BULLETIN OF

ENTOMOLOGY
VOL III

1953-1954

PRINTED BY ORDER OF THE TRUSTEES OF

THE BRITISH MUSEUM (NATURAL HISTORY)
LONDON: 19^3-1954

DATES OF PUBLICATION OF THE PARTS

No. i. 17 June 1953

No. 2. 10 October 1953

No. 3. 20 November 1953

No. 4. ii December 1953

No. 5. 4 December 1953

No. 6. 12 March 1954

No. 7. 27 April 1954

No. 8. 23 November 1954

No. 9. 22 October 1954

No. 10. 23 November 1954

No. 11. 23 November 1954

No. 12. 30 November 1954

No. 13. 21 December 1954

No. 14. 31 December 1954

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GREAT BRITAIN

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CONTENTS

ENTOMOLOGY VOLUME 3

No. i. The sub-genus Stegomyia (Diptera : Culicidae) in the Ethiopian
Region. Part II. Distribution of species confined to the East
and South African sub-region. By P. F. MATTINGLY i

No. 2. A pest of coconut palms in Portuguese East Africa.

Two new representatives of the genus Hypotrabala (Lepidoptera :
Lasiocampidae) from the Belgian Congo. By W. H. T. TAMS (Pis. 1-2) 67

No. 3. A revision of the genus Teriomima Kirby (Lepidoptera : Lycaenidae).

By H. STEMPFFER and N. H. BENNETT (Pis. 3-9) 77

No. 4. Notes on British Ichneumoninae with descriptions of new species

(Hym., Ichneumonidae). By J. F. PERKINS 103

No. 5. On a very remarkable flea from Argentina. By H. E. KARL JORDAN
Descriptions of new and little-known Siphonaptera. By F. G. A. M.
SMIT 177

No. 6. The early literature on Mallophaga. Part III. 1776-86. By

THERESA CLAY and G. H. E. HOPKINS (Pls.io-i2) 221

No. 7. Some sawflies of the European Alps and the Mediterranean Region

(Hymenoptera : Symphyta). By R. B. BENSON 267

No. 8. Neue Notiophygidae (Coleoptera). Von HANS JOHN (Pis. 13-17) 297

No. 9. The Protoptila group of the Glossosomatinae (Trichoptera : Rhyaco-

philidae). By MARTIN E. MOSELY 315

No. 10. A review of the Aedes scutellaris subgroup with a study of variation
in Aedes pseudoscutellaris (Theobald). (Diptera : Culicidae). By
ELIZABETH N. MARKS (PI. 18) 347

No. ii. A new genus and some new species of the Chauliodini (Megaloptera).

By D. E. KIMMINS 415

No. 12. The genera Henricohahnia Breddin, Dicrotelus Erichson, Nyllius Stal,

Orgetorixa China and allied new genera. By N. C. E. MILLER 445

No. 13. Revisional notes on the genus Epitola Westwood (Lepidoptera :

Lycaenidae). By PATRICK ROCHE (Pis. 19-22) 489

No. 14. African species of the genus Pardomima Warren (Lepidoptera : Pyra-

lidae : Pyraustinae). By EDWARD L. MARTIN (PI. 23) 503

Index to Volume 3 523

f.

CORRIGENDA

P. 141, line 13. For " Ctenichneumon rubroator " read " Ctenichneumon rubroater."
P. 149, last line. For " Pseudamblyteles " read " Pseudoambly teles."
P. 160, line 10. For " 8. vm. 1936 " read " 10. x. 1935."
P. 171, column i, last line. For " corruscator " read " coruscator."

P. 173, column 2, line 35. For " Cyclolabus Heinrich 1935 " read " Cyclolabus Heinrich 1936."
P. 181. For " Barreropsyilini " read " Barreropsyllini."
P. 445 (title-page). For " Pp. 445-488 " read " Pp. 445-486."
P. 487 (title-page). For " Pp. 489-502 " read " Pp. 487-502."
P. 503 (title-page). For " Pp. 503-521 " read " Pp. 503-522."

2 4 JUN 1953

THE SUB-GENUS STEGOMYIA
(DIPTERA: CULICIDAE)

IN THE
ETHIOPIAN REGION

(PART II)

P. F. MATTINGLY

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 3 No. i

LONDON: 1953

THE SUB-GENUS STBGOMYIA

(DIPTERA: CULICIDAE)
IN THE ETHIOPIAN REGION

II. DISTRIBUTION OF SPECIES CONFINED TO
THE EAST AND SOUTH AFRICAN SUB-REGION

BY

P. F. MATTINGLY

Pp. 1-65 ; 15 Text-figures

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 3 No. i

LONDON: 1953

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

Parts 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. 3, No. i of the Entomological
series.

PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM

Issued May, 1953. Price Fifteen Shillings.

THE SUB-GENUS STEGOMYIA (DIPTERA
CULICIDAE) IN THE ETHIOPIAN REGION.

II. DISTRIBUTION OF SPECIES CONFINED TO
THE EAST AND SOUTH AFRICAN SUB-REGION

By P. F. MATTINGLY

CONTENTS

INTRODUCTION . .

NOTES ON TAXONOMY ......

DISTRIBUTION RECORDS .....

Doubtful records ......

Records based on misidentifications

Unidentified and misquoted localities

Distribution outside the Ethiopian Region
LIST OF LOCALITIES WITH TOPOGRAPHICAL DETAILS
BIONOMICS IN RELATION TO DISTRIBUTION .

Breeding-places ......

Seasonal distribution .....

Biting-habits ......

ZOOGEOGRAPHY . . . . .

SUMMARY ........

APPENDIX I. Further notes -on species occurring in

Sub-Region ........

APPENDIX II. Rainfall of the Bor-Pibor-Torit area ....

APPENDIX III. Distribution of pale forms of Aedes aegypti in the Ethiopian
Region .........

APPENDIX IV. Corrigenda to Part I .......

ACKNOWLEDGMENTS ..........

REFERENCES

the West African

Page

4

4

18

22

25
25
26
26
28
28
29
30
31
50

51

57

60
6l

62

62

SYNOPSIS

The distribution of Stegomyia spp. occurring in the West African Sub-region has been dis-
cussed in the first paper of this series. The present paper deals with the remaining Ethiopian
species, i.e., with those which are restricted to the East and South African Sub-region. As
before, zoogeography is discussed mainly in relation to rainfall and altitude, and such notes
on bionomics are included as are thought necessary for an understanding of distribution. One
new species closely related to Aedes aegypti is described from Mauritius, and the male of Aedes
woodi, larvae which may prove to be those of pseudonigeria and masseyi and a new subspecies
of dendrophilus from Kenya are described for the first time. Previously unpublished taxo-
nomic data concerning most species are included. An appendix is devoted to the distribution
of pale forms of Aedes aegypti in the Ethiopian Region, and other appendices deal with recent

ENTOM. Ill, I. I

4 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

information concerning the species treated in the first paper and with the rainfall of the Bor-
Pibor-Torit area. The distribution of certain Palearctic Stegomyia is discussed in relation to
the problem of the origins of the Ethiopian fauna and an account is given of a specimen of
Aedes cretinus.

INTRODUCTION

THE present paper follows the pattern of its predecessor (Mattingly, 1952), but is
concerned with those species which have not as yet been recorded from anywhere
in the West African Sub-region. The number of species discussed is greater than
in the first paper but much less is known about them, and this is true of every depart-
ment of mosquito studies. In the field of distribution certain territories, notably
Tanganyika and Nyasaland, which are of vital importance, are almost unknown.
With respect to taxonomy four species, poweri, masseyi, chaussieri and pseudo-
nigeria, are, at the time of writing, known with certainty only from the adult female,
although there is good reason to hope that males and early stages of all of them may
very soon be described. The early stages of woodi are still unknown, but again
there is good hope that these will soon be available. Among these species the
first four are all likely to prove of special value for our understanding of the origins
and affinities of the sub-genus. In the field of bionomics no studies comparable
with those carried out by the yellow fever institutes in Uganda and, to a less extent,
Nigeria are as yet available. The present paper, like its predecessor, attempts to
bring together all available information relating to Stegomyia distribution as a
preliminary to the taxonomic study which will form the subject of further papers
in the series. It is clear that a review of the sub-genus as a whole will be necessary
and this will be undertaken in the third paper of the series. In the present paper
all that is attempted is a partial revision of group A, the principal Ethiopian group,
and some attention is paid to the distribution of certain Palearctic species, since it is
apparent that even this group cannot be fully understood against a purely Ethiopian
background.

To save repetition the list of localities with topographical details and the list of
references have been restricted to information which has not already been published
in Part I. Keys are also omitted since these have already been published. The
map of the distribution of collectors (Mattingly, 1952, fig. i) is not thought to have
been sufficiently affected by new records to be worth reprinting. It can be brought
up to date almost completely by adding those localities listed on p. 56 of the present
paper together with the Kologha Forest, Njombe and the Tzitzikama Mountains.

NOTES ON TAXONOMY

The necessity for a revision of the main groups of the sub-genus as defined by
Edwards (1932) has already been noted (Mattingly, 1952), and it is now possible to
make certain concrete suggestions for a partial revision of the main Ethiopian group,
group A. For reasons given below it is felt strongly that this should be extended
by the inclusion of Aedes mascarensis from Edwards' group B, together with the
closely related species from Mauritius which is here described for the first time (p.
16). At the same time it is felt that if the present groups are to be maintained

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 5

then Aedes chemulpoensis Yamada should be transferred from group B to group A.
This species is at present known only from North-eastern China, the Mukden area
and Korea, but it is a perfectly typical member of group A with respect both to its
scutal markings and its male genitalia. Yamada in his type description (Yamada,
1931) noted the resemblance of the scutal markings to those of simpsoni and poweri,
and Feng (19386) again noted its affinities with group A in the first published descrip-
tion of the male terminalia. Edwards seems to have been unacquainted with the
male terminalia when he drew up his classification, since the only male specimen
in the British Museum of earlier date than Feng's description is undissected. In
its femoral and tibial markings chemulpoensis resembles mttatus and it is thus an-
nectant to group D. The structure of the ventral brush of the larva is somewhat
peculiar, but in other respects this stage also is characteristic of group A.

The occurrence in Eastern Asia of a member of group A, which, if we except the
cosmotropical Aedes aegypti, is otherwise entirely Ethiopian, may at first sight
seem rather startling, but taken in conjunction with such other facts as the occur-
rence of mttatus in Spain, North Africa, the Balearic Islands, Corsica and Sardinia
and of an isolated species of Group C (cretinus) in Crete, Macedonia and Trans-
caucasia (Georgia), it is in reasonable conformity with prevailing ideas concerning
the pre-glacial distribution of Palearctic woodland faunas. Basing his ideas on
the present distribution of loess, Reinig (1936) has pictured the occurrence during
the Glacial Period of a broad belt of desiccation with, at its ends, a Mediterranean
and an East Asian wooded refuge. This author figures certain smaller refuges in
between (loc. cit., fig. 12), but it would seem that among these the Armenio-Persian
was glaciated at least in part during the maximum extension of the ice sheets (com-
pare, for example, Furon, 1943, fig. 20). De Beaufort (1951) summarizes Reinig's
theories but does not mention these smaller refuges. The present distribution of
isolated populations of groups A, C and D in the Palearctic is in close conformity
with Reinig's views. Aedes mttatus exactly occupies the western portion of his
Mediterranean refuge, and chemulpoensis fits squarely into his East Asian refuge.
The Cretan and Macedonian form of cretinus fits well into the eastern portion of his
Mediterranean refuge, but the form (lindtropi Schingarew) from Georgia assigned
by Baschkareva (1931) and Stackelberg (1937) to this species occurs much too far
east and is perhaps to be associated rather with the Armenio-Persian refuge. It
does not seem certain that this form is in fact conspecific with cretinus, although
it may be that the present distribution shows the effect of post-glacial migration,
or that Reinig's Mediterranean refuge ought to be extended to include the southern
fringes of the Black Sea. Unfortunately only a single much damaged specimen of
cretinus from any part of its range is at present available for examination (see
below under Aedes albopictus, p. 18). Examples of affinities between the Mediter-
ranean and East Asian refuges quoted by De Beaufort are the Blue Jay, Cyanopica
cyana, with two subspecies in Spain and Portugal and five in South-East Asia and
Japan, none occurring in between, and the Barbary Ape (Macaco, sylvana = M.
inud] of Gibraltar with relatives confined to Southern Asia and Japan. It is to
be noted that in both these cases, as in the case of mttatus and chemulpoensis, the
East Asian fauna is represented only in the western and not in the eastern part

6 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

of the Mediterranean refuge. In fact it seems that in these cases the western and
eastern portions of the Mediterranean refuge constitute distinct faunal areas. This
discontinuity between the western and eastern Mediterranean refuges is further
illustrated by the case, quoted by De Beaufort later in his book, of the mountain
goat, Capra pyrenaica, which occurs in the Pyrenees and the higher ranges of the
Iberian peninsula and has relatives in Eastern Europe, Crete and the Caucasus
but none in the Alps or Appenines. Finally De Beaufort mentions the work of
Caradja (1934), who traces affinities between the Lepidoptera of South-west Europe
and those of Eastern Asia and observes that they are more primitive than those
occurring in between.

On the basis of these facts it may be said that there is evidence for a much wider
pre-glacial extension of groups A and D, and perhaps also C, in the Southern
Palearctic. It may be noted that cretinus also has a relative (flavopictus of Japan
and Korea) in the East Asian refuge, but it is preferred to postpone discussion of
this point to a later paper dealing with the general problem of affinities within the
sub-genus. All that is intended here is to attempt to justify the inclusion of
chemulpoensis in group A and to show that this is defensible on distributional as
well as on morphological grounds. The eastern rather than western Palearctic
affinities of group A is a matter of some general zoogeographical interest, since a
similar phenomenon has been encountered in some other groups (see, in particular,
Uvarov, in press).

Aedes amaltheus. This very remarkable species possesses mesonotal markings
typical of Edwards' group A and male terminalia of the type found in his groups
B and C. The difficulty of fitting it into Edwards' system has already been noted
(Mattingly, 1952), and it will be necessary to deal with this problem in the paper to
be devoted to the discussion of the taxonomy of the whole sub-genus in relation to
that of the Ethiopian species. Here it may be noted that the only comparable
species (Aedes galloisi Yamada) again occurs in Reinig's Easter Asian refuge, being
confined as far as is known to Japan. It does not appear, however, that the meso-
notal markings of this species are absolutely typical of group A. No specimens are
available for examination, but published figures (e.g., in Stackelberg, 1937) show
the supra-alar patches extending forward to fuse with the anterolateral patches and
backwards to the scutellum. This type of pattern is characteristic of a number of
Palearctic, Ethiopian and northern oriental Finlaya. The male terminalia and the
larva (Sasa and Kano, 1951) appear to be quite typical of group C. The morpho-
logical characters of amaltheus have been well described by De Meillon and Lavoi-
pierre (1944), and nothing can usefully be added here except to note that the larva
possesses certain interesting and possibly primitive characters recalling vittatus on
the one hand and unilineatus on the other, e.g., the branched antennal seta (see
Hopkins, 1952), strongly branched saddle hair, spinose saddle edge and tendency to
development of precratal tufts.

Aedes pseudonigeria. The larva from Magoebaskloof, Transvaal, attributed to
this species by Ingram & De Meillon (1929) and subsequently to demeilloni by
Edwards (1936), seems almost certainly to have been wrongly attributed in both
cases. It was described by the present author (in Hopkins, 1952) as ? pseudonigeria

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 7

purely to avoid introducing another name into the synonymy. In fact, however,
it is virtually identical with the larva of heischi and almost certainly belongs to a
member of that group. Recently larvae collected by Dr. De Meillon in the Karabib
area of South- West Africa have been sent to the British Museum. There is some
reason to believe that they may be those of pseudonigeria. Other identical larvae
have been taken by De Meillon together with adults of pseudonigeria at Francistown,
Bechuanaland (per fide Muspratt). In both cases the larvae are unassociated, and
I do not think it possible to rule out the possibility that they are those of unilineatus,
which they closely resemble. It is true that unilineatus has not been found in
S.W. Africa, but it may well be that collecting there has been confined to altitudes
above its, apparently rather restricted, limits. The rainfall in the Karabib area
appears to be rather low even for pseudonigeria, but might well not be too low for
unilineatus, which is an exceedingly drought-resistant species (see Appendix I).
For these reasons I am unwilling to attribute the Karabib or Francistown larvae to
pseudonigeria until liason material is available for comparison. A character
distinguishing the Karabib larvae from those of unilineatus as at present known is
the very small number of pecten spines (4-5 in the available material), and this may
prove diagnostic should the identification be confirmed. For a final elucidation
of the situation it is still highly desirable that the identity of the Magoebaskloof
larva should be established by breeding out. The male of pseudonigeria is still
unknown. It should be noted that the " white spot " on the middle femur men-
tioned by Edwards (1941) in his key is not a definite spot of the kind found, e.g., in
unilineatus or calceatus, but an irregular patch or streak which is likely to be rather
indefinite in some specimens.

Aedes chaussieri. The male and early stages of this species are still unknown,
but as it has recently been found in gallery forest near Elisabethville (Lips, in litt.}
there is reason to hope that they may soon be obtained. It is difficult to draw any
conclusions as to its relationships until such material is available, and this is unfor-
tunate as the species is likely to prove a very interesting one.

Aedes masseyi. The male is still unknown, but this species has recently been
found in the same forest as chaussieri. In view of its interesting resemblance to
amaltheus on the one hand and to keniensis on the other further material will be
eagerly awaited. The species is discussed below under keniensis.

Aedes keniensis. Van Someren (1946 bis) quotes the presence of pale scaling
round the edges of the pre-scutellar bare space as a distinctive character from
masseyi. An examination of the type series of the latter species shows, however,
that, at least in some cases, the absence of pale scales from this region is due to
rubbing. One paratype from Ruwe clearly shows a thin border of narrow yellow
scales. In the single specimen from Elisabethville in the British Museum even
this very tenuous border is reduced and only one or two yellow scales are visible.
A better distinguishing character concerns the third hind tarsal segment, but it
should be noted that, as indicated by Edwards (1941), this is not always entirely
dark in masseyi, because it has a few pale scales below at the base in the Elisabeth-
ville specimen. The specimens from Nairobi tentatively attributed by Edwards
(1941) to masseyi are quite clearly keniensis. Both they and the type series of

8 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

keniensis have the third hind tarsal pale all round on about the basal \ to . Two
specimens recently received from Njombe have this segment pale above on about
the basal ^ to \ but are more narrowly pale below. The key to adults given in the
first paper of this series should be amended accordingly. For a further note on
these specimens see below under " Doubtful Records " (p. 22). Two unassociated
larvae, recently received from Elisabethville through the kindness of Monsieur Lips,
are believed, from their resemblance to keniensis, to be almost certainly those of
masseyi. They differ from keniensis in having strongly sclerotized pecten spines,
typically with a single strongly developed secondary denticle (occasionally with
1-2 minute ones dorsally or ventrally in addition) and a very short unbranched
saddle hair. On the key to larvae (Mattingly, 1952) they would run down to aegypti
and mascarensis, from which they can be recognized by the structure of the pecten
spines and saddle hair. They came from the township area, and were associated
in the one case with larvae of Aedes schwetzi and in the other with those of aegypti.

A fuller description will be published elsewhere (Mattingly and Lips, in press).

Van Someren (1946 bis) has suggested that keniensis may ultimately prove to be
a subspecies of masseyi, and the occurrence of the former at Njombe strongly suggests
that it is in fact a highland representative of the latter. At the same time it should
be noted that masseyi shows some very interesting resemblances to amaltheus, from
which it differs only in scutal characters. Geographically it is more or less inter-
mediate between the two species. Further light on its affinities may be expected
from the discovery of the male and early stages.

Aedes heischi. As noted above, the larva from Magoebaskloof, Transvaal, attri-
buted by Ingram and De Meillon (1929) to pseudonigeria is indistinguishable from
that of the present species. In view of the very close resemblance of the larva of
keniensis, noted by Van Someren (1951), however, it would be most unsafe to make
a definite attribution until further larvae have been collected and bred out. It
should be noted that, as indicated by Van Someren, heischi is not always separable
with certainty from Aedes demeilloni in the adult stage, since occasional specimens,
among them several in the British Museum, have no pale scales, apart from the
knee-spot on the anterior surface of the mid femur.

Aedes demeilloni. This was treated by Edwards (1941) as a subspecies of deboeri
but it is clearly a distinct species. Differences between the adults are slight but
apparently constant. The larval differences are striking, and there appears to be
a constant difference in breeding-places since demeilloni has so far only been found
in plant axils, especially those of Dracaena. As noted in the previous paper, con-
fusion of this species with dendrophilus (Van Someren, 1946 ; Smithburn & Haddow,
1946 ; Haddow et al., 1947) arose from the misleading description of the latter
given by Edwards (1941), and in particular from this author's failure to mention
the spines which occur on the basal lobe of the male coxite in both species. Further
confusion has arisen from the fact that fig. 6ic of Hopkins (1936), although labelled
" deboeri var. demeilloni," was in fact drawn from a larva of angustus, while fig. 6ie,
labelled " unilineatus," was drawn from a larva of demeilloni.

Aedes subargenteus. The larva of this species has previously been known only
from a single specimen from Nyasaland bred from an egg laid by the type female.

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 9

Through the kindness of Mr. Muspratt, who has recently sent six larvae from
Pondoland to the British Museum, it is now possible to amplify the description given
by Hopkins (1936, 1952). Mr. Muspratt states that the very pale colour of the
head and siphon noted by Hopkins is constant. Antennal seta single. Head
setae A, B and C single, d single or split distally into 2-3 branches. Mentum
rounded, the teeth rather small, 12-13 teeth on either side of the median tooth.
Pleural spines very small and pale in colour. Setae on anterior margin of pro-
thorax normally developed. (They are all missing from the Nyasaland larvae, so
that Hopkins' statement that they were small and inconspicuous seems probably to
have been due to an error of observation.) Comb with 8-10 teeth resembling those
of the Nyasaland larva but none of them bifid. Siphonal index (uncrushed) 2 -4-2 -7.
Pecten with 7-13 teeth proximal to the subventral seta and 1-2 detached teeth
distal to it. Subventral seta bifid and simple -or single and plumose. In one case
both conditions are shown by the same larva. Saddle hair single. Upper caudal
seta with 3 branches, lower with 2. Setae of ventral brush each with at least 2
branches. Distal edge of saddle with only light sculpturing, i.e., without obvious
spiculation. Gills large, sub-equal, sausage-shaped.

Aedes kivuensis. This is still known only from the type specimen in the Congo
Museum at Tervuren. Through the kindness of the Director and Monsieur Basilew-
sky it has been possible to examine this specimen and the following details may
be added to the description given by Edwards (1941) : Anterior median spot of
mesonotum composed of narrow scales. Anterolateral pale patches small and
narrow, their posterior ends terminating well in front of and outside the postero-
lateral lines. The latter strongly marked, deep yellow in colour, as in the case of
the median lines, and reaching forward almost to the scutal angles. Supra-
alar patches small and entirely white. Median lobe of scutellum with a few black
scales posteriorly, lateral lobes entirely white. Scale patch on posterior pro-
notum very small. Abdomen too shrunken to reveal the tergal bands. Fore-leg
with the tibia narrowly but completely ringed at base. First two tarsi uniformly
ringed at base, the second white on almost the basal half. Mid-leg with a well-
developed white spot beyond half-way on the anterior surface of the femur. Femoral
knee-spot small but reaching the tip above. Tibia entirely dark. First and second
tarsi uniformly ringed at base, the second white on at least the basal half. Hind
leg with femur extensively pale at base, with a small white spot just beyond the
tip of the basal pale stripe on the anterior surface and a small knee-spot which
reaches the apex above. Tibia entirely dark. First three tarsi narrowly pale at
base ; the fourth and fifth entirely pale except for a small black spot at the tip
below in each case. In view of our almost complete ignorance of the distribution
of this form it is preferred to treat it for the present as a distinct species.

Aedes woodi. A long series of adult females has now been received from Ganda
through the kindness of Mrs. E. C. C. Van Someren. A full description will be
published in the appropriate taxonomic paper of the present series. All the speci-
mens are immediately distinguishable from those of any other species, having a
double median line of yellow scales on the scutum by the dark-scaled lateral lobes
of the scutellum. Only two males have so far been received and neither of these

,io THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

is complete. The palps resemble those of simpsoni rather than strelitziae in having
a number of long setae on the two distal segments. The lateral lobes of the scutellum
are dark scaled. The scutal ornamentation has been almost entirely obliterated in
both specimens, but one of them appears to have the posterior ends of the sub-
median lines white or whitish and to have the pre-scutellar bare space rather
broadly bordered by white scales in addition to some yellow ones. There is some
variation in both respects in the females and comparable variation occurs in Aedes
simpsoni. Segment II of the abdomen has a few pale scales at the base of the tergite
in one specimen. In the other the tergite is rubbed. The pale basal bands on
tergites III-VI are shallow as in the female. Those on V and VI are markedly
narrower in the males. Sexual dimorphism with respect to the tergal bands is
shown by a number of Ethiopian species of the sub-genus. The male terminalia
appear to be virtually identical with those of simpsoni. The legs of both sexes are
very much as in the type. The femoral knee-spots are very short, and are separated
from the tips of the femora by distinct dark areas. The second mid-tarsal and
the third hind tarsal are pale above on about the basal -| and f respectively and both
are rather narrower below. The fourth and fifth hind tarsi are missing from both
males. In the females they are entirely dark and entirely pale respectively, as in
the type. Attempts to find the breeding-places have hitherto proved unsuccessful
and the early stages are still unknown.

Aedes strelitziae. This has been fully described by Muspratt (1950).

Aedes poweri. The type female is still the only specimen that can be assigned
with certainty to this species, although further specimens collected by Mr.
Muspratt in Cape Province, and discussed below under " Doubtful Records," appear
almost certainly to belong to it. The affinities of the species are uncertain although
it seems to be most nearly related to contiguus, which it resembles very closely.
Certain characters quoted by Edwards are in fact artefacts. Thus the interruptions
to the pale border round the eyes (Edwards, 1923 bis) are due to rubbing as,
apparently, is the absence of pale scales from the border of the pre-scutellar bare
patch (Edwards, 1941). The appearance of the mid-lobe of the scutellum is rather
striking, as it possesses more dark scales than are found in the majority of species.
Such scales are, however, possessed by all the Ethiopian Stegomyia, and their
complete absence from individual specimens is the exception rather than the rule
although they are seldom mentioned in descriptions ; contiguus, in particular, often
shows them well. The most distinctive character shown by poweri is the possession
of a conspicuous pale basal stripe on the lower surface of the middle tibia. This
is between and i of the tibial length. The third mid-tarsal is quite extensively
pale above at the base, but pale scales in this position are not uncommon as an
aberration in a number of species. Since the completion of this paper Mr. Muspratt
has sent descriptions of the larva and male terminalia which confirm the relation-
ship to the langata-contiguus group suggested below under " Distribution." It
seems that the terminalia are indistinguishable from those of contiguus, while the
larva is distinguishable only on variations in the statistical distribution of the
number of branches in head seta d. Variations in colour characters of the adults
include the constant suppression of pale scaling on the distal half of the second

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION n

mid-tarsus in the male, occasional presence of pale scales round the pre-scutellar
bare space and very occasional presence of a pale spot in the middle of one or both
front femora.

Aedes contiguus. As noted by Van Someren (1946 bis), this species closely
resembles langata. It can, however, be separated on the characters given in the
key (Mattingly, 1952). The anterolateral scutal patches are also rather smaller
and less wedge-shaped (see pi. ii, fig. n of Edwards (1941) which apparently repre-
sents contiguus and not poweri as stated). On the basis of these characters it seems
fairly certain that the two rather rubbed females of " contiguus " mentioned by
Edwards (1941) are in fact langata. The presence of the latter species in Southern
Rhodesia (Ndanga) has been confirmed by the dissection of male terminalia. The
presence of contiguus in the Transvaal has also been confirmed by the dissection of
male terminalia. Transvaal specimens tend to have the pale band at the base of
the third hind tarsal rather broader than do those so far seen from Southern
Rhodesia ( the length of the segment in the type male and the specimen from
Ndanga and from ^ to | the tarsal length in those specimens so far received from the
Transvaal) . This band is therefore not always quite so strikingly narrow as Edwards'
description might suggest. A very fine series of Transvaal contiguus has recently
been received from Dr. De Meillon, and the question of variation will be discussed
in more detail in the taxonomic paper of the present series.

Aedes langata. Resemblances to contiguus have been discussed in connection
with that species. The length of the pale basal band on the third hind tarsal is
again variable (about the length of the segment in the Kenya form, \ this length
in the problematical specimen from Nyasaland and \ in the Southern Rhodesian
specimens. This character cannot therefore be safely used for separation from
contiguus. An incomplete female from Gwelo in the British Museum resembles
langata in scutal markings and in the broad abdominal bands and distal expansion
of the ventral pale stripe at the base of the hind tibia, but differs from all other
available specimens in having a pale posterior stripe on about the basal f of the
first mid-tarsal, much as in heischi. This specimen and another, even less complete,
from the same locality cannot be attributed with certainty until further material
is available. The larva of langata closely resembles those of apicoargenteus, soleatus
and schwetzi but, in the material available, it can be separated from the first of
these by having the antennal seta at less than 0-7 x the distance from the base, to
the apex.

Aedes calceatus. The precise status of the various populations attributed to this
species is one of the most troublesome problems connected with the Ethiopian
Stegomyia at the present time. Much more material is needed from certain parts
of the range before the problem can be approached with any confidence. For the
moment the following brief notes on variation are all that can be offered. It should
be borne in mind that not only this species and soleatus but also heischi and the form
from Gwelo here tentatively attributed to langata may exhibit a pale posterior stripe
on the first mid-tarsal. From heischi and soleatus all the known forms of calceatus
can be separated by having an entirely dark fifth hind tarsal, and soleatus also differs
strikingly in its abdominal markings. From calceatus langata differs in its broader

12 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

abdominal bands and in the shape of the pale line at the base of the hind tibia.
In calceatus the Livingstone form differs from the type form in having the femoral
knee-spots on the hind legs shorter with the backward prolongation reduced or
absent, the pale ring at the base of the third hind tarsal segment only about -^ the
length of the segment or less (about ^ the length of the segment in the type form
from Tanganyika) and the second abdominal tergite all dark. A single male from
the Langata Forest in the British Museum, which has terminalia almost precisely
like those of the type form calceatus and is here tentatively attributed to that
species, also has the backward prolongation of the hind femoral knee-spot reduced
and the band at the base of the third hind tarsal only about \ the length of the
segment. In this specimen both the second and third abdominal tergites are dark
and the median pale stripe on the scutum is more strongly indicated than in the type
form, reaching forward as far as the front end of the anterolateral patches. In the
type form this stripe stops short at the posterior end of the anterolateral patches,
and is represented further forward by at most a few pale scales. In the Langata
form the general reduction of pale scaling is also shown in the rather narrower
anterolateral pale patches on the scutum, and in the fact that the second mid-tarsal
is dark all round on about the apical third (pale behind to the tip in the type form).
The pale scales at the base of the third mid-tarsal mentioned by Edwards (1941)
are present in only a few specimens in the type series. They are not shown by the
Langata or the Livingstone form. As already noted, pale scales occur in this
position as an aberration in a number of species of Ethiopian Stegomyia. Mrs.
Van Someren has kindly sent the following note on variation in Kenya Lowland
specimens from Ganda, Gede and Kwale at present tentatively attributed to
calceatus : " Fore tarsi with first segment wholly or only partly white behind and
second segment sometimes white to | behind. Mid-tarsi with the amount of black
on the first tarsal joint very variable ; second segment nearly all white with a
dark spot at tip anteriorly ; third segment nearly always dark but may have a
small patch of white scales at the base behind. Third hind tarsal segment with
the white basal band usually very narrow (Ganda and Gede), but it may extend to
(Kwale). Hind femur with the anterior surface white on the basal ^ then a small
black patch followed by a large silvery white patch at | ; rest black (Ganda and
Gede) or white on just over the basal \, rest black (Kwale). Third abdominal tergite
with a white band or spot (?)." A full discussion of the significance of this variation
must await the taxonomic paper in the present series.

Aedes soleatus. A description of the larva of this species has recently been
supplied by Mrs. E. C. C. Van Someren for inclusion in the new edition of vol. i of
Mosquitoes of the Ethiopian Region (Hopkins, 1952). This description was used
together with the paedotype and other specimens in the preparation of the key to
larvae in the first paper of the present series. Recent collections in Kenya have
shown the adult to be somewhat variable, and Mrs. Van Someren has kindly sent
the following note on variation in specimens from Gede and Taveta : " Mid-tarsi
1-4 usually pale behind and above (Gede) or first with only a white ring at the base
but sometimes also white behind to near tip, second white with variable amount
of black in front at tip, third nearly always black but may sometimes be white

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 13

behind basally (Taveta). Third hind tarsal segment with basal white ring to .
Tergite 2 usually and 3 sometimes without white dorsal bands. ' ' Occasional specimens
of the Taveta form with the first mid-tarsal segment largely dark behind would
run down, on the key to adults included in the first paper of this series, to the
apicoargenteus group, to which soleatus seems to be quite closely related. From the
members of this group it can be distinguished by the size and shape of the antero-
lateral scutal patches and by the less extensive pale patch on the posterior pro-
notum. Other characters which may perhaps prove less reliable are the smaller
knee-spots, and the fact that the median anterior pale spot on the scutum is composed
of narrow scales. In connection with the latter character it should be noted that
Marks (1951) has found that it may vary in the scutellaris group with the temperature
at which the early stages are reared. One specimen from Taveta kindly sent by
Dr. Lumsden has the basal pale patch on the sixth abdominal tergite little more
than half the depth of the segment. Such a specimen, if it possessed the normal
type of first tarsal segment, would run down to heischi. If with a first tarsal which
was largely dark behind, it would run down to couplet 30, but could not be taken
beyond this owing to the combination of pale-ringed fifth hind tarsal segment
and pale spot on the anterior surface of the mid-femur. Confusion between atypical
specimens of soleatus and heischi seems to be the biggest danger and there is, in
fact, some reason to suppose that it may have occurred in the past. The distal
expansion of the basal ventral stripe on the hind tibia is perhaps the best single
diagnostic character from heischi. In the material at present available the larva
of soleatus can be separated from that of apicoargenteus by having the antennal
seta at less than 07 x the distance from base to apex.

Aedes apicoargenteus ssp. denderensis. This form is of particular interest because
it occupies an intermediate position both morphologically and geographically
between the apicoargenteus and calceatus groups. It differs from all other forms
in both groups, except calceatus, in having the larval siphon entirely dark. The
adult shows no constant difference from those of the type form occurring in Uganda
and the central Kavirondo district of Kenya, but differs from West African speci-
mens, other than those from the Cameroons, in having smaller anterolateral scutal
patches and in the more frequent occurrence of pale scales on the lateral lobes of
the scutellum. The third hind tarsal is more narrowly banded than is usual in
other regions, except the Cameroons, and in one specimen the fifth hind tarsal is
almost entirely dark, a feature also known elsewhere at present only from the
Cameroons. It is clear that the distinctive features of denderensis which have so
far been noted, other than the pigmentation of the larval siphon, are merely those
of a highland form. The hind femur is pale behind on about the basal and is
more extensively pale than in any other form so far examined. Van Someren
(1946), however, has recorded specimens from both Kenya and Uganda in which
it is pale on up to a half. Despite the suggestion of Wolfs (1949) to the contrary,
the condition of the male terminalia falls within the range of variation exhibited by
the type form. The character of the larval siphon is considered to be an important
one and, as it appears to be geographically representative, there is felt to be some
justification for naming this form as a sub-species.

14 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

Aedes schwetzi. A description of the larva of this species has recently been
published in the second edition of vol. i of Mosquitoes of the Ethiopian Region, to
which it was contributed by Mr. G. G. Robinson. It closely resembles those of
other members of the group, but is separable from calceatus and denderensis by
having the distal part of the siphon markedly paler than the remainder. The
range of variation in the position of the antenna! seta overlaps that encountered
in apicoargenteus on the one hand and soleatus and langata on the other, since in
some larvae it is at more and in others at less than 07 x the distance from base to
apex. Only a partial separation is therefore possible. From adults of apico-
argenteus it is easily separated by the broad and conspicuous border of pale scales
round the pre-scutellar space, but as already noted under apicoargenteus (Mattingly,
1952) the characters of the male terminalia given by Edwards (1941) are not really
distinctive unless it be that the ninth tergite of schwetzi is a little more deeply
excavated. As far as is known the lateral lobes of the scutellum are always entirely
white scaled, and this should make separation from the majority of apicoargenteus
easy.

Aedes deboeri. This appears to be a member of the dendrophilus group. It
shows resemblances to dendrophilus on the one hand and to bambusae on the other.
The relationship to bambusae is discussed below under that species. The larva
shows no constant difference from that of dendrophilus but, whereas no larvae of
the latter so far seen have more than one detached pecten tooth beyond the siphonal
tuft, deboeri larvae often have two or three. In deboeri larvae the siphonal tuft
is normally bifid, rarely single, whereas Kenya and Uganda dendrophilus normally
have it single. Nigerian and Gold Coast larvae of dendrophilus, however, often
have it bifid. Adults of deboeri can be separated from those of dendrophilus by
the narrower anterolateral pale scutal patches and by having the second mid-tarsal
segment pale above on less than half, usually much less, as well as by the largely
dark fifth hind tarsal.

Aedes bambusae and " ssp. kenyae." Type form bambusae can be recognized
from deboeri by the fact that the anterior horn of the anterolateral pale scutal
patches is continued round the anterior edge of the scutum nearly to the median
anterior pale spot and by the great reduction of the knee-spots on the hind femora,
which are represented at most by one or two pale scales. The yellow colouring of
the pale scutal markings is not absolutely distinctive, as Kenya deboeri often have
these markings pale yellow, and in the single female from Marangu tentatively
attributed to this species they are deep yellow. Ssp. kenyae is intermediate between
the two forms. Its pale scutal markings vary from deep yellow to whitish and it
is thought that the colour may be influenced by climatic factors, though insufficient
evidence is as yet available to establish this. It has well-developed knee-spots
on both mid and hind femora and in this respect resembles deboeri. Taking into
account this fact, its distribution and the known variation in both forms, it seems
more reasonable to regard it as a subspecies of deboeri than of bambusae. More
detailed evidence with regard to distribution is, however, much to be desired.
From deboeri (and from bambusae} ssp. kenyae is readily distinguishable in the larval
stage by having the pecten spines arranged in discontinuous groups with 1-5 spines

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 15

in each. Adult differences from deboeri are less satisfactory, but all specimens at
present available are distinguished by the fact that the second mid-tarsal is pale
above almost to the tip, at least along the mid-line.

Aedes angustus. As already indicated, the larva figured for demeilloni by Hopkins
(1936, fig. 6ic) is in fact that of the present species. Dr. Haddow has collected
further larvae from the Chuya Forest which almost certainly belong to
angustus. Unfortunately, however, the only specimen from the original series
which is now available for examination is a single incomplete pelt. The Chuya
Forest larvae are separable from those of all other Ethiopian mainland Stegomyia,
except unilineatus, the South-west African larvae attributed to pseudonigeria and,
in some cases, amaltheus by the fact that the setae composing the ventral brush
are single. In addition to the differences from unilineatus given in the key (Mat-
tingly, 1952), they can be recognized from all larvae of unilineatus at present
available by the absence of stellate setae from the thorax. This character is,
however, so variable among the Ethiopian Stegomyia that it has to be employed
with very great caution.

Aedes ruwenzori. The breeding-places of this species are still undiscovered, but
Gillett (19516) has obtained eggs by means of a similar technique to that employed
for africanus (Gillet et al., 1950). Larvae were obtained from these eggs, and Mr.
Gillett kindly allowed me the use of his MS. in preparing the following brief account :
Head seta A with 5-8 branches. Thorax with stellate setae. Comb and pecten
spines much as in bambusae. Sub-ventral tuft of siphon with 2-4 branches, usually
bifid. Saddle hair with 3-7 branches. Upper caudal seta with 2-5 branches,
lower trifid. Setae of ventral brush with 2-4 branches. In the key to larvae
(Mattingly, 1952) this species would run down with bambusae, deboeri and dendro-
philus. Gillett notes that it can be recognized from bambusae by the presence of
stellate setae on the thorax. The unreliability of this character has, however, been
noted above, and a better character would perhaps be the large number of branches
in head seta A. All larvae of bambusae so far examined have this seta double, and
those of deboeri and dendrophilus have it double or at most, in rare cases, triple.
Partial differential characters are afforded by the numbers of branches in the sub-
ventral tuft of the siphon and in the saddle hair and caudal setae. From angustus
the larva should be easily distinguished by the branched setae of the ventral brush
as well as by the comb and pecten spines, which are very similar to those of bambusae.
The structure of the comb and pecten spines makes separation from africanus
both easy and certain and this is an advantage, since damaged adults from high
altitudes, such as that from Dendezi (Mattingly, 1952), may be difficult to assign.

Aedes mascarensis. That this species is closely related to aegypti is shown by
the fact that their male terminalia are virtually identical while their larvae can
only be separated with difficulty. Despite this fact Edwards (1932) placed aegypti
in his group A and mascarensis in his group B. In doing so he appears to have been
influenced by the fact that the mesonotal pattern of mascarensis is at first sight
strikingly different from that shown by other members of group A. In the present
author's opinion mascarensis should be transferred to group A together with chemul-
Poensis (see above). This would render both groups more or less homogeneous

i6 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

with respect to their male terminalia. Nor are the mesonotal markings of mascarensis
nearly so aberrant with respect to the rest of the group as the description given by
Edwards (1941) would suggest. It is true that the scales over much of the scutum
(at least on the anterior half) are silvery white, but this condition is approached
by pale specimens of " aegypti var. queenslandensis." What Edwards failed to
point out is that even in mascarensis the typical anterolateral patches, postero-
lateral lines and supra-alar patches of group A are clearly indicated by local aggre-
gations of broader scales. It is true that the median anterior spot and the median
longitudinal pale lines are indistinguishable, but these are normally composed of
narrow scales in aegypti and in pale specimens of queenslandensis they may be very
inconspicuous, while in the so-called " var. mosquito Robineau-Desvoidy " the
median lines are absent. A single female adult recently sent to the British Museum
by Monsieur Vinson from Moka, Mauritius, differs notably from mascarensis in
scutal ornamentation and in the relative widths of the tarsal bands. This seems
quite clearly to be a new species, although the possibility of hybridization between
aegypti and mascarensis ought to be investigated. The description follows.

A'edes (Stegomyia) vinsoni sp. n.

Adult $ : Proboscis wholly dark. Clypeus devoid of scales. Back of head
differs from that of mascarensis in being devoid of dark scales even in the lateral
areas. Posterior pronotum largely covered with broad pale scales. (The precise
condition in mascarensis cannot be ascertained from available specimens owing to
rubbing, but it certainly bears a number of large, broad, loosely attached pale
scales and is not entirely bare as stated by Edwards (1941)). Mesonotum with
anterolateral and supra-alar patches, and posterolateral lines clearly marked by
aggregations of relatively broad scales as in mascarensis, but differs from mascarensis
in having a well-marked median longitudinal stripe of broad scales, tapering
posteriorly, the scales on either side of this stripe very pale fawn rather than white.
Scales around the pre-scutellar bare space also broadened. The whole of the
mesonotum is covered with white or whitish scales. (The normal extent of pale
scaling in mascarensis is not clear. Edwards (1941) and MacGregor (1924, 1927)
are ambiguous in their descriptions and all specimens at present available are
rubbed. It is, however, clear from these specimens that, at least in some cases,
the posterior part of the mesonotum is quite extensively dark.) Scutellum apparently
with all lobes entirely pale scaled, (mascarensis has the usual small patch of dark
scales on the tip of the mid-lobe, although most of these are rubbed away in the
available specimens.) Dorsal surface of abdomen almost entirely pale scaled as
in some aberrant forms of aegypti (Summers-Connal, 1926, 1927 ; Drake-Brockman,
1911). (In mascarensis the tergites have broad, shallow, pale basal bands as in
typical aegypti.) Front femur very largely pale behind, pale in front and above
on about the basal three-fifths. A small but conspicuous white spot above at tip.
Tibia all dark. Tarsi all dark except the first two, which are narrowly banded at
base. The presence of a knee-spot on the front leg is unusual in the Ethiopian
Stegomyia. It is shown by aegypti and by some, but not all, topotypic calceatus

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 17

but is absent in the case of mascarensis, which is almost equally abnormal in having
no knee-spots on any of the legs. Mid-femur largely pale in front and behind but
with a dark stripe above reaching nearly to base. Tibia entirely dark. First tarsal
pale ringed on about the basal , second pale on about the basal ; remainder dark.
(mascarensis has no knee-spot and the tarsi are rather more narrowly ringed
while the femur as a whole is much darker, pale scaling being restricted to a narrow
line on the under surface.) Hind femur pale hi front nearly to tip, pale behind on
about the basal f , with a dark streak above tapering backwards for about the
basal f. Knee-spot well developed. Tibia entirely dark. First two tarsi each
pale on about the basal ^. Third and fourth tarsi pale on about the basal \ and
f respectively. Fifth tarsal entirely pale. (As already noted, mascarensis lacks
the femoral knee-spot. It also has the femur less extensively pale and the tarsi
more narrowly banded except the fourth, which is pale on about the basal f.) All
claws apparently simple as in Group C. (Front and middles claws toothed in
mascarensis as in group A.)

Adult $ and early stages unknown. Holotype $ in British Museum.

Aedes granti. This species was placed by Edwards (1932) in his group C. The
recent description of the male terminalia (Leeson & Theodor, 1948) shows him to
have been correct. All the claws of the female are simple as in group C and some
members of group B. The scutal markings are unlike those of most members of
the group in that there is a complete border of pale scales round the edges. In this
respect they recall paullusi, hakanssoni and scutoscriptus. Knight & Rozeboom
(1946) and Knight & Hurlbut (1949) have recently revised group C, splitting off
the albolineatus sub-group and raising it to the status of a full group. They recognize
three sub-groups typified by scutellaris, albopictus and mediopunctatus respectively,
and they place granti in the second of these. In this they are apparently governed
by the basal position of the median portion of the pale tergal bands, but on the
much more important character of the arrangement of pleural scales (probably
unknown to them since there is no description of the condition in granti in the
literature) this species should clearly go into the scutellaris sub-group. It has,
however, in common with hakanssoni and scutoscriptus (but not paullusi), an extra
stripe of pale scales between the dorsal border of the sternopleura and the lower
edge of the posterior pronotum, as in albopictus, and is thus distinguished from more
typical members of the group. It is possible that these three species should be
placed in a separate sub-group. They are all restricted to rather remote islands
(in the zoogeographical sense), hakanssoni and scutoscriptus being known only from
Truk and Ponape respectively. Further study of the whole group from this point
of view is, however, required, and the matter will be discussed more fully in the
next paper.

Aedes albopictus. As noted above, Knight & Hurlbut (1949) have placed this
in a separate sub-group from scutellaris. In the same sub-group they provisionally
placed unilineatus. Examination of the latter shows that it possesses all the charac-
ters adduced for the albopictus sub-group, but differs from albopictus in having rather
more extensive pleural scaling, and in particular in the presence of a small additional

ENTOM. Ill, I. 2

i8 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

patch of scales in the posteroventral corner of the posterior pronotum. In this
respect it agrees with Biro's specimen of Aedes cretinus from Amari, Crete (Edwards,
19216), which I have been able to examine through the kindness of Dr. Soos of the
Budapest Museum. Males of albopictus from the Seychelles, Madagascar and
Mauritius have been dissected and their terminalia compared with those of other
specimens from Southern India, Java, Celebes, the Philippines and Hong Kong.
In general they show a rather feeble development of the enlarged spines on the
basal lobe of the coxite, but a similar condition is found in other parts of the range
and it does not appear to be taxonomically significant. Only one topotypical
specimen (a male) is available for comparison. This specimen, like those from
Mauritius, shows well-developed white stripes on the undersides of the fore and mid-
tibiae and the first mid-tarsus. Similar stripes are shown in various degrees of
development by most of the specimens from the Seychelles but a few Seychelles
specimens and two from Madagascar (kindly lent by Prof. Peus of the Berlin
Museum) appear to be entirely dark. It seems that discrepancies in the literature
are due to the fact that this character, besides being sometimes difficult to observe,
is variable. Thus ChT Ho (1931), Martini (1931) and Bonne- Wepster & Brug
(1932) all describe the tibiae as entirely dark, despite the fact that white stripes of
varying degrees of distinctness can be observed in specimens in the British Museum
from Malaya (Perak), Sarawak, the Philippines, China (Hong Kong) and Japan.
In other respects the Ethiopian specimens agree well with the topotypical one from
Calcutta, differing mainly in having the tarsal bands, in general, slightly narrower.
This difference is shown best by the first mid-tarsal and the second hind tarsal, which
have the basal bands about to J and J to f the length of the segment respectively
in the Ethiopian specimens as compared to and f respectively in the Calcutta
specimen. All Ethiopian specimens, like the Calcutta specimen, appear to have
the hind tibia entirely dark.

DISTRIBUTION RECORDS

The following list resembles in all respects that given in the first paper of the
series. The same abbreviations are employed with one addition, which is as follows :
Terv. = Congo Museum, Tervuren. Doubtful records are again marked with an
asterisk and these, together with records believed to have been based on misidenti-
fications, are discussed separately below (p. 22).

Aedes amaltheus

N. RHODESIA. Livingstone (De Meillon & Lavoipierre, 1944), Balovale (Robinson,
1948). S. RHODESIA. Bindura*, Darwin*, Shamva* (as poweri, Leeson, 1931),
Ndanga* (B.M.). BECHUANALAND. Kasane (De Meillon, 19470).

Aedes pseudonigeria

ANGOLA. O Wambu (= Nova Lisboa, Theobald, 1910), Bailundo (as wellmani,
Theobald, 1910). Rare in Benguela area, common in western Bihe and eastern
and central Bailundo (Wellman, MS.). BECHUANALAND. Unnamed locality (as

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 19

poweri, Edwards, 1912), Botletle River (Edwards, 1941), Francistown (Muspratt,
in litt.}. S.W. AFRICA. Otjiwarongo (Edwards, 1924^), Kanovlei, Karabib area*,
Okahandja (Muspratt, in litt.}, Okokarara (B.M.).

Aedes chaussieri

BELGIAN CONGO. Sandoa (Edwards, 19236), Elisabethville (Schwetz, 19276),
Keyberg (Elisabethville area, B.M.), Kisanga Gallery Forest (Keyberg, Mattingly
& Lips, in press), Lubilash Valley (L.S.H.). N. RHODESIA. Unnamed locality (as
africanus, Edwards, 1912), Lake Young (Edwards, 1923 bis), Ndola (B.M.).

Aedes masseyi

BELGIAN CONGO. Ruwe (as poweri, Theobald, 1910), Elisabethville (Edwards,
1941), Kisanga Gallery Forest (Keyberg, near Elisabethville, Mattingly & Lips,
in press). N. RHODESIA. Unnamed locality (as poweri, Neave, 1912), Lake Young
(Edwards, 1923 6/s).

Aedes keniensis

KENYA. Nairobi (as sp. near poweri, V. G. L. Van Someren & De Boer, 1926,
as ? masseyi, Edwards, 1941, as keniensis, E. C. C. Van Someren, 1946 6*s), Langata
Forest (Garnham, 1949), Fort Hall, Kerugoya, Kiambu (E. C. C. Van Someren, in
litt.), Eldoret*, Meru* (as simpsoni, Symes, 1935). TANGANYIKA Njombe* (B.M.).

Aedes heischi

KENYA. Taveta (E. C. C. Van Someren, 1951), Gede (B.M.), Mombasa, Shimba
Hills (E. C. C. Van Someren, in litt.). TANGANYIKA. Dar-es-Salaam* (as pseudo-
nigeria, Haworth, 1924), Mombo*, Moshi* (as ? pseudonigeria, Edwards in McHardy,
1932). TRANSVAAL. Magoebaskloof* (as pseudonigeria, Ingram & De Meillon,
1929).

Aedes demeilloni

ZULULAND. Eshowe (as subargenteus, Bedford, 1928, as deboeri var. demeilloni,
Edwards, 1936). NATAL. Amanzimtoti, Impetyeni, Margate (B.M.), Stanger
Beach (Muspratt, in litt.), Dukuduku Forest, St. Lucia (Muspratt in litt.). CAPE
PROVINCE. Hole-in-the-Wall (as deboeri, De Meillon & Lavoipierre, 1944, fide
Muspratt), Port St. Johns (B.M.), Coffee Bay, Embotyi, Keimouth, Mazeppa Bay
(Muspratt, in litt.).

Aedes kivuensis
BELGIAN CONGO. Kibati (as subargenteus ssp. kivuensis, Edwards, 1941).

Aedes woodi

KENYA. Ganda, Kaloleni (B.M.). NYASALAND. Cholo (Edwards, 1922).
MOZAMBIQUE. Vilanculos (Pereira, 1946).

20 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

Aedes subargenteus

KENYA. Taveta (B.M.), Ganda (E. C. C. Van Someren, in litt.}. NYASALAND.
Fort Johnston (Edwards, 1925), Ndala Chikoa (Lamborn, in litt.}, Unnamed locality
(Lamborn, 1939), Maiwale (B.M.). ZULULAND. Train between Empangeni and
Gingindhlovu* (Ingram & De Meillon, 1927), Umfolosi (B.M.), Dukuduku Forest,
Emseleni, St. Lucia (Muspratt, in litt.). NATAL. Melville (Muspratt, in litt.).
CAPE PROVINCE. Embotyi (B.M.), Port St. Johns (Muspratt, in litt.).

Aedes strelitziae

ZULULAND. Dukuduku Forest, Richards Bay, St. Lucia (Muspratt, in litt.).
NATAL. Margate (Muspratt, 1950), Amanzimtoti (Muspratt, in litt.). CAPE PRO-
VINCE. Embotyi, Keimouth, Port St. Johns (Muspratt, in litt.).

Aedes poweri

NATAL. Unnamed locality (Theobald, 1905). CAPE PROVINCE. Groot Rivier
Mouth*, Kologha Forest* (Muspratt, in litt.).

Aedes contiguus

S. RHODESIA. Mashonaland (as africanus, Theobald, 1901), Salisbury (Edwards,
1936), Ndanga* (B.M.). TRANSVAAL. Onderstepoort, Roberts Heights (as poweri ,
Bedford, 1928), Letaba*, Leysdorp Road*, Rolle Siding*, Tzaneen* (Ingram &
De Meillon, 1929), Johannesburg (Edwards, 1936), Fontainbleau, Pretoria, Riet-
fontein, Witkoppen (B.M.), Pietersburg (Liv.).

Aedes langata

KENYA. Langata Forest (E. C. C. Van Someren, 19460), Nairobi (E. C. C. Van
Someren, in litt.). NYASALAND. Maiwale* (H.D.). S.RHODESIA. Salisbury* (as
contiguus, Edwards, 1941), Gwelo*, Ndanga (B.M.).

Aedes calceatus

KENYA. Langata Forest* (as langata, E. C. C. Van Someren, 19460), Ganda
(B.M.), Gede, Kwale (E. C. C. Van Someren, in litt.). TANGANYIKA. Lindi
(Edwards, 19246). N. RHODESIA. Livingstone (De Meillon & Lavoipierre, 1944).
S. RHODESIA. Shamva (Leeson, 1931), Bindura (Edwards, 1941), Darwin (L.S.H.),
Chindamora Reserve*, Ndanga* (Meeser, in litt.). ZULULAND. Ishongwe (Muspratt
in litt.).

Aedes soleatus

KENYA. Gede (Bailey, 1947, E. C. C. Van Someren, 1947), Taveta (Heisch, 1948).
TANGANYIKA. Dar-es-Salaam, Lindi (Edwards, 19246, Harris, 1942), points between
Tanga and Moshi (Harris, 1942). NYASALAND. Mlanje (B.M.). S. RHODESIA.

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 2 i

Shamva (Leeson, 1931), Bindura, Darwin (Edwards, 1941), Chindamora Reserve*,
Ndanga* (Meeser, in litt.}. ZULULAND. Dukuduku Forest (Muspratt in litt.}.

Aedes apicoargenteus ssp. denderensis

BELGIAN CONGO. Costermansville (Wolfs, 1949). RUANDA-URUNDI. Kisenyi*
as apicoargenteus, Seydel, ig2ga).

Aedes schwetzi

BELGIAN CONGO. Elisabethville (Edwards, 1926 ; Schwetz, 1927*1) . He Shashu
(Edwards, 1941), Kipushi (L.S.H.), Costermansville* (Wolfs, in litt.}, Panda (Terv.),
Lubumbashi River (B.M.). N. RHODESIA. Balovale (Robinson, 1948), Ndola
(Robinson, 1950), Lake Bangweulu district, Kasama (B.M.).

Aedes deboeri

KENYA. Nairobi (as poweri, V. G. L. Van Someren & De Boer, 1926, as deboeri,
Edwards, 1926, E. C. C. Van Someren, 19466), Langata Forest (Garnham, 1949),
Kiambu, Ngong (E. C. C. Van Someren, in litt.}. TANGANYIKA. Arusha*, Mombo*,
Moshi* (Harris, 1942), Marangu* (B.M.).

Aedes bambusae

BELGIAN CONGO. Kausi & Biega Mountains (Wolfs, in litt.), Kivu Highlands
at 3000 metres (B.M.). UGANDA. Saddle between Mt. Mgahinga and Mt. Sabinio
(Edwards, 1935), Kanaba (Edwards & Gibbins, 1939), Behungi, Muko (Edwards,
1941), Lugezi (B.M.), Chuya Forest (Haddow, in litt.).

Aedes bambusae ssp. kenyae

KENYA. Elgeyo Escarpment (as deboeri, Edwards, 1941), Kaimosi Forest, Kisii,
Kitale (E. C. C. Van Someren, 19460), Elburgon, Equator, Mau (E. C. C. Van
Someren in Garnham et al., 1946), Cheborget, Eldoret, Muhoroni, Taito (E. C. C.
Van Someren in litt.), Chagroi Forest (fide Muspratt).

Aedes angustus

BELGIAN CONGO. Kausi and Biega Mountains (Wolfs, in litt.). UGANDA. Saddle
between Mt. Mgahinga and Mt. Sabinio (Edwards, 1935), Muko (Edwards & Gibbins,
1939), Chuya Forest (Haddow, in litt.).

Aedes ruwenzori

UGANDA. Bunguha, Kabingo, Kakuka, Kizimba (Haddow & E. C. C. Van
Someren, 1950).

Aedes mascarensis

MAURITIUS. Pamplemousses*, Reduit (MacGregor, 1924), Corps de Garde,
Unnamed localities (B.M.).

22 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

Aedes vinsoni sp. n.

MAURITIUS. Moka (B.M.).

Aedes granti

SOKOTRA. Unnamed locality (Theobald, 1901), Mouri (Leeson & Theodor, 1948).

Aedes albopictus

FR. SOMALILAND. Boat off Djibouti* (as scutellaris Walker, Doreau, 1909).
AMIRANTE Is. Desroche I. (as scutellaris, Theobald, 1912 bis). SEYCHELLES Is.
Unnamed localities on Coetivy, Dennis and Mahe Islands, Anse aux Pins, Capucin,
Cascade Estate, Long Island, Morne Blanc, Porte Victoria, unnamed localities on
Praslin and Silhouette Islands (all as scutellaris, Theobald, 1912 bis), Porte Victoria
region and unnamed localities on Mahe Island (Harper, 1947). MAURITIUS. Un-
named localities (as Culex albopictus, De Grandpre & De Charmoy, 1900, as scutellaris,
Theobald, 1905 bis, as Aedes albopictus, Edwards, 1920). "Everywhere on the
island from sea-level to the highest altitude," Flat Island, Gabriel Island (MacGregor,
1927). MADAGASCAR. Ankasobe, Diego Suarez, Majunga (as lamberti Ventrillon,
Ventrillon, 1904), unnamed localities (as scutellaris, Legendre, 1918, as albopictus,
Edwards, 1941), Antananarivo (as lamberti, Enderlein, 1921). REUNION. St. Denis
(Edwards, 1920), Ste. Rose (B.M.).

DOUBTFUL RECORDS
Aedes amaltheus

S. RHODESIA. Ndanga. This record is based on one female only and therefore
requires confirmation. The records from Bindura, Darwin and Shamva (as poweri,
Leeson, 1931) can no longer be confirmed from specimens. The possibility that
they refer to amaltheus is discussed below under zoogeography.

Aedes psetidonigeria

S.W. AFRICA. Karabib area. This record is discussed above under Taxonomy.

Aedes keniensis

KENYA. Eldoret, Mem (as simpsoni, Symes, 1935). As noted in the previous
paper, records of simpsoni from above about 5000 ft. in Kenya are probably the
result of misidentification. The true identity of the species concerned can no longer
be confirmed from specimens, but Mrs. Van Someren has suggested (in lift.) that
they may have been keniensis, and this seems very probable. TANGANYIKA.
Njombe (B.M.). This record is based on two females only and therefore requires
confirmation. One specimen is, however, in perfect condition and seems typical.
Both have a number of pale scales below at the base of the hind tibia as mentioned
by Mrs. Van Someren (1946, bis) for some Kenya specimens, and one has two or

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 23

three white scales forming a very small white spot beyond half-way on the anterior
surface of the mid femur.

Aedes heischi

TANGANYIKA. Dar-es-Salaam (as pseudonigeria, Haworth, 1924). There is one
female specimen of Haworth's in the British Museum apparently belonging to this
species, but its identity requires confirmation. Mombo, Moshi (as ? pseudonigeria,
Edwards in McHardy, 1932). The identification of these specimens seems to have
given Edwards considerable trouble, and it is not clear whether he came to any
final decision about them. No specimens have been preserved. On the basis of
our present knowledge an attribution to heischi seems possible but specimens are
badly needed from this area. (See also under deboeri below.) TRANSVAAL. Magoe-
baskloof (as pseudonigeria, Ingram & De Meillon, 1929). This record is based on
a whole larva and a larval and a pupal pelt in the British Museum. The larvae
appear to be indistinguishable from those of heischi but adults are required for
confirmation, since it is possible that we may be dealing with an undescribed species
of the demeilloni group.

Aedes subargenteus

ZULULAND. Train between Empangeni and Gingindhlovu (Ingram & De Meillon,
1927). The British Museum has a specimen which is marked as taken in a caboose
at Umfolosi in March, 1927, and presented by Dr. Ingram. It is almost certainly
the specimen to which this record refers. Umfolosi does not, however, lie between
Empangeni and Gingindhlovu, but is a few miles up the line from Empangeni.

Aedes poweri

CAPE PROVINCE. Groot Rivier Mouth, Kologha Forest (Muspratt, in litt.}.
Specimens have not as yet been received for comparison with the type, but from
Mr. Muspratt 's description the assignation to poweri seems a reasonable one. It will,
however, be necessary to have males and early stages from Natal before the identity
of the Cape Province with the topotypic form can be fully established.

Aedes contiguus

TRANSVAAL. Letaba, Leysdorp Road, Rolle Siding, Tzaneen (as poweri, Ingram
& De Meillon, 1929). There is no evidence that, in reassigning Ingram and De
Meillon's material, Edwards (1941) saw specimens from anywhere but Johannesburg
and Pretoria. Records from below 3000 ft. are therefore still open to question,
and it is preferred to ignore them in considering the distribution of contiguus until
they can be confirmed. Dr. De Meillon kindly sent the remaining material (one
specimen each from Letaba and Tzaneen), but, as these specimens are reduced to
the thorax only, their identity cannot be confirmed. S. RHODESIA. Ndanga (B.M.).
This record is based on a unique female and therefore requires confirmation.

24 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION
Aedes langata

NYASALAND. Maiwale (H.D.). This record is based on one female only and
therefore requires confirmation. S. RHODESIA. Salisbury (as contiguus, Edwards,
1941). This record is based on two badly rubbed females and requires confirmation.
The record from Gwelo is discussed above under Taxonomy.

Aedes calceatus

KENYA. Langata Forest (as langata, E. C. C. Van Someren, 1946^). This
record is discussed above under " Taxonomy." S. RHODESIA. Chindamora
Reserve, Ndanga (Meeser, in litt.}. No specimens from either locality are available
for examination and these records must therefore be regarded as provisional.

Aedes soleatus

S. RHODESIA. Chindamora Reserve, Ndanga (Meeser, in litt.). No specimens
from either locality are available for examination and these records must therefore
be regarded as provisional.

Aedes apicoargenteus ssp. denderensis

RUANDA-URUNDI. Kisenyi (as apicoargenteus, Seydel, 19290). Monsieur Seydel
informs me that no specimens are now available, and there appears to be no means
of deciding whether the record should be attributed to the type form or the subspecies.

Aedes schwetzi

BELGIAN CONGO. He Shashu (Edwards, 1941), Costermansville (Wolfs, in litt.).
These records are discussed under Zoogeography.

Aedes deboeri

TANGANYIKA. Arusha, Mombo, Moshi (Harris, 1942). These records cannot
now be confirmed. They may perhaps have referred to heischi. Mr. Swaine kindly
sent some of Harris's specimens from the laboratory at Morogoro but they were
unfortunately destroyed in transit. The record from Marangu is discussed above
under " Taxonomy."

Aedes mascarensis

MAURITIUS. Pamplemousses (MacGregor, 1924). This record was based on a
casual observation of a mosquito in flight and is therefore open to question.

Aedes albopictus

FR. SOMALILAND. Boat off Djibouti (as scutellaris, Doreau, 1909). This record
was based on a casual observation and there is no means of confirming it.

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 25

RECORDS BASED ON MISIDENTIFICATIONS
A edes pseudonigeria

KENYA. Nairobi (V. G. L. Van Someren & De Boer, 1926). This record appears
to have been based on a larva which is now in the British Museum. It belongs to
a species of Aedimorphus. TANGANYIKA. Dar-es-Salaam (Haworth, 1924), Mombo,
Moshi (Edwards in McHardy, 1932). It is most improbable, on distributional
grounds, that these attributions can have been correct. The species concerned is
here provisionally taken to be heischi, but it might have been deboeri or soleatus.
Specimens from all these localities would be very welcome. TRANSVAAL. Magoe-
baskloof (Ingram & De Meillon, 1929). See above, in the section on Taxonomy,
under heischi. ZULULAND. Eshowe (Ingram & De Meillon, 1927). This record
was based on the material subsequently taken by Edwards (1936) as the type series
of demeilloni. In addition to demeilloni it contains one specimen of dendrophilus
(Mattingly, 1952).

A edes demeilloni

UGANDA. Mongiro (Smithburn & Haddow, 1946), Mamirimiri (Haddow et al.,
1947), KENYA. Kaimosi Forest (E. C. C. Van Someren, 1946). All these records
refer to dendrophilus (Mattingly, 1952). ZULULAND. Eshowe (Edwards, 1936).
This material contained one specimen of dendrophilus (Mattingly, 1952).

Aedes poweri

BELGIAN CONGO. Kabinda (Schwetz, 19276) = apicoargenteus, Ruwe (Theobald,
1910) = masseyi. KENYA. Unnamed locality (Neave, 1912) = sp. indet., Nairobi
(V. G. L. Van Someren & De Boer, 1926) = deboeri. N. RHODESIA. Unnamed
locality (Neave, 1912) = masseyi from Lake Young. TRANSVAAL. Roberts Heights
(Bedford, 1928) = contiguus. BECHUANALAND. Unnamed locality (Edwards, 1912)
= pseudonigeria from Botletle River.

Aedes scutellaris

All records from the Ethiopian Region (Doreau, 1909 ; Theobald, 1912 bis ;
Legendre, 1918) appear to refer to albopictus.

UNIDENTIFIED AND MISQUOTED LOCALITIES.

ANGOLA. O Wambu (Aedes pseudonigeria, Theobald, 1910). Apparently a
version of Huambo which as Mr. Exell of this Museum has kindly informed me, is
an old name for Nova Lisboa.

TRANSVAAL. Pietersburg, Rietfontein (Aedes contiguus, B.M. & Liv.). There
are several places with these names in the Transvaal. In the list of localities (p.
26) I have included what appear to be the largest places with the appropriate
names.

26 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

DISTRIBUTION OUTSIDE THE ETHIOPIAN REGION

Aedes albopictus. South Georgia* (probably cretinus or a nearly related species
(lindtropi Schingarew), see Stackelberg, 1937), Chagos Archipelago and throughout
Oriental Region, N.E. China to Manchurian border, Japan to 40 N., Ryuku Islands,
Formosa, Mariana Islands, Dutch New Guinea*, Serang*, Timor*, Hawaiian Islands.
Temporarily established in Port Darwin, where it apparently did not persist.
Records from Polynesia are incorrect. (Theobald, 1912 bis ; Rhoudkhadze, 1926 ;
Kumm, 19316 ; Barraud, 1931 ; Feng, 19380 ; Bohart & Ingram, 1946 ; Farner
et al., 1946 ; Brug & Bonne- Wepster, 1947 ; Chow, 1949, 1950).

Records marked with an asterisk are doubtful and require confirmation.

LIST OF LOCALITIES WITH TOPOGRAPHICAL DETAILS

The remarks made in connection with the list of localities published in the first
paper of this series (Mattingly, 1952) apply equally to the present one. Altitudes
are again given in feet, where possible to the nearest 100 ft., otherwise to the nearest
5ooft., and rainfalls as the mean annual total to the nearest 5 inches. Figures
enclosed in brackets have been read from maps or, in the case of some rainfalls, refer
to an adjacent station. Details of localities included in the previous list are not
repeated.

LIST OF LOCALITIES

Locality.

Ankasobe, Madagascar
Anse aux Pins, Seychelles .
Antananarivo, Madagascar
Arusha, Tanganyika .
Bailundo, Angola
Behungi, Uganda
Benguela, Angola
Botletle River, Bechuanaland
Bunguha, Uganda .
Capucin Point, Seychelles .
Cascade Estate, Seychelles
Chagroi Forest, Kenya
Cheborget, Kenya
Chindamora Reserve, S. Rho-
desia .....
Cholo, Nyasaland
Chuya Forest, Uganda
Coetivy I., Seychelles
Coffee Bay, Cape Prov.
Mt. Corps de Garde, Mauritius .
Darwin, S. Rhodesia .
Dennis I., Seychelles
Desroche I., Amirantes
Diego Suarez, Madagascar .
Djibouti, Fr. Somaliland .

Altitude
(4.5oo)

4,600
4,600

(5.500)
(8,000)

<5o

(3,000)
(6,000)
<5oo
1,000
(6,000)
(6,200)

(4,5oo)
3,000
(5-8,000)
<5oo
<5oo
2,400
3,100
<5oo

<5 00
<5oo

Latitude

18.20 S.

4.41 S.

18.58 S.

3-25 s.

(12.158.)

1.15 S.

12.358.

2O. 12 S.

(o. 43 N.)
(4.488.)
(4.41 S.)
(0.308.)
0.358.

(17-308.)
(16.058.)
(1.158.)
(7.168.)
31.588.
20.16 S.
16.40 S.

(3.488.)
(5.508.)

12.10 S.

n. 34 N.

Longitude

47.10 E.

55-32E.

47-30E.

36.45 E.
(i6.5oE.)

29.48 E.

13.26 E.

24.20 E.
(30.07 E.)
(55-33E.)
(55.29E.)
(35.ioE.)

35.ioE.

(31.20 E.)
(35-o 4 E.)
(29.45 E.)
(56.I6E.)

29.18 E.

57-27E.

31 .30 E.
(55-42E.)
(53-54E.)

49.20 E.

43.09 E.

Rainfall

(55)
(90)
(60)

45
(70)

(55)
(10)

15

(60)
(90)
(100)
(55)
(55)

(35)
60

(50)
(90)
(45)
(65)

25
(90)
(90)
(45)

(5)

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

27

LIST

Locality.

Dukuduku Forest, Zululand
Elburgon, Kenya
Elgeyo Escarpment, Kenya
Emseleni, Zululand .
Equator, Kenya
Flat I., Mauritius
Fontainbleau, Transvaal. See

Johannesburg.
Gabriel I., Mauritius
Ganda, Kenya ....
Gingindhlovu, Zululand
Groot Rivier Mouth, Cape Prov.
Hole-in-the-Wall, Cape Prov.
He Shashu, Belgian Congo .
Impetyeni Forest, Natal
Ishongwe, Zululand .
Johannesburg, Transvaal .
Kabingo, Uganda
Kakuka, Uganda
Kaloleni, Kenya
Kanaba, Uganda
Kanovlei, S.W. Africa
Kasama, N. Rhodesia
Kausi and Biega Mts., Belgian

Congo .....
Keimouth, Cape Province .
Kerugoya, Kenya
Keyberg, Belgian Congo. See

Elisabeth ville.
Kibati, Belgian Congo
Kipushi, Belgian Congo
Kizimba, Uganda
Kologha Forest, Cape Prov.
Kwale, Kenya ....
Long I., Seychelles .
Lubilash Valley, Belgian Congo .
Lubumbashi R., Belgian Congo .
Lugezi, Uganda
Magoebaskloof, Transvaal .
Mahe I., Seychelles .
Maiwale, Nyasaland .
Majunga, Madagascar
Marangu, Tanganyika
Mau, Kenya ....
Mazeppa Bay, Cape Prov. .
Melville, Natal ....
Mt. Mgahinga, Uganda
Mkanduli, Cape Province .
Mlanje, Nyasaland .
Moka, Mauritius
Mombo, Tanganyika .

OF LOCALITIES (cont.)

Altitude

Latitude

<5

28.24 S.

7,900

(o.iSS.)

8,000

i. oo N.

<5o

(27.258.)

9,000

oo.oo

<5

(19.528.)

<5oo

(19.538.)

<5

(3.138.)

<5

(29.01 S.)

<5oo

(33.588.)

<5

32.07 S.

4,800

(2.02 S.)

(4.5<>o) .

(30.40 S.)

(500)

27.25 S.

5.9oo

26.11 S.

6,500

0.12 N.

7,000

0.35N.

<5oo

3-058.

(7.5oo) .

1.238.

(4,000)

19.15 S.

4,400

10.10 S.

6,500

2 . 20 S.

<5oo

(32.41 S.)

(5,000) .

0.30 S.

6,500

1.348.

(4,000)

(11.478.)

(6,000)

0.39N.

(4-5,000) .

(32.308.)

(500) .

4.10 S.

<5

(4.378.)

(2,500) .

(9.35S.)

(4,000)

(11.308.)

7.5oo

1.25 S.

3.500 .

(23.528.)

(0-3,000)

4-45 S.

3,200

(14.258.)

<5

15.408.

4,600

3.198.

8,300

o.io S.

<5

(32.308.)

500

30.398.

(4-9,000) .

(1.238.)

(2,000)

31.468.

4,100

(16.07 S.)

1,100

20.13 8.

1,400

4.528.

Longitude
3 2.i8E

(35-5'E.)
35-ioE.

(32.30E.)
35-34 E.

(57-39E.)

(57.41 E.)
(40.03 E.)
(3I-38E.)
(25.03 E.)

29.15 E.
(28.55E.)
(29-35E.)

32.25 E.

28.04 E.

29.58E.

30.01 E.

39-38E.

29.58 E.

19.20 E.

3i.2 3 E.

28.40 E.

(28.22E.)

37.16 E.

29. 16 E.

(2 7 .I 5 E.)

30.04 E.
(27.20 E.)

39.32 E.
(55-3IE.)
(24.07 E.)
(27.20 E.)

29-34E.
(30.02 E.)

55-3E.
(35-I7E.)

46.20 E.

37-33 E.

35-40 E.
(28.35E.)

30.31 E.
(29.39 E.)

28. 43 E.
(35.47 E.)

57-30E.

38.14 E.

Rainfall
40
45
45
(40)
50
(40)

(40)
(45)

45
(40)
(45)
(55)
(50)
(30)

35
(45)
(65)
(30)
(55)
(20)

50

(60)
(40)
(50)

(60)

45
(55)

40

40

(90)
(55)
(50)
(60)

(35)
(85-150)

(25)
(60)

(50)

55
(50)

40
(60)

25
70

(70)
35

28

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

LIST
Locality.

Morne Blanc, Seychelles
Mouri, Sokotra

Muko, Uganda ....
Ndala Chikoa, Nyasaland .
Ngong, Kenya . < .
Njombe, Tanganyika
Okokarara, S.W. Africa
Otjiwarongo, S.W. Africa .
Pamplemousses, Mauritius
Panda, Belgian Congo = Jadot-

ville.

Porte Victoria, Seychelles .
Praslin I., Seychelles
Reduit, Mauritius
Richards Bay, Zululand
Rietfontein, Transvaal
Rolle Siding, Transvaal
Ruwe, Belgian Congo
Mt. Sabinio, Uganda
St. Denis, Reunion .
St. Lucia, Zululand .
Ste. Rose, Reunion .
Sandoa, Belgian Congo
Shimba Hills, Kenya
Silhouette I., Seychelles
Stanger Beach, Natal
Taito, Kenya .
Umfolosi, Zululand .
Vilanculos, Mozambique
Witkoppen, Transvaal. See

Johannesburg.

OF LOCALITIES (cont.).
Altitude Latitude

800
<5oo
7,500
(1,600)
6,400
6,000
(4,5oo)
4,800

<50o
(0-1,300)
(1,000)
<5oo
5,400
i, 600
(4,000)
(4-12,000)
<5o

<5 00

2,900

1,000

(0-2,500)

(6,500)

(4.398.)

(I2. 3 6N.)

(1.138.)

(14.10 S.)

1.188.

9.20 S.

20.35 S.

(20.27 S.)

(20.09 S.)

4-308.
(4.198.)

(20.12 S.)
28.388.

26.09 S.

(24.498.)

10.41 S.
i .25 S.

20.55 8.
(28.308.)
(21 .07 S.)

9.398.
4.138.
(4.288.)

(29.20 S.)

0.258.

28.25 S.
22.29 S.

Longitude
(55-27E.)
(53-59E.)
(29.50E.)
(35.05 E.)

36.40 E.

3 4 . 4 6E.

17.27 E.
(i6. 39 E.)
(57-35 E.)

55-29E.
(55.43 E.)
(57-33E.)

32.04 E.

28.11 E.
( 3 i.o 7 E.)

25-35E.

29.34 E.

55-30E.
(32.25 E.)
(55-50E.)

22.44 E.

39.26E.
(55.i 4 E.)

( 3 I,.20E.)

35-I5E.
32.i 4 E.
34-51 E.

Rainfall
(i oo)
(15)
(55)
(25)
35
40

(20)

(15)
(50)

85

(90-120)
(60)
45
(30)
(20)

45
(60)

>IOO

(35)

>IOO

(50)

(45)
(90-140)

(40)
(65)

45

25

BIONOMICS IN RELATION TO DISTRIBUTION

General considerations have already been discussed (Mattingly, 1952), and accor-
dingly all that is attempted here is a presentation of the available data.

BREEDING-PLACES.

The same classification into " Preferred " and " Occasional " is adopted as in
the previous paper.

Aedes amaltheus. Known only from tree-holes (De Meillon & Lavoipierre, 1944).
Aedes pseudonigeria. De Meillon's larvae from S.W. Africa were found in tree-holes.
Aedes chaussieri. Breeding-places unknown. Aedes masseyi. Breeding-places
unknown. Aedes keniensis. Known only from tree-holes (E. C. C. Van Someren,
1946 bis). Aedes heischi. Known only from tree-holes (E. C. C. Van Someren,
1951 ; Lumsden, in lift.). Aedes demeilloni. Preferred. Dracaena axils (Hopkins,
1936 ; Muspratt, in litt.). Occasional. Bamboos (Ingram & De Meillon, MS.
It seems possible, however, that this may have referred to dendrophilus) , banana

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 29

and Sirelitzia axils (Muspratt, in Hit.}. Aedes subargenteus. Known only from
tree-holes (Hopkins, 1936). Pondoland larvae, like the paedotype, were obtained
from gravid wild-caught females (Muspratt, in litt.}. Aedes kivuensis. Breeding-
places unknown. Aedes woodi. Breeding-places unknown. Aedes strelitziae.
Preferred. Axils of Strelitzia nicholai (Muspratt, 1950). Occasional. Axils of
cultivated banana (Muspratt, 1950), Dracaena axils (Muspratt, in litt.}. Aedes
poweri. Breeding-places unknown. Cape Province larvae have been obtained
from gravid females (Muspratt, in litt.}. Aedes contiguus. Preferred. Probably
tree-holes. Occasional. A rot-hole in a paw-paw tree. Pandanus axils, a snail
shell (Hopkins, 1936). Aedes langata. Known only from tree-holes (E. C. C. Van
Someren, 1946 ; Meeser, MS.). Aedes calceatus. Preferred. Tree-holes (Muspratt,
1945). Occasional. Crowns of coconut palms (Haworth, 1924. The findings of
Lester (1927) and Wiseman et al. (1939), however, appear to prove conclusively that
the record was due to introduction by a native collector), utensils (Muspratt, 1945),
bamboo pot in a tree (E. C. C. Van Someren, MS.). Aedes soleatus. Preferred.
probably tree-holes (Harris, 1942 ; Lumsden, in litt.}. Occasional. Crowns of
coconut palms (Haworth, 1924, but see above under calceatus}, bamboo stumps
(Harris, 1942), bamboo pots set up as traps (Bailey, 1947). Aedes apicoargenteus
ssp. denderensis. Known only from a tree-hole (Wolfs, 1949). Aedes schwetzi.
Preferred. Tree-holes and bamboo stumps (Robinson, in litt.}. Occasional. A tin,
a hole in cement, a tub (Schwetz, 19270). Aedes deboeri. Known only from tree-
holes (Harris, 1942 ; E. C. C. Van Someren, 1946 bis}. Aedes bambusae. Known
only from bored bamboos (Hopkins, 1936). Aedes bambusae ssp. kenyae. Tree-
holes, rock-holes, bamboo pots (Garnham et al., 1946).

Aedes angustus. Known only from bored bamboos (Hopkins, 1936). The
compressed thorax suggests that it is specially adapted to this type of habitat.
Aedes ruwenzori. Breeding-places unknown. Despite the compressed thorax all
attempts to find larvae in bamboos have failed. (Haddow & Van Someren, 1950).
Larvae have been obtained from wild-caught gravid females (Gillett, 19516). Aedes
mascarensis. Known only from tree-holes (MacGregor, 1924). Aedes vinsoni.
Breeding-places unknown. Aedes granti. Known only from wells (Leeson &
Theodor, 1948). Aedes albopictus. Common in tree-holes, cut bamboos, leaf axils
and coconut shells, rare in ground pools, rock-holes and utensils (Earner et al., 1946).
MacGregor (1927) differs in recording it as common in rock-holes in Mauritius,
but it should be noted that the term " rock-hole " may connote two very distinct
types of breeding-place (Mattingly, 1952). Chow (1950) infers that it is common
in artificial water containers in Formosa, and again it may be noted that the term
" utensils " may include a number of quite distinct ecological niches. Bick (1949)
records it as commoner in artificial containers than in tree-holes.

SEASONAL DISTRIBUTION

So little is known about the species dealt with in the present paper that almost
nothing can be added to the summary already given (Mattingly, 1952). For some
brief notes on Aedes bambusae kenyae see Garnham et al. (1946). For Aedes deboeri

3 o THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

see Garnham, 1949, and for Aedes calceatus and schwetzi see Robinson, 1950. Aedes
albopictus is of special interest because of its more northerly occurrence in the Far
East than in the case of Aedes aegypti or most other species of the sub-genus. The
most northerly record of albopictus appears to be from Pekin where it was found
in early September (Ch'i Ho, 1931). This record seems to be a very exceptional
one, and it is doubtful whether it indicates an indigenous population. Feng (1935),
however, records it as the commonest day-time-biting mosquito a little further
south in Shantung Province. Lamborn (1922) notes its almost complete absence
from Shanghai during a cold spell when the mean daily temperature was 61 F.
Many authors record it as most abundant during the rainy season, and Senior White
(1934) states that it occurs in Calcutta only at this time. The figures given by the
latter author are very small, but they seem to indicate that it makes its appearance
well after the beginning of the rains. The same author (1920) records it as abundant
throughout the year in Ceylon.

BITING-HABITS

As in the previous paper of this series the subject of biting-habits is taken, for
convenience, to include that of vertical distribution. The following species seem
likely to bite man at least on occasion.

Aedes amaltheus. A few specimens were taken biting on the forest floor at Kasane
(De Meillon, 1947^). Aedes pseudonigeria. Wellman, in an unpublished letter to
Austen, describes this as a " persistent and vicious biter." One specimen in the
British Museum, probably the one referred to by Theobald (1910), is marked as
taken out of doors in a village at 5 p.m. and as a common domestic mosquito biting
by day. Others are marked as taken at house or tent lamps at 8 p.m. Mr. Muspratt
informs me that this species was taken biting by De Meillon in S.W. Africa. Aedes
chaussieri. The single specimen from Ndola was taken biting at dusk (Robinson,
in Hit.}. It seems probable that some at least of the Congo specimens were also
taken biting. Aedes masseyi. It seems probable that adults in the British Museum
were taken biting. Aedes keniensis. In a long series of catches in the Langata
Forest Garnham (1949) took only one specimen biting. Aedes demeilloni. De
Meillon and Lavoipierre (1944) took a single specimen biting in forest at Mkanduli.
Aedes subargenteus. In an unpublished report very kindly sent by Dr. Lamborn
he records this species as biting very viciously in dense woodland at about n a.m.,
and again in the afternoon near Ndala Chikoa. Lumsden (in lift.) records it as
biting at Taveta both on the forest floor and in the canopy. Aedes kivuensis. The
only known specimen was presumably taken biting, but there is no definite infor-
mation on this point. Aedes woodi. The Mozambique specimen was taken biting
(Pereira, 1946), and Mrs. Van Someren states (in litt.) that specimens are easily
obtained in the bush at Ganda. There is no information concerning the type
specimen. Aedes strelitziae. This is noted by Muspratt (1950) as a more persistent
daytime biter than simpsoni. Aedes poweri. Muspratt states (in litt.) that he took
a number of females biting in the Kologha Forest. Aedes contiguus. Bedford
(1928) records specimens (as poweri) attempting to bite at Onderstepoort. Ingram
& De Meillon (1929) give larval records only. Aedes soleatus. Bailey (1947) took

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 31

adults on a number of occasions in forest. Lumsden (in litt.) has records of biting
on the ground in bush. Aedes deboeri. Garnham (1949) found this to be the prin-
cipal man-biting Aedes in the Langata Forest. Aedes bambusae. Edwards (1935)
records the capture of a few adults and Haddow states (in litt.} that he observed
biting in the Chuya Forest. Aedes bambusae kenyae. Garnham et al. (1946) took
" a few adults only " in the Kaimosi Forest. They also took some adults in houses
at Kisii. Aedes angustus. Haddow states (in litt.) that he took this as a tree-top
biter in the Chuya Forest. Edwards (1935) appears to have taken it only in the
larval stage but he is not very clear on this point. Aedes ruwenzori. Haddow
and Van Someren (1950) note this as a man-biting species markedly arboreal and
crepuscular in its habits. Aedes mascarensis. MacGregor (1927) states that " it
readily attacks man but is not very persistent in its attack, and it is easily frightened
off completely." Aedes vinsoni. The only known specimen was taken in a house.
Aedes granti. Described by Grant as being " very troublesome " (Forbes, 1903).
Aedes albopictus. There is general agreement among observers from various parts
of the range that this species prefers to bite by day in the shade. Night biting,
though not unknown, is comparatively rare. Although readily entering houses in
search of blood it is less domesticated than Aedes aegypti (Bonne- Wepster & Brug,
1932, and others) and a number of authors (e.g., Bohart & Ingram, 1946) record it
as particularly abundant in woodlands. In the Ethiopian Region Harper (1947)
notes that it attacks man readily in the Seychelles and MacGregor's account of its
behaviour is typical. This author states (1927) that in Mauritius it is a " voracious
and persistent biter in houses, while in woods and forests it frequently attacks in
swarms."

Aedes heischi, langata, calceatus, denderensis and schwetzi are not on record as
biting man.

The available data regarding the biting cycle have been included in the above
summary. Concerning vertical distribution relatively little is known. The only
species which have been recorded as definitely acrodendrophilic are deboeri (Garn-
ham, 1949), ruwenzori (Haddow & Van Someren, 1950), bambusae and angustus
(Haddow, in litt.). Lumsden states (in litt.) that subargenteus has been taken biting
in the canopy but does not indicate whether it is more abundant there than on the
ground. Forms which seem definitely to prefer the ground are bambusae kenyae
(Garnham et al., 1946) and soleatus (Bailey, 1947 ; Lumsden in litt.). With respect
to the vertical distribution of breeding-places we have only the observations of
Garnham et al. (1946), who obtained bambusae kenyae larvae from bamboo pots at
all heights up to 60 ft. (the greatest height investigated), and Bailey (1947), who
obtained soleatus larvae from ground level up to 36 ft. (the greatest height inves-
tigated).

ZOOGEOGRAPHY

Owing to the relative paucity of records this subject cannot be discussed in as
much detail as in the case of species occurring in the West African Sub-region. In
spite of this, however, the conjectural distributional areas of most species, as calcu-
lated empirically from such records as are available, seem reasonably consistent.

32 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

In comparing the two sub-regions the most striking difference is the very much
greater diversity of the East and South African fauna. Thus only one species
(pseudoafricanus) appears to be entirely confined to the West African Sub-region,
although two others, apicoargenteus and fraseri, have such a limited extension
outside it that they may reasonably be treated as West African. Even africanus,
with its vast extension in the Guinean forests, is not clearly West African in origin,
though it may be thought that the available evidence, such as it is, suggests an
original home in the Uganda savanna. Aedes dendrophilus, though now probably
more widespread in the West African savannas than in any other part of its range,
is clearly an ancient species with formerly a much wider extension, and there is
nothing to indicate in which part of its range it originated. The remaining West
African species are either so widespread as to belong to neither sub-region in par-
ticular (luteocephalus, vittatus), or mainly South and East African with minor incur-
sions into the West African savannas (metallicus, unilineatus) , or have been so widely
distributed by man that their natural distribution has been obscured (simpsoni,
possibly South African ; aegypti, possibly non-ethiopian). In short, the available
evidence, meagre though it is, would appear to suggest that the West African Stego-
myia fauna has been almost entirely derived from that of the surrounding highlands
and savannas. This view is also supported by the rather dubious evidence to be
derived from morphological resemblances between present-day species. As against
this the East and South African Sub-region possesses no less than 27 species, which,
as far as we know, are entirely confined within its limits in addition to such others
as may be presumed to be at present undiscovered.

Rhodesian Highland species. Chapin's Rhodesian Highland District has two
indigenous species, chaussieri and masseyi, the latter with a close relative, keniensis,
in the East African Highland District (Fig. i). The record of chaussieri from the
Lubilash valley suggests that it may perhaps occur in the Southern Congo Savanna
District, in which case it would qualify for inclusion among the West African species,
but this record is too imprecise for there to be any certainty on the point, and it
has seemed best to treat it in the present paper, since it is clear that, in any case, its
main area of distribution is almost certainly in the Rhodesian Highlands. Present
records would suggest that both species are confined to the north-eastern part of
the district, but it is probable that in fact both extend for a considerable distance
westwards into Angola, since, apart from the Bihe plateau, this part of Africa is
as yet entirely uncollected. The Bihe plateau is probably above the altitudinal
limits of either species, since neither is at present known from above 4,000 or at
most 4,500 ft. The southward extension of both species would seem to be very
limited to judge by the negative record from Balovale. It is not clear whether
the operative factor here is rainfall or altitude since the two are closely correlated
in this area. The occurrence of chaussieri below 3,000 ft. on the northern face of
the plateau, if this could be confirmed, would suggest that rainfall is the effective
limiting factor, and in this case both species would probably have rainfall limits
resembling those of africanus, Balovale, being a borderline locality. Keniensis
seems clearly to be a highland form of masseyi, which it very closely resembles.
The record from Fort Hall suggests that in the northern part of its range it may

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

33

occur below 4,500 ft., but this is a locality associated with very abrupt changes of
altitude, and in its present form the record is too imprecise to be of value. It is
also possible that masseyi may be precluded from the main distributional area of
keniensis by inadequate rainfall (Njombe, like Balovale, has an average rainfall of
40 in., with 6 dry months). The precise limits of these two species and of amaltheus
in Northern Rhodesia would be an interesting study, particularly if further light

t-0

FIG. i. Distribution of Rhodesian Highland species and of Aed.es keniensis.

could be thrown on their relationships by the discovery of early stages and males
of masseyi.

Among those Rhodesian species which apparently extend into adjacent fauna!
districts is Aedes schwetzi (Fig. 2). The only records of this species from outside
the district are, however, two from the Costermansville area, and these cannot be
finally accepted until specimens are available for examination. Wolfs states (in
litt.) that only one specimen has been taken in Costermansville itself, and it seems
possible that it may be more abundant at the slightly lower level of He Shashu.
The record from this island is due to Edwards, but the condition of his material is

ENTOM. in, i. 3

34

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

unknown. The relationship of schwetzi to apicoargenteus has already been discussed,
and it has been pointed out that the latter does not appear to occur above 3,500 ft.,
at most, along the northern edge of the Rhodesian plateau. In this part of its
range schwetzi is not known from below about 4,000 ft. although further south, at
Balovale, it apparently occurs at about 3,400 ft. It appears therefore that the 3,500-
ft. contour may be taken as a good approximate boundary between the two species.
It will be seen when the East African Highland and the East African Lowland
Districts are discussed that this is a very significant altitude over a large part of
Africa, bearing comparison with the 6,ooo-ft. contour on Ruwenzori and in the

FIG. 2. Distribution of Aedes schwetzi and allied forms.

Kavirondo and other montane areas (Mattingly, 1952). It is possible that it may
prove to be a critical altitude for apicoargenteus along the western face of the Uganda
plateau since, as already noted, the latter is associated with aberrant forms of this
species. It is not, however, absolutely preclusive here, as it appears to be further
south. As in the case of masseyi and chaussieri the rainfall limits of schwetzi are
difficult to assess, but there seems to be no reason to doubt that they are approxi-
mately the same as those of apicoargenteus. It is interesting to note that schwetzi
was found at Balovale at a time when africanus was apparently absent from there,
since there is also some indication that apicoargenteus may be slightly more drought-
resistant than africanus (Mattingly, 1952).

Together with Aedes schwetzi it is convenient to mention the closely related
apicoargenteus ssp. denderensis, although the latter is at present known with certainty
only from the Kivu highlands and is thus a purely East African Highland species.

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 35

Since it is known only from Costermansville little can be said here about its distri-
bution, but it seems reasonable to suppose that it is a highland form of apicoargenteus
restricted to the Kivu region and perhaps to neighbouring areas of comparable
altitude, e.g., in Ruanda-Urundi. The record from Kisenyi cannot be assigned
with certainty as between the type form and sub-species since the distribution of
the former in Tanganyika is uncertain, while the altitude (4,800 ft.) would probably
not preclude it at this latitude (see Mattingly, 1952, fig. 7). In view of the resem-
blance between the larvae of denderensis and calceatus, which suggests some affinity
with the East African Lowland fauna, even if a remote one, it would be particularly
interesting to know more about the distribution of the former. Unfortunately,
however, the Stegomyia fauna of Ruanda-Urundi is at present almost completely
unknown.

\ .Apparently restricted to the southern and western parts of the Rhodesian Highland
District are the very interesting species amaltheus and pseudonigeria, which also
extend into the south-east veld and south-west arid districts respectively (Figs. 3, 4).
In the southern part of its range pseudonigeria appears to be restricted to an area
having only 10 to 20 in. of rain in the year. It is rather surprising therefore that it
should also be known from the Bihe plateau, where the rainfall is much higher.
It seems virtually certain that it does not extend into the eastern part of the
Rhodesian Highland District, since it is a vicious man-biter and could hardly have
been missed in such well collected areas as Elisabethville and Ndola. It seems
reasonable to conclude that the Bihe form is a distinct sub-species, and that it is
probably restricted to altitudes of the order of 5,000 ft. and over (Fig. 3). Unfor-
tunately the available material is quite inadequate for a proper comparative study
of the two forms. It would seem that the S.W. African form is also a highland
mosquito, since it is not known from below about 3,000 ft.

Aedes amaltheus is at present known only from a very restricted range of altitudes
between about 3,000 and 4,000 ft. The lowest mean annual rainfall with which it is
associated is about 20 in. and the highest about 40 in. If these are in fact the distri-
butional limits then the population occurring in the more low-lying parts of Southern
Rhodesia would seem to be separated from that occupying the main part of the
range by the width of the Zambesi valley at Livingstone (Fig. 4). As in the case
of the southern boundary of masseyi and chaussieri it is not clear whether the
northern boundary of this species is to be equated with the 4O-in. isohyet or the
4,ooo-ft. altitudinal contour since the two run close together. This is unfortunate,
since the question of possible upper rainfall limits in Stegomyia is an interesting one.
It can only be hoped that the very scanty records from this area will before long
be supplemented. In the meantime there seems little reason to doubt that the
main distributional area of amaltheus covers a large part of southern Angola (not
shown in Fig. 4, which covers only the eastern part of the putative range), together
with the south-western part of Northern Rhodesia and parts of northern Bechuana-
land and Ovamboland. De Meillon & Lavoipierre (1944) express surprise that so
striking a species should previously have been overlooked, but in point of fact there
were no previous Stegomyia records at that time from any part of the main distri-
butional area as here defined. There were, however, some records from Bindura

36 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

and Shamva in Southern Rhodesia and it is supposed for this reason that the speci-
mens from these localities attributed to poweri (Leeson, 1931) may in fact have
been amaltheus. In view of the fact that proof of the presence of the latter in

\o

Anzxs with
more Tn&n
2.0* Ravn-

FIG. 3. Distribution of Aedes pseudonigeria.

Southern Rhodesia at present rests on only one female specimen this hypothesis
is highly conjectural.

Chapin's South-east Veld District contains at least two very distinct faunal
areas, the high veld of the Transvaal and Southern Rhodesia and the humid coastal

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

37

belt of Zululand, Natal and Pondoland. Both have characteristic species showing
interesting affinities with the fauna of the East African Highlands. A large part
of the area (Orange Free State, Basutoland and most of Eastern Cape Province) is
still unknown as far as its Stegomyia fauna is concerned. Information concerning
the extreme southern limits of such a species as contiguus would be very valuable
in assessing the climatic limits of the sub-genus as a whole an important matter
in any speculation regarding its past history.

over

*vooo'

Lo.rvd under
3000'

ftaln-
Areas w
more than <*<>'
Aai.iv

amaltKe-as.

FIG. 4. Putative distribution of Aedes amaltheus.

Aedes contiguus is the characteristic species of the High Veld and it is known from
nowhere else, but Aedes langata, which appears to be closely related to it and which
overlaps with it in Southern Rhodesia, was first described from the East African
Highlands (Fig. 5). In so far as confirmed records are concerned both would
appear to be highland species with altitudinal limits between 4,000 and 6,000 ft.,
but unconfirmed records suggest that langata at least may occur rather lower, and
3,500 ft. is perhaps nearer the limit for this species. There are also unconfirmed
records of contiguus from lower altitudes in the Transvaal, and further information
is needed before any very critical estimate can be made. It is, however, notable

38 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

that in general the 3,500 ft. contour appears to approximate to the " boundary "
between the East African Highland and East African Lowland faunas. Any
attempt to calculate empirical rainfall limits is also rather hazardous, but it is a

So

FIG. 5. Putative distribution of Aedes langata and Aedes contiguus. Only the unshaded
areas are within the rainfall and altitudinal limits so far recorded.

striking fact that neither species has been found anywhere with less than 20 in.
or more than 40 in. of rain (cp. amaltheus, above, and deboeri, Fig. 6). Applying
these as putative limits together with the altitudinal limits deduced above an
interesting discontinuous distribution is obtained embracing the Abyssinian and

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 39

East African Highlands and the High Veld. This distribution seems a reasonable
one in the light of negative records and of the known distribution of other species
of mosquitoes, and suggests the interesting possibility that these species may have
an upper rainfall limit, though why this should be so is not at all clear.

The relation between the faunas of the Abyssinian and Kenya Highlands is a
particularly interesting one, and it has therefore been thought desirable to include a
map showing rainfall in relation to altitude in this region (Fig. 6). This map is
based, wherever possible, on selected rainfall figures from the summaries published
by the East African Meteorological Service. Few or no records are, however, at
present available from the more northerly part of the area, and here use has been
made of the isohyets provided by the Rainfall Map of East Africa (E.A.F. No. 1518).
The gap between north-eastern Uganda and the area of suitable rainfall in southern
Abyssinia seems probably to be purely an altitudinal one and, if based on the
3,ooo-ft. contour, would be somewhat less extensive than it appears in Fig. 6, where
for convenience the 3,500 -ft contour, is used. Nevertheless it seems likely to be a
more permanent one than the gap between the Guinean Savanna Province and the
wetter parts of Abyssinia, which is purely a rainfall one (Fig. 14). Under exist-
ing rainfall conditions it seems that the topotypical population of langata is isolated
from areas of suitable rainfall and altitude in more northerly parts of Kenya, and
the absence of records from further north than Nairobi is perhaps not entirely fortuit-
ous. It seems likely that this species achieves its most extensive distribution in
Tanganyika.

Aedes poweri apparently requires a rainfall of at least 40 in. (38*3 in. in the case
of the Kologha Forest, fide Muspratt, but all rainfall figures in the present paper,
as in its predecessor, are expressed to the nearest 5 in.). It occurs, therefore, as
far as is known, in the wetter parts of South Africa from Natal to the Eastern Cape
Province, as do such species as demeilloni and strelitziae. It appears to require
very well distributed rainfall (2 months with less than i in. in the case of the Kologha
Forest and no months in the case of the Blaaukrans Forest (Groot Rivier)). Inten-
sive collecting by Mr. Muspratt has failed to reveal it in the coastal lowlands of
Natal or in certain upland forests, and it is still not known in what part of Natal
the type specimen was taken. It seems reasonable to suppose, however, that in
this part of its range it is a highland species, and that it may be regarded on the
basis of distribution as well as of morphology as the third and southernmost link
in the chain langata-continguus (cp. Figs. 5 and 8). It seems not at all unlikely
that this species and contiguus may overlap in some parts of their range, as do
contiguus and langata farther north.

Aedes subargenteus, which occurs widely in the coastal parts of Zululand, Natal
and Pondoland, is represented by a closely related species or subspecies (kivuensis)
in the Kivu Highlands (Fig. 7). It also occurs in the lower parts of Chapin's East
African Highland District at the southern end of Lake Nyasa and at Taveta. So
far as is known the type form does not occur above about 3,000 ft. To judge from
its distribution in South Africa it might be expected to require a rainfall of the
order of 40 in. or more. Its occurrence at Taveta would not be inconsistent with
this, since, as shown in the appendix to the previous paper (Mattingly, 1952), this

4

THE SUB, GENUS STEGOMYIA IN THE ETHIOPIAN REGION

place has a higher effective rainfall than the annual total of 26 in. would suggest.
Without local information it is impossible to explain the Fort Johnston records in

FIG. 6. Details of rainfall and altitude in the Abyssinian and East African Highlands.

similar terms, but it seems possible that here too some unusual local factors may
operate. Regarding kivuensis it seems probable that it is confined to some of the

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 41

Mfumbiro vulcanoes and perhaps to parts of Ruanda-Urundi. It is interesting to
note that Edwards and Gibbins failed to find it on Mgahinga or Sabinio.
Somewhat resembling subargenteus, though perhaps more nearly allied to simpsoni

Areas w UK JLO va.v
Or

FIG. 7. Distribution of Aedes subargenteus, kivuensis and woodi.

and strelitziae, is Aedes woodi, which is known only from three very scattered
localities in the South-eastern Veld, East African Lowland and East African High-
land Districts respectively. It is curious that this species does not extend as far
south as the coastal belt of Natal where other members of the group are so abundant,

42 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

but so little is known about it that speculation on this point would be useless. Of
the other species known from the South-eastern Veld District calceatus and soleatus
are discussed below in connection with the East African Lowland District. Aedes
strelitziae and Aedes demeilloni appear to be entirely confined to the area of Zululand,
Natal and Cape Province, having the very equitably distributed rainfall which has
been shown to be characteristic of dendrophilus (Mattingly, 1952). Aedes strelitziae

.10

30

FIG. 8. Distribution of Aedes poweri, demeilloni and strelitziae. Unshaded areas
agree approximately in rainfall and altitude with the area in which these species
have been found.

does not appear to have been found much above sea-level, but the record of demeil-
loni from the Impetyeni Forest suggests that this species may occur at considerable
altitudes. The record is too imprecise to give much idea of the altitudinal limits,
but the presumptive limit of 5,000 ft. shown in Fig. 8 is probably an extreme one,
and some of the areas shown as possibly suitable may well be in fact too high.

Among the East African Highland species keniensis, langata and kivuensis have
already been mentioned. So large a part of this district is still uncollected that
nothing more can usefully be said about their distribution. The distribution of

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 43

the deboeri group is of particular interest owing to the intergrading of deboeri itself
with the so-called bambusae ssp. kenyae of the East African Montane District. It
seems probable that the latter is in fact either a distinct species or else a subspecies
of deboeri. The striking yellow markings which have led to its being associated
with bambusae appear to be characteristic of species inhabiting the montane bamboo
forests, and are shown equally by angustus, bambusae s. str. and ruwenzori. Under
these circumstances a fuller knowledge of the distributional limits of deboeri and
of its variation in relation to altitude is much to be desired. At present the only
record from below 5,000 ft. which can be checked is that from Marangu, and the
records from Mombo and Moshi therefore appear doubtful, but more material is
badly needed from this part of Africa. The upper limit of deboeri, as far as is known,
is about 6,500 ft. The lower limit of kenyae is apparently about 5,000 ft. on the
western face of the Kenya highlands, but on the eastern face, where it might possibly
overlap with deboeri, it is not known from below about 8,000 ft. (Fig. 9). In this
connection it is interesting to note that at lower altitudes it is recorded as breeding
mainly in tree-holes and shaded rock-holes unlike bambusae s. str., which is known
only from bamboos. In this respect its habits are more in accordance with those of
deboeri. It may also be noted that the two forms are separated by a rather well-
marked rainfall factor, since kenyae, in so far as is known, is restricted to the wetter
western part of Kenya, while deboeri is known only from the drier eastern part (cp.
Figs. 6 and 9). The known rainfall limits of deboeri are 35 in. to 40 in. or 35 in. to
50 in. if the record from Marangu is included. The fact that, unlike keniensis, it has
not been recorded from the area of rather higher rainfall around Mt. Kenya may
suggest that its upper limit is about 40 in. and the Marangu form is a distinct species
or sub-species but much more evidence is required on this point. However, the
Rhodesian Highland affinities of keniensis and those of langata with the fauna of the
South-eastern Veld do seem to be reflected in their respective distributions in
Kenya, where the latter appears to have the same rainfall limits as deboeri. Mor-
phologically deboeri shows some resemblance to heischi, and to demeilloni so that it
might perhaps be regarded as the East African Highland representative of the
East African Lowland fauna. On the other hand, as already indicated, it clearly
has very close affinities with bambusae kenyae, and so might be regarded as a deriva-
tive of the East African montane fauna. At the present time there is too little
evidence to favour either hypothesis. Nor need they be mutually exclusive, and the
view that deboeri has been derived from the heischi group and kenyae, in turn, from
deboeri has much to commend it. Much more material is required, especially from
Tanganyika, before any hypothesis can be given very much substance.

Among the East African Montane forms other than kenyae, Aedes bambusae s.
str. and Aedes angustus are, as far as is known, confined to the mountains of the
Kivu, Mfumbiro and Kigezi regions, while Aedes ruwenzori is isolated further to
the North on the Ruwenzori range. The single record from the Kibati lava plain
suggests that kivuensis may be confined to certain of the Mfumbiro volcanoes or
it may occur in Ruanda-Urundi. These are all highly interesting species, and it
seems certain that others equally interesting await discovery on the many uncollected
mountains of Africa. The importance of montane species for the understanding

44

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

of the relationships of the Ethiopian fauna as a whole are such that any new evi-
dence will be most welcome ; for the present all that can be done is to place the
few available facts on record.

Among East African Lowland species heischi is of particular interest owing to
the diverse affinities suggested by its morphological characters, which recall deboeri,
from the East African Highlands, the calceatus group from the East African Lowlands
and demeilloni from the coastal part of the South-eastern Veld. In assessing its

FIG. 9. Distribution of East African highland and montane species.

rainfall requirements the record from Taveta requires to be approached with caution
for reasons given above, and it would seem, in general, to require at least 40 in.
In this respect it approximates most closely to demeilloni among the species men-
tioned, and it certainly resembles this species most closely on larval characters.
It seems therefore that it probably represents a South-eastern Veld element in the
East African Lowland fauna. The same is true of woodi, although in this case the
affinities are with simpsoni and strelitziae rather than with the demeilloni group.
The other East African Lowland species, calceatus and soleatus, appear to require
less rain, and both are known from localities with only 25 in. Both species have
been recorded from the South-eastern Veld and East African Highland Districts,

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

45

but it is in the East African Lowland District that they appear to attain their
widest distribution. Records from the Chindamora Reserve and from Ndanga
cannot at present be accepted, since no specimens are available from either locality.
The record of calceatus from the Langata Forest almost certainly concerns a distinct
species or subspecies, as explained above under " Taxonomy." Apart from these

O Sol eat as
A KeXsckt

FIG. 10. Distribution of the calceatus group.

the only records from above 3,500 ft. are those from Bindura (Edwards, 1941) and
of soleatus from Mlanje. No specimens from Bindura have been seen, and the
record from Mlanje is based on one female only. This specimen is in good condition
and seems quite typical but its identity requires confirmation. In any case neither
record need necessarily imply occurrence above 3,000 ft., since both localities are
associated with abrupt changes of altitude. It would seem therefore that both
species may be regarded as being limited largely, if not entirely, by the 3,500 ft.
contour (Fig. 10) ; certainly everything at present known about the East African

46 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

Highland and East African Lowland Stegomyia suggests that the two districts are
best denned by an altitudinal boundary of this order. The very interesting
records from Northern Zululand, sent by Mr. Muspratt while this paper was in the
press and not therefore included in Fig. 10, probably represent the southern limits
of the two species. In neither case do the rainfall limits appear to be at all
clearly denned except that 25 in., perhaps, represents a minimum.

There remain for discussion only the island species. Of these Aedes mascarensis
and Aedes vinsoni are apparently restricted to Mauritius. The relative distributions
of the two forms on the island cannot be discussed since the number of precise
locality records is negligible. Their resemblance to pale forms of aegypti is, however,
so striking and of so much interest that it has been thought desirable to show
their distribution in relation to that of such of these forms as occur in the Ethiopian
Region (Fig. n). The forms in question are referred to collectively by Edwards
(1941) as " var. queenslandensis Theo.," but it is preferred to treat them here as
var. queenslandensis Edwards nee Theobald, since there are many pale forms occur-
ring in various parts of the world, and we have as yet no evidence as to their origin
or genetical constitution. Under the circumstances it seems best that the name
queenslandensis Theo. should be restricted to the North Australian form on which
Theobald (1901) based his type description. Mathis (1934) has shown that the
laboratory bionomics of a number of strains from different parts of the world are
similar and there has been a small amount of other work on these lines, but there
would seem to be a case for a more thorough investigation from a genetical stand-
point. Few attempts to cross pale and dark forms appear to have been made,
but Dr. Mara has informed me in conversation that they hybridise readily in
Eritrea. The matter is of particular interest in the present connection because
there are in the British Museum some aegypti from Mauritius which are quite
markedly pale in colour. It seems possible that this may have some bearing on
the curious statement by MacGregor (1927) regarding the distribution of aegypti
on that island. According to MacGregor, " for some inexplicable reason it is
restricted to the coastal belt, in which, moreover, it has locally a very circum-
scribed distribution. In Rodriguez, however, the species is much commoner,
and occurs all round the island up to a level of at least 800 feet." A purely coastal
distribution of aegypti in Mauritius might conceivably be explained by the fact
that the local form was derived mainly from introduced " queenslandensis," since
the distribution of this form on the mainland is almost entirely coastal (Fig. n),
the few exceptions being localities which are in close and constant communication
with ports either by rail or waterway (e.g., Degema, Lokoja, and see Lewis, 1945).
To explain this coastal distribution in terms of altitude or rainfall is not very
easy. Although they seem normally to be restricted to localities at or near sea-
level, pale forms are on record from at least four localities at considerable altitudes
(Gebeit, Harrar, Mecca, Sinkat). These are, however, all in very hot parts of the
range. It is possible that they may represent temporary introductions and here,
even more than in most cases owing to the factor of human transportation, it is
necessary to bear in mind the possibility of seasonal extensions of range. Certainly
the temperature factor seems likeliest in the present case to limit distribution, and

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

47

it is interesting to note that Jepson et al. (1947) have expressed the opinion that
it is through the operation of this factor that Anopheles gambiae is largely excluded
from areas above 1,000 ft. in Mauritius. It would seem that the temperature
gradient on the island must be very steep. The rainfall picture is a com-
plicated one. This is a form showing greater powers of resistance to drought
than any other Stegomyia, not even excepting vittatus, and yet it is known not
only from places with moderately high rainfall (the palest of all the specimens

FIG. ii. Distribution of pale forms of Aedes aegypti and of Aedes mascarensis and vinsoni
in the Ethiopian Region. Inset : Map of Mauritius.

in the British Museum comes from Dar-es-Salaam), but even from those in which
the rainfall figure approaches or exceeds the three-figure mark (Bonny, Degema,
Old Calabar, Porte Victoria, Principe Island). Here the explanation seems almost
certain to be casual introduction from drier areas, and it seems reasonable to
assume that this is by nature a drought-adapted form from the Red Sea littoral
which has been introduced into coastal localities, often with high rainfall, where
it is capable of surviving without immediate reversion to the " typical " colour.
If this explanation is correct than it would seem that we have to deal with a com-
paratively well defined and stable genetical entity rather than with a number
of local aberrations of independent origin. It is not within the province of the
present paper to discuss the distribution of aegypti in non-Ethiopian Africa (for

4 8

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

which see Kumm, 19316, Callot, 1938, and Senevet, 1939), but it may be noted
that the reference to colouring made by Linnaeus (1762) in his type description
suggests that this was based on a pale form. Such forms certainly occur in Egypt,
and the British Museum has a series from Alexandria. Mr. Lewis states (in litt.)
that pale forms occurring inland in semi-desert areas in the extreme north of the
Sudan appear rather different from the coastal forms discussed above. On distri-

O 10 *O 6O 8O tOO 1X0 I*O 160

a

I6O 160 IfcO ti*O no

EZ3 Pistrtku.tton.oJ
area, of Gvop C

FIG. 12. a. Distribution of certain Stegomyia spp. in relation to Reinig's Glacial
Wooded Refuges, b. Distribution of Aedes granti and allied species in relation to
that of Group C as a whole. (Since this figure was prepared Aedes vittatus has been
found in Sardinia).

butional grounds it would not be at all unreasonable to suppose that the Mediter-
ranean and Red Sea populations are distinct. Details of the distribution of pale
forms of aegypti in the Ethiopian Region are given in Appendix III. Returning
to the position in Mauritius, it may be noted that if the conditions observed by
MacGregor still prevail, then mascarensis and vinsoni may well be largely isolated
from aegypti either by rainfall or by temperature barriers, since there are no records
of either species, susceptible of confirmation, from below about 1,000 ft. A local
survey of the relative distribution of the three forms would be of considerable
interest.

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

49

Concerning Aedes granti little can be said, since it is known only from a single
locality. The mesonotal coloration recalls a moderately pale Aedes aegypti, although
on structural characters it appears to be quite unrelated. The abdomen does not
show any pale speckling. The distribution of this species in relation to that of
its supposed nearest relatives has been discussed above under " Taxonomy." Its
distribution in relation to that of Group C as a whole is shown in Fig. 12.

The failure of Aedes albopictus ever to be taken on the mainland of Africa is one
of the mysteries of African zoogeography. It is true that over most of its range
this species is associated with very heavy rainfall, but in parts of India and in
China north of the Yangtse it must encounter rainfalls at least as low as those of
most of the East African coast (e.g., Delhi with 30 in. and 6 dry months) while in

FIG. 13. Recorded distribution of Aedes albopictus.

Mauritius it would seem to occur in coastal localities with less than 40 in. of rain.
Under these circumstances it might reasonably be expected at least from Zanzibar,
Pemba and Mafia Islands, which have more than 70 in. rain. In fact, however,
it seems never to have become established further west than the Seychelles and
Madagascar (Fig. 13). The only explanation which can be offered in terms of
rainfall is that in the Seychelles and western Madagascar the period of heavy rain
is from November to March, whereas in the islands mentioned it is from March to
May. Such considerations do not, however, suffice to explain its absence from the
wetter parts of the Mozambique coast. It would be interesting to know whether
it occurs in the Comoro Islands, and MacGregor's statement that it does not appear
to occur on Rodriguez seems to merit investigation. From rainfall considerations
the most likely areas for further spread are clearly Zanzibar and its adjacent islands
and the small area northwards from Tanga, which has the highest rainfall (about
55 in.) along the whole east coast.

ENTOM. Ill, I. 4

5 o THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

SUMMARY

The distribution of those species of Stegomyia found in the West African Sub-
region has been discussed in a previous paper (Mattingly, 1952). The present
paper deals with those species which have not so far been found in the West African
Sub-region and are believed to be confined to the East and South African Sub-
region. As in the first paper, zoogeography is discussed mainly in relation to rain-
fall and altitude. It is hoped that it may be possible to discuss other factors,
notably temperature and vegetation, in later papers, The present paper, although
it deals with many more species, is shorter than its predecessor because less is known
about the Stegomyia fauna of the East and South African Sub-region than about
that of the West African Sub-region. This applies to all aspects of mosquito
studies. Studies on taxonomy and relationships are hampered by the fact that
males and early stages of a number of species are still unknown, and by the
lack of representative series from more than a very few parts of the range. Very
large areas in the sub-region have still not been visited by collectors, and certain
of these, notably in Tanganyika and Nyasaland, are so situated geographically as
to prevent a proper co-ordination of the knowledge so far gained. There appears
to be an insufficient awareness on the part of collectors that the eggs of this group
are readily obtained by scraping out dry tree-holes and can be easily hatched in
the laboratory. They form, in fact, ideal collector's material, since they are almost
entirely immune from damage during transport. No large-scale studies on ecology
and ethology comparable to those made in Uganda and to a less extent in British
West Africa have been carried out anywhere in the sub-region. The very much
greater altitudinal diversity of the East and South African Sub-region appears to
have led to more extensive speciation than in the West African Sub-region, and here
the task of the taxonomist is rendered an especially difficult one, since there is at
present insufficient evidence to show how far the variation due to altitudinal factors
is at present discontinuous. All the known montane and island Stegomyia are at
present confined to the East and South African Sub-region, which is therefore of
major importance for the study of relationships and the reconstruction of the past
history of the group. Certain facts bearing on the relationship between the
Ethiopian Stegomyia fauna and that of the Palearctic Region are discussed, and
it is shown that Group A, which is at the present time entirely Ethiopian (with the
exception of Aedes aegypti), must at one time have extended into eastern Asia.
This subject will be treated more fully in a later paper, in which an attempt will
be made to relate the taxonomy of the Ethiopian Stegomyia to that of the sub-genus
as a whole.

The present paper includes a description of a new species from Mauritius even
more closely resembling a pale form of A edes aegypti than does mascarensis. Reasons
are given for transferring mascarensis from group B and placing it with the new
species in Group A. At the same time an appendix has been added containing
details of the distribution of pale forms of aegypti in the Ethiopian Region. Other
appendices are devoted to further notes on species discussed in the first paper and

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 51

to the rainfall of the very interesting region lying between Abyssinia and the
Guinean Savanna Province, which was also discussed in that paper.

Other points of interest are the occurrence in the Rhodesias of a recently dis-
covered species (Aedes amaltheus) which is annectent between Groups A and C,
the description, here published for the first time, of a larva from South West Africa
believed to be that of Aedes pseudonigeria, the discovery of Aedes langata (hitherto
known only from Kenya) in Southern Rhodesia, the discovery of Aedes subargenteus
(previously known only from Zululand, Natal and Pondoland and, as a supposed
subspecies, from the Kivu region) in the neighbourhood of Kilimanjaro, the dis-
covery of a highland member of the calceatus group probably a new species or sub-
species, near Nairobi, the discovery, on Kilimanjaro, of a curious yellow form of
deboeri which may eventually throw some light on the interesting problem of the
relationship of this species to " bambusae ssp. kenyae " and the discovery, for the
first time outside the Lagos area, of Aedes pseudoafricanus at Banana near the
mouth of the Congo. The male of Aedes woodi and the adults and larva of a new
subspecies of Aedes dendrophilus are described for the first time. The very recent
discovery of Aedes keniensis at Njombe in the Livingstone Mountains lends support
to the opinion, already formed from its resemblance to masseyi, that it represents
a Rhodesian element in the East African Highland fauna. An unassociated larva
is described which is believed to be that of masseyi. Despite the relative paucity
of records it seems reasonably clear that for the purposes of the present group the
East African Highland, Rhodesian Highland and East African Lowland Districts
may be closely defined by altitudinal boundaries of the order of 3,500 ft. The
East African Montane District seems to be similarly definable by a boundary in
the neighbourhood of 6,000 to 6,500 ft., but it is to be noted that these altitudinal
boundaries appear frequently to be associated with significant rainfall limits.

Short notes on taxonomy and bionomics in relation to distribution are included.
Topographical details and references to literature are confined to those which were
not included in the first paper.

APPENDIX I
FURTHER NOTES ON SPECIES OCCURRING IN THE WEST AFRICAN SUB-REGION

Since the publication of the first paper in this series a number of further distribu-
tion records have been obtained for species occurring in the West African Sub-region.
These are listed below. The abbreviations used are as follows : B.M., British
Museum (Natural History) ; K.I., Koninglijk Instituut voor den Tropen, Amster-
dam ; Terv., Musee du Congo Beige, Tervuren.

Aedes apicoargenteus

FR. CAMEROONS. Oyom-Abang (near Yaounde), Evodoula (B.M.). FR. EQUA-
TORIAL AFRICA. Brazzaville (Grjebine, 1950). BELGIAN CONGO. Kapanga, Matadi,
Mwene-Ditu, Rutshuru (Terv.), Popokabaka (B.M.). UGANDA. Kasunganyanja,
Namalu (Haddow, in litt.}, Ntotoro East, Tokwe (Lumsden, 1951), Ntaya Swamp

52 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

(Smithburn & Haddow, 1951), Nyagak Forest (Lumsden & Buxton, 1951). KENYA.
Kodera (Garnham & McMahon, 1947).

Aedes fraseri

FERNANDO Po. Boloko, Botonos, San Carlos, Santa Isabel (Gil Collado, 1936).
UGANDA. Ntotoro East (Lumsden, 1951).

Aedes dendrophilus

BELGIAN CONGO. Kimilolo River (Terv.), Kisanga River (Mattingly & Lips, in
press), Elisabethville (Muspratt, in litt.}. UGANDA. Ntotoro West (Lumsden,
1951). KENYA. Kwale (B.M.). N. RHODESIA. Serenje (B.M.). NATAL. Scott-
burgh (B.M.).

Aedes africanus

DAHOMEY. Ouidah (Huttel, 1950). FR. CAMEROONS. Oyom-Abang (near
Yaounde), Evodoula (B.M.). BELGIAN CONGO. Banzyville, Eala, Gombi-Masaka-
Kibanzi, Kabila, Kabukulu, Kakulubu, Kambundi, Kianga, Kibulu, Kimilolo
River, Kinkosi, Kisantu, Kitutu, La Kafubu, Leopoldville, Mangembo, Mubanga,
Mulassu-Tugi, Mwela, Popokabaka, Tukisi, Zundu (Terv.), Kasapa River, Kiniama,
Kisanga River (Keyberg), Lofoi River (Kundelungu Plateau), Luano (B.M.).
RUANDA-URUNDI. Usumbura (Terv.). UGANDA. Kaabong, Kasunganyanja, Lab-
wor (Haddow, in litt.}, Lunyo (Gillett et al., 1950), Ntotoro East, Ntotoro West,
Tokwe (Lumsden, 1951), Ntaya Swamp (Smithburn & Haddow, 1951), Nyagak
Forest (Lumsden & Buxton, 1951). ABYSSINIA. Jimma (Giaquinto-Mira, 1950).
KENYA. Kodera (Garnham & McMahon, 1947), Cheborget, Mambwa (E.C.C. Van
Someren, in litt.); TANGANYIKA. Ukara I. (B.M.). N. RHODESIA. Serenje
(B.M.).

Aedes pseudoafricanus
BELGIAN CONGO. Banana (as africanus Wanson, 1935).

