Bulletin of the British Museum (Natural History)

BULLETIN OF

THE BRITISH MUSEUM

(NATURAL HISTORY)

ENTOMOLOGY
Vol. XXIV
1969 — 1970

BRITISH MUSEUM (NATURAL HISTORY)
LONDON: 1971

DATES OF PUBLICATION OF THE PARTS

No. i • « • • • '13 June 1969

No. 2 . , . . . .3 December 1969

No. 3 . . . . .3 December 1969

No. 4 . . . . .22 January 1970

No. 5 . . . ."•'... 30 January 1970

No. 6 . . . . . -30 January 1970

No. 7 . . . . . . .27 May 1970

No. 8 . . . . . .27 May 1970

No. 9 . . . . . . .24 July 1970

Printed in England by Staples Printers Limited at their Kettering, Northants, establishment

CONTENTS

ENTOMOLOGY VOLUME XXIV

PAGE

No. i. A key to the genera of the Menoponidae (Amblycera: Mallophaga:

Mallophaga : Insecta) By T. CLAY i

No. 2. The Type-Material of Tachinidae (Diptera) described by N. Baranov.

By C. W. SABROSKY & R. W. CROSSKEY 27

No. 3. Bombyliidae, and a first record of Nemestrinidae from Sokotra

(Diptera). By D. J. GREATHEAD 65

No. 4. Thysanoptera from the Solomon Islands. By L. A. MOUND 83

No. 5. A revision of the genus Catoptropteryx Karsch (Orthoptera: Tetti-

goniidae). By J. HUXLEY 127

No. 6. A list of the Type-Specimens of Odonata in the British Museum

(Natural History) Part III. By D. E. KIMMINS 171

No. 7. Studies of African Asilidae (Diptera) i. Asilidae of the Congo basin.

By H. OLDROYD 207

No. 8. A list of the Type-Specimens of Plecoptera and Megaloptera in the

British Museum (Natural History). By D. E. KIMMINS 335

No. 9. A revision of the Termites of the genus Macrotermes from the

Ethiopian Region (Isoptera: Termitidae). By J. E. RUELLE 363

Index to Volume XXIV 445

A KEY TO THE GENERA OF THE

MENOPONIDAE
(AMBLYCERA : MALLOPHAGA :
INSECTA)

THERESA CLAY

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 24 No. i

LONDON : 1969

A KEY TO THE GENERA OF THE MENOPONIDAE
(AMBLYCERA : MALLOPHAGA : INSECTA)

BY

THERESA CLAY

British Museum (Natural History)

Pp. 1-26; 7 Plates, 29 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY VoL 24 No. i

LONDON: 1969

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

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

This paper is Vol. 24, No. i of the Entomological
series. The abbreviated titles of periodicals cited follow
those of the World List of Scientific Periodicals.

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

Trustees of the British Museum (Natural History) 1969

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 13 June, 1969 Price

is.

A KEY TO THE GENERA OF THE MENOPONIDAE
(AMBLYCERA : MALLOPHGA : INSECTA)

By THERESA CLAY

SYNOPSIS

A review of the characters which have been used in the diagnosis of the genera of the Meno-
ponidae is given, together with a number not previously used. A discussion of their taxonomic
value is included, followed by a key to the genera and subgenera of the family.

PART I
INTRODUCTION

IN formulating a key to the genera of the Menoponidae an attempt has been made to
use characters which not only define each genus but also place at least some of the
genera in larger groupings; these may prove in some cases to be phylogenetic. The
greater number of species now available has shown that many of the former key
characters cannot always be used or have only a limited application. These include
presence or absence of postpalpal and antero ventral head processes; ctenidia on the
3rd femur and sternites; asters of spiniform setae on sternite II of Myrsidea;
subdivisions of the flagellum ; number of prosternal setae ; shape of head and abdomen ;
the characters of the dorsolateral margin of the head and hypopharyngeal sclerites.
These characters may vary within groups of otherwise similar species or may be
common to otherwise dissimilar genera. It is hoped that this key may give a clearer
conception of the generic characters and their reliability in the taxonomy of the family.
Only those structures of ectodermal origin and which can be seen satisfactorily
without sections have been considered; the task of sectioning and examining a
sufficient number of specimens of each genus would have been impossible and a
number of the genera are represented only by specimens mounted on slides. Some
characters have obviously been missed; also omitted are those which could not be
seen clearly in all the available material or the morphology of which could not be
correctly interpreted. There are also other characters which need further material
and time for study, such as the segments and sensilla of the flagellum.

There follows a review of the characters of various parts of the body with a
discussion of their taxonomic importance. These characters fall into four groups:
those which are found throughout the Menoponidae and which are of little interest
here, but are mentioned when they have been or might be included in generic
descriptions; those which vary throughout a genus and are of specific value only;
those which can be used for generic separation ; and those which group similar genera
together. The same character can of course be specific in one group and generic in
another. Two cases have been found in which a character, apparently unique for the
Amblycera, is of no more than specific or of species-group value (see Comatomenopon
elbeli Emerson abdomen and Meromenopon head setae). It is appreciated that the
term ' generic character ' is not very meaningful considering how the generic concept

ENT. 24, I. I

4 THERESA CLAY

differs among workers on Mallophaga at the present time; elsewhere (Clay, 1947 : 457-
477 i I95I : I7I~I75 ) r953 : 5^i i 1966 : 332-333) the present writer's views on the genus
have been made clear and need not be repeated. It is felt that at present there is
little to be gained by attempting to group the genera into various families and other
divisions.

Head

i. The Head Capsule. The basic structure of the head is similar throughout the
Menoponidae (Buckup, 1959; Haub, 1967), but the form of the dorsolateral margin
(Text-fig. 4) and its junction with the temple margin, the presence or absence of a
preocular notch or slit, position of the eyes and the form of the antennary fossa provide
useful generic characters. The presence or absence of a narrow preocular slit is
usually constant in a group of otherwise similar species, but in Menopon there is
variation from a deep narrow slit (M. interpositum Ansari) to a shallow narrow slit
(M. gallinae] to an approximately straight margin (M. jellisoni Emerson). M.
interpositum, for instance, with the abdominal characters typical of Menopon has the
dorsolateral margin of the head similar to that of Amyrsidea perdicis. Variously
developed ventral sclerotized processes arising near the anterolateral margin may be
present or absent (e.g. Pacifimenopon Price) or occasionally present in genera in which
such processes are not usually found (e.g. Machaerilaemus raggianae Price & Emerson,
Kurodaia quatei Price); they are probably of little phylogenetic value. Processes
arising near the base of the maxillary palp (postpalpal processes, Text-fig. 5) are
usually more constant within genera, but may be present or absent in Nosopon and
Pseudomenopon, for example; in the latter genus and in Colimenopon they may be
present in the nymph and absent in the adult; it is also doubtful whether their
presence or absence forms good generic characters for some of the species infesting
the Galliformes. The presence of these processes is therefore only specific in some
genera, but in others is probably generic (e.g. Kelerimenopon, Apterygon Clay and
Hohorstiella) . The gular region has two or more setae each side and may or may not
have a well-sclerotized plate; the form of this plate is a generic character in
Pseudomenopon and Colimenopon.

The tentorium has been described by Symmons (1952 : 365) who showed its
similarity in the species of Menoponidae she examined belonging to the genera
Colpocephalum, Menopon, Menacanthus, Myrsidea and Ancistrona. In mounted
specimens it is generally possible to see the anterior and posterior pits, the anterior
tentorial arms and the tentorial bridge; differences are shown in the width of the
bridge relative to the size of the head and the size of the posterior pits, but it is doubtful
whether these will provide useful characters for the recognition of genera. In Eureum
and Dennyus the bridge is narrowed centrally, but in some species this tends to be
rather wider and would probably grade into that of some species of Myrsidea, for
instance, which is broader than is usual in Dennyus but less broad than in some
species of other genera. Symmons (1952 : 375) showed that there were some
differences in the tentorium of Trinoton, mainly of muscle attachments, and of
Piagetiella, in which parts of the bridge and anterior arms are difficult to see owing to
some lack of sclerotization and to the heavy sclerotization of parts of the head.

KEY TO GENERA OF MENOPONIDAE 5

The degree of pigmentation of the internal carinae gives a characteristic appearance
to the head and may be correlated with other characters to help in generic groupings.
However, it seems to be a rather variable character; the quill-inhabiting species, for
instance, having a tendency to lighter pigmentation (e.g. Comatomenopori).

2. The Head Setae and Sensilla. These tend to form constant patterns (Clay,
1960 : 573) and the dorsal head setae expecially may be useful for the recognition

23 22

9 9 21

f-4-4

-u

FIGS. 1-3. i, Menoponidae head (dorsal) and pronotum, 2-3, Posterior part of dorsum of
head. 2, Menacanthus stramineus. 3, Colpocephalum sp. a-e. dorsal head sensilla; 8-31.
dorsal head setae; d.p.s. central pronotal setae 1-2; t. transverse pronotal carina.

of genera and generic groupings. They have been used extensively in the key,
being referred to by numbers e.g. seta 23 ; those of taxonomic importance have been
labelled in Text -figs. 1-4. On the anterior region of the dorsum there may be extra
setae in some species of a genus or in one sex only and the setae of this region have
not therefore been used. The lengths of 10 and n (the preocular setae) are useful
specific characters ; the addition of one or more setae to this pair may be specific
(Adornithophilus crinitus Clay) or generic (Meromenopon). Ancistrona has a row of
fine setae along the inner dorsolateral margin (Text-fig. 20) not seen in any other

6 THERESA CLAY

genus. Numbers 14-16 (setal complex and dorsal seta of Price & Beer, 1963 : 851)
usually form a characteristic group, their position in relation to each other often being
of specific importance. The size and position of 17 and 18 (the mid-dorsal head setae)
may form generic (Cuculiphilus] or specific characters (e.g. Colpocephalum) ; the
absence of 18 is a generic character in Myrsidea. The two ocular setae (19 and 20)
seem to be always present : 19 usually lies near the division of the two ommatidia
(when apparent) ; 20 is usually small and often marginal and difficult to see ; in Dicteisia
it is well developed (Price, 1968) . The posterior marginal or submarginal temporal and
occipital setae are numbered from the centre outwards as the first ten form useful
taxonomic characters, whereas the anterior temporal setae are not always constant in
number, and in mounted specimens may get confused with the ventral. The occipital
setae (21-22) are always present and their relative length and thickness may form
specific or generic characters. Seta 23 may lie anterior or anterolateral to 22 (Text-
fig, i) or in a straight line with 21 and 22 (Text-figs. 2-3), often some considerable dis-
tance across the temple; it is important to identify it correctly and not confuse it with
24 or 25. In Myrsidea seta 23 is always absent and in Bucerophagus it may be min-
ute. Of the seven following marginal or submarginal setae (24-30), 29 and 30 in
nearly all species can be recognized as 30 lies near and submarginal to 29, either being
on the outer or on the mediad side ; in Trinoton it is nearer seta 27 ; in Numidicola
it is marginal. The next marginal seta is 28, usually short and fine; followed by 27
and 26 which are either separated or have their alveoli contiguous or nearly con-
tiguous; in this last condition 26 is usually fine and significantly shorter than 27.
The position of these two setae is a constant feature within genera and groups of
genera and is a useful key character. However, it does separate Bucerophagus, in
which 26 and 27 are closely associated, from the otherwise similar genera Chapinia
and Bucerocolpocephalum Elbel in which they are not. This leaves two setae, 25 and
24, one or both of which may be minute to medium to very long; the length of these
setae may be a useful character, but in Machaerilaemus one or both may be missing
and in other genera the lengths may vary. In those species with one long and two
short setae between 23 and 27 (Text-fig, i) it is not always possible to number the
long seta; it is usually submarginal to the two short marginal setae and may vary in
its position relative to them. Ancistrona has extra setae: three long and two short
between 23 and 28. In addition to these temporal setae, one of the setae anterior
to 30 (here called 31) may be long and stout. The number of long setae may be
generic (three in Eomenopon] or specific (two or three in Kurodaia, Price & Beer,
I963 : 851). The lengths of the head setae are not always key characters owing to
the known and potential variation and to the subjective interpretation of ' long ' and
' short '.

The dorsal head sensilla may be five in number (Text-fig, i), but there are more
usually three: a. is not always apparent as it lies on the margin just anterior to seta
9; b. just posterior to 9 is useful for the identification of this seta; c. is usually present
and associated with setae 14 and 15; d. associated with seta 16 and e. with 17 are
less commonly present. The presence or absence of e. is a group character in
Austromenopon, being present only in the species infesting the Alcidae; d. is absent in
Actornithophilus species from the Lari, being present in species from other hosts; in

KEY TO GENERA OF MENOPONIDAE 7

Menacanthus from the Galliformes d. may be present or absent in different species.
The absence of c. may be a generic or group character: in the Colpocephalum-complex
c. is usually present, but appears to be absent in the species infesting the Galliformes
and in a few of those on the Ciconiiformes.

The ventral head setae are mainly constant in number and position throughout
the Menoponidae. The labral setae comprise a posterior row of usually closely
arranged setae and an anterior row often with i, 3, 6 and 7 short and the remainder
longer (Text-fig. 4). An examination of many species, however, shows that there is
no clear distinction between ' long ' and ' short ' setae and there are many exceptions.
This is not therefore usually a useful character, but in Ancistrona these setae are

FIGS. 4-6. 4, Myrsidea sp. head: m. 3 postmental setae; a. the 2 anterior subocular setae;
5. subocular seta; c. subocular comb row; d. dorsolateral margin of head. 5, Menacan-
thus stramineus: part of venter of head; m. 4 postmental setae; mp. base of maxillary
palp; p. postpalpal process. 6, Trinoton sp. antenna.

diagnostic, being all minute with the exception of one long seta each side. In all
species examined there are two setae, one long and one short, at the anterior end
of the ventrolateral margin (Text-fig. 4 a .), posterior to these are one or more setae
(s. subocular setae, Ryan & Price, in press), followed in most genera by the subocular
comb row ( = lateroventral head fringe, Clay 1966 : 330). The first or only seta
in group s. is frequently elongate and sometimes flattened and somewhat hyaline
(e.g. Eidmanniella ; PI. 4, fig. 20) ; in Meromenopon from the Meropidae this seta is
flanged with at least one tooth (PI. 3, figs. 15-16), being unlike any other seta seen
in the Mallophaga ; in the congeneric species from the Coraciidae the seta is normal.
The presence of only one seta in group s. may be a specific or generic character; in
Eomenopon and Pacifimenopon the elongate seta may not be the most anterior one.
In some cases it is not always possible to separate some of the setae of group s.
from the comb row. The subocular comb row (PI. 4, fig. 19) is characteristic of the

8 THERESA CLAY

Menoponidae, but is absent or atypical in Microctenia, Machaerilaemus and A ncistrona ;
in the latter genus its function may have been taken over by the row of setae along
the inner edge of the dorsolateral margin. In some genera (e.g. Colpocephalum)
there is a ventral patch or line of submarginal setae on the anterior region of the
temple (PI. 4, fig. 21) ; this should not be confused with the subocular comb row and
the seta continuous with this row which in mounted specimens may appear submar-
ginal. In Kelerimenopon a band of thickening runs inwards from the ventrolateral
margin with at least one seta. The histology of the setae and integument of the
Menoponidae is discussed by Neuffer, 1954.

3. The Antenna. In all Menoponidae the antenna (PL I, fig. i) comprises the scape,
pedicel and a flagellum of usually two segments, the terminal one sometimes being
subdivided. The distal anterior angle of the scape and pedicel may be produced
laterally (Trinotori) or the pedicel alone (Mimemamenopon Carriker) forming generic
characters; the latter condition may also be only specific (Ciconiphilus, Hohorstiella).
In one species of Eidmanniella three of the distal setae of the pedicel are broad and
hyaline (Ryan & Price, in press). The first segment of the flagellum is always
pedunculate or wineglass-shaped. Ferris (1923 : 57) discussing Menopon and
Tendeiro (1967 : 384) Chapinia, considered that this segment was divided into two,
the first being small and formed by a line across the ' stalk ', the part proximal to
the line being darker in colour. This condition is apparent in many species belonging
to many genera : in Bucerophagus productus there is no sign of an external structural
division (PL I, fig. 2) and no break of the internal marginal thickening; the apparent
segmentation may be due only to the difference in pigmentation. In Myrsidea
cornicis and Pseudomenopon pilosum in which this line is apparent, reconstructions
of the antennae from sections by Buckup (1959) and Haub (1967) show no break in
the pedunculate segment, these authors describing it as a single segment. It would
therefore be more satisfactory to treat it as such. Keler (1958 : 82) in the original
description of Eidmanniella stated that the antenna appeared three-segmented as
there was only a trace of the line of division between the two segments of the
flagellum; stereoscan photographs of this type of antenna (PL i, fig. 4) show a definite
division, but perhaps not so marked as in some other species (PL 3, figs. 13-14). The
second segment of the flagellum, usually referred to as the terminal antennal segment,
may be globose to elongate, the shape frequently being similar in the species of a
genus. The surface of this segment is ridged (PL 2, figs. 7-12) and it is these ridges
when deep and in a straight line which may give a false impression of a subdivision
of the segment. It is possible that the form of these ridges and the sculpturing of
their edges (PL i, fig. 3) may provide further taxonomic characters. The distance
apart of the ridges, visible with the light microscope, has been used in the key to
separate two groups (PL 3, figs. 13-14). The form of the sculpturing of the third
segment may also prove to be a taxonomic character. Distally there are a number of
setae, sometimes cone-shaped (PL 3, fig. 13) and two sensilla which seem to be
sensilla coeloconica (PL 3, fig. 13; PL 7, fig. 42). In species with the last segment
subdivided, one of these sensilla is proximal to or near the dividing line (Cuculiphilus,
PL I, fig. 3), suggesting that the primitive condition may have been a three-segmented
flagellum with a sensillum on each of the two terminal segments. Whether the last

KEY TO GENERA OF MENOPONIDAE 9

segment is subdivided or not has frequently been used as a generic character, but it
now appears that this may not always be reliable. In Cuculiphilus there is a well
marked division internally and externally (PL i, fig. 3). In Bucerophagus productus
there is an internal break in the marginal thickening and the part proximal to this is
slightly more darkly pigmented, but externally there is no definite division (PL i,
fig. i); one of the sensilla lies proximal to the internal division. Some species of
Menacanthus also show a break in the internal marginal thickening of the terminal
segment without any other indication of a division, so that the presence or absence
of an internal break may not be relevant. It would seem that in B. productus the
appearance of a division is partly due to the slight change in pigmentation and
partly to a deep furrow between the ridges (PL 2, figs. 7-9). In B. africanus similar
photographs (PL 2, figs. 10, 12) suggest that there is a definite division; there is also
a marked difference in pigmentation in this species, the part proximal to the line of
division being darker in colour. This segment in some species of Colpocephalum
resembles that of B. productus in having the proximal part more darkly pigmented
and the two sensilla widely separated. In Plegadiphilus the presence of a subdivided
terminal antennal segment has again been incorrectly used as a generic character
(Clay, 1947; Blagoveshtschensky, 1964). In some species of this genus the
pigmentation and the ridges, together with the position of the proximal sensillum and
an internal break in the marginal thickening, gives the impression of one or more
subdivisions (PL 3, fig. 14). In Plegadiphilus plegadis, in which this segment is
short there is no indication of a division and the two sensilla are close together. In
Austromenopon one species (A. affine] shows internally and externally a line of division
with the sensillum just proximal to the line, others show some indication of a line of
division, while others show none. It is not possible therefore to use this character as
a major division in the key.

A preliminary study of the position of the sensilla and their associated setae as
possible generic characters has shown considerable variation within groups of otherwise
similar species. In many species the two sensilla are near each other at the distal end
of the segment and their surface apertures are similar; in Bucerophagus the two
sensilla are widely separated and the proximal one lies in a circular pit (PL 2, fig. n).
Menacanthus stramineus has the terminal antennal segment (PL 3, fig. 13) typical of a
number of species of Menacanthus from the Galliformes and Passerif ormes ; in this
the distal sensillum is on the end surface of the segment with the majority of the
setae, while the proximal is nearby with two or three setae which arise from a slight
indentation. This differs from the elongated last antennal segment of Menopon
and Amyrsidea in which there is no indentation and the two sensilla and the two
lateral setae are close together on the end of the segment. However, other species
of Menacanthus (that from Arborophila, for instance) in which the segment is
elongate, the condition is similar to that of Menopon; and the Menacanthus from
Alectoris in which the last segment is short, the sensilla are close together and the
setae are merged with the group of terminal setae and no longer associated with one
of the sensilla ; this arrangement is also found in many of the species parasitic on the
Passeriformes. Thus, the position of the sensilla is not necessarily dependent on the
length of the segment ; there is some indication that the wide separation of the sensilla

io THERESA CLAY

is associated with a tendency in the group towards subdivision of the segment. In
Austromenopon the proximal sensillum may be near the distal end or at various
positions in the distal part of the segment, being found nearest the base in A. affine,
in which it lies near the line of division. The species of this genus probably all have
two of the stout setae placed near the middle of the lateral margin of the terminal
segment. It is apparent from the above survey that these characters will not
provide any major divisions for the key but further studies of all the antennal
characters may reveal some patterns of taxonomic interest.

4. Eyes. Wundrig (1936) has shown that the Amblycera have two ommatidia
each side and these show all stages from eyes with well-developed biconvex lenses
(PI. i, fig. 6) to those with no lens. As it is doubtful whether without sections it is
possible to state the exact stage of development of the ommatidia, the character of
' eyes present or absent ' has not been used. The lenses are usually located on the
dorsolateral margin but in Pseudomenopon and part of Eidmanniella (new genus,
Ryan & Price, in press and see Clay, 1957 : 143) the eyes are more towards the centre
of the head (PI. i fig. 5).

5. Mouth Parts. With the exception of the hypopharynx these are similar
throughout the Menoponidae (Buckup, 1959; Haub, 1967). The maxillary palp is
four-segmented, the last segment usually having two well-marked sub-terminal
setae, the relative sizes of which may be a specific character (Clay, 1968, PL i, figs.
6-7). The alveoli of these setae are usually contiguous, but may be separated by a
definite gap, this appearing to be a constant character in Dicteisia, Clayia and
Somaphantus; both conditions are found in species of Menacanthus parasitic on the
Passeriformes. In some species these setae are not apparent and perhaps merge
with the terminal setae; in Cuculiphilus sens. lat. there is a third seta associated
with this pair (PL 7, fig. 45). Each labial palp has five anterior setae, in Neomenopon
one is fine and not apparent in all specimens. The number of setae on the prementum
appears to be constant ; the lengths of one of these setae has been used as a species-
group character in Actornithophilus (Clay, 1962 : 237). The postmentum usually
has four setae each side; (Text-fig. 5) ; in a few genera (e.g. Myrsidea, Text-fig. 4) the
most posterior seta each side is missing and in Nosopon milvus Tendeiro it is replaced
by a clump of four to six small setae. The hypopharyngeal sclerites and the
functionally associated epipharyngeal crest show various degrees of development
which appear to be of little phylogenetic importance (Clay, 19626 : 220) : otherwise
similar species may have the sclerites well-developed or reduced (Austromenopon,
Clay, 1959 : 159; Myrsidea, Clay, 1966, PL 2, figs. 1-2). In Neomenopon the form
of the hypopharynx is unusual and may prove to be a good generic character. The
distinctive epipharyngeal organs (Buckup, 1959 : 262) appear to be present in all the
Menoponidae.

Thorax

Especial attention has been paid to the thorax and legs which provide many useful
generic and subgeneric characters. A detailed study has been published by Mayer
(1954) and only those characters of taxonomic value will be discussed here.

KEY TO GENERA OF MENOPONIDAE n

I. Prothorax. The transverse carina of the pronotum (Text-fig, i) is apparent
in all genera with the exception of Rediella ; the vertical carina shows various stages
of development. In some species there is a vertical groove each side of the pronotum
giving the so-called ' winged ' or ' lobed ' prothorax. The postnotum (sens. Mayer,
10,54 = mesonotum sens. Cope, 1941) in the majority of genera is a well pigmented
oblong sclerite (Text-fig. 8), often distorted in mounted specimens; it may not be
apparent (Numidicola) or it may be of a different shape (Ancistrona). The prosternal
plate varies from being well developed to greatly reduced ; in Eidmanniella a posterior

rv

mt,

FIGS. 7-9. 7, Heleonomus sp., mesothorax: i, ist coxa with the 5 posterior coxal setae
2, 2nd coxa; ms. mesosternal plate; mt. metasternal plate. 8, Colpocephalum sp.,
pronotal margin and mesonotum: ps. pronotal marginal setae; pn. postnotum; mn.
mesonotum; as. anterior mesonotal setae. 9, Actornithophilus sp., postnotum and
anterior mesonotal setae.

process of the prosternal plate may be strongly or weakly sclerotized or absent.
There are usually two pairs of central pronotal setae (dps) lying on or near the
transverse carina, but in some species-groups (Clay, 1962 : 237) or genera (Myrsidea)
they are reduced to one pair or absent. Although the number and lengths of the
posterior marginal setae of the pronotum may be useful specific or species-group
characters, they are not necessarily of generic importance. All species have two
small central prosternal setae; additional setae may be of generic importance,
but in some genera both conditions may be found (e.g. Ciconiphilus, Clayia).

2. Mesothorax. The mesonotum is developed to a greater or lesser extent, the
differences not being sufficiently clear cut to use as a generic character. There are
two different types of mesosternum: in the majority of genera (Text-fig. 7) the
sclerite (part of the episternum) bearing the inner articulation of the leg is separated

12 THERESA CLAY

from that of the other side by a distinct mesosternal plate or by an area without a
definite plate. In the other type (Myrsidea, Cuculiphilus] the mesonotum, pleura
and mesosternum are fused to form a sclerotized ring round the body (Clay, 1966,
PI. i, fig. 6). The anterior mesonotal setae may be four in number clustered round
the distal end of the postnotum (Text-fig. 8); in some species of Odoriphila the
setae of the pair each side of the postnotum lie close together and in some specimens
one seta may be hidden below the other giving at low power the appearance of only
two. Less commonly the outer setae may be widely separated from the inner
(Actornithophilus, Text-fig. 9) or there may be only two (Myrsidea). These setae
are constant in position and number within genera and form useful key characters.
In addition, the mesonotum has at least one other seta each side, lateral or postero-
lateral to the anterior mesonotal setae, and it is important to identify these before
deciding whether there are two anterior mesonotal setae or four widely spaced ones.
The centre of the mesosternum usually has four or more setae, but in some genera
there are only two (Austromenopori) ; this is usually a constant character within a
genus or groups of genera but in Bucerophagus there may be two or more setae.

3. Metathorax. In Myrsidea some species may have the metanotum strongly
modified while in other similar species it is normal, this character therefore seems to
be of little phylogenetic importance (Clay, 1966 : 331). In Clayia and at least some
species of Menopon, there is a variously developed central vertical line of thickening
in the anterior part of the metanotum ; however, some of the species of Menopon are
based on specimens not in sufficiently good condition to say whether this thicken-
ing is present, but it may prove to be a good character for these two genera. The
outer seta at each end of the posterior marginal or submarginal row of metanotal
setae is long or the longest of the row ; it is sometimes anterior to the rest of the row.
The presence of many central setae on the metanotum may be a specific character
(Actornithophilus}. There is usually a central metasternal plate with setae.

The species of Trinoton have two large thoracic sternal plates with many setae;
caudad to the posterior plate is a bilobed flap appearing white in untreated specimens
(PI. 5, fig. 25). Species of Eureum also have a white flap arising from what is
probably the metasternum and a similar one on abdominal sternum I (PI. 5, fig. 26)
and in Dennyus on the metasternum and a number of the abdominal sterna. In
Trinoton the surface sculpture (PI. 5, figs. 28, 30) of the flap is similar in specimens
parasitic on species belonging to five genera of the Anseriformes, but differs from that
of the flaps in Eureum (PI. 5, figs. 27, 29) and Dennyus (PL 3, fig. 17). The function
of these areas is unknown. Species of Trinoton also have a conspicuous white area
surrounding the gular plate (PI. 5, fig. 25), but the surface sculpture is quite distinct
from that of the metasternal flap (PI. 3, fig. 18).

Legs

The gross morphology of the legs is similar throughout the Menoponidae (see Mayer,

1954)-

i. Coxa. These are attached to the ventral part of the thorax except in those
genera (e.g. Trinoton, Eureum) with wide sternal plates which cause the articulation

KEY TO GENERA OF MENOPONIDAE

to be more lateral; in the former genus the usual anterior prolongation of the first
coxa is greatly reduced. The first coxa frequently has four or five posterior setae
(Text-fig. 7.), but in certain genera (Austromenopon) or groups of genera there are
more; however some of the species may have only one or two extra setae with the
occasional specimen without these (in Eidmanniella) ; this character cannot therefore
always be used. Those species with many setae on coxa I may also have a greater
number on II and III.

2. Trochanter. Ventrally there are a number of well marked sensilla associated
with two setae (Text-fig. 13), the most usual number on legs II and III being four
to five, but some genera (Bonomiella) or species-groups appear to have a constant
number of three. The number is of doubtful general use in the key as it is sometimes
only specific (Nosopon) or there is the occasional specimen in which it varies on differ-
ent legs; however, in some species groups of Menacanthus (see below, p. 18) the
number is correlated to a certain extent with other characters.

3. Femur. The most important taxonomic character of this segment is the ventral
chaetotaxy of the third femur. This may be in the form of patches of irregularly
arranged setae (brushes, Text-fig, n) or regular rows of stout spine-like setae (combs

13

\

14

FIGS. 10-14. lo-ii, Menoponidae legs. 10, ist, ventral: d. ist outer dorsolateral tibial
seta; v. 2nd. outer ventrolateral tibial seta, n, 3rd, ventral: b. brush of setae. 12,
Bonomiella sp. claw. 13, Trochanter with ventral sensilla and setae. 14, Venter of 3rd
femur with ctenidia.

14 THERESA CLAY

or ctenidia, Text-fig. 14, PI. 6, fig. 31) or a few scattered setae. The form tends to
be constant throughout groups of otherwise similar species and frequently provides
useful generic characters. The presence of ctenidia is a group character in the
Colpocephalum-complex, but they are also found in other genera : Bucerocolpocephalum
with ctenidia is otherwise similar to Bucerophagus and Chapinia without; in
Piagetiella they may be present or absent in different species. Microctenia, in
addition to a brush of setae, has the venter of the third femur covered with rows of
comb-like outgrowths (PI. 6, fig. 32).

4. Tibia. Here again the chaetotaxy shows good taxonomic characters: the
number of outer dorsal setae may form useful specific characters (Clay, 1962 : 195 ;
1966 : 334, Text-fig. 10). Many species in which there are many outer dorsal setae
on tibia I, have only a few on II and III (Text-fig, n), while in others (Colpocephalum-
complex) there are numerous marginal and submarginal setae in this position (PI. 6,
fig. 33). Hoazineus and Heleonomus (PI. 6, fig. 34) have a row of short regular setae
along the outer margin of I-III. In some genera the extra tibial setae may be present
or absent (Falcomenopon, Emerson & Elbel and some species of Kurodaia) and in
Osborniella the number of the tibial setae is fewer than in most species of the
Colpocephalum-complex. There is considerable difference in the size and position
of the terminal ventral tibial setae, but no clear distinctions could be found as they
grade from relatively fine to thick spine-like setae and the distance between them
shows all stages from two of the alveoli being contiguous to all four being widely
spaced; it was decided that this is not a practical generic character. One genus
(Piagetiella) has a tibial spur in the male.

5. Tarsus. Keler (1952, 1955) and Mayer (1954) give descriptions of this joint in
some species of the Menoponidae. The tarsus comprises two segments, the proximal
being small and the distal being longer and of various proportions. The pretarsus
bears two claws which articulate with the well-sclerotized unguifer; the shape of the
claws may be diagnostic (Bonomiella, Microctenia) . The unguitractor in the adult is
in the form of two plates to which is attached the tendon-like apodeme of the
retractor muscle of the claws, usually visible in specimens treated with KOH.
Distally the dorsal part of the unguitractor may be elongated laterally forming two
pointed processes (Neomenopon, Hohorstiella) or there may be a central comb-like
area (Bucerophagus). Arising from the ventral part of the unguitractor on legs II
and III is a hyaline, sometimes tuberculate process (PI. 6, fig. 35), the empodium
(sens. Keler, 1952) ; the shape of this may be diagnostic (Clay, 1966) ; it may be small
and is sometimes not apparent and perhaps absent. Owing to the difficulty of seeing
its true form in mounted specimens, the empodium has not been used here as a
taxonomic character. The first tarsal segment has a pair of setae usually hyaline
and sometimes flattened; just distal to these is a pad-like lobe, the euplantula
(sens, Keler, 1952). Examination of sections and of the whole structure with the
light and scanning electron microscopes (PI. 7, figs. 37-41) suggests that the euplantula,
in at least some species, has an outer ventral membrane covering a honey-combed area
within which is a framework of vertical strands (PI. 7, fig. 41) or of vertical and hori-
zontal strands giving a characteristic banded appearance (PI. 7, fig. 40). The form
of these strands may be useful taxonomic characters and appears to be constant

KEY TO GENERA OF MENOPONIDAE 15

within genera and groups of genera ; all the members of the Colpocephalum-complex,
for instance, have the vertical strands only; other genera not belonging to this
complex also with vertical strands only are Microctenia, Hoazineus, Bonomiella and
Trinoton. However, it is not possible to use this character for basic divisions in
the key as there are a number of genera in which no internal striations can be seen
and in Menopon it has been possible to see the striations in only some of the species.
The euplantula of the second tarsal segment may be similar to that of the first
(e.g. Bucerophagus) or show considerable differences : in some of the species in which
euplantula I shows vertical striations only, II may be elongated to more than half
the length of the tarsus with vertical striations and deeply serrated margins (e.g.
Turacoeca] . In this position in Pseudomenopon there are rows of comb-like processes
(PL 6, fig. 35) ; Keler (1952 : 581) suggested that this structure was homologous with
euplantula II. Its presence enables the nymphs of Pseudomenopon, in which the
characteristic gular plate is not developed, to be recognized generically. Examination
of this area with the scanning electron microscope suggests that in some species the
form of the processes may prove to be a diagnostic character. The characters of
euplantula I, are probably similar on all three legs, but are usually best seen on the
first leg as in mounted specimens this is more often lying in the dorsoventral plane.
Pad-like structures along the inner side of the claw can be considered as pulvilli;
it is not possible to say whether their degree of development is of taxonomic use.
Certain other characters of the tarsus are not used owing to the difficulty of seeing
them in all species or in all specimens of a species.

Abdomen

The abdomen varies greatly in shape and in those species in which the plates are
not heavily sclerotized the proportions can be affected by the treatment of the speci-
mens. The shape of the abdomen (together with the rest of the body) may be an
adaptation to some particular factor of the environment, such as the inside of the
quill (Clay, 1962 : 192; Tuff, 1967 : 247). A genus based on such characters might
have been derived from different stocks (e.g. Somaphantus, see below, p. 19) and the
species might have lost many of the characters showing their affinities: Rediella
with a distinctive appearance, resembles Actornithophilus from the same host order
in the characters of the male genitalia and the spacing of the anterior mesonotal
setae.

There are six spiracle-bearing segments (III-VIII) and two (I-II) anterior to these;
posteriorly to tergite VIII, there is usually a single sclerite, but some species have
two. In Myrsidea females the terga may be strongly modified and tergite I not
apparent or with II, greatly reduced in size; these modifications, as that of the meta-
notum, seem to be no more than specific (Clay, 1966 : 331) ; members of the
Colpocephalum-complex may also show tergal modifications in the female. Sternite
I is usually apparent but is not so in Aegypiphilus; sternites II-VI appear as discrete
central plates; VII may be fused or partly fused with the following sternites to
form a subgenital plate ($ Myrsidea, Clay, 1966, fig. 26; Chapinia, both sexes) or
VII may be separated from the subgenital plate (<$ Myrsidea, Clay, 1966, fig. 27;

16 THERESA CLAY

Actornithophilus, both sexes); in females of the Austromenopon species parasitic on
the Procellariformes both conditions of VII are found, so that this is not necessarily
a generic character. Rarely the males show modifications of the sternites as in
Cacamenopon Price (sternites VI-VII) and Holomenopon goliath Clay (sternites
VIII-IX). Post vulval sclerites may or may not be apparent; their chaetotaxy
is sometimes a constant and generic character (Kurodaia and Nosopori).

The pleurites are usually in the form of discrete plates separated from the sternites
by a membraneous area, frequently sculptured, and from the tergites by a narrow
suture. In Piagetiella there may be some fusion between pleurites and tergites,
either sexual or specific ; in the female Myrsidea with modified abdomens they may be
reduced, absent or modified in various ways. In Comatomenopon elbeli the female
has sucker-like organs on pleurite III. The inner postero ventral angle of some or
all of the pleurites may be prolonged as a process (Text-fig. 26) ; this character
appears to be generic in some groups (Plegadiphilus) , but is present or absent in
others (Menacanthus from the Gallif ormes) . Patterns of internal thickening of the
pleurites and lateral areas of the tergites may be only of specific value (in
Austromenopon for instance) and are perhaps not useful generic characters.

The female anogenital region shows considerable variation: the typical anal
corona may be present or absent in the species of Austromenopon parasitic on the
Procellariformes (Clay & Moreby, 1967 : 158) ; some species of the Galliformes-
infesting Menoponidae may also lack the typical anal corona and show various other
modifications of the venter of the terminal segments; however, these are not always
correlated with other characters showing differences and may not be of any phylo-
genetic significance. The lateral edges of the anus may show various setae-bearing
processes which appear to be constant in certain groups and generic in character
(Turacoeca, Chapinia). The male genitalia may be similar throughout a genus or
genera, with the occasional species being distinct (e.g. Menopon). In dealing with
such males and females it does not seem reasonable or useful to erect a new genus
because one sex shows some unique character, while the other sex is not separable
from the rest of the group. This is illustrated by the genus Menopon in which
there is much diversity of the female anogenital region and the male genitalia. In
M. gallinae the terminal segment of the female abdomen is elongated, the anterior
(ventral) margin of the anus is widely separated from the posterior (dorsal) margin
which is terminal and beset by a row of spine- like setae; in pattens and interpositum
the anal margins are not so widely separated and the anus appears more normal; in
spinulosum the last segment is not elongated, the setae surrounding the anus form a
triangle, the terminal margin of the abdomen does not bear the posterior anal setae,
but has a number of long and short setae. The male genitalia also show considerable
difference between the typical gallinae form and those with the greatly enlarged and
asymmetrical parameres of the spinulosum group.

Various structures associated with the female genital chamber (Clay, 1961, fig. 7«z;
Price, 1966 : 18) and the form of the bursa copulatrix (Clay, 1968 : 207) provide useful
specific and sometimes generic characters. Although the presence or absence of
spermatophores in the male cannot be used as a key character it is possible that
their distribution within the genera of the Menoponidae may be of taxonomic

KEY TO GENERA OF MENOPONID AE 17

interest. They are probably present in all Myrsidea (Clay, 1968 : 207) and have also
been seen in species belonging to other genera (e.g. Austromenopon, Ciconiphilus,
Dicteisia) .

The spiracles usually open on the tergites, but in some species of Myrsidea they
open on the pleurites or the membraneous area between tergite and pleurite; in
Colpocephalum heterosoma the spiracles open on the pleurites in the female and
on the tergites in the male (Price, 1965 : 128). Although the presence of crop teeth
has been used as a generic character they seem to be present in all the Menoponidae :
further dissections of suitable material are necessary to see whether they will show
any taxonomic characters.

The Chaetotaxy of the A bdomen. All species examined have a small anterolateral
seta each side of tergite I and II. At each end of the posterior row of seta, or
somewhat submarginal to it, on tergites II-VIII is the post-spiracular seta with the
two small associated setae (Clay, 1954 : 716) ; on tergite I the seta in this position is
usually long and is included under the post-spiracular setae. A generally constant
character is the presence or absence of a small seta laterad to each of the long outer
setae on tergite I or to the post-spiracular setae on one or more tergites. Setae may
be present or absent on sternite I in different species belonging to the same genus
(e.g. Myrsidea}. The presence of ctenidia on one or more sternites is usually
associated with similar ones on the venter of the third femur (Colpocephalum-
complex); in a few genera (e.g. Piagetiella, Eomenopon) there may be abdominal
ctenidia but none on the femur. The position and the number and size of the setae
in the sternal brushes may be a useful generic character.

Part II

KEY
INTRODUCTION

The key includes all the generally recognized genera and subgenera (with the
exception of those belonging to the Colpocephalum-complex and the Menacanthus-
complex) even where it is considered that there is no advantage in the recognition
of some of these taxa. The Colpocephalum-complex, as interpreted here, comprises
all those genera with ctenidia on the venter of the 3rd femur with the exception of
Cuculiphilus sens, lat., Bucerocolpocephalum, Piagetiella, Turacoeca and Odoriphila;
it is possible that the last genus should be included. Dicteisia may belong to the
complex but is separated in the key. Other characters common to the complex are
the contiguous alveoli of head setae 26 and 27 ; setae 24 and 25 are usually minute to
short, in a few species they are longer, but do not reach to the end of the pronotum;
the presence of a vertical patch of submarginal setae on the temples; head sensilla
c. usually present; four anterior mesonotal setae; usually more than two central
mesosternal setae; mesosternum normal; tibiae II-III usually with submarginal
patch or row of outer distal dorsal setae, euplantula with vertical striations only;
ctenidia on one or more sternites. Baiter (personal communication and see Baiter,
1968) has found that the operculum of the egg opens diagonally in species of the
complex from the Galliformes, Falconiformes, Ciconiiformes, Pelecaniformes,

ENT. 24, I. 2

i8

THERESA CLAY

Psittaciformes and Passerif ormes ; in addition this character is found in the eggs of
Nosopon, but not of Osborniella or any of the species of Ciconiphilus examined.

The Menacanthus-complex comprises Menacanthus, Amyrsidea, Argimenopon
Eichler, Cracimenopon Carriker and Desumenopon Carriker, its species being parasitic
mainly on the Galliformes and Passeriformes. Divisions within this complex are
dependent on which characters are used, these include : width of head and form of its
dorsolateral margin; form of gular and prosternal plates; development of the
hypopharynx; presence of postpalpal processes; shape of the antennal segments;
number of setae on coxa I; number and position of the sternal brushes of setae;
presence of sternal spiniform setae; position of the post-spiracular setae on tergites
I-II; degree of development of the internal tergal and pleural thickening; presence
of a prolongation of the posteroventral corner of the pleurites; terminal segments
of the female abdomen and the male genitalia. A distinctive species group (including

19 20 21 22

FIGS. 15-22. Heads of Menoponidae genera. 15, Odoriphilia. 16, Neomenopon. 17,
Meromenopon. 18, Colimenopon. 19, Pseudomenopon. 20, Ancistrona. 21, Dennyus.
22, Gruimenopon.

the type species of Menacanthus) parasitic on the Passeriformes has long postpalpal
processes; an approximately rectangular, strongly pigmented gular plate with or
without a central thinner area; two to four setae anterior to the subocular comb
row; three ventral sensilla on the trochanter; and spiniform setae laterally on the
posterior margin of the sternites. Another group (e.g. M. alaudae), also parasitic
on the Passeriformes, has short postpalpal processes; gular plate various; four to
five setae anterior to the comb row, two of which are long; four ventral sensilla on the
trochanter; and no spiniform setae on the sternites. This group, in addition to
being found on the Passeriformes, is found on the Picidae (the species may have

KEY TO GENERA OF MENOPONIDAE 19

fewer setae anterior to the comb row and rather longer head processes) and on the
Galliformes (length of processes varies). Amongst other species parasitic on the
Passeriformes are those showing characters intermediate between the two groups:
a species from one of the Parulidae has the characters of the first group but the post-
palpal processes are small ; Menacanthus crateropus has shorter processes than in the
first group, no sternal spiniform setae, three setae anterior to the comb row and three
sensilla on the trochanter; the species on Salpinctes (Troglodytidae) has small
processes, four to five setae anterior to the comb row; no sternal spiniform setae
and three sensilla on the trochanter. Elsewhere (Hopkins & Clay, 1955 : 180) the
possibility has been discussed that some of the species of Menacanthus from the
Galliformes are nearer to species of Amyrsidea than to other species included in
Menacanthus, being separable only by the presence of the postpalpal process.

It is possible that Menopon, Clayia and Somaphantus also belong to the Mena-
canthus-complex. It has already been suggested that the species of Somaphantus
might be derived from more than one Galliformes-infesting stock, the similarity being
due to the environment of the quill inside which they live. This would explain
some of the differences between the species such as the position of the post-spiracular
setae and chaetotaxy of the head ; spencei Emerson is the only species with a circular
structure within the genital chamber. S. kingi Emerson and Price (no specimens
seen) resembles other species of Somaphantus in the tubular abdomen, but differs
in the number of the sternal brushes and does not have the typical Somaphantus
head; its affinities lie perhaps with such species as Amyrsidea elbeli Emerson and
Stojanovich; the female of this latter species resembles Menopon gallinae in the
prolongation of the last segment and the form of the anus. S. kingi is not included
in Somaphantus in the key.

These genera of Menoponidae found on the Galliformes are possibly derivatives
from a single ancestral stock perhaps parasitic on an early Galliformes stock; the
evolution of the parasites may have included not only divergence with the divergence
of their hosts but also perhaps secondary infestations from host to host at the specific
and supra-specific level.

Notes

Illustrations. In preparation of this paper use has been made of the Stereoscan
scanning electron microscope (S.E.M.) and this has made possible the elucidation
of certain structures. Although most of these can be seen with the light microscope,
especially when once identified and elucidated with the scanning electron microscope,
photographs taken with this latter instrument are the most accurate method of
representing small structures.

Authors and Bibliographical References. In order to save space authors of genera
and species appearing in Hopkins & Clay, 1952, 1953, 1955 are not cited and only
those references which do not appear in Keler, 1960 are listed. Published works
consulted but not always acknowledged in detail are the excellent series of papers on
the genera of the Colpocephalum-complex by Price et alia.

Supplementary Characters. These are separately paragraphed in the key; they
are further attributes characteristic of the genus or group of genera and may help to

ENT. 24, I. 2§

THERESA CLAY

confirm the correct placing of the specimen being identified. New species anomalous
in respect of the key characters may still be placed in the correct genus ; the key is an
artificial arrangement and should be made to fit the genus not the genus to fit the
key.

28 !

29

FIGS. 23-29. 23-24, Prosternal plates. 23, Machaerilaemus. 24, Holomenopon. 25,
Eureum, prosternal and part of gular plate. 26, Plegadiphilus , abdominal pleurites
III-IV. 27, Turacoeca, ventral margin of $ anus. 28-29, Microtrichial patch in $
genital chamber. 28, Cuculiphilus (Falcophilus). 29, Cuculiphilus (Cuculiphilus) .

Actornithophilus and Longimenopon. These are placed together in the key,
couplet 49, as it seems probable that the species of Longimenopon will prove to be
generically inseparable from those included in Actornithophilus (see Timmermann,
1965 : 179).

KEY TO GENERA OF THE MENOPONIDAE

1 Alveoli of marginal temporal setae 26 and 27 closely associated (Text-figs. 2-3) 2
Alveoli of marginal temporal setae 26 and 27 not closely associated (Text-fig, i) 30

2 (i) Head seta 26 similar to 27, long and proximally stout.

Mandibles each with 3-4 teeth, some posteriorly tuberculate; deep narrow
preocular slit; prosternum with > 2 central setae; <J with tibial spur; 3rd
femur with or without ctenidia; one or more sternites with ctenidia; large

species, length > 3-8 mm PIAGETIELLA

Head seta 2 6 significantly shorter and finer than 27 (Text-fig. 2) ... 3

3 (2) Venter of 3rd femur with ctenidia (Text-fig. 14)

Setae 24—25 short to medium ........ 4

Venter of 3rd femur without ctenidia ........ 8

4 (3) Sternites without ctenidia; no preocular notch or slit.

Each side of $ tergite VI comb of elongate setae with clubbed ends held
in pocket in tergite VIII . . . , . . NEW GENUS (in press)

One or more sternites with ctenidia ; preocular notch or slit .... 5

5 (4) 2 postpalpal processes each side (Text-fig. 15)

Narrow preocular slit; prosternum with > 2 central setae; 4 anterior
mesonotal setae; sternites III-IV with single full ctenidium each side

ODORIPHILA

KEY TO GENERA OF MENOPONIDAE 21

Without 2 postpalpal processes each side ....... 6

6 (5) 2 anterior mesonotal setae; ventral margin of $ anus with 2 processes each end,

each bearing long spine-like setae (Text-fig. 27).

Occipital setae (21-22) minute, seta 23 long and lateral to 22 ; broad shallow
preocular notch; single ctenidium each side of sternites III-IV or III-V

TURACOECA
4 anterior mesonotal setae; $ anus without ventrolateral processes.

Euplantula with vertical striations only ...... 7

7 (6) Typical oblong strongly pigmented postnotum absent.

Seta 20 medium length, longer than 19; > 2 prosternal setae; pronotum with
scattered minute setae; sternite III-IV with single full ctenidium each side

DICTEISTA

Typical oblong strongly pigmented postnotum present COLPOCEPHALUM-

complex p. 17

8 (3) 2 anterior mesonotal setae ......... 9

4 anterior mesonotal setae ......... 1 1

9 (8) Prosternal plate well-developed, pointed posteriorly (Text-fig. 23) prosternum

with > 2 central setae.

Antennal fossa deep, head broad at temples and preocular expansions;
3rd femur without brush; trochanter with not more than 3 ventral sensilla

MACHAERILAEMUS

Prosternal plate not well developed and not pointed posteriorly; prosternum
with 2 central setae

Seta 23 missing or anterior to 22 . . . . . . . 10

10 (9) Proximal antennal sensillum large, in pit and widely separated from distal

(PI. 2, figs. 10-11) ; claw without large basal process; bilobed process each end
of $ ventral anal margin with long setae, some of which may be stout and
spiniform ; temples expanded ; euplantula banded ; trochanter with more than
3 ventral sensilla.

Seta 23 missing or minute; 3rd femur with or without brushes

BUCEROPHAGUS

Antennal sensilla small, adjacent and terminal; claw with large basal process
(Text-fig. 12); $ anus without ventral processes; temples scarcely expanded ;
euplantula not banded; trochanter with 3 ventral sensilla.

3rd femur without brushes ; seta 24-25 short . . . BONOMIELLA

11 (8) Tibiae I-III with outer submarginal comb of short setae (as in PI. 6, fig. 34);

brushes on sternite IV-V towards centre of segments.

Preocular slit; terminal antennal segment elongate, cylindrical; 3rd femur
with thick brushes; euplantula with vertical striations only; setae 24-25

short HOAZINEUS

Tibiae I-III without such comb of setae; brushes or ctenidia on sternites IV-V

absent or lateral on segments . . . . . . . . 12

12 (n) Gular plate large and tripartite (Text-fig. 19); 2nd tarsal segment of 2nd and

3rd legs with combs of processes (euplantula II, PI. 6, fig. 35).

Seta 24 or 25 long; preocular slit; postpalpal processes present or absent;
some abdominal pleurites with posteroventral corners prolonged

PSEUDOMENOPON
Gular plate and tarsi not as above . . . . . . . .13

13 (12) Gular plate with 4 seta-bearing processes (Text-fig. 18).

Seta 24 or 25 long; head considerably broader than long; antennal fossa deep
and pouch-like; prosternal plate with posterior process and > 2 setae;
tergites I & II with short seta laterad to post-spiracular seta COLIMENOPON
— Gular plate without 4 seta-bearing processes . . . . . . 14

22 THERESA CLAY

X4 (T3) One spinous process near base of antenna and one near base of maxillary palp;
thickening running inwards from anterior end of subocular comb row bearing
at least one seta.

Close set row of subocular setae; head sensilla c. not apparent; tergite I
with short seta laterad to post-spiracular seta; euplantula banded; seta

24-25 short (KELERIMENOPON) 15

Head without such spinous processes and without setae-bearing thickening

running inwards from comb row . . . . . . . . 16

X5 (J4) Abdominal pleurites without prolongation of ventro-posterior corner; broad

deep preocular notch .... sg. LORIMENOPON Price & Emerson

Some abdominal pleurites with prolongation of ventro-posterior corner; narrow

preocular slit sg. KELERIMENOPON

16 (14) Labrum with striated lobe (PI. 4 fig. 23); hypopharynx characteristic (PI. 4,

fig. 22).

Head broad, dorsal preocular margin overlapping ocular margin (Text-fig.
16); sensilla c. not apparent; setae 24-25 short . . NEOMENOPON

Labrum without striated lobe; hypopharynx not as in PL 4, fig. 22 . . 17

17 (16) Dorsum of head with scattered minute alveoli

$ with 1-2 circular or oval structures associated with genital chamber;
setae 24-25 short ........... 18

Dorsum of head without scattered minute alveoli ...... 20

1 8 (17) Distal anterior angle of pedicel not markedly prolonged; posterior part of coxa

I with > 6 setae; single structure in $ genital chamber; <$ genitalia asym-
metrical.

Head semilunar .......... 19

Distal anterior angle of pedicel markedly prolonged; posterior part of coxa I
with < 6 setae; 2 oval structures in $ genital chamber; <$ genitalia symmet-
rical MIMEMAMENOPON

19 (18) Sternite I divided vertically, partially or entirely; gular plate sculptured;

prosternum with only 2 central setae .... EOMENOPON

Sternite I not divided ; gular plate not sculptured ; prosternum with > 2 central

setae PACIFIMENOPON

20 (17) Venter of 3rd femur without brushes; temples with ventral submarginal patch

or row of setae; setae 24-25 short; postvulval sclerite with setae; head broad

at temples, semilunar ........ NOSOPON

Without above combination of characters . . . . . . .21

21 (20) One postpalpal process each side; seta 24-25 short

Sensilla c. not apparent; prosternal plate with well-developed posterior
process or with 3 irregular small processes . . . . . . 22

Either without postpalpal processes or if present then seta 24 or 25 long . . 23

22 (21) Head without preocular slit or notch; pleurites without prolongation of postero-

ventral corners; tergite I without short seta laterad to post-spiracular seta;
well-developed circular structure of cellular appearance associated with $
genital chamber ......... APTERYGON

Head with preocular slit; some pleurites with posteroventral prolongations;
tergite I with short seta laterad to post-spiracular seta; either no or different
type of structure associated with $ genital chamber . . HOHORSTIELLA

23 (21) 3 or more preocular setae (10-11) on at least one side; tibiae II-III with patch

or row of outer dorsal submarginal setae; dorsal preocular margin overlaps
ocular margin (Text-fig. 17).

Head sensilla c. not apparent; euplantula banded; seta 24-25 short

ME R OMENOPON
Without above combination of characters . . . . . . . 24

KEY TO GENERA OF MENOPONIDAE 23

24 (23) Postnotum not apparent; head seta 16-19 all long and stout, reaching at least

to the transverse pronotal carina.

Seta 24 long and stout; no preocular notch or slit . . NUMIDICOLA

Postnotum present; head seta 16-19 not as above. • • • • • 25

25 (24) Brushes on sternite IV only.

No setae between 23 and 27 reaching beyond transverse pronotal carina . 26
Brushes absent or not on sternite IV only . . . . . . . 27

26 (25) The 2 subterminal setae of maxillary palp with definite gap between their

alveoli ; shape of head characteristic (PI. 7, fig. 44) ; abdomen narrow and

tubular SOMAPHANTUS

The 2 subterminal setae of maxillary palp with their alveoli contiguous; shape

and head of abdomen otherwise ...... MENOPON

27 (25) Inner central pronotal setae absent or on posterior part of pronotum; no seta

between 23 or 27 reaching beyond transverse pronotal carina.

Broad or shallow preocular notch ....... CLA YIA

Inner central pronotal setae on or near transverse carina; one seta between 23

and 27 reaching below transverse pronotal carina . . . . . 28

28 (27) Terminal antennal segment without signs of division; ridges numerous and

close together (PI. 3, fig. 13) . . . MENACANTHUS- complex, p. 18
Terminal antennal segment with form of ridges and pigmentation suggesting
one or more divisions, or if no signs of division, ridges are few and widely
separate (PI. 3, fig. 14) .......... 29

29 (28) More than 2 central prosternal setae; pleurites without posterior processes;

tergites with transverse bar . . . . . EUCOLPOCEPHALUM
Only 2 central prosternal setae ; some abdominal pleurites with posterior proces-
ses; no transverse tergal bars ..... PLEGADIPHILUS

30 (i) Thorax with 2 large sternal plates bearing many setae (PI. 5, fig. 25); scape

(and pedicel) with distal anterior prolongations (Text-fig. 6).

Seta ii on protuberance; large species, length > 4-00 mm . TRINOTON
Thorax without such sternal plates; scape without distal anterior prolongation 31

31 (30) 2 anterior mesonotal setae.

Seta 23 absent or anterior to 22 ; at least one of the setae 24, 25 or 26 long;
sensilla c. not apparent; euplantula banded ... 32

4 anterior mesonotal setae (sometimes widely spaced) ..... 36

32 (31) One pair of mid-dorsal head setae; not more than one pair of central pronotal

setae ; no preocular slit or notch ; ? ventral anal margin without lateral setae-
bearing processes ....... • 33

Two pairs of mid-dorsal head setae ; 2 pairs of central pronotal setae ; preocular

slit or notch; $ ventral anal margin with lateral setae-bearing processes . 35

33 (32) Prosternum with 2 central setae; dorsal margin of head without ventral trun-

cated ovoid excavation; seta 23 absent; no central pronotal setae.

Mesothorax with sternum, pleura and tergum fused to form strongly
pigmented ring ....... • MYRSIDEA

Prosternum with > 2 central setae; ventral truncated-ovoid excavation in
dorsal margin of head with thickened anterior rim; seta 23 present; one pair
of central pronotal setae .... ... 34

34 (33) Gular plate horseshoe-shaped (Text-fig. 25); temporal carina not developed

EUREUM

Gular plate not horseshoe-shaped; temporal carina well developed DENNYUS

35 (32) Venter of 3rd femur and sternite IV with ctenidia; terminal antennal segment

with signs of division ; $ anal processes with stout spiniform setae

BUCEROCOLPOCEPHALUM

Venter of 3rd femur and sternite IV without ctenidia; terminal antennal seg-
ment without signs of division ; $ anal processes with long setae . CHAPINIA

24 THERESA CLAY

36 (31) Venter of 3rd femur and lateral areas of some sternites with comb-like out-

growths (PI. 6, fig. 32); distal tarsus swollen; claws delicate with narrow
elongate points (PL 6, fig. 36).

Euplantula with vertical striation only; postnotum not vertically oblong;

preocular notch MICROCTENIA

Venter of 3rd femur and lateral areas of sternites without comb-like out growths ;

tarsi and claws not as above . . . . . . . . . 37

37 (36) Centre of mesosternum with 2 setae.

Tergite I with short seta laterad to post-spiracular setae, may be lacking in
Austromenopon becki . . . . . . . '.'• . . 38

Centre of mesosternum with > 2 setae.

Seta 23 missing or anterior to 22; sternal brushes absent or only on, or
thickest on IV V. . 47

38 (37) Mesothorax as a sclerotized ring formed by fused sternum, pleura and tergum;

temples with ventral submarginal patch or rows of setae.

Seta 1 8 widely posterolateral to 17; group of 3 subterminal setae on maxil-
lary palp (PI. 7, fig. 45); terminal segment of antenna subdivided (PI. i,
fig. 3); head broad with narrow preocular slit . . . . . . 39

Mesothorax not fused as above; temples without ventral submarginal patch or

rows of setae ........... 43

39 (38) Venter of 3rd femur without ctenidia; tergites I-VII with short seta laterad to

post-spiracular seta ...... NEW GENUS (in press)

Venter of 3rd femur with ctenidia; tergites III-VII without short seta laterad
to post-spiracular seta.

Euplantula with vertical striations only . . (CUCULIPHILUS) 40

40 (39) Postpalpal processes present ...... sg. CARRIKERIA

Postpalpal processes absent . . . . . •'..'• • • 41

41 (40) Abdominal sternite I not apparent sg. AEGYPIPHILUS

Abdominal sternite I apparent ......... 42

42 (41) $ genital chamber with vertically elongate patch of micro trichia and with

sclerite each side with thickened inner margin (Text-fig. 28) ; $ with stout

spiniform setae on tergite VII sg. FALCOPHILUS

$ genital chamber with patch of microtrichia not vertically elongate and with-
out such sclerite each side (Text-fig. 29) ; $ without spiniform setae on tergite
VII "... ' . . sg. CUCULIPHILUS

43 (38) Narrow preocular slit.

Euplantula banded .......... 44

Without preocular slit.

Seta 23 anterior to 22 ......... 45

44 (43) Eyes in normal position on dorsolateral margin; dorsolateral and ventrolateral

margins normal (PI. 4, fig. 19) ; seta 23 in line with 22 and 21 ; tibiae II and III
with 1-2 irregular submarginal rows of dorsal setae; tergite II without short
seta laterad to post-spiracular seta. . . NEW GENUS Ryan & Price

in press

Eyes not marginal (PI. i, fig. 5); cavity between dorsolateral and ventrolateral
margins roofed over distally (PL 4, fig. 20) ; seta 23 anterior to 22 ; tibiae II
and III without extra rows of dorsal setae; tergite II with short seta laterad to
post-spiracular seta EIDMANNIELLA

45 (43) Large triangular or rectangular postnotum; dorsolateral margin of head with

row of short setae.

No typical subocular comb row; only one of labral setae long each side; 2
short and 3 long setae between setae 28 and 23 ; venter of 3rd femur without
brushes . ANCISTRONA

KEY TO GENERA OF MENOPONIDAE 25

Normal vertically oblong postnotum ; dorsolateral margin of head without row

of short setae ........... 46

46 (45) Prosternal plate with deeply serrated margin (Text-fig. 24) ; tergite II with short

seta laterad to post-spiracular seta . . . . HOLOMENOPON

Prosternal plate without deeply serrated margin; tergite II without short seta

laterad to post-spiracular seta .... AUSTROMENOPON

47 (37) Tibiae I-III with dorsal row or rows of short stout submarginal setae, signifi-

cantly shorter than outer ventral setae.

At least one long stout seta between 27 and 23 . . . . 48

Tibiae I-III without such setae.

Anterior mesonotal setae separated ....... 49

48 (47) Outer dorsal tibial setae in comb-like row (PI. 6, fig. 34) ; head narrowed

anteriorly; preocular notch backed by heavily pigmented nodus (PI. 4, fig. 24)

HELEONOMUS

— Outer dorsal tibial setae in irregular row or rows; head broadly rounded anter-

iorly; preocular concavity without heavily pigmented nodus (Text-fig. 22)

GRUIMENOPON

49 (47) Without typical pronotal transverse carina or typical oblong postnotum.

Head narrow, elongate, sides approximately parallel without notch or slit
(PI. 7 fig. 43); terminal antennal segment with signs of division; vertical
carina of pronotum well-developed ; 3rd femur without brushes . REDIELLA

Transverse pronotal carina and postnotum typical.

Euplantula banded . ACTORNITHOPHILUS & LONGIMENOPON

ACKNOWLEDGEMENTS

I am greatly indebted to J. H. Calaby, R. E. Elbel, K. C. Emerson, J. A. Ledger,
G. A. Lincoln, R. L. C. Pilgrim and R. D. Price for supplying material; to Professor
R. D. Price for reading the manuscript and making many helpful comments; and
to the staff of the British Museum (N.H.) Electron Microscope Unit.

REFERENCES
BALTER, R. S. 1968. Lice egg morphology as a guide to taxonomy. Med. biol. 18 : 94-95.

— 1968. The microtopography of avian lice eggs. Ibid : 166-179.
BLAGOVESHTCHENSKY, D. I. 1964. Keys to the European part of U.S.S.R. Order Mallo-

phaga, in Bei-Bienko, G. Ya [Ed.]. Opred. Faune SSSR I : 309-323. (Seen in translation,

I.P.S.T., 1967).
CLAY, T. 1961. A new genus and species of Menoponidae (Mallophaga) from Apteryx. Ann.

Mag. nat. Hist. (13) (1960), 3 : 571-576.
— 196212. A key to the species of Actornithophilus. Bull. Br. Mus. nat. Hist. (Ent.) 11 :

189-244.
19626. A new species of Anatoecus from Phoenicopterus ruber. Ent. Ber., Amst. 22 : 220-

226.
1966. Contributions towards a revision of Myrsidea. I. Bull. Br. Mus. nat. Hist.

(Ent.) 17 : 327-395-

— 1968. Contributions towards a revision of Myrsidea. III. Bull. Br. Mus. nat. Hist.
(Ent.) 21 : 203-243.

CLAY, T. & MOREBY, C. 1967. Entomology of Antarctica. Mallophaga and Anoplura. II.

Antarctic Research Series 10 : 157-196. Honolulu.
HAUB, F. 1967. Der Kopf von Pseudomenopon pilosum (Mallophaga- Amblycera). Zool.

Jb.Anat.84 : 493-558-
HOPKINS, G. H. E. & CLAY, T. 1952. A check list of the genera and species of Mallophaga.

British Museum (Nat. Hist), London.

26 THERESA CLAY

KELER, S. VON. 1960. Bibliographic der Mallophagen. Mitt. zool. Mus. Berl. 36 : 146-403.

PRICE, R. D. 1966. The genus Eomenopon Harrison with descriptions of seven new species
(Mallophaga: Menoponidae) . Pacif. Insects 8 : 17-28.

1968. A review of the genus Dicteisia Bedford (Mallophaga : Menoponidae) with des-
cription of a new species. /. med. Ent. Honolulu 5 : 445-452.

PRICE, R. D. & BEER, J. R. 1963. The Kurodaia (Mallophaga : Menoponidae) parasitic on
the Strigiformes, with a key to the species of the genus. Ann. ent. Soc. Am. 56 : 849-857

1965. The Colpocephalum (Mallophaga : Menoponidae) of the Ciconiiformes. Ibid.

58 : 111-131.

SCHARF, W. C. & PRICE, R. D. 1965. A taxonomic study of the genus Cuculiphilus (Mallo-
phaga : Menoponidae). Ann. ent. Soc. Am. 58 : 546-555.

TENDEIRO, J. 1967. Mallophages due Pare National de 1'Upemba (Congo) (Mission G. F.
de Witte). Rev. Est. ger. univ. Mozambique 4 : 361-441.

TIMMERMANN, G. 1965. Die Federlingsfauna der Sturm vogel und die Phylogenese des
procellariiformes Vogelstammes. Abh. Verh. naturw. Ver. Hamburg (8 suppl.) : 1-249.

TUFF, D. W. 1967. Notes on the Mallophagan genus Comatomenopon and descriptions of
two new species. /. Med. Ent. Honolulu 4 : 247-250.

THERESA CLAY D.Sc.

Department of Entomology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON, S.W.7

PLATE i

FIG. i. Bucerophagus productus. Antenna.

FIG. 2. Bucerophagus productus. Base of 3rd antennal segment.

FIG. 3. Cuculiphilus snodgrassi. 4th and 5th antennal segments.

FIG. 4. Eidmanniella sp. n. (in press). 3rd and 4th antennal segments.

FIG. 5. Eidmanniella sp. n. (in press). Dorsum of head.

FIG. 6. Plegadiphilus sp. Dorsum of head. o. ommatidia; d. dorsolateral margin of head-

(S.E.M.)

Bull. Br. Mus. nat. Hist. (Ent.) 24, i

PLATE i

ENT. 24, I.

PLATE 2

FIGS. 7-12.
same seta in figs. 10-12
FIG. 7. B. productus.
FIG. 8. B. productus.
FIG. 9. B. productus.
FIG. 10. B. africanus.
FIG. ii. B. africanus
FIG. 12. B. africanus

Bucerophagus antenna (S.E.M.). x. marks the same seta in figs. 7-9; v. the

Terminal antennal segment.

End-on view of terminal antennal segment.

As fig. 2, enlarged.

Antenna.

Part of terminal segment showing proximal sensillum.

Part of terminal antennal segment. (S.E.M.)

Bull. Br. Mus. nat. Hist. (Ent.) 24, i

PLATE 2

PLATE 3

FIG. 13. Menacanthus stramineus. Terminal antennal segment.

FIG. 14. Plegadiphilus sp. from Geronticus eremita. Terminal antennal segment.

FIGS. 15-16. Meromenopon sp. To show variation (not specific) in subocular seta (s).

FIG. 17. Dennyus sp. Details of metasternal flap.

FIG. 1 8. Trinoton emersoni Clay. Part of white gular area with 2 gular setae. (S.E.M.

Bull. Br. Mus. nat. Hist. (Ent.) 24, i

PLATE 3

PLATE 4

FIG. 19. ' Eidmanniella' aurifasciata. Antennal fossa, s. subocular seta; c. subocular comb
row. (S.E.M.)

FIG. 20. Eidmanniella sp. Antennal fossa. (S.E.M.)

FIG. 21. Colpocephlaum sp. c. subocular comb row; /. submarginal temporal setae. (S.E.M.)

FIG. 22. Neomenopon sp. Hypopharynx. (Light microscope. T.C.)

FIG. 23. Neomenopon sp. Labrum. (Light microscope. T.C.)

FIG. 24. Heleonomus sp. Head. (Light microscope. T.C.)

Bull. Br. Mus. nat. Hist. (Ent.) 24, i

PLATE 4

24

PLATE 5

FIG. 25. Trinoton querquedulae . g. white gular area; m. metasternal flap. (Direct illum-
ination. T.C.).

FIG. 26. Euveuvn cimicoides. Venter of thorax and part of abdomen, m. metasternal flap
(S.E.M.)

FIG. 27. Eureum cimicoides. Metasternal flap. (S.E.M.)

FIG. 28. Trinoton sp. Part of metasternal flap. (S.E.M.)

FIG. 29. Eureum cimicoides. Details of metasternal flap (S.E.M.)

FIG. 30. Trinoton sp. Details of metasternal flap (S.E.M.)

Bull. Br. Mus. nat. Hist. (Ent.) 24, i

PLATE 5

y

/^ (m

PLATE 6

FIG. 31. Colpocephalum sp. Femoral ctenidia. (S.E.M.)

FIG. 32. Microctenia sp. Femoral comb-like processes. (S.E.M.)

FIG. 33. Franciscoloa roseicapillae Price & Beer. 3rd tibia (Phase contrast T.C.)

FIG. 34. Heleonomus sp. Row of setae on outer tibial margin. (S.E.M.)

FIG. 35. Pseudomenopon pilosum. Euplantula II. e. base of eupodium; d. piece of dirt.

(S.E.M.)

FIG. 36. Microctenia sp. Tarsus. (Phase contrast. T.C.)

Bull. Br. Mus. nat. Hist. (Ent.) 24, i

PLATE 6

35

PLATE 7

FIG. 37. Amyrsidea sp. Euplantula I. (S.E.M.)

FIG. 38. Actornithophilus sp. Euplantula I. ? without surface membrane. (S.E.M.)

FIG. 39. Part of fig. 2, enlarged (S.E.M.).

FIG. 40. " Eidmanniella " aurifasciata. Euplantula I, to show horizontal banding. (Phase
contrast. T.C.)

FIG. 41. Falcomenopon sp. Euplantula I, to show vertical striations. (Phase contrast.
(T.C.)

FIG. 42. Plegadiphilus sp. Proximal sensillum. (S.E.M.)

FIG. 43. Rediella mirabilis. Head. (T.C.)

FIG. 44. Somaphantus spencei. Head. (T.C.)

FIG. 45. Cuculiphilus snodgrassi. Subterminal sensory setae of maxillary palp. (S.E.M.)

Bull. Br. Mus. nat. Hist. (Ent.) 24, i

PLATE 7

A LIST OF SUPPLEMENTS
TO THE ENTOMOLOGICAL SERIES

OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

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

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

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

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

1965- £3 5s.

5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156;
475 Text-figures. November, 1965. £2 15$.

6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae & Drosophilidae.
Pp. 129; 328 Text-figures. May, 1666. £3.

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

£33s.

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

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

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rhopal-
ocera). Pp. 322; 233 Text-figures. Coloured frontispiece. September, 1967. £8.

11. MOUND, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 184:
82 Text-figures. May, 1968. £4.

12 WATSON, A. The Taxonomy of the Drepaninae represented in China, with an

Account of their World Distribution (Lepidoptera : Drepanidae). Pp. 151:

293 Text-figures, 14 plates. November, 1968. £5.
13. AFIFI, S. A. Morphology and Taxonomy of the Adult Males of the families

Pseudococcidae and Eriococcidae (Homoptera : Coccoidea). Pp. 210: 52

Text-figures. December, 1968. £5.

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

C: .

THE TYPE-MATERIAL OF

TACHINIDAE (DIPTERA)

DESCRIBED BY N. BARANOV

C. W. SABROSKY

&
R. W. CROSSKEY

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 24 No. 2

LONDON: 1969

THE TYPE-MATERIAL OF TACHINIDAE
(DIPTERA) DESCRIBED BY N. BARANOV

BY

CURTIS WILLIAMS SABROSKY

Systematic Entomology Laboratory, U.S. Department of Agriculture

&

ROGER WARD CROSSKEY

Commonwealth Institute of Entomology
Pp. 27-63

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 24 No 2

LONDON: 1969

THE BULLETIN OF THE BRITISH MUSEUM

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

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

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

This paper is Vol. 24, No. 2, of the Entomological
series. The abbreviated titles of periodicals cited follow
those of the World List of Scientific Periodicals.

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

Trustees of the British Museum (Natural History) 1969

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 3 December, 1969 Price £i is.

(£1-05)

THE TYPE-MATERIAL OF

TACHINIDAE (DIPTERA)

DESCRIBED BY N. BARANOV

BY

By CURTIS W. SABROSKY & R. W. CROSSKEY

CONTENTS

Page

SYNOPSIS ........... 29

INTRODUCTION .......... 29

ACKNOWLEDGEMENTS ......... 33

PART I. Baranov's available species-group names and their types . 35

PART II. Baranov's manuscript names that have been published . . 54

REFERENCES ........... 59

INDEX TO SPECIES-GROUP NAMES ....... 61

SYNOPSIS

An alphabetical catalogue is given of the 156 species and infraspecific taxa of Tachinidae
described by N. Baranov, with an account of all located type-material on which the names
are based. Sixty-one lectotypes are newly designated. Manuscript names of Baranov that
have appeared in print, some of which have been validated by later authors, are enumerated
and briefly discussed. The references given include a complete bibliography of Baranov's
papers on Tachinidae.

INTRODUCTION

BETWEEN 1926 and 1942 the Slav dipterist N. Baranov — who was born in Russia
but worked mainly in Jugoslavia — published a series of papers on the taxonomy of
Calyptrate Diptera, many of them in rather inaccessible Yugoslav journals. In
these he described and named a total of 26 genera, one subgenus, 183 species, and
17 infraspecific taxa, distributed as follows : 19 genera, one subgenus, and 145
species (plus n infraspecific taxa) of Tachinidae ; 6 genera and 32 species (plus 6
infraspecific taxa) of Sarcophagidae ; one genus and 5 species of Calliphoridae ;
and one species of the Muscidae. Our present paper is concerned only with the
Tachinidae, the vast majority of which were described from the Oriental and
Australasian Regions ; Baranov described only a few taxa from the Palaearctic
Region, and none from Africa or the New World. For each species-group taxon
we give an account of all the type-material that we have been able to locate, but we
have not attempted to assign the species to currently recognized genera or to investi-
gate possible synonymy (this will be done by Crosskey at a later stage for the
systematic catalogues of Oriental and Australasian Tachinidae in preparation).
The paper is presented in two parts : Part I contains the properly proposed and

30 C. W. SABROSKY & R. W. CROSSKEY

available species-group names of Baranov, and Part II contains the 32 manuscript
names of Baranov that have appeared in print ; of the latter a few were cited in
synonymy by Baranov himself, but most were published by someone else.
Occasionally the other authors have given some descriptive matter and have thus
made some of the Baranov manuscript names nomenclaturally available under the
International Code of Zoological Nomenclature, 1961, so that authorship must be
credited to them (under Article 10) and not to Baranov. However, as the names
appear in the literature credited to Baranov it is desirable to account for them in
the present work. In addition to the names given in Part II we have discovered,
during preparation of this paper, a number of specimens in different museum
collections that are sometimes labelled as types and bear manuscript names of
Baranov that have never (so far as we can trace) appeared in print ; at least 23
such names are known to us, but as they are unpublished manuscript names only
we are not recording them.

All names given in Parts I and II are listed alphabetically in their original com-
binations. For each nominal species-group taxon in Part I the entry is arranged to
show the following information in the sequence indicated : —

Name ; author ; date and page reference of original publication ; status
and sex of primary type ; authority for lectotype designation (if relevant) ;
data of primary type (when available in the sequence : locality, altitude, date
of collection, host information, name of collector) ; type-depository ; location
of genitalia in the case of male primary types (as ' genitalia in situ ' or ' geni-
talia on slide ').

Number and sex of paralectotypes or paratypes, with data and depository
information as for primary types. (It has not been considered necessary,
however, to specify whether male genitalia are present on or removed from
paralectotypes or paratypes.)

Explanatory comments or annotations if considered necessary.
The type data as we record them do not necessarily conform exactly in spelling
or sequence with the data labels. Geographical names are given as on the data
labels (e.g. Buitenzorg and not the modern equivalent of Bogor), except for the
correction of obvious misspelling and a modern terminology for sovereign states
(e.g. Thailand instead of Siam). The major islands of the former Dutch East
Indies are shown by their well known names such as Celebes, and not by their very
new names. The names of plants and insects cited in the host information are
given as on the data labels and have not been checked for their modern equivalents.
Collectors' names are given with full initials when these are known, even if not all
are shown on the data labels (e.g. R. J. A. W. Lever instead of R. A. Lever).
Baranov spelt his own name with either a terminal ' v ' or ' ff ' in his papers on
Tachinidae, but almost always used the ' ff ' ending on his type labels and deter-
mination labels ; we have not differentiated in this paper but have adopted the
' v ' ending throughout, except when quoting his labels.

Much of Baranov's type-material from H. Sauter's collecting in Formosa is
labelled either ' Kankau ' or ' Kankau (Koshun) ' ; we have uniformly used both
names as the village of Kankau (now Koko) is in the district of Koshun, with the

TYPE-MATERIAL OF TACHINIDAE 31

co-ordinates 22° oo' N. and 120° 49' E. (locality traced from Gazetteer (No. 13)
Formosa (Taiwan), U.S. Navy Department, Hydrographic Office Publication No.
393. 139 PP-. 1944)-

Recognition of type-material and interpretation of its status are often difficult
with Baranov species, especially in the face of variant usage in both practice and
publication. Certain conclusions need some discussion.

Baranov consistently placed an identification label in his handwriting on each
specimen (we interpret the very few exceptions found as due to loss of labels),
usually with ' n.sp. N. Baranoff ' after the specific name. However, sometimes
some of such material was not mentioned in the original publication. We have
felt obliged to exclude from the type-series all material that differs from that speci-
fied in the published data except where there seems to be a reasonable explanation
for a discrepancy, such as a typographical error or a misread label. All such
instances are annotated.

Baranov did not use the term ' holotype ' in publication or on labels, although his
' Typus ' is sometimes that. Usually ' Typus ' was not mentioned in publication,
however. Some series have a ' Typus <$ ' and a ' Typus $ ' and ' Cotypen ', but
some have only ' Cotypen ' so far as we can discover. Occasional examples are
labelled ' Paratypisches Exemplar '. We have regarded all such series as consisting
of syntypes and have designated lectotypes when not already designated ; usually,
when available, we have selected Baranov's ' Typus <$ ' as the lectotype. Each
lectotype has been clearly labelled as such, and 61 lectotypes are newly designated.

Hennig (1941) mentioned many of Baranov's specimens in his list of the Diptera
of Formosa, but he was only a recorder of data on specimens in the Deutsches
Entomologisches Institut collection and not a reviser or designator of lectotypes.
His use of ' Typus ' merely indicates the presence of one specimen, as opposed to
' Typen ' for more than one ; this is clear from his multiple use of ' Typus ' under
those species where there were single specimens from two or more localities.
Townsend (1934-1942, Manual of Myiology, 12 Parts, Itaquaquecetuba) has some-
times cited ' Ht [i.e. holotype] in DEI ', but he did not see Baranov material or
label specimens, and apparently he listed ' holotypes ' by assumption from the
literature. If there is only one male in the series, in such a case, we have accepted,
albeit reluctantly, Townsend's action as fixation of the lectotype. However,
when two or more syntypes of the designated sex are available Townsend's published
designation is not an ultimate restriction to a single specimen and we have then
designated a lectotype.

Designation of a lectotype automatically converts all other syntypes into para-
lectotypes, even if they are never labelled as such and even if not conspecific with
the lectotype. In Baranov's work mixed type-series rarely occur and we have
recorded the few cases known to us. We have labelled all available syntypes
remaining after lectotype designation as paralectotypes, even for mixed series. We
should, however, comment here that there may be other paralectotypes that we
have not seen, and indeed we consider this probable for the following reason :
many of the species described by Baranov from an unstated number of specimens

32 C. W. SABROSKY & R. W. CROSSKEY

were based upon material submitted to him for identification by the Imperial (now
Commonwealth) Institute of Entomology, and in the years from 1932 until 1940 he
received all the material of Oriental Tachinidae coming to this Institute from
British-administered territories ; this often consisted of series of reared specimens
sent by departments of agriculture or forestry in India, Ceylon, Burma, Malaya,
Solomon Islands, or Fiji, and undoubtedly some original specimens were returned
(after identification and description) to collections in the territories of origin. It is
therefore likely that some insect collections in the countries mentioned still contain
specimens that are paralectotypes of species described by Baranov.

The status of certain material is sometimes difficult to determine, and we have
decided each case individually. For some species an expression in the original
publication such as ' Weitere Exemplare ' seems to be merely a way of recording
additional type-material and we have accepted these specimens as syntypes. In
other cases it seems clear to us from Baranov's words that the additional specimens
cited did not form part of his type-series, nor was the description based upon them.
All such cases have been annotated. Should later differences of opinion arise our
lectotype designations will nevertheless stand, and any other specimens later
believed to be part of the type-series will be additional paralectotypes.

Some comment is necessary on the slide preparations of male genitalia which
exist in several museum collections and associate with pinned type-specimens.
Baranov was one of the first workers on the Tachinidae to recognize the great
value of the male genitalia for distinguishing between closely allied species, especially
among the Exoristine and Goniine forms in which few other really reliable characters
exist, and he frequently published figures of the genitalia drawn from permanent
slide-mounts. Many of Baranov's male type-specimens have had the whole hypo-
pygium neatly extracted and mounted on labelled glass slides, and it is usually
possible to associate slides of the genitalia with the actual specimens from which
they were made. In the text for each nominal taxon listed in Part I of the present
work we have indicated whether the genitalia of the male primary type are in situ on
the specimen or separately slide-mounted ; we have been able to locate the
associated slides for almost all primary types from which they have been removed,
but there are a very few (indicated where necessary) for which the slide-mount
appears to be lost. In the course of our work it was found that several slides were
still among Baranov's own collection (now in the U.S. National Museum) although
the associated type-specimens were correctly located elsewhere, and in these cases
the slides have now been sent to the museum collections in which the type-specimens
themselves are deposited. The statement ' genitalia on slide ' given in the text for
any holotype or lectotype therefore implies that the slide will be found in the same
collection as the pinned primary type.

Baranov's type-material is scattered among several museum collections. The

bulk of it is in London, Eberswalde (East Germany) and Washington D.C., but

some type-specimens are in Bogor (Indonesia), Ottawa, Warsaw, Amsterdam,

Dresden, and Brisbane. Specimens mentioned by Baranov as being located in the

Awhere Baranov formerly worked ; and we thank Dr. A. Kaltenbach for confirming"! —

TYPE-MATERIAL OF TACHINIDAE 33

museum at Stettin are now in the Zoological Institute of the Polish Academy of
Sciences, Warsaw. In some of his papers Baranov has mentioned specimens in the
Instituut voor Plantenziekten, Buitenzorg (now the Central Institute for Agricultural
Research, Bogor) but from a detailed list of Tachinidae in the collection of that
Institute (very kindly sent to us by Dr. Ida Njoman Oka : see Acknowledgements)
it appears that no actual syntype specimens are present there. There are, however,
several holotype, lectotype and paralectotype specimens in the Museum Zoologicum
Bogoriense. Specimens mentioned by Baranov as belonging to the Imperial
Institute of Entomology are in the collection of the British Museum (Natural
History). To condense the text we have used the following abbreviations for the
main type-depositories :

BMNH British Museum (Natural History), London.

CNC Canadian National Collection, Ottawa.

DEI Deutsches Entomologisches Institut, Eberswalde.

IZPAN Instytut Zoologiczny, Polska Akademia Nauk, Warsaw.

MZ Museum Zoologicum Bogoriense, Bogor, Indonesia.

USNM United States National Museum, Washington, D.C.

Baranov's own collection was acquired by the United States National Museum in
1960, and the depository abbreviation USNM is applied to material that correctly
belonged in the former Baranov collection.

ACKNOWLEDGEMENTS

It gives us much pleasure to acknowledge the helpfulness and generosity of the
colleagues who have assisted us with the loan of type-specimens or with valuable
information in reply to our enquiries.

We are especially indebted to Dr. Gunther Morge for the loan of the very large
number of Baranov types, together with genitalia slides, from the collection of the
Deutsches Entomologisches Institut, and to Dr. S. Somadikarta for the loan of
types from the Museum Zoologicum Bogoriense, Indonesia.

For confirmation of the presence of types in other collections, and for details of
their data, we are most grateful to Dr. A. Draber-Monko (Zoological Institute,
Polish Academy of Sciences, Warsaw), Dr. W. Ellis (Zoological Museum,
Amsterdam), Dr. R. Hertel (Staatliches Museum fur Tierkunde, Dresden), and to
Mr. G. E. Shewell and Dr. D. M. Wood (Entomology Research Institute, Canadian
Department of Agriculture, Ottawa). Dr. L. P. Mesnil kindly furnished us with
information on a few types from the DEI collection that are temporarily in his care
at the Commonwealth Institute of Biological Control, Delemont, Switzerland, and
Dr. J. d'Aguilar kindly confirmed that the missing holotype of Voria edentata was
not on loan to him.

We thank Dr. D. Mikacic, Department of Parasitology and Parasitic Diseases,
Faculty of Veterinary Medicine, University of Zagreb, for confirming that so far as
he can trace there are now no Baranov type-specimens in Zagreb, Yugoslavia,
none can be found in the Naturhistorisches Museum, Vienna (to which it was

34 C. W. SABROSKY & R. W. CROSSKEY

thought that some material could have been moved from Zagreb during the last
war).

Dr. Ida Njoman Oka provided us with a most valuable list of the Tachinidae
identified by Baranov that are in the collection of the Central Agricultural Research
Institute (Lembaga Pusat Penelitian Pertanian) in Bogor, Indonesia, and we take
this opportunity of expressing our particular appreciation of this in view of the large
amount of work that its preparation entailed.

TYPE-MATERIAL OF TACHINIDAE 35

PART I.— BARANOV'S AVAILABLE SPECIES-GROUP NAMES AND THEIR TYPES

In the following list, LECTOTYPE indicates by present designation.

Actia pulex Baranov, 19386 : 410. LECTOTYPE^, SOLOMON ISLANDS : Tulagi, 19.1^.1934,
on Cocos flower (R. J. A. W. Lever) (BMNH). Genitalia in situ.

Paralectotypes : i (J, i $, same data as lectotype (BMNH). i $, same data as lectotype

(USNM).

Actia takanoi Baranov, 19350 : 557- LECTOTYPE $, PHILIPPINE REPUBLIC : Los Banos,
I9.V.I928 (S. Takano) (USNM).

Paralectotype : i $, same data as lectotype (USNM).

