. ARNOLD
PART I. SPHECIDAE
THIs paper is based on a collection of nearly 700 specimens which Mr. R. B. Benson
of the British Museum has submitted to me for study. It is of considerable interest,
since it is derived from regions from which there have been few new records within at
least the last fifty years. The great majority of the species from West Africa known
up to the date of Dalla Torre’s Catalogus Hymenopterorum, 1897, were made known
chiefly by F. Smith and F. F. Kohl, and after that date by R. E. Turner. The pro-
portion of new species is considerable, especially of the Cercerinae. On the other hand,
the tribes Miscophini, Psenini, and Pemphredonini, of which the members are small
or very small, are barely represented.
In the following list of species I have given only the original reference, which in
many cases applies to only one sex. For descriptions of the other sex, if known, the
reader should refer to my monograph of the African Sphecidae published in the
Annals of the Transvaal Museum, 9-14, 1922-1931, and to several papers published
in the Occasional Papers of the National Museum of Southern Rhodesia.
Types of new species are in the British Museum.
The least magnification required to resolve the sculpture with a stereoscopic
binocular microscope is indicated in brackets, e.g. (40 diameters).
Subfamily TRYPOXYLONINAE
; Pison xanthopus Brullé
1833, Ann. Soc. Ent. Fr. 2: 408.
Kpeve, Gold Coast, June 1942, I ¢.
Pison allonymum Schulz
1906, Spolia Hymen.: 213.
Haramaia, Ethiopia, June 1948, 1 9.
Pison montanum Cameron
1910, Sjéstedt’s Kilimandjaro-Meru Exped. 8: 289.
Dilla (Sidamo) Ethiopia, June 1948, 2 33.
Trypoxylon viduum sp. n.
(Fics. I, Ia-c)
g. 11 mm. long. Black. Wings hyaline, slightly smoky beyond the cells. Clypeus
98 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
and face up to the scutate area, and the temples with silvery, the occiput with short
and black pubescence. Thorax with the usual whitish and erect pubescence, nowhere
dense. Clypeus twice as wide in front as long, the anterior margin feebly convex.
Facial shield entire, slightly longer than wide across the lateral angles, from which is
emitted a carina extending over the inner half of the ocular sinuses. The shield is
closely punctured, the punctures largest at the sides, the interspaces about as wide
as the punctures. Behind the scutate area but not reaching the posterior ocelli, the.
Fic. 1. Trypoxylon viduum 3, head, x15. Fic, 1a. Trypoxylon viduum 6g, antenna, X15.
Fic. 1b. Trypoxylon viduum g, 8th sternite, x27. Fic. 1c. Trypoxylon viduum 4, genitalia, x 27.
Fic. 2. Trypoxylon acutangulum 9°, head, x15. Fic. 2a. Trypoxylon acutangulum 9, epinotum
and petiole, x15. Fic. 3. Philanthus ugandicus 4, genitalia, x21. Fic. 4. Philanthus histrio 4,
genitalia, h; Philanthus histrio, race distinctus, d, X 24.
puncturation is larger and deeper at the sides. Second joint of the flagellum twice as
long as wide, as long as the third, the apical joint acute, four times longer than wide
at the base and nearly as long as the four proceeding joints united. Interocular
distance on the vertex equal to the length of the first three joints of the flagellum
plus half of the fourth. Posterior ocelli as far from the eyes as from each other.
Pronotum and mesonotum nitidulous, the former with sparse punctures and rounded
shoulders, the mesonotum sparsely punctured in the middle but more closely at the
sides, the punctures about as large as those on the facial shield. Mesopleura shining,
a little more finely punctured than the mesonotum. (Scutellum and metanotum
obscured by the pin.) The U-shaped area of the epinotum obliquely rugose over the
basal third, the median groove wide and transversely costate, the rest of the area
finely rugose, the sides of the epinotal dorsum and the declivity very coarsely trans-
versely rugose and with anastomoses. The sides of the epinotum shining, vertically
and not closely striate. Petiole slender, fully six and a half times longer than wide
aw)
n 4350
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 99
across the node, and as long as the second, third, and half the fourth tergites united.
The second tergite is as long as the third, and not quite twice as long as wide behind.
Second abscissa of the radius twice as long as the first. The eighth sternite resembles
that of T. abditum Arnold, but differs in having three stout bristles emitted from each
arm ; the genitalia are like those of I’. cognatum Arnold but the outer paramera are
narrower.
Jowaha, N. of Debra Sina, 6,000 ft., Ethiopia, 1 3.
This could be the ¢ of T. latiscutatum Arnold, but I hesitate to ascribe it to that
species in the absence of females from the same locality.
Trypoxylon acutangulum sp. n.
(Fics. 2, 2a)
®. gmm. long. Black. Tarsi dark brown, the apical joint of the hind pair ochreous.
Wings hyaline, the veins black, the costa brownish-yellow at the base. Clypeus and
lower half of the face with silvery pubescence, the thorax with sparse whitish pubes-
cence. Facial shield and ocular sinuses dull, closely and very finely punctured
(30 diameters), the vertex in front of and between the ocelli less closely punctured
and with a row of larger punctures behind the shield. Anterior margin of the clypeus
convex. Facial shield entire, narrow, three-fourths longer than wide across the lateral
angles, the apical angle acute. A faint trace of a carina is emitted from the angles
into the ocular sinuses. Interocular distance on the vertex equal to the length of the
first two joints of the flagellum plus two-thirds of the third joint. Posterior ocelli
almost contiguous with the eyes. Second joint of the flagellum as long as the third.
Shoulders of the pronotum rounded. Mesonotum and scutellum shining, very finely
punctured, the interspaces fully twice as wide as the punctures on the sides of the
mesonotum, wider on the disk. Mesopleura more finely punctured and more sparsely
than the mesonotum. Median area of the epinotal dorsum U-shaped, one-fourth
wider at the base than long, shallowly grooved down the middle, reticulate-rugose
at the base, transversely and finely rugose elsewhere, the rugae extending over the
lateral areas of the dorsum. Declivity transversely rugose and finely punctured, and
with a deep median longitudinal impression. There is a deep pit at the brow of the
declivity. Sides of the epinotum shining, finely transversely striate. Petiole four and
a half times longer than wide behind, as long as the second tergite plus two-thirds of
_ the third. Second abscissa of the radius two and a half times longer than the first.
Niamey, French West Africa, August 1944, I 9.
Remotely related to T. Rohit Arnold, but smaller, with a narrower facial shield and
a shorter petiole.
Subfamily PHILANTHINAE
Philanthus coarctatus Spinola
1838, Ann. Soc. Ent. Fr.'7: 486.
Ufdem, Ethiopia, August 1944, I 3.
100 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Philanthus rubidus Arnold
P. abyssinicus Arnold, 1932, Occ. Pap. Rhod. Mus. 1: 7.
P. rubidus Arnold, 1946, Occ. Pap. Nat. Mus. S. Rhod. 12: 82.
River Errer, near Harar, Ethiopia, May 1948, 1 .
Philanthus ugandicus Magretti
1908, Bull. Mus. Hist. Nat., Paris, 14: 188.
Adis Ababa, Dessie, and Zuquala (9,000 ft.), Ethiopia, 7 gg.
Philanthus histrio distinctus subsp. n.
(Fic. 4d)
P. histrio Fabricius, 1804, Syst. Piez.: 301. -
g. 12 mm. long. Black, the occiput slightly rufescent. The pale markings on the
head and abdomen are similar to those of histrio i. sp. but the colour is creamy white
instead of yellow, the clypeus has a rectangular black macula in the middle of the
anterior margin, and the maculae on the second tergite are larger and not triangular,
but nearly round.
The thorax is eritirely black. Fore legs brown, middle and hind femora black, their
tibiae and tarsi brownish, the former with the apical third whitish on the outside, and
also the first four tarsal joints. Puncturation on the vertex behind the ocelli larger
and wider apart than in the type of the species ; the puncturation of the mesonotum
also a little coarser, otherwise the sculpture is like that of the type of the species.
The second joint of the flagellum is thinner, being nearly four times longer than the
third. The genitalia differ slightly, the apical third of the outer paramera being
narrower and more acute at the apex. (Fig. 4h, genitalia of histrio i. sp.)
Debra Sina, Ethiopia, November 1945.
Philanthus loefflingi Dahlbom, var. meneliki Arnold
1925, Ann. Transv. Mus. 11: 150.
Bishoftu; Ethiopia, June 1948, 1 3.
Philanthus dichrous dolosus Kohl, var. abyssinicus Arnold
1925, Ann. Transv. Mus. 11: 162.
Goré, Ethiopia, February 1948, 1 g, 2 29. ’
In these specimens the thorax is entirely black, as are also the middle and hind legs. —
Philanthus fuscipennis consimilis Kohl
1891, Ann. naturh. Hofmus. Wien, 6: 349.
Bishoftu and Jowaha, Ethiopia, 3 gg. These three are a slight variety in which —
the hind tibiae are yellow. In one of the specimens from Jowaha the scutellum and
postscutellum are also yellow.
Subfamily CERCERINAE
In Plates 4 and 5 the male genitalia of the majority of the species described here-
under are illustrated. For figures of other species included in the Guichard collection
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA IO
and which are also found in the southern half of the continent the reader should refer
to my paper on Cerceris published in the Journal of the Entomological Society of
Southern Africa, 5, 1942. For the sections into which I have separated the
females, the chapter on the genus in my monograph of the African Sphecidae, in the
Annals of the Transvaal Museum, 14, part 2, should be consulted. These sections
are solely for the purpose of facilitating the identification of the species and are not
natural, as can be seen by a comparison of the genitalia of the corresponding males.
For example, the genitalia of species in which the females have such distinctive
characters as a porrect clypeus lamina, sections B and C, or a median tubercle,
section E, can differ considerably.
I am unable to recognize the validity of the subgenus A piraptrix Shestakov, which
is based on the presence of a basal platform on the second sternite. In some species
there is a more or less distinct median carina on that segment which is sometimes
flattened cephalad. The basal platform appears to have arisen by a gradual reduction
of the carina, but the platform is by no means always distinctly defined. In fact an
examination of a fair proportion of the species shows that there is a gradation from
a raised area with well-defined margins to one which is barely recognizable. In one
case, C. varicincta Cameron, this variability is exhibited within -the species.
Cerceris longiuscula sp. n.
(Fics. 5, 5a; PL. 4, FIG. 1)
g$. 14mm. long. Black. Apical margin of the first tergite and all of the following
tergite pale burnt umber, the second and third paler than the rest. Sternites 2-7 dull
_ yellow, stained with brown over their apical halves. Apical third of the fore and
middle tibiae and all the tarsi, pale reddish-brown. The following parts are creamy
white: clypeus excepting the black anterior third, sides of the face up to the level of
the antennal sockets, lower half of the frontal triangle, a small round spot at the top
of the temples, a small transverse spot on the pronotal shoulders, a triangular spot
on the mesopleura below the tegulae, and a transverse spot on each side of the
scutellum.
Wings pale fuscous. Clypeus dull, very finely rugulose, and with a few shallow
punctures superimposed ; the lateral sclerites with sparse, decumbent, and yellow
pubescence. The rest of the head dull, with a microscopic and dense puncturation,
on which is superimposed a larger and deeper one which becomes finer dorsad, the
interspaces a little wider than the punctures. Between the ocellar area and the
antennal sockets there are also a few longitudinal rugae. The frontal carina is high
and acute. Thorax dull. Anterior face of the pronotum almost smooth, the dorsal
face finely and very sparsely punctured. The puncturation of the mesonotum and
scutellum a little larger and closer than that of the face, that of the mesopleura much
larger and closer, reticulate punctate; the metapleura are costate except over the
middle third. The epinotum, excepting the triangular area and the somewhat shining
middle lower half of the declivity, has a puncturation about twice as large as that of
the mesonotum, but less close ; the triangular area is obliquely costate, eight costae
on each side. Tergites microscopically reticulate and dull, with a very sparse, small,
102 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
and shallow puncturation on the second to sixth tergites, the first tergite with a
sparse and large one.
Pygidial area coarsely punctured, rectangular, twice as long as wide, the apical
margin straight. Sternites shining, shallowly punctured, the sixth with a small tooth
on each side near the apical margin. Median area of the clypeus long, about half as
long again as wide, produced well beyond the lateral sclerites, shallowly grooved in
the middle of the apical half, the apical margin feebly concave. Inner orbits feebly
divergent below, the interocular distance across the clypeus very little greater than
across the vertex. Posterior ocelli twice as far from the eyes as from each other; the
interocular distance on the vertex equal to the length of the first four joints of the
flagellum. The second joint is twice as long as wide at the apex and one-fourth longer
than the third, the apical joint twice as long as wide at the base and obliquely
truncate.
Dorsum of the pronotum not short, as long as the third joint of the flagellum.
Mesonotum wide, four-fifths wider than long. First tergite nearly twice as wide
behind as long, the second two-thirds longer than the first and three-fourths wider
behind than long. Posterior tibiae with six serrations.
Kpeve, Gold Coast, June 1942, I 3.
Not related to any other African species known to me. The outer paramera of the
genitalia are distinctive, being produced into an acute tooth on the inside not far
from the apex.
Cerceris pallida Arnold, var. lutulenta var. n.
C. pallida Arnold, 1935, Bull. Soc. Sci. Nat. Maroc, 15: 5.
2. 15 mm. long. This is a variety in which the colour of the head and thorax is
dull ochreous, the mesothorax pale ferruginous, with the margins of the mesopleura
and the middle of the epinotal declivity blackish. The tergites are ochreous, the first
four black at the extreme base and apex. Legs reddish-yellow.
Boghé, Mauritania, September 1943, I 9.
Cerceris moggionis sp. n.
(Fics. 6, 6a and 60; PL. 4, FIG. 2)
g. 13 mm. long. A streak on the upper half of the median area of the clypeus and
the posterior margins of the lateral sclerites straw-yellow, the rest of the clypeus
brown. Sides of the face, frontal triangle, and the frontal carina also pale yellow, the
vertex reddish-brown at the sides, the rest of the head black. Antennae ferruginous,
the last three joints black. Pronotum dark ferruginous, blackish in the middle and ~
at the lower margins of the sides, the tegulae ferruginous, the rest of the thorax black.
Abdomen pale ferruginous or yellowish-red, the apical half of the first tergite pale
yellow, a median triangular area on the second sternite and the basal halves of the
third to sixth sternites, blackish. Legs pale ferruginous. A small elongate spot on the { |
outside of the apical half of the anterior femora, the apical two-thirds of the underside —
of the middle and hind femora, the upper side of the hind coxae and the upper side
of the middle and hind tibiae, pale yellow.
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 103
Wings pale fuscous, with a violaceous tint, darkest on the costal margin and radial
cell. Head with a decumbent silvery pubescence, less dense on the vertex than else-
where. Epinotum with exserted, fairly long and white pilosity. Abdomen with short,
sparse, and yellowish pubescence. Clypeus dull and smooth, the rest of the head and
the dorsum of the pronotum reticulate-punctate, the punctures about one-third as
wide as the anterior ocellus. Anterior face of the pronotum shallowly punctured, the
sides transversely rugose. Metapleura transversely costate above and below, the rest
Fic. 5. Cerceris guichardi 4, first two tergites, x6. Fig. 5a. Cerceris guichardi 3, head, Xx Io.
Fic. 6. Cerceris moggionis g, clypeus, X10. Fic. 6a. Cerceris moggionis 4, first two tergites, x Io.
Fic. 7. Cerceris insignita g, abdomen, x6 (stippled parts ferruginous). Fic. 8. Cerceris flavo-
nasuta 9, head, x10. Fic. 8a. Cerceris flavonasuta 9, first two tergites, x 10. Fic. 8b. Cerceris
flavonasuta 9, pygidium, x10. 8c. Cerceris flavonasuta 9, first two tergites, x Io.
of the thorax strongly reticulate-punctate, the punctures on the mesothorax a little
larger and deeper than those on the head. Median area of the epinotum with six
large transverse costae. Tergites 1-5 not quite dull, with fairly large punctures, the
interspaces for the greater part twice as wide as the punctures, the sixth finely
punctured and dull, the pygidial area nearly twice as long as wide, the lateral margins
parallel over the basal half, convex outwardly on the apical half, the apical margin
convex. Sternites 1-6 shining, shallowly and sparsely punctured, the sixth with a
yellow fimbria on its hind margin and a triangular tooth on each side. Median area
of the clypeus oval, feebly convex, nearly half as long again as wide, the apical
margin with three large and rounded teeth. Inner orbits parallel. Vertex wide, the
interocular distance there equal to the length of the first five joints of the flagellum.
Posterior ocelli twice as far from the eyes as from each other. Second joint of the
flagellum a little more than twice as long as wide, two-fifths longer than the third,
the apical joint half as long again as wide, transversely truncate at the apex. Shoulders
ENTOM. 2. 3 M
104 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
of the pronotum angular, raised to form a subcarinate tubercle. Mesopleura with a
small tooth below. Scutellum depressed lengthwise in the middle. Epinotal dorsum
short, about as long as the scutellum. First tergite three-eighths wider behind than
long, second tergite seven-ninths wider than long. Hind tibiae with five serrations,
Moggio, Ethiopia, April 1948, 1 g.
Not related to any other Ethiopian species. The tuberculate pronotal shoulders
and the shape of the outer paramera of the genitalia are distinctive. The outer
paramera have on the inside, a little behind the middle of their length, a lamelliform
fimbriated lobe, which can be seen only when those parts are widely splayed out.
Cerceris insignita sp. n.
(Fic. 7; PL. 4, FIG. 3)
3g. 11 mm. long. Black. Antennae, occiput, posterior half of the temples, pro-
notum, tegulae, scutellum, metanotum, sixth and seventh abdominal segments, and
the legs, ferruginous. Mandibles ferruginous, last three joints of the flagellum blackish
above, median area of the clypeus, sides of the face, frontal triangle, and carina,
yellowish-white ; the lateral sclerites of the clypeus pale brown and with a fine yellow-
ish pubescence, the moustache brown. Tergites 2-5 with a narrow pre-apical pale
yellow band which is abruptly dilated laterad, the dilatation almost reaching the
base, the sixth with a yellow spot on each side. Sternites fusco-ferruginous. Wings
hyaline, the veins brown, the pterostigma ochreous, the apical third of the forewing
pale fuscous. Clypeus and yellow areas of the face dull, shallowly, finely and sparsely
punctured. The rest of the head, and the thorax, reticulate-punctate and dull, the
punctures on the vertex about one-third as wide as the anterior ocelli, the punctures
on the mesonotum and scutellum a little larger, and largest on the mesopleura.
Triangular area of the epinotum slightly shining, closely and transversely rugose,
the middle third of the epinotal declivity shining and almost smooth. Tergites 1-6
dull, sparsely punctured, the punctures as large as those on the vertex. Pygidial area
shining, sparsely punctured, nearly parallel-sided but a little wider over the posterior
half, the apical margin convex. Sternites smooth and shining, the sixth with a curved
tooth on each side below which there is a bundle of ferruginous hairs; seventh
sternite arcuately emarginate at the apex, the lateral margins raised or carinate.
Median area of the clypeus convex, suboval, half as long again as wide, the apical
margin tridentate. Inner orbits parallel. Posterior ocelli twice as far from the eyes
as from each other. Interocular distance on the vertex equal to the length of the first
five joints of the flagellum. The second joint is twice as long as wide, one-fourth
longer than the third, the apical joint truncate. Dorsum of the pronotum short, as
long as the first joint of the flagellum, the shoulders subangular. First tergite as long
as wide behind, the second two-thirds wider behind than long. Hind tibiae with five
serrations.
Moggio, Ethiopia, April 1948, 1 g.
The colour pattern of the tergites is unlike that of any other Ethiopian species.
Although so different in colour from C. moggionis, it is allied to that species, but the
genitalia are different. The outer paramera are elongate triangular over the distal
-
——
<M
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 105
third and acute at the apex. Another specimen, also from Moggio, represents a
variety, opulenta var. n., which differs from the type as follows:
g. 12 mm. long. Occiput and temples black, the latter with a small ferruginous
macula on top. The yellow areas on the tergites are more extensive, so much so that
on the second tergite the black area is reduced to a transverse band at the base and
a small triangular pre-apical macula, its base caudad. On the third tergite the black
covers only the middle third of the segment, a little more on the fourth and fifth, but
on the latter the black merges caudad into ferruginous. The first tergite is a little
broader in proportion to its length, and the sculpture on the head and thorax is also
a little shallower.
Cerceris rufiscutis Cameron, var. decolorata Arnold
C. rufiscutis Cameron, 1910, Sjdéstedt’s Kilimandjaro-Meru Exped. 8: 278.
Arnold, 1931, Ann. Tvansv. Mus. 14: 182.
Haramaia, April 1948, 19, 1g; Lake Bishoftu, Ethiopia, August 1946, 1 9;
Moggio, April 1945, I g¢; Lekempti, Ethiopia, June 1946, 1 J.
Cerceris flavonasuta sp. n.
(Fics. 8, 8a—c; PL. 4, FIG. 9)
2. 125 mm. long. Head ferruginous, with a large macula extending from the
antennal sockets to a little distance behind the ocellar area, and the anterior half of
the temples black. Lamina of the clypeus pale yellow, its apical margin blackish ;
the median area below it black. Frontal carina pale yellow, mandibles fusco-ferru-
ginous, antennae ferruginous, the fifth and following joints of the flagellum fuscous
above. Thorax black, the metanotum pale yellow. Tergites 1-4 and the basal half
of the fifth black, the first to third with pre-apical yellow bands, narrowly interrupted
in the middle and moderately dilated laterad, the fourth with a very thin pre-apical
yellow band ; the sternites 1-4 black, the second and third with a large yellow macula
on each side, the fifth and the whole of the apical segment ferruginous. Antennae,
apices of the femora, tibiae, and tarsi ferruginous. The hind coxae with a pyriform
yellow macula on the upper side. Wings hyaline, tinged with dull yellow, the ptero-
stigma ochreous, the veins brown, the tegulae ferruginous.
Labrum wider than long, with a large tubercle on each side. Median area of the
clypeus below the lamina smooth and shining. The lamina is free to its base, as long
as wide, convex transversely and strongly so lengthwise, a little wider in front than
behind, shining, sparsely punctured and broadly carinate lengthwise in the middle,
the apical margin shallowly excised. Lateral sclerites of the clypeus shining and
sparsely punctured.
Head dull and reticulate-punctate, finely on the face, more strongly on the vertex,
with the meshes emphasized longitudinally, and more finely and closely on the
occiput and upper part of the temples; the underside of the head is longitudinally
rugose and fairly strongly punctured in between.
Thorax dull, pronotum closely and finely punctured, mesonotum and scutellum
longitudinally and finely rugose, punctate, the punctures on the middle part of the
106 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
mesonotum largest and not so large as on the upper part of the face. Metanotum
finely and sparsely punctured.
Mesopleura and epinotum closely rugulose-punctulate, with a shallow larger —
puncturation superimposed. Triangular area of the epinotum closely and obliquely
rugulose, finely punctured between the rugae. First tergite shining, with sparse and
large punctures. The second to fifth closely and finely punctured over the basal half,
more sparsely with larger punctures apically.
Pygidial area widened towards the straight apical margin, and half as long again
as wide there. Sternites shining, shallowly and sparsely punctured, the sixth deeply
and narrowly excised. Mandibles with a broad rounded tooth at the middle of the
inner margin. Interocular distance on the vertex equal to the length of the first six —
joints of the flagellum, the second joint five-eighths longer than the third. Ocelli half
as far again from the eyes as from each other. Temples wide, at their widest clearly
wider than the eyes. Pronotal collar short, the shoulders rounded. Mesopleura with
a feeble denticle below. First tergite as long as wide, the second three-fourths wider
than long. Posterior tibiae with seven spiniferous serrations.
g. 9°5 mm. long. Clypeus, frontal triangle and carina and lower two-thirds of the
face, excepting a black streak extending from the antennal sockets to the clypeus,
pale chrome yellow; the mandibles ochreous, black at the apex, the rest of the head
black.
Pronotal collar with a transverse yellow band, narrowly interrupted in the middle.
Apical margin of the fifth abdominal segment, and the whole of the sixth and seventh ~
ferruginous. Posterior coxae and trochanters pale yellow on the underside. Eighth
to eleventh joints of the flagellum infuscated on the upperside. Otherwise coloured
like the 9.
The sculpture is everywhere a good deal more coarse than in the 9. Yellow areas —
of the face and the clypeus dull, shallowly and sparsely punctured, the interspaces.
about three times wider than the punctures. Thorax dull, the abdomen nearly so.
_ Pronotal collar sparsely and shallowly punctured. Mesonotum and scutellum clearly —
longitudinally and finely rugose with a shallow and not close puncturation in between
the rugae. Mesopleura coarsely reticulate-punctate, the punctures about half as wide
as one of the posterior ocelli. Triangular area of the epinotum closely, obliquely, and
finely rugose, grooved down the middle; the rest of the epinotum with a large and ~
deep puncturation. The abdominal segments have a microscopic and close funda-
mental puncturation on which is superimposed a larger one, the punctures as large
as those on the epinotum, but more widely spaced, the interspaces being for the
greater part two and a half times wider than the punctures. Pygidial area slightly
widened towards the transverse apex, half as long again as wide, the sides slightly
convex. Median area of the clypeus slightly flattened, a little longer than wide, the
apex tridentate. Interocular distance on the vertex equal to the length of the first
six joints of the flagellum. Apical joint of the flagellum curved, twice as long as wide,
the apex transversely truncate. First tergite nearly one-fourth longer than wide.
Posterior tibiae with six serrations.
Aselle, Ethiopia, July 1947, 1 2; Lekempti, Ethiopia, May 1946, 2 3d.
This species belongs to the small section B in my key to the species (1931, Aum.
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 107
Transv. Mus. 14: 137), which also includes C. andersoni Turner. It differs from that
species and the other four species by the shape of the clypeal lamina and pygidial
area, and from proteles Brauns, bicolor Smith (Pl. 4, fig. 4), and pictiventris Ger-
staecker by the colour of the abdomen.
Cerceris fulviventris Guérin
(PL. 4, FIG. 4)
1844, Icon. végne anim. 7: 444.
C. bicolor Smith, 1856, Cat. Hymen. B.M.: 447.
Guiglia, 1948, Ann. Mus. Stor. Nat. Genova, 68: 187.
Kaolack, August 1943, I g; Kaffrine, August 1943, I g and I 9.
Guiglia’s redescription of fulviventris Guérin, based on the type, appears to me
sufficient evidence that bicolor Smith should sink in synonymy. The colour of the g
in regard to the yellow markings is variable. In the ¢ from Kaolack all the tergites
have a short yellow streak on each side at the hind margin. It should also be noted
that whereas in the 9 the epinotal triangle is transversely costate, in the gg that area
is coarsely reticulate-rugose. The ¢ is easily recognizable by the large semi-elliptical
lobe, directed ventrad, on the inner inferior margin of the outer paramera of the
genitalia.
Cerceris severini Kohl
1913, Rev. zool. afr. 3: 207.
Brauns, 1926, Ann. Transv. Mus. 11: 289.
Diaforabe, French Sudan, August 1944, 1 J, 1 2; Abbai Gorge, Ethiopia, October
1945, I 9; Niamey, Niger Colony, August 1944, 1 9; Tillembeya, River Niger, French
Sudan, September 1944, 4 gg. The specimen from Abbai Gorge has the first tergite,
excepting the apical margin, black, and third to fifth tergite infuscated in the middle.
It should be observed that there is an error in Dr. Brauns’s translation of the
original description, which runs ‘The fourth (i.e. third) abscissa of the radius is one
and a half times longer than the first three (i.e. two) united, which are very short.’
Kohl says: ‘Der Abschnitt der Radialader, welcher die 3 Cubitalzelle vorne be-
grenzt, ist sehr viel kiirzer als das folgende Stiick, welches mehr als 14 Mal so lang als
wie jenes ist.’
Cerceris zavattarii Guiglia
(PL. 4, FIG. 5)
1939, R. Accad. Ital. Miss. biol. Borana, 8 (2): 58.
I ascribe to this species, known to me only from the description, a specimen to
which I refrain from giving a varietal name, since it differs only in minor details and
greater size from one of the paratypes mentioned by Guiglia.
3g. 14 mm. long. The dark areas of the head and thorax are black, without any
ferruginous parts. The vertex has two small yellow spots behind the ocelli. The
median maculae on the tergites are shorter, not reaching the apical margin, and also
more semicircular, that on the first black and on the following fusco-ferruginous.
The interocular distance on the vertex is equal to a little more than the length of the
108 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
first four joints of the flagellum. The second joint of the flagellum is fairly slender,
or two and three-fifths longer than wide at the apex, the apical joint is short and
obliquely truncate. The dense exserted and yellow pilosity on the sternites, including
even the eighth, is a distinctive character.
Bishoftu, 7,000 ft., Ethiopia, May 1948.
This species bears a strong resemblance to C. mugax Arnold and like it has the
sternites densely pilose. It can be distinguished, however, by the very different
sculpture of the epinotal triangle, by the more slender antennae, the narrower first
tergite, and the shape of the outer paramera of the genitalia, which are more acute
at the apex and strongly dentate on the inner margin.
Cerceris rhodesiae Brauns
(PL. 4, FIG. 6)
1929, Ann. Transv. Mus. 11: 330.
Arnold, 1942, J. Ent. Soc. S. Afr. §: 20.
I reduced this species to the rank of a subspecies of C. monocera Kohl, but on re-
examination, and a more careful comparison of the genitalia, find that the latter
organs are not identical as I believed. Since the stipites or outer paramera are in
most cases curved, it is necessary in comparing them to arrange them in exactly the
same plane. As a result, it can be seen that though the genitalia are very similar,
in those of rhodesiae the dilatation or lobe on the inner margin of the outer paramera
is less abrupt, and the distal portion beyond it is longer and rounder at the apex.
Also the differences in the puncturation and in the colour pattern are too great to be
considered merely subspecific. Furthermore, another character which I overlooked
is to be seen in the radial cell. In rhodesiae it is truncate at the apex, whereas in
monocera the apex is bluntly angular and ends on the costal margin. For these reasons
rhodesiae must be restored to specific rank. A specimen in this collection represents
a subspecies, rhodesiae haramatae subsp. nov.
3g. 11mm. long. In colour this differs from the species in having pale yellow bands,
dilated laterad, on the second to sixth tergites, an elongate macula of the same colour
on each side of the epinotal declivity and the scutellum entirely black. The upper
sides of the femora are dark brown, almost black on the hind pair. The dorsum of
the thorax is moderately shining, the mesonotum more strongly rugose lengthwise,
and the punctures between the rugae deeper. The puncturation of the epinotum is
larger and deeper and less close, the interspaces smooth and fully as wide as the
punctures, whereas in rhodesiae i. sp. the interspaces form raised meshes.
The median area of the clypeus is shorter and does not project so far beyond the
lateral sclerites, it is one-fourth longer than wide, in rhodesiae very nearly one-half
longer than wide. The ocelli are one and three-quarter times farther from the eyes
than from each other, in rhodesiae one and a half. The first tergite is wider, or as
wide as long, and in rhodesiae one-fourth longer than wide. The genitalia barely
differ, the outer paramera being a little less widely dilated anteriorly and slightly
rounded at the apex.
Haramaia, Ethiopia, May 1948, 1 g.
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 109
Cerceris campsomeroides sp. n.
(PL. 4, FIG. 7)
g. 14mm. long. Black. Clypeus, excepting a wide anterior margin, sides of face,
frontal triangle and carina, wide apical bands moderately dilated laterad on the
second, third and fourth tergites and an apical band covering only the middle third
_ on the fifth tergite, chrome yellow. Fore and middle femora black, their apical fourth,
like the tibiae and tarsi dull yellow, the hind legs also of that colour, but the femora
more or less stained reddish on the apical half. Apical joint of the flagellum and
ochreous below. Wing hyaline, the veins reddish ochreous, the pterostigma pale
yellow, the apical margin of both wings faintly fuscous.
Head and thorax with a long, erect, yellowish-grey pilosity, the abdomen with a
shorter, oblique and sparser one, on the sternites somewhat denser on the apical
margins. Yellow area on the head dull, longitudinally rugulose and finely and fairly
closely punctured. The upper part of the face, excepting a small smooth area below
the anterior ocellus, closely punctured and rugulose, the vertex and upper part of the
temples less closely, with larger punctures which are about one-third as wide as one
of the posterior ocelli, the interspaces a little wider than the punctures. Lower half
of the temples strigose lenghtwise and closely punctured, the punctures smaller than
on the vertex. Sides of the pronotum coarsely wrinkled, the mesopleura obliquely
costate. The rest of the thorax strongly punctured, the largest punctures, on the
scutellum and metanotum, are half as wide as one of the posterior ocelli, the inter-
spaces wider than the punctures. Those on the pro- and mesonotum slightly smaller,
and a little closer, on the epinotum equally large but closer, or almost reticulate-
punctate. The triangular area on epinotal dorsum is dull, obliquely rugose and
punctured, its lateral margins not impressed and therefore rather indistinct. First
tergite with a puncturation like that of the epinotum, the second to fifth more strongly
punctured on the apical than on the basal half, the interspaces a little wider than the
punctures ; the sixth and seventh tergites sparsely and coarsely punctured, the pygi-
dial area a little narrower at the base than at the apex and fully twice as long as wide
at the base, the apical margin straight. Tumid areas of the sternites with a shallow
and fairly large puncturation. The whole body fairly dull. Median area of the clypeus
fairly flat, one-seventh longer than wide, the apical margin strongly tridentate.
Temples a little wider than the eyes. Posterior ocelli very little farther from the eyes
than from each other, the interocular distance on the vertex equal to the length of the
first five joints of the flagellum. The apical joint of the flagellum curved, excavated
below, the two preceding joints also concave below and feebly carinate lengthwise.
Dorsum of the pronotum sloping cephalad, as long in the middle as the third joint
of the flagellum, the shoulders rounded. First tergite one-fourth wider behind than
long, the second a little shorter than the first and fully twice as wide behind as long.
Hind tibiae with seven spiniferous serrations. Second abscissa of the radius two-
thirds longer than the first.
Accra, Gold Coast, 1941, I ¢.
Not closely related to any other Ethiopian species. In the colour and narrow
IIo SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
abdomen this insect has a superficial resemblance to the males of some species of the
Scoliid genus Campsomeris, e.g. C. clotho Saussure.
C. iniqua Kohl
1894, Ann. naturh. Hofmus. Wien, 9: 288.
Lekempti, 6,500 ft., Ethiopia, May 1946, 1 J.
Cerceris nobilitata walegae subsp. n.
C. nobilitata Cameron, 1905, Trans. S. Afr. Phil. Soc. 15: 216.
2. 10-5 mm. long. Larger than nobilitata, and differing from that as follows:
Dorsum of the pronotum raised on each side into a transverse pale yellow torus,
lIOb
lOa
Fics. 9 and 9a. Cerceris faceta 2, head, x12. Fic. 9b. Cerceris faceta, first two tergites, x 12.
Fic. 9c. Cerceris faceta, pygidium, x12. Fic. 10. Cerceris electra 9, head, x12. Fic. 10a.
Cerceris electra, first two tergites, x12. Fic. 11. Cerceris maia 3, pronotum, X12. FIG. 12,
Cerceris alcyone g, head, X12. Fic. 12a. Cerceris alcyone, first two tergites, x 12.
similar to that in C. osivis Arnold but larger. Metanotum pale yellow, hind margin
of mesopleura, the epinotum excepting the middle, and the first tergite and sternite
and the second sternite, castaneous red. Second tergite black, the macula at the base
yellow, not interrupted in the middle and larger. The apical band on the third tergite
and the apical macula on the fifth pale ochreous, instead of yellowish-white, and
larger. The puncturation of the abdomen is closer, the interspaces on the second to
fifth tergites being, for the greater part, narrower than the punctures (wider in the
type of the species). The first tergite is widest behind the middle, instead of at the
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA III
middle. The fore and middle femora are stained with black above, and the hind
femora are castaneous red. Otherwise like the type of the species.
Gambeila, 700 ft., in the territory of the Walega and Nuer tribes, Ethiopia,
February, 1948, I 9.
The yellowish parts of the head have been discoloured to reddish-yellow by
cyanide fumes.
In the presence of the tori on the pronotal dorsum this race resembles C. osivis,
which is closely allied to nobilitata.
Cerceris faceta sp. n.
(FIGs. 9, 9a—c; PL. 4, FIG. 8)
2. 9 mm. long. Black. The following parts are bright lemon-yellow: mandibles
excepting the apex, clypeus excepting the median apical margin and the underside
of the median cone, frontal carina, scapes, sides of the face, a small spot behind each
eye, pronotal collar, tegulae, a round macula on the mesopleura below the wing-base,
a round macula on each side of the scutellum, the metanotum, an ovoid macula on
the sides of the epinotum, a band on the first tergite covering nearly the apical half,
a transverse apical macula on each side of the second tergite, a wide apical band on
the third, slightly dilated laterad, a much narrower one on the fourth, the apical half
of the fifth, more or less triangular lateral maculae on the second, third, and fourth
sternites, and the legs. The hind femora on the upper side, and the hind tibiae on the
underside, streaked with black. The flagellum brownish-yellow above, ochreous
below, the first two joints blackish. Wings hyaline, the forewing faintly fuscous at
the apex, the pterostigma reddish-brown. Clypeus, face, and temples and mesopleura
with fairly dense, whitish decumbent pubescence, elsewhere the pubescence is scanty.
Yellow areas of the clypeus and face dull, not very closely punctured. Metanotum
finely and sparsely punctured, metapleura obliquely rugose. Triangular area of the
epinotum smooth and shining, grooved down the middle. The rest of the body is
deeply and closely punctured. Second to fourth tergites are reticulate-punctate with
punctures half as wide as one of the posterior ocelli, a little smaller but less close on
the thorax and fifth tergite, smallest on the face and vertex which are reticulate-
punctate. Pygidial area nearly twice as long as wide, the sides feebly convex, the
apical margin straight, dull, coarsely punctured except over the apical third, which
is rugulose. Sternites sparsely and coarsely punctured on the tumid areas. Mandibles
with a low and wide tooth on the inner margin. Median area of the clypeus inflected
in the anterior third, its anterior margin with a triangular tooth on each side; the
basal two-thirds of the median area raised ventrad and ending in a short acute cone.
Inner orbits feebly divergent above and below. Posterior ocelli half as far again from
the eyes as from each other. Vertex wide, the interocular distance there equal to the
length of the first seven joints of the flagellum. Pronotal dorsum convex transversely
and lengthwise, in the middle as long as the second joint of the flagellum.
First tergite a little wider behind than long, the second as long as the first and
twice as wide as long. Fifth sternite with an acute tooth on each side near the apical
margin. Posterior tibiae with five spiniferous serrations.
ENTOM. 2. 3 N
112 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
3. 7°5mm. long. Apical half of the sixth tergite yellow, the second to fifth sternites
with small yellow maculae on each side; hind femora and tibiae yellowish-brown
on the upper side; otherwise the colour is like that of the 9. Puncturation also as
in the 9.
Median area of the clypeus hexagonal, a little longer than wide, the apical margin
tridentate. Apical joint of the flagellum two-fifths longer than wide, not emarginate
on the inside, very little attenuated towards the rounded apex. The anterior inferior
angles of the pronotum subdentate. Epimeron of the mesopleura swollen, clearly
visible from the dorsal aspect. First tergite as long as wide, widest in the middle.
Otherwise like the 9.
Fatick, Senegal, August 1943, I 9, 1g.
Related to C. rhodesiae Brauns, from which it differs by the smaller size, the smooth
triangular area of the epinotum, the puncturation, and in the ¢ also in the apical ©
joint of the flagellum. .
Cerceris nugax Arnold
(PL. 5, FIG. 17)
1931, Ann. Transv. Mus. 14: 176.
Tillembeya, River Niger, French Sudan, August 1944, 2 gd.
Fatick and Kaolack, Senegal, August 1943, 3 dd.
The specimens from Senegal differ from the type form in having the median
triangular black area of the tergites wider.
A re-examination of the paratype from Dahalak (Dahleck) Island, Red Sea,
indicates that it should be considered a distinct race, not only on account of its
larger size, but also on account of the sculpture of the epinotal triangle and the shape
of the outer paramera of the genitalia. The epinotal triangle is not arcuately and
transversely costate all over as in the type and in the West African specimens listed
above, but is obliquely costate, having about twelve costae divergent caudad from
the base and four or five arcuate costae in the apical angle. The outer paramera are
much blunter at the apex.
This race may be known as imsularis (Pl. 5, fig. 16).
Cerceris yungvei Cameron
(PL. 5, FIG. 18)
1910, Sjéstedts Kilimandjaro-Meru Exped. 8: 273.
Ethiopia, Bishoftu, 1 g, 1 2; Moggio, 2 gg; Akaki, 1 9, 1948; Aselle, 1 g.
Cerceris mazimba Brauns
1926, Ann. Trans. Mus. 11: 295.
Arnold, 1942, J. Ent. Soc. S. Afr. 5: 9.
Tillembeya, River Niger, French Sudan, August 1944, I 2, 3 3.
In the g, 9 mm. long, the yellow markings on the abdomen are larger than in the
type, but the genitalia do not differ.
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 113
Cerceris spinicaudata Cameron
1905, Tvansv. S. Afr. Phil. Soc. 15: 216.
Accra, Gold Coast, 2 99, August 1941; Labadi, Gold Coast, 1 g, March; Kaffrine,
Senegal, 1 ¢, August 1943; Moggio, Ethiopia, 1 g, June, 1946
Cerceris electra sp. n.
(Fics. 10, oa and 6)
2. 8-5 mm. long. Black. The following parts are yellowish-white: mandibles
excepting the black apex, clypeus excepting the black anterior margin of the median
area, sides of the face, frontal triangle and carina, upper side of the scapes, the outer
thirds of the pronotal collar, the outer half of the tegulae, the metanotum, a broad
transverse macula at the base of the second tergite, transverse apical bands on the
third and fifth tergites, that of the third dilated laterad, and a large triangular macula
on each side of the third sternite. Flagellum dark brown above, ochreous below. Fore
and middle femora black above and at the base below, the underside yellowish-white,
the hind femora with a spot of the same colour on the underside at the apex, the
tibiae and tarsi pale yellow, the hind tibiae brownish above, the tarsal joints pale
fuscous apically. Wings hyaline, the apex of the forewing fuscous, the pterostigma
and veins brown. Clypeus and whitish areas of the face with a white and decumbent
pubescence, the thorax and abdomen with sparse, short, and grey pubescence.
Clypeus and pale areas of the face dull, shallowly, finely and sparsely punctured,
the rest of the head with punctures about one-third as wide as the anterior ocellus,
the interspaces hardly wider than the punctures, except on the temples where they
are a little wider.
Mesopleura coarsely reticulate-punctate and dull, the metapleura transversely
striato-rugose, the triangular area of the epinotum shining, impunctate, and grooved
down the middle. The rest of the thorax and the first five tergites moderately shining
and with a strong puncturation. The punctures are about half as wide as the anterior
ocellus, separated on the mesonotum and scutellum by interspaces wider than the
punctures, and as wide on the tergites. Mesonotum impunctate. Tumid areas of the
_ sternites less closely and more shallowly punctured than the tergites. The ventral
margins of the first tergite are raised into a tubercle caudad, and the second sternite
is feebly carinate lengthwise over the basal half. Pygidial area barrel-shaped,
reticulate-rugose, nearly half as long again as wide, the apical margin straight.
Mandibles with two small teeth on the inner margin. Median area of the clypeus
fairly flat, a little wider than long, the anterior margin slightly inflected, the pre-
apical margin forms two very short, wide, and convex lobes extending as far as the
apical margin. Posterior ocelli a little farther from the eyes than from each other.
Interocular distance on the vertex equal to the length of the first six joints of the
flagellum plus half of the seventh. Second joint of the flagellum twice as long as wide.
Anterior coxae produced outwardly in front, but not so much as in C. placida Arnold.
First tergite about as long as wide, the sides feebly convex. Posterior tibiae with six
spiniferous serrations.
I14 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Labadi, Gold Coast, March 1941, I 9.
Allied to C. placida from which it differs by the colour of the scutellum and tergites,
the sparser puncturation of the thorax, the lamina of the pre-apical margin of the
clypeus, and the narrower first tergite.
Cerceris maia sp. n.
(FIG. 11; PL. 5, FIG. 10)
g. 8mm. long. Black. The following parts are pale yellow: clypeus excepting the
black anterior margin, lower half of the face, a spot on the tegulae, apical bands on
the second, third, fourth, and fifth tergites, that of the second dilated laterad, lateral
maculae on the second to fourth sternites. The apical margin of the sternites, except-
ing the first, testaceous. Scapes yellow below, the flagellum ferruginous, paler below
than above, the last four joints fusco-ferruginous above. Tarsi of the fore and middle
legs reddish-yellow, the basal joint pale yellow except at the apex, the tibiae and
trochanters ferruginous, darker above, the tibiae with a narrow reddish streak below.
Trochanters and basal half of the hind femora pale yellow, the apical half black, the
hind tibiae fusco-ferruginous, with a dorsal yellowish-white streak on the apical half,
the hind tarsi reddish-yellow.
Wings smoky hyaline, the forewing beyond the cells and including the radial cell,
pale fuscous, the pterostigma and veins dark brown. Head, dorsum of the thorax,
first tergite, and sternites with a sparse, erect, and whitish pubescence. Clypeus and
yellow areas of the face shining, strongly and closely punctured. Metapleura trans-
versely costate, triangular area of the epinotum shining and smooth, feebly grooved
down the middle; the rest of the body, excepting the basal halves of the sternites,
coarsely reticulate-punctate, the punctures half as wide as the anterior ocellus, except
on the head where they are a little smaller. Median area of the clypeus a trifle longer
than wide, the anterior margin tridentate. Interocular distance on the vertex equal
to the length of the first five joints of the flagellum plus two-thirds of the sixth.
Posterior ocelli nearly half as far again from the eyes as from each other. Second joint
of the flagellum one and a half times longer than wide, the apical third nearly twice
as long as wide at the base, moderately curved and obliquely truncate. Antero-
ventral corners of the pronotum produced into acute triangular teeth, which are
clearly visible when the insect is viewed from above.
Pronotal collar in the middle as long as the second joint of the flagellum. First
tergite widened caudad, the sides convex, one-sixth wider behind than long, and as
long as the second. Pygidial area half as long again as wide, the sides feebly convex,
the apex rectilinear. Posterior tibiae with five serrations. Anterior coxae conically
produced outwards, as in C. Placita Arnold.
Moggio, Ethiopia, April 1948, I 3.
Not related to any other Ethiopian species, and easily distinguished by the teeth
of the pronotum.
Not to be confused with C. inconspicua Arnold, in which it is the prosternum which
is dentate on each side.
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 115
Cerceris alcyone sp. n.
(FIGs. 12, 12a; PL. 5, FIG. 11)
g. 8mm. long. Black. The following parts chrome yellow: the median area of
the clypeus excepting the apical margin and extreme base, a narrow streak on each
side of the face margining the eyes, frontal triangle and carina, an apical band slightly
dilated laterad on the third tergite and on the third sternite, a very thin apical line
on the fifth tergite (a brown one on the sixth), the apical half below of the hind coxae,
and the hind trochanters. Mandibles ferruginous, black at the base and apex.
Antennae dark brown, the first seven joints of the flagellum ferruginous below.
Tegulae ferruginous. Fore legs pale ferruginous, the femora darker above. Middle
femora and tibiae brownish above, red below. The tarsi brown excepting the basal
joint, which is yellow. Hind legs brownish-red. Wings hyaline, the radial cell and
anterior margin beyond it dark brown. Sternites nitidulous, the rest of the body dull.
Clypeus and face not closely punctured, the interspaces about twice as wide as the
punctures. Vertex and temples reticulate-punctate, the punctures one-fourth as
wide as the anterior ocellus. Posterior margin of the sides of the pronotum trans-
versely costate, the metapleura finely and transversely rugose, the triangular area
of the epinotum transversely and arcuately rugose and grooved down the middle;
the rest of the thorax coarsely reticulate-punctate, the punctures half as wide as the
anterior ocellus. The puncturation of the tergites is as large as that of the thorax,
but not quite so close, the interspaces being about half as wide as the punctures. _
The punctures on the tumid areas of the sternites are a little smaller than on the ter-
gites. Median area of the clypeus feebly convex, one-third longer than wide, the
apical margin distinctly tridentate. Interocular distance on the vertex equal to the
length of the first seven joints of the flagellum. The second joint is twice as long as
wide, and the apical two-fifths longer than wide at the base, and subconical. Pos-
terior ocelli slightly farther from the eyes than from each other. First tergite one-
fourth longer than wide, narrower at the apex than at the base. Pygidial area about
half as long again as wide, the apex transverse, the lateral margins feebly convex.
Posterior tibiae with five serrations.
Moggio, Ethiopia, April 1948, I 3.
This species resembles C. macalanga Brauns in the colour of the face and clypeus,
but is not related. It is more closely related to C. spinicaudata Cameron, from which
it differs by the colour pattern, closer puncturation, the rugose triangular area of the
epinotum and the genitalia.
Cerceris merope sp. n.
(Fics. 13, 13a, and 130; PL. 5, FIG. 12)
9. 8-5 mm. long. Black. The following parts pale yellow: mandibles excepting
the apex, clypeus excepting a narrow apical black margin, sides of the face, more
narrowly above than below, upper side of the scapes, the outer thirds of the pronotal
collar, the tegulae, metanotum, a transverse basal macula on the second tergite, an
apical band very narrow mesad but reaching the base laterad on the third tergite, the
tumid half of the third sternite, and an apical band on the fifth tergite dilated laterad,
116 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
the fore and middle tibiae and underside of the femora, the posterior tibiae excepting
an apical infuscation on the underside, the fore metatarsus, and the middle hind
metatarsi excepting the extreme apex. The rest of the tarsal joints ochreous on the
fore and middle legs, blackish on the hind legs.
Clypeus and yellow areas of the face shallowly and not very closely punctured.
Vertex reticulate-punctate, the punctures one-third as wide as one of the ocelli;
17a
Fic. 13. Cerceris merope 9, head, x12. Fic. 13a. Cerceris merope first two tergites, X12.
Fic. 13b. Cerceris merope pygidium, x12. Fic. 14. Cerceris sterope 3, first three tergites, x 12.
Fic. 14a. Cerceris sterope, pygidium, x15. Fic. 15. Cerceris celaeno 3, epinotum and first ter-
gite, x12. Fic. 16. Cerceris macalanga $, epinotum and first tergite, x12. Fic. 17. Cerceris
taygete $, pronotum, X13. Fic. 17a. Cerceris taygete, first two tergites, x 12.
temples closely rugoso-punctate. Pronotal dorsum sparsely and not coarsely punc-
tured, the anterior face a little more closely, the sides rugulose and shallowly punc-
tured. Puncturation of the mesonotum, scutellum, and epinotum excepting the sides
larger than that of the vertex, the interspaces on the mesonotum half as wide as the
punctures, and fully as wide on the epinotum. Mesopleura coarsely reticulate-
punctate, metapleura coriaceous, triangular area of the epinotum shining, grooved
down the middle, very shallowly transversely rugose.
Puncturation of the tergites I-5 as large as on the epinotum, fairly close but not
reticulate-punctate. Pygidial area barrel-shaped, closely rugose, two-thirds longer
than wide, the apical margin straight. First and second sternites carinate lengthwise
medially, the tumid areas punctured like the tergites. Mandibles with a low broad
tooth on the inner margin. Median area of the clypeus as wide as long, its anterior
two-fifths inflected, the inflection not extending to the lateral angles, its upper
Weep > Z
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 117
margin arcuate and slightly tuberculate in the middle. Inner orbits feebly divergent
below. Second joint of the flagellum twice as long as wide, one-third longer than the
third joint.
Posterior ocelli about one-fourth farther from the eyes than from each other.
Interocular distance on the vertex equal to a little more than the length of the first
seven joints of the flagellum. Dorsum of the pronotum as long in the middle as the
first joint of the flagellum, the shoulders rounded. Anterior coxae produced out-
wardly into a blunt cone. Triangular area of the epinotum one-fourth longer than
wide at the base. First tergite as long as wide, widest in the middle, the sides convex,
as long as the second, the latter four-fifths wider behind than long. Posterior tibiae
with five serrations.
g. 8-5 mm. long. Hind tibiae without the apical fuscous streak on the inside, the
hind metatarsus pale yellow, the remaining joints brownish-yellow. The punctura-
tion everywhere coarser than in the 9, especially on the pronotal dorsum, the arcuate
and transverse rugae on the epinotal triangle more distinct than in the 9. The median
longitudinal carina on the first sternite does not reach the apical margin, that on the
second almost obsolete. Pygidial area wider at the base than at the transverse apex,
the sides straight, very little longer than wide at the base. The yellow maculae on the
_pronotal dorsum smaller than in the 9, covering only the outer fifths. Median area
of the clypeus one-fourth longer than wide. Second joint of the flagellum twice as
long as wide at the apex, barely longer than the third, the apical joint slightly longer
than the third, and slightly longer than wide at the base, obliquely truncate. Other-
wise like the 9.
Accra, Gold Coast, September 1941, 1 9; July, 1d.
Allied to C. placida Arnold from which it differs by the colour, shape of the first
tergite, the sculptured triangular area of the epinotum, and in the 9 also in the shape
of the clypeus.
Cerceris sterope sp. n.
(FIGs. 14, 14a; PL. 5, FIG. 13)
3. 7°5-8:5 mm. long. Black. The following parts pale chrome-yellow: clypeus,
face excepting a narrow black line dilated above of each side of the frontal carina,
a very small spot on each side of the pronotal collar, outer third of the tegulae, a
small apical spot on each side of the second tergite, apical bands on the third and
fifth tergites, that of the third dilated laterad, the tibiae excepting a black streak on
the underside and the base of the metatarsi. Fore and middle tarsi ochreous, the hind
pair dark brown. Flagellum dark brown, the second to sixth and the apical joints
pale ferruginous below. Wings hyaline, slightly fuscous at the apex, the veins and
pterostigma black. Clypeus with decumbent yellowish pubescence, the rest of the
body with a sparse, erect, and grey pubescence, long on the head, thorax, and first
two tergites. Clypeus shining, shallowly punctured, the interspaces more than twice
as wide as the punctures on the anterior half of the median area, the yellow areas of
the face a little more closely punctured. Metapleura striato-rugose, the triangular
area of the epinotum rugoso-punctate, grooved down the middle, the sulcus margined
by a carina on each side ; metanotum finely and sparsely punctured. The rest of the
118 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
body coarsely reticulate-punctate, the punctures half as wide as the anterior ocellus, —
except on the upper part of the face, temples, and sternites, where they are a little
smaller. On the scutellum the puncturation is not so close as elsewhere, the inter-
spaces being as wide as the punctures.
Median area of the clypeus one-fifth longer than wide. Second joint of the flagellum
one-third longer than wide, very little longer than the third joint. Posterior ocelli half
as far again from the eyes as from each other.
Pronotal collar in the middle a little longer than the second joint of the flagellum,
the shoulders rounded. First tergite one-fourth longer than wide, nearly as long as
the second, the latter not quite one-third wider than long and two and a quarter times
wider at the apex than at the base. Second sternite with an ill-defined basal plat-
form. Pygidial area two-fifths longer than wide at the transverse apex, its sides —
convex. Posterior tibiae with five feeble serrations.
Haramaia, May 1948, 1 § type; Lekempti, May 1946, 1 g. The specimen from
Lekempti lacks the yellow spots on the pronotum.
In the colour of the legs and scapes, and in the shape of the second tergite, this
species resembles C. macalanga Brauns, but in spite of the difference in the colour of
the first two tergites it is more closely related to C. migrostoma Brauns, having like
that species the median groove of the epinotal triangle margined on each side by a
carina. The genitalia resemble those of nigrostoma and macalanga.
Cerceris celaeno sp. n.
(FIG. 15, PL. 5, FIG. 14)
3. 7mm. long. Deceptively like C. macalanga Brauns, but smaller, and differing
from it in colour and sculpture as follows: Scape pale yellow below, clypeus and face
creamy white or very pale yellow, sparsely and strongly punctured, the interspaces ©
for the greater part three times as wide as the punctures. Triangular area of the
epinotum not distinctly grooved down the middle, only with a thin and indistinct
line, with four or five punctures near the middle and the lateral margins without the —
transversely costate grooves which are present in macalanga. Fifth tergite with a
narrow pale yellow apical band, interrupted in the middle ; in macalanga this segment
is entirely black. Sixth tergite also with an apical yellow band, but much narrower
than in macalanga. Pygidial area parallel-sided (in macalanga wider at the base than
at the apex). Median area of the clypeus more convex than in macalanga, and the
lower half of the orbits are parallel, not divergent as in that species. The dorsal —
aspect of the epinotum is almost semicircular, or about three times wider at the base _
than long, whereas in macalanga it is semi-ovate, or three-fifths wider at the base than —
long (Fig. 16). The first tergite is broader, being only one-third longer than wide,
and is much less contracted caudad. The outer paramera of the genitalia are rela-~
tively shorter, extending barely beyond the apex of the sagittae, and with the widened —
apical portion much shorter. Otherwise like macalanga Brauns. +
Moggio, Ethiopia, June 1946, I 3. |
I gave a corrected description of macalanga 3 in 1942 (J. Ent. Soc. S. Afr. 5: 12).
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 119
Cerceris taygete sp. n.
(Fics. 17, 17a; PL. 5, FIG. 15)
36. 7:5 mm. long. Black, the following parts pale yellow: base of the mandibles,
clypeus excepting a wide black anterior margin, sides of the face, a small spot on
each side of the pronotal collar, outer half of the tegulae, a small subrectangular spot
on each side of the second tergite near its base, a transverse band, slightly dilated
laterad, on the third tergite, a large triangular spot on each side of the third sternite,
the apical half of the sixth tergite, the tibiae, tarsi, and the apex of the fore and middle
femora. The tibiae with a black streak on the underside. Antennae brownish-black,
the underside of the second to seventh joints reddish. Wings hyaline, the veins and
pterostigma brown, the apex of the forewing faintly fuscous. Clypeus with large but
not deep punctures, the interspaces larger than the punctures. Metapleura trans-
versely striate, metanotum shining, with a few small punctures. Tumid areas of the
sternites with a small and shallow puncturation. The rest of the body deeply reti-
culate-punctate ; the largest punctures are on the thorax and tergites and are about
half as wide as the anterior ocellus, those on the vertex and temples a little smaller.
Triangular area of the epinotum with a median furrow, and with somewhat indistinct
rugae divergent from the base outwardly. Median area of the clypeus fairly convex,
one-fifth longer than wide, its apical margin straight and edentate. Dorsum of the
pronotum as long in the middle as the second joint of the flagellum, horizontal, the
shoulders obtusely angular. Prosternum with a denticle on each side near the middle.
First tergite one-third longer than wide, the sides moderately convex, as wide at the
base as at the apex.
Second tergite one-fourth longer than the first, three and a half times wider at the
apex than at the base. Sixth sternite with a sharp tooth on each side. Pygidial area
twice as long as wide, the lateral margins feebly convex, the apical margin straight.
Posterior tibiae with six serrations. Outer paramera of the genitalia dilated at the
apex, their ventral membraneous portion very large.
Gambeila, Ethiopia, February 1948, 1 3.
Related to C. inconspicua Arnold, from which it differs by the narrower face, the
shape of the first tergite, the black femora, and the genitalia. In imconspicua the
median area of the clypeus is only one-ninth longer than wide.
Subfamily ASTATINAE
Of this subfamily there are two examples of two species, of which one is too
damaged to permit of determination. The other is the following:
Astata fuscistigma Cameron
1905, Rec. Albany Mus. 1: 258, 3.
Accra, Gold Coast, 1941, 2 29.
It is remarkable to find the range of this species extending to the tropical belt,
since hitherto it has been recorded only from South Africa.
ENTOM. 2. 3 O
120 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Subfamily NyssONINAE
Alysson guichardi sp. n.
(Fic. 18, 18a, and 18); PL. 5, FIG. 19)
2.9mm. long. Black. Pronotal dorsum, mesonotum, and scutellum fusco-
ferruginous. The median area of the clypeus, excepting a black spot in the middle,
Fic. 18. Alysson guichardi 9, thorax, X15. Fic. 18a. Alysson guichardi, head, x15. Fic. 18},
Alysson guichardi 3, eighth sternite, xc. 34. Fic. 19. Nysson quadricolor 3, flagellum, x27.
Fic. 19a. Nysson quadricolor, seventh tergite, x27. Fic. 20. Brachystegus senegalensis, apical —
joints of the flagellum, x15. Fic. 20a. Brachystegus senegalensis, apical tergites, x 12. Fic. 20b.
Brachystegus senegalensis 2, epinotum, X 12.
a narrow line on each side of the face, the underside of the scapes, a large, subtri-
angular macula, the apex mesad, on each side of the second tergite, the underside of -
the anterior tibiae and the basal two-fifths of the hind femora, creamy white. Last
three joints of the anterior tarsi reddish-yellow. A large macula at the middle of the
mandibles, pale ochreous. Wings fusco-hyaline, the radial cell darker, the veins and
pterostigma blackish. )
The lower half of the face with a sparse whitish pubescence, the clypeus, mandibles, _
declivity of the epinotum, the third and following tergites, and the underside of the
femora with a white and very sparse pilosity. Clypeus smooth and shining. Face
and temples almost dull, finely and very closely punctured, the interspaces about as —
large as the punctures. Vertex shining, sparsely punctured with small punctures —
intermixed with a few larger piligerous ones. Pro- and meso-thorax moderately 4
shining, as finely punctured as the face, but more sparsely, the interspaces for the —
greater part three to four times wider than the punctures; from each of the latter
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 121
there is emitted a small, erect, and black hair. Metapleura smooth and shining, the
metanotum closely, finely, and shallowly punctured. Dorsum of the epinotum one-
third wider than long, as wide at the base as at the apex, its lateral margins feebly
convex ; its median triangular area is defined by strongly raised margins and is as long
as wide at the base. The middle third of the triangular area is margined by two zig-
zagged rugae with transverse rugae between them, and the outer thirds are obliquely
costate. The sides of the dorsum outside the median area are transversely rugose, the
rugae becoming wider apart and stronger caudad. Posteriorly the dorsum is mar-
gined by a strong carina which ends on each side in a broad and low tooth. The
declivity has a median carina which nearly reaches the base and at its upper third is
joined on each side by an oblique carina ; the space between these carinae is coarsely
reticulate-rugose. The sides of the epinotum are closely and obliquely rugose.
Abdomen smooth, shining, and impunctate. Pygidial area broadly triangular,
rounded at the apex, and clothed with adpressed golden setae.
Mandibles with two widely separated teeth on the inner upper margin. Clypeus
short, three and a half times wider than long in the middle, with a deep transverse
impression behind the apical margin. Inner orbits slightly divergent dorsad. Posterior
ocelli half as far again from the eyes as from each other. Interocular distance on the
vertex equal to the length of the first three joints of the flagellum plus two-thirds of
the fourth joint. Flagellum slender, the first joint twice as long as wide, the second
a little more than four times longer than wide at the apex and one-third longer than
the third, all the remaining joints at least twice as long as wide. Pronotum, excluding
the neck, four and a half times wider behind than long in the middle, convex trans-
versely and lengthwise, and with a transverse depression in front of the hind margin.
Mesonotum one and a half times longer than wide at its widest. First tergite a little
wider behind than long, the basal third concave in the middle and divergently
striate, the lateral angles dentate. Posterior tibiae with a row of short spines fringed
with long white hairs on the upper margin. First abscissa of the radius three times
as long as the second, the latter a little longer than the petiole of the second cubital
cell.
g. 6:5 mm. long. The whole of the clypeus and the posterior corners of the pro-
notum, below the tegulae, creamy white. Hind metatarsus also of that colour, except
at the base and apex. Otherwise coloured like the 9. The puncturation of the head
and thorax, in view of the smaller size, is somewhat larger than in the 9. The sculp-
ture of the epinotal dorsum, outside the triangular area, is coarser than in the 9,
and consists of about seven oblique costae with the interspaces transversely rugose.
Mandibles with a small tooth on the inner margin behind the apex. First joint of the
flagellum subglobose, the second joint one and a half times longer than wide, barely
as long as the third, the apical joint truncate at the apex, concave below and twice
as long as wide at the base. Interocular distance on the vertex equal to the length
of the first four joints of the flagellum plus half of the fifth. First tergite one-fifth
longer than wide behind, more deeply concave in the middle of the base than in the 9.
Posterior tibiae only feebly spinose. Apical sternite deeply bifurcate, the apical
tergite longitudinally rugulose, its apex widely rounded. Otherwise like the 9.
Ghedo, Ethiopia, 7,500 ft., May 1946, 2 99, 1 6.
122 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
This species differs considerably from the other African species, guillarmodi Arnold,
and is more closely related to the European ratzeburgi Dahlbom, from which it differs _
in the sculpture of the epinotum and the colour of the scutellum and the legs. It is
noteworthy that both of the African species inhabit mountainous country.
Nysson quadricolor sp. n.
(FIGS. 19, 19a; PL. 5, FIG. 20)
9. 5mm. long. Head black, the base of the mandibles dull yellow, the underside
of the scapes and of the first four joints of the flagellum yellowish-white, the rest of
the flagellum dark brown. Pronotal collar, except in the middle, pale yellow, the tips —
of the epinotal teeth translucent and yellowish, the rest of the thorax dark ferru-
ginous, the anterior third of the mesonotum fusco-ferruginous. Tergites 1-4 black,
the first with a large pale ochreous macula on each side of the apical half, the second
to fourth with apical bands of the same colour, narrowly interrupted in the middle,
the sternites and the last two tergites ferruginous. Legs fusco-ferruginous, the upper
side of all the tibiae yellowish-white, the tarsi reddish-brown. Wings hyaline, the
veins black. The whole body dull and with a very short, decumbent, and whitish
pubescence, scanty on the tergites. Clypeus finely and closely punctured, trans- —
versely impressed behind the straight apical margin. Face with a small and not very
close puncturation, the interspaces closely striate lengthwise. The vertex and temples
are more deeply and more closely punctured, almost reticulate-punctate ; the temples
separated from the foramen by a high carina. Inner orbits strongly convergent
below, the interocular distance on the vertex being twice as great as across the base
of the clypeus, the latter twice as wide as long. Between the antennal sockets there —
is a transverse torus. Scapes obconical, about half as long again as wide at the apex; —
flagellum moderately clavate, the first joint transversely ovoid, the second to eighth
wider than long, the ninth and tenth as wide as long, the apical one and a half times
wider than long. Posterior ocelli very little farther from the eyes than from each
other. Pronotal dorsum feebly punctured, its anterior margin convex, the shoulders
subdentate. Metapleura smooth, feebly and transversely striate at the base. Dorsum —
of the epinotum short, in the middle about half as long as the scutellum, with about
six oblique carinulae, the lateral angles produced into stout conical teeth. The rest of
the thorax coarsely reticulate-punctate. Mesonotum nearly one and a half times wider ~
in front than long, the scutellum twice as wide as long, the sides of the metanotum —
slightly elevated. Tergites strongly and closely punctured, the punctures a little
smaller than those of the mesonotum, the interspaces microscopically striate (40
diameters). Second sternite punctured like its tergite, the other sternites smooth,
excepting their apical thirds which are sparsely punctured. Pygidium triangular, the
apex rounded, about one and a half times longer than wide at the base. First abscissa
of the radius five times longer than the second, a little shorter than the third, the
petiole of the second cubital cell half as long again as the second abscissa ; the cubitus
of the hindwing is emitted from the end of the submedial cell.
3. 5°6mm. long. Clypeus and the underside of the scapes and of the first six joints
of the flagellum pale yellow, the rest of the antennae reddish-brown. Fifth tergite
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 123
with a pre-apical ochreous band as in the preceding tergites, the sixth and seventh
ferruginous, the sternites blackish. Clypeus four times wider than long. Flagellum
clavate, the second to tenth joints more than one and a half times wider than long,
the eleventh wider than long, the apical curved, twice as long as wide at the base,
compressed dorso-ventrally at the apex, which is transverse. Shoulders of the pro-
notum rounded. Apical angles of the seventh tergite spinose. First abscissa of the
radius as long as the third. Outer paramera of the genitalia semi-cylindrical beyond
the base, widened towards the apex; the sagittae with a semicircular dilatation on
the outside at the end of the middle third. Otherwise like the 9.
Tillembeya, French Sudan, August 1944, I 2, 1.
Remotely related to N. braunsit Handlirsch, from which it differs in colour, sculp-
ture, and in the absence of the angular frontal tubercle behind the inter-antennal
torus.
Brachystegus senegalensis sp. n.
(Fics. 20, 20a, and 200; PL. 5, FIG. 21)
2. 7°5-10 mm. long. Head and thorax black. Abdomen reddish-brown, the first
five tergites with a median triangular black area extending from the base to the apex,
and at the basal angles extended to the lateral margins. Clypeus, sides of the face,
and the temples covered with a long, dense, adpressed yellowish silvery pubescence,
concealing the sculpture; the middle of the face and the vertex with less dense,
shorter, and a yellower pubescence, intermixed with setulae. The pronotum, sides
of the epinotal dorsum outside the median area, and the basal half of the first tergite
also with yellowish silvery pubescence, concealing the underlying sculpture on the
epinotal dorsum. A dense adpressed silvery pubescence covers the meso- and
meta-sternum, the underside of the coxae and trochanters, femora, and the sternites.
On the second sternite a sparse and shallow puncturation shows through the
pubescence. Tibiae and tarsi with decumbent silvery pubescence, the hind tibiae also
with a row of short spines on the upper outer margin, and also with white setulae.
Wings pale fuscous. Face and vertex with a shallow and not close puncturation, the
punctures about one-fourth as wide as the anterior ocellus, the interspaces closely
and miscroscopically punctured (25 diameters). Epicnemium glabrous and dull.
Mesonotum, mesopleura and scutellum deeply pitted,! the pits about one-third as
wide as the anterior ocellus and shining, the interspaces narrower than the pits and
closely, microscopically punctured. Metapleura transversely rugose. Median area
of the epinotal dorsum divided by carinae into thirteen unequal areolets which are
shining. The declivity of the epinotum divided by carinae which converge ventrad
into five areolets which are irregularly and transversely costate. Tergites dull, the
first five with a fundamental close puncturation (16 diameters) on which is super-
imposed a larger sparse and shallow puncturation which becomes progressively finer
on each succeeding segment.
Pygidial area closely punctured, nearly twice as long as wide at the base, the apex
narrowly rounded. Anterior third of the clypeus inflected, glabrous and shining, the
? By pits are to be understood depressions having flat floors and vertical sides ; punctures, on the other
hand, have concave floors and sloping sides.
124 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
apical margin feebly convex. Interocular distance on the vertex three-fifths greater
than across the base of the clypeus and equal to the length of the first seven joints of
the flagellum. Face with a cariniform tubercle between the antennal sockets.
Posterior ocelli two-fifths farther from the eyes than from each other. A short
longitudinal carina margins the inner orbits of the posterior ocelli; they are more
clearly visible when the head is viewed from in front. Second joint of the flagellum
one-third longer than wide and as long as the third. Pronotal dorsum in the middle as
long as the first joint of the flagellum, its anterior margin obtusely angular in the
middle, the shoulders dentiform. Mesopleura with an uneven vertical carina in front
of the posterior third, and ending dorsad in a broad, low tooth, just visible from above.
Scutellum two-fifths wider than long, its posterior angles rounded. Posterior angles
of the epinotal dorsum produced into teeth which are as long as wide at the base.
First tergite three-fifths wider than long. As in all the species of the genus, the first
sternite has a median longitudinal and shining ridge, ending abruptly before the
apical margin and flanked by divergent carinae. First abscissa of the radius fully
seven times longer than the second, the latter as long as the petiole of the second
cubital cell. Cubitus of the hindwing emitted at a considerable distance beyond the
end of the submedial cell.
$. 8-2-10°5 mm. long. Pubescence on the clypeus and face pale brassy golden.
Flagellum fusco-ferruginous, the apical joint yellowish. Second to fifth sternites with
apical fimbriae of curved golden hairs, directed cephalad. The black median maculae
on the first, third, fourth, and fifth tergites do not reach the apical margin. The red
lateral areas on the first and second tergites have traces of ochreous patches. Clypeus —
a little more than twice as wide as long in the middle, the glabrous inflected anterior
third rugose and with two very small teeth on each side near the middle.
Inner orbits more strongly convergent below than in the 9, the interocular
distance on the vertex being nearly twice as great as across the base of the clypeus.
Second joint of the flagellum a little longer than wide, the remaining joints except-
ing the last increasing gradually in width and all wider than long, the apical joint
curved, compressed dorso-ventrally, and truncate at the apex. Sixth tergite with a
longitudinal carina on each side near the lateral margins and extending only over
the apical half of the segment. Pygidial area sharply carinate on the lateral margins,
twice as wide at the base as at the apex, strigose lengthwise and closely punctured,
the apex with three strong teeth. Otherwise like the 9.
Diafarabé, French Sudan, August 1944, 2 99, 2 gd.
Nearest to B. decipiens Arnold, from which it is distinguished, inter alia, by the
colour of the wings, legs, and abdomen, the feeble tubercle on the face, and the absence
of teeth on the inferior margin of the mesopleura. ‘
Brachystegus senegalensis var. aurantiacus var. n.
2. 9:5 mm. long. Black. Tibiae and tarsi reddish-brown, the extreme base and
apex of the anterior tibiae pale ferruginous. Tergites 1 and 2 with triangular orange
maculae on each side, occupying more than the apical half of the segments, tergites
3-5 with apical bands, interrupted in the middle and dilated laterad and of the same
colour. Wings pale fuscous. Pubescence and sculpture like that of the species,
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 125
excepting the tomentum on the second sternite which is much less dense and dis-
tributed in little separated patches.
_ g. 7°5 mm. long. Colour as in the 9. The decumbent pubescence on the sides of
the epinotal dorsum is very sparse and absent on the basal half of the first tergite.
Scutellum not so wide as in the type of the species, being only one-fifth wider than
long, and with the lateral margins more convex.
Tillembeya, French Sudan, August 1, 2, September 1944, I 3.
There is so little difference between this variety and the smaller ¢ and @ of the
species, except in the colour of the tergites, that it might be suspected that the colour
in senegalensis has been affected by the vapour of cyanide. However, yellow colours
are turned to bright red by cyanide and not to a reddish-brown. Moreover, in the g
of aurantiacus the sternites are without fimbriae.
Gorytes thalia montivagus subsp. n.
(PL. 5, FIG. 22)
G. thalia Handlirsch, 1895, S.B. Akad. Wiss. Wien, 104: 1023 2; 1901, Verh. Zool. bot. Ges. Wien,
51: 427, 3, 9.
3. 7°5mm. long. This differs from the type of the species as follows. Flagellum
black. Yellow areas on the sides of the face narrower and abruptly contracted above.
The spot on the posterior angles of the mesonotum yellow instead of red. Hind tibiae
black on the inside. Hind tarsi with the basal joint and the basal half of the second
joint pale yellow, the rest black. Apical margin of the fifth tergite rufescent, without
a yellow band. Apical joint of the flagellum longer, three-fifths longer than wide at
the base. Outer paramera of the genitalia narrower and blunter at the apex. The
genitalia are peculiar, having a membranous velum connecting the outer and inner
paramera, and, apparently, also the base of the sagittae. Between the inner and outer
paramera the margin of the velum is produced into a short, curved and chitinized
tooth.
2. 7-5 mm. long. Middle and hind tibiae fusco-ferruginous, yellow on the outside.
Hind tarsi fusco-ferruginous. Pygidial area scutate, narrowly truncate at the apex,
nearly half as long again as wide at the base (as wide there as long in ¢halia i. sp.).
Clypeus shorter, the apical margin straight and with a feeble median emargination.
River Hawash, 6,000 ft., Ethiopia, June, 1946, I g, I 9.
The colour of the legs in the ¢ is like that of G. emery Grib., but the epinotum is
not ‘densely, coarsely punctured, and sub-reticulate’.
Ammatomus spiniferus Du Buysson var. latus var. n.
A. spiniferus Du Buysson, 1897, Ann. Soc. Ent. France, 66: 359, 9.
This variety differs from the type of the species only in having the puncturation of
the mesonotum a little deeper and denser, and in the wider abdomen.
The first tergite is as wide behind as long, and the second three-fourths wider at
the apex than at the base. In spiniferus i. sp. the first is one-fourth longer than wide
behind, and the second is nearly twice as wide at the apex as at the base.
Didessa River, Ethiopia, 4,000 ft., May 1946, 2 99.
126 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Subfamily STIZINAE
tridens group
Stizus braunsii Handlirsch
1894, Ann. naturh. Hofmus. Wien, 9: 287.
Tillembeya, River Niger, August 1939-44, 3 dd, 3 29; River Errer, near Harar,
Ethiopia, May 1948, 2 33.
In Ann. Transv. Mus. 20 (2), 1940, Pl. V, figures were given of the genitalia of ten
species of this very difficult group. Further examination shows that the degree of
chitinization of the basal portion of the outer paramera and the width of the apical
portion are somewhat variable.
Stizus haemorrhoidalis Handlirsch
1900, Verh. Zool. bot. Ges. Wien. 50: 472.
Dilla, Ethiopia, April 1948, 2 g¢.
Stizus multiguttatus sp. n.
(FIGs. 21, 21a, and 21b)
3. 8-5-9 mm. long. Related to S. bvaunsii Handlirsch, but larger and with more
éxtensive and deeper yellow markings. Sculpture like that of braunsit.
Black. The following parts are chrome-yellow: mandibles excepting the apex,
labrum, clypeus, face below the antennal sockets, and a narrow line on each side of
the lower half of the face, scapes, underside of the first two joints of the flagellum,
upper half of the pronotum including the tubercles, lateral margins of the meso-
notum, tegulae, upper two-thirds of the mesopleura, a large more or less polygonal
spot on each side of the scutellum, metanotum, posterior angles of the epinotum, a
more or less semicircular macula on each side of the first tergite not reaching the
apical margin, apical bands on the second to fifth tergites, bisinuate cephalad and
abruptly, widely dilated laterad, an oblong macula on each side of the sixth tergite,
lateral maculae on the second to fifth sternites, that on the second very large, under-
side of the coxae and trochanters, and all the legs excepting the basal third of the
hind femora.
Upper side of the flagellum blackish, the underside, excepting the first two joints,
pale ferruginous. Wings hyaline, the veins dark brown. Clypeus at its widest five-
sevenths wider than long. Interocular distance on the vertex two and three-quarter
times greater than across the base of the clypeus, the inner orbits therefore more
convergent than in braunsii.
Third joint of the flagellum longer than in braunsii, being barely (one-eighth)
shorter than the second, the latter nearly twice as long as wide. The three apical
joints of the flagellum like those of brawnsii, seventh tergite much less attenuated
towards the apex than in braunsii (Fig. 21 c). The outer paramera of the genitalia
much wider than in braunsii.
9. 8-10 mm. long. Second cubital cell not petiolated, the first and second trans-
verse cubital veins meeting on the radius. Sixth tergite entirely black, and all the
a ee a ee
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 127
femora have a black streak on the upper side at the base. Otherwise like the ¢ in
colour, but the apical bands of yellow on the tergites are less deeply bisinuate on their
anterior margin.
In one of the 3 29, the yellow bands are much reduced on the second and following
tergites.
Second joint of the flagellum twice as long as wide, one-fourth longer than the third.
Otherwise like the 3.
Tillembeya, River Niger, French Sudan, August 1944, 5 d¢, 3 29.
2le
2b
Fic, 21. Stizus multiguttatus 3, x6. Fic. 21a. Stizus multiguttatus 3, seventh tergite, xc. 10.
Fic. 21b. Stizus multiguttatus, genitalia, xc. 22. Fic. 21c. Stizus braunsii 3, seventh tergite,
X¢. 10. Fic. 22. Stizus sudanensis g, abdomen, x4. Fic. 22a. Stizus sudanensis, seventh
sternite, xc. 12. Fic. 22b. Stizus sudanensis, genitalia, x 16.
Stizus corpulentus Arnold
1929, Ann. Transv. Mus. 18: 270.
1939, Occ. Pap. Rhod. Mus. 5: 27.
Lake Bishoftu, 7,000 ft., Ethiopia, May 1946, 1 9. A slight variety in which the
clypeus is entirely black, and the pale bands on the abdomen are glaucous white
instead of yellow.
discolor group
Stizus sudanensis sp. n.
(FIGS. 22, 22a, and 220)
6. 12mm. long. Black. Lateral margins of the mesonotum, and of the epinotum,
first tergite excepting a median black T-shaped area at the base, and seventh tergite,
ferruginous. Sternites fusco-ferruginous. The following parts are yellow: mandibles
ENTOM. 2. 3 P
128 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
excepting the apex, labrum, clypeus, face below the antennal sockets, a narrow line
on the lower half of the sides of the face, scapes, upper half of the pronotum, a tri-
angular macula on each side of the scutellum, apical bands on the first six tergites
dilated laterad and also slightly and angularly in the middle, and apical bands on the
second to sixth sternites more or less interrupted medially.
Legs yellow, middle and hind femora flavo-ferruginous. Flagellum dark brown
above, flavo-ferruginous below.
Wings hyaline, slightly yellow, the veins dark brown. Clypeus and lower half of
the face with decumbent silvery pubescence. The puncturation on the head and
thorax, excepting the epinotum, is very shallow and microscopic. On the epinotum
outside the median triangular area the punctures are larger, less oblique, and closer,
the interspaces being as wide as the punctures. The abdomen has the fine oblique —
and shallow puncturation common to the species of the genus. Clypeus twice as wide
in front as long. Interocular distance on the vertex very nearly twice as great as
across the base of the clypeus. Second joint of the flagellum nearly three times longer
than wide and one-sixth longer than the third, the three apical joints as in myasae
Turn. but the spine on the tenth joint is less angularly bent at the apex. Ocelliasfar
from the eyes as from each other. Dorsum of the epinotum short, not longer than the
scutellum. Seventh sternite fairly strongly punctured in the basal half, concave, the _
lateral margins being somewhat swollen, without a median tubercle. Anterior femora —
strongly compressed. First abscissa of the radius three times longer than the second,
Tillembeya, River Niger, French Sudan, September 1944, 1 3.
Like S. nyasae Turner, the other member of this group, this species differs from —
S. discolor Handlirsch in the lack of a median tubercle on the seventh sternite.
Stizus nyasae Turner var. robustus var. n.
(FIG. 23)
S. nyasae, Turner, 1912, Ann. Mag. Nat. Hist. 10: 338.
3. 13mm. long. This variety differs from the type of the species only in the greater
size and extent of the yellow colour. The two small yellow lines on the disk of the
mesonotum which are present sometimes in myasae are here developed into two large _
longitudinal bars extending over the anterior two-thirds of the segment. The
scutellum is black only over its anterior third, and the whole of the epinotum is yellow —
excepting a narrow black basal band on the triangular area, and narrow black
borders outside and adjacent to the lateral margins of that area. In three of the four
specimens the yellow areas of the tergites are not crenulate as in the type of the
species ;' their anterior margins are less indented on the second and third tergites and
on the remaining tergites larger.
The genitalia are identical. The outer paramera have a narrow dorsal face, and
a wider outer face.
9. 12mm. long. One specimen of this sex, from the same locality as the males, —
differs from the ? of typical myasae in having all the black parts replaced with ferru- —
ginous.
Tillembeya, River Niger, French Sudan, September 1944, 4 3d, I 9.
? The heraldic term ‘vair’ accurately describes the arrangement of the colour in myasaei. sp.
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 129
caffer group
Stizus laterimacula fraudulentus subsp. n.
(Fic. 24)
S. laterimacula Handlirsch 1895, S.B. Akad. Wiss. Wien, 104: 980.
g. 11-12 mm. long. Smaller than Jaterimacula i. sp., and with a very different
colour pattern on the abdomen, but with the genitalia barely distinguishable.
_ Puncturation and pubescence as in the type of the species. Black. The following
parts pale chrome yellow: mandibles, labrum, clypeus, face below the antennal
Fic. 23. Stizus nyasae robustus g, genitalia, x12. Fic. 24. Stizus laterimacula fraudulentus Z,
genitalia, x12. Fic. 25. Stizus nigriventris 3, x3. Fic. 26. Stizus bensoni g, head, x6.
Fic. 26a. Stizus bensoni, genitalia, x 12.
sockets and at the sides and a broad median line reaching the anterior ocellus, scapes,
a narrow line on the temples adjacent to the eyes, upper half of the pronotum includ-
ing the tubercles, sides of the mesonotum, upper half of the mesopleura and a tri-
angular macula below it, subtriangular and transverse maculae on each side of the
scutellum, the metanotum, two oblong spots on the median area of the epinotum,
the outer half of the sides of the epinotal dorsum and the lateral margins of the
declivity, lateral semi-elliptical maculae on the first tergite, apical bands on the
second to sixth tergites crenulated and similar to those of S. myasae and S. lon-
culatus, an indistinctly margined median macula on the seventh tergite, small
transverse maculae on the second to sixth sternites and the legs. The femora pale
rufous on the upper side. Flagellum ferruginous. The apical margins of the sternites
are pale brown. Wings hyaline, tinged with yellow, the veins yellowish-red.
The inner orbits are a little more convergent below than in laterimacula and the
130 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
var. euteles, the interocular distance on the vertex being three-fifths greater than
across the clypeus, and the anterior margin of the latter is concave and not straight —
as in the others. As in all the species of this group the hind tibiae are transversely —
compressed and curved outwardly. The genitalia are almost identical with those of
laterimacula i. sp., the only difference being the slightly more rounded apex of the —
outer paramera and the heavier chitinization of the inner margin.
Kaffrine and Kaolack, Senegal, August 1943, 2 gd.
On account of the colour pattern of the abdomen, this insect could easily be mis-
taken for myasae or a small loriculatus. It may be noted here that ewteles Handlirsch,
which I reduced to a variety of laterimacula (1929, Ann. Transv. Mus. 18: 282) differs
from the latter and this race in having the outer paramera of the genitalia densely
pilose on the underside.
fasciatus group
The majority of the species in this group are very similar in the female sex, anda
revision based on the male genitalia is greatly needed, and would probably reduce
some of the supposed species to subspecific, or even varietal, rank. Unfortunately
the $$ of some of them are at present unknown. It is therefore with some hesitation
that I describe as a new species two 99, without the corresponding 3.
Stizus nigriventris sp. n.
(FIG. 25)
9. 20-23 mm. long. Mandibles excepting the black apex, labrum, clypeus, face ~
at the sides and below the antennal sockets and the whole of the outer half of the —
occiput pale ochreous yellow, the inner half of the occiput black. The yellow at the
sides of the face with an extension inwards almost enclosing the anterior ocellus. —
The dorsum of the thorax is mainly reddish ochreous, but the upper two thirds of the
pronotum is pale yellow; the black areas are: the mesonotum excepting the lateral
margins and a large rectangular U-shaped median area extending from the anterior
to the posterior margin, a short triangular band on the base of the median area of the
epinotum and the inner half of each side of the epinotal declivity. Meso- and meta- |
pleura black, excepting a large macula covering the upper two-thirds of the former,
and an oblong macula on the upper half of the metapleura. The underside of the |
thorax, including the coxae and trochanters, is entirely black. First tergite pale
ferruginous, with a large and distinctly defined yellow macula on each side and a
black apical band angularly dilated laterad. Second to fifth tergites black with large
subtriangular maculae on each side, their apices mesad. Sixth tergite ferruginous,
covered with a dense reddish-golden pubescence. Sternites entirely black. Legs and
antennae flavo-ferruginous. Wings flavo-hyaline, the veins flavo-ferruginous, the
radial, second and third cubital cells, and apical half of the second discoidal cell,
fuscous. Clypeus and face with a silvery pubescence, the thorax with a very short
greyish and inconspicuous pubescence. Head and thorax dull, microscopically
reticulate-punctate, the labrum and anterior half of the clypeus less dull and a little
more strongly punctured. Tergites 1 to 5 moderately nitid, very shallowly, finely and
closely punctured. Sternites dull, microscopically reticulate and with a very sparse
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 131
and shallow puncturation superimposed. Clypeus a little more than twice as wide
as long, the anterior margin moderately concave.
Inner orbits subparallel over the lower two-thirds. Interocular distance on the
vertex equal to the length of the first two joints of the flagellum plus four-fifths of the
third and only one-fifth greater than the interocular distance across the base of
the clypeus. Second joint of the flagellum not quite five times longer than wide at the
apex, and fully twice as long as the third. Apical joint slightly curved, a little longer
than wide at the base. Posterior ocelli two-thirds farther from each other than from
the eyes. Pygidial area not marginate, triangular, narrowly rounded at the apex.
First abscissa of the radius slightly longer than the second. Basal joint of anterior
tarsi with six spines.
Tillembeya, River Niger, French Soudan, September 1944, 2 29.
This differs from aurifluus and multicolor, rubroflavus, and rubellus Turner in the
colour of the dorsum of the thorax, especially the V-shaped macula of the meso-
notum, and the entirely black sternites. The clypeus is not deflected in the anterior
half as it is in rubroflavus.
Stizus basalis Guérin
1844, Icon. Régne anim.: 438.
Tillembeya, River Niger, French Sudan, Sept. 1944, I g, 3 29.
Stizus bensoni sp. n.
(Fics. 26, 26a)
g. 17mm. long. Black. The following parts are bright chrome-yellow: basal half
of the mandibles, labrum, clypeus, lower half of the face, a line on the sides of the face
joining a transverse M-shaped bar above, a line on the temples margining the posterior
orbits, two small spots on the posterior margin of the vertex, the upper margin of the
pronotum including the tubercles, the lateral margins of the mesonotum narrowly,
a subtriangular macula on each side of the scutellum, the metanotum, narrow lines
margining the median area of the epinotum but not meeting at the apex, an oblong
spot on the upper angles of the epinotal declivity, transverse bands, widely inter-
rupted mesad and widest laterad on the first six tergites. Seventh tergite fusco-
ferruginous and with pubescence of the same colour. Sternites 1-6 black, the apical
margins brownish-yellow, narrowly on the first three, more widely on the fourth to
sixth, the seventh sternite fusco-ferruginous. Legs and flagellum ferruginous, the
coxae and trochanters black. Wings hyaline tinged with yellow, the veins reddish-
yellow. Head and thorax with exserted yellowish-grey pubescence, fairly long on the
epinotum. Head and thorax dull, the sides of the epinotum nitidulous. Clypeus and
face and vertex microscopically and very closely punctured, pronotum almost smooth,
the rest of the dorsum of the thorax very finely, obliquely and closely punctured, the
interspaces barely larger than the punctures except on the anterior third of the meso-
notum where the puncturation is sparser. Mesopleura as closely punctured as the
mesonotum, but the punctures are round and a little larger. Metapleura and sides
of epinotum almost smooth. Tergites nitidulous, less closely and more obliquely
punctured than the mesonotum, the sternites less closely than the tergites.
132 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Clypeus distinctly convex lengthwise, nearly six-sevenths wider than long, the
anterior margin concave. Face below the antennal sockets slightly raised in the
middle, but not subcarinate. Inner orbits feebly convergent below; the interocular
distance on the vertex equal to the length of the first three joints of the flagellum
plus half of the fourth and one-fourth greater than the least distance between the eyes
below. Posterior ocelli not quite twice as far from each other as from the eyes,
Second joint of the flagellum slender, curved, five times longer than wide at the apex —
and two-thirds longer than the third joint. Apical joints twice as long as wide at the
base, concave below, curved, rounded at the apex. Declivity of the epinotum feebly
concave. Seventh tergite widely rounded at the apex. First cubital vein straight,
first abscissa of the radius a little shorter than the second. Outer paramera of the
genitalia flat and lamelliform in the apical half, the outer margin almost semi- —
circular.
Koundé-Saré, River Niger, French Sudan, August 1945, 1d.
Although the first transverse cubital vein is straight as in the ruficornis group, this
species is evidently closely related to imperialis Handlirsch, as is seen by the shape ~
of the genitalia. In imperialis the outer paramera are not so widely dilated as in this —
species and with the outer margin of that part bluntly angular instead of semicircular —
(1936, Occ. Pap. Rhod. Mus. §: 15, fig. 6). Named after Mr. R. B. Benson, Department
of Entomology, British Museum (Natural History).
Stizus ferrugineus Smith
1856, Cat. Hymen. B.M. 4: 342.
Meisso, Ethiopia, August 1945. Id.
Stizus ritzemae Handlirsch
1895, S.B. Akad. Wiss. Wien, 104: 1000, 9.
Accra, Gold Coast, September 1941, I 9.
This specimen agrees fairly well with Handlirsch’s description, but the first tergite _
is entirely ferruginous.
In the absence of the g a comparison of the genitalia is not possible, but I strongly
suspect that ritzemae is only a colour variety of ferrugineus Sm.
tridentatus group
Stizus amoenus Smith
1856, Cat. Hymen. B.M. 4: 338.
Tillembeya, River Niger, French Sudan, September 1944, I ¢.
Stizus mionii Guérin
1844, Icon. Régne anim. 438.
Koundé-Saré, River Niger, French Sudan, August 1945, I 2, 5 dd.
Subfamily BEMBICINAE
In the descriptions of new species which follow I have not given a lengthy and
detailed account of the colour pattern since I am of the opinion that that can be more
:
ie
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 133
easily appreciated by illustrations in the text-figures. There are, of course, minor
individual variations, but on the whole there is a basic pattern peculiar to each
species.
In the descriptions of new species of this genus I have had occasion to refer to
other African species, comparing the genitalia and the colour-pattern of the tergites.
The reader should therefore refer to my monograph of the African Sphecidae (1929,
Ann. Transv. Mus. 18 (4)) in which figures of the genitalia have been given of the
majority of the species, and to the plate illustrating the pattern of the tergites. In
this genus the outer paramera of the genitalia have a ventral membranous ramus
which in the natural state cannot be seen from the dorsal aspect (see Figs. 27) and
28b) but which is made visible if pressure is applied to the base of the dorsal rami so
as to splay them out.
The collection contains females of three or four possibly new species which, in the
absence of any striking specific character, it would be unwise to name and describe
in the absence of the other sex.
Bembix splendida sp. n.
(FIGs. 27, 27a and 27D)
g. 20mm. long. Mandibles, labrum, clypeus excepting a bi-arcuate black macula
covering the basal two-thirds, sides of the face up to the level of the anterior ocellus,
pale yellow. The rest of the face and the vertex, excepting a quadrate black macula
covering the ocellar area, flavo-ferruginous. Occiput black, the outer part, widening
ventrad, flavo-ferruginous above, merging into pale yellow below. Clypeus with
silvery pubescence, the rest of the head with a long, fairly dense, erect and greyish
pubescence. Pronotum ferruginous, the rest of the thorax black, with the following
parts deep lemon-yellow; a narrow transverse band on the posterior half of the
scutellum, the basal half of the metanotum, a wide V-shaped area margining the
median epinotal area and large triangular maculae, the apex mesad, on the posterior
angles of the epinotal dorsum. Lateral margins of the mesonotum and the tegulae
pale ferruginous. Mesonotum and scutellum with erect yellowish-grey pubescence,
the rest of the thorax, including the pleura and sterna, with a long, erect, abundant
and whitish pilosity. Tergites 1-6 yellow, the extreme base and apex of each one
black, the yellow areas enclosing a black spot on each side of the middle, on the first
and second, the spots on the third to fifth quadrangular and contiguous with the
basal black margin. The base and apex of the sixth and the whole of the seventh,
pale ferruginous. Sternites black, the second to fourth with a small lateral yellow
maculae. Abdomen with a decumbent, short and yellowish pubescence. Antennae
and legs, not including the coxae and trochanters, ferruginous, the apical joints of all
the tarsi, excepting the extreme base, black. Wings clear hyaline, the veins brown.
Labrum and clypeus slightly shining, sparsely and finely punctured. Dorsum of
the thorax dull, densely and very finely punctured, the punctures round. Pleura and
sterna dull, closely punctured, the punctures oblique and a little larger than those of
the mesonotum. Tergites nitidulous, finely, obliquely punctured, the interspaces
about three times wider than the punctures. Sternites shining, the second with a
134 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
smooth, very sparsely punctured median triangular area ending in a short carina,
truncate or dentiform at its apex, the sides are closely and obliquely punctured, the
following sternites less strongly punctured, the sixth with a raised semi-elliptical
platform, acute at the apex, the seventh with a shining median carina. The seventh
tergite is sparsely and coarsely punctured, abruptly attenuated near the apex.
Clypeus twice as wide as long, subcarinate at the base, the apical margin concave.
Inner orbits moderately divergent below the middle. Interocular distance on the
27a ‘
Fic. 27. Bembix splendida g, xc. 2:7. Fic. 27a. Bembix splendida, apical joints of flagellum,
X13°5. Fic. 27b. Bembix splendida, genitalia, «14:5. Fic. 28. Bembix fantiorum 4, thorax,
X7°5. Fic. 28a and b. Bembix fantiorum, genitalia, dorsal and ventral view, x 13°5. Fic. 28¢.
Bembix fantiorum 3, apical tergite, x12. Fic. 28d. Bembix fantiorum , apical joints of the
flagellum, x 13°5.
vertex equal to the length of the first two joints of the flagellum plus four-fifths of the
third. Second joint of the flagellum four times longer than wide, and three-fifths
longer than the third, the sixth and following joints excavated below, the seventh
bidentate on the outer margin, the apical acuminate, two and a half times longer than
wide at the base. Anterior coxae for the greater part pale yellow. Middle metatarsi
dilated below near the base. Middle femora not dentate on the hind margin.
Koundé-Saré, River Niger, French Sudan, August 1945, 3 dd.
Related to ugandensis Turner, from which it differs by its smaller size, ferruginous
legs and antennae, the round puncturation of the thorax, the colour of the last four
tergites, the unarmed middle femora, and the genitalia.
This species belongs to the diversipenmis group.
7 Nad z -
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 135
melanopa group
Bembix fantiorum sp. n.
(Fics. 28, 28a—d)
g. 14-15 mm. long. Black. The following parts are pale yellow: the sides of the
labrum and clypeus to a varying degree, sometimes the former has only the median
third black, and the clypeus may be entirely black, the sides of the face, two small
spots below the ocellar area, the outer orbits, mandibles excepting the apex, pro-
notum, lateral margins of the mesonotum and sometimes two longitudinal streaks on
the anterior half of the same, sometimes the hind margin of the scutellum, the greater
part of the mesopleura, the metapleura excepting the margins, and the anterior and
posterior thirds of the sides of the epinotum.
First five tergites in the middle with transverse bands of glaucous white, becoming
yellower laterad, bisinuate cephalad, and more or less interrupted in the middle; the
sixth tergite with two whitish spots. The shape of these bands is like that of melanopa,
and as in that species the yellow band on the second tergite sometimes encloses a
black spot on each side (see Aun. Transv. Mus. 18, 1929, Pl. 4, figs. 16 and 17).
Sternites 2-5 with triangular yellow maculae at the sides. Antennae black, the
last three joints ochreous below. Legs yellow, coxae and trochanters partly black,
the underside of the fore and middle tarsi black, the anterior femora on the upper side
and the anterior tibiae both above and below with black streaks, the middle and hind
femora with small black streaks above near the apex and at the base below, the middle
and hind tibiae with black streaks above and below. Wings clear hyaline.
Labrum and clypeus with a few scattered punctures.
Thorax dull, closely and obliquely punctured, tergites nearly dull, a little more
finely punctured than the thorax. Sternites shining, coarsely and obliquely punc-
tured, the second more densely so than the rest.
Clypeus very convex, two-thirds wider than long. Inner orbits slightly divergent
below. Second joint of the flagellum three times longer than wide and half as long
again as the third, the fifth to ninth spinose, the last three concave below. Middle
femora with about eight teeth on the apical half of the hind margin. Tarsal comb
with yellow spines, of which there are six on the basal joints. Second and sixth
sternites as in melanopa, the seventh more narrowed in the apical part than in that
species. The seventh tergite, coarsely punctured, is more narrowed apically than in
that species and the apex is widely rounded.
9. 15 mm. long. Mesonotum with a large U-shaped yellow macula, rectangular at
the base. Hind margin of the scutellum and metanotum, and the posterior half of the
median area of the epinotal dorsum, yellow. First tergite sometimes with two trian-
gular yellow maculae on the declivious anterior face. The yellow bands on the second
and third tergites usually enclose transverse black maculae, and the band on the first
is usually entire. The sixth tergite is entirely black and more coarsely and more
closely punctured than in the 4, its lateral margins not sinuate, the apex narrower
than in the other sex. The second sternite is closely punctured at the sides, much
more coarsely and very sparsely in the middle. The black streak on the upper side
of the hind femora is longer and wider than in the ¢.
ENTOM. 2. 3 Q
136 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Otherwise like the g in colour, sculpture, and pilosity.
Second joint of the flagellum three and a half times longer than wide, and about
one-third longer than the third joint. Apical joint moderately curved, compressed
dorso-ventrally at the apex, and nearly three times longer than wide at the base.
Tarsal comb fusco-ferruginous.
Labadi, Gold Coast, March—May, 22 gd, 8 99; Kaolack, Senegal, August, 1 4,
2 2°; Tillembeya, River Niger, September, 2 gg, 2 29.
This is not melanopa, race forcipata Handlirsch (which is also richly yellow on the
pleura), since, according to Handlirsch, the genitalia do not differ from those of the
type of the species, but it is more closely related to melanopa i. sp. than to any other,
in spite of the difference in the genitalia.
The species is named after the major native tribe of the Gold Coast.
Bembix guigliae sp. n.
(Fics. 29, 29a-c)
3g. 14-16 mm. long. Black. Mandibles excepting the apex, labrum, clypeus at the
sides and on the anterior margin, outer orbits more widely below than above, sides —
of the face, the bottom of the face above the antennal sockets, and a transverse bar
with irregular margins below the ocellar area, yellow.
Underside of the scapes yellow, the upper side and the flagellum excepting the last
three black joints ferruginous.
The yellow markings on the thorax are very much the same as those of the 9 of the
preceding species, fantiorum q.v., excepting that the U-shaped macula on the meso-
notum is absent, and the maculae on the pleura are less extensive. The yellow is also
of a deeper colour, deep chrome-yellow. The pattern of the deep chrome-yellow
markings on the first five tergites is also very similar to those of fantiorum, but the
enclosed transverse black maculae are often continuous with the basal black area on
the second tergite and always so on the third. Sixth tergite black at the base, the
apical third ferruginous, seventh tergite fusco-ferruginous, with the basal third black. —
|
é
.
Sternites 1-6 fusco-ferruginous in the middle, blackish laterad, the second to fifth ©
with triangular yellow maculae at the sides; sixth sternite black over the basal half, —
and ferruginous on the apical. Legs yellow, the anterior tibiae with an oblong black
macula on the underside, the fore and middle femora with a black streak above, and
a short one on the underside at the base ; the hind femora have only a black and short
macula at the base. Wings hyaline, tinged with yellow, the forewing slightly darker
over the basal half but not distinctly infuscated. Clypeus with adpressed silvery —
pubescence, the rest of the head, the thorax, and base of the first tergite with a long
erect and yellowish-white pilosity.
The sculpture of the head, thorax, and tergites is like that of B. fuscipennis Le
Peletier but a little finer. The puncturation of the sternites is deeper and more dense,
especially on the second which has a ferruginous, compressed, and curved tooth like
that of fuscipennis. The raised scutate platform on the sixth is narrower at the base
than in fuscipennis.
The seventh sternite differs considerably from that of fuscipennis ; the basal half
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 137
is slightly flattened in the middle and the apical half is depressed, marginate over the
basal half of its sides, and subcarinate in the middle. Basal joint of middle tarsi
dilated basad as in fuscipennis. The antennae as in that species, but the tenth and
eleventh joints of the flagellum are distinctly shorter. Middle femora without teeth
on the hind margin.
2. 14mm. long. Sixth abdominal segment ferruginous, the yellow margin on the
sides of the face and the transverse band below the ocellar area often broken up into
29
Fic. 29. Bembix guigliae 3, genitalia, x15. Fic. 29a. Bembix guigliae g, apical tergite, x15.
Fic. 29b. Bembix guigliae 3, 7th sternite, x15. Fic. 29c. Bembix guigliae 3, apical joints of
flagellum, x15. Fic. 30. Bembix fuscipennis g, genitalia, x 20. Fic. 30a, Lateral view of apex of
inner paramera.
three or four spots, flavo-ferruginous, the black maculae on the underside of the
femora larger than in the g, otherwise like the g in colour, sculpture, and pilosity.
Second joint of the flagellum four times longer than wide and twice as long as the
third. Inner orbits feebly divergent below. Clypeus twice as wide as long.
Labadi and Accra, Gold Coast, March—May, 9 gg; Kaffrine, Senegal, August, I 3;
Labadi and Accra, April and August, 6 99; Kaffrine, Senegal, 1 2; Tillembeya, River
Niger, August, I 3.
Named after Dottoressa Delfa Guiglia of the Genoa Museum who has published
valuable papers on the African Sphecidae.
Closely related to B. fuscipennis, but with a different colour-pattern and suffi-
ciently different genitalia. The outer paramera are wider than in fuscipennis i. sp.
and much more densely pilose on the underside. In this species and in fuscipennis the
inner paramera in the normal position and viewed from above have the appearance
138 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
of cylindrical rods. They are actually rod-shaped basad, but are expanded and
spatulate apically, with the anterior margin strongly concave.
On account of the yellow fasciae and maculae on the thorax this is not the B.
stevensont Parker. I take this opportunity to reply to some criticisms by Professor
Parker (1942, Proc. Ent. Wash. 44: 208). In my monograph of the African Sphecidae
(1931, Ann. Transv. Mus., 14: 214) I stated that stevensoni was only a clear-winged
variety of fuscipennis. I have a fairly large series before me, taken in many cases in
the same localities and at the same times as the typical form. Professor Parker, in
the paper referred to, contrasts in Pl. 18 the shape of the outer paramera with those
of a specimen of fuscipennis 1. sp. which he received from a Mr. Stevenson and stated
to be determined by myself. He also says that this illustration, Fig. 15, is ‘in all
essentials identical with that given for this species by Handlirsch’. In view of the fact
that labels are often misplaced, I accept no responsibility for determinations which
do not carry my printed determination label and which are not obtained direct from
me. Furthermore, the illustrations in Handlirsch’s monograph are by no means
reliable, being limited in most cases to the upper ramus of the stipes and so devoid of
detail as to be semi-diagrammatic. His figure for fuscitpennis is inaccurate, and I
therefore give in Fig. 30 an illustration of the genitalia of that species. It will be seen
that, on comparing this with Parker’s figure of the genitalia of stevensoni (1929,
Proc. U.S. Nat. Mus. 75, Pl. 14, fig. 197), the only difference, a very slight one,
is that the apical part of the stipes on this inner margin is very obtusely angular,
whereas in fuscipennis 1. sp. it is almost straight. Parker’s Fig. 199 of the seventh
tergite agrees with that of fuscipennis.
Bembix agrestis Parker
1929, Proc. U.S. Nat. Mus. 75: 135.
Dessie, November 1945, 1 3; Lake Bishoftu, Ethiopia, 7,000 ft., February 1948,
229;
Bembix massaica Cameron
1910, Sjdstedt’s Kilimandjaro-Meru Exped. 8: 290.
‘River Hawash, S. of Adama, Ethiopia, June 1946, 12 99.
Bembix intermedia Dahlbom
1845, Hymen. Eur. 1: 491.
Danakil, Mille River, 1,500 ft., Ethiopia, July 1946, 1 9.
Smaller than South African specimens, 12 mm. long.
Bembix cultrifera ypsilon subsp. n.
B. cultrifera Arnold 1929, Ann. Transv. Mus. 18: 339.
2. 16mm. long. Disk of mesonotum with U-shaped yellow macula, the base
touching the hind margin but the arms not reaching the anterior margin. The labrum
is even shorter than in the type of the species, being but little longer than wide at the
base. The flagellum is thinner, the second joint fully four times longer than wide,
instead of three and a half. Wings short, the forewing not extending back beyond the
apex of the fourth tergite.
Ufdem, Ethiopia, August 1945, I @.
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 139
Subfamily AMPULICINAE
Dolichurus ignitus contractus subsp. n.
D. ignitus Smith 1869, Trans. Ent. Soc. Lond.: 304.
9. 85-10 mm. long. This race differs from ignitus i. sp. as follows. Apex of the
clypeus, mandibles, tibiae, and tarsi blackish-brown. Last three abdominal segments
and apical margin of the third fusco-ferruginous.
Face with faint traces of rugae at the sides, otherwise smooth, and, like the vertex,
with a few very shallow punctures. The dorsum of the pronotum is more narrowed
cephalad and less depressed in the middle, so that the posterior tubercles are less
prominent, the transverse rugae on the anterior margin are reduced to faint striae,
and the shoulders are distinctly dentiform and not merely rectangular. The dorsum
of the epinotum is much more narrowed caudad, the base being three-fifths wider
than the apical margin, whereas in ignitus i. sp. it is only one-seventh wider. The
rugosity on the epinotal dorsum is much reduced, the transverse rugae being fewer.
In the larger specimen (type) the hind margin is raised at the corners into blunt
triangular teeth, but on the smaller specimen the margin is only slightly higher there.
Dilla (Sidamo), Ethiopia, April, 2 29.
Ampulex toroensis Turner
1919, Ann. Mag. Nat. Hist. 4: (9) 45.
Adis Ababa, 1 ¢. |
Ampulex sp.
Accra, Gold Coast, December 1941, I g.
Possibly the ¢ of one of the several species of Kohl described only from 99.
Subfamily SPHECINAE
Sceliphron (Pelopeus) fulvo-hirtum Arnold
1928, Ann. Transv. Mus. 12: 246.
Lekempti, Ethiopia, May 1946, 4 3.
Sceliphron (Hemichalybion) brachystylus Kohl
1888, Verh. Zool. bot. Ges. Wien, 88: 154 9.
1918, Ann. naturh. Hofmus. Wien, 82: 83 g.
Lake Bishoftu, Ethiopia, May 1946, I 3.
Sceliphron (Chalybion ) planatum sp. n.
(Fic. 31)
6: 14mm. long. Metallic blue, the abdomen steel blue, femora and tibiae purplish-
blue, tarsi dark brown. Antennae black. Mandibles fusco-ferruginous. Wings
hyaline, the apex beyond the cells narrowly fuscous. Clypeus and face with long
silvery pubescence. The rest of the head and the thorax with outstanding, long, and
whitish pilosity. The abdomen with adpressed greyish pubescence.
Clypeus distinctly convex, subcarinate medially on the anterior half, the apical
140 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
margin tridentate. Face shining, fairly strongly, but sparsely punctured, the vertex
shining with a few punctures, the temples impunctate. Thorax shining, the pro-
mesonotum very sparsely punctured, a little more finely than the face, the pronotum
bituberculate. Mesopleura and mesonotum with very shallow, widely spaced punc-
tures which, seen at a certain angle, form vertical lines. Metapleura not sunken, the
surface on the same plane as the mesopleura and sides of the epinotum, with which
they are so completely fused that the margins are almost obliterated ; they are finely,
33b
Fic. 31. Sceliphron planatum 3, x7}. Fic. 32. Ammophila barbarorum 9, head, x 7}. Fic. 324.
Ammophila barbarorum 2, pronotum, x 73. Fic. 33. Ammophila caprella 3, head, x 74. Fic. 33a.
Ammophila caprella 3, pronotum, x7}. Fic. 33b. Ammophila caprella 3, genitalia, x18.
Fics. 34 and 34a. Ammophila argyrocephala 2, pronotum, front and side views, x 12.
shallowly, and sparsely punctured. Seen from the side, the outline of the mesopleura
is almost a straight line, as in S. heinit Kohl.
Epinotum very long, transversely striate, the dorsum three times longer than the ~
only slightly lower declivity. The dorsum is almost flat, not grooved down the middle.
Petiole moderately curved, as long as six-sevenths of the hind metatarsus. Inter-
ocular distance on the vertex equal to the length of the second joint of the flagellum —
plus three-fourths of the third, and hardly greater than across the clypeus. Second
“
F
e
joint of the flagellum barely longer than the third, and not quite three and a half —
times longer than wide at the apex. Third abscissa of the radius one-fifth longer than
the second.
Danakil, Mille River, 1,500 ft., Ethiopia, July 1946, 2 3d.
This species belongs to the section of Chalybion in which the metapleura are not
sunken, and is not to be confused with S. heiniz Kohl in which they are distinctly so.
—
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 141
Ammophila (Podalonia) tydei Le Guillou v. capensis Le Peletier
1845, Hist. Nat. Ins. Hymén, 3: 368.
Lake Bishoftu, Lekempti, Haramaia, and Diredawa, Ethiopia, 4 $3, 5 99, May—
September.
Ammophila gracillima Taschenburg
1869, Z. ges. Naturgesch. 84: 433.
Sardo, Ethiopia, July 1946, 1 dg.
Ammophila beniniensis Palisot de Beauvois
1837, Ins. vec. en Afr. et Amér.: 48.
Moggio, Ethiopia, April 1948, I ¢.
Ammophila bonae-spei Le Peletier v. ferrugineipes Le Peletier
1845, Hist. Nat. Ins. Hymén. 8: 383.
Dilla, Ethiopia; Accra and Labadi, Gold Coast; Kaffrine, Senegal: 3 gg, 5 $9.
Ammophila punctaticeps Arnold
1919, Proc. Rhod. Sci. Ass. 18: 52.
Tillembeya, River Niger, French Sudan, September 1948, 1 9.
Ammophila barbarorum sp. n.
(FIGS. 32, 32a)
9. 21 mm. long. Black. Mandibles excepting the apex, tegulae, the apex of the
first tergite above and the whole of the underside, the second tergite and sternite, and
the sides of the third tergite ferruginous. Legs ferruginous, excepting the upper side
of the fore and middle tibiae and the hind coxae, trochanters, and femora, which are
black. The metatarsus of the hind legs and the underside of the scapes fusco-
ferruginous. Wings hyaline, slightly tinged with yellow, the apex of the forewing
faintly fuscous, the veins brown.
Clypeus, lower half of the face, and the temples with a greyish silvery pubescence,
and a long whitish pilosity. Thorax, including the trochanters and the anterior
femora, with a similar pilosity, and on the pleura, mesonotum, and sides of the epi-
notum a yellowish-grey tomentum which does not obscure the sculpture. Head dull,
face and clypeus finely and shallowly punctured, the interspaces almost twice as wide
as the punctures on the clypeus, upper half of face, and vertex, and about as wide on
the lower half of the face. Temples almost impunctate.
Pronotum finely punctured, the dorsum slightly shining. Mesonotum transversely
striate, closely punctured between the striae, the sculpture less strong in the middle
posteriorly. Scutellum and metanotum closely and longitudinally striate. The pleura
finely punctured, less closely than the mesonotum, and with traces of transverse
striae on the sterna. Scutate area of the epinotum closely, arcuately, transversely
striate, widest across the middle, one-fourth longer than wide at the base. Clypeus
feebly convex, two-thirds wider than long, the apical margin convex. Inner orbits
142 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
barely convergent below. Interocular distance on the vertex equal to the length
of the first joints of the flagellum plus one-fourth of the fourth joint. Second joint
of the flagellum four times longer than wide at the apex and five-eighths longer
than the third. Dorsum of the pronotum about as long as the third joint of the ~
flagellum, very slightly depressed on each side of the middle, but much less widely
so than in dolichocephala Cameron, three times wider behind than long in the middle,
the anterior face nearly vertical, not oblique as in dolichocephala Cameron. First —
joint of the petiole as long as the second, the second tergite conical, nearly twice as
long as wide behind. Fifth to sixth tergites with a decumbent and silvery pubescence.
The third abscissa of the radius in the type is only one-third as long as the second,
and in the paratype even less, so that the second and third transverse cubital veins
are almost contiguous on the radius. In the paratype the pronotum is a little shorter
than in the type and very feebly depressed on each side of the middle.
Adis Ababa, November 1945, 2 29.
Related to dolichocephala Cameron, from which it may be distinguished by the
slope of the pronotum, the much less produced clypeus, and the shorter joints of the
flagellum. In dolichocephala the second joint of the flagellum is fully five times longer
than wide, and the interocular distance on the vertex is equal to the length of the
first two joints of the flagellum plus half of the third.
Ammophila caprella sp. n.
(Fics. 33, 334, and 336)
g. 23mm. long. Black. Mandibles excepting the apex, the fore legs, the underside
of the middle femora, and the middle and hind tibiae ferruginous; the underside of
the hind tibiae blackish in the apical half. The underside of the first tergite (second
joint of the petiole), the second tergite and sternite, and the sides of the third tergite
fusco-ferruginous. Wings hyaline, the apex faintly fuscous, the veins dark brown.
Face and clypeus with a dense, dull golden yellow pubescence, the temples with —
silvery pubescence. Head and thorax with an abundant yellowish-grey pilosity,
fairly short on the mesothorax and epinotum, long elsewhere. Below the pilosity is
a dense, pale yellowish tomentum, obscuring the sculpture except on the dorsum of |
the thorax. Upper part of the face and the vertex dull, shallowly punctured, closely
on the face, with a small area in front of the ocelli impunctate. Thorax dull, the
pronotum finely and sparsely punctured, the mesonotum closely punctured and
finely transversely striate. Scutellum and metanotum punctured and with a trace of —
longitudinal striae. Scutate area of the epinotum closely transversely rugose and
punctured between the rugae.
Clypeus flat, more than one-third of its length projects beyond the level of the
bottom of the eyes, the anterior margin straight. Interocular distance on the vertex
equal to the length of the first three joints of the flagellum, and one-fourth greater
than interocular distance across the clypeus.
Second joint of the flagellum nearly four times longer than wide and about one-
third longer than the third joint. Posterior ocelli nearly twice as far from the eyes
as from each other. Pronotal collar very convex transversely and lengthwise, three
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 143
times wider behind than long in the middle. Scutate area of the epinotum almost
parallel-sided, one-fourth longer than wide. First joint of the petiole a little longer
than the second. Second tergite three-fourths longer than wide at the apex.
2. 24mm. long. Pilosity on the head and thorax not so dense as in the g. Clypeus
less produced than in the g, three-fourths wider than long, the anterior margin feebly
convex, and of the same shape as in barbarorum 9.
Posterior ocelli two-thirds farther from the eyes than from each other. Interocular
distance on the vertex equal to the length of the first three joints of the flagellum
plus one-third of the fourth. Second joint of the flagellum nearly twice as long as the
third. Scutate area of the epinotum widened caudad, one-sixth longer than wide.
Otherwise like the ¢.
Adis Ababa, April 1948, 1 §; Managasha, Ethiopia, November 1945, I 9.
Related distantly to A. insignis Smith, race transvaalensis Cameron, from which it
differs by the long pilosity and colour of the pubescence on the face, and in the $ by
the clypeus being wider and flat in the apical part, and in both sexes by the much
shorter joints of the petiole. The shape of the clypeus in the 9 also differs consider-
ably from that of zusignis.
Ammophila argyrocephala sp. n.
(Fics. 34 and 34a)
2. 19-21 mm. long. Black. Basal half of the mandibles, tegulae, the underside
of the petiole, second and third tergites at the base and sides, and the second and third
sternites, ferruginous. Legs dark ferruginous, the upper side of the coxae, trochanters,
and femora more or less black. Wings hyaline.
Clypeus, face, and temples with a dense silvery pubescence, which on the face
appears to end abruptly below the vertex, where the pubescence is much shorter.
The temples have in the middle a fringe consisting of a single row of long, white hairs.
Thorax with a yellowish-silvery and dense tomentum, obscuring the sculpture on the
pleura and epinotum. Pronotum smooth, almost impunctate. Mesonotum indis-
tinctly transversely striate, the striae absent from the disk. Epinotal dorsum closely
transversely rugulose.
Clypeus five-sixths wider than long, the apical margin feebly convex. Inner orbits
_ parallel. Interocular distance on the vertex equal to the length of the first two joints
of the flagellum plus three-fourths of the third.
Second joint of the flagellum four and a half times longer than wide and fully half
as long again as the third. Posterior ocelli nearly one and a half times farther from
the eyes as from each other. Pronotal collar four times wider behind as long, the
dorsal face merging gradually into the sloping anterior face ; seen from the front it is
much narrowed dorsad, or broadly triangular with the apex rounded. The pro-pleura
have a single row of long hairs. Scutate area of the epinotal dorsum gradually
widened from the base to near the apex, then narrowed, one-third longer than wide
at the base. First joint of the petiole a trifle longer than the second. Second abscissa
of the radius twice as long as the third.
River Errer, May 1948, 1 2; Meisso, Ethiopia, September 1945, 2 29.
ENTOM. 2. 3 R
144 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Sphex (Parasphex) senegalensis sp. n.
(Fic. 35)
g. 13mm. long. Black. This species is so much like S. lividocinctus Costa, a north
Mediterranean species, that one would be inclined to consider it a subspecies but for
the fact that the habitats are separated by the Mediterranean Sea and the Sahara.
Kohl’s description of Costa’s species (1890, Ann. naturh. Hofmus. Wien, 52339,
es
wy
Fic. 35. Sphex senegalensis, genitalia, x21. Fic. 36. Tachytes basilica, genitalia, x18. FIG. 37.
Tachytes guichardi, genitalia, X18. Fic. 38. Tachytes habilis, genitalia, x18. Fic. 39. Tachytes
lamentabilis, genitalia, x18. Fic. 40. Tachytes griseola, genitalia, x21. Fic. 41. Tachytes
rufitibialis, genitalia, x 21.
Pl. VIII, fig. 11) applies in nearly every respect to this species except in the following
details:
First tergite dark brown, the apical margin of the second to fifth tergites reddish
testaceous.
Head and thorax with a long yellowish-white pilosity which on the face, temples, and
epinotum is so dense as to hide the underlying sculpture. First tergite with a thin
decumbent pubescence of the same colour.
Vertex shining, finely and sparsely punctured. Mesonotum shining, punctured,
the interspaces for the greater part two or three times wider than the punctures.
Scutellum shining with a few small punctures and distinctly grooved down the middle.
Mesopleura closely punctured, the punctures larger than those of the mesonotum.
Epinotum closely, transversely, and somewhat arculately rugose, on the dorsum
posteriorly also punctate. The proportions of the antennal segments, the interocular
rr
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 145
distance on the vertex, and the length of the petiole are the same as in lividocinctus
Costa, also the venation. Claws with four distinct teeth.
Koalack, Senegal, August 1943, I J.
Sphex (Isodontia) Simoni du Buysson
1897, Ann. Soc. Ent. Fr. 356.
Dilla, Ethiopia, April 1948, I 9.
Sphex (Proterosphex) cinerascens Dahlbom
1845, Hym. Eur. 1: 25 and 436. .
Labadi, Gold Coast, and Tillembeya, River Niger, French Sudan, April and
September, 5 33,5 29.
Sphex (Proterosphex) incomptus Gerstaecker
1871, Arch. Naturgesch. 87: 352.
Aburi, Gold Coast, Dec. 1941, 2 99; Dilla, April 1948, and River Hawash,
Ethiopia, June 1946, 2 99.
Sphex (Proterosphex) nigrohirtus Kohl
1895, Ann. Naturh. Hofmus. Wien, 10: 66.
S. convadti Berland, 1927, Bull. Mus. Hist. Nat., Paris, 1927 (2): 154.
Aburi, Gold Coast, December 1941, I 9.
This specimen agrees in every detail with the description of conradtt. An examina-
tion of the material in the National Museum, Southern Rhodesia, comprising two 99
from Abak, Nigeria, and two 99 from Uganda, shows that the depth of the depression
on the clypeus varies, and convadii are merely specimens in which it is deep.
Sphex (Proterosphex) haemorrhoidalis Fabricius
1781, Spec. Insect. 1: 443.
This species is given to considerable variation in the colour of the flagellum,
clypeus, and legs, all of which may be more or less red instead of black as in the typical
form. Four g¢ from Aburi, Gold Coast, and Jowaha and River Didessa, Ethiopia,
do not diverge from the type, but one 2 from Aburi has the hind tibiae fusco-ferru-
ginous. Two other 99 from Aburi and Kpeve, Gold Coast, are intermediate between
the varieties volubilis Kohl and pulchripennis Mocsarz.
Sphex (Proterosphex) rufiscutis laevigatus subsp. n.
S. rufiscutis Turner, 1918, Ann. Mag. Nat. Hist. (9) 1: 359.
Q. 19 mm. long. This differs from the typical form as follows. The epinotum has
a thin, short, and greyish pilosity, very scanty on the dorsum, and the declivity lacks
the decumbent pubescence which, with the pilosity, obscures the sculpture in the
type. The sculpture of the dorsum consists of a very close and fine transverse stria-
tion only, larger rugae being absent ; and is less narrowly caudad. There is a shallow
median longitudinal impressed line. The interocular distance on the vertex is greater,
being very nearly equal to the length of the first three joints of the flagellum.
The petiole is shorter, being only a little longer than the second joint of the flagellum
146 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
(in the type of the species equal to the length of the second plus half of the third).
The pronotal collar is shorter and flatter.
Near Sofara, French Sudan, September 1944, I 9.
Possibly a distinct species, but in the absence of the ¢ it is best to treat it asa
subspecies.
Subfamily LARRINAE
Genus Tachytes Panzer
The collection contains a number of species of which a large proportion are repre-
sented by only one sex, and by specimens which are defective, for which reason they
cannot be described although apparently new species. In all the collections which
I have seen, the males outnumber the females. The reason is that the former are, by
their habits, much more conspicuous. They visit flowers far more frequently than
the females, and when not so occupied are in the habit of sunning themselves in a
particular spot to which they return again and again if disturbed. The females make
their burrows more often in hard soil, such as, for example, native foot-paths, than
in loose sand. I cannot recall ever having found them nesting in the sandbanks of
rivers, which are favourite nesting-sites for Bembix. The great differences in the
length of the tongue and galea amongst the species might lead one to suppose that
they are oligotropic, but this supposition cannot be maintained in view of the fact
that species with those organs short, medium, and long have been taken visiting the
same flowering bushes, such as Combretum and Zisyphus.
Tachytes observabilis Kohl
1894, Ann. naturh. Hofmus. Wien, 9: 295.
Lake Anasa, Ethiopia, April 1948, I 3.
Tachytes basilica Guérin
(Fic. 36)
1844, Icon. régne anim.: 440.
Tillembeya, River Niger, French Sudan, September 1944, I 3.
Tachytes guichardi sp. n.
(Fic. 37)
dg. 13mm. long. Black. Abdomen flavo-ferruginous, the anterior face of the first
tergite and the greater part of the middle of the first three sternites blackish. Tibiae
and tarsi pale reddish-brown. Wings hyaline, slightly yellow, the apex beyond the
cells pale brown. Head, thorax, and base of first tergite with yellowish pilosity.
Tergites with a pale yellow decumbent pubescence arranged in a chequered pattern
as in the basilica group. Vertex, mesonotum, and scutellum very finely punctured,
the interspaces barely larger than the punctures. Epinotum and mesopleura very
finely reticulate-punctate. Sternites moderately shining, very finely and closely
punctured.
Galea more than two and a half times longer than the scapes. Second joint of the
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 147
flagellum three times longer than wide, the three to six joints moderately arcuate
below.
Interocular distance on the vertex a little shorter than the second joint of the
flagellum. Dorsum of the epinotum about one and a half times wider at the base than
long. Seventh tergite trapezoidal, the apical margin wide and feebly convex. Basal
joint of anterior tarsi with five to six spines. The outer paramera of the genitalia
densely villose below, the membranous apical portion narrow and very long.
Dakar, Senegal, August 1943, I 3.
Allied to neavet Turner from which it differs by the wider vertex and the flavo-
ferruginous abdomen. In the colour of the abdomen it resembles basilica Guérin,
which, however, is a larger insect with a shorter galea.
Tachytes dilaticornis Turner
1917, Aun. Mag. Nat. Hist. (8) 20: 23.
Jowaha, N. of Debra Sina, 6,000 ft., Ethiopia, August 1946, I dg.
Tachytes hamiltoni Turner?
Loc. cit., supra: 17.
Q. 15 mm. long. This is probably the hitherto undescribed 9 of this species. The
pilosity and pubescence of the head and thorax is brassy golden. The femora, except-
ing the basal half of the middle pair, are flavo-ferruginous. First four tergites with
apical fasciae of silvery pubescence, the fifth and the sides of the fourth with a pre-
apical row of long, porrect, and brown setae, the pygidial area with dark coppery
pubescence. Second sternite over the basal two-thirds dull and microscopically,
closely punctured, the apical third, like the following sternites, smooth, shining, and
with a few large setigerous punctures. Interocular distance on the vertex very little
shorter than the second joint of the flagellum, the latter three times longer than wide.