Aedes simpsoni

FERNANDO Po. Biapa, Musola, Rebola (Gil Collado, 1936). BELGIAN CONGO.
Bili (De Meillon & Lavoipierre, 1944), Albertville, Inongo, Kazungeshi, Kibati,
Kimilolo River, Komi, unnamed locality between Libenge and Lisala, Ngbandi,
Thysville (Terv.), Kinanyira (Uvira area) (B.M.). RUANDA-URUNDI. Usumbura
(Terv.). UGANDA. Kaabong, Kasunganyanja (Haddow, in litt.}, Ntotoro East,
Ntotoro West, Tokwe (Lumsden, 1951), Koich-Kenya River Junction (Lumsden
& Buxton, 1951), Bageza (near Mubende), Bugazi (on Kome Island), Bunono,
Buwaya, Kitubulu, Seguku (all near Entebbe), Hakitengya (Gillett, I95i). ABYS-
SINIA. Jimma (Giaquinto-Mira, 1950). TANGANYIKA. Mofu (Gander, 1951).
NATAL. Port Shepstone, St. Winifred's (B.M.). CAPE PROVINCE. Port St.
Johns (B.M.), East London, Mazeppa Bay (Muspratt, in litt.}.

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 53

Aedes luteocephalus

BELGIAN CONGO. Banana (Wanson, 1935), Elisabeth ville (Mattingly & Lips,
in press). UGANDA. Namalu (Haddow, in litt.}, Mongiro (Lumsden, 1952).

Aedes unilineatus

ZULULAND. Ishongwe (Muspratt in litt.}.

Aedes metallicus

BELGIAN CONGO. Elisabeth ville (Mattingly & Lips, in press). KENYA. Kodera
(Garnham & McMahon, 1947). ZULULAND. Ishongwe (Muspratt in litt.}.

Aedes vittatus

DAHOMEY. Abomey (Huttel, 1950). FR. EQUATORIAL AFRICA. Brazzaville
(Grjebine, 1950). BELGIAN CONGO. Albertville, Banzyville, La Lufira, Lubum-
bashi River, Ngbandi, Sesenge-gadin (Terv.). UGANDA. Kaabong, Kamion
(Haddow, in litt.}, Ntotoro West (Lumsden, 1951). SUDAN. Between Suakin
and Erkowit (as sugens, King, 1908). ARABIA. Jebel Jihaf (B.M.). ABYSSINIA.
Moyale (La Face, 1939), Guder (Giaquinto-Mira, 1950). ERITREA. Barentu
(Giaquinto-Mira, 1950). IT. SOMALILAND. Burhacaba (Zavattari in Tedeschi &
Scalas, 1934). S. RHODESIA. Msonneddi (Edwards, 1940), Domboshawa (K.I.).

Aedes aegypti

CAPE PROVINCE. Additional southerly records kindly sent me by Mr. Muspratt
are East London and Port Alfred. He notes that this species has not so far been
found in Port Elizabeth or further westward. Dr. Mara has informed me in
conversation that he cannot accept my suggestion that his record of aegypti from
Mt. Bizen indicates a casual introduction. He tells me that it has since been
found there repeatedly and appears to be well established.

Records from outside the Ethiopian Region additional to those given in the first
paper are as follows :

Aedes unilineatus

INDIA and PAKISTAN. Karachi (Hicks & Diwan Chand, 1936), Nilgiri Hills
(Russell & Mohan, 1942), Kohat (Qutubuddin, in litt.}.

Aedes vittatus

BALEARIC Is. Raxa-Caubet (Mallorca) (Canamares, 1951). CORSICA. Bucalojo
River Gorge, San Nicolao, West Coast near Ajaccio (Aitken, in litt.}. SARDINIA.
Cantoniera Ovile Cannas, Picocca River Gorge, Villanovatulo (Aitken, in litt.}.
INDIA. Hyderabad City (Deccan) (Qutubuddin, 1951), Nedumangad, Kalkulam
(lyengar 1938).

54 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

The discovery of specimens of dendrophilus in the Tervuren Museum, where
they had been placed under Aedes fraseri, is of interest as this species has
only very recently been recorded for the first time from the Katanga (Mattingly,
1952). The specimens, unlike that on which the previous record was based, are
in good condition and seem typical, although males and early stages are still required
for final confirmation. In the appendix to my first paper it was suggested that the
occurrence of this species so far outside its known rainfall limits might be due to
the presence of gallery forest, but Monsieur Lips informs me that this forest and
others from which Aedes africanus is recorded above are of a special type known
locally as Muhulu, which is denser and more humid than ordinary gallery forest
and is associated mainly with the sources and head-waters of streams. There is
no doubt that it is an important distinction, and the resemblance to the type of forest
in which Aedes africanus was found at Taveta is striking (see appendix to first
paper). Synecological observations of this kind are felt to be of great value, and
it is considered that they may well have an important part to play in the further
development of mosquito research. Mr. Muspratt's record from Elisabethville
is based on a unique female in the S.A.I.M.R. collection. The records of dendro-
philus and africanus from Serenje are each based on a unique female specimen.
The rainfall here appears to have an even less equable seasonal distribution than
at Elisabethville since there are, on an average, 7 months with less than i in. of
rain. Serenje is, however, situated on a high plateau, and it seems possible that
forest of the Muhulu type may occur there. The record of dendrophilus from
Kwale is associated with one of the isolated areas shown in Fig. 9 of the previous
paper as having a type of rainfall suited to this species. It may well represent
a distinct subspecies. The taxonomic status of this and other isolated populations
will be discussed in a later paper in the series. As noted in the previous paper,
the record of africanus from Ukara Island was based on a single larva. It has now
been possible to confirm it from a series of ten female adults. The record of
unilineatus from Karachi is of special interest since it is associated with detailed
rainfall data. It is based on a mosquito survey, which may be taken for practical
purposes to have lasted from the beginning of May to the end of October, 1935.
Rainfall during this period was as follows : May o-o, June o-o, July 1-04 in.,
August 0-30 in., September 0-15 in., October 0*02 in. During the period in question
adults were found during August only. No larvae appear to have been found.
The mean annual rainfall over the previous six years was 10-2 in., with a mini-
mum of 0-69 in. in 1931 and a maximum of 20-82 in. in 1933. 7-17 in. fell in 1934.
The number of months with less than i in. of rain was 12 in 1931, 8 in 1933 and
10 in 1934. The record of this species from the Nilgiri Hills is interesting as
supporting the conclusion already drawn from Barraud's record from Bombay
that this species may occur in India in association with higher rainfalls than any
as yet recorded for it in Africa. Mr. D. J. Lewis has kindly told me that in his
opinion the altitude given for Erkowit in the first paper is too high and that it is
in fact of the order of 3,500 ft. Various altitudes appear in the literature. Hurst
& Black give 3,600 ft. and the East and South African Handbook gives 3,800 ft.
Certainly a figure of this order would be in better accordance with those recorded

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 55

for this species from elsewhere than the figure of 4,300 ft. which I have quoted
(see Mattingly, 1952, pp. 262 and 289). The records of vittatus from the Balearics
and from Corsica and Sardinia are of great interest since they fully confirm the
locality recorded for the type by Bigot, on which some doubt has been thrown.
The record from Gilder is interesting because of the high altitude involved (cp.
Mara's negative record from Mt. Bizen). " Chell's Third Camp " quoted in the first
paper as an unidentified locality for this species can now be said probably to have
been in the Marsabit area (see Edwards, 1941, p. 465). The record from Brazza-
ville is interesting in view of the paucity of records from this part of Africa noted
in the first paper. The record from Jebel Jihaf is associated with an altitude of
circa 7,100 ft. This is in good accord with its distribution elsewhere, in contrast
to Patton's negative record from behind Aden (see Mattingly, 1952, p. 291). The
record of pseudoafricanus from Banana is the first from outside the Lagos area.
It is based on a long series of specimens in the Congo Museum. This series contained
24 complete males, all of which were dissected and all of which proved to be pseudo-
africanus. The scutal markings of these and of the accompanying females agree
with those of specimens from the Lagos area and may be regarded as diagnostic
(see Chwatt, 1949, and Mattingly, 1952). In the Congo Museum the specimens
were placed under africanus, but it seems that this species probably does not occur
at Banana, which has a rainfall of only 32^ in. with 4 dry months (Vandenplas,
1943). The new records of africanus do not add materially to our knowledge of
its distribution except for that from Usumbura, which provides one of the rare
cases of its occurrence in an area with less than 40 in. of rain (35 in. but with only
3 dry months according to Vandenplas, 1943), and that from Kaabong, which
probably has a similar rainfall, although this is at present uncertain. It may be
noted that experimental evidence is now available to show that the eggs of this
species can withstand desiccation for at least 74 days (Gillett et al., 1950). The
record from Kaabong is of particular interest in relation to the known occurrence
of africanus in Abyssinia (see Mattingly, 1952). The record of apicoargenteus from
Kapanga is from the locality of that name in the Katanga and not from the one
in the Leopoldville area (per fide Basilewsky). It is of interest as representing the
highest altitude from which apicoargenteus has been recorded in this part of its range
(see Mattingly, 1952). The records of this species from Rutshuru and Usumbura
are based on one and two female specimens respectively. Larvae from these
localities would be very welcome for comparison with those of denderensis. The
relatively low rainfall in combination with very equitable seasonal distribution
at Usumbura has been noted in connection with Aedes africanus. Its significance
would be easier to assess if more recent detailed figures were available. Gillett 's
paper should be consulted for some further notes on the anomalies in the biting
behaviour of Aedes simpsoni discussed in the first paper of the present series. (See
also Bruce-Chwatt, 1950.) The record of this species from East London extends
its known distribution considerably to the southward.

The following are details of the localities listed above other than those already
tabulated in the first paper. Certain localities in the Belgian Congo could not be
placed with accuracy without making local inquiries, and as they affect only species

56 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

having a very wide distribution in this part of Africa it has been considered sufficient
to give a rough indication of their whereabouts. Gombi-Masaka-Kibanzi (Coll.
Henrard) and Kinkosi and Tukisi (Coll. Fain) have not been traced. There are
at least three places by the name of Bili in the Belgian Congo, and De Meillon gives
no indication as to which is implied (Coll. Liegeois).

<5o

y.oSN.

2.04 E.

(40)

2,500

5-568.

2Q.I3E.

50

<5

5-59S.

12.27 E.

30

[1,500) .

4-iyN.

21 . 12 E.

(70)

(2,000)

i5.o8N.

37-35 E.

(15)

[ 1,000)

(3-20N.) .

(8. 45 E.) .

<IOO

(1,000)

(3-25N.)

- (8. 45 E.) .

<IOO

(1,000)

(3-35 N.)

(8. 4 oE.) .

<IOO

(500)

(2. 45 N.)

. ( 44 .05E.) .

(25)

(1,700) .

(17.368.)

. ( 3 i.o8E.) .

(35)

<5

33-01 8.

. 2 7 . 5 8E. .

35

1,100

4.00 N.

ii . 10 E.

(6 5 )

(7,000) .

(9.00 N.)

- (37-45E.) .

(50)

(3,5oo) .

(0.43 N.)

. (30.o 4 E.) .

(55)

(3,000-

i.i 5 N.

29.47 E.

(50)

4.500)

(5.700) .

(7-40N.)

(36.50 E.) .

(55)

5,000

3-33N.

34.06E.

(35)

Locality Altitude Latitude Longitude

Abomey, Dahomey

Albertville, Belgian Congo .

Banana, Belgian Congo

Banzyville, Belgian Congo .

Barentu, Eritrea

Biapa, Fernando Po .

Bili, Belgian Congo. Not identified

(see above).

Boloko, Fernando Po .
Botonos, Fernando Po
Burhacaba, It. Somaliland .
Domboshawa, S. Rhodesia .
East London, Cape Prov.
Evodoula, Fr. Cameroons
Guder, Abyssinia
Hakitengya, Uganda .
Mt. Homa, Belgian Congo .

Jimma, Abyssinia
Kaabong, Uganda
Kabila, Belgian Congo. Bakali-

Inzia District.
Kabukulu, Belgian Congo. Bakali-

Inzia District.
Kakulubu, Belgian Congo. Wamba-

Bakali District.
Kambundi, Belgian Congo .
Kamion, Uganda
Kapanga, Belgian Congo
Kasapa R., Belgian Congo .
Kasunganyanja, Uganda
Kazungeshi, Belgian Congo .
Kianga, Belgian Congo. "Wamba-

Bakali District.
Kibulu, Belgian Congo. Kwango

Prov.

Kimilolo R., Belgian Congo . (4,000) . (11.438.) . (27.31 E.)

Kiniama, Belgian Congo . . (4,000) . (11.308.) . (28.23 E.)
Kitutu, Belgian Congo. Bakali-

Inzia District.

Kodera, Kenya . . . .. (4.500) . (0.308.) . (34.20 E.)
Koich-Kenya River Junction,

Uganda (2,500) . (3-34**.) . (3I.2QE.)

Komi, Belgian Congo . . . (1,500) . (3.208.) . (23.50 E.)

Rainfall

(3,000) .

(5.488.)

(I7.20E.)

(65)

(3,000) .

(3-45N.)

- (34-I5E.) .

(35)

(3.ooo) .

8.20 8.

22.36E.

(55)

(4,000)

(11.278.)

. (27.I9E.)

(50)

(3.500) .

(0.20 N.)

. (30.20E.) .

(50)

(4,800) .

(8.578.)

. (27.22E.)

(45)

(50)
(50)

(60)

(50)
(70)

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 57

Locality Altitude Latitude Longitude Rainfall

Labwor, Uganda . . .. (5,000) . (2.35 N.) . (33.50 E.) . (45)

La Kafubu, Belgian Congo =

Elisabeth ville.

La Lufira, Belgian Congo . . (4,500) . n.ooS. . 26.59 E. . (50)

Lofoi R., Belgian Congo . . (5,300) . (10.268.) . (27.53 E.) (5)

Luano, Belgian Congo . . (4,000) . (11.368.) . (27.36 E.) . (50)

Lunyo, Uganda. See Entebbe.

Mambwa, Kenya . . . (6,000) . 0.37 8. . 35.07 E. . (55)

Mangembo, Belgian Congo . . (1,500) . (4.348.) . (I4.I9E.) . (55)

Mobanga, Belgian Congo . . (3,000) . (5.538.) . (17.34 E.) (5)

Mofu, Tanganyika . . . (2,000) . (8.108.) . (36.ioE.) . (45)

Mulassu-Tugi, Belgian Congo.

Bakali-Inzia District.
Musola, Fernando Po. Not iden-
tified.

Mwela, Belgian Congo . . (3,500) . (6.028.) . (i7.4oE.) . (65)

Mwene-Ditu, Belgian Congo . (3,000) . (7.00 S.) . (23.25 E.) . (55)

Namalu, Belgian Congo . . (5,000) . (i.45N.) . (34.40 E.) . (50)

Ngbandi, Belgian Congo. Ubangi

District.

Ntaya Swamp, Uganda . . (3,500) . (0.41 N.) . (30.03 E.) . (55)

Ntotoro East and West, Uganda.

See Ntotoro Valley.

Nyagak Forest, Uganda . . (5,000) . (2.27 N.) . (30.57 E.) . (50)

Ouidah, Dahomey . . . <5OO . 6.2oN. . 2.05 E. . (40)

Port Alfred, Cape Prov. . . <5OO . 33-378. . 26.58 E. . (30)

Port Shepstone, Natal . . <5OO . (30.45 S.) . (30.27 E.) . (45)

Rebola, Fernando Po. Not identi-
fied.

Rutshuru, Belgian Congo . . 4,200 . 1.098. . 29.25 E. . 65

St. Winifreds, Natal . . . <5oo . (30.06 S.) . (30.51 E.) . (40)

San Carlos, Fernando Po . . <5OO . (3.25 N.) . (8.33 E.) . <ioo

Scottburgh, Natal . . . <5oo . 30.178. . 30.44 E. , (45)

Serenje, N. Rhodesia . . . 5,200 . 13.02 S. . 30.58 E. . 45

Sesenge-Gadin, Belgian Congo . (3,000) . (3.4oN.) . (29.40 E.) . (60)

Tokwe, Uganda .... (3,000) . (0.45 N.) . (30.02 E.) . (55)

Uvira, Belgian Congo . . . 2,600 . 3.248. . 29.08 E. . 35

Zundu, Belgian Congo. Bakali-
Inzia District.

APPENDIX II

RAINFALL OF THE BoR-PiBOR-ToRiT AREA

When plotting putative distributional limits of Aedes apicoargenteus in this area
it became apparent that the rainfall boundaries, based on empirical limits of 25 in.
with 3 dry months, 30 in. with 4 dry months or 45 in. with 5 dry months, instead
of following the isohyets and so denning the Guinean and Abyssinian areas of high
precipitation as isolated blocks tended to turn towards one another, while the dis-
tribution of patches of " forest " as shown by maps suggested that there might exist a
connecting bridge in the form of an area of exceptionally well distributed rainfall
(Mattingly, 1952). Such a bridge would be of considerable interest to zoogeo-
graphers as helping to explain the occurrence of a well-marked Guinean element

58 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

in the Abyssinian fauna. Thanks to Mr. D. J. Lewis it has been possible to obtain
additional rainfall figures for this area, and it has therefore seemed worth while to
prepare a detailed rainfall map (Fig. 14). It will be seen that purely on the basis

FIG. 14. Rainfall of the south-eastern Sudan and adjacent parts of Abyssinia,
Kenya, Uganda and the Belgian Congo.

of rainfall figures, two interpretations are still possible (Fig. 15), and the evidence
for the existence of such a bridge is therefore still largely that afforded by the
distribution of " forest " patches on the map. It seems clear, however, that such
a bridge must in any event have existed in comparatively recent times even if we
assume only very small long-term fluctuations in rainfall and that, subject to
edaphic factors, it would provide a forested connecting path between the two

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

FIG. 15. Alternative interpretations of the rainfall data illustrated in Fig. 14. Of
two a seems the more probable on account of the distribution of forest.

the

regions. The rainfall figures provided by Mr. Lewis with the assistance of the
Sudan Government Meteorologist represent mean annual rainfalls up to 1950.
As they have not previously been published they are given below :

Locality

Akobo

Boma

Bor.

Gilo

Isoke

Kapoeta .

Katire

Kongor .

Lafon

Lerua

Loa

Loelli

Nagishot .

Nimule .

Okaru

Palataka .

Pibor Post

Torit

Latitude Longitude

7-47
6.10

6. 12

See

33-01
34-29
31-33
Katire

4.16
4.46

32.02
33-35

4-03

32.47

7.09

5-02

31.22
32.28

4.00
3-47

;s-i8)

4. 16

32.34
31-58
(34-45)
33-34

3-37

- 32-03

4.29
4.02
6.48

32.09
32.26
33-08

4.24

32-33

Rain

(mms.)

972

1317

860

2263

777
1530
1009

716

1068

603

1017

1189

I2II

1395
880
988

Rain

Months

(Ins. to

with less

nearest

than

(5 in.)

i in.

40

4

55

2

35

4

90

2

55

3

30

3

60

3

40

5

3

5

60

3

45

4

25

3

40

3

50

2

50

4

55

2

35

3

40

3

Years

of
Average

3

8

30
8
6

13
ii

9

3

10
6
6

30
8

7

30
28

Other rainfall figures shown in Fig. 14 are taken from Hurst & Black (1943), and
from the summaries of rainfall for the year 1948 (including normals to date) pub-
lished by the East African Meteorological Department.

60 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

APPENDIX III

DISTRIBUTION OF PALE FORMS OF Aedes aegypti IN THE ETHIOPIAN REGION

NIGERIA. Bonny, Old Calabar (Theobald, ignc), Lagos (Summers-Connal,
1926, 1927), Lokoja (Simpson, 1913), Degemma (B.M.).

PRINCIPE I. Unnamed locality (B.M.).

SUDAN. Port Sudan, Suakin, Tokar (Edwards, 1941), Aqiq, Dongonab, Gebeit,
Halaib, Kassala, Mohammed Gul, Sinkat (Lewis, 1945).

ARABIA. Aden (Edwards, 1941), Kameran I., Mecca (Lewis, 1945), Amd, Tarim
(B.M.).

ABYSSINIA. Harrar (Giaquinto-Mira, 1950).

ERITREA. Archiko, Massawa (Lewis, 1943). Mara (in Jannone et aL, 1946)
notes that inland the species is represented by the " typical " (dark) form whereas
in coastal areas this is replaced by the pale form.

BR. SOMALILAND. Zeila (Edwards, 1941). G. R. C. Van Someren (1943) records
both forms and notes the occurrence of intergrading but does not give details of
their respective distributions.

KENYA. Mombasa (B.M.).

TANGANYIKA. Dar-es-Salaam (Edwards, 1941).

ZANZIBAR. Unnamed localities (B.M.).

ZULULAND. Ntambanana (B.M.).

NATAL. Durban (B.M.).

SEYCHELLES. Unnamed localities (Theobald, 1911), Porte Victoria (B.M.).
Harper (1947) particularly notes the absence of aegypti from all localities other than
Porte Victoria, which he visited.

MAURITIUS. Port Louis (B.M.).

ALDABARA. Picard I. (B.M.).

Details of localities not already listed are given below :

Locality Altitude Latitude Longitude Rainfall

Amd, Eastern Aden Prot. . . 3,100 . is.soN. . 48.ioE. . ? (10)

Aqiq, Sudan .... <soo . 18.14!?. . 38.nE. . 5

Archiko, Eritrea .... <5oo . (15.32 N.) . (39.25 E.) . (10)

Degema, Nigeria .... <5oo . 4.45 N. . 6.48 E. . (90)

Dongonab, Sudan . . . <5<x> . 2i.o6N. . 37.05 E. . <5

Gebeit, Sudan .... 2,500 . 18.57 N. 36.51 E - 5

Halaib, Sudan .... <5oo . 22.I2N. . 36.36E. . <5

Kameran I., Arabia . . . <5oo . 15.25]^. . 42.35 E. . (5)

Massawa, Eritrea . . . <5oo . 15.37]^. 39-29 E - . 5

Mecca, Arabia .... 2,000 . 2I.25N. . 39.54 E. . (5)

Mohammed Gul, Sudan . . <5oo . (20.45 N.) . (37.ooE.) . <5

Picard I., Aldabara . . . <5oo . (9.218.) . (46.05 E.) . ? (90)

Port Louis, Mauritius . . . <5<x> . 20.188. . 57.31 E. . (35)

Port Sudan, Sudan . . . <5<x> . 19.37 N. 37- I 3 E - 5

Principe I., G. of Guinea . . 0-3,000 .. 2.ooN. . 7-3oE. . 100

Sinkat, Sudan . . . . 3,000 . iS.soN. . 36.50 E. . 5

Suakin, Sudan . . -. . <5<>o . 19.07 N. . 37.20 E. . 5

Tarim, E. Aden Prot. . * 2,200 . 16.02 N. . 49.00 E. . (25)

Tokar, Sudan . . . ' . <5<x> . 18.25 N. 37-45 E - 5

Zeila, Br. Somaliland . . . <5oo . n.2oN. . 43.28 E. . (5)

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 61

APPENDIX IV

CORRIGENDA TO PART I

The receipt of further material of Aedes keniensis and the publication of a descrip-
tion of the larva of Aedes ruwenzori (Gillett, 19516) and of the adult of Aedes vinsoni
(supra) necessitate certain alterations to the keys published in the first paper of
the series. Further alterations are also required to cover variation in soleatus as
revealed by Mrs. Van Someren's studies of Kenya Lowland material, but it is not
felt that information concerning variation in calceatus is as yet sufficient to warrant
altering the keys. It should, however, be borne in mind that considerable difficulty
may be experienced with atypical specimens of this species, and the possibility of
confusion of an occasional aberrant specimen of soleatus with heischi should also
be noted. Alterations to the keys can conveniently be made as follows :

p. 246. First line of couplet 20. For " all round " read " above." For " ith "
read " ^th." Couplet 23 should be rewritten as follows :

23. Anterolateral scutal patches pointed behind and reaching margin of

scutum for some distance in front. First mid-tarsal often extensively

pale behind. Scale patch on ppn very small soleatus Edwards.

Anterolateral patches rounded. First mid-tarsal without posterior exten-
sion of pale scaling (may be pale above for up to f ths). Scale patch on

ppn extensive 24.

p. 247. After couplet 36 insert :

37. All femora with well marked knee-spots. Mid femur largely pale in front

vinsoni Mattingly.
Femora without knee-spots. Mid-femur with pale scaling restricted to

a narrow line on the under surface mascarensis MacGregor.

p. 249. Second half of couplet 20. For " bambusae, dendrophilus , deboeri " read
"24." After couplet 23 insert

24. Head seta A with 5-8 branches ruwenzori.

This seta with at most 3 branches, usually less

bambusae, deboeri, dendrophilus.

Corrections to the list of localities on pp. 262-263, suggested to me by persons
with local knowledge, are as follows :

Altitude of Erkowit to read 3600 ft. Altitude of Fort Ternan to read 5800 ft.
Latitude and longitude of Kayembe-Mukulu to read (9.01 S.) (23.59 E.). Altitude
of this locality to read (3300). In the second line of the summary on p. 293 the
words " No species " should read " Only one species." Mr. Muspratt informs me
that De Meillon and Lavoipierre's record of simpsoni from Cape Province should
be interpreted as from Hole-in-the-Wall rather than from Mkanduli. The last
entry under this species on p. 253 should be amended accordingly.

62 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

ACKNOWLEDGMENTS

In addition to those persons and institutions already listed in the first paper of
this series, many of whom have contributed also to the second, I am indebted to
Monsieur P. Basilewsky and the Director of the Congo Museum for the many
facilities afforded me there ; to Dr. J. Bonne- Wepster for showing me the collection
of African Culicidae at the Indisch Instituut, Amsterdam ; to Dr. T. H. G. Aitken
of the Rockefeller Foundation for allowing me to publish his very interesting records
of Aedes vittatus ; to Dr. Giaquinto-Mira of the World Health Organization, Teheran,
Persia, for topographical data from Abyssinia ; to Dr. De Barros Machado of the
Museu do Dundo, Lunda, Angola for topographical data ; to Dr. Elizabeth Marks
of the University of Queensland for the use of a manuscript copy of her thesis ;
to Dr. W. Peters of the Colonial Development Corporation, Njombe, Tanganyika,
who has sent me much material new to that territory ; to Dr. A. Soos of the Hun-
garian National Museum for the loan of a specimen of Aedes cretinus ; to Dr. B. P.
Uvarov of the Anti-Locust Research Centre, London, for allowing me to consult
his manuscript while his paper was in the press ; to Monsieur Vinson of the Govern-
ment Service, Mauritius for the very interesting new species of Stegomyia from that
island ; to Mr. Qutubuddin of the Malaria Institute of Pakistan for the record of
unilineatus from Kohat, to Dr. N. L. Corkill, A.D.M.S., Mukalla and Dr. L.
Merucci of Tarim for Arabian aegypti, and to Dr. L. Mara of the World Health
Organization for information regarding Eritrean aegypti.

REFERENCES

BARRAUD, P. J. 1931. Notes on some Indian mosquitoes of the subgenus Stegomyia, with

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BASCHKAREVA, A. D. 1931. Some facts on malaria and mosquitoes in the Sotschi district.

Mag. Parasit., Moscow, 2 : 55-58.
BICK, G. H. 1949. Notes on mosquitoes from Leyte, Philippine Islands (Diptera, Culicidae).

Nat. Hist. Misc. Chicago, No. 41.
BONNE-WEPSTER, J., & BRUG, S. L. 1932. The subgenus Stegomyia in Netherland India.

Geneesk. Tijdschr. Ned.-Ind., Bijbl. ii : 39-119.
BRUCE-CHWATT, L. J. 1950. Recent studies on insect vectors of yellow fever and malaria

in British West Africa. /. trop. Med. Hyg. 53 : 71-79.
CALLOT, J. 1938. Contribution a l'6tude des moustiques de Tunisie et en particulier du

Sud de la R6gence. Arch. Inst. Pasteur, Tunis, 27 : 133-183.
CANAMARES, F. T. 1951. La presencia del " Aedes (Stegomyia) vittatus Big." en al Medi-

terr&neo y algunes observaiciones sobre el mismo. Rev. Sanid. Hig. publ., Madrid, 25 :

435-443-
CARADJA, A. 1934. Herkunft und Evolution der palearktischen Lepidopterenfauna. Int.

ent. Z. 28 : 217-224.
CHOW, C. Y. 1949. Culicine mosquitoes collected in Western Yunnan, China during 1940-42.

Proc. ent. Soc. Wash. 51 : 127-132.
1950. Collection of Culicine mosquitoes (Diptera, Culicidae) in Taiwan (Formosa),

China, with description of a new species. Q. J. Taiwan Mus. 3 : 281-287.
DE BEAUFORT, L. F. 1951. Zoogeography of the Land and Inland Waters. London.
DE GRANDPRE, A. D., & DE CHARMOY, o'E. 1900. Les moustiques. Port Louis.
DRAKE-BROCKMAN, R. E. 1911. Stegomyia fasciata in North-east Africa. Bull. ent. Res.

2 : 179.

THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 63

EDWARDS, F. W. 1920. Notes on the mosquitoes of Madagascar, Mauritius and Reunion.
Ibid. 11 : 133-138-

1922. Mosquito notes : III. Ibid. 13 : 75-102.

1923 bis. Four new African mosquitoes. Ibid. 13 : 397-399.

19240. Some mosquitoes from Ovamboland, S.W. Africa and from the Cape Province.

Ann. S. Afr. Mus. 19 : 159-163.

19246. Description of two new species of mosquitoes from coconut palms in East Africa.

Trans. R. Soc. trop. Med. Hyg. 18 : 187.

1925. Mosquito Notes : V. Bull. ent. Res. 15 : 257-270.

1935- Mosquito Notes : XII. Ibid. 26 : 27-136.

1940. A new plain-winged Anopheles from Rhodesia. Ann. trop. med. Parasit. 34 :

93-96.

ENDERLEIN, G. 1921. Die Culiciden fauna Madagascars. Wien. ent. Ztg. 38 : 47-52.
FENG, L. C. 1935. Notes on some mosquitoes collected from Shantung Province, North

China. Chinese med. J. 49 : 359-365.

i 9380. A critical review of literature regarding the records of mosquitoes in China :

Pt. II. Peking nat. Hist. Bull. 12 : 285-318.

19386. The tree hole species of mosquitoes of Peiping, China. Chinese med. J., Suppl.

II : 503-525-

FORBES, H. O. (Ed.) 1903. The Natural History of Sokotra and Abd-el-Kuri. Liverpool.
FURON, R. 1943. Manuel de Prehistoire Generale. Paris.
GANDER, L. 1951. Experimentelle und Oekologische Untersuchungenuber das Schlupfvermogen

der Larven von Aedes aegypti L. (Thesis for the Ph.D. degree, University of Basel.)
GARNHAM, P. C. C. 1949. Acrodendrophilic mosquitoes of the Langata Forest, Kenya.

Bull. ent. Res. 39 : 489-490.

and McMAHON, J. P. 1947. The eradication of Simulium neavei, Roubaud, from an

Onchocerciasis area in Kenya Colony. Bull. ent. Res. 37 : 619-628.

GiAQuiNTO-MiRA, M. 1950. Notes on the geographic distribution and biology of " Anophe-

linae " and " Culicinae " in Etiopia. Riv. Malarial. 29 : 281-313.
GILLETT, J. D. I95ia. The habits of the mosquito Aedes (Stegomyia) simpsoni Theobald in

relation to the epidemiology of yellow fever in Uganda. Ann. trop. Med. Parasit. 45 :

IIO-I2I.

19516. The larva, pupa and adult male of Aedes (Stegomyia) ruwenzori Haddow and

van Someren (Diptera, Culicidae). Ibid. 45 : 195-198.

GRJEBINE, A. 1950. Moustiques du Moyen-Congo. Bull. Inst. Etud. centrafr., N.S. 1 :

25-48.

HARPER, J. O. 1947. A mosquito survey of Mahe, Seychelles. E. Afr. med. J. 24 : 25-29.
HICKS, E. P., & DIWAN CHAND. 1936. A mosquito survey of Karachi airport. Rec. malar.

surv. India, 6 : 515-535.
Ho, Cn'I. 1931. Study of the adult Culicids of Peiping. Bull. Fan Inst. Biol. Peking 2 :

107-175.
HOPKINS, G. H. E. 1952. Mosquitoes of the Ethiopian Region. I. Larval Bionomics of

Mosquitoes and Taxonomy of Culicine Larvae. 2nd ed. London : Brit. Mus. (Nat.

Hist.).
HUTTEL, V. 1950. Note sur la repartition des moustiques dans le Bas-Dahomey. Bull.

Soc. Path. exot. 43 : 563-566.
IYENGAR, M. O. T. 1938. Studies on the epidemiology of Filariasis in Travancore. Indian

med. Res. Mem. No. 30.
JEPSON, W. F., MOUTIA, A., & COURTOIS, C. 1947. The malaria problem in Mauritius ; the

bionomics of Mauritian anophelines. Bull. ent. Res. 38 : 177-208.
KING, H. H. (1908) Report on economic entomology. Rep. Wellcome trop. Res. Lab. 3 :

201-248.
KNIGHT, K. L., & HURLBUT, H. S. 1949. The mosquitoes of Ponape Island, eastern Carolines.

/. Wash. A cad. Sci. 39 : 20-34.

64 THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION

KNIGHT, K. L., and ROZEBOOM, L. E. 1946. The Aedes (Stegomyia) albolineatus group

(Diptera, Culicidae). Proc. biol. Soc. Wash. 59 : 83-96.
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12 : 401-409.

1939. The medical entomologist's report. Annu. med. Rep. Nyasald. (1938).

LEGENDRE, J. 1918. Note sur les Stegomyia de Tamatave. C. R. Soc. Biol. Paris, 81 :

832-833-
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mosquitoes. /. trap. Med. Hyg.' 30 : 136-145.

LINNAEUS, C. 1762. D. F. Hasselquists . . . Reise nach Palestina. Rostock.
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man, in Bwamba County, Uganda. Bull. ent. Res. 42 : 317-330.
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A study in relation to the sylvan epidemiology of yellow fever. Ibid. 42 : 721-760.

& BUXTON, A. P. 1951. A study of the epidemiology of yellow fever in the West Nile

district, Uganda. Trans. R. Soc. trap. Med. Hyg. 45 : 53-78.

MACGREGOR, M. E. 1924. Aedes (Stegomyia) mascarensis, a new mosquito from Mauritius.

Bull. ent. Res. 14 : 409.

McHARDY, J. W. 1932. Annu. med. (sanit.} Rep. Tanganyika (1928) : 45.
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(Theobald). (Thesis submitted for the degree of Ph.D. at Cambridge University.)
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moustique de la fievre jaune. C. R. Soc. biol. 117 : 878-880.
MATTINGLY, P. F. 1952. 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. Bull. Brit. Mus. (Nat. Hist.)

Ent. 2 : 235-304.

and LIPS, M. Notes on the Culicini (Diptera, Culicidae) of the Katanga. Rev. Zool.

Bot. afr.. (In Press.)

PEREIRA, M. DE C. 1946. Culicideos da Provincia do Sul do Save (Africa Orientale Portu-

guesa). Ann. Inst. Med. trop., Lisbon, 3 : 341-364.
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bionomics as observed during 1943-45. Pakistan J. Hlth. 1 (2) : 26-32.
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special reference to Plasmodium gallinaceum. J. Parasit. 28 : 127-129.
SASA, M., and KANO, R. 1951. Description and comparative studies on the larva of Aedes

(Stegomyia} galloisi Yamada (Diptera, Culicidae). Jap. J. exp. Med. 21 : 109-113.
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THE SUB-GENUS STEGOMYIA IN THE ETHIOPIAN REGION 65

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VAN SOMEREN, V. G. L., & DE BOER, H. S. 1926. Report on the mosquito breeding areas

of Nairobi. Kenya med. J. 2 : 275-298.
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Paris, 10 : 550-555.
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15: 575-585.

YAMADA, S. 1921. Description of ten new species of Aedes found in Japan, with notes on
the relation between some of these mosquitoes and the larva of Filaria bancrofti Cobbold.
Annot. zool.jap. 10 : 45-81.

PRESENTED

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ENTOM. Ill, I.

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' '^

A PEST OF COCONUT PALMS IN
PORTUGUESE EAST AFRICA

TWO NEW REPRESENTATIVES OF

THE GENUS HYPOTRABALA

(LEPIDOPTERA: LASIOCAMPIDAE)

FROM THE BELGIAN CONGO

W. H. T. TAMS

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 3 No. 2

LONDON: 1953

A PEST OF COCONUT PALMS IN
PORTUGUESE EAST AFRICA

TWO NEW REPRESENTATIVES OF THE

GENUS HYPOTRABALA (LEPIDOPTERA :

LASIOCAMPIDAE) FROM THE

BELGIAN CONGO

BY

W. H. T. TAMS

I

Pp. 67-75; Pis. 1-2; Text-figures 1-14, 9-16

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A PEST OF COCONUT PALMS IN
PORTUGUESE EAST AFRICA

By W. H. T. TAMS

IN the spring of 1952 Mr. V. Sleptzow, one of the London representatives of the
Sociedade Agricola do Madal, Quelimane, Africa Oriental Portuguesa, brought me
a photograph of larvae, cocoons and moths of a destructive pest that was attacking
coconut palms on a plantation at Micaune, Mosambique.

At that time I could do no more than state that the insect involved was a moth
belonging to the family Limacodidae, and I asked Mr. Sleptzow if he could procure
specimens from the plantation. A consignment was sent by air from Quelimane
on gth June, 1952, but there were no moths, and the small bottles containing larvae
had become loose and were smashed, completely ruining the whole consignment,
which arrived in a state of putrefaction. The larvae and cocoons were quite un-
known to me, and I asked Mr. Sleptzow if he would be good enough to try again.
He at once communicated with the African management of the company, with the
result that in the middle of September I received by air mail another consignment
of larvae and cocoons, and two battered crippled female moths, which had emerged
on the journey. I carefully nursed the cocoons at home, and succeeded in rearing
a number of moths of both sexes.

After a close study of the species I found that not only was it unknown to me,
but I could not find any close relative with which I could associate it generically.
Accordingly, I sent several examples to Dr. E. Martin Hering of the Berlin Museum.
Dr. Hering, who is the recognized authority on the family Limacodidae, informed
me that he fully agreed with me that the species did not belong to any known genus,
but appeared to be related to the genera in which the hind tibia bears two spurs
only, and the tarsi are clothed with scales rather than with the more delicate hair-
scales. It may be related to the Zinara group, and has some features suggestive
of Sporetolepis Karsch, which, however, has at the end of the fore wing cell the
fork of the parting-vein deep and closed distally by a cross-vein, whereas in
the genus here described the fork is shallow and open.

TROGOCRADA gen. n.
Typus generis Trogocrada deleter sp. n.

Antenna in $ pectinate, the pectinations decreasing slightly in length distad.
Antenna in $ simple. Palpus stout, upcurved before frons, 3rd segment small,
almost spherical, inconspicuous. Hind tibia with only one pair of spurs ; mid
tibia with a bushy covering of long scales, all tarsi covered with scales (not fine
hair-scales) (fig. 3, a, b, c, d). Fore wing with veins Sc, Ri and R2 separate and

ENTOM. Ill, 2. 6

7 o

COCONUT PALMS IN PORTUGUESE EAST AFRICA

almost straight ; veins R3 + R4 + R5 stalked, medials separate, M2 nearer to M3
than to Mi ; parting-vein within the cell terminating in an open shallow fork.
Hind wing with vein Sc anastomosing with middle of anterior margin of cell ; vein
Rs stalked with vein Mi for proximal fourth ; lower angle of cell not appreciably
nearer termen than upper angle. (Figs. 4, 5.)

FIGS. 1-3. Trogocrada deleter sp. n. Fig. i, male ; fig. 2, female ; fig. 3, legs (a) fore, (b) mid, (c)
hind, to show vestiture, (d) hind, to show tibial spurs.

Trogocrada deleter sp. n.

(Figs, i, <$, 2 ?.)

c. Antenna warm buff, degraded with fuscous. Palpus mainly warm blackish
brown, warm buff apically. Frons warm blackish brown, vertex apricot buff.
Thorax warm to apricot buff, sometimes degraded with fuscous. Abdomen fuscous,
lighter near base. Pectus and legs light to apricot buff, shaded with warm blackish
brown in front. Venter fuscous, lighter medially. Fore wing apricot buff, proximal
three-fourths heavily over-shaded with fuscous to warm blackish brown, with a
wavy appearance between the velvety blackish-brown lunules medially and a
strong, velvety, blackish-brown curved postmedial fascia (cf. fig. i), the distal
portion of the wing suffused with tawny. Hind wing fuscous to fuscous black.

Underside of both fore and hind wings fuscous to fuscous black. Expanse :
22-24 mm.

COCONUT PALMS IN PORTUGUESE EAST AFRICA 71

$. Similar in pattern to $ ; the predominating colour may be apricot buff with
tawny to fuscous shading beyond the postmedial fascia on the fore wing, the medial
markings and postmedial fascia velvety blackish brown, or the whole wing may be
suffused with fuscous. Underside may be apricot buff or fuscous. Expanse :
26-27 mm.

st.

FIGS. 4, 5. Trogocrada deleter sp. n. Fig. 4, fore-wing venation ; fig. 5, hind-wing venation.

< genitalia. The male genitalia exhibit a robust structure of segments IX-X,
the division between the terga being difficult to discern. The various structures
are better displayed by figures than by descriptions. The valva is most unusual
in shape and possesses distally a sort of loose " corona " of woolly hair-scales, and
two fairly stout whip-like setae (figs. 8-13).

$ genitalia. A very lightly sclerotised ostium bursae ; ovipositor lobes broad
and prominent.

72 COCONUT PALMS IN PORTUGUESE EAST AFRICA

Larva (fig. 6). The larva is green with velvety darker green shading and a variable
line down the middle of the dorsum, frequently white edged with reddish, or entirely
reddish. In shape it is quite unlike any Limacodid larva known to me.

Cocoon (fig. 7). The larva makes an almost spherical cocoon (9 mm. X 8 mm.),
whitish with irregular blackish markings.

FIGS. 6, 7. Trogocrada deleter sp. n. Fig. 6, larva ; fig. 7, cocoon.

Holotype <$, allotype $ and 8 paratypes:

Portuguese East Africa, Micaune, destroying coconut plantations. Presented
by the Sociedade Agricola do Madal, through Messrs. E. Isen (Quelimane), M.
Terestchenko and V. Sleptzow, to whom I am grateful for the opportunity to study
such an interesting moth.

COCONUT PALMS IN PORTUGUESE EAST AFRICA

73

14

FIGS. 8-14. Trogocrada deleter sp. n., <J genitalia. Fig. 8, complete armature, valvae (shaded)
unspread ; fig. 9, complete armature, valvae spread ; fig. 10, valvae, anellus and aedoeagus
separated from segments IX-X ; fig. n, segments IX-X ; fig. 12, sternum X (gnathus) ;
fig. 13, terga IX-X (tegumen and uncus) ; fig. 14, sternum IX (vinculum).

TWO NEW REPRESENTATIVES OF

THE GENUS HYPOTRABALA

(LEPIDOPTERA : LASIOCAMPIDAE)

FROM THE BELGIAN CONGO

By W. H. T. TAMS

IN 1925 I drew attention (Ann. Mag. Nat. Hist. (9) 16 : 552-561) to the fact that
certain species of African Lasiocampidae were not associated in their proper genera,
and I dealt particularly with the genera Pseudometa Aurivillius and Hypotrabala
Holland. I have since discovered that Pseudometa schultzei Aurivillius is not a
Pseudometa but belongs in the genus Mallocampa Aurivillius.

Some time ago Monsieur L. A. Berger of the Congo Museum at Tervuren, Belgium,
brought me two further examples of the genus Hypotrabala, and these are described
below. I have given in this paper a few figures of the eighth sternum in various
species of this genus, as this structure may be readily observed if the scales are
carefully removed from the underside of the terminal abdominal segments, and it is
an important aid to identification.

Hypotrabala regalis sp. n.

(PI. i, figs. 3, <?, 4, $ ; PL 2, figs. 5, 6.)

c. Antenna honey yellow, the shaft with some chocolate scales. Head, palpus,
thorax, abdomen and legs light orange yellow, with some chocolate shading. Fore
wing pale orange yellow enriched with light orange yellow, the dark pattern indicated
in the photograph chocolate. At the end of the cell a large glossy white oval spot,
finely edged with chocolate, with a much smaller spot of like appearance between it
and the costa, from which it is slightly more distant. Hind wing pale orange yellow
without pattern. Underside pale orange yellow. Expanse 82 mm. (77 mm. from
tip-to-tip).

$. Similar, much larger, richer in colour (light orange yellow), with some fuscous
shading in the subterminal markings ; a single smaller discocellular spot, chocolate
with or without a few white scales centrally.

Expanse 114 mm. (104 mm. from tip-to-tip).

Belgian Congo, Lulua, Kapanga (F. G. Overlaet).

Holotype <$: i8.xii.i933, 4 $ paratypes dated 13. ix. 1932, x. 1933 and xi.i933
(2) respectively. Allotype $ : 8.xii.i932, 2 $ paratypes dated 12, I4.xii.i932,
respectively.

The $ genitalia are unlike those of the other Hypotrabala species known to me
in having a single medial process on sternum VIII (PI. 2, fig. 5). The $ genitalia
are shown in PL 2, fig. 6 ; they are strongly sclerotized, with a robust bar before
the ostium.

Bull. B.M. (N.H.) Entomology III. 2.

PLATE i

Bull. B.M. (N.H.) Entomology III, 2.

PLATE 2

HYPOTRABALA.
GENITALIA.

THE GENUS HYPOTRABALA FROM THE BELGIAN CONGO

75

Hypotrabala guttata fontainei ssp. n.
(PI. i, fig. i, <$ ; PL 2, figs. 7, 8.)

$. Slightly larger than H. guttata guttata Aurivillius (PL i, fig. 2), with a larger
discal glossy white spot. <$ genitalia (PL 2, figs. 7, 8, text-fig. 10) with sternum
VIII bearing two long straight processes, the shape of the sternum being quite
different from that of H. guttata guttata (text-fig. 9), which came from the Gold
Coast. Expanse 70 mm. (66 mm. from tip-to-tip).

Holotype <$ : Belgian Congo, Lusambo, 13. x. 1949 (Dr. M. Fontaine).

16

FIGS. 9-16. (J VIHth sternum in the genus Hypotrabala. 9. Hypotrabala guttata guttata. 10. H.
juttata fontainei. 1 1 . H. porphyria. 12. H. horridula. 13. H. dollmani. i^.H.neavei. 15. H. joiceyi.
1 6. H. horridul i seydeli.

EXPLANATION OF PLATES AND FIGURES

PL. I, FIGS. 1-4. Hypotrabala guttata fontainei <J (fig. i), H. g. guttata <$ (fig. 2) ; Hypotrabala
regalis cJ (fig. 3), ? (fig. 4).

PL. II, FIGS. 5-8. Hypotrabala regalis $ genitalia (fig. 5), and ? (fig. 6) ; H. guttata fontainei (J
genitalia (fig. 7), <$ sternum VIII (fig. 8).

ENTOM. Ill, 2. 7

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25 NOV 1953

A REVISION OF THE GENUS

TERIOMIMA KIRBY

(LEPIDOPTERA: LYCAENID AE)

H. STEMPFFER AND N. H. BENNETT

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 3 No. 3

LONDON : 1953

A REVISION OF

THE GENUS TERIOMIMA KIRBY
(LEPIDOPTERA: LYCAENIDAE)

BY

H. STEMPFFER AND N. H. BENNETT

V

Pp. 77-104 ; Pis. 3-9 ;

BULLETIN OF
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ENTOMOLOGY Vol. 3 No. 3

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THE BULLETIN OF THE BRITISH MUSEUM
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issued in five series corresponding to the Departments
of the Museum, and an Historical Series.

Parts appear at irregular intervals as they become
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hundred pages, and will not necessarily be completed
within one calendar year.

This paper is Vol. 3, No. 3 of the Entomological
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PRINTED BY ORDER OF THE TRUSTEES OF
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Issued November, 1953 Price Seven Shillings and Sixpence

A REVISION OF THE GENUS TERIOMIMA
KIRBY (LEPIDOPTERA LYCAENIDAE)

By H. STEMPFFER AND N. H. BENNETT

SYNOPSIS

Teriomima, a small genus of African butterflies, of the family Lycaenidae, is distributed
throughout the E. and S.E. regions. Little attention has been paid to them since 1918, when
Aurivillius, in Seitz : The Macrolepidoptera of the World, 13 : 327-328, dealt with the few species
then recognized. An exhaustive study of the male genitalic structure has led the authors to
describe two additional genera, eight new species and one new subspecies, all contained within
the earlier conception of the genus.

i. HISTORY OF THE GENUS

THE Liptenine genus Teriomima was described by Kirby in 1887 (Ann. Mag. Nat.
Hist. (15) 19 : 364-367). In it he placed Pieris erastus Hewitson (1866) and the
following seven new species which he described at the same time, namely Teriomima
subpunctata (the type species), T. puella, T. tenera, T. similis, T. erasmus, T. flaveola,
T. dispar and T. hildegarda. In 1888 H. H. Druce described and added Teriomima
melissa (Ent. Mon. Mag. 25 : 109). In 1890 Kirby (Ann. Mag. Nat. Hist. (6) 6 :
268-271) added a further six new species, T. decipiens, T. delicatula, T. serena, T.
modesta, T. cordelia and T. dubia, and transferred Hewitson's Liptena adelgitha
(1874) to the genus. In the same year W. J. Holland (Psyche, 5 : 429) added Terio-
mima leucostola, T. xanthostola and T. xanthis. Two years later Staudinger (7ns,
D. ent. Zeit. 4 : 219-220) added Teriomima adelgunda and T. alberta, both new, and
in 1894 Smith and Kirby (Rhop. Ex. 2 : 115, Afr. Lye., pi. 25, figs, i and 2) added
T. freya. In 1895 T. galenides Holland (Ent. News Philad. 6 : 167) appeared. In
1898 Grose Smith (Novit. Zool. 5 : 355) extended the list with T. pusio and T.fuscula.

Aurivillius, in Rhopalocera Aethiopica (1898), included in Teriomima the following
species : T. subpunctata Kirby, T. delicatula Kirby, T. puella Kirby, Durbania
puellaris Trimen (Proc. Zool. Soc. Lond. 1894 : 59-60), T. hildegarda Kirby with var.
freya Smith & Kirby, Liptena aslauga Trimen (Trans. Ent. Soc. 1873 : 117), Durbania
pallida Trimen (Trans. Ent. Soc. 1898 : 12) and Durbania micra Grose Smith (Novit.
Zool. 5 (1898) : 356). He excluded from the genus Pieris erastus Hewitson, T.
tenera Kirby, T. pusio Kirby, T. similis Kirby and T. serena Kirby, attributing them
to the genus Citrinophila on neuration. T. erasmus Kirby and T. flaveola Kirby
he treated as synonyms of P. erastus Hewitson.

T. xantha was described by Grose Smith in 1901 (Rhop. Ex. 3 : 140, Afr. Lye.,
pi. 29, figs. 13, 15).

In Seitz (Macrolep. 13 : 327-328, 1918), Aurivillius listed the following as Terio-
mima species : T. subpunctata Kirby, T. delicatula Kirby, T. puella Kirby, T.

ENTOM. Ill, 3. 8

8o A REVISION OF THE GENUS TERIOMIMA KIRBY

puellaris Trimen, T. micro, Grose Smith, T. aslauga Trimen, and treated hildegarda
Kirby and pallida Trimen as forms of aslauga owing to the great variability of their
wing patterns. Hawker Smith described T. parva, T. minima and T. minima
latimarginata in 1933 (Stylops, 2 : 6-7) ; Talbot added T. minima f. stygia in 1935
(Ent. Mon. Mag. 71 : 72) ; Riley described T. woodi in 1943 (Entomologist, 76 :
225-226) ; van Son described T. zuluana in 1949 (Ann. Transv. Mus, 21(2) : 211),
placing this species closest to T. puellaris Trimen.

This appears to be the history of the genus to date. Of the thirty-seven names
cited seven have been absorbed into Citrinophila, viz., tenera, similis, erasmus,
erastus, flaveola, serena and pusio ; seven are now placed in Liptena, viz., decipiens,
modesta, leucostola, xanthostola, xantha, xanthis and galenides ; four in Epitolina,
viz., melissa, dispar, Cordelia and dubia ; four are now in Micropentila, viz., adel-
githa, adelgunda, alberta andfuscula.

2. METHOD OF REVISION

The dissection of a number of specimens labelled " aslauga " revealed that several
species were involved. Late in 1949 each of the authors commenced, by arrange-
ment, an independent survey of the genitalic forms within the genus. Each having
access to a different source of material, and employing varying techniques, it was
hoped that the possibility of errors of interpretation might thereby be greatly
reduced. When the two sets of results were correlated an encouraging degree of
unanimity was at once apparent. Where divergences occurred they were usually
due to a shortage of material, always the main obstacle to this kind of investigation.
An extensive exchange of specimens, genitalic preparations, etc., soon confirmed the
mutual realization that there were differences of structure of sufficient diversity
and stability to warrant regrouping into three genera, two of which are new, as
indicated below :

1. TERIOMIMA Kirby, to include subpunctata, puella, etc.

2. BALIOCHILA genus nov., to include aslauga, hildegarda, etc.

3. CNODONTES genus nov. to include pallida, etc.

At an early stage both workers agreed that it is impossible, except in a few in-
stances, to make reliable specific determinations from external appearances. This
difficulty is well illustrated by the fact that Aurivillius, for all his experience, placed
the bulk of our species of Baliochila and Cnodontes under a single specific name in
Seitz, wherein hildegarda and pallida are treated as subspecies or seasonal forms of
aslauga. Even when long series have been positively identified by dissection, such
is the intergradation of wing pattern and coloration, especially in Baliochila,
that it would still be hazardous to name most specimens without examining the
genitalia, even with the rich material (eleven hundred specimens) available for
comparison in the British Museum. The descriptions given later in this work might
suggest a great diversity of form, but what frequently occurs is no more than a
confusing reversal of ground-colour and pattern. In plate 9 an attempt is made to
illustrate the extremes of variation within the species B. hildegarda and B. dubiosa.

A REVISION OF THE GENUS TERIOMIMA KIRBY 81

Also shown are examples of B. neavei and C. vansomereni, which differ greatly in
appearance from the selected type-specimens.

From the foregoing it will be appreciated that the formulation of a key based
on external appearances would be a waste of time. A key to the genitalia could
be readily produced, but it is considered that the complete set of illustrations of these
organs which accompanies this work is so self-explanatory as to render a key super-
fluous.

The effectiveness of this revision was greatly enhanced by Professor Varley, of
the Hope Department of Entomology, University Museum, Oxford, who, through
the kind offices of Professor Hale Carpenter, lent the whole of the Hope Department
collection of Teriomima, amounting to more than three hundred and fifty specimens,
to facilitate this undertaking. We are indebted to Dr. V. G. L. van Someren for
a similar act of co-operation. Our thanks are also due to T. H. E. Jackson, Esq.,
for numerous contributions of material, as well as for his " on-the-spot " knowledge
of the genus, so readily placed at our disposal ; to E. B. D. Barnes, Esq., of Umtali,
who provided examples of the new species B. barnesi and B. lipara, and to Dr. van
Son, who furnished some examples of B. aslauga. We are also indebted to W. H. T.
Tarns, Esq., for much sound advice on genitalic problems.

3. SYSTEMATIC ACCOUNT OF THE GENERA AND SPECIES

Genus TERIOMIMA Kirby (sensu stricto).
Kirby : Ann. Mag. Nat. Hist. (5) 19 : 364 (1887).

Type of the genus : Teriomima subpunctata Kirby (1887) designated by Kirby.
Eyes bare ; palpi rather long, ascending, projecting beyond the frons, clothed
with adpressed scales, the second joint stout, laterally compressed, the third joint
slender, acuminate ; antennae rather short, with a distinct, subcylindrical club ;
male forelegs with unsegmented tarsi, clothed beneath with fine spines. Wing
shape. Fore wings : costa evenly arched, rather rounded apex, outer margin strongly
convex. Hind wings : oval, anal angle not very distinct. Neuration : Fore wing
with twelve veins ; cell short ; vein 4 from the lower angle of the cell ; posterior
discocellular concave ; vein 5 nearer to 6 than to 4 ; 6 and 7 arising from a common
stalk from the upper angle of the cell ; 8 and 9 branching from 7 ; 10 and n free,
from the upper edge of the cell, 10 arising very close to the upper angle. Hind
wing : cell rather short ; 3 arising a little before the lower angle of the cell ; 4
from the angle ; posterior discocellular concave ; 5 nearer to 6 than to 4 ; 6 from
the upper angle of the cell ; 7 from just before the upper angle.

MALE GENITALIA : GENERAL DESCRIPTION. Uncus bifid, consisting of two more or
less pointed finger-like processes ; no subunci ; tegumen fairly wide ; vinculum of
moderate width ; anellus fused to the bases of the valvae, forming a narrow sheath
to the base of the aedoeagus ; valva oblong, with a pointed, more or less falcate
apex, sometimes having a curved, finely pointed harpe extending over the distal
third of its length ; aedoeagus fairly stout, the distal end having one or two ventrally-
directed lobes which may be long and curved, as in the subpunctata-group, or rather
short and blunt, as in the micra-group.

82 A REVISION OF THE GENUS TERIOMIMA KIRBY

(i) Teriomima subpunctata Kirby (1887)
(PI. 3, fig. i, neallotype <$ ; PI. 5, fig. 26, $ genitalia)

Teriomima subpunctata Kirby, 1887, Ann. Mag. Nat. Hist. (5) 19 : 364, W. Africa (?).
syn. Teriomima delicatula Kirby, 1890, Ann. Mag. Nat. Hist. (6) 6 : 269, Usagara.

(NoxE. The treatment of delicatula as a synonym is the result of a careful exami-
nation of over 150 specimens ; it is impossible to find any constant points of differ-
entiation in facies or genitalia.)

Holotype ($) in the British Museum (N.H.). <$ not previously described.

c. Frons brown with two lateral fine white lines ; second joint of the palpi
laterally compressed, clothed with white scales at the centre, brown at the extremi-
ties, third joint white tipped ; antennae black, ringed with white, club black with
reddish-brown tip ; tarsi black, ringed with white.

UPPERSIDE FORE WING. White, with a costal margin of light brown scales ex-
tending one-third from the base, a minute patch of the same colour between the
extremities of veins n and 12 ; a triangular apical patch, rather darker, from
the extremity of 10 on the costal margin to midway between 2 and 3 on the outer
margin.

UPPERSIDE HIND WING. White, with a very small patch of light brown scales
at the tip of each vein.

UNDERSIDE FORE WING. Ground-colour white, yellowish at the costal margin
and in the apical area ; lightly patterned with small patches of mixed pale and dark
brown scales arranged as follows : five evenly spaced patches from the base to mid-
way along the costal margin ; three evenly spaced patches from the base of the
cell to the discocellulars ; a straight row of three patches disposed between veins
9 and 10, 6 and 5 and 5 and 4 ; a curved row of five spots corresponding to the
inner edge of the apical patch on the upperside, the lowermost between veins 3 and
4 ; two rows, marginal and submarginal, of very small spots from the apex to vein 3.

UNDERSIDE HIND WING. Ground-colour white, with a rather more evenly dis-
tributed pattern of light and dark brown patches than on the fore wing ; although
very irregular, these patches may be placed in six rows, the fourth from the base
being the most heavily marked.

Fringes of both wings white except at the apex of the fore wing, where they are
brown with interruptions of white.

LENGTH OF FORE WING. -15-6 mm.

J GENITALIA. Uncus bifid, lobes dilated at the base and tapering to blunt points,
tips ventrally inclined, sparsely hairy ; no subunci ; tegumen fairly large ; vinculum
of moderate width ; anellus sheathing the base of the aedoeagus, which is long and
robust, curved in the form of a flattened S and bearing, near the distal end, two ven-
trally directed lobes or flaps, the distal end blunt ; valva flask-shaped, with a long,
curved harpe, apex weakly falcate.

Neallotype $ : Brit. E. Afr., Uchweni Forest, 1-2. iii. 1912 (S. A. Neave), in the
British Museum (N. H.) B.M. Type No. Rh. 15684.

DISTRIBUTION. Kenya, Tanganyika.

A REVISION OF THE GENUS TERIOMIMA KIRBY 83

(2) Teriomima puella Kirby (1887)
(PL 3, fig. 2, neallotype $ ; PI. 5, fig. 27, <$ genitalia)
Teriomima puella Kirby, 1887, Ann. Mag. Nat. Hist. (5) 19 : 365, W. Africa.

Holotype (<$} in the British Museum (N.H.). $ not previously described.

$. External characters as in the male. A long series from Nyasaland, Mlanje,
show considerable variation of the costal markings of the upperside fore wing and
in the spotting of the underside hind wing.

Length of fore wing : 17 mm.

J GENITALIA. Very similar to those of subpunctata, but with narrower valvae,
having shorter and more sharply curved harpes.

Neallotype $ : Nyasaland, Mlanje, 2200', 8.xi.i3 (S. A. Neave), in the British
Museum (N.H.). B.M. Type No. Rh. 15685.

DISTRIBUTION. Nyasaland, N.E. Rhodesia, Tanganyika Territory.

(3) Teriomima zuluana van Son (1949)
(PI. 5, fig. 28, genitalia)

Teriomima zuluana van Son, 1949, Ann. Transv. Mus. 21 (2) : 211, pi. 3, figs. 5-8, text-fig, i,
Hluhluwe, Zululand.

Types in the Transvaal Museum.

For convenience the author's descriptions are given below. A male and a female
of this species have been made available for study by the courtesy of Mr. Pennington.

" $. Head : frons black, eyes edged with ochraceous white ; palpi black, third
joint white at tip ; antennae black, shaft chequered with white, club black above,
below with seven terminal joints reddish-brown. Thorax : black, clothed with
antimony yellow (Ridgway, pi. XV) scales and hairs above, and scales only of the
same colour below ; legs black, chequered with white, and with some antimony
yellow chequering on coxae and femora. Wings : antimony yellow above and below.
Upperside : fore wing, costa blackened from the base to less than one-third the
length of the cell, closely followed by a bar opposite the middle of the cell, another
opposite two-thirds of the cell, and a third at the middle of the costa (absent in
most specimens) ; apical area with the inner edge very oblique (almost parallel
with the inner margin) to vein M x , narrowed below the vein and reaching the middle
of area Cuj. Hind wing unmarked. Underside : fore wing : seven equidistant
small bar-like black dots, the first being placed a little before the level of the middle
of the cell and the third, very minute, a little before the level of the end of the cell ;
under magnification most of these dots appear to consist of separate minute bars
which are often unequally developed on the right and left side ; an interrupted
oblique bar from the sixth costal dot to vein M 1? (absent in many paratypes) ; a
black marginal line, thickened at the apex, interrupted between the veins and not
reaching below vein Cu 2 , where it is very indistinct. Hind wing : a black dot at
the middle of area RS ; postdiscal black dots in areas M 1( Cu 1} Cu 2 ; there is no post-
discal dot in area A 2 ; a rather diffuse discocellular dot tinged with orange, and a

ENTOM. Ill, 3. 8

84 A REVISION OF THE GENUS TERIOMIMA KIRBY

dot before middle of cell ; a black interrupted marginal line from vein Cu 2 to anal
angle (in many paratypes the dots of the underside of the hind wing are reduced or
absent, and are best marked in January specimens). Length of fore wing : 14-6
mm. Antenna-wing ratio : 0-42.

Description of the female allotype.

Larger than the male, antennal club, with a few greyish scales below, and
without any reddish brown. Fore wing : black apical area a little larger than
in the male and projecting inwards along vein M x ; otherwise like male.

Length of fore wing. 16 mm. Antenna-wing ratio : 0-42.

3 GENITALIA. Uncus double, with the prongs bent down at an obtuse angle at
the middle. Valve : very elongate, gradually widened from the narrow base to
two-thirds of its length, then rather abruptly narrowed and produced distally into
an acute, slightly down-curved spine ; ventral rim with a long claw-like process
arising midway from base to apex and directed distad, with the tip slightly down-
curved and reaching beyond the base of the apical spine ; its length is half the
length of the valve. Aedoeagus : half as long as the valve, stout, apically rounded,
with two elongate triangular prongs directed downwards and which are shorter than
the width of the aedoeagus ; vesica forming two elongate sacs."

DISTRIBUTION. Zululand.

(4) Teriomima puellaris Trimen (1894)

(PI. 5, fig. 29, (J genitalia)
Durbania puellaris Trimen, 1894, Proc. Zool. Soc. 1894 : 59, pi. 6, fig. 14, ?, Manica, Vunduzi R.

Types not examined. Presumably in the South African Museum, Cape Town.

<$ GENITALIA. Of the subpunctata-type, but with uncus lobes more slender ;
harpes shorter ; aedoeagus with very distinctive ventral lobes, long, sinuate and
tapering, distal end of aedoeagus forming a small point.

DISTRIBUTION. Mashonaland.

(5) Teriomima micra Grose Smith (1898)

(PI. 6, fig. 30, (J genitalia)
Durbania micra Grose Smith, 1898, Novit. Zool. 5 : 356, B.E.A., Tana River (Dr. Ansorge).

Types in the British Museum (N.H.).

(NOTE. The author described the male as the female and vice versa.)

$ GENITALIA. Uncus lobes straighter than in the subpunctata-type armatures,
more slender and sharply pointed ; no subunci ; tegumen quite long ; vinculum
and anellus as in the preceding species ; aedoeagus terminating in a sharp point
which arises from a zone thickly clad with fine hairs, ventral face deeply excised
near the distal end, then broadening to a blunt, ventrally directed lobe ; valva
without harpe, apex strongly falcate, with a pronounced angle on the ventral margin ;
uncus and distal half of valvae hairy.

DISTRIBUTION. Kenya, coast region.

A REVISION OF THE GENUS TERIOMIMA KIRBY 85

(6) Teriomima parva Hawker Smith (1933)
(PI. 3, fig. 3, neallotype ? ; PL 6, fig. 31 $ genitalia)

Teriomima parva Hawker Smith, <$ (nee ? = minima minima], 1933, Stylops, 2:6, 14 m.
N.W. of Mombasa, Rabai, 23.1.1904 (K. St. A. Rogers).

Holotype (J) in the British Museum (N.H.) ; $ not previously described.

$. UPPERSIDE fore wing and hind wing somewhat paler than in the male ; fore-
wing costal markings reduced to three pale brown spots and a very short subapical
band ; apical dark brown band extending to the tornus, but little more than half
as wide as in the male ; hind wing as in the male, apart from the slightly paler ground
colour. Underside fore wing and hind wing as in the male.

Length of fore wing : $, 12 mm.

$ Neallotype : 14 m. N.W. of Mombasa, Rabai, i3.vi.O3 (K. St. A. Rogers), in
the British Museum (N.H.), B.M. Type No. Rh. 15686.

c GENITALIA. Differ only very slightly from those of micra. The valvae are
less strongly falcate and there is no marked angle on the ventral edge to break the
smooth inward curve of the apex.

DISTRIBUTION. Kenya ; Port. E. Afr. ; Moa, Tanganyika Territory.

Genus BALIOCHILA gen. nov.

Type of the genus : Liptena aslauga Trimen, 1873.

Eyes, palpi, antennae, legs, wing-shape and neuration as in Teriomima. The
generic name was suggested by the spotted costa found in these species.

<J GENITALIA : GENERAL DESCRIPTION. All but one of the species included in
this genus share a common character, this being so distinctive that, in the authors'
estimation, other quite considerable divergences may be disregarded ; a pair of
processes, the base of each being strongly attached to the dorsal side of the anellus
on its inner margin and less strongly, with a half-membranous lesion to the tegumen,
on its outer margin. The processes rest more or less parallel with the axis of the
abdomen, midway between uncus and aedoeagus ; they are usually crowned with
strong spines, but in some species are completely covered with short, stiff bristles.
It is difficult to decide the exact homology of these processes ; they cannot be classed
as subunci as they are not articulated at the level of the uncus-tegumen suture, nor
can they be termed anellus lobes since they are also fused to the tegumen ; the
noncommital term " special processes " will therefore be used in the ensuing specific
descriptions. It is possible that they correspond with the asymmetrical processes
found in Ornipholidotos, which are articulated to the vinculum at a point approxi-
mating to the suture of the tergite and sternite.

Four groups of species are readily separated within the genus by the formation
of the uncus and the shape of the special processes :

(a) Uncus bifid, the two arms widely separated, more or less slender ; special
processes rather short ; aslauga Trimen, barnesi sp. nov., neavei
sp. nov., hildegarda Kirby, dubiosa sp. nov., nyasae sp. nov., stygia
Talbot.

86 A REVISION OF THE GENUS TERIOMIMA KIRBY

(b) Uncus a long flattened stalk, excised at the apex ; special processes long

and slender ; woodi Riley, fragilis sp. nov.

(c) Uncus forked, the arms slender and tapering, curved in both lateral and

ventral planes ; special processes long and robust ; minima Hawker
Smith, lipara sp. nov.

(d) Uncus forked, the arms mounted on a long central stalk ; no special pro-

cesses, but a pair of long, slender anellus lobes are found in their
place ; singular is sp. nov.

Other parts of the genitalia are similar to those of Teriomima ; the aedoeagus,
however, never has the distinctive ventral expansions found in the subpunctata-
group.

(7) Baliochila aslauga Trimen (1873)

(PI. 6, fig. 32, $ genitalia)
Liptena aslauga Trimen, 1873, Trans. Ent. Soc. 1873 : 117, Natal, Pinetown, near Durban.

Type not examined. Stated by Trimen to be in the collection of W. Morant,
Esq.

c GENITALIA. Uncus formed of two short, spiny arms, at the base of each a
small, acute triangular apophysis ; no subunci ; tegumen wide ; special processes
short and crowned with spines ; vinculum broad, anellus sheathing the lower part
of the aedoeagus ; valva flask-shaped with a falcate pointed apex ; aedoeagus long,
slightly excised on the ventral face, near the distal end.

DISTRIBUTION. S. Africa, Natal, Delagoa Bay, etc., in the British Museum (N.H.),
Zululand, Hluhluwe, in coll. Stempffer. False Bay, in coll. Jackson.

(8) Baliochila barnesi sp. nov.
(PI. 3, fig. 4, holotype <$ ; fig. 5, allotype ? ; PI. 6, fig. 33, < genitalia)

Types in the British Museum (N.H.).

(J. Frons grey or blackish-grey with two lateral fine white lines ; second joint of
the palpi laterally compressed, clothed with fuscous scales, with a scattering of
white on the underside, third joint white-tipped ; antennae fuscous, ringed white,
club fuscous, orange-tipped ; legs fuscous, ringed white.

UPPERSIDE FORE WING. Orange-yellow with dark brown markings ; costal border
extending about two-thirds from the base, with two clearly-marked dots of the ground-
colour between costa and margin ; occasionally an additional one or two less clearly-
marked dots may be present ; some fuscous scales in the cell ; one fuscous bar from
the costal border covering the discocellulars ; another, larger curved bar from the
costal border to vein 3 ; apex broadly fuscous and extending as a tapering marginal
band to the tornus, the inner edge of this band rather irregular.

UPPERSIDE HIND WING. Coloration as in the fore wing ; the dark crescentic
marginal border extending from the apex to the anal angle, about 2 mm. wide at its
centre ; inner edge irregular.

A REVISION OF THE GENUS TERIOMIMA KIRBY 87

Fringes fuscous, with white patches between veins 5-6-7 of the fore wing ; irre-
gular white patches from the apex to the anal angle of the hind wing.

UNDERSIDE FORE WING. Ground-colour less orange than on the upperside and
shading to pale yellow towards the inner margin ; all markings of the same tint as
the hind wing ground-colour ; costal border with four evenly spaced yellow dots
between the costa and the margin ; from the costal border run four bars, one crossing
the cell near its base, a second crossing the cell about midway, a third covering the
discocellulars, the fourth and largest curving from the margin to vein 3 ; apical
pattern consisting of a triangular patch based on the costal margin curving outward
and terminating at vein 4 ; outside this an antemarginal band, crenulate and tapering
from the costa to vein 2 ; a marginal line tapering from the apex to vein 2.

UNDERSIDE HIND WING. Ground colour dark violet-grey with a boldly marked
pattern consisting of numerous orange spots disposed in five irregular bands, in
certain examples the third band accentuated by a heavy fuscous outline to the
outer side of each spot ; between the veins a submarginal series of triangular orange
markings.

Fringes as on upperside.

( GENITALIA. Uncus formed of two arms, longer and more slender than in aslauga,
the apices slightly expanded and curving ventrally, with numerous strong spines ;
at the base of each arm a small, laterally-directed apophysis ; tegumen wide ;
special processes strongly curved as in aslauga, the apices broadened, bearing numer-
ous strong spines ; valvae typical of the genus, apices falcate, distal end of the
aedoeagus bearing a sharp tooth on either side of the dorsal surface ; ventral curve
deeply excised.

Readily distinguished from the armature of aslauga by the longer arms of uncus
and by the toothed aedoeagus.

$. Frons, palpi, antennae and legs as in the male.

UPPERSIDE FORE WING. Orange yellow with fuscous markings, rather lighter
than the male ; costal border extending about two-thirds from the base, narrower
than in the male, not encroaching upon the cell ; small yellow marginal dots varying
as in the male ; no bar over the discocellulars ; a small bar extending from the
termination of the costal border to midway between veins 6 and 5 ; apex and
marginal border as in the male, but the latter reduced in width between the ex-
tremities of veins 3 and i.

UPPERSIDE HIND WING. Marginal border narrower than in the male, extending
from the apex to the anal angle.

Fringes of both wings fuscous, interrupted with white.

UNDERSIDE FORE AND HIND WING. Coloration as in the male, but the two bars
crossing the base and middle of the fore-wing cell do not quite reach its lower border.

Length of fore wing : <$, 14 mm.
?, 13 mm.

Holotype <^ : Mineni Valley, Manica, S.E. Trop. Afr., in the British Museum
(N.H.), B.M. Type No. Rh. 15687.

Allotype $ : Mineni Valley, Manica, S.E. Trop. Afr., in the British Museum
(N.H.), B.M. Type No. Rh. 15688.

88 A REVISION OF THE GENUS TERIOMIMA KIRBY

Paratypes : Mineni Valley, 4$ ; Chirindi Mt., Mashonaland 2$ ; Chirindi
Forest, Gazaland i^ ; Umtali, Mashonaland i$ ; Mezimbiti,
Port. E. Afr. 2^, all in B.M. (N.H.). S. Rhodesia, Vumba
Mtns., !( i? in Stempffer coll. S. Rhodesia, Vumba Mtns.,
i(J i$ in Jackson coll.

(9) Baliochila neavei sp. nov.

(PI. 3, fig. 6, holotype J ; fig. 7, allotype ? ; PL 6, fig. 34, <$ genitalia ; PI. 9, fig.

65, c, Mozambique)

Types in the British Museum (N.H.).

(. Frons grey with two lateral fine white lines ; palpi, antennae and legs as in
B. barnesi.

UPPERSIDE FORE WING. Orange yellow with dark brown markings as follows :
costal border interrupted with two small yellow dots ; some dark scales at base
and middle of the cell near the upper edge ; a streak covering the discocellulars ;
a curved patch from the costal border to vein 4 ; some dark scales between veins
4 and 3 ; a large apical patch prolonged to the tornus, its inner edge irregular.

UPPERSIDE HIND WING. Orange yellow with a wide dark brown border from the
costa to the anal angle.

Fringes of both wings dark brown, interrupted with white.

UNDERSIDE FORE WING. Orange yellow shading to very pale yellow at the
inner margin, with shining violet grey markings ; four small yellow dots spaced
along the costal margin ; three transverse streaks crossing the cell, one at its base,
one at the middle, the third covering the discocellulars ; a larger, curved patch
from the costa to vein 2 ; in the apical zone an oblique line running from the costa
to vein 2 ; a crenulate line, parallel with the margin, from costa to tornus ; a shining
grey marginal line, wide at apex, narrowing to the tornus.

UNDERSIDE HIND WING. Ground-colour shining violet grey, with six transverse
rows of orange-yellow spots, outlined with darker scales ; the first row at the base
of the wing, the second, in which the spots tend to fuse together, crossing the cell,
the third, irregular, at the outer end of the cell, the fourth, also irregular, the fifth,
parallel with the margin, the sixth consisting of a series of triangular submarginal
spots.

Fringes as on the upperside.

<$ GENITALIA. Uncus consisting of two slender, curved arms, slightly expanded
at the apices, which are thickly clad with strong spines ; tegumen fairly wide ;
special processes short and resembling those of aslauga ; vinculum wide ; valvae
of the usual shape with strongly falcate apices ; aedoeagus long, the distal end bearing
a pair of sharply-pointed triangular lobes on its dorsal surface and a deep excision
on the ventral face. This species is readily separable from B. barnesi by the absence
of the triangular apophyses at the base of the uncus arms.

$. Frons as in the male ; second joint of the palpi laterally compressed, clothed
with white scales and a scattering of dark ones, third joint almost wholly white ;
antennae and legs as in the male,

A REVISION OF THE GENUS TERIOMIMA KIRBY 89

UPPERSIDE FORE WING. Orange yellow with dark brown markings ; costal border
reduced, not invading the cell, with three yellow costal dots ; a small streak from
the costa covering the upper discocellular ; another from the costa to a little below
vein 6 ; apical patch and marginal border narrower than in the male, very reduced
towards the tornus.

UPPERSIDE HIND WING. The dark marginal border greatly reduced in width,
almost linear towards the anal angle, running inwards from the tips of the veins.

Fringes as in the male.

UNDERSIDE FORE WING. Orange yellow shading to pale yellow at the inner
margin, with violet grey markings ; costal border interrupted by four small yellow
dots ; four transverse streaks extending inward from the costa, the first at the base,
the second crossing the middle of the cell, the third over the discocellulars, and the
fourth curving and reaching vein 3 ; in the apical zone an oblique streak, with two
small costal yellow dots, extending to vein 4 ; a band of spots, confluent, from the
apex to vein 6, then triangular and separated from vein 5 to vein 3 ; a shining
grey marginal line, wide at the apex, tapering to the tornus.

UNDERSIDE HIND WING. As in the male, but the violet grey ground-colour a
little lighter and the orange spots larger.

Fringes as on the upperside.

Length of fore wing : $, 15-8 mm.
?, 15 mm.

Holotype <$ : Mlanje, Nyasaland, 16.^.1913, (S. A. Neave), in the British
Museum (N.H.), B.M. Type No. Rh. 15689.

Allotype $ : Mlanje, Nyasaland, 21 .iv. 1913, (S. A. Neave), in the British Museum
(N.H.), B.M. Type No. Rh. 15690.

Paratypes : A long series of both sexes from Mlanje ; Brit. Cent. Afr. ; Mandala,
2<? ; Port. E. Afr., Ruo Valley, 6^, i$ all in B.M. (N.H.) ; E. Afr.,
Monapo, 8$ 2$ in Oxford University Museum. Mozambique,
Tendo du Sunque, Gorongoza Dist., i<$ in coll. Stempffer.

This species exhibits considerable variation in facies, for instance, the example
from Mozambique, pi. 9, fig. 65, in which the genital armature is typical, is more
slender in build, has the ground-colour much paler on the upperside, with the dark
markings reduced and lighter in shade ; the underside hind wing has the ground-
colour of a very light violet-grey shade with the yellow spots reduced in number
and of paler tint.

(10) Baliochila hildegarda Kirby (1887)

(PI. 6, fig. 35, <$ genitalia (lateral), fig. 36, $ genitalia (ventral) ; PI. 9, figs. 55-60,

c and $ forms)

Teriomima (?) hildegarda Kirby, 1887, Ann. Mag. Nat. Hist. (5) 19 : 367, Ashanti. Figured in

Rhop. Ex. 1, Afr. Lye. pi. 4, figs. 7 and 8.
syn. Teriomima frey a Smith and Kirby, 1894, Rhop. Ex. 2 : 115, Afr. Lye. pi. 25, figs, i and 22.

Type in the British Museum (N.H.).

In the absence of abdomen and forelegs the sex of the holotype is debatable.
In facies it resembles a number of males from the Mombasa region, with reduced

90 A REVISION OF THE GENUS TERIOMIMA KIRBY

costal markings ; on the other hand, it may well be a female from almost any
locality within the range of distribution of the species. Fig. 55, PI. 9, shows a $ under-
side resembling that of the type specimen. The locality " Ashanti " is an obvious
error, as none of the species of this group have been recorded outside the Eastern
half of S. Africa. The species is very variable, with a striking form from the
coastal region, very heavily marked, the costal border almost flowing into the apical
patch and the hind- wing margins very wide and dark, (PI. 9, fig. 58.). Two females
are also shown in PL 9, figs. 56 and 57, illustrating upper and underside of a lightly
marked example with dark underside hind wing, figs. 59 and 60 a heavily marked
upperside with light underside hind wing.

$ GENITALIA. Uncus consisting of two short, broad arms, strongly toothed at
the apices and covered with small spines. In what is assumed to be the typical
form, from the Mombasa region, the two arms are splayed widely apart, but from
other regions the uncus shows a number of modifications in this respect, the opposite
extreme being reached in certain specimens from N.E. Rhodesia and British Central
Africa, wherein the uncus lobes tend to converge slightly at the tips. No subunci ;
tegumen wide ; special processes shorter than in aslauga crowned with strong
spines ; vinculum, anellus and valvae as in aslauga ; aedoeagus, long, excised
on the ventral face near tip. Distal half of valvae hairy.

DISTRIBUTION. Kenya, Tanganyika, Nyasaland, Rhodesia.

(n) Baliochila dubiosa sp. nov.

(PI. 3, fig. 8, holotype $ ; fig. 9, allotype $ ; PI. 7, fig. 37, holotype <$ genitalia ;
figs- 38, 39, 40, uncus variations ; fig. 41, $ genitalia ; PI. 9, figs. 61-64, c? forms)

Types in the British Museum (N.H.).

Assembled under this name are a number of forms, probably akin to B. hildegarda.
In facies they exhibit a considerable amount of variation, but all share a peculiar
formation of the uncus. This shape is not absolutely constant from specimen to
specimen but, however great the variation, the affinity of the dissected examples is
at once apparent. In view of this, the genitalic description is given first and that
of the facies later.

<$ GENITALIA. Uncus bifid, clothed with strong spines ; the two arms are deeply
cleft at their apices and each bears, at its base a small, lateral triangular apophysis,
as in aslauga and hildegarda ; tegumen, special processes, vinculum, valvae and
aedoeagus as in hildegarda. As previously stated, the uncus shape is rather variable,
so a number of characteristic forms are figured. The small lateral apophyses are
present in specimen no. 3052 from Kilifi, Kenya, fig. 38, but absent in specimen no.
3062, fig. 39 also from Kilifi, as well as in specimen no. 2637, fig. 40 from Meru.
Sometimes the two arms are asymmetrical, the" apical notch more or less deeply
cleft, and so on, but these variations do not appear to indicate racial distinctions.

c. Frons dark brown with two fine lateral white lines ; second joint of the palpi
laterally compressed, clothed with white and black scales, third joint white tipped ;
antennae black, ringed with white, club black ; legs black ringed with white.

A REVISION OF THE GENUS TERIOMIMA KIRBY 91

UPPERSIDE FORE WING. Yellow, with dark brown markings ; costal border
narrow, not extending into the cell, with three small costal yellow dots ; some black
scales at the base of the cell and in the centre ; a small streak covering the disco-
cellulars ; a curved bar running from the costa to vein 2 ; a large apical patch
continued down the border to the tornus, its inner edge forming an even curve.

UPPERSIDE HIND WING. Yellow, with a narrow dark brown marginal border, the
inner edge tending to run inward along the veins.

UNDERSIDE FORE WING. Orange-yellow shading to pale yellow towards the
inner margin, with markings of shining dark grey ; costal border interrupted by
four small costal white dots ; three transverse bars, one at the base of the cell,
one crossing the middle and extending beyond the origin of vein 2, the third covering
the discocellulars ; from the termination of the costal border, a broad curved bar,
rather irregular in outline, reaching vein 2 ; a shining grey apical zone with two
minute white costal spots, prolonged by a tapering marginal border to the tornus ;
between the veins the marginal border has a double row of small orange spots, de-
creasing in size towards the tornus.

UNDERSIDE HIND WING. Ground-colour violet grey with five transverse rows of
orange-yellow spots, outlined with darker scales, the first row at the base of the wing,
the second crossing the middle of the cell, the third, very irregular, at the end of
the cell, the fourth and fifth parallel with the margin ; between the veins a series of
submarginal orange-yellow spots.

Fringes of both wings shining grey.

It seems impossible to find any two specimens exactly alike, even those examples
shown by dissection to possess genitalia of identical pattern exhibit bewildering
variations of wing marking. Sometimes the dark markings of the fore wing upper-
side are reduced while the dark marginal border of the hind wing is developed, as
in no. 3052 from Kilifi. The costal border of the fore wing sometimes invades the
whole of the cell and so the yellow ground colour is reduced to less than half of
the wing surface ; the hind wing underside ground colour varies between shining
dark violet grey, almost black, as in no. 2637 from Meru, to a very light grey, as in
no. 3052 from Kilifi. Only by examination of the genital armature can this species
be determined with certainty. The specimens referred to above, together with two
other examples, are figured on pi. 9, figs. 61-64.

$. Frons, palpi, antennae and legs as in the male.

UPPERSIDE FORE WING. Orange yellow, rather deeper than in the male, with
dark brown markings ; costal border very narrow, with two, occasionally three or
four very small costal yellow spots ; a transverse patch running from the apex of
the costal margin to the origin of vein 7 ; apical patch and marginal border as in
the male.

UPPERSIDE HIND WING. Orange yellow, with a linear dark brown margin, some-
times a little broader and tending to run inwards along the veins.

Fringes of both wings interrupted with orange.

UNDERSIDE FORE WING. Orange-yellow shading to pale yellow towards the inner
margin ; four transverse shining violet grey streaks running from the costa, the
first at the base of the cell, the second crossing the middle, the third covering the

92 A REVISION OF THE GENUS TERIOMIMA KIRBY

discocellulars, the fourth reaching vein 4 ; in the apical zone an oblique streak
reaching vein 4, an anteterminal line from near the apex to vein 2, crenulate ;
marginal line wide at the apex, tapering to the tornus.

UNDERSIDE HIND WING. As in the male, ground-colour and markings varying
to a similar degree.

Length of fore wing : <$, 14 mm.

?, H'S mm.
Holotype <$ : Brit. E. Afr., Uchweni Forest, nr. Witu, 25-27.11.1912 (S. A.

Neave). B.M. Type No. Rh. 15691.
Allotype $ : Brit. E. Afr., Uchweni Forest, nr. Witu 25-27.^.1912 (S. A.

Neave). B.M. Type No. Rh. 15692.

Paratypes : Brit. E. Afr., Uchweni Forest nr. Witu, Feb. -Mar. 1912 (S. A.
Neave). 12^ 10$. All in British Museum (N.H.).

(12) Baliochila nyasae sp. nov.
(PI. 3, fig. 10, holotype $ ; PI. 4, fig. n, aUotype $ ; PL 7, fig. 42, < genitalia)

Types in the British Museum (N.H.).

cJ. Frons blackish grey with two lateral fine white lines ; second joint of the palpi
laterally compressed, clothed with black and white scales ; third joint white at
the base and tip ; antennae black, ringed white, club black, orange tipped ; tarsi
black, ringed white.

UPPERSIDE FORE WING. Orange yellow with dark brown markings ; costal border
running from the base to the extremity of vein n, where it nearly joins the apical
patch, leaving two small costal dots of the yellow ground-colour, one between
vein 12 and the costa, the other between veins n and 12 ; some dark scales in
the cell, one small streak covering the upper discocellulars, another streak extending
from the border to vein 4 ; a broad apical patch, with a wide marginal border reaching
the tornus, inner edge rather irregular.

UPPERSIDE HIND WING. Orange yellow with a dark brown marginal border about
3'5 mm. wide in the centre and running from the apex to the anal angle ; inner
edge rather irregular.

Fringes black, interrupted with white between the veins.

UNDERSIDE FORE WING. Ground-colour orange yellow, paler towards the inner
margin, with markings of lustrous dark violet grey ; costal border has five small
dots of the ground-colour, four evenly spaced along the border between the base
and the extremity of vein 12, the fifth between veins n and 12 ; from the costal
border runs a streak at the base of the cell, another at its middle and a third covers
the discocellulars ; a larger, curved marking runs from the costa to vein 3 ; in the
apical area an oblique line runs from the costa to vein 3 ; another, crenulate, lies
parallel with the marginal border from the apex to vein 2 ; a marginal line, of fairly
even width, extends from the apex to the tornus.

UNDERSIDE HIND WING. Ground-colour lustrous violet grey, with numerous
isolated orange-yellow spots arrange in five irregular bands, also a submarginal
series of interneural orange spots.

A REVISION OF THE GENUS TERIOMIMA KIRBY 93

Fringes as on upperside.

<$ GENITALIA. Uncus formed of a pair of slender arms fused together almost to
their extremities, leaving a notch at the distal end ; the suture between the arms
is clearly discerned under the microscope ; tegumen wide ; special processes short
with long fine spines at their ends ; valvae of the usual shape, with slightly falcate
apices ; aedoeagus rather short, thick, weakly excised below the tip on the ventral
face.

$. Frons, palpi, antennae and legs as in the male.

UPPERSIDE FORE WING. Orange yellow, rather paler than in the male, with dark
brown markings ; costal border narrower than in the male, leaving two or three
small yellow dots along the costa ; a small dark spot running from the costal border
and ending between veins 6 and 5 ; broad apical patch merging into a wide outer
margin which tapers to the tornus, its inner edge irregular.

UPPERSIDE HIND WING. Coloration as in the fore wing, with a narrower marginal
border than in the male, about 1-5 to 2 mm., with a very irregular inner edge.

Fringes as in the male.

UNDERSIDE FORE WING AND HIND WING. As in the male, but with the lustrous
dark violet grey of the fore wings slightly reduced and the ground-colour of the
hind wings rather lighter and duller greyish violet.

Fringes as on the upperside.

Length of fore wing : <$, 13 mm.

?, 15 mm -
Holotype $ : Mlanje, Nyasaland, 6.i.i3 (S. A. Neave). In British Museum

(N.H.), B.M. Type No. Rh. 15693.
Allotype $ : Mlanje, Nyasaland, 6.i.i3 (S. A. Neave). In British Museum

(N.H.), B.M. Type No. Rh. 15694.
Paratypes : Mlanje, Nyasaland, Nov.-April, n^ 10$, Ruo Valley, Nyasaland,

1$, all in the British Museum (N.H.).

(13) Baliochila stygia Talbot (1935)
(PL 4, fig. 12, neallotype $ ; PI. 7, fig. 43, $ genitalia)

Teriomima minima f. stygia Talbot, 1935, Ent. Mon. Mag. (3) 21 : 72, pi. i, fig. 4, 3, Kenya.
Rabai, 700', 14 m. N.W. of Mombasa (K. St. A. Rogers).

c. Holotype in the Hope Department of Entomology, University Museum,
Oxford.

$. Not previously described.

UPPERSIDE FORE WING. Orange yellow with dark brown markings ; wide costal
border covering most of the cell, leaving three small costal yellow dots ; a large
round spot flowing from the costal border and covering the discocellulars ; an ex-
tensive apical patch prolonged by a wide apical border, tapering to the tornus, the
inner edge of the apical patch and of the marginal border forming an even curve ;
some black scales scattered over the orange-yellow area.

94 A REVISION OF THE GENUS TERIOMIMA KIRBY

UPPERSIDE HIND WING. Colouring as in the fore wing, but with the marginal
border a little less wide, with an irregular and ill-defined inner edge.

Fringes of both wings black with some yellow hairs.

UNDERSIDE FORE WING. Ground-colour yellow, becoming paler towards the
inner margin, with dark grey markings as follows ; costal border leaving four small
costal white dots ; a spot at the base of the cell, one in its middle ; a streak covering
the discocellulars ; an oblique stripe curving from the costa to vein 3 ; a large apical
patch leaving two small costal white dots, divided by two series of confluent, in-
distinct orange yellow spots parallel with the outer margin ; marginal line wide at
the apex, tapering to the tornus.

UNDERSIDE HIND WING. Ground-colour shining violet grey with the usual six
transverse bars of orange-yellow spots, outlined with darker scales, the fourth row
from the base being more strongly accentuated than the rest ; marginal line shining
grey.

Fringes as on the upperside.

The general appearance of the female underside is lighter than that of the male ;
the hind wing appears almost unmarked unless closely inspected, owing to the
dull tinge of the markings.

Length of fore wing : $, 11-2 mm.

<$ GENITALIA. Uncus consisting of two pairs of short and slender points ; no
subunci ; tegumen very wide ; special processes stout, rather short, clothed with
bristles ; vinculum broad ; anellus sheathing the base of the aedoeagus ; valvae
oblong, with falcate pointed apices ; aedoeagus small, deeply excised ventrally near
the distal end.

Holotype $ : Mombasa, 14 m. N.W. of Rabai, 700', 2i.xii.og (St. A. Rogers).
Oxford Univ. Mus.

Neallotype $ : Zanzibar, 23.ix.24, #D.M. Cat or coll., in British Museum (N.H.),
B.M. Type No. Rh. 15695 ; in addition there are three more fe-
males bearing the same data in the British Museum (N.H.).

(14) Baliochila woodi Riley (1943)
(PI 7, fig. 44, <$ genitalia)

Teriomima woodi Riley, 1943, Entomologist, 76:225, Nyasaland, Mt. Mlanje, 12.11.38, 2500',
(R. C. Wood).

Types in the British Museum (N.H.).

$ GENITALIA. Uncus in the form of a flattened lobe, narrow at the base, ex-
panding towards the apex, where it is slightly excised and inclined ventrally ; no
subunci ; tegumen of moderate size ; special processes very long and slender,
clothed with short bristles ; vinculum wide ; valvae of the usual shape with slightly
falcate, slender and acute apices ; aedoeagus long, tapering evenly to a truncate
tip. Some hairs on the uncus, valvae hairy.

DISTRIBUTION. Nyasaland, Mlanje.

A REVISION OF THE GENUS TERIOMIMA KIRBY 95

(15) Baliochila fragilis sp. nov.
(PL 4, fig. 13, holotype <$ ; fig. 14, allotype $ ; PI. 7, fig. 45, < genitalia)

Types in the British Museum (N.H.).

cJ. Frons blackish brown with two lateral fine white lines ; second joint of the
palpi laterally compressed, clothed with white scales with a scattering of black
ones ; third joint white tipped ; antennae black, ringed with white, club black,
orange tipped ; tarsi black, ringed with white.

UPPERSIDE FORE WING. Orange yellow with dark brown markings, ground-colour
paler in some examples ; some dark scales at the base of the wing ; a small dark
streak over the discocellulars, absent in some specimens ; a dark oblique marking
extending from the costa, leaving a small yellow dot between the costa and vein
n, to vein 4, sometimes 5 ; apical patch with an irregular inner edge, prolonged
by a broad marginal border ending at the extremity of vein 3, but continued in the
form of small interneural patches to the tornus.

UPPERSIDE HIND WING. Ground-colour orange yellow, sometimes pale yellow,
with a dark marginal border, linear in most examples, occasionally as much as one-
and-a-half millimetres in width, with an irregular inner edge.

Fringes of both wings dark brown, interrupted with white on the hind wings.

UNDERSIDE FORE WING. Orange yellow, paler towards the inner margin, with
dark brown markings as follows : costal border with four, sometimes three, small
yellow dots ; three small bars crossing the cell at its base, centre and over the dis-
cocellulars respectively ; an oblique, curved streak running from the costa to vein 3,
sometimes 4 ; apical patch prolonged by a marginal border tapering to the tornus ;
two indistinct rows of interneural orange spots running parallel with the outer
margin, the inner row very small, in some examples obsolete.

UNDERSIDE HIND WING. Ground-colour greyish brown, sometimes lighter, some-
times darker, with six rows of small, indistinct orange-yellow spots, the fourth
row from the base being the most prominent, the fifth and sixth parallel with
the margin. In some specimens the spots can only be seen with the aid of a
microscope.

Fringes brown, interrupted with white.

c GENITALIA. Uncus in the form of a flattened oval disc, deeply excised at the
apex, supported on a longish stalk or neck, and bearing a number of short, sturdy
spines ; tegumen triangular ; special processes very long and slender, almost straight,
clothed with long, fine spines over their whole length ; vinculum wide ; valvae of
the usual shape, distal ends narrow, strongly falcate ; aedoeagus short and thick,
distal end truncate. In facies the males are somewhat variable, but their genitalia
are absolutely constant.

?. Frons, palpi, antennae and tarsi as in the male.

UPPERSIDE FORE WING. Yellow with dark brown markings ; costal border
narrow, with two small costal yellow dots ; an oblique spot from the costa to vein 5 ;
dark apical spot and marginal border as in the male.

UPPERSIDE HIND WING. Yellow, with a linear dark marginal border from the
costa to the anal angle.

9 6 A REVISION OF THE GENUS TERIOMIMA KIRBY

Fringes blackish brown interrupted with white near the apex of the fore wings
and the hind wings.

UNDERSIDE FORE WING. Yellow, paler towards the inner margin, with dark
brown markings as follows : three small yellow dots spaced along the costa, some
dark scales in the cell near its base, at its centre and over the disocellulars ; a dark
oblique spot from the costa to vein 4 ; apical patch with an irregular inner edge,
with the marginal border ending between veins 2 and 3 ; two rows of orange yellow
spots running parallel with the outer margin, rather more clearly marked than in
the male.

UNDERSIDE HIND WING. Ground colour slightly darker than in the male, the
transverse rows of orange spots paler in colour and appearing more distinct than
in the male.

Fringes as on the upperside.
Length of fore wing : $, 12 mm.

?, 13-6 mm.
Holotype $ : Kenya, Meru, 6000', May, 1946 (T. H. E. Jackson), in the British

Museum (N.H.), B.M. Type No. Rh. 15696.
Allotype $ : Kenya, Meru, 6000', May, 1946 (T. H. E. Jackson), in the British

Museum (N.H.), B.M. Type No. Rh. 15697.

Paratypes : Kenya, Nairobi, 7^ 2$ ; Kibwezi, 3000', 9^ i$ ; Chuka, 5600',
loj 2$ ; Makindu, 3300', i<$ ; Masongaleni, 3000', 1$ ; Gheni,
Ukamba, i<$ ; Meru, 56oo'-6ooo', 3$ i$ ; Samburu, i<$ ;
Muani, i$ ; Hills of Teita, i$ ; Mto-ya-mawe, i$ ; Kedai,
i(2 ; Somaliland, i<$ i$, all in British Museum (N.H.) ; Nairobi,
i(2, in Univ. Mus. Oxford. Meru, i<$ 2$ in coll. Jackson.
Kibwezi, 5(2 ; Bura Hill, 1$, in coll. van Someren. Meru,
2(2 i? ; Kima (Kenya), 1$, in coll. Stempffer.

(16) Baliochila minima minima Hawker Smith (1933)
(PI. 4, fig. 15, neallotype $ ; PI. 7, fig. 46, <$ genitalia)

Teriomima minima Hawker Smith, 1933, Stylops, 2 : 6, Kenya, Uchweni Forest, Witu, Feb.
1912 (S. A. Neave).

Holotype $ in the British Museum (N.H.).

$ not previously described.

UPPERSIDE FORE WING. Yellow, tinged with orange, markings blackish brown ;
some dark scales at the base of the costa and of the cell ; a small costal spot at the
extremity of vein 12, another extending obliquely from the costa to vein 5 ; apical
patch prolonged by the marginal border, which tapers to the tornus, the inner edge
irregular.

UPPERSIDE HIND WING. Marginal border linear, with an ill-defined and irregular
inner edge.

Fringes dark brown, interrupted with yellow.

UNDERSIDE FORE WING. Yellow, with silvery-grey markings ; costal border
with three small dots ; a spot at the base of the cell, one in the middle and a streak

A REVISION OF THE GENUS TERIOMIMA KIRBY 97

over the discocellulars ; a curved streak from the costa to vein 3 ; a streak parallel
with the outer margin from near the apex to vein 2 ; a marginal line wide at the
apex and tapering to the tornus.

UNDERSIDE HIND WING. Yellow, with five transverse silvery-grey stripes, the
first near the base, the second crossing the cell, the third irregular, the fourth and
fifth parallel with the marginal border ; marginal line silvery grey.

Fringes of fore wing dark brown interrupted with yellow, of hind wing yellow
towards the apex, mixed with brown towards the anal angle.

Length of fore wing : $, 10-1 mm.

Neallotype - : Kenya, Uchweni Forest, nr. Witu, 25-27. xi. 1912 (S. A. Neave),
in British Museum (N.H.). B.M. Type No. Rh. 15698.

<$ GENITALIA. Uncus forked from an oval base, the two prongs slender and
sinuate ; no subunci ; tegumen rather small ; special processes long and slender,
clothed with stiff bristles ; vinculum wide ; anellus sheathing the base of the aedo-
eagus ; valvae oblong, with falcate apices ; aedoeagus short and thick, with a cleft
tip.

DISTRIBUTION. Kenya, Uchweni Forest, near Witu.

(17) Baliochila minima latimarginata Hawker Smith (1933)

(PI. 4, fig. 16, neallotype $ ; PI. 7, fig. 47, genitalia (ventral) ; PL 8, fig. 48.

<$ genitalia (lateral))

Teriomima minima latimarginata Hawker Smith, 1933, Stylops, 2 : 7, Kenya, Rabai, Mombasa,
icJ, 1906 (K. St. A. Rogers).

Holotype $ in the British Museum (N.H.). $ not previously described.

<$ GENITALIA. Closely resembles m. minima save in the relative length of the
special processes and the lobes of the uncus. Whereas in m. minima all four tips
appear of equal length, in latimarginata the tips of the special processes project
well beyond those of the uncus.

$. The description given of the female of m. minima would answer equally well
for the form now considered, but there are some examples in which the marginal
borders of the upperside hind wings are nearly as broad as in the male.

Length of fore wing : $, 9-5 mm.

Neallotype $: B.E.A., Rabai, nr. Mombasa, 15.^.1912 (S. A. Neave) in the
British Museum (N.H.).

DISTRIBUTION. Rabai, Mombasa ; Moa, Tanganyika Terr. ; B.E.A., nr. Wangi ;
Jilore, 19 miles W. of Malindi ; Shimba Hills ; Kwale ; Kilifi.

(18) Baliochila minima amanica ssp. nov.

(PI. 4, fig. 17, holotype $ ; fig. 18, allotype $ ; PI. 8, fig. 49, uncus and special

processes)

Types in the British Museum (N.H.).

<$. UPPERSIDE FORE WING. Orange yellow with blackish-brown markings ;
costal border not invading the cell, bearing four small costal yellow dots ; some dark

9 8 A REVISION OF THE GENUS TERIOMIMA KIRBY

scales in the base of the cell, in the middle and over the discocellulars ; running
from the costal border the large apical patch is prolonged by a marginal border
tapering to the tornus, the inner edge of apical patch and marginal border irregular.

UPPERSIDE HIND WING. Orange yellow with the marginal border reduced to a
series of isolated spots at the extremities of veins 4 to ib.

Fringes of fore wings black interrupted with orange yellow, of hind wings orange
yellow with black at the extremities of the veins.

UNDERSIDE FORE WING. Orange yellow shading to pale yellow towards the inner
margin, with lustrous black markings as follows : six transverse stripes running
from the costa, the first crossing the base of the cell, the second its middle, the third
covering the discocellulars, the fourth reaching vein 4 , the fifth, oblique, also reaching
vein 4, the sixth, linear, from near the apex to vein 3 ; a black marginal line near
the tips of the wings and near the tornus.

UNDERSIDE HIND WING. Orange yellow with five transverse stripes of lustrous
blackish tint, the first from vein 8 to la, crossing the base of the cell ; the second
from the costal to the abdominal border, crossing the middle of the cell ; the third
very irregular, also crossing the whole wing and covering the discocellulars ; the
fourth and fifth parallel with the outer margin ; black marginal spots at the end
of the veins. The general appearance of the underside gives an impression of vivid
contrast between ground-colour and markings.

Fringes as on the upperside.

$ GENITALIA. Differ from those of minima minima only in the relative length
of the special processes, which are short and thick, projecting considerably less than
the tips of the uncus.

From 100 miles N.W. of Mombasa, Sagalla Mtn., 3500', a series of males and females
have been collected of a race identical in genitalic structure but differing in facies.
In the males the marginal border of the upperside hind wing is more developed,
irregular but continuous from the end of vein 6 to the anal angle ; the underside
markings are of much lighter tint, shining grey. The general aspect is much less
contrasted. In the female the ground colour is lighter than in the male, the costal
border reduced to small costal black dots, apical patch as in the male, no marginal
border to the hind wings ; fringes as in the male. The underside resembles that
of the male ; fringes of fore wings dark brown interrupted with orange yellow, of
hind wings ochre yellow.

Length of fore wing : <, 12 mm.
?, 12 mm.

Holotype $ : Tanganyika Terr., Amani, May-Sept., 1945 (native collector),
T. H. E. Jackson. In British Museum (N.H.) B.M. Type No.
Rh. 15700.

Allotype $ : Brit. E. Afr., Sagalla, 3500', c. 100 miles N.W. of Mombasa (St. A.
Rogers). In British Museum (N.H.), B.M. Type No. Rh. 15701.

Paratypes : Hills of Teita, Kilimanjaro, 2$ ; Amani, 4$ ; Sagalla, i^, all in
B.M. (N.H.) ; Sagalla, 11$ 4$ ; Amani, 3^ ; Arabuka Forest,
i^, all in coll. Oxford Univ. Mus. Sagalla, I ; Amani, i$ ;
Kasigau, i$ in coll. van Someren. Amani, 2$ 2$; Turiani,
Tanganyika Terr., i$ in coll. Jackson.

A REVISION OF THE GENUS TERIOMIMA KIRBY 99

(19) Baliochila lipara sp. nov.
(PI. 4, fig. 19, holotype $ ; fig. 20, allotype $ ; PI. 8, fig. 50, $ genitalia)

Types in the British Museum (N.H.).

c. Frons blackish grey with two fine lateral white lines ; second joint of the palpi
laterally compressed, clothed with brown and white scales, third joint white at the
base and the tip ; antennae black, ringed with white, club black, orange tipped ;
tarsi brown, ringed with white.

UPPERSIDE FORE WING. Orange yellow, of a deeper tint than in the associated
species, with dark brown markings ; costal border narrow, not invading the cell,
two small orange costal dots ; a small rectangular spot from the costa to vein 7,
almost running into the apical patch ; the apex broadly dark brown, marginal
border tapering to a line at the tornus, its inner edge irregular.

UPPERSIDE HIND WING. Deep orange yellow, the marginal border indistinct,
almost linear, from the costa to the anal angle. In an example from Umtali (coll.
Stempffer) the marginal border is of a reddish tinge.

Fringes of both wings dark brown, interrupted with white.

UNDERSIDE FORE WING. Orange yellow, paler towards the inner margin, with
shining dark violet-grey markings ; costal border with four small yellow dots ;
running from the costal border are four patches, one over the base of the cell, one
over the middle, one covering the discocellulars, the fourth larger, curved, reaching
vein 4 ; apical patch broad, with two yellowish-white costal spots ; marginal border
tapering to the tornus, its inner edge rather irregular ; a series of three oblique
orange streaks ; a submarginal series of interneural orange spots.

UNDERSIDE HIND WING. Ground-colour is the same as the markings of the fore
wing underside, with six transverse rows of small orange spots, the first row at the
base of the wing, the second crossing the cell, the third broken and indistinct, the
fourth and fifth parallel with the outer margin, the sixth submarginal ; the. fourth
row has the outer edges of its spots outlined with darker scales, and forms an angled
line crossing the wing from the costa to the abdominal margin. In some specimens
the orange markings are greatly reduced and the general impression is then of a
uniformly marbled grey.

Fringes as on the upperside.

c GENITALIA. Uncus formed of two small lobes with blunt apices ; tegumen wide ;
special processes highly developed as in minima, densely covered with long, fine
spines ; vinculum wide ; valvae oblong, with falcate apices ; aedoeagus long, its
distal end deeply notched, the dorsal lobe cut obliquely, the ventral lobe curved and
slightly expanded at the tip.

The genitalia plainly indicate the affinity of this species with minima, despite
the discrepancy in size.

$. Frons, palpi, antennae and tarsi as in the male.

UPPERSIDE FORE WING. Orange yellow, paler than in the male, with similar dark
brown markings.

UPPERSIDE HIND WING. As in the male. The Umtali example in the Stempffer
coll. has the marginal border even more reddish than in the male from that locality.

ioo A REVISION OF THE GENUS TERIOMIMA KIRBY

Fringes of fore wings dark brown interrupted with white, of hind wings whitish
towards the apex, dark brown towards the anal angle.

UNDERSIDE FORE WING. Orange yellow, paler towards the inner margin with
lustrous dark violet-grey markings ; costal border with four small yellow dots ;
four transverse markings as in the male ; apical zone not so dark as in the male, the
orange spots being larger and the marginal border ending at vein 3.

UNDERSIDE HIND WING. Ground-colour a little lighter than in the male, with the
orange spots larger ; the transverse, angled line, no. 4 from the base, quite indis-
tinct.

Fringes of both wings brown, interrupted with white.
Length of fore wing : $, 15 mm.
,, ,, $, 14 mm.

Holotype <$ : Nyasaland, Mlanje, 31 .xii. 1912 (S. A. Neave) in the British Museum

(N.H.), B.M. Type No. Rh. 15702.
Allotype $ : Nyasaland, Mlanje, 20. xi. 1912 (S. A. Neave) in the British Museum

(N.H.), B.M. Type No. Rh. 15703.

Paratypes : Nyasaland, Mlanje, 3^ 2$ ; Zomba, 2<$ ; Plains of Lake Chilwa,
1$ ; Kenya, Rabai, i<3 2$ ; Mashonaland, Umtali, 2$ ; all in
the British Museum (N.H.) ; Kenya, Rabai, i^ 2$ in Oxford
Univ. Mus.; Kenya, Sagalla, 2$ i$ ; Kasigau, i$ ; Kibwezi,
i$ in coll. van Someren. Kenya, Mombasa, i$ in coll. Jackson.
S. Rhodesia, Umtali, ig i$ in coll. Stempffer.

(20) Baliochila singularis sp. nov.
(PL 4, fig. 21, holotype $ ; PL 5, fig. 22, allotype $ ; PL 8, fig. 51, $ genitalia)

Types in the British Museum (N.H.)

(So named because of its isolated position in the genus.)

c. Frons blackish grey with two fine lateral white lines ; second joint of the palpi
laterally compressed, clothed with brown and white scales ; third joint dark brown,
white tipped ; antennae black, ringed with white, club black with a few white
scales ; tarsi brownish black, ringed with white.

UPPERSIDE FORE WING. Pale orange yellow with light brown markings as follows :
An oblique streak running from the costa to vein 4, well separated from the apical
patch ; apical patch smaller than in the related species, triangular, terminating
at the extremity of vein 4 ; no marginal border between 4 and the tornus.

UPPERSIDE HIND WING. Pale orange yellow without markings.

Fringes of fore wing brown, interrupted with white towards the apex, of hind wing
dirty white.

UNDERSIDE FORE WING. Pale yellow, orange tinged towards the costa ; markings
as on the upperside ; apical patch very pale brown.

UNDERSIDE HIND WING. Ground-colour pale brown, with a sprinkling of yellow
scales ; the only marking, scarcely visible, is a transverse, angled, darker line from
the costa to the abdominal border.

A REVISION OF THE GENUS TERIOMIMA KIRBY 101

Fringes of fore wing light brown, of hind wings light brown at the base, degraded
white in the distal half.

c? GENITALIA. Uncus consisting of two curved lobes mounted upon a stalk or
neck, the latter exceeding the former in length ; tegumen triangular ; no special
processes ; vinculum wide ; valvae of the usual shape, with strongly falcate apices ;
aedoeagus long, subcylindrical, with a pointed, ventrally directed distal end ; anellus
lobes highly developed, long, slender and excurved at the apices, which are heavily
spinous. It is considered that the formation of this armature indicates that singu-
laris is a link between the preceding species and those of the genus following. The
special processes are lacking, but the anellus lobes are highly developed. The uncus,
however, exhibits no fusion with the eighth tergite.

$. Frons, palpi, antennae and tarsi as in the male.

UPPERSIDE FORE WING. Ground-colour pale yellow, some light brown scales
at the base of the costa ; an oblique streak running from the costa to vein 4, light
brown from the costa to vein 10, darker from vein 10 to vein 4 ; apical patch as in
the male.

UPPERSIDE HIND WING. Pale yellow, no markings.

Fringes of both wings pale brown, interrupted with white.

UNDERSIDE FORE WING. As in the male.

UNDERSIDE HIND WING. Ground-colour lighter than in the male, markings similar.

Fringes as on the upperside.

Length of fore wing : <$, 14 mm.
?, 15-8 mm.

Holotype <$ : S. Africa, Durban, I5.ix.23, ex coll. W. Feather, in British Museum
(N.H.), B.M. Type No. Rh. 15704.

Allotype $ : S. Africa, Durban, I5.ix.23, ex coll. W. Feather, in British Museum
(N.H.), B.M. Type No. Rh. 15705.

Paratypes : S. Rhodesia, Umtali, 3.viii.27 (ex coll. Feather) ij ; Matabele-
land, Filabusi, 2.ix.23, i<$, all in British Museum (N.H.) ;
Durban, i<$ in coll. T. H. E. Jackson.

Genus CNODONTES gen. nov.

Type of the genus : Durbania pallida Trimen, 1898.

Eyes, palpi, antennae, legs,wingshape and neuration as in Teriomima and Baliochila.

$ GENITALIA. Exhibit a peculiarity of structure unknown to the authors in any
other genus of African or Holarctic Lycaenidae. The uncus is fused to the eighth
tergite, which forms a sort of guard or sheath. This feature suggested the name of
the genus. There are no subunci ; tegumen triangular with two long, digitate
processes springing from its base, these processes in no way connected with the
anellus ; vinculum wide ; valvae as in the preceding genus ; aedoeagus long ; a
pair of long, slender arms arising from the dorsal surface of the anellus and resting
parallel with the two processes from the base of the tegumen ; uncus and valvae
pilose. It seems that, in this genus, the " special processes " of Baliochila have
divided into two independent parts, one remaining attached to the tegumen, the
other to the anellus.

102 A REVISION OF THE GENUS TERIOMIMA KIRBY

(21) Cnodontes pallida Trimen (1898)
(PL 5, fig. 23, neallotype $ ; pi. 8, fig. 52, <$ genitalia)

Durbania pallida Trimen, 1898, Trans. Ent. Soc., 1898 : 12, pi. i, fig. 7, <j, fig. 8, ?. Mashona-
land, Gadzima, Umfuli R. (G. A. K. Marshall).

Types in the British Museum (N.H.).

The specimen described by Trimen as the female of this species is, in fact, another
male, the genitalia identical with those of the holotype. This error well illustrates
the danger of determination on superficial characters in this group. Very few ex-
amples of this species are available, not a single female from Mashonaland among
them. Having a male and a female from N. Bechuanaland, Ghanzi, Mongalatsila,
both taken by the same collector on the same day (i8.iii.i925, J. Maurice), and
bearing a strong superficial resemblance to one another, it is considered reasonable
to describe this female as the neallotype, the male having proved a true pallida
on dissection.

UPPERSIDE FORE WING. Ground-colour rich orange yellow, markings dark brown ;
costal band not invading the cell, except for a few dark scales on the discocellulars ;
three small costal yellow dots ; a patch of darker brown running from the costa
to vein 4 ; apical patch continued as a tapering marginal border to the tornus,
inner edge broken.

UPPERSIDE HIND WING. Ground-colour as on fore wing ; an indistinct, linear
marginal border from the apex to the anal angle.

Fringes of fore wing brownish, interrupted with white near the apex ; of hind
wings white interrupted with brown from the apex to vein 4, then brown to the anal
angle.

UNDERSIDE FORE WING. Deep orange yellow at the costal margin, paler towards
the inner margin ; costal and apical bands pale brown, the costal with four small
yellowish dots and a broken line of white scales on the anterior margin ; in the
cell are three evenly spaced dark brown patches ; a strongly defined dark patch
from the costa to vein 4 ; running through the apical patch and parallel with the
outer margin are two rows of weakly defined orange spots, diminishing in size from
the apex towards the tornus and merging into the ground-colour between veins
4 and 3.

UNDERSIDE HIND WING. Ground-colour pale fuscous, with six irregular transverse
rows of orange-yellow spots, the fourth from the base being most conspicuous as
it is outlined with an irregular zone of darker scales.

Length of fore wing : 17 mm.

Neallotype?: N. Bechuanaland, Ghanzi, Mongalatsila, i8.iii.25 (J- Maurice),
in the British Museum (N.H.), B.M. Type No. Rh. 15706.

$ GENITALIA. Uncus a simple lobe with a rounded apex, fused to the eighth
tergite as stated in the general description ; no subunci ; processes of the tegumen
long and slender, with hooked apices ; vinculum wide ; valvae of the usual shape,
apices weakly falcate ; aedoeagus long, subcylindrical, distal end pointed ; anellus
lobes very long and slender, slightly spatulate at the apices. Uncus, tegumenal
processes and valvae hairy.

A REVISION OF THE GENUS TERIOMIMA KIRBY 103

DISTRIBUTION. Rhodesia, E. bank of the Loangwa River, N. of Zumbo; N.
Bechuanaland ; Port. E. Africa.

(22) Cnodontes vansomereni sp. nov.

(PI. 5, fig. 24, holotype <$ ; fig. 25, allotype $ ; PI. 8, fig. 53, <$ genitalia (ventral) ;
fig- 54> <? genitalia (lateral) ; PI. 9, fig. 66, $, Katanga)

Types in the British Museum (N.H.).

(J. Frons blackish grey with two lateral fine white lines ; second joint of the
palpi laterally compressed, clothed with white scales and a sprinkling of brown ones ;
third joint white at the base and the tip ; antennae black, ringed with white, club
black ; tarsi dark brown, ringed with white.

UPPERSIDE FORE WING. Pure yellow, not orange tinged, with dark brown mark-
ings ; some dark scales between the costa and vein 12, a dark oblique streak from
the costa to vein 4, leaving a small yellow dot between the costa and vein u, almost
running into the apical patch ; apical patch triangular, with a small yellow dot
between veins 7 and 8, inner edge of patch irregular ; marginal border greatly
reduced in width from the end of vein 4 to the tornus, where it is linear.

UPPERSIDE HIND WING. Pure yellow, no dark markings.

Fringes very dark brown, interrupted with white near the apex of the fore wings
and round the hind wings.

UNDERSIDE FORE WING. Yellow, paler towards the inner margin ; brown costal
border extending to vein n, with two small yellow spots, one between vein 12
and the costa, the other between veins n and 12 ; some brown scales in the cell
near the upper border, some over the discocellulars ; an oblique streak extending
from the costa almost to vein 4, brown from the costa to vein 6, darker brown from
6 to its lower edge, the outer edge of this streak almost confluent with the apical
patch ; apex and marginal border light reddish brown, the border tapering to a line
at the tornus, the inner edge of the apical patch and the border forming an even
curve ; a series of very indistinct orange spots between the veins and parallel with
the margin.

UNDERSIDE HIND WING. Ground-colour light reddish brown, varying to a light
yellowish grey, with nearly obsolete rows of yellow spots, the row most visible
being outlined with darker scales and reaching from the end of vein 8 on the costa
to the abdominal border, forming almost a right angle at the level of vein 5.

Fringes blackish brown.

A number of examples, notably a male from Katanga, pi. 9, fig. 66, look very
different from the typical form, being much paler both on upperside and underside,
but the genitalia establish their identity beyond doubt.

cJ GENITALIA. Uncus small, triangular, with a rounded apex ; fused to it is the
eighth tergite, which forms behind it a hood or guard ; the uncus is attached to
a tegumen of moderate size by a semi-transparent unsclerotised membrane ; from
the base of the tegumen arise two digitate lobes with rounded apices, covered with
long, fine hairs. These lobes are not connected to the anellus. Vinculum wide ;
valvae of the usual shape with serrated apices terminating in fine curved points ;

IO4

A REVISION OF THE GENUS TERIOMIMA KIRBY

aedoeagus long, distal end pointed and ventrally inclined ; anellus lobes long and
slender, apices slightly spatulate and curving towards the special processes.

$. Frons, palpi, antennae and tarsi as in the male.

UPPERSIDE FORE WING. Lighter than in the male, similarly marked.

UPPERSIDE HIND WING. Without markings.

UNDERSIDE FORE WING. Yellow, paler towards the inner margin, markings as
in the male.

UNDERSIDE HIND WING. Light reddish brown in the allotype, but varying as in
the male, with an almost invisible pattern of orange-yellow spots.

Fringes brown with a few whitish interruptions.

Length of fore wing : <^, 15 mm.
?, 15-8 mm.

Holotype $ : Migori Valley, 4200', S. Kavirondo, Kenya (T. H. E. Jackson),
in British Museum (N.H.), B.M. Type No. Rh. 15707.

Allotype $ : Migori Valley, 4200', S. Kavirondo, Kenya (T. H. E. Jackson),
in British Museum, (N.H.), B.M. Type No. Rh. 15708.

Paratypes : Migori Valley, 4200', 2$ 2$ ; Nyasaland, Shire Valley, 3^ i$ ;
Tanganyika Terr., District of the Great Craters, 2$ ; N. Rho-
desia, Luangwa Valley, 3OOo'-i8oo', 4^ i$ ; N. Rhodesia,
Chambezi Valley, 4500', 2$ (T. A. Barnes), ig (S. A. Neave) ;
Belgian Congo, Kasenga, 3$ 2$ ; Katanga, i^ i$ ; Lualaba
River, i$ ; Angola, Kalweba River, 1$ ; all in British Museum
(N.H.). E. Afr., Kongwa, c. 3000', 3$ ; St. Michael's Mission,
ic? ; Itigi, c. 150 m. E. of Tabora, i<$ ; N.E. Rhodesia, Loangwa
Valley, 24^ 9$ ; Chambezi Valley, Chinsali Dist. and Kasama
Dist., 7(J 2$ ; all in Oxford Univ. Mus. Hope Fountain, 2$ in
coll. Jackson. Kenya, Suna, 1$; Belgian Congo, Katanga,
Lupweshi River, i<J i$ ; in coll. Stempffer. S. Kavirondo,
Suna, 5(J 3$ ; in coll. van Someren.

4. CONCLUSIONS

While the Teriomima sensu stricto have typical Liptenine genitalic armatures,
the two new genera, with their special developments, may well belong to a different
group. Whether they have a common origin, or constitute an outstanding example
of convergence of external characters, is debatable. The existence of B. singularis
rather suggests that the former is the correct assumption, as this species has the
uncus formation of Baliochila, the anellus lobes of Cnodontes, but no "special
processes." The wealth of new forms from Mlanje, Nyasaland, revealed by the
intensive collecting of Dr. S. A. Neave in 1912, 1913 and 1914, suggests that similar
intensive methods, employed in other localties favourable to this group, might well
be rewarded by the discovery of many more.

PRESENTED

Bull. B. M. (N.H.) Entom. Ill, 3

PLATE 3

<

>

\ /

PLATE 3

Uppersides (left) and undersides (right) of the following : (i) Teriomima subpunctata Kirby,
neallotype <$. (2) Teriomima puella Kirby, neallotype ?. (3) Teriomima pdrva Hawker Smith,
neallotype ?. (4) Baliochila barnesi gen. et sp. nov., holotype <$. (5) Baliochila barnesi allo-
type $. (6) Baliochila neavei sp. nov., holotype (J. (7) Baliochila neavei allotype ?. (8)
Baliochila dubiosa sp. nov., holotype c?. (9) Baliochila dubiosa, allotype $. (10) Baliochila
nyasae sp. nov., holotype <$.

Bull. B. M. (N.H.) Entom. Ill, 3

PLATE 4

21

PLATE 4

(n) Baliochila nyasae, allotype $. (12) Baliochila stygia Talbot, neallotype ?. (13) Balio-
chila fragilis sp. nov., holotype <$. (14) Baliochila fragilis, allotype $. (15) Baliochila minima
minima Hawker Smith, neallotype ?. (16) Baliochila minima latimarginata Hawker Smith,
neallotype ?. (17) Baliochila minima amanica ssp. nov., holotype <$. (18) Baliochila minima
amanica, allotype $. (19) Baliochila lipara sp. nov., holotype <$. (20) Baliochila lipara,
allotype ?. (21) Baliochila singularis sp. nov., holotype ^.