Alophora albopunctata Baranov, 19350 : 559- Holotype $, JAPAN : Hokkaido, Sapporo,
17. x. 1923 (S. Takano) (USNM).

Paratypes : i <$, JAPAN : Hokkaido, Moiwa, 19. ix. 1923 (S. Takano) (USNM) [head
missing].

Argyrophylax nigrotibialis Baranov, 19350 : 552- Holotype °., FORMOSA : Koshun,
Kankau, ix.i9i2 (H. Sauter) (DEI).

Paratypes : i $, same data as holotype, except date 7-viii.i9i2 (USNM). i $, FORMOSA :
Tainan, Shinkwa, I3.vii.i926 (S. Takano) (USNM). i £, JAPAN : Kanazawa, 2i.viii.i93o
(S. Takano) (USNM) ; i $, same data, except date I9.viii.i93o (USNM). i <$, MALAYA:
Sungai Siakap, 2. hi. 1930 (H. T. Pagden) (BMNH).

No paratypes have been traced from the China, Hangchow, locality mentioned in the
original description.

Arrhinodexia eumorphophaga Baranov, 19340 : 48- Holotype $, MALAYA : Kuala
Lumpur, 5.ix.i927, ex Eumorphus marginatus F. (G. H. Corbett) (BMNH). Genitalia on
slide.

Arrhinomyia issikii Baranov, 19350 : 557- Holotype $, JAPAN : Yumotu, 8.viii.i934 (S.
Issiki) (USNM). Genitalia in situ.

Asiocarcelia pseudocaudata Baranov, 19340" : 4°7- Holotype <$, FORMOSA : Tainan,
iv.igio (H. Sauter) (USNM). Genitalia on slide.

The holotype is labelled by Baranov as ' Carcelia pseudocaudata n.sp. N. Baranoff
TYPUS '.

Bactromyia crassiseta Baranov, 19386 : 409. Holotype $, AUSTRALIA : Queensland,
Biloela [publ. as Biloala], 14.41.1927 (G. A. Currie) (BMNH).

Bactromyia fransseni Baranov, 19340 : 45- Lectotype $, by designation of Crosskey
(1963 : 6), CEYLON : Peradeniya, 8.viii.i928, pupal par. of Psara bipunctalis (J. C. Hutson)
(BMNH). Genitalia on slide.

Paralectotypes : 2 $, 7 $, same data as lectotype (BMNH). i $, same data as lectotype
(USNM). i $, CEYLON : Peradeniya, 2i.vi.i9ig, ex Nacoleia annubilala (J. C. Hutson)
(BMNH). i (J, i ?, CEYLON : Kalutara, 3o.viii.i929, larval par. of Lamprosema diemenalis
on Calapogonium (J. C. Hutson) (<$ in USNM, $ in BMNH). 3 $, JAVA : Buitenzorg,
2g.iii.i932, par. on Cnaphalocrocis medinalis (C. Franssen) (two in MZ, Bogor, one in USNM).

Baranov annotated the original description as follows : ' Originalfundort Java. Cotypus
<J in der Sammlung des Instituuts voor Plantenziekten in Buitenzorg. Cotypus $ in meiner
Sammlung ' ; nevertheless we consider that the material listed before the description is part of
the syntype series, as there is no evidence to show that the description has not been partly
based upon it, and we hold Crosskey 's (1963 : 6) designation of a lectotype from Ceylon as
valid. The female ' Cotypus ' from Java alluded to by Baranov as in his collection is now
in the USNM (see list of paralectotypes above) and in fact bears a Baranov label reading
' Allotypus $ '. We have been unable to confirm whether Baranov's male ' Cotypus ' from
Java with the date 29.iii.i932 is in the collection at Bogor (formerly Buitenzorg), but the
slide of the male genitalia from this specimen is at present in the USNM collection.

36 C. W. SABROSKY & R. W. CROSSKEY

Bactromyia fransseni solomonica Baranov, 19380 : 170. LECTOTYPE <$, SOLOMON
ISLANDS : Russell Island, vi.ig32 (R. J. A. W. Lever) (BMNH). Genitalia on slide.

Paralectotype : i $, same data as lectotype (USNM).

There was no evidence from the original description of solomonica that Baranov had more
than one specimen, and Crosskey (1963 : 7) assumed that the single specimen in the BMNH
collection was the holotype. It has now been found that Baranov's collection in the USNM
contains a second specimen with identical data, so that present designation of a lectotype is
necessary (see above) . The lectotype is in poor condition with loss of one wing, several legs,
and the abdomen is separately card-mounted ; Baranov's original label on the lectotype
reads ' solomonicola ', but solomonica was the published spelling.

Bezziomyiobia nigripes Baranov, 19380 : 172. Holotype $, SOLOMON ISLANDS : Tulagi, 16
[publ. as 6]. xii.i934 (R- J- A. W. Lever) (BMNH).

Blepharipoda eutachinoides Baranov, 19320 : 92. LECTOTYPE <J, FORMOSA : Sokutsu,
ix.i9i2 (H. Sauter) (DEI). Genitalia on slide.

Paralectotype : i <$, same data as lectotype (USNM).

Cadurcia leefmansi Baranov, 1933 : 153. Holotype (as ' Protograph ', figured specimen)
(J, JAVA : Buitenzorg, ex caterpillar of Brachartona catoxantha Hampson (Leefmans) (probably
lost, whereabouts not traced, but slide-mount of genitalia in USNM).

Paratypes : i <£, i $, same data as holotype (USNM). i $, same data as holotype (MZ,
Bogor) .

This species was described from five specimens of both sexes (number of each not specified),
of which one male was referred to by Baranov as the ' Protograph ' and the other four as
' Cotypen '. The unfamiliar term ' protograph ' as used by Baranov is clearly equivalent
to holotype and refers to the single specimen from which the illustration of the male genitalia
was drawn (after removal of the hypopygium) ; this specimen was stated by Baranov to be
in the Instituut voor Plantenziekten in Buitenzorg, Java (now the Lembaga Pusat Penelitian
Pertanian, Bogor) and the slide preparation of its genitalia in his own collection. The
' Protograph ', i.e. holotype, specimen cannot now be found in this institution in Bogor, or
at the Museum Zoologicum Bogoriense, and it is not among Baranov's own collection, and
must be considered probably lost ; but the slide of the genitalia, labelled ' Protograph ',
from Baranov's own collection, is present in the U.S. National Museum.

Of the ' Cotypen ' specimens (i.e. paratypes) two are in USNM collection and one (a com-
plete but teneral male) has been located in the Museum Zoologicum Bogoriense. Baranov
noted in the original publication that C. leefmansi had earlier been identified by Bezzi as
' Degeeria albiceps Macquart ', and the paratype specimens in USNM and MZ all bear this
name as well as Baranov's original labels.

In an earlier work Crosskey (1963) referred to four syntypes of Cadurcia leefmansi in error :
as noted above, there were five original specimens of which one is acceptable as holotype.

Cadurcia vanderwulpi Baranov, 19386 : 410. Holotype $, INDIA : U. P., Haldwani,
Chakrata Range, 4 [publ. as i8].vi.i93o, ex pupa of Hapalia machaeralis (S. N. Chatterjee)
(BMNH).

Baranov published the name vanderwulpi as a ' nom. nov. ' for the misidentified ' Argyro-
phylax zetterstedti, v. d. Wp., nee. B. B., nee Villeneuve '. It is not a replacement name for
a junior homonym, but is an available name for a nominal species based upon the three-line
description given by Baranov ; the sole cited specimen (data above) is the holotype.

Calotheresia (Calotheresiopsis) orientalis Baranov, 19321? : 214. Holotype $, CELEBES :
Tomboekoe [publ. as Tomboegoe] (USNM). Genitalia on slide.

Carcelia aberrans Baranov, 19310 : 27. Holotype $, FORMOSA : Koshun, Kankau, 7-viii.i9i2
(H. Sauter) (DEI). Genitalia on slide.

TYPE-MATERIAL OF TACHINIDAE 37

Carcelia buitenzorgiensis Baranov, 19310 : 45. Lectotype $, by designation of Crosskey
(19676 : 103), JAVA : Buitenzorg, 1919 (W. Roepke) (USNM). Genitalia on slide.

Paralectotypes : 2 <$, same data as lectotype (BMNH & USNM : USNM specimen
represented by genitalia slide only).

Carcelia caudata Baranov, 19310 : 41. LECTOTYPE $, FORMOSA : Koshun, Kankau,
7.viii.i9i2 (H. Sauter) (DEI). Genitalia on slide.

Paralectotypes : i o*. same data as lectotype, except date ix.igia (USNM). i <£,
FORMOSA : Toa Tsui Kutsu, v.i9i4 (H. Sauter) (DEI).

Carcelia caudatella Baranov, 19320* : i. Holotype <£, SUMATRA : Siberut Island, ix.ig24
(C. B. K. &> N. S.) (MZ, Bogor). Genitalia on slide.

The Baranov collection in USNM contains a male specimen of caudatella labelled by
Baranov as ' n.sp. ', but it has the data ' Karimen Djawa, v.ig26 (Dammerman) ' (not cited
in the original publication) and is not a type-specimen.

Carcelia distincta Baranov, 19310 : 32. Holotype <$, FORMOSA : Sokutsu, ix.i9i2 (H.
Sauter) (DEI). Genitalia on slide.

At the time of writing the holotype is temporarily in the collection of Dr. L. P. Mesnil,
at Delemont, Switzerland.

Carcelia frontalis Baranov, 19310 : 43. Holotype Q*. FORMOSA : Toa Tsui Kutsu, v.1914
(H. Sauter) (DEI). Genitalia on slide.

Carcelia hirsuta Baranov, 19310 : 38. LECTOTYPE <J, FORMOSA : Koshun, Kankau,
7.viii.i9i2 (H. Sauter) (DEI). Genitalia in situ.

Paralectotype : i <$, same data as lectotype, except date 7.vii.i9i2 (DEI) [abdomen
missing] .

Carcelia malayana Baranov, 1934^ : 4°4- Holotype <£, MALAYA : Malay Peninsula, Kuala
Lumpur, 2. v. 1932 (BMNH). Genitalia on slide.

Carcelia octavo Baranov, 19310 : 35. LECTOTYPE J, FORMOSA : Koshun, Kankau, ix. 1912
(H. Sauter) (DEI). Genitalia in situ. Lectotype designated from ' octava A ', see discussion
below.

Paralectotypes : i $, same data as lectotype (USNM). 3 <£, same data as lectotype (one
in DEI, two in BMNH). i <£, same data as lectotype, except date 7.viii.i9i2 (USNM).
3 6*. same data as lectotype, except date viii.i9i2 (DEI), i <$, same data as lectotype,
except date 7.ix.i9i2 (DEI). 2 o*, same data as lectotype, except date iv.igi2 (DEI), i $,
same data as lectotype, except date iii.igi3 (DEI), i $, same data as lectotype, except
date 22. vi. 1912 (BMNH).

Baranov described octava in two forms, A and B. The lectotype and all above-listed
paralectotypes are of ' form A ' and are so labelled by Baranov. The DEI collection contains
in addition one paralectotype of ' form B ', labelled as such by Baranov, and with the same
data as the lectotype, except for the date viii.i9i2.

Carcelia pilosa Baranov, 19310 : 29. LECTOTYPE $, JUGOSLAVIA : Bosnia, Sarajevo
(USNM). Genitalia on slide.

We have not seen the second original syntype, locality unknown to us, stated by Baranov
to be in the Riedel collection, but Mesnil (1944 : 29) has noted that it is very close to
Carcelia excisa (Fallen).

Carcelia pilosella Baranov, 19310 : 37. LECTOTYPE <J, FORMOSA : Koshun, Kankau.

7.vii.i9i2 (H. Sauter) (DEI). Genitalia on slide.

Paralectotypes : i $, same data as lectotype, except date vi.i9i2 (USNM). i $, same

data as lectotype, except date ix.i9i2 (DEI).
Carcelia prima Baranov, 19310 : 31. LECTOTYPE $, FORMOSA : Koshun, Kankau, 22. vi.

1912 (H. Sauter) (DEI). Genitalia in situ. Lectotype designated from 'prima B ', see

following discussion.

38 C. W. SABROSKY & R. W. CROSSKEY

Paralectotypes : Form prima B : i <$, same data as lectotype (DEI), i <$, same data
as lectotype, except date vii.igia (DEI), i <$, same data as lectotype, except date viii.igia
(USNM). 2 c£, same data as lectotype, except date ix.igi2 (USNM & BMNH). Form
prima A : i <$, same data as lectotype (DEI). 2 <J, same data as lectotype, except date
iv.igi2 (DEI), i $, same data as lectotype, except date vi.iQi2 (USNM). 3 $, same data
as lectotype, except date viii.i9i2 (two in DEI, one in BMNH). i $, same data as lecto-
type, except date 7.viii.i9i2 (DEI), i <J, 2 $, same data as lectotype, except date ix.igi2
(DEI), i <J, same data as lectotype, except date 7.ix.i9i2 (DEI), i <$, same data as lecto-
type, except date 7.xi.igi2 (DEI), n $, FORMOSA : Sokutsu, ix.iQi2 (six in DEI, three in
BMNH, two in USNM). i <J, i $, FORMOSA : Taihorinsho, ix.igog (DEI) ; i <j>, same data
(USNM).

In the original description Baranov treated Carcelia prima as two forms, prima A in which
there are two reclinate orbital setae and prima B in which there is only one reclinate orbital
seta ; he did not state the number of specimens of either form, but the specimens we have
located and recorded above must represent most of the original syntype material for both
forms. In a later paper Baranov (19340" : 396-39?) considered A and B to be distinct
species and he retained the name prima for the species represented by B ; we have therefore
selected a specimen of form B as the lectotype of Carcelia prima (see above). Baranov
(19340") treated his form A in synonymy with Eucarcelia kockiana (Townsend), but we are
not able to say at this time whether this is correct.

We have not seen the specimen of prima A from Tsingtau cited by Baranov in the original
publication, but this will be another paralectotype if it is ever located.

It should be noted that the female paralectotype from Taihorinsho in the DEI collection
is wrongly associated and has the hind coxa bristled and a ventral submedian seta on the
middle tibia (in all the other material the hind coxa is bare and the mid tibia lacks a v sub-
median seta) . The lectotype, all paralectotypes from prima B, and most of the paralectotypes
from prima A, have the basicosta a clear yellow-orange colour, but in some of the paralecto-
types of prima A the basicosta is darker and distinctly browned on the fore margin (so it is
possible that prima A paralectotypes could be an admixture of specimens from two very
closely allied species). The lectotype and paralectotypes of prima B have only one pair of
reclinate orbital setae, and the paralectotypes of prima A have two pairs of reclinate orbital
setae, but it is not fully certain, despite Baranov's (19340") treatment, whether this is evidence
that two species are involved.

Carcelia quarta Baranov, 193 10 : 33. Holotype <J, FORMOSA : Gebiet des Sh'shastammes,
v-vi.igi2 (H. Sauter) (DEI). Genitalia on slide.

Carcelia quinta Baranov, 19310 : 33. LECTOTYPE £, FORMOSA : Koshun, Kankau,
ix.i9i2 (H. Sauter) (DEI). Genitalia in situ. Lectotype designated from ' quinta A ', see
discussion below.

Paralectotypes : i Q*. same data as lectotype, except date y.xi.1912 (BMNH). i $, same
data as lectotype, except date 22. vi. 1912 (USNM). i <$, same data as lectotype, except
date vi.igi2 (DEI).

Baranov described quinta in two forms, A & B, denned by differences in the relative widths
of the interfrontal area and parafrontals ; the number of original specimens of each form
was not stated, but so far as we can tell the four specimens cited above form the complete
type-series for both forms. The lectotype and each of the paralectotypes in USNM and
BMNH is labelled by Baranov as ' quinta A ', and the paralectotype in DEI collection is
labelled by Baranov as ' quinta B '.

Carcelia rasella Baranov, 19310 : 44. LECTOTYPE <$, JUGOSLAVIA : Serbia, Golubac.
i. v.i 927 (USNM). Genitalia on slide.

Paralectotypes : 2 <$, same data as lectotype (USNM).

TYPE-MATERIAL OF TACHINIDAE 39

Carcelia rasoides Baranov, 19310 : 42. LECTOTYPE $, FORMOSA : Koshun, Kankau,
22. vi. 1912 (H. Sauter) (DEI). Genitalia on slide.

Paralectotypes : i <$, same data as lectotype (DEI). 2 o\ same data as lectotype, except
date ix.igi2 (DEI & BMNH).

Carcelia rufa Baranov, 19310 : 33. LECTOTYPE Q*. FORMOSA : Macuyama, vi.i9i4 (H.
Sauter) (DEI). Genitalia in situ.

Paralectotypes : 3 $, FORMOSA : Koshun, Kankau, 7. v. 1912 (H. Sauter) (DEI, USNM &
BMNH) ; i $, same data (DEI), i <J, FORMOSA : Koshun, Kankau, 22. vi. 1912 (H. Sauter)
(DEI) ; i ?, same data (USNM).

Carcelia rutilloides Baranov, 19310 : 29. Holotype $, FORMOSA : Chosokei, 1914 (H.
Sauter) (DEI).

Carcelia secunda Baranov, 19310 : 31. Holotype $, FORMOSA : Sokutsu, ix.igi2 (H.
Sauter) (DEI). Genitalia on slide.

Carcelia septima Baranov, 19310 : 35. LECTOTYPE <J, FORMOSA : Koshun, Kankau,
viii.i9i2 (H. Sauter) (DEI). Genitalia on slide.

Paralectotypes : i 5*. same data as lectotype (USNM). i $, same data as lectotype
(BMNH). i $, same data as lectotype, except date ix.igi2 (DEI).

Carcelia setosella Baranov, 19310 : 44. Holotype Q*. FORMOSA : Sokutsu, v.i9i2 (H. Sauter)
(DEI). Genitalia on slide.

Carcelia sexta Baranov, 19310 : 34. Holotype <$, FORMOSA : Taihorinsho, ix.igog (H.
Sauter) (DEI). Genitalia on slide.

Carcelia tertia Baranov, 19310 : 32. Holotype (J, FORMOSA : Taihorinsho, ix.igog (H.
Sauter) (DEI). Genitalia on slide.

Catacarcelia rondaniella Baranov, 19340* : 392. LECTOTYPE Q", FORMOSA : Koshun,
Kankau, 7.vii.igi2 (H. Sauter) (USNM). Genitalia on slide.

Paralectotypes : i <$, FORMOSA : Takao, ig.xii.igo7 (H. Sauter) (USNM). i °., same
data as lectotype (USNM).

The two paralectotypes each bear a label with the manuscript name ' Exorista carcelioides
Baranov n.sp. ' and a second label ' Carcelia rondaniella n.sp. N. Baranoff ', both in Baranov's
writing.

Chaetexorista solomonensis Baranov, 1936 : 101. Holotype $, SOLOMON ISLANDS :
Shortland, Korovo, 23.^.1934 (H. T. Pagden) (BMNH). Genitalia in situ.

Chaetoptiliopsis burmanica Baranov, 19386 : 411. Holotype <$, BURMA : Northern Shan
States, Panghai Res., Namtu, R.O., 26^.1934, ex C. [alopepla] leayana (BMNH). Genitalia
in situ.

Paratype : i $, same data as holotype, except date 25^.1934 (BMNH).

The USNM Collection contains a male and a female from the same rearing, with similar
data except for the date 28^.1934, but these are not recorded in the original description and
are not type-material.

Senior White, Aubertin & Smart (1940 : 82) erroneously placed this genus and species in
the Calliphoridae.

Cnephalia sillemi Baranov, 1935^ : 407. Holotype <$, CHINA : Sinkiang, Karakorum Range,
Karakash Valley, between Kawak Pass and Sanju Pass [approx. 36° 30' N. and 78° 15' E.],
3700-3200 m, i6.ix.-5.x.ig29 (/. A. Sillem) (Zoologisch Museum, Amsterdam). Genitalia
in situ.

Ctenophoroceropsis yerburyi Baranov, 19386 : 409. Holotype^1, SOUTH YEMEN REPUBLIC :
Aden, 2i.ii.i8g5 (Yerbury) (BMNH). Genitalia in situ.

Paratypes : i <?, same data as holotype (BMNH). 2 <$, same data as holotype, except
dates g.ii.i8g5 and 26.ii.i8g5 (BMNH) (both headless), i <J, same data as holotype, except
date 2o.ii.i8g5 (USNM).

40 C. W. SABROSKY & R. W. CROSSKEY

Dexiomimops ruflpes Baranov, 19350 : 557. Holotype $, JAPAN : Maoka, Karafuto,
2i.viii.ig23 (•$• Takano) (USNM). Genitalia in situ.

The Baranov collection in USNM contains two males from Formosa (no other data) that are
here not considered to be part of the type-series. After listing the type, Baranov stated
' Auch von Formosa mir bekannt ', without giving details and without clearly including the
specimens in his type-series.

Doleschalla solomonensis Baranov, 19346 : 182. Holotype $, SOLOMON ISLANDS :
Guadalcanal, Lunga, iv.i932 (R. J. A. W. Lever) (BMNH). Genitalia in situ.

Paratypes : i <$, SOLOMON ISLANDS : Tulagi, 23.vii.i933 (R. J. A. W. Lever) (BMNH) ;
i (£, same data, except date 3o.vii.i933 (USNM).

Baranov (i934c : 475), in a second paper in the same month (August 1934), published
records of four males of D. solomonensis, giving two localities not mentioned for the original
series ; these males appear to be additional material, and are presumed to be not type-
material. In this second paper (Baranov, 1934^) there is a figure of the male genitalia, but
no diagnosis in words, and under the International Code of Zoological Nomenclature, 1961
(Article i3a (i)) the name solomonensis would be nomenclaturally unavailable from this
second paper even if it should be found to be the earlier of the two publications.

Dolichocolon australe Baranov, 19386 : 405. LECTOTYPE $, AUSTRALIA : Queensland,

Gympie, iii.1932, ex Spodoptera exempta (W. H. T. Summerville) (BMNH). Genitalia in situ.

Paralectotypes : i $, same data as lectotype (BMNH). 1^,1$, same data as lectotype

(USNM). 2 $, AUSTRALIA : Queensland, Gadgarra, 6.^.1932, ex Spodoptera (J. Harold

Smith) (USNM).

Dolichocolon orbitale Baranov, 19386 : 406. LECTOTYPE <J, INDIA : C.P., Rahatgaon,
Hoshangabad, 25. ix. 1926, ex Hapalia machaeralis (S. N. Chatter jee) (BMNH). Genitalia in
situ.

Paralectotypes : i $, same data as lectotype, except date 4. x. 1926 (USNM). i $ (genitalia
slide only), same data as lectotype, except date i6.viii.i926 (USNM). i °-, same data as
lectotype, except date i6.viii.ig26 (USNM). i °-, same data as lectotype, except date
2.x. 1926 (BMNH).

Beeson & Chatter] ee (1935 : 173) had already published this name, prior to Baranov's
description, and had given a figure of the entire fly together with an account of the life
history, but these authors gave no description and the name is therefore nomenclaturally
unavailable from Beeson & Chatterjee (1935) under Article i3a (i) of the International Code
of Zoological Nomenclature, 1961.

Dolichocolon quadrisetosum Baranov, ig35a : 555. LECTOTYPE $, FORMOSA : Koshun,
Kankau, vii.igi2 (H. Sauter) (DEI).

Paralectotypes : i $, same data as lectotype, except date 7.vii.igi2 (DEI). 3 $, same
data as lectotype, except date 7.viii.igi2 (two in USNM, one in DEI). 2 Q*, FORMOSA :
Takao, 31. x. and 8.xii.igo7 (H. Sauter) (USNM). i $, FORMOSA : Takao, 8.xii.igo7 (H.
Sauter) (IZPAN, Warsaw).

We accept the two males from Takao (in USNM collection) as syntypes as they are labelled
as ' n.sp. ' by Baranov, but it should be noted that only the female was mentioned in the
original description. We have not seen the Formosan specimen with the data ' Shinkwa,
12. iv. 1932 (S. Takano) ' mentioned by Baranov, but this will be another paralectotype if
ever located.

Dolichocolon rufescens Baranov, 19386 : 406. LECTOTYPE <£, AUSTRALIA : New South
Wales, Yantabulla, 25^.1916 (BMNH). Genitalia in situ.

Paralectotype : i $, same data as lectotype, except date 25. vi. 1916 (Siddens) (USNM).

Echinomyia praeceps aestivalis Baranov, ig2ga : 14. LECTOTYPE $, JUGOSLAVIA :
Macedonia, Kavadar, I5.vi.ig27 (USNM).

TYPE-MATERIAL OF TACHINIDAE 41

Paralectotypes : i 9, same data as lectotype, except date ig.vii.igay (USNM). i $, same
data as lectotype, except date 5.vii.ig27 (CNC). i 9. same data as lectotype, except date
5.vii.i927 (USNM). i $, JUGOSLAVIA : Macedonia, Skoplje, n.ix.i928 (CNC).

We accept Baranov's ' cotypus ' specimens from Kavadar with the date 5.vii.iQ27 as
paralectotypes, but it should be noted that in the original publication Baranov cited this date
for specimens from Skoplje : we consider that Baranov probably made a slight error in
recording the data. Baranov noted specimens from Belje (22.vi.-5.vii.i923), Bitol (9.ix.
1928) and Struga (y.ix.i928) in the original publication but none of these have been located ;
they will be additional paralectotypes when found.

Echinomyia praeceps vernalis Baranov, 19290 : 13. LECTOTYPE <$, JUGOSLAVIA :
Serbia, Golubac, 9^.1927 (USNM). Genitalia in situ.

Paralectotypes : 2 <J, same data as lectotype (USNM & CNC).

The specimens from Skoplje (14. iv. & 17^.1927) and TopCider (8 & 14^.1924) mentioned
in the original publication have not been located, but will be additional paralectotypes if
later found.

Erycia bezzii Baranov, 19340 : 44. LECTOTYPE 9, MALAYA : Kuala Lumpur, 24 [publ. as
29].viii.i93i, ex Telicota palmarum Moore (BMNH).
Paralectotype : i 9, same data as lectotype (USNM).

There was no evidence from the original description of bezzii that Baranov had more than
one specimen, and Crosskey (19676 : 103) cited the specimen in BMNH collection as holo-
type. It has now been found that Baranov's collection in USNM contains a second specimen
with identical data, so that present designation of a lectotype is necessary (see above).

Erycia intermedia Baranov, 1939 : in. Holotype 9, JAPAN : Hokkaido, Sapporo, i3.viii.
1923 (K. Tamanuki) (USNM).

Erycia nigricosta Baranov, 1936 : 99. Holotype <$, SOLOMON ISLANDS : Guadalcanal,
Kaukau [publ. as Kankau], 22.viii.i934 (R- ]• A. W. Lever) (BMNH). Genitalia on slide.

Erycia nymphalidophaga Baranov, 1936 : 112. LECTOTYPE <$, INDIA : U.P., Dehra
Dun, 5.x. 1929, ex nymphalid on Citrus aurantium (N. C. Chatter jee) (BMNH). Genitalia
in situ.

Paralectotypes : 3 9. same data as lectotype, except dates 6, 10 and n.x.i929 (BMNH).
i 9. same data as lectotype, except date io.x.i92g (USNM).

Beeson & Chatterjee (1935 : 174) published this name prior to Baranov's description, but
gave only a three-line note on the life history and no description ; the name is nomen-
claturally unavailable from Beeson & Chatterjee (1935) under Article i3a (i) of the Inter-
national Code of Zoological Nomenclature, 1961.

Erycia palpata Baranov, 1936 : 113. Holotype 9. FORMOSA : Toa Tsui Kutsu, v.i9i4 (H.

Sauter) (USNM).
Erycia rufofemorata Baranov, 1936 : 112. Holotype 9, JAVA : Buitenzorg, ii.i933 (R- W.

Paine) (BMNH).

Paratype : i 9, same data as lectotype, except date 1.1933 (USNM).

Erycia takanoi Baranov, 1939 : in. LECTOTYPE <J, JAVA : Pasoeroean, iii.i926 (S.

Takano) (USNM). Genitalia in situ.

Paralectotype : i 9, same data as lectotype (USNM).

Eucarcelia caspica Baranov, 1934^ : 390. Holotype $, ? U.S.S.R. or IRAN : Caspian Sea
region, Talysch [? = Talish], 1897 (Korb) (USNM). Genitalia on slide.

Eucarcelia dammermani Baranov, 1934^ : 393. LECTOTYPE $, JAVA : Idjen, 1850 m,
Ongop-Ongop, v.ig24 (Dammerman) (USNM). Genitalia on slide.
Paralectotype : i 9, same data as lectotype (MZ, Bogor).

Eucarcelia grossa Baranov, 1934^ : 393- Holotype <J, FORMOSA : Tainan, iv.igio (H.
Sauter) (USNM). Genitalia on slide

42 C. W. SABROSKY & R. W. CROSSKEY

Eucarcelia indica Baranov, 19340* : 394- Holotype <$, INDIA : Silhar Kalhar, 27. vi. 1911
(USNM). Genitalia on slide, head missing.

Eurystaea leveriana Baranov, 19346 : 182. Holotype $, SOLOMON ISLANDS : Malaita, Su'u,
iv.i933 (R- J- A. W. Lever) (BMNH).

Eutachina argenteostriata Baranov, 19380 : 171. Holotype $, SOLOMON ISLANDS :
Guadalcanal, Kovagombi, i.v.1936 (R. J. A. W. Lever) (BMNH).

Eutachina aureifrons aureifrons Baranov, 1936 : 107. LECTOTYPE <$, JAVA : Idjen,
Kendeng, 1400 m, 111.1924 (Dammermari) (MZ, Bogor). Genitalia in situ.

Paralectotype : i <$, JAVA : Idjen, Blawan, 950 m, vi.i924 (Dammerman) (USNM).

In the original description Baranov cited the locality simply as ' Java ' without additional
data, and stated ' Typus im Zoologischen Museum in Buitenzorg '. The collection of the
Zoological Museum in Bogor contains one male specimen but it is actually labelled as ' Para-
typus ', and the male specimen in USNM collection is labelled as ' Typus '; however, the
slide preparation of the genitalia from the USNM collection is labelled as ' Cotypus ' and
not as ' Typus '. We consider that the two males are syn types, and the specimen in MZ,
Bogor has been designated as lectotype to conform with Baranov 's statement of the
depository.

The USNM collection contains a female specimen with the same data as the lectotype
(except that the date is given ambiguously as both iii and vi), but as only the male sex was
mentioned in the description and as the female specimen is labelled as ' aureifrons mihi N.
Baranoff ' (i.e. not with an original ' n.sp. ' label) we consider that it is not an original syntype.

Eutachina aureifrons sumatrana Baranov, 1936 : 107. LECTOTYPE $, SUMATRA :
Selemoekae, viii.1925 (O. Posthumus) (USNM). Genitalia on slide.

Paralectotype : i $, SUMATRA : Goen-mongko, 7.viii.i925 (O. Posthumus) (USNM).

Eutachina aureisquamosa Baranov, 19386 : 410. Holotype $, SOLOMON ISLANDS :
Guadalcanal, Oreke, 700 ft., I4.xii.i934 (R. J. A. W. Lever) (BMNH). Genitalia in situ.

Eutachina aurichalcea Baranov, 1936 : 100. Holotype $, BOUGAINVILLE ISLAND : Kieta,
v.1934 (/• L. Froggatt) (BMNH).

Eutachina basalts Baranov, 19320 : 86. Holotype $, FORMOSA : Koshun, Kankau, ix.i9i2
(H. Sauter) (DEI). Genitalia on slide.

Eutachina civiloides Baranov, 19320 : 84. LECTOTYPE <$, FORMOSA : Koshun, Kankau,
vii.i9i2 (H. Sauter) (DEI). Genitalia on slide.

Paralectotypes : i <J, i °-, same data as lectotype, except date 7.viii.igi2 (USNM & DEI).

Eutachina fuscipennis Baranov, 19320 : 90. LECTOTYPE <J, FORMOSA : Koshun, Kankau,
viii.i9i2 (H. Sauter) (DEI). Genitalia on slide.

Paralectotypes : i <£, same data as lectotype (USNM). i ^, same data as lectotype,
except date iv.i9i2 (DEI).

The DEI collection contains two females from Kankau, date ix.i9i2, determined by
Baranov as fuscipennis, but these are not part of the original type-series.

Eutachina hyalipennis Baranov, 19320 : 88. LECTOTYPE <$, FORMOSA : Chipun, vii.
1912 (H. Sauter) (DEI). Genitalia on slide.

Paralectotypes : i <$, FORMOSA : Koshun, Kankau, 7.viii.i9i2 (H. Sauter) (DEI) ; i <$,
same data, except date ix.i9i2 (USNM).

A female specimen determined by Baranov as hyalipennis and having similar data
(Kankau, vi.i9i2) is in DEI collection but is not an original syntype.

Eutachina ladelli Baranov, 1936 : 108. Holotype $, THAILAND : Hua Hin, iv.i926
(W. R. S. Ladell) (BMNH). Genitalia on slide.

TYPE-MATERIAL OF TACHINIDAE 43

Eutachina tnungomeryi Baranov, 19386 : 410. LECTOTYPE $, AUSTRALIA : Queensland,
Gordonvale, 2.111.1936, ex Laphygma exempta Walker (R. W . Mungomery) (BMNH). Genitalia
in situ.

Paralectotypes : i $, same data as lectotype (BMNH). i <$, same data as lectotype
(USNM). i $, same data as lectotype, except collector (7. W. Buzacott) (USNM).
Eutachina quadriseta Baranov, i932a : 91. Holotype <$, FORMOSA : Sokutsu, ix.i9i2
(H. Sauter) (DEI). Genitalia on slide.

Baranov (19380! : 171) later recorded specimens from the Solomon Islands as Eutachina
quadrisetosa Bar. and compared them with Formosan quadrisetosa, but this is a lapsus and
clearly an erroneous subsequent spelling of quadriseta Baranov.

Eutachina rusticella Baranov, 1936 : 108. LECTOTYPE <$, FORMOSA : Takao, 28. xi. 1907
(H. Sauter) (IZPAN, Warsaw). Genitalia on slide.

Paralectotypes: i Q\ same data as lectotype (USNM) . i $, SUMATRA: [no other locality
data] 8.ix.i93o (USNM : genitalia preparation only).

In the original description Baranov recorded ' Sumatra (Imperial Institute of Entomology) '
among the original material seen, and there must therefore have been at least one syntype
from Sumatra. We have been unable to locate a complete specimen from Sumatra, but the
Baranov collection in USNM contains a slide preparation of the male genitalia of a specimen
of rusticella labelled by Baranov as from Sumatra, with the date 8.ix.i93o, and with the
abbreviated words ' Imper. Inst. ', and we consider this slide to confirm that there was at
least a male syntype from Sumatra ; we therefore record it above as one of the paralectotypes.
Tachinid material handled by Baranov from the Imperial (now Commonwealth) Institute of
Entomology was normally deposited in the British Museum (Natural History) collection or
returned to the sender, but in this case (as the body of the syntype has not been found) the
genitalia slide preparation is retained in the USNM collection, whence it can be returned if
the associated specimen is found.

The USNM collection also contains a male specimen of rusticella from the same locality as
the lectotype, but with the date 6.xii.i9O7 ; this date does not fit with the original published
data and the specimen is considered not to be a syntype.

Eutachina tenuiforceps Baranov, 19320 : 87. Holotype Q*, FORMOSA : Koshun, Kankau,
vii.i9i2 (H. Sauter) (DEI). Genitalia on slide.

The DEI collection contains three females from Kankau determined by Baranov as
tenuiforceps, but this species was described only from the male and these females are not
original type-material (though listed as ' Typen ' by Hennig, 1941 : 194).

Euthelairosoma siamense Baranov, 19386 : 411. Holotype <J, THAILAND : Siam [also
bearing some other undecipherable data] (BMNH). Genitalia in situ.

Euvespivora orientalis Baranov, 1942 : 162. Holotype <£, JAVA : Delawa, io.v.i937, teak
forest ex larva of Vespa analis (L. G. E. Kalshoven) (USNM). Genitalia in situ.

Euvespivora salotnonica Baranov, 1942 : 163. Holotype $, SOLOMON ISLANDS : Tulagi,
17.11.1938 (R. J. A. W. Lever) (BMNH).

Exorista distincta Baranov, 19316 : 120. LECTOTYPE <$, FORMOSA : Koshun, Kankau,
7.xi.i9i2 (H. Sauter) (DEI). Genitalia in situ.

Paralectotypes : i <$, same data as lectotype (USNM). i $, same data as lectotype,
except date 7.viii.i9i2 (BMNH). 2 9, same data as lectotype, except date 7.viii.i9i2 (DEI
& USNM). i 9, same data as lectotype, except date ix.igi2 (BMNH).

In the original publication Baranov mentioned a total of five specimens (2 <$, 3 9) but did
not mention the month date ' xi '. We have traced a total of six specimens (see above), two
of which have the month date ' xi ' ; all are labelled as ' n.sp. ' by Baranov and we consider
all to be original syntypes. We consider the discrepancies to be due to inadvertent lapse by
Baranov in recording the material.

Exorista phrynoides Baranov, 1939 : no. Holotype <$, JAPAN : Hokkaido, Sapporo,
22. vi. 1924 (S. Takano) (USNM). Genitalia in situ.

44 C. W. SABROSKY & R. W. CROSSKEY

Exorista picta Baranov, 19350 : 553- LECTOTYPE ^, FORMOSA : Koshun, Kankau,
y.vii.igia (H. Sauter) (DEI). Genitalia in situ.

Paralectotypes : i 9, same data as lectotype (BMNH). 2 $, same data as lectotype,
except date ix.igia (DEI & USNM). 2 $, same data as lectotype, except date ix.igia
(DEI), i o*, same data as lectotype, except date vii.igia (DEI), i <$, same data as lecto-
type, except date 4.viii.i9i2 (USNM). i <J, i $, same data as lectotype, except date 7. viii.
1912 (BMNH). i $, same data as lectotype, except date y.xi.i9i2 (USNM). 3 $, FORMOSA :
Tainan, v.i9i2 (H. Sauter) (DEI), i $, FORMOSA : Shinkwa, 19^1.1932 (S, Takano) (USNM).

Exorista quadrimaculata Baranov, 19340 : 43- Lectotype <$, by designation of Crosskey
(19676 : 104), MALAYA : Klang, 9.11.1931 (BMNH). Genitalia in situ.

Paralectotypes : i $, same data as lectotype (BMNH). i 5*, i °-, MALAYA : Estate K.
Lipis, 30. iv. 1 93 1, host Psychidae ? Clania variegata (G. H. Corbett) (USNM). i <$, SUMATRA :
E.G., Pematang Siantar, 13.1.1932 (R. I. Nel) (BMNH) ; i <$, same data, except date 21. xi.
1931 (USNM). i o*. CEYLON : Ratnapura, 27. vi. 1922, ex psychid (/. C. Hutson) (BMNH).

Although Baranov headed the original description of quadrimaculata with a ' 6* ' sex
symbol only, we have accepted two female specimens as part of the original syntype series and
have listed them as paralectotypes above. Our reasons for this are, firstly, that these
females have similar data to the males and bear Baranov's usual type of ' n.sp. ' label ;
and, secondly, that elsewhere in the same paper Baranov has introduced a note on the
' Weibchen ' in descriptions headed ' 6* ' only, in such a way that the descriptions appear to
be based only on the male but were actually drawn from both sexes. We therefore infer that
Baranov had both sexes before him but that the symbol ' $ ' was inadvertently omitted.

Exorista seniorwhitei Baranov, 19386 : 408. Holotype $, INDIA : [Assam] Khasia Hills,
Mawphlang [publ. as Mauphlong], io.x.i92o (R. Senior-White) (BMNH). Genitalia in situ.
Paratype : i <J, INDIA : [Assam, Khasia Hills] Laitlyngkot, i6.x.i92o (R. Senior-White)
(USNM).

The localities of the type-material are not easy to trace in atlases, but are mentioned by
Senior- White (1922) in his paper on the Diptera of the Khasia Hills.

Exorista vicinalis Baranov, 19316 : 123. LECTOTYPE <$, FORMOSA : Koshun, Kankau,
viii [publ. as vii].i9i2 (H. Sauter) (USNM).

Paralectotype : i $, same data as lectotype (USNM).

The lectotype lacks the genitalia and the slide preparation has not been located.

Fabriciella pandellei Baranov, i g2ga : 19. LECTOTYPE £, JUGOSLAVIA : Bosnia, Trebevie
(USNM). Genitalia in situ.

Paralectotype : i <$, JUGOSLAVIA : Croatia, Sejeme, 19. vi. 1918 (USNM).

In addition to the two specimens above mentioned the USNM collection contains a male
specimen with data ' Sejeme, Croatia, 2. viii. 1929 ', but this is not considered to be an
original syntype ; Baranov's description mentions only specimens with month dates ' VI.—
VII.' and collection in August would probably be too late for citation of the material in a
paper published in the same year.

Formosia mirabilis solomonicola Baranov, 1936 : 101. LECTOTYPE 6*. SOLOMON
ISLANDS : Guadalcanal, Kaukau [publ. as Kankau], 21. viii. 1934 (R-J- A. W. Lever] (BMNH).
Genitalia in situ.

Paralectotypes : i °-, BOUGAINVILLE [publ. as Guadalcanal] : Kieta, v.i934 (/• L. Froggatt)
(BMNH). i (J, i $, SOLOMON ISLANDS : New Georgia, Segi, Morovo Lagoon, 5^.1934
(H. T. Pagden) (USNM). i °-, SOLOMON ISLANDS : Tulagi, Ridge, 31.111.1934 (H. T. Pagden)
(BMNH). i $, SOLOMON ISLANDS : Montgomery, Tetipari, 12. v. 1934 (H. T. Pagden)
(BMNH). i $, SOLOMON ISLANDS : Isabel [publ. as Isabbel], 111.1932 (R. J. A. W. Lever)
(BMNH).

We have not seen the specimens (other paralectotypes from Solomon Islands) mentioned
by Baranov with the data ' Renodova, 14^.1934, leg. H. T. Pagden ' and ' Calwel, 1.1932 '
(a $ and Q* respectively), and their location is unknown to us.

TYPE-MATERIAL OF TACHINIDAE

45

Goniophthalmus dubiosus Baranov, 19360 : 555. LECTOTYPE <$, JAVA : Pasoeroean,
io.iii.ig26 (S. Takano) (USNM). Genitalia in situ.
Paralectotype : i $, same data as lectotype (USNM).

There is a female specimen in the USNM collection labelled as new species but with the
data ' Dammerman N.O. Soemba, Kambera, 4.111.1925 '. No such specimen is mentioned
in the original description and it is not a syntype.

Gymnodexia atkinsoni Baranov, 19340 : 49. LECTOTYPE <$, BURMA : Maymyo,
Mandalay District, 13^1.1930, ? ex Curculionidae, on Phyllanthus emblica (D. J. Atkinson)
(BMNH). Genitalia in situ.

Paralectotypes : i <$, same data as lectotype (USNM). i o*> i ?, same data as lectotype,
except dates 8.vi. and 17^1.1930 respectively (BMNH).

Although Baranov headed the original description of atkinsoni with a ' <$ ' sex symbol
only, we have accepted one female specimen as part of the original syntype series and have
listed it above as a paralectotype. Our reasons for this are, firstly, that the specimen has
identical data with the lectotype (except for slight difference in date) and bears the usual
type of Baranov ' n.sp. ' label ; and, secondly, that elsewhere in the same paper Baranov
has introduced a note on the ' Weibchen ' in descriptions headed ' $ ' only, in such a way that
the descriptions appear to be based only on the male but were actually drawn from both sexes.
We therefore infer that Baranov almost certainly had the female before him at the time of
description, and accept it as an original syntype (see similar situation discussed under Exorista
quadrimaculata Baranov) .

Hapalioloemus machaeralis Baranov, I934/" : l62- Holotype <$, INDIA : C.P., Rahatgaon,
y.viii.1926 (S. N. Chatter jee) (BMNH). Genitalia in situ.

Hemidegeeria villeneuvei Baranov, 19340 : 44. LECTOTYPE $, BURMA : Shwegu Res.,
Bhamo, 25.111.1930, ex Acacia pruinescens (D. J. Atkinson) (BMNH). Genitalia in situ.

Paralectotypes : 3 $, same data as lectotype, except dates 29-iii., 27.lv. and 24^.1930
(BMNH). i o*, i $, same data as lectotype, except dates i.iv. and 2i.iv.i93o respectively
(USNM). i °-, same data as lectotype, except date 3.^.1930 (USNM).

Ilia tnirabilis Baranov, 19380 : 172. Holotype ?, SOLOMON ISLANDS : Guadalcanal, Lunga
Estate, 4.vi.i935 (R. J. A. W. Lever] (BMNH).

The USNM collection contains a specimen of this species bearing the label ' Ilia mirabilis
g.n. sp.n. N. Baranoff ' in Baranov's writing, but it is not mentioned in the original publica-
tion and is not a type-specimen. The data are : $, SOLOMON ISLANDS : Savo Island,
Tasimania, 500-700 ft., 24^1.1935 (R. J. A. W. Lever).

Isocarceliopsis hemimacquartioid.es Baranov, 19340" : 406. LECTOTYPE $, FORMOSA :
Toa Tsui Kutsu, v.i9i4 (H. Sauter) (DEI).

Paralectotypes : 2 <$, FORMOSA : Takao, 8.xii.i9O7 (H. Sauter) (IZPAN, Warsaw) ;
i (J, same data, except date 2i.xii.i9O7 (USNM) ; i <J, same data, except date 6.xii.i9O7
(BMNH). In addition the following paralectotypes that are not conspecific with the lecto-
type (see discussion) : 2 $, FORMOSA : Tainan, iv.igio (H. Sauter) (DEI & USNM).

It is not clear from the format in the original publication how many specimens were avail-
able to Baranov at the time of description, for he cited only the following information :
' FORMOSA : einige Exemplare im Deutschen Entomologischen Institut in Berlin-
Dahlem. — Tainan, iv.igio und Takao, xii.igo7 (H. Sauter) ; Museum in Stettin '. Normally
this punctuation would indicate in Baranov's work that he saw some specimens in the DEI
collection from unspecified localities, and also specimens in Stettin from Tainan and Takao ;
in fact the Stettin collection (now in Warsaw) contains specimens only from Takao, and the
DEI collection contains specimens from Tainan and Toa Tsui Kutsu (a locality not actually
cited by Baranov at the time of description, but recorded later by Hennig, 1941 : 198 as
the locality of one of Baranov's types). In addition there are specimens from Baranov's
own collection that are now in the USNM or BMNH collections, and altogether we have
located seven specimens (all males) that we consider to be original syntypes and have listed

46 C. W. SABROSKY & R. W. CROSSKEY

above ; these include the specimen from Toa Tsui Kutsu which we consider to be one of
' einige Exemplare ', and have selected as lectotype.

It is important to note that hemimacquartioides type-material is mixed, and consists of two
very closely similar species : in one species the male head is without proclinate orbital setae
and the scutellum has small upwardly-directed crossed apical setae, and in the second species
the male head has two pairs of proclinate orbital setae and the scutellum lacks apical setae.
The latter species, because of the proclinate orbital setae, has the male head extremely like
that of the normal female Tachinid in which these setae are present, and we think that
Baranov cited ' $ ' as well as ' £ ' at the head of the description of hemimacquartioides in
error (as we have found no female specimens among the syn types). In the original descrip-
tion Baranov wrote of the scutellum that ' Die gekreuzten, aufgerichteten Apikalborsten
sind fein ', and we have therefore selected the lectotype from specimens showing this charac-
ter (i.e. from the species in which the male lacks the proclinate orbital setae). The specimens
in which the male possesses proclinate orbital setae are both from the Tainan locality cited
in the original description ; one is in the DEI collection and the other in USNM (see above) .

The lectotype lacks the genitalia and the slide preparation has not been located. How-
ever, it may be noted that the conspecific male paralectotypes from Takao in the USNM
and BMNH collections are both intact, and that although both males from Takao in the
IZPAN collection at Warsaw are without genitalia there is a slide preparation in that collec-
tion that must associate with one of the two specimens.

Kosempomyiella rufiventris Baranov, I934/ : J^5- LECTOTYPE <J, FORMOSA : [no
other locality data] (H. Sauter) (DEI). Genitalia in situ.

Paralectotypes : 2 $, same data as lectotype, but with month date ' i ' (USNM). 4 $,
FORMOSA : Kosempo, v.igi2 (H. Sauter) (DEI, two ; BMNH ; USNM).

Baranov's original description reads as though all syntypes (number unstated) of both
sexes were from Kosempo, but the specimen bearing his red handwritten ' Typus ' label
(here designated as lectotype) has no locality data other than ' Formosa I. ' ; only the female
syntypes have the locality Kosempo on the data labels. Since there is more than one male,
and none from Kosempo that we have been able to locate, Townsend's (1938 : 41) citation
of ' Ht male ' does not provide a valid fixation of a lectotype.

Leiosiopsis maculibasis Baranov, 19350 : 553. Holotype 6*. JAPAN : Hokkaido, Sapporo

(S. Takano) (USNM). Genitalia in situ.

Leverella institutiimperialis Baranov, 19346 : 474. Lectotype $, by fixation of Townsend
(1939 : 43), SOLOMON ISLANDS : Guadalcanal, Doma, iii.i933 (R. J. A. W. Lever) (BMNH).
Genitalia on slide.

Paralectotype : i $, same data as lectotype (and mounted on same pin) (BMNH).

The original material consists of one male and one female syntype, of which Townsend
(1939 : 43) cited the male as ' Ht ' (= holotype) and the female as ' At ' (— allotype).
Townsend's action restricts the name to a single primary type, and we accept it as providing
valid fixation of a lectotype.

Leverella novaeguineae Baranov, 1934^ : 474- Holotype $, INDONESIAN NEW GUINEA
(WEST IRIAN) : Fakfak (USNM). Genitalia on slide.

Macrozenillia townsendi Baranov, 19350 : 553. Holotype <$, FORMOSA : Sokutsu, ix.igia
(H. Sauter) (DEI).

Paratype : i <$, JAPAN : Hokkaido, Hattaribetsu, 2y.vii.i924 (S. Takano) (USNM).
The holotype lacks the genitalia and the slide preparation has not been located.

Masicera oculata Baranov, 19350 : 554. Holotype $, FORMOSA : Koshun, Kankau, y.vii.
1912 (H. Sauter) (DEI).

Paratypes : 2 <$, same data as lectotype, except dates vii. and 7.viii.i9i2 (USNM). i $,
FORMOSA : Shinkwa, 3. vii. 1926 (S. Takano) (USNM).

TYPE-MATERIAL OF TACHINIDAE 47

In the original publication there is definite reference only to specimens of oculata from
Formosa, but Baranov added that the species was known to him from Java and South
India (' Auch von Java mir bekannt aus Baoris bada Moore geziichtet und aus Siidindien
aus Plusia sp. gezogen '). We have not located any specimens from these additional localities
that could have been seen by Baranov, but we consider that they would in any case be
without type status (as there is no evidence that they were available to Baranov at the time
of description).

Meigenia tnutabilis nobilis Baranov, 19266 : 168. LECTOTYPE <$, JUGOSLAVIA : Serbia,
Golubac, 3o.iv[publ. as vi].ig25 (USNM). Genitalia on slide.

The original publication shows no evidence of how many specimens Baranov saw at the
time of description, but the single male of nobilis in Baranov's collection conforming to the
original data bears a red Baranov label reading ' Typi ' ; we deduce from this that he had
more than one original specimen, and accordingly designate the available specimen as lectotype.

Meigenia tnutabilis pilosa Baranov, 19266 : 168. Holotype <$, JUGOSLAVIA : Serbia,
Golubac, 24. vi. 1925 (USNM). Genitalia on slide.

There is no evidence that the original type-material of subspecies pilosa consisted of more
than one specimen, and the male in the USNM collection having the data as published by
Baranov is considered to be the holotype.

Meigenia mutabilis vulgaris Baranov, 19266 : 168. LECTOTYPE <$, JUGOSLAVIA : Serbia,
Golubac, 30. iv. 1925 (USNM). Genitalia on slide.

Paralectotype : i $, same data as lectotype, except date 12. vi. 1925 (USNM).

We have not located any syntype with the data ' Topfiider, 9^.1924 ' that Baranov recorded
in the original publication.

Micropalpus vulpinoides Baranov, 1932^ : 2. LECTOTYPE <$, SUMATRA : Deli, Siriaria
2.viii.i928 (/. C. v. d. Meer Mohr) (MZ, Bogor). Genitalia on slide.
Paralectotype : i <^, same data as lectotype (USNM).

Monoleptophaga caldwelli Baranov, 19386 : 412. LECTOTYPE 3, AUSTRALIA : Queens-
land, Nambour [70 mi. North of Brisbane], x.i937, ex Monolepta rosea adult (N. E. H.
Caldwell) (BMNH). Genitalia in situ.

Paralectotypes ; 2 9, same data as lectotype (BMNH & USNM). 4 <$, 4 $, same data as
lectotype (Dept. of Primary Industries Coll., Brisbane, Queensland).

Myiobia bezziana Baranov, 19386 : 411. LECTOTYPE <$, INDIA : Bengal, Darjeeling,
5000 ft., 1923, (/. C. M. Gardner] (BMNH). Genitalia in situ.

Paralectotypes : 2 $, same data, except date 7^1.1923 (BMNH) ; i $, same data as
lectotype, except host Zeuzera multistrigata given (USNM) ; i $, same data, except date
ii.i923 (USNM).

Baranov did not state the number or sex of his original specimens, but gave the date as
i6.viii.i923- We consider all the above-listed specimens to be original syntypes, but it
should be noted that none of them bears the day and month date ' i6.viii ' cited in the
original publication.

Beeson & Chatterjee (1935 : 177) published this name before Baranov's description, but
gave only four lines on the life history and no description ; the name is nomenclaturally
unavailable from Beeson & Chatterjee (1935) under Article i3a (i) of the International Code of
Zoological Nomenclature, 1961.

Myiofijia bezziana Baranov, I934C : 478. LECTOTYPE <$, FIJI : Taveuni, ii.i934- bred
from host moth larva in coconut tree (R. W. Paine) (BMNH). Genitalia in situ.

Paralectotypes : i $, i ?, same data as lectotype (USNM) . i $, same data as lectotype,
except date 1.1934 and ' bred from host cocoon in coconut ' (USNM). i $, FIJI : Ura,
Taveuni, 1.1934 (R- w- Paine) (BMNH) ; i $, same data, except date xi.i933 (BMNH).
i ?, FIJI : Nabokoyia, Taveuni, iii.i934, ex pyralid cocoon on coconut tree (R. W. Paine)
(BMNH).

48 C. W. SABROSKY & R. W. CROSSKEY

Townsend (1941 : 291), for M. bezziana, cited ' Ht ' (= holotype) male and ' At ' ( =
allotype) female in London without further information ; since there are at least two male
original syn types Townsend 's statement does not restrict the name to a single specimen and
is not a valid lectotype fixation. A lectotype is therefore here newly designated.

Myiostoma magna Baranov, 193513 : 557. Holotype $, JAPAN : Hokkaido, Sapporo,
7-ix.i923 (S. Takano) (USNM).

Parexorista latistylata Baranov, 19340" : 405. Holotype $, FORMOSA : [Formosan locality
unknown] i. (H. Sauter) (USNM). Genitalia on slide.

The holotype bears Baranov's label reading ' Carcelia latistylata n.sp. N. Baranoff TYPUS '.

Phorocera imperator Baranov, 1936 : 109. Holotype <$, CELEBES : S. Celebes, Samanga,
xi.i895 (H. Frilhstorfer) (BMNH, ex coll. Wainwright). Genitalia in situ.

Phorocera isabeli Baranov, 19380 : 171. Holotype $, SOLOMON ISLANDS : Isabel, Tatamba,
n.vii [publ. as vi].i935 (R. /. A. W. Lever) (BMNH). Genitalia in situ.

Phorocera magna Baranov, 19340 : 46. LECTOTYPE $, INDONESIA : Moluccas, Batjan,
viii.1929 (W. Roepke) (USNM : genitalia slide only).

This species was described from material obtained by W. Roepke in July and August,
1929, while investigating the parasites of Thosea moluccana Roepke, 1935 (Lepidoptera :
Limacodidae), a serious pest of coconut in Batjan island, Moluccas. The description of
Phorocera magna by Baranov (19356) was published as an unpaginated appendix following
the last page (p. 38) of Roepke's (1935) account of his work in Batjan, and was based upon
four syntypes (2 <$, 2 $) with the data cited by Baranov as ' Molukkeninsel Batjan, vii, 1929
(W. Roepke) '; this description, which Baranov intended to be the original description, had
however already appeared in print in an earlier paper (Baranov, 19340) while Roepke's work
was still in press. The name Phorocera magna is therefore nomenclaturally available from
the 1934 paper here cited.

Baranov (19340), as we^ as repeating the description, mentioned two small specimens of
P. magna from Ceylon, but we do not accept these specimens (which, in any case, we have
been unable to locate) as original syntypes because the description was made only from the
Moluccan material.

Unfortunately we have been unable to locate any of the four original syntypes, but a slide
preparation of the male genitalia of one of them is in USNM (ex Baranov collection) and this
we have fixed as lectotype. However, there are four specimens of P. magna in the Museum
Zoologicum Bogoriense and four specimens in the Central Agricultural Research Institute,
Bogor, which were all collected by Roepke in Batjan and reared from Thosea moluccana ;
none of these other eight topotypic specimens existing in Bogor have been labelled by
Baranov and all have the date ' xii.i92g ' (the original material was collected in July or
August), and there is no evidence that any of them are syntypes. Furthermore, each of them
has a locality label reading ' Penamboean, Batjan ', and it is unlikely that Baranov would
have omitted the first of these words in his unusually full citation of data for P. magna if
some of these specimens were his original material. Thus we are convinced that none of the
specimens now in Bogor represent type-material. It is especially surprising that Baranov's
own collection at the U.S. National Museum does not contain at least one complete syntype,
as Baranov normally retained part of his type-series if this consisted of several specimens.
Finally, we should record that Baranov (19340) cited the month date of the original syntypes
as ' VIII ', but in the later description published (Baranov, 19356) he gave it as ' VII '. In
the absence of syntypes we are unable to say for certain which is correct, but the lectotype
genitalia slide has the August month date and we accept this as correct. The month date
' xii ' on specimens in Bogor might also possibly be in error for ' vii ' or ' viii ', but we have
no evidence on this.

The specimens of P. magna in the Museum Zoologicum Bogoriense are three females and
one male, but we do not know how many of each sex are present among the four specimens
in the Central Agricultural Research Institute, Bogor.

TYPE-MATERIAL OF TACHINIDAE 49

Phorocera magna form maxima Baranov, 1936 : 105. LECTOTYPE $, FORMOSA :
Sokutsu, ix.i9i2 (H. Sauter) (USNM). Genitalia in situ.

Paralectotypes : i <$, same data as lectotype (USNM). i <$ (genitalia slide only), same
data as lectotype (USNM).

Phorocerosoma anomala Baranov, 1936 : 99. Lectotype $, by designation of Crosskey
(1966 : 108), FORMOSA : Koshun, Kankau, y.viii.igia (H. Sauter) (DEI).

Paralectotypes : i $, same data as lectotype, except date 22. vi. 1912 (USNM). 3 $, same
data as lectotype, except date vi.i9i2 (DEI), i 9, same data as lectotype, except date
ix.i9i2 (DEI), i $, same data as lectotype, except date y.ix.i9i2 (DEI), i $, 2 $, FORMOSA:
Toa Tsui Kutsu, v.i9i4 (H. Sauter) (J ? USNM, ? DEI).

The paralectotype series is mixed and consists of two species of Phorocerosoma Townsend
(see Crosskey, 1966 : 108-109). The two females in the DEI collection with the data ' Toa
Tsui Kutsu ' and ' Kankau ix.i9i2 ' are actually specimens of Phorocerosoma vicarium
(Walker) and are not conspecific with the lectotype of anomala and most of the paralecto-
types (which have four strong marginal setae on the third abdominal tergite and are actually
specimens of Phorocerosoma postulans (Walker), the senior synonym of anomala). The
lectotype and all paralectotypes of anomala bear the name ' Exorista anomala ' in Baranov's
writing.

Baranov (1936 : 99) makes it clear that the description of anomala is based solely on
Formosan specimens, although he cited the data (' i $, Ugi Is., 6.V.I934, leg. R. A. Lever ')
of a specimen from the Solomon Islands. This specimen is in the BMNH collection and
was cited by Crosskey (1966 : 108) as a syntype of anomala : we now consider, however,
that it has no type-status.

At present the name anomala is in synonymy with postulans, but it should be noted that if
again treated as valid in combination with Phorocerosoma the correct spelling will be anomalum
(under Article 34(b) of the International Code of Zoological Nomenclature, 1961).

Plagioderophagus niger Baranov, 19386 : 412. LECTOTYPE <$, INDIA : U.P., Dehra
Dun, 22. iv. 1930, ex larva of Plagiodera rufescens defoliating] Flacourtia Ramnotchi (S. N.
Chatter jee) (BMNH). Genitalia in situ.

Paralectotypes : i <$, 2 ?, same data as lectotype (<£ $ USNM, $ BMNH).

Beeson & Chatterjee (1935 : 177) had already published this name, prior to Baranov's
description, with a note on the life-history, but they gave no description and the name is
therefore nomenclaturally unavailable from Beeson & Chatterjee (1935) under Article I3a
(i) of the International Code of Zoological Nomenclature, 1961.

Platerycia compressa Baranov, 1936 : in. LECTOTYPE 6*, FORMOSA : Tainan, v.i9i2

(H. Sauter) (DEI). Genitalia on slide.

Paralectotypes : i 6*. i ?, same data as lectotype (USNM). i 6*. FORMOSA : Koshun,

Kankau, 7.vii.i9i2 (H. Sauter) (DEI), i ?, FORMOSA : Hoozan, 7.^.1911 (H. Sauter) (DEI).
Prosopodes leveri Baranov, 19386 : 410. Holotype £, SOLOMON ISLANDS : Russell Island,

Karamola, 2o.ix.i934 (R. ]. A. W. Lever) (BMNH).

Protonemoraea japanica Baranov, 19350 : 556. Holotype <$, JAPAN : Hokkaido, Sapporo,
10. ix. 1923 (S. Takano) (USNM). Genitalia on slide.

The USNM collection also contains the female specimen noted by Baranov in the original
publication as being probably conspecific with the holotype. Its data are : JAPAN :
Hokkaido, Kamuikotan, 5.viii.i93o (S. Takano).

Protonemoraea takanoi Baranov, 19350 : 557. Holotype ?, JAPAN : Hokkaido, Sapporo,
I5.viii.ig24 (S. Takano) (USNM).

Stomatomyia bezziana Baranov, 19340 : 48. Lectotype $, by designation of Crosskey
(19660 : 673), CEYLON : Batticaloa, 13.^.1922, ex Nephantis serinopa (J. C. Hutson)
(BMNH). Genitalia on slide.

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

50 C. W. SABROSKY & R. W. CROSSKEY

Sturmia hell a oceanica Baranov, 19380 : 170. Holotype ?, SOLOMON ISLANDS : San
Cristobal, Waiai, 8 [publ. as S].v.i935 (R. J. A. W. Lever) (BMNH).

Sturmia bisetosa Baranov, 19326 : 75. Holotype <$, FORMOSA : Sokutsu, ix.igia (H.
Sauter) (DEI).

The holotype lacks the genitalia and the slide preparation has not been located.

Sturmia chatterjeeana Baranov, 1934^ : 484- Holotype $, INDIA : U.P., Dehra Dun
17. iv. 1934, parasitic on Euproctis bipunctapex (S. N. Chatterjee) (BMNH). Genitalia in situ.
The BMNH collection contains two male specimens with same data as holotype except
for date i8.vi.i934 and three female specimens with same data as holotype except for date
2o.iv.i934 ; the USNM collection contains a male with same data as the holotype and a
male with date i8.iv.i934 '> they bear Baranov's determination labels as chatterjeeana but
they are not type-material.

Sturmia hutsoni Baranov, 19340 : 42- LECTOTYPE $, CEYLON : Mawanella, 7.^.1928,
larval parasite of Earias fabia feeding on Hibiscus esculentus (J. C. Hutsori) (BMNH).
Genitalia in situ.

Paralectotypes : 3 <$, same data as lectotype (2 o* BMNH, i <$ USNM).

One of the paralectotypes in BMNH lacks the abdomen.

Sturmia inconspicuella Baranov, 19326 : 79. Lectotype <$, by designation of Crosskey

19676 : 57, 59), FORMOSA : Koshun, Kankau, viii.i9i2 (H. Sauter) (DEI). Genitalia in situ.

Paralectotypes : 25 <$, 6 $, same data as lectotype, except several dates from iv.-xi.i9i2

(21 6*, 5 ? DEI ; i o* BMNH ; i ? USNM). i 6", FORMOSA : Sokutsu, ix.i9i2 (H. Sauter)

(DEI).

There is also one male paralectotype in the collection of the Department of Agriculture,
Pakistan, but the data are not known to us.

Sturmia inconspicuoid.es Baranov, 19326 : 80. Lectotype $, by designation of Crosskey
(19676 : 50, 51), FORMOSA : Koshun, Kankau, 7.viii.i9i2 (H. Sauter) (DEI).

Paralectotypes : i <$, 3 $, same data as lectotype (DEI). 2 $, same data as lectotype,
except date ix.i9i2 (DEI & USNM). 5 <$, FORMOSA : Sokutsu, ix.i9i2 (H. Sauter} (2 <J
DEI ; 2 c? USNM ; i <J BMNH).

The lectotype lacks the genitalia and the slide preparation has not been located. Two
of the paralectotypes (?) in DEI collection with the same data as the lectotype and the female
in DEI with data ' Kankau ix.igi2 ' are believed to be misidentified and not conspecific with
the lectotype (see Crosskey, 19676 : 51) ; it is probable also that the female specimen in
USNM with the same data ' Kankau ix.i9i2 ' is likewise not conspecific with the lectotype.

Sturmia latiforceps Baranov, 19326 : 78. Lectotype <$, by designation of Crosskey (19676 :
72-73), FORMOSA : Koshun, Kankau, 7.viii.i9i2 (H. Sauter} (DEI). Genitalia in situ.

Paralectotypes : 6 Q*. same data as lectotype (4 Q* DEI ; i <$ USNM ; i $ coll. Mesnil,
Delemont). i <$, same data as lectotype, except date ix.i9i2 (BMNH). 2 Q", same data as
lectotype, except date iv.i9i2 (DEI & USNM). i $, same data as lectotype, except date
v.1912 (DEI). 2 o*> FORMOSA : Sokutsu, ix.i9i2 (H. Sauter} (DEI).

In the original publication Baranov mentioned thirteen males from Kankau and Sokutsu ;
all have been located and are accounted for above.

Sturmia latistylata Baranov, 19326 : 79. LECTOTYPE <J, FORMOSA : Koshun, Kankau,
7.ix.igi2 (H. Sauter) (DEI). Genitalia in situ.

Paralectotypes : 3 <$, i $, same data as lectotype, except no day date (DEI). 2 ^, i $,
same data as lectotype, except date 7.viii.igi2 (o* DEI, $ $ USNM). i <$, same data as
lectotype, except date vii.igi2 (USNM). 4 $, FORMOSA : Kanshizei, v.igoS (H. Sauter}
(z 6* DEI, 2 (J BMNH). i <J, FORMOSA : Sokutsu, ix.i9i2 (H. Sauter} (DEL).

TYPE-MATERIAL OF TACHINIDAE 51

In the original publication Baranov mentioned fourteen specimens (n <J, 3 $) ; we have
located this total, but 12 <J and 2 $. One of the males in USNM collection has a pair of
proclinate orbital setae and may have been mistaken for a female, thus accounting for the
discrepancy. The female paralectotype from Kankau in DEI collection, and one of the
males from Kanshizei in the same collection, have lost the abdomen. The male paralecto-
type from Sokutsu is mis-associated with the lectotype, and is a specimen of Zygobothria
atropivora (Robineau-Desvoidy) ; the male paralectotype in DEI with date 7.viii.i9i2 from
Kankau is also mis-associated and is a specimen of a Sisyropa species.

Sturmia macrophallus Baranov, 19326 : 76. Lectotype $, by designation of Crosskey
(19676 : 105), FORMOSA : Koshun, Kankau, ix.igi2 (H. Sauter) (DEI). Genitalia on slide.

Paralectotypes : i £, same data as lectotype (USNM). i $ FORMOSA : Tainan, v.igi2
(H. Sauter) (DEI.)

All specimens mentioned by Baranov in the original publication have been located and are
listed above. In addition the Baranov collection in USNM has another male with data
identical to those of the lectotype, but it is not mentioned as part of the original syntype
series and is not therefore a paralectotype.

Sturmia nigribarbis Baranov, i934a : 42. LECTOTYPE $, BURMA : Upper Thaungyin,
Moulmein, I7.viii.i93i, parasitic on Hapalia machaeralis pupa (D. J. Atkinson) (BMNH).
Genitalia in situ.

Paralectotypes : 2^,2$, same data as lectotype (<£<£ USNM, $$ BMNH) . 3 $, same data
as lectotype, except date I4.viii.i93i (i in BMNH, 2 in USNM).

Sturmia oculata Baranov, 19326 : 80. Holotype <$, FORMOSA : Tainan, v.igi2 (H. Sauter)
(location not traced, possibly lost).

The holotype of oculata should be in the DEI collection, and was recorded as present in
that collection by Hennig (1941 : 199). It is now missing from DEI collection and has not
been located.

Sturmia painei Baranov, i934a : 42. Lectotype <£, by designation of Crosskey (1967^ :
81-82), JAVA : [Javanese locality unknown] 1929-30, ex Tirathaba sp. (R. W. Paine)
(BMNH). Genitalia in situ.

Paralectotypes : 2 <$, 3 $, same data as lectotype (i £, 2 $ BMNH ; i Q*, i $ USNM). i <$
(genitalia slide only), same data as lectotype (USNM).

Sturmia paradoxalis Baranov, 19326 : 80. Holotype <£, FORMOSA : Sokutsu, ix.igi2 (H.
Sauter) (DEI). Genitalia on slide.

Mesnil's (1951 : 200) citation of the month date of the holotype as ' November ' is in error.

Sturmia picta Baranov, 19326 : 77. LECTOTYPE £, FORMOSA : Koshun, Kankau, 7.viii.
igi2 (H. Sauter) (DEI). Genitalia in situ.

Paralectotypes : 2 <$, same data as lectotype (DEI & USNM). i $, i 9, same data as
lectotype, except no day date (<$ DEI, $ USNM). 3 ^, i $, same data as lectotype, except
date vi.i9i2 (2 $ DEI ; i <J, i $ USNM). i <J, same data as lectotype, except date ix.igi2
(BMNH).

Hennig (1941 : 199) recorded 16 ' Typen ' in DEI collection, but some of these have not
been traced.

Sturmia sumatrana Baranov, 1932^ : i. Holotype $, SUMATRA : N.O. Sumatra, Medan,
iv.i928 (/. C.v.d. Meer Mohr) (USNM).

Sturmia trisetosa Baranov, 19326 : 78. Lectotype (J, by designation of Crosskey (19676 :
105), FORMOSA : Koshun, Kankau, vii.igi2 (H. Sauter) (DEI). Genitalia on slide.

52 C. W. SABROSKY & R. W. CROSSKEY

Paralectotypes : i $, same data as lectotype (USNM). i ^, 2 $, same data as lectotype,
except date j.vm.igi2 (<$ BMNH, 2 $ DEI), i <£, same data as lectotype, except date
viii.i9i2 (USNM). i <£, 3 $, same data as lectotype, except date ix.igia (i <£, 2 $ DEI, i $
BMNH). i <J, FORMOSA : Sokutsu, vi. 1912 (//. Sauter) (DEI), i $, FORMOSA : Taihorinsho,
ix.igi2 (H. Sauter) (DEI).

The paralectotype from Taihorinsho (DEI) and the male paralectotype from Kankau
ix.igi2 in DEI collection have lost the abdomen.

Sturmia trisetosoides Baranov, 19326 : 78. Lectotype <$, by designation of Crosskey
(19676 : 105), FORMOSA : Tainan, iv.igio (H. Sauter) (DEI). Genitalia on slide.

Paralectotype : i <$, same data as lectotype (USNM).

Baranov based the original description on two males and a female, all from Tainan. The
female syntype has not been located. The Baranov collection in USNM contains four males
that have the same data as the lectotype and are all labelled by Baranov as ' n.sp. N.
Baranoff ' in his usual style of label. Not all of these can be regarded as syntypes, since the
original description mentions only two males (i.e. one in addition to the lectotype) ; one of
them, however, has had the hypopygium extracted and it is considered that this one is the
second specimen of the original pair (it is therefore listed as male paralectotype above, and
has been labelled as such).

Sturmia unguicularis Baranov, 19340 : 480. Holotype <J, JAVA : Ngantang (Frau Neuhaus)
(USNM).

The holotype lacks the genitalia and the slide preparation has not been located. In the
original description Baranov mentioned a second specimen from New Caledonia which he
expressly excluded from the type-material by the following statement : ' Vorliegende
Beschreibung bezieht sich auf das javanische Exemplar und das neukaledonische Exemplar,
welches in ziemlich schlechten Zustande ist, betrachte ich nur als Idiotype und nicht als
Cotype '. This specimen, in poor condition as Baranov said, is in the BMNH collection
and has the following data : <£, NEW CALEDONIA : Noumea, 17^.1928 (T. D. A. Cockerell).

Sturmia unisetosa Baranov, 19326 : 75. Lectotype <£, by designation of Crosskey (19670 :
68-69), FORMOSA : Koshun, Kankau, 7.viii.i9i2 (H. Sauter) (DEI).

Paralectotypes : i $, same data as lectotype, except date 7.ix.igi2 (DEI), i $, same
data as lectotype, except date ix.igi2 (USNM).

The lectotype lacks the genitalia and the slide preparation has not been located.

Sturmia vicinella Baranov, 19326 : 79. Holotype $, FORMOSA : Tainan, iv.igio (H. Sauter)
(DEI).

The holotype lacks the genitalia and the slide preparation has not been located.

Sturmia wainwrighti Baranov, 19327 : 100. Holotype <$, INDIA : Assam, Khasia Hills,
2i.iii.i9ii (C. B. Antram) (BMNH, ex coll. Wainwright). Genitalia in situ.

Paratypes : 7 $, 8 $, same data as holotype (BMNH). i $, same data as holotype (USNM).
3 c?, 2 $, same data as holotype, except date 2o.iii.igi i (BMNH). i $, same data as holotype,
except date 2o.iii.igii (USNM). i $, same data as holotype, except date 23.iii.ign (BMNH).
i <$, same data as holotype, except date n.iii.igio (BMNH).

Not all of the dates of the paratype material were recorded by Baranov in the original
publication, but we are nevertheless satisfied that all the specimens (other than the holotype)
listed above are acceptable as paratypes.

Sumatrodexia incisuralis Baranov, 19320 : 215. Holotype^, CHINA : Szechwan, Tatsienlu
(Exp. Stotzder) (Staatliches Museum fur Tierkunde, Dresden). Genitalia on slide.

Sumatrodexia montana Baranov, 19326 : 215. LECTOTYPE <$, JAVA : Tjibodas, 1400 m,
xii.i927 (USNM). Genitalia in situ.

Paralectotypes : 3 <$, same data as lectotype (2 in MZ, Bogor & one in BMNH).

Both paralectotypes in the Bogor collection lack the genitalia, but an associated slide
preparation from one of them is present in the same collection.

TYPE-MATERIAL OF TACHINIDAE

53

Sumatrodexia vittata Baranov, 19320 : 215. Holotype $, JAVA : Surabaja (USNM).
Genitalia in situ.

Takanoella parvicornis Baranov, i935a : 559. Holotype $, JAPAN : Hokkaido, Sapporo,
28. iv. 1924 (S. Takano) (USNM). Genitalia in situ.

Tamanukia japanica Baranov, 19350 : 551. Holotype $, JAPAN : Hokkaido, Obihiro,
3o.viii.i924 (S. Tamanuki) (USNM). Genitalia on card mount below locality label.

Trichoformosomyia sauteri Baranov, I934/ : l64- LECTOTYPE $, FORMOSA : Formosa
I. (H. Sauter) (DEI). Genitalia in situ.

Paralectotypes : i <$, FORMOSA : Sokutsu, vi.igi2 (BMNH). i <$, FORMOSA : Suisharya,
X.IQII (USNM).

Baranov did not state the number of male specimens on which the description was based,
but Townsend (1939 : 145) assumed that the specimen in DEI collection was the ' Ht ' ( =
holotype). We do not accept this as a valid lectotype fixation because there are other male
syntypes with Baranov's ' n.g., n.sp. ' labels in other collections, and we therefore here fix
the lectotype of T. sauteri by present designation.

Tricholyga psychidarum Baranov, 19340 : 47. Holotype $, SUMATRA : Pematang Siantar,
Naga Hoeta Estate, 1750 ft., io.x.i93i, ex larva of bagworm (R. I . Nel) (BMNH). Genitalia
on slide.

Paratypes : 2 $, same data as holotype (USNM). i <$, same data as holotype, except
date given as 6-io.x.i93i (BMNH). i °-, same data as holotype, except date 14. x. 1931
(USNM). i $, SUMATRA : Pematang Siantar, Simpang Raja Estate, 2200 ft., 17.1.1932
(R. I. Nel) (BMNH) ; i $, Pematang Siantar, Bah Kapoel Estate, 1500 ft., 2.1.1932, ex
larva of bagworm (USNM). i $, SUMATRA : Pematang Siantar, Mardjandi Estate, 2500 ft.,
24.1.1932, ex larva of bagworm (R. I. Nel) (BMNH).

In the original description Baranov characterized psychidarum as having hairy eyes, but
mentioned that most specimens of the type-material had the hairing rubbed off (' bei den
meisten Exemplaren Behaarung abgerieben '). From our own examination of the holotype
and paratype material we are sure that two separate species are involved, one having the eyes
virtually bare in the natural state and the other having the eyes conspicuously long haired,
for the eye difference is associated with quite different forms of male genitalia ; other material
of the complex now available also confirms this conclusion. The holotype has bare eyes, and
the name psych-idarum therefore applies to the species with this character. Most of the
above-listed paratypes have bare eyes and are conspecific with the holotype, but two of the
female paratypes are hairy-eyed and mis-associated : these are the USNM specimen with
date 2.1.1932 and the BMNH specimen from Mardjandi Estate, 24.1.1932. No applicable
name is known to us for the hairy-eyed species, which appears at present to be undescribed.
The BMNH collection contains a hairy-eyed male with data ' Sumatra, Naga Hoeta Est.,
P. Siantar, 8.xii.3o R. I. Nel ' but it bears a determination label in Baranov's writing as
' Eutachina psychidarum Baranoff ' and is evidently not an original paratype. The same
collection also has a hairy-eyed female with data ' Malaya, Chebiot Estate, Labu, 28.9.1928 ',
but is also labelled by Baranov as ' Eutachina psychidarum Bar. ' and is presumed not to be
the paratype mentioned in the original publication from ' Malaya, 28. ix. 1928 ', which has
not been seen.

Vibrissina hokkaidensis Baranov, 19350 : 554. LECTOTYPE $, JAPAN : Hokkaido,
Sapporo, 19 [publ. as 12]. x. 1923 (S. Takano) (USNM). Genitalia in situ.
Paralectotype : i 9, same data as lectotype (USNM).

Voria edentata Baranov, 19320 : 83. Holotype <$, FORMOSA : Koshun, Kankau, 22. vi. 1912
(H. Sauter) (not located and possibly lost : formerly in DEI).

Baranov described this species only from the single specimen recorded above and cited the
DEI collection as the type depository. The holotype was later seen and recorded by Hennig
(1941 : 192), who also cited the locality and date as ' Kankau, VI ' for the type and men-
tioned three other specimens in the DEI collection (another specimen from Kankau and two

54 C. W. SABROSKY & R. W. CROSSKEY

just with the data ' Formosa '). We have been unable during the preparation of the present
work to locate the holotype specimen and consider that it is possibly lost ; furthermore, the
DEI collection now contains only two of the three other specimens recorded by Hennig (viz.
a male with data ' Kankau y.vii.igia H. Sauter ' and female with data ' Formosa I Sauter ',
both determined by Baranov as Voria edentatd).

In a revision of Old World Voriini the data of the ' type ' of edentata has been cited by
d'Aguilar (1957 : 2^2) as ' 1 '^e de Taiwan : Kankau, recolte en juillet-aout 1912, par H.
Sauter ', but Baranov clearly cited the holotype date as ' 22. vi. 1912 ' in the original publica-
tion and Hennig also found the month date to be June. Thus probably d'Aguilar did not
see the true holotype from DEI, but may have seen the later determined specimen from
Kankau in DEI collection (recorded above), although this specimen bears no month date
August. Dr. d'Aguilar (personal communication) has kindly confirmed that the specimen
seen by him was returned to DEI, and Dr. L. P. Mesnil has confirmed that the holotype is
not on loan to him at Delemont.

It should be noted that the slide preparation of the male genitalia from the specimen of
V. edentata with the date ' Kankau y.vii.1912 ' is also in the DEI collection, but not the
slide preparation from the holotype.

Winthemia diver sa solomonica Baranov, 19386 : 405. Holotype £, SOLOMON ISLANDS :
Tulagi, 4-ix.i934, ex psychid cocoon (R. J. A. W. Lever] (BMNH). Genitalia in situ.

Winthemia diversoides Baranov, 19320 : 47. Holotype <J, FORMOSA : Sokutsu, ix.igiz
(H. Sauter} (DEI). Genitalia on slide.

At the time of writing the holotype is temporarily in the collection of Dr. L. P. Mesnil,
at Delemont, Switzerland.

Winthemia hokkaidensis Baranov, 1939 : no. Holotype $, JAPAN : Hokkaido, Sapporo,
10. ix. 1923 [publ. as 1929] (S. Takano) (USNM).

Winthemia mallochi Baranov, 19320 : 46. Holotype <$, FORMOSA : Koshun, Kankau,
vi.i9i2 (H. Sauter) (DEI). Genitalia on slide.

At the time of writing the holotype is temporarily in the collection of Dr. L. P. Mesnil, at
Delemont, Switzerland.

Zenillia caldwelli Baranov, 19386 : 409. Holotype $, AUSTRALIAN NEW GUINEA : Admiralty
Islands, Manus, 1932 (N. E. H. Caldwell) (BMNH).

Zenillia roseanella Baranov, 1936 : 104. LECTOTYPE <$, FORMOSA : Sokutsu, ix.i9i2
(H. Sauter) (USNM). Genitalia on slide.

Paralectotypes : none traced.

In the original publication Baranov recorded a specimen from New Britain that is now in
the BMNH collection and has the following data : $ (without abdomen) NEW BRITAIN :
Rabaul, 29.1.1934, ex lepidopterous larva (/. L. Froggatt). This specimen, however, has only
a Baranov determination label as ' Zenillia roseanella Baranoff ', and it is not a syntype
because Baranov expressly stated that the description was based on ' Exemplare ' from
Formosa which he was publishing in the 1936 paper (' hier an dieser Stelle ') as it had not yet
appeared elsewhere. Thus the name is based on syntypes from Formosa, of which only one
(here fixed as lectotype) has been located.

PART II.— BARANOV'S MANUSCRIPT NAMES THAT HAVE BEEN PUBLISHED.

We bring together in the following alphabetical list all those manuscript names of Baranov
that have appeared in print, either as nomina nuda or as species-group names made available
by the action of later authors. The available species-group names are shown in bold type.

Actia mallochiana Gardner, 1940 : 178 (Actia mallochiana Baranov MS). Available name,
attributable to Gardner.

TYPE-MATERIAL OF TACHINIDAE 55

Gardner described the puparium of a species of Tachinid fly under Baranov's hitherto
unpublished manuscript name Actia mallochiana. Under Article 10 of the International
Code of Zoological Nomenclature, 1961, the name mallochiana is made available by, and must
be attributed to, Gardner.

Allophora dubiosa Baranov in Hennig, 1941 : 187. Nomen nudum.

Hennig listed without description (as ' Phasia dubiosa Baranoff ') a specimen in DEI
collection from Formosa that bears a Baranov label as ' Allophora dubiosa n.sp. N. Baranoff ' ;
the name was not published by Baranov and remains a nomen nudum.

Argyrophylax rufitibialis Baranov in Hennig, 1941 : 196. Nomen nudum.

Hennig listed, under this name and without description, a specimen from Hoozan in
Formosa in DEI collection ; the name was not published by Baranov and remains a nomen
nudum. Mesnil (1944 : 27) also cited the name, but did not validate it by description.

Bactromyia compsiluroides Baranov in Baranov, 19386 : 409. Nomen nudum.

Baranov mentions compsiluroides as a Formosan species in his original description of
Bactromyia crassiseta Baranov, 1938, but without characterizing it, and the name remains a
nomen nudum.

Cossidophaga kalshoveni Baranov in Baranov, 1934/1 161. Manuscript name cited in
synonymy, unavailable.

This name was cited by Baranov in synonymy under Cossidophaga atkinsoni (Aubertin), a
species described by Aubertin (1932) as Podomyia atkinsoni. Baranov (19347 : I^i), when
describing the new genus Cossidophaga for atkinsoni, abandoned his intended specific name
kalshoveni for this species but published the latter name in synonymy ; it is unavailable
under Article n (d) of the International Code of Zoological Nomenclature, 1961.

Crocuta taiwanica Baranov in Hennig, 1941 : 195. Nomen nudum.

Hennig listed without description (as ' Siphona taiwanica Baranoff ') four specimens in the
DEI collection from Macuyama and Toa Tsui Kutsu in Formosa ; the name was not published
by Baranov and remains a nomen nudum.

Ctenophorocera sturmioides Mesnil, 1950 : 126 (Prosopaea sturmioides Baranov MS).
Available name, attributable to Mesnil.

Mesnil (1950 : 126) described Ctenophorocera sturmioides as a new species based upon a
male holotype specimen (in DEI collection) labelled by Baranov as ' Prosopaea sturmioides
n.sp. N. Baranoff' and with the data : — Formosa : Sokutsu, vi.igia (H. Sauter). Mesnil
placed the species in his subgenus Parapales Mesnil, and the holotype is labelled (in addition
to Baranov's label) as ' Parapales sturmioides Mesn. ' in Mesnil's writing.

Hennig (1941 : 196) listed the name Prosopaea sturmioides Baranoff but gave no descrip-
tion ; he mentioned three specimens in DEI collection from Formosa, one without further
data, one from Toa Tsui Kutsu, and one from Sokutsu, of which only the last was seen by
Mesnil (i.e. the holotype cited above). It should be noted that the month date of the holo-
type specimen from Sokutsu is ' vi ' as given by Hennig, and not ' v ' as given by Mesnil
(1950 : 127).

Cuphocera varia form formosana Baranov in Baranov, 1936 : 98. Manuscript name cited in
synonymy, unavailable.

This name was cited by Baranov in synonymy under the true Cuphocera varia (Fabricius),
and is unavailable under Article 1 1 (d) of the International Code of Zoological Nomenclature,
1961.

Cuphocera varia form malayana Baranov in Baranov, 1936 : 98. Manuscript name cited in
synonymy, unavailable.

This name was cited by Baranov in synonymy under the true Cuphocera varia (Fabricius),
and is unavailable under Article 1 1 (d) of the International Code of Zoological Nomenclature,
1961.

56 C. W. SABROSKY & R. W. CROSSKEY

Dolicholon ater Gardner, 1940 : 177 (Dolichocolon ater Baranov MS). Available name,
attributable to Gardner.

Gardner described and figured the puparium of a species of Tachinid fly under Baranov's
hitherto unpublished manuscript name Dolichocolon ater. Under Article 10 of the Inter-
national Code of Zoological Nomenclature, 1961, the name ater is made available by, and must
be attributed to, Gardner.