Galea as in the 3, twice as long as the scapes. Basal joints of the anterior tarsi with
six spines on the outer margin, the pubescence on the middle and hind tibiae golden.
Accra, Gold Coast, July 1946, I 9.
The type of the species, a f, was taken in British East Africa, so this 2 may perhaps
be a distinct species, but the main characters suggest that the identification is correct.
Tachytes habilis Turner
(Fic. 38)
Loc. cit., supra: 35.
Koundé-Saré, French Sudan, August 1945, 4 ¢¢.
The locality of the type of the species is near Fort Johnstone, Nyasaland.
Tachytes versatilis Turner
Loc. cit., supra: 34.
Moggio, Ethiopia, April 1948, 1 3.
148 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Tachytes neglecta Turner
Loc. cit., supra: 32.
Tillembeya, River Niger, French Sudan, September 1944, I g.
The genitalia resemble those of habilis Turner, but the apical portion of the outer
paramera are dilated, or spoon-shaped, and not parallel-sided as in that species.
Tachytes lamentabilis sp. n.
(Fic. 39)
g. 14mm. long. Black. Face and clypeus with pale golden pubescence and pilosity.
The rest of the head, the thorax, and base of first tergite with dull yellow pilosity,
short on the mesonotum. First four tergites with apical fasciae of grey pubescence,
sternites 2-4 with narrow, thin, and yellowish-grey apical fasciae, the fifth and sixth
with brown fasciae. The seventh tergite closely punctured and with fulvous pubes-
cence. Wings flavo-hyaline, without any apical infuscation, the veins brown. Vertex
with a few small and shallow punctures. Dorsum of thorax closely and microscopi-
cally punctured. Second sternite shallowly and sparsely punctured in the middle,
more closely and very finely at the sides, the following sternites closely and strongly
punctured in the apical half, the basal half smooth and shining. Galea half as long
again as the scapes. Clypeus subcarinate lengthwise in the middle. Interocular
distance on the vertex equal to the length of the first two joints of the flagellum.
The second joint is only twice as long as wide. Dorsum of the epinotum short, as
long as the scutellum. Seventh tergite subtriangular, the apex narrowly truncate.
Basal joints of the anterior tarsi with six spines.
2. 16mm. Clypeus very convex lengthwise, but not subcarinate. Galea as long
as the scape. Interocular distance on the vertex equal to a little more than the length
of the second joint of the flagellum. The latter nearly three times longer than wide.
Otherwise like the ¢.
Aburi, Gold Coast, December 1941, I 3, I 9.
Owing to the scanty pilosity and pubescence this species looks very different from
the other species of the genus. To the naked eye, it looks more like some of the larger
species of Livis (Motes) of the sepulchralis group.
Tachytes compacta sp. n.
Q. 12-14 mm. long. Black. First tergite at the sides and on the apical margin,
the apical margin of the second tergite, the base of the second sternite, and apical
margin of the third sternite, somewhat reddish.
Clypeus, face, temples, and the thorax with a dense, pale brassy pubescence, that of
the thorax paler or almost silvery, and very dense on the pleura. The long erect
pilosity present in most species of the genus is absent. First four tergites and sternites
with apical faciae of yellowish-grey pubescence, the pygidial area with a dense, fine,
and golden pubescence.
Extreme apex of the hind femora and the basal third of the hind tibiae with pale
golden pubescence, the apical two-thirds with silvery pubescence. Wings short,
hyaline, the veins ochreous. Puncturation of the vertex and thorax and second
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 149
sternite close and microscopic, the second sternite dull except on the apical margin,
the third to fifth sternites shining and very sparsely punctured, but with the apical
fourth dull and finely punctured, the sixth shining, sparsely and fairly coarsely
punctured. Galea shorter than the scape. Apical margin of the clypeus narrowly
reflected. Interocular distance on the vertex equal to the length of the second joint
of the flagellum plus half of the first. Epinotal dorsum as long as the scutellum. Basal
joint of the anterior tarsi with six yellow spines on the outer margin.
Kaffrine, Senegal, August 1943, 2 99.
Closely related to T. erynnis Turner, from which it differs by the colour of the
pubescence, the absence of pilosity, the colour of the first two abdominal segments,
and the shining third sternite. The smaller specimen has the first two tergites paler,
or yellowish-red.
Tachytes fucata sp. n.
9. 15mm. long. Head and thorax black, abdomen ferruginous, the base of the
third and fourth tergites, the whole of the fifth, black. The middle and apical margin
of the fourth and fifth sternites and the whole of the sixth blackish. Legs ferruginous,
the upper side of the fore and middle femora blackish, the posterior coxae and tro-
chanters fusco-ferruginous. Wings clear hyaline, the veins ochreous. Pubescence
on the head and thorax dense, yellowish-silvery, on the pleura and sterna silvery,
the face and epinotum also with a short, exserted whitish pilosity. First four tergites
with apical fasciae of silvery pubescence. Sternites shining, the puncturation micro-
scopic. The pygidium elongate triangular, four-fifths longer than wide at the base,
clothed with dark golden pubescence.
Galea half as long as the scape. Interocular distance on the vertex equal to very
little more than the length of the second joint of the flagellum ; the second joint two
and one-third times longer than wide at the apex. The erect pilosity on the dorsum of
the epinotum is confined to the sides, which gives a false impression of that part
having a median triangular field. Basal joint of the anterior tarsi with five spines
on the outer margin.
Aleg, S. Mauritania, September 1943, 1 9; Tillembeya, River Niger, French Sudan,
August 1944, I 9.
_ Possibly related to T. sjéstedti Cameron, but that species has only the three basal
abdominal segments red, and the abdomen much broader at the base, or conical,
whereas in this species it is lanceolate. The specimen from Tillembeya has the wings
slightly tinged with yellow, and the third and fourth abdominal segments less stained
with black.
Tachytes griseola sp. n.
(Fic. 40)
3. 7°5-8:5 mm. long. Black. Tarsi brownish-ochreous, the apical joint paler.
Sixth to ninth joints of the flagellum red below. Wings hyaline, the veins and the
tegulae pale ochreous. Clypeus and face with a silvery pubescence and a scanty grey
pilosity. Thorax with a sparse decumbent pubescence and a long, white pilosity,
longest on the sides of the thorax and epinotum. The anterior and middle femora also
150 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
with the same and dense pilosity. Abdomen with a fairly long, adpressed greyish
pubescence, directed inwards on the basal half of the tergites and outwards on the
apical half, where it is denser and forms transverse fasciae. Seventh tergite with
silvery pubescence. Vertex, mesonotum, and scutellum very finely and closely
punctured (30 diameters), the dorsum of the epinotum closely and somewhat obliquely
rugulose-recticulate (16 diameters). The declivity dull, closely and transversely
striolate. Sternites shining, the second closely and finely punctured, the following
segments more coarsely and less closely.
Galea shorter than the scape. Vertex wide, the interocular distance at the base of
the eyes being only two and a half times greater than on the vertex, where it is equal
to very little more than the length of the first two joints of the flagellum. The second
joint of the flagellum is short, being not quite twice as long as wide at the apex.
Dorsum of the epinotum as long as the scutellum and metanotum united. Basal
joint of the anterior tarsi with three spines on the outer margin.
2. 10mm. long. The extreme apical margins of the first five abdominal segments
fusco-ferruginous. Pygidial area clothed with a dark golden pubescence. Second
sternite fairly dull, closely and microscopically punctured. Interocular distance on
the vertex equal to the length of the first two joints of the flagellum. Basal joint of
the anterior tarsi with five white spines on the outer margin. Otherwise like the 3.
Labadi, Gold Coast, March 1941, 5 gg, 1 9; Ufdem, Ethiopia, August 1945, I 4,
19. These are a little larger than the specimens from the Gold Coast, 9, Io mm.,
I 9, 10-5 mm. long.
This species and the one described below belong to a group of small Tachytes in
which the tarsi are pale reddish-yellow and the vertex is wide. It includes T. lepida
Arnold, T. nigroannulata Bischoff, and T. stlverlockt Turner, from all of which it
differs by the genitalia.
Tachytes rufitibialis sp. n.
(Fic. 41)
3, 7°5-g mm. long. Closely related to preceding species, T. griseola, from which it
differs as follows. Third to ninth joints of the flagellum dark brown above, reddish-
yellow below. Apices of the femora and the tibiae reddish-yellow, like the tarsi.
Pubescence on the clypeus and face pale golden. Pubescence and pilosity on the
thorax yellowish-grey. Tergites 1-4 with yellowish-grey fasciae covering the apical
third of the segments, the basal portion with only a microscopic black pubescence,
Sculpture of the epinotum like that of griseola, but stronger, the declivity moder-
ately shining.
Interocular distance at the base of the eyes two and a third times greater than on
the vertex, where it is equal to the length of the first two joints of the flagellum plus
one-fourth of the third.
Pygidial area more rounded at the apex, covered with pale golden pubescence.
Outer paramera of the genitalia with the apical portion more acute, not bluntly
angular on the outer margin but evenly convex. Otherwise like griseola.
Tillembeya, River Niger, French Sudan, August 1944, 5 gd.
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 151
Tachytes discrepans sp. n.
(FIG. 42)
g. 9°5 mm. long. Black. Face and clypeus with silvery pubescence. Thorax with
a scanty yellowish-grey pubescence, the spinotum also with a sparse pilosity of the
same colour. Wings hyaline, the veins ochreous. Tergites 1-4 with apical fasciae of
grey pubescence, the seventh tergite with silvery. Vertex and dorsum of the thorax
very finely recticulate-punctate. Sternites not quite dull, very finely punctured, the
44b
Fic. 42. Tachytes discrepans, genitalia, x27. Fic. 43. Tachytes falcigeva, genitalia, x 18.
Fic. 43a. Tachytes falcigera, apical joints of the flagellum, x18. Fics. 44 and 44a. Odontosphex
bidens 3, head, x18. Fic. 44b. Odontosphex bidens 3, flagellum, x27. Fic. 44c. Odontosphex
bidens 3, wings, X12.
second, third, and fourth with a thin greyish pubescence, more abundant on the apical
margins, forming fasciae. Anterior margin of the clypeus produced into a very short
lobe over the median fourth. Galea shorter than the scape.
Ninth, tenth, and eleventh joints of the flagellum compressed dorso-ventrally on
their inner halves and strongly carinate on the inner margin, the second joint twice
as long as wide at the apex.
Interocular distance at the base of the eyes five-sevenths wider than on the vertex,
where it is equal to the length of the first two joints of the flagellum. Anterior meta-
tarsi with four white spines on the outer margin. Apical portion of the outer para-
mera of the genitalia very thin, lamelliform, twisted, and slightly dilated close to the
apex.
2. 10mm. long. Tarsi, excepting the basal joint, fusco-ferruginous. Pubescence
ENTOM. 2. 3 S
152 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
and pilosity on the head and thorax yellowish-silvery, on the underside of the fore
and middle femora silvery. Tergites 1-4 with a thin greyish pubescence on the basal
part, and with broad apical silvery fasciae.
Pygidial area with golden pubescence. Second sternite dull, microscopically
punctured, the remaining sternites shining, fairly coarsely and very sparsely punc-
tured. Interocular distance on the vertex slightly less than the length of the first two
joints of the flagellum. The second joint of the latter two and a half times longer than
wide, the ninth to eleventh joints not dilated nor carinate. Anterior margin of the
clypeus without a median lobe, otherwise like the ¢.
Kaolack, Senegal, August 1943, I 3, I 9.
The ¢ differs from all the other African species in the shape of the genitalia and,
excepting iustabilis and the following species, by the ninth to eleventh joints of the
flagellum. The 9 resembles T. stlverlockt Turner and T. disputabilis Turner, from both
of which it can be distinguished by the colour of the pubescence on the sixth tergite,
the sculpture of the thorax, particularly of the epinotal dorsum, and by the number of
spines on the outer margin of the anterior matatarsi. From instabilis Turner 9 it may
be distinguished by its smaller size and wider vertex, which in instabilis is equal to
only the length of the second joint of the flagellum.
Tachytes falcigera sp. n.
(Fics. 43, 434)
3. 14mm. long. Black. Wings hyaline, and faintly tinged with yellow, the veins
ochreous. Face and clypeus with yellowish-silvery pubescence. Thorax with a rather
sparse yellowish-grey pubescence, and on the epinotum with a sparse pilosity of the
same colour. Tergites 1-4 with apical fasciae of silvery pubescence, the seventh with
yellowish silvery.
Vertex, mesonotum, and scutellum very closely and finely punctured, the following
sternites shining, sparsely and coarsely punctured. Galea a little more than half as
long as the scapes. Anterior margin of the clypeus flattened.
Interocular distance at the base of the eyes twice as great as on the vertex, where
it is equal to the length of the second joint of the flagellum. The second joint is two
and a third times longer than wide at the apex, and the ninth to eleventh joints are
compressed on their inner halves and carinate, as in the preceding species.
Dorsum of the epinotum a little shorter than the scutellum and metanotum united.
Anterior metatarsi with five white spines on the outer margin. Outer paramera of the
genitalia abruptly attenuated apically, the attenuated part sickle-shaped and slightly
widened at the apex, the inner rod-shaped paramera emit two long and thin hairs at
the apex.
Aleg, S. Mauritania, September 1943, I J.
In my key to the species (1923, Aun. Transv. Mus. 9) this runs down close to
T. instabilis Turner, but the interocular distance is shorter than in that species. The
genitalia of the specimen from Willowmore, Cape Province, which I identified as
instabilis, are quite unlike those of falcigera, resembling those of T. bulawayoensis
Bischoff.
ee ee wig
ER gga me ae
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 153
Tachytes seminuda sp. n.
. 10-5 mm. long. Black. First two abdominal segments and the third excepting
the apical margin red. Last three joints of the anterior tarsi and apical joint of the
lower and hind tarsi, ferruginous. Wings hyaline, the veins pale brown. Clypeus
and sides of the face and the bottom of the temples with a short and by no means
dense silvery pubescence. On the thorax there is a little grey pubescence on the
lateral margins of the mesonotum and on the epinotum, but the outstanding pilosity
present in nearly all other species of the genus is absent.
The first three tergites have inconspicuous apical fasciae of silvery pubescence,
and the pygidial area has dark golden pubescence. Vertex microscopically and closely
punctured (30 diameters), the mesonotum closely and transversely rugulose (25
diameters), the dorsum of the epinotum reticulate-punctate and obliquely striate at
the sides of the base (30 diameters). Second sternite dull, the third and following
sternites shining, with a few large punctures near the apical margin. The abdomen
is widest in the middle, or lanceolate, instead of being obconical or widest across the
basal segment, as in the other species of the genus.
Galea less than half as long as the scapes. Eyes widely divergent below, the inter-
ocular distance at the base of the eyes being very nearly three and a half times greater
than on the vertex, where it is equal to the length of the second joint of the flagellum.
Clypeus short, nearly four times wider than long. Flagellum slender, the second
joint fully two and a half times longer than wide at the apex, all the following joints
at least twice as long as wide. Second to fifth sternites and the fifth tergite with a row
of brown setae on the apical margin. Anterior metatarsi with five yellow spines on
the outer margin. Pygidial area triangular, longer than wide at the base.
Tillembeya, River Niger, French Sudan, August 1944, 2 99.
This species differs from all the others known to me by the paucity of pubescence
and pilosity, and the shape of the abdomen. Superficially it looks much more like a
Tachysphex, e.g. T. miniatulus Arnold, than a Tachytes.
PHYTOSPHEX gen. n.
Characters
9. Anterior tarsi without a comb, the metatarsus with a few small spines on the
underside and at the apex. Hind tibiae and metatarsi with a row of long spines on
the upper side. Pygidium without a marginate dorsal face.
g. Anterior metatarsi without long spines on the outer margin, the hind tibiae and
metatarsi as in the 9. In both sexes the vertex is wide, the eyes not being strongly
convergent above as in Tachysphex, and the sculpture of the head and thorax is
strong, consisting of large and widely spaced punctures on the head and thorax, and
a coarse reticulation on the epinotum. Anterior margin of the clypeus produced in
the middle into an acute, wide, and triangular tooth. Otherwise like Tachysphex.
Genotype: Tachysphex turnert Arnold.
Not only does this species differ from Tachysphex in the sculpture, but the nesting
habits are, according to the late Dr. Brauns’s observations, totally different. Instead
of burrowing in the ground, it nests in hollow stems of Aloe, Datura, &c., and the
partitions between the cells are formed of earth and little pebbles.
154 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Phytosphex turneri Arnold
1923, Ann. Transv. Mus. 9: 165.
A slight variety, differing from the type of the species in having the second to
fourth tergites blackish in the middle and on the apical third of the segments.
Adis Ababa, Ethiopia, April 1948, I 9.
ODONTOSPHEX gen. n.
Characters
36. Mandibles not excised on the outer margin. Clypeus short, not longer than the
distance between its hind margin and the antennal sockets, the anterior margin with
two long and thin teeth near the middle. Eyes strongly convergent above, the vertex
narrower than the anterior ocellus. Posterior ocelli oval, flattened, and almost
obsolete. Second to seventh joints of the flagellum short, wider than long, the seventh
and tenth swollen behind. Dorsum of the pronotum linear, lying well below the level
of the mesonotum. Episternal suture absent. Dorsum of the epinotum with a defined
and broad median area. Eighth sternite emarginate and bidentate. Anterior tarsi
without cilia or spines. Middle tibiae with two calcaria, the inner one very small.
Posterior tibiae and tarsi without long spines on the upper side, the tibiae with a row
of very short spines hardly longer than the pubescence. Forewings with three cubital
cells, the third less than twice as long on the cubitus as on the radius, the second
receiving both recurrent veins. Hindwing: the cubitus emitted well before the end
of the submedial cell.
Genotype: O. bidens.
The genus is remotely related to Tachysphex, from which it differs, inter alia, by
the venation, the absence of the episternal suture, and the form of the flagellum.
Odontosphex bidens sp. n.
(FIGS. 44, 44a-c)
3. 6mm. long. Black. Middle third of the mandibles rufescent. Flagellum
ochreous, the first four joints slightly infuscated. Apical margins of the abdominal
segments testaceous. Tibiae, tarsi, and the hind femora pale reddish-ochreous.
Wings hyaline, tegulae and pterostigma pale yellow, the veins pale brown. Clypeus
and face up to the anterior ocellus, temples, underside of the anterior femora, sides
and anterior angles of the mesonotum, mesopleura, and base of the epinotal dorsum,
covered with silvery pubescence. A small patch of similar pubescence on each side
of the scutellar and metanotal disks.
Sides of the epinotum, and the declivity excepting the middle, with exserted and
fairly dense silvery pilosity. Abdomen with a very fine, grey, and decumbent
pubescence, and on the apical margins of the first five tergites fasciae of silvery
pubescence. Face, from the antennal sockets to the anterior ocellus, dull, micro-
scopically and closely punctured, slightly convex but without any swellings. Ocellar
area and vertex shining, sparsely punctured. On the vertex a carinula on each
side margins the eyes.
Mesonotum, scutellum, and metanotum shining, finely punctured, the interspaces
three or four times wider than the punctures, except on the posterior fourth of the
-
ooo
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 155
mesonotum and on the metanotum, where they are smaller. Mesopleura dull, closely
punctured (35 diameters), a short transverse crest extends from the hind margin
forwards, just above the middle coxae. Dorsum of the epinotum nitidulous, a little
more strongly and more closely punctured than the scutellum, the margins of the
median area indicated by a narrow depressed line. A median carina extends from
the base to near the apex of the area. Declivity dull, with a wide pit below the brow.
Metapleura and sides of the epinotum fairly dull, closely punctured (45 diameters).
Tergites and sternites almost dull, very closely and finely punctured (30 diameters).
Head wider than the thorax. Clypeus five times wider than long, the anterior margin
feebly convex and with an acute tooth, longer than wide at the base, on each side of
the middle. Eyes strongly divergent below, the interocular distance across the base
of the clypeus being eight times greater than across the vertex, where it is equal to
the length of the second joint of the flagellum. First joint of the flagellum very small,
not globose, the second joint three-fourths wider than long, the first to seventh wider
than long, the seventh to tenth swollen below, the apical joint one and a half times
longer than wide. Scutellum twice as wide as long, dorsum of the epinotum about
three times wider at the base than long, as long as the scutellum, the junction with
the subvertical declivity rounded. First tergite about one and a half times wider
behind than long.
Basal joint of the anterior tarsi thin and very long, as long as the remaining joints
united.
Hind femora and tibiae slightly compressed transversely, the tibiae with a row of
about twelve very small scalariform teeth on the upper margin (45 diameters), apical
margin of seventh tergite transverse. First abscissa of the radius slightly shorter than
the third, and a little longer than the second.
Aleg, S. Mauritania, September 1943, I ¢.
Tachysphex fluctuatus Gerstaecker
1857, Mber. Akad. Wiss. Berl.: 510.
Arnold, 1945, Sphecidae of Madagascar: 97.
Labadi, Gold Coast, March 1941, 1 ¢; Meisso, Ethiopia, September 1945. The
second specimen has the epinotum much more strongly and more widely reticulate,
and more pilose than in the typical form.
Tachysphex bruneiceps Arnold
1923, Ann. Transv. Mus. 9: 153.
River Hawash, Ethiopia, June 1946, I 9.
Tachysphex egregius Arnold
1924, Ann. Transv. Mus. 11: 273 3.
1929, Ann. Tvansv. Mus. 18: 384 9.
Labadi, Gold Coast, May 1941, 1 g, 1 9; Tillembeya, River Niger, French Sudan,
August 1944, I 9.
7
:
156 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Tachysphex theseus sp. n.
(Fie. 45)
9. 10mm. long. Black. Apical margin of the fourth and the whole of the fifth and
sixth abdominal segments ferruginous. Tarsi ferruginous, the apical joints a little
darker. Wings hyaline, the veins pale brown. Face and clypeus with pale golden
pubescence, thorax with a very short, sparse, and grey pubescence. Clypeus and face
dull, very finely and closely punctured, the vertex nitidulous and not quite so closely
punctured as the face, the temples smooth and shining.
48
Fic. 45. Tachysphex theseus 9, Xc. 10. Fic. 46. Gastrosericus fluviatilis 9, x16. Fic. 46a.
Gastrosericus fluviatilis 3, clypeus, X16. Fic. 46b. Gastrosericus fluviatilis 92, wing, xc. 10.
Fic. 47. Gastrosericus senegalensis 9, X16. Fic. 48. Palarus rothschildi croesus 2, pygidium, x Io.
Mesonotum shining, punctured, the punctures about twice as large as those on the
face, the interspaces from two to three times wider than the punctures, except on the
margins which are very closely punctured. Scutellum and metanotum shining, their
puncturation a little smaller on the face. Mesopleura finely and very closely punc-
tured. Dorsum and declivity of the epinotum dull, fairly closely reticulate, the sides
of the epinotum shining, transversely striate. Abdomen dull, impunctate, the
pygidial area twice as long as wide at the base, the apex narrowly truncate.
Median area of the clypeus nearly twice as wide as long, impressed behind the
anterior margin which is feebly concave in the middle. Supra-antennal tubercles
prominent. Interocular distance on the vertex very nearly as long as the first two
joints of the flagellum. Behind the ocellar area there is a well-defined angular depres-
sion. Flagellum slender, the second joint nearly three times longer than wide at the
apex, the third joint a little longer than the second. Dorsum of the epinotum two
and two-fifths wider at the base than long. Fourth joint of all the tarsi very short,
yeaa
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 157
wider than long, the claws asymmetrical, the outer claw of the anterior tarsi larger
than the inner, and smaller than the inner of the middle and hind tarsi.
Accra, Gold Coast, September 1941, I 9.
Related to T. hippolyta Arnold and T. harpax Arnold and having the claws
asymmetrical as in them, but differing from both by the length and sculpture of the
epinotum, the slender flagellum, and the colour of the apical abdominal segments.
Tachysphex panzeri van der Linden var. aethiopicus Arnold
1923, Ann. Transv. Mus. 9: 167.
Labadi, Gold Coast, March 1941, 3 3d.
Atelosphex lugubris Arnold
1924, Ann. Transv. Mus. 11: 72.
Accra, Gold Coast, September 1941, I 9.
This differs to a slight degree from the type of the species, but hardly enough to
justify a varietal name. The puncturation of the mesonotum in the middle is sparser,
the dorsum of the epinotum is more reticulate, and the pygidial area is narrower.
In the coarse puncturation and the absence of spines along the length of the hind
metatarsi, the genus shows some affinity with Phytosphex Arnold, but in the latter the
epicnemial suture is present.
Parapiagetia capensis Brauns var. ferox Arnold
Brauns, 1910, Disch. ent. Z.: 666.
Arnold, 1922, Ann. Transv. Mus. 9: 135.
Ufdem, Ethiopia, August 1945, 1 9; Aleg, S. Mauritania, September 1943, I 9,
Dakar, Senegal, August 1943, I 9.
The specimen from Dakar is a slight variety, having the last three abdominal
segments blackish.
Gastrosericus neavei reversa subsp. n.
G. neavei Turner, 1913, Tvans. Ent. Soc. Lond.: 754.
9. 8mm. long. The corners of the median lobe of the clypeus are not rectangular
as in neavet i. sp. but produced into blunt teeth, and the wings are slightly darker.
Otherwise it does not differ except in one very remarkable character in which it also
differs from all other Ethiopian Sphecids known to me.
In neavet i. sp. the dorsum of the epinotum is glabrous, but in this race on each side
of the median transversely rugose area there is a coarse, obliquely exserted and silvery
pubescence, directed cephalad. In the ¢ this pubescence is less abundant, but plainly
apparent if the insect is looked at caudad. Otherwise like the type of the species.
Tillembeya, River Niger, French Sudan, August 1944, 4 292, 446.
Gastrosericus fluviatilis sp. n.
(Fics. 46, 46a and 46D)
2. 8mm. long. Black. The following parts pale yellow: basal half of the mandibles,
the apex of the scapes, tegulae, apical half of the anterior femora, apical third of the
158 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
middle and extreme apex of the hind femora, tibiae and first two joints of tarsi, the
last three joints pale ferruginous. Wings hyaline, faintly smoky, the veins brown.
Clypeus, lower half of the face, temples, dorsum of the pronotum, mesosterna, and
lower half of the mesopleura with silvery pubescence. Epinotum with short, erect
and white pubescence. Tergites 1-4 with apical fasciae of grey pubescence, the
pygidial area with dark golden pubescence. Sternites 3-6 shining and impunctate,
the rest of the body dull, closely and microscopically punctured, the largest punctures
being on the scutellum and epinotum (30 diameters). Clypeus short, five times wider
than long, carinate lengthwise in the middle, and with a very short lobe on the
anterior margin ; between the lobe and the lateral angles there are two small teeth on
each side. Interocular distance at the base of the eyes two-thirds greater than across
the vertex, where it is equal to the length of the first five joints of the flagellum.
Temples with a triangular tooth near the middle.
Second joint of the flagellum slightly shorter than the third and not quite twice
as long as wide. Lower third of the face excavated, the upper margin of the excavated
area continued as a groove margining the eyes as far as the level of the anterior
ocellus.
Propleura with a transverse torus on each side in front of the hind margin. Anterior
coxae triangular, the anterior margin raised and thickened. Pronotal collar barely
as long as the first joint of the flagellum and lying much below the level of the meso-
notum. Dorsum of the epinotum fully twice as wide at the base as long, joining the
oblique declivity in a wide curve. First tergite a little longer than wide at the apex,
and like the second tergite a little swollen in front of the hind margin. Pygidial area
elongate-triangular, slightly shining and sparsely punctured at the base. Anterior
metatarsi with six short spines on the outer margin. Second discoidal cell petiolate,
the first abscissa of the radius fully three times longer than the second.
g. 6mm. long. Middle and hind tibiae with a fuscous macula on the inner side,
the last four joints of the hind tarsi black. Pubescence on the clypeus and face golden.
Clypeus produced in the middle into an acute tooth. Joints of the flagellum very
short, the second one and a half times longer than wide. Temples without a tooth,
anterior coxae less thickened in front than in the 9. Otherwise like the 9.
Tillembeya, River Niger, French Sudan, August 1944, I g, 6 99.
Related to ¢urnert Arnold, from which it differs, inter alia, by the shape of the
7
om
1 ee yr
ee mS gem yt a=
clypeus and anterior coxae, the much finer puncturation of the thorax, and the ©
venation.
Gastrosericus senegalensis sp. n.
(Fic. 47)
9. 6:5 mm. long. Head and thorax black, abdomen pale ferruginous. Mandibles
yellow at the base, the apex of the scapes ochreous on the upper side. Femora fusco-
ferruginous, the fore and middle pairs yellow at the apex, tibiae and tarsi pale ferru-
ginous, the tibiae pale yellow on the upper side. Wings hyaline, the veins ochreous.
Head and thorax with a very short adpressed and yellowish-silvery pubescence,
without any long pilosity. Tergites with a thin and microscopic greyish pubescence.
The whole body dull and, excepting the pygidial area, very closely and microscopically
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 159
punctured (50 diameters), the dorsum of the pronotal collar with traces of transverse
rugulosity. Pygidial area shining, sparsely punctured, twice as wide at the base
as long.
Clypeus three times wider than long, subcarinate in the middle, the anterior margin
convex and without teeth. Eyes divergent below, the interocular distance at their
base half as great again as across the vertex, where it is equal to the length of the first
three joints of the flagellum plus half of the fourth. The second joint about twice as
long as wide, and as long as the third. Temples unarmed. Dorsum of the pronotum
narrowed anteriorly, fairly long, as long as the first joint of the flagellum. Dorsum of
the epinotum twice as wide at the base as long, with a very indistinct basal triangular
area. Anterior metatarsi with five long spines on the outer margin. First abscissa of
the radius twice as long as the second, the recurrent veins contiguous on the second
cubital cell.
Dakar, Senegal, August 1943, I @.
Smaller than the Egyptian G. waltii Spinola and without the long pubescence on
the head and thorax and rugose sculpture of the epinotum present in that species.
It differs from G. capensis Brauns in the much finer sculpture and dull surface of the
head and thorax and by the absence of a long and woolly pubescence on those parts.
Liris (subgen. Motes) nugax Kohl
1894, Ann. naturh. Hofmus. Wien, 9: 302.
Dilla, Ethiopia, April 1943, 1 9; Didessa River, Ethiopia, June 1946, 1 9.
Liris (Motes) croesus Smith
1856, Cat. Hymen. B.M. 4: 284.
Diafarabé, Senegal, August 1944, I 3.
Liris (Motes) solstitialis Smith
1856, Cat. Hymen. B.M. 4: 283.
Aburi and Accra, Gold Coast, August to September 1941, I g, I 9.
Liris (Motes) rugifera Turner
1918, Aun. Mag. Nat. Hist. (9), 5: 362.
Accra, Gold Coast, September, I 9.
Liris (Motes) thysanomera Kohl
1894, Aun. naturh. Hofmus. Wien, 9: 302.
Accra and Aburi, Gold Coast, December 1941, 3 29.
Liris (Motes) antaca transvaalensis Cameron
Arnold, 1945, Sphecidae of Madagascar: 126.
Adis Ababa and River Hawash, Ethiopia, August and June, 2 29.
ENTOM. 2. 3 T
160 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Liris (Motes) egregia Arnold
1929, Ann. Transv. Mus. 18: 395.
Adis Ababa and Dilla, Ethiopia, June, 2 33.
Liris (Motes) setigera Arnold
1940, Ann. Transv. Mus. 20: 123.
Tillembeya, River Niger, French Sudan, August 1944, I 3.
Liris (Motes) denticulata Turner
1920, Ann. S. Afr. Mus. 17: 496. ;
Accra and Labadi, Gold Coast, September and March 1941, 2 29.
These represent a slight variety in which the tarsi are ferruginous instead of
brownish-black.
Liris (Motes) flavitincta Arnold
1940, Ann. Transv. Mus. 20: 141.
Accra, Gold Coast, September 1941, I 9.
Liris (Motes) abyssinica Arnold
1933, Ann. Mag. Nat. Hist. (10) 77: 353.
Fiché, Ethiopia, October 1945, I 3.
Liris (Motes) gracilicornis Arnold
1923, Ann. Transv. Mus. 9: 250.
Labadi, Gold Coast, March 1941, I Q.
Liris nigricans Walker race reticulata Saussure
1871, List Hymen. Egypt: 21.
Arnold, 1945, Sphecidae of Madagascar: 132.
Dilla, Ethiopia, April 1948, 2 3.
Palarus rothschildi croesus subsp. n.
(Fic. 48)
P. vothschildi Magretti, 1908, Bull. Mus. Hist. Nat. Paris: 189, 3.
The insect described hereunder agrees in the main with the description of roths-
childi, but there are differences in structure as well as in colour. Not much importance
attaches to the latter, and since it appears to me improbable that Magretti over-
looked the structural characters it seems more prudent to treat this as a subspecies.
3. 95mm. long. The yellowish-white markings on the scutellum are confined to
the posterior angles, the epinotum is entirely black, the pale maculae on the tergites
are present on the fifth and sixth as well as on the four basal tergites. Mesopleura
without any transverse striation. Excepting a small round and smooth area in front
of the hind margin, the whole of the epinotal dorsum is strongly reticulate-rugose.
Sixth sternite with an acute curved tooth on each side, the seventh bluntly triangular,
the apical without two small teeth. Otherwise like rothschildi i. sp. In the original
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 161
description ‘supra ocellum magnum’ is evidently a lapsus calami, the triangular
swelling on the face being below, and not above, the anterior ocellus.
2. 12mm. long. Anterior half of the mesopleura dirty yellow, stained in parts with
red. The dorsum of the epinotum with a yellowish-white macula on each side. The
pale maculae on the first five tergites cover the greater part of the segments but do
not reach the basal and apical margins. Second to fourth sternites ferruginous,
blackish at base and apex and with a narrow yellow apical line, interrupted in the
middle. Sixth tergite and sternite ferruginous.
Supra-antennal swelling less triangular than in the g. Interocular distance on the
vertex equal to a little more than the length of the first joint of the flagellum. The
second joint is a little more than two and a half times longer than wide and one-
fourth longer than the third. First tergite with a short and acute tooth on each side
of the vertical anterior face and near its dorsal boundary, the sides of the dorsal face
not angular posteriorly as in the g. The transverse crest on the second sternite is not
broad and flat on its vertical face as in the g, but is low and has a trenchant edge
which bears seven small teeth. Pygidium long, the sides with large elongate punc-
tures, the dorsal face dull, very finely, closely, and longitudinally striate, at least
three times longer than wide at the base.
Ufdem, Ethiopia, I 3, I 9.
Related to P. O’Neili Brauns, from which it differs in the g by smaller size, the
colour, the shape of the first tergite, and in the 9 by the colour, the sculpture, and
shape of the epinotal declivity.
Palarus obesus sp. n.
(Fic. 49)
2. 12mm. long. Mandibles pale yellow at the base, ferruginous at the apex.
Clypeus yellowish-white, the rest of the head and the thorax black, the posterior
margin at the pronotum excepting the middle, the tegulae, a spot on the mesopleura
below the tegulae, the posterior lateral margin of the scutellar area and the meta-
notum, yellowish-white. First and second abdominal segments pale ferruginous, the
third to fifth blackish, the sixth ferruginous, the first to fifth tergites with pale yellow
transverse bands, those of third, fourth, and fifth emarginate in the middle of the
posterior margin. Fore and middle femora yellowish-white below, the anterior pair
black above, the middle pair ferruginous above. Hind femora ferruginous, the extreme
apex yellowish, tibiae and tarsi pale ferruginous. Wings hyaline, the veins ochreous.
Clypeus, face, and thorax, excepting the epinotum, with silvery pubescence, very
sparse on the thorax ; temples and epinotum with long white pilosity. Apical margins
of the abdominal segments with a few long yellowish hairs.
Clypeus dull, closely and finely punctured, subcarinate lengthwise in the middle,
half as wide again as long, the anterior margin convex. Face dull, closely and finely
punctured, the space behind the anterior ocelli shining, sparsely punctured, and
sharply carinate. Eyes contiguous on the vertex.
Mesothorax and mesonotum nitidulous, sparsely, finely, and very shallowly punc-
tured. Metapleura smooth. Epinotum dull, microscopically reticulate. Abdomen
162 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
shining, finely and sparsely punctured. Sixth tergite dull, lanceolate, the sides
strongly marginate and with a median carinate area which is longitudinally rugose,
the space between the median area and the sides obliquely rugoso-punctate ; sixth
sternite coarsely punctured, and with a strong median carina.
Flagellum moderately clavate, the second joint nearly three times longer than wide
52a
Fic. 49. Palarus obesus 2, pygidium, Xc.12. Fic. 50. Paranysson oscari 4, first sternite, xc. 12.
Fic. 50a. Paranysson oscari 3, maxillary palp, x27. Fic. 51. Dasyproctus basifasciatus 9,
petiole, x18. Fic. 52. Dasyproctus stevensonianus occidentalis 3, epinotum and petiole, x 18.
Fic. 52a. Dasyproctus stevensonianus occidentalis 3, genitalia, x 24. Fics. 53 and 53a. Oxybelus
coniferus 9, clypeus, X15. Fic. 53b. Oxybelus coniferus, abdomen, x7}. Fic. 53c. Oxybelus
contferus 3, clypeus, X15.
and twice as long as the third, the latter a little longer than wide, the fourth and apical
as long as wide, the other joints wider than long. Dorsum of the epinotum a little
shorter than the scutellum, shallowly depressed in the middle; the declivity vertical,
with a median triangular depression, the apex ventrad. Pygidial area nearly twice
as long as wide at the base. Anterior tarsi with a well-developed comb, the basal
joint with six yellow spines on the outer margin. Second cubital cell with a very short
stalk, the second abscissa of the radius two-thirds longer than the first. The first
recurrent vein meets the first cubital cell not far from the first transverse cubital vein,
the second meets the second transverse cubital vein. Radial cell not appendiculate.
Podor, S. Mauritania, September 1943, 2 29.
Apparently more closely related to the Indian P. comberi Turner than to any of
the African species, differing from it by the shape of the yellow markings on the
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 163
abdomen, the coarse puncturation of the mesonotum, and the colour of the legs.
The shape of the pygidial dorsum appears to be the same, judging by Turner’s
description.
Paranysson oscari Turner
(Fics. 50 and 50a)
1914, Ann. Mag. Nat. Hist. (8) 14: 341, 9.
Aburi, Gold Coast, December 1941, 3 dd.
In this species the maxillary and labial palpi are densely pubescent, and the basal
sternite is very convex, rising caudad, and is sharply truncate a little in front of
the hind margin. In P. helioryctoides Turner the basal sternite is similar, but as it does
not rise so high caudad, the face of the truncation is shorter.
Subfamily PEMPHREDONINAE
Psenulus pauxillus laevior subsp. n.
P. pauxillus, Arnold, 1947, Occ. Pap. Nat. Mus. S. Rhod. 2: 159.
g. 6:5 mm. long. Femora brown. Mesonotum one and a half times wider than
long, wider than in pauxillus i. sp. in which it is only one and two-fifths wider, less
strongly and less closely punctured, and without the feeble transverse ridges between
the punctures. The dorsum of the epinotum is only feebly rugulose at the sides and
the Y-shaped groove is narrower. The interocular distance on the vertex is shorter,
being equal to the length of the first three joints of the flagellum plus half of the
fourth.
Gambeila, Ethiopia, February 1948, 1 g.
Subfamily CRABRONINAE
All the specimens in this collection belong to the species-group Dasyproctus. For
the validity of genera and subgenera in this subfamily, as opposed to species-groups,
I would refer the reader to my remarks on the subject in my Monograph of the
Sphecidae of Madagascar, 1945, p. 157.
Crabro (Dasyproctus) oedignathus Arnold
1933, Ann. Mag. Nat. Hist. (10) 11: 355, 3.
Adis Ababa, Wondo, Akaki, Mount Chillala (8,000-9,000 ft.), Ethiopia, 11 gd,
3 £8.
Q. 11 mm. long (hitherto undescribed). Pubescence on the clypeus pale golden,
the sixth tergite entirely black. The puncturation on the vertex behind the brow of
the face finer, and the sculpture of the epinotum is also finer, the median triangular
area is without longitudinal rugae and is only reticulate-rugose. Mandibles of the
usual shape, not angularly dilated near the base as in the g. Second joint of the
flagellum two and two-thirds longer than wide (twice as long as wide in the @);
the fifth joint not dilated below. Otherwise like the ¢.
164 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Crabro (Dasyproctus) pullatus uniguttatus subsp. n.
C. pullatus Arnold, 1944, Occ. Pap. Nat. Mus. S. Rhod. 2: 30.
g. 8-8-5 mm. long. Pronotal tubercles straw-yellow. The vertex in front of the
anterior ocellus is much more strongly punctured than in pullatus i. sp. The punctures
are about one-fourth as wide as the anterior ocellus and the interspaces are two anda
half times wider than the punctures. The node of the petiole is a little thicker.
Wondo, Ethiopia, April 1948, 4 33.
Crabro (Dasyproctus) basifasciatus sp. n.
(Fic. 51)
9. 95mm. long. Black. The following parts lemon-yellow: scapes, pronotal
dorsum except in the middle, pronotal tubercles, a small spot on the anterior angles
of the scutellum, transverse and narrow bands, interrupted in the middle, at the
extreme base of the third and fourth tergites, the upper sides of the fore and middle
tibiae, and a spot near the apex of the hind tibiae. Tarsifusco-ferruginous. Mandibles
ferruginous. The sculpture hardly differs from that of C. dubiosus Arnold, but under
a high magnification (30 diameters) the mesonotum and scutellum have minute
punctures with interspaces three to four times wider than the punctures. The
posterior corners of the mesonotum and the sides of the scutellum have a few very low
longitudinal rugae.
Clypeus like that of dubiosus, the median lobe a little more deeply excised. The
second joint of the flagellum a little thicker, being two and a third times longer than
wide (nearly three times in dubiosus). The petiole of the abdomen is much shorter
than in dubiosus or in any other species of the group excepting C. ruficaudis Arnold
and C. barkeri Arnold; it is two and a half times longer than wide across the node,
whereas in dubiosus it is nearly three times longer.
Moggio, Ethiopia, April 1948, I 9.
Closely related to dubiosus, from which it differs by the shorter and thicker petiole
and the ground colour of the legs, and the position of the yellow bands on the tergites.
Crabro (Dasyproctus) stevensonianus occidentalis subsp. n.
(Fics. 52 and 52a)
C. stevensoni Arnold, 1926, Ann. Transv. Mus. 2: 369.
C. stevensonianus Arnold (nom. nov.), 1940, ibid. 20: 135.
$. 7mm. long. This race differs from the type of the species as follows: scapes with
a brown macula on the inner side below. Anterior tibiae yellowish-red below, dark
brown above, all the tarsi brownish-black, all the rest of the legs black. The yellow
maculae on the second tergite duller and smaller. The reticulations on the epinotum
are much higher and wider apart.
As in the type of the species 3, the petiole is five times longer than wide across the
node.
Aburi, Gold Coast, December 1944, I 3.
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 165
Crabro (Dasyproctus) bipunctatus Lepeletier var. avius var. nov.
D. bipunctatus Lepeletier, 1834, Ann. Soc. Ent. Fr. 3: 802.
3g. 7:°5-8:5 mm. long. This differs from the type of the species as follows. Tegulae
black. Fore and middle tibiae and tarsi fusco-ferruginous, all the rest of the legs
black. Wings hyaline, slightly smoky, the veins and pterostigma black. Pronotal
collar shorter, shorter than the first joint of the flagellum (distinctly longer in
bipunctatus i. sp.), the median groove much shallower and narrower.
9. 8-5 mm. long. The face and middle tibiae ferruginous. Otherwise like the ¢.
Adis Ababa, Zuquala {9,000 ft.), and Aselle, 4 $$, Lake Bishoftu, Ethiopia, 1 9.
Subfamily OXYBELINAE
Oxybelus coniferus sp. n.
(Fics. 53, 534-0)
9. 6:5-7°8 mm. long. Black. The following parts yellowish-white: a short and
transverse streak on each side of the pronotal dorsum, small spots, sometimes absent,
on the anterior corners of the scutellum, the metanotum, transverse and more or less
lanceolate maculae on each side of the first four tergites. Anterior tibiae dirty white
on the upper side. Apical joint of the tarsi pale brown. Wings hyaline, slightly
smoky, veins dark ochreous to black. Tegulae fusco-ferruginous. Mandibles some-
times yellowish-red on the basal half. Mandibles, clypeus, lower half of the face and
temples with silvery pubescence.
Metanotum and scutellum with erect, brownish-grey pubescence, the pleura,
sterna, and femora below with white and thin pubescence. Median area of the clypeus
shining, the anterior margin convex and with a blunt tooth on each side, the base with
a transversely compressed, and slightly curved conical protuberance, its apex
rounded. The space behind the scapes shining and very finely punctured, the rest
of the head dull, very closely punctured, the interspaces not wider than the punctures ;
on the occiput and temples striated between the punctures. Pronotal dorsum very
short, almost linear in the middle, the lateral angles acute. Mesonotum and scutellum
shining, a little more strongly punctured than the vertex, the interspaces narrower
than the punctures and slightly raised longitudinally, or subrugose. Metapleura
nitidulous, coarsely reticulate-rugose. Metapleura strongly, sides of epinotum finely
and transversely striate. Metanotum shining, longitudinally and not closely costate.
The lateral squama black, rostrate and acute at the apex. Epinotal process canali-
culate, narrowed towards the apex which is excised, half as long again as wide at the
base. Median triangular area of the epinotal dorsum with eight transverse costae
which extend over the lateral areas, the interspaces dull, very finely reticulate.
Declivity of the epinotum dull and transversely rugose outside the median triangular
space.
Abdomen shining, the tergites very sparsely punctured on the yellow areas at
the sides, fairly closely over the basal half and at the apical margin. Pygidial area
dull, closely punctured, triangular, the apex truncate. Second sternite sparsely
and coarsely punctured, excepting the sides which are closely and finely so. Eyes
166 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
moderately divergent above and below, the interocular distance on the vertex equal
to the length of the flagellum, less the apical joint. Posterior ocelli twice as far from
each other as from the eyes. Between the posterior ocelli and the eyes there is a low
and oblique torus. Comb of the anterior tarsi ochreous, the basal joint with six spines
on the outer margin.
3g. 6:2 mm. long. Femora with a dull yellow macula on the upper side at the base.
Face and mesonotum with a slight aeneous tint. Pale maculae on the first to fifth
tergites, smaller than in the 9 and of a chrome-yellow colour. Median area of the
clypeus with a longitudinal carina, wide at the base, projecting as a tooth on the
concave anterior margin. Pygidial area trapezoidal. Otherwise like the 9.
Adis Ababa, Wondo, Ethiopia, August and September, 1945, 12 29, 9 dé.
Related to O. pilosus Arnold, from which it differs in both sexes by the greater size,
in the ¢ by the shape of the clypeus and its basal process, black pronotal tubercles,
stronger puncturation of the mesonotum and tergites, entirely black flagellum, and
colour of the tarsi. In the 3, it differs from pilosus by the colour of the legs and of the
flagellum.
Oxybelus acutissimus Bischoff var. propinquus Arnold
1927, Ann. Transv. Mus. 12: 83.
Haramaia, Ethiopia, May, 1948, 3 99.
Oxybelus pictus Arnold
1927, Ann. Transv. Mus. 12: 115.
Tillembeya, River Niger, French Sudan, September 1944, I 9.
This specimen is a slight variety, differing from the type of the species by the less
close puncturation of the mesonotum, by the stronger sculpture of the lateral areas
of the epinotal dorsum, and by the presence of narrow pre-apical yellow bands on the
third and fourth tergites as well as on the first and second.
Oxybelus curviscutis Arnold
1927, Ann. Transv. Mus. 12: 109.
Accra, Gold Coast, Tillembeya, River Niger, French Sudan, Dakar, Senegal,
and Haramaia, Ethiopia, 5 99°, 4 3d.
The 2 from Tillembeya is a variety with a reddish pygidium.
Oxybelus pallidus Arnold
1927, Ann. Transv. Mus. 12: tor.
Ufdem, Ethiopia, August 1945, 2 dd.
PART II, POMPILIDAE
This interesting collection of Pompilidae made by Mr. K. M. Guichard and now the ©
property of the British Museum (Natural History) comprises 162 specimens of which —
a considerable proportion is too defective, lacking antennae or tarsi or wings, to
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 167
permit of identification ; but it has, nevertheless, been possible to work out the great
bulk of it quite satisfactorily.
Subfamily PEPSINAE
Genus Hemipepsis Dahlbom
1843, Hymen, Eur. 1: 123.
Arnold, 1932, Ann. Transv. Mus. 14: 318-319.
Arnold, 1948, Occ. Pap. Nat. Mus. S. Rhod. 14: 233.
The collection contains twelve specimens of which seven are males. These do not
agree with any of the species I have seen, and in view of our present knowledge of the
genus and also of the absence of the corresponding females, it would be unwise to
describe them as new. I have elsewhere (1932) expressed the opinion that the genus
is of recent origin on account of the uniformity of the male genitalia. It is therefore
difficult to correlate the sexes unless they are taken in copula. The copulatory act
is probably of very short duration, as in the Sphecidae, for in nearly forty years of
collecting in S. Rhodesia I have only once taken a Pompilid species copulating,
Anoplius fuscus montivagus Arnold. Both sexes may occasionally be seen on trees of
which the flowers have short corollas, such as Rhus and Zizyphus, but as the repro-
ductive phase in both sexes is then past, they pay no attention to one another, so that
their proximity is of little use in correlating the sexes.
Hemipepsis aethiops Kohl
1913, Rev. Zool. Afr. 8: 193.
Didessa River, western Ethiopia, 25 May 1946, 1 9. The type was taken at Mpika,
N. Rhodesia, not Belgian Congo as stated in Kohl’s work.
Hemipepsis fallax Saussure
1892, in Distant: Nat. Tvansvaal: 221.
Didessa River and Lekempti, Ethiopia, May 1946, 2 gd.
Hemipepsis iodoptera meridionalis Arnold
Arnold, 1932, Ann. Transv. Mus. 14: 361.
Kpeve, Gold Coast, June 1942, I ¢.
A variety of this race in which the colour of the head and pronotum is castaneous
red.
Hemipepsis glabrata anchietae Radoszkovsky
Radoszkovsky, 1881, J. Sci. Mat. Phys. Nat. Lisboa, 8: 213.
Meisso; Ethiopia, 4 September 1945, I 3.
Cyphononyx flavicornis Fabricius var. antennatus Smith
1781, Spec. Insect, 1: 450.
Smith, 1855, Cat. Hymen. B.M. 3: 186.
Lake Bishoftu, 7,000 ft., May 1946, 2 92; Asba Tafari, Ethiopia, 7,800 ft., Sep-
tember 1945, I .
ENTOM. 2. 3 U
4
168 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Cyphononyx optimus Smith
1855, Cat. Hymen. B.M. 8: 141.
Ghedo, 7,000 ft., May 1946, 1g; Goré, Ethiopia, 6,000 ft., February 1948, 1 g.
Cryptocheilus impressiceps Arnold
1932, Ann. Transv. Mus. 14: 376.
Gorgora, Lake Tana, 11 December 1945, I 9.
A slight variety in which the red parts of the head and thorax are much darker
than in the type of the species. It should be noted that in this species the width of
the vertex is slightly variable. In the type it is equal to the length of the second joint
of the flagellum which is five times longer than wide at the apex ; in the paratopotype
(in the Nat. Mus., S. Rhod.) the interocular distance is fully one-fourth greater than
the length of the second joint, and in specimens from Matetsi, S. Rhodesia (in the
Nat. Mus., S. Rhod.), it is nearly one-sixth greater, or equal to the length of the first
two joints of the flagellum.
Cryptocheilus rhodesianus senegalensis subsp. n.
C. rhodesianus Arnold, 1932, Ann. Tvansv. Mus. 14: 372.
2. 12-5 mm. long. This form differs from the type of the species as follows: wings
less deeply clouded, the fuscous cloud on the basal vein absent, the distal cloud covers
only the lower two-thirds of the second cubital cell and the middle of the upper half
of the second discoidal cell. The flagellum is thinner, the second joint being nearly
five times longer than wide at the apex. The mandibles and clypeus are entirely
black. Interocular distance on the vertex equal to the length of the second joint of
the flagellum plus half of the first. Each of the scales on the hind tibiae overhang
only one spine, not two as in the type.
Diafarabé, French Sudan, 14 August 1944, type; Tillembeya, River Niger, French
Sudan, August 1944, I 9.
Priocnemis modestus sp. n.
(Fic. 54)
2. 9:5 mm. long. Black. Scapes, second joint of the flagellum, and base of the third
joint ferruginous. Legs, excepting the coxae and trochanters, pale ferruginous.
Wings smoky, with a fuscous cloud over the apex of the medial cell, and another
fuscous cloud covering the radial, second, and third cubital cells, excepting their
upper margin, and the outer upper half of the second discoidal cell. Last two joints
of the tarsi brown. Mandibles with a few long yellowish hairs. Basal half of the sixth
tergite with a dense brown pilosity. Apical margin of the clypeus slightly reddish,
not quite dull, and with a transverse row of large punctures. The rest of the clypeus,
the face, vertex, promesonotum, and mesopleura dull, reticulate-punctate, the face
and scutellum more strongly punctured (30 diameters) than the other parts. Upper
half of the metapleura strongly and obliquely striate, the lower half very finely and
closely striato-punctate. Epinotum dull, closely transversely rugulose, the rugae
stronger and farther apart on the declivity, which is also somewhat reticulate.
Abdomen shining, with a microscopic brown pubescence. Clypeus two and two-thirds
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 169
wider than long, the apical margin feebly concave. Inner orbits slightly divergent
below. Interocular distance on the vertex equal to the length of the second joint of
the flagellum. Posterior ocelli two and a half times farther from the eyes than from
each other. Second joint of the flagellum nearly five and a half times longer than wide
Fic. 54. Priocnemis modestus 9, wings, Xc.7. Fics. 55 and 55a. Marimba guichardi 2, pronotum,
dorsal and lateral view, X12. Fic. 55b. Marimba guichardi 9, wings, x7. Fic. 56. Pseudagenia
accraensis 2, head, X12. Fic. 56a. Pseudagenia accraensis 9, first tergite, x 12.
at the apex and fully one-third longer than the third joint. Temples nearly as wide
as the eyes. Postnotum with a median groove, as long as the metanotum. The
junction of the epinotal dorsum with the declivity not clearly delimited, the two
forming in profile a low and unbroken arc, the whole segment one-seventh longer than
wide. First tergite a little longer than wide at the hind margin, four and a half times
wider there than at the base. Second abscissa of the radius four times longer than the
first and one-third longer than the third. The first recurrent vein meets the cubitus
a little before the middle of the second cubital cell, the second meets the third cubital
cell at the end of its proximal third. Nervulus distinctly postfurcal. Cubitus of the
hindwing emitted a little beyond the end of the submedial cell. Hind tibiae with a
row of nine scales each overlapping a spine, and with a row of spines on each side of
the row of scales. Claws with an acute tooth in the middle.
Dilla, Ethiopia, April 1948, 1 9.
170 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Not closely related to any other Ethiopian species. The colour resembles that of
P. inermis Arnold, which is a larger species and with much stronger sculpture.
Subfamily CTENOCERATINAE
Genus Marimba Pate
Pate, 1946, Trans. Amer. Ent. Soc. 72: 93 and 120.
Turner (Cryptosalius), 1918, Ann. Mag. Nat. Hist. (9) 1: 354-355.
Arnold (Pavapompilus), 1932, Ann. Transv. Mus. 15: 89, 104-106.
Marimba guichardi sp. n.
(Fics. 55, 55@ and 550)
Q. 11-5 mm. long. Black, head and prothorax very dark reddish-brown. First
six joints of the antennae, legs, and extreme apex of sixth tergite, fusco-ferruginous.
Wings hyaline, bifasciated, the inner fascia covering the apical two-fifths of the
medial, first submedial and anal cells and the base of the second submedial and first
discoidal cells, the outer fascia covering the radial cell excepting the apex, the second
and third cubital, the apex of the first cubital cell and first discoidal cell, and the
second discoidal cell excepting the base. Flagellum, mesosternum, and coxae covered
with a decumbent, very fine and silvery pubescence. Lower corners of the epinotum
with exserted silvery pubescence, the apical margins of the first three tergites with
decumbent silvery pubescence at the sides. Labrum semicircular, the galea extending
beyond the closed mandibles, the latter with a tooth behind the apex. Clypeus
microscopically and closely punctured, with a greasy lustre, nearly three times wider
than long, the anterior margin straight. Face finely and very closely punctured
(25 diameters), the interspaces as wide as the punctures. Inner orbits parallel. On
each side of the bottom of the face there is a deep circular pit, half on the face and
half on the clypeus. Inter-antennal tubercle prominent; a narrow impressed line
extends from it to the anterior ocellus. Posterior ocelli very little farther from each
other than from the eyes. Interocular distance on the vertex equal to the length of
the second joint of the flagellum plus two-ninths of the third joint ; the second joint
is slightly more than three times longer than wide at the apex and the third joint is
one-sixth longer than the second. The puncturation of the dull promesonotum and
scutellum is like that of the face but a little larger. The dorsum of the pronotum
merges gradually into the anterior face, the sides are nitidulous, with a few large
punctures, and the transverse groove is wide. Mesonotum three times longer than the
pronotum. Scutellum very convex, subcarinate lengthwise in the middle.
Postnotum much shorter than the metanotum and depressed. Mesopleura coarsely
rugose, pitted between the rugae. Metapleura obliquely costate. Dorsum of the
epinotum two-thirds wider than long, shining, very coarsely and transversely rugose,
with numerous anastomoses or almost reticulate-rugose, the interspaces wide, about
as wide as the ocelli, the sides more closely rugose, the declivity oblique, shining, and
with seven transverse rugae. Abdomen lanceolate, two and two-thirds longer than
wide across the middle, the first tergite as long as wide behind and four times wider
there than at the base. Sixth tergite longer than wide at the base, the apex subacute.
Transverse groove of the second sternite between the first and second third of the
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 171
segment. Second abscissa of the radius a trifle longer than the third and four times
longer than the first. Cubitus of the hindwing emitted a little before the end of the
submedial cell. Claws bifid.
Dilla, Ethiopia, April 1948, 1 9.
Apparently closely related to M.indocilis Turner, which I have not seen, but differ-
ing therefrom according to Turner’s description as follows: the smaller size, the colour
of the legs and basal joints of the antennae, the shorter second abscissa of the radius,
and the close puncturation of the promesonotum. From M. contristans Turner it
differs also by the puncturation of the pro- and mesothorax, the smaller size, and the
colour of the wings. These two species and two others from Africa were described by
Turner (1918) and placed in his genus Cryptosalius (genotype C. vava Bingham). In
my monograph of the Ethiopian Pompilidae I placed Cryptosalius in synonymy with
Parapompilus. However, Pate (1946) indicated that Turner’s African species were
not congeneric with C. vava and proposed for them the new generic name Marimba.
These species and the one described above differ from Parapompilus in the absence
of a tarsal comb, the fasciated wings, and the long and very coarsely sculptured
epinotum. Another distinction mentioned by Pate, ‘the humeral angles of the pro-
notum not broadly rounded as in Parvapompilus’, does not hold good, as in this
species, guichardi, they are rounded. Until the males are known, the relationship of
Marimba with Parapompilus cannot be decided.
Subfamily PSEUDAGENIINAE
Pseudagenia pseudocyphononyx Turner
1918, Ann. Mag. Nat. Hist. (9) 1: 352.
Adis Ababa, 3 August 1945; Zuquala, Ethiopia, October 1945, 2 99.
Somewhat smaller and with shorter wings than in the type and specimens from
S. Rhodesia.
Pseudagenia accraensis sp. n.
(Fics. 56, 56a)
2. 16mm. long. Mesopleura, metanotum, metapleura, epinotum, and the first
three abdominal segments black, the rest of the body, the antennae, and legs pale
ferruginous. Wings very pale yellowish-brown, slightly darker over the radial,
second cubital, and the upper half of the second discoidal cells, but not fasciated.
Face and clypeus with a microscopic reddish pubescence. Mesopleura closely and
very finely reticulate rugose, metapleura closely striate, epinotum closely, trans-
versely, and finely rugose, the rugae strongest on the posterior half of the epinotal
dorsum. Clypeus very convex transversely, nearly half as wide again as long in the
middle, the anterior margin convex, with a slight projection in the middle. Inter-
ocular distance across the base of the clypeus one-third greater than across the vertex,
where it is equal to three-fourths of the length of the second joint of the flagellum.
Posterior ocelli as far from the eyes as from each other. Flagellum long and slender,
all the joints excepting the first at least six times longer than wide, the second joint
nearly seven times longer than wide at the apex and one-seventh longer than the third.
172 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Scutellum fairly flat, two-thirds as long as wide at the base. Postnotum shining,
finely and transversely striate, as long as the metanotum. The whole epinotum is as
long as wide at the base, the dorsum merges gradually into the declivity, the whole
forming a low curve in profile. First tergite one-fifth longer than wide, very narrow
basad. Second abscissa of the radius three times longer than the first, and half as
long again as the second. The wings extend back to the apex of the abdomen.
Accra, I94I, I 9.
The species nearest to this one is P. lwjae Arnold, from which it differs by the
longer clypeus, the narrower vertex, the much more slender antennae, the longer
peduncle of the first tergite, and the colour of the wings.
Pseudagenia stigmalis R. Lucas
1898, Deutsch Ost-A fr. (3) 4: 61.
Arnold, 1934, Ann. Transv. Mus. 15: 342, 346.
Adis Ababa, 3 August 1945, 2 99.
It is with some hesitation that I ascribe these two specimens to this species, since
the original description is quite inadequate. They appear to differ only in having the
hind femora entirely ferruginous, and the face also of that colour excepting a black
streak adjacent to the antennal sockets.
Pseudagenia laevigata Smith
1855, Cat. Hymen. B.M. 8: 143.
Accra, Gold Coast, June 1941, 2 99.
Pseudagenia flavotegulata Bingham
1902, Ann. Mag. Nat. Hist. (7) 9: 208.
Kaolack, Senegal, August 1943, I 9.
A slight variety in which the base of the hind coxae on the upper side, the first
abdominal segment and the sides and pre-apical bands on the second and third
tergites, and the middle of the second and third sternites are black.
Pseudagenia infantula Kohl
1894, Ann. naturh. Hofmus. Wien, 9: 307.
Lake Bishoftu, Ethiopia, July 1945, I 9.
This specimen represents a slight variety in which the vertex is a little narrower,
being equal to the length of the second joint of the flagellum, and the first tergite as
long as wide behind. The distinctive shape of the clypeus is exactly like that of
infantula i. sp.
Pseudagenia albosignata sp. n.
(Fics. 57, 57a and 575)
g. 8mm. long. Black. Clypeus, excepting the basal half of the middle, and the
sides of the face to a little beyond the level of the antennal sockets, dirty white. Legs
ferruginous, the coxae, trochanters, and tarsi brown. Wings hyaline, with a pale
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 173
fuscous cloud over the basal vein and nervulus, and an outer fuscous cloud covering
the basal third of the radial cell, the greater part of the second cubital cell and a small
area on the upper part of the second discoidal cell; the apex of the forewing also
slightly fuscous. Clypeus and face with a sparse brownish pilosity, the temples,
coxae, posterior half of the mesosternum and posterior angles of the epinotum with
a decumbent silvery pubescence, the latter and the mesosternum and metapleura
. : “60
Fic. 57. Pseudagenia albosignata 9, head, X15. Fic. 57a. Pscudagenia albosignata 3, head, X15.
Fic. 57b. Pseudagenia albosignata 3, genitalia, xc. 45. Fic. 58. Poecilagenia longicollis 2, thorax,
X15. Fic. 59. Poecilagenia rugosa g, pronotum, X12. Fic. 59a. Poecilagenia rugosa 6,
epinotum and basal tergites, x12. Fic. 60. Paracyphononyx funebris 3, genitalia, x 30.
also with some longer, exserted, and white hairs. Postnotum shining, arcuately and
transversely striate, depressed in the middle and as long as the metanotum. The rest
of the thorax and the head dull, very finely reticulate-punctate (30 diameters), the
epinotum also transversely rugulose, feebly so over the basal third. Tergites with a
microscopic greyish pubescence.
Clypeus very convex, nearly half as wide again as long, the anterior margin convex.
Inner orbits slightly convergent below, the interocular distance on the vertex equal
to a trifle more than the length of the first two joints of the flagellum. Posterior
ocelli twice as far from the eyes as from each other. Second joint of the flagellum five
and a half times longer than wide and barely longer than the third.
Dorsum of the pronotum half as long as the mesonotum, the anterior face
174 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
sub-vertical, the shoulders rounded. Scutellum very convex. Dorsum and declivity of
the epinotum not distinctly delimited, the two parts forming a low arc in profile; the
dorsal face with a shallow median longitudinal groove, the whole segment as long as
wide. First tergite nearly twice as long as wide behind, and a little more than twice
as wide there as at the base, the sides straight. Second abscissa of the radius two-
thirds longer than the first, the third two-fifths longer than the second ; the nervulus
almost interstitial with the basal vein.
2. 9mm. long. Face and clypeus entirely black ; antennae fusco-ferruginous. Sixth
tergite broadly triangular, flat, shining, and sparsely punctured. Otherwise like the g
in colour, sculpture, and pubescence. Clypeus two-fifths wider than long, the anterior
margin produced into a blunt triangular lobe. Face as wide below as on the vertex,
where the interocular distance is equal to a very little less than the length of the first
two joints of the flagellum. Posterior ocelli not quite twice as far from the eyes as
from each other. Flagellum slender, the second joint five times longer than wide at
the apex. First tergite wider caudad than in the g, only three-sevenths longer than
wide behind. Otherwise like the 2.
Adis Ababa, August-September 1945, 2 gg, I 9; Lekempti, Ethiopia, May
1946, 1 3.
Related to P. rubrirostris Arnold, from which it differs by the colour of the legs, the
paler fasciae of the wings, and the more acute lobe of the clypeus. It also resembles
P. amita Arnold, but is smaller, differently coloured, and with a less prominent lobe
on the clypeus.
Poecilagenia longicollis sp. n.
(Fic. 58)
9. 8:5 mm. long. Head, pro-mesonotum, scutellum, metanotum, anterior half of
the meso- and metapleura, and anterior legs, pale ferruginous. Posterior half of the
mesopleura, the metapleura, epinotum, and abdomen, black. Middle and hind tibiae
fusco-ferruginous. Antennae pale ferruginous at the base, becoming gradually darker
to fuscous from the fifth joint onwards. Wings hyaline, with a pale fuscous
cloud covering the basal third of the radial cell, distal half of the second cubital,
proximal half of the third cubital cell, and the distal upper corner of the second
discoidal cell. Mesopleura and posterior corner of the epinotum with decumbent
silvery pubescence, the tergites with sparse, very fine and grey pubescence. Clypeus
slightly shining, very shallowly punctured. Face dull, closely and finely reticulate-
punctate, the vertex nitidulous and with a few punctures. Pronotum not quite
dull, closely and transversely striate. Mesonotum transversely, the scutellum longi-
tudinally and closely rugulose and nearly dull. Mesopleura obliquely, metapleura
transversely rugose. Epinotum dull, coarsely reticulate-rugose, and with a fine
median longitudinal carina. Abdomen shining and impunctate. Mandibles with a
small tooth just behind the apex. Maxillary palpi long, the last three joints united
fully as long as the first three joints of the flagellum. Clypeus nearly two and a third
times wider than long, the apical margin convex. The interocular distance on the
vertex is very little less than it is across the bottom of the eyes and is equal to the
length of the first two joints of the flagellum. Posterior ocelli a little farther from
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 175
the eyes than from each other. First joint of the flagellum half as long again as wide,
the second joint three and three-fifths longer than wide at the apex, four-fifths longer
than the third which is twice as long as wide. Dorsum of the pronotum widest across
the middle, twice as wide there as long, the anterior margin straight, the shoulders
subrectangular. Mesonotum three-eighths longer than the pronotum. Scutellum and
disk of the metanotum feebly convex. Postnotum shallow, in the middle one-third
as long as the metanotum. Dorsum of the epinotum parallel-sided, about one-ninth
wider at the base than long, the declivity about half as long as the dorsum and oblique.
First tergite about one-fourth longer than wide at the posterior margin, and one-third
longer than the second tergite.
Second abscissa of the radius twice as long as the first, a little longer than the third
which is half as long as the fourth. Transverse anal vein of the hindwing prefurcal.
Aburi, Gold Coast, December 1941, I 9.
Differs from P. reversa Bischoff by the longer pronotum and epinotum, narrower
vertex, and paler red ‘colour.
Poecilagenia rugosa sp. n.
(Fics. 59, 594)
3g. 8mm. long. Head, pro- and meso-sternum, the middle third lengthwise of the
epinotum, abdomen, coxae, trochanters, and the hind legs black. The thorax above
and at the sides dark red. Apex of the mandibles ferruginous, the acute inferior edge
of the scapes yellowish, the apical tergite pale yellow. Fore and middle legs, excepting
the coxae and trochanters, fusco-ferruginous. Wings hyaline, with a pale brown cloud
covering the greater part of the radial, second, and third cubital cells and the distal
corner of the second discoidal cell. Clypeus and face with a short, coarse, and yellow-
ish pubescence, the temples with long greyish pilosity. Dorsum of the thorax except-
ing the base of the epinotum and the pronotum nearly glabrous, the rest of the thorax
and the coxae and trochanters with a coarse silvery pubescence intermixed with a
long pilosity of the same colour, sufficiently dense to obscure the sculpture on the
pronotum and epinotum. Abdomen with a decumbent, very fine and grey pubescence.
Clypeus and face dull, strongly and closely punctured, the temples, vertex, and
occiput more sparsely so and not quite dull. Pronotum and mesopleura coarsely
reticulate-rugose. Mesonotum dull, reticulate-punctate, the rugae thicker trans-
_ versely, the scutellum and metanotum with a similar but larger sculpture. Postnotum
half as long as the metanotal disk, with two or three costae on each side of the median
depression. Metapleura obliquely costate. Epinotum very coarsely reticulate-rugose,
the majority of the interspaces fully as wide as the ocelli. Hind coxae coarsely rugoso-
punctate and swollen at the base on the upper side.
Clypeus a little more than twice as wide as long, the anterior margin feebly convex.
Antennal sockets twice as far from each other as from the eyes, in consequence of
which the supra-antennal tubercle is wide. Interocular distance on the vertex equal
to the length of the first four joints of the flagellum and one-fifth greater than across
the base of the clypeus. Ocellar area tumid, the posterior ocelli as far from the eyes
as from each other. Temples narrow. Scapes transversely compressed, with a sharp
edge on the underside. First joint of the flagellum a little longer than wide, the
ENTOM. 2. 3 x
176 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
second not quite twice as long as wide and as long as the third. Dorsum of the pro-
notum fairly convex transversely, two and a half times wider in the middle than long,
the anterior margin nearly straight, the anterior face vertical, the shoulders sub-
rectangular. Mesonotum twice as long as the pronotum. Scutellum convex trans-
versely, as long as wide at the base. The dorsum and the very oblique declivity of the
epinotum form a very low arc in profile; the whole segment is one-seventh wider at
the base than long, and the sides are feebly convex. Abdomen clavate, the first
tergite long and narrow, transversely compressed, the sides vertical and flat, three
times longer than wide at the apical margin which is less than twice as wide as the
base. The rest of the abdominal complex is narrow and five-sixths longer than the
first tergite. The length of the first three abscissae of the radius are as 5:12:10.
Cubitus of the hindwing interstitial with the transverse anal vein.
Diafarabé, French Sudan, 5 August 1944, I 3.
This specimen is badly damaged, the head and pro-mesonotum being separated
from the thorax, and parts of the legs are missing. However, objectionable as it is
to base a new species on such specimens, I have done so in this case, as the species is
so distinct from the other known males of the genus that it can be recognized without
difficulty.
Cyemagenia rubrozonata Cameron
1910, Sjdstedt’s Kilimandjaro-Meru Exped. 8: 253.
Dilla and Haramaia, Ethiopia, April 1948, I 2, 4.4¢.
Subfamily POMPILINAE
Atopopompilus venans mlanjiensis Arnold
1937, Ann. Transv. Mus. 19: 24.
Accra, Gold Coast, 13 July 1941, I 9.
This specimen is an aberration in which the second and third abscissae of the radius
are of about equal length. Otherwise it agrees in all respects with the type of the
subspecies.
Paracyphononyx affinis Haupt
1929, Mitt. Zool. Mus. Berl. 15: 170.
Aburi, Gold Coast, December 1941, I 3.
Paracyphononyx difficilis Bischoff
1913, Arch. Naturgesch. 79: 55.
Tillembeya, River Niger, French Sudan, 14 August 1944, 1g; Accra, May 1941, I ¢.
Paracyphononyx laboriosus Arnold
1936, Ann. Transv. Mus. 18: 438.
Asba Tafari, Ethiopia, 7,800 ft., September 1945, I 9.
Paracyphononyx metemmensis Magretti
1884, Ann. Mus. Stor. Nat. Genova, 21: 565.
Tillembeya, River Niger, French Sudan, August 1944, 1 $; Accra, Gold Coast,
December 1941, I ¢.
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 177
Paracyphononyx funebris sp. n.
(Fic. 60)
g. 7°5-9°5 mm. long. Black. Flagellum ferruginous below, brownish-black above,
scape dirty yellow on the underside. Wings fuscous. Clypeus, face, and underside
of the coxae with greyish-silvery pubescence, the head and pronotal collar also with
black, erect hairs. Epinotal declivity and base of the first tergite with a dense, coarse,
erect whitish pubescence, the extreme base of the second, third, and fourth tergites
with a decumbent bluish-grey bloom. Sixth tergite white.
Clypeus nearly two and a half times wider than long, separated from the bottom of
the eyes by a length equal to two-thirds of the cheeks. Interocular distance on the
vertex very slightly less than the interocular distance across the base of the clypeus,
and equal to the length of the first two joints of the flagellum plus two-thirds of the
third joint. The fourth to eleventh joints of the flagellum are dilated below near the
base, and the second joint is a little shorter than the third. Posterior ocelli two-thirds
farther from each other than from the eyes. Pronotum two and a half times wider
across the posterior angles than long. Dorsum of the epinotum twice as wide at the
base as long, twice as long as the declivity, the junction of the two faces rounded.
First tergite as long as wide behind. First abscissa of the radius a little longer than
the second ; third cubital cell petiolate, the petiole variable in length, but sometimes
as long as the second abscissa of the radius.
' Haramaia, Ethiopia, May 1948, on Myosotis flowers, 19 3d.
Related to difficilis Bischoff, from which it differs by the colour of the legs, the
darker wings, wider clypeus, and the genitalia. In the latter the outer paramera are
not spatulate as in difficilis, but narrow and strap-like, with six long hairs emitted
from about the middle of the inner margin.
Paracyphononyx zonatus Illiger
1802, Mag. Insektenkunde, 1: 193.
Diafarabé, French Sudan, 14 August 1944, I 3.
A slight variety with the outer paramera of the genitalia narrower in the middle.
In addition to the species listed above, the collection contains seven females,
representing two or three species of the genus Paracyphononyx, which do not agree
with any of the species known to me. It would be unwise to describe these as new in
the absence of specimens which could be indubitably considered the opposite sex.
In this genus the males can easily be determined by the genitalia, but the females
are far more difficult to determine.
Episyron histrio Lepeletier
1845, Hist. Nat. Ins. Hymén, 3: 440.
Tillembeya, River Niger, French Sudan, 14 August 1944, 1 9; Abbai Gorge and
River Didessa, Ethiopia, 23 October 1945, 25 August 1946, I 9, I d.
The specimen from the Didessa, 11 mm. long, is not only larger than the typical of
the species but differs considerably in colour, the first four abdominal segments being
178 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
black, excepting the yellow bands on the second and third tergites. The genitalia,
however, are identical.
Batazonellus capensis Dahlbom
1843, Hymen. Eur. 1: 49.
Batazonellus Arnold, 1937, Ann. Transv. Mus. 19: 1.
Lake Bishoftu, Ethiopia, 12 May 1946, I 9.
Schistonyx umbrosus Klug
1834, Symb. phys., Dec. 4, T. 39, F. 4.
Dakar, Senegal, 1 August 1945, 2 99.
Elaphrosyron insidiosus Smith
1879, Descr. new sp. Hymen.: 143.
Kaolack, Senegal, 1 9; Diafarabé, French Sudan, 1 9; Dilla, Ethiopia, 2 g¢.
GUICHARDIA gen. n.
Characters
3. Palpi short, the last three joints of the maxillary palpi not more than three
times longer than wide. Labrum exposed. Mandibles with a tooth behind the apex.
Clypeus feebly convex, as wide as the bottom of the face. Cheeks very short. Face
without a supra-antennal tubercle. Occiput not concave, its junction with the vertex
rounded. Pronotum not much shorter than the mesonotum, convex lengthwise.
Lateral margins of the mesonotum feebly reflexed posteriorly. Sides of the scutellum
steep, raised above the level of the metanotum. Postnotum linear. Dorsum of the
epinotum convex lengthwise and transversely, forming in profile a low and unbroken
arc with the very short declivity, its posterior angles dentiform. Apical ventral valve
produced into a long spine. Femora of the trachyscelid type. Claws bifid, the pul-
villus small and with about ten short cilia. Wings with three cubital cells, the third
much wider on the cubitus than on the radius; nervulus slightly prefurcal. Cubitus
of the hindwing emitted a little beyond the end of the submedial cell, the basal lobe
small.
Genotype: G. macilenta Arnold.
The subdentate posterior angles of the epinotal dorsum are very similar to those of
Epiclinotus Haupt, but the wide clypeus and the rounded junction of the slightly
convex occiput with the vertex show that the genus belongs to the tribe Pompilini.
Guichardia macilenta sp. n.
(Fics. 61, 61a—d)
g$. 10°5-II mm. long. Black. Head and thorax with a fine, decumbent grey
pubescence and without erect pilosity. First three abdominal segments (or four in
the paratype) with a dense, long, greyish-silvery pubescence, the remaining segments
with a thin black pubescence and some longer and erect black hairs. Wings almost
hyaline over the basal fourth, the rest moderately fuscous with traces of darker
clouds over the basal vein area and the radial and second cubital cells. Clypeus
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 179
two-thirds wider than long, the lateral margins long, the anterior margin straight.
Inner orbits parallel below, slightly divergent above the middle and convergent on
the vertex, the interocular distance there no greater than across the base of the
clypeus, and equal to the length of the second and third joints of the flagellum united.
Posterior ocelli slightly nearer to the eyes than to each other. Second joint of the
flagellum as long as the third, and nearly two and a half times longer than wide.
Fic. 61. Guichardia macilenta 3, head, X12. Fic. 61a. Guichardia macilenia, head and pro-
notum, x12. Fic. 61b. Guichardia macilenta, epinotum and first tergite, x12. Fic. 61c.
Guichardia macilenta, epinotum, x12. Fic. 61d. Guichardia macilenta, genitalia, x27. Fic. 62.
Anoplius octomaculatus 9, epinotum, x6. Fic. 62a. Anoplius octomaculatus, abdomen, x 6.
Pronotum convex lengthwise, seven-ninths wider across the posterior angles than long
in the middle, moderately narrowed cephalad, the hind margin obtusely angular.
The horizontal part of the scutellum one-fifth longer than wide, and about three
times longer than the metanotal disk. Epinotum moderately convex lengthwise and
transversely, about one-eighth longer than wide at the base, the lateral margins
convex ; the dorsum is about four times longer than the very oblique declivity, its
posterior angles subdentiform. Abdomen narrow, the first tergite about one-seventh
longer than wide at the apical margin. First and third abscissae of the radius of
equal length, and half as long as the second. The third cubital cell three and two-
thirds wider on the cubitus than on the radius; both recurrent veins meet the second
and third cubital cells a little beyond the middle.
Agogo, Gold Coast (type), 30 January 1942; Aburi, Gold Coast, 21 January 1941.
180 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Anoplius octomaculatus sp. n.
(Fics. 62 and 62a)
9. 15mm. long. Black. First four tergites with large yellowish-red maculae on
each side. Wings dark fuscous. Clypeus with a sparse grey pubescence, the face with
a small patch of silvery pubescence on each side, adjacent to the eyes and at about
the level of the antennal sockets, the rest of the face and the temples with a very
sparse black pilosity. The posterior corners of the mesopleura, the inner half of the
upperside of the middle and hind coxae with dense and brilliant silvery pubescence.
The setae on the apical abdominal segment long and black. Clypeus two and a
quarter times wider than long, the apical margin feebly convex. The inferior inter-
ocular distance a little greater than across the vertex, where it is equal to slightly less
than the length of the second joint of the flagellum. Posterior ocelli one-third farther
from the eyes than from each other. Antennae slender, the second joint of the
flagellum six times longer than wide at the apex and one-third longer than the third
joint. Temples half as wide as the eyes. Pronotum three and a half times wider
between the posterior angles than long in the middle. Scutellum three-fourths as long
as the mesonotum. Disk of the metanotum flat, truncate behind. Postnotum de-
pressed, half as long as the metanotal disk, shining and transversely striate. Dorsum
of the epinotum more than twice as wide at the base as long, with a median longi-
tudinal impressed line, the declivity steep, compressed at the sides and not much
shorter than the dorsum. Second abscissa of the radius five-eighths longer than the
first, the third cubital cell petiolate, the petiole very short, or about one-fifth the
length of the first abscissa of the radius. Both recurrent veins meet the cubital cells
beyond their middle. Transverse anal vein of the hindwing postfurcal.
Tillembeya, River Niger, French Sudan, 26 September 1944, I 9.
Not closely related to any other Ethiopian species, and easily recognized by the
silvery pubescence on the mesopleura and coxae and the large red maculae on the
tergites.
Anoplius successor Cameron
1910, Sjéstedt’s Kilimandjaro-Meru Exped. 8: 251.
A. subfasciatus Arnold, 1937, Ann. Transv. Mus. 19: 63.
River Didessa, May 1946, and Debra Sina, Ethiopia, September 1945, 2 99. The
specimen from Debra Sina is a slight variety in which the wings are darker and the
pubescence entirely black.
Anoplius morosus Smith
1855, Cat. Hymen. B.M. 8: 140.
Akaki and Asba Tafari, Ethiopia, February 1948, I 9, 2 gd.
Anoplius fuscus L.
1761, Fauna Suec, ed. 2: 412.
Arnold, 1937, Ann. Transv. Mus. 19: 65.
In my monograph of the Ethiopian Pompilidae I remarked on the variability of
this species in the Ethiopian region and named some varieties. There are four speci-
mens in the collection, from different parts of Ethiopia, of which one agrees with the
Sern ioe pdeek see
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 181
var. acutiangulus Cameron, and the others differ only in a minor degree from the
varieties montivagus, johannis, and brachcerus Arnold. It would serve no useful
purpose to give names to these specimens since the variations grade one into another.
Pompilus plumbeus Fabricius
1787, Mant. Ins. 1: 278.
Labadi, Gold Coast ; Fatick, Senegal ; Tillembeya, River Niger, French Sudan, and
River Hawash, Ethiopia, 1 3, 4 29.
The specimens from Fatick and Hawash represent a slight variety in which the
first four joints of the anterior tarsi and the second to fourth of the middle and hind
tarsi are yellowish-red.
Pompilus rutilus Klug subsp. lutarius Saussure
1834, Symb. phys. 4: T. 38.
Saussure, 1891, Mitt. Schweiz. Ent. Ges. 81: 265.
Accra and Labadi, Gold Coast ; Tillembeya, River Niger, French Sudan ; Diafarabé,
French Sudan, 3 99, 8 g¢.
Pompilus cinnamomeus Arnold
1937, Ann. Transv. Mus. 19: 52.
Accra, Gold Coast, September 1941, 6 99.
Pompilus contrarius sp. n.
(Fic. 63)
Q. 8-5 mm. long. Head and thorax black, the anterior half of the clypeus and the
neck and lateral margins of the pronotum, flavo-ferruginous. Mandibles ferru-
ginous, the apex black. Abdomen and legs pale ferruginous, antennae fusco-ferru-
ginous. Wings hyaline, tinged with yellow; the forewing beyond the cells, but not
including the extreme apical margin, faintly fuscous. Head and thorax with a dull
white decumbent pubescence, long and dense on the epinotum where it is arranged
in close transverse rows directed laterad on each side of the middle line; below the
pubescence there are traces of transverse striae. Clypeus two and a half times wider
than long, the apical margin shallowly concave. Inner orbits parallel, slightly con-
vergent on the vertex. Interocular distance at the base of the eyes one-fifth greater
than on the vertex, where it is equal to the length of the first two joints of the
flagellum plus one-sixth of the third joint. Posterior ocelli three-fifths farther from
each other than from the eyes. Flagellum slender, the second joint nearly five and
a half times longer than wide at the apex. Pronotum about two and a third times
wider than long (measured tangentially), the hind margin arcuate. Postnotum one-
fourth as long in the middle as the metanotal disk, transversely striate. Epinotum
with a fine median longitudinal carina, the dorsum gibbous, merging gradually into
the steep declivity which is about twice as long as the dorsum, and flat. The dorsum
is three times wider than long. The proportions of the first three abscissae of the
radius are as 3:11:8; the third cubital cell is twice as long on the cubitus as on the
radius. Transverse anal vein of the hindwing prefurcal.
182 SPHECIDAE AND POMPILIDAE (HYMENOPTERA) COLLECTED
Tillembeya, River Niger, French Sudan, 14 August 1944, I 9.
Easily recognized by the peculiar arrangement of the pubescence of the epinotum
and its flattened and wide declivity.
64
Fic. 63. Pompilus contrarius 9, epinotum, xc. 15. Fic. 64. P. (Pilompus) levis 9, head, Xc. 15.
Fic. 64a. P. (Pilompus) levis, epinotum, xc. 15. Fic. 646. P. (Pilompus) levis, epinotum and
first tergite, xc. 15. Fic. 64c. P. (Pilompus) levis, wing, X12. Fic. 65. Ceropales variolosus 6,
left stipes, dorsal view, x 27.
PILOMPUS subgen. n. (anagram Pompilus)
Characters
9. Anterior tarsi without a comb, all the claws bifid, the pulvillus small witha —
comb of about nine cilia. Clypeus not strap-like and much wider than long as in
Pompilus, but narrow, more or less hexagonal (Fig. 64), and as wide as the bottom of
the face. First tergite wide at the base, not much narrower there than at the apex.
Otherwise like Pompilus.
Subgenotype, P. levis Arnold.
Pilompus levis sp. n.
(Fics. 64, 64a-c)
9. 8:5 mm. long. Black. Palpi ochreous. Wings hyaline, the forewing beyond the ~
cells pale fuscous. Head, thorax, and legs with a very fine pruinose pubescence ; the
BY MR. K. M. GUICHARD IN W. AFRICA AND ETHIOPIA 183
basal two-thirds of the first tergite and the basal half, more or less, of the second to
fourth tergites with a similar pubescence. Clypeus two and a third times wider than
long, the anterior margin concave and twice as long as the lateral margins. Palpi
short. Inner orbits divergent over the middle third of their length. Interocular
distance on the vertex equal to the length of the first two joints of the flagellum plus
one-third of the third joint. Posterior ocelli twice as far from each other as from the
eyes. Flagellum fairly stout, the second joint barely three times longer than wide at
the apex and as long as the third joint.
Occiput nearly flat, its junction with the vertex widely rounded. Pronotum twice
as wide behind as long in the middle, the hind margin obtusely angular, the dorsal
and anterior face forming an unbroken arc in profile. Mesonotum as long as the
pronotum. Postnotum about half as long in the middle as the feebly convex meta-
notal disk, linear at the sides. Epinotal dorsum widest across the middle, seven-
ninths wider there than long, nearly twice as long as the oblique declivity which is
nearly flat transversely. First tergite as long as wide behind, only three-fourths wider
there than at the base. Anterior tarsi with a few short spines on the outside ; posterior
tibiae with two rows of three spines each on the upperside. Third cubital cell petiolate,
the petiole half as long as the second abscissa of the radius; the proportions of the
three abscissae are as 6:4:10. First transverse cubital vein strongly curved inwards,
the pterostigma very small, the nervulus postfurcal. Transverse anal vein of the
hindwing slightly prefurcal.
Aburi, Gold Coast, December 1941, I 9.
Subfamily CEROPALINAE
Ceropales variolosus Arnold
(Fic. 65)
1947, Ann. Transv. Mus. 19: 87, 9.
3. 7:5 mm. long (hitherto undescribed). Sixth and seventh abdominal segments
yellow. Puncturation of the face less close than in the 9, the punctures arranged in
transverse rows, the spaces between the rows about twice as wide as the punctures.
Clypeus two and a half times wider than long. The proportions of the first three
abscissae of the radius are as 5:11:15. The outer paramera of the genitalia have the
shape of a flattened fish-hook, with the acute, triangular, and transparent barb on
the inside.
Accra, Gold Coast, September 1941, 2 §¢, 1 9; Diafarabé, French Sudan, August
1944, I 9.
The colour is variable. In the other $ specimen and in the 2 from Diafarabé the
red parts of the head and thorax are replaced with black.
Ceropales latifasciatus Arnold
1937, Ann. Transv. Mus. 19: 92.
Lekempti, Haramaia, and Cencia, Ethiopia, April-May 1948, 3 dé.
ENTOM. 2. 3 | ef s Y
Fic, I.
Fic. 2.
Fic. 3.
Fic. 4.
Fic. 5.
Fic. 6.
BiG. -7.
Fic. 8.
Fic. 9.
PLATE 4
GENITALIA
Cerceris guichardi, x 25.
Cerceris moggionis, X25.
Cerceris insignita, X 30.
Cerceris bicolor, x 28.
Cerceris zavattarii var., X 33.
Cerceris rhodesiae, X 36.
Cerceris campsomeroides, X 30
Cerceris faceta, X 42.
Cerceris flavonasuta, X 30.
PLATE 4
Bull. B.M. (N.H.) Ent. II, 3
rm
GENITALIA
PLATE 5
GENITALIA
Fic. 10. Cerceris maia, X 42.
Fic. 11. Cerceris alcyone, X 42.
Fic. 12. Cerceris merope, X 40.
Fic. 13. Cerceris sterope, X 42.
Fic. 14. Cerceris celaeno, x 60.
Fic. 15. Cerceris taygete, x 40.
Fic. 16. Cerceris nugax insularis, x 25.
Fic. 17. Cerceris nugax, X25. ‘
Fic. 18. Cerceris yungvei, X 40.
Fic. 19. Alysson guichardi, x 60.
Fic. 20. Nysson quadricolor, x 40.
Fic. 21. Brachystegus senegalensis, X25.
Fic. 22. Gorytes thalia montivagus, x 42.
PLATE 5
Bull. B.M. (N.H.) Ent. II, 3
a =
or 0 rom mee os aay 5 RE
3,
AY EEN
GENITALIA
: xj Ss
Jay, My
rete
PRESENTED
17 DEC 1951
eee ee.
PRINTED IN
GREAT BRITAIN
AT THE
UNIVERSITY PRESS
OXFORD
CHARLES BATEY
PRINTER
TO THE
7 FEB 1992
a CO PItON OF FLEAS
Peom THE BODIES OF
BRITISH BIRDS
WITH NOTES ON THEIR DISTRIBUTION
AND HOST PREFERENCES
MIRIAM ROTHSCHILD
: BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 2 No. 4
LONDON : 1952
me WcLeCriON OF FLEAS
FROM THE BODIES OF BRITISH BIRDS
WITH NOTES ON THEIR DISTRIBUTION
AND HOST PREFERENCES
BY
MIRIAM ROTHSCHILD )
Xf
Pp. 185-232; 3 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 2 No. 4
LONDON : 1952
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, 1s issued
in five series, corresponding to the Departments of the
Museum.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four hundred
pages, and will not necessarily be completed within one
calendar year.
This paper is Vol. 2, No. 4, of the Entomology series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued February 1952 Price Ten Shillings
mACILLEC TION OF FLEAS PROM THE BODIES
Or BRITISH BIRDS, WITH NOTES ON THEIR
SrsteibUTION AND HOST PREFERENCES
By MIRIAM ROTHSCHILD
With three text-figures
SYNOPSIS
Ten species of fleas (143 specimens) were collected from the bodies of 36 species (74 specimens) of British
birds. There are single records of Ceratophyllus styx (2 specimens), C. hirundinis (10 specimens), C.
columbae (1 specimen), C. vagabunda (2 specimens), Nosopsyllus fasciatus (1 specimen), and Orchopeas
wickhami (1 specimen). There are 2 records of C. rusticus (26 specimens), 3 records of C. garei (4 speci-
mens), 32 records of C. gallinae (52 specimens), and 36 records of Dasypsyllus g. gallinulae (44 specimens).
The geographical distribution and host preferences of C. vagabunda, C. garei, C. gallinae, and D. g.
gallinulae are discussed, and in addition the ratio of fleas found on the body and in the nest of the host
and the sex-ratios of the three last-mentioned species are considered. It is shown that fleas may be
collected from the bodies of birds at all times of the year, and not only in the breeding-season.
INTRODUCTION
THE fleas discussed in this paper were obtained by Miss Theresa Clay and Colonel
Richard Meinertzhagen from the bodies of British birds and have recently been
presented to the British Museum (Natural History). The collection numbers 143
specimens, more than double the total bird-fleas taken from the bodies of their hosts
(as opposed to nests) in the combined Rothschild, Waterston, and British Museum
cabinets. It thus forms the finest collection of its kind in existence, and in an effort
to throw some light on various interesting points which it raises, especially concern-
ing the host preferences and distribution of the three commonest bird-fleas, all the
British records of these species have been reviewed and are discussed in the following
pages.
It is still a general belief that fleas occur only occasionally on the bodies of adult
avian hosts. As recently as 1949 O’Mahony writes: ‘Two species of fleas have been
collected, both from the bodies of the birds—a rare enough circumstance in itself.’
It is, of course, true that bird-fleas are essentially nest-dwellers, but in small numbers
they occur much more frequently on the host than is generally realized, and this
becomes apparent if birds are sealed in containers impregnated with chloroform
immediately they are shot and subsequently searched.
The material in the British Museum (N.H.), including the C. Rothschild and
J. Waterston Collections, has been listed and the data arranged in Tables 2, 3, and 9
for comparison. Table 3 incorporates records generously supplied by various collectors
(see acknowledgements) and additional information obtained from a thorough search
of the literature prior to 1947.
Until recently authors have tended to record the common species of bird-fleas in
188 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
very general terms, which makes an accurate comparison of host—parasite relation-
ships almost impossible. Thus, for example, Jeffrey (1922), who collected over a
thousand fleas for Charles Rothschild, records Ceratophyllus gallinae ‘in great
abundance’ from the nests of the Fowl (Gallus domesticus), Sparrow-Hawk (Accipiter
nisus), Starling (Sturnus vulgaris), and Long-Eared Owl (Asio otus) ; ‘in large num-
bers’ from the Swift (Apus apus), and so forth. The actual number of positive nests
is not supplied, and negative records are ignored altogether. Nor is it possible to
arrive at the correct figures by examining the material in the Rothschild Collection
itself, for many specimens have been given away and exchanged. Some records of
material in the Rothschild Collection were passed on to other collectors for publica-
tion. Thus in 1936 I passed all my records of bird-fleas, including those in the
- C. Rothschild Collection, to G. B. Thompson, who published a quantity of them in
his paper ‘The Parasites of British Birds and Mammals’ (1937c). Unfortunately,
for some inexplicable reason, he omitted many of them and all the localities and
dates. It should be noted that although he included the Honey Buzzard in his list,
he did not enter any flea-species under this host! This may have been due to some
query raised at the proof stage concerning this very interesting record. The where-
abouts of the nest was kept secret by the late Mr. Meade Waldo, who discovered
it in Cornwall. The present paper includes all the records of Ceratophyllus gallinae,
C. garet, C. vagabunda insularis, and Dasypsyllus g. gallinulae which I had passed
to G. B. Thompson already at that time. The records of C. borealis with full data
have been republished by me (Rothschild, 1948). Several of the Scottish records
(specimens collected by F. J. Cox and others) were published by Waterston (1914),
and consequently care has to be exercised to avoid duplication, since the data on
the slides are not always the same as the original data as published (see Table 9).
Some slight differences will also be found between the Waterston material listed in
Table 9 and in the original papers published by Waterston himself. These discrepancies
can be explained by the following facts:
1. A number of specimens must have been given away by Waterston himself,
during his lifetime, to other collectors.
2. Some specimens are present in the collection that have not been recorded in
print.
3. A few of Waterston’s determinations were incorrect. These are principally
records of Ceratophyllus fringillae, which on re-examination of the specimens
proved to refer to C. gallinae. Two of these misdeterminations were noted by
Waterston himself (1909: 227).
4. Some mislabelling has occurred, particularly when the original labels became
illegible through age. Thus in one instance, Passer domesticus has been rewritten
on the outside of the tube as Phasianus colchicus!
In Table 3, column 2, some specimens given away in exchange between 1935 and
1950 are included, as well as material at present in the collection. In Table 9 the
specimens actually in the British Museum (N.H.) in June 1950 are listed.
Except in the case of C. vagabunda, which is not a common species, records from
the continental literature have not been taken into consideration. There are few
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS 189
fleas more difficult to separate than the closely related bird-Ceratophylli, and
consequently a large proportion of such records would require corroboration before
they could be used. Moreover, the available records for the three common bird-fleas
in Britain are quite extensive.
In this paper the host-nomenclature and the arrangement of the families of birds
have been adapted from Witherby’s Handbook of British Birds. Scientific names and
authors’ names are supplied in the tables, but sometimes omitted in the text, to
avoid unnecessary repetition. In recent years many collectors name the subspecies
of the bird host, but it has been thought better to adopt a binomial nomenclature
in Tables 2, 3, and 9 because many of the subspecific names are open to suspicion.
Thus, for example, several specimens of C. gallinae in the British Museum collection
are recorded from nests of Corvus monedula monedula from Northampton and Norfolk
in April and May; since this subspecies of the Jackdaw only occurs in Britain as
a rare autumn migrant, it is safe to assume that an error has been made either in
the identification of the host or in writing out the label. In the case of the collection
made by Miss Clay and Colonel Meinertzhagen the identification of the host can be
_ accepted without question, so in Table 1 the trinomials supplied by the collectors
are listed in column I.
Jo Bossanyi and the author made a search of the British literature prior to 1947,
including many of the minor publications. After that date only the principal entomo-
logical journals have been consulted. There can be little doubt, therefore, that some
records have been missed, but it is hoped that the list of authors and references will
assist any future research concerned with British bird-fleas. Papers containing
British records are starred.
ACKNOWLEDGEMENTS
The following collectors have kindly sent me lists of British bird-fleas from their
own collections, many of which records have not hitherto been published, and I
should like to thank them for their extremely generous help: H. Britten (45 records) ;
R. B. Freeman (34 records) ; G. D. Morison (19 records) ; E. O'Mahony (57 records) ;
G. B. Thompson (19 records) ; P. Tate (44 records from the Molteno Institute col-
lection, most of which were determined by G. H. F. Nuttall or Charles Rothschild),
and F. A. Turk (a number of negative records). I should also like to thank the follow-
ing museums and institutions for information concerning bird-fleas: Marischal
College, Aberdeen; Department of Zoology, University College of Wales, Aberyst-
wyth ; The Museum, Altrincham ; The Museum, Belfast ; The City Museum, Birming-
ham ; The Museum, Blackburn ; The Museum, Bradford ; The City Museum, Bristol;
Sexey’s School Museum, Bruton; The Molteno Institute, Cambridge ; The National
Museum of Wales, Cardiff ; The Museum, Carlisle ; The Chelmsford and Essex Museum,
Chelmsford ; The Department of Zoology, University College, Cork; The Museum,
Dunfermline ; The Museum, Glasgow ; The University, Glasgow ; The Public Museum,
Gloucester ; The Belle Vue Museum, Halifax ; The Educational Museum, Haslemere ;
190 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
The Union Lodge Museum, Hawick; The Tolson Memorial Museum, Huddersfield ;
The Mortimer Museum, Hull; The Fitz Park Trust, Keswick; The Museum, Ketter-
ing; The Museum, Kilmarnock; The City Museums, Leeds; University College,
Leicester ; Littlehampton Museum, Littlehampton ; The School of Hygiene Museum,
University of Liverpool, Liverpool; The Museum, Mansfield; Marlborough College
Natural History Society, Marlborough; The Hancock Museum, Newcastle-upon-
Tyne; The Natural History Museum, Nottingham ; The Museum, Peebles; The City
of Perth Museum, Perth; The Harris Museum, Preston ; The Department of Zoology,
University of Reading, Reading; Denstone College Museum, Rochester; The Hert-
fordshire County Museum, St. Albans; The City Museum, Sheffield; The Essex
Museum of Natural History, Stratford; The Museum, Sunderland; The County
Museum, The Royal Institution of South Wales, Swansea; The County Museum,
Truro; The City Museum, Wakefield; The Museum, Warrington; The Museum,
Warwick ; The Museum, Worthing.
I am also extremely grateful to Jo Bossanyi for searching eeugl 142 of the
lesser-known journals and periodicals dealing with British Natural History up to
December 1946, and to G. H. E. Hopkins and F. G. A. M. Smit for criticizing the manu-
script and supplying records from the literature and unpublished data. G. P. Holland
has kindly sent me the new record of C. vagabunda from Canada, and with his usual
helpfulness and enterprise K. Jordan has succeeded in obtaining several species of
bird-fleas from Russia, in order that he could himself verify interesting records of
I. Ioff. My remarks on distribution have been criticized by O. W. Richards, and
I am most grateful to him for his kindness.
DESCRIPTION OF THE COLLECTION
The collection made by Miss Clay and Colonel Meinertzhagen is listed in Table 1.
Ten species of fleas were found distributed among thirty-six species of birds, and
of these four are host-specific, and are dealt with first.
TABLE I
Fleas collected by Colonel Richard Meinertzhagen and Miss Theresa Clay
from the Bodtes of British Birds
Number
Host Species of flea and sex Locality and date
CoRVIDAE
Corvus c. corax L. : : . Ceratophyllus gallinae 1Q Shetland,
Raven Schrank (1803) Aug. 1939
Dasypsyllus gallinulae gal- 1Q Shetland,
linulae Dale (1878) Aug. 1939
Corvus f. frugilegus L. : . C. gallinae 1Q Orkneys,
Rook Aug. 1938
(18 specimens shot)
Corvus monedula spermologus Vieill. C. gallinae 22 Orkneys,
Jackdaw Aug. 1938
(36 specimens shot) Ceratophyllus vagabunda CaP Cornwall,
insularis Roths. (1906) Mar. 1946
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS tor
TABLE I—(cont.)
Number
Host Species of flea and sex Locality and date
STURNIDAE
Sturnus vulgaris zetlandicus Hart. C. gallinae 1Q Shetland,
Shetland Starling Aug. 1938
(135 specimens shot) C. gallinae Id Shetland,
5 Aug. 1939
Ceratophyllus garei Roths. 22 Orkneys,
(1902) Aug. 1938
C. gallinae 26 North Uist,
Zo Aug. 1941
FRINGILLIDAE
Fringilla coelebs gengleri D. g. gallinulae 26 Dartmoor,
Kleinschmidt Apr. 1940
British Chaffinch D. g. gallinulae Id Reading,
May 1940
Emberiza citrinella nebulosaGengler C. gallinae se County Kerry, Ireland,
Yellow Hammer Oct. 1945
(56 specimens shot) C. gallinae Id Cambridge,
May 1941
C. garet 2 Cornwall,
May 1946
D. g. gallinulae re Arran, Scotland,
Oct. 1943
Emberiza c. calandra L. D. g. gallinulae eS Cumberland,
Corn-Bunting Apr. 1941
PLOCEIDAE
Passer d. domesticus (L.) C. gallinae re Hampshire,
House-Sparrow Nov. 1944
(35 specimens shot) C. gallinae I6 Arran, Scotland,
Oct. 1943
C. gallinae ry Shetland,
Aug. 1939
ALAUDIDAE
Alauda a. arvensis L. . C. garet Ig Cumberland,
Skylark Ta? Apr. 1941
Alauda arvensis theresae D. g. gallinulae aed Co. Sligo, Ireland,
Meinertzhagen Oct. 1950
MOTACILLIDAE
Anthus pratensis (L.) . D. g. gallinulae Id Dartmoor,
Meadow Pipit 1Q Apr. 1940
Anthus spinoletta meinertzhagent . OD. g. gallinulae 1g Sutherlandshire,
Bird. Apr. 1944
Hebridean Rock-Pipit
CERTHIIDAE
Certhia familiaris britannicus C. gallinae Id Wales,
Ridgw. Jan. 1942
British Tree-Creeper
PARIDAE
Parus major newtoni Prazak C. gallinae Ig Cumberland,
British Great Tit 2 Apr. 1941
(66 specimens shot) C. gallinae Id Cumberland,
29 Apr. 1941
D. g. gallinulae 1Q Cumberland,
Apr. 1941
192. A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
TABLE 1—(cont.)
Number
Host Species of flea and sex Locality and date
Parus major newtoni (cont.) D. g. gallinulae 1Q Buckinghamshire,
July 1941
D. g. gallinulae 1Q Reading,
June 1941
D. g. gallinulae 1g Hampshire,
Apr. 1941
D. g. gallinulae 1d Cumberland,
Apr. 1941
Orchopeas wickhami 1d Berkshire,
Baker (1895) Mar. 1946
Parus coeruleus obscurus Prazak . C. gallinae 49 Cumberland,
British Blue-Tit Apr. 1941
MUSCICAPIDAE
Muscicapa s. striata (Pall.) . C. gallinae 19 Hampshire,
Spotted Flycatcher June 1941
SYLVIIDAE
Acrocephalus schoenobaenus (L.) D. g. gallinulae 1Q Hampshire,
Sedge-Warbler June 1941
Sylvia n. nisoria (Bechstein) C. gallinae 1Q Shetland,
Barred Warbler Aug. 1939
Sylvia b. borin (Bodd.) C. gallinae 1d Shetland,
Garden-Warbler Aug. 1939
Sylvia a. atricapilla (L.) D. g. gallinulae i> Winchester,
Blackcap June 1941
Sylvia c. cinerea Bechstein . D. g. gallinulae 2° Reading,
Whitethroat May 1940
TURDIDAE
Turdus philomelos hebridensis C. gallinae 19 North Uist,
Clarke. Aug. 1941
Hebridean Song-Thrush
Turdus m. merula L. D. g. gallinulae Id Co. Kerry, Ireland,
Blackbird Oct. 1945
D. g. gallinulae Id Arran, Scotland,
Oct. 1943
Oenanthe o. oenanthe (L.) D. g. gallinulae 78 Orkneys,
Wheatear Aug. 1938
Saxicola torquatus theresae Meinertz. OD. g. gallinulae Id Scilly Isles,
Hebridean Stonechat ao Mar. 1943
Erithacus rubecula melophilus Hart. C. gallinae Ig Hampshire,
British Robin Apr. 1941
C. gallinae Id Arran, Scotland,
Oct. 1943
D. g. gallinulae Id Arran, Scotland,
Oct. 1943
D. g. gallinulae 1d Arran, Scotland,
Oct. 1943
D. g. gallinulae 2Q Cumberland,
Apr. 1941
D. g. gallinulae 1d Hampshire,
Apr. 1941
D. g. gallinulae Id Co. Kerry, Ireland,
Oct. 1945
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
Host
PRUNELLIDAE
Prunella modularis occidentalis
(Hart.)
British Hedge-Sparrow
(90 specimens shot)
Prunella modularis hebridium
Meinertz.
Hebridean Hedge-Sparrow
TROGLODYTIDAE
Troglodytes t. troglodytes (L.)
Wren
Troglodytes t. hebridensis Meinertz.
Hebridean Wren
HIRUNDINIDAE
Delichon u. urbica (L.).
House-Martin
(7 specimens shot)
Riparia r. riparia (L.).
Sand-Martin
(2 specimens shot)
PIcIDAE
Dryobates major anglicus (Hart.)
British Great Spotted Wood-
pecker
STRIGIDAE
Athene noctua vidalii Brehm. , .
Little Owl
FALCONIDAE
Accipiter n. nisus (L.) .
Sparrow-Hawk
ENTOM. II. 4
TABLE I—(cont.)
Species of flea
Cy
. gallinae
io)
. gallinae
C. gallinae
D. g. gallinulae
dS
. g. gallinulae
. g. gallinulae
. g. gallinulae
D
D
C. gallinae
C. gallinae
D
. g. gallinulae
D. g. gallinulae
D. g. gallinulae
D. g. gallinulae
Nosopsyllus fasciatus
Bosc. (1800)
D. g. gallinulae
Ceratophyllus rusticus
Wagner (1903)
C. rusticus
Ceratophyllus hirundinis
Curtis (1826)
Ceratophyllus styx Roths.
(1900)
C. gallinae
C. gallinae
C. gallinae
Aa
Number
and sex
r¢
193
Locality and date
Scilly Isles,
Mar. 1945
Co. Galway, Ireland,
Jan. 1947
Winchester,
June 1941
Winchester,
June 1941
Cumberland,
Apr. 1941
Cumberland,
Apr. 1941
Co. Galway, Ireland,
Jan. 1947
Harris, Outer Hebrides,
Nov. 1949
N. Uist, Outer Hebrides,
Nov. 1949
N. Uist, Outer Hebrides,
Nov. 1949
Sutherlandshire,
Oct. 1944
Co. Kerry, Ireland,
Oct. 1945
Hampshire,
June 1941
Arran, Scotland,
Oct. 1943
Rodel, Harris, Outer
Hebrides,
Nov. 1949
Wiltshire,
Aug. 1946
Wiltshire,
Aug. 1946
Wiltshire,
Aug. 1946
Inverness,
Aug. 1941
Berkshire,
Mar. 1946
Hampshire,
June 1941
Suffolk,
date unknown
194 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
TABLE I—(cont.)
Number
Host Species of flea and sex Locality and date
HyYDROBATIDAE
Fulmarus g. glacialis (L.) . . Dz. g. gallinulae 19 Orkneys,
Fulmar Petrel Aug. 1938
CoLUMBIDAE
Columba I. livia Gm. . : . C. gallinae vag Shetland,
Rock-Dove Aug. 1939
C. gallinae ed North Uist,
Aug. 1941
Ceratophyllus columbae Id Ross of Mull,
Walck. & Gerv. (1844) Feb. 1944
RALLIDAE
Gallinula c. chloropus (L.) . . Dz. g. gallinulae 1Q Orkneys,
Moorhen Aug. 1938
TETRAONIDAE
Lagopus s. scoticus (Lath.) . . C. gallinae z¢ Orkneys,
British Red Grouse Aug. 1938
A. HOSsT-SPECIFIC SPECIES
1. Ceratophyllus styx Rothschild, 1900. This strictly host-specific flea swarms in
the burrows of the Sand-Martin (Riparia r. riparia), and according to Evans (1904)
and Waterston (1916), both sexes overwinter in the nest as pupae or imagines. Jolley
and Storer (1945) record that 20 per cent. of nestling Sand-Martins were found to
be infested, but the two males collected by Colonel Meinertzhagen (Table 1) appear
to be the first recorded from the adult bird. Owing to the large numbers of C. styx
present in individual nests, and the fact that it is sometimes found ‘free’ in sand
quarries far removed from a nest, it is surprising that it is not collected more often
as a straggler on other hosts. There are four records from nests of the Dipper (Cinclus
cinclus) in Britain (Rothschild, 1915; Britten, 1920; and Walsh, 1938), and Mr.
Edward Armstrong informs me that this bird’s habits might well bring it into contact
with this flea, since it has been known to roost in disused Sand-Martin burrows.
Mr. H. Britten has kindly sent me a record of C. styx from the nest of the Redstart
(Phoenicurus phoenicurus) collected by F. Taylor at New Mill, Cheshire, 1921,
but unfortunately neither he nor I have had the opportunity of examining this
particular specimen.
Further collecting from the nests of the Sand-Martin, especially in northern Scot-
land, is most desirable. Up to the time of writing C. styx is the only species recorded
from this host in Britain. Recently Miss Clay has collected several specimens of
C. riparius Jordan & Rothschild, 1920, and Frontopsylla lapponica Nordberg, 1934,
from the nest of the Sand-Martin in north-east Sweden. The former species has
hitherto been known as the host-specific flea of the Sand-Martin (Riparia r. riparia)
in North America (including Canada), and there are also records from Trans-Baikal
(several specimens in the C. Rothschild collection received from I. loff and the
determinations checked by K. Jordan). It is also possible that C. riparius will be
discovered in burrows of the Sand-Martin in northern Scotland, since the ranges of
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS 195
this species and of C. styx probably overlap at some point in north Europe. F.
lapponica may also occur in Scotland ; the only other record is from Finland.
2 and 3. Ceratophyllus hirundinis Curtis, 1826, and Ceratophyllus rusticus Wagner,
1903. It is a well-known fact that the nests of the House-Martin (Delichon u. urbica)
are richer in fleas, both as regards species and actual numbers, than those of any
other bird in Britain (Waterston, 1910; M. Rothschild, 1947; and Allan, 1950).
It is therefore not surprising to find that the highest number of fleas collected off
an individual host came from the House-Martin: Colonel Meinertzhagen found
twenty-five specimens of C. rvusticus on one bird and ten of C. hirundinis on another.
It is a point of considerable interest that C. hirundinis is confined to the House-
Martin in Great Britain, although it is apparently found in nests of the Swallow
(Hirundo rustica) on the continent of Europe (Wagner, 1930). In the British Museum
collection there are three specimens of C. hirundinis and one of C. rusticus obtained
in different parts of Germany in nests of the Swallow, apart from the numerous
records from the continental literature which include an exceptionally heavy infesta-
tion of C. hirundinis—2,000 specimens—from one nest. It seems unlikely that
all the existing records from this host can be attributed to careless collecting or
misidentification of the host. The same may be said of C. rusticus, which has only
once been recorded from the Swallow in Britain (Rothschild, 1947), and until recent
years was considered a rare flea in this country.
The distribution of House-Martin fleas in general presents certain baffling problems.
Judging from their flea fauna, it seems possible that relatively small communities
of these birds must remain isolated over long periods of time, and the distribution
of C. hirundinis and C. rusticus suggests the same may apply in a lesser degree to
the Swallow. In the east of Scotland Orneacus waterstoni (Jordan) 1925 and Fronto-
psylla laetus (Jordan & Rothschild) 1920 have been recorded from the nests of the
House-Martin. These fleas appear to be absent from England and Eire and over
the larger part of Europe, but occur in the Swiss Alps. A related species of Orneacus
is recorded from the Himalayas. The presence of Orneacus and Frontopsylla in these
nests and their absence elsewhere may be due to climatic changes in the past, and
could also be governed by some temperature or humidity requirements of the larvae,
but it nevertheless seems unlikely that this factor alone accounts for the distribution
of these fleas and prevents a secondary spread southwards on the House-Martin.
It is obvious in the case of C. hirundinis, which in Britain pullulates in the nests of
the House-Martin breeding in the same district as the Swallow, that this cannot
be the only explanation why it is not found in nests of the latter species.
It is also worth noting that the exchange of fleas between Sparrows and House-
Martins appears to be entirely one-sided in Britain. Ceratophyllus gallinae (Schrank)
1803 and Ceratophyllus fringillae (Walker) 1856 are found from time to time in the
nest of the latter (Evans, 1906; Waterston, 1906; Bagnall, 1921; and Rothschild,
1947), but no House-Martin fleas seem to survive on Sparrows. There is, however,
one specimen of C. hirundinis from a House-Sparrow in Germany in the C. Roths-
child collection.
4. Ceratophyllus columbae (Walckenear & Gervais) 1844. The Domestic Pigeon and
the Rock-Dove (Columba I. livia) are infested, in Europe, with a host-specific flea,
196 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
C. columbae, which is absent, however, from the dove-cots of the U.S.A. (Jordan,
1923; Rothschild, 1952 in press). The collection contains one male of this species
from a Rock-Dove shot in Ross of Mull. On two occasions C. gallinae was taken
from the Rock-Dove, and it is this species which infests the domestic pigeon in the
Eastern United States. It should be noted that C. columbae is not a parasite of the
Wood-Pigeon (Columba palumbus L.) as stated by G. B. Thompson (1937). There
are no records from this bird, except Thompson’s own record from a London Wood-
Pigeon, which had presumably come into close contact with domestic pigeons.
B. SPECIES WHICH ARE NOT HOST-SPECIFIC, BUT SHOW SOME HOST-PREFERENCE
Ceratophyllus vagabunda insularis Rothschild, 1906. Two male specimens of C.
vagabunda insularis were obtained from a Jackdaw (Corvus monedula spermologus
Vieillot) shot in Cornwall (see also O’Mahony, 1948: 89). This flea has an interesting
distribution, and also displays a somewhat peculiar choice of hosts, favouring three
widely separated groups of birds, the Corvidae, Laridae, and Order Falconiformes.
Since C. vagabunda has broken up into subspecies (Jordan, 1926) of which C. v.
insularis is peculiar to the British Isles, we may deduce that it is a relatively ancient
form. The species as a whole displays an alpine-boreal distribution, occurring in
Alaska, Canada, Spitzbergen, Great Britain, Sweden, Finland, Turkestan, Siberia,
and the Swiss Alps (Jordan, 1932). Recently there has been a record from Belgium
(Cooreman, 1947), but F. G. A. M. Smit and F. Peus (personal communications)
have failed to find it in nests in Holland and Germany, although between them they
have examined over one hundred Gulls’ nests and also Jackdaws’ nests. This type
of distribution is reminiscent of that of C. borealis, only C. vagabunda is not markedly
restricted to islands (Rothschild, 1948).
A glance at Table 2, listing the specimens in the British Museum and other British
records together with their hosts, shows that a wide variety of birds may be infested.
In the ‘Synopsis of the British Siphonaptera’ (Rothschild, 1915) this flea is described
as ‘a rare species frequenting the nests of sea birds’, and the statement is widely
quoted in the literature. There are, however, five other records from the Jackdaw
in Britain (O’Mahony, 1948, and Table 2), and it will be noted that those listed in
Table 2 were taken far inland in Northamptonshire, Midlothian, and Herefordshire.
The male from which the British subspecies of the flea was originally described
(Rothschild, 1906) was obtained from a hollow tree in Reading, Berkshire, which
had been used successively by Woodpecker, Owl, and Starling. It is more than
likely that this hole had also been occupied by Jackdaws at one time or another.
The same may be said of the hollow tree at Hampton Court, Middlesex, in which
a specimen of C. vagabunda was found inside a Crataegus seed in the store of Mus sp.
(Waterston, 1923). There is also a record of this species from the Jackdaw in Sweden
(Wahlgren, 1903 and 1907) and Belgium (Cooreman, 1947), one from the Magpie
(Pica pica) at Ashton, Peterborough, one from the Raven (Corvus corax) on Inish-
trahull (O’Mahony, 1941), and the host of C. v. alpestris in the Swiss Alps is also a
Corvid, the Mountain Chough (Pyrrhocorax graculus). In Finland Nordberg records
it from no less than roo per cent. of the Jackdaws’ nests which he examined—fifty-
seven in all—and Cooreman concluded from this that the Jackdaw was probably
ene ht eS =
—-- -
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS _ 197
TABLE 2
Records of C. vagabunda Boheman (1866)
Total No.
Name of host of records
(all from nests unless for each Location of First published
otherwise stated) Localities host specimens vecora
CoRVIDAE
Corvus corax L. Inishtrahull Isles, Co. x Nat. Mus. Dublin O’Mahony, 1941
Raven Donegal (Eire)
Corvus monedula L. Cornwall (England) I R. Meinertzhagen O’Mahony, 1948
Jackdaw Coll. (det. M.
Rothschild)
Treago Park, Hereford 2* E. O’Mahony Coll. O’Mahony, 1948 and
(England) present paper
Colinton, Midlothian 3T B.M. (J. Waterston Waterston, 1906 (as
(Scotland) Coll.) C. insularis Roths.)
Ashton, Peterborough 3 769 B.M. (C. Rothschild Present paper,t and
° (England) Coll.) O’Mahony, 1948
(Recorded as Coloeus Sweden I Probably Wahlgren Wahlgren, 1907 (as
monedula) Coll. C. monedulae)
(Recorded as Coloeus Towns and villages in 57 Unknown Nordberg, 1934
monedula) Finland
(Recorded as Coloeus Godinne (Belgium) 2§ Cooreman, 1947
monedula)
Pica pica (L.) Ashton, Peterborough I B.M. (C.R. Coll.) Present paper
Magpie (labelled (England)
Pica rusticus)
Pyrrhocorax graculus L.
Mountain Chough
Above Findelen, Zer-
matt (Switzerland)
B.M. (C.R. Coll.)
(Type of Alpine sub-
species C. vaga-
bunda alpestris J.
1926)
Jordan & Rothschild,
1920
STRIGIDAE
Bubo bubo (L.) Jomala (Finland) I Unknown Nordberg, 1934
Eagle-Owl
Strix aluco L. Arniston, Midlothian I B.M. (J.W. Coll.) Present paper
Tawny Owl (Scotland)
FALCONIDAE
Falco peregrinus Tunst.
Peregrine Falcon
ACCIPITRIDAE
Aquila chrysaétus (L.)
Golden Eagle
(From nest and nest-
lings)
Tintagel, Cornwall
(England)
Finland
Rapids, Alaska
B.M. (C.R. Coll.)
Unknown
Rocky Mountain
Lab., Hamilton,
Montana (det. Karl
Jordan; seen by
M. Rothschild) ||
* One of these records has not been published by E. O’Mahony.
t In Waterston’s (1906) paper three separate collections are given with dates and the comment
“common in the nest of Corvus monedula’. The specimens in the British Museum are dated 7.5.06 and
24.5.06, and those of 19.6.06 are missing.
{ There is no way of ascertaining whether Thompson (1937) included this record or not in his list,
since he does not indicate the source of his data. This also applies to several of the other specimens
which are entered in this column as ‘first published record in the present paper’.
§ The record reads ‘found in numbers in the nests of Coloeus monedu
it as two records.
Present paper
Nordberg, 1936
Phillip, 1938
la’, but we have only counted
|| Unless otherwise stated only the specimens in the British Museum collection have been seen by the
author.
1988 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
Name of host
(all from nests unless
otherwise stated)
Buteo buteo (L.)
Common Buzzard
Accipiter nisus (L.)
Sparrow-Hawk
Pernis apivorus (L.)
Honey-Buzzard
ANATIDAE
Anser brachyrhynchus
Baillon
Pink-footed Goose
(Recorded as Melan-
onyx brachyrhynchus)
Branta leucopsis
(Bechst.)
Barnacle-Goose
PHALACROCORACIDAE
Phalacrocorax carbo (L.)
Cormorant
(Recorded as Cor-
morant)
Phalacrocorax aristotelis
(L.)
Shag
(From nest and nest-
lings)
PROCELLARIIDAE
Fulmarus glacialis (L.)
Fulmar Petrel
COLUMBIDAE
Columba livia Gm.
Rock-Dove
LARIDAE
Larus argentatus Pont.
Herring-Gull
TABLE 2—(cont.)
Localities
Finstrom, Finland
Jomala, Finland
Cornwall (England)
Advent Bay, Spitz-
bergen
Spitzbergen
Advent Bay, Spitz-
bergen
Farne Islands,
Northumberland
(England)
Bodd and Stoil, Canna
Isle (Scotland)
Lambay Island (Eire)
St. Kilda (Scotland)
Cave in cliff, Todhead,
Kinneff (Scotland)
Burwick, Orkney Isles
(Scotland)
Cliffs near Todhead,
Kinneff (Scotland)
Ireland’s Eye, Co. Dub- 6**
lin (Eire)
Casana Rock, Howth,
Co. Dublin (Eire)
Lambay Isle (Eire)
Cruden Bay (Scotland)
Portlethen, Kincar-
dineshire (Scotland)
Muchalls, Kincardine-
shire (Scotland) ft
Total No.
of records
for each Location of
host specimens
I Unknown
I Unknown
I B.M. (C.R. Coll.)
B.M. (C.R. Coll.)
Unknown
B.M. (C.R. Coll.)
I R. S. Bagnall Coll.
I Glasgow Univ. Coll.
(det. K. Jordan)
2
I E. O’Mahony Coll.
1 BM. (J.W. Coll.)
1 BM. (J.W. Coll.)
ri) B.M. (J.W. Coll.)
2 B.M. (J.W. Coll.)
E. O'M. Coll.
I E.O'M. Coll.
r 23
2** E.O'M. Coll.
6
3
2
First published
record
Nordberg, 1934
Nordberg, 1934
Present paper
Jordan, 1932
Dampf, 1911
Jordan, 1932
Bagnall, 1921
Carrick, 1939
O’Mahony, 1945
Waterston, 1906 (as
C. insularis)
Present paper
Waterston, 1906 (as
C. insularis)
Waterston, 1910
O’Mahony, 1941, and
present paper
O’Mahony, 1939
O’Mahony, 1939, and
present paper
Allan, 1950
Allan, 1950
** Five of these records and one of those from Lambay Isle are published here for the first time.
tt The record says ‘nests’, but as no numbers are given this is treated as two records.
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
Name of host
(all from nests unless
otherwise stated)
Larus hyperboreus
Gunn.
The Glaucous Gull
(Recorded by Wagner
as Larus glaucus)
Rissa tridactyla (L.)
Kittiwake
Pagophila eburnea
(Phipps)
Ivory Gull
(From nestling in nest)
ALCIDAE
Fratercula arctica (L.)
Puffin
DomEsTIc PoULTRY
Gallus domesticus L.
Domestic Fowl
(From nest ?)
MAMMALIAN Host
Apodemus tscherga (=
Apodemus s. tscherga
Kasts.)
UNCLASSIFIED Hosts
A hole in a tree occupied
successively by Star-
ling, Woodpecker,and
Owl
Inside seed of Crataegus
in store of Mus sp. in
hollow tree
Nests of sea-birds
Coming down the rocks
On the ground ? (no
data)
Off clothing
tt The author has not seen this specimen, but jud
appears doubtful.
TABLE 2—(cont.)
Localities
North Siberia
Kidliut Bay, North-
West Territory
(Canada)
Isle of May (Scotland)
Great Skellig, Co. Kerry
(Eire)
Spitzbergen
Sheep Island, off Bal-
lintoy (Eire)
St. Kilda (Scotland)
Djarkent, Turkestan
Narankol, Turkestan
Djarkent, Turkestan
Near Reading, Berk-
shire (England)
Hampton Court,
Middlesex (England)
Shetland Isles (Scot-
land)
St. Kilda Isle (Scot-
land)
Farne Islands, North-
umberland (England)
Klass Billen Bay, Spitz-
bergen
Total No.
of records
for each
host
+
I
72
4
2
1{3
I
Several
I
I
i
Location of
specimens
Unknown. (Type of
subspecies C. vaga-
bunda orientalistt
Wagner, 1929)
B.M. and G. P. Hol-
land Coll. (det.
G. P. Holland and
K. Jordan)
B.M. (C.R. Coll.)
Nat. Mus. Coll. Dublin
B.M. (C.R. Coll.)
G. B. Thompson
Coll.
B.M. Coll. (received
from D. J. McCrae)
B.M. (C.R. Coll.)
B.M. (C.R. Coll.)
B.M. (C.R. Coll.).
(Type specimen of
British subspecies
C. vagabunda insu-
lavis Rothschild,
1906)
B.M. (J.W. Coll.)
B.M. (J.W. Coll.)
B.M. (J.W. Coll.)
B.M. (C.R. Coll.)
B.M. (C.R. Coll.)
199
First published
record
Wagner, 1929
Present paper
Rothschild, 1915
O’Mahony, 1941
Present paper
Thompson, 1937
Present paper
Waterston, 1923, and
present paper
Waterston, 1923
Rothschild, 1906
Waterston, 1923
Waterston, 1914
Waterston, 1906 (as
C. insularis)
Present paper
Present paper
ging from the description its status as a subspecies
200 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
TABLE 2—(cont.)
Total No.
Name of host of vecords
(all from nests unless for each Location of First published
otherwise stated) Localities host specimens record
Jumping about on the Kap Todsen, Isfjorden, 4 Reichsmus., -Stock- Boheman, 1866 (as
ground Spitzbergen holm. (Type of C. digitalis Wahl-
arctic subspecies gren, 1903)
C. vagabunda vaga-
bunda Boheman,
1866; seen by M.