Bull. B. M. (N.H.} Entom. Ill, 3

PLATE 5

29

PLATE 5

(22) Baliochila singularis, allotype $. (23) Cnodontes (gen. nov.) pallida Trirnen, neallotype
?. (24) Cnodontes vansomereni sp. nov., holotype c?. (25) Cnodontes vansomereni allotype $.
(26) Teriomima subpunctata Kirby, <$ genitalia (aedoeagus detached). (27) Teriomima puella
Kirby, $ genitalia (aedoeagus detached). (28) Teriomima zuluana van Son, <$ genitalia (aedoe-
agus detached). (29) Teriomima puellaris Trinien, $ genitalia (uncus and aedoeagus detached).

Bull. 13. M. (N.H.) Entom. HI, 3

PLATE 6

PLATE 6

(30) Teriomima micra Grose Smith, <$ genitalia (aedoeagus detached). (31) Teriomima
parva Hawker Smith, <J genitalia (aedoeagus detached). (32) Baliochila (gen. nov.) aslauga
Trimen, $ genitalia (aedoeagus detached). (33) Baliochila barnesi sp. nov., <$ genitalia (aedoe-
agus detached). (34) Baliochila neavei sp. nov., <$ genitalia (aedoeagus detached). (35) Balio-
chila hildegarda Kirby, J genitalia (lateral aspect) . (36) Baliochila hildegarda Kirby, <J genitalia
(aedoeagus detached).

Bull. B. M. (N.H.) Entom. Ill, 3

PLATE 7

PLATE 7

(37) Baliochila dubiosa sp. nov., genitalia of holotype <J (aedoeagus detached). (38 B. dubiosa,
uncus (Kilifi, Kenya). (39) B. dubiosa, uncus (Kilifi, Kenya). (40) B, dubiosa, uncus (Meru,
Kenya). (41) B. dubiosa, <$ genitalia (aedoeagus detached). (42) Baliochila nyasae sp. nov.,
<? genitalia (aedoeagus detached). (43) Baliochila stygia Talbot, $ genitalia (aedoeagus deta-
ched). (44) Baliochila woodi Riley, <J genitalia. (45) Baliochila fragilis sp. nov., <J genitalia.
(46) Baliochila minima minima Hawker Smith, uncus and special processes. (47) Baliochila
minima latimarginata Hawker Smith, <y genitalia (valvae and aedoeagus detached).

Bull. B. M. (N.H.) Entom. Ill, 3

PLATE 8

PLATE 8

(48) Baliochila minima latimarginata Hawker Smith, genitalia (lateral aspect). (49) Balio-
chila minima amanica ssp. nov., uncus and special processes. (50) Baliochila lipara sp. nov.,
<$ genitalia (aedoeagus detached). (51) Baliochila singularis sp. nov., <$ genitalia. (52) Cno-
dontes (gen. nov.) pallida Trimen, <$ genitalia. (53) Cnodontes vansomereni sp. nov. <$ geni-
talia (aedoeagus detached). (54) Cnodontes vansomereni sp. nov., $ genitalia (lateral aspect).

Hull. B. M. (N.H.) Entom. Ill, .?

PLATE y

PLATE 9

(55) Baliochila hildegarda Kirby, Mombasa, <$ underside. (56) B. hildegarda Kirby, Mom-
basa, 9 upperside. (57) Underside of fig. 56. (58) B. hildegarda Kirby. Sagalla, B.E.A.,
cj underside (heavily marked form). (59) B. hildegarda Kirby, Fort Jamieson, N.E. Rhodesia,
$ upperside (heavily marked form). (60) Underside of fig. 59. (61) Baliochila dubiosa sp.
nov., nr. Mombasa, 30 m. N.W. of Rabai, $ underside. (62) B. dubiosa sp. nov., Kilifi, Kenya,
c? underside. (63) B. dubiosa sp. nov., Nairobi, Kenya, <$ underside. (64) B. dubiosa sp. nov.,
Meru, Tanganyika, <$ underside. (65) Baliochila neavei sp. nov., Gorongoza, Mozambique
cJ underside. (66) Cnodontes vansomereni gen. et sp. nov., Katanga, R. Lupweshi, <$ underside.

PRESENTED

25 NOV B53

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

17 DEC 1953 NOTES ON

BRITISH ICHNEUMONINAE
WITH DESCRIPTIONS OF NEW

SPECIES

(HYM., ICHNEUMONIDAE)

J. F. PERKINS

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 3 No. 4

LONDON: 1953

NOTES ON BRITISH ICHNEUMONINAE WITH
DESCRIPTIONS OF NEW SPECIES

(HYM, ICHNEUMONIDAE)

BY

J. F. PERKINS

>. 103-176 ; 41 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 3 No. 4

LONDON: 1953

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

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

This paper is Vol. 3, No. 4 of the Entomological
series.

PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM

Issued December, 1953. Price Fifteen Shillings.

NOTES ON BRITISH ICHNEUMONINAE WITH

DESCRIPTIONS OF NEW SPECIES

(HYM., ICHNEUMONIDAE)

By J. F. PERKINS

CONTENTS

INTRODUCTION ........... 105

I. NOTES ON THE LlNNEAN TYPES OF ICHNEUMONINAE . . . IO6
II. THE STEPHENS TYPES OF ICHNEUMONINAE .... 107

III. THE DESVIGNES TYPES OF ICHNEUMONINAE . . . .113

IV. TYPES OF BRITISH ICHNEUMONINAE IN THE BRIDGMAN, MARSHALL

AND MORLEY COLLECTIONS . . . . . . .115

(fl) The Bridgman types of Ichneumoninae . . . . .116
(6) The Morley types of British Ichneumoninae . . . . 116
(c) The Marshall type of Mesostenus maurus . . . . .117

V. THE L. A. CARR COLLECTION OF ICHNEUMONIDAE . . . 117

The types of Ichneumoninae in the L. A. Carr collection . . 118

VI. SPECIES TO BE DELETED FROM THE LIST OF BRITISH ICHNEUMONINAE 118

VII. SPECIES RECORDED IN THE ADDENDA OF MORLEY'S British Ichneu-
mons ........... 135

VIII. CHANGES IN THE TRIVIAL NAMES OF BRITISH ICHNEUMONINAE . 135

IX. CHANGES IN GENERIC PLACEMENT OF SPECIES OF BRITISH ICHNEU-
MONINAE .......... 136

X. ADDITIONS TO THE LIST OF BRITISH ICHNEUMONINAE. . . 137

XI. DESCRIPTIONS OF NEW SPECIES OF BRITISH ICHNEUMONINAE . 145

XII. CHECK LIST OF BRITISH ICHNEUMONINAE ..... 169

XIII. REFERENCES .......... 175

SYNOPSIS

The paper deals with the British species of Ichneumoninae to be added to, and deleted from
the list given by Kloet and Hincks in their Check List of British Insects. Two genera and
sixteen species are described as new and three new names are proposed. In all, no trivial
names are deleted from the list and 105 are added ; this includes changes due to direct synonymy.
Some new synonymy is included. Finally a new check list of British Ichneumoninae is given.

INTRODUCTION

THE present paper was written in order to clear the way for a handbook on the
British species of Ichneumoninae. I have therefore given almost no notes on the
characters for distinguishing the species which are added to the British list of Ich-

ENTOM. Ill, 4. 9

106 NOTES ON BRITISH ICHNEUMONIN AE

neumoninae as given in Kloet and Hincks (1945), as these will appear in the keys in
the handbook. I have, however, given some information on certain of the species
that I consider should be excluded from the British list, where these have a wider
interest.

I have been much aided in this work by the very kind co-operation of many
workers who have sent me material and given me information. I wish to thank
the Director of the Institut Royal des Sciences Naturelles de Belgique for allowing
me the opportunity of examining the Wesmael collection, and Mons. A. Collart for
the facilities he placed at my disposal ; the officials of the Linnean Society for
allowing me to examine certain Linnean types ; Mr. E. A. Ellis of the Castle Museum,
Norwich, for the loan of the Bridgman collection of Ichneumoninae ; the officers
of the Suffolk Naturalists who presented the Morley types of Ichneumonidae to
the British Museum and who have lent me the Morley collection of Ichneumonidae ;
Mr. A. A. Gumming of the City Museum and Art Gallery, Plymouth, for allowing me
to examine and borrow material from the Bignell collection ; Mr. H. C. S. Halton
of the Natural History Museum, Wollaton Hall, Nottingham, for facilities to examine,
and to borrow, types and material from the L. A. Carr collection of Ichneumonidae ;
Miss G. Roche of the National Museum of Ireland for the loan of specimens from the
Johnson collection ; Prof. C. H. Lindroth of the Lund University Museum for the
loan of specimens from the Thomson collection ; Mr. A. W. Stelfox for the loan of
his collection of Phaeogenini ; Dr. O. W. Richards for the loan of his collection of
Ichneumoninae ; Dr. W. D. Hincks for the loan of material of Platylabini ; and Mr.
G. J. Kerrich and Mr. G. Heinrich for information on various points, including
types.

Throughout the paper names which are given after an equals sign are the valid
names, while those given after syn. are synonyms.

References to original descriptions are not quoted in the text, but they can be
found by consulting Dalla Torre (1902), Morley (1903) and Schmiedeknecht (1902
and 1928-1932) ; references to species described after 1932 will be found in section
XIII at the end of the paper.

I. NOTES ON THE LINNEAN TYPES OF ICHNEUMONINAE

Roman (1932) has discussed the Linnean types of Ichneumonidae, but more
recent work has made it necessary to re-assess certain of them, and the results are
given below, in so far as they affect the British list of Ichneumoninae.

Ichneumon fossorius L., 1758

= Amblyteles fossorius (L., 1758).

syn. Amblyteles viridatorius (Gravenhorst, 1820). (syn. nov.)

Roman has synonymized fossorius with Amblyteles subsericans (Gravenhorst).
The type of fossorius is a male, and this can be distinguished at once from subsericans
by the ventral fold on the sternites. The Linnean specimen has a fold on sternites
2-4. I have seen no specimen of Amblyteles fossorius (L.) from the British Isles.

NOTES ON BRITISH ICHNEUMONIN AE 107

Ichneumon molitorius L., 1761.

Roman suggested that the Linnean species belonged to the form described as
crassifemur by Thomson. However, I regard crassifemur as being a species distinct
from molitorius L. Thomson, distinguished not only in the characters of the hind
leg given by Thomson, but also in having the groove on the underside of the middle
femur with only very sparse hairs. The Linnean specimen agrees with Thomson's
interpretation of molitorius. I have examined type material of crassifemur from the
Thomson collection.

Ichneumon comitator L., 1758.

= Coelichneumon comitator (L., 1758).

syn. Ichneumon restaurator F. Wesmael, 1844.

Ichneumon lineator F. Gravenhorst, 1820, nee F., 1781.
Ichneumon ferreus Gravenhorst, 1829.

I regard Ichneumon ferreus as the form of C. comitator with the femora and tibiae
red, but distinct from Coelichneumon purpurissatus (see p. 138).

II. THE STEPHENS TYPES OF ICHNEUMONINAE

I have re-examined the types of Stephens that had already been selected by
Claude Morley (see Morley, 1902), and also I have been able to recognize further
types which previously had remained unknown. I am therefore giving a complete
list of the species of this sub-family described by Stephens in 1835 in his Illustrations
of British Entomology : or a synopsis of Indigenous Insects : Mandibulata, 7 : 126-
207 and 269-273.

The following changes of nomenclature are necessary :

Coelichneumon nigerrimus (Stephens, 1835) for Coelichneumon derasus

(Wesmael, 1844).
Cratichneumon albifrons (Stephens, 1835) for Cratichneumon gravenhorstii

(Boyer de Fonscolombe, 1847).
Cratichneumon fabricator maculifrons (Stephens, 1835) for this British

sub-species.
Coelichneumon eximius (Stephens, 1835) for Coelichneumon coeruleus (Cres-

son, 1864).

The notes on the types of Ichneumon and Trogus are given in the order in which
the species are placed in Stephens' work.

Ichneumon fasciatus

Type Hym. 3b 1580 <$ selected by Morley.

= Hepiopelmus leucostigmus (Gravenhorst, 1820).

The same specimen is the type of Ichneumon maculiventris Desvignes.

io8 NOTES ON BRITISH ICHNEUMON IN AE

Ichneumon nigerrimus

Type Hym. 3b 1721 <$ selected by Morley.
= Coelichneumon nigerrimus (Stephens, 1835).

syn. Coelichneumon derasus (Wesmael, 1844). (syn. nov.)
This name is not now invalidated by Ichneumon nigerrimus (Scopoli).
(= Sphex nigerrimus Scopoli = Anoplius nigerrimus (Scopoli)).

Ichneumon compunctor

Type Hym. 3b 1814 ^ selected by Morley.
= Ichneumon cessator Mueller, 1776.

Ichneumon rufipes

Type Hym. 3b 1582 selected by Morley.

= Polytribax curvus (Schrank, 1802). (syn. nov.} (comb, nov.}

syn. Ichneumon curvus Schrank, 1802.
Cryptus rufipes Gravenhorst, 1829.
Phygadeuon curvus (Schrank) ; Gravenhorst, 1829.
Plectocryptus curvus (Schrank) ; Thomson, 1873 and 1874.
Microcyptus curvus (Schrank) ; Thomson, 1883.

Mesocryptus (lapsus for Microcryptus} curvus (Schrank) ; Thomson, 1897.
Plectocryptus curvus (Schrank) ; Schmiedeknecht, 1905.

Confusion has been caused in the interpretation of this species by the fact that
one specimen from Stephens' collection (not the type) has the head and thorax of
Polytribax curvus male, and on to this has been glued the abdomen of a female
Eurylabus torvus Wesmael. The species with oval spiracles which were placed in
Microcryptus by Thomson (1883) and the related species which were placed by
Schmiedeknecht in Plectocryptus are congeneric with the North American species of
Polytribax.

Ichneumon maculifrons

No type.

= Cratichneumon fabricator maculifrons (Stephens, 1835).

The specimen selected by Morley as the type has the hind femur red and thus does
not agree with the original description, but from the description this species is most
probably Cratichneumon fabricator (F.) as stated by Morley, and I propose to use
this name for the British subspecies of fabricator. The specimen which Morley
selected, however, does belong to the same subspecies.

Ichneumon maculicornis

Type Hym. 3b 1585 $ selected by Morley.
= Phaeogenes maculicornis (Stephens, 1835).
syn. Phaeogenes scutellaris Wesmael, 1844.

NOTES ON BRITISH ICHNEUMONIN AE 109

Ichneumon cognatus

No type. <$. Given as a synonym of Ambly teles subsericans (Gravenhorst, 1820)
by Desvignes (1856). Males of this species as well as of the closely related A.
elongatus Brischke are present in the Stephens collection.

Ichneumon crassicornis Stephens nee Rossi, 1794.

Type Hym. 3b 1725 $ selected by Morley.
= Probolus concinnus Wesmael, 1853.

Ichneumon femorator

Type Hym. 3b 1581 $ selected by Morley.
= Probolus culpatorius (L., 1758).

syn. Probolus alticola (Gravenhorst, 1820).

Ichneumon fulvipes

No. type. Given as a synonym of annulator F., 1793 (= culex Mueller, 1776) by
Desvignes (1856). This agrees with the description.

Ichneumon albifrons

Type Hym. 3b 1565 $ selected by Morley.
= Cratichneumon albifrons (Stephens, 1835).

syn. Ichneumon gravenhorstii Boyer de Fonscolombe, 1847, nee Wesmael, 1836, nee
Gue"rin-Men6ville,i838. (syn. nov.)

It is not conspecific with Ichneumon impugnator Wesmael, of which I have
examined the syntypes, which has a conspicuously longer malar space.

Ichneumon binotatus

No type.

= Stenichneumon lineator (F., 1781). (syn. nov.).

Given as a synonym of Ichneumon leucomelas Gmelin (= Barichneumon albilineatus
(Gravenhorst) by Desvignes, but the description cannot refer to that species. The
male of Stenichneumon lineator (F.) agrees with the description and I regard binotatus
as a synonym of this species.

Ichneumon bipunctorius

No type. Not given by Desvignes (1856). From the description = Barichneumon
albilineatus (Gravenhorst, 1820).

Ichneumon cingulipes

Type Hym. 3b 1576 $ selected by Morley.
= Amblyteles oratorius (F., 1793).

no NOTES ON BRITISH ICHNEUMONIN AE

Ichneumon microcephalus

Type Hym. 3b 1579 $ selected by Morley.
= Ichneumon formosus Gravenhorst, 1829. C s y n - n ov.}

I follow Wesmael, who examined the type of formosus, in the synonymy of 7.
obsessor Wesmael with that species.

Ichneumon fulvoscutellatus

No type. The description obviously refers to a discoloured specimen, and since
this was obtained in May, most probably to a species of Ichneumon that had hiber-
nated. I have been unable to determine the species from the description. Morley
has suggested that the name is a synonym of /. terminatorius Gravenhorst.

Ichneumon quadrinotatus

No type. The specimen which had been selected as the type by Morley is from
the Desvignes collection. However, there seems no doubt that the species is a
synonym of /. gracilicornis Gravenhorst, as given by Desvignes (1856).

Ichneumon concinnatorius

No type.

= Ichneumon terminatorius Gravenhorst, 1820.

Ichneumon dimidiatus

Type Hym. 3b 1574 $ selected by Morley.

= Amblyteles pallidicornis (Gravenhorst, 1829).

Ichneumon diver sorius

No type.

= Amblyteles armatorius (Forster, 1771). $.

Not quoted by Desvignes (1856).

Ichneumon triangulator

No type.

= Amblyteles trifasciatus (Gravenhorst, 1829) teste Morley.

Not given by Desvignes (1856).

Ichneumon eximius

Type Hym. 3b 1817 $ selected by Perkins.

= Coelichneumon eximius (Stephens, 1835).

syn. Coelichneumon coeruleus (Cresson, 1864). (syn. nov.)

There is a female of the North American Coelichneumon coeruleus from the Stephens
collection bearing a label in the writing of F. Smith saying that it stood in the
Stephens British collection. This specimen agrees entirely with Stephens' descrip-
tion and it thus seems certain that it is in fact the type of eximius.

NOTES ON BRITISH ICHNEUMONIN AE in

Ichneumon erythrogaster Stephens nee Gmelin, 1790.

Type Hym. 3b 1720 $ selected by Morley.
= Protichneumon coqueberti (Wesmael, 1848).

Ichneumon melanopyrrhus

Type Hym. 3b 1815 <$ selected by Perkins.

= Coelichneumon orbitator (Thunberg, 1822). (syn. nov.)
syn. Coelichneumon liocnemis (Thomson, 1888).

Morley has synonymized this species with Coelichneumon castaneiventris (Graven-
horst), but I have seen no specimens of that species from Britain with the hind
femur and tibia red, as is given in the original description of melanopyrrhus.

Except that Stephens does not mention the small white marks on the thorax,
there would be no doubt that the species here described was C. orbitator. However,
the single male of orbitator from the Stephens collection has the thorax covered in
dirt and these marks obscured and, moreover, has the apex of the 3rd tergite red
and the base of the 4th tergite black as given in Stephens' original description. I
have therefore no doubt that this was the specimen that Stephens was describing
and have selected it as the type.

Roman (1912) synonymized C. liocnemis (Thomson) with /. orbitator Thunberg ;
in 1914 he synonymized I. ruficauda Wesmael with this species, but ruficauda, of
which I have seen the type series, is completely distinct. As Thunberg states that
the legs are red, I have accepted Roman's synonymy of orbitator with liocnemis,
ruficauda having legs for the most part black in the specimens I have examined.

Ichneumon castanopyga

Type Hym. 3b 1577 $ selected by Morley.
= Amblyteles castanopyga (Stephens, 1835).
syn. Amblyteles rubriventris Wesmael, 1854.

Ichneumon pyrrhopus

No type.

= Cratichneumon fabricator maculifrons (Stephens, 1835). (syn. nov.)
This species has been synonymized with Cratichneumon fugitivus (Gravenhorst),
but from the description it is unlikely to be that species, as I have never seen a
specimen of fugitivus with red hind femora. However, the earlier authors mixed
this species with the forms of fabricator with a red abdomen, and it is most probable
that the Stephens species is a synonym of C. fabricator maculifrons (Stephens) .

Ichneumon gasterator

Type Hym. 3b 1567 $ selected by Morley.

= Cratichneumon corruscator (L., 1758).

In the Stephens collection there is also a specimen of C. fugitivus (Gravenhorst)

H2 NOTES ON BRITISH ICHNEUMONIN AE

labelled as gasterator ; this no doubt is the form mentioned by Stephens with " pos-
terior tibiae sometimes pitchy at apex, and reddish at the base."

Ichneumon femorator

Type Hym. 3b 1581 $ selected by Morley.
= Colpognathus celerator (Gravenhorst, 1807).

Ichneumon rufator

Type Hym. 3b 1698 $ selected by Morley.

= Phaeogenes semivulpinus (Gravenhorst, 1829). (syn. nov.)

Ichneumon abdominator

Type Hym. 3b 1816 $ selected by Perkins.

= Diadromus troglodytes (Gravenhorst, 1829). (syn. nov.)

The specimen was unlabelled as to species but had a label " New species " in
Smith's handwriting. It is a peculiarly coloured specimen of troglodytes agreeing
with Stephens' description.

Ichneumon picipes

No type. I have been unable to recognize this species from the description,
though most probably it is a female Phaeogenes.

Ichneumon ruficollis

No type.

= Barichneumon sanguinator (Rossi, 1794).

This synonymy was given by Desvignes (1856).

Ichneumon rufescens

Type Hym. 3b 1564 $ selected by Morley.
= Aoplus ratzeburgii (Hartig, 1838).

syn. Stenichneumon pictus (Gravenhorst) Morley.

Ichneumon cinctorius Stephens nee Fabricius 1775

Type Hym. 3b 1813 $ selected by Perkins.

= Ambly teles indocilis Wesmael, 1844.

This type was not previously indicated. Morley (1902) attributes the species
to Desvignes (who only re-described it), and selected a so-called type from the latter's
collection. The two specimens are, however, conspecific.

Trogus atrocaudatus

No type. From the description = Callajoppa cirrogastra (Schrank, 1781). I have
never seen a British specimen of C. exaltatoria (with which Morley synonymised it)
in which the 4th tergite is testaceous.

NOTES ON BRITISH ICHNEUMONIN AE 113

Trogus dissimilator

No type. From the description = Ichneumon didymus Gravenhorst, 1829 (syn.
bisignatus Gravenhorst), as suggested by Morley.

III. THE DESVIGNES TYPES OF ICHNEUMONIN AE

Morley (1902) has given some notes on the Desvignes types, and all those types,
which are in the British Museum, were selected by him. Heinrich (1937) has given
further information on them. However, there were various problems left outstand-
ing and it has now been possible to re-assess these. I have therefore included a
complete list of the Desvignes types. The following names are at present valid :

Platylabus obator (Desvignes, 1856).

syn. Ichneumon obator Desvignes, 1856.

Ichneumon lautatorius Desvignes, 1856.

Ichneumon minutorius Desvignes, 1856.

syn. /. captorius Thomson, 1887. (syn. nov.)

Chasmias paludator (Desvignes, 1854.)

syn. Chasmodes paludicola Wesmael, 1857.

In the list of types which follows all but the last two species were described by
Desvignes in 1856 in the Catalogue of the British Ichneumonidae in the British Museum.

Ichneumon maculiventris

Type Hym. 3b 1580, ?.

= Hepiopelmus leucostigmus (Gravenhorst, 1820).

This specimen is also the type of Ichneumon fasciatus Stephens.

Ichneumon obator

Type Hym. 3b 1583, <?.

= Platylabus obator (Desvignes, 1856).

This is a distinct species and not a synonym of P. pedatorius (F.). It differs from
Pedatorius in having the epicnemia strongly raised on each side of the middle line,
the distance between thyridiae : breadth of a thyridia = i : at most 1-5 ; stigma
black, hind femur black apically and segment I of hind tarsus sub-equal to segments
2 + 3+4- I have taken it abundantly in June, flying around Galium in hedgerows.

Ichneumon crassorius

Type Hym. 3b 1693, $.

= Ichneumon didymus Gravenhorst, 1829. (Heinrich, 1937).
/. inquinatus Wesmael (syn. brevigena Thomson) which Morley considered as
conspecific with /. crassorius, is a completely distinct species.

Ichneumon relucens

Type Hym. 3b 1573, ?.

= Amblyteles indocilis Wesmael, 1844.

n 4 NOTES ON BRITISH ICHNEUMONIN AE

Ichneumon cubicularis

Type Hym. 3b 1571, ?.

= Amblyteles fabricii (Schrank, 1802).

syn. Amblyteles truncicola Thomson, 1888. (syn. nov.)

Ichneumon lautatorius

I have examined the type in the Curtis collection.

Ichneumon ancipterus

Type Hym. 3b 1722, $.

= Amblyteles pattiatorius (Gravenhorst, 1829.) (syn. nov.)

Ichneumon dubitatus

Type Hym. 3b 1572, #.

= Amblyteles palliatorim (Gravenhorst, 1829). (syn. nov.)

Ichneumon flavocinctus

Type Hym. 3b 1570, <J.

= Ctenichneumon panzeri (Wesmael, 1844). (Heinrich, 1937).

Ichneumon minutorius

Type Hym. 3b 1702, ^.

syn. Ichneumon captorius Thomson, 1887.

The female of this species was recorded erroneously as rufidens Wesmael by
Desvignes. It is one of the species that was formerly lumped under " Ichneumon
raptorius." I have examined 9 female and 2 male syntypes of captorius from the
Thomson collection.

Ichneumon binotatus

Type Hym. 3b 1566, <?.

= Cratichneumon corruscator (L., 1758).

Ichneumon niveatus

Type Hym. 3b 1690, $.

= Aptesis arridens (Gravenhorst, 1829).

Ichneumon cinctorius

This species was described by Stephens (1835). It has been attributed to Des-
vignes by Morley (1902), but Desvignes only re-described it. The name is a primary
homonym and the species is Amblyteles indocilis Wesmael.

NOTES ON BRITISH ICHNEUMONIN AE 115

Ichneumon rubedinis

Type Hym. 3b 1569, ?.

= Ichneumon walkeri Wesmael, 1848. (syn. nov.)

This species is distinct from /. vulnemtorius Zetterstedt, 1838 (syn. dahlbomi
Wesmael), differing from it in having a weak but distinct scopa on the hind coxa, the
central area of the post-petiole strongly and rather evenly, longitudinally striate,
and the clypeus with coarse, widely spaced punctures. I am unable to distinguish
/. polyonomus Wesmael, 1859, from walkeri except in colour and these probably
represent only northern and alpine subspecies.

Hoplismenus semirufus

Type missing.

= Platylabops apricus (Gravenhorst, 1820). (Heinrich, 1937).

Ichneumon paludator

Desvignes, 1854, Trans. R. ent. Soc. Lond. n.s. 3 : 44.

Type Hym. 3b 1723, ?.

= Chasmias paludator (Desvignes, 1854) .

syn. Chasmodes paludicola (Wesmael, 1857). (Heinrich, 1937).

Ichneumon cambriensis

Desvignes, 1867, Ent. mon. Mag. 4 : 130.

Type Hym. 3b 1602, <$.

= Phaeogenes stipator Wesmael, 1855.

IV. TYPES OF BRITISH ICHNEUMONIN AE IN THE BRIDGMAN,
MORLEY AND MARSHALL COLLECTIONS

The Bridgman and Morley types, for the most part, have not been examined by
other workers. By far the greater number of the names refer to species which had
already been described. The following names are at present valid :

Dicaelotus cameroni Bridgman, 1881.

Phaeogenes distinctus (Bridgman, 1887). (comb, nov.)

syn. Herpestomus distinctus Bridgman, 1887.
Platylabops pulchellatus (Bridgman, 1889).

syn. Ichneumon pulchellatus Bridgman, 1889.
Ichneumon rufidorsatus Bridgman, 1887.
Barichneumon heracleanae (Bridgman, 1884).

syn. Ichneumon heracleanae Bridgman, 1884.
Platylabus transversus Bridgman, 1889.

syn. Platylabus lativentris Thomson, 1894. (syn. nov.)

n6 NOTES ON BRITISH ICHNEUMONINAE

(a) The Bridgman Types of Ichneumoninae
The Bridgman types of Ichneumonidae are in the Castle Museum, Norwich.

Ichneumon pulchellatus Bridgman, 1889

= Platylabops pulchellatus (Bridgman, 1889).

This is probably a synonym of I. eupitheciae Brischke, 1879.

Ichneumon rufidorsatus Bridgman, 1887
A distinct species, belonging to the genus Ichneumon s.s.

Ichneumon heracleanae Bridgman, 1884

= Barichneumon heracleanae (Bridgman, 1884).

Closely related to peregrinator L., and with this perhaps better placed in a separate
genus.

Platylabus transversus Bridgman, 1889
syn. Platylabus lativentris Thomson, 1894.

Dicaelotus cameroni Bridgman, 1881
A distinct species of Dicaelotus.

Herpestomus striatus Bridgman, 1881
= Oiorhinus pallipalpis Wesmael, 1844.

Herpestomus distinctus Bridgman, 1887
= Phaeogenes distinctus (Bridgman, 1887).

Diadromus formosus Bridgman, 1881
= Aethecerus longulus Wesmael, 1844.

Phaeogenes nitidus Bridgman, 1886
= Cratichneumon magus (Wesmael, 1855), <. (syn. nov.)

Phaeogenes similis Bridgman, 1881

= Thyraeella collaris (Gravenhorst, 1829). (syn. nov.)
This is the form with little red on the thorax.

(b) The Morley Types of British Ichneumoninae
Dinotomus spinosus Morley, 1903

Type Hym. 3b 1818, <J.

= Hoplismenus bidentatus (Gmelin, 1790). (syn. nov.)

NOTES ON BRITISH ICHNEUMONIN AE 117

Ctenichneumon plicatus Morley, 1903

Type Hym. 3b 1819, $.

Spilichneumon occisorius (F., 1793).

Nematomicrus elliotti Morley, 1903

Type Hym. 3b 1820, <j>.

= Eriplatys ardeicollis (Wesmael, 1844). (syn. nov.)

(c) The Marshall Type of Mesostenus maurus.
Mesostenus maurus Marshall, 1873

Type Hym. 3b 1563, ?.

= Hoplismenus bidentatus (Gmelin, 1790). (syn. nov.)

V. THE L. A. CARR COLLECTION OF ICHNEUMONID AE

Through the kindness of Mr. H. C. S. Halton of the Natural History Museum,
Wollaton Hall, Nottingham, I have been able to examine the L. A. Carr collection
of Ichneumoninae, and to borrow for study specimens of special interest. In the
course of this work it has become clear that specimens of continental origin were
mixed with the Lichfield specimens. Under Protichneumon fuscipennis (Wesmael) (=
Amblyjoppa fuscipennis} are a female labelled " Lichfield 1915 teste Habermehl "
and a male, " Lichfield 1916 teste Roman " of the continental form of this species
with almost black wings ; in the Harwood collection also were I male and 3 females
" Lichfield 1917 " and i female " Lichfield 1919 " of the same form received from
L. A. Carr. This form is abundantly distinct, and in the many specimens of this
species that I have seen from the British Isles none has approached this wing colour.
There were also normally coloured British specimens in the Carr collection. Under
Ctenichneumon inspector (Wesmael) were a single pair ; the female agrees completely
with two specimens of a very red, central European form of Ctenichneumon messorius
(Gravenhorst) which were bought by the late D. S. Wilkinson from Schmiedeknecht,
and which were also named inspector by Schmiedeknecht. I have seen much
material of messorius from this country, and never any specimens approaching this
form in colour ; the male is a specimen of Ambly teles uniguttatus (Gravenhorst), a
species of which I have seen no British specimens from other collections. Under
Ambly teles uniguttatus there is a specimen of the ordinary British form of Ctenich-
neumon messorius.

I feel, therefore, that at present it is better to exclude from the British list those
species which are included solely on the basis of the Carr collection. It is noteworthy
that most of the doubtful specimens are of red forms, whereas in England specimens
are most usually darker than central European forms. Up to 1919 Carr used short
black pins for pinning his material ; from 1921 to 1923 all are pinned on continental
steel pins.

n8 NOTES ON BRITISH ICHNEUMONIN AE

It is of interest to note that the male determined as Eupalamus lacteator (Graven-
horst) by Pfankuch is in reality a male of Cratichneumon clarigator (Wesmael) ; I
have, however, seen material of the true lacteator from the British Isles. Otherwise,
I have not re-examined the general records from the Carr collection.

The Types of Ichneumoninae in the L. A. Carr Collection

Three species of Ichneumoninae were described from the L. A. Carr collection.
All are synonyms.

Cratichneumon fallax Habermehl, 1923

= Cratichneumon varipes (Gravenhorst, 1829). (syn. nov.)

i

syn. Ichneumon anglicanus Schmiedeknecht, 1929.

Some of these specimens are a little discoloured by cyanide. Cratichneumon
varipes stands in the collection under varipes, fallax and magus.

Barichneumon carri Habermehl, 1923

= Barichneumon gemellus (Gravenhorst, 1829). (syn. nov.)
This is rather a dark form, as are most British specimens.

Amblyteles duplicator Roman, 1923

= Hybophorellus injucundus (Wesmael, 1854). (syn- nov.}

The type of duplicator is a female. There is also a single male of Hybophorellus
injucundus in the collection. (A number of the very interesting species present in
the collection are represented by a single pair.) I have not seen the female type of
injucundus, which was described from a specimen sent to Wesmael from the Stock-
holm Museum ; this is described as having the hind legs darker than in the above
specimen, but otherwise, in structure and sculpture, agreeing very well. I have
seen two other specimens of the male of this species, both from Schmiedeknecht, the
one purchased by Wilkinson and the other from the Morley collection.

Hybophorellus injucundus is, of course, quite distinct from H. aulicus (Gravenhorst,
1829).

VI. SPECIES TO BE DELETED FROM THE LIST OF
BRITISH ICHNEUMONINAE

In the course of revising the British species of Ichneumoninae it has been neces-
sary to re-assess the records of certain of the earlier British authors. No material
could be named at all adequately until Wesmael published his Tentamen disposi-
tionis methodicae Ichneumonum Belgii in 1844. Prior to that date the descriptions
of Ichneumoninae were given with almost no idea of any comparsion of species, and
the groupings were based on colour. Wesmael was the first author to divide the
subfamily on structural characters, and revolutionized the systematics of the group.
The next major advance is found in the works of C. G. Thomson, whose discovery
of further structural characters pointed the way to a much sounder appreciation of

NOTES ON BRITISH ICHNEUMONI N AE 119

the group. Unfortunately the workers who immediately followed Thomson made
no use of many of the characters that he indicated as of great significance, and the
study of the Ichneumoninae continued with the description of great numbers of
species, described with the use of a very limited range of characters and with great
emphasis on colour. Further, almost no examination of types was undertaken.
In recent years Gerd Heinrich, who has correlated many of the European species,
genera and tribes with exotic forms, has again made a great contribution to the
study of this subfamily.

For myself, I have found it best to base my determination of species, in greater
part, on the work of Thomson, together with Wesmael's descriptions. Also Wesmael
examined much of the type material of the Gravenhorst species, which are, at the
present time, not available for study, and thus the value of his collection is greatly
enhanced for research purposes. I was most fortunate in being able to spend a
fortnight in Brussels studying the Wesmael collection, which naturally gave me
much greater confidence in the interpretation of these species. A point of consider-
able interest was that in the majority of cases it had been possible to determine the
Wesmael species correctly from the published work of Thomson and Wesmael. A
certain number of these species I had failed to determine from the keys of Ber-
thoumieu and Schmiedeknecht ; and from the material that has been sent me by
other workers, and the determinations that I have seen given by them, I have found
that others, too, have experienced the same difficulties.

Desvignes, in 1856, examined the Stephens collection which had come to the
British Museum in 1853. It is evident from the labels on certain of the specimens
that where Desvignes did not agree with the Stephens determinations, he placed
the specimens under the species that he thought correct, and if otherwise unknown
to him, he excluded these species from his catalogue. Where I have seen no further
specimens of these species in the collections that I have examined, I also propose
to exclude them from my consideration of British species. In addition, I propose
to exclude those species first recorded by Marshall, the present whereabouts of which
is unknown.

Stephens frequently gave descriptions of the other sex of species, drawn from
Gravenhorst's description, if he thought that he had recognized one sex of the species.
Where it was later shown that Gravenhorst had wrongly associated the sexes,
Marshall recorded both names in his list, based on the descriptions given by Stephens.
Hence a number of species were brought into the British list for which no British
specimens ever existed.

I have only been concerned with those species of which no subsequent material
has been obtained. There are a number of species which were incorrectly determined
by the earlier authors but of which genuine material has been captured in recent
years.

Below are given the names of the 86 species that I consider should now be dropped
from the list as given by Kloet and Hincks (1945).

Trogus spinosus Morley, 1903
= Hoplismenus bidentatus (Gmelin, 1790).

ENTOM. Ill, 4. 10

120 NOTES ON BRITISH ICHNEUMONIN AE

Hoplismenus bispinatorius (Thunberg, 1822)

The type of this species is conspecific with H. perniciosus Gravenhorst, 1829,
according to Roman (1912). I follow Thomson in regarding perniciosus as distinct
from bidentatus (Gmelin). It is the latter species which is present in Britain.

Hoplismenus maurus (Marshall, 1873)
= Hoplismenus bidentatus (Gmelin, 1790).

Hybophorellus aulicus (Gravenhorst, 1829)

This species was introduced by Marshall, but the location of the specimen on which
the record was based is unknown to me. Reference to Wesmael's description of
Amblyteles injucundus will show that it is not a synonym of aulicus. It is described
as having the cheeks obliquely striate, and I regard Amblyteles duplicator Roman as
being a synonym of injucundus. I have not seen the types of injucundus or aulicus.
A single male of Amblyteles injucundus (Lichfield, 1923 ; teste Schmiedeknecht) and
the type of Amblyteles duplicator Roman (Lichfield, 1922) are in the Carr collection.

Protichneumon fusorius (L., 1761)

This name seems originally to have been included in the British list as the identi-
fication of Amblyjoppa fuscipennis (Wesmael), and this latter species appears as
fusorius in the early collections that I have seen and also in the Morley collection.

1 believe that the inclusion of similatorius (F.) by Marshall was due to his thinking
this name correct for part of the fusorius of British authors. Bignell had a male of
/. primatorius Forster from Kilmore, Ireland, and a female of Amblyjoppa fusci-
pennis, ex Ch. porcellus, iS.vi.iSSy (/. H. Wood], under this name. In the Carr
collection the single male (1915 ; teste Habermehl) and a single female (1916 ; teste
A. Roman) are specimens of pisorius (L.), and it would appear therefore that Carr
wrongly transcribed the name.

Although the female of fusorius is completely distinct from that of Amblyjoppa
fuscipennis, the males of these two species are difficult to differentiate. The best
characters on which to distinguish them are the form of the lower margin of the
mesopleurum which, when viewed dorsolaterally, is more strongly sinuate in fusorius
than in fuscipennis, and the pronotal collar which centrally is sub-equal in length
to the distance between the posterior ocelli, whereas in fuscipennis it is conspicuously
shorter than this distance ; fusorius has the ventral fold clearly developed on sternites

2 and 3 of the gaster, whereas in fuscipennis it is only clearly developed on sternite
2 ; also the hind tarsus is distinctly infuscate apically in fusorius. P. fusorius male
is of course at once distinguished from the other Western European species of Pro-
tichneumon in having no ventral plica on sternite 4 of the gaster.

Protichneumon disparis (Poda, 1761)

Stephens recorded this species under Trogus flavatorius Gravenhorst, 1829. The
species, however, was not included by Desvignes in his catalogue and I also exclude
it.

NOTES ON BRITISH ICHNEUMONIN AE 121

Coelichneumon sugillatorius (L., 1758)

Stephens recorded C. cyaniventris (Wesmael, 1858) as this species. The Desvignes
specimens and those in the Morley and Bignell collections were also all cyaniventris.

Coelichneumon sinister (Wesmael, 1848)

First recorded by Marshall, who included it as being the male which Gravenhorst
associated with the female of leucocerus and which Wesmael later described as
sinister. Stephens quoted a description of both male and female of leucocerus, and
it appears that Marshall included the species on this evidence. The male of sinister
has the flagellum marked with white centrally ; the female has a scopulate tubercle
beneath the hind coxa, and the flagellum not " rolled " apically in dead specimens.

Coelichneumon nothus (Holmgren, 1880)

This is included by Morley (Brit. Ichs. 1 : 29) as a variety of comitator (L.) (recte
auspex (Mueller)), but he does not state that he knew of any British specimens of
this form.

Coelichneumon periscelis (Wesmael, 1844)

This species was first recorded by Marshall as being the female of the species which
Gravenhorst had associated with the male of his Ichneumon pallifrons. Stephens
gives a description of both sexes of pallifrons and it appears that Marshall included
the species on this evidence. Specimens that I have seen determined as this species
are incorrectly named. That recorded by Morley (Brit. Ichs. 1 : 27) from the
Bignell collection is a male of Stenichneumon militarius (Thunberg) ; it is in fact the
specimen that Bignell recorded as Ichneumon pistorius Gravenhorst (= militarius}
from Bickleigh 2o.viii.i88i.

In addition to the characters given by Wesmael and Thomson for this species,
it differs from desinatorius (Thunberg, 1822) (syn. subguttatus (Gravenhorst, 1829))
in having the gena more weakly and more sparsely punctate and the malar space
shorter.

Coelichneumon funebris (Holmgren, 1864)

= biannulatus Gravenhorst, 1820, sec Thomson.

g = derasus Wesmael, 1844, sec Thomson = nigerrimus Stephens (1835).

There has been much confusion in the association of funebris (Holmgren) male
with its correct female. The specimens that I have seen so named from this country
are mostly the males of Coelichneumon nigerrimus (Stephens) (e.g., the specimens in
the Cambridge Museum cf. Kerrich, 1935, Trans. Soc. Brit. Ent., 2 : 38). The single
specimen in the Carr collection (1921 ; teste Schmiedeknecht) is a male of Coelich-
neumon purpurissatus Perkins (see p. 138). I accept Thomson's interpretation of
biannulatus, and this species is unknown to me from the British Isles. Thomson
states that the male of derasus (Wesmael) stood in several examples under /. funebris
in Holmgren's collection (Thomson, 1893, Opusc. ent. 18 : 1907) ; on the same page
he gives notes on the true male of biannulatus.

122 NOTES ON BRITISH ICHNEUMONIN AE

Coelichneumon moestus (Gravenhorst, 1829)

This species was first recorded by Marshall. I have seen no specimens from
Britain, and no specimen of it is present either in the Marshall or Fitch collection
(the latter collection contains a number of specimens from Marshall).

Coelichneumon puerulus (Kriechbaumer, 1890)

This species was first recorded on a misidentified male of Barichneumon gemellus
(Gravenhorst) (see Kerrich, 1935, Trans. Soc. Brit. Ent. 2 : 38). C. puerulus is a
species completely unknown to me.

Cratichneumon externus (Berthoumieu, 1895)

The specimen determined as this in the Hancock collection is a female of Barich-
neumon Acceptor (Scopoli). C. externus is a species completely unknown to me.

Cratichneumon fallax Habermehl, 1923
This is a synonym of Cratichneumon varipes (Gravenhorst, 1829).

Cratichneumon dissimilis auctt. angl.

= Cratichneumon jocularis (Wesmael, 1848).

The male of Aoplus ochropis (Gmelin) has been much confused with this species.
In the Johnson collection the specimens from Newcastle, Co. Down were jocularis,
those from Coolmore, Co. Donegal were ochropis.

In the Morley collection are 7^ Cratichneumon nigritarius (Gravenhorst), 2^
Cratichneumon fb'rsteri (Wesmael), 2$ Aoplus ochropis and i$, The Mound, 8 . viii . 1900
( Yerbury) of Cratichneumon jocularis. In the Carr collection was a <$ Aoplus ochropis,
Chobham, 19. ix. 1891 (Beaumont) and 4 Cratichneumon nigritarius (2 teste Roman,
i var. teste Habermehl and i var. teste Pfankuch), all from Lichfield.

There has been much confusion, systematically, between dissimilis, which I have
not seen from Britain, and jocularis. I have followed Wesmael as being the first
reviser of Ichneumon dissimilis Gravenhorst ; he selected the female, of which he
saw the Gravenhorst specimen, as being the type. The male which Gravenhorst
associated with dissimilis female is Cratichneumon jocularis (Wesmael). The two
species can be distinguished as follows :

A. $. Mesoscutum polished, with at most only a weak indication of microsculpture between
the punctures on the disc, the lateral lobes with sparse, shallow, irregularly spaced punctures ;
hind coxa, beneath, largely smooth in the apical two-thirds with only a few coarse, scattered
punctures ; hind femur and hind tibia with a conspicuous, black, apical mark ; thyridiae a
a little narrower.

o*. Face, cheeks (at least on the orbits) and the outer orbits below, yellow ; front coxa and
trochanter broadly marked with yellow, middle coxa and trochanter usually with yellow marks ;
lateral lobes of the mesoscutum more finely and shallowly punctate than the middle lobe in

NOTES ON BRITISH ICHNEUMON IN AE 123

front of the disc ; hind femur broadly black apically, hind tibia black in apical half ; tarsi
black; antenna with no white band ..... dissimilis (Gravenhorst, 1829).

(syn. Ichneumon citrinops Wesmael, 1857 <$ ; citrinops Wesmael Kriechbaumer, 1893
?c? ; zephyrus Wesmael, 1857 (sec Wesmael) ; dissimilis Gravenhorst Wesmael, 1859 ?).

B. $. Mesoscutum dull, the punctures of the lateral lobes similar to those of the central
lobe before the disc, at least for the greater part coriaceous between the punctures ; hind coxa
with the punctures only a little more widely spaced in the apical half than basally ; hind femur
and tibia entirely red ; thyridiae a little broader.

J. Face, and usually the outer orbits below, ivory, the cheeks black ; front coxa and tro-
chanter at most weakly marked with ivory ; middle coxa and trochanter rarely with ivory
marks ; lateral lobes of the mesoscutum coarsely punctate, the punctures comparable with
those on the middle lobe before the disc ; hind femur weakly, narrowly infuscate at the apex ;
hind tibia infuscate at most in apical quarter ; tarsi often, at least in part, pale ; antenna
most usually with a white ring ....... jocularis (Wesmael, 1848).

(syn. Ichneumon dissimilis Gravenhorst $ nee $, 1829 ; punctifrons Holmgren $ 1864 ;
dissimilis Gravenhorst Taschenberg, 1866 ; dissimilis Gravenhorst Holmgren <J$, 1880 ;
dissimilis Gravenhorst Kriechbaumer, 1893.)

Eupalamus oscillator Wesmael, 1844

This was first recorded from Bignell's collection ; his 2 females are Ichneumon
didymus Gravenhorst, without locality. Other British specimens that I have seen
determined as oscillator have all been E. wesmaeli Thomson. E. oscillator differs
from wesmaeli not only in colour, but in the female, in the proportions of the seg-
ments of the middle tarsus (Figs, i and 2).

Melanichneumon erythraeus (Gravenhorst, 1820)

Specimens from Stephens, Morley, Carr and Cambridge Museum collections are
all females of Barichneumon ridibundus (Gravenhorst). Bignell had i$ Aoplus
ochropis (Gmelin), Plym Bridge, 26. v. 1886, and i$ Aoplus ruficeps (Gravenhorst),
Hartlepool (/. Gardner], standing under this name. The specimen in the Johnson
collection is Barichneumon sanguinator (Rossi) (fide Stelfox).

The male is immediately distinct from all British species of this group in having
the thorax in greater part red. The female, however, might be confused with san-
guinator, but has the thyridiae broader and the front and middle coxae conspicuously
marked with ivory ; in sanguinator the coxae are sometimes marked with red or in
very small part dirty yellow. It is at once distinguished from ridibundus in lacking
the tubercle in the pronotal furrow and in having the post-petiole longitudinally
striate centrally.

It should be noted that the species recorded under Melanichneumon from the
British Isles should all be placed in Barichneumon according to Heinrich.

Melanichneumon albipictus var. obsoletus (Berthoumieu, 1895)

The specimen recorded by Johnson (1931, Ent. mon. Mag. 67 : 53) is Barich-
neumon gemellus (Gravenhorst). M. albipictus is probably better placed in Stenich-

I2 4

NOTES ON BRITISH ICHNEUMONIN AE

I 2

FIGS. 1-3. Mid tarsal segments i to 4 : fig. i, Eupalamus oscillator, $ ;
fig. 2, E. wesmaeli, $. Hind claw, fig. 3, Patroclus sputator, $.

neumon, as at present understood, though the thyridiae are less broad than in other
species of that genus.

Barichneumon sexalbatus (Gravenhorst, 1820)

This was recorded by Morley. His specimen from Lyndhurst, Hants, 12 . viii . 1901
is a male of Barichneumon coxigfyptus Heinrich.

Wesmael examined the female type of this species from Piedmont. He states
(1859) that he was unable to find any structural difference between this species and
bilunulatus Gravenhorst, and in fact, that they differed only in the colour of the
femora. Wesmael in 1848 (Bull. A cad. Belg. Cl. Sci. 15. i : 182) placed his derivator
as bilunulatus var. 6, but for the present I regard bilunulatus as being distinct from
praeceptor (Thunberg) (syn. derivator Wesmael) (cf. Thomson, 1893, Opusc. ent.
18 : 1963) . In the female, bilunulatus has conspicuous spines on the hind tibia which
are almost lacking in praeceptor. I have not seen the former species from Britain.

Barichneumon bilunulatus auctt. angl.

= Barichneumon praeceptor (Thunberg, 1822).

For notes on this species, see above, under sexalbatus.

NOTES ON BRITISH ICHNEUMONIN AE 125

Barichneumon incubitor auctt. angl.

= Barichneumon coxiglyptus Heinrich, 1951.

Roman (1932, Ent. Tidskr. 53 : 7) pointed out that Ichneumon incubitor L. was a
Gambrus, and the same species as G. ornatus (Gravenhorst) . Barichneumon incu-
bitor auctt. angl. is Barichneumon coxiglyptus Heinrich, though Barichneumon citator
(Thunberg) also occurs in this country. For the differences between these two
species see Heinrich (1951, Bonn. zool. Beitr. 3-4 : 271).

Barichneumon earn* Habermehl, 1923
= Barichneumon gemellus (Gravenhorst, 1829).

Barichneumon eupitheciae (Brischke, 1879)

The recorded specimen bred from Eupithecia helveticaria by Evans is not present
in the Morley collection. From the notes which Morley gives, this specimen was
most probably a male of Cyclolabus pactor (Wesmael). It is probable that Ichneu-
mon pulchellatus Bridgman is a synonym of /. eupitheciae Brischke, but the type of
the latter requires examination.

Barichneumon angustatus (Wesmael, 1848)

The specimens that I have seen determined as this species have all been males and,
for the most part, rather strongly marked specimens of Barichneumon deceptor
(Scopoli). It seems, originally, to have been introduced to the British list on the
strength of the Stephens description of the male of Ichneumon militaris Graven-
horst. The male recorded as angustatus by Morley is no longer present in his col-
lection. A male in the Johnson collection is Barichneumon chionomus (Wesmael).
In the Carr collection 2 males teste Schmiedeknecht are Barichneumon deceptor and
I male teste Habermehl, is Barichneumon coxiglyptus Heinrich.

No female has ever been recorded from Britain. This sex is abundantly distinct,
with its abdomen strongly narrowed apically and with conspicuous, ivory, apical
spots on tergites 4-7, but the male is very similar to deceptor ; it differs from that
species in having the post-petiole black, rarely reddish at the apex ; the sub-alar
callus more rounded and by no means carinate except at the extreme posterior end
and usually completely, or almost completely, ivory ; the head a little less narrowed
behind the eyes ; mesoscutum a little more shining, with the punctures a little
coarser and the interspaces a little wider ; tergites 5-7 with apical, ivory bands or
spots.

Ichneumon haesitator Wesmael, 1844

This was originally recorded by Marshall. Stephens quoted the description of
both sexes of Ichneumon latrator from Gravenhorst. The female of latrator Fabricius
Gravenhorst is /. haesitator, and hence Marshall recorded this species erroneously.

/. haesitator belongs to the latrator group and has no ivory spots on the scutellum
or on the apical tergites, the vertex with a yellow spot on the orbits, the stigma

126 NOTES ON BRITISH ICHNEUMONIN AE

fuscous, the thyridia conspicuously broader than the distance between the thyridiae,
the hind femur black, and the mesoscutum with distinct microsculpture between
the punctures and thus dull.

Ichneumon eremitatorius Zetterstedt, 1838

According to Roman, stigmatorius Zetterstedt, 1838, which is given by Kloet and
Hincks as a synonym of eremitatorius, is a synonym of vulneratorius Zetterstedt,
1838 ; and eremitatorius is the male of alpestris Holmgren, 1864. This latter species

1 have not seen from Britain.

Ichneumon amphibolus Kriechbaumer, 1888

This determination was by A. Roman (Johnson, 1920, Irish Nat. 29 : 19-20).
No specimen under this name is present in the Johnson collection. It is a species
that is completely unknown to me.

Ichneumon quadrialbatus Gravenhorst, 1820

Barichneumon perscrutator (Wesmael) males were determined as this species by
Stephens and Desvignes. The male stated to have been bred by Col. Partridge
from Geometra smaragdaria, in the Morley collection and recorded by him, is an
Exephanes occupator (Gravenhorst) ; there appears to have been some error in label-
ling this specimen.

/. quadrialbatus is related to gracilicornis Gravenhorst, but in the female has at
most a very small, ivory spot on tergite 5, the femora completely red, the tibiae
red with the hind tibia infuscate apically, the scutellum strongly convex, tergites

2 and 3 castaneous, most usually marked with black. The male also has the femora
completely red, and in the only specimen that I have examined the tergites are com-
pletely black. There is a female of this species sent by Gravenhorst to the Linnean
Society and now in the British Museum collection.

Ichneumon quaesitorius L., 1761

This species was first introduced by Stephens in his Catalogue where he gave
it as ? quaesitorius. A specimen so labelled from the Stephens collection is a female
of /. gracilentus Wesmael with tergites 2 and 3 infuscate ; his description in the
' Illustrations ' is taken from Gravenhorst. Marshall wrongly synonymized Chasmias
paludator (Desvignes) with Ichneumon quaesitorius and it was the former species
to which he was referring.

I. quaesitorius resembles primatorius Forster in structure, but the propodeum is
less strongly excised before the dentiparal spines and the post-petiole is coarsely,
longitudinally striate ; in colour it is of course completely distinct.

Ichneumon piceatorius Gravenhorst, 1820

This was recorded by Marshall, apparently from the specimen determined as this
by Desvignes. It is a male of Coelichneumon haemorrhoidalis (Gravenhorst). The
identity of piceatorius Gravenhorst is still in doubt.

NOTES ON BRITISH ICHNEUMONIN AE 127

Ichneumon haglundi Holmgren, 1864

Specimens so named from the British Isles are Ichneumon formosus Gravenhorst,
1829. I am indebted to Mr. G. J. Kerrich for information concerning the type of

haglundi.

Ichneumon submarginatus Gravenhorst, 1829

This species was recorded by Stephens but is not included in Desvignes' catalogue.
Bignell had placed under this name i$, no data, Eupalamus wesmaeli Thomson and
i $, Bickleigh, 28.vi.i88i, Stenichneumon militaries (Thunberg).

This species is related to nereni Thomson. In the female the thyridia is distinctly
narrower than the distance between the thyridiae, the mesoscutum is closely punctate
with microsculpture in the interspaces on the disc, the scutellum is strongly convex
and yellow, tergite 2 black or piceous, 3 black, both narrowly red on the apical
margin ; tergites 6 and 7 have a yellow, apical spot. I do not know the male of this
species.

Ichneumon languidus Wesmael, 1844

Both Marshall and Bridgman determined specimens of Ichneumon tuberculipes
Wesmael as this species. In the Carr collection the male teste Schmiedeknecht is
also a specimen of tuberculipes, but the one teste Habermehl is a male of Coelichneumon
leucocerus (Gravenhorst) .

/. languidus is one of the species with the female having elongate basal flagellar
segments, the flagellum strongly attenuate apically, the malar space long, but the area
supero-media elongate ; it has no scopulate tubercle on the hind coxa.

Ichneumon rufidens Wesmael, 1844

This in actual fact is a most interesting species with a unidentate mandible and a
very large clypeus. Marshall appears to have based his record on a single specimen,
so named by Desvignes, which is a female of Ichneumon minutorius Desvignes (syn.
captorius Thomson).

Ichneumon silaceus Gravenhorst, 1829
This was recorded by Stephens but is not in Desvignes' catalogue.

Ichneumon subcylindricus Gravenhorst, 1829

This also was recorded by Stephens but is not in Desvignes' catalogue. I do not
know this species, the type of which was re-described by Wesmael (1859).

Ichneumon tempestivus Holmgren, 1864

= Ichneumon albiger Wesmael, 1844.

As pointed out by Thomson, tempestivus is only the form of albiger which has the
hind tibia more or less marked with yellow ; this form is common in Britain.

128 NOTES ON BRITISH ICHNEUMONI N AE

Ichneumon militaris Gravenhorst, 1820

This was recorded by Stephens but is not in Desvignes' catalogue. Specimens so
determined by Bridgman belong to the form of /. extensorius L. with the hind tibia
completely infuscate.

Ichneumon caedator Gravenhorst, 1829

This was recorded by Stephens but is not in Desvignes' catalogue. The single
male (Lichfield, 1921 ; teste Schmiedeknecht) and female (Lichfield, 1922 ; teste
Roman) are correctly determined but the record requires confirmation.

Ichneumon grains Wesmael, 1855

This was recorded by Johnson (1929, Ent. mon. Mag. 65 : 135). This specimen
is a female of Chasmias motatorius (F.).

Ichneumon ffthomsoni Holmgren, 1864

The specimen from Scotland so determined by Roman is a female of Ichneumon
rufidorsatus Bridgman. For notes on thomsoni see Heinrich (1951, Bonn. zool. Beitr.
3-4:259).

Ichneumon inquinatus Wesmael, 1844
syn. Ichneumon brevigena Thomson, 1886.

Marshall wrongly synonymised /. crassorius Desvignes 1856, (= didymus Graven-
horst, 1829) with this species. Specimens named inquinatus by Morley that I have
examined are females of I. didymus. The single female in the Carr collection is
correctly determined (Lichfield, 1923 ; teste Habermehl) ; it is pinned on a Carlsbad
pin, other specimens of this date are all on continental steel pins. The record
requires confirmation.

Ichneumon multipictus Gravenhorst, 1820

Marshall apparently recorded this species from a specimen determined by Des-
vignes. It is a female of Ichneumon exilicornis Wesmael. The specimens named
multipictus in the Bignell collection are 2 females of Ichneumon validicornis Holm-
gren, no data, and Cann Wood, 6.iii. 1887 (/. Keys). The specimen so named in the
Carr collection (Lichfield, 1919 ; teste Roman) is a female of Barichneumon lepidus
(Gravenhorst) .

Ctenichneumon melanocastaneus auctt. angl.

= Ctenichneumon rubroater (Ratzeburg, 1852).

C. rubroater differs from melanocastaneus (Gravenhorst) in having the yth segment
of the flagellum quadrate, not elongate and the head with the temples broader and
only very weakly converging behind the eyes. C. melanocastaneus is probably only
a form of C. repentinus (Gravenhorst) with a black scutellum (Heinrich, 1929).

NOTES ON BRITISH ICHNEUMONINAE 129

Ctenichneumon repentinus auctt. angl.

= Ctenichneumon devylderi (Holmgren, 1871).

C. devylderi differs from repentinus (Gravenhorst, 1820) in having the spiracles
of the ist tergite much smaller, the length : breadth being at most 1-5 : I.

Ctenichneumon sputator (F., 1793)

= Patroclus sputator (F., 1793). (comb, nov.)

This was recorded by Stephens but is not in Desvignes' catalogue. The specimens
named sputator in the Johnson collection are males of Stenichneumon culpator
(Schrank). The dark male (Lichfield, 1923 ; teste Schmiedeknecht) in the Carr
collection is Cratichneumon fabricator maculifrons (Stephens) and the pair of speci-
mens with red banded abdomen (Lichfield, 1923 ; teste Habermehl) are correct, but
I question the provenance of these two specimens.

This species I consider to belong to Patroclus, which was originally described from
Central America. This genus has the claws with upstanding pectinations, at least
basally (Fig. 3) ; also the ist tergite of the gaster is strongly raised dorsally at
about the line of the spiracles.

Ctenichneumon flavocinctus (Desvignes, 1856)
= Ctenichneumon panzeri (Wesmael, 1844).

Amblyteles uniguttatus (Gravenhorst, 1829)

The female of Ctenichneumon panzeri (Wesmael) has been misidentified as this on
numerous occasions. In the Carr collection are present I $ Probolus culpatorius (L.),
i$ (teste Pfankuch) Ctenichneumon messorius (Gravenhorst) and 2<J (one con-
spicuously red marked, the other without red markings) (teste Schmiedeknecht)
which are correctly determined ; under messorius, Carr has another male of uni-
guttatus (concerning which, see p. 117). The species still requires confirmation as
British.

Amblyteles conspurcatus (Gravenhorst, 1829)

This was recorded by Stephens but is not in Desvignes' catalogue. A specimen
from the Stephens collection labelled as this is a male of Ichneumon validicornis
Holmgren.

Amblyteles viridatorius (Gravenhorst, 1820)
= Amblyteles fossorius (L., 1758).

syn. Amblyteles atratorius (F., 1793).

This was recorded by Stephens but is not in Desvignes' catalogue. A Stephens
specimen so labelled is a female of Amblyteles glaucatorius (F.). The single pair
(teste Schmiedeknecht) recorded by Carr as atratorius (F.) are a pair of Amblyteles
(vadatorius Illiger 1807) =pictus (Schrank, 1776) obviously recorded under atratorius

I 3 o NOTES ON BRITISH ICHNEUMONIN AE

due to an error of transcription. Otherwise AmUyteles quadripunctorius (Mueller)
has been mistaken for this species.

A . fossorius can be distinguished from quadripunctorius in the female in that it
lacks long spines on the extensor surface of the hind tibia, in having the temples
less narrowed behind the eyes and the lower tooth of the mandible minute, as well
as in colour ; and in the male in having the ventral plica of the abdomen infuscate,
the lower tooth of the mandible minute, the hind femur entirely red, and in the
penis valves having no inflated subapical area beneath (Figs. 4 and 5).

FIGS. 4, 5. Lateral view of apex of penis valve : fig. 4, A mblyteles fossorius ;
fig. 5, A. quadripunctorius.

Amblyteles cerinthius (Gravenhorst, 1820)

The specimens so named by Marshall are a pair of Amblyteles longigena Thomson.
Specimens so named in the Morley collection are also longigena. The British
specimens do not agree with Gravenhorst's description of cerinthius. The species
was first introduced to the British list by Stephens, but was deleted by Desvignes ;
I have found no specimen so labelled among the Stephens material.

Amblyteles microcephalus (Stephens, 1835)
= Ichneumon formosus Gravenhorst, 1829.

Amblyteles duplicator Roman, 1923

= Hybophorellus injucundus (Wesmael, 1854). (comb, nov.)
syn. Amblyteles injucundus Wesmael, 1854.

The records of these are based on a single male (Lichfield, 1923) (injucundus} and
a single female (Lichfield, 1922) (duplicator type) in the Carr collection. (See under
Hybophorellus aulicus, above.)

Amblyteles fossorius (L., 1758)

The type of this species has been wrongly determined. It is a male of the species
previously known as Amblyteles viridatorius (Gravenhorst). (See above.) Ambly-
teles fossorius auctt. angl. = Amblyteles subsericans (Gravenhorst, 1820).

NOTES ON BRITISH ICHNEUMONIN AE 131

Ctenamblyteles homocerus (Wesmael, 1854)

Bignell first recorded this species but his collection now contains no specimen
under this name.

Ctenamblyteles is closely related to Patroclus. Specimens with dark wings are
considered by Heinrich to be the typical form, which has the teeth on the shaft of
the claw evenly spaced and large. I have seen a single specimen from the Swiss
Alps, which appears to be homocerus var. noskiewiczi Heinrich, 1926, having almost
clear wings and the hind tibia only very narrowly infuscate apically, internally ;
this form is distinguished from homocerus not only in the slight colour difference,
but also in having the pectinations on the shaft of the claw thinner and contiguous ;
when more material can be examined this may well prove to be a distinct species.

Anisobas hostilis (Gravenhorst, 1820)

This was recorded by Stephens but is not in Desvignes' catalogue. Of the spec-
mens recorded by Morley, the one taken by R. C. Bradley in the New Forest is not
in Morley's collection, but the one taken by himself is a male of Barichneumon
tergenus (Gravenhorst). Also under this name in the Morley collection is another
male of tergenus and 2 females of Barichneumon gemellus (Gravenhorst).

Stenolabus daemon (Wesmael, 1844)

= Asthenolabus daemon (Wesmael, 1884).

The 2 females (Lichfield, 1921 ; teste Schmiedeknecht) under this name in the
Carr collection are Platylabus concinnus Thomson. British specimens of concinnus
are very largely black.

Apaeleticus inclytus Wesmael, 1853

= Apaeleticus bellicosus Wesmael, 1844. (syn. nov.)

I have examined the type material of these species in the Wesmael collection.

Eurylabus rufipes (Stephens, 1835)
= Polytribax curvus (Schrank, 1802).

Ischnopsidea truncator (F., 1798)

= Rhexidermus truncator (F., 1798).

The single female in the Carr collection (Lichfield, 1921 ; teste Habermehl) is cor-
rectly determined, but the species requires confirmation as British.

Heterischnus rufipes (Wesmael, 1848)

= Rhexidermus nigricollis (Wesmael, 1844). (syn. nov.)

I have examined the type material of these species in the Wesmael collection.
Heterischnus rufipes auctt. = Heterischnus pulex (Mueller, 1776), following Wesmael.

I 3 2 NOTES ON BRITISH ICHNEUMONIN AE

Dicaelotus pusillator (Gravenhorst, 1807)

This was recorded by Marshall. Specimens so named by him are Epitomus parvus
Thomson, 1891 (syn. Hemiteles pygmaeus Brischke, 1890, nee Brischke, 1888). The
Bignell collection also had Epitomus parvus under this name. In the Carr collec-
tion are I female (Lichfield, 1921 ; teste Habermehl) Dicaelotus rufoniger Berthoumieu
and i female (Lichfield, 1917; teste Pfankuch) Cratichneumon varipes (Gravenhorst).
T have been unable to identify pusillator (Gravenhorst) female, which Wesmael
placed tentatively in Dicaelotus after he had seen the type ; the males described by
Gravenhorst for this species are mixed ; the specimen sent to Wesmael was Herpe-
stomus arridens (Gravenhorst).

Dicaelotus rufilimbatus (Gravenhorst, 1820)

I have been unable to satisfy myself as to the identity of this species. Wesmael
examined the male type from the Gravenhorst collection, but was uncertain of its
identity, though expressing the opinion that it might be the male of D. erythrostoma
Wesmael. It was recorded by Stephens, but it is not in Desvignes' catalogue.

Colpognathus jucundus Wesmael Thomson, 1891 nee Wesmael, 1844.

This was first recorded as Phaeogenes jucundus by Marshall. I have seen a speci-
men so named from his collection and it is Phaeogenes ischiomelinus (Gravenhorst).
The specimen recorded by Morley from the Piffard collection is a female of Colpog-
nathus divisus Thomson. The Bradley specimen quoted by Morley is completely
doubtful and has not been traced. The Bignell specimen, Bickleigh, 8.ix.i882, is
a female of Diadromus varicolor Wesmael.

Ph: jucundus Wesmael, is a true Phaeogenes related to modestus Wesmael ;
montanus Thomson, is probably a synonym of it.

Centeterus major Wesmael, 1844

The single specimen collected by E. A. Butler, now in the Morley collection, is a
male of Colpognathus divisus Thomson.

Herpestomus furunculus Wesmael, 1844
= Herpestomus nasutus Wesmael, 1844. (syn. nov.)

syn. Herpestomus intermedius Wesmael, 1844. (syn. nov.)

I have seen the type material of these species ; Herpestomus intermedius Wesmael
I consider also to be a synonym of nasutus.

Diadromus guttulatus (Gravenhorst, 1829)
= Diadromus candidatus (Gravenhorst, 1829). (syn. nov.)

Diadromus conciliator (Wesmael, 1859)

This was recorded by Marshall, who included it as being the male which Graven-
horst associated with Ichneumon opprimator female and which Wesmael later

NOTES ON BRITISH ICHNEUMONIN AE 133

described as conciliator. Stephens quoted a description of both male and female of
opprimator, and it appears that Marshall included the species on this evidence.

Diadromus prosopius Holmgren, 1889

The specimen in the Carr collection (teste Habermehl) is a male of Phaeogenes
yusticatus Wesmael.

Notosemus albibuccus (Kriechbaumer, 1890)
= Notosemus bohemani (Wesmael, 1855).

Aethecerus pallicoxa Thomson, 1891

In the Johnson collection the specimen det. Roman as ? pallicoxa is a male of
Oiorhin^s pallipalpis Wesmael. In the Carr collection are also a pair labelled as
pallicoxa (teste Roman), which are likewise pallipalpis.

Mevesia similis (Bridgman, 1881)
= Thyraeella collaris (Gravenhorst, 1829).

Baeosemus mitigosus (Gravenhorst, 1829)

syn. Herpestomus phaeocerus Wesmael, 1844.

This was recorded by Stephens but is not in Desvignes' catalogue. A specimen
so named from the Stephens collection is a female of Phaeogenes fuscicornis Wesmael.
Gravenhorst recorded Ph. mitigosus var. I <$ from Netley, but this, from the descrip-
tion, cannot be conspecific with mitigosus and is probably a male Phaeogenes.

Phaeogenes nigridens Wesmael, 1844

The Johnson specimen is a male of Ph. ophthalmicus Wesmael from Newcastle,
Co. Down. In the Carr collection is a specimen labelled "Ph. ? nigrinus Berth.
teste Schmiedeknecht ? nigridens " which is a female of Aethecerus discolor Wesmael.

There has been considerable confusion concerning this species. It is related to
planifrons, heterogonus and curator, but differs in the female in having the fore wing
with the basal plates infuscate, not yellow ; the hind coxa with a transverse carina
beneath which extends to the inner, ventro-lateral margin where it is only slightly
raised, and behind this carina the coxa is a little depressed ; the head is mere
quadrate.

Phaeogenes limatus Wesmael, 1844

= Phaeogenes fulvitarsis Wesmael, 1844.

The specimen in the Carr collection (Lichfield, 1923 ; teste Schmeideknecht) is a
female of Phaeogenes flavidens Wesmael. Ph. limatus Wesmael Thomson is quite
another species which I regard as being conspecific with Phaeogenes infimus Wesmael.

Phaeogenes homochlorus Wesmael, 1844
= Phaeogenes invisor (Thunberg, 1822). (syn. nov.)

I 3 4 NOTES ON BRITISH ICHNEUMONIN AE

Phaeogenes lascivus Wesmael, 1855

The specimen in the Carr collection (Lichfield, 1921 ; teste Schmiedeknecht) is a
female of Phaeogenes fuscicornis Wesmael.

Phaeogenes minimus Berthoumieu, 1901

The specimen in the Carr collection (Lichfield, 1922 ; teste Habermehl) is a female
of Herpestomus nasutus Wesmael.

Phaeogenes nitidus Bridgman, 1886.
= Cratichneumon magus (Wesmael, 1855).

Phaeogenes coryphaeus Wesmael, 1844

This species was first recorded by Bridgman on a specimen that he examined from
Champion's collection. There is no specimen of this species in the Bridgman col-
lection nor amongst the Champion material which came to the British Museum.
Specimens so named in Johnson's collection are Ph. fulvitarsis Wesmael nee auctt.
(syn. ruficoxa Thomson) .

The form of the head in coryphaeus is quite distinct from the other species of the
fulvitarsis group, being comparable in shape with planifrons, and having the occipital
carina about as near to the posterior ocellus as the distance between the posterior
ocelli.

Phaeogenes ruficoxa Thomson, 1891

= Phaeogenes fulvitarsis Wesmael, 1844, nee auctt. (syn. nov.)

The species which has previously been determined as fulvitarsis by British authors

is Phaeogenes rusticatus Wesmael. The species previously determined as rusticatus

I am describing as a new species.

Phaeogenes minutus Wesmael, 1844

The two specimens in the Carr collection are labelled " Lichfield, 1921 ; teste
Schmiedeknecht," which is a male of Oiorhinus pallipalpis Wesmael, and " Lich-
field, 1923 ; teste Schmiedeknecht," which is a female of Phaeogenes ischiomelinus
(Gravenhorst) .

The type of this species is in bad condition, but I believe it to be a small female of
infimus Wesmael.

Phaeogenes ?? acutus (Gravenhorst, 1829)

This species was recorded by Marshall. Specimens so named from his collection
are 2 females of Aethecerus discolor Wesmael.

Phaeogenes socius Holmgren,

= Phaeogenes osculator (Thunberg, 1822), <$ (sec Roman).

The male and female, Lichfield, 1921, in the Carr collection, teste Schmiedeknecht,
are a pair of osculator.

NOTES ON BRITISH ICHNEUMONIN AE 135

Melanomicrus elliotti Morley, 1903.
= Eriplatys ardeicollis (Wesmael, 1844).

VII. SPECIES RECORDED IN THE ADDENDA OF MORLEY'S
BRITISH ICHNEUMONS

In the Addenda to Morley's British Ichneumons, 1 : 292-293, the following species
are listed. I have examined the specimens and include my identifications.

Phaeogenes socius Holmgren, 1889, $; the Foxall specimen is Aethecerus nitidus
Wesmael, the Piffard specimen is Aethecerus discolor Wesmael.

Phaeogenes macilentus Wesmael, 1844, det. Morley is Oiorhinus pallipalpis Wesmael.

Diadromus tenax Wesmael, 1844, det. Morley is Ischnus nigricollis Wesmael.

Misetus oculatus Wesmael, 1844, is correct.

Phaeogenes murcifer Holmgren, 1889. I am unable to find this specimen in the
Morley collection.

Colpognathus armatus Thomson, 1891, det. Morley is Cotyognathus divisus Thomson.

VIII. CHANGES IN THE TRIVIAL NAMES OF
BRITISH ICHNEUMONINAE

The following changes in specific names due to homonymy and synonymy have
to be made :

Trogus pictus (Kriechbaumer, 1882).

= Psilomastax pyramidalis Tischbein, 1868.

Cercodinotomus Uchida, 1940, type species Psilomastax pictus Kriechbaumer,
1882, is a direct synonym of Psilomastax Tischbien, 1868, type species Psilomastax
Pyramidalis Tischbein, 1868, the two type species being conspecific. (syn. nov.)

Protichneumon erythrogaster (Stephens, 1835) nee (Gmelin, 1790).

= Protichneumon coqueberti (Wesmael, 1848).
Amblyjoppa laminatoria (Fabricius, 1798).

= Amblyjoppa proteus (Christ, 1791).
Coelichneumon subguttatus (Gravenhorst, 1829).

= Coelichneumon desinatorius (Thunberg, 1822).
Coelichneumon derasus (Wesmael, 1844).

= Coelichneumon nigerrimus (Stephens, 1835).
Stenichneumon trilineatus (Gmelin, 1790).

= Stenichneumon lineator (Fabricius, 1781).
Platylabus semirufus (Desvignes, 1856).

= Platylabops apricus (Gravenhorst, 1820).

Cratichneumon gravenhorstii (Boyer de Fonscolombe, 1847) nee (Wesmael, 1836)
nee (Guerin Meneville, 1838).

= Cratichneumon albifrons (Stephens, 1835).
Cratichneumon liostylus (Thomson, 1887).

= Cratichneumon infidus (Wesmael, 1848).
Melanichneumon nudicoxa (Thomson, 1888).

= Barichneumon digrammus (Gravenhorst, 1820).

ENTOM. Ill, 4. II

136 NOTES ON BRITISH ICHNEUMONIN AE

I have examined a male of digrammus sent by Gravenhorst to the Linnean Society
and now in the British Museum.

Melanichneumon perscrutator (Wesmael, 1844) nee (Thunberg, 1822).

= Barichneumon maculicauda notn. nov.
Melanichneumon albolineatus (Gravenhorst, 1829).

= Barichneumon albilineatus (Gravenhorst, 1820).
Barichneumon deceptor (Gravenhorst, 1820).

= Barichneumon deceptor (Scopoli, 1763).
Ichneumon obsessor Wesmael, 1844.

= Ichneumon formosus Gravenhorst, 1829.

I accept this synonymy, given by Wesmael (1859), after he had examined the
Gravenhorst type.
Ichneumon confusorius Gravenhorst, 1829.

= Ichneumon confusor Gravenhorst, 1820.
Chasmias paludicola (Wesmael, 1857).

= Chasmias paludator (Desvignes, 1854).
Exephanes hilaris (Gravenhorst, 1829) nec (Say, January 1829).

= Exephanes ischioxanthus (Gravenhorst, 1829).
Amblyteles vadatorius (Illiger, 1807).

= Amblyteles pictus (Schrank, 1776).
Amblyteles negatorius (Fabricius, 1793).

= Amblyteles pulchellus (Christ, 1791.) (syn. nov.)
Diadromus rubellus (Gravenhorst, 1829, <$) nec (Gmelin, 1790).

= Diadromus quadriguttatus (Gravenhorst, 1829).
Epitomus pygmaeus (Brischke, 1890, nec 1888).

= Epitomus parvus Thomson, 1891.
Phaeogenes melanogonus (Gravenhorst, 1820) emend.

= Phaeogenes melanogonos (Gmelin, 1790).
Phaeogenes scutellaris Wesmael, 1844.

= Phaeogenes maculicornis (Stephens, 1835).
Phaeogenes tibiator (Thunberg, 1822) nec (Gravenhorst, 1820).

= Phaeogenes callopus Wesmael, 1844.

IX. CHANGES IN GENERIC PLACEMENT OF SPECIES OF
BRITISH ICHNEUMONINAE

Protichneumon fuscipennis (Wesmael, 1844) to Amblyjoppa Cameron, 1902.

Stenichneumon ochropis (Gmelin, 1790), castaneus (Gravenhorst, 1820), and ratze-
burgii (Hartig, 1838) to Aoplus Tischbein, 1874.

Cratichneumon ruficeps (Gravenhorst, 1829) an< ^ rubricosus (Holmgren, 1864) to
Aoplus Tischbein, 1874.

The authenticated British species included under Melanichneumon Thomson, 1893,
to Barichneumon Thomson, 1893.

Barichneumon magus (Wesmael, 1855) to Cratichneumon Thomson, 1893.

NOTES ON BRITISH ICHN EUMONIN AE 137

Barichneumon pulchellatus (Bridgman, 1889) to Platylabops Heinrich, 1950.
Barichneumon semirufus (Gravenhorst, 1820) to Cratichneumon semirufus
(Gravenhorst, 1820). (comb, nov.j

Barichneumon rufidorsatus (Bridgman, 1887) to Ichneumon L., 1758.
Barichneumon humilis (Wesmael, 1857) to Aoplus Tischbein, 1874.
Chasmias lugens (Gravenhorst, 1829) to Ichneumon L., 1758.
Ctenichneumon haereticus (Wesmael, 1854) to Ichneumon L., 1758.
Spilichneumon fabricii (Schrank, 1802) to Amblyteles Wesmael, 1844.
Platylabus rubellus (Gmelin, 1790) and exhortator (F., 1787) to Ectopius Wesmael,

1859-
Platylabus infractorius (L., 1761) to Pristiceros Gravenhorst, 1829.

Stenolabus Heinrich, 1936, nee Schulthess-Rechberg, 1910 = Asthenolabus
Heinrich, 1951.

Eurylabus dims Wesmael, 1853, to Zimmeria Heinrich, 1933.

Ischnopsidea Viereck, 1914 = Rhexidermus Foerster, 1868.

Herpestomus distinctus Bridgman, 1887, to Phaeogenes Wesmael, 1844.

Proscus cephalotes (Wesmael, 1844) and suspicax (Wesmael, 1844) to Phaeogenes
Wesmael, 1844.

X. ADDITIONS TO THE LIST OF BRITISH ICHNEUMONINAE

The list that follows contains those species which have been described previously
and of which I have now seen material from Britain. In a few cases these additions
are due only to the fact that the majority of the specimens previously determined
under certain names were incorrectly identified, and thus a species will have to be
deleted at the same time (e.g., Ichneumon languidus auctt. angl. for the most part
is Ichneumon tuberculipes Wesmael and no specimens of languidus have been seen
from Britain).

Certain species, too, which were formerly given as synonyms, are reintroduced as
distinct species, as further material and information concerning them has now been
assembled. But allowing for this, some 48 species are added as completely new
additions to the list given by Kloet and Hincks, and in all, 65 names are added.
These numbers do not include the species described as new later in this paper, nor
changes of name due to direct synonymy. I have seen the types or syntypes of
species marked with an asterisk.

LlSTRODROMINI

Anisobas platy stylus Thomson, 1888

ENGLAND: Staffs; Maer Woods, i$, 18.^.1947 (H. W. Daltry) det. G. J.
Kerrich. (H. W. Daltry Coll.)

PROTICHNEUMONINI
Coetichneumon falsificus (Wesmael, 1844)*

ENGLAND : Essex ; Colchester, i$, 1910 (P. Harwood) ; Devon ; Newton Abbot,
i$, 27 . vi . 1935 (R. C. L. Perkins) . (B.M. Coll.)

138 NOTES ON BRITISH ICHNEUMONIN AE

Coelichneumon solutus (Homgren, 1864)

SCOTLAND : Inverness ; Cairn Gorm, 3000 ft., 1$, 27. vi. 1934 (R. B. and /. E.
Benson), det. A. Roman. (B.M.Coll.)

Coelichneumon purpurissatus nom. nov.

. syn. Ichneumon nigrator Fabricius, 1793, nee Mueller, 1776.
Ichneumon lineator varr. 1-5 Wesmael, 1844.
Coelichneumon lineator auctt. ex parte.

I am using this name for the species which is very closely related to comitator L.,
but in which the hair on the propodeum is fuscous and not pale, and the abdomen
more clearly metallic. Trentepol in his redescription of nigrator described the hind
femur as being red, with a fuscous line outwardly, the front and middle femora
being black. This would apply better to the present species than to comitator.
Coelichneumon subviolaceiventris (Pic) (comb, nov.} (=Ichneumon subviolaceiventris
Pic, 1908, ^change, 24 : 67) is very similar to purpurissatus but can at once be dis-
tinguished by the lack of the scopa on the hind coxa.

Various other names are given by Dalla Torre as synonyms of Ichneumon lineator
auctt. Of these biguttatus Thunberg (syn. biguttorius Thunberg) is Stenichneumon
lineator (F.) ; narrator F. is more probably a specimen of Coelichneumon comitator
(L.) ; fuscatorius Thunberg is uncertain as to position, the type was not found by
Roman, and it is described as having the legs red with the femora black ; /. caeru-
lescens Tischbein is described as having the apical angles of the ist tergite marked
with ivory, and I have seen no specimen agreeing with his description of this species,
which may be distinct. I have therefore been forced to propose a new name for
this form.

ENGLAND : Cornwall ; Gurnards Head, i^, 1-8. viii. 1936, on umbels of Smyrnium
(G. D. Hale Carpenter) ; Polperro, i$ (T. A. Marshall] ; St. Minver, i$, i. viii. 1910
(Le Marchant) ; no locality, i$, bred i.vii.igio ex Dianthoecia barrettii (H. M.
Edelsten) (B.M.Coll.) ; Devon; Prawle Point, i<$, 8. v. 1938 ex. Dianthoecia bar-
rettii (A. J. L. Bowes) ; Cornwall; nr. Bude, 1$, 23. vi. 1910 (F. C. Woodforde) ;
Hants; New Forest, Ashurst Walk, i<j>, 9-10. vi. 1912 (F. C. Woodforde) (Hope
Department, Oxford). Cornwall ; Carbis Bay, 2$, 29. viii. 1928 and 13. ix. 1931 (A.
Thornley) (C. Morley Coll.).

Coelichneumon serenus (Gravenhorst, 1820)

This I regard as a distinct species. Of it I have seen only i male, Desvignes Coll.,
and 2 females from the old British collection in the British Museum.

Coelichneumon truncatulus (Thomson, 1886)*

I am now also regarding this species as distinct. ENGLAND : Lanes ; Kent ;
Surrey ; Herts ; Bucks. 13^, 11$, v-vi. (B.M.Coll.)

NOTES ON BRITISH ICHNEUMONIN AE 139

HERESIARCHINI
Heresiarches eudoxius (Wesmael, 1844)*

ENGLAND : Hants ; New Forest, Minstead Wood, i<$, I5.vii.i934 (C. Morley) ;
, i? (Capron) (cf. Morley, British Ichs. 1 : 203, footnote). (C. Morley Coll.)

ICHNEUMONINI

Hoplismenus bidentatus (Gmelin, 1790)
syn. Hoplismenus bispinatorius auctt. angl.

Aoplus defraudator (Wesmael, 1844)*

SCOTLAND : Inverness and Perthshire, and IRELAND : Co. Wicklow ; 3^, n$,
vi ix (B.M.Coll.)

Aoplus altercator (Wesmael, 1855)

ENGLAND : Bucks and Somerset. SCOTLAND : Perthshire and Inverness. IRE-
LAND : Co. Wicklow. 2$, 9$, vi viii. (B.M.Coll.)

Aoplus virginalis (Wesmael, 1844)*

ENGLAND : Suffolk ; Staverton Thicks, 2$, n.vii.i9i4 (C. Morley). (C. Morley
Coll. det. as Barichneumon eupitheciae Brischke.)

Cratichneumon clarigator (Wesmael, 1844)*
ENGLAND : Surrey ; Hunts ; Herts ; Kent. 15,^, 2$, v-vii. (B.M.Coll.)

Cratichneumon jocularis (Wesmael, 1848)
syn. Cratichneumon dissimilis auctt. angl.

IRELAND : Co. Down ; Newcastle, 2$ (W. F. Johnson] (Dublin Museum).
ENGLAND : N. Yorkshire. SCOTLAND : Nairn and Aberdeen. IRELAND : Co.
Wicklow. 6<$. (B.M.Coll.) SCOTLAND: "The Mound," i$, 8. viii. 1900 (Yerbury).
(Morley Coll.)

Cratichneumon pseudocryptus (Wesmael, 1857)*

ENGLAND : i$. (Capron) (In C. Morley Coll. under Plectocryptus tinctorius
(Gravenhorst)).

Barichneumon leucocheilus (Wesmael, 1844)*

ic?, labelled " 734 transfuga " from British Collection. ENGLAND : Surrey ;
Byfleet, i$, 26-29^.1949 (R. B. Benson). (B.M.Coll.)

Barichneumon praeceptor (Thunberg, 1822)
syn. Barichneumon bilunulatus auctt. angl.

I 4 o NOTES ON BRITISH ICHNEUMONIN AE

Barichneumon coxiglyptus Heinrich, 1951

syn. Barichneumon incubitor auctt. angl.

A rather common species. In the B.M. Coll. is material from Surrey ; Suffolk ;
Devon ; Cambs ; Herts ; Hereford ; Kent, v-vi, viii-ix.

Barichneumon citator (Thunberg, 1822)

ENGLAND: Devon; Dartmoor, i<$, i.vi.i935 (R. C. L. Perkins); Lydford, i$,
8-10 . ix . 1946 (/. F. Perkins) . (B.M.Coll.)

Ichneumon exilicornis Wesmael, 1857*
There is a single $ of this species from the Desvignes collection. (B.M.Coll.)

Ichneumon tuberculipes Wesmael, 1848*

This species has most usually been recorded as languidus Wesmael in Britain. In
the British Museum collection there is material from Devon, Hants and Essex.

Ichneumon megapodius Heinrich, 1949

A parasite of Amathes alpicola and I have examined 4 males and 3 females, all
reared from pupae of this host from Perthshire (K. Todd) and Inverness (P. Harwood}.
British specimens are much more extensively marked with red than those from the
Alps.

Ichneumon fuscatus Gravenhorst, 1829

There is a single female which may belong to the above species, and which is quite
distinct from any other species known to me, from Surrey ; Chobham, 9 . vi . 1910
(G. Le Marchant). (B.M.Coll.)

Ichneumon eurycerus Thomson, 1890

SCOTLAND: Inverness; Nethy Bridge, i$, 30. v. 1931; Perth; Rannoch, i$,
v.1922; ENGLAND: W. Suffolk, i$, 1928 (P. Harwood). (B.M.Coll.)

Ichneumon crassifemur Thomson, 1886*

I have only seen material of this species from the Desvignes and Stephens collec-
tions. It was previously regarded as a synonym of molitorius L.

Ichneumon melanotis Holmgren, 1864

This species, too, I regard as distinct from molitorius. In the British Museum
collection there is material from Hants, Bucks and Inverness.

Ichneumon septentrionalis s. sp. atrifemur nom. nov.

syn. scelestus Perkins, 1952, nee Cresson, 1864.
For information concerning this species see Perkins, 1952, Bull. ent. Res. 48 : 361.

NOTES ON BRITISH ICHNEUMONIN AE 141

Ichneumon lautatorius Desvignes, 1856*
This species is now regarded as being distinct from sarcitorius L.

Exephanes amabilis Kriechbaumer, 1895

IRELAND : Co. Meath, i$, ix. 1922 (G. L. R. Hancock] det. G. Heinrich. ENGLAND :
Cambs ; Adventurers fen, i<$, 8. vii. 1924 (G. L. R. Hancock}. (B.M.Coll.) Norfolk ;
Wroxham, i$, 3.ix.i9O2 (G. A. K. Marshall}. (Hope Department, Oxford).

Exephanes caelebs Kriechbaumer, 1890

IRELAND : Co. Cavan ; Lisney, i$, 31. .1936 ; Sandpit, Arva Road, i$, 21. v.
1934 (R. C. Farris} ; Co. Wicklow ; Enniskerry, i$, 25.viii.i935 (/. F. Perkins}.
(B.M.Coll.)

Thyrateles camelinus (Wesmael, 1844)*

ENGLAND : Essex, Loughton, i$ (G. C. Champion) ; i$, Stephens Coll. (B.M.Coll.)
Both specimens were previously determined as Ctenicheunom castigator (F.).

Ctenichneumon rubroator (Ratzeburg, 1852)
syn. Ctenichneumon melanocastaneus auctt. angl.
Not uncommon. ENGLAND : Devon ; Essex ; Cornwall ; vi vii. (B.M.Coll.)

Ctenichneumon devylderi (Holmgren, 1871)

syn. Ctenichneumon repentinus auctt. angl.

There has been confusion between this species and repentinus (Gravenhorst), of
which I have seen no specimens from Britain. ENGLAND : Bucks and Essex, 6<^,
5$. (B.M.Coll.)

Spilichneumon johansoni (Holmgren, 1871)
ENGLAND : Northants ; Herts ; Kent ; Bucks ; Surrey. $<$, 4$. (B.M.Coll.)

Spilichneumon stagnicota (Thomson,

ENGLAND : Gloucestershire ; Forest of Dean, High Meadow Woods, i$, 9. vi. 1936
(E. B. Britton and /. F. Perkins} ; Hants; New Forest, i$, 15. vi. 1912 (G. T.
Lyle) ; Hereford ; Malvern Hills, i$, 20-21. iv. 1935 (R. B. Benson) ; Kent ; Tun-
bridge Wells, i?, viii.1920 (C. G. Nurse). (B.M.Coll.)

Ambly teles gradatorius (Thunberg, 1822)
syn. Amblyteles egregius (Gravenhorst, 1829).

SCOTLAND: Perth; i<$, vi.i9i4, ex Eurois occulta (P. Harwood} ; Rannoch,
, vi.i927 (P. Harwood}. (B.M.Coll.)

I 4 2 NOTES ON BRITISH ICHNEUMONIN AE

Atnbly teles longigena Thomson, 1888.
syn. Avnblyteles cerinthius auctt. angl.

SCOTLAND : Forfar ; Perth ; Isle of Arran. ENGLAND : Kent ; Devon. 6<$, 7$.
(B.M.Coll.)

Amblyteles elongatus Brischke, 1878
syn. Amblyteles subsericans var. elongatus Brischke sec Heinrich.

This is a common species which used to be determined as subsericans (Gravenhorst),
and of which the male appears to be much rarer than the female. Heinrich has
described the genus Limerodops for this species.

Amblyteles subsericans (Gravenhorst, 1820)
syn. Amblyteles fossorius auctt. angl.

This is a more robust species than elongatus and is rarer. I have seen material
of it from ENGLAND : Bucks, Herts and Glos., the male being more frequently
captured than the female. A . subsericans has been given as a synonym of fossorius
L., but I have re-examined the male type of that species and found it to be A.
viridatorius (F.), a species unknown to me from Britain.

Acolobus sericeus Wesmael, 1844*

ENGLAND : Essex and Hants, 3$, 5$, the Hants specimens bred from Ectropis
luridata (syn. extersaria). (B.M.Coll.)

Probolus concinnus Wesmael, 1853*

This is a distinct species. ENGLAND : Hants ; W. Suffolk ; Northants ; Corn-
wall. 6, 4$, vii-ix. (B.M.Coll.)

PLATYLABINI

Rhyssolabus arcticus Hellen, 1942

SCOTLAND : Inverness ; Cairn Lochain, 3-4,000 ft., ig, 3.^.1934 (R. B. and/. E.
Benson) ; Aviemore, ex Psodos coracina, i<j>, 22^.1948 (A.Richardson). (B.M.Coll.)
I am indebted to Mr. G. Heinrich, who pointed out the probable identity of this
species.

Platylabus intermedius Holmgren, 1871

ENGLAND: Devon; Bickleigh, i<?, 19. vi. 1880 (G. C. Bignell), previously det.
as pedatorius (F.) (Bignell Coll.) ; Cornwall; Botusflemming, i<? (T. A. Marshall).
SCOTLAND : Inverness ; Cairn Lochain, 3-4,000 ft., i<$, 3.vii.i934 (R. B. and/. E.
Benson) ; Ross-shire ; Glenshiel, i<$, i.vii.i934 (0. W. Richards). (B.M.Coll.)

Platylabus opaculus Thomson, 1888

ENGLAND : Essex ; Colchester (Harwood) ; IRELAND : Co. Cavan ; Farnham,
29. vi. 1935 (R. C. Farris) 2,$, 6$. (B.M.Coll.)

NOTES ON BRITISH ICHNEUMONIN AE 143

Platylabus gigas Kriechbaumer, 1886.

ENGLAND: Hants; New Forest, itf, 29. iv. 1912 ex Selenia tetmlunaria (F. C.
Woodforde). (Hope Department, Oxford).

Platylabus punctifrons Thomson, 1888*

ENGLAND : Essex ; Colchester, i^, i<j> (Harwood) ; Frinton, 2$, viii and ix.ixjig
(C. G. Nurse) ; Suffolk ; Sudbury, i$, 1916 (P. Harwood) ; Kent ; Halstow Pier
(flying south over Thames estuary), i$, 25. ix. 1949 (/. F. Burton). (B.M.Coll.)
3$ ex Eucymatoge subnotata, 18. viii. 1879 (previously det. as pedatorius (F.)) (G. C.
Bignell Coll.).

Platylabus obator Desvignes, 1856*

This is a distinct species, recognizable from pedatorius (F.) by the black stigma
and more strongly raised epicnemia, and in the female by the conspicuously black
apex to the hind femur. ENGLAND : Devon ; Hereford ; Herts ; Surrey ; Glos. ;
Cornwall. 49^, 7$, v-vii. (B.M.Coll.) In 1941 and 1942 the male of this species
was quite common flying around Galium molugo in the lanes around Newton Abbot.

Platylabus rufiventris Wesmael, 1844.*
ENGLAND : Devon ; Dartmoor, i<$, 18. viii. 1934 (R. C. L. Perkins). (B.M.Coll.)

Platylabus concinnus Thomson, 1888

ENGLAND : Devon ; Braunton, i<$, 5.vii.i93O (Wright) ; Cornworthy, i$ (T. A.
Marshall) ; Essex ; Colchester, 1$, 1909 (Harwood) ; Suffolk ; i$, bred 4. viii. 1911
(C. G. Nurse). Both the females were previously det. as dolorosus (Gravenhorst).
(B.M.Coll.) Lichfield, 2$, 1921, det. as Platylabus daemon by Schmiedeknecht.
(Carr Coll.) Heinrich has placed this species in Asthenolabus, but in spite of its
rather small thyridiae, I consider it better retained in Platylabus.

Asthenolabus latiscapus Thomson, 1894
ENGLAND : Devon and Suffolk ex Euphyia cuculata, 7$, 3$. vi-vii. (B.M.Coll.)

PHAEOGENINI
Dicaelotus orbitalis Thomson, 1891

ENGLAND: Suffolk; Assington, i<j>, i6.v.i9O2 (C. Morley). (B.M.Coll.)
IRELAND: Co. Kildare ; Rye Water, i?, 8. vii. 1945 (A. W. Stelfox). (Stelfox Coll.)

Dicaelotus pudibundus (Wesmael, 1844)*

ENGLAND: Bucks; Slough, i$, 9.^.1937 (0. W. Richards}. (B.M.Coll.) Isle
of Wight ; Ryde, i$, n. viii. 1902 and i$, 17. viii. 1903 (det. previously as rufilim-
batus (Gravenhorst)). (C. Morley Coll.)

144 NOTES ON BRITISH ICHNEUMONI N AE

Dicaelotus pictus (Schiedeknecht, 1903). (comb, nov.)

syn. Deloglyptus pictus Schmiedeknecht.

ENGLAND : i<$, Desvignes Coll. (previously det. as Herpestomus arridens (Graven-
horst)) ; Herts ; Boxmoor, i$, n.v.i935 (R. B. and/. E. Benson). (B.M.Coll.)

Dicaelotus punctiventris (Thomson, 1891). (comb, nov.)

syn. Deloglyptus punctiventris Thomson.

ENGLAND : Bucks ; Burnham, in empty barn, i$, 2i.vii.i94i (0. W. Richards).
(B.M.Coll.) Suffolk; Barton Mills, i$, 5.vi.i9i6 (C. Morley) previously det. as
Melanomicrus elliotti Morley. (C. Morley Coll.)

Dicaelotus erythrostomus Wesmael, 1844

ENGLAND : Hants ; Sopley Common, on sand, i$, i6.vii.i949 (0. W. Richards) ;
IRELAND: Co. Galway ; Bencorr, 2,000 ft., i$, 3.^.1946 (R. A. Lever}. (B.M.
Coll.)

Dicaelotus cameroni Bridgman, 1881

This I regard as a distinct species. I have seen only the type from Britain and a
single female from Germany (Ruthe Coll.).

Dicaelotus rufoniger Berthoumieu, 1896
syn. Dicaelotus pumilus var. rufoniger Berthoumieu.

The majority of the material of this species that I have examined has been from
Ireland. In the B.M.Coll. are specimens from Co. Wicklow, Co. Sligo, Co. Kildare
and Co. Wexford (A. W. Stelfox), and a $ from Herts ; St. Albans (T. A. Marshall).
I have also examined 3$ and 20$ in the A. W. Stelfox Coll.

Dicaelotus inflexus Thomson, 1891
ENGLAND : i$ (Desvignes). (B.M.Coll.) IRELAND : i$. (Stelfox Coll.)

Dicaelotus morosus Wesmael, 1855
ENGLAND : Bucks ; Kent. IRELAND : Co. Leitrim. 27$. (B.M.Coll.)

Eparces grandiceps Thomson, 1891*

No locality, i$, 14. iv. 1900, det. as C. opprimator C.M., ix. 1902. (C. Morley Coll.)
This is probably the specimen recorded by Morley from Bramford Marshes, near
Ipswich (Brit. Ichs. 1 : 281).

Micrope macilenta (Wesmael, 1844)

IRELAND : Co. Kildare; Skerries Bog, i$, 23. ix. 1948 (A. W. Stelfox). (Stelfox
Coll.) I have followed Thomson's interpretation of this genus and species.

NOTES ON BRITISH ICHNEUMONINAE 145

Diadromus tenax Wesmael, 1844*
ENGLAND : !<, Desvignes Coll. (B.M.Coll.)

Diadromus albinotatus (Gravenhorst, 1829)

ENGLAND: Devon; Newton Abbot, 2$, 3.viii.i94i (/. F. Perkins) and i$,
6.viii.i94i (R. C. L. Perkins). (B.M.Coll.)

Phaeogenes curator (Thunberg, 1822)

ENGLAND : i<j>, Desvignes Coll. ; Monmouth ; Trelleck Beacons, i<j>, io.vi.i936
(E. B. Britton and /. F. Perkins). (B.M.Coll.)

Phaeogenes elongatus Thomson, 1891

ENGLAND: Herts; Bricket Wood, i$, 17^.1936 (R. B. Benson). (B.M.Coll.)
No locality, 1$, bred ex E. nigricostana (Bignell) ; Suffolk ; Monks Soham Garden,
ic?, 14.^^.1917; i$, without data. All previously det. as Centererus opprimator
(Gravenhorst). (C. Morley Coll.)

ALOMYINI
Alornya semiflava Stephens, 1835*

A rather common species, appearing later in the year than debellator (F.) (see
Perkins, 1952, Bull. ent. Res. 48 : 363).

XI. DESCRIPTIONS OF NEW SPECIES OF
BRITISH ICHNEUMONINAE

Below are given the descriptions of the species that I believe to be previously
undescribed. Due to the lack of grouping of the many species that have already
been described, it is very difficult indeed to be at all certain of this. When the
opportunity arises for someone who has very extensive collections of the European
species of this subfamily, to see all the types that are scattered throughout Europe,
it is certain that many species that are now thought to be distinct will prove to be
already described under other names.

Ichneumon quartanus sp. nov.

Similar to Ichneumon insidiosus Wesmael (of which I have examined the series
of syntypes) in general facies and in the relatively short basal segments of the flagel-
lum, but at once distinguished from that species in having a large, ivory, apical spot
on tergite 5 of the gaster.

.. Head distinctly narrowed behind the eyes ; malar space long, longer than the post-anellus
(malar space : breadth of base of mandible about 1-2:1); mandible strongly narrowed to the
apex, the lower tooth distinct ; inner orbits sub-parallel centrally ; antenna with 42-44 seg-
ments ; flagellum with the basal segments somewhat short (Fig. 6), conspicuously widened
beyond the middle and strongly attenuate apically, beneath, at its widest, with the flattened

I 4 6

NOTES ON BRITISH ICHNEUMONIN AE

ventral area of the segments a little broader than long ; post-anellus length : breadth about
1-5:1; irons, dorsally, coarsely rugose-punctate, with only indistinct microsculpture ; face
rather coarsely punctate, irregularly striate beneath the antennal sockets and, in part, on the
epistoma ; clypeus rather coarsely, rather sparsely punctate with an apical row of closer
punctures.

Thorax similar in sculpture to insidiosus, but the pronotum, laterally, with coarser punctures ;
propodeum with the area superomedia a little transverse ; area petiolaris rugose-punctate,
not divided from the more coarsely rugose areae internae ; middle tarsus a little stout as in
insidiosus (Fig. 7) ; hind femur (Fig. 8) a little thinner than in insidiosus, length : breadth
about 3-9 : i (in insidiosus about 3-6 : i) ; hind coxa closely punctate, with no scopa.

Gaster with the central area of post-petiole with the striae rather close and rather regular ;
tergite 2 with moderately broad thyridiae (breadth : distance between thyridiae about i : 1-25),
rather closely punctate, in part striate between the gastrocoeli and centrally, tergite 3 basally
with the punctures comparable with those at the apex of tergite 2, becoming sparser apically,
but still clearly impressed.

FIGS. 6-8. Ichneumon quartanus, $ ; fig. 6, Antennal segments 3 and 4
fig. 7, middle tarsus ; fig. 8, hind femur.

Colour : head black, inner orbits sometimes yellow or red ; scape and pedicel black, segments
3-8 of antenna red, 9-15/16 ivory, the following segments fuscous or black ; thorax black,
scutellum largely ivory ; gaster with the tergites black, the post-petiole in part and tergites
2-3 red (3 sometimes fuscous apically) ; tergites 4-7 with an apical ivory spot, that on 4 being
narrower than the spot on 5 ; coxae and trochanters black, trochantelli red ; femora red with
the hind femur broadly black dorsally and apically ; tibiae and tarsi red, the hind tibia narrowly
infuscate apically and the apical hind tarsal segments infuscate ; wings yellowish clouded,
costal vein and stigma pale testaceous.

Length : 12 mm.
. Unknown.

HOLOTYPE. $. Hym. 3b 1821. ENGLAND
20. vi. 1937 (A. E. Wright}. (B.M.Coll.)

PARATYPE. $. SCOTLAND : Inverness ; Nethy Bridge,
(B.M.Coll.)

Westmorland ; Langdale Pikes,
1931 (P. Harwood).

Ichneumon caproni sp. nov.

Closely related to Ichneumon nereni Thomson and to minutorius Desvignes (=
captorius Thomson). (I have examined the type or syntypes of all these species.)
It differs from both these species in having a thicker hind femur and in having the

NOTES ON BRITISH ICHNEUMONIN AE

head less sharply and more convexly narrowed behind the eyes ; the post-petiole
is less strongly striate as in nereni, but has no white spot at the apex of tergite 5 (in
the seven examples seen), whereas in nereni it is most usually present.

$. Head convexly narrowed behind the eyes (Fig. 9) ; malar space long, but slightly shorter
than the post-anellus ; mandible strongly narrowed to the apex, the lower tooth distinct ;
inner orbits sub-parallel centrally ; antenna with 32-34 segments, flagellum with the basal
segments elongate, distinctly widened beyond the middle and attenuate apically ; post-anellus
length : breadth about 2:1; ninth antennal segment about as long as broad ; frons rugose-
punctate above, with microsculpture ; face punctate, striate-punctate beneath the antennal
sockets, and with sparser, coarser punctures on the epistoma, clypeus basally with the punctures
similar to those on the epistoma, at most with a few scattered punctures in the apical half.

II

12

FIGS. 9-12. Head dorsally, hind femur laterally : figs. 9 and n, Ichneumon caproni,

figs. 10 and 12, I. nereni, $.

Thorax similar in sculpture to nereni, though the punctures on the whole are a little smaller
and closer ; propodeum with the area superomedia a little transverse, the area petiolaris not
margined laterally, punctate or punctate-rugose, becoming rugose, narrowly, dorsad, but
clearly differentiated from the more coarsely rugose areae internae ; front and middle tarsi a
little broader than in nereni ; the hind femur a little incrassate, length : breadth about 3-5:1
(Fig. n) ; hind coxa a little more finely and a little more closely punctured than in nereni,
with no scopa.

Gaster with the post-petiole rather finely striate ; tergite 2 rather coarsely punctate, some-
what striate between the gastrocoeli and centrally, the thyridiae rather narrow (breadth :
distance between thyridiae about i : 1-75) ; tergite 3 basally, similar in sculpture to the apical
half of tergite 2, the punctures very shallow or erased apically.

Colour : head black, inner orbits red, or yellow or at least with pale marks on the frontal
orbits ; scape black, sometimes marked with red, pedicel fuscous ; antennal segments 3-8 red,
sometimes marked with fuscous, segments 8/9-14/15 ivory, the following segments fuscous or
black ; thorax black, the sub-alar callus sometimes marked red, scutellum for the greater part
ivory ; gaster with the tergites black, except the apex of the post-petiole marked with red ;
tergites 2-3 red, sometimes marked with yellow, 6 and 7 with a large, ivory, apical spot ; legs

148 NOTES ON BRITISH ICHNEUMONIN AE

with the coxae and trochanters black, the trochantellus red ; front and middle femora red,
broadly marked with black ; hind femur black narrowly red basally ; front and middle tibiae
and tarsi testaceous, the apex of middle tibia and apical tarsal segments sometimes infuscate ;
hind tibia testaceous, fuscous apically ; hind tarsus fuscous or black, sometimes paler basally ;
wings very weakly smoky, the costal vein pale testaceous, fuscous apically, the stigma
testaceous.

Length: 9-11 mm.

o*. Unknown.

HOLOTYPE. ?. Hym. 3b 1822. ENGLAND : (Capron Coll.] (B.M.Coll.)
PARATYPES. ENGLAND : i$ (Capron Coll., det. as gracilicornis by Capron) ;
i$, B.M. British Coll. (det. as ammonius in Desvignes' catalogue). GERMANY : i,
Ant., 6.x (Buchecher) ; i$, Lag., 4.viii.i865 (Buchecher), France: i$, Castellane,
g.iv. 1931 (C. Morley). (B.M. Coll.) ENGLAND : i$, Capron Coll. (det. as gracilicornis
by Capron). (C. Morley Coll.)

Ichneumon aquilonius sp. nov.

Related to vulneratorius Zetterstedt, walkeri Wesmael and polyonomus Wesmael
(this latter I regard as a subspecies of walkeri, from the Alps). It differs from vul-
neratorius in having the clypeus remotely punctate in the apical half and the central
area of the post-petiole much more strongly, longitudinally striate ; from walkeri
and polyonomus in having the hind coxa much more coarsely, though closely punctate
beneath and with no trace of a scopa ; superficially it differs from all three in having
an orange-coloured scutellum (though this may well vary). As with most of these
northern species, the colour will probably show great variation.

$. Head strongly narrowed behind the eyes ; malar space long, about i times as long as
the breadth of base of mandible ; mandible strongly narrowed to the apex, the lower tooth
distinct ; inner orbits diverging a little ventrad ; antenna with 33 segments ; flagellum with
the basal segments elongate, conspicuously widened beyond the middle and attenuate apically,
at the widest with the flattened, ventral area of the segments slightly wider than long ; post-
anellus length : breadth = 1-7:1; flagellar segment 5/6 quadrate ; frons, dorsally rugose-
punctate and in part with microsculpture on the rugae and in the interspaces between the
punctures ; face, for the most part, with clear punctures becoming rugose-punctate beneath the
antennal sockets and in part centrally ; clypeus with rather sparse punctures, very sparse in
the apical half but with an apical row of more or less distinct, rather close punctures.

Thorax similar in sculpture to vulneratorius ; propodeum with the area superomedia quadrate
or a little transverse, more or less closed posteriorly, the area petiolaris not margined laterally,
punctate dorsad, rugose ventrad and thus little differentiated in sculpture from the areae
internae ; front and middle tarsi thin, as in vulneratorius ; hind coxa closely punctate beneath
tending to rugose-punctate before the apex, with no scopa ; hind femur moderately thin as
in vulneratorius.

Gaster with the central area of the post-petiole coarsely, longitudinally striate ; tergite 2
with broad transverse thyridiae (breadth : distance between thyridiae about 1-8 : i) ; tergite 2
closely punctate, in part striate between the gastrocoeli and centrally ; tergite 3 closely punctate
basally, becoming more sparsely punctate apically.

Colour : head black with the clypeus, epistoma and inner orbits red ; scape red, fuscous
basally ; pedicel fuscous ; basal flagellar segments red or marked with fuscous, antennal seg-
ments 8/9-13/14 ivory, beyond this black ; pronotum black with the collar centrally, and
the upper margin laterally, red ; mesosternum and mesopleurum black with the sub-alar callus

NOTES ON BRITISH ICHNEUMONIN AE

149

marked with red ; mesoscutum red, more or less marked with fuscous anteriorly ; scutellum
orange ; post-scutellum red ; propodeum black ; gaster red, the 4th segment sometimes black,
the following segments black ; legs red with the coxae and trochanters black (in part, slightly
reddish in the type) ; femora sometimes conspicuously marked with black ; apex of hind
femur and apical tarsal segments sometimes marked with fuscous ; wings yellowish, the costal
vein and stigma pale testaceous.

Length : 8 mm.

o*. Unknown.

HOLOTYPE. $. Hym. 3b 1823. SCOTLAND: Perthshire; Killin, 15-21^.1932
(R. B. Benson}.

PARATYPE. $. ENGLAND: Westmorland; Fairfield, c. 2,500 ft., 5.^.1938
(G. J. Kerrich).

THYRATELES gen. nov. (Ichneumonini)

Clypeus flat ; the occiput somewhat impressed behind the ocelli ; mandibles strongly
narrowed to the apex ; pronotal collar rather narrow, the transverse groove deeply impressed ;
scutellam strongly convex ; propodeum with the basal groove distinct, the area basalis with
no central tubercle, the area superomedia more or less quadrangular, no dentiparal spines ;
post-petiole with the central area longitudinally striate or rugose-striate, and not conspicuously

FIG. 13. Thyrateles camelinus, $ : 2nd tergite.

raised at the line of the spiracles ; gastrocoeli deep and broad, thyridiae distinct (Fig. 13) ;
claws impectinate.

$ with the hypopygium as in A mblyteles ; the flagellum strongly elongate.

Type species Amblyteles camelinus Wesmael.

This genus belongs to that group represented in the Holarctic region by Patroclus,
CtenamUytdes (syn. Pseudambly teles) and Hybophorellus. All these genera have

150

NOTES ON BRITISH ICHNEUMONIN AE

females in which the hypopygium is like Ambly teles, but which have conspicuous and
often large thyridiae. A few species in Amblyteles have small thyridiae, but Cteni-
chneumon, with which certain of the above genera have previously been confused
in Europe, has the thyridiae completely absent. Both Patroclus (to which the
European Ctenichneumon sputator (F.) belongs) and Ctenamblyteles have the claws
pectinate at least towards the base (this pecten is not to be confused with the row
of oblique, contiguous, basal teeth). Hybophorellus has shallow gastrocoeli, a
strongly convex clypeus and dentiparal spines. Amongst the material in the
British Museum I have noted no species outside this area which would be referable
to this group. Descriptions most usually fail to mention the presence or absence
of thyridiae which seem important in this section of the Ichneumoninae.

Spilichneumon celenae sp. nov.

This species belongs to the section of Spilichneumon having, in the female, robust
mandibles with the lower tooth small and much shorter than the enlarged upper
tooth, the hypostomal carina conspicuously raised and the front and middle tibiae
coarsely spinulose.

16

FIGS. 14-17. Antennal segments 2 and 3 : fig. 14, Spilichneumon celenae, $ ; fig. 15,
S. simplicidens, $. Mandible, fig. 16, and head dorsally, fig. 17, 5. celenae, $.

$. Head with the temples distinctly narrowing behind the eyes (Fig. 17) ; malar space a
little shorter than the breadth of base of mandible ; mandible with the sides of the shaft sub-
parallel, only very weakly expanded beyond the middle, the upper tooth larger than the lower
(Fig. 1 6) ; antenna very little widened beyond the middle, strongly attenuate apically, with
36 segments, the post-anellus short, length : breadth about 1-2:1 (Fig. 14), the 4th antennal
segment sub-quadrate ; frons dorsally, coarsely punctate and weakly coriaceous ; face rather
coarsely punctate, somewhat striate dorsad and with the punctures becoming finer and sparser
towards the malar space ; clypeus (which is relatively a little smaller than in stagnicola) rather
coarsely punctate, the punctures more sparse centrally, towards the apex.

Thorax : sculpture similar to that of stagnicola ; pronotal collar short ; propodeum with the
strongly elongate area superomedia open behind, the lateral carinae erased posteriorly and the
area petiolaris divided from the areae internae ; front and middle tarsis a little expanded
(mid tarsus with segment 2 about twice as long as broad), hind femur thickened (length : breadth
about 3-2 : i).

Caster with the post-petiole finely striate ; tergite 2 with small and very shallow gastrocoeli,
thyridiae absent, rather finely, rather sparsely punctate, the punctures sparser apically, only
striate at the extreme base ; 3rd tergite basally similar in puncturation to the apex of the 2nd,

NOTES ON BRITISH ICHNEUMONIN AE 151

apically with the punctures becoming very fine and very sparse ; 4th tergite basally similar
in puncturation to the apex of the 3rd, apically, together with the remaining segments with only
very fine punctures ; sternites 2 and 3 with a ventral fold.

Colour : head black, sometimes with the inner and vertical orbits more or less red-marked ;
antenna black or fuscous, with segments 8-13 ivory or marked with ivory ; thorax black,
pronotal collar sometimes red-marked centrally, scutellum for the greater part ivory ; legs with
the coxae black, trochanters black, narrowly yellow apically ; front and middle trochantelli
black or piceous, femora red, black basally, tibiae and tarsi testaceous, the apex of the tarsi
somewhat infuscate ; hind trochantellus red, fuscous dorsally, femur black, reddish apically
and at extreme base, tibia pale testaceous, fuscous apically, tarsus testaceous with the apices
of the segments infuscate ; gaster black with tergites 2 and 3 red, 4 basally, laterally, sometimes
red ; tergites 6 and 7 with a large, ivory apical spot ; wings with the costal vein pale testaceous,
fuscous apically, stigma pale testaceous.

o*. Head with the temples distinctly narrowing behind the eyes ; malar space about half as
long as the breadth of base of mandible ; mandible with the sides of the shaft narrowing distally ;
antenna with tyloidae on segments 5/6-17/19, that on 5 when present being weak; frons
rugose-punctate above ; face coarsely punctate, striate in part below the antennal sockets ;
clypeus coarsely punctate, weakly rounded apically ; thorax more coarsely and closely
punctured than in the female ; propodeum with the area superomedia elongate and closed
posteriorly, the lateral carinae complete, the area petiolaris divided by strong carinae from the
areae internae ; hind femur, length : breadth about 3-7 : i.

Gaster with the post-petiole more coarsely and irregularly striate than in the $ ; tergite 2
with the gastrocoeli ill-differentiated and shallow, thyridiae absent, rather closely punctured,
striate between the gastrocoeli and extending more or less to the apex of the segment ; tergite
3 sub-quadrate, a little more finely and less closely punctate than tergite 2, centrally, basally
striate-rugose ; sternites 2 and 3 with a median fold ; hypopygium with a sub-acute, central,
apical projection.

Colour : head black ; face yellow, black centrally, varying to black with the orbits yellow ;
clypeus yellow sometimes marked with black ; scape marked with yellow beneath ; thorax
black, collar sometimes red centrally, scutellum marked with yellow, tegula pale testaceous ;
coxae black, trochanters black, narrowly yellow apically ; trochantelli testaceous, fuscous
dorsally ; front and middle femora red, broadly marked with black ; front and middle tibiae
and tarsi testaceous, the tarsi in part infuscate ; hind femur black, narrowly red basally and
apically ; hind tibia pale testaceous, fuscous apically ; hind tarsus fuscous, testaceous basally ;
gaster black ; extreme apex of post-petiole, and tergites 2 and 3 testaceous ; tergite 4 testaceous
laterally, basally ; wings with costal vein pale testaceous, infuscate apically, the stigma pale
testaceous.

Length : $ 12 mm., $ u mm.

HOLOTYPE. $. Hym. 3b 1824. IRELAND : ex Celena haworthii, 1884 (W. V. de
F. Kane). (B.M.Coll.)

ALLOTYPE. $. IRELAND: Co. Wicklow; Torilagee Mt., 1-2,000 ft., 22. viii. 1937
(A. W. and G. M. Stelfox). (B.M.Coll.)

PARATYPES. SCOTLAND: i$, Sutherland; R. Traligal, Glen Dubh, 19. vi. 1948
(J. Balfour-Browne). IRELAND : i<$, same data as allotype. (B.M.Coll.) SWEDEN :
i$. (Thomson Coll., Lund University Museum).

This species is closely related to simplicidens (Thomson) and agrees with it in the
form of the mandible, but simplicidens differs in having the basal flagellar segments
more elongate (post-anellus length : breadth about 1-6 : I, 4th antennal segment
about 1-3 : i), the hypostomal carina a little less raised, the post-petiole more coarsely
striate, tergite 3 in greater part fuscous and the legs much more broadly infuscate.

ENTOM. in, 4. 12

152

NOTES ON BRITISH ICHNEUMONIN AE

The males from Skane, Ringsjon, which Thomson had associated with the single
female type of simplicidens I believe to belong to celenae.

Amblyteles (Triptognathus) propinquus sp. nov.

Much confusion has existed in Britain between Amblyteles uniguttatus (Gravenhorst)
and conspurcatus (Gravenhorst) and Ctenichneumon panzeri (Wesmael). I have now,
however, seen material from England which does belong to the uniguttatus group of
species with the mandible unidentate, but which appears to be distinct from described
species. It may be a subspecies, but this has yet been impossible to determine. I

21

FIGS. 1 8-2 1. Propodeum : fig. 18, Amblyteles conspurcatus, $ ; fig. 19, A. propinquus,
fig. 20, A. uniguttatus, $. Apex of penis valve, fig. 21, A, propinquus.

NOTES ON BRITISH ICHNEUMONIN AE 153

have also seen material of the same form in the Wesmael collection, from the dunes
near Ostend. Only the male is known to me.

cJ. More robust than uniguttatus, less robust than conspurcatus. Similar to uniguttatus, but
with the vertex deeply and more sharply impressed behind the ocelli ; mesoscutum at least in
part a little less closely punctate, the propodeum deeply excised before the dentiparal spines
(Fig. 19) (the excision a little less abrupt than in conspurcatus) ; mesopleurum with the speculum
clearly punctate (as in uniguttatus) ; thyridiae distinct but small, in breadth subequal to the
length of the 4th middle tarsal segment (in uniguttatus broader than this ; in conspurcatus
the thyridiae absent).

Colour : black ; clypeus sometimes with an ivory spot on each side ; pronotum with the
collar centrally and the mark in the hind angles ivory ; sub-alar callus and scutellum in greater
part ivory ; gaster with tergites 2-4 testaceous, 4 marked with fuscous ; apical tergites and
genital claspers not marked with ivory ; sternites 2-3 and in part 4, testaceous ; front and
middle femora red, yellow apically and sometimes dorsally, often marked with black posteriorly
and beneath, the tibiae and tarsi pale testaceous, the tarsi in part infuscate ; hind coxa some-
times with an ivory, dorsal mark, hind femur red, varying to black but most usually retaining
a yellow dorsal stripe (in the darkest specimen seen the femur is black with a reddish dorsal
stripe), hind tibia testaceous, narrowly infuscate apically, in dark specimens with the fuscous
markings more extensive, hind tarsus fuscous, usually pale at the base ; wings weakly smoky,
darker apically, with the costal vein fuscous becoming testaceous in basal , stigma testaceous,
margined with fuscous.

Penis valves similar to those of uniguttatus and conspurcatus (Fig. 21).

Length : 16-18 mm.

HOLOTYPE. <$. Hym. 30 1825. ENGLAND: Cambs ; Croydon, 17^1.1945
(C. E. Tottenham). (B.M.Coll.)

PARATYPES. ENGLAND: Cambs; 5$, Croydon, 17-23. vi. 1945. (C. E. Totten-
ham). Suffolk; Sudbury, Gull Lane, I7.vii.i922 (P. Harwood). (B.M.Coll.)

Platylabus stolidus sp. nov.

This species belongs to the group of pedatorius (F.), having the strongly expanded
hypostomal carina ; it differs from the other species of the group that are known
to me in having the temples relatively strongly enlarged.

?. Head with the temples large, sub-parallel directly behind the eyes (Fig. 22) ; frons dorsad
distinctly punctate ; face coarsely punctate, the punctures becoming finer towards the orbits
and sub-rugose on the epistoma, the inter-antennal tubercle weak ; clypeus closely punctate
basally, more sparsely punctate in apical third, little convex (conspicuously flatter than in
pedatorius) ; antenna very little widened before the apex, 39 segments, similar to that of
pedatorius ; hypostomal carina broad.

Thorax with the sculpture similar to that of pedatorius ; the epicnemia not strongly raised ;
notauli vaguely impressed anteriorly ; dentiparal area with no apical tooth ; the area supero-
media a little transverse ; area petiolaris clearly differentiated by lateral carinae, trans-striate-
rugose ; hind femur rather stout, length : breadth about 3-2 : i (of pedatorius about 3-7 : i) ;
hind tarsus with segment i a little longer than segments 2 + 3 + 4.

Gaster with segment i similar in form and sculpture to that of pedatorius ; tergite 2 with the
distance between the thyridiae a little greater than in pedatorius (thyridiae : distance between
thyridiae about 1-5 : i) and a little more oblique than in that species, punctate or rugose between
the gastrocoeli and behind the thyridiae, otherwise with clear punctures which become shallow

154 NOTES ON BRITISH ICHNEUMONIN AE

and ill-defined towards the hind margin, covered in coriaceous sculpture as are the following
segments ; tergite 3 with clear, shallow punctures basally, fading out in the apical third.