Euhapalivora indica Gardner, 1940 : 179 (Euhapalivora indica Baranov MS). Available
specific name, attributable to Gardner.

Gardner described and figured the puparium of a species of Tachinid fly under Baranov's
hitherto unpublished manuscript name Euhapalivora indica. The specific name indica,
even though published in combination with an unavailable genus-group name, is available
under Article n (g) (ii) of the International Code of Zoological Nomenclature, 1961, and is
attributable to Gardner under Article 10. The generic name Euhapalivora (as mentioned by
Crosskey, 19670: 13) is not accompanied, however, by a definition of the generic taxon and
is a nomen nudum which is unavailable under Article 13 (a).

The species indica has been assigned to the genus Pseudoperichaeta Brauer & Bergenstamm
by Crosskey (19676).

Exorista apicalis Baranov in Hennig, 1941 : 193. Nomen nudum.

Hennig listed without description 20 specimens in DEI collection from Kankau and Sokutsu
in Formosa ; the name was not published by Baranov and remains a nomen nudum. Mesnil
(1950 : 153) cited Exorista apicalis Baranov as a synonym of Sisyropa soror Mesnil, 1944, but
it is nevertheless unavailable from Mesnil under Article n (d) of the International Code of
Zoological Nomenclature, 1961.

Exorista grisellina Gardner, 1940 : 177 (Exorista grisellina Baranov MS). Available name,
attributable to Gardner.

Gardner described and figured the puparium of a species of Tachinid fly under Baranov's
hitherto unpublished manuscript name Exorista grisellina. Under Article 10 of the Inter-
national Code of Zoological Nomenclature, 1961, the name grisellina is made available by, and
must be attributed to, Gardner.

Exorista maculiventris Baranov in Hennig, 1941 : 194. Nomen nudum.

Hennig listed, under this name and without description, two specimens in DEI collection
from Kankau in Formosa ; the name was not published by Baranov and remains a nomen
nudum. Mesnil (1950 : 154) cited Exorista maculiventris Baranov as a synonym of Sisyropa
thermophila (Wiedemann, 1830), but it is nevertheless unavailable from Mesnil under Article
ii (d) of the International Code of Zoological Nomenclature, 1961.

Exorista pulchra Baranov. See Zenilliana pulchra Mesnil.

Exorista simulator Baranov in Hennig, 1941 : 194. Nomen nudum.

Hennig listed, under this name and without description, three specimens in DEI collection
from Formosa, two without further data and the other one from Toa Tsui Kutsu ; the name
was not published by Baranov and remains a nomen nudum. Mesnil (1949 : 66) cited the
name ' simulator Bar. (in litt.) ' as a synonym of Phorocerosoma forte Townsend without
noting the generic name used by Baranov ; it is nevertheless unavailable under Article ii
(d) of the International Code of Zoological Nomenclature, 1961.

Exorista winthemioides Baranov. See Nemosturmia winthemioides Mesnil.

Gymnodexia orientalis Baranov in Baranov, 19340 : 49. Nomen nudum.

Baranov (19340 : 49), in the original description of Gymnodexia atkinsoni, made the
statement : ' Am nachsten zu orientalis mihi (in litteris) ', but nowhere published a descrip-
tion of orientalis. Hennig (1941 : 191), under the name Gymnodexia orientalis Baranoff but
without description, listed three specimens in the DEI collection from Formosa. The name
remains a nomen nudum.

TYPE-MATERIAL OF TACHINIDAE 57

Hyalomyodes orientalis Baranov in Hennig, 1941 : 189. Nomen nudum.

Hennig listed, under this name and without description, a specimen in DEI collection
from Tainan in Formosa ; the name was not published by Baranov and remains a nomen
nudum.

Kosempomyia sauteri Baranov in Baranov, I934/ : 165. Manuscript name cited in synonymy,
unavailable.

This name was cited by Baranov in synonymy with Kosempomyiella rufiventris Baranov,
and is unavailable under Article 1 1 (d) of the International Code of Zoological Nomenclature,
1961. Hennig (1941 : 187) listed the name without description, and mentioned three speci-
mens in DEI collection, two from Formosa without other data and one from Kosempo in
Formosa.

Leskia deaurata Baranov in Hennig, 1941 : 190. Nomen nudum.

Hennig listed, under this name and without description, a specimen in DEI collection from
Sokutsu in Formosa ; the name was not published by Baranov and remains a nomen nudum.

Masicerella indistincta Gardner, 1940 : 178 (Masicerella indistincta Baranov MS). Available
specific name, attributable to Gardner.

Gardner described and figured the puparium of a species of Tachinid fly under Baranov's
hitherto unpublished manuscript name Masicerella indistincta. The specific name indistincta,
even though published in combination with an unavailable genus-group name, is available
under Article n (g) (ii) of the International Code of Zoological Nomenclature, 1961, and is
attributable to Gardner under Article 10. The generic name Masicerella (as mentioned by
Crosskey, 19670 : 18) is not accompanied, however, by a definition of the generic taxon and
is a nomen nudum which is unavailable under Article 13 (a).

Medinodexia formosana Baranov in Hennig, 1941 : 190. Nomen nudum.

Hennig listed, under this name and without description, a specimen in DEI collection from
Tainan in Formosa ; the name was not published by Baranov and remains a nomen nudum.

Meigenia setosa Baranov in Hennig, 1941 : 193. Nomen nudum.

Hennig listed, under this name and without description, a specimen in DEI collection from
Hoozan in Formosa ; the name was not published by Baranov and remains a nomen nudum.

Nemosturmia winthemioid.es Mesnil, 1949 : 76 (Exorista winthemioides Baranov MS).
Available name, attributable to Mesnil.

Mesnil (1949 : 76) described Nemosturmia winthemioides as a new species based upon a
male holotype specimen from Formosa in the DEI collection labelled by Baranov as
' Exorista Winthemioides ' ; Mesnil erroneously attributed authorship of the name win-
themioides to Baranov, but Baranov did not publish the name, and authorship of the name
winthemioides is attributable to Mesnil.

Hennig (1941 : 194) listed the name ' Exorista winthemioides Baranoff ' and mentioned
the single specimen in DEI collection (now the holotype of Nemosturmia winthemioides
Mesnil), but he gave no description.

Paradionaea orientalis Baranov in Hennig, 1941 : 189. Nomen nudum.

Hennig listed, under this name and without description, a specimen in DEI collection
from Kankau in Formosa ; the name was not published by Baranov and remains a nomen
nudum.

Phasia dubiosa Baranov. See Allophora dubiosa Baranov.

Phorinia flavipalpis Baranov in Hennig, 1941 : 194. Nomen nudum.

Hennig listed, under this name and without description, five specimens in DEI collection
from Kankau in Formosa ; the name was not published by Baranov and remains a nomen
nudum.

Phoriniophylax fetnorata Mesnil, 1957 : J4 (Phoriniophylax femorata Baranov MS). Avail-
able name, attributable to Mesnil.

58 C. W. SABROSKY & R. W. CROSSKEY

Hennig (1941 : 196) listed, under this name but without description, two specimens in
DEI collection from Tainan in Formosa that are determined by Baranov as ' Phoriniophylax
femorata ', but Baranov never published this name. Later Mesnil (1944 : 27) placed the
name femorata Baranov in combination with Argyrophylax Brauer & Bergenstamm, but
without validating it by description so that it remained at that time (1944) a nomen nudum.
In a more recent work, however, Mesnil (1957 : 14) appended to the description of his new
subspecies Argyrophylax nova novella Mesnil the following remark : ' il est probable que
Phoriniophylax femorata Baranov est congenerique de cette espece [Argyrophylax nova
Mesnil]. Elle s'en distingue par ses pattes jaunes a tarses noirs '. We consider that this
statement purports to give characters differentiating the taxon femorata, and that the name
is nomenclaturally available under Article 13 (a) (i) of the International Code of Zoological
Nomenclature, 1961 ; under Article 10 it takes the authorship and date of Mesnil, 1957.

Prosopaea sturmioides Baranov. See Ctenophorocera sturmioides Mesnil.

Siphona taiwanica Baranov. See Crocuta taiwanica Baranov.

Sumpigaster formosensis Baranov in Hennig, 1941 : 190. Nomen nudum.

Hennig listed, under this name but without description, three specimens in DEI collection
from Formosa (no other data) ; the name was not published by Baranov and remains a
nomen nudum.

Zenilliana pulchra Mesnil, 1949 : 68 (Exorista pulchra Baranov MS.) Available name,
attributable to Mesnil.

Mesnil (1949 : 68) described Zenilliana pulchra as a new species based upon a male holotype
specimen from Sokutsu, Formosa, in the DEI collection, but erroneously attributed author-
ship of the name to Baranov ; however, Baranov did not publish it, and authorship of the
name pulchra is attributable to Mesnil.

Hennig (1941 : 194) listed the name ' Exorista pulchra Baranoff ' and mentioned the
single specimen in DEI collection (now the holotype of Zenilliana pulchra Mesnil), but he
gave no description ; hence Exorista pulchra Baranov in Hennig is a nomen nudum.

TYPE-MATERIAL OF TACHINIDAE 59

REFERENCES

Note : the references that follow contain a complete bibliography for Baranov's papers on
Tachinidae, although it has not been necessary to cite all of these in the foregoing text.

AUBERTIN, D. 1932. A tachinid fly parasitic on the bee-hole borer of teak. Stylops 1 :

35-36.
BARANOV, N. 19260. Die in Serbien gesammelten Dexiinae. Encycl. ent. Serie B II, 3 :

56-60.

- 19266. Prilog poznavanju srpskih Tachina (Beitrag zur Kenntnis der serbischen Tachini-
den). Letopis poljoprivredne, ogledne i kontrolne stanice u Topcideru, No. i : 153-184.
[In Serbian and German].

- 1927. Die nach Hypopygiumbau geordneten in Serbien gesammelten Tachinidae.
Encycl. ent. Serie B II, 4 : 31-44.

- 1928. Tachinidensammlung des zoologischen Museums in Zagreb. Glasn. hrv. prirodosl,
Drust. 39-40 : 196-200.

— 19290. Studien an pathogenen und parasitischen Insekten I. Die Jugoslavischen
Arten der Tachinidengruppe Echinomyia. Arb. parasit. Abt. Inst. Hyg. Zagreb, No. i :
1-23.

- 19296. Studien an pathogenen und parasitischen Insekten II. Beitrag zur Kenntnis
der Phasiinen mit besonderer Beriicksichtigung der Gruppe Ocyptera (Diptera. Tachin.).
Arb. parasit. Abt. Inst. Hyg. Zagreb, No. 2 : 1-22.

— I929C. A contribution to the morphology of the tachinid flies bred from Pyrausta nubilalis
Hb. Scient. Rep. int. Corn Borer Invest. 2 : 128-130.

— 19300. Nekoliko rije£i o Tachinama, koje parazitiraju na Pyrausta nubilalis Hb. Acta
Soc. ent. jugosl. 3-4 (1928-1929) : 103-108.

— 19306. Die wahre Ceromasia senilis Mg. und juvenilis Girschn. (Dipt., Tachin.). Kono-
wia 9 : 34-36.

— 19300. Die Sternitenkette des Abdomens bei den parasitaren Raupenfliegen und ihre
systematische Bedeutung. Z. Parasit Kde 2 : 506-534.

— 19310. Studien an pathogenen und parasitischen Insekten III. Beitrag zur Kenntnis
der Raupenfliegengattung Carcelia R.D. Arb. parasit. Abt. Inst. Hyg. Zagreb, No. 3 : 1-45.

— 19316. t)ber Raupenfliegen, welche durch die Farbe des Abdomens Exorista confinis
Fall, ahnlich sind. Wiener, ent. Ztg 48 : 117-124.

— I93ic. Carcelia ambigua Villen.= Carcelia rasella Baranoff. Konowia 10 : 290.

— 19320. Neue orientalische Tachinidae. Encycl. ent. Serie B II, 6 : 83-93.

— 19326. Zur Kenntnis der formosanischen Sturmien (Dipt. Larvaevor.). Neue Beitr.
syst. Insektenk. 5 : 70-82.

— 19320. Zur Kenntnis der orientalischen Winthemia-A.rten (Dipt. Larvaev.). Ent.
NachrBL, Troppau 6 : 45-47.

— 19320". Larvaevoridae (Ins. Dipt.) von Sumatra, I. Miscnea zool. sumatr. 66 : 1-3.
1932^. Zur Kenntnis der orientalischen De#z'o-ahnlichen Arten. Wiener ent. Ztg 49 :

212-216.

— 19327. Bestimmungstabelle der orientalischen Sturmia- Arten der scutellata-Gruppe
(Dipt. Larv.). (Mit Beriicksichtigung einiger verwandten Gattungen). Ent. NachrBL,
Troppau 6 : 100-101.

- 1932^. O Eutachina civilis Rnd., parazitu metlice. Acta Soc. ent. jugosl. 5-6 (1930-1931) :
62-65.

— 1933. Cadurcia leefmansi, eine neue orientalische Raupenfliege (Dipt. Tach.). Treubia
14 : 153-154.

— 19340. Mitteilungen uber geziichtete orientalische Larvaevoriden. (Insecta, Diptera).
Ent. NachrBL, Troppau 8 : 41-49.

60 C. W. SABROSKY & R. W. CROSSKEY

BARANOV, N. 19346. Zur Kenntnis der Raupenfliegenfauna der Salomon-Inseln (Dipt.,
Tachinidae). Stylops 3 : 181-184.

- I934C- Zur Kenntnis der parasitaren Raupenfliegen der Salomonen, Neubritanniens,
der Admiralitats-Inseln, der Fidschi-Inseln und Neukaledoniens, nebst einer Bestimmungs-
tabelle der orientalischen Sturmia-Arten. Vet. Arh. 4 : 472-485.

- 1934^- Obersicht der orientalischen Gattungen und Arten des Carcelia-Komplexes
(Diptera : Tachinidae). Trans. R. ent. Soc. Land. 82 : 387-408.

— 19340. Ein interessanter Fall von Sphecoidie bei der Larvaevoride Vespocyptera petiolata
Townsend. Encycl. ent. Serie B II, 7 : 157-160.

- I934/- Neue Gattungen und Arten der orientalischen Raupenfliegen (Larvaevoridae) .
Encycl. ent. Serie B II, 7 : 160-165.

- 19350- Neue palaarktische und orientalische Raupenfliegen (Dipt., Tachinidae). Vet.
Arh. 5 : 550-560.

- 19356- Eme neue orientalische Raupenfliege. Meded. LandbHoogesch. Wageningen
39 (i) : [39-40]-

- I935C. Larvaevoridae (= Tachinidae, Dipt.). Wiss. Ergebn. niederl. Exped. Karakorum
1 : 407-409.

- 1936. Weitere Beitrage zur Kenntnis der parasitaren Raupenfliegen (Tachinidae =
Larvaevoridae) von den Salomonen und Neubritannien. Ann. Mag. nat. Hist. (10) 17 :
97-H3-

— 19380;. Weiteres iiber die Tachiniden (s.l.) der Salomon-Inseln. Vet. Arh. 8 : 170-174.

— 19386. Neue indo-australische Tachinidae. Bull. ent. Res. 29 : 405-414.

- 19380. Raupenfliegen (Tachinidae s.l.) welche auf der Adria-Insel Pag beim Trinken von
Meerwasser gefangen wurden. Encycl. ent. Serie B II, 9 : 103-107.

- 1939. Sechs neue Raupenfliegen aus der Sammlung Takanos. Ent. NachrBL, Troppau
12 (1938) : 110-112.

- 1942. Ein neuer Vespidenparasit von Java und eine mit ihm verwandte Fliege von den
Salomon-Inseln. Vet. Arh. 12 : 161-163.

BARANOV, N. & HERGULA, B. 1928. Uber die systematische Stellung der aus Pyrausta
nubilalis Hb. erzogenen Tachine Ceromasia senilis (Mg.) auct. nov. Glasn. hrv. prirodosl.
Drust. 39-40 : 192-195.

BEESON, C. F. C. & CHATTERJEE, S. N. 1935. On the biology of the Tachinidae (Diptera).
Indian Forest Rec. (Entom.) N.S., 1 : 169-184.

CROSSKEY, R. W. 1963. A systematic review of the Oriental and Australasian species of
Argyrophylax Brauer and Bergenstamm, including the description of a new species from
New Britain (Diptera : Tachinidae). Ann. Mag. nat. Hist. (13) 6 : 1-16.

- 1966. New generic and specific synonymy in Australian Tachinidae (Diptera). Proc. R.
ent. Soc. Lond. (B) 35 : 101-110.

- 19670. An index-catalogue of the genus-group names of Oriental and Australasian
Tachinidae (Diptera) and their type-species. Bull. Br. Mus. nat. Hist. (Ent.) 20 : 1-39.

- 19676. New generic and specific synonymy in Oriental Tachinidae (Diptera). Proc. R.
ent. Soc. Lond. (B) 36 : 95-108.

— 19670. A revision of the Oriental species of Palexorista Townsend (Diptera : Tachinidae,
Sturmiini). Bull. Br. Mus. nat. Hist. (Ent.) 21 : 35-97.

D'AGUILAR, J. 1957- Revision des Voriini de 1'ancien monde (Dipt. Tachinidae). Annls
Epiphyt. Serie C, 8 : 235-270.

GARDNER, J. C. M. 1940. The puparia of some Indian Tachinidae (Diptera). II. Indian
J. Ent. 2 : 177-181.

HENNIG, W. 1941. Verzeichnis der Dipteren von Formosa. Ent. Beih. Berl.-Dahlem 8 :
1-239.

MESNIL, L. P. 1944. Larvaevorinae (Tachininae) . In Lindner, Fliegen palaearkt. Reg.
64g : 1-48.

TYPE-MATERIAL OF TACHINIDAE

61

MESNIL, L. P. 1949. Larvaevorinae (Tachininae) . In Lindner, Fliegen palaearkt. Reg. 64g :
49-104.

1950. Larvaevorinae (Tachininae). In Lindner, Fliegen palaearkt. Reg. 64g : 105-160.

1951. Larvaevorinae (Tachininae). In Lindner, Fliegen palaearkt. Reg. 64g : 161-208.

- 1957. Nouveaux Tachinaires d'Orient (Deuxieme serie). Mem. Soc. r. ent. Belg. 28 : 1-80.

1960. Larvaevorinae (Tachininae). In Lindner, Fliegen palaearkt. Reg. 64g : 561-608.

1962. Larvaevorinae (Tachininae). In Lindner, Fliegen palaearkt. Reg. 64g : 705-752.

ROEPKE, W. 1935. De slakrupsenplaag op het Molukken-eiland Batjan. Meded. Landb-

Hoogesch. Wageningen 39 (i) : 1-38 (with two-page unpaginated appendix).
SENIOR-WHITE, R. 1922. Notes on Indian Diptera. Mem. Dep. Agric. India ent. Ser. 7 :

i-vi + 83-169.
, AUBERTIN, D. & SMART, J. 1940. Family Calliphoridae. Fauna Br. India, Diptera 6,

288 pp., London.
TOWNSEND, C. H. T. 1938. Manual of Myiology. Part VII. 434 pp. Itaquaquecetuba,

Sao Paulo.

— 1939. Manual of Myiology. Part IX. 270 pp. Itaquaquecetuba, Sao Paulo.
1941. Manual of Myiology. Part XL 342pp. Itaquaquecetuba, Sao Paulo.

INDEX TO SPECIES-GROUP NAMES

aberrans, Carcelia, 36
aestivalis, Echinomyia praeceps, 40
albopunctata, Alophora, 35
anomala, Phorocerosoma, 49
apicalis, Exorista, 56
argenteostriata, Eutachina, 42
ater, Dolichocolon, 56
atkinsoni, Cossidophaga, 55
atkinsoni, Gymnodexia, 45, 56
atkinsoni, Podomyia, 55
atropivora, Zygobothria, 51
aureifrons, Eutachina aureifrons, 42
aureisquamosa, Eutachina, 42
aurichalcea, Eutachina, 42
australe, Dolichocolon, 40

basalis, Eutachina, 42
bella, Sturmia, 50
bezziana, Myiobia, 47
bezziana, Myiofijia, 47
bezziana, Stomatomyia, 49
bezzii, Erycia, 41
bisetosa, Sturmia, 50
buitenzorgiensis, Carcelia, 37
burmanica, Chaetoptiliopsis, 39

caldwelli, Monoleptophaga, 47
caldwelli, Zenillia, 54
caspica, Eucarcelia, 41
caudata, Carcelia, 37
caudatella, Carcelia, 37
chatter jeeana, Sturmia, 50

civiloides, Eutachina, 42
compressa, Platerycia, 49
compsiluroides, Bactromyia, 55
crassiseta, Bactromyia, 35

dammermani, Eucarcelia, 41
deaurata, Leskia, 57
distincta, Carcelia, 37
distincta, Exorista, 43
diversa, Winthemia, 54
diversoides, Winthemia, 54
dubiosa, Allophora, 55
dubiosa, Phasia, 55, 57
dubiosus, Goniophthalmus, 45

edentata, Voria, 53
eumorphophaga, Arrhinodexia, 35
eutachinoides, Blepharipoda, 36
excisa, Carcelia, 37

femorata, Phoriniophylax, 57
flavipalpis, Phorinia, 57
formosana, Cuphocera varia f., 55
formosana, Medinodexia, 57
formosensis, Sumpigaster, 58
fransseni, Bactromyia, 35
frontalis, Carcelia, 37
fuscipennis, Eutachina, 42

grisellina, Exorista, 56
grossa, Eucarcelia, 41

62

INDEX TO SPECIES-GROUP NAMES

hemimacquartioides, Isocarceliopsis, 45
hirsuta, Carcelia, 37
hokkaidensis, Vibrissina, 53
hokkaidensis, Winthemia, 54
hutsoni, Sturmia, 50
hyalipennis, Eutachina, 42

imperator, Phorocera, 48
incisuralis, Sumatrodexia, 52
inconspicuella, Sturmia, 50
inconspicuoides, Sturmia, 50
indica, Eucarcelia, 42
indica, Euhapalivora, 56
indistincta, Masicerella, 57
institutiimperialis, Leverella, 46
intermedia, Erycia, 41
isabeli, Phorocera, 48
issikii, Arrhinomyia, 35

japanica, Protonemoraea, 49
japanica, Tamanukia, 53

kalshoveni, Cossidophaga, 55
kockiana, Eucarcelia, 38

ladelli, Eutachina, 42
latiforceps, Sturmia, 50
latistylata, Parexorista, 48
latistylata, Sturmia, 50
leefmansi, Cadurcia, 36
leveri, Prosopodes, 49
leveriana, Eurystaea, 42

machaeralis, Hapalioloemus, 45
macrophallus, Sturmia, 51
maculibasis, Leiosiopsis, 46
maculiventris, Exorista, 56
magna, Myiostoma, 48
magna, Phorocera, 48
malayana, Carcelia, 37
malayana, Cuphocera varia f., 55
mallochi, Winthemia, 54
mallochiana, Actia, 54
maxima, Phorocera magna f., 49
mirabilis, Formosia, 44
mirabilis, Ilia, 45
montana, Sumatrodexia, 52
mungomeryi, Eutachina, 43
mutabilis, Meigenia, 47

niger, Plagioderophagus, 49
nigribarbis, Sturmia, 51
nigricosta, Erycia, 41

nigripes, Bezziomyiobia, 36
nigrotibialis, Argyrophylax, 35
nobilis, Meigenia mutabilis, 47
nova, Argyrophylax, 58
novaeguineae, Leverella, 46
novella, Argyrophylax nova, 58
nymphalidophaga, Erycia, 41

oceanica, Sturmia bella, 50
octava, Carcelia, 37
oculata, Masicera, 46
oculata, Sturmia, 51
orbitale, Dolichocolon, 40
orientalis, Calotheresia, 36
orientalis, Euvespivora, 43
orientalis, Gymnodexia, 56
orientalis, Hyalomyodes, 57
orientalis, Paradionaea, 57

painei, Sturmia, 51
palpata, Erycia, 41
pandellei, Fabriciella, 44
paradoxalis, Sturmia, 51
parvicornis, Takanoella, 53
phrynoides, Exorista, 43
picta, Exorista, 44
picta, Sturmia, 51
pilosa, Carcelia, 37
pilosa, Meigenia mutabilis, 47
pilosella, Carcelia, 37
postulans, Phorocerosoma, 49
prima, Carcelia, 37
pseudocaudata, Asiocarcelia, 35
psychidarum, Tricholyga, 53
pulchra, Exorista, 56, 58
pulchra, Zenilliana, 58
pulex, Actia, 35

quadrimaculata, Exorista, 44
quadriseta, Eutachina, 43
quadrisetosa, Eutachina, 43
quadrisetosum, Dolichocolon, 40
quarta, Carcelia, 38
quinta, Carcelia, 38

rasella, Carcelia, 38
rasoides, Carcelia, 39
rondaniella, Catacarcelia, 39
roseanella, Zenillia, 54
rufa, Carcelia, 39
rufescens, Dolichocolon, 40
rufipes, Dexiomimops, 40
rufitibialis, Argyrophylax, 55

INDEX TO SPECIES-GROUP NAMES

rufiventris, Kosempomyiella, 46, 57
rufofemorata, Erycia, 41
rusticella, Eutachina, 43
rutilloides, Carcelia, 39

salomonica, Euvespivora, 43
sauteri, Kosempomyia, 57
sauteri, Trichoformosomyia, 53
secunda, Carcelia, 39
seniorwhitei, Exorista, 44
septima, Carcelia, 39
setosa, Meigenia, 57
setosella, Carcelia, 39
sexta, Carcelia, 39
siamense, Euthelairosoma, 43
sillemi, Cnephalia, 39
simulator, Exorista, 56
solomonensis, Chaetexorista, 39
solomonensis, Doleschalla, 40
solomonica, Bactromyia fransseni, 36
solomonica, Winthemia diversa, 54
solomonicola, Formosia mirabilis, 44
soror, Sisyropa, 56
sturmioides, Ctenophorocera, 55
sturmioides, Prosopaea, 58
sumatrana, Eutachina aureifrons, 42
sumatrana, Sturmia, 51

taiwanica, Crocuta, 55
taiwanica, Siphona, 55, 58
takanoi, Actia, 35

takanoi, Erycia, 41
takanoi, Protonemoraea, 49
tenuiforceps, Eutachina, 43
tertia, Carcelia, 39
thermophila, Sisyropa, 56
townsendi, Macrozenillia, 46
trisetosa, Sturmia, 51
trisetosoides, Sturmia, 52

unguicularis, Sturmia, 52
unisetosa, Sturmia, 52

vanderwulpi, Cadurcia, 36
varia, Cuphocera, 55
vernalis, Echinomyia praeceps, 41
vicarium, Phorocerosoma, 49
vicinalis, Exorista, 44
vicinella, Sturmia, 52
villeneuvei, Hemidegeeria, 45
vittata, Sumatrodexia, 53
vulgaris, Meigenia mutabilis, 47
vulpinoides, Micropalpus, 47

wainwrighti, Sturmia, 52
winthemioides, Exorista, 56, 57
winthemioides, Nemosturmia, 57

yerburyi, Ctenophoroceropsis, 39
zetterstedti, Argyrophylax, 36

DR. CURTIS WILLIAMS SABROSKY, A.B., M.S., Sc.D.(Hon.

SYSTEMATIC ENTOMOLOGY LABORATORY,

U.S. DEPT. OF AGRICULTURE,

c/o U.S. NATIONAL MUSEUM,

WASHINGTON, D.C., 20560,

U.S.A.

DR. ROGER WARD CROSSKEY, D.Sc., A.R.C.S., F.I.Biol.,
COMMONWEALTH INSTITUTE OF ENTOMOLOGY,
c/o BRITISH MUSEUM (NATURAL HISTORY),
CROMWELL ROAD,
LONDON, S.W-7, ENGLAND.

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?ffi :

BOMBYLIIDAE, AND A FIRST

RECORD OF NEMESTRINIDAE

FROM SOKOTRA (DIPTERA)

D. J. GREATHEAD

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 24 No. 3

LONDON: 1969

BOMBYLIIDAE, AND A FIRST RECORD

OF NEMESTRINIDAE
FROM SOKOTRA (DIPTERA)

BY

DAVID JOHN GREATHEAD

-V t v.X.

East African Station, C.I. B.C., Kampala, Uganda

Pp. 65-82 ; 8 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 24 No. 3

LONDON : 1969

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Parts will appear at irregular intervals as they
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pleted within one calendar year.

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

This paper is Vol. 24, No. 3 of the Entomology
series. The abbreviated titles of periodicals cited
follow those of the World List of Scientific Periodicals.

World List abbreviation :
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Issued 3 December, 1969 Price Eleven Shillings

BOMBYLIIDAE, AND A FIRST RECORD OF
NEMESTRINIDAE FROM SOKOTRA (DIPTERA)

D. J. GREATHEAD

SYNOPSIS

The Bombyliidae of Sokotra are reviewed and new species described as the result of the
examination of a new collection of specimens from the island made by K. M. Guichard in 1967.
The first record of a Nemestrinid, Atriadops cinnamonea Brunetti, is also reported.

THE first collections of Diptera from Sokotra were made by W. R. O. Grant, pub-
lished by Ricardo (1903), and by O. Simony, published by Becker (1910), both in
the same year, 1899. No subsequent collections of Diptera were made until 1967
when a collecting expedition to Southern Arabia and Sokotra was made by K. M.
Guichard. New material of all the described species from the island has thus
become available as well as material of other species both new and not previously
known from the island. This new material was collected between March and May,
whereas previous visitors who collected Bombyliidae were there in December to
February. This difference in timing has not, as far as can be inferred from the
Bombyliidae, made a marked difference in the species in flight.

All three collectors travelled in the relatively flat northern part of the island. No
specimens of Bombyliidae are available from the higher parts of the Haggier moun-
tains or the southern coastal area. Popov (1957) has provided a description of the
vegetation, which can be classified as subdesert and, like the fauna, includes a high
proportion of endemics.

The Bombyliidae now known from the island comprise 16 or 17 species, of which
9 are not known from outside Sokotra and Abd-el-Kuri. The remainder are all
recorded from the arid adjoining areas of the African mainland and four from
Southern Arabia. Of the endemic species only one, Bombylius socotrae, is not
clearly allied to species known from neighbouring parts of Africa and Arabia. As,
however, very little is known of the Bombyliidae of the Somali Peninsula any
detailed comparison is premature. Examination of the material leaves a general
impression of a tendency to greater size and more striking colouring than mainland
counterparts, thus Petrorossia sokotrae is on average much larger than any African
species and the specimens of Hemipenthes inauratus and Anthrax fuscipennis are all
at the upper limit of size found on the mainland. Villa dioscoridae and Exoprosopa
punctipennis both show the latter tendency to a striking extent.

The record of a Nemestrinid from the island is of interest both because it adds a
new family to the faunal list and because it is apparently the same as a species
described from Malawi. However, the extent of the distributions of Nemestrinidae
cannot be assessed as they are seldom captured.

68 D. J. GREATHEAD

Family BOMBYLIIDAE
Bombylius mollis Bezzi

Bombylius mollis Bezzi, 1921 : 15.

The specimen listed as Bombylius sp. by Ricardo (1903) has been examined ;
although the proboscis is missing and the dorsal surface largely denuded, there is
no doubt as to its identity. B. mollis is a widespread species ranging from Ethiopia
to Transvaal.

SOKOTRA : Adho Dimellus, 3500 ft, i <£, 9.11.1899 (W. R. 0. Grant), BM(NH).

Bombylius socotrae sp. n.

A single female Bombylius, the only other specimen of the genus from Sokotra,
represents a remarkable new species with a combination of characters not fitting
into any of the groupings erected by Bezzi (1924). It is nearest to the B. minor
group in its pale whitish pubescence, pale legs with black spines, hyaline wings and
black body. These characters, as well as the brown antennae, black proboscis
and complete absence of black hairs or bristles except on the legs distinguish it
from other species.

Head : black with heavy grey tomentum except for the buccal margin which is yellowish
grey. Eyes separated by almost three times the width of the ocellar tubercle at the vertex,
which is tumid and separated by a groove from the frons. All hair and scales white, shaggy
on frons and face, shorter elsewhere and with a broad band of scales surrounding the eyes.
Antennae with third segments missing, first and second reddish brown to greyish, first three
times the length of second, hairs on first white, second bare except for minute yellowish spicules.
Proboscis black, with a reddish base twice the length of the head, palpi reddish brown with
colourless spicules. Thorax : black, heavily dusted with grey-brown tomentum on the dorsal
surface and grey elsewhere, densely covered in greyish white hair, macrochaetae glistening
white. Legs : coxae black, remainder brown, lighter on the femora and tibiae, darker on the
tarsi which are blackish at their tips. Hair on femora and scales white, spines and spicules
well developed, black. Fore and mid femora unarmed, hind with a row of six anteroventral
spines as well as apical spines above and a single anterodorsal and a single posterodorsal one
near the apex. Claws mostly missing, posterior ones black with a reddish base, pulvilli white,
almost as long as the claws. Wings : hyaline but tinged opaque yellowish at the base in
costal cell and in first basal cell. Veins reddish brown at fore border, paler towards the posterior
margin where they are yellowish brown. Venation ; middle crossvein almost at middle of
discoidal cell, first posterior cell blunt at apex and stalk long, longer than penultimate section
of vein. Squamae and alulae white with white fringes. Hal teres brownish tinged. Abdomen:
black with a brownish grey tomentum, tergites reddish at their sides and yellowish on the
posterior margins, more broadly at the apex than base. Hairs and bristles white, the latter
inconspicuous, weak, hairlike. Ovipositor concealed in a dense tuft of yellow hair.

Length of body 9 mm ; of wing 8 mm ; of proboscis 4 mm.

Holotype $. SOKOTRA : Kalinsiya, sea-level, 26.^.1967 (K. M. Guichard),
BM(NH).

BOMBYLIIDAE, AND NEMESTRINIDAE FROM SOKOTRA 69

Systoechus somali Oldroyd
Systoechus somali Oldroyd, 1947 : 105.

This species, which is well known as a predator of the desert locust, Schistocerca
gregaria Forskal, is common in Somalia, the Ogaden in Ethiopia and northern
Kenya (Hynes, 1947 ; Greathead, 1958). Only a single specimen was collected in
Sokotra, but it shows no differences from mainland specimens.

The species belongs to the group with hyaline wings, entirely pale bristles on the
legs and no abdominal bristles in the male. The male is distinguished from other
species by the entirely blackish hair on the frons and face, dark brown pleural hair,
and pale, almost white, silky abdominal hair. The female has white facial hair,
three grey vittae on the thorax and the abdomen with pale hairs and dense rows of
dark bristles across the tergites.

SOKOTRA : Hammadero, noo ft, i $, 18.^.1967 (K. M. Guichard], BM(NH).

Geron sp. $

Hesse (1938) has shown that the species of Geron are remarkably similar in external
characters and can only be distinguished with difficulty except by the characters
of the male genitalia. He has also shown that, at least in southern Africa, there are
many more species than was formerly believed. It is thus impossible in the present
state of our knowledge to identify females from other regions reliably. The present
specimens however run to G. nigrifacies in Hesse's (1938) key and approximate to
his description. From G. nigrifacies they differ in that the fore tibiae are paler,
the brassy scaling extends over the scutum and the hyaline wings have paler yellowish
to yellowish brown veins.

SOKOTRA : Hammadero, noo ft, 2 ?, 8.^.1967 (K. M. Guichard), BM(NH) and
author's collection.

These specimens may be conspecific with the pair identified by Becker (1910)
as G. gibbosus Meigen, which it has not been possible to trace.

Phthiria sp.

Ricardo (1903) reported two male Phthiria sp. from Sokotra. No specimens of
this genus are present in Guichard's material and unfortunately the specimens
collected by Grant can no longer be found in the British Museum.

Chiasmella sica sp. n.

The genus Chiasmella was erected by Bezzi (1924) for the single species brevipennis
Bezzi from Southern Yemen. Doubt has been expressed whether the genus was
distinct from Chionamoeba Sack, but Greathead (19670:) showed that it is a distinct
genus of uncertain affinities, possibly closest to the Tomomyzinae in which he pro-
visionally included it. The present species agrees in most characters with C.
brevipennis, differing most notably in that the wing has an infuscated fore border,
and that the tomentum and hair on the frons is gold in the male. It is a darker,
more powerfully built species resembling a There vid in appearance.

70 D. J. GREATHEAD

<J. Head : black, but frons red-brown and buccal rim yellowish. Occiput above and vertex
covered in brown tomentum and blackish brown hair, frons with dense glittering gold tomentum
and fine gold hairs, face with greyish tomentum and white hair and lower part of occiput with
white hair, which is flattened and adpressed at the margins of the eyes. Antennae with first
segment brown, second and third blackish, hairs pale yellowish. Styliform part of third longer
than in C. brevifacies, equal in length to that of the rest of the antenna. Proboscis and palpi
black, labella brown, hairs colourless. Thorax : black ; dorsal surface with dark brown tomen-
tum, pale yellowish hairs on the anterior half, blackish ones on the posterior half, and with
thin reddish gold hairlike scales. Notopleural stripe consists of an ill-defined band of white
scaly hairs continuing around the margin of the scutellum as small white scales. Pleurae
with white tomentum and white scaly hair. Macrochaetae strong, reddish gold. Legs : coxae
and basal three-quarters of femora black, remainder yellow-brown but tarsi darker. Basal
part of legs, to apices of femora, with dense white hair and scales, apices of femora and tibiae
with yellow scales, tarsi with brown scales. Spines and spicules black, fore femora unarmed,
mid with spines only in apical third, hind in apical two-thirds. Wings : hyaline with a brown
infuscation in the costal, sub-basal, first basal, basal half of marginal and basal part of first
submarginal cells to just beyond the middle crossvein. Venation as C. brevifacies, except that
the second vein originates at a distance of about twice the length of the middle crossvein from
that vein. Halteres yellow brown, with apices of knobs paler. Abdomen : black, hair at
sides of first tergites and curved margins of succeeding tergites white. Median part of first
and dorsal part of succeeding tergites with dark reddish gold scales and a posterior fringe of
white scales. Lateral and ventral surfaces with white scales, hind margins of segments with
tiny black hairs, longer and more conspicuous dorsally and towards the apex. Sternites
almost entirely concealed by the recurved tergites. Hypopygium not examined.

Length of body 10 mm, of wing 8 mm.

$. As male except that the head is entirely black and the gold on the frons is replaced by
sparse whitish tomentum and pale yellowish white hair. Ovipositor concealed by a mass of
pale gold hair.

Holotype^. SOKOTRA : Jebel Ommari, Hadibo Plain, 600 ft, 22.111.1967 (K. M.
Guichard), BM(NH).

Paratypes. SOKOTRA : Hadibo Plain, 2 $, 111.1967 (K. M. Guichard), BM(NH)
and author's collection.

Petrorossia sokotrae (Ricardo) comb. n.
(Text-figs. 1-2)

Anthrax sokotrae Ricardo, 1903 : 367.
Thyridanthrax sokotrae (Ricardo) Bezzi, 1924 : 22.

This species, which clearly belongs to the genus Petrorossia, was unaccountably
listed by Bezzi (1929) under Thyridanthrax. It is one of the largest species of the
genus but very variable in size ; the present specimens range between 5 and
12 mm. It belongs to the P. fulvipes Loew section of the genus but lacks a distinct
wing pattern, the wings being tinged brownish, darker at the base and along the
fore border and identical in both sexes.

Head : black, with grey tomentum except on the vertex and upper part of the frons, which
are velvety black. Ocellar tubercle prominent, vertex almost three times its width in the male
and only slightly wider in the female, and with a deep sulcus behind the ocellar tubercle separating
the inflated halves of the occiput. Hairs on vertex, frons except for a few at the base of the
antennae, and on the upper part of the sides of the occiput to the level of the antennae, black,

BOMBYLIIDAE, AND NEMESTRINIDAE FROM SOKOTRA 71

hairs at the base of the antennae and on the face reddish golden, remaining parts of occiput with
gold hairs and with short pale yellowish hairs below. Antennae black with a reddish tinge and
grey tomentum, hairs black, first segment large, almost as long as second and basal part of third
combined, third with a short conical basal part sharply marked off from an elongate stylar
part 3 times its length, style jointed at two-thirds of the distance from its base and with a short
pale spicule at the apex. Proboscis brown, with golden hairs, broad, only slightly longer than
the buccal cavity. Palpi short, black, with long golden hairs. Thorax : black with heavy
grey tomentum on the pleura. Hairs of collar, humeral callus and mesepimeron rich golden,
hair on metapleuron pale gold, scalelike, macrochaetae and fine scattered hairs on dorsal surface
black. Dorsal surface also with short adpressed golden scalelike hairs. Legs : coxae blackish
brown with grey tomentum and golden hair. Remainder red-brown, with the tips of the
tarsi darkened. Hairs and scales golden except for an admixture of black scales on the hind
femora in the middle above, and towards the apices of the hind tibiae in the female ; in the
male this blackening is more extensive with black scales and even black cuticle at the base of
the femora. Spines and spicules black. Middle and hind femora with antero- and postero-
ventral rows of small black spines. Claws black with brownish bases. Pulvilli well developed,
whitish. Wings : greatly narrowed at base, alula absent, anal lobe extremely narrowed at
base. Membrane brownish glistening darker at the base and along the fore-border. Veins
brown. Upper branch of third vein sometimes with a short appendix. Venation normal.
Squamae small, brownish tinged, with whitish fringes. Halteres with brown stalks and paler
brown to ivory knobs, flattened. Abdomen elongate, flattened, reddish with black on tergites
above except for their hind margins and the apical, sixth, tergite which is entirely red. First
tergite covered with dense golden hair, and with dense flattened hair of the same colour below
the scutellum and along the hind margin. Remaining tergites with sparse fine black hair and
elongate gold scaly hairs. Sternites with gold hair and scales. Hypopygium (Text-figs.

0- 2 mm.

FIGS. 1-2. Petrorossia sokotrae. i, Lateral view of hypopygium ; 2, apical view of

telomere.

72 D. J. GREATHEAD

i & 2) red. Telomere large and with unusual thick bristles on the dorsal surface and at
the apex. Aedeagus and accessory structures less elaborate than in other species so far illus-
trated. Dense gold hair in the apical cavity concealing the ovipositor.

Length of body 7-13 mm, of wing 6-12 mm.

SOKOTRA : Adho Dimellus, 2800 ft, 5 <£, i <j>, 24.^.1967 (K. M. Guichard), i $
with same data except caught on 25.^.1967, BM(NH) and author's collection.

Ricardo's type-series (loc. cit.) were captured at Homhil, Hadibo Plain, Addah
Valley, as well as Adho Dimellus during January and February.

Anthrax fuscipennis (Ricardo)

Argyramoeba fuscipennis Ricardo, 1903 : 366.
Anthrax fuscipennis (Ricardo) Becker, 1910 : 132.

Becker (1910) placed his A. dentata as a synonym of A. fuscipennis ; as thus
accepted the species is widespread in the Mediterranean region, Arabia and north
eastern Africa. Although many species of Anthrax are widespread and subject to
variation in details of the colour pattern, this interpretation of the species should be
treated with caution until the hypopygia of males from a number of localities can
be checked with those of males from Sokotra. Unfortunately this is not yet possible
as no male specimens have been captured on the island. The form of the hypopygium
of material from northern Ethopia has been illustrated (Greathead, 19670).

SOKOTRA : Hadibo Plain, foothills 500 ft, i $, 30.^.1967 (K. M. Guichard},
Hadibo Plain, i $, iv. 1967 (K. M. Guichard), author's collection.

Ricardo's type-series were captured at Adho Dimellus and on the Hadibo Plain
during January and February and Becker reported the species from Ras Shoab
during the same months.

Anthrax aygulus Fabricius
Anthrax aygulus Fabricius, 1805 : 121.

This species is now known to be widespread in the northern part of the Ethiopian
Region. It has not previously been reported from Sokotra.

SOKOTRA : Hadibo Plain, i <J, ^.1967 (K. M. Guichard), BM(NH).

Spogostylum sp.? ventrale Bezzi

Spogostylum ventrale Bezzi, 1924 : 174.

Three female specimens of a species of Spogostylum, in rather poor condition,
appear to belong to 5. ventrale Bezzi ; however as the species of this genus are
notoriously difficult to identify on external characters and many are apparently
restricted to small areas and replaced by closely similar species in adjoining areas,
the identification is regarded as provisional.

ABD-EL-KURI : Jebel Saleh, 1000-1500 ft, 3 $, 7^.1967 (K. M. Guichard},
BM(NH) and author's collection.

BOMBYLIIDAE, AND NEMESTRINIDAE FROM SOKOTRA 73

Hemipenthes inauratus (Klug)

Anthrax inauratus Klug, 1832 : No. i.
Thyridanthrax inauratus (Klug) Bezzi, 1924 : 204.
Thyridanthrax inauratus (Klug) ; Engel, 1932-7 : 534.
Hemipenthes inauratus (Klug) Bowden, 1964 : 98.

H . inauratus is now known from localities in Ghana (Bowden, 1964) east to Somalia
(Bezzi, 1924) and from Arabia (Klug, 1832) but has not previously been reported
from Sokotra. It is one of the most beautiful Bombyliidae and is at once recognized
by the purplish black head and thorax, dimidiate black wing pattern and golden
scaling of the dorsal surface of the abdomen. The Sokotra specimens fall within
the variation indicated in the quoted descriptions and are thus not morphologically
distinct from continental specimens.

SOKOTRA : Suk, sea-level, i <£, i <j>, 2^.1967 (K. M. Guichard), BM(NH).

Villa dioscoridae sp. n.

(Text-figs. 3-5)

Anthrax hottentotta Linnaeus, 1758 : 590 ; Ricardo, 1903 : 368 [Mis-identification].
Hemipenthes circumdatus Meigen, 1820 : 143 ; Becker, 1910 : 133 [Mis-identification].

Recent work, taking a more critical view of Villa spp. and making use of the
characters of the hypopygium, has suggested that there are more species in the
hottentotta - circumdata group than was formerly recognized (Lyneborg, 1965).
Ricardo (1903) even regarded circumdata as a synonym of hottentotta. For the
present purpose it is sufficient to state that the Sokotran species is distinct and not
a form of one of the Palaearctic members of the complex. Neither is it conspecific
with the allied African species galla Greathead (1967 a) to which it is closest.

From all the other species of the complex it differs in the brighter, more golden
and less yellow pubescence, and the more clear cut, contrasting black and yellow
banding of the abdomen, in having black hair on the face, black scales only on the
legs, and black bands of scales across the sternites.

Holotype <£. Head : black, as V. hottentotta (sensu Engel, 1932-7) or V. galla, except that
all the hairs and scales on the frons and face are black, only those on the occiput being yellow
and that the third antennal segment is tinged reddish. Thorax : similar to the other species
but the hairs shorter than that of V. hottentotta and more richly coloured than either species ;
that of the collar, dorsal surface and prealar callus deep golden yellow with a reddish tinge,
that on remaining parts of the pleura straw-coloured. Legs : dark reddish, clothed in black
scales and hairs except coxae ; fore and mid coxae with yellow hairs on basal half and black
hairs and bristles on apical half, hind coxae mainly with yellow pubescence and only a few
blackish bristles at the apex. Wings : veins darker, blackish, basal comb covered in black
scales except for a patch of silvery ones at the base in addition to the silvery ones of the
' epaulette '. Venation normal. Squama opaque white with a pale yellow fringe. Halteres
pale yellow-brown with a cream knob. Abdomen : hair short as V. galla but scales more
distinctly yellow and denser, so that the pattern of light and dark bands is sharper. First tergite
with shiny black scales, second with pale straw-yellow scales on basal third, remainder black,
third with a narrow band of pale scales at the base, remainder black, fourth basal two-thirds
pale, apical third black, fifth and sixth black with fringes of pale scales and tufts of long black

74

D. J. GREATHEAD

ones at the sides, seventh with pale scales fringed with long black ones. Sternites : only
fourth entirely covered in pale scales, first to third with black ones in middle and fifth to seventh
black with pale fringes. Hypopygium, (Text-figs. 3, 4 & 5), with black hair, very similar to the
other closely allied species but the accessory structures broader and with deeper wing-like
flanges.

Length of body 18 mm, of wing 16 mm.

Paratype $$. Head : as male but vertex slightly wider and scales on face beneath the
black hair yellow. Thorax : hair longer and denser. Legs : tending to less extensively black
vestiture on coxae and in some with a few yellow hairs and scales at the bases of the femora.
Wings : as male but pale scales at base of comb white and ' epaulette ' blackish. Abdomen :
hairs longer and denser, pattern as male but with a tendency for the pale bands to be broader
and wider at the sides than in the middle. Ovipositor reddish with yellow hair and about
eight reddish spines on each side.

Length of body 16-19 mm, of wing 9-17 mm.

Holotype <^
BM(NH).

Paratypes.
i $, 1.11.1899
i $, 7.11.1899

SOKOTRA : Adho Dimellus, 2800 ft, 25.iv.i967 (K. M. Guichard),

SOKOTRA : Alilo Valley, 3000 ft, i <?, i.xi.iSgg [sic] ; Alilo Valley,
Homhil, 1500 ft, 4 $, 20, 22, 25.1.1899 ; Adho Dimellus, 3500 ft,
Goahal Valley, i $, 16.1.1899 (all collected W. R. 0. Grant], all in

BM(NH) ; Hamadero, noo ft, i $, 8.^.1967 (K. M. Guichard}, in author's collection.

Two females from Abd-el-Kuri representing an isolated population have yellow
scales on the frons as well as the face, yellow hair on the face except on the midline,
hair shorter and paler more straw-coloured and (dorsal surfaces denuded) abdominal
pattern less sharp with scattered yellow scales among the black ones. From above
these females bear a close resemblance to V. galla.

ABD-EL-KURI : 2 $, 8^.1967 (K. M. Guichard), BM(NH) and author's collection.

FIGS. 3-5. Villa dioscoridae. 3, Lateral view of aedeagus ; 4, apical view of accessory

structures ; 5, dorsal view of same.

BOMBYLIIDAE, AND NEMESTRINIDAE FROM SOKOTRA 75

Thyridanthrax argentifrons (Becker) comb. n.

Hemipenthes argentifrons Becker, 1910 : 133.

Becker (loc. cit.) described the species from two males. The present material
consists of three rather denuded females, which agree more or less with Becker's
description except that there are few silver scales on the posterior abdominal
tergites. As such characters are liable to sexual dimorphism (e.g. T. cunamae
Greathead, 19670) there is little doubt that the present specimens represent the
female of T. argentifrons. It is one of the group of Paragus-\ike species, as noted
by Becker, but is at once distinguished from other known species by the heavy
silver scaling on the frons combined with the absence of yellow hair or scaling from
the abdomen with its contrasting black and white pattern. The following des-
cription is based on the females and includes differences from the males as described
by Becker.

Head : black, facial cone bluntly rounded at its apex, face convex, frons two and a half
times the width of the ocellar tubercle at the vertex (in male twice the width) . Hair on frons
and face short, sparse, glistening black. Scales on lower half of frons large, dense, silver,
projecting over bases of antennae, on upper part of frons and occiput small, silvery, opalescent,
on face similarly opalescent but appearing glistening black when viewed from some angles.
Antennae black, first and second segments cylindrical with short black hair except on the
underside of first where it is longer, third one and a half times length of first and second, broad
at base and closely applied to the second, tapering rapidly to a thick styliform part about
two-thirds the length of the segment and terminating in a minute spicule. Proboscis black
at base, brown towards apex, tips of labella projecting beyond buccal cavity. Palpi dark
brown with brown hairs. Thorax : black with a brown tomentum. Hair of collar and noto-
pleural tuft pale brownish yellow, not red-brown as described by Becker for the male. Dorsal
surface denuded but according to Becker the males have red-brown hair and metallic golden
scales. Notopleural stripes well developed, white, metapleural tuft and plumula also white.
Remaining areas of pleura bare or possibly denuded. Prosternum with a row of stiff black
hairs. Macrochaetae long, robust, black. Legs : black. Coxae with stiff black hair, femora
with sparse fine brown hair below, spines and claws black and scales glistening black. Fore
and mid femora unarmed, hind with three to four spines on the underside at the middle and one
subapical one. Wings : hyaline. Veins brown, basal hook and vestigial basal comb black.
Venation ; middle cross vein at basal third of discoidal cell, which is small and narrow, so that
the posterior cells are long, contact of third and fourth posterior cells with discoidal cell about
equal. Squama brownish at base, yellowish at margin, with a fringe of translucent white
scales. Abdomen : black with glossy black scales and stiff black hair at the sides and across
the hind margins of the tergites, except for the first which has white hair and two bands of
white scales, a narrow one along the hind edge of the first tergite [on second according to Becker]
and a broader one across the anterior third of the third tergite ; however the fourth to seventh
tergites are badly denuded and there are traces of silver scales intermixed with the black ones.
Sternites black, first four with opalescent whitish scales and brownish hair, remainder with
glossy black scales and black hair. Ovipositor exposed, with five reddish spines on each side.

Length of body 5-7 mm, of wing 5-7 mm.

SOKOTRA : Jebel Omari, Hadibo Plain, 600 ft, 2 ?, 22.iii.ig67 (K. M. Guichard) ;
Kalinsiya, sea-level, I $, 26.1^.1967 (K. M. Guichard), BM(NH) and author's
collection.

The syntypes, two males, were collected at Ras Shoab in January.

76 D. J. GREATHEAD

Thyridanthrax sp. ? argentifrons (Becker)

Three of the four female specimens from Abd-el-Kuri identified by Ricardo (1903)
as Anthrax sp. have been examined. They are all badly denuded, rather faded and
greasy, which makes certain identification impossible, but the general close resem-
blance to the preceding species suggests that they probably belong to it or to a
divergent subspecies developed in response to isolation on Abd-el-Kuri.

They differ from T. argentifrons females in that they are larger, 9-10 mm, the
cuticle is not completely black but tends to have reddish areas, antennae dark reddish,
the upper part of the pleura are densely covered in yellowish brown hair, the legs
are dark reddish, the hair at the base of the abdomen is pale yellowish, the terminal
abdominal segments show signs of extensive silver scaling and that the sternites are
all covered in pale hair and silvery scales.

ABD-EL-KURI, 2 ?, 22.11.1899 ; i ?, S.xii.iSgS (W. R. 0. Grant), BM(NH).

Thyridanthrax alatus (Becker) comb. n.

Hemipenthes alatus Becker, 1910 : 134.

The species was described from a single female and is represented by a female
in the present collection. It is a pale-haired species with brown areas on the body
and a shiny black vertex with indistinct ocellar tubercle, a combination of characters
allying it with T. capella Greathead and T. pallescens Greathead, a species-group
which seems, as far as it has been recognized, to be characteristic of the desert
areas bordering the Palaearctic and Ethiopian regions (Greathead, in press).

Head : irons to region of antennae, and a triangular area between the antennae, genae and
occiput black, lower parts red-brown, face conical, eye indentation barely indicated, bisecting
line very short, vertex broad, three times the distance between the posterior ocelli in width,
ocellar tubercle indistinct. Vertex glossy black, bare except for sparse black hairs, frons with
sparse black hair on the black area and fine opalescent scales, face and pale parts with short
yellowish hair and opalescent scales except for a tuft of short black hairs at the apex of the
facial cone, occiput with short pale yellowish hair and also with opalescent scales. Antennae ;
first segment red, remainder black, hairs on first and second segments black above, brownish
yellow below, third broad, strap-like, tapering sharply to a minute style at the apex, twisted so
that it is vertically flattened at the base and horizontally flattened at the apex. Proboscis
projecting, labral part red, labial part black, palpi brown elongate with yellowish hairs. Thorax :
black but with extensive brownish areas on the pleura, and scutellum red except for the extreme
base. Hairs of collar and pleura pale white, tinged yellowish, dorsal surface denuded but
showing traces of short golden brown hair and fine scales. Scales among the hairs at the sides
of the dorsal surface white, tinged buff. Macrochaetae yellow-brown. Metapleura and
hypopleura bare. Legs : black, except for upper surfaces of femora and coxae, which are
brown. Hair and scales on coxae white, spines, spicules and scales on remaining parts black.
Claws black with red bases. Wings : base, entire costal, basal and anal cells and marginal
and discoidal cells to level of middle crossvein tinged pale yellow, remainder smoky hyaline.
Veins yellow-brown. Upper branch of third vein straight at base and bent at right angles
at the first bend. Discoidal cell truncate, the vein between it and the second basal cell long
almost as long as the cell is wide at the wing margin, contact of discoidal and fourth posterior
cells equal to that of the latter with the second basal cell. Basal hook brown, comb black with
whitish scales. Squama and alula translucent yellowish with white fringes. Abdomen : black
along the midline and bases of tergites, and sternites merging to red-brown on the remainder

BOMBYLIIDAE, AND NEMESTRINIDAE FROM SOKOTRA 77

giving the appearance of red-brown with an ill denned black median stripe. Hair on first
segment and sides of basal half of second segment white, fine scattered hair on dorsal surface
of segments 2-4 and hair rows at the margins of tergites on the dorsal surface black, those at
the sides and on the apical segments yellowish ; scales at the bases of the tergites white gradually
becoming yellow at the apex, however the pattern is difficult to distinguish as the specimen is
rather denuded. Sternites with white scales and fine sparse yellowish hairs. Ovipositor
red-brown with three weak spines at each side.
Length of body n mm, of wing 9 mm.

ABD-EL-KURI : Jebel Saleh, 1000-1500 ft, i $, 7^.1967 (K. M. Guichard),
BM(NH).

Becker's type is from Sokotra, February 1899.

Exoprosopa punctipennis Ricardo

(Text-figs. 6, 7)
Exoprosopa punctipennis Ricardo, 1903 : 364.

This seems to be one of the commonest and most striking Bombyliidae from
Sokotra. It is very similar in appearance to E. punctulata, a common and wide-
spread species on the African mainland. It differs in being larger (10-15 rnm),
more brightly coloured and heavily marked, in having dark-haired pleura, in that
the abdomen is only obscurely reddish at the sides, and in that a short appendix
juts into the discoidal cell from the recurrent angle of the vein separating it from the
third posterior cell. None of these differences is an important character, and as it
has spicules on the fore tibiae, like E. punctulata but unlike most other species of
the genus, it is concluded that its resemblance to this species is one of close relation-
ship and that E. punctipennis is probably derived from it. Ricardo (1903) draws
attention to the appendix jutting into the discoidal cell from the vein dividing it
from the third posterior cell. This character, like many minor instabilities of wing
venation, does not invalidate the proposed relationship with E. punctulata, as an
examination of a series of this species shows that this vein is variable in length and
angularity and in the development of an appendix in one or both wings. Similarly
E. disrupta (Walker) which also belongs to the E. punctulata group (Greathead,
1967^ ; E. disrupta also has spiculate front tibiae, a point not previously mentioned)
shows a tendency to the development of an appendix into the discoidal cell.

Head : brownish black with a reddish buccal rim and a heavy red-brown tomentum. Face
conical, its sides and base forming an equilateral triangle in profile. Eyes separated by three
and a half to four times the width of the ocellar tubercle in both sexes. Eye indentation
barely indicated, bisecting line very short. Hairs sparse, black on vertex, frons, face and under-
side of head, on frons and face with scattered opalescent scales beneath, occiput with very short
brown hairs and dense silvery scales at the posterior margins of the eyes. Antennae with first
segment red-brown, two and a half times the length of the second, which is dull brown, shorter
than wide, third dull brown, as long as first and second segments together, with a rounded
base and tapering to an elongate red-brown style three-quarters the length of the third segment,
style with a distinct spicule at its apex. Proboscis black shagreened, as long as the greatest
length of the head, thus it is conspicuously projecting. Palpi black with fine black hairs.
Thorax : black except apical two-thirds of scutellum, which is red heavily dusted with brown
tomentum. Hair of collar above between notopleural stripes pale yellow, the hairs darker

78 D. J. GREATHEAD

yellow at their tips, sparse hair on dorsal surface, hair on pleura except upper parts of meta-
pleural tuft, black, also with dense stripes of white scaly hairs along notopleura, upper part of
metapleural tuft and plumula white. Scales on dorsal surface very sparse opalescent, macro-
chaetae black. Legs : black including all hairs scales and spines. Fore tibiae spiculate,
only hind femora with spines below, these with a complete row. Wings : base, costal cell,
first basal cell, submarginal cell except broad apical part, basal half of submarginal and second
basal cells, extreme base of first posterior cell, basal third of anal and base of axillary cell all
infuscated chocolate-brown; also with chocolate-brown spots at the bases of the second and
third submarginal, discoidal, second, third and fourth posterior cells. Basal hook and comb
black, squama white, its fringe and fringe of alula also white. Venation with veins separating
submarginal cells angularly bent, not smoothly curved, vein between discoidal and third basal
cell long and contorted, bent twice almost at right angles into the third basal cell (thus giving
a broad apex to the discoidal cell) and into the discoidal cell where a short appendix is emitted
into that cell. Abdomen, <$ : conical, tergites black, obscurely reddish at the sides, first tergite
with white hair except for a line of black hairs along its margin on the dorsal surface, second
tergite with white hair at the sides of the basal two-thirds and with a narrow band of white

FIGS. 6-7. Exoprosopa punctipennis. 6, Lateral view of hypopygium
aedeagus showing outline of accessory structure.

7, dorsal view of

BOMBYLIIDAE, AND NEMESTRINIDAE FROM SOKOTRA 79

scales at the base, otherwise with black hair and glossy black scales, third, fifth and sixth
tergites also with white basal bands, otherwise tergites with black hair and scales. Sternites
dark red with black hair and scales. Hypopygium (Text-figs. 6 & 7) red with black hairs.
Very similar to that of E. punctulata (Hesse, 1956 : fig. 254) differing only in minor details of
shape. $ : as male but flattened, not conical, and ovipositor red with five black spines on
each side.

Length of body 10-15 mm, of wing 8-14 mm.

SOKOTRA : Hammadero, noo ft, 2 ^, i $, 8.^.1967 ; Kalinsiya, sea-level,
2 c£, i °-, 26.111.1967 ; Hadibo Plain, foothills, 500 ft, i <£, 30.^.1967 ; Jebel Omari,
Hadibo Plain, 600 ft, i $, 22.111.1967 (all collected K. M. Guichard], BM(NH)
and author's collection.

Ricardo's (1903) type-series were collected at Homhil, Adho Dimellus and Hadibo
Plain, December - February. Becker (1910) recorded it from Sokotra, January
and February.

Exoprosopa insularis Ricardo

(Text-figs. 8, 9)
Exoprosopa insularis Ricardo, 1903 : 365.

Ricardo (1903) compared this species with E. disrupta but as shown above E.
dismpta is related to E. punctulata and E. punctipennis while E. insularis lacks the
special characters of this group ; in contrast the front tibiae are not spiculate,
the proboscis is short and the abdomen black, broad and flattened. Ricardo's
other comparison with E. venus Wied. [sic], in fact E. venosa Wied. sensu Loew
(described by Hesse, 1956, as E. leucothyrida), is more apposite. E. insularis has
the highly contorted venation between the submarginal cells, narrowed first posterior
cell, unstable venation tending to emit appendices either into the discoidal or third
posterior cells and concentration of the infuscation along the margin of the veins,
dark colouring and other body characters of species of the subgenus Acrodisca Bezzi.
It seems to belong more properly with this grouping than with the E. punctulata
group. As suggested by Bowden (1964) and accepted by Greathead (19670;),
there is room for a regrouping of species within the genus, relying more on general
similarities than single venational characters which are not always reliable. On
this basis both E. insularis and E. leucothyrida belong in a grouping centred on
Bezzi's subgenus Acrodisca.

Head : black with brown tomentum, brown around ocellar tubercle and in two stripes running
from it to the antennae, genae and buccal rim yellow-brown. Sparse short hairs on occiput
above, vertex, frons and face black. Hair at sides of occiput and fringing the posterior margin
pale yellow. Scales on vertex and frons glossy black, on face mixed glossy black and dull gold,
and on occiput dull gold. Eyes separated at vertex by three times the width of the ocellar
tubercle, which is set forward of the narrowest point at the top of the head. Eye indentation
straight-sided, shallow, so that the emargination is symmetrical and angular, bisecting line
short. Facial cone triangular in profile, the apical angle being almost a right angle. Antennae ;
first segment red with black hair, second black about half length of first with black hair, third
slightly longer than first two segments together, upper surface straight, lower sloping upwards
so that it is conical, with the apex over the circumference of the base, lower surface grooved,
black above narrowly red below, style about two-thirds length of the third segment, black and

8o

D. J. GREATHEAD

with a reddish spicule at the apex. Proboscis with only the tips of the labella projecting,
reddish black with a shiny black heavily shagreened labial sclerite below. Palpi black with
black hairs. Thorax : black with brown tomentum and a paler median stripe of tomentum
along the length of the scutum. Dorsal surface with sparse black hair and shiny black scales.
Notopleural stripe and margin of scutellum marked by dull golden elongate scales. Hair on
collar and pleura dull brownish gold with an admixture of black hair on the prosternum and
lower part of propleura, and also a few fine black hairs on the mesepisternum. Metapleura
bare except for the metapleural tuft. Plumula and prealar tuft pale yellowish. Macrochaetae
long, fine, black. Legs : dark reddish heavily overlaid with shiny black scales, spines and
spicules black except that the fine hairs on the fore tibiae and tarsi and the apical segments of
the other tarsi are paler yellowish, hair black except for an admixture of dull gold ones on the
hind coxae. Claws red at their bases, black apically. Fore femora unarmed, mid with four
anterolateral and two posterolateral spines and hind with a complete row of about eight postero-
lateral spines. Wings : opaque greyish, all veins broadly bordered with dark brown, so that
only the centres of the wider cells are clear, praediscoidal spot and a fine border running from
it to the base of the third basal cell along the vein, and another running along the first vein
and ending in a spot at the root of the second, grey. Venation as in Defilippia with the apical
vein of the discoidal cell sinuous and parallel with the margin but with an angular bend into
the third posterior cell, emitting an appendix on some specimens and others with an appendix
into the discoidal cell at the bend towards the base. Basal hook and comb black. Squama
yellowish grey with a yellow fringe. Abdomen : black with obscure reddish areas at the
margins of the tergites and on the sternites. First tergite and sides of second and third with
dull golden hair and scales (on the reflexed margin below they are black) , remaining sparse hair
and fringe of scales at the margins of the sclerites black. Scales on second tergite black, with
white scales at the anterior corners bordered with yellow scales which extend in a band across
the base at the middle ; third segment with the scales on the basal half yellow, darker, more
golden where they join the black scales on the posterior half ; remaining segments with inter-
mixed black and golden scales, the latter denser towards the posterior margins, tending to give

FIGS. 8-9. Exoprosopa insularis. 8, Lateral view of hypopygium ; 9, dorsal view, outline
of paramere and aedeagal accessory structure.

BOMBYLIIDAE, AND NEMESTRINIDAE FROM SOKOTRA 81

an irregular banded appearance. Sternites with sparse yellow hair and golden scales. Hypopy-
gium (Text-figs. 8 & 9), dark red with golden hair. Ovipositor with five strong red spines at
each side and sparse black hair.

Length very variable, the two males 8 mm and 10-5 mm, wings 8 mm and n mm. The
females 8 mm and 14 mm and their wings 8 and 14 mm also.

SOKOTRA : Adho Dimellus, i <$, 24. iv. 1967 (K. M. Guichard} ; Hammadero,
i c£, 2 ?, 18.^.1967 (K. M. Guichard), BM(NH) and author's collection.

Ricardo's type was caught at Jenaagahan in January and Becker (1910) reported
it from Sokotra in January and February.

Family NEMESTRINIDAE
Atriadops cinnamonea Brunetti

Atriadops cinnamonea Brunetti, 1929 : 5.

This species was described from Malawi (Nyasaland) and the only other record
is a single male from Tanzania, Ilonga, at light, 20.11.1965 (/. A. D. Robertson),
the hypopygium of which has been illustrated by Greathead (19676).

SOKOTRA : Hammadero, noo ft, 2 <$, 18.^.1967 (K. M. Guichard), BM(NH)
and author's collection.

REFERENCES

BECKER, T. 1910. Dipteren aus Siidarabien und von der Insel Sokotra. Denkschr. Akad.

Wiss. Wien. 71 : 131-160, 4 figs.
BEZZI, M. 1921. On the Bombyliid fauna of South Africa (Diptera) as represented in the

South African Museum. Ann. S. Afr. Mus. 18 : 1-180, 2 pis.

— 1924. Bombyliidae of the Ethiopian Region, viii -)- 390 pp., 46 figs. London.
BOWDEN, J. 1964. The Bombyliidae of Ghana. Mem. ent. Sac. sth Afr. 8 : 159 pp., 145

figs.

BRUNETTI, E. 1929. New African Diptera. Ann. Mag. nat. Hist. (10) 4 : 1-35.
ENGEL, E. O. 1932-37. Fliegen palaearkt. Reg. 25 Bombyliidae. IV, 3. 619 pp., 15 pis.,

239 figs.

FABRICIUS, J. C. 1805. Systema Antliatorum. 372 + 30 pp. Brunswick.
GREATHEAD, D. J. 1958. Observations on two species of Systoechus (Diptera : Bombyliidae)

preying on the desert locust, Schistocerca gregaria (Forskal), in eastern Africa. Entomo-

phaga 3 : 3-22, 29 figs., i photo.

— 19670. The Bombyliidae (Diptera) of northern Ethiopia. /. nat. Hist. 1 : 195-284,
i map, 96 figs.

— 19676. The genus Trichopsidea Westwood, with a discussion of its relation to other
genera of Nemestrinidae (Diptera). /. nat. Hist. 1 : 305-313, 10 figs.

— (In Press). Notes on Bombyliidae (Diptera) from the southern borderlands of the Sahara
with descriptions of new species. /. nat. Hist.

HESSE, A. J. 1938. A revision of the Bombyliidae (Diptera) of southern Africa. (Part I)
Ann. S. Afr. Mus. 34 : 1053 pp., 332 figs.

82 D. J. GREATHEAD

HESSE, A. J. 1956. A revision of the Bombyliidae (Diptera) of southern Africa. Parts II

and III. Ann. S. Afr. Mus. 35 : 972 pp., 2 pis., 286 figs.
HYNES, H. B. N. 1947. Observations on Systoechus somali (Diptera Bombyliidae) attacking

the eggs of the desert locust (Schistooerca gregaria (Forskal)) in Somalia. Proc. R. ent.

Soc. Lond. (A) 22 : 79-85, 3 figs.
KLUG, F. 1832. In EHRENBERG. Symb. phys. Dec. 3. No. 12. Anthrax. 5 pp., pi. xxx.

Berlin.

LINNAEUS, C. 1758. Sy sterna Naturae. Vol. I. loth Ed. 824 pp. Stockholm.
LYNEBORG, L. 1965. A revised list of Danish Bombyliidae (Diptera), with a subspecific

division of Villa circumdata Meig. Ent. Meddr 64 : 155-166, 13 figs.
MEIGEN, J. W. 1820. Systematische Beschreibung der bekannten Europdischen zweiflugeligen

Insekten. Vol. 2. x + 365 pp., pis. 12-21. Aachen.
OLDROYD, H. 1947. A new species of Systoechus (Diptera : Bombyliidae), bred from eggs

of the desert locust. Proc. R. ent. Soc. Lond. (B) 16 : 105-107, 2 figs.
POPOV, G. B. 1957. The vegetation of Socotra. /. Linn. Soc. Lond. (Bot.) 55 : 706-720,

i map, 1 8 figs.
RICARDO, G. 1903. The flies of Sokotra. In FORBES, H. O. The Natural History of

Sokotra and Abd-el-Kuri, pp. 359-378, i pi. Liverpool.

DAVID JOHN GREATHEAD, B.Sc., A.R.C.S., Ph.D.,

Entomologist-in-Charge,

EAST AFRICAN STATION,

COMMONWEALTH INSTITUTE OF BIOLOGICAL CONTROL,

c/o P.O. Box 7065,

KAMPALA, UGANDA.

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2. NIXON, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera :
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THYSANOPTERA FROM THE
SOLOMON ISLANDS

L. A. MOUND

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 24 No. 4

LONDON : 1970

THYSANOPTERA FROM THE
SOLOMON ISLANDS

BY

LAURENCE ALFRED MOUND

Pp. 83-126 ; i Map, 43 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 24 No. 4

LONDON : 1970

THE BULLETIN OF THE BRITISH MUSEUM

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

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

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

This paper is Vol. 24, No. 4, of the Entomological
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those of the World List of Scientific Periodicals.

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

Trustees of the British Museum (Natural History), 1970

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 22 January, 1970 Price £i 6s.

THYSANOPTERA FROM THE
SOLOMON ISLANDS

By L. A. MOUND

CONTENTS

Page
INTRODUCTION .......... 85

THRIPIDAE ........... 89

PHLAEOTHRIPIDAE .......... 90

PHLAEOTHRIPINAE .......... 90

MEGATHRIPINAE . . . . . . . . . .116

REFERENCES ........... 124

INDEX ............ 125

SYNOPSIS

Thirty species of Thysanoptera are recorded from the Solomon Islands. Nine of these
species are widespread across the world, two are widespread in the Western Pacific, two are
known from other parts of Melanesia, and seventeen are known only from the Solomons.
Fifteen new species are described, four new genera are defined, and seven new synonymies
are established. The new species were collected in leaf-litter and a discussion is included on
the structure, collecting methods and distribution of leaf-litter species.

INTRODUCTION

ONLY two species of Thysanoptera have been described from the Solomon Islands
(Euoplothrips carcinoides Hood, 1937 and Mecynothrips snodgrassi Hood, 1952), and
the present author has seen published records of only three other species from this
area (Haplothrips priesnerianus Bagnall in Mound, 1968, and Thrips tabaci
Lindemann and Selenothrips rubrocinctus (Giard) in Lever, 1968). In the present
account a further twenty five species are recorded, of which fifteen are described as
new, and notes are included of fragmentary material of several other species which
have also been seen during the course of this work.

Most of the material discussed here was extracted from leaf-litter on the ground
through berlese funnels, either by the government entomologist, Dr. John Greenslade
(P.J.M.G.), or by Mr. P. N. Lawrence (P.N.L.) who visited the Islands with the
Royal Society Expedition in 1965, or by collectors from the Bernice P. Bishop
Museum, Hawaii. A single gall containing very large numbers of a single species
was collected by Mr. J. Grant on the Royal Society Expedition, and three species
were submitted for identification to the Commonwealth Institute of Entomology by
the Solomon Islands Department of Agriculture. Unless stated to the contrary the
material referred to in this paper is deposited in the British Museum (Natural
History).

Thrips were extracted from moss, soil, leaf-litter and plant debris through eight-
inch diameter plastic funnels. Several samples were taken at each site and these
were placed in funnels beneath electric lights. The insects dropped into tubes of

86

L. A. MOUND

MAP i. The Solomon Islands (N.B., the Florida Islands are also known as the Nggela Group).

THYSANOPTERA FROM SOLOMON ISLANDS 87

alcohol pushed on to the bottoms of the funnels. The extractions were carried out
at Kukum Agricultural Research Station, near Honiara, Guadalcanal, not at the
various collecting sites, and Dr. Greenslade has suggested that at least some of the
species were attracted either to the lights or to the alcohol of the funnels. This is
probably true of some thripid specimens, but most of the Phlaeothipidae recorded
here are apparently leaf -litter dwelling species and some were accompanied by larvae
and apterae.

Because most of the species discussed here were extracted from leaf -litter, it is
not possible to discuss the thrips fauna of the Solomon Islands in general terms.
The few species known from surrounding areas were mainly collected from flowers or
green leaves or dead wood, and these habitats have not been examined for thrips in
the Solomons. Conversely the thrips of leaf -litter are unknown from Fiji and New
Guinea, although some species have been described from Indonesia. However
Moulton (1944) does not record the genera Ecacanthothrips, Machatothrips and
Mecynothrips from Fiji, and in view of the large size of the species and the frequency
with which they have been taken in the Melanesian area, it would be reasonable to
expect collectors to find them if they were present. These forms may not extend
across the Pacific, although Rhaebothrips is known from Formosa to Samoa and
Eastern Australia. The apparent absence of the conspicuous flower-living Aeolo-
thripidae from the Melanesian area is rather surprising, as this family is well
developed in Australia and also occurs in New Zealand, New Caledonia and Southern
India. Similarly no Merothripidae were taken in the Solomon Islands, although
these inconspicuous thrips are readily taken in leaf-litter in Australia and also
occur in New Zealand.

Some species that live in leaf-litter have a rather unexpected geographical range.
Bradythrips hesperus is now known from Guyana, Southern India and the Solomon
Islands, and species of the genus Psalidothrips are known from Java, the Solomon
Islands and Brazil. The author has recently collected unidentified species of
Allothrips and Pseudocryptothrips in Australia and both these genera occur in
North America. These distributions may be natural but could be the result of
man's activity. Leaf-litter thrips feed on the fungi associated with dead leaves and
it seems likely that such fungi have extensive ranges. Unlike phytophagous
insects which are limited by the range of their host plants, these fungus-feeding
thrips have a continuous habitat wherever there are dead leaves and a suitably
drained soil. However it is likely that man has also influenced the distribution of
some species during the last three or four hundred years. The present author has
commented elsewhere (1968 : 140) on the possibility that Nesothrips propinquus
has been distributed around the Southern Hemisphere in hay used on board ships,
and it is suggested below that Hoplandrothrips flavipes has similarly been distributed
in firewood on ships. Lindroth (1957) in an account of the faunal relationships of
North America and Europe has produced evidence to show that some species,
particularly of carabid beetles, have been distributed by man in the soil and gravel
used as ballast on sailing ships. Similar studies have not been made in tropical
regions but ballast was carried by all sailing ships in varying quantities in order to

88 L. A. MOUND

maintain vessels at a suitable level in the water, thus more ballast was needed with
a light cargo than a heavy cargo. Captain Cook records loading the Endeavour
with eight tons of ' iron ballast ' before leaving England in 1768, but he also records
taking on twenty tons of ' stone ballast ' at Tahiti, and unspecified amounts in New
Zealand, Eastern Australia and Java (Beaglehole, 1955). Such stone ballast
probably included quantities of soil and plant debris. In North Atlantic ports
ships were not permitted to throw ballast into the water, as it could have blocked
the harbour, but had to offload it on to the quayside. Similar regulations were
probably in operation in tropical ports. Once the ballast was on the quay, it was
then available to the next ship, so creating the opportunity for organisms to be
moved in ballast from port to port. Although European settlers and plant collectors
undoubtedly moved very large numbers of tubs of soil containing plants from place
to place, this traffic in ships' ballast is probably more important to zoogeographers
because of its sheer bulk. Lindroth (1957 : 161) states that according to the records
of Poole Harbour in Southern England, 1180 tons of ballast were supplied to ships
sailing from Poole to Newfoundland in the year 1815. With the vast tonnage of
sailing vessels in operation during the eighteenth and nineteenth centuries there
were thus ample opportunities for soil-living organisms to be moved around the
world.

There are several morphological characters which seem to be associated with the
leaf-litter habitat in the Phlaeothipinae. The species of Baphikothrips, Psalido-
thrips, Mystrothrips and Solomonthrips are bicoloured, usually yellow with one or
more transverse brown markings. The wings of most of these species are narrow,
not constricted medially, with widely spaced cilia and few or no accessory cilia.
Baphikothrips, Psalidothrips and Solomonthrips species have a narrow bell-shaped
pelta as in Adraneothrips and Hoplandrothrips species. Surface reticulation is
frequent in leaf -litter species such as Solomonthrips, Mystrothrips and the Glypto-
thripini, but it is not restricted to such species nor are all leaf -litter species reticulate,
e.g. Psalidothrips. An interesting negative characteristic is the apparent absence of
oedymerous forms in these species. Fungus-feeding Phaleothripinae living under
bark frequently show a very great range of body form, e.g. Ecacanthothrips. The
leaf-litter species are also fungus-feeders but none of the species referred to here are
known to produce strongly oedymerous individuals. The major males of Solomon-
thrips greensladei described below have larger femora with tubercles than the minor
males but they do not show any great difference in body size.

Most of the specimens referred to in this paper were treated briefly with 5%
sodium hydroxide solution prior to dehydration and mounting in balsam. This
treatment facilitates clearing but destroys the hypodermal pigments, and so the
notes on colours of the species refer only to cuticular colour. The head lengths
which are quoted are based on measurements of total head length from the base of
the head to the interantennal projection. The text-figures were drawn at various
magnifications, the heads and pronota using a Wild drawing tube and the remainder
using a Zeiss camera lucida. Mr. B. R. Pitkin drew text-figures 10-12, 24, 25 and
28-32, and the others were drawn by the author. This paper could not have been

THYSANOPTERA FROM SOLOMON ISLANDS 89

completed without the advice and generous loan of material from Dr. H. Priesner
of Linz, Miss Kellie O'Neill of the U.S.D.A., Washington, and Dr. Paul Arnaud of
the California Academy of Sciences.

THRIPIDAE

Included among the few thripids taken from leaf-litter were two female Scirto-
thrips from Mt. Austen, Guadalcanal, one female Pseudodendrothrips from New
Georgia, and four females and one male of the Thrips /Taeniothrips group from
Guadalcanal and Kolombangara. The condition of these specimens was too poor
for accurate study below the generic level at the present state of knowledge of the
Melanesian fauna.

Chirothrips spiniceps Hood

Chirothrips spiniceps Hood, 1915 : 12-15.
Chirothrips spiniceps Hood ; zur Strassen, 1960 : 175.

This species is probably established in the Solomon Islands. It is known from
North America, Mexico and Hawaiian Islands according to zur Strassen.

Material studied. GUADALCANAL : on Rice, i $, xi-xii.igGs and i $, 4.xii.i965
(M. McQuillan) ; on IBrachiaria miliiformis, i $, 17^.1966 (M. McQuillan}.

Microcephalothrips abdominalis (Crawford)

Thrips abdominalis Crawford, 1910 : 157-159.

This species is widely distributed in the tropics on composite flowers, and may be
a pest on sunflowers.

Material studied. GUADALCANAL : Kukum, i $, I4.vii.ig66 (P.J.M.G.) ; Mt.
Popanamisiu, i $, x.ig65 (P.J.M.G.).

Plesiothrips perplexus (Beach)

Sericothrips ? perplex a Beach, 1896 : 216.

Plesiothrips perplexus (Beach) ; Stannard, 1968 : 333-337.

The postocellar chaetotaxy of the specimens listed here compares favourably
with American specimens collected on grass in Washington, B.C. The species is
widespread on grasses ; the author has collected it in Eastern Australia, but
members of the genus need careful study as indicated by Stannard.

Material studied. GUADALCANAL : on Rice, 2 $, xi-xii.i965 ; i $, 3.xii.i965 ;
i $, 4.xii.i965 (M. McQuillan] ; on Eleusine indica, i $, 14^.1966 (M. McQuillan).

Selenothrips rubrocinctus (Giard)
Physopus rubrocinctus Giard, 1901 : 263-265.

The red-banded cocoa thrips is spread throughout the tropics as a pest on the
leaves of cocoa trees.

go L. A. MOUND

Material studied. GUADALCANAL : Mt. Austen, i <j>, ^.1966 (P.J.M.G.).
RENDOVA : 10 larvae on Cacao leaves, 1957 (E. S. Brown).

Thrips tabaci Lindemann
Thrips tabaci Lindemann, 1888 : 61-75.

This cosmopolitan species is recorded from the Solomon Islands by Lever (1968 :
8) as a result of thrips having been seen on onions. No material has been studied.

Thrips unispinus Moulton
Thrips (Epithrips) unispinus Moulton, 1940 : 252.

The type specimens of this species have not been studied but the male and female
listed below agree well with the description based on material from Koitaki, New
Guinea.

Material studied. GUADALCANAL : Mt. Austen, i $, i <$, I3.xi.ig64 (P.J.M.G.).

PHLAEOTHRIPIDAE

In addition to the species which are discussed below under the two subfamilies
Phlaeothripinae and Megathripinae, a number of rather poor specimens have been
seen which cannot be adequately distinguished. In the author's opinion, no useful
purpose is served by describing species on inadequate material in unrevised genera
which are known to include highly variable species. Two specimens of undescribed
Androthrips and fifteen unidentified Haplothrips have been seen, also single speci-
mens of Adraneothrips, Dicer atothrips, Horistothrips and Karnyothrips. Of more
interest, but still not worth describing, were three specimens of a genus between
Nesothrips and Dicer atothrips, and three specimens of a genus close to Malacothrips.

PHLAEOTHRIPINAE
BAPHIKOTHRIPS gen. n.

Type-species : Baphikothrips color atus sp. n.

Weakly sclerotized slender species with cuticle bicoloured brown and yellow. Head faintly
sculptured dorsally ; eyes large, longer on dorsal surface than ventral ; postocular setae
behind inner margin of eyes, shorter than dorsal length of eye ; cheeks weakly incut behind
eyes, without major setae ; fore ocellus directed forwards ; mouth cone long, not sharply
pointed ; stylets retracted into head, maxillary bridge present. Antennae eight-segmented,
VIII not sharply constricted at base. Pronotal setae well developed, epimeral sutures usually
incomplete ; praepectus absent ; probasisternal plates with one seta at anterior external
angle ; mesopraesternum broadly boat-shaped but weakly sclerotized. Fore tarsi unarmed,
legs slender. Lateral mesonotal setae well developed. Metanotum with longitudinal band of
reticulation, median setae wide apart. Fore wings slender, weakly constricted medially ;
cilia widely spaced, 2-4 accessory cilia ; sub-basal setae well developed. Pelta bell-shaped ;
tergites II-VII with two pairs of wing-retaining setae ; tergite IX with accessory seta well
developed between BI and Bg : tube shorter than head.

THYSANOPTERA FROM SOLOMON ISLANDS 91

This new genus resembles Baphothrips in many ways but in the opinion of the
present author this is probably superficial. Baphikothrips is very close to Adraneo-
thrips but has larger eyes, which are more extensive dorsally than ventrally, and
moreover has the postocular setae rather close together. Adraneothrips and
Baphikothrips both have a maxillary bridge, whereas in Baphothrips and Malaco-
thrips the maxillary guides approach each other at their anterior ends as in more
typical members of the Phlaeothripina (Priesner, 1960). It seems more likely that
Adraneothrips and Baphikothrips are derived from Haplothrips-like stock by the loss
of the praepectus and degeneration of the fore wings in association with their cryptic
habitat, than that they have evolved from members of the Phlaeothripini and
developed independantly a maxillary bridge. Moreover, these two genera, in
common with Haplothrips, do not have antennal segment eight sharply constricted
into a basal neck as is found in Baphothrips and Phlaeothripina such as Phlaeothrips,
Ecacanthothrips and Hoplandrothrips. For these reasons the author would place
Adraneothrips and Baphikothrips in the Haplothripini.

KEY TO THE SPECIES OF BAPHIKOTHRIPS

i Antennae very long, segment III about four times as long as wide with two sense
cones ; head brown, pronotum and fore legs yellow ; antennal III not much
paler than IV ; male not known ..... antennatus sp. n. (p. 93)
Antennae shorter, III about twice as long as wide with three sense cones ; head
yellow with brown margins, pronotum and fore legs with brown markings ;
antennal III yellow, IV brown but V and VI brown with basal third yellow ; male
with no fore tarsal tooth, sternite VIII with glandular areas . coloratus sp. n. (p. 91)

Baphikothrips coloratus sp. n.

(Text-figs. 3 & 4)

$ (macropterous) . Colour yellow with brown markings ; head and pronotum yellow with
brown margins ; mesothorax, sides of metathorax, and sides of abdominal segments II-III
and V-VI brown ; tube brown ; mid and hind femora and tibiae brown medially ; fore legs
with brown shadings along margins ; antennal III and basal third of V and VI yellow ; wings
shaded except at extreme apex and median constriction ; major setae pale.