Rothschild)
Herbage (about 50 ft. Muchalls, Kincardine I G. D. Morison Coll. Present paper
below gull’s nest) (Scotland)
the ‘normal host for this flea’. Nordberg also found it in one nest each of the Eagle
Owl (Bubo bubo), the Buzzard (Buteo buteo), the Peregrine Falcon (Falco peregrinus),
and the Sparrow-Hawk (Accipiter nisus).
The tendency to favour birds of prey (Falconidae and Accipitridae) may be
observed in Great Britain, where C. vagabunda has been collected from the nests of
the Peregrine Falcon and the Honey-Buzzard (Pernis apivorus). In Alaska it was
taken from a nest of the Golden Eagle (Aquila chrysaétus). An equally favoured
group of birds appears to be the Gulls. There are twenty-two records from the
Herring-Gull (Larus argentatus) in Britain, two from the Kittiwake (Rissa tridactyla),
one from the Ivory-Gull (Pagophila eburnea) in Spitzbergen, and the Glaucous Gull
(Larus hyperboreus) in Siberia, and in Canada (see Table 2).
Characteristics common to these hosts are difficult to find. A large proportion of
them build on precipitous rocks or cliffs, and the Corvidae, certain birds of prey,
and various Gulls, as well as the Shag and Cormorant (Table 2), construct large,
bulky nests. It is obvious, however, from a glance at the list of hosts that neither
of these requirements is essential to C. vagabunda.
It is often assumed that species which display a so-called fringing or relict distribu-
tion, like C. vagabunda, were once distributed widely over the Continent, but were
restricted to their present limited area during or following one of the ice ages. With
this possibility in mind it is tempting to speculate on the identity of the true host
of C. vagabunda in the past. The Choughs (Pyrrhocorax) instantly spring to mind.
These birds, which build bulky nests on rocky and precipitous cliffs, were undoubtedly
more widely distributed in the past. Moreover, the Mountain Chough is the host
of C. v. alpestris in the Alps, and the fairly closely related Jackdaw is the favoured
host inland in Europe today.
It would be a relatively easy task, and an interesting one, to trace the distribution
of C. vagabunda on the Jackdaw in Europe. It seems possible that at the present
time there is a secondary spread of this flea southwards again on this particular
host. Another point of interest is whether the north European subspecies, C. v.
vagabunda, is present on migrating Jackdaws which visit this country. There is
every reason to suppose that it is, and during the winter months both subspecies
are probably to be found in Great Britain.
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS 201
It is perhaps worth noting that a relatively high proportion of records of C. vaga-
bunda are of ‘free’ specimens, that is to say specimens collected away from any
particular host: ‘Jumping about on the ground’; ‘Off clothing’; ‘Coming down the
rocks’; ‘Inside a Crataegus seed’, and so forth (Boheman, 1866; Waterston, 1923,
and Table 2). The impression is created that certain areas such as sea cliffs in
particular districts, for example the east coast of Ireland (O’Mahony, 1949) or the
Tintagel district in Cornwall, are infested with these fleas, which possibly migrate
from the nests if the temperature falls, or when they become hungry (Waterston,
1910; Nordberg, 1936; and Ioff, 1946), and in due course wander on to other species
of birds nesting in the vicinity which will also serve as hosts.
If indeed C. vagabunda is a ‘relict’ species, reduced to a mere fringe of its former
distribution either by climatic changes or by the gradual elimination of its principal
host, it may owe its survival to the crowded breeding conditions on rocky cliffs along
the coast, and to its ability to spread without being actually carried on the body
of the host to its nest.
C. SPECIES WHICH ARE NOT HOST-SPECIFIC AND SHOW LITTLE OR NO
HostT-PREFERENCE
1. Ceratophyllus garei Rothschild, 1902. In this collection there are only three
records of C. garei (Table 1), compared with thirty-three of C. gallinae and thirty-
five of Dasypsyllus gallinulae gallinulae (Dale) 1878. The ratio of C. garet to the other
two common bird-fleas in this collection is therefore unusually low, approximately
1:22 instead of the expected 1:6, as shown in the overall records in Table 4. There
are two probable explanations to account for this fact. Out of the thirty-six species
of birds shot by Colonel Meinertzhagen, twenty-nine are of the Order Passeriformes.
It will be seen from Table 4 that out of a total of 108 records of C. garet only 35
per cent. are from Passerine birds, whereas 86 per cent. of records of C. gallinae
and no less than gz per cent. of records of D. g. gallinulae are from this order. The
second reason for the relatively small number of C. garei is the fact that this species
is apparently less frequently found on the bodies of its hosts than either of the two
other common British bird-fleas (Table 4).
The host-preferences of C. gavei appear to be governed more by the situation of the
nest than by any other factor. Nevertheless there are certain features concerning
its choice of host which are difficult to explain on the basis of the nesting site or
microclimate alone. It must not be forgotten that we only guess at the reason why
damp sites are chosen. Since Buxton’s excellent work on the biology of Xenopsylla
cheopis (Buxton, 1938) the most favoured explanation centres round the humidity
requirements of the larvae, but no actual experiments have been carried out with
C. garei. Moreover, very little indeed is known about the actual conditions in the
nests of different species of birds, and there are no reliable data available by which
we could compare the microclimate of a Linnet’s or a Robin’s nest.
It will be seen from Table 3 that C. garei is the only species of flea which has been
recorded from the Anatidae (sixteen records in Britain). There are also thirteen
records of this species from the Scolopacidae, from which family there are no records
of C. gallinae and only two of D. g. gallinulae. C. garei is also the dominant species
ENTOM. Il. 4 Bb
British Bird Hosts of C. gallinae, C. garei, and D. g. gallinulae
TABLE 3
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
(from the British Museum collections, the literature, and unpublished sources)
202
>
:%
Oo 8
Host “as
Corvus corax L. Ors.*
Raven
Corvus corone L, B.M.
Carrion-Crow Ors.
Corvus frugilegus L. Ors.
Rook
Corvus monedula L. B.M.
Jackdaw ee
Pica pica (L.) B.M.
Magpie
Garrulus glandarius (L.) Ors
Jay
Sturnus vulgaris L. B.M.
Starling oe
Coccothraustes cocco-
thraustes (L.) Ors.
Hawfinch
Chloris chloris (L.) B.M
Greenfinch Ors.
Carduelis carduelis (L.) B.M.
Goldfinch
Carduelis spinus (L.) Ors.
Siskin
Carduelis cannabina (L.) B.M.
Linnet Ors.
Fringilla coelebs L. B.M.
Chaffinch Ors.
Pyrrhula pyrrhula (L.) Ors.
Bullfinch
Emberiza citrinella L. B.M.
Yellow Bunting Ors.
Emberiza calandra L. B.M.
Corn-Bunting Ors.
Emberiza schoeniclus (L.) B.M.
Reed-Bunting Ors.
Passer domesticus (L.) B.M.
House-Sparrow Ors.
Passer montanus (L.) Ors.
Tree-Sparrow
Alauda arvensis L. B.M.
Skylark Ors
Anthus trivialis (L.) Ors
Tree-Pipit
Anthus pratensis (L.) B.M.
Meadow-Pipit Ors
C. gallinae
Nn
Nw Oo
29
13
38
C. garet
Ln law
—
lla lo law
Jan
2
I
» D.g. gallinulae
«|
N
Skeet
ee See (ee ag bear
2
>
ie
ss
Host “as
Anthus spinoletta (L.) B.M.
Rock-Pipit Ors.
Motacilla alba yarrelli
Gould B.M.
Pied Wagtail
Tunst.
Grey Wagtail
Motacilla cinerea cinerea oa
Certhia familiaris (L.) Von
Tree-Creeper
Sitta europoea L.
Nuthatch
Parus major L. B.M.
Great Tit Ors.
Parus coeruleus L. B.M.
Blue Tit Ors.
Parus ater L. Ors.
Coal-Tit
Parus atricapillus L. Ors.
Willow-Tit
Aegithalos caudatus (L.) B.M
Long-tailed Tit on
Panurus biarmicus (L.) B.M.
Bearded Tit
Lanius collurio L. Ors.
Red-backed Shrike
Muscicapa striata (Pall.) B.M.
Spotted Flycatcher Ors.
Muscicapahypoleuca(Pall.) Ors.
Pied Flycatcher
Regulus regulus (L.) Ors.
Goldcrest
Phylloscopus collybita B.M.
(Viell.)
Chiff-chaff
Phylloscopus trochilus (L.) { BM.
Willow-Warbler Ors.
Phylloscopus sibilatrix B.M.
(Bechst.) Ors
Wood-Warbler
Locustella naevia (Bodd.) Ors.
Grasshopper-Warbler
Acrocephalus schoenoboenus { B.M.
(L.) Ors.
Sedge-Warbler
* Includes various unpublished records as well as those from the literature.
¢ Doubtful host, designated as Passer sp.
C. gallinae
| v4
| m4
C. garet
ww D.g. gallinulae
HNHNHOA
~
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
>
Sy
= §
Host MY 8
Sylvia nisoria (Bechst.) Ors
Barred Warbler
Sylvia borin (Bodd.) B.M.
Garden Warbler Ors.
Sylvia atricapilla (L.) Ors.
Blackcap
Sylvia communis Lath. B.M.
Whitethroat Ors.
Sylvia curruca (L.) B.M.
Lesser Whitethroat
Turdus viscivorus L. B.M.
Mistle-Thrush Ors.
Turdus ericetorum Turton B.M.
Song-Thrush Ors.
Turdus merula L. B.M.
Blackbird Ors.
Oenanthe oenanthe (L.) B.M.
Wheatear Ors.
Saxicola torquata (L.) B.M.
Stonechat Ors.
Phoenicurus phoenicurus B.M.
(L.) Ors
Redstart
Phoenicurus ochrurus (L.) Ors
Black Redstart
Erithacus rubecula (L.) B.M.
Robin Ors.
Prunella modularis (L.) B.M.
Hedge-Sparrow Ors.
Troglodytes troglodytes (L.) { B.M.
Wren Ors.
Cinclus cinclus (L.) B.M.
Dipper Ors.
Hirundo rustica L. B.M.
Swallow Ors.
Delichon urbica (L.) B.M.
House-Martin Ors.
Riparia riparia (L.) B.M.
Sand-Martin
Apus apus (L.) B.M.
Swift oe
Picus viridis L. B.M.
Green Woodpecker
Dryobates major (L.) Ors.
Great Spotted Wood-
pecker
Jynx torquilla L. B.M.
Wryneck
Athene noctua (Scop.) B.M.
Little Owl Ors.
Asio otus (L.) B.M.
Long-eared Owl Ors.
C. gallinae
Lal
Lal
RW OUR HW ORO UW H Ca) hay De PUL as ek agian
HW
TABLE 3—(cont.)
D g. gallinulae
Host
Strix aluco (L.)
Tawny Owl
— Tyto alba (Scop.)
—_- — Barn-Owl
noe Accipiter nisus (L.)
Sparrow-Hawk
Anas platyrhyncha L.
Mallard
Anas acuta L.
Pintail
Spatula clypeata (L.)
Shoveler
Aythya fuligula (L.)
Tufted Duck
al
||
ee oH
Common Eider
Melanitta nigra (L.)
Common Scoter
Mergus serrator L.
LTT Ta toww II
re I pe te eae
Shag
Hydrobates pelagicus (L.)
Storm-Petrel
Fulmarus glacialis (L.)
Fulmar Petrel
Columba palumbus L.
Wood-Pigeon
Columba oenas L.
Stock-Dove
Columba livia Gm.
Rock-Dove
Scolopax rusticola L.
Woodcock
Capella gallinago (L.)
Snipe
Calidris alpina (L.)
Dunlin
Tringa totanus (L.)
Redshank
Charadrius hiaticula L.
Ringed Plover
Vanellus vanellus (L.)
Lapwing
|
Sterna macrurva Naumann
Arctic Tern
Larus ridibundus L.
Black-headed Gull
Larus canus L.
Common Gull
Larus fuscus L.
Somateria mollissima (L.)
Red-breasted Merganser
Phalacrocorax aristotelis (L.)
Lesser Black-backed Gull
hHU C. gallinae
Lal
[epececateimeaare las fines
C. garei
Lal
nN
nd
°
Ww
D. g. gallinulae
204 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
TABLE 3—(cont.)
: :
3
Ss 3 § %&» 8 = & %&
i ea s; 8 iG
Host aN OO A Host aN oS On
Larus marinus L. Ors — 1 — ~~ Lagopus scoticus (Lath.) BM. — 2 2
Great Black-backed Gull Red Grouse Ors. I (3\noe
Rissa tridactyla (L.) Ors I—- — Phasianus colchicus L. B.M. 2 5.
Kittiwake Pheasant Ors. — 2 —
Stercorarius parasiticus(L.) BM. — «1 — Perdix perdix (L.) BM — 4ft —
Arctic Skua Common Partridge Ors. — 2f —
Fratercula arctica (L.) Ors. j heey Me: v Alectoris rufa (L.) BM — «1 —
Puffin Red-legged Partridge
Crex crex (L.) BM. — 2 — Gallus domesticus L. BM. 13 — —
Corn-Crake Fowl pes 23 I —
Gallinula chloropus (L.) B.M. i 4 4
Moorhen Ors. — «I 2
¢ This is an under-estimate since two of these six records were ‘from several nests’.
in the nests of the Phasianidae and Tetraonidae (Game-birds), from which there are
nineteen records as opposed to six of D. g. gallinulae and three of C. gallinae. It is
also found more frequently in the nests of the Laridae than either of the other two
species. In Britain it is the only flea recorded from the Linnet (Carduelis spinus)
and the Bearded Tit (Panurus biarmicus), and despite the paucity of records it may
be considered the dominant species in the nests of the Reed-Bunting (Emberiza
schoeniclus). This list is sufficient to show C. garet’s predilection for ground-nesting
species which favour damp sites and build open nests. Nevertheless, it has been
collected from the nests of a great variety of birds, and Table 3 shows that it has been
recorded more than once from the Starling (Sturnus vulgaris), Blackbird (Turdus
merula), and Song-Thrush (Turdus ericetorum). There are some curious omissions
from its list of hosts in Britain. One would expect a number of records from the
Swan, the Herring-Gull, Dipper, Wren, and Robin, but C. gave has never been
recorded from any of these birds in Britain, although there is one specimen in the
C. Rothschild collection from Tyvoglodytes troglodytes in Germany.
It appears that nests of the Wren (Tvoglodytes troglodytes) and Robin (Evithacus
vubecula) present optimum conditions for D. g. gallinulae. Whatever these condi-
tions may be—and the difference between the microclimate presented by a Robin’s
nest and a Linnet’s nest appears superficially to be rather slight—they possibly
prove unsuitable for C. gavei. In these circumstances competition from D. g. galli-
nulae may prove too strong. On the other hand, host preference of the imago and
not the requirements of the developmental stages may in part account for this
distribution. It is less curious that there is only one record from the ground-bird
par excellence, the domestic fowl, since it nests under exceptionally dry conditions.
As is pointed out in Table 4, C. garei occurs less frequently on the bodies of its
hosts than either C. gallinae or D. g. gallinulae. Unfortunately the relevant data
are often omitted both in the literature and on labels on slides, but certain ento-
mologists such as James Waterston always noted whether specimens came from
4
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS 205
the body or nest of their host. In other cases collectors like W. H. Pollen and
F. J. Cox can supply the necessary data. Generally speaking, if a flea is collected
from the body of a bird the fact is considered of interest and is therefore noted, so
most of the material without specific information on this point comes from nests.
In the Charles Rothschild collection (Table 9) there is a batch of male and female
C. garei which bears this label: ‘Found by taxidermist in Natural History Museum,
London. Land-Rail.’ Although there is no mention of the body of the host, in this
TABLE 4
Comparative Records of C. gallinae, C. garei, and D. g. gallinulae (compiled
from all known British records)* —
Species of flea . E : : : ; C. gallinae C. garet D. g. gallinulae
Total number of records . ; ; : 363 108 266
Number of known species of bird-hosts in
Britain. , A ; ‘ : ‘ 65 49 59
Percentage on Passerine hosts . : ‘ 86 35 92
Number of records from body of host. (Note: 58 10 57
Many of these records are of more than one (16 per cent. of (9 per cent. of (21 per cent. of
specimen) total records) total records) total records)
Proportion of sexes on body of host . ‘ 60 2 89 339
373 23 315
Proportion of sexes in nest (only collections 3152 772 2169
in B.M. of under 20 specimens counted) 18849 695 126d
Percentage of collections (from individual 31 per 30 per 13 per
nests) in B.M. with over 20 specimens per cent. cent. cent.
nestt
* These include unpublished records kindly supplied by various authors (see acknowledgements).
Records away from a host, from mammalian hosts, or from unspecified birds’ nests are omitted.
t In actual fact the percentage is probably much higher, since collectors often preserve a small fraction
only of the specimens they find in a nest.
case I have assumed that these specimens were collected from the bird itself. Apart
from this one batch of fleas, only specimens definitely known to have come from
the host are counted in line 5 of Table 4. Since the totals in line I contain a number
of records in which the relevant data are missing altogether, the figures for the number
of fleas from the bodies of birds may be on the low side, but this should not affect
the relative frequency of the three species concerned. Furthermore, there are a
number of records in which no host is named at all: ‘Found in old bird’s nest’, or
‘From old nesting box’ are all-too-familiar labels. Since these have been omitted
as well as the records of ‘free’ specimens, there is probably adequate compensation
for the odd flea which may have come from the body of its host and for which there
are no available data. Altogether it seems likely that these figures, 9 per cent., 16
per cent., and 21 per cent., give a fair picture of the relative frequency of the three
common bird-fleas on the bodies of their hosts.
There are inadequate numbers (ten specimens only) for comparing the ratio of
the sexes for this species. With this point in mind, fifty-six collections of C. garet
(from the British Museum) from nests of named bird hosts were examined. Thirty-
206 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
nine contained less than twenty specimens per nest, and the sex-ratio was 69 males
to 77 females. The remaining seventeen collections contained more than twenty fleas
per nest, and three collections out of these showed a great preponderance of females.
One of the heavy infestations came from the nest of the Bearded Tit (Panurus
biarmicus), and others were from Anatidae (four), Perdix perdix (three), and Phalacro-
corax aristotelis (two).
In a previous paper (Rothschild, 1948) I described C. garet as a ‘circumpolar
species extending southwards to central Europe and through Alaska and Canada
to the western United States’. It is well distributed on the mainland of Britain,
but according to O’Mahony (1948) is relatively scarce in Ireland. This point is
further discussed on page 212.
2. Ceratophyllus gallinae (Schrank) 1803. There are thirty-two separate records
of this flea in the collection, but in actual numbers it exceeds D. g. gallinulae—fifty-
two specimens compared with forty-four. C. gallinae has a sound claim to be the
commonest bird-flea in Britain. It occurs on a greater variety of hosts, and the
total number of records from birds in Britain (excluding the Fowl) about equals
TABLE 5
Selected Hosts on which C. garei is the Dominant Species of Flea
C. garei C. gallinae OD. g. gallinulae
Name of host records vecovds records
Carduelis cannabina (L.) . ° 3 fo) fe)
Linnet
Panurus biarmicus (L.) : 2 fe) °
Bearded Tit
Anas platyrhyncha L. “ 4 fo) °
Mallard
Somateria mollissima (L.) . ‘ 4 fo) fo)
Common Eider
Capella gallinago (L.) ; ‘ 4 ° °
Snipe
Vanellus vanellus (L.) ; : 5 ° I
Lapwing
Perdix perdix (L.) . : 6 fe) °
Common Partridge (three of
these records are from ‘several
nests’)
the combined collections of both C. garei and D. g. gallinulae. Moreover, a higher
proportion of C. gallinae infestations may be described as heavy. It will be seen
from Table 4 that 16 per cent. of the total records were from the body of the host,
less than in the case of D. g. gallinulae but significantly more than in that of C. garei.
It is also interesting that in this collection the number of female fleas on the host —
outnumbers the males by approximately 3:2, whereas they are about equal in the
case of D. g. gallinulae. In eighty-one collections from named birds’ nests (see Table
4) the sex-ratio was also approximately 3:2.
In the ‘Synopsis of the British Siphonaptera’ (Rothschild, 1915) the hosts of
C. gallinae are given as ‘the fowl and many birds’, and again ‘in the nests of most
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS 207
birds and hen-houses’. In 1930 Wagner was more precise and lists: ‘ Birds, principally
Gallus, Passer, Parus’. Freeman (1941) takes a wider view and considers that
C. gallinae favours ‘almost all birds’.
It is shown in Table 4 that (excluding collections from the Fowl) 86 per cent. of
British records of C. gallinae are from Passerine birds. It does not occur on the
Anatidae (two specimens recorded in the continental literature from ducks have not
been checked and are probably errors in determination) or Scolopacidae and there
is only one record each in Britain from the Rallidae and Laridae. The wild Jungle
Fowl (Gallus gallus L.) is not infested with C. gallinae, and Jordan (1928) suggests
that the domestic fowl is a secondary host and that the primary hosts in Europe
may be the Paridae.
If we examine the list of records in Table 3 it will be noted that, despite the wide
range of hosts, C. gallinae is much more common in the nests of birds which build
in holes and some distance from the ground. These nests (as well as hen-houses)
are almost certainly drier than those built in low situations. Thus C. gallinae is
the dominant species in the nests of such birds as the Starling, House-Sparrow,
Jackdaw, and Blue-Tit (see Tables 3, 6, and 9).
TABLE 6
Selected Hosts on which C. gallinae is the Dominant Species of Flea
C. gallinae C. garei D. g. gallinulae
Name of host vecovds vecords records
Corvus corone L. : ; : y | I I
Carrion-Crow
Sturnus vulgaris L. . : q 42 2 6
Starling
Passer domesticus (L.) ; : 46 fo) 3
House-Sparrow
Parus major L. : ‘ ° 25 fo) 9
Great Tit
Parus coeruleus L. . : > 36 fe) 4
Blue-Tit
Strix aluco L. : “ 6 fo) fe)
Tawny Owl
Athene noctua (Scop.) : ° 5 ° fe)
Little Owl
Accipiter nisus (L.) . ° : 5 fe) ; I
Sparrow-Hawk
Out of the thirty-eight collections (from the British Museum) which contain more
than twenty specimens per nest, eighteen came from Parus spp., six from Passer
domesticus, and five from Sturnus vulgaris.
C. gallinae is widely distributed in Europe and in parts of Asia. There are speci-
mens in the British Museum collection from France, Holland, Switzerland, Germany,
Austria, Hungary, Roumania, Russia, north China, Mongolia, and east Turkestan.
The species has probably been introduced into the United States and certainly into
New Zealand, and has passed on to various species of indigenous hosts. Its range in
208 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
the United States is restricted to the eastern States. It is found more frequently than
other bird-fleas as a straggler on mammals. It is also recorded quite frequently away
from any host (see Table 8), and has been noted by Waterston (1910) migrating from
nests in impressive swarms. Bacot (1914) and Ioff (1946) have shown that this
species can survive over a year in empty birdnests. Although it seems clear that C.
gallinae may pass the winter on the body of its host and thus be transported back to
the new nest when it is built, it is likely that birds can also be reinfested at their
nesting-sites in the spring by fleas which have over-wintered under the bark of trees
and in piles of rubbish, dead leaves, or old nesting-holes.
3. Dasypsyllus gallinulae gallinulae (Dale) 1878. Miss Clay and Colonel Meinertz-
hagen collected thirty-five records of this interesting species. It is shown in Table 4
that D. g. gallinulae occurs in a higher proportion on the bodies of birds than the
other two common species, and also that 92 per cent..of its hosts are Passerines.
These two facts are sufficient to account for its preponderance in the collection.
It will be noted, for instance, that if all records of fleas from Parus major are pooled
(Table 3), we find that C. gallinae exceeds D. g. gallinulae on this host by about 3:1,
yet in this collection (Table 1) the reverse is true, and the ratio is 1:2.
The distribution of this species is one of considerable interest. The genus Dasy-
psyllus is essentially Neotropical, and D. g. gallinulae is the one representative in
Britain of a characteristic South American genus. There are five species recorded
from ground-nesting birds in Chile, Brazil, and the Argentine ; one species in Sumatra
on rats (probably an accidental host) ; and D. stejnegeri is known from Behring Isle
and Commander Isle, Siberia, St. Paul Island (Pribilov Group), Alaska, and Langara
Isle, Queen Charlotte’s (British Columbia), and is probably parasitic on sea-birds,
though the host is unknown (Jordan, 1929, and Holland, 1949). D. g. gallinulae is
represented in Mexico, the U.S.A., and Canada by a subspecies, D. g. perpinnatus
Baker, 1904. It is confined to the Pacific coast of the North American continent,
although many of its bird hosts extend their range far eastwards. It is suggested
by Holland (1949: 30) that ‘this flea requires a type of nest (the exact genus or species
of the bird host not being important) in country supporting a climate with a rela-
tively high mean humidity, and perhaps certain temperature limits. As the nests
of most Passerines are open and exposed directly to the atmosphere, it may be quite
readily seen that it is possible for the humid atmosphere of the coast to present
suitable breeding conditions for these fleas, whereas the dry climate and more
extreme temperatures of the interior of British Columbia, even though the same
birds were nesting, would prohibit development. Limitation of distribution of this
type thus is almost undoubtedly controlled by factors which affect the flea larva
rather than the adult.’ This seems a sound hypothesis, and the distribution of
D. g. gallinulae among bird hosts in Britain gives us a hint that it, too, may not
be able to survive in dry nests.
C. gallinae is the dominant species in relatively dry nests which are built in holes
or in the open but some distance from the ground, and C. garei is characteristic of
ground-nesting species, especially those which favour damp sites, whereas D. g.
gallinulae is found most commonly in the nests of Passerine birds building near the
ground. The flea fauna of the Paridae in Britain present an excellent example of
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS 209
this fact, since C. gallinae is predominant in nests of the Blue-Tit (Parus coeruleus)
and Great Tit (Parus major), C. garei in the nests of the Bearded Tit (Panurus
biarmicus), and D. g. gallinulae in the nests of the Long-tailed Tit (Aegithalos cauda-
tus). D. g. gallinulae is also dominant in nests of the Robin, Redstart, Willow-
Warbler (Phylloscopus trochilus), and Greenfinch (Chloris chloris). There are no
records at all of this flea from the Hirundinidae, Strigidae, Laridae, or Anatidae,
and none, either, from the Fowl ; however, it exceeds all others on the Sylviidae, with
23 records against 7 of C. gallinae and 2 of C. garei, and on the Turdidae with 85
records against 44 and 8 respectively.
Since we know nothing of the relative humidity of any nests, this suggestion must
not be pushed too far. Moreover, the number of records from various hosts is too
small to be significant. It is also obvious that the overlap between all these three
species of fleas, particularly C. gallinae and D. g. gallinulae, is considerable (see Table
3), and there are several instances which run directly contrary to this theory. Thus
there are more records of D. g. gallinulae than of C. garei in the nests of both the
Moorhen and Dipper. There is an equal number of records of C. gallinae and D. g.
gallinulae from the Mistle-Thrush (Turdus viscivorus), although the former has a
10:9 majority over D. g. gallinulae in nests of the Song-Thrush (Turdus ericetorum),
Despite these and numerous other contradictions, the data contained in Table 3,
meagre though they are, suggest that all these three common species of bird-flea,
not only C. gare, have a preference for birds favouring certain types of nesting sites.
TABLE 7
Selected Hosts on which D. g. gallinulae 1s the Dominant Species of Flea
D. g. gallinulae C. gallinae C. garei
Name of host records records records
Chloris chloris (L.) . ; “ 9 2 oO
Greenfinch
Fringilla coelebs L. . 7 é 10 5 I
Chaffinch
Phylloscopus trochilus (L.) : 8 fo) fo)
Willow-Warbler
Sylvia communis Lath. . : 5 I fe)
Whitethroat
Turdus merula lL. . ps 3 a7 22 4
Blackbird
Erithacus rubecula (L.) . : 33 4 oO
Robin
Troglodytes troglodytes (L.) ‘ 16 12 fo)
Wren
Gallinula chloropus (L.) . : 6 I 5
Moorhen
D. g. gallinulae is not found in such large numbers in the nests of birds as either
C, gallinae or C. garei (see Table 4). Out of ninety-three batches from the nests of
named bird hosts in the British Museum collection, only 13 per cent. contained more
ENTOM. II. 4 CO
z1o A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
than ten specimens of either sex. The hosts which are most heavily infested
numerically are the Warblers and the Robin, which are also the hosts with the
relatively largest number of records. One heavy infestation came from the nest of
the Moorhen, proving that this species can breed successfully in what appear to be
especially damp nests.
The numbers of the sexes of D. g. gallinulae on the body of the host is almost
equal (see Table 4), but in the nest there is a heavy preponderance of females—
216:126 in the eighty batches counted.
The range of D. g. gallinulae on the continent of Europe is not known with any
accuracy. In the British Museum collection there are specimens from France,
Holland, Germany, Switzerland, Portugal, Italy, Roumania, Iceland, the Azores,
Madeira, and a pair from the Nepal-Sikkim frontier of northern India.
TABLE 8
British Records of C. gallinae, C. garei, and D. g. gallinulae
collected away from any Host or Nest
Species of flea Location Locality Date Collector Collection
C. gallinae Refuse dump N. Ireland 1939 Received from Rothschild Coll.
(Bidi3.2) Dept. of Agri- (B.M.)
culture
C. gallinae Dead leaves in Kennetpans, 1914 British Museum *
(x 9) orchard Kincardine on
Forth
C. gallinae Beneath beech Hylton Bridge, 7.xi05 R.S. Bagnall on
(x Q)* bark Sunderland
C. gallinae In rotten oak- Sherwood 30.v.27. C. E. Stott -
(1 9) tree
C. gallinae In hollow tree Wroxeter, 23.vViii.i2 G. A. K. Marshall bs
(1d, 3 ) Uppington,
Shropshire
C. gallinae Found ina Newport, X.49 J. Maclagan rf
(many) garden Shropshire
C. gallinae From keeper’s Leicester 3.iv.1899 H. B. Headley sp
(x1 g, 1 Q) larder
C. gallinae On palings Leicester 17.iv.1899 H. B. Headley #
(many)
C. gallinae In moss Oxford 6.iii.1897 J. Shipp a
(r 8)
C. gallinae Warehouse Manchester 1930 H.R. P. Collett H. Britten
C. gallinae Under bark of Hylton, Co. v.09 R. S. Bagnall One specimen
(many) beech-trees Durham Rothschild Coll.
(B.M.)
C. gallinae Cruciferous Gibside, Co. v.09 R. S. Bagnall Bagnall Coll.
(2 3, 3 9) plant Durham
C. gallinae Stump of Sorbus Drum, Aberdeen- 15.v.42 G. D. Morison Morison Coll.
(x 9) aucuparia shire
C. gallinae Dead leaves in Brodie, Nairn, 20.11.45 a e
(1 9) glasshouse Morayshire
* Although there is only one specimen in the Rothschild Collection, Bagnall (1921) records it ‘in
numbers under beech tree bark’ (Bagnall, R. S., Tvans. Nat. Hist. Soc. Northumberland & Durham, 1921,
5: 181-198).
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS air
TABLE 8—(cont.)
Species of flea Location Locality Date Collector Collection
C. gallinae In barns Holton, Suffolk 1913 Strickland and) f
(in numbers) Merriman
C. gallinae Hole in an oak- Holton, Suffolk 1913 Strickland and
tree Merriman
C. gallinae From Vespa Harpenden, 17.iii.2t R. Stenton British Museum
(1 9) vulgaris Herts. (Min. of Agr.)
C. garei Sandhills Hynish, I. of Ti- 30.iv.12 H.Donisthorpe Rothschild Coll.
(1 3) ree, Argyllshire (B.M.)
C. garet In a stack of Funton, nr. 19.iii.98 J. J. Walker a
(x 9) reeds Iwade, Kent
C. garet Haystack refuse Hunstanworth, ? R. S. Bagnall Bagnall Coll.
(1 g, 2 9) Co. Durham
D. g. galli- In acave onsea Donegal, Ireland vii.22 E. E. Austen Rothschild Coll.
nulae (2 9) cliffs (B.M.)
D. g. galli- Among stones in Near Portsalon, 8.viii.22 G.C. C. Damant fe
nulae (3 9) cave in sea cliffs Lough Swilly,
Co. Donegal
D. g. galli- Read’s Cavern Burrington 4.v.40 G,. A, Walton 3
nulae Coombe, Men-
(3 3, 5 Q) dip Hills, N. Som.
D. g. galli- In grass Rainow, Cheshire 7.vi.19 F. Neave H. Britten
nulae
D. g. galli- Swept off Vac- Long Hill, Co. 21.v.23 A. W. Stelfox Nat. Mus.,
nulae cinium Wicklow Dublin
D. g. galli- Dry bark of Craibstone, 5.iv.43 G. D. Morison Morison Coll.
nulae (1 Q) cherry-tree Aberdeen
D. g. galli- Herbage Craibstone, I.v.41 r PP
nulae (1 9) Aberdeen
D. g. galli- Pine log Drum, 15.V.42 os ¥
nulae (1 Q) Aberdeen
D. g. galli- Among herbage Newtonhill, 21.V.49 .
nulae (1 9) Kincardine
+ These records were published in Parasitology, 1913, 6 (1):
by G. H. F. Nuttall, C. Strickland, and G. Merriman), but the whereabouts of the specimens is not
known.
D. STRAGGLERS FROM MAMMALIAN HOSTS
I-19 (Observations on British rat-fleas,
1. Nosopsyllus fasciatus (Bosc. D’Antic) 1801. One specimen was collected from
Troglodytes t. troglodytes in Arran. The normal host is the Brown Rat (Rattus norve-
gicus). There are three females and one male of this species in the British Museum
collection from a Blackbird’s nest in Holland (Smit, 1949).
2. Orchopeas wickhami (Baker) 1895. One specimen of the Grey Squirrel flea,
which was originally imported into this country from the U.S.A. with its host,
Sciurus carolinensis, was collected from Parus major newtoni Prazak. This species
of flea is a more frequent straggler on to avian hosts. In the Charles Rothschild
collection there are specimens collected from ‘a Magpie’s and Hawk’s nest’, and an
unnamed bird’s nest in the New Forest, Hampshire. An exchange of fleas between
birds and squirrels is, in fact, a usual accident, and Monopsyllus sciurorum (Schrank)
1803 has been known to breed in large swarms in an Owl’s nest (Waterston, IgIo),
212 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
while C. gallinae has frequently been recorded breeding in Squirrels’ dreys in Britain,
although not in large numbers (Freeman, 1941).
SOME GENERAL CONSIDERATIONS RELATING TO C. GALLINAE,
C. GAREI, AND D. G. GALLINULAE
Geographical distribution. With regard to the three common species of bird-fleas
C. gallinae, C. garet, and D. g. gallinulae in Britain, the maps in Figs. I-3 all tell
much the same story. These fleas are widely distributed throughout the British Isles
and certain obvious concentrations mark the home territories of well-known col-
lectors such as C. Rothschild, H. G. Jeffrey, R. Newstead, H. Britten, and J. Water-
ston. In Ireland, according to E. O’Mahony, C. garez is relatively scarce, but unless
lists of negative records are provided such statements may be somewhat misleading.
If a long series of Ducks’ nests, or Snipes’ nests, yielded no specimens of C. garet
this would be significant. In Cornwall, Turk (1946) considered C. gallinae ‘not so
common as it seems to be in other counties. Its place seems to be taken by D. galli-
nulae, which is a very common species. ... I have records from many small Passerine
birds and from Blackbirds, the domestic pigeon and the Wood-Pigeon.’ However,
if we pool all records for the British Isles (see Table 3), D. g. gallinulae is seen to be
the dominant species in the nests of the Blackbird (G. B. Thompson, 1937, records
350 specimens from one nest), and the same may apply to the unnamed ‘small
Passerines’ to which Turk refers earlier. Since going to press Dr. Turk has sent mea
list of these species. They include the Robin, Wren, and Tree-Creeper, in which
D. g. gallinulae is usually dominant; the Thrush, Mistle-Thrush, and Goldcrest, in
which nests both species have been recorded an approximately equal number of times;
and the Great Tit, Blue-Tit, Jackdaw, Carrion-Crow, Raven, and Rook, in which
nests C. gallinae is usually the dominant species of flea. In all, Dr. Turk found
D. g. gallinulae ‘in nearly a hundred nests’. In Ireland O’Mahony (1939) has come to
a somewhat similar conclusion with regard to these two species, namely that D. g.
gallinulae is ‘the commoner of the two in point of numbers’, but again the seven
species of birds’ nests he has listed in which D. g. gallinulae outnumber C. gallinae
include the Wren, the Greenfinch, the Chaffinch, the Mistle-Thrush, the Song-Thrush,
and the Blackbird. D. g. gallinulae is usually dominant in five of these, and in the
nest of the Song-Thrush the two species are about equally represented (see Tables
3 and 6). He did not record D. g. gallinulae at all from nests of the House-Sparrow
or the Blue-Tit—records which would have been far more significant. It would be
of much interest if further collecting in the west of England and Ireland confirmed
the suggestions made by O’Mahony and Turk, since the distribution of D. g. gallinulae
among its hosts in Britain and the distribution of the subspecies D. g. perpinnatus in
North America suggest that it requires rather a high mean humidity in the nest.
Thus it might possibly increase the range of its Passerine hosts where the mean
annual rainfall is greater, and also occur in larger numbers in the nests of certain
species where it is numerically sparse in drier conditions. In the meantime, how-
ever, the distribution maps of C. gallinae, C. garei, and D. g. gallinulae (Figs. 1, 2,
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS 213
and 3) show the distribution of the collectors rather than that of the fleas, and
indicate that all three species occur where they are looked for.
A few observations on the distribution of C. vagabunda and the Martin fleas are
included in the description of the collection on pages 195 and 196. With all the
species concerned further collecting is necessary before any conclusions can be
drawn. From the little we know, however, it is obvious that the flea fauna of
Scotland is of considerable interest. There are several species, such as C. borealis,
Orneacus waterstoni, and Frontopsylla laetus, which display a so-called alpine-boreal
type of distribution, occurring in Scotland and the Swiss Alps and in some cases
along the northern fringe of Europe, but not apparently in England or the inter-
vening areas of the Continent. This type of distribution is even more striking in
the case of fleas than in other insects, since the host itself is widely distributed.
On the other hand, C. hivundinis occurs on the House-Martin throughout its range.
It is, however, clear, as we have seen in the case of D. g. perpinnatus, that other
factors limit the distribution of the fleas, presumably requirements of the develop-
mental stages. Although many of the Passerine hosts of bird-fleas migrate to South
Africa and other warm countries in the winter, and (we may surmise) take their
fleas with them, the Ceratophyllid bird-fleas have remained a group which is charac-
teristic of temperate climates.
In east Africa over a period of fourteen years, G. H. E. Hopkins and a team of
his friends examined over 2,000 birds shot for food or scientific purposes. Ecto-
parasites were collected by the method described in the introduction. Not a single
specimen of Ceratophyllus was found, thus confirming the fact that the apparent
absence of this genus on birds, in tropical Africa at least, is genuine. It should be
noted that a number of migrants were included in this collection. Thus fifteen
specimens of the Sand-Martin (Riparia riparia) and six specimens of the Swallow
(Hirundo rustica) were among the negative records. There were also smaller numbers
of birds such as the European Cuckoo (Cuculus c. canorus), the Whinchat (Saxicola
rubetra), the Green Sandpiper (Tvinga ochropus), and the Wood Sandpiper (Tringa
glareola).
Host-preference. Among the three species of common bird-fleas which are not
host-specific we can discern two different trends of host-preference. On the one hand
there is a marked predilection on the part of D. g. gallinulae and C. gallinae for the
Order Passeriformes, and on the part of C. garei for the Charadriiformes, Anseri-
formes, and Galliformes. Within this framework we can—despite a big overlap and
many exceptions—discern a preference for different types of nesting sites. Broadly
speaking, C. garei favours damp nests on the ground, D. g. gallinulae those built near
the ground, and C. gallinae nests built in drier situations some distance from the
ground. There is also one generalization which may be made regarding the host
relationship of all these three common species: those birds which are the most
frequently infested and may be termed the favoured hosts are also the most heavily
infested. Thus C. gallinae, which can breed in such a wide variety of nests, is found
regularly in greatest numbers and obviously ‘does best’ in the nests of Passer
domesticus, Sturnus vulgaris, Parus, and certain Strigidae. This fact is reflec-
ted in the number of specimens of C. gallinae collected from individual birds by
60°
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Fic. 1. Distribution map of C. garei in Britain, based on 108 records.
In the case of multiple records from one locality, for example twenty-three separate records of C. gallinae from
Blue-Tits’ nests in Cambridge, only one circle is used. This accounts for the fact that there are more records
than circles on each map. Massed records of this type merely indicate intensive collecting in the area concerned,
and have no bearing on the question of distribution.
12° 11° 10° 9 ° 8° 7 ° 6° 5 ° 4° ‘2 oa 7° fe) ° it
Fic. 2. Distribution map of C. gallinae in Britain, based on 363 records.
oO 1 2 3 4 5 6
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Fic. 3. Distribution map of D. g. gallinulae in Britain, based on 266 records.
a’
a
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS 217
Meinertzhagen and Pollen, thus six, four, and four specimens from Starlings, four,
three, and two specimens from Parus spp., and nine and six specimens respectively
from individual Little Owls (Tables 1 and 9).
Bacot (1914) has pointed out that in some cases egg-production of Nosopsyllus
fasciatus falls off in drought, and Buxton (1938) has shown that in the case of
X. cheopis ‘even at one temperature and within tolerable limits of humidity the
production of the first filial generation of fleas is closely dependent on humidity,
the production being less as the fatal limit is approached’. Edney (1947) has shown
that the life of the adult X. brasiliensis is shortened if the pupal stage is kept at low
humidities, and these types of phenomena may account for the fact that all three
common bird-fleas occur in smaller numbers in some nests than in others, only the
favoured hosts providing the optimum conditions. It is of course possible that we are
concerned with a combination of many factors, and not only the humidity require-
ments of the developmental stages. However, in this fact we can see indications of
incipient or vestigial host-specificity—either a legacy from the past or a hint of
things to come.
There are also indications of a third type of host-preference (such as we find in
C. vagabunda) which at times seems to cut across the two main trends already
mentioned. Undoubtedly certain birds appear to be more attractive or susceptible
than others to one or all of the species concerned. Thus the Shag (Phalacrocorax
aristotelis) and Cormorant (Phalacrocorax carbo) are placed in the same genus, and
build the same type of nest in the same type of habitat, frequently in colonies which
adjoin. There are three records of C. garei from the former (including one very heavy
infestation where over 500 fleas were counted) and none at all from the latter.
There are also more than six records from the Partridge (Perdix perdix)—a bird
which is not infested with any other flea (Table 5). However, both C. gallinae and
D. g. gallinulae as well as C. garei have been collected from the Pheasant (Phasianus
colchicus) and Red Grouse (Lagopus scoticus). Here again there would appear to be
little if any difference between these three nests, which are no more than sheltered
scrapes lined with grass; in fact Pheasants and Partridges not infrequently lay in
the same nest. It is also curious that D. g. gallinulae should have a marked predi-
lection for the Moorhen (six records), since this bird is not a Passerine and (more
important still) favours unusually damp nesting-sites.
The records from the Wood-Pigeon (Columba palumbus) suggest that this may
also be a case in which the bird itself is attractive to fleas, since the type of
nest it builds is exceptionally unsuitable for nidicoles. In Finland Nordberg
(1936) has examined the nests of over fifty species of birds. He has shown that
nests built in holes and what he terms ‘half-holes’ (halbhélen Nester) support the
heaviest populations of nidicoles generally, and also the greatest variety of species.
The Stock-Dove (Columba oenas) is one of the most heavily infested of all, with a
total of fifty-eight species of nidicoles and 6,573 specimens per I dm.® of nesting
material. The Wood-Pigeon, with one species and only twenty specimens per I dm.?,
is unquestionably the least infested out of all those he examined in Finland. In
Britain there are only four records of C. gallinae from the Stock-Dove, but from the
Wood-Pigeon we have records of the following species: C. gallinae (I—but 6 females
ENTOM. Il. 4 Dd
218 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
TABLE 9
C. gallinae, C. garei, and D. g. gallinulae im the British Museum (Natural History)
from named Bird Hosts
CORVIDAE
Corvus corone L. (*Carrion-Crow)
C. gallinae, 2 2, N. Wales, vii.30f (N. H. Joy),
nest; 7 g, 11 9, Ashton, Oundle, 1919 (N. C.
Rothschild), nestt; Ashton, Oundle, summer,
1919 (J. Stuart), nest ; 3 9, Ashton Wold, Oundle,
vi.26 (F. J. Cox), nest; < 3g, < 9, Layer Marney,
Essex, v.36 (M. Rothschild), nest ; Layer Marney,
Essex, ix.36 (M.R.), nest; 1 9, Reading, vii.31
(N. H. Joy), nest; all Rothschild Coll.
Corvus monedula L. (Jackdaw)
C. gallinae, 4 3, 2 9, Colinton, Midlothian,
24.v.06 (J. Waterston), nest, B.M. 1914.517;< 6,
< 9, Tring, Herts., summer, 1919 (N.C.R.),
nest; Tring, Herts., viii.26 (K. Jordan), nest;
1 9, Bushey Park, 2.iv.06 (A. H. Bishop) ;
3 6, 2 §, Windsor Park, 22.xi.28 (H. Donis-
thorpe), nest; 1 g, 7 2, Norfolk, 24.iv.34 (W. H.
Pollen), nest; 3 ¢, Ross-shire, 20.iii.34 (W. H.
Pollen), body; 1 g, 1 9, Northampton, 1.v.35
(W. H. Pollen), body; D. gallinulae, 5 3, 4 9,
Duncarn Hill, Fifeshire, 6.xi.o9 (H. Ashworth),
Coll.; 5 9, Woodburn House, near Dalkeith,
12.v.08 (J. Waterston (?)), nest; B.M. 1914.517;
I g, Wiltshire, 14.iv.34 (W. H. Pollen), body;
Rothschild Coll. ; 2 3, 2 9, Northampton, 1.xii.34
(W. H. Pollen), body; Rothschild Coll.; 1 3g,
10 Q, Colinton, Midlothian, 19.v.09 (J. Water-
ston), nest; 10 g, < 9, Arniston, Midlothian,
28.v.o9 (A. Bennett), nest; 8 g, < 9, Loudoun,
Ayrshire, 17.vi.og (J. Gloag), nest; all B.M.
1914.517; 7 gd, 8 9, Armitage, Staffs., 24.iii.28
(C. E. Stott), nest; 2 g, 2 9, Ince, Cheshire, 1890
(R. Newstead); < g, < 9, Ashton; Oundle,
5.vi.o6 (F. J. Cox); Ashton, Oundle, 11.v.0g
(F. J. Cox); Ashton, Oundle, v.10 (F. J. Cox),
nest; Ashton, Oundle, vii.26 (F. J. Cox), nest;
Ashton, Oundle, v.27 (F. J. Cox), nest; 2 9,
Boxmoor, Herts. (A. Piffard); < g¢, < 9, Tring,
7.vii.1goo (N.C.R.), nest; Tring, vi.or (N.C.R.),
nest; Tring, 10.vii.or (N.C.R.), nest; 1 g, 6 Q,
Netherfield, Sussex, I.vii.o7 (W. R. Butter-
field) ; 8 g, 6 9, Hastings, Sussex, 1909 (W. R.
Butterfield) ; 3 g, 1 9, Reading, Berks., 9.vi.39
(B. T. Parsons), nest; 1 3, Brockenhurst, Hants.,
28.v.06 (B. Piffard); < 3, < 9, Newport,
1.0.W., v.10 (H. G. Jeffrey), nest; Newport,
1.0.W., 1912 (H. G. Jeffrey), nest; C. garei, 1 g,
Hockley, Essex, 25.v.03 (F. W. Frohawk) ;
D. gallinulae, 1 3,12 9, Kirkwall, Orkney, 10.v.06
(J. Waterston), nest; all Rothschild Coll. ;
Kirkwall, Orkney, 8.vi.o6 (J. Waterston), nest;
1 9, Lochgelly, 25.vi.og (J. Brown); 3 g, 5 2
Arniston, Midlothian, 28.v.og (A. Bennett),
nest; all B.M. 1914.517.
nest; all Rothschild Coll.
Pica pica (L.) (Magpie)
C. gallinae, 2 9, New Forest, 5.v.98 (B. G.
Rye), nest, B.M. 98.88; 1 9, Essex, 16.1.35
(W. H. Pollen), body, Rothschild Coll.
STURNIDAE
Sturnus vulgaris L. (Starling)
C. gallinae, 3 3, 2 9, Little Roe Island, vii.1o
(J. Waterston), nest; << g, < Q, Ollaberry, vi.10
(J. Waterston), nest; Ollaberry, 16.iii.12 (J. Wa-
terston), nest; Ollaberry, vi.12 (J. Waterston),
nest; all B.M. 1914.517; 1 g, Aberdour, Fife-
shire, 20.iv.07 (J. Waterston), nest; Rothschild
FRINGILLIDAE
Coccothraustes coccothraustes (L.) (Hawfinch)
D. gallinulae, 1 3, 1 9, Eaton, nr. Chester,
5.v.o1 (R. Newstead), nest; Eaton, nr. Chester,
g.v.o1 (R. Newstead), nest; all Rothschild Coll.
* Name of host as in Witherby and not necessarily as on slides. Thus Turdus musicus becomes Turdus
evicetorum since Waterston’s publications make it clear that the Song-Thrush is the host from which he
obtained the specimens in question. Certain specimens from Corvus corone are merely marked ‘Carrion-
Crow’.
+ With regard to material in the British Museum, the registered number is appended in all cases
unless the material referred to pertains to the Rothschild Collection (1923. 615). It must be explained
that after Charles Rothschild’s death in 1923 material continued to flow in to Karl Jordan, which he
worked out and then incorporated in the Rothschild Collection. Although technically British Museum
material, the author has made no attempt to separate it from the rest, since she feels that the date is
sufficient indication of the fact that it was incorporated after the Rothschild Collection passed into the
absolute control of the Trustees.
+ When it is known for certain if specimens were collected from the body or nest of the host this in-
formation is added—whether or not it is present on the slides or tubes.
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS 219
TABLE 9—(cont.)
Chloris chloris (L.) (Greenfinch)
C. gallinae, 1 g, 1 9, Colinton, Midlothian,
I1.v.06 (J. Waterston), nest; D. gallinulae, 1 9,
Lothianburn, 26.v.06 (?); 1 g, 19, Arniston,
Midlothian, 28.v.og (A. Bennett), nest; Arnis-
ton, Midlothian, 8.vi.og (A. Bennett), nest; all
B.M. 1914.517; 2 2, Abbey St. Bathans, Berwick-
shire, 14.v.07 (J. Waterston), nest; 3 9, Tresco,
Isles of Scilly, 12.vi.o7 (F. J. Cox), nest; all
Rothschild Coll.
Carduelis carduelis (L.) (Goldfinch)
C. gallinae, < 3, < 9, Redland, Bristol, 1917
(P. Stonelake) ; C. gare, 2 g, 2 9, Fancott, nr.
Luton, i.44 (R. B. Laurence), nest; all Roths-
child Coll. (B.M.).
Carduelis cannabina (L.) (Linnet)
C. garei, < fg, < 9, Newport, 1.0.W., vi.1o
(H. G. Jeffrey), nest; Rothschild Coll. (B.M.).
Fringilla coelebs L. (Chaffinch)
C. gallinae, 1 3, Ross-shire, 16.iii.34 (W. H.
Pollen), body; Rothschild Coll.; 2 9, Arniston,
Midlothian, vioog (A. Bennett), nest; 3 Q,
Loudoun, Ayrshire, summer, 1909 (? collector) ;
D. gallinulae, 8 3, 10 9, Inchture, Perthshire,
vii.o6 (J. H. McNair), nest; Inchture, Perth-
shire, viii.o6 (J. H. McNair), nest; 1 g, Lahill,
Largo, 29.x.12 (L. J. Rintoul, E. V. Baxter),
body; all B.M. 1914.517; 8 g, 3 2, Lesmahagow,
Lanarkshire, 31.vi.o6 (J. Clark, per Watkins and
Doncaster), nest; Rothschild Coll.; 2 g, 9 9,
Loudoun, Ayrshire, vi.og (J. Gloag), nest; B.M.
1914.517.
Emberiza citrinella L. (Yellow Hammer)
C. gallinae, 1 9, Nethercraigs, nr. Paisley,
4.v.09 (J. Waterston), nest; B.M. 1914.517;
< 9, Tring, Herts., 19.vii.og (N.C.R.), nest;
Rothschild Coll. (B.M.); D. gallinulae, 1 3, 3 9,
Rilla Mill, Cornwall, vi.o5 (K. Jordan), nest;
Rothschild Coll. (B.M.).
Emberiza calandra L. (Corn Bunting)
C. garei, 1 9, Todhead, Kinneff, Kincardine-
shire, 16.vi.og (J. Waterston), body; B.M. 1914.
517.
Emberiza schoeniclus (L.) (Reed Bunting)
C. garet, 6 3, 3 9, Tring, Herts., vii.o3 (N.C.R.),
nest; Tring, Herts., vi.o6 (F. J. Cox), nest;
D. gallinulae, 1 9, Windsor Forest, Surrey,
23.vi.28 (F. W. Frohawk) ; all Rothschild Coll.
PLOCEIDAE
Passer domesticus (L.) (House-Sparrow)
C. gallinae, 3 3, 5 2, Carie, Rannoch, 24.vii.o8
(F. J. Cox), nest; Rothschild Coll. (B.M.);
<4, < 9, Mull, Argyllshire, 28.viii.r2 (L. G.
Esson) ; Rothschild Coll. (B.M.) ; 8 3,9 9, Wilber-
lea, St. Andrews, Fife, 18.viii.o8 (J. Waterston),
nest; 3 9, Auchinblae, Kincardineshire, 27.v.09
(J. Campbell); 4 g, 10 9, Kennetpans, Clack-
mannan, 8-10.viii.o8 (J. Waterston), nest; 1 4,
Gorebridge, nr. Dalkeith, 17.vii.o6 (J. Water-
ston), nest; 8 g, < 9, Colinton, Midlothian,
7.v.06 (J. Waterston), nest; Colinton, Mid-
lothian, 11.v.06 (J. Waterston), nest; 2 gf, 6 9,
Dunlaverock, Coldingham, viii.og (J. F. Cor-
mack) ; Dunlaverock, Coldingham, 7.ix.og (J. F.
Cormack); Dunlaverock, Coldingham, ix.og
(J. F. Cormack) ; 1 3, 1 9, Dunlaverock, Colding-
ham, 2.viii.1o (J. F. Cormack), nest; < g, < 9,
Coldingham, Berwickshire, 25.ix.o8 (J. F. Cor-
mack), nest; Coldingham, Berwickshire, viii.o9
(J. F. Cormack), nest; all B.M. 1914.517; 1 9,
Armitage, Staffordshire, x.31 (L. C. E. Stott) ;
4 6, I 9, Penarth, Glamorgan, 20.ix.28 (H. M.
Hallett); 2 ¢, 3 9, Whipsnade, Bedfordshire,
nest; < g, < 9, Tring, Herts., 1901 (N.C.R.),
nest; Tring, Herts., vi.o6 (K. Jordan), nest;
Tring, Herts., ix.1o (K. Jordan), nest; < 4,
< 9, London, xi.og (N.C.R.), nest; all Roths-
child Coll.; 2 g, 5 9, Woking, Surrey; B.M.
1910.266; 2 9, Pylewell Park, Lymington,
Hants, 1906 (R. Newstead); < g, < 9, New-
port, I.0.W., v.1o (H. G. Jeffrey), nest; New-
port, I.0.W., 1912 (H. G. Jeffrey), nest; all
Rothschild Coll.; D. gallinulae, 1 3, Arniston,
Midlothian, 28.v.o9 (A. Bennett) ; B.M.1914.517;
4 6, 2 9, Longwood, Southwick, Sunderland,
25.vi.10 (F. Palin); 3 g, 2 9, Park Mill, Gower
Peninsula, Glamorgan, 23.v.19 (F. J. Cox), nest;
all Rothschild Coll.
ALAUDIDAE
Alauda arvensis L. (Skylark)
C. gallinae, 1 3, Burwick, Orkney, 11.vi.06
(J. Waterston), nest; Rothschild Coll. (B.M.) ;
r 9, Aberlady, 13.vi.o6 (R. Cochrane), nest;
B.M. 1914.517; C. garei, 11 g, 3 9, Wallasey
sandhills, Liverpool, 18.vi.o6 (Watkins and
Doncaster) ; Rothschild Coll.; D. gallinulae, 1 Q,
Ollaberry, 8.vii.12 (J. Waterston), body; 1 9,
East of Silverside, Ollaberry, 10.vi.10 (J. Water-
ston), body; 1 g, 6 9, Tankerness, nr. Kirkwall,
Orkney, 8.vi.o6 (J. Waterston), nest; 1 Q,
Berwick, 11.vi.o6 (? collector), nest; all B.M.
I914.517.
220 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
TABLE 9—(cont.)
MOTACILLIDAE
Anthus pratensis (L.) (Meadow-Pipit)
C. gallinae, 1 9, Cowdenbeath, Fifeshire,
10.vi.o7 (J. Waterston), nest; B.M. 1914.517;
C. garvei, 1 9, Lochhead, Lochgelly, 21.x.09
(J. Brown), body; B.M. 1914.517; < 3, < Q,
Newport, I.0.W., ix.10 (H. G. Jeffrey), nest;
1 g, Foulness Island, 28.v.34 (W. H. Pollen),
body; D. gallinulae, 1 9, Walls, Shetland, vi.o6
(J. Clark, per Watkins and Doncaster), nest;
3 6, 7 &, Braid Hills, nr. Edinburgh, 2.vi.o6
(J. Waterston), nest; 1 9, Lesmahagow, Lanark-
shire, 22.vii.o6 (J. Clark, per Watkins and Don-
caster), nest; all Rothschild Coll.
Anthus spinoletta (L.) (Rock-Pipit)
C. gallinae, 1 9, Ollaberry, Shetland, 18.x.10
(J. Waterston), body; D. gallinulae, 1 g, 1 9,
nr. Haylor, Ronas Voe, Cumberland, 13.vii.12,
nest; all B.M. 1914.517.
Motacilla alba yarrelli Gould (Pied Wagtail)
C. gallinae, 5 3, 6 9. Kennetpans, Clack-
mannan, 6.viii.o3 (J. Waterston), nest; 3 g, 8 9,
Arniston, Midlothian, 8.vi.og (A. Bennett), nest ;
all B.M. 1914.517; 2 6, Westfield, Sussex,
2.viii.o7 (W. R. Butterfield) ; C. garei, 3 g, < Q,
Tring, Herts., 1902 (N.C.R.), nest ; Tring, Herts.,
vii.o3 (N.C.R.), nest; all Rothschild Coll.; D.
gallinulae, 2 3, 7 2, Hillend, Fife, 26.v.06 (J.
Waterston), nest; 5 9, nr. Loganlea Waterfall,
25.v.07 (R. Cochrane), nest; 1 4, Arniston,
Midlothian, 28.vi.og (A. Bennett), nest; all B.M.
I1914.517; 10 g, 5 9, Lesmahagow, Lanarkshire,
20.vi.06 (J. Clark, per Watkins and Doncaster),
nest; I ¢, Ross-shire, 20.iii.34 (W. H. Pollen),
body; all Rothschild Coll.
Motacilla cinerea cinerea Tunst. (Grey Wagtail)
D. gallinulae, 1 3, Arniston, Midlothian, 29.vi.og
(J. Waterston), nest; B.M. 1914.517; 2 9, Coal-
burn, Lanarkshire, vi.o6 (J. Clark), nest; Roths-
child Coll. (B.M.).
CERTHIIDAE
Certhia familiaris (L.) (Tree-Creeper)
D. gallinulae, 1 9, Howick, Northumberland,
1.xi.34 (W. H. Pollen), body; Rothschild Coll.
(B.M.).
SITTIDAE
Sitta europoea L. (Nuthatch)
C. gallinae, 6 3, 6 9, Hastings, Sussex, 1909
(W. R. Butterfield) ; Rothschild Coll. (B.M.).
PARIDAE
Parus major L. (Great Tit)
C. gallinae, < 3, < 9, Arniston, Midlothian,
4.vi.og (A. Bennett), nest; B.M. 1914.517; 1 9,
Ashbourne, Derbyshire (F. C. R. Jourdain) ; 2 g,
4 , Kingsland, Herefordshire, 19.v.(? year) (R.
Williams) ; < g, < 9, Tring, Herts., ix.21 (F. J.
Cox), nest; 3 g, < 9, Chingford, London, 1910
(C. Nicholson) ; 1 9, Surrey, 13.vi.og (H. Russell) ;
<<, <9, Shere, Guildford, 13.vio9; < g,
< 9, Sussex, 22.iv.08 (W. R. Butterfield); all
Rothschild Coll.; 1 9, Perch and Nagshead,
Gloucester, 28/29.vii.45 (received from Com.
Inst. Ent. per Dr. K. Jordan), nest; B.M. 1914.
517; <3, < 9, Wembdon, nr. Bridgwater,
1902 (G. Gare); Wembdon, nr. Bridgwater, no
date (G. Gare) ; 2 g, Ross-shire, 20.iii.34 (W. H.
Pollen), body; < ¢, < 9, Bere Regis, Dorset,
I5.vii.19 (H. Russell); < 3, < 9, Dorset, vi.14
(H. Russell) ; all Rothschild Coll.; D. gallinulae,
2 3, 2 3, Cowdenbeath, 10.vi.o7 (J. Waterston),
nest; 1 g, 1 9, Aberdour, Fife, v.o7 (J. Water-
ston); 1 g, Arniston, Midlothian, 4.vi.og (A.
Bennett), nest; all B.M. 1914.517.
Parus coeruleus L. (Blue-Tit)
C. gallinae, 8 3, < 9, Kennetpans, Clack-
mannan, 8-10.viii.o8 (J. Waterston), nest;
< 6, < Q, Arniston, Midlothian, vi.og (A.
Bennett), nest; all B.M. 1914.517; < 3, < 9,
Rosehill, Cheadle, Staffordshire, xii.13 (J. R. B.
Masefield); < 4, < 9, Cardiff, 12.vi.33
(H. M. Hallett), nest; < ¢, < 9, Ashton,
Oundle, ix.oz (N.C.R.), nest; Ashton, Oundle,
vi.1o (J. Stuart), nest; 1 9, Hockley, Essex,
18.v.03 (F. W. Frohawk); < 3,< 9, London,
xi.og (N.C.R.), nest; 1 g, 3 9, Chiswick, ii.or
(A. Sich); 8 g, 6 9, Hastings, Sussex, 1909
(W. R. Butterfield); < g, < 9, Reading,
Berks., 9.vi.39 (B. T. Parsons), nest; 1 ,
I 9, Rilla Mill, Cornwall, vio5 (K. Jordan),
nest; D. gallinulae, 1 9, Ross-shire, 20.ii1.34
(W. H. Pollen), body; 3 ¢g, 2 9, Crickleigh,
Gloucester, 24.iv.04 (R. Newstead) ; all Roths-
child Coll.
Aegithalos caudatus (L.) (Long-tailed Tit)
D. gallinulae, 5 3, Essex, 1910 (H. Russell) ;
Rothschild Coll.
Panurus biarmicus (L.) (Bearded Tit)
C. garei, < 3, < 9, Hickling Broad, Norfolk,
8.v.16 (H. Russell), nest; 6 g, 4 9, Horeton
Broad, Cromer, vii.1896 (F. H. Barclay), both
Rothschild Coll.
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS 221
TABLE 9—(cont.)
MUSCICAPIDAE
Muscicapa striata (Pall.) (Spotted Flycatcher)
C. gallinae, t 9, Kinneff, Kincardineshire,
15.vi.og (J. F. Cormack), nest; 2 2, Gorebridge,
Dalkeith, 12.vii.o6 (? collector), nest; 9 9,Arnis-
ton, Midlothian, 3.viii.og (A. Bennett) ; D. galli-
nulae, 5 9, Kinneff, Kincardineshire, 5.vi.og
(J. F. Cormack), nest ; 3 9, Gorebridge, 12.vii.o6
(J. Waterston), nest ; all B.M. 1914.517; 1 9, Les-
mahagow, Lanarkshire, 5.viii.o6 (J. Clark, per
Watkins and Doncaster), nest; 1 g, 1 9, Tring,
Herts., vii.tg (K. Jordan), nest; 1 3g, 1 Q,
Newport, I.0.W., ix.1goo (H. G. Jeffrey), nest;
all Rothschild Coll.
SYLVIIDAE
Phylloscopus collybita (Vieill.) (Chiff-Chaff)
C. gallinae, 2 3, 1 9, Devon, vi.35 (G. M.
Spooner), nest; D. gallinulae, 8 3, < 9, Lesma-
hagow, Lanarkshire, vii.o6 (J. Clark, per Wat-
kins and Doncaster), nest; 3 9, Reigate, Surrey,
12.vi.or (C. Reeves) ; all Rothschild Coll.
Phylloscopus trochilus (L.) (Willow-Warbler)
D. gallinulae, 2 3, 3 2, Glencorse Pond, 3.vii.o7
(R. Cochrane), nest; 1 g, 3 9, Arniston, Mid-
lothian, 3-4.vi.og (A. Bennett), nest; all B.M.
914.5175 °< ¢, < 9, Tring; Herts:, vi.28(K.
Jordan), nest; < g, < 9, Tring, Herts., vii.28
(K. Jordan), nest; all Rothschild Coll.
Phylloscopus sibilatrix (Bechst.) (Wood-Warbler)
D. gallinulae, << 3, < 9, Arniston, Midlothian,
vi.og (A. Bennett), nest; B.M. 1914.517.
Acrocephalus schoenobaenus (L.) (Sedge-Warbler)
D. gallinulae, 2 g, Tring, Herts., v.34 (L.
Goodson), nest; Rothschild Coll.
Sylvia borin (Bodd.) (Garden Warbler)
D. gallinulae, 3 3, 4 9, Arniston, Midlothian,
25.vi.og (A. Bennett), nest; B.M. 1914.517.
Sylvia communis Lath. (Whitethroat)
D. gallinulae, 1 9, Skokholm Bird Observa-
tory (R. M. Lockley), body; Rothschild Coll.
Sylvia curruca (L.) (Lesser Whitethroat)
D. gallinulae, 9 3, 10 , Tansor Wold, Oundle,
v.06 (F. J. Cox), nest; Rothschild Coll.
TURDIDAE
Turdus viscivorus L. (Mistle Thrush)
C. gallinae, 1 3, 1 9, Colinton, Midlothian,
24.v.06 (J. Waterston) ; D. gallinulae, 1 g, 6 9,
Hillend, Fife, 25.v.08 (R. Cochrane), nest;
2 2, Loganlea Waterfall, 25.v.07 (R. Cochrane),
nest ; all B.M. 1914.517.
Turdus ericetorum Turton (Song-Thrush)
C. gallinae, 1 g, Colinton, Midlothian, 7.v.06
(J. Waterston), nest; B.M. 1914.517; I Q,
Nethercraigs, Paisley, 4.v.0o9 (J. Waterston),
joint nest with T. merula; B.M. 1914.517; < 6,
< 9, Loch Ranza, Arran Isle, N.B., vi.27 (F. J.
Cox), nests; 2 9, Tring, Herts., 10.xi.14 (E.
Hartert); all Rothschild Coll.; D. gallinulae,
2 4, Kinneff, Kincardineshire, 14.vi.og (J. Water-
ston), body; 1 g, i Q, Colinton, Midlothian,
7.v.06 (J; Waterston), nest; 1 gf, 3 9, Nether-
craigs, Paisley, 4.v.09, joint nest with T. me-
rula; all B.M. 1914.517; I 4, 1 9, Crickleigh,
Gloucester, 20.iv.06; Rothschild Coll.
Turdus merula L. (Blackbird)
C. gallinae, 1 9. St. Margaret’s Hope, South
Ronaldshay, Orkney, vi.o6 (J. Waterston), nest ;
B.M. 1914.517; 2 6, 4 §, Carie, Rannoch,
23.vii.o8 (F. J. Cox), nest; Rothschild Coll. ;
1 9, Woodburn House, nr. Dalkeith, 12.v.08
nest; I g, 3 9, Colinton, Midlothian, 15.iv.07
(J. Waterston), nests; Colinton, Midlothian,
26.vii.og (J. Waterston), body; 1 9, Nether-
craigs, Paisley, 4.v.09 (J. Waterston), joint nest
with T. ericetorum; all B.M. 1914.517; 1 Q,
Armitage, Staffordshire, i.33 (L. C. E. Stott),
nest; 5 9, Tring, Herts., vii.1g00 (N.C.R.), nest;
Tring, Herts., 15.vii.11 (H. Mugford); 4 3, 7 9,
Reigate, Surrey, 12.vi.18 (C. E. Stott); < ¢,
< 9, Newport, I.0.W., v.10 (H. G. Jeffrey),
nest; Newport, I.0.W., ix.1o (H. G. Jeffrey),
nest; < 9, St. Mary’s, Scilly Isles, 11.vi.o7
(F. J. Cox), nest; all Rothschild Coll. ; C. garez,
1 g, Park House, Crathorne, Yarm-on-Tees,
Yorks. (? collector), nest; B.M. 1920.166 (pre-
sented by S. Hirst) ; 1 g, Newport, I.0.W., iv.1o
(H. G. Jeffrey) ; D. gallinulae, 3 3, 3 2, Kilronan,
Arran Is., Ireland, vii.o6 (M. Mullin, per Watkins
and Doncaster); 1 g, 1 9, Hillend, Fife, 18.v.06
(J. Waterston), nest; all Rothschild Coll. ;
4 6, < Q, Flotterstone, Fifeshire, 13.v.06 (J.
Waterston), nest; 4 ¢, 5 9, Aberdour, Fife,
20.iv.07 (J. Waterston), nest; 1 9, Burntisland,
8.x.09 (Rettie); 2 g, 1 9, St. Margaret’s, nr.
Queensferry, 9.vi.og (W. H. Elder), nest; all
B.M. 1914.517; 3 6, 8 Q, Colinton, nr, Edin-
burgh, 24.v.06 (J. Waterston); Colinton, nr.
Edinburgh, 25.v.06 (J. Waterston); all Roths-
child Coll.; 4 g, 2 9, Arniston, Midlothian,
3.vi.og (A. Bennett), nest ; Nethercraigs, Paisley,
4.V.09, joint nest with T. musicus (= ericetorum) ;
all B.M. 1914.517; 1 g, 1 9, Tring, Herts.,
222 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
TABLE 9—(cont.)
15.vii.11 (H. Mugford); 1 ¢, Newport, I1.0.W.,
iv.1o (H. G. Jeffrey), nest; all Rothschild Coll.
Oenanthe oenanthe (L.) (Wheatear)
D. gallinulae, 1 9, Gluss Voe, west side, Olla-
berry, 28.iv.13 (J. Waterston), body; B.M.
1914.517.
Saxicola torquata (L.) (Stonechat)
D. gallinulae, < 3, < 9, Loudoun Estate, Ayr-
shire, 12.vi.og (J. Gloag), nest; B.M. 1914.517.
Phoenicurus phoenicurus (L.) (Redstart)
C. gallinae, 1 9, Gorebridge, nr. Dalkeith
(? collector), nest; < g, < 9, Arniston, Mid-
lothian, 25.vi.og (? collector), nest; D. gallinulae,
9 6, < 9, Arniston, Midlothian, I.vii.o6 (A.
Bennett), nest; Arniston, Midlothian, 28.v.o9
(A. Bennett), nest; all B.M. 1914.517.
Erithacus rubecula (L.) (Robin)
C. gallinae, 1 3, Auchinblae, Kincardineshire,
17.iv.09 (J. Duffus), nest; D. gallinulae, 1 4,
1 9, Auchinblae, Kincardineshire, 17.iv.09 (J.
Duffus), nest ; all B.M. 1914.517; 3 4, 3 9, Hillend,
Fife, 29.v.06 (? collector), nest ; 3 9, Flotterstone,
Fifeshire, 29.v.06; 5 g, 2 9, Colinton, Midlothian,
vi.o6 (J. Waterston), nest; all B.M. 1914.517;
2g, 8 9, Carie, Rannoch, N.B., vii.o8 (F. J. Cox),
nest; 1 9, Banbury, 4.xii.34 (W. H. Pollen), body ;
2 4, < 9, Loch Ranza, Arran Isle, vii.27 (F. J.
Cox), nests; < g, < 9, nr. Park Mill, Gower
Peninsula, Glamorgan, v.19 (F. J. Cox), nest;
<6, <Q, Tring, Herts., 7.viiior (N.C.R.),
nest; Tring, Herts., 3.ix.o1 (N.C.R.), nest ; Tring,
Herts., 17.ix.01 (N.C.R.), nest; Tring, Herts.,
5.x.01 (N.C.R.), nest; 1 g, 2 9, Newport, I.0.W.,
ix.1o (H. G. Jeffrey), nest; Newport, I.0.W.,
1912 (H. G. Jeffrey), nest ; all Rothschild Coll.
PRUNELLIDAE
Prunella modularis (L.) (Hedge-Sparrow)
C. gallinae, 1 9, Quixewood, Berwickshire,
14.v.07 (J. Waterston), nest; 5 9, Elveden,
Suffolk, 14.viii.o7 (F. J. Cox), nest; 1 9, Tring,
Herts., 1901 (N.C.R.), nest; 4 3, 1 9, Crickleigh,
Gloucester, 21.iv.06 (R. Newstead) ; 1 9, North-
ampton, 1.xii.34 (W. H. Pollen), body; all
Rothschild Coll. ; C. garei, 1 3, Newport, I.0.W.,
v.10 (H. G. Jeffrey) ; Rothschild Coll.; D. galli-
nulae, 4 3, 9 Y, Hillend, Fife, 26.v.06 (J. Water-
ston), nest; 2 g, 3 9, Edinburgh, 12.v.08 (J.
Waterston), nest; 5 g, 10 9, Colinton, Mid-
lothian, 13.v.06 (J. Waterston) ; Colinton, Mid-
lothian, 24.v.06 (J. Waterston); Colinton,
Midlothian, 19.v.og (J. Waterston); Colinton,
Midlothian, 31.vi.o9 (J. Waterston), body; all
B.M. 1914.517; 1 9, Dumbarton, Scotland,
21.ix.34 (W. H. Pollen), body; Rothschild Coll. ;
33,4 9, Arniston, Midlothian, vi.og (A. Bennett),
nest; B.M. 1914.517; < 6, < 9, Talacre, nr.
Prestatyn, 16.v.03 (R. Newstead) ; 1 9, Newport,
1.0.W., v.10 (H. G. Jeffrey), nest ; all Rothschild
Coll.
TROGLODYTIDAE
Troglodytes troglodytes (L.) (Wren)
C. gallinae, 1 9, Lamba, Shetland, 25.ix.12
(J. Waterston), nest; B.M. 1914.517; 6 g, 7 9,
Ashton, Oundle, vi.o6 (F. J. Cox), nest; 2 4,
< 9, Tring, Herts., 5.x.o1 (N.C.R.), nest; D.
gallinulae, 2 3, 1 9, St. Kilda, 1900 (W. Eagle
Clarke); all Rothschild Coll.; 1 9, Gluss Voe,
Ollaberry, 20.ix.10 (J. Waterston), body; 4 3,
< 9, between Hillend and Flotterstone, Fife-
shire, 29.v.06 (J. Waterston), nest; 2 g, 3 Q, nr.
Loganlea Waterfall, 25.v.07 (J. Waterston), nest;
all B.M. 1914.517; 1 9, Lesmahagow, Lanark-
shire, 5.viii.o6 (J. Clark, per Watkins and Don-
caster), nest; Rothschild Coll.; 1 ¢, Pettadale
Water, nr. R. Roe, 24.iv.12 (J. Waterston),
body; B.M. 1914.517; 1 g, 1 9, Cardiff, 14.vi.31
(H. M. Hallett, A. H. Salmon); 2 9, Tring,
Herts., vi.or (N.C.R.), nest; all Rothschild Coll.
CINCLIDAE
Cinclus cinclus (L.) (Dipper)
C. gallinae, 1 2, Auchinblae, Kincardineshire,
14.iv.og (L. Duffus), nest; 1 g, Abbey St. Ba-
thans, Berwickshire, 26.v.07; all B.M. 1914.517;
I g, 1 9, Rothbury, Sunderland, vii.o6 (per
Watkins and Doncaster), Rothschild Coll.; D.
gallinulae, 9 3, < 9, Torduff Reservoir, Mid-
lothian, 12.v.o9 (J. Waterston), nest; B.M. 1914.
517; 1 9, nr. Granthouse, 14.v.07 (J. Waterston),
nest; B.M. 1914.517; 1 9, Nantyffrith, nr. Min-
era, Flintshire, 5.v.or (R. Newstead); Roths-
child Coll.
HIRUNDINIDAE
Hirundo rustica L. (Swallow)
C. gallinae, 2 3, 7 2, Dirleton, East Lothian,
2.ix.og9 (J. Gardiner), nest; 3 g, 7 9, Howletburn,
Galston, Ayrshire, 4.ix.10 (J. Gloag), nest; all
B.M. 1914.517; 8 3, < 9, Newport, 1.0.W.,
1912 (H. G. Jeffrey), nest; Newport, I.0.W.,
1914 (H. G. Jeffrey), nest; all Rothschild Coll.
Delichon urbica (L.) (House-Martin)
C. gallinae, 7 3, 5 Q, Mull, Argyllshire, ix.12
(L. G. Esson); Mull, Argyllshire, 7.i.13 (L. G.
Esson) ; all Rothschild Coll. ; 2 3, 4 9, Gladhouse,
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
223
‘TABLE 9—(cont.)
Midlothian, 13.viiiio6 (J. Waterston), nest;
B.M. 1914.517; < 3, < 9, Dunlaverock, Ber-
wickshire, ix.o6 (J. Waterston), nest ; Rothschild
Coll.; 9 3g, 10 9, Dunlaverock, Coldingham,
24.ix.06 (J. F. Cormack), nest; Dunlaverock,
Coldingham, viii.og (J. F. Cormack), nest;
Dunlaverock, Coldingham, ix.og (J. F. Cor-
mack), nest; 1 g, 12, Dunlaverock, Coldingham,
2.viii.1o (J. F. Cormack); all B.M. 1914.517;
10 g, 6 9, Ashton, Oundle, viii.o6 (F. J. Cox),
nest ; Rothschild Coll.
Riparia riparia (L.) (Sand-Martin)
C. gallinae, 5 3, Reigate, Surrey, ix.25 (L.C. E.
Stott) ; Rothschild Coll.
APODIDAE
Apus apus (L.) (Swift)
C. gallinae, 3 g, 8 2, Newport, I.0.W., 1913
(H. G. Jeffrey), nest; 2 g, 5 9, Wembdon, nr.
Bridgwater, 3.vii.or (G. Gare); Wembdon, nr.
_ Bridgwater, 1902 (G. Gare) ; all Rothschild Coll.
PICIDAE
Picus viridis L. (Green Woodpecker)
C. gallinae, 1 2, Northampton, 25.v.34 (W. H.
Pollen), body; Rothschild Coll.
Jynx torquilla (L.) (Wryneck)
C. gallinae, < 3, < 9, Wembdon, nr. Bridg-
water, 1902 (G. Gare); C. garei, 1 9, Wembdon,
nr. Bridgwater, 1902 (G. Gare); all Rothschild
Coll.
STRIGIDAE |
Asio otus (L.) (Long-eared Owl)
C. gallinae, < 3, < 2, Newport, I.0.W., ix.1o
(H. G. Jeffrey), nest; Newport, I.0.W., 1912
(H. G. Jeffrey), nest; all Rothschild Coll.
Tyto alba (Scop.) (Barn-Owl)
C. gallinae, 2 3, 4 9, Ashton Wold, Oundle,
vi.26 (F. J. Cox), nest; < g, < Q, Tring, Herts.,
summer, 1919 (F. J. Cox), nest; 2 ¢, Reigate,
Surrey, 1I0.viii.tg0o (C. Reeves); 9 g, 10 Q,
Hastings, Sussex, 1909 (W. R. Butterfield) ; all
Rothschild Coll.
Strix aluco L. (Tawny Owl)
C, gallinae, < g, < 9, Arniston, Midlothian,
iv.og (A. Bennett), nest; Arniston, Midlothian,
vi.og (A. Bennett), nest; all B.M. 1914.517;
1 g, 1 2, Upton, nr. Chester, 25.v.05 ; 4 9, Tring,
Herts. (M.R.), nest; 2 g, 7 9, Tring, Herts.,
1936 (M.R.); 1 3, Northampton, 21.iv.34 (W. H.
Pollen), body; all Rothschild Coll.
Athene noctua (Scop.) (Little Owl)
C. gallinae, 1 9, Northamptonshire, 22.v.34
(W. H. Pollen), body; 5 g, 4 2, Northampton-
shire, 12.iv.34 (W. H. Pollen), body; all Roth-
schild Coll.
FALCONIDAE
Accipiter nisus (L.) (Sparrow-Hawk)
C. gallinae, 1 3, Bradford, Yorks., viii.29
(N. H. Joy), nest; < g, 2 9, Tring, Herts.,
summer, 1919 (N.C.R.), nest; < g, < 9, New-
port, I.O.W., 1912 (H. G. Jeffrey), nest; 1 9,
Norfolk, 18.iv.34 (W. H. Pollen), body; D. galli-
nulae, 1 6, Beaconsfield, Buckinghamshire,
16.vii.34 (W. H. Pollen), body; all Rothschild
Coll.
ANATIDAE
Anas platyrhyncha L. (Mallard)
C. garei, 2 3, 3 2, Lochgelly, Fifeshire, 25.v.o9
(J. Brown), nest; B.M. 1914.517 < g, < 9,
Tring, Herts., vii.o3 (F. J. Cox), nest; Tring,
Herts., v.o5 (N.C.R.), nest; all Rothschild Coll.
Anas acuta L. (Pintail)
C. garei, 6 3, 3 2, Orkney Islands, vi.11 (N. H.
Joy), nest; Rothschild Coll.
Aythya fuligula (L.) (Tufted Duck)
C. garei, < 3,< 9, Merton, Oxford, vi. (? date)
(Lord Walsingham) ; 1 9, Thetford, Norfolk, 2.vi.
(? date) (per British Museum) ; all Rothschild Coll.
Somateria mollissima (L.) (Common Eider)
C. garei, 4 3, Orkney Islands, vi.11 (N. H.
Joy), nest; Rothschild Coll.; 1 g, 1 9, Tanker-
ness, Pomona, 8.vi.o6 (J. Waterston), nest;
B.M. 1914.517; 8 3, < 9, Bass Rock, Firth of
Forth, vii.26 (F. J. Cox), nest; Rothschild Coll.
Melanitta nigra (L.) (Common Scoter)
C. garei, < 3, < Q, Little Roe, Shetland,
vii.1o (J. Waterston), nest; Rothschild Coll.
Mergus serrator L. (Red-breasted Merganser)
C. gavei, 1 9, Belfast, Ireland, 2.vi.34 (W. H.
Pollen), body; Rothschild Coll.
PHALACROCORACIDAE
Phalacrocorax aristotelis (L.) (Shag)
C. gareit, < 3, < 9, Lamba Island, Shetland,
Vii.12, nest; << g, < 9, Lamba Island, Shetland,
xii.z3, nest; all B.M. 1914.517; <¢, < 9,
Orkney Islands, viir (N. H. Joy), nest;
Rothschild Coll.
224 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
TABLE 9—(cont.)
HYDROBATIDAE
Hydrobates pelagicus (L.) (Storm-Petrel)
C. gallinae, 1 3, Borough of Copister, SE. end
of Yell, opposite Mossbank, Shetland, 25.vii.13
(J. Waterston and Captain Jensen), nest; B.M.
I1914.517.
Fulmarus glacialis (L.) (Fulmar Petrel)
C. gallinae, 1 9, Ross-shire, 5.1.35 (W. H.
Pollen), body; Rothschild Coll.; D. gallinulae,
2 g, Uyea Island, Shetland, 24.iv.12 (J. Water-
ston), body; Uyea Island, Shetland, vi.12 (J.
Waterston), near nest; all B.M. 1914.517.
COLUMBIDAE
Columba palumbus L. (Wood-Pigeon)
C. gallinae, 6 9, Berkshire, 11.iv.34 (W. H.
Pollen), body ; Rothschild Coll.
Columba oenas L. (Stock-Dove)
C. gallinae, 1 2 Colinton, Midlothian, 19.v.09
(J. Waterston), nest; Rothschild Coll.
Columba livia Gm. (Rock-Dove)
C. gallinae, < 3, < Q, Little Roe, Yell Sound,
Shetland, 17.vii.11 (J. Waterston), nest; 1 9,
Caves in cliffs, Todhead, Kinneff, 15.vi.og (J.
Waterston), nest ; all B.M. 1914.517; 1 9, Belfast,
Ireland, 24.v.35 (W. H. Pollen), body; C. garez,
4 6, < Q, Little Roe, Shetland, 17.vii.11 (J.
Waterston), nests; D. gallinulae, 4 g, 5 9, ina
cave NE. end of Little Roe, Shetland, 17.vii.11
(J. Waterston), nest; all Rothschild Coll.
SCOLOPACIDAE
Scolopax rusticola L. (Woodcock)
D. gallinulae, 8 3, < 9, Aberdour, Fifeshire,
25.v.07 (J. Waterston), nest; B.M. 1914.517.
Capella gallinago (L.) (Snipe)
C. garei, 1 9, Northmavine, Shetland (R. H.
McNair); 5 9, Tankerness, Orkney, 8.vi.o6
(J. Waterston), nest; < g, < 9, Lochgelly,
Fifeshire, 25.v.09 (J. Brown), nest; all B.M.
1914.517.
Calidris alpina (L.) (Dunlin)
C. garei, < g, < 9, Tinga Skerry, Yell Sound,
Shetland, 11.vii.12 (J. Waterston), nest; B.M.
1914.517.
Tringa totanus (L.) (Redshank)
C. garei, 1 3, 2 9, Lochgelly, Fifeshire, 25.v.09
(J. Brown), nest; B.M. 1914.517.
Charadrius hiaticula L. (Ringed Plover)
C. garei, 1 9, Ollaberry, Shetland, 22.ix.10
(J. Waterston), body; 1 9, Aberlady, nr. Edin-
burgh, 22.v.09 (J. F. Cormack), nest; all B.M.
1914.517.
Vanellus vanellus (L.) (Lapwing)
C. garei, 2 3, Tents Moor, Fifeshire, 12.vi.og
(J. F. Cormack), nest; 4g, 5 2, Lochhead, Loch-
gelly, Fifeshire, 25.iii.o9 (J. Brown), nest; 5 dg,
3 9, Aberlady, nr. Edinburgh, 22.v.o9 (J. F.
Cormack), nests; Aberlady, nr. Edinburgh,
10.v.10 (J. F. Cormack), nest; all B.M. 1914.517.
LARIDAE
Larus ridibundus L. (Black-headed Gull)
C. gavei, 1 9, Orkney Islands, vi.r1 (N. H.
Joy), nest; Rothschild Coll.
Larus canus L. (Common Gull)
C. garei, < g, < , Little Roe, Shetland, v.11
(J. Waterston), nest; B.M. 1914.517.
Larus fuscus L. (Lesser Black-backed Gull)
C. garei, 3g, 792, Walls, Shetland, viii.o6
(per Watkins and Doncaster), nest; Rothschild
Coll.
STERCORARIIDAE
Stercorarius parasiticus (L.) (Arctic Skua)
C. garei, < g, < 9, Hascussay, Shetland,
7.vii.11 (M. Meade-Waldo) ; Rothschild Coll.
Crex crex (L.) (Corn-Crake)
C. garei, < 3, < 9, found by taxidermist in
Nat. Hist. Mus., London; body; Rothschild
Coll.
RALLIDAE
Gallinula chloropus (L.) (Moorhen)
C. gallinae, 1 3, 5 9, Ashton, Oundle, v.06
(F. J. Cox), nest; Rothschild Coll.; C. garei,
< 4g, 9 9, Arniston, Midlothian, 26.vii.o6 (A.
Bennett), nest; 1 g, 3 9, on the Eye, Grant-
house, Berwickshire, 14.v.07, nest; all B.M.
1914.517; 2 9, Ashton, Oundle, vi.o6 (F. J. Cox),
nest; 2 g, 5 9, Tring, Herts., vii.oz (N.C.R.),
nest; all Rothschild Coll.; D. gallinulae, 5 9,
Glencorse Pond, 29.v.06 (J. Waterston), nest;
I g, 4 Q, Arniston, Midlothian, 29.vi.og (J. Wa-
terston), nest; all B.M. 1914.517; 2 g, Ashton,
Oundle, v.06 (F. J. Cox), nest; Rothschild Coll.
TETRAONIDAE
Lagopus scoticus (Lath.) (Red Grouse)
C. garei, 2 3, 6 9, Tankerness, Orkney, 8.vi.06
(J. Waterston), nest; B.M. 1914.517; 1 g, IQ,
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS 225
TABLE 9—(cont.)
Perth, Scotland, 2.vii.o7 (A. E. Shipley); D.
gallinulae, 1 9, Tankerness, nr. Stromness, Kirk-
wall, Orkney, 4.vi.o6 (J. Waterston), nest; 1 4,
(A. E. Shipley) ; all Rothschild Coll.
PHASIANIDAE
Phasianus colchicus L. (Pheasant)
C. gallinae, t 3, 1 2, Cowdenbeath, Fifeshire,
10.vi.o7 (J. Waterston), nest; B.M. 1914.517;
3 9, Windsor Forest, v.30 (H. Donisthorpe),
nest; Rothschild Coll.; C. garei, 2 3, 5 9, Loch-
gelly, Fifeshire, 25.v.09 (J. Brown), nest; 1 9,
Arniston, Midlothian, 28.v.o9 (A. Bennett), nest ;
I 4,1 9, Aberlady, nr. Edinburgh, 22.v.09 (J. F.
Cormack), nest; all B.M. 1914.517; 1 9, Tring,
Herts., summer, 1919 (F. J. Cox), nest; 1 6,
t 9, Windsor Forest, v.30 (H. Donisthorpe),
nest ; D. gallinulae, 1 9, Ross-shire, 4.1.35 (W. H.
Pollen), body; 1 9, Aberdour, Fifeshire, 25.v.07
(J. Waterston), nest; all Rothschild Coll.
<3, <Q, Ashton Wold, Oundle, vi.1g9 (J.
Stuart), nest;9g, << 9, Newport, I.0O.W., ix.1g00
(H. G. Jeffrey) ; all Rothschild Coll.
Alectoris rufa (L.) (Red-legged Partridge)
C. garei, < g, < Q, Tring, Herts., summer,
1gi1 (F. J. Cox), nest; Rothschild Coll.
DOMESTIC POULTRY
Gallus domesticus L. (Fowl)
C. gallinae, 1 g, 3 9, Kinneff, Kincardine-
shire, 14.vi.og (J. R. Frazer), nest; 2 g, 6 9,
Dirleton, East Lothian, 2.ix.og (J. Gardiner),
nest; 1 9, Colinton, Midlothian, 24.v.06 (J. Wa-
terston), nest; 5 g, 9 9, Biggar, Lanarkshire,
10.ix.09, nest; all B.M. 1914.517; 1 9, Bucking-
ham, 29.v.or (L. W. Rothschild); < g, 2 Q,
Tring, Herts., 7.xii.1899 (N.C.R.), nest; Tring,
Herts., 1901 (N.C.R.), nest; all Rothschild Coll. ;
6 6, < 9, Woking, Surrey; B.M. 1910.266
(Edward Saunders Coll.); 4 9, Richmond, viii.
36; <g, < 9, Newport, I1.0.W., 1912; all
Rothschild Coll.
Perdix perdix (L.) (Common Partridge)
C. garei, 1 9, banks of Bervie, Arbuthnot,
Kincardineshire, 14.vi.og, nest; B.M. 1914.517;
from one bird), C. garet (1), D. g. gallinulae (several), C. farrent (2), C. rusticus (1),
C. columbae (1), and M. sciurorum (1). No other bird in Britain has such an im-
pressive list of ‘stragglers’ except the Puffin (Fratercula arctica) which nests in
burrows under conditions particularly favouring straggling.
A bird, on the other hand, which may possibly be unattractive to fleas is the
Swift (Apus apus). It is classified among Nordberg’s hole or half-hole nesters, and
has a fairly high nidicolous fauna. Furthermore the sites chosen, the tendency to
return year after year to the old nest and to breed in colonies, and also the close
texture of the nest, should all contribute—as indeed they do in the case of the House-
Martin—to a rich flea-fauna, both with regard to the number of species and to the
size of the flea population. The very reverse is true, since the only fleas found in
the nest of the Swift are C. gallinae and C. fringillae, which are no doubt acquired
from the old Sparrows’ nests of which the Swift often makes use if they are situated
in suitable holes or cracks (Witherby, 1938).
Another curious fact is the absence of C. gavei from the nests of the Herring-Gull.
This flea has been recorded from L. ridibundus, L. canus, L. marinus, and L. fuscus
in Britain, but the only record of C. garei from the Herring-Gull (Waterston, IgIo)
has proved on re-examination of the specimens to refer to atypical females, probably
of C. borealis (Rothschild, 1948). Owing to the search for C. vagabunda we have
a number of negative records of the Herring-Gull’s nest—negative as far as C. garet
is concerned. There are sixteen such records from R. M. Allan and a further nine
(at least) from E. O’Mahony. Is there some unknown factor about the Herring-Gull
or its nest which suits C. vagabunda but not C. garei?
Swans, also, seem unattractive to bird-fleas, and although their nests are accessible
ENTOM. Il. 4 Ee
226 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
and frequently searched no fleas have been recorded from the genus Cygnus in any
part of the world. The same applies to the Kingfishers (Alcedo).
Seasonal variation in numbers. The monthly records of fleas found on the bodies
of British birds by the three chief collectors are set out in Table 10. Fleas are seen
to occur in every month of the year, but the lack of negative records and the small-
ness of the numbers make it impossible to compile seasonal frequencies. Colonel
Meinertzhagen has informed me that most of his collecting was done in the months
of April, August, and October. Thus, for example, out of 131 specimens of Sturnus
v. zetlandicus, 111 were shot in August. The record of ten infested birds in March
(Table 10) is nevertheless interesting, since nesting of the species concerned has
scarcely commenced, although some individuals may be visiting old sites. It is to
be expected that the peak of infestation occurs in the breeding-season. Three
additional midwinter records not included in Table 10 are available, C. gallinae from
the Siskin (Carduelis spinus) in February (Dale, 1878), the same species—a heavy
infestation—from the Black Redstart (Phoenicurus ochrurus gibraltariensis) in
November (Williamson, 1939), and a specimen of C. gavei from the Blackbird also in
November (Turk, 1946). These winter records, although few in number, do prove
conclusively that fleas may be present on the bodies of birds all the year round.
Fleas on passage migrants. Two of Colonel Meinertzhagen’s records come from
migrant birds—a male C. gallinae from Sylvia b. borin and a female C. gallinae from
Sylvia n. nisoria shot in the Shetlands in August, 1939. Since Colonel Meinertzhagen
was at the time collecting specimens for the purpose of studying the British races
of birds, he avoided as far as possible shooting migrants, and the total is therefore
deceptive. Mr. O’Mahony has kindly sent me two unpublished records of a male
and a female D. g. gallinulae collected from two migrant Wheatears (Oenanthe
oenanthe) from Fair Isle in August 1950. Two separate records of D. g. gallinulae
(without dates) from the body of Sylvia communis taken by R. M. Lockley at
Skokholm Bird Observatory were from migrants. The Black Redstart referred to
above was one of a party of migrants which reached the Isle of Man on 12 November.
There is no reason to suppose that migrant birds are free from fleas, and in fact the
contrary is indicated.
Time spent on the host’s body. It is a matter of interest that D. g. gallinulae is
more closely associated with the body of the host than either of the other two
common bird-fleas (Table 4) and probably spends relatively less time in the nest.
This may be one of the reasons why it has succeeded in spreading from the New
World to the Old. In r91r K. Jordan and C. Rothschild found several larvae of
D. g. gallinulae feeding on the epidermis under the wings of a dead nestling Black-
bird, another piece of evidence possibly suggesting a closer association with the host
itself than is usual with bird-fleas. Living nestlings are rarely examined for flea
larvae. Another matter concerning this species which is worth investigating is the
question of its breeding-season. It was first shown by Bacot (1914) and later con-
firmed by Ioff (1939) that C. gallinae, unlike many mammal fleas, has a fixed breed-
ing-season, which coincides with the spring and summer nesting-season of its hosts,
and which cannot be altered under laboratory conditions by a rise or fall in tem-
perature. Dasypsyllus, unlike Ceratophyllus, is a tropical genus and it would be of
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS 227
interest to know whether D. g. gallinulae has acquired a fixed spring breeding-season
like C. gallinae in the Palearctic Region.
TABLE I0
Monthly Records of C. gallinae,* C. garei, and D. g. gallinulae from
the Bodies of Bird Hosts in Britain
Collectors Nie i. M. A, M. Ve VE A; S: O. N. iD;
T. Clay & R. 3 I 5 16 4 8 I 22 fe) Fz, 5 fe)
Meinertzhagen
W. H. Pollen 3 fe) 5 5 5 I I fe) I fe) I 3
J. Waterston fo) fe) fe) 3 I 5 I o 3 2 fo) fe)
TOTAL 6 I Io 24 be) 14 3 22 4 14 6 3
* Records from domestic fowl omitted.
Sex-rvatio. In all large collections of Ceratophyllid bird-fleas from nests the females
are found to predominate, and there is probably a real divergence in the sex-ratio
of these species. In the case of mammal fleas the sex-ratio is often estimated from
the relative numbers on the bodies of the host, and in such cases the difference may
be more apparent than real. It has been shown, for instance, that the sex-ratio of
X. cheopis on the bodies of rats alters according to climatic conditions, males pre-
dominating on hot days and females on cold days (Cole, 1945), although there is
evidence that in the population as a whole the sexes occur in equal numbers. Mam-
mals are continually returning to their flea-infested lairs, nests, burrows, and runs,
and thus have the opportunity of acquiring fresh fleas or reacquiring specimens that
have moved off previously. In the case of birds, once the breeding-season is over the
chances of acquiring or ‘changing’ fleas must be greatly reduced. We may therefore
expect the sex-ratio of fleas on the bodies of birds to remain fairly constant through-
out the year. The marked difference between the proportion of males of D. g. gallinulae
on the bird and in the nest may in this case reflect a divergence in the feeding-habits
of the sexes. There is another point worth remembering in connexion with bird-
fleas. The sexes copulate freely on emerging from their cocoons, with or without
a blood meal, and therefore a single female carried through the winter on the body
of a host may be capable of laying fertile eggs in the nest in the following spring
without the presence of a male at that time, since she was probably fertilized the
previous spring in the nest. It has been shown that sperm is stored in the sperma-
theca and released when the eggs are about to be laid. It would be interesting to
know how long the sperm of bird-fleas remains viable in nature. In the laboratory
Pulex irritans has been known to lay fertile eggs sixty-five days after copulation
(Bacot, 1914).
Proportion of birds infested with fleas. Another question which springs to mind
concerns the actual proportion of birds which carry fleas on their bodies. This must
vary according to the species. We can deduce that House-Martins and Starlings,
for example, are more heavily infested than most species, and that the peak of
infestation occurs during the breeding-season. The figures in Table 4 do not give us
228 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
any information on this subject, because there are no negative records available.
Apparently few British flea collectors, unlike the collectors in East Africa (see p. 213),
keep negative records. The relative data for nine species have been supplied by
Miss T. Clay (Table 11). The infestation rate may be slightly higher than usual, since
TABLE II
Proportion of Birds infested with Fleas
(Collected by Colonel Meinertzhagen and Miss T. Clay)
Name of bird Total shot Total infested
Corvus f. frugilegusL. . : , 18 I
Rook
Corvus monedula spermologus Vieill. . 36 2
Jackdaw
Sturnus vulgaris zetlandicus Hart. . 135 4
Shetland Starling
Emberiza citrinella nebulosa Gengler . 56 4
Yellow-Hammer
Passer d. domesticus (L.) . é Z 35 3
House-Sparrow
Parus major newtoni Prazak . : 66 7
British Great Tit
Prunella modularis occidentalis (Hart.) go 6
British Hedge-Sparrow
Delichon u. urbica (L.) . - : a 2
House-Martin
Riparia r. riparia (L.) . : : 2 I
Sand-Martin
a large proportion of the collecting was done in April and August. The monthly
returns for the Great Tit (Table 12) show that in the case of this bird collecting —
happened to be rather more evenly spread out, and the preponderance of specimens shot
in April is not so marked. This gives a truer picture of the infestation rate, but un-
TABLE 12
Proportion of Great Tits (Parus major newtoni) infested with Fleas
(Collected by Colonel Meinertzhagen and Miss T. Clay)
Parus major newtont Prazak
British Great Tit as Bs AG IM fe Poe N AN ee 0. | Nu |e
Number uninfested . oe | ake 2 9 | I2 3 2 3 I 8 7
Number infested ; -| — {| — I 5} — I I —!/—]-};-f{]-
Total shot. - - fut (ites 2 10 T7 2 3 4 5 I 3 8 *
fortunately positive winter records are altogether lacking for this species. It will
be seen from Table 11 that nearly 7 per cent. of 445 Passerines examined carried
fleas on their bodies, and this is probably a fair estimate for the whole order in
Britain. In the case of the Hirundinidae the proportion may well be greater.
A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS 229
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* Papers containing records of British bird-fleas.
230 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
Torr, I. G. 1941. The ecology of fleas in connection with theiy epidemiological importance (in
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Saratov, 116 pp.; 157 figs.
JEFFERY, H. G. 1923. *Diptera: Pulicidae (Fleas). Proc. Is. Wight Nat. Hist. Soc. 1: 123-124.
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Jotiey, A. E., and Storer, R. 1945. *Report on the Sand-Martin Enquiry, 1944. Leicester-
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Jorpan, K. 1925. New Siphonaptera. Novit. zool. 32: 96-112, figs. 1-46.
1926. New Siphonaptera. Novit. zool. 38: 385-394; figs. 1-22.
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1937. Three new bird fleas from Kashmir. Novit. zool. 40: 299-306; figs. 87-92.
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NorRDBERG, S. 1934. Aphanipterologisches aus Finnland. I. Verzeichnis der bisher in Finnland
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A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS 231
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WATERSTON, J. 1906. *On some Scottish Siphonaptera. Ann. Scot. Nat. Hist.: 211-214.
1909. *On some Scottish Siphonaptera. Ann. Scot. Nat. Hist. 18: 226-228.
1910. *On some habits and hosts of bird Ceratophylli taken in Scotland in 1909, with
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WILLIAMson, K. 1940. *Numbers of Black Redstarts on passage in Man. Brit. Birds, 38 (9):
252-254.
* Papers containing records of British bird-fleas.
232 A COLLECTION OF FLEAS FROM THE BODIES OF BRITISH BIRDS
LIST OF TABLES
TABLE 1: Fleas collected by Colonel Richard Meinertzhagen and Miss
Theresa Clay from the bodies of British birds.
TABLE 2: Records of C. vagabunda Boheman (1866).
TABLE 3: British bird hosts of C. gallinae, C. garei, and D. g. gallinulae.
TABLE 4: Comparative records of C. gallinae, C. garei, and D. g. galli-
nulae (compiled from all known British records).
TABLE 5: Selected hosts on which C. gavei is the dominant species of
flea.
TABLE 6: Selected hosts on which C. gallinae is the dominant species
of flea.
TABLE 7: Selected hosts on which D. g. gallinulae is the dominant
species of flea.
TABLE 8: British records of C. gallinae, C. garei, and D. g. gallinulae
collected away from any host or nest.
TABLE 9: C. gallinae, C. garei, and D. g. gallinulae in the British
Museum Collection from named bird hosts.
TABLE 10: Monthly records of C. gallinae, C. garei, and D. g. gallinulae
from the bodies of bird hosts in Britain.
TABLE 11: Proportion of birds infested with fleas. (Collected by Colonel
Meinertzhagen and Miss T. Clay.)
TABLE 12: Proportion of Great Tits (Parus major newtont) infested with
fleas. (Collected by Colonel Meinertzhagen and Miss T.
Clay.)
PRESENTED
7 FEB 1952
OXFORD
| ae
CHARLES BATEY
PRINTER
TO THE
UNIVERSITY
17 APR 1952
THE SUB-GENUS STEGOMYIA
(DIPTERA : CULICIDAE)
IN THE
ETHIOPIAN REGION
(Part I)
teen) PINGLY
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 2 No. 5
LONDON : 1952
THE SUB-GENUS STEGOMYIA
(DIPTERA: CULICIDABE)
IN THE ETHIOPIAN REGION
I A PRELIMINARY STUDY OF THE DISTRIBUTION OF
SPECIES OCCURRINGIN THE WEST AFRICAN SUB-REGION
WITH NOTES ON TAXONOMY AND BIONOMICS
BY
re", MATIUNSIY
Yui
po
7
Pp. 233-304; 16 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 2 No. 5
LONDON : 1952
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is issued
in five series, corresponding to the Departments of the
Museum.
Parts appear at irregular intervals as they become ready.
Volumes will contain about three or four hundred pages,
and will not necessarily be completed within one calendar
year.
This paper is Vol. 2, No. 5 of the Entomology series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued April 1952 Price Fifteen Shillings.
THE SUB-GENUS STEGOMYIA (DIPTERA,
CULICIDAE) IN THE ETHIOPIAN REGION
I. A PRELIMINARY STUDY OF THE DISTRIBUTION OF
SPECIES OCCURRING IN THE WEST AFRICAN SUB-REGION
WITH NOTES ON TAXONOMY AND BIONOMICS
By P. F. MATTINGLY
CONTENTS
INTRODUCTION . ‘ ~ A F ‘ z 235
NotTEs oN TAXONOMY : ; ce F : 31 235
Key to adults : : : ; : ; ae |
Key to larvae ; . : : : : eA
DISTRIBUTION RECORDS . ; : F ‘ - 249
Doubtful records . . : : , : af 257
Records based on misidentifications : z = i258
Unidentified and misquoted localities : , « 258
Distribution outside the Ethiopian Region : - 259
List oF LOCALITIES WITH TOPOGRAPHICAL DETAILS . 259
BIONOMICS IN RELATION TO DISTRIBUTION . : - 269
Breeding-places_. ? ‘ ; F : 2 269
Seasonal distribution . ; : : ‘ 2 270
Biting-habits - . : : : : “1 275
ZOOGEOGRAPHY : A : : . 5 eo
SUMMARY 3 : : A : : ss = 293
ACKNOWLEDGEMENTS : : ‘ - : <- 4204
REFERENCES . : : ‘ ‘ ° : - 295
APPENDIX ; F : A é 5 : 303
SYNOPSIS
This paper deals with the distribution of the species of Stegomyia occurring in the West African sub-
region. All the species of wide distribution in the Ethiopian Region are, however, included and the topo-
graphical data therefore cover most of the region. Aédes aegypti is treated very briefly since it is regarded
as a special case meriting a separate paper. Zoogeography is discussed mainly in relation to rainfall and
altitude. Temperature and vegetational relationships will be discussed in later papers. Such notes on
taxonomy and bionomics are included as appear to be necessary for an understanding of distribution.
These also will be discussed at greater length in later papers in the series.
INTRODUCTION
THE objects of the present study are firstly to render possible a more rational approach
to the taxonomy of the sub-genus Sfegomyia, and incidentally of other African
insects, and secondly to discover what light may be thrown by this comparatively
well-known group on general problems concerning the distribution of the Ethiopian
fauna. It is perhaps not too much to hope that some small quota may be added to
236 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
the growing accumulation of evidence regarding past climates in Africa which is of
fundamental importance for the understanding of phylogeny and of the human,
plant, and animal geography of the continent.! Ernst Mayr (1942) has clearly shown
that it is as a population that the species is encountered in nature, and it is upon
population studies that our knowledge must ultimately be based. A taxonomy based
solely on the arbitrary selection of morphological types, while indisputably essential,
can only be of ancillary importance. In the author’s opinion the greatest need for
mosquito studies at the present time is the application of genetical methods (on this
point see Bates, 1949).
The upper limit to our knowledge of distribution is imposed by the supply of
collectors. In the present instance records are available from more than 600 localities
(Fig. 1), but not all of these have been investigated with equal thoroughness. In
general the British territories have rendered the best service, but it is unhappily
true that the most serious gap in our knowledge at the present time is provided by
Tanganyika, of which only a narrow fringe along the northern border and along the
coast is known. Vast areas of unknown territory exist in Angola, the Cape Province,
and French Equatorial Africa, but these are so situated that discoveries there would
be likely to add to rather than to integrate our existing knowledge. From the
phylogenetic point of view most knowledge could probably be gained from the study
of high-altitude faunas. At present only those of Ruwenzori, the Kigezi district, and
a few parts of Kenya are known. .
The present paper deals only with those species occurring in the West African
sub-region. Those which are known only from the East and South African sub-regions
will be dealt with in a later communication. This does not mean that only the West
African districts are considered. All the Stegomyia species of wide distribution in
Africa enter this sub-region, and these species must necessarily be considered in
relation to their range as a whole if their distribution is to be understood. The dis-
tribution of one species, A édes aegyPti, is so artificial and so many records are available
that to deal with these in full, as has been done for the other species, would serve
little useful purpose, besides taking up a wholly disproportionate amount of space.
Details of the distribution of this species may be found in Kumm (19310) supplemented
by Edwards (1941) and by the notes on recent records given below. Some small
attention is paid, in the section on breeding-places, to the occurrence of wild popula-
tions. It is possible that some idea of the natural distribution of the species, which is
almost certainly Ethiopian in origin, might be gained if more records of this kind were
available.
Apart from one or two important collections which the author has not as yet had
the opportunity to examine, all known sources of distribution records have been
tapped. It is not believed that the collections in question will add to any great extent
to our knowledge, as much of the material which they contain has been published.
Great difficulty has been experienced in tracing many localities and in ascertaining
the known facts regarding their climate and topography. In order to save others this
labour the best available data have been included in tabular form. The major
factors affecting distribution may be considered to be rainfall, temperature, humidity,
t For a summary of this work see Zeuner (1944).
IN THE ETHIOPIAN REGION 237
and vegetation. The last named is itself dependent to so large an extent on the other
factors that it is preferred to discuss it separately in a later communication. It is
here referred to only where, as in the case of the Guinean forests, it appears to con-
stitute a gross physical barrier to distribution. Attention has been paid to the
possible influence of human transportation, but this is considered to be only a minor
io ° .
1° ° ° x0 39 ipo° 50°
PEM
“T
20° 10° 0° "10° 20° 30° ba? 50°
Fic. 1. Distribution of collectors of Stegomyia spp. other than Aédes aegypit.
factor. Its importance for the mosquitoes is very liable to exaggeration and only the
merest handful of species can be shown ever to have become established outside their
natural boundaries, while very striking cases of failure to become established even
under apparently favourable conditions can be quoted. Among the species here
discussed only the domestic Aédes aegypti, and Aédes simpsoni which commonly
breeds in the leaf axils of food plants, show any sign of having been affected in this
way. Man as a destroyer of forests is perhaps more important, but this factor has
relatively little significance for the present study, which is concerned rather with
distribution at the present time and the factors by which this is controlled than with
distribution in the past. The consideration of temperature and humidity has had to
238 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
be postponed for a subsequent communication owing to the relative inaccessibility of
the necessary data, but altitude is considered for the sake of the light which it throws
on temperature limits. Rainfall, for which quite abundant figures have recently
become available, is considered in full, and rainfall boundaries are found to provide
a surprisingly, in some cases almost dramatically, accurate fit with known Stegomyia
distributions, especially when the seasonal distribution of rain is taken into account.
It must, however, be emphasized that the observations here recorded are purely
empirical. It is not claimed that rainfall per se governs distribution. On the contrary,
such evidence as is available suggests that for the most part it operates indirectly
through its influence on vegetation.
Detailed taxonomic study of the sub-genus is in progress and is not yet complete.
The treatment here adopted is therefore largely conventional except as regards
dendrophilus, which has been the subject of much confusion. Work already carried
out has, however, revealed a considerable number of errors in published descriptions,
and these are corrected in the notes on taxonomy. The keys to adults and larvae
cover all Ethiopian members of the sub-genus and have been made as up-to-date as
possible. No descriptions of new species are at present known to be in the press, but
the position of a new form from the East African Highlands is under consideration
and will be discussed in the second paper.
The section on bionomics is confined to those aspects of the subject having an
obvious and immediate bearing on distribution. In this connexion it may be said
that the most serious gap in our knowledge concerns seasonal distribution, the proper
understanding or even the mere recording of which would illuminate every aspect of
mosquito studies.
NOTES ON TAXONOMY
Edwards (1932) divided the sub-genus Stegomyza into four groups. Of these, group
A contains all the purely Ethiopian mainland species and the cosmotropical A édes
aegypti. Group B contains certain Oriental and Palearctic species and Aédes mas-
carensis of Mauritius. Group C contains Oriental and Australasian species and A édes
albopictus, granti, and unilineatus, part of whose range is Ethiopian. Group D contains
only Aédes vittatus, which is Palearctic, Oriental, and Ethiopian, and shows certain
characters anectant to the sub-genus Aédimorphus. In so far as this grouping was
based on the male terminalia it appears to have been a good one, but colour charac-
ters in Aédes are somewhat unsatisfactory and mascarensis should perhaps have been
placed in group A. In any case a new system of grouping has been rendered necessary
by the discovery of Aédes amaltheus (De Meillon & Lavoipierre, 1944), which is
anectant between groups A and B, having the colour characters of one and the male
terminalia of the other, the discovery of certain aberrant species of Stegomyza in the
Mariana Islands (Bohart & Ingram, 1948), and the proposed transfer to this sub-
genus of certain species hitherto assigned to Armigeres (Brug & Bonne-Wepster,
1947). Such a regrouping is, however, beyond the scope of the present paper, and the
following notes refer only to matters of detail.
Aédes apicoargenteus. Specimens from Kampala have a number of pale scales on
the lateral lobes of the scutellum. Edwards (1941) states that in the ‘typical form’
IN THE ETHIOPIAN REGION 239
these lobes are entirely black-scaled but gives no further explanation. This has led
to some confusion. Other specimens with the lateral lobes white-scaled are recorded
from Kenya (Van Someren, 1946), and Wolfs (1949) notes that they are white-scaled
in ssp. denderensis from Costermansville. This is a rather unstable species, perhaps of
comparatively recent origin, and the occurrence of this type of peripheral variation
does not seem to have much taxonomic significance. It is, however, noteworthy that
pale scaling has only been recorded from the higher parts of the range, above 3,000
ft. As will be shown below, apicoargenteus seems to be more strictly confined to low
altitudes than, for example, africanus, whose range is otherwise similar. Its highland
counterpart, schwetzi, has white lateral lobes. The occurrence of pale scales on the
lateral lobes in apicoargenteus unfortunately renders separation from fraser rather
more difficult since this species frequently exhibits a number of dark scales in this
position (see below). The extent of the pale area on the posterior surface of the hind
femur is used by Edwards (1941) as a key character, but this is so variable in different
parts of the range as to be extremely unreliable. The best diagnostic character from
fraseri, as pointed out by Van Someren (1946), is the metallic sheen on the pale
. markings of the femora; schwetzi is easily separated by the very extensive pale
scaling in the pre-scutellar region, but the differences in the male terminalia given by
Edwards (1941) do not appear to be significant. The twisting of the style observed
_ in schwetzi is due merely to the angle from which this was drawn and apicoargenteus
also shows it when drawn from the same angle. The number and length of the bristles
on the basal lobe is very variable in apicoargenteus and overlaps with that found in
schwetzi. An average figure would perhaps be 8, but some specimens may have as
many as 12. West coast specimens sometimes have fewer bristles than those from
farther East, but the difference is not constant. The only genitalic character which
seems to be of much diagnostic value concerns the [Xth tergite, which is rather more
deeply excavated in schwetzi, but even here there is some variation in both species.
It is rather doubtful whether schwetzi should be treated as more than a sub-species,
but further material is required from intermediate localities before the point can be
settled.
The larvae of apicoargenteus, frasert, and schwetzi are all very similar and closely
resemble those of the East African calceatus, soleatus, and langata. As far as can be
ascertained from available material, apicoargenteus and fraseri can generally be
separated from the other species on the position of the antennal bristle. In fraseri
this arises at 0-82—0-85 x the distance from base to apex of the antenna. In apico-
argenteus the corresponding figure is 0-67—0-82. Specimens of this species from
Uganda show a rather greater average displacement from the base than do those
from Nigeria, Cameroons, and Belgian Congo, and this may be an altitudinal effect
comparable to that described in connexion with the scutellar lobes. Average positions
in material so far available are 0-80 for Uganda, 0-74 for Nigeria, and 0-75 for Belgian
Congo and Cameroons, but the samples are too small for proper statistical analysis.
In all specimens of the calceatus group examined the figure is less than 0-70. schwetzi
overlaps the two groups. As noted by Van Someren (1946), those larvae of fraseri
so far examined can be separated from apicoargenteus by the greater reduction of the
median denticle of the comb spines (with proportional emphasis of the lateral fringe),
240 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
but all specimens so far available come from Kenya and it is by no means certain
that this character would be diagnostic for West Coast larvae. From all other
members of the group, except ssp. denderensis, calceatus can be separated by the
possession of a wholly dark siphon. This character is not mentioned in the type
description of denderensis (Wolfs, 1949), but it is shown by both larvae available for
examination and is very probably constant. It is a striking character, and although
the differences from the type form with respect to male terminalia given in the type
description are not constant, it is considered that this form should be named as a
sub-species. Further material from the Kivu Highlands and Ruanda-Urundi would,
however, be very welcome. Differences in colour characters between the sub-species
and West Coast specimens of the type form include smaller anterolateral scutal
patches, more frequent occurrence of pale scaling on the lateral lobes of the scutellum,
narrower pale bands on third hind tarsi, and reduced pale scaling of fifth hind tarsi.
None of these are, however, absolutely diagnostic from Kenya and Uganda specimens
of the type form.
Aédes fraseri. This species has already been discussed, but a further word is
perhaps necessary regarding the scaling of the lateral lobes of the scutellum. Edwards
(1941) states that the scales in this position are ‘always white’, a statement belied
by his figure (Fig. 39). In fact some dark scales are usually present. It is probable
that West Coast apicoargenteus can be reliably distinguished from frasert by having
the lateral lobes wholly or very largely black, but the distinction does not hold good
in the eastern part of their range. As noted by Edwards (1941) and Van Someren
(1946), the shape of the anterolateral pale scutal patches and the extent of the pale
area on the posterior surface of the hind femur are subject to much variation.
Aédes dendrophilus. This species has been misidentified by Van Someren (1946)
and others by reason of the misleading description given by Edwards (1941). This
description should be ignored and that given by Van Someren (loc. cit.) substituted
for it. Edwards appears to have been misled by the very striking sexual dimorphism
occurring in dendrophilus. This is an unusual phenomenon in Stegomyia, although a
small degree of dimorphism with respect to, for example, the abdominal tergites is
common. An even greater degree of dimorphism characterizes some oriental Finlaya,
however (see photographs in Barraud, 1934). The characteristic markings of the male
as described by Van Someren for the Kenya and Uganda forms are also shown by the
Gold Coast form, and Edwards’s type 3, although damaged, shows them well. By
reason of the same error of observation Edwards failed to recognize the identity of
dendrophilus with Aédes trinidad. Through the kindess of Dr. Ceballos and Dr. Peris
of the Madrid Museum it has been possible to examine the two cotype males of this
species and it is clearly conspecific with dendrophilus. It should be noted that the
white spot on the clypeus referred to in the type description of trinidad (Gil Collado,
1935) is due to frosting of the integument and not to the presence of scales. Aédes
aegypti thus remains distinguishable from all other Stegomyia by the presence of
scales in this position. A careful comparative study reveals that Edwards’s type series
of Aédes demeilloni from Eshowe, Zululand, contains a specimen of dendrophilus
which can be recognized by-the character given in the key. The correctness of this
diagnosis was suggested by the fact that the specimen in question was bred out from
IN THE ETHIOPIAN REGION 241
a tree-hole, whereas all those of demezliont came from axils of Dracaena or ‘ Fern Tree’,
Both the identity of the Eshowe specimen with dendrophilus and the distinctiveness
of the breeding-places of the two species have been confirmed by the receipt of further
specimens of both sexes kindly sent by Dr. De Meillon, and Mr. Muspratt has made
extensive collections of both in Natal which add further confirmation. This curious
but by no means unique distribution is discussed more fully in the appropriate
section. The Kenya—Uganda, Sierra Leone—Gold Coast—Nigeria, Fernando Po, and
South African forms all show minor differences in colour markings, and the latter at
least should perhaps be named as a sub-species. The point will be discussed in a
later paper to be devoted to taxonomy. The problem is complicated by the fact that
Edwards’s type series of dendrophilus consists of a number of dwarfed specimens,
some with slightly atypical markings. These have the appearance of specimens bred
out in the laboratory on an inadequate larval diet. The presence of thickened, spine-
like setae on the inner angle of the basal lobe of the male terminalia of dendrophilus
is not sufficiently indicated either by Edwards’s drawing or by his figure. Such spines
are characteristic of a number of species in this group (dendrophilus, deboeri, bambusae,
heischi, demeilloni, keniensis, angustus) which cannot safely be separated either by
the degree of development of the spines or the shape of the lobe, both of which are
very variable. The larva of Aédes deboert is not constantly separable from that of
dendrophilus. Kenya and Uganda specimens of the latter commonly have the sub-
ventral seta of the siphon single (usually double in deboert), but it is frequently double
in the Gold Coast form. The best partial character appears to be the possession of
more than one pecten tooth beyond the subventral seta. This is of freqent occurrence
in deboert but has never been observed in dendrophilus. In bambusae the subventral
seta is commonly bifid but may be single, and there may be either 1 or 2 pecten teeth
beyond it.
Aédes africanus. Edwards (1941) gives a number of striking colour differences
between this species and luteocephalus. Most of these differences are, however, very
inconstant and this has led Haddow and Mahaffy (1949) and others to suggest that
the two forms are interbreeding sub-species. With this opinion the present author is
in strong disagreement. An examination of a large series of both species from many
parts of their joint range shows that the extent of the pale basal marking on the hind
femur affords an absolutely constant diagnostic character. Even in the most extreme
forms the difference is striking, and in almost the whole of the material it is so gross
as to be easily detectable with the naked eye (Fig. 2). Of the other characters pro-
posed by Edwards the most reliable is perhaps the extent of the antero-median
silver stripe on the mesonotum. Unfortunately, however, too many of the available
specimens are rubbed in this position to be certain on the point. A fairly reliable
character is the extent of the pale scaling at base of the 4th hind tarsal. This nearly
always forms a complete ring in africanus, but in a few specimens from peripheral
parts of the range the ring is interrupted ventrally. In luteocephalus the whole
segment is commonly dark, but a small basal pale spot may be present on the dorsal
surface, and in some specimens from the West Nile Province of Uganda this is fairly
extensive and resembles the rather narrow pale interrupted band sometimes found
in africanus. The remaining characters are of local value only, although their totality,
ENTOM, Il. 5 Gg
242 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
in the great majority of cases, renders identification both certain and easy. The
larval characters proposed by Hopkins (1936) are very largely unreliable and should
be ignored except where experience has shown them to be locally applicable.
Aédes pseudoafricanus. The small difference from africanus with respect to scutal
markings noted by Chwatt (1949) appears to be constant in all available specimens.
The whole of these, however, come from the Lagos area, and it is by no means certain
that the character would prove constant should the species be discovered elsewhere.
All male specimens attributed to africanus in the British Museum collection have been
dissected and these have yielded only a few pseudoafricanus from Yaba. The resem-
blance of the male terminalia to those of fraseri is striking but it is not absolute. The
os. i 4
_ ——— m2?
a c
steppe ase DR
ae i
d PEM
Fic. 2. Left hind-femur. a—b. Aédes africanus; c—d. Aédes luteocephalus. a from Ndola, 6 and ¢
from Godia, d from Likoma Island. d is very exceptional.
posterior edge of the IXth tergite, for example, is quite different in shape in the two
species. The combination of colour characters resembling those of one species with male
terminalia resembling those of another is not unique among the Ethiopian Stegomyia.
It is also shown strikingly by amaltheus (De Meillon & Lavoipierre, 1944). In view of the
very curious results of cross-breeding experiments between certain species of Stego-
myia, such cases must necessarily carry a suggestion of naturally occurring hybridiza-
tion, and this is lent some colour in the case of pseudoafricanus by the distribution of
the species concerned. In the experiments concerned Toumanoff (1937, &c.) showed
that hybrid offspring of Aédes aegypti and Aédes albopictus exhibit all the morpho-
logical characters of the female parent in whichever direction the cross is made. This
was confirmed for the aegypti 9 x albopictus 3 cross by Hoang Tich Try (1937) and
for the reverse cross by Downs and Baker (1949). The genetical and cytological
background is obscure, although some light is thrown on the problem by the discovery
that in the mosquitoes, or at least in those species so far examined, there is no
differentiation of sex chromosomes (Sutton, 1942 ; Callan & Montalenti, 1947 ; White,
1949). Gilchrist & Haldane (1947) consider that sex in these insects is determined by
a single dominant gene for male. Downs & Baker (1949) have suggested that the
phenomenon is one of parthenogenesis following mechanical fertilization of the ovum.
This would not, however, explain the very interesting observation made by Bonnet
(r950) that hybrid offspring of aegypti 2 x albopictus 3, although themselves identical
IN THE ETHIOPIAN REGION 243
with aegypti, produce offspring with albopictus characters when back-crossed to pure
line albopictus gs. While these observations are of considerable interest in relation to
the phylogeny of the sub-genus it is by no means certain that hybridization takes
place in nature. Rozeboom and his fellow workers are of the opinion that it does not
since females of aegypti and albopictus in cage colonies show an exclusive preference for
males of their own species when given the choice, even though they may mate with
males of the other species when no others are available (Mattingly e¢ al., in press). In
the present author’s opinion the most probable explanation of the occurrence of these
intermediate forms is that they are genuinely anectant in the phylogenetic sense.
The Lagos populations of africanus and pseudoafricanus are effectively sympatric
at the present time, since, although the larval breeding-places are segregated, adults
of both sexes of both species are taken together near the fringes of the mangrove
swamps (Mattingly, 19490). It does not, however, follow that this is a case of sym-
patric speciation. The area of low rainfall occurring between Lagos and Cape Three
Points and serving to separate the Upper from the Lower Guinean forests is commonly
supposed to result from a comparatively slight change in the direction of the coastline
in relation to the direction of rain-bearing winds. If this is true, then small changes
in the level either of the sea or of the land may well have served in the past to isolate
extensive areas of mangrove ‘forest’ from the true forest with consequent speciation
of mangrove and true forest Stegomyia, an interesting example of the possible effect
of small changes of elevation on the climate, flora, and fauna.
Aédes simpsoni. Haddow, Van Someren, eé al. (in the press), note that this is
a, very variable species and express doubts as to the usefulness of applying the name
‘var. lilit’ to one among many varieties. It would seem, however, that the simpsonz of
the Transvaal, Natal, and Cape Province may be largely isolated from those occurring
in the rest of Africa (Fig. 12), and it is felt that if good representative collections
could be assembled these would repay careful study. It is proposed to consider the
matter from this angle in the course of the taxonomic work on the group. For a note
on variation in this species see Edwards (1941).
Aédes luteocephalus. This species has already been discussed under africanus. The
type locality is wrongly quoted in the type description (see under ‘ Doubtful Records’,
p. 257).
Aédes unilineatus. Difficulties experienced in identifying the larvae of this species
(De Meillon & Lavoipierre, 1944; Lewis, 1945) were due in part to an error in the
legend to Fig. 61 of the first volume of Mosquitoes of the Ethiopian Region (Hopkins,
1936). An examination of the original drawings for this figure and the specimens on
which they were based shows that the comb and pecten spines attributed to unilineatus
were in fact drawn from a specimen attributed by Ingram & De Meillon (1929) to
pseudonigeria and reassigned by Edwards (1936) to demeillont. The identity of this
specimen is still uncertain, but it is certainly not unilineatus. The larva illustrated,
in the same figure, as demezilloni is in fact angustus, and neither demeillonit nor uni-
lineatus is figured. Typical comb spines of wnilineatus have very small secondary
denticles which are almost or quite invisible under low powers of the microscope.