Colour : black ; malar space apically ivory ; frontal orbits marked with yellow, outer orbits
sometimes marked with yellow centrally ; antenna with segments 10/11-14/15 with an ivory
dorsal stripe, scape with a small, yellow, basal spot beneath ; mandibles red towards apex ;
tegulae sometimes marked with pale testaceous ; gaster with the apices of the tergites piceous,
tergite 2 piceous basally ; trochantelli narrowly red apically sometimes in part with piceous
markings, that of the front leg pale, sometimes testaceous beneath ; front and middle femora
and tibiae red, their tarsi fuscous, in part testaceous ; hind femur red, distinctly though rather

22

FIG. 22. Platyldbus stolidus, $ : head dorsally.

narrowly black apically, hind tibia black, usually red in basal third, hind tarsus black, the seg-
ments reddish basally and segments i and 2 with a small red apical spot ; wings with the costal
vein pale testaceous, fuscous apically and the stigma black.

Length : 9-11 mm.

cJ. Unknown.

HOLOTYPE. $. Hym. 3b 1826. ENGLAND : Essex ; Colchester (Harwood).
(B.M.Coll.)

PARATYPES. BELGIUM: i$, Steinbach, i8.viii.i875 ; i<j>, Francorchamps,
3.viii.i899 (Severiri). (Coll. Tosquinet, Brussels Museum).

Differs from pedatorius in the broader temples, the black stigma, the black apex to
the incrassate femur, the flatter clypeus.

Platylabus odiosus sp. nov.

This species belongs to the pedatorius (F.) group.

$. Head with temples strongly narrowed behind the eyes, a little convex ; frons, above,
coriaceous, with shallow, ill-defined punctures and striae ; face strongly punctate becoming
rugose-punctate on the epistoma ; the inter-antennal tubercle weak ; clypeus strongly punctate
basally, very sparsely punctate in apical third, convex ; antenna hardly widened before the
apex, with 39-40 segments ; hypostomal carina broad.

Thorax with the puncturation similar to pedatorius, but with distinct microsculpture between
the punctures on the lateral lobe ; the epicnemia not distinctly raised, notauli vaguely impressed
anteriorly ; dentiparal area with no apical tooth ; area superomedia less transverse than in

NOTES ON BRITISH ICHNEUMONIN AE

155

pedatorius ; area petiolaris clearly differentiated by lateral carinae, striate-rugose ; hind femur
with length : breadth about 3-8 : i ; hind tarsus a little thinner than in pedatorius, and with
segment i subequal to segments 2 + 3 + 4 (Fig- 2 4)-

Gaster a little narrower than in pedatorius, the ist segment similar in form to pedatorius ;
tergite 2 with thyridiae : distance between thyridiae about 2 : I (cf. Fig. 26), rugose between
the gastrocoeli, otherwise punctate, the punctures becoming very feeble apically, mostly
coriaceous, as are the following segments ; tergite 3 with clear, shallow punctures in basal half.

Colour : black ; apex of malar space ivory, frontal orbits and usually external orbits centrally,
yellow ; antenna with segments 11/12-14/15 with an ivory dorsal stripe, the scape sometimes

26

FIGS. 23-26. Hind tarsal segments 1-4 : fig. 23, Platyldbus pedatorius, $ ; fig. 24,
P. odiosus, <j>. Tergite 2 : fig. 25, P. pedatorius, <J ; fig. 26, P. odiosus, Q*.

yellowish basally, beneath ; mandibles red towards apex ; scutellum towards apex and usually
post-scutellum, marked with yellow ; tegula usually marked with yellow ; gaster with the tergites
somewhat piceous apically, tergite 2 also piceous basally ; sternites in part somewhat piceous
marked ; trochanters very narrowly reddish apically ; front and middle trochanteUi, femora,
tibiae and tarsi red, the tarsi in part somewhat infuscate ; hind trochantellus in part red,
femur red, rarely with an extremely narrow fuscous mark apically, tibia black, red in basal
third, tarsus black, with the bases of the segments usually more or less reddish and segments
i and 2 with a small, dorsal, apical, red spot ; wings with the costal vein pale testaceous, fuscous
apically, stigma black.

o*. Similar to the 9 but more slender, the punctures of the abdomen deeper and more distinct,
the ist segment of hind tarsus shorter than segments 2 + 3 + 4 ; the face and clypeus marked

156

NOTES ON BRITISH ICHNEUMONIN AE

with yellow, the antenna with no white stripe above, the scape broadly yellow beneath, the
femur narrowly fuscous apically.
Length : 6* 10 mm., $ 8-10 mm.

HOLOTYPE. $. Hym. 3b 1827. ENGLAND: Devon; Newton Abbot, 8.x. 1941
(R. C. L. Perkins). (B.M.Coll.)

ALLOTYPE. $. BRITISH ISLES (T. A. Marshall). (B.M.Coll.)

PARATYPES. ENGLAND: 4$, Cornwall; Botusfleming (T. A. Marshall); i$
(Cameron); i$ (Capron). GERMANY: 2$, Apfl., I4.viii. (Buchecher). (B.M.Coll.)

This species has the interspaces between the punctures on the mesoscutum less
strongly coriaceous than in the opaculus Thomson group, and also has the hypo-
stomal carina strongly raised as in pedatorius. It differs from pedatorius in the
sculpture of the mesoscutum, the black stigma and the different proportions of the
hind tarsus (in pedatorius $, tarsus i is distinctly longer than 2+3+4).

Cyclolabus dubiosus sp. nov.

The species related to factor (Wesmael) are most obscure, and it is possible that
more than two species (pactor and dubiosus) occur in the British Isles. In dealing
with our species I have felt it necessary, however, to distinguish this species from
pactor. I give the differences in the form of a table. I have examined the syntypes
of pactor in the Wesmael collection.

pactor (Wesmael).

$6*- Thyridiae very broad ; breadth of
thyridia : distance between thyridiae
about 1-9:1 (Fig. 28).

Scutellum at most with the lateral keels
present basally.

Area superomedia usually receiving the
costula behind, or at the middle.

$. Lateral lobes of the mesonotum,
centrally, polished between the punctures.

o*. Mesopleurum more sparsely punc-
tate.

dubiosus sp. nov.

Thyridiae narrower ; breadth of thy-
ridia : distance between thyridiae about
1-5:1 (Fig. 27).

Scutellum with the lateral keels ex-
tending to the middle, where they are
inwardly curved.

Area superomedia receiving the costula
before the middle.

Lateral lobes of the mesonotum with
surface sculpture between the punctures.

Mesopleurum in part sub-rugosely punc-
tate.

FIGS. 27, 28. Tergite 2 : fig. 27, Cyclolabus dubiosus, $ ; fig. 28, C. pactor,

NOTES ON BRITISH ICHNEUMONIN AE

157

HOLOTYPE. $. Hym. 3b 1828. ENGLAND : Essex ; Colchester, 1902 (Har-
wood). (B.M.Coll.)

ALLOTYPE. <$. ENGLAND : Essex ; Colchester, 1903 (Harwood). (B.M.Coll.)

PARATYPES. ENGLAND : Glos. ; 4^, Staunton, 4-11. 1.1936 (E. B. Britton and
J. F. Perkins}. Essex ; Colchester, 2<$ (one ex Eucymatoge subnotata) (Harwood).
Surrey ; i$, Claygate (G. C. Champion). Berks ; Ascot, i^ ex Eupithecia goossen-
siata, x.1934 (E. A. Cockayne). W. Suffolk; i<j>, bred i8.viii.i9ii (C. G. Nurse).
Somerset ; Crowcombe, i$, ex Eupithecia expallidata, 9. 1.1949 (E. C. P. C.).

I have seen material of pactor bred from Eupithecia insigniata and Eupithecia
arceuthata.

Herpestomus wesmaeli sp. nov.

Related to nasutus Wesmael, but with the head and thorax conspicuously dorso-
ventrally flattened, thus approaching the form found in Eriplatys ardeicollis (Wes-
mael), but less depressed than in that genus, and without the intumescent first
segment of the gaster.

29

FIG. 29. Herpestomus wesmaeli, $ : head and thorax laterally.

.. Head with the temples rather short but strongly convex, the vertex broadly and rather
deeply excised, and with the plane of the face almost at right angles to that of the frons, which
is not intumescent, and with very small antennal scrobes ; malar space about 0-8 times breadth
of base of mandible ; mandible with the lower tooth small and well removed from the apex
of the upper tooth ; genal carina meeting the hypostomal carina at a distance from the base
of the mandible which is less than the malar space ; frons closely punctate, tending to striate-
punctate, particularly ventrad ; vertex finely punctate, becoming more finely and more
remotely punctate on the temples ; face closely punctate, becoming more finely and more
closely punctate on the malar space and with the epistoma convex, sub-quadrate ; clypeus
with coarser punctures than the face, convex, and with a narrow flattened anterior margin ;
antenna with 24-26 segments, similar in general form to that of nasutus.

Thorax distinctly dorso-ventrally flattened (Fig. 29) ; pronotal groove rather shallow, the
collar, centrally, broader than in nasutus and with the anterior margin centrally not reflexed,
laterally with the furrow transcostate, the hind margin transcostate, punctate dorsad and in
the hind angle ; mesoscutum with very weak short notauli, clearly punctate, polished between
the punctures except on the middle lobe anteriorly ; scutellum more finely punctured than the
mesoscutum centrally, flattened ; mesopleurum punctate, somewhat striate posteriorly, ventrad,
the speculum more finely punctate ; sternaulus rather shallow, transcostate, extending to
about the middle ; mesosternum elongate, a little more finely punctate than the mesopleurum,
viewed laterally with the lateral margin almost straight from the epicnemia to the hind margin.

158 NOTES ON BRITISH ICHNEUMONIN AE

the prepectus short ; propodeum dorsally flattened with the broad area petiolaris subequal in
length to the large, almost pentagonal area superomedia, the area basalis obsolete or absent,
the costula short, strong, the area externa narrow, the area spiracularis slightly widening at the
costula ; metapleurum punctate with the clearly denned area coxalis rugose ; hind coxa weakly,
obliquely transcostate in the dorsal furrow, finely and closely punctate beneath, the punctures
becoming a little coarser and a little sparser apically ; front wing with the 2nd intercubital
unpigmented ; hind wing with the nervellus antefurcal.

Caster with ist segment rather short, similar to that of nasutus, with distinct, somewhat
coarse punctures extending almost from the base nearly to the apex and sometimes, in part,
somewhat striate ; tergite 2 with rather broad thyridiae, breadth of a thyridiae : distance
between thyridiae about 1-5 : i, gastrocoeli rather short, rugose, the segment otherwise clearly
punctate, the punctures obsolete apically, in far the greater part with no microsculpture between
the punctures ; tergite 3 with the punctures similar in size to those of tergite 2, obsolete apically ;
tergite 4 a little more finely and less deeply punctate, the punctures obsolete in apical half.

Colour : black ; usually with the epistoma yellow-marked dorsally, and the face with a
yellow spot between the antennal sockets, and the orbit, epistoma, clypeus and apex of malar
space often piceous or reddish ; mandible in part reddish ; antenna with the scape, in part,
reddish beneath, basal flagellar segments sometimes in part piceous beneath ; pronotal collar,
centrally, sometimes yellow or red marked ; tegula yellow marked posteriorly ; coxae black,
trochanters narrowly yellowish apically, trochantelli reddish, in part yellowish, rest of the legs
reddish, marked with fuscous, the hind legs more extensively fuscous than the front ones,
varying to the legs almost completely infuscate ; tergites 2-4 piceous, darker centrally, varying
to almost completely black with only the apices of the segments red.

cJ. Similar to the $ but narrower ; antenna with 23-24 segments, without tyloidae ; scape
and pedicel marked with yellow beneath, face and clypeus yellow, mandibles in greater part
yellow ; pronotal collar often marked with yellow centrally ; hind angle of pronotum usually
marked with yellow ; sub-alar callus usually marked with yellow ; tegula yellow ; front and
middle coxae and trochanters conspicuously marked with yellow, otherwise the legs similarly
coloured to $ ; tergites black with the apices of the segments (except i) narrowly yellow.

Length : Q* 4 mm., $ 4-5 mm.

HOLOTYPE. $. Hym. 3b 1829. ENGLAND: Bucks; Brickhill, S.vi
(R. B. Benson). (B.M.Coll.)

ALLOTYPE. $. Same data as type. (B.M.Coll.)

PARATYPES. 2<^ i$, same data as type (B.M.Coll.) Suffolk ; i$, Barton
Mills, fir, 5.vi.i9i6 (C. Morley}. (C. Morley Coll.) IRELAND: Co. Kildare ;
i? (A. W. Stelfox). (A. W. Stelfox Coll.)

Dicaelotus fitchi sp. nov.

This species belongs to the pumilus (Gravenhorst) group (Group A, Thomson,
1891, Opusc. Ent. 15:i6i8).

$. Head with the temples sub-parallel directly behind the eyes, strongly convex ; malar
space very short, the length about one-third of the breadth of base of mandible (Fig. 30), the
genal carina almost straight, meeting the hypostomal carina at a distance from the base of the
mandible about equal to half the breadth of base of mandible ; frons, dorsad, with the clear
punctures in the main twice as far from each other as the diameter of the punctures, the vertex
and temples with finer punctures, the frons, ventrad, trans-striate ; face with the epistoma
very short and transverse and strongly convex, divided from the clypeus by a distinct epistomal
groove ; face striate-punctate beside the sparsely punctate epistoma, and becoming sparsely

NOTES ON BRITISH ICHNEUMONIN AE

159

punctate towards the orbits ; clypeus broad, a little convex, very weakly, broadly projecting
centrally apically, basally with rather coarse, sparse punctures ; antenna with 25-26 segments ;
basal segments of flagellum very short, post-anellus sub-quadrate, segment 2 a little elongate,
3 sub-quadrate.

Thorax : pronotum centrally with only a faint indication of the transverse groove, laterally
with the furrow trans-striate (as is the posterior margin in greater part), dorsad punctate, the
collar punctate ; mesoscutum with the notauli present anteriorly, clearly punctate, polished
between the punctures except on the central lobe anteriorly ; scutellum more finely, more
closely punctate, usually with lateral carinae extending to the middle, the scutellar fovea trans-
striate ; mesopleurum coarsely punctate, striate-punctate posteriorly, ventrad, the speculum
only punctate dorsad ; sternauli extending to about two-thirds, straight, coarsely trans-striate ;
mesosternum much more finely and sparsely punctate, the posterior carina broadly interrupted
in front of the middle coxae ; propodeum similar to that of pumilus ; hind coxa punctate,
striate beneath on the inner margin, outwardly more sparsely punctate apically.

Gaster with post-petiole punctate laterally, sometimes with scattered punctures centrally,
apically, but still with a distinct, unpunctured, central area ; tergite 2 (with no gastrocoeli or
thyridiae) punctate basally (punctate-striate laterally), the punctures becoming weaker and
sparser posteriorly ; tergite 3 with the punctures, basally, similar to those in the middle of tergite

FIG. 30. Dicaelotus fitchi,

30

head from in front.

2, becoming obsolete apically ; tergite 4 with indistinct punctures in basal two-thirds ; hind
wing with the nervellus antefurcal.

Colour : black ; clypeus red apically ; mandible pale testaceous ; antenna with scape red-
marked beneath and basal flagellar segments marked with red ; pronotum usually with the
collar red-marked centrally, hind angles usually red-marked ; legs red, the coxae sometimes
infuscate basally, and the hind coxa sometimes entirely black ; femora and tibiae sometimes
fuscous marked ; hind tarsus sometimes weakly infuscate ; gaster with tergite I red laterally
id apically ; tergites 2 and 3 red with a pair of central, fuscous spots ; tergite 4 red at the
base varying to the tergites all black with only the apices of the segments narrowly red or
jiceous.

?. In general similar to the $, but the face longer with the epistoma less transverse, the
malar space even shorter (conspicuously shorter than in pumilus) , the hypostomal carina arcuately
iflexed, but less strongly so than in pumilus ; the punctures generally coarser and closer than
the $, gaster with the post-petiole evenly punctate, tergite 4 clearly punctate almost to apex ;
itenna with 29 segments, tyloidae on segments 9-16 (those on 15 and 16 small in the single
:imen seen).

Colour similar to the $, but with the scape yellow beneath, the clypeus entirely yellow, the
landible in greater part yellow (facial orbits with an indication of a yellow mark), pronotum
irith the collar centrally, and the hind angles marked with yellow, tegula yellow ; front and
xiddle coxae and all the trochanters marked with yellow.
Length : <$ 6 mm., $ 5-6 mm.

160

NOTES ON BRITISH ICHNEUMONIN AE

HOLOTYPE.
F. B. White).
with red.

ALLOTYPE.

PARATYPES.

$. Hym. 3b 1830. BRITISH ISLES : ex Depressaria apiella (Dr.
(Fitch Coll.) This is a specimen with the abdomen little marked

<$. BRITISH ISLES (Marshall). (Fitch Coll.)
2$, same data as Allotype. (Fitch Coll.) ENGLAND : Suffolk ; i<j>,
Ipswich District, g.vi.iSgy (C. Morley). (B.M.Coll.)

I have also examined 3 females which may represent a distinct species, differing
from the darkest of the above in having the tibiae fuscous, ivory basally. I have,
however been unable to discover any other reliable differences. Two are from
IRELAND: Co. Dublin; The Slade of Saggart, 8.vii.i936 (A. W. Stelfox) (A. W.
Stelfox Coll.), and the third from GERMANY (Ruthe] (B.M.Coll.). This latter
specimen is very small, being about 3-5 mm. long.

D. fitchi is superficially most similar to inflexus Thomson, but differs in the
shorter malar space, and the 4th antennal segment being relatively a little shorter.

Dicaelotus suspectus sp. nov.

This species, also, belongs to the pumilus (Gravenhorst) group.

<j>. Head with the temples sub-parallel directly behind the eyes, strongly convex ; malar
space moderate, about 0-6 times the breadth of base of mandible, the genal carina conspicuously
incurved ; frons coarsely punctate, the scrobes polished, vertex and temples with finer punctures,
which are sparser except behind the posterior ocelli ; face with the epistoma short, transverse

31

FIG. 31. Dicaelotus suspectus, $ : mesopleurum.

and strongly convex, divided from the clypeus by a strong epistomal groove ; epistoma only
punctate laterally, the face beside this striate-punctate, the punctures becoming very sparse
towards the orbits ; clypeus weakly rounded apically, a little convex, sparsely punctate
basally ; antenna with 25 segments, the post-anellus sub-equal to the following segment and
sub-quadrate, the sub-apical segments conspicuously transverse.

Thorax : pronotum centrally with a weak, transverse groove, laterally with the furrow
trans-striate, as in the posterior margin in greater part, punctate dorsad ; mesoscutum with the
notauli very short but rather deep, clearly, rather sparsely punctate, polished between the
punctures except on the median lobe anteriorly ; scutellum much more finely and more closely

NOTES ON BRITISH ICHNEUMONIN AE 161

punctate, carinate laterally to about the middle ; mesopleurum coarsely punctate anteriorly,
punctate-striate posteriorly, ventrad, the speculum finely, sparsely punctate ; sternaulus rather
broad, coarsely trans-striate, extending to about two-thirds the length of mesopleurum, and
with a coarsely trans-striate mesopleural furrow joining it, and extending forward to the epic-
nemia (Fig. 31) ; mesosternum more finely and sparsely punctate, the posterior carina at most
very narrowly interrupted at the lateral angle ; propodeum similar to that of pumilus, but the
sculpture weaker ; hind coxa punctate, the puncture sparser apically.

Gaster with the post-petiole sparsely punctate, usually punctate-striate laterally, apically ;
tergites 2 and 3 with clear punctures, becoming obsolete apically ; the following tergites rather
sparsely pubescent ; hind wing with the nervellus antefurcal.

Colour : black ; clypeus red, usually infuscate basally ; mandible with the shaft pale red ;
antenna with the scape red, infuscate dorsally, pedicel and base of flagellum testaceous or
marked with testaceous ; tegulae pale testaceous ; pronotal collar centrally sometimes marked
with pale testaceous ; legs red, marked with pale testaceous, hind coxa infuscate at least basally,
hind femur usually infuscate at least apically, hind tibia sometimes marked with fuscous apically,
apical mid and hind tarsal segments usually infuscate.

Length : 4-5 mm.

<J. Unknown.

HOLOTYPE. ?. Hym. 3b 1831. ENGLAND: Suffolk; Bentley Woods, 16 . vi . 1902
(C. Morley}. (B.M.Coll.)

PARATYPES. ENGLAND : i$ (Capron). (Morley Coll.) Surrey ; i$, Bagshot
Heath, 20. viii. 1934 (0. W.Richards}. (B.M.Coll.) SWEDEN: Skane; i$, Loderup,
22.vii.i938 (/. F. and D. M. S. Perkins). (B.M.Coll.)

This species is of considerable interest, as not only does it possess a well-developed
sternaulus, but also an oblique furrow running between this and the epicnemia, as
is found in certain Hemiteles (s.l.) . In many Porizonini it appears that the anterior
part of the sternaulus is missing, and thus only a " mesopleural furrow " is present,
such as is found in many Braconidae. This character, together with the very
narrowly interrupted posterior carina of the mesosternum, allows this species to
be readily distinguished from all others within the pumilus group.

Epitomus proximus sp. nov.

This species is very closely related to Epitomus parvus Thomson and therefore I
give the description in the form of a comparative table with that species.

proximus parvus

?o*- Genal sulcus very weakly impressed. Genal sulcus distinctly impressed.

Gena with coriaceous sculpture behind Gena polished behind the genal sulcus.
the genal sulcus.

Face with rather even coriaceous sculp- Face with a polished area above the

ture laterally. clypeus towards the genal sulcus.

Head more shallowly impressed along the Head deeply impressed along the outer

outer margin of the eye. margin of the eye.

<J. Lower frontal orbits sometimes con- Lower frontal orbits not marked with

spicuously marked with yellow. yellow.

, 18$, England, Ireland, Sweden. 30^, 73$, England, Ireland, Ger-

many, Sweden.

162

NOTES ON BRITISH ICHNEUMONIN AE

HOLOTYPE. ?. Hym. 3b 1832. ENGLAND : Glos ; Forest of Dean, High
Meadow Woods, 9^.1936. (E. B. Britton and /. F. Perkins}. (B.M.Coll.)

ALLOTYPE. <$. IRELAND: Co. Kildare ; 29.viii.i937 (A. W. Stelfox). (B.M.
Coll.)

PARATYPES. 16$, 17$, ENGLAND, IRELAND, SWEDEN. (B.M.Coll. and A. W.
Stelfox Coll.)

The form of the genal sulcus is usually a good character in separating species of
Ichneumonidae, and it is for this reason that I have retained these forms as distinct,
in spite of the paucity of other characters.

Mevesia guttata sp. nov.

Agreeing with arguta (Wesmael) in the form of the genal carina, which in both
meets the hypostomal carina directly behind the base of the mandible, the large,
deep clypeal pits, and also in having no white band on the flagellum ; it differs
from arguta in having the clypeus inflexed apically but with no flattened apical
margin, the notauli very short, the frons more weakly intumescent, the petiolar
area less strongly excavate, the genae more finely punctate ventrad, posteriorly,
the abdomen with more distinct sculpture though strongly shining and with more
distinct, large, sparse punctures ; the male is more strongly sculptured than the
female.

$. Head with the temples sub-parallel behind the eyes and then broadly rounded to the
vertical carina ; frons and vertex closely punctate and in part coriaceous, the punctures becom-
ing finer and sparse and with no microsculpture on the temples and very sparse on the genae ;

32

FIGS. 32, 33. Face to show clypeus : fig. 32, Mevesia arguta,

33

fig. 33, M. guttata, ?.

antennal scrobes trans-striate ; face with the epistoma weakly convex and slightly elongate, more
finely and more remotely punctured than the frons, the punctures becoming sparser on the face,
laterally ; clypeus convex, inflexed apically, with weak microsculpture and remotely punctate,
and clearly separated from the epistoma (Fig. 33) ; malar space coriaceous and sparsely
punctate, about half as long as breadth of base of mandible ; antenna with 19-20 segments,
the basal flagellar segments elongate, becoming sub-quadrate, quadrate at the nth segment,
the post-anellus with length : breadth about 2-5 : i.

Thorax : pronotal collar rather short, the transverse groove deep and somewhat crenulate,
the lateral groove and the hind margin striate, becoming punctate towards the upper margin and
the hind angle ; mesoscutum with the punctures similar to the vertex, but becoming close

NOTES ON BRITISH ICHNEUMONIN AE 163

anteriorly on the middle lobe, where there is microsculpture between the punctures ; notauli
deep anteriorly but shorter than in arguta : scutellum much more finely and more sparsely
punctate than the disc of the mesoscutum ; mesopleurum striate-punctate, the speculum polished,
narrowly punctate above ; sternaulus crenulate, extending nearly to the middle ; mesosternum
finely, closely, shallowly punctate ; propodeum with the basal groove narrowed centrally,
but broad on either side of the projection ; area superomedia finely rugose, little transverse,
rounded anteriorly, receiving the distinct costula at about the middle ; all the areae enclosed
except the basal area ; area interna punctate-rugose, area dentipara rugose, area spiracularis
rugose-striate, area petiolaris punctate-rugose, distinctly separated from the coarsely rugose
areae externae, moderately excised ; metapleurum with the area coxalis clearly delimited,
striate-rugose ; hind coxa with the dorsal groove obliquely striate, closely punctate laterally,
the punctures becoming sparser apically, and somewhat weakly striate on the inner margin,
beneath.

Gaster somewhat compressed apically ; with the post-petiole coriaceous, sometimes with the
coriaceous sculpture tending to run into longitudinal striae, and with very sparse, coarse punc-
tures, the extreme apex polished, no delimited central area ; tergite 2 with very broad thyridiae
which are rather narrowly separated centrally, coriaceous, with the sculpture becoming obso-
lescent apically and with scattered punctures ; the sculpture of the following tergites similar
in kind to that of the 2nd, but much weaker ; of the 5th and following tergite very weak indeed ;
hind wing with the nervellus opposite.

Colour : black ; clypeus piceous, pale testaceous apically, mandible yellow with the teeth
piceous, flagellum dull testaceous, fuscous above, basally ; pronotal collar marked with piceous
centrally and laterally, hind angle yellow marked, tegula yellow ; legs red with the coxae black,
the front coxa marked with pale testaceous beneath, the middle coxa narrowly pale marked
apically, the trochanters infuscate dorsally, the femora sometimes more or less infuscate and
the tarsi infuscate apically ; gaster with the extreme apex of tergite i piceous to pale testaceous ;
tergite 2 testaceous with sub-apical, fuscous spots or band, which laterally join the lateral
basal stripe, somewhat infuscate basally ; tergite 3 with sub-apical spots or band ; tergite
4 fuscous with a basal and apical testaceous band, the following segments more or less piceous
apically ; wings hyaline.

<J. In general similar to the $, but with the sculpture of the abdomen (which is not compressed
apically) more distinct ; malar space shorter than in the $ (about one-third breadth of base of
mandible) ; antenna with 22 segments, with rather small tyloidae on segments 9-1 1 ; propodeum
a little shorter, more coarsely sculptured and with the carina a little more strongly raised than
in the $.

Colour ; antenna black or fuscous, yellowish beneath apically ; face and clypeus yellow, the
genae not marked with yellow ; hind angle of pronotum marked with yellow, tegula yellow ;
front and middle coxae mostly yellow, trochanters and trochantelli yellow, femora pale testa-
ceous, very finely marked with light fuscous, tarsi pale testaceous, infuscate apically ; hind
coxa with a yellow apical spot, trochanter and trochantellus yellow, femur fuscous, some-
what testaceous basally, tibia and tarsus infuscate, the tibia darker basally and apically ;
gaster with the thyridiae, which extend almost completely across the segment, yellow ; tergite
I narrowly testaceous apically, tergite 2 testaceous apically, tergite 3 testaceous basally and
apically, tergite 4 testaceous apically ; the following tergites more or less testaceous apically.

Length : < 6 mm., $ 5-6 mm.

HOLOTYPE. $. Hym. 3b 1833. ENGLAND: Suffolk; Monks Soham, 8. v. 1911
(C. Morley) (B.M.Coll.) (Previously del. as Diadromus collaris.}

ALLOTYPE. 3. ENGLAND : Capron Coll. (B.M.Coll.)

PARATYPES. ENGLAND: Herts; Tring, i$, 2. .1937 (R. B. Benson). (EM.
Coll.) i$, 1052 (/. B. Bridgman). (Previously det. zsmacilentus Wesmael.) (Nor-
wich Museum.)

i6 4

NOTES ON BRITISH ICHNEUMONIN AE

PARAETHECERUS gen. nov. (Phaeogenini)
The single known species is elongate and of the general facies of Micrope Thomson.

Head with the mandibles bidentate, the shaft with the sides sub-parallel, and the lower
tooth a little smaller and not far removed from the upper tooth ; frons coriaceous with scattered
punctures, with no antennal scrobes ; face somewhat inflexed, the epistoma sub-quadrate and
clearly divided from the clypeus by a rather deep groove ; genal carina meeting the hypostomal
carina (which is not very strongly raised) at a distance from the base of the mandible ; gena not
excavate posteriorly ; clypeus strongly produced in the centre, with a wide excision in the
lower margin of this protuberance, the apical margin flattened (Fig. 34) ; the face and malar
space short ; antenna weakly clavate, the scape not expanded basally nor flared apically.

FIGS. 34, 35

34

<j+*

. Paraethecerus elongatus, $ : fig. 34, face to show clypeus ; fig. 35, tergites
i and 2 of gaster.

Thorax with a long pronotal collar ; mesoscutum with short, weak notauli ; mesosternum
flattened, the sternauli reaching to about one-third the length of the mesosternum ; posterior
carina of mesosternum narrowly, but distinctly interrupted at the lateral angle ; propodeum
with a distinct dorsal and apical face, the area superomedia closed posteriorly, and the general
sculpture of the propodeum becoming rugose ; legs with the claws simple ; wings rather narrow,
front wing with the 2nd intercubital vein present but unpigmented, hind wing with the nervellus
postfurcal.

Gaster somewhat clavate ; tergite i not intumescent and not punctate ; thyridiae large and
distinct, well removed from the base of tergite 2, the gastrocoeli undifferentiated (Fig. 35) ;
ovipositor sheaths rather broad and a little exserted ; the hypopygium, apically, well removed
from the apex of the ovipositor.

NOTES ON BRITISH ICHNEUMONIN AE 165

Only the $ known.

Type species Paraethecerus elongatus sp. nov.

Differs from Aethecerus in the form of the frons, scape and gena. In general
facies and in the form of the frons, similar to Micrope, but that genus has an evenly
convex clypeus and a more elongate and less inflexed face and longer malar space,
and a strong, complete posterior carina of the mesosternum. Paraethecerus also
has a resemblance to Centeterus and Eparces in general form, but differs in the
clypeus and in the strong thyridiae.

Paraethecerus elongatus sp. nov.

$. Head with the temples parallel behind the eyes, rather long and convex ; face coriaceous
with scattered punctures and tending to striation beneath the antennal sockets ; clypeus with
coarse punctures basally ; genae and temples in greater part coriaceous, with scattered punc-
tures ; antenna with 17-18 segments, segment 3 a little shorter than 4 (3, length : breadth about
1-8 : i ; 4, length : breadth about 2 : i).

Thorax : pronotum laterally with the groove weakly striate, evenly striate posteriorly,
coriaceous dorsad with fine, indistinct punctures ; mesoscutum clearly punctate, the punctures
sparser in the middle of the lateral lobes, with sculpture between the punctures on the central
lobe and on the disc ; scutellum more finely and less closely punctate than on the disc of the
mesoscutum, polished between the punctures ; mesopleurum more closely and more coarsely
punctate, the speculum punctate, a little more finely and less closely punctate on the meso-
sternum ; propodeum with the area superomedia pentagonal, 1-6 to 2 times as long as broad
at the junction with the more or less distinct costulae, which are received before the middle ;
the lateral carinae erased at least posteriorly ; metapleurum coarsely, very closely punctate,
the coxal area not differentiated ; petiolar area not or rather weakly differentiated from the
areae laterales ; hind femur about 3-8 times as long as broad ; hind coxa punctate, the punctures
sparser apically, clear on the lower, inner margin, the dorsal groove coriaceous.

Gaster with the tergites coriaceous, the sculpture becoming very weak on tergites 5-7, tergites
2-4 with scattered punctures ; post-petiole with the central area more or less indicated.

Colour : black ; clypeus in part red, mandibles in greater part pale testaceous, antenna
testaceous, infuscate apically, sometimes also fuscous marked above, basally ; pronotum some-
times with the collar red marked, hind angles sometimes red marked, apex of prosternum some-
times red marked, tegula pale testaceous ; legs red, coxae sometimes infuscate basally, middle
and hind tibiae narrowly infuscate basally and apically, hind femur infuscate apically, the
tarsi with the apical segment infuscate.

Length : 5 mm.

<J unknown.

HOLOTYPE. $. Hym. 3b 1834. ENGLAND: Suffolk; Brandon, 7^1.1903
(C. Morley). (B.M.Coll.) (det. as Aethecerus longulus Wesmael by C. Morley.)
PARATYPE. $. GERMANY (Ruthe). (B.M.Coll.)

Phaeogenes foveolatus sp. nov.

This species belongs to the Phaeogenes fulvitarsis group (section F. of Thomson),
and is most closely related to fulvitarsis Wesmael (syn. ruficoxa Thomson) and rusti-
catus Wesmael (syn. fulvitarsis auctt. angl.) ; it is probably the species that Thomson
determined as rusticatus. It differs from fulvitarsis in having the inner tooth of
the carina of the hind coxa arising distinctly before the hind margin, the 4th

1 66

NOTES ON BRITISH ICHNEUMON IN AE

segment of the antenna a little shorter than the distance between the posterior
ocelli, the flagellum stouter, and the notauli at most faintly indicated behind the
pit on the anterior margin of the mesoscutum.

$. Head with the temples roundedly narrowed behind the eyes ; hind ocellus about twice
as far from the occipital carina as its greatest diameter ; frons with the antennal scrobes weakly,
obliquely striate, clearly punctate dorsad and on the vertex, the punctures sparser on the temples ;
epistoma convex, not sharply differentiated from the rest of the face, punctate, the punctures
becoming sparser towards the orbits ; gena more clearly punctate than the temples, malar
space in part coriaceous and punctate, about half as long as breadth of base of mandible ; cly-
peus convex, with a sub-apical ridge which is interrupted centrally by punctures and coriaceous

36

37

FIGS. 36-39. Basal antennal segments : fig. 36, Phaeogenes foveolatus, <j> ; fig. 37,
P. fulvitarsis, . Hind coxa of P. foveolatus, $ : fig. 38, lateral ; fig. 39, ventral.

sculpture (as in all species of this group) ; hypostomal carina strongly raised, the genal carina
meeting this at a distance behind the base of the mandible less than the length of the malar
space ; gena not excavate ; antenna somewhat stout apically, the post-anellus at most equal
to the distance between the posterior ocelli (Fig. 36) ; antenna with 23-25 segments (24 in 9
out of 14 specimens).

Thorax : pronotal collar well developed, the transverse groove distinct, the lateral groove
and the lower part of the hind margin striate, above this rather coarsely punctate ; notauli
represented by a deep anterior pit, behind this rarely with a very short, hardly traceable impres-
sion ; mesoscutum with rather coarse, clear punctures on the disc, more finely and sparsely
punctate on the lateral lobes ; scutellum a little more finely punctate than the disc of the meso-
scutum, more coarsely punctate than the lateral lobes ; mesopleurum punctate, becoming
punctate-striate behind, the speculum for the most part with sparse, rather coarse punctures ;
sternauli shallow and inconspicuously crenulate, hardly extending to half ; mesosternum more
finely punctured than the pleurum ; propodeum with all the areae distinct, the area superomedia
elongate, hexagonal, receiving the costula well before the middle, weakly rugose ; area interna

NOTES ON BRITISH ICHNEUMONIN AE 167

and area dentipara weakly rugose and punctate, area petiolaris trans-striate-punctate, aera
externa rugose, area spiracularis striate-punctate ; metapleurum punctate, striate in part ;
hind coxa with an oblique, sub-apical carina, which is very weakly, broadly raised externally,
and more sharply raised internally, the area before the carina very finely punctate with only a
faint indication of oblique striation (Figs. 38-39).

Gaster with the post-petiole weakly coriaceous in part, sometimes in part longitudinally
striate, and sometimes with a few scattered punctures ; thyridiae broad, and about as far from
the base of the 2nd tergite as their length ; tergites 2 and 3 coriaceous with few, widely scattered
punctures ; tergite 4 similarly but more weakly sculptured, the following segments becoming
progressively smoother ; hind wing with the nervellus opposite, or slightly postfurcal.

Colour : black ; mandibles pale testaceous except for the teeth ; antenna with the scape and
pedicel black, varying to broadly testaceous beneath ; 3 or 4 basal flagellar segments testaceous ;
a white stripe usually on antennal segments 9-12, sometimes reduced, and in 2 German speci-
mens on segments 8-12 ; legs red, pale testaceous in part, the hind femur and sometimes the
hind coxa infuscate apically, the hind tibia rather narrowly infuscate basally and apically, and
the apical tarsal segments infuscate ; gaster with tergite i narrowly pale apically ; tergites 3-5
entirely red, the following segments narrowly piceous apically.

(J. Similar to the $ in general sculpture, and also with the notauli only clearly represented by
the anterior pit ; antenna with 27 segments, tyloidae on segments 10-13/14, the basal segments
a little shorter than in rusticatus ; clypeus marked with yellow ; legs similar in colour to those
of the $ but the coxae and hind trochanter can be extensively black marked, and the hind
tarsus is fuscous, with the basal segments narrowly pale basally ; gaster with only the apices
of the central segments narrowly testaceous, and more strongly coriaceous and punctate than
in the $.

Length : (J 5 mm. ; $ 5-6 mm.

HOLOTYPE. $. Hym. 3b 1835. ENGLAND: Surrey; Boxhill, 9. v. 1891
(T. R. Billups}. (B.M.Coll.)

ALLOTYPE. <$. ENGLAND: Surrey; Horsley, 24^11.1949 (/. F. Perkins)
(B.M.Coll.)

PARATYPES. ENGLAND: Kent; Deal, 4$, 2.viii.i88o; i$, 8.viii.i88o
(T. R. Billups) ; Surrey ; i,^, same data as allotype. SCOTLAND : Inverness ;
Aviemore, i$, 4. vi. 1952 (R. B. Benson). IRELAND : Co. Dublin ; Slade of Saggart,
i$, 4 . x . 1936 (A . W. Stelfox} ; Co. Wicklow ; Dunran, i$, 25 . v . 1937 (A . W. Stelfox) .
SWEDEN: Skane ; Ringsjon, i$, 4^.1938 (/. F. Perkins). GERMANY: 3$
(Riithe). (B.M.Coll.) FINLAND: Kuusamo ; Paanajarvi Rajala, i$, 5.vii.i935,
beaten from Picea (G. J. Kerrich). (Helsinki Museum.)

Phaeogenes (Proscus) coriaceus sp. nov.

Similar to elongatus Thomson, but differing in the more coarsely coriaceous
sculpture of the gaster, with only sparse punctures on the 2nd tergite.

?. Head with the temples slightly diverging directly behind the eyes, rather strongly convex ;
frons and vertex coarsely punctate, the punctures becoming finer on the temples, the antennal
scrobes trans-striate ; face with the epistoma convex, punctate at least laterally and dorsad, the
rest of the face punctate with the punctures becoming coarser and sparser towards the orbits ;
clypeus strongly transverse, sparsely punctate ; hypostomal carina narrow ; lower tooth of
mandible a little smaller and shorter than the upper ; antenna with 20-22 segments, the post-
anellus very slightly shorter than the following segment.

ENTOM. Ill, 4. 13

1 68

NOTES ON BRITISH ICHNEUMONIN AE

Thorax : pronotum with a broad collar (Fig. 40) centrally with the transverse groove very
shallow, the lateral furrow irregularly trans-striate, the posterior margin more or less striate,
strongly punctate dorsad ; mesoscutum. with the notauli very short, rather coarsely punctate,
polished between the punctures except on the middle lobe anteriorly ; scutellum much more
finely punctate than the disc of the mesoscutum ; mesopleurum coarsely punctate, punctate-
striate anteriorly, ventrad, the speculum with punctures only along the posterior margin ; stern-
aulus crenulate extending almost halfway to the posterior carina ; mesosternum more finely
punctate, the posterior carina broadly interrupted before the lateral angle ; propodeum (Fig. 41)
with the area superomedia elongate, hexagonal, receiving the costula well before the middle ;
all the areae clearly differentiated ; basal area with the tubercle distinct, punctate basally
becoming rugose laterally and apically, the area superomedia weakly or very weakly, irregularly
trans-striate, the area petiolaris punctate-trans-striate and the area coxalis punctate ; hind
coxa with the dorsal furrow almost devoid of sculpture, rugose dorsally at the base, punctate
outwardly and beneath, the punctures becoming a little coarser and sparser apically and finer

FIGS. 40, 41. Phaeogenes coriaceus, $ : fig. 40, thorax dorsally to show
pronotal collar ; fig. 41, propodeum laterally.

on the inner margin, unarmed ; hind femur about three times as long as broad ; wings with the
nervellus postfurcal.

Gaster with the post-petiole more or less coriaceous and longitudinally striate ; tergites 2 and
3 very strongly coriaceous, 2 with sparse punctures, 3 with punctures closer, basally ; tergite 4
a little less strongly coriaceous and with the punctures similar, though weaker, to those on 3 ;
the following tergites weakly coriaceous, 5 and 6 weakly punctate basally.

Colour : black ; face usually with a yellow spot on the lower, outer part of the antennal
socket, sometimes also marked with red laterally ; clypeus red apically, often ivory in the lateral
angles ; mandibles in large part pale testaceous ; scape sometimes reddish beneath, basal
flagellar segments more or less marked with red, segments 1 1 and 1 2 sometimes white marked ;
thorax with the pronotal collar more or less marked with red or yellow centrally, tegula pale
testaceous ; legs red, with the front and /or middle coxae infuscate basally, hind coxa usually
black basally ; the trochanters often in part more or less yellow, hind trochanter sometimes
infuscate ; hind femur infuscate apically, hind tibia infuscate at base and apex ; tarsi with the
apical segment infuscate and on the hind leg usually with all the segments more or less fuscous

NOTES ON BRITISH ICHNEUMONIN AE 169

marked ; wings hyaline, very weakly smoky apically, the stigma black ; gaster with the apex
of tergite i and tergites 2-4 red, the following tergites piceous apically.

<J. Similar to the $>, but with the temples less broad, the face longer and the epistoma hardly
differentiated, the malar space shorter ; antenna with 26 segments, segments 7-15 with large
tyloidae ; thorax less elongate ; hind femur a little thinner (length : breadth about 3-4 : i) ;
gaster with the post-petiole shining, more rugose.

Face, clypeus and malar space yellow ; scape and pedicel yellow marked beneath, flagellum
fuscous, yellowish beneath ; front and middle coxae and trochanters yellow, the femora and
tibiae marked with yellow ; hind coxa black basally though red to yellow apically, hind trochan-
ter and trochantellus yellow, hind femur not fuscous apically ; gaster with tergites 2, 3 and 4
black or fuscous marked dorsally.

Length : <J 7 mm., <j> 5-7 mm.

HOLOTYPE. $. Hym. 3b 1836. ENGLAND : Hants ; New Forest, Denny, on
dead beech 6.vi.iQ34 (C. Morley). (B.M.Coll.) (Aethecerus dispar Wesmael det.
C. Morley.)

ALLOTYPE. $. ENGLAND : Hants ; New Forest, Denny wood, on dead beech,
7.vii.i940 (C. Morley). (B.M.Coll.) (Centeterus opprimator Gravenhorst det.
C. Morley).

PARATYPES. ENGLAND: Hants; New Forest, i$, i7.vii.38 (Phaeogenes suspi-
cax det. C. Morley) ; i$, 9^.1934 (Aethecerus dispar det. C. Morley) (C. Morley).
(C. Morley Coll.) GERMANY : i$, (Ruthe). (B.M.Coll.)

XII. CHECK LIST OF BRITISH ICHNEUMONINAE

The changes to the British List of Ichneumoninae, which have been dealt with
in the preceding pages, are here brought together. So many points remain unsolved
that this list can, at most, be considered as a basis for further change. There are
also certain matters which should be made clear. I have used Ambly teles in the
sense of Thomson and have not accepted the segregates that are recognized by
Heinrich. I have examined the male genitalia of those European and North
American species which are available to me for dissection and find that in some
cases these show quite striking group characters ; however, in certain of these
groups I have so far been unable to correlate the differences with other satisfactory
characters, so I prefer to place the species in one genus as they all have a number
of characters in common, and to wait for a much wider investigation before accepting
the segregates. It should, perhaps, be pointed out that under Triptognathus other
workers have assembled two quite divergent elements ; and that the male genitalia
of Ambly teles subsericans and elongatus are essentially similar, though showing also
the only reliable differences of which I know for separating the males of these two
species, so that either both have to be included in Limerodops or both retained in
Ambly teles.

Certain placements of genera within tribes are obviously unsatisfactory ; Hypo-
mecus, Apaeleticus, Goedartia, Hepiopelmus and Acolobus I have retained in the
tribes in which they have been placed by recent authors ; Tricholabus, which seems
to me to be a completely anomalous genus, I have included in the Ichneumonini.
The Phaeogenini, too, contains very diverse elements, and are grouped together on

1 7 o

NOTES ON BRITISH ICHNEUMONIN AE

very superficial characters. I have also retained the Heresiarchini as a separate
tribe ; if these are placed in the Protichneumonini as Heinrich has proposed, then
this latter name will be a synonym of Heresiarchini, though the position concerning
the names of the higher categories has not yet been completely clarified by the
International Commission on Nomenclature.

It will be seen that a few dates, particularly earlier ones, differ from those given
in Kloet and Hincks' list ; in these cases I have followed the findings of Sherborn.
In addition to this, I have accepted the date of Holmgren's fascicle on Phaeogenini
as being 1889 as given by Kloet and Hincks, and not 1890 as quoted by Dalla Torre
and on the title-page to the complete volume of Holmgren's Ichneumoninae ;
Sharp, in the Zoological Record, decided, on the evidence available to him, that
1889 was correct.

TROGINI

TROGUS Panzer, 1806.
lapidator (Fabricius, 1787).

PSILOMASTAX Tischbein, 1868.

CERCODINOTOMUS Uchida, 1940.
pyramidalis Tischbein, 1868.

pictus (Kriechbaumer, 1882).

CALLAJOPPA Cameron, 1903.
cirrogastra (Schrank, 1781).
exaltatoria (Panzer, 1804).

PR O TICHNE UMONINI

PROTICHNEUMON Thomson, 1893.
pisorius (Linnaeus, 1758).
coqueberti (Wesmael, 1848).
erythrogaster (Stephens, 1835).
nee (Gmelin, 1790).

AMBLYJOPPA Cameron, 1902.
fuscipennis (Wesmael, 1844).
proteus (Christ, 1791).

laminatoria (Fabricius, 1798).

COELICHNEUMON Thomson, 1893.
cyaniventris (Wesmael, 1859).
desinatorius (Thunberg, 1822).

sitbguttatus (Gravenhorst, 1829).
leucocerus (Gravenhorst, 1820).
falsificus (Wesmael, 1844).
auspex (Mueller, 1776).
nigerrimus (Stephens, 1835).

derasus (Wesmael, 1844).
bilineatus (Gmelin, 1790).
haemorrhoidalis (Gravenhorst, 1820).

truncatulus (Thomson, 1886).
solutus (Holmgren, 1864).
impressor (Zetterstedt, 1838).
consimilis (Wesmael, 1844).
purpurissatus Perkins, 1953.

nigrator (Fabricius, 1793).

nee (Mueller, 1776).
comitator (Linnaeus, 1758).

ferreus (Gravenhorst, 1829).
microstictus (Gravenhorst, 1829).
serenus (Gravenhorst, 1820).
ruficauda (Wesmael, 1844).
orbitator (Thunberg, 1822).

HERESIARCHINI

HERESI ARCHES Wesmael, 1859.
eudoxius (Wesmael, 1844).

LIS TRODROMINI

NEOTYPUS Foerster, 1868.
nobilitator (Gravenhorst, 1807).

LISTRODROMUS Wesmael, 1844.
nycthemerus (Gravenhorst, 1820).

ANISOBAS Wesmael, 1844.
cingulatorius (Gravenhorst, 1820).
platy stylus Thomson, 1888.

ICHNEUMONINI

HOPLISMENUS Gravenhorst, 1829.
bidentatus (Gmelin, 1790).

spinosus (Morley, 1903).

maurus (Marshall, 1873).

bispinatorius auctt. angl.

NOTES ON BRITISH ICHNEUMONIN AE

171

Ichneumonini cont.

albifrons Gravenhorst, 1829.
armatorius (Fabricius, 1787).
nee (Forster, 1771).

STENICHNEUMON Thomson, 1893.
culpator (Schrank, 1802).
militarius (Thunberg, 1822).
rufinus (Gravenhorst, 1820).
lineator (Fabricius, 1781).

trilineatus (Gmelin, 1790).
scutellator (Gravenhorst, 1829).

AOPLUS Tischbein, 1874.
alienator (Wesmael, 1855).
defraudator (Wesmael, 1844).
castaneus (Gravenhorst, 1820).
mtzeburgii (Hartig, 1838).
virginalis (Wesmael, 1844).
ochropis (Gmelin, 1790).
rubricosus (Holmgren, 1864).
ruficeps (Gravenhorst, 1829).
humilis (Wesmael, 1857).

PLATYLABOPS Heinrich, 1950.
apricus (Gravenhorst, 1820).

semirufus (Desvignes, 1856).
pulchellatus (Bridgman, 1889).

CRATICHNEUMON Thomson, 1893.
magus (Wesmael, 1855).

nitidus (Bridgman, 1886).
varipes (Gravenhorst, 1829).

fallax Habermehl, 1923.
jocularis (Wesmael, 1848).

dissimilis auctt. angl.

punctifrons (Holmgren, 1864).
rufifrons (Gravenhorst, 1829).
sicarius (Gravenhorst, 1829).
foersteri (Wesmael, 1848).
luleiventris (Gravenhorst, 1820).
versator (Thunberg, 1822).
nigrilarius (Gravenhorst, 1820).
albifrons (Stephens, 1835).

gravenhorstii (Boyer de Fonscolombe, 1847).
nee (Wesmael, 1836).
nee (Guerin-MeneVille, 1838).
fabricator maculifrons (Stephens, 1835).
infidus (Wesmael, 1848).

liostylus (Thomson, 1887).
culex (Mueller, 1776).
clarigator (Wesmael, 1844).
fugitivus (Gravenhorst, 1829).
corruscator (Linnaeus, 1758).

semirufus (Gravenhorst, 1820).
pseudocryptus (Wesmael, 1857).
albilarvatus (Gravenhorst, 1820).
lanius (Gravenhorst, 1829).

EUPALAMUS Wesmael, 1844.
wesmaeli Thomson, 1886.
lacteator (Gravenhorst, 1829).

ERISTICUS Wesmael, 1844.
clericus (Gravenhorst, 1829).

BARICHNEUMON Thomson, 1893.
anator (Fabricius, 1793).
gemellus (Gravenhorst, 1829).

carri Habermehl, 1923.
albilineatus (Gravenhorst, 1820).

albolineatus (Gravenhorst, 1829).
bimaculatus (Schrank, 1776).
saturatorius (Linnaeus, 1758).
monostagon (Gravenhorst, 1820).
maculicauda Perkins, 1953.

per scrutator (Wesmael, 1844).

nee (Thunberg, 1822).
dumeticola (Gravenhorst, 1829).
faunus (Gravenhorst, 1829).
albosignatus (Gravenhorst, 1829).
digrammus (Gravenhorst, 1820).

nudicoxa (Thomson, 1888).
sanguinator (Rossi, 1794).
leucocheilus (Wesmael, 1844).
peregrinator (Linnaeus, 1758).
heracleanae (Bridgman, 1884).
praeceptor (Thunberg, 1822).

bilunulatus auctt. angl.
derogator (Wesmael, 1844).
chionomus (Wesmael, 1844).
plagiarius (Wesmael, 1848).
deceptor (Scopoli, 1763).

vestigator (Wesmael, 1844).

nee (Thunberg, 1822).
lepidus (Gravenhorst, 1829).
ter 'genus (Gravenhorst, 1820).
callicerus (Gravenhorst, 1820).
coxiglyptus Heinrich, 1951.

incubitor auctt. angl.
citator (Thunberg, 1822).
basiglyptus (Kriechbaumer, 1890).
locutor (Thunberg, 1822).
ridibundus (Gravenhorst, 1829).

CHASMIAS Ashmead, 1900.
motatorius (Fabricius, 1775).
paludator (Desvignes, 1854).
paludicola (Wesmael, 1857).

172

NOTES ON BRITISH ICHNEUMONINAE

Ichneumonini cant.

ICHNEUMON Linnaeus, 1758.
lugens Gravenhorst, 1829.
sarcitorius Linnaeus, 1758.
lautatorius Desvignes, 1856.
xanthorius Forster, 1771.
deliratorius Linnaeus, 1758.
molitorius Linnaeus, 1761.
crassifemur Thomson, 1886.
melanotis Holmgren, 1864.

macrocerus Thomson, 1886.
confusor Gravenhorst, 1820.

confusorius Gravenhorst, 1829.
bucculentus Wesmael, 1844.
septentrionalis atrifemw Perkins, 1953.
ligatorius Thunberg, 1822.
suspiciosus Wesmael, 1844.
extensorius Linnaeus, 1758.

militaris auctt. angl.
gracilentus Wesmael, 1844.
albiger, Wesmael, 1844.

tempestivus Holmgren, 1864.
validicornis Holmgren, 1864.
albicollis Wesmael, 1857.
stramentarius Gravenhorst, 1820.
terminatorius Gravenhorst, 1820.
computatoriits Mueller, 1776.
eurycerus Thomson, 1890.
fuscatus Gravenhorst, 1829.
rufidorsatus Bridgman, 1887.
memorator Wesmael, 1844.
latrator Fabricius, 1781.
subquadratus Thomson, 1887.
analis Gravenhorst, 1829.
spurius Wesmael, 1848.
primatovius Forster, 1771.
bellipes Wesmael, 1844.
didymus Gravenhorst, 1829.
tuberculipes Wesmael, 1848.
megapodius Heinrich, 1949.
formosus Gravenhorst, 1829.

obsessor Wesmael, 1844.

haglundi Holmgren, 1864.

microcephalus Stephens, 1835.
quartanus Perkins, 1953.
gracilicornis Gravenhorst, 1829.
emancipatus Wesmael, 1844.
exilicornis Wesmael, 1857.
caloscelis Wesmael, 1844.
insidiosus Wesmael, 1844.
minutorius Desvignes, 1856.

captorius Thomson, 1887.

nee Thunberg, 1822.
caproni Perkins, 1953.

nereni Thomson, 1887.
ignobilis Wesmael, 1855.
aquilonius Perkins, 1953.
vulneratorius Zetterstedt, 1838.
walkeri Wesmael, 1848.
haereticus (Wesmael, 1854).
cessator Mueller, 1776.

THYRATELES Perkins, 1953.
camelinus (Wesmael, 1844).

PROBOLUS Wesmael, 1844.
culpatorius (Linnaeus, 1758).
concinnus Wesmael, 1853.

CTENICHNEUMON Thomson, 1894.
edictorius (Linnaeus, 1758).
divisorius (Gravenhorst, 1820).
inspector (Wesmael, 1844).
rubroater (Ratzeburg, 1852).

melanocastani4S auctt. angl.
nitens (Christ, 1791).
devylderi (Holmgren, 1871).

repentinus auctt. angl.
messorius (Gravenhorst, 1820).
funereus (Geoffrey, 1785).
panzeri (Wesmael, 1844).

flavocinctus (Desvignes, 1856).
castigator (Fabricius, 1793).

SPILICHNEUMON Thomson, 1894.
occisorius (Fabricius, 1793).
johansoni (Holmgren, 1871).
stagnicola (Thomson, 1888).
celenae Perkins, 1953.
raptorius (Linnaeus, 1758).
septemguttatus (Gravenhorst, 1829).

AMBLYTELES Wesmael, 1844.
punctus (Gravenhorst, 1829).

nee (Shaw, 1798).
fabricii (Schrank, 1802).
armatorius (Forster, 1771).
propinquus Perkins, 1953.
crispatorius (Linnaeus, 1758).
glaucatorius (Fabricius, 1793).
pictus (Schrank, 1776).

? labor atorius (Mueller, 1776).

vadatorius (Illiger, 1807).
pallidicornis (Gravenhorst, 1829).
pulchellus (Christ, 1791).

negatorius (Fabricius, 1793).
equitatorius (Panzer, 1786).
mercatorius (Fabricius, 1793).

NOTES ON BRITISH ICHNEUMONINAE

173

Ichneutnonini cont.

monitorius (Panzer, 1801).
quadripunctonus (Mueller, 1776).
castanopyga (Stephens, 1835).
amatorius (Mueller, 1776).
luctatorius (Linnaeus, 1758).
longigena Thomson, 1888.

cerinthius auctt. angl.
gradatorius (Thunberg, 1822).
trifasciatus (Gravenhorst, 1829).
palliatorius (Gravenhorst, 1829).
margineguttatus (Gravenhorst, 1829).
oratorius (Fabricius, 1793).
indocilis Wesmael, 1844.
subsericans (Gravenhorst, 1820).
elongatus Brischke, 1878.

subsericans var. elongatus Brischke, 1878.

LIMERODES Wesmael, 1844.
arctiventris (Boie, 1841).

EXEPHANES Wesmael, 1844.
ischioxanthus (Gravenhorst, 1829).
hilaris (Gravenhorst, 1829).

nee (Say, Jan. 1829).
occupator (Gravenhorst, 1829).
caelebs Kreichbaumer, 1890.
amabilis Kreichbaumer, 1895.

ACOLOBUS Wesmael, 1844.
albimanus (Gravenhorst, 1829).
sericeus Wesmael, 1844.

HEPIOPELMUS Wesmael, 1844.
variegatorius (Panzer, 1800).
leucostigmus (Gravenhorst, 1820).

TRICHOLABUS Thomson, 1894.
strigatorius (Gravenhorst, 1829).

EURYLABINI

GOEDARTIA Boie, 1841.
alboguttata (Gravenhorst, 1829).

EURYLABUS Wesmael, 1844.
larvatus (Christ, 1791).
torvus Wesmael, 1844.
tristis (Gravenhorst, 1829).

ZIMMERINI

ZIMMERIA Heinrich, 1933.
dirus (Wesmael, 1853).

PLATYLABINI

RHYSSOLABUS Berthoumieu, 1896.
arcticus Hell6n, 1942.

PRISTICEROS Gravenhorst, 1829.
serrarius Gravenhorst, 1829.
infractorius (Linnaeus, 1761).

PLATYLABUS Wesmael, 1844.
nigrocyaneus (Gravenhorst, 1829).
tenuicornis (Gravenhorst, 1829).
histrio Wesmael, 1855.
variegatus Wesmael, 1844.
dolorosus (Gravenhorst, 1829).
rufus Wesmael, 1844.
gigas Kreichbaumer, 1886.
transversus Bridgman, 1889.
punctifrons Thomson, 1888.
intermedius Holmgren, 1871.
opaculus Thomson, 1888.
pedatorius (Fabricius, 1793)-
obator (Desvignes, 1856).
stolidus Perkins, 1953.
odiosus Perkins, 1953.
iridipennis (Gravenhorst, 1829).
pumilio Holmgren, 1871.
rufiventris Wesmael, 1344.
vibratorius (Thunberg, 1822).
decipiens Wesmael, 1*48.
tricingulatus (Gravenhorst, 1820).
concinnus Thomson, 1888.

ASTHENOLABUS Heinrich, 1951.
STENOLABUS Heinrich, 1936.

nee Schulthess-Rechberg, 1910.
vitratorius (Gravenhorst, 1829).
latiscapus (Thomson, 1894).

CYCLOLABUS Heinrich, 1935.
nigricollis (Wesmael, 1844).
pactor (Wesmael, 1844).
dubiosus Perkins, 1953.

ECTOPIUS Wesmael, 1859.
rubellus (Gmelin, 1790).
exhortator (Fabricius, 1787).

HYPOMECUS Wesmael, 1844.
quadriannulatus (Gravenhorst, 1829).

APAELETICUS Wesmael, 1844.
inimicus (Gravenhorst, 1820).
bellicosus Wesmael, 1844.
ivclytus Wesmael, 1853.

NOTES ON BRITISH ICHNEUMONIN AE

PHAEOGENINI

RHEXIDERMUS Foerster, 1868.

ISCHNOPSIDEA Viereck, 1914.
nigricollis (Wesmael, 1844).

rufipes (Wesmael, 1848).
thoracicus (Gravenhorst, 1829).

HETERISCHNUS Wesmael, 1859.
pulex (Mueller, 1776).
rufipes auctt.

MISETUS Wesmael, 1844.
oculatus Wesmael, 1844.

HEMICHNEUMON Wesmael, 1857.
elongatus (Ratzeburg, 1852).

TRACHYARUS Thomson, 1891.
corvinus Thomson, 1891.

NEMATOMICRUS Wesmael, 1844.
tenellus Wesmael, 1844.

STENODONTUS Berthoumieu, 1896.
marginellus (Gravenhorst, 1829).

ERIPLATYS Foerster, 1868.

MELANOMICRUS Morley, 1903.
ardeicollis (Wesmael, 1844).

elliotti (Morley, 1903).

HERPESTOMUS Wesmael, 1844.
wesmaeli Perkins, 1953.
nasutus Wesmael, 1844.

furunculus Wesmael, 1844.

intermedius Wesmael, 1844.
arridens (Gravenhorst, 1829).

xanthops (Gravenhorst, 1829).
brunnicornis (Gravenhorst, 1829).

DICAELOTUS Wesmael, 1844.
(incl. DELOGLYPTUS Foerster, 1868).
pudibundus (Wesmael, 1844).
punctiventris (Thomson, 1891).
pictus (Schmiedeknecht, 1903).
parvulus (Gravenhorst, 1829).
orbitalis Thomson, 1891.
ruficoxatus (Gravenhorst, 1829).
fitchi Perkins, 1953.
erythrostomus Wesmael, 1844.
inflexus Thomson, 1891.

? rufilimbatus (Gravenhorst, 1820).
rufoniger Berthoumieu, 1896.

pumilus var. rufoniger Berthoumieu, 1896.

earner oni Bridgman, 1881.
pumilus (Gravenhorst, 1829).
morosus Wesmael, 1855.
suspectus Perkins, 1953.

EPITOMUS Foerster, 1868.
parvus Thomson, 1891.

pygmaeus (Brischke, 1890, nee i
proximus Perkins, 1953.

ORONOTUS Wesmael, 1844.
binotatus (Gravenhorst, 1829).

NOTOSEMUS Foerster, 1868.
bohemani (Wesmael, 1855).

albibuccus (Kreichbaumer, 1890).

MEVESIA Holmgren, 1889.
arguta (Wesmael, 1844).
guttata Perkins, 1953.

THYRAEELLA Holmgren, 1889.
collaris (Gravenhorst, 1829).
similis (Bridgman, 1881).

DIADROMUS Wesmael, 1844.
quadriguttatus (Gravenhorst, 1829).

rubellus (Gravenhorst, 1829).

ex parte.

nee (Gmelin, 1790).
troglodytes (Gravenhorst, 1829).
subtilicornis (Gravenhorst, 1829).
tenax Wesmael, 1844.
varicolor Wesmael, 1844.
albinotatus (Gravenhorst, 1829).
candidatus (Gravenhorst, 1829).

guttulatus (Gravenhorst, 1829).

COLPOGNATHUS Wesmael, 1844.
celerator (Gravenhorst, 1807).
divisus Thomson, 1891.

CENTETERUS Wesmael, 1844.
confector (Gravenhorst, 1829).
opprimator (Gravenhorst, 1820).

EPARCES Foerster, 1868.
grandiceps Thomson, 1891.

MICROPE Thomson, 1891.
macilenta (Wesmael, 1844).

OIORHINUS Wesmael, 1844.
pallipalpis Wesmael, 1844.

NOTES ON BRITISH I CH N E UMONI N AE

175

Phaeogenini cont.
PARAETHECERUS Perkins, 1953.
elongatus Perkins, 1953.

AETHECERUS Wesmael, 1844.
dispar Wesmael, 1844.
nitidus Wesmael, 1844.
placidus Wesmael, 1844.
discolor Wesmael, 1844.
longuhts Wesmael, 1844.

OROTYLUS Holmgren, 1889.
mitis (Wesmael, 1848).

PHAEOGENES Wesmael, 1844.
(incl. PROSCUS Holmgren, 1889).
semivulpinus (Gravenhorst, 1829).
planifrons Wesmael, 1844.
heterogonus Holmgren, 1889.
curator (Thunberg, 1822).

crassidens Thomson, 1891.
melanogonos (Gmelin, 1790).

melanogonus (Gravenhorst, 1820), emend.
ophthalmicus Wesmael, 1844.
stipator Wesmael, 1855.
maculicornis (Stephens, 1835).

scutellaris Wesmael, 1844.
invisor (Thunberg, 1822).

homochlorus Wesmael, 1844.
mysticus Wesmael, 1855.

callopus Wesmael, 1844.

tibiator (Thunberg, 1822).

nee (Gravenhorst, 1820).
fulvitarsis Wesmael, 1844.

ruficoxa Thomson, 1891.
rusticatus Wesmael, 1844.

fulvitarsis auctt. angl. ex parte.
foveolatus Perkins, 1953.
cephalotes Wesmael, 1844.
elongatus Thomson, 1891.
coriaceus Perkins, 1953.
suspicax Wesmael, 1844.
flavidens Wesmael, 1844.
modestus Wesmael, 1844.
impiger Wesmael, 1844.
ischiomelinus (Gravenhorst, 1829).
eques Wesmael, 1844.
distinctus (Bridgman, 1887).
infimus Wesmael, 1844.
fuscicornis Wesmael, 1844.
trepidus Wesmael, 1844.
bellicornis Wesmael, 1844.
osculator (Thunberg, 1822).

socius Holmgren, 1889.

ALOMYINI

ALOMYA Panzer, 1806.
semiflava Stephens, 1835.
debellator (Fabricius, 1775).

XIII. REFERENCES

CARR, L. A. 1924. Trans. N. Staffs. Fid. CL, 58 : Appendix.

DALLA TORRE, C. G. DE 1902. Catalogus Hymenopterorum 3 : pt. 2. Leipzig.

HEINRICH, G. 1926. Dtsch. ent. Z. 1926 : 256.

1929. Ibid. 1928 : 318.

1933. Mitt. zool. Mus. Berl. 19 : 159.

1934. Ibid. 20 : 1-263.

1936. Dtsch. ent. Z. 1935 : 197.

1937- Ann. Mag. nat. Hist. ser. 10, 20 : 279.

1949. Mitt, munch, ent. Ges. 35-39 : 1-127.

1950. Mitt. Sammelst. Schmarotzerbest., 27 : 4-5.

1951. Bonn. zool. Beitr., (1951) : 235-290.
LELLEN, W. 1942. Notul. ent., Helsingl. 22 : 77.

KERRICH, G. J. 1935. Trans. Soc. Brit. Ent. 2 : 38.

:OET, G. S., and HINCKS, W. D. 1945. A Check List of British Insects. Stockport.
IORLEY, C. 1902. Ent. mon. Mag. 38 : 119-121.

1903. Ichneumonologia Britannica. The Ichneumons of Great Britain ; Ichneumoninae.
Plymouth.

ROMAN, A. 1912. Zool. Bidr. Uppsala, 1 : 229-293.
1914. Ark. Zool. 9, No. 2 : 1-40.
1932. Ent. Tidskr. 53 : 1-16.

ENTOM. Ill, 4. 14

1 7 6

NOTES ON BRITISH ICHNEUMON IN AE

SCHMIEDEKNECHT, O. IQO2. Opuscula Ichneumonologica, 1. Blankenburg in Thiiringia.

1928-1932. Ibid. Suppl.-Band 1. Blankenburg in Thiiringia.

SHERBORN, C. D. 1901. Index Animalium, Part i. Cambridge.

1922-1933. Ibid., Part 2. London.

UCHIDA, T. 1940. Insecta Matsum. 15 : 9.

Note. Where the references to original descriptions are not quoted above, they may be found
by consulting Dalla Torre (1902), Morley (1903), and Schmiedeknecht (1902) and (1928-1932).

PRESENTED

1 7 DEC 1953

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

1 7 DEC 1953

ON A VERY REMARKABLE FLEA
FROM ARGENTINA

H. E. KARL JORDAN

DESCRIPTIONS OF NEW AND
LITTLE-KNOWN SIPHONAPTERA

F. G. A. M. SMIT

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 3 No. s

LONDON: 1953

ON A VERY REMARKABLE FLEA

FROM ARGENTINA COLLECTED BY

DR. J. M. DE LA BARRERA

BY

H. E. KARL JORDAN

DESCRIPTIONS OF NEW AND LITTLE-KNOWN

SIPHONAPTERA

BY

F. G. A. M. SMIT

Pp. 177-219 ; 67 Text-figures.

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 3 No. 5

LONDON : 1953

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

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

This paper is Vol. 3, No. 5 of the Entomological
series.

Issued December, 1953.

Price Ten Shillings.

ON A VERY REMARKABLE FLEA FROM
ARGENTINA COLLECTED BY DR. J. M. DE
LA BARRERA

By H. E. KARL JORDAN

(Tring, Herts)
With 5 text -figures

The rapid increase of our knowledge of the Flea fauna of the Republic of Argentina
during recent years is due to Dr. J. M. de la Barrera, Director of the Institute of
Hygiene at Buenos Aires, and we students of Siphonaptera in the Tring Department
of the British Museum (Natural History) are profoundly grateful to him for the invalu-
able collections submitted to the British Museum (Nat. Hist.) for study and report.
The latest collection, received in the autumn of 1952, contained, inter alia, a number
of interesting species from Bariloche, Nahuel Huapi, Rio Negro, where Dr. de la
Barrera has a summer residence. Among these fleas is a species which excels in
interest every other new one Dr. de la Barrera has previously discovered. When
the specimens were cleared and mounted the very conspicuous and quite unexpected
kind of distinction of this species took my breath away, and my two colleagues
reacted in the same way : incredible! The Incredible Flea is the subject of this
paper. A new species, new genus, new tribe. I name the genus Barreropsylla in
appreciation of the great services Dr. de la Barrera has rendered and is continuing
to render to the study of Fleas, an order of parasites so closely linked to his researches
on the diseases of the wild mammals of his country. I expect that students of
Siphonaptera will be as astonished at the somewhat spectacular distinctions of the
species as we were, and that the extraordinarily wide gap, in some somatics, between
Barreropsylla excelsa and the related species will arouse interest beyond the sphere
of Siphonapterology. A short survey of the main characteristics will be sufficient
to prove that the species provides an ample field of meditation for the taxonomist
and geneticist In evaluating the characteristics of B. excelsa we must bear in
mind that the species, apart from its special features, is an American Stephanocircid
fitting well into the subfamily of Craneopsyllinae ; there is no doubt about that.
Confining the discussion, for the sake of brevity, to four of the obvious distinguishing
features of B. excelsa, we can group them into two phylogenetically contrasting
categories :

(a] Characters ancestral for the Stephanocircidae and (b) characters highly speciali-
zed. The first category is exemplified in the postantennal section of the head (usually
termed occiput in our writings on fleas) (Fig. 3) : the dorsal margin is medianly
somewhat rounded dilated (dt) ; this is an early step in the evolution of a sclerotized
band which extends some distance down the side in all other species of the family,
Cleopsylla Rothschild, 1914, taking an intermediate position. Similarly primitive

ENTOM. Ill, 5. 15

i8o

VERY REMARKABLE FLEA FROM ARGENTINA

are the arrangement and number of the setae on the occiput : they form three rows,
in which Barreropsylla agrees with many fleas outside the family, but differs from
the known species of Stephanocircidae, there being four rows in Cleopsylla and five
or six in the other genera (for Cleopsylla see F. Smit's paper on fleas in this issue
of the Bulletin, Figs. 13, 14). Category (b) is represented by the modifications of
the genal comb and the pronotum : whereas the known species of helmet fleas have
a genal comb of four or more spines (sometimes much shortened), we find in Barrero-
psylla at the genal edge one to three dark knobs (Gc), which I interpret as being the
remnants of a three-spined ancestral comb ; an advance in evolution by reduction
which is no rarity in fleas. The modification of the pronotum is an extreme and
probably not surpassed by any other flea existing ; it transcends the pronotal

FIG. I. Pronotum of Nonnapsylla rothschildi Wagn., 1938, $ from Peru.

FIG. 2. Pronotum of Meringis altipecten Traub & Hoff, 1951, <J from New Mexico.

development of all fleas. The dorsal area is drawn out into a long process which
reaches to the end of the mesonotum and bears a comb, the spines of which project
downward. It is a feature which would seem to be more appropriate for a species
of Membracidae than for a flea. In order to make it easier to understand what has
happened to the pronotum of Barreropsylla we figure the pronotum of two other
fleas (Figs, i and 2) for comparison. In contrast with the new genus the helmet
fleas have a vertical pronotal comb more or less similar to that of Nonnapsylla
Wagn. 1938 (Fig. i) ; in no species of the family is there an indication of a develop-
ment in the direction of the Barreropsylla pronotum. In other groups of fleas,
however, species occur in which the pronotum is dorsally somewhat widened back-
wards, the comb being curved ; and a small number of species are now known in
which the posterior margin is longer than the anterior one, the projecting posterior
angle being raised well above the mesonotum (Fig. 2 ; see also figs. 54, 55 in F.
Smit's paper). Comparing Fig. 3 with Fig. i it is evident that the development of

VERY REMARKABLE FLEA FROM ARGENTINA 181

the mantilla has had various effects on the pronotum, one of them being a considerable
reduction in its extension downwards. The true occiput, which lies partly within
the pronotum, seems likewise to be affected ; the line which represents its upper
surface in a lateral aspect is broader than usual, looking like a rod emanating from
the subapical vertical sclerotized ridge ; its apex touches the inner side of the pro-
notal exoskeleton and is surrounded by a pale halo (dap], which give one the impres-
sion of being a condylus and socket a very interesting point. However, the detail
is too much obscured in both specimens and the observation requires confirmation
by further material.

Though we see what has happened to the pronotum, we have no evidence of how
and why the mantilla has come into existence. Nevertheless, the wide difference
between the usual pronotum of a helmet flea (Fig. i) and that of Barreropsylla
(Fig. 3) and the occurrence here and there among other fleas of a pronotum like
Fig. 2 suggest an evolution by stages. It is therefore reasonable to assume that
intermediate stages have existed and perhaps one or the other of them still exists,
which gives us hope that Dr. J. M. de la Barrera will discover the missing link or
links. Meanwhile the new flea has to be described and its proper position in the
Craneopsyllinae to be ascertained.

The distinctions of Barreropsylla being a mixture of primitive somatics, which
put the genus at the bottom of the phylogenetic tree of the family, and very highly
specialized ones, which place it at the top, it is obvious that the flea followed its
own line of evolution and represents a tribe which branched off below the
other branches.

Barreropsyilini tribus nova Craneopsyllinarum

Dorsal margin of occiput with a short downward widening in middle. Pronotum
dorsally produced backwards into a process which bears a comb each side ; no
vertical pronotal comb. This diagnosis is kept short as it may possibly have to
cover in future one or several less strongly modified species.

Barreropsylla gen. nov.

(Fig. 3)

($. Spines of comb of helmet long, except the first (uppermost), which is reduced
in length and width, the ventral spine being the longest ; anterior area of helmet
narrow, with short incrassations, many of which are obtuse, and with a row of small
and rather indistinct bristles ; upper third of this area without incrassations ;
ventrally the area projects down as a nose which is about as long as the base of the
ventral spine is broad (in <$ the nose broken away). Helmet not distinctly separated
from genal area, but there is below the spines of the comb a pale stripe from the
apex of the second spine downwards ; at the posterior side of this stripe there are
numerous small bristles, and from its anterior side some narrow, apically rounded-
dilated sclerotized streaks extend obliquely across it in an upward direction. Maxilla
rounded-truncate, not pointed. Occiput with only three rows of bristles and a

1 82

VERY REMARKABLE FLEA FROM ARGENTINA

VERY REMARKABLE FLEA FROM ARGENTINA 183

single long bristle between second and third row. In the only known species the
pronotal mantilla (or hood) on each side with a comb of ten spines, which gradually
become shorter, the last one being little longer than broad.
Type : Barreropsylla excelsa sp. nov.

Barreropsylla excelsa sp. nov.
(Figs. 3, 4, 5)

$$. Comb of helmet with 8 spines (as in Cleopsylla townsendi Roths., 1914),
lowest spine (measured from the upper basal angle) one-half longer than the second
spine from above, basal margins of spines more or less deeply sinuate. Much behind
pharynx and close to the sclerotized anterior margin of the antennal fossa one long
bristle, in front of which appears a short branch of the tentorium ; no second long
bristle between helmet and antennal fossa, but several small ones ; the small spoon-
like sclerotizations under the spines of the helmet number 6 or 7. Remnants of
genal comb (Gc) : on left side of head of $ a knob at lower angle of genal margin
(Fig. 3), on right side a row of 3, in $ 3 on both sides. Segment I of antenna nearly
straight, a little more than one-half longer than broad, II with a row of 5 or 6 bristles,
of which in <$ some (4?) reach beyond middle of club, whereas in $ 5 extend to apex
of club or a little beyond it. The widening of the middle of the dorsal margin of
the occiput is not distinctly more sclerotized than the rest of the margin, and there
are no indications of other tubers ; the anterior row of occiput contains 2 bristles
in (J and 3 in $, the second and third rows in $ and $ 5 each side, the second row
being continued dorsally by one or two much smaller bristles. The labial palp has

5 segments and reaches to the trochanter (J) or nearly ($), being much longer than
the maxillary palp. Pronotum with 2 rows of bristles, the posterior row containing
10 long ones on the two sides together. For bristles of meso- and metanotum see
Fig. 3. Pale central area of metepisternum twice as long as broad ; on metepimere
2 rows of bristles, in <$ containing 6, 5 on one side and 5, 4 on the other, in $ 6,

6 and 6, 4, besides a long bristle at apical margin in <$ and $. Abdominal tergites
likewise with 2 rows, the numbers of bristles being on III and IV in < and $ 13 or
14, 17 or 18 (the two sides together), on V in ^ 4, 17, in $ 4, 15, on VI in
(J 4, n, in $ i, 12, on VII in $ o, 10, in $ 5, 6. Antepygidial bristles long and
stout, in ( i each side, in $ 2 equal in length. Basal abdominal sternum with a
pair of ventral marginal bristles, sterna III to VI with a single row, numbering
(the two sides together) in <? on III 10, IV 9, V 8, VI 7, in $ on III n, IV to VI
8 each.

Femora, on outer surface, with a few lateral bristles in apical third and i to 3
further forward, besides the usual ventral bristles near base and apex; on inner
surface only the ventral subapical one, no lateral ones ; on outer side of fore-, mid-
and hind-tibiae a row of lateral bristles numbering respectively, in $ 6, 7, 9 or 10,
and in $ 7, 10 or n, and 12 ; the bristles at the dorsal margin and close to it counted
from the base to the middle of the apical margin number the same in ^ and $ ; in
fore-tibia 19 and in mid- and hind-tibia 22 ( i) ; the row of bristles at the apical
margin is interrupted in all three tibiae, not forming a comb,

184

VERY REMARKABLE FLEA FROM ARGENTINA

Terminalia. <$. (Fig. 4). As VII. t. is totally and VIILt. partially telescoped
into Vl.t., it is not possible to determine accurately the outline of each segment and
to recognize the homology of every sclerite. In front of the antepygidial bristle
lies a narrow curved sclerite which bears on the convex anterior side a row of rounded

Vl.st.

FIG. 4. Barreropsylla excelsa, terminalia of <$.

humps ; it has evidently been shifted from its proper place ; it may be a part of
VIl.t. or it may not. When a second $ becomes available for comparison the nature
of this conspicuous sclerite may become clear. The margin of VIILt. runs across
the sensilium and in consequence the shape of the spiracle (Spi) is obscured. VIII. st.
bears only three bristles each side, two of them small, the third fairly large ; the
extension dorsal of VHI.st. is not clear at all, and the dotted line to indicate this
extension may be in the wrong place ; IX.t. is much longer than broad ; it is anted-

VERY REMARKABLE FLEA FROM ARGENTINA

185

orly produced into two broad apodemes ; the lower one is the manubrium (M), the
upper margin of which (anterior margin in figure) is nearly straight and not much
longer than the manubrium is broad at the base ; ventral margin evenly convex.
The sinus above the manubrium round and the apodeme above it almost the inverse
of it. The clasper (Cl) ends with a conical process (P), which is about twice as long
as broad and bears a row of three stout and two slender bristles at the anterior (or
dorsal) margin, two bristles at the apex and two at the posterior side, there being

FIG. 5. Barreropsylla excelsa, terminalia of

in addition a long one on the inside at the base of the process. Below P and forming
with it a round sinus there is a narrow, almost spine-like projection, which may
possibly not belong to the clasper ; it is placed close to the clasper's sclerotized
swelling, which bears the movable digitoid F. This is claw-like, narrow, curved
towards P and pointed and bears a submarginal row of 4 bristles, of which the
lowest is the stoutest ; then follow on the margin of F 6 very thin ones (some quite
short) and at some distance from apex a long one about the thickness of the third
of the submarginal row but longer. The upper end of the anterior arm of IX. st. is
curved backwards as shown in Fig. 4 ; the posterior arm is straight, broadest at the

186 VERY REMARKABLE FLEA FROM ARGENTINA

base, gradually narrowed, the apical half being about as broad as the basal fourth of
the antepygidial bristle and bearing two slender bristles. X.t. triangular in lateral
aspect, with an oblique row of thin bristles on the side and several more bristles
further forward, the base partially overlapped by the sensilium. Most of the distal
sclerites of the phallosome (Pti) are hidden and not indicated in Fig. 4 ; the proximal
portion is bulbiform and more convex dorsally than ventrally. Some distance
proximally of the digitoid F lies, each side of body, a sclerite which resembles the
blade of a woodcutter's axe (ax) and is conspicuous by its shape ; it is evidently
a part of the phallosome.

$. (Fig. 5.) VII. t. with 13 bristles in a row and in front of them 8 small ones
(on the two sides together) ; its apical margin nearly straight. Cavity of spiracle
(Spi) of VHI.t. curved upwards, being a little more than twice as long as broad ;
from the spiracle downwards there are 19 bristles on the left side and 14 on the right,
the upper bristle of the second row being the largest ; on the inside of the apical
margin there is a cluster of 3 short and stiff bristles and none further down. X.t.
bears 7 bristles each side, the dorsal median one being larger than the apical bristle
of the stylet (Sty) ; X.st. is evenly convex below, and has a small bristle at the upper
margin and a long one near the lower part of the apical margin. Stylet about three
times as long as broad. The spermatheca (R.s.) resembles that of Tiampsylla
titschacki Wagn., 1937, its orifice being ventral.

Length <$ 2-2 mm., $2-5 mm.

One pair from the Republic of Argentina : collected by Dr. J. M. de la Barrera
at Bariloche, Nahuel Huapi, Rio Negro, 780 mm., 3 vii. 1952, on Akodon sp., type <.
In the coll. of the British Museum (Nat. Hist.).

DESCRIPTIONS OF NEW AND LITTLE-KNOWN

SIPHONAPTERA

By F. G. A. M. SMIT

SYNOPSIS

Six new fleas are described in this paper and two imperfectly known species are redescribed.
Unless stated otherwise, all type specimens are in the Rothschild collection and the British
Museum collection of Siphonaptera at Tring, Herts.

Family VERMIPSYLLIDAE

Dorcadia ioffi, n. sp. 1

(Figs, i, 3)

TYPE MATERIAL. Holotype male and allotype female : Issyk-Kul, Tien Shan,
Turkestan, 1943, ex Ovis aries, received from I. G. loff. Paratypes : two females
with same data as holotype and allotype.

DIAGNOSIS. Closely related to the hitherto only known representative of the
genus, Dorcadia dorcadia (Rothschild), 1912, but readily distinguishable in both
sexes by the much larger size of the abdominal spiracular fossae ; the male differs
in the shape and proportions of the modified segments, the female likewise in the
terminal segments, though less markedly so.

DESCRIPTION. Apart from the larger size of the spiracular fossae and the details
of the terminalia, there do not seem to be any major differences in the two species of
Dorcadia.

MODIFIED ABDOMINAL SEGMENTS : MALE (Fig. i, cf. Fig. 2). Manubrium rather
narrow, about five times as long as broad at its apically broadest point, in D. dorcadia
about two and a half times as long as broad. The posterior margin of the clasper
above the movable process bears about half a dozen long setae at or near the margin
which is truncate just above the movable process ; in D. dorcadia only 2 of the
marginal setae are elongated and the margin above the movable process forms an
acute process. Along the margin of the clasper below the movable process 3 very
long curved setae ; in D. dorcadia there are 4-5 such setae. On the outer side
of the clasper a vertical patch of about 6 setae, on the corresponding part, but on
the inside, about 16 small setae ; in D. dorcadia on the outer side about 12 setae,
but none on its inner side. Movable process proportionally longer than that of

1 The specimens of this new species (Dorcadia) and the next one described here (Chaetopsylla) were
among material received in 1946 from Prof. I. G. loff of the Parasitological Laboratory at Saratov,
U.S.S.R., in exchange for material from the Tring collection. Since then it has become practically
impossible to communicate with colleagues in Russia, and I have not been able to inform Professor
loff of the status of the specimens which are now here described as new. To describe new species without
the consent of the colleague who submitted the material for study and comments would normally be
unethical, but in this case the increase of our knowledge seemed to outweigh this consideration a view
which is shared by my colleagues at Tring.

i88 DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

FIGS, i, 2.

Fig. i, Dorcadia ioffi n. sp. Terga VII, VIII and clasper (tergum IX) and anal segment.
(Holotype.) Fig. 2, Dorcadia dorcadia (Rothschild). Terga VII, VIII and clasper
(tergum IX) and anal segment. (Yen-an Fu, Shensi.)

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 189

FIGS. 3-5.

Fig. 3, Dorcadia ioffi n. sp. Terminalia of female. (Allotype.) Fig. 4, Dorcadia dor-
cadia (Rothschild). Tergum VII. (Yen-an Fu, Shensi.) Fig. 5, Dorcadia dorcadia
(Rothschild). Terminalia of female. (Yen-an Fu, Shensi).

igo DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

the other species and with more numerous setae. In both species the dorsal anal
lobe (tergum X) is fused with tergum IX ; the ventral anal lobe (sternum X) is
distinctly longer in the new species than in D. dorcadia. The very peculiar phallo-
some is extremely similar in the two species.

FEMALE (Fig. 3, cf. Figs. 4, 5). This is a larger flea than D. dorcadia (although
the reverse appears true owing to the lesser degree of expansion of the available
females of D. ioffi), and the fact that its tergal and sternal sclerites are larger (cf.
Figs. 3 and 4, 5) may be due to this fact, but they are also distinctly broader and
with a more convex anterior margin, as shown in the figures. The spiracular fossae
are actually and relatively larger ; above the spiracular fossa of tergum VIII about
8 upwards-pointing setae, while in D. dorcadia there are only 2 or 3 such setae ;
the setae behind and below the sensilium and on the ventral anal lobe are also more
numerous than in D. dorcadia. The spermathecae of the two species are very
similar.

LENGTH OF MALE, 3 mm. ; the length of the female cannot be given in a satis-
factory way, owing to the different stages of expansion of the specimens.

REMARK. This new species is named in honour of the great Russian Siphon-
apterologist, Prof. I. G. I off.

Chaetopsylla caucasica, n. sp.

(Figs. 6, 8, 10, 12)

TYPE MATERIAL. Holotype male and allotype female : Kisha, N.W. Caucasus,
1940, ex Maries martes, received from Prof. I. G. loff.

DIAGNOSIS. The new species is closely related to Chaetopsylla matina Jordan,
and differs from it in the male by the shape of the corpus of the clasper, the straight-
ness of the movable process and the greater length of the distal arm of sternum IX ;
the female is difficult to distinguish from that of C. matina, except for a difference in
the shape of the spermatheca.

DESCRIPTION : HEAD. Frontal tubercle deciduous ; labial palp consisting of
5 segments, not extending beyond the apex of the fore coxa. Without a strong seta
on the genal process immediately below the hinder part of the eye.

THORAX. Metepimeron with 2 rows of setae, the anterior row being well developed
and composed of 4 or 5 setae.

ABDOMEN. The setae of the terga are not notably long and coarse ; for the
arrangement of setae below or behind the spiracular fossae of terga II-VII see
Fig. 12 ; in the female these spiracular fossae are situated well in front of the short
main tergal row of setae and there are in this sex two setae above the spiracular
fossa of tergum II, and one above those of terga I II-VII.

MODIFIED ABDOMINAL SEGMENTS : MALE (Figs. 6, 8, cf. Figs. 7, 9). Corpus of
clasper about one and a half times as long as broad, as in C. matina, but its posterior
margin is strongly convex ; the setae along this margin far more numerous than in
C. matina ; setae on the lower part of the inner side of the corpus longer and more
distinctly grouped together than in C. matina. Movable process apparently straight

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 191

FIGS. 6, 7.

Fig. 6, Chaetopsylla caucasica n. sp. Clasper and sternum IX. (Holotype.) Fig. 7,
Chaetopsylla niatina Jordan. Clasper and sternum IX. (Cauterets, Hautes Pyrenees.)

I 9 2 DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

12

FIGS. 8-12.

Fig. 8, Chaetopsylla caucasica n. sp. Aedeagal apodeme. (Holotype.) Fig. 9, Chaetopsylla
matina Jordan. Aedeagal apodeme. (Cauterets, Hautes Pyrenees.) Fig. 10, Chaeto-
psylla caucasica n. sp. Sternum VII, spermatheca and sclerotized part of the ductus
bursae copulatricis. (Allotype.) Fig. n, Chaetopsylla matina Jordan. Sternum VII,
spermatheca and sclerotized part of the ductus bursae copulatricis. (Cauterets, Hautes
Pyrenees.) Fig. 12, Chaetopsylla caucasica n. sp. Terga II- VII of male. (Holotype.)

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 193

(that of the right-hand side is curved in the only male available, but this shape
seems to be due to distortion) ; the process narrows markedly at two-thirds and is
then of uniform width to near the apex, where it tapers gradually ; the acetabulum
is situated at the anterior margin of the corpus, while in C. matina it is placed well
away from this margin. Manubrium with a straight upper and a convex lower
margin. Proximal arm of sternum IX more distinctly angulate than that of C.
matina, while the distal arm is slenderer and relatively much longer than in the
latter species and bears apically a few rather short setae (not long ones as in C.
matina). Aedeagal apodeme much broader than that of C. matina.

FEMALE (Fig. 10, cf. Fig. n). Sternum VII very much like that of C. matina,
but the ventral lobe appears to be sharper, although females of C. matina are rather
variable in this respect. Tergum VIII with a main row of n setae on one side,
12 on the other, on each side the 6 lower and larger setae of this row are situated
below the spiracular fossa. Appendix of spermatheca relatively stouter than that
of C. matina, while the reservoir also appears to be much larger in relation to the
length of the appendix. The sclerotized part of the ductus bursae copulatricis is
less strongly bent than that of C. matina, but whether this shape is genuine or not
cannot be ascertained from the only female available.

LENGTH OF MALE, 2| mm. ; female approximately 3^ mm.

Family STEPHANOCIRCIDAE
Subfamily CRANEOPSYLLINAE
Cleopsylla monticola, n. sp.

(Figs. 13, 15, 17, 19, 21)

TYPE MATERIAL. Holotype male and allotype female : Pichincha, Ecuador,
ist October, 1931, ex Caenolestes fuliginosus, collected by F. Spillmann. Paratypes :
one male and one female, Chimborazo, Ecuador, April, 1931, ex Sigmodon sp. ;
one male, Iliniza, Ecuador, April, 1931, ex Thomasomys sp. ; these paratypes were
also collected by F. Spillmann.

DIAGNOSIS. This new species from Ecuador differs from the hitherto only known
representative of the genus, Cleopsylla townsendi Rothschild (occurring in Peru), by
the straightness of the anterior margin of the frons, the strongly reduced number of
small setae bordering the antennal fossa dorsally, and by the chaetotaxy of the hind
tibia. The modified abdominal segments and genitalia of the male are extremely
similar in the two species, but the females are most easily distinguishable by the
shape of the spermatheca.

DESCRIPTION : HEAD (Fig. 13, cf. Fig. 14). Dorsal margin of frons anteriorly
strongly rounded ; the anterior margin of the frons slopes strongly backwards ;
it is straight for most of its extent and in its lower portion rounded to form the
genal margin, which is slightly longer than that of C. townsendi. On the frons
there is a vertical ctenidium, consisting of 7 spines in the male, and 8 in the female,
as is also the case in the other species ; the two species also agree in having the lowest

ENTOM. Ill, 5. l6

194 DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

spine elongated and curved downwards pre-apically ; the bases of these spines in
C. monticola form an imaginary line which is parallel to the straight frontal margin,
while this is not the case in C. townsendi owing to this species having usually a
markedly convex anterior frontal margin. In front of the ctenidium a row of 5

FIGS. 13-16.

Fig. 13, Cleopsylla monticola n. sp. Head and pronotum of male. (Iliniza, Ecuador.)
Fig. 14, Cleopsylla townsendi Rothschild. Head and pronotum of male. (Carhuamayo,
Peru.) Fig. 15, Cleopsylla monticola n. sp. Spermatheca (Allotype.) Fig. 16, Cleopsylla
townsendi Rothschild. Spermatheca and genital ducts. (Tacna, Peru.)

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 195

FIGS. 17, 170, 18, i8a, 19, 20.

Fig. 17, Cleopsylla monticola n. sp. Clasper and sternum IX. (Iliniza, Ecuador.) Fig.
170, Cleopsylla monticola n. sp. Movable process of clasper. (Iliniza, Ecuador.) Fig. 18,
Cleopsylla townsendi Rothschild. Clasper and sternum IX. (Carhuamayo, Peru.)
Fig. iSa, Cleopsylla townsendi Rothschild. Movable process of clasper. (Carhuamayo,
Peru.) Fig. 19, Cleopsylla monticola n. sp. Sternum VII of female. (Allotype.)
Fig. 20, Cleopsylla townsendi Rothschild. Sternum VII of female. (Tacna, Peru.)

ig6 DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

setae, the fourth seta from above being the longest. The genal ctenidium consists
of 5 spines, the lowest of which is a little longer than the others and partly concealed
by its neighbour, while the small most dorsal spine is placed on the broad, rounded
genal process and is rather variable in size. Clava of antenna somewhat broader
than that of C. townsendi ; antennal fossa dorsally bordered by one or two small
setae, in contrast with C. townsendi, where there are many more such setae. Chaeto-
taxy of the postantennal region of the head virtually the same in the two species,
except that in the new species the setae, like most of the other setae, are slightly
longer than in C. townsendi.

THORAX. Pronotum (Fig. 13, cf. Fig. 14) with two rows of setae and a ctenidium
of 25-28 spines in the male and 25-27 in the female (in C. townsendi, male with 22-25,
female with 21-26 spines) ; the sharply pointed straight pronotal spines gradually
decrease in length from above downwards in both species. Pronotum dorsally
longer than the uppermost spines, while the reverse is true for C. townsendi ; meso-
thorax and metathorax as in the latter species.

LEGS (Fig. 21, cf. Fig. 22). A row of short spiniform setae on the inner side of
the hind coxa. The chaetotaxy of the posterior margin of the tibiae of the two
species is different and the differences described and figured here are taken from the
hind tibia : the two median groups of strong setae arising from notches in the posterior
margin consist of 3 setae in both species, but in the new species the innermost seta is
displaced downwards and has become marginal, thus causing the tibia to possess a
false comb, while in C. townsendi the innermost seta of the two groups is in line
with the other two setae and therefore the more primitive condition is retained here.

ABDOMEN. Terga I-V in the male with 2-4, 2-4, 3, 3, 2-3 apical spinelets respec-
tively on each side, in the female 2-3, 3, 1-2, 1-3, 1-2 respectively ; for C. townsendi
these figures are : in the male 2-3, 2-3, 1-2, 1-2, i, and in the female 2-3, 2-3,
1-2, 1-2, i. The chaetotaxy of the first 7 abdominal segments is virtually the same
in the two species.

MODIFIED ABDOMINAL SEGMENTS : MALE (Fig. 17, cf. Fig. 18). Movable process
of clasper (Fig. 170) with its apex expanded both anteriorly and posteriorly, the
posterior expansion forming a conspicuous lobe which is almost absent in C. towns-
endi (Fig. i8<z). The distal arm of sternum IX tends to be slightly broader than
that of C. townsendi. In other respects the modified segments, as also the phallo-
some, are very much alike in the two species.

FEMALE (Figs. 15, 19, cf. Figs. 16, 20). Posterior margin of sternum VII with a
long and shallow sinus, similar to that in C. townsendi though shorter ; the row of
setae extends upwards to a point level with the lower margin of the lobe of the
posterior margin of the sternum, while in C. townsendi it extends to above this
point. Spermatheca markedly different from that of the Peruvian species, its tail
being much shorter and stouter and internally finely striated (instead of reticulate)
and the striae on the inside of the reservoir are less dense. The genital ducts could
not be made out in the two females available.

LENGTH of male, 2 mm. ; female, 2| mm.

REMARKS. Cleopsylla townsendi is known to occur in places at high altitudes in
Peru (between 12,000 and 16,000 ft.) and the same seems to be true for the new

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 197

species described above, for all specimens of the latter were collected on three of the
highest mountains in Ecuador, namely Chimborazo (altitude i 20,500 ft.), Pichincha
(altitude 16,000 ft.) and Iliniza (altitude 17,500 ft.). Though the altitude is
not given on the slide labels, it seems safe to assume that the specimens were collected
well above 10,000 ft., near the timber line.

It should be noted that the paratypes of Cleopsylla monticola were recorded by
Jordan (1931) under the name of Cleopsylla townsendi, and also that (contrary to
his belief at that time) the supposed inaccuracy in his drawing of the tail of the
spermatheca of C. townsendi does not exist, the drawing being perfectly accurate
and the difference in the spermatheca being specific.

Plocopsylla phyllisae, n. sp.

(Figs. 25, 26, 28, 30)

TYPE MATERIAL. Holotype male from Guamani, Ecuador, July, 1931, ex Oryzomys
sp., collected by F. Spillmann.

DIAGNOSIS. The new species is related to Plocopsylla phobos Jordan, but differs
in the male by the spines of the genal ctenidium being much more strongly reduced,
by the typical abdominal terga not being rounded off dorso-posteriorly, and by having
a broader clasper and a differently shaped movable process and spiniform on the
inner side of the latter ; moreover, the sterna IX of the two species are not alike
and the hind tibiae differ in their chaetotaxy. Female unknown.

DESCRIPTION : HEAD (Fig. 25, cf. Figs. 23, 24). Prectenidial part of head narrower
than the " helmet "- ctenidium, which consists of 14 spines on each side. In other
respects the head is extremely like that of P. phobos, except that the four genal
spines are strongly reduced in size, which is not the case in the male of any other
known Plocopsylla, and in the female sex such small genal spines are only found in
P. phobos (which could imply that the females, attributed to P. phobos, are not that
species).

THORAX. Very similar to that of P. phobos ; in both species the pronotal cteni-
dium consists of 14 spines.

LEGS (Fig. 30, cf. Fig. 31). The setae at the posterior (dorsal) margin of the
hind tibia are much thinner and therefore grouped more closely together than in
P. phobos, while the fourth notch from the apex bears only 2 setae as against 3 in
P. phobos.

ABDOMEN (Fig. 26, cf. Fig. 27). Chaetotaxy of terga I-VII and of sterna II-VII
similar to that of P. phobos, but the setae are much shorter than in the latter species.
The shape of the terga is quite normal (Fig. 26), but stress should here be laid upon
the shape of those of P. phobos : in the male of that species the dorsal part of the
posterior margin of terga 1 1- VI is rounded and forms no distinct angle with the
dorsal margin (Fig. 27). This is an interesting form of sexual dimorphism, since in
the female of P. phobos the terga are normally shaped (as in Fig. 26). Similar
modified terga are found in the males of P, wolffsohni (Rothschild) and P, chiris

ig8 DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

26

FIGS. 21-27.

Fig. 21, Cleopsylla monticola n. sp. Hind tibia of male. (Iliniza, Ecuador.) Fig. 22,
Cleopsylla townsendi Rothschild. Hind tibia of male. (Carhuamayo, Peru.) Fig.
23, Plocopsylla phobos Jordan. Gena of male. (Holotype.) Fig. 24, Plocopsylla
phobos Jordan. Gena of female. (Guamani, Ecuador.) Fig. 25, Plocopsylla phyllisae
n. sp. Gena of male. (Holotype.) Fig. 26, Plocopsylla phyllisae n. sp. Terga III-V.
(Holotype.) Fig. 27, Plocopsylla phobos Jordan. Terga III-V. (Holotype.)

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 199

FIGS. 28, 29.

Fig. 28, Plocopsylla phyllisae n. sp. Modified abdominal segments. (Holotype.) Fig.
29, Plocopsylla phobos Jordan. Modified abdominal segments. (Holotype.)

200 DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

FIGS. 30-33.

Fig. 30, Plocopsylla phyllisae n. sp. Hind tibia of male. (Holotype.) Fig. 31, Ploco-
psyllaphobos Jordan. Hind tibia of male. (Holotype.) Fig. 32, Craneopsylla minerva
minerva (Rothschild). Apex of clasper. (Villa Carolina, Rio Lavallen, Argentina.)
Fig. 33, Megarthroglossus jamesoni n. sp. Head, thorax and coxae. (Holotype.)

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 201

(Jordan), and the former is also sexually dimorphic in this respect, like P. phobos,
while the sole known female of P. chins lacks the abdomen. Terga I-V with 4-5,
3-4, 3-4, 3-4, i-i apical spinelets respectively (in P. phobos : 3-4, 3-4, 2-3, 1-2,
o respectively). Of the two antesensilial setae the upper one is a thin and very
small seta, and the lower, which in the other species of Plocopsylla (except P. ender-
leini Wagner, where antesensilial setae are absent in the male) is quite normally
developed, though sometimes rather short, is vestigial.

MODIFIED ABDOMINAL SEGMENTS OF MALE (Fig. 28, cf. Fig. 29). Apodeme of
tergum IX and manubrium less broad than in P. phobos, but the corpus of the clasper
is distinctly broader than in the latter, its dorsal margin being almost straight ;
the dorso-apical margin is bordered by a fringe of 12 long setae. Movable process
of clasper far less broad basally than that of P. phobos, and with a sharply pointed
apex ; the modified spiniform seta on the inner side of the movable process differs
markedly from those in all other representatives of the genus by having a downward-
pointing triangular thin extension. As is usual in all members of this genus and
also in those of the other genera of the subfamily Craneopsyllinae, a distinct non-
setiferous process is attached to the inside of the corpus of the clasper near the dorsal
part of the base of the movable process. This lateral process is always associated
with the latter, and a useful hint as to its origin and function is given by the
structure of the clasper of Craneopsylla minerva minerva (Rothschild) (Fig. 32).
Here we see that on the inner side of the clasper, along the margin of the dorso-apical
angle, the surface is partly sclerotized and striated ; opposite this the movable
process has a striated patch on its outer side. In Plocopsylla the clasper is extended
further backwards and the apex of the movable process does not reach the apex
of the clasper ; since the inner striated sclerotization is so closely associated with
the movable process, it has moved inwards and developed into a rodlike structure. The
function of this process could be (a) to act as a locking device for the movable process
or (b) to serve as a stridulatory organ (in connection with the corresponding striated
area of the movable process). 1 Posterior margin of proximal arm of sternum IX
more irregular than that of P. phobos ; the distal arm bears 2 modified setae, the
one at the ventral margin being spiniform with a slightly curved apical part, the
other is extremely short and has a very blunt apex.

LENGTH of male, 2 mm.

REMARKS. If the relative development of the genal spines in the two sexes of
this new species is the same as it is in P. phobos (where these spines are well developed
in the male and strongly reduced in the female), the unknown female of P. phyllisae
might be expected to possess at most vestiges of genal spines, a condition confined
in known Stephanocircidae to Barreropsylla excelsa Jordan (see the paper by K.
Jordan in the same issue of this Bulletin).

The new species is named after Miss Phyllis T. Johnson, Entomologist of the
Walter Reed Army Medical Center, Washington, D.C., in appreciation of her great
interest in the flea fauna of South America.

1 Though the latter suggestion seems less likely than the first one, it would be of interest if South
American entomologists would try out the experiments which Ossiannilsson performed with Auchenor-
rhynchous Homoptera (Opusc. ent., 1946, 82-84) in order to prove whether or not these fleas can sing.

202 DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

Family HYSTRICHOPSYLLIDAE

Subfamily ANOMIOPSYLLINAE

Megarthroglossus j antes oni, n. sp.

(Figs. 33-36)

TYPE MATERIAL. Holotype male and allotype female : Pine Nut Mountains,
Douglas County, Nevada, U.S.A., 25th March 1951, from the nest of Neotoma
cinerea, collected by E. W. Jameson, Jr.

DIAGNOSIS. The new species belongs to the group of species of Megarthroglossus
which normally have a pronotal ctenidium consisting of 16 spines, i.e., M. sicamus
Jordan & Rothschild, M. bisetis Jordan & Rothschild and M. becki Tipton & Allred,
and is easily separable from these in the male by the shape of sternum VIII, the
posterior margin of which has no marked ventral incision, and moreover its posterior
margin is almost straight and does not form a lobe ; the female differs from the others
of the group by the combination of the following characters : the possession of 3
antesensilial setae each side, sternum VII without a lateral sinus, and the appendix
of the spermatheca being less than twice as long as its reservoir, which has no distinct
" collar."

DESCRIPTION. HEAD AND THORAX (Fig. 33) similar to those of the other members
of the genus ; pronotal ctenidium consisting of 16 spines ; abdominal terga I and II
each with i-i apical spinelets respectively in the male ; in the female there is only
i spinelet on one side of tergum I.

MODIFIED ABDOMINAL SEGMENTS : MALE (Fig. 34). Sternum VIII without a
trace of a ventral incision of its posterior margin ; the posterior hyaline part of this
sternum with an almost straight posterior margin which forms no lobe ; the length
of the entire ventral margin of sternum VIII equals that of sternum VII. Clasper
with 6 slender setae along its dorsal margin, a short row of about 6 small setae on its
inner side from above the acetabulum to the dorso-apical angle and with n long
and medium-sized setae distributed along the whole of the posterior margin. Movable
process elongate, smoothly and slightly convex posteriorly, its apex obliquely
truncate ; none of its setae stout.

FEMALE (Figs. 35, 36). Tergum VII with 3 antesensilial setae each side. Posterior
margin of sternum VII with a ventral sinus but without a lateral sinus ; with a main
row of 5 setae and in front of the lower setae about 3 additional ones. Appendix
of spermatheca about one and a half times as long as the reservoir, the " collar " of
which (the part which is striated on the inside) only bulges dorsally, while the lower
margin of this collar is straight and in line with the lower basal margin of the
appendix.

LENGTH of male, i| mm. ; female, 2 mm.

REMARK. This new species of the Western North American genus Megarthro-
glossus is named after my colleague Dr. E. W. Jameson, Jr., of the University of

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 203

California, who himself collected the two specimens, and most kindly allowed me to
describe them and to incorporate the holotype and allotype in the British Museum
collection at Tring.

35

FIGS. 34-36.

Fig. 34, Megarthroglossus jamesoni n. sp. Clasper, sterna VIII and IX. (Holotype.)
Fig. 35, Megarthroglossus jamesoni n. sp. Spermatheca. (Allotype.) Fig. 36, Megar-
throglossus jamesoni n. sp. Sternum VII and spermatheca. (Allotype.)

20 4 DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

Subfamily CTENOPHTHALMINAE

Palaeopsylla similis peusi, n. ssp.

(Figs. 37 and 39)

TYPE MATERIAL. Holotype male and allotype female : Zerma, Rui Mountain,
East Serbia (on the border of Bulgaria, 42 54' N. 22 33' E.), 25th July, 1935, ex
Talpa europaea pancici (= Talpa europaea europaea), collected by N. Zemcuznikov.
Paratypes : two males and one female with same data as holotype and allotype.

DIAGNOSIS. Separable from the nominotypical subspecies, Palaeopsylla simiUs
similis Dampf, in the male by the convex dorsal margin of the clasper, by the wider
base and narrower apex of the movable process and by the more strongly serrate
posterior margin of the proximal arm of sternum IX ; in the female by the shape
of sternum VII, the posterior margin of whiclj has no sinus below its dorsal lobe.

DESCRIPTION. Except for differences in the terminal segments, the new sub-
species is indistinguishable from the nominotypical subspecies.

MODIFIED ABDOMINAL SEGMENTS : MALE (Fig. 37, cf. Fig. 38). Dorsal margin
of corpus of clasper distinctly convex, its dorso-apical angle rather acute and pro-
jecting a little above the point of insertion of the seta on the posterior margin.
Manubrium similar to that of the nominotypical subspecies, and like that of the
latter showing a certain amount of variation in its width. Movable process of clasper
straight, with a broad triangular base ; the anterior (or dorsal) margin narrows
abruptly near the apex, and from the apical angle the margin rounds off to continue
as the posterior margin ; in the nominotypical subspecies the anterior margin is
smoothly rounded near the apex, but the apical and posterior margins meet at a
right angle. Chaetotaxy of the movable process in the two subspecies rather
similar. Sternum IX in subspecies peusi is very similar to that of the nominotypical
subspecies, but the posterior margin of the proximal arm is much more strongly
serrate over a somewhat longer distance ; also the apex of the distal arm widens
a little less abruptly.

FEMALE (Fig. 39, cf. Fig. 40). Sternum VII with a row of 5 or 6 stout setae,
as in the nominotypical subspecies, but the uppermost seta is shifted more forwards
(backward from the row) ; the posterior margin of this sternum has a broad dorsal
lobe below which the margin slants backwards, but without forming a distinct
sinus, as is the case in the nominotypical subspecies. The reservoir of the sperma-
theca seems to be narrower than that of P. similis similis, but such forms also occur
in the latter, where likewise the part near the orifice may sometimes be distended
ventrad.

LENGTH of male, if-2 mm. ; female, 2,\ mm.

REMARKS. Palaeopsylla similis similis Dampf is widely distributed in Eastern
Europe and is known to occur in East and South-east Germany, Austria, Hungary,
Northern Yugoslavia, Roumania, Bulgaria and European Russia ; throughout its
range of distribution it is a specific parasite of the mole (Talpa europaea). 1 P.

1 The occurrence of one specimen, which is in the Tring collection, on Sorex araneus, caught on the
Glatzer Schneeberg, Silesia, is beyond any doubt accidental, since other specific parasites of the mole
are occasionally found on Sorex and vice versa.

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 205

similis peusi is so far only known from the Rui Mountain on the Serbian-Bulgarian
border, and there also is a specific parasite of the mole. The Rui Mountain may well
be one of the most north-eastern points of distribution of P. s. peusi, since P. s.

FIGS. 37, 37a, 38, 3805.

ig- 37> Palaeopsylla similis peusi n. ssp. Clasper and sternum IX. (Holotype.)
Fig. 37a, Palaeopsylla similis peusi n. ssp. Movable process of clasper. (Holotype.)
Fig. 38, Palaeopsylla similis similis Dampf. Clasper and sternum IX. (Nove Stbske,
Pleso, High Tatra.) Fig. 38a, Palaeopsylla similis similis Dampf. Movable process
of clasper. (Nove Stbske, Pleso, High Tatra.)

206 DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

similis has been found in south-west Bulgaria at Rhodope and Karlik, which localities
are situated south-easterly of the Rui Mt. The new subspecies probably occurs in
Albania and the northern part of Greece.

I have great pleasure in naming this new subspecies after my friend and colleague
Prof. Dr. F. Peus, who, through his excellent work, has contributed greatly to our
knowledge of European fleas.

Family ISCHNOPSYLLIDAE
Subfamily ISCHNOPSYLLINAE

Description of the hitherto unknown male of Ischnopsyllus consimilis
(Wahlgren), 1904.

(Figs. 41, 43, 44)

Lt.-Col. R. Traub, Chief of the Department of Entomology, Walter Reed Army
Medical Center in Washington, D.C., most kindly sent me a male of an Egyptian
bat flea of the genus Ischnopsyllus for study and comparison with the representatives
of this genus in the Tring collection. Though this male does not agree with any
described male of Ischnopsyllus, albeit closely related to /. octactenus (Kolenati),
I thought it could well be the unknown male of Ischnopsyllus consimilis (Wahlgren),
1904, which was described from a single female collected in the neighbourhood of
Cairo, Egypt, ex Rhinopoma microphyllum ; unfortunately this type-specimen must
be considered lost. From Wahlgren's description one deduces that consimilis must
be extremely closely related to octactenus, and it was even thought that it might
be a synonym of the latter. The description of /. consimilis deals mainly with such
unsatisfactory characters as counts of setae, which agree rather well with the numbers
of the Egyptian male (taking into account small differences which may be sexual) ;
the numbers of spines in the ctenidia of the two sexes agree very well. Since both
the female holotype of I. consimilis and the Egyptian male Ischnopsyllus are closely
related to /. octactenus, and differ from it in similar respects, it seems a safe assump-
tion that the male belongs to Wahlgren's species.

I am greatly indebted to Colonel Traub and also to the collector of this male,
Mr. P. Quentin Tomich, for allowing me to describe the specimen. It is charac-
teristic of Colonel Traub's great generosity that he has presented this unique specimen
to the British Museum collection of Siphonaptera at Tring.

DATA. One male ex Pipistrellus kuhli, 5 miles west of Simbellawein (i.e., 12 miles
south of Mansura), Dakahlia Province, [Lower] Egypt, 26th February, 1947, collected
by P. Quentin Tomich.

DIAGNOSIS. Closely related to Ischnopsyllus octactenus (Kolenati), but differs in
the following characters : mesonotum and metanotum each with only a few semi-
erect setae forming a " mane " ; dorso-apical angle of clasper obtuse, strongly
rounded ; sternum VIII broader, with an oblique and rounded apex ; sternum IX

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 207

Figs. 39-42.

Fig- 39, Palaeopsylla similis peusi n. ssp. Sternum VII and spermatheca of two females.
(Zerma, Rui Mt., Serbia.) Fig. 40, Palaeopsylla similis similis Dampf. Sternum
VII and spermatheca. (Moscow, Russia.) Fig. 41, Ischnopsyllus consimilis (Wahl-
gren). Mesonotum and metanotum of male. (Simbellawein, Egypt.) Fig. 42,
Ischnopsyllus octactenus (Kolenati). Mesonotum and metanotum of male. (S. Feliu
de Guixols, Spain.)

ao8 DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

differs in details of the apical half of the distal arm ; crochet of phallosome with an
upturned apex. Female : From the information available (i.e., Wahlgren's original
description) the only difference from /. octactenus seems to be in the number of spines
in the pronotal ctenidium, namely 26 as against 29-32 in /. octactenus, though this
difference may not be genuine, since Wahlgren stated, " Da ich nur ein Exemplar
zu untersuchen gehabt habe, sind die Stachelzahlen der grosseren Kamme (der
beiden thorakalen und des zweiten und dritten abdominalen) insoweit unsicher, als
ein Irrtum bis auf zwei Stacheln nicht vollig ausgeschlossen ist."

43

FIGS. 43, 44.

Fig. 43, Ischnopsyllus consimilis (Wahlgren). Clasper. (Simbellawein, Egypt.) Fig.
44, Ischnopsyllus consimilis (Wahlgren). Aedeagal crochet, sternum IX and sternum
VIII. (Simbellawein, Egypt.)

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 209

DESCRIPTION. The number of spines in the eight ctenidia of /. consimilis and
7. octactenus are compared below :

/. consimilis $ I . octactenus <$ I. consimilis $ 7. octactenus $

Pronotum 25 . 28-31 . 26 . 29-32

Metanotum 25 . 28-32 . 24 . 25-31

Tergum I . 12 11-18 . 12 . 11-15

II .22 . 20-28 . 20 . 21-27

III . 20 . 19-22 . 18 . 16-22

IV . 17 . 15-18 . 18 . 13-18
V 13 11-16 . 14 . 9-14

VI . 13 . 9-13 . 12 . 9-13

HEAD. Practically indistinguishable from that of /. octactemis.

THORAX. Pronotum like that of /. octactenus, except for the number of spines
in the ctenidium ; mesonotum and metanotum each with about 4 or 5 semi-erect
setae near the dorsal margin (Fig. 41), while in 7. octactenus these are much more
numerous (Fig. 42). In other respects the thorax, as also the legs, show no notice-
able differences between the two species under discussion.

ABDOMEN. Segments I-VII similar to those of 7. octactenus ; for the numbers
of spines in the tergal ctenidia, see above.

MODIFIED ABDOMINAL SEGMENTS OF MALE (Figs. 43, 44, cf. Figs. 45, 46). Apodeme
of tergum IX rather broad ; manubrium curved upwards, narrowing towards the
apex ; the upper margin of the manubrium near the ventro-anterior part of the
margin of the apodeme shows no lobe as in I. octactenus, but this lobe is also quite
frequently absent, or at most indicated, in the latter species, so it is likely that
specimens of 7. consimilis possessing this lobe may be found in the future. The
dorso-apical angle of the clasper is strongly obtuse and rounded off ; along the dorsal
margin near the angle there are the usual 3 small setae ; posterior margin of clasper
almost straight ; of the 2 acetabular setae the lower is slightly stouter than the
upper ; the ventral margin of the clasper forms a rather straight line. Movable
process of clasper crescentic, the lower half of the posterior margin fairly straight,
while in 7. octactenus it is more regularly rounded ; chaetotaxy of the movable
process similar to that of the latter. Sternum VIII fairly broad and straight, its
proximal part being widened ; dorsal margin of the distal part concave, but much
less so than in 7. octactenus ; its apical margin obliquely rounded, and at the angle
where the dorsal and apical margins meet there is a minute spiniform seta (in
7. octactenus a small thin seta) . On the apico- ventral part of the sternum a curved
row of 6 blade-shaped long setae and in front of these, at the ventral margin, 2 more
flattened setae, but like the corresponding ones in 7. octactenus these are heavily
pigmented ; further proximad along the margin some 6 curved setae. Sternum IX
with the distal arm apically divided into a setiferous upper lobe and a lower non-
setiferous one ; the latter lobe, which is the outer, is elongated and widest in its
middle (in 7. octactenus it is triangular and widest at its base) ; the upper part of
the widened apical portion of the inner lobe much longer (from the point of junction

ENTOM. in, 5. 17

210 DESCRIPTIONS OF NEW AND LITTLE- KNOWN SI PHON AFTER A

with the inner lobe to the apex) than the lower part (in /. octactenus the upper
and lower parts are subequal in size). Aedeagus not substantially different from
that of /. octactenus, but the crochet differs by having amongst ether things an
upturned apex ; other differences can be observed by comparing Fig. 44 with
Fig. 46.
LENGTH of male, 2 mm.

FIGS. 45, 46.

Fig. 45, Ischnopsyllus octactenus (Kolenati). Clasper. (Newport, Isle of Wight.)
Fig. 46, Ischnopsyllus octactenus (Kolenati). Aedeagal crochet, sternum IX and
sternum VIII. (Newport, Isle of Wight.)

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 211

REMARK. In view of the desirability of stability in nomenclature I designate the
above described Egyptian male Ischnopsyllus as neotype of Ischnopsyllus consimilis
(Wahlgren), 1904.

Family AMPHIPSYLLIDAE
Subfamily MESOPSYLLINAE

Redescription of Caenopsylla assimulata (Weiss), 1913, in comparison with
Caenopsylla mira Rothschild, 1909.

(Figs. 47-53, cf. Figs. 54-62)

Through the great kindness of Prof. E. Seguy, Sous-Directeur au Museum National
d'Histoire Naturelle, Paris, I have been able to study the male holotype of Caenopsylla
assimulata (Weiss), which is the only known specimen of this species. Although
Weiss (1913) described it at length, its true morphology and status could not be
ascertained owing to the fact that the few sketchy drawings published by Weiss
do not show the true diagnostic characters. It is therefore not surprising that
Wagner & Wassilieff (1933) remark that ". . . le C. assimulata Weiss a ete trouvee
en Tunisie, a Matmata, sur le Macroscelides rozeti, mais un seul exemplaire insuffisam-
ment decrit par A. Weiss." It is, however, not really a matter of the species being
insufficiently described, since Weiss devoted nine full pages to the description of the
sole specimen, but lack of detailed figures is liable to render any description, however
lengthy, practically useless in the case of creatures such as fleas, where so much
depends on the shape of the details of complicated structures. I fully agree with
Dampf (1945), who advocates more exactness in drawings, but I would like to add
that all taxonomically important parts of fleas should be treated in such a way.
Detailed and accurate drawings replace, in a much more satisfactory way, large
numbers of pages of descriptive matter a fact which unfortunately is not yet
wholly appreciated by many editors of periodicals. The extra expense of repro-
ducing figures is probably more than offset by the saving in the number of pages
required for a description, to say nothing of the fact that in nearly all instances
descriptions of fleas unaccompanied by good and adequate figures are practically
useless.

The holotype of Caenopsylla assimulata is not well cleared and its dorsum is
much contracted ; the specimen is in the collection of the Museum National d'Histoire
Naturelle at Paris.

Only one other species of Caenopsylla is known, C. mira Rothschild, 1909, and
therefore it seemed advisable to redescribe C. assimulata in comparison with C. mira.

Caenopsylla assimulata (Weiss), <J Caenopsylla mira Rothschild, $

DIAGNOSIS

Pronotum rather narrow dorsally, its Pronotum much longer dorsally than ven-

ctenidium straight. Hind tibia without a trally, its ctenidium strongly curved. Hind

well-developed false comb of setae at the tibia with a distinct false comb. Corpus

posterior margin. Corpus of clasper with of clasper narrowest in its middle part, the

subparallel dorsal and ventral margins ; fixed process being drawn out into a long

2 acetabular setae. lobe ; 4 acetabular setae.

ENTOM. Ill, 5. I7

212 DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

48

FIGS. 47-50.

Fig. 47, Caenopsylla assimulata (Weiss). Head and prothorax and fore coxa. (Holo-
type.) Fig. 48, Caenopsylla assimulata (Weiss). Mesothorax, metathorax and tergum
I. (Holotype.) Fig. 49, Caenopsylla assimulata (Weiss). Hind tibia. (Holotype.)
Fig. 50, Caenopsylla assimulata (Weiss). Last hind tarsal segment. (Holotype.)

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 213

53

FIGS. 51-53.

Fig. 51, Caenopsylla assimulata (Weiss). Clasper and sternum IX. (Holotype.) Fig.
52, Caenopsylla assimulata (Weiss). Sternum VIII. (Holotype.) Fig. 53, Caeno-
psylla assimulata (Weiss). Phallosome. (Holotype.)

214 DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

FIGS. 54-57-

FIG. 54, Caenopsylla mira Rothschild. Head, prothorax and fore coxa of male. (Holo-
type.) Fig. 55, Caenopsylla mira Rothschild. Head and pronotum of female. (Biskra,
Algeria.) Fig. 56, Caenopsylla mira Rothschild. Hind tibia. (Holotype.) Fig. 57,
Caenopsylla mira Rothschild. Last hind tarsal segment. (Holotype.)

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 215

6O

FIGS. 58-60.

Fig. 58, Caenopsylla mira Rothschild. Mesothorax, metathorax and tergum I of male.
(El Kantara, Algeria.) Fig. 59, Caenopsylla mira Rothschild. Clasper and sternum
IX. (Holotype.) Fig. 60, Caenopsylla mira Rothschild. Phallosome. (Holotype.)

216 DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

FIGS. 61, 62.

Fig. 61, Caenopsylla mira Rothschild. Terminalia of male. (Holotype.) Fig. 62,
Caenopsylla mira Rothschild. Terminalia of female. (Biskra, Algeria.)

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 217

Caenopsylla assimulata (Weiss),

Caenopsylla mira Rothschild,

DESCRIPTION

HEAD (Fig. 47). The chaetotaxy, as well
as the other characteristics of the head, can
be seen in the figure, but stress should be
laid upon the presence of 2 genal spines
(though normal in the subfamily), the upper
one of which is much thinner than the lower ;
of the five-segmented labial palp the 2nd
and 3rd segments are subequal in length
and each is about half the length of the 4th
or 5th segments. The clava of each antenna
is broken off, and although on the slide label
is written that they are somewhere in the
preparation, I have been unable to find
them ; pedicellus with 3 short setae.