Head with faint sculpture dorsally (Text-fig. 3) ; eyes longer on dorsal surface than on
ventral ; antennals III and IV with three sense cones (Text-fig. 4). Pronotum weakly sculp-
tured near posterior margin ; epimeral sutures usually not quite complete ; epimeral setae
little longer than other major setae. Mesonotal lateral setae about 20^1 long, expanded at
apex ; metanotal setae pointed but not acute, 25|x long, 40^ apart. Fore wing narrow, maximum
width 55(jt, weakly constricted medially, distal width 35^1 ; cilia widely spaced, two to four
accessory cilia ; sub-basal setae in a straight line, 30^, 30(0. and 45^1 long with expanded apices.
Pelta bell-shaped, faintly sculptured ; seta BI on tergites broadly expanded at apex, more
than 50(1 long ; tergites with faint sculpture laterally. Sternites with transverse row of about
10 accessory setae SJJL long.

Measurements in microns of holotype with range from four paratypes in parentheses. Body
length 1300 (1150-1400). Hind tibia 135 (120-). Fore wing 580 (510-). Head, length 175
(160-) ; width 160 (150-) ; postocular seta 32 (26-35). Pronotum, length 97 (88-100) ;
width 190 (180-200) ; epimeral seta 35 (32-40). Tergite IX, BI 55 (49-) ; accessory seta 42 ;
B2 58 (49-). Tube, length 97 (88-) ; terminal setae 80 (70-). Antennal segments length
26 (23-) ; 35 (-39) ; 52 (49-) ; 52 (49~) ', 42 (39-) ; 39 ; 26 (-29).

L. A. MOUND

FIGS. 1-7. Figs, i & 2. Baphikothrips antennatus : i, Head and pronotum (sculpture
omitted from pronotum). 2, Left antenna. Figs. 3 & 4. Baphikothrips coloratus :
3, Head. 4, Right antenna. Figs. 5-7. Pelta of Ecacanthothrips species : 5, spinipes
gynaecoid female. 6, spinipes oedymerous female. 7, sanguineus.

THYSANOPTERA FROM SOLOMON ISLANDS 93

Holotype <j>. CHOISEUL : Vasu River, iG.xi.igGs (P.J.M.G.}, collected with
14 $ paratypes.

Other material. FLORIDA Is. : Vatilau, 18 <j>, 2.xii.i965 (PJM.G.}. GUADAL-
CANAL : Mt. Austen, i $, 8.11.1966, i $, 11.11.1966 (PJM.G.} ; Mt. Popanamisiu,
at 7000 feet in moss forest, i °-, 6.xi.i965 (P.N.L.).

The bicoloured antennae and the weak fore wings of this species are similar to
Adraneothrips species, but very large eyes are not found in that genus except when
the eyes are extended on the ventral surface.

Baphikothrips antennatus sp. n.

(Text-figs, i & 2)

$ (macropterous) . Bicoloured ; head and antennal segments I and II dark brown ; lateral
sclerites of ptero thorax, abdominal segment IV and tube brown ; anterior margin of abdominal
segment V, posterior margin of III, and segments VIII and IX light brown ; pronotum and
fore legs yellow ; middle femora dark brown in basal half, hind femora and middle and hind
tibiae largely yellow ; wings shaded except at median constriction, dark around sub-basal
setae, cilia rather pale ; major setae pale except on dark sclerites ; antennal segments III
and IV light brown, V-VIII darker but V and VI with basal stem yellow.

Head rather long with faint sculpture dorsally (Text-fig, i) ; eyes larger on dorsal surface
than on ventral ; antennal segments very long, sense cones small, two sense cones on III and V,
three on IV (Text-fig. 2). Pronotal epimeral sutures not complete (Text-fig, i) ; mesonotal
lateral seta 30^ long with expanded apex ; metanotal setae finely acute, 35[x long, 6o(z apart.
Legs long and slender. Fore wing narrow, maximum width 6o(i, weakly constricted medially,
distal width 401*, cilia widely spaced, 2-4 accessory cilia ; sub-basal setae in straight line,
45ji, 5O[ji and yojj. long. Pelta narrow and bell shaped ; tergites with very faint sculpture
laterally ; BI and 62 on IX with apices weakly expanded. Sternites transversely reticulate
with about six accessory setae 2O[i long.

Measurements in microns of holotype. Body length 1650. Hind tibia 190. Fore wing
700. Head, length 225 ; width 180 ; postocular seta 55. Pronotum, length 130 ; width 240 ;
epimeral seta 45. Tergite IX, BI 100 ; accessory seta 56 ; 62 100. Tube, length 115 ;
terminal setae 115. Antennal segments 32 ; 42 ; 100 ; 90; 95 ; 60 ; 49 ; 30.

c£ (macropterous). Colour rather paler than female, particularly the abdomen. Chaetotaxy
and sculpture similar to female but 62 on tergite IX short and stout ; sternite VIII with an
irregular pair of glandular areas laterally ; pseudovirga long and slender as in many Haplothrips
species.

Measurements in microns of allotype. Body length 1450. Hind tibia 175. Fore wing 600.
Head, length 225 ; width 150 ; postocular seta 45. Pronotum, length 120 ; width 185 ;
epimeral seta 42. Tergite IX, BI 100 ; accessory seta 65 ; 62 32. Tube 105. Antennal
segments 30 ; 38 ; 100 ; 100 ; 100 ; 65; 50; 30.

Holotype °-. SANTA YSABEL : South East Coast, 1000 Ships Bay opposite
Lillininia Island, leaf-litter on shore, 20.ix.i965 (P.N.L.).

Allotype $, and 5 °., 4 <$, 2 larvae collected with holotype.

Other material. SANTA YSABEL : Tatamba, in mangrove litter, i $, I4.X.I9&5
(P.N.L.) ; San Jorge Island, in litter in gulley, i ?, 22.ix.i965 (P.N.L.). CHOISEUL :
Malangona, i $, 4.111.1964 (P. Shanahan) in Bishop Museum Collection.

Because of the very long antennae this species shows less resemblance to Adraneo-
thrips than does coloratus.

94 L. A. MOUND

BAPHOTHRIPS Priesner
Baphoihrips Priesner, 1933 : 69-70. Type-species : B. tricolor Priesner, by monotypy.

The unique female, from soil in Java, upon which this genus is based has been
studied and compared with the new species described below. The genus may be
defined as follows : —

Cuticle bicolored, brown and yellow ; wings weakly banded. Cheeks incut behind large
compound eyes ; postocular setae long ; first ocellus directed forward between bases of
antennae ; maxillary stylets close together in middle of head, maxillary bridge absent.
Antennae long, eight-segmented, segment IV longest, VIII constricted at base ; three sense
cones on III and IV. Pronotum transverse, epimeral sutures complete ; anteroangular setae
rather close to midlaterals ; praepectus absent, mesopraesternum well developed but weakly
sclerotized. Lateral mesonotal setae well developed. Metanotum reticulate, one pair of long
setae wide apart near anterior margin. Fore wings moderately broad, weakly constricted
medially, with accessory cilia ; three sub-basal setae in straight line. Pelta weakly sclerotized ;
tergites II-VII with two pairs of wing-retaining setae and several lateral setae anterior to BI ;
tergite IX with accessory seta between BI and 62 well developed, more than half as long as BI ;
tube shorter than head.

This genus belongs in the Phlaeothripina close to Malacothrips, but most of the
species placed in that genus are not known to the present author. Baphothrips
apparently has larger, more rounded eyes, the cheeks are less constricted behind the
eyes, and the maxillary stylets are retracted further into the head. Adraneothrips
and Baphikothrips are readily distinguished by the presence of a maxillary bridge
and the weaker fore wings. Moreover these two genera do not have the eighth
antennal segment constricted into a basal neck and the pronotum is relatively
longer.

KEY TO THE SPECIES OF BAPHOTHRIPS

i Dorsal surface of head reticulate, cheeks strongly narrowed to base ; metanotum
with longitudinal band of reticulation ; mesonotal lateral setae less than half as
long as pronotal posteroangular setae ; head yellow, brown only on anterolateral
margins, mesothorax with anterior margin brown ; abdominal segments VIII
and IX yellow ......... tricolor Priesner

— Dorsal surface of head not reticulate, cheeks weakly narrowed to base ; metanotum
weakly reticulate ; mesonotal lateral setae long, more than half as long as pronotal
posteroangulars ; head and abdominal segments VII-IX brown, anterior margin
of mesothorax yellow ........ leios sp. n. (p. 94)

Baphothrips leios sp. n.

(Text-figs. 8-10)

$ (macropterous). Bicoloured ; pterothorax, abdominal segments V-VI, all tibiae and
tarsi yellow ; dark brown on head and pronotum, antennal segments I and II, tube and
abdominal segments VII-IX, II-III and anterior margin of IV ; femora shaded brown medially,
fore femora darkest ; antennal segments I II-VII I light brown ; major setae light brown ;
basal half of wing darker than distal half, cilia dark.

Head weakly sculptured laterally and between ocelli ; postocular setae longer than eyes
(Text-fig. 9) ; maxillary stylets retracted deeply into head, close together medially, maxillary

THYSANOPTERA FROM SOLOMON ISLANDS

95

\

n

\

10

FIGS. 8-12. Figs. 8-10. Baphothrips leios : 8, Left antenna. 9, Head and pronotum.
10, Tergite IV. Figs, n & 12. Mystrothrips dilatus : u, Left antenna. 12, Tergite
III.

96 L. A. MOUND

guides well developed, bridge absent ; ventral surface of head with a pair of long setae basally
and another pair between the eyes ; antennae long, sensorium on II near apex (Text-fig. 8) ;
mouth cone extending two thirds across prothorax, acute but with broadly rounded labrum.

Pronotum short and wide (Text-fig. 9) ; probasisternal plates well developed with five
setae along anterior margin. Mesonotal lateral setae 50(0. long, apex expanded. Metanotal
setae finely acute, 65^ long, IOO(JL apart. Fore legs long, without armature. Wing broadest
(8o[x) just beyond sub-basal setae, weakly constricted medially, with seven accessory cilia ;
sub-basal setae long (65(0. ; 8o[i ; IOOJJL), expanded apically.

Pelta weakly sclerotized, reticulate, broadly bell-shaped ; tergite II with an irregular group
of 12 or more setae laterally anterior to BI ; setae in this position on tergites III-VIII in an
irregular transverse row (Text-fig. 10), but posterior tergites with fewer setae (3) than anterior
(8) ; BI and ~Bz on IX very weakly expanded at apex ; sternites with transverse row of about
24 rather long fine setae (40^).

Measurements in microns of holotype with one paratype in parentheses. Body length
2200 (1750). Hind tibia 260 (245). Fore wing 850 (700). Head, length 275 (260) ; width
210 (190) ; postocular seta 105 (90). Pronotum, length 130 (125) ; width 320 (300) ; epimeral
seta 84 (78). Tergite IX, BI 155 (155) ; accessory seta 105 (105) ; 62 160 (160). Tube,
length 180 (160) ; terminal setae 160 (145). Antennal segments length 42 (35) ; 52 (49) ;
84 (84) ; 80 (80) ; 100 (97) ; 80 (80) ; 65(65) ; 49 (45) ; terminal seta 60 (60).

Holotype $. GUADALCANAL : Forest litter in root mat, 5.vii.ig65 (P.N.L.).

Other material. GUADALCANAL : near Honiara, Poha River, grasses and weeds,
i ?, I9.xi.ig65 (P.N.L.) ; Mt. Austen, i $, 2i.ix.ig65 (P.J.M.G.).

Bradythrips Hesperus Hood & Williams

Bradythrips hesperus Hood & Williams in Hood, 1925 : 68-69.
Bradythrips hesperus Hood & Williams ; Hood & Williams, 1927 : 4-5.
Bradythrips hesperus Hood & Williams ; Ananthakrishnan, 1966 : 5.

Type material of this species has not been studied but the specimens listed below
compare closely with the published descriptions. All the specimens from the
Solomon Islands however are macropterous, whereas the original specimens from
Guyana and the recently collected ones from Quilon in Southern India are all
apterous. At the apex of the tube there is a pair of fine dorsal setae which are less
than one quarter as long as the four major anal setae, and laterally on the meta-
thorax there is a large seta on a tubercle.

Material studied. GUADALCANAL : Mt. Austen, beaten from dead sticks on
ground, 9 $, ii.ii.ig66, and 2 $ with 8 larvae, 8.ii.ig66 (P.J.M.G.). SAN CRISTOVAL :
forest litter, i $, 5.viii.ig65 (P.N.L.) .

Ecacanthothrips spinipes (Bagnall)
(Text-figs. 5 & 6)

Phloeothrips spinipes Bagnall, 1908 : 195-196.
Ecacanthothrips spinipes (Bagnall) Mound, 1968 : 90.
Ormothrips inermis Buff a, 1909 : 168. Syn. n.
Ecacanthothrips bagnalli Priesner, 1930 : 364-365. Syn. n.
Ecacanthothrips guineaensis Moulton, 1947 : 176-177. Syn. n.

Bagnall described the unique holotype of this species, which was dry on a card
point, as having yellow antennae. This specimen has now been mounted into

THYSANOPTERA FROM SOLOMON ISLANDS 97

balsam and not only are the distal antennal segments brown but also segments three
and four have light brown shadings near their apices. The antennae of inermis were
also described as being yellow. Buffa's unique holotype has not been studied but
the description does not separate it from spinipes, and as this species is apparently
common in New Guinea, inermis is here regarded as a synonym. The unique
holotype of bagnalli, which is a medium-sized male, and the type series of guinea-
ensis, three very small females, have been studied and compared with the material
listed below from New Guinea and the Solomon Islands.

This species can be distinguished from sanguineus Bagnall, the type-species of the
genus, by the unarmed fore femora of the female and the broader reticulations
laterally on the pelta (Text-figs. 5-7). The male has two subapical fore femoral
teeth, a small and variable dorsal one overlaying the larger ventral tooth (cf. fig. in
Priesner, 1930). However the species varies considerably in size, even within a
population, and the smallest individuals can appear very different from the larger
ones, as is known in sanguineus. In large females the fore femur bears a series of
stout thorn-like setae on the inner dorsal surface, the fore tibia has about five
tubercles on the inner margin, the sense cones on antennal segment three are dark,
and the pelta has broad lateral reticulations. Medium-sized females are similar to
the large ones, but very small females have no stout setae on the fore femur, no
tubercles on the fore tibia, pale sense cones on the antenna, and the lateral margins
of the pelta are eroded into small chitinous islets (Text-fig. 5).

There are four other nominal species of Ecacanthothrips with unarmed fore femora
in the females. Neither piceae Ishida nor inarmatus Kurosawa from Japan have
been studied, although according to the descriptions they are very similar to each
other. The unique holotype of leai Moulton from Malaya, which is a very small
female, has been compared with a rather large female paratype of coniger Priesner
from Borneo, but because of the size-difference and the poor state of the leai holo-
type, it is not possible to state how these species differ. From spinipes they differ
in having the third antennal segment brown.

Material studied. Holotype $. NEW GUINEA : Dorey (Wallace}.

NEW GUINEA : Holotype $ and 2 $ paratypes of guineaensis, with i °-, I ^ labelled
as bagnalli by Moulton, Finschhafen, on bark, i6.iv.i944 (E. S. Ross) ; Mafnn Bay,
on bark, 3 $, i $, ix.iQ44 (E. S. Ross) ; Mt. Lamington, Owen Stanley Range at
1500 feet, 2 $ (C. T. McNamara), all in Californian Academy of Sciences collection ;
north of Wau, Bulolo Gorge at 900 m, bark of fallen log, 12 $, i <$, 4 larvae (with
sanguineus), 4^.1968 (R. Rice via F. Bianchi) ; Aiyura, on grasses at 5400 feet,
i °., 22.^.1962 (J. H. Barrett}.

?KEI ISLANDS : Holotype <$ of bagnalli, in Dr. Priesner's collection.

SOLOMON ISLANDS : Guadalcanal, Mt. Austen, under bark, 4 $, 2 $, 24.xi.i965
(P.J.M.G.) ; Tambeluse, in camp kitchen, i $, io.xi.ig65 (P.N.L.).

Euoplothrips crassipes Hood

Euoplothrips crassipes Hood, 1937 : 599~6°4-

Species of this genus have a stout curved process on the inner margin of the fore

98 L. A. MOUND

femora. These thrips cause the leaves of their host plants to roll along the margins,
and the species are recorded from Samoa, Tonga, Solomon Islands, Eastern Australia,
New Guinea and India.

Material studied. SOLOMON ISLANDS : Tulagi, Big Florida Island, in rolled-up
leaf, i $ paratype, i8.iii.i936 (R. A. Lever).

Haplothrips gowdeyi (Franklin)
A nthothrips gowdeyi Franklin, 1908 : 724.

This species is very widely distributed on flowers in the tropics, including the
Pacific islands, and it probably breeds in the flowers of grasses.

Material studied. GUADALCANAL : Kukum Agricultural Station, on grasses and
flowers, 2 $, 2.xi.i965 (P.N.L.).

Haplothrips priesnerianus Bagnall

Haplothrips priesnerianus Bagnall, 1933 : 327—328.
Haplothrips priesnerianus Bagnall ; Mound, 1968 : 114.

This species appears to be widespread on grasses and other Gramineae in the
tropics and subtropics. It has been recorded from Sudan, Libya, Pakistan, India,
and the Solomon Islands.

Material studied. GUADALCANAL : on rice, 26 $, g<$, xi-xii.i965 (M. McQuillan) ;
Ilu, on ? Brachiaria mutica, n $, 17 and 27^.1965 (M. McQuillan) ; Kukum
Agricultural Station, on grasses and flowers, i $, i <$, 21. xi. 1965 (P.N.L.) ; Mt.
Popanamisiu, i $, x.ig6^ (P.J.M.G.) ; Mt. Austen, i $, 24.viii.i965 (P.J.M.G.).

Hoplandrothrips flavipes Bagnall

Hoplandrothrips flav ipes Bagnall, 1923 : 628-629.

Phloeothrips gracilicornis Priesner, 1927 : 72-73. Syn. n.

Phlaeothrips claratibia Moulton, 1937 : 4M- Syn. n.

Phloeothrips indicus Ramakrishna & Marghabandu, 1939 : 43-44. Syn. n.

Phlaeothrips (Hoplandrothrips) flavitibia Moulton, 1944 : 300-302, regarded as a synonym of

indicus R. &. M. in Ananthakrishnan, 1964 : 101—104.
Hoplandrothrips flavipes Bagnall ; Mound, 1968 : 120.

The holotype of gracilicornis from Nigeria has not been examined but the material
listed below under this name was determined by Dr. Priesner. The characters
given by Moulton to distinguish the Fijian species flavitibia from flavipes are not
correct, and although the types have not been studied, the synonymy given by
Ananthakrishnan is accepted here. Specimens determined by Dr. Ananthakrishnan
as indicus have been studied but not the unique holotype of that name. Moulton
(1947 : 175) has recorded a female of claratibia from Finschhafen, New Guinea ;
Bianchi (1953 : 106) records both sexes from Samoa, and the species is here recorded
for the first time from the Solomon Islands, Sumatra, and Malaya. There is no
record of the species from the Neotropics but it is now known from Hawaii to West

THYSANOPTERA FROM SOLOMON ISLANDS 99

Africa. It probably feeds on fungus growing on dead wood and may well have been
distributed round the world on the wood stored in ships as firewood. The Indian
specimens listed below have fewer accessory wing cilia (6-8) than African or Solomon
Islands specimens (10-12), and Moulton states that flavitibia has sixteen. The
tubercle at the apex of the fore femur is not developed in very small males.

The species is placed in Hoplandrothrips rather than Phlaeothrips because the fore
wings are weakly constricted medially. The antenna with the four fat sense cones
on segment three has been figured by Ananthakrishnan, Priesner and Mound, and
the following notes are intended to amplify the other descriptions.

Colour brown, median abdominal segments paler ; all tibiae, tarsi and femoral apices yellow,
extreme base of mid and hind femora pale ; major pronotal setae dark ; wings weakly shaded,
cilia dark ; antennals III, IV, V and sometimes VI shaded apically but much paler in teneral
specimens. Head reticulate ; postoculars long, broadly expanded at apex ; vertex with about
six pairs of fine submedian setae, cheeks with one pair of stout sub-basal setae ; stylets close
in centre of head. Antennal III with four fat sense cones ventrally, IV with four stout sense
cones. Pronotum reticulate near posterior margin particularly in large individuals ; pronotal
setae long, expanded ; praepectus absent, mesopraesternum absent medially ; female with
minute fore tarsal tooth, male with large tooth. Lateral mesonotal setae well developed ;
metanotum reticulate. Pelta bell-shaped ; abdominal tergites with two pairs of wing-retaining
setae ; tergite IX with BI and 62 more than two thirds as long as tube, weakly expanded at
apex ; terminal setae as long as tube.

Material studied. Holotype °- [KENYA: Kijalie, Kikuyu Escarpment, xii. 1911
(Alluaud & Jeannel)] 27.

GUINEA : Conakry, on Kola nuts, 27 $, 2 $, determined by Priesner as gracili-
cornis, xii. 1898 (Maclaud).

HAWAIIAN ISLANDS : Oahu, Kipapa, in wind trap, holotype $ of daratibia,
26.^.1934, in California Academy of Sciences collection.

INDIA : Madras, on palm leaf sheath, i $, 23.x. 1960 ; on coconut sheath, i $,
i (£, 8.iv.i964 (Ananthakrishnan).

SUMATRA : Lampongs, i $, 25. xi. 1921 (Karny).

MALAYA : Pahang, on Palaquium guiia, i $, I4.ix.i925 (Corbett).

SOLOMON ISLANDS : Kolombangara, 30 m, Pepele, i £, i <£, 13.^.1964, and i $,
1 1. 11.1964 (P. Shanahan), in Bishop Museum collection, Hawaii ; San Cristoval,
7 miles South of Wainoni, in forest leaf -litter, i $, 26.^.1965 (P.N.L.).

MYSTROTHRIPS Priesner

Mystrothrips Priesner, 1949 : 117, Type-species : Sagenothrips dammermanni Priesner, 1933,

by monotypy.
Mystrothrips Priesner ; Stannard, 1955 : 92-93.

The following redefinition of this genus is based on a paratype of dammermanni
from Java and the new species described below. According to Stannard, clavatoris
Hood from Brazil, the only other species in the genus, has knobbed setae on the
antennae and legs. The present author has taken at least one species of this genus
in leaf-litter in Queensland, Australia.

ioo L. A. MOUND

Body strongly reticulate, all major setae broadly expanded except on antennae. Head
longer than broad, concave at posterior dorsal margin, deeply incut behind eyes ; postocular
setae short and expanded ; one pair of postocellar and mid-dorsal setae present ; cheeks
tapering slightly to base of head, each with about five fine recurved setae ; maxillary stylets
wide apart, retracted about halfway into head ; antennae eight-segmented, segments sculp-
tured, three sense cones on III and IV, terminal seta longer than VIII. Pronotum reticulate,
epimeral sutures complete ; anteroangular and midlateral setae about as far apart as the
length of one seta ; praepectus very weak but apparently present ; mesopraesternum weakly
sculptured, broadly boat-shaped ; fore tarsal claw slender, slightly curved, length equal to
half the tarsal width. Mesonotum with lateral setae well developed, broadly expanded.
Median setae of metanotum slender ; three pairs of basal wing setae with expanded apices.
Pelta broad, reticulate, withdrawn into concave anterior margin of tergite II. Tergites II-VII
with two pairs of sigmoid wing-retaining setae ; BI and 62 on IX long with broad round apices,
BS acute ; tube faintly sculptured with overlapping scales, terminal setae shorter than tube ;
sternites with a row of about twelve accessory setae, marginal seta BI shorter than 62.

Mystrothrips dilatus sp. n.

(Text-figs, ii & 12)

$ (micropterous) . Colour yellowish, shaded brown on tergite II, at sides of head, apex of
tube, and antennal segments I, II, VI-VIII and apical half of V. Head very similar to dammer-
manni (see fig. in Stannard, 1955 : 101); postocellar setae fan-shaped, little smaller than
postoculars ; sensorium on antennal II near apex (Text-fig, n). Tergites and sternites fully
sculptured (Text-fig. 12) ; tube with faint sculpture, like overlapping scales.

Measurements in microns of holotype. Body length 1900. Hind tibia 160. Fore wing
180. Head, length 210 ; width 1 60 ; postocular seta 20. Pronotum, length no ; width 270 ;
epimeral seta 20 long, 13 wide at apex. Tergite IX, BI no ; 62 130. Tube, length 135 ;
terminal setae 80. Antennal segments : 42 ; 50 ; 70 ; 68 ; 68 ; 58 ; 42 ; 42.

Holotype $. KOLOMBANGARA : near Kuzi, 1500 ft., leaf -litter in mossy wood,

Although this new species resembles dammermanni not only in the characters
given in the generic definition but also in the colour pattern, it can be distinguished
by the shorter, more broadly expanded, fan-shaped setae particularly on the head
and pronotum. In dammermanni the postocellar setae are small with acute apices
and the prothoracic epimeral setae about three times as long as broad apically.

PSALIDOTHRIPS Priesner

Psalidothrips Priesner, 1932 : 61-62. Type-species : P. amens Priesner, by monotypy.

This genus was erected for the species amens from Java, which was based on a
single female, and the author is grateful to Dr. Priesner for the loan of this holotype.
The following definition of the genus is based on a comparison of amens with the
two new species from the Solomon Islands described below, but the other four
species in Psalidothrips, described by Hood (1955) from Belem, Brazil, have not
been studied.

Slender, weakly sclerotized, bicoloured thrips. Head as broad as long, cheeks rounded,
narrowed to base, eyes well developed ; ventral surface with only one pair of long setae, arising
between tentorial pits not at base of head ; postocular setae long, close to eye, postocellars

THYSANOPTERA FROM SOLOMON ISLANDS 101

about as long as ocellar traingle ; maxillary stylets not deeply retracted into head, without
maxillary bridge ; maxillary palps very small, scarcely longer than labial palps ; mouth cone
short and rounded. Antennae eight-segmented ; sensorium on II near apex. Pronotum with
anteromarginal and anteroangular setae minute, posteroangulars about twice as long as
epimerals ; epimeral sutures complete ; mesopraesternum complete, praepectus absent with
cuticular islets arranged linearly in a regular pattern. Meso- and metanotum faintly sculptured,
without major setae ; ventral thoracic setae minute. Fore femora thickened in both sexes,
fore tarsus unarmed in female, with stout tooth in male. Fore wing cilia widely spaced, distal
half of wing parallel-sided without accessory cilia ; two pairs of small sub-basal wing setae,
BI minute. Pelta not constant in shape, margin frequently broken up into cuticular islets.
Tergites II-VII with two pairs of wing-retaining setae, these setae curved but not sigmoid ;
BI and 62 on IX almost as long as tube in both sexes ; terminal setae shorter than tube ;
fustis of female short and weak. Sternal accessory setae less than iO[j. long, each sternite with
less than six accessory setae ; sternite VIII of male with transverse glandular area ; sternite VI
of male frequently with a pair of deeply reticulate areas.

Priesner originally placed this genus in the Haplothripini near to Adraneothrips.
However the wing form of Psalidothrips is probably an adaptation to the leaf-litter
habitat. The genus has subsequently been placed in both the Phlaeothripina and
Mesothripina, but it is probably related to Sagenothrips, another Melanesian leaf-
litter form, and should come into the Hoplothripina. Most Phlaeothripidae have a
pair of long setae near the posterior ventral margin of the head, but in Psalidothrips
these setae are short and there is a pair of long setae between the tentorial pits.
The genus is also unusual in the small size of the maxillary palps and the anterior
setae of the pronotum, as well as the few sternal accessory setae.

The three species in this genus known to the author may be distinguished by
means of the following key. Unlike many fungus-feeding species of thrips, the two
new species are remarkably constant in body size and the lengths of their major
setae.

1 Antennal segments with lines of sculpture ; segments III and IV with two sense

cones ; maxillary stylets subparallel, retracted about half way into head ; head,
antennae, mesothorax and abdominal segment II brown, rest of body yellowish,

male micropterous grandis sp. n. (p. 103)

Antennal segments without sculpture ; III and IV with three sense cones, the ventral
one smaller than the lateral ones ; maxillary stylets wide apart, V-shaped low in
head ; abdominal segments shaded laterally, II not distinctly browner than
other segments ....... 2

2 Sense cones on antennal IV about two thirds as long as segment ; postocular and

epimeral setae with apices expanded ; head brown ; male not known

atnens Priesner

Antennal sense cones shorter, on IV about one half as long as segment ; major
setae acute or softly rounded at apex ; head pale medially, deeply shaded between
eyes and along cheeks ; male macropterous . . . minor sp. n. (p. 103)

According to the descriptions given by Hood (1955) the four species from Brazil
may be distinguished as follows ; dissidens and umbraticus have three sense cones
on antennal III and four on IV, and dissidens is almost uniformly brown ; retifer
has three sense cones on both III and IV but the head is reticulate all over ; con-
ciliatus has two sense cones on III and IV as in grandisbut the segments are apparently
not reticulate and the sense cones are more than half as long as the segments.

L. A. MOUND

15

FIGS. 13-17. Figs. 13 & 14. Sophikothrips malaitae : 13, Head and pronotum. 14,
Left antenna. Figs. 15 & 16. Psalidothrips grandis : 15, Left antenna. 16, Head.
Fig. 17. Psalidothrips minor, left antenna.

THYSANOPTERA FROM SOLOMON ISLANDS 103

Psalidothrips grandis sp. n.

(Text-figs. 15 & 16)

$ (macropterous) . Colour yellow ; head, abdominal tergite II, anterolateral sclerites of
mesothorax, and antennal segments brown, base of antennal III and distal half of II paler ;
head sometimes pale at base ; mid and hind coxae, and sides of tube shaded ; fore wings
shaded except at median constriction.

Head weakly sculptured at base (Text-fig. 16), ventral surface smooth ; antennal segments
sculptured, lateral sense cones on III i^-i6[i, ventral sense cone not developed (Text-fig. 15).
Anteroangular and anteromarginal setae of pronotum 6{x long ; midlateral seta 5O(x, epimeral 30(1,
posteroangular 70-90(1.. Median setae of metanotum weak, less than 15^ long, about 40^
apart. Basal wing setae small, BI, 6(jt, B2 and 63 15-30^. Pelta broadly bell-shaped ; tergites
very faintly sculptured ; tube rather strongly narrowed, about 1305^ long, yojj. wide at base,
25(i at apex.

Measurements in microns of holotype with range from five paratypes in parentheses. Body
length 1850 (1750-1900). Hind tibia 195 (180-200). Fore wing 830 (730-900). Head, length
210 (195-) ; width 195 (180-). Pronotum, length 155 (145-160) ; width 260 (240-). Tergite
IX, BI 135 (125-) ; 62 165 (145-). Tube length 130 (125-140). Antennal segments length
39 (-42) ; 39 (-42) ; 65 (-68) ; 52 (45-55) ; 52 (49-55) ; 55 (52-) ; 49 (45-52) ; 49 (45-52) ;
terminal seta 70.

<J (micropterous) . Colour and chaetotaxy similar to female ; fore femora thicker than
female, fore tarsal tooth more than two thirds as long as width of tarsus. Glandular area on
abdominal sternite VIII 30(x long extending across full width of sternite ; sternite VI with
paired submedian areas of deeper reticulation sometimes present.

Measurements in microns of allotype. Body length 1500. Hind tibia 175. Fore wing 210.
Head, length 190 ; width 160. Pronotum, length 155 ; width 230. Tergite IX, BI 120 ;
B% 80. Tube length 100. Antennal segments 35 ; 35 ; 58 ; 45 ; 52 ; 52 ; 45 ; 45 ; terminal
seta 60.

Holotype $. KOLOMBANGARA : North of Kuzi at 1000 feet, 6.ix.i965 (P.N.L.).

Allotype <$, 12 $ and 7 $ paratypes taken with holotype.

Other material. KOLOMBANGARA : North of Kuzi at 250 feet, 12 $, 4 <$ and i
larva, 6.ix.i965 (P.N.L.). GUADALCANAL : Nuhu at 1000 feet, i $, 28.x. 1965
(P.N.L.). VANGUNU : i ?, io.vi.ig66 (P.J.M.G.).

Although the females of this species are macropterous, many specimens had the
distal half of the fore wings broken. This condition was so common that it is
possible the wings are broken naturally in the field.

In the larva the anterior margin of the pronotum bears very small setae as in the
adult.

Psalidothrips minor sp. n.

(Text-fig. 17)

$ (macropterous) . Colour pale yellowish brown ; dark brown at anterior and lateral margins
of head, and anterior and lateral margins of pterothorax ; antennae shaded brown, also anterior
corners of abdominal tergites ; wings shaded except at median constriction, cilia dark.

Head almost without sculpture, very similar to grandis ; surface of antennal segments
smooth (Text-fig. 17), ventral sense cone on III and IV about ioy. long. Midlateral seta of
pronotum weakly expanded, all other major setae acute or softly rounded at apex. Tergal
sculpture weaker than in grandis ; tube less conical, about 115^ long, 6opt wide at base, 25[z at
apex.

io4 L- A- MOUND

Measurements in microns of holotype with range from five paratypes in parentheses. Body
length 1750 (1550-1800). Hind tibia 175 (170-). Fore wing 730 (700-750). Head, length
170 (-180) ; width 165 (-175). Pronotum, length 135 (130-140) ; width 225 (210-230).
Tergite IX, BI 100 (-115) ; B£ 140 (130-). Tube length 115 (no-). Antennal segments
length 30 (-32) ; 40 ; 58 (54-) ; 52 (45-) ; 49 (45~) ; 49 (45-52) ; 39 (42-) ; 42 (39-) ;
terminal seta 50.

cj (macropterous) . Colour and chaetotaxy similar to female ; fore femora thickened,
fore tarsal tooth about two thirds as long as tarsal width. Glandular area on sternite VIII
about I5fji long, extending fully across width of sternite ; sternite VI with a pair of submedian
deeply reticulate areas ; Ba on tergite IX much stouter than BI at base.

Measurements in microns of allotype. Body length 1600. Hind tibia 175. Fore wing 700.
Head, length 165 ; width 140. Pronotum, length 130 ; width 210. Tergite IX, BI 100 ;
Ba 90. Tube length 100. Antennal segments 26 ; 35 ; 55 ; 45 ; 49 ; 49 ; 35 ; 39 ; terminal
seta 42.

Holotype $. GUADALCANAL : Mt. Austen, 2i.iv.ig65 (P.J.M.G.}.
Allotype <$ collected with holotype.

Other material. GUADALCANAL : Mt. Austen, i $, 8.11.1966 (P.J.M.G.) ; Nuhu
at 1000 feet, i $, i <$, 28.X.I965 (P.N.L.). KOLOMBANGARA : ? locality, 3 <j>, 3 $,
9.vi.ig65 (P.J.M.G.} ; North of Kuzi at 500 feet, in forest leaf -litter, i <J, 6.ix.i965
(P.N.L.). WAGINNA : u <j>, 10 $ and i larva, 3.vii.ig66 (P.J.M.G.}. CHOISEUL :
Malangona, i $, 8.111.1964 (P. Shanahan) in Bishop Museum Collection.

This species is more closely related to amens Priesner, the type-species of the
genus from Java, than to grandis with which it was collected at one site in the
Solomon Islands.

SOLOMONTHRIPS gen. n.

Type-species : Solomonthrips greensladei sp. n.

Small slender species, light brown or frequently yellowish with brown markings, metanotum
and at least part of head reticulate. Usually macropterous but micropterous and hemimacrop-
terous individuals found. Antennae eight-segmented, almost moniliform, VIII always distinct,
usually constricted at base ; two sense cones ventrolaterally on III and IV, sensorium on II
near apex ; apical seta on VIII very long, usually longer than VII -f- VIII. Eyes well developed,
cheeks sharply incut behind eyes without major setae ; vertex reticulate at least in part,
postocular setae well developed ; mouth cone short and rounded apically, maxillary palps
small ; stylets deeply or barely retracted into head, usually with maxillary bridge.

Pronotum transverse, weakly sculptured, epimeral sutures complete ; only four pairs of
major setae present, anteroangulars (? or midlaterals) absent, posteroangulars not close to
epimeral sutures ; major setae broadly expanded apically. Mesonotum reticulate, without
major or elongate setae. Metanotum with elongate triangular band of reticulation, median
setae not enlarged. Praepectus present, sometimes weak ; probasisternum large and pig-
mented ; mesopraesternum not small but weakly sclerotized. Fore femora moderately en-
larged, sometimes with indistinct irregular hump on inner margin ; fore tibiae unarmed in
female ; fore tarsi armed in both sexes. Fore wings weakly expanded at base, distal two thirds
slender and parallel-sided ; cilia widely separated except around apex, no duplicated cilia ;
third sub-basal wing seta distant from one and two.

Pelta reticulate, bell-shaped ; tergites III to VIII with two pairs of wing-retaining setae,
only one wing-retaining seta on tergite II ; tergites laterally frequently with sculpture. Setae
on tergite IX shorter than tube, BI and 62 expanded apically ; terminal cilia of tube short

THYSANOPTERA FROM SOLOMON ISLANDS

105

22

FIGS. 18-23. Figs. 18-22. Heads of Solomonthrips species : 18, intermedius. 19,
greensladei. 20, striatus. 2i,fimbrii. 22, setifer. Fig. 23. Fore leg of S. greensladei
male.

and weak. Male without glands on sternites ; 62 on tergite IX not reduced to a stout seta,
little different from B2 of the female.

Although superficially resembling both Malacothrips and Adraneothrips, this

io6 L. A. MOUND

new genus can be distinguished from them both by the presence of the praepectus.
Mystrothrips is also similar but has three sense cones on the third and fourth antennal
segments, and has the lateral mesonotal setae well developed. The thoracic
chaetotaxy of Solomonthrips is unique and moreover the development of wing-
retaining setae on the eighth tergite is unusual. However the genus is probably
derived from a genus like Sagenothrips. The author has examined the unique holo-
type of Sagenothrips gracilioornis through the courtesy of Dr. Priesner, and this
specimen has no major mesonotal setae and the third sub-basal wing seta is more than
twice its length from the second. The pronotal anteroangular and midlateral
setae are very reduced, less than ion l°ng- Contrary to Stannard (1955 : 79) the
anteromarginal setae are present, i6ji long, the right hand one being displaced
submarginally and the left hand one partially obscured by the pigment of this
imperfectly cleared individual. Although the heads of both genera are more or
less reticulate, neither Sagenothrips nor Solomonthrips belong in the Glyptothripini.
The reticulation of the body and the weak structure of the wings are probably
functional adaptations to the leaf -litter habitat.

KEY TO THE SPECIES OF SOLOMONTHRIPS

1 Tergite VIII with BI not large and expanded as on tergite VII but acute and curved,

closely parallel to marginal wing-retaining setae (Text-fig. 24) ; fore femur in both
sexes with irregular hump on inner margin (Text-fig. 23) ; major males with
subapical fore tibial spur (male not known in striatus) ; two pairs of long ventral
interocular setae, as long as antennal III ....... 2

Tergite VIII with chaetotaxy closely similar to tergite VII, BI major straight with
expanded apex (Text-fig. 25) ; fore femur without an irregular hump on inner
margin, male without a subapical fore tibial spur ; ventral interocular setae not
as long as antennal III although sometimes longer than remaining ventral head
setae .............. 3

2 Sternal reticulations on V-VII anterior to median row of accessory setae with internal

longitudinal striations (Text-fig. 30) ; median area of vertex between postocular
setae clearly reticulate (Text-fig. 20) ; antennal III much paler than IV, head
without median longitudinal dark stripe ..... striatus sp. n. (p. 113)
Sternal reticulations without internal longitudinal striations ; median area of vertex
between postocular setae without sculpture or with very faint reticulations (Text-
fig. 19) ; antennal III not much paler than IV, head usually with median longitudinal
dark stripe greensladei sp. n. (p. 107)

3 Antennals III and IV with dorsal setae expanded apically (Text-fig. 27) ; head fully

reticulate both dorsally and ventrally (Text-fig. 22) ; ventral interocular setae no
longer than remaining ventral head setae ; probasisternum fused into single plate ;
abdominal sternites with more than twelve long accessory setae ; major body
setae very broadly expanded, lateral abdominals almost as broad as long ; maxil-
lary stylets barely retracted into head, maxillary bridge absent . setifer sp. n. (p. in)
Antennal segments with major setae acute ; head not fully reticulate ; ventral
interocular setae longer than remaining ventral head setae ; probasisternal plates
separate ; abdominal sternites with accessory setae small, less than a quarter as
long as sternal marginals ; lateral abdominal setae longer than width of apical
expansion ...... ...... 4

4 Upper surface of head with median longitudinal band of reticulation (Text-fig. 18) ;

stylets deeply retracted into head, maxillary bridge about one third of head

THYSANOPTERA FROM SOLOMON ISLANDS 107

width ; major setae short with expanded smooth apex, basal wing setae less than
half basal width of fore wing ; tergite IX setae BI rather short, two thirds of B2.

intertnedius sp. n. (p. no)

Upper surface of head not sculptured between postocular setae but with transverse
band of reticulations at base (Text-fig. 21) ; stylets low in head, maxillary bridge
more than half of head width ; major setae long, expanded apices fimbriate, basal
wing setae more than half basal width of fore wing ; tergite IX setae BI more than
two thirds as long as 62 ....... fimbrii sp. n. (p. no)

The development of the major seta BI on tergite eight as an additional wing-
retaining seta in greensladei and striatus is not always complete. Although weak
and sigmoidal in all the specimens studied, this seta in a few individuals is very
weakly expanded or " soft " at the apex instead of acute. The characters given
in the key suggest that setifer is quite distinct from the other species, but it appears
to be merely an extreme specialization within the group. The probasisternal
plates are large and heavily pigmented in the other species and so their fusion in
setifer is not particularly surprising. Similarly the placement of the maxillary
stylets and the extent of the reticulation on the vertex is variable between the
other species and the condition in setifer can be regarded conveniently as an extreme
of a series.

Solomonthrips greensladei sp. n.

(Text-figs. 19, 23, 24, 26, 34, 35)

The specimens on which this species is described are variable both in colour and
the lengths of some major setae. These specimens were collected at several different
sites in the Solomon Islands and although the two main colour forms were collected
together on both Guadalcanal and Kolambangara there is a possibility that local
populations may eventually be found to be specifically distinct. For this reason a
male has been selected as holotype, and this individual is weakly oedymerous and
has the aedeagus exposed (Text-fig. 35).

<J (macropterous) . Colour yellowish, thoracic sternites and coxae brown, head with lateral
margins and median longitudinal stripe brown ; antennae light brown, darkening distally ;
fore and hind wings and major setae shaded ; mid and hind tarsi yellow, rest of legs variable
pale brown ; abdominal segments weakly shaded posterolaterally, tube darker apically.

Head reticulate at base, weakly sculptured medially (Text-fig. 19) ; maxillary stylets low ;
antennals III, IV and V with two ventrolateral sense cones, a small dorsal sense cone on V
(Text-fig. 26). Pronotum wider than long, weakly scupltured medially (Text-fig. 19), epimeral
sutures complete. Fore femora weakly incrassate, interior margin with indistinct rugose
hump, fore tibia with apex thickened or with a spur in major males, fore tarsus with stout
claw (Text-fig. 23). Mesonotum reticulate near posterior margin, without long or expanded
setae. Metanotum reticulate medially with one pair of weak setae near anterior margin.
Fore wing typical of genus.

Pelta longer than wide, weakly reticulate (Text-fig. 34) ; tergites III-VIII with two pairs of
wing-retaining setae, the posterior pair much stouter than the anterior pair on anterior seg-
ments ; major seta BI on tergite VIII fine and curved, usually with acute apex, parallel to
wing-retaining setae (Text-fig. 24) ; BI on tergites II-VII with broadly expanded assymetric
apex, sculpture anterior to BI with stout dentate microtrichia, tergites weakly reticulate
medially ; BI on IX shorter than or sub-equal to B% with moderately expanded apex, B2

io8

L. A. MOUND

acute ; tube with faint sculpture like overlapping tiles, terminal setae shorter than tube.
Sternites transversely reticulate anterior to accessory setae, reticules sometimes with one or

24

26

FIGS. 24-29. Solomonthrips species. Figs. 24 & 25. Tergite VIII : 24, greensladei.
25, fimbrii. Figs. 26-28. Antennae : 26, greensladei. 27, setifer. 28, fimbrii. Fig.
29. Pelta of fimbrii.

THYSANOPTERA FROM SOLOMON ISLANDS 109

two faint longitudinal markings ; median sternal setae arise submarginally, about three times
as long as accessory setae.

Measurements in microns of holotype with range from three paratypes in parentheses. Body
length 1370 (1130-1500). Hind tibia 120 (105-145). Fore wing 570 (480-700). Head,
length 150 (130-175) ; width 140 (125-170) ; postocular seta 45 (39-65). Pronotum, length
97 (74-105) ; width 240 (190-255) ; epimeral seta 32 (-65). Tergite IX, BI 72 (65-95) ;
62 90 (80-110). Tube, length 97 (80-110) ; terminal setae 60 (50-80). Antennal segments
length 32 (29-32) ; 39 (-45) ; 68 (58-75) ; 55 (48-68) ; 61 (55-71) I 5° (45-) ; 32 (29-) ;
32 (29-) ; terminal seta 65 (58-97).

$ (macropterous) . Colour similar to male, but abdomen typically with sternite V as dark as
thoracic sternites ; several females have abdomen brownish yellow, as in typical males without
a dark band on segment V ; antennae variable, usually with apex of III not much paler than
IV. Chaetotaxy and sculpture of female very similar to male, fore tibiae not armed.

Measurements in microns of allotype with range from four paratypes in parentheses. Body
length 1650 (1400-1780). Hind tibia 175 (145-180). Fore wing 720 (570-). Head, length
160 (145-180) ; width 155 (145-170) ; postocular seta 45 (38-49). Pronotum, length 105
(80-120) ; width 225 (205-250) ; epimeral seta 35 (32-42). Tergite IX, BI no (84-) ; B2
IX5 (97-)- Tube, length 120 (105-125) ; terminal setae 70 (58-74). Antennal segments
length 39 (29-) ; 45 (39-) ; 77 (68-) ; 74 (64-) I 5« (52-) ; 39 (35~43) I 35 (32-39) ; terminal
seta 90 (80).

Holotype^. GUADALCANAL : Mt. Gallego at 2500 feet, 12^11.1965 (P.N.L.).
Allotype $ and paratype $ taken with holotype.

Other material. GUADALCANAL : Umasani River, forest litter, 2 °., i <$, 5.vii.ig65
(P.N.L.) ', Nuhu, i (£, 31. x. 1965 (P.N.L.) ; Mt. Austen, forest litter, i $, 19. xi. 1963
(P.J.M.G.) ; Mt. Austen, beaten from sticks on ground, 5 $ and i larva, 11.11.1966
(P.J.M.G.) ; Mt. Popanamisiu, montane forest litter at 5000 feet, i $, x.1965
(P.J.M.G.) ; Malekuna, forest soil, i $ ,9-iv. 1966 (P.J.M.G.}. MALAITA : Puama-
wara, forest soil, i $, ii.vii.i966 (P.J.M.G.}. NEW GEORGIA : Canonygu, forest
litter, i Cj, i8.lv. 1966 (P.J.M.G.} ; Barakoma, forest litter near airstrip, i $, 6. v. 1966
(P.J.M.G.}. RENDOVA : i $, 6^.1966 (P.J.M.G.). Gizo : i ?., i #, ii.ix.ig65
(P.N.L.). KOLOMBANGARA : near Kuzi ; forest litter on coral limestone, 2 $,
8.1x.i965 (P.N.L.) ; litter in mossy wood at 1500 feet, i °-f 4.ix.i965 (P.N.L.) ;
valley litter at 50 feet, 8.1x.i965 (P.N.L.) ; forest litter at 1000 feet, i $, i <$ and
3 larvae, 6.ix.i965 (P.N.L.).

The following material was mounted from tubes in the Bishop Museum, Hawaii.
CHOISEUL : Kitipi River at 80 m, 9 °-, 5 <£ and 7 larvae, 20.111.1964, and Malangona,
i <£, 4.111.1964 (P. Shanahan). KOLOMBANGARA : Pepele at 30 m, i <£, i $, 11.11.1964,
and i $, 13.11.1964 (P. Shanahan) ; Iriri at 5 m, 2 °., i <$, 4^11.1964 (/. Sedlacek).
MALAITA : Dala at 30 m, i ^ and 2 larvae, 14^1.1964 (J. Sedlaceck). GUADALCANAL :
Gold Ridge at 500 m, i $, 24.^.1956 (/. L. Gressitt).

Larvae taken with this species have four tubercles (or two bifid tubercles) on
the head just underneath the antennae. Larvae taken with striatus have similar
tubercles but the greensladei larvae have a smooth head and pronotal shield, and
abdominal segment nine is brown only in the distal half.

no L. A. MOUND

Solomonthrips fimbrii sp. n.

(Text-figs. 21, 25, 28 & 29)

cj (macropterous) . Colour light brown, antennals I and II and bases of III and IV yellowish ;
mid and hind tarsi and median half of tube pale ; fore wings uniformly shaded.

Head reticulate basally (Text-fig. 21), maxillary stylets low in head ; postocular and other
major setae with expanded fringed apices ; antennae shorter than in greensladei (Text-fig. 28)

Pronotum similar to greensladei but epimeral and midlateral setae twice as long as antero-
marginals and posteroangulars ; praepectus and mesopraesternum weak ; fore femora without
a tubercle, fore tarsi with a stout claw. Mesonotum as in greensladei, metanotum broadly
reticulate in median area. Fore wings typical of genus, sub-basal setae rather long.

Pelta with basal portion rather broad (Text-fig. 29) ; tergite VIII with two pairs of wing-
retaining setae, BI broadly expanded apically (Text-fig. 25). Tergal sculpture rather weak,
sternites without reticulations.

Measurements in microns of holotype with one paratype in parentheses. Body length 1150
(900). Hind tibia 130 (115). Fore wing 500 (450). Head, length 135 (125) ; width 125 (120) ;
postocular seta 48 (39). Pronotum, length 84 (?) ; width 175 (160) ; epimeral seta 45 (35).
Tergite IX, BI 45 (52) ; 62 60 (58). Tube, length 87 (72) ; terminal seta 39 (35). Antennal
segments length, 19 (23) ; 32 (32) ; 58 (52) ; 55 (48) ; 55 (48) ; 48 (42) ; 35 (26) ; 32 (29) ;
terminal seta 64 (58).

$ (macropterous). Colour of allotype and $ from San Cristoval similar to but darker than
holotype ; 2 $ from Guadalcanal dark brown with brown tube. Sculpture and chaetotaxy
very similar to male, pronotum more clearly sculptured than in greensladei.

Measurements in microns of allotype with one paratype in parentheses. Body length 1350
(1300). Hind tibia 140 (140). Fore wing 570 (550). Head, length 130 (130) ; width 130
(130) ; postocular seta 49 (49). Pronotum, length 80 (80) ; width 195 (195) ; epimeral seta
45 (45)- Tergite IX, BI 71 (71) ; Bg 77 (77). Tube, length 97 (97) ; terminal seta 48 (48).
Antennal segments length 26 (26) ; 35 (35) ; 61 (61) ; 58 (55) ; 55 (55) ; 48 (45) ; 32 (29) ;
35 (32) ; terminal seta 72 (64).

Holotype^. GUADALCANAL : Mt. Austen, forest litter, 2i.iv.i965 (P.J.M.G.).
Allotype $ with same data as holotype.

Other material. GUADALCANAL : Mt. Austen, forest litter, 2 $, 8.^.1966
(P.J.M.G.) ; litter in river forest hollow, i <$, 5^.1965 (P.N.L.). SAN CRISTOVAL :
near Wainoni, litter on ridge at 1000 feet, i $, 8.viii.i965 (P.N.L.). CHOISEUL :
Kitipi River, i $, 2 <£, 20.^.1964 (P. Shanahan) in Bishop Museum Collection.

The two dark brown females with the unicolorous brown tube referred to above
do not appear to differ in structure from the other specimens although the colour
difference is quite distinctive. The extent of the sculptured band near the posterior
margin of the vertex is not identical in any of the specimens.

Solomonthrips intermedius sp. n.

(Text-fig. 18)

cj (macropterous). Colour brown, antennals II and III, hind tarsi, and median area of
tergites yellow, also a pale longitudinal stripe behind each eye on vertex ; fore wings shaded.

Head reticulate with a pale unsculptured longitudinal stripe behind each eye (Text-fig. 18) ;
maxillary stylets retracted almost as far as eyes ; antennae slender, sense cones lateral on III
and IV. Pronotum irregularly sculptured, major setae short ; fore tarsal claw recurved but
small ; meso- and metanotum similar to greensladei ; Bs on fore wing more than five times its

THYSANOPTERA FROM SOLOMON ISLANDS in

length from Bg. Pelta stout basally as in fimbrii ; abdominal tergites reticulate antero-
medially, with dentate microtrichia laterally ; tergite VIII with two pairs of wing-retaining
setae, BI with moderately expanded rounded apex as other major setae. Sternal accessory
setae very small, less than one fifth as long as sternal marginals.

Measurements in microns of holotype. Body length 1250. Hind tibia 130. Fore wing 570.
Head, length 160 ; width 135 ; postocular seta 17. Pronotum, length 74 ; width 190 ;
epimeral seta 22. Tergite IX, BI 45 ; 6265. Tube, length 105 ; terminal setae 55. Antennal
segments length 26 ; 35 ; 64 ; 64 ; 64 ; 48 ; 29 ; 32 ; terminal seta 85.

§ (macropterous) . Rather paler than male but sculpture and chaetotaxy very similar.

Measurements of allotype. Body length 1400. Hind tibia 145. Fore wing 610. Head,
length 1 60 ; width 145 ; postocular seta 17. Pronotum, length 71 ; width 210 ; epimeral
seta 26. Tergite IX, BI 45 ; B2 71. Tube, length no ; terminal setae 52. Antennal seg-
ments length 29 ; 32 ; 68 ; 64 ; 64 ; 55 ; 26 ; 29 ; terminal seta 87.

Holotype ^. GUADALCANAL : Mt. Popanamisiu, montane litter at 5000 feet,
x.ig65 (P.J.M.G.).

Allotype $ with same data as holotype.

Solomonthrips setifer sp. n.

(Text-figs. 22, 27, 32 & 33)

cJ (micropterous) . Colour light brown, femora, tibiae, tube and lateral margins of head
darker ; base of antennal III yellow, and a yellow longitudinal stripe behind each eye.

Head fully reticulate (Text-fig. 22), ventral head setae all subequal ; maxillary stylets
barely retracted into head, bridge not visible ; antennae rather short, dorsal setae on III,
IV and V blunt or expanded (Text-fig. 27).

Pronotum sculptured (Text-fig. 22), probasisternal plates fused medially ; fore tarsal claw
slender as in intermedius . Mesonotum as in greensladei ; metanotum short, fore wings very
short but with three basal setae ; mesosternum with about 12 pairs of setae, metasternum
with about 24 pairs of setae.

Pelta rather broad, probably foreshortened as a result of microptery (Text-fig. 33) ; tergites
with sculpture well developed, lateral setae very short and broadly expanded (Text-fig. 32) ;
tergite VIII with two pairs of wing-retaining setae, BI short with broadly expanded apex ;
tergite IX with BI probably expanded and assymetric at apex ; seta between BI and 62
more than half as long as BI. Sternites transversely reticulate over whole area posterior to
antecostal ridge, with about twelve accessory setae each about equal in length to sternal marginal
setae.

Measurements in microns of holotype. Body length 1050. Hind tibia 120. Fore wing 115.
Head, length 145 ; width 125 ; postocular seta 26. Pronotum, length 64 ; width 185 ;
epimeral seta 26. Tergite IX, BI 58 ; B2 68. Tube, length 74 ; terminal setae 42. Antennal
segments length 26 ; 35 ; 55 ; 45 ; 45 ; 45 ; 32 ; 29 ; terminal seta 48.

$ (macropterous). Colour similar to male ; fore wing weakly shaded, dark only around
sub-basal setae at anterior margin. Sculpture and chaetotaxy very similar to male ; pro-
basisternal plates fused medially ; metanotum reticulate ; pelta rather broader than similar
species ; fore wing thickened on anterior margin between sub-basal setae. Sternal marginal
setae less than i -5 times as long as accessory setae.

Measurements in microns of allotype. Body length 1300. Hind tibia 145. Fore wing
570. Head, length 160 ; width 140 ; postocular seta 26. Pronotum, length 74 : width 195 ;
epimeral seta 29. Tergite IX, BI 74 ; B2 90. Tube, length 100 ; terminal setae 55. Antennal
segments length 32 ; 42 ; 58 ; 52 ; 55 ; 48 ; 35 ; 32 ; terminal seta 48.

Holotype $. WAGINNA : forest litter, 3.vii.i966 (P.J.M.G.).
Allotype $ collected with holotype.

L. A. MOUND

FIGS. 30-35. Solomonthrips species. Figs. 30 & 31. striatus : 30, Sternite VI. 31,
Tergite IV. Figs. 32 & 33. setifer : 32, Tergite V. 33, Pelta. Figs. 34 & 35, green-
sladei : 34, Pelta. 35, Aedeagus tip.

THYSANOPTERA FROM SOLOMON ISLANDS 113

Solomonthrips striatus sp. n.

(Text-figs. 20, 30 & 31)

§ (macropterous) . Colour yellowish, antennals IV-VIII dark brown ; sternite VI and
tergites VI and VII with brown markings, thoracic sternites and tube apex brown ; fore wings
shaded with median area paler ; major setae dark.

Head reticulate (Text-fig. 20), maxillary stylets retracted half way into head ; antennae
very similar to greensladei. Thorax and appendages, and abdominal sculpture and chaetotaxy
(Text-fig. 31) similar to greensladei ; anterior half of sternites V-VII with numerous longitudinal
striations within the reticles (Text-fig. 30).

Measurements in microns of holotype. Body length 1700. Hind tibia 160. Fore wing
700. Head, length 160 ; width 160 ; postocular seta 28. Pronotum, length 100 ; width
225 ; epimeral seta 29. Tergite IX, BI 103 ; 62 no. Tube, length 205 ; terminal setae 80.
Antennal segments length 35 ; 42 ; 80; 80 ; 74 ; 55 ; 35 ; 35 ; terminal seta 80.

$ (hemimacropterous). Very similar to macropterous female but darker, tube brown and
head shaded medially ; fore wing 550^ long, without cilia.

Holotype °- SAN CRISTOVAL : near Wainoni, in leaf-litter on ridge at 1000 feet,
S.viii.1965 (P.N.L.).

Paratype $ hemimacropterous and 3 larvae, SAN CRISTOVAL : 6 miles S.E. of
Wainoni, in moss-forest litter at 2325 feet, 3.viii.i965 (P.N.L.).

This species is very similar to greensladei although the sternal reticulations are
apparently quite distinctive. These striations may not be evident in specimens
that have not been fully cleared however.

Larvae taken with this species are very similar to larvae taken with greensladei
but have abdominal segment nine all brown, the head and pronotal shield bear
minute microtrichia, and the major setae are shorter and more broadly expanded
at the apex.

SOPHIKOTHRIPS gen. n.

Type-species : Sophikothrips malaitae sp. n.

Head small, broader than long ; eyes large ; postocular setae absent ; stylets retracted into
base of head ; antennae eight-segmented, III with three sense cones, IV with four. Pronotum
twice as long and twice as broad as head ; praepectus very weak, mesopraesternum absent ;
fore tarsus with tooth ; fore wing without accessory cilia ; major setae short, apices expanded.
Pelta divided into a small dark anterior portion and a transverse poorly defined posterior
sclerite ; sternites with no accessory setae ; tergites with one pair of straight wing-retaining
setae ; tube as long as head, apical setae shorter.

Nine species have been described in the genus Sophiothrips Hood, and some
authors place Nanothrips Faure with three species and Zaxenothrips Crawford with
one species in synonymy with this genus. Seven of the Sophiothrips species were
described as having two long dorsolateral sense cones on antennal segments three
and four, although both bicolor and vorticosus apparently have only one sense cone
on segment three. The species included in Zaxenothrips and Nanothrips are des-
cribed as having one sense cone on segment three and two on four, and these sense
cones are shorter than those found in the other species. Eleven of these thirteen

114 L- A- MOUND

species have well developed interocellar setae and a laterally displaced stout post-
ocular seta ; however 5. vorticosus is described as having small postocular and
interocellar setae, and N. breviceps is figured as having three small pairs of postocular
setae. The small head and wide pronotum may be an adaptation to the leaf-litter
habitat. The new species described below, although similar to this group, has
three long sense cones on antennal segment three and four on segment four, no
postocular setae except a minute one on the cheek, and moreover the maxillary
stylets are not restricted to the mouth cone.

Sophikothrips malaitae sp. n.

(Text-figs. 13 & 14)

$ (macropterous) . Colour, head, thorax and anterior half of pelta brown, rest of abdomen
yellow with apex of tube shaded ; mid and hind femora and tibiae shaded brown medially,
fore tibiae shaded along external margins, fore femora shaded at base ; antennals I and II
yellow, III-VI pale basally and shaded at least in apical half, VII and VIII brown ; major
setae pale ; fore wing shaded in basal third, pale distally. Body weakly sclerotized with
only very faint indications of sculpture.

Head small, broader than long, eyes well developed (Text-fig. 13) ; stylets retracted into base
of head, mouth cone rounded, maxillary palps well developed ; no postocular setae, intero-
cellars moderate. Antennae eight-segmented (Text-fig. 14), sensorium on II in distal half ;
three sense cones on III, four on IV, two on V and VI.

Pronotum very broad, anteroangular setae not developed, other major setae rather small
(Text-fig. 13) ; epimeral sutures complete ; praepectus weakly indicated by fusion of several
chitinous islets, gular sclerite anterior to praepectus well developed ; probasisternum broad,
spinasternum and mesopraesternum absent ; fore femora broader than head length, fore tibia
not armed, fore tarsi with moderate curved tooth at inner apical margin. Mesonotum without
major setae ; metanotum broader than long, median setae fine, 30(1 long. Fore wing bent
through an angle of about 5° in basal third, almost parallel-sided, cilia widely spaced except
around apex, sub-basal wing setae expanded, i6\i long.

Pelta divided into an irregularly oval anterior portion and a very weakly sclerotized, appar-
ently dumb-bell shaped posterior portion ; wing-retaining setae on tergites III-VI I strong,
almost straight, on tergite VIII very weak ; submedian tergal setae minute (3^) ; tergal
seta BI with expanded rounded apex, two thirds as long as wing-retaining seta ; lateral tergal
setae not enlarged ; BI and 62 on tergite IX weakly expanded at apex ; sternites without
accessory setae.

Measurements in microns of holotype. Body length 700. Fore wing 450. Head, length
70 ; width 130 ; maxillary palp 26. Pronotum, length 130 ; width 320 ; epimeral seta 26.
Tergite IV, wing-retaining seta 45 ; seta BI 32. Tergite IX BI 29. Tube, length 65 ; ter-
minal seta 50. Antennal segments 22 ; 32 ; 39 ; 39 ; 32 ; 29 ; 25 ; 23.

Holotype $. MALAITA : Givarin, 24.1.1965 (P.J.M.G.).

Tolmetothrips granti sp. n.

(Text-figs. 36 & 37)

$ (macropterous). Colour brown, head and particularly tube dark brown ; all tibiae and
tarsi yellow ; antennals I and VIII dark brown, VII and base of II paler, apex of II and III-VI
yellow, or VI shaded brown ; the yellow on legs and antennae is a rich golden yellow, not a
bright citron-yellow ; major setae shaded but not dark except at apex of tube ; fore wings

THYSANOPTERA FROM SOLOMON ISLANDS 115

deeply shaded in distal two thirds except for a pale longitudinal stripe near the posterior margin,
cilia dark.

Head about as wide as long, eyes not directed laterally and rather smaller ventrally than
dorsally (Text-fig. 36) ; cheeks project behind eyes, with several pairs of small setae ; dorsal
surface reticulate, postocular setae long, IOOJJL in holotype ; ventral surface of head without
sculpture, no long setae between eyes or near posterior margin, one pair of long setae just
posterior to tentorial pits ; maxillary stylets retracted as far as eyes, approaching each other
in middle of head, maxillary guides well developed ; mouth cone broadly rounded, maxillary
palps about 50^ long. Antennae eight-segmented, VIII weakly constricted at base but without
a basal neck (Text-fig. 37) ; sensorium on II in apical half of segment ; one sense cone on III,
three on IV, two (+ one) on V and VI, but the two external sense cones on IV frequently
replaced by one larger cone.

Pronotum broad, at least anterior half reticulate, epimeral sutures complete (Text-fig. 36) ;
all five pairs of major setae well developed (on holotype, AM 42^ ; AA 6o(j. ; ML 65^ ; Ep ioo[/. ;
PA SOJJL). Praepectus absent, mesopraesternum reduced, frequently absent medially. Meso-
notum reticulate, lateral setae about SOJJL long with expanded apex. Metanotum reticulate,
median setae acute, 2O|A long, 6o(ji apart and 70^ from anterior margin of sclerite. Fore femora
weakly expanded, tibiae and tarsi unarmed ; mid and hind tibiae with a stout apical seta on
external margin. Fore wing almost evenly wide, accessory cilia variable 5-10 ; three sub-basal
setae in straight line, about 70-80^ long with expanded apices.

Pelta bell-shaped but posterior flange broad. Tergites reticulate laterally, lines of sculpture
bear dentate microtrichia ; II-VII with two pairs of sigmoid wing-retaining setae, anterior
pair only half as long as posterior pair ; marginal seta BI long, about IOO[JL ; tergite IX BI and
62 with expanded apices, accessory setae small ; tube weakly constricted in basal third, apical
setae rather short ; fustis very reduced. Sternites not sculptured, with a median transverse
row of 7-10 accessory setae 2o\i long.

Measurements in microns of holotype. Body length 2500. Hind tibia 240. Fore wing 980.
Head, length 225 ; width 220. Pronotum, length 170 ; width 350. Tergite IX, BI 145 ;
BZ 140. Tube, length 225 ; basal width 98 ; apical width 49 ; longest terminal setae 160.

<J (macropterous) . Colour, sculpture and chaetotaxy very similar to female ; fore tarsus
with a stout median tooth more than one third of tarsal width long ; tergite IX seta B2 very
short and stout with a broad round apex ; sternite VIII with an irregular oval glandular
area anterior to accessory setae, about ioo[i wide.

Measurements in microns of allotype. Body length 2200. Hind tibia 200. Fore wing 750.
Head, length 190 ; width 195. Pronotum, length 160 ; width 320. Tergite IX, BI 160 ;
B2 40. Tube, length 195 ; basal width 100 ; apical width 50 ; longest terminal seta 160.

Holotype <j>. SAN CRISTOVAL : confluence of Warahito and Pogato Rivers,
from a convoluted (? leaf) gall, six inches in diameter, on a liana hanging from a
banyan, 24.vii.i965 (/. Grant).

Allotype (J and numerous specimens of both sexes and immature stages taken
with the holotype (69 $ $ and 16^,^ mounted on slides).

The gall in which this species was collected was spherical, about six inches in
diameter, composed of a solid mass of highly convoluted non- woody tissue. This is
apparently the largest gall caused by thrips which has been recorded. Only one
species has been found in the gall but there may be 10,000 individuals in this single
colony. The body size was found to be remarkably constant.

The leaf-feeding Phlaeothripinae of the Melanesian region are poorly known and
the generic classification of this group is rather difficult to interpret. The type-
species of Eothrips is not known to the present author but annulicornis Karny has

n6 L. A. MOUND

been studied and this genus apparently is distinct on account of the striate metano-
tum, long sense cones, and unarmed fore tarsi of the male. The new species granti
cannot be placed in Teuchothrips or Gynaikothrips (sensu strictu) on account of the
short head and reduced eyes, and moreover the short-headed Gynaikothrips citritibia
Moulton, 1940 from New Britain should be placed in Eothrips (comb. n). In
citritibia, of which the author has recently studied the holotype and allotype, the
sense cones on antennals III and IV are three quarters as long as the segments,
and the fore tarsus of the male does not have a tooth. Similarly granti cannot be
placed in Eugynothrips as the species of that genus have long sense cones and short
anterior pronotal setae.

The antennae, the sculpture and the chaetotaxy of granti are very similar to
smilacis Priesner, which is a common species, rolling the leaves of a climbing plant
Smilax australis in Eastern Australia. This is the only species at present placed in
the genus Tolmetothrips Priesner, 1953. In smilacis the maxillary stylets are wide
apart, low in the head and the eyes are not so reduced as in granti.

Priesner (1960) places the leaf-feeding, gall-forming species of the Gynaikothrips I
TeiichothripsjLiothrips complex in the same subtribe as the fungus-feeding species
of Hoplothrips. In the opinion of the present author, Hoplothrips can be dis-
tinguished from the leaf-feeding forms not only by its biology but also by the greater
number of sense cones on the third antennal segment and the constricted neck at
the base of segment eight. In these respects this genus is similar to the genera
around Hoplandrothrips, and Hoplothrips is probably a reduced and highly successful
offshoot from the fungus-feeding Phlaeothripina.

MEGATHRIPINAE

Atractothrips solomoni sp. n.

(Text-figs. 38, 39 & 43)

$ (macropterous) . Colour medium brown, lateral margins of head dark brown ; antennals
I-II and VI-VIII dark, III-V light brown with yellow bases ; all tarsi pale, mid and hind
femora and tibiae yellow at base and apex ; fore wings with two longitudinal shadings ; all
setae pale.

Antennae eight-segmented (Text-fig. 38) ; sensorium on II in apical half ; two (or three)
sense cones on III, four on IV, major dorsal setae with weakly expanded apices. Preocular
projection of head short (Text-fig. 39) ; interocellar setae large ; eyes rounded ; postocular
and mid-dorsal setae well developed ; maxillary stylets widely separated ; ventral surface of
head with less than ten pairs of fine setae.

Pronotum with five pairs of major setae (Text-fig. 39), anteroangulars and midlaterals on
an elongate tubercle ; epimeral sutures complete ; praepectus present, mesopraesternum
well developed. Fore tarsus with a minute tooth. Anterior angles of mesothorax not expanded ;
fore wing slightly bent before middle, evenly wide to apex, without accessory cilia, basal setae
E$2 and 63 stout.

Pelta very broad, curving away from tergite II laterally ; tergites II-VII with one pair of
wing-retaining setae near posterior margin (Text-fig. 43) ; tergite IX with BI and 62 stout ;
tube weakly constricted at apex. Sternal accessory setae in regular transverse row ; sternite
surface anterior to accessory setae reticulate, reticles with internal markings.

Measurements in microns of holotype. Body length 2450. Hind tibia 260. Fore wing

THYSANOPTERA FROM SOLOMON ISLANDS

117

36

37

38

(\

FIGS. 36-39. Figs. 36 & 37. Tolmetothrips granti : 36, Head and pronotum. 37,
Right antenna. Figs. 38 & 39. Atmctothrips solomoni : 38, Right antenna (external
sense cones on III abnormal). 39, Head and pronotum.

n8 L. A. MOUND

950. Head, length 320 ; postocular seta 55. Pronotum, length 135 ; width 300. Tergite IX,
length 97 ; BI 70 ; ~Bz 80. Tube, length 290 ; terminal setae 130. Antennal segments
length 45 ; 60 ; 71 ; 65 ; 58 ; 55 ; 40 ; 32.

cJ (apterous). Colour rather darker than female. Head and pronotum as in female, ocelli
absent ; fore tarsus with stout claw. Tergites and sternites more heavily sculptured than
female, without wing-retaining setae.

Measurements in microns of allotype. Body length 1500. Hind tibia 200. Head, length
290 ; postocular seta 60. Pronotum, length 130 ; width 260 ; epimeral seta 71. Tergite IX,
BI 65 ; 62 80. Tube, length 225 ; terminal setae 130. Antennal segments length 50 ; 60 ;
68 ; 60 ; 55 ; 48 ; 40 ; 32.

Holotype $. CHOISEUL : Vasu River, i6.xi.iQ65 (P.J.M.G.}.
Allotype (£ and two paratype $ taken with the holotype.

This is the second species to be included in the genus Atractothrips. It differs
very considerably from bradleyi Hood, 1938 from Panama, and may eventually
have to be placed in a separate genus (see Stannard, 1957 ; 93-94). Four paratypes
of bradleyi have been studied and that species differs from solomoni as follows :

Antennal segment II with sensorium and two stout setae at apex ; sense cones on III minute ;
antennal segments long and slender, dorsal setae small and acute ; lateral ocellar setae long,
interocellars small ; eyes reduced, angular in outline ; postocular setae not distinct, dorsal
surface of head with about 10 pairs of setae ; pronotal midlateral setae very small, close to
anteroangulars ; three small setae along anterior margin of epimeral suture ; praepectus
absent ; fore tarsi unarmed in both sexes ; basal wing setae absent ; anterior angles of meso-
thorax projecting ; lateral metanotal sclerite with large expanded setae ; tergites with three
pairs of setae at margin including wing-retaining seta and posteroangular seta ; median tergal
setae and pores large ; tergites sculptured in front of antecostal ridge ; tube very long ; BI on
IX about one quarter as long as segment ; sternal accessories not in a regular transverse row.

MACHATOTHRIPS Bagnall

Machatothrips Bagnall, 1908 : 189. Type-species : M. biuncinatus by monotypy.
Adiaphorothrips Bagnall, 1909 : 536-537. Type-species : A. simplex by monotypy.
Machatothrips Bagnall ; Priesner, 1932 : 339-344.
Machatothrips Bagnall ; Mound, 1968 : 133-135.

Females in this genus have a series of tubercles on the fore femur although these
are not present in the males. Because the type-species of Adiaphorothrips is the
male of the type-species of Machatothrips, several species which were described in
Adiaphorothrips, particularly from Australia, are now referred to the older genus
although they do not belong there. The following species are at present placed in
Machatothrips.

antennatus (Bagnall, 1915), from West Sarawak.

artocarpi Moulton, 1928, from Formosa.

biuncinatus Bagnall, 1908, from New Guinea.

braueri Karny, 1912, from West Africa.

celosia Moulton, 1928, from Formosa.

haplodon Karny, 1925, from Uganda.

= spatiata Priesner, 1932, from Congo. Syn.n.

heveae Karny, 1921, from Java.

THYSANOPTERA FROM SOLOMON ISLANDS 119

tisshikii Ishida, 1932, from Japan.
montanus Priesner, 1932, from Sarawak.
quadrudentatus Moulton, 1947, from New Guinea.

Kurosawa (1968) has recently published the following synonymy of species
from Japan:

M. femoralis Ishida, 1932 = Docessissophothrips frontalis Bagnall.

M. ohtai Ishida, 1932 = H oplothrips flavipes Bagnall.

M. ipomoeae Ishida, 1932 — Rhaebothrips lativentris Karny.

One paratype of spatiata from Dr. Priesner 's collection has been studied and
compared with the type specimens of haplodon. M. isshikii is not recognizable
as a member of this genus from its description. The species artocarpi, celosia,
heveae and montanus are not at present clearly denned, and the unique holotype of
celosia has not been studied, although type material of all the other species has
been used in the preparation of the following key.

1 Mid-dorsal head setae very small, less than one quarter as long as postoculars . . 2
Mid-dorsal head setae usually well developed, more than one quarter as long as

postocular setae ............ 4

2 Fore femur of female with about 20 small tubercles 6(i in length ; anterior angles of

pronotum with several stout thorn-like setae .... antennatus Bagnall
Fore femora with few stout tubercles ......... 3

3 Fore femur of female with four tubercles ; antennal III much paler than IV

quadrudentatus Moulton

Fore femur of female with six or more tubercles ; antennal III dark with a lighter
area at base and at apical exterior margin ; pronotal midlateral setae pale, more
than two thirds as long as epimeral setae ..... haplodon Karny

4 Distal tubercles on fore femur fused into a ridge .... biuncinatus Bagnall
Fore femoral tubercles all distinct ......... 5

5 Antennal III dark, only basal membrane paler ; tubercles on fore femur slender, close

set, forming a regular series decreasing in length towards apex, 45[x-io(ji

braueri Karny
Antennal III yellow at base ; fore femoral tubercles stouter, more widely separated,

not forming a regular series decreasing in length evenly ..... 6

6 Fore femur with four (or five) tubercles, no tubercles on basal half of femur ; antennal

III sharply yellow at extreme base, apex uniformly dark . artocarpi Moulton (p. 119)
Fore femur otherwise .......... 7

7 Fore femora with three tubercles montanus Priesner

Tubercles present on most of inner margin of femora . . . heveae Karny

Machatothrips artocarpi Moulton
Machatothrips artocarpi Moulton, 1928 : 322-325.

The statement in the original description that the mid-dorsal head setae are
' very small, hardly one fourth the length of postoculars ' is not correct. In the
holotype these setae are almost erect but their length has been calculated by using a
microscope with the vertical movement calibrated in microns. The mid-dorsal
setae are about Son and the postoculars i8ou.

The species is apparently widespread in the Western Pacific, but as indicated
above it is not clearly different from heveae or montanus. The females from the

120 L. A. MOUND

Solomon Islands have four tubercles on the fore femora, but of three females taken
together in New Guinea one has four tubercles and the other two have five.

Material studied. Holotype $ and allotype <$, FORMOSA : Kagi, on Artocarpus,
io.viii.iQ27 (R. Takahashi). NEW GUINEA : Maffin Bay, on bark, 3 $, i <£, ix.iQ44
(E. 5. Ross) ; in California Academy of Sciences.

SOLOMON ISLANDS : Guadalcanal, in dried calyx of shrivelled coconut, 2 $, 2 <$,
x.i93i (R. J. Lever) ; Umasani River, Areca macrocalyx dead fronds, 2 $?, i $,
9^.1965 (P.N.L.) ; Kukum, i $ 3 #, 1962 (P.J.M.G.).

Mecynothrips snodgrassi Hood

Mecynothrips snodgrassi Hood, 1952 : 294.
Mecynothrips snodgrassi Hood ; Mound, [in press].

This species was described on a single oedymerous male from the SOLOMON
ISLANDS : Big Florida Island, on a shrub along jungle trail, 29. xi. 1944 (H. E.
Milliron). Species of Mecynothrips are also known from New Guinea, Kei Islands,
Manus Island, New Britain and Eastern Queensland. The genus has recently
been revised and snodgrassi, which is only known from the Solomon Islands, can be
distinguished by the apical expansion of the fore tibia lying parallel to the fore
tarsal claw.

Material studied. BIG FLORIDA ISLAND : Tulagi, i <£, 2.ix.i96o (C. W. O'Brien).
GUADALCANAL : Gold Ridge at 600 m, i $, 22. vi. 1956 (/. L. Gressitt), in the Bishop
Museum collection, Hawaii.

OMMATIDOTHRIPS gen. n.

Type-species : Ommatidothrips lawrencei sp. n.

This genus is erected for a species in which the head is strongly sexually dimorphic,
but in which both males and females have an area of clear cuticle on each cheek
posterior to the compound eyes, apparently representing a single isolated
ommatidium. Eyes of this type have not previously been described in the Thysano-
ptera, but apterous females of a second species of this genus have been collected by
the author on dead grass in Northern Queensland, Australia.

Large brown, non-sculptured species, feeding on fungal spores. Head long, dorsally elevated,
eyes rather small, with one ommatidium separate on each cheek ; cheeks convex with stout
spines in male, weakly concave with fine setae in female ; postocellar and postocular setae
long ; stylets approach each other in middle of head, mouth cone rounded with large maxillary
palps. Fore tarsus armed in both sexes ; wings, when present, of equal width with few
accessory cilia. Pronotum emarginate anteriorly, anteroangular setae not arising at margin ;
epimeral sutures complete, praepectus present. Tube almost as long as head, with many
fine setae ; both sexes with B2 on tergite IX less than half as long as BI ; abdominal tergites
with one pair of wing-retaining setae.

The relationships of this new genus within the Megathripinae are not clear.
The long head with the maxillary stylets approaching each other medially suggests
Megalothrips, but in that genus there is a stout seta on the cheek behind each eye

THYSANOPTERA FROM SOLOMON ISLANDS

40

FIGS. 40-43. Figs. 40-42. Ommatidothrips lawrencei : 40, Head, pronotum and fore
leg of female. 41, Head and fore leg of male. 42, Left antenna. Fig. 43. Atractothrips
solomoni, tergite IV.

122 L. A. MOUND

and the males have a pair of drepanae on abdominal segment six. The new species
is unusual in the subfamily in having the seta 62 on tergite nine of the male less
than half as long as BI, and in this character it resembles Abiastothrips. Several
genera which Priesner (1960) places in the Cryptothripina are regarded by Stannard
(1957) as being quite unrelated and as belonging in the subfamily Phlaeothipinae.
Certain of these genera have the head elevated in the midline and do not have
three equally long pairs of setae on tergite nine in the males. The present author
considers that at least Abiastothrips belongs in the Megathripinae, near to Crypto-
thrips, and it is to these forms that the new genus is most closely related.

Ommatidothrips lawrencei sp. n.

(Text-figs. 40-42)

$ (macropterous) . Colour medium brown, increasingly dark toward posterior, tube black
in basal two thirds ; major setae light brown ; legs yellowish, fore femora sometimes darker ;
basal half of antennal III and apex of II yellow, rest of antenna light brown, wings weakly
shaded.

Head long, weakly elevated in midline, with very faint sculpture (Text-fig. 40) ; cheeks
weakly expanded behind eyes, bearing an isolated ommatidium on each side ; cheeks parallel-
sided or weakly concave, with about seven pairs of setae ; one pair of postocular and postocellar
setae almost as long as head-width, middorsal head setae about one third as long as head width ;
maxillary stylets broad, band-like, approaching each other in midline ; mouth cone rounded,
maxillary palps IOOJA long. Antennae on a short preocular process, eight-segmented ; sen-
sorium on II in apical half of segment ; two sense cones on III, four on IV ; VIII constricted
at base (Text-fig. 42).

Pronotum not sculptured, anterior margin thickened and deeply concave ; epimeral sutures
complete ; epimeral setae more than three times as long as anteromarginal setae, remaining
prothoracic setae about two thirds as long as epimerals ; praepectus narrow, transverse ;
probasisternal plates large with four long setae on anterior margin ; mesopraesternum shallow
boat-shaped. Mesonotum with three pairs of fine setae less than 20^ long, with faint lines of
sculpture. Metanotum not sculptured medially, median setae about 5O(x long. Fore femora
moderately thickened, external margin with long fine setae ; fore tibiae slender ; fore tarsal
tooth shorter than width of tarsus. Fore wings almost parallel-sided, about goy. wide medially
and loopi. wide subapically ; 12-15 accessory cilia ; only two pairs of major sub-basal wing
setae, about 130^ long, BI fine and hair like 30^ long.

Pelta broad, not trilobed, weakly reticulate in anterior half. Tergites III-V with one pair
of sigmoid wing-retaining setae, tergites II and particularly VI and VII with weak almost
straight wing-retaining setae ; tergites III-VI laterally with a group of about 12 fine setae
anterior to BI and Bg, these groups of setae extend medially across the tergites of the posterior
segments so that IX bears two irregular transverse rows of fine setae ; 62 on IX less than
half length of BI ; sides of tube straight with numerous fine setae about 30^1 long, base of tube
i30jj., apex 55[x wide, longest terminal seta about 230^. Sternites with transverse row of about
15 accessory setae 40^1 long, sternites VII and VIII with accessory setae in two transverse rows ;
BI on sternites at least twice as long as Bg, arising well in front of posterior margin.

Measurements in microns of holotype with range from five paratypes in parentheses. Body
length 3400 (-3800). Hind tibia 450. Fore wing 1250. Head, length 450 (-470) ; width.
behind eyes 240 ; postocular seta 230 (210-240). Pronotum, length 130 ; width 350 (-390) ;
epimeral seta 190 (-225). Tergite IX, BI 350 (340-360) ; 62 115 (-160). Tube length 420
(-450). Antennal segments III-VIII, 160 (-170) ; 130 (-135) ; 115 ; 80 ; 70(65-) ; 80(68-).

(J (apterous). Colour similar to female, setae brown on cheeks and base of fore femora.
Head weakly elevated in midline, without sculpture (Text-fig. 41), cheeks convex medially

THYSANOPTERA FROM SOLOMON ISLANDS 123

with about 10 stout setae ; ocelli absent, otherwise similar to female. Pronotum as in female
except that posteroangulars almost equal epimerals, setae particularly long in oedymerous
males ; fore femur broad with about 6 stout cyathiform setae at posterior external angle ;
fore tibia with a small tubercle close to apex ; fore tarsus stout, longer than tarsal width (Text-
fig. 41). Abdominal tergites with reduced wing-retaining setae, VI-IX with transverse row
of fine setae ; sternites, tergite IX and tube similar to female, although tube comparatively
short, narrowing from 1305^. to 5O[x.

Measurements in microns of allotype with range from three paratypes in parentheses. Body
length 3400 (3200-3500). Hind tibia 400 (-420). Head, length 410 (-460) ; width behind
eyes 220 (200-) ; postocular seta 225 (-240). Pronotum, length 130 (-200) ; width 450 (-560) ;
epimeral seta 250 (-320). Tergite IX, BI 330 (320-360) ; B2 115 (95-). Tube length 350
(320-360). Antennal segments III-VIII, 145 (-160) ; 105 (-120) ; 95 (-105) ; 65 (-70) ;

65 (58-) ; 70.

Holotype $. GUADALCANAL : near Honiara, Poha River, ig.xi.i965 (P.N.L.).

Allotype $. GUADALCANAL : Mt. Austen, in litter between buttress roots of tree,
4.vii.i966 (P.J.M.G.), collected with 2 $, 3 $ paratypes and 2 larvae.

Other material. GUADALCANAL : Umasani River, in forest leaf-litter, i $ and

1 larva, 5^.1965 (P.N.L.) ; Mt. Austen, in forest litter, 4 $, i ^ and i larva,
29.xi.i963 (P.J.M.G.) ; Mt. Austen, 2 $, 8.^.1966 (P.J.M.G.). KOLOMBANGARA :

2 c?, 9-vi.i965 (P.J.M.G.). NEW GEORGIA : Munda, 3 $ and n larvae, 15.^.1966
(P.J.M.G.). RENDOVA : i <£, 6.V.I966 (P.J.M.G.). CHOISEUL : Vasu River,
i 2, i c?, i6.xi.i965 (P.J.M.G.).

Rhaebothrips lativentris Karny

Rhaebothrips lativentris Karny, 1913 : 128-129.

The genus Rhaebothrips was erected by Karny for the species lativentris, based on a
single male from Formosa. Two further species have been described in the genus,
major Bagnall, 1928 from Samoa, and fuscus Moulton, 1942 from Guam, Fiji and the
Torres Straits. Karny (1924) recorded lativentris from Queensland, and there are
specimens determined as that species by Karny in Dr. Priesner's collection from
Ponape. Moulton (1944) recorded major from Fiji but did not refer to his species
fuscus in that paper, although his remarks on the variation in major would appear
to include fuscus as a variety. The unique holotype of major is lost (Mound, 1968),
but Bianchi (1953) states that Moulton's specimens from Hawaii and Fiji can be
separated from new specimens available from Samoa.

At present it is not possible to be certain of the specific identity of the Solomon
Islands representatives of this genus, but it seems that there is just one widespread
species involved. The specimens from Ponape mentioned above vary greatly in
size, both within and between sexes. In the specimens listed below, it has been
noted that the tube is shorter than the head in micropterae but longer than the head
in macropterae. However the tube is relatively longer in males than in females.

Material studied. NEW GEORGIA : Munda, i $, i ^ (mac.), 15.^.1966 (P.J.M.G.).
GUADALCANAL : Mt. Austen, under bark, i $ (mic.), 24.xi.i965 (P.J.M.G.) ; Lunga,
i $ (mic.), 15.^.1963 (P.J.M.G.). SAN CRISTOVAL : 10 miles south of Wainoni,

I24 L. A. MOUND

litter 3 feet up palm stump, i <£ (mic.), 27.vii.ig65 (P.N.L.) ; 6 miles S.E. of Wainoni,
in ground moss of forest, i $ (mic.), 3^.1965 (P.N.L.). VATILAU : i $ (mac.),
(PJ.M.G.}.