An occasional spine may have one or two large denticles, but such spines are clearly
aberrant and should cause no difficulty. An extreme example is shown in Fig. 3
244 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
together with a normal comb spine. The pecten spines are very variable. The larva
of unilineatus is very distinctive and should not be confused with any other occurring
on the mainland. It can be recognized by the characters given in the key. The
degree of development of the barred area of the ventral brush varies even in specimens
from the same locality, but it is never as well developed as in the majority of species.
Barraud (1934) notes that in Ethiopian wnilineatus the saddle hair is usually 5-
branched while in Indian larvae it is usually bifid. Such South African larvae as
have been examined, however, show the bifid condition. Sudan and West Nile
larvae have 4-5 branches, but Gold Coast larvae appear to produce either type of
=
SS
a La
b
PEM
Fic. 3. Aédes unilineatus. a. Comb spines. b. Pecten spines. c. IXth tergite of male. The lower
comb spine is aberrant. This condition is only seen in an occasional spine.
saddle hair at random. The descriptions of the male terminalia given by Edwards
(1941) and Barraud (1934) are misleading since both appear to have overlooked the
presence of two small, very strongly sclerotized setiferous lobes on the outer edges of
the IXth tergite (Fig. 3). These are not shown in the only dissection left by Edwards
in the British Museum owing to the way in which this is mounted, but they show
clearly in fresh mounts from the same locality (Accra) and from Gadau, Erkowit,
and Lahore.
KEY TO ADULTS
1. Scutum with 3 pairs of small white spots; tibiae white ringed near middle;
proboscis usually pale inmiddle : 2 : . vittatus Bigot"
Scutum and tibiae otherwise; proboscis dark. : : : : 2
2. Proboscis and tibiae with white lines . : ‘ . grantt Theobald
Proboscis all dark ; tibiae dark except, in some cases, at base ; 3
3. Front half of scutum white scaled : : ‘ mascarensis MacGregor
Scutum otherwise marked . : A ; f ; 4
4. Front half of scutum with a median white stripe only : ‘ ; 5 5
Scutum otherwise marked . 5 : i : ; : , . 6
1 Species dealt with in the present paper are shown in heavy type.
Io.
rT.
TZ:
13.
I4.
15.
16.
ty B
18.
I
IN THE ETHIOPIAN REGION 245
. Middle femur with a white spot in middle in front . unilineatus Theobald
Middle femur without such a spot } : . albopictus Skuse
. Middle femur with a white stripe in front reaching nearly to tip; clypeus
of female scaly . ; ; : . aegypti Linnaeus -
Middle femur without such a stripe; clypeus bare ; : ; : 7
. Third hind tarsal much more broadly pale than the others. , , 8
Third hind tarsal only narrowly pale . ‘ II
. Hind femur with a well-marked pale patch at base broadly expanded below
and behind (Fig. 2) ; ; : luteocephalus Newstead
Hind femur at most with a small pale spot at base : 9
. Posterolateral lines of scutum broad, deep yellow, conspicuous, reaching
forward almost or quite to the posterior ends of the anterolateral patches ;
thorax laterally compressed. ; A ‘ . ruwenzort Haddow &
E. C. C. Van Someren
These lines shorter, less conspicuous; thorax of normal width , =. 210
Anterior median stripe of scutum very short; antero-lateral patches on
scutum slightly crescentic, concave behind : . africanus Theobald
This stripe longer ; scutal patches wedge-shaped, straight-edged behind .
pseudoafricanus Chwatt
Fourth hind tarsal segment all black . , ‘ : ‘ ‘ ee.
Fourth hind tarsal segment pale at least at base . : 14
Lateral lobes of scutellum white scaled ; anterior median spot on scutum
composed of broad or narrow scales . ; 13
Lateral lobes of scutellum dark scaled ; this spot composed of narrow - scales
only os : : , : , : . woodt Edwards
Hind tibia all donk helow , ‘ : : ; simpsoni Theobald
Hind tibia extensively pale below at base : ; . Strelitziae Muspratt
Prescutellar area of scutum covered with broad metallic scales
metallicus Edwards
This area with narrow, non-metallic scales . : 15
Posterior half of scutum with pale scales forming three or four n narrow
longitudinal lines . ; ; a 16
Posterior half of scutum with pale scales not ageregated into lines
chaussiert Edwards
Median area of scutum with paired yellow longitudinal lines inside the
usual posterolateral lines ‘ : : ‘ , , : ee
This area with a single median line -. ; ‘ ; ee
Fourth hind tarsal segment white at base only : : subargenteus Edwards
This segment all white : ; . subargenteus ssp. kivuensis Edwards
Each tibia with a narrow pale ring slightly removed from base
pseudonigeria Theobald
White marks on tibiae confined to extreme base (except sometimes on the
hind leg) ; 19
g. Hind tibia all dark (occasionally with a ‘very small number of pale scales at
base below in keniensis) . ‘ : ; ; : ‘ é i 26
2
246
20.
ZX.
22.
23.
24.
25.
26.
27.
28.
29.
30.
aT.
THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
Hind tibia with a well-marked Pale patch at base, at least on the under
surface : : : > ae
Third hind tarsal pale all round on at least the basal 1th
honioasis E. C. C. Van Someren
Third hind tarsal all dark or, at most, with one or two pale scales at base
below . : 21
Median longitudinal line of scutum white; prescutellar bare patch witha a
conspicuous border of pale scales. amaltheus De Meillon & Lavoipierre
Median longitudinal line yellow; pale scales round prescutellar bare patch
reduced or absent . : : masseyt Edwards
Pale basal patch on 6th abdominal tergite prolonged backwards nearly to
the apex of the segment : Tk ea
This patch shallower, at most half the depth of the segment . , eB
First mid-tarsal pale behind pk to the tip, pale in front on only about the
basal 3th : : : soleatus Edwards
This segment pale above on at most the basal Sths, usually much less,
more narrowly pale below, without any notinealde extension of pale
scaling behind ; : ; : ; : ae |
Pale markings of femora snowy white . : 5 ‘ fraseri Edwards
These markings metallic silver . ; 25
Prescutellar bare patch with a broad border of yellow scales ; ; 5th hind tarsal
entirely dark . ; schwetzi Edwards
Prescutellar bare patch with at most a “very | narrow border of pale scales ;
5th hind tarsal at least narrowly pale at base above (except sometimes in
ssp. denderensis) . . apicoargenteus Theobald
First mid-tarsal segment pale behind for most of its length . : 27
This segment more or less uniformly ringed at base, without any cetmentle
extension of pale scaling on the posterior surface : : ot Lae
5th hind tarsal entirely dark : ‘ ‘ calceatus Edwards
5th hind tarsal pale at base. : heischi E. C. C. Van Someren (in part)
Hind femoral knee-spot suppressed, represented by at most two or three
pale scales ; mid-femoral knee-spot, where present, at most very small,
not reaching to the tip of the femur : 29
Hind femoral knee-spot well developed ; mid-femoral knee- -spot reaching t to
the tip of the femur above. 30
Thorax laterally compressed ; mid- femur nearly always with a white spot on
the anterior surface at about half-way between base and apex
angustus Edwards
Thorax of normal width; mid-femur without such a spot bambusae Edwards
Mid-femur with a conspicuous white spot on the anterior surface at about
two-thirds the distance from the base; 5th hind tarsal dark . wag.
Either with the anterior surface of the mid-femur dark except for the knee-
spot or with the 5th hind tarsal pale at base or both . , ,\ wae
Middle tibia pale below on at least the basal 4th . : | powerk Theobald
Middle tibia usually all dark, at most with a small pale spot at base oe
32.
33:
34.
35.
36.
IN THE ETHIOPIAN REGION 247
Hind tibia pale below on at least the basal #,ths, this pale stripe expanded
distally ; median longitudinal pale line af scutum expanded in front of
prescutellar bare patch and fused with the pale border round this patch ;
posterolateral lines of scutum yellow ; second mid-tarsal pale above on at
least a half, usually more . ‘ langata E. C. C. Van Someren
Hind tibia pale on at most 4, the pale stripe at most very slightly expanded
distally ; median pale line of scutum of uniform width throughout, not
fused with the pale scales at the sides of the prescutellar bare patch;
posterolateral lines usually white or whitish; second mid-tarsal pale
above on at most 4, usually less : ; P contiguus Edwards
Fifth hind tarsal largely dark, at most narrowly pale at base; anterior
surface of mid-femur dark except for knee-spot ‘ . deboeri Edwards
Fifth hind tarsal pale on at least the basal 4rd, often more, or, “if not, then
anterior surface of mid-femur with a silvery spot at about 3rds ; 34
Pale streak at base of hind tibia at most jth the tibial denptly usually less ;
2nd mid-tarsal broadly pale above on at most the basal half, apical half
dark or at most with a very narrow dorsal line of pale scales. 35
Either with the pale streak at the base of the hind tibia much more than
sth the tibial length or with the 2nd mid-tarsal broadly pale above nearly
to tip or both : : : i ae
Anterior surface of mid-femur usually with a . spot of silvery scales
heischt E. C. C. Van Someren (in part)
Anterior surface of mid-femur usually dark except at tip demeilloni Edwards
Anterolateral scutal patches pointed in front, varying in colour from
bright yellow to creamy white; basal bands on abdominal tergites
yellowish . ‘ ; . bambusae ssp. kenyae E. C. C. Van Someren
Anterolateral scutal patches rounded in front, snowy white; tergal bands
white . es ; : . ; : . @endrophilus Edwards
KEY TO FOURTH-STAGE LARVAE
Ventral brush with at least the great majority of setae single : : 2
Ventral brush with at least the great majority of setae branched . ; 5
Lateral seta of saddle and subventral seta of siphon single. . angustus
These setae each with at least 2 branches . : 3
Comb spines with a small number of well-developed lateral denticles, at
least the submedian ones easily visible under low powers of the micro-
scope ; antennal seta with 2-4 branches . . . . amaltheus (in part)
Comb spines with the lateral denticles less strongly developed, invisible
or visible only with difficulty under low powers of the microscope;
antennal seta single : ; 4
. Barred area of ventral brush absent or poorly developed : saddle hair often
with more than 2 branches. : . unilineatus
Barred area of ventral brush well developed : saddle hair bifid . albopictus
. Ventral brush with 2 or more isolated tufts arising proximal to the barred
area. : ; $ : : : : ; ; ; : 6
248
IO.
DE.
12.
13.
14.
BS:
16.
17.
THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
Ventral brush without unpaired tufts proximal to the barred area . : 7
. Antenna lightly spiculate ; setae of ventral brush with numerous branches ;
comb spines with lateral denticles only moderately developed. vittatus
Antenna smooth; setae of ventral brush single or bifid ; comb spines with
submedian teeth strongly developed, often appearing trifid amaltheus (in part)
Comb consisting of elongated scales fringed around the apex . africanus,
pseudoafricanus, luteocephalus
Comb consisting of spines with the median denticle more strongly developed
than the lateral denticles ‘ 8
Comb spines with a small number of rather coarse lateral denticles of which
the submedian are usually visible under low powers . ; 9
Comb spines with a fringe of more delicate denticles on either side of the
main median denticle, these lateral denticles usually invisible under low
powers and often almost invisible under high é : 19
Pecten spines very deeply cleft, typical spines with two or more » large
median denticles subequal in size; saddle hair and subventral seta of
siphon single , : subargenteus
Pecten spines otherwise ; saddle hair usually and ‘subventral seta always
with at least two branches , ‘ 10
Pecten teeth with poorly developed secondary denticles invisible ‘under
low powers of the microscope : II
Pecten teeth with well-developed secondary denticles visible under low
powers of the microscope : ‘ : : ; é eta:
Lower caudal seta with 2 or more branches ae : < ; ee
Lower caudal seta single . ; 13
Typical pecten teeth short and broad, ‘not more than 4xas long as s their
breadth at base. : . stmpsoni, demerllont
Typical pecten teeth longer and narrower, ‘at least 6xas long as their
breadth at base. ‘ ‘ : : . strelitziae
Spines at bases of pleural setae large and curved . ‘ j ‘ heischt
These spines smaller . , é . keniensis
Distal edge of saddle with a group of c coarse ‘spicules near the base of the
caudal setae, easily visible under low powers; antennal seta with 2-4
branches ; : . amaltheus (in part)
Distal edge of saddle smooth or at most with very minute spicules not or
scarcely visible under low powers ; antennal seta single ; ‘ 15
Saddle hair with at least 3 branches; pleural spines very large and con-
spicuous : ; . metallicus
Saddle hair with at most 2 branches; pleural spines smaller ‘ ; oe
Submedian denticles of comb spines strongly developed, giving the
majority of these spines a trifid appearance under low powers. . ge Sy
Submedian denticles not so strongly developed, only an occasional spine
appearing trifid under low powers . ‘ : ‘ ‘ ; . 18
Pecten with 8-12 spines. . ; : : : ‘ ; mascarensis
Pecten with 14-20 spines . ‘ : : : i ; : aegypti
IN THE ETHIOPIAN REGION 249
18. Upper caudal seta with about 6 branches; saddle hair with 3-4. grants
Upper caudal seta with 2-4 branches ; saddle hair bifid . ‘ . contiguus
Ig. Pecten spines arranged in discontinuous groups . ; bambusae kenyae
Pecten spines more or less equally spaced, at most with one or two
terminal spines detached : 20
20. Pecten spines with at least one secondary denticle near the base easily
visible under low powers : aI
Pecten spines often almost simple, secondary denticles, when present,
visible only under high power . : . bambusae, dendrophilus, deboert
21. Siphon more or less uniformly dark. j : .. R2
Siphon dark only on about the basal ths, apical } 1th pale : : 23
22. Subventral seta of siphon with 3-4 branches ; antennal seta situated at not
more than ;jths the distance from base to apex; saddle hair bifid or
trifid . ‘ ; . calceatus
Subventral seta of siphon ‘bifid: antennal seta at more than ;jths the
distance from base; saddle hair single or bifid apzcoargenteus op: denderensis
23. Median denticle of comb spines much reduced, lateral denticles well
developed forming a fringe which is visible under low power 5 frasert
Median denticle strongly developed in relation to the fringe which is
invisible under low power . apicoargenteus, schwetzi, langata, soleatus
DISTRIBUTION RECORDS
The list which follows is believed to contain all published records except those
which are merely repetitive and do not indicate that a species has been found more
than once in a particular locality but merely that the same record has been published
on more than one occasion. Doubtful records are marked with an asterisk and are
discussed in a separate section. Records clearly based on misidentifications are
omitted and are discussed in another section. Some of the records are based on
unpublished specimens in various collections. For these collections the following
abbreviations are employed. B.M. British Museum (Natural History) ; Berl. Berlin
Museum; H.D. Hope Department, Oxford; H.I. Tropical Institute, Hamburg ;
L.S.H. London School of Hygiene and Tropical Medicine; Liv. Liverpool School of
Tropical Medicine ; M.L. Entomological Laboratory, Morogoro, Tanganyika. Records
from the Washington Museum were kindly supplied by Dr. Stone. These are marked
W.M. Allreferences are listed in full in the bibliography. Addresses of correspondents
who have sent records are given in the acknowledgements.
Aédes apicoargenieus
SIERRA LEONE. Daru (Simpson, 1913 ; Evans, 1925), Batkanu, Koinadugu (Butler
et al., 1915), Mabang (Gordon, 1929), Moyamba, Pujehun (Davey, MS.). LIBERIA.
Du River (Bequaert, 1930). Gorp Coast. Kumasi, Obuasi (Theobald, 1909), Lorha
(Carter, 1916), Nsawam (Macfie & Ingram, 1923a), Accra (Macfie & Ingram, 19236),
Aburi, Sunyani (Edwards, 1941), Asuboi (Mattingly, 1947). ToGoLAND. Klein Popo
(= Anecho, as Aniella togoensis, Enderlein, 1923). NIGERIA. Lagos (Wesché, 1910;
ENTOM. II. 5 Hh
250 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
Dunn, 1927); Philip, 1931), Yaba (Graham, 1911; Dunn, 1926; Mattingly, 1947),
Bende (Simpson, 19120), Kaduna*, Katagum*, Zungeru* (Johnson, 1919), Aba,
Ibadan, Ikeja (Mattingly, 1947), Itowolo (Mattingly, 19490), Agbor (L.S.H.). Br.
CAMEROONS. Misselele (Zumpt, 1937). FR. CAMEROONS. Kribi or Wuri (as Kingia
maculoabdominalis, Enderlein, 1923), Yaoundé (B.M.). BELGIAN Conco. Unnamed
locality (as Kingia albertit, Theobald, 1912), Kayembe Mukulu (Rodhain & Bequaret,
1913), Kabinda (Edwards, 1926), Leopoldville (Duren, 1929), Stanleyville (Schwetz,
19304; Liégeois, 1944), Lomami River between Obenge-Benge and Isangi (Schwetz,
19306), Eala, Kisantu, Ponthierville (Edwards, 1941), Mongbwalu (Schwetz, 1944),
Aba, Aketi, Ango, Basoko, Bayenga, Bondo, Buta, Irumu, Kimvula, Libenge, Nian-
gara, Titule, Yangambi (Liégeois, 1944), Coquilhatville (Wolfs, 1946), Maluku, Nta,
Pongwema (Fain & Henrard, 1948), Kimbao (Edwards,.MS.), Luluabourg (B.M.),
Katompe (Liv.). RUANDA-URUNDI. Kisenyi* (Seydel, 1929a). UGANDA. Unnamed
locality (Neave, 1912), Entebbe (Bequaert, 1930), Kilembe (Edwards & Gibbins,
1939), Chagwe, Kampala (Edwards, 1941), Bundibugiyo (Haddow, 1945a), Bubuk-
wanga (Haddow, 19450), Mongiro (Smithburn & Haddow, 1946), Mamirimiri (Had-
dow, 1947), Zika (Entebbe area, Haddow & Dick, 1948), Budonga and Siba Forest,
Bugala I., Bugoma Forest, Bujuko Forest, Busi I., East and West Busoga Woodlands
(Jinja and Busia areas), Buvuma I., Bwamba Lowlands and North Spur Ruwenzori
to 5,000 ft., Fort Portal, Itwara Forest, Kalinzu Forest, Kasyoha and Kitomi Forests,
Kiansonzi, Kibale-Hima Forest, Kitgum, Kome I., Kyenjojo, Masaka, Masindi
Port, Mubende, Mukono, Sango Bay Forests (Kaiso, Malambigambo, Namalala and
Toro) (Haddow, in litt.), Mpumu, Namanve Payida (B.M.). SuDAN. Khor Kobkwa
( = Khor Kobwa, as Kingia maculoabdominalis, Theobald, 1913). KENYA. Kaimosi
Forest, Kisii (Garnham eé al., 1946), Kakamega (E. C. C. Van Someren, 1946),
Kabwoch, Sondu River (E. C. C. Van Someren, in litt.). TANGANYIKA. Bukoba
(McHardy, 1932; Harris, 1942).
Aédes fraseri
FRENCH GUINEA. Dubreka (B.M.). SIERRA LEONE. Freetown (as fraseri, Bacot,
1916; as frasert and blacklocki, Evans, 1926), Daru (as blacklockt, Evans, 1925), Njala
(Edwards, 1941), Mabang, Moyamba Town (Liv.). GoLp Coast. Takoradi (Edwards,
1941; Mattingly, 1947). NicEr1A. Unnamed locality in the south (Edwards, 1917),
Oshogbo (L.S.H.). UGanpa. Unnamed locality (Neave, 1912), Mpumu (Edwards,
1912), Kampala (Hancock, 1930), Mamirimiri, Mongiro (Haddow, 1947), Bubuk-
wanga (Smithburn eé al., 1949), Bukakata, Bwamba lowlands, and North Spur
Ruwenzori to 5,000 ft., Bwigi I., Masindi Port (Haddow in litt.). Kenya. Kaimosi
Forest, Kakamega (E. C. C. Van Someren, 1946), Sondu River (E. C. C. Van Someren
in litt.).
Aédes dendrophilus
SIERRA LEONE. Freetown (as fraseri, Edwards, 1917; as dendrophilus, Edwards,
1921a), Kissy (Liv.). Gortp Coast. Aburi, Nsawam, Oblogo (Edwards, 19214),
Dodowah (Macfie & Ingram, 1923). NicER1IA. Lagos (Philip, 1931), Ibadan
(Edwards, 1941). FERNANDO Po. Santa Isabel (as trinidad, Gil Collado, 1936).
IN THE ETHIOPIAN REGION 251
UGANDA. Bwamba County (as deboert demeilloni, E. C. C. Van Someren, 1946),
Mongiro (as deboeri demeilloni, Smithburn & Haddow, 1946), Mamirimiri (as deboert
demeilloni, Haddow, 1947), Bubukwanga (Smithburn é al., 1949), Bugoma Forest,
Bujuko Forest, Busi I., Bwamba lowlands to 5,000 ft., Kitgum (fide Harper) (Haddow
in litt.). Kenya. Kaimosi Forest (as deboeri demeilloni, E. C. C. Van Someren, 1946).
ZULULAND. Eshowe (as deboeri demeilloni, Edwards, 1936). NATAL. Isipingo Beach,
Margate (B.M.), Amanzimtoti (Muspratt in litt.). CAPE Province. Embotyi
(Muspratt in litt.).
Aédes africanus
FRENCH GUINEA. Kindia (Wilbert & Delorme, 1927). SIERRA LEONE. Freetown
(Theobald, 1901 ; Evans, 1926; Davey, 1933), Koinadugu (Butler e¢ al., 1915), Daru
(Evans, 1925), Mabang (Gordon, 1929), unnamed localities (Davey, 1939), Moyamba,
Port Loko, Pujehun (Davey, MS.), Segbwema (Liv.), Bo (B.M.). LiBerta. Kaka
Town (Bequaert, 1930), Roberts Field, Kakata (Briscoe, 1947). GoLp Coast.
Ashanti (Macfie & Ingram, 1916b), Tamale, Yape (Simpson, 1918), Broomassie*
(Liv.). DAHomMEy. Bismarckburg (as Antella africana, Enderlein, 1923). NIGERIA.
Old Calabar (Theobald, 1901), Bonny, Duke Town (Theobald in Annett e¢ al., Igor),
Lagos (Theobald, rg01; Wesché, 1910; Connal, 1916; Dalziel, 1920; Dunn, 19270;
Philip, 1931; Kerr, 1933), Oshogbo (Mayer, 1911), Yaba (Graham, 1911; Dalziel,
1920 ; Dunn, 1926; Philip, 1929 ; Mattingly, 1947), Bende, Forcados (Simpson, 19120),
Opobo (Connal, 1915), Afaka, Kaduna, Ubassa (Johnston, 1916), Brass, Ikom
(Connal, 1917), Ikoyi (Dunn, 1926; Kerr, 1933), Ibadan (Kumm, 1931a@; Kerr, 1933),
Iwonran, Mushin (near Yaba, Kerr, 1933), Kateri (Edwards, 1941), Aba, Apapa,
Ikeja (Mattingly, 1947), Itowolo (Mattingly, 1949a), Ogbomosho (Bugher e¢ al., in
press), Sapele (L.S.H.). Fr. CAMEROoNS. Duala and unnamed localities (Griinberg,
1905), Yaoundé (B.M.). FR. EQUATORIAL AFRICA. Brazzaville (Sicé & Vaucel, 1928).
BELGIAN Conco. Ruwe (Theobald, 1907), Kabinda (Schwetz & Carter, 1915),
Elisabethville, R. Vakila (Schwetz, 1927a), Stanleyville (Schwetz, 1927); Liégeois,
1944), Tshela, Thysville (Schwetz, 1938), Coquilhatville, Lualaba River, Matadi
(Edwards, 1941), Ilambi (Schwetz, 1942), Aba, Aketi, Bondo, Buta, Inongo, Irumu,
Kimvula, Niangara, Titule, Yangambi, Yatolema (Liégeois, 1944), Balumi, Ban-
ningville, Black River, Dumi-Mato, Edudu, Pongwema, Tsunikitoko (Fain & Henrard,
1948), Costermansville, Lembwa River, Luluabourg (B.M.). RUANDA-URUNDI.
Dendezi (B.M.). UGANDA. Kampala (as Kingia africana, Theobald, 19114; as Aédes
africanus, Hancock, 1930), unnamed locality (= Chagwe, Mabira Forest, Neave,
1912), Bugomolo (Hancock, 1930), Kilembe (Edwards & Gibbins, 1939), Entebbe
(Edwards, 1941; Haddow & Dick, 1948), Bundibugiyo (Haddow, 1945a), Bubuk-
wanga (Haddow, 19450), Mongiro (Smithburn & Haddow, 1946), Mamirimiri
(Haddow, 1947), Kitinda, Zika (both near Entebbe, Smithburn eé al., 1948), Amag-
burugburu, Manjuguja North, Mbango (Smithburn e¢ al., 1949), Kyarumba (Haddow
& Van Someren, 1950), Dwaji I. (Gillett in litt.), Budonga & Siba Forests, Bugala I.,
Bugoma Forest, Bujuko Forest, Bukasa I., Busi I., East and West Busoga Wood-
lands (Jinja and Busia areas), Buvuma I., Bwamba Lowlands, and North Spur
Ruwenzori to 5,000 ft., Bwigi I., Fort Portal, Itwara Forest, Jana I., Kalinzu Forest,
252 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
Kasyoha and Kitomi Forests, Kiansonzi, Kibale-Hima Forest, Kiryandongo,
Kitgum, Kome I., Kyenjojo, Masaka, Masindi Port, Mityana, Mubende, Muhangi
Forest, Mukono, Nyamugasani Valley, Sango Bay Forests (Kaiso, Malambigambo,
Namalala and Toro) (Haddow in litt.), Godia, Kabarole, Kitasa, Midia, Payida
(B.M.), Jinja, Mboga Stream (Jinja—Mumias Road) (Liv.). Supan. Li Rangu, Sopo
(Lewis, 1945). ABYSSINIA. Sidamo Prov. (Bevan, 1937). KENyA. Chepalungu, Fort
Ternan, Kaimosi Forest, Kisii (Garnham eé al., 1946), Kabwoch, Kakamega,
Kibigori, Kisumu, Kitale, Koru, Lela, Maseno, Mugusi River, Muhuroni, Sondu
River (E. C. C. Van Someren in litt.). TANGANYIKA. Bukoba (McHardy, 1932; Harris,
1942), Ukara I.* (B.M.). Mozampigue. Unnamed localities* (Howard, 1912).
NYASALAND. Zomba* (Theobald, 1901). ANGOLA. Bihé (as Stegomyia africana and
Stegomyia dubia), Katemar (= Katema’s, Bihé area, as S. dubia) (Theobald, 1910),
Kibokolo do Zombo (Gamble, 1912). N. RuopeEs1A. Ndola (Robinson, 1950).
Aédes pseudoafricanus
NIGERIA. Yaba (as africanus, Graham, 1911), Lagos area (as africanus, Connal,
1915, &c.; Kerr, 1933; Edwards, 1941; as pseudoafricanus, Chwatt, 1949), Mushin
(near Yaba, as africanus, Kerr, 1933), Itowolo (as africanus, Mattingly, 1949a),
lado, Middle Point (both in Lagos area, Chwatt in litt.), Apapa, Gbohun—Gbohun,
Ikoyi, Onigbongbo (Lagos area) (B.M.).
Aédes simpsont
GAMBIA. Bathurst (Edwards, 1941). SIERRA LEONE. Freetown (Kennan, 1915;
Bacot, 1916; Evans, 1926), Daru (Blacklock, 1925), Kaballa (L.S.H.), Makeni, Port
Loko, Pujehun (Davey, MS.). Gorp Coast. Dodowa (Simpson, 1914), Accra (Macfie,
1917), Takoradi (Pomeroy, 1932), Aburi (Edwards, 1941), Asuboi, Kumasi (Mat-
tingly, 1947). NicErtaA. Afaka, Kudemsa (Johnston, 1916; Johnson, 1919), Yaba
(Dunn, 1926; Philip, 1929; Mattingly, 1947), Ibadan (Philip, 1931; Kumm, 19314;
Kerr, 1933), Oshogbo, Shaki (Philip, 1931), Lagos (Kerr, 1933), Gadau (Taylor,
1934), Ikeja, Kaduna, Port Harcourt (Mattingly, 1947), Ogbomosho (Bugher e¢ al.,
in press). Br. CAMEROONS. Kumba (fide Macnamara), Babango (L.S.H.). FR.
CAMEROONS. Yaoundé (B.M.). GABoon. Mayumba Forest (Galliard, 1931). BELGIAN
Conco. Kasongo, Tshumbiri, Wathen (Newstead ef al., 1907), Leopoldville (Newstead
et al., 1907; Duren, 1929), Matadi (Newstead ef al., 1907; Duren, 1938), Kivanda
Kapepula (Rodhain & Bequaert, 1913), Kabinda (Schwetz & Carter, 1915), Stanley-
ville (Schwetz, 1927); Liégeois, 1944), Katanga Prov. (Seydel, 1929b), Popokabaka
(Schwetz, 1938), Karimi (as Karumi, Uganda, Edwards, 1941), shores of Lake Kivu
(Schwetz, 1944), Costermansville (Schwetz, 1944; Wolfs in litt.), Aba, Aketi, Basoko,
Bondo, Buta, Irumu, Kimvula, Niangara, Titule, Yangambi, Yatolema (Liégeois,
1944), Coquilhatville (Wolfs, 1946), Kingangati, Kwamouth (Fain & Henrard, 1948),
Kimbao (Edwards in litt.). RuANDA-URuNDI. Usumbura (Edwards, 1941). UGANDA.
Kampala (as bromeliae, Theobald, 1g11a; as simpsont, Hancock, 1930), unnamed
locality (Neave, 1912), Kilembe (Edwards & Gibbins, 1939), Bundibugiyo (Haddow,
19454), Bubukwanga (Haddow, 19456), Bundinyama (Smithburn & Haddow, 1946),
Soroti (Hopkins, MS.), Budongo and Siba Forests, Bugala I., Bugoma Forest,
IN THE ETHIOPIAN REGION 253
Bujuko Forest, Bukakara, Bukasi I., Busi I., East and West Busoga Woodlands
(Jinja and Busia areas), Bwamba Lowlands and North Spur Ruwenzori to 5,000 ft.,
Entebbe, Fort Portal, Itwara Forest, Jubiya Forest, Kalinzu Forest, Kalungu,
Kasyoha and Kitomi Forests, Kiansonzi, Kibale-Hima Forest (= Mpanga Forest
auctt.), Kiryandongo, Kitgum, Kome I., Kyenjojo, Masaka, Masindi Port, Mityana,
Mubende, Muhangi Forest, Mukono, Nyamugasani Valley to 5,000 ft., Sango Bay
Forests (Kaiso, Malambigambo, Namalala, and Toro) (Haddowin litt.), Godia, Metu,
Midia, Rumogi (B.M.). SUDAN. Bor (as lilii, Theobald, 1910), Khartoum* (Whitfield,
1939), Juba (Edwards, 1941), Kau (Findlay, Kirk, & MacCallum, 1941), Heiban,
Kauda, Malakal, Meridi, Roseires, Sennar (Lewis, 1945), Torit (Knight in litt.).
AByssINIA. L. Hardin (Edwards, 1941). ERITREA. Nefasit (Lewis, 19430). KENYA
Digo, Eldoret*, Kiambu*, Meru*, Nairobi* (Symes, 1935), Kisumu (Symes, 1935;
Haddow, 1942), Kakamega, Kilifi (Symes, 1935; E. C. C. Van Someren in litt.).
Mombasa (Symes & Roberts, 1937; Wiseman ef al., 1931; Teesdale, 1941), Fort
Ternan, Kaimosi, Kisii (Garnham e¢ al., 1946), Gede (Bailey, 1947), Taveta (Heisch,
1948), Butere, Kerio Valley, Luanda, Makindu, Malindi, Msambweni, Tiwi, Yala
(E. C. C. Van Someren, in litt.). TANGANyIKA. Unnamed locality (Neave, 1912),
Dar-es-Salaam (Pomeroy, 1920; Edwards, 1923; McHardy, 1927; Harris, 1942),
Lindi, Tanga (Haworth, 1924), Moshi (McHardy, 1932), Mtibwa Forest (Harris,
1942), Morogoro (Harris, 1942; Hocking, 1947), Amani (Berl.), Mwanza (M.L.,
E. C. C. Van Someren in litt.). PeEmsBa I. Unnamed locality (Edwards, 1941).
_ ZANZIBAR. Various localities including Zanzibar Town (Aders, 19170). MOZAMBIQUE.
Mozambique, Nampula (De Meillon & Rebelo, 1941), Loureng¢o Marques, Umbeluzi
(Rebelo & Pereira, 1943), Busi R., Guara Guara, Mambone, Nhemancone, Quelimane
(W.M., fide Stone), Chindio (Liv.). NyAsALAND. Unnamed locality (Neave, 1912),
unnamed locality in uplands (Lamborn, 1930), Fort Johnston (Edwards, 1941), Fort
Maguire, Likoma I. (L.S.H.). N. Ruopesra. Livingstone (Muspratt, 1945), Balovale
(Robinson, 1948), Ndola (Robinson, 1950). ANGOLA. Bihé (Theobald, 1910), San
Salvador (Gamble, 1914). S. RHODESIA. Shamva, Umtali (Leeson, 1931), Salisbury
(Edwards, 1941), Bulawayo, Ndanga, Sinoia, Umtali, Victoria Falls (Meeser in litt.).
TRANSVAAL. Unnamed locality (Theobald, 1905), Abundant in low veldt (Bostock
& Simpson, 1905), Onderstepoort (Theobald, 1911), Roberts Heights (Pratt- Johnson,
1921), Leysdorp Road, Nelspruit (Ingram & De Meillon, 1929). ZULULAND. Empan-
geni (Ingram & De Meillon, 1927), Mhlatuze, Ntabanana (Bedford, 1928). NATAL.
Durban (Pratt-Johnson, 1921 ; Ingram & De Meillon, 1927 ; Bedford, 1928), Margate
(Muspratt 1950), Amanzimtoti (Muspratt in litt.). BECHUANALAND. Kasane (De
Meillon, 1947a). CAPE PRoviNcE. Mkanduli (De Meillon & Lavoipierre, 1944).
Aédes luteocephalus
GAMBIA. Bathurst (Edwards, 1941). FRENCH GUINEA. Dubreka (B.M.). SIERRA
LEONE. Freetown (Bacot, 1916; Evans, 1926; Davey, 1933), unnamed localities
(Davey, 1939), Bonthe, Makeni, Moyamba, Port Loko, Pujehun (Davey, MS.),
Kissy (Liv.), Aberdeen (L.S.H.). LiBerta. Roberts Field (Briscoe, 1947). GoLp
Coast. Accra (Macfie & Ingram, 1916a), Ashanti (Macfie & Ingram, 19168), Christians-
borg (Accra area, Ingram & Macfie, 1917), Tamale, Yape (Simpson, 1918), Nsawam
254 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
(Macfie & Ingram, 1923a), Saltpond (Edwards, 1941), Aburi (B.M.). NicEr1A. Bende
(Simpson, 1912b), Yaba (Simpson, 19120; Dunn, 1926; Kerr, 1933; Mattingly, 1947),
Lagos (Connal, 1915, &c.; Dalziel, 1920; Dunn, 1927a; Philip, 1931; Kerr, 1933),
Adun (Connal, 1917), Ibi (Connal, 1924), Ikoyi (Lagos area, Dunn, 1926; Kerr,
1933), Oshogbo (Kumm, 19310), Ibadan (Kumm, 1931a@; Kerr, 1933; Mattingly,
1947), Iwonran, Kano, Mushin (near Yaba) (Kerr, 1933), Gadau (Taylor, 1934),
Lokoja (Edwards, 1941), Ikeja (Mattingly, 1947), Ogbomosho (Bugher eé¢ al., in press),
Badagry (Liv.). BELGIAN Conco. Kumba* (Newstead e¢ al., 1907), Jadotville (De
Meillon, 1943), Kasongo (Liv.). UGANDA. Mpumu (as Kingia luteocephala, Theobald,
Ig10), unnamed locality (Neave, 1912), Mamirimiri (Haddow, Smithburn, eé¢ al.,
1948), Bwamba Lowlands and North Spur Ruwenzori to 5,000 ft. (Haddow in litt.),
Godia, Laufori, Metu, Rumogi (B.M.). SUDAN. Bor (Theobald, 1910), Juba (Hewer,
1934; Edwards, 1941), Khartoum* (Whitfield, 1939), Heiban (Lewis, 19432),
Danaglia, Malakal, Sennar, Wad el Magdub (Lewis, 1945), Jebel Marra (Abbott, 1948),
Singa (B.M.). ERITREA. Nefasit (Lewis, 19430). KENyA. Fort Hall, Kerio Valley,
Kisumu (E. C. C. Van Someren in litt.). N. RHoDEsIA. Balovale, Livingstone (Robin-
son, 1948), Ndola (B.M.). NyaAsALAND. Likoma I. (L.S.H.). S. RHopEstA. Victoria
Falls (Meeser in litt.). BECHUANALAND. Kasane, Maun (De Meillon, 19472).
Aédes unilineatus
GoLpD Coast. Accra (Macfie, 1916), Tamale* (Ingram, 1919), Aburi (Edwards,
1941). NiGcERIA. Kaduna, Katagum, Zungeru (Johnson, 1919), Ibadan (Philip,
1931), Gadau (Taylor, 1934), Baro, Lokoja (Edwards, 1941), Ogbomosho (Bugher
et al., in press). UGANDA. Metu, Rumogi (B.M.). SupAN. Bahr-el-Ghazal (Theobald,
1906), Jebelein (Theobald, 1910; Lewis, 1948), Erkowit, Juba (Edwards, 1941),
Heiban (Lewis, 1943a), El Fasher, Jebel Moya, Sennar, Tembura, Wad el Magdub,
Wad Medani (Lewis, 1945). KENyA. Unnamed locality (Neave, 1912), Dolo, Juba
River (Edwards, 1941). NyASALAND. Port Herald (Edwards, 1941). N. RHODESIA.
Livingstone (De Meillon & Lavoipierre, 1944). S. RHopEsiA. Fort Victoria (De
Meillon, 1943). TRANSVAAL. Letaba, Tzaneen (Ingram & De Meillon, 1929).
Aédes metallicus
SENEGAL. Dakar (Kartman eé al., 1947). GoLD Coast. Accra (Macfie, 1916),
Navarro (Ingram, 1919), Asuboi (Mattingly, 1947). NicER1A. Kano (Kumm, 19314),
Gadau (Taylor, 1934), Maiduguri (Mattingly, 1947). BrLG1AN Conco. Jadotville
(De Meillon, 1943). UGANDA. Mbarara (Edwards, 1941), Soroti (Haddow in litt.).
SUDAN. Bor (as dubia, Theobald, 1g10), Darfur Prov., Juba (Edwards, 1941), Nuba
Mts., Wad el Magdub (Lewis e¢ al., 1942), Heiban (Lewis, 1943a), Abu Usher, Dagu,
El Fasher, Kadugli, Kalora, Kassala, Khartoum, Kheiwok, Kudring, Malakal, Mug,
Nahud, Raga, Roseires, Sennar, Shaloat, Sheikh Gamal, Sheikh Karim, Wad Banda,
Wad Ganatir, Wad Medani (Lewis, 1945), Dueim, Kosti (Lewis, 1948), Yambio
(Abbott in litt.), Umm Sunt (B.M.). Kenya. Mombasa (Wiseman ef al., 1939;
Teesdale, 1941), Kisumu (Haddow, 1942), Fort Ternan (Garnham é al., 1946), Isiolo
(Heisch, 1947), Gede (Bailey, 1947), Garissa, Garsen, Gotani, Kakamega, Kerio
Valley, Kibos, Kilifi, Kipini, Koru, Langata Forest, Makindu, Malindi, Maseno,
IN THE ETHIOPIAN REGION 255
Msambweni, Taveta, Witu (E. C. C. Van Someren in litt.). TANGANYIKA. Dar-es-
Salaam (Edwards, 1923 ; Scott, 1926 ; McHardy, 1927), Tanga (Nixon, 1923 ; Haworth,
1924), Lindi (Haworth, 1924), Morogoro (Harris, 1942), ‘Widely distributed from
coast to Moshi and Mwanza’ (Harris, 1942), Mwanza (M.L., E. C. C. Van Someren in
litt.). ZANZIBAR. ‘Various localities’ (Aders, 19174), Zanzibar Town (Aders, 19170),
Mnazi Moja (= European Club, Zanzibar Town, Edwards, 1941). MOZAMBIQUE.
Nova Choupanga (Séguy, 1931). N. RHopEsiA. Livingstone (De Meillon & Lavoi-
pierre, 1944). ANGOLA. Bihé (as Quasistegomyia dubia, Theobald, 1910). S. RHODESIA.
Bindura, Gatooma, Shamva (Leeson, 1931), Fort Victoria, Odzi, Salisbury (Meeser
in litt.). TRANSVAAL. Komatipoort, Letaba (Ingram & De Meillon, 1929). ZULULAND.
Mhlatuze, Ntabanana (Bedford, 1928). BECHUANALAND. Francistown, Kasane,
Maun, Nata (De Meillon, 1947a). S. W. AFrRicA. Fransfontein, Grootfontein, Karabib,
Okahandja, Okimbahe, Omatjette, Otjimbingwe, Outjo, Usakos (Muspratt in litt.).
Aédes vittatus
GamBIA. Bathurst (Theobald, 1903). SIERRA LEONE. Freetown (Theobald, 1901 ;
Butler e al., 1915; Bacot, 1916; Evans, 1926; Davey, 1939), Batkanu, Benikoro,
Firiwa, Kaballa, Serakolia, Sonkonia, Tirikoro, Waliki (Simpson, 1913), Koinadugu
(Butler e¢ al., 1915), Daru (Edwards, 1941), Kamabai, Yiraia (B.M.), Boala Karafaia,
Kamasiki (L.S.H.), Bo, Kailahun, Makeni, Moyamba, Port Loko (Davey, MS.),
Dunkiawallia, Pujehun (Liv.). LiBertA. Monrovia (Kumm, 1931a@), Firestone
Plantation (Du Group) (Barber e¢ al., 1932), Roberts Field (Briscoe, 1947). Ivory
Coast. Bobo-Dioulasso (B.M.). GoLtp Coast. Bole (Ingram, 1912), Cape Coast,
Tamale (Simpson, 1914), Wa (Carter, 1916), Batiasan, Dogankade, Gambaga, Jefisi,
Kulmasa, Lilixia, Mayoro, Nacon, Nandaw, Nandawli, Nangudi, Pinna, Sambolugu,
Savelugu, Tanina, Tishi, Tumu, Ulu, Winduri, Yeji, Zouragu (Ingram, Ig19),
Sunyani (Edwards, 1941), Accra, Kintampo (Liv.). NIGERIA. Oshogbo (Mayer,
Ig11; Mattingly, 1947), Baro (Simpson, I912a), Zungeru (Simpson, 1912a; Johnson,
1919), Lokoja (Connal, 1915), Kakuri (Johnston, 1916), Katagum (Johnson, 1919),
Kaduna (Johnson, 1919, 1920), Gadau (Taylor, 1930), Abeokuta, Funtua, Ibi, Kano,
Katsina, Shaki, Yola (Kumm, 1931a@), Ibadan (Kumm, 1931@; Mattingly, 1947),
Ikoyi (Kerr, 1933), Ogbomosho (Bugher e# al., in press). FR. CAMEROONS. Yaoundé
(B.M.). BreLGian Conco. Kisui (as sugens, Newstead et al., 1907), Leopoldville,
Matadi (Newstead e¢ al., 1907; Duren, 1929), Elisabethville (Schwetz, 19272),
Stanleyville (Schwetz, 19276, 1930a; Liégeois, 1944), Boma (Duren, 1929), Kinshasa
(Bequaert, 1930), Lukula (Schwetz, 1938), Kaparata (Schwetz, 1944), Aba, Bondo,
Buta, Libenge, Niangara, Titule (Liégeois, 1944), Coquilhatville* (as sugens, Wolfs,
1946). UGANDA. Entebbe (Theobald, 1907), unnamed locality (Neave, I912), Sipi
(Hancock & Soundy, 1929; Hargreaves, 1932), Kyere, Mukongoro (Hancock, 1930),
Busi I., Fort Portal, Kitgum, Masindi Port, Soroti (Haddow in litt.), Ntotoro Valley
(Lumsden in litt.), Buvuma I. (Gillett in litt.), Chagwe, Jinja, Metu, Rumogi (B.M.).
SUDAN. Erkowit (Theobald 1910), Darfur Prov., Juba, Mongalla Prov. (Hewer, 1934),
Nuba Mts. (Hewer, 1934; Lewis, 1943a), Malakal (Hewer, 1934; Lewis, 1945),
Khartoum* (Bedford, 1938), Dar Shol, Delami, Kajo Kaji, Talodi (Edwards, 1941),
Jebel Meidob (Edwards, 1941 ; Lewis, 1945), Jebelein (Edwards, 1941; Lewis, 1948),
256 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
Wau (Edwards, 1941; Abbott in litt.), Damazin Cataract, Doka, El Obeid, Gedaref,
Geneina, Jebel Gurein, Jebel Mandera, Kassala, Kosti*, Roseires, Sennar, Umm
Ruaba (Lewis, 1945), El Fasher, Jebel Marra (Abbott, 1948), Rumbek, Yambio
(Abbott in litt.), Loka (Liv.). AraBria. Aden, Nobat (= Nobat Dakim), Sheikh
Othaman, Ulub (Patton, 1905), Wadiyain (as Wadi Yain, Edwards, 1941), Jebel Jihaf
(B.M.). Soxotra I. Hadibo, Kallansiya, Kathub (Leeson & Theodor, 1948). ABys-
SINIA. Harrar (as brumpti, Neveu Lemaire, 1905), L. Metahara area* (as Stegomyia
sp., Doreau, 1909), Meta Gafersa (La Face, 1939). ERITREA. Arisateb Valley, Tes-
senei, Ugaro (Lewis, 19430), Ras Malalu, east slopes of central massif, western plains
(Jannone et al., 1946), Mt. Ramlu at 3,000-4,000 ft. (B.M.). BRITISH SOMALILAND.
Buran (Edwards, 1941), Gedeis, Upper Sheikh (G. R. C. Van Someren, 1943), Las
Anod (B.M.). KENyA. Shimoni (as sugens, Edwards, 1914), Digo, Kisumu, Meru
(Symes, 1935), Mombasa (Symes, 1935 ; Wiseman é al., 1939), Nairobi (Symes, 1935 ;
H. I., fide Weyer, E. C. C. Van Someren in litt.), Fort Hall, Kakamega, Kilifi (Symes,
1935; E. C. C. Van Someren in litt.), Central Kavirondo (Symes & Roberts, 1938),
‘Third Camp’ (Edwards, 1941), Kaimosi Forest (Garnham é al., 1946), Isiolo
(Heisch, 1947), Kisumu (Grainger, 1947), Eldoret, Gotani, Kericho, Kiambu, Kisii,
Kitale, Kitui, Langata, Makindu, Malindi, Mara R., Maragua, Maseno, Narok,
Taveta, Thika, Turbo, Wesu (E. C. C. Van Someren in litt.), Malakisi (fide Leeson).
TANGANYIKA. Lindi (Haworth, 1924), Morogoro (Scott, 1926; Harris, 1942), Mwanza
(McHardy, 1932; Harris, 1942), ‘Widely distributed from coast to Bukoba’ (Harris,
1942), ‘All along shores of Mwanza Gulf and on Ukara Island’ (Smith in litt.), Dar-es-
Salaam (B.M.). PEmBa I. Unnamed locality (Edwards, 1941). ZANZIBAR. Zanzibar
Town (Aders, 1917a). MozAMBIQUE. Unnamed localities (Howard, 1912), Mocuba,
Quelimane (De Meillon & Rebelo, 1941), Namaacha (Rebelo & Pereira, 1943),
Ressano Garcia (W. M., fide Stone). NyASALAND. Uplands (Lamborn, 1930), Fort
Johnston (B.M.), ‘Abundant above 3,000 ft.’ (Lamborn, MS.). N. RHopesiA. Living-
stone (Robinson, 1948). ANGOLA. Bihé (Theobald, 1910). S. RHoDEsIA. Mashonaland
(Theobald, 1901), Gatooma, Salisbury, Shamva (Leeson, 1931), Bindura, Chibi, Mt.
Darwin, Fort Victoria, Goromonzi, Gwelo, Maranka Reserve, Matopo Hills, Msoneddi,
Ndanga, Triangle Estate, Umtali, Victoria Falls, Zaka (Meeser in litt.), Epworth
(L.S.H.). TRANSVAAL. Onderstepoort (Theobald, 1911b; Bedford, 1928), Brits
(Ingram & De Meillon, 1927), Yokeskei River (De Meillon, 1928), Komatipoort,
Letaba, Louis Trichardt, Nelspruit, Skukuza, Tomango, Waterpoort, Waterval
Boven (Ingram & De Meillon, 1929), Pietersburg (Liv.). ZULULAND, Begamuzi
Drift (Ingram & De Meillon, 1927), Ntabanana (Bedford, 1928). NATAL. Weenen
(Bedford, 1928), Margate (Muspratt in litt.). BECHUANALAND. Kasane (De Meillon,
1947a). S.W. AFricA. Karabib, Okimbahe, Otjimbingwe, Outjo (Muspratt in litt.).
Aédes aegypti
Apparently more or less throughout the region except at high altitudes and,
perhaps, in the extreme south and south-west. The following records will be found
to fill most of the gaps in the table given by Edwards (1941). CAPE PROVINCE.
Grahamstown, Port St. Johns (Edwards, 1941), Kimberley (De Meillon, 1942),
Upington (De Meillon, 1943). S.W. AFrica. Tsumeb, Windhoek (De Meillon, 1943),
IN THE ETHIOPIAN REGION 257
_ Fransfontein, Karabib, Okimbahe (Muspratt in litt.). BECHUANALAND. Kasane, Nata
(De Meillon, 1947a). N. RuopestA. Balovale, Livingstone (Robinson, 1948), Ndola
(Robinson, 1950). TANGANYIKA. ‘Throughout’ (Harris, 1942). SuDAN. Widespread
in Darfur Province (Abbott, 1948).
DOUBTFUL RECORDS
Aédes apicoargenteus
NIGERIA. Kaduna, Katagum, Zungeru. Johnson does not state from which of these
localities his specimens came. Katagum seems much too dry. RUANDA-URUNDI.
Kisenyi. No specimens are at present available for examination. In the distribution
maps (Figs. 4 and 8) the record is ascribed to ssp. denderensis.
Aédes africanus
GOLD Coast. Broomassie. The specimen in question is not marked with the locality
but was contained in a box with other specimens similarly labelled and known to
have come from Broomassie. There is nothing intrinsically unlikely about the record.
TANGANYIKA. Ukara Island. This record is based on a single larva and requires
confirmation. MozAMBIQUE and NYASALAND. No specimens can now be found by
means of which these interesting records could be confirmed or disproved. They are
discussed below under zoogeography.
Aédes simpsont
SuDAN. Khartoum. This record is based on a single specimen found in an aeroplane
and may be presumed to be the result of casual introduction. KENYA. Records given
by Symes (1935) from localities above about 5,000 feet have not been confirmed and
are thought to have been due to misidentification, although, in the absence of speci-
mens, this cannot be proved.
Aédes luteocephalus
BELGIAN Conco. Kumba. In his type description Newstead states that a single
specimen was taken at Kumba. This is clearly an error since the Liverpool School
collection contains two cotypes both marked by Newstead on the actual mounts as
taken at Kasongo. In his type description of Aédes (Dunnius) albomarginatus, in the
same paper, Newstead states that two specimens were taken at Kasongo, but the
Liverpool collection contains only a single holotype marked as taken at Kumba. It is
quite evident that Newstead confused the data relating to the two species in his
published descriptions and that the type locality of luteocephalus is Kasongo. The
point is of some importance since a record from Kumba would be entirely at variance
with what is known of the distribution of this species, while Kasongo is perfectly
acceptable. Edwards (1941) sought to equate Kumba with Tumba, but this appears
to have been quite gratuitous. Kumba is, in fact, a small village whose position is
clearly shown on Newstead’s map (see also under misquoted localities infra). The
reference by Theobald (1910) to a ‘type 3’ of luteocephalus in the British Museum
concerns a specimen of King’s from the Sudan apparently chosen by Theobald as a
lectallotype. SUDAN. Khartoum. This record is based on a single specimen found in
a plane and may be taken to represent a casual introduction.
ENTOM. II. 5 Ii
258 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
Aédes unilineatus
GoLpD Coast. Tamale. This record was based on larvae only and is queried by
Ingram. He was, however, familiar with the larva of unilineatus which he helped to
describe (Ingram and Macfie, 1917) and the record seems perfectly acceptable.
Aédes vittatus
BELGIAN CONGO. Coquilhatville. Wolfs quotes this record from a report of the
hygiene service at Coquilhatville for 1941. It is not supported by specimens and
vittatus has not been found again in Coquilhatville. SuDAN. Khartoum and Kosti.
Lewis (1945) considers that these were chance introductions. ABySssINIA. Lake
Metahara area. The species in question is stated to have been found breeding in
large numbers in rock-holes and it appears very probable that it was vittatus although
its identity cannot now be established.
RECORDS BASED ON MISIDENTIFICATIONS
The following records have been rejected.
Aédes frasert
SIERRA LEONE. Freetown (Edwards, 1917). = dendrophilus (see Edwards, 19214).
S. RHODEsIA. Unnamed localities (Garnham é al., 1946). = langata.
Aédes africanus
N. RHopEsIA. Unnamed locality (Edwards, 1912). = chaussieri from Lake Young
area. S. RHODESIA. Mashonaland (Theobald, 1901). = contiguus.
UNIDENTIFIED AND DOUBTFUL LOCALITIES
SIERRA LEONE. Dunkiawallia, Kamasiki, Waliki. Records of Aédes vittatus from
these localities are not thought to be of sufficient interest to merit an intensive search.
It is evident that this species is widely distributed throughout the territory. Kumm
(19310) takes Waliki to be the name of a place, but from Simpson’s account it seems
equally possible that it is the name of a river.
FR. CAMEROONS. Wuri (A. apicoargenteus). This is presumed to refer to some
locality on the Wuri River, probably at or near Duala.
BELGIAN ConcGo. Kimba (A. luteocephalus, Theobald, 1910a) and Tumba (A.
luteocephalus, Schwetz, 19276). These are clearly inaccurate renderings of the type
locality, Kumba, as given by Newstead e¢ al. (1907) (for a further discussion see above
under doubtful records). Lualaba River (A. africanus, Edwards, 1941). In its
present form this record is too vague to be of any value. No specimens are at present
available.
KENYA. ‘Third Camp’ (A. vittatus). The significance of this record could only be
ascertained by reference to relevant correspondence or records. These are not
available to the author. The specimen was collected by Dr. Chell in about 1973.
? NYASALAND. Zoruta (A. africanus, Theobald, 1910). This is presumed to be a
misspelling of Zomba from which Theobald (1901) has a previous record.
IN THE ETHIOPIAN REGION 259
ARABIA. Wadi Yain (A. vittatus, Edwards, 1941). The correct rendering should be
Wadiyain. Mr. Philby, who collected the specimens, kindly supplied this information
together with the approximate map reference given below.
DISTRIBUTION OUTSIDE THE ETHIOPIAN REGION
Aédes unilineatus. Punjab, Delhi Prov., Bombay area (Barraud, 1934).
Aédes vittatus. Corsica (Bigot, 1861, not since found there) ; Spain (Gil Collado,
1935; Clavero, 1946) ; India, Ceylon (Barraud, 1934) ; Cochin China, Annam (Borel,
1930) ; Algeria (Senevet, 1936).
Aédes aegypti. Throughout tropics and sub-tropics (Barraud, 1928; Kumm,
19310; Farner et al., 1946; Brug & Bonne-Wepster, 1947).
LIST OF LOCALITIES WITH TOPOGRAPHICAL DETAILS
The list is as accurate as it has been possible to make it from readily available
sources. It could undoubtedly be improved, especially by persons with local know-
ledge. Figures enclosed in brackets have been read from maps or, in the case of
rainfall figures, refer to an adjacent station. In the case of regions or provinces an
average figure is given and such figures are also enclosed in brackets. Where more
than one rainfall station is available for a particular locality an average figure is
given. Published figures and figures sent in litt. are unenclosed. Altitudes are given
in feet, where possible to the nearest 100 ft., otherwise to the nearest 500 ft. Rainfall
is given as the mean annual total to the nearest 5 inches and all references in the text
should be read in this sense. For many of the data included in the table the author
is indebted to correspondents. Aside from this the main sources have been Gossweiler
& Mendong¢a (1939), Grandidier (1934), Hurst & Black (1943), Nash (1948), Vanden-
plas (1943), The Times Gazetteer, the Atlas Général du Congo Belge, the Atlas do
Portugal Ultramarino, the Gold Coast Atlas, the Atlas of the Tanganyika Territory,
the East and South African Handbooks, the Boletim Mensal das Observacoes
Meteorologicas (Lourengo Marques), the annual rainfall summaries of the British
East African Meteorological Department, the climatological summaries of the
Rhodesia and Nyasaland Meteorological Service, the summary of rainfall normals
issued by the South African Department of Irrigation, and the handbooks and reports
of various government departments. Finally, where no published figures were
available, reference was made to the excellent large-scale maps of nearly every part
of Africa prepared during the last war for use by the forces and now available to the
public. Among these the Rainfall Map of Eastern Africa prepared by the East
African Survey Group (E. A. F. No. 1518 of 1943) deserves special mention.
List oF LOCALITIES
Locality | Altitude Latitude Longitude Rainfall
Aba, Belgian Congo ; , : : (3,500) 3.50 N. 30:13:15. (55)
Aba, Nigeria - : : 3 4 < 500 5.07 N. RD ik OF 85
Abeokuta, Nigeria . . ‘ ‘ ; < 500 7.09 N. 3.27 EB. 40
Aberdeen, Sierra Leone . = : : < 500 8.30 N. 13.18 W. < 100
260 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
List oF LOCALITIES—(cont.)
Locality Altitude Latitude Longitude Rainfall
Aburi, Gold Coast . 1,400 5.51 N 0.10 W. 40
Abu Usher, Sudan . (1,200) (14.54 N.) (33.14 E.) (z0)
Accra, Gold Coast . < 500 5.33 N 0.13 W. 25
Aden, Aden Prot. < 500 12.45 N 45.04 E. (5)
Adun, Nigeria < 500 5.59 N 8.09 E. (80)
Afaka, Nigeria (2,000) (10.41 N.) (7-23 E.) (50)
Agbor, Nigeria < 500 6.15 N 6.14 E. (80)
Aketi, Belgian Congo (1,500) 2.42 N 23.52 E. (65)
Amagburugburu, Uganda (3,000) (0.45 N.) (30.03 E.) (55)
Amani, Tanganyika 3,000 5.05 S 38.40 E. 75
Amanzimtoti, Natal < 500 30.03 S 30.53 E. (40)
Anecho, Togoland . < 500 6.15 N 1.39 E. (30)
Ango, Belgian Congo (2,500) 4.03 N 25.53 E. (75)
Apapa, Nigeria D < 500 (6.20 N.) (3.19 E.) (70)
Arisateb Valley, Eritrea . (2,000) (15.13 N.) (36.45 E.) (15)
Asuboi, Gold Coast. < 500 (5.46 N.) (0.22 W.) (50)
Babango, Br. Cameroons. (5,000) (6.20 N.) (10.30 E.) (70)
Badagry, Nigeria < 500 6.23 N 2.58 E. (60)
Bahr el Ghazal, Sudan (1,200) (9.30 N.) (30.00 E.) (30)
Balovale, N. Rhodesia 3,400 (13.34 S.) (23.07 E.) 40
Balumi, Belgian Congo . : (2,000) 4.108 15.50 E. (55)
Lake Bangweulu, N. Rhodesia . 3,800 11.208 30.00 E, (50)
Banningville, Belgian Ae I,100 3.195 ¥7-23-E: 50
Baro, Nigeria < 500 8.35 N. 6.18 E. (55)
Basoko, Belgian Congo (1,500) T.21-N. 23.40 E. (75)
Bathurst, Gambia . < 500 13.28 N 16.40 W. 45
Batiasan, Gold Coast (1,000) (10.43 N.) (1.44 W.) (40)
Batkanu, Sierra Leone (500) (9.05 N.) (12.27 W.) IIo
Bayenga, Belgian Congo . (1,500) 2.19 N 21.46 E. (70)
Begamuzi Drift, Zululand (3,000) (28.10 S.) (31.20 E.) (25)
Bende, Nigeria . < 500 5.32 N 7.36 E. (90)
Benikoro, Sierra Leone (1,500) (9.43 N.) (11.27 W.) (80)
Bihé, Angola (5,500) 12.18S 17.00 E. (70)
Bindura, S. Rhodesia 3,700 (I7:21'S:) (31.21 E.) (35)
Bismarckburg, Dahomey 2,300 8.12 N 0.51 E. 55
Mt. Bizen, Eritrea . : (5—8,000) (15.18 N.) (39.05 E.) (15)
Black River, Belgian Congo (1,500) 3.535 15.55 E. (55)
Bo, Sierra Leone - < 500 7.55 N 11.44 W. 120
Boala Karafaia, Sierra Leone ; (1,500) (9.39 N.) (11.04 W.) (80)
Bobo-Dioulasso, Ivory Coast 1,400 TT.12/N: 4.17 W 40
Bole, Gold Coast ; I,000 9.02 N 2.29 W (40)
Boma, Belgian Congo < 500 5.508 13.09 E * (35)
Bondo, Belgian Congo (1,500) 3.55 N 23.55 E (65)
Bonny, Nigeria < 500 4.25 N 7.13E > 100
Bonthe, Sierra Leone < 500 (7.33 N.) (12.37 W.) 150
Bor, Sudan I,700 6.12 N 31.33 E 35
Brass, Nigeria < 500 4.19 N 6.15 E > 100
Brazzaville, Fr. Eq. Africa (1,000) 4.158 15.20 E 60
Brits, Transvaal : 3,600 25.3858 27.47 E 25
Broomassie, Gold Coast . (500) 8.21 N 0.58 W (45)
Bubukwanga, Uganda (3,000) (0.45 N.) (30.03 E.) (55)
Budongo Forest, Uganda (3,500) (1.45 N.) (31.34 E.) (60)
Bugala I., Uganda . ; (3,800) (0.24 S.) (32.16 E.) (80)
Bugoma Forest, Uganda . (4,000) (1.20 N.) (31.00 E.) (55)
Bugomolo, Uganda (3,500) 1.31 N 31.27 E (55)
Bujuko Forest, Uganda . (4,000) 0.25 N 32.15 E (45)
IN THE ETHIOPIAN REGION 261
List oF LOcALITIES—(cont.)
Locality Altitude Latitude Longitude Rainfall
Bukakata, Uganda (4,000) 0.15 S. 32.02 E. 55
Bukasa I., Uganda (3,800) (0.26 S.) (32.30 E.) (75)
Bukoba, Tanganyika 3,800 ¥.20 5. 31.48 E. 80
Bulawayo, S. Rhodesia 4,400 20.09 S. 28.37 E. 25
Bundibugiyo, Uganda 3,600 0.42 N. 30.04 E. 55
Bundinyama, Uganda (3,000) (0.47 N.) (30.05 E.) (55)
Buran, Br. Somaliland 3,000 10.13 N. 48.47 E. 10
Busia, Uganda (4,000) (0.28 N.) (34.05 E.) (60)
Busi I., Uganda (3,800) (0.03 N.) (32.20 E.) (60)
Busi R., Mozambique < 500 (19.55 S.) (34.20 E.) (40)
Buta, Belgian Congo (1,200) 2.48 N. 24.46 E. 60
Butere, Kenya (4,500) 0.13 N. 34.30 E. (70)
Buvuma I., Uganda (3,800) o.11 N. 33.18: E. 65
Bwigi I., Uganda (3,800) (0.28 S.) (32.24 E.) (75)
Calabar, Nigeria. : < 500 4.57 N. 8.20 E. 125
Cape Coast, Gold Coast . < 500 5.00 N. 1.16 W. 25
Chagwe, Uganda (4,000) (0.25 N.) (33.00 E.) (55)
Chepalungu, Kenya 5,500 (1.00 S.) (35.15 E.) (35)
Chibi, S. Rhodesia . (4,000) 20.19 S. 30.25 E. (30)
Chindio, Mozambique. < 500 1739'S: 35.14 E. (35)
Christiansborg, Gold Coast. Sée Accra.
Coquilhatville, Belgian Congo . 1,200 0.04 N. 18.20 E. (70)
Costermansville, Belgian Congo 5,300 2:30:'5) 28.51 E. (55)
Dagu, Sudan . (2,000) (13.39 N.) (30.08 E.) (15)
Dakar, Senegal : < 500 14.40 N. 17.26 W. 20
Damazin Cataract, Sudan (1,000) (11.44 N.) (34.25 E.) (30)
Danagla, Sudan (1,300) (14.25 N.) (33.31 E.) (15)
Dar-es-Salaam, Tanganyika < 500 6.50 S. 39.17 E. 45
Darfur Prov., Sudan (2,500) (13.00 N.) (25.00 E.) (15)
Dar Shol, Sudan (2,500) 11.54 N. 30.28 E. (30)
Daru, Sierra Leone . : 500 (7.59 N.) (10.50 W.) go
Mt. Darwin, S. Rhodesia. 4,900 16.45 S. BT.30.be (35)
Delami, Sudan (2,300) (11.52 N.) (30.20 E.) (30)
Dendezi, Ruanda- Urundi (6,000) (2.28 S.) (29.03 E.) (55)
Digio, Kenya < 500 1227'S: 40.56 E. (35)
Diredawa, Abyssinia (4,000) 9.40 N. 41.59 E. (30)
Dodowa, Gold Coast ; (500) 5.53 N. 0.05 W. (40)
Dogankade, Gold Coast . (500) (8.42 N.) (0.29 W.) (45)
Doka, Sudan . (1,500) 13.30 N. 34.45 E. (25)
Dolo, Kenya . : (1,000) (4.08 N.) (41.48 E.) (10)
Du River, Liberia . (0-1, 500) (6.15 N.) (10.25 W.) > 100
Duala, Fr. Cameroons < 500 4.03 N. 9.41 E. 160
Dubreka, Ivory Coast < 500 9.48 N. 13.32 W. (170)
Dueim, Sudan 1,500 13.590 N. 32.20 E. ste)
Duke Town, Nigeria < 500 (4.55 N.) (8.19 E.) > 100
Dumi-Mato, Belgian Congo. (2,000) 4.30 S. 15.45 E. (50)
Dunkiawallia, Sierra Leone. Not identified.
Durban, Natal < 500 29.52 S. 31.03 E. 45
Dwaji I., Uganda (3,800) (0.00) (32.56 E.) (80)
Eala, Belgian Congo I,000 o.o1 N. 18.30 E. 70
Edudu, Belgian Congo (2,000) (3.10 S.) (16.30 E.) (60)
Eldoret, Kenya 6,900 0.32 N. 35.16 E. 40
El Fasher, Sudan : 2,800 13.38 N. 25.21 E. 10
Elisabethville, Belgian Congo 5 4,000 11.40 S. 27.34 E. 50
El Obeid, Sudan 2,200 13.51 N: 30.14 E. 15
Embotyi, Cape Province | < 500 (31.30 S.) (29.50 E.) (45)
262 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
List oF LocaLiTIES—(cont.)
Locality Altitude Latitude Longitude Rainfall
Empangeni, Zululand < 500 ' 28.468. 31.55 E. 40
Entebbe, Uganda 3,900 0.05 N. 32.29 E. 60
Enugu, Nigeria : 700 6.27 N. 7.29 E. 7°
Epworth, S. Rhodesia (5,000) (17.56 S.) (31.36 E.) (35)
Erkowit, Sudan. 4,300 18.46 N. 36.06 E. (5)
Eshowe, Zululand . : 1,700 28.53 S. 31.28 E. 60
Firestone Plantation, Liberia < 500 (6.18 N.) (10.25 W.) > 100
Firiwa, Sierra Leone (1,500) (9.20 N.) (11.18 W.) (80)
Forcados, Nigeria . < 500 5.20 N. 5.26 E. 130
Fort Hall, Kenya 4,100 0.42 S. 37.10 E, 55
Fort Johnston, Nyasaland 1,700 14.25 S. 35.16 E. 25
Fort Maguire, Nyasaland 1,600 13.40 S. 34.57 E. (30)
Fort Portal, Uganda 5,100 0.40 N. 30.17 E. 55
Fort Ternan, Kenya : 7,000 O.12'S. 35.20 E. 50
Fort Victoria, S. Rhodesia 3,700 20.04 S. 30.46 E. 25
Francistown, Bechuanaland 3,300 24.135, 27.30 E. 20
Fransfontein, S.W. Africa (3,500) (20.13 S.) (15.01 E.) (10)
Freetown, Sierra Leone < 500 8.30 N. 13.10 W. 160
Funtua, Nigeria (2,500) (11.32 N.) (7/T5°E:) (55)
Gadau, Nigeria (1,000) (11.50 N.) (10.12 E.) 30
Gambaga, Gold Coast I,300 10.33 N. 0.27 W. (45)
Garissa, Kenya (500) 0:27'S. 39.38 E. Io
Garsen, Kenya : < 500 2,10'9; 40.06 E, (30)
Gatooma, S. Rhodesia 3,800 18.20 S. 29.51 E. 30
Gbohun-Gbohun, ee See Lagos.
Gedaref, Sudan 2,400 14.02 N. 35.24 E. 25
Gede, Kenya . < 500 3.19 S. 40.01 E. 45
Gedeis, Br. Somaliland (3,500) (10.03 S.) (45.30 E.) (15)
Geneina, Sudan 3,100 13.29 N. 22.29 EB: 25
Godia, Uganda c (4,000) 3.25 N. 30.58 E. (60)
Goromonzi, S. Rhodesia . (4,800) (17.51 S.) (31.22 E.) (35)
Gotani, Kenya a: (1,000) 3.47 S. 39.32 E. (45)
Grahamstown, Cape Province : 1,800 33.18 S. 26.32 E. 30
Grootfontein, S.W. Africa (4,000) 19.31 S. 18.06 E. 20
Guara Guara, Mozambique < 500 (25.00 S.) (32.30 E.) (25)
Gwelo, S. Rhodesia 4,700 19.28 S. 29.48 E. 25
Hadibo, Sokotra I. , < 500 (12.37 N.) (54.05 E.) (15)
Lake Hardin, Abyssinia . (3,000) (8.52 N.) (39.50 E.) (40)
Harrar, Abyssinia . 5,500 9.22 N. 42.02 E. 35
Heiban, Sudan 3,400 11.05 N. 30.05 E. 35
Ibadan, Nigeria 700 7.23 N. 3.50 E. 50
Ibi, Nigeria < 500 8.11 N. 9.43 E. (45)
Ikeja, Nigeria < 500 (6.29 N.) (3.14 E.) (60)
Ikom, Nigeria 3 ; < 500 6.00 N. 8.40 E. (90)
Ikoyi, Nigeria. See Lagos
Ilado, Nigeria. See Lagos.
Ilambi, Belgian Congo (1,500) 0.34 N. 24.15 E. 80
Inongo, Belgian Congo I,100 1.55 S. 18.30 E. 65
Irumu, Belgian Congo (3,000) 1.25 N. 30.00 E. 50
Isangi, Belgian Congo (1,000) 0.42 N. 24.15 E. (80)
Isiolo, Kenya (4,000) 0.22 N. 37-35 E. 20
Isipingo, Natal < 500 30.00 S. 30.55 E. (40)
Itowolo, Nigeria < 500 (6.28 N.) (3.27 .) (60)
Itwara Forest, Uganda : (5,000) (0.45 N.) (30.25 E.) (55)
Iwonran, Nigeria. See Lagos.
Jadotville, Belgian Congo (4,300) 11.00 S. 26.44 E. (50)
IN THE ETHIOPIAN REGION 263
List oF LocALiTies—(cont.)
Locality Altitude Latitude Longitude Rainfall
Jano I., Uganda (3,800) (0.15 S.) (32.36 E) (75)
Jebelein, Sudan. 1,500 12.37-N: 32.50 E. (15)
Jebel Gurein, Sudan (1,400) (13.39 N.) (34.41 E.) (25)
Jebel Jihaf, Aden Prot. 7,100 (13.45 N.) (44.40 E.) (10)
Jebel Mandera, Sudan (1,300) (14.25 N.) (33-31 E.) (15)
Jebel Marra, Sudan (5-12,000) 13.00 N. 24.20 E. 25
Jebel Meidob, Sudan (2,500) (15.18 N.) (26.25 E.) sae)
Jebel Moya, Sudan (1,500) (13.28 N.) (33.22 E.) (15)
Jefisi, Gold Coast (1,000) 10.45 N. 2.16 W. (40)
Jinja, Ugana . 3,900 (0.26 N.) (33.11 E.) 45
Juba, Sudan . ? 1,800 4.51 N. 31.37 E. 35
Jubiya Forest, Uganda (4,000) (0.15 S.) (31.59 E.) (55)
Kaballa, Sierra Leone. (1,500) 9.39 N.. 11.41 W. 80
Kabarole, Uganda. See Fort Portal.
Kabinda, Belgian Congo . 3,100 6.08 S. 24.30 E. 65
Kabwoch, Kenya (4,000) - 0.45 S. 34.27 E. (65)
Kadugli, Sudan 2,000 11.00 N. 29.43 E. 30
Kaduna, Nigeria 1,900 10.30 N. 7.28 E. 45
Kailahun, Sierra Leone (1,000) (8.16 N.) (10.36 W.) 120
Kaimosi, Kenya 5,500 0.05 N. 34.50 E. 70
Kajiado, Kenya (6,000) Te5ELSs 36.47 E. (20)
Kajo Kaji, Sudan . 4,100 3.53 N. 31.40 E 50
Kaka Town, Liberia (1,000) 6.34 N. 10.20 W 100
Kakamega, Kenya . 5,100 (0.14 N.) (34.44 E.) 70
Kakata, Liberia (1,000) (6.30 N.) (10.25 W.) 100
Kakuri, Nigeria ‘ (1,500) (10.26 N.) (7.26 E.) (45)
Kalinzu Forest, Uganda . (4,000) (0.25 S.) (30.05 E.) (55)
Kallansiya, Sokotra I. < 500 (12.39 N.) (53.38 E.) (15)
Kalora, Sudan (4,000) (11.55 N.) (31.18 E.) (25)
Kalungu, Uganda . (4,000) (0.20 S.) (31.46 E.) (45)
Kamabai, Sierra Leone . (1,500) (9.20 N.) (11.58 W.) (100)
Kamasiki, Sierra Leone. Unidentified.
Kampala, Uganda . 3,900 0.19 N. 32.35 E 45
Kano, Nigeria I,500 12.02 N. 8.32 E 35
Kaparata, Belgian Congo 3,800 (0.25 N.) (29.33 E.) (60)
Karabib, S.W. Africa 3,800 (21.55 S.) (15.40 E.) 5
Karimi, Belgian Congo (4,500) (0.07 N.) (29.47 E.) (55)
Kasane, Bechuanaland (3,000) 17.49 S. 25.09 E 20
Kasongo, Belgian Congo . (2,000) A275: 26.39 E (50)
Kasyoha Forest, Uganda. (4,000) (0.20 S.) (30.10 E.) (55)
Kassala, Sudan ; 2,000 15.28 N. 36.24 E 15
Katagum, Nigeria . ; (1,000) 12.18 N. 10.20 E 20
Kateri, Nigeria ‘ 1,700 (9.37 N.) (7.25 E.) (60)
Kathub, Sokotra I. < 500 (12.36 N.) (54.00 E.) (15)
Katompe, Belgian Congo (2,500) 6.06 S. 26.24 E (40)
Katsina, Nigeria 1,800 13.01 N. 7.30 E 35
Kau, Sudan . (1,000) (10.40 N.) (31.30 E.) (25)
Kauda, Sudan ; (2,500) (11.01 N.) (30.33 E.) (35)
Kayembe-Mukulu, Belgian Congo (2,500) (9.30 S.) (25.36 E.) (50)
Kericho, Kenya 6,400 0.23 S. 35.17E 65
Kerio, Kenya . (4,000) 0.38 N. 35.37 E (45)
Khartoum, Sudan . I,500 15.35 N. 32.35 E 5
Kheiwok, Sudan . (2,000) ~ 12.42 N. 29.09 E (15)
Khor Kobwa, Sudan (2,500) (4.02 N.) (30.35 E.) (55)
Kiambu, Kenya 6,300 Lit3:5 36.52 E 40
Kiansonzi, Uganda (4,000) (0.14 N.) (31.45 E.) (35)
264 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
List oF LOCALITIES—(cont.)
Locality Altitude Latitude Longitude Rainfall
Kibale Forest, Uganda (4,500) (0.30 N.) (30.26 E.) (50)
Kibigori, Kenya. (4,000) 0.05 S. 35.03 E. (65)
Kibokolo do Zombo, Angola 3,800 6.22 S. 15.17 E. 50
Kibos, Kenya 3,800 0.04 S. 34.51 E. 45
Kilembe, Uganda 4,500 (0.25 N.) (29.59 E.) (60)
Kilifi, Kenya , < 500 3337-5. 39.52 E. 40
Kimbao, Belgian Congo ; (3,500) (5.30 S.) (17.30 E.) 65
Kimberley, Cape Province 4,000 28.42 S. 24.59 E. 15
Kimvula, Belgian Congo . (2,500) (5.44 S.) (16.04 E.) (50)
Kindia, Fr. Guinea . 1,400 10,03 N. 12.50 W. 80
Kingangati, Belgian Congo : (2,500) (4.22 S.) (15.50 E.) (50)
Kinshasa, Belgian Congo. See Leopoldville
Kintampo, Gold Coast I,200 8.04 N. 1.42 W. (55)
Kipini, Kenya < 500 2.32 8. 40.32 E. (45)
Kiryandongo, Uganda 4,000 1.53 N. 32.03 E. 50
Kisantu, Belgian Congo . 1,900 5.07 S. 15.07 E. (50)
Kisenyi, Ruanda-Urundi 4,800 1.308. 29.10 E. 50
Kisii, Kenya . ; 5,700 0.41 S. 34.47 E. 65
Kissy, Sierra Leone < 500 (8.29 N.) (13.12 W.) > 100
Kisui, Belgian Congo (1,500) (0.22 S.) (25.38 E.) (70)
Kisumu, Kenya 3,800 0.06 S. 34.45 E. 45
Kitale, Kenya , 6,200 1.02 N. 35.00 E. 45
Kitasa, Uganda. See Entebbe.
Kitgum, Uganda . 3,600 3.20 N. 32.53 E. 50
Kitinda, Uganda. See Entebbe.
Kitui, Kenya ‘ ; 3,900 121;5: 38.01 E. 30
Kivanda-Kapepulu, Belgian Congo : (4,000) (10.00 S.) (24.00 E.) (50)
Koinadugu, Sierra Leone (1,500) (9.30 N.) (11.25 W.) (80)
Komatipoort, Transvaal . 500 25.26 S. 31.57 E. 25
Kome I., Uganda (3,800) 0.05 S. 32.45 E. (80)
Koru, Kenya 5,900 OTS: 35.16 E. 60
Kosti, Sudan 1,500 13.10 N 32.40 E. I5
Kribi, Fr. Cameroons < 500 2.59 N 9.56 E. IIo
Kudemsa, Nigeria . (2,000) (10.23 N.) (7.33 E.) (45)
Kudring, Sudan. (2,700) (11.39 N (30.32 E.) (35)
Kulmasa, Gold Coast (1,000) (9.42 N.) (2.25 W.) (40)
Kumasi, Gold Coast (800) 6.43 N 1.37 W. 7°
Kumba, Belgian Congo (1,500) (1.50 S:) (25.50 E.) (70)
Kumba, Br. Cameroons . 800 (4.40 N.) (9.35 E.) > 100
Kwamouth, Belgian Congo 1,600 3.12'9 16.15 E. 60
Kyarumba, Uganda (5,500) 0.09 N 29.58 E. (50)
Kyenjojo, Uganda 4,800 0.36 N 30.39 E. 45
Kyere, Uganda 3,500 1.29 N 33.37-E. 50
Lado, Sudan . (1,500) (5.01 N.) (31.43 E.) (35)
Lagos, Nigeria < 500 6.20 N 3.20 E. 70
Langata Forest, Kenya (5,500) r:20'S 36.45 E. (35)
Las Anod, Br. Somaliland (2,500) (8.31 N.) (47.26 E.) (5)
Laufori, Uganda (2,500) 3.36 N 31.35 E. (50)
Lela, Kenya . (4,500) 0.02 S 34.36 E. (65)
Lembwa River, Belgian Congo (4,000) 6.01 S 17.06 E. (60)
Leopoldville, Belgian Congo 3 I,100 4.208 15.18 E. 55
Letaba, Transvaal P ‘ 1,800 23.52S 30.18 E. (25)
Leysdorp Road, Transvaal (2,000) (24.00 S.) (30.22 E.) 30
Libenge, Belgian Congo . I,300 3.40 N 18.39 E. 55
Likoma I., Nyasaland 1,600 12.04 S 34.44 E. (75)
Lilixia, Gold Coast . (1,000) 10.46 N 2.06 W. (40)
IN THE ETHIOPIAN REGION 265
List oF LOCALITIES—(cont.)
Locality Altitude Latitude Longitude Rainfall
Lindi, Tanganyika . < 500 10.00 S. 39.42 E. 35
Li Rangu, Sudan (2,000) (4.40 N.) (28.21 E.) (55)
Livingstone, N. Rhodesia 3,200 17.50 S. 25.49 E. 25
Loka, Sudan i (1,000) 4.14 N. 31.01 E. (50)
Lokoja, Nigeria < 500 7.48 N. 6.44 E. 50
Londiani, Kenya . 7,500 0.10 S. 35.30 E. 45
Lorha, Gold Coast . ‘ < 500 10.39 N. 2.52 W. (45)
Louis Trichardt, Transvaal 3,200 23.03 S. 29.54 E. 30
Lourengo Marques, Mozambique < 500 25.58 S. 32.36 E. 25
Luanda, Kenya (4,500) 0.02 N. 34.34 E. 65
Lukula, Belgian Congo < 500 522 5. 12.59 E. 45
Luluabourg, Belgian ial 2,100 5.50 S. 22.198 Es. 65
Lumbwa, Kenya 6,300 0.12 S. 35.29 E. 50
Mabang, Sierra Leone < 500 (8.22 N.) (12.52 W.) (130)
Mabira Forest, Uganda (4,000) (0.28 N.) (32.58 E.) (55)
Maiduguri, Nigeria . 1,200 11.49 N. 13.09 E. 20
Makeni, Sierra Leone < 500 (8.55 N.) (12.05 W.) 120
Makindu, Kenya 3,300 21795 37.49 E. 25
Malakal, Sudan I,500 9.32 N. 91.30 E: 30
Malakisi, Kenya (4,000) (0.25 N.) (34.20 E.) (70)
Malindi, Kenya < 500 3.179: 40.07 E. 40
Maluku, Belgian Congo (1,000) 4.04 S. 15.33 E. (55)
Mambone, Mozambique . < 500 20.58 S. 35.00 E. (35)
Mambrui, Kenya < 500 3.07 S. 40.09 E. (40)
Mamirimiri, Uganda (3,000) (0.48 N.) (30.09 E.) (55)
Manjuguja North, Uganda (3,000) (0.45 N.) (30.03 E.) (55)
Mara River, Kenya (6,500) 1.079: 35.08 E. (45)
Maragua, Kenya . ‘ (4,000) 0.47 S. 37.071. (35)
Maranka Reserve, S. Rhodesia (4,000) (19.15 S.) (32.15 E.) (35)
Margate, Natal ‘ < 500 (30.51 S.) (30.22 E.) (45)
Masaka, Uganda 4,300 0.20 S. 31.44 E. 45
Maseno, Kenya ‘ r - : x 6,000 0.01 S. 34.36 E. 50
Masindi Port, Uganda . F Z > 3,400 1.42 N 32.05 E. 40
Matadi, Belgian Congo . < 500 5.50S 13:32 E: 45
Matopo Hills, S. Rhodesia (4,000—5,000) 20.40S 28.30 E. (25)
Maun, Bechuanaland P ‘ ‘ (3,000) 19.59S 23.25 E. (15)
Mayoro, Gold Coast ve i Fi (500) (10.56 N.) (1.04 W.) (40)
Mayumba Forest, Gaboon . < 500 3:24 5 10.37 E. (60)
Mbango, Uganda (3,000) (0.45 N.) (30.03 E.) (55)
Mbarara, Uganda ‘ : : 4,700 0.37S 30.39 E. 35
Meridi, Sudan . - ‘ ; a 2,900 4.52N 29.28 E. 55
Meru, Kenya . . 6,200 0.03 N 37-39 E. 60
Meta Gafersa, Abyssinia . (5,000) (4.50 N.) (38.45 E.) (30)
Lake Metahara, Abyssinia (3,000) 8.52 N 39.51 E. (40)
Metu, Uganda : (3,000) 3.39 N 31.47 E. (50)
Mhlatuze Settlement, Zululand (1,000) (28.40 S.) (31.45 E.) 55
Middle Point, Nigeria. See ee
Midia, Uganda (4,000) 3.27 N 31.03 E. (50)
Misselele, Br. Cameroons < 500 (4.06 N.) (9.25 E.) 85
Mityana, Uganda . (4,000) (0.24 N.) (32.05 E.) (45)
Mnazi Moja, Zanzibar. See Zanzibar Town.
Mocuba, Mozambique (500) (16.49 S.) (37.01 E.) 40
Mombasa, Kenya < 500 4.03 S. 39.40 E. 50
Mongalla Prov., Sudan (1,500) (5.00 N.) (32.00 E.) (35)
Mongbwalu, Belgian eee A ‘ 4,000 (1.54 N.) (30.04 E.) (55)
Mongiro, Uganda . i " ; (3,000) (0.49 N.) (30.10 E.) (55)
ENTOM. II. 5 Kk
266 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE
List OF LOCALITIES—(cont.)
Locality Altitude Latitude Longitude Rainfall
Monrovia, Liberia . < 500 6.18 N. 10.45 W. > 100
Morogoro, Tanganyika 1,700 6.50 S. 37-47E. 35
Moshi, Tanganyika . 2,700 3:20 S. 37.32 E. 40
Moyale, Abyssinia . ; 3,900 3.30 N. 39.02 E. 25
Moyamba, Sierra Leone . < 500 8.09 N. 12.22 W. 100
Mozambique, Mozambique < 500 15.00 S. 40.42 E. 40
Mpumu Forest, Uganda . (4,000) 0.12 N. 32.51 E. (55)
Msambweni, Kenya < 500 4.299. 39.27 E. 60
Msoneddi, S. Rhodesia (4,800) (17.08 S.) (30.53 E.) (35)
Mtibwa Forest, Tanganyika (2,000) (6.00 S.) (37.45 E.) (35)
Mubende, Uganda . 5,100 0.35 N. 31.22 E. 45
Mug, Sudan ‘ (2,000) (12.42 N.) (29.09 E.) (15)
Mugusi River, Kenya (4,500) (0.30 S.) (34.42 E.) (50)
Muhangi Forest, Uganda. (4,000) 0.50 N. 30.45 E. (50)
Muhoroni, Kenya 4,300 0.09 S. 35.12 E. 60
Mukongoro, Uganda 3,300 1.20 N. 33-53 E. 55
Mukono, Uganda 3,900 0.21 N. 32.45 E. 55
Mumias, Kenya (4,500) 0.21 N. 34.28 E. 70
Mushin, Nigeria. See Yaba.
Mwanza, Tanganyika 3,700 2.32:9. 32.53 E. 40
Nacon, Gold Coast . (1,000) (10.50 N.) (1.25 W.) (40)
Nahud, Sudan 2,100 12.42 N. 28.26 E. 15
Nairobi, Kenya 5,500 ¥17:9. 36.50 E. 35
Naivasha, Kenya : 6,200 0.43 S. 36.21 E. 30
Namaacha, Mozambique . (800) 25.58 S. 32.08 E. 40
Namanve, Uganda . 3,700 0.21 N. 32.41 E. 50
Nampula, Mozambique (2,000) 15.06 S. 39.17 E. 45
Nandaw, Gold Coast (1,000) (10.44 N.) (2.18 W.) (40)
Nandawli, Gold Coast (1,000) (10.19 N.) (2.35 W.) (45)
Nangudi, Gold Coast (1,000) (10.51 N.) (0.40 W.) (40)
Nanyuki, Kenya 6,400 0.01 N. 37.05 E. 30
Narok, Kenya : (6,500) 1.05 S. 35.51 E. 25
Nata, Bechuanaland (4,000) 25.15 S. 25.15 E. (15)
Navarro, Gold Coast (1,000) (10.41 N.) (1.50 W.) (40)
Ndanga, S. Rhodesia (4,000) 20.11 S. 31.20 E. (40)
Ndola, N. Rhodesia 4,100 12.59 S. 28.38 E. 50
Nefasit, Eritrea (5,000) (15.17 N.) (39.03 E.) (30)
Nelspruit, Transvaal 2,300 25.27 5S. 30.58 E. 40
Nhemancone, Mozambique (500) (17.00 S.) (37.00 E.) (45)
Niangara, Belgian Congo (2,500) 3.41 N. 27.55 E. (65)
Njala, Sierra Leone < 500 (8.05 N.) (12.10 W.) (110)
Nobat Dakim, Aden Prot. I,100 (13.15 N.) (44.40 E.) (5)
Nova Lisboa, Angola : 5,700 12.45 S. 15.48 E. 60
Nova Choupanga, Mozambique (500) (17.09 S.) (34.40 E.) (30)
Nsawam, Gold Coast < 500 5.48 N. 0.20 W. (45)
Nta, Belgian Congo (2,500) 4.10 S. 15.45 E. (55)
Ntabanana, Zululand (1,000) (28.35 S.) (31.40 E.) (30)
Ntotoro Valley, Uganda . (2,000—4,000) (0.50 N.) (30.00 E.) (55)
Nuba Mts., Sudan . . ‘ (2,000—4,000) 11.00 N. 30.30 E. (35)
Nyamugasani Valley, Uganda . (3,000—12,000) (0.01 S.) (29.54 E.) (50)
Obenge-benge, Belgian Congo . (1,500) r.33'S. 25.05 E. (75)
Oblogo, Gold Coast 3 < 500 (5.36 N.) (0.17 W.) (40)
Obuasi, Gold Coast (700) 6.12 N. 1.40 W. (60)
Odzi, S. Rhodesia . 3,100 18.55 S. 32.25 E. (35)
Ogbomosho, Nigeria d (1,100) 8.06 N. 4.12 E. 40
Okahandja, S.W. Africa . 4,400 21.59 S. 16.56 E. (15)
IN THE ETHIOPIAN REGION 267
List oF LocaALiTIEs—(cont.)
Locality Altitude Latitude Longitude Rainfall
Okimbahe, S.W. Africa . (2,500) (21,22 S.) (15.23 E.) (10)
Omatjette, S.W. Africa . (3,500) 21.03 S. 15.32 E. (10)
Onderstepoort, Transvaal 4,200 (24.47 S.) (31.15 E.) 30
Opobo, Nigeria < 500 4.34 N. 7,25 Ee. 100
Oshogbo, Nigeria. 1,000 7.47 N. 4.29 E. 45
Otjimbingwe, S.W. Africa (2,500) (22:21 8.) (16.08 E.) (10)
Outjo, S.W. Africa . 4,000 (20.07 S.) (16.09 E.) 15
Payida, Uganda 5,000 2.25 N. 30.59 E. 55
Pemba I. : < 500 5.00 S. 39.50 E. 80
Pietersburg, Transvaal 4,200 23-54 S. 29.25 E. 20
Pinna, Gold Coast . (1,000) 10.53 N. 1.45 W. (40)
Pongwema, Belgian Congo (2,000) 4.15 S. 16.00 E. (55)
Ponthierville, Belgian Congo I,600 0.22 S. 25.28 E. (75)
Popokabaka, Belgian Congo (1,500) 5.42 S. 16.40 E. (55)
Port Harcourt, Nigeria < 500 4.40 N. be oh OF role)
Port Herald, Nyasaland < 500 17.029. 35.15 E. 35
Port Loko, Sierre Leone < 500 8.460 N. 12.44 W. 105
Port St. Johns, Cape Province < 500 31.38'S. 29.33 E. 50
Pretoria, Transvaal : 4,400 25.45 S. 29:12i1:, 30
Pujehun, Sierra Leone < 500 7.35 N. 11.06 W. > 100
Quelimane, Mozambique . 500 (17.51 5.) (36.54 E.) 55
Raga, Sudan . 1,800 8.28 N. 25.41 E. 45
Mt. Ramlu, Eritrea (1,000-6,000) | (13.15 N.) (41.45 E.) (20)
Ras Malalu, Eritrea ; < 500 (15.12 N.) (39.51 E.) (10)
Ressano Garcia, Mozambique . < 500 25.26 S. 32.00 E. (25)
Roberts Field, Liberia < 500 (6.50 N.) (11.22 W.) 100
Roberts Heights, Transvaal (5,000) 25.47 S. 28.08 E. 25
Roseires, Sudan 1,800 11.51 N. 34.23 E. 30
Rumbak, Sudan I,900 6.48 N. 29.42 E. 40
Rumogi, Uganda (3,000) 3.34 N. 31.21-E. (50)
Salisbury, S. Rhodesia 4,800 17.50 S. 31.01 E. 35
Saltpond, Gold Coast < 500 5.12 N; 1.05 W. (30)
Sambolugu, Gold Coast (1,000) (10.48 N.) (0.53 W.) (40)
Sango Bay Forests, Uganda (4,000) (0.50 S.) (31.40 E.) (45)
San Salvador, Angola 1,900 6:20'S. 14.47 E. 50
Santa Isabel, Fernando Po < 500 3.46 N. 8.46 E. > 100
Sapele, Nigeria ; < 500 5.53 N. 5.46 E. (100)
Savelugu, Gold Coast (500) 9.38 N. 0.50 W. (40)
Segbwema, Sierra Leone . < 500 (7.59 N.) (10.58 W.) (100)
Sennar, Sudan 1,700 13.34 N. 33-34 E. 20
Serakolia, Sierra Leone (1,500) (9.37 N.) (11.08 W.) (80)
Shaki, Nigeria (1,500) (8.34 N.) (3.19 E.) (40)
Shaloat, Sudan (2,000) (12.42 N.) (29.09 E.) (15)
Shamva, S. Rhodesia 3,200 17.20 S. 31.37 E- (35)
Sheikh Gamal, Sudan (2,000) (12.42 N.) (29.09 E.) (15)
Sheikh Karim, Sudan. (2,000) (11.01 N.) (30.33 E.) (35)
Sheik Othaman, Aden Prot, < 500 (12.53 N.) (45.03 E.) (5)
Shimoni, Kenya < 500 4.38 S. 39.23 E. (55)
Sidamo Prov., Abyssinia 5,900 6.10 N. 38.45 E. (40)
Singa, Sudan : (1,400) 13.10 N. 33-55 E. 25
Sinoia, S. Rhodesia. 3,800 1722'S. 30.11 E. 30
Sipi, Uganda . 6,700 I.20N. 34.22 E. (70)
Skukuza, Transvaal (1,000) 24.59 S. 31.36 E. 20
Sondu River, Kenya (6,000) 0,31 S. 35.04 E. (60)
Sonkonia, Sierra Leone (1,500) (9.45 N.) (11.26 W.) ; (80)
Sopo, Sudan . (2,000) (26.08 E.) | (45)
(8.06 N.)
268 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
List oF LOCALITIES—(cont.)
Locality Altitude Latitude Longitude Rainfall
Soroti, Uganda 3,700 1.43 N. 33-37 E. 50
Stanleyville, Belgian Congo 1,400 0.26 N. . 25.14 E. 65
Sunyani, Gold Coast 1,200 7.22 N. 2.21 W. 60
Tagoi, Sudan : (3,500) (11.53 N.) (30.55 E.) (25)
Takoradi, Gold Coast < 500 4.53 N. 1.46 W. 40
Talodi, Sudan 2,000 10.37 N. 30.21 E. 30
Tamale, Gold Coast 600 9.28 N. 0.51 W. 45
Tanga, Tanganyika < 500 5.05 S. 39.08 E. 55
Tanina, Gold Coast. (1,000) 9.55 N. 2.28 W. (45)
Taveta, Kenya 3,500 3.24 S. 37.40 E. 30
Tembura, Sudan (2,000) (5.36 N.) (27.30 E.) (55)
Tessenei, Eritrea (2,000) (15.08 N.) (36.39 E.) (15)
Thika, Kenya : ‘ 4,900 1.03 S. 37.05 E. 35
‘Third Camp’, Kenya. Unidentified.
Thysville, Belgian Congo : 2,000 5.16 S. 14.54 E. (50)
Tirikoro, Sierra Leone (1,500) (9.05 N.) (10.50 W.) (90)
Tishi, Gold Coast (1,000) (11.02 N.) (0.26 W.) (35)
Titule, Belgian Congo (1,500) 3.15 N 25.35 E. (75)
Tiwi, Kenya . < 500 4.158 39.35 E. 65
Tomango, Transvaal "2,400 (25.27 S.) (30.58 E.) (30)
Torit, Sudan ; 2,500 (4.25 N.) (32.33 E.) 40
Triangle Estate, S. Rhodesia (2,500) (21.00 S.) (31.30 E.) (25)
Tshela, Belgian Congo 800 4.58S 12.56 E. 50
Tshumbiri, Belgian Congo (1,500) (2.51 S.) (16.15 E.) (60)
Tsumeb, S.W. Africa 4,200 19.148 17.42 E. 20
Tsunikitoko, Belgian Congo (2,000) 4.20S 16.20 E. (55)
Tumu, Gold Coast . : I,000 10.52 N 1.59 W. (40)
Turbo, Kenya 5,900 0.38 N 35.03 E. 50
Tzaneen, Transvaal. 2,500 23.50S 30.10 E. 30
Ubassa, Nigeria (1,500) . (10.35 N.) (7.20 E.) (45)
Ugaro, Eritrea : (2,000) (14.46 N.) (37.20 E.) (20)
Ukara I., Tanganyika 3,900 5.528. 33.03 E. 60
Ulu, Gold Coast (1,000) (10.42 N.) (2.47 W.) (45)
Ulub, Aden Prot. (3,000) (13.30 N.) (44.42 E.) (5)
Umbeluzi, Mozambique . (500) 26.02 S. 32.20 E. 25
Umm Ruaba, Sudan 1,800 12.53 N. 31.13 E. 15
Umm Sunt, Sudan . (1,500) (14.20 N.) (33.35 E.) (15)
Umtali, S. Rhodesia 3,700 18.58 S. 32.40 E. 30
Upington, Cape Province ; 2,600 28.27'S. 21.15 E. be)
Upper Sheikh, Br. Somaliland . 4,700 (9.56 N.) (45.12 E.) 20
Usakos, S.W. Africa : 2,900 22.01 S. 15.35 E. 5
Usumbura, Ruanda-Urundi 2,600 3.26.8. 29.15 E. 35
River Vakila, Belgian Congo 5,100 (8.30 S.) (27.15 E.) (50)
Victoria Falls, S. Rhodesia 3,000 17.5258. 25.51 E. 20
Wa, Gold Coast I,100 10.05 N. 2.27 W. (45)
Wad Banda, Sudan : (2,000) 13.06 N. 27.55 E. (15)
Wad el Magdub, Sudan . (1,300) (14.23 N.) (33.24 E.) (15)
Wad Ganatir, Sudan (2,000) (12.35 N.) (28.37 E.) (15)
Wadiyain, Arabia . (1,000) (15.30 N.) (47.00 E.) (5)
Wad Medani, Sudan P 1,600 14.24 N. 33.31 E. 15
Waliki, Sierra Leone. Unidentified.
Waterpoort, Transvaal 2,600 (22.53 S.) (29.37 E.) (15)
Waterval Boven, Transvaal 4,800 25.38 S. 30.20 E. 30
Wathen, Belgian Congo . (2,000) (5.08 S.) (14.42 E.) (50)
Wau, Sudan . I,700 7.42 N. 28.01 E. 45
Weenen, Natal 2,800 28.51 S. 30.05 E. 25
IN THE ETHIOPIAN REGION 269
List oF LOCALITIES—(cont.)
Locality Altitude Latitude Longitude Rainfall
Wesu, Kenya . (5,000) 4.22'S: 38.20 E. 60
Windhoek, S.W. Africa 5,500 22.34 S. 17.05 E. 15
Winduri, Gold Coast (1,000) (10.43 N.) (0.50 W.) (40)
Witu, Kenya < 500 2.22 S. 40.27 E. 45
Wuri, Fr. Cameroons. See Duala.
Yaba, Nigeria < 500 (6.24 N.) (3.20 E.) (70)
Yala, Kenya (4,500) 0.06 N. 34.32 E. 65
Yambio, Sudan ; 2,900 4.34 N. 28.24 E. 55
Yangambi, Belgian Congo (1,000) 0.50 N. 24.15 E. 79
Yaoundé, Fr. Cameroons 2,300 3.52 N. 11.38 BE. 65
Yape, Gold Coast (500) 9.08 N. 1.10 W. (40)
Yatolema, Belgian Congo (1,500) 0.18 N. 24.32 E. (80)
Yeji, Gold Coast ; < 500 8.13 N. 0.39 W. (50)
Yiraia, Sierra Leone (1,500) (9.27 N.) (11.20 W.) (80)
Yokeskei R., Transvaal . (5,000) (26.05 S.) (28.00 E.) (30)
Yola, Nigeria ‘ 900 9.13 N. 12.29 E. 40
Lake Young, N. Rhodesia (4,500) (11.14 S.) (31.44 E.) (40)
Zaka, S. Rhodesia . ; : , ; (3,000) (20.21 S.) (31.24 E.) (40)
Zanzibar Town, Zanzibar ; : : < 500 6.10 S. 39.14 E. 65
Zika, Uganda. See Entebbe.
Zomba, Nyasaland Z : : ‘ 3,100 15.21.S; 35.25 E. 55
Zouragu, Gold Coast F : F : (1,000) 10.47 N. 0.47 W. (40)
Zungeru, Nigeria . : : : : (600) 9.45 N. 6.05 E. 45
BIONOMICS IN RELATION TO DISTRIBUTION
The following notes are intended only to illustrate certain aspects of bionomics of
particular importance to the study of distribution. The subjects discussed include
breeding-places, seasonal distribution, and biting-habits (including vertical distribu-
tion). It will be seen that our knowledge of all these matters, with the possible
exception of breeding-places, is still very rudimentary. It could have been much less
so had collectors realized the importance of publishing full data wherever possible.
Observations which may by themselves appear insignificant often acquire consider-
able significance when related to our knowledge as a whole.
BREEDING-PLACES
The fact that the majority of Stegomyia breed for the most part in tree-holes or
plant axils has had an important bearing on the nature of the distribution records
available. Much collecting has been carried out in Africa in the course of malaria
surveys, but in work of this kind attention is commonly confined to such breeding-
places as ground pools and swamp or stream edges. None of the Ethiopian Anopheles
is known to breed in tree-holes. As against this, however, it is interesting to recall
that two of the Ethiopian Stegomyia, Aédes africanus and Aédes trinidad, were first
obtained from ground pools. Records of this kind are also available for some other
species, e.g. luteocephalus (Briscoe, 1950) and apicoargenteus (Evans, 1925), but,
except where they refer to rock-pools, they may be assumed to be due almost cer-
tainly to flushing of tree-holes by heavy rain. All the records quoted are from places
with very heavy rainfall. The converse process, contamination of receptacles by
270 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
ground-pool or stream breeding larvae, also occurs quite commonly. One calabash
found by the author in a Ju-Ju shrine in Southern Nigeria contained a thriving
population of Anopheles gambiae, Megarhinus brevipalpis, and Aédes aegypti. Such
cases serve to emphasize the point that, save for certain highly specialized forms,
mosquito larvae are capable of a higher degree of tolerance than might be expected
from the comparatively restricted range of breeding-places in which they are normally
found. The restriction results, in fact, not from the preferences of the larva but from
those of the ovipositing female, though no doubt the latter are adapted to providing
the larva with optimum conditions for development (see Russell & Rao, 1942, and
summary in Bates, 1949). It is probable that no Ethiopian Stegomyia breeds exclusively
in one type of habitat, and cases in which only a single type of breeding-place is
known are probably attributable to insufficient collecting. Nevertheless with the
exception of certain highly plastic species (e.g. aegypti, albopictus) it is generally
possible to distinguish a breeding-place of choice from others which are merely
occasional or accidental. The whole question of the selection of breeding-places is
a complex one and has never been systematically studied, at least in the present
group. It would appear that the adoption of an unusual larval habitat may result
from diametrically opposite conditions, namely the absence of suitable breeding-
places or the existence of such ideal conditions that a teeming population is produced
which ‘overflows’ into every usable niche. |
The list of breeding-places which follows makes no pretence to completeness, but
an effort has been made to include the preferred breeding-place of each species, where
this is known, together with such records of occasional breeding-places as seem
necessary to give an idea of the adaptability of the species in question. The ecology
of the various classes of breeding-place is discussed briefly under seasonal distribution.
Aédes apicoargenteus. Preferred. Tree-holes (Hopkins, 1936 ; Garnham ef al., 1946).
Occasional. Rock-holes, utensils, bamboo stumps (Hopkins, 1936; Harris, 1942).
Aédes frasert. Preferred. Tree-holes (Hopkins, 1936 ; Garnham e¢ al., 1946). Occasional.
Rock-holes (Garnham ¢é al., 1946), rot-hole in mangrove (Holstein in litt.). Aédes
dendrophilus. Preferred. Tree-holes (Hopkins, 1936; Van Someren, 1946). Occasional.
Banana axils, banana and bamboo stumps (Hopkins, 1936), rock-holes (Garnham
et al., 1946). Aédes africanus. Preferred. Tree-holes (Hopkins, 1936, also mentions
it as common in bamboo stumps). Occasional. Utensils (Hopkins, 1936), rock-holes
(Garnham é al., 1946), bamboo pots (Robinson, 1950). Aédes pseudoafricanus.
Preferred. Rot-holes in Avicennia mangrove (Chwatt, 1949). Occasional. No other
breeding-places have been recorded. Aédes simpsoni. Preferred. Plant axils, especially
Colocasia, Gonja banana, and pineapple (Haddow, 1948). May be locally abundant in
tree-holes (Hopkins, 1936) or in utensils (Lewis, 1943a). Occasional. Coco-nut shells,
tubs (Hopkins, 1936), rock-pools (Lewis, 1943a), a concrete basin (Abbott, 1948),
bamboo pots (Robinson, 1950), axils of Dracaena and Strelitzia (Muspratt, 1950 and
in litt.). Aédes luteocephalus. Preferred. Tree-holes (Hopkins, 1936; Lewis, 19432).
Occasional. Cut bamboos, utensils, crab-holes (? accidental) (Hopkins, 1936), rock-
holes (Hopkins, 1936; Abbott, 1948), temporary pools (almost certainly accidental)
(Briscoe, 1950). Aédes unilineatus. Preferred. Tree-holes (Hopkins, 1936; Lewis,
1943a; Barraud, 1934). Occasional. Screw-palm axils (Ingram & De Meillon, 1929),
IN THE ETHIOPIAN REGION 271
rot-hole in paw-paw (Hopkins, 1936), bamboo pots (Robinson, 1950), bamboo stumps
(author, unpublished). Aédes metallicus. Preferred. Tree-holes (Hopkins, 1936).
Occasional. Utensils, coco-nut shells (Hopkins, 1936), rock-pools (Lewis, 1943),
banana axils (Teesdale, 1941), a concrete basin (Abbott, 1948), bamboo pots (Robin-
son, 1950). Aédes vittatus. Preferred. Rock-pools (including holes in coral, Wiseman
et al., 1939) (Hopkins, 1936). Occasional. Utensils, hoof-prints, boats, wells (Hopkins,
1936), tree-holes (Kerr, 1933; Rageau in litt.), bamboo pots (Harris, 1942). Aédes
aegypti. Common in all kinds of utensils, gutters, boats, water-tanks, &c. May be
locally abundant in wells (Hopkins, 1936). Dalziel (1920) records it, surprisingly, as
abundant in crab-holes.and this has been confirmed by Dunn (1927, &c.) and by
Riqueau (1929). Apart from this the dictum enunciated by Carter (1924), that the
species is never found breeding right through from oviposition to adult in pools all
the sides of which, at the waterline, are of mud, appears to hold good. Berner (1947)
gives records from residual pools in earth drains and the author has had similar
experiences, but it is extremely difficult in these cases to rule out contamination.
Some interesting wild populations have been described, notably by Haddow (19452,
1948) and Garnham é al. (1946). Haddow found larvae most commonly in tree-holes,
less commonly in plant axils. Garnham and his fellow workers found them most
commonly in rock-holes and in holes in recently felled trees, less commonly in true
tree-holes (for a discussion of this point see below under seasonal distribution).
Robinson (1950) found a high proportion of the tree-hole population to consist of this
species. He also gives records from rock-pools which seem to constitute a dry season
refuge. Adults of Aédes aegypti recently received from Dr. Fain were bred out from
holes in old weathered lava on the lava plain north of Lake Kivu near Sake. These
holes were said to be about 10 km. from the nearest human settlement.
SEASONAL DISTRIBUTION
A number of scattered records are available, but their sum total is surprisingly
small and little can be gained from attempting to correlate them. Most of our
knowledge therefore rests on a few studies pursued continuously over a period in
particular localities. There can be no question but that this is a most profitable field
for study, and good figures, were they available, would throw much light both on
bionomics and on distribution. As it is, so little is known regarding differences
between individual species that the subject is best discussed in relation to breeding-
places. For this purpose the latter are classified in accordance with common practice
as tree-holes, bamboos, plant axils, rock-holes, and artificial containers. It must,
however, be emphasized that this is not a natural classification and that even in the
present state of our knowledge further subdivision is required.
Tree-holes. Two distinct types were recognized by Garnham é¢ al. (1946), who found
that the large horizontal tree-hole not uncommonly encountered in fallen tree-trunks
was unattractive to most Stegomyia although apparently preferred to the smaller
type of tree-hole by Aédes aegypti. This is in accordance with the author’s experience
in Southern Nigeria except that such holes were not found to contain even Aédes
aegypti. The ‘dominant species’ appeared to be Culex macfiet. These observations
appear to indicate that the attraction for the ovipositing female is not purely a
272 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
chemical one depending on the nature of the walls of the breeding-place but is to some
extent conditioned by the degree of exposure of the water surface to insolation. If
this is so, then it might be expected that tree-holes with a more or less vertical
opening and deeply shaded interior would be more attractive to certain species than
those in the forks of trees with more or less horizontal opening. It is clear also that
the situation of the breeding-place relative to shade trees will be of importance.
Dunn (1928) made a quantitative study of the fauna of a tree-hole with a small
opening vis-a-vis one with a large opening. His observations appear to indicate a
general preference on the part of Stegomyza for the latter, but being based on only
a single tree-hole of each type they must be regarded as inconclusive. A number of
investigators (Dunn, 1927a; Harris, 1942; Garnham é al., 1946; Hocking, 1947;
Bailey, 1947) have employed ‘artificial tree-holes’, made by sawing off joints of
bamboo, as traps, and Dunn showed these to be more attractive than containers made
of metal. It would be interesting to compare the attractiveness of a container with
deeply shaded interior, such as a narrow-necked calabash, with that of a bamboo pot,
but this does not appear to have been done.
Larvae of Aédes unilineatus, metallicus, aegypti, simpsoni, and fraseri have all been
obtained by immersing dried material from tree-holes, and the larva of Aédes
subargenteus was first obtained in this way. It is presumed that these species survive
through the dry season by means of drought-resistant eggs (Taylor, 1934; Hopkins,
1936; Garnham eé¢ al., 1946; Gillett e al., 1950), but it appears highly probable from
the distribution studies discussed below that there is some variation in this respect
between individual species. Robinson (1950) appears to regard survival in the adult
state as more probable in Northern Rhodesia, and Taylor (1934) has an interesting
negative record for luteocephalus. The possibility of obtaining larvae by immersion
of tree-hole scrapings during dry-season surveys should be borne in mind by the
collector particularly as such scrapings are much more easily transported than
living adults or early stages.
Hatching does not usually take place immediately the first rain falls. Nor is it
to be expected that all the eggs in a tree-hole will hatch on the first immersion.
Ingram & Macfie (1916d) found larvae at Accra in March, ie. right at the beginning
of the rainy season, but Taylor, working at Gadau, noted that ‘although a few
showers fell during May and June no rainwater remained more than a few hours in
any of the tree-holes until the beginning of July’, and Muspratt (1945) notes that at
Livingstone, where the rainy season lasts from October to May, larvae are usually
found from the middle of November onwards. Robinson (1950) states that at
Livingstone he found larvae in tree-holes from December to May inclusive and that
‘one to one and a half inches of rain falling over a period of one to two weeks is
sufficient to initiate breeding in tree-holes, although most holes do not get sufficient
water before about three inches have fallen’. It is to be presumed that the last eggs
_ laid in a large tree-hole during the previous year would be near the bottom and would
therefore be flooded well before the tree-hole was full, but this point does not appear
to have been investigated. In view of Robinson’s findings in Northern Rhodesia it
seems almost certain that a number of species of Stegomyia can survive the dry
season as adults in wetter parts of the region. There can be no doubt, however, that
IN THE ETHIOPIAN REGION 273
the adult population is very drastically reduced even under quite favourable condi-
tions. Thus in Bwamba County during a prolonged dry spell lasting from the end of
December to the end of March 20 catches made between 2 February and 11 March
yielded only 7 adults of apicoargenteus as compared with 198 from 20 catches made
during the preceding wet season (Haddow et al., 1947). At Itowolo 7 catches made
_ between 21 November and 16 May yielded 24 africanus as compared with 501 from
15 catches between 12 June and 25 October. In this case the number of adults taken
declined very sharply even during the short dry spell between the large and small
rains and 2 catches made between 24 August and 7 September yielded only 7 africanus
(Mattingly, 19494). Adults do not, of course, appear in greatest numbers until some
time after hatching takes place. At Itowolo greatest numbers of africanus were
taken shortly before the period of maximum rainfall in July and a second peak
occurred shortly after the heaviest rainfall during the small rains. In this case the
seasonal distribution of africanus was closely paralleled by that of the two species of
Diceromyia which were the only other tree-hole breeders taken, and it was considered
that the type of breeding-place must be the governing factor rather than any intrinsic
difference between the species. As against this the figures given by Kerr (1933) for
africanus and luteocephalus in the Yaba area, although small, do suggest the possiblity
of a specific difference. In this case luteocephalus showed a much smaller peak in
relation to the small rains and a correspondingly larger concentration during the
period May to July. In addition the average catch obtained during July was more
than twice that obtained during June (slightly smaller during July than during June
for africanus at both Yaba and Itowolo). In Bwamba the wet and dry season catches
referred to in connexion with apicoargenteus yielded 429 and 34 africanus respectively.
Bamboos. Among the Ethiopian Stegomyia the only specialized bamboo breeders
are confined to the East African Highlands and cannot therefore be discussed here.
Most of the Guinean species appear to breed readily in bamboo pots and several of
them are known from bamboo stumps, so that it is clear that this type of habitat
closely resembles the small tree-hole. Bored bamboos clearly present a distinct type
of habitat, but no information appears to be available regarding their utilization by
Guinean species. In Southern Nigeria the most characteristic bamboo breeders
appear to be Dunnius spp.
Plant axils. These are the subject of a most informative paper by Haddow (1948)
devoted to the breeding-places and seasonal distribution of Aédes simpsoni in Bwamba
County. This author found that at Bundibugiyo with a rainfall of about 55 inches,
local rainfall had little direct effect on the numbers of larvae obtained since a light
shower was sufficient to fill the plant axils to capacity and heavy rain could not fill
them further. Three or four showers a month appeared to be sufficient under normal
circumstances to maintain the larval population. Increased rainfall favoured breed-
ing indirectly, however, through its effect on the growth of the plants. A monthly
survey of the axils of Xanthosoma sagittifolium showed that these only dried out
completely for a short period at the beginning of February and the larval population
was renewed within ro days after the first light rain. During the dry season larvae
appeared to be able to survive in the merest film of water. During the short period
of drought they could be obtained by immersing scrapings from the dry axils,
ENTOM. II. 5 Ll
274 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
presumably from drought-resistant eggs. It seems that in localities with a higher or
better distributed rainfall or in a wetter year larvae would occur all the year round
and the necessity for a drought-resistant egg would not exist. At Mombasa, with a
mean annual rainfall of only 45 inches, Wiseman e¢ al. (1939) found that some banana
axils held water throughout the dry season. They found adult simpsoni to be most
numerous during the drier months and suggested that heavy rainfall might wash the
larvae out of pineapple crowns and leaf axils.
Rock-holes. Garnham et al. (1946) showed that these are of two different kinds,
each supporting an entirely different fauna. Well-shaded holes in boulders in the
Kaimosi Forest were found to harbour typical tree-hole breeding species such as
frasert, africanus, and dendrophilus, but exposed pools outside the forest contained
only aegypti and vittatus. Residual pools in stream beds must also be distinguished
from those which are directly dependent for their water-supply on rain. Robinson
(1950) notes that at Livingstone the occurrence of such pools in combination with
tree-holes may serve to provide Aédes aegypti and vittatus with natural breeding-
places for nine months of the year, whereas tree-holes alone were available only for
six months. In general it would seem that the latter species is likely to be most
abundant during the dry season when the water level in stream beds is low (see also
Gil Collado, 1935). A rain-filled rock-hole exposed to sunlight is in general more liable
to rapid desiccation than a tree-hole, and species utilizing such breeding-places must
be capable of very rapid larval development. Such a capacity is certainly possessed by
vittatus, pupae of which have been found in rock holes only 3 days after the onset of
rain (Lamborn, 1930). Aédes aegypti is also capable of very rapid development, at
least when adequate food is available. On a particularly rich diet this species was
found to be capable of completing its development from the moment of hatching to
the moment of emergence in 85 hours at normal laboratory temperature (author’s
unpublished experiments at Yaba).
Artificial containers. These do not form a natural group. Their attractiveness for
particular species depends in part on their intrinsic nature and in part on their
situation. Shade and proximity to dwellings are important extrinsic factors. Most
species, other than aegypti, very seldom enter dwellings although some, e.g. africanus,
simpsont, luteocephalus, are occasionally recorded. Even aegypti has been shown to
prefer natural breeding-places when these are available in sufficient numbers, e.g. in
the case of villages situated near forests or shambas (Haddow, 1945a; Garnham et al.,
1946; Dunn, 1927a@ and c). Receptacles made of natural organic matter, e.g. the
bamboo pots used as traps or calabashes, to some extent mimic tree-holes and form
a class by themselves. The only species regularly found in small containers made of
metal, pottery, or other inorganic substances or in containers in or immediately
about houses is Aédes aegypti, and it is evident that its ability to utilize such breeding-
places accounts in large measure for its very wide distribution. Nevertheless it
cannot account entirely for this phenomenon. It may explain transportation,
especially by sea, and occurrence in urban centres with piped or other permanent
water-supplies, but it cannot explain occurrence in rural areas where the presence of
water in small containers is as much a seasonal phenomenon as its presence in tree-
holes. There can be no question that this is a highly adaptable species both with
IN THE ETHIOPIAN REGION 275
respect to temperature and to rainfall requirements, probably the most adaptable of
all the Ethiopian Stegomyia with the possible exception of vittatus.
In a special class are those very large containers used for conservation, e.g.
tanks, cisterns, concrete wells, hollow baobab trees, and these may perhaps be
essential to the survival of certain species occurring in very arid parts of the region.
Thus Patton (1905) records A édes vittatus only from such places in the neighbourhood
of Aden. Even here, however, fig-trees and residual and other rock pools might be
expected to provide natural breeding-places at least in some years.
BITING-HABITS
Under this heading are included three distinct aspects of mosquito behaviour,
namely host preference, the biting-cycle, which is an expression of the concentrations
of any particular species found biting at various times of day and night, and vertical
distribution, which is an expression of the concentrations found at various heights
above ground. All these, no doubt, are related reciprocally to distribution, but in the
present state of our knowledge the nature and extent of this relationship can only be
dimly apprehended. They are accordingly discussed very briefly.
Very little is known concerning host selection in this group. It seems reasonable to
suppose that forest species may be, in general, more specific in their preferences than
those inhabiting the savannah, and certain observations communicated privately by
Dr. Chwatt suggest that africanus may be more exclusively addicted to primate
hosts than luteocephalus. A detailed investigation of this point would be of consider-
able interest. On the whole the vast and confused literature relating to zoophily and
anthropophily in mosquitoes suggests that the majority of species are plastic in the
sense that host specificity occurs, if at all, at the level of the strain or biotype rather
than that of the species (see, in particular, Galliard, 1936, and Wanson & Nicolay,
1937). In this connexion it may be noted that ‘Culex molestus’, usually regarded as
a highly anthropophilic form, besides showing a preference for birds over man when
given the choice will, like Culex pipiens and Culex fatigans, produce many more eggs
on a meal of bird blood than on one of mammal blood (Mattingly e¢ al., in press, and
author’s unpublished observations on Lagos fatigans). Woke (1937) studied the same
question using A édes aegypti and found that of the various hosts employed man and
monkey were the least satisfactory. The figures he obtained, expressed as eggs per
mg. of ingested blood, were Monkey 28-2, Man 29-2, Canary 42-4, Turtle 46:3,
Rabbit 48-9, Guinea-pig 52-4, and Frog 52-6. As against this Toumanoff (1949)
obtained more numerous eggs and greater longevity with human blood than with
that of two species of lizards (Gecko and Calotes spp.). There seems to be no reason
to suppose that most mosquito species are any less plastic in this respect than in their
host preferences. If this is so, then the species population may be expected to adapt
itself to whatever selection of hosts is available in a given locality and we ought
not to be surprised if a species which readily bites man in one part of its range refuses
to do so in another. A number of inconsistencies of this kind are in fact to be found
in the literature relating to the Ethiopian Stegomyia. Thus De Meillon (1947a) states
that in South Africa Aédes vittatus does not bite man, but Robinson notes (in litt.)
that it bites in Northern Rhodesia. Garnham é al. (1946) noted that ‘Adults have
276 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
rarely been found, biting, in catches in the vicinity of breeding places’ in the Kaimosi
Forest, while, on the other hand, Leeson has an interesting unpublished record from
Malakisi, not far from Kaimosi, where vittatus bit in abundance in a wooded ravine
but were entirely absent from native huts at a similar distance from the breeding-
places. Jannone e¢ al. (1946) record it as biting in Eritrea, and Lewis (1943) considered
that it was probably the most important vector of yellow fever in the Nuba Mountains
because ‘it existed in great numbers and bit people by day and in the evening near
their homes’. Gil Collado (1935) records the Spanish form as biting man avidly.
Similarly Haddow (19454) and Garnham ¢é al. (1946) note that adults of forest
populations of aegypti are very rarely taken biting man. Aédes simpsoni has proved
equally elusive in Nigeria and the Gold Coast. Full length 24-hour catches at various
heights above the ground failed to yield a single adult although larvae were taken
abundantly in nearby banana groves (Bugher e¢ al., in press). Short-period catches
at ground level in banana groves at Asuboi also failed to yield any adults although
larvae were abundant (author’s unpublished observation). On the other hand, it is
understood from Dr. F. N. Macnamara that this species was quite often taken biting
in the British Cameroons, while Haddow (1945c) was able to take 596 adults in the
course of ten 24-hour catches in Bwamba County. Bostock & Simpson (1905) state
that this is one of the commonest species in the Low Veld region of the Transvaal,
suggesting by implication that here also it bites readily. Aédes unilineatus appears
very rarely to bite man. The only records of its biting or attempting to bite are two
from the Sudan (Lewis, 1947). This author failed to observe it biting in the Nuba
Mountains although larvae were common. A édes metallicus is recorded by Lewis (1943)
as biting only very near its breeding-places in the Sudan. Bailey (1947) rarely took
adults in Kenya; Wiseman e¢ al. (1939) found it to bite freely at Mombasa. Robinson
found it to the extent of 2 per cent. among just over 2,000 mosquitoes taken biting man
at Livingstone. Aédes apicoargenteus was taken by Haddow e¢ al. (1947) in Bwambaand
Garnham ef al. (1946) found it to be ‘the dominant species in many adult catches’ in
the Kaimosi Forest. In Nigeria only 7 specimens were taken during a year’s catching
(Mattingly, 1949b). Garnham é¢ al. (1946) took adults of fraserz in abundance in deep
forest, but Haddow e¢ al. (1947) record only one specimen from a considerable number
of catches in Bwamba. Haddow, Van Someren, e¢ al., however, record it as biting not
uncommonly in this part of its range (in press). Garnham é¢ al. (1946) took no adults
of dendrophilus in Kaimosi, but Haddow, Van Someren, é al., record it as ‘quite
common’ in Bwamba. Aédes africanus seems to bite man readily in all parts of its
range, and the same is true of pseudoafricanus and luteocephalus.
Regarding the biting-cycle it will suffice for present purposes to point out that
some species are markedly crepuscular with a strongly marked biting-peak shortly
after sunset. These include africanus (Haddow et al., 1947; Mattingly, 1949@; Kerr,
1933), pseudoafricanus (Mattingly, 1949); Chwatt, 1949), luteocephalus (Kerr, 1933;
Lewis, 1943a; Bugher e¢ al., in press), and metallicus (Lewis, 1947). Others feed quite
readily during the daytime, e.g. simpsont (Haddow, 1945c), vittatus (Lewis, 19434),
apicoargenteus (Garnham et al., 1946; Haddow et al., 1947), dendrophilus and frasert
(Haddow, Van Someren, e¢ al., in press), and possibly unilineatus (Lewis, 1947),
Concerning hosts other than man little is known, but Haddow & Dick (1948) give
IN THE ETHIOPIAN REGION 277
an interesting account of africanus and apicoargenteus feeding on anaesthetized
monkeys. De Meillon (1947a) took luteocephalus on a freshly killed baboon but failed
to take any in a Magoon trap baited with a monkey either at 50 ft. or at ground level.
Bedford (1928) gives records of simpsont and metallicus from horses and cattle.
With respect to vertical distribution evidence is available only for a few species.
Acrodendrophilic species, i.e. those biting in greatest numbers in the canopy, include
africanus (Haddow et al., 1947 ; Mattingly, 19494; Garnham eé¢ al., 1946), luteocephalus
(Bugher e¢ al., in press; Haddow, Van Someren, ef al., in press; De Meillon, 1947),
apicoargenteus (Garnham e¢ al., 1946; Haddow et al., 1947), and frasert (Garnham eé¢
al., 1946). It is very dangerous to base any estimate of the size of the population of
such species as africanus on ground-level catches. It must be emphasized that such
catches can give little or no idea of the relative abundance of a canopy species. Thus
in a series of catches in Bwamba Haddow et al. (1947) took only 22 out of 502 africanus
at ground level, while the author (1949a@) took only 39 at ground level out of a total
of 525 at Itowolo. It is true that Kerr (1933) obtained considerable numbers of
‘africanus’ on the ground, but this was almost certainly due to the presence of
pseudoafricanus as a contaminant. The presence of small numbers of the latter
species, which is less addicted to the canopy than africanus, may also have been
responsible for the rather higher percentage taken on the ground at Itowolo than in
Bwamba (7-4 per cent. as compared with 4:4 per cent.). It is also essential that
catches should be made during the peak biting-time. Species biting mainly on the
ground include dendvophilus (Haddow, Van Someren, ef al., in press), simpsoni
(Haddow, 1945c), aegypti, and metallicus (Bailey, 1947). Still less is known regarding
the vertical distribution of breeding-places. Bailey (1947) obtained simpsoni in
bamboo pots at ground, level only, and Garnham ef al. (1946) give a similar observa-
tion on forest-dwelling aegypti. Teesdale (1945), however, obtained the latter in
bamboo pots placed on the tops of coco-nut palms. Garnham ef al. (1946) obtained
fraseri and dendrophilus at all heights up to 60 ft. (the greatest height investigated),
but note that africanus, while breeding at heights up to 50 ft., was five times commoner
at ground level than higher up.
ZOOGEOGRAPHY
The existence of two major elements in the Ethiopian fauna, a West African on
the one hand and an East and South African on the other, has long been recognized.