HEAD (Figs. 54, 55). Chaetotaxy very
much the same, but the posterior row on
the postantennal region is oblique and sub-
parallel to the row on the pronotum. The

2 genal spines are slightly stouter ; segments
2-4 of the five-segmented labial palp are all
of subequal length and slightly shorter than
segments i and 5. The antenna is shown
in the figures ; the pedicellus also bears only

3 small setae.

THORAX (Figs. 47, 48). Pronotum of
normal shape, with one row of 12 setae
(6 each side) ; pronotal ctenidium consisting
of 14 spines, the uppermost of the group
formed on each side by the 3 lowest spines,
which all lie closely against each other, is
much the stoutest in the ctenidium and
points obliquely downwards ; the upper
4 spines on each side are slightly curved and
the 2nd and 3rd from above are somewhat
spatulate ; they are all directed downwards
like the third spine from below.

Mesonotum with a main row of 10 setae
(5 each side), preceded by a row of 10 much
smaller setae, both rows practically vertical.
Mesosternosome with 6 setae on one side
and 7 on the other ; the anterior margin of
the mesepisternum is long and almost
vertical.

THORAX (Figs. 54, 55, 58). Pronotum in
both sexes much longer dorsally than ven-
trally, with one row of 12 setae ; pronotal
ctenidium consisting of the same number of
similar spines, but the ctenidium is strongly
curved and the 4 dorsal spines on each side
are directed backwards and downwards, also
the second and third from above are much
less spatulate.

Mesonotum with a main row of 10 setae
on both sides together, preceded by a row
of 10 smaller setae ; these two rows are
arranged obliquely. Mesosternosome with
8-9 setae ; anterior margin of mesepi-
sternum more oblique.

Metanotum with a main row of 12 setae
preceded dorsally by only 4 small setae ;
its posterior margin bears dorsally two
apical spinelets on each side ; metepi-
sternum with one long and one short seta ;
metepimeron with 5 setae on one side and
6 on the other.

LEGS (Figs. 49, 50). Hind tibia with an
indistinct false comb of 8 setae along the
posterior margin ; the five pairs of lateral
plantar setae on the fifth segment of the
hind tarsus are rather slender, on the plantar
surface about 20 minute setae.

Metanotum with a main row of 10 setae,
preceded by a row of 8-10 setae ; its pos-
terior margin bears dorsally 3 apical spinelets
on each side ; metepisternum also with one
long and one short seta ; metepimeron
extending upwards much nearer to dorsum,
with 6 setae.

LEGS (Figs. 56, 57). The setae of the
hind tibia forming a false comb number 12
and are stouter, hence the false comb is very
distinct ; the five pairs of lateral plantar
setae are stouter and there are about 30
minute setae on the plantar surface.

2i8 DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA

Caenopsylla assimulata (Weiss),

Caenopsylla mira Rothschild,

DESCRIPTION

ABDOMEN. Terga I-III with 2-2, i-i,
i -i apical spinelets respectively ; terga
I-VII normally respectively each with 4, 6,
6, 6, 6, 6, 6 setae in the main row on each
side ; the setae on tergum VII are all about
equally spaced. In front of the main row
of terga I-III respectively 2-3, 3-4, 1-3
small setae, none at all on terga IV-VII.
Similar antesensilial setae. Basal sternum
with one seta along the ventral margin on
each side, sterna III-VII each with 2 or 3
setae on each side, sternum VIII as in Fig. 61,
its apical margin not sinuate.

MODIFIED SEGMENTS (Figs. 59, 61).
Tergum VIII roughly triangular, its dorsal
margin undulate, and apically this margin
and the ventral one meet at a sharp angle ;
2 long setae near the dorsal margin. Manu-
brium much broader and tapering strongly
apically. Corpus of clasper with a deep
sinus in its dorsal margin, which divides off
the fixed process as a very distinct lobe,
bearing at its apex 5 slender setae ; most of
the posterior margin of the clasper slightly
convex ; 4 acetabular setae. Movable pro-
cess elongate, four times as long as its
maximum width ; chaetotaxy similar, but
the long seta at the posterior margin is
inserted at one-fourth the length of the
posterior margin from the apex. Apex of
proximal arm of sternum IX triangular, the
narrow arm widening only a little towards
its junction with the distal arm. The distal
arm bears ventrally a large number of small,
medium-sized and long setae ; its divided-off
apical part triangular, very differently
shaped to that of the other species, details
as in Fig. 59 ; the membranous flap is much
larger. Aedeagal apodeme much narrower ;
for details of the phallosome, which is very
different, see Fig. 60.

LENGTH. 6*. 2-2 i mm - ' ? 2 i~ 2 i mm. 1
HABITAT. Biskra and El Kantara, both
in Algeria.

HOST. Ctenodactylus gundi (occurring in
Libya, Tunis, Algeria, and west to the
Moroccan Atlas) ; the occurrence of one
specimen on EUomys quercimts munbyanus
may be accidental.

1 Although the female sex of this species is not dealt with here, it seemed useful to give a figure of its
terminalia (Fig. 62).

ABDOMEN. Terga I and II each with one
apical spinelet each side ; terga I-VII res-
pectively each with 4, 6, 6, 6, 6, 5, 4 setae in
the main row on each side ; on tergum VII
the gap between the lowest seta and its
neighbour is twice as long as the space
between the ist and 2nd or 2nd and 3rd.
In front of the main row one seta on terga I
and II, none at all on terga III-VII. Tergum
VII with 3 antesensilial setae the lowest of
which is very small. Basal sternum appar-
ently without setae, sterna III-VII on each
side with 2 setae, sternum VIII as in Fig. 52,
its apical margin with a sinus.

MODIFIED SEGMENTS (Fig. 51). Shape of
tergum VIII not visible in the specimen, but
it bears 3 long setae. Manubrium of
clasper long and narrow, not tapering in its
apical portion. Corpus of clasper with sub-
parallel dorsal and ventral margins, its apical
half directed upwards ; 4 slender setae along
the dorsal margin near the apex ; posterior
margin of clasper slightly concave ; 2 aceta-
bular setae. Movable process spindle-shaped,
two and a half times as long as its maximum
width ; besides a number of small setae it
bears a fairly large one at the posterior
margin at one-third its length from the apex.
Apex of proximal arm of sternum IX squarish ;
the greater part of this arm very narrow but
it widens to a triangle with a rather long
base. The distal arm bears ventrally 5 long
setae, and its structure is complicated and is
shaped as shown in Fig. 51 ; the membranous
flap projecting from behind the apex may
belong to this sternum or it may be the
aedeagal crochet ; this could not be ascer-
tained, no material being available for
dissection. Aedeagal apodeme broad
details of the phallosome, see Fig. 53.

for

LENGTH.
HABITAT.

i mm.

Matmata, South Tunisia.

HOST. Elephantulus rozeti (which was
collected by Weiss in March, 1913) ; it may
be expected to occur on the Gundi.

DESCRIPTIONS OF NEW AND LITTLE-KNOWN SIPHONAPTERA 219

REMARKS

The difference in the development of the pronotum is of interest : in C. assimulata
we see that the pronotum is still of a more or less normal shape, but its ctenidium
shows a clear tendency towards the condition found in C. mira, where the pronotum
is dorsally widened backwards, and as a result of this the ctenidium is strongly
curved and the upper spines tend to point downwards. These are obviously two
stages in the evolution of the pronotum in the genus Caenopsylla, and it would be
of interest to know whether a species of this genus with the original straight pronotal
spines still exists, and also whether or not the evolution in another species has gone
further, producing a pronotum more or less like that of Barrewpsylla. In order to
find the answer to this question it will be necessary to collect intensively throughout
North Africa, which may unfortunately mean a delay of many years.

REFERENCES

DAMPF, A. 1945. Notas sobre Pulgas. Rev. Soc. Mex. Hist, nat., 6 : 68.

JORDAN, K. 1931. Further records and descriptions of fleas from Ecuador. Novit. Zool.,

37 : 138.
WAGNER, J., & WASSILIEFF, A. 1933. Tableaux analytiques pour la determination des puces

rencontrees en Algerie et en Tunisie. Arch. Inst. Pasteur Tunis, 21 : 452.
WAHLGREN, E. 1904. Zwei neue Puliciden aus Aegypten. Results Swed. zool. Exped. Egypt

and White Nile 1901 : 3-4.
WEISS, A. 1913. Contribution a 1'etude des pulicides. Une puce nouvelle de la Tunisie

(Typhlopsylla assimulatus n. sp.). Arch. Inst. Pasteur Tunis, 3 : 187-196, figs. 1-3.

PRESENTED

1 7 DEC 1953

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

1 6 MAR 1954

THE EARLY LITERATURE
ON MALLOPHAGA

(PART III)

THERESA CLAY

AND

G. H. E. HOPKINS

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 3 No. 6

LONDON: 1954

THE EARLY LITERATURE ON MALLOPHAGA

BY

THERESA CLAY

AND

G. H. E. HOPKINS

PART III. 1776-1786

Pp. 221-266 ; Pis. 10-12 ; 68 Text-figures.

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 3 No. 6

LONDON: 1954

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

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

This paper is Vol. 3, No. 6 of the Entomological
series.

Issued March, 1954. Price Ten Shillings.

THE EARLY LITERATURE OF MALLOPHAGA

By THERESA CLAY AND G. H. E. HOPKINS

(PART III, 1776-1786)

SYNOPSIS

This part, the third of a series of papers in which it is intended to review the species of Mallo-
phaga described between 1758 and 1818, deals with the twelve species described by F. Schrank
in 1776 and 1781 ; the two species by O. F. Miiller in 1776 ; the single species by J. C. Fabricius
in 1777 ; the seven species by De Geer in 1778 ; the seven species by O. Fabricius in 1780 ; the
three species by A. F. Fourcroy in 1785 ; the single species described by Mohr in 1786. 1 As in
previous instalments, the references following the headings are to the original work.

SCHRANK, 1776
(Beytrdge zur Naturgeschichte, Leipzig : 113-120.)

WITH Schrank we reach much firmer ground than we have trodden hitherto, for all
his species are quite reasonably well figured and there can be little doubt about the
identity of any of them. All have, of course, been renamed (singly or in groups) by
Nitzsch. The new species, all of which were published in perfectly valid form, are
discussed below.

Pediculus curuccae (p. 113, PI. 5, fig. i).

The host is " Motacilla curucca " (= Sylvia curucca), and the figure shows a
Menoponid resembling Menacanthus rather than Myrsidea. No subsequent author
has anything particularly useful to say about it, but Nitzsch (1818 : 300) renamed it
Liotheum (Menopon) minutum. As no material from the type host is available, neo-
types of Menacanthus curuccae (Schrank) cannot be erected.

Pediculus rubeculae (p. 115, PI. 5, fig. 4)

No host is mentioned in the description, but the species is called " Rothkehlchen-
laus," which is sufficient indication that the host was Erithacus rubecula ; in a later
work (1781 : 507) Schrank gives the host as Motacilla rubecula. The figure is a
reasonably good representation of a Ricinus and could not be mistaken for anything
else.

1 In the last instalment of this series we remarked (Bull. Brit. Mus. (Nat. Hist.) Entomology, 2 : 4)
that " K61er 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." This remark is clumsily worded and suggests a reproach to Dr. von Keler that we certainly did not
intend. In ascribing the authorship of equi to Linne he was in conformity with nearly all recent writers
on Mallophaga, since, with extremely few exceptions all have either ignored or overlooked the fact that
Pediculus equi Linnd is a nomen nudum.

ENTOM. Ill, 6. 1 8

224

THE EARLY LITERATURE ON MALLOPHAGA

The species was again described by Olfers (1816 : 91) as Nirmus pterocephalus,
a name which covers several species but which we are restricting to Ricinus rubeculae,
and by Nitzsch (1866 : 121) as Physostomum agonum, both from the same host as
Schrank's material. Harrison (1916 : 18, 68) correctly restores Schrank's name for
the species, but then (p. 103) wrongly mentions it in Philopterus as well, probably
through confusion with Philopterus rubeculae (Denny), which Harrison omits and
which is not congeneric with rubeculae Schrank.

This species is at once distinguished from both dolichocephalus (Scopoli) (Clay and
Hopkins, 1951 : 10) and fringillae De Geer (see below) by the form of the mandibles
(Text-fig. 4). It is further distinguished from the former species by the shape of
the head (PI. 10, figs, i, 2), the terminal segments of the female abdomen (Text-fig. 3)

FIGS, i, 2. Ricinus rubeculae (Schrank). i. Gular and thoracic plates, female.

2. Male genitalia.

FIGS. 3, 4. Ricinus rubeculae (Schrank). 3. Terminal segments of female abdomen.

4. Mandible.

and prothorax, the form of the gular and prothoracic plates (Text-fig, i), the male
genitalia (Text-fig. 2), and by the absence of the colour pattern characteristic of
dolichocephalus. From fringillae it is distinguished in the female by the shape
of the head and prothorax, by the form of the gular and prothoracic plates, the
number of gular setae, and by the chaetotaxy of the terminal segments of the female
abdomen. No males of fringillae have been seen.

Neotype male (PI. 10, fig. i ; Text-fig. 2) and neallotype female (PL i, fig. 2 ; Text-
figs, i, 3, 4) of Ricinus rubeculae (Schrank) in the British Museum (Nat. Hist.) (slide
no. 562) from Erithacus r. rubecula from Moravia, Czechoslovakia. Neoparatypes :
17 males and 50 females from Erithacus rubecula melophilus Hart, from the British
Isles.

THE EARLY LITERATURE ON MALLOPHAGA

225

MEASUREMENTS IN MM.

Head .
Prothorax
Abdomen
Total .
Genitalia

Male

Length
0.58

Breadth

o-55

0-70

2-34
0-38

Female

Length Breadth
0-67 0-63

0-52
0-88

3-02

FIG. 5. Menacanthi4s alaudae (Schrank), male.

226

THE EARLY LITERATURE ON MALLOPHAGA

Pediculus alaudae (p. 115, PL 5, figs. 5, 6)

Another obvious Menacanthus. Although there is no explicit host-record, Schrank
calls the species " Feldlerchenlaus " and in 1781 (p. 506) he gives the host as Alauda
arvensis.

The same species was redescribed, from the same host, by Piaget (1880 : 446) as
Menopon parviceps (see Clay, 1949 : 904).

FIG. 6. Menacanthiis alaudae (Schrank), male genitalia.
MEASUREMENTS IN MM. :

Male

Female

r "

Length

Breadth

0-25
0-12

o-39

0-28

0-83

o-55

1-22

r

Length

Breadth

0-27

0'43

0-15

0-32

0-40

1-03

0-72

i-55

Head .

Prothorax

Metathorax

Abdomen

Total .

Genitalia

Neotype male (Text-figs. 5, 6) and neallotype female (Text-fig. 7 ; PI. i, fig. 3) of
Menacanthus alaudae (Schrank) in the Meinertzhagen Collection (British Museum
(Nat. Hist.)) (slide no. 327oa) from Alauda a, arvensis Linn, from Ushant, France.

THE EARLY LITERATURE ON MALLOPHAGA

227

Neoparatypes : 9 males and 33 females from various subspecies of Alauda arvensis
from France, British Isles, Sweden and Yugoslavia.

Pediculus citrinellae (p. 116, PL 5, fig. 7)

The figure represents a Philopterus, apparently a nymph, and the host (" Ammer,
Goldammer, Aimerling ") is given by Schrank in 1781 (p. 507) as Emberiza citrinella.

FIG. 7. Menacanthus alaudae (Schrank), terminal segments of female abdomen.

8

FIGS. 8, 9. Philopterus c. citrinellae (Schrank), male. 8. Anterior region of head. 9. Genitalia.

Nitzsch (1818 : 290) proposed Philopterus (Docophorus) communis zsanomen novum
for Ricinus Emberizae De Geer, Pediculus curvirostrae Schrank, Pediculus Pynhulae,
Citrinellae and Chloridis Schrank, and Nirmus globifer Olfers. As all these names
are nomenclatorially valid, the only effect of Nitzsch's action was to create confusion
in the nomenclature and to compromise hopelessly the name communis, Ricinus

228

THE EARLY LITERATURE ON MALLOPHAGA

emberizae De Geer is a synonym of Pediculus citrinellae Schrank, and Nirmus globifer,
though composite, is primarily a new name for the same species. As three of the
names combined by Nitzsch under communis refer to citrinellae, we think it only
reasonable to restrict communis to the Philopterus found on Emberiza citrinella ;
we therefore select as neotype of Philopterus communis Nitzsch, 1818, the neotype

a

10

FIG. 10. Philopterus c. citrinellae (Schrank), genital plates of four females taken from one host

individual.

If

FIG. ii. Philoptenis c. citrinellae (Schrank), female genital region, a. Last sternal plate of

P. citrinellae curvirostrae (Schrank).

of Pediculus citrinellae Schrank, 1776. Philopterus communis Nitzsch thus becomes
a synonym of P. citrinellae (Schrank), and it will only receive a bare mention under
its other original components. Ricinus emberizae de Geer and Nirmus globifer Olfers,
both of which are synonyms of citrinellae, will be dealt with in discussing the work
of these authors. The identities of the insects described by Schrank as Pediculus

THE EARLY LITERATURE ON MALLOPHAGA 229

citrinellae, P. curvirostme, P. Pyrrhulae and P. chloridis will be discussed together
after our account of the last-named.

Neotype male (PI. 10, fig. 4; Text-figs. 8, 9) and neallotype female (Text-figs. 10, n)
of Philopterus citrinellae (Schrank) in the British Museum (Nat. Hist.) (slides nos.
565-6) from Emberiza c. citrinella Linn, from Saxony, Germany. Neoparatypes :
28 males and 42 females from various subspecies of Emberiza citrinella from Yugo-
slavia, Estonia, France and the British Isles.

Pediculus curvirostrae (p. 117, PI. 5, fig. 8)

The description and figure are those of a Philopterus and the host is given as Loxia
curvirostra Linn. No early author adds anything to our knowledge of the species,
but it is one of the components of Philopterus communis Nitzsch, 1818. Piaget in
1880 described Docophorus compar from the same host ; his syntypes, two males
and two females (slides nos. 538-9), agree with our neotypes of curvirostrae.

Neotype male (as described below) and neallotype female (Text-fig. n#) of Philop-
terus curvirostrae (Schrank) in the Meinertzhagen Collection (British Museum (Nat.
Hist.) (slides nos. 4112 and 4111)) from Loxia c. curvirostra Linn, from Poland. Neo-
paratypes : 24 males and 36 females from the same host form from Poland, Estonia
and England.

Pediculus pyrrhulae (p. 117, PL 5, fig. 9)

The host is " Blutfink " (= Pyrrhula pyrrhula) and the louse is a nymph of a
Philopterus. It is one of the components of Philopterus communis Nitzsch. Piaget
(1880 : 58) described independently a Docophorus communis var. pyrrhulae from
Pyrrhula vulgaris (= Pyrrhula pyrrhula). Piaget's type material in the British
Museum (Nat. Hist.) (one male and two females, slide no. 536) agrees with the neo-
types of Philopterus pyrrhulae (Schrank).

Neotype male and neallotype female (as described below) of Philopterus pyrrhulae
(Schrank) in the Meinertzhagen Collection (British Museum (Nat. Hist.) (slide no.
1042^)) from Pyrrhula pyrrhula europaea Vieillot from France. Neoparatypes : 12
males and 36 females from various subspecies of Pyrrhula pyrrhula from France,
Sweden, Estonia and the British Isles.

Pediculus chloridis (p. 118, PL 5, fig. 10)

Another Philopterus nymph, from " Griinfink," the host given by Schrank later
(1781 : 506) as Loxia Moris ; it is also another of the components of Philopterus
communis Nitzsch.

Neotype male and neallotype female (as described below) of Philopterus chloridis
(Schrank) in the Meinertzhagen Collection (British Museum (Nat. Hist.)) (slide no.
290) from Chloris c. chloris (Linn.) from Norfolk, England. Neoparatypes : 32
males and 60 females from the same host form from the British Isles and Estonia.

230 THE EARLY LITERATURE ON MALLOPHAGA

Philopterus citrinellae, P. curvirostrae, P. pyrrhulae and P. chloridis

These names were given to specimens of Philopterus taken respectively from Em-
beriza citrinella, Loxia curvirostra, Pyrrhula pyrrhula and Chloris Moris and can be
considered together. While discussing their status the names will be used to refer
to populations from these hosts even though in the conclusion they may be con-
sidered as indistinguishable.

The characters on which the species of Philopterus from the small European
Passeres can be distinguished have been discussed in Part II of the present work
(Clay & Hopkins, 1951). The four forms listed above parasitize the bird family
Fringillidae and the possible distinguishing characters between these populations
are discussed below.

HEAD. This is of the typical Philopterus form similar to that figured in Clay &
Hopkins, 1951 : 7 for P. coarctatus, from which it differs in the median indentation
of the hyaline margin (not always apparent in mounted specimens) and the somewhat
deeper indentation of the anterior plate (clypeal signature) (Text-fig. 8) . P. frin-
gillae from Passer domesticus also has both the hyaline margin and anterior plate
indented, but the pre-antennal region and the length of the anterior plate are pro-
portionally shorter. Specimens from the four hosts under consideration show no
constant differences in shape of the anterior plate nor in the proportions 1 of the
head. A comparison of the size of the head (as shown by the absolute breadth at
the temples, Tables i, 2) shows that there is some variation in specimens taken from
one host species, but the means fall into a number of groups. Thus, in the males
citrinellae, curvirostrae and chloridis fall into one group, with pyrrhulae tending to be
larger, but the number of specimens of the latter measured is rather small. In the
females of P. citrinellae and curvirostrae the mean tends to be smaller than in those
of chloridis and pyrrhulae. Further material will have to be examined before it
can be shown whether or not these differences are significant.

MALE GENITALIA (Text-fig. 9). The characters show individual variation as well
as differences in appearance due to the position in which the genitalia are viewed.
This individual variation affects the shape of the parameres and endomeres, number
and position of the setae on the mesosome and the presence (and size) or absence of
the dorsal sclerotized area on the mesosome. If sufficient specimens are examined
genitalia of similar appearance will be found irrespective of the host from which the
specimens were taken. However, the endomeres are minute structures and their
true form can probably only be reconstructed from sections, so it may prove that
these structures will show subspecific variation when more thoroughly examined.
On the available material there seem to be no characters in the genitalia by which

1 To find the various proportions of the head (e.g., length of preantennal region to that of the post-
antennal region or breadth of various parts of the preantennal region to the length of preantennal
region) entails the making of a number of detailed measurements from a large number of specimens,
and these measurements tend to be inaccurate owing to the difficulty of delineating the different regions
of the head. However, if the image of a typical specimen from one host is projected on to a piece of
paper and an outline of the head made, it is possible to project other heads on to this outline, and by
adjusting the magnification to get one fixed breadth (say that of the temples) the same ; from this it
is possible to compare the various proportions of the head and find the variation within a number of
specimens from one host and between those from different hosts.

THE EARLY LITERATURE ON MALLOPHAGA 231

the populations from the four hosts can be separated. The genitalia of P. coarc-
tatus (Scopoli) (see Clay and Hopkins, 1951 : 6) and P. fringillae (Ibid : 32) are
readily separable on the characters of the mesosome.

The diagnostic characters and the type of variation found in the female genital
region have been discussed under coarctatus (see Clay & Hopkins, 1951 : 6).
Specimens from the four hosts under discussion cannot be separated by the shape
of the genital plate, although the mean of the breadth of the plates of citrinellae is
smaller than in the other three. In general proportions the plate is similar to that
of coarctatus and differs from that of fringillae as discussed under that species (see
Clay & Hopkins, 1951 : 32). There is some variation in the shape of the genital
plate, as shown by the four examples taken from one host individual in Text-fig. 10,
and also in the size and shape of the last sternal plate.

CHAETOTAXY. There is some individual variation found throughout the different
populations in the number of setae (especially in those on the abdomen), but there
are three groups of setae which may show subspecific characters ; these are the setae
of the metasternum, those of the male genital plate, and the group each side of the
last sternal plate of the female abdomen. In citrinellae 80-4% of 46 males and females
examined had 3 metasternal setae, 17-4% had 4, and 2-2% na d 5. In pyrrhulae
91-2% of 34 examined had 3 metasternal setae and 8-8% had 4. In chloridis 84-6%
of 52 specimens had 3 setae and 15-4% had 4. In curvirostrae 4-2% of 48 examined
had 3 setae, 45-8% had 4, 45-8% had 5 and 4-2% had 6. Thus, while curvirostrae
has 95-8% of specimens with 4 or more setae, the other three have over 80% of
specimens with only three setae.

The male genital plate in citrinellae has two anterior and two posterior setae as in
P. coarctatus (see Clay & Hopkins, 1951, fig. 4) ; the two anterior setae are found in
100% of specimens of citrinellae (29 examined), pyrrhulae (13 examined) and chloridis
(26 examined). In curvirostrae, however, 100% of the 21 specimens examined had
3 (or rarely 4) anterior setae on the male genital plate.

Each side of the last sternite of the female abdomen there are in citrinellae usually

2 normal setae and one spine-like seta (Text-fig, n). In citrinellae 97-2% of the
36 specimens examined had these 3 setae (rarely reduced to 2) on at least one side
of the abdomen, only 2-8% had 4 setae on both sides of the abdomen. In pyrrhulae
95' l0 /o of the 41 examined had 3 or less setae on at least one side, and 4-9% had 4
setae on both sides. In chloridis 100% of the 64 examined had 3 setae or less on at
least one side. In curvirostrae 100% of the 32 examined had 4 or more setae on
both sides (Text-fig, na). Thus, while 100% of specimens of curvirostrae had 4 or
more setae on both sides of the abdomen, the other three had 95% or more with only

3 on at least one side.

On the evidence discussed above, populations from the four hosts obviously form
a single species. The populations from Loxia curvirostra can be distinguished in
the male in 100% of specimens (on the material examined) on one character and in
80% of specimens on a second character ; the females from this host can be dis-
tinguished in 95% of cases on one character and in 80% on a second character.

The populations from Loxia curvirostra can, therefore, be recognized as a sub-
species. If subsequent investigations reveal subspecific differences in the other

232 THE EARLY LITERATURE ON MALLOPHAGA

populations Schrank's names are available for these, but for the present we are unable
to recognize them. The host distribution of the two subspecies we recognize is as
follows :

Philopterus citrinellae citrinellae (Schrank). Type host : Emberiza c. citrinella.
Also occurs on Pyrrhula pyrrhula and Chloris Moris.

Philopterus citrinellae curvirostrae (Schrank). Type host : Loxia c. curvirostra.

MEASUREMENTS IN MM. of neotypes of P. c. citrinellae :

Male Female

Length Breadth Length Breadth

Head . . . 0-52 0-47 . 0-55 0-53

Prothorax . 0-28 0-30

Pterothorax . 0-43 0-47

Abdomen . . 0-62 . 0*75

Total ... 1-46 . 2-67

Genitalia . . 0-22

Pediculus sturni (p. 118, PI. 5, figs. 11-14)

A nymph of Sturnidoecus from " Staar " (given by Schrank in 1781 as Sturnus
vulgaris}. Schrank considered that his fig. 12 represented a different species, but
it is merely a still younger nymph. Nitzsch (1818 : 290) unnecessarily renamed the
species Philopterus (Docophorus) leontodon, and it was usually known by this name
until Harrison (1916 : 104) restored the name given to it by Schrank.

MEASUREMENTS IN MM. :

Male Female

Length Breadth Length Breadth

Head . . . 0-52 0-53 . 0-55 0-57

Prothorax . 0-30 0-32

Pterothorax. . 0-47 0-52

Abdomen . . 0-71 0-70 . 0-96 0-83

Total . . . 1-52 . 1-80

Genitalia 0-35

Neotype male (Text-figs. 12-13 ; PI. 10, fig. 5) and neallotype female (Text-fig. 14)
of Sturnidoecus sturni (Schrank) in the Meinertzhagen Collection (British Museum
(Nat. Hist.)), slide no. 15663, from Sturnus v. vulgaris Linn, from Cornwall, England.
Neoparatypes : 138 males and 175 females from various subspecies of Sturnus
vulgaris from the British Isles and Estonia.

O. F. MOLLER, 1776

(Zoologiae Danicae Prodromus. Havniae : pp. 184-186)

This is quite the worst of the early works, but the names contained in it are bi-
nomial and must be considered. Fortunately there are only two new names in it,

THE EARLY LITERATURE ON MALLOPHAGA

233

which both might be thought to apply to Mallophaga and also are accompanied by a
description or an indication. Harrison (1916) adds a third, Pediculus strigis, which
he considers to be a nomen nudum, but it is neither a nomen nudum nor a new name,
but a reference to strigis Pontoppidan, q.v. (Clay & Hopkins, 1951, p. 29).

12

FIG. 12. Sturnidoccits sturni (Schrank), male genitalia. a. Distal ends of parameres in natural

position.

Pediculus alcae arcticae (p. 185)

The entire description is " e cinereo caerulescens," but in the case of both this and
the following species there are references to " Isl.R." (= Olafsen, 1772). We have
not been able to consult the original of Olafsen's work but have seen a German
translation dated 1774, on p. 322 of which there are much fuller descriptions of both

234

THE EARLY LITERATURE ON MALLOPHAGA

the species in question, under vernacular names. Olafsen's description of his
" Lunda-Luus, Pediculus (Alcae arcticae) " includes a vivid account of its habits
which leaves no room for doubt that it is a tick.

Pediculus colytnbi grylle (p. 185)

Miiller's entire description is " capite fusco, abdomine glauco," but Olafsen's
fuller description of his "Hofu-Luus " states, among other points, that it is rather
large, flat and oblong like Hippobosca (now Melophagus) ovina. This description
could not be applied to any of the Mallophaga found on Alcidae, but would very well
fit a tick.

13

FIGS. 13, 14. Sturnidoecus sturni (Schrank) . 13. Male head.

abdomen, ventral.

14. Terminal segments of female

J. C. FABRICIUS, 1777*
(Genera insectorum eommque characteres naturales, etc. Chilonii, pp. 309-310)

Of the new names proposed in this work all, with one exception, are unnecessary
renamings of Scopoli's species and have already been dealt with under the latter
author. The descriptions are quoted from Scopoli, and the neotypes we have erected
for Scopoli's species are also neotypes of Fabricius' nomina nova. The single excep-
tion is dealt with below.

Pediculus phasiani (p. 310)

The host-record is " Habitat in Phasiano Motmot," and besides the very brief
description there is a reference to " Red.Oper.I." Redi's plate i shows three species,
belonging to as many genera, stated to be " Pollini dell' astore." In spite of Redi's
belief that all of these were hawk-parasites, the reference has enabled us to interpret
Fabricius' name, for his inadequate description does not fit the upper figure on the

* According to Engelmann, Bibl. Hist. -Nat., 1846, p. 479.

THE EARLY LITERATURE ON MALLOPHAGA 235

plate and the right-hand lower figure is a Craspedorrhynchus , a genus which does
not occur on the Galliformes. But Fabricius' description does fit the left-hand
lower figure on the plate, which represents a Chelopistes, a genus known to occur on
the Ortalidae. As we have no material from Ortalis motmot we v are unable to erect
neotypes for Chelopistes phasiani (J. C. Fabricius).

DE GEER, 1778

(Memoires pour servir a I'histoire des insectes. Stockholm. Vol. 7, Des Ricins,

pp. 69-82)

The descriptions in this work are good and the figures for the most part excellent,
but we were inclined at first to think De Geer's classification not binomial (though
certainly binary) and the names invalid in consequence. This is because the generic
name in each case is followed by a long descriptive phrase or diagnosis, with or without
a comma after the name of the host (which always immediately follows the generic
name Ricinus and is placed in the genitive case). We have, however, consulted
Dr. Jordan on this point, and he kindly informs us that in his opinion the names are
valid. He states : " The Latin diagnosis of the various species of Ricinus differs
from the diagnoses of all other insects in vol. 7 and previous volumes in the trivial
names of the species not being put into brackets, and in the case of Ricinus the
genitive of the host-name is in italics like the Latin text of the diagnosis, not in
Roman type. The work was issued after De Geer's death, and the omission of the
brackets and the consequent italicizing of the trivial names was probably due to the
editor. The diagnosis of species i of Ricinus should read : "Ricinus (Fringillae)
pallide-fuscus , corpore . . . ' We accept Dr. Jordan's opinion on this point
with all the more pleasure because De Geer's work is so outstandingly better than
that of his contemporaries that it would be very regrettable if technical invalidity
of the names he proposed should deprive him of the credit for it.

With regard to the statement of Harrison (1916 : 10-20) that Pediculus Linn.,
Ricinus De Geer and Nirmus Hermann " must be treated as equivalent, inasmuch
as each included all Mallophaga," and his action in rejecting (e.g.] Ricinus fringillae
De Geer as a homonym of Pediculus fringillae Scopoli, the revised Rules of Zoological
Nomenclature make it clear that Harrison's opinion was incorrect and that both
these names may be used.

Ricinus fringillae (p. 71, PI. 4, figs. 5-8)

The description and figures are good, and undoubtedly represent a species belong-
ing to the genus Ricinus s. sir. (= Physostomum Nitzsch). The question of the host,
however, is one of some difficulty. De Geer's description in Latin is preceded by
one in French, in which he calls the louse " Ricin du Ph^on " (= Fringilla coelebs},
but he later states that he found it in March on " 1'oiseau nomme Bruant " (=
Emberiza citrinella], and that it also lives " sur les Pingons & d'autres petits oiseaux ";
the species of Ricinus found on Fringilla coelebs and on Emberiza citrinella, though very
closely related, are not the same, and De Geer's figures resemble the latter rather than

236

THE EARLY LITERATURE ON MALLOPHAGA

the former in the proportions of the head. Nevertheless, if there were no complica-
tions we might have taken the view that De Geer's calling the louse " Ricin du
Pingon " compelled us to apply his name to the species found on Fringilla coelebs, but
there is a very strong reason for reluctance to adopt this attitude : Nitzsch (1818 : 302)
published the name Liotheum (Physostomum) nitidissimum as a nomen novum for
Ricinus fringillae De Geer, with Ember iza citrinella as host, and the name Liotheum
(Physostomum) irascens (a nomen nudum) with Fringilla coelebs as host ; irascens was
published with a description by Burmeister (1838 : 442) and all later authors have
used nitidissimus and irascens for the species of Ricinus found on Emberiza citrinella
and Fringilla coelebs respectively. To select Fringilla coelebs as type-host of Ricinus
fringillae De Geer would mean that the host of nitidissimum Nitzsch, 1818 (whose

FIG. 15. Ricinus fringillae De Geer, Gular and thoracic plates, i'emale.

sole claim to validity is the reference to De Geer), would of necessity be the same bird
and the Ricinus of Emberiza citrinella would be left without a name. We consider
that this course would cause so much confusion that we must select the species from
Emberiza citrinella to bear the name that De Geer gave to the composite of both
species. De Geer's species is also a component of Nirmus pterocephalus Olfers, 1816,
but we consider this latter composite name to refer mainly to Ricinus rubeculae
(Schrank).

The characters by which R. fringillae can be distinguished from rubeculae are given
above under that species. This species is separated from others occurring on the
Fringillidae by the shape of the head and prothorax (PI. 10, fig. 6) and by the pro-
thoracic plate (fig. 15), and from some species by the characters of the mandibles
(fig. I?)-

THE EARLY LITERATURE ON MALLOPHAGA
MEASUREMENTS IN MM. :

237

Female

Head .
Prothorax
Abdomen
Total

Length
0-70

Breadth
0-68
0-52
1-07

Neotype female (Text-figs. 15-17 ; PL 10, fig. 6) of Ricinus fringillae De Geer in
the British Museum (Nat. Hist.) (slide no. 563) from Emberiza c. citrinella Linn,
from Saxony, Germany. Neoparatypes: 4 females from the same host species from
England, Estonia and Czechoslovakia.

16

FIGS. 1 6, 17. Ricinus fringillae De Geer, female. 16. Terminal segments of abdomen.

17. Tips of mandibles.

Ricinus emberizae (p. 74, pi. 4, figs. 9, 10)

The host is " Bruant," Emberiza or "Groning " (= Emberiza citrinella), and the
description and figure are those of a Philoptems. We consider this to be a synonym
of Philopterus c. citrinellae (Schrank). Doubtless De Geer was misled into thinking
it new by the fact that Schrank's species was described from a nymph.

Neotype of Philopterus emberizae (De Geer) a male in the British Museum (Nat.
Hist.) collection (slide no. 564) from Emberiza c, citrinella Linn, from Yugoslavia,
which agrees with the neotype of Philopterus c. citrinellae (Schrank).

Ricinus cornicis (p. 76, PI. 4, fig. n)

De Geer places " Pediculus Corvi Coracis Linn." as a synonym, but the species has
nothing to do with Philopterus corvi (Linn.), both figure and description very clearly
referring to a Myrsidea. The host-record is " Corneille " (= Corvus cor one comix}.

Nitzsch (1818 : 300) renamed the species Liotheum (Menopon) mesoleucum, quoting

ENTOM. Ill, 6. 10

238 THE EARLY LITERATURE ON MALLOPHAGA

De Geer and giving the host-record "Corvi Cornicis." Harrison (1916 : 13) erro-
neously considered Ricinus cornicis De Geer (a Myrsidea) to be preoccupied by Pedi-
culus cornicis J. C. Fabricius (a Philopterus)', he also took the erroneous view that
mesoleucum Nitzsch, 1818, is a nomen nudum, and in consequence the species has
commonly been known as Myrsidea subaequalis (Haan*). The neotypes we erect for
Myrsidea cornicis (De Geer) are necessarily also neotypes of M. mesoleuca (Nitzsch).

/ // l/^HH^f "\\ \

I II /NJIXA \\ \

18

FIG. 1 8. Myrsidea cornicis (De Geer), female.

This species can be distinguished from other Myrsidea occurring on the European
species of Corvus by the greater modification of tergites I-II (Text-fig. 18) in the
female and by the sclerite of the genital sac in the male (Text-fig. 20). The
chaetotaxy of sternite II (Text-fig. 22) in both sexes resembles most closely that of
the species on Corvus monedula, but cornicis can be distinguished by the shape of the
head in the male (PI. n, fig. i) and by the characters given above.

* The authorship of the name is often incorrectly ascribed to Nitzsch or to Lyonet.

THE EARLY LITERATURE ON MALLOPHAGA

239

MEASUREMENTS IN MM.

Head .

Prothorax

Metathorax

Abdomen

Total .

Genitalia

Male

Length

o-33

1-89
0-66

Breadth
0'6o
0-40

o-54
0-67

Female

Length

o-35

2-14

Breadth
0-65
0-41

0-68
0-87

Neotype female (Text-fig. 18) and necdlotype male (Text-figs. 19-22, PL 10, fig. i) of
Myrsidea cornicis (De Geer) in the Meinertzhagen Collection (British Museum (Nat.
Hist.)) (slide no. i6oi2a) from Corvus corone comix Linn, from Sweden. Neopara-
types : 174 males and 143 females from the same host form from Sweden, Estonia, E.
Prussia, Hungary, Yugoslavia and the British Isles.

23

Myrsidea cornicis (De Geer), male. 19. Male genitalia. 20. Sclerite of genital
sac. 21. Tip of paramere enlarged. 22. Second abdominal sternite.
Male heads, dorsal. 23. Anaticola crassicornis (Scopoli). 24. A. mergiserrati

(De Geer).

Ricinus lari (p. 77, PI. 4, fig. 12)

There are references to " Pediculus Sternae Hirundinis Linn." and " Redi Experim.
Tab. 9," though neither of these species is even congeneric with that described by De
Geer, both his figure and his description being quite certainly those of a Trinoton.
The host-record " sur les Mouettes " (= Larus spp.) is definitely erroneous, for neither
Trinoton nor any genus closely resembling it occurs naturally on the Laridae. 1

1 We considered the large type of Actornithophilus occuring on the Laridae, but the figure does not
agree with this.

240

THE EARLY LITERATURE ON MALLOPHAGA

J. C. Fabricius (1781 : 481) erroneously placed De Geer's name as a synonym of
Pediculus sternae Linn., doubtless misled by De Geer's reference to this species.
Nitzsch (1818 : 300) considered De Geer's species to be probably the same as his
Liotheum (Trinotori) lituratum (a nomen nudum in 1818), from Mergus albellus, pro-
bably because another of De Geer's species is from Mergus serratus.

We see no reason whatever to adopt Nitzsch's belief, De Geer's figure not being
sufficiently detailed to refer to one species of Trinoton rather than another, and the
common-sense procedure with regard to a species described from a straggler is ob-
viously to make it a synonym of some earlier name if possible. Fortunately Trinoton
querquedulae (Linn.) is a much earlier valid name and there is nothing in De Geer's
figure and description to suggest that this was not the species he had.

27

25

28

FIGS. 25-28. 25. Anaticola c. mergiserrati (De Geer), terminal segments of male abdomen,
ventral. 26. Female vulva. 27. A. crassicornis (Scopoli), female meso-metasternal
plate. 28. A. c. mergiserrati, female meso-metasternal plate.

We accordingly designate as male neotype of Trinoton lari (De Geer) a specimen
in the Meinertzhagen Collection (British Museum (Nat. Hist.)) (slide no. 8018) from
Anas. c. crecca (Linn.) from Nepal, which agrees with Text-figs. 26-28 and PL 2,
fig. 2 (in Clay & Hopkins, 1950) of Trinoton querquedulae ( (Linn.).

Ricinus mergi serrati (p. 78, PL 4, fig. 13)

The description and figure are unquestionably those of an Anaticola and the host
is carefully given as " le Plongeon nomme Mergus serratus Linn. Faun. Ed. 2 no.
136 " (an important point, because " plongeon " normally means a member of the
Gaviidae, or divers).

THE EARLY LITERATURE ON MALLOPHAGA 241

The species was renamed Pediculus mergi by J. C. Fabricius (1781 : 480) and Philop-
terus (Lipeurus] temporalis by Nitzsch (1818 : 292) ; both names are mere nomina
nova for De Geer's species and our neotypes of mergiserrati must also be neotypes of
mergi and temporalis. De Geer's figure seems to represent a nymph but is not so
good as most of his figures. Ricinus mergiserrati De Geer passed under Nitzsch's
name until 1916, when Harrison (p. 138) mentioned it as Esthiopterum mergiserrati
De Geer and correctly placed mergi Fabricius and temporalis Nitzsch as synonyms.

De Geer carefully separates, but does not name, another "Ricin" from the mesa
host, drawn as fig. 14 of his plate 4. This is undoubtedly an Anatoecus, and will be
dealt with under Philopterus icterodes Nitzsch, 1818.

All the specimens of Anaticola from the Anseriformes examined are very similar,
having male genitalia differing only in proportions and the female genital region in
the chaetotaxy ; all have the bilobed spermatheca in the female. Distinguishing
characters are found in the proportions of the head, meso-metasternal plate and male
genitalia, and in the chaetotaxy of the prothorax and male and female genital
regions. A. mergiserrati (which should probably be considered as a subspecies of
A. anseris (Linn.) or certainly of A. crassicornis (Scopoli)) is distinguished from anseris
by the proportions of the head and male genitalia, by the characters of the pre-
antennal region of the head, and by the presence of only one dorsal submarginal
prothoracic seta each side. Both crassicornis and mergiserrati tend to have a greater
number of setae on the genital region of the male (Text-fig. 25) and on the vulva
of the female (Text-fig. 26) than anseris too few specimens of this latter species
are available for a more exact comparison. A. mergiserrati is distinguished from
crassicornis with certainty only by the broader marginal carinae of the head (Text-
figs. 23-24) and the longer penis. The meso-metasternal plate (Text-figs. 27, 28) is
shorter and broader in mergiserrati.

MEASUREMENTS IN MM. :

Male Female

Length Breadth Length Breadth

Head . . . 0-67 0-43 . 0-72 0-47

Prothorax . . 0-32 . 0-35

Pterothorax . 0-41 0-47

Abdomen . . 1-50 0-49 . 2-00 0-57

Total . . .2-70 . 3-20

Genitalia . .0-65

LENGTH OF PENIS IN MM. (number of specimens in brackets): 0-20 (i), 0-22 (6),

23 (3).

Neotype male (Text-figs. 24, 25 ; PI. n, fig. 2) and neallotype female (Text-figs. 26,
28 ; PI. u, fig. 3) of Anaticola crassicornis mergiserrati (De Geer) in the Meinertzhagen
collection (British Museum (Nat. Hist.)) (slide no. I92g8a) from Mergus senator Linn,
from North Uist, Scotland. Neoparatypes : 46 males and 28 females from the same
host species from the British Isles.

242

THE EARLY LITERATURE ON MALLOPHAGA

Ricinus gallinae (p. 79, PL 4, fig. 15)

The host is " Poule " (= Callus domesticus) and the description and figure repre-
sent a Goniocotes. The name is not preoccupied by Pediculus gallinae Linn., which
is a Menopon.

Nitzsch (1818 : 294) renamed the species Philopterus (Goniodes) hologaster. Bur-
meister (1828 : 431) placed it in his new subgenus Goniocotes, of which it was sub-
sequently selected as type species.

30

29

FIGS. 29, 30. Goniocotes gallinae (De Geer). 29. Male. 30. Male genitalia.

MEASUREMENTS IN MM.

Head .

Prothorax

Pterothorax

Abdomen

Total .

Genitalia

Male

Female

Length

Breadth

0-28

o-37

0-22

o-34

0-50

0-51

0-90

0-42

1

Length

Breadth

0-40

0-51

0-90

0-31
0-46
0-8o

1-44

Neotype male (Text-figs. 29, 30) and neallotype female (Text-figs. 31, 32) of Gonio-
cotes gallinae (De Geer) in the British Museum (Nat. Hist.) Collection (slides nos.
567-8) from Gattus domesticus from Scotland. Neoparatypes : 4 males and 25 females
from the same host species from the British Isles,

THE EARLY LITERATURE ON MALLOPHAGA

243

Ricinus cams (p. 81, PI. 4, fig. 16)

The host is " chien " (= Canis familiaris) , and the description and not very good
figure unquestionably represent the species of Trichodectes commonly found on this
host.

O. Fabricius (1780 : 215) gives a completely independent description of a Pediculus
canis, from Canis familiaris, which has given much trouble because the description

31

FIGS. 31. Gomocotes gallinae (De Geer), female.

is so bad. Nitzsch (1818 : 296) gave a doubtful reference to canis Fabricius under
his Trichodectes subrostratus (a nomen nudum in 1818), Harrison ignored it, and other
authors who mention it mostly accept Nitzsch's suggestion that it was subrostratus.
It must receive serious consideration because, as it is not a primary homonym of
Ricinus canis De Geer, it could be the earliest valid name for Felicola subrostratus
(Burmeister).

244

THE EARLY LITERATURE ON MALLOPHAGA

Turning to the original description of Fabricius, we find that numerous structural
characters exclude any dog-parasites except Mallophaga and it is stated to occur in
large numbers, excluding any likelihood that it was not a true dog-parasite. We
fully agree with Keler (1938 : 422) that with one exception (the mention of a " ros-
trum subcylindricum ") the characters agree excellently with the Trichodectes of the
dog, this former character being certainly what led Nitzsch to suggest that it might
be subrostratus (found on the domestic cat). But we again find ourselves in full
agreement with Keler that the other characters exclude the cat-louse, and that the
" rostrum subcylindricum " must have been a portion of a hair gripped between the
mandibles. Accordingly we shall select a neotype for Trichodectes canis (O. Fabri-
cius) in such a way as to make it a synonym of T. canis (De Geer), of which it is also
a secondary homonym. Trichodectes lotus Nitzsch (1818 : 296) is a nomen novum for
De Geer's species and therefore an absolute synonym, based on the same types.

32

FIG. 32. Goniocotes gallinae (De Geer), terminal segments ot female abdomen, ventral.

Neotype male and neallotype female of Trichodectes canis (De Geer), which agree
with the description and figures published by Werneck under the same name (Wer-
neck, 1936 : 502-6, figs. 130-135), in the British Museum (Nat. Hist.) (slide nos.
581-2) from Canis familiaris from Sao Paulo, Brazil. Neoparatypes : 32 males and
31 females from the same host species from Brazil, British Isles and Germany.

O. FABRICIUS, 1780

(Fauna Groenlandica, Hafniae & Lipsiae, pp. 215-220)

Most of the descriptions in this work are new and independent, but there are a few
redescriptions. The descriptions are poor and there are no figures, but the names
are in unquestionably valid form.

There is some confusion regarding the correct interpretation of the hosts from
which Fabricius collected his specimens, and we are greatly indebted to Dr. Finn
Salomonsen for assistance in identifying these correctly.

Pediculus canis (p. 215)

This species has been fully discussed under Ricinus canis De Geer, of which it is a

synonym.

THE EARLY LITERATURE ON MALLOPHAGA

243

Neotype male of Trichodectes canis (Fabricius) in the British Museum (Nat. Hist.)
(slide no. 583) from Canis familiaris from Sao Paulo, Brazil. This specimen agrees
with the neotype of Trichodectes canis (De Geer).

33

FIG. 33. Saemundssonia grylle (O. Fabricius), male genitalia. a. Head of paramere.

Pediculus strigis (p. 216)

Fabricius thought his species, taken from Strix nyctea (= Nyctea scandiaca), was
the same as P. strigis Miiller, which we have already discussed as P. strigis Pontop-
pidan. It is unlikely that this is the case, but as the name is a primary homonym
we do not propose to discuss it further.

246

THE EARLY LITERATURE ON MALLOPHAGA

Pediculus corvi (p. 217)

This name is attributed to Syst.Nat. (i.e., Linne), and there is no need to discuss
it except to note that as the description is that of a Philopterus and the host is
Corvus comx it could be regarded as a restriction of Linne's name to the species found
on the raven. We have already restricted Philopterus corvi (Linn.) in the same sense.

34

36

FIG. 34. Saemundssonia grylle (O. Fabricius), terminal segments of female abdomen.
FIGS. 35, 36. Saemundssonia lari (O. Fabricius). 35. Anterior region of male head, dorsal.

36. Genital region of female.

Pediculus clangulae (p. 217)

Fabricius gave the host of this species as Anas clangula, but (as confirmed by Dr.
Salomonsen) the species which Fabricius had must have been Bucephala islandica
(Gmelin). In addition to the description there is a queried reference to Pediculus
querquedulae Linn. It was doubtless this reference which led Harrison (1916 : 12)

THE EARLY LITERATURE ON MALLOPHAGA 247

to refer clangulae to Trinoton, but the description cannot possibly apply to Trinoton
or to any other duck-parasite except an Anaticola, which it fits very well. The
species appears never to have been renamed or redescribed.

As we have no specimens of Anaticola from Bucephala islandica (Gmelin) we are
unable to erect neotypes for Anaticola clangulae (O. Fabricius).

Pediculus grylle (p. 218)

Although this purports to be a renaming of P. colymbi grylle Miiller (a tick), the
brief independent description " Pediculus niger, fasciis albis, abdomine ovato " can-
not apply to a tick, none of which are banded, but does apply fairly well to the
Saemundssonia, which occurs on Cepphus grylle, and not at all to any other known
parasite of this bird.

Denny (1842 : 44, 86, PI. 5, fig. 6) redescribed the species as Docophorus mega-
cephalus from a teneral specimen obtained from the same host, 1 and the species was
known under this name until Harrison (1916 : 12, 14) restored the name given to it
by Fabricius.

Specimens examined : 28 males and 38 females from Cepphus g. grylle (Linn.) from
the British Isles. As we have no material of Saemundssonia from Cepphus g. arctica
Brehm.), the subspecies from which Fabricius took his specimens, we are unable to
erect neotypes of Saemundssonia grylle (O. Fabricius). We have, however, included
figures (Text-figs. 33, 34 ; PI. n, fig. 4) of what is almost certainly this form drawn
from specimens taken from Cepphus g. grylle (Linn.). It should be noted that there
is individual variation in the shape of the female genital plate and chaetotaxy of
the genital region.

MEASUREMENTS IN MM. :

Male Female

Length Breadth Length Breadth

Head . . . 0-52 0-45 . 0-55 0-52

Prothorax . . 0-28 0-32

Pterothorax . . 0-36 0-40

Abdomen . .0-62 0-59 . 0-67 0-65

Total . . .1-28 . 1-42

Genitalia . .0-46

Neotype of Docophorus megacephalus Denny, a male (Text-figs. 33, 34 ; PI. n,
fig. 4), in the Meinertzhagen Collection (British Museum (Nat. Hist.)), (slide no.
16698), from Cephus g. grylle (Linn.) from N. Uist, Outer Hebrides.

Pediculus bassani (p. 218)

Thompson (1940 : 372-381) has dealt fully with the identity and synonymy of
Pectinopygus bassani (O. Fabricius) and has erected neotypes for it (p. 380).

Denny's type material of this species is not in the Brit. Mus. (N.H.) collection.

248

THE EARLY LITERATURE ON MALLOPHAGA

Pediculus lari (p. 219)

Harrison (1916 : 16) considered this name to be preoccupied by lari De Geer, but
the latter was published in Ricinus and is now in Trinoton, so lari O. Fabricius, which
is a Saemundssonia, is valid. The description is much better than the rest and
mentions a triangular head, acuminate in front and with a bifid rostrum, that the
insect is wholly black except for white blotches above and below, besides other

38

FIGS. 37-39. Saemundssonia lari (O. Fabricius). 37. Male genitalia. 38. Head of paramere
enlarged. 39. Mesosome enlarged (from another specimen).

details which make it perfectly clear that Fabricius had before him a female Saemunds-
sonia. The original host-record is " Habitat in laro glauco et quidem macro " ;
Larus glaucus is a synonym of L. hyperboreus Gunnerus. The species was re-
described as 5. lariphaga by Timmermann (1949 : 8). Docophorus lari Denny, 1842 ;
D. lari Grube, 1851 ; D. lari T. Miiller, 1927 and Pediculus lari Larranaga, 1928, are
all referable to Saemundssonia and are, therefore, homonyms of 5. lari (Fabricius).
5. lari (O. Fabricius) is the earliest name for any species of Saemundssonia from

THE EARLY LITERATURE ON MALLOPHAGA 249

the genus Lams, and, as shown by Timmermann (1951), the Saemundssonia from
nearly all the species of this genus must be considered as subspecies of lari.

In 5. /. lari the tergal plates of abdominal segment II are joined medianly in both
sexes and the dorsal abdominal setae of the female do not form a continuous line
across the segment, occurring only along the posterior margins of the tergal plates.
In the male genitalia there is no sclerotized cross-bar at the distal end of the basal
plate (Text-fig. 37) and the endomeral projections are fused medianly.

MEASUREMENTS IN MM. :

Male Female

Length Breadth Length Breadth

Head . . .0-60 0-62 . 0-62 0-68

Prothorax . . 0-34 0-37

Pterothorax . . 0-48 0-53

Abdomen . . 0-90 0-85 . 1-16 0-98

Total . . .1-76 . 2-06

Genitalia . .0-68

Neotype male (Text-figs. 35, 37-39, PI. n, fig. 5) and neallotype female (Text-fig.
36) of Saemundssonia lari lari (O. Fabricius) in the British Museum (Nat. Hist.)
Collection (slide no. 569) from Larus hyperboreus Gunnerus from Iceland. Neopara-
types : 51 males and 48 females from the same host species from Iceland, Jan Mayen
Is. and the British Isles.

Pediculus tringae (p. 219)

This species is stated to occur on Tringa striata, which at once presents us with a
difficulty, because this name has been applied to the redshank and also (incorrectly)
to the purple sandpiper ; but the former does not occur in Greenland, so the host
of P. tringae must have been Erolia maritima. The insect is described as white with
a red dot on the abdomen (obviously ingested blood) and a suborbicular body, the
size is that of the head of a pin, the antennae are like very minute hairs, the thorax
is oval and the abdomen almost circular. Obviously the specimen was a very young
nymph. Harrison regards the species as unrecognizable and we have much sym-
pathy with this view, but it is one only to be adopted in the last resort. The mention
of an almost orbicular abdomen at once rules out any genera found regularly on the
Charadriiformes except Saemundssonia, Austromenopon and perhaps Actornitho-
philus, but not only are nymphs of the two latter genera brownish-yellow rather than
white, but also their antennae are concealed and the palps (sometimes mistaken for
antennae) far from conspicuous, so that Fabricius' insect must have been a Saemunds-
sonia ; only one Saemundssonia appears to occur normally on Erolia maritima.

Through the kindness of Dr. Rene Malaise it has been possible to remount and
examine the syntypes of Docophorus arcticus Mjoberg (1910 : 122) said to have been
taken from " Tringa sp. (maritima ?)." The syntypes, which comprise one male and
two females, are conspecific with the neotypes of S. tringa from Erolia maritima ; S.
arctica can, therefore, be considered as a synonym of tringae (Fabricius).

250

THE EARLY LITERATURE ON MALLOPHAGA

This species is distinguished from other Saemundssonia on the Charadrii by the
size and shape of the head (PI. n, fig. 6) and anterior plate (clypeal signature) (Text-
fig. 40) and by the male genitalia (Text-figs. 42-44). The tergal plates of abdominal
segment II are joined medianly in both sexes and the dorsal abdominal setae do not
form a continuous line across each segment, occurring only along the posterior
margins of the tergal plates. In the male genitalia there is a sclerotized cross-bar
at the distal end of the basal plate and the endomeral projections are fused medianly
(Text-fig. 42).

40

FIGS. 40, 41. Saemundssonia tringae (O. Fabricius). 40. Anterior region of male head, dorsal.
41. Terminal segments oi female abdomen.

MEASUREMENTS IN MM.

Head .

Prothorax

Pterothorax

Abdomen

Total .

Genitalia

Male

Length Breadth

0'55 0-5I

0-28

0-38

0-75 0-65

1-47

Female

Length
0-62

0-97

I-6 7

Breadth

o-59
o-33
0-47
0-83

Neotype male (Text-figs. 40, 42-44, PI. n, fig. 6) and neallotype female (Text-fig.
41) of Saemundssonia tringae (O. Fabricius) in the Meinertzhagen Collection (British
Museum (Nat. Hist.)), slide no. 2346, from Erolia maritima (Briinnich) from Scotland.
Neoparatypes : 16 males and 16 females from the same host species from the British
Isles, Spitzbergen and Finland.

THE EARLY LITERATURE ON MALLOPHAGA

251

Lectotype of Saemundssonia arctica (Mjoberg) : $ in the Naturhistoriska Riks-
museum, Stockholm, from "Tringa sp. (maritimaP) from Lilla Pendulum- On "
(6.vii.i899).

Pediculus hiaticulae (p. 220)

The host is Charadrius hiaticula, and the only points of much value in the descrip-
tion are that the head is triangular, the oblong body sublinear and the whole insect
mouse-coloured, but this description is sufficient to give us the genus at once, for the

42

44

FIGS. 42-44. Saemundssonia tringae (O. Fabricius). 42. Male genitalia. 43. Head of paramere.
44. Mesosome enlarged (from another specimen).

only genus occurring on the genus Charadrius in which the body could possibly be
described as sublinear is Quadraceps. Two species of this genus occur on Charadrius
hiaticula, and both are fairly common ; Quadraceps fissus (Burmeister) is a relatively
stout species, while the other is very slender. This latter species was discussed at
some length by Waterston (1915 : 35) without his being able to find a satisfactory
name for it, but Hopkins (1942 : 115) expressed the opinion that the slender species
is Quadraceps hiaticulae (O. Fabricius). As Fabricius describes his species as sub-
linear we think there can be no question that this is the species he had before him,

252

THE EARLY LITERATURE ON MALLOPHAGA

and we reject Harrison's placing oifissus Burmeister as a synonym (Harrison : 1916 :
15, 113). Harrison ascribes the authorship to Miiller, but the latter only mentioned
Pediculus hiaticulae as a nomen nudum, so the author is Fabricius.

MEASUREMENTS IN MM. :

Male

Female

Head .

Prothorax

Pterothorax

Abdomen

Total .

Genitalia

Length
0-42

1-16
1-83
0-36

Breadth
0-23
0'17
0-25
0-3I

Length
0-42

1-26
1-90

Breadth
0-23

0-18

0-24
0-30

45

FIGS. 45, 46. Quadraceps hiaticulae (O. Fabricius), male. 45. Anterior region of head, dorsal.

46. Terminal segments of abdomen.

Neotype male (Text-figs. 45-50 ; PI. 12, fig. i) and neallotype female (Text-fig. 51,
PI. 12 fig. 2) of Quadraceps hiaticulae (O. Fabricius) in the Meinertzhagen Collection
(British Museum (Nat. Hist.)), slide no. 10514, from Charadrius h. hiaticula Linn,
from Ireland. Neoparatypes : 116 males and 159 females from the same host form
from the British Isles and 17 males and 9 females from Charadrius h. tundrae (Lowe)
from the Sudan.

THE EARLY LITERATURE ON MALLOPHAGA

253

Pediculus lagopi (p. 220)

This appears to be a redescription of Linne's species of the same name and the
host (Tetrao lagopus) is also the same.

(Species Insectorum

J. C. FABRICIUS, 1781

. . Hamburgi & Kilonii, Vol. 2, pp. 478-484)

This is little more than a list of names, and all the new names proposed in it have
already been discussed. It need not detain us.

48

FIGS. 47, 48. Quadraceps hiaticulae (O. Fabricius). 47 Male genitalia. 48. Paramere.

SCHRANK, 1781

(Enumemtio insectomm Austriae indigenorum.

pp. 499-507, pi. i.)

Augustae Vindelicorum,

We must confess to a great feeling of relief in getting away from the work of J. C.
and O. Fabricius to that of Schrank, for (as before) his descriptions are for the most
part accurate and are nearly all accompanied by figures which enable his species to be
easily recognized.

ENTOM. HI, 6.

254

THE EARLY LITERATURE ON MALLOPHAGA

Pediculus porcelli (p. 500, PI. i, fig. i)

After being mentioned by Linne without indication, definition or description, this
name was copied by almost every author with whom we have dealt, still as a nomen
nuduni, until Schrank finally described the species. The host is Mus Porcellus, the
description is good and the figure excellent, considering the date. Schrank himself
(1803 : 186) renamed the species Pediculus saviae, Olfers (1816 : 83) redescribed it as
Pediculus bifurcatus, and Nitzsch (1818 : 304) proposed Gyropus gracilis as a nomen
novum for it. Later authors have either used Nitzsch's name or referred the name
porcelli to Linne, but the authorship of this name must be ascribed to Schrank.

50

49

51

FIGS. 49, 50. Quadraceps hiaticulae (O. Fabricius). 49. Mesosome, ventral. 50. Distal end of

mesosome, dorsal.
FIG. 51. Quadraceps hiaticulae (O. Fabricius), terminal segments of the female abdomen.

The species has been excellently described and figured by Werneck (1936 : 398-
402, figs. 1-9), and our neotypes agree with his description and figures.

Neotype male and neallotype female of Gliricola porcelli (Schrank) in the British
Museum (Nat. Hist.) Collection (slides nos. 584-5) from tame guinea-pig, Cavia por-
cellus (Linn.), from Rio de Janeiro, Brazil. Neoparatypes : 14 males and 30 females
from the same host species from Brazil, British Isles and S. Africa.

Pediculus ovis (p. 502, PI. i, figs. 8, 9)

This is a somewhat similar case to that of porcelli, since Linne mentioned the name
without description and with only a queried reference to Redi, which does not save

THE EARLY LITERATURE ON MALLOPHAGA 255

the name from being a nomen nudum. In passing it seems worth mentioning that
the presence of the query suggests very strongly that what Linne had before him
was not Redi's insect, for the latter's figure is not at all bad ; we suspect that Linne
may have had a sucking louse. In any case the authorship of ovis must be assigned
to Schrank.

The host is Ovis aries (domestic sheep) and the species is well known. Schrank
(1803 : 187) renamed it Pediculus ovisarietis, Olfers (1816 : 85) described it as Pedi-
culus sphaerocephalus, and Nitzsch (1818 : 296) proposed the new name Trichodedes
sphaerocephalus for " Fed. Ovis Linn. Fabric." (nomina nuda], Schrank's species and
the left-hand figure on Redi's pi. 22. Our neotypes of ovis Schrank are also
automatically neotypes of the nomina nova proposed by Schrank (1803) and Nitzsch
(1818), but are not neotypes of sphaerocephalus Olfers because this has an indepen-
dent description.

The species has been excellently described and figured by Werneck (1936 : 537-
540, figs. 169-174), and the neotypes agree with his figures and description.

Neotype male and neallotype female of Damalinia ovis (Schrank) in the British
Museum (Nat. Hist.) Collection (slides nos. 586-7) from domestic sheep, Ovis aries
Linn, from Rio de Janeiro, Brazil. Neoparatypes : 29 males and 37 females from
the same host species from Brazil, E. Africa and New South Wales, Australia.

Pediculus anatis (p. 503, PL i, figs. 2, 3)

The excellent description and the figures represent an Anaticola, and there is a
reference to the upper figure of Redi's plate 10, which is Anaticola anseris (Linn.).
The host is given as Anas boschas varietas fera (= Anas p. platyrhynchos) , but fig. 2
of the plate is labelled P. anseris, which perhaps means that this specimen was taken
from a goose. There can be little doubt that anatis Schrank is a mixture of Anaticola
anseris (Linn.) and A. crassicornis (Scopoli), and it is probably the basis for Olfers'
record of both Anas and Anser as hosts of the latter species and the subsequent
appearance in the literature of a non-existent " crassicornis Olfers." In view of
Schrank's host-record and choice of specific name, anatis must be held to apply
primarily to Anaticola crassicornis (Scopoli), of which it is a synonym.

Neotype male of Anaticola anatis (Schrank) in the Meinertzhagen Collection (British
Museum (Nat. Hist.)), slide no. 4242, from Anas p. platyrhynchos Linn, from N.E.
Poland. This specimen agrees with the description and figures (Clay & Hopkins,
1951 : 17) of A. crassicornis (Scopoli).

Pediculus ortygometrae (p. 503)

This species, from Rallus Crex = Crex crex (Linn.), is not figured and the descrip-
tion is most unsatisfactory. The species was identified by Burmeister (1838 : 428)
and by Denny with Nitzsch's manuscript name Nirmus attenuatus, and there are
important reasons why this identification should be accepted if at all possible. The
position is that attenuatus first appeared in print (Nitzsch, 1818 : 291) as a nomen
nudum, a reference to Pediculus ortygometrae Schrank being queried. Burmeister
gave no description of the insect but included the same reference without the query,

256 THE EARLY LITERATURE ON MALLOPHAGA

thus making Nirmus attenuatus Burmeister a nomen novum for P. ortygometme
Schrank. Denny (1842 : 52, 134, PL 10, fig. 2) described material from Crex pratensis
(= Crex crex (Linn.)) as Nirmus attenuatus " Nitzsch," and " N. attenuatus Nitzsch "
as subsequently described by Giebel from Nitzsch's manuscript (1874 : 170, PL 6,
fig. i) appears to be congeneric with Denny's species. To interpret P. ortygometrae
otherwise than was done by Burmeister and by Denny would raise serious difficulties
with regard to the type species of Rallicola.

Schrank's description of ortygometrae is as follows : "Insectum oblongum. Caput
cordatum, parte acuminata ab abdomine aversa ; antennae breves moniliformes.
Thorax angustus, longus apice latior, apicis medio productiore. Abdominis seg-
menta, demta thorace octo setosa, utrinque spiraculis magnis maculatis instructa.
Pedes thoraci affici, articulis tribus elliptoidibus constant, articulo tertio acuto,
breviori. In capite varia transparent intestina ; ejusque apices utrinque setam
longiusculam emittunt."

We considered the possibility that the various internal structures appearing in the
head might be the conspicuous gular plate of Pseudomenopon, but this genus is ex-
cluded by the cordate head and the fact that the " apex " of the thorax is stated to
be produced in the middle. We think Fulicoffula to be excluded by the statement
that the insect is oblong and by the cordate head. Incidifrons is not known from
Crex crex, but we have examined /. fulicae (Linn.) and this, in common with Pseudo-
menopon, has two long setae arising from the temporal angles (Schrank's " apices "
of the head), whereas Rallicola has only one. Summing up, the whole of the very
vague description could apply to Rallicola and the presence of only one long temporal
seta is peculiar to this genus. It is legitimate to use as confirmation the fact that
Rallicola is common on Crex crex and the other genera rare or unknown.

This species is distinguished from R. cuspidatus (Scopoli) (see Clay & Hopkins,
1951 : 25) by the shape of the head in both sexes, the absence of marked sexual
dimorphism of the antennae, the male genitalia and the female genital region.

MEASUREMENTS IN MM. :

Male Female

Length Breadth Length Breadth

Head . . 0-44 0-33 0-45 0-33

Prothorax . . 0-22 0-22

Pterothorax . . 0-29 . 0-28

Abdomen . .0-86 0-42 . 0-92 0-42

Total . . . 1-53 . 1-66

Genitalia . .0-28

Neotype male (Text-figs. 52, 53) and neallotype female (Text-figs. 54, 55) of Ralli-
cola ortygometrae (Schrank) in the British Museum (Nat. Hist.) Collection (slide nos.
570-1) from Crex crex (Linn.) from Scotland. Neoparatypes : 5 males and 3 females
from the same host species from Scotland and France.

The material described by Denny as Nirmus attenuatus " Nitzsch " (one male in
the British Museum (Nat. Hist.) Denny Collection) agrees with the neotype of Ralli-

THE EARLY LITERATURE ON MALLOPHAGA

257

cola ortygometrae (Schrank). There are no specimens of " Oncophorus attenuatus
N.," as described by Piaget (1880 : 214, PI. 18, fig. i) in the Piaget Collection in the
British Museum (Nat. Hist.).

FOURCROY, 1785

(Entomologia Parisiensis, Paris. Part 2, pp. 517-522)

The section of this work which deals with Anoplura and Mallophaga is in two diffe-
rent parts ; pages 517-519 contain a list of species of Pedicidus, sometimes with

52

FIGS. 52, 53. Rallicola ortygometrae (Schrank). 52. Male. 53. Male genitalia.

brief descriptions or quotations of the descriptions of other authors (whose names are
not mentioned), and all the names are in valid binomial form. But on pages 520-
522, under a separate heading " pediculi Rhedi & Linnaei " there is a list in which
every name which is not a nomen nudum is a descriptive phrase and has a partial

258 THE EARLY LITERATURE ON MALLOPHAGA

duplicate: "Pediculus accipitris, abdomine oblongo" and "Pediculus accipitris, abdo-
mine ovato," for instance. We conclude that these phrases on pp. 520-522 are not
to be taken as names and are only in Latin because the whole work is in that language.
On the other hand, the names on pp. 517-519 are undoubtedly valid and must be
dealt with, because in three instances Geoffrey's descriptive phrases are first shortened
to valid form in this work.

54

FIGS. 54, 55. Rallicola ortygometrae (Schrank). 54. Female.

abdomen.

55. Terminal segments of female

Pediculus circi (p. 518)

Proposed for Geoffrey's Pediculus circi, fuscus oblongus ... , the identity of
which has already been briefly discussed (Clay & Hopkins, 1950 : 270). Geoffrey's
host-record is " Busard des marais, circus Bellon " (= Circus aeruginosus (Linn.)).

The same species was named Pediculus milvi by Schrank (1803 : 193) ; Liotheum
(Laemobothrion) giganteum Nitzsch (1818 : 301) is composite and we shall restrict

THE EARLY LITERATURE ON MALLOPHAGA

259

the name to the present species. L. nigrolimbatum Giebel was described from speci-
mens taken from Circus cineraceus (= C. pyargus (Linn.)) and Circus aeruginosus,
and in view of the loss of the types is best assumed to be the same as circi (Fourcroy) .
The species of Laemobothrion from the Falconiformes can be divided into three
main groups as follows (see also Keler, 1937, and Eichler, 1943) :

1. Gular setae and lateral setae on prosternal plate absent . . tinnunculi group.
-. Gular setae and lateral setae on prosternal plate present . . . 2.

2. Majority of abdominal tergites with central uncoloured area (Text-fig. 68)

(vulturis group) 3.
-. Abdominal tergites without such areas .... maximum group.

3. Lateral margins of anterior abdominal segments with dark, stout spine-

like setae ......... glutinans N.

-. Lateral margins of anterior abdominal segments without such setae

rest of vulturis group.

56

58

59

61

FIGS. 56-58. Posterior setae of ist femur. 56. Laemobothrion tinnunculi (Linn.)

57. L. circi (Fourcroy). 58. L. vulturis (J. C. Fabricius).
FIGS. 59-61. Proximal part of and femur. 59. L. tinnunculi. 60. L. circi. 61. L. vulturis.

In addition the members of the tinnunculi group have the shape of the head dis-
tinctive, have only a few setae on the anterior margin of the prothorax, and never
have a complete vertical line of setae along the outer edges of the patches of minute
comb-like structures on abdominal sternites IV-V. 1 The vulturis group is further
characterized by the presence in the female of a lateral pigmented spot each side
posterior to the vulva (Text-fig. 67, x), and by the presence in both sexes of dark,
stout spine-like setae on the margin of the first femur and on the margin and dorsal

1 These are similar to those found in Microtenia,

a6o

THE EARLY LITERATURE ON MALLOPHAGA

surface of the proximal part of the second femur (Text-figs. 58, 61) ; the third femur
may or may not have these setae. In general, members of the vulturis group tend
to be larger and to have a greater number of stouter setae. In the maximum group
some forms have (Text-figs. 57, 60) a few of these spine-like setae on the femora,
smaller than those of the vulturis group ; in the tinnunculi group the second femur

FIG. 62. Laemobothrion circi (Fourcroy), male head, dorsal.

64

63

FIGS. 63, 64. Laemobothrion circi (Fourcroy). 63. Female prosternal and meso-metas-
ternal plates ( unmounted specimen). 64. Proximal part of prosternal plates of three
individuals to show variation in shape and number of setae (mounted Canada balsam).

(only) may have one or two small spine-like setae (Text-figs. 56, 59). Each of these
groups will probably prove to be a polytypic species, the subspecies of which seem
to be distinguished by the colour pattern (apparently correlated to a certain extent
with the plumage colour of the host), the shape of the gular plate, the number and

THE EARLY LITERATURE ON MALLOPHAGA

261

size of the setae, and the total length in the females. There is some individual varia-
tion in the numbers of gular and lateral setae of the prosternal plate which makes
these characters unreliable for the separation of related forms ; similarly the shape
of the prosternal plate tends to vary (Text-figs. 63, 64). The position of the setae
on the meso-metasternal plate (see Eichler, 1943 : 210) is also variable, due to varia-
tion in the posterior prolongation of this plate. The shape and size of the head, even
where the total length is a distinguishing character, tend to be similar in related
forms. It seems doubtful whether the male genitalia show any differences through-
out the members of one group, but a greater amount of material is necessary in order
to judge the amount of variation in the shape of the sclerites within populations from
one host species.

65

FIG. 65. Laemobothrion circi (Fourcroy) male, terminal segments of abdomen.

a. Anus (internal).

Of the four species described up to this date 1785 L. tinnunculi (Linn.) is quite
distinctive (see Clay & Hopkins, 1950 : 228). L. vulturis (J. C. Fabricius) was re-
described (Clay & Hopkins, 1951 : 33) from a small amount of mounted material,
but now through the kindness of Mr. B. K. Tandan (University of Lucknow) it has
been possible to examine fresh material of this species and thus to amplify the descrip-
tions and figures. It differs from other forms parasitizing the Aegypiinae mainly
in size and possibly in colour pattern, but the available specimens of Laemobothrion

262

THE EARLY LITERATURE ON MALLOPHAGA

from this subfamily are not in a condition to show this last character fully. The
number and size of the setae tend to be greater in both sexes than in circi (cf. Text-
figs. 66 and 67), but there is considerable individual variation in this character. In
our first description (1951 : 34) it was suggested that the number of lateral prosternal
setae was a diagnostic character, but the number of these in L. vulturis may vary, as
shown for the following 4 males and 4 females (setae on left and right side of each
prosternal plate) :

Male .
Female

4 + 5; 4
3 + 4; 4

3; 5 + 3; 7 + 6
6; 6 + 7; 7 + 6

66

FIG. 66. Laemobothrion circi (Fourcroy) female, terminal segments of abdomen.

This species has the form of the tergal plates (Text-figs. 68) and femoral setae
(Text-figs. 58, 61) characteristic of the vulturis group. L. maximum (Scopoli) (see
Clay & Hopkins, 1951 : 34) and L. circi (Fourcroy) belong to the maximum group,
but owing to lack of material of maximum the differences (if any exist) between these
two forms cannot be given. L. circi differs from L. titan Piaget (host: Milvusm.
migrans) in the lesser degree of pigmentation and in the female by the shorter abdo-
men. In both sexes of circi abdominal tergites I-IV show signs of a median division.

THE EARLY LITERATURE ON MALLOPHAGA
MEASUREMENTS IN MM. :

263

Male

Head (midline

(laterally)
Prothorax
Pterothorax .
Abdomen
Total .
Genitalia

Length

1-44
1-70

4'95
8-15

2-20

Breadth

i-54

1-20
2 -2O

Female

Length Breadth
1-52
1-78 1-65

6-00
9-40

1-87

2-80

67

FIG. 67. Laemobothrion vulturis (J. C. Fabricius), Terminal segments of female abdomen,

ventral, x. Post-vulval pigmented area.
FIG. 68. Laemobothrion vulturis (J. C. Fabricius), third abdominal tergite, female.

Neotype male (Text-figs. 62, 65; PI. 12, figs. 3, 4) and neallotype female (Text-figs.
57, 60, 63, 64, 66; PI. 12, fig. 5) of Laemobothrion circi (Fourcroy) in the British
Museum (Nat. Hist.) Collection (slide no. 609) from Circus a. aeruginosus (Linn.)
from Kenya. Neoparatypes : 26 males and 20 females from the same host form
from Kenya, Yugoslavia and India.

Pediculus passeris (p. 519)

A binomial name for Geoff roy's descriptive phrase " Pediculus subflavescens :
abdomine ovato ..." (see Clay & Hopkins, 1950 : 270), but with a brief descrip-
tion which may perhaps be independent. The point is not of much importance, as

264 THE EARLY LITERATURE ON MALLOPHAGA

both descriptions clearly indicate a Philopterus and the host is the same : "moineau
franc " (= Passer domesticus).

The earliest name for the Philopterus from Passer domesticus is P. fringillae
(Scopoli) (see Clay & Hopkins, 1951 : 32) and passeris, therefore, becomes a synonym
of this latter name.

Neotype male of Philopterus passeris (Fourcroy) in the Meinertzhagen Collection
(British Museum (Nat. Hist.)), slide no. 15778 from Passer d. domesticus (Linn.) from
Cornwall, England. This specimen agrees with the figures and description (Clay &
Hopkins, 1951 : 32 ; fig. 43 ; PI. 3, fig. 3) of P. fringillae (Scopoli).

Pediculus corvi (p. 519)

Proposed for Geoff roy's " Pediculus albo nigroque varius ..." The host is
"corbeau," which may mean crow or raven, though usually the latter. Geoffrey's
species is certainly a Philopterus.

Philopterus corvi (Fourcroy) is preoccupied both in Pediculus and in Philopterus,
for if not the same as Philopterus corvi (Linn.) it is a homonym of the latter. It is
therefore unnecessary to try to fix it more closely.

MOHR (N.), 1786

(Fors0g til en Islandsk Naturhistorie. Kiebenhavn)

Pediculus procellariae (p. 103)

There is a short Latin description but no host-record other than that provided by
the name, which is preoccupied by Pediculus procellariae J. C. Fabricius, 1775.

We consider Pediculus procellariae Mohr, 1786, nee J. C. Fabricius, to be completely
unrecognizable.

ACKNOWLEDGMENTS

We are indebted to the Trustees of the British Museum for permission to publish
Text-figs. 12, 33, 53, drawn by Mr. A. J. E. Terzi, to the Zoological Society of
London for permission to publish figures 52-55, and to Colonel Meinertzhagen for
permission to publish Text-figs. 29 and 31, drawn by Mr. R. S. Pitcher. Text-figs.
5 and 18 were drawn by Mr. A. Smith, and the remaining figures by Miss T. Clay.

TABLE I. Breadth (in mm.} of Head at Temples of Male Philopterus populations

with Number of Specimens.

0-45-0-46 0-47-0-48 0-49-0-50 0-51-0-52

citrinellae 3 22 2

curvirostrae . . i 16 4

Pyrrhulae . . 246

chloridis . 18 6 2

THE EARLY LITERATURE ON MALLOPHAGA

265

TABLE II. Breadth (in mm.} of Head of Female Philopterus populations with

Number of Specimens

0-47-0-48 0-49-0-50 0-51-0-52 0-53-0-54 0-55-0-56 0-57-0-58 0-59-0-60
citrinellae i 3 6 16 4

curvirostrae . . I II 12 4 I

pyrrhulae . 2 3 2 5 19 I

chloridis . 2 I 7 8 12

LIST OF SPECIES
The synonymy of the following names has been established :

Specific name
agonum Nitzsch .
alaudae Schrank
anatis Schrank
arcticus Mjoberg
attenuatus Burmeister .
bassani O. Fabricius .
bifurcatus Olfers
canis De Geer
canis O. Fabricius
chloridis Schrank
circi Fourcroy
citrinellae Schrank
clangulae O. Fabricius
communis Nitzsch
compar Piaget
cornicis De Geer
curuccae Schrank
curvirostrae Schrank .
emberizae De Geer
fringillae De Geer
gallinae De Geer
giganteum Nitzsch
globifer Olfers
gracilis Nitzsch .
grylle O. Fabricius
hiaticulae O. Fabricius
hologaster Nitzsch
lari De Geer
lari O. Fabricius
lariphaga Timmermann
latus Nitzsch
leontodon Nitzsch
megacephalus Denny .
mergi J. C. Fabricius .
mergiserrati De Geer .
mesoleucum Nitzsch
milvi Schrank

Present status Page.

Ricinus rubeculae (Schrank) .... 224
Menacanthus alaudae (Schrank) . . . 226
Anaticola crassicornis (Scopoli) . . . 225
Saemundssonia tringae (O. Fabricius) . . 249
Rallicola ortygometrae (Schrank) . . . 255
Pectinopygus bassani (O. Fabricius) . . 247
Gliricola porcelli (Schrank) .... 254
Trichodectes canis (De Geer) . . . .243
Trichodectes canis (De Geer) .... 244
Philopterus citrinellae (Schrank) . . 229, 232
Laemobothrion circi (Fourcroy) . . .258
Philopterus citrinellae (Schrank) . . 227, 232
Anaticola clangulae (O. Fabricius) . . . 246
Philopterus citrinellae (Schrank) . . . 228
Philopterus curvirostrae (Schrank) . . . 229
Myrsidea cornicis (De Geer) . . . .237
Menacanthus curuccae (Schrank) . . . 223
Philopterus curvirostrae (Schrank) . . 229, 232
Philopterus citrinellae (Schrank) . . . 237
Ricinus fringillae De Geer .... 235
Goniocotes gallinae (De Geer) . . . .242
Laemobothrion circi (Fourcroy) . . . 258
Philopterus citrinellae (Schrank) . . . 228
Gliricola porcelli (Schrank) .... 254
Saemundssonia grylle (O. Fabricius) . . 247
Quadraceps hiaticulae (O. Fabricius) . -251
Goniocotes gallinae (De Geer) . . .242

Trinoton querquedulae (Linn.) . . . 239
Saemundssonia lari (O. Fabricius) . . . 248
Saemundssonia lari (O. Fabricius) . . . 248
Trichodectes canis (De Geer) .... 244
Sturnidoecus sturni (Schrank) . . . 232

Saemundssonia grylle (O. Fabricius) . .247
Anaticola c. mergiserrati (De Geer) . . 241

Anaticola c. mergiserrati (De Geer) . . . 240
Myrsidea cornicis (De Geer) . . . .237
Laemobothrion circi (Fourcroy) . . .258

266

THE EARLY LITERATURE ON MALLOPHAGA

Specific name
minutum Nitzsch
nigrolimbatum Giebel .
nitidissimus Nitzsch .
ortygometrae Schrank .
ovis Schrank
ovisarietis Schrank
parviceps Piaget
passeris Fourcroy
phasiani J. C. Fabricius
porcelli Schrank .
procellariae Mohr
pterocephalus Olfers
pyrrhulae Schrank
pyrrhulae Piaget
rubeculae Schrank
saviae Schrank .
sphaerocephalus Olfers
sphaerocephalus Nitzsch
sturni Schrank .
subaequalis Haan
temporalis Nitzsch
tringae Fabricius

Present status Page

Menacanthus curuccae (Schrank) . . . 223
Laemobothrion circi (Fourcroy) . . . 259
Ricinus fringillae De Geer .... 236
Rallicola ortygometrae (Schrank) . . .255
Damalinia ovis (Schrank) .... 254
Damalinia ovis (Schrank) .... 255
Menacanthus alaudae (Schrank) . . . 226
Philopterus fringillae (Scopoli) . . . 263
Chelopistes phasiani (J. C. Fabricius) . . 234
Gliricola porcelli (Schrank) .... 254
Unrecognisable . ... 264

Ricinus rubeculae (Schrank) .... 224
Philopterus citrinellae (Schrank) . ,. 229, 232
Philopterus citrinellae (Schrank) . . .229
Ricinus rubeculae (Schrank) .... 223
Gliricola porcelli (Schrank) .... 254
Damalinia ovis (Schrank) .... 255
Damalinia ovis (Schrank) . . . .255
Sturnidoecus sturni (Schrank) . . . 232

Myrsidea cornicis (De Geer) .... 238
Anaticolac. mergiserrati (De Geer) . . .241
Saernundssonia tringae (O. Fabricius) . . 249

REFERENCES

CLAY, T., & HOPKINS, G. H. E. 1950. The early literature on Mallophaga : Pt. /. Bull. Brit.

Mus. (Nat. Hist.) Ent. 1 (3) : 223-272.

1951. Pt. //. op. cit. 2 (i) : 1-36

EICHLER, W. 1943. Balkan-Mallophagen. I : Laemobothrion. Mitt, naturw. Inst. Sofia, 16 :

207-213.
HOPKINS, G. H. E. 1942. Stray notes on Mallophaga: V. Ann. Mag. nat. Hist, (n) 9:

108-119.
KELER, S. 1937. Uber einige neue und interressantere Mallophagen des Deutschen Ento-

mologischen Instituts in Berlin-Dahlem. Arb. morph. tax. Ent. Berlin-Dahlem, 4 : 312-324.
THOMPSON, G. B. 1940. Notes on species of the Genus Pectinopygus. Ann. Mag. nat. Hist.

(n), 5 .-372-381.
TIMMERMANN, G. 1949. Beitrage zur Kenntnis der Ektoparasitenfauna Islandischer Sauge-

tiere und Vogel. I. Greinar Visind. Isl. 2 (3) : 1-32.

1951. Die Mowenkneifer. Par asitological News, Reykjavik, (2) 1 : 1-12.

WATERSTON, J. 1915. On some Mallophaga in the Kgl. Zoologisches Museum, Konigsberg.

Zool. Jb. (Abt. Syst.), 39 : 17-42.
WERNECK, F. L. 1936. Contribuicao ao conhecimento dos Mallophagos encontrados nos mam-

miferos sul-americanos. Mem. Inst. Osw. Cruz, 31 : 391-589.

PRESENTED

1 6 MAK 1954

EXPLANATION OF PLATES

PLATE 10

FIG. i. Ricinus rubeculae (Schrank), . x 28.

FIG. 2. Ricinus rubeculae (Schrank), $. x 28.

FIG. 3. Menacanthus alaudae (Schrank), $. x 55.

FIG. 4. Philopterus citrinellae citrinellae (Schrank). ^. x 62

FIG. 5. Sturnidoecus sturni (Schrank), <$. X 51.

FIG. 6. Ricinus fringillae De Geer, $. X 25.

PLATE ii

FIG. T. Myrsidea cornicis (De Geer), $. x 45.

FIG. 2. Anaticola mergiserrati (De Geer), <$. x 37.

FIG. 3. Anaticola mergiserrati (De Geer), $. x 29.

FIG. 4. Saemundssonia grylle (O. Fabricius), cJ. X 38.

FIG. 5. Saemundssonia lari (O. Fabricius), ^. x 38.

FIG. 6. Saemundssonia tringae (O. Fabricius), (J. x 66.

PLATE 12

FIG. i. Quadraceps hiaticulae (O. Fabricius), $. x 47.

FIG. 2. Quadraceps hiaticulae (O. Fabricius), $. x 47.

FIG. 3. Laemobothrion circi (Fourcroy), <$. x n.

FIG. 4. Laemobothrion circi (Fourcroy), $ genitalia. x 42.

FIG. 5. Laemobothrion circi (Fourcroy), $. X n.

Bull. B. M. (N.H.) Entom. Ill, 6.

PLATE ro

s~*

-

Bull. B. M. (N.H.) Entom. Ill, 6.

PLATE ii

Hull. n. M. (N.H.) Entom. Ill, 6.

PLATE 12

m.

x-*v- 1

d-\ i

r*

t

v-

rm

PRESENTED
1 6 MAR 1954

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

> 1 1 7 MAY 1954

SOME SAWFLIES
OF THE EUROPEAN ALPS

AND THE
MEDITERRANEAN REGION

(HYMENOPTERA : SYMPHYTA)

R. B. BENSON

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 3 No. 7

LONDON: 1954

SOME SAWFLIES

OF THE EUROPEAN ALPS
AND THE MEDITERRANEAN REGION

HYMENOPTERA : SYMPHYTA

BY

ROBERT B. BENSON

H

Pp. 267-296 ; 31 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 3 No. 7

LONDON : 1954

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

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

This paper is Vol. 3, No. 7 of the Entomological
series.

Issued April, 1954. Price Six Shillings.

SOME SAWFLIES OF THE EUROPEAN ALPS

AND THE MEDITERRANEAN REGION
I (HYMENOPTERA : SYMPHYTA)

By ROBERT B. BENSON

SYNOPSIS

The paper deals mainly with new or little-known sawflies (Hym. Symphyta) collected in
recent years in Cyprus and other parts of the Mediterranean Region with a few from the
European Alps. In all 16 new species and 5 new subspecies are described, and 4 are re-
instated from the synonymy.

A list is also included of the sawflies collected for the Museum in Cyprus by Mr. G. A.
Mavromoustakis.

THE following species or subspecies of sawflies are nearly all from the Central Euro-
pean Alps or the lands in or bordering the Mediterranean, and most of them are
described here for the first time. The specimens on which the new forms are being
based have come to me from many sources and have been accumulating in the
British Museum (Natural History) awaiting description over several years, although
not all of them belong to that institution. I should like here to thank all those
who have sent me the specimens and especially those who have waited patiently
a long time for results.

At the end of the paper I have included a list of all the sawflies collected in Cyprus
for the British Museum by Mr. G. A. Mavromoustakis. Three of these species
were also found by Dr. Hakan Lindberg during his stay in Cyprus in 1939 and I
am indebted to him for lending me his material for study. As virtually no records
of sawflies from Cyprus have been published before, except for some Cephidae by
Benson (1946), most of these records are new ; and the list probably includes all
the species that have yet been collected on the island.

I am using here, as I have before, the expression "(statu novo}" or "(stat. nov.}"
after a species name to indicate that I am changing its status, raising for example
a subspecies to the level of a species. I now propose to introduce still another
of these expressions, "(species revocata)" or "(sp. rev.}", to draw attention to
the fact that the species referred to is being recalled from wrong synonymy ; " (genus
revocatum) " and " (nomen revocatum) " may be used in a similar way.

POL : OOL is a well known abbreviation in Hymenoptera to indicate the com-
parative distance between the posterior ocelli (i.e., posterior ocellar line = POL)
and the distance between one of these ocelli and the eye margin (i.e., ocellar ocular
line = OOL). In the same way I am using the abbreviation OO-CL to indicate
the distance between a hind ocellus and occipital-carina or the hind margin of the
head where this carina would be if it were developed (i.e., ocellar occipital-carina
line).

ENTOM, III, 7 2I

270 SOME SAWFLIES OF THE EUROPEAN ALPS

CEPHIDAE
Calameuta festiva sp. n.

$. Black except for the following parts which are yellow : apical half of the front
femur, the front tibia and the front tarsus, extreme apex of the middle femur,
sides of the 3rd tergite, whole of the 4th and 5th except medially behind, sides of
the 6th and yth, 8th except the extreme lateral margins, also spot each side of
3rd, 4th and 5th sternites. Wings hyaline with stigma and venation piceous except
for extreme base of C in forewing which is yellow.

Head with eyes slightly closer together in front than the height of one ; distance
between antennal sockets about f as long as the distance between an antennal socket
and the anterior tentorial pit on the same side (ant. ant. : ant. tent, as i-o : 1-5).
Antenna 2i-segmented ; club thickening from 9th to I3th segment ; penultimate
segments about f as long as broad. Thorax and abdomen normal but legs with sub-
bifid tarsal claws and hind tibia with 2 pre-apical spines ; cerci about ^ as long as
sawsheath and reaches back as far ; sawsheath \ as long as basal plate.

Pubescence on head and thorax above dark ; pale and evenly distributed on under-
thorax and whole of abdomen ; on head, pronotum and anterior and lateral lobes of
mesonotum the pubescence is about \ as long as the diameter of an ocellus. Puncta-
tion : head shining between the small follicles on the face but at the back of the
head these are dense and conspicuous ; prothorax likewise has only very fine follicles,
but medial lobe of the mesonotum is dull with fine dense punctures ; lateral lobes
shining with large widely-separated punctures in the middle and behind though
on the anterior J and the outer lateral margin the punctures are a little smaller and
closer together though much larger than those on the anterior lobe, and the inter-
spaces are still shining ; scutellum almost impunctate ; mesopleura covered with
large shallow follicles. Abdomen with the surface roughened by the dense follicle.
Length 9-5 mm.

CYPRUS: Yerasa, 1000 ft., i $ (Holotype), 2.iv.i945 (G. A. Mavromoustakis)
(British Museum).

This species is superficially like C. gaullei (Konow), from Algeria, which, however,
is distinguished by its much more heavily infuscated wings and by the punctation
of the lateral mesonotal lobes (dull with dense punctures as is the anterior lobe),
by the punctation of the anterior margin of the scutellum, and by its ant. ant. :
ant. tent, ratio of i-o : 2-0 (see Benson, 1946). Structurally it is much more like
C. apicicornis Pic (on the basis of a $ collected at Jerusalem, 2.iv.i94i, by Mr. H.
Bytinski-Salz) which has very similar sculpture, but this species can be distinguished
by its rich marking of yellow on the face and the thorax, and by the longer saw-
sheath (sawsheath : basal plate as i-o : 1-8) with its broad subtruncate apex when
viewed from above. C. idolon Rossi is distinguished by its rich yellow colouring
on face and pronotum, its yellow costa and stigma in the forewing, and the thicker
club of its antenna (the subapical segments of which are about twice as broad as
long), and its single pre-apical spine on the hind tibia as well as by its ant. ant. :
ant. tent, ratio of i-o : 2-0.

SOME SAWFLIES OF THE EUROPEAN ALPS 271

ARGIDAE
Kokujewia palestina sp. n.

$. Head black. Thorax with mesonotum including scutellum reddish yellow
(except for the depressed lateral and hind margins of the side lobes and the hind
margin of the scutellum which are black) ; underside, metanotum and legs entirely
black. Abdomen reddish yellow except for ist tergite and sawsheath which are
black. Wings smoky with a black spot under the base of the stigma ; stigma and
venation black.

In structure not distinguishable from K. ectrapela Konow except that the saw-
sheath is broadly rounded behind (Figs, i and 2) (instead of narrowing to a rounded
point) and the hind basitarsus is about as long as the three following tarsal segments
together (instead of clearly longer than this). Length 10 mm.

PALESTINE : Wadi Umbaghik, larva on ? Rumex i? (Holotype) emerged iii.i945
(H. Bytinski Salz) (British Museum) .

I ect.

FIGS. 1-2. Sawsheath of Kokujewia spp. from above : (i) ectrapela ', and (2) palestina.

I am indebted to the late Dr. Gussakovskii for giving to the British Museum a
paratype of K. ectrapela (Caucasus and Transcaucasia) with which I have been able
to compare the specimen from Palestine. K. ectrapela differs, in addition to the
characters in the form of the sawsheath and tarsi mentioned above, in having a
black scutellum and a red supra-clypeal area. K. dementi Zirngiebl (Anatolia),
the only other described species in the genus, is said to differ from K. ectrapela in
having a black medial stripe on the mesonotum and red on the frons.

CIMBICIDAE
Abia plana sp. n.

?. Colour dark metallic green ; antenna with apex of 3rd segment, 4th, and
obscurely 5th and 6th brown ; ist, 2nd, base of 3rd and yth black ; labrum and
mouthparts piceous ; legs with coxae, trochanters and femora (except for apices)
black ; apices of femora, tibiae and tarsi yellowish brown, slightly fuscous beneath
on the hind legs. Wings as in A. sericea L. but that the apical cloud is obsolete.

272 SOME SAWFLIES OF THE EUROPEAN ALPS

Head as in A. sericea L. but that the surface is entirely dull with fine coriaceous
sculpture round the orbits and between the punctures. Thorax dull with fine
coriaceous surface sculpture between the punctures which are mostly smaller than
the interspaces between them. Legs as in A. sericea L., but that the inner tooth
of the tarsal claw is broader and longer than the end tooth (Fig. 3). Abdomen
with the tergites flat, not longitudinally arched as they are in A . sericea ; and dull
all over with dense hair follicles in the middle parts of each tergite while the margins
and sides are densely coriaceous and the whole is covered in a fine dense pile ;
hypopygium simple (not produced slightly in the middle as it is in A. sericea ) ;
sawsheath parallel-sided in dorsal view and trifid at apex ; cerci long and extending
back almost as far as the apex of the sawsheath ; saw very similar to that of A .
fulgens. Length 13 mm.

c. As in $, but the eyes above approach to each other to a distance that is little
more than | the diameter of an ocellus, the 4th, 5th and 6th abdominal tergites
each have the central quarter modified as in A. sericea and covered with a dense
fine mat of short black tomentum surrounded by a shining glabrous margin, but,
unlike A. sericea, etc., the modified areas are scarcely depressed and are (as in
A. fulgens) in the same plane as the lateral portions of the same tergites. Length
11-5 mm.

HUNGARY: Retyezath, 1,200-1,800 ft., ig, i? (Holotype), 6-7.^.1937, i <$,
24.v-4.vi.i937 (B. Lipthay) (British Museum).

This species is very close to A . fulgens Zaddach as is shown by the very similar
saws (vide Kangas, 1946, fig. la) and by the fact that in these 2 species alone of the
European species known to me 1 the tomentum patches on the modified portions
of the 4th to 7th abdominal tergites in the male are not depressed below the level
of the rest of the tergite (in A. sericea L., nitens L., candens Konow, melanocera
Cameron, imperialis Cameron and vitilisi Turner the tomentum patches are in
depressions). The new species is, however, at once distinguishable from A. fulgens
by its heavier punctation and by its tarsal claws, which have the inner tooth minute
in A. fulgens but larger than the end tooth in the new species (cf. Figs. 3 and 4).

Corynis semisanguinea (Pic.)

The description by Pic (i9i6[i]) of this species from Algiers and of C. subcarinata
from Greece (1916(2]) seem generally to have been overlooked ; Gussokovskii
(1947), for example, omitted them from his key to the genus. I had in fact already
prepared a description of the specimens before me as a new species and informed
the collector about this when I came upon Pic's paper quite by chance. So far as
Pic's description goes it covers Guichard's specimens very well and I think it probably
refers to the same species, but as Pic's description is very incomplete as well as being
generally inaccessible I include a fuller description herewith :

1 Excluding those species which Kangas places in Abia (Aenoabia) and which really belong to Zaraea
as defined in Benson, 1951.

SOME SAWFLIES OF THE EUROPEAN ALPS 273

$. Reddish yellow with the following parts bronzey black : antennae and head
(except for the fronto-clypeal area below the antennae), longitudinal fleck in the
middle of each of the lateral lobes of the mesonotum, sunken lateral parts of the
mesonotum beside the mesoscutellum, metanotum (except for the meta-post-
scutellum), mesosternum, mesepimeron, front and lower part of mesepisternum,
metapleura, coxae, trochanters, basal half of fore and middle femora, and extreme
base of hind femur, ist and 2nd tergites except at sides above, a medial fleck on
each of the 3rd, 4th and 5th tergites, the medial part of the margins between these
segments, a broad broken band on the hind margin of the 8th tergite, almost the
whole of the underside of the abdomen with the sawsheath (except the apex of the
hypopygium and the gth sternite).

Wings hyaline with stigma costa and rest of venation yellow.

Antenna (Fig. 6) with 3rd segment twice as long as 4th ; 3rd -f 4th = 5th ;
5th a little more than i| times as long as broad (i-o : 1-7). Head with malar space
about as long as | diameter of front ocellus ; POL : OOL as i-o : 0-8 ; POL :
OO-CL as i-o : 0-6. Pubescence on head and thorax grey and upstanding, the longest
hairs being about the same as the diameter of an ocellus ; on abdomen fine, dense
and recumbent.

Punctation : the larger punctures over almost the whole of the upper surface
are interspaced with more numerous very fine punctures ; on the middle of the
mesonotum, scutellum, middle of mesopleura, and middle of the abdomen, the
larger punctures are mostly separated from each other by from 1-2 diameters ;
on the head the punctures are coarser than on the body, and, round the orbits,
between the antennae and on the postocellar area are very densely spaced and partly
confluent ; on the sides of the apical tergites the punctures are also very densely
set. Length 6-5 mm.

TRIPOLITANIA : 75 km. S. of Bou Ngem, 5 $, 4.11.1952 (K. M. Guichard) (British
Museum).

Superficially this species is very like C. sanguinea (Vollenhoven) of which I have
before me i $ from the Canary Islands (Tho. V. Wollaston Coll., B.M. 1869-65).
The punctation in C. sanguinea, however, is very much denser ; on the meso-
notum, for example, many of the larger punctures are contiguous or with interspaces
little more than a diameter. Furthermore in C. sanguinea the 3rd segment in the
antenna is only about i| times as long as the 4th, and the club, forming the 5th
segment, is about twice as long as broad (cf. Figs. 5 and 6). The abdomen is also
paler above in colour, and the stigma and apex of the costa are blackish brown
instead of yellow.

Corynis reticulata sp. n.

$. Black except for the yellowish white tarsi, tibiae and extreme apices of femora.
Wings hyaline ; stigma and rest of venation yellowish brown.

Antenna (Fig. 7) with 3rd segment i| times as long as 4th ; 5th (club) = 3rd
+ 4th ; 5th about twice as long as wide. Head with malar space very short, scarcely

ENTOM. Ill, 7. 2i

274

SOME SAWFLIES OF THE EUROPEAN ALPS

FIGS. 3-4. Hind tarsal claw in Abia spp. : (3) piano,', and (4) fulgens.

FIGS. 5-8. Antenna in Corynis spp. : (5) sanguined, ; (6) semisanguinea ; (7) reticulata ; and

(8) fulvicrtts.

FIGS. 9-10. Penis valve in Dolerus spp. : (9) romanus ; and (10) ihovacicus.

SOME SAWFLIES OF THE EUROPEAN ALPS 275

longer than the greatest breadth of the apical segment of the maxillary palp ;
supra-clypeal area concave above but below and on clypeal area slightly concave
medially ; front margin of clypeus with an excision about the size of the front
ocellus ; POL : OOL is as i-o : 0-7 ; POL : OO-CL as i-o : 0-5. Claws bifid ; inner
front tibial spur about f as long as basitarsus, and outer spur \ as long as inner
spur. Abdomen with 8th tergite unmodified. Punctation : whole insect reticulate
in appearance, so densely covered in punctures that the only interspaces larger
than a puncture are : one immediately adjoining each ocellus, a few in the middle
of the front lobe of the mesonotum and a few on the front of the scutellum. Pubes-
cence outstanding and silvery ; on head and upper mesopleura about as long as
diameter of front ocellus ; on rest of thorax and abdomen about | this length.
Length 5 mm.

PALESTINE : Shapat near Jerusalem, i $ (Holotype), 27.1^.1918 (E. E. Austen)
(British Museum).

This species appears to be nearest to C. andrei Konow and C. similis Mocsary
and runs to the couplet containing these two species in Gussokovskii's key (1947),
but in both these species the antennal club is much shorter than the 3rd and 4th
antennal segments combined. C. similis (known from Crete, Cyprus, Syria, etc.)
has very much shorter pubescence on the whole body, except on the head it is
nowhere as long as | the diameter of the front ocellus ; and the punctures on the
head and thorax are much less dense with abundant shining interspaces larger than
individual punctures ; and it also has a longer malar space (about as long as the
diameter of the front ocellus in the $ and about this diameter in the <$). C.
andrei (Konow) (from Oran) I have not seen but it is described as having the pubes-
cence on the head and mesonotum fuscous and the apical 3rd of the hind femur pale.

Corynis fulvicrus sp. n.

$. Black with the following parts reddish yellow : labrum, mouthparts, suffused
fleck on underside of antennal club, apical \ of front and middle femora above, and
hind femur (except for extreme base above and basal | below) hind tibia and tarsus
(except for the apical tarsal segments which are brownish). Wings hyaline ; stigma,
C. and Sc. yellow ; rest of venation brown.

Antenna (Fig. 8) with 3rd segment f longer than 4th ; 5th (club) = 3rd + 4th.
Head with clypeus excised in front to a depth of about f diameter of front ocellus ;
malar space very short (about as long as greatest breadth of apical segment of the
maxillary palp ; POL : OOL as i-o : 0-8 ; POL : OO-CL as i-o : 0-6. Legs with
inner front tibial spur about f as long as basitarsus ; outer spur about ^ the inner
spur. Abdomen with 8th tergite unmodified.

Punctation mostly very dense with interspaces as large as individual punctures
only on clypeus, middle of supra-clypeal area, in frontal basin adjoining front
ocellus and beside each lateral ocellus, middle of mesonotum and mesoscutellum,
upper mesopleura, and most of 6 basal tergites. Pubescence, dense silvery up-stand-
ing and about as long as diameter of front ocellus, on head mesonotum and whole
mesopleura. Length 7-5 mm.

276 SOME SAWFLIES OF THE EUROPEAN ALPS

ALGERIA: Hamman Ben Hadjar, i $ (Holotype), 31.111.1910 (F. D. Morice)
(British Museum) ; Misserghim, I ?, 1929 (Allauaud and Jeannell) (Paris Museum).

Another $ from Algeria : Chellala, 1895 (de Vauloger) (Paris Museum) agrees
with the above in colour and structure except that the whole punctation is sparser ;
the punctures on all the mesonotum (except the margins), including the scutellum,
and most of the middle of the mesepisternum are separated by interspaces as
large or larger than individual punctures.

TENTHREDINIDAE

SELANDRIINAE

Selandria serva fuscitarsis subsp. n.

This differs from the typical subspecies in that the 4 apical tarsal segments of
the hind legs and middle legs are infuscate and that in the forewings the costa
has little more than the basal \ yellow and almost the apical | black (in the typical
subspecies the basal f of the costa are yellow and only the apical \ black).

CORFU : i <$ (Holotype), S.iv.igia (F. D. Morice) (British Museum).

ITALY : Romagne, 2 <$, i $, 1945 (P. Zangheri) (i <, i $ in Zangheri Coll. ; i $
in B.M.) ; Bologna, Gaibola, i $, 24.^.1950 (G. Grandi) (in B.M.) and i $, 30. iv.
1951 (G. Grandi).

Strongylogaster lineata cypria subsp. n.

This form differs from the typical S. lineata (Christ) in that the hind femora are
entirely pale yellow (instead of infuscate at base) and the two basal antennal seg-
ments are entirely black (instead of yellow).

CYPRUS : near Platania Forest station, 3,500-4,000 ft., 2 $ (including Holotype),
7. v. 1945 (G. A. Mavromoustakis) ; Mt. Troodos, 5,500-6,000 ft., i $, 28. vi. 1937
(G. A. M.) (British Museum).

Since writing this I have seen 3$ of this subspecies from LEBANON : Falonka,
17. v. 1953 (G. A. M.) (British Museum).

Dolerus romanus sp. n.

cJ. Black ; wings hyaline with black stigma and venation.

Head contracted behind the eyes ; clypeus with front half inflexed along a medial
transverse carina and anterior excision not so deep as half total height of clypeus ;
antenna about as long as vein C of forewing, 8th segment being about four times as
long as its basal breadth ; head densely and rather coarsely punctured above without
interspaces larger than the punctures except adjoining each of the lateral ocelli and
each side of the post-ocellar area which is margined laterally by a deep pit ; hind
ocelli nearly as far apart as the distance between an ocellus and the occipital carina
(POL : OO-CL as i-o : 1-2) ; occipital furrow behind the temples and carina well
developed.

SOME SAWFLIES OF THE EUROPEAN ALPS 277

Mesonotum with front lobe heavily punctured, dense at the sides with an impunc-
tate area in front and in the middle with interspaces between the punctures larger
than the punctures ; lateral lobes with smaller shallow punctures spacer towards
the front and sides and denser behind ; scutallum flat and except for a shining
sparcely punctate area in front is coarsely and very densely punctured ; post-tergite
of scutellum with obsolescent surface sculpture and no medial carina ; meso-
pleura very densely and rather finely punctured above, but with the punctures
thinning out below and the mesosternum is almost impunctate.

Abdomen with the ist tergite shining and sparcely punctured, but with the other
tergites densely transversely coriaceously sculptured all over except for a small
medial apical bare patch on the 8th tergite ; $ penis-valve as Fig. 9.

Pubescence very dense and grey and long on head above and the whole of the
thorax, the longer hairs being about as long as i| the diameter of the front ocellus ;
abdomen with a hair patch each side of the middle line on each of the first 4 tergites,
those on the ist tergite being very long ; 5th to 8th tergites clothed all over except
in the middle line. Length 8 mm.

ITALY: Ermilia, Rivola, Fuenza, i $ (Holotype), 18. ii.i95i (P. Zangheri) (in
the Zangheri Collection).

This most interesting new species would run to the megaptems-asper couplet in
my recent key to the British species (Benson, 1952, p. 77) but differs from both these
species and from D. thoracicus Fallen (cf. Figs. 9 and 10) in the form of the penis
valve, in its much denser punctation on the mesonotum and base of the abdomen,
and the much denser and longer pubescence on the abdomen. The penis-valve
appears to be closest to that of the N. American D. sericeus Say but it differs from
that species in almost every other character not common to all the " black " Dolerus,
lacking, for instance, the strong tubercle on the apex of the 8th tergite, the deep
excision of the clypeus and the coarse punctation of the mesopleura. In the pubes-
cent clothing the new species resembles D. nigratus Miiller and the possibility that
it might prove to be the unknown male of the Mediterranean D. rufotorquatus Costa
had to be considered. D. rufotorquatus $ is not known to differ in any way struc-
turally from D. nigratus $, but it would appear that the new species is far too densely
sculptured on the thorax and base of the abdomen to be the male of D. rufotorquatus
which now appears to me to be no more than a southern race of D. nigratus with a
red instead of black pronotum and front lobe of the mesonotum. (Dolerus rufotor-
quatus Costa = D. nigratus rufotorquatus stat. nov.).

BLENNOCAMPINAE
At h alia cuspidata sp. n.

$. Head black except for the clypeus, labrum, mouthparts and underside of the
antennae which are yellow. Thorax and abdomen yellow except for the following
parts which are black : front lobes of mesonotum, a spot covering the posterior
\ of the raised part of each of the lateral lobes, the post-tergite of the scutellum
together ^ with the sunken lateral parts of the metanotum, the mesosternum, the
extreme apices of the front and middle tarsal segments and the sawsheath, Wings

278 SOME SAWFLIES OF THE EUROPEAN ALPS

hyaline ; stigma and apical | of vein C as well as Sc + R i piceous ; rest of venation
yellow.

Impunctate except only for faint hair follicles. Head with clypeus very short
(laterally only about as long as the 2nd antennal segment) and slightly produced
medially (where it is about as long as the ist antennal segment) and sparsely pubes-
cent ; malar space less than \ the diameter of the front ocellus ; distance between
antennal sockets about the same as the distance between an antennal socket and
the nearest eye margin. Antenna i2-i3-segmented ; 3rd segment greater than
4th + 5th ; 6th onwards broader than long. Legs with tibial spurs broad and very
short (inner hind tibial spur about ^ as long as basitarsus and about f as long as
apical breadth of tibia) ; claws with a small middle tooth in addition to the end
tooth. Abdomen with hypopygium as in A. cordata Lep. (see Benson, 1952, p. 82,
fig. 254) and saw (Fig. n) with prominent and sharp marginal teeth very like those
of A. cordata (I.e., fig. 252).

Pubescence on head and mesonotum long and grey but rather sparse, on the
mesopleura it is evenly spread but it becomes very sparse on the mesosternum ;
abdomen entirely glabrous above. Length 6-7 mm.

c. Coloured as in $ but that the whole upperside of the thorax is black (except
only for the declivous parts of the mesonotum round the wing bases) and that the
black spreads also from the mesosternum to cover the lower parts of the espisternum
and the whole of the epimerum to the base of the wings ; the metasternum and the
middle of the ist tergite are also i infuscate.

In structure as in $ except for the sexual segments, and that the malar space is
linear, that the pubescence on the thorax is much denser and covers the under-
thorax evenly ; hypopygium entire behind.

PALESTINE: Jerusalem, i $ (Holotype) i6.iv.ig43, 2 $, i <$, i.v.ig4i, i $,
7 . v . 1943 (H . Bytinski-Salz) (Holotype, i $ and i $ paratype in British Museum ;
2 $ in Bytinski-Salz collection).

There are only 3 species of Athalia with toothed claws previously known (I do
not regard A. galericulata Kontuniemi 1 as anything more than a dark form of A.
scutellariae Cameron such as are often to be found in Britain with the typical form) ;
and of these I can find no structural differences between the darker A. scutellariae
(Europe) and the paler A. flammula Zhelochovtsev (E. Asia), so that I believe they
are but races of the same species. [Athalia galericulata Kontuniemi, 1951 =
scutellariae Cameron, 1880, syn. nov. A. flammula Zhelochovtsev, 1927 = A.
scutellariae flammula Zhel. stat. nov.]

A. scutellariae and also A. dimidiata Konow (Transcaucasia) differ from the new
species in their longer tibial spurs and in the form of their antennae (which have
only the gth segment onwards transverse). The new species is otherwise very
similar to A. dimidiata in structure, having a very similar hypopygium and saw to
that species, though in colour A. dimidiata is very different in having an entirely
black thorax and ist tergite.

1 Likewise A thalia cordatoides Kontuniemi, 1951, and A. longifoliae Kont., 1951, are synonyms of
A. lineolata Lepeletier, 1823, syn. nov.

SOME SAWFLIES OF THE EUROPEAN ALPS 279

Athalia glabricollis meridiana subsp. n.

This subspecies differs from the typical subspecies (see Benson, 1952, p. 81)
in that approximately the basal \ of the veins C and Sc -f- R are yellow (instead of
only about the basal |) and that the mesonotum is more densely pubes cent.

PERSIA : Suva, 2 <, 8 $ (including Holotype), Escalera Coll. (British Museum
1900-61). TURKEY : Ockmen, i $, I2.viii.i939 (F. S. Bodenheimer) (Brit. Mus.) ;
Aksehir, i <, 8.viii.i95i (Wahrman Coll.). PALESTINE : Jaffa, 2 <J, 2 ?, 24. in. 1951
(H. Bytinski-Salz) ; Jerico, i $, 3 . iv . 1943 (H. B.-S.) ; Jordan, Al Maghtas, i ?,
24. ii. 1942 (H. B.-S.).

Empria persephone sp. n.

$. Black except for the following parts which are brownish white to brown :
labrum, mouthparts, the apices of the front and middle femora, fore and lower
side of front tibia and tarsus, fore side of middle tibia, i bases of middle and hind
tarsal segments, and a fleck each side of tergite 2, 3, 4, 5 and 6.

Wings infuscate ; stigma and venation piceous brown.

Head contracted behind eyes ; the whole covered with dark pubescence (about as
long as the diameter of a lateral ocellus) arising from minute tubercles surrounded
by shining interspaces ; occipital carina reaches from mandible almost to level of
top of eyes ; clypeus about as long as the distance between the hind ocelli, sub-
truncate in front, very slightly emarginate, with a small medial tooth continued
back almost to the base of the clypeus as a longitudinal rib ; eyes about \ longer than
broad ; malar space about equal to length of 2nd antenna! segment ; frons as a
raised platform ; frontal furrow very shallow and ill-defined ; hind ocelli further
apart than distance of each from hind margin of head (POL : OO-CL = i-o : 0-8) ;
POL : OOL as i-o : 1-2 ; postocellar area about twice as wide as long.

Thorax shining and impunctate though in places covered with minute tubercles ;
the whole with dense pubescence mainly fuscous in colour ; hind tibial spurs about
as long as apical width of tibia ; claws with a minute medial tooth in addition to
the end tooth. Wing venation normal, with vein m-cu missing in hind-wing.
Abdomen with hypopygium slightly emarginate medially ; penis- valve as in figure
16. Length 6 mm.

FRANCE: Var, Les Args, i $ (Holotype), 15.^.1939 (W. Fassnidge) (British
Museum).

This species would run in Conde's key to European Empria (Conde, 1940) to
E. liturata (Gmelin) and in my key to the British species to couplet 10 which includes
E. liturata (Benson, 1952, p. 86-90). It differs from E. liturata by its infuscate
wings, its very dark colour pattern, its longer antenna (in liturata the subapical
segments are less than 3 times as long as broad), in its flat table-like frontal area, in
its almost truncate clypeus and in its different penis-valve (cf. fig. 16 with fig. 275
in Benson, I.e.).

280

SOME SAWFLIES OF THE EUROPEAN ALPS

Q.

"^i.

CJ

FIGS. 11-15. Ninth and loth marginal teeth of saw of : (n) Athalia cuspidata ; (12)
Monophadnus pallescens ; (13) M. monticola ; (14) M. alpicola ; and (15) Paracharactus
hyalinus.

FIG. 16. Penis valve of Empria persephone.

SOME SAWFLIES OF THE EUROPEAN ALPS 281

Monophadnus alpicola sp. n.

$. Black with the following parts brown to brownish white : labrum, tegula,
apices of femora of all legs, tibia of front and middle legs, and base of tibia of hind
legs and bases of tarsal segments. Wings hyaline ; stigma with the upper |
black and the lower | brown ; rest of venation brown.

Head swollen behind the eyes ; and eyes small so that the temples in dorsal view
appear as long as the eyes ; genal carina short and fading out at about level of the
bottom of the eyes ; clypeus slightly emarginate in front and faintly punctate ;
malar space about \ length of front ocellus ; antenna about as long as costa of
forewing, with 2nd segment about as long as broad, 3rd about \ longer than 4th
segment, 4th-6th of almost equal length, so are yth-gth ; frontal area flat with its
sides continued forwards to join, on the antennal sockets, the conspicuous supra-
antennal crests which border the deep concave antennal furrow ; post-ocellar
area about twice as wide as long ; hind ocelli about as far apart as each is from the
hind margin of the head ; hind orbits with a deep furrow from the top with a line
of coarse irregular punctures in the furrow.

Thorax impunctate ; prepectus to mesopleura absent. Wing venation and legs
as in M . pallescens Gmelin but that the tarsal claws have each a definite middle
tooth in addition to the end tooth. Abdomen mostly impunctate except for very
faint coriaceous sculpture in places ; sawsheath almost twice as long as basal plate
(i-o : 0-6), parallel-sided in dorsal view and truncate at the apex where it is about
as wide as the apex of the apical tarsal segment ; ovipositor about as long as 4
basal tarsal segments ; saw with sharp marginal teeth (Fig. 14). Pubescence pale
and covering whole insect including mesosternum (though it is sparser here) except
for the 4 basal tergites of the abdomen which are glabrous. Length 4-5-6-5 mm.
$ unknown.

SWITZERLAND: Valais, Arolla, 7,000 ft., 2 $ (including Holotype), i8.vi.iQ35,
14 $$, 29. vi. 1935 (/. E. 6- R. B. Benson) ; Les Hauderes, 4-5,000 ft., 2 ?, 6-
27. vi. 1935 (/. E. & R. B. B.}.

Monophadnus, as restricted by Benson (1952, p. 97-98), includes 5 or 6 previously
described species of which 3 are nearctic and 2-3 European. M. alpicola sp. nov.
is distinguished at once from any of these species by its small eyes (so that when
the head is viewed from above the length of the temple behind the eye appears
about as long as the eye from that viewpoint). 1

From M. pallescens Gmelin and M. monticola Hartig it is also distinguished by its
toothed claws, by the form of the teeth on the saws (cf. Figs. 12, 13 and 14), and by
the pubescent clothing of the underthorax, which in these two species is interrupted
by a broad glabrous band at the junction of the mesosternum and episternum and
is extremely sparse on the mesosternum. M. semicinctus Hartig is so different
from any of the species already mentioned that it may perhaps represent a different
generic group, having very short antennae (scarcely f as long as costa of forewing),

1 It is interesting that Empria alpina Benson, another high alpine sawfly we collected in Switzerland
at the same time but which also occurs in arctic regions such as Lapland and the mountain tops of
Scotland, likewise differs from all other species in its genus by its similarly small eyes.

282 SOME SAWFLIES OF THE EUROPEAN ALPS

an elongate 2nd antennal segment, very large eyes having very short temples behind
(in dorsal view eyes about twice as long as temples behind the eyes), and an up-
turned apex to vein A 3 in the forewing.

Superficially the new species is very similar to Paracharactus hyalinus (Konow)
which we collected at the same time and place as the new species. In P. hyalinus
the pubescent clothing is the same, the eyes are not so reduced in relation to the
length of the temple (in dorsal view about 1-3 : i-o), the claws are very similar, but
its elongate antennae at once distinguish it (segments 3, 4 and 5 are subequal in
length) and the saws are different (cf. Figs. 14 and 15).

It would seem that the genera Monophadnus and Paracharactus (-f- Phymato-
cer&psis) are extremely closely related and may even really belong to one series.
The presence or absence of a prepectus to the mesopleura is not the clear-cut charac-
ter that recent writers would have us think. In Paracharactus longicornis (Hartig)
(comb, nov.) (= Monophadnus longicornis Hartig of previous authors) theprepectal
furrow is reduced to a very short pit, in P. hyalinus it is obsolete and only indicated
by an ill-defined depression, while in Dicrostema gracilicornis (Zaddach) the prepectus
itself is reduced to a very narrow flange ; all these three were yet treated by Enslin
(1912-18) as lacking a prepectus.

I characterise Paracharactus as follows :

Blennocampini having antennal segments 3, 4 and 5 of almost equal length,
claws without a basal lobe, a post-genal carina developed on the head below, the
stub to vein A 3 of the forewing simple (not bifid or turned up at apex) and with a
prepectus to mesopleura i defined.

Paracharactus longicornis (Hartig) is attached to Helleborus and this associaton
again suggests the view that Paracharactus and Monophadnus are closely related,
for Monophadnus is so far as is known entirely associated with Ranunculaceae.

Eutomostethus gagathinus meridionalis subsp. n.

This form differs from typical Eutomosthethus gagathinus (Klug) of Europe in
being on the average larger (6-5-7-5 mm. : 5-5-6-5 mm.) and in that the apical
antennal segment is i| to twice as long as the 8th segment (in E. gagathinus gaga-
thinus the apical segment is about \ times as long as the 8th) and the mesonotum
is more densely pubescent.

CYPRUS : Chiffliccondia, near Limassol ; 3 <$, 4 $, 13. in. 1946 ; 2 <$, 3 $ (including
Holotype), 20.iii.i946 ; 3 #, 5 $, 21.111.1946 ; 2^,28.^.1946; and 2 $, 31.111.1946
(G. A. Mavromoustakis) (British Museum).

TENTHREDININAE
Tenthredopsis convergens sp. n.

cJ. Black : except for the labrum, mandibles, yth segment of the antenna and
probably also 8th and 9th (which are missing in the type) which are white ; and
except for the following parts which are reddish brown : palps, femora of all the
legs, tibiae and tarsi of front and middle legs (tibia of hind legs piceous).

SOME SAWFLIES OF THE EUROPEAN ALPS 283

Wings hyaline ; stigma white at the extreme base, but with the apex and the
rest of the venation piceous.

Head clearly contracted behind the eyes, which are large and strongly converging
in front, where they are closer together than the height of an eye (i-o : 1-2) ; malar
space only about as long as the width of the apical segment of the maxillary palp ;
clypeus subtruncate in front and slightly emarginate medially ; antennal sockets
moderately expanded on their inner margins (as in T. excisa Thomson), but the
medial fovea is not deep and is separated behind from the 3-pronged frontal con-
cavity adjoining the front ocellus ; occipital carina well-developed throughout,
but most prominent behind post-ocellar area and genae ; POL : OOL as i-o : 1-8
and POL : OO-CL as 1-0:1-4; postocellar area about twice as broad as long and
defined laterally by very deep furrows. Above, the head is smooth and almost
impunctate except for the hair follicles though the genae and hind orbits are rough ;
and the pubescence is short and dark. Thorax above shining and smooth except
for the follicles and except for some clear punctures on the posterior half of the meso-
scutellum ; mesopleura dull with coarse irregular surface puncturing. Legs normal,
with basitarsus of hind legs equal to three following tarsal segments together.
Wings normal, with external vein surrounding hind pair. Abdomen transversely
coriaceous and evenly clothed in pubescence ; ist tergite with slight medial carina ;
penis valve as in T. excisa group (see Benson, 1952, fig. 310). Length 9-5 mm.