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— 1966. Thysanopterologica Indica IV. Bull. Ent. Loyola Coll. 7 : 1-12.

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Soc. Northumb. 3 : 183-217.

— 1909. On some new and little known exotic Thysanoptera. Trans, nat. Hist. Soc.
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597-

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1928. On some Samoan and Tongan Thysanoptera, with special reference to Ficus

gall-causers and their inquilines. Thysanoptera. Insects Samoa 7 : 55-76. British

Museum (Nat. Hist.), London.
1933- A contribution towards a knowledge of the Thysanopterous genus Haplothrips

Serv. Ann. Mag. nat. Hist. (10) 11 : 313-334.
BEACH, A. M. 1896. Contributions to a knowledge of the Thripidae of Iowa. Proc. Iowa

Acad. Sci. 3 : 214-227.
BEAGLEHOLE, J. C. 1955. The Journals of Captain James Cook on his voyages of discovery.

I. The voyage of the Endeavour 1768-1771. Hakluyt Society, Cambridge.
BIANCHI, F. 1953. Thysanoptera of Samoa. Proc. Hawaii ent. Soc. 15 : 93-108.
BUFFA, P. 1909. I Tisanotteri esotici esistenti nel Museo Civico di Storia Naturale di Geneva.

Redia 5 : 157-172.
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2 : 153-170-

FRANKLIN, H. J. 1908. On a collection of Thysanopterous insects from Barbados and St.

Vincent Islands. Proc. U.S. natn. Mus. 33 : 715-730.
GIARD, A. 1901. Sur un Thrips (Physopus rubrocincta nov. sp.) nuisable au Cacaoyer. Bull.

Soc. ent. Fr. 1901 : 263-265.
HOOD, J. D. 1915. Descriptions of new American Thysanoptera. Insecutor Inscit. menstr.

3 : 1-40.

— 1925. New Neotropical Thysanoptera collected by C. B. Williams. Psyche 32 : 48-69.

— 1937. A new Euoplothrips (Thysanoptera) from the Solomon Islands. Ann. Mag. nat.
Hist (10) 20 : 599-604.

— 1938. A new genus and species of Phlaeothripidae (Thysanoptera) from Palmetto.
Proc. biol. Soc. Wash. 51 : 27-32.

1955. Brasilian Thysanoptera VI. Revta bras. Ent. 4 : 51-160.

HOOD, J. D. & WILLIAMS, C. B. 1927. A synopsis of the Thysanopterous family Urothripidae.

Ann. ent. Soc. Am. 20 : 1-8.

ISHIDA, M. 1932. Fauna of the Thysanoptera in Japan. Insecta matsum. 7 : 1-16.
KARNY, H. 1912. Ueber einige afrikanische Thysanopteren. Fauna exot. 2 : 19-26.

— 1913. H. Sauter's Formosa-Ausbeute. Thysanoptera. Supplta ent. 2 : 127-134.

— 1921. Beitrage zur Malayischen Thysanopterenfauna IV. Thysanopteren von Hevea.
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— 1924. Results of Dr. E. Mjoberg's Swedish Scientific Expeditions to Australia 1910-1913.
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THYSANOPTERA FROM SOLOMON ISLANDS 125

KUROSAWA, M. 1968. Thysanoptera of Japan. Insecta matsum. Supplement 4. 94 pp.
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MOULTON, D. 1928. New Thysanoptera from Formosa. Trans, nat. Hist. Soc. Formosa

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— 1937. Further notes on Hawaiian thrips with descriptions of new species. Proc. Hawaii,
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— 1940. Thysanoptera from New Guinea and New Britain. Occ. Pap. Bernice P. Bishop
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MOUND, L. A. 1968. A review of R. S. Bagnall's Thysanoptera collections. Bull. Br. Mus.
nat. Hist. (Ent.) Supplement 11. 181 pp.

— [in Press). A review of the Melanesian genus Mecynothrips (Thysanoptera). /. nat.
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PRIESNER, H. 1927. Neue und wenig bekannte Thysanopteren, gesammelt in West Afrika,
von Prof. Dr. F. Silvestri. Boll. Lab. Zool. gen. agr. R. Scuola Agric. Portici 21 : 61-83.

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— 1960. Das System der Tubulifera (Thysanoptera). Am. ost. A had. Wiss. Mathematische-
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RAMAKRISHNA AYYAR, T. V. & MARGABANDU, V. 1939. Notes on some Indian Thysanoptera

with descriptions of new species. Indian J. Ent. 1 : 35-48.
STANNARD, L. J. 1955. On some reticulate-headed genera of the tribe Glyptothripini Priesner

(Thysanoptera : Phlaeothripidae) . Trans. Am. ent. Soc. 81 : 77-101.

- 1957. The phylogeny and classification of the North American genera of the sub-order
Tubulifera (Thysanoptera). Illinois biol. Monogr. 25 : 200 pp.

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INDEX

abdominalis, 89 bagnalli, 96

Adiaphorothrips, 118 Baphikothrips, 90

Adraneothrips, 90 Baphothrips, 94

arnens, 101 Bradythrips, 96

Androthrips, 90 Chirothrips, 89

antennatus, 93 claratibia, 98

artocarpi, 119 coloratus, 91

Atractothrips, 116 crassipes, 97

126

L. A. MOUND

dammermanni, 100
Diceratothrips, 90
dilatus, 100
Ecacanthothrips, 96
Eothrips, 116
Epithrips, 90
Eugynothrips, 116
Euoplothrips, 97
fimbrii, no
flavipes, 98
flavitibia, 98
gowdeyi, 98
gracilicornis, 98
grandis, 103
granti, 114
greensladei, 107
guineaensis, 96
Haplothrips, 89; 98
hesperus, 96
Horistothrips, 90
indicus, 98
inermis, 96
intermedius, no
Karnyothrips, 90
lativentris, 123
lawrencei, 122
leios, 94

Machatothrips, 118
Malacothrips, 90
malaitae, 114
Mecynothrips, 120

Microcephalothrips, 89
minor, 103
Mystrothrips, 99
Nesothrips, 90
Ommatidothrips, 120
Ormothrips, 96
perplexus, 89
Pleisothrips, 89
priesnerianus, 98
Psalidothrips, 100
Pseudodendrothrips, 89
Rhaebothrips, 123
rubrocinctus, 89
Sagenothrips, 99
Scirtothrips, 89
Selenothrips, 89
setifer, in
smilacis, 116
snodgrassi, 120
solomoni, 116
Solomonthrips, 104
Sophikothrips, 113
spiniceps, 89
spinipes, 96
striatus, 113
tabaci, 90
Taeniothrips, 89
Thrips, 90
Tolmetothrips, 114
tricolor, 94
unispinus, 90

LAURENCE ALFRED MOUND, B.Sc., D.I.C., D.T.A.,

Department of Entomology,

BRITISH MUSEUM (NATURAL HISTORY),

CROMWELL ROAD,

LONDON, S.W-7

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an account of their world distribution. Pp. 151 : 14 plates, 293 text-figures.
November, 1968. £5.

13. AFIFI, S. A. Morphology and Taxonomy of Adult Males of the families
Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210 : 52 text-
figures. December, 1968. £5.

14. CROSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa
and its Islands. Pp. 198 : I plate, 331 text-figures. July, 1969. £4 155.

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

£4-

16. GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe
(Hymenoptera : Chalcidoidea). Pp. 908 : 686 text-figures. November, 1969.
£19-

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A REVISION OF THE

GENUS CATOPTROPTERYX KARSCH

(ORTHOPTERA : TETTIGONIIDAE)

J. HUXLEY

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 24 No. 5

LONDON : 1970

A REVISION OF

THE GENUS CATOPTROPTERYX KARSCH
(ORTHOPTERA : TETTIGONIIDAE)

BY

JOHN HUXLEY .

British Museum (Natural History) \

Pp. 127-170; i Plate, 95 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 24 No. 5

LONDON : 1970

THE BULLETIN OF THE BRITISH MUSEUM

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

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

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

This paper is Vol. 24 No. 5 of the Entomological
series. The abbreviated titles of periodicals cited
follow those of the World List of Scientific Periodicals.

World List abbreviation
Bull. Br. M^4s. nat. Hist. (Ent.).

Trustees of the British Museum (Natural History) 1970

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 30 January, 1970 Price £i IDS.

A REVISION OF

THE GENUS CATOPTROPTERYX KARSCH
(ORTHOPTERA : TETTIGONIIDAE)

By JOHN HUXLEY

CONTENTS

Page

INTRODUCTION ........... 129

ACKNOWLEDGEMENTS ......... 130

MATERIAL ........... 130

TREATMENT ........... 131

TAXONOMIC STRUCTURE ......... 135

BIOLOGY ............ 137

Catoptropteryx Karsch . . . . . . . . . 137

Key to the species ......... 140

Descriptions of the species . . . . . . . . 147

REFERENCES .......... 170

SYNOPSIS

The African genus Catoptropteryx Karsch is fully revised and a key is given to the species.
Three specific synonyms are newly established and seven new species are described.

INTRODUCTION

KARSCH erected the genus Catoptropteryx in 1890 for a single new species, C.
guttatipes, from Cameroon. In 1896 he described a further six new species and
suggested that closely related to these were three of the four African species described
in Caedicia Stal, of which, however, he formally transferred only Catoptropteryx afra
(Karsch). Kirby, in his catalogue of 1906, listed eleven species of Catoptropteryx
Karsch and included all the African species of Caedicia Stal. No new species has
been recorded since 1896, with the exception of C. latipennis Chopard, 1955, which
is considered here to belong to another genus (see below). Although up to the
present time the genus Catoptropteryx Karsch has been remarkably homogeneous
and easily recognized, there has always been considerable difficulty in the separation
of the species. The present work, while attempting to remove this difficulty,
inevitably extends the morphological and geographic boundaries of the group,
making it more heterogeneous and less readily characterized. Over 1200 specimens
have been examined, including all the type material. Seven new species are
described, three specific synonyms are newly established and lectotypes are desig-
nated for C. signatipennis Karsch and C. neutralipennis Karsch. In the course of
examining the ovipositor as a source of taxonomic characters it was necessary to
homologize its parts in order to establish a terminology generalized enough for use
in taxonomic studies on other Tettigoniidae.

130 J. HUXLEY

Catoptropteryx latipennis Chopard, as mentioned above, is incorrectly placed in
this genus and is removed from it by the following synonymy, based on an examina-
tion of the type-specimens :

Symmetropleura plana (Walker)

Phaneroptera plana Walker, 1869 : 339. Holotype <$, SOUTH AFRICA : Natal (BMNH)

[examined].

Tylopsis plana (Walker) Kirby, 1906 : 441.
Symmetropleura plana (Walker) Ragge, 1964 : 298.
Catoptropteryx latipennis Chopard, 1955 : 267. Holotype <$, SOUTH AFRICA : Cape Province,

Tzitzikama Forest, Stormsrivierpiek (UZI Lund) [examined]. Syn. n.

ACKNOWLEDGEMENTS

I want to express my gratitude to the following specialists who, directly or
indirectly, have made available to me type specimens and other material from
their respective institutions :

Mr. P. Basilewsky, Dr. W. Bazyluk, Dr. M. Beier, Dr. I. J. Cantrall, Mr. R. H.
Carcasson, Dr. M. Descamps, Mr. M. Donskoff, Mr. Y. Gillon, the late Dr. H. J.
Grant, Dr. K. K. Giinther, Dr. R. Kumar, Mr. M. Lamotte, Professor C. H. Lindroth,
Miss J. M. McKay, Mr. E. Morales Agacino, Dr. E. C. G. Pinhey, Dr. D. C. Rentz,
Dr. C. H. F. Rowell, the late Dr. G. van Son, Mr. L. D. E. F. Vesey-Fitzgerald,
Mr. J. A. Whellan.

My especial thanks are due to Dr. D. R. Ragge for valuable criticism and advice
during the preparation of this work ; to Professor G. G. E. Scudder for his comments
on the interpretation of the ovipositor homologies ; and to Dr. M. C. Eluwa for
observations on the biology of Catoptropteryx Karsch and the use of his photograph
of the eggs (Plate i, fig. i).

MATERIAL

The museums and other institutions listed below have provided the material for
this study. The abbreviations used in the text to refer to type-depositories are
given in parentheses.

Academy of Natural Sciences of Philadelphia (ANS) ; British Museum (Natural
History) (BMNH) ; California Academy of Sciences, San Francisco (CAS) ; Coryn-
don Museum, Nairobi (CM) ; Institute Espanol de Entomologia, Madrid (IEE) ;
Instytut Zoologiczny of the Polska Akademia Nauk, Warsaw (IZPAN) ; Labora-
toire d'Entomologie, Office de la Recherche Scientifique et Technique Outre-Mer,
Abidjan (ORSTOM) ; Makerere University College, Kampala (MUC) ; Musee
Royal de 1'Afrique Centrale, Tervuren, Belgium (MR AC) ; Museum National d'His-
toire Naturelle, Paris (MNHN) ; Museum of Zoology, University of Michigan,
Ann Arbor (MZUM) ; National Museum, Bulawayo (NM) ; Naturhistorisches
Museum, Vienna ; Transvaal Museum, Pretoria (TM) ; Universitetets Zoologiska
Institution, Lund, Sweden (UZI) ; University of Ghana, Accra (UG) ; Zoologisches
Museum of the Humboldt-Universitat, Berlin (ZMHU).

REVISION OF CATOPTROPTERYX 131

TREATMENT

It has been found necessary in attempting to provide a reliable means of identifica-
tion of the species of Catoptrapteryx Karsch to make use of a few characters hitherto
not commonly employed in taxonomic work on the Tettigoniidae. One class of
characters that may deserve comment is that based on the coloration of the integu-
ment. Many workers still tend to regard colour characters as unreliable in the
sense that they are so often subject to individual variation as to be of dubious value
in differentiating the species. However, the level of significance of differences in
coloration, as with any other type of structural difference, varies from group to
group, and there is no doubt that within this genus alone a particular feature of the
colour pattern may show a consistent difference between a certain two species while
in other species the same difference is embraced by intraspecific variation. In
using the present revision, therefore, where a colour character is in the same state
in all the examined material of a species, the value of this should not be under-
estimated but assessed as a function only of the number of specimens on which the
description is based.

Much work has been done, especially in recent years, on the acoustic behaviour of
Orthoptera and the high taxonomic significance of song patterns is widely
recognized. However, although the nature of sounds produced by Ensifera is
clearly dependent in part on the structure of the stridulatory organ, and although
this structure can be fairly readily examined in dead specimens, very little use has
been made of this as a source of characters in systematic work on the group. The
present work makes use of the three most easily examined attributes of the stridula-
tory file, namely its gross appearance, length and number of teeth. The drawings
of files were made from stained and cleared preparations of actual specimens, using
a microprojector. The measurement of tooth number and file length was made on
nitrocellulose replicas of a subsample of each species. The technique for making
the replicas was essentially that described by Ragge (1969 : 172). A drop of 5%
ammonia solution containing a trace of detergent as a wetting-agent was used to
relax the wing base. To obtain sufficiently robust replicas from this genus it was
necessary to coat the whole of the proximal centimetre of the ventral surface of the
wing. The replicas were examined by diffuse transmitted light. ' Tooth number '
(T) includes all the teeth, however rudimentary, that can be discerned under the
low power (xioo) of a microscope. ' File length ' (F), which was measured using
a screw micrometer eyepiece, is the linear distance between the first and last tooth.
The measurements made on replicas do not differ significantly from those made on
the wing itself. Both tooth number and average tooth density (T/F) were found
to be useful characters. The interspecific and infraspecific variation in these
variates is shown in the scatter diagrams of tooth number on file length (Text-fig, i).
In any one species, there is a broad correlation between tooth number and file
length, as might be expected, but for the genus as a whole there is no such significant
correlation owing to the large variation of average tooth density between species.
For example, the sample mean of this last variate is 24-56 (teeth per mm) for C.
aurita sp. n. and 63-51 for C. nanus sp. n., producing widely separated clusters of

132

100-

80

60

40

20

J. HUXLEY

100^

80-

60-

40-

20-

• *\ . \A

cP

D

U D D D

n n

*V* fT<Y Y

X v/0( /~\ \_J
X **— '

punctulata X
guttatipes O
ambigua 0
occidentalis A

KEY

neutralipcnnis A
extensipes D

V5

2-0

x xx

2'5

0

naevia X

serrifera O
(K=sthn.forrn)

nanus 0

capreola A

apicalis A

aurita •

KEY

1-5

file length

2-0

2-5

FIG. i. Scatter diagrams of tooth number plotted against file length for the left male
stridulatory file of Catoptropteryx spp.

REVISION OF CATOPTROPTERYX 133

points for these two species. While there is of course considerable overlap, lor the
samples plotted each species does have a characteristic distribution of points and
many pairs of species can be clearly differentiated by simultaneous consideration
of tooth number and file length.

Although the form of the ovipositor valves is frequently used in the systematics
of the Phaneropterinae, little use is made of the more complex assembly of sclerites
at their base. This constitutes a mechanism for the control of the movements of
the valves, in particular during oviposition, and its structure probably relates
closely to oviposition behaviour. This aspect is peripheral to the present study
except insofar as it is suggested that the form of the basal mechanism, like that of
the valves themselves, is likely to be maintained within narrow limits of variability,
and for this reason may be expected to provide reliable characters for the identifica-
tion of the species. The shape of the basal sclerites is complex in many Phanerop-
terinae, and almost always so in those genera, like Catoptropteryx Karsch, in which
the valves are abnormally small, and whatever the functional significance of the
form may be its variation from species to species is often discrete and readily per-
ceived. Too commonly in the Tettigoniidae the males of a group of species may be
easily distinguished by clear differences in their genitalia, while identification of the
females relies heavily on their topographic association with identifiable males.
In many such cases study of the basal mechanism of the ovipositor reveals good
specific characters, and on the assumption based on this observation, that further
use will be made of this structure in future work on phaneropterine taxonomy,
I have considered it advisable to establish a morphological nomenclature at once
rather than lay the foundation of a confusing diversity of terms by constructing an
ad hoc terminology on a purely descriptive basis. Apart from the retention of the
terms ' dorsal valves ' and ' ventral valves ', already commonly used and un-
ambiguous in meaning, in preference to ' gonoplac ' and ' first gonapophysis ',
the ovipositor is described using the nomenclature of Scudder (19610, 19616, 1964).
Scudder and Harz intend to propose the terms ' supragonangulum ' and ' infra-
gonangulum ' for the anterodorsal and posteroventral divisions of the gonangulum
of the Tettigoniidae (Scudder in lit.} and I am anticipating the publication of their
work by adopting these terms here. The structure, interrelationships and homologies
of the sclerites of the ovipositor of Catoptropteryx Karsch were determined by examin-
ation of examples of all the species well represented in the available material. The
abdomens of dried specimens were macerated in 10% potassium hydroxide and
studied under isopropyl alcohol and in cleared and stained slide preparations.
The results of this investigation are summarized diagrammatically in Text-fig. 2.
The most useful specific differences in the basal mechanism are to be found by
examining the form and relative sizes of the lobe of the infragonangulum (iga) and
the first gonocoxa (gc i), and the spatial relationship of these sclerites to each other.
The supragonangulum (sga) and second gonocoxa (gc 2) also show specific differences
but these tend to be less well defined and the sclerites are not always so readily
examined in dried material. The subtlety of shape of the ovipositor parts makes
verbal diagnosis difficult and this has therefore been minimized in the species

134

J. HUXLEY

descriptions, reliance being placed almost entirely on the text-figures for comparison
of ovipositor characters. When comparing specimens with the text-figures it is
essential to bear in mind the possible effects of a number of factors, the operation of
which leads to a misleading diversity in the appearance of ovipositors of the same
species. All the drawings have been made from dried specimens in which the process
of drying causes variable distortion of sclerites, variable lateral displacement of
the subgenital plate, and variation in the degree to which the anterior part of the
basal mechanism is concealed by the ninth and tenth abdominal tergites. Further,
as the ovipositor in life consists of a system of moving parts, the positions of the
parts relative to one another are subject to variation ; for example, in Text-fig. 2,
anticlockwise rotation of the plate formed by the fused gonangulum and first gonocoxa
would result in an anterior displacement of the second gonocoxa and dorsal valve,
posterior displacement of the ventral valve, and partial concealment of the supra-
gonangulum beneath the ninth abdominal tergite. This sort of change produces
an unexpectedly large diversity in the gross appearance of the ovipositor.

Every species description includes the ranges in the material examined of certain
linear measurements commonly found to be useful in tettigoniid taxonomy at this
level. All measurements are given in millimetres. Means are not given for two
reasons : the samples are extremely heterogeneous and the relative contributions
of geographic and intrapopulational variation to the value of the mean are not

ep

ad

cd

FIG. 2. Diagram of the ovipositor of a generalized Catoptropteryx sp. Membrane is shown
stippled, ad, apodeme ; aiv, anterior intervalvula ; c, cercus ; cd, condyle ; ep,
epiproct ; gap i, first gonapophysis (= ventral valve) ; gap 2, second gonapophysis ;
gc i, first gonocoxa ; gc 2, second gonocoxa ; gp, gonoplac (= dorsal valve) ; iga,
infragonangulum ; piv, posterior intervalvula ; pp, paraproct ; sga, supragonangulum ;
sp, subgenital plate ; su, suture ; VIII, IX, X, terga.

REVISION OF CATOPTROPTERYX 135

assessed ; even in a (biologically) homogeneous sample knowledge of the mean
alone, with no estimate of the variance or skewness of the distribution, is not very
useful. Measurements were made using calipers, with the exception of the file
length (see above) and the median length of the pronotum, for both of which a
screw micrometer eyepiece was used. ' Total length ' means the distance from the
most anterior part of the head (excluding its appendages) to the tip of the flexed
hind wings. ' Length of fore wings ' is the distance from the distal margin of the
second axillary sclerite to the apex of the wing, and is conveniently measured by
inserting one point of the calipers into the small notch in the wing extending distad
from this sclerite.

In descriptions of the legs and antennae the terms ' internal ', ' external ', ' dorsal '
and ' ventral ' refer to these appendages in standard positions, the antennae and
fore legs directed forwards and the mid and hind legs directed backwards.

The wing- venation is described using the nomenclature of Ragge (1955). The
veins used in the diagnosis of Catoptropteryx spp. are illustrated and named in Text-
fig. 65.

TAXONOMIC STRUCTURE

It may be necessary to stress that throughout this paper, where expressions
such as ' affinity ' and ' related ' are used without qualification no phyletic meaning
is intended, such terms referring only to the degree of morphological resemblance
observed. Whether phyletic information can be inferred from the evidence of the
few morphological characters studied in this revision is another matter.

Text-fig. 3 is a schematization of the affinities of the species of Catoptropteryx
Karsch based on an intuitive selection of characters from the ovipositor, fore wing
venation, eyes, tympanic organs and male cerci. It is not based on a rigorous
phenetic analysis but is merely an attempt to give visual form to a largely subjective
assessment of the relations among the species. It can be seen that the genus falls
into three principal species-groups comprising respectively i) extensipes, 2} aurita
and serrifera, and 3) the remaining twelve species. The second group is linked to
the third both morphologically and geographically by C. serrifera sp. n. The third
and largest group subdivides itself fairly readily into the following three subgroups :
i) nanus, ramulosa, capreola and naevia ; ii) apicalis, afra and neutralipennis ;
iii) ambigua, nigrospinosa, punctulata, guttatipes and occidentalis. Of these, ii)
and iii) are strongly linked through C. neutralipennis Karsch.

C. aurita sp. n. is the only species in which the internal auricle of the tympanic
organs is at all developed. Although in the Phaneropterinae this most commonly
has the status of a generic character, the inclusion of C. aurita sp. n. in this genus,
with the consequent heterogeneity of the latter, is justified by the observation that
C. serrifera sp. n., which has no tympanic auricles, is rather more closely related to
the auriculate species than it is to the rest of the genus. I can find no real justification
for the attachment of special taxonomic significance to the difference in the tympanic
organ of these two species. Morphogenetically, the difference might readily be
assumed to be trivial in that the auriculate form could be simply derived from the

136

J. HUXLEY

open form by a slight expansion of the ventral internal rim of the tympanic opening.
The reverse transformation is equally feasible. A decision as to which may be the
primitive and which the derived form of this structure might be of some biogeo-
graphic interest, as the auriculate species is the only one with an East African
distribution. This decision clearly cannot be based on the kind of evidence available
here. However, one purely speculative possibility is that the auriculate form is
primitive and the open form derived from it. It might be supposed for example
that in an environment in which predation pressure is high, selection would favour
reduction of the auricle to give completely open tympana, which could be the more

FIG. 3. Schematic diagram of the affinities of the species of Catoptropteryx Karsch. The
thickness of the line joining two species is a measure of their morphological resemblance.
(See text, p. 135).

REVISION OF CATOPTROPTERYX 137

efficient condition for nondirectional defence responses. If, moreover, the population
density were high, reduction of mating efficiency by a reduction of the efficiency of
directional responses to acoustic stimuli might not be a dominant selective factor.
On the other hand, under conditions of low predator density and sparse populations,
directional response capabilities may be favoured over nondirectional, and the
auricle retained.

BIOLOGY

Nothing has been published relating to the biology of any of the species of Cato-
ptropteryx Karsch. Moreover, since all the properly documented material examined
was collected in light-traps, I have virtually no evidence in respect of ecological
information. Enquiries made of collectors have not enabled any conclusions to be
reached on this issue. The collecting localities almost all lie within areas of moist
lowland forest or forest-savanna mosaic and several collectors suggest that it is
likely to be a forest-dwelling genus. Dr. M. C. Eluwa, formerly of the University of
Ibadan, however, has suggested (in lit.) that the principal habitat of the genus
may be the bushes characteristic of farm lands. A few of the specimens examined
had feculae attached and microscopic examination of these has revealed plant
material only, without any trace of animal parts. In his laboratory Dr. Eluwa
has fed specimens of an unidentified species of Catoptropteryx Karsch with tender
leaves of Ficus sp. The dense sensory bristles of the valves of the ovipositor and
their lack of teeth in the majority of species indicate that eggs are not usually
deposited within plant tissues, and the small size of the valves argues against the
use of deep crevices in soil or bark as oviposition sites. Dr. Eluwa supports this
deduction with a single observation of oviposition in the laboratory. A row of
29 eggs was deposited along a vein on the abaxial surface of a Ficus leaf (see Plate i,
fig. i).

CATOPTROPTERYX Karsch

Catoptropteryx Karsch, 18906 : 361. Type-species, by monotypy, Catoptropteryx guttatipes
Karsch, 1890.

c£ $. Small to medium size. Integument generally smooth and shiny.

Length of antennae about i£ times total length ; margin of antennal scrobes not, or only
slightly, extending above fastigium of frons. Fastigium of vertex compressed, narrow, width
less than half width of first antennal segment, horizontal or very slightly declinate, sulcate
dorsally. Fastigium of frons triangular, acuminate, almost meeting ventral surface of fastigium
of vertex. Eyes large to very large, more or less prominent, sub-globose or slightly elliptical.

Pronotal disk more or less planar, lateral margins diverging posteriad, anterior margin straight
or slightly concave, posterior margin moderately to strongly convex ; lateral lobes of pronotum
more or less planar, meeting disk at very slightly rounded right-angle, height usually exceeding,
sometimes about equal to, length. Fore wings extending well beyond hind knees, length about
i \ times hind femur length ; Sc and R contiguous in proximal half ; Rs arising slightly proximad
of centre ; cross-veins usually somewhat regular and parallel ; archedictyon fairly open-
textured, membrane transparent. $ stridulatory organ small, without mirror ; stridulatory
rib not prominent dorsally ; file with 30-110 teeth. $ stridulatory apparatus present, consisting
of one or more thickened, transverse veinlets near posterior margin of cubito-anal area of right

138 J. HUXLEY

fore wing, each bearing dorsal row of small, regular, hook-shaped spines with points directed
distad. Hind wings fully developed, extending beyond fore wings in flexed position by about
-J- fore wing length. Fore coxae with long dorsal spine of circular transverse section. Fore
femora with about 1-4 very small internal ventral spinules (often o in C. extensipes Karsch).
Mid femora with about 1-4 very small external ventral spinules (often o in C. extensipes Karsch).
Hind femora with about 3-6 internal and 3-6 external ventral spinules ; hind knees with 2 small
spinules on each lobe. Fore and mid tibiae sulcate dorsally, each with about 1-5 internal
and 1-5 external ventral spurs ; fore tibiae with usually o, sometimes 1-2 external dorsal spurs in
addition to apical ; mid tibiae with 0-3 internal dorsal spurs in addition to apical. Hind
tibiae flat or somewhat convex dorsally, with well developed dorsal carinae, about 4-10
internal and 10-20 external ventral spurs, and about 20-35 dorsal spinules on each side ; apex
on each side with 2 large curved ventral spurs and i large dorsal spur. Tympanic organs
externally without auricles, internally auricle absent or (C. aurita sp. n.) more or less well
developed.

Abdominal tergites simple, without spines ; tenth tergite of $ unmodified. Epiproct simple,
triangular. <$ cerci unbranched, unarmed except for small apical or subapical spinule. $ cerci
abruptly narrowed in distal quarter. Ovipositor very small, length about f to £ median length
of pronotum ; margins of valves mostly smooth, occasionally finely crenulate or serrate at apex
of dorsal valve ; basal mechanism well developed, ist gonocoxa usually as large as or larger than
infragonangulum. $ subgenital plate with shallow triangular excision in distal margin ; styles
present, usually very small, sometimes apparently not articulated. $ subgenital plate simple,
with apical margin entire.

General coloration pale green to olive-green, sometimes overlaid with purple-brown over fore
wings, and with variable brown or black markings often present on lateral lobes of pronotum
and base and apex of fore wings. Hind wings hyaline or faintly fumose (markedly fumose in
C. extensipes Karsch). Dark areas of variable size usually present at some or all of following
loci : post-ocular regions of head ; dorsal surface of antennae ; fore tibiae in region of tympanic
organ, especially in lateral sulci ; hind tibiae proximally at two points on each side and one
point dorsally, and around bases of ventral spurs. All spurs and spinules blackened, at least
apically.

While Catoptropteryx Karsch is rather easily recognized (see Plate i, figs. 2-3),
it is not easy to make a brief list of diagnostic features distinguishing it from its
closest relatives. It resembles quite closely Caedicia Stal from which it seems to be
principally distinguished by its more globular eyes (or very much larger eyes in the
case of C. extensipes Karsch) and less prominent antennal scrobes. Confusion of
these genera is in practice unlikely as Caedicia Stal has an Australian and Austro-
oriental distribution. Dapanera Karsch, however, is extremely similar and has a
similar distribution to Catoptropteryx Karsch. The former genus may be dis-
tinguished by the fore wings which are broader and have a denser, more opaque
archedictyon, by the well developed ovipositor, and (except from C. aurita sp. n.)
by the auriculate inner tympanum. The most useful characters distinguishing
the species of Catoptropteryx Karsch are undoubtedly to be found in the basal
mechanism of the ovipositor (see p. 133), the male cerci and stridulatory organ
(see p. 131) and the coloration of the fore wings and pronotal lobes (see p. 131). It is
essential, however, for reliable determination to take into account all the characters
treated in the diagnostic descriptions.

The distribution of the genus falls into two distinct areas of tropical Africa (see
Text-fig. 4). C. aurita sp. n. has an East African distribution, while the rest of

REVISION OF CATOPTROPTERYX

139

the genus has a distribution characteristic of many West African Tettigoniidae,
extending from Sierra Leone and Guinea to Angola, Congo and Uganda.

FIG. 4. Map showing the distribution of Catoptropteryx aurita sp. n. (circles), and the
approximate distribution of the rest of Catoptropteryx Karsch.

I40

J. HUXLEY

KEY TO THE SPECIES

The characters employed in the key are chosen partly for convenience in use, and are not
always those on which the most weight is placed in the diagnoses of the species. Hence, the
key does not provide a complete diagnosis of each species, this being a function of the descrip-
tions. Every specimen, therefore, needs to be compared with the description indicated by the
key before the identification can be considered reliable.

1 Eyes very large, elliptic in outline, as in Text-figs. 5-6, 8-9. Hind tibiae deep brown

to black with 2 broad, well defined, very pale bands in proximal third

C. extensipes Karsch (p. 168)

Eyes moderately large, sub-globose, as in Text-figs. 7, 10.
black then not with 2 pale bands in proximal third .

2 Auricles present on internal side of tympanic organs

developed (Text-figs. 12-13). (East Africa) .

Hind tibiae if brown or

rudimentary to quite well

C. aurita sp. n. (p. 167)

Tympanic auricles totally absent ; tympanic organs open on both sides (Text-fig. 1 1).

(West and Central Africa) ..........

Base of fore wings with no dark marking (Text-fig. 14)

Base of fore wings with brown or black mark, sometimes small and inconspicuous,

extending distad from region of second axillary sclerite (Text-figs. 15-17) .

1 mm

FIGS. 5-10. Left eye, 5-7 viewed from the side, 8-10 viewed at about 90° to the plane of
insertion of the eye on the head ; 5, 8, Catoptropteryx extensipes Karsch (Cameroon) ;
6, 9, C. extensipes Karsch (Ghana) ; 7, 10, C. apicalis (Bolivar).

REVISION OF CATOPTROPTERYX

141

Lateral lobes of pronotum with no dark brown or black marking .... 5

Lateral lobes of pronotum dorsad with dark brown to black marking, from single

small dark spot to broken or continuous longitudinal stria .... 6

Lateral lobes of pronotum as in Text-fig. 37, clearly higher than long. Apex of

dorsal valves of ovipositor smooth (Text-fig. 54) C. nigrospinosa (Brunner) (p. 160)
Lateral lobes of pronotum as in Text-fig. 42, not, or only very slightly, higher than

long. Apex of dorsal valve of ovipositor with about 4 very small irregular

teeth (Text-fig. 58) C. serrifera sp. n. (p. 165)

1 mm

FIGS. 11-13. Right tympanic organ viewed from the internal side of the tibia of u,
Catoptropteryx apicalis (Bolivar) ; 12, C. aurita sp. n. (Mozambique) ; 13, C. aurita sp. n.
(Zanzibar).

2nd

axillary

Sc R

15

16

17

FIGS. 14-17. Diagram showing the types of coloration of the fore wing base of Catoptro-
pteryx Karsch : dorsolateral view of left female fore wing of 14, C. capreola Karsch ;
15, C. guttatipes Karsch ; 16, C. punctulata (Karsch) ; 17, C. afra (Karsch).

I42

J. HUXLEY

6 Venation of fore wings as in Text-figs. 60, 61 or 62 ; pattern of parallel veins in

distal half of wing ; RI with more than 2 branches ; MA branched or unbranched 7

Venation of fore wings as in Text-fig. 63 ; veins in distal half not parallel ; RI

bifurcate ; MA unbranched .......... 8

7 Venation of fore wings as in Text-fig. 60 ; MA bifurcate or unbranched ; Rs

usually bifurcate. Area between pronotal disk and longitudinal stria of lateral

lobe orange-yellow (Text-fig. 18) ..... C. nanus sp. n. (p. 147)

18

19

20

21

22

23

24

25

26

27

28

29

30

33

FIGS. 18-33. Lateral view of the pronotum of 18, Catoptropteryx nanus sp. n. ; 19, C.
ramulosa sp. n. ; 20-25, C. capreola Karsch ; 26-30, C. naevia sp. n. ; 31, C. apicalis
(Bolivar) ; 32, C. afra (Karsch) ; 33, C. neutralipennis Karsch.

REVISION OF CATOPTROPTERYX

143

34

35

36

37

38

39

40

41

42

43

44

45

46

47

FIGS. 34-47. Lateral view of the pronotum of 34-36, Catoptropteryx ambigua sp. n. ; 37,
C. nigrospinosa (Brunner) ; 38-39, C. punctulata (Karsch) ; 40, C. guttatipes Karsch ;
41, C. occidentalis sp. n. ; 42, C. serrifera sp. n. ; 43-45, C. auritasp. n. ; 46, C. extensipes
Karsch (Cameroon) ; 47, C. extensipes Karsch (Ghana).

144

J. HUXLEY

Venation of fore wings as in Text-figs. 61-62 ; MA with about 6 branches generated
pectinately ; Rs with 3 branches. Lateral lobes of pronotum as in Text-fig. 19 ;
cuticle same colour above stria as below . . . C. ratnulosa sp. n. (p. 148)

Coloration of pronotal lobes as in Text-figs. 20-25. External spinules of hind
femora blackened only apically (Text-fig. 66). Stridulatory file with about
40-54 teeth. . . . . . . . . C. capreola Karsch (p. 151)

Coloration of pronotal lobes as in Text-figs. 26-30. Some at least of external
spinules of hind femora wholly black, and usually set in dark patches on femoral
carina (Text-fig. 67). Stridulatory file with about 75-106 teeth.

C. naevia sp. n. (p. 153)

48

49

50

52

53

FIGS. 48-53. Lateral view of the ovipositor of 48, Catoptropteryx ramulosa sp. n. ; 49,
C. capreola Karsch ; 50, C. naevia sp. n. ; 51, C. apicalis (Bolivar) ; 52, C. neutralipennis
Karsch ; 53, C. ambigua sp. n.

REVISION OF CATOPTROPTERYX

10

Fore wings as in Text-fig. 64 ; scattered fuscous or fuscescent cells, isolated or in
small groups, in areas R, RI or Rs. Cerci of <$ depressed in apical half (Text-fig.
74). Stridulatory file as in Text-fig. 88, in posterior quarter abruptly sinuose
with very closely spaced teeth. Valves of ovipositor unusually slender (Text-fig.
55) . . . . . . . C. punctulata (Karsch) (p. 161)

Fore wings with no isolated dark cells in areas R, RI or Rs. Cerci of <$ not depressed.
Stridulatory file with no abrupt sinuosity. Valves of ovipositor not unusually
slender ............. 10

Basal mark of fore wings of type shown in Text-fig. 15 : black stripe runs distad

from beneath second axillary sclerite into area M, not centred on M . . . u

Basal mark of fore wings not as above . . . . . . . . 12

54

55

56

57

58

59

FIGS. 54-59. Lateral view of the ovipositor of 54, Catoptropteryx nigrospinosa (Brunner) ;
55, C. punctulata (Karsch) ; 56, C. guttatipes Karsch ; 57, C. occidentalis sp. n. ; 58, C.
serrifera sp. n. (with view of apex of dorsal valve, x 35) ; 59, C. aurita sp. n.

146

J. HUXLEY

64

FIGS. 60-65. Right fore wing of 60, Catoptropteryx nanus sp. n., $ ; 61-62, C. ramulosa
sp. n., $ ; 63, C. capreola Karsch, $ ; 64, C. punctulata (Karsch), ^ ; 65, C. extensipes
Karsch, Q\

REVISION OF CATOPTROPTERYX 147

1 1 Cerci of $ short, strongly sinuose viewed in plane of principal curvature (Text-fig. 75).

Ovipositor as in Text-fig. 56 . . . . . C. guttatipes Karsch (p. 163)

Cerci of $ long and slender, moderately sinuose viewed in plane of principal curvature.

Ovipositor as in Text-fig. 57 . C. occidentalis sp. n. (p. 164)

12 Basal mark of fore wings of type shown in Text-fig. 17 : proximally not extending

as far as second axillary sclerite ..... C. afra (Karsch) (p. 156)
Basal mark of fore wings of type shown in Text-fig. 16 : black stripe runs distad

from beneath second axillary sclerite along centre of M . . . . . 13

13 Hind tibiae wholly dark brown to black ; hind tarsi black or very dark green.

Cerci of <$ black dorsally. Basal mechanism of ovipositor as in Text-fig. 53

C. ambigua sp. n. (p. 159)
Hind tibiae green or yellow-brown. Cerci of <$ red- or yellow-brown, not black

dorsally. Basal mechanism of ovipositor as in Text-figs. 51-52 . . . 14

14 Cerci of <$ asymmetrically inflated at apex (Text-fig. 71). Basal mechanism of

ovipositor as in Text-fig. 51 : inferior margin of lobe of infragonangulum emargin-
ate from lateral viewpoint ..... C. a pica I is (Bolivar) (p. 155)

Cerci of <$ not inflated at apex (Text-fig. 72) . Basal mechanism of ovipositor as in
Text-fig. 52 : inferior margin of lobe of infragonangulum not emarginate from
lateral viewpoint. ..... C. neutralipennis Karsch (p. 157)

DESCRIPTIONS OF THE SPECIES

Catoptropteryx minus sp. n.

(Text-figs, i, 18, 60, 68, 82)

(J. Eyes moderately large, sub-globose, strongly prominent.

Lateral lobes of pronotum as in Text-fig. 18 ; height clearly exceeding length. Fore wings as
in Text-fig. 60 ; RI with about 4 branches arranged pectinately ; Rs usually bifurcate ;
branches of RI and Rs straight and parallel, archedictyon between them denser and with
smaller cells along centre of each area ; MA bifurcate or unbranched. Stridulatory file as in
Text-fig. 82, narrow, almost straight, teeth closely spaced. Tympanic organs without auricles.

Cerci as in Text-fig. 68 ; short, arcuate, circular in transverse section, almost straight viewed
in plane of principal curvature ; apex acute, bearing spinule.

Sides of thorax, and often external surface of hind femora in proximal half, whitish (probably
white in living insect). Lateral lobes of pronotum whitish, with longitudinal black stria, usually
unbroken, dorsad ; area between black stria and disk orange-yellow. Fore wings without
basal or apical markings ; membrane blackened only in cells adjacent to MP + Cui& ; incon-
spicuous striped pattern generated by venation of distal half of wings. Hind wings hyaline
except for apical archedictyon. External spinules of hind femora blackened only apically.

66 67

FIGS. 66-67. Lateral view of part of the left hind femur of 66, C. capreola Karsch ; 67,

C. naevia sp. n.

148 J. HUXLEY

9. As <J except for stridulatory organ and abdominal terminalia. Ovipositor very similar
to that of C. ramulosa sp. n. (Text-fig. 48), but posterior angle of infragonangulum more rounded
and obtuse, and supragonangulum somewhat less salient dorsad ; apical margin of dorsal
valves smooth.

Males Females

Number of specimens examined : 26 25

Total length : 29-1-30-7 32-8-35-0

Median length of pronotum : 3-2-3-6 3'4~3'9

Length of hind femur : 15-1-17-2 16-3-18-0

Length of fore wing : 20-0-21-9 23-2-25-5

Stridulatory file —

number examined : 15

number of teeth (T) : 76-110

length (replica) (F) : 1-43-1-75

T/F : 53-1-71-0

C. nanus sp. n. is very closely related to C. ramulosa sp. n., from which it can be
distinguished by the venation of the fore wings and the coloration of the pronotum ;
the striped pattern of the fore wings is much less marked than in the latter species.
The relation between tooth number and file length is distinctive (see Text-fig, i),
though whether it differs significantly from that in C. ramulosa sp. n. is not yet
known.

The species is known only from Cameroon.

Holotype $. CAMEROON : Efulen, 15.1.1923 (Weber) (ANS Philadelphia).

Paratypes. CAMEROON : — , 1898-1899 (Conradt) i <$ (IEE Madrid) ;

Efulen, 17. xi. 1920 (Weber] i $ (BMNH), 2.vii.i922 (Weber) i <$ (BMNH), 24.x. 1922
(Weber] i J (ANS Philadelphia), i ? (BMNH), io-2i.xi.i922 (Weber] i <£, i $ (BMNH),
4 (J, 2 $ (ANS Philadelphia), xii.i922 (Weber] 3 $, 3 $ (BMNH), 6 £, 9 $ (ANS Phila-
delphia), 1.1923 (Weber] i <£, i $ (BMNH), 3 ^, 3 ? (ANS Philadelphia), 10.11.1923
(Weber] i $ (BMNH), i <?, 2 $ (ANS Philadelphia), 12-22.111.1923 (Weber] i $ (BMNH),
i $ (ANS Philadelphia), ^.1923 (Weber) i $ (ANS Philadelphia), 5.^.1923 (Weber)
i c? (BMNH).

Catoptropteryx ramulosa sp. n.

(Text-figs. 19, 48, 61-62)

cJ. Not known.

$. Eyes moderately large, sub-globose, strongly prominent.

Lateral lobes of pronotum as in Text-fig. 19 ; height clearly exceeding length. Fore wings
as in Text-figs. 61 and 62 ; branches of R\, Rs and MA generated pectinately, sub-parallel,
archedictyon between them denser and with much smaller cells along centre of each area ;
RI with 3 or 4 branches, Rs with 3 branches, MA with about 6 branches. Tympanic organs
without auricles.

Ovipositor as in Text-fig. 48, posterior angle of infragonangulum only slightly greater than
90° ; apical margin of dorsal valves smooth.

Sides of thorax pale green to ivory. Lateral lobes of pronotum not whitish, with somewhat
variable broken black stria dorsad ; cuticle same colour above stria as below. Fore wings

REVISION OF CATOPTROPTERYX

149

68

69

71

70

72

73

74

77

78

FIGS. 68-8 1. Left male cercus viewed (left) from above, at 90° to its plane of principal
curvature, and (right) from the outside, in the plane of principal curvature : 68,
Catoptropteryx nanus sp. n. ; 69, C. capreola Karsch ; 70, C. naevia sp. n. ; 71, C.
apicalis (Bolivar) ; 72, C. neutralipennis Karsch ; 73, C. ambigua sp. n. ; 74, C.
punctulata (Karsch) ; 75, C. guttatipes Karsch ; 76, C. occidentalis sp. n. ; 77, C.
serrifera sp. n. ; 78, C. aurita sp. n. ; 79, C. extensipes Karsch (Cameroon) ; 80, C.
extensipes Karsch (Ivory Coast) ; 81, C. extensipes Karsch (Ghana).

150

J. HUXLEY

82

FIGS. 82-93. Left stridulatory file of 82, Catoptropteryx nanus sp. n. ; 83, C. capreola
Karsch ; 84, C. naevia sp. n. ; 85, C. apicalis (Bolivar) ; 86, C. neutralipennis Karsch ;
87, C. ambigua sp. n. ; 88, C. punctulata (Karsch) ; 89, C. guttatipes Karsch ; 90, C.
occidentalis sp. n. ; 91, C. serrifera sp. n. ; 92, C. aurita sp. n. ; 93, C. extensipes Karsch.
The posterior end of the file is to the left in the drawings.

REVISION OF CATOPTROPTERYX 151

without basal or apical markings ; membrane blackened only in cells adjacent to MP -\-
striped pattern generated by venation, particularly marked in distal half of wings. Hind wings
hyaline except for apical archedictyon. External spinules of hind femora blackened only
apically.

Females

Number of specimens examined : 2

Total length : 33'O-35'O

Median length of pronotum : 3 -5-3 -6

Length of hind femur : 17-3-17-8

Length of fore wing : 23-8-25-5

C. ramulosa sp. n. is very closely related to C. nanus sp. n., from which it can be
readily distinguished by the many-branched MA , pectinate Rs and more conspicuous
pattern of the archedictyon of the fore wings, and by the coloration of the pronotal
lobes. C. nanus sp. n. and C. ramulosa sp. n. may possibly be shown by the evidence
of further material to be geographic variants of the same species. However, in
the absence of unequivocal evidence in this direction at the present, and since the
two forms are clearly delimited in the available material, I prefer to treat them here
as separate species rather than give them subspecific status.

C. ramulosa sp. n. is known only from eastern Congo (Kinshasa).

Holotype $. CAMEROON : 39 km S. of Walikale, 700 m, 25.xii.ig57 (Ross &
Leech) (CAS San Francisco).

Paratype $. CAMEROON : 39 km S. of Walikale, 700 m, I4«ix.i957 (Ross & Leech)
(BMNH).

Catoptropteryx capreola Karsch
(Text-figs, i, 14, 20-25, 49, 63, 66, 69, 83)

Catoptropteryx capreola Karsch, 1896 : 332. Holotype <$, CAMEROON : Lolodorf (ZMHU

Berlin) [examined].
Catoptropteryx immaculipennis Karsch, 1896 : 333. Holotype ^, CAMEROON : Lolodorf (ZMHU

Berlin) [examined]. Syn. n.

cJ. Eyes moderately large, sub-globose, strongly prominent.

Lateral lobes of pronotum as in Text-figs. 20-25 ; height clearly exceeding length. Rs of
fore wings with 2 or rarely more branches as in Text-fig. 63; #1 bifurcate ; MA unbranched.
Stridulatory file as in Text-fig. 83, broad, straight, teeth widely spaced. Tympanic organs
without auricles.

Cerci as in Text-fig. 69 ; long and slender, circular in transverse section, arcuate, not or only
very slightly sinuose viewed in plane of principal curvature ; apex acute, bearing small
spinule ; length somewhat variable.

Lateral lobes of pronotum dorsad with longitudinal dark brown to black stria continuous or
broken and of variable length : most common variants as in Text-figs. 20-22, 24-25 ; Text-fig.
23 illustrates rarer pattern. Fore wings without dark markings at base or apex ; membrane
usually blackened for variable distance on either side of MP + Cwia, and occasionally over whole
of cubito-anal area, but without isolated dark cells in areas R, RI or Rs. Hind wings hyaline
except for apical archedictyon. External spinules of hind femora blackened only apically, as
in Text-fig. 66, and not set in dark patches on femoral carina ; hind femora with numerous
brown spots in distal half. Cerci, epiproct and parts of tenth abdominal tergite often red-
brown or dark brown.

152 J. HUXLEY

?. As <$ except for stridulatory organ and abdominal terminalia. Ovipositor as in Text-fig.

49 ; apical margin of dorsal valves smooth ; infragonangulum large, inflated, usually some-
what flattened dorsolaterally.

Males Females

Number of specimens examined : 184 100

Total length : 3i'5-39'5 34-8-40-8

Median length of pronotum : 3-6-4-5 3'9-4'9

Length of hind femur : 15-9-21-5 17-5-20-4

Length of fore wing : 23-2-30-5 25-1-30-4
Stridulatory file —

number examined : 17

number of teeth (T) : 4°~54

length (replica) (F) : 1-46-1-84

T/F : 27-4-34-4

C. capreola Karsch has a very strong affinity with C. naevia sp. n., from which it
is most reliably distinguished by the coloration of the external spinules of the hind
femora. It also differs from that species in the coloration of the pronotal lobes,
the shape of the male cerci viewed in their plane of principal curvature, and the
number and distribution of teeth in the stridulatory file (see Text-fig, i). The
differences between the two species in the gonangulum of the ovipositor are not
always as clear as in the specimens figured here, being of a similar magnitude to
intraspecific variation and variation in distortion through drying.

The known range of C. capreola Karsch extends across West and Central Africa
from western Liberia to Uganda and northern Angola.

Holotype $. CAMEROON: Lolodorf (Conradt) (ZMHU Berlin).

GUINEA : N.E. end Nimba Range, 4 mis N.W. of Nzo, vii.i963, i 9 ; Nimba,
Ziela, 11.1957, i <$, iii.i957, 3 <$ ; Nimba, vii-xii.i95i, i £. LIBERIA : Marshall
Territory, iii.1955, i 9, ^.1955, i 9, v.io.55, 2 <?, i 9, vi.igss, 3 $, i 9, 11.1956,
i 9, iv.i956, i <?, x.i956, i 9, xii.i956, i <$, iv.1957, i <$ ; N. of Monrovia, Bomi Hills,

5 mis N.E. of mines, Forest Reserve Rest House, vii.1963, 3 <$, 4 9- IVORY COAST :
Adiopodoume, ^.1955, i <$, ix.ig63, i <£, x.1963, i <$, vii.i965, i $, viii.1965, i <$,
ix.i965, 2 9> x.i965, i <$, xi.ig6^, i 9 ; Toumodi, Lamto, 1.1952, i 9, ^.1964, 8 <£,

6 9, x.1966, i $, xii.i966, 2 <$ ; Tai, i.i955, i 9 ; Grabo, 1.1955, i $ ', Nimba, Yale,
380 m, ^.1964, i $ ; Foret du Banco, x.1963, 18 $, 10 9 ; Reserve du Banco, 5 <£,
i 9- GHANA : Tafo, xi.i96i, 8 <$, 3 $, xii.i96i, 5 £, ^.1962, 3 <£, ^.1962, 2 $, i 9,
iv.i957» 3 cJ, 8 9, v.ig57, 8 9, vi.igs;, i $, 3 9, v-vi.ig54, 5 ^, 2 9 ; W. Region, nr.
Wiawso, 30 mis N.W. of Tano Lodge, x.igbo, 5 ^ ; Ashanti Region, nr. Kubease,
Bobiri Forest Res., x.igbo, i <$, Ashanti Region, Bobiri Forest Res., 23 mis S.E. of
Kumasi, xi.1959, i ^ ; Ashanti Region, Bekwai Dist., Numia Forest Res., N. of
Prasu R. Bridge, vii.i962, i $. NIGERIA : W. Prov. n mis E. of Ondo, Owenna
Forest Rest Ho., xii.igGo, 3^, i 9 i W. Prov. University Coll. Ibadan, xii.i96o, i 9 ;
E. Prov., 20 mis N.E. of Calabar, Forest Res., 1.1961, i ^ ; W. Prov., 24 mis S. of
Benin, Sapoba Forestry Sta., 1.1961, i <$ ; nr. Ibadan, Gombar, 1.1965, 5 <$, i 9-
CAMEROON : Lolodorf (Conradt} i ^ (ZMHU) Berlin (holotype of Catoptropteryx
immaculipennis Karsch) ; Efulen, xi.i92o, i ^, vii.i922, i °-» X.IQ22, 3 J, xi.ig22,

REVISION OF CATOPTROPTERYX 153

14 <£, 7 ?, xii.1922, 12 <?, 3 ?, 1.1923, 4c£, i ?, ".1923, i ?, 111.1923, 6^, 4 ?, ^.1923,
i <J, i 9 ; Kribi, 1908, i <$, i $ ; Etandac, xi.ig22, i ^ ; Abong Mbang, X.IQ46,
i r£ ; Ja R., Bitye, i $ ; Victoria, i <$ ; Kumba, ¥1.1959, i <£ ; - — , 1950, i ^ ;

Mt. Cameroon, Post & Telegraph Road, 55 ft, xn.ig6o, i $ ; - — , 1898-1899,

i $. FERNANDO Poo : Basile, 1.1933, i <$. CENTRAL AFRICAN REPUBLIC : Fort
Sibut, 1375 ft, x.1934, i ^. CONGO (BRAZZAVILLE) : M'boko Sogho, i $ ; Sanga R.,
Nola, 1300 ft, x.1934, i <$, 2 $ ; Mayumbe, Dimonika, 1.1964, i $ ; Mts. du Chaillu,
Mbila, xii.i963, 4 <$. CONGO (KINSHASA) : 39 km S. of Walikale, 700 m, ix.i957,
i <$, xii.1957, i <J, 2 $ ; Ubangi, Budjalibala, 11.1949, i <$ ; Region des Lacs, i $,
i $ ; Flandria, xii.i93O, i $ ; Equateur, Boende, vi.i926, i<$ ; Equateur, Bokuma,
1951, i <$ ; Idiofa, Mwilambongo, 1947, i $ ; Boma-Yanga, x.i9i2, i $ ; Hte
Tshuapa, Skela, 1936, i $ ; Bas Congo, Luki, vi-vii.i952, i $ ; Avakubi, x.i9i2,
i c£ ; Kivu, Costermansville, 1951, i $ ; W. Kivu, Walungu, 1939, i <$ ', Eala,
iv-v.i932, i $ ; Kibali-Ituri Dist., 10 mis W. Epulu R. Ferry, Irumu-Avakubi Rd.,
Saidi, 2800 ft, ix.i934, 3^, i $ ; Kibali-Ituri Dist., betw. Mambasa & Saidi, Epulu R.
Ferry, 2500 ft, ix.i934, 2$ ; Medje, 2°25' N., 27°i5' E., viii.igio, i $. UGANDA :
— , i <$, i 9. ; Zika Forest, viii.1963, 2 <$, x.1963, i •$ ; Zika Forest, 7 mis from
Entebbe, 40 ft level, iii-vi.i96i, i <$, 80 ft level, iii-vi.igGi, i ^, vii.ig6i, i $ ;
Entebbe, x.1914, i $ ; Mfanga Forest Res., viii.1964, i $. ANGOLA : Congulu,
iv.1934, i cJ, i ?.

Catoptropteryx naevia sp. n.

(Text-figs, i, 26-30, 50, 67, 70, 84)

cj. Eyes moderately large, sub-globose, strongly prominent.

Lateral lobes of pronotum as in Text-figs. 26-30 ; height clearly exceeding length. Rs of
fore wings with 2 or rarely more branches as in Text-fig. 63 ; RI bifurcate ; MA unbranched.
Stridulatory file as in Text-fig. 84, broad, slightly curved, teeth closely spaced. Tympanic
organs without auricles.

Cerci as in Text-fig. 70 ; long and slender, circular in transverse section, arcuate, more or
less strongly sinuose viewed in plane of principal curvature ; apex acute, bearing small spinule ;
length somewhat variable.

Lateral lobes of pronotum as in Text-fig. 26, with single black spot dorsad in anterior half ;
less commonly with additional dark brown to black markings in longitudinal line as in Text-
figs. 27-28, and rarely as in Text-figs. 29-30, but spot corresponding in position to that in
commonest variant always most conspicuous of pronotal markings. Fore wings without dark
markings at base ; membrane usually blackened for variable distance on either side of MP +
Cwia, but without isolated dark cells in areas R, RI or Rs ; small fuscous spot occasionally
present at apex. Hind wings hyaline except for apical archedictyon. Some at least of external
spinules of hind femora wholly black and usually set in dark brown to black patches on femoral
carina, as in Text-fig. 67 ; hind femora with numerous brown spots in distal half. Cerci,
epiproct and parts of tenth abdominal tergite often red-brown or dark-brown.

$. As (J except for stridulatory organ and abdominal terminalia. Ovipositor as in Text-fig.
50 ; apical margin of dorsal valves smooth ; infragonangulum large, inflated, usually roughly
prolate-spheroidal .

154 J- HUXLEY

Males Females

Number of specimens examined : 117 32

Total length : 37-5-44-2 37'5-45'5

Median length of pronotum : 4'5-5'6 4-4-5-4

Length of hind femur : 20-1-23-2 19-1-23-5

Length of fore wing : 28-6-34-3 27-9-33-6

Stridulatory file —

number examined : 15

number of teeth (T) : 75-106

length (replica) (F) : 1-66-2-18

T/F : 39-5-547

This species is very similar to C. capreola Karsch, from which, however, it shows a
consistent distinction in the coloration of the external spinules of the hind femora,
and also in the number and distribution of teeth in the stridulatory file (see Text-
fig, i). Other real though perhaps less clear-cut differences exist in the coloration
of the pronotal lobes and the shape of the male cerci. The infragonangulum of the
ovipositor tends to be more inflated than in C. capreola Karsch, but this is a rather
subtle distinction that may be masked by the distortion of the basal sclerites due to
drying. Although there is an overlap of the ranges of all the dimensions measured,
C. naevia sp. n. is appreciably larger on average than C. capreola Karsch.

C. naevia sp. n. is distributed in West Africa from Sierra Leone to Ghana.
Holotype^. LIBERIA : Marshall Territory, 30. vii. 1955 (Fox) (ANS Philadelphia).

Paratypes. GUINEA : Forest de Dieke, 20 mis N. of Dieke, 24. vii. 1963 (Jago)
i <$ (BMNH) ; Nimba, Ziela, 25^.1957 (Lamotte, Amiet & V anderplaetsen) i <£
(MNHN Paris). SIERRA LEONE : Kenema, 31.^1.1946 (Jordan) i $ (BMNH) ;
Njala, i8.iv.i929 (Hargreaves) i $ (BMNH). LIBERIA : Marshall Territory, 27. xi.
1954 (Fox) i $ (BMNH), 27.xii.i954 (Fox) i $ (BMNH), 17-28.1.1955 (Fox) i <?,
i $ (BMNH), 3 <$, i $ (ANS Philadelphia), i2-i6.ii.i955 (Fox) i <$ (BMNH), 2 <$,
i $ (ANS Philadelphia), i-28.iii.i955 (Fox) 2 £ (BMNH), 8 # (ANS Philadelphia),
I9~27.iv.i955 (Fox) i 3 (BMNH), 3 $ (ANS Philadelphia), 17^.1955 (Fox) i $
(BMNH), 16-27^1.1955 (Fox) 2 <$ (BMNH), 3 <J (ANS Philadelphia), 2-3O.vii.i955
(Fox) 2 <$ (BMNH), 6 c?, i $ (ANS Philadelphia), 20-28. viii. 1955 (Fox) i £ (BMNH),
2£ (ANS Philadelphia), 6-22.ix.i955 (Fox) i $, i ? (BMNH), i <J (ANS Philadelphia),
n-23.x.i955 (Fox) i £ i ? (BMNH), 4 <£ (ANS Philadelphia), 5-i6.xi.ig55 (Fox)
i c? (BMNH), i(?,i$ (ANS Philadelphia), 4-25.xii.ig55 (Fox) i <? (BMNH), 2 £, 2 $
(ANS Philadelphia), 4-17.1.1956 (Fox) i <$ (BMNH), 3 <$, i $ (ANS Philadelphia),
i-i8.ii.igs6 (Fox) 2 <$, i $ (BMNH), 6 <J, 2 $ (ANS Philadelphia), I5.iii.ig56 (Fox)
i ^ (BMNH), 4-25.iv.igs6 (Fox) i ^ (BMNH), 4^ (ANS Philadelphia), 3-ig.v.igs6
(Fox) i c? (BMNH), i & i ? (ANS Philadelphia), i.vii.igs6 (Fox) i $ (ANS Philadel-
phia), 28-3I.X.I956 (Fox) i £ (BMNH), 2 £ (ANS Philadelphia), 7.xi.igs6 (Fox)
i $ (BMNH), i c? (ANS Philadelphia), is.xii.igs6 (Fox) i $ (ANS Philadelphia),
6-31.1.1957 (Fox) i ^ (BMNH), 2 ^ (ANS Philadelphia), 3-28.ii.ig57 (Fox) i £
(BMNH), 2 ^ (ANS Philadelphia), ig-2i.iii.ig57 (Fox) i £ (BMNH), i £ (ANS
Philadelphia), 8.iv.ig57 (Fo*) i <J (BMNH), i-2o.v.ig57 (Fox) i <J (BMNH), i J
(ANS Philadelphia) ; N. of Monrovia, Bomi Hills, 5 mis N.E. of mines, Forest Res.

REVISION OF CATOPTROPTERYX 155

Rest Ho., 23. ¥11.1963 (Jago) 4 <$, 2 $ (BMNH) ; Bindah, forest edge, 6.1v.i92o
(Barrett] i <$ (BMNH). IVORY COAST : Adiopodoume, 24.111.1890 (Vuillaume) i $
(ORSTOM Abidjan) ; Foret du Banco, I5.X.I963, i $ (BMNH), i <J (ORSTOM
Abidjan), I2.V.I964, i^ (ORSTOM Abidjan). GHANA : Tafo, 2-30. xi. 1961 (Gardner]
3 <?, i $ (BMNH), i-26.xii.i96i (Gardner] i <J, 2 $ (BMNH), 1-29.1.1962 (Gardner]
X £, 2 $ (BMNH), 3-25.11.1962 (Gardner] 2 £, 3 $ (BMNH), 3.111.1962 (Gardner) i $
(BMNH), I9.iv.i957 (Eastop) i $ (BMNH), 31^.1957 (£asfo/>) i <? (BMNH), 7-2i.vi.
1957 (Eastop) 2 c? (BMNH), v-vi.i954 (Williams) 5 <J (BMNH) ; Ashanti, Bobiri
Forest Res., 23 mis S.E. of Kumasi, 21. xi. 1959 (Jago) i $ (UG Accra) ; E. Region,
Kade Agr. Res. Sta., 9.vii.i963 (Acheampong) i $ (UG Accra) ; E. Region, Tafo,
W.A. C.R.I., x.i96i (Jago) i J (UG Accra). GHANA ? : Fantee Country, Bosso
(Jones) i £ (BMNH).

Catoptropteryx apicalis (Bolivar)
(Text-figs, i, 7, lo-n, 31, 51, 71, 85. Plate i, fig. 2)

Caedicia apicalis I. Bolivar, 1893 : 177. Holotype $, IVORY COAST : Assinie (IEE Madrid)

[examined] .
Catoptropteryx signatipennis Karsch, 1896 : 333. Lectotype <J, CAMEROON : Victoria (ZMHU

Berlin) [examined]. Syn. n.
Catoptropteryx apicalis (Bolivar) Kirby, 1906 : 416.

cj. Eyes moderately large, sub-globose, strongly prominent.

Lateral lobes of pronotum as in Text-fig. 31 ; height clearly exceeding length. Rs of fore
wings as in Text-fig. 63, with 2 or rarely 3 branches ; R\ bifurcate ; MA unbranched. Stri-
dulatory file as in Text-fig. 85 ; broad, gently curved, teeth closely spaced. Tympanic organ
without auricles.

Cerci as in Text-fig. 71 ; long, robust, arcuate, circular in transverse section, sinuose viewed
in plane of principal curvature ; apex inflated behind large terminal spinule.

Lateral lobes of pronotum usually unmarked but very rarely with faint trace of longitudinal
stria dorsad. Basal mark of fore wings of type shown in Text-fig. 16 : continuous black stripe
about 1-0-2-5 mm l°ng runs distad from beneath second axillary sclerite along centre of M ;
variable number of fuscous cells on either side of MP + Cu\& ; no isolated dark cells in areas
R, RI or Rs ; fuscous spot at apex usually well developed, very rarely absent. Hind wings
hyaline except for apical archedictyon. Spinules of hind femora yellow or light brown at base,
black at tip and set in black patches on femoral carina, or wholly black with or without dark
area surrounding base. Tenth abdominal tergite, epiproct and cerci dark red or red-brown.

$. As <J except for stridulatory organ and abdominal terminalia. Ovipositor as in Text-fig.
51 ; infragonangulum prominent ventrad, with well developed ventral concavity causing
inferior margin of lobe to appear more or less deeply emarginate from lateral viewpoint ;
apex of dorsal valves smooth.

Males Females

Number of specimens examined : 168 131

Total length : 41-0-46-0 44-0-50-6

Median length of pronotum : 4'5-5'° 4'8-5'7

Length of hind femur : 20-2-23-4 21-4-25-9

Length of fore wing : 30-0-33-8 32-8-36-4

Stridulatory file —

number examined : 20

number of teeth (T) : 79-108

length (replica) (F) : 2-06-2-48

T/F : 35-4-46-0

156 J. HUXLEY

C. apicalis (Bolivar) is very similar to C. neutralipennis Karsch. The males of
the two species are distinguishable clearly only by the form of the apex of the cerci.
Female specimens may be very difficult to distinguish and it is sometimes necessary
to place some reliance on their association with identifiable males. See also dis-
cussions under C. neutralipennis Karsch and C. afra (Karsch).

The known range of C. apicalis (Bolivar) extends across tropical Africa from
Sierra Leone to Uganda.

Lectotype Designation. Of the two syntypes of Catoptropteryx signatipennis
Karsch, I have selected and labelled the male specimen as the LECTOTYPE.

Holotype 9- IVORY COAST : Assinie (Alluaud) (IEE Madrid).

GUINEA : Nimba, Ziela, 11.1957, i $. SIERRA LEONE : Freetown, xi.igGo, i 9>
xii.i966, i 9- LIBERIA : N. of Monrovia, Bomi Hills, 5 mis N.E. of mines, Forest
Reserve Rest House, vii.i963, 7 $, i $. IVORY COAST : Foret du Banco, x.1963,
26 c£, 14 9, V.I9&4, 3 (J, 3 9 ; Reserve du Banco, i <£ ; Mokta, vi.i964, i ^ ; Adio-
podoume, v.1954, 2 £, iv.i955, i $, x.ig6^, 2 $, i $, xi.i963, i $, ^.1964, i $, v.1965,
5 <$, i 9, vi.i965, i <J, viii.1965, 2 <J, ix.igGs, 4 $, 2 9> x.i965, i $, xi.igGs, 2 $, 1.1966,

3 <£, 2 9, ii.i966, i <$, xii.i966, i £ ; Lamto, Toumodi, 111.1964, i $, ^.1964, 5 <?,
23 9, xii.i965, i (?, 2 9> x.1966, 2 ^ ; Niangon, 111.1967, i $, i $ ; Apolo Piege, v.igGo,

1 $ ; Barrage d'Ayame, ^.1964, i $. GHANA : Accra, xii.1963, i ^ ; Kade Agr.
Res. Sta., vii.1963, 3 <$, 5 $ ; Kumanin, 3 mis from Kade Agr. Res. Sta., i ^, i $ ;
Volta Region, Amedzofe, xi.i963, i $ ; Trans- Volta Togoland, Kpandu Rest
House, xii.i959, i $ ; W. Region, near Wiawso, 30 mis N.W. of Tano Lodge, x.ig6o,

2 $ ; Bibianaha, ix-xii.igog, i $ ; Tafo, v-vi.i954, 2 $, vi.i955, i ^, iv.i957, i 9,
V.I957, i (J, 4 ?, vi^QS?. J3 cJ. 5 ?. xi.ig6i, 8 $, 2 ?, xii.igGi, 5 cJ, 2 ?, 1.1962, i <J,

4 $, 11.1962, 6 $, 3 9> 111.1962, 16 ^, 4 9- NIGERIA : near Ibadan, Gombar, 1.1965,

5 J, 7 9 ; Ibadan, x.i965, i 9- CAMEROON : Efulen, xi.i92O, 1^,1$, ix.i922,
i 9, X.IQ22, i <J, 2 9, xi.i922, 5 ^, 3 9, xii.i922, 15 ^, 4 9, 1.1923, 5 ^, 111.1923, 3 c?
iv.i923, i (J, v.1923, i 9 i Etandac, xi.i922, i J ; Victoria, Muyuka, vi.i949, i 9 ;
Victoria, Mabete, v-vi.i949, i 9 ; Victoria (Preuss) i ^ (lectotype of Catoptropteryx
signatipennis Karsch) (ZMHU Berlin) ; Lolodorf (Conradt) i 9 (paralectotype of
Catoptropteryx signatipennis Karsch) (ZMHU Berlin) ; Jabassi, iv.i85o, i 9 i
Johann-Albrechtshohe, 1.1896, i 9- SPANISH GUINEA : Rio Manyani, vi.igig, i J.
CONGO (BRAZZAVILLE) : Mts. du Chaillu, Mbila, xii.1963, i <$, 3 9 ', Brazzaville,
xi.i963, 2 9. 1.1964, i c?, 11.1964, i 9 ', Bassin de la Sangha, 1899, i ^. CONGO
(KINSHASA) : Kibali-Ituri Dist., 10 mis W. Epulu R. ferry, Irumu-Avakubi Rd.,
Saidi, ix.i934, i 9 ; Congo River, left bank, Lukolela, i°5' S., 1.1921, i 9- UGANDA :
7 mis from Entebbe, Zika Forest, ground level on steel tower, Hi. -vi. 1961, i <$,
40 ft level, i (J.

Catoptropteryx afra (Karsch)

(Text-figs. 17, 32)

Caedicia afra Karsch, 1889 : 446. Holotype $, NIGERIA : Benue1 (IZPAN Warsaw)
[examined].

1 The original description gives this locality, though the holotype itself is labelled merely ' Niger '.

REVISION OF CATOPTROPTERYX 157

Catoptropteryx afra (Karsch) Karsch, 1896 : 335 (footnote).

$. Not known.

$. Eyes moderately large, sub-globose, strongly prominent.

Height of lateral lobes of pronotum clearly exceeding length . Rs of fore wings bifurcate, as
in Text-fig. 63 ; RI bifurcate ; MA unbranched. Tympanic organ without auricles.

Ovipositor not distinguishable from that of C. neutralipennis Karsch (Text-fig. 52) ; inferior
margin of infragonangulum lobe not emarginate from lateral viewpoint ; apex of dorsal valves
smooth.

Lateral lobes of pronotum without dark markings. Basal mark of fore wings of type shown
in Text-fig. 1 7 : brown to black (in one specimen very faint, reddish) mark at base of M, about
0-4-0-8 mm long (0-4 mm in holotype), proximally not extending as far as second axillary
sclerite. Cells on either side of MP + Cum darkened ; apex without dark spot. Hind wings
hyaline except for apical archedictyon. Spinules of hind femora black only at tip, not set in
dark patches on femoral carina.

Holotype ? Other $?

Number of specimens examined : i 4

Total length : 50-3 40-2-46-2

Median length of pronotum : 6-2 4-7-5-2

Length of hind femur : 24-6 19-7-23-0

Length of fore wing : 37-4 29-5-34-1

The identity of the holotype is not at all plain. It resembles C. apicalis (Bolivar)
in size and general appearance, but the form of the basal mechanism of the ovi-
positor is not distinguishable from that of C. neutralipennis Karsch, and in the
coloration of the fore wing base it differs slightly from either of these species. C.
afra (Karsch) is almost certainly a synonym of either C. apicalis (Bolivar) or C.
neutralipennis Karsch (in fact, the last two may also prove to be one species — see
discussion under C. neutralipennis Karsch) ; however, without males that could be
confidently associated with the holotype, I have been unable to resolve the problem
here. The four other females that I have included under this species agree with the
holotype in the form of the ovipositor and the coloration of the fore wings and
hind femora, but are considerably smaller.

C. afra (Karsch) is known only from Nigeria.

Holotype $. NIGERIA : Benue (IZPAN Warsaw).

NIGERIA : Ibadan, x.ig56, i $, ^.1965, i $, xii.i966, i $, viii.1967, i $.

Catoptropteryx neutralipennis Karsch
(Text-figs, i, 33, 52, 72, 86)

Catoptropteryx neutralipennis Karsch, 1896 : 334. Lectotype <$, TOGO : Misahohe (ZMHU
Berlin) [examined].

<$. Eyes moderately large, sub-globose, strongly prominent.

Lateral lobes of pronotum as in Text-fig. 33 ; height clearly exceeding length. Rs of fore
wings bifurcate as in Text-fig. 63 ; RI bifurcate ; MA unbranched. Stridulatory file as in
Text-fig. 86 ; broad, gently curved, teeth closely spaced. Tympanic organ without auricles.

Cerci as in Text-fig. 72 ; long, slender, arcuate, circular in transverse section, more or less
sinuose viewed in plane of principal curvature ; apex more or less obtuse, but without con-
spicuous inflation, bearing small terminal spinule.

I58 J. HUXLEY

Lateral lobes of pronotum without dark marking. Basal mark of fore wings of type shown
in Text-fig. 16 : continuous black stripe, about 1-0-2-5 mm l°ng> runs distad from beneath
second axillary sclerite along centre of M ; variable number of fuscous cells on either side of
MP -\- Cum ; no isolated dark cells in areas R, R± or Rs ; fuscous spot at apex usually poorly
developed or absent, rarely well developed. Hind wings hyaline except for apical archedictyon.
Spinules of hind femora black at tip only, at tip and base, or wholly black, sometimes set in
dark patches on femoral carina. Cerci red- or yellow-brown.

$. As $ except for stridulatory organ and abdominal terminalia. Ovipositor as in Text-fig.
52 ; inferior margin of lobe of infragonangulum not emarginate from lateral viewpoint ; apex
of dorsal valves smooth.

Males Females

Number of specimens examined : 73 60

Total length : 37'8-43'3 37'7-44<(>

Median length of pronotum : 4-1-4-9 4-3-4-8

Length of hind femur : 18-6-21-8 19-5-22-3

Length of fore wing : 27-7-31-5 28-5-31-9

Stridulatory file —

number examined : 21

number of teeth (T) : 64-101

length (replica) (F) : i'73-2'37

T/F : 34-0-42-6

This species can be distinguished with certainty from C. apicalis (Bolivar) only
by the shape of the cereal apex in the male. Identification of female specimens
may be impossible, for while the ovipositors of the two species may have quite
different superficial appearances, their structure is essentially similar and many
specimens appear intermediate when compared with the figures given here. The
differences may be due to the effects of very slight differences in the relative growth
rates of the basal sclerites on the manner of distortion of the integument during
development. Such small differences in relative growth rates need not of course be
between species, and are quite likely to be intraspecific. They are also likely to be
related to the absolute dimensions of the adult insect, and since C. apicalis (Bolivar)
is larger on average than C. neutralipennis Karsch, a size-related feature could falsely
appear to be a specific character. There are some grounds for believing that this is
probably the case here, and since the females have been segregated entirely on the
appearance of the basal mechanism of the ovipositor, the division into ' species '
may be in effect no more than a separation of the larger specimens from the smaller,
and as such does not necessarily correspond in any way to the division between the
two male forms, which is based on a discontinuity in the form of the cerci. In both
sexes the two species are so similar in all other features that the possibility must be
borne in mind that they and C. afra (Karsch) may all belong to one species, dimorphic
in the male sex over part of its range, and highly variable in size. At present,
however, there is not sufficient evidence to justify synonymizing the three names.

C. neutralipennis Karsch is distributed in West Africa from Sierra Leone to
Cameroon.

Lectotype Designation. Of the two syntypes of Catoptropteryx neutralipennis
Karsch, I have selected and labelled the male specimen as the LECTOTYPE.

REVISION OF CATOPTROPTERYX 159

Lectotypec?. TOGO : Misahohe, 26. vi. 1894 (Baumann) (ZMHU Berlin).
Paralectotype °.. TOGO : Bismarckburg, 3o.x.-5.xi.i893 (Conradt) (ZMHU
Berlin).

GUINEA : Nimba, vii-xii.i95i, i $ ; Nimba, Ziela, 11.1957, i <$, 111.1957, 4 <$,

1 $, v.igsy, 6 (J, 2 ?. SIERRA LEONE : Freetown, 1.1956, i $, X.IQ66, i $ ; Njala,
viii.1926, i <£. LIBERIA : N. of Monrovia, Bomi Hills, 5 mis N.E. of mines, Forest
Reserve Rest House, vii.1963, 3 <£, i ? ; Marshall Territory, 111.1955, i $, iv.i955,
ic?,i$, V.IQ55, 3c?, 2 ?, vi.i955, 3^, vii.igss, 2 $, ix.igss, 3^, X.IQSS, 2 ?, 111.1956,

2 ?, iv.1956, 3 (J, i ?, v.i956, i $, vi.ig56, i ?, 1.1957. * <J. m'-i957> * ?• IVORY
COAST : Foret du Banco, X.I9&3, 1^,1$; Lamto, Toumodi, ^.1964, 4 ^, 2 $ ;
Yale, ^.1964, i $ ; Adiopodoume', 11.1967, i $ ; S^guela, xii.i964, i $ ; Mokta,
vi.i964, i $. GHANA : Trans- Volta Togoland, Kpandu, xii.1959, i <$ ; E. Region,
Kade Agr. Res. Sta., vii.1963, 4 <$, 8 $ ; Kumanin, 3 mis from Kade Agr. Res.
Sta., vii.1963, i $ ; W. Region, Sefwi-Bekwai, Shell filling sta., x.i96o, i <$ ; Bibia-
naha, ix-xii.igog, i <$ ; Ashanti, near Nabaume, 6°37'N, i°i7'W, viii.1957, i $ ;
Tafo, v-vi.i954, i <J, 2 $, ^.1957, 3 ^, 2 $, v.i957, 2 ^, 7 $, vi.i957, 8 <J, 9 $, xi.igSi,

3 (J, xii.i96i, 2 (J, 3 $, 1.1962, i <^, 111.1962, i ^, i $. NIGERIA : W. Province, Ibadan,
xii.i96o, i $ ; 24 mis S. of Benin, Sapoba Forestry Sta., 1.1961, i $ ; Bende, i $.
CAMEROON : Efulen, xi.i922, 2 <$, i $, 111.1923, i $.

Catoptropteryx ambigua sp. n.

(Text-figs, i, 34-36, 53, 73, 87)

(£. Eyes moderately large, sub-globose, strongly prominent.

Lateral lobes of pronotum as in Text-figs. 34-36 ; height clearly exceeding length. Rs of
fore wings as in Text-fig. 63, bifurcate ; .7?i bifurcate ; MA unbranched. Stridulatory file as
in Text-fig. 87, narrow, almost straight, teeth closely spaced. Tympanic organs without
auricles.

Cerci as in Text-fig. 73 ; moderately long, circular in transverse section, arcuate, sinuose
viewed in plane of principal curvature ; apex obtuse, bearing small spinule.

Lateral lobes of pronotum dorsad with variable longitudinal stria, as in Text-figs. 35-36,
black to faint red, occasionally totally absent. Basal mark of fore wings of type shown in
Text-fig. 1 6 : continuous black stripe runs distad from beneath second axillary sclerite along
centre of M ; broad band of fuscous cells on either side of MP + Cui& ; no isolated dark cells
in areas R, RI or Rs ; fuscous spot at apex usually very well developed, more rarely faint or
absent. Hind wings hyaline except for apical archedictyon. Spinules of hind femora wholly
black and set in black patches on femoral carina, as in Text-fig. 67 ; hind femora usually with
numerous fuscous spots in distal half ; hind tibiae wholly dark grown to black, sometimes
lighter ventrally ; hind tarsi black to very dark green. Cerci red-brown ventrally, black
dorsally.

$. As (J except for Stridulatory organ and abdominal terminalia. Ovipositor as in Text-fig.
53 ; lobe of infragonangulum small compared with first gonocoxa ; apical margin of dorsal
valves smooth.

i6o

J. HUXLEY

Number of specimens examined

Total length :

Median length of pronotum :

Length of hind femur :

Length of fore wing :

Stridulatory file —

number examined :
number of teeth (T) :
length (replica) (F) :
T/F :

Males

7

35-4-38-9
37-4-3
19-1-20-6
25-7-28-1

6

63-80
1-47-1-83
35-0-44-4

Females

3
37-8-40-5

4-3-4-7

19-0-22-2

27-6-30-8

C. ambigua sp. n. is similar in many respects to C. neutralipennis Karsch and to
C. pundulata (Karsch). There is no possibility of confusion with the latter species
which has a unique coloration of the fore wings. It may be distinguished from
C. neutralipennis Karsch by the ovipositor, the coloration of the hind legs, and the
pronotal stria when this is present. The ovipositor is rather similar to that of
C. nigrospinosa (Brunner) but may be distinguished easily enough. The locality
may be helpful in identifying difficult specimens (e.g. badly discoloured males),
since the range of the species does not appear to overlap those of its most easily
confused relatives.

C. ambigua sp. n. is known only from Cameroon, Congo (Kinshasa) and Uganda.
Holotype <j>. UGANDA : Bwamba, Ntandi, ii.igGS (BMNH).

Paratypes. CAMEROON : Ja River, Bitye, vi-vii.igog, dry season, i $ (ANS
Philadelphia). CONGO (KINSHASA) : Sankuru, Komi, vii.i93o (Ghesquiere] i <$
(MRAC Tervuren) ; Busira, vi.i936 (Ghesquiere} i <$ (BMNH) ; 73 mis E. of Kama,
viii.1957 (Ross & Leech] i $ (BMNH) ; Flandria, xii.ig3O (Hulstaert] i $ (MRAC
Tervuren). UGANDA :— — , i <$ (BMNH) ; Bwamba, Ntandi, ^.1968, i $
(BMNH) ; Bwamba, v.1956 (Carcasson) i <$ (CM Nairobi) ; 7 mis from Entebbe,
Zika Forest, lakeside swamp, 40 ft level on steel tower, iii-vi.igGi (Corbet] i <£
(BMNH).

Catoptropteryx nigrospinosa (Brunner)
(Text-figs. 37, 54)

Caedicia nigro-spinosa Brunner, 1891 : 97. Holotype $, CAMEROON (IZPAN Warsaw).

[examined].
Catoptropteryx nigrospinosa (Brunner) Kirby, 1906 : 415.

o*. Not known.

$. Eyes moderately large, sub-globose, strongly prominent.

Lateral lobes of pronotum as in Text-fig. 37 ; height clearly exceeding length. Rs of fore
wings with 2 branches as in Text-fig. 63 ; R\ bifurcate ; MA unbranched. Tympanic organs
without auricles.

Ovipositor as in Text-fig. 54 ; apical margin of dorsal valves smooth.

Fore wings without dark markings at base ; membrane blackened for short distance on
either side of MP + Cui&, but without isolated fuscescent cells in areas R, RI or Rs ; small
fuscous spot at apex. Hind wings hyaline except for apical archedictyon. Spinules of hind
femora wholly black and set in dark brown to black patches on femoral carina ; hind femora
with numerous brown spots in distal half. Hind tibiae somewhat darkened dorsally.

REVISION OF CATOPTROPTERYX 161

Females

Number of specimens examined : 2

Total length : 41 -0-43-0

Median length of pronotum : 4-5-4-8

Length of hind femur : 20-3-20-4

Length of fore wing : 29-4-31-0

The structure of the ovipositor shows an affinity with C. ambigua sp. n., but the
coloration of C. nigrospinosa (Brunner) clearly distinguishes it from all the other
species of the genus.

This species is known only from Cameroon.

Holotype $. CAMEROON (IZPAN Warsaw).

CAMEROON : Efulen, 4.xii.i922, i $ (ANS Philadelphia).

Catoptropteryx punctulata (Karsch)
(Text-figs, i, 16, 38-39, 55, 64, 74, 88)

Caedicia punctulata Karsch, 18900 : 260. Holotype $, CAMEROON : Kribi (ZMHU Berlin) (in

alcohol) [examined].
Catoptropteryx maculipennis Karsch, 1896 : 333. Holotype $, CAMEROON : Lolodorf (ZMHU

Berlin) [examined]. Syn. n.
Catoptropteryx punctulata (Karsch) Kirby, 1906 : 415.

o\ Eyes moderately large, sub-globose, strongly prominent.

Lateral lobes of pronotum as in Text-figs. 38-39 ; height clearly exceeding length. Fore
wings as in Text-fig. 64 ; Rs bifurcate ; RI bifurcate ; MA unbranched. Stridulatory file as
in Text-fig. 88 ; broad, curved, teeth widely spaced over three quarters of length ; in posterior
quarter, file abruptly sinuose, with closely spaced teeth. Tympanic organs without auricles.

Cerci as in Text-fig. 74 ; short, arcuate, depressed in apical half, strongly sinuose viewed in
plane of principal curvature ; apex acute, bearing terminal spinule.

Lateral lobes of pronotum as in Text-figs. 38-39 ; variable, continuous or broken, longi-
tudinal brown to black stria dorsad, very rarely faint or absent. Basal mark of fore wings of
type shown in Text-fig. 16 : continuous black stripe usually about 6 mm long, sometimes
shorter, runs distad from beneath second axillary sclerite along centre of M and MP ; band of
fuscous cells on either side of MP + Cu\& usually broad and giving fore wings purple-brown
appearance ; in areas R, RI and Rs variable number of scattered fuscous to fuscescent cells,
isolated or in small groups ; fuscous spot at apex usually very well developed. Hind wings
usually faintly fumose, darker towards margin of anal fan, sometimes hyaline, except for apical
archedictyon. External spinules of hind femora usually wholly black, but not set in dark
patches on femoral carina. Hind tibiae dorsally more or less fuscescent, with darker area at
base of each spinule ; rarely, entirely fuscous. Tenth abdominal tergite with pair of large,
dark brown spots, or single median dark area ; epiproct dark brown ; cerci yellow-brown,
dark brown to black dorsally.

$. As o* except for stridulatory organ and abdominal terminalia. Ovipositor as in Text-fig.
55 ; first gonocoxa large, very slightly concave ; valves slender ; apex of dorsal valve smooth.
Basal mark of fore wings generally shorter than in <$, about 0-5-5-0 mm long.