Chapin (1932), basing his conclusions mainly on the birds, has mapped out an
approximate boundary for the West African sub-region and Edwards (1941) attempted
to fit this to the distribution of the mosquitoes. The latter author gives a list of
species supposed to be confined to the forested part of this area or to spread only into
the adjacent savannah. Among the Stegomyia he includes Aédes apicoargenteus,
frasert, luteocephalus, and dendrophilus. Of these, however, only apicoargenteus
occurs within the area of more or less continuous closed canopy forest where it is
accompanied by Aédes africanus, a species which Edwards does not include and
which is certainly found well outside the sub-region. Aédes luteocephalus is not a
forest species at all. It is a savannah species which does not enter closed forest. It
extends far beyond the boundaries of the sub-region. Of the four species listed by
278 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
Edwards only frasert and apicoargenteus are even approximately confined to the West
African sub-region and even these are found in certain isolated forest areas around the
Kavirondo Gulf which lie in Chapin’s East African Highland District. Aédes dendro-
philus resembles fraseri in possessing a restricted forest-fringe type of distribution
but, like some birds and Lepidoptera, this species also occurs in the forested parts of
Natal, an interesting distribution suggesting that it is a more ancient species than
the others, probably a relict from some past pluvial period when the southern forests
were more nearly continuous with those of central Africa (see Chapin, 1932: 376,
and Moreau, 1933). For general notes on mosquito distribution in the region see
Haddow (19454), De Meillon (19470), Lewis (1947), and Abbott (1948).
| 19° 9° ro 2.00 30°
oe >
, = 9
fl So , vt ° L10°
he 8
ve *s °
®
ef
es e e*. e \ NN
] e M7 7, as
0°: ’/] e *e e . re OM ~ Lo
© i e@e A tO, Ca
fu
F 2 7
°
a @ at gee Nt Ip
\
a H J
10- a abicoargenteus. 7
BS ssp. denderensis. ee 4 ¥
o Fraseri. Te = (a)
dB schwetzi. 4
Y West African sub-region. 0
Ta Sudanese Savanna District,
I b East African Highland District. >
Me Rhedeseen igh wat secede PEM
10° o° 10° 20° 30°
Fic. 4. Distribution of the Aédes apicoargenteus group.
Of all the West African species only apicoargenteus can be said to be typical of the
sub-region in the sense that it occupies the whole or a large part of it and extends for
only short distances beyond its boundaries (Fig. 4). This species is therefore discussed
first.
Aédes apicoargenteus. In so far as our knowledge goes, the distribution of this
species fits Chapin’s boundaries very well. The only certain records outside them are
from Kitgum and a few localities round the Kavirondo Gulf, all in the East African
Highland District, and Lorha in the Sudanese Savanna District. There is also a
IN THE ETHIOPIAN REGION 279
probable record from Kaduna in this district. All these localities have mean annual
rainfalls of at least 45 in., but if the 45-in. isohyet is taken as the limit of distribution,
further exceptions must be made, since Accra, Anecho, and Masindi Port certainly,
and Katompe and Kiansonzi probably, have a lower rainfall than this. The explana-
tion appears to be that, within limits, the monthly distribution of rain is as important
in determining distribution as its total amount. Accra and Anecho lie in the curious
dry belt extending approximately from Cotonou to Cape Three Points, where the coast
runs parallel to the path of the south-west monsoon instead of cutting sharply across
it as it does farther east and west. Although the rainfall in this area is low the
15° 10° 5° 9° 5° 10° :
4s"
aa > aaa
:
e cet
10% ; Le
°
. ; 18
. , : hs
e
e e x
so] ° abicoargenteus. — 4D : Ie
© Negative Records.
eo? Doubtful Record. a5"
PEM;
Fic. 5. Distribution of Aédes apicoargenteus in West Africa. The shaded area receives less than
1 in. of rain during each of six or more months in the year. Rainfall contours after Nash (1948).
proximity of the sea maintains a high atmospheric humidity and the seasonal dis-
tribution of rain is very equable, there being, on an average, only 4 dry months (i.e.
months with less than 1 in. of rain) at Anecho and 3 at Accra. Even so, however, it is
unlikely that apicoargenteus occurs permanently within the area since in some years
the rainfall is almost certainly too low (e.g. 10°84 in. at Accra in 1926). It is note-
worthy that the record from Accra (Macfie & Ingram, 19236) was for December 1918,
i.e. the end of a 3-year period of exceptionally heavy rain (41-05 in. in 1916, 44:20 in.
in 1917, and 32-37 in. in 1918). Deducing empirical limits from the data available
it may be said that apicoargenteus is not known to occur anywhere with less than
25 in. and 3 dry months, 30 in. and 4 dry months, or 45 in. and 5 dry months. On
this basis we might expect to find the species everywhere in the western part of its
range with 5 dry months or less, and such a distribution is in good accordance with
the data available. Reliable negative records are few and, in general, no use is made
of these in the present paper. Those shown in Fig. 5 (Gadau and Kano) seem to be the
only ones of much value in the present instance.
In the eastern and southern parts of its range the distributional limits of apico-
argenteus are probably mainly altitudinal. Over almost the whole of Uganda rainfall
280 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
is high and extremely equable in its annual distribution (Masindi Port and the
Kiansonzi area have only one dry month), but there is some reason to suppose that
the species may be absent from the extreme northern part of the territory. Thus
Lumsden in a recent survey failed to find it beyond Payida in the West Nile Province
although other localities visited (Godia, Laufori, Metu, Midia, Rumogi) would seem,
to judge from nearby rainfall stations, to have plenty of rain (3 dry months in each
case). It cannot, however, be said that reliable negative records have been established
a 26° 31°
4ow
eoove® Kee)
e
5 o"
fe @vreeeK?
Sai %
x
@ abicoaurgenteus.
=Fovest.
oto 6 Putative rainfall limit.
—~- Ysohyets.
x Rainfall Stations.
26°
Fic. 6. Distribution of Aédes apicoargenteus in north-east Uganda and the Sudan. Only those
rainfall stations are shown which are in the critical area.
for this area, since the survey was a very short one. When the empirical limits
deduced for the western part of the range are applied here an exceedingly interesting
picture is obtained (Fig. 6). The boundary turns east in the neighbourhood of Juba
and again farther north in the neighbourhood of Bor. How far east it runs is unknown
since no figures are available from this comparatively little-known part of the Sudan,
but it is tempting to suppose that it may join with a similar boundary delimiting
the wetter parts of Abyssinia. The similarity of the Abyssinian and West African
faunas has often been a matter for comment (see, e.g., Carpenter, 1935), and whether.
a bridge still exists in this region or not, it seems reasonable to suppose that a relatively
slight change of climate would suffice to recreate one. Aédes apicoargenteus is not
known from Abyssinia, but so little collecting has been done there that negative
records are quite valueless. Aédes africanus is known from the Sidamo Province so
that the existence of a fairly recent link between the Abyssinian and Guinean
Stegomyia may be safely inferred. The more northerly part of the Bor—Pibor region
IN THE ETHIOPIAN REGION 281
is largely swampy and is stated by Lewis (1947) to possess very few trees, but the map
(NB 36 of the International Series) shows small isolated patches of forest having the
appearance of relicts of a continuous belt running north-east from Mongalla to the
Abyssinian border. Collections from these patches of forest as well as from others in
the Bor, Mongalla, and Juba areas would be of great interest.
From Mt. Elgon southwards the distribution of apicoargenteus is limited well to the
5 . :
e abicoargenteus.
« Ossp.denderensis.
1) —] Landover 5000’
imal Land over 3000%
", i ad ; ‘ i
Za actaecitsiessthan Ho!
& ° nf
ier st on at he A
15° a BOP 35°
Fic. 7. Distribution of Aédes apicoargenteus in the southern and eastern parts of its range.
PEM,
west of the 45-in. isohyet by altitudinal factors (Fig. 7). Haddow, Van Someren, e¢ al.
(in press), report that it does not occur above about 5,500 ft. on the western slopes
of Ruwenzori, and Garnham é al. (1946) record it up to approximately the same
altitude in Kavirondo. This is a critical level for many plants and animals at this
latitude (see Chapin, 1932). Considerable areas of northern Tanganyika, while
having less than 45 in. of rain in the year, have a sufficiently well-distributed rainfall
to support the species, at least on the basis calculated for West Africa. No collecting
has, however, been done except at and near Mwanza, from which there is a negative
record.’ Once again surveys of isolated patches of forest would be of considerable
interest. In the Kivu Highlands apicoargenteus appears to extend up to at most
5,000 ft. Above this it is replaced by schwetzi (very rare as far north as this) or more
1 See Appendix.
ENTOM. II. 5 Mm
282 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
commonly by ssp. denderensis. Farther south still, as noted by Chapin for the birds,
the altitudinal limit is much lower and it is not known to occur above 2,500 ft. or
possibly 3,000 ft. (Kayembe-Mukulu). In the Katanga and the Rhodesian Highlands
it certainly does not reach 4,000 ft. (Elisabethville, Ndola), and it is replaced at these
altitudes and, farther south still, at 3,500 ft. (Balovale) by schwetzi. It seems im-
19° 9° 19° 3o° 35°
ma os 4 |
Te 10°
~ sa
roe Lo?
: he CALT%
CO} eAedes Fraseri. ; - 0°
(70 Closed Forest.
Area of insufficient rainfall. a
V7AZ Land over S000’, . ==
= vi 109 30° 35°
r 0° 10° 20° 30°
7
10 orl
oO
Oo 7) ~
© r)
0- i C ro?
orn Yy
192 0° 10° 20° 40°
Fic. 8. Distribution of Aédes fraseri in the western and eastern parts of its range and putative
range as a whole.
probable that this species would be found anywhere in the southern Congo above
about 3,000 ft.
Summarizing the distribution of apicoargenteus it may be said that it shows a
much less clear-cut relationship to rainfall and altitude than do a number of other
species. The explanation may be that it requires for permanent survival in an
area a more purely equatorial type of forest than the dry or mixed forests which
suffice for, for example, africanus. This matter is, however, beyond the scope of the
present paper and must be left for discussion in a later one.
Aédes fraseri. As already noted this species has a peripheral or fringing distribution
IN THE ETHIOPIAN REGION 283
relative to the main Guinean forests (Fig. 8). Haddow states (in litt.) that, unlike
apicoargenteus and africanus, it is never taken more than a short distance inside the
Semliki Crown Forest. Garnham éé al. (1946) found it in the depths of the Kaimosi
Forest, but it would appear that this is a comparatively poor type of relict forest.
Chapin (1932: 232) discusses the distribution of birds having a similar habit and
states that they follow the northern border of the Congo forests in a narrow band and
sometimes extend southwards along the eastern edge of the forests or occur in places
along its southern margin. Except at Dubreka fraser is not known to occur any-
where with a rainfall of less than 40 in. or more than 3 dry months. Dubreka has 4
dry months with a rainfall of over 150 in., and here the species is at present known
only from mangrove. Applying the limits of 40 in. with 3 dry months or Ioo in.
with 4 dry months a putative distribution is obtained which is in very good agreement
with the known distribution of fraseri and the type of distribution described by
Chapin (Fig. 8). An extensive area with comparable rainfall also exists in the southern
Congo, and it is possible that frasert may be found there although at present there
are no records. Similar altitudinal limits appear to apply to this species in Uganda
and Kenya to those given for apicoargenteus (Garnham ef al., 1946; Haddow, Van
Someren, ef al., in press). It is to be noted that the dry area in the Gold Coast and
Togoland and, farther east, the Adamawa Highlands both constitute serious inter-
ruptions to the distribution and it would seem that the Upper and Lower Guinean
populations may at times be discontinuous.
Aédes dendrophilus. In the northern part of its range this species appears to have
a fringing distribution very similar to that of fraseri (Fig. 9), but its putative rainfall
limits are even more restricted since it is not known from anywhere with more than
2 dry months in the year.’ In addition it is also known from an area in Natal with the
same type of rainfall and from Fernando Po which suggests that it is a rather ancient
species with, at one time, a much wider distribution. This view is also supported by
its morphological characters. Its altitudinal limits appear to be much the same as
those cited for fraseri (Garnham e al., and Haddow, Van Someren, é al., loc. cit.).
Various small, isolated areas in the southern part of the continent appear to possess
an adequate rainfall for the species and not all of them are at impossible altitudes.
Among the more promising are the Haenertsberg—Magoebaskloof area of the Trans-
vaal, the Mbabane region of Swaziland, parts of Mozambique (Vila Paiva d’Andrada
and Chinde—Pebane areas), the Inyanga, Morgenster, Bikita, Mt. Silinda, and
Umtali regions of Southern Rhodesia, and certain localities in southern Tanganyika
at the northern end of Lake Nyasa (Makete, Musekera, Mwitika, Kyela).
Aédes africanus. Like apicoargenteus but unlike fraseri and dendrophilus this
species is found in the depths of closed canopy forest which does not therefore con-
stitute a barrier to its distribution. Among mosquitoes, as noted by Chapin for the
birds, this capacity to penetrate thick closed canopy forest is the exception rather
than the rule. Unlike apicoargenteus, africanus has not been found in the dry part of
the Gold Coast. It is, however, able to surmount the Rhodesian Plateau and in
consequence its distribution is very closely and completely defined by the 4o-in.
isohyet except to the east of Lake Victoria where there is a complete altitudinal
t See Appendix.
284 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
barrier and south of Lake Nyasa where the rainfall is sufficient but its distribution is
unsuitable (Fig. 10). The only records from outside the distributional area so defined
are those from Zomba (Theobald, 1901) and Mozambique (Howard, 1912), of which
the former certainly and the latter very probably fall within an isolated area of
adequate rainfall which may well support a relict population (Fig. 11). It is
~ if 9 20° 30° ed
ad ? rid)
&
y— =
ey a)
AREY |
: E J.
0+ | % «¢ Lo
? y
Be
G.
e Aédes dendrobhilus.
; Areas with less than Ho"
UL be tapanpban than a dvy
mon ths.
“~~
$.
&
UH hand over 5000’.
=, Closed canopy Fovest. REA.
20° 10° o° 10° 20 30° Ao?
Fic. 9. Distribution of Aédes dendrophilus.
unfortunate that these records can no longer be checked. The putative distributional
limits of 40 in. with 4 dry months, 45 in. with 5 dry months, or 50 in. with 6 dry
months, deduced empirically from the known distribution, fit this distribution so well
that it can scarcely be doubted that they are significant. As to whether their effect
on the species is direct or indirect, through the effect on vegetation, is not at present
known.! The existence of a relict population of this species in Abyssinia has already
been referred to. Altitudinal limits appear to be similar to those of apicoargenteus,
but with a decidedly greater tolerance of high altitude especially in the southern part
of the range. Known limits include 5,500 ft. on Ruwenzori (Haddow & Van Someren,
? See Appendix.
IN THE ETHIOPIAN REGION 285
1950), about 6,000 ft. in the Kavirondo region (Garnham é¢ al., 1946), about 6,000
ft. in the Kivu Highlands (Costermansville) and in Ruanda (Dendezi), about 5,500
ft. on the Kibara Massif (R. Vakila) and on the Bihé Plateau (Bihé), and about
4,000 ft. on the Rhodesian Plateau (Elisabethville, Ndola). The record from the
Sidamo Province of Abyssinia (Bevan, 1937) is associated with an altitude of
5,900 ft.
UZLLIZTTZZ, 7 7 4
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10° 0° 10° 20° 30° ho" 4o*
Fic. 10. Distribution of Aédes africanus.
Aédes simpsont. This is the only species, in addition to those already discussed,
which occurs in the area of continuous closed canopy forest. Its presence here is
almost certainly the result of introduction by man, since Haddow (1945c, 1950, &c.)
has clearly shown that it is a mosquito of the forest edges and especially of plantations
and shambas, and its occurrence in the forest proper is confined to stray individuals
and is so rare as to be without significance. Haddow has pointed out, in conversation,
that very much traffic takes place in young pineapple and banana plants which
frequently contain larvae. As a plant-axil breeder it flourishes in areas of very much
lower rainfall than can be tolerated by the species so far discussed. The minimum
average rainfall tolerated is, as far as present records go, 20 in. with 8 dry months
(Sennar), and the 20-in. isohyet fits the known distribution very well (Fig. 12). These
rainfall requirements appear to exclude it entirely from Chapin’s Sudanese and
Somali Arid Districts and from his South-West Arid District (except in the Bulawayo
area). Otherwise it occurs in all the main faunal districts. Its altitudinal limits
appear to be much the same as those of africanus and it is known from up to about
286 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
5,500 ft. on Ruwenzori, 5,000 ft. in the Cameroons and in Eritrea, 6,000 ft. in the
Kivu Highlands, 5,500 ft. on the Bihé Plateau, 4,000 ft. on the Rhodesian Plateau,
and 4,500-5,000 ft. in the Transvaal (Roberts Heights) and Southern Rhodesia
(Salisbury, Bulawayo). It is rare on the Rhodesian Plateau, but Robinson (1950)
considers this to be due to the absence of banana plantations. Garnham ¢é al.
(1946) appear to have found it up to about 6,000 ft. in Kavirondo but, unfortunately,
26° 28° 30° 32° 3%e 36° 38°
83 UTangangihoa a5 SBN - =Lee
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020
a
104 Q sa
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Area withunsuitable rainfall. “5 “a
of Doubtful Records. 45 0 ee.
s NS <a
26° 219° 30° 32° 34° 36° 36° ei
Fic. 11. Distribution of Aédes africanus in the south-eastern part of its range showing rainfall
and number of dry months,
its distribution in other parts of the Kenya Highlands is not known with certainty
since no records from altitudes above about 4,500 ft. can now be confirmed. The
virtual isolation of the Transvaal and Natal population by a dry belt in Southern
Rhodesia and Mozambique has been noted above.
Aédes luteocephalus (Fig. 13). This species appears, if anything, to be even more
reluctant to enter closed forest than st¢mpsoni. Haddow has stated, in conversation,
that it will not enter the edges of the Semliki Forest even though it may be biting
freely in the open. As a corollary to this it appears to be entirely absent from the
Upper and Lower Guinean Forests even though it has succeeded in establishing
itself in coastal areas where the former has been cleared (e.g. Roberts Field). It has
also been recorded from two localities (Adun, Bende) in the western extension of the
IN THE ETHIOPIAN REGION 287
Lower Guinean Forest beyond the Cross River. These records cannot now be con-
firmed from specimens and may have referred to africanus. It is in any case probable
that the forest in this area has by now been so broken up as to constitute only a
partial barrier. It appears to be even more resistant to drought than simpsont, the
19° 0° 10°
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° ®
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Td Southwest Arid District. ,
[304 Tle Southeast Veld District. -30
Area With 20 of rain or less.
PEM.
20° 10° 0° 10° 20° 30° eo®
Fic. 12. Distribution of Aédes simpsoni.
lowest rainfall recorded anywhere in the northern part of its range being 15 in. with
8 dry months (Danagla and Wad el Magdub in the Wad Medaniarea). In the southern
part of its range it is also known from Maun, which has only 15 in. The record from
Nefasit shows that in this part of its range it can occur up to about 5,000 ft., but
elsewhere it is rare above about 3,500 ft. In western Kenya it has been recorded
from two localities at about 4,000 ft. (Kerio, Fort Hall), but on the Uganda Plateau
it is very rare and the only records which can be confirmed are from about 3,000 ft.
in the Semliki Valley and from between 3,000 ft. and 4,000 ft. in the West Nile
288 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
Province. The record from Mpumu cannot now be confirmed and may have referred
to africanus. As against this it seems possible that luteocephalus occurring in this
region may occasionally have been misidentified as africanus. On the Rhodesian
Plateau it has been recorded from Balovale (3,400 ft.) and Ndola (4,100 ft.). At Ndola
it varies in abundance from year to year and in one year only a single specimen was
_t9° i 10° 20° me 40°
ae Se
te
ow)
12 pel f=5 Lot
e luteocebhalus.
FS Land above 3500’,
Avea.of continuous closed
ad esi caneby forest. ;
TI} Aveawith less than 15" rain. )
10° 0° 10°
Fic. 13. Distribution of Aédes luteocephalus.
taken (Robinson in litt.). The eastern limit of its distribution cannot be inferred
since no collecting has been done in western Tanganyika. There is a single specimen
from Likoma Island off the eastern shore of Lake Nyasa in the London School of
Hygiene collection. Nor is it known whether it occurs in Mozambique, but it may be
taken as virtually certain that it does not extend nearly as far south as simpsont.
Aédes unilineatus. This species is of interest as being one of the few Ethiopian
Stegomyia which also occurs in the Oriental Region. Like luteocephalus it is purely
a savannah mosquito, but its resistance to drought appears to be even higher and it
is on record from several places with rainfalls of about 10 in. (Dolo, El Fasher,
Erkowit). Unlike luteocephalus it does not occur in deforested areas with high rainfall
such as Freetown and it is not, in fact, known from anywhere in Africa with more
IN THE ETHIOPIAN REGION 289
than 55 in. per annum (Tembura). It is not known whether it occurs in those parts of
Arabia with adequate rainfall and there are no acceptable negative records even
from the Aden region since it is not clear that anybody has searched in tree-holes.
It is stated by persons with local knowledge that the latter are by no means rare in
the wetter parts of Arabia, especially in fig-trees growing in the wadis. It isnot known
from anywhere above 4,000 ft. except Erkowit, which is in the hottest part of its
5° 0 15° 30° 45° 60° 15°
\ =
30° ? : PF
e Aédes unilineatus.
Area with less than 10"rain.
lag E53 Area with more than 60°rain.
TD Land over 3500".
BEM.
15° o° 15° 30° 45° 60° 15°
Fic. 14. Distribution of Aédes unilineatus. Inset: Putative distribution of unilineatus with that
of two related species.
range. Elsewhere the highest altitude recorded is at Fort Victoria (3,700 ft.), and the
3,500-ft. contour probably provides a good approximate boundary. It does not appear
to enter forest and this probably serves to exclude it even from those parts of the
Congo Basin with less than the empirical limit of 55 in. of rain per annum. The
putative distribution is shown inset in Fig. 14. As to whether the southern populations
of the Zambesi and Limpopo valleys are linked with the northern population of the
Sudanese and Guinean savannahs is not known, but, in view of the fact that it
occurs as far east as Dolo, it seems possible that it may extend southward along the
edges of the central plateau. The Indian records seem to indicate similar climatic
limits with the exception of that from the Bombay area. The rainfall here is about
70 in. per annum. It appears that unilineatus is a relict of a fauna which was at one
time widespread over much of Asia Minor. Related species still occurring in this
ENTOM. II. 5 Nn
290 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
area include granti, which is known only from Sokotra, and cretinus, which has been
recorded from Crete (Edwards, 1921b), Macedonia (as delta, Séguy, 1924), and
Transcaucasia (as albopictus, Rhoudkhadzé, 1926; as lindtropi, Schingarew, 1927,
and see Stackelberg, 1937).
Aédes metallicus. This species has been recorded from only four localities in the
a 19° 9° 108 20° 30° nae
e e
Y e +. e of
ee e > e e 9
10% ~ “ 10°
°
e
e
0 \
o ‘4 aS
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AL, Boundavy of West African rs 14 s:
Sub-region,
30; 30)
PEM.
20° 10° 0° 10° 20° 30° 40° 50°
Fic. 15. Distribution of Aédes metallicus.
West African sub-region. Two of these (Mbarara and Yambio) are peripheral. The
other two (Accra, Asuboi) are in the dry area of the Gold Coast and Togoland (Fig.
15). Its resistance to drought appears to be greater even than that of unilineatus and
it is the only species so far discussed which occurs widely in Chapin’s South-West
Arid District. It is not, however, known from Arabia. Despite its almost complete
absence from the West African sub-region it is known from several localities in
East Africa with quite high rainfall, e.g. Kakamega with 70 in. and Tanga with 60 in.
It seems probable that temperature plays a greater part in controlling the distribution
of this species than in the case of those previously discussed. Its altitudinal limits
IN THE ETHIOPIAN REGION 291
vary in different parts of the range. In Kenya it has been recorded from an indefinite
altitude in the Fort Ternan area (Garnham ¢é al., 1946), apparently about 6,000 ft.,
and from 5,500 ft. in the Nairobi area. There are no records from the Kivu High-
lands and only one from the Rhodesian Plateau (Jadotville, 4,300 ft.). On the Bihé
Plateau it apparently occurs at about 5,500 ft. and in South-West Africa it occurs
up to about 4,500 ft. (Okahandja). It appears to be absent from the higher parts of
the Transvaal where it has not been recorded above about 2,000 ft. (Letaba), but in
Southern Rhodesia it occurs up to 4,800 ft. (Salisbury). Morphologically it is unique
among the Ethiopian Stegomyia in lacking a ventral process from the paraprocts,
a feature which recalls the oriental members of the sub-genus. Its mesonotal
ornamentation is also unique and suggests a comparison with the Aédes longipalpis
group vis-d-vis the other Ethiopian Finlaya.
Aédes vittatus. This is the most widely distributed of the species here discussed
(with the exception of aegypti). It combines drought resistance, due at least in part
to its ability to breed in rock-pools, with a marked ability to withstand low tempera-
tures. It is, however, by no means equally common everywhere. In the Lagos area
it is extremely rare, having only been recorded once. In Bwamba only a single speci-
men has been taken (Haddow, Van Someren, eé al., in press). It appears to be absent
from or extremely rare in a large area of the Belgian Congo, a fact which may perhaps
be related to the nature of the surface rocks (Lubilash sandstones, see Chapin, 1932:
30). It is not clear to what extent it enters forest, but Garnham e# al. (1946) specifically
mention its absence from shaded forest rock-holes and there are many indirect
references in the literature to suggest that it is largely or entirely a mosquito of the
open country. That this does not preclude it, as it apparently precludes luteocephalus,
from the forested eastern part of the Congo is readily understood from the fact that
its breeding-places of preference are commonly found in the exposed beds or edges
of streams. The occurrence of this species in Spain is of particular interest since it
implies the ability to contend with two sets of adverse conditions, the long, cold
winters and the torrential nature of the breeding-places during spring and summer.
Gil Collado (1935) notes that it is most abundant in the autumn. It is not known in
what stage the winter is passed in this part of the range, although it seems almost
certain that it must be in the egg. On the basis of the Ethiopian records alone, this
would seem to be more resistant to low temperature than any of the other species
discussed, except perhaps aegypti, if only because it occurs in a number of places
in the Kenya Highlands well above 6,000 ft. (Eldoret, Kericho, Kiambu, Kitale,
Mara R., Maseno, Meru, Narok). There is also an interesting record from Sipi
(6,700 ft., Hancock & Soundy, 1929), on the Uganda side of Mt. Elgon. As against
this there are some rather striking negative records. Patton (1905) notes that it has
not been found on the Jebel Jihaf above Ulub. Mara (1945) failed to find it on Mt.
Bizen. (This may have been a coincidence, but it seems to be a very common
mosquito at lower levels.) It appears to be very rare on the Rhodesian Plateau, where
the only records are from Elisabethville and Livingstone, and there is no record from
the Kivu Highlands. The likeliest explanation seems to be that the egg is highly
resistant to low temperatures as well as to desiccation but the adults and larvae are
not. In this case survival would depend on the coincidence between suitable
292 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
temperature conditions and a low water level in the streams. Records from over
4,000 ft. in the southern part of the range include Bihé, Salisbury, Goromonzi, Gwelo,
Msonneddi, Onderstepoort, Yokeskei River, Waterval Boven, and Pietersburg.
It is interesting to note that it has not been found as high up as metallicus in South-
West Africa or as simpsoni in Eritrea.
e 35° Go 1
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ey
[awd
+ °
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“a ®
(~
° a 4
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ag Nec
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Say
5 °
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@ Aé.vittatus. : 4 °° /
@
ae. . se
PEM.
0° 30° 60° qo*
Fic. 16. Distribution of Aédes vittatus.
Aédes aegypti. For reasons already given it is not proposed to discuss the dis-
tribution of this species in detail. Some notes on its altitudinal limits may, however,
be of interest as bearing on the general problem of Stegomyia distribution and the
relationship between the Ethiopian and Palearctic elements in the sub-genus. The
most remarkable record of aegypti from the Ethiopian region is perhaps that of Mara
(1945), who found it at nearly 8,000 ft. on top of Mt. Bizen, and this during by no
means the warmest part of the year. That this was a temporary introduction is
highly probable, but it seems that larvae at any rate can survive for short periods
at temperatures well below those which broadly limit the distribution of the species.
Some unpublished records from comparatively high altitudes in Kenya, kindly sent
by Mrs. Van Someren, include the following: Eldoret (6,900 ft.), Fort Ternan (7,000
ft.), Kajiado (about 6,000 ft.), Kiambu (6,300 ft.), Londiani (7,500 ft.), Lumbwa
(6,300 ft.), Maseno (6,000 ft.), Naivasha (6,200 ft.), Nanyuki (6,400 ft.), Narok
(6,500 ft.). Symes (1935) notes that it is particularly abundant at Meru (6,200 ft.).
Southerly records from comparatively high altitudes include Salisbury (4,800 ft.,
ee
Te
IN THE ETHIOPIAN REGION 293
Theobald, 1910), Pretoria (4,400 ft., Theobald, 1910), and Windhoek (5,500 ft.,
De Meillon, 1943). MacGregor (1927) makes the curious observation that in Mauritius
aegypti seems to be restricted to the coastal belt and is very local while on Rodriguez
it is much commoner and is found up to ‘at least 800 feet’.
SUMMARY
The le paper deals with the distribution of those species of Stegomyra which
have been found in the West African sub-region. No species is entirely confined to
the sub-region as defined by Chapin. On the basis of this sub-genus it would be
reasonable to extend the limits of the sub-region to include all land below 6,000 ft.
around the Kavirondo Gulf and, farther north, to a point somewhere east of Kitgum.
All published records are listed together with such unpublished records as are at
present available. It has been possible to verify, and where necessary amend, the
majority of doubtful records by reference to preserved specimens. Records for which
this has not been possible are discussed individually. Full distributional records of
Aédes aegypti are not given since the data available for this species are so numerous as
to require separate treatment. Records of the species discussed in this paper are
available from rather more than 500 different localities and these are listed with
their altitude, longitude, latitude, and rainfall as nearly as these can be ascertained.
Some notes on taxonomy are given, including corrections to published descriptions
and some reassignment of specimens and new synonymy. The keys to the sub-genus
at present available are extremely misleading and new keys to adults and larvae have
therefore been prepared. Full taxonomic treatment is reserved for a later paper.
Bionomics are discussed in relation to distribution with particular emphasis on
seasonal distribution, the study of which is regarded as crucial for our understanding
of every aspect of the group. Zoogeography is discussed principally in relation to
rainfall. Temperature will be discussed in a later paper. It is here considered only
in so far as its influence is manifested in altitudinal distribution. By far the most
serious hindrance to our understanding of the distribution of the group is the absence
of records from almost the whole of Tanganyika and Nyasaland. Some points of
general interest which have emerged in the course of the work include the existence
of a climatic ‘bridge’ between the Ubangi—Uelle and Abyssinian Highland districts,
across the south-eastern extremity of the Sudan, the existence of which may help to
explain the strong Guinean element in the Abyssinian fauna. The existence of an
isolated population of Aédes dendrophilus in Natal and Pondoland supports the
hypothesis of a much wider extension of the African forests in the past already
inferred from the distribution of other groups. It is shown that large parts of the
Guinean region have the same type of annual rainfall distribution as the part of
South Africa in question. Old records, which unfortunately can no longer be checked
from specimens, indicate the presence of an isolated population of Aédes africanus
in southern Nyasaland and a part of Mozambique. It is shown that the area in
question possesses a type of rainfall suited to africanus but is separated from that
part of Northern Rhodesia in which this species is known to occur by a broad belt in
which the annual rainfall distribution is unsuitable. The Upper Guinean Forest,
although by no means continuous, appears to be sufficiently so to constitute a
294 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
complete barrier to all but five species, and two of these, aegypti and simpsoni, may
have been introduced into its interior by man. Aédes vittatus occurs only in the
higher northern and eastern parts of the forest area. Its exclusion from the remainder
may be the result of geological factors. It is probable that penetration of those areas
in which it does occur has been along stream and river beds since it does not appear
normally to enter forest. Aédes africanus and apicoargenteus are the only species
normally found in the heart of the high, closed canopy, equatorial forest, and both of
these extend far into the park savannah; africanus also occurs in the Rhodesian
dry forest; frasert and dendrophilus appear to be restricted to fringing and relict
forests of an undefined type. The remainder are savannah species. The relationship
of the various species to their characteristic plant associations is almost wholly
obscure and will not be understood until much careful work has been done on both the
entomological and the botanical side. This subject will be discussed in a later paper.
ACKNOWLEDGEMENTS
I am especially indebted to Dr. A. J. Haddow of the Virus Research Institute,
Entebbe, and Mrs. E. C. C. Van Someren of the Medical Research Institute, Nairobi,
on whose experience of African mosquitoes I have drawn freely during the prepara-
tion of this paper and who have been most generous in the matter of specimens and
of information. Others who have very kindly supplied me with specimens or with
climatological or topographical data or unpublished distribution records or have
otherwise assisted me are Dr. P. H. Abbott of the Civil Hospital, Li Rangu, Sudan;
Lt.-Col. M. S. W. Bisdee of Medical Dept. H.Q., British Military Administration,
Eritrea; Prof. P. A. Buxton and Mr. H. S. Leeson of the London School of Hygiene
_ and Tropical Medicine ; Prof. G. D. H. Carpenter and Prof. G. C. Varley of the Hope
Department, Oxford; Dr. G. Ceballos and Dr. S. V. Peris of the Instituto Espafiol
de Entomologia, Madrid; Dr. L. J. Chwatt of the Yellow Fever Research Institute,
Yaba, Lagos; Prof. T. H. Davey, Prof. R. M. Gordon, and Dr. F. O’Rourke of the
Liverpool School of Tropical Medicine ; Monsieur A. M. L. Deleplanque of the Service
de l’Hygiéne Publique, Luluabourg, Belgian Congo; Dr. B. De Meillon and Mr. J.
Muspratt of the South African Institute for Medical Research, Johannesburg ;
Dr. A. Duren, Inspecteur Général de l’Hygiéne, Ministére des Colonies, Brussels;
Mr. A. W. Exell and Dr. Hugh Scott of this Museum; Dr. A. Fain of the Congo
Medical Service ; Monsieur M. Holstein of the Service Général d’Hygiéne, Bobo-Diou-
lasso; Lt.-Cmdr. K. L. Knight of U.S. Naval Medical Research Unit No. 3, Cairo,
Egypt; Dr. W. A. Lamborn of Fort Johnston, Nyasaland; Mr. D. J. Lewis of the
Gezira Research Farm, Wad Medani, Sudan ; Monsieur M. Lips of the Section d’Etudes
Antimalariennes, Elisabethville ; Dr. W. H. R. Lumsden and Mr. J. D. Gillett of the
Virus Research Institute, Entebbe; Mr. C. V. Meeser of the Research Laboratory,
Salisbury, S. Rhodesia; Mr. H. Merlin of Cirencester, Gloucestershire ; Prof. F. Peus
of the Zoologischen Museums der Universitat, Berlin ; Mr. H. St. J. Philby of Jeddah,
Saudi-Arabia; Monsieur J. Rageau of the Service de l’Hygiéne, Yaoundé, Fr. Came-
roons ; Mr. G. G. Robinson of the Department of Health, N. Rhodesia ; Prof. J. Schwetz
of the Université Libre de Bruxelles ; Mr. A. Smith of the East African Medical Survey
and Filariasis Research, Mwanza, Tanganyika; Dr. A. Stone of the U.S. Dept. of
IN THE ETHIOPIAN REGION 295
Agriculture Bureau of Entomology, Washington, D.C.; Mr. G. Swaine of the Agri-
cultural Entomological Laboratory, Morogoro, Tanganyika; and Monsieur J. Wolfs
of the Service de l’Hygiéne, Costermansville, Belgian Congo. Among many who have
indirectly supplied me with information I owe a special debt to the librarians of the
Commonwealth Institute of Entomology, the Imperial Institute, the Royal Air Force
Meteorological Library, and the Science Museum, London.
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296 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
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——aE
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— 1903. Report on a collection of mosquitoes or Culicidae, etc., from Gambia and a descrip-
tion of a new species. Mem. Lpool. Sch. trop. Med. 10, App.: i-ix
1905. New Culicidae from India, Africa, British Guiana and Australia. J. econ. Biol. 1:
18-36.
—— 1906. Report on economic entomology. Pt. 1, Rep. Wellcome trop. Res. Lab. 2: 67-83.
— 1907. A monograph of the Culicidae of the world, 4, London.
1909. Description of the new mosquitoes collected by Dr. Graham in Ashanti. Colon. Rep.
misc. Ser., Lond. 287.
— 1910. A Monograph of the Culicidae of the World, §, London.
— 1g911a. Uganda Culicidae, including thirteen new species. Novae Culicidae, 1: 9-35.
— 1911b. The Culicidae or mosquitoes of the Transvaal. Rep. vet. Res. S. Afr. 1.
—— 1912. A new species of Culicidae. Rev. zool. afr. 2: 78.
— 1913. New Culicidae from the Sudan. Amn. trop. Med. Parasit. 9: 591-602.
TouMANOFF, C. 1937. Essais préliminaires d’intercroisement de S#. albopicta Sk. avec St.
argentea Poiret. Bull. Soc. méd.-chir. Indochine, 15: 964-970.
1938. Nouveaux faits au sujet de l’intercroisement de St. albopicta Skuse avec St. argentéa
(St. fasciata) Theob. Rev. méd. frang. Extr.-Orient, 17: 365-368.
1939. Les races géographiques de St. fasciatus et St. albopictus et leur intercroisement. —
Bull. Soc. Path. exot. 82: 505-509.
1949. L’hémophagie variée et l’activité reproductrice chez Aédes aegypti L. et Aides
albopictus Skuse. Ibid. 42: 466-470.
1950. L’intercroisement de l’Aédes (Stegomyia) aegypti L. et Aédes (Stegomyia) albopictus
Skuse. Observations sur la mortalité dans la descendance des générations hybrides F, et Fy
de ces insectes. Ibid. 48: 234-240.
VANDENPLAS, A. 1943. La pluie au Congo Belge. Mém. Inst. R. mét. 16.
VAN SOMEREN, E. C. C. 1946. Ethiopian Culicidae: notes and descriptions of some new “=
and hitherto unknown larvae and pupae. Tvans. R. ent. Soc. Lond. 96: 109-124.
Van SoMEREN, G. R. C. 1943. Notes on the mosquitoes of British Somaliland. Bull. ent. Res.
84: 323-328.
Wanson, M., & Nicotay, F. 1937. Biologie de Culex fatigans dans le Bas-Congo. Aun. Soc.
belge Méd. trop. 17: 111-122.
Wescut, W. 1910. On the larval and pupal stages of West African Culicidae. Bull. ent. Res. 1:
7-59.
WuitE, M. J. D. 1949. Cytological evidence on the phylogeny and classification of the Diptera.
Evolution, 3: 252-261.
WHITFIELD, F. G. S. 1939. Air transport, insects and disease. Bull. ent. Res. 80: 365-442.
WizBeErt, R., & DELorME, M. 1927. Sur un spirochétose ictérohémorragique du Chimpanzé
transmissible 4 ’homme. Ann. Inst. Pasteur, 41: 1139-1155.
ie aes
IN THE ETHIOPIAN REGION 303
Wiseman, R. H., Symes, C. B., McManon, J. C., & TEESDALE, C. 1939. Report on a malaria
survey of Mombasa. Nairobi.
Woke, P. A. 1937. Comparative effects of the blood of different species of vertebrates on egg
production of Aédes aegypti. Amer. J. trop. Med. 17: 729-745.
Wo rs, J. 1946. Note sur les moustiques de Coquilhatville (Deuxiéme note). Ann. Soc. belge
Méd. trop. 26: 95-104.
— 1949. Aédes (Stegomyia) apicoargenteus var. denderensis var. n. (Culicidae). Rev. Zool.
Bot. afr. 42: 190-192.
ZeuNER, F. E. 1944. Dating the past. London.
ZumptT, F. 1937. Stechmiickenstudien im Pflanzungsgebiet des Kamerunberges. Tropenpflanzer,
40: 366-383.
APPENDIX
SINCE going to press, a number of specimens have been received which are of so much
interest from the point of view of distribution that they are felt to warrant special
mention. The new records are as follows:
Aédes apicoargenteus. TANGANYIKA. Ukara Island (B.M.).
Aédes africanus. KENYA. Taveta (B.M.).
Aédes dendrophilus and Aédes simpsont. B. Conco. Elisabethville area (B.M.).
The records of apicoargenteus and dendrophilus are each based on a single incomplete
female and are therefore subject to confirmation. Both specimens are, however, in
quite good condition and agree so closely with the types that little doubt is felt as
to their identity. Of the three records the most startling is that of Aédes africanus
of which several good specimens have been sent by Dr. Lumsden. Taveta has a mean
annual rainfall of only 26 inches with five dry months and is thus far outside the limits so
far recorded for africanus. The explanation, suggested to me by Dr. Haddow, appears
to be the presence in this locality of ‘dense, humid, evergreen forest’ depending, not
on rainfall, but on ‘innumerable springs rising through the volcanic ash’. From this
it is clear that the presence of africanus in this locality is dependent, not on the local
rainfall, but on a subsoil water content conditioned by the rainfall some considerable
distance away. It is tempting to infer from this that the 40 inches or so which are
normally required are necessary rather for the maintenance of a suitable type of
forest than for the provision of adequate supplies of water in the breeding-places. It
must, however, be remembered that under the conditions described humidity will be
high and temperature low, and evaporation from tree-holes within the forest will be
correspondingly reduced. Accordingly the effective rainfall will be higher than in
other localities where the actual rainfall is similar. Nevertheless it would seem that
this most interesting discovery lends strong support to the belief, elsewhere expressed,
that rainfall is primarily of importance for its indirect effect on vegetation. The
Taveta population is quite clearly an isolated relict one comparing in this respect with
those in the Sidamo Province, of Abyssinia and, perhaps, in the Zomba area and
adjacent parts of Mozambique.
The record of apicoargenteus from Ukara Island, for which I am indebted to Mr.
Smith, extends the known distribution of the species in this area very much to the
south. It does not in any way affect the empirical rainfall limits deduced from
304 THE SUB-GENUS STEGOMYIA (DIPTERA, CULICIDAE)
existing data, but it suggests the possibility of an interesting comparative study on
seasonal distribution here and in the neighbouring border-line area around Mwanza.
The record of dendrophilus from Elisabethville, sent by Monsieur Lips, comes from
;
far outside the known rainfall limits of the species. It seems that it may well be ©
explained by the presence of extensive gallery forest in this region, in which case the
record would be comparable with that of africanus from Taveta. Further details on
this point are awaited from the collector, but it is known that Aédes chaussieri and
masseyt have recently been found in gallery forest in this area (Lips in litt. and speci-
men of chaussieri now in the British Museum). This record, if confirmed, would seem ~
to render almost certain the presence of dendrophilus in the South Congo savannah as
suggested above.
The record of stmpsont from the Elisabethville area does not appear to call for
special comment.
PRESENTED
1 7 APR 1952
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A REVEEW, AND A REVISION
IN GREATER PART, OF
meee CLENISCINI OF THE
OLD WORLD
(HYM., ICHNEUMONIDAE)
G. J. KERRICH
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. II, No. 6
LONDON::1952
A REVIEW, AND A REVISION
IN GREATER PART, OF
2nE CTENISCINI OF THE OLD WORLD
(HYM., ICHNEUMONIDAE)
BY
G. J. KERRICH
(Commonwealth Institute of Entomey
Pp. 305-460; Pls. 4-7; 86 Text-figs.
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. II, No. 6
LONDON: 1952
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is issued in
jive series, corresponding to the Departments of the Museum.
Parts appear at irregular intervals as they become ready.
Volumes will contain about three or four hundred pages, and
will not necessarily be completed within one calendar year.
This paper is Vol. II, No. 6 of the Entomology series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued December 1952 Price Thirty Shillings
A REVIEW, AND A REVISION IN GREATER
Pani, OF THE CTENISCINI OF THE OLD
WORLD (HYM., ICHNEUMONIDAE)
By G. J. KERRICH
COMMONWEALTH INSTITUTE OF ENTOMOLOGY, AND HONORARY ASSOCIATE OF THE BRITISH MUSEUM
(NATURAL HISTORY)
[Received for publication on 10 May 1951]
INTRODUCTION . ‘ ‘ : : j : , : : : : = fs 21307
DESCRIPTIVE TERMS . ‘ ‘ : . : ‘ : ‘ ; ‘ ‘ ‘ - 309
DISCUSSION OF SOME CHARACTERS, AND OF METHODS ‘ E é < 2 - « “311
DISCUSSION OF GENERA 5 : : > hay . : : : : é ; + 313
Kery To GENERA : : “ : - 3 ‘ : , : : r ; s 316
GENUS ACROTOMUS HOLMGREN . : ; : : : - A é : : - 319
GENUS ANISOCTENION FORSTER . : : F ; A ; 2 : A p + 325
Key to species of Anisoctenion . : , ‘ : . , , . . . - 350
Genus EXENTERUS HartTIG : : ; ‘ ; : ; ; : ‘ - - 354
Key to species of Exenterus : : : : : : : : é ; . - 364
Grnus EXYSTON SCHIODTE . . : ‘ F ; F 3 is F : - ae 307
Key to species of Exyston : ‘ : : : : : : : : ; - 389
GENuS PAREXYSTON, GEN. NOV. . : ; : : : : ‘ : ; : « 392
GENuUs SMICROPLECTRUS THOMSON ‘ ; : : 4 ; ‘ : : ‘ - 393
Key to species of Smicroplectrus : : : ; : : : : ° . - 412
GrENnus EUDIABORUS, GEN. NOV. ‘ , : 3 ; : 2 5 ; 3 wats
GENUS CTENISCUS (Hartipay MSS.), Curtis, THOMSON . ‘ : : : : é . 420
Key to ‘black-bodied’ species of Cteniscus . : : , : ‘ : ‘ ; - 443
CONCLUSION : : ‘ : : . : : : ‘ : ; ; : - 451
SUPPLEMENT : ; A , ; ; F 5 : 5 : : : : . Ad2
REFERENCES. : : : ; : E : : : : : : : - 455
APPENDIXES : : . : : ; : : : : : : : : - 458
SYNOPSIS
The systematic position and characters of the tribe are briefly reviewed, and there follow discussions of
some descriptive terms and some characters used in this study. The genera are discussed from the points
of view both of taxonomy and nomenclature, and a new key to genera is given, taking account of the
Old World species and of those from the New World known to the author. The genera Anisoctenion,
Exyston, Smicroplectrus, and the ‘black-bodied’ species of Cteniscus are fully revised, with descriptions
of old and new species and with keys. Much fresh European information on the genus Eventerus is given,
with a key to the Old World species. The remaining sections are less fully treated. Altogether two new
generic names are proposed, and ten new species are described, with one new variety and one aberra-
tion named. Some new synonymy is given. The paper is illustrated by eighty-six text-figures and
four plates.
INTRODUCTION
THE subfamily Tryphoninae, to which the Cteniscini belong, has recently been re-
defined by Townes & Townes (1949) to include all those ectoparasitic genera in which
there is a large or rather large stalked egg, of which only the anchor passes down the
308 A REVIEW AND A REVISION OF
channel of the ovipositor. The ovipositor (stylet) is not dorsally notched near the
apex, or only occasionally weakly so. Thus the subfamily now includes the Cteniscini,
Tryphonini, Sphinctini, Eclytini (= Eclytus+the ‘Thymaridini’) and'a number of
tribes mostly not included by the earlier authors ; and it excludes not only the Meto-
piinae and Diplazoninae, but also almost the whole of the ‘Mesoleptini’ of authors.
The steps leading to this position were reviewed by Kerrich (1942, pp. 64-65).
The Cteniscini differ from the other Tryphonine tribes, as from all other Ichneu-
monidae, in having the middle tibia with a single spur, and the hind tibia without
a spur or with a very short one, much shorter than the apical breadth of the tibia
where it issues. Usually the female of the species has more bright coloration than
the male, whereas in Ichneumonidae generally the reverse is the case. They are
most closely related to the genus Polyblastus of the Tryphonini, as recorded by
Thomson (1883) and by Townes & Townes (1949).
As in other Tryphoninae, the female of the species is often to be found carrying
an egg on the ovipositor. In the genera Exyston and Smicroplectrus the egg is often
seen to be nearly or quite ready to hatch. This recalls the case of Polyblastus (see
Kerrich, 1936), but only a single egg is carried at a time. It argues that a female
individual must be long-lived to deposit all her eggs.
I have been able to see a vastly greater amount of material of this tribe than has
ever before been studied by any one taxonomist in Europe; and the confidence with
which species have been separated or synonymized, and the assessment made of the
range of variation within species, could not have been attained in any other way.
For the loan, or in a few cases the gift, of such material I am much indebted to
Dr. Kjell Ander, Dr. L. Berland, Dr. J. G. Betrem, Dr. H. C. Bléte, Dr. Z. Bouéek,
Mr. H. Britten, Monsieur E. Cavro, Professor G. Ceballos, Monsieur A. Collart through
co-operation of Dr. J. Leclercq, Mr. H. W. Daltry, Mr. E. A. Ellis, the late Dr. E. O.
Engel, Dr. Ch. Ferriére, the late Professor Fr. Gregor, Mrs. F. H. Haines, Rektor K.
Hedwig, Herr Gerd Heinrich, phil. Mag. W. Hellén, Mr. W. D. Hincks, Herr Rolf
Hinz, Dr. R. Krogerus, Dr. Th. Kupka, Professor Dr. O. Lundblad, Herr B. Maixner,
Dr. R. Malaise, Dr. L. Masi, Professor H. W. Miles, Mr. C. Morley, Lektor A.
Nordstrém, Mr. J. F. Perkins, the late Herra A. Saarinen, Dr. Erich Schmidt, Mr.
A. W. Stelfox, Dr. H. G. M. Teunissen, Professor E. Tortonese, Rev. C. E. Tottenham,
Dr. H. K. Townes, and Professor T. Uchida.
Outstandingly valuable to me have been the collection of the British Museum
(Natural History), which I was also privileged to study before obtaining my present
professional position, the Wesmael-Tosquinet collection of the Royal Natural History
Museum, Brussels, and the collection of Mr. A. W. Stelfox. Dr. Th. Kupka’s collection
was especially helpful in the earlier stages of the work, as were specimens given me
by Rektor K. Hedwig, which I could compare with Gravenhorst specimens and then
retain. Dr. H. K. Townes and Mr. W. R. M. Mason have kindly lent me nearctic
material, which has better enabled me to assess generic limits.
The material is mostly in the private collections of the persons mentioned, or in
the museum collections of which they have charge, except that some specimens have
now been presented to or exchanged with the British Museum (Natural History).
The British Museum contains (apart from some British material sent in exchange to
THE CTENISCINI OF THE OLD WORLD 309
Cambridge in 1931) the J. F. Stephens, T. Desvignes, T. A. Marshall, Ruthe,
Buchecker, and R. von Stein collections; and all material here recorded as collected
by Dr. R. C. L. Perkins, Mr. and Mrs. J. F. Perkins, Mr. and Mrs. R. B. Benson, and
Mr. G. E. J. Nixon. Other Swiss material is mostly from the Geneva museum ; while
the Paris museum contains the collections of O. Sichel, J. Giraud, J. de Gaulle, and
A. Seyrig. The specimens in the de Gaulle collection were mostly without determina-
tion labels, and so required re-identification.
I also wish to thank those who have received me in their homes, or in the museums
of which they have charge. My visits to Breslau (Wroclaw) were made in 1936 and
1938, and the observations and lectotype selections here recorded were mostly made
in the latter year.
I wish to express my particular gratitude to the late Dr. A. Roman, with whom
my first real progressin the study of the Ichneumonidae, particularly the Tryphoninae,
was made; to Mr. J. F. Perkins, whose incisive comment has frequently illumined
the passage of this work; and to Mr. W. R. M. Mason, who is revising the nearctic
species, for his co-operation in matters of mutual concern.
Herr Rolf Hinz kindly suggested my incorporating his observations on the biology
of three of the species (two of which he had already identified), and was so good as
to add that, so placed, they would be most readily found by future workers.
I am indebted to Mr. R. B. Benson for much information on the sawfly hosts of
these insects, and on problems of distribution.
I wish to thank Dr. T. H. C. Taylor for reading my manuscript and advising on a
number of points, and Mr. N. D. Riley for discussing with me the complicated
problems of nomenclature.
Text-figures I to 5 and the four beautiful plate figures were drawn by Mr. Arthur
Smith, and I wish to express my appreciation of the artist’s skill in executing them.
In so far as material studied was captured on an expedition to Finland in 1935, this
paper is a result of that expedition, of which the author’s costs were defrayed equally
by the Royal Society and by the Balfour Managers (Cambridge).
DESCRIPTIVE TERMS
It has been my constant endeavour to avoid descriptive terms that are morpho-
logically inaccurate (e.g. 1942, pp. 43-45). Despite advancing knowledge the diffi-
culty remains, however, that it is still not possible to delimit all morphological areas ;
and, in any case, the entities of taxonomic significance do not always correspond with
the morphological entities. The following points seem to call for comment in intro-
ducing this paper.
Considering the facial aspect of the head, writers on Ichneumonidae have termed
that part between the median ocellus, eyes, and antennae the ‘frons’, and that part
between the antennae, eyes, and clypeus the ‘face’. In 1942 it was proposed to
follow Pratt (1940) by substituting ‘ postfrons’ and ‘antefrons’ respectively: actually,
however, Pratt restricted the term ‘antefrons’ to the more or less raised, median part,
and excluded the orbital regions (parafrontals). After consultation with other
310 A REVIEW AND A REVISION OF
specialists working in the British Museum (Nat. Hist.), I now use the terms ‘upper
face’ and ‘lower face’ respectively, and retain ‘epistoma’ as part of the latter.
The term ‘genal costa’ is retained as applied by earlier authors (Thomson, 1873),
but ‘oral costa’ is replaced by the more appropriate “hypostomal costa’ (Michener,
1944). These structures are of particular importance in the Cteniscini.
It was pointed out in 1942 (pp. 43-44) that earlier authors ignored the annellus at
the base of the antennal flagellum, and the practice was then started of including
it in reckoning the number of flagellar segments. The objection is that it is not always
clearly defined in the Braconidae, and it is uncertain whether it should be regarded
as a separate segment or as a secondary specialization. I have now agreed with
J. F. Perkins to treat the flagellum as having a basal segment composed of an annellus
and a postannellus: these, then, together constitute the first flagellar or third antennal
segment. An analogous case is that of the trochanterellus, which is regarded by
morphologists as a specialized basal part of the femur.
The pair of ventrally placed plates of the prothorax were thought by earlier workers
to represent a divided prosternum (Pfankuch, 1919). O. W. Richards, in a paper on
the Sphecoid genus Trypoxylon (1934), termed such plates the proepisterna ; but he
tells me it is uncertain whether this is morphologically accurate throughout the
higher Hymenoptera, and it seems safer to call them the propleura. It is well known
that in higher Hymenoptera the true prosternum is greatly reduced and usually
largely concealed.
The main plate of the mesonotum, anterior to the scutellum, was simply termed
the mesonotum by earlier workers. A nearer approach to accuracy is now achieved
by calling it the mesoscutum, but it must be realized that it includes the mesopre-
scutum. This last is supposed by some workers to be the area between the notaulices ;
but this interpretation cannot be regarded as proven and, in any case, such area is
not defined posteriorly.
Snodgrass (1935) introduced the term ‘notaulices’ as being etymologically more
correct than ‘notauli’, and the present writer adopts this change and also substitutes
‘sternaulices’ for the old ‘sternauli’. The ventral plate of the mesothorax, between
the sternaulices, is still termed the mesosternum ; for it is necessary to refer to this
area, whether it corresponds with the true mesosternum or not.
The ‘scutellum metathoracis’ has been called by most British systematists the”
‘postscutellum’, a term employed in a different sense by morphologists. The structure
is here referred to as the ‘metascutellum’.
It has long been realized that there are two body-segments between the mesothorax
and the waist ; and the use of the term ‘propodeum’ (or ‘propodaeon’) for the true
first abdominal segment is well-established practice. It may, however, be worth
pointing out that the ‘area pleuralis’, though treated with the propodeum in taxo-
nomic descriptions, seems really to belong to the metapleuron (Pratt, 1940).
The term ‘gaster’ is again used to include the whole of the body posterior to the
propodeum (see Kerrich, 19400), and thus corresponds exactly with the ‘abdomen’
of earlier authors. It was termed the ‘metasoma’ by Michener (1944).
As has been pointed out to me, the ‘ovipositor sheaths’ (Bohrerklappen of German
authors) do not always sheath the ovipositor. I continue to refer to these structures
THE CTENISCINI OF THE OLD WORLD 311
as the ‘third valvulae’ (Snodgrass, 1935), although this term is not in general use
among systematists, and although it has also been pointed out to me that they are
not morphologically equivalent to the first and second valvulae.
The term ‘nervellus’ is applied here, as by most authors, to a transverse vein of
the hind wing; and reference is made in descriptions to where it is intercepted by
a longitudinal vein above, below, or about in the middle. Morley (1903) applied this
term to the longitudinal vein; but he changed this practice in his four succeeding
volumes.
The trochanterellus is treated as a separate entity and, in descriptions, is not
included in either the trochanter or the femur.
DISCUSSION OF SOME CHARACTERS, AND OF METHODS
Just as, in the main, the terms used in this paper follow those of previous authors,
so, in the main, do the characters used in describing species and in separating them ;
though the attempt is always made to improve the precision with which they are
used. Two sets of characters, however, appear to be quite new, and a third set has
been greatly developed in comparison with previous work. Some characters are found
to be more variable and therefore less valuable than has often been supposed.
It has already been stated that the form of the genal and hypostomal costae is of
particular importance in the Cteniscini ; and indeed this may apply to the whole form
of the gena. The use of these characters was begun by C. G. Thomson and A. Roman;
but they did not carry them very far. Recently Townes & Townes (1949) have found
them of use, particularly in the genus Monoblastus. They did not, however, illustrate
them ; nor did they find such striking differences as are now disclosed in palaearctic
Exyston and the two genera most closely related to it. Palaearctic Smicroplectrus
must be considered as in the majority of cases indeterminable if the hinder gena
cannot be viewed.
The shape of the antennal scape does not seem to have been studied previously.
This varies to some extent, as do other characters; but I find it of such taxonomic
value that the European species of Anisoctenion and Exyston could almost be deter-
mined on their antennal scapes alone. The differences in Exenterus are of lesser degree,
though very serviceable ; but in Smicroplectrus there seems to be greater variability,
and I cannot use them for diagnosis except in one species.
The development of a pair of piliferous lobes on the anterior angles of the meta-
notum, recalling the cenchri of sawflies, has not previously been recorded. They are
well developed in species of Exyston and Smicroplectrus studied ; whereas in Cteniscus
and in Diaborus auctt. they are represented by much weaker, simple projections,
which may to a lesser degree be piliferous. To this extent the character has been used
as an aid to generic diagnosis. The same structures are developed in Exenterus
species, though not so strongly as in Exyston and Smicroplectrus. Intermediate stages
are to be found in Acrotomus and Anisoctenion which, in this respect, are less con-
sistent than the other genera mentioned.
Holmgren (1858) used the presence or absence of an ‘apical’ fovea on the scutellum
(i.e. as distinct from the normal basal scutellar fovea) three times in the course of his
312 A REVIEW AND A REVISION OF
dichotomous key to the species included in Exenterus ; and Schmiedeknecht followed
by using it to separate two quite large groups of species in Cteniscus. The greatest
weight assigned to this character was by Davis (1897), who based a new genus
Excavarus upon an admittedly larger excavation than I have seen in any European
insect. Walley (1937) has already discussed this matter, and has shown that Exca-
varus is not a valid genus. No doubt each species has its own measure of weakness of
the cuticle of the disk of the scutellum, resulting in a tendency to buckling ; and this
tendency is sometimes mentioned in descriptions in the present work. It is necessary,
however, to find quite other characters for use in dichotomous keys.
It was colour characters that were most used by the older authors to distinguish
insect species. Although it has long been recognized that these did not provide
a sufficient basis for species discrimination, it is my firm conviction that they should
not be rejected out of hand. A colour difference between two forms, if consistent
throughout an adequate series of specimens, may indicate that structural differences
should be sought. It may often happen that, when two species have been placed in
synonymy because they have been found to differ only in one or more apparently
trivial colour characters, the synonymized species will have to be reinstated when
structural differences have been discovered.
In the present work the coloration of the species has been studied almost as
intensively as the structure. My conclusion is that each species, or sex of a species,
has a colour pattern that is fundamentally consistent however much it may vary,
firstly in the degree of development and the intensity of the brighter colours, secondly
in the extent and intensity of infuscation, and thirdly in the intensity of the melanic
pigment in the parts of the cuticle that are regularly black or blackish.
Colour characters, then, must be used with discretion. They vary with type of
locality, and also quite independently of locality. A character very frequently used
in the past has been the presence or absence of a large apical pale mark on the
scutellum. This proves consistent in the separation, as good species of Cteniscus, of
two forms that previously have been treated as varieties; yet such mark may be
absent or quite distinct within another species of Cteniscus and also within one of
the new species of Smicroplectrus. When the coloration of a species has been investi-
gated with the aid of adequate material, it can be seen which colour characters are
sound, and can be used as additional means of distinguishing the species. When that
stage has been reached, it also becomes possible to give probable interpretations of
older colour diagnoses, as has also been done in the present paper.
Where proportions are quoted in descriptions, they have been calculated from
micrometer measurements. It is surprising how difficult it is to estimate length to
breadth accurately by eye, and allowance should on this account be made for possible
inaccuracy in older descriptions. The proportions of the petiolar segment have been
measured in most of the specimens from which descriptions and redescriptions have
been made, particular care being taken not to overlook extreme cases. Proportions
of the radial cell have been measured on a fair sample of the specimens, the others
then being compared with these.
Figures are drawn from female specimens unless the contrary is stated.
The synonymic tables are not intended to be complete, and for some species are
THE CTENISCINI OF THE OLD WORLD 313
not given at all. References are given in all cases where the synonymy is new, or has
been confirmed by reference to the types; and also in some cases where this is desir-
able to explain current usage, or where the reference is recent, or contains information
to which attention is particularly drawn.
For locality records, the island of Ireland is treated as a single zoo-geographical
unit. For other countries the boundaries taken are those that existed in 1937, as in
most cases these give the country in which the locality was when the recorded speci-
men was captured. No political implications should be attached to this procedure.
DISCUSSION OF GENERA
The genus Anisoctenion was, as noted by Schmiedeknecht (1911), founded by
Forster for species differing from Acrotomus in having the clypeus broadly apically
rounded and the claws strongly pectinate. He adds that the first character is not
sufficient to separate the genera, for the last three species placed in Acrotomus, i.e.
rubiginosus Grav., binotatus Thoms., and parvulus Thoms., have the clypeus more
rounded than truncate ; and that, if the second difference is insufficient, then Aniso-
ctenion can be united with Acrotomus. Certainly this second difference is insufficient,
for a number of species placed in Acrotomus by Schmiedeknecht have the claws more
or less strongly pectinate. However, the form of the clypeus is very characteristic
in the lucidulus—succinctus group. I propose to restrict the genus Acrotomus to this
group, and to transfer all the other species generally placed in Acrotomus to the genus
Anisoctenion (see below), though laetus Grav. may perhaps be considered as a tran-
sitional species.
Viereck (1912) designated lucidulus Grav. as genotype of Acrotomus Hlgr. 1858, but
Schmiedeknecht (1911) gives lucidulus Hlgr. = succinctus Grav. If lucidulus Hlgr. is
not the same species as lucidulus Grav., then lucidulus Grav. was not an originally
_ included species, and Viereck’s type selection would be invalid. As Roman has
already pointed out (1914, 1917), Holmgren had these two species mixed. Roman
and I have since (1939) examined Holmgren’s specimens, and find that all those of
lucidulus Grav. are referable to Holmgren’s varieties 2 and 3. We selected female and
male lectotypes of lucidulus Hlgr., specimens from Smaland, and these are succinctus
Grav. as given by Schmiedeknecht.?
The genera Exyston and Smicroplectrus have been treated by authors as though
very distinct. Smucroplectrus has, at. least in Europe, been treated as though its
species were scarcely to be distinguished except by colour differences, and it was not
until 1913 that it was noted by Roman that guinquecinctus Grav. has the genal and
hypostomal costae strongly elevated as in Exyston. It has not previously been noted
that guinquecinctus Grav. has the epicnemial carina elevated, though much less
strongly than in that genus, and that pratorum Woldst. (= brevipetiolatus Thoms.)
* The difficulty that arises over the identity of the type species of Acrotomus is due to the misidentifica-
tion of a species. Such cases were considered by the International Commission on Zoological Nomenclature
at its Paris meeting in 1948 (1950, Bull. Zool. Nomenclature 4: 158-159, 6th Meeting, minute 38). I there-
fore propose to invite the Commission to consider this case, and in the meantime to treat Acrotomus
succinctus Gravenhorst (= Acrotomus lucidulus Holmgren, 1858, non Gravenhorst 1829) as the type
species of the genus.
ENTOM. 2, 6 Qq
314 A REVIEW AND A REVISION OF
has vomeriform third valvulae and finely pectinate claws. The generic characters
were found so to intergrade that it proved difficult to decide at which point the species
should be separated into genera ; in fact, it might have seemed more natural to place
the whole series of species into one genus were it not that the extreme forms, cinctulus
Grav. and jucundus Hlgr., would have so little in common. At one time I thought it
would be better to restrict the genus Exyston to the species with long petiole and
simple claws, and to transfer pratorum Woldst. to Smicroplectrus, which would become
characterized by the short petiole and pectinate claws; but the discovery of the
species which has proved to be phaeorrhaeus Hal., and the subsequent examination
of albicinctus Grav., species largely intermediate between genalis and pratorum,
dispelled this notion. It seems best to erect a new genus for calcaratus Thoms. This
species has been studied independently, from two Finnish specimens, by Hellén
(1944), who suggested that it might be better placed in a separate genus ‘die in die
Nahe von Exyston und Smicroplectrus zu stellen ware’, but did not mention the form
of the hypostomal costa. Hellén referred to Schmiedeknecht, but the latter’s sug-
gestion (1912) was for a genus to contain pratorum as well as calcaratus.
Walley (1937) considers Excavarus Davis not to be a valid genus. He places
E. annulipes Cress. in Smicroplectrus and E. viticollis Cress. in Diaborus auctt.
Hincks (1944) has most opportunely reopened the question of the status of the
names Cteniscus (Haliday MSS.) Curtis and Exenterus Hartig, and the further
question as to whether the two genera, as generally understood by authors, can be
maintained as distinct.
It is necessary to dispose of the second question in the first place. As observed by
Schmiedeknecht (1911), Holmgren (1858)! distinguished only three genera: all species
not referable to Exyston or Acrotomus he placed in Exenterus. Forster’s work (1868)
was modified by Thomson (1883), who produced the first satisfactory table of genera.
The citations of Dalla Torre (1901-1902) and of Viereck (1914) were, no doubt,
derived more from a study of the literature than of the insects. Hincks, though
himself an accomplished student of Ichneumonoidea, has purposely avoided duplicat-
ing my study of the Cteniscini, and his citations here and in the ‘Check List’ (Kloet &
Hincks, 1945) are derived from his interpretation of existing literature. Morley (1911:
203) has united the two genera; but I have no hesitation in following Thomson’s
interpretation and maintaining them as distinct as, in particular, Roman, Schmiede-
knecht, and Cushman have done.
Accepting, then, the two genera in the Thomsonian sense, we have to consider the
status of the name Cteniscus. Hincks writes: ‘Personally I am inclined to regard the
genus Cteniscus as dating from 1832 (Haliday in Curtis), and C. aurifluus Haliday is
therefore the genotype by original designation.’ The same view is implied by Morley
(IQII: 204).
Haliday published diagnoses of his genus Cteniscus and of his three new species
in October 1838. Westwood (1840) listed the species and selected curtisit Hal. as
™ Most authors incorrectly quote Holmgren 1855, the date on which this paper was communicated
to the Academy. The paper actually appeared in two parts, one included in each of the two parts of the
first volume of this series, being the parts issued for the years 1855 and 1856 respectively. The whole
volume is dated 1858, which is now quoted as the date of publication.
ae
THE CTENISCINI OF THE OLD WORLD 315
type. This species has been very little understood: in the present paper its identity
is clarified but, as shown below, other events stand prior to its selection as type by
Westwood, and earlier by Curtis (1837). It is at the present time reckoned as con-
generic with aurifluus Hal., but I do not regard it as typical of the genus as a whole.
Curtis’s selection was an evident oversight, for he had already stated that aurifluus
was the type, and in any case it was prior to the valid publication of the species then
selected.
It is from the second edition of Curtis’s Guide (1837) that publication of Cteniscus
Hal. has usually been dated. The genus is there listed (column 98) with the three
species subsequently diagnosed (October 1838). Later authors, in thus validating
the genus, were, no doubt, under the misapprehension that the diagnoses of the
species had been published previously. It appears to me that the publication in
Curtis’s Guide, ed. 2, cannot be valid unless aurifluus Hal. can be accepted, with its
genus, as valid from 1832. This depends upon whether ‘occurs on Willows from
July to Sept.’ can be accepted as a valid publication of the genus and species.
As stated by Hincks (1944), the acceptance of this standpoint would leave the status
quo undisturbed, and Cteniscus would take as genotype by original designation
the species aurifluus Hal., a distinctive species of the typical general form and colour
prevalent in the genus.
Since these considerations were first committed to paper, it has been pointed out
to me by W. R. M. Mason, who is studying the New World species of this tribe, that,
if the publication of the species aurifluus Hal. be not accepted as valid from 1832,
the genus Cteniscus would nevertheless be validated on the same page by inclusion
of the described species sexlituratus Grav., which was mentioned as type species by
Stephens (1835: 231). We cannot know for certain what species this is until the genus
to which it belongs has been adequately revised and a lectotype selection carefully
made; for it is possible that Gravenhorst’s series was mixed. But we can accept it
from Pfankuch (1906: 92) that it is a Diaborus Forst. Thoms. which, in these circum-
stances, would have to take the name Cteniscus.
The name Diaborus is also in question, for, as W. R. M. Mason and later G. Heinrich
pointed out to me, the type is sedulus Woldstedt 1877 by designation of Viereck
(1912). The collection from which this species was described is presumed to be in the
Leningrad museum. Schmiedeknecht (1912: 2344) placed the species in Cteniscus
Thoms. and stated that the male was unknown. My interpretation of the original
description is that it was made from not less than one female and two males; and
that, though it may refer to some species of Cteniscus Thoms. such as marginatus
Thoms., it contains nothing to preclude Exyston genalis Thoms., which is redescribed
in the present paper.
Much now depends upon whether Cteniscus aurifluus Haliday 1832 can be con-
sidered as valid. In the hope that it can, I propose the name Eudiaborus gen. n., with
type pallitarsis Thomson 1883, for the genus characterized by Thomson (1883: 882,
885) under the name Diaborus Foérst. This name seems appropriate, seeing that
Roman (1939) showed that Thomson applied Férster’s name to the genus that author
actually had.
If aurifluus Hal. 1832 be rejected, then the name Cteniscus must be transferred to
316 ‘A REVIEW AND A REVISION OF
the genus Diaborus Forst., Thoms. non Woldst., and take as type a species not yet
clarified. In that case another name will be required for Cteniscus Hal. Thoms.
Woldstedt’s original material of sedulus would have to be examined and a lectotype
selected in order to determine whether Diaborus Férst. Woldst. were correct or not.
Anecphysis Foérst., Schmiedeknecht (1911: 2281 ; 1912: 2343-2344) might have served,
had not Davis (1897) misused the name for a synonym of one of his own species of
Exyston. But W.R. M. Mason tells me that, from his study of the nearctic fauna, he finds
that the species grouped under a certain other genus of Forster, having line priority
over Diaborus, are not generically separable from Cteniscus auctt.; and herein lies
a possible alternative solution of the problem.
Since going to press I see that Mason has now (1951 April: 227) published his
acceptance of Cteniscus Haliday as valid from 1832,.and has duly placed Evidolius
Forster in synonymy with the genus used in this sense (type Ct. aurifluus Hal.).
He earlier wrote to me (in litt.) that he would not regard pygmaeus Hlgr., type
species of Evidolius Forst., as a satisfactory type species of the whole genus as he now
understands it.
The action of Stephens (1835) in citing Tryphon sexlituratus Grav. 1829 as type of
Cteniscus has never been followed, and to adopt this usage now would cause great
confusion. All things considered, I propose to invite the International Commission
on Zoological Nomenclature, by the exercise of their plenary powers, to set aside the
citations by Stephens of sexlituratus Grav. and by Westwood and Viereck of curtisii
Hal., and either to confirm Cteniscus aurifluus Haliday in Curtis 1832, or alternatively
to select aurifluus Haliday 1838 as type species of Cteniscus Curtis 1832. Meanwhile
I intend to continue the use of Cteniscus in that generally accepted sense.
Reasons are adduced below for placing all interpretations of Picroscopus in syno-
nymy with Exenterus. It is also open to question whether ‘Tricamptus’ apiarius
Grav. merits generic separation ; but, beside the distinctive appearance, there may
‘be a biological difference in this case. The species of Exenterus, as that genus has
recently been constituted, are all, so far as is known, parasites of Diprionidae, while
apiarius has been recorded by Forsius (1911) as occupied with full-grown larvae of
Croesus septentrionalis L. (Nematinae). I was prepared to maintain this species in
a distinct genus so long as I believed such genus to have a valid name; but, as shown
on another page, the name Tvicamptus Foérst. must be applied to pratorum Woldst.
(Exyston). I do not, however, think it opportune to propose a new generic name, and
therefore replace the species in Exenterus where Schigdte and Holmgren had it.
KEY TO GENERA OF CTENISCINI
1. Lower mandibular tooth distinctly larger and more prominent than upper, and
usually much longer (Figs. 1-3). , ; , - ‘ ; \ 2
Mandibular teeth sub-equal (Figs. 4, 5) ; : : ‘ ; i 3
2. Clypeus almost pentagonal, not strongly transverse; apically sub-truncate or,
rather, broadly and shallowly emarginate between the distinct apical angles
(Figs.6,7) . 4 : ‘ ; , ‘ : . Acrotomus Higr.
THE CTENISCINI OF THE OLD WORLD 317
Clypeus sub-elliptical, strongly transverse ; apically more or less rounded, rarely
at all emarginate and not apically angulate (Figs. 8,9) | Anisoctenion Forst.
Fics. 1-5. Right mandible of Fig. 1. Acrotomus succinctus Grav. Fig. 2. Anisoctenion pumilio
Higr. Fig. 3. Anisoctenion triangulatorius Grav. Fig. 4. Cteniscus gnathoxanthus Grav. Fig. 5.
Cieniscus aurvifluus Hal. Drawn by Mr. Arthur Smith. As indicated by the arrow, the direction
of illumination for these five figures is half right, not half left.
3. Area superomedia transverse, generally distinctly so: body rather coarsely, and
for the most part closely punctate ; in particular, the vertical region and temples
of the head are more or less coarsely punctate: hind tibia without, or with a very
minute spur, much shorter than one-third the breadth of the tibia where it
issues: propodeum and first two segments of gaster usually distinctly rugose
318 A REVIEW AND A REVISION OF
(Pl. 4): gaster ere thick-set and often broadly bright yellow-banded
; Exenterus Hartig
Area superomedia very seldom transverse: body rather finely, and generally for
the most part sparsely punctate ; in particular, the vertical region and temples
of the head are more or less finely punctate: in case of doubt the hind tibia
almost always bearing a short spur, whose length is about one-third the breadth
of the tibia where it issues: propodeum and first two segments of gaster usually
coriaceous or shining (Pls. 5, 6, 7), at most partly finely rugulose, or very seldom
in large part distinctly rugose: gaster generally not thick-set and not broadly
bright yellow-banded é ; ; . : : é ; ; 4
> soe
“ — oe
Fics. 6-9. Clypeus of Fig. 6. Acrotomus lucidulus Grav. Fig. 7. Acrotomus succinctus Grav.
Fig. 8. Anisoctenion laticeps Grav. Fig. 9. Anisoctenion alacer Grav.
4. Body strongly hairy: hypostomal costa almost always strongly raised: metathorax
with anterior angles raised to form a pair of piliferous lobes (recalling the cenchri
of sawflies). Hind tibia often bearing a short spur: genal costa and epicnemial
carina strongly raised in most non-spurred species : -
Body not strongly hairy: hypostomal costa not strongly raised: ‘metathorax with- |
out such piliferous lobes. Hind tibia without a spur: genal costa and epicnemial
carina not strongly raised. ; ‘ ; ; , , : : 7
5. Head and thorax rather coarsely and irregularly, and also somewhat sparsely, ~
hairy: petiolar segment often markedly elongate and narrowed: epicnemial
carina strongly raised: hind tibia without a spur in all known palaearctic
species (Pl. 5) . ‘ ‘ Exyston Schigdte —
Head and thorax finely and regularly, and also n more densely, hairy: petiolar
segment not markedly elongate and narrowed: hind tibia with a short spur in
most species (Fig. 62, p. 398) . ; ‘ ‘ : . ‘ . 6
6. Hypostomal costae curved inwards, almost meeting behind mouth-parts (Fig. 59,
P. 393) - ‘ : : : : : : : Parexyston gen. n.
THE CTENISCINI OF THE OLD WORLD 319
Hypostomal costae not nearly meeting behind mouth-parts: epicnemial carina
sometimes distinctly, but never strongly raised . . Smicroplectrus Thoms.
7. Tergite II with a pair of well-defined diagonal impressed furrows at base
Eudiaborus gen. nov.
Tergite II without such well-defined furrows . ; ; ; ; : 8
8. Fore wing with areolet closed ‘ . Cteniscus (Hal. MSS.) Curt., Thoms.
Fore wing with areolet open ; ‘ : : ; Eridolius Forst.
The above table of genera attempts to achieve, so far as possible, a natural classifi-
cation. I maintain the opinion that a fuller understanding of the genera and species
should precede the construction of a ‘practical’ key of a simpler and more artificial
nature, designed for speedier identifications.
It will be seen that the separation of the genera is not such an easy matter as may
appear at first sight. They are based on groups of species with striking combinations
of characters, yet the species will not all fall easily into watertight compartments.
The genera represent, rather, aggregates of species with the tendency to possess such
combination of characters. In each main genus there are one or more species which
approach the unspecialized Cteniscus type: pumilio Higr. in Anisoctenton, pratorum
Woldst. in Exyston, ictericus Grav. in Exenterus, and jucundus Hlgr. in Smicroplectrus.
Conversely, the male of Cteniscus hofferi Gregor approaches Diaborus auctt., while
Ct. unicinctus Higr. is decidedly suggestive of Acrotomus.
Genus ACROTOMUS Holmgren
1858 Acrotomus Holmgren, Svensk Vet.-Akad. Handl. 1: 222.
In this genus as now restricted (see pp. 313, 316) I can, at present, distinguish only
two species, lucidulus Grav. and succinctus Grav.
Roman (1914: 2-3) synonymized sexcinctus Grav. and auriculatus Thoms. with
lucidulus Grav. An intensive study of large material from diverse localities should
reveal whether any valid differences exist. My study of a number of very variable
species within the tribe leads me to expect that all the material I have seen in this
particular aggregate will fall within the range of variation of a single species ; but at
the present stage the concept is being left as that of an aggregate species.
Hellén (1941: 48) doubted the synonymy of auriculatus Thoms., and recorded under
that name a male specimen which he characterized, and which he compared with
triangulatorius Grav., a species that has not, till now, been well understood, at least
under that name (see p. 323). He most kindly sent me this recorded specimen on loan,
which has enabled me to establish that it does belong to the Jucidulus aggregate.
The identity of sexcinctus Grav. Morley is also established in the present paper
(see p. 336). See also my supplement, p. 452 below.
A. succinctus Grav. is abundantly distinct from the lucidulus aggregate, and can be
recognized with the aid of the following table:
320 A REVIEW AND
succinctus Grav.
Clypeus (see Fig. 7)
Antennae scarcely incrassate, with scape elon-
gate and slender (Fig. 11).
Notaulices extending to middle of mesoscutum,
though indistinctly.
Petiolar segment without distinct basal lobes;
less strongly transversely impressed in apical
A REVISION OF
lucidulus Grav. agg.
Clypeus (see Fig. 6)
Antennae distinctly incrassate, with scape
shorter, stouter, and sharply incised (Fig. —
Io).
Notaulices generally short.
Petiolar segment with basal lobes strong ; more
' strongly transversely impressed in apical
third.
i
third.
||
Fics. 10-12. Fig. 10. Left antennal scape and following segments, in sinistro-lateral view, of
Acrotomus lucidulus Grav., drawn from a specimen compared with the type series in 1936. Fig. 11.
The same of Acrotomus succinctus Grav. Fig. 12. Apical segments of gaster of Acrotomus succinc-
tus Grav. in similar view.
Female hypopygium broader, not folded mid-
ventrally, terminating in a short point.
Female hypopygium sharply folded mid-
ventrally, at least in apical quarter.
Female third valvulae broad in ventral view,
and densely hairy beneath ; not or very little
projecting beyond apex of tergite 8.
Female third valvulae narrow in lateral and in
ventral view; projecting strongly beyond
apex of tergite 8 (Fig. 12).
Male gaster seldom pale yellow laterally,
generally in ventral view only, apart from
the hind margins of the tergites.
Male gaster clearly pale yellow in lateral view,
before the reflexed lateral margins of the ter-
gites.
Acrotomus lucidulus Gravenhorst
Pfankuch (1906: 93) states that there are two females in the Gravenhorst collection:
this is correct if intended to exclude the ‘varieties’. Of these two I select as lectotype
the specimen in which the pronotum does not appear to have a continuous antero-
fs ry
Bee
THE CTENISCINI OF THE OLD WORLD 321
lateral yellow margin: the first in the collection. Pfankuch states ‘Var. 2 fehlt’, but,
though not so labelled, two specimens are present.
I have specimens of this species compared with the Gravenhorst material in 1938.
Specimens of the aggregate species seen from England, Belgium, Norway, Sweden,
Finland, Spain, France, Switzerland, and Germany. Also reliably recorded from
Holland (Teunissen, 1948) and Czechoslovakia (Gregor, 1920).
Biology: ENGLAND, Beds., Luton, I g em. vii.1940, ex larva of Cladius difformis
Panz. (larva 20.vi.1940), V. H. Chambers. This confirms the records of Brischke and
de Gaulle. Brischke’s record of sexcinctus Grav. from the Danzig district, and of the
breeding of it from Hemichroa australis Lep. (= alni auctt. non L.), quoted by Morley
and Schmiedeknecht under Acrotomus, is very likely to refer to Cteniscus curtisii Hal.
(see p. 426 below).
Acrotomus succinctus Gravenhorst
1829 Tryphon succinctus Gravenhorst, Ichneumonologia Europaea, 2: 166-167 (non Exenterus
succinctus Holmgren, 1858, see p. 421 below).
1883 Delotomus lucidulus Thomson, Opuscula Entomologica, fasc. 9: 884 (non Gravenhorst,
1829).
1906 Acrotomus succinctus Pfankuch, Z. Hym. Dipt. 6: 94-95.
Mandible (Fig. 1).
In the Gravenhorst collection I select as lectotype the male specimen corresponding
to the description of ‘Individui prope Barterodam’ (this locality is in Hannover
province).
There is no doubt that Thomson’s description refers to succinctus Grav., though
there is also a female of the true Jucidulus in his series from PaAlsjé.
I have a pair of specimens given me by Rektor Hedwig and compared with the
Gravenhorst material in 1938.
Specimens seen from Ireland (numerous, from several counties), England, Belgium,
Sweden, Finland, France, Switzerland, Germany, Austria. Also reliably recorded
from Holland (Teunissen, 1948) and Czechoslovakia (Gregor, 1929).
Genus ANISOCTENION Forster
1868 Anisoctenion Forster, Verh. naturh. Ver. preuss. Rheinl. 25: 194.
Anisoctenion yakui Uchida
1932 Acrotomus yakui Uchida, Ins. Mats. 6 (4): 163+pl. v, fig. 12.
Female. Head moderately narrowed and rounded behind eyes; rather shallowly
emarginate behind: on vertical region and temples moderately and rather closely
punctate, sparsely so on genae except near mandibular base: upper face moderately,
rather closely and sharply punctate in middle, more sparsely so at sides: lower face
closely and rather coarsely punctate, with epistoma scarcely convex. Cheeks short
and not at all buccate. Clypeus distinctly alutaceous, rather sparsely beset with
punctures of mixed sizes, fine to coarse; sharply delimited basally ; weakly convex ;
slightly impressed just before the moderately rounded apex. Malar space very short,
less than half width of pedicellus below. Hypostomal costa weakly raised: genal
ENTOM, 2, 6 RI
322 A REVIEW AND A REVISION OF
costa scarcely raised, meeting it some way behind mandibular base, farther behind
than distance between mandibular apices.
Antennae with scape (Fig. 26) rather elongate, moderately excised externo-
ventrally ; with postannellus 1-6 times length of following segment; with flagellum
stout, filiform, 34-segmented.
Pronotum very little raised before dorsal transverse furrow which is narrow,
irregular, and shallow ; with epomiae very weak ; laterally rather finely, shallowly and
confusedly punctate above, longitudinally rugose below. Propleura finely and closely
punctate. Mesoscutum moderately and rather evenly convex ; with notaulices moder-
ately impressed, not sharp but broad and finely rugose ; moderately punctate discally,
more finely so peripherally. Scutellum very strongly convex; margined to about
middle, rather finely punctate except on elevated centre. Mesopleura closely and
rather coarsely punctate below, moderately so anteriorly above, and with an exten-
sive speculum: sternaulices broad but superficial: mesosternum almost flat, rather
finely and closely punctate in front, finely and more sparsely so behind: mesolcus
narrow and weakly impressed, closed behind: prepectus closely rugose-punctate:
epicnemial carina very weakly raised. Propodeum long, scarcely raised from in front
and not at all sharply declived ; finely punctate above, obsoletely rugose behind ; on
area pleuralis rather closely punctate, rather finely so above, moderately so below:
area posteromedia transverse but not strongly so, extending a good third the way
forward, with a percurrent median costa: area superomedia narrow, very much longer
than its greatest breadth, expanded to where it emits costulae before middle, not
sharply delimited basally.
Petiolar segment 2-6 times length of its greatest breadth ; almost regularly explan-
ate from near base to near apex; laterally weakly margined, but with spiracular
tubercles rather prominent; without distinct dorsal keels, but medially raised in
anterior half and with a median impression behind ; rather finely, closely and shallowly
‘punctate. Tergite 2 with thyridia conspicuous, very little broader than long, and ~
widely separated ; finely and closely punctate in basal half: gaster more and more
finely and sparsely punctate toward apex. Female hypopygium sharply folded mid-
ventrally. Female third valvulae rather narrow, with ventral margin slightly sinuate
and decurved, projecting strongly beyond apex of tergite 8 (as in triangulatorius
Grav., Fig. 15).
Fore wings with basal and disco-cubital veins parallel; with stigma and radial cell
unusually elongate ; with areolet sub-petiolate ; with nervus parallelus emitted about —
in middle of brachial cell. Hind wings with nervellus slightly antefurcal, intercepted
a little below middle.
Hind tibiae slender in about basal two-thirds, thereafter strongly expanded to
apex. Tarsal claws moderately and rather regularly pectinate, but not to near
apex of claw.
Length 9 mm.
Head black, with the following parts deep, bright flavous margined with fulvous:
mouth-parts except mandibular apices, clypeus, lower face except for a median mark
above, and a small mark between eye and mandibular base: clypeo-frontal suture
dark. Antennae pale ferrugineous above, merging to fulvous in apical half; bright
THE CTENISCINI OF THE OLD WORLD 323
flavous below, merging to fulvous beyond post-annellus ; with annellus and extreme
base of postannellus pale all round. Thorax and propodeum black, with mark below
fore wings and conspicuous marks on apical half of scutellum and on metascutellum
bright flavous; with hind margins of scutellum and metascutellum obscurely pale:
tegulae mainly stramineous. Fore and mid coxae, trochanters and trochanterelli pale
flavous, the coxae dark at base: the remaining segments of these legs weakly rufe-
scent: hind coxae, trochanters and trochanterelli flavous ; with coxae at base and in
part on outer side, and trochanters and trochanterelli above, blackish margined with
fulvous: hind femora, tibiae and tarsi rufous, with the following blackened—femora
weakly at base and above, tibiae more strongly above, and first four tarsal segments
except at base and apex. Gaster with tergites 1 and 2 black, pale along hind margin
and with a conspicuous pale spot near mid apex ; with thyridia pale ; with remaining
tergites rufous, but tergite 3 infuscate in about basal half.
Redescribed from the holotype 2 kindly sent me on loan by Professor Uchida:
JAPAN, Hokkaido, Sapporo, 9, 15.vi.1929, H. Yaku.
According to the original description, the male differs as follows—cheeks yellow:
scutellum amd metascutellum yellow only at apex: hind coxae and femora almost
black, the coxae yellowish at apex: gaster almost black, the tergites with pale apical
margins.
In the general colouring, shape of head, form of clypeus, short malar space, stout
flagellum, form of propodeum, petiolar segment, and female third valvulae this species
appears most closely related to triangulatorius Grav. ; but it differs in not having the
hypostomal costa sharply raised nor the tarsal claws strongly pectinate. It shares
with vidibundus Grav. and pronotalis nov. the character of having the hypopygium
sharply folded mid-ventrally. For the cheeks to be yellow in the male but only with
a small yellow spot between eye and mandibular base in the female is very peculiar
in the Cteniscini.
Anisoctenion triangulatorius Gravenhorst
1829 Tryphon triangulatorius Gravenhorst, Ichneumonologia Europaea, 2: 205-206.
1835 Tryphon mesoleptoides Stephens, Illustrations of British Entomology: Mandibulata, 7: 245,
syn. n.
1858 Acrotomus coarctatus Holmgren, Svensk Vet.-Akad. Handl. 1: 224-225.
1906 Acrotomus triangulatorius (Grav. non Hlgr., 2 non g) Pfankuch, Z. Hym. Dipt. 6: 222-223.
The following notes and description of triangulatorius Grav. and the freehand
drawings (reproduced as Figs. 13-15) were made in Breslau in August 1938:
‘I select as lectotype the specimen corresponding to Gravenhorst’s colour descrip-
tion, a female, which is placed third in the collection. The allotype is placed eighth.
Two other males in the collection differ from the above in having the propodeum
rather shining ; with strong costulae ; with area superomedia not nearly parallel-sided:
these would agree with specimens typically placed as coarctatus Hlgr.
“Lectotype and allotype:—Head slightly narrowed behind eyes, rather weakly
emarginate behind. Vertex moderately finely and densely punctate, with strong
impressions lateral to the lateral ocelli: face moderately coarsely and densely punc-
tate, not shining. Clypeus coarsely punctate, apically sparsely so; very deeply
324 A REVIEW AND A REVISION OF
delimited from face ; apically almost truncate; moderately projecting when seen in
side view. Lower mandibular tooth much longer than upper. Oral costa strongly
raised where met by genal costa. Cheeks very short.
‘Antennae with scape oval, scarcely one and a half times as long as broad ; post-
annellus curved as in laticeps (Grav.) Thoms.
‘Pronotum above and mesoscutum finely and densely punctate: pronotum coarsely
longitudinally rugose below: notaulices. moderately strongly impressed, reaching
Fics. 13-15. Figures drawn freehand in Breslau in 1938 from type of Anisoctenion triangu-
latorius Grav. Fig. 13. Propodeum. Fig. 14. Petiolar segment, with base of following. Fig. 15.
Apex of gaster, in sinistro-lateral view.
middle of mesoscutum. Scutellum moderately raised, not margined to middle. Meso-
pleura strongly punctate, with a large shining speculum: epicnemia slightly raised:
mesolcus deeply impressed. Propodeum densely punctate and rather dull: area
petiolaris finely rugose: area superomedia almost parallel-sided and almost three
times as long as broad; costulae very weak.
‘Petiolar segment more than twice as long as broad ; with prominent spiracles and
moderately strong keels ; strongly punctate and not shining except at apex. Thyridia
large, about half width of space between them (very slightly foreshortened in figure).
Third valvulae rather strongly projecting, strongly hairy.
‘Radius curved at apex.
‘Tarsal claws strongly pectinate in both sexes.’
THE CTENISCINI OF THE OLD WORLD 325
I have compared with this description long series of specimens, principally in the
British Museum and Manchester Museum collections, named as mesoleptoides Steph.
or coarctatus Hlgr., including the type of mesoleptoides and can find no satisfactory
separation.
Schmiedeknecht (1911: 2285) separated triangulatorius and coarctatus as follows:
Gaster black: segments 2 and 3 red or brown-red or blackish with red incisures.
Propodeum with 5 distinct upper areae . . ‘ . coarctatus Hlgr.
Segments 1 and 2 or 1-3 black, their hind borders and the remaining segments red.
Propodeum shining: costula absent or weak . ; . triangulatorius Grav.
Habermehl (1925) recorded different specimens under the names ériangulatorius
and coarctatus, but there is no evidence that he ever compared them, and I do not
know of any record published by Pfankuch under either of the two later names.
Schmiedeknecht (1911: 2291) merely quoted Pfankuch’s description of ¢triangula-
torius. In the L. A. Carr duplicate collection there is a specimen labelled as having
been determined by Schmiedeknecht as coarctatus, but which would run down better
in his key to triangulatorius: it agrees with my description of triangulatorius except
for the shape of the area superomedia, which has the lateral costae strongly divergent
from base to middle. Indeed, the area superomedia is seldom at all nearly parallel-
sided, at least in English specimens: yet that is one of the characters that is variable.
I have examined two female specimens from the same locality in Sweden (Skane,
HG6r district, vi.1938, D. M. S. and J. F. Perkins) in which the area superomedia is to
be described as very nearly parallel-sided: in one the costula is to be described as
very weak, and in the other it is not. I now feel confident that we are dealing with
a single species which is variable, inter alia, in the shape of the area superomedia, the
strength of the areolation of the propodeum, the prominence of the keels and spira-
cular tubercles of the petiolar segment, the convexity of the scutellum and even,
within the limits of its general form, of the shape of the clypeus.
The species is now described as follows.
Head slightly narrowed and rounded behind eyes, though sometimes more strongly
so in smaller specimens; broadly and shallowly emarginate behind: vertical region
and temples rather strongly and densely punctate, cheeks confluently so just above
mandibular base: lower and upper face strongly and densely punctate, lower face
almost confluently so just below scrobes, and upper face somewhat transversely
rugose-punctate. Clypeus strongly transverse ; coarsely punctate, sparsely so in lower
half; very deeply delimited basally; with apical margin not broadly rounded but
strongly depressed below the tentorial pits, and more or less nearly truncate in middle ;
not strongly convex, shallowly impressed before apical margin. Malar space very
short, shorter than basal breadth of front metatarsus. Where met by genal costa, the
hypostomal costa is strongly raised, often at an acute angle. Genal costa inflected:
gena almost angulate behind lower base of mandible. Mandible with lower tooth very
much longer and stronger than upper (Fig. 3).
Antennae elongate, not much attenuate to apex; with scape (Fig. 27) short,
globose, weakly excised; with postannellus distinctly curved; with flagellum 32- to
36-segmented.
326 A REVIEW AND A REVISION OF
Pronotum with epomiae sharp and strong; behind them finely to moderately and
rather closely punctate above, more or less strongly longitudinally rugose below.
Propleura shining, shallowly and finely punctate, more coarsely so anteriorly. Meso-
scutum normally convex anteriorly ; with notaulices moderately, sometimes rather
sharply, impressed ; moderately punctate discally, more finely and densely so lateral-
ly and anteriorly. Scutellum not strongly convex; seldom margined to middle;
very finely punctate at sides and almost impunctate above and behind. Mesopleura
with a shining speculum; moderately punctate anteriorly above, strongly and, in
part, almost confluently so below: sternaulices not deeply, though sometimes rather
sharply impressed: mesosternum rather finely and densely punctate; prepectus
rugose-punctate: epicnemial carina very slightly elevated: mesolcus narrow and
sharply impressed, closed behind. Propodeum more or less finely punctate above,
with area pleuralis more strongly so, rugose-punctate to finely rugose behind; with
costae weak or rather weak, the hindmost the strongest, with costulae generally com-
plete but sometimes more or less obsolete: area basalis not delimited apically: area
superomedia sometimes more or less parallel-sided, more often distinctly contracted
from middle to base: a triangular vestige of the area interna (Thomson, 1873, fasc. 5,
pl. fig. 16) is often present.
Petiolar segment elongate, 1-9 to 2:3 times length of its greatest breadth in female,
2:I to 2:7 times in male; laterally sharply margined ; with dorsal keels distinct to
beyond middle, often strong; with spiracular tubercles more or less prominent ; in
basal half indeterminately rugose-punctate, in apical half rather strongly punctate,
sometimes rugose-punctate ; apically smooth and shining in middle. Tergite 2 with
thyridia very distinct, not more than twice as broad as long. Remainder of gaster
more and more finely punctate towards apex. Female third valvulae rather strongly —
projecting, decurved (usually more distinctly than is indicated by Fig. 15).
Fore wings with basal and disco-cubital veins parallel, or even slightly convergent —
to point of ramellus; with radius upcurved at apex; with areolet usually shortly —
petiolate but sometimes with a long petiole ; with nervus parallelus emitted somewhat —
below middle of branchial cell. Hind wings with nervellus slightly antefurcal and
intercepted below middle. |
Legs rather slender, though with femora rather short (Fig. 17). Claws of all tarsi —
very strongly pectinate. ,
Length 7 to Ir mm. j
Head black or blue-black, with the following parts deep bright flavous ; occasion- —
ally paler: longitudinal marks near apices of genae, sometimes wanting, clypeus —
except marginally, and lower face with two large, triangular marks. These marks ~
extend along the orbits and in one specimen are confluent below; but normally they
reduce the black on the lower face to parts below the outer margins of the scrobes
and a medial, more or less triangular part, which is often more nearly parallel-sided —
above. Mandibles pale stramineous, occasionally brighter, merging to almost black
near apex. Palpi dull fulvous. Antennae with flagellum fulvo-testaceous, darker
above towards base ; with annellus generally entirely pale ; with scape and pedicellus
black above, below more or less obscurely pale in female and pale fiavous in male.
Thorax and propodeum black, with humeral angle of pronotum and occasionally
THE CTENISCINI OF THE OLD WORLD 327
a small mark below base of fore wing pale ; with tegulae stramineous to pale flavous ;
with large apical marks of scutellum and metascutellum bright flavous. Legs of
female with coxae and trochanters black or brownish, almost always pale at apex,
except for the hind coxae which are generally totally black: fore and mid trochan-
terelli, femora, tibiae and tarsi dull flavous, the femora in large part nearer fulvous
and the mid tarsi sometimes darkened: hind legs beyond the trochanters more or less
rufous, sometimes paler, often almost black, especially above ; hind tibiae, however,
always pale flavous at base. The male differs in having the fore and mid legs flavous,
except for the dark coxal bases, the femora scarcely tending to fulvous: the hind
trochanterelli are more or less flavous. Wings with veins and stigma ferrugineous to
deep fuscous, basally dull flavous. Petiolar segment black. Remainder of gaster
16
<2 oo
Fics. 16-17. Left hind femur, in sinistro-lateral view, of Fig. 16. Amisoctenion alacer Grav.
Fig. 17. Anisoctenion triangulatorius Grav.
varying from rufous, with the hinder tergites indefinitely infuscate in part, to black,
with only the thyridia, broad margins of tergites 2 and 3, and very narrow margins
of remaining tergites rufous, and the sternites mainly rufo-testaceous: typically
tergites 2 and 3 and the genitalia are more or less rufous, and the male is the
darker sex.
Redescribed from the following material: IRELAND, Co. W. Mayo, Old Head,
I g, 21.v.1942; Co. Wicklow, Devil’s Glen, 1 9, 23.vi.1929, Glenmore Castle, I 9,
16.vi.1934, Powerscourt Demesne, 2 33, 25.v.1932, on oak, Powerscourt Deerpark,
2 $d, 27.v.1932, on oak, A. W. Stelfox: ENGLAND, unlocalized, 1 2, 2 gg; Essex,
Colchester, 1 2, 3.vi.1909, Harwood ; Staffs., Lichfield district, 9 29,6 33, L. A. Carr;
Yorks., Grassington, I g, 19.v.1945, Beedale, 1 9, 9.vi.1945, W. D. Hincks: SWEDEN,
Skane, Hé6r district, 2 99, 2-17.vi.1938, D. M.S. & J. F. Perkins: CZECHOSLOVAKIA,
Aussig on Elbe, 1 3, 18.v.1930, Th. Kupka.
The lectotype ? and allotype of coarctatus Hlgr., ‘in Bahusia’ (SWEDEN, Bohuslan),
_C. Boheman, have been compared and found to agree with the above description and
a few of the specimens from which it was made.
Further material seen: ENGLAND, &c., 31 specimens in British Museum collection,
including the Stephens type; Surrey, Dorking, Ranmore, 1 9, 18.v.1948, G. J. K.:
BELGIvuM, unlocalized, 2 $3, C. Wesmael ; Vivier d’Oie, 2 gg, J. Tosquinet: HOLLAND,
Venlo, Daeden, Plasmolen, 3 3g, 4 99, H. G. M. Teunissen: FRANCE, Maisons Laffitte,
St. Cloud, &c., 26.iv-29.v., J. de Gaulle; Haut-Rhin, Wattwiller, A. Seyrig (alto-
gether 22 $4, 10 29 in Paris Museum) : GERMANY, material in Gravenhorst collection ;
328 A REVIEW AND A REVISION OF
Stolzenau, I 9, 13.v.1945, Hannover, I g, 14.v.1948, I 9, 15.v.1948, R. Hinz;
Schwerin, I g, 14.v.1882, I g, 17.v.1884, S. Brauns.
Biology. Herr Rolf Hinz has observed this species flying together with Tryphon
compunctor Grav. from the beginning to the middle of May, and that both species
parasitize larvae of Apythemus serotinus Mill. living on oak. He found the females
hunting for larvae that had crawled into cracks in the bark or into the foliage for
their last larval ecdysis.
Anisoctenion rubiginosus Gravenhorst
1829 Tryphon rubiginosus Gravenhorst, Ichneumonologia Europaea, 2: 145.
1858 Acrotomus insidiator Holmgren, Svensk Vet.-Akad. Handl. 1: 224 (placed in synonymy
with rubiginosus Grav. by Pfankuch, 1906).
1858 Exenterus morio Holmgren, Svensk Vet.-Akad. Handl. 1: 233 (placed in genus Cteniscus by
Schmiedeknecht, 1912), syn. n.
1883 Delotomus binotatus Thomson, Opuscula Entomologica, fasc. 9: 886 (placed as a variety of
rubiginosus Grav. by Gregor, 1929, by authorization of A. Roman).
1883 Delotomus parvulus Thomson, Opuscula Entomologica, fasc. 9: 886, syn. n.
1906 Acrotomus rubiginosus Pfankuch, Z. Hym. Dipt. 6: 90-91.
1937 Acrotomus binotatus Hellén, Notul. Ent. 17: 124 (non parvulus Hellén, 1941).
The following notes were made in Breslau in August 1938:
‘rubiginosus Grav. I select the male specimen in the Gravenhorst collection as
lectotype, as Pfankuch has already done by implication. In addition to the characters
given by Schmiedeknecht, I have noted the following: head of female broadened
behind eyes. Face with a strong longitudinal impression lateral to each scrobe:
clypeus finely transversely impressed in apical half. Epicnemia moderately strongly
raised, scarcely incised medially. Petiolar segment finely aciculate-punctate, scarcely
shining, slightly broadened near base. Radialis II sinuate. Tarsal claws weakly and
irregularly pectinate. Third valvulae finely hairy, with ventral margin sinuate,
decurved.’
I do not think that anyone will quarrel with Roman’s selection of a female lectotype
of insidiator Hlgr. ; nor with my own selection as lectotype of binotatus Thoms., from
the three female specimens thus placed in the Thomson collection, of that one with
best developed black spots on tergite 2 and most nearly sessile areolet. The situation
regarding the other two names is more involved.
In the Swedish collection in the Stockholm museum stand the following specimens,
all taken by Boheman and labelled morio Holmgren: 1 ¢ from Stockholm and I ¢
from Lapland, each labelled ‘Typus’, one ¢ from Stockholm labelled ‘Allot.’, and
1 9 from Lapland labelled ‘Parat.’ I think the male specimen from Stockholm
labelled ‘Typus’ should be considered as the holotype. I believe that the transfer of
morio Higr. from the genus Cteniscus has not previously been published ; and I dissent
from the opinion of Roman (1914: 16) regarding it.
In the Thomson collection, under parvulus Thoms., there stand two specimens.
The second is a female from Farhult in Skane, which I have determined as pumilhio
Hgr.: it is evidently one of the specimens to which Thomson referred on p. 1255. It
7"
THE CTENISCINI OF THE OLD WORLD 329
does not run to the species-group of binotatus in Thomson’s key, and I reject it from
consideration in this connexion.
The first specimen is a diminutive female, about 5 mm. in length, belonging to this
species-group but labelled ‘Scan’ (= Skane). The fourth and last specimen under
binotatus is a male from Ostergétland ; it fits the diagnosis of parvulus about equally
well, except that it is not notably smaller than the first three (female) specimens. I
prefer to leave open, at the present time, the choice between either of these two
specimens, or the possibility that the type is lost.
In all the material now examined, I can find very little divergence from the
characters noted in Breslau. In some specimens the epicnemial carina is strongly
incised medially: I should now describe it as strongly elevated between the sternau-
_lices, in front of which a blunt angulation of the prepectus suggests a lateral closing
_ of the coxal cavities. The sculpture of the petiolar segment I should now describe
as rugulose-punctate, though the rugulosities are sometimes so regularly longitudinal
as to suggest aciculation. The degree of development of the keels of the petiolar
segment is decidedly variable. The decurving of the ventral margin of the third
valvulae is less marked in smaller specimens. The diminutive female standing under
parvulus in the Thomson collection is of lighter body-colour, and has the head
appreciably narrowed behind the eyes, a condition that might be expected in an
ill-nourished individual. But I cannot find any convincing specific separation. Hav-
ing made the redescription given below, I feel satisfied that we are dealing with a
single, variable species.
Head generally a little broadened behind eyes, particularly in female, in larger
specimens strongly so; distinctly longer behind eyes than eye length seen from above;
rather shallowly emarginate behind ; strongly longitudinally impressed beside scapes ;
on vertical region and orbits finely or very finely punctate, grading to moderately or
rather finely so on genae and epistoma ; with upper face densely punctate. Clypeus
very strongly transverse; finely to moderately and somewhat sparsely punctate;
not deeply delimited basally; very weakly convex in basal half and somewhat
impressed in apical half; very broadly rounded at apex. Malar space short but not
very short, about equal to greatest width of pedicellus in side view in female, rather
less in male. Where met by genal costa, the hypostomal costa is raised into a sharp
or rather sharp tooth. Genal costa inflected: gena neither at all angulate nor much
excavate behind lower mandibular base. |
Antennae short, usually not much attenuate to apex; with scape short, weakly
excised ; with flagellum 24 to 30, usually 26- to 28-segmented.
Pronotum with dorsal transverse furrow rather strongly impressed and often trans-
_ costate at sides ; with epomiae short ; laterally finely to moderately and rather closely
punctate above, more or less longitudinally rugose below. Propleura finely punctate
and shining. Mesoscutum moderately or rather strongly convex; with notaulices
short and weakly impressed ; moderately punctate discally, finely to very finely so
laterally and anteriorly. Scutellum finely to moderately punctate in front, finely so
behind ; in larger specimens almost flat above, often margined to beyond middle,
falling almost perpendicularly behind ; in smaller specimens more ordinarily shaped,
but the same conformation suggested. Mesopleura closely and rather coarsely
ENTOM. 2, 6 ss
330 A REVIEW AND A REVISION OF
punctate, moderately so above and behind, often alutaceous in male, rugose by ante-
rior margin and below wing-base: sternaulices broad, rather shallowly impressed:
mesosternum very finely punctate: epicnemial carina extending about half-way up
mesopleuron, strongly raised between sternaulices: prepectus moderately and closely
punctate at sides, bearing traces of a pair of keels forming lateral margins to the
strongly excavate coxal cavities: mesolcus strong and deeply impressed, closed
behind. Propodeum rather short, sharply raised from behind ; on area externa finely
or rather finely punctate and on area pleuralis rather finely to moderately so, other-
wise finely rugulose and rather shining, though occasionally rugose; with costae
rather weak, the costulae very weak and usually incomplete, sometimes absent or
scarcely indicated: area posteromedia not extending to middle: area superomedia
rather elongate, generally strongly contracted from junction with costulae to base,
seldom basally delimited.
Gaster of female spindle-shaped, of male more parallel-sided. Petiolar segment
appearing rather short, of female 1-4 to 1-8 times, of male 1-5 to 2-2 times length of
its greatest breadth ; with distinct basal lobes, but with lateral margins not prominent ;
with dorsal keels weak, but often distinct for about three-quarters length of segment ;
finely or, more often, moderately punctate, usually in part rugose-punctate, very
finely punctate behind. Tergite 2 with thyridia usually short, and in female incon-
spicuous: remainder of gaster more and more finely punctate. Female third valvulae
projecting, with ventral margin decurved.
Fore wings with stigma broad ; with basal and disco-cubital veins parallel or slightly
convergent; with areolet petiolate to sub-sessile; with nervus parallelus emitted
a little below middle of brachial cell. Hind wings with nervellus antefurcal, inter-
cepted below middle.
Legs rather stout. Tarsal claws weakly pectinate, bearing a few weak comb-teeth
near base.
Length 5 to 84 mm.
Head black; with mandibles, except at apex and sometimes at base, and palpi
flavous to ochreous; with clypeus usually castaneous at apex. Antennae with scape
and pedicellus black or almost so, paler at apical margins; with annellus castaneous
or paler; with remainder of flagellum fulvous to ochreous or darker below, darker,
often mainly ferrugineous, above. Thorax black, with hind margin of mesopleura |
paler: tegulae stramineous or more white, occasionally darker. Legs mainly dull
flavous to fulvous, more the latter; with coxae and trochanters black or almost so,
pale at apex; with trochanterelli largely darkened; with fore and mid femora
darkened near base and more extensively below, and hind femora black or almost so
except at apex and extreme base; with hind tibiae more ferrugineous, especially
below, or in male even darker, but always pale at base ; with tarsi more or less dark-
ened, and fore and mid tibiae occasionally darkened near base. Wings with veins
and stigma ferrugineous or paler, basally dull flavous. Petiolar segment black, rufous —
at extreme apex. Remainder of gaster may be entirely rufous, occasionally cas-
taneous, but is generally modified in the following ways: first, it is often more
flavous beneath, and sometimes has ill-defined flavous hind margins of tergites:
secondly, it is usually to a greater or lesser extent overspread with infuscation. Most
THE CTENISCINI OF THE OLD WORLD 331
frequently this is developed on tergite 2 as a median mark or as a pair of marks (the
form described by Thomson as binotatus) ; but in darkest forms little remains pale in
dorsal view but the thyridia and hind margins of tergites. Male gaster as thus
described, but the darker forms relatively frequent.
Redescribed from the following material: ENGLAND, unlocalized, 1 3, 2 99, Desvignes
coll. (g under subnitidus Grav., see Morley, 1911: 193) ; Devon, Dartmoor, Lustleigh,
I Q, 30.vi.1934, R. C. L. Perkins; Somerset, Bristol, Long Ashton, 1 g em., I.v.1935,
ex Cladius difformis Panz. (Tenthredinidae), C. L. Walton (Kerrich, 1942: 63) ;
Kent, Eynsford, 1 9, 6.vi.1939, G. E. J. Nixon; Wye, 1 g, 10-13.v.1949, Surrey,
Byfleet, 1 3, 26-29.v.1949, Herts., Royston, 1 2, 24.v.1949, R. B. Benson ; Cambridge,
I g, v.1931, G. J. K., Gog-Magog Hills, 1 3, 19.v.1918, G. T. Lyle, Devils Dyke,
I g, 5.vi.1925, H. T. Pagden; Yorks., Anstey, 1 g, 25.v.1930, J. W. Saunt, Grassing-
ton, I 9, 19.v.1945, W. D. Hincks: BELGIvum, unlocalized, 6 ¢g, 2 92, Diest, I g, I ,
1850, C. Wesmael; Arlon, 2 $¢, 20.v.1870, J. Tosquinet: SWEDEN, Skane, Ringsj6,
2 $3, 7.vi, I 9, 12.vi, H66r district, 1 g, 13.vi, 9, 18.vi, Skaralid, 1 ¢, 3.vii.1938,
D. M.S. & J. F. Perkins; Stockholm district, 2 gg (one the lectotype of morto
Higr., the other labelled allotype), 1 9 (type of insidiator Hlgr.), C. Boheman ; Lapland
(southern), Tarna, 1 g, 18.vii, I 9, 19.vii (paratypes of morio Higr.), C. Boheman:
SWITZERLAND, Geneva, Peney, 3 ¢4, 1886, 1 3, 9 29, 18.v.-13.vi, 1876-1889, Tour-
nier; Valais, Ferpécle, 1 g, 21-27.vi, Arolla, 1 9, 30.vi, I 9, 6.vii.1935, R. B. & J. E.
Benson: GERMANY, unlocalized, 1 9, Ruthe coll.; unlocalized (probably Germany)
I g, 1 2, O. Schmiedeknecht ; Schladen, 1 9, 10.v.1946, Celle, 1 3, vi.1942, R. Hinz:
CZECHOSLOVAKIA, Karov, I 9, 9.vi.1907, coll. Holik; Trojacka, 1 3g, 6.vi.1934, Brno,
I 9, 18.v.1937, Fr. Gregor.
Further material examined: ENGLAND, Surrey, Clandon Downs, 10 ¢4, 8 929,
5-7.vi.1950, J. F. Perkins: HoLianp, Tilburg, 1 g, 4.vi.1942, A. Adriaanse (coll.
H. G. M. Teunissen under parvulus Thoms.) : SWEDEN, material in Thomson collection
(see above): FINLAND, Tvadrminne, 1 3, L. v. Essen; Impilaks, 1 g, R. Forsius;
Helsingfors, 1 g, W. Hellén: FRANCE, Maisons Laffitte, St. Cloud, la Bourboule, J. de
Gaulle ; Haut-Rhin, A. Seyrig (numerous specimens in mus. Paris): GERMANY, Hol-
stein, Trittau, 1 g, v.1946, Bayrische Wald, Waldmiinchen, 1 3, vi.1948, G. Heinrich:
_AustriA, Styria, Admont district 1,200 m., 3 29, II.vi-I.vii.1950, G. Heinrich.
Biology. Reared from Cladius difformis Panz. (Tenthredinidae) on strawberry
(Kerrich, 1942: 63).
Anisoctenion pumilio Holmgren
1858 Exenterus pumilio Holmgren, Svensk Vet.-Akad. Handl. 1: 240.
1888 Cteniscus (or Delotomus) pumilio Thomson, Opuscula Entomologica, fasc. 12: 1255.
1912 Cteniscus pumilio Schmiedeknecht, Opuscula Ichneumonologica, 5: 2339-2340.
1929 Acrotomus moravicus Gregor, Zprdvy ésk. Stat. Redl. Gymn. Nové Jicin, 1929: 4 (separate
pagination), nomen nudum, syn. 0D.
1941 Acrotomus parvulus Hellén, Notul. Ent. 20: 48 (non Thomson, 1883).
Head (Fig. 18) strongly transverse ; strongly narrowed behind eyes and rounded ;
shallowly emarginate behind: on vertical region and temples extremely finely punc-
tate, usually near occiput and on genae clearly less finely so: with lower face finely,
332 A REVIEW AND A REVISION OF
and upper face finely and closely punctate. Clypeus not strongly transverse ; sparsely
beset with punctures, rather fine to moderate ; rather deeply delimited basally ; dis-
tinctly projecting in side view, but weakly convex ; apically broadly rounded, almost
truncate ; distinctly transversely impressed before margin. Cheeks short and not at
all buccate, but in frontal view slightly concave. Malar space about two-thirds
distance between mandibular apices. Hypostomal costa moderately raised to a very
obtuse point, where it slopes strongly outward: genal costa distinctly raised, inflected.
Mandible (Fig. 2).
Antennae filiform; with scape rather elongate, moderately excised externo-
ventrally ; with pedicellus abnormally elongate (Fig. 28); with flagellum 22- to 25-
segmented.
Pronotum with dorsal transverse furrow narrow and irregular, but behind it in
middle strongly shining; with epomiae of moderate strength, not curtailed below;
behind epomiae extremely finely punctate above, rugulose or rugose below. Pro-
pleura very finely punctate, rugose anteriorly and by inner margin. Mesoscutum very
strongly convex, rising sharply from pronotum, though almost flat discally; with
notaulices moderately and rather sharply impressed; very finely punctate, closely
so anteriorly. Scutellum rather weakly to strongly convex, often buckled ; not mar-
gined to middle; falling rather sharply behind; extremely finely punctate, sparsely
so discally. Mesopleura rather finely to finely punctate below, extremely finely so
above, and with an extensive speculum: sternaulices broad and deeply impressed:
mesosternum finely and not closely punctate, its halves notably convex: mesolcus
of moderate width, deeply impressed, partly closed by a bituberculate ridge behind:
prepectus rugose-punctate: epicnemial carina slightly elevated, sometimes almost
complete. Propodeum short, curving strongly downward from base, and descending
sharply from costulae; with costae of moderate strength; very finely, and for the
most part closely punctate, only on and around area posteromedia conspicuously
shining, sometimes rugulose behind spiracle: area superomedia distinctly longer than
broad, nearly parallel-sided or a little expanded to costulae, not basally delimited:
area posteromedia not extending to middle.
Petiolar segment 1-6 to 2-0 times length of its greatest breadth; almost regularly
explanate from near base to very near apex in female; laterally distinctly but not
strongly margined ; with dorsal keels not strong, but distinct to rather near apex;
finely or very finely punctate and finely rugulose beside keels, smoother between
them, and with a median longitudinal depression between them on post-petiole.
Remainder of gaster extremely finely punctate. Tergite 2 with thyridia conspicuous,
short and broad. Female third valvulae in side view narrow, projecting strongly
beyond apex of tergite 8, with ventral margin straight or almost so (Fig. 20).
Fore wings with basal and disco-cubital veins slightly divergent; with radius
distinctly upcurved apically ; with areolet sessile or sub-sessile ; with nervus parallelus
emitted below middle of brachial cell. Hind wings with nervellus slightly antefurcal,
intercepted far below middle.
Tarsal claws not, or weakly and irregularly pectinate.
Length 5 to 64 mm. .
Head black, with mandibles, except near apex, and palpi stramineous: gena with
THE CTENISCINI OF THE OLD WORLD 333
a pale mark, stramineous or darker, at apical margin by lower articulation of man-
dible: clypeus with a broad, rather triangular mark, castaneous or paler, by the
almost truncate middle part of the apical margin. Antennae with flagellum ochreous
to rufo-ferrugineous, darker above than below: scape and pedicellus generally notably
darker, but with their apical margins pale, as also annellus and extreme base of
postannellus. Thorax and propodeum black, with the following parts paler: scutellum
and metascutellum at apex and along hind margins, fore and hind margins of meso-
pleura, marks below tegulae, and hindmost angles of propleura: humeral angles of
pronotum and tegulae stramineous: ventral margin (collar) of pronotum, more broadly
in middle and at lower angles, distinctly or obscurely stramineous. Legs mainly
fulvo-testaceous, with tarsi and hind tibiae darkened at apex, and hind knees some-
times darkened: hind tibiae otherwise paler: trochanters and trochanterelli, hind
coxae at apex, and mid and fore coxae more extensively, stramineous. Wings with
veins and stigma ochreous to ferrugineous, basally stramineous. Gaster with tergite I
black, with tergites 2 and 3 sometimes almost black in basal half, and with remainder
gradually paler towards apex, which is often almost ochreous; with apical margins
of all tergites of female and some of male, and with thyridia, yellowish ; with venter
stramineous or darker.
Redescribed from the following material: ENGLAND, NE. Yorks., Mulgrave Woods,
I 9, 3.v.1936, H. Britten ; Westmorland, Witherslack, 1 3, 24.vi.1941, A. E. Wright:
GERMANY, Bavaria, Gérkau, I 2, vi.1930, Th. Kupka; Silesia, Schillersdorf, 1 9,
1.ix.1935, I 9, 4.ix.1937, Th. Kupka; unlocalized (probably Germany), 1 9, O.
Schmiedeknecht: CZECHOSLOVAKIA, Moravia (unlocalized), 1 9; Moravia, Suchdol,
I 4, 27.viii. 1930, Fr. Gregor; Bohumin, Haatsch, 1 4, 28.viii. 1936, Th. Kupka:
SWEDEN, Skane, Hdér district, 1 g, 11.vi, Ringsjé, 1 g, 19.vi.1938, D. M. S. &
J. F. Perkins: FINLAND, Tavastia australis, Sadksm&ki, 1 9, mid-vi.1g21, W. Hellén
(Hellén, 1941, under A. parvulus Thoms.) ; Hattula, 1 2, L. v. Essen.
Additional material seen: SWEDEN, Stockholm, type 9; Skane, 1 g, C. Boheman ;
Skane, Farhult, 1 @ in Thomson coll. (Thomson, 1888: 1255) redetermined by
G. J. K.: FRANcE, Mesnil, nr. Paris, 1 2, 19.vii.1891, J. de Gaulle: GERMANY, Hann-
over, I g, 21.viii. 1946, R. Hinz: Austria, Carinthia, 1 specimen, C. Boheman.
I examined the Holmgren type in 1939, and formed the opinion that the species
should be transferred to Anisoctenion. Six days later, the Moravian pair was received
in Manchester from Professor Gregor: they agreed with the diagnosis made and with
my memory of the Holmgren type. The synonymy was confirmed by direct com-
parison in 1947.
Professor Gregor wrote to me that he had taken the species commonly in warm
localities. He found there were great individual differences between different speci-
mens and that, in smaller specimens, there was very little difference in length between
the mandibular teeth. He regarded the species as a connecting link between Acro-
tomus and Cteniscus, but had not published a valid description lest one should already
exist in an unsuspected place as, in fact, has proved to be the case.
In Schmiedeknecht’s key to Acrotomus (5: 2284-2286) this species runs down with
alacer Grav. ; indeed, the Schmiedeknecht specimen in the British Museum collection
was so determined by that author. Pumilio differs from alacer principally in having
334 A REVIEW AND A REVISION OF
the head short, strongly narrowed behind the eyes; malar space much shorter than
distance between mandibular apices; mesopleura finely punctate; sternaulices and
mesolcus strongly impressed; gaster spindle-shaped in female; ovipositor sheaths
strongly projecting (Fig. 20) ; tarsal claws scarcely pectinate ; in having none of the
features mentioned (p. 350) as special for alacer; and, of course, in the coloration
and much smaller size and number of antennal segments. It has a much greater
I9
Fics. 18-19. Head in dorsal view of Fig. 18. Anisoctenion pumilio Hlgr. Fig. 19. Antsoctenion
alacer Grav.
a PR)
Fics. 20-21. Apex of gaster, in sinistro-lateral view, of Fig. 20. Anisoctenion pumilio Hgr. '
Fig. 21. Anisoctenion laticeps Grav.
resemblance to a small specimen of rubiginosus Grav. which, however, has the head ~
slightly broadened behind the eyes and the mesopleura rather coarsely punctate ; and —
from which it can also be separated by the key characters.
Anisoctenion tenuis sp. n.
Head not so transverse as in pumilio Hlgr., but strongly narrowed and rounded ~
behind eyes; moderately emarginate behind: on vertical region and temples finely ~
and rather sparsely punctate; on genae finely or very finely and very sparsely
punctate above, rather finely and much more closely so below; on upper and lower
face moderately and closely punctate, more finely and sparsely so towards orbits.
Clypeus as described for pumilio Higr., but less sparsely punctate and not sharply
or at all deeply delimited. Cheeks and malar space as described for pumilio Hlgr.:
hypostomal and genal costae also similar, but- hypostomal weakly raised.
Antennae with scape stout, rather weakly but sharply excised externally (Fig. 29) ;
THE CTENISCINI OF THE OLD WORLD 335
with pedicellus of normal proportions; with flagellum filiform and slender, 27- to
30-segmented.
Pronotum with dorsal transverse impression broad; with epomiae of moderate
strength ; laterally sparsely and very finely punctate above, finely reticulate to rugose
below. Propleura shallowly punctate ; finely and rather closely so by outer margin,
elsewhere more coarsely and sparsely so. Mesoscutum not rising sharply from pro-
notum, but rather strongly and evenly convex, though flattened discally behind
middle; with notaulices moderately and rather sharply impressed anteriorly, quite
distinct to flattened part; finely and more or less sparsely punctate. Scutellum
weakly convex and not falling sharply behind ; sharply margined to beyond middle ;
finely and sparsely punctate discally, very finely and more closely so at sides
and especially behind. Mesopleura rather finely to moderately, rather shallowly and
sparsely punctate below, obscurely punctate anteriorly above: sternaulices broad and
rather deeply impressed: mesosternum finely and not closely punctate: mesolcus of
moderate width and moderately impressed, closed behind: prepectus finely rugose:
epicnemial carina sharply though not strongly raised. Propodeum long and not at all
sharply declived ; with costae of moderate strength in front, stronger behind ; for the
most part very finely to finely punctate and elongately griseo-pilose: area postero-
media strongly transverse, extending barely a third the way forward ; area supero-
media narrow, very much longer than its greatest breadth, basally delimited but not
sharply so, expanded to where it emits costulae before middle.
Petiolar segment very elongate and slender, 3-5 to 3-8 times length of its greatest
breadth ; laterally distinctly but not strongly margined, with spiracular tubercles
prominent ; with dorsal keels not strong, not reaching base and evanescent well before
apex; finely and sparsely punctate beside keels behind spiracles; with sparse, elon-
gate pilosity. Tergite 2 with thyridia distinctly broader than long ; finely punctate in
basal half: gaster more and more finely and sparsely punctate towards apex.
_ Fore wings with basal and disco-cubital veins nearly parallel; with areolet sub-
sessile or sessile ; with nervus parallelus emitted above middle of brachial cell. Hind
wings with nervellus almost opposite, intercepted about in middle.
Tarsal claws very weakly pectinate.
Length 6 to 7 mm.
Head black, with mouth-parts except mandibular apices, and clypeus except
marginally, pale flavous: clypeus deeper flavous in Burmese type. Antennae mostly
- flavous to base of post-annellus; beyond that dull fulvous to pale ferrugineous,
distinctly darker towards apex. Thorax and propodeum rufo-ferrugineous to black ;
_ with ventral margin of pronotum, propleura, fore and hind margins of mesopleura,
and hind margins of scutellum and metascutellum distinctly paler; with scutellum
above and propodeum behind dull rufous in Burmese type. Legs mainly fulvous ;
with fore and mid coxae, trochanters and trochanterelli whitish, the coxae fulvous
around base ; with hind legs slightly darkened in part in Formosan specimen. Wings
with veins and stigma ferrugineous; with stigma slightly and veins conspicuously
paler at base. Gaster with tergite 1 rather ferrugineous except at mid apex, and with
thyridia pale: remainder rufous in Burmese type but in Formosan specimen mainly
infuscate, though with tergites dark rufous apically.
336 A REVIEW AND A REVISION OF
Described from the following material: NE. Burma, Kambaiti, 7,000 ft., 1 3,
24.V.1934, R. Malaise (holotype): Formosa, Hassenzan, I g, 24.vi. 1934, L. Gressitt
(paratype).
Holotype in Royal Natural History Museum, Stockholm: paratype in collection
of Dr. H. K. Townes.
This species is clearly most closely related to the European pumilio Higr. The
advance in knowledge of the taxonomy and distribution of the genus gained by
describing the species is held to outweigh the disadvantage of having to do this from
the male sex alone.
W. R. M. Mason has sent me a single female specimen of a species related to pumilio
Hlgr. and ¢enuis sp. n., from a locality at a height of 3,500 ft. in the Nilgiri Hills in
southern India. This is a further wide extension of the known distribution of the
genus.