PALESTINE: Elon, i <$ (Holotype), i6.vii.i9~-. (N. Bytinski-Salz) (in British
Museum) .

This species is readily distinguished at once from every other known species in the
genus by its strongly converging eyes in front where they are closer together than
the height of an eye and, correlated with this, by the exceptionally short malar
space and the strongly narrowed head behind the eyes in dorsal aspect. On all
other counts it is a typical Tenthredopsis and there seems no reason to erect for it a
distinct genus.

Sciapteryx costalis corcyrensis subsp. n.

Differs from S. costalis costalis F. in the $ in that the whole of the inner orbits are
white-margined (instead of only the upper half) and that the lower | of the outer
orbits are also white-margined and that the supra-clypeal area is banded across with
white ; the <$ likewise is more profusely marked with white than is 5. costalis cos-
talis, having the whole of the face below the eyes white (except along the post-genal
carina and the tips of the mandibles), as well as a white spot on the mesapisternum.

CORFU : i ^ and i $ (Holotype) (S. S. Saunders Coll.) (British Museum, 1886-
19).

The colour pattern on the head of the new subspecies is similar to that of 5.
sorror Konow which, however, differs from all S. costalis not only in its black veins
C and Sc + R of the forewing but also in the fact that its hindwings are as smoky in
colour as its forewings (all forms of S. costalis have the hindwings subhyaline in
contrast to the smoky forewings).

284 SOME SAWFLIES OF THE EUROPEAN ALPS

Sciapteryx cleopatra sp. n.

$. Black with the following parts yellowish white : palps, labrum (except front
margin) front | of clypeus (except front margin), inner orbits and lower | of outer
orbits, front of tegula and hind margin of pronotum, line along upper side of all
femora and front side of all tibiae (except apical J in hind tibiae and extreme apex in
the other legs), apical and lateral margins of all tergites of abdomen and apical
margins of sternites. Wings yellowish subhyaline ; basal f of stigma, C and Sc + R
of forewing orange in colour ; rest of venation piceous.

Head shining with very dense coarse punctures becoming rugose on frons, thinning
out on post-ocellar area and temples behind the eyes where the punctures are sepa-
rated by large shining interspaces. Malar space about as long as greatest width of
2nd antennal segment ; POL : OOL as i-o : 1-4 ; POL = OO-CL ; postocellar area
about as long as its greatest breadth ; postgenal and occipital carinae continued to
level with top of eyes. Antenna as in 5. costalis F.

Thorax shining between the punctures which are sparse and fine on the medial
parts of the front lobes of the mesonotum, coarse and tending to fuse at sides of the
lobes, dense and small on the front of the lateral lobes, becoming coarse and irregular
behind, large and distinct on the front of the scutellum, smaller behind, but here
the interspaces between them are densely sculptured with fine surface reticulations ;
mesopleura densely rugose above, shining between scattered punctures below ;
legs with tibial spurs short (on hind legs the inner spur is scarcely longer than the
apical breadth of the tibia). Wings normal. Abdomen with transverse alutaceous
sculpture above ; sawsheath and saw very similar to that of S. costalis. Length
7-8 mm.

PALESTINE : Jerusalem, i $ (Holotype), 1929 (5. Tahudhi) " Sciapteryx costalis F.,
c, del. R. Forsius " (British Museum).

EGYPT : Alexandria, i $, 1902 (/. de Joannis) (Paris Museum).

This species is close to 5. costalis but distinguishable from it at once by its sub-
hyaline instead of smoky forewings and by its different punctation (for in S. costalis
the whole of the mesonotum and mesopleura are dull, densely covered with small
irregular punctures and with the interspaces between these punctures dull with
irregular coriaceous sculpture). In 5. costalis the malar space is also shorter (in the
$ about as long as the greatest breadth of the ist antennal segment). 5. cleopatra
is also much more profusely marked with white on the face of the female than is S.
costalis, in fact it is almost as pale here as male 5. costalis and the holotype was
actually identified by Forsius as a male S. costalis.

S. levantina Andre has its wings coloured as in 5. cleopatra but has a different type
of punctation from both species, with small regular wide-spaced punctures on the head
and thorax and dense reticulate surface sculpture between.

Elinor a flaveola (Gmelin) and E. dominiquei (Konow)

In addition to the differences in colour and the differently shaped and segmented
antennae in these two species (flaveola has longer and thinner 9-segmented antennae ;

SOME SAWFLIES OF THE EUROPEAN ALPS

285

dominiquei has shorter more compressed and 8-segmented antennae) there are note-
worthy differences in the shape of the clypeus which have not been mentioned
before :

E. dominiquei (Gmelin) has the front lobes of the clypeus convex with declivous

sides (Fig. 18).

E. flaveola (Konow) has these lobes flat and the sides as though pressed out (Fig.
17).

flav.

mont.

dom.

mac.

guich. 23

FIGS. 17-20. Clypeus of Elinora spp. : (17) flaveola; (i 8) dominiquei ; (19) maculata ; and (20)

guichardi.

FIGS. 21-23. Meso-scutellum of Macrophya spp. : (21) montana ; (22) aphrodite ; and (23) cyrus.

Elinora corynetes (Kirby) (comb, nov.)

Macrophya corynetes Kirby (1882) 1: 264-5, and pi. 10, fig. 3.

Mr. K. M. Guichard collected at Jebel Soda and also at 75 km. south of Bou
Ngem in Tripolitania on 2nd~4th March, 1952, a series of 16 $ of a species of Elinora
very like E. pectoralis Kriechbaumer except that the wings are entirely hyaline
(instead of yellowish) that the vein Sc + R of forewing is piceous (instead of yellow)
and that the 2nd antennal segment is black. 14 of these specimens agree closely
with the type specimen of Macrophya corynetes from Tunis and are presumed to

286 SOME SAWFLIES OF THE EUROPEAN ALPS

belong to that species. Two specimens, however, taken at Jebel Soda on 2nd
March in company with normal specimens differ in that the outer spur of the middle
tibia has a small tooth half-way down the inner side and a flange from the tooth to
the base of the spur ; the spur is, in fact, modified in the same way for cleaning the
antennae as the inner front spur, though to a lesser extent (cf. Figs. 27 and 28).
As no other differences in these insects seem to be correlated with this form of spur it
is assumed to be aberrational, but in E. maculata Kriech. (= syriaca Andre) and in
the two new species which follow here the middle outer spur is apparently normally
modified in this way like a front spur. 1 For the saw of this species compared with
that of E. caspia and guichardi sp. n. see (Figs. 24-26).

Elinora saharensis sp. n.

$. Yellowish white marked with black as follows : head above antennae (except
fleck on inner orbits above and streak continuing from genae to temples behind eyes
almost to vertex), antenna (except basal segment), apical segment of labial palp and
basal parts of labium together with apex of mandible, margins of supraclypeal area,
axis, mesonotum (except for a streak each side of front each side of front lobe) as well
as whole of scutellum (except its post-tergite), metanotum (except for the post-
scutellum) upper edge of mesepisternum, mesosternum and streak on mesepimerum
and mesopleura, legs with a touch on the outer apex of the tibiae the apices and a
line on the outer side of the tarsal segments, basal tergite (except laterally) and front
part of each of the following tergites though the black thins out laterally to end
before reaching the lateral edge of the segment and is smaller on each succeeding
tergite so that on the 8th and gth it occupies no more than the narrow front margin.
Wings hyaline ; stigma, C and Sc (i.e., front half of the fused Sc + R) yellow ; rest
of the venation brown except for the yellow extreme bases.

Head broadened behind the eyes and face very flat ; malar space about equal to
diameter of front ocellus ; antenna 8-segmented with 3rd longer than 4th + 5th ;
clypeus (cf. Fig. 20) almost in one plane and scarcely incurved along the margins,
almost glabrous and smooth except for a few coarse but shallow punctures on the
lobes ; frons and temples smooth between minute scattered punctures ; POL : OOL
as about i-o : 1-5. Thorax shining between very fine scattered punctures which,
however, become thicker on lower part of mesopleura and mesosternum ; outer spur
on middle tibia modified as in corresponding spur on front tibia being stout with a
strong tooth near the apex. Abdomen dull with fine transverse alutaceous surface
sculpture ; sawsheath and saw similar to that of E. pectoralis.

Pubescence white and on head and mesonotum about as long as malar space ;
on mesopleura longer, outstanding and apically curved, becoming adpressed on
mesosternum. Length 10-5 mm.

SAHARA DESERT : Ahaggar Mountains, Oued Tamanrusset, 10 E., 24 N., at
about 1,300 m., i $, 5.^.1928 (Paris Museum).

1 Captain D. B. Baker tells me that in several species of Eucera as well as in some other genera of
bees the middle spur is likewise modified similarly to the front spur and that this is sometimes accompanied
by several modifications of the basitarsus, but I am not aware that the middle spurs are ever so modified
in other Hymenoptera.

SOME SAWFLIES OF THE EUROPEAN ALPS 287

This species is extremely close to E. coynetes which differs in having shorter
pubescence on the mesopleura (not longer than malar space), a simple or but slightly
modified inner spur on the middle tibia, a slightly longer malar space and no dark
colour on the sternites. It may well be that E. saharensis will ultimately be treated
as a fourth subspecies of E. pectoralis together with lindbergorum (Forsius) (Atlas
Mountains), pectoralis (Kriechbaumer) (Algiers), and corynetes (Kirby) (Tunisia and
Tripolitania) ; E, dominiquei (Konow) (W. Europe) and flaveola (Gmelin ) (C. and
S.E. Europe) are also closely related.

E. saharensis is, however, of particular interest in that it is the first sawfly recorded
from the Ahaggar Mountains, probably because so few collectors go there early enough
in the year.

E. saharensis is very similar in general appearance to E. maculata Kriechbaumer
(= syriaca Andre) which has its outer middle tibial spur modified in the same way.
E. maculata is, however, readily distinguished by its thickened clypeus with slightly
projecting anterior lobes and declivous margins (cf. Figs. 19 and 20), as well as by its
slightly convex and more strongly punctured scutellum.

Elinora guichardi sp. n.

$. Black with yellowish white on the base of the mandible, the labrum, a stripe
on the gena, the tegula, and sometimes the ist perapterum, an oblique streak
behind on the mesepisternum and another on the metapleura ; and with reddish
yellow to yellowish white in middle of mandibles, at least front of clypeus, some-
times basal segment of antenna, at least on hind angles of pronotum, legs (though
infuscate throughout and at least on bases and on posterior sides of coxae, apices
of tibiae and most of tarsi), hind margins of tergites laterally with the ventral por-
tions of them entirely, together with the broad hind margins of the sternites. Wings
hyaline , stigma (except lower margin), C and Sc (front half of Sc + R) yellowish
brown ; lower margin of stigma and rest of venation piceous. Otherwise as in E.
saharensis Bens. sp. n., but that the malar space is only about as long as | diameter
of a front ocellus, the 3rd antennal segment is about as long as the 4th -f- the 5th
and about | of 6th. The inner spur of the middle tibia bears a medial inner tooth
and from the tooth a flange runs to the base of the spur (cf. Figs. 27 and 28). Saw
with sharp ventral teeth (Fig. 25).

<. Colour extremely variable as in $ and though the abdomen may be entirely
black (except i for whitish hypopygium and margins to the sternites) it may be
reddish yellow to entirely reddish yellow and in the palest forms the reddish yellow
spreads over the 4th and 5th tergites at least posteriorly, and sometimes even across
the posterior margins of the other tergites also. Wings as in $.

Structure, except for sexual segments, as in $ ; but the malar space is very short
(only about ^ diameter of front ocellus) and in one of the males (out of 8 <$) the inner
tibial spur is un-modified. Length 7-9-5 mm.

TRIPOLITANIA : 75 km. south of Bou Ngem, 2 $ (including Holotype), and i <$
(with simple middle tibial spurs), 4.^.1952 (K. M. Guichard) (British Museum) ;
Jebel Soda (Wadi Ghodaifa), 5 <, 3.^.1952 (K. M. G.) ; Wadi Tonzist (51 miles
south of Bou Ngem), i $, 8.iii.i952 (K. M. G.).

288

SOME SAWFLIES OF THE EUROPEAN ALPS

Structurally this very variable species appears to be closest to E. corynetes which
is distinguished from it, however, by having a longer malar space (about as long as
front ocellus in $) and a saw with teeth that are much less sharp and prominent
(cf. Figs. 24-26).

FIGS. 24-26. Ninth and loth marginal teeth of saw in Elinora spp. : (24) corynetes ; (25)

guichardi ; and (26) caspia.

FIGS. 27-28. Inner tibial spurs of Elinora guichardi : (27) foreleg ; and (28) middle leg.

Rhogogaster arctica Kiaer.

Dr. E. Enslin has most kindly presented to the British Museum (Natural History)
i $ Rhogogaster arctica Kiaer which he collected himself in the Frankische Jura of
Bavaria, on 2 . vii . 1933 ; this species was previously only known from arctic and sub-
arctic Europe and is therefore an addition to the known fauna of Central Europe.
Superficially it is very like P achy protasis rapae L., but that the antennae are shorter
with the 3rd segment about i| times as long as the 4th, the stigma of the forewing
is pale, and, of course, except for the generic characters, the flatter and more broadly
emarginate labrum, and the tibia longer than the femur and with shorter spurs.

Macrophya orientalis Mocsary (stat. nov.) and M. rufipes (L.).

Macrophya rufipes var. orientalis Mocsary, 1891, p. 156.

Mocsary described this form as a variety of M. rufipes L. differing from the typical
form in having an entirely black abdomen instead of one banded with red. A series
in the British Museum from S.W. PERSIA, B.M. 1900-61 (Escalera Coll.) evidently
belonging to the form described by Mocsary differs so markedly from M. rufipes in

SOME SAWFLIES OF THE EUROPEAN ALPS 289

sculpture as to indicate that the two belong to different species, distinguishable as
follows :

M. orientalis Mocsary. Head with strong and dense punctures on the frontal
area which become finer on the inner orbits ; the surface between the punctures
is shining. Abdomen black with lateral flecks on 6th and yth tergites, dull above
with dense transverse striae, and the pubescence sparse and very short. Wings
smoky. Inner hind tibial spur shorter (spur : basitarsus as i-o : 1-8).

M. rufipes (L.). Head shining between shallow setiferous punctures, the punc-
tures becoming smaller and sparser on the orbits and temples. Abdomen black
with i reddish yellow band on the 3rd and 4th tergites, and in $ with a large
lateral fleck on each side of the 6th tergite, a smaller one on each side of the 7th
and on the middle of the Qth, shining above without surface sculpture between
the follicles and the adpressed hairs from these follicles are longer than the
distance between them so that they overlap. Wings yellowish hyaline. Inner
hind tibial spurs very long (spur : basitarsus as i-o : 1-4).

Macrophya aphrodite sp. n.

$. Black with the following parts yellow : mouthparts, labrum, clypeus, hind
angles of pronotum, tegula, meso-scutellum (except hind margin), and its post-
tergite, fleck on mesopleura, legs (except for black hind coxae, bases of fore and middle
coxae, apical \ of hind femur ; and brownish extreme apices of fore and middle
tibia and tarsal segments ; and for reddish brown hind tibia and tarsus) ; ist tergite
of abdomen almost entirely, 3rd, 4th, 5th and 6th tergites each with a lateral fleck
each side increasing in size progressively so that on the 6th the flecks almost meet
dorsally, yth with a small lateral fleck and gth entirely. Wings yellowish hyaline ;
venation piceous though the lower part of the disc of the stigma is brown.

Head with malar space very short (only about ^ the transverse diameter of the
front ocellus), densely punctured on frons, orbits, vertex and genae, with interspaces
alutaceous and smaller than the punctures ; on temples the punctures are much
sparser so that the interspaces are in places larger than the punctures and with the
surface smooth and shining. Thorax : mesonotum very thickly covered with small
punctures dull with alutaceous sculpture between ; scutellum (fig. 22) in front tumid
and shining, with sparse punctures, but with the posterior quarter depressed and
densely punctured and without a medial keel ; post-tergite about as long as the width
of a cencher, and shining with only 2 or 3 vague punctures and no medial keel ; under-
thorax with dense fine punctures and a few scattered interspaces larger than punc-
tures and with alutaceous surface sculpture. Legs with hind tibia about as long as
hind tarsus ; basitarsus longer than following tarsal segments (as about 1-2 : i-o) ;
inner hind tibial spur more than | as long as basitarsus (as about i-o : 1-6). Fore
wing : anal cell with very short cross vein in the middle. Abdomen with dense
transverse alutaceous sculpture ; saw not distinguished from that of M. moniana.

<$. Coloured as in $ but that the front and middle coxae are black only at the
extreme apex and the hind coxae are only black above on the basal f , the hind tibia

2QO SOME SAWFLIES OF THE EUROPEAN ALPS

and tarsus are on the other hand entirely black (except for a yellow apical outer
spot on the tibia, and on the basitarsus, and for the mainly yellow 2nd and 3rd
tarsal segments), that the yellow flecks on the abdomen are smaller (that on the ist
tergite is medially contracted and the Qth tergite is only yellow at the extreme apex),
Structurally as in $ except for the sexual segments and that the malar space is linear.
Length $ 10-11-5 mm. ; $ 9-5-10-5 mm.

CYPRUS : Episcopi, 7 $ (including Holotype), 9 <$, 14-30. v. 1937 (G. A. Mavro-
moustakis) ; Flatus, i $, 19. vi. 1937 (G. A. M.) ; and Flatus, 3,800 ft., i $, lo.viii.
1937 (G. A. M.).

This species is apparently most closely related to M. montana Scopoli which it
much resembles in colour ; but the stigma is brown instead of piceous and in the
$ the hind tibia and tarsus are reddish brown instead of being black with yellow
flecks (as they are, however, in its <^). In sculpture the head of the new species is
far more densely and finely punctate all over (though not so densely and finely
punctate as is M. postica Brulle). The scutellun (Fig. 22), however, is different in
form, being tumid and smooth and almost impunctate in front, without a medial
keel behind or on the post-tergite (in M. montana it is flatter and dull, with numerous
punctures except on the front ^, and there is a medial keel over the posterior | and
continued across the anterior 3rd of the post-tergite (Fig. 21)).

Macrophya cyrus sp. n.

$. Black with the following parts yellow : mouthparts, labrum, clypeus, i ist,
2nd, and base of 3rd antennal segments, hind angles of pronotum, tegula, meso-
scutellum (except hind margin and post-tergite), fleck on mesopleura, legs (except
coxae and i femora especially inner side of hind pair, but that on the hind legs the
yellow colour has an orange tinge, and the tibia and tarsal segments are brown at
their apices), ist tergite of the abdomen almost entirely, broad apical margins of
3rd to gth tergites (broken medially on the 3rd and 4th, and laterally on 7th and 8th),
also i medial apical flecks and narrow apical margins of sternites except hypopygium.
Wings subhyaline with the forewings slightly infuscate apically (the infuscation
occupies cell 3Rx and i overflows the margins of the surrounding cells) ; stigma,
C, Sc and anal veins of forewing yellow ; rest of venation brown to piceous.

Head with frons and area beside ocelli heavily punctures though with interspaces
as large as punctures, giving way to shining and parsely punctured lower face, orbits
and temples with a large impunctate area each side of post-ocellar region, the inter-
spaces and impunctate areas being without surface sculpture byeond the hair follicles ;
malar space very short (about % diameter of front ocellus). Thorax with mesonotum
and mesopleura heavily punctured and with coriaceous sculpture on the interspaces
which are in places as large as the punctures ; meso-scutellum (Fig. 23) with anterior
f tumid, rounded and almost impunctate ; while the posterior J and post-tergite
are dull with dense coarse punctures and coriaceous sculpture between the punctures,
and are transected by a sharp medial longitudinal keel ; the post-tergite is extremely
short (only about \ width of a cencher). Fore wings with the anal cell constricted
medially for about as long as is the greatest width of the apical portion of the cell.

SOME SAWFLIES OF THE EUROPEAN ALPS 291

Abdomen and legs as in M. aphrodite, but that the saw has very acute marginal teeth.

<. Coloured as in $ but that the fore and middle legs are entirely pale and the
hind femora are only black on the inner side, though the hind tibia may be i infus-
cate below as well as on the inner side, and on the abdomen the ist and 3rd-6th
tergites and all the sternites together with the hypopygium are yellow (except at
the extreme bases, and the 3rd and 4th tergites medially). Structurally as in $
except for the sexual segments and that the malar space is linear. Length $ 10-11
mm. ; <J 8-5-10-5 mm.

S.W. PERSIA : K. Send, 6 $ (including Holotype), 10 $ ; and Bazuft, 2 $, 2 <J.
(Escalera Coll.) (British Museum 1900-61).

This species belongs to the same group as the preceding but forms a sub-group,
distinguished by the yellow stigma, C and Sc of forewing, together with M. postica
Brulle, M. superba Tischbein and M. ottomana Mocsary all occurring in S.E. Europe
and the E. Mediterranean. It is closest to M. superba with which it agrees in general
colour and sculpture including details of the mesoscutellum and its abnormally short
post-tergite. It differs, however, in its wings being subhyaline with apical infus-
cations (uniformly yellowish hyaline in superba) in its much shorter malar space in
the $ ( diameter of front ocellus instead of I) and in the constriction of the anal cell
of the forewing (superba has a short cross- vein). M. postica has much denser punc-
tation on the head with the interspaces coriaceous, a scutellum as in M. aphrodite
Benson sp. n. (see above), uniformly yellowish hyaline wings with but a narrow con-
striction of the anal cell of the forewing and a saw with blunt marginal teeth. M.
ottomana (if I have correctly interpreted this species) was represented by i ^ in the
Escalera Collection from S.W. Persia, K. Sefid, which agrees well with the original
description of this species so far as that goes ; it differs from the new species in having
a scutellum like that already described above for M. aphrodite, with a normal post-
tergite (cf. Figs 22 and 23), wings coloured as in M. cyrus but with anal cell constricted
for only a short way (as in M. postica), a head with denser punctation (as in M.
postica), but without microsculpture between the punctures, antennae entirely black,
hind femur with basal \ yellow and apical | black on both outer and inner sides and
abdomen with much less extensive yellow colouring as follows : a yellow lateral
fleck each side joined by a narrow apical margin on the ist tergite, lateral flecks on
the 3rd to yth tergites (those on the 5th and 6th being much larger than the others
but still interrupted medially), and the apex of the last tergite ; while the hypo-
pygium and sternites are black.

NEMATINAE
Cladius ordubadensis Konow (species revocata)

Mr. Mavromoustakis has taken in Cyprus a long series of this form which was
originally described by Konow (1891, p. 211-12) from the Caucasus and is known
also in the Crimea. In this series there is scarcely any variation in the structure of
the male or female antennae or deviation from Konow's description ; and Zhelochovt-

2Q2

SOME SAWFLIES OF THE EUROPEAN ALPS

sev's view (1952) that this species, together with C. difformis Panzer and C. comari
Stein, are but forms of C. pectinicornis Geoffrey is not acceptable without much
stronger evidence.

opac

FIGS. 30-31.

FIG. 29. Penis valve of Mesoneura lanigera.

Saw of Mesoneura spp. with the clothing of flattened pubescence omitted
(30) lanigera and (31) opaca.

Mesoneura lanigera sp. n.

$. Yellowish brown with the following parts black or piceous : head above
clypeus (except for the supra-clypeal area), antennae, axis, a broken vitta on each
lateral lobe of the mesonotum, suture beside the mesoscutellum, mesosternum,
sunken and lateral parts of the metanotum, bases of coxae and extreme base of front

SOME SAWFLIES OF THE EUROPEAN ALPS 293

and middle femora, apex of hind tibia and hind tarsus, sternites and sawsheath
of abdomen. Wings hyaline ; stigma, C and Sc (i.e, front half of Sc + R) yellow,
rest of venation piceous.

Pubescence : head and thorax covered with dense woolly pubescence, much of it
as long as the long inner spur on the fore tibia, and yellowish white in colour. Abdo-
men with long woolly pubescence below but this is very short and sparse above.

Punctation : head and thorax smooth and shining except for the fine surface
follicles ; abdomen above with dense alutaceous surface and transverse striae.

Head with malar space linear ; clypeus medially excised in front to about | its
total length ; frontal area slightly concave and carinate laterally, but with the frontal
basin confluent in front with the deep antennal groove ; POL : OOL as i-o : 0-7 ;
POL : OO-CL as about i-o : 0-8 ; postocellar area defined laterally with short deep
pits. Antenna almost as long as C of forewing ; 3rd segment a little shorter than
4th and obliquely truncate apically ; 4th-5th ; 6th onwards progressively shorter.

Thorax, wing venation and abdomen as in Mesoneura opaca F., but inner hind tibial
spur about as long as apical width of tibia, sawsheath broadly emarginate behind
about as broad as the apex of the hind femur and saw with more marginal teeth
(cf. Figs. 30 and 31).

$. Black with the following parts yellowish white : mouthparts, base of mandibles,
labrum, front | of clypeus, prothorax, tegulae, i trochanters, apical ^ of front and
middle femora, and hind femur (except extreme base and a line beneath basal |),
tibiae (except apex of hind pair), fore and middle tarsi, and 2nd, 3rd and 4th ter-
gites and front of 5th. Otherwise as in $ except for sexual segments ; hypopygium
broadly truncate at apex, very slightly emarginate medially ; penis valve (Fig. 29).
Length 7-8 mm.

CYPRUS : Pera Pedi, 2,000 ft., 3 $ (including Holotype), 4.^.1952 (G. A. Mavro-
moustakis ; Potamitissa, 3,000 ft., i,<^, 25-26. iii. 1944 (G. A. M.) (in British Museum).

Mr. Mavromoustakis tells me, in a letter dated 19.^.1953, that these specimens
were all captured at flowers of Quercus infectoria Oliv., which is probably the food-
plant as the genus is associated so far as is known entirely with Quercus.

This new species can be distinguished from M. opaca F. by the pale colour of the
abdomen in the $, by the saws (cf. Figs 30 and 31), and by the long woolly golden
pubescence on the head and thorax in both sexes ; in M. opaca this pubescence is
nowhere as long as the inner front tibial spur. The colour pattern and pubescence
would serve also to distinguish the new species from the two other described species
in the genus, M. arquata Klug (C. Europe) and M. macroptera Takeuchi (Japan),
but no male of any species in the genus has been found before.

SAWFLIES OF CYPRUS

CEPHIDAE.

Syrista parreyssi Spinola. (S. Europe and E. Mediterranean to Caucasus.) Mt.
Troodos, Krios River, 5 ^, i $, 15-20. vi.i937. Platus, 3,800 ft., i <^, 3 $,
5-12. vii. 1937.

294 SOME SAWFLIES OF THE EUROPEAN ALPS

Pachycephus smyrnensis J. P. F. Stein. (Balkans and E. Mediterranean.) Amathus,

Kyrenia, Larnaca and Limassol, iii-iv.i93i-5.
Trachelus armenius (Konow). (E. Mediterranean.) Amathus, Larnaka, Limassol,

Mesagitonia and Yermasogia, iii-iv . 1935-46.
T. tdbidus (F.) (C. Europe and Mediterranean to Caucasus.) Amathus, Limassol,

Mesagitonia and Yermasogia, iii-iv. 1935.
Calameuta idolon (Rossi). (Mediterranaean to Caucasus.) Fasoulla, 400-500 ft.,

8 <, 8 $, iii-iv. 1940, on flowers of Sinapis.
C.festiva Benson sp. n. (Cyprus.) Yerasa, 1,000 ft., i $, 2.^.1945.

SlRICIDAE.

Urocems gigas gigas L. (W. Palaearctic.) Limassol, I ?, iv.1932.

Sirex noctilio F. (Holarctic.) 5 c, 3 ?, 25.x. 1927, H. M. Morris, ex pine log.

ARGIDAE

Arge ochropus Gmel. (Mediterranean and Europe to C. Asia.) Flatus, 3,800 ft.,

2 ?, 20. vi. 1937; Pera Pedi, 2,000 ft., i $, i8.vi.i937; Kykko, 2 <$, i ?,

15-17. vii. 1939 (H. Lindberg).
A. cyanocrocea syriaca (Mocsary). (Asia Minor.) Amathus, i $, 23.1^.1935 ; Kato

Amiandos, 3,500 ft., ^.1946; Flatus, 3,800 ft., 2 $, 3 <$, io.vi.i937; Mt.

Troodos, Krios River, 4,500-5,000 ft., 3 <$, i $, 15-17. vi. 1937 ; Yermasogia,

4 $, I2.iii-28.iv.i935 ; Yersa, 1,000-1,200 ft., iii-iv . 1945-47 ; Galata, i <$,

15-21. vi. 1939 (H. Lindberg).
A. nigritarsis Klug. (E. Mediterranean to N. Persia.) Limassol, i <$, i$ , 10-17.

iii.i93i ; Flatus, 3,800 ft., 18 $, 6 $, 11-19^1.1937 ; Pera Pedi, 4,000 ft., i $,

i8.vi.i937; Mt. Troodos, Krios R., 4,500-5,000 ft., 4 <$, 2 ?, 15. vi. 1937;

Yermasogia, 4 $, 13-14.^.1935.
A. proximo, Andre. (E. Mediterranean to Turkmen.) Mandria, 2 $, i6.vi.i937;

Pera Pedi, 2,500 ft., 7 $, 7 $, i8.vi.ig37 ; Flatus, 3,800 ft., n <$, 6 ?, 11-19.

vi.i937 ; Mt. Troodos, Krios R., 4,500-5,000 ft., 10 ^, 16-17. vi. 1937 '> Kykko,

2 c, 3 ?, 15-17. vii. 1939 (H. Lindberg).

ClMBICIDAE.

Corynis similis (Mocsary). (E. Mediterranean.) Amathus and Limassol, iii-iv.
I93I-35-

TENTHREDINIDAE

SELANDRIINAE

Strongylogaster lineata cypria Benson subsp. n. (Cyprus.) Near Platonia Forest
Station, 3,500-4,000 ft., 2 ?, 7. v. 1945; Mt. Troodos, 5,500-6,000 ft., i $,
28. vi. 1937.

SOME SAWFLIES OF THE EUROPEAN ALPS 295

BLENNOCAMPINAE.

Athalia cor data Lepeletier. (W. Palaearctic.) Limassol and Mesagitomia, xii. 1934-

iii.i935; Mt. Koznos, 2,500 ft., 3^.1936 ; Kato, Amiandos, 3,500 ft.,

4.iv.ig46.
Allantus balteatus Klug. (C. Europe and Mediterranean.) Amathus, Limassol,

Mesagitomia, Sphalogiotissa, Yesmasogia, xi-iv and vii. 1933-35.
Empria archangelskii Dovnar-Zapolski. (E. Mediterranean to Caucasus.) Koznos

Mts., 2,500 ft., in. 1936 ; and Ayia, Izini R., 7 miles from Limassol, i <, i $,

25. xii. 1947.
Eutomostethus gagathinus meridionalis Benson subsp. n. (Cyprus.) Chifliccondia,

nr. Limassol, 13-31.1^.1946 ; Akrotiri Bay, 2 <, I $, 26.^.1947.

TENTHREDININAE

Macrophya aphrodite Benson sp. n. (Cyprus.) Episcopi and Flatus, iv-v.i937.
Flatus, 3,800 ft., i <J, 19. vi. 1937, i ?, 10. vii. 1937.

NEMATINAE

Cladius ordubadensis Konow. (E. Mediterranean to Caucasus.) Limassol, Mesagi-
tomia, Sphalagiotisa, and Yermasagia, xi-vii.

Mesoneura lanigera Benson sp. n. (Cyprus.) Pera Pedi, 2,000 ft., 3 $, 4.^.1952 ;
Potamitissa 3,000 ft., i $, 25-26.^1.1944.

Nematus lucidus Panzer. (Palaearctic.) Kellaki, 2,000 ft., i ?, 28.^.1952; Pera
Pedi, 2,000 ft., i $, 4^.1952.

Pristiphora ? sp., near biscalis Forster. Pera Pedi, 2,000 ft., i $, 4.^.1952.

REFERENCES

BENSON, R. B. 1946. Classification of the Cephidae (Hymenoptera Symphyta). Trans. R.
ent. Soc. Lond. 96 (6) : 89-108.

1951-52. Handbooks for the Identification of British Insects, 6 (2-6), Hymenoptera Sym-
phyta, London (R. ent. Soc.)

CONDE, O. 1940. Eine Revision der mir bekannten Empria Arten (Hym. Tenthr.) . . .

Dtsch. ent. Z. 1939-40 : 162-168
GUSSAKOVSKII, V. V. 1935-47. Faune de I'URSS, N.S. Insecta Hym., 2, Chalastogastra,

pt. i (1935) and pt. 2 (1947)-
KANGAS, E. 1946. Uber die Gattung Abia Leach (Hym. Tenthredinidae) im Lichte ihrer

europaischen Arten. Ann. ent. fenn. 12 : 77-122.
KIRBY, W. F. 1882. List Hym. Brit. Mus. 1. London.
KONOW, F. W. 1891. Bemerkungen und Nachtrage zum Catalogus Tenthredinidarum

Europae. Dtsch. ent. Z. 35 : 209220.
KONTUNIEMI, T. 1951. Zur Kenntnis des Lebenszyklus der Sagewespen (Hymenoptera,

Symphyta) in Finland. Act. ent. fenn. 9 : 1-92.

MOCSARY, A. 1891. Tenthredinidae et Siricidae novae. Term. Fuz. 14 : 155-159.
Pic, M. 1916. (i) Sur le genre Amasis Leach, Exchange 32 : 14-15.

1916. (2) Sur le genre Amasis Leach (Deuxieme article). Tom. cit. : 17-18.

ZHELOKHOVTSEV, A. N. 1952. [Revision of the Cladiinae (Hym. Tenthredinidae) of the

U.S.S.R.]. Zool. Zh. 31 : 257-269. [In Russian].

ZIRNGIEBL, L. 1949. Ein Beitrag zur Kenntnis der Tenthrediniden (Hym.1. Mitt. Munch,
ent. Ges. 35-39 : 283-290.

PRESENTED

1 7 MAY 1954

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

24NOV 1954

NEUE NOTIOPHYGIDAE
(COLEOPTERA)

HANS JOHN

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 3 No. 8

LONDON: 1954

NEUE NOTIOPHYGIDAE
(COLEOPTERA)

(Bad Nauheim)

Pp. 297-313 ; Pis. 13-17

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 3 No. 8

LONDON: 1954

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

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

This paper is Vol. 3, No. 8 of the Entomological
series.

PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM

Issued November, 1954. Price Seven Shillings and Sixpence.

Von Hans JOHN

(Bad Nauheim)
SYNOPSIS.

New Notiophygidae.

The paper is based mainly on material collected in South Africa by R. E. Turner, and in
Ceylon and the Far East by Dr. M. Cameron and R. V. de Salvaza.

Two new genera, Pondonatus from S. Africa and Profallia from Singapore, are described,
each with one new species.

Six new species of Aphanocephalus and four of Cephalophanus are also described and figured.

The holotypes of the new species are in the British Museum (N.H.), with the exception of
Aphanocephalus pseudatomus and A. tonkinensis, which are represented by paratypes, the
holotypes being deposited in the Paris Museum.

EINE Sendung Notiophygidae aus den nicht determinierten Bestanden des Briti-
schen Museums in London ergab eine solche Fiille wenig bekannter und neuer
Spezies, dass es geboten erscheint, die Ergebnisse zusammenzufassen und gemeinsam
zu besprechen. Fur die freundliche Bereitstellung des Materials und seine Aus-
sonderung spreche ich Mr. E. B. Britton und Miss C. M. F. von Hayek meinen
verbindlichsten Dank aus.

Die Familie der Notiophygidae (= Discolomidae) umfasst bisher 13 Gattungen,
von denen nur bei Notiophygus, Aphanocephalus und Discoloma Spezies in grosserer
Zahl bekannt geworden sind. Aber auch diese waren meist auf Grand weniger
oder einzeln vorliegender Exemplare determiniert worden, wobei die korperlich
grosseren, 7 bis 8 mm. messenden Gattungen sich in der Vereinzelung ihres Auftre-
tens nicht von den kleinen, i bis 3 mm. messenden unterscheiden. Um so auf-
schlussreicher war daher die vorgelegte Kollektion, welche von einigen Spezies
verschiedener Gattungen ganze Reihen enthielt. Es darf daher angenommen
werden, dass fur die Zahl der erbeuteten Tiere nicht ihre Seltenheit, sondern die
Methode des Sammelns ausschlaggebend ist. Hier mochte ich nicht unterlassen,
dankbar die Namen der Herren R. E. Turner, R. V. de Salvaza und Dr. Cameron
zu erwahnen. Leider fehlen noch fast alle Beobachtungen iiber die Lebensweise
der Tiere und Larven sind bisher erst von zwei Gattungen bekannt geworden,
(Fritz van Emden, Zool. Anz. 101 (1/2), 1932 : Discoloma cassideum Reitter und Arb.
morph. tax. Ent. 5 (2), 1938 : Notiophygus hessei John).

Die Zugehorigkeit fraglicher Kafer zur Familie kann ganz kurz dahin beantwortet

ENTOM. Ill, 8. 23!

300 NEUE NOTIOPHYGIDAE (COLEOPTERA)

werden, dass Notiophygidae an alien Beinen drei Tarsen und dazu drei Paar gleich-
gestaltete kugelige Hiiften haben. Durch die kugeligen Hiiften sind sie auch leicht
von denjenigen Colydiidae zu trennen, welche ebenfalls nur 3 Tarsen besitzen.
Denn die Erwartung Horns (Fauna Hawaii. 3 (5) : 431 1908), dass moglicherweise
Gattungen mil kugeligen Hiiften, wie Discoloma, aber tetrameren Tarsen und anders
gebildeter Fiihlerkeule als tibergange zu den Colydiidae gefunden werden konnten,
hat sich bisher nicht erfiillt. Nach Untersuchungen iiber die anatomischen Unter-
schiede der Colydiidae und Notiophygidae sind solche Ubergange auch nicht zu
erwarten. Dagegen ist anzunehmen, dass nicht nur die Spezieszahl iiberall noch
erheblich steigen wird, sondern dass auch noch neue Gattungen auftreten werden.
Denn Gebiete wie Sudamerika und Australien, aber auch die weiter von der Kiiste
entfernten Lander Asiens sind in dieser Hinsicht noch nicht erschlossen. Daher
sei schon hier gesagt, dass in der vorgelegten Kollektion auch 2 neue Gattungen
gefunden wurden. Im Ubrigen verteilt sich das Material recht ungleichmassig
auf folgende Gattungen : Notiophygus Gory, Discoloma Er., Cassidoloma Kolbe,
Parafallia Arr., Aphanocephalus Woll. und Cephalophanus John.

Notiophygus Gory

N. canus John, Arb. morph. taxon. Ent., Berl. 2.1.1935, p. 16 und 5.2.1938, p. 130.

Die Spezies wurde auf Grund eines <$ Exemplars festgelegt (Typus im Britischen
Museum). Spater wurden 6 Exemplare und jetzt 48 Exemplare determiniert,
die alle am gleichen Ort (Mossel Bay, C. P.) von R. E. Turner gesammelt waren.
Abbildungen : I.e.

Discoloma Erichson

D. sancatarinae John, Beitrage Ent. (Bln-Friedrichshagen), 2.6.1952, p. 618.

Die Spezies wurde nach 8 Exemplaren, leg. Liiderwald, Fundort S. Catarina,
Brasilien, determiniert (Typus ffi und Paratypen im Deutsch. Ent. Inst.).
Material : 1$, nicht ganz ausgefarbt, im Britischen Museum, leg. Plaumann, S.
Catharina (!), Nova Teutonia. Abbildungen : I.e.

Cassidoloma Kolbe

Ct. hymaloides Reitter, Dtsch. ent. Z. 22, 1878, p. 125 (Discoloma).

Material : i Exemplar leg. L. Burgeon, Fundort : Haut Uelle, Moto, im Britischen
Museum. Abbildungen : Arb. morph. taxon. Ent., Berl. 7, 4, 1940, Taf. 7 und 8.

Parafallia Arrow
P. simoni John (Ann. Mag. nat. Hist. (12) 5, 1952 ; 156).

Die Spezies wurde nach 4 Exemplaren der Coll. Grouvelle des Pariser Museums
beschrieben. (Typen im Pariser Museum, Paratypen im Deutschen Entomologi-
schen Institut). Vom Britischen Museum wurden vorgelegt :

12 Exemplare aus Ceylon, Colombo und 19 Exemplare aus ^Singapore, leg.^Dr.

NEUE NOTIOPHYGIDAE (COLEOPTERA) 301

Cameron, Nov./Dez., 1915, welche zu einigen Erganzungen der Beschreibung
Veranlassung geben.

Auf p. 157 I.e. heisst es : . . . die 2. Pore(des Pronotums) liegt dicht an der
Leiste auf einer schmalen ovalen Flache. Die Befunde an 31 Exemplaren ergaben :
Die 2. Pore liegt frei neben der Randleiste, oft ist ihr Rand etwas verbreitert und
bildet einen flachen Ring, seltener ist dieser Ring zu einer kleinen Flache verbreitert,
welche zum Discus hin rund, zum Seitenrand hin spitz ausgebildet ist, ohne mit der
Randleiste zu verschmelzen. Diese letzte Form wurde unter 12 Tieren aus Colombo
3 mal, unter 19 Exemplaren aus Singapore i mal beobachtet. Die Reihe aus Colombo
umfasst Grossen von n X 0*9 mm. bis 1*3 X 1*05 mm. Die Serie aus Singapore
umfasst Grossen von i X 0*85 mm. bis i'2 X i mm. Von beiden Reihen wurden
identische Penisse prapariert. Abbildungen : Taf. 17, fig. la-ic.

Ein Exemplar tragt einen Zettel mit einem Hinweis auf die Lebensweise : ' ' Fruit
of Jaradaga".

Neue Spezies von Aphanocephalus Wollaston

tiber die von Wollaston (1873) in Ent. Mon. Mag. aufgestellte Gattung haben
verschiedene Autoren gearbeitet, zuletzt Grouvelle (Notes Leyden Mus. 34 (2)
1912.). Dabei wurden eine Anzahl Spezies dieser Gattung zugeschrieben, welche
nicht dorthin gehoren. Auf Grund von Material aus dem Britischen und Pariser
Museum konnte ich die Verhaltnisse klar stellen und die Ergebnisse in einer " Re-
vision der Gattung Aphanocephalus " niederlegen, welche zurzeit noch nicht ge-
druckt vorliegt. Soweit Spezies der Gattung Parafallia Arrow zuzuweisen waren,
habe ich dies in der " Parafallia-Studie " Ann. Mag. nat. Hist. (12) 5, 1952, p. 152
erwahnt. Die hier aufgefiihrten neuen Spezies von Aphanocephalus sind in Art
der Beschreibung der " Revision " angeglichen.

Aphanocephalus pseudatomus sp. n.

Die Spezies wurde auf Grund von 4 Exemplaren aus dem Besitz des Pariser
Museums aufgestellt, eine Beschreibung aber bisher nicht veroffentlicht. Die jetzt
in Anzahl aus dem Besitz des Britischen Museums vorliegenden Exemplare, welche
R. V. de Salvaza an gleicher Ortlichkeit sammelte wie A. de Cooman, gaben die
Moglichkeit, die Geschlechter zu trennen.

Kurz behaart. Die Spezies ist in Grosse und Farbung atomus Grouvelle sehr
ahnlich doch durch die starkere Ausbildung der Seitenrander und die viel kraftigere,
Punktierung, besonders der Elytren, sehr gut von ihr zu trennen. Pseudatomus ist
etwas breiter im Umriss und die Basis des Pronotums ist starker konvex als bei
atomus. Zudem ist die Partie an den Vorderecken nur etwas rotlich transparent,
im ganzen aber dunkel, nicht farbig aufgehellt wie bei atomus. Die Punktierung
des Pronotums ist ziemlich kraftig, die der Elytren noch starker und die Pseudo-
poren treten besonders im Basalteil oft zu Gruppen zusammen. Die Behaarung
der Oberflache ist etwa dreimal langer als bei atomus. In der Mitte der Elytren
befindet sich ein rundlicher rotbrauner Fleck mit verschwommenen Randern.
Die Randleiste ist kraftig entwickelt, in ihrer Flache schrag liegend und lasst die

302 NEUE NOTIOPHYGIDAE (COLEOPTERA)

Tuberkelporen besonders im Basalteil deutlich sehen. Die Spezies gehort zu den
Formen, deren Penis distal nicht in zwei seitliche Teile gespalten ist und deren
Peniszunge an Stelle einer distalen Verbreiterung spitz geformt ist und eine heraus-
streckbare chitinisierte Rohre als Fiihrung des sac intern besitzt (Vergl. amterus
John, Ent. BL 37 (5/6), 1941, Taf. 8, fig. 10). Grosse : 17 X 1-3 mm. Material :
Typen und 2 Paratypen im Pariser Museum, 7 Paratypen im Deutschen Entomo-
logischen Institut, 19 Paratypen im Londoner Museum. Fundort : Tonkin, Hoa
Binh, leg. A. de Cooman und R. V. de Salvaza. Abbildungen : Taf. 14, fig. 2-6
(atomus Grouvelle, fig. 7).

Aphanocephalus tonkinensis sp. n.

Kurz behaart, von ovaler Umrissform, glanzend dunkelbraun bis rotbraun,
das Pronotum im ganzen etwas heller. Die Elytren haben einen breiten Seitenrand,
der sich aber zur Spitze hin stark verschmalert. Die leicht erhohten Porenoffnungen
der Tuberkel sind gut sichtbar. Die Punktierung des Pronotums ist von mittlerer
Feinheit, die der Elytren lasst sehr deutlich zwei Formen erkennen, die zarteren
Punkte, aus denen die Haare entspringen und die Pseudoporen, welche wesentlich
grosser, iiber die ganze Flache verteilt, aber im Basalteil besonders gross und haufig
sind. Die Behaarung ist sehr fein und kurz und scheint allgemein auf den Elytren
noch schwacher zu sein als auf dem Pronotum. Die Unterseite ist dunkel, das "i."
Sternit ist gleichmassig grob punktiert, die ubrigen dagegen feiner. Der Penis ist
weichhautig, einfach gebaut, die Zunge (Einlage) ist fast % langer als der Penis-
korper und am Ende breit abgerundet. Grosse : 1-5 X 1*05 mm. Material :
Typen $$ und 21 Paratypen im Pariser Museum, 2 Paratypen im Britischen
Museum, 6 Paratypen im Deutschen Entomologischen Institut. Fundort : Tonkin,
Hoa Binh, leg. A. de Cooman und R. V. de Salvaza. Abbildungen : Taf. 14, fig. 11-14.

Aphanocephalus decoomani sp. n.

Kurz behaart. Die Oberflache ist stark konvex und bei seitlicher Ansicht zeigt
sich die starkste Krummung in der Mitte des Korpers, nach vorn und zur Elytren-
spitze hin ist die Schwingung vermindert. Die Rander am Pronotum und besonders
an den Elytren sind kraftig abgesetzt, die Farbe ist tiefschwarz, am Kopfausschnitt
und am Seitenrand des Pronotums rotbraun transparent und gelegentlich, besonders
im Basalteil der Elytern, mit einem zarten metallischen Hauch, gelblich oder blau-
lich, iibergossen. Die Punktierung der ganzen Oberflache ist fein und gleich-
massig, auf den Elytren sind die Pseudoporen grosser, sehr gleichmassig iiber die
Flache verteilt, finden sich aber in der Nahe des Scutellums oft zu einer Gruppe
zusammen. Bei einer ganzen Anzahl der Tiere zeigt sich auf dem Pronotum eine
Besonderheit. Sie besteht in vier zarten Furchen, welche strahlenformig von der
Mitte der Basis ausgehen und durch eine Anzahl grosserer Punkte ausgezeichnet
sind. Die beiden mittleren Furchen fehlen haufig,manchmal ist uberhaupt nur eine
leichte Schwingung der Oberflache nebst ein paar grosseren Punkten iibriggeblieben.
Im Gegensatz zu hemisphaericus Wollaston und birmanus Dodero ist die ganze Unter-
seite nebst Beinen so dunkel wie die Oberseite. Der Peniskorper ist dorsal an der

NEUE NOTIOPHYGIDAE (COLEOPTERA) 303

Spitze geschlitzt, beide Enden sind mehrfach gezipfelt. Die Zunge hat einen
geschwungenen dicken Rand, der nach hinten (innen) in zwei freistehende scharfe
Dome auslauft. Grosse : 2*25 X 1*7 bis 2*5 X 2 mm. Material : 12 Exemplare
(Typus c$ und Paratypen) im Pariser Museum, 4 Paratypen im Deutschen Entomo-
logischen Institut, I Paratypus im Britischen Museum. Fundort : Tonkin, Hoa
Binh, leg. A de Cooman, 1926, i Exemplar Laos, " betw. Vientiane and Luang
Prabang " 1919, leg. R. V. de Salvaza. Abbildungen : Taf. 14, figs. 15-18.

Aphanocephalus prophysus sp. n.

Das einzelne Exemplar gehort zu den in Gestalt und Farbe einander ahnlichen
Spezies mit je einem Fleck auf den Elytren. Diese mogen hier prophysus in Stich-
worten gegeniiber gestellt werden :

bimaculatus Grouvelle (= austerus John), fast doppelt so gross und starker sculptiert

Australian
modiglianii Grouvelle, pronotum kurzer, der gelbliche Fleck auf den Elytren 1st sehr gross

Engano, Malaconni

vitreus Matthews, transparent, pronotum schmaler, Elytren mit grossem aufgelosten roten Fleck

" China "
atomus Grouvelle, pronotum kurzer und schmaler, Randleiste der Elytren schmaler, Punktierung

von gleicher Feinheit Mana Riang, Ranau,

Palembang, Sumatra, Padang und Mentawai.
binotatus Grouvelle, .... nur auf den Seyshellen .......

pseudatomus n. sp., wie atomus, doch mit viel starkerer Punktierung . . Tonkin, Hoa Binh.

Haare winzig. Gegeniiber den genannten Spezies hat prophysus ein auffallend
grosses und breites Pronotum, der Umriss ist daher verkehrt eiformig. Die Ober-
flache ist ebenso spiegelnd blank wie bei atomus Grouvelle, die etwas sparsamer gesetzte
Punktierung ist ein wenig scharfer, tragt aber ebenso kurze Haare. Die Basis
des Pronotums ist viel starker geschwungen als bei atomus. Auf den Elytren sind
die Pseudoporen weich eingesetzt und erscheinen dadurch etwas grosser. Der
Schulterbuckel ist schwach, die Randpartie ist kraftig aufgekippt, verschwindet
aber an der Spitze fast, die Randleiste ist abgeschragt und zeigt die 6 Tuberkelporen
deutlich. Die iiber der Randpartie befindliche Einschniirung der Elytren, welche
bei den anderen Spezies eine fortlaufende Reihe grosser eingestochener Punkte
tragt, ist hier nur mit wenigen schwachen Punkten besetzt. Der Fleck auf jeder
Elytre ist klein, transparent braunrot und etwa um die Lange seines Durchmessers
von der Sutura entfernt.

Grosse : 1*6 X ri5 mm. Material : i Exemplar (Typus) im Britischen Museum.
Fundort : Singapore, leg. Dr. M. Cameron. Abbildungen : Taf. 14, figs. 8-10.

Vom Britischen Museum erhielt ich 6 Exemplare von Kafern der Gattung Aphano-
cephalus, von denen Grouvelle je zwei mit den Namen "niger" und " sculpturatus "
bezeichnet hatte, wahrend das dritte Paar als potamophilus Lea, cotype bezettelt
war. Beschreibungen sind nicht erschienen. Bei zwei Spezies konnte ich die
Namen belassen, bei sculpturatus i. 1. Grouvelle stelle ich fest, dass ich die Spezies
nach einem einzelnen Exemplar desselben Fundortes und Sammlers bereits als

ENTOM. Ill, 8. 23

304 NEUE NOTIOPHYGIDAE (COLEOPTERA)

pellitus n. sp. in einer Revision der Gattung Aphanocephalus beschrieben hatte.
(Die Arbeit erscheint im 2. Heft der Ent. Bl. 1954).

Aphanocephalus potamophilus sp. n.

Drei australische Spezies sind bisher bekannt geworden : bimaculatus Grouvelle
(= austerus J.), hackeri J. und perlucidus J., welche in Cairns, Brisbane und Rock-
hampton gefunden wurden. Zu ihnen kommt eine von Lea, 1921, gesammelte
Spezies, die Lea selbst als "potamophilus" benannt, aber nicht beschrieben hat.
Leider liegt als Fundort nur die allgemeine Angabe " Australia " vor, doch stammen
die Tiere wahrscheinlich aus den Ausbeuten von den Lord Howe und Norfolk Inseln.

Lang behaart. Gelbbraun bis kastanienbraun, ungefleckt. Das Pronotum
ist im ganzen etwas heller als die Elytren, es ist trapezformig, am Kopfausschnitt
fast gerade geschnitten und die schragen Seitenlinien sind vom Basaltuberkel ab
scharf nach innen umgebogen. Die Basis springt medial zum Scutellum vor.

Die Punktierung ist dicht und kraftig, die Lange der Haare betragt etwa das
5-6-fache der Zwischenraume dieser Punkte.

Der Seitenrand der Elytren ist schmal, aber deutlich bis zur Elytrenspitze vom
Discus abgesetzt durch eine mitlaufende Einschniirung, welche mit grossen Punkten
perlschnurartig besetzt ist. Die Randleiste ist dick und tragt je 6 Tuberkelporen.
An der Elytrenspitze lasst eine fast geradlinige Verbindung dieser Poren den Umriss
kantig erscheinen.

Die Haarpunkte sind nur wenig kleiner als auf dem Pronotum, aber weitlaufiger
gesetzt, die Pseudoporen sind zahlreich, verschieden gross und tief eingelassen,
die grosseren neigen zu ovaler Form.

Der Penis gleicht in seiner Anlage dem von bimaculatus Grouvelle ( austerus J.,
Ent. Bl. 37 (5/6), 1941, p. 193, fig. 10). Der distal spitz endende Peniskorper und
die distal nicht gespaltene Zunge (Einlage) weichen im Bau so weit von dem Typ
ab, der sich bei alien nicht australischen Spezies dieser Gattung bisher gefunden
hat, dass man von einem gesonderten australischen Formkreis sprechen kann.
Grosse 1-9 X 1*4 mm. Material : 2 Exemplare (Typus ^$) im Britischen Museum,
Fundort : Australia, leg. Lea, 1921. Abbildungen : Taf. 15, figs. 1-4, Umriss,
Penis, Pronotum, Sculptur.

Aphanocephalus niger sp. n.

Die bereits von Grouvelle mit dem Namen "niger" bezeichnete aber nicht beschrie-
bene Spezies ist die zweite mir bekannt gewordene Spezies aus Indien.

Kurz behaart. Schwarz glanzend, am Rande des Pronotums und an der Basis
der Elytren mit einem Stich ins rotbraune, breit oval, mit leichter Einschniirung
der Umrisslinie an der Basis des Pronotums und deutlicher Abflachung am Kopf-
ausschnitt. Die Basis ist gegeniiber dem Scutellum stark konvex. Die Randleiste
der Elytren ist schmal, aber bis zur Spitze hin deutlich, nach aussen leicht abgerundet.
Sie tragt je 6 Tuberkelporen in schwachen Verbreiterungen der Leiste. Die Ein-
schnumng des Discus oberhalb der Randleiste ist massig und zeigt nur gelegentlich
kleine punktformige Vertiefungen. Die Punkte des Pronotums, aus denen die Haare

NEUE NOTIOPHYGIDAE (COLEOPTERA) 305

entspringen, sind klein und weich eingesetzt, die Lange der Haare betragt kaum
die Halfte der Zwischenraume dieser Punkte. Auf den Elytren sind die Haar-
punkte fast doppelt so gross, untermischt mil etwas grosseren Pseudoporen, beide
Formen sind weich in die Oberflache eingesenkt, die Lange der Haare ist hier etwas
geringer. Die Farbe der Unterseite ist braun, in der Mitte verdunkelt, Beine,
Fiihler und Mundteile sind gelbbraun. Die Behaarung ist fein, aber doppelt so
lang wie auf der Oberseite. Beim $ ist die mediale Partie des Metasternums vor
den Coxae 3 stark konvex, beim $ dagegen ziemlich flach. Grosse : 1-95 X 1-6
mm. Material : 2 Exemplare (Typus <^$) im Britischen Museum. Fundort :
India, leg. Bowring, 63-47+ un< i Sarda, Bengal, F.W.C., 1927. Abbildungen :
Taf. 15, figs. 5-8, Umriss, Penis, Pronotum, Sculptur.

Neue Spezies von Cephalophanus John

John : Ent. Bl. 36 (3), 1940, p. 81 und 38 (5/6), 1942, p. 171.

Die hier beschriebenen 4 neuen Spezies erganzen in bemerkenswerter Weise die
Kenntnis der Gattung, von welcher bisher nur 2 Spezies (I.e.) aus Chien-Hoa, Tonkin
und Si-Rambe, Sumatra bekannt waren. Die Gattung ist durch zwei Eigentiim-
lichkeiten ausgezeichnet, welche ihr innerhalb der Familie allein zukommen :
die <$<$ haben an den Tarsen des i. Beinpaares gespaltene Klauen und im Innern
ihres Korpers findet sich ein mehr oder weniger langer Chitinfaden, welcher medial
an der Analseite des " I." Sternits entspringt und zum Teil bis in das Metasternum
hineinragt. Da das Ende dieses elastischen Fadens unter der Basis de Penis liegt,
ist seine Funktion wohl zu vermuten, konnte bisher aber nicht eindeutig festgelegt
werden. Die neuen Spezies stellen aber noch andere Probleme. Bisher konnte
Cephalophanus zu den Gattungen gezahlt werden, welche regelmassig 6 Tuberkel
auf dem Seitenrand jeder Elytre besitzen. Aber die in 2 Exemplaren (<$) vor-
liegende Spezies lewisi besitzt jederseits 7 Tuberkel und es fragt sich nun, ob Cephalo-
phanus sich hier wie Notiophygus Gory verhalt, bei welcher Gattung nicht nur die
Zahl der elytralen Tuberkel oft uber 6 hinausgeht, sondern auch haufig eine Zahl-
variation der linken und rechten Elytre gefunden wird, oder ob Cephalophanus,
falls eine Vermehrung der Tuberkelzahl eintritt, diese auf beiden Seiten stets gleich-
massig ausbildet. Weiterhin wurde bei der Spezies dohertyi (1$, 2??) bei einem $
ein abweichend geformtes " i." Sternit gefunden, welches in der Art seiner Aus-
bildung einen vb'llig normalen Eindruck gemacht hatte, wenn nicht die gattungs-
gebundene Form bereits bekannt gewesen ware (Abb. Taf. 13, fig. id). Es handelt
sich hier nicht um eine Deformation, wie sie gelegentlich beobachtet wurde und
durch Verletzungen der Puppe erklart werden kann (Suppl. ent. I.e. 17, 1929, Taf. 9
bei fig. 2) sondern anscheinend um eine Mutante, da die Abweichung beiderseits
vb'llig regelmassig ausgebildet ist. Sie besteht in einem doppelten bogigen Aus-
schnitt des normalerweise zwischen die Coxae 3 zum Metasternum vorspringenden
" i." Sternits, wobei in diesem Fall in der Mitte der Intercoxalprocess als schmaler
Vorsprung stehen bleibt, aber keine Verbindung mehr mit dem Metasternum be-
sitzt. Es entwickelt sich ein tief in den Korper reichender Spalt, der nur durch
eine zarte Haut ausgekleidet ist. Ich erwahne dies so ausfiihrlich, weil ich eine

306 NEUE NOTIOPHYGIDAE (COLEOPTERA)

ahnliche Erscheinung bei der Gattung Solitarius m. (Ent. Bl. 39 (1/2), 1943, p. 28)
als normal angesehen, beschrieben und abgebildet habe. Dort handelt es sich
ebenfalls um zwei tiefe Ausschnitte, die aber zu beiden Seiten der Mittelpartie am
Vorderrand des Mesosternums liegen (S. schaumi m.). Damals stand mir nur i$
zur Verfiigung. Eine Nachprufung bei der zweiten zu Solitarius gehorenden Spezies
des schon von Matthews beschriebenen Aphanocephalus impunctatus (Ann. Ic.
Mag. nat. Hist. (5), 19, 1887) konnte leider nicht erfolgen, da das vorgelegte Exemplar
bereits so stark geklebt war, dass ein volliges Zerfallen zu befiirchten war. Es
taucht also hier der Verdacht auf, dass das der Beschreibung zu Grunde liegende
Exemplar von Solitarius schaumi ebenfalls eine Mutation war. Wenn es stets eine
missliche Sache ist, eine Beschreibung nach einem Unikum auszufiihren, so liegen
die Verhaltnisse bei den Notiophygidae leider so, dass die Tiere nur ausnahmsweise
in grosserer Anzahl gefangen werden, meist werden am selben Ort nur i oder 2
Exemplare zufallig mitgesammelt und gleiche spater nie mehr gefunden.

Cephalophanus bryanti sp. n.

Die Spezies hat eine fast halbkugelige Gestalt, die Farbe ist dunkelbraun, auf
jeder Elytre befinden sich 2 grosse, goldbraune Flecke. Am Pronotum ist die Partie
beiderseits des Kopfausschnittes ebenfalls fleckartig braun, doch kann diese Farbe
sich auch seitlich bis zum 2. Tuberkel ausdehnen und vorn am Kopfausschnitt
zusammenfliessen. Die vordere Ecke ist abgerundet, die dort beginnende Rand-
leiste ist um die Tuberkelpore herum verbreitert und flach, wird im mittleren Teil
schmal und hoch und hat vom 2. Tuberkel ab mehr als die doppelte Breite ihres
mittleren Teiles. Die eingestochenen Punkte stehen im 3- bis 5-fachen Abstand
ihrer Durchmesser und sind auf dem Discus kleiner als auf den Randpartien. Zwi-
schen ihnen stehen sehr zahlreich die kleineren Haarpunkte. Auf den Elytren
fehlen die Basal- und Randflecke der Tuberkel, das Basaltuberkel ist nicht, der
Schulterbuckel ist nur schwach erhoht. Die Randpartie ist gegen den Discus bis
kurz vor die Elytrenspitze abgesetzt, ihre Randleiste ist besonders im vorderen
Teil stark erhoht und tragt jederseits 6 Tuberkelporen in kleinen Anschwellungen.
Die Pseudoporen sind kleiner und stehen weiter voneinander entfernt, als die ein-
gestochenen Punkte des Pronotums, sie sind auf dem Discus sehr zart, verstarken
sich aber zu den Seiten hin und sitzen auch auf der Randpartie. Die Behaarung
der Oberflache ist diinn, aber ziemlich lang, auf den dunklen Teilen ist sie dunkel,
auf den Flecken glanzt sie goldig. Bei den Fiihlern ist das Basalglied braun,
die Geissel samt Keule ist fast schwarz, die Behaarung ist im ganzen dunkel, nur
an der Keule zum Teil auch farblos.

Grosse : 2-85 X 2-6 mm. Material : i<$, i$ (Typen) im Britischen Museum,
Fundort : Mt. Matang, W. Sarawak, leg. G. E. Bryant, Dezember, 1913. Abbil-
dungen : Taf. 13, fig. 3a-y, Taf. 14, fig. la-c.

Cephalophanus dohertyi sp. n.

Die Spezies ist ahnlich clipeoexcisus m. (Ent. Bl. 36, (3), 1940, p. 82), doch besitzt
sie 4 rundliche Flecke auf jeder Elytre. Die Farbe der Oberflache ist dunkelbraun,

NEUE NOTIOPHYGIDAE (COLEOPTERA) 307

die Flecke sind triib rotlichgelb bis gelbbraun, gelegentlich verschwinden sie unter
der langen grauen Behaarung, welche aus langeren und kiirzeren einfachen Haaren
besteht. Das Pronotum hat einen flach-konkaven Kopfausschnitt, der tJbergang
zum Seitenumriss ist vollig abgerundet, wahrend zur Basis hin der Seitenrand mit
der Basis scharf winklig zusammentrifft. Der Seitenrand besitzt eine breite, am
Kopfausschnitt flachgedriickte Leiste. Kurz vor ihrem Anfang sitzt ganz aussen
am Rande die apikale Tuberkelpore. Beim 2. Tuberkel verbreitert sich die Rand-
leiste bogig um die Pore herum und lauft dann in doppelter Breite ihres vorderen
Teiles bis zur basalen Ecke. Die Basis selbst ist beiderseits konkav, in der Mitte
konvex und ist gegeniiber dem Scutellum kurz quer abgestumpft. Im ganzen
ist das Pronotum sehr schwach konvex, basal gegen das 2. Tuberkel hin sogar
schwach konkav. Die Elytren sind breiter als die Basis des Pronotums. Sie
haben eine kleine vorgezogene Schulterecke und besitzen einen schwachen Schulter-
buckel und ein flaches Basaltuberkel. Vom Schulterbuckel her setzt sich eine
flache Randpartie durch eine zum Aussenrand parallele Punktreihe ab. Sie bildet
eine kraftige Randleiste aus, iiber welcher eine 2. Punktreihe dicht entlang lauft,
wahrend der Raum dazwischen im allgemeinen von Punkten frei bleibt. Hier
sitzen nicht erhoht jederseits 6 gelbbraune Tuberkelflecke, wahrend die zu-
gehorigen Poren vor ihnen aussen auf der Leiste liegen. Vom Pronotum ist nur
der mittlere Teil des Discus dunkel gefarbt, Kopfausschnitt und Seiten sind gelb-
braun. Ebenso gefarbt sind auf den Elytren jederseits : das Basaltuberkel, dessen
Pore sich in einigem Abstand von der Basis offnet, 6 Tuberkelflecke auf der Rand-
partie und 4 rundliche Discoidalflecke, die mehr oder weniger deutlich sind. Die
Behaarung ist auf dem Pronotum zarter und infolge aufrechter Stellung der Haare
weniger sichtbar, als auf den Elytren, wo sie mehr anliegt und infolge haufiger
tiberdeckung einen zarten gelbbraunen Pelz bildet. Auf dem Pronotum finden
sich starkere und viel schwachere eingestochene Punkte, aus denen die Haare ent-
springen, die langeren Haare haben zudem kleine kuglige Kornchen als Basis,
welche am Rande der grossen eingestochenen Punkte sitzen. Auf den Elytren
sind die Pseudoporen so gross, wie die eingestochenen Punkte der Seitenteile des
Pronotums. Ihr Abstand voneinander betragt das 4- bis 6-fache ihres Durch-
messers, das Innere liegt flach und zeigt keine Perforationspunkte. Die zwischen
ihnen stehenden Haarpunkte sind kleiner als die kleinen Punkte des Pronotums.

Grosse : 3-95 X 3*25 mm. Material : 2 Exemplare (Typus ^$) im Britischen
Museum, i$ (Paratypus) im Deutschen Entomologischen Institut. Fundort :
Borneo, Peugaron, leg. Doherty. Abbildungen : Taf. 13, fig. ifl-ig.

Cephalophanus fasciatus sp. n.

Bei dieser Spezies ist der Gegensatz zwischen dem kleinen Pronotum und dem
grossen Elytrenteil noch starker als bei dohertyi sp. n. Die Farbe des Tieres ist
dunkelbraun, die Rander des Pronotums und Flecke der Elytren sind lebhaft gelb-
braun. Die vorderen Discoidalflecke der Elytren fliessen zusammen, verbinden
sich mit dem i. und 2. Tuberkelfleck der Randpartie und beriihren sich an der
Sutura in einer erloschenden Fortsetzung, so dass eine deutliche Binde quer iiber

3o8 NEUE NOTIOPHYGIDAE (COLEOPTERA)

die Elytren entsteht. Ebenso fliesst der im Spitzenteil der Elytren befindliche
Fleck mit dem 6. Tuberkelfleck zusammen. Die Behaarung ist etwas kiirzer und
weniger dicht als bei dohertyi sp. n., aber im Gegensatz zu dieser Spezies sind hier
auf den dunklen Stellen der Oberflache die Haare braunlich, auf den Flecken aber
gelbgrau. Beim Pronotum bildet sich am Ubergang zum Seitenrand eine stumpfe
Ecke, in der die apikale Tuberkelpore sitzt. Bereits etwas davor, vom Kopfaus-
schnitt her, beginnt, breit und flach einsetzend, die dicke Randleiste, welche im
mittleren Teil schmaler wird, aber um die basale Tuberkelpore und dann bis zur
scharf geschnittenen Ecke sich wieder auf das doppelte verbreitert. Die Rand-
partie neben der basalen Pore ist etwas aufgetrieben. Die eingestochenen Punkte
des Pronotums, welche die grosseren Haare tragen, sind auf den Seitenteilen etwas
vermehrt und grosser als auf dem Discus. Die Haare stehen excentrisch, ihre
Basis ist nicht so ausgepragt wie bei dohertyi. Die iibrigen kleineren Haarpunkte
fallen wenig auf. Die Elytren haben einen rund erhohten Schulterbuckel und ein
flach aufgetriebenes Basaltuberkel. Die Randpartie ist nicht sehr deutlich abgesetzt,
aber im vorderen Teil bis zum 3. Tuberkelfleck leicht aufgetrieben. Zwischen
den mittleren und hinteren Discoidalflecken ist die Partie um die Sutura bis kurz
vor die Elytrenspitze leicht eingesenkt, so dass hier jede Elytre schwach fur sich
kumuliert. Alle Flecke sind im Umriss verschwimmend eingesetzt. Die Pseudo-
poren sind so gross wie die grossen Punkte des Pronotums, ihre Innenflache ist eben
und die Perforation ist gelegentlich sichtbar. Sie sitzen im 3-bis 6-fachen Abstand
ihres Durchmessers und sind sehr leicht in die Oberflache eingebettet. Die Fiihler,
Mundteile und Beine sind braun, die Fiihler sind im ganzen hell behaart. Grosse :
3'75 X 3'45 mm - Material : i$ (Typus) im Britischen Museum, Fundort : Perak,
leg. Doherty. Abbildungen : Taf. 13, fig. 2a-2c.

Cephalophanus lewisi sp. n.

Ahnlich clipeoexcisus m. (Ent. Bl. 36 (3), 1940, p. 82), doch mit breiterer Basis
der Elytren, kiirzerem Pronotum und 7 elytralen Tuberkeln. Auf den Elytren
fehlen die Flecke am Basaltuberkel, und die Tuberkelflecke der Rander sind nur um
die Pore herum angedeutet. Die 3 Discoidalflecke heben sich schwach gelbbraun
bis kraftig orangefarben von der dunkelbraunen Oberflache ab. Ebenso gefarbt
sind die Rander des Pronotums und die aussere Randleiste der Elytren. Der
Kopfausschnitt am Pronotum ist schwach konkav und lauft rund in die Seitenlinie
iiber. Die breite, oben abgeflachte Randleiste tragt vorn die apikale Tuberkelpore,
ist im mittleren Teil gegen die Innenflache kraftig erhoht und verdoppelt ihre
Breite von der Pore des Basaltuberkels bis zur scharfgeschnittenen Ecke. Der
Discus ist sehr schwach gewolbt, wird zu den Randpartien flacher und ist gegen den
Vorderabschnitt der Randleiste schwach konkav. Die Oberflache ist dicht mit
grossen und kleinen eingestochenen Punkten besetzt, von denen die grossen zu den
Seiten hin ihren Durchmesser verdoppeln. Die Elytren besitzen einen gut gewolbten
Schulterbuckel, aber nur ein massig erhabenes Basaltuberkel. Die Randpartie ist
deutlich abgesetzt und kurz vor der dicken Randleiste kraftig aufgekippt. Sie tragt
jederseits 7 schwache Tuberkel, deren Poren aussen auf der Randleiste sitzen.

NEUE NOTIOPHYGIDAE (COLEOPTERA) 309

Die Pseudoporen der Oberflache sind etwas kleiner als die grossen Punkte von den
Randpartien des Pronotums, sie sind um das 2- bis 4-fache ihrer Durchmesser
voneinander entfernt. Bei giinstiger Beleuchtung sind in ihrer Mitte Perforations-
punkte zu sehen. Die Behaarung besteht aus leicht gebogenen starkeren und
schwacheren Haaren von stumpf grauer oder auch dunkler Farbe, die aber auf den
Flecken gelblich-rotlich aufglanzen. Die Unterseite mit Fiihlern, Mundteilen und
Beinen ist einheitlich braun, die Mitte des Metasternums und die Epipleuren sind
dunkelbraun, die Sternite sind lang und anliegend behaart. Die Fiihler sind in
ganzer Lange hell behaart und die schlanke Keule ist auf ihrer Vorderseite ein-
geschniirt. Grosse : 3 X 2-8 mm. Material : 2 Exemplare (Typus <$$) im Britischen
Museum, Fundort : Kuching, leg. I. E. A. Lewis, 1910. Abbildungen : Taf. 13,
fig. 4~4^.

Pondonatus gen. nov.

Die Gattung steht Notiophygus Gory nahe, hat aber g-gliedrige Fiihler, Fliigel
und andere Merkmale der korperlich kleinen Gattungen der Familie. Beim Pro-
notum fehlt die innere Randleiste, welche bei Notiophygus die Randpartie vom
Discus trennt, an der Basis der Elytren fehlt das Basaltuberkel und auf der Oberflache
ist nur eine Haarform vorhanden. Die Seiten der Elytren sind von der Schulter
bis zum letzten Drittel fast parallel und die Schenkel iiberragen den Korperumriss
nicht. Die Oberflache ist mit tiefen grubenformigen Punkten besetzt, welche wie
bei den anderen Gattungen auf dem Pronotum als " eingestochene Punkte " den
Haaren als Basis dienen, auf den Elytren dagegen " Pseudoporen " sind, welche
in ihrer Mitte eine feine Perforation besitzen, wahrend die Haare in den Zwischen-
raumen entspringen. Das Pronotum hat eine gleichmassig schwach gewolbte
Oberflache und zeigt in einiger Entfernung vom Seitenrand eine mitlaufende leichte
Schwellung (dort, wo bei Notiophygus die innere Randleiste sitzt) und die so gebildete
Randpartie ist in ihrer Flache konkav. Die beiden Tuberkelporen sitzen auf
kleinen Verbreiterungen des Aussenrandes. Pronotum und Elytren sind eng an-
einander geschmiegt, doch ist die Basis der Elytren etwas breiter als die des Pro-
notums und die Schulterecke ist ein wenig vorgezogen. Die Randpartie der Elytren
ist konkav, bis zur Spitze deutlich und vom Discus durch eine fortlaufende Reihe
unregelmassig gesetzter tiefer Punkte getrennt. Die 6 Tuberkel des Seitenrandes
bilden schwache Schwellungen und ihre Poren sitzen aussen am Rande. Die
Haare entspringen auf dem Pronotum innen am Rande der eingestochenen Punkte.
Seitlich betrachtet, bilden sie kleine Hakchen, doch von oben gesehen spalten
sie sich im letzten Drittel und ihre Enden sind weit gespreizt, sie stellen also eine
Haarform dar, die bisher in der Familie noch nicht beobachtet wurde. Auf der
Unterseite steckt der nach vorn gerichtete Kopf bis iiber die Augen im Ausschnitt
des Prosternums. Die Fiihler sind Q-gliedrig und besitzen eine dicke rundliche
Keule. Die Mundteile sind denen von Notiophygus sehr ahnlich. Die Mandibeln
sind 3-zahnig und besitzen eine Lacinia mobilis mit Kammzahnen, die Maxillen
haben eine distal beborstete Lacinia und Galea sowie 4-gliedrige Palpi. Das Labium
hat 3-gliedrige Palpi, welche nahe beieinander eingelenkt sind. Bei beiden Palpi

310 NEUE NOTIOPHYGIDAE (COLEOPTERA)

ist das 2. Glied das grosste. Bei den Beinen erreichen nur die Schenkel des 3. Bein-
paares den Aussenrand des Korpers und die Tibien haben die Lange der Schenkel.
Die mittleren und vorderen Beinpaare haben kiirzere Schenkel und ihre Tibien
sind kiirzer als die Schenkel. Die ersten beiden Tarsenglieder sind zusammen so
lang wie das dritte. Vom Prosternum ab ist die Unterseite mit grossen eingesto-
chenen Punkten besetzt, die in der Korpermitte die Neigung zum Schwinden zeigen.
Von den Epimeren sind nur Reste schmaler Streifen in den Seitenwinkeln des
Meso- und Metasternums sichtbar. Das " Ventralstiick " des 7. Tergits ist schmal.
Die Epipleuren sind mit einer fein punktierten Leiste gegen den Korper abgesetzt.
Die Behaarung der Unterseite ist einfach und sparlich. Die hautigen Fliigel haben
im wesentlichen nur Costa und Media, ein Anallappen fehlt. Der Penis gleicht
dem von Notiophygus, doch ist die Peniszunge (Einlage) starker rohrenformig
gebildet. Die Gattung lebt in Pondoland, Siid-Afrika,. Typus : Pondonatus turnrie.

Pondonatus turneri sp. n.

Die Spezies ist schlank, stumpf braunschwarz und uberall mit mehr oder weniger
tiefen punktformigen Gruben besetzt. Die Behaarung ist locker gestellt, braunlich
bis farblos, transparent. Das Pronotum ist medial am starksten konvex, seine
Randpartien sind durch einen schwachen Wulst vom Discus getrennt und zum
Rande hin konkav ausgebildet. Der Kopfausschnitt ist von oben gut sichtbar
und lasst den Kopf bis zum Fiihleransatz hervortreten. Die Vorderecken am
Seitenumriss, ebenso die Hinterecken sind abgerundet. Bei den beiden Tuberkel-
poren (s. Zeichnung -), welche aussen am Rande liegen, ist der Rand etwas verdickt
und am basalen Tuberkel ist auch die daneben liegende Randpartie etwas auf-
getrieben. Discus und Randpartien sind mit kleinen eingestochenen Punkten
besetzt, die in Grosse und Tiefe etwas variieren. An den Randern dieser Punkte
entspringen die hakenformig gebogenen, distal gespaltenen Haare. Das Scutellum
ist klein, etwas eingesenkt. Die Elytren uberragen seitlich das Pronotum und die
Schulterecke ist dazu ein wenig vorgezogen, so dass der Eindruck entsteht, dass die
Basalecke des Pronotums dort die Basis der Elytren iiberschneidet. Doch sind
beide Teile nur sehr eng aneinander geschmiegt. Der Seitenumriss der Elytren
ist nur schwach konvex, fast gerade, im Spitzenteil lauft die Randlinie halbkreis-
formig zusammen. Der Schulterbuckel ist gut entwickelt, ein Basaltuberkel fehlt
aber (s. Notiophygus). Die Pseudoporen der Oberflache sind bedeutend grosser als
die eingestochenen Punkte des Pronotums und die Haare entspringen zwischen
ihnen. Die Randpartie ist schmal, am Rande konkav und bis zur Spitze deutlich
abgesetzt, sie tragt 6 schwach erhabene Tuberkel beiderseits, deren Poren an der
Aussenseite des Randes liegen. Die Unterseite ist im ganzen schwarz bis schwarz-
braun glanzend, Pro-, Meso-, Metasternum und Sternite sind mit grossen einge-
stochenen Punkten besetzt, aus denen sehr zarte einfache Haare entspringen.
Das " i." Sternit ist sehr breit, die iibrigen schmal, ohne besondere Merkmale.

Grosse : 2-05 X 1-4 bis 2-2 X 1-6 mm. Material : 41 Exemplare (Typus ffi,
Paratypen) im Britischen Museum, 8 Exemplare (Paratypen) im Deutschen Entomo-
logischen Institut. Fundort : 24 Exemplare Port St. John, Pondoland ; 18 Exem-

NEUE NOTIOPHYGIDAE (COLEOPTERA) 311

plare E. Cape Prov., Katberg ; 5 Exemplare Eshowe, Zululand ; 2 Exemplare Natal :
Kloof, leg. R. E. Turner. Abbildungen : Taf. 16, figs. 1-8.

Im Gegensatz zu den Verhaltnissen bei Notiophygus, wo jeder Fundort eine neue
Spezies ergibt, liegt hier ein auffallend weitraumiges Siedlungsgebiet einer Spezies
vor. Es diirfte wohl naheliegen, diese Ausbreitung mit dem Besitz von Fliigeln in
Verbindung zu bringen. Gestiitzt wird diese Ansicht durch die ebenfalls weitrau-
mige Verbreitung der zur gefliigelten Gattung Pmviclava John (Mitt, dtsch.
ent. Ges. 10, (9/10), 1941, p. 103) gehorenden Spezies corrosa m., welche aus der Ge-
gend des Victoria Sees, Insel Ukerewe und aus Kamerun, Makak, bekannt geworden
ist.

Profallia gen. nov.

Als Sharp die amerikanische Gattung Pallia aufstellte (Biol. cent.-amer. Col. 2
(i), 1902, p. 629), betonte er die Verwandtschaft mit Aphanocephalus Woll. und
setzte sich mit den Versuchen auseinander, diese Gattung zu placieren. Er lehnt
Matthews Vorschlag ab, eine Familie der Pseudocorylophidae zu bilden, bezweifelt
aber auch Horns Familie der Discolomidae. Er halt alle diese Formen fur abwei-
chende Colydiidae.

Inzwischen ist die Familie der Notiophygidae (= Discolomidae) nicht nur an-
erkannt worden, sondern ist auf bisher 13 Gattungen angewachsen und ihre Trennung
von den Colydiidae ist auf Grand eindeutiger Merkmale zweifelsfrei durchgefuhrt.
Es zeigte sich, dass die Notiophygidae in Beziehung auf ihren inneren Korperbau,
einschliesslich der Sexualorgane, eine weit grossere Analogic besitzen, als die Coly-
diidae. Sie konnen zudem auf Grand von 2 ausseren Merkmalen auch von den-
jenigen Colydiidae getrennt werden welche innerhalb allgemein tetramerer Gattungen
trimer sind.
Diese Merkmale sind :

(1) Der Besitz von 3 Paar gleichgebildeten kugeligen Hiiften.

(2) Der Besitz offener Poren am Korperrand und zwar wenigstens 2 am Seitenrand
des Pronotums und wenigstens 6 am Seitenrand der Elytren (oberseits oder unter-
seits) .

Die neue asiatische Gattung steht Pallia Sharp naher als Pamfallia Arrow zu Pallia.
Leider sind nur 2 Exemplare einer Spezies vorhanden, doch zweifle ich nicht, dass
diese Gattung sich ebenso verhalten wird, wie ihre nachsten Verwandten, d. h. die
verschiedenen Spezies werden so geringe Unterschiede aufweisen, dass unbedenklich
auch nach einer Spezies die Gattungscharaktere festgelegt werden konnen.

GATTUNGSDIAGNOSE.

Die Grosse der Kafer ist ungefa.hr die gleiche wie bei der Gattung Pallia. Der
eliptische Umriss des Korpers ist bei Pallia leicht eiformig, wobei das stumpfe
Ende vorn am Pronotum liegt. Zugleich ist der Kopfausschnitt mehr oder weniger
breit quer abgeschnitten und gibt dadurch dem Umriss ein rundlich-kantiges Aus-
sehen. Die Gestalt von Profallia ist dagegen schmal-eliptisch, vorn und hinten
vollig abgerandet, vorn ein wenig schmaler als hinten, aber in keiner Weise ei
formig. Von der Seite betrachtet, fallt die Abweichung der Riickenlinie gegen-

312 NEUE NOTIOPHYGIDAE (COLEOPTERA)

iiber Pallia besonders auf : Profallia ist fast | niedriger bei gleicher Lange. Bei
Seitenansicht bildet die Randlinie der Elytren mil der Randlinie des Pronotums
an der Schulterecke bei Pallia einen Winkel von ca. I35oder i rechte Winkel,
bei Profallia ist dieser Winkel bedeutend stumpfer und liegt etwa bei 160. Das
Pronotum bildet fast einen vollen Halbkreis, wenn man die Basis als Durchmesser
betrachtet. Der Kopfausschnitt ist von oben her nicht sichtbar, bei genauerer Prii-
fung findet sich aber ein leichter, sehr flachwinkliger Kerb und von vorn gesehen
bildet der Kopfausschnitt einen sehr flachen Bogen, der ohne Ecke rund in die Seiten-
linie iibergeht. Dort setzt eine schmale Randleiste an, welche bis zur Basis reicht.
Am Ansatz dieser Leiste liegt die nach vorn geoffnete i. Pore, wahrend die 2. etwas
vor der Basalecke in einer schwachen Verbreiterung der Randleiste liegt. Die
Oberflache des Pronotums ist glatt und gleichmassig gekriimmt, an den Seiten,
besonders aber vor dem Kopfausschnitt ist die Krummung etwas starker. Die
Basis ist medial schwach konvex, aber vor den Basalecken konkav. Das Scu-
tellum ist etwas breiter als lang. Die Elytren sind im ganzen starker konvex als
das Pronotum und ihre Schulterecke ist abgerundet. Die dort plotzlich breit ein-
setzende Randleiste ist mehrfach breiter als die des Pronotums und zeigt an ihrem
Ansatz einen Kerb, in welchem die Basalecke des Pronotums ruht (wie bei Pallia
und Aphanocephalus). Die Flache dieser gegen den Discus scharf abgesetzten
Leiste ist etwas schrag nach aussen geneigt und zeigt sehr deutlich 6 Poren, welche
auf Verbreiterungen der Leiste liegen. Seitlich betrachtet krummt sich die Rucken-
linie vor der Spitze soweit rucklaufig, dass die Randleiste dort von oben her nicht
mehr zu sehen ist. Das Pronotum ist mit sparlich stehenden eingestochenen Punkten
besetzt, welche sehr feine kurze Haare tragen (bei 6o-facher Vergr. zu sehen). Die
Oberflache der Elytren zeigt deutlich 2 Formen von Gruben, die kleineren Haar-
gruben und die grosseren Pseudoporen. Der Kopf ragt bis zum Fuhleransatz
aus dem Pronotum heraus, er ist sparlich aber viel langer behaart als die Oberflache.
Der Clipeus ist durch eine scharfe Naht von der Stirn getrennt. Die Fiihler sind
8-gliedrig, das Basalglied ist seitlich etwas kantig, die Keule ist so breit wie lang,
rundlich-dreieckig und flachgedriickt. Das 4. Glied zeigt eine Abschniirung, die
darauf hinweist, dass auch, wie bei Pallia, Spezies mit 9 Gliedern gefunden werden
konnten. Die Mundteile zeigen eine weitgehende Ubereinstimmung mit denen
der nachststehenden Gattungen. Die Mandibeln sind 3-zahnig und besitzen eine
Lacinia mobilis, die Maxillen haben eine distal beborstete Galea und Lacinia und
4-gliedrige Palpi. Die Unterlippe hat 3-gliedrige Palpi. Die Unterseite zeigt im
Bau den Familientyp, das " i." Sternit bildet eine glatte Flache. Die Behaarung
ist sparlich aber langer als auf der Oberseite. Die Schenkel erreichen den Umriss
des Korpers nicht, die Tibien des i. und 2. Beinpaares sind kiirzer als die Schenkel,
die Tibien des 3. Beinpaares sind so lang wie die Schenkel. Bei den Tarsen ist
die dritte so lang wie die beiden ersten zusammen. In Ubereinstimmung mit den
iibrigen Gattungen ist auch die Verlangerung der Hiiften im Innern des Korpers.
Auch hier sind alle drei Paare von gleicher Bauart. Bei den Sterniten ist die ur-
spriingliche Zahl von 7 Sterniten durch Verschmelzen der ersten drei auf fiinf
reduziert und dieses " i." Sternit zeigt im Innern bei vielen Gattungen durch stehen-
gebliebene Leisten seine Herkunft aus 3 Sterniten deutlich an. Bei der vorliegenden

NEUE NOTIOPHYGIDAE (COLEOPTERA) 313

Gattung 1st aber diese Verschmelzung so weitgehend, dass eine Dreiteilung nicht
mehr sichtbar 1st. Die Gattung 1st gefliigelt. Die Alae besitzen nur Costa und
Media, ein Basallappen fehlt. Der Penis gleicht im Grundschema dem von Pamfallia
(der Penis von Pallia ist noch nicht bekannt), doch ist die Basis der Peniszunge
(Einlage) hier so stark ausgebildet, dass sie vom Gelenk ab im Bogen aus der dorsal
geschlitzten Basis des Peniskorpers heraustritt, ehe sie als Rohre ventral hervor-
kommt. Dazu teilt sich von ihrem Grunde her ventral ein schmaler Streifen ab,
welcher stark chitinisiert ist und vielleicht als Fuhning dient. Habitat : Malakka,
Typus : Profattia cameroni.

Profallia cameroni sp. n.

Die Beschreibung ist im wesentlichen in der Gattungsdiagnose enthalten. Hier
folgen noch einige Angaben, welche fur die Trennung von Spezies entscheidend sein
konnen.

Die Gesamtfarbe ist dunkelbraun. Die Elytren tragen in ihrer Mitte je einen
roten verschwommenen Fleck, der die Seite und die Sutura nicht erreicht und
welcher so dunkel ist, dass er nur schwer sichtbar wird. Bei den Elytren sind die
Haarpunkte kraftiger als auf dem Pronotum und sind nebst den Pseudoporen
weich in die Oberflache eingebettet, erscheinen daher oft zu Zweien oder Dreien
zusammenfliessend. Seitlich gesehen, liegt die hochste Erhebung der Korper-
krummung hinter der Mitte, und das Pronotum ist etwas schwacher konvex als der
Spitzenteil der Elytren. Die Randpartie der Elytren zeigt eine deutliche scharfe
Kante als Aussenlinie und auf ihr sitzen die 6 Poren. Im letzten Drittel der Elytren
wolbt sich der Discus ein wenig iiber die Randlinie hinaus, das heist diese Linie unter-
lauft den Discus. Die Fuhler sind 8-gliedrig. Beim 4. Fiihlerglied ist eine Teilung
angedeutet, doch ist keine echte Articulation vorhanden. Die breite Keule zeigt
eine leichte Abschnurung.

Grosse : 1-42 X 0-9 mm. Material : 2 Exemplare (Typus $, Paratypus) im Bri-
tischen Museum, Fundort : Singapore, Dezember, 1915, leg. Dr. Cameron. Ab-
bildungen : Taf. 17, figs, ^a-y und 4-7.

PRESl

24 NOV 1954

TAP. 13.

FIG. la-ig. Cephalophanus dohertyi sp. n. ; ia, Umriss mit Fleck- und Farbverteilung ;
ib, Pronotum von oben ; ic, Unterseite ; id, abweichende Bildung des " i." Sternits bei
einem $ ; le, Sculptur von Pronotum (Pr) und Elytren (El) ; if, gespaltene Klauen eines <$,
nur am ersten Beinpaar vorhanden ; ig, Penis, a, lateral ; b, ventral ; c, halb von hinten ;
d, Penisspitze ventral.

FIG. 2-2c. C. faciatus sp. n. za, Umriss mit Fleck- und Farbverteilung ; 26, Pronotum
von oben ; 20, Sculptur von Pronotum (Pr) und Elytren (El).

FIG. $a-y. C. bryanti sp. n. 30, Umriss mit Fleck- und Farbverteilung ; 36, Pronotum
von oben ; y, Sculptur von Pronotum (Pr) und Elytren (El).

FIG. qa-qd. C. lewisi sp. n. qa, Umriss mit Fleck- und Farbverteilung ; 46, Pronotum
von oben ; 40, Sculptur von Pronotum (Pr) und Elytren (El) ; ^d, Penis : a, lateral, b, ventral,
(Stellung der Seitenteile im Wassertropfen) ; c, ventral (Lage in situ) ; d, Basis dorsal.

Bull. B. M. (N.H.) Entom. Ill, 8.

PLATE 13

TAF. 14.

FIG. i. Cephalophanus bryanti sp. n. Penis, a, lateral; b, ventral; c, mittlere Spitze
halb dorsal.

FIGS. 2-6. Aphanocephalus pseudatomus sp. n. 2, Umriss mit Farbverteilung und Sculptur ;
3, Antennenkeule ; 4, Penis, a, lateral, b, ventral ; 5, Sculptur, Pr, Pronotum, El, Elytren ;
6, Pronotum unverkurzt.

FIG. 7. A. atomus Grouv. : Sculptur zum Vergleich mit pseudatomus, Pr, Pronotum; El,
Elytren.

FIGS. 8-10. A. prophysus sp. n. 8, Umriss mit Farbverteilung und Sculptur ; 9, Sculptur,
Pr, Pronotum, El, Elytren ; 10, Pronotum unverkurzt.

FIGS. 11-14. A. tonkinensis sp. n. u, Umriss mit Farbverteilung und Sculptur; 12,
Sculptur, Pr, Pronotum, El, Elytren ; 13, Penis, a, lateral, b, ventral ; 14, Pronotum un-
verkurzt.

FIGS. 15-18. A. decoomani sp. n. 15, Umriss mit Farbverteilung und Sculptur ; 16, Sculp-
tur, Pr, Pronotum, El, Elytren ; 17 Pronotum unverkurzt ; 18, Penis, a, lateral, b, Spitze halb
dorsal, c, Spitze ventral.

Oberhalb der kreisformigen Sculpturzeichnungen sind die zugehorigen Haarlangen auf
kurzen wagrechten Strichen angegeben.

Bull. B.M. (N.H.) Entom. Ill, 8.

PLATE 14

TAP. 15.

FIGS. 1-4. Aphanocephalus potamophilus sp. n.
FIGS. 5-8. A. niger sp. n.

Butt. B. M. (N.H.) Entom. Ill, 8.

PLATE 15

TAP. 16.
Alle Figuren beziehen sich auf Pondonatus turneri gen. n., sp. n.

FIG. i. Mundteile, a, Labrum ; b, Mandibeln mit den Chitinverstarkungen der Muskeln
des Abductors und Adductors und der Lacinia mobilis ( / ) ; c, Maxille mit 4-gliedrigem Palpus ;
d, Labium mit 3-gliedrigen Palpi.

FIG. 2. Penis, a, lateral ; b, ventral.

FIG. 3. Riickenpartie (Mesonotum, Metanotum, 7 Tergite) nebst Alae und Elytren.

FIG. 4. Aufsicht, rechts ist die Sculptur der Oberflache eingezeichnet, links geben die Pfeile
die Stellen an, an welchen sich aussen am Rande die Tuberkelporen offnen, Nebenzeichnungen :
Die beiden Poren am Rande des Pronotums.

FIG. 5. Unterseite, 5 Sternite, das " i." Sternit entspricht den ersten 3 Tergiten. Zwischen
Analspalte und Elytrenspitze ist das " Ventralstiick" des 7. Tergits sichtbar .

FIG. 6. Kopf, darunter der Clipeus.

FIG. 7. Q-gliedriger Fuhler.

FIG. 8. Haare. a, lateral ; b, dorsal.

Bull. B. M. (N.H.) Entom. Ill, 8.

PLATE 16

TAP. 17.

FIG. la. Parafallia simoni John, ib, links die normale Umrandung der basalen Tuberkel-
pore, rechts eine seltene Abweichung der Umrandung. ic, Penis, /, lateral, v, ventral, dazwi-
schen : Penisspitze halb dorsal.

FIG. 2a. Pallia schmidti John. 26, Pronotum (Ent. Bl. 40, (3/4), 1944, p. 88), beide
Figuren zum Vergleich mit Figs. 3 und 36.

FIG. 30. Profallia cameroni gen. n., sp. n. Umriss, 36, Pronotum ; y, Kopf von oben und
8-gliedriger Fiihler.

FIG. 4. P. cameroni, Unterseite.

FIG. 5. P. cameroni, Riickenpartie mit innerer Verlangerung der Huften (c), Penis (P)
und Fliigel.

FIG. 6. P. cameroni, Mundteile. a, Oberlippe ; 6, Mandibel mit Lacinia mobilis ( / ) ;
c, Maxille mit 4-gliedrigem Palpus, d, Unterlippe mit 3-gliedrigen Palpi.

FIG. 7. P. cameroni, Penis, /, lateral, v, ventral, dazwischen die durch Kreis und Kreuz
bezeichneten Teile vergrossert, ventral gesehen.

Anschrift des Verfassers : Bad Nauheim, Frankfurterstrasse 54, Germany.

Bull. B. M. (N.H.) Entom. Ill, 8.

PLATE 17

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

THE PROTOPTILA GROUP OF THE
GLOSSOSOMATINAE

(TRICHOPTERA : RHYACOPHILIDAE)

MARTIN E. MOSELY

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 3 No. 9

LONDON: 1954

THE PROTOPTILA GROUP OF THE
GLOSSOSOMATINAE

(TRICHOPTERA : RHYACOPHILIDAE)

MARTIN E. MOSELY.

Pp. 315-346 ; 86 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 3 No. 9

LONDON: 1954

THE BULLETIN OF THE BRITISH MUSEUM

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

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

This paper is Vol. 3, No. g of the Entomological
series.

PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM

Issued October, 1954 Price Ten Shillings

THE PROTOPTILA GROUP OF THE
GLOSSOSOMATINAE

(TRICHOPTERA : RHYACOPHILIDAE)
By MARTIN E. MOSELY*

IN 1937, when describing some Mexican Hydroptilidae (Trans. R. ent. Soc. Lond.
86 : 151-190), I remarked of Protoptila : "It is perhaps open to question whether
Protoptila should not be placed in the Rhyacophilidae rather than the Hydroptilidae."
Following this suggestion Dr. H. H. Ross, writing in 1938 (///. Nat. Hist. Surv. Bull.
21 : 112), placed the genus in the Rhyacophilidae, subfamily Glossosomatinae.
For my part I retained the genus in the Hydroptilidae for the time being, as I had
a large amount of Mexican material still unsorted and considered that there was
a possibility that more new genera belonging to the group might be found when
I should be able to spare the time for a further examination. When describing
Brazilian Hydroptilidae (1939, Novit. Zool. 41 : 218-224), I erected two new genera
akin to Protoptila, still retaining the group in the Hydroptilidae. Subsequently
I received a letter from Dr. G. Ulmer in which he wrote "I am now sure that Morto-
niella and Protoptila do not belong to the Hydroptilidae but to the Glossosomatinae
of Rhyacophilidae. ... If Mortoniella and Protoptila are Glossosomatinae, then the
genera Mexitrichia, Canoptila and Antoptila will belong also to this subfamily,
perhaps Padunia Mart. (?)". On receipt of this letter I determined to complete
my examination of the Mexican material and to deal with all the genera concerned.
It was my own inclination to make a new subfamily of the Rhyacophilidae to take
these kindred genera, but as both Ulmer and Ross consider that there is close rela-
tionship in the earlier stages to genera in the Glossosomatinae, I accepted their
view and make the transfer accordingly.

The genus Protoptila, the first to be described in the group, probably contains
a great number of species, but in this paper I propose to confine myself to those
Mexican and Brazilian genera and species with which I am personally acquainted.
Dr. Ross has described many more from North America in his papers on Illinois Trich-
optera. Other genera in the group are Mexitrichia (Mexico and Brazil), Mortoniella
(Brazil), Antoptila (Brazil), Canoptila (Brazil) and a new genus from Mexico, described
in this paper under the name Culoptila, found in the material which I had set aside
for further examination. With regard to Padunia Martynov, if this genus should
really prove to be associated with Protoptila and its allies (of which Ulmer does not
seem to be entirely convinced), attention should be drawn to the far more complete
neuration of the posterior wing, as figured by Martynov, than is to be found in any

* Mr. Martin E. Mosely died in 1948.
ENTOM. Ill, 9. 24

3i8 THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

of the other genera in the group, although that of the anterior wing agrees tolerably
well with at least Antoptila.

All the genera in the group are characterized by a pale streak at the anastomosis,
covering the cross-veins from the radius to the media, which are always aligned,
the remaining cross-veins of the anastomosis generally continuing the line down
to the post-costa. The new genus Culoptila is remarkable for the extraordinary
development of the tegulae which, in the male sex, are enlarged to form hollow
caps, at the base of which is situated a structure that no doubt functions as a scent-
organ. More will be found about this structure in the generic account.

All the species in these genera have a ground-colour of chestnut brown ; in
some of the Brazilian species the anterior wings are decorated with silver bars,
particularly along the anastomosis. When denuded, it is seen that the membrane
at the anastomosis is coloured yellow. It may be added that in the anterior wing
of Protoptila there is a short arc of stiff bristles situated on the membrane between
Cu 2 and the anal veins. In the other genera of the group this arc is represented by
a short straight row of similar bristles arising from the posterior margin of Cu 2 .
All species have a ventral process to the sixth sternite, that of Culoptila being
somewhat peculiar in form.

The Mexican species described in this paper were collected by Dr. Adolph Dampf,
and all the types are deposited in the British Museum (Nat. Hist.).

Finally, I must acknowledge the help given me by Mr. D. E. Kimmins, of the Dept.
of Entomology, who has not only drawn all the figures illustrating this paper, but
has also put his younger eyes at my disposal in making out the very intricate and
obscure structures of the genitalia of this difficult group.

Protoptila Banks

Banks, 1904, Proc. ent. Soc. Wash. 6 : 215. Ulmer, 1906, Notes Leyden Mus. 28 : 99 ; id.,

1907, Gen. Insect. 60a : 220. Betten, 1934, N.Y. Mus. Bull., 292 : 149. Mosely, 1937, Trans.

R. ent. Soc. Lond. 86 : 152. Ross, 1938, III. Nat. Hist. Surv. Bull. 21 : 112 ; id., 1944, op.
cit., 23 : 41.

In the (, the genital characters are very obscure and it is scarcely possible to
homologize them with corresponding parts in other Trichoptera species. In the
descriptions I have already made, I have attempted to follow McLachlan's nomen-
clature of the genital parts, but my naming of such organs as " intermediate appen-
dages," "penis-sheaths " is necessarily arbitrary. I have failed to recognize any
process that could be termed an inferior appendage, though Ross describes these
processes in some of his species. In many of the species, the ventral margin of the
eighth segment is modified to such an extent that it is difficult to avoid the conclusion
that this segment functions as a structure of the genitalia. The description "penis-
sheaths " for the telescoped structures terminating in a strong and often sinuous
spine is perhaps open to criticism, since penis-sheaths of this form are, so far as I
know, unknown in the Trichoptera. I have hesitated as to whether they may not
function as inferior appendages, but their origin seems too near the dorsal half of
the ninth segment for them to be so regarded. What I have termed " intermediate

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

319

appendages ' ' might equally be superior appendages or even processes of the dorsal
plate . In a paper of this nature, it is immaterial what the parts are termed, provided
that the connection between the descriptions and the figures is clearly indicated.

FIGS. 1-3. Neuration of wings of i, Protoptila locula, $ ; 2, P. liqua, $ ; 3, P. resolda, $.

In the wings, the anterior is characterized by an unbroken line of nervures at
the anastomosis, and the variation in the neuration seems practically to be confined
to the base of apical fork no. 2, where in some species it is narrow and pointed and
in others wider and blunt. In the posterior wing, where the neuration is always
reduced, there are numerous irregularities in the different species.

A close consideration of the descriptions of the genitalia and neuration given in

320

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

my papers of Mexican and Brazilian species shows that the genus falls into two
groups, in which the presence or absence of penis-sheaths and variations in the base
of apical fork no. 2 in the anterior wing may be co-related. Thus what I will term
the locula group with narrow base to fork no. 2 and penis-sheaths present contains
the following species : locula, piacha, malica, techila, alexanderi, dubitans, resolda.

piacha

6

techila

FIGS. 4-8. Neuration of posterior wings of 4, Protoptila cavdela, <J ; 5, P. delaca, $ ;
6, P. piacha, $ ; 7, P. techila, <J ; 8, P. malica, $.

What I have termed the liqua group, with the base of fork no. 2 broad and penis-
sheaths absent contains liqua, guata, tojana, delaca, lorada, rota, salta, cardela. Ixtala
is an exception and here the penis-sheaths are present while the base of fork no. 2
is broad in most of the examples examined. I do not consider these small differences

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

321

13

FIGS. 9-13. Protoptila locula, $. g, Genitalia, lateral ; 10, genitalia, dorsal ; n, genitalia,
ventral ; 12, penis, lateral ; 13, penis, ventral.

322 THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

to be of sufficient value to warrant the separation of the two groups into distinct
genera.

Protoptila locula sp. n.

(Figs, i, 9-13)

Insect brownish, neuration as figured.

GENITALIA, <J. Margin of the eighth tergite truncate, the sternite without the
long production found in P. salta, ixtala, rota and resolda. Ninth segment consider-
ably withdrawn within the eighth, from the side only the dorsal and ventral distal
apices being visible. Beyond the margin of the eighth tergite can be seen the centre
of the distal margin of the ninth segment, forming a small, triangular projection.
Intermediate (?) appendages two-jointed ; from above, basal joint visible as an
out- turned, triangular projection, distal joint with a scroll-like head, broadening
towards the margin of the eighth segment. Seen from the side, the appendage is
directed downward, basal joint with an acute, distally directed apex ; distal joint
triangular, directed vertically downward, wide at the base, its apex a pointed,
in-turned hook. Penis-sheaths three-jointed, short, terminal joint the longest.
Penis very complicated ; from the side with an irregularly-formed distal part, a pro-
jecting, blunt hook on the lower margin, and towards the base two pairs of blackened
processes, the distal the larger. The basal pair form two short rods with fringed,
truncate apices, the distal pair fused to make a plate as shown in the figure. There
is the usual enormous, axe-headed extension of the upper surface at the base.

Length of anterior wing, <$, 2 mm.

MEXICO: Guerrera, Cocula, 28.^1.1935, Iguala, 29.1^.1935, 21.1.1936; Micho-
acan, Caracuara, 1197 m., 24-29.^, 3-5^.1935, Nocupetara, 9^.1935; Mexcala,
13 . xii . 1929.

Type, (microscope preparation), Caracuara, 24-29.^.1935.

Protoptila piacha sp. n.

(Figs. 6, 14-20)

Neuration, posterior wing as figured, anterior as in P. locula.

GENITALIA, <. From above, the margin of the eighth tergite slightly convex
and serrate, fringed with long hairs ; sternite strongly produced to nearly as far as
the end of the penis, narrowing to a shallowly excised apex, upper margin from the
side strongly serrate. Ninth segment almost completely withdrawn within the
eighth, only the pointed tip of the ventral margin visible in the side view. Inter-
mediate appendages two- jointed ; from above, basal joint broad and triangular ;
terminal joint arises from a broad base and narrows to a pair of bands curling under
the penis and, as seen from beneath, with the apices divided in two small forks.
From the side, the second joint is long, lower margin sinuous, apex in the form of a
down-turned hook. Penis deep, with the distal margin strongly excised, lower
portion broad, separated from the upper by a shallow, lateral groove in which lies

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

323

the penis-sheath. Lower penis-cover as figured. Penis-sheaths with a very long,
sinuous terminal joint, basal joints obscure, probably three-jointed.

Length of anterior wing, $, 2-25 mm.

MEXICO : Chiapas, Huixtla, 21. xi. 1930 ; Guerrera, Tierra Colorada, i8.xii.i929.

Type, $ (microscope preparation), Tierra Colorada, i8.xii.i929.

FIGS. 14-20. Protoptila piacha, $. 14, Genitalia, lateral ; 15, a penis-sheath ; 16, geni-
talia, dorsal ; 17, genitalia, ventral ; 18, apex of intermediate appendage, dorsal ;
19, penis, lateral ; 20, penis, ventral.

ENTOM. Ill, 9. 2 4

324 THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

Protoptila malica sp. n.

(Figs. 8, 21-26)

Neuration of the posterior wing as figured, anterior as in P. locula.

GENITALIA, <. Distal margin of the eighth segment fringed with stiff hairs,
somewhat serrate towards the distal end of the sternite, which projects only slightly
beyond the margin of the ninth ventral segment. From beneath, the distal margins
of both the eighth and ninth segments are widely excised, the excision of the eighth
being more shallow than the ninth ; from above the eighth tergite is slightly and
widely excised, the margin of the ninth showing beyond it. From above, the basal
joints of the intermediate appendages are large and triangular ; from the side the
appendage is directed downward, both joints rectangular, the basal the shorter and
with a large, curved, lateral lobe obscuring the connection of the joints. Penis from
above broad, apex excised to form two out-turned stout horns ; from beneath,
the excision is wider and the distal surface of the apical joint of the intermediate
appendage is seen to be densely covered with small corrugations and its upper
margin slightly produced to form a hook. Other structures of the penis are shown
in the figures, but I am not venturing to suggest their functions nor to name the
parts. Penis-sheaths three- jointed and very short, the basal joint much the longest.

Length of anterior wing, $, 3 mm.

MEXICO: Colima, 27.1.1930, i < (microscope preparation).

Protoptila techila sp. n.

(Figs. 7, 27-31)

Insect brown. Antennae wanting in the single example before me. There are
differences in the neuration of the posterior wing between this and other species
in the genus, as may be seen by a comparison of the figures. In the anterior wing,
neuration is normal and corresponds with that in P. locula.

GENITALIA, $. Ventral margin of the eighth sternite enormously produced in
a wide plate with a serrate distal margin ; from the side the margin is truncate, with
the upper angle produced in a strong spur. Ninth segment withdrawn into the
eighth, leaving the dorsal portion protruding beyond it so that, from above, it
appears with the centre produced in a small, subacute triangle. The ventral
margin of the ninth segment is completely concealed within the eighth ; from the
side it is much produced and narrows to a sinuous point. The upper penis-cover
from above appears as a pair of long, sinuous processes with strongly dentate outer
margins, inner margins with angular indentations at the centre. Intermediate
appendages two-jointed ; from above, basal joints stout, distal curving inwards
with bifurcate apices, the forks short and pointed, the lower the longer. Penis-
sheaths three- jointed, terminal joint spine-like and sinuous, second joint the smallest,
first joint sinuous. Penis with the usual axe-headed projection of the upper margin
at the base ; it is complicated and fashioned after the pattern of locula, with various
processes on the lower surface (lower penis-cover ?), of which the most conspicuous is

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

325

a pair of strong, up-turned hooks towards the centre, and at the base a pair of rectan-
gular, distally directed processes with fringed apices.

Length of anterior wing, <J, 2-5 mm.

MEXICO: Oaxaca, Rio Chiltepec, 2i.iii.i932, i ^ (microscope preparation).

24

FIGS. 21-26. Protoptila malica, $. 21, Genitalia, lateral ; 22, apex of an intermediate
appendage, from beneath ; 23, genitalia, dorsal ; 24, genitalia, ventral ; 25, penis,
lateral ; 26, penis, ventral.

326

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

FIGS. 27-31. Protoptila techila, $. 27, Genitalia, lateral ; 28, penis-sheath ; 29, genitalia,
dorsal ; 30, penis, lateral ; 31, penis, ventral.

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE 327

Protoptila alexanderi Ross

Ross, 1941, Trans. Amer. ent. Soc. 67 : 48, pi. 2, fig. 14.

Numerous examples of this species were taken in Nuevo Leon, Tamaulipas,
5.vi.i93i.

FIGS. 32-35. Protoptila liqua, <$. 32, Genitalia, lateral ; 33, intermediate appendage,
lateral ; 34, penis, lateral ; 35, penis, ventral.

Protoptila liqua sp. n.

(Figs. 2, 32-37)

Insect brown. The neuration of the posterior wing differing from that of P. techila
as shown in the figure.

328

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

GENITALIA, $. Margin of the eighth tergite armed with an evenly and closely
arranged fringe of long, stiff hairs. Ninth segment completely withdrawn within
the eighth and invisible except in a balsam preparation. Intermediate appendages

r

FIGS. 36-37. Protoptila liqua, <$. 36, Genitalia, dorsal ; 37, seventh, eighth and ninth

segments, ventral.

two-jointed, basal joint from above short and wide, terminal joint armed with two
strongly chitinized spines, the upper from above sinuous and slender, directed
upward, inward and distally, the lower much stouter and directed downward,

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

329

terminating in an acute beak. Penis with the usual axe-headed extension of the
basal upper margin, apex membranous with a pair of large, stout spines at the base
of the membrane, directed downward, a single sinuous spine under the apex, also

39

FIGS. 38-43. Protoptila guata, $. 38, Genitalia, lateral ; 39, genitalia, dorsal ; 40,
genitalia, ventral ; 41, interm ediateappendage, ventral ; 42, penis, lateral ; 43, penis,
ventral.

330

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

directed downward. Towards the base of the penis are two pairs of branches
terminating in beak-like apices, the outer directed inwards, the inner distally ;

FIGS. 44-48. Protoptila tojana, <$. 44, Genitalia, lateral ; 45, genitalia, dorsal ;
46, genitalia, ventral ; 47, penis, lateral ; 48, penis, ventral.

still nearer the base is a pair of chimneypot-shaped processes, apices fringed with
stiff hairs. Penis-sheaths wanting. Lower margin of the eighth sternite furnished
with four pairs of spine-like processes ; from beneath, the shortest pair at the centre

THE PROTOPTILA GROUP OF THE GLOSSOSOMATIN AE 331

with a longer pair outside ; the other two pairs, which are the longest, arise near the
pleurite. Seen from beneath, the outer of these lateral pairs is calliper-shaped, the
inner straight, directed inwards with the apices touching.

Length of anterior wing <$, 4 mm.

MEXICO: Liquidamber, 1130 m., 19.111.1931. La Prusia, 1075 m., 25.111.1931.

Type, <$ (microscope preparation), Liquidamber, 19.111.1931.

Protoptila guata sp. n.

(Figs. 38-43)

Neuration, anterior wing as in P. liqua, posterior as in P. delaca.

GENITALIA, <^. Distal margin of the eighth tergite straight, serrate and fringed
with long hairs ; the sternite is produced in a pair of rather short processes with
truncate apices separated from each other by a wide excision. The upper angle
of the sternite is produced in a long, sinuous spine, sparsely clothed with short hairs
towards the apex. Ninth segment with only its upper margin visible, forming a
wide, shallow, angular projection from above and a short spur from the side ; lower
margin, only to be seen in a cleared specimen, produced in a pair of sharp processes
divided from each other by a wide excision. Intermediate appendages two-jointed,
basal joint large, in side view with its lower angle produced in an up-curved, sickle-
shaped hook ; the second joint is bifurcate, upper fork long, slender and sinuous,
curving downward, lower fork stout, curving upward from the side, somewhat
sausage-shaped and sloping inwards from beneath. Penis dilated at the apex and
excised, with a process on its under surface, wide at the base, terminating in a
chitinous hook directed downward. There is a pair of hooks arising about the centre,
also on the under side and curving downward. Lower penis-cover as figured.
Penis-sheaths wanting.

Length of anterior wing, <, 3 mm.

MEXICO: Sinaloa, Badiraguata, 30.^1.1932, i <$ (microscope preparation).

Protoptila tojana sp. n.

(Figs. 44-48)

Neuration, anterior as in P. liqua, posterior as in P. delaca.

GENITALIA, $. Distal margin of the eighth tergite widely and deeply excised
to show the rounded distal margin of the ninth segment in the excision ; the margin
is strongly fringed, dentate and pigmented. The sternite is only slightly produced
beyond the tergite and bears on each lateral margin a very long spine directed
distally and sparsely fringed towards the apex ; the distal margin of the segment is
not excised but merely slightly concave. Ninth segment partly withdrawn within
the eighth to leave just the apical margin above and beneath visible, the upper
showing as a blunt knob, the lower as a stout, upwardly directed hook. Intermediate
appendages with the terminal joint produced from a broad, stout base in a pair of
strongly chitinized, superimposed, sinuous spines, the upper the shorter ; basal

332

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

FIGS. 49-53. Protoptila delaca, $. 49, Genitalia, lateral ; 50, genitalia, dorsal
51, genitalia and apex of intermediate appendage, ventral ; 52, penis, lateral
53, penis, ventral.

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE 333

joint of the appendage obscure. Penis with a membranous apex containing a
hooked spine arising from its under surface ; from above and beneath, the penis
widens to an excised apex. Lower penis-cover as shown in the figure. Penis-
sheaths wanting.

Length of anterior wing, $, about 2-5 mm. (wing broken at base).

MEXICO: Chiapas, Jonata, 2.xi.i932, i $ (microscope preparation).

Protoptila delaca sp. n.

(Figs. 5, 49-53)

Neuration, posterior wing as figured, anterior as in P. liqua.

GENITALIA, <. Distal margin of the eighth tergite rounded and serrate, fringed
with long, stiff hairs. Sternite cleft, forming a wide, deep excision, base rounded,
the sides of the excision with truncate apices, of which the outer angles are produced
in three or four long, pointed fingers or teeth of varying length. From the side
it may be seen that the lowest of these dentate processes is separated from the others
by a rounded excision. Ninth segment with the dorsal margin withdrawn into the
eighth, ventral margin from beneath appearing in the excision of the eighth as a
stout, blackened, bifurcate structure, the apices of the forks obliquely truncate.
Intermediate appendages two-jointed, curling down and under the penis ; basal
joint broad from above, rather long and narrow from the side ; terminal joint with
a rounded outer lobe and an inner, in-turned spur extending nearly half-way across
the penis ; a second small spur between this spur and the rounded lobe can be made
out in a balsam preparation. Penis from above with the apex dilated in two pairs
of superimposed lobes, the upper the smaller ; arising from the margin of the lower
is a pair of corkscrew spines, the apices directed outwards ; from beneath is seen a
short central spine directed distally. Lower penis-cover as shown in the figures.
Penis-sheaths wanting.

Length of anterior wing, <$, 3-5 mm.

MEXICO : Vera Cruz, Cardel, 3.viii.i932, i $ (microscope preparation).

Protoptila lorada sp. n.

(Figs. 54-58)

Neuration, anterior wing as in P. liqua, posterior as in P. cardela.

GENITALIA, $. Eighth tergite with the distal margin straight and slightly dentate ;
the margin of the sternite with a narrow, rounded and not very deep excision, near
which it is produced in a pair of shortish and very acute projections or spurs. Ninth
segment entirely withdrawn into the eighth ; in a cleared preparation, the dorsal
margin is undulating, ventral produced at its centre and shallowly excised at the
centre of the produced portion. Intermediate appendages two- jointed, basal joint
somewhat rectangular, apical joint bearing three sinuous spines of which the upper
is the longest and, from the side, rather widely separated from the two lower. Penis
with a membranous, excised apex, armed on its lower surface, as seen from the side,

334

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

with three stout spines, of which the most distal is considerably curved, the central
longer and nearly straight, the basal short and beak-like. Structures towards
the base, including two more hooked processes, probably form part of the lower
penis-cover. Penis-sheaths wanting.

FIGS. 54-58. Protoptila lorada, $. 54, Genitalia, lateral ; 55, genitalia, dorsal
56, genitalia, ventral ; 57, penis, lateral ; 58, penis, ventral.

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

Length of anterior wing, $, 2-25 mm.

MEXICO: Guerrera, Tierra Colorada, i8.xii.i92g.

Type, c, mounted as a microscope preparation.

335

FIGS. 59-63. Protoptila cardela, <$. 59, Genitalia, lateral ; 60, genitalia, dorsal
61, genitalia, ventral ; 62, penis, lateral ; 63, penis, ventral.

336 THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

Protoptila cardela sp. n.

(Figs. 4, 59-63)

Neuration, posterior wing as figured, anterior as in P. liqua.

GENITALIA, <. Ninth segment only visible as a small ventral hook, the remainder
being completely withdrawn into the eighth. The dorsal margin of the latter is
straight, fringed with very long hairs ; from beneath it is deeply excised to leave
prominent outer angles separated by a deep, round-based excision. From the side
the distal margin is obliquely truncate, the extreme distal angle appearing consider-
ably above the hooked termination of the ninth segment. Intermediate appendages
unusual in construction, apparently single-jointed, with a broad base, lateral outer
angle produced in a very long, sinuous spine. Penis from above broad and rectan-
gular, upper surface with the apex excised, lower surface projecting beyond it ;
imbedded in the membranous apex is a sinuous spine arising form a broad base.
Other structures of the penis, possibly a complicated form of lower penis-cover,
as shown in the figures. Penis-sheaths wanting.

Length of anterior wing, <, 275 mm.

MEXICO : Vera Cruz, Cardel, 3.viii.i932, i $ (microscope preparation).

A very unusual form of Protoptila, with no apparent penis-sheaths and a single-
jointed intermediate appendage.

Culoptila gen. n.
(Figs. 64-66)

This genus has so far only been recorded from Mexico. Its most striking feature
is the development of the tegulae, which are enlarged in the male from mere plates
to huge caps, containing at their bases concertina-shaped structures which probably
function as scent-organs. Whilst such structures are generally extensile or ever-
sible, I believe that in Culoptila the organ is fixed as in the shape shown in the
figure. The material at my disposal is plentiful and in the large number of examples
passed through KOH, there is no sign of any expansion or extension of the structures.
As may be seen in the figures, the edges of the folds are armed with bristles. Culoptila
may also be distinguished from Protoptila and other genera in the group by the
neuration and characters of the genitalia.

In the anterior wing, forks nos. i, 2, 3 and 4 are present, fork no. 4 being absent
in Protoptila and other allied genera. As with these allied genera, there is a short
row of stiff bristles arising from the posterior edge of Cu 2 , which is considerably
thickened. In the posterior wing the neuration is variable and possibly even
occasionally aberrant, as is shown in the figures of C. aluca. Ocelli present. Spurs
o, 4, 4. There is always a prolongation of the upper distal margin of the ninth seg-
ment covering a pair of in-turned hooks of varying shape. There is a large, shield-
shaped upper penis-cover. The penis is very obscure ; there are, in every species,
two black spines of unequal length and the presence of an apparent duct at the base
of one of them suggests that these may be penis and sheath. The structure which

THE PROTOPTILA GROUP OF THE GLOSSOSOM ATINAE

337

I have doubtfully termed the lower penis-cover is in all species bifurcate, and in
cleared examples is seen, in the side view, to be connected far back in the fifth seg-
ment with the upper penis-cover. The ventral process of the sixth sternite is
peculiar in shape in all species and from the side is seen as a rounded or oval lobe
attached to the sternite by a short stem.

FIGS. 64-66. 64, Head and thorax of Culoptila aluca, <$, dorsal ; 65, tegula of C. aluca,
showing scent-organ ; 66, head and thorax of Allotrichia pallicornis Eaton.

Type species, Culoptila aluca sp. n.

Culoptila aluca sp. n.

(Figs. 67-69)

Insect brownish, neuration as figured.

GENITALIA, $. Ninth dorsal segment produced. Beyond it there is a large
dorsal plate which, from above, entirely obscures the genitalia. Arising from its
underside, near the base, is a down-turned process terminating in a bifurcate apex,
outer fork long and down-curved, inner rounded, armed at its apex with a few bristles.

338

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

There is a very large, shield-shaped upper penis-cover and a pair of asymmetric
spines, the longer of which may be the penis, the shorter its sheath. At the centre
of the margin of the eighth sternite is a process which may be the lower penis-cover,
with outer wings and a central plate bearing, on each side, a blunt projection of its
apical margin, armed with a pair of bristles. Inferior appendages (?) short, slightly
curved.

Length of anterior wing, $, 2 mm.

MEXICO : Guerrera, Cocula, 28.111.1935, Tierra Colorada, i8.xii.i929 ; Michoacan,
Caracuaro, 1197 m., 24-29.^, 3-5^.1935, Nocupetaro, 9^.1935 ; Huetamo, 8.vii.

1935-

Type, $ (microscope preparation), Caracuaro, 24-29.^.1935.

67a

67

FIGS. 67 67a. Culoptila aluca, <$. Wings ; 6ja, apex of posterior wing of another example.

Culoptila amberia sp. n.

(Figs. 70-72)

Anterior wing as figured, both posterior wings too incomplete for description.

GENITALIA, <. Ninth segment extending downwards only slightly below the
pleurite of the eighth. Dorsal plate produced, apex truncate, directed downward
in side view ; from its underside arises a pair of stout, in-curving hooks with their
sub-acute apices nearly meeting. Upper penis-cover from beneath wide at the base,
becoming somewhat abruptly sub-acute at the apex, the formation of the structure
obscure ; from the side it is broad, with a truncate apex and there are indications
that it may consist of two structures superimposed. Penis and sheath in the form
of two blackened spines, the longer being probably the penis. Inferior appendages
invisible except in cleared material ; they lie above what I suppose to be the lower
penis-cover and are short and sinuous, curving slightly outward as seen from beneath.
Lower penis-cover consisting of a pair of large arms separated from each other by
a wide, rounded excision, each apex armed with one or two stout bristles.