1 62

J. HUXLEY

Males
150

34-3-38-5
4-0— 4-6
17-6-20-8
25-5-29-6

20

32-43
1-23-1-56

Females

73

35-8-40-8
4-0-4-8
18-8-21-6
25-8-29-7

Number of specimens examined

Total length :

Median length of pronotum :

Length of hind femur :

Length of fore wing :

Stridulatory file —

number examined :
number of teeth (T) :
length (replica) (F) :
T/F :

This is one of the most clearly defined species in the genus, being conspicuously
unique in the coloration of the fore wings, and the form of the male cerci, stridulatory
file and ovipositor valves. The basal mechanism of the ovipositor is very similar
to that of C. guttatipes Karsch, differing from this chiefly in the deeper concavity of
its first gonocoxa. It is best distinguished from this species by the more slender
ovipositor, the depressed cerci of the male, and the type of basal mark of the fore
wings. The number and depth of colour of the dark cells of the fore wings are
very variable, and the isolated spots characteristic of the species are not always
easily discerned ; a piece of white paper held behind the wing is sometimes helpful
in detecting these in badly faded specimens.

The known range of C. punctulata (Karsch) extends across tropical Africa from
Sierra Leone to Uganda.

Holotype $. CAMEROON : Kribi, xii.i8882 (Morgen) (ZMHU Berlin) (in alcohol).

GUINEA : Nimba, vii-xii.i95i, 1^,1$, 1951, i <$ ; Nimba, Zie"la, 11.1957, i <$,
iii.i957, 6 <$, v.1957, 7 <$ ; Nimba, Camp Gouan, 1.1957, i $ ; Nimba, Keoulenta,
xii.i956, i <$. SIERRA LEONE : Freetown, Mt. Aureol, i.i956, i $ ; Njala, x.1928,
i <$, iii.i933, i (£ ; Monts Loma, Foret Camp, 1070 m, v.i963, i <$. LIBERIA :
Mt. Coffee, iii.i897, i <$ ; N. of Monrovia, Bomi Hills, 5 mis N.E. of mines, Forest
Reserve Rest House, vii.1963, i ^ ; Marshall Territory, ^.1955, i $, iv.1955, i $,
vi.i955, i (?, i $, vii.i955, 3 <?, viii.1955, i ?, ix.igss, i <?, 1.1956, i <£, 11.1956, i £,
iv.i956, 3 $, x.i-956, i <£, 1.1957, i <$, v.i957, i <£. IVORY COAST : Nimba, ^.1964,
i <$ ; Foret du Banco, x.i963, 5 $, 3 $, v.1964, i <$ ; Reserve du Banco, 2 $ ; Adio-
podoume, 1.1953, i ?, xii.i953, i $, iv.igss, i $, xi.i963, i <$, v.ig6$, 2 <$, vi.io.65,
1$, viii.1965, i^, x.1965, 2^, 111.1967, i $ ; Lamto, Toumodi, ^.1964, i<^ ; N'dzida,
vi.i952, i $ ; 6 km from Tai, 1.1955, 2 $ ; Azaguie, xi.i963, i °-. GHANA : W.
Region, nr. Wiawso, 3 mis N.W. of Tano Lodge, x.i96o, 2 $ ; Ashanti, Bobiri
Forest Reserve, 23 mis S.E. of Kumasi, xi.1959, i <$, i 9 i nr- Kumain, Bobiri
Reserve, vii.1957, i <£ ; Bibianaha, ix-xii.igog, i $ ; E. Region, Kade Agr. Res.
Sta., vii.i963, i ^ ; Tafo, i $, v-vi.i954, 3 <$, iv.igtf, 8 $, 6 $, v.i957, 5 <J, 8 $,
vi.i957, 10 (J, ii $, xi.i96i, 11 ^, 6 $, xii.i96i, 12 ^, i 9, 1.1962, 4 $, 2 $, 11.1962,
ii ^, 3 $, 111.1962, 2 c£, 5 <j>. NIGERIA : Oban Dist., i <$ ; nr. Ibadan, Gombar,

2 The date 23^.1890 also appears on the label of the holotype but is clearly not the date of collection.

REVISION OF CATOPTROPTERYX 163

1.1965, i $. CAMEROON : Efulen, X.IQ22, i <$, xi.i922, 2$, xii.igaz, 3^, i $, 1.1923,
3 <£, 111.1923, i c£, ^.1923, i (J, i $ ; Douala, v.i96i, i ^ ; Mundame, viii.ign, 2 $ ;
Lolodorf (Conradt) i $ (ZMHU Berlin) (holotype of Catoptropteryx maculipennis
Karsch). CONGO (BRAZZAVILLE) : Oka, 1200 ft, 11.1948, i<£ ; Odzala, x.i963, i<$;
Mayumbe, Dimonika, 1.1964, i $ ; Mts. du Chaillu, Mbila, xii.i963, i <^. CONGO
(KINSHASA) : Eskibondo, betw. Nigangara & Dungu, Uele, 2650 ft, x.i934, i ^ ;
Kai Bumba, X.I92O, i <$ ; Region des Lacs, i $ ; 39 km S. of Walikale, 700 m, xii.
1957, i $ ; Ubangi, Budjalibala, 11.1949, I ? '> Eala, iv.i935, i $ ; Janda Sundi,
x.i920, i °- UGANDA : Mpanga Forest, viii.1965, i °-, X.I9&5, i <$ ; i ml E. of
Bundebugyo, viii.1964, i $ ; Bugoma Forest, vi.i933, i $.

Catoptropteryx guttatipes Karsch
(Text-figs, i, 15, 40, 56, 75, 89)

Catoptropteryx guttatipes Karsch, 18906 : 362. Holotype <$, CAMEROON : Barombi-Station
(ZMHU Berlin) [examined].

J. Eyes moderately large, sub-globose, strongly prominent.

Lateral lobes of pronotum as in Text-fig. 40 ; height clearly exceeding length. Rs of fore
wings bifurcate as in Text-fig. 63 ; 7?i bifurcate ; MA unbranched. Stridulatory file as in
Text-fig. 89 ; narrow, moderately curved, teeth widely spaced. Tympanic organs without
auricles.

Cerci as in Text-fig. 75 ; short, arcuate, circular in transverse section, not depressed, strongly
sinuose viewed in plane of principal curvature ; apex obtuse, bearing small spinule.

Pronotal lobes usually with no dark markings ; very rarely with very faint trace of longi-
tudinal stria dorsad. Fore wings in region of stridulatory organ, including area M, black, red-
brown or yellow-brown with black membrane ; basal mark of fore wings of type shown in
Text-fig. 15 : continuous black stripe, about i mm long, or continuous with main dark region
of area M, runs distad from beneath second axillary sclerite into area M, not along centre of
M ; membrane fuscous in cells on either side of MP -\- Cuia. ; no isolated dark cells in areas
R, RI or RS ; apical fuscous spot usually small, sometimes faint or absent. Hind wings hyaline
except for apical archedictyon. External spinules of hind femora usually wholly black and set
in dark spots on femoral carina, less commonly not set in dark spots, rarely blackened only at
tip ; hind femora rarely with one or two fuscous bands in distal half (see below). Hind tibiae
dorsally usually more or less fuscous with darker spot at base of each spinule. Tenth abdominal
tergite and epiproct with variable brown to black markings, former often with pair of black
spots ; cerci blackened dorsally.

°.. As <J except for stridulatory organ and abdominal terminalia. Ovipositor as in Text-fig.
56 ; first gonocoxa large, fairly strongly concave ; apex of dorsal valves smooth. Usually no
dark markings at base of fore wings except short black stripe in area M, as in Text-fig. 15 ;
rarely archedictyon fuscous over whole of cubito-anal area.

Males Females

Number of specimens examined : 32 25

Total length : 38>6-43'7 36-7-44'1

Median length of pronotum : 4-2-4-7 4-1-4-8

Length of hind femur : 18-9-21-9 19-1-21-7

Length of fore wing : 28-6-32-5 28-6-32-4

Stridulatory file —

number examined : 30

number of teeth (T) : 33-82

length (replica) (F) : 1-40-2-11

T/F : 21-6-39-4

164 J. HUXLEY

The ovipositor of this species is extremely similar to that of C. punctulata (Karsch),
differing chiefly in its more strongly concave first gonocoxa and much less slender
valves. Male specimens could be confused with C. occidentalis sp. n. and may
best be distinguished by their shorter, more strongly sinuose cerci. Two male
specimens from Congo (Kinshasa) and one female from Cameroon have two con-
spicuous fuscous bands, one apical, in the distal half of the hind femora. The males
are set apart from the rest of the material examined also by the high number of
teeth (79 and 82) in the stridulatory file. These two males are linked tenuously to
the other 27 examined, among which the highest tooth number recorded was 55,
by one specimen with 63 stridulatory teeth (see Text-fig, i), and with no trace of
dark bands on the hind femora. None of these unusual individuals differs sufficiently
from the other material in any other characters to be considered specifically distinct,
and the variation seems to have no broad geographic significance, so no formal
nomenclatural recognition is indicated.

C. guttatipes Karsch appears to be sparsely distributed in Central Africa from
eastern Nigeria to Uganda, but is also recorded here from two localities in southern
Guinea.

Holotype^. CAMEROON: Barombi-Station (Preuss) (ZMHU Berlin).

GUINEA : W. of Irie, Col de Seredou, vii.i963, i $ ; Nimba, vii-xii.igsi, i $.
NIGERIA : Eastern Prov., 20 mis N.E. of Calabar, Forest Reserve, 1.1961, i $.
CAMEROON : - — , 3^, i $ ; Johann-Albrechtshohe, iv.i896, i^, viii.i896, i 9 ;

Tiko PL, Matute, v.1949, i <$ ; Mundame, i $ ; Kumba, xi.i938, i $, vi.i959, i $ ;
Ja River, Bitye, 2 $, vi-vii.igog, dry season, i $, i $ ; Victoria, 4 $ ; Efulen,
ix.i922, i $, xi.i922, 4 c£, xii.i922, i ^, 4 $, 1.1923, 2 <$, ^.1923, i $. CONGO (BRAZZA-
VILLE) : Sanga R., Nola, 1300 ft, x.1934, i <$ ; Mts. du Chaillu, Mbila, xii.1963, 2 <$,
i $ ; Odzala, x.1963, i <$. CONGO (KINSHASA) : 39 km S. of Walikale, ix.i957,
i (£ ; Ituri Forest, 4000 ft, iv.i93O, i ^ ; Bangala, Loka, xii.i93i, i <£ ; Kivu,
Costermansville, 1951, i $ ', Mayumbe, Makungu, xi.i9i2, i $ ; Tshuapa, Flandria,
xi.i94O, i(£ ; Yambata, ii-iii.i9i4, i $ ; Libenge, i.i-937, i $ ; Stanleyville, vii.i9i2,
i $ ; Binga, iii.i932, i $ ; Kibali-Ituri Dist., Irumu-Avakubi Rd., 10 mis W. Epulu
R. ferry, Saidi, 2800 ft, ix.i934, i $.

Catoptropteryx occidentalis sp. n.

(Text-figs, i, 41, 57, 76, 90)

cj. Eyes moderately large, sub-globose, strongly prominent.

Lateral lobes of pronotum as in Text-fig. 41 ; height clearly exceeding length. Rs of fore
wings bifurcate as in Text-fig. 63 ; RI bifurcate ; MA unbranched. Stridulatory file as in
Text-fig. 90, broad, moderately curved, teeth widely spaced. Tympanic organs without
auricles.

Cerci as in Text-fig. 76 ; long and slender, circular in transverse section, arcuate, somewhat
sinuose viewed in plane of principal curvature ; terminating in small spinule.

Fore wings in region of stridulatory organ, including area M, red-brown to yellow-brown with
black membrane and sometimes black Cu% ; basal mark of fore wings of type shown in Text-
fig. 15 : continuous black stripe, about 1-5 mm long, runs distad from second axillary sclerite

REVISION OF CATOPTROPTERYX 165

into area M, not along centre of M ; membrane fuscous in cells on either side of MP + C«ia ;
no isolated dark cells in areas R, RI or Rs ; fuscous or fuscescent spot, usually well developed,
at apex. Hind wings hyaline except for apical archedictyon. External spinules of hind femora
usually wholly black, sometimes also set in dark patches on femoral carina. Hind tibiae
dorsally more or less fuscescent, with darker area at base of each spinule. Tenth abdominal
tergite, epiproct and cerci red-brown ; cerci blackened dorsally in proximal half.

$. As (£ except for stridulatory organ and abdominal terminalia. Ovipositor as in Text-fig.
57 ; form of basal mechanism highly distinctive : first gonocoxa very large, with ventro-
posterior angle about 90° containing small, deep depression ; second gonocoxa particularly
prominent ; apical margin of dorsal valves smooth. No dark coloration at base of fore wings
except basal stripe in area M.

Males Females

Number of specimens examined : 4 5

Total length : 41-8-43-8 40-9-42-8

Median length of pronotum : 4-8-5-0 4-7-5-0

Length of hind femur : 21-0-21-7 20-0-21-4

Length of fore wing : 31-2-32-7 3°'7-32<3

Stridulatory file —

number examined : 4

number of teeth (T) : 41-46

length (replica) (F) : 1-73-1-98

T/F : 22-2-26-6

C. occidentalis sp. n. is nearest morphologically to C. guttatipes Karsch with which
the males could sometimes be confused ; the cerci, however, are usually much longer
and rather less sinuose than in the latter species. The females are easily dis-
tinguished by the form of the basal mechanism of the ovipositor, which is quite
unlike that of any other species.

This species is known only from Liberia and Ivory Coast.

Holotype $. LIBERIA : N. of Monrovia, Bomi Hills, 5 mis N.E. of mines,
Forest Reserve Rest House, 23^^.1963 (Jago) (BMNH).

Paratypes. IVORY COAST : Foret du Banco, 15. x. 1963, 2 ^, i $ (BMNH), i <$
(ORSTOM Abidjan), 12^.1964, i $ (ORSTOM Abidjan) ; Tai, 21.1.1955, i $
(ORSTOM Abidjan) ; Foret sud, Ndzida, 31^.1952, i ? (ORSTOM Abidjan) ;
— , 6.ii.i96o (Cachan) i $ (ORSTOM Abidjan).

Catoptropteryx serrifera sp. n.

(Text-figs, i, 42, 58, 77, 91, 94-95)

<J. Eyes moderately large, sub-globose, strongly prominent.

Lateral lobes of pronotum as in Text-fig. 42 ; length usually about equal to, or slightly
exceeding height ; rarely length very slightly less than height. Rs of fore wings with 2 or
rarely 3 branches as in Text-fig. 63 ; RI bifurcate ; MA unbranched. Stridulatory file as in
Text-fig. 91, usually almost straight, teeth closely spaced. Tympanic organ without auricles.

Cerci as in Text-fig. 77 ; short, arcuate, circular in transverse section, more or less sinuose
viewed in plane of principal curvature ; terminating in small spinule which is sometimes
unguiform.

No dark markings apart from blackened tips of spurs and spinules. External spinules of hind
femora black or dark brown apically, or wholly so, but with no dark area around their bases.
Hind wings hyaline except for apical archedictyon.

1 66

J. HUXLEY

?. As <$ except for stridulatory organ and abdominal terminalia. Ovipositor as in Text-fig.
58 ; apex of dorsal valve crenulate, bearing about 4-5 very small irregular teeth.

Number of specimens examined

Total length :

Median length of pronotum :

Length of hind femur :

Length of fore wing :

Stridulatory file —

number examined :
number of teeth (T) :
length (replica) (F) :
T/F :

Males

13

40-3-47-3
4-6-5-6

21-2-22-g
30-5-36-2

13

58-77
1-55-2-05

33-2-47-4

Females

16
40-5-45-0

4-7-4-5
20-6-23-5

3I-4-35-4

The form of the lateral lobes of the pronotum, the absence of black markings
from the pronotum and from the base of the fore wings, and the absence of tympanic
auricles, taken in combination distinguish this species from its congeners. The
female is additionally characterized by the distinctive, though somewhat variable,
form of the basal sclerites of the ovipositor, and its toothed dorsal valves.

This species presents a morphological link between the typical species of the
genus and the East African species with its tympanic auricles, its affinity with the
latter species being manifest in the toothed ovipositor, the form of the basal sclerites
and the shape of the pronotal lobes.

When the material from north of the Congo River is compared with that from the
south side, differences in dimensions, and subtle differences in the ovipositors, are
apparent. In the southern group the apical teeth of the ovipositor are bigger, the
appearance of the basal sclerites is somewhat different and the length of the fore
wings relative to that of the hind femora is less (see Text-fig. 95). It would not be

o
o

o 90

32 33

FORE WING

94

95

FIGS. 94-95. Catoptropteryx serrifera sp. n. : 94, map showing the distribution ; 95, scatter
diagram of length of hind femur plotted against length of fore wing. Black disks
represent specimens from south, open circles specimens from north, of the Congo River.

REVISION OF CATOPTROPTERYX 167

unreasonable to suppose that the Congo River might sufficiently restrict gene-flow
between the northern and southern populations for stable subspecific distinctness
of the two groups to be maintained. Carcasson (1964) stated : ' Very often the
ranges of two vicariants will be separated by a partial geographic barrier which
may not have been the cause of the original disjunction, but may help to stabilize
their respective ranges . . . The Congo River and the Rift Valleys of East Africa
are also important dividing lines, though not necessarily effective barriers '. In the
present case, however, I consider there is not sufficient justification for introducing
subspecific names into nomenclature, and any conclusions based on the morphological
discontinuity between the groups must have regard to the small number of specimens
examined.

C. serrifera sp. n. is known only from the lowland forest zone of Central Africa
(Text-fig. 94).

Holotype 9- CAMEROON : Efulen, io.xi.i922 (Weber] (ANS Philadelphia).

Paratypes. CAMEROON ? :— — , 23^.1890 (Morgen) i $ (ZMHU Berlin) (in
alcohol). CAMEROON : Metet, 2i.iii.i942, i <$ (ANS Philadelphia) ; Sangamelima,
Fulasi, V.I92O (Evans) i $ (MZUM Ann Arbor) ; Lolodorf, 21.1.1919 (Reis) i $
(MZUM Ann Arbor) ; Efulen, i6.xi.i922 (Weber] i <$ (ANS Philadelphia), 4-23. xii.
1922 (Weber] 2 <£, i ? (ANS Philadelphia), i $ (BMNH), 15.1.1923 (Weber] i $
(BMNH). CONGO (BRAZZAVILLE) : Chaillu Mts., Mbila, xii. 1963 (Descarpentries
6- Villiers] 2 $, i $ (MNHN Paris), i $ (BMNH) ; Sibiti, xi.i963 (Descarpentries
6- Villiers) i 9 (MNHN Paris), i 9 (BMNH) ; Mayumbe, Dimonika, 1.1964 (Descar-
pentries & Villiers) i $ (MNHN Paris). CONGO (KINSHASA) : Simba, x.i9i2 (Mayne)
i $ (MRAC Tervuren) ; Lac Leopold II, Bolobo, 1955 (Viccars] i <£ (BMNH) ;
Equateur, Flandria, 1929 (Hulstaert] 2 $ (MRAC Tervuren) ; Oshwe, xii. 1913 (Maes]
i 9 (MRAC Tervuren) ; Stanley Pool, 3-io.x.i957 (Brien, Poll & Bouillon] i 9
(MRAC Tervuren) ; N'Kele, Kunungu, 1937 (Schouteden] i 9 (MRAC Tervuren) ;
Maniema, 1936 (Henrard] i 9 (MRAC Tervuren) ; Tolo, xii. 1913 (Maes] i 9 (BMNH) ;
Eala, xi.i934 (Ghesquiere] i 9 (BMNH) ; Equateur, Bokuma, 3O.viii.i934 (Hulstaert]
i 9 (BMNH).

Catoptropteryx aurita sp. n.

(Text-figs, i, 4, 12-13, 43-45. 59. 78> 92- Plate i, fig. 3)

c£. Eyes moderately large, sub-globose, strongly prominent.

Lateral lobes of pronotum as in Text-figs. 43-45 ; approximately as high as long. Ifs of
fore wings as in Text-fig. 63, bifurcate, or rarely with 3 or 4 dichotomous branches ; R± with 2
or 3 branches ; MA unbranched. Stridulatory file as in Text-fig. 92 ; narrow, moderately
curved, teeth widely spaced. Tympanic organs as in Text-figs. 12-13 '• internal side with
auricle more or less well developed.

Cerci as in Text-fig. 78 ; short, circular in transverse section, arcuate, somewhat sinuose
viewed in plane of principal curvature ; terminating acutely in small spinule. Styles com-
paratively large, with clear articulation with subgenital plate.

Lateral lobes of pronotum without dark marking or with partially developed black stria
dorsad, as in Text-figs. 43-45. Fore wings without dark markings at base ; large fuscescent
spot at apex ; membrane dark in cells adjacent to MP + Cwia, but no isolated dark cells in

168 J. HUXLEY

areas R, R\ or R^. Hind wings hyaline except for apical archedictyon. Spinules of hind femora
apically or wholly black.

$. As (J except for stridulatory organ and abdominal terminalia. Ovipositor as in Text-fig.
59 ; apex of dorsal valves serrate, bearing about 5-7 small teeth.

Males Females

Number of specimens examined : 10 10

Total length : 41-3-54-3 40-5-52-2

Median length of pronotum : 5-5-6-2 5 -2-5 -9

Length of hind femur : 21-4-25-5 20-7-26-0

Length of fore wing : 29-8-40-1 31-1-45-7

Stridulatory file —

number examined : 10

number of teeth (T) : 4!-53

length (replica) (F) : 1-68-2-21

T/F : 23-5-26-5

C. aurita sp. n. is very clearly distinct from the rest of the genus, having a rudimen-
tary to quite well developed auricle on the internal side of the tympanic organ.
The distribution has no intersection with that of any other species. C. serrifera
sp. n. provides a geographical and morphological bridge from this to the typical
species, having the unusual form of the pronotal lobe in common with C. aurita
sp. n., an ovipositor in many respects intermediate, and a tympanic organ of the
typical form. In C. aurita sp. n. the teeth of the ovipositor are larger and more
regular in shape than in C. serrifera sp. n., and the form of the basal mechanism,
though similar, can be distinguished easily enough ; however, the best distinguishing
feature of specimens with poorly developed tympanic auricles is the number of
teeth in the stridulatory file. The variations in coloration, size, and form of ovi-
positor and tympanic organ have no apparent geographic correlations.

C. aurita sp. n. is known only from East Africa, ranging from southern Kenya
to southern Rhodesia and Mozambique (Text-fig. 4).

Holotype $. RHODESIA : Lundi, 3-5.^.1964 (van Son & Vdri) (TM Pretoria).

Paratypes. KENYA : Mombasa, iv.igss, i $ (BMNH). TANZANIA : E. Usam-
bara Mts., Amani, 1950 (Verdcourt) I $ (CM Nairobi) ; Zanzibar, Mtoni, v.1954
(Brown) i <J (BMNH). ZAMBIA : Abercorn, i8-22.xi.i963 (Vesey-Fitzgerald) i <J
(BMNH);- —(Trenewith) i 9 (TM Pretoria). RHODESIA : Lundi, 3-5.^.1964

(van Son 6- Vdri) i <?, i $ (BMNH), 3 <J (TM Pretoria) ; Vumba, 2-9.xi.ig59 (van
Son) i<? (BMNH) ; S. Melsetter, Vimba, X.IQSS, i $ (NM Bulawayo). MOZAMBIQUE :
Kruger National Park, Pafuri, 24~28.iv.i96i (Rorke) i $ (TM Pretoria) ; Chiluvo
Hills, X.I9&3 ,i c? (BMNH), 3.xi.i963, 2 $, 2 $ (NM Bulawayo) ; Savanie Forest,
x. 1963, i $ (NM Bulawayo).

Catoptropteryx extensipes Karsch
(Text-figs, i, 5-6, 8-9, 46-47, 65, 79-81, 93)

Catoptropteryx extensipes Karsch, 1896 : 334. Holotype <$, CAMEROON : Lolodorf (ZMHU

Berlin) [examined].

cJ. Eyes as in Text-figs. 5-6, 8-9 ; very large, elliptic, moderately, rarely strongly,
prominent.

REVISION OF CATOPTROPTERYX 169

Lateral lobes of pronotum as in Text-figs. 46-47 ; height slightly greater than length. Fore
wings as in Text-fig. 65 ; appreciably narrowed in distal half ; Rs bifurcate ; RI with 2 to 4
branches ; MA unbranched. Stridulatory file as in Text-fig. 93 ; broad, moderately curved,
teeth widely spaced. Tympanic organs without auricles.

Cerci as in Text-figs. 79-81 ; strongly arcuate to U-shaped, near apex abruptly narrowed and
bent downward slightly ; apex acute.

General coloration green. Frons, labrum and clypeus suffused with brown of variable
intensity. Pronotal lobes unmarked, or with longitudinal brown to black stria dorsad as in
Text-fig. 47, not extending as far as anterior margin. Basal mark of fore wings similar to type
shown in Text-fig. 16 but very reduced in length and width, about 0-5 mm long ; cells in
posterior half of wing fuscescent but not distributed in isolated groups in areas R, RI or #5.
Hind wings somewhat fumose, darker toward margin of anal fan. Herring-bone pattern of
hind femora picked out in dark brown. Hind tibiae deep brown, paler or more reddish dorsally,
with 2 broad, well defined, very pale yellow or green bands in proximal third. Tarsi and apex
of tibiae almost black, with some small paler markings. Abdominal tergites with fuscous spot
dorsally, tenth with broad fuscous or black patch ; epiproct and paraprocts dorsally with
variable brown to black markings ; cerci wholly black.

$. Not known.

Males

Number of specimens examined : 1 1

Total length : 44-6-50-8

Median length of pronotum : 5'3-6-3

Length of hind femur : 25-7-30-0

Length of fore wing : 31 -9-36-9
Stridulatory file —

number examined : 1 1

number of teeth (T) : 49~79

length (replica) (F) : 2-07-2-71

T/F : 20-5-35-3

This species differs from the rest of the genus in so many characters, the most
striking of which are the form of the eyes, cercus and fore wings, that one may
suspect that the discovery of the female could make the erection of a new genus
desirable for it. As it is, without the evidence of the female, and more particularly
of the structure of the ovipositor, the species rests not too uncomfortably in Catoptro-
pteryx Karsch, and it is most conveniently left here for the time being. There is
some geographic variation in the pronotal coloration and the form of the cerci and
eyes. At the western and eastern edges of the range the pronotal stria is absent,
and in specimens from Cameroon and Fernando Poo the eyes are narrower (Text-
figs. 5 and 8) and the cerci less strongly curved (Text-fig. 79).

Catoptropteryx extensipes Karsch has a West African distribution, ranging from
Sierra Leone to Cameroon and Fernando Poo.

Holotype <$. CAMEROON : Lolodorf (Conradt) (ZMHU Berlin).

GUINEA : Nimba, Ziela, iii.1957, 3 <$. SIERRA LEONE : Freetown, Mt. Aureol,
vi.i956, i c£. LIBERIA : N. of Monrovia, Bomi Hills, 5 mis N.E. of mines, Forest
Reserve Rest House, vii.i963, i $. IVORY COAST : Foret du Banco, x.1963, i $.
GHANA : W. Region, near Wiawso, 3 mis N.W. of Tano Lodge, x.i96o, i <$.
CAMEROON : Efulen, xi.i92o, i $, xii.i922, i <$. FERNANDO Poo : 1901, i <$.

170 J. HUXLEY

REFERENCES

BOLIVAR, I. 1893. Voyage de M. Ch. Alluaud dans le territoire d'Assinie (Afrique occidentale)

en juillet et aout 1886. i4e Memoire. Orthopteres. Annls Soc. ent. Fr. 62 : 169-184, i pi.
BRUNNER VON WATTENWYL, C. 1891. Additamenta zur Monographic der Phaneropteriden.

Verh. zool.-bot. Ges. Wien 41 : 1-196, 2 pis.
CARCASSON, R. H. 1964. A preliminary survey of the zoogeography of African butterflies.

E. Afr. Wildl. J. 2 : 122-157, 6 figs-
CHOPARD, L. 1955. In HANSTROM, B., BRINCK, P. & RUDEBECK, G. [Edd.], 5. Afr. Anim.

Life 2 : 266-300 (Orthoptera Ensifera), 26 figs. Stockholm.
KARSCH, F. 1889. Orthopterologische Beitrage III. Berl. ent. Z. 32 : 415-464, i pi.

- 18900. Neue westafrikanische durch Herrn Premierlieu tenant Morgen von Kribi ein-
gesendete Orthopteren. Ent. Nachr. 16 : 257-276, 4 figs.

- 18906. Verzeichniss der von Herrn Dr. Paul Preuss auf der Barombi-Station in Deutsch-
Westafrika, 1890, gesammelten Locustodeen aus den Familien der Phaneropteriden,
Mekonemiden und Gryllakriden. Ent. Nachr. 16 : 353-369, 4 figs.

- 1896. Neue Orthopteren aus dem tropischen Afrika. Stettin, ent. Ztg 57 : 242-359, 44
figs.

KIRBY, W. F. 1906. A synonymic catalogue of Orthoptera 2, Pt. i, viii + 562 pp. London.
RAGGE, D. R. 1955. The wing-venation of the Orthoptera Saltatoria 159 pp., 106 figs. London.

- 1964. A revision of the genus Tylopsis Fieber (Orthoptera : Tettigoniidae) . Bull. Br.
Mus. nat. Hist. (Ent.) 15 : 297-322, 52 figs.

- 1969. A revision of the African species of Pseudorhynchus Serville (Orthoptera : Tetti-
goniidae). Bull. Br. Mus. nat. Hist. (Ent.) 23 : 167-190, 38 figs.

SCUDDER, G. G. E. i96ia. The comparative morphology of the insect ovipositor. Trans. R.
ent. Soc. Lond. 113 : 25-40, 10 figs.

- 19616. The functional morphology and interpretation of the insect ovipositor. Can.
Ent. 93 : 267-272, 5 figs.

- 1964. Further problems in the interpretation and homology of the insect ovipositor.
Can. Ent. 96 : 405-417, n figs.

WALKER, F. 1869. Catalogue of the specimens of Dermaptera Saltatoria in the collection of the
British Museum 2 : 225-423. London.

JOHN HUXLEY

Department of Entomology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON, S.W-7

PLATE

PLATE i

FIG. i. Eggs of Catoptropteryx sp. on abaxial surface of Ficus leaf.
FIG. 2. C. apicalis (Bolivar), $.
FIG. 3. C. aurita sp. n., <$.

Bull. BY. Mus. nat. Hist. (Ent.) 24, 5

PLATE i

A LIST OF SUPPLEMENTS
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OF THE BULLETIN OF
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2. NIXON, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera :
Braconidae). Pp. 284 : 348 text-figures. August, 1965. £6.

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

4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera,
Termitidae) from the Ethiopian Region. Pp. 172 : 500 text-figures. Sep-
tember, 1965. £3 55.

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

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

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

£33s.

8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the

world species of Cleora (Lepidoptera : Geometridae). Pp. 119 : 14 plates, 146
text-figures, 9 maps. February, 1967. £3 los.

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

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

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

12. WATSON, A. The Taxonomy of the Drepaninae represented in China, with
an account of their world distribution. Pp. 151 : 14 plates, 293 text-figures.
November, 1968. £5.

13. AFIFI, S. A. Morphology and Taxonomy of Adult Males of the families
Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210 : 52 text-
figures. December, 1968. £5.

14. CROSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa
and its Islands. Pp. 198 : i plate, 331 text-figures. July, 1969. £4 155.

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

£4-

16. GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe
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£19-

Printed in England by Staples Printers Limited at their Kettering, Northants, establishment

A LIST OF THE TYPE-SPECIMENS
OF ODONATA IN THE BRITISH
MUSEUM (NATURAL HISTORY)

PART III

D. E. KIMMINS

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 24 No. 6

LONDON: 1970

A LIST OF THE TYPE-SPECIMENS

OF ODONATA IN THE BRITISH

MUSEUM (NATURAL HISTORY)

PART III

BY

DOUGLAS ERIC KIMMINS

•

Jt

Pp. 171-205

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 24 No. 6

LONDON: 1970

THE BULLETIN OF THE BRITISH MUSEUM

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

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

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

This paper is Vol. 24 No. 6 of the Entomological
series. The abbreviated titles of periodicals cited
follow those of the World List of Scientific Periodicals.

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

Trustees of the British Museum (Natural History), 1970

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 30 January, 1970 Price £i

A LIST OF THE TYPE-SPECIMENS OF

ODONATA IN THE BRITISH MUSEUM

(NATURAL HISTORY) PART III

Families Platystictidae, Protoneuridae, Platycnemididae, Coenagriidae,
Pseudostigmatidae, Perilestidae, Synlestidae, Lestidae, Megapodagriidae,
Pseudolestidae & Polythoridae.

By D. E. KIMMINS

CONTENTS

Page

ADDENDA TO PART I ......... 173

PLATYSTICTIDAE . . . . . . . . . .174

PROTONEURIDAE .......... 175

PLATYCNEMIDIDAE .......... 179

COENAGRIIDAE .......... 181

PSEUDOSTIGMATIDAE ......... 193

SYNLESTIDAE .......... 193

PERILESTIDAE ........... 193

LESTIDAE ........... 194

MEGAPODAGRIIDAE .......... 196

PSEUDOLESTIDAE .......... 198

POLYTHORIDAE .......... 199

REFERENCES ........... 200

SYNOPSIS

A list of the type-specimens of Odonata in the British Museum (Natural History), belonging
to the eleven above-mentioned families has been prepared. Four hundred and two taxa are
dealt with and lectotypes are designated for one hundred and thirteen of these taxa.

This list completes those published in 1968, Bull. Brit. Mus. nat. Hist. (Ent.) 22
(7) : 277-305, and 1969, op. cit. 23 (7) : 287-314. The present list was completed
in May 1969.

ADDENDA TO PART I

peruviana Cowley (Aeschnosoma), 1934 : 93~94> ngs I-2- Holotype °-. 4206. Aeschno)
soma peruviana Cowl. °-. Peru, Prov. Moyombamba, Dep. S. Martin, Rioja (P. Martin
HOLOTYPE. 1934, Stylops, 3 : 93, f. i, 2. det. Cowley, coll. Cowley.

serva Kirby (Trithetnis), 1900 : 69-70, text-fig, i. LECTOTYPE $. Sierra Leone, Free
Town, 5.ix.[i8]99 (E. E. Austen), with on reverse, 'T. serva K' / Trithemis serva Kirby, 5*
Lectotype, D. E. Kimmins det. 1967.

The lectotype lacks segments 7-10, and appears to be the example figured by Kirby.

174 D- E- KIMMINS

tricolor Kirby (Orthetrum), 1894 : 555-556. Holotype <$. Kandy, 3o.vi.[i8]92 / Ceylon,
Yerbury Coll. / Orthetrum tricolor [WFK's writing].

This species was accidentally omitted from Kimmins (1968), probably in the course of
typing the manuscript.

CORRECTION TO PART II
p. 307. multinervosa Eraser should be (Devadatta), not (Diphlebia).

PLATYSTICTIDAE

actaeon Laidlaw (Drepanosticta), 1934 : 558-559, fig. 3. Holotype $. Brit. N. Borneo,
Mt Kinabalu, Kabayau, 600 ft, 8. v. 1929 (H. M. Pendlebury) / Drepanosticta actaeon sp. n.
Holotype, Type [Laidlaw's writing] / Drepanosticta actaeon Laidl.
One inferior appendage detached and glued to card.

anascephala Eraser (Drepanosticta), 1933. Kimmins, 1966 : 177.
antelopoides Eraser (Protosticta), 1931. Kimmins, 1966 : 178.

apicalis Kirby (Platysticta), 1894 : 561-562, pi. 42, fig. i. LECTOTYPE 6*- [Ceylon],
Belihul-Oya, 6.vi.[i8]92 /apicalis, Kb. type / Platysticta apicalis Kirby, $ Lectotype, D. E.
Kimmins det. 1968.

davenporti Eraser (Protosticta), 1930. Kimmins, 1966 : 188.

dentifera Kimmins (Drepanosticta), 1936 : 101-103, fig- I7- Holotype <$. At Waterfall /
Sarawak, Mt. Dulit, 3300 ft, Primitive forest / Drepanosticta dentifera sp. n. $, det. D. E.
Kimmins.

dulitensis Kimmins (Drepanosticta), 1936 : 98, fig. 15. Holotype $. Sarawak, Mt Dulit,
3700 ft, 20. x. 1932, on rocks below top crossing / Drepanosticta dulitensis <$, sp. n., det. D. E.
Kimmins.

Jorficula Kimmins (Drepanosticta), 1936 : 100-101, fig. 16. Holotype $. Sarawak, Mt
Dulit, 4000 ft, moss forest, i6.x.i932 / Drepanosticta forficula sp. n. <$, det. D. E. Kimmins.

gracilis Kirby (Protosticta), 1889 : 302. Holotype <?. [Celebes], Tondano / Menado (Wallace)
I Protosticta gracilis type [WFK's writing].

kinabaluensis Laidlaw (Protosticta), 1915^ : 37-38, text-fig. 56. Holotype <J. Borneo
(/. C. Moultori) I Protosticta kinabaluensis $ Laidlaw Type, Kina Balu, Borneo (/. C. Moulton),
5-ix.i9i3 [Laidlaw's writing].

greeni Kirby (Platysticta), 1891 : 204-205, pi. 20, figs 3, 3a. Holotype $. Ceylon / Ceylon,
Pundaloya, viii.iSSg (E. E. Green) / greeni (Type) [WFK's writing] / Platysticta greeni
Kirby, $ Holotype.

Currently placed as a synonym of Platysticta maculata Selys.

hamadryas Laidlaw (Drepanosticta), 1931 : 187, fig. 2. Holotype <J. D. hamadryas
Kuala Tahan, Pahang, F.M.S., 300 ft, 2o.xi.[i9]2i (H. Pendlebury) [in Fraser's writing].
This specimen came to the BMNH with the Fraser Bequest.

hearseyi Fraser (Protosticta), 1922. Kimmins, 1966 : 195.
lankanensis Fraser (Ceylonosticta), 1931. Kimmins, 1966 : 200.
lindgreni Fraser (Protosticta), 1920. Kimmins, 1966 : 201.

lymetta Cowley (Drepanosticta), 1936 : 161-163, figs 13-14. Holotype $. Surigao, Min-
danao, Baker j Philippine Is. (C. F. Baker) / Drepanosticta sp. <$, H. Campion det. / Drepanos-
ticta lymetta sp. n. HOLOTYPE, prothorax drawn August, det. J. Cowley, 1935.

marsyas Lieftinck (Drepanosticta), 1965 : 178-180, figs 3-7. Holotype £. Pahang,
Cameron's Highlands, 4-5000 ft, i5-iv.[i935] (H. M. Pendlebury) / Drepanosticta marsyas $
Type [Laidlaw's writing] / Drepanosticta marsyas Lieft., HOLOTYPE, M. A. Lieftinck det. 1964.

TYPE-SPECIMENS OF ODONATA IN BMNH 175

tnegametta Cowley (Drepanosticta), 1936 : 163-167, figs 15-22. Holotype Q*. Surigao,
Mindanao (Baker) / Drepanosticta megametta sp. nov., £ HOLOTYPE, prothorax, thoracic
pattern and anal apps drawn, det. J. Cowley, August 1935.

tnortoni Fraser (Protosticta), 1924. Kimmins, 1966 : 205.

mylitta Cowley (Drepanosticta), 1936 : 160-161, figs 1-12. Holotype 6*- Drepanosticta
mylitta sp. n. $ HOLOTYPE. Borongan, Samar, Philippine Is. (C. F. Baker) det. J. Cowley,
1935 / details drawn, J. Cowley, 1935.

nietneri Fraser (Ceylonosticta), 1931. Kimmins, 1966 : 206.

pan Laidlaw (Drepanosticta), 1931 : 185-187, fig. i. Lectotype Q* (Lieftinck, 1965 : 174).
Perak, Batang Padang, Jor Camp, June 1923 (F. N. Chasen) / Drepanosticta pan <$ Type
[Laidlaw's writing] / Lectotype selected by Lieftinck, 1964.
The apical abdominal segments are detached and glued to card.

paulina Drury (Libellula), 1773 : pi. 40, fig. 4.

Calvert (1932 : 4) was unable to trace the type of this species. McLachlan's collection,
however, contained a single example, lacking head and abdomen, without locality label but
bearing one of his type labels and labelled by him 'Palaemnema paulina Drury'. The apical
brown wing markings are rather more extensive than in Drury's figure, extending basally
by about the length of the pterostigma. If it is Drury's specimen, I suspect that it has been
re-pinned. I have no information as to how this example came into McLachlan's possession,
but see Kimmins (1969 : 306) under titia. Selys (1886 : 146) states 'Patrie: Mexique.
Honduras d'apres Drury; coll. Selys, Mac-Lachlan.' There is no indication that McLachlan's
specimen was Drury's type, and I am of the opinion that it may have been labelled Type by
McLachlan in error. I am retaining it in our type-collection in case information should come
to light as to its history.

paulitoyaca Calvert (Palaemnema), 1931 : 47-49, pi. 5, figs 49, A-G", pi. 18, figs 74-76.
Holotype <$. Atoyac, Vera Cruz, May (H. H. S[mith]). / Palaemnema paulina Drury, P. P.
Calvert det. 1903, B. C. A. Neur. p. 136, Original of pi. 5, fig. 49 / Palaemnema paulitoyaca
Calv. TYPE o\ Original of figs 74, 76, pi. 18, Tr. Am. Ent. Soc., LVII, P. P. Calvert det. 1931.

rufostigma Kimmins (Protosticta), 1958 : 349, fig. i. Holotype <$. S. India, Tinnevelly Dt.,
Naraikadu, 2500-3000 ft, 3-8. x. 1938 (G. M. Henry). Protosticta rufostigma Kimmins, $
TYPE, D. E. Kimmins det. 1947.

It may be pointed out that the date of capture is incorrectly given in the original description
as September.

sanguinostigma Fraser (Protosticta), 1922. Kimmins, 1966 : 212.

silenus Laidlaw (Drepanosticta), 1934 : 557~55^, fig- 2. Holotype <£. Perak, Larut Hills,
4500 ft, 20. ii. 1932 [no collector's name] / Drepanosticta silenus <$ Laidlaw [labels D.E.K.].

Lieftinck (1965 : 176) designates the above example as lectotype, but this is not necessary
as Laidlaw states that the BMNH Q* is the holotype. Lieftinck also states that the $ allotype
was not recovered. This I do not understand, as the allotype is in the Type Collection with
the Holotype Q*.

stevensi Fraser (Protosticta), 1922. Kimmins, 1966 : 214.

versicolor Laidlaw (Protosticta), 1913 : 78. Holotype $. Sarawak, Lawas, I5.ix.[i9]og
(J. C. Moulton) I Protosticta versicolor Laidlaw, $ Type.

viridis Fraser (Drepanosticta), 1922. Kimmins, 1966 : 218.
walli Fraser (Ceylonosticta), 1931. Kimmins, 1966 : 218.
wheeleri Fraser (Drepanosticta), 1942. Kimmins, 1966 : 218.

PROTONEURIDAE
acuta Kimmins (Elattoneura), 1938 : 300, figs 6, A-D. Holotype $. Nigeria, Lagos (G.

176 D. E. KIMMINS

Strachari) / Elattoneura acuta $ Kimm. Type, det. D. E. Kimmins, 1968.
Apex of abdomen, penis cleared and mounted in Canada balsam.

amelia Calvert (Neoneura), 1903 : 138, pi. 5, fig. 36, pi. 6, fig. 8. LECTOTYPE <J.
[Guatemala], Vera Paz, Cubilguitz ([G. C.] Champion) / Neoneura amelia Calvert, P. P.
Calvert det. 1903, B.C. A. Neur., p. 138, Orig. of part of pi. 6, fig. 8 / Neoneura amelia Calvert,
<J Lectotype, D. E. Kimmins det. 1968.

Two examples were the basis of the figures of this species. As the second example is now
fragmentary, I have selected the above example (which has carried a Museum type-label for
many years) as the Lectotype.

annandalei Eraser (Caconeura), 1921. Kimmins, 1966 : 178.
apicalis Eraser (Chloroneura), 1924. Kimmins, 1966 : 178.

atrocyana Lieftinck (Notoneura), 1960 : 117, figs 1-5. Holotype <$. N. Dutch New Guinea,

Waigeu, Camp Nok, 2500 ft, iv.i938 (L. E. Cheesman) / Notoneura atrocyana Lieftinck, <$

Holotype, det. D. E. Kimmins, 1968.

The original determination labels on the paper envelopes were accidentally destroyed.
auricolor Eraser (ssp. Caconeura dorsalis), 1927. Kimmins, 1966 : 181.
autumnalis Eraser (Caconeura), 1922. Kimmins, 1966 : 181.
balli Kimmins (Elattoneura), 1938 : 296, figs. 3, A-E. Holotype $. Sierra Leone, Bafodea,

13. iv. 1912 (Jas J. Simpson) / Disparoneura pruinosa Selys, <$, Determined by H. Campion /

Elattoneura balli Kimm. <$ Type, det. D. E. Kimmins.
banksi Tillyard (Isosticta), 1913 : 433-434, pi. 47, fig. 4. LECTOTYPE 6*- N[orth]

Queensland], Banks Is., I2.ii.[i9]io (H. Elgner) / Isosticta Banksii Till., <$ Type, R.J.T. /

Isosticta banksi Till., $ Lectotype, D. E. Kimmins det., 1968.

The spelling of the name 'Banksii' was corrected to 'banksi' in the original publication.
bilineata Eraser (Melanoneura), 1922. Kimmins, 1966 : 182.
botti Eraser (Caconeura), 1922. Kimmins, 1966 : 183.

burmanensis Eraser (ssp. of Caconeura verticalis), 1933. Kimmins, 1966 : 184.
cacharensis Eraser (ssp. of Disparoneura campioni), 1933. Kimmins, 1966 : 184.
campioni Eraser (Disparoneura), 1922. Kimmins, 1966 : 185.
canescens Tillyard (Neosticta), 1913 : 435-438, pi. 47, fig. 6. LECTOTYPE <?. N[ew]

S[outh] W[ales], Illawarra, x.[i9]o7 (R. J. Tillyard) / Neosticta canescens Tillyard, 6* TYPE /

Neosticta canescens Till., ^ Lectotype, D. E. Kimmins, 1968.
canningi Eraser (Caconeura), 1919. Kimmins, 1966 : 185.
[cara Calvert (Protoneura), 1903 : 143, pi. 5, figs 38, 45, pi. 6, fig. 9.

The specimen in the BMNH from Guerrero, La Venta is fragmentary, and although it

was the original of part of pi. 6, fig. 9, in view of the fact that there were two other males in

the Calvert collection, I am assuming that the holotype is one of these and have marked our

fragmentary $ and an incomplete $ from Dos Arroyos as paratypes.]
coelestina Tillyard (Alloneura), 1906 : 184-186, pi. 17, figs 5a, b. LECTOTYPE <J. N.

Queensland], Cairns, [Redlynch], i.[i9]o5 (R. J. Tillyard) / Alloneura coelestina Till., <J TYPE,

R.J.T. / Alloneura coelestina Till., $ Lectotype, D. E. Kimmins det. 1968.

Currently placed in the genus Notoneura.

cyaneovittata Eraser (Esme), 1922. Kimmins, 1966 : 187.
[dorrigoensis Tillyard (race of Neosticta canescens), 1913 : 437-438. Type not in the

collection of Odonata bequeathed to BMNH by Tillyard but is in the Australian National

Collection at Canberra (C.S.I.R.O.).]
dorsalis Kimmins (Elattoneura), 1913 : 299, text-figs 5, A-E. Holotype <J. Sierra Leone,

Yana, 3o.iii.i9i2 (Jas J. Simpson) / Elattoneura dorsalis Kimmins, <J Type, det. D. E.

Kimmins.

TYPE-SPECIMENS OF ODONATA IN BMNH 177

evelynae Lieftinck (Notoneura), 1960 : 120, fig. 6. Holotype <$. N. Dutch New Guinea,
Waigeu [Isl.], Camp Nok [Mt Buffelhoorn], 2500 ft, iv.iQ38 (L. E. Cheesman) / lat. stripe
pale blue, under thorax and abd. apex light brown [collector's note] / Notoneura evelynae
Lieftinck, $ Holotype, det. D. E. Kimmins, 1968 [original determination on paper envelope
accidentally destroyed].

fieldi Tillyard (Austrosticta), 1908 : 765-766, pi. 42, figs 6-9. LECTOTYPE 6*. [Australia],
N[orthern] Territory], Tennant's Creek, iv.[i9]o6 (D. Field] / Austrosticta Fieldi Till.,
o* TYPE, R.J.T. / Austrosticta fieldi Till., Q* Lectotype, D. E. Kimmins det. 1968.

filicicola Tillyard (Oristicta), 1913 : 438-440, pi. 440, fig. 7. LECTOTYPE <J. N[orth]
Queensland], Cooktown, i.[i9]o8 (R. J. Tillyard) / Oristicta filicicola Till., 6* TYPE, R.J.T. /
Oristicta filicicola Till., £ Lectotype, D. E. Kimmins det. 1968.

fletcheri Fraser (Disparoneura), 1919. Kimmins, 1966 : 192.

hyperythra Selys (Alloneura), 1886 : 180-181. Holotype <$. Type [McL. label] / Labuan /
Alloneura hyperythra Selys, <$ Borneo.

Currently placed in the genus Prodasineura. The type now lacks the right anterior wing.

klugi Cowley (Protoneura), 1941 : 145-148, figs 1-24. Holotype o*. Peru, [Dept. Loreto],
Iquitos, v.1938 (G. C. Klug) / Protoneura klugi Cowley, Q* Holotype, det. J. Cowley, 1940.
Holotype (with head detached) mounted on card.

longistyla Fraser (Esme), 1931. Kimmins, 1966 : 202.
tnackwoodi Fraser (Caconeura), 1919. Kimmins, 1966 : 202.

marshalli Ris (Chlorocnemis), 1921 : 291, pi. 8, fig. 8, text-fig. 20. Lectotype <$ (Pinhey, 1962 :

104). Mashonaland, Mazoe, 4000 ft, 24.11.1905 (G. A. K. Marshall) / Chlorocnemis nov. spec.

Det. Dr F. Ris /Chlorocnemis marshalli Ris, Type, described 1921, det. Miss C. Longfield.

Pinhey, 1962, in effect designated the BMNH example as lectotype. The specimen now

has only two wings, one pair possibly retained by Ris.

moultoni Laidlaw (Disparoneura), 1912 : 98. Holotype <$. Sarawak, n.v.ign (/. C.
Moulton) I Disparoneura nov. spec. [Ris' writing] / Disparoneura moultoni Laidlaw, Type Q*.
Currently placed in the genus Prodasineura, as a synonym of P. hyperythra (Selys).

mudiensis Fraser (Esme), 1931. Kimmins, 1966 : 205.

mutata Selys (Disparoneura), 1886 : 164-165. LECTOTYPE <J. Type [McL. label] /
Magila / Disparoneura mutata ^ Selys, Magila / Disparoneura mutata Selys, $ Lectotype,
D. E. Kimmins det. 1968.

Currently placed as a synonym of Elattoneura glauca (Selys) .

nigra Fraser (Caconeura), 1922. Kimmins, 1966 : 206.

nigra Kimmins (Elattoneura), 1938 : 297, figs 4, A-E. Holotype £. S. Nigeria, Ibadan,
n.iii.[i9]i9 / Elattoneura nigra Kimm. <$ Type, det. D. E. Kimmins. /
Penis mounted in Canada balsam.

nigripes Selys (Chlorocnemis), 1886 : 141. Lectotype $ (Gambles, 1967 : 197).

oculata Kirby (Disparoneura), 1894 : 562-563. Holotype 9- [Ceylon], Kottawa, 19. iv.
[i8]92 [Yerbury] / oculata type [WFK].

Currently placed as a synonym of Elattoneura centralis Hagen.

pauli Longfield (Chlorocnemis), 1936 : 470-471, fig. 2. Holotype <$. Central Africa,
Uganda, Toro Distr., Kibale Forest, 2.iii.i934, By river (C. E. Longfield) / Chlorocnemis
pauli sp. nov. Type $, det. Miss Longfield.

peramans Calvert (Protoneura), 1902 : 231; 1903 : 141-142, pi. 5, figs 48, 49. Holotype £.
Guatemala, Vera Paz, Panima ([G. C.] Champion) / Protoneura peramans Calv., P. P. Calvert
det. 1903, B.C. A. Neur. p. 141, Orig. of Tab. 5, fig. 48.

No type was indicated by Calvert in 1902 in his brief description in society proceedings.

178 D. E. KIMMINS

A pair in coitu only is referred to and strictly one of these should have been chosen as type.
Calvert (1903 : xxviii) states that the types of his new species described in B.C. A. Neur.
are the specimens marked as originals of figures, and as the above-mentioned example has
been considered as type for many years, I have accepted Calvert as having designated this
specimen as type.

peramoena Laidlaw (Disparoneura), 1913 : 76-77. LECTOTYPE <$. Sarawak, Lawas,
15. ix. 1909 (/. C. Moultori) I Disparoneura peramoena Laidlaw, Type <$. / Disparoneura
peramoena Laidl., <$ Lectotype, D. E. Kimmins det. 1968.
Currently placed in the genus Prodasineura.

peruviensis Fraser (Protoneura), 1946. Kimmins, 1966 : 209.
protostictoid.es Fraser (Protoneura), 1946. Kimmins, 1966 : 210.
ramburi Fraser (Indoneura), 1922. Kimmins, 1966 : 211.

remissa Calvert (Protoneura), 1903 : 144-145, pi. 5, fig. 43. Holotype $ (Calvert, 1903 :
xxviii). [Mexico], Tabasco, Teapa, ii (H. H. Smith}} / Protoneura remissa Calv., P. P.
Calvert det. 1903, B. C. A. Neur. p. 144. Orig. of Tab. V, fig. 43.
Currently placed in the genus Psaironeura.

salomonis Selys (Alloneura), 1886 : 188, 189. Holotype <J. Type [McL. label] / Solomon
Ids (Mathew) / Alloneura salomonis Selys <$, I. Salomon.
Currently placed in the genus Notoneura.

sita Kirby (Disparoneura), 1894 : 563. LECTOTYPE $ (teneral, lacking head). [Ceylon],
Hot Wells, Trincomali, I3.ix.[i8]9i ([Yerbury]} / sita, type [WFK] / Disparoneura sita
Kirby, $ Lectotype, D. E. Kimmins det. 1968.

Fraser (1923 : 223) refers to the adult type-specimen in the BMNH and a teneral co-type
male. The latter specimen is the one fully described by Kirby and the adult is only very
briefly referred to. The teneral specimen bears Kirby's determination label and has been
chosen as the type. Currently placed in the genus Prodasineura.

solitaris Tillyard (Alloneura), 1906 : 182-184, pi. xvii, figs 4a, b. LECTOTYPE <$. N.
Queensland], Kuranda, i.[i9]o5 (R. J. Tillyard) / Alloneura solitaria Till., <$ TYPE, R.J.T. /
Alloneura solitaris Till., $ Lectotype, D. E. Kimmins det. 1968.

Currently placed in the genus Notoneura. The specific name is given four times as solitaris
in the text but on the type it is spelled solitaria.

souteri Fraser (Disparoneura), 1924. Kimmins, 1966 : 213.

subnodalis Selys (Disparoneura), 1886 : 162, 163. LECTOTYPE $. Type [McL. label) /
Old Calabar (R\utherford~\) 86 / Disparoneura subnodalis Selys <£ Old Calabar / Disparoneura
subnodalis Selys, <$ Lectotype, D. E. Kimmins det. 1968.

Pinhey, 1962 : 107 quotes <J $ cotypes in BMNH and i damaged $ in Brussels, type not
designated.

theebawi Fraser (Caconeura), 1922. Kimmins, 1966 : 216.

tillyardi Campion (Isosticta), 1921 : 38-41, figs i, 3. Holotype^. Mt. Canala, I3.vi.[i9]i4,
Coll. P. D. Montague, New Caledonia Exped. / Isosticta tillyardi Campion, $ Holotype.
Determined by H. Campion.

vittata Selys (Alloneura sg. Disparoneura), 1886 : 174-175. LECTOTYPE <$. Type
[McL. label] / Camaroons (R\utherford]) , 2 / Disparoneura vittata Selys, <$ Lectotype, D. E.
Kimmins det. 1968.

Currently placed in the genus Elattoneura.

wallacei Selys (Alloneura), 1886 : 183-184. LECTOTYPE <?. Type [McL. label] / New
Guinea / Alloneura wallacei Selys <$ / Alloneura wallacei Selys, £ Lectotype, D. E. Kimmins
det. 1968.

The original description states that it is based upon two specimens, collected by Wallace,

TYPE-SPECIMENS OF ODONATA IN BMNH 179

one from ? New Guinea, the other from ? Jobi, in coll. Selys. These specimens, with de
Selys' labels, were actually in the McLachlan collection (now in BMNH).

PLATYCNEMIDIDAE

alatipes McLachlan (Psilocnetnis), 1872 : 1-2, i text-fig. Holotype <$. Type [McL. label] /
[Interior of] Madagascar / Psilocnemis alatipes McL.
Currently placed in the genus Platycnemis.

arachnoides Fraser (Pseudocopera), 1922. Kimmins, 1966 : 179.

atomaria Selys (Psilocnemis), 1886 : 122-123. LECTOTYPE <J. Type [McL. label] /
Labuan / Ps. atomaria <$ [Selys' writing] / Psilocnemis atomaria Selys, <J Lectotype, D. E.
Kimmins det. 1968.

There are two males in the McLachlan collection bearing Selys' identification labels and of
these the larger (abdomen, 33 mm) has been chosen as Lectotype. There are also two
females, det. McLachlan, from N.W. Borneo, but these have no type labels. Possibly the
allotype is in the Selys collection. Currently placed in Platycnemis.

bilineata Selys (Trichocnemis), 1869 : 28. LECTOTYPE <J. Type [McL. label] / Sey-
chelles (Wright) I Trichocnemis bilineata de Selys $ / Hemicnemis bilineata Selys $ Seychelles /
Trichocnemis bilineata Selys, <J Lectotype, D. E. Kimmins det. 1968.

The lectotype $ now lacks its head. Currently placed in the genus Leptocnemis, as a
synonym of L. cyanops (Selys).

borneensis Selys (Trichocnemis), 1886 : 116. Holotype 9. Type [McL. label] / North
Borneo / Elopura, M[ar]ch, [i8]84 / Trichocnemis borneensis S. [McL's writing].

campioni Laidlaw (Coeliccia), 1917 : 224-225, text-figs 3-4. Holotype £. Borneo, Lio
Matu, 31. x. 1914 / Coeliccia campioni Type $ [Laidlaw's writing].

chaseni Laidlaw (Calicnemis), 1928 : 136. Holotype J. Jor, Pahang, 4-vi.i922 (F. Chasen) /
Calicnemis chaseni n. sp. <$ holotype [Laidlaw's writing].

The original locality label (written in the BMNH Setting Room) erroneously gave F. F.
Laidlaw as the collector. This label has been replaced. Currently placed in the genus
Calicnemia, a replacement name for Calicnemis [preocc.]. The anal appendages have been
mounted in canada balsam on celluloid.

cyaneothorax Kimmins (Coeliccia), 1936 : 89, fig. 10. Holotype <J. Sarawak, Mt Dulit,
R. Koyan, 2500 ft, Primary forest, i8.xi.i932 / Riverside / (B. M. Hobby &> A. W. Moore) /
Coeliccia cyaneothorax, <J sp. n., det. D. E. Kimmins.

cyanops Selys (Trichocnemis), 1869 : 28. Holotype Q*. Type [McL. label] / Seychelles
(Wright) I Trichocnemis cyanops (J de Selys / Hemicnemis cyanops Selys <$ Seychelles.
Currently placed in the genus Leptocnemis.

dinoceras Laidlaw (Coeliccia), 1925 : 562-563, text-figs 2-3. Holotype <£. Philippines,
Mindanao, [Landanao], Kolambugan, 18^.1914 (E. A. Wileman) / Coeliccia dinoceras <J
Laidlaw.

The type now lacks head and anal appendages.

dorothea Fraser (Coeliccia), 19336 : 466-467, figs 4a, b; 1936 : vii. LECTOTYPE <J.
TYPE, C. dorotheae $, Haldibari T[ea] E [state], Duars, Bengal (H. V. O'Donel), 3o.x.[ig]3i.
det. F. C. Fraser, sp. nov. / Coeliccia dorothea Fraser $ Lectotype, D. E. Kimmins det. 1969.
Fraser (1936 : vii) restricted the type to material in BMNH. This species was published
as dorothea, although Fraser's label gives it as dorotheae. This species was accidentally omitted
from my 1966 list of Fraser's types, probably during retyping.

flavostriata Laidlaw (Coeliccia), 1917 : 223-224, text-figs 1-2. Holotype #. Borneo, Mt
Merinjak, 28^.1914 / Coeliccia flavostriata n. sp. $ Type [Laidlaw's writing].

i8o D. E. KIMMINS

fraseri Laidlaw (Coeliccia), in Eraser, 1932 : 655-656; Laidlaw, 1932^ : 14, pi. i, figs 15, 22 ;
pi. 2, figs 9-10; pi. 3, fig. 8.

This taxon has a strange history. Strict application of priority of publication would
make it Coeliccia fraseri Fraser, since a description of it appeared in Fraser's Indian Dragon-
flies, published on 15 Feb. 1932, with reference to Laidlaw, 1931. Laidlaw's paper was
delayed and did not appear until March 1932. Fraser states 'type in B.M.' and Laidlaw 'At
present in my collection ; the types will be deposited in the British Museum.'

The BMNH has a specimen, presented by Fraser in 1937, with all the labels in Fraser's
writing, which he had marked as type ; the year of capture does not agree with the description,
and I am considering it at present as a possible syntype. I understand that part of Laidlaw's
collection was acquired by the late John Cowley, and it is therefore possible that additional
syntypes or even the actual type may be amongst his papered material (now in BMNH). I
propose therefore to defer any decision on the type of this taxon until it has been possible
to examine the Cowley Collection in detail.

hainanense Laidlaw (ssp. Coeliccia scutellum), 1932 : 23. LECTOTYPE $. Hainan, Mt
Wuchi, iy.v.[i9]o3 / Coeliccia scutellum hainanense holotype [typewritten] / Coeliccia
scutellum hainanense Laidlaw, $ Lectotype, D. E. Kimmins det. 1968.
The date of the lectotype was incorrectly published as 'ig.v.o^'.

incisa Kimmins (Risiocnemis (Prionocnemis)), 1936 : 91. Holotype $. Prionocnemis
incisa sp. n. $, det. D. E. Kimmins / Risiocnemis incisa Kim., $ Holotype, det. D. E. Kimmins,
1969.

This species was published under the name Risiocnemis (Prionocnemis), as Risiocnemis was
a newly proposed replacement name for Prionocnemis (preocc.).

kitnminsi Lieftinck (Idiocnemis), 1958 : 270-272, text-figs 56-60. Holotype $, New
Britain (A. Willey), Reg. Mar. 1.1898 / 68 / Idiocnemis nov. spec. Det. Dr. F. Ris / Idiocnemis
inornata Selys, det. H. Campion. Figured and described Entom. Hi, p. 247 (1919) / Idioc-
nemis kimminsi Lieft. det. M. A. Lieftinck, 1958, HOLOTYPE.

Lieftinck quotes the number as 89, but by comparison with the allotype $, which has the
number on the right hand end of the label, I think that the type should be 68.

[lieftincki Laidlaw (Coelicci a), 1932 : 32-33. In May 1931, prior to publication, a $ and $
were presented to BMNH by Laidlaw, who stated in his letter (3^.1931) 'C. lieftincki is at
present a MSS name.' It was naturally assumed that these were the types and they have
been so marked since that date. In the preparation of these type-lists, it was noticed that
collecting dates differed from those given in the description, and that the types were stated
to be in the Lieftinck Collection. Dr Lieftinck has informed me that he does have the types,
and the BMNH examples are therefore merely additional material, not of the type-series.
The male is in very poor condition, lacking most of abdomen.]

loogali Fraser (Coeliccia), 1932 : 652-653; Laidlaw, 19320! : 26-28, pi. I, figs 4, 27, pi. II,
17-18, pi. Ill, i. LECTOTYPE ,$. Burma, Maymyo, 3i.vii.[i9]24 (Wall) / Coeliccia loogali
Fras. COTYPE / Coeliccia loogali Fraser, Q* Lectotype, D. E. Kimmins det. 1968.

This taxon is in a position diametrically opposed to that of Coeliccia fraseri (q.v.). This
was a Fraser MSS name which was intended to be published by Laidlaw but which, through
delay in publication was antedated by Fraser himself, and in this case should be attributed
to Fraser.

macrostigtna Laidlaw (Coeliccia), 1918 : 225-227, text-figs 5-6. Holotype <$. Borneo,
Baram, 2O.x.[i9]io / Coeliccia macrostigma $ Type / Coeliccia macrostigma Laidlaw.

Holotype <$ specified by Laidlaw as 'Type $ in the British Museum'.
tnontana Fraser (Coeliccia), 1931. Kimmins, 1966 : 204.

nemoricola Laidlaw (Coeliccia (Trichocnemis)), 1912 : 95-96; 1915 : 37. Lectotype^ (by
designation of Laidlaw, 1915). Sarawak, Mt Batulawi, 27^.1911 (/. C. Moulton) / Coeliccia
(Trichocnemis) nemoricola Laidlaw, $ Type [Laidlaw's writing].

TYPE-SPECIMENS OF ODONATA IN BMNH 181

I consider that Laidlaw (1915) by his reference to 'The type of C. nemoricola ... to be

deposited in the British Museum' in effect designated the example, now in BMNH and

labelled by him as $ Type, as lectotype, especially as he distinguished the second example

as 'Co-type'.
nigrohamata Laidlaw (Coeliccia), 1918 : 228. Holotype 6*- Borneo, Mt Merinjak, a6.v.

1914 / Type, Mt Merinjak, 26/5/15 / Coeliccia nigrohamata <J Type [Laidlaw's writing] /

Coeliccia nigrohamata Laidlaw.
nipalica Kimmins (Calicnemia), 1958 : 350-351. Holotype 6*. Nepal, Phewa Tal, nr.

Pokhara, 2500 ft, 8.v. 1954 (/• Quinlari) / Calicnemia nipalica Kim., <$ Type, D. E. Kimmins

det. 1958.
pachy stigma Selys (Allocnemis), 1886 : 138-139. LECTOTYPE <J. Type [McL. label] /

Old Calabar, R[utherford], 72 / Allocnemis pachystigma Selys, $ Old Calabar / Allocnemis

pachystigma Selys, 6* Lectotype, D. E. Kimmins det. 1968.

Of the four syn types in the McLachlan collection (2 <$, Sierra Leone, 2 <J, Old Calabar), I

have chosen the most complete as Lectotype. Currently placed in Stenocnemis.

poungyi Eraser (Coeliccia), 1924. Kimmins, 1966 : 209.
pruinosa Eraser (Metacnemis), 1928. Kimmins, 1966 : 210.

pyriformis (Eraser MS) Laidlaw (Coeliccia), 1932 : 26. Holotype <$. Tonkin, Thai-Nien,
Basin of Fleuve Rouge, 2i.xii.ig24 (H. Stevens) / Coeliccia pyriforma sp. nov., 6* (Type) [in
Eraser's writing].

rectangulata Laidlaw (Calicnemia), 1932 : 97-98, fig. 2a. LECTOTYPE <J. Pahang,
F.M.S., Cameron Highlands, Kuala Boh, 3i.v.[i9]3i [no collector's name given] / Calicnemia
rectangulata $ Type [Laidlaw's writing].

Laidlaw lists two examples but gives no published designation as to holotype. The only
example in BMNH is marked Type by him and has been chosen as the Lectotype.

reflexa Kimmins (Risiocnemis), 1936 : 92-93, fig. 12. Holotype $. N.W. Borneo / Prionoc-
nemis reflexa $ sp. n., det. D. E. Kimmins / Risiocnemis reflexa Kim., $ Holotype, det. D. E.
Kimmins, 1969.

ruflpes Selys (Metacnemis), 1886 : 139-140. Holotype <J. Type [McL. label] / Camaroons,
R[utherford] 37 / Alloneura rufipes Selys n. sp. <$ Camaroons.
Currently placed (with doubt) in Platycnemis.

salomonis Kimmins (Lieftinckia), 1957 : 3I4~3I5. text-fig, i. LECTOTYPE <J. Solomon
Islands, Guadalcanal, Tapenanje, 10-23. xii. 1953 (J- D. Bradley) / Lieftinckia salomonis
Kimm., $ Type, D. E. Kimmins det. 1955 / Lieftinckia salomonis Kim., <J Lectotype, D. E.
Kimmins det. 1969.

Although the above example was labelled Q* Type, no type-designation was made in the
original description.

scutellum Laidlaw (Coeliccia), 19320. : 22, pi. Ill, fig. 18. LECTOTYPE <J. Tonkin, Bao
Ha, ao.ix.[i9]23 (H. Stevens) / Coeliccia scutella [sic] $, sp. nov. (Type) [in Eraser's writing] /
Coeliccia scutellum Laidlaw, <J Lectotype, D. E. Kimmins det. 1968.

Of the two examples listed, only the above lectotype has been traced. The date of this is
incorrectly given in the original description as 2o.iv.24-

subaequistyla Eraser (Copera), 1928. Kimmins, 1966 : 214.
superplatypes Eraser (Copera), 1927. Kimmins, 1966 : 215.

COENAGRIIDAE

abercornensis Pinhey (ssp. Aciagrion steelae) 1958 : 103-104, fig. 2. LECTOTYPE $.
N[orthern] R[hodesia], Abercorn, Lake Chila, iv.i954 (E. Pinhey) / Aciagrion steelae abere-

i82 D. E. KIMMINS

cornensis Pinhey 1956, TYPE <$ / Aciagrion steelae abercornensis Pinhey, $ Lectotype, D. E.
Kimmins det. 1969.

adarnsi Calvert (Argia), 1901 : 80-81, pi. iv. figs 35, 353. Holotype $. [Panama], Bugaba,
800-1500 ft (Champion) / Argia adamsi Calv. COTYPE, P. P. Calvert det. 1901. Original of
flf. 35, 355, pi. iv. B.C. A. Neur.

adytum Perkins (Agrion), 1899 : 69. LECTOTYPE <J. Hawaiian Islands (R.C.L. Perkins] /
Kauai, Waimea, 4000 ft, vi.i894 (Perkins) / Agrion adytum Prk. / Agrion adytum Prk.,
Type / Agrion adytum Prk., $ Lectotype, D. E. Kimmins det. 1969.

Tenth abdominal segment of lectotype detached and mounted on card. Currently placed
in Megalagrion.

albistigma Eraser (Ischnura), 1927. Kimmins, 1966 : 177.

alcyone Laidlaw (Agriocnemis), 19310 : 249-250. LECTOTYPE <J. Brit. N. Borneo,
Bettotan, 26.vii.i927 (H. M. Pendlebury) / Agriocnemis alcyone <$ [Laidlaw's writing] /
Agriocnemis alcyone Laidl., $ Lectotype, D. E. Kimmins det. 1969.

The locality label of this specimen was inaccurate and has been replaced, using information
from the original description. The type series comprised 3 $, i $, and the above example
was given to BMNH as the holotype, although not labelled as such. It has therefore been
designated Lectotype.

aluensis Campion (Teinobasis), 1924 : 614, fig. 2. Holotype <J. Solomon Is., Alu (C. M.
Woodford) / Teinobasis aluensis Campion, $ Holotype, det. D. E. Kimmins, i8.ix.i933.

amaurodytum Perkins (Agrion), 1899 : 66-68. LECTOTYPE <J. Hawaiian Islands /
Molotai Mts, 4000 ft +, viii.i8g3 (Perkins) / Agrion amaurodytum Prk. Type / Agrion
amaurodytum Perk., $ Lectotype, D. E. Kimmins det. 1969.
Currently placed in Megalagrion.

angolense Longfield (Agriocnemis), 1947 : 15-17, figs 6 A-E. Holotype <J. Angola
M[ission] S[cientifique] S[uisse], 1933 / SangeVe1, n.ii.[i933] / Agriocnemis angolense Type $,
det. Miss C. Longfield.

There are also allotypes of the homochrome and heterochrome females.

[apicale Selys (Acanthagrion), 1876 : 306-307.

This species was based upon three $ examples ; one from Para (Bates) in the Selys Collection
and two from Peba in the McLachlan collection. As the name apicale was a Bates MSS
name, I am considering the <J example in the Selys collection as the LECTOTYPE and the
BMNH examples as paralec to types.]

argentea Tillyard (Agriocnemis), 1906 : 192-193, pi. 17, figs loa, b. LECTOTYPE (J.

N. Queensland], Kuranda, xii.[ig]o4 (R. J. Tillyard) / Agriocnemis argentea Till., $ TYPE
R.J.T. / Agriocnemis argentea Till., <J Lectotype, D. E. Kimmins det. 1969.

This species is placed by Eraser (1960) as a synonym of Agriocnemis exsudans Selys.

armstrongi Eraser (Amorphostigma), 1925. Kimmins, 1966 : 179.
arthuri Eraser (Mortonagrion), 1942. Kimmins, 1966 : 179.

asteliae Perkins (Agrion), 1899 : 66. LECTOTYPE <J. Honolulu, Oahu, 3000 ft, vii.iSgs
(Perkins) / Agrion asteliae Prk., Type / Agrion asteliae Perkins, <J Lectotype, D. E. Kimmins
det. 1969.

Currently placed in Megalagrion.

attenuatum Eraser (Aciagrion), 1928. Kimmins, 1966 : 180.

aureofrons Tillyard (Psuedagrion), 1906 : 189-191, pi. 17, fig. 8. Lectotype <$. (Watson
1969 : 71). N[orth] Queensland], Atherton, i.[i9]o5 (R. J. Tillyard) / Pseudagrion aureo-
frons. cJ Lectotype, Tillyard, 1906, design by J. A. L. Watson, 1969.

auricolor Eraser (Amorphostigma), 1927. Kimmins, 1966 : 180.

TYPE-SPECIMENS OF ODONATA IN BMNH 183

[australis Selys (Agriocnemis), 1877 : 155-156.

The unique $ (Queensland) is said by Selys to be in the McLachlan collection, but it has
not been traced.]

azureum Eraser (Aciagrion), 1922. Kimmins, 1966 : 181.

beadlei Pinhey (Pseudagrion), 1961 : 33-34, pi. 2, fig. n. Holotype <J. Uganda, Jinja,
vi.i949 (E. Pinhey} [ Pseudagrion beadlei Pinhey 1956, Type $, HOLOTYPE.

bellona Laidlaw (Ceriagrion), 19150 : 274. Holotype (J. Sarawak, Mt Matang, 4.xii.i9i3
(/. C. Moultori) I Ceriagrion sp. nov. [H. Campion's writing] $ Type [Laidlaw's writing] /
Ceriagrion bellona Laidlaw, $ Holotype, D. E. Kimmins det. 1969.

bidens Kimmins (Nesobasis), I958« : 239-241, figs 1-2. Holotype #. New Hebrides,
Aneityum, Red Crest, 1200 ft, 3 miles N.E. of Anelgauhat, vi.i955 (L. E. Cheesmari) / Neso-
basis bidens Kim., (J TYPE, D. E. Kimmins det. 1957.

bidentatum Eraser (Ceriagrion), 1941. Kimmins, 1966 : 182.

blackburni McLachlan (Megalagrion), 1883 : 238-239. LECTOTYPE <J. Type [McL.
label] / Hawaiian Islands, [at the head of Wailuku Valley, Mani] (Blackburn) / 26 / Megal-
agrion Blackburni, McL. / Megalagrion blackburni McL., <J Lectotype, D. E. Kimmins det.
1969.

The lectotype lacks the apical third of the right fore wing and the whole of the left hind
wing.

bradleyi Kimmins (Teinobasis), 1957 : 315-316, fig. 2. LECTOTYPE <J. Solomon Is.,

Guadalcanal, Tapenanje, io-23.xii.i953 (/. D. Bradley) / Teinobasis bradleyi Kim., $ TYPE,

D. E. Kimmins det. 1955 / Teinobasis bradleyi Kim., $ Lectotype, D. E. Kimmins det. 1969.

Although the type was labelled as such, no holotype was specified in the original description.

brisbanense Tillyard (Agrion), 1917 : 477-479, pi. 23, figs 13-14. Holotype <J. Q[ueens-
land], Brisbane, [Kedron Brook], 22.i.[i9]i3 (R. J. Tillyard) / Agrion brisbanensis Till.,

<J TYPE, R.J.T.

Currently placed in the genus Coenagrion.
buxtoni Eraser (Ischnura), 1927. Kimmins, 1966 : 184.

calliphya McLachlan (Agrion?), 1883 : 236-237. Holotype (J. Type [McL. label] / Hawaiian
Islands [Lanai, about 2000 ft (Blackburn)] / 55 / Agrion? calliphya, McL.

Currently placed in the genus Megalagrion.

cardinalis Eraser (Oxyagrion), 1946. Kimmins, 1966 : 185.

cardinalis Kimmins (Ischnura), 1929 : 224-225, 3 text-figs. Holotype <J. Society Is., N.

Raiatea, 1000 ft, 31^.1925 ([L. E.] Cheesman) / Ischnura cardinalis Kimmins, $ Type, det.

D. E. Kimmins.
castellani Roberts (Coenagrion), 1948 : 63-68, 2 text-figs. Holotype <$. Italy, Rome,

Acilia, i.vi.i946 (O. Castellani) j C. castellani Rob. ^ Type [F. C. Eraser's writing] / Penis

detached and mounted on card.

Owing to Dr Roberts' illness, Col. F. C. Eraser undertook the preparation of Roberts'

manuscript for press, which is also the reason for the determination label in Eraser's hand-
writing.
cervula Selys (Ischnura), 1876 : 262-263. LECTOTYPE <J. Type [McL. label] / California

(Edwards) / 219 / Ischnura cervula Selys <J / Ischnura cervula Selys, <J Lectotype, D. E.

Kimmins det. 1969.

Of the pair mentioned by Selys, the $ now lacks abdominal segments 6-10 and the left

hind wing. The allotype $ also lacks the abdominal segments 6-10 and most of both left wings.

ceylanica Kirby (Archibasis), 1891 : 205-206, pi. xx, fig. 4. Holotype <J. Ceylon, Kandy,
viii.i888 (E. E. Green) / Kandy, Aug. 88 / A. ceylanica Kb. type.
Currently placed in Pseudagrion.

184 D. E. KIMMINS

cheesmanae Fraser, (Ischnura), in Mumford, 1942 : 646, figs 1-3. Holotype (J. Tahiti,
Hitiaa, io.vii.i925 (Cheesman [L. £.]) / Ishnura sp. [sic.] / I. cheesmani [sic] Fraser in Mum-
ford (Type), det. F. C. Fraser, 1959. Locality [sic] of type given in error as in the Morton
colln. F.C.F.

chelata Calvert (Argia), 1902 : 88, pi. iv. figs 47, 473. Holotype $. [Costa Rica, Volcan de]
Irazu, 6-7000 ft (H. Rogers] / Argia Lachrymans H. $ [Selys' writing] / Argia chelata Calv.
$ TYPE, P. P. Calvert det. 1901. Original of ff. 47, 473, pi. iv, B.C. A. Neur.

circulatutn Selys (Enallagma), 1883 : 133-135. LECTOTYPE <?. Type [McL. label] /
Japan (Lewis) / cobalt blue, 2O.viii.[i8]8i / Agrion circulatum Selys, $ Japon / Enallagma
circulatum Selys, <$ Lectotype, D. E. Kimmins det. 1969.

Although labelled 'Agrion' by Selys, it was published as 'Enallagma' '.

citrinutn Campion (Ceriagrion), 1914 : 278-279. Holotype $. [S. Nigeria], Lagos,
(Strachan) / Ceriagrion citrinum Campion, Holotype. Determined by H. Campion.

clausen i Fraser (Agriocnemis) , 1922. Kimmins, 1966 : 186.

coelestis Longfield (Pseudagrion) , 1947 : 9-10, fig. 3. Holotype $. Angola, M[ission]
S[cientifique] S[uisse], 1933 / Mupa, viii / Pseudagrion coelestis Type <$, det. Miss Longfield.

coeruleiceps Longfield (Pseudagrion), 1959 : 19-21, figs 3, d, g, h. Holotype <$. Nigeria,
Vom, Stock Farm, 7-ix.i956 (R. M. Gambles) / Pseudagrion coeruleiceps Type <$, det. Miss
Longfield.

coeruleum Tillyard (Pseudagrion), 1908 : 739-741, pi. 30, figs 13-14. LECTOTYPE <$.
W. A[ustralia], Perth, i.[i9]o7 (R. J. Tillyard) / Pseudagrion coeruleum Till., <$ TYPE, R. J. T. /
Pseudagrion coeruleum Till., <$ Lectotype, D. E. Kimmins det. 1969.
Currently placed in the genus Austroagrion.

collopistes Calvert (Telebasis), 1902 : 116-117, pi- v> n8s 27-28. Holotype $. Mexico,
Teapa, i (H. H. S[mith]) / Telebasis collopistes Calv., P. P. Calvert det. 1902, Original of
Tab. V, figg. 27, 28.

corallinum Campion (Ceriagrion), 1914 : 279-281. Holotype <$. Sierra Leone, Port Lokko,
i.v.1912 (/. /. Simpson) / Ceriagrion corallinum Campn., Holotype. Determined by H.
Campion.

crocops Selys (Pseudagrion), 18760 : 512-513. Holotype <$. M6nado [Celebes] / Agrion
crocops Selys (pres de pruinosum) <£ / Pseudagrion crocops Selys $.

cupraurea Calvert (Argia), 1902 : 85, pi. iv, figs 24, 42. Holotype <£. Panama, David
(Champion) / Argia cupraurea Calv. (J, P. P. Calvert det. 1901, Original of f. 42, pi. iv. B.C. A.
Neur.

deceptor McLachlan (Agrion?), 1883 : 235-236. Holotype <J. Type [McL. label] / Hawaiian
Islands [Oahu, (Blackburn)] / 67 / Agrion? deceptor McL.
Currently placed in the genus Megalagrion.

decoloratum Fraser (Libyagrion), 1928. Kimmins, 1966 : 188.

descendens Fraser (ssp. Acanthagrion apicale), 1946. Kimmins, 1966 : 189.

digiticollis Calvert (Telebasis), 1902 : 118, pi. v, fig. 21. Holotype $. [Mexico], Tabasco,
Teapa, i (H. H. S[mith]) / Telebasis digiticollis Calv. P. P. Calvert det. 1902. Original of
Tab. v, fig. 21.

dolorosa Fraser (Pacificagrion), 1953. Kimmins, 1966 : 189.

[dorothea Fraser (Coenagrion), 1923 : 462-463, figs. 8-8c.

A female example in BMNH has been labelled Allotype, since there was only one female
in the type-series. The male type presumably went to the Indian Museum collection,
although Fraser does not mention the location of any types. It may be mentioned that the
specific name is pre-occupied by Coenagrion dorothea (Fourcroy, 1785).]

TYPE-SPECIMENS OF ODONATA IN BMNH 185

[draconis Barnard (var. of Pseudagrion kersteni), 1937 : 2I3-2I5. fig- J4 g-

The BMNH has one example, received from Barnard, labelled 'kersteni var. draconis Type'.
Pinhey (1964 : 26-27) has studied <J, $ types and describes a 'holotype <$' and 'allotype $'.
I regard his action as in effect designating a $ lectotype, and our example is therefore only
one of the type-series.]

dubium Laidlaw (Pseudagrion?), 1912 : 97-98, fig. 5. Holotype <$. [Sarawak], Mt Batu
Lawi, 27.v.[i9]n, Pseudagrion [deleted] Stenagrion dubium Type $ [in Laidlaw's writing] /
Gen. spec ? (Pseudagrion ??).

Currently placed in the genus Stenagrion.

dundoense Longfield (Pseudagrion), 1959 : 24-25, fig. 2. Holotype <$. N. Angola, Lunda
Prov., Dundo, iii.i949 / Pseudagrion dundoense Type $, det. Miss C. Longfield.

ecornutum Selys (Agrion), 1872 : 44-45. LECTOTYPE <J. Type [McL. label] / Amur
Land / 272 / Agrion ecornutum de Selys <$ / Agrion ecornutum Selys, <$ Lectotype, D. E.
Kimmins det. 1969.

Currently placed in Coenagrion.

eudytum Perkins (Agrion), 1899 : 68. LECTOTYPE £. Hawaiian Islands (R, C. L.
Perkins) / Kauai, Lihue, [about 1000 ft], 1896 (Perkins) / Agrion eudytum $ Prk., $ Lectotype,
D. E. Kimmins det. 1969.

Currently placed in Megalagrion.

euphorbia Fraser (ssp. Argia extranea), 1966. Kimmins, 1966 : 191.

exclamationis Campion (Austroagrion), 1915 : 106-108, i fig. Holotype <$. [Australia],
N[orthern] Territory], Koolpinyah, 6.iii.[i9]i3 (G. F. Hill) / I[mperial] B[ureau of] E[nto-
mology] 421 / Austroagrion exclamationis Cmpn., $ Holotype, Determined by H. Campion.

fallax Perkins (var. of Agrion amaurodytum) , 1899 : 67-68. LECTOTYPE <J. Hawaiian
Islands (R. C. L. Perkins) / Hawaii, Kauai, 4000 ft, x.i8g6 (Perkins) / Agrion amaurodytum,
var. of var. peles Prk. / Set out as type of fallax by C. H. Kennedy. Fallax types missing &
Perkins suggested they might be under peles. = fallax description and is marked by Perkins
as 'var. of var. peles.' See top label [Kennedy's writing] / Agrion amaurodytum var. fallax
Prk., (J Lectotype, D. E. Kimmins det. 1969.
Currently placed in Megalagrion.

floridense Fraser (ssp. Acanthagrion gracile), 1946. Kimmins, 1966 : 192.
forficula Fraser (ssp. Argia extranea), 1946. Kimmins, 1966 : 193.

fragilis Tillyard (Ischnura), 1906 : 186-187, P1- J7. figs 6a. b- LECTOTYPE <$. N[orth]
Queensland], Atherton, i.[ig]o5 (R. J. Tillyard) / Aciagrion fragilis Till., <$ TYPE, R.J.T. /
Ischnura fragilis Till., £ Lectotype, D. E. Kimmins det. 1969.

Although originally described as Ischnura, Tillyard appears to have written new deter-
mination labels when he transferred the species to Aciagrion in 1912. The lectotype now
lacks the right fore wing.

gangetica Laidlaw (Ischnura), 191301 : 235-236, text-fig. Holotype <$. N. India, Shamket,
Kamaon, i6.v.[i9]i2 (A. D. Imms) / For. Zool. Coll., Shamket, Kamaon, i6.v. 1916 / Ischnura
gangetica Type <$ [Laidlaw's writing].

The type was returned to the Forest Research Institute and subsequently presented to
BMHN by Dr. A. D. Imms. Currently placed as a synonym of Ischnura for cipata Morton.

gautama Fraser (Indagrion), 1922. Kimmins, 1966 : 194.

granti McLachlan (? Ischnura), 1903 : 402, pi. 24a, figs 1-2. LECTOTYPE <J. Sokotra,

Dahamis, feet, I9.xi.[i8]g8 (W. R. O. Grant) / Ischnura granti TYPE <J / Ischnura (?)

granti McL., <$ Lectotype, D. E. Kimmins det. 1969.
Currently placed in the genus Enallagma.