Anisoctenion ridibundus Gravenhorst
1829 Tryphon ridibundus Gravenhorst, Ichneumonologia Europaea, 2: 188-189.
1906 Acrotomus ridibundus Pfankuch, Z. Hym. Dipt. 6: 219.
1911 Acrotomus sexcinctus Morley, The Ichneumons of Great Britain, 4: 197 (non Gravenhorst
1829).
Head moderately narrowed behind eyes in female, more strongly so in male, and
strongly rounded; rather shallowly emarginate behind: for the most part finely
punctate, rather closely so above, sparsely so on most of gena; with lower and upper
face moderately punctate, and generally with lower margin of gena densely and a little
more coarsely so. Clypeus sparsely beset with punctures of mixed sizes, very fine to
rather coarse; elongately pilose near apical margin; sharply delimited basally;
strongly projecting in side view, but not strongly convex; not broadly rounded
apically in female, though often rather so in male; and not or scarcely impressed
before margin. Cheeks not at all buccate, but in frontal view slightly concave. Malar
space of female about three-quarters distance between mandibular apices; that of
male short, about equal to greatest width of fore metatarsus, seen from above near
apex. Hypostomal costa rather weakly raised: genal costa scarcely raised, inflected.
Antennae relatively slender; with scape strongly convex interno-dorsally and
weakly excised externo-ventrally ; with pedicellus almost barrel-shaped, but more
elongate than in Jaticeps Grav. (Fig. 30); with flagellum 26- to 29-, most usually
28-segmented ; having the ultimate segment usually elongate, but sometimes imper-
fectly and sometimes fully divided (thus sometimes having a segment more in one
antenna than in the other) ; and having some middle segments incurved externally
as in laticeps Grav.
Pronotum with dorsal transverse furrow shining; with epomiae sharp but not ©
strong ; behind epomiae finely punctate above, rather strongly longitudinally rugose
below. Propleura irregularly, rather finely and sparsely punctate. Mesoscutum —
strongly convex, rising rather sharply from pronotum; with notaulices weakly or
moderately, seldom at all sharply impressed ; finely punctate discally, less finely so
peripherally. Scutellum strongly convex, and falling rather sharply behind; not
margined to middle ; finely to very finely punctate. Mesopleura moderately and not
THE CTENISCINI OF THE OLD WORLD 337
closely punctate below, finely, sparsely and irregularly so above: sternaulices broad,
rather shallowly impressed: mesosternum finely to moderately and closely punctate,
prepectus rather coarsely so: mesolcus sharply impressed, widening a little posteriorly,
almost always well closed behind: epicnemial carina very slightly elevated, extending
about half way up mesopleuron. Propodeum finely punctate above and on area
pleuralis, elsewhere weakly rugose; with costae of moderate strength at sides and
behind, usually weak and often incomplete above: area basalis excavate: area supero-
media longer than broad.
Petiolar segment 1-6 to 2:0 times length of its greatest breadth; finely punctate
above, but almost impunctate by hind margin; sharply but not strongly margined
laterally ; with dorsal keels not strong, but distinct to well beyond spiracles; not
canaliculate between dorsal keels where these are distinct, but with a median depres-
sion behind. Remainder of gaster finely to very finely punctate. Tergite 2 strongly
transverse ; with a pair of broad, not sharply defined, furrows extending diagonally
outward from base; with thyridia short, sharply impressed and conspicuous, about
one-half breadth of intervening space. Female hypopygium sharply folded mid-
ventrally. Female third valvulae rather broad, closely punctate ; with ventral margin
slightly sinuate but, on the whole, slightly upcurved; projecting well beyond apex
of tergite 8.
Fore wings with basal and disco-cubital veins parallel or nearly so; with radius
slightly upcurved at apex and slightly sinuate ; with areolet sub-sessile ; with second
recurrent almost regularly arcuate; with nervus parallelus emitted a little below
middle of brachial cell. Hind wings with nervellus slightly antefurcal, intercepted
below middle.
Legs rather stout. Hind tibiae distinctly swollen before and very distinctly
narrowed to base. Claws of all tarsi strongly or rather strongly pectinate.
Length 64 to 9 mm.
Female. Head black, with the following parts flavous, with some fulvous margins:
palpi, mandibles except at apex, clypeus, genae except broadly near occiput, and
whole of lower face, except for a short median mark just below antennae. Antennae
with scape and pedicellus dull flavous to rufous below, rufous to ferrugineous above ;
with annellus more or less rufous; with remainder of flagellum fulvous to testaceous,
more or less ferrugineous near base. Thorax and propodeum black, with the following
_ parts flavous, with some fulvous margins: scutellum at apex and metascutellum,
humeral angles of pronotum and ventral margin rather broadly at sides, propleura
mainly, marks below fore wings, tegulae, and metategulae in part; usually with the
following parts more or less ferrugineous: fore and hind margins of mesopleura in part,
lateral extensions of scutellum and metascutellum, and some margins of propleura ;
sometimes with paler marks beside mesolcus. Legs mainly fulvous, with hind tibiae
decidedly paler at base ; with fore and mid coxae, and all trochanters and trochanter-
elli, mainly stramineous to pale flavous, the mid coxae often flushed with fulvous ;
with hind coxae except at apex, and hind femora except narrowly at base and apex,
dark ferrugineous. Wings with veins and stigma ferrugineous, paler at base. Petiolar
segment black, sometimes dark ferrugineous towards apex. Remainder of gaster
mainly dark ferrugineous above ; with apex and sides and with thyridia fulvous ; with
ENTOM. II. 6 Tt
338 A REVIEW AND A REVISION OF
slightly broad apical margins of tergites 2 and 3, usually with narrow margins of
remaining tergites, and with most of venter paler.
The male differs as follows: genae with flavous coloration much more restricted, '
often only narrowly flavous by the mandibular articulation ; clypeus with a-pair of
dark marks below tentorial pits; lower face with a broad, percurrent, median black
band and with eye margins black for some distance below antennal sockets, the black |
margin sometimes curved inward towards tentorial pits. Propleura not pale-coloured '
except by hind margins. Hind tarsi and tibiae dark above, the latter conspicuously _
pale at base: fore and mid femora and tibiae often paler than in female. Gaster of
darker appearance, scarcely fulvous at apex and only narrowly so at sides.
Redescribed from the following material: ENGLAND, Desvignes coll., 1 2 (previously
twice misdetermined) ; Surrey, Abinger, 1 9, 18.vili.1947, G. J. K.: BELGIum, unlocal-
ized, Groenendael, Rouge Cloitre, 9 $g,7 29, Wesmael and Tosquinet coll.: SwITZER-
LAND, Grisons, Tarasp. I 9, Ch. Ferriére: GERMANY, unlocalized (probably Germany)
I g, O. Schmiedeknecht; nr. Berlin, r 9, Ruthe coll.; Munich and other localities,
I g, 20.vi, I 3, 2 9Y, 16.vii. Buchecker coll.; Bavaria, Oberstdorf, Birgsau, 3 992,
viii.1936, G. E. J. Nixon; Silesia, Ransern, 1 g, I 2, 7.vi.1934, K. Hedwig: CzEcHo-
SLOVAKIA, Bohemia, I g, J. Sekera; Breudeis, 1 3g, coll. Salvator; Trojacka, 1 g,
6.vii.1928, Fr. Gregor.
The Silesian pair, kindly given me by Rektor K. Hedwig, was compared with the ~
male type in Breslau in September 1936.
From the key to species and description given by Morley (1911) under Acrotomus
sexcinctus Grav. I could tell that he had an Anisoctenion. In response to my request —
for the loan of one of the specimens taken by Capron, most probably around Shere
in southern Surrey, Mr. Morley most generously gave me a male. This proved, as
I had suspected, to be of this species, which has not before been properly recorded
as British. ‘
Further material seen: FRANCE, 5 specimens, Sévres, St. Cloud, &c., 12.vi.—28.vii,
J. de Gaulle: GERMANY, Gottingen, I g, I 9, 14-19.vii, I 9, 28.viii.1946, R. Hinz:
AusTrRiA, Carinthia, 19, C. Boheman. See also supplement, p. 452 below.
‘
Anisoctenion pronotalis sp. n.
Head distinctly broader than thorax, strongly transverse; slightly and almost
rectilinearly narrowed behind eyes, then strongly rounded ; moderately emarginate
behind ; in facial view strongly narrowed to mouth, with epistoma not strongly con-
vex and with cheeks not at all buccate: with temples and cheeks rather finely punc-—
tate; with lower face rather coarsely, but at sides and near midline not densely so;
with upper face broadly at sides moderately punctate: vertical region very finely
punctate and upper face in middle almost impunctate. Clypeus not strongly trans-
verse ; sparsely beset with moderate punctures; basally distinctly but not deeply
delimited ; apically moderately rounded; in side view very weakly convex in basal, —
and almost flat in apical half. Malar space a little less than distance between mandi-
bular apices. Hypostomal and genal costae sharply but not strongly raised, meeting
not far behind mandibular base, at a distance of about twice their greatest height.
Antennae (Fig. 32) with scape rather elongate, not quite twice the length of its
ay
THE CTENISCINI OF THE OLD WORLD 339
greatest breadth, moderately excised externo-ventrally ; with postannellus unusually
elongate, just twice the length of the following segment, both measured along inner
margin ; with flagellum 33- to 34-segmented, elongate, slender, and finely tapering.
Pronotum strongly raised before the sharply impressed dorsal transverse furrow ;
with epomiae strong but short; laterally very finely punctate above, very strongly
shining below ; with lateral margins at about level of epomiae suddenly expanded to
produce broad flanges, which present a distinct ventral surface overhanging the
propleura. Propleura shining and finely punctate. Mesoscutum normally convex,
rather dull; with notaulices weakly impressed but traceable to disk; rather finely
punctate, between tegulae and notaulices more coarsely and closely so. Scutellum
rather flattened ; with margins not at all sharp, yet percurrent, separating a hinder
face from the sides, and weakly tuberculate at junction ; moderately punctate above
at base, impunctate on disk, very finely punctate on sides and behind. Mesopleura
moderately punctate except on and before speculum: sternaulices rather strongly
impressed: mesosternum almost flat, shallowly, but for the most part coarsely and
very closely punctate: mesolcus narrow for more than half its length, then broadened
and deepened, largely closed by an incised costa behind: prepectus with a pair of
sharp keels which form lateral margins to the coxal cavities, and a pair of blunter
outwardly directed keels. Propodeum with a conspicuous covering of elongate,
silvery hairs ; rather shining and, for the most part, very finely rugulose-punctate, but
with the area pleuralis finely to moderately and closely punctate; with costae of
moderate strength: area basalis not distinctly delimited apically: area superomedia
elongate, greatly expanded to the costulae, which are emitted far before middle,
decidedly narrow behind: area posteromedia not extending to middle.
Petiolar segment elongate, about 2-8 times length of its greatest breadth; not
laterally margined through the spiracles, which are more or less prominent; with
sharp basal auricles ; rather strongly convex, but with dorsal keels scarcely indicated ;
smooth near mid line, moderately punctate at sides. Tergite 2 about as long as its
apical breadth ; with a pair of strong, though not sharply defined, diagonal impressed
furrows at base; with thyridia large and conspicuous. Segment 3 of female with
_ distinct epipleura margined to near apex. Female hypopygium sharply folded mid-
ventrally. Female third valvulae rather broad, with ventral margin slightly up-
curved: projecting strongly beyond apex of tergite 8.
Fore wings with stigma rather elongate and narrow; with basal and disco-cubital
veins convergent ; with radius upcurved at apex ; with areolet sub-sessile ; with second
recurrent strongly sinuate ; with nervus parallelus emitted about in middle of brachial
cell. Hind wings with nervellus slightly antefurcal, and intercepted a little below
middle.
Claws of all tarsi strongly pectinate from base to apex.
Length of female about 12} mm., of male 113 mm.
Head bluish-black, with the following parts bright flavous, narrowly margined
with fulvous: mouth-parts except mentum and mandibular apices, clypeus, genae
except near occiput, and lower face. Antennae with scape dull rufous above, with an
infuscation near apex and extending along outer edge, flavous beneath; with pedi-
cellus and annellus partly pale rufous, partly infuscate, paler beneath ; with remainder
340 A REVIEW AND A REVISION OF
of flagellum ferrugineous, paler beneath, towards apex, and at apices of basal seg- — |
ments and base of postannellus; with postannellus otherwise almost black above.
Thorax and propodeum black, but with mesoscutum bluish-black like the head, and
propleura inclining to ferrugineous ; with flavous marks below fore wings: tegulae dull
castaneous, in part dull flavous. Fore and mid legs with coxae and trochanters
stramineous, the mid coxae to some extent infuscate behind; with femora more or
less rufous, paler in front ; with tibiae and tarsi pale rufo-testaceous: hind legs with
coxae and trochanters castaneous, the trochanters and the coxae at extreme base dull —
stramineous beneath, the coxae also with paler marks above and below near base;
with tarsi and apical fourth of tibiae rufous below, darker above, the tibiae in basal
three-fourths pale beneath. Wings with veins and stigma ferrugineous, pale at base.
Gaster rufous ; with tergite 1, except at apex, almost black ; with tergite 2 dark at
base except on thyridia.
The male differs in having the hind trochanters entirely, and trochanterelli in part,
dull stramineous.
Described from the following material: NE. Burma, Kambaiti, 6,000 ft., 1 9
(paratype), I.vi.1934, R. Malaise ; Kambaiti, 7,000 ft., 1 2 (holotype), 1 g (allotype),
8.vi.1934, R. Malaise.
Holotype and allotype in Royal Natural History Museum, Stockholm: paratype in —
British Museum (Natural History).
The strongly pectinate claws and form of the female third valvulae place this
species in the laticeps species-group. The elongate scape, strongly sinuate second
recurrent vein, and general coloration relate it to laticeps Grav.; but it resembles
vidibundus Grav. in having diagonal furrows on the second tergite and a sharply
folded hypopygium. The flattened scutellum is reminiscent of laetus Grav., and the
elongate petiolar segment also of ¢tviangulatorius Grav. The form of the prepectus is —
suggested in rubiginosus Grav., but that of the pronotum is quite novel among
palaearctic Cteniscini.
From Dr. H. K. Townes, through Mr. W. R. M. Mason, I have received a specimen —
with the following data: Formosa, Musha, I 9, 21.v.1932, L. Gressitt. This differs — 4
from the above-described northern Burmese specimens as follows: hypostomal costa —
weaker, and genal costa almost absent, though a vestige is just discernible on left side of
head. Flagellum 32-segmented. Pronotum laterally moderately punctate above and
with epomiae not short. Scutellum of essentially similar form, but almost conical and
sharply bituberculate, moderately rugose-punctate at sides, and with discal impunc-
tate area greatly reduced. Punctation of area pleuralis and petiolar segment obscure.
Diagonal impressed furrows of tergite 2 not discernible. Female third valvulae not
so broad. Basal and disco-cubital veins very little convergent: areolet sub-petiolate.
Length 10 mm. Head and thorax above a deeper black: pale colour of head a deeper,
mustard-yellow: antennae almost unrelieved dull ferrugineous, apart from the cas- —
taneous annellus and extreme base of postannellus: tegulae darker: legs rather
darker, in particular the mid coxae in greater part infuscate: tergites 2 more exten-
sively and 3 at base weakly infuscate.
This form is so essentially similar to the northern Burmese specimens that I cannot
regard it as specifically distinct. The differences in size and colour are less than are
Pree i he ee
THE CTENISCINI OF THE OLD WORLD 341
known to occur in other species within Europe, and the smaller number of flagellar
segments and narrower third valvulae are clearly correlated with the smaller size.
The form of the scutellum and obsolescence of the genal costa I regard as minor
malformations: since I have seen specimens of Smicroplectrus gucundus Higr. with
the genal costa incompletely developed, I can regard this as not indicating a specific
separation. Moreover, Townes & Townes (1949: 408) record that, in Monoblastus
kaniacensis Hall, the genal costa is usually obsolete but sometimes complete. These
and other differences will indicate the range of variation to be sought, just as the
record itself is a valuable indication of the distribution.
Anisoctenion laticeps Gravenhorst
1829 Tryphon laticeps Gravenhorst, Ichneumonologia Europaea, 2: 214-215.
1883 Delotomus laticepbs Thomson, Opuscula Entomologica, fasc. 9: 884.
1906 Acrotomus laticeps Pfankuch, Z. Hym. Dipt. 6: 280.
The two specimens standing under this name in the Gravenhorst collection are not
in very good condition. They are not laticeps Thoms. et auctt., but were seen in
Stockholm in 1939 and determined by A. Roman as Cteniscus dorsator Thunb.
var. punctipleuris Thoms. The first specimen is a male; the second has the gaster
broken off. As Rektor K. Hedwig observed in a letter to me, these do not agree with
Gravenhorst’s description, and cannot be reckoned as types. Pfankuch (1906) stated
all too briefly ‘die Type ist ein Weibchen’. It is, of course, not impossible that he
made a mistake; but laticepbs Thoms. agrees well with Gravenhorst’s description.
Rektor Hedwig wrote to me that he could not find anywhere in the collection
a specimen which could be the type, which must be reckoned as lost. The interpre-
tation of Thomson, as first reviser, has, therefore, been accepted.
A redescription of the species having been made in 1938 from Thomson’s series and
other material, a lectotype was accordingly selected in 1934, a female specimen from
Palsj6. In 1948 it was found necessary to revise the description to conform with the
standard adopted in the present paper; but the lectotype, and a specimen from
Ortofta designated allotype, are among the material from which the 1948 description
has been made.
Head broadened behind eyes, generally very distinctly so; narrowly and rather
_ Strongly emarginate behind: on temples and cheeks moderately punctate, near occi-
put and malar space more densely so; with upper face very densely punctate; with
lower face rather coarsely punctate, and on epistoma very densely so. Cheeks dis-
tinctly buccate. Clypeus strongly transverse ; rather sparsely and rather coarsely to
coarsely punctate; sharply delimited basally, with tentorial pits particularly con-
Spicuous ; apically broadly rounded (Fig. 8) ; moderately projecting in side view, and
scarcely at all impressed before apical margin. Malar space about three-quarters
distance between mandibular apices. Hypostomal costa weakly raised: genal costa
scarcely raised, inflected.
Antennae (Fig. 33) with scape abnormally elongate, more than twice the length
of its greatest breadth, moderately excised externo-ventrally ; with pedicellus almost
barrel-shaped ; with post-annellus rather strongly curved externally and some middle
342 A REVIEW AND A REVISION OF
segments distinctly so; with flagellum 28- to 32-segmented, rather stout, explanate
to about two-thirds its length and thereafter tapering to apex.
Pronotum with epomiae not strong; finely punctate before and rather finely to
very finely punctate behind them ; with longitudinal rugosities below rather weak to
very weak or virtually absent. Propleura shallowly, but generally somewhat coarsely
punctate, much more finely so near outer margin; strongly rugose anteriorly and by
inner margin. Mesoscutum not rising sharply from pronotum, more or less weakly —
convex ; with notaulices short, sometimes not reaching anterior margin ; somewhat
coarsely punctate discally, more finely so laterally and anteriorly. Scutellum moder-
ately convex ; margined to about middle; finely punctate discally and very finely so
peripherally. Mesopleura shining, somewhat finely punctate above, rather coarsely —
but not closely so below: sternaulices broad, moderately impressed: mesosternum
finely and closely punctate, prepectus rather coarsely so: mesolcus strong and deeply
impressed, partly closed behind: epicnemial carina very slightly elevated, extending
about half-way up mesopleuron. Propodeum rather short; sharply raised from —
behind, and often also from in front, to base of area superomedia; with costae and —
especially costulae rather strong: rugulose-punctate above, mainly more shining on ~
area superomedia and behind, but somewhat rugose before petiole ; with area pleuralis
mainly finely punctate: area superomedia hexagonal, not greatly longer than broad: ©
area posteromedia extending to beyond middle.
Petiolar segment not notably elongate, normally 1-6 to 2-0 times length of its
greatest breadth; obscurely rugose-punctate to near apex; laterally distinctly mar-
gined, with spiracular tubercles very prominent ; with dorsal keels strong, though not
nearly attaining apex, distinctly canaliculate between them. Tergite 2 with thyridia —
somewhat indistinct, especially in paler specimens. Remainder of gaster more and
more finely punctate towards apex. Female third valvulae very broad, with ventral
margin slightly sinuate and, on the whole, slightly upcurved ; projecting well beyond
apex of tergite 8 (Fig. 21).
Fore wings with basal and disco-cubital veins parallel or slightly convergent ; with
radius slightly upcurved at apex and often slightly sinuate; with areolet shortly
petiolate ; with second recurrent. strongly sinuate; with nervus parallelus emitted
about in middle of brachial cell. Hind wings with nervellus slightly antefurcal, inter-
cepted in or slightly below middle.
Claws of all tarsi strongly pectinate.
Length 8 to 10 mm.
Head bluish-black, with the following parts deep flavous, margined with fulvous: ~
mouth-parts except mandibular apices, clypeus, genae except near occiput, lower face
except for an incomplete median darker line, margins a little way up and a median
mark on upper face. Antennae deep flavous to pale ferrugineous, darker above except
for annellus and base of postannellus. Thorax and propodeum bluish-black, with the
following parts deep flavous margined with fulvous: wide anterior marginal mark and
humeral angles of pronotum, propleura in part (though often not at all flavous in ©
male), and conspicuous marks of scutellum and metascutellum: often there are pale —
marks beneath the wings, and ferrugineous hind margins of scutellum, metascutellum —
and mesopleura: tegulae flavous. Fore and mid legs mainly more or less fulvous; on
THE CTENISCINI OF THE OLD WORLD 343
fore and sometimes mid legs the coxae, except at base, and the trochanters and tro-
chanterelli, are decidedly paler; mid coxae darker at base: hind legs rufous; with
coxae mainly, and trochanters, trochanterelli and femora above, pale ferrugineous to
almost black. Wings with veins and stigma ferrugineous, much paler at base. Gaster
black, typically with the following parts rufous: tergites 1 at extreme apex, 2 and 3
and most of 4, pygopods and most of sternal region: but the rufous coloration is
occasionally less and sometimes more extensive.
Redescribed from the following material: ENGLAND, Cambs., Wicken Fen, I 4,
3I. Vii. 1925, H. T. Pagden, 1 9, 26.viii. 1931, I 9, 26.vii. 1933, G. J. K.; Norfolk,
Catfield, 2 99, 12-16.vii.1920, J. J. F. X. King: Betcium, unlocalized, 6 33, 4 29,
Diest, 2 33, 2 29, vili.1849, C. Wesmael: GERMANY, Silesia, Breslau, 1 2, 8.vii.1937,
I 4, 19.vii.1938, K. Hedwig; Ruthe coll., r g, 1 9, Buchecker coll., 1 g, 13.vii;
unlocalized (probably Germany), 2 $3, O. Schmiedeknecht : CZECHOSLOVAKIA, Bohu-
min, Haatsch, 1 9, 24.vii.1938, Th. Kupka: SWEDEN, Skane, Palsj6, 1 9 (lectotype),
Ortofta, 1 g (allotype), C. G. Thomson.
Additional material seen: FRANCE, Arras, Chaville (near Paris), 6 specimens,
23.vi-ix, J. de Gaulle: GERMANY, Schladen, 1 g, 20.viii.1945, R. Hinz; Bavaria,
Haag-an-der-Amper, I 9, I5.viii.1947, I 3, v. 1948, G. Heinrich: Russia, Perm, I 9,
25.vii.1926, Lubischew (coll. A. Seyrig).
Time of appearance. On the data available I do not think that Heinrich (19490: 121)
was justified in postulating two generations, for I have June and a number of July
dates for this species. It is, however, possible that there are two generations that
nearly or quite overlap. .
The elongate form of the antennal scape is very characteristic, and enables this
species to be separated at once from species of Cteniscus of similar appearance, even
when the mandibles cannot be seen.
Anisoctenion laetus Gravenhorst
1829 Mesoleptus laetus Gravenhorst, Ichneumonologia Europaea, 2: 50-51.
1906 Acrotomus laetus Pfankuch, Z. Hym. Dipt. 6: 23.
Three forms have been separated in the key to species, and further distinguished
below. These are here treated as varieties of a single species /aetus Grav.; though
Thomson (1883) treated all three as separate species.
The species as a whole was described by Gravenhorst (1829) as Mesoleptus laetus.
Pfankuch (1906) refers to a label of Férster’s as Delotomus, and states that the
description of Acrotomus orbitatorius Schigdte by Holmgren corresponds with the type.
The form that is here regarded as typical and described in full agrees fairly well
with the description of Tvyphon cephalotes Gravenhorst. It was thus identified by
Thomson (1883: 885) : the present redescription is partly based on a pair of specimens
from Thomson’s series, and also a female thus named in the Ruthe collection.
Pfankuch (p. 296), in placing cephalotes in synonymy with Jaetus, stated that the
type was missing from the collection. He gave the sex as female, which it must have
been, according to the genal coloration, if it really was the Anisoctenion species.
The first reviser of cephalotes, however, was Stephens (1835, 7: 250-251), who had
344 A REVIEW AND A REVISION OF
not an Anisoctenion but a species of Cteniscus (also mixed with cephalotes in the Ruthe :
collection). A single female specimen bears Stephens’s original label: this was placed
in the British Museum collection, perhaps by C. Morley, under Cteniscus pictus Grav.,
together with several other Stephens specimens which that author, who made no
mention of pictus, may have placed as cephalotes (see Morley, 1911: 194).
I have worked through Gravenhorst’s original description to see which of the two —
interpretations, that of Stephens or that of Thomson, can be better substantiated ;
but the result has been inconclusive. It seems preferable to disturb the present
nomenclatorial usage as little as possible, pending a solution of the problems here
considered ; so the form of the species now described in full will be referred to as var.
cephalotes Grav. Thoms.
Thomson did not recognize laetus Grav. The other two forms he described as new —
species, marginatus and calcaratus. Gravenhorst’s description of laetus was evidently
made from a series of males; and it represented the variation within the species as he
saw it. The name lJaetus cannot, therefore, be used in a restricted sense for any one ©
of the three varieties until the Gravenhorst series has been re-examined and a lecto-
type recognized.
A full redescription has been made of the form marginatus Thoms., in order to
compare it most carefully with cephalotes Grav. Thoms. One of the specimens thus
studied was a female from Thomson’s series, which was both previously and subse-
quently compared with the type pair from Isgarde on Oland. Schmiedeknecht added
nothing to our knowledge of marginatus. Roman, in arranging the Swedish collection __
in the Stockholm museum, followed Thomson’s nomenclature, but made marginatus
a variety of cephalotes.
The male type of calcaratus has remained unique and unstudied since being de-
scribed by Thomson (1883: 885).
Variety cephalotes Gravenhorst, Thomson
1829 ? Tryphon cephaoltes Gravenhorst, Ichneumonologia Europaea, 2: 246-247.
1883 Delotomus cephalotes Thomson, Opuscula Entomologica, fasc. 9: 885 (non Tryphon cepha- —
lotes Stephens, 1835).
1906 Acrotomus laetus = cephalotus Pfankuch, Z. Hym. Dipt. 6: 296.
Head (Fig. 22) distinctly, generally strongly, broadened behind eyes; moderately
emarginate behind: more or less finely and closely punctate, more closely so on medial
parts of lower and upper face and on genae. Clypeus (Fig. 23) not strongly transverse ;
with moderate punctures, which are separated by about their own diameters above,
but are a little coarser and considerably sparser below; sharply delimited basally ;
with apical margin not broadly rounded, but in middle almost truncate or slightly —
emarginate (not, however, with distinct apical angles as in Acrotomus s. str.) ; very
weakly convex, weakly transversely impressed before apical margin. Malar. space
nearly half basal width of mandible, and a little shorter than distance between mandi-
bular apices. Hypostomal costa rather strongly, but very obtusely and roundedly
raised: genal costa distinctly raised, inflected.
Antennae with scape of normal proportions, moderately excised externo-ventrally: _
THE CTENISCINI OF THE OLD WORLD 345
flagellum 28- to 32-segmented, almost filiform, tapering slightly to apex ; with post-
annellus rather strongly curved externally.
Pronotum with dorsal transverse impression broad and shining; with epomiae
strong ; behind epomiae shining and moderately punctate above, strongly longitudin-
ally rugose below (seldom very incompletely so). Propleura shining and rather finely
punctate; rugose anteriorly and by inner margin. Mesoscutum strongly convex to
middle, though not very sharply raised anteriorly; more or less finely and closely
punctate; with notaulices short, distinct near but not reaching anterior margin.
Scutellum of rather flattened appearance ; with margins distinct to far beyond middle
and almost appearing to extend round apex; very finely punctate, very sparsely so
on disk, more densely near periphery. Mesopleura rather evenly and strongly punc-
tate, mesosternum rather finely so: sternaulices broad but shallow: mesolcus not deep,
narrow in anterior half then greatly broadening, partially closed by a deeply incised
ridge behind: epicnemial carina distinctly elevated, dorsally incomplete. Propodeum
above finely and somewhat densely punctate like the mesoscutum, on area pleuralis
more strongly and densely so, elsewhere finely rugose ; with costae including costulae
of moderate strength: area basalis excavate, generally not fully delimited: area
superomedia distinctly longer than broad: area petiolaris not extending to middle.
Gaster nearer spindle-shaped than clavate, but much more strongly narrowed to
apex than to base: petiolar segment rather strongly and densely punctate except
at apex; remainder of gaster much more finely so. Petiolar segment of female
1-7 to 2:3 times, of male 2-0 to 2:4 times as long as broad: laterally rather sharply
margined, with spiracular tubercles not very prominent; with dorsal keels distinct
to well beyond middle but not very strong; outside the keels rather strongly and
densely punctate, but between them more sparsely and irregularly punctate to
rugose-punctate, smooth behind. Tergite 2 with thyridia often well defined but
seldom conspicuous, not broad. Female third valvulae projecting very little beyond
apex of tergite 8; not broad, with ventral margin slightly sinuate or almost straight.
Fore wings with basal and disco-cubital veins slightly convergent; with second
abscissa of radius very slightly sinuate or almost straight ; with areolet more or less
shortly petiolate; with nervus parallelus emitted about in middle of brachial cell.
Hind wings with nervellus slightly antefurcal and intercepted below middle.
Tarsal claws strongly pectinate in female, rather strongly in male.
Length 74 to Io mm.
Female. Head black, with the following parts stramineous to flavous, margined
with ochraceous: genae largely, clypeus and lower face mainly. The pale coloration
on the genae is confluent with that on the lower face, and extends about a quarter
the way up the outer orbit, and also broadly to the lower articulation of the mandible:
dark coloration extends along clypeo-frontal suture, and on lower face to a variable
extent down mid line and from outer parts of antennal sockets, and up from the
clypeo-frontal suture towards these marks. Mandibles stramineous to flavous, merg-
ing through ochraceous to the black tips. Antennae fulvo-testaceous, darker above
and towards base ; the scape and pedicellus generally black above. Thorax and pro-
podeum black, with margins of some sclerites, and often with apical marks of scutellum
and metascutellum castaneous ; with tegulae more or less pale rufous ; with humeral
ENTOM. II. 6 uu
346 A REVIEW AND A REVISION OF
angle and usually with lower margin of pronotum pale. Fore and mid legs rather
pale rufous, becoming paler towards apex: hind legs darker rufous, merging to cas-
taneous or almost black at base: fore coxae and trochanters largely flavous below;
all trochanters paler at apex. Wings with veins and stigma ferrugineous, basally dull
flavous. Petiolar segment black, rufous usually only at extreme apex. Remainder of
gaster rufous, tending to paler below, somewhat infuscate near base.
Male. Head black, sometimes with a pair of pale marks on epistoma; with pale
marks of inner orbits extending from about level of antennae, usually beyond lowest
level of eye but never quite to mandibular base. Clypeus black at base and sides, with
rufous or paler colour by apical margin, or more extensive. Lower margin of pro-
notum often not pale. Coloration otherwise as in female.
Redescribed from the following material: SWEDEN, mus. Stockholm, 5 29, 3 3d,
provinces of Smaland, Ostergétland, Bohuslan, Stockholm, Gastrikland, and Dalarne,
det. Holmgren as orbitatorius Schigdte and det. Roman as cephalotes Grav.: GERMANY,
I 9, Ruthe coll.: SwitzERLAND, Engadine National Park, Val Munster, 2 gg, 2 29,
3.vii.1922, Ch. Ferriére; Peney 1 g, 18.v.1878, Tournier.
Variety marginatus Thomson
1883 Delotomus marginatus Thomson, Opuscula Entomologica, fasc. 9: 885.
Differs from the form cephalotes Grav. Thoms. as follows: head (Fig. 24) rather
shallowly emarginate behind ; for the most part moderately punctate. Clypeus rather
finely punctate; at apex rather broadly almost truncate (Fig. 25), the truncation
broader than in cephalotes. Antennal flagellum 27- to 31-segmented.
Pronotum irregularly rugose-punctate below. Propodeum with hinder lateral
costae of the area superomedia usually weak or absent.
———— .
Fics. 22-25. Anisoctenion laetus Grav. Head in dorsal view of Fig. 22. var. cephalotes Grav.
Thoms. Fig. 23. var. marginatus Thoms. Clypeus of Fig. 24. var. cephalotes Grav. Thoms. Fig.
25. var. marginatus Thoms. ;
THE CTENISCINI OF THE OLD WORLD 347
Petiolar segment on the average a little more elongate than that of cephalotes:
laterally usually not sharply margined; with dorsal keels usually distinct to well
beyond middle, but so weak as scarcely to interrupt the general convexity of the
segment.
Tarsal claws rather weakly pectinate in female, weakly so in male.
Length 6 to 8 mm.
Female with flavous coloration not predominating on lower face, extending only
a little way on to the genae and scarcely at all up outer orbits: male with pale margins
of the inner orbits often short.
Redescribed from the following material: SWEDEN, mus. Lund, 1 9, Gotland,
placed as marginatus Thoms. ; mus. Stockholm, 1 g, 1 2, Gotland, det. A. Roman as
cephalotes var. marginatus Thoms.: HOLLanpD, Venlo, 1 g, 28.vi.1942, H. G. M.
Teunissen, det. A. Roman as marginatus Thoms. : SWITZERLAND, Geneva, Peney, I g,
20.v.1889, Tournier: SPAIN, 4 gg, province Madrid, province Avila, Sierra Morena,
J. Abajo and A. Seyrig: ENGLAND, Cornwall, Botusfleming, 2 $3, Marshall coll. ;
Devon, Ideford, 1 g, J. F. Perkins: IRELAND, Co. Sligo, Toberscanavan, I 9, 22.vil.
1933. Co. Clare, Clooncoose, I g, 22.vii.1924, A. W. Stelfox.
Variety calcaratus Thomson
1883 Delotomus calcaratus Thomson, Opuscula Entomologica, fasc. 9: 885.
The unique male type agrees with the description of cephalotes Grav. Thoms. in
almost all respects ; but it is a large dark form with the sculpture more than ordinarily
pronounced. The length is Io mm., and the petiolar segment is just over twice as long
as broad. The tarsal claws may be described as strongly pectinate. The notaulices
do reach the anterior margin of the mesoscutum. As indicated by Thomson, the area
superomedia is not much longer than broad. The hind tibiae each bear a minute spur ;
but the right-hand spur is more than double the length of the left-hand one.
This specimen may be only a freak form of cephalotes Grav. Thoms.
Further material examined: of var. cephalotes Grav. Thoms.: BELGIUM, Rouge
Cloitre, 1 9, 1849, C. Wesmael: GERMANY, Bavaria, Haag-an-der-Amper, I g, v. 1948,
G. Heinrich (recorded as ? rubiginosus Grav., Heinrich, 19490: 122). Of var. margin
atus Thoms.: IRELAND, Co. Dublin, Slade of Saggart, 1 9, 22.vii.1936, A. W. Stelfox:
ENGLAND, Surrey, Byfleet, 1 9, 26-29.v.1949, R. B. Benson: FRANCE, Maisons
Laffitte, &c., 5 specimens, J. de Gaulle ; Alpes-Maritimes, Peira-Cava, I,500—2,000 m.,
55 specimens, 1931, A. Seyrig. Of intermediate form: BELGium, unlocalized, 2 33,
C. Wesmael: Russi, Orel, I g, 25.vi.1943, R. Hinz.
The varieties cephalotes and marginatus have been kept separate at the present
stage because it has proved possible to place most specimens into one or the other
with confidence ; but intermediate forms have now been found, as above recorded.
When really adequate material is available from the whole European range of this
generally scarce species, it seems likely that the varietal status will prove untenable.
348 A REVIEW AND A REVISION OF
Anisoctenion alacer Gravenhorst
1829 Tryphon alacey Gravenhorst, Ichneumonologia Europaea, 2: 132.
1858 Acrotomus xanthopus Holmgren, Svensk Vet.-Akad. Handl. 1: 223.
1906 Acrotomus alacer Pfankuch, Z. Hym. Dipt. 6: 86.
1907 Anisoctenion xanthopus Schmiedeknecht, Die Hymenopteren Mitteleuropas, 618.
1911 Anisoctenion alacer Schmiedeknecht, Opuscula Ichneumonologica, 5: 2295.
Head (Fig. 19) sub-cubical, having a length behind the eyes, as seen from above,
much greater than the eye length in same view ; scarcely broader than thorax ; moder-
ately narrowed and very little rounded behind eyes; very broadly and shallowly
emarginate behind, the margin often perceptibly sinuate: with vertex and temples
rather finely punctate; with cheeks moderately and upper face also more densely
punctate ; with lower face moderately, or near middle more coarsely punctate ; malar
space more or less coriaceous. Cheeks long; convex below in front view. Clypeus
(Fig. 9) elongate-pilose ; sparsely and for the most part rather coarsely punctate in
upper half, virtually impunctate in lower half except for a row of piliferous punctures
just before the margin; basally not deeply delimited; with apical margin varying
from very broadly rounded, as in the specimen figured, to rather narrowly so; in side
view rather strongly or strongly convex, and not always impressed before apical
margin. Malar space about equal to distance between mandibular apices, sometimes
rather less. Hypostomal costa weakly raised: genal costa scarcely raised, meeting
hypostomal costa far behind mandibular base.
Antennae elongate and slender; with scape strongly excised externo-ventrally ;
with flagellum 34- to 41-segmented.
Pronotum abnormally forwardly produced, narrowly rounded in front, and dis-
tinctly punctate on the fore part; with dorsal transverse furrow shallow, but broad ~
and more or less longitudinally costate in middle; with epomiae rather strong in
middle but indeterminate below ; laterally moderately punctate above, rather strongly
longitudinally rugose behind and below, more finely sculptured in male. Propleura —
rather coarsely punctate ; anteriorly coarsely reticulate-rugose ; laterally very sharply
margined. Mesoscutum rather strongly and evenly convex; with notaulices short, —
but strongly impressed and transcostate; finely to moderately and rather closely
punctate. Scutellum weakly to strongly convex; not margined to middle; finely to
moderately punctate, usually impunctate and shining in centre. Mesopleura coarsely
punctate below, moderately so above: sternaulices broad and very superficial: meso-
sternum moderately and closely punctate, prepectus rather coarsely so: mesolcus —
indicated by a transcostate line or ridge; scarcely impressed anteriorly, broadening
and merging into a wider concavity posteriorly ; sometimes appearing to be closed by
a strong transcosta, but continuing between cavities of mid coxae: epicnemial carina
scarcely elevated, extending well up mesopleura. Propodeum elongate, rather
strongly griseo-pilose ; obscurely rugose discally, more or less shining near periphery ;
finely punctate on area externa and moderately so on area pleuralis; with costae —
rather weak and indefinite above, strong behind: area basalis somewhat excavate, but —
not sharply defined: area superomedia two to three times as long as broad, weakly —
arcuately trans-rugose: area posteromedia broader than long, shining, sometimes
divided by a distinct median keel.
THE CTENISCINI OF THE OLD WORLD 349
Petiolar segment 1-5 to 1-8 times length of its greatest breadth in female, 1-9 to 2-2
times in male; almost evenly explanate from near base to apex, weakly so in male,
rather strongly so in female; laterally distinctly margined, with spiracular tubercles
more or less prominent; with dorsal keels rising very sharply at base, margining
a strongly raised medial area, approaching closely before or between spiracles, but
becoming indistinct not far behind middle of segment; rather finely punctate, or in
part finely rugose, beside the keels, finely punctate between and behind them. Ter-
gite 2 with thyridia conspicuous, in male large: thyridia of tergite 3 distinct. Re-
mainder of gaster very finely punctate. Female hypopygium large. Female third
valvulae rather broad but not large; not or scarcely reaching apex of tergite 8 ; with
ventral margin scarcely curved.
Fore wings with basal and disco-cubital veins decidedly divergent ; with areolet
_ sessile or very shortly petiolate; with nervus parallelus emitted below middle of
brachial cell. Hind wings with nervellus rather strongly antefurcal, intercepted far
below middle.
Legs elongate and slender. Claws of all tarsi rather strongly to strongly pectinate.
Length of female 8 to 10, of male 8} to 11 mm.
Head black, with the following parts stramineous to flavous, margined by fulvous
(the colour often appearing to have been dulled by cyanide vapour): clypeus, except
_ for a broad basal margin, and mandibles except at apex. Palpi more or less rufo-
_ testaceous. Antennae ferrugineous or darker, darker above; usually with a series of
_ basal segments at apical margin, and annellus mainly, distinctly paler. Thorax and
_ propodeum black; with humeral angles of pronotum, tegulae, and metategulae
_ stramineous to flavous, the pronotal marks usually margined with fulvous. Legs
_ mainly rufous to rufo-testaceous ; with fore and mid tarsi at apex, hind tarsi largely,
and hind tibiae sometimes at apex, darker ; with coxae often darker in part ; with fore
coxae and fore and mid trochanters in front often flavous. Wings with veins ferru-
gineous to testaceous, much paler at base ; with stigma pale testaceous. Gaster black ;
with thyridia castaneous ; with tergites 2 to 7 usually finely pale-margined: in male
to some extent pale beneath, especially on fore part ; in female with hypopygium and
region behind it rufous to ferrugineous, and with preceding sterna stramineous to
_ ochreous or citreous.
Redescribed from the following material: ENGLAND, Stephens coll., 1 g, 2 99;
Devon, Newton Abbot, 3 $3, 1 2, R. C. L. Perkins and J. F. Perkins; Lancs., Man-
chester, Mersey banks, 1 9, L. Nathan, 6 33, 2 29, G. J. K.: Betcrum, Sysseele,
Groenendael, Stockal, Cortenaeken, Charleroy, 12 3g, 2 99, Wesmael and Tosquinet
coll.: GERMANY, ? Bavaria, 3 $3, 2 99, Buchecker coll.; Silesia, Mihlbach, 1 2,
K. Hedwig.
Further material seen: ENGLAND and GERMANY, 25 specimens in British Museum
collection: SWEDEN, material in Holmgren collection: FRANCE, Haute Marne, Clerc ;
Rambouillet, J. de Gaulle (altogether 4 specimens): AUSTRIA, holotype. Recorded
from Holland (Teunissen, 1948) and Czechoslovakia (Gregor, 1929). The Woldstedt
specimen recorded from Finland is a female Cteniscus similis Hlgr.
All the dates, where recorded, are in the months of September and October, usually
in the first three weeks of October.
350 A REVIEW AND A REVISION OF
The Silesian male, kindly given me by Rektor K. Hedwig in 1936, was compared
with the type in the Gravenhorst collection. The synonymy of xanthopus Hlgr. was
confirmed in Stockholm in 1937.
Morley (1911) quoted the description of Tryphon anceps Stephens (1835), which he
considered to be ‘almost indubitably synonymous’. Stephens’s specimen is not to
be found in the British Museum collection; but from the description and date of
capture, I consider such synonymy to be not at all probable.
This is a rather isolated species within the genus. The elongate, greyish-black body
caused some earlier workers to confuse it with Dyspetes praerogator Grav., which it
superficially resembles. Peculiar features are the forwardly produced pronotum,
the ill-defined, transcostate mesolcus, and the elongate legs, including particularly
slender hind femora (Fig. 16). Notable also are the transcostate notaulices, the
elongate areola, and the sharply raised medial area of the petiolar segment. The sub-
cubical head and colour pattern of the clypeus it has in common with an apparently
new species, of which two males are available, and which is here briefly diagnosed but —
not validated. But that species does not share with it any of the other features men- —
tioned in the present paragraph.
Biology. Herr Rolf Hinz has taken this species in the Hannover district in October,
flying between grass in the woods. He finds that it parasitizes species of Tenthredopsis.
The egg is laid externally not far behind the head of the host larva. This makes no
defensive movements, and the parasite may ride it for a considerable time.
Clausen (1932) studied the life-history of a species determined as A. alacer Grav.
It was parasitizing a species of ‘ Allantus’ on oak at Suigenin Korea. R. B. Benson tells
me this would probably be Allantus (= Emphytus auctt.) togatus Panz. or some
related species.
Anisoctenion genalis Thomson
This species may be diagnosed by the sum of the characters leading to it in the key
to species (pp. 350-354).
Diagnosed from the following material: ENGLAND, Devon, Dartmoor, I 3, 8.vi.1935,
R. C. L. Perkins: Austria, Styria, Bésenstein Massif, 1,800 m., I 3g, 8.vi.1950,
G. Heinrich: see also supplement (p. 454 below).
This species forms a connecting link with the genus Cteniscus. When further
material was seen in 1952, including both sexes, I was able to determine it. A
description is given in the supplement (p. 452 below).
KEY TO PALAEARCTIC SPECIES OF ANISOCTENION FORSTER
1. Female third valvulae more or less distinctly decurved (Fig. 15): mesopleura more
or less coarsely and closely punctate and either epicnemial carina strongly
raised or malar space very short: hypostomal costa raised into a rather sharp
tooth in known European species. . 2
Female third valvulae not thus distinctly decurved: neither epicnemial carina
strongly raised nor female malar space very short: if the male malar space is
very short the mesopleura are rather finely and sparsely punctate: hypostomal :
costa, if rather strongly, then very obtusely and roundedly raised ; 4
THE CTENISCINI OF THE OLD WORLD 351
. Clypeus very strongly transverse, not coarsely punctate nor deeply or sharply
delimited: malar space not very short: flagellum 24- to 30-segmented: epic-
nemial carina strongly raised between sternaulices: hypostomal ‘costa raised
into a rather sharp tooth where met by genal costa: larger specimens are
rather stoutly built : ‘ . rubiginosus Grav.
Clypeus less transverse, coarsely punctate or at least with some coarse punctures,
and deeply or at least sharply delimited: malar space very short in both sexes:
flagellum 32- to 36-segmented: epicnemial carina very slightly raised: species
of rather slender build . ; : ‘ : ‘ ; , ; 3
. Hypostomal costa raised into a rather sharp tooth where met by genal costa (as
in rubiginosus Grav.): female hypopygium not sharply folded mid-ventrally:
claws of all tarsi very strongly pectinate: antennal scape shorter (Fig. 27):
Europe. ; : . triangulatorius Grav.
Hypostomal costa weakly raised, not at all ‘dentate’: female hypopygium
sharply folded mid-ventrally: tarsal claws no more than moderately pectinate:
antennal scape longer (Fig. 26): Japan, unknown in Europe yakwi Uchida
. Head not broadened or sub-cubical, not longer behind eyes than eye length, seen
from above: either mesopleura rather finely and sparsely punctate or tergite 2
and female hypopygium as in second half of couplet 5 : 5
Head broadened or sub-cubical, distinctly longer behind eyes than eye length,
seen from above: mesopleura strongly punctate in lower half: tergite 2 and
female hypopygium not so ; : ; , y : - kO
. Mesopleura finely or rather finely punctate: sternaulices strongly impressed:
tergite 2 and female hypopygium not as in alternate . : ; ‘ 6
Mesopleura moderately or strongly punctate below: sternaulices no more than
moderately impressed: tergite 2 with a pair of diagonal furrows extending
outward from base: female hypopygium sharply folded mid-ventrally . 9
. Head strongly narrowed behind eyes (Fig. 18): tarsal claws with pectination
absent or weak: female third valvulae not as in alternate . : 7
Head no more than moderately narrowed behind eyes: tarsal claws moderately
and very distinctly pectinate: female third valvulae moderately broad in side
view and very broad as seen from above, almost ‘vomeriform’, shortly and
densely hairy below: NE. Burma, Formosa ; , : : 2: ep
. Propodeum strongly raised to base of area superomedia, which is little longer
than broad: petiolar segment not 1} times as long as broad :
cf. Cteniscus rufonotatus Hier.
Propodeum not thus strongly raised, with area superomedia distinctly longer
than broad: petiolar ee more than 14 times length of its greatest
breadth : : ‘ i ; R ‘ : : 8
. Pedicellus abnormally elongate Fig. ss narrower below than width of malar
space: flagellum 22- to 25-segmented: scutellum not margined to middle:
propodeum short, with area posteromedia roughly quadrate: petiolar segment
up to twice length of its greatest breadth: Europe. . pumilio Higr.
352 A REVIEW AND A REVISION OF
Pedicellus short and stout (Fig. 29), broader below than width of (male) malar space:
flagellum 28- to 31-segmented: scutellum sharply margined to beyond middle:
propodeum long and not at all sharply declived, with area posteromedia
strongly transverse: petiolar segment more than three times length of its
greatest breadth: female unknown: NE. Burma, Formosa . tenuis sp. n.
~ . 26
al
28
tenuis a ie A a 9
ridib.
30
31
potty?
Fics. 26-31. Antennal scape and following segments, in sinistro-lateral view, of Fig. 26. Aniso-
ctenion vakui Uchida. Fig. 27. Anisoctenion triangulatorius Grav. Fig. 28. Anisoctenion pumilio
Higr. Fig. 29. Anisoctenion tenuis sp.n. Fig. 30. Anisoctenion ridibundus Grav. Fig. 31. Aniso-
ctenion laetus Grav. var. marginatus Thoms. ;
g. Pronotum and prepectus not with abnormal developments as in alternate: post-
annellus about one and a third times length of the following antennal segment,
measured along inner margin (Fig. 30): petiolar segment not more than about
twice the length of its greatest breadth ; with dorsal keels distinct though short:
tergite 3 almost entirely black or blackish: length 9 mm. or less: Europe
‘ , : : . ; ; , : ‘ ridibundus Grav.
Pronotum with lateral margins suddenly expanded to produce broad flanges:
; THE CTENISCINI OF THE OLD WORLD 353
= prepectus with a pair of sharp keels which close the fore coxal cavities later-
ally: postannellus twice the length of the following antennal segment, similarly
measured : petiolar segment 2:8 times length of its greatest breadth ; with dorsal
keels scarcely indicated: tergite 3 almost entirely red: length 10 mm. or more:
NE. Burma, Formosa. : . - : ‘ . pronotalis sp. n.
10. Head more or less broadened and strongly rounded behind eyes (Figs. 22, 23):
clypeus not coloured as in alternate . ; 3 aed
Head sub-cubical, slightly and almost rectilinearly narrowed (Fig. 10): clypeus
stramineous to flavous, margined with fulvous; and with a broad, blackish
band along basal margin. ; ; ‘ : : : ; - ae
Se. sty i eae cr oe.
Fa a, OO ae
_ Fics. 32-33. Antennal scape and following segments, seen from above, of Fig. 32. Anisoctenion
_ pronotalis sp.n., illustrating especially the elongate postannellus. Fig. 33. Anisoctenion laticeps
Grav., illustrating especially the elongate scape.
_ It. Scape very elongate (Fig. 32): clypeus as in Fig. 8: female third valvulae very
broad, extending well beyond apex of tergite 8 PB. 21): male genae, like those
of female, largely flavous : , . laticeps Grav.
Scape not unusually elongate (Fig. 31): clypeus as in Figs. 24, 25: female third
valvulae not broad, scarcely projecting beyond apex of tergite 8; pale colora-
tion scarcely touching genae of male (/aetus Grav. agg.) : Ee
_ 12. Head rather shallowly emarginate behind (Fig. 23): clypeus at apex rather
| broadly almost truncate (Fig. 25): malar space much shorter than distance
between mandibular apices: tarsal claws more weakly pectinate in each sex
respectively: female with flavous coloration not predominating on lower face,
extending only a little way on to the genae and scarcely at all up the outer
orbits . . . laetus Grav. var. marginatus Thoms.
Head moderately émateinate behind (Fig. 22): clypeus with apical margin in
middle almost truncate (Fig. 24), more narrowly so than in var. marginatus : malar
space not much shorter than distance between mandibular apices: tarsal claws
more strongly pectinate in each sex respectively: female with flavous coloration
predominating on lower face, extending broadly on to the genae and about
a quarter the way up the outer orbits : ; , ‘ RS
13. Area superomedia distinctly longer than broad: hind tibiae with no spur .
; , laetus Grav. var. cephalotes Grav. Thoms.
ENTOM. II. 6 xx
354 A REVIEW AND A REVISION OF
Area superomedia not much longer than broad: hind tibiae each bearing a minute
spur: unique male . : : ; laetus Grav. var. calcaratus Thoms.
14. Pronotum strongly produced forward: mesolcus not distinctly impressed, indi-
cated by a transcostate line or ridge: legs, particularly hind femora (Fig. 16),
elongate and slender: antennal flagellum 34- to 41-segmented: a rather large
species, length 8 to 11 mm., with male of elongate and slender build, and with
gaster mainly greyish- black ; ; : ; alacer Grav.
Pronotum not strongly produced forward: mesolcus distinctly, though not deeply
impressed: legs, particularly hind femora, not nearly so elongate and slender:
antennal flagellum less than 34-segmented : , ; a ae ee
15. Pronotum laterally above and mesoscutum rather coarsely and closely punctate ;
gaster shallowly, but for the most part moderately and closely punctate:
gaster with tergites 2 to 4red . ; ‘ ‘ . genalis Thoms.
Pronotum laterally above and mesoscutum finely punctate; gaster for the
most part extremely finely punctate: gaster blackish or deep castaneous, with
tergites finely pale-margined at apex: male unknown to me ‘ °
cf. Cteniscus similis Hier.
Genus EXENTERUS Hartig
1837 Exenterus Hartig, Arch. Naturgesch. 3 (1): 156.
Tricamptus Forster 1868, Thomson 1883 non Woldstedt 1874, syn. n.
Picroscopus Forster 1868, syn. n.
Picroscopus Thomson 1883 non Forster.
Picroscopus Davis 1897, synonym by Roman 1913.
Like the majority of Forster’s genera, Picroscopus was founded without the citation
of any species. It was adopted by Thomson for the single species Tryphon ictericus —
Grav., and has been understood thus by later authors.
Forster’s diagnosis of Picroscopus was extracted from his key to genera and quoted —
by Schmiedeknecht as follows: ‘Fliigel mit Areola; Hintertibien ohne Sporn; FuB-
klauen gekammt; Hinterleib breit sitzend, das 1. Segment gleich hinter der Basis
stark erweitert und von da nach der Spitze hin fast unmerklich erweitert; Area
superomedia nicht scharf umgrenzt, breiter als lang.’ It has been overlooked that
ictericus Grav. has non-pectinate claws: it disagrees, therefore, with the original
generic description and, according to the interpretation of the rules generally accepted
in Europe, it cannot serve as type. |
Picroscopus Davis 1897 has already been synonymized with Exenterus by Roman
(1913). Davis included two species, Exenterus canadensis Provancher 1883 and Exen-—
terus hullensis Provancher 1886. In his key to genera (p. 227) Davis placed Picro-—
scopus in a section with tarsal claws pectinate ; but Cushman (1940), in his review
of Exenterus, placed hullensis Prov. in a section with tarsal claws not pectinate in
female. If the claws are simple in the male also, hullensis Prov., like ictericus Grav., ©
could not serve as type.
THE CTENISCINI OF THE OLD WORLD 355
Cushman accorded canadensis Prov. fuller treatment, but the identity of that
species has been in doubt. In case any attempt should be made to upset Cushman’s
opinion, I propose to clinch the matter by proposing an alternative type.
Schmiedeknecht continues: ‘Aus Forsters hinterlassenen Schriften habe ich nun
ersehen, daB er als Typus dieser neuen Gattung den Exenterus oriolus betrachtet
hat ; darauf wiirde wohl Niemand kommen, denn bei allen Exemplaren dieser Art,
die ich untersucht habe, ist der Metathorax deutlich gefeldert, speciell die Area
superomedia gut umgrenzt [italics mine], weit besser wiirden sich E. cingulatorius und
ictericus als Typen fiir die Gattung eignen. Abgesehen von der Felderung des Meta-
thorax paBt E. oriolus ganz gut auf die Diagnose.’
Now the nature of the bordering of the area superomedia is not easily amenable to
exact definition, for it is not comparable to that occurring in, for example, Coelich-
neumon, but consists of rugosities of inconstant shape. Having before me three speci-.
mens of oviolus, I found that in one specimen the rugosities that border the area
superomedia are raised distinctly above the others, in another they are not. While
one may agree with Schmiedeknecht that the area superomedia is well bordered, I do
not think it can be asserted that it is always sharply bordered. Since the evidence
is that for Picroscopus Forster meant oriolus Hart. I think we should give that author
the benefit of the doubt. Accordingly, as a precautionary measure, I select Exenterus
oriolus Hartig 1838 as type of the genus Picroscopus, which falls into synonymy with
Exenterus.
I intend to treat ictericus Grav. as a species of Exenterus. Apart from the coloration,
it is said to differ from other known European species in having the petiolar segment
angularly projecting at the base and not auriculate, and in the absence of a costula.
A noteworthy character is that tergite 2 bears coarse but well-separated punctures,
and is not or scarcely at all rugose. However, in hullensis Prov. and in smaller speci-
mens of adsperus Hartig and other species, tergite 2 bears well-separated punctures
over a large part of the area, and is rugose only at base and in middle. Cushman
(1940) separated off a group of four North American species, including canadensis
Prov., in which the basal carina is absent. In the shape of the petiolar segment I
recognize no generic separation which appears, therefore, to break down. I am
much indebted to Dr. H. K. Townes for sending me specimens of canadensis Prov.
and hullensis Prov. and thus enabling me to check the points at issue. This new
synonymy has been published by Mason (1951, April: 229)
The position of ‘Tricamptus’ apiarius Grav. has been considered in the discussion
of genera (p. 316).
The genus Exenterus has recently been revised by Cushman (1940). His paper is,
of course, a very notable advance on the work of Schmiedeknecht, who began his
key to species with a consideration of the coloration of the fore-legs ; but even Cush-
man has not made adequate use of the characters afforded by the anterior part of
the mesosternum. Moreover, an examination of the main European museum collec-
tions has yielded further results previously unpublished. The key to species that
follows is based on Cushman’s work, but incorporates such additions and modifica-
tions as I have found desirable. The determination of single males, however, remains
often a matter of considerable difficulty or uncertainty.
356 A REVIEW AND A REVISION OF
Exenterus apiarius Gravenhorst
Most of the salient characters have been mentioned by other authors, but the
following should be added: epicnemial carina distinctly, rather weakly or moderately
raised: antennal scape elongate, deeply excised ventrally, its dorsal about, usually just
over, twice its ventral length. The hypostomal costa is strongly raised and falls
gradually to occipital foramen; and the double genal costa is similar to that of —
Exyston genalis Thoms. (Fig. 51). In well-developed specimens the scutellum has, in
about apical half, a saucer-like depression, with a distinct margin in front and at sides.
Recorded in each month from May to September; the earliest day of a specimen
seen by me being 17.v.1854 (male, Ruthe coll.), and the latest 10.ix (Rebais, nr.
Fontainebleau, de Gaulle).
Specimens seen from Wales, England, Holland, Belgium, France, Germany,
Sweden, Poland (Przepatkowo, viii.1925, G. Heinrich, mus. Paris), Russia (Perm, &,
Lubischew, mus. Paris).
Exenterus abruptorius Thunberg
The hind margin of the cavities of the fore coxae is raised, but not bidentate:
therefrom radiate strong keels to the epicnemial carina, weaker in male. Tergite 2 of
gaster is generally slightly less than twice as broad as long. Antennal scape rather
weakly excised externo-ventrally. For other characters, see comparison under oviolus
Hart., and also key to species. See also Pl. 4.
' Biology. This has been studied by Clausen (1932, under coreensis Uchida) and by
Morris (1937). The species has been reared from Diprion pint L. and Neodtprion
sertifer Geoftr.
The dates of emergence of reared specimens are obviously affected by laboratory
conditions: in the field the adults are known to appear in the last ten days of June
and in early July.
Specimens seen from England (see Kerrich, 1935), Germany (mus. Munich),
Sweden (Holmgren and Thomson collections; Morris, Cameron, & Jepson), Finland
(V. Saarinen; G. J. K.), France (Braunschweig and Alpes Maritimes, mus. Paris),
Spain (Valencia, Cafiada, Giner Mari).
Reliably recorded also from Czechoslovakia (Gregor, 1929). Doubtfully recorded
from Belgium (Tosquinet, 1890). Not yet found in Holland. Introduced to Canada,
but apparently failed to become established.
Exenterus simplex Thomson
Thomson (1883) made the statement ‘mesosterno et -unguiculis simplicibus’.
Roman (1912, 1913) did not see Thomson’s types, but found in Thunberg’s collection —
specimens which appeared to agree with Thomson’s description, and he regarded
simplex as a dwarf form of abruptorius Thunb. Thomson’s mention of locality leaves
no doubt that the type is correctly identified. Examination of this type under the
microscope at a favourable angle and with good illumination shows that the hind —
—
THE CTENISCINI OF THE OLD WORLD 357
_ margin of the cavities of the fore coxae is, in fact, very strongly raised and strongly
bidentate. The keels towards epicnemial carina are very weak and only basally
developed.
Another specimen stands in the Thomson collection and has for locality only
‘Suecia’. Dr. Kjell Ander informs me that the two small coloured labels indicate that
it was collected by J. W. Zetterstedt in Gotland in July-August 1841.
Exenterus oriolus Hartig
1838 Exenterus oriolus Hartig, Jahresber. Fortschr. Forstw. 1: 270.
1883 Exenterus flavellus Thomson, Opuscula Entomologica, fasc. 9: 887.
This synonymy is correct. Thomson’s unique type was taken by Dahlbom on a pine
needle beside a larva of Diprion similis Hart. ; and the published locality is Kjeflinge,
which is north-west of Lund.
Cushman (1940) could find no structural characters to separate this species from
abruptorius Thunb., and wrote that it might be a colour form of it. The proportions
of the head and malar space (see key to spp., pp. 364-365) and the structure of the meso-
sternum appear to me to render it distinct. The margin of the cavities of the fore
coxae is not raised behind, but only weakly at sides, and the keels towards epicnemial
carina are absent or very few. In female, the yellow colour extends about three-
quarters way up eye behind it, as against about a quarter the way up in abruptorius.
In male, the yellow bands of the first two tergites are much broader than those of
following: not so in abruptorius.
E. oriolus appears to be a slightly shorter and stouter species than abruptorius: the
' hind legs are noticeably a little shorter and stouter relatively, and tergite 2 is slightly
more than twice as broad as long in a few female specimens measured. The scape is
a little more deeply excised in oriolus (but not sufficiently so for diagnosis) ; and the
average number of flagellar segments is smaller, ranging from 25 to 29 as compared
with 27 to 32 in abruptorius. The tarsal claws are less strongly pectinate.
Finally, the suggestion made to me that this is a southern colour-form of abrup-
torius is now further contra-indicated by the distributional data; for I know abrup-
torius from near the Arctic Circle down to the Spanish province of Valencia. The
female specimen taken in that province is rather more extensively and brightly
yellow-coloured than those from more aay latitudes, but does not approach
female orviolus in colour pattern.
Biology. Reared from Diprion pint L. and Neodiprion sertifer Geofir. ; and it seems
likely that D. similis Hart. is also involved: Schmiedeknecht (1911) mentions also
Gilpinia variegata Hart. and frutetorwm Fabr.
Material seen from Sweden (Thomson coll.), Germany (Ruthe coll., Th. Kupka
coll.), Holland (H. G. M. Teunissen coll.: Teunissen, 1948), Spain (Teruel, Orihuela
del Tremedal, ex D. pini L.). Also recorded from France (Maneval, 1925) and
Hungary (Cushman, 1940). Material obtained by Farnham House Laboratory
staff, perhaps from Hungary, also seen.
Dates: Maneval’s specimen was captured on 28.vii.19g24; and the recorded emer-
gences are on dates later in the year.
358 A REVIEW AND A REVISION OF
Exenterus ictericus Gravenhorst
1829 Tryphon ictericus Gravenhorst, Ichneumonologia Europaea, 2: 208.
1858 Exenterus ictericus Holmgren, Svensk Vet.-Akad. Handl. 1: 235.
1883 Picroscopus ictericus Thomson, Opuscula Entomologica, fasc. 9: 888.
The cheeks and malar space are short. The margin of cavities of fore coxae is
strongly raised, and very strongly emarginate medially, as in amictorius Panz.:
laterally it extends strongly to the fore margin behind lower angle of pronotum:
ventrally there often extend from the margin to the epicnemial carina a series of weak
keels producing the appearance of a row of large quadrangular punctures. The
mesolcus is closed behind. The clypeus sometimes bears a pale spot: the pale anterior
marks of mesoscutum are frequently missing.
Schmiedeknecht stated that the male was unknown: in fact there is very little
sexual dimorphism, but in the male the dorsal keels of the petiolar segment are
distinct almost to apex.
In one male specimen from the Swiss Jura the costulae are present, though weak
and incomplete.
Biology. Bred from the Diprionid sawfly, Monoctenus junipert L. (Forsius, 1911).
Material examined as follows: FINLAND, Kuusamo, Paanajarvi, Ruskeakallio, 1 9,
14.vili.1935, G. J. K. (compared with type viii.1938): SWEDEN, Jamtland, 3 gd,
mus. Stockholm ; Holmgren coll.; Thomson coll.: BELGIum, Béverloo, 3 gg, I 9,
26.vi.1867, J. Tosquinet: GERMANY?, I g, 2 29, Buchecker coll.: SwITZERLAND,
Geneva, Peney, 2 3d, 3 29, 10.vi-30.vii, 1885-1889, Tournier; Engadin National
Park, 4 36, 24.vii-2.viii ; Swiss Jura, le Crit, long series, vii.1935, A. Seyrig: FRANCE,
I 2, holotype; Alpes-Maritimes, long series, 1931, A. Seyrig: SPAIN, I 4, viii.1904,
J. Dusmet: ITALy, Piedmont, Biella, Val Chiobbia, 2 specimens, viii.1928, F. Capra:
YUGOSLAVIA, Prenj Mts., I 3, 15.vili.1936, V. & E. Martino.
Recorded from Germany (Thuringia: Schmiedeknecht, 1911) ; also from Czecho-
slovakia (Ruthenia, UbuSin, vi.1928: Gregor, 1929) and from Bulgaria (Rila Planina,
15-16.vii.1931: Gregor, 1933).
Dates of appearance of this species thus range from the end of June to early
September.
Exenterus amictorius Panzer
This species has been generally known as Exenterus marginatorius Fabr. For the
rejection of Ichneumon marginatorius Fabricius 1793 as a homonym of Ichneumon
marginatorius Rossi 1790, and the substitution of the next available name, I follow
Cushman (1943). He has reproduced Panzer’s original description and figure, and
I agree that these must have been intended to represent the species in question. The
hind tibial spurs may have been added as an afterthought, their absence not having
been noted at that time.
a i
~ i weytinnys
Hind margin of cavities of fore coxae strongly raised and bidentate: keels therefrom —
to epicnemial carina absent or very weak. Antennal scape very weakly excised
externo-ventrally (Fig. 34). The extent of yellow coloration on the gaster is very
variable. The tarsal claws are stated by Schmiedeknecht to be simple: they are,
prewes
THE CTENISCINI OF THE OLD WORLD 359
however, pectinate, at least in the female, though the pectination is weaker and less
regular than in female abruptorius. Petiolar segment (Fig. 39).
Biology. Recorded as reared from Diprion pini L., D. similis Hart., Neodiprion
sertifer Geofir., Gilpinia pallida Klug, virens Klug and polytoma Hart. Oviposition on
D. pini L. described by Scheidter (1934).
Dates of captured specimens range from early June to the end of September.
Material seen from England (see Kerrich, 1935), Belgium (Wesmael-Tosquinet
coll.), Germany (Hartig coll., Ruthe coll.), Austria (G. Heinrich), Czechoslovakia,
Poland, Sweden, Finland, Switzerland, France, Spain. Also recorded from Holland
(Teunissen, 1948). Introduced to Canada, where it has become established.
Exenterus claripennis Thomson
1883 Exenterus claripennis Thomson, Opuscula Entomologica, fasc. 9: 887 (non Cushman, 1940).
The Thomson specimen under this name in the Lund collection is labelled ‘ Nerike’
and is referable to amictorius Panz.: the type from Wittsi6 in northern Skane appears
to be lost (cf. Roman, 1913). The other specimen standing under that name in the
collection was taken by R. Malaise around Tornetrask, and is a later addition.
Two specimens standing under claripennis in the Stockholm museum were exam-
ined in 1935, and regarded as distinct from others then known to me. One bears the
determination label of A. Roman who had, evidently, recognized it subsequently to
the appearance of his 1913 paper, but had not, I believe, published any note to this
effect. R. Malaise kindly sent me this female specimen for further study, and I now
agree this species to be the true claripennis for the reasons which follow.
Roman wrote (1913) that he found, in Thomson’s description of claripennis, no
reliable difference from adspersus Hart., that Thomson had not mentioned this species
with its synonym /epidus Higr., and that he thought Thomson had redescribed the
‘to him unknown’ adspersus as claripennis. In actual fact he redescribed adspersus
as lavicinus, and used the words ‘... ab E. lepido pictura uberiore distinctus’. So his
conception of /epidus may have been the species now described as confusus.
Thomson gave for claripennis ‘E. marginatorio . . . valvula ventrali 2 apice magis
acuminata discedens’. Now the degree of pointedness of the female hypopygium is,
when not extreme, a difficult character to use, for the appearance depends largely
on the degree of folding or flattening of that organ: in amictorius (marginatorius) it
seems inconstant ; but the condition in the species here identified as claripennis might
fit ‘magis acuminato’, whereas that in claripennis Cush., I think, would not, and
tricolor has the female hypopygium beyond all comparison more pointed. Had
Thomson had éricolor, it is unlikely that he would have omitted to mention the red
coloration of the underside of the gaster. Thomson wrote ‘mesosterni acetabulis haud
bidentatis’: in this species the acetabulae (margining the fore coxal cavities) are
distinctly developed and laterally raised but not ventrally bidentate, whereas in
claripennis Cush. and in tricolor they are absent or rudimentary.
Very closely related to amictorius Panz., from which it differs as given in key to
species (p. 366).
Another authority to mistake claripennis was Kriechbaumer: two specimens he
360 A REVIEW AND A REVISION OF
determined thus prove to be vellicatus Cush. They were bred from the same host as
were Cushman’s types.
E. claripennis Thoms. diagnosed from the following material: SWEDEN, Dalarne,
Fulufjall, r 9, vii.tg27, on snow field, K. H. Forsslund (neotype) ; Smaland, 1 9,
Boheman ; det. A. Roman; ?Blekinge, Aryd, 1 3, 1 9, ex Neodiprion sertifer Geoftr.,
Morris, Cameron, & Jepson, det. G. J. K.
Further material seen: AUSTRIA, Styria, Bésenstein massif, 1,800 m., I 3, 7.vii.1950;
Ardning, 600 m., I 9, II.viii.1950, G. Heinrich: FINLAND, Kangasala, 1 gf, R. Frey.
Exenterus adspersus Hartig
1838 Exenterus adspersus Hartig, Jahresber. Fortschr. Forstw., 1: 271 (non Morris, Cameron, &
Jepson, 1937).
1858 Exenterus lepidus Holmgren, Svensk Vet.-Akad. Handl. 1: 231 (non Thomson, 1888).
1888 Exenterus laricinus Thomson, Opuscula Entomologica, fasc. 12: 1254-1255.
1911 Exenterus adspersus Schmiedeknecht, Opuscula Ichneumonologica, 5: 2307.
1913 Exenterus adspersus Roman, Ent. Tidskr. 34: 127.
1940 Exenterus adspersus Cushman, U.S. Dept. Agric. Misc. Publ. 354: 6-7.
Holmgren (1858) himself suggested the possibility of his species being synonymous
with adspersus Hartig. Thomson (1888), however, redescribed adspersus as laricinus
(see below), which he compared with lepidus Hlgr. To which species he attributed
lepidus must remain in doubt, for none is to be found under that name in his collection.
Apart from Thomson, no one questioned the synonymy until in 1935 the present
author wrongly, as will be seen below, placed another species as lepidus (Kerrich,
1942).
Dr. E. O. Engel very kindly directed my search for the type in the Zoologische
Statssammlung in Munich, but we were unable to find it. Cushman (1940) gave the
first clear picture of this species, including a figure of the apex of the gaster: he
also identified the species with laricinus Thoms., which latter name he placed in
synonymy. I can support this procedure with further evidence, for I was able to
borrow from Munich a specimen (dated 9.ix.1884) placed as adspersus by Kriech-
baumer who, more than anyone, should have been familiar with Hartig’s work. This
specimen I found, by direct comparison in Lund, to agree with Thomson’s laricinus.
It agrees, moreover, with the characters given by Cushman (1940).
Dr. Roman, when arranging the collection of Swedish Ichneumonidae in the Stock-
holm museum in the years 1937-1940, selected lectotypes of the Holmgren species.
The lectotype of Exenterus lepidus is a female specimen taken by Boheman in the
Swedish province of Dalarne: by direct comparison in 1947, I find it specifically
identical with the above-mentioned Munich specimen, though rather more strongly
sculptured.
Further material examined: ENGLAND, Hants, New Forest, Matley Bog, 1 g em.
1935, ex Diprionid cocoon (viii.1934), A. J. Duarte (Kerrich, 1942, as lepidus Hlgr.) ;
Surrey, Dorking, Westcott, 1 g, 1.ix.1948, G. J. K.; Beds., Clophilland Apsley Heath,
2 $3 em. iv.1948, ex Diprion similis Hart. (larvae ix.1947), V. H. Chambers: BEL-
GIUM, I 4, coll. Wesmael: SWEDEN, material in Holmgren coll.; Norrbotten, Boden,
I 9, 9.vii.1938, K. J. Hequist: FINLAND, Karislojo, material reared from Gilpinia
THE CTENISCINI OF THE OLD WORLD 361
pallida Klug by R. Forsius (Forsius, 1911: 100; Roman, 1911: 201) ; Karislojo, 1 g,
r 9, ex Diprion pini L., R. Forsius (mus. Paris): GERMANY, 5 29, coll. Ruthe;
Schirmer, I 2, 2 $4, ix.1905, Buckow (mus. Stockholm) ; Saxony, Vogtland, 1 3, ex
Diprionid cocoon (mus. Paris): SWITZERLAND, Nyon, I g, Cheyr: FRANCE, Toulouse,
2 6d em. iv.1925, ex Diprionid cocoons, Delmas.
For a note on the British specimen recorded as lepidus Hlgr. see below under
confusus. ;
Biology. Bred from Diprion pini L., D. similis Hart., and Gilpinia pallida Klug (see
above) : recorded also from D. nipponicum Rohwer, G. polytoma Hart. and Neodiprion
sertifer Geoffr. (Cushman, 1940).
Exenterus tricolor Roman
For characterization, see key to species and subsequent discussion of differences
between the males. I find no reliable difference from adspersus Hart. in the amount
of yellow coloration of the males, such as is suggested by Roman (1913: 127) ; but
an average difference might be demonstrable if the two species were taken together
in several different localities.
Material examined as follows: FINLAND, Finstrém, 1 3, 24—29.vii.1946, W. Hellén;
Vichtis, r g, R. Frey: SWEDEN, type material of A. Roman; Skane, Skaralid, 1 3,
I.vii.1938, D. M.S. & J. F. Perkins: GERMANY, I 9, coll. Hiendlmayr (mus. Munich),
compared with type ; Kéln, 1 9, 16.viii., I g, 23.vili.1946, W. Aerts: CZECHOSLOVAKIA,
Riesengebirge and Tyssa, 5 29, Morris, Cameron, & Jepson: AUSTRIA, Styria, Admont,
Kaiserau, 1,100 m., I g, 10.vi.1950, G. Heinrich.
Biology. Reared from Gilpinia polytoma Hart. in Finland (Forsius, 1932) ; and from
this species and Neodiprion sertifer Geoffr. in Czechoslovakia and Saxony by Morris,
Cameron, & Jepson (1937). These three authors give an extensive account of the
life-history.
Exenterus vellicatus Cushman —
For characterization, see key to species and subsequent discussion of differences
between the males.
Material examined as follows: GERMANY, I 9, 30.vil.1884, I 2, 23.vii.1886, both ex
Gilpima polytoma Hart., 1 g, I1.viii.1888, coll. Kriechbaumer (mus. Munich) ;
Holstein, Trittau, Hahnheide, 2 99, ix.1945, 3 $d, vi.1946, G. Heinrich: CzEcHO-
SLOVAKIA, several localities as recorded, 5 gd, 11 99, ex Gilpinia polytoma Hart.,
Morris, Cameron, & Jepson (1937: 377, as Exenterus sp.). Recorded as established
in Canada (Mason, 1951 April: 2209).
Exenterus confusus sp. n.
1888 ?? Exenterus lepidus Thomson, Opuscula Entomologica, fasc. 12: 1255 (non Holmgren,
1858).
1937 Exenterus adspersus Morris, Cameron, & Jepson, Bull. ent. Res. 28: 375-376 (non Hartig,
1838).
1940 Exenterus claripennis Cushman, U.S. Dept. Agric. Misc. Publ. 354: 3, 9 (non Thomson,
1883).
ENTOM. II. 6 + ah
362 A REVIEW AND A REVISION OF
This species was taken in several specimens in northern Finland in 1935 by the
present writer. It runs down in Schmiedeknecht’s key to adspersus Hartig, and was
determined thus in September 1935; with how much assistance from Dr. Roman
I cannot now remember, but at least with his concurrence.
It is clearly the species identified by Cushman (1940) as claripennis Thoms. It has
the posterior face of the propodeum generally less nearly perpendicular than in
amictorius (marginatorius), as stated by Cushman; whereas in the species here
identified as claripennis that face is rather more so. Reasons for rejecting Cushman’s
interpretation are given above.
As Cushman (op. cit.) supposed, this is indeed the species reported by Morris, —
Cameron, & Jepson as adspersus Hartig: I saw part of their material, and bear part
responsibility for the identification. Later, after seeing the true adspersus, I supposed
that this species must be lepidus Holmgren. The males, however, had still not been
separated: a British specimen of adspersus was misidentified and recorded as lepidus
in 1942; but in the absence of further explanation, the usual synonymy was followed
by Kloet & Hincks (1945).
Diagnosis. Species with the characters given by Cushman (1940) for claripennis
Thomson, and with those given for this species in the key below. In the female sex
very distinct from adspersus in having the hypopygium not acute or ventrally
folded, the third valvulae not reaching the apex of the gaster, and the mesosternum
and pleura not largely yellow-marked.
Head moderately to strongly narrowed behind eyes, shallowly emarginate behind:
on vertical region, temples and most of genae rather coarsely to coarsely and closely
punctate; near malar space, and on lower face in greater part (except in smallest
specimens) and upper face reticulate-punctate: upper face often with a distinct
median keel. Clypeus generally distinctly alutaceous, rather finely to rather coarsely
punctate above the pre-apical impression ; rather strongly delimited basally ; rather
narrowly rounded apically. Malar space short, as usual in the genus. Hypostomal
costa slightly raised; genal costa meeting it a good quarter the way back to the
foramen magnum.
Antennae with scape (Fig. 38) weakly convex dorsally, weakly excised externo-
ventrally ; with flagellum 31- to 36-segmented.
Pronotum with dorsal transverse furrow broad and shining ; with epomiae strong ;
laterally shallowly and rather closely punctate above, generally strongly longitud-
inally rugose below. Propleura rather sparsely punctate, more closely so at sides.
Mesoscutum rather strongly convex (some variation in this) ; with notaulices just indi-
cated ; rather shallowly but more or less coarsely punctate discally, more finely punc-
tate behind. Scutellum not margined to middle; weakly convex and moderately to
coarsely punctate above; smoother and falling rather abruptly behind. Mesopleura
in upper half shining, moderately punctate in front, finely so around speculum; in
lower half coarsely and closely punctate: sternaulices more or less superficial: meso-
sternum and prepectus moderately and closely punctate: epicnemial carina scarcely
elevated: mesolcus not strongly impressed ; narrow in front, then broadening and
sometimes transcarinate; generally well closed behind. Propodeum with costae
mostly rather weak, but usually normally complete ; for the most part weakly reticu-
THE CTENISCINI OF THE OLD WORLD 363
late-rugose, but.on area pleuralis shining and finely punctate in front ; with posterior
face generally less abruptly declived than in amictorius Panz. and adspersus Hart.:
area superomedia strongly contracted from costulae to base and not basally delimited.
_ Gaster with tergites I and greater part of 2 rather strongly and closely reticulate-
rugose, the sculpture somewhat weaker and diffuser towards apex of tergite 2; with
tergite 3 in about basal half and at sides, and sometimes base of 4, closely reticulate,
sometimes suggesting fine longitudinal rugosities; occasionally the reticulation is
more extensive, but otherwise tergites 3 to 6 rather coarsely punctate, progressively
more sparsely and shallowly so; with tergites 7 and 8 almost always finely and
sparsely punctate. Petiolar segment 1-2 to 1-7, generally 1-4 to 1-5 times length of its
greatest breadth; with dorsal keels of moderate strength and distinct for about
three-quarters length of segment in female, rather stronger and almost percurrent in
male. Apex of female gaster much as in amictorius Grav., but third valvulae perhaps
on the average a little broader than in that species, and decidedly broader than as
delineated by Cushman for that species (1940:3, fig. c): hypopygium not acute and
not or scarcely folded in mid line: third valvulae not reaching apex of gaster.
Fore wings with basal and disco-cubital veins slightly divergent ; with areolet sub-
sessile or shortly petiolate ; with nervus parallelus emitted about in middle of brachial
cell. Hind wings with nervellus opposite or slightly antefurcal, intercepted about
in middle.
Tarsal claws with no more than a trace of pectination at base, not evident in male.
Length 6 to Io mm.
Head black, with the following parts deep, bright flavous: palpi, mandibles except
at apex and, in male, often at base, clypeus except along suture and sometimes at
apex, and lower face except for a median band. The yellow coloration extends on
inner orbits at least a little way above antennal sockets ; and also through malar space
on to the genae, where it adjoins the mandibular base and extends a little way up
outer orbits, but does not reach hypostomal costa nor approach genal costa. Antennae
black, merging to ferrugineous above, dull fulvous below; with scape and sometimes
pedicellus flavous-marked beneath. Thorax and propodeum black, with the following
deep, bright flavous: greater part of scutellum and metascutellum ; pair of roughly
triangular marginal marks on mesoscutum covering notaulices, occasionally reflected
on pronotum ; median pair of marks on anterior margin of pronotum, often coalescing
and extending towards lower angles; paired marks always below tegulae, often just
above epicnemial carina, occasionally beside hind end of mesolcus, and occasionally
by hind angle of area dentipara. Legs black, with the following deep, bright flavous,
paler on hind leg: fore coxa extensively below and often round apex ; mid coxae less
extensively, sometimes only slightly below; trochanters and trochanterelli, except
that those of hind leg are dark-marked; fore and, less extensively, mid femora in
front and at apex; fore and mid tibiae and tarsi mainly; hind tibiae in about
basal two-thirds except at extreme base; hind tarsal segments progressively more
narrowly at base: hind coxae not pale marked. Wings with veins and stigma ferru-
gineous, basally paler. Gaster black, with the following bright flavous in female:
rather broad apical bands, which often do not approach the sides, on first two tergites ;
and usually a series of much narrower apical bands on tergites 3 to 6: in male the
364 A REVIEW AND A REVISION OF
narrower bands may be developed on tergites 3 to 5 only; and in darkest forms even
the band on tergite 2 is undeveloped.
Described from the following material: CZECHOSLOVAKIA, Kunzak, 1 ? (type) from
cage, 18.x.1933, I 2 em. 4.xi1.1933, I 2 em. 29.i1.1934; Hradec, 9 go, 7 29 em.
vi-vii.1936 ; Dobris, I 3, 4 29 em. vi.1936; Plese, 2 gg, I 2 em. vi.1936; Allrestice,
3 PY em. vi.1936 ; Tyssa, 2 $3; unlocalized, 7 33, 3 29; Riesengebirge, 1 9 (undated),
Morris, Cameron, & Jepson: GERMANY, unlocalized (probably Germany), 2 gg, O
Schmiedeknecht (mus. Geneva); Heilbrunn, 1 9, 8.vi.1888, J. Kriechbaumer ; Bay-
rische Wald, Waldmiinchen, 1 g, 9.vi.1948, 1 g, 1-8.ix.1948; Holstein, Trittau,
Hahnheide, 1 9, 29.vi.1946, G. Heinrich: SWEDEN, Lapland, Tornetrask, R. Malaise
(mus. Paris): FINLAND, Kuusamo, Liikasenvaara, 3 $3, I 9, 9.vii.1935 ; Mutkatun-
turi, I 9, 3I.vll.1935, G. J. K. All the above, except the type, have been designated
paratypes.
Additional material seen: AUSTRIA, Styria, Admont, Kaiserau, 2 $4, 2 29, 1-8.vii.
1950, G. Heinrich. Recorded (under claripennis Thoms.) as established in Canada
(Mason, 1951 April: 229).
Biology. The Czech material was reared by Morris, Cameron, & Jepson from
Gilpinia polytoma Hart. with the exception of the specimen from the Riesengebirge
which was from Neodiprion sertifer Geoffr.: these authors give considerable detail of
their work (1937). The Finnish specimens were beaten from Picea on cool days.
KEY TO EUROPEAN SPECIES OF EXENTERUS HARTIG
1. Petiolar segment trisinuate (Fig. 40): hind tibia bearing a very minute spur:
genal costa double, conformed much as in Exyston genalis Thoms. (Fig. 51):
fore wings conspicuously clouded along fore margin . . aptarius Grav.
Petiolar segment not thus trisinuate (e.g. Fig. 39): hind tibia without a minute
spur: genal costa not double: wings hyaline, or moderately and evenly infu-
mate . : : ; ; : ‘ : ; ‘ ‘ ‘ 2
2. Clypeus broadly rounded at apex and neither transversely impressed nor re-
flexed: antennae in female barely half as long as body, in male much shorter
than body: gaster broad, with tergite 2 much broader at base than long (in
female aac twice as broad as long): claws in female usually pectinate.
(abruptorius group) 3
Clypeus more or less distinctly transversely impressed before the strongly
rounded and reflexed apex: antennae in female much more than half as long
as body, in male nearly as long as body: gaster generally narrow, with tergite 2
nearly or quite as long as basally broad : claws in female not or weakly pectinate
(amictorius group) 5
3. Margin of cavities of fore coxae not raised, except weakly at sides: in side view,
greatest length of head behind eye a little less than eye length: malar space
almost equal to distance between mandibular apices: gaster in female more
than half yellow: mesosternum conspicuously yellow-marked: yellow of first
tergite extending far towards base laterally ; ‘ , ortolus Hart.
Margin of cavities of fore coxae raised: in side view, greatest length of head —
THE CTENISCINI OF THE OLD WORLD 365
behind eye a little more than eye length: malar space only about half distance
between mandibular apices: gaster in female more than half, and mesosternum
black: yellow of first tergite not or little wider laterally than medially . . 4
oo
amict,
ae =>
\ adsp,
_ Fics. 34-38. Antennal scape, &c., in sinistro-lateral view, of Fig. 34. Exenterus amictorius Panz.
¢ Fig. 35. Exenterus claripennis Thoms. Fig. 36. Exenterus adspersus Hart. Fig. 37. Exenterus
; vellicatus Cushm. Fig. 38. Exenterus confusus sp. n.
vellic, conf.
88.
_ Fics. 39-40. Petiolar segment of Fig. 39. Exenterus amictorius Panz. 9. Fig. 40. Exenterus
apiarius Grav. 9.
4. Margin of cavities of fore coxae not bidentate: claws in female pectinate: larger
species, 8—II mm. ; ‘ . abruptorius Thunb.
Margin of cavities of fore coxae strongly raised and strongly bidentate: claws
simple: smaller species, 7 mm. ‘ ‘ . simplex Thoms.
5. Gaster broad, as in abruptorius group, mainly chestnut-brown i in colour, and with
segments I—2 strongly punctate but not rugose: costulae of propodeum absent,
or weak and incomplete . ‘ ‘ . tctericus Grav.
Gaster relatively narrow, as described before, black or black and yellow, and with
366 A REVIEW AND A REVISION OF
segments I-2 strongly rugose-punctate (except in smallest specimens of some
species): costulae of propodeum distinct . : ; é . : 6
6. Margin of cavities of fore coxae raised. ;
Margin of cavities of fore coxae not raised, or only with traces of keels laterally 8
7. Margin of cavities of fore coxae bidentate, emarginate between the teeth and
appearing discontinuous in middle: scape very weakly excised externo-ven-
trally (Fig. 34) : ‘ amictorius Panz. (= marginatorius Fabr.)
Margin of cavities of fore coxae not bidentate, appearing continuous and almost
straight: scape rather weakly excised externo-ventrally (Fig. 35)
: : . : : . : claripennis Thoms.
8. Males: see discussion immediately following.
Females . : ; : : ; : : : : ‘ : 9
g. Gaster laterally beyond tergite 3, genitalia and hypopygium rufo-castaneous:
hypopygium very acute (Cushman, 1940, fig. B), and without a distinct, broad
stramineous or white hind margin: head moderately to strongly narrowed
behind eyes, not as in female adspersus Hart. . ‘ . tricoloy Roman
Gaster not laterally rufo-castaneous: hypopygium not as above. .- =
1o. Temple strongly convex, nearly as long as short diameter of eye: hypopygium ~
sharply folded and moderately acute (Cushman, 1940, Fig. A): third valvulae
very distinctly projecting beyond apex of tergites 7 and 8 (they project slightly —
beyond in ¢vicolor Roman but not in other described European species)
‘ : ‘ , : : : , adspersus Hart.
Not as above . , ' ‘ : . ‘ ; ; ; |
11. Hypopygium pinched ventrally near apex, its lower margin in profile very
strongly curved, its apex thin, wavy, broadly white (Cushman, 1940, fig. D):
mesosternum and especially propodeum largely yellow-marked
vellicatus Cush, |
Hypopygium not at all as above, not dissimilar from that of amictorius (mar-
ginatorius) (Cushman, 1940, fig. C), though ovipositor sheaths broader than as
there represented: mesosternum and propodeum not largely yellow-marked
confusus sp. Nn.
SEPARATION OF MALES OF Four SPECIES
Whereas the species adspersus Hart., tricolor Roman, vellicatus Cush., and confusus
nov. are clearly distinguishable in the female sex upon the form of the hypopygium
and third valvulae, and certain other features, such good external characters are not
available for the separation of the males. Thus I find that in male adspersus the head
may be shorter and distinctly narrowed behind the eyes; and some males that I
believe to be vellicatus Cush. have the gaster laterally rufo-castaneous as in tricolor
Roman. This condition should be distinguished from that obtaining in some small
specimens of other species, in which the whole gaster, except the yellow-coloured —
parts, is dull castaneous rather than black.
Taking specimens and the description of confusus nov. as a starting-point, the —
THE CTENISCINI OF THE OLD WORLD 367
other species have been compared with these in both sexes. It is hoped that the
resulting observations will enable the majority of specimens to be placed with confid-
ence: the greatest difficulty appears to be the separation of males of tricolor Roman
and vellicatus Cush.
E. adspersus Hart. differs from confusus as follows: lower face reticulate to some
extent below antennae, but in greater part discretely punctate: clypeus finely to
rather coarsely and rather sparsely punctate, on the average less strongly punctate
than in confusus: scape somewhat strongly convex dorsally (Fig. 36): scutellum
rather weakly to moderately convex, generally much more evenly so than in confusus,
i.e. seldom falling at all abruptly behind: posterior face of propodeum more abruptly
declived: propodeum more strongly, often quite strongly reticulate-rugose: yellow
coloration of gena reaching hypostomal costa, and reaching or approaching genal
costa: male mesoscutum more often, like that of female, with a pair of discal yellow
marks between tegulae: propleura sometimes pale-marked: mid coxae and femora of
male extensively pale-marked, as much so as on fore leg of confusus: hind coxae
sometimes pale-marked beneath.
E. vellicatus Cush. is best compared with adspersus, with the colour differences of
which from confusus it very much agrees, though the extent of genal coloration is
_ often intermediate, as is the form of the scape (Fig. 37). Epistoma alutaceo-punctate,
very little raised: notaulices a little more distinct than in adspersus or tricolor:
_ sternaulices more distinctly impressed : mesosternum decidedly flattened, with mesol-
cus more sharply impressed, scarcely broadening behind: area pleuralis of propodeum
weakly rugose behind and below (quite strongly so in adspersus). The head is often
distinctly raised behind the ocelli, but unfortunately this is not invariably so. In
female a yellow mark runs along the sternaulices, and in male there is usually a trace
of this.
E. tricolor Roman differs from vellicatus as follows: scape weakly convex dorsally,
as in confusus: epistoma not thus flattened, usually smooth between punctures: genal
colour as in adspersus or intermediate, as in vellicatus: mesosternum and pleura as
described for adspersus, but speculum extends farther forward and pleuron is more
finely punctate in front of it.
Genus EX YSTON Schigdte -
1839 Exyston Schigdte, Guérin’s Mag. Zool. (2) 1: 12 (separate pagination).
1877 Tricamptus (Forster) Woldstedt, Bull. Acad. Sci. St. Pétersbourg, 28: 454.
_ Our knowledge of this genus in Europe has been in a most unsatisfactory condition.
Unfortunately I have not examined the Gravenhorst types. The following specific
names require consideration: albicinctus Grav., albocinctus Grav., tricolor Grav., sub-
nitidus Grav., cinctulus Grav., sponsorius (Fabr. non Jurine) Grav., carinatus Thoms.,
genalis Thoms., calcaratus Thoms., phaeorrhaeus Hal., pratorum Woldst. = brevipetio-
latus Thoms. = triangulatorius Hlgr. non Grav. (Anisoctenton).
Of the above I am, in the present work, making calcaratus Thoms. the type of a new
genus, Parexyston.
One of the two commonest European species has probably been sufficiently charac-
368 A REVIEW AND A REVISION OF
terized for its recognition, and has been best known as brevipetiolatus Thoms. Roman
(1939) thought it to be Cteniscus phaeorrhaeus Haliday but, as will be shown below,
that conclusion is now disproved. Hellén (1937) placed it in synonymy with pratorum
Woldst. ; and there is not, to my knowledge, any valid prior name.
The second of the two most common species, which is very variable in colour, has
the genal costa continuous or almost so; a fact first recognized by Thomson (1883),
who took it to be the typical species and placed it under Gravenhorst’s name cinctulus.
The type of cinctulus Grav. 1820 is not to be found in the Turin museum (see Kerrich,
1950), so a lectotype should be selected from the Gravenhorst collection at Wroclaw ;
but I accept the descriptions of 1820 and of 1829 as referring to this species, and I
find no clear indication that any of the colour varieties described should refer to
genalis Thoms. (see below). An intensive study of cinctulus Grav. confirms what had
already been suspected, namely that the form described as carinatus Thoms. is within
its limits of variation and is not specifically distinct.
Consideration must now be given to the name sponsorius. Pfankuch and Schmiede-
knecht place sponsorius Grav. as a variety of cinctulus, but Gravenhorst attributed
the name to Fabricius who would, of course, have priority. Dr. Kai Henriksen wrote
to me that the type of sponsorius is not in the museum at Copenhagen. I am indebted
to Kustos Dr. Olaw Schréder for information from Kiel. It is not known whether
Pfankuch ever examined the Fabrician collection there. No specimen labelled Ich-
neumon sponsorius is present. The type of Cryptus sponsor is present, labelled in ©
Fabricius’s own writing: it is a female with a distinctly exserted ovipositor, and may
well have been correctly identified by later authors. There is also a specimen labelled
by Fabricius ‘sponsatorius’: on this the mid and hind tibiae each bear two spurs, so
it can be no Cteniscine. It is a large species, about 15 mm. in length excluding appen-
dages, and does not appear to agree with Fabricius’s descriptions of sponsorius. It —
seems best to drop the name sponsorius unless or until another type can be discovered.
Exyston sponsorius Grav. as identified by most later authors is a form of cinctulus
Grav. ; and I find no clear indication in Gravenhorst’s descriptions that it could refer
to any other species. F
Thomson’s diagnosis of genalis (1883: 883) was quite inadequate, and has failed to
lead subsequent authors to its recognition. In his collection are three specimens, all
female and all belonging to the species figured and redescribed as genalis in the present
work. Unfortunately, no one of these can with assurance be selected as lectotype:
one, however, is labelled ‘coll. Ros.’ ; and since, as Dr. Kjell Ander informs me, Rosen-
schidld collected in Vastergétland, this specimen could be the type. Thomson’s
words, ‘Long. 3-34 lin. E. cinctulo colore statura et magnitudine simillimus’, cor-
roborate this interpretation of the species. I was able to tell A. Roman that this was
the species placed as genalis in the Lund collection; and his acceptance of this is”
evidenced by the series he arranged in the Swedish collection at Stockholm. He had
known the species as a species, and he told me he had recognized it among Forster’s
material in the Vienna museum under the manuscript names apicalis (9) and aberrans
(3); but he had been misled by Thomson’s diagnosis. The species he recorded as
genalis from Kamtchatka (Roman, 1931) is another, nearer related to pratorum
Woldst. (see below). .
THE CTENISCINI OF THE OLD WORLD 369
Until 1946, albocinctus Grav. and tricolor Grav. were both unknown to me, and
I wondered whether they would prove to be mere forms of the variable cinctulus
Grav., as other authors had supposed. Pfankuch examined the types and amplified
the descriptions (1906). Later (1925: 57), he recorded both forms from North Slesvig ;
though he wrote of albocinctus ‘Wohl nur ein Varietat von E. cinctulus’. Schmiede-
knecht (1912) mentioned norecord of either subsequent to those of Gravenhorst (1829),
though de Gaulle listed both in his catalogue (1908).
H. G. M. Teunissen sent me a list of the Cteniscine species known to him to
occur in Holland, including Exyston albocinctus Grav. He most kindly responded to
my request for the loan of this species by sending a Dutch male taken in 1887, and
since recorded (Teunissen, 1948). Though only about 74 mm. in length and, in
general, of rather paler coloration, this specimen agrees very well with the descrip-
tions given by Gravenhorst and by Pfankuch. It is unquestionably a good species.
The petiolar segment is, as stated by Gravenhorst, relatively less narrow than in
cinctulus. | am pleased to acknowledge the credit due to Dr. Teunissen for the re-
discovery of this species.
A small sending received shortly afterwards from R. Malaise contained another
male of this species, bearing no determination label, and a female which I associated
with it. Then I discovered that this female agrees very well with the descriptions of
tricolor Grav. I have no hesitation in maintaining this association of the sexes, so
tricolor Grav. falls in synonymy with albocinctus Grav. It may be objected that
Gravenhorst recorded his specimen of éricolor as a male, and that Pfankuch did not
disturb this interpretation; but ‘vomeriform’ third valvulae of a female Cteniscine
are, when withdrawn, folded over, and they present an appearance very similar to
that of a pair of male claspers. In such case the closest scrutiny is required to ensure
correct sexing.
Specimens in de Gaulle’s collection were unlabelled, but have now been redis-
covered. Now that I know the species, I am satisfied that the description of Ichneu-
mon albicinctus Grav. 1820 refers to it, though the type cannot be found in the Turin
museum.
Cteniscus phaeorrhaeus Haliday was duly included by Morley, in the fourth volume
of his monograph (Ig91I: 207), under Exenterus (= Exenterus+Cteniscus as here
accepted). When examining the Haliday collection in the Dublin museum two years
later, Morley selected a lectotype, gave a brief description of it, and placed it next
Cteniscus mitigosus Grav., but did not identify it with any species described elsewhere
(1913: 261). At the same time he declared that the record given in his monograph
was an error. Apart from a record by Haines (1931), the matter rested there until
Roman (1939: 195), after correspondence with A. W. Stelfox, identified it with
Exyston brevipetiolatus Thoms.
Upon my inquiring whether Roman’s synonymy could be supported by fuller and
more positive evidence than was contained in his rather brief note, Stelfox sent me
Morley’s lectotype for examination, and wrote that in his opinion it could not be the
type, and that Roman had agreed with this. It is a female but, in other respects, fits
Morley’s diagnosis. It runs down in Schmiedeknecht’s key to Cteniscus dorsator
Thunb. (= limbatus Hlgr.), and fits the description of that form. It must be rejected
ENTOM. It. 6 ZZ
370 A REVIEW AND A REVISION OF
as type of phaeorrhaeus Hal. since, as indeed is stated by Morley, the fore legs are
not basally black, and the gaster is not fulvous at the apex but red discally and bright
flavous banded near the apex. More than one twentieth-century specialist of repute
has mistaken a female for a male Cteniscine, yet Haliday recorded both sexes of Ct.
aurifluus and may be presumed to have distinguished them, especially as, so Stelfox
tells me, he often dissected specimens when describing species. Furthermore, another
female specimen of dorsator Thunb. in the collection bears the label ‘cephalotes’ in
Haliday’s writing.
A. W. Stelfox has now given me the following fuller account of his correspondence
with A. Roman. He sent a description and figure of Morley’s lectotype, which
Roman identified as dorsator Thunb. He had failed to find in Haliday’s collection
a specimen which could, in his opinion, be the type, so.he sent from his own collection
a male specimen of a species he had taken frequently in Ireland, had not identified
with any other author’s description, but thought most likely to be Haliday’s phaeor-
rhaeus. Roman identified it with brevipetiolatus Thoms., and agreed with the con-
clusion. I have now seen the actual specimen and agree with the identification.
Indeed, the conclusion appeared highly probable. The species agrees with Haliday’s
description in the male sex diagnosed. It is of interest that the specimen recorded as
Exenterus phaeorrhaeus from Woodstreet by Haines in his Hants & Dorset list (1931)
does belong to brevipetiolatus Thoms. Haines used Morley’s monograph, and the
diagnosis there published, which Morley later (1913) declared to have been an error,
seemed to me to refer to the same species. Mr. Morley kindly sent me the specimen
on loan, which enabled me to confirm this supposition, and to see from a second label
that he had placed the specimen correctly, later in 1911.
On first glancing at the box from the part of the Haliday collection relevant to
the present study containing, among other Tryphoninae, the Cteniscini, I looked for —
brevipetiolatus Thoms. and caught sight of what I took to be a large dark male.
Upon closer examination, I was astonished to find it was a male of a species I had
received from Czechoslovakia from Professor Gregor, and which I had believed to
be undescribed. This specimen I designate the type. The pinhead bears green sealing-
wax: Haliday’s mode of indicating an Irish insect. I have examined every Cteniscine
in that box from which brevipetiolatus is absent and, in my view, no other specimen
could possibly serve as type. Moreover, it fits Haliday’s diagnosis even better than
does brevipetiolatus: the fore and mid femora, except internally, and also the tro-
chanters are distinctly ferrugineous, and the coxae are black or almost so. The hinder
part of the gaster is much more outstandingly fulvous: in this respect the specific
name may be taken as indirect supporting evidence, for ‘ phae-orrhaeus’, as I am told
by classical scholars at the University of Manchester, means ‘bright-coloured rump’.
Finally it may be noted that Morley (1911) provided a false clue by misinter-
preting Haliday’s manuscript catalogue, in which there stands against phaeorrhaeus
the mark for ‘certainly Irish’ but not the mark indicating ‘common or in many ~
collections’.
Tryphon subnitidus Grav. was placed in Exyston by Pfankuch (1906). Schmiede- —
knecht (1911), being unable to place the species, omitted mention of it; but Morley —
(1911) recorded it as British from an old specimen in the British Museum. This
fe
=
THE CTENISCINI OF THE OLD WORLD 371
proves to be a small, dark male Antsoctenion rubiginosus Grav., as is a specimen so
placed in the Melville-Bury collection, now in the Manchester Museum. C. Morley
very kindly sent me on loan two specimens from the series taken by him in Suffolk
in 1919 and recorded as subnitidus (1936), but these agree with my description of
Parexyston calcaratus Thoms.
In 1949, when sorting specimens for the redescription of phaeorrhaeus Hal., I found
that a second, closely related, species was mixed with them, represented by three
specimens from Switzerland and one from Spain. At last a species had been dis-
covered whose male I could reconcile with Gravenhorst’s description and Pfankuch’s
diagnosis of subnitidus.
Exyston cinctulus Gravenhorst
1820 Ichneumon cinctulus Gravenhorst, Mem. Accad. Sci. Torino, 24: 359-361.
1820 Ichneumon sponsorius Gravenhorst, ibid.: 362.
1829 Mesoleptus cinctulus Gravenhorst, Ichneumonologia Europaea, 2: 37-41.
1839 Exyston cinctulus Schiedte, Guérin’s Mag. Zool. (2) 1: 12 (separate pagination),
1883 Exyston cinctulus Thomson, Opuscula Entomologica, fasc. 9: 882.
1883 Exyston carinatus Thomson, ibid.: 882-883.
1906 Exyston cinctulus Pfankuch, Z. Hym. Dipt. 6: 21.
Head moderately to rather strongly narrowed behind eyes and strongly rounded ;
moderately emarginate behind: on inner orbits finely, on vertex and temples finely
to moderately punctate, and on genae more coarsely so: epistoma more or less
coarsely reticulate or rugose-punctate, occasionally in small specimens discretely
punctate: upper face generally with a distinct median keel, reticulate-punctate to
rugose. Clypeus normally not strongly transverse ; rather sparsely beset with punc-
tures of different sizes, fine to coarse ; sharply delimited basally ; narrowly to broadly
rounded at apex. Malar space about equal to distance between mandibular apices
in male, distinctly longer in female. Hypostomal costa strongly raised, the elevation
- extending well back towards foramen magnum: genal costa continuous (Fig. 50) ;
strongly raised in female, less strongly so in male. A weak costa is generally de-
veloped, running outward from the hypostomal costa toward the genal; but this
may be broken up into a series of transverse rugae.
Antennae with scape deeply excised externo-ventrally (Fig. 53) ; with postannellus
about 1-9 times length of the following segment ; with flagellum 28- to 37-, usually
_ 32- to 34-segmented.
Pronotum with epomiae sharp and generally fairly strong; laterally rather finely
to moderately, occasionally quite finely punctate above, finely punctate to rugose
below. Propleura rather finely to moderately punctate. Mesoscutum rather weakly
to strongly and evenly convex ; with notaulices always indicated but extremely super-
ficial; rather finely punctate medially, finely to very finely so laterally. Scutellum
rather weakly to rather strongly convex ; margined to near middle ; rather coarsely
punctate or rugose-punctate in front, finely reticulate-rugose behind. Mesopleura
rather coarsely punctate in greater part, generally rather finely so-on region of
speculum: sternaulices of moderate strength, rather sharply impressed: mesosternum
closely and finely punctate: epicnemial carina strongly raised in female, moderately
372 A REVIEW AND A REVISION OF
or rather strongly so in male ; undeveloped above sternaulices, medially emarginate:
mesolcus moderately impressed, generally transcostate in male, closed behind. Pro-
podeum with costae weak but normally complete; finely to moderately punctate
and shining above, merging to rugose, especially on spiracular and posteromedial
areas, but with area pleuralis usually discretely punctate: area superomedia usually
distinctly longer than broad but sometimes subquadrate.
Gaster much more spindle-shaped than clavate. Petiolar segment generally elon-
gate and slender, of female 2-2 to 3-0, usually 2:5 to 2-8, of male 1-8 to 3-0, usually
2:2 to 2°6, times length of its greatest breadth ; bearing lateral hairs distinctly longer
than half its least breadth; generally moderately to strongly auriculate basally, but
occasionally rather weakly so; with dorsal keels usually not sharp, distinct for half
to more than four-fifths length of segment; rather finely to moderately punctate
above, often in part rugose-punctate. Sculpture of remaining tergites as described
for genalis Thoms. (p. 376 below). Female third valvulae not vomeriform.
Fore wings with basal and disco-cubital veins convergent; with areolet shortly
petiolate ; with nervus parallelus emitted in middle of brachial cell. Hind wings with
nervellus almost opposite, usually intercepted well above middle.
Hind femur varying considerably i in proportion (Figs. 41-42), from 4°5 to 5:2 times
length of its greatest breadth in Irish specimens alone. Tarsal claws not pectinate.
Length 6 to II mm.
Female. Head coloured as in genalis Thoms., but the yellow coloration less exten-
sive behind eyes, the black not more remote from lower mandibular articulation than
about length of malar space. Antennae coloured as in genalis; but with scape and
pedicellus often dull ferrugineous marginally and below, and only the membrane
conspicuously pale. Thorax coloured as in female genalis ; but the yellow coloration
generally a little less extensive around epicnemial carina and on pronotum, sometimes
much more restricted on or even absent from those parts and beneath wing-base.
Legs with the following bright flavous: hind coxae largely below, fore and mid coxae
except at base, and trochanters and trochanterelli; the hind and sometimes the mid —
trochanters above and the hind trochanterelli are dark-marked ; the coxae are other-
wise black or infuscate, or in brighter colour-forms partly rufous: femora, tibiae and
tarsi fulvous to rufous; with femora flavous at apex and hind tibiae conspicuously
stramineous at base; with the following parts infuscate: fore and mid tarsi at apex
and hind tarsi largely, hind tibiae at apex and before base, hind femora at base and
before apex, and sometimes fore and mid femora slightly before base. Wings with —
stigma and costa ochreous ang the other veins ferrugineous, more or less yellow
at base.
Colour pattern of gaster essentially of the type described for genalis, but more
diverse and irregular: frequently there is no infuscation on tergites 2-4, and in —
brightest females the petiolar segment is rufous, only a little infuscate in basal half:
in darkest females the rufous and yellow colouring is rather indefinitely suffused with —
infuscation, but in darkest males the tergites are black or infuscate from base, the
infuscation so spreading as to reduce the rufous colouring to a narrow region before
the pale yellow part.
Male. Head coloured as in male genalis Thoms., or the pale coloration further
i
THE CTENISCINI OF THE OLD WORLD 373
reduced in varying degree: in extreme case the black invades the clypeus, and the
facial pale marks are reduced to narrow orbital lines running to the upper mandibular
articulation ; but commonly the orbital marks broadened above the tentorial pits,
and a central mark remaining on the epistoma. Thorax black ; normally with tegulae
wholly pale and scutellum conspicuously pale-marked at apex, and sometimes with
metascutellum more or less obscurely pale-marked ; but sometimes the tegulae are
partly infuscate, and in darkest colour-forms the scutellar mark is absent or greatly
reduced. Coxae on the average more infuscate: in brightest forms the hind coxae are
yet broadly flavous below ; but in darkest the fore and mid coxae are only a little pale
at apex and the hind coxae not at all pale, and the rufous colouring of the hind femora
and tibiae is largely replaced by infuscation.
Redescribed from the following material: IRELAND, S. Kerry, Coumloughra Reeks,
I g, Darrynane, 1 2, Westmeath, Lough Owel, 1 3, Wicklow, Glenmacnass, I 3, I 9,
Greystones, I 9, Coan, 3 gg, Kings River, 1 3, Co. Dublin, Glenasmole, 1 g, I 9,
8.vi-15.vii, 1928-1937, A. W. Stelfox e¢ al.: SCOTLAND, Ross-shire, Loch Duich, 1 3,
29.V1.1934, O. W. Richards: ENGLAND, unlocalized, 3 ¢¢, Stephens coll., 1 g, I 9,
Desvignes coll., 1 2, 1929-1930, F. S. J. Hollick; Devon, Lustleigh, 1 g, 7.vi.1933,
_R.C. L. Perkins ; Monmouth, Trelleck Beacons, 1 4, 10.vi.1936, E. B. Britton & J. F.
Perkins: BELGiIuM, unlocalized, 11 ¢¢, 6 29, Cortenaeken, 1 g, C. Wesmael; Steen-
brugge, I g, 24.v.1875, Vivier d’Oie, I g, 29.v.1886, St. Croix, I g, 22.viii.1871, Ath,
I 9, 19.vili.1876, Severin, I 2, 10.viii.1899, J. Tosquinet: HOLLAND, Venlo, I 4, 19.vi,
I 9, 20.vi.1942, H. G. M. Teunissen: SwEDEN, Skane, Degaberga, 2 gg, 3 29,
Q-12.vii.1938, D. M. S. & J. F. Perkins: FRANCE, Central Massif, la Bourboule, 1 2,
J. de Gaulle: SwiTzERLAND, Geneva, Peney, 1 3, 4 99, 20.v.—10.vi, 1876-1889, I 6,
20.vil1.1889, Tournier ; Champel, 1 3g, 25.iv, 1 gf, 18.vi, Buess; Valais, les Haudéres,
I g, 6-27.vi.1935, R. B. Benson; Engadine National Park, 2 34,5 99, 6.vii-—6.viii,
1919-1923, Ch. Ferriére: GERMANY, Berlin dist., 1 ¢, 3 29, Ruthe coll. ; Bavaria, 1 4,
3 29, 7.vi, I Q, 8.vi, I B, 15.vi, I Y, 25.vili, I g without data, Buchecker coll. ; Erzge-
birge, Gorkau, 3 29, vii.1932, Th. Kupka; Schwerin, 1 9, 11.vi.1880, I g, 30.v.1884,
S. Brauns; Celle, 1 3, 21.vi.1942, R. Hinz: CZECHOSLOVAKIA, Bohumin, I 9, viii.1937,
Th. Kupka: Rumania, 2 $3, 1 9, M. Jaquet.
Further material seen: FINLAND, Tenala, 2 $3, v. Essen; Jaaski, 1 9, K. Valle;
Taipalsaari, 1 g, W. Hellén: SwEDEN, good series from many parts in Stockholm
_ museum ; material under cinctulus and carinatus in Lund museum: GERMANY, Upper
Bavaria, Berchtesgaden, 1 3, 30.v, I 3, 30.vi, I 9, 20.vi.1947, G. Heinrich: FRANCE,
Maisons Laffitte, Fontenay, la Bourboule, &c., J. de Gaulle; Brout-Vernet, H. de
Buysson ; Haut-Rhin, Alpes Maritimes, A. Seyrig (altogether 24 specimens in Paris
museum): AUSTRIA, Styria, Admont, Kaiserau, 1,200 m., I 9, vi.1950, G. Heinrich:
60 further specimens in British Museum (Natural History).
An inordinately variable species. To decide whether it was a single species, the
method adopted was to write a description first from the Irish material, which could
be assumed to be conspecific; and then to modify this by reference to more than
seventy other specimens, which was as diverse a collection as could be assembled.
374 A REVIEW AND A REVISION OF
VARIATIONAL TENDENCIES
A dark form is found in Ireland, the west of England and Scotland, and the high
Alps. This tends to have the clypeus rather broadly rounded at apex. An inter-
mediate has been studied from southern Sweden, Degaberga, having the same data
as the normal form; and also from La Bourboule, French Central Massif.
A brighter, stouter form is found in Holland, Belgium, Switzerland, and Bavaria.
This has the head rather less narrowed behind the eyes, and the petiolar segment
relatively less elongate.
Small specimens of the species tend to have the clypeus more transverse and the
malar space a little shorter.
Exyston cinctulus Grav. ab. approximatus, ab. n.
Differs from the normal form of cinctulus in having genal costa approaching rather
close to mandibular base but thereafter inflected (Fig. 49).
Russia: Samara (on R. Volga), 1 9, vili.1924, Lubischew (collection of A. Seyrig).
Holotype in Museum d’Histoire Naturelle, Paris.
4 |
42
Fics. 41-42. Left hind femur, in outer side view, of Exyston cinctulus Grav. These illustrate the
great intraspecific variation, and it is instructive to compare them with Figs. 16-17 (p. 327) of
two species of Anisoctenion, where a good inter-specific difference is indicated. Fig. 41 is drawn
from a female specimen from Czechoslovakia, and Fig. 42 from the type of ab. approximatus n. —
from Russia.
A second specimen has since been definitely attributed to this aberration: FINLAND,
Tammerfors, C. Lundahl: this is a fine female specimen of 11 mm. length.
At first this was thought to be a good species, closely related to cinctulus Grav. but —
having secondary differences correlated with the primary character. When, however,
cinctulus was studied intensively from a series of over eighty specimens, as diverse as
could be obtained, it was found that the supposed differences of the Russian specimen
were within or but little outside the great range of variation in cinctulus to the west
of Russia.
Other specimens were found, with the genal costa inflected but the intervening
space narrower. Yet other specimens have a mere trace of an inflected genal costa ;
and one particularly instructive specimen (in British Museum) has the genal costa —
Pen. te ie
es =
THE CTENISCINI OF THE OLD WORLD 375
split into an outer branch running almost normally as in Fig. 50 and an inner branch
inflected in the manner of the aberration. It appears that in the normal form the
genal costa, though having the condition described as continuous, continues towards
the hypostomal costa as a lamina contiguous with the genal margin backing the
mandibular articulation.
Exyston cinctulus Grav. subsp. clavatus Cresson
1864 Cteniscus clavatus Cresson, Proc. Ent. Soc. Philadelphia, 8: 284.
1897 Exyston clavatus Davis, Trans. Amer. ent. Soc. 24: 238.
1944 Exyston clavatus Townes, Mem. Amer. ent. Soc. 11 (1): 167-168.
Altogether less strongly hairy than the European form: genal costa not so strongly
elevated or hairy: yellow coloration more extensive behind eyes: antennal scapes
largely dull fulvous, with a yellowish spot beneath: flagellum also dull fulvous above in
apical half: thorax more richly yellow-marked: propodeum decidedly tending to dull
rufous (among European material I have only seen this in a few old specimens which
may have suffered fading) : coxae darkened to rufous only, not at all black: petiolar
segment scarcely infuscate at base, but there is some infuscation on other tergites,
particularly 3 and 4.
Diagnosis based on 1 9, Iosco Co., Michigan, 16-17.vii.1939, R. R. Dreisbach ; det.
H. K. Townes.
W. R. M. Mason kindly informs me that clavatus is very much less variable than
the European form: also that.it is evidently not an introduced species, for it is known
from remote parts of northern Ontario; yet it is geographically isolated, being un-
known west of the Rocky Mountains. These facts strengthen my belief that it isa true
geographical sub-species.
He has since written that he agrees with my treatment of clavatus, and has sent me
a further two females and a male for study. These substantially confirm the above
diagnosis; but the male has the hind coxae and petiolar segment pale castaneous
except at apex. He has recorded this new synonymy in the North American catalogue
(1951 April: 229).
Exyston genalis Thomson
1883 Exyston genalis Thomson, Opuscula Entomologica, fasc. 9: 883 (non Roman 1931, nec
Teunissen 1948).
Head rather strongly to strongly narrowed behind eyes; moderately emarginate
behind: on vertex, temples and inner orbits rather finely to moderately punctate ;
on genae more coarsely, irregularly and in part confluently so: epistoma coarsely
reticulate-punctate: upper face with a distinct median keel, moderately punctate,
more or less rugose on triangle between median ocellus and antennal sockets. Clypeus
rather strongly transverse, distinctly more so than in cinctulus Grav. ; rather sparsely
beset with punctures of different sizes, fine to coarse ; rather deeply delimited basally ;
more or less narrowly rounded at apex. Malar space about equal to distance between
mandibular apices in female, distinctly less in male. Hypostomal costa strongly
raised, the elevation falling gradually to the foramen magnum. Genal costa strongly
376 A REVIEW AND A REVISION OF
raised, double (Fig. 51): the outer costa almost continuous, the inner strongly in-
flected, meeting the hypostomal costa at the point where the latter is most strongly
raised ; the intervening space traversed by a series of ridges: outer and inner costae ~
crenulate ; usually both are upstanding in the female ; but sometimes in that sex, and
usually to a greater extent in the male, the inner costa is bent outward, so that the
intervening space is in varying degree narrowed.
Antennae with scape weakly excised externo-ventrally (Fig. 54) ; with postannellus
about 1-8 times length of the following segment ; with flagellum 31- to 37-segmented.
Pronotum with epomiae of moderate strength; laterally moderately punctate
above, finely punctate to rugose below. Propleura moderately, or rather finely and
closely punctate. Mesoscutum strongly and almost evenly convex; with notaulices
very superficial and not clear-cut ; more or less finely punctate medially, very finely
punctate and shining laterally. Scutellum moderately convex; rather finely to
moderately and often rather closely punctate (in the specimen from the Tatra Mts.
largely rugose) ; not margined to middle. Mesopleura moderately punctate, in part
rugose-punctate, in front ; more finely punctate behind: sternaulices quite distinctly,
though seldom strongly, impressed: mesosternum densely and very finely punctate:
epicnemial carina strongly raised, indistinct above sternaulices, medially emarginate
but not nearly interrupted: mesolcus very sharply impressed, partly closed behind.
The male differs in having the epicnemial carina only moderately raised, and the
mesolcus more or less distinctly transcostate, not sharply impressed. Propodeum
with costae rather weak but normally complete, though area basalis sometimes in-
completely delimited apically ; with area pleuralis finely and closely punctate ; else-
where more or less finely rugose.
Gaster distinctly clavate. Petiolar segment elongate and slender, 2-5 to 3-1 times
length of its greatest breadth near apex ; bearing lateral hairs distinctly longer than —
half its least breadth; weakly or moderately auriculate basally; with dorsal keels
rather sharp, distinct for quite three-quarters length of segment; almost smooth ~
between the keels, finely punctate and obsoletely rugose beside and behind them.
Tergite 2 distinctly raised on mid line at base, and impressed to either side of this
elevation; rather finely to moderately punctate and often rugulose on basal half.
Remainder of gaster more and more finely punctate. Female third valvulae small,
vomeriform. .
Fore wings with basal and disco-cubital veins strongly convergent; with areolet —
usually shortly petiolate but sometimes sub-sessile or sessile ; with nervus parallelus
emitted in middle of brachial cell. Hind wings with nervellus almost opposite, inter-
cepted a little above middle. .
Hind femur 4-1 to 5-2 times length of its greatest breadth. Tarsal claws not —
pectinate.
Length 6 to 10 mm.
Female. Head black above a line running roughly transversely around, just below
antennal sockets and foramen magnum ; below this line stramineous to flavous, the
pale coloration also extending some way up orbits of upper face: mandibles con-
colorous except at apex. Antennae with scape, pedicellus and postannellus mainly —
black, the scape almost always conspicuously pale-marked below: with annellus cas- _
THE CTENISCINI OF THE OLD WORLD 377
taneous; beyond postannellus changing to ferrugineous above, dull fulvous below.
Thorax black, with the following parts pale-coloured as on head: propleura mainly,
ventral margins of pronotum at sides (the marks sometimes interrupted), region of
epicnemial carina with backward extensions on sternaulices and beside mesolcus,
small marks below wing-base, tegulae, mark at apex of scutellum and usually another
on metascutellum. Legs rufous; with trochanters and trochanterelli, fore and mid
coxae mainly, and hind coxae nearly always below, bright flavous ; with knees rather
indefinitely pale flavous; with the following parts infuscate: hind tarsi, hind tibiae
at apex and before base, and hind femora before apex and at base: hind coxae gener-
ally in greater part black and usually rufous-marked, but sometimes in greater part
rufous. Wings with veins and stigma pale ferrugineous, more or less yellow at base.
Petiolar segment back, with a small pale mark at mid apex. Remainder of gaster
normally mainly rufous; with tergite 2 black at base and sides, and with lateral black
marks at base of tergites 3 and 4; with tergites having ill-defined but successively
broader pale yellow apical margins, which occupy most of tergites 6 and 7: the
yellow coloration may be more extensive, or the pale parts of the tergites may be to
a greater or lesser extent overspread with infuscation.
Male differs as follows. Black coloration more extensive, reaching behind eye to
a line between eye and lower articulation of mandible ; often developed at mid base of
mandible, along clypeo-frontal suture, and extending farther on to epistoma: orbits
of upper face not pale. Scape with pale mark smaller or almost undeveloped. Thorax
black, with tegulae usually wholly or partly pale, but without other pale marks. Fore
and mid coxae to a large extent black or blackish at base: hind coxae almost entirely
black. Yellow margins of tergites narrower and often more sharply defined: in one
_ specimen from Bavaria tergites 2 to 6 mainly black above, with just a trace of rufous
colouring on 3 and 4.
The male specimen from the French Ardennes has much of the female structure and
coloration. This species is illustrated on PI. 5.
Redescribed from the following material: Norway, Sognsvatnet, Aker, I Q,
31.vii.1935, I. Soot-Ryen (mus. Stockholm, recorded as E. sp. ign. by Roman, 1942:
14): SWEDEN, Dalarne, 1 3g, 16.vii, A. Roman; Dalarne, Orsa W., I 9, 5.viii.1938,
_ G, J. K.: FINLAND, Keuru, 1 4, J. Sahlberg ; Karislojo, 2 $4, 11.vi.1934, R. Krogerus ;
Ruovesi, I 3, 26.vi.1931, V. Saarinen ; Tampere, Aitolahti, 1 g, 26.vi, I 2 28.vi.1933,
_ A. Saarinen; Kuusamo, 1 4, 6.vii. 1935, R. Krogerus; Kuusamo, Ruskeakallio, 1 3,
_17.Vii.1935, R. Krogerus, 1 9, 26.vii.1935, G. J. K.: SWITZERLAND, Nyon, I 9, Cheyr:
FRAncE, Ardennes, Verdun, I 4, 10.viii.1925, Benoist: BELGIUM, Severin, I 3, 22.vi.
1899, Bastogne, I g, 23.viii.1887, J. Tosquinet: GERMANY, Schwerin, I Q, viii.1887,
S. Brauns (Nat. Mus. Ireland) ; Kéln, 1 9, vii.1945, Aerts; Goslar, I 9, 21.viii.1945,
Celle, 1 2, 26.vii.1942, R. Hinz; Berlin dist., 1 g, Ruthe coll. ; Oderberg-Liepe, 1 9,
19. viii.1938, G. J. K.; Bavaria, Erzgebirge, Gérkau, I 9, II.vi.1932, Th. Kupka:
MIDDLE Europe (unlocalized), 1 2, 4.vii.1g00, R. von Stein coll.: CZECHOSLOVAKIA,
Bohemia, Chodau, 1 4, 10.vi.1878, R. von Stein coll. ; Erzgebirge, Aussig, I 2, II.vi.
1932, Th. Kupka: PoLanp, Tatra Mts., Zakopane, 1 g, 28.vi.1932, D. Aubertin &
E. Trewavas.
Additional material seen: ITaty, Liguria, 1 ? in Spinola coll. in Turin museum:
ENTOM. II. 6 34
7
378 A REVIEW AND A REVISION OF .
GERMANY, Bodmann am Bodensee, I 4, 25.v., I g, 18.vi.1949; Bildhausen, Krs Kis-
singen, I g, 15.vili, I 9, 22.viii.1946; Holstein, Trittau, 1 g, 3.vii.1946, G. Heinrich: —
HOLLAND, Hulshorst, 1 g, 26.vi.1935, Lindemans: SWEDEN, Stockholm mus. coll.,
good series from wide range of latitudes; Thomson coll., 1 9, unlocalized, Rosen-
schiéld (possibly the type) ; 1 9, province Narcke; also 1 9 ‘Finl.’ (presumably from
Finland).
Since the above was written, I have seen the series of 30 §g, 12 9? taken in Styria,
Austria, by G. Heinrich in June-July 1950. These confirm the characters used in my
two keys to species of the genus.
Exyston phaeorrhaeus Haliday
1838 Tvyphon (subgenus Cteniscus) phaeorrhaeus Haliday, Ann. Mag. nat. Hist. (1) 2: 113 (non
Morley 1911, vec Morley 1913, nec Roman 1939).
Head rounded and strongly narrowed behind eyes ; moderately emarginate behind:
between ocelli and eyes rather coarsely and sparsely punctate; before occiput very
finely punctate above ; on temples and cheeks for the most part moderately punctate ;
on inner orbits more densely so, and on malar space more finely and much more
densely punctate: epistoma reticulate-punctate to rugose: upper face in middle more
or less trans-rugose, with a median keel that is often distinctly bifurcate before median
ocellus. Clypeus smooth in contrast with the epistoma ; sparsely beset with punctures
of different sizes, fine to coarse; rather deeply delimited basally; weakly convex;
rather narrowly to rather broadly rounded apically. Malar space at least four-fifths
distance between mandibular apices, the gena distinctly angularly raised before
mandibular articulation. Genal costa (Fig. 52) laminate, running forward to a point,
then deeply emarginate before it again runs forward to meet the hypostomal costa,
which is raised to a sharp point well forward. Outer genal costa bulging strongly
outward, seen in frontal view as in Fig. 43, the bulge forming a very distinct angle
with the normal plane of the gena. Between inner and outer costae is a cavity,
traversed by a costa running outward from below point of inner genal costa, and
usually a weaker one dorsal to this (below it in Fig. 52). The keel that, in this view,
descends from the point of the hypostomal costa runs close behind the lower articula-
tion of the mandible and appears almost to close the cavity in front.