Length of anterior wing, <$, 2-75 mm.

MEXICO: Liquidamber, 19.111.1931, i < (microscope preparation).

THE PROTOPTILA GROUP OF THE CxLOSSOSOMATINAE

339

FIGS. 68-69. Culoptila aluca, <$. 68, Genitalia, lateral ; 69, genitalia, ventral.

34<>

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

70

FIG. 70. Culoptila amberia, . Anterior wing.

72

FIGS. 71-72. Culoptila amberia, <. 71, Genitalia, lateral ; 72, genitalia, ventral.

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE 341

Culoptila rusia sp. n.

(Figs. 73-74)

Insect brownish, neuration as in C. aluca.

GENITALIA, ^. Dorsal plate much produced and very narrow at the sub-acute
apex, which is directed slightly downward ; at the base, on the underside, are two
large, in-curving hooks or processes with blackened apices, sinuous from the side.

74

FIGS. 73-74. Culoptila rusia, . 73, Genitalia, lateral ; 74, genitalia, ventral.

Upper penis-cover shield-shaped, apex sub-acute, from the side with the lower
margin produced downward in a large triangular flap, apex produced and acute.
Penis and sheath in the form of two blackened spines, the shorter being probably

342 THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

the penis. Inferior appendages short, curving downward. Lower penis-cover
with a pair of widely divergent, lateral arms and a quadrate plate dimly seen beneath
it.

Length of anterior wing, $, 2 mm.

MEXICO : La Prusia, 1075 m., 25.1^.1931 ; Belisario, Dominguez (San Geronimo),
Chiapas, 6. v. 1935. (The place is situated near the swift-running Huixtla River,
with cataracts and waterfalls.)

Type, $ (microscope preparation), La Prusia, 25.1^.1931.

Culoptila saltena sp. n.

(Figs. 75-76)

Neuration as in C. aluca.

GENITALIA, <. Ninth segment produced dorsally, slightly shallower than the
eighth, lower distal margin terminating in a rounded knob as seen from the side.
There is a dorsal plate which, from beneath, is seen to bear on the under surface
two in-turned claws enclosing an oval space. Apical margin of the plate arises from
a narrowish stem, broadened towards the distal, truncate margin, of which the upper
angle is produced in a sharp spur, lower forming an inwardly directed hook. Upper
penis-cover shield-shaped as in C. aluca. There are two black, asymmetric spines,
of which the longer is probably the penis. Inferior appendages short and curved.
Lower penis-cover broad, with divergent, narrow, triangular, lateral arms, apices
notched from the side ; behind it a quadrate plate whose excised distal margin
appears between the arms.

Length of anterior wing, <$, 2mm.

MEXICO: Huixtla, 21. xi. 1930; Saltenango de la Paz, 15.111.1931; Morelos,
Cuernovaca, 1542 m., southern slopes of the Central Tableland, many brooks in
deep-cut valleys, 8-n.vi.i932.

Type, $ (microscope preparation), Huixtla, 2i.xi.i930.

Mexitrichia meralda sp. n.

(Figs. 77-79)

Insect very similar in general appearance and neuration to M. leroda Mosely.

GENITALIA, $. To the margin of the ninth dorsal segment is jointed a large plate,
probably fused superior appendages, with a rounded excision of the distal margin,
the outer angles produced, blackened and incurved like a pair of horns. From the
side the plate is wide at the base, with a sinuous lower margin, the upper being
straight until it approaches the apex, where it is abruptly bent downward ; the
apex of the plate terminates in a blackened, distally and slightly downwardly
directed beak. Upper penis-cover long, spine-like, straight from above, curving
slightly upwards from the side, with a slightly thickened base. Penis broad and
fleshy, not exserted in the type as is shown in leroda; the two terminal black spines,
so conspicuous in leroda, are doubtfully shown in meralda and appear to be directed

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

343

basally. Sheaths long, somewhat ribbon-shaped. Lower penis-cover from below
with two outstanding wings as in leroda, and a very long, twisted spine arising from
the centre and extending beyond the penis ; the surface is covered throughout its
length with a few small teeth.

76

FIGS. 75-76. Culoptila saltena, <J. 75, Genitalia, lateral ; 76, genitalia, ventral.

Length of anterior wing, <$, 3 mm.

MEXICO: Huixtla, 21. xi. 1930; Esmeralda, 545 m., 19. xi. 1930; Morelos,

344

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

7, 2i.iv.i932; Montecristo de Guerrero, n. 111.1931; Michoacan, Caracuaro,

25- iv. 1935-

Type, <$ (microscope preparation), Huixtla, 21. xi. 1930.

FIGS. 77-79. Mexitrichia meralda, <$. 77, Genitalia, lateral ; 78, genitalia, dorsal

79, genitalia, ventral.

THE PROTOPTILA GROUP OF THE GLOSSOSOMATINAE

345

Mexitrichia rancura sp. n.

(Figs. 80-86)

Insect brown. In the neuration there are slight differences between rancura
and the type species leroda as shown in the figure.

GENITALIA, $. Dorsal margin of the ninth segment widely excised as in leroda.
Beyond it is jointed a large plate, apical margin above produced at its centre in an
acute triangle, sides bent downward, with acute, blackened apices which, from the
side, appear as a beak-like hook directed downward ; from this aspect, the produced
centre of the plate appears as a large, distally directed beak. Upper penis-cover long,
from the side sinuous, from above broadened before the apex. Penis very broad,
roughly oblong from the side, from beneath strongly constricted at the middle,

80
FIG. 80. Mexitrichia rancura, <J. Wings.

tapering from a broad base ; arising towards its apex is a pair of laterally outstretched,
short, curved horns, densely covered with minute setae. There are two sinuous
and strongly chitinized spines, shaped like staples, back to back on each side of
the penis. These may be penis-sheaths ; their attachment to the penis and lower
penis-cover is very obscure. Lower penis-cover with outstretched wings as in the
other species of the genus, but with no apparent central spine in the single example
available.

Length of anterior wing, $, 4 mm.

MEXICO: Barranca Honda, 24.11.1931, i $ (microscope preparation).

In the figures of the species, I have included certain structures apparently connected
with the penis and the lower penis-cover. These structures are very obscure as
to their attachments. They appear also in meralda, but as in this species they are
even more obscure, I have omitted to figure them or to refer to them in the descrip-
tion of either species. I believe that homologous structures, equally obscure, occur
also in the genera Protoptila and Culoptila. It is perhaps beyond the scope of this
purely taxonomic paper to deal with these anatomical features.

346

THE PROTOPTILA GROUP OF THE GLOSSOSOMATIN AE

86

^^^^

FIGS. 81-86. Mexitrichia rancura, 3. 81, Genitalia, lateral ; 82, genitalia, dorsal ; 83,
genitalia, ventral ; 84, lateral portion of dorsal plate, ventral ; 85, penis with its upper
and lower covers, lateral ; 86, penis with upper and lower covers, ventral.

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

24 NOV 1954

A REVIEW OF THE

AEDES SCUTELLARIS SUBGROUP

WITH A STUDY OF VARIATION IN

AEDES PSEUDOSCUTELLARIS

(THEOBALD)

(DIPTERA : CULICIDAE)

ELIZABETH N. MARKS

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 3 No. 10

LONDON: 1954

A REVIEW OF THE AEDES SCUTELLARIS

SUBGROUP WITH A STUDY OF VARIATION IN

AEDES PSEUDOSCUTELLARIS (THEOBALD)

(DIPTERA : CULICIDAE)

BY

ELIZABETH N. MARKS

YvA

(Department of Entomology, Queensland University) ^

Pp. 347-414 ; Plate 18 ; 19 Text-figures ; i Map

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 3 No. 10

LONDON: 1954

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

Parts 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. 3, No. 10 of the Entomological
series.

PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM

Issued November 1954 Price Eighteen Shillings

A REVIEW OF THE AEDES SCUTELLARIS

SUBGROUP WITH A STUDY OF VARIATION

IN AEDES PSEUDOSCUTELLARIS (THEOBALD).

(DIPTERA : CULICIDAE)

By ELIZABETH N. MARKS M.Sc. (Qux), PH. D. (CANTAB.), F.R.E.S.

(Department of Entomology, Queensland University)

CONTENTS

PAGE

I. INTRODUCTION .......... 350

II. REVIEW OF THE scutellaris SUBGROUP . . . . . 351

(1) General survey . . . . . . . . 351

(2) Taxonomic characters ....... 357

(a) General ......... 357

(b) Head 358

(c) Thorax ......... 358

(d) Legs 360

(e) Abdomen ........ 361

(/) Male genitalia ........ 362

(3) Zoogeography . .... 374

(4) Plate and key for identification of adults .... 379

(5) Interspecific relationships ....... 383

(6) Immature stages ........ 386

III. EXPERIMENTAL STUDIES OF VARIATION IN Aed.es pseudoscutellaris

(THEOBALD) ......... 387

(1) Introductory ......... 387

(2) Material ' 388

(3) Methods 389

(a) Rearing ......... 389

(b) Sampling ......... 390

(c) Examination of specimens ...... 391

(d) Variation of larval environment ..... 392

(4) Results 395

(a) Tables III and IV : Length of wing and of hind femur . 395

(b) Table V : Pale scales under the proboscis . . . 396

(c) Table VI : Scutal pattern ..... 397

(d) Table VII : White scales on either side of the prescutellar

bare area ........ 398

(e) Table VIII : White basal patches on fore and mid tarsi . 399
(/) Tables IX and X : Extent of white anterior streak and

ventral dark line on the hind femur . . . 400

(g) Tables XI-X VIII : The bands on the hind tarsal segments 401
(h) Table XIX : Distance between lateral patches on tergites

V and VI ........ 407

(i) Table XX : Specialized setae on the basal lobe . . 408
ENTOM. Ill, 10. 23

350 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

PAGE

(5) Summary of experimental results ...... 409

(6) Conclusions ......... 410

IV. ACKNOWLEDGMENTS . . . . . . . . .410

V. BIBLIOGRAPHY . . . . . . . . . .411

SYNOPSIS

Eighteen described and three undescribed members of the Aedes scute llaris subgroup are
recognized from the Australasian region and eastern part of the Oriental region. The systematic
status accorded to members is reviewed, the diagnostic specific characters critically examined,
and the geographical distribution of the subgroup illustrated and discussed.

An original pictorial chart for identification of members of the subgroup indicates also the
geographical distribution of various taxonomic characters, the implications of which are con-
sidered. The general trend is from west to east but one character shows a north-south distri-
bution. A key to adults is provided for use with the chart, and the inadequate state of present
knowledge of immature stages is briefly indicated.

Source and experimental methods of rearing a colony of Aedes pseudoscutellaris Theobald are
described. Results are tabulated of a biometrical study of twenty-eight characters, in five
series of pseudoscutellaris adults bred from larvae whose environment had been subjected to
controlled variation of either temperature or salinity, and in one series of adults from different
stock, also bred in controlled conditions. The records are analyzed and their implications
discussed.

The results support the specific status accorded to members of the scutellaris subgroup, in
that the observed range of experimental variation is not generally of the same magnitude as
interspecific differences in the same character. Lack of variation in the basal lobe of the male
coxite is evidence of its value in defining species. Two characters should probably be used
with caution for delimiting species, since pseudoscutellaris can exhibit both extremes of their
development. A complete interruption to the white bands on hind tarsal segments II-IV
can be produced by low temperatures of larval environment. Presence or absence of a white
streak under the proboscis may be due to differences in hereditary constitution, though larval
environment also has some effect.

I. INTRODUCTION

IN recent years, many closely related mosquitoes which were, or would have been,
regarded by earlier workers as varieties or subspecies, have been given full specific
status. This is partly due to more adequate material being available for study,
but in certain well-investigated groups, biological studies have suggested or proved
the distinctness of the forms concerned, where morphological differences are slight
or obscure. At the same time, cross-breding experiments have shown a few forms
to be no more than subspecies. The classical example is the Anopheles maculipennis
complex in Europe ; others are the Anopheles maculipennis complex in North
America and the Anopheles gambiae complex in Africa. Culicine mosquitoes have
not received the same attention as Anophelines, but the Culex pipiens complex is
now being widely studied.

All the foregoing groups of mosquitoes have a continental distribution. Where
the members of a complex are geographic races, largely confined to different island
groups, there might appear to be a strong case for regarding some, at least, of them
as subspecies. Such a complex is the Aedes scutellaris subgroup, widely distributed
in the Australasian region and the eastern part of the Oriental region. Of the

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 351

eighteen described members of the subgroup, seventeen have been treated by recent
authors as full species ; cross-breeding experiments have involved only four members,
one of which, as a result, is regarded as a subspecies. There is a little evidence
indicating biological differences between sympatric members of the subgroup, but
no intensive research on the subject has been made. In many cases quite small
morphological differences are used to separate the species.

Huxley (1942) described variation as "... a study of the differences between
organisms. On analysis, these differences may turn out to be due to differences in
environment ... or ... to differences in hereditary constitution ... or ... to a
simultaneous difference both in environment and in constitution . . . The important
fact is that only experiment can decide between the two." Simpson (1944) pointed
out that "... no morphological character is inherited as such. What is inherited
is a complex of potentialities for development, and the ultimate morphological
expression of the same hereditary characters may differ markedly."

It appeared that some light might be shed on the systematic status of members
of the scutellaris subgroup by a study of the amount of variation that could be
induced in one member by controlled variation of the environment. Comparison
of specimens from different stocks of the same species, reared in identical conditions,
might indicate differences due to hereditary constitution. At the same time, the
reliability of certain taxonomic characters could be tested. In order to evaluate
the results of the experimental work, it was first necessary to review the scutellaris
subgroup as a whole, and to examine the characters in use for distinguishing the
species.

In addition to laboratory-reared material, these studies are based on the examina-
tion of numerous specimens in the collections of the British Museum (Natural
History), London School of Hygiene and Tropical Medicine, Bernice P. Bishop
Museum, Honolulu (E. C. Zimmerman collection), and University of Queensland,
and on notes on specimens in the U.S. National Museum made by Dr. Alan Stone.

II. REVIEW OF THE SCUTELLARIS SUBGROUP

(i) General Survey

The subgenus Stegomyia Theobald of genus Aedes Meigen is practically confined
to the tropical and subtropical regions of the old world, chiefly the Ethiopian,
Oriental and Australasian regions. Aedes (Stegomyia) aegypti Linn, has spread to
the new world by man's agency.

A group of species within the subgenus Stegomyia is known as the scutellaris
group. Its exact limits have been differently interpreted.

Edwards (1932) divided the subgenus Stegomyia into four groups. In what he
termed " Group C (scutellaris group) " he included ten species from the Oriental
and Australasian regions, Crete and Africa. Three of these were known from the
Australasian region, A. (S.) albolineatus (Theobald), A. (S.) albopictus (Skuse) and
A. (S.) scutellaris (Walker) (of which he listed five varieties).

The albolineatus-complex, removed from Group C by Knight & Rozeboom (1946),
was named Group E (albolineatus group) by Knight & Hurlbut (1949). The latter

352 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

authors divided Group C, thus modified, into three subgroups, viz. : Subgroup I,
scutellaris s.str. Subgroup II, albopictus, and Subgroup III, mediopunctatus. The
term " the scutellaris group " as used by Earner & Bohart (1945) and more recent
authors has, in fact, referred to practically the same complex of species as Knight
& Hurlbut's Subgroup I. 1

The scutellaris subgroup of the present paper is synonymous with Knight &
Hurlbut's Subgroup I, which the authors define as follows : " Characterized by
having the more mesal portions of the abdominal tergal markings sub-basal. In
addition, post-spiracular scales are lacking, the scutal longitudinal median line is
relatively slender, and the pleural scale patches are arranged in two rather well-
defined longitudinal bands (not true of gurneyi, however.)"

The following list of the species included by Knight & Hurlbut under their defini-
tion, with the addition of one species and one subspecies since described is arranged
in chronological order of the date of publication of the names (as either varietal or
specific) . The original designation is given in parenthesis.

1. scutellaris Walker, 1859 (Culex scutellaris).

2. pseudoscutellaris 2 Theobald, 1910 (Stegomyia pseudoscutellaris) .

3. tongae Edwards, 1926 (Aedes variegatus var. tongae).

4. andrewsi Edwards, 1926 (Aedes variegatus var. andrewsi).

5. alorensis Bonne- Wepster & Brug, 1932 (Aedes (Stegomyia} variegatus var.

alorensis} .

6. horrescens Edwards, 1935 (Aedes (Stegomyia} scutellaris var. horrescens}.

7. gurneyi Stone & R. Bohart, 1944 (Aedes (Stegomyia} gurneyi}.

8. marshallensis Stone & R. Bohart, 1944 (Aedes (Stegomyia} marshallensis).

9. guamensis Earner & R. Bohart, 1944 (Aedes (Stegomyia} guamensis}.

10. pernotatus Earner & R. Bohart, 1944 (Aedes (Stegomyia} pernotatus}.

11. quasiscutellaris Earner & R. Bohart, 1944 (Aedes (Stegomyia} quasiscutel-

laris).

12. hensilli Earner, 1945 (Aedes (Stegomyia) hensilli}.

13. paullusi Stone & Earner, 1945 (Aedes (Stegomyia} paullusi).

14. riversi R. Bohart & Ingram, 1946 (Aedes (Stegomyia) riversi}.

15. scutoscriptus R. Bohart & Ingram, 1946 (Aedes (Stegomyia} scutoscriptus}.

16. hakanssoni Knight & Hurlbut, 1949 (Aedes (Stegomyia} hakanssoni).

17. scutellaris katherinensis Woodhill, 1949 (Aedes scutellaris katherinensis} .

18. polynesiensis Marks, 1951 (Aedes (Stegomyia) polynesiensis} .

1 A . (S.) gurneyi is often placed with albopictus ; scutoscriptus is also aberrant. Stone & Earner
(1945) found A. (S.) galloisi Yamada difficult to place, from the description, and omitted it from their
key ; Knight & Hurlbut (1949) place it in Subgroup II, albopictus.

2 Marks (1951) demonstrated that two species had previously been confused under the name " pseudo-
scutellaris." In this paper " pseudoscutellaris " is used for authors' references which do not discriminate
between the two forms. Where the identity of the form is beyond doubt it is referred to under the
appropriate name pseudoscutellaris or polynesiensis.

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 353

In addition there are three other forms known but not named. 1 The form of
A. (S.) scutellaris described from Andaman Is. by Barraud (1928, 1934) is recognized
by recent authors as probably distinct, and from my own observations of specimens
I feel satisfied that this is so. Edwards (1929) noted a form from Rotuma I., north
of Fiji, which differed in certain aspects from " pseudoscutellaris." I have seen this
also and it appears to represent a distinct race. Both these forms await review
when more adequate material is available. Bohart & Ingram (19466) gave some
details of "Aedes sp. in scutellaris group " from Palau Group, Caroline Islands, and
Dr. Alan Stone (in lift.) considers it is a valid species. He has sent me additional
particulars of it which confirm his finding.

The scutellaris subgroup is equivalent to a " species group " of Zeuner (1943),
or to a " superspecies " of Mayr (1942), if the latter concept were modified, as seems
reasonable, to include groups of species which are mainly allopatric but in which
the ranges of a few species overlap. 2

Mattingly (1953) considers that A. (S.) granti (Theobald), which Knight & Hurlbut
(1949) placed in Group C, Subgroup II albopictus, belongs to the scutellaris sub-
group, which it resembles in pleural markings. Mattingly uses the presence of an
extra stripe of pale scales between the dorsal border of the sternopleuron and the
lower edge of the posterior pronotum to link granti with hakanssoni and scutoscriptus ,
and to distinguish them from " more typical members of the group." However, a
similar stripe may occur in scutellaris, in pseudoscutellaris and, reduced in length,
in polynesiensis. It is most likely to be seen in large unrubbed specimens and might
well be found in other species if suitable series were available. Descriptions indicate
that granti differs from all the species here included in the scutellaris subgroup in
markings of proboscis, female palps, wings, legs, scutellum and abdominal tergites
(which have the white bands basal medially). While some species of the subgroup
do not conform in certain characters with the general pattern, there is an overall
strong indication of close relationship. In my opinion, inclusion of granti would
destroy the homogeneity of the subgroup, which could no longer be regarded as
representing a superspecies, whereas, if granti were placed by itself in a separate
subgroup of Group C, the affinities with the scutellaris subgroup indicated by the
pleural pattern would not be obscured.

The full synonymy of the various species of the scutellaris subgroup, first dealt
with by Edwards (1917), has been fully covered in recent papers on the group (1944
onwards), and finally cleared up by the examination of a topotypic male of Aedes
scutellaris (Walker), by Stone (1947). In references to earlier papers, species are
treated here under the names by which they are at present recognized.

Table I records the treatment which species described before 1940 have received

1 Since this paper went to press, Dr. R. M. Bohart has informed me of a fourth undescribed species
from Korror I., in the Palau Group, and has very kindly supplied details which will enable it to be
distinguished from others in the subgroup. Descriptions of the two species referred to here as Palau
sp. and Korror sp. will be published by Dr. Bohart at an early date. In addition Mr. Mattingly informs
me that he has recently examined a specimen of the scutellaris subgroup from the Maldive Is., a description
of which is in the press. In the majority of characters, including the basal lobe of the male coxite, it
closely resembles scutellaris scutellaris but it differs in having white scales on the under side of the
proboscis.

1 The concept has recently been so modified (Mayr et al., 1953, P- 2 9)-

354

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

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A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

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356 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

from those authors responsible for changes in nomenclature or systematic status.
Forms described since 1940 have suffered no change in the status ascribed to them
by their original authors.

The first recognition of the scutellaris subgroup as such (i.e., that A. scutellaris
as then recognized comprised a complex of closely related forms was by Edwards
(1926), prompted by Buxton & Hopkins who had observed constant differences
between the anal gills of larvae from Samoa and from the New Hebrides. He found
that " there are at least five distinct varieties distinguishable by small differences
of colour and also by the male hypopygium, especially in the form of the basal lobe
of the sidepiece. The characters are fairly well defined, but are perhaps best treated
as varietal rather than specific, especially as their significance appears to be mainly
geographical."

Barraud (1928), Edwards (1929) and Bonne- Wepster & Brug (1932) gave details
of forms from Andaman Is., Rotuma I. and Taroena (= paullusi) respectively which
differed from those already described but did not give them varietal names ; the
last authors, however, described a sixth variety, alorensis.

Following Edwards, varietal status was ascribed to the named forms until Knight,
Bohart & Bohart (1944) listed them as subspecies, without comment on the change.

Farner & Bohart (1945) on the basis of male genitalia differences, regarded the
known Australasian members of the scutellaris subgroup as separate and distinct
species occupying similar ecologic niches. The occurrence of more than one from
the same area lent support to their contention.

Farner (1945) observed, in describing hensilli, that it was possibly a subspecies
of guamensis but until further material was available and in view of the differences
in tarsal and abdominal banding it seemed best to regard it as specifically distinct
from guamensis.

Since 1944, most authors have described new forms as full species without comment
on their status. The exception is scutellaris katherinensis to which Woodhill (1949^)
gave subspecific rank on the grounds of its ability to hybridize with scutellaris scutel-
laris.

The description in recent years of many new forms in the scutellaris subgroup
reflects an increase in the quantity and quality of collections, and the examination
of males from localities whence only females were known before. Additional
characters have been brought into use for identifying the different species but it
cannot be said that the recognition of the various forms as full species is due to
the discovery of new and more significant characters. They are still distinguished
from one another chiefly by differences previously in use for separating those forms
recognized as varieties. It is the significance placed on these characters by taxono-
mists that has altered.

The treatment of the different forms (except s. katherinensis) as full species is
now general, and will probably remain so unless it can be proved incorrect. Not
all culicidologists are convinced that this treatment is in all cases justified, and where
forms replace one another geographically it may yet be possible to demonstrate
that some are, in fact, only subspecies. Morphological differences between some
species appear to be no greater than those between the subspecies s. scutellaris and

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 357

s. katherinensis. However, to quote Mayr (1942), " Considerably more material
must be examined in order to recognize subspecies, than is needed for the description
of good species."

It may be, on the other hand, that, with further knowledge of geographical varia-
tion, what is now recognized as a single monotypic species will be broken up into
two or more subspecies. Bohart & Ingram (19466), for instance, found that speci-
mens of hensilli from Ulithi had the last tarsal segment usually about half white,
whereas in Truk specimens it varied from two-thirds white to all white ; there was
also a greater tendency for complete abdominal bands in Truk females. There
may be some slight differences between tongae from Tonga and from Sikiana in
degree of development of specialized setae on the basal lobe of the male coxite.

I have seen a series of eight specimens of s. scutellaris from Admiralty Is., two of
which are normal, four have a line of white scales on the anterolateral margin of the
scutum, one has hind tarsal V with a dark patch at the tip, and one has hind tarsal
bands reduced in width and segment V dark on the apical half. A male similar to
the last and a normal male were reared from the same larval collection from Aneityum
I., New Hebrides, by Dr. Marshall Laird. Male genitalia of the three types appear
identical. These variations have not been recorded from elsewhere in the range of
s. scutellaris. Male genitalia of the three types appear identical.

The few hybridization experiments recorded, viz. : s. scutellaris X s. katherinensis
(Woodhill, 19490), s. scutellaris and s. katherinensis X pseudoscutellaris 1 (Woodhill,
1950), and s. scutellaris X pernotatus (Perry, 1950), have only served to confirm the
status already ascribed to the forms concerned. 2 Smith- White (1950) discussed the
genetical significance of non-reciprocal fertility between s. scutellaris and s. katherin-
ensis and suggested further lines of investigation.

(2) Taxonomic Characters
(a) General

Doleschall (1858) and Walker (1859, 1861) made no comparison with other species
in their descriptions, which all apply to s. scutellaris.

Theobald (1901, 1903, 1907, 19100) included specimens of both albopictus and
members of the scutellaris subgroup in his concept of scutellaris. When (19106) he
described pseudoscutellaris he distinguished it by the characters in which it differs
from what he called " scutellaris ", but we now know to have been albopictus, i.e.,
by the typical scutellaris subgroup pleural markings of white lines. He also observed
the curving white lateral patches on the abdominal tergites. Edwards (1917)
included several forms of the scutellaris subgroup, among them pseudoscutellaris, in
one monotypic species and it was not until 1926 that he realized they represented a
complex of distinct forms.

1 I have confirmed by examination of specimens the identity of Woodhill's colony as pseudoscutellaris
and not polynesiensis.

2 Woodhill (1954) has crossed pseudoscutellaris from Fiji with polynesiensis from Tahiti and obtained
small numbers of fertile hybrids, showing that the two species are not genetically isolated, though they
do not mate readily in laboratory conditions. This suggests that polynesiensis may be a subspecies of
pseudoscutellaris. However, repetition of the experiment using Fijian polynesiensis and a study of the
habits of the two forms, where they occur together in nature, would appear desirable in order to jtidge
whether there are barriers to their natural hybridization.

358 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

The characters discussed here include all that have been found of importance by
Edwards and later authors in identification of specimens of the scutellaris subgroup
and some which have been suggested but are not in general use. A consideration
of their occurrence and variability supplements the information given in Plate I.
They do not, however, include all characters which may be noted in descriptions.
The first author to use a character is indicated. A name is put in inverted commas
where it is obvious that an author was including more than one form under it.

Colour differences have been noted in some cases. Not enough is known about
factors causing differences in scale colour, but generally in mosquito taxonomy,
such small differences as between white and creamy white or yellowish are of doubtful
value.

(6) Head

Bonne-Wepster & Brug (1932) noted in the median white stripe on the vertex,
a difference in the width towards the nape between " scutellaris " and alorensis ;
this character is not in general use.

The same authors observed that Taroena specimens of " scutellaris " (= paullusi)
had a distinct white line under the proboscis ; this feature is now known to occur in
a number of species, sometimes in the male only ; in some cases it appears to be very
variable.

Stone & Farner (1945) noted the white stripe in paullusi males, but only a few
pale scales under the female proboscis ; I have seen a male of this species from
Samar I. which lacks the white stripe. The following species have a white stripe :
riversi <$, $ ; quasiscutellaris $, $ ; tongae <$, $ (Stone & Farner (1945) observed
this in <$, it is also present in all $$ I have seen) ; horrescens $ ($ with some pale
scales) ; pernotatus $, $ (but I have seen pernotatus <$<$ in which it was lacking).
It is present also but has not previously been noted in Andamans sp. <$, $, and
alorensis $ ; and I have observed that there may be some pale scales under the pro-
boscis in andrewsi $, scutoscriptus (Dr. Alan Stone informs me (in lift.) that scuto-
scriptus has a faint line of yellowish brown scales on the underside of the proboscis)
and Rotuma sp. <$. Bohart & Ingram (19466) noted that the proboscis of " pseudo-
scutellaris " $<$ sometimes has some pale scales beneath ; this was also observed
by Marks (19510) in " pseudoscutellaris " (= polynesiensis) <$<$ and $$. Both sexes
of pseudoscutellaris may have proboscis with a white stripe beneath, or entirely
dark ; variation in this is shown in Table V.

Stone & Bohart (1944) found that marshallensis had the white markings on the
male and female palps considerably reduced ; this is the only species thus distin-
guished.

(c) Thorax

Scutum. Edwards (1926) noted that the scales of the median stripe might be
creamy rather than white. Bonne-Wepster & Brug observed that the stripe in
alorensis was narrower than in " scutellaris," and Edwards (1935) that in horrescens
it was broader than in "pseudoscutellaris " but no measurements have been made

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 359

from comparable series of different species. Environmental conditions can affect
this character in pseudoscutellaris. 1 The median line often continues into a denfiite
or faint fork on either side of the prescutellar bare area : Bonne- Wepster & Brag
(1932) noted some whitish scales in this position in " scutellaris," a distinct fork
occurs in hakanssoni and scutoscriptus , and I have observed it also in Andaman Is.,
form and in alorensis ; Woodhill (19490) found no fork in the great majority of
specimens of s. katherinensis ; most authors refer to it only in general terms. Varia-
tion of the amount of white scaling in this position in pseudoscutellaris is shown in
Table VII. It may be a character of which more use could be made.

Bonne- Wepster & Brug (1932) observed a narrow white line on the anterolateral
margin of the scutum in Taroena specimens of " scutellaris " (= paullusi) ; Stone
& Farner (1945) observed a similar line of fine yellowish scales in quasiscutellaris
(not noted in the original description); scutoscriptus has a broad line in this position ;
a narrow line occurs in hakanssoni, pseudoscutellaris, and in some specimens of s.
scutellaris from Admiralty Is. In all except probably scutoscriptus these lines are
variable and may be incomplete, particularly in females. In pseudoscutellaris they
may be reduced to patches of scales on the scutal angles. Species which usually
have the scutal angle dark scaled may sometimes have two or three pale scales in
this position though this has not been recorded in descriptions. I have observed
this in polynesiensis, andrewsi, alorensis, guamensis, horrescens and Rotuma sp.
The number of pale scales on the scutal angle is important in distinguishing between
specimens of pseudoscutellaris and polynesiensis and would possibly prove of use in
other cases if details were available.

The white scales above the wing root are usually all broad. In hakanssoni this
is not specifically stated in the description and, in fact, I have observed that they
are mainly narrow-curved. In scutoscriptus there are some narrow-curved scales in
addition to the broad ones noted in the description. It is possible that they have
been overlooked in other species also, and it is a character that is worth further
investigation.

Scutellum.

The characteristic scaling of the scutellum in the scutellaris subgroup is white,
with a small patch of black scales at the apex of the midlobe. Knight & Hurlbut
(1949) found that hakanssoni has much more extensive black scaling. Gurneyi is
described on differences from albopictus and by inference has all lobes entirely
white scaled, but Dr. Stone informs me (in litt.) that there are a few dark scales at
the tip of the midlobe in unrubbed specimens.

Pleuron.
Stone & Bohart (1944) observed that the white scales on the pleura of gurneyi

1 Mr. P. F. Mattingly drew my attention to a character which Edwards found useful in separating
African species of Stegomyia, viz., the presence of broad scales medially on the anterior margin of the
scutum. A cursory examination of the experimental series of pseudoscutellaris showed that these scales
might be present or absent and suggested that they are affected by environment though this was not
thoroughly investigated. The character therefore does not appear a promising one for use in the
scutellaris subgroup.

3 6o A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

are arranged in patches rather than the straight lines characteristic of other members
of the scutellaris subgroup (in this it resembles albopictus}. The description of
scutoscriptus indicates that only the upper pleural line is complete in this species.

Specimens of some species have the upper and lower white patches on the mesepi-
meron fused, in others they are separate ; usually no note of this is given. I have
observed both conditions in pseudoscutellaris and it does not appear a promising
diagnostic character, though it would need further investigation to be certain.

Wings. These are dark scaled in all species. Wing length is often given as a
measure of size but not as a diagnostic character. In some species a white spot
occurs at the base of the costa ; it has possibly been overlooked in others ; it is a
practically constant character in pseudoscutdlaris. The ratio of the length of the
fork cells to their stems, a character used in several culicine groups, is omitted
from most descriptions.

Halteres. The scaling of the halteres is not recorded for all species, but in most
the knob appears to be mainly dark scaled, whereas in scutoscriptus Farner & Bohart
(1946) observed it to be entirely pale scaled and in hakanssoni it is largely pale.

(d) Legs

Fore femur. Stone & Bohart (1944) noted the absence of a white apical spot
on the fore femur of marshallensis but it apparently occurs in all other species.

Mid femur. Bonne- Wepster & Brug (1937), who had a mixed collection of
" scutellaris," observed that sometimes the mid femur had a thin longitudinal white
stripe on the outer (= anterior) side. Stone & Farner (1945) noted this as one oi
the diagnostic characters of paullusi ; Woodhill (19490) found that it was a constant
character by which s. katherinensis could be distinguished from s. scutellaris. I have
found that a similar white stripe occurs in alorensis ; this had previously been over-
looked. Of five adults of andrewsi examined, one female had an anterior line on the
basal two-thirds of one femur ; possibly this was an aberrant specimen, or else in
this species the character may be variable.

Fore and mid tibiae in some species have scattered pale scales posteriorly, but
this has not been used as a diagnostic character.

Fore and mid tarsi. Most species have white patches at the base of segments I
and II on fore and mid tarsi. Edwards (1935) noted that horrescens had white
scales more frequently on fore and mid tarsal III than did " pseudoscutellaris " ;
Farner & Bohart (1944) found that males of guamensis had the fore tarsus entirely
dark and distinguished pernotatus by the presence of basal patches on III and
sometimes IV and V ; quasiscutellaris is similar to horrescens. I have observed
that in andrewsi only fore and mid tarsal segment I has a basal white patch. The
development of this character in pseudoscutellaris is shown in Table VIII.

Hind femur. Edwards (1926) noted differences in the size of the white spot at
the tip of the hind femur, but this is not a useful character. Bohart '& Ingram
(19466) drew attention to two distinct types of pattern in the marking of the anterior
surface of the hind femur ; there is a longitudinal pale area which either forms a
tapered white line (in most species), or trails off ventrally towards the tip rather

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 361

than tapering to a sharp point (in guamensis, 1 scutoscriptus , hakanssoni and sp. from
Palau, also in some specimens of marshallensis (see p. 382)).

I have observed an apparent difference between pseudoscutellaris and polynesiensis
in the distance from the base of the femur to the beginning of the dark ventral
scaling, but the character needs further investigation.

Hind tarsi. The white bands on the hind tarsi are one of the most important
distinctive characters, and are usually measured in terms of the ratio of the length
of the band to the total length of the segments ; most species have V entirely white.
Edwards (1926) observed differences in the widths of the bands and gave ratios
for segment IV. Bonne- Wepster & Brug (1932) observed that in " scutellaris "
the band on I was incomplete (i.e., interrupted by dark scales beneath) and gave
ratios for the bands on II-IV, noting that Taroena specimens (= paullusi) had wider
bands on IV. Stone & Bohart (1944) found marshallensis had V dark on the apical
half or more ; hakanssoni is similar ; in hensilli the character shows geographical
variation, Ulithi specimens have the apical half dark, whereas in Truk specimens
not more than one- third is dark and the segment may be all white. Two specimens
of s. scutellaris (one from Admiralty Is. ; one from Aneityum I., New Hebrides) have
been seen with apical half of V dark and bands on other segments reduced.

Farner & Bohart (1944) observed that guamensis had the white bands completely
interrupted by dark scales on the inner side. Stone & Farner (1945) noted that
andrewsi 2 had the bands on IV interrupted on the dorsal surface, but this appears
to be variable and I have seen two females with the band quite complete. The
majority of species have the band on I interrupted beneath though it is recorded
as complete in a few and not noted in others. It was complete in specimens of
alorensis and Andamans sp. examined. In some species the extent to which the
remaining bands are interrupted has been found to be quite variable. Tongae was
previously recorded as having complete bands on II-IV, but I have seen one specimen
from Tonga with I-V, another with IV and V, and one from Sikiana with IV
completely interrupted. Complete bands are also recorded for pernotatus but a
specimen from Aneityum I., New Hebrides, has the bands on I-IV completely
interrupted beneath. Variation in this character in pseudoscutellaris is shown in
Tables XIII-XVIII. Farner (1945) gave measurements of the inner as well as
the outer lengths of the white bands in hensilli. Stone & Farner (1945) reversed
the usual convention by giving the width of the dark band on segment IV at its
widest dimension, but no other authors have followed this formula.

(e) Abdomen

The characteristic markings of the abdominal tergites in the scutellaris subgroup
are lunate lateral white patches curving away from the base of the segment into
forwardly directed hooks which may extend dorsally to form a broken or complete
sub-basal band across the segment. The characteristic markings of a species
usually show best on tergites IV-VII but may be found on II and III in some.

1 Not noted in the original description.

2 Stone & Farner said they had not examined specimens of andrewsi and it was included in their key
on the basis of the original description ; however, the latter does not mention this character.

362 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

Edwards (1926) observed the degree of completeness of these bands on the different
segments of the abdomen ; this continues to be one of the principal diagnostic
characters employed, but in some species is rather variable. He noted the distance
the bands were removed from the bases of the tergites ; Bonne- Wepster & Brug
(1932) also used this character, and Knight & Hull (1952) found differences in it
between paullusi and s. scutellaris. It is possible that if long series of other species
were examined and the measurement could be made from the actual base of the
tergite, and not from the apex of the preceding one, distinctive differences might
be found. In practice, with dried specimens which have been in various conditions
and ages when killed and have shrunk to different extents, the character has not
been proved a generally useful one, the more so as not all authors have noted it
when describing species. Edwards (1926) also noted whether the lateral patches
were creamy or white and (1935) on what segments they were visible dorsally ;
the latter character is rather unreliable. Farner & Bohart (1944) observed sub-basal
lateral patches on guamensis which are not of the typical lunate shape, but rather
triangular. There is possibly some variation in the shape of the lateral patches in
other species, but no long enough series have been examined to be sure of this.

Edwards (1926) recorded the scale pattern of the sternites, whether those of the
basal segments were all white, or had apical black bands ; the latter is the more
common condition. This is not one of the principal diagnostic characters but
might be used to supplement them in some cases. It may prove rather variable
and in dried specimens the sternites are often difficult to see. Some $$ of s. katherin-
ensis which have narrow sub-basal white bands on sternites V and VI and a submedial
band on VII appear distinct from s. scutellaris which usually has wide basal bands on
V and VI, and often only small lateral patches on VII ; other specimens are much
more like s. scutellaris.

(/) Male genitalia

Edwards (1926) first recognized that members of the scutellaris subgroup (" races
of Aedes variegatus ") could be distinguished " by the male hypopygium, especially
in the form of the basal lobe 1 of the sidepiece." The parts that have been used in
distinguishing members of the scutellaris subgroup include the ninth tergite, coxite,
style and basal lobe of coxite. The complete genitalia of pseudoscutellaris are
illustrated in Fig. i. There are various terminologies in use for the descriptioi
of the parts of the male genitalia. That followed here is given by Edwards (1941),
and all except direct quotations from other authors have been translated to thes
terms. The male genitalia of the mosquito rotate 180 during the first 24 houi

1 The term basal lobe is in general use for this structure in subgenus Siegomyia ; it bears no resemblance
to, for instance, the basal lobe in subgenus Ochlerotatus. In the scutellaris subgroup, its anatomical
position indicates that it is homologous with the harpago (claspette of Edwards) of Ochlerotatus and
Finlaya, though it lacks the terminal appendage present in those subgenera. Edwards (194*) i n hi
general description of mosquito genitalia describes the basal lobe as a modification of the coxite on the
area between the ventral root and the midsternal connection of the coxites. " In some genera part or
the whole of this basal lobe has more the form of a subsidiary appendage, which has been called the
claspette." In his description of subgenus Stegomyia he says " claspettes in the form of a hairy basal
lobe or plaque attached to sternal side of coxite, this plaque in a few species with the inner part forming
a small process bearing modified bristles." This is in contrast to his earlier (i93 2 ) description of the
subgenus "... basal lobe (plaque) present and hairy ; no claspettes."

A KEVIEW OF THE AEDES SCUTELLARIS SUBGROUP

363

Fig. i. (J Genitalia of Aedes pseudoscutellaris (Theobald), complete, tergal view (E. 7).

ENTOM. Ill, 10.

26

364

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

after emergence. Thus the 8th and gth tergites become ventral, the 8th and gth
sternites dorsal. The ventral aspect referred to in the literature is therefore tergal.
The left coxite, of which the basal lobe is usually illustrated, is that on the left side
of the insect after rotation and therefore on the left side of figures of whole mounts

Fig. 2. <J genitalia of Aedes pseudoscutellaris (Theobald) bisected, inner lateral view (H. 33).

of genitalia which are conventionally drawn from the tergal aspect with the distal
portions towards the top of the page. In this paper the view from the mid line of
the genitalia is referred to as " inner lateral view " (illustrated in Fig. 2), and that
of the side of the basal lobe nearest the coxite as " outer lateral view."

Certain of the setae of the basal lobe may be enlarged, thickened or flattened
and arise from tubercles ; these are quite distinct in appearance from the remainder

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

365

1 \ 1 1 '

\ l l'i \

\M'i\

\\\

FIGS. 3 and 4. Left basal lobe of $ coxite of Aedes pseudoscutellaris (Theobald). Fig. 3.
Tergal view (B .7). Fig. 4. Sternal view (B. 7).

366

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

and are here referred to as specialized setae. Other authors have given them various
designations (not always very clearly denned, but usually interpretable by reference
to accompanying figures). The term does not include the very long hairs at the

FIGS. 5 and 6. Left basal lobe of coxite of Aedes pseudoscutellaris (Theobald). Fig. 5.
Inner lateral view (B. i). Fig. 6. Outer lateral view (B. i).

tip of the lobe in some species which are thickened and elongated by reason of their
size but do not differ in form from the majority these are the " thickened bristles "
of some authors. In several species a scale has occasionally been observed on the
basal lobe (Fig. 7) ; it appears to be an aberration of no taxonomic significance.

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

367

Edwards (1926) found differences in the length of the coxite compared with its
breadth at the base ; this ratio has not been recorded for most of the recently
described species. Though the character by itself is not diagnostic it might be
advantageous to have some note of it as it would give a basis of comparison where,
as in many cases, the only part of the genitalia illustrated is the basal lobe.

8

FIGS. 7 and 8. Left basal lobe of <$ coxite of Aedes polynesiensis Marks. Fig. 7. Tergal
view (the scale is aberrant). Fig. 8. Sternal view. <$, Taveuni, Fiji.

Edwards also observed differences in the style, whether slender or moderately
stout, with tip swollen to varying degrees, and with its spine slender or stout, blunt
or pointed. Such differences of degree are not very useful diagnostic characters, in

368

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

particular if other forms are not available for comparison. The appearance of
the style depends to some extent on the way the specimen has been mounted. If
the ratio of length of style to length of coxite were recorded, differences might be
found between species. The spine varies even more with the mounting, as it is

FIGS. 9 and 10. Left basal lobe (? coxite of Aedes polynesiensis Marks. Fig. 9. Inner
lateral view. Fig. 10. Outer lateral view. <J, Suva, Fiji

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

369

liable to shrink in diameter, and I have seen it appear quite different in two specimens
of the one species.

Another character used by Edwards (1926) was the form of the ninth tergite
whether convex or emarginate it is emarginate in quasiscutellaris, convex in most
other species in which it has been described. The sclerotized portion of the tergite
is narrow and very apt to become twisted in a mounted preparation, so that its shape
is not always easy to interpret.

FIGS, ii, 12. Left basal lobe of $ coxite of sp. in scutellaris group from Andaman Is.
Fig. ii. Inner lateral view. Fig. 12. Outer lateral view.

The character which Edwards observed as most important, the basal lobe, is
now often the only part of the genitalia described or figured, since it has been found
the most distinctive of all diagnostic characters used in separating the species of
the scutellaris subgroup.

Edwards noted that the basal lobe differed in shape, the tip rounded, square-ended
or conical ; also in the length and disposition of the setae at the tip. He did not
mention the occurrence on the basal lobe in any forms of stouter specialized setae,
though his figure of andrewsi suggests they were observed in this species. The figures
suggest (as do Edwards' preparation of genitalia of British Museum specimens)
that the genitalia were mounted undissected ; they were drawn in tergal view. He
distinguished the basal lobe of tongae from that of " pseudoscutellaris " by the fact

37

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

that the former was hairy at the tip only and the latter hairy on the apical half or
more. Edwards' figure shows the setae extending half-way down the tergal aspect
of the basal lobe of " pseudoscutellaris," though in fact they occur on the sternal
aspect. The specimen figured was from Samoa (= polynesiensis) .

Barraud (1928) was the first to record the presence of specialized setae on the basal
lobe. He observed in Andaman Is. form that a few of the setae at the apex of the

13

14

FIGS. 13, 14.

Left basal lobe of $ coxite of Aedes andrewsi Edwards from Christmas I.
Fig. 13. Inner lateral view. Fig. 14. Outer lateral view.

basal lobe are " stouter and more blade-like than the majority." Bonne-Wepster
& Brug (1932) recognized the distinctive form of the basal lobe in alorensis and the
presence of " a peculiar claspette-like, long and slender structure with two filaments
at tip." (In my opinion really part of the basal lobe bearing specialized setae (see
figure in Plate 18).

Farner & Bohart (1944) described and figured " enlarged bristles " (i.e., specia-
lized setae) on the basal lobes of pernotatus and scutellaris. They made an important
contribution to the methods of investigating the subgroup by the introduction of
figures of the basal lobe in lateral as well as in tergal view. (They figured lateral

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 371

views for scutellaris and guamensis, but the latter from comparison with specimens
is more a tergal than truly lateral view.) They also stated " the thickened bristles
of the basal lobe in guamensis are less developed than those in pseudoscutellaris 1 ."
This might have been interpreted as a reference to specialized setae in which case
the specimens of "pseudoscutellaris" would be true pseudoscutellaris. However,
their illustration is of polynesiensis and Dr. Alan Stone (in litt.) informs me that the
setae of the basal lobe in the latter are all slightly heavier than in guamensis.

Farner & Bohart (1945) referred to quasiscutellaris as having " a row of somewhat
thickened bristles " and " no specialized setal group." They described the basal
lobe of tongae as " without a specialized group of stout setae ; large bristles confined

FIGS. 15, 16. Left basal lobe of $ coxite of Aedes tongue Edwards from Sikiana, Solomon
Is. Fig. 15. Inner lateral view. Fig. 16. Outer lateral view.

to apex, particularly in ventral view." They had seen one male from Vavau, Tonga
Is., and figured the lobe in dorsal and ventral view. I have examined the basal
lobe of a male of tongae from Sikiana, Solomon Is. in lateral view and it has 9-10
specialized setae (Fig. 15). It is not easy to judge with specimens mounted in
tergal view but another Sikiana male appears to have 6 or 7. Three males from
Tonga have 5 to 7 specialized setae which appear less strongly developed and more
tapered than in those from Sikiana.

The same authors distinguished between " simple " (e. g., tongae) and " complex "
(e.g., quasiscutellaris) basal lobes and noted whether the lobe is covered with minute
setae tergally.

1 Cf. Bohart & Ingram (19466) who describe the basal lobe of " pseudoscutellaris " (= polynesiensis)
as " largely covered with bristles, none of which appear as thickened setae."

372

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

Farner (1946) described the basal lobe of hensilli as similar to guamensis " but
apparently with thickened bristles extending more basad." Bohart and Ingram
(1946) distinguished two types of arrangement of specialized setae : " in a clump "
in scutellaris as compared with " in a row " in hensilli.

The distinction is not a satisfactory one since an inner lateral view of the basal
lobe of s. scutellaris shows the specialized setae in a row though in other views they
often appear clumped. The basal lobe of s. scutellaris is square-ended in inner

17

FIGS. 17, 18. Left basal lobe of $ coxite of Aedes guamensis Farner & R. Bohart from
Rota, Marianas. Fig. 17. Inner lateral view. Fig. 18. Outer lateral view.

lateral view and the specialized setae are grouped on the sternal angle of the tip
(this occurs also in andrewsi) whereas in hensilli and species with similar basal lobes
the tip is more rounded and the row of specialized setae extends down the sternal
aspect of the lobe. Woodhill (19490;) described the genitalia of s. katherinensis as
indistinguishable from those of s. scutellaris, " the basal lobe of the coxite carrying
a series of hairs at the apex with several longer hairs joined to form a spine. The
degree of development of this spine varies in both subspecies . . . ." I have examined

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 373

specimens of s. katherinensis from Woodhill's colony. The " spine " is the group
of specialized setae which, as in s. scutellaris, often appear clumped, but in inner
lateral view they are seen as a row of about six setae. From this aspect the lobe
differs somewhat in shape from s. scutellaris ; the tip is more conical without a distinct
angle sternally, and the row of specialized setae extends down the sternal aspect.

Knight and Hurlbut (1949) figured only the tergal view of the genitalia of hakan-
ssoni and described it as " quite similar to river si and hensilli " which presumably
means that specialized setae are present though this is not indicated in figure or
text. I have examined a paratype of this species, and, though difficult to make out,
there do appear to be 3 or 4 broader setae in a row among the non-specialized ones.

It was the observation of constantly-occurring specialized setae on the basal lobe
of the coxite of specimens from the laboratory colony at Cambridge which first drew
attention to the distinctness of this form (pseudoscutellaris) from the widely distri-
buted Polynesian form (polynesiensis) usually referred to when the name " pseudo-
scutellaris " was used.

There is usually one aspect of the basal lobe which shows the characteristic shape
of the lobe, and the specialized setae to best advantage.

In species with complex basal lobes, which appear expanded in tergal view, such
as quasiscutellaris, pernotatus, horrescens and alorensis, this aspect shows all that is
needed for identification. In a species such as paullusi with a simple truncate lobe,
the setae all visible tergally and none of them specialized, the same view is again
sufficient. Most of the species with basal lobe simple or expanded in lateral view
have only a few long setae at the tip visible tergally. Thus Fig. i which shows this
aspect of pseudoscutellaris is an equally good illustration of polynesiensis and tongae,
and shows nothing except in proportionate lengths of the parts of the genitalia,
to distinguish it from Rotuma sp. and possibly also from hensilli, hakanssoni, river si,
guamensis and andrewsi. On these same basal lobes, the unspecialized setae may
occur only towards the apex or may extend varying distances towards the base
if the latter they may do so on the sternal aspect of the lobe, as in polynesiensis
(when they are adequately illustrated from an inner lateral or, if there are no
specialized setae, a sternal view), or more towards the outer lateral aspect as in
pseudoscutellaris.

Practice has varied in illustrating the lateral aspect of the basal lobe. Thus
figures of the lateral view of s. scutellaris in Farner & Bohart (1944, 1945), Bohart
& Ingram (19466) and Knight & Hull (1952) and of s. katherinensis in Woodhill
(i949#) all appear to be of the outer lateral aspect ; whereas those of hensilli and
riversi in Bohart and Ingram (19466) appear to be of the inner lateral aspect. Where
specialized setae occur in a row, the illustration must show them in file to display
their number and shape ; this, in all species examined, is best seen from the inner
lateral view. I have drawn the outer lateral aspect for a number of species and find
that it shows quite distinctive arrangements of the unspecialized setae which would
be valuable adjuncts to the characters of the specialized setae. Figs. 3-18 demon-
strate the necessity for dissection of the genitalia in forms with a simple basal lobe,
and for illustrations of more than one aspect of it, in order to show all the distinctive
features.

374 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

Where the basal lobe is simple it might be useful to record its length relative to
the length of the coxite, taking them from a given level (e.g., the outer lateral angle
at the base of the coxite). This ratio appears to differ between pseudoscutellaris,
polynesiensis and tongae on the one hand and Rotuma sp. on the other, though one
would need to see more than the single available specimen of the latter to be satisfied
on this point.

A few authors have recorded the number of apical teeth on the phallosome but
there is insufficient information to suggest whether there may be specific differences
in this character.

(3) Zoogeography

Mosquitoes of the scutellaris subgroup are found in the eastern part of the Oriental
region and in almost all parts of the Australasian region that lie within the tropics,
though their distribution on the tropical portion of the Australian mainland is
apparently rather limited. They range from the Andaman Is. in the west, north-
east to Okinawa and south-east to Mangareva Is. and Pitcairn I., i.e., from approxi-
mately 93 E. to 130 W. and from 26 N. to 25 S. 1

The present known distribution of the various members of the subgroup is shown
in the accompanying map. 2

In addition to well-established records, a distinction has been made on the map
between definite records of the scutellaris subgroup, where it is the identity of the
species that is in doubt, and comprehensive records which include a whole island
group within the range of a species, though the localities given for it cover only a
small portion of that range. Where the range of two species overlap, the known
limits of the overlap are shown.

Most species of the scutellaris subgroup are confined to islands or groups of islands
and are plainly examples of the process of speciation by geographical isolation.
The origins of the insect faunas of Pacific islands have been discussed by, amongst
others, Buxton (1935) and Zimmerman (1948). They are derived mainly from
the west and are largely Indo-Malayan in origin. Farner & Bohart (1945) con-
sidered that superficially the scutellaris subgroup " appears to be of Polynesian or
Melanesian origin, having spread westward into the insular parts of the Oriental
region " but they acknowledged that with further knowledge of the subgroup, it
might be necessary to modify or discard this suggestion. There seems no reason
to suppose that members of this subgroup have spread naturally in a different
direction from other mosquitoes and other insects. Moreover, they include the
easternmost representatives of the subgenus Stegomyia, which extends throughout
the Oriental and Ethiopian regions, and this in itself is strong evidence that the
subgroup originated somewhere in the western part of its range, where also it overlaps
the range of the closely allied albopictus subgroup.

1 The specimen of scutellaris subgroup from Maldive Is., reported by Mattingly (see footnote, p. 353)
extends the range westward to approximately 74 E.

2 A copy of an earlier version of this map appears in Manson-Bahr & Muggleton (1952). The original
contained no reference to Anopheles punctulatus farauti, and the indication in the published map, that
its southern boundary of distribution is coincident with that of s. scutellaris, is incorrect except in
New Hebrides.

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376 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

Though many of the species of the scutellaris subgroup are isolated from their
fellows, there are several cases where the ranges of two or three overlap. If the
distribution is treated in terms of island groups, the overlap appears considerable,
but when details of locality records are examined, it is found that in some cases,
as far as at present known, it is of small extent.

Apart from the records of recognized forms Andamans sp., andrewsi from
Christmas I., alorensis from Alor and s. scutellaris from Ceram, Aroe Is. amd Moluccas
there are numerous records of " scutellaris " from islands of the Malay Archipelago
which have not been reassessed since the existence of many distinct species in the
scutellaris subgroup was recognized. Brug (1932), Bonne- Wepster & Brug (1932,
1937) and Brug & Bonne-Wepster (1947) record mosquitoes of the scutellaris sub-
group from Sumatra, Java, Lesser Paternoster Is., Celebes, Boeton, Kaboena,
Adonare, Tenimber, Boeroe, Sanana, Saparoea and Sumba.

Brug & Bonne-Wepster (1947) also record paullusi from Celebes, Ceram, Amboina,
Sanana and Sumatra. Since it has now been found that alorensis has an anterior
white line on the mid femur, it seems possible that some of these records may refer
to alorensis. For this reason, until further information is available, these localities
have not been included in the range of paullusi on the map, but remain in the area
of species of unconfirmed identity. Paullusi is known with certainty from Sangir
and the southern half of Philippine Is. ; in the latter it overlaps the range of s.
scutellaris which occurs throughout the group. To the north the subgroup extends
to Okinawa, where one species, riversi, is found.

The Caroline Islands stretch from east to west over 2,000 miles and five species 1
are known from them s. scutellaris, hensilli, scutoscriptus, hakanssoni and sp.
from Palau group. In the latter islands, at the western end of the chain, s. scutellaris
is the only species recorded from Pulo Anna and Angaur ; on Peleliu and Garakayo
both hensilli and " sp. in scutellaris group " have been found. To the north-east
hensilli is the only species known from Ulithi Atoll, and still further east it is found
on five islands of Truk Atoll, on four of which scutoscriptus also occurs. At the
eastern end of the Carolines chain, hakanssoni is the only species of the group found
on Ponape. It is a reasonable supposition that hensilli was formerly confined to
Ulithi and perhaps other islands as yet uncollected, and has subsequently extended
its range both east and west to overlap in each direction that of another form.
North of Caroline Is., a single species, guamensis, is known from Mariana Is., and
to the east another, marshallensis , from Marshall and Gilbert Is.

The Philippines and Carolines records of s. scutellaris represent the north-west
limits of its range. It extends south through New Guinea and I have seen specimens
from Hammond Is., Torres Straits, within about 20 miles of Cape York. In the
eastern islands of Torres Straits it occurs on Murray, Darnley, Nepean and Yorke
Is. The subspecies s. katherinensis is known from Northern Territory and northern
West Australia. The situation in north Queensland is interesting and requires
elucidation. Mackerras (1946) mentioned the occurrence of scutellaris in unsettled
parts of Cape York Peninsula adjacent to Thursday I. ; his specimens have not been
available for examination. Of two females from the Red Island Point area near

1 Dr. R. M. Bohart has a sixth species, from Korror I. in the Palau group.

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 377

Cape York, one is apparently s. scutellaris with mid femora entirely dark anteriorly ;
the other has three white scales scattered along the anterior surface of its remaining
mid femur. Of five females from the Coen district, about 200 miles south of Cape
York, two from Skull Creek are badly damaged ; the only remaining mid femur
has three or four white scales scattered along its anterior surface. There are seven
to fourteen white scales in this position in specimens from Coen and from Musgrave
Station. The fifth specimen from Blue Mountains goldfield, has distinct but incom-
plete narrow lines of white scales on the anterior surface of its mid femora. In this
it resembles one of Woodhill's s. scutellaris x s. katherinensis F 1 hybrids, rather
than s. katherinensis. It must be remembered that s. katherinensis is known chiefly
from specimens descended from one batch of eggs from Katherine, N.T., only six
other specimens from two localities being recorded. It might be possible for the
white line to vary to this extent in different stocks (cf., variation in white scaling
on the proboscis of different stocks of pseudoscutellaris shown in Table V). On the
other hand, Cape York Peninsula is the most likely place for a naturally interbreeding
population of the two subspecies to occur. Whether or not this specimen is a hybrid,
it indicates that s. katherinensis does occur on the Peninsula.

The north-east limits of s. scutellaris are Admiralty Is. and Bismarck Archi-
pelago. As already noted, some Admiralty Is. specimens have atypical scutal or
tarsal markings. Laird (1946) records s. scutellaris from New Britain, but Hill's
(1925) record of " variegatus " from New Ireland has not been re-checked. Laird
(1952) found quasiscutellaris on Nissan I., half-way between New Ireland and
Solomons.

In the south-east s. scutellaris extends to New Hebrides where it occurs on
numerous islands and overlaps the range of the local species, pernotatus. A possible
physiological difference between s. scutellaris from New Guinea and New Hebrides
is suggested by its incrimination as the vector of jungle dengue fever in New Guinea
(Mackerras, 1946), whereas in New Hebrides Perry (1948) considered that it did
not appear to be a vector of dengue in nature under normal conditions (though
Daggy (1944) thought it a probable vector on epidemiological grounds). North of
New Hebrides, Farner & Bohart (1945) quoted identifications of s. scutellaris from
Rennell 1 and Bellona Is. in south-west Solomons, and of s. scutellaris and tongae
from Nupani I. in Santa Cruz group. They considered these records needed confir-
mation by further collections.

The islands have been doubtfully included in the range of these species on the
map.

In Solomon Is. both quasiscutellaris and the aberrant species, gurneyi are found ;
the latter has been recorded only from Bougainville and Guadalcanal. Tongae has
been collected on Sikiana an outlying island east of the Solomons, but is the only
species known both from there and from Tonga.

In the eastern part of the subgroup's range there are several species, females of

1 The identification of further specimens from these islands would be of particular interest. Professor
G. D. Hale-Carpenter informs me (in litt.) that in its butterflies of the genus Euploea, Rennell differs
considerably from other islands of the Solomons group, and it seems that it got some at least of its fauna
directly from New Guinea's eastern tip via the Louisiade archipelago as stepping stones.

378 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

which are very difficult to distinguish from one another. Pseudoscutellaris and
polynesiensis have until recently been treated together as " pseudoscutellaris " 1
and Rotuma sp. females 2 may be indistinguishable from polynesiensis, as also are
some females of horrescens. It has been possible to check many of the records by
examination of males and polynesiensis appears to be the only widely distributed
species in this area. Specimens collected by Dr. Marshall Laird on Nukonono I.
have confirmed that polynesiensis occurs in the Tokelau group. Until males have
been seen, some doubt must be felt in particular about records from Ellice Is. and
Wallis I., which are nearest to the ranges of other members of the subgroup.

Theobald's (1907) record of " scutellaris " from Pitcairn I. seems to have been
overlooked by later authors. Though Theobald at that time included both albo-
pictus and scutellaris in his concept of " scutellaris," it is very unlikely that this
record referred to albopictus and highly probable that it was polynesiensis.

In Fiji three species occur ; polynesiensis is here at the western limit of its range
but pseudoscutellaris and horrescens are not known from elsewhere. Edwards (1935)
suggested that horrescens might be the native Fiji form, and that " pseudoscutellaris "
had been more recently introduced from Samoa. While it is likely that the wide-
ranging polynesiensis is an introduction, not enough is known of the distribution
of pseudoscutellaris to cast doubt on its endemicity. Though the most obvious
explanation is that these sympatric species originally developed in geographical
isolation, comparatively little is known of differences in their ecology which might
affect speciation. 3 Horrescens does show differences in habits from "pseudo-
scutellaris," but now that the latter name is known to have covered two species,
the subject needs re-examination.

The question arises why, when most of the species of the scutellaris subgroup
have, as far as we know, a fairly restricted distribution, a few should have much
more extensive ranges. The larval ecology of all is essentially similar, why should
some species apparently be much more biologically aggressive than others ?

A study of the literature suggests that there may be some correlation between
a species' ability to extend its range and a combination of avidity for human blood
and ability to colonize artificial containers. The only forms for which both latter
characteristics are recorded are s. scutellaris, "pseudoscutellaris " (= polynesiensis) ,
hensilli, quasiscutellaris and riversi. With the exception of the last two, each of
these species overlaps in its range at least two other members of the subgroup.
Table II gives a summary of the recorded habits of the species. 4

The inference to be drawn is that the activities of man have had much to do with
the spread of these species and also that almost certainly they originally developed

1 Taylor's (1914) and Breinl's (1915) records of " pseudoscutellaris " from Papua can be assumed to
refer to s. scutellaris.

2 In one ? from Rotuma the ratio of the length of the white band on hind tarsal segment IV was
0-86 ; in one $, 0-79. This is longer than is usual in pseudoscutellaris and polynesiensis, and might
prove a distinguishing character.

3 Zimmerman (1948), as a result of his observations of insects on Pacific islands, considers that certain
organisms appear to have selected certain environmental conditions. Rather than being what they are
as the result of environmental influences, certain mutant forms seek out the niches best suited to them.

4 Dr. R. M. Bohart informs me that Korror sp. breeds in pitcher plants and has a very distinctive
larva.

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 379

in geographic isolation somewhere within their present range. Buxton and Hopkins
(1927) have put forward a strong case for the distribution of " pseudoscutellaris "
(= polynesiensis) through the islands of Polynesia by the agency of Polynesian
canoes. They similarly explain the occurrence of tongae on Sikiana ; little has
been recorded of the habits of tongae. Marks (1950) found that a percentage of
pseudoscutellaris eggs could survive up to three weeks immersion in sea water,
suggesting the possibility of dispersal in small floating vessels, such as coconut
husks ; however, it is now doubtful whether pseudoscutellaris is widely distributed.
The fact that certain forms of the scutellaris subgroup are known from the same
localities without apparent intergrades justifies their treatment by taxonomists
as full species, but further investigation is needed of forms which replace one another
geographically. Huxley (1940) has emphasized the necessity for detailed mapping
of the boundaries and range changes of species and subspecies. Many more records
are needed before this can be attempted for the scutellaris subgroup.

(4) Plate and Key for Identification of Adults

Plate 1 8 includes the twenty-one known forms in the scutellaris subgroup 1 and,
as indicated, eighteen have been actually examined, though in two of these male
genitalia were not seen. The data included are based on a study of all descriptions
and figures available, supplemented by direct observations on specimens, and by
notes on gurneyi, marshallensis, scutoscriptus , hakanssoni and sp. from Palau group,
kindly made by Dr. Alan Stone from specimens in U.S. National Museum.

The previous discussion of the diagnostic characters of the scutellaris subgroup
indicates the variability of these in some species. Each figure therefore represents
what, as far as can be judged from the information available, would be the condition
in an average specimen of that series. 2 In several cases where characters vary
considerably and both extremes are well represented, this has been shown by dupli-
cation of the parts concerned ; in the case of pseudoscutellaris some of the variation
obtained experimentally has been indicated thus.

All drawings, except those of the basal lobe, are conventional. For example, no
attempt has been made to indicate the distance of the white band from the base of
the tergite. The lengths of the tarsal segments are in their correct proportions,
but the width has been exaggerated. The width of the white bands on the tarsal
segments is the outer (i.e., greatest) width ; where the bands are interrupted by
dark scales this is shown in the same figure though in the actual specimens the
interruption (except in andrewsi) is on the inner side.

The descriptions from which the figures are drawn vary in detail. Hensilli and
s. katherinensis are fully described. Marshallensis is described on the basis of

1 Dr. R. M. Bohart has supplied the following details for Korror sp. of characters figured in Plate i.

Proboscis : no white scales on under side in female, a somewhat distinct line in male. Scutum : no
white scales along anterolateral margin. Scutellum : silver scales on all lobes, black at apex of mid
lobe. Mid femur: no anterior white line. Hind femur: anterior white marking sloping off ventrally,
not tapering. Hind tarsus : white bands complete on II-V, covering III, IV, V. Tergite V :
band usually, but not always, incomplete. Basal lobe : somewhat as in riversi but row of specialized
setae in apical rather than subapical position.

* The faint line of yellowish brown scales under the proboscis of scutoscriptus has not been interpreted
as a white streak.

ENTOM. III. 10. 27

380

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

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A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 381

differences from s. scutellaris and gurneyi on the basis of differences from albopictus,
but Dr. Stone has cleared up doubtful points for me. He has also provided a descrip-
tion of the characters illustrated for sp. from Palau group, of which only brief details
were given by Bohart and Ingram (19466) . The illustration of the basal lobe of
this species is copied from a sketch supplied by Dr. Stone ; he describes the lobe as
follows : " The apex is somewhat angular, almost as in marshallensis, but lacking
the two stout spines ; the hairs of the apex of the lobe (mediad) are thickened and
curved and form a very distinct curved tuft." Bohart and Ingram's notes on the
hind tarsus differ from Dr. Stone's, so both have been illustrated.

The differences in the figures of the basal lobe are probably in some cases more
apparent than real and in others more real than apparent since they are from various
sources. The drawing of the basal lobe of scutoscriptus is enlarged from a small
figure of the complete genitalia and possibly suggests a closer resemblance to perno-
tatus and alorensis than actually exists. Dr. Stone informs me (in litt.) that an
examination of the type slide of riversi shows the basal lobe to be more as in s.
katherinensis than would appear from the figure in the chart.

Some species show the distinctive shape and specialized setae of the basal lobe
best in tergal view. In others an inner lateral view is essential for correct inter-
pretation. It is believed that the figures are drawn to meet these requirements.
Unfortunately, sp. from Rotuma does not show the specialized setae in tergal view,
and since a second male was not available from which a mount in lateral aspect
could be made, the basal lobe is figured in sternal view ; this figure is therefore
not directly comparable with any of the others.

It was thought proper to show a distinction between freehand and scale drawings.
The former were made during visits to the British Museum (Natural History) and
though they are believed to give an accurate representation, it was not possible to
give the same time and care to details as in specimens which were examined in the
laboratory in Cambridge or Brisbane ; the latter were drawn to scale.

The species are arranged in Plate 18 as far as possible in their geographical relation-

* No information is available on the habits in nature of spp. from Andaman Is., or Rotuma I., nor of
andrewsi, alorensis, pseudoscutellaris or s. katherinensis ; laboratory colonies of the last two feed readily
on man. Bahr (1912) had both polynesiensis and pseudoscutellaris amongst the mosquitoes used when he
proved "pseudoscutellaris " the vector of filariasis in Fiji (see Marks, 1951).

f Bohart & Ingram (19466) report that adults of guamensis " are rarely seen in nature and have never
been observed to bite man even in heavily breeding areas ". Reeves & Rudnick (1951) record 21 speci-
mens of guamensis in a collection of 7767 mosquitoes taken biting man. Farner & Bohart (1945) quote
Fullaway's report in 1912 of this species as " very abundant and troublesome in the forests " but it seems
unlikely that this referred to guamensis. Laboratory experiments of Travis (1947) showed that guamensis
would not feed readily on a dog but that in 68% of those which did, larvae of Dirofilaria immitis developed
to the infective stage.

J Manson-Bahr & Muggleton (1952) found that adults of horrescens reared in the laboratory fed reluc-
tantly on human blood and concluded from their experiments that this species was unlikely to be a
suitable host for the local non-periodic filaria.

Buxton & Hopkins' (1927) records of breeding places of " pseudoscutellaris " refer to polynesiensis,
the only species of the subgroup known from Samoa. Paine (1943) gives a long list of breeding places
of " pseudoscutellaris " in Fiji but these may refer either to pseudoscutellaris or to polynesiensis. Perry
(1950) quotes a personal communication from Lever reporting that " pseudoscutellaris " was an important
agent in the transmission of dengue in Fiji this might apply to either or both pseudoscutellaris or
polynesiensis.

If Backhouse & Heydon (1950) found scutellaris inhospitable to Wuchereria bancrofti in the Rabaul
region.

382 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

ships, so that those next to one on the plate are those nearest to it on the map.
This was more difficult with species in the eastern part of the range of the group,
and for interpretation of apparent trends in characters, the plate should be used in
conjunction with the map.

It is hoped that the plate may overcome certain disadvantages of keys and tables.
The visual presentation directs attention to resemblances and differences and the
geographical arrangement brings out trends in characters and suggests relationships,
some of them quite unexpected, which might otherwise escape notice. If an unde-
scribed form were encountered, the plate should indicate the fact more strongly
than a key can. Like keys and tables, however, the plate is not more than a pre-
liminary guide and final identifications should always be made by reference to full
published descriptions.

Though the plate was designed for use on its own, it might prove more useful in
the field if a key were available or a preliminary guide. The following key has
therefore been prepared. It gives a good indication of the necessity for examination
of male genitalia for certain identification of a species. The plate obviates the need
for a key to male genitalia.

KEY TO ADULTS OF THE scutellaris SUBGROUP

(Median white line on scutum relatively slender ; hind tarsal segments I-IV
basally banded, V basally banded or all white.)

1 . Abdominal tergal markings basal ; pleural markings in patches . albopictus subgroup.
Abdominal tergal markings with more mesal portions basal ; pleural markings in

longitudinal bands ; costa with white line on basal third .... granti.

Abdominal tergal markings with more mesal portions sub-basal ; costa all dark or

with small basal white patch ..... scutellaris subgroup 2.

2. Pleural markings in patches ......... gurneyi.

Pleural markings in longitudinal bands . . . . . . . . 3.

3. Hind femur with white marking on anterior surface sloping off ventrally towards apex 1 4.
Hind femur with white marking on anterior surface tapering towards apex . . 7.

4. Abdominal tergites with triangular white lateral markings .... guamensis.
Abdominal tergites with lunate white lateral markings . . . . . 5.

5. Scutellum with patches of dark scales on all lobes ; hind tarsal V with apical half or

more completely dark 2 ........ hakanssoni.

Scutellum with small patch of dark scales at apex of mid lobe only ; hind tarsal V

white to apex on outer aspect . . . . . . . . . 6.

6. Scutum with wide white line along anterolateral margin ; abdominal tergites with

lateral patches only ......... scutoscriptus.

Scutum dark on anterolateral margin ; some abdominal tergites with complete

bands .... .... ^. ... Palau sp.

1 Since going to press I have seen four ?$ of marshallensis collected by Dr. M. Laird on Teaoraereke I.,
off Tarawa, Gilbert Is., in which the ventral dark scaling on hind femur is reduced ; in two specimens
the white marking on anterior surface still appears tapering, but in two it slopes off ventrally towards
apex; the latter would key to the second half of couplet 5, being distinguished by the dark apical half
of a hind tarsal V.

2 Dr. R. M. Bohart has supplied the following note : Korror sp. keys out to couplet 5 but has hind
tarsal V about | dark all around. Otherwise it agrees with the second half of the couplet and goes on
to couplet 6 where it differs from either part by tergites sometimes having lateral patches only and
scutum always dark on anterolateral margin.

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 383

7. Mid femur with longitudinal white line on anterior surface ..... 8.
Mid femur dark on anterior surface . . . . . . . . .10.

8. Scutum with narrow white line along anterolateral margin .... paullusi.
Scutum dark on anterolateral margin ........ 9.

9. Proboscis with white streak ventrally ........ alorensis. 1

Proboscis dark ventrally ........ scutellaris katherinensis

10. Scutum with complete or incomplete narrow white or yellowish line along antero-

lateral margin (at least, not less than five white scales) . . . . . II.
Scutum dark on anterolateral margin (at most, less than five white scales on scutal

angle) ............. 12.

11. Some abdominal tergites with complete bands ..... quasiscutellaris. 2
Abdominal tergites never with complete bands .... pseudoscutellaris.

12. Hind tarsal V with black apex ; some abdominal tergites with complete bands . 13.
Hind tarsal V white to apex on outer aspect . . . . . . .14.

13. Length of white band on hind tarsal IV, 0-2-0-4 ; on hind tarsal V, 0-2-0-5 length

of segment ........... marshallensis.

Length of white band on hind tarsal IV, 0-3-0-6 ; on hind tarsal V, 0-4-1-0 length

of segment ........... hensilli. 3

14. Some abdominal tergites with complete bands ....... 15.

Abdominal tergites with incomplete dotted bands, or with lateral patches . . 16.

15. Proboscis with white streak ventrally. ...... Andamans sp.

Proboscis dark ventrally ....... scutellaris scutellaris.

1 6. Some abdominal tergites with almost complete dotted bands ; proboscis with white

streak ventrally ............ 17.*

Abdominal tergites with lateral patches only ....... 18.

17. White scaling on one or more of hind tarsals III-V interrupted by dark scales on

inner aspect .......... tongae (in part) .

White scaling on hind tarsals III-V not interrupted river si, horrescens, tongae (in part).

18. Proboscis with white streak ventrally. ....... pernotatus. 6

Proboscis dark ventrally . . . . . . . . . . .19.

19. White band on hind tarsal IV interrupted by dark scales on outer aspect . andrewsi.
White band on hind tarsal IV not interrupted . . . . . . .20.

20. Length of white band on hind tarsal IV, not less than 0-8 length of segment Rotuma sp.
Length of white band on hind tarsal IV usually less than 0-8 length of segment

polynesiensis*

(5) Interspecific Relationships

In considering the relationships of the species it can be seen from Plate 18 that
different characters show trends in different directions. Where the affinities thus
implied conflict with one another, it is impossible to say which is indicative of closer
relationship. The tendency might be to regard the genitalia characters as of greatest
importance. However, Woodhill (1950) has shown that differences in the basal
lobe between s. scutellaris and pseudoscutellaris are not mechanical barriers to cross

1 Alorensis is known only from one $; it is probable that paullusi ?$ with reduced anterolateral
markings on scutum would be difficult to separate from alorensis $$.

2 Aberrant s. scutellaris from Admiralty Is. would key out here ; quasiscutellaris with reduced line
on scutum may key to s. scutellaris.

3 Aberrant s. scutellaris from Admiralty Is. and New Hebrides would key out here ; hensilli with hind
tarsal V completely white will key to 5. scutellaris.

4 Some specimens of s. scutellaris have dotted bands and would key to couplet 16, but have no ventral
white streak on proboscis.

5 Some specimens of horrescens and polynesiensis may key out here ; some specimens of pernotatus
will key to andrewsi but the interruption to the hind tarsal bands is on the inner aspect,

6 Some specimens of horrescens, pernotatus and andrewsi may key out here.

384 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

fertilization ; such differences therefore must be regarded from the same morpho-
logical standpoint as any other characters.

The Aedes (Finlaya] kochi group of mosquitoes has a similar distribution to that
of the scutellaris subgroup though it does not extend quite so far into the Pacific.
Marks (1947) observed that neither general coloration, nor structure of genitalia in
the kochi group showed any obvious relationship to geographical distribution, but
suggested that the affinities from New Guinea east might be traced in the larvae
since the structure of the teeth of the lateral comb did appear to be related to geo-
graphical distribution. It would be exceedingly difficult to trace affinities in the
larvae of the scutellaris subgroup, since many of them are so alike that they can
only be identified doubtfully, if at all ; adult characters are a more promising field.

The white scaling on the proboscis is frequently a variable character and the trends
it shows in distribution may be open to question. Species with a white stripe under
the proboscis extend from Andaman Is. east to Alor (alorensis) and Philippines
(pauttusi) and north of the latter to Okinawa (riversi) ; there is a gap in distribution
then as far as Solomon Is. 1 (quasiscutellaris) , to the east of which the character
appears in Sikyana and Tonga (tongae), New Hebrides (pernotatus) and Fiji (horres-
cens and sometimes pseudoscutellaris) .

White scaling on the anterolateral margin of the scutum links paullusi in the
Philippines east to the Caroline species scutoscriptus on Truk and hakanssoni on
Ponape, south to aberrant s. scutellaris in Admiralty Is. and quasiscutellaris in
Solomon Is., and east again to pseudoscutellaris in Fiji.

Since hakanssoni is the only species with distinctive scutellar scaling this character
cannot be considered.

Species with a white anterior streak on the mid femur have a north-south distri-
bution from Philippines to northern Australia, these are paullusi, alorensis and
s. katherinensis. Dr. Alan Stone informs me (in litt.} that the mid femur of the type
specimen of riversi " has a very narrow border of pale yellowish scales along the
lower margin." Most species have pale scaling along the lower posterior margin
of the mid femur and this sometimes extends on to the anterior margin. I have seen
a specimen of s. scutellaris with a distinct white border along the lower anterior
margin of the mid femur and probably the condition in riversi is comparable with this
and not with the medially placed anterior streak of paullusi.

Four species are linked by having the white anterior scaling on the hind femur
sloping off apically, instead of tapered as in the remaining seventeen forms. This
character is found only in specimens from Marianas (guamensis] and Carolines
(sp. from Palau, scutoscriptus from Truk and hakanssoni from Ponape). 2

On the hind tarsi, the absence of a dark interruption to the band on segment I
brings together species in the western part of the subgroup's range, Andamans sp.,
alorensis, s. katherinensis with gurneyi from Solomon Is. in the east, and sp. from
Palau and riversi from Okinawa in the north.

There is a reduction in the width of the white hind tarsal bands linking species in

1 The occurrence of related forms in Moluccas area and in Solomon Is. without intermediates in New
Guinea is known in Lepidoptera (Zeuner, 1943) and Odonata (Lieftinck, 1949) and has been explained
by "Zeuner (I.e.) in terms of the theory of continental drift.

2 Also in some specimens of marshallensis from Gilbert Is. (see p. 382).

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 385

a west-east line, hensilli in the western Carolines, scutoscriptus on Truk, hakanssoni
on Ponape and marshallensis from Marshall and Gilbert Is. ; these, except scuto-
scriptus, also have in common a black apical half to segment V (white in all other
species and sometimes in hensilli).

Species which may have segments III-V or IV-V interrupted by a line of dark
scales beneath, extend south-east from Marianas (guamensis) through Carolines (sp.
from Palau, scutoscriptus from Truk, hakanssoni from Ponape) to Sikyana in Solo-
mons, and Tonga (tongae) ; pseudoscutellaris in Fiji may show this character in
certain environmental conditions, and it may occur in aberrant pernotatus. Far
west of these species, andrewsi may have the band on segment IV interrupted, but
dorsally, not ventrally.

Complete abdominal bands are found in species in the western and central part
of the subgroup's range, from Andaman Is. sp., alorensis (Alor), s. katherinensis
(N. Australia), paullusi (Philippines), hensilli and sp. from Palau (Carolines), s.
scutellaris (New Guinea, etc.) to quasiscutellaris and gurneyi in Solomons, and north-
east to marshallensis (Marshalls and Gilberts) . On the outskirts of this distribution
almost complete abdominal bands are found to the north, in riversi (Okinawa) and
to the east in tongae (Sikiana and Tonga) and horrescens (Fiji).

The species which have only curving lateral patches on the abdominal tergites
are found from Caroline Is. (scutoscriptus, hakanssoni) south-east to Rotuma sp.
and pseudoscutellaris (Fiji) with polynesiensis ranging east from Fiji, and to the
west, pernotatus in New Hebrides. North of Caroline Is., guamensis also has lateral
patches but these are of distinctive shape. The exception to this distribution
pattern is andrewsi from Christmas Is., far removed from the other species with
lateral patches.

When one examines the form of the basal lobe, there is one particularly well-
defined type, a simple lobe with a row of specialized setae along its sternal aspect.
This can be traced from Okinawa (riversi) south-east through Marianas (guamensis),
Caroline Is. (hensilli in the east, hakanssoni in the west) to Sikyana and Tonga
(tongae) and Fiji (pseudoscutellaris). Rotuma sp. may be allied to this type and
polynesiensis is essentially similar but has lost the specialized setae. Other species
with a simple basal lobe are found in the western parts of the subgroup's range.
The specialized setae are at the apex in Andaman Is. sp. and andrewsi, while paullusi
has only non-specialized setae arising from its truncate apex. Though slightly
expanded in lateral view the basal lobes of s. scutellaris and s. katherinensis are
essentially similar to the simple type. There is a fairly close resemblance between
andrewsi and s. scutellaris, both of which have the specialized setae set on the sternal
angle of the tip ; Dr. Stone has noted a resemblance between riversi and s. katherin-
ensis.

Quite a different form of lobe is that with an elongated somewhat flattened apex,
with specialized setae borne on its inner projection (i.e., towards midline of genitalia).
This is found in the East Indian species alorensis (Alor) and in pernotatus (New
Hebrides) ; scutoscriptus (Caroline Is.) appears to be similar ; quasiscutellaris
(Solomons) and horrescens (Fiji) though rather different, might be derived from this
form ; gurneyi which lacks specialized setae could be derived either from this, or

386 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

from further flattening of the apex of a lobe of paullusi type. Marshallensis has a
similar inner projection, but the apex of the lobe is produced, not flattened, and it
could derive almost equally well from the simple lobe first discussed ; sp. from Palau
is apparently most like marshallensis but without specialized setae on the inner
projection.

Although the distribution of the different characters may suggest different affinities,
it is clear that (with one exception) there is a very definite west-east trend (as already
pointed out, it is unlikely to be east-west), which is in accordance with known
relationships of the fauna of Pacific islands. The notable exception is the north-south
distribution of mid-femur pattern, but this is in the Indo-Australian portion of the
range, where distribution of the fauna is a more complex problem.

(6) Immature Stages

Although, as already detailed, it was differences between larvae from different
areas that first led to the recognition of the scutellaris subgroup, Farner & Bohart
(1944) justly observed "An effective systematic revision of the larvae must await
the availability of greater amounts of reared and associated material as well as a
critical study of the taxonomic characters." No one has yet produced such a
revision.

No details have been recorded of larvae of andrewsi, alorensis, gurneyi, tongae or
spp. from Rotuma, Palau group and Andaman Is., though all except possibly
andrewsi, alorensis and Rotuma sp. have been reared from larvae. Descriptions
available for larvae of the other forms vary in their completeness from Woodhill's
(19490) note on the larvae of s. katherinensis " indistinguishable from those of Aedes
scutellaris scutellaris " to Knight & Hurlbut's (1949) detailed account of the larva
of hakanssoni.

Various authors have found characters for distinguishing the larvae where two
species occur in the one locality. Edwards (1935) tabulated characters for separating
larvae of horrescens 1 and " pseudoscutellaris " (Fiji) and indeed it was the observation
of two distinct larval forms which led to the recognition of horrescens as a distinct
species. Larvae of pseudoscutellaris and polynesiensis can be separated on the
relative lengths of their gills (Marks, 19516) so that the three Fijian species are identi-
fiable. Perry (1944) gave key characters for distinguishing the larvae of s. scutellaris
and pernotatus (New Hebrides). Bohart & Ingram (19466) were able to distinguish
hensilli larvae from those of scutoscriptus and sp. from Palau (Caroline Is.).

Belkin (1950) provided a system of nomenclature for the complete chaetotaxy of a
culicine larva. With the aid of this and adequate representative larval material
of the various species, it would very likely be found that many of the larvae could
be identified on a combination of characters, as are the adults, and possibly some
trends in the distribution of characters would appear. For example, examination
of a small number of larvae of pseudoscutellaris and polynesiensis has suggested that
hair 2 of abdominal segments III-VII is frequently single in pseudoscutellaris and

1 Edwards gives a series of distinctive characters. The presence of multi-branched hairs on thorax
and abdomen is not by itself sufficient to identify a specimen as horrescens, since occasionally these hairs
are fairly heavily branched in larvae of pseudoscutellaris and polynesiensis.

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 387

rarely so in polynesiensis. Long series from varied localities are essential for evalua-
tion of characters such as this. Our present knowledge is quite inadequate for
preparation of a key to the larvae of the subgroup or for useful discussion of the
specific larval characters.

No attempt has been made to separate species on pupal characters and the pupal
stages have in general been ignored though Knight & Hurlbut (1949) give details
of the pupa of hakanssoni, and Penn (1949) describes that of s. scutellaris.

The eggs of all species, so far as is known, appear identical but no comparative
measurements for different species have been recorded.

III. EXPERIMENTAL STUDIES OF VARIATION IN AEDES
PSEUDOSCUTELLARIS (THEOBALD)

(i) Introductory

The characters by which a culicidologist distinguishes the various members of
the scutellaris subgroup from one another have already been discussed in detail.

Descriptions of some species indicated a certain amount of variation in several
of the diagnostic characters. It was decided therefore to take a single species
and investigate the extent to which such morphological characters of the adults
could be affected by controlled variation of the larval environment. The results
would give some indication of the reliability of the different characters for taxonomic
purposes, and the factors involved in their phenotypic variation. They might
thereby provide some evidence on the validity of the specific status accorded to the
form investigated.

The common diagnostic characters of the subgroup were chosen for biometrical
study. These all (except on the genitalia) concern the proportions of white scaling
to black. Some additional characters of this type were studied, in view of their
possible taxonomic use and also to discover whether they varied in a similar fashion
to the rest. The measure usually given to indicate size of mosquitoes is the wing-
length. This was recorded, and also for comparison, the length of the hind femur.
It is a well-known zoological fact that size varies considerably with the temperature
of the animal's environment, as well as from other causes such as, in mosquitoes,
abundance of larval food.

In addition to these two measurements, the following characters were examined
(each is discussed in further detail in conjunction with its appropriate table) :

White scales under the proboscis ; white scales on the anterolateral margin of the
scutum ; white scales in front of the prescutellar bare area ; white patches at the
bases of fore and mid tarsal segments ; extent of anterior white streak and of dark
ventral scaling on hind femur ; greatest length of white bands on hind tarsi ; and
least length (i.e., the extent to which they are interrupted beneath) ; distance
between the lateral patches on abdominal tergites V and VI ; number of specialized
setae on the basal lobe of the male coxite.

Variation was apparent also in several characters which were not investigated
biometrically. In females bred at 15-16 C., the median white stripe on the scutum
was the width of about five scales across, whereas at 30-32 C., it was 9 or 10 scales

388 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

across. At 30-32 C., broad appressed white scales were developed in certain sites
where at lower temperatures the white scales were narrow curved. 1 This was
particularly marked on the anterior margin of the scutum at the commencement
of the median white stripe (some specimens from | sea-water were similarly affected);
a few broad scales were frequently present also at the posterior extremity of the
median stripe, and on the scutal angle.

The studies of variation described herein were made on specimens derived from a
laboratory colony, a small population which had been inbred for many generations.
Though it seems probable that small populations are favourable to relatively rapid
evolution (Mayr, 1942 ; Ford, 1949), they often show reduced variability due to
accidental gene loss or genetically limited ancestry. It is therefore likely that the
amount of variation observed in this stock is less than would occur in a natural
population subject to the same conditions ; this is confirmed by a comparison with
material from a colony of different origin. However, the specimens have been
reared in controlled conditions, so that what variation does occur can be related to
the variation in the environment.

Conditions were not controlled rigidly and the experiments were spread over a
period of approximately sixteen months. It is therefore not surprising that when
some of the series were tested for homogeneity in several cases appreciable hetero-
geneity was found. In this kind of exploratory work it is not necessary to make
very precise measurements of the effect on each series of changing one factor in the
environment and the interest is mainly in the qualitative differences. Nevertheless,
comparison between Series F, C, B and E, in which the temperature was progressively
raised while other conditions were the same, is very constant wherever any positive
effect occurs at all.

The effects of the different treatments are illustrated in Tables III-XX. In
certain cases where these were not clear-cut and it was a matter of interest to
examine them further, their significance was tested statistically. In the tables,
where specimens fall on the limits of two ratio groupings, they have in all cases
been placed in the lower category.

(2) Material

The species chosen for investigation was Aedes (Stegomyia) pseudoscutellaris
(Theobald) of which a culture was obtained from the following source :

The late Mr. D. W. Amos in August, 1948, sent by air larvae (all stages) of "pseudo-
scutellaris " from Fiji to Sir Philip Manson-Bahr in London. No particulars are
now available of the number of specimens with which the colony was started, nor
of the exact locality from which they had been collected, though it is believed to
have been Suva. Mr. P. G. Shute established the laboratory colony and it was
subsequently maintained as a continuously breeding population in the Parasitology
Department of the London School of Hygiene and Tropical Medicine. From thence

1 It would be interesting to know what factor is responsible for the development of a scale in one or
other of two such distinct shapes. The distinction between narrow and broad scales on a given site is
often used to separate species or even subgenera of mosquitoes.

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 389

a sample of all stages from egg to pupa was obtained in November, 1949. Approxi-
mately 350 adults were reared from it and their progeny are regarded as the first
(Cambridge) generation. The studies of variation were made on specimens from this
stock. Whether the original stock from Fiji was a pure culture of pseudoscutellaris
is unknown, but there is no doubt that the sample brought to Cambridge was a pure
culture of this species.

A second stock of pseudoscutellaris was received from Mr. B. A. O'Connor in April,
1951, direct from Fiji. This consisted of a batch of about 120 eggs obtained from
wild-caught males and females, collected ioth-i6th March at Naduruloulou Agricul-
tural station (about 12 miles in a direct line east of north from Suva). Approximately
60 adults were obtained from this sample, and their progeny, reared under standard
conditions, were compared with the other stock.

(3) Methods
(a) Rearing

The design of the experiments required that the conditions in which larvae were
reared should be controlled, though highly refined techniques and rigorous control
of all conditions were deemed unnecessary. A continuous breeding colony was not
maintained ; egg batches were hatched as required and the larvae reared with
the following standard technique.

Breeding bowls were kept at the required temperature in thermostatically con-
trolled incubators or in a constant temperature room. Straight sided glass bowls
of varying capacity were used.

Larvae. The liquid contents of the bowl in which the eggs were to be submerged
consisted of tap-water, " larval essence " and a small amount of larval food.

" Larval essence," used as a hatching stimulus, 1 was the filtered liquor in which a
previous culture of larvae had been reared. It was stored in a jar and used as
required in amounts of approximately 30-100 c.c. per bowl. The quantity was
not standardized since there was no means of standardizing the quality.

The size of bowl and the volume of tap- water used were adjusted to the number
of eggs to be hatched so that overcrowding of larvae was avoided.

Larval food consisted of a mixture of equal parts by volume of finely ground
Bemax and two types of dog-biscuit, one of which contained meat-meal. It was fed
either in aqueous suspension or the powder sprinkled directly on the surface of the
medium. Larvae were fed as necessary, usually daily or every second day. Enough
food was added to keep the liquid slightly cloudy, but to avoid formation of a scum.
The medium soon developed a rich culture of micro-organisms and it is likely that
these formed the bulk of the material actually ingested by the larvae.

Salinity. Specific gravity measurements made with an immersion hydrometer
indicated no increase of salinity in tap-water culture media at completion of rearing
at 25-28 C., and in essence, when compared with tap-water at similar temperatures.

1 Numerous workers have found that the presence of micro-organisms is one of the many factors
that stimulate hatching in Aedes eggs, including Buxton and Hopkins (1927) in their studies of " pseudo-
scutellaris " (= polynesiensis). The effect of the " essence " may have been through the micro-organisms
it introduced to the medium.

390 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

When rearings were made in diluted sea-water, the salinity was estimated by the
method described by Harvey (1928) of titration with silver nitrate solution. The
corrections he gives were applied and the results recorded in parts per thousand
(% ) to the first decimal place. The salinity of selected samples from other con-
ditions was measured in the same way.

Air. Compressed air was bubbled slowly through the larval medium ; if this
was not done a scum formed on the surface and larvae and pupae drowned.

Pupae were collected daily or every second day and transferred to a dish of clean
water in a cage ; in other respects the pupae were subjected to the same variations
in environment as the corresponding larvae. When all pupae in the dish had
emerged, the cage was transferred to a constant temperature room at 24-26 C.

Adults of all series were kept in cages (8 in. cubes or larger) in a constant tem-
perature room at 24-26 C. The humidity of the room varied, but glass bowls
with moist filter paper or porous earthenware pots of water were placed in the cages
to provide moisture. Most specimens required for subsequent examination were
killed 24-48 hours after emergence.

No food was given other than blood meals. The source of these was usually man,
less frequently rabbit and once chicken. A blood meal was offered when the majority
of females in the cage were 3-5 days old and continued long enough for numerous
specimens to engorge (usually 20-30 minutes). Sometimes a second meal was
given on a later day. Porous earthenware pots containing water were provided
for oviposition. They were removed within 7 days of the blood meal and the water
decanted, but the pots were kept moist for a further 1-2 days. They were stored in
the constant temperature room at 24-26 C. and shards with the required quantity
of eggs broken off when needed. Where egg batches were obtained from individual
adults, these were laid on moist filter paper in a glass tube.

Specimens for study were usually pinned through the side as this was least likely
to damage the characters studied. Each specimen received a label with its series
letter (indicative of the conditions of rearing) and a serial number.

Records. Day by day records were kept for each batch of larvae reared. These
included source and approximate or exact number of eggs ; date submerged ;
particulars of medium ; number of pupae collected ; emergence of adults (in certain
samples the proportions of the sexes were recorded from a count of pupal skins) ;
date and source of blood meal ; date of oviposition.

(b) Sampling

Sampling was at two levels, firstly from the total of adults reared in a batch,
secondly from the collection of pinned specimens. In all cases damaged specimens
were rejected. In the case of progeny of individual females, all suitable adults
reared were pinned. With large collections of adults of mixed parentage, if none
were needed for breeding, all were killed and the required number of specimens
pinned. If it was desired to breed from the adults, each specimen for pinning was
captured by hand in a glass tube. This method was found more satisfactory than
an aspirator for the purpose of obtaining perfect specimens.

Pupation of a batch of larvae extended over a period of days. The pupae (and

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 391

adults emerged therefrom) in lots representing i, 2 or 3 days pupation were kept in
separate cages until samples had been taken. The majority of males emerged in
the first few days, while the proportion of females gradually increased. The pro-
portion of each sex to be pinned from any particular cage was in several cases known
accurately from a count of sexes of the pupal skins. In most cases the numbers
were estimated in the light of this experience, by direct observation of the adults
to be sampled. The serial number of a pinned specimen indicated the batch from
which it came and also the collection of pupae within that batch. The specimens
examined represented proportionately the number pinned from each pupal collection
within the batch. In this way a representative cross-section of the population
was obtained.

It should be made clear, however, that though equal numbers of each sex from a
batch might be pinned and equal numbers examined, the numbers of each sex
actually reared in that batch were not necessarily equal.

From the males examined, a smaller number were selected on the same principle
for dissection of the genitalia.

(c) Examination of specimens

The characters of the external morphology of the specimens were examined with
a binocular dissecting microscope. Measurements were read on an eyepiece micro-
meter with divisions of o-i mm. ; using X 7 eyepieces and X 5 objective, 38 divisions
of the scale represented i mm.

A rotating insect stage was used for orientating specimens for examination of
characters.

Only one leg or wing (the one most easy to observe) was measured on each
specimen.

Genitalia. The technique employed was a simple one which facilitated the
preparation of a large number of mounts.

The specimen was relaxed and the terminal segments cut off and transferred to a
small tube of 10% potassium hydroxide. This was set in a beaker of water, which
was heated to boiling-point, and it was allowed to remain in the hot water for
5 minutes. The genitalia were transferred to a watch glass of distilled water, plus
acetic acid. After five minutes they were transferred to cellosolve on an excavated
slide and then dissected. With a straight surgical needle the genitalia were divided
longitudinally into two halves, the basal lobe of one half was detached with the
aid of fine steel pins. 1

The parts were transferred to chloral gum medium on a slide, arranged so that
both basal lobes were in lateral view, and covered with a piece of glass coverslip.
They were examined with a monocular microscope using ^ in. objective.

When required, scale drawings were made with the aid of a squared eyepiece
micrometer.

1 It is recommended that in specimens where the basal lobes are to be examined in lateral view, one
lobe should be left attached to the coxite so that it may readily be determined which is the inner and
which the outer lateral view.

392 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

(d) Variation of larval environment

Only one factor was varied in the larval environment in any one series of experi-
ments.

An arbitrary " standard " series was reared as already described, in a tap- water
medium at 25-28 C. (Series B).

The following series were reared similarly but the temperature was varied :

i5-i6 C. (Series F).
i9-22 C. (Series C).
30-32 C. (Series E).

After preliminary experiments it was found that larvae could be reared satis-
factorily in a medium containing ^ sea-water. Series D was reared in ^ sea-water
at 25-28 C.

Specimens of Naduruloulou stock (Series H) were reared similarly to Series B.

Attempts to rear larvae at 10 C., and at temperatures above 32 C. were unsuc-
cessful.

The following summarizes the particulars for each series :

Series B. 2^-28 C.

About 40 batches representing nine generations were reared. Many of these
were for the purpose of maintaining the stock, but 1,394 specimens (658 $$, 736 $$)
were pinned from 19 batches, either mass rearings or individual egg batches. Speci-
mens representing nine of these batches were studied ; they had been reared
from December, 1949, to July, 1950. The 152 males examined came from
six batches (ist generation mass rearing, 54 ; 3rd generation individual batches,
9, 9 ; 4th generation individual batches, 36, 14, 30). From the same batches
respectively (excluding one 3rd generation batch) females were taken in the following
numbers, 20, 10, 10, 10, 10 ; and in addition from a 3rd generation mass rearing
(20) and 5th and 6th generation individual rearings (10, 10), making a total of 100,
from eight batches. The mean temperature reading for the larval medium for one
of these batches was 27-4 C. (6 readings). Genitalia of 25 males were dissected.

Series F. i5-i6 C.

One batch was reared in February-March, 1951, 350 specimens pinned (202 <
148 $$) and 100 of each sex examined. The batch was gth generation (from Series C
eggs, laid by females of 2nd generation bred at I9-22C.). Mean temperature
was 15-2 C. (30 readings).

Genitalia of 25 males were dissected.

Series C. i.g-22 C.

Six batches were reared from February, 1950, to March, 1951, 500 specimens
pinned (250 , 250 $$) and 100 of each sex examined. The first three batches
(50, 50, 100 pinned adults) were from eggs from Series B (ist, 4th and 7th generations
respectively). From the third of these, three succeeding generations at i9-22 C

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 393

were bred (100 adults pinned from each). Twenty per cent of each lot of pinned
specimens was examined, the sexes being equally divided.

The following are the mean temperatures of the medium for each batch, with the
number of readings in parenthesis : 20-5 C. (8), 20-3 C. (9), 21-3 C. (3), 20-5 C. (19),
207 C. (19), 20-5 C. (10).

Gemitalia of 25 males were dissected.

Series E. 3O-32 C.

Four batches were reared from December, 1950, to March, 1951, 469 specimens
pinned (lots of 60, 88, 300 and 21 totalling 260 $$, 199 $$), and 100 of each sex
examined. From the above batches respectively were taken 35, 30, 21 and 14 males
and 18, 28, 48 and 6 females. The first two batches were 8th generation eggs from
Series B ; the third, gth generation eggs from Series C (laid by female of 2nd
generation bred at i9-22 C.) ; the fourth, the progeny of the third. Mean
temperatures of the medium for each batch (number of readings in parenthesis) were
31-1 C. (12), 307 C. (22), 30-6 C. (23), 29-9 C. (8).

Genitalia of 25 males were dissected.

Series D, 25-28 C., sea-water.

Five batches were reared from June to November, 1950, in a medium of sea-water
at 25-28 C., 500 specimens pinned (250 $<$, 250 $$) and 100 of each sex examined.
The first batch was 6th generation eggs from Series B and the remainder were
succeeding generations bred from these ; from each 100 adults were pinned, 20%
of which were examined, the sexes being equally divided. For two batches the mean
temperatures (number of readings in parenthesis) were 27-1 C. (4), 25-9 C. (9).

The following are the salinity measurements at the beginning and end of each
rearing :

Initial Number Final

salinity. of days. salinity.

il ) (l \

\ tool \ loo/

11-7 . 23 . 16-2

n-9 . 15 . 13-1

I2-I . 13 . I2'7

n-9 . 13

11-7 . 15 . 15-0

The sea-water used was the laboratory stock from Lowestoft ; its salinity,
measured on one occasion, was 34'6% .
Genitalia of 25 males were dissected.

Series H. 25-28 C.

One batch, the 2nd (Cambridge) generation of the Naduruloulou stock, was reared
for study in April-May, 1951, 97 adults pinned (47 <$<$, 50 $$) and 25 of each sex
examined. Two temperature readings of the medium were 27 C.

Genitalia of 10 males were dissected.

394

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

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A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

395

(4) Results
(a) Tables III and IV : Length of wing and of hind femur

Hind femur

Series F. i5-i6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3O-32 C.
Series D, 25-28 C.,

seawater
Series H, 25-28 C.,

different stock

Females

Series F, 15-! 6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C.,

different stock

TABLE IV. Aedes pseudoscutellaris

Length (38 units = i mm.)
Length of femur.*

Number of 48- 51- 54- 57- 60- 63- 66- 69- 72- 75- 78- 81- 84- 87-
specimens 50 53 56 59 62 65 68 71 74 77 80 83 86 89

i 21 56 20 2

IOO

i

21

IOO

2

7

12

52

152

2

24

49

25

IOO

n

11 26

2O

25

9

2

IOO

2

22

49

21

6

19

25 .

IOO
IOO
IOO
IOO
IOO

25

_ 3 6

2 3 5 12 32 25 16 5
7 12 37 29

4 10 29 39 16
2 13 23 38 21
21 42 24 4

i

. 1676 8

* Numbers of specimens expressed in percentages.

The measurements have not been converted from the units in which they were
made, since tabulation of these shows clearly the effects of different larval conditions.

Wing-length was measured from the large sclerite at the base 1 to the tip of the
wing, excluding the fringe. The table shows that the longest wings are found in
specimens bred at lowest temperatures, and conversely the shortest in those bred
at the highest temperatures, with a gradient between. The wings of both males
and females reared in sea- water (Series D) are significantly shorter (at 0-1% level)
than those reared in tap-water at the same temperature (Series B) . Wings of females
are longer than those of males in the same series.

The length of the hind femur was measured from its upper basal margin to the
tip of the white patch at the apex. It shows a similar gradation to the wing-length
in relation to temperature except that this is not apparent between females bred
at i5-i6 C. and i9-22 C. In Series D from sea-water the femur is shorter

1 At the wing base there is a complicated system of small sclerites ; the largest of these is a prominent
dark shield-shaped sclerite lying just posterior to the base of the costa and articulating distally with
the remigium (the united bases of veins of the radial complex). Prashad (1918) in his description of
the wing-joint in Anophelines calls it the " epaulette " ; S. R. Christophers (personal communication)
regards it as homologous with the 2nd axillary of Snodgrass (1935). It was found convenient to measure
the wing-length from the proximal edge of the well-defined sclerite, rather than from the actual base
of the costa which was not always easy to observe. The difference between measurements taken from
these two points is less than 2% of the total wing-length and therefore of negligible importance io
comparisons between species.

ENTOM. Ill, io. 28

396

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

than in Series B from tap-water ; the difference is highly significant. The femur
is longer in females than in males from the same series.

The ratio of the length of the wing to that of the hind femur was plotted for
several series. This showed some correlation but there was considerable scatter of
the points, so that one could not be deduced from the other with any precision.

(b) Table V : Pale scales under the proboscis

TABLE V. Aedes pseudoscutellaris

Proboscis

Males

Series F, i5-i6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C.,

different stock

Females

Series F, i5-i6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C.,

different stock

Number of
specimens

Pale streak on under side
Number of pale scales*

i-io 11-20 21-30 31-40 41-50 51-60 61-70

IOO

64

25

8

3

IOO

32

52

13

3

152

20

43

30

6

i

IOO

16

48

20

13

3

IOO

i?

69

13

i

25

16

IOO

93

7

IOO

67

28

5

IOO

58

3i

6

4

IOO

64

3i

5

IOO

66

3i

3

25 24 12 24 20

* Numbers of specimens expressed in percentages.

16

The amount of pale scaling under the proboscis was recorded by an actual count
of the number of pale scales. These observations led to the conclusion that a count
of more than ten could be taken to represent the presence of a pale streak under
the proboscis.

Ignoring for the moment the implications of the counts for Series H, the table
illustrates that a pale streak is of much more frequent occurrence in males than in
females. The variation of larval environment has had little effect on the amount
of pale scaling in the female, except at the lowest temperature, i5-i6, when a
much higher proportion have the proboscis completely dark. This applies also to
males at i5-i6 C., and there is a progressive reduction in the number of males
with a recognizable streak at temperatures below 25-28 C., but not an appreciable
increase above. Series D in | sea-water also shows a reduction in numbers with a
streak.

Series H, from different stock, has a very high proportion of both sexes with a
streak. This emphasizes that though the observations of variation in one stock
may suggest certain conclusions, it would be unsafe to interpret them as applying
rigidly to all members of the species, let alone to related species.

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 397

(c) Table VI : Scutal pattern

Scutum

Males

Series F, i5-i6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

| sea-water
Series H, 25-28 C.,

different stock

Females

Series F, 15-! 6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

$ sea-water
Series H, 25-28 C.,

different stock

TABLE VI. Aed.es pseudoscutellaris

Extent of white scaling on anterolateral margin
Scutal pattern * (see Fig. 19).

Number of
specimens

100

100
151

IOO
IOO

25

IOO
IOO
IOO
IOO
IOO

25

B-

65

II

69

II

5

i
i

30
35

4
4

i

12

29
61
16
32

18

20

A-

4
43

M
29

28

40

2
24

59
32
55

52

* Numbers of specimens expressed in percentages.

B

i

ii
82
46
65

40

A +

20
6

4

20

30 5

25

28

A-

A A +

FIG. 19. Variation in white scaling on anterolateral margin of scutum of Aedes pseudo-
scutellaris (Theobald). (B -, F. 268 ; B, C. 44 ; A -, C. 152 ; A, D. 488 ; A +, E. 123 ;
all ??.)

The line of white scales along the anterolateral margin of the scutum distinguishes
pseudoscutellaris from the closely similar species polynesiensis. The line may be
definite, or faint and interrupted, but there are always some pale scales present.

For biometrical interpretation the amount of white scaling was divided into
five categories, B -, B, A -, A, and A +, B - having the least and A + the greatest

ENTOM. Ill, 10. 2 8

398

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

development of white scales. These categories are illustrated from typical specimens
in Fig. 19.

B : white scales confined to a small patch on the scutal angle.

B : white scales on the scutal angle and extending forward slightly along the
margin ; or else only on the scutal angle, with 2 or 3 scales further forward.

A : white scales extending along the anterolateral margin, but the line inter-
rupted to a greater or lesser extent, sometimes complete on one side and incomplete
on the other.

A : white scales forming a narrow continuous line along the anterolateral margin.

A+ : white line on anterolateral margin quite thick and well marked.

There is no line of demarcation between these categories and interpretation is
purely a matter of judgment, but by allotting the specimens among them, they do
present a general picture of the variation in this character.

From the table it appears that at 15 C. the amount of white scaling is reduced
to a minimum and the pattern is similar in both sexes ; with increase of temperature
the white scaling increases, but to a greater extent in males than in females ; a larval
environment of ^ sea- water does not appreciably affect it.

The presence of white scales on the anterolateral margin of the scutum is the only
character known to distinguish females of pseudoscutellaris from polynesiensis.
Specimens of the latter occasionally have 1-3 pale scales on the scutal angle. A
count of the number of white scales on the scutal angle in specimens of pseudo-
scutellaris recorded as B from all series, (165 out of a total of 1,101 examined)
gave a range of 1-13 with a mean of approximately 6 scales for 299 scutal angles
(one side of a specimen was sometimes rubbed or obscured) .

(d) Table VII : White scales on either side of the prescutellar bare area

TABLE VII. Aedes pseudoscutellaris

Scutum

Males

Series F, 15-! 6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25^28 C.,

different stock

Females

White scales on either side of prescutellar bare area

Total number of white scales on both sides.*

Number of , ^

specimens o 1-4 5-8 9-12 13 or over

IOO

31

54

M

IOO

27

55

15

135

6

61

27

IOO

26

55

15

> IOO

18

54

23

Series F, 15-! 6 C.

IOO

50

39

10

Series C, i9-22 C.

IOO

61

33

5

Series B, 25-28 C.

, IOO

4i

42

13

Series E, 3o-32 C.

IOO

60

37

3

Series D, 25-28 C.,

IOO

52

42

5

\ sea-water

Series H, 25-28 C.,

: 21

76

24

4

different stock

* Numbers of specimens expressed in percentages.

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

399

In certain species the white median line on the scutum divides into a definite fork
on either side of the prescutellar bare area. In recording this character an actual
count of the total number of white scales on both sides of the prescutellar bare area
was made, and from these observations it was concluded that a count of more than
eight represented a more or less complete fork.

The table shows comparatively little variation in this character and nothing
that could be safely correlated with variation in larval environment. Only a very
small percentage of specimens show a tendency .to form a complete fork ; males,
however, do have a few more white scales than females.

The evidence suggests that this character is less subject to variation than some
others concerned in scale pattern.

(e) Table VIII : White basal patches on fore and mid tarsi

Fore and mid tarsi

Males

Series F, i5-i6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C..

J sea-water
Series H, 25-28 C.,

different stock

Females

Series F, i5-i6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C.,

different stock

TABLE VIII. Aedes pseudoscutellaris

Number of
specimens

100
100
151

IOO
IOO

25

White basal patches
Some white scales at base of one or both segments

Fore tarsus
II III

IV

I

IOO
IOO
IOO
IOO
IOO

IOO
IOO
IOO
IOO
IOO

IOO
IOO
IOO
IOO
IOO

Mid tarsus
II III IV

98
99

97

IOO

99

IOO -

IOO

IOO

IOO

20

IOO

IOO

IOO

IOO

IOO

12

IOO

IOO

IOO

IOO

IOO

58

IOO

IOO

4

IOO

IOO

IOO

92

I

23

IOO

IOO

21

IOO

IOO

IOO

66

3

13

IOO

IOO

II

25 . ioo ioo 72 4 8 . 100 100 16

* Numbers of specimens expressed in percentages.

One or more white scales at the base of a segment were interpreted as a white
patch. Usually both tarsi of the pair were examined and a patch was recorded if
there were white scales on one or both.

The table shows that pseudoscutellaris normally has white patches at the base
of fore and mid tarsal segments I and II ; the male seldom has more and in these
characters is less subject to variation than is the female.

Females frequently have a white patch at the base of fore tarsal III and though
there is no positive correlation with temperature at i5-i6 C. and i9-22 C.,
there is a marked increase in the number with this character at 25-28 C. and
again at 30-32 C.; larval environment of sea-water does not produce any notable

400

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

effect. At 3O-32 C. and in ^ sea-water an occasional specimen has a patch on
IV, and a higher proportion show one on V. The different stock of Series H show
these characters at 25-28 C. Below 25- 28 C. patches are absent from mid
tarsal III ; at this temperature few specimens show them, but there is a marked
increase at 30-32 C. and a lesser one in ^ sea-water ; Series H again resembles
the latter two series.

The more frequent occurrence of white patches on the fore tarsal segments than
on the mid is rather unusual amongst mosquitoes and does not appear to have been
observed in other members of the scutellaris subgroup. The usual tendency is for
the amount of white scaling to be least on the fore tarsus, somewhat more extensive
on the mid tarsus, and greatest on the hind tarsus.

(/) Tables IX and X : Extent of white anterior streak and ventral dark line on the
hind femur

TABLE IX. Aedes pseudoscutellaris

Hind femur

Males

Series F, i5-i6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea- water
Series H, 25-28 C.,

different stock

Females

Series F, 15-! 6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C.,

different stock

Number of
specimens

100
100
152

IOO
IOO

25

IOO
IOO
IOO
IOO
IOO

25

Length of anterior white streak
Ratio of length of streak to total length of femur.*

o-75-
0-80

0-80-
0-85

32
30
16
29

12

44

58
37
49
44
30

44

0-85-
0-90

67
68
84
70
84

56

42
62

54
69

56

0-90-
o-95

o-95-

I -OO

* Numbers of specimens expressed in percentages.

The hind femur at its base joins the trochanter, which projects ventrally a short
distance taken from the base of the femur. All measurements taken from the base
of the femur were taken towards the dorsal side ; if taken from beneath they would
have been slightly shorter.

The length of the white anterior streak has not been used taxonomically, nor is it
likely to be of value in that respect since many species have it apparently equally
developed. However, the opportunity was taken to investigate whether this
character varies in the same way as other characters of white scaling. Table IX
shows no apparent correlation between the length of the streak and the conditions
in which larvae of the different series were reared.

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

401

TABLE X. Aedes pseudoscutellaris

Hind femur

Males

Series F, 15-! 6 C.
Series C, i9-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28c.,

different stock

Females

Series F. 15-! 6 C.
Series C, i9-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

\ sea-water
Series H, 25-28 C.,

different stock

Number of
specimens

100
100
152

IOO

99
25

IOO
IOO
IOO
IOO
IOO

25

18
25

12

3

13

34

3i

19

7

6

Distance from base, of femur to beginning
of dark ventral scaling

Ratio of distance to total length of femur.*

o-oi-
o- 10

O- IO-
0-20

44
48
18

19

8

39
32

28

25
29

O-2O-
0-30

34
25
48

65
59

76

22

35
43
56
55

72

0-30-
0-40

21

18

6

12

9
16

* Numbers of specimens expressed in percentages.

A possible difference was observed between pseudoscutellaris and polynesiensis in
the extent of the dark ventral scaling on the hind femur. The amount of variation
in this character in pseudoscutellaris was therefore investigated. The measurement
was made from the base of the femur, as above, to the beginning of the dark scaling.
If a line of dark scales reached right to the base ventrally, it was counted as zero,
though owing to the trochantal projection, the micrometer reading was a small
number. This explains the low frequency in the o-oi-o-io ratio in Table X, as
compared with that in Groups o and 0-10-0-20. This fault was not serious from a
point of view of comparison with other species. Table X shows that this character
follows the general trends already shown for others. The amount of dark scaling is
greatest at low temperatures and following a gradient, becomes least at 30-32
(though this gradation is not found between males at 15 C. and i9-22 C.) ; sea-
water apparently does not affect it.

It may be added here that of 33 specimens of polynesiensis from various localities
examined for this character 3 fell in the category 0-20-0-30, 9 in 0-30-0-40, 20 in
0-40-0-50, and i in 0-50-0-60. This suggests a difference between the two species,
but it may be a geographically variable character ; insufficient specimens of poly-
nesiensis from Fiji were examined to indicate whether it would be a useful character
there.

(g) Tables XI-XVIII : The bands on the hind tarsal segments
Each band was measured at its greatest length, i.e., from the base to the further

402

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

extremity of the white scales ; the bands were longest on the upper anterior (outer)
surface of the segments.

The band on hind tarsal segment I was always interrupted by a dark line beneath.
The bands on II-IV were measured also at their least length, i.e., from the base
of the segment to the nearest dark scaling which was continuous to the apex. Some-
times a band was slightly interrupted by dark scales at its base but formed a con-
tinuous ring distal to these ; such interruptions were ignored. In other words, the
measurements of least length are of the difference between the greatest extent of
continuous dark scaling and the total length of the segment. Bands were narrowest
on the lower posterior (inner) surface of the segment.

A continuous line of dark scales beneath, even if only one scale in width was
regarded as a complete interruption ; sometimes these scales were greyish rather
than black, but contrasted with the white.

Table XI : White band on hind tarsal segment I

TABLE XI. Aedes pseudoscutellaris

Hind tarsal, Segment I

Males

Series F, i5-i6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

$ sea-water
Series H, 25-28 C.,

different stock

Number of
specimens

100
100
151

100
IOO

25

Greatest length of white band
Ratio of white band to total length of segment.*

0-15-

O-2O

82
72
13

3

15

0-25-
0-30

ii
25
85
7i
75

72

0-30-
o-35

2

25
IO

28

o-35-
0-40

Females

Series F, 15-! 6 C. 100

Series C, i9-22 C. 100

Series B, 25-28 C. 100

Series E, 3o-32 C. 100

Series D, 25-28 C., 100

sea-water

Series H, 25-28 C., . 25

different stock

82

73
ii

i
6

27
79
57
80

84

10
39
14

16

* Numbers of specimens expressed in percentages.

The length of the band varies in relation to the temperature gradient, being least
at i5-i6 C., and greatest at 30-32 C. It is apparently not affected by a larval
environment of ^ sea-water. Females from the different stock, Series H, do not
differ from others reared at 25-28 C., but the males resemble those reared at
30-32 C. The length of the band varies little between males and females of the
same series.

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 403

Tables XII and XIII : White band on hind tarsal segment II
TABLE XII. Aedes pseudoscutellaris

Hind tarsal, Segment II

Males

Series F, i5-i6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3O-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C..

different stock

Females

Series F i5-i6c.
Series C, i9-22 C.
Series B, 25-28 C.
Series E 3o-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C.,

different stock

Number of
specimens

100

100
152

IOO
100

25

IOO
IOO
IOO
IOO
IOO

25

Greatest length of white band
Ratio of white band to total length of segment.*

o-2o-
0-25

4
4

i

0-25-
0-30

81

60

15

5

8

82

50

9

I
4

0-30-
o-35

15
34
62

53

66

72

15
46
60
23
47

68

o-35-
0-40

2

22
40
26

28

I
I

31
71
51

28

0-40-

o-45

* Numbers of specimens expressed in percentages.

TABLE XIII. Aedes pseudoscutellaris

Hind far sal, Segment II

Males

Series F, i5-i6 C.
Series C, i9-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C.,

different stock

Females

Series F, 15-! 6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3O~32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C.,
different stock

Number of
specimens

IOO
IOO

150

IOO
IOO

25

IOO
IOO
IOO
IOO
IOO

76

30

I

2
2

78
20

Least length of white band
Ratio of white band to total length of segment*

o-oi-
0-05

0-05-

O'lO

10

5

i

O-IO-

0-15

13

28

9

2
IO

12

31

5

I

O- IS-
C' 20

34
49

52
44

4

4
4i

20

35

25 36

* Numbers of specimens expressed in percentages.

0-20-
0-25

2

35

46

34

5

49
53

57

0-25-
0-30

5

26
8

64

The band, both in its greatest and in its least length, follows the temperature
gradient ; the amount of white scaling increases from i5-i6 C. up to 3O-32 C,

4 o 4

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

Considering the greatest length of the band, males appear unaffected by sea-water
but females show a slight increase in white scaling.

Females reared at 3O-32 C. and in sea-water in both measurements show more
white scaling than males of the same series. 1

In regard to the least length of the band the most noteworthy effect of the varied
conditions is the appearance at low temperatures of a high proportion of specimens
with the band completely interrupted by dark scales. Females from sea- water
show a slight increase in the white scaling but males are not affected. The different
stock of Series H also has slightly more white scaling.

Tables XIV and XV : White band on hind tarsal segment III

TABLE XIV. Aedes pseudoscutellaris

Hind tarsal. Segment III

Males

Series F, i5-i6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C..

\ sea-water
Series H, 25-28 C.,

different stock

Females

Series F, i5-i6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C..

different stock

Greatest length of white band
Ratio of white band to total length of segment*

Number of
specimens

100

100

152

IOO
100

25

IOO
IOO
IOO
IOO
IOO

0-30-
o-35

o-35-
0-40

0-40-
o-45

o-45-
0-50

0-50-
o-55

16

68

25 4 72 20

* Numbers of specimens expressed in percentages.

o-55-

O'OO

55

36

7

13

61

26

2

43

49

5

i

8

50

30

12

19

64

13

4

O'OO-

0-65

0-65-
0-70

16

46 49

4

6 48

46

5

72

19

4

i

23

39

30

6

_.,, ^ 2

39

3

ll

2

The bands show the same correlation with temperature as do those on Segments I
and II. Both males and females from sea-water show an increase in the greatest
length of the band ; the females do so in the least length also but it is not affected
in the males. Males of different stock (Series H) have slightly longer bands, but not
the females ; neither differ much in the least length from the original stock. In
all series the females have slightly longer bands than the males and less extensive
dark scaling beneath. At i5-i6 C. a high proportion of both sexes have the white
band completely interrupted.

1 This effect is no doubt partly due to the fact that, as many females emerge later than the males,
their immature stages have had longer in the abnormal conditions. It was observed that in a proportion
of specimens from these two series the tarsal segments did not become fully extended after emergence ;
grossly abnormal specimens were not used for measurements, but in others this had probably occurred to
a less marked degree.

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

4<>5

TABLE XV. Aedes pseudoscutellaris

Hind tarsal, Segment III

Males

Series F, i5-i6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3O-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C.,

different stock

Females

Series F, 15-! 6 C.
Series C, i9-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C.,

different stock

Number of
specimens

100
100
152

IOO
IOO

25

IOO
IOO
IOO
IOO

IOO

Least length of white band
Ratio of white band to total length of segment*

O-OI- O-IO-

O-2O-

0-30-

0*40-

o 50- o 60-

o

O-IO O-2O

0-30

0-40

0-50

0-60 0-70

66

3 21

IO

12

I 14

50

23

II

87

2

3

65

32

I

14

81

4

12

84

4

4 8

4 25

23

3

8

40

49

I

85

14

30

67

2 I

2

58

40

25 . 4 92

* Numbers of specimens expressed in percentages.

4

Tables XVI and XVII.: White band on hind tarsal segment IV
TABLE XVI. Aedes pseudoscutellaris

Hind tarsal, Segment IV

Males

Series F, 15-! 6 C.
Series C, i9-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C.,

different stock

Females

Series F, i5-i6c.
Series C, i9-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C.,

different stock

Greatest length of white band
Ratio of white band to total length of segment*

1

Number of o 50

o-55-

0-60-

o 65-

0-70-

o-75-

0-80-

0-85

specimens 0-55

0-60

0-65

0-70

o-75

0-80

0-85

0-90

IOO . 2

12

64

21

i

.

IOO .

5

33

48

13

i

152 .

3

16

51

29

i

IOO .

i

15

57

24

2

i

IOO .

2

2O

52

24

I

i

25

4

56

36

4

IOO

13

40

4

7

IOO .

i

14

65

19

i

IOO .

3

3i

5i

15

IOO .

i

4

36

48

9

2

IOO .

5

44

44

6

I

25

28

16

* Numbers of specimens expressed in percentages.