186 D. E. KIMMINS

guichardi Kimmins (Pseudagrion), 1958 : 351-352, figs 1-2. Holotype <J. Ethiopia, nr
Cencia, 29.111.1948 (K. M. Guichard) / Pseudagrion guichardi Kim., $ TYPE, D. E. Kimmins
det. 1958.

haemastigma Eraser (Ischnura), 1927. Kimmins, 1966 : 194.
hamulata Eraser (Argia), 1946. Kimmins, 1966 : 194.

hawaiiense McLachlan (Agrion?), 1883 : 232-234. LECTOTYPE $. Hawaiian Islands /
66 $ / [Oahu, at no great elevation above the sea (Blackburn)] / Agrion? hawaiiense McL. /
Agrion? hawaiiense McL., $ Lectotype, D. E. Kimmins det. 1969.

Currently placed in the genus Megalagrion. McLachlan quotes 'No. 16', but this must be
a printer's error as the three syntypes are numbered '66'.

herberti Calvert (Argia), 1902 : 82, pi. iv. figs 37, 375. Holotype <$. [Mexico], Guerrero,
Amula, 6000 ft (H. H. Smith) / Argia herberti Calv. <J Type, P. P. Calvert det. 1901. Original
of ff. 37, 375, B.C.A. Neur.

heterogamias Perkins (Agrion), 1899 : 77, pi. v, fig. 3. LECTOTYPE $. Hawaiian Islands
(R. C. L. Perkins) f Kauai, Waimea, 4000 ft, v.i8g4 (Perkins) / Agrion heterogamias Prk.,
<J Type / Agrion heterogamias Prk., <$ Lectotype, D. E. Kimmins det. 1969.
Currently placed in Megalagrion.

huallaga Eraser (Argia), 1946. Kimmins, 1966 : 196.

hyacinthus Tillyard (Agriocnemis), 1913 : 457-459, pi. 48, figs 15, 16. LECTOTYPE <J.

N[ew] S[outh] W[ales], Sydney, I2.xii.[i9]o8 (R. J. Tillyard) / Agriocnemis hyacinthus Till.,
^ TYPE, R.J.T. / Agriocnemis hyacinthus Till., $ Lectotype, D. E. Kimmins det. 1969.

ignifer Tillyard (Pseudagrion), 1906 : 188-189, pi. 17, figs 7a, b. LECTOTYPE <J. N[orth]
Queensland], Kuranda, xii.[i9]o4 (R. J. Tillyard) / Pseudagrion ignifer Till., $ TYPE, R.J.T. /
Pseudagrion ignifer Till., <J Lectotype, D. E. Kimmins det. 1969.

ignitum Campion (Ceriagrion), 1914 : 281-282. Holotype (J. Gold Coast, Aburi, 1912-13
(W. H. Patterson) / 628 / Ceriagrion ignitum Campion, <$ Holotype, Determined by H.
Campion.

immsi Laidlaw (Enallagma), igi^a : 236-237, i text-fig. Holotype <J. India, Centr[al]
Provinces], Sonder Bhandara, 5.xii.i9i2 (Dr A. D. Imms) j For. Zool. Coll. Sonder Bhandara,
5.xii.i9i2 / Ischnura immsi Type $ [Laidlaw's writing].

Holotype returned to Indian Forest Research Institute and subsequently presented to
BMNH by Dr Imms. Currently placed as a synonym of Enallagma sparsum Selys.

indicatrix Calvert (Argia), 1901 : 82-83, pi- iv> ngs 23» 39. 39s- Holotype <$. [Mexico],
Tabasco, Teapa, ii (H. H. S[mith]) / Argia indicatrix Calv. <J TYPE, P. P. Calvert det. 1901.
Original of ff. 39, 393, pi. iv, B.C.A. Neur.

indicum Fraser (Pseudagrion), 1924. Kimmins, 1966 : 197.
infrequentula Fraser (Argia), 1946. Kimmins, 1966 : 197.
interrupta Fraser (Agriocnemis), 1927. Kimmins, 1966 : 198.

interruptum Selys (Acanthagrion), 1876 : 314-316. LECTOTYPE <J. Valparaiso / 13? /
Acanthagrion interruptum Selys <J [Selys label] / Acanthagrion interruptum Selys, <$ Lecto-
type, D. E. Kimmins det. 1969.

The McLachlan collection contained 2 <$ examples (one incomplete) labelled Valparaiso,
and 2 (J, i $ labelled Chili. They did not bear McLachlan type labels, but as Selys does not
mention any examples in his own collection, the lectotype has been chosen from the two from
Valparaiso. The other examples are considered as paralectotypes.

jacksoni Pinhey (Pseudagrion), 1961 : 37-38, pi. 2, figs i, 10. Holotype <J. Uganda,
Aswa R[iver], Karamoja, iii.[i9]52 (T. H. E. Jackson) / Pseudagrion jacksoni Pinh., HOLD-
TYPE [<?].

TYPE-SPECIMENS OF ODONATA IN BMNH 187

jugorum Perkins (Agrion), 1899 : 72-73, pi. v, figs 2, u, na. LECTOTYPE <J. Hawaiian
Islands (R. C. L. Perkins) / West Maui Mts, 4000 ft, iv.i8g4 (Perkins) / A. jugorum, 12 /
Agrion jugorum Prk., Type / Agrion jugorum Perk., <$ Lectotype, D. E. Kimmins det. 1969.
Currently placed in Megalagrion.

kauaiense Perkins (Agrion), 1899 : 75-76. $ LECTOTYPE. Hawaiian Islands (R. C. L.
Perkins] / A. kauaiense / Agrion Kauaiense Perk., Type / Agrion kauaiense Prk., $ Lectotype,
D. E. Kimmins det. 1969.

Currently placed in the genus Megalagrion.

kibalense Longfield (Pseudagrion), 1959 : 22-23, ng- I- Holotype <$. Central Africa,
Uganda, Toro Dist., Kibale Forest, Buzirasagama, 21.1.1938 (C. E. Longfield) / Pseudagrion
kibalense Longfield, <$ Type, det. Miss C. Longfield.

lachrytnosa Eraser (Paciflcagrion), 1926. Kimmins, 1966 : 200.

laidlawi Kimmins (Teinobasis), 1936 : 95-97, figs 13, 14. Holotype ^. Brit[ish] N[orth]
Borneo, Bettotan, 23.vi-i7.viii.[i9]27 (C. B[oderi\ K[loss] & H. M. P[endlebury~\) / Teinobasis
sp. n.?, compared with type of T. kirbyi Laidl. <$, 4.11.1935, J. Cowley / Teinobasis laidlawi
Kimmins <$, det. D. E. Kimmins / Teinobasis laidlawi Kim., <$ Holotype, D. E. Kimmins
det. 1969.

[lanceolatutn Selys (Agrion), 1872 : 43-44. In the BMNH are three rather battered syn-
types from the McLachlan collection. In view of their condition, it seems that a lectotype
should be selected from material in the Selys Collection.]

laterals Selys (Acanthagrion), 18760 : 317-319. LECTOTYPE <J. New Granada (Nolcken) /
Acanthagrion laterale de Selys <$ / Acanthagrion laterale Selys, $ Lectotype, D. E. Kimmins
det. 1969.

leoninum Selys (Argiagrion), 18760 : 530-531. Holotype $. Type [McL. label], Sierra

Leone, 32 [3 labels on one card] / Telebasis leonina de Selys / Argiagrion Leoninum Selys $.

The holotype now possesses only the right mid leg.
leptodemas Perkins (Agrion), 1899 : 70. Holotype^. Hawaiian Islands (R. C. L. Perkins) /

Oahu, Halemano, 2000 ft, 11.1893 (Perkins) / A. leptodemas / Agrion leptodemas Prk. Type.

Currently placed in Megalagrion.
leveri Kimmins (Nesobasis), 1943 : 698-700, 5 figs. Holotype $. Fiji, Nadarivatu, 17.1.

[1943] (N.I54) (R. A. Lever) / Nesobasis leveri Kim., £ TYPE, det. D. E. Kimmins, vii.i943.
longfieldae Fraser (Enallagma), 1947. Kimmins, 1966 : 201.
[louisae Laidlaw (Amphicnemis), 1913 : 71. There is in BMNH the female syntype, which

now lacks the apex of the abdomen. I have no information as to the location of the <$ syntype.

It is possible that it may have passed into the John Cowley collection from Laidlaw, and I

therefore refrain from designating a lectotype until the Cowley Collection (now in BMNH)

can be thoroughly examined.]
lyelli Tillyard (Agrion), 1913 : 449-451, pi. 49, figs i, 2, 20. LECTOTYPE $. Vic[toria],

Gisborne, 22.xii.[ig]o8 (R. J. Tillyard) / Agrion lyelli Tillyard, £ TYPE, R. J. T. / Agrion

lyelli Till., $ Lectotype, D. E. Kimmins det. 1969.

Currently placed in the genus Coenagrion.
maclachlani Selys (Agriocnemis), 1877 : 152-153. LECTOTYPE <$. Type [McL. label] /

Gaboon / 249 / Agriocnemis MacLachlani de Selys [Selys' writing] / Agriocnemis maclachlani

Selys <J Lectotype, D. E. Kimmins det. 1969.

Selys states that a couple were given to him by McLachlan, but he must have seen more

examples, because McLachlan's collection contains 2 <J, i $ bearing Selys' labels.
madelenae Laidlaw (Amphicnemis), 1913 : 71. LECTOTYPE <$. Sarawak, Matang Road,

H.xi.[i9]o9 (/. C. Moulton) / Amphicnemis madelenae Laidlaw <$ Type / Amphicnemis

madelenae Laidl., $ Lectotype, D. E. Kimmins det. 1969.
The lectotype now lacks its head.

188 D. E. KIMMINS

magnanimum Selys (Pseudagrion), 18760 : 516-517. Holotype <$. Aru / Telebasis mag-
nanimum De Selys £ / Pseudagrion magnanimum Selys <$.
Currently placed in the genus Papuagrion.

makoka Eraser (Argia), 1946. Kimmins, 1966 : 202.
tnalabaricum Fraser (Pseudagrion), 1924. Kimmins, 1966 : 203.

tnaldivense Laidlaw (Enallagma), 1902 : 221-222. LECTOTYPE $ (very incomplete).
Maldives, Hulule, 2O.vi.[i9]oo (Gardiner] / Enallagma(?) maldivense sp. n. Type [Laidlaw's
writing] / Enallagma(?) maldivense Laidl., $ Lectotype, D. E. Kimmins det. 1969.

The lectotype now consists of the thorax and the right posterior wing. There is also a
$ paralectotype, lacking head and apex of abdomen. These examples have lost their BM
register labels, so that there is no record as to how or when they came to the Museum.

malekulana Kimmins (Nesobasis), 19360 : 73-74, figs. 5-6. Holotype Q*. New Hebrides,
Malekula, Ounua, iii-iv.i929 (L. E. Cheesmari) / Nesobasis malekulana sp. n., <$ Holotype,
det. D. E. Kimmins.

melanicterum Selys (Pseudagrion), 18760 : 492-493. LECTOTYPE <£. S[ierra] Leone /
Pseudagrion melanicterum Selys, D. E. Kimmins det. 1961. I believe this to be the 6* type /
Pseudagrion melanicterum Selys, <$ Lectotype, D. E. Kimmins det. 1969.

melidora Lieftinck (Palaiargia), 1953 : 241-244, fig. 4. Holotype (J. N. Dutch New Guinea,
Waigeu, Camp Nok, 2500 ft, iv.i938 (L. E. Cheesman) / lat. stripes of thorax pale gr[een],
dorsally blue, mac[ulae] of Abd[omen] blue / Palaiargia melidora Lft., <J HOLOTYPE.

microdernas Perkins (ssp. Agrion calliphya), 1899 : 71. LECTOTYPE $. Hawaiian
Islands (R. C. L. Perkins) j Hawaii, Kona, 3000 ft, ix.i892 (Perkins) / Agrion calliphya
McLach. race microdernas = Hawaii specimens / Agrion calliphya var. microdernas Prk. /
Agrion calliphya var. microdernas Perk., $ Lectotype, D. E. Kimmins det. 1969.
Currently placed in Megalagrion.

mildredae Fraser (Ischnura), 1927. Kimmins, 1966 : 204.

miniopsis Selys (Oxyagrion), 1876 : 299-300. Holotype <J. Bogota (Nolcken) / Telebasis
miniopsis Selys $ / Oxyagrion miniopsis Selys $ / Oxyagrion miniopsis Selys, <$ Holotype,
D. E. Kimmins det. 1969.

This example was not labelled Type by McLachlan and was only recognized as the holo-
type during the preparation of this type-list.

mishuyaca Fraser (Argia), 1946. Kimmins, 1966 : 204.
mollusca Fraser (Argia), 1946. Kimmins, 1966 : 204.

molokaiense Perkins (Agrion), 1899 : 73. LECTOTYPE 6*. Hawaiian Islands (R. C. L.
Perkins) / Molokai Mts, 4000 ft (Perkins) / A. molokaiense / Agrion molokaiense Prk. Type /
Agrion molokaiense Prk., $ Lectotype, D. E. Kimmins det. 1969.
Currently placed in the genus Megalagrion.

monardi Longfield (Pseudagrion), 1947 : 12-15, ng- 5- Holotype <J. Angola, Miss[ion]
sc[ientifique] suisse, 1932-33 / Ebanga, xi / Pseudagrion monardi Type Q*, det. Miss C. Long-
field.

moorei Longfield (Ceriagrion), 1952 : 44-46, fig. 3. Holotype^. Gold Coast, N. Territories,
Yapi, (/. /. Simpson) / Ceriagrion moorei sp. n. Type <$, det. Miss C. Longfield / figured.

[nahuana Calvert (var. of Argia agriodes), 1902 : 99, pi. 4, fig. 6233.

There is a series of cotypes in BMNH, but none has an indication that it was the original
of pi. 4, fig. 6253, and therefore the type must be in some other collection.]

naia Fraser (Agriocnemis), 1932. Kimmins, 1966 : 205.

nesiotes Perkins (Agrion), 1899 : 72. LECTOTYPE <J. Hawaiian Islands (R. C. L. Perkins)
I Hawaii, Puna, xii.i896 (Perkins) / Agrion nesiotes Prk. / Agrion nesiotes Prk., Type /

TYPE-SPECIMENS OF ODONATA IN BMNH 189

Agrion nesiotes Prk., £ Lectotype, D. E. Kimmins det. 1969.
Currently placed in Megalagrion.

nigra Campion (Teinobasis), in Laidlaw, 1928 : 136-138, figs 2, 3. Holotype <£. [Philip-
pines], Luzon, Mt Makiling (Baker) / Teinobasis nigra, <$ Holotype, Determined by H.
Campion.

nigroflavum Fraser (Ceriagrion), 1933. Kimmins, 1966 : 206.

nigrolineatum Perkins (var. of Agrion nigrohatnatum), 1899 : 65. LECTOTYPE $.
Hawaiian Islands (R. C. L. Perkins] / Oahu, Waianae Mts, iv.iSga (Perkins) / A. nigroha-
matum Blk., race nigrolineatum / Agrion nigrohamatum var. nigrolineatum Prk. / Agrion
nigrohamatum var. nigrolineatum Prk., $ Lectotype, D. E. Kimmins det. 1969.

Although not marked as Type by Perkins, the example designated as Lectotype bears a
manuscript determination label and has for many years been labelled as type in the BMNH.
Currently placed in Megalagrion.

novaehispaniae Calvert (ssp. Enallagma coecum), 1907 : 381. Holotype $. Vera Cruz,
Atoyac, iv (H. H. S[mith]) / Enallagma coec[um] nov[ae] hisp[aniae] Calv. P. P. Calvert
det. 1907. B.C. A. Neur. p. 381 (113). TYPE.

obsoletum Selys (Leptagrion?), 18766 : 985-986. LECTOTYPE <J. Obydos (Traill),
24.i.[i8J74 / Leptagrion obsoletum Selys, <$ Obydos / Leptagrion ? obsoletum Selys, <J Lecto-
type, D. E. Kimmins det. 1969.

oceanicum McLachlan (Megalagrion), 1883 : 239-240. Holotype <J. Type [McL. label] /
Hawaiian Islands [Oahu, at no great elevation above the sea (Blackburn)] / 67 / Megalagrion
oceanicum, McL.

[olivaceum Laidlaw (Ceriagrion), 1914 : 345-346, pi. 16, fig. 9.

The BMNH has a pair, marked cotypes. The types however should be in the Indian
Museum and our examples are considered as paratypes.]

oresitrophum Perkins (Agrion), 1899 : 60-70. LECTOTYPE <$. Hawaiian Islands (R. C. L.
Perkins) / Kauai, Halemanu, 4000 ft, v.i8g5 (Perkins) / Agrion oresitrophum Prk., Type /
A. oresitrophum / Agrion oresitrophum Prk., $ Lectotype, D. E. Kimmins, det. 1969.

Currently placed in Megalagrion.

orobates Perkins (Agrion), 1899 : 70. Holotype $. Hawaiian Islands (R. C. L. Perkins) /
Kauai, Waimea, v.i8g4 (Perkins) / Agrion orobates Perk., Type.

Currently placed in Megalagrion.

pacificum McLachlan (Agrion?), 1883 : 234-235. LECTOTYPE <$. Type [McL. label] /
Hawaiian Islands [Lanai and Maui (incorrectly Oahu), at various elevations] / 53 / Agrion?
pacificum McL. / Holotype, C. H. Kennedy. Sept. 1925 / Agrion ? pacificum McL., $ Lecto-
type, D. E. Kimmins det. 1969.

I have chosen as Lectotype the <$ labelled holotype by Kennedy, 1925, and which has for
many years carried a BM Type label. Currently placed in the genus Megalagrion.
pollens Calvert (var. Argia violacea), 1902 : 98, pi. iv. figs 25, 61, 6is. Allotype ?. [Mexico],
Guadaljara, Jalisco, vii (Schumann) / Argia violacea pallens Calv., $ Cotype, P. P. Calvert
det. 1901. Original of fig. 25, pi. iv, B.C. A. Neur.

Although Calvert labels this cotype, in common with others of the type-series, as the
original of a figure, I consider it as allotype 9-
pallidum Fraser (Ceriagrion), 1933. Kimmins, 1966 : 208.
paludensis Fraser (Aciagrion), 1922. Kimmins, 1966 : 208.

peles Perkins (var. Agrion amaurodytum) , 1899 : 67. LECTOTYPE (J. Hawaiian Islands
(R. C. L. Perkins] / Hawaii, Kau, 4000 ft, ix.i8g5 (Perkins) / Agrion amaurodytum Prk., race
peles Prk. / Agrion amaurodytum var. peles Prk. Type / Agrion amaurodytum var. peles
Perk., <J Lectotype, D. E. Kimmins det. 1969.
Currently placed in Megalagrion.

IQO D. E. KIMMINS

pendleburyi Laidlaw (Ceriagrion), 1931 : 198-200, fig. 6. LECTOTYPE <J. F[ederated]
M[alay] S[tates], Perak, Batang, Jor Camp, 1800 ft, i.vi.ig23 (H. M. Pendlebury) / <$ $ in
cop. / Ceriagrion pendleburyi Types <$ § / Ceriagrion pendleburyi Laidlaw, $ Lectotype,
D. E. Kimmins det. 1969.

The lectotype <J and allotype $ both now lack the apical segments of the abdomen, and the
right fore wing of the lectotype is also missing.

percellulata Calvert (Argia), 1902 : 74-75, pi. iv, figs 5, 27. Holotype <?. [Mexico], Vera
Cruz, Atoyac, v (H. H. S[mith]) / Argia percellulata Calv. <$ COTYPE, P. P. Calvert det. 1901.
Original of f. 27, pi. iv, B. C. A. Neur.

Although this example is labelled COTYPE, it is to be considered as holotype by Calvert's
statement (1908 : xxviii) 'The type specimens of the new forms described in this work are
those which have been figured on the Plates, the male preferably to the female ..."

[perparva McLachlan Mss, in Selys (Ischnura), 1876 : 263-265.

LECTOTYPE <$. Texas occ[identale] / Ischnura perparva McLachl. $ [Selys' writing] /
Ischnura perparva McL., $ Lectotype, D. E. Kimmins det. 1969.]

McLachlan's collection contained two battered syn types (i <J, i $), although they were not
marked Type by him. By courtesy of Dr G. Demoulin, I have seen two more syntypes from
the Selys collection in Brussels, an incomplete <$ and a complete $. Of the two, I have chosen
as Lectotype the Brussels <$, as it is the more mature. Abdominal segments 5-10, one
anterior, one median and both posterior legs are missing.

platystigma Eraser (Ceriagrion), 1951. Kimmins, 1966 : 209.

plana Calvert (var. Argia vivida), 1902 : 96, pi. iv, fig. 58. Holotype (J. [Mexico], Guerrero
Sierra de la Aguas Escondidas, vii (H. H. Smith) / Argia vivida plana Calv., <J COTYPE, P. P.
Calvert det. 1901. Original of f. 58, pi. iv., B.C. A. Neur.

popoluca Calvert (Argia), 1902 : 82, pi. iv, figs 38, 383. Holotype 3. [Mexico], Tabasco,
Teapa, iii (H. H. Smith) / Argia popoluca Calv. <J TYPE, P. P. Calvert det. 1901. Original
of ff. 38, 383, pi. iv, B.C.A. Neur.

praeclarum Eraser (Pseudagrion), 1924. Kimmins, 1966 : 209-210.

prothoracicum Kimmins (Telagrion), 1945 : 187-189, 3 figs. Holotype £. Ecuador,
Intaj / Telagrion prothoracicum Selys, n. sp., $ [Selys' writing] / Telagrion prothoracicum
Kim., (J TYPE.

pruinescens Tillyard (Agriocnemis), 1906 : 191-192, pi. 17, fig. ga. LECTOTYPE $.
N[orth] Queensland], i.[ig]o5 (R. J. Tillyard) / Agriocnemis pruinescens Till., $ TYPE, R.J.T. /
Agriocnemis pruinescens Till., $ Lectotype, D. E. Kimmins det. 1969.
Currently placed in the genus Ischnura.

pseudelongatutn Longfield (Enallagma), 1936 : 475-477, figs 3, 4. Holotype <$. Central
Africa, Uganda, Kigezi Dist., Birunga Mts, swamp in bamboo forest, 7500 ft, 8. iii. 1934
(C. E. Longfield) / Enallagma pseudelongatum sp. nov., Type <J. E in cop. with F, det. Miss

C. Longfield.

quadrigerum Selys (Agrion), 1883 : 136-138. LECTOTYPE <J. Type [McL. label] / Japan
(Pryer) / 145 / Agrion quadrigerum Selys, Japon / Agrion quadrigerum Selys, £ Lectotype,

D. E. Kimmins det. 1969.

The (J lectotype has had the 7-10 segments of the abdomen re-attached with glue at some
time, probably by McLachlan. It differs from the original description in having a forked
black mark at the apex of the 8th segment, as in sieboldi. We have in our collection an
example labelled by Asahina 'Cercion sieboldii Selys = quadrigerum Selys, det. Asahina,
!953' so presumably this is a variable character.

Currently placed as a synonym of Cercion sieboldi (Selys), 1876.

TYPE-SPECIMENS OF ODONATA IN BMNH 191

rajah Laidlaw (Teinobasis), 1912 : 97. LECTOTYPE <J. Sarawak, Limbang, 22.vi.ign
(/. C. Moultori) I Teinobasis rajah $ Type [Laidlaw's writing] / Teinobasis rajah Laidlaw,
$ Lectotype, D. E. Kimmins det. 1969.

I have no information as to the location of the second syntype <J.

[ranauense Schmidt (ssp. Pseudagrion pruinosum), 1934 : 34^, fig- 4?b.

There is in BMNH a^ labelled 'Typus', which was presented in 1934 by Dr F. F. Laidlaw.
It has been labelled 'Type' but I consider that it can be no more than a syntype, probably
only a paratype.]

rarurn Longfield (Ischnuragrion), 1947 : 6-9, figs 1-2. Holotype <J. Angola Miss[ion]
sc[ientifique] suisse, 1932-1933 / Lunda, ix / Ischnuragrion rarum, <J genotype, det. Miss C.
Longfield.

retniger Laidlaw (Atnphicnemis), 1912 : 96-97. Holotype $. Sarawak, Kuala Madalam,
Limbang R., II.V.IQII (/. C. Moultori) / Amphicnemis remiger Laidlaw, $ Type [Laidlaw's
writing] .

rhoadsi Calvert (Argia), 1902 : 92, pi. iv, figs 55, 553. Holotype <£. Mex[ico], Nuevo Leon,
Monterey, 25.^.1899 (Rhoads) / Argia rhoadsi Calv., <$ TYPE, P. P. Calvert det. 1901. Original
of ff. 55, 555, pi. iv, B.C. A. Neur.

rogersi Calvert (Argia), 1901 : 83, pi. iv, figs 40, 403. Holotype <$. Costa Rica, Cach£ (H.
Rogers) / Argia rogersi Calv. £ Type, P. P. Calvert det. 1901. Original of ff. 40, 405, pi. iv,
B.C.A. Neur.

rubricauda Tillyard (Agriocnemis), 1913 : 459-460, pi. 47, fig. 8, pi. 48, figs 17, 18. LECTO-
TYPE (J. N[orth] Queensland], Cookham, xii.[i9]o7 (R. J. Tillyard). Agriocnemis rubri-
cauda Till., <$ Lectotype, D. E. Kimmins det. 1969.

rubroviridis Pinhey (Pseudagrion), 1956 : 23-24, text-figs 2, 3. Holotype <J. N[orthern]
R[hodesia], Livingstone, [Maramba River, near] Victoria Falls, x.1953 (E. Pinhey) / Male and
Female in Copula / Pseudagrion rubroviridis Pinh., HOLOTYPE.

rufocinctum Pinhey (Pseudagrion), 1956 : 24-26, text-figs 3, 4. Holotype Q*. Uganda,
Kamengo, 13^.1952 (E. Pinhey) / Face & thorax above orange-red; sides of thorax green /
Pseudagrion rufocinctum Pinh. 1954, HOLOTYPE.

rufostigma Longfield (Pseudagrion), 1947 : 11-12, fig. 4. Holotype <$. Angola, Miss[ion]
sc[ientifique] suisse, 1932-1933 / Sangeve, ii / Pseudagrion rufostigma Type <$, det. Miss C.
Longfield.

samoensis Fraser (Pseudagrion), 1925. Kimmins, 1966 : 212.

sobrina McLachlan (Telebasis), 1873 : 37-37. Holotype <$. Type [McL. label] / New
Zealand / Telebasis sobrina McL. / Xanthagrion sobrinum McLach. <$ [Selys label].
Currently placed in Xanthocnemis.

somalicum Longfield (Enallagma), 1931 : 277-278, 2 text-figs. Holotype <$. Brit. Somali-
land : Marojeh, 10.54 N., 48.59 E., 2250 ft, 5-7. xi. (C. L. Collenette) / Flying over waterhole /
Enallagma somalicum Longf., Q* Holotype, D. E. Kimmins det. 1969.

[spencei Fraser (Pseudagrion), 1922. Kimmins, 1966 : 214. Holotype $ not traced.]

steelae Kimmins (Aciagrion), 1955 : 109-110, fig. i. Holotype $. N[orthern] Rhodesia,
Lake Bangweulu, near Monfuli, from stream, i.x.i94& (M. Steele) / Aciagrion steelae, <$ Type,
D. E. Kimmins det. 1955.

stellatum Martin (Pseudagrion), 1915 : 46-49. LECTOTYPE <J. British East Africa,
Edge of Kenia Forest, SE Side, 7.11.1911 (/. T. Anderson) / Pseudagrion stellatum R. Martin,
c? type [Martin's writing] / Pseudagrion stellatum Martin <J Lectotype, D. E. Kimmins det.
1969.

Martin states that the description was based on material sent to him by Campion and that
the 6* types are in BMNH and Martin collections. Under these circumstances, the lectotype

ig2 D. E. KIMMINS

should be selected from the BMNH material and I have chosen the one which has for years
carried a BM type label.

Currently placed as a synonym of Ps. bicoerulans Martin.

talamanca Calvert (Argia), 1907 : 371, pi. viii, figs 34, 345. Holotype <J. Costa Rica,
Carillo (Underwood) / Costa Rica, Carillo (C. F. Underwood) / Argia talamanca Calvert, <$
Type. Orig of figs 34, 343, pi. viii, B.C. A. Neur.

tarascana Calvert (Argia), 1902 : 90-91, pi. iv, figs 14, 513, 5153. Holotype Q". Mexico,
[Guanajualo], Acambaro, 3o.iii.i889 (Rhoads) / Argia tarascana Calv., <$ TYPE, P. P. Calvert
det. 1902, B.C. A. Neur. p. 91. Original of pi. iv, f. 5153.

tasmanica Tillyard (race of Ischnura heterosticta), 1913 : 451. LECTOTYPE $. Tas-
[mania], Cressy, 3.i.[i9]o9 (R. J. Tillyard) / Ischnura heterosticta tasmanica Till., <£ Lecto-
type, D. E. Kimmins det., 1969.

The BMNH has three <$ examples of the type-series, from which the above example has been
selected. They were over the label tasmanica in his collection, but bore no determination
labels.

tinctipennis McLachlan (Erythromma), 1894 : 436 ($); 1896 : 373-4 (<?). LECTOTYPE $.
Type [McL. label] / Ta-chien-lu / Erythromma tinctipennis McL. / Pyrrhosoma tinctipenne
McL. / Erythromma tinctipennis McL., $ Lectotype, D. E. Kimmins det. 1969.

Of the three syntype females, two are now in BMNH and one has been chosen above as
Lectotype. The BMNH also has the <$ allotype.

torresiana Tillyard (Ischnura), 1913 : 452-453, pi. 48, figs 5-6. LECTOTYPE <$. N[orth]
Queensland], Cooktown, i.[i9]o8 (R. J. Tillyard) / Ischnura torresiana Till., <J TYPE, R. J. T. /
Ischnura torresiana Till., <$ Lectotype, D. E. Kimmins det. 1969.

triangularis Laidlaw (Pericnemis), 1913^ : 248. Holotype $. Borneo, Bettotan, i3.vii.
[ig]27 (C. Boden Kloss) / Pericnemis triangularis n. sp. $ type [Laidlaw's writing].
The holotype now lacks abdominal segments 8-10.

trifoliata Fraser (Argia), 1946. Kimmins, 1966 : 216.

truncatipenne Calvert (Anisagrion), 1902 : 106, pi. v, fig. 17. Holotype <$ (incomplete).
Guatemala, El Reposo [Pacific coast-region] (Champion) / Anisagrion truncatipenne Calv.
TYPE, P. P. Calvert det. 1902. B.C. A. Neur., p. 106, Original of pi. v, fig. 17.

ultneca Calvert (Argia), 1901 : 80, pi. iv, figs 9, 34, 343, 34!. Holotype <^. Vera Cruz,
Atoyac, v (H. H. S[mith]) / Argia ulmeca Calv. COTYPE <$. P. P. Calvert det. 1901. Original
of ff. 34, 343, pi. iv, B.C. A. Neur.

umbriaca Fraser (Argia) 1946. Kimmins, 1966 : 217.

underwoodi Calvert (Argia), 1907 : 370, pi. 8, figs 36, 37, 373. Holotype <$. Costa Rica,
Carillo (Underwood) / Costa Rica, Carillo (C. F. Underwood) / Argia underwoodi Calv. <$ TYPE,
P. P. Calvert, det. 1907, B.C. A. Neur. p. 370. Original of pi. viii, ff. 37, 373.

vagabundum Perkins (Agrion), 1899 : 75. LECTOTYPE <$. Hawaiian Islands (R. C. L.
Perkins) / Kauai, Lihue (Perkins) / Agrion vagabundum Prk. / Agrion vagabundum Prk.,
Type / Agrion vagabundum Prk., $ Lectotype, D. E. Kimmins det. 1969.
Currently placed in Megalagrion.

vansomereni Pinhey (Enallagma), 1956 : 26-27, text-fig. 4. Holotype <$. Uganda, Acholi,
Paimal, iv.i952 (T. H. E. Jackson) / Enallagma vansomereni Pinhey, 1954. HOLOTYPE.

vansomereni Pinhey (Pseudagrion), 1961 : pi. 2, figs 2, 9. Holotype <$. Uganda, Aswa
R[iver], Karamoja, iii.[i9]52 (T. H. E.Jackson) / Pseudagrion vansomereni Pinh. [<$] Holotype.

varralli Fraser (Mortonagrion), 1920. Kimmins, 1966 : 217.
versicolor Fraser (Telebasis), 1946. Kimmins, 1966 : 217.
victoria Fraser (Agriocnemis), 1928. Kimmins, 1966 : 217.

TYPE-SPECIMENS OF ODONATA IN BMNH 193

violacea Fraser (Coenagrion), 1924. Kimmins, 1966 : 218.

waianeanum Perkins (var. of Agrion amaurody turn) , 1899 : 67. LECTOTYPE Q*. Hawaiian

Islands (R. C. L. Perkins) / Oahu, Waianae Mts, 2000 ft, iv.i892 (Perkins) / Agrion amauro-

dytum var. waianeanum Prk., Type / A. amaurodytum Prk., race waianaeanum / Agrion

amaurodytum var. waianeanum Prk., $ Lectotype, D. E. Kimmins det. 1969.

Currently placed as Megalagrion.
wallacei Campion (Teinobasis), 1924 : 613-614, fig. i. $ Holotype. N[ew] Guin[ea]

(Wallace) / N / Teinobasis wallacei Campion, £ Holotype, det. D. E. Kimmins, i8.ix.i933.
whellani Longfield (Ceriagrion), 1952 : 42-43, fig. 2. Holotype <$. S. Rhodesia, Dept.

Agric., Melsetter Distr., Chibudzana R., i6.xi.i948 / J. A. Whellan, collector / Ceriagrion

whellani sp. n. Type <$, det. Miss C. Longfield / figured.
whellani Pinhey (Pseudagrion), 1956 : 18-22, text-figs 1,2. Holotype <$. Uganda, Acholi,

Madi Opei, iii.i952 (T. H. E. Jackson) / Male and Female in copula / Pseudagrion whellani

Pinh., 1954, HOLOTYPE.

williamsoni Fraser (Pseudagrion), 1922. Kimmins, 1966 : 219.
zelandica McLachlan (Telebasis), 1873 : 35-36. LECTOTYPE 6"- Type [McL. label] /

New Zealand / Xanthagrion Zelandicum Selys Q* / Telebasis zelandica McL. / Telebasis

zelandica McL., <$ Lectotype, D. E. Kimmins det. 1969.
Currently placed in the genus Xanthocnemis.

PSEUDOSTIGMATIDAE

abnorme McLachlan (Anomisma), 1877 : 87. Holotype 3. Type [McL. label] / E. Peru? /

Rio Napo / Anomisma abnorme, McL.
buckleyi McLachlan (Mecistogaster), 1881 : 32. LECTOTYPE <J. Type [McL. label] /

Ecuador, R. Bobonaza (Buckley) / Mecistogaster buckleyi McL., / Mecistogaster buckleyi

McL., (J Lectotype, D. E. Kimmins det. 1968.

calcipennis Fraser (Microstigma), 1946. Kimmins, 1966 : 185.

sincerus McLachlan (race of Mecistogaster jocaste Hagen), 1877 : 88; 1881 : 32. LECTO-
TYPE $. Type [McL. label] / Pebas, Amazons (Hauxwell) / 13 / Mecistogaster sincerus
McLachl. $ / Mecistogaster sincerus McL. 9 Lectotype, D. E. Kimmins det. 1968.
Currently placed as a var. of M. jocaste Hagen.

terminatum McLachlan (Microstigma), 1877 : 87. Holotype $. Type [McL. label] / E.
Peru? / Rio Napo / Microstigma terminatum McL. / Anomisma abnorme McL. $.

Synonymized with Anomisma abnorme McL., by McLachlan, 1881 : 31 as 'first reviser'.

PERILESTIDAE

bispinus Kimmins (Perilestes), 1958 : 353-354, fig. 3. Holotype $. [Brazil, Rio Negro],
Thomar / Perilestes bispinus Kim., <$ Type, D. E. Kimmins det., 1958.

risi Morton (Chorismagrion), 1914 : 170-172, pi. 9, figs 1-3. Holotype cj. Cape York, N.
Queensland / Nov. gen., nov. spec., det. Dr F. Ris / Chorismagrion risi Mort. TYPE [in Eraser's
writing] / One pair of wings in Ris collection, teste F. C. Fraser [D.E.K.].

This example was presented to BMNH in 1953 by F. C. Fraser, with other type material.

SYNLESTIDAE

albicauda Tillyard (Synlestes), 19130 : 238-241, pi. 15, figs 10-11. LECTOTYPE <$.

Queensland], [Mount] Tambourine, 2. i. [19] 13, in cop. (R. J. Tillyard) / Synlestes albicauda
Till., TYPE <£, RJ.T. / Synlestes albicauda Till., <$ Lectotype, D. E. Kimmins det. 1968.

Tillyard ( : 240) specified a pair in cop. as 'Types <$, ?', but did not indicate which was the
holotype ; therefore the above lectotype designation has been made. Currently placed in the
genus Episynlestes.

194 D. E. KIMMINS

nigrescens Tillyard (ssp. of Synlestes weyersi), 1917 : 472-473, fig. 8b. LECTOTYPE <J.

N[ew] S[outh] W[ales], Illawarra, u.ii.[i9]n (JR. J. Tillyard) j Synlestes weyersi nigrescens

Till., TYPE (J, R.J.T. / Synlestes weyersi nigrescens Till., $ Lectotype, D. E. Kimmins det.

1968.

Fraser (1960 : 26) treats this taxon as a species.
selysi Tillyard (Synlestes), 1917 : 473-475, pi. 13, text-figs 7-8. LECTOTYPE £. N[ew]

S[outh] W[ales], Hornsby, 3i.iii.[i9]i7 (R. J. Tillyard) / Synlestes selysi Till., TYPE 6*, R.J.T. /

Synlestes selysi Till., <J Lectotype, D. E. Kimmins det. 1968.
tropicus Tillyard (Synlestes), 1917 : 475, text-figs 8d, 9. Holotype $. N. Queensland],

Kuranda, 2o.xii.[i9]i2 (F. P. Dodd) / Synlestes tropica [sic] Till., TYPE <J. R.J.T.

LESTIDAE

albicauda (Selys MS) McLachlan (Lestes), 1896 : 23-24. LECTOTYPE <$. 'N'[New Guinea]
/ Lestes albicauda Selys n. sp. <$ / Lestes albicauda McL. / Lestes albicauda McL., $ Lectotype,
D. E. Kimmins det. 1968.

Lieftinck, 1951 : 6 designated a 'holotype', but as a holotype can only be designated in
the original publication, I have here designated the same example as Lectotype. Currently
placed in Lestes (Indolestes).

alleni Tillyard (Austrolestes), 1913 : 425, pi. 45, figs 11-12. LECTOTYPE <?. N. Q[ueens-
land], Cairns, ix.[i9]o5 (E. Allen) / Lestes Alleni Till., <$ TYPE, R.J.T., with in pencil 'albicauda
McL.' / Lestes alleni Till., $ Lectotype, D. E. Kimmins det. 1968.

Currently placed in the subgenus Indolestes. The pencil annotation on Tillyard's label
may be an error for Austrolestes albicauda Ris, nee McL., which has been placed as a synonym
of A . alleni Tillyard.

aridus Tillyard (Lestes), 19080 : 762-764, pi. 42, figs 4-5. LECTOTYPE <?. [Australia],
N[orth] Territory], Tennant's Creek, i.[i9]o6 (/. Field) / Lestes aridus Till., $ TYPE, R.J.T. /
Lestes aridus Till., <$ Lectotype, D. E. Kimmins det. 1968.

Currently placed as Lestes (Austrolestes}.

aruanus Lieftinck (Lestes), 1951 : 8-9, fig. 4. Holotype <$. Aru / Type [McL. label] / Lestes
albicauda $, Selys, sp. n., Aru [in Selys' writing] / Lestes albicauda McL. [with paratype label
attached] / Lestes aruanus Lieftinck, Holotype, det. M. A. Lieftinck, 1947.

The holotype lacks its head. Currently placed in the subgenus Indolestes.
californica (Selys MS) McLachlan (Archilestes), 1895 : 20-21. Holotype <£. Type [McL.
label] / California ([Henry] Edwards) / Archilestes californicus Selys <J n. sp. [Selys' writing] /
Archilestes californica McL.

The locality and collector's labels, being very small, have been assembled as one label.
cheesmanae Kimmins (Austrolestes), 1936 : 69-70, text-figs 1-4. Holotype $. New
Hebrides, Erromanga, vii.i93o (L. E. Cheesman) / Austrolestes cheesmanae sp. n. <$.

Currently placed as Lestes (Austrolestes).

colensonis White (Agrion), 1846 : pi. 6, fig. 3; Selys, 1862 : 328-329; White, 1874 : 25.
LECTOTYPE $. New Zealand, on reverse a BM register number [i8]45-6i / Agrion
colensonis White, $ Lectotype, D. E. Kimmins det. 1968.

The first reference is to a figure, published with a name in 1846; Selys gave a description
(1862) under the name Lestes colensonis Adam White and in 1874 White gave references to
the original publication and that of Selys and stated that the type was in the BMNH.
Currently placed in the genus Austrolestes.

davenporti Fraser (Ceylonolestes), 1930. Kimmins, 1966 : 188.

decipiens Kirby (Lestes), 1894 : 565-566. LECTOTYPE <?. Mahagany, 2o.ix.[i8]gi /
Ceylon, Yerbury Coll. / decipiens type [WFK] / Lestes decipiens Kirby, $ Lectotype, D. E.
Kimmins det. 1968.

Currently placed as a subspecies of Paralestes praemorsa (Selys).

TYPE-SPECIMENS OF ODONATA IN BMNH 195

disartnatus Fraser (Lestes), 1961. Kimmins, 1966 : 189.
dorothea Fraser (Lestes), 1924. Kimmins, 1966 : 190.

excelsa Fraser (Orolestes), 1933. Kimmins, 1966 : 191.

At the time when this type deposited in the BMNH (1953), Fraser informed me that one
pair of wings had been presented to the Ann Arbor Museum, Michigan. During the prepara-
tion of this paper however, this pair of wings was discovered in his collection, and they have
now been re-attached to the type.

Helena Fraser (Indolestes), 1922. Kimmins, 1966 : 195.
indica Fraser (Indolestes), 1922. Kimmins, 1966 : 197.

insularis Tillyard (Austrolestes), 1913 : 425-426, pi. 45, figs 13-14. Holotype £. [Queens-
land, Torres Straits], Banks Is., i6.ii.[i9]io (H. Elgner) / Austrolestes insularis Till., R.J.T.,
c? TYPE.

Currently placed as Lestes (Indolestes).

nigriceps Fraser (Lestes), 1924. Kimmins, 1966 : 206.

obscurus Kirby (Lestes), 1898 : 245. Holotype <$. [Transvaal], Barberton / obscurus
patype [sic] Kirby [in Distant's writing] / Lestes obscurus Kirby, <$ Holotype, D. E. Kimmins
det. 1968.

Currently placed as a synonym of Lestes plagiatus Burm.

paludosus Tillyard (Lestes), 1906 : 181-182, pi. 17, figs 3a, 3b. LECTOTYPE <$. N.
Queensland], Townsville, i.[i9]o5 (R. J. Tillyard) / Lestes paludosus Till., $ TYPE, R.J.T. /
Lestes paludosus Till., $ Lectotype, D. E. Kimmins det. 1968.
Currently placed as a synonym of Lestes concinnus Hag.

psyche Selys (Lestes), 1962 : 329.

In the BMNH there is a damaged $ syntype (ex McLachlan collection), labelled Type by
him, which is possibly the $ referred to in the original description.

pulcherritna Fraser (Ceylonicolestes), 1924. Kimmins, 1966 : 210.

quercifolia Selys (Lestes), 1878 : 318; 1891 : 64. LECTOTYPE $. Type [McL. label] /
Menado / Lestes quercifolia Selys $ (race de Praemorsa) / Lestes quercifolia Selys, $ Lecto-
type, D. E. Kimmins det. 1968.

Whether the Menado form of praemorsa is worthy of a name, I will leave for others to
decide; the name does not appear to have come into general use. L. praemorsa Hagen is
currently placed in Paralestes.

selysi McLachlan (Orolestes), 1895 : 22-23. LECTOTYPE 6*. Type [McL. label] / Dar-
jeeling / Orolestes Selysi McL. / Orolestes selysi McL., 6* Lectotype, D. E. Kimmins det. 1968.
The lectotype now lacks the right hind wing. Fraser (1929 : 838-839, fig. 2) refers to the
two examples in the McLachlan coll., and states that it has not been re-discovered at Darjeeling.
He also gives a photograph purporting to be of the wings of this species, which does not
appear to have been taken from either of the two McLachlan examples. In 1923 : 25-27,
Fraser repeats his statement but gives wing figures from a different example, collected by
Dr A. Kerr in Laos, 1932.

sikkima Fraser (Lestes), 1929. Kimmins, 1966 : 213.

simulatrix McLachlan (Lestes), 1895 125-27. Holotype^. Type [McL. label] / Madagascar /
Lestes simulatrix McL.

Currently placed in the genus Paralestes.

tenuissimus Tillyard (Austrolestes), 1906 : 179-181, pi. 17, figs 2a, 2b. LECTOTYPE <$.
N. Queensland], Cairns, i.[i9]o5 (R. J. Tillyard) / Lestes tenuissimus Till., <$ TYPE, R.J.T. /
Lestes tenuissimus Till., D. E. Kimmins det. 1968.
Currently placed in Lestes (Indolestes).

IQ6 D. E. KIMMINS

tridens McLachlan (Lestes), 1895 : 24-25. Holotype <$. Type [McL. label] / Delagoa Bay /
Lestes tridens McL.

Currently placed in the genus Paralestes.

veronica Fraser (Indolestes), 1924. Kimmins, 1966 : 217.

unicolor McLachlan (Lestes), 1895 : 27-28. LECTOTYPE <J. Type [McL. label] / Tama-
tave, Madagascar / Lestes unicolor McL. / Lestes unicolor McL., <$ Lectotype, D. E. Kimmins
det. 1968.

The lectotype has part of the left fore wing attached to a card.

wallacei Kirby (Lestes), 1889 : 302-303. Holotype <j>. Sar[awak], Jo. / (Wallace) / Wallacei
type [WFK's writing] / Lestes wallacei Kirby, $ Holotype. D. E. Kimmins det. 1968.
Now placed in the genus Orolestes.

MEGAPODAGRIIDAE

aequatoriale Selys (Heteragrion), 1886 : 63-64. LECTOTYPE <J. Type [McL. label] /
Bogota / Heteragrion aequatoriale Selys <$, Bogota / Heteragrion aequatoriale Selys, $
Lectotype, D. E. Kimmins det. 1968.

McLachlan's marked type-series includes two from New Granada, which are not included
in the original description, although they bear determination labels in Selys' writing.

albescens Tillyard (ssp. Argiolestes griseus), 1913 : 414-415. LECTOTYPE <$. [South]
Queensland, Stradbroke Id., 2.x.[i9]n (H. Hacker) / Argiolestes griseus albescens $ TYPE,
R.J.T. / Argiolestes griseus albescens Till., £ Lectotype, D. E. Kimmins det. 1968.

alpinus Tillyard (Argiolestes), 1913 : 417-419, pi. 44, figs 7, 8. LECTOTYPE (J. N[ew]
S[outh] W[ales], Ebor, [4600-4800 ft], 3.i.[i9]i2 (R. J. Tillyard) / Argiolestes alpinus $ TYPE.
R.J.T. / Argiolestes alpinus Till., <J Lectotype, D. E. Kimmins det. 1968.

angustipennis Selys (race of Heteragrion aequatoriale), 1886 : 64-65. LECTOTYPE <$.
Ecuador / Heteragrion angustipennis Selys, <$ Ecuador / Heteragrion angustipennis Selys,
6* Lectotype, D. E. Kimmins det. 1968.

Selys described the <J as lacking the three last segments of the abdomen, but since then
these segments have been glued on, possibly by McLachlan.

aureus Tillyard (Argiolestes), 1906 : 178-179, pi. 17, figs la, ib. LECTOTYPE <J. N.
Queensland], Kuranda, xii.[i9]o4 (Jf?. /. Tillyard) / Argiolestes aurea [sic] Till., $ TYPE,
R.J.T. / Argiolestes aureus Till., <$ Lectotype, D. E. Kimmins det., 1968.

carrillica Calvert (Philogenia), 1907 : 356-357, pi. 7, figs 6, 7, 12. Holotype^. Costa Rica,
Carrillo (Underwood) / Philogenia carrillica Calv., P. P. Calvert det. 1907. B.C. A. Neur.,
p. 356. TYPE. Original of PI. VII, figs 6, 7, 12.

championi Calvert (Philogenia), 1901 : 61, pi. 5, figs 3, 4. Holotype $. [Panama],
V[olcan] de Chiriqui, below 4000 ft, ([G. C.] Champion) / Gizzard removed / Original of figs 3,
4, 4a, PL V, B.C. A. Neur., Philogenia championi Calv., $ TYPE, P. P. Calvert det. 1901.

chrysoides Tillyard (Argiolestes), 19130 : 237-238, pi. 15, figs 8, 9. Holotype (J. Q[ueens-
land, Blackall Ranges], Montville, 6.x.[ig]i2 ([^4.] /. Turner) / Argiolestes chrysoides TYPE $,
R.J.T.

The locality label incorrectly gives the collector's name as E. J. Turner.

coartans Lieftinck (Argiolestes), 1956 : 81-84, n8s I0> J4> T5- Holotype <$. N. Dutch New
Guinea, Waigeu [Island], Camp Nok, 2500 ft, iv.i938 (L. E. Cheesman) / Argiolestes coartans
Lieftinck, Holotype, det. M. A. Lieftinck, 1955.

curtum Selys (Megapodagrion), 1886 : 46, 47. Holotype $. Type [McL. label] / Ecuador,
R. Bobonaza (Buckley) / Megapodagrion curtum Selys, <£•

The original description has the generic and specific names misspelt 'Megapodagriom
curtue'.

TYPE-SPECIMENS OF ODONATA IN BMNH 197

duodecimo Calvert (Paraphlebia), 1901 : 60, 61, pi. 5, fig. 2. Holotype <$. [Guatemala],
Vera Paz, Purula ([G. C.] Champion) / Paraphlebia duodecima Calv., P. P. Calvert det. 1901.
TYPE. Original of PI. V, fig. 2, B.C. A. Neur.

eboracus Tillyard (ssp. Argiolestes griseus), 1913 : 413. LECTOTYPE $. N[ew] S[outh]
W[ales], Ebor, i8.xii.[i9]n (R. J. Tillyard) / Argiolestes griseus eboracus Till., <$ TYPE,
R.J.T. / Argiolestes griseus eboracus Till., 5" Lectotype, D. E. Kimmins det. 1968.

ephippiatus Lieftinck (Argiolestes), 1956 : 97-98, figs 35, 36. Holotype <$. [N.] Papua,
Kokoda, 1200 ft, ix.i933 (L. E. Cheesmari) / Argiolestes ephippiatus Lieft., HOLOTYPE, det.
M. A. Lieftinck, 1955.

erythrogastrum Selys (Heteragrion), 1886 : 61-62. LECTOTYPE <$. Type [McL. label] /
Chiriqui / Heteragrion erythrogastrum Selys, Q* Chiriqui [Selys' writing] / Heteragrion
erythrogastrum Selys, $ Lectotype, D. E. Kimmins det. 1968.

I do not regard Calvert's statement (1901 : 65) concerning the incomplete example from
McLachlan's collection (labelled 'Type') and studied by Calvert, as a designation of lectotype,
since McLachlan so labelled all the type-series in his collection. I do not know of any later
type-designation and have therefore designated the above complete example as Lectotype.

flaviceps Eraser (Burmargiolestes), 1933. Kimmins, 1966 : 191.

fontanus Tillyard (Argiolestes), 1913 : 417-419, pi. 44, figs 9-10. LECTOTYPE <?. N[ew]

S[outh] W[ales], Dorrigo, n.xii[i9]n (R. /. Tillyard) / Argiolestes fontanus <$ TYPE, R.J.T. /
Argiolestes fontanus Till., $ Lectotype, D. E. Kimmins det. 1968.

intermedius Tillyard (race of Argiolestes griseus), 1913 : 412-413. LECTOTYPE Q*.

Vic[toria], Alexandra, xii.[i9]o6 (R. J. Tillyard) / Argiolestes griseus intermedius Till.,
<J TYPE, R.J.T. / Argiolestes griseus intermedius Till., Q* Lectotype, D. E. Kimmins det. 1968.

leucorrhinum Selys (Heteropodagrion Mesagrion), 1885 : cxliv; 1886 : 50-51. Holotype

o*, Type [McL. label] / Bogota / Mesagrion leucorrhinum $ Selys, Bogota, with on under-
side, Heteropodagrion [deleted] leucorrhinum $ Selys, Bogota.
maclachlani Selys (Allolestes), 1869 : 97-98. Holotype ?. Type [McL. label] / Seychelles

(Wright) I Allolestes maclachlani $ Selys.

Legs glued to card, first six segments of abdomen only present, abdomen glued on upside

down.
malgassica Kirby (Tatocnemis), 1889 : 302, 2 text-figs. Holotype Q\ Madagascar,

Betsileo (Deans Cowan)] / Tatocnemis malgassica type [WFK's writing].

An additional locality label, giving full details, has been added.
tnicrostigma Lieftinck (Argiolestes), 1956 : 99-101, figs 40, 41. Holotype <$. [C.] Papua,

Mafulu, 4000 ft, 1.1934 (•£•• E- Cheesman) / Argiolestes microstigma Lieft., HOLOTYPE, Det.

M. A. Lieftinck, 1955.
minimus Tillyard (Argiolestes), 1908 : 735-737, pi. 35, figs n, 12. LECTOTYPE 6*. W.

A[ustralia], Bridgetown, i.[i9]o7 (R. J. Tillyard) / Argiolestes minima [sic] Till., $ TYPE,

R.J.T. / Argiolestes minimus Till., $ Lectotype, D. E. Kimmins det. 1968.
nigra Kimmins (Bornargiolestes), 1936 : 87-88, figs 9, A-D. Holotype <J. Sarawak, Mt

Dulit, 3000 ft, 24. x. 1932 (Native collector) / Bornargiolestes nigra ^ Type, Kimmins, det.

D. E. Kimmins.
nobilis Tillyard (ssp. Argiolestes icteromelas) , 1913 : 410. LECTOTYPE <J. N[ew]

S[outh] W[ales], Ebor, [4000-5000 ft], 6.i.[i9]i2 (R. J. Tillyard) / Argiolestes icteromelas

nobilis Till., Q* TYPE, R.T.J. / Argiolestes icteromelas nobilis Till., Q* Lectotype, D. E. Kimmins

det. 1968.

pallidistigma Eraser (Calilestes), 1926. Kimmins, 1966 : 208.
prothoracalis Lieftinck (Argiolestes), 1956 : 98-99, figs 33, 34. Holotype <$. [N.] Papua,

Kokoda, 1200 ft, vi.i933 (L. E. Cheesman) / Argiolestes prothoracalis Lieft. HOLOTYPE,

Det. M. A. Lieftinck, 1955.

igg D. E. KIMMINS

quint a Calvert (Paraphlebia) , 1901 : 60, pi. 5, fig. i. Holotype ^. Guatemala, [Vera Paz],

Panima (\G. C.] Champion) / Paraphlebia quinta Calvert, P. P. Calvert det. 1901. TYPE.

Original of pi. v, fig. i, B.C. A. Neur.
raphaella Selys (Philogenia), 1886 : 37. Holotype 6*. Type [McL. label] / Bogota / Philo-

genia raphaella Selys.
roseonotata Selys (Podopteryx), 1871 : 415-416. Holotype $. Aru Isl[and, (Wallace)'] /

Podopteryx roseonatatus de Selys (unique) .
setigerum Selys (Megapodagrion), 1886 : 42-43. Holotype^. Type (McL. label] / Ecuador,

Intaj / Megapodagrion setigerum <$ Selys.

The type now lacks its head.
tennis Tillyard (ssp. Argiolestes griseus), 1913 : 413-414. Holotype <$. N[ew] S[outh]

W[ales], [Hornsby], N[ear] Sydney, i8.xii.[i9]og (R. J. Tillyard) / Argiolestes griseus tenuis

Till., c? TYPE, R.J.T.
tricellulare Calvert (Heteragrion), 1901 : 63, pi. 5, fig. 5. Holotype <$. [Guatemala],

S[an] Geronimo, 3000 ft, ([G. C.] Champion) / D taken with C / Heteragrion tricellulare Calv.

P. P. Calvert det. 1901. B.C. A. Neur., p. 63. TYPE. Original of pi. 5, fig. 3.

Example C (taken with above) is not in our collection.
trinervis Selys (Neurolestes), 1885 : cxliv; 1886 : 71-72. LECTOTYPE <J. Type [McL.

label] / Old Calabar, R[utherford] 73 / Nevrolestes trinervis Selys J, Old Calabar / Neurolestes

trinervis Selys, $ Lectotype, D. E. Kimmins det. 1968.

Selys (1886 : 72) gives the locality as 'Old Thai', probably a printer's error for 'Old

Calabar'.

PSEUDOLESTIDAE

inopinata McLachlan (Thaumatoneura), 1897 : 131. Holotype <$ (incomplete). [No
locality data] / Thaumatoneura inopinata McL. (Type).

The specimen was purchased by McLachlan, without locality data and with abdominal
segments 5-10 lacking.

mirabilis Kirby (Pseudolestes), 1900 : 538-539, pi. 12, fig. 3. LECTOTYPE <$. Hainan,
5-fingered Mt. [/. Whitehead] / Pseudolestes mirabilis <$ Kirb[y] (type) / Pseudolestes mirabilis
<$ Kirby <J Lectotype, D. E. Kimmins det. 1968.

The lectotype lacks segments 7-10 of abdomen. The name J. T. Thomasson on the label
is that of the donor; the material was collected by J. Whitehead. There is a seventh ex-
ample, by the same collector, in the BMNH from the Crowley Bequest. This is complete, but
cannot be included in the type-series. In view of this additional example, I have labelled the
paralectotypes.

pellucida Calvert (Thaumatoneura), 1094 : 216; 1908 : 355. LECTOTYPE <$. Costa
Rica, Carrillo (Underwood) / Thaumatoneura pellucida Calv., P. P. Calvert det. 1907, B.C. A.
Neur., p. 355. COTYPE / Thaumatoneura pellucida Calv., $ Lectotype, D. E. Kimmins det.
1968.

In describing this species, Calvert states 'Mr F. D. Godman has recently acquired for the
Biologia Centrali- Americana six males of this genus, taken at Carrillo, Costa Rica, by Mr F. C.
Underwood. Four are T. inopinata McLachlan, two are new.' The specimen in our collec-
tion is part of the BCA collection, which was acquired by the BMNH in 191 1. It was labelled
COTYPE by Calvert, but one would assume that Godman would have had the type in the
BCA collection. I do not know of any earlier selection of type for this species and I have
therefore designated the BMNH example as Lectotype.

LESTOIDEIDAE

conjuncta Tillyard (Lestoidea), 1913 : 428-429, pi. 46, figs 1-2, pi. 47, fig. i. Holotype £.
[N. Queensland], Kuranda, i.[ig]o8 (E. Allen) / Lestoidea conjuncta Till., <$ TYPE, R.J.T.

TYPE-SPECIMENS OF ODONATA IN BMNH 199

POLYTHORIDAE

adjuncta Fraser (ssp. Polythore derivata), 1946. Kimmins, 1966 : 177.

aequatorialis Selys (race of Thore picta), 1873 : 500. Holotype <j>. Type [McL. label) /
Ecuador / Thore aequatorialis $ de Selys, race de picta?

Currently placed in the genus Polythore. The type lacks most of the abdomen.

alcyone Selys (Cora), 1873 : 503. Lectotype $ (Montgomery, 1967 : 142). Type [McL.
label] / Bogota / Cora alcyone Selys <$ [Selys' writing] / Lectotype, Montgomery, 1967, D. E.
Kimmins.

The penis has been removed for study.

ambigua Fraser (ssp. Polythore derivata), 1946. Kimmins, 1966 : 177.

beata McLachlan (Thore), 1869 : 28. Lectotype $ (Montgomery, 1967 : 151). Type [McL.
label] / Up[per] Amaz[ons], Pebas (Hauxwell) j Thore beata McL. / Lectotype, Montgomery,
1967, D. E. Kimmins.

Currently placed in the genus Polythore. Montgomery lists Type <$ and paratypes in
BM(NH), and in effect selected the example bearing a type label as the lectotype.

boliviano McLachlan (Thore), 1878 : 89. Holotype <J. Type [McL. label] / Bolivia, Chairo /
Thore boliviana McL. [McL. label] / boliviana McLach. £ [Selys' label].
Currently placed in the genus Polythore.

chirripa Calvert (Cora), 1907 : 348. Lectotype <$ (Montgomery, 1967 : 143). Costa Rica,
Carillo (Underwood] / Cora chirripa Calvert <J Cotype. P. P. Calvert, det. 1907, B.C. A.
Neur., p. 348 / Lectotype, Montgomery, 1967, D. E. Kimmins.

concinna McLachlan (Thore), 1881 : 28. LECTOTYPE <?. Type [McL. label] / Ecuador,
R. Bobonaza (Buckley) / Thore concinna McL. / Thore concinna McL., <$ Lectotype, D. E.
Kimmins det. 1968.

Currently placed in the genus Polythore.

cyane Selys (Cora), 1853 : 71, 72. Lectotype 3 (Montgomery, 1967 : 144), Venezuela / Cora
cyane de Selys <$ [Selys' writing] / Cora cyane Selys, Lectotype, Montgomery, 1967.

derivata McLachlan (Thore), 1881 : 27. LECTOTYPE 3. Type [McL. label] / Ecuador,
R. Bobonaza (Buckley) / Thore derivata McL. / Thore derivata McL., 3 Lectotype, D. E.
Kimmins det. 1968.

Currently placed in the genus Polythore.

dualis McLachlan (Cora), 1878 : 90. LECTOTYPE 3. Type [McL. label] / Ecuador, Intaj /
Cora dualis McL. [McL. writing] / Cora dualis McL., 3 Lectotype, D. E. Kimmins det. 1968.
The type-series was restricted by Montgomery (1967 : 144) to 'the types and paratype 3
in the Br. Mus.' The penis of the lectotype has been removed for study.

jocosa McLachlan (Cora), 1881 : 30. Holotype <£. Type [McL. label] / Ecuador, Rio Bobon-
aza (Buckley) / Cora jocosa McL.

Currently placed in the genus Josocora. The penis of the type has been removed for study.

mirabilis McLachlan (Euthore), 1878 : 87. LECTOTYPE 3. Type [McL. label] / Ecuador,
Intaj, / Euthore mirabilis McL. / Euthore mirabilis McLachlan, 3 Lectotype, D. E. Kimmins
det. 1968.

Montgomery (1967 : 148) restricted the choice of lectotype to the series in the BM(NH).

munda McLachlan (Cora), 1878 : 91. Lectotype 3 (Montgomery, 1967 : 146). Type [McL.
label] / Ecuador, Intaj [C. Buckley'] / Cora munda McL. / Lectotype 3, Montgomery, 1967,
D. E. Kimmins.

I consider that Montgomery, by his statement 'Type: 3- Intaj, Ecuador .... In Br. Mus.,
ex coll. McLachlan' in effect designated the BM(NH) example (which then carried a holotype
label) as the Lectotype.

200 D. E. KIMMINS

tnutata McLachlan (Thore), 1881 : 29. LECTOTYPE <J. Type [McL.] / Ecuador, R.

Bobonaza (Buckley] / Thore mutata McL. / Thore mutata McL., <$ Lectotype, D. E. Kimmins

det. 1968.

Currently placed in the genus Polythore.

originata Fraser (ssp. Polythore derivata), 1946. Kimmins, 1966 : 208.
plagiata Selys (Euthore), 1873 : 501. Holotype <j>. Type [McL. label] / Rio Negro / 206 /

Euthore plagiata Selys $.
pulchella Kirby (Sapho), 1889 : 300. LECTOTYPE <J. Colombia [WFK] / S[apho]

pulchella type [WFK] / Sapho pulchella Kirby, $ Lectotype, D. E. Kimmins det. 1968.
Currently placed in Polythore. The locality of this species was originally given as

'Cameroons' but the label was later changed to 'Colombia' by Kirby. The collection from

which it was described included material from both countries and when Kirby realized that

he had mistakenly placed the species in an African genus, he must have changed the locality

label.
saundersi Selys (Thore), 1853 : 70. LECTOTYPE <J. Amaz[ons] / 54 / Thore saundersii

Selys / Thore saundersi Selys, $ Lectotype, D. E. Kimmins det. 1968.

Currently placed in the genus Polythore, as a synonym of Polythore picta (Rambur).
scintillans McLachlan (Chalcopteryx), 1870 : 169-170. LECTOTYPE #. St. Paulo /

Chalcopteryx scintillans McL., <$ Lectotype, D. E. Kimmins det. 1968.

Montgomery (1967 : 142) makes two slight errors. McLachlan states that he examined 6,

not 9, examples, of which 5 are now in the BM(NH). The register number of the specimens

is 65/3, not 115/3, a mis-reading of the badly written figure 6.
skinneri Calvert (Cora), 1907 : 349. Holotype <J. Costa Rica, Tablazo, 1300-1600 m,

vi.[i9]o5 (Biolley) / Cora skinneri Calv. <J TYPE, P. P. Calvert det. 1907. B.C.A. Neur.

P- 349-

The penis of the type has been removed for study.

terminalis McLachlan (Cora), 1878 : 92. LECTOTYPE <J. Type [McL. label] / Bolivia,
Unduavi / Cora terminalis McL. [McL. label] / Cora terminalis McL. [Selys' label] / Cora
terminalis McL., <J Lectotype, D. E. Kimmins det. 1968.

The penis of the lectotype has been removed for study. The allotype $ lacks most of its
abdomen.

terminate Fraser (ssp. Polythore derivata), 1946. Kimmins, 1966 : 216.
victoria McLachlan (Thore), 1869 : 28. Holotype <J. Type [McL. label] / Bolivia / Thore
victoria 6* McLachl. [Selys' label].

Currently placed in the genus Polythore.

REFERENCES

BARNARD, K. H. 1937. Notes on dragonflies (Odonata) of the S. W. Cape, with descriptions
of the nymphs and of new species. Ann. S. Afr. Mus. 32 : 169-260, 32 text-figs.

CALVERT, P. P. 1901-08. Odonata. Biologia cent.-am. 50 : 17-420, pis 2-10.
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1931. The generic characters and the species of Palaemnema (Odonata : Agrionidae).

Trans, am. ent. Soc. 57 : i-m, 21 pis, 2 text-figs.

CAMPION, H. 1914. Three new species of Ceriagrion from West Africa (Order Odonata).
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— — 1915. A new agrionine dragonfly from Northern Australia. Ann. Mag. nat. Hist.

(8) 16 : 105-108, I text-fig.

1921. Odonata collected in New Caledonia by the late Mr Paul D. Montague. Ann.

Mag. nat. Hist. (9) 8 : 33-67, text-figs i-n, pis 8-9.
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(9) 14 : 612-614, 2 figs.

TYPE-SPECIMENS OF ODONATA IN BMNH 201

COWLEY, J. 1934. Notes on Odonata (Corduliidae) of the Neotropical Region. Stylops

3 : 91-95, 2 text-figs.

— 1936. Descriptions of three new species of Drepanosticta (Odonata) from the Philippine
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— 1941. A new species of Protoneura from Peru, and a review of the group of Protoneura
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FRASER, F. C. 1923. Zoological results of the Percy Sladen Trust Expedition to Yunnan
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— 1929. Indian Dragonflies. xxxiv. /. Bombay nat. Hist. Soc. 33 : 834-850, 2 pis

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— 1932. Indian Dragonflies. xxxix. /. Bombay nat. Hist. Soc. 35 : 645-656, i pi., i
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— 1933. Fauna Br. India, including Ceylon and Burma. Odonata, vol. i. xiii -+- 423, 180
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— I933a- A revision of the genus Orolestes (Order Odonata). Rec. Indian Mus. 35 : 175-
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!933&- Additions to the Dragonfly (Odonate) Fauna of India, with descriptions of new

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— 1936. Fauna Br. India, including Ceylon and Burma. Odonata, vol. III. xi -f- 461,
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- 1960. A Handbook of the Dragonflies of Australasia, with keys for the identification of
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KIMMINS, D. E. 1929. Ischnura cardinalis sp. n. (Family Agrionidae) : an addition to the
fauna of the Society Islands. Entomologist 62 : 224-225, 3 text-figs.

— 1936. The Odonata of the Oxford University Sarawak Expedition. /. fed. Malay St.
Mus. 18 (i) : 65-108, 17 text-figs.

— 19360. Odonata, Ephemeroptera and Neuroptera of the New Hebrides and Banks
Island. Ann. Mag. nat. Hist. (10) 18 : 68-88, n text-figs, pi. 3.

— 1938. New African species of Elattoneura (Odonata). Ann. Mag. nat. Hist, (n) 1 : 294-
302, 6 text-figs.

— 1943. A new species of Odonata from Fiji. Ann. Mag. nat. Hist, (n) 10 : 698-700,

5 figs-

— 1945. A new species of Odonata from Ecuador. Ann. Mag. nat. Hist, (n) 12 : 187-189,

3 figs-

- 1955. Three new species of African Odonata. Entomologist 88 : 109-113, 3 figs.

— 1957. Odonata collected by Mr J. D. Bradley on Guadalcanal Island, 1953-54. Bull.
Br. Mus. nat. Hist. (Ent.) 5 (8) : 309-320, 5 figs.

— 1958. New species and subspecies of Odonata. Bull. Br. Mus. nat. Hist. (Ent.) 7 (7) :
347-358, 7 text-figs.

— 19580. Miss L. E. Cheesman's Expedition to New Hebrides, 1955. Orders Odonata,
Neuroptera and Trichoptera. Bull. Br. Mus. nat. Hist. (Ent.) 6 (9) : 237-250, 6 text-figs.

— 1966. A List of the Odonata Types described by F. C. Fraser, now in the British Museum
(Natural History). Bull. Br. Mus. nat. Hist. (Ent.) 18 (6) : 173-227.

— 1968. A list of the Type-Specimens of Libellulidae and Corduliidae (Odonata) in the
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204 D- E- KIMMINS

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WATSON, J. A. L. 1969. Taxonomy, ecology and zoogeography of dragonflies from the

north-west of Western Australia. A ust. J. Zool. 17 : 65-112.
WHITE, A. 1843. In Dieffenbach, E. Travels in New Zealand; with contributions to the

geography, geology, botany and natural history of that country. 2, App. : 177-296. London.
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Captain Sir James Clark Ross, R.N., F.R.S., during the years 1839 *° I^43- Ed. Richardson,

J. & Gray, J. E. London [Insects, p. 25].

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Printed in England by Staples Printers Limited at their Kettering, Northants, establishment

STUDIES OF AFRICAN ASILIDAE

(DIPTERA) I. ASILIDAE OF THE

CONGO BASIN

H. OLDROYD

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 24 No. 7

LONDON: 1970

STUDIES OF AFRICAN ASILIDAE (DIPTERA)
I. ASILIDAE OF THE CONGO BASIN

BY

HAROLD OLDROYD

Pp. 207-334 ; 96 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 24 No. 7

LONDON : 1970

THE BULLETIN OF THE BRITISH MUSEUM

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In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
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This paper is Vol. 24 No. 7 of the Entomological
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World List abbreviation
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Issued 27 May, 1970 Price £3 155.

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STUDIES OF AFRICAN ASILIDAE (DIPTERA)
I. ASILIDAE OF THE CONGO BASIN

By H. OLDROYD

CONTENTS

Page
INTRODUCTION ........... 209

ABBREVIATIONS . . . . . . . . . .210

ASILIDAE ........... 210

THE ASILIDAE OF THE CONGO BASIN ....... 214

Key to the Tribes of the African Asilidae . . . . . 215

ACKNOWLEDGEMENTS ......... 329

REFERENCES ........... 330

INDEX ............ 330

SYNOPSIS

A preliminary survey is made of the genera and species of Asilidae known to occur within the
geographical area of the Congo Basin. Keys are given to tribes and genera : wherever possible,
keys are given to all the African species of the genus (S. of the Sahara), but in some genera at
this stage it is possible only to give keys to the species known to occur in the Congo Basin. A
few big genera — notably Ommatius and Neolaparus — are so poorly known that keys are useless
until a generic revision has been undertaken. Three new genera and 39 new species are
described.

INTRODUCTION

THE present paper originated in an invitation to examine and report upon the
Asilidae taken in the Garamba National Park, in the extreme north-east corner of
the Congo Basin. At the time I had on loan a good deal of material from the
Muse"e Royal de 1'Afrique Central, in Tervuren, and from the Institut Royal des
Sciences Naturelles de Belgique, and collections from Urundi and from Kasai made
by my friend Monsieur Frans Francois. All the authorities concerned kindly
agreed that I should write a combined report on their material, which should be
published in the series of volumes of the Exploration du Pare National de la
Garamba : Mission H. De Saeger.

Unfortunately changed circumstances made it impossible for the paper to appear
in this form, and the Editors of the Bulletin of the British Museum (Natural History)
generously agreed to accept it for publication in their series. The collections of the
British Museum (Natural History) are rich in Asilidae from most parts of Africa
except the Congo Basin, and the present collections fill a noticeable gap, in the
centre of the African continent. It is hoped, therefore, that this preliminary

210 H. OLDROYD

review will give some indication of what genera and species of Asilidae occur within
the Congo Basin, and form convenient links with other papers on genera and species
from other parts of Africa, south of the Sahara, which will be necessary before the
Asilidae in the British Museum (Natural History) can be adequately identified.

The expression ' Congo Basin ' is used loosely to cover the geographical area
drained by the Congo River. Biologically the fauna of the north-eastern corner
(where the Garamba National Park is situated) has links with that of Uganda and
Kenya ; species of the Congo Basin proper also extend into the rain forest areas
of the Cameroons and southern Nigeria ; while the fauna of the Katanga uplands
is linked with the Rhodesia highlands rather than with the lowland rain forest.

ABBREVIATIONS

The following abbreviations are used for various institutions and collections:
IPNC = Institut des Pares Nationaux (du Congo), Brussels ; MRAC = Muse"e
royale de 1'Afrique centrale, Brussels (Tervuren) ; IRSNB = Institut R. des
Sciences naturelles de Belgique, Brussels ; FJF = collection of Monsieur F. J.
Francois, Brussels ; BMNH = British Museum (Natural History), London ;
SMNS = Staatliches Museum fur Naturkunde in Stuttgart (Lindner collection) ;
MCZH = Museum of Comparative Zoology, Harvard ; SAIMR = South African
Institute for Medical Research, Johannesburg ; MHNP = Museum national
d'Histoire naturelle, Paris. Collectors' names are given, except in the material
from the Garamba National Park, where all specimens not otherwise credited were
collected by H. De Saeger.

ASILIDAE

Asilidae (Robber Flies) are attractive to the collector of insects because they are
usually easily seen, and because they behave in interesting ways. All known
members of the family are predatory, adults of both sexes feeding by sucking the
body-fluids of other insects, and occasionally of spiders. Most of them capture
their prey in flight, grasping it with the fore legs, piercing it with the proboscis,
and sucking it dry. They feed either in continued flight, or after first alighting
with their prey.

That this habit is well-established in the evolution of the Asilidae is shown by
the degree to which their structure is adapted to an aerial, predatory life. The
head, in particular, is highly functional in its structure. All Asilidae have the eyes
separated, though sometimes closely approximated at one point. A characteristic
of the family, by which Asilidae may be distinguished from, say, Therevidae, is the
way in which the vertex of the head is sunk beneath the level of the eyes. Another
way of expressing this is to say that the eyes are raised above the level of the head,
and this is a more significant description, indicating that the eyes are specially
developed in this family.

The eyes are of such a shape that a considerable area of each eye faces forwards,
and these areas are equipped with facets larger than the rest (Text-figs 56, 57, 60).
The appearance is such as to suggest a special acuity of vision in a forward direction,
correlated with the need to identify prey, and to catch it in the air. Compared
with many other Diptera, the eyes of Asilidae are generally flattened in front, thus

ASILIDAE OF THE CONGO BASIN 211

exposing a greater proportion of the facets to the front and relatively few to the
side, though these lateral facets are strongly curved, and so command a wide field
of view.

These characteristics of the eyes are most exaggerated in the tribe Xenomyzini,
where the effect of large, flattened eyes and tiny frons and face is usually described
as being ' goggle-eyed '.

The legs of Asilidae are developed for the pursuit of prey. In general, they are
stout and strong, in contrast to the fragile legs of such aerial insects as Bombyliidae.
Nearly always they are covered with hairs and strong bristles, which obviously
fulfil a practical function in holding the struggling prey, and preventing it from
injuring its captor ; the characteristic ' moustache ' of the Asilidae, a mass of
bristles just above the base of the proboscis, similarly keeps the prey away from the
eyes and antennae of the robber fly.

In a few genera the legs have acquired additional devices for grasping prey.
The most advanced example is in the genus Gonioscelis, where the fore femur is
expanded basally and its ventral surface is heavily armed with spines and bristles,
which meet the tibia to form a crab-like prehensile organ (Text-fig. 49). In
Hoplopheromems the mid-femur and tibia are equipped with long, spiny bristles.
Many genera have the hind femur swollen and strong, with its curved tibia closing
firmly on to it. This development has occurred, obviously independently, in genera
of all tribes, sometimes, as in Hoplistomerus, with the spines carried on warts or
other processes of the femur.

One of the most enigmatic of such devices is the fore tibial spur of certain genera
of Saropogonini, including Saropogon itself. This formed the basis of Hermann's
group Acanthocnemini : a convenient device for quick identification, but an
unsatisfactory basis for a classification, since we still cannot say whether this
tibial spur is a practical device — thus showing similarity of habit, or of prey — or a
functionless relict from some ancestor, and thus an indicator of affinity. Some
species of Microstylum (Mimoscolia) have a similar spur and terminal process on
the mid- tibia (Text-fig. 39).

The wings of Asilidae are of primary importance in the taxonomy of the family,
though the range of venational pattern is comparatively restricted. A discal cell is
always present, and an almost full set of longitudinal veins : Sc, R\, RZ+Z, ^4+5, MI,
MZ, MZ, M4, Cui, i A. Variations in venational pattern arise from the meeting of
adjacent veins before they reach the wing-margin, and thus the closure of certain
cells, of which the most important is the marginal cell, lying between the veins R\
and RZ+S (Text-fig. 69).

The marginal cell is closed, with a short terminal stalk, in Asilini, Ommatiini,
Atomosiini, and most Laphriini ; it is open, with veins RI and RZ+S reaching the
wing-margin separately, in Leptogasterini, Saropogonini, Stichopogonini and
Xenomyzini. In certain genera related to Laphystia this cell is normally open,
though vein R2+3 has a characteristic retrograde curve. These genera comprise
Hermann's group Prytaniinae and Hull's tribe Laphystiini, transitional between the
old subfamilies of Laphriinae and Dasypogoninae. In most respects they are most
closely related to Laphriini, and are here treated as belonging to that tribe.

212 H. OLDROYD

It is generally assumed that open cells are a more primitive condition, and closure
of cells is a specialization ; though of course this specialization has occurred in-
dependently in many different lines of evolution. The most generalized venation
occurs in the Leptogasterini, where often all the cells are open, including the anal
cell. The last is particularly variable, being sometimes divergently open, almost as
in some Nematocera, but sometimes closed far from the wing-margin. Leptogasterini
have lost the alula, and the shape of the wing and of individual forks and cells show
clearly that being generalized cannot necessarily be equated with being primitive.

Leaving aside the Leptogasterini, which are a peripheral group, whether primitive
or advanced, the central group of the family is the Saropogonini with Stichopogonini
and Xenomyzini as specialized offshoots. Saropogonini include a diversity of
genera, all of which have an open marginal cell, even though other venational
specializations occur, e.g. in Microstylum.

The generally robust structure of Laphriini extends to the wings, where additional
strength is often provided by the closure of both the first and fourth posterior cells
(Text-figs 18-20). These must give considerable rigidity along the posterior area
of the wing. On the other hand the actual wing margin, which in Asilidae is
normally sclerotized into an ' ambient vein ', is membranous in a number of genera,
especially in Laphystia and related genera.

Among Asilini the conspicuous peculiarity lies in the genera Promachus, Alcimus,
Philodicus and Apoclea, where an additional vein unites the fork of R^+5 with the
stem of Rz+3, producing three submarginal cells. Since Asilini are clearly an
advanced tribe, which at specific level show signs of recent evolution, it would be
reasonable to visualize this additional vein as a new structure, perhaps giving
additional rigidity to the radial field of the wing. However, many Brachycera in
different families have a stump vein on the radial fork, and sometimes a complete
cross-vein : for example, some Bombyliidae, including the entire tribe Exoprosopini.

There has therefore been a temptation to look upon three submarginal cells as a
primitive condition in Brachycera, from which the more usual two submarginal
cells are derived by reduction. Shannon and Bromley (1924) regarded the
Promachus-group of genera as evidence that the proper numbering of the branches
of the radial sector should be : RI, RZ, ^3+4, ^5. In this way Shannon & Bromley
accepted it as axiomatic that the additional vein in these genera is a true vein,
one of the ancestral complement.

In fact, supernumerary veins often occur in Brachycera. In Nemestrinidae they
are a regular feature of certain genera. In Syrphidae the ' spurious vein ' is a
family characteristic. Among Asilidae it is quite common to find supernumerary
cross-veins, sometimes only in one wing. The small anterior cross-vein (r-m) is
often duplicated. There is evidence that such veins appear during the pupal
period, and are the result of sclerotization of a fold. On this interpretation it is
suggested that the stump vein from the radial fork, when this is present, and even
the complete vein in Promachus and its allied, are no more than this, and give no
indication of phylogeny.

The thorax and abdomen of Asilidae show no remarkable features. The abdomen
varies from slender and elongate in Leptogasterini, most Asilini and many other

ASILIDAE OF THE CONGO BASIN 213

genera, to the extremely squat abdomen of genera such as Sisyrnodytes, in which
the abdomen is short, broad, flattened, and bee-like in form.

With few exceptions, the genitalia of both sexes are exposed, and readily examined
for purposes of classification. Female terminalia may be simple, or they may take
the form of an ovipositor. Although an ovipositor may appear highly characteristic
it is an adaptive character, suited to the requirements of egg-laying. The larvae of
Asilidae live either in soil or in wood, but the eggs are laid in a variety of special
sites. They may be simply dropped on to the ground, in which case the ovipositor
is of very simple, lamellate structure. Many eggs are concealed in flowers, leaf-axils,
or even inside a slit cut into the tissue of a plant : it seems that the object is conceal-
ment and protection of the egg, and possibly of the first instar larva. Many of the
more striking ovipositors, which are most frequent in the advanced tribe Asilini, are
adapted to the circumstances of oviposition. The manner of oviposition may be
quite different in related genera, or even within a single genus, and so ovipositors
are unreliable indicators of relationships.

Male genitalia present a complicated problem. They are constructed mainly
from the ninth segment. The ninth tergite, or epandrium, is sometimes entire,
sometimes split into the superior forceps. From the ninth sternite arises a pair of
inferior forceps (gonopods) which often appear double because each bears on its inner
surface an accessory lobe, the clasper. Within the two claspers appear the aedeagus
and its two parameres. The basic pattern is thus that of a trifid median structure,
flanked by three pairs of lobes, and topped (dorsally) by a median pad formed from
the paired anal lamellae (proctiger).

The variations upon this basic structure are extensive, and are made more
confusing by partial or complete inversion. Thus in Laphriini the typical genital
structure is boat-shaped, the apparent hull of the boat consisting of the undivided
ninth tergite (epandrium), which has become inverted and appears to house the
other (ventral) appendages (Text-figs 3-36).

The problem of rotation of male genitalia in Diptera, leading to inversion, has
received undue prominence in textbooks, more especially in regard to muscoid flies.
Rotation may be a source of confusion in comparative morphology, but — at least in
Asilidae — it is usually possible to distinguish between the true dorsal aspect, with
the lamellae, and the true ventral aspect, with the gonopods and claspers. As an
indication of relationships, rotation is an unsafe guide. In many genera rotation
occurs during mating : i.e. the genitalia of the two sexes are engaged, and then the
male changes position, twisting the genitalia in the process. This rotation may be
partial or total, depending upon the mating attitudes, and may or may not persist
during later life, or after death. Consequently no attention should be paid to
rotation per se, beyond ensuring that the correct orientation of the parts is under-
stood before they are compared with others.

The specific identification of Asilidae is much more difficult than is generally
thought, and possibly for this reason this attractive family has not been as exten-
sively studied as, say Syrphidae. No large scale revision of African Asilidae has
ever appeared, and in every genus the student is faced with a dilemma : if few
species are known, then material is scarce, and comparison difficult ; if material is

214 H. OLDROYD

abundant, as in Ommatius and Neolapams, the variation is so great that specific
identification is not possible until the whole genus has been revised.

Substantial progress in knowledge of African Asilidae will come only from com-
plete revisions of successive genera.

THE ASILIDAE OF THE CONGO BASIN

The collections of the British Museum (Natural History) include much material
from Africa south of the Sahara, but for historical reasons little of this comes from
the area of the Congo Basin. The writing of the present paper seemed to provide
an opportunity to review the Asilidae of this larger and important natural region
of central Africa, which would be complementary to later studies that it was hoped
to make of Asilidae from the rest of the Ethiopian Region.

Unfortunately as the work progressed it became evident that a number of genera
were insufficiently known for a definitive review to be possible. The outstanding
example is the genus Neolaparus, flies of deceptively simple appearance which so
far have resisted all attempts at a classification. Only a thorough study of all the
African species, with an attempt to find new taxonomic characters, will make it
possible to rationalize this genus. Ommatius is more tractable, but there is an
abundance of species, many of which are clearly of wide distribution. These and
several other genera must be revised for the entire region before it will be possible
to deal adequately with any faunistic survey.

The material described in the present paper, therefore, consists of the collections
of the Mission H. De Saeger from the Pare National Du Garamba, together with such
other material as could be handled satisfactorily at the same time. Whenever
possible I have given a key to species, from this collection, or from the Congo Basin,
and in some genera a key to all the known species from the Ethiopian Region.

One fact that clearly emerges from this study is that there is no fauna of Asilidae
that is characteristic of the Congo Basin, or even of the equatorial forest belt as a
whole. The distribution of Asilidae resembles that of Tabanidae : both families
seem to flourish best in savanna country, with an open, mixed vegetation, which
provides a multiplicity of habitats.

There are interesting parallels between the tribe Laphriini of the Asilidae and the
genus Chrysops of the Tabanidae. Chrysops spp. are known as ' deer flies ' in North
America because their typical habitat is the open woodland in which deer abound,
and Laphriini, with their wood-living larvae, are characteristic of this habitat.
Chrysops has invaded the high forest by way of the tree-canopy, coming down to
ground level where the canopy is broken by rivers, by clearings, or by the out-
cropping of rocky or mountainous areas which bring ground level up to the canopy
of the surrounding forest. It would be interesting to discover whether the high
canopy of the Congo Basin also supports a large population of Laphriini, which fly
among the tree-tops and prey upon the Hymenoptera and other insects feeding on
the flowers of the forest trees.

The Pare National du Garamba is not situated in high forest, and has not much
open woodland. It is mostly park savanna, i.e. predominantly grassland with
scattered trees (Exploration du Pare National du Garamba ; Afdelung Introduction,

ASILIDAE OF THE CONGO BASIN 215

p. 101 ; pis. III-XII). It is noteworthy that Laphriini are poorly represented
within the confines of the Pare, though they are numerous among collections from
elsewhere in the Congo Basin.