Antennae with scape strongly excised externally (Fig. 56) ; with pedicellus strongly
rounded internally, as seen from above; with postannellus about 1-5 times length of ©
following segment; with flagellum 25- to 28-segmented.
Pronotum with distinct shining dorsal furrow; with epomiae rather strong and
sharp; laterally finely punctate above, more or less longitudinally rugose below. —
Propleura smooth and very finely punctate. Mesoscutum very strongly convex though
somewhat flattened discally; with notaulices very weak but distinct, extending
almost as far back as hind end of tegulae; finely punctate between notaulices,
sparsely and very finely punctate at sides. Scutellum rather weakly to rather strongly
convex ; not margined to middle; rather finely to moderately punctate, densely
behind. Mesopleura with longitudinal rugosities just behind those on pronotum;
otherwise moderately punctate below, sparsely and finely so above: sternaulices.
THE CTENISCINI OF THE OLD WORLD 379
shallow, punctate or rugose-punctate: mesosternum finely punctate, near mid line
rather closely, laterally rather sparsely so: mesolcus moderately deeply impressed,
broad and transcostate, closed behind: epicnemial carina strongly raised near mid
line, medially deeply incised, thus bilobed ; laterally moderately raised, disappearing
just above lower angle of pronotum.
Propodeum with costae strong ; with areae externa, basalis and superomedia shin-
ing and very finely punctate ; with area pleuralis often mainly reticulate-rugose, and
with other areae rather strongly rugose: area basalis generally delimited apically,
falling sharply forward between strongly raised keels: area posteromedia elongate,
extending forward far beyond middle.
Gaster strongly clavate. Petiolar segment 2-0 to 2-4 times length of its greatest
breadth ; moderately auriculate basally ; with dorsal keels distinct, sometimes to near
apex ; sharply raised from margins to keels, flattened, sometimes slightly concave,
between the keels ; rather obscurely punctate medially near apex, elsewhere obscurely
rugose. Tergites 2 to beyond middle and 3 at base closely beset with punctures of
moderate strength on an alutaceous background, the interspaces tending to form
longitudinal rugulosities: tergite 2 appreciably raised medially at base: thyridia
sharply outlined and conspicuous, almost semicircular. Remainder of gaster normally
finely punctate. Female third valvulae small, vomeriform.
Fore wings with basal and disco-cubital veins convergent, generally very slightly
so; with areolet varying from distinctly petiolate (as in type) to sessile ; with nervus
parallelus emitted about in middle of brachial cell. Hind wings with nervellus
slightly antefurcal, intercepted about in middle.
Tarsal claws with such stout hairs as to suggest pectination.
Length 54 to 7 mm.
Female. Head black, with the following parts dull yellowish-ivory: palpi, man-
dibles except at apex and medially at base, clypeus, lower face except for a transverse
line below antennal sockets, and genae largely. Antennae dull fulvous to ochreous
beneath, with a few basal segments mainly black ; more extensively black above, and
becoming paler more gradually towards apex ; with scape at apex above, annellus and
extreme base of post-annellus castaneous. Thorax black, with scutellum at apex,
propleura, and usually the lower angle of pronotum, yellowish-ivory ; with tegulae
_ sometimes pale castaneous behind. Legs mainly castaneous to black, with troch-
anters and femora pale at extreme apex: fore and mid legs with coxae and trochanters
mainly yellowish-ivory anteriorly and femora and tibiae mainly flavous anteriorly.
Wings with veins and stigma ferrugineous, basally paler. Gaster with petiolar seg-
ment black; with tergites 2 to 4 dull castaneous to black, with pale apical margins,
and usually largely fulvo-castaneous at sides; with tergites 5 or 6 and following
much paler, generally bright flavous: sternal region flavous to fulvous.
Male differs as follows. Head black, with palpi dull fulvous; with mandibles pale-
coloured across middle: clypeus black at sides and more or less broadly inside suture,
otherwise ivory-white, except that in type it is broadly fulvo-castaneous at apex.
Thorax without any pale marks. Legs coloured as in female, but the coxae and tro-
chanters are not pale except at extreme apex.
Redescribed from the following material: IRELAND, unlocalized, type g, A. H.
380 A REVIEW AND A REVISION OF
Haliday: ENGLAND, unlocalized, 1 3, 1907; Devon, Dartmoor, 3 $4, 22.vi.1935,
Lydford, 1 9, 20.vi.1944, R. C. L. Perkins: HoLLAND, Den Dungen, I 4, 15.v.1942,
H. G. M. Teunissen (Teunissen, 1948, under E. genalis Thoms.): SwiTZERLAND,
Geneva, Peney, 3 99, 28.v.1884, I 9, 1886, I 9, 5.vi.1889, Tournier ; Champel, 1 9,
coll. Buess; CZECHOSLOVAKIA, Moravia, Brno Hady, I 9, 13.vi.1936, A. Hoffer.
Further material seen: FRANCE, Paris district, St. Michel, O. Sichel; Haut-Rhin,
30.v.1920, A. Seyrig (4 specimens in mus. Paris): GERMANY, Schleswig, Kiel, 1 ¢,
22.v.1949, G. Heinrich: AusTRIA, Styria, Admont district, 1,200 m., on Alnus incana,
I 9, 25.vi.1950, G. Heinrich: TuRKEy-in-Europe, Istanbul, 1 3, coll. de Gaulle; this
is a peculiar specimen, with the spiracular tubercles of the petiolar segment sharp and
very prominent.
Exyston subnitidus Gravenhorst
1829 Tryphon subnitidus Gravenhorst, Ichneumonologia Europaea, 2: 144-145 (non Morley rort,
nec Morley 1936).
1906 Exyston subnitidus Pfankuch, Z. Hym. Dipt. 6: 90.
This species is closely related to, but much less strongly hairy than, phaeorrhaeus
Hal.
Head rounded and rather strongly narrowed behind eyes (decidedly less so than in
phaeorrhaeus Hal.); moderately and rather angularly emarginate behind: before
occiput finely punctate above ; on temples and cheeks for the most part moderately
punctate, sparsely so between eyes and ocelli; on inner orbits rather finely punctate
and on malar space densely so: epistoma of female rather finely punctate above to
rather coarsely so below, on an alutaceous background, the punctures for the most
part well separated; of male reticulate-punctate to rugose, the punctures not well
separated except in mid line: upper face with median keel absent or very short ; rather
coarsely punctate, the punctures for the most part well separated. Clypeus as de-
scribed for phaeorrhaeus Hal. Malar space about three-quarters distance between
mandibular apices, the gena not angularly raised before mandibular articulation.
Genal region of the type described and illustrated (Fig. 52) for phaeorrhaeus Hal.,
differing as follows: outer costa seen in frontal view as in Fig. 44, not forming a very
distinct angle with the normal plane of the gena, not bulging so strongly outward,
so that the genal cavity is less wide: inner costa running forward to a sharper point,
but incomplete internally, so that the cavity is partly open behind: hypostomal costa
more strongly and laminately raised, distinctly bent outward: the keel that arises
from point of hypostomal costa does not run very close behind mandibular articula-
tion nor appear to close the cavity in front: the keel that crosses the cavity from poms
of genal costa itself raised to a point.
Antennae with scape weakly excised externo-ventrally (Fig. 57) ; with postannellus
about 1-7 times length of the following segment ; with flagellum 25- to 27-segmented.
Pronotum as described for phaeorrhaeus Hal., but laterally moderately punctate
above. Propleura rather finely punctate. Mesoscutum rather strongly convex ; with
notaulices scarcely traceable ; medially rather finely punctate, at sides more finely
and anteriorly densely so. Scutellum strongly convex; rather finely to moderately
=x
THE CTENISCINI OF THE OLD WORLD 381
punctate ; not margined to middle. Mesopleura not much rugose anteriorly ; moder-
ately punctate above, rather coarsely so below, and closely so almost throughout ;
sternaulices broad and rather shallow and, like the mesosternum, rather finely and
closely punctate: mesolcus rather deeply and sharply impressed, closed by an incised
ridge behind: epicnemial carina much as described for phacorrhaeus Hal. Propodeum
with costae moderately strong: finely to very finely punctate above, moderately so
on area pleuralis: area basalis not falling sharply from apex: area posteromedia rugose,
_ not elongate as in phaeorrhaeus Hal. and thus not extending far beyond middle.
Gaster not strongly clavate as it is in phacorrhaeus Hal. Petiolar segment 2:0 to 2-1
times length of its greatest breadth ; moderately auriculate basally ; with dorsal keels
Fics. 43-44. Head in facial view of Fig. 43. Exyston phaeorrhaeus Hal. Fig. 44. Exyston subni-
tidus Grav.
not distinct to near apex, nor marking such an abrupt change of slope as in the
previous species ; conspicuously punctate, or in part rugose-punctate, in apical two-
thirds. Tergite 2, like those following, normally finely punctate and shining ; almost
evenly convex between the thyridia, which are not very conspicuous. Female third
valvulae weakly vomeriform.
Fore wings with basal and disco-cubital veins extraordinarily strongly convergent,
so that a continuation of the lower abscissa of the disco-cubital, from point of ramel-
lus, would meet the stigma before the point of emission of the radius; with areolet
very distinctly petiolate; with nervus parallelus emitted slightly below middle of
_ brachial cell. Hind wings and tarsal claws as described for phaeorrhaeus Hal.
Length 6 to 7 mm.
Female. Head coloured as in female phaeorrhaeus Hal., but the pale coloration
more distinctly yellowed and rather less extensive; region of tentorial pits black.
Antennae mainly black at base ; to some extent paler towards apex above; in greater
part dull fulvous below. Thorax black, with scutellum near apex, tegulae, and lower
angle of pronotum yellowish-ivory: propleura black with inner margins flavous, or
flavous with outer margins black. Gaster mainly castaneous, or with petiolar segment
and bases of following tergites blackish: tergites somewhat paler at sides. Wings
with costa yellow, with other veins and stigma more or less ferrugineous, paler at base.
Male. Head coloured as in male phaeorrhaeus Hal., but neither specimen before me
has more than a sub-triangular apical mark pale on clypeus. Thorax black with
382 A REVIEW AND A REVISION OF
tegulae more or less castaneous. Tergites 2 and following with distinct pale margins,
these becoming broader and paler towards apex, which is mainly bright flavous.
Legs coloured much as in phaeorrhaeus Hal., sex for sex; but the mid and hind
coxae and the trochanters are blackish, a little paler beneath.
Redescribed from the-following material: SwITZERLAND, Engadine national park,
Zernetz dist., I 9, 25.vii.19g21, I g, 23.vii.1922, Grupshum, 1 g, 18.vii.1925: SPAIN,
Valle de Ordesa, I , viii.1931, J. M. Dusmet.
Exyston albicinctus Gravenhorst
1820 Ichneumon albicinctus Gravenhorst, Mem. Accad. Sci. Torino, 24: 375 (non Ichneumon
albicinctus Gravenhorst 1829 = Barichneumon locutoy Thunb.).
1829 Tryphon albocinctus Gravenhorst, Ichneumonologia Europaea, 2; 204-205, syn. 0.
1829 Tryphon tricolor Gravenhorst, ibid. 2: 207-208, syn. n.
1878 Exyston cinctulus 3 var. Brischke, Schrift. naturf. Ges. Danzig, 4: 106.
1906 Exyston albocinctus Pfankuch, Z. Hym. Dipt. 6: 222.
1906 Exyston tricolor Pfankuch, ibid. 6: 223.
Head slightly narrowed or very slightly broadened behind eyes, and then strongly
rounded ; shallowly or rather shallowly emarginate behind: from level of lateral ocelli
shining, with temples rather finely punctate: genae moderately to rather coarsely
punctate, the punctures in many cases separated by less than their own diameters:
malar space and inner orbits rather finely punctate: lower face coarsely, and in large
specimens very closely punctate, the punctures somewhat squashed out of shape by
their mutual proximity: upper face very closely beset with moderate punctures, often
almost trans-rugose between antennal bases and median ocellus, and with a median
keel running to the median ocellus. Clypeus irregularly punctate, rather coarsely so —
above, rather finely and sparsely so below; scarcely convex; very superficially, and
very seldom at all sharply, delimited basally, and not set distinctly at an angle with
lower face; apically distinctly bi-tuberculate. Malar space about equal to or, more
usually, less than distance between mandibular apices in male, about one-fifth greater
than such distance in female. Genal costa moderately raised ; inflected much as in
pratorum Woldst. (Fig. 48), but more angularly so in female: the gena not angled
between costa and mandibular base, in position where in other species there is an
outer costa. Hypostomal costa, where met by genal, bent outwards and raised rather
to a point.
Antennae with scape (Fig. 55) moderately excised externally ; with postannellus
about twice length of the following segment, both measured along inner margin:
flagellum 31- to 35-segmented.
Pronotum with epomiae of moderate strength or rather weak, not sharp; laterally
moderately punctate above, closely punctate to rugose-punctate below. Propleura
moderately and closely punctate, but rather sparsely so in the small specimen from
Finland. Mesoscutum moderately and almost evenly convex; with notaulices not
clearly indicated ; finely to moderately punctate, sparsely so across disk, more closely -
so in front and behind. Scutellum rather strongly and evenly convex, though some-
times slightly buckled ; rather coarsely and closely punctate ; not distinctly margined
beyond basal furrow. Mesopleura in front coarsely and closely punctate, often in part
THE CTENISCINI OF THE OLD WORLD 383
rugose-punctate; behind more moderately and less closely punctate, but without
a distinct impunctate speculum: sternaulices broad but generally superficial: meso-
sternum densely and rather finely punctate: epicnemial carina moderately raised
between and along fore margin of sternaulices, then with margin curving inward ;
medially emarginate: mesolcus sharply impressed, often weakly transcostate, with
either a strong costa closing it behind or else a series of weaker transverse costae in
that position. Propodeum with areolation weak and incomplete in middle ; in particu-
lar, the area superomedia is not or very indistinctly bordered basally and laterally,
and the costulae are usually incomplete though always indicated: area posteromedia
well bordered and transversely rugose: area angularis longitudinally rugose: area
pleuralis moderately and rather closely punctate: area externa more finely and other
lateral areas more coarsely punctate, all less closely so: area superomedia represented
by a more or less distinctly shining patch, impunctate or sparsely punctate, often
flanked by weak transverse rugosities.
Petiolar segment of female 2-2 to 2-3 times, of male 1-9 to 2-1 times length of its
greatest breadth ; very strongly auriculate at base ; with dorsal keels distinct in about
basal third of segment, thereafter not or scarcely indicated in male, though with
clearer indications in female ; for the most part moderately punctate, but in middle
third between positions of keels shining, and not or sparsely punctate. Remainder of
gaster with tergite 2 discretely and for the most part moderately punctate; with
remaining tergites more and more finely punctate towards apex. Female third
valvulae vomeriform.
Fore wings with basal and disco-cubital veins strongly convergent ; with areolet
distinctly petiolate; with nervus parallelus emitted below middle of brachial cell.
Hind wing with nervellus almost opposite, intercepted about in middle.
Tarsal claws not pectinate.
Length 63 to 114, normally 8 to Io mm.
Female. Head black above a line running roughly transversely around, just below
antennal sockets and foramen magnum ; below this line ivory-white, much modified
by flavous or fulvous, the pale coloration also extending some way up orbits of upper
face: mandibles concolorous except at apex. Antennae with scape, pedicellus and
postannellus mainly black, the scape conspicuously pale-marked in apical half below;
with annellus mainly castaneous; with remainder of flagellum fulvous, ferrugineous
for a short distance at base beneath and a much greater distance above. Thorax
black, with the following parts pale-coloured as on head: pronotal collar, widely near
hind angles, propleura totally, tegulae, small mark below fore wings and hind margin
of mesopleura narrowly above, and large apical marks of scutellum and metascu-
tellum ; also on mesothorax a pair of marks that go broadly down lower halves of fore
margins of pleura, then narrow on upper halves of sternaulices, then greatly broaden
along fore margin of mesosternum, then continue more narrowly backward beside
mid line to closure of mesolcus. Legs with the following parts pale flavous: coxae,
trochanters and trochanterelli, fore and mid femora anteriorly, and knees; with fore
and mid femora behind, and with hind femora fulvous to castaneous; with tibiae and
tarsi flavous to fulvous, the tarsi somewhat darkened, and the hind tibiae darkened
above near apex and with a broad darker ring near base ; with dark marks on femora,
;
384 A REVIEW AND A REVISION OF
hind trochanters and mid coxae at base, and with hind coxae mainly almost black
above. Wing veins pale ferrugineous, except for the yellow costa: stigma ochreous.
Petiolar segment black, with a median apical spot coloured as on following: remainder —
of gaster bright, pale fulvo-castaneous, the tergites more and more broadly margined
apically and laterally with yellowish-white, tergites 6 and 7 being mainly so coloured,
but the colour margins are not sharply defined: tergite 2 blackish at base, with yellow
thyridia: sterna mainly yellowish.
Male differs as follows. Head for the most part black, including a very broad band
inside basal margin of clypeus; with pale colour on remainder of clypeus, on middle
and along basal margins of mandibles, and on maxillary and labial palps. Antennal
scape without a pale mark beneath, or with an inconspicuous mark that does not
reach apex of segment. Thorax black; with a small pale apical mark on scutellum,
and sometimes an obscurer one on metascutellum: tegulae partly ferrugineous. Legs —
coloured as in female, but with fulvous coloration darkened ; with coxae wholly or for —
the most part, trochanters usually in part and hind femora behind, black or almost
so. Gaster coloured as in female, but usually tergites 3 to 5 are blackish at base and —
tergite 2 more than half black: the palest coloration is always less extensive and often
more yellow.
Redescribed from the following material: BELGIuM, Steenbrugge, I g, 24.v.1875,
Ghlin, 5 $6, 17.vi.1877, Mont 1|’Enclus, 1 3g, 26.v.1893, J. Tosquinet: HoLLanp,
Exaeten, i ¢, vi.1884, I g, 5.vi.1887, A. Wasmann: GERMANY, Schwerin, I 4,
II.vi.1889, S. Brauns: SWITZERLAND, Geneva, Peney, I 9, 29.v.1876, I g, 5.vi.1889, |
I g, 20.vi.1889, Tournier ; Champel, 1 4, coll. Buess; without data, 1 2 : HuNGarRy,
Jaszberény, 1 2, Moczar (mus. Stockholm): FinLanp, Abo district, Uskela, 1 3,
E. J. Bonsdorff, coll. Maklin: Europe, unlocalized, 1 g bought from Staudinger
(Brit. Mus.).
Additional material seen: FRANCE, Paris district, Maisons-Laffitte, Rueil, Nantesse;
Varennes, St. Baume, J. de Gaulle (4 specimens in mus. Paris): GERMANY, Crefeld,
I 9, vi.1928, A. Ulbricht.
Originally described from Piedmont, Italy, and the female synonym from the
Rhineland. Brischke’s insect was from the Danzig district: from his diagnosis I have
no doubt that it belonged to this species. Evidently a very scarce species: Morley has
already, quite rightly, rejected Stephens’s British records.
Exyston sp.
1931 Exyston genalis Roman, Ark. Zool. 28A (6): 14 (non Thomson 1883).
It is thought desirable to correct an existing record, but inopportune to validate _
this species with a name based on a single imperfect male. In order, however, to
stimulate the search for further specimens, and to increase our understanding of the
genus as a whole, the species has been separated in my key to palaearctic species, and
is here further diagnosed.
A larger, more elongate species than pratorwm Woldst.: length of unique male
ro mm. Upper face reticulate-punctate, not rugose. Clypeus sharply delimited
basally. Malar space a little less than distance between mandibular apices. Post-
ye
THE CTENISCINI OF THE OLD WORLD 385
annellus 1-7 times length of following segment. Pronotum laterally moderately punc-
tate above. Notaulices short and extremely superficial. Mesopleura coarsely punctate
in front. Propodeum with costae normally complete and rather strong. Petiolar
segment about 2-5 times length of its greatest breadth; weakly auriculate basally ;
with dorsal keels strong and almost percurrent. Tergite 2 reticulate-rugose in basal
half, thereafter rugose-punctate. Fore wings with basal and disco-cubital strongly
convergent. Tarsal claws not pectinate.
So closely related to the nearctic E. boreotis Davis (of which I have seen two pairs
determined by H. K. Townes) that I thought it might be a form of that species.
In the American specimens, however, the temples are finely or rather finely punctate ;
whereas in the Siberian they bear coarse punctures separated by less than their own
diameters. I have been allowed to submit the specimen to W. R. M. Mason, who is of
the opinion that it is not boreotis Davis but that, like that species, it belongs to the
group of variatus Prov.
Exyston pratorum Woldstedt
1858 Exenterus triangulatorius Holmgren, Svensk Vet.-Akad. Handl. 1: 235 (non Gravenhorst
1829).
1874 Exenterus pratorum Woldstedt, Bidr. Kann. Finl. Natur Folk, 21: 52-53.
_ 1877 Cteniscus (Tricamptus) pratorum Woldstedt, Bull. Acad. Sci. St. Pétersbourg, 28: 454.
_ 1883 Exyston brevipetiolatus Thomson, Opuscula Entomologica, fasc. 9: 883 (synonym by Hellén
1937).
_ 1911 Exyston brevipetiolatum Morley, The Ichneumons of Great Britain, 4: 192-193.
_ i911 Exenterus phaeorrhoeus Morley, ibid. 4: 207 (non Haliday 1838).
1911 Exyston triangulatorius Schmiedeknecht, Opuscula Ichneumonologica, §: 2300-2301.
y 1937 Exyston pratorum Hellén, Notul. Ent. 17: 124.
——
1939 Exyston phaeorrhaeus Roman, Ent. Tidskr. 60: 195 (non Haliday 1838).
Head slightly to moderately narrowed behind eyes and strongly rounded ; rather
shallowly to moderately emarginate behind: vertical region and temples very finely
punctate, genae finely to moderately so; upper face finely to moderately and closely
_ punctate, sometimes transrugulose just above antennae; epistoma rather coarsely
punctate, sometimes reticulate in part. Clypeus for the most part rather coarse-
ly punctate ; sharply and rather deeply delimited basally ; more or less weakly convex ;
often quite broadly rounded at apex. Malar space of female a little less than distance
_ between mandibular apices ; of male much shorter, about equal to width of annellus.
Genal costa moderately raised, strongly but seldom angularly inflected (Fig. 48),
often bearing short carinae in front: hypostomal costa strongly and roundedly raised,
bent slightly outward. The gena is roundedly angulate between costa and mandibular
base, in the position in which in other species there is an outer costa (see again Fig. 48):
in male this angulation is carried farther back (dorsal), is much sharper near mandi-
bular base, and bounds an intervening area that is more distinctly excavate behind
articulation of mandible and is often distinctly though imperfectly transcarinate.
Antennae strongly hairy ; with scape weakly excised externo-ventrally (Fig. 58) ;
with postannellus generally less than 1-5 times length of following segment; with
flagellum of female 23- to 26-, of male 22- to 25-segmented.
Pronotum with epomiae rather strong and sharp; laterally very finely punctate
ENTOM. It. 6 3B
386 A REVIEW AND A REVISION OF
above, rugose in part below. Propleura finely punctate. Mesoscutum very strongly
convex anteriorly, flattened discally; with notaulices shallowly impressed, but
extending to beyond middle; more or less finely punctate, often with some faint —
rugulosity. Scutellum weakly to moderately convex ; not margined to middle ; gener-
ally finely punctate above, closely punctate or rugose-punctate behind, longitudinally
rugose at sides. Mesopleura each with a large, almost impunctate area above; finely
to moderately punctate below, moderately so above: sternaulices broad and moder-
ately impressed: mesosternum very finely punctate: epicnemial carina moderately
45 46 47
Fics. 45-47. The very variable petiolar segment of Exyston pratorum Woldst. Fig. 45 drawn
from a male from Orsa, middle Sweden, resembling the type and illustrating the applicability of
Forster’s diagnosis of Tvicamptus to this species. Fig. 46 drawn from a normal female from
Paanajarvi, NE. Finland, and Fig. 47 from a male from the New Forest in southern England,
illustrating the other extreme condition.
raised between and along fore margin of sternaulices, then bent upward on to meso-
pleura, medially almost completely interrupted: mesolcus moderately to strongly
impressed, broadening posteriorly, closed behind. Propodeum with costae of moder-
ate strength; mainly more or less finely rugose, but with some upper areae shining”
and very finely punctate, the area pleuralis generally discretely punctate in front,
and the area posteromedia finely reticulate-rugose: area superomedia not distinctly
delimited basally.
Petiolar segment (Figs. 45-47) short for an Exyston, 1-3 to 1-7 times length of its
greatest breadth ; dorsally reticulate-rugose, sometimes finely so; not strongly auricu-
late basally ; with dorsal keels broad and flattened at base, thereafter not very strong
though sometimes almost percurrent: spiracular tubercles undeveloped or more or
less prominent. Remainder of gaster moderately punctate, finely so towards apex;
with tergites 2, generally to beyond middle, and 3 at base longitudinally finely rugose.
Female third valvulae small, vomeriform.
Fore wings with basal and disco-cubital veins almost parallel; with areolet sessile
or sub-sessile ; with nervus parallelus emitted below middle of brachial cell. Hind
wings with nervellus strongly antefurcal, intercepted below middle.
Tarsal claws finely pectinate.
Length 5 to 74 mm.
Female. Head black, with the following parts ivory-white, usually in part suffused
THE CTENISCINI OF THE OLD WORLD 387
with flavous and margined with fulvous: palpi, mandibles except at apex, clypeus,
genae broadly, and lower face largely. The pale colour on the gena usually extends
beyond the angulation and on to the hypostomal costa ; it extends broadly more than
half-way up the sides of the lower face, which also bears a rather heart-shaped mark
almost covering the epistoma: the pale marks on the lower face may be so largely
confluent as to confine the lower black coloration to the tentorial pits. Antennae with
basal segments mainly black, often castaneous at apex and sometimes more exten-
sively below: beyond postannellus ferrugineous above, often paler towards base, dull
fulvous below, often darker at apex. Thorax black, with the following parts pale-
coloured as on head: propleura largely, tegulae, and scutellum and metascutellum at
apex ; usually lower angles of pronotum and often an extension along ventral margin,
and generally a pair of marks on anterior margin of mesopleura; occasionally also
marks on epicnemia, below bases of fore wings and down posterior margins of meso-
pleura. Fore and mid legs with coxae, trochanters and trochanterelli similarly pale-
coloured, the trochanters dark-marked above, the coxae darkened usually only at
base but sometimes more extensively ; with femora fulvo-castaneous and with knees,
tibiae, and tarsi fulvous to flavous: hind legs with coxae, trochanters, and trochan-
terelli black or almost so, pale at apex and on trochanterelli beneath ; with femora,
tibiae, and tarsi ferrugineous, the femora and tibiae more or less dull castaneous in
middle or almost entirely, the tibiae sometimes much paler than the femora, and the
knees conspicuously stramineous-white. Wing veins and stigma ferrugineous to fus-
cous, basally paler. Petiolar segment black. Remainder of gaster normally having
tergite 2 black at base and sides, and 3 with isolated black marks at mid base and at
anterior angles ; with tergite 2 mainly rufocastaneous, the following similar but succes-
sively duller; with tergites having pale margins, yellowish to ivory, broader on
successive segments to apex, covering most of tergite 6. In some specimens the black
coloration is much more extensive, covering all of tergite 2 but the apical margin and
thyridia ; and in some the pale apical margins are much less pronounced.
Male differs as follows. Head without the pale marks on genae and sides of lower
face, with heart-shaped mark on epistoma smaller, and with clypeus narrowly black
along sutural margin. Thorax black; with tegulae generally pale ; sometimes (as in
type) with a transverse pale mark near apex of scutellum. Mid, and generally fore,
coxae dark except at extreme apex. Gaster sometimes mainly black dorsally ; pale
margins of segments, on the average, narrower and sharper.
Redescribed from the following material: IRELAND, S. Tipperary, Ballinacourty,
I 9, W. Cork, Glengarriff, 1 2, Waterford, Dungarvan, I 9, Queens Co., Portarlington,
I $, Woodbrook, 1 9, N. of Emo Park, 2 99, Wicklow, Clara, 3 gd (one seen by
A. Roman, see Roman, 1939), 1 2, Arklow dist.,1 3, Ballinclea, 1 3, Deputy’s Pass, 1 9,
Athdown, 2 929, 4.vi-26.vi and 26.vii—16.viii.1925-1945, A. W. Stelfox: SCOTLAND,
Argyll, Puck’s Glen, I 3, 9.vi.1925, J. J. F. X. King; Inverness-shire, Aviemore, I 4,
26.vi.1934, R. B. & J. E. Benson: ENGLAND, Devon, Cornworthy, 1 9, T. A. Marshall
coll.; Kingsteignton, 2 ¢¢, 1 9, Hilber, 1 3, Lustleigh, 1 g, Newton Abbot, 1 2,
3-20.Vi., 1933-1942, J. F. Perkins ; Hants, New Forest, 1 g, 26.v.1912, G. T. Lyle, 2 29,
17-24.vi.1923, H. Scott; New Forest, Lyndhurst, 1 9, 18.vi.1907, C. Morley (see
above, p. 370) ; Surrey, Oxshott, 1 3, 14.viii.1890, T. A. Marshall coll. ; Abinger, 1 9,
388 A REVIEW AND A REVISION OF
28.viii.1947, G. J. K.; Herts., Brickett Wood, 4 33, I 2, 26.v.-9.vi., 1936-1937, R. B.
Benson ; Lancs. mid, Carnforth, Leighton Moss, I 9, 5.vi.1941, H. W. Miles; West-
morland, Foulshaw Moss, 1 g, I 9, 8.vi.1941, G. J. K.: BeLcrum, unlocalized, x ¢,
4 22, C. Wesmael ; Vivier d’Oie, 1 9, 18.v.1883, I 9, 2.vi.1881, Orval, 1 9, 6.viii.1892,
Stockel, 1 9, 13.viii.1890, J. Tosquinet: SwEDEN, Dalarne, Orsa, 1 g, 4.viii.1938,
G. J. K.: FINLAND, Ladoga-Karelia, Sortavala, 1 3, 27.vi, Woldstedt (type) ; Kuusamo,
Liikasenvaara, I 9, 3.viii.1935, Paanajarvi, Ruskeakallio, 1 2, 17.vii.1935, G. J. K.:
FRANCE, Paris district, Claman, I 9, 5.vi.1892, Mesnil-le-Roi, 1 9, J. de Gaulle:
SWITZERLAND, Buchholz, I 9, 12.vili.1g07, ? captor; Berisal, 1 9, vii.1gog, J. de
Gaulle; Valais, les Haudéres, 1 3, I 9, 6-27.vi.1935, Ferpécle, I g, 21-27.vi.1935,
R. B. & J. E. Benson: Germany, Eisenberg, 1 2, Th. Kupka: CZECHOSLOVAKIA,
Aussig, I 3, 20.V.1935, I 9, 19.vi.1932, Th. Kupka.
Further material seen: SWEDEN, material in Stockholm and Lund museums, includ-
ing Thomson’s type series: FRANCE, Mesnil-le-Roi, Maisons-Laffitte, Sévres, J. de
Gaulle, Vendresse, A. Seyrig, 8 specimens in Paris museum: GERMANY, Crefeld,
1 specimen, A. Ulbricht (Paris museum); Bodmann am Bodensee, I 3, 26.v.1949,
G. Heinrich; Helzen, 1 g, 17.v.1943, Stolzenau, I 9, 3.vi.1945, Géttingen, 1 9,
13.ix.1949, R. Hinz: Austria, Styria, Admont district, 1 g, 1 2, Io.vi.1950, I Q,
3.vil.1950, G. Heinrich.
The last-mentioned female specimen has the gena like that of the male, and has
comparatively small yellow marks on the propleura.
This is a more than ordinarily variable species ; yet, after examining all the above
diverse material, I am satisfied that it does belong to a single species. The male —
specimen from Orsa, middle Sweden, has been presented to the British Museum
collection: it agrees very well with the type, except that it has the petiolar segment
more distinctly auriculate at base.
Fig. 45, of the petiolar segment, demonstrates that this species does not disagree —
with Forster’s diagnosis of Tvicamptus, so that Woldstedt’s selection is valid whether —
the rules require agreement with the diagnosis or not. Tricamptus falls, therefore, as
a synonym of Exyston Schigdte. This new synonymy has been published by Mason —
(r951, April: 230)
Time of appearance. Ulbricht (1926: 21) records that his dates of capture of this
species were in May and the beginning of June, and then again in August. The dates —
of capture I have been able to record lead me to believe that there are two genera-
tions a year in most districts; for they range from 17.v to 3.vii and from 26.vii to
13.ix. The only exceptions are a single unspecified July date for a Swiss capture ; and
a mid- July date from NE. Finland, where the climate is such that one would expect
a species normally having two generations a year to have only one.
Exyston ‘triangulatorius Hlgr. var. melanurus’ Ulbricht -
Ulbricht’s variety melanurus was based on a specimen or specimens he had
captured, and which he attributed to the last of the three varieties brought forward
by Schmiedeknecht (1911: 2301). He gave as description ‘Hinterleibspitze schwarz
gefleckt’, within inverted commas; but this was not an exact quotation of Schmiede- —
knecht, whose words were ‘Var. 3.g. Die hinteren Segmente schwarzlich gefleckt.’
THE CTENISCINI OF THE OLD WORLD 389
Moreover, the varieties presented by Schmiedeknecht were not always original, but
‘may often be traced to an earlier author. These three varieties may be traced to
Holmgren (1858: 235), whose actual relevant words were ‘var. 3 ¢: abdomine nigro-
notato’.
Through the kindness of Herr B. Maixner I have been able to see the Ulbricht
material of Exyston in the Crefeld museum. All specimens standing under this species
are correctly determined to species. Three stand under the species name, unqualified,
and others under three varietal numbers. Two of these segregates may be considered
to correspond with varieties mentioned by Schmiedeknecht, if numbers be transposed
and the sexes reversed [actually Schmiedeknecht’s var. 1 is the usual condition for
the male, but I have never seen a female to correspond]. But the specimens under
Ulbricht’s remaining number, his var. 2, are not mutually consistent in colour pattern,
not even approximately so. One of the specimens not arranged as a variety is labelled
‘var. 2 Higr.’ So I feel able to select as lectotype of melanurus Ulbricht another speci-
men not arranged as a variety. This is a small male specimen having a small
blackish spot on tergite 6, to the right of the midline, and also correspondingly at
extreme base of tergite 7. This mark is clearly individual and of no real varietal
significance. The only other specimen that might have served is a male arranged
under var. 2, having tergite 5 mainly infuscate above but not tergite 3. No doubt it
Was a specimen marked in some such way as that that Holmgren placed as his var. 3.
KEY TO PALAEARCTIC SPECIES OF EXYSTON SCHI@DTE
/ 1. Genal costa single and not remote from mandibular base (Figs. 49, 50): female
third valvulae not vomeriform: scape deeply excised externally (Fig. 53). 2
Genal costa double, or single and remote from mandibular base: female third
valvulae vomeriform: scape not deeply excised externally except in phacorrhaeus
Hal. (easily distinguished in facial view, Fig. 43) . : ‘ ; : 3
2. Genal costa continuous or almost so (Fig. 50). ; ‘ . cinctulus Grav.
Genal costa inflected, though not remote from mandibular base (Fig. 49)
cinctulus Grav. ab. approximatus nov.
3. Genal costa double, the outer costa continuous or almost so, the inner costa
inflected ‘ F ; ; ; 4
Genal costa single and ‘strongly inflected (e.g. Fig. 48) 4 F , ; 6
4. Genal costae crenulate, rather regularly curved (Fig. 51), the intervening space
traversed by a series of ridges: petiolar segment elongate and slender, at least
2°5 times length of its greatest breadth near apex: flagellum more than
_-30-segmented . : . genalis Thoms.
Inner genal costa very strongly sinuate and pointed (e. g. Fig. 52): petiolar segment
not 2°5 times length of its greatest breadth: flagellum less than 30-segmented 5
5. A very strongly hairy species, with gaster strongly clavate: gena (Figs. 43, 52)
as in detailed description (p. 378): antennal scape strongly excised externally
(Fig. 56): tergite 2 to beyond middle closely beset with moderate punctures
and not shining: fore wings with basal and disco-cubital not extraordinarily
strongly convergent : ; ' ; : : phaeorrhaeus Hal.
Fics. 48-52. Head in hind view of Fig. 48. Exyston pratorum Woldst. Fig. 49. Exyston cinctulus
Grav. ab. approximatus nov. (lower tooth of right mandible malformed). Fig. 50. Exyston cinctulus
Grav., normal. Fig. 51. Exyston genalis Thoms. Fig. 52. Exyston phaeorrhaeus Hal.
Fig. 48 drawn freehand.
THE CTENISCINI OF THE OLD WORLD 391
A much less strongly hairy species, with gaster scarcely clavate: gena (Fig. 44)
differing as detailed in description (p. 380): antennal scape weakly excised
externo-ventrally (Fig. 57): tergite 2 finely and rather sparsely punctate and
shining: fore wings with basal and disco-cubital oo ee strongly con-
vergent . : ; subnitidus Grav.
6. Clypeus very snfecticiatly delimited basally énd not set distinctly at an angle with
lower face: propodeum with areolation weak and incomplete in middle ; in par-
ticular, the area superomedia is not or very indistinctly bordered basally and
laterally: tergite 2 discretely punctate and shining. albicinctus Grav.
=. 3 he sims 4 . a)
cide. ge". albic.
ows
Subn,
_ Fics. 53-58. Left antennal scape, &c., in sinistro-lateral view, of Fig. 53. Exyston cinctulus Grav.
Fig. 54. Exyston genalis Thoms. Fig. 55. Exyston albicinctus Grav. Fig. 56. Exyston phaeorrhaeus
Hal. Fig. 57. Exyston subnitidus Grav. Fig. 58. Exyston pratorum Woldst.
Clypeus quite deeply and sharply delimited and set distinctly at an angle with
lower face: propodeum with areolation normally complete and moderately
strong: tergite 2 at least finely rugose in about basal half. . : 7
7. A larger, more elongate species, with length of unique male 10 mm.: antennae
normally hairy: pronotum laterally moderately punctate above and mesopleura
coarsely punctate in front: petiolar segment about 24 times length of its greatest
breadth: fore wings with basal and disco-cubital strongly convergent: Kam-
tchatka, R. Malaise (genalis Roman 1931 non Thomson) : > Se:
A moderate-sized species, not more than about 7 mm. in length: antennae very
strongly hairy: pronotum laterally very finely punctate above and mesopleura
no more than moderately punctate: petiolar segment about 14 times length of
its greatest breadth: fore wings with basal and disco-cubital almost parallel:
a rather common European species . ; : ‘ pratorum Woldst.
ALTERNATIVE KEY TO EUROPEAN SPECIES OF ExyYSTON SCHIGDTE
In the foregoing key a natural arrangement has been attempted, and the fullest use
made of diagnostic characters. The following may serve for the readier separation of
species known from Europe, especially where few specimens are available, and the
hinder head and female valvulae cannot be clearly observed:
1. Antennal flagellum almost always more than 30-segmented ‘ j : 2
Antennal flagellum less than 29-segmented . ‘ ‘ ‘ ‘ , 4
392 A REVIEW AND A REVISION OF
2. Clypeus very superficially delimited basally: areolation of propodeum decidedly
incomplete in middle: petiolar segment not 2-5 times length of its greatest
breadth . ; albicinctus Grav.
Clypeus quite deeply and sharply delimited basally: areolation of propodeum nor-
mally complete: petiolar segment often more than 2-5 times length of its
greatest breadth near apex (e.g. Pl. 5) , , ; ; ‘ . 3
3. Antennal scape deeply excised externally (Fig. 53): hind trochanter dark- or dull-
marked above; scutellum pale-marked at apex except in the darkest males
; ; , : : ; : ‘ ; ; cinctulus Grav.
Antennal scape weakly excised externo-ventrally (Fig. 54): female hind trochanter
usually, and male scutellum immaculate . : ; . genalis Thoms.
4. Antennal scape strongly excised externally: gena strongly angularly raised, bulg- —
ing outward as seen in facial view in Fig. 43: gaster strongly clavate
: phaeorrhaeus Hal.
Antennal’ scape not strongly excised : gena not thus raised: gaster not strongly
clavate . ; ‘ ‘ - ; . : ‘ ; ; . 5
5, Antennae normally hairy: tergite 2 sparsely and finely punctate, not rugose: fore —
wings with basal and disco-cubital piste Ts strongly convergent: hind
tibiae not whitish at base ; : subnitidus Grav.
Antennae very strongly hairy: tergite 2 finely longitudinally rugose to beyond
middle: fore wings with basal and disco-cubital almost parallel: hind tibiae con-
spicuously stramineous-white at base. ‘ , . pratorum Woldst.
Genus PAREXYSTON, gen. n.
Genotype Exyston calcaratus Thoms. (monobasic).
Reference to this new genus is made in the discussion of genera (pp. 313-314), and
the characters are defined in the key to genera (pp. 316-319). The single described
species has previously been placed in Exyston ; but I think it is more closely related —
to Smicroplectrus, and particularly to S. clémenti sp.n. (p. 404).
Parexyston calcaratus Thomson
When studying the series in the Thomson collection in 1938 I noted the following
characters additional to those given by Thomson (1883): head shining; narrowed
behind eyes but not much rounded; very broadly emarginate behind. Cheeks very
long. Hypostomal and genal costae strongly elevated: the genal costa is not double,
but the cheek is angulate therefrom to the lower articulation of the mandible.
Antennae incrassate almost to apex, with apical segment pointed. Thorax shining:
A
epomiae very long and strong: notaulices very strong: scutellum margined to beyond ~
middle, but not sharply: epicnemial carina strongly raised: sternaulices broad: mesol-
cus deeply impressed, open behind. Propodeum dorsally rugose, shining on area
pleuralis ; with costae strong: area superomedia sub-hexagonal, emitting costulae in
middle: area petiolaris extending to middle. Petiolar segment rugose; with basal
lobes ; with dorsal keels not strong but almost percurrent. Tergite 2 with sculpture
= ' -—? i a” Se
a ey ~ eet — wa Lhe
THE CTENISCINI OF THE OLD WORLD 39
basally rugose, apically finer; thyridia large, about half breadth of space between
them. Remaining tergites shining.
Description from 2 29, 4 ¢¢ in Thomson collection.
Since that date, the following material has been examined: IRELAND, Co. Wicklow,
Monagloch, 1 9, 8.vi.1926, A. W. Stelfox: ENGLAND, Devon, Kingsteignton, Starks
Ponds, I 3, 2 29, 29.v—I.vi.1942, Highweek, 1 g, I 9, 22.v.1942, Newton Abbot, 1 g,
vi.1940, J. F. Perkins; Notts., Teversall, 1 2, 26.vi.1913, J. W. Carr (recorded as
brevipetiolatus Thoms.) ; Suffolk, Monks Soham, flying at blackthorn, 2 29, 23.v.1920,
C. Morley (recorded as subnitidum Grav.): HOLLAND, Oisterwijk, 1 9, 18.vi.1943,
oo
Fic. 59. Head in hind view of Parexyston calcavatus Thoms.
H. G. M. Teunissen: SWEDEN, Skane, HG6r district, 1 3, 28.vi.1938, D. M. S. & J. F.
Perkins: FINLAND, Keuru, 1 2, W. Hellén (recorded as pratorum ab. nigriceps Hellén) ;
Hattula, 1 3, L. v. Essen: GERMANY, Berlin district, 1 2, Ruthe coll.
Of the above, the Ruthe collection specimen is 6} mm. in length, and has the thorax
and propodeum largely rufous, as in the syntypes. With one exception, the remaining
specimens are of a much smaller form, 4 to 44 mm. in length, and have the ground
colour of the thorax and propodeum black or blackish. This form looks so distinct
that I would have thought it merited varietal status ; but the specimen from Holland
is an intermediate, having a length of 54 mm. and the thorax notably paler than in
_ this form. The large rufous and the small dark forms are not or not widely geo-
‘ graphically separated, for both have been taken in the southern Swedish province
_ of Skane.
The second segment of the maxillary palps is strongly swollen in this species.
The earlier British record of this species (Carr, 1924: 53) was based on three
specimens of Smicroplectrus quinquecinctus Grav.
Genus SMICROPLECTRUS Thomson
1883 Smicroplectyrus Thomson, Opuscula Entomologica, fasc. 9: 881-882, 888.
If our knowledge of the genus Exyston in Europe has been in an unsatisfactory con-
dition, our knowledge of European Smicroplectrus has been quite rudimentary.
ENTOM. II. 6 a
394 A REVIEW AND A REVISION OF
Though there are other good characters available for the separation of some of the
species, it is the form of the hinder gena that is of prime importance, and this has
never before been properly studied. Roman (1913: 128), from the study of a single
female specimen in the Stockholm museum, discovered that in S. guinquecinctus Grav.
there is a double genal costa, the inner costa being sharply raised and inflected: in
all other forms known to him he found no outer costa. He failed to observe the differ-
ence in the form of the gena of jucundus Hlgr. from that in bohemani Higr., and con-
sidered that there was no more than one species described under bohemant, jucundus,
and costulatus.
After prolonged study, I recognize eight European species ; and of these, only one
remains a unique type, and this one is very distinct. All seven others are represented
in my material by series of over twenty specimens. It is hoped that this revision will
provide a sound basis for the study of at least the European species: I now find I
can place well over go per cent. of the specimens confronting me. Doubt of the
identity of some of the males, especially smaller ones, is no cause for wonder ; but it
is vexing as well as perplexing to be still unable to place four female specimens. One
of these specimens, however, looks like an intermediate between two of the recog-
nized species. Since Smicroplectrus species are, so far as is known, almost all attached
to Nematine sawflies on catkin-bearing plants, predominantly Salix, it seems quite
possible that hybridization would sometimes occur.
The difficulty of discriminating the species was greatly increased by the existence
of small, pale forms, which I recognize in five species, of which the majority of speci- —
mens are larger and darker. These forms are of smaller size and have fewer antennal
segments than normal individuals: the general darker body colour is altogether paler,
though the pale colour may be brighter ; the punctation is weaker, though the grosser
sculpture may be more outstanding. Whether these differences are due to the use of —
different hosts, to nutritional differences in individuals, or to rate of development at
a particular time of year is not known. They do, however, intergrade with the normal,
and cannot even be regarded as distinct varieties.
Apart from these small, pale forms, which do not seem previously to have been ~
noted, the European species are very uniform in general colour pattern. Thus it has
happened that species have not been described on concomitant colour characters, as
happened in most Ichneumonid genera, and several are now new. For the eight
European species recognized, only five names are available ; and I place two of these
as synonyms, having examined all the types concerned. The colour characters that
have been used, such as the colour of the hind coxae, and the presence or absence of
a pale apical mark on the scutellum, are found to break down; and are unlikely to do ~
more than provide negative evidence in some cases and within defined regions. The
degree of development of the costula is quite unreliable for separating the species.
The type species, S. qguinquecinctus Grav., is also regarded as the typical species,
and others are to a large extent described in comparison with it.
Tryphon incisus Grav. was placed in Exenterus by Pfankuch (1906), but transferred
to this genus by Schmiedeknecht (1911) on account of the mention of a relatively long
hind tibial spur. I have seen the single female specimen in the Gravenhorst collection,
and can state that the hind tibial spur is about two-fifths the length of the metatarsus, __
4
THE CTENISCINI OF THE OLD WORLD 395
and that the mid tibia bears two spurs. The species can, therefore, be excluded not
only from this genus but from the tribe Cteniscini. I did not, at the time, identify it
positively ; but, from my recollection of it, I suppose it to belong to the Polyblastina.
Smicroplectrus quinquecinctus Gravenhorst
1820 Ichneumon quinquecinctus Gravenhorst, Mem. Accad. Sci. Torino, 24: 374.
1829 Tryphon quinquecinctus Gravenhorst, Ichneumonologia Europaea, 2: 165-166.
1858 Exenterus erosus Holmgren, Svensk Vet.-Akad. Handl. 1: 227.
1906 Smicroplecirus quinquecinctus Pfankuch, Z. Hym. Dipt. 6: 94.
Head (Figs. 63, 64) not strongly transverse ; moderately, sometimes rather strongly,
narrowed behind eyes in female and strongly so in male, slightly rounded ; moderately
emarginate behind: with vertical region and orbits finely and sparsely punctate;
before occiput and on temples and cheeks moderately so; cheeks below and upper
face much more coarsely and densely punctate, and epistoma still more coarsely and
densely so, bearing punctures separated by much less than their own diameters:
sometimes the sculpture is rugose on mid line above and below antennal bases.
Clypeus sparsely and rather finely punctate below, more coarsely and densely so
above ; rather deeply delimited, with tentorial pits conspicuous ; not strongly convex
basally, and rather shallowly impressed before the narrowly rounded apex. Malar
space longer than distance between mandibular apices in female, about as long in
male. Genal costa strongly laminately elevated, not or scarcely angularly inflected:
gena with an outer costa elevated at least part way back from mandibular base,
thence completed as an angulation running togenal costa. Hypostomal costa strongly
raised, evenly rounded below junction of genal costa. Genal pit enclosed by these
costae distinctly broader along genal costa than its length along hypostomal (Fig. 69).
Antennae with scape (Fig. 60) weakly excised externo-ventrally: flagellum 26- to
32-segmented, moderately explanate in middle.
Pronotum with dorsal transverse furrow shining; with epomiae sharp and of
moderate strength ; laterally strongly punctate above, finely rugulose-punctate below.
Propleura strongly punctate. Mesoscutum strongly convex anteriorly ; with notau-
lices strongly impressed anteriorly, running far beyond middle and falling into a
median depression; rather finely to moderately punctate, much more densely so
anteriorly between notaulices. Scutellum strongly convex ; finely and sparsely punc-
tate above, more coarsely and densely so at sides and behind; margined usually to
about middle but sometimes well beyond; more or less longitudinally impressed on
disk. Mesopleura rather coarsely to coarsely punctate, rather closely below, more
moderately so above, and with a shining speculum: sternaulices broad, rather
shallowly impressed: mesosternum finely and densely punctate, prepectus strongly
so: epicnemial carina dorsally incomplete, distinctly elevated in female and slightly
so in male: mesolcus narrow but deeply impressed, a little broadened posteriorly,
_ closed behind. Propodeum with costae moderately strong, sometimes strong; with
dentiparal areae weakly produced and with costulae complete or incomplete ; finely
rugose to rugose-punctate, with the area pleuralis moderately and discretely punctate,
396 A REVIEW AND A REVISION OF
and the area posteromedia more or less rugose, generally transversely so: area basalis
not delimited apically: area superomedia more or less explanate to near apex, moder-
ately rounded behind.
Gaster on tergites I and 2 mainly, and 3 at least at base, coriaceo-rugose ; elsewhere
mainly alutaceo-punctate, generally strongly so, giving the surface a decidedly matt
appearance; with the hinder segments smoother and more shining towards apex.
Petiolar segment (Fig. 74) 1-4 to 1-8, often less than 1-6 times length of its greatest
breadth ; strongly margined, with basal lobes pointed, sometimes acutely so; with
dorsal keels rather strong and almost percurrent. Tergite 2 with diagonal basal .
impressions fairly distinct but ill-defined.
Fore wings with stigma and radial cell of normal, rather elongate shape: stigma —
about 4:0 times (varying 3:6 to 4:3), radial cell about 2:8 times as long as broad ; with
basal and disco-cubital veins nearly parallel ; with areolet rather small and sub-sessile —
or shortly petiolate ; with nervus parallelus emitted in middle of branchial cell. Hind
wings with nervellus opposite or slightly antefurcal and intercepted about in middle,
or a little below.
Legs of normal, slender form: hind metatarsus about 7 times length of its greatest —
breadth in inner side view in female, 6 to 7 times in male. Tarsal claws moderately,
sometimes weakly pectinate.
Length of normal form 7 to 12 mm.
Female. Head black, with the following parts ivory-white, often in large part pale —
flavous, sometimes merging to fulvous, particularly on palpi: clypeus, genae, lower
face, except narrowly below and between antennae, and mouth-parts except mandi-
bular apices. Antennae dark above, below with scape pale flavous, and the segments —
that follow pale flavous, merging to fulvous in apical half of antenna. Thorax and
propodeum black, with the following parts pale-coloured as on head: anterior margin —
at sides and humeral angle of pronotum, propleura, tegulae, laminate margins of
mesoscutum behind tegulae, wide margin of prepectus, sometimes a pair of marks ©
beside mesolcus, line below fore wings, mesepimera, scutellum at apex and meta- ©
scutellum and their lateral extensions, and piliferous lobes of metanotum. The scu-—
tellum is often more or less dull fulvous discally, and the propodeum may be dull
fulvous marked. Legs mainly fulvous to rufous; with fore and mid coxae and tro-
chanters and fore femora anteriorly flavous, merging to fulvous: apical segment of
mid tarsus and hind femora at apex infuscate: hind tibiae and tarsi almost black,
sometimes paler below: hind coxae and trochanters occasionally darkened. Wings —
slightly infumate, with veins and stigma fuscous to ferrugineous, basally pale. Gaster
black: tergite I sometimes with a median apical spot, and following tergites more and
more broadly margined apically and laterally with ivory-white, the margins often
bordered anteriorly with pale fulvous: the white colouring covers more than half
the exposed surfaces of tergites 6 and 7.
Male. Differs from the female as follows: genae more narrowly pale, usually black
between outer and inner costae ; lower face occasionally black in mid line. Propleura
generally quite black, occasionally pale near mid line and behind: pale margins of
pronotum and prepectus absent or reduced to mere dots. Hind coxae and trochanters
usually in greater part black: fore and mid coxae sometimes dark-marked (especially : |
a
iy
THE CTENISCINI OF THE OLD WORLD 397
in British specimens). Pale margins of tergites becoming broader towards apex of
gaster much more gradually ; the white colouring usually covering less than half the
exposed surfaces of tergites 6 and 7.
This species is illustrated on Pl. 6.
Redescribed from the following material: SCOTLAND, Inverness-shire W., Arisaig,
I 4, 9.vii.1939, A. W. Stelfox: ENGLAND, Dorset, Studland, South Haven Peninsula,
12-acre Wood, I 9, 22.vi.1934, G. J. K. (compared with Gravenhorst material and
with type of evosus Hgr.); Surrey, Chobham, 1 9, 16.v.1891, Melville-Bury coll. ;
Essex, Colchester, 3 $3, 1914; Suffolk, Sudbury, 1 g, 1921, W. B. or B. S. Harwood;
Bucks., Princes Risborough, I 9, 21.vi—4.vii.1942, Brickhill, 1 $, 21.vi.1937, Herts.,
Brickett Wood, 1 4, 9.vi.1936, R. B. Benson; Staffs., Lichfield district, 2 $3, I 92,
1923, L. A. Carr (one recorded as Exyston calcaratus Thoms.) ; Yorks., Ayton Wood,
I 9, 10.vii.1921, F. Pepper: BELGIUM, unlocalized, 1 2, C. Wesmael ; Groenendaele, 1 9,
27.vi.1880; Ormeignies, I 9, 29.vili.1880; Hestreux, 2 99, 6.vili.1884; Rhode St.
Genése, I 3, 27.vi.1884; Vivier d’Oie, I 9, 19.viii.1888; Espinette, I g, 26.v.1890;
Verrewinkel, 1 3, 4.vi.1891, J. Tosquinet: HoLLanp, Venlo, I 9, 21.vi, I 3, 24.viii,
I Q, 29.viii.1942 ; Meiendel, x 9, 13.vii.1939, I g, I 9, 6.vi.1940; Schayk, I 2, 19.vi.
1948, H. G. M. Teunissen: SWEDEN, Skane, H6or district, 1 g, 13.vii; Réstanga, I 9,
5.vii, Degaberga, 1 9, 10.vii.1938, D. M. S. & J. F. Perkins; Smaland, 1 9, C. Bohe-
man (type of evosus Higr.): FINLAND, Lojo, 1 9, R. Krogerus; I 9, 17.viii.1918, Hakan
Lindberg: FRANCE, Sévres, I 9, 3.vi.1895, J. de Gaulle: SWITZERLAND, Nyon, I 9,
Cheyr: GERMANY, Gottingen, I 9, 29.vi.1948, Helzen, I 2, 26.v.1943, I g, 28.v.1945,
R. Hinz; Bavaria, Bodmann am Bodensee, I 9, 19.vi.1949, Bayrische Wald, Wald-
miinchen, 800 m., I 9, 16.vi.1948, G. Heinrich; Gérkau, I 9, vii.1930, Th. Kupka:
CZECHOSLOVAKIA, Bohumin, I 9, 14.vii.1938, Th. Kupka; Chodau, I 9, 27.vii.1886,
R. von Stein: Russi, Orel, 1 9, 22.vi, I gf, 26.vi.1943, R. Hinz.
Further material seen: FRANCE, Maisons Laffitte, 3 specimens, J. de Gaulle.
The small, pale form of this species has the costae of the propodeum and petiolar
segment very strongly developed, but the punctation in parts relatively poorly so.
Apart from the paler colour, especially of legs and gaster, it agrees with the descrip-
tion of the species given above. Antennal flagellum 26- to 27-segmented: length 6}
_ to7mm. Described from the following: BELGIuM, I 3, 1 9, C. Wesmael: GERMANY ?;
_ I 9, O. Schmiedeknecht: SWEDEN, Skane, Kivik, 1 9, 19.vii.1938, D. M.S. & J. F.
Perkins.
_ The Irish record of this species (W. F. Johnson, 1915: 132) is based on a male of
Acrotomus succinctus Grav.
I do not think it will be possible to determine, from Gravenhorst’s description of
1820, which species of Smicroplectrus he then had before him. Since the original
- material is not to be found in the Spinola collection in Turin (see Kerrich, 1950), we
must accept the monograph of 1829 as the first revision. The England, Dorset, speci-
men was compared by me with the specimens in Breslau in 1938, and was found to be
the same species. In 1947 I compared it with the type of evosus Hlgr. and thus con-
firmed the synonymy suggested by Pfankuch (1906).
398 A REVIEW AND A REVISION OF
Smicroplectrus excisus sp. n.
Head (Fig. 65) not strongly transverse; sharply and, in type form, almost recti-
linearly narrowed behind eyes; shallowly emarginate behind: finely to moderately —
punctate, seemingly on the average more finely so in male; cheeks below, lower face
and upper face densely and rather coarsely punctate, occasionally rugose-punctate.
Clypeus more sparsely punctate and usually apically less narrowly rounded than in
guinquecinctus Grav., the punctures fine to moderate; rather deeply delimited, with
tentorial pits conspicuous; in type form not strongly convex basally, and rather
= 60 ae
ms amos Us
Fics. 60-61. Left antennal scape, in sinistro-lateral view, of Fig. 60. Smicroplectrus quinque-_
cinctus Grav. Fig. 61. Smicroplectrus excisus sp.n. Fig. 62. Left hind metatarsus in inner side
view of Smicroplectrus excisus sp. n.
shallowly impressed before apex. Malar space about equal to distance between man-
dibular apices in female, distinctly less in male. Genal costa laminately elevated,
though less strongly than in quinquecinctus Grav., angularly inflected: gena with
outer costa and genal pit as described for guinquecinctus. Hypostomal costa strongly
raised, obtusely round-pointed below junction of genal costa.
Antennae with scape (Fig. 61) strongly convex above and very deeply. excise
externo-ventrally: flagellum normally 25- to 30-segmented, moderately explanate in
middle.
Pronotum with dorsal transverse shining impression narrow, rather indeterminate ;
with epomiae rather strong; laterally rather strongly punctate above, more finely
rugulose-punctate below. Propleura moderately to rather coarsely but shallowly
punctate. Mesoscutum in type form weakly convex anteriorly, but often rather
strongly so; with notaulices broad but not deeply impressed anteriorly, running far
beyond middle and falling into a median depression ; moderately and sparsely pun
tate, rather more finely and densely so anteriorly. Scutellum strongly convex, norm-
ally not margined to middle; moderately and usually sparsely punctate discally,
much more densely punctate to rugose-punctate peripherally ; slightly impressed on ~
disk. Mesopleura rather coarsely punctate, with a shining speculum: sternaulices —
broad, moderately impressed: mesosternum finely and densely punctate, more
sparsely so behind: prepectus coarsely alutaceo-punctate, deeply excavate: epi
nemial carina moderately elevated: mesolcus broad, moderately to deeply impressed,
J
THE CTENISCINI OF THE OLD WORLD 399
almost always transcostate, closed by a strong costa behind. Propodeum with costae
rather strong ; with costulae complete ; finely punctate to rugose, with area pleuralis
discretely punctate except below ; with area posteromedia much more strongly rugose:
area basalis delimited apically: area superomedia usually almost regularly hexagonal,
emitting costulae about in middle, but sometimes much more strongly narrowed to
base and with costulae well behind middle.
Gaster on tergites I mainly, and 2 and 3 at base, coriaceo-rugose, but petiolar
segment sometimes almost smooth between the dorsal keels ; elsewhere usually finely
and rather sparsely punctate and strongly shining. Petiolar segment normally 1-3
to 1:6 times length of its greatest breadth ; moderately strongly margined ; with basal
_ lobes pointed, sometimes acutely so ; with dorsal keels of moderate strength to strong,
_ almost percurrent. Tergite 2 with diagonal basal impressions usually distinct and
_ rather strong, but ill-defined.
Wings as described for quinquecinctus Grav., but stigma and radial cell on the
_ average about Io per cent. relatively broader.
Legs relatively stouter than in guinquecinctus Grav., notably the hind tibiae and
tarsi: hind metatarsus about 54 times length of its greatest breadth in inner side view
_ (Fig. 62). Tarsal claws moderately pectinate.
Length 54 to Io mm.
Differs little in coloration from quinquecinctus Grav. The scape and pedicellus
_ below have the pale coloration generally duller and more restricted. The lower face
_ is often more broadly black below the antennae and more distinctly on the mid line
_ above, and the clypeo-frontal suture is dark: by contrast the propleura are pale-
marked. In female, the pale marks of pronotum are less conspicuous, and the pro-
_ pleura are to some extent dark at sides: in male the pale margin of prepectus is often
_ well developed, though narrower than in female. Coloration of legs similar in female:
_ in male the hind coxae and trochanters are mainly fulvous. The pale margins of the
_ tergites are less extensive. Wings similar.
Described from the following material: BELGIUM, unlocalized, 1 g, 1 9, coll. Wes-
; mael; unlocalized, 1 g, 12.vii.1881, Hestreux, 3 99, 5-6.viii.1884, Orval, I 9, 7.viii.
=
) 1892, J. Tosquinet: HOLLAND, Voorburg, on Salix, I 9, 5.viii.1939, Biesbosch, I 9,
_ 23.vi.1943, H. G. M. Teunissen: SWEDEN, Skane, Palsj6, 2 33, 4 29, coll. C. G. Thom-
son ; Réstanga, 2 29 (one the holotype), 1 ¢ (allotype), 4.vii.1938, Hor district, 2 $4,
_ 26-28.vi.1938, D. M.S. & J. F. Perkins; province Halland, 3 $3, 23-27.vi, I1907—-I9II
- (mus. Stockholm): FRANCE, ?France, 1 g (under C?. sexcinctus Grav.), J. de Gaulle;
Ardennes, Verdun, I 9, I0.viii.1925, Benoist: SWITZERLAND, unlocalized, 1 g, D.
Imhoff (mus. Turin): GERMANY, Upper Bavaria, Haag-an-der-Amper, I g, v.1948,
_ G. Heinrich ; Celle, 1 9, 4.vii.1942, Géttingen, 1 g, 7.viii.1946, R. Hinz; 2 29, prob-
ably Berlin district, Ruthe coll. ; Warnemiinde, I 9, vili.1886, S. Brauns (Nat. Mus.
Ireland) ; unlocalized, r 9, O. Schmiedeknecht (mus. Genoa): POLAND, Pomorzia,
Sepolno, 2 $d, 23-24.vi.1936, G. Heinrich: Russia, Orel, 2 $¢, 26.vi.1943, R. Hinz.
Small pale form: GERMANY, I 4, 8.ix.1854, Ruthe coll. ; Warnemiinde, I 9, viii.1886,
S. Brauns.
Holotype in British Museum (Natural History).
This species is variable in a number of respects. In the type form the head is
400 A REVIEW AND A REVISION OF
almost rectilinearly narrowed behind the eyes (Fig. 65), the clypeus is not strongly
convex basally, the mesoscutum is weakly convex anteriorly, and the scutellum is not
strongly margined: in other specimens the head is decidedly rounded behind the eyes,
the clypeus is much more strongly convex basally, the mesoscutum is rather strongly
convex anteriorly, the scutellum is more strongly margined, and the body colour is
rather paler. In the small pale form the body colour is altogether paler, the gaster —
is mainly ferrugineous, and the hind tibiae and tarsi are ochreous to ferrugineous: the
flagellum is 23- to 26-segmented, and the length is 44 to 54 mm. Such differences
might be due to the use of different host species, or to the nutrition of the individual —
hosts. The Thomson collection contains a series of five females and five males, all —
from Palsjé: 1 2, 3 gd are of the small, pale form ; and both other forms are included
but are not sharply separable. All agree in the peculiar conformation of the scape and —
the gena, and in the possession of relatively short and thick hind legs: these characters —
distinguish the species most readily from its nearest known relative guinquecinctus —
Grav.
This species was identified by C. G. Thomson and Brauns as erosus Hlgr. but, as
shown above, the synonymy of evosus Hlgr. with guinquecinctus Grav. has now been
confirmed.
Smicroplectrus trianguligena sp. n.
Head not strongly transverse ; moderately to rather strongly narrowed behind eyes |
in female, rather strongly to strongly so in male, less rounded than in guinquecinctus
Grav. ; rather shallowly emarginate behind: on vertical region and temples finely or ~
rather finely punctate, and on genae and orbits of lower face moderately so ; on epi-
stoma rather coarsely punctate, sometimes in part confluently so: upper face rather
finely to moderately and densely punctate, often transversely rugose-punctate above
antennal sockets. Clypeus as described for guinquecinctus Grav. Malar space about
equal to distance between mandibular apices in female, a little less in male. Genal
costa moderately raised, a little less so than in bohemani Higr., more or less angularly
inflected: hypostomal costa much as described for qguinguecinctus Grav. Gena (Fig.
470) sharply angulate in position of outer costa, the angulation sometimes a little
raised near mandibular articulation: the enclosed genal pit usually longer, along —
hypostomal costa, than its breadth along genal, and often traversed by one or two ~
weak costae.
Antennae much as described for guinquecinctus Grav.: flagellum 28- to 33-seg-
mented. :
Thorax and propodeum structure differing from that described for guinquecinctus —
Grav. as follows: propleura moderately alutaceo-punctate: mesoscutum more evenly —
convex: epicnemial carina less elevated and sternaulices very superficial: propodeum ~
with costae generally strong; with area superomedia sometimes weakly delimited
basally: area pleuralis with punctation weak or rather weak.
Gaster with tergites 1 and 2 mainly, and 3 at base, coriaceo-rugose ; elsewhere
shining, more and more finely punctate towards apex: otherwise as described for
guinquecinctus Grav., but petiolar segment on the average a little longer, 1-6 to 1'9
times length of its greatest breadth.
ap ee ee
Bie
THE CTENISCINI OF THE OLD WORLD 401
Wings as described for guinquecinctus Grav., but areolet usually longer petiolate.
Legs slender as in quinquecinctus Grav.: tarsal claws rather weakly pectinate.
Length 73 to Ir mm.
_ Coloration very much as described for guinquecinctus Grav., but male propleura
_ regularly pale-marked near mid line, and sometimes more extensively so. Male hind
coxae not darkened in any but the Irish specimens. Pale margins of tergites some-
times less extensive.
} Described from the following material: ENGLAND, Staffs., Lichfield dist., 1 3, 1921,
L. A. Carr (recorded as jucundus Hgr., Carr, 1924: 53): BELGIUM, unlocalized, 2 3d,
I 9, C. Wesmael; Vivier d’Oie, r g, 29.v.1891, I 2, 17.vii.1898 (?), J. Tosquinet:
HOLLAND, Meiendel, 1 g, 2.vi.1937, H. G. M. Teunissen: FRANCE, Maisons-Laffitte,
14 (no date), St. Cloud, 2 gg, 11-12.v.1887, Vincenne, I 3, 30.iv.1890, Mendon, 1 9,
_ 23.vi.1891, Mesnil, 1 2, 20.v.1892, Forét de St. Germain, I 3, 5.v.1894, Fontainebleau,
_ 19, 17.vi.1898, J. de Gaulle ; Nantes, 2 99, T. A. Marshall; Mayenne, la Sanbee, 1 9,
24.V.1926, Coulans, I 2, 24.v.1926, Benoist ; Yonne, Mezilles, 1 2, v.1940, I 9, vi.1946,
_ E. Cavro; ‘St. Dii.N.’ (? France), 2 $3, 15.v.1921, Haut-Rhin, Bollwiller, 2 29,
_ 30.v.1920 (one of the type), Roderv, 1 3, 12.vi.1932, A. Seyrig: SwITZERLAND, Nyon,
_ IQ, Cheyr; Peney, 1 g, 1886, Tournier: GERMANY, Rhineland, Geisenheim Notgottes,
_ Forsthaus Riidesheim, 1 g, 13.v.1923, E. Schmidt; Gottingen, I g, 10.v.1947, R.
_ Hinz; Bodmann am Bodensee, 1 2, 17.v.1949, G. Heinrich: CZECHOSLOVAKIA, Erzge-
birge, Aussig, I 3g, 26.v.1935, Th. Kupka: PoLanp, Wlodzimierz, 1 9, 17.vi.1944, R.
_ Hinz: Europe, 1 (no interpretable data).
Type 2 in Muséum d’Histoire Naturelle, Paris.
_ This species comes between quinquecinctus Grav. and bohemani Hlgr.; and small
specimens, especially males, may be difficult to separate from both. It does, however,
_ differ from both in having the greater part of the gaster not alutaceo-punctate but
_ strongly shining. The male hind coxae are not darkened in any but the doubtfully
associated Irish specimens mentioned below, whereas those of quinquecinctus are
considerably darkened in all western European material seen.
_ Provisionally I place here four small male specimens having the following data:
_ IRELAND, Co. Wicklow, Powerscourt, 2 gg em. 4-II.v.1932, ex sawfly larvae, G. C.
_ Scott ; George’s Bridge, 1 3, 19.v.1940, Co. Dublin, Glenasmole, 1 g, 21.v.1939, A. W.
_ Stelfox. Length of these specimens 6 to 8 mm.: flagellum 23- to 27-segmented. In
- correlation with the small size, however, the characters are weakly developed ; and
_ confirmation of the occurrence of this species in Ireland by the capture of the female
is much to be desired. Mr. Stelfox has kindly presented one of the Powerscourt
_ specimens to the British Museum (Natural History).
_ Confirmation of the single English record would also be very acceptable.
Smicroplectrus bohemani Holmgren
1858 Exenterus Bohemani Holmgren, Svensk Vet.-Akad. Handl. 1: 226-227.
1911 Microplectron Bohemani Schmiedeknecht, Opuscula Ichneumonologica, §: 2310, 2311.
1913 Smicroplectrus Bohemani Roman (pars), Ent. Tidskr. 34: 128.
Head not strongly transverse; slightly to moderately narrowed behind eyes in
female and moderately to rather strongly so in male, rather strongly rounded pos-
ENTOM. II. 6 3D
402. > A REVIEW AND A REVISION OF
teriorly ; shallowly emarginate behind: on vertical region and temples rather finely —
and sparsely punctate, more densely so before occiput: genae rather coarsely and
more or less densely, upper face densely and rather coarsely, and epistoma coarsely _
punctate; upper face and epistoma usually alutaceous and more or less confluently —
punctate. Clypeus coarsely and rather densely punctate above, sparsely and much
more finely so below; deeply delimited; usually moderately convex basally and
rather shallowly impressed before apex, which generally is narrowly rounded. Malar
space about equal to distance between mandibular apices. Genal costa distinctly but —
not strongly laminately elevated, not angularly inflected, meeting hypostomal costa —
a moderate distance behind mandibular base, about as in quinquecinctus Grav.
(Fig. 69): gena with no outer costa, but strongly rounded in that position. Hypo-
stomal costa less strongly elevated than in quinquecinctus Grav., rounded or very
obtusely pointed below junction of genal costa.
Antennae with scape rather weakly or moderately excised externo-ventrally:
flagellum 26- to 32-segmented.
Pronotum with dorsal transverse shining impression narrow, deeply impressed;
with epomiae strong and sharp; laterally rather coarsely and very densely punctate
above, merging to alutaceo-punctate below. Propleura densely and rather coarsely —
punctate. Mesoscutum normally convex anteriorly ; with notaulices deeply impressed —
and often transcostate anteriorly, running far beyond middle and falling into a
median depression ; finely to moderately punctate, more finely and densely so an-
teriorly between notaulices. Scutellum moderately convex; usually rather strongly
punctate above, densely so behind; slightly impressed on disk. Mesopleura coarsely
and more or less densely punctate, less coarsely so above, and witha shining speculum:
sternaulices superficial: mesosternum rather finely and densely punctate, more
sparsely so posteriorly: prepectus coarsely and very densely punctate: epicnemial —
carina weakly elevated: mesolcus narrow anteriorly, broadened posteriorly, moder-
ately impressed, incompletely closed behind. Propodeum generally with costae strong |
and costulae complete; with dentiparal areae often rather strongly produced ; finely
to moderately rugose, with the area posteromedia usually strongly so, and the area’
pleuralis for the most part discretely punctate: area basalis not delimited apically:
area superomedia rather shining and often somewhat excavate. ,
Gaster as described for guinguecinctus Grav., but usually less strongly alutaceo- ~
punctate and with diagonal basal impressions indistinct: petiolar segment 1-3 to 1-7
times length of its greatest breadth. ,
Fore wings as described for guinquecinctus Grav.; but hind wings with nervellus
often intercepted well below middle.
Legs slender as in guinquecinctus Grav.: tarsal claws weakly to moderately
pectinate.
Length of normal form 7 to 124 mm.
Coloration very much as described for quinquecinctus Grav., but the ivory-white
usually purer, with much less flavous. Lower face of female sometimes black more
than half-way down in mid line ; of male sometimes black except for inner orbits and”
a pair of pale marks on epistoma: male genae only narrowly pale. Trochanterelli
usually mainly flavous, and hind coxae often distinctly flavous at apex: hind femora ~
THE CTENISCINI OF THE OLD WORLD 403
sometimes extensively darkened ; male coxae and trochanters less often and on the
average less extensively darkened than in guinquecinctus Grav.: male hind kneés
inconspicuously pale. Pale margins of tergites often less extensive.
Redescribed from the following material: ENGLAND, unlocalized, 1 2, Desvignes
coll.; Herts., Brickett Wood, I 3, 5.v.1950, R. B. Benson; Notts., Oxton Bogs,
I 9, 14.vii.1915, J. W. Carr: HOLLAND, Venlo, I 9, 21.vi.1942, H. G. M. Teunissen:
SwEDEN, Skane, Lund, 1 9, 31.v.1939, W. Hellén; Smaland, ‘Osst. Lars’, 1 9,
17.vi.Ig1I (mus. Stockholm); Stockholm, 2 gg, C. Boheman (one the type);
Dalarne, Orsa, I 9, 4.viii.1938, G. J. K.: FINLAND, Karislojo, 1 ¢, 13.vi, R. Forsius;
Helsinge, 1 2, W. Hellén; Tampere, Aitolahti, 1 9, 22.vi.1932, V. Saarinen: SWITZER-
LAND, Ziirich, 1 9, Buchecker coll.: GERMANY, probably Berlin district, 1 9, Ruthe
coll.; Celle, r g, 25.v.1942, 2 29, 27.vi.1942, Munster, I 9, 2.viii.1942, R. Hinz;
Upper Bavaria, Haag-an-der-Amper, 3 gd, 2 29, v.1948, G. Heinrich: CzEcHO-
SLOVAKIA, Bohemia, Chodau, I 9, I.vi.1882, R. von Stein; Bohumin, Haatsch, 1 9,
24.vii.1938, Th. Kupka: SIBERIA, Kamtchatka, 2 99, R. Malaise.
Small pale form has body colour altogether paler: antennal flagellum 22- to 27-
segmented: length 54 to 7 mm.: BELGium, Mont St. Aubert, 1 9, 1.vi.1878, J. Tos-
quinet: SWEDEN, Smaland, Lommaryd, 1 2 (mus. Stockholm): Europe, unlocalized,
I 9, O. Schmiedeknecht.
Smicroplectrus sibiricus sp. n.
Head rather strongly transverse, more so than in perkinsorum sp.n. (Fig. 66) ; strongly
rounded and almost broadened behind eyes; shallowly emarginate behind: finely to
moderately and rather densely punctate ; lower face more coarsely so. Clypeus rather
coarsely punctate; evenly rounded apically; deeply delimited, with tentorial pits
conspicuous ; scarcely convex basally, but distinctly impressed before apex. Malar
space slightly less than distance between mandibular apices. Genal and hypostomal
costae much as described for bohemani Hlgr.; but the hypostomal less strongly
elevated, and the genal meeting it relatively nearer mandibular base. Mandibles with
lower tooth contracted, as in jucundus Hlgr.
Antennae with scape weakly excised externo-ventrally: flagellum 23-segmented,
very little explanate in middle, but rather strongly hairy, and with the segments very
distinctly separated.
Pronotum with dorsal transverse shining impression narrow, deeply impressed ;
with epomiae sharp and of moderate strength ; laterally rather coarsely but shallowly
and not densely punctate above, longitudinally rugose just behind epomiae and below.
Propleura alutaceo-punctate. Mesoscutum very strongly convex anteriorly; with
notaulices weakly impressed; moderately punctate, more finely and densely so
anteriorly between notaulices. Scutellum rather strongly convex anteriorly, falling
rather gradually behind; not margined to middle; shining and almost smooth on
disk, finely rugose-punctate at sides and along hind margin. Mesopleura moderately
and rather sparsely punctate, with a shining speculum: sternaulices broad, rather
superficial: mesosternum shining and finely punctate, sparsely so posteriorly: epic-
nemial carina very distinctly elevated, not quite complete dorsally: mesolcus deep,
transcostate, almost closed behind. Metascutellum moderately raised. Propodeum
404 A REVIEW AND A REVISION OF
with costae rather strong and costulae complete; with area externa rugulose-punc-
tate, elsewhere mainly rather strongly rugose: area basalis not delimited apically: —
area superomedia slightly rounded at sides and behind.
Gaster on tergite I mainly, and on 2 to beyond middle, finely rugose; elsewhere —
shining and moderately punctate, more finely so towards apex. Petiolar segment about
1°5 times length of its greatest breadth (female) ; distinctly margined ; with anterior
angles not acutely pointed ; with dorsal keels distinct and almost percurrent, though
not strong. Tergite 2 with basal diagonal impressions not distinct.
Fore wings with stigma and radial cell relatively broad: stigma about 2:8 times,
radial cell about 2-3 times as long as broad. The areolet is relatively a little larger, but —
in other respects the wings are as described for quinquecinctus Grav.
Hind tibiae without the minute spur characteristic of the genus. Tarsal claws ©
weakly pectinate, with only two or three slender teeth.
Length 6 mm. ,
Coloration of head and body much as in quinquecinctus Grav., but the pale colour
is purer ivory-white, and the pale margin of the prepectus is reduced to a relatively
narrow band on the ventral surface and a pair of rather large lateral marks. Wing
veins paler than in guinquecinctus Grav. Fore and mid legs mainly pale fulvous; with —
coxae, trochanters and trochanterelli ivory-white, the coxae darker marked at
extreme base: hind legs mainly pale ferrugineous, but with coxae mainly almost —
black ; with femora at apex and tibiae at base conspicuously ivory-white ; with coxae
at extreme apex, trochanters and trochanterelli in part, and first four tarsal segments —
basally, ivory-white.
SIBERIA, Yeniseisk, I 9, J. Sahlberg.
Holotype in Royal Natural History Museum, Stockholm.
The white hind knees and absence of a hind tibial spur are characters shared with -
annulipes Cresson and velox Walley. H. K. Townes has very kindly sent me a~
female specimen of each of these nearctic species. Both differ from szbivicus as follows:
costae of propodeum strong: petiolar segment with dorsal keels very strong; with —
basal lobes acute-pointed and very prominent: tergite 2 with strong longitudinal
rugosities ; with diagonal impressions very strong and deep. S. annulipes differs also
in having the dentiparal area of the propodeum strongly produced, the tarsal claws
strongly pectinate, the thorax ventrally red, and the hind tibiae very broadly and
distinctly white in the middle. The area posteromedia of annulipes I should describe
as moderately rugose transversely, not as aciculate, as given by Davis (1897: 233).
Smicroplectrus clémenti sp. n.
Head strongly transverse, more so than in ferkinsorum sp.n. (Fig. 66); rather —
strongly narrowed and rounded behind eyes; rather shallowly emarginate behind: on ~
vertical region and temples finely punctate, towards and on greater part of genae
rather finely so ; near mandibular base, and on inner orbits and upper face moderately
punctate, and on epistoma rather coarsely and densely so. Clypeus rather finely
punctate above, almost impunctate below; rather deeply delimited ; rather strongly —
and almost evenly convex ; moderately rounded at apex. Malar space about three- —
—"
THE CTENISCINI OF THE OLD WORLD 405
quarters distance between mandibular apices. Genal costa moderately laminately
elevated, bulging outward (Fig. 73), not angularly inflected: gena with no outer costa,
but roundedly angled in that position. Hypostomal costa strongly and rather sharply
raised. :
Antennae with scape weakly excised externo-ventrally: flagellum 24-segmented.
Pronotum with epomiae of moderate strength; laterally rather finely punctate
above, finely so below. Propleura very shallowly, moderately to rather coarsely punc-
tate. Mesoscutum rather strongly convex ; with notaulices rather deeply impressed
for a short distance near, but not reaching anterior margin; finely to rather finely
punctate. Scutellum moderately convex, rather irregularly punctate, margined to
about middle. Mesopleura moderately and not densely punctate below, rather finely
so above, with speculum rather extensive: sternaulices broad, moderately impressed:
_ mesosternum finely and densely punctate: epicnemial carina rather strongly raised:
mesolcus sharply impressed, closed behind. Propodeum with costae of moderate
strength and costulae complete ; rather finely rugose, with area pleuralis in greater
part and area externa discretely punctate: area superomedia a little broader than
long, basally delimited.
Gaster with tergites I and 2 mainly, and 3 at base, weakly coriaceo-rugose ; else-
where shining and finely punctate. Petiolar segment relatively long, 2-0 times length
of its greatest breadth ; distinctly margined, with basal lobes pointed and with spira-
cular tubercles rather prominent ; rather much raised in middle, but with dorsal keels
_ not strong and not distinct much beyond half length of segment.
Wings as described for quinquecinctus Grav., but basal and disco-cubital veins
distinctly divergent, the basal meeting the first cubital at an acute angle.
Legs rather short and stout, about as in perkinsorum sp.n.: hind tibiae without the
minute spur characteristic of the genus: tarsal claws not pectinate.
Length 64 mm. ;
Differs in coloration from guinquecinctus Grav. as follows: pale colour mostly bright
flavous, extending some way up orbits of upper face, that on antennae scarcely
_ merging to fulvous: colour pattern of thorax as described for perkinsorwm sp. n.
(p. 407). Leg colour much as described for sibivicus sp.n. (p. 404), but fore and mid
coxae and trochanters mainly pale flavous, hind knees not nearly so conspicuously
pale, and hind tibiae broadly luteous in middle. Gaster pale ferrugineous, the margins
_ bright flavous.
AustTrIA, ‘Carinth. alp.’, 1 9, C. Boheman.
Holotype in Royal Natural History Museum, Stockholm.
Dedicated to the late Ernst Clément, in recognition of his pioneer work on the
intensive revision of groups of European Ichneumonidae.
It seems very likely that this unique specimen is a small pale form of a species
normally larger and darker.
Smicroplectrus perkinsorum sp. n.
Head (Fig. 66) rather strongly transverse; moderately narrowed and rounded
behind eyes; rather shallowly emarginate behind ; in side view (Fig. 67): on vertical
406 A REVIEW AND A REVISION OF
region and temples finely and rather closely punctate, the punctures becoming gradu-
ally coarser towards mandibular articulation: genae rather finely to moderately and —
epistoma moderately punctate ; upper face densely and rather finely so. Clypeusas _
described for guinguecinctus Grav., but more sharply delimited basally, and usually
more broadly rounded at apex (considerable variation in this): the punctation also —
is weaker in smaller specimens. Malar space slightly less than distance between
mandibular apices in female, distinctly less in male. Genal costa sharply carinate —
but not laminate, not angularly inflected: gena falling to a deep, longitudinal pit
beside hypostomal costa; the pit bounded externally by a sharp angulation of the —
gena. Hypostomal costa (visible in Fig. 67) exceptionally strongly elevated and
acutely, or almost acutely, round-pointed below junction of genal costa; falling
evenly to foramen magnum.
Antennae with scape weakly or rather weakly excised externo-ventrally: flagellum
21- to 26-segmented, relatively short and thick, and not much attenuate to apex. |
Pronotum with dorsal transverse shining impression narrow; distinctly rugose-
punctate before it; with epomiae sharp and rather strong ; moderately and densely ~
punctate above, more or less longitudinally rugose below. Propleura rather coarsely __
but shallowly punctate. Mesoscutum rising rather sharply from pronotum; with
notaulices broad and strongly impressed anteriorly, running far beyond middle and ~
falling into a median depression ; finely to rather finely punctate. Scutellum weakly
convex ; moderately and rather irregularly punctate ; margined to about middle ; more —
or less impressed on disk. Mesopleura rather coarsely punctate, with speculum large: ©
sternaulices broad, rather shallowly impressed in female, shallowly in male: meso- —
sternum rather finely punctate, sparsely so behind: prepectus rugose-punctate at sides, —
very deeply excavate: epicnemial carina distinctly but not at all strongly raised: —
mesolcus very deeply impressed, incompletely closed behind. Metascutellum short, —
moderately raised. Propodeum with costae relatively weak or rather weak; with —
costulae complete or incomplete, always indicated but sometimes indefinite internally —
and usually so externally ; finely rugose including the area pleuralis, which is often
somewhat shining but not discretely punctate; with area posteromedia rather more —
strongly rugose, having a median longitudinal carina at least indicated posteriorly —
but usually almost complete: area basalis delimited or not delimited apically: area
superomedia of rather inconstant shape, more or less explanate to near apex, ~
moderately rounded to almost truncate behind, usually emitting costulae well
behind middle.
Gaster on tergite I mainly, and 2 and 3 at base, coriaceo-rugose ; elsewhere finely
alutaceo-punctate, not matt but generally not strongly shining. Petiolar segment
(Fig. 75) relatively elongate, 1-8 to 2-2 times length of its greatest breadth; not —
strongly margined ; with basal lobes very bluntly rounded, or at any rate of similar
general shape; with dorsal keels rather weak, extending to rather near apex. Ter-—
gite 2 with diagonal basal impressions indistinct.
Fore wings with stigma and radial cell relatively broad: stigma about 3-0 times
(varying 2-8 to 3-3), radial cell 2-1 to 2-6 times as long as broad: with areolet some-
times longer petiolate: in other respects the wings are as described for guinguecinctus
Grav.
THE CTENISCINI OF THE OLD WORLD 407
Legs relatively stouter than in guinquecinctus Grav., particularly the tibiae: tarsal
claws distinctly, but weakly or rather weakly pectinate.
Length of normal form 6 to 8 mm.
Differs in coloration from quinquecinctus Grav. as follows: genae on the average
more narrowly yellow: pale colouring frequently extending well up orbits of upper
face in female: scape and pedicellus below with pale coloration usually duller and
more restricted. Propleura often largely black in female: pale coloration of pronotum
more restricted: pale margin of prepectus represented by a narrow transverse line in
middle and a pair of lateral dots, but these sometimes undeveloped in female, as
almost always in male: no fulvous colouring on thorax. In both sexes the fore and
mid coxae are usually to a greater or lesser extent dark, and the hind coxae and tro-
chanters mainly black or almost so: all trochanterelli flavous: hind femora often
d extensively darkened: hind tibiae and tarsi not nearly black, but ferrugineous or
paler. Pale margins of tergites rather less extensive than in guinquecinctus, the pale
coloration stramineous to pale flavous.
Described from the following material: ENGLAND, unlocalized, 1 3, 3 99, T.
Desvignes coll. ; Cornwall, Botusfleming, 1 9, T. A. Marshall; Devon, Lustleigh, 1 9,
II.vii.1934, R. C. L. Perkins (type), 1 J, 23.v, I 3, I 9, 3.vi.1933, J. F. Perkins ; Bovey
Tracey, I 9, viil.1933, Dartmoor, 2 3g, I 9, 8-25.vi.1935, I dg, vi.1938, R. C. L.
Perkins; Newton Abbot, 1 4, 22.vi.1938, R. C. L. Perkins, 3 $d, vi.1941, I 3g, I 9,
—-I0.vi, I g, 2 92, 14.vi, I Q, 16.vi.1941, I Q, vi.1942, J. F. Perkins (some taken on wild
rose) ; Kingsteignton, Starks Ponds, 10 $4, 2 29, 24.v—5.vi.1942, J. F. Perkins ; High-
week, 9 gd, I 2, 22.v—-9.vi.1942, D. M. S. Perkins and J. F. Perkins; Dunsford, 14,
vi.1941, Milber, 1 9, vii.tg41, J. F. Perkins; Dorset, Studland Heath, 1 4, 6.vi.
1928, R. B. Benson; Oxon., Henley, 1 9, 25.vi.1906, H. Scott; Surrey, Oxshott, 1 ¢,
28.v.1933, J. F. Perkins; Bucks., Beaconsfield, 1 3, 27.v.1934, J. F. Perkins; Herts.,
Brickett Wood, 1 3, 17.vi.1936, Brickhill, 1 9, 21.v.1937, R. B. Benson; Suffolk,
Sudbury, r 9, 1926, Essex, Wivenhoe, 1 9, 25.v.1915, W. B. or B. S. Harwood;
Yorks., Holmhouse Wood, I 9, 15.vi.1935, J. Wood: BELGIuM, Stockel, 1 9, 10.vi. 1890,
Steinbach, I 9, 12.vi.1896, Rhode St. Genése, I 9, 27.vi.1884, Mont l’Enclus, 1 Q,
4.vii.1890, J. Tosquinet: HOLLAND, Babberich, 1 9, 14.vi.1941, Venlo, 2 gd, 30.v—
I.vi.1942, H. G. M. Teunissen: FINLAND, Kirjavalaks, 1 g, W. Hellén; Isthmus
karelicus, Rajajoki, 1 9, 17.vi.1934, K. Lahtivirta: FRANCE, Paris district, Maisons
Laffitte, 1 9, 24.v, J. de Gaulle: GERMANY, K6ln, I g, 12.v.1942, W. Aerts; Celle, 1 Z,
25.v.1942, Hannover, 1 g, 8.vi.1948, R. Hinz; Blankenburg-i-Thiiringen, I Q, I.vi.
1924, H. Hedicke (mus. Paris); Odenwald, 1 g, 20.v.1939, Th. Kupka; Bavaria,
Erzgebirge, Gérkau, 1 g, Th. Kupka; Bodmann am Bodensee, 1 9, 28.v.1949, G.
Heinrich.
The small, pale form of this species has the body colour altogether paler than that
of the typical form: the hind tibiae are pale ochreous; and the gaster is pale ferru-
gineous, with the margins of the tergites sometimes bright flavous. Agreeing in
structure with the typical form. Antennal flagellum 22- to 25-segmented: length 5 to
63 mm. Described from the following: ENGLAND, unlocalized, 1 9, T. Desvignes:
CZECHOSLOVAKIA, Erzgebirge, Aussig, 2 29, 17.vi.1934; unlocalized, 2 gg, Th. Kupka.
Holotype ¢ in British Museum (Natural History).
408 A REVIEW AND A REVISION OF
Structurally, this is the most distinct species of Smicroplectrus in Europe. It is also
distinct in having been taken on wild rose; but it has not yet been bred, and this
association remains to be investigated.
I have much pleasure in dedicating this species to Dr. R. C. L. Perkins and to Mr.
and Mrs. J. F. Perkins.
Smicroplectrus heinrichi sp. n.
1949 Smicroplectrus jucundus Heinrich, Mitt. Miinchen. ent. Ges. 85-89: 121 (non Holmgren —
1858).
Head moderately to rather strongly narrowed and rounded behind eyes; rather
shallowly emarginate behind: on vertical region rather finely, and on temples, cheeks,
inner orbits, and upper face moderately and densely punctate, confluently so just —
above antennae; epistoma coarsely and usually confluently punctate, often longi-
tudinally rugose medially. Clypeus rather sparsely punctate, moderately so above,
more finely so below; sharply delimited ; distinctly but not strongly convex in basal
half; usually narrowly rounded at apex. Malar space almost equal to distance
between mandibular apices in female, distinctly less in male. Genal costa carinate —
but not laminate, except near junction with hypostomal; not angularly inflected,
meeting hypostomal costa far behind mandibular base (Fig. 72) ; gena with no outer
costa, evenly rounded in that position as seen from behind, almost straight-sided in —
lateral view: genal pit sometimes indicated. Hypostomal costa moderately raised,
somewhat pointed below junction with genal costa. Second segment of maxillary
palpi strongly swollen in female.
Antennae with scape weakly or rather weakly excised externo-ventrally, generally
rather short: flagellum 23- to 27-segmented.
Pronotum with dorsal transverse furrow rather broad, shining; with epomiae of
moderate strength; laterally moderately and densely punctate. Propleura densely —
and rather coarsely punctate. Mesoscutum much as described for bohemani Hlgr., but
the punctation finer. Scutellum rather finely to moderately and not sparsely punc-
tate above, usually more coarsely and densely so behind; usually strongly convex
and not margined beyond scutellar fovea, but often less convex and more or less.
longitudinally impressed on disk. Mesopleura rather coarsely and densely punctate
below, not much less so above, and with a shining speculum: sternaulices weakly or
moderately impressed: mesosternum finely and densely punctate, prepectus coarsely
and densely so: epicnemial carina scarcely raised: mesolcus usually rather weakly
impressed, greatly broadened in posterior half, more or less closed behind. Propo-
deum with costae of moderate strength or rather strong, and with costulae generally
complete but occasionally absent ; in greater part rugose, very finely to moderately
so; sometimes rugose on greater part of area pleuralis, but area externa generally —
discretely, extremely finely punctate: area superomedia usually smooth, not de- —
limited basally.
Gaster on tergites I and 2 mainly, and 3 at base, coriaceo-rugose; elsewhere
strongly shining, moderately punctate on tergite 3, more and more finely so towards
THE CTENISCINI OF THE OLD WORLD 409
apex. Petiolar segment as described for guinguecinctus Grav.; 1-4 to 2-2, usually
about 1? times length of its greatest breadth.
Wings as described for guinquecinctus Grav., but radial cell sometimes relatively
short in male.
Legs slender, as in guinquecinctus Grav.: tarsal claws generally (but not in Swedish
specimen) rather strongly pectinate, with about four rather long, outwardly pointing
comb-teeth.
Length 6 to 8$ mm.
Differs in coloration from quinquecinctus Grav. as follows: the ivory-white purer,
much less suffused with flavous. Pale colouring sometimes extending up orbits of
upper face in female, and sometimes much more restricted on antenna. Pale apical
marks of scutellum and metascutellum small or, more often, undeveloped in female,
not developed in any male seen: no fulvous colouring on thorax. Leg colour much
as described for bohemant, but male hind coxae on the average darker. Pale margins
of tergites sometimes less extensive.
Described from the following material: IRELAND, Co. West Mayo, Old Head, 1 4,
3 2 (one the type), 28.v.1942, Co. Kildare, Sallins, 1 2, 17.v.1933, A. W. Stelfox ;
Co. Wicklow, Athdown, I 4, I.v.1933, J. F. Perkins; George’s Bridge, I g, I.v.1949,
A. W. Stelfox ; Co. Dublin, Glenasmole, I 3, 23.v.1930, 2 $d, 3.V1.1934, I 3, 2I.V.1939,
I 4, 20.v.1949, A. W. Stelfox, Dublin dist., 1 9, ex sawfly larva, em. c. 4.v.1932, G. C.
Scott: SCOTLAND, Cairngorm Mts., Aviemore, 1 g, G. C. Champion (recorded as
jucundus Higr., Bridgman, 1889: 434): BELGIUM, I Q, coll. C. Wesmael: HOLLAND,
Tilburg, 1 9, 4.iv.1941, coll. Adriaanse ; Kijfhoek, 1 9, 17.v.1936, H. G. M. Teunissen
' (recorded as jucundus Higr., Teunissen, 1948): SWEDEN, Skane, Skaralid, 1 9, 3.vil.
1938, J. F. Perkins: FINLAND, Munksnas, 1 9 ex Nematine sawfly larva, em. 5.iv.1940,
E. Lindqvist : SWITZERLAND, Valais, Arolla, 1 9, ex larva Amauronematus sp. on Salix,
em. iv.1936, R. B. Benson: GERMANY, Helzen, 1 34, 13.v.1943, R. Hinz; Bavaria,
Haag-an-der-Amper, 5 3d, 5 29, v.1948, Bodmann am Bodensee, I 3, 25.iv.1949,
G. Heinrich.
Holotype 2 in British Museum (Natural History), kindly presented by Mr. A. W.
Stelfox.
I first received this species in good series from Ireland from A. W. Stelfox: later,
when a good Alpine series came from G. Heinrich, I was able to appreciate its chief
_ characteristic. Heinrich’s rich material of Smicroplectrus has been especially valuable
in the elucidation thus far of this difficult genus.
Smicroplectrus jucundus Holmgren
1858 Exenterus jucundus Holmgren, Svensk. Vet.-Akad. Handl. 1: 227 (non Heinrich 1949).
1883 Smicroplectrus jucundus Thomson, Opuscula Entomologica, fasc. 9: 888.
1883 Smicroplectrus costulatus Thomson, ibid.: 888.
1913 Smicroplectrus Bohemanit Roman (pars), Ent. Tidskr. 384: 128.
Head moderately transverse (more so than in guinquecinctus Grav.) ; slightly to
moderately narrowed and strongly rounded behind eyes; deeply emarginate behind ;
in lateral view relatively short, and with vertical region rather strongly elevated
(Fig. 68): shining, moderately, or in small specimens rather finely, and somewhat
ENTOM., II, 6 3E
4
410 A REVIEW AND A REVISION OF 2
densely punctate; with gena less densely and upper face much more densely so;
epistoma usually distinctly alutaceous, rather coarsely to coarsely punctate, often
in larger specimens rugulose-punctate (but not in lectotype): eyes very distinctly
divergent in female but much less so in male. Clypeus almost impunctate below,
sparsely punctate, with coarser and finer punctures, above ; rather narrowly rounded
apically ; not strongly convex and often not deeply delimited, though with tentorial
pits conspicuous. Malar space much longer than distance between mandibular apices
in female, slightly shorter in male. Genal costa single, carinate ; not strongly inflected,
but meeting the hypostomal at less than a right angle (Fig. 71). Hypostomal costa
distinctly and more or less laminately elevated, but not at all strongly so for a Smicro-
plectrus, its height being only about a quarter the distance of the junction with the
genal costa from the base of the mandible. Mandibles with lower tooth contracted.
Antennae with scape very weakly excised externo-ventrally: flagellum normally —
21- to 24-segmented, relatively short and thick, and very little attenuate to apex.
Pronotum with dorsal transverse shining impression narrow, rather deeply im-
pressed, usually longitudinally costate ; with epomiae sharp but rather weak ; laterally
moderately to rather coarsely and closely punctate above, rugulose-punctate below.
Propleura rather densely and coarsely punctate. Mesoscutum rather weakly to
moderately convex anteriorly, but rather regularly rounded, not so much flattened
before the scutellum as in other species here described ; with notaulices moderately or
rather strongly impressed and often transcostate anteriorly; finely to moderately
punctate, more finely and densely so anteriorly between notaulices; generally
strongly shining but sometimes conspicuously alutaceous (it is so in type of costulatus
Thoms.). Scutellum very strongly convex ; moderately punctate above, weakly and
irregularly rugose-punctate at sides, finely rugose behind; not or weakly margined
to middle. Mesopleura above moderately, below coarsely punctate; sternaulices
broad and not deeply impressed: mesosternum densely and finely, prepectus densely
and rather coarsely punctate: mesolcus usually not deeply impressed, very narrow
till near hind end of sternum, where it is usually crossed by several transverse rugosi-
ties: epicnemial carina not elevated; but in female the prepectus is somewhat bent —
over ventrally. Propodeum with costae moderately strong; with dentiparal areae
more or less weakly produced and with costulae complete or incomplete, sometimes —
scarcely indicated ; finely coriaceous, but with the area pleuralis often distinctly finely
punctate (it is so in type of costulatus Thoms.) and the area posteromedia rugose: area
basalis not or scarcely delimited apically. |
Gaster on tergites 1 and 2 mainly, and 3 and sometimes 4 at base, coriaceous ; else-
where alutaceo-punctate towards base of segments, at least in larger specimens, finely
and sparsely punctate and shining towards apex. Petiolar segment 1-4 to 1-9 times
length of its greatest breadth, but with the longer form predominating ; with basal
lobes generally obtusely pointed or almost rounded ; with dorsal keels not very strong,
usually indefinite in apical third. Tergite 2 with diagonal basal impressions indistinct.
Fore wings with stigma about 3-3 times, radial cell about 2-6 times as long as broad:
hind wings with nervellus intercepted distinctly below middle: wings otherwise as
described for guinquecinctus Grav.
Legs of normal, slender form in female, rather stouter in male. Hind tibiae with
THE CTENISCINI OF THE OLD WORLD 411
spur relatively short even for this genus. Tarsal claws rather strongly but finely
pectinate.
Length normally 6 to 8} mm. Length of small, pale form 44 to 6 mm.: flagellum
may be 18- to 19-segmented.
Differs in coloration from quinquecinctus Grav. as follows: ivory-white usually
much purer, with very little flavous colouring. In female, the pale facial colour
usually extends up the orbits of upper face almost to the top of the eye (though some-
times leaving a detached pale spot) ; but in male it is more restricted, often to a pair
of incomplete bands on the orbits of the lower face and a median heart-shaped mark
above the clypeus: pale colour of genae very restricted in male. In female the whole
fore margin of the mesopleuron may be pale, and the pale markings of scutellum and
metascutellum very restricted or entirely absent: in male the usual pale markings of
the thorax are mostly or entirely absent, though the tegulae are stramineous in both
sexes. The hind coxae and trochanters are fulvous in the English and black in the
majority of continental specimens, flavous at apex: hind femora and tibiae fulvous
to ferrugineous. Wing veins and stigma tending to be rather paler, especially in male.
As described by Holmgren, tergites 5 to 7 of female may be almost wholly ivory-
white.
Redescribed from the following material: ENGLAND, Surrey, Byfleet, ex larva
Amauronematus amplux Kon. (Tenthredinidae) on Betula (larva vi.1947), I Q em.
iv.1948, R. B. Benson; Herts., Brickett Wood, 2 33, 2.v.1949, I 2, 23.V.1950, I Qem.
iii.1945 ex larva Amauronematus sp. on Salix, R. B. Benson; Hunts., Woodwalton,
I 2, 21-23.v.1949, R. B. Benson: SwEDEN, Lapland, 4 33, 4 29, including I g, 2 22
of small, pale form ‘ad Ovickjock (D. D. Boheman et Wahlberg), ad Umefors (ipse) ’
(Holmgren, 1858: 227) ; Lapland (no further data), 2 Jd, 4 29, including type to the
description, under jucundus Hlgr. in Thomson coll. (one female a small, pale form),
I 2 (unique type of costulatus Thoms.): FINLAND, Pihtipudas, 2 99, 7.vi, I 2, 9.Vi.
1945, Borga, 1 9, Ivalo, 1 g, 2 92, Parikkala, 2 99, Haapajarvi, I 9, 3.vi.1945, W.
Hellén; Saana, 1 g, Nordman; Kouvola, I g, 24.v.1940, V. Karvonen; Tampere,
P. Pirkkala, 1 9, 27.v.1934, A. Saarinen, Aitolahti, 1 9, 12.vi.1932, V. Saarinen:
SWITZERLAND, Valais, Ferpécle, 5,000-8,000 ft., 4 gd, 2 292, 14-27.vi.1935, R. B.
Benson: GERMANY, Bavaria, Bodmann am Bodensee, I 4, 25.iv, I 2, 23.v.1949, G.
Heinrich.
Through the kindness of Dr. Kjell Ander I was able in 1947 to take the Thomson
material from Lund, and make direct comparison with the Holmgren material in
Stockholm. A. Roman’s lectotype is a female specimen taken by P. Wahlberg.
Records of jucundus from Belgium, Holland, Scotland, and the northern midlands
of England are, in this paper, referred to other species. The European distribution
as now known to me is arctic and sub-arctic, Alpine and sub-Alpine, and both sides
of London in southern England. How much importance can be attached, in the
present state of our knowledge, to negative evidence from the middle European
plains remains to be seen; but R. B. Benson points out to me that this corresponds
with the Siberian type of distribution (see Benson, 1950).
Three female and three male specimens from Mt. Rainier, Washington, U.S.A.,
taken by H. K. Townes and determined by him as albilineatus Walsh, are extremely
Fics. 63-66. Head from above of Fig. 63. Smicroplectrus quinquecinctus Grav. 9. Fig. 64.
Smicroplectrus quinquecinctus Grav. g. Fig. 65. Smicroplectrus excisus sp. n. Fig. 66. Smicroplec-
trus perkinsorum sp. n.
r.emd.
Fics. 67-68. Head in dextro-lateral view of Fig. 67. Smicroplectrus perkinsorum sp.n. Fig. 68.
Smicroplectrus jucundus Hlgr. hy.c. = hypostomal costa, y.md. = right mandible.
69 | 70
7\ de
Fics. 69-73. Head in hind view of Fig. 69. Smicroplectrus quinquecinctus Grav. Fig. 70. Smicro-
plectrus trianguligena sp. n. Fig. 71. Smicroplectrus jucundus Higr. Fig. 72. Smicroplectrus
heinrichi sp.n. Fig. 73. Smicroplectrus clémenti sp.n. Figs. 69-72 to same scale, Fig. 73 larger.
GS. 74-75. Petiolar segment of Fig. 74. Smicroplectrus quinquecinctus Grav. Fig. 75. Smicro-
plectrus perkinsorum sp. 0.
414 A REVIEW AND A REVISION OF
similar to the European material on which the description of jucundus was based, and
agree with the description in all respects. I do not, however, propose the placing of
albilineatus Walsh in synonymy ; but leave the problem with W. R. M. Mason, for
reasons that will emerge when his revision of the nearctic species is published.
KEY TO PALAEARCTIC SPECIES OF SMICROPLECTRUS THOMSON
1. Male antennae pale-banded: female unknown: Japan. quadriceps Uchida —
Female known, and male of all but two species: antennae not pale-banded . 2
2. Hypostomal costa exceptionally strongly laminate (to be seen in Fig. 67), acute-
angled or almost so below junction of genal costa, which is not laminate: gena
falling to a deep, longitudinal pit beside hypostomal costa, bounded externally
by a sharp angulation of the gena: head strongly transverse (Fig. 66): fore wings
with stigma and radial cell relatively broad (see description): petiolar segment
relatively elongate, with basal lobes rounded (Fig. 75), or at any rate of similar
shape. ‘ : : perkinsorum sp. nov.
If the gena has an outer costa or is ; angulate or r strongly rounded in that position,
the genal costa is laminately elevated: head not so transverse except in non- —
spurred species (couplet 7): in other respects not as above . : ‘ 3
3. Genal costa carinate, but not laminate, neither double nor angled in poe of
outer costa: no distinct genal pit ‘
Genal costa moderately or strongly laminate, not as in the two following species
(Figs. 71 and 72) . ‘ . ; : : 5
4. Head, notably eyes (viewed laterally), gosh (Fig. 68, cf. Fig. 67), with vertical
region rather strongly elevated: genal costa not strongly inflected, but meeting ~
hypostomal costa at distinctly less than a right angle (Fig. 71): hypostomal
costa weakly elevated, the height being only about a quarter the distance of the
junction of the genal costa from mandibular base ; . jucundus Higr.
Head not so short (similar to Fig. 67), nor vertical region thus elevated: hypo-
stomal costa moderately elevated, met by genal costa at nearly a right angle
far behind mandibular base (Fig. 72) , ; ; heinricht sp. nov.
5. Gena with no outer costa, but seis rounded in that position: genal costa
moderately laminate ' ‘ ; 6
Gena with an outer costa, at least in part, or sharply angulate i in ‘that position 8
6. Head less transverse than in perkinsorum sp.n. (Fig. 66): genal costa normally
curved, meeting hypostomal a moderate distance behind mandibular base, —
about as in guinquecinctus Grav. (Fig. 69): hind tibia with the minute spur
characteristic of the genus: [hind knees not conspicuously pale]
‘ ; , ‘ - : : : hohemdéil Hier.
Head more transverse than in perkinsorum sp.n. (Fig. 66): genal costa meeting
hypostomal much nearer mandibular base: hind tibia without the minute spur
characteristic of the genus i : ‘ : , ; : ; 7
7. Genal costa not bulging outward as in alternate: petiolar segment about 1} times
THE CTENISCINI OF THE OLD WORLD 415
length of its greatest breadth: basal and disco-cubital veins (as in most species)
nearly parallel: [hind knees conspicuously pale] . : . sibivicus sp.n.
Genal costa bulging very, distinctly outward (Fig. 73): petiolar segment about
twice length of its greatest breadth: basal and disco-cubital veins distinctly
divergent : : : : ; ‘ : : . clémenti sp. n.
8. Genal costa moderately raised: genal pit (Fig. 70) triangular, usually longer along
hypostomal costa than its length along genal: [differs from guinquecinctus Grav.
in having the gaster mainly shining, and from excisus sp.n. in the shapes of the
antennal scape and hind metatarsi] . , trianguligena sp.n.
Genal costa strongly laminately raised: genal pit distinctly broader along genal
costa than its length along hypostomal (Fig. 69) . : F : ‘ 9
g. Scape of normal form, weakly excised externo-ventrally (Fig. 60): legs of normal
slender form, the hind metatarsus about 7 times length of its greatest breadth
in inner side view in female, 6 to 7 times in male: gaster mainly alutaceo-
punctate, more or less matt . ‘ . guinquecinctus Grav.
Scape strongly convex above and very deeply excised externo-ventrally, its
ventral being less than half its dorsal length (Fig. 61): legs relatively stout, the
hind metatarsus about 54 times length of its greatest breadth in same view
(Fig. 62): gaster mainly strongly shining . ; : ‘ €XCISUS SP. N.
Genus EUDIABORUS, gen. n.
Diaborus Forster, Thomson 1883 non Woldstedt 1877.
This genus was diagnosed by Forster (1868) without included species. It was taken
up by Thomson (1883) ; and Roman (1939) showed that it was the genus that Forster
_ actually had. However, as has been recorded earlier in the present paper, Woldstedt
had, in the meantime (1877), included two species neither of which belongs to the
genus in question ; and Diaborus has to take sedulus Woldst. as type, by designation
_ of Viereck 1912. To fill the lacuna the present writer has (p. 315 above) proposed
Se
Eudiaborus as a new genus, taking as type the best-known species pallitarsis Thoms.
Thomson included four species he described as new; and a fifth, with normal third
valvulae and seemingly attached to conifer woods, which he interpreted as litura-
torius L. It is, however, evident that Thunberg (1822, as recorded by Roman, Ig12),
Zetterstedt (1840), Holmgren (1858), Brischke (1878), and later Schmiedeknecht
(r911) had more than one species mixed under the name lituratorius.
Gravenhorst (1829) described a species, Tvyphon scalaris, based on males; and
_ a second form, based on females, two with similar data, which he called Tryphon
sexlituratus. He placed scalaris, which had page priority, as questionably the male of
sexlituratus; and placed sexlituratus doubtfully in synonymy with lituratorius L.
Pfankuch (1906) associated these Gravenhorst species with the present genus, and
placed both in synonymy with lituratorius.
Roman (1909) described an arctic species glutiniatus, with a variety inversus which
he later (1931) recognized as a good species. Gregor (1937) described a new form
moravica, which he placed as a subspecies of lituratorius ; but he added that it might
416 A REVIEW AND A REVISION OF
well be a good species. The above I believe to be the sum total of European species —
described.
Roman (1932), in the course of a revision of the types in the collection of the
Linnean Society of London, showed that lituratorius L. does not belong to this tribe —
at all, but is actually Hypsantyx impressus Grav. He put forward scalaris Grav. as
the first available name for lituratorius auctt.
Later (1939) he revised the Thomson species. He considered lituratorius Thoms.
as an atavistic form of lituratorius auctt. with normal third valvulae, and added
filipalpis Thoms. to this aggregate. He also placed nigrifrons Thoms. in synonymy
with pfallitarsis Thoms. I do not think that these interpretations will be upheld ; for
J. F. Perkins has, in the British Museum collection, arranged provisionally under the
four Thomson species, forms that accord with Thomson’s diagnoses and which do
seem to be good species. Roman’s identification of pallitarsis Thoms. as a species to
be found in abundance parasitizing Nematus (Pteronidea) ribesit Scop. was, however, —
helpful.
Now Schmiedeknecht was evidently unable to comprehend those parts of Thom-
son’s monograph written in the Swedish language. He merely translated (1911)
Thomson’s Latin diagnoses of the four new species ; and he erected a barrier to further
understanding of the genus by giving a key to species, according to which the lower
face should not be wholly yellowish-white except in the female of ‘/itwratorius’. This —
was completely contrary to what Thomson wrote in his remarks on the genus. It is
clear that lituratorius or, since 1932, scalaris, has come simply to signify any Diaborus
Thoms. that any student of Ichneumonidae could not place in any of the other —
described species.
The genus requires complete revision with the aid of larger material than has been
available and, when this has been done, the Gravenhorst and Thomson types should |
again be examined. I do not, however, see any reason to identify scalaris Grav. with
lituratorius Thoms. non L. I think it far more likely to be pallitarsis Thoms. ; and it
should not stand in any publication as a separate species without other interpretation.
In order to make Thomson’s introduction to the genus (p. 888) more widely avail-
able to students of Ichneumonidae, I give below an English translation of it. I do
so by courtesy of Hakan Ohlssons Boktryckeri, Lund; and Dr. Kjell Ander has very
kindly checked it for me, and corrected it in one particular. I have thought it better
to give as nearly as possible a literal translation, so that the English words can be
compared with those used by Thomson.
‘Note. The following colour-pattern is common to all species: the face together
with the cheeks and mouth-parts, a line on the pronotum in front of the tegula and
one on the lower margin, a mark on the scutellum and also the under side of the
antennal scape are white-yellow ; almost white are the coxae and trochanters of the
fore legs and also the median triangular marks on the hind margins of the dorsal seg-
ments, the femora are more reddish, the tarsi and tibiae sometimes of the same colour, —
the under side of the breast is in the female almost without exception yellowish and —
closely pilose. All the specimens of D. frontalis, which were found on a willow-bank
at Ortofta, bred from Nematus Westermanni show no variation in the yellow-white —
orbits of the frons nor in the almost rose-red hind margin of the second dorsal segment,
THE CTENISCINI OF THE OLD WORLD 417
wherefore I believe that the colour in these species is constant and can be used as
a basis for species-determination.’
Eudiaborus glutiniatus Roman
FINLAND, Kuusamo, Paanajarvi, Ruskeakallio, bog behind cliff, 1 9, 9.viii.1935,
ee
I give a figure of the under side of the apex of the female gaster (Fig. 76) to illustrate
the peculiar glutiniae (third valvulae) in this species.
Fic. 76. Apical segments of gaster, from beneath, of Eudiaborus glutiniatus Roman, illustrating
especially the glutiniae (third valvulae).
A. Roman told me he regarded this species as not uncommon in the Arctic, but
that he did consider his capture of it at about 62° latitude as remarkable.
Mason (1951, April: 228) has now recorded this as a North American species.
Eudiaborus boreoalpinus sp.n.
1940 Diaborus nigrifrons var. pallitarsis Hedwig, Z. Ent. Breslau, 18 (4) (non Thomson 1883).
Head (Fig. 77) more nearly cubical than in pallitarsis Thoms. and other palaearctic
species seen; seen from above about two-thirds length of its greatest width across
eyes, and sometimes as much as 1:5 times eye length behind eye; only slightly nar-
rowed behind eyes till near hind margin, but slightly more rounded in male; deeply
emarginate behind: on vertical region, temples and upper genae rather coarsely but
quite discretely punctate, on lower genae and upper face much more closely so ; lower
face coarsely and, in part, confluently punctate on an alutaceous background, some-
times a little smoother medially. Clypeus sparsely beset with punctures, fine to rather
coarse; fairly sharply and not deeply delimited; more or less weakly convex and
usually rather narrowly rounded apically. Hypostomal costa not so strongly bent
_ outwards as in pallitarsis Thoms. ; rather evenly raised to where met by genal costa,
a little farther behind mandibular base than width of malar space. Malar space about
equal to distance between mandibular apices, or a very little wider.
Antennae with scape of moderate proportions, rather more elongate than in palli-
_ tarsis Thoms., and with a rather sharp ventro-lateral excision (but some variation
in this): with flagellum rather slender, 24- to 28-segmented.
Pronotum with dorsal transverse furrow deeply and sharply impressed; with
ENTOM. It. 6 , 3F
418 A REVIEW AND A REVISION OF
epomiae of moderate strength, sometimes sharp; laterally moderately and rather
closely punctate above, rugose below. Propleura rather closely punctate, moderately
so behind, more coarsely in front. Mesoscutum moderately to strongly convex, but
not rising sharply from pronotum; with notaulices rather short, rather weakly or
moderately impressed ; moderately and unevenly punctate. Scutellum usually weakly
convex; finely and often rather sparsely punctate above, moderately so on sides;
not margined far beyond the broad scutellar fovea. Mesopleura moderately and
closely punctate, more coarsely and sparsely so in middle: sternaulices broad, shal-
lowly impressed: mesosternum less concave than in fallitarsis Thoms., rather finely
and very closely punctate, prepectus moderately and closely so: epicnemial carina —
almost rectilinear between notaulices, very little incised medially: mesolcus not
deeply impressed, more or less closed behind. Propodeum in greater part finely —
rugose in female, more coarsely so in male; with area externa finely or rather finely
punctate and conspicuously shining, and area pleuralis moderately and closely punc-
tate; with costae including costulae sharp and of moderate strength: area supero- —
media distinctly longer than broad, emitting costulae about in middle, not or not
strongly delimited basally.
Petiolar segment (Fig. 78) 1-4 to 1-5 times length of its greatest breadth in female,
I°5 to 1-7 times in male, unusually elongate and regularly explanate for this genus;
with basal lobes weak or scarcely apparent ; laterally weakly margined in female, more
strongly so in male ; with dorsal keels in female weak, distinct for about half length of
segment, longer and stronger in male; dorsally coriaceo-rugose, smooth at mid apex. —
Remainder of gaster shining: tergite 2 with diagonal basal furrows no more than
moderately impressed, moderately and closely though shallowly punctate between
them: gaster more and more finely punctate towards apex. Female hypopygium bent
at a definitely obtuse angle, and sharply keeled mid-ventrally beyond the bend (cf.
pallitarsis Thoms. in which it is bent at about a right angle and is not thence sharply
keeled). Female third valvulae of the same general type as in pallitarsis Thoms., but
more elongate, less broad, and less excavate beneath.
Fore wings with basal and disco-cubital veins not nearly parallel: with areolet
always distinctly petiolate; with nervus parallelus emitted well below middle of ©
brachial cell: radial cell 2-6 to 2-9 times length of its greatest breadth. Hind wings
with nervellus antefurcal and intercepted far below middle.
Legs elongate and slender ; notably the hind femora (Fig. 79), which are much more
so than in pallitarsis Thoms.: the males, however, less characteristic in this respect.
Tarsal claws regularly and rather strongly pectinate in female, but with much shorter —
comb-teeth in male.
Length 6 to 10 mm.
Female. Head black, with the following parts pale, bright flavous: mandibles
except at apex, clypeus, lower face except for a median mark in upper part, and genae
very broadly, the pale colour crossing the hypostomal costa: palpi more fulvous. The
pale colour does not extend up the inner orbits above the top of the antennal sockets:
the region of the clypeo-frontal suture is more or less distinctly dark-marked between
the tentorial pits. Antennae ferrugineous, merging to black at base above, paler
below; with scape pale, bright flavous below; with pedicellus similar, or usually at
THE CTENISCINI OF THE OLD WORLD 419
least conspicuously pale below. Thorax and propodeum black, with the following
bright flavous: lower angle of pronotum and margin some way forward thereof,
humeral angle and margin well forward thereof, propleura, marks below fore and hind
wings, rather small mid apical marks of scutellum and metascutellum, and lateral
flanges of scutellum: tegulae stramineous, scarcely suffused flavous: mesosternum not
pale-marked. Legs rufous, notably the hind tibiae, which are not at all blackened, or
only very weakly so at extreme apex ; with fore and mid femora and fore tibiae paler
<n ay, rg 78
oan O
Fics. 77-79. Eudiaborus boreoalpinus sp.n. Fig. 77. Head from above. The head is tilted farther
backward than in corresponding figures of species of other genera, otherwise the hind emargina-
tion would appear deeper. Fig. 78. Petiolar segment. Fig. 79. Left hind femur in sinistro-lateral
view.
in front and hind trochanters below: fore and mid coxae, trochanters and trochan-
terelli flavous ; with those of mid leg extensively, and fore coxae on outer side, rufous-
marked: tarsi somewhat darkened at apex. Wing veins pale ferrugineous; with
axillary sclerites pale flavous: stigma pale ochreous. Gaster black above, with sides
and venter flavous: tergites with pale flavous triangular marks at mid apex, increas-
ing in size from 1 to 5; and usually with very fine pale hind margins: third valvulae
darkened at apex.
Male differs as follows. Pale coloration distinctly not reaching level of top of
antennal sockets, and on gena much more restricted, not nearly reaching hypostomal
costa: lower face with a broad, median black band: clypeus broadly black basally
between tentorial pits. Thorax with tegulae and upper margin of pronotum coloured
as in female, but other pale markings absent or much reduced: in particular, the
scutellum is not pale-marked at mid apex except obscurely in one specimen, and the
propleura are black. Legs with coxae, trochanters and trochanterelli blackish in place
of rufous, and rather more extensively so, the fore trochanters blackish-marked
420 A REVIEW AND A REVISION OF
above: mid and hind femora sometimes blackish below near base: hind tibiae at apex
usually, and hind tarsi always, extensively darkened. Pterostigma sometimes darker.
Triangular pale marks sometimes undeveloped on tergites 1 and 2: venter blackish
in part. —
Described from the following material. SCOTLAND, Inverness-shire, Cairngorm Mts.,
3,000 ft., 1 9, 18.vili.1925, F. C. Best (type): SWEDEN, Jemtland, 1 2; Lapland, 1 9,
25.vii, C. Boheman: SWITZERLAND, Engadine National Park, I] Fuorn, La Drosa, 1 9,
10.viii.1921, Ch. Ferriére: Austria, Styria, Kreuzkogel southern slope, 1,800 m.,
I g, 7.viii.1950 ; Niedere Tauern, 2,000 m., I 9, 21I.vili.1g50, G. Heinrich: GERMANY,
Bavaria, Allgau, Osterachtal, 1,600 m., 2 gg, I 9, 8.viii.1949, G. Heinrich; Silesia,
Glatzer Schneeberg, I 3, 2.vili, I 9, II.vili.1g21, K. Hedwig (recorded as nigrifrons
Thoms. var. pallitarsis Thoms. as determined by A. Roman, Hedwig 1940).
Holotype 2 in British Museum (Natural History).
In the female sex, this species is very distinctive in the long, almost straight-sided
head ; the ventrally almost straight, scarcely incised epicnemial carina ; the shape of
the petiolar segment ; the long, slender hind legs with slender femora and clear red
tibiae, which are not blackened or scarcely so at extreme apex ; and the black meso-
sternum. The male is more weakly characterized.
This species is of particular interest on account of the distribution, which is of the
wide Arctic type (Benson, 1950: 93, 95).
Further material seen: see supplement, p. 454 below.
Genus CTENISCUS (Haliday MSS.), Curtis, Thomson
1832 Tryphon, subgenus Cteniscus (Haliday MSS.) Curtis, British Entomology, 9: 399.
1838 Tryphon, subgenus Cteniscus Haliday, Ann. Mag. nat. Hist. (1) 2: 113.
1883 Cteniscus Thomson, Opuscula Entomologica, fasc. 9: 881-882.
I began my work on this genus with a study of the two Haliday species that had
been proposed as genotypes. It was soon found necessary, not only to redescribe
curtisit Hal., which has been very little understood, but also to distinguish it more
critically from the species most closely allied to it. This work was then extended to
become a revision of most of the European ‘black-bodied’ species of the genus.
Among the red-bodied species, Roman (1913, 1931) has pronounced a number of
the forms, described as species by Holmgren and others, to be no more than races or
mere colour varieties of favomaculatus Grav. This synonymy, which has been largely
followed by the present and other authors, was, however, carried too far. Thus, the
majority of forms have the head rather strongly punctate and not narrowed behind
the eyes, whereas pumilus Hlgr. and limbatellus Higr. have the head very finely
punctate and shining, distinctly narrowed behind the eyes. Other good structural
differences may be found, to separate forms at present distinguished only by colour —
characters. A. Roman rearranged the collection of Swedish Ichneumonidae in the
Stockholm museum, but he did not omit to select lectotypes of the forms described
by Holmgren.
THE CTENISCINI OF THE OLD WORLD 421
Cteniscus hofferi Gregor
1858 Exenterus succinctus Holmgren, Svensk Vet.-Akad. Handl. 1: 230 (non Gravenhorst 1829).
1914 Cleniscus gnathoxanthus var. Roman, Ark. Zool. 9 (2): 16 (non Gravenhorst 1829).
1937 Cteniscus hofferi Gregor, Festschrift E. Sivand, Riga, 2: 416-417.
Head moderately narrowed and scarcely rounded behind eyes; rather deeply and
almost angularly emarginate behind: shining, with vertical region and temples moder-
ately, and cheeks generally finely punctate ; lower face more coarsely, and upper face
medially also more densely so, obsoletely alutaceous. Clypeus sparsely punctate;
rather narrowly rounded apically; rather deeply delimited, with tentorial pits con-
spicuous ; strongly convex in basal and shallowly impressed in apical half: in male it
is generally more evenly convex, and very shallowly impressed before the projecting
apex. Malar space shorter, in male much shorter, than distance between mandibular
apices. Hypostomal costa slightly raised to where it is met by genal costa, at a
strongly acute angle a good way behind mandibular base. Upper mandibular tooth
longer than lower.
Antennae with scape rather short, moderately excised externo-ventrally: flagellum
normally 27- to 31-, but in Holmgren’s small northern specimen only 22- segmented,
tapering gradually to apex.
Pronotum with dorsal transverse furrow narrow and sharply impressed; with
epomiae usually weak ; shining and moderately punctate above, more or less longitu-
dinally rugose below. Propleura shining and moderately punctate, anteriorly strongly
rugose. Mesoscutum rising sharply from pronotum, rather strongly convex; with
notaulices short but strongly impressed, giving it a distinctly trilobed appearance ;
discally often rather coarsely punctate, peripherally more finely so; between notau-
lices generally very weakly rugose. Scutellum generally rather strongly convex; |
sparsely punctate ; margined scarcely beyond scutellar fovea. Mesopleura closely and
rather strongly punctate, much more sparsely so above; with a shining speculum:
sternaulices very superficial: mesosternum towards mid line more sparsely and irregu-
larly, almost rugose-punctate, smooth and shining behind: prepectus strongly
punctate: epicnemial carina very slightly elevated: mesolcus moderately impressed,
narrow anteriorly: and widened posteriorly, scarcely closed by the transverse
rugosities behind. Metascutellum smooth and rather prominent. Propodeum shining
and very weakly punctate above, with the area pleuralis much more clearly and
densely punctate ; with costae including costulae moderately strong, in male strong
and sharp: area basalis not delimited apically: area superomedia usually longer than
broad, emitting costulae before middle.