406

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

TABLE XVII. Aedes pseudoscutellaris

Hind tar sal, Segment I V

Males

Series F, i5-i6 C.
Series C, ig -22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea- water
Series H, 2 5 -28 C.,

different stock

Females

Series F, i5-i6 C.
Series C, ig -22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

\ sea-water
Series H, 25-28 C.,

different Stock

Number of
specimens

100
100
152

IOO
TOO

25

IOO
IOO
IOO
IOO
IOO

25

Least length of white band
Ratio of white band to total length of segment

o-oi- o-io- 0-20- 0-30- 0-40-
o-io 0-20 0-30 0-40 0-50

75
6

7 13 3 i

5 6 18 44

22

17

3

18

2

I
2

50 I 5 14 16 14

i 5 43

5

24

0-50- 0-60- 0-70-
0-60 0-70 o 80

21

65
60

73

48
61

21

39
52

ii
39

60

I

34

72 6

58

24

* Numbers of specimens expressed in percentages.

The greatest and least length of the band is affected by temperature similarly
to the bands of the preceding segments. Specimens reared in ^ sea- water show some
increase in white scaling, though this is not very marked in the least length of the
band of the males. Series H is not greatly different from the original stock. The
bands are longer, by both measurements, in females than in males of the same series.
A high proportion of specimens reared at i5-i6 C. have the bands completely
interrupted.

In pseudoscutellaris this segment is all white above, but occasionally has one or
two dark scales at the base on its under side ; these seldom form any obvious inter-
ruption to the white scaling. Their presence was noted, and if they extended an
appreciable distance along the segment, an estimate of this was made but no exact
measurement.

The table records the percentages of specimens in which dark scales were present
and also the percentage (of the total number of specimens) in which there was an
observable interruption to the white scaling by a dark line beneath. The length
of the dark line was measured from the base of the segment ; occasional dark scales
at the tip were ignored. The categories are unequal and, except for total interrup-
tion, approximate.

Above 22 C. there is no great difference between the series though females at
25-28 C. tend to have dark scales more often than those at 3O-32 C. or in ^ sea-
water ; males from Series H have them more often than those of the original stock,
but females do not differ. However, none of these series show any appreciable
interruption by dark scales.

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

407

Table XVIII : Dark scaling on hind tarsal segment V

TABLE XVIII. Aedes pseudoscutellaris

Hind tarsal, Segment V

Males

Series F, i5-i6 C.
Series C, iQ-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

sea-water
Series H, 25-28 C.,

different stock

Females

Series F, i5-i6 C.
Series C, i9-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C..

\ sea-water
Series H, 25-28 C..

different stock

Number of
specimens

100
100
152
99
100

25

IOO
IOO
IOO
IOO
IOO

25

A mount of dark scaling beneath

Ratio of dark line to total
Dark scales* length of segment*

t i

0-25-0-50 0-50-0-99 i-oo

Absent

Present

4

96

39

61

99

i

94

6

92

8

72

28

10

90

60

40

89

ii

99

i

99

i

88

51

2

12

1

* Number of specimens expressed in percentages.

(h) Table XIX. Distance between lateral patches on tergites V and VI
TABLE XIX. Aedes pseudoscutellaris

Abdominal tergites V and VI

Tergite V

Males

Series F, i5-i6 C.
Series C, i9-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

$ sea-water
Series H, 25-28 C.,

different stock

Females

Series F, i5-i6 C.
Series C, i9-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

^ sea-water
Series H, 25-28 C.,

different stock

Horizontal distance between lateral white patches
Tergite VI

Distance between
patches*

1

Distance between
patches*

Number of 5 or more Number of
individuals 1-4 scales scales individuals

5 or more
1-4 scales scales

IOO
IOO

148

IOO
IOO

2
2

3

8

4

98
98

97
92
96

IOO
IOO

144

IOO
IOO

2
2

8
3

IOO
9 8
9 8
92

97

25

9

91

25

12

88

IOO

IOO

IOO

IOO

IOO

IOO

IOO

IOO

IOO

TOO

IOO

IOO

IOO
IOO

i

99

IOO

IOO
IOO

3

2

97
98

25

25

96

* Numbers of specimens expressed in percentages.

4 o8 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

At i9-22 C. there is a marked increase in the number of specimens with dark
scales and a few males have a measureable dark line ; at i5-i6 few specimens
are without dark scales and a notable proportion, particularly of males, have an
incomplete dark line beneath, while in a few the interruption is complete.

Tergites V and VI were selected for the study of variation in abdominal markings
as in all members of scutellaris subgroup the characteristic pattern of the species is
most frequently found on one or both of these segments. In order to discover
whether there was any tendency under different treatments for the lateral patches
of pseudoscutellaris to extend and form a complete transverse band, the greatest
horizontal distance between the lateral white scales was measured. Where the white
scales were confined to continuous patches this is the distance between the patches,
but in many cases beyond the tip of the continuous marking (i.e., towards the mid
line) there were one or two isolated white scales.

Since the upper surface of the tergite is curved to a variable extent the measure-
ment actually represents the chord of the arc, but the difference would be unim-
portant.

Measurements were made with a micrometer, 38 divisions of which equalled i mm.
It was frequently observed that single scales measured in width approximately one
of these divisions. The distances have therefore been expressed in the table as
number of scales, in order that they may be easily interpretable. For evaluation
of the results a distance of 4 or less was taken to represent a tendency to form a
complete band (a condition comparable with the dotted bands of riversi, horrescens
and tongae), but in no case was a complete band observed.

The table shows that there is very little tendency to form complete bands and
this is scarcely affected by different treatments though there is some increase in
white scaling at 30-32 C., and males of all series show a little more white scaling
than females ; males of the different stock resemble those bred at 3O-32 C.

(i) Table XX : Specialized setae on the basal lobe

TABLE XX. Aedes pseudoscutellaris

Basal lobe of coxite of male Number of specialized setae

Number of specialized setae*

Number of Number of , -*- >

specimens lobes 23456

2 16 46 30 6

20 54 24 2

28 54 18

Series F, i5-i6 C.
Series C, ig-22 C.
Series B, 25-28 C.
Series E, 3o-32 C.
Series D, 25-28 C.,

25
25
25
25
25

50
50
5
50
r ,o

2 22 52 20 4

2 14 50 26 8

sea-water

Series H, 25-28 C., . 10 . 20 . 35 60 5

different stock

* Number of lobes expressed in percentages.

The specialized setae on the basal lobe of the coxite are of particular interest as
they are the most important character by which pseudoscutellaris is distinguished
from polynesiensis (in which they are absent). Large variations in the number of

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 409

these would cast considerable doubt on their value in identification and delimitation
of species.

As already recounted, the basal lobe was mounted in lateral view (see Fig. 5).
The specialized setae are broad and well defined, but at the distal end of the row
there may be one or two more slender setae which appear slightly flattened and it is
often difficult to decide whether these should be included in the count or not. In
order to avoid as far as possible any differences between series due to different
interpretations of such setae at different times, a slide of genitalia mounts (2 or
3 per slide) was examined from each series in turn.

The results are shown in Table XX. They have been examined statistically
and the difference between the series are not significant.

It may be added that although no records were made, there appeared also to be
very little variation in the arrangement of the non-specialized setae on the basal
lobe. An occasional specimen has a scale amongst the setae towards the tip, but
this is an aberration that occurs also in polynesiensis (see Fig. 7) .

(5) Summary of Experimental Results

The following is a summary of the results obtained from a biometrical study of
specimens of Aedes pseudoscutellaris reared under controlled conditions in which
one factor of the larval environment was varied.

1. Size of adults decreases with increase in temperature or salinity of the larval
environment.

2. In general, the amount of white scaling increases with rise in temperature ;
conversely the amount of dark scaling increases with fall in temperature. The
reverse has not been observed but some characters are not affected.

3. Increase in temperature effects an increase in white scaling in the following
positions : under side of proboscis ; anterolateral margin of scutum ; fore and
mid tarsal segments III, fore tarsals IV-V, hind tarsals I-IV ; abdominal tergites V
and VI. Many of these characters follow a parallel gradient to the temperature ;
others are affected only at low or at high temperatures, or to different degrees in
the two sexes. The dark ventral line on the hind femur increases in extent with
decrease in temperature.

4. A larval environment of' sea-water effects a decrease in white scaling under
the proboscis of males ; and an increase in white scaling on fore tarsal segments
IV-V and hind tarsal II of females and on hind tarsals III-IV of both sexes. Other
scale characters are apparently not affected.

5. The extent of white scaling on either side of the prescutellar bare area and the
length of the anterior white streak on the hind femur do not appear to vary in relation
to larval environment.

6. Males tend to have more white scaling on the under side of proboscis, antero-
lateral margin of scutum and lateral patches of abdominal tergites, than do females
reared in the same conditions.

7. Females tend to have more white scaling on fore tarsal segments III-V, mid
tarsal III and hind tarsals III-IV, than do males reared in the same conditions.

410 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

8. A white streak under the proboscis may be absent or variable in one stock and
well developed in another stock of the same species.

9. The characteristic white scaling on the anterolateral margin of scutum is
retained in varied environments.

10. Although the greatest lengths of white bands on the hind tarsi vary with
larval environment, the total variation is insufficient to affect seriously their use
as taxonomic characters.

n. Low temperatures may produce partial or complete interruption of the white
bands on the hind tarsal segments.

12. The characteristic markings of the abdominal tergites are retained in varied
environments.

13. The number of specialized setae on the basal lobe of the male coxite is not
varied significantly by alteration of larval environment.

(6) Conclusions

It would be unsafe to regard the results of these studies on a small population
as applicable to pseudoscutellaris as a whole, let alone to other species of the scutellaris
subgroup. Nevertheless, they do support the specific status given to members of
the scutellaris subgroup by showing that, although currently accepted diagnostic
characters can be varied by environmental conditions, with few exceptions the range
of variation is not of the same magnitude as the differences between species hi the
same character. They also show that these characters remain relatively constant
through numerous generations. In particular, the lack of variation in the basal
lobe of the male coxite is evidence of the value of this structure in defining species.

The experiments suggest that two characters should be used with some caution
for delimiting species since pseudoscutellaris can exhibit both extremes of their
development. Complete interruption to the white bands on hind tarsal segments
II-IV can be produced by low temperatures of larval environment. The presence
or absence of a white streak under the proboscis may be due to differences in here-
ditary constitution, but there is also, in males, a greater tendency for a streak to
be present in specimens reared at higher temperatures.

It is possible that in other species of the scutellaris subgroup, these characters
may be genetically fixed. In this connection it may be noted that Waddington
(1952) has shown experimentally, in Drosophila melanogaster , that in the course of
selection, a genetic constitution may be synthesized, which under normal conditions
produces the same effect as was originally found only as a response to the stimulus
of an abnormal environment.

IV. ACKNOWLEDGMENTS

Grateful acknowledgment is made of generous interest and help received from
many quarters, without which these studies could have made little progress.

Thanks are due for their advice and assistance to members of the staff of the
Department of Zoology, University of Cambridge, where the work was carried out,
and in particular to Professor V. B. Wigglesworth for facilities of the Sub-department

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 411

of Entomology, to Mr. F. R. Parrington for laboratory accommodation in the Museum
of Zoology, and to Dr. John Smart for his stimulating supervision and constructive
criticisms ; I am indebted also to Miss M. Haslop for her technical assistance.
Dr. F. J. Anscombe of the Statistical Laboratory advised on mathematical aspects
of the studies, and Mrs. E. H. Lawrie tested some of the results statistically.

Portion of the work was done in the British Museum (Natural History), and I am
grateful to Mr. N. D. Riley for facilities in his department and to Mr. P. F. Mattingly
for the loan of specimens, for practical help and profitable discussions and for
generously undertaking to see this paper through the press.

Thanks are due to Sir Philip Manson-Bahr for stock from his colony of pseudo-
scutellaris ; to Sir Rickard Christophers for much valuable advice, particularly in
establishing the colony in Cambridge ; to Professor P. A. Buxton for the loan of
specimens from the London School of Hygiene and Tropical Medicine ; to Dr. E. C.
Zimmerman for the loan of his collection of mosquitoes from south-east Polynesia ;
to Dr. A. R. Woodhill for specimens from his laboratory colonies ; to Messrs. B. A.
O'Connor and I. M. Rakai for eggs of species of the scutellaris subgroup from Fiji ;
and to Dr. and Mrs. Marshall Laird for numerous specimens from Pacific Islands
collected by them in the course of a project initiated by the Royal New Zealand
Air Force and supported by a grant from the N.Z. Department of Scientific and
Industrial Research. Invaluable help has been received from Dr. Alan Stone who
answered many queries and supplied detailed notes on specimens in the U.S.
National Musum.

Finally, I am indebted to Queensland's National Mosquito Control Committee
for granting me leave of absence to pursue these studies, and allowing some additional
work to be completed in the Department of Entomology, University of Queensland.

V. BIBLIOGRAPHY

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1934- Fauna of British India, Diptera. 5. Family Culicidae, Tribes Megarhinini and

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412 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

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& BONNE-WEPSTER, J. 1947. The geographical distribution of the mosquitoes of the

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BUXTON, P. A. 1927. Sur les moustiques de Tahiti et du groupe des iles de la Societ6. Bull.
Soc. Etud. oceanien 21 : 306-308.

1935- Insects of Samoa. Part 9, Fasc. 2. (Summary : pp. 33-104.) London : Brit.

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& HOPKINS, G. H. E. 1927. Researches in Polynesia and Melanesia. Parts 1-4. London :

Lond. Sch. Hyg. & Trop. Med.
BYRD, E. E., ST. AMANT, L. S., & BRCMBERG, L. 1945. Studies on filariasis in the Samoan

area. Nav. Med. Bull. Wash. 44 : 1-20. [Cited in Rev. appl. Ent. 34 : 38-39, 1946.]
CHEESMAN, L. E. 1927. Notice sur les moustiques de Tahiti. Bull. Soc. Etud. oceanien 19 :

245-247-
DAGGY, R. H. 1944. Aedes scutellaris hebrideus Edwards ; probable vector of dengue in

New Hebrides. War Med. 5 : 292-293. [Cited in Trop. Dis. Bull. 42 : 32, 1945.]
DOLESCHALL, C. L. 1858. Derde Bijdrage tot de kennis der Dipteren Fauna van Nederlandsch

Indie. Natuurk. Tijdschr. Ned.-Ind. 17 : 73-128.
EDWARDS, F. W. 1917. Notes on Culicidae with descriptions of new species. Bull. ent.

Res. 7 : 201-229.
1924. A synopsis of the adult mosquitoes of the Australasian region. Ibid. 14 : 351-401.

1926. Mosquito notes : VI. Ibid. 17: 101-131.

1927. Diptera Nematocera from the South Pacific collected by the "St. George"

Expedition, 1925. Ann. Mag. nat. Hist. (9) 20 : 236-244.

1929. Mosquito notes : VIII. Bull. ent. Res. 20 : 321-343.

1932. Diptera. Family Culicidae. Genera Insect. 194 : 1-258.

1933- Mycetophilidae, Culicidae and Chironomidae and additional records of Simuliidae

from the Marquesas Islands. Bull. Bishop. Mus. Honolulu, 114 : 85-92.

1935- Mosquito notes : XII. Bull. ent. Res. 26 : 127-136.

1941. Mosquitoes of the Ethiopian region. III. Culicine Adults and Pupae, Pp. 499.

London : Brit. Mus. (Nat. Hist.).
FARNER, D. S. 1945. A new species of Aedes from the Caroline Islands (Diptera, Culicidae).

Proc. biol. Soc. Wash. 58 : 59-61.

& BOH ART, R. M. 1944. Three new species of Australasian Aedes (Diptera, Culicidae).

Ibid. 57 : 117-122.

1945- A preliminary revision of the scutellaris group of the genus Aedes. Nav.

med. Bull., Wash. 44 : 37-53.
DICKE, R. J., SWEET, G., ISENHOUR, L., & HSAIO, T. Y. 1946. The distribution of

mosquitoes of medical importance in the Pacific area. Nav. Med. 983. Washington :

Bureau of Medicine and Surgery, Navy Dept.
FORBES, J., & HORSFALL, W. R. 1946. Biology of a pest mosquito common in New Guinea.

Ann. ent. Soc. Amer. 39 : 602-606.
FORD, E. B. 1949. Early stages in allopatric speciation, [in] Jepson, G. L., Mayr, E. &

Simpson, G. G., Genetics, Palaeontology and Evolution. Princeton, N.J.
HARVEY, H. W. 1928. Biological Chemistry and Physics of Sea Water. Cambridge University

Press.
HILL, G. F. 1925. The distribution of Anopheline mosquitoes in the Australian region, with

notes on some Culicine species. Proc. Roy. Soc. Viet. 37 : 61-77.

HULL, W. B. 1952. Mosquito survey of Guam. U.S. Forces med. J. 3 : 1287-1295.
HUXLEY, J. 1942. Evolution : The Modern Synthesis. London.
JACHOWSKI, L. A., & OTTO, G. F. 1952. Filariasis in American Samoa. II. Evidence of

transmission outside of villages. Amer. J. trop. Med. Hyg. 1 : 662-670.

A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP 413

KNIGHT, K. L., BOHART, R. M., & BOHART, G. E. 1944. Keys to the Mosquitoes of the Austra-
lasian Region. Washington : National Research Council.

& HULL, W. B. 1952. The Aedes mosquitoes of the Philippine Islands. II. Subgenera

Skusea, Christopher siomyia, Geoskusea, Rhinoskusea and Stegomyia (Diptera, Culicidae).
Pacif. Sci. 6 : 157-189.

& HURLBUT, H. S. 1949. The mosquitoes of Ponape Island, eastern Carolines. /. Wash.

Acad. Sci. 39 : 20-34.
& ROZEBOOM, L. E. 1946. The A edes (Stegomyia) albolineatits group (Diptera, Culicidae).

Proc. biol. Soc. Wash. 59 : 83-98.
LAIRD, M. 1946. A report on material obtained during a mosquito survey at Palmalmal,

New Britain. Trans, roy. Soc. N.Z. 75 : 465-478.

1952. Notes on the mosquitoes of Nissan Island, Territory of New Guinea. Pacif. Sci.

6 : 151-156.

LEE, D. J. 1944. An Atlas of the Mosquito Larvae of the Australasian Region. Tribes Mega-

rhinini and Culicini. Melbourne : Australian Military Forces (Restricted).
LEVER, R. J. A. W. 1945. Entomological notes. Agric. J. Fiji, 16 : 46-49.
LIEFTINCK, M. A. 1949. Synopsis of the Odonate fauna of the Bismarck Archipelago and

the Solomon Islands. Treubia, 20 : 319-374.
MACKERRAS, I. M. 1946. Transmission of dengue fever by Aedes (Stegomyia} scutellaris Walk.

in New Guinea. Trans. R. Soc. trop. Med. Hyg. 40 : 295-312.
MANSON-BAHR, P., & MUGGLETON, W. J. 1952. Further research on filariasis in Fiji. Trans.

R. Soc. trop. Med. Hyg. 46 : 301-326.
MARKS, E. N. 1947. Studies of Queensland mosquitoes. Part I : The Aedes (Finlaya) Aoc/w

group, with descriptions of new species from Queensland, Bougainville and Fiji. Pap.

Dep. Biol. Univ. Qd., 2 (5) : 1-66.
1950. Note in proceedings of meeting. Proc. R. ent. Soc. (C), 15 : 51.

i95ia. Mosquitoes from south-eastern Polynesia. Occ. Pap. Bishop Mus. 20 : 123-130.

19516. The vector of filariasis in Polynesia : a change in nomenclature. Ann. trop.

Med. Parasit. 45 : 137-140.

MATTINGLY, P. F. 1953. The subgenus Stegomyia (Diptera, Culicidae) in the Ethiopian

Region. II. Distribution of species confined to the East and South African sub-region.

Bull. Brit. Mus. (Nat. Hist.) Ent. 3 : 1-65.

MAYR, E. 1942. Systematics and the Origin of Species. New York.
, LINDLEY, E. G., & USINGER, R. L. 1953- Methods and principles of systematic

zoology. New York.
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the Cook Islands with notes on the breeding habits of Stegomyia pseudoscutellaris. Trans.

R. Soc. trop. Med. Hyg. 19 : 138-149.
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V. Int. Congr. Ent. 2 : 431-450.
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Med. 4 : 1-57.
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Agric. Fiji, 22 : 135.
PENN, G. H. 1947. The larval development and ecology of Aedes (Stegomyia) scutellaris

(Walker, 1859) in New Guinea. /. Parasit. 33 : 43-50.

1949. The pupae of the mosquitoes of New Guinea. Pacif. Sci. 3 : 3-85.

PERRY, W. J. 1944. Keys to the Mosquitoes of the Neiv Hebrides. U.S. Navy (restricted).
1948. The dengue vector on New Caledonia, the New Hebrides and the Solomon Islands.

Amer. J. trop. Med. 28 : 253-259.
1949. The mosquitoes and mosquito-borne diseases of the Treasury Islands (British

Solomon Islands). Ibid. 29 : 747-758.
1950. Biological and cross-breeding studies on Aedes hebrideus and Aedes pernotatus

(Diptera, Culicidae). Ann. ent. Soc. Amer. 43 : 123-136.

4 i4 A REVIEW OF THE AEDES SCUTELLARIS SUBGROUP

PRASHAD, B. 1918. The thorax and wing of the mosquito (Anopheles}. Indian J. med.

Res. 5 : 610-640.
REEVES, W. C., & RUDNICK, A. 1951. A survey of the mosquitoes of Guam in two periods

in 1948 and 1949 and its epidemiological implications. Amer. J. trap. Med. 31 : 633-658.
SIMPSON, G. G. 1944. Tempo and Mode in Evolution. New York.
SMITH-WHITE, S. 1950. A note on non-reciprocal fertility in matings between subspecies of

mosquitoes. Proc. Linn. Soc. N.S.W. 75 : 279-281.

SNODGRASS, R. E. 1935. Principles of Insect Morphology. New York.
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ent. Soc. Wash. 49 : 85.
& BOHART, R. M. 1944. Studies on mosquitoes from the Philippine Islands and

Australasia (Diptera, Culicidae). Proc. ent. Soc. Wash. 46 : 205-225.
& FARNER, D. S. 1945. Further notes on the Aedes scutellaris group (Diptera, Culicidae).

Proc. biol. Soc. Wash. 58 : 155-162.
TAYLOR, F. H. 1914. Contribution to a knowledge of Australian Culicidae No. i. Proc.

Linn. Soc. N.S.W. 39 : 454-468.
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Brit. Mus. (Nat. Hist.).

19106. The Culicidae of Fiji, including two new species. Entomologist, 43 : 155-159.

TRAVIS, B. V. 1947. Relative efficiency of six species of mosquitoes from Guam, M.I., as

developmental hosts for Dirofilaria immitis. J. Parasit. 33 : 142-145.
WADDINGTON, C. H. 1952. Selection of the genetic basis for an acquired character. Nature,

169 : 278.
WALKER, F. 1859. Catalogue of the Dipterous insects collected in the Aru Islands by Mr.

A. R. Wallace, with descriptions of new species. /. Linn. Soc. (Zool.) 3 : 77-131.
1861. Catalogue of the Dipterous insects collected at Dorey, New Guinea, by Mr. A. R.

Wallace, with descriptions of new species. Ibid. 5 : 229-254.
WOODHILL, A. R. i949a. A new subspecies of Aedes (Stegomyia) scutellaris Walker (Diptera,

Culicidae) from Northern Australia. Proc. Linn. Soc. N.S.W. 74 : 140-144.
19496. A note on the experimental crossing of Aedes (Stegomyia) scutellaris scutellaris

Walker and Aedes (Stegomyia) scutellaris katherinensis Woodhill (Diptera, Culicidae).

Ibid. 74 : 224-226.
1950. Further notes on experimental crossing within the Aedes scutellaris group of species

(Diptera, Culicidae). Ibid. 75 : 251-253.
1954- Experimental crossing of Aedes (Stegomyia) pseudoscutellaris Theobald and Aedes

(Stegomyia) polynesiensis Marks (Diptera, Culicidae). Ibid. 79 : 19-20.
ZEUNER, F. E. 1943. Studies in the systematics of Troides Hubner (Lepidoptera, Papilionidae)

and its allies ; distribution and phylogeny in relation to the geological history of the

Australasian Archipelago. Trans, zool. Soc. Lond. 25 : 107-184.
ZIMMERMAN, E. C. 1948. Insects of Hawaii, i . Introduction. Honolulu.

PRES1

24 NOV 1954

PLATE 18

Comparative figures of species of the Aedes
(Stegomyia) scutellaris group, arranged approx-
imately in their geographical relationships.

All drawings, except basal lobe, diagrammatic
O - from published descriptions and figures.
specimens seen, basal lobe from published figures.
O - specimens seen, basal lobe from freehand drawings.

4

C;

riversi R. Bohart & Ingram

guamensis Farner &. R. Bohart <D

I

sp. from Andaman Is.

paullusi Stone & Forncr <D

hensilli Farner O

andrewsi Edwards <D
KEY FIGURE

Basal lobe of left coxlte of j
T - tergal (ventral) view
S sternal (dorsal) view
L - lateral (Inner) view

Segments I V of Hind tarsus
'showing greatest width of white

bands and interruption of bands

by dark scales.

ero-lateral

Lateral view of
abdominal terglteV

olorensis Bonne-Wepster & Brug

scutelloris scutelloris (Walker)

4

scutellaris katherinensis Woodhill

pernotatus Farner i R. Bohart <D

scutoscriptus R. Bohart & Ingram

siscuteilaris Farner R. R. Bohart

hakonssoni Knight &. Hurlbut <D

sp. from Rotuma I

pseudoscutellaris (Theobald)

marjhallensis Stone &. R. Bohart O

4

polynesiensis Marks

gurneyi Stone & R. Bohart

horrescens Edwards

tongae Edwards

^SENTED

24NOV 1954

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

<- - <::;<

2 A NOV 1954

A NEW GENUS AND SOME NEW

SPECIES OF THE CHAULIODINI

(MEGALOPTERA)

D. E. KIMMINS

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 3 No. n

LONDON: 1954

A NEW GENUS AND SOME NEW SPECIES
OF THE CHAULIODINI (MEGALOPTERA)

BY

D. E. KIMMINS
'

Pp. 415-444 ; 22 Text-figures

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 3 No. n

LONDON : 1954

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

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

This paper is Vol. 3, No. no/ the Entomological
series.

PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM

Issued November, 1954 Price Eight Shillings

A NEW GENUS AND SOME NEW SPECIES OF

THE CHAULIODINI (MEGALOPTERA), WITH

NOTES ON CERTAIN PREVIOUSLY

DESCRIBED SPECIES

By D. E. KIMMINS

SYNOPSIS

The greater part of this paper is devoted to descriptions of eleven new species or subspecies
and a new genus from Oriental, Australian and Chilean localities. The types of some species
described by Walker, McLachlan and Weele have been re-examined, and new figures of genitalia
prepared, and an artificial key for the identification of the genera of the Chauliodini is given.

THE tribe Chauliodini may be distinguished by the following diagnosis : Antennae
moniliform, serrate or pectinate in the male, moniliform, subserrate or, more rarely,
pectinate in the female. Normally not more than three cross-veins between the
radius and its sector. In the hind wing the basal r-m cross-vein is present, obsoles-
cent or entirely absent. Cuj with only one branch beyond the inter-cubital cross-
vein.

Male genitalia with only one pair of appendages. Ninth tergite moderately
sclerotized, somewhat hood-shaped. Sternite generally less sclerotized than tergite,
forming a variously shaped subgenital plate, whose apical margin is often produced
in a transparent triangular lobe. Tenth tergite forming a pair of anal plates or
claspers, differing in shape in the various species and variously armed on the inner
surface with black spinules or comb-like teeth. There is a group of trichobothria
forming a more or less convex wart on the outer surface. Tenth sternite generally
membranous although occasionally (Ctenochauliodes) it forms a sclerotized plate.
Aedeagus hinged to the lower basal angle of the ninth tergite, trough- or scoop-like,
simple or bifid at its apex.

$ GENITALIA. Eighth sternite more or less sclerotized, sometimes produced
apically as a subgenital plate. Ninth tergite deep, its lateral margins produced
downwards almost across the segment ; to their lower angles are attached a pair of
foliate lateral gonapophyses. Ninth sternite apparently membranous. Tenth
segment forming a pair of cercoid anal plates, of varying form, each with a group of
trichobothria.

The known distribution of the tribe is somewhat discontinuous, although further
collecting and study may cause us to modify our conclusions. There appears to be
a small Antarctic element consisting of two genera, with representatives in Aus-
tralia, New Zealand and Chile, one of the genera also possibly extending northward
into California. There are two genera occurring in South Africa, one of which is

ENTOM. Ill, ii. 29

418

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

fairly closely related to Protochauliodes from Australia and Chile and the other has
a near relative in Madagascar.

The Nearctic region contains at least four genera in addition to possible representa-
tives of the Antarctic genus referred to above. The Oriental region appears to be
richest in described species and subspecies and contains four genera, of which three
spread northward into the Manchurian-Japanese sub-region of the Palaearctic
fauna. The majority of the species occur in the Indo-Chinese sub-region.

Certain genera exhibit sexual dimorphism in the antennae and while such charac-
ters will serve to separate the males genetically, it has not been easy to find vena-
tional characters which will serve equally for both sexes. The wing venation of
the tribe is rather uniform, although at the same time subject to aberration. In
the generic key, separate male and female characters have been used to distinguish
Protochauliodes from Neohermes, but with Parachauliodes and Neochauliodes I have
failed to find a satisfactory character to separate the females. It should be pointed
out that in both sexes the abdomen shrinks considerably in drying and that the
form of the genitalia can often only be clearly made out after removal and treatment
in KOH solution. All the genitalia drawings in this paper have been made from
dissections so treated.

KEY TO GENERA OF CHAULIODINI

i. Posterior branch of Rs in both wings forked ; anterior branch of M in hind wings

forked ......... DYSMICOHERMES Munroe.

-. Posterior branch of Rs in both wings simple ; anterior branch of M in hind wings

forked or simple ............ 2.

3A 2A
Archichauliodes

3A 2A
Protochauliodes

2A

IA

Chauliodes

Platychauliodes

3A 2A
Anachauliodes

Dysmicohermes

Platychauliodes

Ctenochauliodes

FIG. i. Anal area of fore wing, a, and anterior basal area of hind wing, p.

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI 419

2. In fore wing, anterior branch of 2A partly fused with lA, this branch appearing

to arise irom lA ; a pigmented area immediately basad of this apparent origin . 3.
-. In fore wing, anterior branch of 2 A not fused with i A, but either it or the stem of 2 A

connected to lA by a cross- vein . . . . . . . . .5.

3. Anterior branch of M in hind wing forked . . . . . . . .4.

-. Anterior branch of M in hind wing simple . . TAENIOCHAULIODES Esben-Petersen.

4. Male antenna without a whorl of erect hairs on each segment ; lateral gonapophyses

of ninth segment of female with a small apical cercus. No cross-vein in marginal
fork of R 3 in fore wing (sometimes absent in Neohermes) . PROTOCHAULIODES Weele. 1
-. Male antenna with erect hairs on each segment other than the basal ; lateral gona-
pophyses of female without small apical cercus : cross-vein in marginal fork of
R 3 in fore wing generally present ...... NEOHERMES Banks.

5. Species from Australia, New Zealand, Chile, S. Africa or Madagascar . . .6.
-. Species with a Nearctic or Oriental distribution ....... 7.

6. Species from Australia, New Zealand or Chile ; wings generally long and narrow

ARCHICHAULIODES Weele. 2
-. Species from S. Africa or Madagascar ; wings relatively broader

PLATYCHAULIODES Esben-Petersen. 3

7. Nearctic species ............ 8.

-. Oriental species ............ 9.

8. Wings mainly blackish brown, with opaque creamy white markings NIGRONIA Banks.
-. Wings greyish hyaline, venation with fuscous spots . . CHAULIODES Latreille.

9. lA in fore wing with three or four branches . . . ANACHAULIODES gen. n.
-. lA in fore wing with only two branches ........ 10.

10. In fore wing, fork of 2 A with a definite footstalk . . . . . .11.

-. In fore wing, fork of 2A sessile ..... CTENOCHAULIODES Weele.

11. Male antenna serrate, anal plates horizontally bilobed . . PARACHAULIODES Weele.
-. Male antenna pectinate, anal plates simple .... NEOCHAULIODES Weele.

Archichauliodes guttiferus (Walker).
Figs. 2, 3.

Hermes guttiferus Walker, 1853, List Neur. Ins. B.M. 2 : 204.

Chauliodes guttiferus (Walker), McLachlan, 1867, /. Linn. Soc. Zool. 9 : 260 ; McLachlan, 1869,

Ann. Mag. Nat. Hist. (4) 4 : 39. Weele, 1907, Notes Leyden Mus. 28 : 252, figs. 19, 20 ;

pi. 4, fig. 2.
Archichauliodes guttiferus (Walker), Weele, 1910, Cat. Coll. Zool. Selys, fasc. 5(i) : 48. Tillyard,

1926, Ins. Aust. N. Z. : 313.

Examination of the material over this name in the British Museum (N.H.) revealed
the presence of three species with almost identical whig markings but entirely
distinct genitalia in both sexes. This raised the question to which of the three

1 The unique $ type of P. dubitatus (Walker) differs from the true Protochauliodes in having the basal
r-m cross-vein completely lacking in both hind wings. In view of the fact that it is a unique, from an
unknown locality (variously suggested to be New Zealand or S. America) and possibly aberrant, it is
proposed to leave it for the present in Protochauliodes. One must not overlook the possibility of its
being an aberrant Neohermes, the transverse fuscous band in the basal part of the fore wing at the level
of the cubital fork certainly resembling that found in Neohermes.

2 In Archichauliodes diver sus (Walker) from New Zealand the basal r-m cross- vein in the hind wing
appears to be very unstable and often obsolete.

3 Paulian's figure of the venation of his Madachauliodes torrentialis shows no basal r-m cross-vein in
the hind wing, and would thus run out in this key to Platychauliodes , from which it appears to differ
in a longer fork to Cu! in both wings.

420

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

should the name guttiferus Walker be applied. Walker's type has for many years
lacked antennae and abdomen and I have seen no statement in print as to its sex.
The female example figured by Weele (1907, pi. 4, fig. 2) is certainly not the type.

FIG. 2. Archichauliodes guttiferus (Walker). Male genitalia. a, lateral ; b, left anal plate,
dorsal ; c, ninth sternite and aedeagus, ventral.

FIG. 3. Archichauliodes guttiferus (Walker). Female genitalia. a, lateral ; b, eighth
sternite, ventral ; c and d, anal plate and lateral gonapophyses, lateral, from another
specimen.

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI 421

The antennae and abdomen may even have been missing at the time of the original
description, since neither part is mentioned and Walker quotes the length of the
body with a question-mark.

Before attempting a decision as to the identity of Walker's guttiferus, preparations
of the abdomens of a number of females (excluding A. deceptor sp. n. from Queens-
land) were made and these, while showing some individual variation, seemed to fall
into two groups on the structure of the eighth abdominal sternite. Comparison
of the wing pattern of these two groups then revealed some not very easily defined
differences, and by comparing Walker's type with these two groups, it appeared to
resemble most closely the group the males of which agreed with the figures of the
genitalia given by Weele (1907, figs. 19-20). It is fortunate that we are thus able
to retain Weele's interpretation of this species. To facilitate identification of these
very similar-looking species I am describing and figuring Walker's guttiferus as well as
the two new species. Examination of long series of Australian material may perhaps
prove that what I have considered to be individual variations are really specific
differences, but this is a matter which can only be decided by Australian students.-

Body-colour fuscous. Antennae, palpi and bases of ocelli piceous. Pronotum
a little longer than broad, tapering slightly anteriorly. Legs uniformly fuscous.
Abdomen reddish black, anal appendages piceous. Wings elongate, apices ovate ;
membrane slightly brownish hyaline, venation pale fuscous. Fore wing lightly
speckled with small brownish dots between the veins, and with rather larger ones at
the origin of Rs, and along the costal margin before and after the pterostigma. Hind
wing with a few brownish dots in the apical part of the wing, and with four larger
spots, two along the costal margin (one at each end of the pterostigma) and the
other two between Rs and M, surrounding the f acetic spots. The basal of these
spots usually overlaps both Rs and M. The hind wing spots sometimes enclose
paler areas, giving them a pupillate appearance.

$ GENITALIA. Ninth sternite about as long as tergite, forming a broad subgenital
plate. Anal plates about one and one-half times as. long as ninth segment. From
the side, each is slender, tapering gradually to apex and slightly down-curved, with
a group of trichobothria at its base. From above it is broad at its base, abruptly
narrowed about midway to half its width and then gradually narrowing to apex,
inner margin spinose. Aedeagus with an arched base, centre of its apical margin
produced in an asymmetric, obliquely truncate lobe. There appears to be some
individual variation in the length and shape of this lobe.

$ GENITALIA. Eighth sternite divided longitudinally into two broad sclerites,
linked by membrane. Ninth tergite deep, lower angles obliquely truncate ; lateral
gonapophyses short, sub-quadrate, upper margin convex, apical margin slightly
concave, upper apical angle more or less produced. Anal plates long, slender.

Length of fore wing, 23-30 mm.

Holotype (lacking antennae and abdomen, sex doubtful, locality unknown)
in Brit. Mus. (N.H.). Additional examples in this collection from AUSTRALIA :
New South Wales, Upper Murrumbidgee River, Rule's Point, 4,450 ft., 20.xii.i934
(R. J. Tillyard) and Mt. Kosciusko, 2,700 ft., 22.1.1885, 16.1.1888. Victoria,
Melbourne.

422 A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

Archichauliodes plomleyi sp. n.
Fig. 4.

AUSTRALIA: New South Wales, Mt. Irvine, 2,300 ft., 25.xii.iQ34 (N. J. B.
Plomley), holotype $, allotype $ and 2 $ paratypes ; Heathcote, bred 26.xii.i9i6,
(R. J. Tillyard), i $ paratype ; no other data, Saunders, 68-3, i paratype (without
abdomen).

Victoria, Melbourne, i paratype (lacking apex of abdomen). Australia, no other
data, 3 $, i ? (lacking abdomen), paratypes. All the above examples in Brit.
Mus. (N.H.).

FIG. 4. Archichauliodes plomleyi sp. n. Genitalia, male a-d ; female e-g. a, lateral ;
b, left anal plate, dorsal ; c, ninth sternite, ventral ; d, aedeagus, dorsal ; e, lateral ;
/, eighth sternite, ventral ; g, anal plate and lateral gonapophyses of another example.

General appearance as in A. guttiferus (Walker). The fore wing spots are perhaps
a little less dense. Hind wing with four or five large fuscous spots, usually smaller
than in guttiferus and with the basal spot usually not or only slightly overlapping
Rs and M. Spots not pupillate. Fifth spot, when present, over the inter-cubital
cross-vein.

$ GENITALIA. Ninth sternite rather shorter than the overhanging tergite, forming

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

423

a broad subgenital plate, its apical margin flattened and semi-membranous. Anal
plates short, from above sharply bent inwards, apices clavate and spinose ; there is
a large, rounded, basal group of trichobothria. Aedeagus forming a truncate basal
arch, from which arise two stout, up-curved, unequal spines, one acute, the other
truncate at apex.

$ GENITALIA. Eighth tergite entire, sclerotized, from beneath parallel-sided,
apical angles produced in rounded lobes. Ninth tergite deep, lateral gonapophyses
forming flattened, ovate or pyriform valves. Anal plates short, conical, varying
somewhat in width at base.

Length of fore wing, 24-28 mm.

Holotype <, allotype $, each with abdomen mounted in Canada balsam. Apart
from the differences already mentioned above, A. plomleyi may be separated from
guttiferus by the short, incurved anal plates and the two asymmetric spines of the
aedeagus in the male and by the entire eighth sternite with rounded apical lobes and
the shorter anal plates in the female.

The collector informs me that Mt. Irvine is one of the Western mountain range,
about seventy miles west of Sydney.

Archichauliodes deceptor sp. n.

Fig. 5-

AUSTRALIA: Queensland, Toowoomba, 2,000 ft., io.xii.i884, i $, holotype;
no other data (F. P. Dodd), i $, allotype, i <j> paratype. All in Brit. Mus. (N.H.).

FIG. 5. -Archichauliodes deceptor sp. n. Genitalia, male a-d, female e. a, lateral,

b, left anal plate, dorsal ; c, aedeagus, ventral ; d, ninth sternite, ventral ; e, lateral.
ENTOM. Ill, n. 2Q

424 A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

In general appearance scarcely distinguishable from A . guttiferus and A . plomleyi.
The femora and tibia are generally lighter than the tarsi, with a darker patch at the
apex of the femora.

c GENITALIA. Ninth tergite hooded, its apical margin slightly convex. Ninth
sternite about as long as tergite, apical margin rounded, not membranous. Anal
plates short, stout, somewhat incurved, with a large lateral group of trichobothria.
Aedeagus long, slender, up-curved, not asymmetric, apex rounded from the side,
pointed from beneath.

$ GENITALIA. Eighth sternite sclerotized, pigmented, before the apex elevated
in two low, rounded, hairy mounds. Ninth segment moderately deep, lateral
gonapophyses about as long as segment, rather narrow, directed obliquely upwards,
apex bent somewhat tailward and rounded. Anal plates short, conical, each with
a large lateral group of trichobothria.

Length of fore whig : $ 25 mm. ; $ 27 mm.

Holotype $, allotype $ each with abdomen mounted in Canada balsam. In both
male and female genitalia this species is more nearly related to plomleyi than gutti-
ferus. The male differs from the former in its less incurved anal plates and in the
long single spine of the aedeagus. The female differs in the less developed eighth
sternite and in the shape of the anal plates and lateral gonapophyses.

Archichauliodes australica sp. n.
Fig. 6.

AUSTRALIA : New South Wales, Upper Murrumbidgee River, Rule's Point,
4,450 ft., 20.xii.i934 (R. J. Tillyard), i $, holotype, in Brit. Mus. (N.H.).

Body-colour dark fuscous. Mandibles pale yellowish. Antennae filiform, the
segments very slightly inflated on inner side but scarcely subserrate. Lateral
margins of head behind eyes with a fringe of short silvery hair. Abdomen with
obscure, transverse, fulvous lines.

Wings elongate, acutely rounded at apices. Membrane greyish hyaline with
brownish venation, fore wing with three conspicuous brown markings in the stigmal
area and with numerous small brown spots along the veins and encroaching on the
membrane. Hind wing practically without markings other than the brownish
stigmal area. Venation much as in guttiferus but costal cross-veins in fore wing
less numerous and straighter. In the right wing there is an aberrant cross-vein
connecting the basal r-m cross-vein with M.

c GENITALIA. Ninth tergite large, hooded, its apical dorsal margin with a wide
shallow triangular excision ; sternite short, broad, apical margin forming a flat-
tened ellipse. Anal plates stout, in side view with a rounded lobe on lower margin
about midway, then tapering to a rounded apex. The group of trichobothria are
situated on a projecting ovate wart close to the base. From above the inner
margin produced in a triangular lobe, set with spinules. Aedeagus forming an arched
plate, in dorsal view the basal arms spirally twisted about midway. The centre of
the arch is produced in two slightly incurved plates, their upper basal margins fused
and elevated in a triangular lobe as seen from the side.

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

425

Length of fore wing, 25 mm.

Type (J with abdomen mounted in canada balsam. This specimen may well be
one of those referred to by Tillyard (1926, Ins. Aust. N.Z. : 313) " the mountain
forms [of A. guttifems] closely resemble the New Zealand species." After the
publication of this work he took typical A. guttiferus at the same locality as this
new species, which differs from A . guttiferus (Walker) in the almost complete absence
of spots, apart from those along the fore wing veins, the more pointed wings, less
moniliform antennae and shorter, blunter anal plates.

FIG. 6. Archichauliodes australica sp. n. Male genitalia. a, lateral ; b, left anal plate,

dorsal ; c, aedeagus, dorsal.

Archichauliodes chilensis sp. n.

Figs. 7, 8.

S. AMERICA : Chili (Calvert), i $, holotype, i $, allotype ; (Reed), 3 $ paratypes ;
(McLachlan collection), i (lacking abdomen), paratype. S. Chili, Araucania (R. M.
Middleton), i $, paratype. All in Brit. Mus. (N.H.).

Antennae moniliform in both sexes, dull blackish brown, the two basal segments
shining reddish brown. Body dull reddish or blackish brown, head with a patch
in front of each antenna, and the back of the head, orange-brown. Wings elongate-
elliptical, apices a little less acute in male than in female, membrane pale brownish
hyaline, venation brownish, some of the longitudinal veins in hind wing narrowly
bordered with brownish. In fore wing the membrane is lightly speckled with darker

426

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

brown adjoining the veins, and in the apical half of the wing there is a brownish
streak running longitudinally through the cells. The pterostigma is more heavily
marked with two brownish spots, and there is a brownish cloud over the fork of 2A.
<$ GENITALIA. Ninth sternite from beneath parabolic, with a semi-membranous,
triangular process extending beyond its apex (much as in A. diver sus (Walker)).
Anal plates short and stout, in side view slightly curved downward near the apex,
which is truncate. From above, the apex is rounded and clothed internally with
dense black spinules. Beneath these appendages is the aedeagus, hinged at its
basal angles to the lower angles of the ninth tergite. From above it resembles an
inverted "V," with a blunt, bifid apex, which in the holotype is somewhat asymmetric.

Sc + R.

Ib *- u lo

FIG. 7. Archichauliodes chilensis sp. n. Male wings.

The outer margin of each apical lobe is armed with a small acute tooth. From the
side these lobes are directed obliquely upward.

$ GENITALIA. Eighth sternite more sclerotized than its tergite, somewhat pro-
duced apically in a subgenital plate. Ninth tergite as deep as eighth segment,
mainly sclerotized but with a wide, apical, membranous area simulating an excision.
The lower basal margins of the tergite overlap the eighth sternite ; lateral gona-
pophyses elongate and slightly tapering from base to apex. Ninth sternite mem-
branous. Anal plates short, each rather stout at the base and bearing a large,
rounded group of trichobothria, apex finger-like and somewhat flexible.

Length of fore wing : $ 27 mm. ; $ 29-42 mm.

This species in general appearance resembles Protochauliodes cinerascens (Blan-
chard), but may be separated from it by the stalked condition of the fork of 2 A
in the fore wing and by the more basal position of the first fork of Rs in both wings.

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

427

From A . diversus (Walker) it may be separated by the absence of any brown speckling
along the branches of Rs in the hind wing and by the shape of the anal plates, the
aedeagus beneath them and of the ninth sternite.

FIG. 8. Archichauliodes chilensis sp. n. Genitalia, male a-d, female, e. d, lateral ; b,
left anal plate, dorsal ; c, aedeagus, dorsal ; d, ninth sternite, ventral ; e, lateral.

ANACHAULIODES gen. n.

c?. Antenna pectinate. Wings with apical branches of Rs somewhat curved
posteriorly near apices. In fore wing, lA with three or four branches, fork of 2A
sessile, anterior branch not fused with lA. In hind wing, basal r-m cross-vein
obsolete.

<$ GENITALIA. Ninth segment not more sclerotized than eighth. Tenth segment
forming two large anal plates, apical margin excised, inner surface with a few strong
teeth on a small projection. Aedeagus large, strongly sclerotized, hooked down-
ward.

$ unknown.

Type species, Anachauliodes tonkinicus sp. n.

428 A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

In general appearance, this genus recalls the North American genus Chauliodes,
from which it differs in the more branched lA and the more curved apical branches
of Rs in the fore wing. The latter character recalls Neochauliodes, from which it
differs in the obsolete r-m cross-vein at the base of the hind wing, as well as the
more branched lA in the fore wing.

Anachauliodes tonkinicus sp. n.
Figs. 9, 10.

TONKIN : Ngai-Tio, 4,800 ft., 22^.1924 (H. Stevens), i <, holotype, Brit. Mus.

(N.H.).

FIG. 9. Anachauliodes tonkimcus gen. sp. n. Male wings.

Body-colour light fuscous. Antenna piceous, strongly pectinate. Labrum,
anteclypeus and mandibles fulvous. Pronotum about as broad as long, slightly
narrowed anteriorly. Legs fuscous, femora with a fulvous stripe above. Wings
greyish hyaline, venation of fore wing luteous with fuscous interruptions, hind wing
mainly fuscous. Fore wing with a dark fuscous patch in costal area basad of the
creamy pterostigma, several fuscous spots in subcostal area, a fuscous patch near
base of wing, a paler fuscous cloud across the centre of the wing and with fuscous
irrorations spreading out from the vein markings. Costal cross-veins fuscous.
Hind wing practically without markings.

$ GENITALIA. Ninth segment scarcely more sclerotized than eighth, tergite
somewhat extended in a hood, sternite forming a short, rounded subgenital plate.
Anal plates large, each with a central group of trichobothria. Apical margin of anal

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

429

plate produced in two rounded lobes, separated by an excision. The upper lobe is
the smaller and carries on its lower inner surface a short, dentiferous projection.
Lower lobe about twice as deep as upper. Aedeagus large, heavily sclerotized,
from the side curved downward, with an acute, hooked apex. The upper surface
is produced to form two recurved, triangular plates, from the side appearing as a
hook. From above, the aedeagus arises from a wide, arched base, parallel-sided
as far as the recurved plates, then narrowed and tapering to the down-turned apex.

Length of fore wing, 43 mm.

Type with abdomen mounted in Canada balsam.

FIG. 10. -Anachauliodes tonkinicus gen. sp. n. Male genitalia. a, lateral ; b, left anal
plate, dorsal ; c, aedeagus, dorsal ; d, aedeagus, lateral.

CTENOCHAULIODES Weele

Weele, 1909, Notes Leyden Mus. 30 : 263 ; id., 1910, Coll. Zool. Selys, 5 (i) : 70.
Type species, Chauliodes nigrovenosus Weele, 1907.

Antennae pectinate in female and probably in male also. In hind wing, basal
r-m cross-vein strong and re-connected to M by a cross-vein. In the genitalia of
the presumed male, there is a sclerotized plate beneath the anal plates, probably
the tenth sternite. It is possible that in some other genera (such as Neohermes)
this plate has become fused to the aedeagus.

Ctenochauliodes forcipatus sp. n.

Fig. n.

CHINA : [Szechwan?], Kwanshien, July 1930 (G. M. Franck), $ holotype.

W. CHINA : Chin-Fu-San (W. A. Maw), $ paratype. (Both in Brit. Mus. (N.H.)).

Body-colour reddish brown. Antennae broken, with only two basal segments

43

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

remaining (probably pectinate). Labrum reddish piceous, inter-ocellar space
piceous, palpi fuscous. Legs dark fuscous. Membrane of wings brownish hyaline,
fore wing with dark fuscous markings forming vague clouds, more particularly at
base and apex of costal area. Venation reddish brown, costal cross-veins in fore
wing sinuously bent in basal half of wing.

cJ GENITALIA. Ninth tergite short and deep, sternite forming a wide, rounded
subgenital plate, its apical margin produced in a triangular, membranous lobe.
Anal plates from side deep at base, upper angle produced in a stout finger, incurved,
lower apical angle of the finger further produced inward in a triangular lobe. Group
of trichobothria set low down on anal plate. Beneath the anal plates is a pigmented,
sclerotized plate, probably the tenth sternite, its lower angles projecting in rounded,
hairy processes. Aedeagus forming a pair of thin, rounded plates with slender
bases. These plates are slightly asymmetric, fused for a short distance medially
and slope downwards in roof-fashion from the centre line.

Length of fore wing, 27 mm.

FIG. ii. Ctenochauliodes forcipatus sp. n. Male genitalia. a, lateral ; b, left anal
plate, dorsal ; c, tenth sternite, from behind ; d, aedeagus, dorsal.

Holotype and paratype each with apex of abdomen mounted in Canada balsam.
The paratype has the wings damaged, as also is the apex of the abdomen, but as
far as can be ascertained it appears to be conspecific with the type. This species
differs from C. nigrovenosus (Weele) in the much more uniformly coloured wings.

PARACHAULIODES Weele

Weele, 1909, Notes Leyden Mus., 30 : 250 ; id., 1910, Coll. Zool. Selys, fasc. 5 (i) : 58.
Type species (original designation), Chauliodes japonicus McLachlan, 1867.
= Metachauliodes Weele, 1910, Coll. Zool. Selys. fasc. 5 (i) : 46, 55, 61. (Lapsus calami.}

The figures and descriptions of the male genitalia given by Weele were based upon
dried material and are not entirely satisfactory. New figures and descriptions have
been made, from preparations cleared in KOH solution, of the two species. It should

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI 431

be noted that the actual illustrations nos. 47 and 48 in Weele's monograph on the
Sialidae have been transposed, the lateral view of continentalis appearing with the
ventral view of japonicus.

Parachauliodes japonicus (McLachlan)

Fig. 12.

Chauliodes japonicus McLachlan, 1867, /. Linn. Soc. Zool. 9 : 232.

Parachauliodes japonicus (McLachlan) Weele, 1909, Notes Leyden Mus. 30 : 259 ; id., 1910,
Coll. Zool. Selys. fasc. 5 (i) : 58, figs. 46, 48 ; pi. 4, fig. 29.

FIG. 12. Parachauliodes japonicus (McLachlan) and P. continentalis Weele (holotype).
Male genitalia. a, lateral ; b, aedeagus, lateral ; c, aedeagus, dorsal ; d, left anal
plate, ventral.

$ GENITALIA. Ninth tergite with its apical margin produced, its centre slightly
concave ; lower apical angle produced and rounded, covering the aedeagus in side
view. Ninth sternite less sclerotized than eighth, short, broadly rounded, with a
small, transparent, triangular lobe projecting beyond its centre. Anal plates stout,

432 A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

longer than deep, each with a shallow, horizontal excision of the apex, dividing it
into a rounded upper part and a shallow, right-angled lower lobe. This excision
is more noticeable in dried examples. The upper part is armed on its inner surface
with black spinules. Aedeagus forming a short, broad tongue, in dorsal view con-
stricted about midway, its apex rounded, scarcely up-turned, and with its lateral
margins elevated.

Parachauiiod.es continentalis Weele
Fig. 12.

Parachauliodes continentalis Weele, 1909, Notes Ley den Mi*s. 30 : 259 ; id., 1910, Coll. Zool.
Selys. fasc. 5 (i) : 60, fig. 47, pi. 4, fig. 30.

c GENITALIA (from holotype). Ninth tergite produced in a rounded hood, from
the side with a definite excision of the apical margin below the attachment of the
anal plates, the lower apical angle so formed being subtriangular, obscuring the
aedeagus. Ninth sternite small, semi-membranous. Anal plates large, from the
side obliquely truncate at apex and divided into an upper and a lower portion by
a horizontal excision. From beneath, the lower portion is relatively short, about
one-third as long as upper, which carries a round area of black spinules on its lower
inner surface. The group of trichobothria is situated rather low on the outer
surface. Aedeagus short, with divergent, basally directed arms, a pair of triangular
lobes on its anterior upper surface, beyond which the apex is produced in a thin,
up-curved tongue.

KOREA.

NEOCHAULIODES Weele
Weele, 1909, Notes Leyden Mus. 30 : 259 ; id., 1910, Coll. Zool. Selys, 5 (i) : 60.

Type species (by designation of Weele, 1910), Chauliodes sinensis Walker, 1853.

This genus differs from Parachauliodes chiefly in the pectinate antennae of the <
(serrate in Parachauliodes) and in the simple anal plates, less produced lateral
angles of the ninth tergite and the longer aedeagus of the male. The wing venation
is variable, some species having the branches of Rs running almost straight to the
wing margin, whereas in others (such as the sinensis group) some of these branches
are sharply curved posteriorly before reaching the margin. I know of no reliable
characters by which to separate the females of the two genera and had it not been
that the antennal difference in the males is to some extent confirmed by differences
in the genital structure, I should have considered the two genera as synonymous.

Sufficient material has not been available to determine to what extent the external
genitalia of the female can be relied upon as specific characters. Differences occur
in form of the eighth sternite, anal plates and lateral gonapophyses, but in the
absence of long series of authentic material, it has been decided to figure the female
genitalia of those species only, of which the female is the holotype.

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI 433

Neochauliodes simplex (Walker)
Fig. 13-

Chauliodes simplex Walker, 1853, List Neur. Ins. B.M. : 200.

Neochauliodes simplex (Walker), Weele, 1910, Coll. Zool. Selys, 5 (i) : 61, fig. 49 ; pi. 4, fig. 31.

Weele's figure and description of the male genitalia were made from a dried example
of Walker's type series. This example, bearing Weele's determination label, has
now had the abdomen cleared in KOH solution and new figures and description
prepared.

fletcheri

FIG. 13. Neochauliodes simplex (Walker) paratype, and N. fletcheri sp. n. holotype.
Male genitalia. a, lateral ; b, aedeagus, dorsal ; c, left anal plate, dorsal.

<$ GENITALIA. Apical margin of the ninth tergite convex, sternite broadly rounded,
its centre produced in a thin, triangular tongue. Anal plates short and stout, from
the side subquadrate, rather longer than deep, with rounded angles, from above
ovate, with rows of comb-like teeth on the inner surface. Aedeagus long, spatulate,
slightly up-curved, its apex a little dilated in side view. From above it tapers
gradually to a broadly rounded apex.

Length of fore wing, $ 25-28 mm., $ 27-34 mm.

INDIA : Assam, Sylhet, Shillong, Khasi Hills ; United Provinces, Gahrwal.

434 A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

Neochauliodes fletcheri sp. n.
Fig. 13-

INDIA: Assam, Shillong, 5,000 ft., 26 . vi-io . vii . 1928 (T. Bainbrigge Fletcher),
3 <J, i $ ($ holotype, 8. vii, $ allotype, 30. vi). All in Brit. Mus. (N.H.).

In coloration and markings similar to N. simplex (Walker) but rather larger.
The chief structural difference is in the male genitalia. The anal plates in side view
are deeper than long. The aedeagus is dark reddish brown, from above with the
lateral margins slightly sinuous (straight in simplex), somewhat convex about mid-
way. From the side the dorsal margins are also more sinuous, tapering to a narrow
apex.

Length of fore wing : <$ 30-32 mm., $ 38 mm.

Holotype with abdomen mounted in Canada balsam.

Neochauliodes indicus (Weele)
Fig. 14.

Chauliodes indicus Weele, 1907, Notes Leyden Mus. 28 : 255, figs. 21-22, pi. 4, fig. 3.
Neochauliodes indicus Weele, 1910, Coll. Zool. Selys. 5 (i) : 62.

The specimen here dealt with agrees quite well with Weele's general description
and figure of the wings, but there are differences in male genitalia. These may be
specific, but after consideration of other instances it is considered that the differences
are not more than might be expected in comparing cleared preparations with dried
material.

$ GENITALIA. Apical margin of ninth tergite straight or slightly concave, some-
what depressed at its centre. Ninth sternite moderately short, broad, the centre
of its apical margin produced in a transparent, triangular lobe. Anal plates stout,
subquadrate from the side, apical margin widely but shallowly excised, so that the
apical angles are unusually pronounced. From above the anal plate is thick, its
apex truncate, inner surface set with rows of black spinules. Aedeagus broad at
base, tapering to a rather narrow, spatulate apex. From the side it is up-curved,
stout basally, apex thin and ligulate.

The chief differences between this example and Weele's figure lie in the excised,
less obliquely truncate apex of the anal plate and the rather broader apex of the
aedeagus. The specimen from which the above description was taken is labelled
Sikkim, Gopaldhara, Rongbong Valley, H. Stevens, and I have also seen a female
from Darjeeling and another from the Khasi Hills.

Neochauliodes tonkinensis (Weele)
Fig. 14.

Chauliodes tonkinensis Weele, 1907, Notes Leyden Mus. 28 : 260, pi. 5, fig. 3.
Neochauliodes tonkinensis Weele, 1910, Coll. Zool. Selys, 5 (i) : 67, fig. 51.

In the McLachlan Collection are one male and two females from the Ruby Mines
district of Burma, which I believe to be Weele's tonkinensis (described from a female).

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

435

The two females at present under discussion differ somewhat in size (fore wing,
39-50 mm.), and the smaller one and also the male have the wing pattern somewhat
resembling indicus but with the basal half of the fore wing heavily spotted. The
costal area is also heavily spotted and for part of its length has two rows of spots.
In the larger female the contrast between the spots and the wing membrane in the
distal half of the wings is much greater.

Indicus

FIG. 14. Neochauliodes indicus (Weele) and N. tonkinensis (Weele). Male genitalia.
a, lateral ; b, aedeagus, dorsal ; c, left anal plate, dorsal.

$ GENITALIA. Ninth tergite with apical margin truncate, sternite semi-mem-
branous, forming a short, broad subgenital plate, very slightly excised at its centre,
its sides a little concave. Anal plates quadrate from the side, inner apical angle
bearing rows of short, comb-like teeth. Aedeagus about twice as long as broad,
tapering to a rounded apex in distal half, lateral margins in this area slightly ele-
vated.

Length of fore wing : $ 39 mm., $ 39-50 mm.

436 A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

Neochauliodes umbratus sp. n.
Fig. 15.

INDO-CHINA : A. Vuillet, i <$, holotype, in Brit. Mus. (N.H.).

Head dark fuscous, labrum and occiput reddish brown, antennae piceous. Pro-
notum orange-brown, meso- and meta-nota, legs and abdomen fuscous. Fore wing
extensively clouded with fuscous, leaving two indefinite, hyaline bands running
obliquely basad from the costal margin, one midway and the other in the apical third.
Pterostigma dark brown, long, interrupted by a cream band. Venation brownish,
costal cross-veins bordered with fuscous. Hind wing mainly hyaline, with a fuscous
cloud at apex and another just beyond the middle, not reaching to the posterior
margin.

FIG. 15. Neochauliodes umbratus sp. n. (holotype). Male genitalia. a, lateral ; b,
aedeagus, dorsal ; c, left anal plate, dorsal.

$ GENITALIA. Ninth tergite only slightly produced apically, from the side with
lower apical angle very obtuse. Ninth sternite short and broad, its apical margin
produced in a lightly sclerotized, thin, triangular lobe or tongue, clothed in micro-
scopic setae. Anal plates globose, short, inner surface with the usual comb-like
teeth. From the side the anal plate is ovate, upper margin flattened. Aedeagus
elongate, narrow, its lateral margins sinuate and subparallel, and with a rounded,
apico-dorsally directed lobe on each side, situated at about one third before the
apex, which is shallowly excised.

Length of fore wing, 33 mm.

Type with abdomen mounted in canada balsam. N. umbratus may be easily
distinguished from the sinensis group by the presence of lateral lobes of the aedeagus.
These lobes are more apically directed than in N. bowringi, to which it is closely
related in genital structure.

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI 437

Neochauliodes bowringi (McLachlan)
Fig. 16.

Hermes sinensis Walker, 1853, List. Neur. Ins. B.M. : 203 (nom. preocc.).
Chauliodes sinensis Brauer, 1865 (nee Walker, 1853), Reise Novara, 1 : 102.
Chauliodes bowringi McLachlan, 1867, /. Linn. Soc. Zool. 9 : 260 ; Weele, 1907, Notes Leyden

Mus. 28 : 259, figs. 23-24 ; pi. 5, fig. 2.
Neochauliodes bowringi (McLachlan) Weele, 1910, Coll. Zool. Selys, 5 (i) : 66.

$ GENITALIA (holotype). Eighth sternite sclerotized, apical margin produced
in a parabolic subgenital plate. Ninth tergite sclerotized, short and deep. Lateral
gonapophyses as large as anal plates, flattened laterally, quadrate, directed upwards.
Anal plates stout, parallel-sided, then abruptly tapered to an acute apex.

CHINA : Hong Kong.

FIG. 1 6. Neochauliodes bowringi (McLachlan) and N. koreanus Weele. Female genitalia,

lateral, of hole-types.

Neochauliodes koreanus Weele
Fig. 16.

Neochauliodes koreanus Weele, 1909, Notes Leyden Mus. 30 : 261 ; id., 1910, Coll. Zool. Selys.
5 (i) : 65, pi. 4, fig. 39.

This opportunity is taken to call attention to two errors in the published locality
data of this species. In his original description, Weele writes " Two females from
Korea (the types) are in the British Museum." In his Selysian monograph he
alters this (without comment) to one female from Korea, Seoul, and one from Hong
Kong. The Korean example, which is here designated holotype, bears an additional
label (quoted by Weele) " Hong Kong Peak, Happy Valley." I feel sure that this
is an error and that this label should in fact belong to the Hong Kong example,
which incidentally is the one figured by Weele on plate 4.

$ GENITALIA. Eighth sternite parabolically produced apically. Lateral gona-
pophyses of ninth sternite foliate, from side with upper margin obtusely angled,

438

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

lower slightly convex, apex acutely rounded. Anal plates parallel-sided, lower
margin excised before the apex.

In the British Museum there is a second example labelled Hong Kong, from the
McLachlan Collection, in which the wings are even more heavily suffused with
fuscous.

Neochauliodes sinensis occidentalis Weele

Figs. 17, 18.

Chauliodes sinensis (Walker) Weele, 1907, Notes Leyden Mus. 28 : 262 (partim).
Neochauliodes sinensis occidentalis Weele, 1909, Notes Leyden Mus. 30 : 260 ; id. 1910, Coll.
Zool. Selys. 5 (i) : 64, pi. 4, fig. 40.

FIG. 17. Neochauliodes sinensis sinensis (Walker) and N.s. occidentalis Weele, holotypes.
Male genitalia. a, lateral ; b, aedeagus, dorsal ; c, left anal plate, dorsal.

<$ GENITALIA (holotype). Closely resembling those of N. s. sinensis (Walker)
but differing as follows : Anal plates from the side with apical angles more rounded,
apical margin obliquely convex. From above they are rather more pointed. Aedea-
gus from the side more slender, from above somewhat constricted basally, apex

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI 439

rounded, not excised. Ninth sternite shorter, with a triangular membranous tongue
at its apex.

$ GENITALIA (allotype). Eighth sternite produced in a subgenital plate, moder-
ately pointed apically in side view. Lateral gonapophyses rather small, shorter than
anal plates, flattened and pointed apically. Anal plates rhomboidal in side view,
stout in dorsal view, the group of trichobothria almost ventral, not visible from
above.

Very closely allied to sinensis Walker and probably correctly considered by Weele
as a subspecies of it. No holotype appears to have been selected from the type
series, and I therefore designate the male in the British Museum (N.H.), with abdo-
men now mounted in Canada balsam, and the female in the same collection as holo-
type and allotype respectively. Two other males in the same collection (and the

FIG. 18. Neochauliodes sinensis sinensis (Walker) and N.s. occidentalis Weele (allotype).

Female genitalia, lateral.

series in the Berlin Museum, not seen by me) become paratypes. The above speci-
mens are all from Omei Shan, Szechwan. In the British Museum (N.H.) are two
further males from N. W. CHINA (ex McLachlan collection) which agree in wing
pattern with the types. Also in the British Museum are a male and female from
W. CHINA, Chin-Fu-San, and a male from TONKIN, Ngai-Tio, which differ consider-
ably in wing markings and which may represent two further subspecies of sinensis.
The male genitalia, however, show only slight comparative differences from N.s.
occidentalis and it seems wiser therefore to leave them undescribed until more
abundant material is available.

Neochauliodes sinensis sinensis (Walker)
Figs. 17, 18.

Chauliodes sinensis Walker, 1853, List. Neur. Ins. B.M. : 199 ; Weele, 1907, Notes Leyden Mus.

28 : 261, figs. 25-26.
Neochauliodes sinensis (Walker) Weele, 1910, Coll. Zool. Selys. 5 (i) : 63.

$ GENITALIA (holotype). Ninth tergite with apical margin slightly produced,

440 A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

rather more sclerotized than sternite. The latter is semi-membranous, like the other
segments, forming a short, wide subgenital plate, apical margin slightly excised and
projecting beyond it is a triangular, membranous lobe. Anal plates short, quadrate
from the side, more or less globular from above. The inner surface is set with black
spinules, arranged in short, curved, comb-like rows. Aedeagus large, strongly
sclerotized, forming an elongate, tapering scoop or tongue, up-curved from the side,
its apex shallowly notched.

$ GENITALIA (MONGOLIA, McLachlan Collection). Eighth sternite with its apical
margin produced in a subgenital plate. Ninth tergite short and deep, lightly
sclerotized, its apical part membranous. Lateral gonapophyses foliate, semi-
membranous, lightly sclerotized and pigmented basally. Anal plates a little larger
than gonapophyses, narrow from above, tapering to an acute apex from the side.

The genitalia of the type differ from those figured by Weele (1907) in the form of
the aedeagus, which is less dilated apically from the side, and less sharply angled
upwards. In dorsal view the lateral margins are evenly convex in apical half, not
sinuous. It is possible that these differences may be due to the fact that the present
figures have been made after clearing the genitalia in KOH solution.

Neochauliodes sinensis truncatus ssp. n.
Fig. 19.

INDIA: Assam, Khasi Hills, Shillong, 5,000 ft., 13-15. vi. 1928 (T. Bainbrigge
Fletcher), holotype <, allotype $ and i $ paratype, all in Brit. Mus. (N.H.).

General appearance much as in the typical sinensis but in the fore wing the oblique,
transverse band is largely broken into rounded dots. The head, particularly in the
female, is distinctly darker and browner than the orange pronotum.

^ GENITALIA. Ninth tergite with its apical margin scarcely produced, rather more
sclerotized than the sternite, which forms a short, wide, semi-membranous sub-
genital plate, apical margin parabolic, and with a triangular, membranous lobe
projecting beyond its apex. Anal plates short, stout, apical margin rounded in
lateral and dorsal views, its inner apical surface set with rows of black, comb-like
spines. Aedeagus strongly sclerotized, large, forming a broad, up-curved plate.
From above it tapers but slightly to a wide, truncate apex with rounded angles.

$ GENITALIA. Eighth sternite produced apically in a broad, shallow, truncated
triangle, angles rounded, sides slightly concave. Ninth tergite moderately sclero-
tized. Lateral gonapophyses flattened, broadly pyriform in side view. Anal plates
about as long as gonapophyses, in side view with lower margin straight, upper
convex, apex rounded. Trichobothria placed laterally.

Length of fore wing : $ 33 mm. ; 9 37-44 mm.

The male genitalia are of the sinensis pattern but differ from the typical sub-
species in the more rounded anal plates and the wider, truncate apex of the aedeagus.

Protochauliodes reedi sp. n.

Figs. 20, 21.
S. AMERICA : Chili (Reed), 5 $, i $ (Calvert), i $, 2 $. Holotype $ (Reed, with

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

441

abdomen mounted in Canada balsam and left wings between celluloid), allotype $
(Calvert) and paratypes in Brit. Mus. (N.H.).

Body fuscous : labrum and clypeus fulvous, mandibles fulvous basally, reddish
brown apically. A piceous area around the ocelli, a fulvous spot between the

FIG. 19. Neochauliodes sinensis truncatus ssp. n. Genitalia. a-c, male, a, lateral ; b,
aedeagus, dorsal ; c, right anal plate, dorsal ; d, female, lateral.

FIG. 20. -Protochauliodes reedi sp. n. Male, anterior wing.

antennae, and back of head marked with fulvous. Antennae dark castaneous
basally, shading to fuscous apically. Thorax with obscure fulvous markings.
Wings greyish hyaline, venation brownish. Fore wing with the veins heavily

442

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

bordered with brownish dots or short, divergent fasciae, especially along the radius
and its sector and at the pterostigma. The centres of the cells are lightly clouded
with a darker grey, but not usually reaching the marginal dots. The attachments
of the wings fuscous, not orange. Venation much as in P. cinerascens, but in the
fore wing most of the costal cross-veins are distinctly angled distad.

cJ GENITALIA. Ninth sternite triangularly produced at centre of apical margin,
beyond rounded shoulders. Anal plates stout, slightly dilated apically and truncate

FIG. 21. Protochauliodes reedi sp. n., Or-e and P. cinerascens (Blanchard),/-A. Genitalia.
a, male, lateral ; b, male left anal plate, dorsal ; c, aedeagus, dorsal ; d, male ninth
sternite, ventral ; e, female, lateral ; /, aedaegus, dorsal ; g, male anal plate, lateral ;
h, male anal plate, lateral, from another example.

or slightly emarginate, with rounded angles in side view, curving inwards in dorsal
aspect. Aedeagus with a broad, arched base, subquadrate in dorsal aspect, its
apex with a triangular excision between two rounded lobes. There are two raised,
divergent ridges or lobes on the upper surface about midway, curving backward
and leaving a wide excision between them. From the side the apical lobes curve
slightly upwards and the median ridges appear as triangular elevations.

$ GENITALIA. Eighth sternite pigmented and sclerotized, a triangular area at

A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

443

centre of apical margin semi-membranous. Ninth tergite deep, lateral gonapophyses
stout, laterally compressed, extending beyond the tergite for about one-third of its
length, basal two-thirds pigmented, apical third membranous and carrying a very
small cercus. Anal plates finger-shaped, about as long as lateral gonapophyses,
apices rounded.

Length of fore wing : <$ 35-39 mm. ; $ 41-47 mm.

This species is closely allied to P. cinerascens (Blanchard) but differs from it in
the pattern of the wings, angled costal cross-veins of fore wings, fuscous wing bases,
the form of the aedeagus in the male and the rounded anal plates of the female.
My interpretation of P. cinerascens is based upon the male in the British Museum
(Nat. Hist.), described and figured by Weele in Coll. Zool. Selys, 5 (i) : 49-50, figs.
35-36, pi. 3, fig. 23. The form of the anal plates appears to be variable in cinerascens,

FIG. 22. Protochauliodes biconicus sp. n. Male genitalia. a, lateral ; b, left anal plate,
dorsal ; c, aedeagus, dorsal ; d, ninth sternite, ventral.

one example having the lower angle produced and with a small point. It is possibly
a little more clavate apically but does not offer any good characters by which the
dried insects may be separated. In cinerascens the aedeagus has its apex divided
into two truncate lobes, separated by an acute excision, and the mid-dorsal lobes
or ridges are much smaller.

Protochauliodes biconicus sp. n.

Fig. 22.

AUSTRALIA : New South Wales, Nowra, 8.x. 1928 (F. A. Rodway), i $, holotype,
in Brit. Mus. (N.H.).

444 A NEW GENUS AND NEW SPECIES OF THE CHAULIODINI

The unique male is somewhat damaged, one antenna and part of the other missing,
and the base of the right-hand wings defective. Body-colour dull reddish brown,
head with paler markings on labrum, clypeus, between antennae and on vertex.
Antennae subserrate, the anterior margin of each segment somewhat dilated, mainly
fuscous. Pronotum anteriorly orange-fulvous. Legs fuscous. Wings with mem-
brane pale brownish hyaline, fore wing with scattered brownish spots, darkest
along costal border ; hind wing with a few spots apically and along costal border.
Venation brown, arrangement of veins much as in P. cinerascens (Blanchard) .

$ GENITALIA. Ninth tergite with apical margin widely and shallowly excised.
Ninth sternite rather broad, lateral margins convex, apex truncate or slightly sinuous.
Anal plates stout, from the side with upper margin convex, lower sinuous, lower
apical angle produced, short and conical. From above this cone is directed back-
ward and slightly outward. Inner margin sinuous, armed with dense, fine spinules
and longer hairs. Aedeagus forming a subtriangular plate, lateral margins excised
basally, apex very slightly bilobed. From the side the aedeagus is abruptly angled
downwards near its base, then curved tailward to a moderately dilated apex with a
small, acute projection on its upper surface.

Length of fore wing, 32 mm.

Type with abdomen mounted in balsam. This is, I believe, the first Protochauli-
odes to be recorded from Australia. Until the type locality of Chauliodes dubitatus
Walker is known, one must not overlook the possibility that the present species
may be the male of dubitatus. It differs however in its narrower wings, different
pattern and the fact that in dubitatus the basal r-m cross-vein in the hind wing is
completely absent. From P. cinerascens it may be separated by the differently
shaped anal plates and by the form of the aedeagus.

APPENDIX

Since the galley-proofs of this paper were corrected, I have seen a copy of Mr.
E. F. Riek's paper ("Australian Megaloptera, or Alder-flies," 1954, Austral. Journ.
Zool. 2 (i) : 131-142, i pi., 3 text-figs.). It seems probable that some of the new
species described in the present paper will prove to be the same as species of Archi-
chauliodes described by Riek, but in the absence of genitalia figures of his species
it must be left for Australian entomologists to decide the matter. His Austro-
chauliodes dubitatus (Walker) may well be the same as my Protochauliodes biconicus,
but his photograph of the female wings shows a definite basal r-m cross-vein in the
hind wing, which is lacking in the type of dubitatus Walker.

D. E. KIMMINS,

6th Sept., 1954.

--T3

2 t ;!OV 1954

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

..,,954 THE GENERA
HENRICOHAHNIA BREDDIN,
DICROTELUS ERICHSON,

NYLLIUS STAL,

ORGETORIXA CHINA AND

ALLIED NEW GENERA

N. C. E. MILLER

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 3 No. 12

LONDON: 1954

THE GENERA HENRICOHAHNIA BREDDIN,

DICROTELUS ERICHSON, NYLLIUS STAL,

ORGETORIXA CHINA AND ALLIED

NEW GENERA

BY

N. C. E. MILLER

Pp. 445-488 ; 21 Text-figures.

'BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 3 No. 12

LONDON: 1954

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

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

This paper is Vol. 3, No. 12 of the Entomological
series.

PRINTED BY ORDER OF THE TRUSTEES OF
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Issued November, 1954 Price Twelve Shillings and Sixpence

THE GENERA HENRICOHAHNIA BREDDIN,

DICROTELUS ERICHSON, NYLLIUS STAL,

ORGETORIXA CHINA AND ALLIED

NEW GENERA

(HEMIPTERA-HETEROPTERA, REDUVIIDAE,
HARPACTORINAE, DICROTELINI)

By N. C. E. MILLER, F.R.E.S., F.Z.S., F.E.S.S.A.

(Commonwealth Institute of Entomology, London)

SYNOPSIS

This paper contains a revision of the genus Henricohahnia Breddin, descriptions of new allied
genera, also descriptions and figures of the genus Nyllius Stal, new genera allied to it and new
species of Orgetorixa China.

THE genus Henricohahnia was erected by Breddin in 1900 (Deutsche ent. Zeit. :
184-185) for the reception of a somewhat bizarre reduviid from the Tengger Moun-
tains of East Java. He gave it the specific name of wahnschaffei.

In 1903 Distant erected the genus Forestus (Ann. Mag. Nat. Hist. 12 : 251-253)
and placed in it four species typicus, montanus, spinosus and inermis but, in 1904
(Fauna Brit. India. Rhynchota 2 : 389) he transferred them to Henricohahnia.
In the same year and publication (loc. cit.) he described a new species Henrico-
hahnia gallus and in 1909 another new species H. badgleyi (Ann. Soc. ent. Belg.

53 : 373).

A critical examination of the species described by Distant, however, has revealed
that the species inermis and badgleyi should not have been placed in Henricohahnia.
Accordingly I have erected a new genus Karenocoris for them.

In the present paper I have also described two more new allied genera Tapiro-
coris and Malaiseana both of which are allied to Henricohahnia.

I have revised all previous descriptions of Henricohahnia species with the excep-
tion of that of wahnschaffei which is given in the original. A few remarks supple-
mentary to Breddin's generic description, however, are added.

All species are figured for the first time except Henricohahnia typica (Distant).

All the known genera in the Dicrotelini are keyed on pages 466 and 474.

Nothing appears to be known about the ecology of Henricohahnia or of any of
the other new genera described herein, but it is probable, to judge by their habitus,
that they pass some portion of their lives under the loose bark of dead trees.

The types and paratypes of all the new species are in the British Museum (Natural

ENTOM. Ill, 12. 30

448 HENRICOHAHNIA BREDDIN AND ALLIED GENERA

History) with the exception of the type and paratype of Neonyllius echinus gen. et
sp. n., which are in the Vienna Museum.

For the genera Nyllius and Dicrotelus, Stal proposed the tribal name of Dicro-
telini (1859, Ofv. Vet. Ak. Fork. 16 (8) : 366). To this tribe I consider that the
genera Neonyllius gen. nov., Paranyllius gen. nov., Orgetorixa China, and Henri-
cohahnia Breddin also belong.

I express my thanks to Dr. Stevan Keler of the Zoological Museum Berlin, for
his kindness in allowing me to examine the types of Dicrotelus prolixus Erichson
and Nyllius asperatus Stal. I am also indebted to Dr. W. E. China of the British
Museum (Nat. Hist.) for his assistance in elucidating the synonymy of Sphagiastes
Stal.

Henricohahnia Breddin

Henricohahnia Breddin, 1900, Deutsche ent. Zeit. : 184-185.

Forestus Distant, 1903, Ann. Mag. Nat. Hist. (12) ; 251-253, type F. typicus Distant ; Distant,
1904, Fauna Brit. India, Rhynchota, 2, 389 ; Distant, 1909, Ann. Soc. ent. Belg. 53 : 373.

Capite a super o viso longe cylindrico, pronoto fere aequilongo, antrorsum in processum com-
pressum angustum, a latere viso acutangularem longe prominente ; capite postoculari quam parte
anteoculari cum oculis ipsis nonnihil longiore, retrorsum levissime angustato ; capitis impressione
superiore pone oculos minusculos obsoleta, fere nulla, ocellis punctiformibus ab oculis sat remotis.
Capite supra granulis paucis setigeris, haud tamen spinulis instructo, infra utrimque spinulis
paucis seriatis armato. Rostri articulo basali marginem posticum oculorum haud attingente,
quam articulo secundo plus duplo breviore. Antennis breviusculis, capite pronotoque simul sumptis
brevioribus, dimorphis ; articulo primo subclavato capite postoculari multo breviore et articulis
secundo tertioque singulis in $ breviore, in $ subaequilongo ; primo tuberculis compressis et obtu-
satis sublobiformibus horrente, secundo in $ supra paucis eiusmodo dentibus armato, in <J inermi,
piloso. Antepronoto tuberculis nonnullis acutiusculis, post-pronoto granulis sparsis armatis,
pone humeros retrorsum sat longe producto. Scutelli apice sat angusto. Hemelytrorum areola
discali valde elongata atque angusta. Membranae area interiore valde parva, quam area exteriore
bdsi ter angustiore eademque area duplo breviore ; area exteriore retrorsum distincte dilatata. Ab-
dominis marginibus in lobos mediocres dilatatis. Prostethio in sulci lateribus utrimque tuberculis
nonnullis, mesostethio antice latera versus nodulo distincto armatis. Venire medio subtiliter
carinato. Femoribus quam tibiis multo longioribus, anterioribus subincrassatis. Femoribus
omnibus supra et extus spinulis fortibus numerosis seriatis, anterioribus subtus serie unica sub-
obliqua spinularum paucarum armatis. Tibiis anterioribus supra spinulis crassis nonnullis
minoribus, anticis subtus paucis maioribus instructis.

Breddin omits to mention in his generic description that the coxae and trochanters
are spinose and that the basal areas of the membranal cells are more or less coriaceous.

At the conclusion of his description of the genus, Breddin adds : " genus singular e
inter Blaptonem et Sclominam ordinandum." However, I consider the relationship
of Henricohahnia to these two genera to be very remote, for the following reasons.
In Sclomina Stal (1861, Ent. Zeit. Stett. 22 : 137) all segments of the antennae are
slender and the basal segment is approximately twice as long as the head, the rostrum
is more or less curved (as in the majority of the Harpactorinae), the head is not
cylindrical, the transverse sulcus is deep and wide and the vertex is not produced
anteriorly. The spines on the head in Sclomina are long and slender. The pro-
notum has long slender spines and the posterior margin of the posterior lobe is

HENRICOHAHNIA BREDDIN AND ALLIED GENERA 449

almost straight. In the hemelytra the internal cell of the membrane is about
two-thirds as long and one-third wider than the external cell and the venation of
the corium is different in having a small, triangular cell adjacent to the discal cell.

Blapton was established by Spinola in 1850 (separata from Mem. Matem. fisica,
Soc. ital. Sci. 25 (i), 1852, 88 and 144).

Spinola states that " ha per tipo una specie di Cayenna discritta dai Signori
Amyot e Serville, vedi Hist, des Hemypt. 376, 113, Sinea punctipes Amyot e Serville."
The type-species of the genus Blapton Spinola, 1850, is thus, undoubtedly, Sinea
punctipes Amyot & Serville, 1843. But this S. American species is congeneric
with the Brazilian Milyas ornaticeps Stal, 1858, which is the type species of the
genus Milyas Stal, 1858, which, because it was preoccupied, was re-named Pselliopus
by Bergroth (1905, Rev. Ent., : 112). Blapton Spinola, 1850, thus has priority
over Pselliopus Bergroth, 1905, and the South African genus Blapton auctt. nee
Spinola takes the next available name, which is Sphagiastes Stal (1853, Ofu. Vet.
Ak. Fork. : 43), type 5. horrificus 1853 = ramentaceus Germar, 1837 = dregei
Spinola, 1850.

There is one other species which has been placed in the genus Blapton, namely
Arilus pilipes Stal (1855, Ofv. Vet. Ak. Fork. : 43). In 1912 Schouteden (Rev. Zool.
afr. 2 : 114) considered that B. pilipes should be placed, at least, in a sub-genus
of Blapton for which he proposed the name Lepton. In 1913 (loc. cit. : 437), having
discovered that Lepton was preoccupied, Schouteden substituted the name Lerton.
This genus is allied to, but is not a subgenus of Sphagiastes.

The affinity of Henricohahnia to Sphagiastes is remote. There is a slight similarity
in the shape of the head and rostrum, but the armature of the head, the extraor-
dinary spinose pronotum, particularly its posterior lobe, the unique structure of the
connexivum and the tuberculate ventral surface of the abdomen, all indicate that
these two genera are by no means closely related. Furthermore, in Sphagiastes
the antennal segments and legs are slender and have compact tufts of short thick
setae as well as normal setae. The abdomen in Sphagiastes is not carinate mid-
ventrally, the membranal cells are more or less equal in area and the membrane has
irregularly ramate veins apically.

Henricohahnia, I think, is allied to Nyllius Stal (1859, Ofv. Vet. Ak. Forh. 16
( 8 ) 355) an( i to Orgetorixa China (1925, Ann. Mag. Nat. Hist. (9) 15 : 486-488) with
which it has the following characters in common, namely the produced vertex,
more or less cylindrical head, tuberculate and spinose pronotum with the posterior
margin medially excavate.

Dicrotelus Erichson (1842, Arch. 8 (i) : 284) has certain affinities with these three
genera but differs in having a slender habitus, a smooth integument and non-
tuberculate connexivum and also in being brachypterous.

The venation of the hemelytra of Henricohahnia and Nyllius differs from that of
Orgetorixa in having a relatively small but distinct internal membranal cell which
Orgetorixa lacks. The presence, absence, or size of this cell, however, may not be
of great importance, since it appears to vary somewhat and may be absent in indi-
viduals of species in which it is normally present. For example, the cell was absent
from a paratype of Nyllius asperatus which I examined.

ENTOM. Ill, 12 30

450 HENRICOHAHNIA BREDDIN AND ALLIED GENERA

Key to Henricohahnia species

1. Humeral angles strongly spinouslj- or conically produced ..... 4.
-. Humeral angles not or hardly at all produced . . . . . . . 2.

2. Head and pronotum without black pattern ....... indica sp. n.

-. Head and pronotum with black pattern . . . . . . : . 3.

3. Produced portion of vertex acute apically in lateral view . . . vitticeps sp. n.
-. Produced portion of vertex broadly rounded apically in lateral view . gallus Distant

4. Humeral angles spinous ........... 5.

. Humeral angles conical, thick, strongly recurved ....... 6.

5. Posterior margin of posterior pronotal lobe narrowly and deeply incised medially

cauta sp. n.
-. Posterior margin of posterior pronotal lobe widely and shallowly incised medially

typica (Distant).

6. Produced portion of vertex truncate apically in lateral view ..... 7.
-. Produced portion of vertex rounded apically in lateral view . . . . . 8.

7. Spines at humeral angles truncate apically . . . . . . . vittata sp. n.

-.. Spines at humeral angles sub-acute apically ........ 9.

8. Posterior margin of posterior pronotal lobe undulate with moderately deep and wide

median incision . . . . . . . , . montana (Distant)

-. Posterior margin of posterior pronotal lobe undulate with very shallow median

incision .......... wahnschaffei Breddin.

9. Posterior margin of posterior pronotal lobe undulate with very wide, shallow median

incision ........... spinosa (Distant)

-. Posterior margin of posterior pronotal lobe regularly undulate . . tinctoria sp. n.

Henricohahnia wahnschaffei Breddin

(Fig. i)
Deutsche ent. Zeit. 1900 : 184-185.

Colour : Ferruginescenti vel fuscescenti-cinerea, griseo pilosa tomentoque cinereo dense induta.
Rostri apice, spinulis gulae, antennarum articulo primo apicem versus quartoque toto nee non
femoribus supra tibiarumque maxima parte, ut videtur semper, nigris vel nigricantibus. Rostri
articuli primi apice secundoque toto, tibiarum annulo distincto medio alioque subbasali obsoletiore,
tarsis articulorum antennarum secundi et tertii parte basali quartique ima basi ferruginescenti-
albidis.

Structure : Capite infra pone oculos spinulis utrimque binis juxtapositis et basin versus tuberculis
utrimque singulis armato. Post-pronoto basi late sinuato vel apertangulariter exciso ; antice
carinis duobus et magis extrorsum lineis utrimque singulis dense pilosis longitudinalibus, omnibus
retrorsum diverge ntibus, inter humeros carinula utrimque transversali ex angulo humerali exeunte
et intus mox evanescente instructo ; hoc angulo sublobiformiter prominulo, spina extrorsum et
valde retrorsum producta armato. Tuberculis spiniformibus femorum in series quatuor vel quinque
ordinatis, anterioribus praeterea subtus spinis tribus vel quatuor armatis. Tibiis anticis subtus
apicem versus spina forti ceterisque longiore calcariformi nonnihil extrorsum directa munitis.
Abdomen in mare hemelytris clausis non multo in femina multo latiore, rhomboideo. Apice
segmenti sexti connexivalis cum basi septimi in lobum apertangularem, apice segmenti quinti cum
ima basi sexti in lobum mediocrem (in mare irregulariter quadrangular em postice extus acuminatum,
in femina trapezoideum apice truncatum) dilatis, in femina angulo apicali segmenti etiam quarti
apertangulariter extrorsum prominente. Abdominis margine toto tuberculis minutis setiferis,
venire connexivoque granulis sparsis instructis.

The measurements given by Breddin, are total length, 13-15-5 mm. ; greatest
pronotal width, 4-4-75 mm.

FIG. i. Henricohahnia wahnschaffei Breddin.

452 HENRICOHAHNIA BREDDIN AND ALLIED GENERA

The specimens on which Breddin based his descriptions were collected by Jung-
huhn in the Tengger Mountains of East Java. In the British Museum there are also
specimens collected by F. C. Drescher in other localities in Java, namely Gunong
Slamet at an altitude of 2,200 ft., Gunong Tankoeban Prahoe between 4,000 and
5,000 ft., Gunong Boerangrang at about 4,000 ft. and Gunong Malabar at a similar
altitude.

Henricohahnia gallus Distant
(Fig. 2)

Henricohahnia gallus Distant, 1904, Fauna Brit. Ind. Rhynchota 2 : 389.

Colour : Dark testaceous. Vertex with three longitudinal parallel stripes, postocular dorsally
with two longitudinal parallel stripes not extending to base, laterally with an irregular stripe
connecting posteriorly with dorsal stripes, black. Rostrum pale testaceous ; segments i and 2
basally, segment 3 apically, suffused with dark brown. Anterior lobe of pronotum with black
pattern as in Fig. 2. Corium suffused with piceous apically ; membrane hyaline, pale testa-
ceous ; venation testaceous ; base of veins of cells with a raised pale luteous spot. Apex of
scutellum pale luteous ; depressed area of disc piceous. Spines on legs pale yellow.

Structure : Basal segment of antennae strongly tuberculate, a little more than twice as long
as produced portion of vertex ; tylus with tubercles ; produced portion of vertex apically
from above sub-acute ; laterally sub-truncate. Head sparsely tuberculate. Anterior lobe of
pronotum with sparse erect and sub-erect tubercles more or less regularly arranged ; posterior
lobe medially depressed with a somewhat obscure, short carina bearing low tubercles, sub-
dorsally ; lobe very sparsely tuberculate ; humeral angles sub-conical and with low setigerous
tubercles ; posterior margin undulate. Disc of scutellum somewhat deeply depressed ; apex
produced, horizontal, tuberculate. Hemelytra extending very little beyond apex of abdomen.

Total length, 9-00 mm.

Hemelytra, 5-50 mm.

Greatest pronotfl width, 2-30 mm.

ic? (holotype), i^, India, Kotagiri, in Brit. Mus. (Nat. Hist.).

Henricohahnia vitticeps sp. n.

(Fig. 3)

Colour : Testaceous. Vertex with a spot laterally, postocular with a wide, median stripe,
very narrowly divided medially, and narrow lateral stripes ; anterior lobe of pronotum with
linear suffusion, disc of scutellum, black.

Structure : Basal segment of antennae less than twice as long as produced portion of vertex,
with short, conical and long cylindrical tubercles bearing setae ; segment 2 with few setigerous
tubercles. Produced portion of vertex acute apically. Head and anterior lobe ot pronotum
sparsely tuberculate. Posterior lobe of pronotum somewhat deeply medially depressed with a
carina on each side of depression ; humeral angles not produced, rectangular, rounded ; lobe
very sparsely tuberculate ; posterior margin undulate with median incision very broad and
shallow. Disc of scutellum feebly transversely depressed posteriorly.

Total length, u-oo mm.

Hemelytra, 6-50 mm.

Greatest pronotal width, 3.00 mm.

I? (holotype), Assam, Utakamund, in Brit. Mus. (Nat. Hist.).

FIG. 2. Henricohahnia gallus Distant.

454

HENRICOHAHNIA BREDDIN AND ALLIED GENERA

FIG. 3. Henricohahnia vitticeps sp. n.

FIG. 4. Henricohahnia indica sp. n.

FIG. 5. Henricohahnia spinosa (Distant).

HENRICOHAHNIA BREDDIN AND ALLIED GENERA 457

Henricohahnia indica sp. n.

(Fig. 4)

Colour : Segments i and 2 of antennae piceous ; remaining segments brown. Head, anterior
lobe of pronotum, scutellum piceous ; anterior lobe of pronotum with two suffused yellowish
spots medially basally ; posterior lobe of pronotum, propleura, acetabula and abdomen ven-
trally, testaceous ; meso- and metapleura and sterna piceous ; meso- and metasternum with a
median testaceous suffusion. Abdomen dorsally brown ; connexivum suffused with black ;
tubercles on ventral surface brown ; apex of yth segment blackish. Corium dark testaceous
with the cross- vein at base of internal cell, external half of basal vein of external cell and apex
of vein R pale luteous ; membrane pale infumate with a coppery lustre ; metathoracic wings
hyaline, iridescent. Legs blackish ; base of anterior tibiae testaceous ; spines on anterior
femora dark yellow ; median and posterior tibiae testaceous with dark brown annulations.
Tubercles on anterior lobe of pronotum yellowish.

Structure : Antennae with abundant, moderately long forwardly directed setae and sparse
longer setae ; basal segment feebly curved, thicker towards apex and thrice as long as produced
portion of vertex ; tuberculate on lower surface, the apical tubercles the longest. Head
sparsely tuberculate ; produced portion of vertex from the side rounded apicallv and with the
sides sub-parallel ; tylus tuberculate not greatly produced. Lateral angles of collar obtusely
conical ; anterior lobe of pronotum with a few moderately long setigerous tubercles sub-dorsally
and laterally ; humeral angles obtusely conical ; posterior lobe of pronotum with very sparse,
low tubercles ; posterior margin sinuate with median incision very wide and shallow. Disc of
scutellum shallowly depressed. Hemelytra extending just beyond apex of abdomen. External
apical angle of connexival segment 5 lobately produced, of segment 6 obtuse angulate. Anterior
femora and tibiae with moderately long and short setigerous spines ; median and posterior
femora with low, setigerous tubercles.

Total length : $, 9-50 mm. ; $, 10-60 mm.

Hemelytra : $, 6-50 mm. ; $, 6-50 mm.

Greatest pronotal width : $, 2-50 mm. ; ?, 3-00 mm.

ic (holotype) and 3^, 2$ (paratypes), S. India (no precise locality) (B.M. 1930-
599) ; i<2, i? (paratypes), S. India, Lovedale, Nilgiri Hills (B.M. 1915-60) ; i$
(paratype), S. India ; Kodai Kanal (B.M. 1926-171) ; all collected by T. V. Camp-
bell.

Henricohahnia spinosa (Distant)
(Fig. 5)

Forestus spinosus Distant, 1903, Ann. Mag. Nat. Hist. (7) 11 : 252.
Henricohahnia spinosa (Distant), 1904, Fauna Brit. Ind. Rhynchota 2 : 388.

Colour : Pale testaceous. Vertex with two pe rallel, narrow, longitudinal stripes, postocular
with a wide median longitudinal stripe, black. Humeral spines light brown. Veins of discal
cell of corium and apical margin, except extreme apex, pale luteous. Costal margin of corium
suffused with piceous. Membrane pale infumate ; venation testaceous. Segments 2, 5 and 6
apically and base of segment 7 of connexivum piceous ; marginal tubercles mostly black.
Spines on femora whitish, very narrowly brown apically. Metathoracic wings infumate ;
venation dark testaceous.

Structure. Basal segment of antennae sparsely tuberculate, a little more than twice as long
as produced portion of vertex. Tylus more or less vertical with an apical tubercle. Produced
portion of vertex rounded apically. Head and pronotum sparsely tuberculate. Posterior
lobe of pronotum with indications of .short carinae sub-dorsally ; humeral angles spinously
produced ; posterior margin undulate with the median incision moderately deep. Disc of

ENTOM. Ill, 12. 3<>

FIG. 6. Henricohahnia tinctoria sp. n.

HENRICOHAHNIA BREDDIN AND ALLIED GENERA 459

scutellum damaged ; apex moderately produced, horizontal. Hemelytra extending just
beyond apex of abdomen.

Total length, 12-50 mm.

Hemelytra, 8-50 mm.

Greatest pronotal width, 3-50 mm.

i$ (holotype), Assam ; Sikkim (Atkinson Coll., B.M. 1892-6) ; 2$, same locality
(Distant Coll., B.M. 1911-383) ; 2$, Bengale, Kurseong (Distant Coll., B.M.,
1911-383) ; i$, Dharmoto, Kumaon, 5,600 ft., 9.^.1912 (A. D. Imms) ; i$,
Chaurata. The ovum of H. spinosa is cylindrical, rather strongly curved at oper-
cular end. Total length, 2-30 mm.

Henricohahnia tinctoria sp. n.

(Fig. 6)

Colour : Testaceous. Head with two longitudinal, parallel black stripes from base of pro-
duced portion of vertex to transverse sulcus and a wide black stripe thence almost to base.
Ocelli margined with red. Anterior lobe of pronotum with piceous pattern as in Fig. 6. Spines
at humeral angles piceous. Tubercles on both pronotal lobes brownish or piceous. Dorsal
surface of abdomen, except connexivum, base of clavus broadly and of corium narrowly, red ;
corium apically with a U-shaped raised luteous spot ; remainder of corium faintly suffused
with red ; vein Sc of metathoracic wing red. Connexivum with segments 2-6 dorsally with
a large irregular piceous spot apically. Mesopleural episternum anteriorly suffused with
piceous. Tubercles on pleura and abdomen ventrally piceous.

Structure : Basal segment of antennae more or less straight, not very strongly tuberculate
and somewhat narrower towards apex ; segment 2 about one-third longer than i and without
tubercles. Head sparsely tuberculate. Basal segment of rostrum extending very little beyond
anterior margin of eyes. Humeral angles spinously produced, the spines rounded apically and
directed posteriorly somewhat ; posterior lobe of pronotum with sparse, very low, setigerous
tubercles ; posterior margin regularly undulate, the median incision moderately deep. Disc of
scutellum very deeply depressed ; carina with a few low tubercles. All segments of connexivum
with short, marginal tubercles ; segment 7 of abdomen dorsally transversely rugose in apical
half. Hemelytra extending just beyond apex of abdomen.

In the female paratype the basal antennal segment is more strongly tuberculate and segment
2 is relatively shorter and feebly tuberculate.

Total length : $, 12-00 mm. ; $, 13-50 mm.

Hemelytra : <$, 7-50 mm. ; ?, 13-50 mm.

Greatest pronotal width : <J, 3-50 mm. ; $, 4-00 mm.

1$ (holotype), Sikkim, Rongli Chu, 3,000 ft., 29.^1.1924 (B.M. 1924-386) ; i<,
i$ (paratypes), Sikkim, Singhik, 3,500 ft., 23.^.1924 (B.M. 1924-386) ; coll. Maj.
R. W. G. Kingston (B.M. Everest Expedition).

Henricohahnia vittata sp. n.

(Fig. 7)

Colour : Testaceous. Posterior lobe of pronotum with a diagonal, longitudinal stripe within
depressions, pro- and mesopleura with a transverse stripe, metapleural acetabula and apex
of tibiae piceous ; coxae with suffused piceous spot. Corium testaceous ; apical half of clavus,
membrane and metathoracic wings infumate.

Structure : Basal segment of antennae twice as long as produced portion of vertex, feebly

4 6o

HENRICOHAHNIA BREDDIN AND ALLIED GENERA

FIG. 7. Henricohahnia vittata sp. n.

HENRICOHAHN1A BREDDIN AND ALLIED GENERA

461

FIG. 8. Henricohahnia cauta sp. n.

462 HENRICOHAHNIA BREDDIN AND ALLIED GENERA

curved and strongly tuberculate. Head with abundant, erect, moderately long setae, sparsely
tuberculate. Posterior lobe of pronotum with very strong sub-dorsal carinae and less strong
lateral carinae ; humeral angles tuberculately produced ; median incision on posterior margin
of lobe regularly concave. Disc of scutellum deeply depressed ; carina and apex irregularly
tuberculate. Head, body, legs with abundant setae. Anterior coxae and trochanters strongly,
median coxae and trochanters, moderately tuberculate.

Total length, 18-00 mm.

Hemelytra, 12-00 mm.

Greatest pronotal width, 3-80 mm.

i$ (holotype), Indo-China ; Laos, Xieng Khouang, lo.v.igig (R. V. de Salvaza)
(B.M. 1917-98). An ovum dissected from this female is dark brown with the
differentiated portion of the chorion whitish. It is cylindrical, glabrous, somewhat
narrower at the opercular end and with one side straight. Total length, 2-40 mm.

Henricohahnia cauta sp. n.
(Fig. 8)

Colour : Testaceous. Humeral angles of pronotum and apex of corium suffused with brown.
Membrane and metathoracic wings faintly infumate, iridescent. Pygophore and metapleura
suffused with piceous. Lateral tubercles of prosternum piceous.

Structure : Basal segment of antennae with short, conical setigerous tubercles and two
longer cylindrical tubercles on upper surface, apically ; remaining segments with sparse,
moderately long, abundant, short setae. Produced portion of vertex from above very narrowly
triangular, acute apically ; from the side triangular, acute apically.

Total length : $, 12-00 mm. ; $, 12-10 mm.

Hemelytra : <$, 7-00 mm. ; $, 7-00 mm.

Greatest pronotal width : $, 3-50 mm. ; $, 3-30 mm.

ic? (holotype), Indo-China, Haut Mekong, Nam Mat, 15. iv. 1918; 3$ (para-
types), Laos, Xieng Khouang, 7-17^.1918 (R. V. de Salvaza) (B.M. 1918-1).

Henricohahnia typica (Distant)
(Fig. 9)

Forestus typicus Distant, 1903, Ann. Mag. Nat. Hist. (7) 11 : 251.
Henricohahnia typica (Distant), 1904, Fauna Brit. Ind. Rhynchota 2 : 387.

Colour : Testaceous, except dorsum of abdomen, pale reddish ; tubercles on connexivum
mostly piceous ; segments 5-7 dorsally suffused with black. Basal half of clavus and extreme
base of corium reddish ; base of veins of membranal cells and an irregular suffusion on corium
apically pale luteous ; membrane faintly infumate ; venation darker. Coxae with piceous
spots. Vein SC of metathoracic wing pale yellow, except apical fourth violaceous ; wing faintly
infumate, iridescent ; remaining veins dark brown. In the male, the pygophore has two large,
suffused piceous spots.

Structure Basal segment of antennae strongly tuberculate and with a long tubercle on upper
apical margin ; segment less than half as long as produced portion of vertex ; tylus with
moderately long, setigerous tubercles ; produced portion of vertex from above and from the
side acute apically. Head sparsely tuberculate. Anterior lobe of pronotum with conical,
acute, moderately long and short tubercles ; lateral angles of collar conically produced ;
posterior lobe with strong carinae anteriorly, sub-dorsally ; humeral angles conical, strongly

HENRICOHAHNIA BREDDIN AND ALLIED GENERA

463

FIG. 9. Henricohahnia typica (Distant).

464 HENRICOHAHNIA BREDDIN AND ALLIED GENERA

recurved, tuberculate ; posterior margin of lobe feebly undulate. Hemelytra extending very
little beyond apex of abdomen.

Total length : <$, 11-50 mm. ; ., 13-50 mm.

Hemelytra : <J, 7-00 mm. ; $, 8-00 mm.

Greatest pronotal width : Q", 3-00 mm. ; $, 3-80 mm.

i$ (holotype), Assam, Sikkim, 5^, 3$, same locality (Atkinson Coll., B.M. 1892-
6) ; ic, N. Khasia, 1,500-3,000 ft. (Distant Coll., B.M. 1911-383).

Henricohahnia montana (Distant)
(Fig. 10)

Forestus montanus Distant, 1903, Ann. Mag. Nat. Hist. (7), 11, 252.
Henricohahnia montana (Distant), 1903, Fauna Brit. India Rhynchota 2, 388).

The following is a description of the male :

Colour : Head and thorax piceous ; anteocular laterally with a light brown spot in front of
eyes ; postocular in front and between ocelli, extreme base of head, anterior lobe of pronotum,
propleural epimeron, metapleural acetabula anteriorly and rostrum, light brown. Posterior
margin of posterior lobe of pronotum suffused with brown. Meso- and metasternum and
abdomen ventrally, light brown with faint reddish suffusion. Femora piceous ; tibiae brown
with a pale yellowish annulation interrupted on outer surface. Corium brown with an irregular
luteous spot at base of veins of membrane and with piceous suffusion ; membrane and meta-
thoracic wings faintly infumate, iridescent. Setae and pubescence pale fulvous.

Structure : Basal segment of antennae a little less than twice as long as produced portion of
vertex, feebly tuberculate ; remaining segments without tubercles and with dense, short setae
and sparse longer setae. Head sparsely tuberculate except basally, laterally ; ventro-lateral
margin with moderately long tubercles. Basal segment of rostrum extending to middle of
eyes. Tubercles on anterior lobe of pronotum short, conical except tubercles sub-dorsally,
basally which are moderately long ; posterior lobe of pronotum sparsely tuberculate, except at
humeral angles ; lobe with an arcuate carina sub-dorsally and a short transverse carina from
humeral angles ; posterior margin sinuate ; spines at humeral angles thick, rounded apically
and directed posteriorly somewhat. Disc of scutellum deeply depressed ; carina with a few
tubercles. Segment 7 of abdomen dorsally transversely striate in apical half ; segments 5
and 6 of connexivum produced ; external margin of segments with low, setigerous tubercles ;
connexivum ventrally very narrow ; spiracles with ostiole directed outwards ; lateral margin
of abdomen ventrally carinate.

Total length : <$, 13-00 mm. ; $, 14-00 mm.

Hemelytra : <J, 7-50 mm. ; 9. 8-00 mm.

Greatest pronotal width : <J, 4-00 mm. ; $, 4-50 mm.

The holotype of H. montana, a female from Mungphu, Assam (Distant Coll.,
B.M. 1913-451), in the British Museum, differs in coloration from the male in having
the head and body dark testaceous and the apex of the corium luteous. The 3rd
antennal segment is pale yellow in the basal half. The basal antennal segment is
strongly tuberculate and segment 2 is relatively shorter and has a few tubercles.
The spiracles are normal and the connexivum wide ventrally. In the British
Museum there is also a male from E. Garo, Assam, collected at an altitude between
1,500 and 2,500 ft. (Distant Coll., B.M. 1911-383), and a male and a female from
N. Khasia from a similar altitude (B.M. 1914-383). There are also females from
Shillong, Assam, altitude 4,900 ft. (B.M. 1913-451).

HENRICOHAHNIA BREDDIN AND ALLIED GENERA

n

FIG. 10. Henrtcohahnia montana (Distant).

466 HENRICOHAHNIA BREDDIN AND ALLIED GENERA

Key to Genera allied to Henricohahnia.

1 . Basal antennal segment with moderately long cylindrical tubercles ; abdomen mid-

ventrally carinate ; coxae and trochanters spinose . . Henricohahnia Breddin.

-. Basal antennal segment with low, rounded tubercles ; abdomen not carinate ventrally ;

coxae sometimes with a very few spines ; trochanters without spines . . . 2.

2. Head and anterior lobe of pronotum with strong black pattern . Tapirocoris gen. nov.
-. Head and anterior lobe of pronotum without pattern . . . . . . 3.

3. Head and pronotum with abundant, low tubercles . . . . . . 4.

-. Head and pronotum with sparse low tubercles . . . . . . . 5.

4. Humeral angles narrowly rounded or conical ; anterior tibiae with few or no spines

Karenocoris gen. nov.

5. Humeral angles broadly rounded ; anterior tibiae with six or more spines

Malaiseana gen. nov.

Key to Species of Karenocoris gen. nov.

1. Head and pronotum strongly tuberculate ; humeral angles conical . . . . 2.
-. Head and pronotum moderately tuberculate ; humeral angles rounded . pustulatus sp. n.

2. Posterior margin ol posterior pronotal lobe undulate with deep median incision

badgleyi (Distant).
-. Posterior margin of posterior pronotal lobe undulate . . . inermis (Distant).

Karenocoris gen. nov. 1

Size small. Basal segment of antennae tuberculate, longer than anteocular. Postocular
longer than anteocular with sides parallel ; constricted basally. Head cylindrical, shorter than
pronotum, ventro-laterally spinose ; entirely tuberculate, except ventrally, basally and a
narrow area on vertex, smooth ; vertex produced anteriorly. Ocelli small, widely separated.
Basal segment of rostrum extending to middle of eyes, a little more than half as long as segment
2 ; segments 2 and 3 straight. Pronotum longer than wide ; anterior lobe shorter than posterior
lobe ; both lobes with a median sulcus and tubercles which are arranged in rows on anterior
lobe ; posterior margin of posterior lobe undulate. Scutellum longer than wide, produced
apically. Pleura, abdomen ventrally and lateral margins of connexival segments tuberculate.
Hemelytra with internal cell of membrane very small ; corium pubescent ; base of cells of
membrane sclerotized. Femora smooth ; anterior and median femora with tubercles on upper
and lateral surfaces ; anterior femora with a row of spines on lower outer surface and a single
spine on inner lower surface ; median femora with a spine on lower surface ; anterior and
median tibiae tuberculate ; anterior tibiae sinuate apically and with a projection sub-apically ;
tarsi tuberculate.

Type species : Karenocoris pustulatus sp. n.

Karenocoris pustulatus sp. n.

(Fig. n)

Colour : Antennae, head, thorax, legs, abdomen ventrally and corium, dark testaceous ;
head, pronotum and propleural epimeron suffused with piceous. Abdomen dorsally black ;
connexival segments with a small yellowish spot basally. Tubercles piceous. Tibiae suffused
with piceous apically

Structure : Basal segment of antennae feebly curved and feebly tuberculate, the tubercles
setigerous. Produced portion of vertex from above narrow with the sides parallel. Transverse

1 Karen, a Burmese people.

468 HENRICOHAHNIA BREDDIN AND ALLIED GENERA

sulcus on vertex situated behind eyes and interrupted medially. Ocellar interspace less wide
than distance between an ocellus and an eye. Anterior margin of anterior pronotal lobe
strongly tuberculate ; sulcus on anterior lobe very narrow, not extending to anterior or posterior
margins ; sulcus on posterior lobe very feebly indicated. Disc of scutellum deeply foveolate ;
produced apical portion horizontal. Abdomen dorsally smooth, except apical third of yth
segment, transversely rugose.

Total length, 11-30 mm.

Hemelytra, 7-60 mm.

Greatest pronotal width, 3-10 mm.

ic (holotype), Upper Burma, Seinghka Valley, Aduna confluence, 28-5 ^-97-35
E., 4,500, 3. v. 1926 (E. Kingdon Ward) (B.M. 1926-400).

Karenocoris inermis (Distant)
(Fig. 12)

Forestus inermis Distant, 1903, Ann. Mag. Nat. Hist. (7) 11 : 253.
Henricohahnia inermis (Distant), 1904, Fauna Brit. Ind. Rhynchota 2 : 389.

Colour : Testaceous. Dorsal surface of abdomen piceous ; connexivum dark testaceous
with a yellowish spot basally. Membrane infumate.

Structure. Basal segment of antennae feebly curved with short curved setae arising from
tubercles. Produced portion of vertex relatively short, thick, narrowly rounded apically.
Posterior lobe of pronotum rugose, tuberculate ; humeral angles sub-conical ; posterior margin
of lobe undulate with feeble median incision. Disc of scutellum damaged ; apex sub-acute.
Anterior tibiae with two spines sub-basally ; anterior femora with 7 spines on lower inner
surface and i spine on lower outer surface. Hemelytra extending just beyond apex of abdomen.

Total length, 13-00 mm.

Hemelytra, 7-50 mm.

Greatest pronotal width, 3-10 mm.

The type of Karenocoris inermis, a female, is from Mungphu (Atkinson Coll.,
Brit. Mus. 1892-6). In the British Museum there is also a female from Sikkim
(B.M. 1911-383).

Karenocoris badgleyi (Distant)

(Fig. 13)
Henricohahnia badgleyi Distant, 1909, Ann. Soc. ent. Belg. 53 : 373.

Colour : Antennae, head, posterior lobe of pronotum, piceous ; anterior lobe of pronotum
and rostrum, brown. Pleura, sterna, legs light brown ; pleura suffused with piceous. Abdo-
men dorsally black ; connexivum brown with a yellow spot on each segment basally ; abdomen
ventrally testaceous with brown suffusion laterally and dark brown tubercles. Corium brown ;
membrane hyaline, infumate.

Structure : Basal segment of antennae feebly curved with short, curved, spatulate setae
arising from tubercles. Produced portion of vertex moderately thick, about half as long as basal
antennal segment. Both lobes of pronotum smooth, tuberculate ; anterior lobe feebly de-
pressed medially ; tubercles on posterior lobe arranged more or less in transverse rows ;
humeral angles sub-acute ; median incision on posterior lobe of pronotum deep, acute angular.
Hemelytra extending just beyond apex of abdomen.

Total length, n-oo mm.

Hemelytra, 7-00 mm.

Greatest pronotal width, 3-00 mm.

FIG. 12. Karenocoris inermis (Distant).

FIG. 13. Karenocoris badgleyi (Distant).

HENRICOHAHNIA BREDDIN AND ALLIED GENERA 471

The type of Karenocoris badgleyi, a male, is from Assam ; collector W. F. Badgley
(B.M. 1906-185). There is also a male from the same source.

Malaiseana gen. nov. 1

Size small. Basal segment of antennae tuberculate, longer than anteocular ; postocular
longer than anteocular, somewhat globose then narrowed. Head cylindrical, shorter than
pronotum ; dorsally tuberculate ; vertex produced anteriorly ; gular region with spines.
Ocelli small, widely separated. Basal segment of rostrum extending beyond anterior margin
of eyes, less than half as long as segment 2 ; segments 2 and 3 straight. Pronotum wider than
long ; transverse sulcus indistinct medially ; both lobes tuberculate and medially longitu-
dinally sulcate, the sulcus not extending to posterior margin of either lobe ; posterior margin
of posterior lobe undulate. Scutellum longer than wide, triangular with apex produced.
External apical angle of connexival segments 5 and 6 produced ; external margins of segments
partly tuberculate. Abdomen tuberculate ventrally. Hemelytra with internal cell of mem-
brane very small ; base of cells of membrane sclerotized. Femora nodulose ; anterior femora
with spines on lower surface ; median femora with a single spine on lower surface ; tibiae
shorter than femora ; anterior tibiae with spines. Corium setose.

Type species : Malaiseana annulipes sp. n.

Malaiseana annulipes sp. n.

(Fig. 14)

Colour : Piceous. Segment 2 of rostrum dark brown. Acetabula, suffusion and spots on
pleura and posterior lobe of pronotum, testaceous. Connexival segments with a yellowish spot
basally ; abdomen ventrally testaceous with suffusion and tubercles piceous ; pygophore
piceous. Tibiae and posterior femora with a median yellowish annulation.

Structure : Basal segment of antennae with short, forwardly directed setae arising from
low tubercles. Produced portion of vertex sub-acute, feebly curved downwards, cylindrical.
Vertex medially and inter-ocellar area, smooth, without tubercles. Ocellar interspace less
wide than distance between an ocellus and an eye. Lateral angles of collar with low, conical
tubercles. Disc of scutellum feebly depressed. Hemelytra extending a little beyond apex of
abdomen. Anterior tibiae with 5 spines on outer and inner surfaces. Pale areas of con-
nexivum not tuberculate.

Total length, 10-00 mm.

Hemelytra, 6*50 mm.

Greatest pronotal width, 3-00 mm.

i( (holotype), N.E. Burma, Kambaiti, 7000 ft., i8.v.i934 (R. Malaise) (B.M.
1935-630).

Tapirocoris gen. nov. 2

Size small. Basal segment of antennae tuberculate, longer than anteocular, postocular
somewhat globose, constricted basally. Head shorter than pronotum ; dorsally and laterally
tuberculate ; vertex produced anteriorly. Ocelli small, widely separated. Basal segment of
rostrum half as long as segment 2, not extending to anterior margin of eyes ; segments 2 and
3 straight. Pronotum wider than long ; transverse sulcus interrupted medially ; anterior
lobe with setigerous tubercles laterally and some low, tubercles on disc ; posterior lobe smooth ;

1 Dedicated to Dr. R. Malaise of the Stockholm Museum.
* In allusion to the shape of the head.

FIG. 14. Malaiseana annulipes sp. n.

FIG. 15. Tapirocoris limbatus sp. n.

474 HENRICOHAHNIA BREDDIN AND ALLIED GENERA

anterior lateral angles acutely produced ; posterior margin of posterior lobe undulate ; anterior
lobe with a median, longitudinal sulcus. Scutellum longer than wide ; apex produced. Heme-
lytra with internal cell of membrane very small. Femora smooth, tuberculate ; anterior
femora with spines on upper and lower surfaces ; median femora with spines on lower surface ;
anterior tibiae with spines on lower surface ; anterior coxae tuberculate. External apical
angle of connexival segments 5 and 6 produced.

Type species : Tapir ocoris limbatus sp. n.

Tapirocoris limbatus sp. n.

(Fig. 15)

Colour : Testaceous. Head with three longitudinal stripes on vertex, produced portion of
vertex, postocular, except a spot basally, a spot between ocelli, and an elongate spot laterally,
pattern on anterior lobe of pronotum, sub-lateral area of posterior lobe, disc of scutellum, spots
on coxae and pleura, shining black. Abdomen dorsally suffused with red ; connexival segments
with a large piceous spot ; abdomen ventrally, meso- and metapleura with sparse small brown
spots. Base of veins of metathoracic wing suffused with red. Tubercles and spines on legs
piceous.

Structure. Basal segment of antennae with short, sub-erect setae arising from tubercles.
Produced portion of vertex sub-acute apically, curved downwards feebly ; sides feebly convex.
Vertex with a few low tubercles in linear rows on vertex and a few on vertex laterally ; trans-
verse sulcus wide. Ocellar interspace a little less wide than distance between an ocellus and
an eye. Posterior lobe of pronotum feebly, transversely rugose. Disc of scutellum depressed
and transversely sulcate ; apex very feebly produced. Hemelytra extending beyond apex of
abdomen. Anterior tibiae with 6 spines on outer and 5 spines on inner surface. External
apical angle of connexival segments 5 and 6 feebly produced, rounded.

Total length, 12-50 mm.

Hemelytra, 8-50 mm.

Greatest pronotal width, 3-50 mm.

i$ (holotype), Burma, Ruby Mines (Doherty) (B.M. 1911-383).

Key to Dicrotelus, Nyttius, Orgetorixa and Allied Genera

1. Brachypterous. Body narrow, elongate. Head and thorax without tubercles.

Segment 9 of abdomen in female strongly lobately produced . Dicrotelus Erichson.
-. Fully alate. Body elongate elliptical. Head and thorax tuberculate. Segment 9

of abdomen in female moderately lobately produced . . . . . . 2.

2. Internal cell of membrane absent ...... Orgetorixa China.

-. Internal cell of membrane present . . . . . . . . . 3.

3. Internal cell of membrane small, elongate, triangular .... Nyllius Stal.
-. Internal cell of membrane very small ......... 4.

4. Connexivum strongly tuberculate . . ' . . . . Neonyllius gen. nov.
-. Connexivum with a large tubercle at external apical angle of segments . . . 5.

5. Segments 7 and 9 of Connexivum produced, broadly rounded . . Paranyllius gen. nov.

Nyllius Stal

Nyllius Stal, 1859, dfv. Vet. Ak. Fork. 16 (8) : 365.

Nyllius Wygodzinsky, 1950, Proc. Linn. Soc., N.S.W. 75 : 87.

Corpus elongatum. Caput cylindricum, inter antennas sat longe spinoso productum. Antennae
geniculatae, articulo I capite paullo breviore, crassiusculo, reliquis gracilibus. Rostrum articulo

HENRICOHAHNIA BREDDIN AND ALLIED GENERA 475

basali brevi, secundo illo duplo longiore. Thorax media constricto, lobo antico muiti-spinuloso,
postico ante scutellum sat profunde sinuato, angulis lateralibus horizontaliter extrorsum spinoso-
productis. Scutellum apice acute productum. Tegmina alaeque completa. Pedes mediocres,
femoribus anticis incrassatis, posticis sub-aequelongis, subtus infra medium spina armatis ; tibiis
anticis femoribus sub-brevioribus. Abdomen margine spinulis obtusiusculis instructo ; segmento
apicali producto fisso .

Dicrotelo affine genus, corpore aspero, thorace constricto tegminibus completis distinctum.

Nyllius asperatus Stal

(Fig. 16)

Nyllius asperatus Stal, 1859, dfa. Vet. Ah. Fork. 16 (8) : 366.
Testaceus vel Jusco testaceus, parce sericeus.
Stal's description quoted above omits reference to the following characters :

Produced portion of vertex acute apically in lateral view ; produced portion of tylus some-
what narrow, rounded apically. Tubercles on head moderately long, slender. Spines on
collar, on anterior lobe of pronotum and at humeral angles, slender, sub-acute. Produced
portion of scutellum with erect, conical elevation apically ; apex sub acute and somewhat
curved downwards ; disc narrowly deeply depressed ; carinae moderately strongly tuberculate.
Hemelytra extending almost to apex of abdomen.

Stal's colour description does not agree with that of the type which is ferruginous with the
corium paler. Additional colour details are, clavus and membrane infumate, connexival
tubercles white and black, abdomen mid-ventrally pale testaceous.

Another specimen, without any locality label, but with a paratype label, differs from the type
in having only one membranal cell and is much paler in coloration.

Total length, n-oo mm.

Hemelytra, 6-50 mm.

Greatest pronotal width, 2-00 mm.

Stal's type, a female in the Stockholm Nat. Riksmuseum, is labelled Nova Hol-
landia, without precise locality.

Paranyllius gen. nov.

Size small. Basal segment of antennae shorter than head. Head longer than pronotum ;
almost smooth with very sparse tubercles ; vertex and tylus produced. Ocelli small. Basal
segment of rostrum extending to middle of eyes. Pronotum wider than long ; anterior lobe
longer than posterior lobe ; lateral angles of collar and humeral angles produced ; both lobes
smooth with a few tubercles ; posterior lobe with carinae sub-dorsally, anteriorly ; posterior
margin excised. Scutellum longer than wide ; produced apically ; carinae of disc tuberculate.
Hemelytra not extending to apex of abdomen ; internal cell of membrane very small. External
apical angle of connexival segments 2-6 tuberculately produced ; external apical angle of
segments 7 and 9 lobately produced ; abdomen ventrally tuberculate. Anterior femora incras-
sate and with a spine on lower surface, sub-apically ; all femora with a few tubercles ; anterior
and median tibiae shorter than femora.

Type species : Paranyllius turneri sp. n.

Paranyllius turneri sp. n.

(Fig. 17)

Colour : Testaceous. Metathoracic wings hyaline