Gaster stoutly spindle-shaped in female, behind petiolar segment almost parallel-
sided in male: petiolar segment very obscurely rugose, but with distinct longitudinal
rugosities by apical angles and smooth and very shining in mid third of hind margin:
remaining segments smooth and shining. Petiolar segment I-o to 1-3 times length of
its greatest breadth in female, 1-3 to 1-7 times in male: rather broad at the base, but
without basal lobes; moderately explanate and strongly margined ; with keels dis-
tinct for more than half length of segment, stronger and more nearly percurrent in
male ; with furrows between keels and margins more or less developed ; with a strong
422 A REVIEW AND A REVISION OF
lateral furrow leading to the glymma: spiracles situated well before middle, not
prominent. Tergite 2 with thyridia distinct but narrow in female, broader and more
conspicuous in male ; with diagonal basal impressions distinct in male, but superficial
and ill-defined. Female hypopygium almost pointed apically: third valvulae normally
not nearly reaching apex of tergite 8.
Fore wings with basal and disco-cubital veins not nearly parallel ; with areolet very
broad and shortly petiolate ; with nervus parallelus emitted in middle of brachial cell.
Hind wings with nervellus strongly antefurcal and intercepted far below middle.
Legs with the black hind tibiae relatively thick (Fig. 81). Claws of all tarsi strongly
pectinate.
Length 7 to 114 mm.
Female. Head black, with the following parts ivory. to pale flavous: genae except
near costa, lower face except for a median mark in upper part, whole of clypeus, often
a small orbital spot just below level of median ocellus, and most of mouth-parts
except mandibular apices. Antennae ferrugineous; basally darker, especially above.
Thorax and propodeum black, with the following parts stramineous to bright flavous
margined with fulvous: humeral angle of pronotum and upper margin some distance
forward thereof, whole of its lower margin at sides and a median anterior mark (these
two sometimes continuous), propleura except anteriorly and near mid line, tegulae,
line below fore wings, generally two pairs of small marks on prepectus, another before
bases of mid coxae, large quadrangular apical mark and lateral extensions of scutel-
lum, mark below hind wings extending down hind margin of mesopleura, and meta-
scutellum. Legs with fore and mid coxae, trochanters and trochanterelli stramineous
to flavous, suffused fulvous: fore and mid tibiae and tarsi, all femora, and hind coxae
and trochanters fulvous: hind tibiae and tarsal segments almost black, fulvous at
base, and often paler beneath ; hind tibiae with a pale sub-basal mark. Wings slightly
infumate, with veins and stigma fuscous to ferrugineous, the veins basally flavous
and the stigma basally luteous. Gaster black above, the tergites having pale apical
and lateral margins; stramineous to fulvous below, with black marks on some ster-
nites, but hypopygium pale: thyridia castaneous.
Male differs from the female as follows. Head with the pale parts more whitish
stramineous, but with the clypeus in apical half generally suffused fulvous. Antennae
deep ferrugineous, almost black ; scape pale flavous beneath, pedicellus and annellus —
pale beneath, postannellus pale at base and apex and next few segments at apex.
Lower margin of pronotum forming a complete pale collar. Hind coxae and tro-
chanters paler. Pale margins of tergites 1-3 triangularly broadened.
Redescribed from the following material: IRELAND, Co. Waterford, Dungarvan,
Maghgiha, I 9, 17.viii.1928 ; Co. West Mayo, Old Head, wood, at larva of Croesus on
Betula; Co. Wicklow, Glending Wood, 1 9, 15.ix.1938; Leitrim Barracks (Glen of
Imaal), 1 3, I1.viii.1940 ; Co. Dublin, Slade Brook, ex larva of Hemichroa crocea Geofir. |
(larva 21.ix.1941), I ¢ em. 6.viii.1942, A. W. Stelfox: ENGLAND, unlocalized, I g
(under curtisii Hal.), T. Desvignes coll.: BELGIuM, unlocalized, 1 3,5 22, C. Wesmael;
Sichem, 3 29, 14-15.ix.1886; Verrewinkel, 1 3, 15.ix.1848; Sysseele, I 2, 22.ix.1876; —
Carlsbourg, I 9, 9.ix.1898 ; Severin, I 9, 28.ix.1897, J. Tosquinet: SWEDEN, Skane,
Skaralid, 1 3g, 4.ix.1938, G. J. K.; Vasterbotten, Tavelsjé, 1 9, 4.ix, C. Boheman ~
;
s
a
0
THE CTENISCINI OF THE OLD WORLD 423
(recorded as succinctus Grav., Holmgren 1858): FINLAND, Lojo, 1 9, 1934, R. Kro-
gerus, I 2, W. Hellén; Karislojo, 1 9, R. Forsius: GERMANY, unlocalized, 1 2, Ruthe
coll. (under Tvyphon sexlituratus Grav.) ; G6ttingen, I 2, 13.ix.1946; Hannover, I 9,
13.ix.1948, R. Hinz: Austria, Pettenbach (S. of Wels), 1 9, 7.x.1941, Czerny: ITALY,
Piedmont, 2 29, 1859, O. Sichel.
Additional material seen: HOLLAND, Brummen, 3 99, vii, round Nematus larvae on
Sorbus aucuparia, Snellen van Vollenhoven (1872: 167, recorded as Exenterus suc-
cinctus Grav.).
Roman (1914) interpreted the unique female in the Holmgren collection as an
extraordinarily white variety of Ct. gnathoxanthus Grav. In later years he arranged
it in the Swedish collection of the Stockholm museum under brveviventris Thoms.,
which species is, however, placed otherwise in the present paper (p. 437).
Since going to press I have, through the kindness of Dr. Z. Bouéek, received the
type of hofferi Greg. on loan. I had no real doubt that the species was correctly
identified, but there were certain discrepancies in an otherwise good original descrip-
tion. It has been my frequent experience that subsequent micrometer measurement
belies an estimate of the length to breadth of a structure made by eye; and the same
difficulty must affect other authors, for in this specimen the length to greatest breadth
of both area superomedia and petiolar segment are I: 1-3 to the nearest o-r. As
suspected, ‘Weiss sind . . . Hiiften und Trochanteren II. und III.’ was a lapsus for
.l. und II.’ The head of the type had been lost, but the thorax and abdomen and
their appendages conform to my description and agree closely with specimens from
which it was made.
Biology. As recorded above, A. W. Stelfox took a parasitized larva of Hemichroa
crocea Geoffr. on 21 September 1941, and bred an adult male parasite, which emerged
in early August the following year. He also found the adult female parasite investiga-
ting Croesus larvae on birch.
R. Hinz took an adult female on 13 September 1948, and sustained it with sugar
water. It attacked a larva of Avge ustulata L. but laid no egg. It attacked a larva of
Hemichroa crocea Geofir., and so wounded it that body fluid oozed out, but again no
egg was laid. When presented with larvae of Croesus septentrionalis L. and latipes
Vill., it oviposited on them without further ado on the writing-table. The main
breeding experiments were carried out, however, with three species of Nematus.
The procedure prior to oviposition was observed as follows: the ovipositor is bent
free from the sheaths, and the egg is pushed to the tip of it, while pulsating movements
of the abdomen take place ; the ovipositor then resumes the resting position, and the
insect is prepared for oviposition. The egg has no real stalk, but a crochet apparatus ;
and is affixed between the first and second or second and third body segments of the
host. Eggs were laid at intervals of about 10 minutes, sometimes longer. Thus on
19th September 1948, 8 eggs were laid between II a.m. and 12.15; while on the
15th, 10 eggs were laid altogether.
Eggs were obtained on 25 larvae of Nematus (Pteronidea) pavidus Lep., on 16 of
{ N. (Pt.) miliaris Panz., and on 11 of N. (Pt.) melanocephalus Hart. The larvae hid
away in preserving tins containing some sand. From the pavidus larvae, 24 cocoons
were obtained ; but these were only found to contain remains of the host larvae. The
424 A REVIEW AND A REVISION OF
same kind of result was obtained with the miliaris. By contrast, the batch of melano-
cephalus gave 10 cocoons; and from these there were hatched in due season 2 adult
females of the sawfly and 8 adult males of the Cteniscus. Considering the difficulty
of obtaining the right conditions for rearing sawfly larvae, Herr Hinz does not wish
to base any facile conclusions on these results; but he observes that the melano-
cephalus is the largest larva of the three Nematus species used.
Cteniscus lineiger Thomson
1858 Exenterus exstirpatorius Holmgren, Svensk Vet.-Akad. Handl. 1: 238 (non Gravenhorst
1820).
1883 oe lineiger Thomson, Opuscula Entomologica, fasc. 9: 894.
Head moderately narrowed and rounded behind eyes ; about as long behind eyes as
eye length, seen from above, in female, shorter in male ; shallowly emarginate behind:
on vertical region, temples and upper face moderately and rather densely punctate ;
on sides of lower face less densely so, and on genae broadly much more finely and
sparsely punctate: epistoma rather coarsely punctate, obsoletely alutaceous. Clypeus
somewhat closely beset, mostly with moderate punctures, closely alutaceous near
mid apex; sharply delimited basally ; moderately convex in basal and broadly and
shallowly impressed in apical half; moderately rounded at apex. Malar space about
three-quarters distance between mandibular apices. Hypostomal costa very slightly
and evenly raised: sides of proboscideal fossa bent strongly outwards.
Antennae with basal segments very much as in Cé. gnathoxanthus Grav. (see descrip-
tion, p. 441 and Fig. 83): flagellum 24- to 27-segmented.
Pronotum with dorsal transverse furrow moderately impressed, coarsely trans-
carinate ; with epomiae sharp but short ; laterally moderately punctate above, exten-
sively longitudinally rugose below. Propleura moderately and closely punctate.
Mesoscutum rising sharply from pronotum, strongly convex; with notaulices rather
short, moderately impressed; for the most part shining and finely to moderately
punctate, but between notaulices closely punctate and obsoletely rugose. Scutellum
weakly convex, and to some extent buckled discally; not margined much beyond
scutellar fovea ; very finely punctate at sides and behind, almost impunctate discally.
Mesopleura rather coarsely and closely punctate below, but shining between the
punctures, not much more finely punctate above, and with a small, shining speculum:
sternaulices moderately and rather sharply impressed: mesosternum rather finely to
moderately and closely punctate, prepectus strongly so: epicnemial carina slightly
but distinctly elevated: mesolcus sharply impressed and narrow, broadened pos-
teriorly, partly closed behind. Propodeum very finely punctate and shining, in part
obsoletely rugose, with area pleuralis not much less finely punctate in greater part, —
distinctly rugose below; with costae sharp and of moderate strength: area postero-
media with a percurrent median keel: area superomedia distinctly broader than long,
distinctly delimited basally.
Gaster stoutly spindle-shaped in female, behind petiolar segment almost parallel-
sided in male: petiolar segment in greater part distinctly though usually weakly
rugose, smooth to a greater or lesser extent near mid apex: tergite 2 coriaceous at
THE CTENISCINI OF THE OLD WORLD 425
base between thyridia: remainder of gaster smooth and shining. Petiolar segment
I-r to 1-3 times length of its greatest breadth in female, 1-3 to 1-5 times in male;
rather strongly explanate and strongly margined; without basal lobes, but with
spiracular tubercles rather prominent ; with dorsal keels distinct for rather more than
half length of segment, and with a distinct median pit just behind them. Tergite 2
with thyridia narrow and inconspicuous in female, broader in male but less so than
in male of hofferi Greg. Female hypopygium blunt-pointed at apex: third valvulae
rather small, but stout and densely hairy, extending a little beyond apex of tergite 8.
Fore wings with basal and disco-cubital veins more nearly parallel than in hoffert
Greg., and both more nearly vertical ; with areolet broad and shortly petiolate ; with
nervus parallelus emitted below middle of brachial cell: stigma and radial cell rela-
tively broad, the latter 2-2 to 2-6 times length of its greatest breadth (in hofferi 2-6
to 3:0 times). Hind wings with nervellus antefurcal, intercepted well below middle.
Legs relatively stout, stouter than in rufonotatus Hlgr., but not so strongly ex-
panded in apical quarter as in hofferi Greg. Tarsal claws with pectination of moderate
strength, not extending to apex.
Length 6 to 8 mm.
Head coloured as in hoffert Greg., but lower face with a complete median black
band, and genae not pale to so near costa. Antennae with scape and pedicellus
blackish above, paler at apex, flavous marked below in female and more distinctly so
in male; with annellus and extreme base of postannellus more or less pale ; beyond
that ferrugineous above, paler below. Thorax and propodeum black, with the follow-
ing parts stramineous to bright flavous: pronotum as described for hofferi Greg., large
quadrangular apical mark and lateral extensions of scutellum and metascutellum,
tegulae, and marks below fore and hind wings: propleura usually only obscurely pale
on hind margin, never in greater part. Legs with fore and mid coxae, and all tro-
chanters and trochanterelli, mainly stramineous to pale flavous; with hind coxae
blackish at base, especially above: fore and mid femora, tibiae and tarsi pale rufous,
_ the femora flavescent towards apex: hind legs with femora rufous and tibiae and tarsi
dull rufous, the tibiae broadly towards apex and tarsi largely infuscate ; sometimes
_ with femora at apex and tibiae a little distance from base obscurely infuscate. Wings
_ with veins and stigma ferrugineous, pale at base. Gaster with petiolar segment black
_ in about basal two-thirds and at hinder angles, sometimes more extensively ; with
_ remainder of tergite 1, tergites 2 to 4 mainly, and sometimes 5 at mid base or more
_ extensively, rufous; with remainder of dorsum and with third valvulae blackish,
_ but tergites 2 to 3 or 4 with rather obscure flavous hind margins, and remaining ter-
_ gites margined ivory-white: female hypopygium blackish, sometimes partly rufe-
' scent, narrowly pale-margined: male with last three visible sternites similar, the first
_ of these more broadly pale-margined.
_ Described from the following material: BELGIuM, unlocalized, 1 9, C. Wesmael:
_ FINLAND, province Nyland (Uusimaa), Perna, 1 3, 3 99, A. Nordstrém ; Pargas, 1 9,
_ E. Reuter. Further material seen: SWEDEN, unlocalized, 2 gg, coll. Ljung (Thomson
coll.); Ostergétland, Mjélsefall, 1 3, 5.viii (type); Aby, 1 g, A. v. Goés. These
_ males are darker on gaster than as above described. Thomson’s female specimen
_ from PaAlsjo is missing.
ENTOM. II. 6 3G
426 A REVIEW AND A REVISION OF
Ct. lineiger var. nordstromi var. n.
Differs from the type form in having the gaster not largely rufous but black, apart
from the pale hind margins: tergites 2 and 3 have hind margins of a duller colour
yet more conspicuous against the black background, and tergite I is obscurely pale-
margined at apex: thyridia castaneous.
Described from the following material: FINLAND, Perna, 3 gg, I 2 (type), A.
Nordstrém. !
Recorded dates of capture of type form and variety range from 13.vii to 23.viii.
W. Hellén first sent two males of the black-bodied form of this species for my
opinion. I told him I believed the species to be new, and that I would place it nearest
to hofferi Greg., and gave him the main differences I found from that species.
Later he wrote me that he had since associated the female sex with the male, and
also a red-bodied form of the species with the black-bodied. He added that the captor,
Lektor A. Nordstrém, wished to present me with specimens, and that he himself
‘ would like me to describe the species in the course of my paper.
However, I placed the red-bodied form tentatively as exstirpatorius Hlgr. non
Grav.; and in 1952 I was able to confirm this by comparison with the type in
Stockholm and with two specimens in Thomson’s collection. But to Dr. Hellén
belongs the credit for being, as I believe, the first to associate real red- and black-
bodied forms of a European species of Cteniscus. The black-bodied form has been
captured exclusively by Lektor Nordstrém, and I have pleasure in naming this
seemingly distinct variety after him.
Cteniscus curtisii Haliday
1838 Tryphon (subgenus Cteniscus) curtisii Haliday, Ann. Mag. nat. Hist. (1) 2: 113.
1855 Exenterus sexcinctus Holmgren, Svensk Vet.-Akad. Handl., 1: 229-230 (non Gravenhorst
1829).
1871 ? poe sexcinctus Brischke, Schrift. Phys.-dkon. Ges. Kénigsberg, 11: 97.
1878 ? Cteniscus sexcinctus Brischke, Schrift. naturf. Ges. Danzig, 4: 104 (= sexcinctus var. I, —
Schmiedeknecht 1911: 2288).
1913 Cteniscus curtisii Roman, Ent. Tidskr. 32: 129.
1929 Cteniscus curtisii Gregor, Zpravy ¢sk. Stadt. Real. Gymn. Nové Jicin, 1929: 4-5 (separate
pagination).
Head somewhat strongly narrowed and, in larger specimens, scarcely rounded be-
hind the eyes; moderately and rather evenly emarginate behind; distinctly shorter
behind eye than eye length, seen from above: shining, with vertical region and temples
for the most part finely punctate, but near occiput and on cheeks less finely so;
upper face moderately and lower face rather coarsely punctate. Clypeus more or less
coarsely, rather shallowly and sparsely punctate; deeply delimited ; generally some-
what broadly rounded apically ; strongly convex in basal and more or less impressed in
apical half. Malarspacein female about three-quarters and in male about one-half dis-
tance between mandibular apices. Hypostomal costa slightly raised to beyond junction |
with genal costa, which is not very far behind mandibular base: gena with a slight
concavity outside this junction. Upper mandibular tooth slightly longer than lower.
Antennae with scape short, somewhat strongly convex dorsally and moderately
THE CTENISCINI OF THE OLD WORLD 427
excised externo-ventrally ; with post-annellus 1-4 to 1-5 times length of following
segment: flagellum 27- to 32-segmented (32 in type), slightly expanded in middle.
Pronotum with dorsal transverse impression sharp, transcostate; with epomiae
rather strong above; shining and rather finely punctate above, merging to trans-
versely rugose below. Propleura moderately and rather closely punctate, anteriorly
finely rugose. Mesoscutum moderately convex ; with notaulices strongly and sharply
impressed ; closely and for the most part moderately to rather coarsely punctate.
Scutellum not strongly convex, often rather flattened discally; sparsely punctate
above and on sides, more closely so behind; with margins not sharp much beyond
scutellar fovea. Mesopleura rather closely and coarsely punctate below, more finely
and much more sparsely so above; sternaulices broad and rather superficial but
quite distinct: mesosternum moderately and somewhat closely punctate in front,
more finely and sparsely so behind: prepectus closely and rather strongly punctate:
epicnemial carina slightly elevated: mesolcus moderately impressed and parallel-
sided for the greater part of its length, suddenly enlarged before the transverse costa
which partly closes it behind. Metascutellum smooth, normally raised. Propodeum
relatively strongly griseo-pilose and closely, finely to moderately punctate, with the
area pleuralis most distinctly punctate ; shining, but with some areas having obsolete
rugulosity, the area posteromedia very distinctly rugulose ; with costae rather strong
but costulae weaker, sometimes incomplete: area superomedia emitting costulae
behind middle, broadest at that point; relatively broad, with the almost straight
sides converging strongly from costulae to base; basally delimited.
Gaster stoutly spindle-shaped in female, very much less rounded in male: petiolar
segment, except near apex, coriaceous, or in part weakly rugose: gaster otherwise
smooth and shining. Petiolar segment of female broad, its breadth being approxi-
mately as great as its length (39: 39 in type, tending to be less in smaller and greater
in larger specimens) ; of male 1-2 to 1-3 times length of its greatest breadth; with
sides almost evenly rounded and not very strongly margined in female, the spiracles
not prominent ; with keels distinct to middle, to which it is strongly raised. Tergite 2
with thyridia rather narrow in female, broad and conspicuous in male ; with diagonal
basal impressions discernible. Female hypopygium pointed apically: third valvulae
extending distinctly beyond apex of tergite 8.
Fore wings with basal and disco-cubital veins not nearly parallel; with areolet
shortly or scarcely petiolate, not very broad, its outer vein almost interstitial with
the second recurrent ; with nervus parallelus emitted distinctly a little below middle
of brachial cell. Hind wings with nervellus antefurcal and intercepted rather far
below middle. ‘
Legs with the hind tibiae and tarsi very thick, thicker than in hoffert Greg. (Fig.
81), the hind metatarsus in inner side view about 54 times length of its greatest
breadth. Claws of all tarsi strongly pectinate, but more sparsely so in male.
Length 64 to 9 mm.
Head black, with the following parts bright stramineous, margined with rufous:
mandibles except at apex, clypeus except narrowly round base, and broad lateral
marks on the lower face extending through full width of the malar space down to the
genae: these marks extend on the genae from eye margin about half-way to costa;
428 A REVIEW AND A REVISION OF
and they leave the lower face black rather broadly below antennae to eye margin,
broadly down mid line, and, usually narrowly, along clypeo-frontal suture. Palpi
stramineous to rufous. Antennae dull ferrugineous, with basal segments usually
almost black except at their apices: sometimes rather paler. Thorax and propodeum
black, with the following parts bright flavous: humeral angle and mark in lower angle
of pronotum, a mark below hind wings, and quadrangular apical marks and hind
margins of scutellum and metascutellum: tegulae usually mostly rufous, sometimes
flavous. Legs red; with fore and mid legs in part, especially on trochanterelli, tend-
ing to dull flavous ; with hind tibiae and tarsi black, the tibiae with some infusion of
reddish colour, especially beneath and in smaller specimens, distinctly stramineous
at base and indistinctly fulvous at extreme base. Wings slightly infumate, with
veins and stigma ferrugineous, much paler at base. Gaster black above, the tergites
having pale apical margins narrow or very narrow: sternites usually mainly black,
the female hypopygium black with narrow pale margin: thyridia castaneous.
Redescribed from the following material: IRELAND, I 9, Haliday coll. (type):
Co. Wicklow, Meeting of the Waters below Rathdrum, I 9, 6. viii. 1925, A. W.
Stelfox: ENGLAND, 1 2, Desvignes coll. (previously undetermined), 1 9, Smith
coll.; Surrey, Oxshott, I 9, 6.vii.1892, Melville-Bury coll. (Manchester Museum) ;
Essex, Colchester, 2 92, 1903, 1914, Harwood ; Lancs. S., Chat Moss, I 9, 21.ix.1941,
H. Britten: FRANCE, I Q (probably France), de Gaulle; Corréze, Bort-les-Orgues,
I 9, 15-23.vi.1934, M. E. Mosely: Iraty, Piedmont, 1 9, 1859, O. Sichel: BELGium,
Vivier d’Oie, I 34, 27.v.1889, I 92, 4.viii.1880: Hestreux, 2 29, 5-9. viii.1884;
Liége, 2 99, 1862, coll. J. Tosquinet: HOLLAND, N. Brabant, Burgst, 1 g, vi.1913,
Smits van Burgst: GERMANy, Hamburg, Trittau, I 9, 7.vil.1945, G. Heinrich:
CZECHOSLOVAKIA, Chodau, 2 99, R. von Stein coll.; Ubutin, I 9, 7.viii.1g24, Fr.
Gregor: FINLAND, Parikkala, 1 9, W. Hellén.
Additional material seen: SWEDEN, Skane, Palsj6, 1 $; I 2 without data, Thomson
coll.; x 2 province Darlarne, 1 9 Lapland, Boheman: POLAND, Pomorzia, Sepolno,
I 2 25.vili.1939, G. Heinrich.
This species has been very little understood since its description, partly owing,
no doubt, to its apparent scarcity. It is hoped that the characters now given in the
key to species will suffice to separate it from its nearest relatives. As shown by
Roman (1913), it was misidentified by Holmgren as sexcinctus Grav., and it has stood
under that name in the Tosquinet and other collections. Characters given by Gregor
(1929) have been unavailable to those ignorant of the Czech language: having received
a translation of this paper, I am able to render the passage in question in the follow-
ing words: ‘abdomen black, with thin pale borders to the segments. Scutellum and
metascutellum posteriorly yellow-marked, with a white border going to the wings.
Mandibles with teeth of equal length (in contradistinction to Acrotomus sexcinctus
and succinctus). The face has two white trapezium-shaped spots touching the eyes
and connected with the white genae. Hypopygium black. Hind tarsi and tibiae
(except at base) black. Abdomen shining, finely punctate.’ The species placed as
curtisit in the Desvignes collection is a male of hoffert Gregor (q.v.). 7
Biology. Since the above was written, I have seen a specimen reared from Hemichroa
australis Lep. (= alni auctt. non L.): ENGLAND, Beds., Flitwick Moor, 1 2 ex larva
THE CTENISCINI OF THE OLD WORLD 429
taken 13.vii.1947 on Betula, em. 26.viii.1947, V. H. Chambers. As indicated in the
synonymic table, it is most probably this species, recorded as Cteniscus sexcinctus,
that was reared from ‘Dineura alni’ by Brischke (1871, 1878).
Cteniscus romani sp. n.
1871 ? Cteniscus sexcinctus var. Brischke, Schrift. Phys.-ékon. Ges. Kénigsberg, 11: 97 (non
Gravenhorst 1829).
1878 ? Cteniscus sexcinctus var. Brischke, Schrift. naturf. Ges. Danzig, 4: 104 (= sexcinctus var. 2
Schmiedeknecht 1911: 2288).
1948 Cieniscus curtist var. Teunissen, Tijds. Ent. 89: 13 (non Haliday 1838).
Head moderately to strongly narrowed behind eyes, moderately or rather shallowly
emarginate behind; about as long behind eyes as eye length, seen from above:
punctate on vertical region, temples and cheeks as described for curtisii Hal. ; on upper
face rather finely and on lower face moderately, sometimes in middle rather coarsely
punctate, on both much more sparsely so than in curtisii Hal., and with an impunctate
area above each antennal socket. Clypeus rather sparsely beset with punctures of
mixed sizes, fine to rather coarse; deeply delimited basally ; moderately convex in
basal half, and somewhat impressed before the rather narrowly to moderately rounded
apex. Malar space nearly three-quarters distance between mandibular apices in
female, about half such distance in male. Hypostomal costa very slightly raised,
bent distinctly outwards; its junction with genal costa farther behind mandibular
base than in curtisiz Hal.
Antennae with scape rather short, and with a sharp lateral excision (Fig. 84);
with postannellus about 1-4 times length of following segment in female, about 1-3
times in male: flagellum 28- to 33-segmented.
Thorax and propodeum structure differing from that described for curtisiz Hal. as
follows: pronotum narrower before dorsal impression: propleura rather sparsely
punctate: mesoscutum less coarsely and closely punctate: mesopleura rather finely
to moderately and sparsely punctate below, finely so above: sternaulices rather
weakly impressed: mesosternum less closely punctate: prepectus moderately and,
except at sides, sparsely punctate, more excavate than in curtisii Hal.: propodeum
having area externa more finely and area pleuralis more distinctly punctate; area
posteromedia smoother; costulae indicated but seldom developed, emitted from
about middle of area superomedia, which is relatively narrow, distinctly longer
than broad, with sides somewhat rounded and not strongly converging, and not
basally delimited.
| Gaster stoutly spindle-shaped in female, behind petiolar segment almost straight-
sided in male, its structure differing from that described for curtisii Hal. as follows:
petiolar segment mainly smooth, but to some extent obscurely rugose at sides; a
little longer, 1-1 to 1-3 times length of its greatest breadth in female, I-1 to 1-4 times
in male; more strongly narrowed to base; more strongly raised to middle, having
dorsal keels distinct not nearly to middle: basal diagonal impressions of tergite 2 more
evident: female hypopygium less pointed, and third valvulae projecting scarcely
beyond apex of tergite 8.
430 A REVIEW AND A REVISION OF
Structure of legs and fore wings as described for curtisit Hal.: hind wings with
nervellus strongly antefurcal and intercepted very low.
Length 64 to Io mm.
Head coloured as in curtisit Hal., but pale, bright flavous in place of stramineous,
and a little more extensively pale on genae: palpi also mainly flavous. Female
antennae with scape, pedicellus, and annellus black to ferrugineous, paler at apex and
beneath ; beyond annellus pale rufous, darkened in about apical half: male antennae
much duller. Thorax differs from curtisii Hal. in having scutellum and metascutellum
without conspicuous quadrangular pale marks at mid apex: scutellum obscurely dull
red at mid apex, bright flavous only at sides of hind margin, where metascutellum is
obscurely pale: tegulae scarcely rufous: fore and hind margins of mesopleuron and
sub-tegular mark often dull castaneous. Legs as described for curtisi: Hal. but fore
and mid legs more yellowed, especially in front ; hind femora often for the most part
much darkened, and hind tarsi often infused reddish. Wing colour as described for
curtisii Hal. Pale margins of tergites 6 and 7 and of female hypopygium rather
broader than in curtisi Hal.
Described from the following material: IRELAND, Co. Dublin, Slade of Saggart,
I 9, 16.ix.1942 ; Co. Wicklow, Deputy’s Pass, 1 g, 11.ix.1932, A. W. Stelfox: FRANCE,
Ardennes, Vendresse, I 3, 30.ix.1912, R. Benoist: BELGIuM, unlocalized, 5 29, coll.
Wesmael ; Vivier d’Oie, I 9, 9.x.1883, J. Tosquinet (type): HoLLAND, Tilburg, 1 9,
16.ix.1941, H. G. M. Teunissen (Teunissen 1948): GERMANY, Radbruch, I 9, 25.1x.
1938, R. Hinz; Holstein, Trittau, Hahnheide, 2 gg, 2 99, ix.1945, G. Heinrich:
SWEDEN, Smaland, Lommaryd, I 4, 3I.viii, 1 9, 12.ix. (mus. Stockholm, ? year and
collector): FINLAND, Sakyla Kolva, 1 2, W. Hellén; Lojo, 1 g, 1938, R. Krogerus.
Dedicated with respect and gratitude to the late Dr. A. Roman.
Cteniscus elegans Stephens
1835 Tryphon elegans Stephens, Illustrations of British Entomology, '7: 239.
1871 ? Cteniscus gnathoxanthus var. 2 Brischke, Schrift. Phys.-dkon. Ges. Kénigsberg, 11: 98 (non
Gravenhorst 1829).
1878 ? Cteniscus gnathoxanthus var. 2 Brischke, Schrift. naturf. Ges. Danzig, 4: 105 ( = Ct.
gnathoxanthus var. 2 Schmiedeknecht 1912: 2329).
1911 Exenterus elegans Morley, The Ichneumons of Great Britain, 4: 204-205.
1914 Cteniscus aulicus Roman, Ark. Zool. 9 (2): 14-16, syn.n.
1948 Cteniscus elegans Teunissen, Tijds. Ent. 89: 13.
The unique female type of Cteniscus aulicus Roman was taken by Boheman in the
southern Swedish province of Blekinge. Roman first labelled it ‘aulicus Rn n.sp.?
= fulvipes Krb. 9’. Evidently added at a later date is a second label in Roman’s ~
handwriting ‘? elegans Stph.’ Having with me a pair of Stephens’s species, previously —
compared with the type, I have been able to compare Roman’s type with them, and
can establish their specific identity.
Head slightly but rather sharply narrowed behind eyes, then rounded ; about as long
behind eyes as eye length, seen from above ; shallowly emarginate behind: on vertex,
temples and cheeks finely and rather sparsely punctate, more densely so near occiput
and lower genal margin: less finely and sparsely punctate than temples is the lower
THE CTENISCINI OF THE OLD WORLD 431
face, particularly on epistoma: upper face still more closely punctate, with strong
scrobal impressions, and usually with a distinct Y-shaped furrow below median
ocellus. Clypeus somewhat sparsely beset with punctures of mixed sizes, fine to
moderate or rather coarse, denser near basal margin; rather deeply and sharply
delimited basally; strongly convex in basal half, strongly impressed before the
reflexed and narrowly rounded apex. Malar space equal to distance between mandi-
bular apices in female, rather less in male. Hypostomal costa bent outward, strongly
and roundedly raised to where met by genal costa rather close behind mandibular
base, thence falling gradually to foramen magnum.
Antennae with scape moderately excised externo-ventrally: flagellum 27- to 20-
segmented, slightly expanded in middle, therefrom tapering gradually to apex.
Pronotum with dorsal transverse furrow moderately impressed, broader in middle ;
with epomiae weak or very weak; laterally rather finely punctate above, more or less
rugose or rugulose below. Propleura rather finely punctate, quite densely so at sides.
Mesoscutum rising sharply from pronotum, rather strongly or strongly convex in
front, but more flattened behind ; with notaulices sharply impressed ; very finely and
rather sparsely punctate discally, less finely and more densely so peripherally, and
especially in front. Scutellum moderately convex, somewhat flattened discally ;
finely and for the most part sparsely punctate; not margined beyond scutellar
fovea. Mesopleura moderately punctate below, more finely so above: sternaulices
weakly to moderately impressed: mesosternum and prepectus moderately punctate:
mesolcus normally impressed; posteriorly a little widened, and transcostate but
scarcely closed. Propodeum more or less rugulose, or on area posteromedia partly
rugose; on area externa finely, and on area pleuralis moderately and densely
punctate ; with costae normally complete, but mostly rather weak: area superomedia
about as broad as long, strongly contracted from costulae to base, generally (but not
in type) basally delimited.
Gaster spindle-shaped in female, behind petiolar segment parallel-sided or some-
what narrowed in male. Petiolar segment I-2 to 1-3 times length of its greatest breadth
in female, 1-4 to 1-6 times in male; with basal lobes distinct but not strong; not
strongly margined, with spiracles well before middle and not prominent; rather
strongly raised in middle, but not as much so as in curtisiit Hal.; with dorsal keels
moderate or rather strong, distinct to beyond middle ; smooth at mid apex and be-
tween keels; very obscurely rugose beside the keels, finely and distinctly so near
apical angles. Remainder of gaster smooth and shining: tergite 2 with thyridia
distinct, very narrow in female, more-crescentic in male. Female hypopygium much
as in hoffert Gregor: female third valvulae more pointed than in that species, reaching
to about apex of tergite 8.
Wings as described for hoffert Gregor, but nervellus not so strongly antefurcal.
Claws of all tarsi rather strongly pectinate.
Length 7 mm.
Head black; with palpi, mandibles except at apex, and clypeus stramineous-
white: the palpi and mandibles are to some extent suffused fulvous, and the clypeus
is fulvous along extreme apical margin. Antennae with scape and pedicellus black or
almost so, marginally paler; with annellus and extreme base of postannellus pale ;
432 A REVIEW AND A REVISION OF
with remainder of flagellum dull fulvous below, ferrugineous above, the basal seg-
ments apically paler. Thorax and propodeum black, with lower angles of pronotum
pale ; with humeral angles of pronotum and with tegulae stramineous ; with scutellum
and metascutellum stramineous to flavous at apex, discally rufo-fulvous. Legs with
all coxae and’ femora bright fulvous, the femora paler at apex: trochanters and tro-
chanterelli stramineous, the hind trochanters basally fulvous: fore and mid tibiae and
tarsi more or less pale dull fulvous, the tarsi dark at extreme apex: hind tibiae broadly
at apex and tarsi ferrugineous, the tibiae mainly dull fulvous, broadly dull strami-
neous at base. Wings with veins and stigma ferrugineous, basally stramineous.
Gaster coloration as described for hofferi Gregor.
Redescribed from the following material: ENGLAND, Kent, Darenth Wood, type
Q, J. F. Stephens; Devon, Barnstaple, 1 9, T. A. Marshall coll. ; Staffs., Lichfield
dist., 1 g, I 2, 1922, L. A. Carr: GERMANY, Koln, 1 9, 13.v.1940, W. Aerts: CzECHO-
SLOVAKIA, Moravia, Pouzdfany, 6.vi.1937, Fr. Gregor.
Additional material seen: SWEDEN, Skane, Esperéd, I 34, 20.vi, I 2 (probably
also from Skane), C. G. Thomson; Blekinge, 1 2, C. Boheman (type of aulicus
Roman). Recently recorded from Holland (Teunissen 1948).
Brischke’s Cteniscus gnathoxanthus var. 2’, recorded from the Danzig district
(1878), may very well be this species. It was reared from Periclista RP USERS Zadd.,
a sawfly species attached to Quercus.
Cteniscus pectoralis Hellén
1951 Cteniscus pectoralis Hellén, Notul. ent., Helsingf. 31: 31.
Head rather strongly transverse, a little shorter behind eyes than eye length, seen
from above; very slightly narrowed behind eyes and then rounded ; rather shallowly
emarginate behind: on vertical region and temples for the most part very finely and
sparsely punctate, but near occiput and on cheeks less finely and sparsely so, and by
lower genal margin moderately alutaceo-punctate: lower face moderately to rather
coarsely and densely punctate: upper face rather finely and densely so, with trans-
verse rugosities just above antennal sockets, and with a distinct inter-antennal keel.
Clypeus rather sparsely beset with punctures of mixed sizes, fine to rather coarse ;
sharply but not deeply delimited basally ; strongly projecting though weakly convex ;
scarcely impressed before the moderately rounded apex. Malar space (female)
clearly greater than distance between mandibular apices. Hypostomal costa more
than slightly but not strongly raised ; met by genal costa at about a right angle, as
far behind mandibular base as length of malar space.
Antennae with scape somewhat strongly convex dorsally, about as much so as in
curtisii Hal. but more elongate, moderately excised externo-ventrally (both broken
off beyond postannellus in unique type).
Pronotum with dorsal transverse furrow sharply impressed, bisected by a median
keel; with epomiae short and weak; laterally finely punctate and shining before
humeral angle, but in greater part obsoletely rugose. Propleura shallowly, moderately
to rather coarsely and densely punctate. Mesoscutum moderately convex; with
notaulices strongly and sharply impressed ; shallowly but coarsely and closely punctate
THE CTENISCINI OF THE OLD WORLD 433
and obsoletely rugose between notaulices, merging to much more finely punctate
peripherally and behind. Scutellum rather strongly convex; very finely punctate;
not margined beyond scutellar fovea. Mesopleura rather finely to moderately and
not closely punctate; with a large impunctate speculum: sternaulices very super-
ficial: mesosternum moderately punctate in front, more finely and sparsely so behind:
prepectus moderately punctate: epicnemial carina indicated but incomplete medially:
mesolcus moderately impressed, greatly broadened and with several transverse
costae but not closed behind. Propodeum for the most part obsoletely rugose, but
with areae superomedia, dentipara, and posteromedia near mid apex, quite strongly
so ; with areae externa and pleuralis finely punctate ; with costae quite strong at sides
and behind, but weaker and irregular above, including the costulae: area postero-
media not reaching middle: area superomedia about as long as broad, not delimited
basally.
Petiolar segment elongate, 1-6 times length of its greatest breath; with a narrow
waist near base and then strongly explanate (rather as in Eclytus fontinalis Higr. but
not strongly contracted again just before apex); sharply margined laterally; with
dorsal keels strongly raised, sharp to about level of spiracles but continuing blunter
to about last quarter of segment, bounding a median longitudinal pit and falling
sharply to lateral furrows; finely rugulose, smooth near mid apex. Remainder of
gaster elongate and slender: tergite 2 a little expanded to apex, with thyridia small
but distinct: remaining tergites contracted to apex, tergites 3 to 5 each the length
of its basal breadth: tergites 2 and 3 finely alutaceo-punctate, the remainder very
finely punctate and shining. Female hypopygium elongate, folded mid-ventrally but
not sharply so: female third valvulae elongate, narrow in ventral view, somewhat
pointed in side view, strongly hairy, extending far beyond apex of tergite 8.
Fore wings with radial cell almost three times as long as broad; with basal and
disco-cubital veins almost parallel; with areolet shortly petiolate, not broad; with
nervus parallelus emitted below middle of brachial cell. Hind wings with nervellus
strongly antefurcal, and intercepted well below middle.
Legs, notably hind femora and tibiae, very elongate and slender. Claws of hind
tarsi each with three, rather small, widely spaced comb-teeth.
Length 114 mm.
Head black, with the following parts stramineous, tinged with flavous: labium and
maxillae, clypeus, and lower face continuous with genae broadly: there are fulvous
upper margins to pale part of genae and lower face, and a small median mark of the
same colour on upper part of lower face: mandibles more ochreous, black at apex.
Antennae with scape, pedicellus and annellus pale ferrugineous to castaneous above,
merging to deep flavous below: postannellus deep ferrugineous. Thorax and propo-
deum black, with the following rufo-fulvous above, pale dull flavous below: pro-
notum, scutellum, metascutellum, mark above epicnemial carina extending down
prepectus, and mark above mid coxa, these last two faintly connecting across meso-
pleuron: propleura pale flavous, tinged with fulvous: humeral angle of pronotum and
mark below fore wing pale bright flavous, tegulae duller: mark below margin of scutellar
fovea and marks on propodeum dull rufous: hind margins of mesopleuron, scutellum
and metascutellum obscurely pale. Fore and mid coxae, trochanters and trochan-
ENTOM. Il. 6 3H
434 A REVIEW AND A REVISION OF
terelli coloured as propleura: fore and mid femora fulvous, obscurely paler at base
and apex, and their tibiae and tarsi paler: hind coxae rufous, as also hind trochanters
and trochanterelli in inner side view and below; but the last two infuscate in outer
side view and pale at base and apex: hind femora, tibiae, and tarsi deep ferrugineous ;
with femora and first four tarsal segments pale at extreme base and apex; with
tibiae pale at base and extreme apex; with fifth tarsal segment, and fourth below,
pale rufous. Wings with veins and stigma ferrugineous, the stigma very pale at base.
Gaster black above, stramineous to pale flavous below: tergites 1 to 6 with narrow,
pale apical margins and tergite 7 with an obscure, triangularly broadened one:
third valvulae pale, partly infuscate above, and rufous towards apex.
Redescribed from the unique holotype, FINLAND, Esbo, 1 9, R. Frey.
Most closely related to Cz. unicinctus Hlgr. in the elongate, slender body, and parti-
cularly the legs; in the ventrally folded (though not sharply folded) hypopygium ;
and in the pale pectoral coloration. It differs notably from that species, however, in
having the head slightly shorter behind eyes than eye length, seen from above; and
is unique among known palaearctic species in having the gaster so elongate that
tergites 4 and 5 are as long as their breadth at base. As to whether the imperfectly
formed epicnemial carina is a good specific character or a mere individual variation,
remains to be seen.
I wish to thank Dr. W. Hellén for his close co-operation in allowing me, before his
description had appeared in print, to make a study of his very distinct and interesting
species for inclusion in my paper.
Cteniscus unicinctus Holmgren
1858 Exenterus unicinctus Holmgren, Svensk Vet.-Akad. Handl. 1: 234.
1858 Exenterus macrocephalus Holmgren, ibid.: 243 (synonym by Roman 1914).
1896 Cteniscus macrocephalus Jacobs & Tosquinet, Aun. Soc. ent. Belg. 34: 108.
1914 Cteniscus unicinctus Roman, Ark. Zool. 9 (2): 17.
1942 Cteniscus unicinctus Kerrich, Trans. Soc. Br. Ent. 8 (2): 63-64.
Head much longer behind eyes than eye length, seen from above; distinctly
broadened and rounded behind eyes, in well-developed specimens strongly so;
somewhat strongly and sometimes rather angularly emarginate behind: on vertical
region, temples and inner orbits finely to rather finely and rather sparsely punctate, on
upper face closely so; genae and epistoma rather finely to moderately and closely
punctate. Clypeus strongly transverse, more than twice as broad as long; fairly
closely beset with punctures of different sizes, fine to rather coarse; not deeply de- —
limited basally ; weakly convex before the slightly reflexed and very broadly rounded
apex. Malar space a little less than distance between mandibular apices in female,
a little more than half this distance in male. Hypostomal costa bent strongly out-
ward, very slightly and evenly raised ; met by genal costa, at nearly a right angle, as
far behind mandibular base as distance between mandibular apices. Mandible with
lower tooth more prominent than upper, rather in the manner of Acrotomus and Antso-
ctenion, but always a little shorter.
Antennae with scape (Fig. 82) strongly convex dorsally, rather weakly excised
externo-ventrally ; with pedicellus relatively elongate ; with postannellus about one-
THE CTENISCINI OF THE OLD WORLD 435
eighth longer than following segment ; with flagellum filiform, decidedly slender, 23-
to 26-segmented.
Pronotum strongly produced forward, as in Anisoctenion alacer Grav. ; with dorsal
transverse furrow strongly impressed, broad and shining ; with epomiae of moderate
strength, sharp, not shortened below; laterally rather finely and closely punctate
above, with some rugosity developed below. Propleura finely punctate ; with outer
hind angles strongly produced and reflected. Mesoscutum rather strongly convex ;
with notaulices of moderate strength, extending well back on to disk; very finely
punctate or, to a greater or lesser extent, not so finely so. Scutellum moderately
convex ; finely and closely punctate above and behind, very finely and sparsely so
at sides; not margined beyond scutellar fovea. Mesopleura finely punctate above,
rather finely to moderately and not closely so below (never so closely as in curtisit
Hal.): sternaulices rather shallow, but broad and very distinct: mesosternum finely
and closely punctate; prepectus rather coarsely and very closely so, bearing a
pair of short keels just behind lower angles of pronotum: epicnemial carina quite
distinctly though not strongly elevated: mesolcus sharply impressed, partly closed
behind. Propodeum relatively elongate and not abruptly declived; with pilosity
long but not thick; generally finely punctate on area externa, elsewhere very finely
so, shining, seldom with much rugulosity ; with costae sharp at sides and behind and
with costulae complete: area superomedia about one and a half times as long as
broad, or longer, not basally delimited.
Petiolar segment elongate, 1-8 to 2-3 times length of its greatest breadth ; almost
regularly expanded from near base to near apex, though sometimes the spiracles are
prominent ; with dorsal keels sharp, approaching rather closely before and between
spiracles, distinct for about two-thirds length of segment ; very obscurely rugose be-
tween and beside keels, smooth and shining behind. Remainder of gaster smooth
and shining: tergite 2 with thyridia distinct. Female hypopygium sharply folded
medially in apical half: female third valvulae elongate pilose, projecting beyond
apex of tergite 8, small, varying considerably in shape.
Fore wings with basal and disco-cubital veins not strongly divergent ; with areolet
sessile to shortly petiolate ; with nervus parallelus emitted a little below middle of
brachial cell. Hind wings with nervellus distinctly antefurcal and intercepted far
below middle.
Legs elongate and slender. Tarsal claws with short, fine pectination, weaker in
male.
Length 7 to 8} mm. in specimens redescribed, but down to 4 mm. in a female
specimen from Lapland.
Female. Head colour as described for Exyston genalis Thoms. (p. 376), but pale
coloration sometimes extending on to upper face right round antennal sockets.
Antennae bright ochreous; with scape and pedicellus pale beneath, ferrugineous
marked at sides; with annellus pale ; with following few segments ferrugineous, pale
at apex. Thorax and propodeum black, with the following parts stramineous to pale
flavous, to some extent flushed with rufous: tegulae, pronotum very largely, pro-
pleura and mesosternum totally, mesopleura below tegulae, along front and hind
margins, and largely below; often pale-marked also on metascutellum and propodeum:
436 A REVIEW AND A REVISION OF
scutellum usually distinctly but sometimes rather obscurely pale-marked, near mid
apex or more extensively on disk, at sides of hind margin, and below margins of
fovea. Similarly pale-coloured are the coxae, trochanters, trochanterelli, and base
and apex of hind femora: legs otherwise more testaceous, but with hind femora
mainly ferrugineous, hind coxae above and usually hind tibiae above and at apex
marked with the same colour. Wing colour as described for C¢. elegans Steph. Gaster
mainly black above, flavous to testaceous below, including the hypopygium, and
sometimes also the whole of tergites 6 to 8: tergite 1 with a median pale mark very
near apex ; and remaining black tergites with pale apical margins, which are usually
triangularly produced forward conspicuously on 2 and 3, and sometimes more ob-
scurely on 4: thyridia small, but pale and conspicuous.
Male differs as follows: pale coloration of head just reaching up to antennal sockets,
and extending to a line going from a point on hypostomal costa rather close behind
mandibular base to a point about a quarter the way up outer orbit. Antennal
flagellum darker, sometimes in greater part ferrugineous. Pronotum black, except
for lower and humeral angles: propleura usually only marginally pale: mesopleura
below and mesosternum black, except sometimes for margin of prepectus and a Y-
shaped mark by sternaulices: scutellum and metascutellum not or obscurely pale-
marked in middle, and propodeum unmarked. Hind tibiae and also hind tarsi above
often more strongly ferrugineous marked: hind coxae sometimes mainly ferrugineous.
Black coloration extending nearer lateral margins of tergites 3 and following, and
tergites 6 and 7 only very narrowly pale at apex: in one southern Swedish specimen
the pale marks on tergites 2 and 3 greatly reduced; and, according to Holmgren
(p. 234), only the second tergite conspicuously pale-banded in type material from
Lapland.
Redescribed from the following material: ENGLAND, Hants, New Forest, 19,
vi.1902, D. Sharp: FRANCE, Paris district, Maisons Laffitte, 1 J, Forét de St. Germain,
I 9, 11.v., Fontainebleau, I 9, 12.vi.1g00, J. de Gaulle: BELGIum, Groenendaele, I 9,
27.vi.1892, I 9, 8.vii.1880, J. Tosquinet: GERMANY, Berlin district, 1 9, 1.vi.1856,
Ruthe coll.: SWEDEN, Skane, H6G6r district, 2 ¢¢, 8.vi, I g, 10.vi.1938, D. M.S. &
J. F. Perkins.
Further material seen: SWEDEN, type and other specimens in Royal Natural History
Museum, Stockholm (see Roman, 1914, Kerrich, 1942).
Cteniscus consobrinus Holmgren
Head shape in dorsal view as described for wnicinctus Hlgr.: punctation much as
described for that species, but in general somewhat finer and denser. Eyes in male
slightly convergent. Clypeus as described for wnicinctus Hlgr., but deeply delimited
basally, and in male not very broadly rounded at apex. Malar space about equal to
distance between mandibular apices in both sexes. Hypostomal and genal costae as
described for unicinctus Hlgr., but meeting at much less than a right angle. Mandible
with lower tooth in same plane as and distinctly shorter than upper.
Antennae with scape of normal proportions, rather weakly excised externo-
ventrally; with postannellus about 1-3 times length of following segment; with
flagellum not relatively slender, 24- to 25-segmented.
THE CTENISCINI OF THE OLD WORLD 437
Pronotum much less produced forward than in wnicinctus Hlgr., sharply declived
to the dorsal transverse furrow; with epomiae rather weak, evanescent below;
laterally finely and not very distinctly punctate above, strongly rugose below.
Propleura as described for uwnicinctus Hlgr. Mesoscutum rising sharply from pro-
notum, moderately convex ; with notaulices rather strongly impressed but short, not
quite reaching anterior margin; finely to very finely and obscurely punctate.
Scutellum moderately convex; very finely punctate; margined a little beyond
scutellar fovea. Mesopleura finely and not closely punctate below, sparsely and very
finely so above: sternaulices broad but superficial: mesosternum very finely punctate,
prepectus rather finely so: epicnemial carina slightly elevated: mesolcus shallowly
impressed anteriorly, deeply so posteriorly, partly closed behind. Propodeum rugulose-
punctate, but with area pleuralis in large part shining ; with costae including costulae
sharp: area superomedia distinctly longer than broad, almost parallel-sided, not dis-
tinctly delimited basally, emitting costulae before middle.
Petiolar segment 1-4 times length of its greatest breadth in type female, 1-7 to
2:0 times in male ; with dorsal keels of moderate strength, distinct to beyond middle ;
dorsally coriaceo-rugose, smooth only near mid apex. Tergite 2 weakly coriaceous
at base; with thyridia distinct. Remainder of gaster smooth and shining. Female
hypopygium pointed but not sharply folded: female third valvulae broad, rather
strongly pointed, projecting beyond apex of tergite 8.
Fore wings as described for wnicinctus Hlgr.: hind wings with nervellus less ante-
furcal and intercepted not so far below middle.
Tarsal claws with pectination of moderate strength in female, weaker in male.
Length 7 to 74 mm.
Head coloured as in Ct. romani nov. (p. 430). Antennae dull ferrugineous, some-
what paler beneath; with scape, pedicellus and basal flagellar segments paler at
apex; with scape and pedicellus beneath and with annellus conspicuously pale.
Thorax and propodeum black, with the following pale: humeral angles of pronotum
and tegulae distinctly, hind margin of scutellum at sides and mark below hind wing,
and outer hind angle of propleura obscurely. Legs pale rufous ; with trochanters and
trochanterelli, and fore and mid coxae at least in part, pale flavous; with coxae
otherwise darker, the hind coxae largely blackish, and with hind trochanters dark-
marked above. Wing colour as described for Ct. elegans Steph. Gaster colour as
_. described for Ct. gnathoxanthus Grav. except for female hypopygium which is
EEE —_
ferrugineous, pale at mid base and with narrow pale margin.
Redescribed from the following material: IRELAND, Co. Wicklow, Glen of Imaal,
I 3, 27.v.1926, A. W. Stelfox: SwEDEN, Lapland, I 9, 25.vii (holotype), 1 ¢ (allo-
type), C. Boheman.
Cteniscus rufonotatus Holmgren
1858 Exenterus rufonotatus Holmgren, Svensk Vet.-Akad. Handl. 1: 243.
1883 Cteniscus breviventris Thomson, Opuscula Entomologica, fasc. 9: 890, syn. n.
1896 Exenterus fulvipes Kriechbaumer, Ent. Nachrichten, 22: 369, syn. n.
Head strongly transverse, a little shorter behind eye than eye length, seen from
above; rather strongly or strongly narrowed and rounded behind eye; shallowly
438 A REVIEW AND A REVISION OF
emarginate behind: on vertex, temples, cheeks, and inner orbits very finely punctate,
for the most part rather sparsely but on cheeks quite sparsely so; on upper face
finely, very seldom in part closely punctate; on epistoma rather finely and not
closely punctate. Clypeus sparsely beset with rather coarse punctures; sharply
delimited basally ; strongly convex to near the reflexed and more or less narrowly
rounded apex. Malar space about five-eighths distance between mandibular apices.
Hypostomal costa slightly and almost evenly raised, met by genal costa not close
behind mandibular base. Mandibles with lower tooth distinctly a little longer and
more prominent than upper, suggesting the condition in genus Anisoctenion.
Antennae with scape as described for gnathoxanthus Grav. ; with flagellum rather
stouter and more coarsely hairy, as in gibbulus Hlgr., 21- to 25-segmented.
Pronotum with dorsal transverse furrow strongly and quite sharply impressed,
distinctly transcarinate ; with epomiae rather strong; laterally finely or very finely
punctate above, weakly to more or less strongly rugose below. Propleura finely and
rather densely punctate. Mesoscutum rising sharply from pronotum, rather strongly
convex, though somewhat flattened on disk; with notaulices rather strongly and
sharply impressed ; for the most part very finely punctate, more sparsely so on disk.
Scutellum as described for gnathoxanthus Grav. Mesopleura rather closely beset with
moderate punctures below, finely punctate above, and with speculum extending
far forward: sternaulices usually ill-defined and rather superficial: mesosternum
finely and closely punctate in front, more finely and sparsely so behind, its halves
distinctly convex: prepectus closely punctate, distinctly excavate: epicnemial
carina slightly elevated: mesolcus strongly impressed, wider and deeper posteriorly,
partly closed behind. Propodeum strongly raised to base of area superomedia;
shining and for the most part obsoletely rugose in front, quite strongly rugose behind,
including about half area pleuralis ; with costae including costulae strong: area postero-
media with median keel usually strong and percurrent: area superomedia a little
longer than broad, emitting costulae about in middle, strongly contracted therefrom
to base, where it is more or less distinctly delimited.
Gaster stoutly spindle-shaped in female, with sides decidedly rounded even in
male. Petiolar segment of female 1-1 to 1-2 times length of its greatest breadth;
strongly narrowed to base ; without basal lobes, but with sides almost evenly rounded
and sharply margined, the spiracles seldom at all prominent ; dorsally weakly rugose
to coriaceous or even smoother, smooth near mid apex ; moderately raised in middle,
with dorsal keels distinct for about three-quarters length of segment: petiolar seg-
ment of male about 1-3 times length of its greatest breadth ; with spiracular tubercles
prominent ; dorsally more distinctly rugose, with keels sharp and generally almost
percurrent. Remainder of gaster for the most part smooth and shining; tergite 2,
however, slightly elevated in about basal half between thyridia ; often weakly coria- —
ceous upon this elevation, occasionally more strongly or extensively sculptured. —
Female hypopygium broadly rounded at apex, appearing almost truncate in middle
when sides are folded: female third valvulae narrow (Fig. 86), projecting slightly
beyond apex of tergite 8.
Wings as described for curtisii Hal. but areolet may be longer petiolate: stigma and ~
radial cell broad, the latter about 2-1 times as long as broad.
THE CTENISCINI OF THE OLD WORLD 439
Hind legs relatively stouter than in guathoxanthus Grav. and gibbulus Hlgr.; but
tibiae not so much thickened as in hoffert Greg. (Fig. 81), and metatarsi not thickened.
Tarsal claws rather weakly and sparsely pectinate, more weakly so in male.
Length 44 to 7 mm.
Female. Head black, with the following parts bright stramineous, margined with
rufous: palpi, mandibles except at apex, clypeus, and broad marks on genae: these
approach, or more often traverse, the hypostomal costa just behind lower articula-
tion of mandible, and. they extend through malar space usually up to the tentorial
pits, and thence triangularly about half way up orbits of lower face. Occasionally
the lower face is colour-patterned as in curtisii Hal., or less extensively. Antennae
rufous to ferrugineous, paler towards apex and below. Thorax and propodeum black,
with the following parts stramineous to flavous: humeral angle of pronotum and mark
in lower angle, which extends some way forward, propleura, and tegulae: hind margin
of scutellum and metascutellum at sides, and mark below hind wing obscurely pale:
scutellum sometimes dull castaneous behind. Legs mainly clear red, to some extent
rather paler ; with fore and mid coxae, except at base, trochanters and trochanterelli
flavous ; with hind tarsi darkened extensively but hind tibiae at apex only ; sometimes
with hind femora at apex and hind coxae blackish. Wings distinctly infumate ; with
veins and stigma pale ferrugineous, much paler at base. Gaster black above, the
tergites with yellowish apical and lateral margins: tergite 2 sometimes, as in Holm-
gren’s type, with a more or less triangular red mark and with thyridia castaneous:
venter mainly pale, the sternites with large, paired dark marks; the hypopygium
dark, with narrow pale margin.
Male differs as follows: pale mark of gena and lower face less extensive. Propleura
pale at apex only.
Redescribed fromthe following material: IRELAND, Co. Wicklow, Rathdrum,
I 9, 13.vili.1g25, A. W. Stelfox: BELcrum, unlocalized, 4 99, C. Wesmael; Genck,
2 29, 25.vili.1886, I 9, 21.viii.1892, Sichem, 2 $¢ (one the type of fulvipes Kriechb.),
I 2 (hereby selected as allotype of fulvipes Kriechb.), 19.ix.1885, Schaffen, 2 99,
18.ix.1885, I 9, 10.ix.1886, Molenstedt, 299, 23.ix.1886, Cagervine, I 9, 25.ix.1885,
J. Tosquinet: SWEDEN, Skane, Torekov, I 4, vii.1860 (lectotype of breviventris
Thoms.), Raften, 1 9, 12.viii (allotype of breviventris Thoms.), ? C. D. E. Roth,
C. G. Thomson coll.; Smaland, 1 9 (type), C. Boheman: SWITZERLAND, Geneva,
Peney, I 9, 15.viii.1884, Tournier; Champel, 1 9, coll. Buess: Spain, Orviedo,
Gijon, I g, viii.1900: Russia, Dniepropetrovsk, I 3g, 7.1x.1943, R. Hinz.
Material of this species stood in the Brussels museum collection under the Holmgren
and the Kriechbaumer names, and included the Kriechbaumer type: the two series
were associated, and the synonymy was confirmed by examination of the Holmgren
type. The clue to synonymy of Thomson’s name was given by specimens, determined
independently and from literature, by Ch. Ferriére and A. W. Stelfox ; and this was
confirmed by examination of the two syntypes.
Additional material examined: FINLAND, Hangé, Keuru, 2 29 (Hellén, 1937: 125;
1941: 48): HoLLAND, Ermelo, 1 9, 9.ix.1936, J. Lindemans (this specimen has a pair
of pale marks flanking the costa that partly closes the mesolcus) ; Made, 1 9, 2.ix.
1942, Oploo, r 9, 16.ix.1947, H. G. M. Teunissen. Dr. Teunissen came independently
440 A REVIEW AND A REVISION OF
to the conclusion that breviventris Thoms. and fulvipes Kriechb. were conspecific
with rufonotatus Hlgr.; but I must contest his placing of rufonotatus and breviventris
as varieties of Ct. lineola Steph. (1948: 13). The unique male type of Jineola is in the
British Museum (Natural History) and is in very poor condition ; but it is undoubtedly
a different species.
Ct. rufonotatus Hlgr. and dahlbomi Hlgr.
Roman (1914: 16), from a study of the two uniques, placed rufonotatus Higr. in
synonymy with dahlbomi Higr.: I do not understand the reference to Schmiede-
knecht, who merely compiled a description from the two diagnoses given by Holmgren,
whereas Roman had the type before him. Having now been able to study rufonotatus
from adequate material, I must dissent from the synonymy given in 1914. CZ.
dahlbomi differs principally in the following respects.
Head not thus strongly transverse, more strongly punctate: mandibles not as
described for vufonotatus but strongly narrowed to middle and then expanded:
notaulices rather weakly impressed: mesopleura more coarsely and less closely
punctate below: propodeum not short, not strongly raised from base nor falling so.
steeply behind: area superomedia much longer than broad, emitting costulae before
middle and not strongly contracted to base: female hypopygium pointed at apex.
The lower face is bright flavous, apart from a rather narrow, median dark band.
The scutellum is distinctly pale-marked at mid apex. The gaster is dull castaneous
above: tergites 2 to 8 have bright flavous apical margins, which on 2 to 5 are preceded
by large, triangular rufous marks: the petiolar segment has a trace of such pale marking
at mid apex. The species would, therefore, reckon as a red-bodied species according
to the definition given at the head of the key to species (p. 443 below). Stigma and
radial cell are broad, as in rufonotatus.
The male type of gracilis Hlgr. corresponds well with the above, and I think that
dahlbomi Higr. may well prove to be synonymous with it: the lower face, cheeks, and
lower angle of pronotum are not pale-marked. On the other hand, I reject the syno- —
nymy (Roman, 1909: 285) of gracilis Hlgr. with funebris Hlgr., which I would
classify as a black-bodied species: it is a larger and more coarsely sculptured form, —
with stouter scape and petiolar segment, with nervellus intercepted not far below
middle, with scutellum not pale-marked at mid apex, and with tergites of gaster
not rufous marked but merely dull castaneous, with rather obscure pale apical
margins.
Cteniscus gnathoxanthus Gravenhorst
Head moderately to rather strongly narrowed behind eyes in female and strongly
so in male; somewhat strongly emarginate behind: on vertex, temples, and cheeks
rather closely, generally finely but on upper genae in larger specimens moderately
punctate, on epistoma and upper face moderately and more or less closely, and on —
inner orbits very finely punctate: eyes in male distinctly convergent. Clypeus rather
sparsely beset, mostly with moderate punctures; deeply and sharply delimited
basally ; strongly convex in basal half, and more or less strongly impressed before
apex, which is moderately, or sometimes rather broadly rounded. Malar space about.
THE CTENISCINI OF THE OLD WORLD 441
three-quarters distance between mandibular apices in female, about, usually less
than, half such distance in male. Hypostomal costa slightly raised to where met by
genal costa, not close behind mandibular base. Mandible (Fig. 4).
Antennae with scape (Fig. 83) rather elongate, moderately excised externo-ven-
trally ; with postannellus rather elongate, about 1-5 times length of following segment ;
with flagellum 25- to 29-segmented.
Pronotum with dorsal transverse furrow strongly impressed; with epomiae of
moderate strength ; laterally finely or rather finely punctate, with a greater or lesser
amount of rugosity below. Propleura closely and rather finely punctate. Meso-
scutum rather weakly convex, almost flattened on disk; with notaulices sharp and
rather strongly impressed ; rather finely punctate discally, finely to very finely so
peripherally. Scutellum generally rather weakly but sometimes more strongly convex,
flattened and generally to some extent buckled discally ; finely punctate, generally
rather sparsely so above and at sides; not margined beyond scutellar fovea.
Mesopleura closely and rather finely punctate below, finely and rather sparsely so
above: sternaulices very superficial: mesosternum rather finely and closely punctate,
prepectus moderately and closely so: mesolcus moderately impressed, broadened
beyond middle, usually well closed behind. Propodeum shining and obsoletely rugose
above, generally quite clearly punctate on area pleuralis; with costae of moderate
strength but costulae irregular : area superomedia seldom delimited basally, distinctly
longer than broad, emitting costulae a little before middle.
Petiolar segment of female 1-1 to 1-3 times length of its greatest breadth; not.so
broadly based as that of curtisit Hal. but more distinctly explanate, rather roundedly
_ so and generally not strongly margined ; with spiracles situated well before middle
and normally not prominent; dorsally obscurely to moderately rugose, with keels
distinct to well beyond middle, where it is quite strongly raised: petiolar segment of
male longer, 1-3 to 1-8 times length ofits greatest breadth ; with strong, nearly straight-
sided margins and more or less prominent spiracles ; dorsally distinctly rugose, with
keels strong and sharp and extending about three-quarters length of segment.
Remainder of gaster smooth and shining: tergite 2 with thyridia usually narrow in
female, broader and more conspicuous in male. Female hypopygium generally rather
pointed (considerable variation in this): female third valvulae broad, distinctly
finely punctate, normally projecting beyond apex of tergite 8 (Fig. 85).
Wings as described for curtisii Hal., but the areolet may be longer petiolate, and
the nervellus is often intercepted very far below middle: stigma and radial cell not so
elongate as in species with thickened hind tibiae, but more so than in the closest
relatives, radial cell about 2-7 times length of its greatest breadth.
Hind tibia and metatarsus of normal proportions (fig. 80) : claws of all tarsi rather
strongly pectinate.
Length 6 to 9 mm.
Female. Head black, with the following bright stramineous, margined with rufous:
palpi, mandibles except at apex, clypeus, and mark going through malar space usually
up to the hypostomal costa: lower face black, with only slight extensions of malar
marks up the orbits, or with greater or lesser development of similar pale colour ; in
extreme case black only in mid line, rather broadly below antennae to eye margin,
ENTOM, II. 6 31
442 A REVIEW AND A REVISION OF
and almost always just above tentorial pit. Antennae black merging to ferrugineous
above, paler below; with basal segments paler at apex. Thorax and propodeum
black, with humeral angle of pronotum and tegulae stramineous to flavous ; with hind
margin of scutellum only at sides and with mark below hind wing stramineous; with
ventral margin of pronotum distinctly or obscurely pale at sides. Legs mainly red,
the fore and mid tibiae and tarsi rather paler; with fore and mid trochanters and all
trochanterelli flavous, and fore and sometimes mid coxae somewhat flavous-marked ;
with hind tibiae black, rufous in part beneath, distinctly stramineous at base and
rufous at extreme base ; with hind tarsi black, the segments paler at apex and some-
times beneath. Wings with veins and stigma ferrugineous, paler at base: axillary
sclerites flavous. Gaster black above, the tergites having narrow pale apical and pale
lateral margins, broader on the hinder segments: venter.mainly pale, the hypopygium
with or with traces of a broad, V-shaped dark mark well inside margin, and the
other sterna with large, paired dark marks: thyridia castaneous.
Male differs as follows. Lower face may be entirely pale except for a small median
band in upper part; and pale colour may extend up orbits above antennal sockets.
Scape sometimes pale beneath. Legs with a great infusion of flavous into the rufous
parts, the coxae being generally mainly flavous: Irish males, however, have the hind
coxae mainly black and the hind femora dark. Pale apical margins of tergites 2 and
3 may be broader: hypopygium and previous sternite mainly black.
Redescribed from the following material: IRELAND, Co. Kildare, Landenstown,
I 9, 20.vili.1941, Skerries Bog, 3 29, 26.ix.1946; Co. Wicklow, Rathdrum, 1 9, 6.viii.
1927, Athdown, I 4, 20.viil.1944, A. W. Stelfox ; Co. Dublin, Howth Castle, in wood,
I g, 10.viii.1939, F. J. O’Rourke: ENGLAND, unlocalized, 1 9, T. Desvignes coll. ;
Norfolk, Catfield, 1 9, 16.vii.tg20, J. J. F. X. King: Betcium, unlocalized, 1 9
C. Wesmael; Vivier d’Oie, 1 9, 13.vii.1884, I 9, 23.vii.1880, I 9, 28.ix.1880;
St. Croix, 1 9, 4.x.1876; Schaffen, 2 99, 22.ix.1885; Schelle, I 9, 11.viii.1885;
Carlsbourg, I 9, 12.vi.1880 ; Groenendaele, I 9, 15.ix.1891; ? locality, I 9, 14.ix.1885,
J. Tosquinet : SWEDEN, Skane, Fjallfota sj6, on Salix, 2 29, 7.viii.1938, D.M.S. & J.F.
Perkins ; Halsingland, Bergvik, bred from under bark, 3 $g,5 29, em. I16-19.vi.1947,
O. Lundblad: FINLAND, Sortavala, Rauskunvuori, I 9, 2I.vili.1935 ; Ladoga, Valamo, —
Pienennukkananlahti, I 9, 31.vill.1935, G. J. K. (see Kerrich, 1940a): FRANCE, un- —
localized, ? France, 1 9, coll. Puton (Mus. Paris): GERMANY, unlocalized, ? Germany,
2 22, O. Schmiedeknecht; probably Berlin district, 5 99, Ruthe coll.; Celle, 1 2, —
29.viii.1942; Gottingen, 1 9, I1.ix.1946, R. Hinz; Bavaria, Gérkau, I 9, vii.1930, —
Th. Kupka: CZECHOSLOVAKIA, Bohemia, Chodau, I 9, 5.ix.1885, I 9, 19.vll.1925,
R. von Stein; Erzgebirge, Aussig, I 9, .27.viii.1933; Teschen, Rosenberg, I Q,
7.vii.I936; Bohumin, Haatsch, 1 9, 23.viii, 1 2, 28.viii, I 9, 20.ix.1936, Th. Kupka.
A black-bodied species without thickened hind legs, and with scutellum not pale-
marked except at sides of hind margin. Characteristic features are the rather elon-
gate scape and post-annellus, the closely and rather finely punctate lower part of the
mesopleura, the broad or rather broad third valvulae, and the distinctly convergent
male eyes. The male is very scarce in collections: Professor Lundblad’s bred series,
part of which he has presented for the British Museum collection, enabled me to
associate the sexes with confidence.
THE CTENISCINI OF THE OLD WORLD 443
Cteniscus gibbulus Holmgren
Characters diagnosed from a pair taken 7m coitu on a leaf of Alnus at Hédr, Skane,
Sweden, 25.vi.1938, J. F. Perkins: these were thus determined by him, and also
fitted my recollection of the type material.
To those characters given in the key to species I add the following: head less strongly
narrowed than in gnathoxanthus Grav., sex for sex: mesolcus very strongly impressed :
propodeum with keels stronger: petiolar segment with dorsal keels strong, in female
almost straight-sided: hind legs rather stouter than in gnathoxanthus Grav.: wings
with stigma and radial cell not nearly as broad as in rufonotatus Hlgr. yet broader
than in gnathoxanthus Grav., radial cell 2:5 times as long as broad: hind coxae and
femora mainly black in both sexes.
KEY TO PALAEARCTIC BLACK-BODIED SPECIES OF
CTENISCUS HALIDAY, THOMSON
Black-bodied species, i.e. species with tergites of gaster black or dull castaneous ;
very often with more or less narrow pale hind margins, which sometimes are trian-
gularly produced forward ; in one species often with a rather extensive rufous mark
on tergite 2; but ot with two or more tergites mainly red.
1. Hind tibiae distinctly thickened, as in hoffert Greg. (Fig. 81) or more so; in case
of doubt scutellum distinctly pale-marked at mid apex ; d | 2
Hind tibiae and tarsi of normal proportions; in case of doubt scutellum not pale-
marked at mid apex : , : : : : : ; , 5
2. Propodeum normally pilose, shining, and not closely punctate: petiolar segment
not strongly raised in middle: hind legs thickened about as in Fig. 81, in inner
side view the hind metatarsus about 7} times length ofits greatest breadth 3
Propodeum strongly griseo-pilose, dull and closely punctate: petiolar segment
strongly raised in middle: hind legs more thickened, in inner side view the
hind metatarsus about 54 times length of its greatest breadth ; Pee:
3. Mesosternum irregularly, rather sparsely and coarsely, almost rugose-punctate:
sternaulices very superficial: female third valvulae normally not reaching
apex of tergite 8: radial cell usually more than 2-6 times length of its greatest
breadth: lower face pale, with only a median dark mark above: female
hypopygium pale. ‘ . hoffert Greg.
Mesosternum regularly, rather finely to moderately and closely punctate:
sternaulices moderately impressed: female third valvulae extending a little
beyond apex of tergite 8, rather small but stout and densely hairy: radial cell
2:6 times length of its greatest breadth or usually less: lower face with a com-
plete median black band: female TERE, mainly black, with narrow
pale margin . é ‘ ‘ . Lineiger Thoms.
4. Head from above distinctly eee dian eye tage behind eye: upper face moderately
punctate: mesopleura rather coarsely and closely punctate below: area
superomedia distinctly delimited basally: third valvulae projecting distinctly
444 A REVIEW AND A REVISION OF
beyond apex of tergite 8: scutellum with panirigas quadrangular yellow
mark at mid apex . . curtisit Hal.
Head from above about equal to eye length behind eye: upper face rather finely
and sparsely punctate: mesopleura rather finely and sparsely punctate below:
80
8!
Fics. 80-81. Left hind leg, in sinistro-lateral view, of Fig. 80. Cteniscus gnathoxanthus Grav.
Fig. 81. Cteniscus hofferi Greg.
area superomedia not delimited basally: third valvulae broader, projecting
scarcely beyond apex of ise 8: scutellum with no more than a dull red
mark at mid apex . ‘ : ; ; . romani sp. n.
5. Petiolar segment with distinct ‘cel lobes : R , ; . —.
Petiolar segment without distinct basal lobes ; : a.
6. Head about as long behind eyes as eye length, seen nd pian hypostomal
THE CTENISCINI OF THE OLD WORLD 445
costa strongly raised: area superomedia about as broad as long: female gaster
quite stoutly spindle-shaped: scutellum apr emai pale-marked at mid
apex: female hypopygium pale . ; j . elegans Steph.
Head much longer behind eyes than eye length, seen from above: hypostomal
costa distinctly but not strongly raised: area superomedia distinctly longer
than broad: gaster much more elongate: scutellum not pale-marked at,mid
apex: female hypopygium blackish . : . similis Hlgr.
7. Gaster elongate and slender, with tergites 4 and 5 as ie as their breadth at
base . ‘ , pectoralis Hellén
Gaster of more normal proportions, with tergites 4 and 5 strongly transverse 8
8c
NN
83
Pesce ececnce SER
84
Fics, 82-84. Left antennal scape and following segments of Fig. 82. Cteniscus unicinctus Higr.
Fig. 83. Cteniscus gnathoxanthus Grav. Fig. 84. Cteniscus romani sp. n.
8. Head much longer behind eyes than eye length, seen from above; at least in well-
developed specimens distinctly broadened: petiolar cau iain more
elongate than in alternate i 9
Head narrowed and not much longer behind eyes ‘than eye length, seen ‘from
above: petiolar segment not elongate, not more than about 14 times length
of its greatest breadth in female or 13 times (normally) in male. . 50
g. Antennae with scape strongly convex dorsally and with flagellum decidedly
slender (Fig. 82): mandibles with lower tooth more prominent than upper,
though a little shorter: female hypopygium sharply folded medially in apical
half, pale: male malar space short: lower face without a median dark band:
thorax mainly pale below in female . ; ‘ ; unicinctus Hlgr.
446 A REVIEW AND A REVISION OF
Antennae with scape of normal proportions and with antennae not relatively
slender: mandibular teeth in same plane: female hypopygium not sharply
folded, dark with pale margin: malar space not short in either sex: lower
face with a median dark band: thorax scarcely at all pale below
‘ : . ‘ ; : : : ‘ ‘ consobrinus Hlgr.
10. Head strongly transverse: upper face finely, seldom in part closely punctate:
lower mandibular tooth distinctly a little longer and more prominent than
upper: female third valvulae narrow (Fig. 86): stigma and radial cell relatively
broad, the latter about 2-1 times as long as broad: hind tibiae mainly clear
red, darkened only at apex and not stramineous at base: female propleura
largely pale rufonotatus Hlgr.
(= = breviventns Thoms., fulvipes Kriechb.)
85 3 86
Fics. 85-86. Apex of gaster, in sinistro-lateral view, of Fig. 85. Cteniscus gnathoxanthus Grav.
Fig. 86. Cteniscus rufonotatus Hlgr.
Head not strongly transverse: upper face moderately and more or less closely
punctate: lower mandibular tooth not distinctly longer or more prominent than
upper: female third valvulae broad (e.g. Fig. 85): stigma and radial cell not
nearly so broad: hind tibiae mainly black, distinctly dull stramineous at
base: propleura scarcely pale-marked : ; ‘ ; ‘ . ta
11. Antennae longer and more finely hairy, with postannellus about 1-5 times length
of following segment: mesoscutum longer, rather weakly convex: mesopleura —
closely and rather finely punctate below: propodeum longer, with area supero-
media longer than broad: female hypopygium mainly pale, with or with ©
traces of a broad See dark mark: eyes in male distinctly convergent —
below .. ; : . gnathoxanthus Grav. —
Antennae shorter and 1 more coarsely hairy, with postannellus about 1-3 times —
length of following segment: mesoscutum shorter, rather strongly convex:
mesopleura rather coarsely and not so closely punctate below: propodeum
shorter, with area superomedia broader than long: female hypopygium mainly —
dark, with pale apical margin: eyes almost parallel . gibbulus Hgr.
Cteniscus aurifluus Haliday
1832 Tryphon (subgenus Cteniscus) aurifiuus Haliday in Curtis, British Entomology, 9: 399.
1838 (Oct.) Tryphon (subgenus Cteniscus) aurifiuus Haliday, Ann. Mag. nat. Hist. (1) 2: 113.
1839 Exenterus geniculosus Schiodte, Guerin’s Mag. Zool. 9: 11-12 (synonym by Morley 1913).
THE CTENISCINI OF THE OLD WORLD 447
1858 Exenterus approximatus Holmgren, Svensk Vet.-Akad. Handl. 1: 239 (synonym by Roman
1914).
1858 Exenterus geniculosus Holmgren, ibid. 241.
1912 Cteniscus geniculosus Schmiedeknecht, Opuscula Ichneumonologica, 5: 2343-2344.
1913 Exenterus aurifiuus Morley, Entomologist, 46: 261.
1914 Cteniscus geniculosus Roman, Ark. Zool. 9 (2): 16.
Head little broader than thorax ; very slightly broadened behind eyes ; broadly and
rather shallowly emarginate behind ; much longer behind eyes than eye length, seen
from above: shining, with vertical region and temples sparsely and very finely
punctate; with cheeks, upper face and sides of lower face about twice as densely
and more coarsely so, and with epistoma still more densely and coarsely so. Frons
with a deep impression above each antennal socket, and with distinct interantennal
keel. Clypeus rather coarsely and, in apical half, sparsely punctate ; broadly rounded
apically ; deeply delimited ; strongly convex ; slightly impressed before apical margin.
Malar space about as long as distance between apices of mandibular teeth in female,
distinctly shorter in male. Genal costa not continuous, weakly raised apically:
hypostomal costa slightly more strongly raised. Upper mandibular tooth a little
larger than lower (Fig. 5).
Antennae with scape rather deeply excised externo-ventrally: flagellum 29- to
35-segmented (often with a segment more in one antenna than the other, 34 : 34 in
holotype), tapering gradually to apex.
Pronotum with dorsal transverse impression broad and shining ; with epomiae of
moderate strength; laterally rather finely to moderately punctate above, to some
extent longitudinally rugose below. Propleura shining, finely punctate, rugose near
inner margin. Mesoscutum dorsally rounded to level of tegulae, thereafter nearly
flat to before scutellar fovea; finely punctate, discally somewhat sparsely, peripher-
ally about twice as densely so; between notaulices very weakly rugose: notaulices
deeply impressed anteriorly, thereafter very superficial. Scutellum strongly convex
and rather sparsely punctate ; not margined to middle; its sides irregularly finely to
coarsely rugose. Mesopleura and mesosternum moderately punctate, mesopleura
sparsely so above; prepectus more coarsely so: epicnemial carina slightly elevated:
sternaulices rather strongly impressed: mesolcus broadened posteriorly, completely
or partially closed behind. Propodeum finely punctate and shining, more or less
rugose behind ; with costae strong: basal area distinctly delimited: area superomedia
elongate-hexagonal, emitting costulae rather before middle.
Gaster not clavate; with tergites I, except near mid apex, and 2 and 3 at base
very finely leathery in female, rugose in male ; elsewhere smooth and shining. Petiolar
segment about 1-3 times length of its greatest breadth in female, about 1-5 times in
male: very strongly margined ; with distinct basal lobes ; with spiracles prominent in
male ; medially strongly raised ; with strong keels running from basal excavation for
two-thirds to three-quarters the length of the segment and bounding a shallow furrow,
deeper in male. Tergite 2 with thyridia rather deep but narrow and inconspicuous ;
with diagonal basal impressions ill-defined but extensive, in male rather deep and
also indicated on tergite 3. Female hypopygium rather narrowly to broadly rounded
apically: female third valvulae of moderate breadth.
448 A REVIEW AND A REVISION OF
Fore wings with basal and disco-cubital veins not nearly parallel: with areolet
broad and shortly petiolate: with nervus parallelus emitted very low on brachial
cell. Hind wings with nervellus antefurcal and intercepted far below middle.
Legs with tarsi relatively short. Tarsal claws not pectinate, or with no more than
a trace of pectination.
Length 7 to Io mm.
Female. Head black, often inclining to blue-black ; with large genal and very large
lateral facial marks white, these sometimes continuous through malar space: clypeus
in apical half, sometimes mainly, white, merging to dull straw-coloured or darker
marginally: mandibles in basal half white, merging through dull reddish to black at
extreme apices: palpi pale reddish. Antennae basally black, with flagellum ferru-
gineous (relatively pale in type, which may be faded). Thorax and propodeum black,
sometimes with scutellum medially and metascutellum dull red: tegulae straw-
coloured, often with an outer darker mark: legs red, with apices of hind femora and
tibiae and of all tarsi black. Wings scarcely infumate ; with wing-veins and stigma
fuscous, basally pale. Gaster with segments I to 3 and sometimes base, or the whole,
of 4 and even sternite 5 red, tergite 1 being black usually only at extreme base;
remainder black: tergites 6 and 7 broadly, and sometimes 3 to 5 narrowly white-
margined at apex: ovipositor red.
Male. Differs in absence or great reduction of pale genal marks: clypeus duller:
segment 1 of gaster black in about basal half, or sometimes almost to apex: apical
segments with narrower pale margins.
This species is illustrated on Pl. 7.
Redescribed from the following material: IRELAND, Co. Sligo, Carrowgarry, I 9,
7.X.1940 ; Co. Kildare, Kilkea, 1 3, 10.ix.1940 ; Landenstown (marsh), 2 99, 4 $3, 29.ix.
1946, A. W. Stelfox ; Haliday collection, 4 29, 4 $d, including holotype and allotype
(see below), and including 2 29, 2 gg with green-marked pinheads indicating de-
finite Irish origin: ENGLAND: Cambs., 1 9, L. Jenyns; Wicken Fen, I 9, 3.vi.1934,
I 4g, 18.viii.1931, G. J. K.; Chesterton, r 9, 16.vii.1934, F. J. Simmonds; Yorks.,
V.C. 64, Askham Bog, I 3, 15.vii.1944, W. D. Hincks.
Additional material seen: IRELAND, 27 specimens from ten different counties,
A. W. Stelfox: BELcium, Tronchiennes, I 9, 10.viii.1867, La Hulpe, 1 ¢, 3.viil.
1879, Genval, I 9, 9.vii.1g01, J. Tosquinet: SWEDEN, series in Holmgren collection,
including unique type of approximatus Hlgr. (Stockholm district, 23.viii, C. Boheman):
GERMANY, Schwerin, I 4, viii.1889, S. Brauns: Russia, Samara, I g, vii.1929,
Lubischew (coll. A. Seyrig).
Time of appearance: the dates of capture of specimens seen range continuously
from 6 July to 12 October. I have wondered whether my own early June date could
have been a Japsus for July, but Morley (1911) has also recorded the species in early
June.
I agree with Roman’s placing of approximatus Higr. in synonymy (1914). The
unique male type appears to be a freak form, in which the petiolar segment has the
keels and the spiracular tubercles very powerfully developed, and in which the
scutellum is very much buckled. The sculpture is mostly developed rather more
strongly than usual; and the general form is unusually squat, in conjunction with
THE CTENISCINI OF THE OLD WORLD 449
which the basal and disco-cubital veins are nearly parallel. But in other respects the
specimen agrees so closely with the normal form in both structure and colour pattern
that I am convinced it is not specifically distinct.
Selection of types. In the Haliday collection are ten specimens which appear to have
been attributed to aurzfluus: these may be numbered 1 to 10. No. I, a male, bears the
label ‘Named by Claude Morley. Exenterus aurifluus, Hal. $9. vi.1913’: this I take
to be a general determination label covering the ten specimens. No. 7, a female,
bears three labels ‘Type C. M.’, ‘Haliday’, and ‘aurifluus’, the last-mentioned in
Haliday’s own writing. No. 8, a male, bears the single label ‘Haliday’. No. 9, a
female, bears the two labels ‘Type C. M.’ and ‘Named by Claude Morley. Cteniscus
aurifluus, Hal TYPE vi.1913’. Now Nos. 1 to 8 inclusive belong to the species that
has been generally understood as aurifluus Hal. (= geniculosus Schigdte), and agree
with the description given above ; but Nos. 9 and 1o-are female and male respectively
of a different species, which runs down in Schmiedeknecht’s key to pictus Grav. and
fits his description of that form, apart from having an indefinite dark mid line on the
yellow lower face.
Why did C. Morley select two types of aurifluus and label them as he did? It
seemed probable that it was on account of Haliday having quoted both sexes in his
diagnosis, and that Morley spared himself the effort of duplicating Haliday’s specific
label. A. W. Stelfox, who had charge of the collection, wrote to ask whether this
were so, pointed out that the second in order had dark hind femora, and asked which
of the two should be the first lectotype. In the course of his reply, Mr. Morley
wrote that he presumably put a type label on the second aurifluus because he had
no cotype labels ; and that he evidently did not consider the pedal coloration specific.
From this it is to be inferred that the first in order, No. 7, is to be taken as Morley’s
first lectotype.
Distinctions, based on specimens 9 and 10, of Cé. pictus Grav. Schmied. from the
first species are given below. I think there can be no doubt, on both positive and
negative evidence, that Haliday’s diagnosis refers to specimens 1-8 and not to 9-Io.
Accordingly, I designate specimen No. 7, bearing Haliday’s own manuscript label,
as holotype: thus, the accepted interpretation of Haliday’s species is confirmed.
Specimen No. 1, a male of definite Irish origin, I designate as allotype.
Cteniscus pictus Gravenhorst, Schmiedeknecht
In the pair misplaced with the series of aurifluus in the Haliday collection, I find
the following distinctions from that species. Head distinctly broader than thorax,
much more transverse. Clypeus narrowly rounded apically, almost truncate in
middle of margin. Genae more buccate. Pronotum below more strongly and ex-
tensively rugose. Mesopleura distinctly alutaceous: mesolcus narrower and more
determinate. Propodeum with costae weaker: area superomedia not basally de-
limited. Tergite 1 more strongly and regularly narrowed to base. Tergite 3 smooth
throughout and tergite 2 almost so. Thyridia conspicuous but not deep. Fore wings
with stigma rather broad: with basal and disco-cubital veins nearly parallel. Tarsal
claws rather weakly but distinctly pectinate.
ENTOM. II. 6 3K
450 A REVIEW AND A REVISION OF
Head in both sexes with lower face, except for an indefinite median line, and with
clypeus, except around apical and sutural margins, yellow: genae in both sexes with
large yellow marks confluent with the yellow of the lower face. Thorax with the —
following conspicuously flavous: humeral angle of pronotum, hind margin of meso-
scutellum at sides, and line below hind wing: also front margin of pronotum above and
at sides in male. Legs flavous except as follows: fore and mid coxae at extreme
base, and hind coxae and femora mainly, ferrugineous to black: fore and mid coxae
mainly, hind coxae apically in male, and all trochanters and trochanterelli, stra-
mineous to flavous. Gaster of female with whole venter red: dorsum mainly red,
tergite 1 black except at apex, tergites 6 to 8 mainly and 4 to 5 at sides dark; ter-
gites 5 to 8 apically white-margined: gaster of male more extensively dark.
1. gnathoxanthus Grav. See above, p. 440.
ditto var.2 ? = elegans Steph., p. 430.
2. fulvipes Kriechb. = rufonotatus Hlgr., p. 437.
3. gtbbulus Hlgr. see p. 443.
4. breviventris Thoms. = rufonotatus Hlgr., p. 437.
5. consobrinus Hlgr. see p. 436.
6. wunicinctus Hgr. see p. 434.
7. similis Hlgr. see p. 445.
8. pullus Hgr.
9g. morio Hlgr. = Anisoctenion rubiginosus Grav., p. 328.
to. funebris Hlgr. see p. 440.
Ir. gracilis Hlgr. see p. 440.
12. pachysoma Steph.
13. pratorum Woldst. = Exyston pratorum, p. 385.
14. pictus Grav. see p. 449.
15. marginatus Thoms.
16. borealis Hlgr.
17. alpicola Higr. placed in synonymy with borealis Higr. by Roman
(1913).
18. deletus Thoms.
19. ustulatus Hgr. placed in synonymy with flavomaculatus Grav.
by Roman (1931). ;
20. wmbellatarum Woldst. placed in synonymy with flavomaculatus Grav. |
by Roman (1931).
21. rufilabris Hlgr.
22. flavilabris Higr. placed in synonymy with flavomaculatus Grav.
by Roman (1931).
23. connatus Higr. placed in synonymy with flavomaculatus Grav.
SUMMARY OF THE SPECIES TREATED UNDER CTENISCUS
BY SCHMIEDEKNECHT (1912)
by Roman (1931).
THE CTENISCINI OF THE OLD WORLD 451
punctipes Thoms.
pumilio Higr.
pumilus Hlgr.
lineiger Thoms.
mitigosus Grav.
approximatus Hlgr.
frigidus Hlgr.
hostilis Hgr.
= Anisoctenion pumilio, p. 331.
see p. 420.
see p. 424.
= aurifiuus Hal., p. 446.
placed in synonymy with flavomaculatus Grav.
by Roman (1931).
32. signifer Thoms.
33. geniculosus Schigdte = aurifluus Hal., p. 446.
34. sedulus Woldst. see p. 315.
35. punctipleuris Thoms. considered to be var. of dorsator Thunb. by A.
Roman.
36. limbatus Hlgr. = dorsator Thunb., see Roman (1912).
37. brevigena Thoms.
38. flavomaculatus Grav. see p. 420.
minutulus Pfank. placed in synonymy with flavomaculatus Grav.
by Roman (1931).
40. limbatellus Hlgr. see p. 420.
41. genalis Thoms.
42. dahlbomi Hlgr. see p. 440.
43. macrocephalus Hlgr. = unicinctus Hgr., p. 434.
44. autumnalts Brischke
45. vufonotatus Hlgr. see p. 437.
46. quadrinotatus Thoms. placed in synonymy with flavomaculatus Grav.
by Roman (1931).
47. albicoxa Thoms.
48. T-nigrum Thoms.
CONCLUSION
The author is very conscious that much remains to be done ; but time could not be
spared in the near future for carrying this through, and others await the results so
far attained. The genera Acrotomus s.str. and Diaborus auctt. require intensive
revision, and in the latter genus the types should be examined. The red-bodied species
of Cteniscus merit equal treatment with the black-bodied ones; and it remains to be
seen for how many species there are both red and black-bodied forms. The work of
Peck (1937) should be followed up and extended, with particular reference to the
male genitalia of Exenterus. W. R. M. Mason, who is making a study of the New
World species, has discovered further fresh characters widely applicable within
the tribe ; and when these have been fully worked out and published, they should be
studied on Old World material. Larvae have been described only in the genus
Exenterus, so ample scope remains in this tribe for extending the work of Beirne
(1941).
ENTOM. II. 6. 3K2
SUPPLEMENT
Genus ACROTOMUS Holmgren
Compare pp. 319-321 above. I re-examined the material in the C. G. Thomson
collection in 1952, and confirmed that Thomson had succinctus Grav. under the name
lucidulus, apart from one specimen to which reference has been made. Under auricu-
latus he had a good series of lucidulus Grav. agg. ; and these are not of any obviously
distinct form, despite his record of their occurrence mostly in conifer woods.
Genus ANISOCTENION Forster
Anisoctenion ridibundus Gravenhorst
Compare pp. 336-338 above. Two male specimens, contained in a sending re-
ceived in 1952 from Professor Sachtleben, bear the following data: GERMANY,
Mecklenburg, 6.vi.1894, F. W. Konow. One of these has the broad, percurrent,
median black band of the lower face, which has been considered diagnostic for the
male of this species; but the other is without it, as in the female.
Anisoctenion genalis Thomson
1883 Cteniscus genalis Thomson, Opuscula Entomologica, fasc. 9: 894.
1888 Cteniscus (or Delotomus) genalis Thomson, Opuscula Entomologica, fasc. 12: 1255-1256.
Head, seen from above, about 1} times eye length behind eye; slightly narrowed
and rounded behind eyes; shallowly emarginate behind: with punctures of mixed
sizes, fine to moderate, rather dense on vertical region but much sparser on temples
and genae, especially in female: upper face and epistoma densely and rather coarsely
punctate, sides of lower face more sparsely and moderately so. Clypeus sparsely
beset with coarse and very fine punctures; basally rather deeply but not sharply
delimited ; moderately or rather strongly convex ; broadly rounded at apex in female,
moderately so in male. Malar space about equal to distance between mandibular
apices. Hypostomal costa rather weakly raised (about as in ridibundus Grav.): genal
costa weakly but sharply raised, inflected.
Antennae with scape decidedly short and stout, weakly excised externo-ventrally:
flagellum 27— to 32-segmented, distinctly expanded in middle in female, with segments
distinctly separated in male.
Pronotum with dorsal transverse impression rather weak, transrugose; with
epomiae sharp and of moderate strength; laterally closely, moderately to rather
coarsely punctate above, coriaceo-rugose below. Propleura rather finely and closely
punctate, shining. Mesoscutum rather weakly to moderately convex ; with notaulices
weak, but sometimes sharply impressed ; for the most part moderately and rather
closely punctate, or in larger specimens rugulose-punctate, more finely and densely
Ne
en - er
THE CTENISCINI OF THE OLD WORLD 453
punctate anteriorly between notaulices. Scutellum moderately convex in female,
strongly so in male, slightly flattened in larger specimens ; margined to beyond scutel-
lar fovea ; finely to moderately punctate above and behind ; obscurely rugose-punctate
at sides in female, more strongly so in male. Mesopleura rather coarsely punctate
below, moderately so above: sternaulices moderately impressed: mesosternum rather
finely to finely punctate, prepectus moderately rugose-punctate: epicnemial carina
slightly elevated: mesolcus not strongly yet very distinctly impressed, broadened
and transcostate posteriorly, more or less closed behind. Propodeum finely rugose,
with area externa finely to rather finely, and area pleuralis for the most part rather
finely to moderately punctate, with costae including costulae normally complete, weak
in female but of moderate strength in male: area superomedia not delimited basally,
about twice as long as broad, broadest where it emits the costulae, before or behind
middle.
Gaster slender spindle-shaped or weakly clavate in female, more parallel-sided
behind petiolar segment in male. Petiolar segment 1-4 to 1-5 times length of its
greatest breadth in female, 1-4 to 1-6 times in male; laterally sharply margined,
strongly so in male; with basal lobes more or less strong and spiracles more or less
prominent ; with dorsal keels distinct to about middle in female, not much farther in
male, and with a shallow median depression behind them ; finely to moderately punc-
tate, smoother between keels in female, and smooth at mid apex. Tergite 2 with
thyridia broad but inconspicuous. Punctation on tergite 2 rather fine to moderate,
on remainder of gaster gradually finer to apex. Female hypopygium quite sharply
pointed at apex: third valvulae of moderate size and breadth, with ventral margin
almost straight, extending slightly beyond apex of tergite 8.
Fore wings with basal and disco-cubital veins weakly or rather weakly divergent ;
with areolet usually distinctly petiolate ; with nervus parallelus emitted well below
middle of brachial cell. Hind wings with nervellus slightly antefurcal, intercepted
distinctly below middle.
Legs of normal build. Claws of all tarsi moderately pectinate, the pectination
distinctly not reaching apex.
Length 6 to 84 mm.
Head black, with palpi dull rufous, and with the following parts stramineous to
pale flavous, margined with dull fulvous: mandibles except at apex, and clypeus
except at base. The dark basal margin of clypeus may be less clearly defined than in
alacer Grav., and may be darkened only to castaneous. Antennae with flagellum dull
rufous in female, more ferrugineous in male; with scape and pedicellus usually con-
colorous below, but distinctly darker above and at sides ; with several basal segments
distinctly paler at apex. Thorax and propodeum black, with the following parts in
varying degree paler: humeral angles narrowly, marks beside base of scutellar fovea,
hind margins of scutellum, and small marks below fore wings: tegulae stramineous.
Legs in female with coxae, trochanters, and trochanterelli varying from pale dull
rufous, on fore coxae beneath, to more or less infuscate, the trochanters and tro-
chanterelli dull flavous at apex: hind femora rufous, very slightly infuscate at apex:
fore and mid femora pale rufous, sometimes flavescent in part: tibiae and tarsi rufo-
testaceous, with very slight infuscation. Male differs as follows: coxae, trochanters,
454 A REVIEW AND A REVISION OF
and trochanterelli much more infuscate, the hind coxae blackish: hind femora
varying to ferrugineous: hind tibia and mid and hind tarsi with infuscation stronger.
Wing veins and stigma dull testaceous to ferrugineous, paler at base. Gaster with the
following parts rufous: tergites 2-4 and female hypopygium and third valvulae,
usually also tergite 5, and sternite of petiolar segment and its tergite in about apical
third ; with tergite 1 in about basal two-thirds or wholly, tergites 5 or 6-8, and last
two sternites of male strongly infuscate ; often with rather weak, pale hind margins
to these sclerites: sternal region otherwise stramineous to pale rufous.
Redescribed from the following material: ENGLAND, Devon, Dartmoor, 1 4,
8.vi.1935, R. C. L. Perkins: Norway, Buskerud West, Al, 1 9, 8.vii.1898, E. Strand
(Deuts. Ent. Inst.): SWEDEN, Skane, Alnarp, I 2 (type), Palsj6, 2 99; Norrland
(unlocalized), 1 g, C. G. Thomson. The Austrian specimen (see p. 350 above) I
determined as Antsoctenion sp. near alacer Grav. (G. Heinrich 1951, Bonn. Zool:
Beitr. 2: 246).
Genus EUDIABORUS mihi
Following the statement (pp. 415-417 above) on the state of our knowledge of
Eudiaborus (= Diaborus auctt.), I made a preliminary study of this genus in the
early summer of 1952, followed by visits to Lund, Stockholm, and Helsingfors. I
believe that the results attained should not be withheld from publication until it is
possible to complete a full-scale revision ; therefore I intend to communicate them to
the Editor of Opuscula Entomologica, Lund, and hope that he may be able to publish
them in 1953.
Eudiaborus boreoalpinus mihi
This species was described (pp. 417-420 above) on account of the distinctive struc-
ture, at least in the female, and on account of the boreo-alpine distribution. Further
material has been seen in 1952, and I have been asked to publish the additional
records.
Further material seen: GERMANY, Silesia, Glatzer Schneeberg, I 3, 2.viii, I d,
II.vili, 2 99, 8.vili.1g21, K. Hedwig (Deuts. Ent. Inst.): Norway, Buskerud East,
Krgdsherad, I 9, E. Strand (Deuts. Ent. Inst.): SWEDEN, Lapland, Karesuando, r 9,
J. W. Zetterstedt (= lituratorius var. b, Zetterstedt 1840, p. 389) ; Tornetrask, 2 99,
28.vii, I Q 9.viii, R. Malaise: FINLAND, Pelkosenniemi, 1 92, Clayhills; Kuusamo,
I 9, 19.vil. 1934 (ref. 360), R. Krogerus; Kuolajarvi, 2 99 27.vii.1934 (ref. 145/6),
R. Krogerus. .
In smaller female specimens the head may be, as in the males, rather more narrowed
and rounded behind the eyes than in the type specimen figured above (Fig. 77).
The female propleura may have the outer margin quite broadly black in anterior
half. In one Silesian male, the median black band of the lower face is so broadened
as to leave only a pair of triangular pale marks going about two-thirds the way
up orbits of lower face from the tentorial pits.
In this species the mesosternum of the female is very little concave, whereas in
pallitarsis Thoms. it is strongly concave.
ee
THE CTENISCINI OF THE OLD WORLD 455
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sawfly (Diprion polytomum, Htg.) in Europe. Ibid. 28: 341-393.
Mor teEy, C. 1903. The Ichneumons of Great Britain, 1, Plymouth.
1911. The Ichneumons of Great Britain, 4, London.
—— 1913. On the Ichneumonidae of the Dublin Museum. Entomologist, 46: 259-263.
1936. The Hymenoptera of Suffolk. Portiosecunda. Trans. Suffolk Nat. Soc.3 (2) : 132-162.
PEck, O. 1937. The male genitalia in the Hymenoptera (Insecta), especially in the Ichneu-
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PrankucH, K. 1906-1907. Die Typen der Gravenhorstschen Gattungen Mesoleptus und
Iryphon. Z. Hym. Dipt. 6: 17-32, 81-96, 217-224, 289-296; 7: 17-24, 145-155.
1919. Der aussere Kérperbau der echten Schlupfwespen (Ichneumonidae). Abh. naturw.
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1925. Ein Beitrag zur Ichneumoniden-Fauna Nordschleswigs. Z. wiss. Insekt. Biol. 20:
9-13, 57-61, 150-156, 175-182.
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1912. Die Ichneumonidentypen C. P. Thunbergs. Zool. Bidr. Uppsala, 1: 229-293.
—— 1913. Neubeschreibungen und Synonyme zur nérdlichen Ichneumonidenfauna Schwedens.
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1914. Beitrage zur schwedischen Ichneumonidenfauna. Ark. Zool. 9 (2): I-40.
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THE CTENISCINI OF THE OLD WORLD 457
Roman, A., 1931a. Entomologische Ergebnisse der schwedischen Kamtchatka-Expedition 1920-
1922. Ark. Zool. 28A(6): 1-32.
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TOSQUINET, J. 1890. See Jacobs & Tosquinet.
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VIERECK, H. L. 1912. Tryphoninae—a review. Proc. ent. Soc. Wash. 14: 175-178.
1914. Type species of the genera of Ichneumon Flies. Bull. U.S. nat. Mus. 88: 1-186.
WALLEY, G. S. 1937. The genus Smicroplectrus in Canada (Hymen.: Ichneumonidae). Canad.
Ent. 69: 131-136.
WEstwoop, J. O. 1838-40. Amn Introduction to the Modern Classification of Insects; founded on
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Wotpstept, F. W. 1877. Beitrag zur Kenntniss der um St. Petersburg vorkommenden Ich-
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ZETTERSTEDT, J. W. 1840. Insecta Lapponica. Lipsiae.
458 A REVIEW AND A REVISION OF
APPENDIX I
Table of the Occurrence of Species in certain European Countries
+ signifies specimen(s) seen by author. © signifies record requiring confirmation.
xX signifies record accepted by author. — signifies record rejected.
Ireland
Acrotomus lucidulus Grav. agg.
Acrotomus succinctus Grav.
Anisoct. triangulatorius Grav.
Anisoct. rubiginosus Grav.
Anisoct. pumilio Hgr.
Anisoct. ridibundus Grav.
Anisoct. laticeps Grav.
Anisoct. laetus Grav. : ‘ : : 1+
Anisoct. alacey Grav.
Exenterus apiarius Grav.
Exent. abruptorius Thunb.
Exent. simplex Thoms.
Exent. oriolus Hart.
Exent. ictericus Grav.
Exent. amictorius Panz.
Exent. claripennis Thoms. : : ; :
Exent. adspersus Hart. ; : : - 2 +/+
Exent. tricolor Roman
Exent. vellicatus Cushm.
Exent. confusus sp.n. :
Exyston cinctulus Grav. : " ‘ ~ | +
Exy. genalis Thoms. : : . : :
Exy. phaeorrhaeus Hal. ; ‘ ‘ ‘ ~ {+
Exy. subnitidus Grav.
Exy. albicinctus Grav.
Exy. pratorum Woldst. +
Parexy. calcavatus Thoms. . : : : t+
Smicro. quinquecinctus Grav. —
Smicro. excisus sp.n. :
Smicro. trianguligena sp.n. . . . : 2 Ae
Smicro. bohemani Hlgr.
Smicro. perkinsorum sp.n.
Smicro. heinrichi sp.n. : s A A t+
Smicro. jucundus Hlgr. ‘ .
Eudiab. boreoalpinus sp.n.
Cteniscus hofferi Greg. : es > t+
Ct. curtisii Hal. 7 : ‘ ‘ : ~| +
Ct. romani sp.n. +
Ct. elegans Steph.
Ct. similis Higr. °
Ct. unicinctus Hlgr.
Ct. consobrinus Hlgr.
Ct. rufonotatus Hlgr.
Ct. gibbulus Hlgr.
Ct. gnathoxanthus Grav.
Ct. aurifiuus Hal.
++ x x |Holland
++ +4 |Finland
+ + + +4 | Switzerland
+++4+++4++4+4++4+ [England
O+t4¢44+ 4+4++4++4 |Belgium
+x+xXx xX
+4+4+4++++++4++4++4 | France
x+xx+4+4++4+4 x x |Czechoslovakia
+
eb
x +
+ ++x
+ ++
+x+
x
tet FHFFHEHHEHH++HE+ F++4+ |Sweden
i+ +
hb
t+tt +44 +4+4+44+4+ 4X14
+++4+4Xt+ 444444
b+t++++4x+ +44
O +O+++ ++ +++
l++++4++ ++
+++ +4+4++ 444444
x+ ++
ae a we
HEHEHE HHH HEHEHE FEXFHEHHHEHH FEE F++44+4+4+4+4+4+4 |Germany
+ + ++ + +44 444+
xxX+
Xf ++ + F+4+44+4+444
+
+ xX
++ 4+ ++ 444
++t+tt4+tt+4¢tt+4¢+ + +444
++ ++
OCOO +0 ++
-b
a
A more detailed analysis of the British fauna will be communicated to the Society for British
Entomology. A few additions from other countries were made in paged proof.
THE CTENISCINI OF THE OLD WORLD 459
APPENDIX II
Flight Table of European Species
This table is compiled from records known or accepted, and is set out in thirds of a month.
I have ignored all breeding records, on account of the influence of laboratory conditions; and
high Alpine and far northern records, north of 63°, on account of the influence of the shorter
summer season.
Dots indicate that the day of the month was not given. It will be appreciated that many of
the gaps in the table represent gaps in our knowledge. Some species are omitted, for which there
are very few dated records. Some species of Exenterus have been more often bred than caught,
and dates of capture are correspondingly few.
Apr. May June July Aug. Sept. Oct. Nov.
Acr. lucidulus agg.
Acr. succinctus
An. triangulatorius .
An. rubiginosus
An. pumilio . 2 . — —_—_—| — sh
An. ridibundus :
An. laticeps . : ‘ ° e
An. laetus agg. : : —_
An.alacer . é : =| eee
Exent. apiarius . P —
Exent. abruptorius . : ——
Exent. oriolus : : —
Exent. ictericus : x ——
Exent. amictorius . : _
Exent. adspersus . : —
Exent. tricolor ‘ : — —_ —
Exent. confusus . : — oo —
Exy. cinctulus ‘
Exy. genalis . : : _—
Exy. phaeorrhaeus .
Exy. albicinctus
Exy. pratorum
Parexy. calcaratus . F e
Smicro. 5-cinctus
Smicro. excisus ; , e —+——_—s« |— —
Smicro. trianguligena : —_—
Smicro. bohemani 5
Smicro. perkinsorum < ‘
Smicro. heinrichi . a —| —__—- —
Smicro. jucundus . : _ Cs
Ct. hofferi : : :-|—
Ct. curtistt : ;
Ct. romani : 3
Ct. elegans. ‘ ; —_—~+«+-—
Ct. unicinctus :
Ct. rufonotatus ‘ F °
Ct. gnathoxanthus . . —
Ct. aurifiuus 2 : —
2 2 DEC 1952
ee Noe hie eee Bie oe li a ge a ee ee re dy oe “Ce Te ee Cae
: : ; ee ths Ty) LS ee eg Se
x
a
a Py
PLATE 4. Exenterus abruptorius Thunberg.
PLATE 5. Exyston genalis Thomson.
PLATE 6. Swetzvopthctinds jidbatiecthitic Gravenhorst.
PLATE 7. Cteniscus cunitves Haliday.
ll. B.M. (N.H.) Ent. II, 6 ea st le
EXENTERUSUABRUPTORIUS THUNBERG
Bull. B.M.(N.H.) Ent. IT, 6
EXYSTON GENALIS THOMSON
PLATE §
ch.
ull. B.M. (N.H.) Ent. II, 6
SMICROPLECTRUS QUINQUECINCTUS
GRAVENHORST
PLATE:6
Bull. B.M (N.H.) Ent. II, 6 PLATE 7
CTENISCUS: AURIPLUUS., HALIDAY
PRESENTED
22 DEC 1952
INDEX TO VOLUME II
New taxonomic names ave printed in bold type
abruptorius, Exenterus _ es 365, pl. 4
abyssinica, Motes . : - es
abyssinicus, Philanthus . : 3 200
aceraensis, Pseudagenia . 169 (figs.), 171-172
Acrotomus . 311, 313, 314, 316-321, 452
acutangulum, Trypoxylon - nae
adspersus, Exenterus 360-361, 365 (fig.),
366, 367
Aédes . 3 235-304
Aedimorphus , ‘ ; < = 233
aegypti, Aédes ; "236-204 (passim)
aethiopicus, Tachysphex ; : », 257
aethiops, Hemipepsis . : ~ G67
africanus, Aédes 250- 294 (passim)
agrestis, Bembix . : “538
alacer, Anisoctenion 318 (fig.), 327 (fig.), 334,
348-350, 354
albicinctus, Exyston
albopictus, Aédes .
albosignata, Pseudagenia
aleyone, Cerceris
allonymum, Pison . : ; : io “207
Alysson = ; 120-122, pl. 5
amaltheus, Aédes . 238, 246, 247, 248
amictorius, Exenterus 358-359, 364, 365 (fig.),
382-384, 391, 392
238, 245, 247, 270
366
Ammatomus. : : ; : » 25
Ammophila . 141-143
amoenus, Stizus . ? ; ‘ 132
Ampulex : . Fi . : + 139
Ampulicinae . : : ‘ : » 339
anchietae, Hemipepsis . ‘ 167
angustus, Aédes : ey x a6, 247
Anisoctenion. 3II, 313, a ears 452-454
Anopheles. : » 270
Anoplius 180-181
antennatus, Cyphononyx = Gy
apiarius, Exenterus
apicoargenteus, Aédes
argyrocephala, Ammophila
356, 364, 365 (fig.)
238-294 (passim)
140 (fig-), 143
Armigeres . - ; F é = 238
Astata . ‘ ‘ : 3 a 3) £50
Astatinae : ; : . DTS
aurantiacus, Brachystegus . 124-125
aurifluus, Cteniscus 317 (Me. a; BOS pl. 7
avius, Dasyproctus . 3 . 165
bambusae, Aédes 241, 246, 249
barbarorum, Ammophila ~ (fig.), 141-142
basalis, Stizus ; ‘ - «if t3%
basifasciatus, Dasyproctus
basilica, Tachytes .
Batazonellus .
Bembicinae .
Bembix é
beniniensis, Ammophila A
bensoni, Stizus
bidens, Odontosphex
bohemani, Smicroplectrus
boreoalpinus, Eudiaborus
Brachystegus
brachystylus, Sceliphron.
braunsii, Stizus :
brevipalpis, Megarhinus .
bruneiceps, Tachysphex .
calcaratus, Anisoctenion
calcaratus, Parexyston
calceatus, Aédes
campsomeroides, Cerceris
capensis, Ammophila
capensis, Batazonellus
caprella, Ammophila
celaeno, Cerceris :
cephalotes, Anisoctenion
Cercerinae
Cerceris
Ceropalinae
chaussieri, Aédes
cinctulus, Exyston
cinerascens, Sphex .
cinnamomeus, Pompilus .
claripennis, Exenterus
clavatus, Exyston .
clementi, Smicroplectrus
coarctatus, Anisoctenion
coarctatus, Philanthus
compacta, Tachytes
confusus, Exenterus
coniferus, Oxybelus
consimilis, Philanthus
consobrinus, Cteniscus
contiguus, Aédes
contractus, Dolichurus
contrarius, Pompilus
corpulentus, Stizus
Crabro .
162 (fig.), 164
144 (fig.), 146
. + 298
132-138
133-138
: p » TBE
129 (fig.), 131-132
151 (fig.), 154-155
401-403, 415
417-420, 454
123-125, pl. 5
- 139
126, 127 (fig.)
‘ 270
155
347) 354
. 392-393
239, 240, 246, 249
109-110, pl. 4
141
178
140 (6g), 142-143
118, pl. 5
344- 346, 347, 353
. . 100-119
"100-119, pls. 4, 5
: : 183
245, 258
371-375, 389, 390 (figs.),
391 (fig.), 392
145
$ F . EST
359-360, 365 (fig.), 366
. : 375
404-405, 413 (fig.),
415
323, 325, 327
. - 99
148-149
361-364, 365 (fig.), 366,
367
162 (figs.), 165-166
100
436-437, 446
247, 249, 258
139
181-182
127
163-165
462
Crabroninae .
cretinus, Aédes
croesus, Motes
croesus, Palarus
Cryptocheilus 3
Cteniscini
Cteniscus
Ctenoceratinae
Culex :
curtisii, Cteniscus .
curviscutis, Oxybelus
Cyemagenia .
Cyphononyx .
dahlbomi, Cteniscus
Dasyproctus .
deboeri, Aédes
decolorata, Cerceris
delta, Aédes .
demeilloni, Aédes .
denderensis, Aédes
dendrophilus, Aédes
denticulata, Motes .
Diaborus
Diceromyia
dilaticornis, Tachytes
discrepans, Tachytes
distinctus, Philanthus
Dolichurus
dorsator, Cteniscus
Dunnius
egregia, Motes
egregius, Tachysphex
Elaphrosyron
electra, Cerceris
elegans, Cteniscus .
Eridolius . , >
Episyron “ °
Eudiaborus
Excavarus
excisus, Smicroplectrus
Exenterus
Exyston
faceta, Cerceris
faleigera, Tachytes .
fallax, Hemipepsis .
fantiorum, Bembix
ferox, Parapiagetia
ferrugineipes, Ammophila
ferrugineus, Stizus .
Finlaya = -
flavitincta, Motes .
flavonasuta, Cerceris
flavotegulata, Pseudagenia
Fleas, from British Birds
fluctuatus, Tachysphex .
240, 24r, 243, 247, 248
239, 240, 249, 278 (fig.),
INDEX
: 163-165
289 (fig.), 290
« 4§9
156 (fig), 160-161
- 168
3°7-459
311, 312, 314-317, 319, 341, 343,
359 351, 354 420-451
170-171
271, 275
Sense 444
s) 3566
> Ego
167-168
- 440
162, 163-165
sil 247, 249
105
290
281 (fig.), 282
238-294 (passim)
» 160
311, 314-316, 319
273
147
I51I-152
100
: 139
. . + 341
. 273
. . + 160
‘ 155
. ° - 178
II13-I14
439-432, 445
. =, Bae
177-178
"315, 319, 415-420, 454
312, 314, 315
"398-400, 412 (fig.), 415
- 311, 312, 318, 319, 354-367
308, 311, 313-316, 318, 367-392
I1I-112, pl. 4
151 (fig.), 152
- 167
134 (fig.), 135-136
. 157
141
. F wiv 532
240, 291
. 160
. 105-107, pl. 4
‘ on 2X92
187-232
155
fluviatilis, Gastrosericus
fraseri, Aédes
fraudulentus, Stizus
fueata, Tachytes
fulviventris, Cerceris
fulvo-hirtum, Sceliphron
funebris, Paracyphononyx
fuscipennis, Bembix
fuscistigma, Astata
fuscus, Anoplius . °
gambiae, Anopheles
genalis, Anisoctenion
genalis, Exyston
gibbulus, Cteniscus
glutiniatus, Eudiaborus .
gnathoxanthus, Cteniscus
Gorytes
gracilicornis, Motes
gracillima, Ammophila
granti, Aédes.
238, 244, 249, 289 (fig.), 290
156 (figs.), 157-158
239-294 (passim)
: 129-130
- 149
107, pl. 4
F) 139
173 (fig.), 177
- 137 (fig.)
.. Tag
180-181
270
350, 354, 452-454
375-378, 389, 390 (fig.), 391
(fig.), 392, pl. 5
443, 446
- re
317 (fig. ), 440-442,
444 (fig.), 445 (fig-), 446
125, pl. 5
- 160
I4I
Gripopterygidae [Index to names in Kim-
mins’s paper]
griseola, Tachytes .
guichardi, Alysson .
guichardi, Marimba
guichardi, Tachytes
Guichardia
guigliae, Bembix
habilis, Tachytes
haemorrhoidalis, Sphex .
haemorrhoidalis, Stizus .
hamiltoni, Tachytes
heinrichi, Smicroplectrus
heischi, Aédes
Hemipepsis
histrio, Episyron
hofferi, Cteniscus
ictericus, Exenterus
impressiceps, Cryptocheilus
incomptus, Sphex .
infantula, Pseudagenia
iniqua, Cerceris :
insidosus, Elaphrosyron .
insignita, Cerceris .
intermedia, Bembix
jucundus, Smicroplectrus
kaniacensis, Monoblastus
keniensis, Aédes
kenyae, Aédes ‘
kivuensis, Aédes
laetus, Anisoctenion
‘ 2 3 oe
144 (fig.), 149-150
120-122, pl. 5
169 (figs.), 170-171
144 (fig.), 146-147
178-179
136-138
144 (fig.), 147
. - 145
126
‘ ‘ . 149
408-409, 413 (fig.),
" 414
241, 246, 247, 248
167
‘ 177-178
421-424, 443, 444 (fig)
358; 365
168
145
172
110
178
104-105, pl. 4
: . 538
341, 409-414
ps5
241, aas, 246, 248
247, 249
. - 245
343-347, 352 (fig.)
INDEX 463
laevigata, Pseudagenia . ‘ ; a (192
laevigatus, Sphex 145-146
laevior, Poenulus . : , i > £63
lamentabilis, oe 144 (fig.), 148
Larrinae - 146-163
langata, Aédes ‘ 230, 247, 249, 258
laticeps, Anisoctenion 318 (fig.), 324, 334 (fig.),
336, 340, 341-343, 352 (fig)
latifasciatus, Ceropales . : ‘ 3; 285
latus, Ammatomus : 5 : ak eX
levis, Pilompus ; 182-183
lindtropi, Aédes_. P ‘ ‘ +) 290
lineiger, Cteniscus . 424-425, 443
Liris’ . : ; - 159-160
longicollis, Poecilagenia - 173 (fig.), 174-175
longipalpis, Aédes . A : : a 20
longiuscula, Cerceris IOI—102, pl. 4
lucidulus, Acrotomus 2 (*e.), 319-321
lutarius, Pompilus . “ - 3X8r
- luteocephalus, Aédes "241-201 (passim)
lutulenta, Cerceris . . ‘ ‘ sc ROR
macfiei, Culex : : . ‘ . Oar
macilenta, Guichardia 178-179
maia, Cerceris 114, pl. 5
Mallophaga [List of species in Clay and
Hopkins’s paper] : : 35-36
marginatus, Anisoctenion 346-347, 352 (fig.),
353
Marimba . 170-171
mascarensis, Aédes, aii 244, 248
massaica, Bembix . : ‘ 138
masseyi, Aédes_.. . é : - 246
mazimba, Cerceris . : x ‘ 1T2
Megarhinus . ‘ ‘ = A 3% 5270
melanurus, Exyston z F - 388-389
meneliki, Philanthus : : > 100
meridionalis, Hemipepsis . » 167
merope, Cerceris a yet 17, pl. 5
mesoleptoides, Anisoctenion 25
metallicus, Aédes “24 5-292 (passim)
mionii, Stizus . ‘ PRE i Y-
modestus, Priocnemis 168-170
moggionis, Cerceris 102-104, pl. 4
Monoblastus . : 311, 341
montanum, Pison . , ‘ P : 97
montivagus, Gorytes 125, pl. 5
morosus, Anoplius , ‘ ‘ 3/1380
Motes . ‘* 4 - . 159-160
multiguttatus, Stizus 126-127
neglecta, Tachytes . 148
Nemouridae [Index to names in "Kim.
mins’s paper] 93
129 (fig.), 130-131
nigriventris, Stizus .
nigrohirtus, Sphex . A «. 545
nordstromi, Cteniscus . ; , - 426
nugax, Cerceris : . 121, pl. 5
nugax, Motes §. : * . - 159
Nysson ' ‘ : » 422-123, pl. 5
Nyssoninae 120-125
obesus, Palarus ? : 161-163
observabilis, Tachytes . é ‘ . 3146
occidentalis, Dasyproctus 162 (figs.), 164
octomaculatus, Anoplius 179 (figs.), 180
Odontosphex . ; 154-155
oedignathus, Dasyproctus : ; «. “£63
cptimus, Cyphononyx . : ‘ + 168
oriolus, Exenterus . + 357,364
oscari, Paranysson 162 (figs.), 163
Oxybelinae 165-166
Oxybelus 165-166
Palarus : : ‘ 160-163
pallidus, Oxybelus . : 2 ; . 166
Paracyphononyx . : ; - a 7
Paranysson . . . : » 163
Parexyston ‘318, 367, 392-393
pectoralis, Cteniscus 432-434, 445
Pemphredoninae . : : . - 163
Pepsinae 167-170
perkinsorum, Smicroplectrus 405-408, 412 (fig.),
413 (fig.), 414
phaeorrhaeus, Exyston 378-380, 381 (fig.), 380,
pias 391 (fig.), 392
Philanthinae . 4 Z F 99-100
Philanthus 99-100
Phytosphex 153, 154
pictus, Cteniscus ; : - 449-450
pictus, Oxybelus . : ; : . 166
Pilompus : : : ; . 182-183
Pison . = 2 : oy 07
planatum, Sceliphron z 139-140
plumbeus, Pomphilus . : : o LOE
Poecilagenia . : c : - 174-176
Polyblastus . - 2 ; : - 308
Pompilidae 166-183
Pompilus ‘ . 81-182
pratorum, Exyston 38 5-388, 390 (fig.), 391,
392
Priocnemis 168-170
pronotalis, Anisoctenion errs a38-a4n 352-3 53
propinquus, Oxybelus
Pseudagenia . eae
Pseudageniinae : 171-174
pseudoafricanus, Aédes 242, 243, 245, 252,
270, 276
pseudocyphononyx, Pseudagenia . ee
pseudonigeria, Aédes > 243, 245
pumilio, Anisoctenion 317 (fig.), 331-334, 336,
351
punctaticeps, Ammophila : = EE
punctipleuris, Cteniscus . é : - 34t
quadriceps, Smicroplectrus. : - 414
quadricolor, Nysson 122-123, pl. 5
quinquecinctus, Smicroplectrus 395-397, 398
(fig.), 412 (fig.), 413 (fig.), 415, pl. 6
464 INDEX
reticulata, Liris . : . ‘ - 160
reversa, Gastrosericus. ‘ , ee
rhodesiae, Cerceris . 108, pl. 4
ridibundus, Anisoctenion 323, 336-338, 340,
352, 452
ritzemae, Stizus_. P ‘ <0) aga
robustus, Stizus 128, 129 (fig.)
romani, Cteniscus . 429-430, 444, 445 (fig.)
rubidus, Philanthus ‘ - Foo
rubiginosus, Anisoctenion 328-331, 334, 340,
351
rubrozonata, Cyemagenia ° a AG
rufitibialis, Tachytes 144 (fig.), 150
rufonotatus, Cteniscus 351, 437-440, 446
rugifera, Motes. : - 159
rugosa, Poecilagenia 173 (figs.), 175-176
ruwenzori, Aédes . ‘ ‘ + 245
Sceliphron . . 139-140
Schistonyx . Mes,
schwetzi, Aédes_. 230, 246, 249, 278 (fig.), 281
seminuda, Tachytes : 1 £63
senegalensis, Brachystegus 123-124, pl. 5
senegalensis, Cryptocheilus < é 2 | 50S
senegalensis, Gastrosericus 156 (fig.), 158-159
senegalensis, Sphex 144-145
setigera, Motes ; : : = . 160
severini, Cerceris . : : oe EOg
sibiricus, Smicroplectrus . 403-404, 415
similis, Cteniscus 354, 445
simoni, Sphex ; : a A - 145
simplex, Exenterus 350-357
simpsoni, Aédes 237-294 (passim)
Smicroplectrus 308, sc 319, 341, 393-415
soleatus, Aédes 239, 246, 249
solstitialis, Motes . Z . - 159
Sphecidae : 97-166
Sphecinae : : : 139-146
Sphex . : 5 : - 144-146
spinicaudata, Cerceris ; , : AON i
splendida, Bembix . 133-134
Stegomyia 235-304
sterope, Cerceris ° Mare 18, pl. 5
stigmalis, een = : - TES
Stizinae & - 126-132
Stizus . : “ ; - 126-132
strelitziae, Aédes 245, 248
subargenteus, Aédes 245, 248, 272
subnitidus, Exyston 380-382, 391, 392 —
successor, Anoplius : : ; . en
succinctus, Acrotomus 317 (fig.), 318 (fig.),
319-321
sudanensis, Stizus . 127-128
Tachysphex . ‘ > 155-157 2
Tachytes . F . 146-153
taygete, Cerceris 119, pl. 5 3
tenuis, Anisoctenion 334-336, 352
theseus, Tachysphex 150-157
thysanomera, Motes ‘ . ‘3 - ae
toroensis, Ampulex . ° : -- 36
transvaalensis, Motes . a - ESO
irinngulatoring, Anisoctenion 317 (fig.), 323-
328, 340, 351, 352 (fig.)
trianguligena, si tila
(fig.), 415
tricolor, Exenterus P . 361, 366, 367
trinidad, Aédes : 240, 269 —
Trypoxylon . * - 97-99, 310
Trypoxyloninae
turneri, Phytophex
97-99
. . ~) 2a
.
ugandicus, Philanthus ., . A ~ Leo
umbrosus, Schistonyx . < ‘ . 59S
unicinctus, Cteniscus , 434-436, 445
uniguttatus, Dasyproctus is ; . 1G
unilineatus, Aédes .
variolosus, Ceropales. . 182 (fig.), 183
vellicatus, Exenterus 361, sie (fig.), 366, 367
versatilis, Tachytes . TAF
viduum, Trypoxylon u 97-99
vittatus, Aédes, 2 " 238-204 (passim)
walegae, Cerceris . - "
woodi, Aédes ‘ Z : ; » <a
xanthopus, Pison . : : % | ee
yakui, Anisoctenion falar 351, 352 (fig.) 4
yungvei, Cerceris . 112, pl. 5
ypsilon, Bembix . : ‘ . » 238
zavattarii, Cerceris . é‘ 107-108
zonatus, Paracyphononyx , : ee
PRESENTED
15 OCT 1955
400-401, 413 _ 3
238-291 (passim)
T10-11IT
OXFORD
BY
CHARLES BATEY
PRINTER
UNIVERSITY
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