The Asilidae of the Pare National du Garamba are mainly Saropogonini,
Stichopogonini, Xenomyzini and Asilini, and belong to genera which have a wide
distribution throughout the savanna areas of Africa. It is these genera (e.g.
Neolaparus, Ommatius) that cannot be comprehensively treated in the present paper,
because the Garamba fauna is a small and unrepresentative sample of a very ex-
tensive and difficult range of species.

KEY TO THE TRIBES OF AFRICAN AsiLiDAE1

1 Marginal cell of the wing open .......... 2

- Marginal cell of the wing closed .......... 6

2 Pulvilli absent. Very narrow, elongate flies . . LEPTOGASTERINI (p. 215)
Pulvilli nearly always present ; if absent, flies of different shape .... 3

3 Prosternum isolated, and surrounded by membrane (between front coxae). Female

with ninth tergite divided into spine-bearing plates (acanthophorites)

SAROPOGONINI (p. 257)
Prosternum complete, with little or no membranous area ..... 4

4 Vertex of head saddle-shaped. Dusty grey flies, living in sand or on rocks. (Text-

figs 53,-56, 57, 60) . ... STICHOPOGONINI (p. 281)

Vertex not saddle-shaped ........... 5

5 Flies with small face and frons and very large ' goggle ' eyes. XENOMYZINI (p. 285)
Head and eyes not of this shape . some LAPHRIINI (LAPHYSTIINI) (p. 224)

6 Antennae blunt or club-shaped. Often with mesopleural bristles .... 7
Antennae with slender arista, sometimes feathered ...... 8

7 Vein M$ straight and parallel with outer vein of discal cell. Small flies, rather like

sawflies ATOMOSIINI (p. 257)

Vein MS curved, not parallel with outer margin of discal cell . LAPHRIINI (p. 224)

8 Arista of antennae feathered ....... OMMATIINI (p. 313)

Arista of antennae bare ........ ASILINI (p. 297)

Tribe LEPTOGASTERINI

Members of this tribe have a characteristic habitus, which sets them apart from
nearly all other Asilidae. The abdomen is long and slender, usually slightly clubbed
apically. The wings are generally narrow, with the anal lobe and alula evanescent,
and with a simplicity of venation in which even the anal cell is often wide open.
The legs are slender and elongate, especially the hind pair, of which the femora and
tibiae may be clavate. Pulvilli are always absent, and the empodium may be
absent or greatly reduced. The metanotum and postmetacoxal area are flattened
obliquely, in such a way that the coxae are pushed forwards, and the base of the
abdomen raised, contributing to what has been called the ' agrionid ' appearance
of these flies. The thorax of many Leptogasterini is further distorted by having
the dorsum compressed into a hump, arid by the development of a pair of processes
on the mesonotum which overlap the pronotum ; in the genus Euscelidia a process
of the pronotum lies between them.

1 For a fuller account of tribal classification, and a bibliography see Oldroyd (1963).

216 H. OLDROYD

To a great extent these features of Leptogasterini are clearly adaptive. The
tarsi, for instance, are adapted for curling securely round a grass stem, in which
process they are actively assisted by the long, curved claws, and by the loss of
pulvilli.

Dipterists who have specially studied Leptogasterini have been led to emphasize
the differences between these and all other Asilidae. Janssens (1954 : 114) stated
an extreme view : ' Je considere . . . qu'il y avait lieu peut-etre d'instituer pour
Leptogaster et quelques voisins une famille qui formera avec les Asilidae une super-
familie ASILOIDEA '. Martin (1968) developed this view at some length, but
Oldroyd (1969) examined Martin's argument in detail, and endorsed the view of
Hull (1962 : 296) that : ' All these characters are collectively found in Lepto-
gasterinae, but are singly shared with other asilids '.

The subdivision of Leptogasterini into genera is also unexpectedly difficult.
The key given by Oldroyd (1963 : 8) is unsatisfactory in some particulars, and fails
to locate generically any species with the following combination of characters :
occiput with fine hairs only, without bristles ; no bifid pronotal process ; and no
swollen or plumate hind legs. Such specimens might be regarded either as Lepto-
gaster without occipital bristles or as Euscelidia without the pronotal process ;
sometimes the general impression is of the former, sometimes of the latter.

In the present paper the genera are interpreted as in Janssens' papers, and all
species which lack the pronotal process, and have the anal cell widely open, are
classed as Leptogaster, whether or not they have strong occipital bristles.

KEY TO THE AFRICAN GENERA OF LEPTOGASTERINI

1 Prothorax with a bifid process, which arises between the two processes of the meso-

notum EUSCELIDIA Westwood (p. 219)

Prothorax without any process between the two processes of the mesonotum . . 2

2 Legs swollen or ornamented .......... 3

Legs neither swollen nor ornamented ......... 4

3 Hind femora and tibiae strongly swollen, with thick, erect hairs

LASIOCNEMVS Loew (p. 223)
Hind femora not strongly swollen, but distinctly elongate. Hind tibiae and tarsi

with conspicuous plumes .... DOLICHOSCIUS Janssens (p. 223)

4 Anal cell of the wing open. Mostly small, rather sombre flies

LEPTOGASTER Meigen (p. 216)
Anal cell of the wing closed and stalked. Mostly large, or very large, brightly

coloured flies AMMOPHILOMIMA Enderlein (p. 219)

LEPTOGASTER Meigen

Leptogaster Meigen, 1803 : 269. Type-species : Asilus cylindricus De Geer, 1776, monotypic

as Asilus tipuloides Fabricius, 1775.
Gonypes Latreille, 1805 : 309. Type-species : Asilus cylindricus De Geer, 1776, monotypic as

Asilus tipuloides Fabricius, 1775

Very few species of Leptogaster had been recorded from the Congo Basin until
Janssens described twenty-five species in a series of papers (1952-57). Many of
these are known from unique specimens, and only a study of much more extensive

ASILIDAE OF THE CONGO BASIN 217

material will make it possible to indicate the extent of intraspecific variation, or
the distribution of species within the area. In the meantime the following key,
compiled from the published descriptions, is offered as a guide to provisional
identification. In addition to Janssens' species there are L. ludens Curran (1927 : i)
— a single female from Banana, at the Congo mouth — L. bicingulata Bezzi (1905 :
279) described from Eritrea and recorded by Janssens from Kitega ; and L. picti-
pennis Loew (1857 : 353) described from S. Africa, and also recorded by Janssens
from Bururi.

KEY TO THE SPECIES OF LEPTOGASTER DESCRIBED BY JANSSENS

1 Mesonotum matt, sometimes with pattern of tomentum ..... 2

- Mesonotum shining, or with shining stripes . . . . . . . 10

2 Empodium very small, or absent. Arista as long as all three antennal segments

together .......... f aristalis, 1957 : I0

(both from Eala, and difficult to separate) .... \ealensis, 1954 : 3°3

- Empodium distinct, usually about half as long as claws ..... 3

3 Mesonotum matt brown or black, with or without indistinct pattern. Hind femora

with darker tip or band .......... 4

Mesonotum red or brown, with three darker stripes. Hind femora uniform . . 8

4 Moustache composed of eight long, black hairs. Occipital bristles unusually strong

melanomystax, 1954 : X33
Moustache pale, occipital bristles weak ........ 3

5 Wings darkened in basal two-thirds, leaving tip pale . . . fumosa, 1954 : 302
Wings either clear, or almost uniformly tinted, with tip sometimes darker . . 6

6 Hind femora gradually darkened towards tip. Wings smoky . . plebeja, 1957 : 9

- Hind femora with distinct subapical dark ring ....... 7

7 Face and proboscis dark. Abdomen pitch-black . . evanescens, 1954 : I27

- Face grey and proboscis red-brown. Abdomen olive-grey with grey pubescence,

especially posteriorly ........ iimndiana, 1953 : 7

8 Empodium half as long as claws. Vein Rz+3 curved forwards at tip ... 9

- Empodium tiny, and vein RZ+Z curved backwards at tip . . velutina, 1954 : 127

9 Occiput with a row of unusually stiff bristles. Body generally covered with silky

grey or whitish hairs ......... puella, 1953 : 6

- Occiput with a few dark hairs. Body generally dark, without silky appearance

rufescens, 1954 : 125 (p. 218)

10 Hind femora uniformly coloured ......... 1 1

- Hind femora banded, striped, or merely becoming progressively darker towards tip 14

11 Wings clear ............. 12

- Wings more or less tinted .......... 13

12 Hind tibiae thickest in middle, tapering towards base and tip pallipes, 1953 : 9 (p. 218)

- Hind tibiae thickest at tip. Thorax with two strong bristles posteriorly

rufa, 1953 : 9, schoutedeni, 1954
13 Thorax black .......... sericea, 1954

- Thorax clear, translucent yellow-brown ..... pellucida, 1954

401

131
132

14 Mesonotum with dark pattern of stripes or spots . . . . . . 15

- Mesonotum without pattern, though sometimes paler at sides, or with a bare stripe 17

15 Mesonotum with silvery spots of tomentum, and also with numerous small bare spots

hermelina, 1954 : 130

- Mesonotum without spots .......... 16

16 Mesonotum brilliant yellow, with triple black stripe. Hind femora yellow with

indistinct band .......... vindex, 1954 : 15

2i8 H. OLDROYD

- Mesonotum dark brown, with a darker stripe. Hind femora reddish brown with

dark band . . . . . . . , . . pilicnemis, 1954 : I29

17 Wings brown at tip. Mesonotum black with grey tomentum, which leaves a bare

stripe. .......... apicalis, 1954 : J32

- Wings not brown at tip, clear, or uniformly tinted . . . . . . 18

18 Bronze and sepia species, hind femora with longitudinal stripe . upembana, 1954 : I2&

- Without bronze colouration or longitudinal stripe on femora . . . . 19

19 Hind femora and tibiae with a strongly marked dark ring near tip . . . 20

- Hind femora only indistinctly darker towards tip . . . . . . 21

20 Mesonotum black with yellow tomentum, which becomes silvery on pleura. A

robust species of 10 mm. Hind femora conspicuously knobbed, and with two

rows of black spines ventrally .... penicillata, 1954 : T3O (P- 2J8)

- Mesonotum red, with weak yellowish tomentum on pleura. A fragile species of

6 mm. Hind femora not conspicuously knobbed, and without ventral spines

stichosoma, 1957 : 6

21 Empodium vestigial ........... 22

- Empodium more than half as long as claws ..... pyragra, 1957 : 8

22 Hind basitarsus as long as preceding four tarsomeres together tarsalis, 1954 : 29 (p. 218)

- Hind basitarsus normal ....... basilewskyi, 1955 : 304

The following species were represented in the collections from the Garamba
National Park :

Leptogaster pallipes Janssens

Leptogaster pallipes Janssens, 1953 : 9.

GARAMBA NATIONAL PARK : P.N.G., 1849, i <£, i $, 31^.1951 (/. Verschuren),
3612, Iso III, i $, ii.vi. 1952 (IPNC).

Leptogaster penicillata Janssens

Leptogaster penicillata Janssens, 1954 : 130.

GARAMBA NATIONAL PARK : P.N.G., 195, i/c/2'", i <j>, 3.1.1950 ; 1276, III/gd/2,
i <$, 19.11.1951 ; 1334, II/gd/4, i ?, 6.iii.i95i ; 2861, II/hd/4, i $, 6.xii.i95i ;
2944, i ex., II/gd/4, 27.xii.i95i ; 3077, II/gd/io, i ex., 30.1.1952 (IPNC).

Leptogaster tarsalis Janssens

Leptogaster tarsalis Janssens, 1953 : 129.

The following three specimens may probably be assigned to this species, though
the third specimen, a female, is larger and more robust than the others. All three
show a characteristic narrowing of the second submarginal cell, veins #4 and R$
both being curved and approximated at the wing-tip.

GARAMBA NATIONAL PARK : P.N.G., 199, I /a/3, i <£, 7.ix.i95o ; 2991, II/fd/i7,
i ex., 3.1.1952 ; 529, Akam., i <j>, 19^.1950 (IPNC).

Leptogaster rufescens Janssens

Leptogaster rufescens Janssens, 1954 : I25-

GARAMBA NATIONAL PARK : P.N.G., 1474, II/ge/2, i ?, 31.^.1951 ; 2102,
II/fd/3, i $, i6.vii.i95i ; 2107, II/hd/4, J ?> I7-vni-I95I» 2 ex. ; 2160, Il/gd/n,

ASILIDAE OF THE CONGO BASIN 219

i <$, 28.vii.i95i ; 2223, II/fd/4, i #, S.viii.igsi ; 3488, Inimvua, i <j>, 2O.V.I952 ;
3606, Mt Tungu (S), i c£, g.vi.1952 ; 3964, II/gd/4, i <$, i °., 23^111.1952 (IPNC).

AMMOPHILOMIMA Enderlein

Ammophilomima Enderlein, 1914 : 155. Type-species : Ammophilomima imitatrix Ender-
lein, 1914, by original designation.

? Lagynogaster Hermann, 1917 : 12. Type-species : Lagynogaster fuliginosa Hermann, 1917,
by original designation.

In his revision of Ammophilomima, Janssens (19536 : 1-12) discussed the sup-
posed differences between Enderlein's genus Ammophilomima and Hermann's
Lagynogaster, and concluded that probably these two '. . ne sont qu'un seul et
meme genre '. Janssens described two species from the collections of the Staats-
sammlung in Munich, where they bore manuscript names given them by Hermann,
but which had never been published.

Although eumenoides was based upon a specimen from Malawi, and imitatrix
from the Cameroons, Janssens records all the species in his key from the region of
Eala, in the Congo Basin. Here is a translation of his key :

KEY TO SPECIES OF AMMOPHILOMIMA

1 Hind femora encircled with a yellow band ........ 2

Hind femora not encircled with a yellow band ....... 7

2 Hind tibiae with a yellow apex .......... 3

- Hind tibiae without any yellow apex ......... 5

3 Stigma filling the whole of the subcostal cell ....... 4

Stigma confined to tip of subcostal vein ..... imitatrix Enderlein

4 Empodium longer than half length of claws. Wings brown. Hind legs black and

yellow ........... straeleni Janssens

- Empodium less than half length of claws. Hind legs brown and bistre (sooty brown).

Wings hyaline ......... evanescens Janssens

5 Hind basitarus largely bistre .......... 6

- Hind basitarus entirely brown ....... ghesquierei Janssens

6 Mesonotum unicolorous ....... aequinoctialis Janssens

- Mesonotum bearing a large black spot in the shape of a trident . basilewskyi Janssens

7 Antennae uniformly reddish yellow .... eumenoides Janssens (p. 219)

Cauripennis Janssens

- Antennae darker .........< •;

^ montana Janssens

The collections from the Garamba National Park contained specimens of A.
eumenoides.

Ammophilomima eumenoides Janssens

Ammophilomina eumenoides Janssens, 1953 : 4.

GARAMBA NATIONAL PARK : P.N.G., 3311, PPk. 73/d/g, 2 ?, 8.iv.52 ; 3623,
Iso II/2, i ?, i8.vi.i952 (IPNC).

EUSCELIDIA Westwood
Euscelidia Westwood, 1849 : 232. Type-species : Euscelidia rapax Westwood, monotypic.

220 H. OLDROYD

Species of Euscelidia are usually recognizable by their general appearance, as
robust and shining Leptogasterines. The type-species and a few others have hairy
hind legs, which could lead to confusion with Lasiocnemus, but the existence of a
vertical process of the pronotum projecting between the two processes of the
mesonotum is diagnostic.

About eight or nine species of Euscelidia have been recorded from the Congo
Basin, most of them described in various papers by Janssens. The differences in
colour and pattern are more subtle than in Leptogaster, and it is not practicable to
attempt a key compiled from descriptions. Only a detailed study of species both
in the Congo Basin and outside it, with comparisons of genitalia, will make such a
key possible. Meanwhile species must be named by direct comparison with
Janssen's descriptions and figures.

The collection from the Garamba National Park contained two of Janssens
species and two others that are clearly new.

Euscelidia bicolor Janssens

Euscelidia bicolor Janssens, 1954 : I23-

GARAMBA NATIONAL PARK : P.N.G., 3844, Mt Moyo, 2 $, 29.vii.52 (IPNC).
The wings are more heavily infuscated than is indicated in the original description.

Euscelidia f estiva Janssens

Euscelidia festiva Janssens, 1954 : I23-

GARAMBA NATIONAL PARK : P.N.G., 529, Akam, i $, 19^.1950 ; 848, I/b/2,
i $, 27.ix.5o (G. Demouliri) ; 895, Napokomweli, i <£, i <j>, iS.x.so (G. Demoulin) ;
3612, Iso III, i (J, n.xi.52 ; 3642, Iso/II/n, i ex., i6.vi.i952 (IPNC).

Euscelidia dorata sp. n.

(Text-fig, i)

A yellow-brown species, greatly resembling Leptogaster rufescens Janssens, but
with a distinct pronotal process, and without any strong occipital bristles. Its
nearest relative among Janssens' species of Euscelidia is E. bicolor, from which
dorata differs in having the thorax yellow-brown, tomented, and a very sparse
moustache.

6* Head. Frons with dense golden brown tomentum ; ocellar tubercle black, with sparse
golden tomentum, and no ocellar bristles. Face tomented brownish, silvery when seen
obliquely ; almost plane, with a tiny tubercle on mouth-margin, which bears a sparse moustache
of about four brownish bristles. Proboscis and palpi clear brown. Occiput with dense tomen-
tum, grey-brown, with no bristles at all, and only sparse, short, yellowish brown hairs. Anten-
nae brown with black hairs ; third segment same length as arista, each equal to sum of first
two segments.

Thorax yellow-brown both in ground colour and in tomentum. Mesonotum quite strongly
humped, tomented, without distinct pattern, but with an indistinct yellowish median stripe.
Bristles yellow (almost all broken off in specimens available) : i notopleural, i intra-alar.

ASILIDAE OF THE CONGO BASIN 221

i ? postalar : fine, short, yellow hairs disposed in humeral areas and along lines of dorsocentrals.
Scutellum with a distinct discal furrow, yellowish brown in ground colour, with whitish tomen-
tum, short yellow hairs, and no discal bristles. Pleura yellow-brown with dense whitish tomen-
tum and only sparse, fine, yellow hairs.

Abdomen. First five segments tubular, yellow-brown, shining through thin yellow tomen-
tum, and with fine black hairs ; sixth and following segments more clavate, dark brown, with
hairs longer and mostly yellow ; venter similar, but more yellowish and with yellow hairs.
Genitalia (Text-fig, i) mahogany-brown.

Legs clear yellow-brown : only tips of tarsi and of hind tibiae rather deeper brown. Hind
femora and tibiae distinctly clavate. All legs clothed with rather long hairs, which are mostly
clear, but may appear black when silhouetted against the light.

Wings almost clear of pigment, but extensively covered with microtrichiae, especially in
apical half. Halteres with yellow stalk and brownish knob.

Length of body 10 mm ; of wing 7 mm.

Holotype <$. GARAMBA NATIONAL PARK : P.N.G., 2644, II/fd/i5, 22.ix.5i
(IPNC).

Paratypes. GARAMBA NATIONAL PARK : 208, i/b/i, i $, 15.11.1950 (G. Demouliri) ;
998, II/d/6, i ?, 2i.xii.5o (/. Verschureri) ; 949, II/c/, i <$, 2i.xii.5o (/. Verschureri) ;
1260, II/fc/6ar, i $, 16.11.1951 (/. Verschureri) ; 1272, II/ed/i4, i <j>, 17.11.1951 ;
1576, II/fb/4, i ?, I9.iv.i95i (/. Verschureri) ; 1633, II/ee/7, 6 $, 27.^.1961 (/.
Verschureri) ; 2024, II/gd/i4, 3 $, 30^1.1951 ; 2051, II/ge/6, i ?, io.vii.i95i ;
2071, Il/gd/n, i c£, I2.vii.i95i ; 2464, II/fd/i5, i ?, I2.ix.ig5i ; 2780, II/gd/4,
i <£, 23.ix.i95i ; 2935, II/fd/io. i $, 2o.xii.i95i ; 2841, II/fc/6, i <j>, 26.xii.i95i
(/. Verschureri) (IPNC).

FIG. i. Euscelidia dorata. Male genitalia.

222

H. OLDROYD

Euscelidia moyoensis sp. n.

(Text-fig. 2)

A species with shining black thorax, and legs extensively marked in black, distin-
guished from the other species known to me by the distinct black band at the
extreme tip of the hind femora. From E. lucida Oldroyd it differs in the much less
evident pattern of tomentum on the thorax, the more prominent silky hairs of the
anterior sternopleuron, and the more abundant pale hairs of the moustache. It
also has close affinities with E. datis (Walk.) from Sierra Leone, but cannot be
compared in great detail because the unique type of the latter is too badly damaged :
the fore and middle legs of moyoensis seem to be much more heavily infuscated.

cj Head. Frons and face with golden tomentum ; moustache a row of 15-20 white bristles
on mouth-margin : no other facial or frontal hairs. Antennae (broken), palpi and proboscis
black, with black hairs. Occiput with golden tomentum and fine yellow hairs.

Thorax. Mesonotum mostly shining black, with yellow tomentum only at extreme sides,
and particularly anteriorly, and on humeral slopes ; long, golden hairs lie vertically, and long,
yellowish white hairs arise from tomented areas ; rest of dorsum with short, black hairs.
Scutellum covered with yellowish white tomentum, with erect, curved yellowish hairs on disc
and on margin. Pronotum, including median lobe and twin lobes of mesonotum, covered with
whitish tomentum, which also covers pleura. Mesopleuron, pteropleuron and anterior part of
sternopleuron also with conspicuous tufts of long, snow-white hairs.

A bdomen in ground colour black, with yellow basally on third and fourth segments, the whole
covered with whitish tomentum. Hairs mostly white, some black dorsally, longer laterally
and on sternites, which are a little more yellow than tergites. Male genitalia (Text-fig. 2) black.

FIG. 2. Euscelidia moyoensis. Male genitalia.

ASILIDAE OF THE CONGO BASIN 223

Legs. A pattern of whitish yellow, deep honey-yellow, and black-brown. Black-brown
covers extreme tips of fore and hind femora, most of mid femora except bases, a median ventral
patch on inner face of each hind femur, anterior and posterior stripes on all tibiae, tips of all
basitarsi, and all other tarsomeres. Hind femora with narrow, whitish yellow stem, deep
yellow, elongate knob, and small, apical, black-brown band. All basitarsi conspicuously
whitish yellow, except for extreme tip, which is black-brown. White bristles on tibiae, but
tarsal bristles black.

Wings. Uniformly, but faintly smoky. Halteres red-brown.

Length of body 12 mm ; of wing 8 mm.

Holotype $. GARAMBA NATIONAL PARK : P.N.G., 3844, Mt Moyo, 2g.vii.52
(IPNC).

DOLICHOSCIUS Janssens

Dolichoscius Janssens, 1953 : 2. Type-species : Dolichoscius francoisi Janssens, 1953, mono-
typic.

Janssens has described two species of Dolichoscius in two separate papers. He
clearly intended the type-species to be longipes, from the Upemba National Park,
but unfortunately this paper (1954) did not appear until after the description of
francoisi from Urundi, which becomes the type-species by monotypy.

Dolichoscius francoisi Janssens, 1953 : 2, Urundi.

Dolichoscius longipes Janssens, 1954 : 120, Upemba.

LASIOCNEMUS Loew

Lasiocnemus Loew, 1951 : 2. Type-species : Lasiocnemus obscuripennis Loew, 1851, by
original designation.

Although Janssens (1952 : 6-10) records several species of Lasiocnemus from the
Congo Basin, including such species as L. lugens Loew, which were originally des-
cribed from distant parts of Africa, there were no species among the material from
the Upemba National Park (Janssens, 1954), and none in the collections from
Garamba.

For the sake of completeness in this review, therefore, I can only quote Janssens'
key to the species recorded by him from the Congo Basin.

KEY TO SPECIES OF LASIOCNEMUS FROM THE CONGO BASIN

1 Wings clear, except for a stigma at tip of subcostal vein . . hyalipennis Janssens

- Wings smoky ............. 2

2 Wings uniformly smoky ........ hermanni Janssens

- Wings with some kind of pattern ......... 3

3 Wings with separated clear flecks ......... 4

Wings with a hyaline crossband .......... 6

4 Legs uniformly black, or dark brown ......... 5

- Legs clear brown, with brown markings on hind femora and tibiae. (Wings with two

large clear spots which almost form a crossband)

fascipennis Engel & Cuthbertson, 1939 : 185

5 Face with brown pubescence ; halteres black ; empodium equal to three-quarters

of length of claws ........ lugens Loew 1857 : 353

224 H- OLDROYD

Face with white pubescence ; halteres pale yellow ; empodium almost as long as

claws ......... obscuripennis Loew, 1851 : 2

6 Transverse clear band of wings in the form of an elongate triangle. Pubescence of

hind tibiae grey apically .... griseicinctipes Speiser, 1913 : 141

- Transverse clear band of wings irregular. Pubescence of hind tibiae reddish apically.

Empodium not half as long as claws . . . anthracinus Janssens, 1952 : 9

Tribe LAPHRIINI

More than one third of the species recorded in the present paper belong to the
tribe Laphriini. Although this proportion is exaggerated by the fact that certain
big genera of other tribes — notably Neolaparus and Ommatius — cannot at present
be reported upon so fully, yet the abundance of Laphriini underlines the fact that
these genera flourish among trees. Many, perhaps most of them, spend their larval
life in the stumps of fallen logs, often in the burrows made by wood-boring beetles
and carpenter-bees.

Most Laphriini are distinctive in appearance, broadly built, often with the ab-
domen basally constricted ; wings strong, with stout veins, marginal and fifth
posterior cells closed ; legs often stout, with inflated femora and curved tibiae ;
and antennae stout, often clavate. The body surface is toughly sclerotized, and
often punctate, giving the appearance of a strong, aggressive insect, with more than
a superficial resemblance to a wasp or a bee.

Laphriini can usually be recognized by the combination of closed marginal cell
and styliform antennae, in contrast to the aristiform antennae of the Asilini, and the
open marginal cell of other tribes. It has been explained above (Introduction) that
the present concept of Laphriini includes not only the tribes Andrenosomini and
Ctenotini of Hull (1962), but also the more controversial ' Prytaniinae ' of Hermann,
the tribe Laphystiini of Hull and others. On the other hand the Atomosiini,
apparently only slightly separated by the small details of wing-venation, sustain
this distinction on a world scale, though they are scarce in tropical Africa, and only
one species is included in the present paper (see below).

KEY TO GENERA OF AFRICAN LAPHRIINI

1 Proboscis flattened into a blade like a paper-knife, with its edges dorsal and ventral 2

- Proboscis triangular in cross-section, with a flat surface ventrally. Sometimes

curved upwards into a sickle-shape ........ 4

2 Face gently swollen up to base of antennal tubercle. Large, hairy, bee-like flies

DASYLLINA Bromley (p. 231)

- Face abruptly swollen into a knob that occupies only lower half .... 3

3 Antennae conspicuously elongate. Abdomen constricted between segments 2-3,

giving a wasp-like appearance. Margin of scutellum with very short hairs, or
none at all. Hind femora with tuberculate spines as in Hoplistomerus

STORTHYNGOMERUS Hermann (p. 231)

- Antennae not conspicuously elongate ; third segment often rather plump. Abdo-

men seldom constricted, and then it is between 1-2 segments. Margin of scutellum
usually with long hairs or fine bristles .... LAPHRIA Meigen (p. 226)

4 Costa of wing not thickened as far as tip of vein R$, and entire hind margin mem-

branous. First posterior cell closed and stalked . . . ... 5

- Costa of wing thickened at least as far as vein M^, or beyond .... 9

ASILIDAE OF THE CONGO BASIN 225

5 Claws blunt at tip. Vein M% of wing nearly always cut short before reaching wing-

margin. Palpi with only one segment (Text-fig. 25). Lower margin of occiput

produced into a rim CTENOTA Loew

Claws pointed at tip. Vein M% of wing reaching margin, or almost so. Palpi with

two segments (Text-fig. 24). Lower margin of occiput not produced . . 6

6 Hind femora very strongly swollen. Third antennal segment hairy above

LAXENECERA Macquart (p. 235)
Hind femora not strongly swollen. Third antennal segment bare above . . 7

7 Scutellum with long marginal bristles. Face smoothly rounded, with a moustache

of strong bristles extending up to antennae. Third antennal segment clavate,
with two-segmented style . . NUSA Walker (DASYTHRIX Loew) (p. 243)

Scutellum without marginal bristles ......... 8

8 Abdomen with a clump of strong bristles on first segment only. Moustache consist-

ing of a row of strong bristles along mouth margin, and a mass of soft, silky hairs
above these . . . PERASIS Hermann (SAUCROPOGON Hull) (p. 244)

- Abdomen with strong lateral bristles on several other segments beyond first.

Moustache consisting of hairs and bristles mingled. Often no pulvilli

GLYPHOTRICLIS Hermann

9 Pulvilli absent ; claws long and slender. Vein M^ parallel to outer end of discal

cell, and often in line with it ANYPODETUS Hermann

- Pulvilli present ............ 10

10 Costa extends round hind margin of wing at most as far as vein Cu + I A : axillary

cell has no vein along its outer margin ........ 1 1

Costa extends round axillary cell ......... 16

11 First posterior cell of wing open on margin. Palpi thicker at tip than at base . 12
First posterior cell of wing closed on or before wing-margin . . . . . 13

12 Hind femora slender LAPHYSTIA Loew

Hind femora distinctly swollen GERROLASIUS Hermann

13 Hind femora conspicuously swollen ......... 14

Hind femora not conspicuously swollen . . . . . . . . 15

14 Hind femora with strong, spine-bearing tubercles ventrally. Hind basitarus as long

as next three segments together. Proboscis as long as face, which is not swollen

HOPLISTOMERUS Macquart (p. 246)

Hind femora swollen, and with bristles, but not arising from tubercles. Hind
basitarsus not much longer than one segment. Proboscis shorter than face,
which is distinctly swollen TRICHARDIS Hermann (p. 247)

15 Palpi large and inflated, ovoid . . AFROMELITTODES Oldroyd & van Bruggen
Palpi pointed at tip some LAPHYSTIA Loew

1 6 Lower occiput with a backwardly-projecting flange. Palpi with only a single

segment (Ctenotini of Hull, 1962) . ........ 17

Lower occiput rounded, sometimes slightly bulbous, but never with a flange. Palpi

with two segments ........... 19

17 Claws noticeably blunt at tip .......... 18

Claws pointed. Abdomen club-shaped, becoming broader posteriorly. Legs

elongate, especially hind pair ....... LAMYRA Loew

1 8 Vein MZ cut short before it reaches the wing-margin. Ground colour of body

mostly obscurred by scaly hairs. Third antennal segment short, club-shaped,
little longer than first two segments together .... CTENOTA Loew

Vein Mz complete. Ground colour of body clearly visible between sparse hairs.
Third antennal segment elongate, 3-4 times as long as first two segments together

PARACTENOTA Engel ; STIPHROLAMYRA Engel

19 Veins closing discal cell parallel to, or even in line with vein, M^, which closes fourth

posterior cell ORTHOGONIS Hermann

These veins not parallel ; vein My distinctly curved ...... 20

226 H. OLDROYD

20 Face with prominent knob, at least on mouth-margin. Palpi leaf-like . . . 21

- Face without prominent knob. Palpi cylindrical ...... 23

21 Plump, hairy flies, mimicking carpenter-bees (Xylocopa). Legs short, densely

fringed with hairs . HYPERECHIA Schiner (p. 249) ; SYSTROPALPUS Hull

— Elongate, bare or only moderately hairy flies, with long, slender legs ... 22

22 Proboscis curved upwards like a scimitar . . . PROAGONISTES Loew (p. 251)
Proboscis straight ANDRENOSOMA Rondani (p. 255)

23 Metanotal callosities hairy or bristly

KATHARMA Oldroyd ; CENOCHROMYIA Hermanns

- Metanotal callosities bare .......... 24

24 Third antennal segment hairy dorsally. Hind femora swollen

LAXENECERA Macquart (p. 235)

- Third antennal segment without hairs dorsally. Hind femora slender

SMERYNGOLAPHRIA Hermann (p. 234)

LAPHRIA Meigen

Laphria Meigen, 1803 : 270. Type-species : Asilus gibbosus Linnaeus, by designation of
Latreille, 1810.

Although nearly thirty species of Laphria have been recorded from the Ethiopian
Region (Hull, 1962 : 323) this genus is not well represented in collections, either
from the equatorial forest or from the savannah areas. The flies are among the
most conspicuous of Asilidae, and easily attract the attention of collectors. To
some extent their apparent scarcity may be a reflection of their feeding habits,
which may take them in pursuit of wasps and bees high up in the trees.

The genus Laphria shares with Storthyngomerus and Dasyllina a characteristic
type of proboscis, laterally flattened, with an acute edge both dorsally and ventrally,
like a paper-knife. Presumably this has some relation to the nature of the prey,
though its significance is not obvious : other Laphriine robber-flies, notably the
genera Proagonistes and Hyperechia, will attack big Hymenoptera, but these have
a proboscis of quite a different shape.

No species of Laphria is among the material that I have received from the Pare
National du Garamba, but I have seen a number of species from other localities in
and around the Congo Basin. It seemed useful, therefore, to give details of these,
and to present a key that includes as many as possible of the species of Laphria
recorded from the Ethiopian Region.

KEY TO THE SPECIES OF LAPHRIA OF THE ETHIOPIAN REGION

1 Scutellum and at least posterior part of mesonotum conspicuously covered with

dense, recumbent, yellow hairs, which conceal ground colour .... 2

- Scutellum bare, or with scattered fine hairs only, not obscuring ground colour . 6

2 Legs distinctly patterned, or entirely red ........ 3

- Legs either entirely black, or indistinctly reddish at extreme base ... 4

3 Mesonotum entirely obscured by dense orange hairs, paler posteriorly

bella Loew, 1857 : 356

— Mesonotum mostly dull brown, with outline pattern only

lateralis Fabr., 1805 : 157 (bequaerti Bromley), 1947 : 409

2 See under Smeryngolaphria pallida.

ASILIDAE OF THE CONGO BASIN 227

4 A rather small species (14 mm). Basal half of wings distinctly hyaline. Moustache

mainly black . aureopilosa Ricardo, 1901 : 171 (variabitis Bromley), 1935 : 112

- Larger species, wings paler only at extreme base. Moustache partly yellow . . 5

5 All segments of abdomen densely covered with bright orange hairs, which obscure

the ground colour ...... auricorpus Hobby, 1948 : 140

Only first four segments of abdomen with golden hairs, which are sparse on disc and
do not obscure ground colour ; last three segments blackish with short black hairs

aurifer Ricardo, 1925 : 280 (p. 228)

6 Legs distinctly patterned in black and red, or entirely red or yellow ... 7

- Legs entirely black, or with indistinct reddish joints only . . . . . 15

7 Hairs of moustache mainly golden ......... 8

Hairs of moustache black, though there may be a tuft of yellow hairs at each side 9

8 Marginal bristles as well as short hairs of scutellum yellow

later alis Fabr., 1805 : 157 (bequaerti Bromley, 1935 : 409)
Marginal bristles and short hairs of scutellum black, or with only isolated yellow ones

contristans Hobby, 1948 : 139 (p. 228)

9 Moustache with some orange hairs or bristles as well as black ones . . . 10
Moustache without orange hairs ; with black bristles and white hairs, or entirely

black 12

10 Mesonotum bare, shining blue-black. Hind femora entirely orange. Smaller

species (15 mm) ........ ctenoventris sp. n. (p. 228)

- Mesonotum dulled with blackish or brownish tomentum. Hind femora not entirely

orange. Larger species (20-25 mm) • • • • • • • • II

11 Hind femora entirely black. Occiput with grey tomentum as well as black hairs.

Abdomen black with black hairs ..... her a Bromley, 1935 : 408

- Hind femora with basal half red, apical half black. Occiput with black hairs but

no grey tomentum. Abdomen black with orange hairs at tip

maynei Janssens 1953 : 207 (p. 230)

12 Abdomen reddish brown, first segment mostly obscure dark brown, median areas of

tergites with an obscure blackish brown stripe . schoutedeni Bromley, 1935 : 408

Abdomen black ............ 13

13 Dorsum of thorax and scutellum black with scattered golden hairs

tola Bromley, 1935 : 409 (p. 230)

- Dorsum of thorax and scutellum black, with short black hairs .... 14

14 Femora bright orange, rather swollen, only extreme base and an apical ring of

variable breadth are black. Thorax and abdomen smooth and shining metallic
blue ; black hairs not arising from obvious pits . flavipes Wiedemann, 1821 : 208
Fore and middle femore entirely black, slender ...... sp. n.

15 First three abdominal segments red, rest black ; abdomen rather club-shaped

carbonaria Ricardo, 1925 : 282 (consistens Bromley, 1935 : 405) (p. 228)

- Abdomen entirely black, or metallic blue-black . . . . . . . 17

1 6 Large black or metallic blue-black species, well over 20 mm long . 16
Length 10-20 mm ........... 19

17 Abdomen entirely black ....... 18

- Segments 2, 3 of abdomen each with a broad white band, half as long as segment,

narrowed or interrupted in middle. Wings pale at base luctuosa Macquart, 1847 : 37

1 8 Wings entirely black-brown. Metapleural bristles (just before halteres) entirely

black .... bipenicillata Bigot, 1891 : 370 (? metalli Walker, 1851 : 108)
Wings black-brown, but clear at base. Upper metapleural bristles black, lower

ones white ........ serpentina Bezzi, 1908 : 378

19 Mesonotum and scutellum black, with thin, sparse covering of yellow hairs

nigrescens Ricardo, 1925 : 279 (? fortipes Walker), 1857 : 128
Mesonotum and scutellum may have tomentum but no yellow hairs, except possibly

on scutellar margin ........... 2O

228 H. OLDROYD

20 Abdomen entirely black .... nigribimba Bromley, 1935 : 406 (p. 230)

- Abdomen black, with a pattern of grey tomentum ricardoi Bromley, 1935 : 4°7 (P- 23°)

Laphria aurifer Ricardo

(Text-fig. 3)
Laphria aurifer Ricardo, 1925 : 280.

LULUA : Kapanga, i <£, i <j>, X.IQ33 (F. G. Overlaet) ; MAYUMBE : Lundu, i $,
24.1.1925 (A. Cottart) ; HAUT-UELE : Yebo Moto, i <$, xii.i926 (A. Corbisier)
(MRAC).

Laphria carbonaria Ricardo

Laphria carbonaria Ricardo, 1925 : 282.
Laphria consistens Bromley, 1935 : 405.

LUKUGA : Bena Bendi, i ^ v. 1915 (R. Mayne) ; KIBALI : Watsa a Niangara,
i <£, vii.i920 (L. Burgeon) ; UELE : Bambesa, i $, 20.ix.i933 (H. J. Bredo} ; Bam-
besa, i $, 20.X.I953 (/. Leroy) ; STANLEYVILLE : Yangambi, i $, 1940, I.N.E.A.C.
(MRAC).

Laphria contristans Hobby
(Text-figs 4-6)

Laphria contristans Hobby, 1948 : 139.

UELE: Bambesa, i <$, i $, io.ix.i937 (/. Vrydagh) ; Bambesa, 10^, 2 $, ix.-x.i933
(H. J. Bredo) ; Bambesa, 5 <£, 3 $, x.ig33 (/. Leroy) ; 2 $, 14^.1933 (P. Henrard) ;
Bambesa, i <£, io.ix.i937 (/. Vrydagh) ; i <£, vi.i937 ; i <$, 10.1939 ; terr. Wamba,
Bayenge, i $, i2/22.viii.i956 (R. Castenau) ; Binza, i ^, 1954 (A. Bosma) ;
COQUILHATVILLE : Eala, i $, 1.1936,1 ^, i $, x.1936 (/. Ghesquiere) ; GABON :
Libreville, i ?, xi.i9i3 (Don G. Babault) (MRAC).

Laphria ctenoventris sp. n.

Allied to maynei Janssens and hera Bromley, but distinguished from both by
having the mesonotum and scutellum bare, shining blue-black, except for small
areas of brown tomentum at extreme sides. Hind femora typically entirely orange,
though experience in other species of Laphria suggests that some variation in leg-
colour is likely.

9 Head. Black. Frons with black hairs and bristles ; face with yellowish tomentum and
long, yellow hairs below antennae and at sides, but with moustache principally black ; about
three stout orange bristles ventrally on each side of moustache. Antennae black, hairs mixed
orange and black. Palpi and proboscis black with orange hairs. Beard orange, but post-
ocular and occipital hairs and bristles black.

Thorax. Mesonotum and scutellum bare, shining, blue-black ; brown tomentum only around
humeri and postalar calli, and before base of scutellum ; covered with short, black hairs and
with black bristles ; a number of weak humerals, one strong and one weak supra-alars, three
or four strong postalars, and six strong marginal scutellars : of the dorsocentrals only a weak

ASILIDAE OF THE CONGO BASIN

229

pair of prescutellars. Pleura covered with rather thin, brassy yellow tomentum ; hairs mostly
orange. Metapleural tuft mainly black.

Abdomen. Dorsum like thorax, shining blue-black, with short black hairs, and only a narrow

FIGS 3-9. Laphria spp. 3, L. aurifer, $ genitalia. 4, L. contristans, $ genitalia. 5, L.
contristans, head. 6, L. contristans, $ genitalia. 7, L. maynei, head. 8, L. nigribimba,
$ VIII sternite. 9, L. nigribimba, $ genitalia.

230 H. OLDROYD

band of brown tomentum at base of each segment. Venter similar, but with dull yellow-brown
band on hind margin of each segment.

Legs. Coxae and trochanters black. Rest of legs entirely bright orange, including all tarsal
segments, pulvilli, and bases of claws ; only apical three-quarters of claws black.

Wings. Almost uniformly smoky brown, indistinctly lighter towards base. Halteres
yellow, knob not darkened.

Length of body 15 mm ; of wing 12 mm.

Holotype $. LULUA : Kapanga, x.ig32 (F. G. Overlaet) (MRAC).

Paratypes. COQUILHATVILLE : Lukolela, i <j>, 2i.xii.ig2o (H. Schouteden) ;
' Congo Beige ', i ? (Don Gilson) (MRAC).

i $, Bomba (A. Henriori) differs in having yellow hairs on mesonotum and ven-
trally on abdomen.

Laphria tola Bromley

Laphria iola Bromley, 1935 : 409

LOMAMI : Lusuku, i £, xii.i93o (P. Quarre) ; LULUA : Kapanga, i <j>, x-ii.i93i/2
(G. F. Overlaet) (MRAC).

Laphria maynei Janssens

Laphria maynei Janssens, 1953 : 207.

Holotype $. STANLEYVILLE : Yangambi, 6.^.1952 (R. Mayne) (IRSNB).

UELE : Bambesa, i <j>, 9^.1938 (P. Henrard) ; SANKURU : Kondue, i <£, i ?
(Ed. Luja) ; LULUA : Kapanga, Itonde, i <£, ix.i932 (G. F. Overlaet} ; LEOPOLD-
VILLE : Lukolela, i $, xi.i934 (Dr Ledoux) (MRAC).

The hitherto undescribed male of this species closely resembles the female. It
is somewhat larger and has the hairs of the legs much longer, especially on ventral
surfaces of femora and tibiae.

Laphria nigribimba Bromley

(Text-figs 8-9)
Laphria nigribimba Bromley, 1935 : 406.

UELE : Bambesa, 2 $, I5.ix.ig33 (H. J. Bredo) ; Bambesa, i <j>, I5.ix.ig33 (/. V.
Levy) ; STANLEYVILLE : Basoko, Yacharo, i $, iv.ig4g (P. L. G. Benoit) ; UBANGI :
Binga, i $>, 5/12.^.1932 (H. J. Bredo) ; LULUA : Kasai, i $, 1928 (Dr. Walker) ;
COQUILHATVILLE : Flandria, i <J, iii.ig32 (R. P. Hulstaert) ; Eala, i <$, 2 $, 1932
(A. Corbisier) ; KUNDELUNGU : riv. Kalumbulwa, i $, 22.10.1951 (G. Marlier) ;
TSHUAPA : Bokuma, i $>, 1953 (R. P. Lootens) (MRAC).

Laphria ricardoi Bromley
Laphria ricardoi Bromley, 1935 : 407.

COQUILHATVILLE : Eala, i $, vi.ig32 (A. Corbisier) ; MAYUMBE : Zobe, i ^, i $,
i.igi6 (R. Mayne) ; UBANGI : Libenge, i £, i.ig27 (Leontovitch) ; TANGANIKA :

ASILIDAE OF THE CONGO BASIN 231

Kamens, 1400 m, riv. Kinga, I <£, 1.1953 (H. Bomans) ; LEOPOLDVILLE : Thysville
i $ (Dt. Houssiaux) (MRAC).

DASYLLINA Bromley

Dassylina Bromley, 1935 : 412 [lapsus for Dasyllina]. Type-species : Dasyllina fulvithorax
Bromley, loc. cit.

The original diagnosis of this genus makes it clear that the name indicates its
general resemblance to the American genus Dasyllis, and therefore the spelling
Dassylina is incorrect.

Dasyllina fulvithorax Bromley

Dasyllina fulvithorax Bromley, 1935 : 413.

BAS-CONGO : Kimwenza, i <£, i-iv.ig56 (R. P. Van Eyeri) (MRAC).

STORTHYNGOMERUS Hermann

Storthyngomerus Hermann, 1919 : 357, note ; Engel, 1924 : 106 ; Lindner, 1955 : 35> n§- 3-
Type-species : Dasypogon tridentatus Fabricus, by original designation.

Nusina Curran, 1927 : 7 ; Bromley, 1935 : 411. Type-species : Laphria dymes Walker, by
original designation.

Each of these two genera was described in a brief note, indicating the type-species,
but saying little about the limits of the genus, or of its affinities. The interpretation
that I have put upon Storthyngomerus is that given by Lindner (1955).

The type-specimen of Laphria dymes Walker is in the BMNH, so that Nusina
can be denned precisely. There can be no doubt that N. dymes is conspecific with
Storthyngomerus tridentatus (Fabr.), and that only females have so far been assigned
to these. On the other hand, Nusina aurea Bromley is known only from males,
and it seems clear from material in the BMNH that most of these are also con-
specific with dymes and tridentatus. Confusion is caused by the sexual dimorphism
of the legs, the femora being all orange in the females and heavily darkened in the
males. One male in the BMNH, with legs entirely orange like the females, has
distinctive genitalia and is clearly specifically distinct.

The fact that the orange-legged females are conspecific with the dark-legged
males and not with the orange-legged male is supported by the other characters
mentioned in the key below.

Laphria testacea Macquart was transferred to the genus Storthyngomerus by Engel
(1924), who did not give any reason for doing so, and it is difficult to understand
why. The description of testacea Macquart agrees quite well with specimens of
Trichardis testacea Hermann (q.v.), whereas the statement that the antennae are
' assez courtes ' does not suggest Storthyngomerus.

Lindner (1955) described Storthyngomerus minor, a very much smaller, black
species, which seems incongruous with the much bigger typical species. Yet it
shares the generic characters of Storthyngomerus which, besides the elongate antennae,
include the absence of scutellar bristles and the presence of spiny tubercles beneath
the hind femora, somewhat like those of Trichardis (q.v.).

232 H. OLDROYD

KEY TO THE SPECIES OF STORTHYNGOMERUS

1 Tiny species not more than 7 mm long. First antennal segment elongate, about as

long as height of face. Facial knob small . . minor Lindner (Kenya) (p. 234)

- Much longer specimens, 11-15 mm long. First antennal segment much shorter, only

twice as long as second, and much shorter than height of face. Facial knob short

but strongly prominent ........... 2

2 Proboscis slightly longer than height of head. Mesopleuron with well-marked

anterior border of white tomentum (Text-fig. 13). Femora orange in females, but
with broad black bands in males, at least on mid and hind femora. Genitalia of
male as in Text-fig, n ; lower forceps curled round tip of clasper, and without a
small apical spine, but with two long bristles . . tridentatus Fabricius (p. 232)

- Proboscis no more than three-quarters height of head. Mesopleuron with anterior

pale margin inconspicuous or absent (Text-fig. 14). Femora orange in males.
Genitalia of male as in Text-fig. 2 ; lower forceps not curled round tip of clasper,
and with a small apical spine but only one long bristle . toroensis sp. n. (p. 233)

Storthyngomerus tridentatus (Fabricius)
(Text-figs 10, n, 13)

FIGS 10-15. Storthyngomerus spp. 10, S.
clasper of $ ; 12, 5. toroensis, clasper of <$ ;
14, 5. toroensis, mesopleuron and spiracle ;

tridentatus, $ genitalia ; n, S. tridentatus,
13, S. tridentatus, mesopleuron and spiracle ;
15, 5. minor, $ genitalia.

ASILIDAE OF THE CONGO BASIN 233

Dasypogon tridentatus Fabricius, 1805 : 167
Laphria dymes Walker, 1849 : 377.
Nusina aurea Bromley, 1935 : 411.

GARAMBA NATIONAL PARK : 1949-52, i $ (no detailed locality or number) (IPNC).
KATANGA : Elisabeth ville, i $, 3O.ix.i926 (Dr. M. Bequaert) ; SANKURU : Komi,
2 $, v.xii.i93O (/. Ghesquiere) ; STANLEYVILLE : Stanleyville, i $, i3/23.viii.i928
(A. Collart) ; COQUILHATVILLE : Eala, i $, viii.35 (/. Ghesquiere) ; Eala, i <$,
iv.i933 (A. Corbisier) ; BAS-CONGO : Lemfu, i $, vi.i945 (Rev. L. De Beir) ;
TSUAPA : Ikela, i 9, 1955 (R. P. Lootens) ; KWANGO : Mwilambongo, i $, ix.i949
(Vanden Borght) ; UELE : Bambesa, i $, io.viii.i937 (/. Vridagh) (MRAC).

Storthyngomerus toroensis sp. n.

(Text-figs 12, 14)

The solitary known male of this species is superficially very similar to males of
S. tridentatus, but differs in having the legs entirely orange ; in S. tridentatus the
females have orange legs, but the males have at least the middle and hind femora
with broad black bands. The proboscis is distinctly shorter than in tridentatus,
being only three-quarters as long as the height of the head instead of rather longer
than this distance. In corroboration there are very clear differences in the male
genitalia. (Text-figs 12, 14.)

o* Head. Facial knob prominent, occupying lower half of face and separated by a distinct
hollow from the prominent tubercle on which the antennae stand. First antennal segment
twice as long as second (third segment broken off). General colour of head black. Frons,
vertex and upper occiput bare, shining black, with sparse black bristles. Face with yellowish
grey tomentum except for a bare patch on facial knob. Hairs yellow, more scaly on middle of
eye-margins. Stronger bristles of moustache black. Proboscis and palpi black : palpi with
black hairs, base of proboscis with longer white hairs. Behind the eyes a broad strip of yellowish
white tomentum, with black bristles on dorsal two-thirds, fine yellow hairs ventrally.

Thorax. Pronotum black with about eight strong black bristles. Mesonotum and scutellum
black, uniformly covered with short, fine, black, bristly hairs and short, golden, silky hairs.
Humeri with some longer black hairs, and prehumeral hollows with white tomentum ; one
notopleural and one supra-alar black bristle ; no long marginal scutellars. Pleura black,
covered with white tomentum except for mesopleuron, which has a large, bare, shining black
area almost reaching to anterior margin, and so with the anterior white margin much less
obvious than in S. tridentatus ; one black mesopleural bristle. Metapleural tuft before halteres
black, bristles strong and dense.

Abdomen. Dorsum with reduced first segment and entire second segment black, with short
black hairs ; second segment laterally with yellow hairs and a tuft of two or three strong orange
bristles. Rest of abdomen, including genitalia, orange, with yellow or pale yellow hairs.
Genitalia as in Text-fig. 12, distinct from those of <$ tridentatus in having lower forceps flattened
and acutely tipped, with one long bristle and a short, subapical spine. No dorsal process
curving round clasper. (Genitalia rotated through 180. °)

Legs. Trochanters black ; rest of legs entirely orange, with nearly all hairs and bristles
yellow, except for a few on tarsi and a large patch of black dorsally on femora (S. tridentatus <$
has yellow hairs even over the black femoral patches).

Wings. Infuscation— both staining and microtrichiae — almost uniform all over, without
clearer centres in basal, anal and axillary cells. Halteres orange.

Length of body 15 mm ; of wing 13 mm.

234

H. OLDROYD

Holotype <£. UGANDA : Southern Toro, Mbarara, Fort Portal Road, 3,800-
4,200 ft, 22-24. x. 1911 (-S- A. Neave) (BMNH).

Storthyngomerus minor Lindner

(Text-fig. 15)
Storthyngomerus minor Lindner, 1955: 35

Holotype <j>. TANGANYIKA : Dar-es-Salaam, xi.-xii.ig5i (E. Lindner) (SMNS).
KENYA : Diani Beach, vii.5i (N. L. H. Krauss) (BMNH).

SMERYNGOLAPHRIA Hermann

Smeryngolaphria Hermann, 1912 : 226 ; Bromley, 1935 : 410 ; Hull, 1962 : 331. Type-
species : Laphria melanura Wiedemann, 1828, by original designation.

Smeryngolaphria is a small and rather ill-defined genus from South and Central
America. Bromley (1935) considered that it should be extended to include the
Oriental species for which Hermann had erected the genera Anisosis and Orthogonis ;
and the BMNH collection contains a number of red-and-black Laphriine species
that were placed in Smeryngolaphria by Miss Ricardo.

Bromley also described Smeryngolaphria pallida from a single male from the
eastern Congo. A second specimen from the same area is before me, and there is
no doubt that it belongs to pallida Bromley. The generic assignment is less certain.

FIGS 16, 17. Smeryngolaphria pallida. 16, head ; 17, tarsus and claw.

ASILIDAE OF THE CONGO BASIN 235

According to Hull (1962), and to specimens in the BMNH, it seems at least equally
likely that it should be placed in the genus Cenochromyia Hermann (1912 : 115),
another genus of yellow or reddish Laphriine flies from New Guinea and adjacent
areas. It will not run to Smeryngolaphria in my key to African genera (Oldroyd,
1963 : n) because the metanotal slopes are hairy. In Hull's interpretation these
slopes should be bare in Smeryngolaphria, though Bromley considered that they
might be either hairy or bare. Bromley specifically mentions that they are hairy
in his species pallida.

A clear decision about the genus of this fly cannot be made in isolation, and will
have to await a more general study of the Laphriini.

Smeryngolaphria pallida Bromley

Smeryngolaphria pallida Bromley, 1935 : 410.

Holotype $. Kivu : Walikale, 1.1.1915 (/. Bequaert] (MCZH). W. Kivu :
Lubongola, pr. Shabunda, i <j>, 1939 (Dr. Hautman) (MRAC).

LAXENECERA Macquart

Laxenecera Macquart, 1838 : 77 ; Hermann, 1919 : 337-358. Type-species : Laxenecera
albibarbis Macquart (an Indian species), by designation of Hermann, 1919, loc. cit.

Acurana Walker, 1851 : 107. Type-species : Acurana sexfasciata Walker, monotypic.

Dyseris Loew, 1857 : 357 ; 1860 : 122. Type-species : Laxenecera andrenoides Macquart,
by designation of Loew, 1860 : 122.

Laxenecera is most easily recognized by the fairly long, narrow third antennal
segment, which bears a distinct fringe of hairs dorsally as well as ventrally.
Macquart says that the name expresses the hairiness of the antennae. Unfortunately
the antennae of Asilidae are prominent, and are easily destroyed in dried specimens,
and then it is necessary to rely upon a combination of other characters. Moreover,
dorsal hairs on this antennal segment also occur in some Hoplistomerus and
Trichardis.

Laxenecera has the proboscis triangular in cross-section, the palpi very slightly
clubbed at the tip, and usually the hind femora swollen with the hind tibiae corres-
pondingly curved. Many of the species look like small bees.

There is one widespread, common, and variable species (albicincta Loew), and a
number that are more locally distributed. There is considerable variation between
individuals, and this is one of the most difficult genera to present in the form of a
key to species. The male genitalia can be examined to some extent without dis-
section, and show some differences between species, but I have not found these
much use in identification. A confusing factor is the very great sexual dimorphism
that is generally present : e.g. in albicincta the male is a slender black fly, with
white hairs and no abdominal bands ; the female is a stout fly with rusty yellow
hairs and bristles, and conspicuous bands on the abdomen. The leg-pattern is the
most reliable way of associating males and females, but individual variation must
be allowed for.

236 H. OLDROYD

The two species originally included by Macquart were from India, but as we
know it today the genus is predominantly an African one. Loew erected the genus
Dyseris for some of the African species as a result of an error in the numbering of a
Plate in one of Macquart's works ; the error was pointed out by Hermann (1919).

KEY TO THE SPECIES OF LAXENECERA OF THE CONGO BASIN

1 Mesonotum with distinct long bristles, prominent in contrast with the much shorter

clothing hairs. Sides of abdominal tergites usually with strong bristles ... 2

- Mesonotum either furry, with bristles hidden among long, soft hairs, or else with

very short clothing hairs and almost no long bristles. Sides of abdomen usually
without strong bristles ........... 15

2 Legs partly orange or yellow, with extensive paler areas on tibiae or femora. . 3

- Legs almost entirely black, no more than faintly reddish on tibiae and at base of

hind femora ............ 8

3 Femora black, except perhaps for base and tip of hind femora .... 4

- Femora extensively orange, especially hind femora ...... 7

4 Abdomen clothed uniformly with yellow hairs, or with a few white ones at sides, but

no obvious posterior bands on abdominal segments . $ auricomata Hermann (p. 238)

- Abdomen with obvious posterior bands on each segment ..... 5

5 First three abdominal segments with broad posterior bands of dense whitish hairs;

other segments contrasting, almost black . . <$ abdominalis sp. n. (p. 238)

- All abdominal segments with posterior bands of pale tomentum and hairs . . 6

6 Only tarsi and dorsal surfaces of tibiae orange. Occipital hairs generally including

a black tuft. Wings lightly but uniformly smoky. Q* hind femora and tibiae
without conspicuous black ventral fringe rufltarsis Bezzi (funditor Curran) (p. 239)

- Fore and middle tibiae mainly orange. Occipital hairs generally entirely yellowish.

Wings strongly infuscated along veins. $ hind femora and tibiae swollen, and

with a conspicuous fringe of black hairs ventrally . . misetna sp. n. (p. 239)

7 Hind femora red at base, black apically, divided transversely pulchella sp. n. (p. 239)

- Hind femora with red and black horizontal stripes

albicincta Loew, undescribed form

8 Abdominal segments with a well-defined band of tomentum on posterior margin . 9

- Abdominal segments without a posterior band of tomentum, or with one at sides only 13

9 Scutellum with black hairs on disc, though marginals may be pale. Wings dark

brown ($). Middle legs with black hairs ventrally, white hairs dorsally (<$)

dimidiata Outran (p. 240)

- Scutellum with pale hairs and bristles. Wings pale . . . . . . 10

10 Hind femora and tibiae with conspicuously strong bristles, especially antero-apically n
Hind femora and tibiae with long, fine hairs and bristles, but not conspicuously

strong ones. ............ 12

11 Abdomen with dense golden tomentum on posterior bands, and almost entirely

covering first two or three segments .... francoisi sp. n. (p. 240)

- Abdomen with ashy grey tomentum on posterior margins, otherwise black-brown

9 albicincta Loew (p. 241)

12 Mesonotum with short, recumbent, golden hairs, almost lost in a mass of very long,

fine hairs, giving a bee-like appearance. Hind tibiae with a brush of long black
hairs ventrally ........ chrysonema sp. n. (p. 242)

- Mesonotum with recumbent golden hairs, but almost no long hairs or bristles.

Whole insect bare and rather deficient in long hairs and bristles sororcula Karsch

13 Short, recumbent hairs of abdomen yellow, as are lateral bristles. Hairs of head

and legs predominantly yellow . . . . . $ auricomata Hermann

- Short, recumbent hairs of abdomen almost entirely black, except on small lateral

ASILIDAE OF THE CONGO BASIN

237

grey triangles ; lateral bristles of abdomen black. Hairs of head and legs in
black or white clumps ........... 14

14 Pale hairs of head and thorax silvery white. Moustache predominantly black

<$ albicincta Loew
Pale hairs of head and thorax yellowish. Lower half of moustache yellowish ($),

or moustache entirely pale ($) . <$ $ albicincta Loew form stuhlmanni Roder

15 Abdominal segments without posterior bands, uniformly covered with yellow or

whitish hairs ........ 9 auricomata Hermann

Abdominal segments showing distinct posterior bands or lateral triangles . . 16

1 6 Mesonotum very bare, with a covering of recumbent black hairs mixed with scaly

golden ones, and with a very few long bristles, unusually slender. Mesopleuron
with a broad bare stripe reaching to dorsal margin . . . gymna sp. n. (p. 242)
Mesonotum furry, bee-like flies. . . . . . . . . . 17

FIGS 18-25. Laxenecera spp. 18, L. mollis, wing ; 19, L. andrenoides, wing ; 20, L.
engeli, wing ; 21, L. albicincta, <$ genitalia ; 22, L. andrenoides, $ genitalia ; 23, L.
auricomata, <$ genitalia ; 24, L. albicincta, maxilla and 2-segmented palpi ; 25, Ctenota
mollitrix, maxilla and i-segmented palp.

238 H. OLDROYD

17 Hairs of mesonotum (though not of pleura) entirely black in <J, partly grey in $.

Scutellum shining black with yellow tomentum at base, and hairs entirely black

chapini Curran (p. 243)

- Hairs of mesonotum entirely yellowish, or with a few black hairs only. Scutellum

with long yellow hairs ........... 18

1 8 Moustache, both male and female, with black hairs in lower part. Mesopleuron

mostly tomented, with only a small bare black area nigrociliata Hermann (p. 243)

- Moustache either yellow (<$) or pale yellowish ($), but without an obvious black tuft.

Mesopleuron shining black for about half its area, tomented on dorsal and posterior

fdasypoda Speiser
margins ..........< **1 Or

\scopifera Speiser

Laxenecera auricomata Hermann

(Text-fig. 23)
Laxenecera auricomata Hermann, 1919 : 351.

Kivu : Tshibinda, 3 $, i <j>, xii.i927 (Ch. Seydel) ; Muhunga, Tshibinda, i $,
xi.igsi ; ITURI : Luburo, i $, 1958 (Mme van Kiel] ; HAUT-UELE : Yebo Moto,
i <j>, 1926 (L. Burgeon] (MRAC). Also found in UGANDA.

Laxenecera abdominalis sp. n.

The male is distinguished from all other Laxenecera known to me by having the
first three abdominal segments mainly or entirely covered with whitish hairs, while
the rest of the abdomen has the hairs almost exclusively black.

o* Head. Clothed with golden hairs, except for a black tuft on each side of frons, close to
antennae, and a black tuft in middle of occipital margin. First antennal segment with yellow
hairs, other two with black hairs.

Thorax black. Ground colour of mesonotum and scutellum almost obscured by short,
recumbent, whitish or yellowish hairs. Bristles longer, fine and hair-like, black on disc of
mesonotum, whitish laterally and posteriorly, and on margin of scutellum. Pleura mostly
obscured by brownish tomentum. Mesopleuron shining black, with a vertical posterior band
of yellowish tomentum, and some black some yellow bristles.

Abdomen black, obscured by short, recumbent yellow hairs. On first segment and a broad
posterior band on second and third, these are silky white, but on anterior half of second and
third segments, and almost entirely on other segments, they are black. Venter black-brown
with mainly yellowish hairs.

Legs mainly black, but tibiae and tarsi of fore and middle legs are extensively orange, with
numerous yellow hairs and bristles. Hind femora black, with mainly black hairs, but with a
few white ones dorso-apically, hind tibiae with dense and moderately long white hairs dorsally
and ventrally ; hind tarsi with black hairs ventrally and a conspicuous fringe of dense silvery
ones dorsally.

Wings rather smoky towards tip, no special features.

Length of body 14 mm ; of wing 10 mm.

Holotype <£. TANGANIKA : Albertville, iii.iS-i.ig (MRAC).
Paratypes. Same data, 2 £ (MRAC).

ASILIDAE OF THE CONGO BASIN 239

Laxenecera rufitarsis Bezzi

Laxenecera rufitarsis Bezzi, 1908 : 377
Laxenecera funditor Curran, 1927 : 8.

URUNDI : Butetsi (Moso), i $, 27^.1950 ; Terr, de Rutana, colline Kisikara
(Bunyambo), alt. 1,600 m, i <j>, 2O.vi.i952 ; Bururi, 2,000-2,250 m, 7 <£, 7 <j>,
g.v.1949 ; Kitaba, 1,850 m, 2 J, io.vi.i949 (FJF).

Laxenecera misema sp. n.

Closely allied to rufitarsis Bezzi (funditor Curran), from which it is distinguished
in the male by the conspicuous fringes of the hind legs. In both sexes it is further
distinguished by having the fore and middle tibiae orange, without a black stripe,
and by having the occipital bristles all yellow, without a black tuft.

cJ Head with some black hairs on second and third segments, and on tips of palpi, otherwise
all hairs of head yellow.

Thorax. Mesonotum dull black, with narrow margin of yellowish tomentum at extreme
sides, and whitish tomentum anteriorly. Short hairs golden, and closely recumbent, longer
hairs and fine bristles mostly golden, but some black ones. Supra-alar and postalar bristles
strong, yellowish, postalar callus a little reddish. Pleura, except for the usual bare mesopleural
area, with brassy yellow tomentum and yellow hairs and bristles.

Abdomen black, each segment becoming dull reddish towards posterior margin. On hind
margin of each is a triangular patch of whitish hairs at each corner, extending towards middle
line ; adjacent to these is a band of yellow hairs, which cover whole of first and second tergites,
but the others have black hairs over most of disc.

Legs of male with conspicuously swollen hind femora, and stout, bowed hind tibiae, with a
brush of black hairs ventrally on apical half of femora and on full length of tibiae. Hind femora
black with red apex and base ; tibiae red ; both femora and tibiae obscured by short black
hairs. A few strong yellow bristles stand out among black hairs, especially on tibiae. Femora
and tibiae both with white hairs posteriorly. Hind tarsi orange, with dense tufts of hair on each
segment, silvery white on anterior face, black on posterior face. Fore and middle femora black
with red base and tip ; their tibiae and tarsi entirely orange.

Wings greyish, distinctly infuscated along veins.

Length of body 15 mm ; of wing n mm.

Holotype $. LOMAMI : Luputa (Bouvier) (MRAC).

Paratypes. i <$, i $, same data. One specimen captured from the wasp Bembex
(MRAC).

Laxenecera pulchella sp. n.

A variable species, recognized by the red antennae, swollen and mainly red hind
femora, and in particular by the grey band along the transverse suture of the thorax.
This species of north-eastern Africa is included for comparison with misema.

Head. Second and third antennal segments dull red, with black hairs ; first segment black,
with pale yellowish hairs. Palpi with some black hairs, otherwise hairs of head all pale yellowish,
or a little golden towards tips.

Thorax shining black, with a strip of grey tomentum along transverse suture, interrupted in
middle. Sides of mesonotum with tomentum, which may be grey, but in some specimens is

240 H. OLDROYD

golden yellow ; silky hairs and bristles may be pale yellowish or deep golden. Pleura evenly
covered with grey tomentum and with pale yellowish hairs and bristles.

Abdomen dorsally shining black with black hairs, and with broad posterior bands of grey
tomentum with white hairs. Sometimes ground colour under these bands is reddish, especially
at sides. In some specimens each segment also has a fore-border of grey tomentum. Strong
lateral bristles on anterior segments.

Legs. Fore and middle femora almost entirely black, with only extreme bases and tips
yellow ; fore and middle tibiae yellow dorsally, black ventrally, their tarsi orange. Hind
femora swollen, red, with a black area of variable extent at tip ; hind tibiae orange-yellow, a
little dusky on posterior face ; tarsi red. Hairs of legs white, and bristles pale yellowish, with
short black bristles ventrally on hind femora.

Wings uniformly faintly greyish, veins yellow towards base.

Length of body 13 mm ; of wing 10 mm.

Holotype <$. SOMALILAND : 30 m. S. of Shillawa, 23.11.53 (Greathead) (BMNH).

Paratypes. Same data, i $ ; KENYA : W. side of Turkwell Valley, 2 $, i.v.54
(Greathead) (BMNH).

Laxenecera dimidiata Curran

Laxenecera dimidiata Curran : 7.

ITURI : d'Obougena a Utike (Collart) ; Stanleyville area (Lang & Chapin) ;
KATANGA : Lukagu (de Witte] (MRAC).

Laxenecera francoisi sp. n.

In pattern rather like the female of albicincta, but distinguished by the more
golden yellow colour of the tomentum and by having the abdominal segments
liberally covered with yellow hairs, which almost entirely obscure the posterior
bands on the first two segments.

Head. Black, with black bristles on and and 3rd antennal segments and on palpi ; other-
wise all hairs yellow, or beard indistinctly white.

Thorax black. Mesonotum with thin golden tomentum, leaving a faint indication of a
divided median stripe. Hairs yellow, mixed with a few black ones ; long and short hairs not
greatly different in length. Scutellum with yellow hairs and numerous long, yellow marginal
bristles. Pleura shining black in part, otherwise tomentum, bristles and hairs yellow.

Abdomen dorsally shining black in ground colour, each segment with a broad posterior band
of golden tomentum, which extends forwards along side-margins. These yellow bands clothed
with golden bristles and hairs. On first two, or perhaps three, tergites the disc of the segment
is also covered with brownish yellow tomentum and clothed with golden hairs ; on other
segments tomentum more blackish, and there are more black hairs.

Legs mainly black. Fore and middle femora rather indistinctly yellow dorsally ; hind
femora red at extreme base. Hairs and bristles all yellowish.

Wings faintly smoky, more yellowish towards base.

Length of body 13-14 mm ; of wing 10 mm.

Holotype <$. BAS-CONGO : Tshela-Mata, 20.V.58 (Francois) (FJF).

Laxenecera albicincta Loew

(Text-figs 21, 24)

Laxenecera albicincta Loew, 1852 : 659.
Laxenecera apiformis Walker, 1855 : 571.

ASILIDAE OF THE CONGO BASIN 241

Laxenecera nigrocuprea Walker, 1855 : 572.
Dyseris zonata Loew, 1857 : 358
Laxenecera stuhlmanni v. Roder, 1893 : 205.
Laxenecera splendida Hermann, 1919 : 341.

The male and female of this species are normally so different that they might be
mistaken for quite distinct species. Typically the male is black, with almost
entirely black bristles, and no more than a trace of whitish bands laterally on the
abdomen. The female is a more bulky insect, with obvious bands of whitish grey
or yellowish grey on the abdomen, and with the hairs of the head, thorax and legs
predominantly rusty yellowish. In particular the postalar tuft is rusty yellowish,
and stands on a tawny callus.

The female has quite a close resemblance to the bee Megachile felina Gerst.

The variation in this species is most obvious in the male, where the presence of
any white or yellowish hairs is more conspicuous than it is in the female. Hermann
(1919) gave it as his opinion that stuhlmanni v. Roder was no more than a variety
of albicincta, distinguished from the typical form by the somewhat differently
coloured hairs, especially those of the legs. In Hermann's own variety splendida
there is less difference between the sexes than in typical albicinda, and some males
from the Congo Basin have taken on much of the superficial appearance of the
females, especially in the rusty yellow postalar bristles, and in the pale moustache.
Somewhat similar variation occurs in specimens from the Gold Coast collected by
Dr John Bowden. These males seem to have genitalia indistinguishable from
those of the typical form.

Laxenecera albicinda is common and widely distributed in eastern and southern
Africa, from ETHIOPIA to NATAL and ZULULAND, and round to the mouth of the
River Congo. Although it occurs round the fringes of the Congo Basin, and
occasionally on the river itself, it is not properly a forest species.

P.N.G., 2332 n/gd/4, i ?, 30.vii.i95i ; 585 i/a/M, i $, 7.^.1950 ; 473 Akam,
i ?, 3-V.I950 (IPNC).

BAS CONGO : Lemfu, 16^, 12 ?, 1.1915 (Rev. P. De Beit] ; Kisantu, 2<£, 29-3O.xii.
1953 (P. Basilewsky) ; RUANDA : Kagera, Gahinga, i $, 29.^.1937 (H. J. Bredo) ;
Rutshuru, i (£, 28.V.36 (L. Lippens) ; LUALA : Kapanga, i <?, iv.i932 (G. F.
Overlaet) : Magidi, i $, 1942 (Rev. P. Van Even) ; de Tenka a Dilolo, Km 109, i <£,
iv.i932 (Dr Ritschard) (MRAC).

KATANGA : Elisabeth ville, i <j>, 27.iv.i9i2 (M. Bequaert] ; i $, 1928-29 (P.
Quarre) ; ITURI I Arac-Aru, i $, vii.1952 (M. Winand] ; Jadotville, Numbi, 2^, 2 ?,
v.1957 (R. P. Th. de Caters] (MRAC).

URUNDI : Terr, de Bubanza, Colline Gihanga (Ruzizi), alt. 850 m, 3 <?, 3 £,
6.iii.i952 ; Gihanga, Plane de la Ruzizi, 850 m, 5 $, 3 $, 9.^.1932 ; Butetsi (Moso),

1 (J, 22^.1950 ; Ruyigi, 1,600 m, i <J, iii.i955 '> Mishiga, i <$, 15^.1957 ; Terr, de
Butana, Colline Ntangusa, Moso, alt. 1,350 m, i $, 2i.vi.i952 ; Kibunbu, 2,100 m,

2 ?, 25.vi.i95o (FJF).

MOZAMBIQUE : Chemba, 1931 (A Ravet] (MRAC).

242 H. OLDROYD

Laxenecera chrysonema sp. n.

A small, black species, with conspicuous golden hairs on all parts of the body,
distinguished from sororcula Karsch by the way in which the recumbent hairs of
the mesonotum are almost hidden in a mass of long, fine hairs.

(J Head. Pubescence long and golden. Antennae have golden hairs, which are mixed with
black on second and third segments ; a tuft of black hairs on occipital fringe ; palpi with
black hairs.

Thorax black with some greyish tomentum on pleura and sides of mesonotum. Mesonotum
with short, recumbent golden hairs, longer towards sides, mixed with very long, erect, fine
hairs which are yellow and black. Scutellum with yellow hairs on disc and long yellow marginal
bristles. Supra-alar tuft dense, yellow, but above and before wing-base there are several long
black bristles. Pleura with yellow hairs.

Abdomen black-brown. Dense yellow hairs form a fringe on sides, and on hind margins of
segments. Hairs on disc of each segment sparser, but mainly yellow.

Legs all black, with long yellow hairs and bristles. Venter of each tibiae had black hairs and
bristles and there are a few black bristles elsewhere on hind tibiae and tarsi.

Wings faintly and uniformly smoky.

Length of body 10 mm ; of wing 8 mm.

$ similar but less golden.

Holotype #. KATANGA : Lubumbashi, g.xii.23 (Seydel) (MRAC).

Paratypes. KATANGA : Lubumbashi, i $, 9.xii.23 (Seydel) ; Mwema, 2 <$
(Bayet) ; Nyamgwe, i $ (R. Mayne) (MRAC).

Laxenecera gymna sp. n.

A rather elongate and bare, black species, the mesonotum clothed only with very
short, recumbent hairs, and almost without strong bristles.

$ Head black with black bristles on occiput, ocellar tubercle and sides of frons. Moustache
with black bristles mingled with many white hairs, especially thick laterally. Palpi and pro-
boscis black ; palpi with stiff black bristles and fine whitish hairs. Antennae black, with short
but numerous black hairs and bristles ; third segment more than ij times as long as first two
together.

Thorax black. Mesonotum bare with only small prehumeral patches of grey tomentum ;
uniformly covered with short, curly black hairs, interspersed with short, silky yellow hairs.
Only supra-alar bristles moderately strong, mixed black and yellow ; other bristles few and
very slender, including slender yellow marginal scutellars. Pleura with whitish tomentum,
leaving a broad vertical band of shining black which reaches to upper margin of mesopleuron.
Metapleural hairs mainly black.

Abdomen shining black, with small triangles of greyish tomentum in posterior angles. Short
clothing hairs black, except for a few white or yellow ones on hind margins of segments, and
entirely covering sixth and seventh tergites. Venter with grown tomentum and longer yellowish
hairs.

Legs black, with yellow hairs and bristles.

Wings uniformly stained brownish. Halteres dark brown.

Length of body 15 mm ; of wing 12 mm.

Holotype $. KASAI : Terr, de Dekese, Itunda, x.5Q (F. J. Francois) (F.J.F.).

ASILIDAE OF THE CONGO BASIN 243

Laxenecera chapini Curran

Laxenecera chapini Curran, 1927 : 9.

P. N. DU GARAMBA : 422 I/a/3, i $, 17.^.1950 ; 717 I/a/3 amont. i <j>, 24.^1.1950;
895 Napokomweli, i $, iS.x.igso ; 1525, 11/91/4, i $, io.iv.i95i ; 1576, II/fb/4,

1 cJ, i <j>, ig.iv.igsi ; 1726, II/fd/i7, i <j>, 14^.1951 ; 1810, II/cf/i2, i <j>, 23.11.1951 ;
1915, II/fd/i8, i ?, 15.^.1951 ; 2015, II/ge/6, i ?, 29.^.1951 ; 2447, II/fd/n, i ?,
i8.ix.i95i ; 3429, II/fd/i8, 2 <£, 6^.1952 ; 4076, Mt Moyo, i <j>, 25.ix.ig52 (IPNC).

ITURI : Mongbwelu, Kilo, i <$, vii.i938 (Mme Scheitz) (MRAC).

Laxenecera nigrociliata Hermann

Laxenecera nigrociliata Hermann, 1919 : 356.

LOMAMI : Luputa, i $, 1.1935 (Dr Bouvier) ; RUANDA : Kibungu, 1^,1$,
x-xii.i937 (R. Verhulst) ; Mahamba, 1400 m, terr. Nyanza, i $, 13/15.1.53 (P.
Basilewsky) ; Gahiro, 1300 m, ten. Blumba, i $ (P. Basilewsky) ; URUNDI :
Maleka, i $, 3i.xii.i932 (L. Burgeon) ; Kivu : Costermansville, i <$, 1951 (H.
Bomans) ; KATANGA : Lugombo, i $, xii.i929 (Ch. Seydel) ; Elisabeth ville, 2 J,

2 $, ii-iv.i938 (De Loose) ; Lac Albert, Ishra, i<£, ix.i935, (H. J. Bredo) ; Kapanga,
i $, ix.i932 (F. G. Overlaet) ; KIBALI-!TURI : Djugu, i <$, I3.viii.i93i (Mme. L.
Lebrun) ; ITURI : Bambili, i <£, 5.xii.i9i3 (Z)r Rodhairi) ; BAS-CONGO : Mahagi,
Nirembe, i <^, ix.-x. 1935 (CA. Scops) (MRAC). URUNDI : Kisenyi, alt. 1350 m,
i cJ, 2 $, v.1955 ; 4 $ iv-v.1957 ; terr. de Muhinga, colline Nyabisudo, (Bugesera),
alt. 1200 m, i $, 3.vi.i952 (FJF).

NUSA Walker

Nusa Walker, 1851 : 105. Type-species : Nusa aequalis Walker, by designation of Hermann,

1912 : 239.
Dasythrix Loew, 1851 : 21. Loew, 1860 : 124. Type-species : Laphria (Dasythrix) inornata

Loew, by original designation.
Halictosoma Rondani, 1873 : 298. Type-species : Halictosoma puella Rondani, monotypic.

Hermann (1912) was misled by the original drawing of Nusa aequalis Walker into
accepting this as an earlier name for Andrenosoma Rondani, but Ricardo (1927)
after studying the type in the BMNH, rightly removed Nusa into synonymy with
Dasythrix Loew, 1851, over which it was shown to have three months' priority.

Loew divided his genus Dasythrix into two sections, making inornata, a South
American species, type of the first section, and infumata, a South African species,
type of the second. Loew gave no distinctions between his two sections, and I
have not seen the S. American inornata, which clearly must be the type of any
restricted genus Dasythrix. Nusa aequalis Walker is from India, and is congeneric
with the African species. Hence, if at a later date it should prove that Loew's
two sections are not congeneric with each other, section i, type inornata, would
retain the name Dasythrix, and section 2 would still be called Nusa.

Ricardo (1925 : 278) described 'Nusa africana <$, Hermann in litt., sp. n.', and
stated that this species was congeneric with Andrenosoma boranica Corti from East

244 H- OLDROYD

Africa, accepting at that time Hermann's view of the synonymy between Nusa
and Andrenosoma. Later (1927 : 206), when she realised the true identity of
Nusa Walker, Miss Ricardo renamed her species Andrenosoma africana, but examina-
tion of the series in the BMNH, including the types, shows that even this is in-
correct. Nusa africana Ric. belongs to the genus Proagonistes , and is not congeneric
with boranica Corti, which is a true Andrenosoma.

Nusa, as correctly denned, is distinctive in appearance. The wing-margin is
membranous shortly beyond R^, the first and fourth posterior cells typically closed
with a long, or very long stalk, and the veins tending to fade before reaching the
wing-margin. The characteristic appearance of flies of this genus is enhanced by
the stiff bristles on all parts of the body, including the head.

Species have been recorded in the genus Dasythrix from all the zoogeographical
regions except the Nearctic, but it is by no means certain that they all belong
properly in Nusa. Among the African species there is extensive sexual dimorphism,
the males tending to be distinctly blacker than the females, and with black hairs
everywhere instead of white ones. There is, however, variation in this, and among
a complex of specimens there are many males with black moustache and some with
the moustache white, though no clear differences can be seen in the genitalia. It
seems likely that Loew's three species from the Cape — infumata <$, brachyptera $
and stenura $ — may be synonyms, but this cannot be determined until types can
be examined.

No species of Nusa has yet been recorded from the Pare national du Garamba.
The genus is unlikely to occur in the forested areas, but may be found in the savanna
fringe close to the border with the Sudan. Besides the S. African infumata-com-
plex, mentioned above, which extends as far north as the Katanga, there are five
other described species from the Ethiopian Region : nigrapex Bigot from Natal ;
dispar Gerstaecker from Zanzibar ; vittipes Bezzi from Somaliland ; ruficornis
Wulp from South Yemen and albicans Engel from Rhodesia.

PERASIS Hermann

Perasis Hermann, 1905 : 37 ; Engel, 1924 : 104-5. Type-species : Perasis sareptana Hermann,

1905, monotypic.
Saucropogon Hull, 1962 : 103. Type-species : Perasis transvaalensis Ricardo, 1925, by original

designation.

This genus was clearly defined by Hermann, but has been misinterpreted by Hull
(1962 : 93), who removed the S. African Perasis transvaalensis Ricardo to be the
type of his new genus Saucropogon (1962 : 103). Hull saw the type and other
specimens of transvaalensis in the BMNH, and he also gives drawings of sareptana
Hermann, though he did not see this species in the BMNH, where it is not represented.

The characters given by Hull (1962 : 103) to define Saucropogon, and to differ-
entiate it from Perasis, concern mainly the wing-venation and the head-structure,
and in fact are the very characters by which Hermann himself defined Perasis
(1905 : 37). Hermann's fig. 28 shows the venation that Hull says is characteristic

ASILIDAE OF THE CONGO BASIN 245

of Saucropogon. It seems evident that Hull has based his interpretation of Perasis
on specimens that were not P. sareptana Hermann, since, among other things, he
says that they have lateral bristles on all the abdominal segments, whereas
Hermann's Perasis has bristles on the first segment only.

The type material of transvaalensis Ricardo agrees with Hermann's definition of
Perasis, and must therefore revert to that genus. This makes Saucropogon Hull,
type transvaalensis Ricardo, an absolute synonym of Perasis Hermann.

The species of Perasis are few and poorly known, scattered from Transcaspia
across North Africa. Hermann (1920 : 177) gave a list of four names in addition
to the type-species : postica Becker and violacea Becker, both from Algeria ;
Dasypogon maura Macquart from Oran ; and Perasis meridionalis Hermann from
the Transvaal. Engel (1925 : 104-5) explained that this last name was a nomen
nudum, no description having been published, and that it appeared to refer to
specimens in Hermann's collection which at the time Engel saw them bore the
manuscript name capensis. In the same year Ricardo (1925 : 245) described
Perasis transvaalensis from the Transvaal, and also recorded meridionalis Hermann
as a nomen nudum.

The theory that Dasypogon maura Macquart, 1849, *s really a Perasis appears to
come from Hermann (1920 : 177) but he gives no reasons for believing this. Bigot
(1878 : 221), in whose collection the type was deposited, thought that it might
belong to Habropogon.

No Perasis is known to me from within the boundaries of the Pare National du
Garamba, but the BMNH possesses a specimen from the Bunyoro district of Uganda
that is specifically distinct from transvaalensis, and Monsieur Francois collected a
number of the same species from Urundi.

Perasis carpenteri sp. n.

Closely allied to Perasis transvaalensis Ricardo, but with several constant colour-
differences. Most obviously, the pleura are uniformly covered with yellow-grey
tomentum over a black and red ground colour ; whereas the pleura of transvaalensis
are distinctly striped, with the anterior-facing areas of mesopleuron and sterno-
pleuron shining black through thin brown tomentum, and the posterior-facing
areas covered with thick yellow-grey tomentum. In carpenteri the whitish hind
margins of the abdominal segments are uniform in breadth, whereas in transvaalensis
they are expanded laterally and constricted in the middle.

$ Head black. Frons and face with a dense tomentum or pile, which is white when the light
falls from one direction. Moustache limited to about ten strong black bristles on mouth-
margin and a few scanty hairs above this ; most of face with tomentum only, no strong hairs
or bristles. A few white hairs in beard, otherwise hairs and bristles of head are black. Anten-
nae black or a little reddish, palpi and proboscis black, all with black bristles.

Thorax. Mesonotum black, reddish only on humeri and postalar calli. These areas have
some whitish tomentum, which spreads across hind margin of mesonotum, immediately before
scutellum. Mesonotum otherwise with very short, adpressed, dark brown hairs ; scutellum,
in contrast, with yellowish adpressed hairs. Pleura reddish in ground colour, but with a big
black area anteriorly on meso- and sternopleura ; the whole pleural area covered uniformly

246 H. OLDROYD

with yellowish grey tomentum, without the rather bare vertical stripe that is seen in trans-
vaalensis. Metapleural tuft of black bristles.

Abdomen. Dorsum dull black, obscured by very short, adpressed, black hairs. Each
segment posteriorly with a narrow band of uniform width, reddish in ground colour, with whitish
tomentum. Venter similar, black hairs rather longer.

Legs. Coxae reddish in ground colour, with pale yellowish hairs. Femora black with short
black hairs ; tibiae and tarsi dull reddish, covered with yellowish white hairs, but all leg-
bristles black.

Wings. Veins black, not partly yellowish as in tmnsvaalensis. Fairly uniformly stained
brown, only slightly paler towards wing-margin. Venation as in transvaalensis, but with stalks
of closed cells a little longer.

Length. Body 13 mm ; wing n mm.

$ Generally similar, but with a few differences. Frons and face with more yellowish white
hairs, especially in space between membrane and bases of antennae. Mesonotum covered with
adpressed yellowish grey hairs like those of scutellum, and with a very narrow median line and a
pair of postscutellar spots formed from dark brown hairs. Femora dull reddish in ground
colour and covered with adpressed pale hairs instead of dark.

Holotype^. UGANDA : Bunyoro, v.igaS (G. D. H. Carpenter] (BMNH).
Paratypes. URUNDI : Kinanga, Ruzizi, 3 $, 8 <j>, xi.igsi (F. J. Francois] (FJF).

HOPLISTOMERUS Macquart

Hoplistomerus Macquart, 1838 : 59, Oldroyd, 1940 : 307. Type-species : Laphria serripes
Fabricius, 1805, monotypic.

In 1940 (loc. cit.) I published a synopsis of the species of this genus, demonstrating
how the genus extends over the Ethiopian Region, with the exception of the
equatorial forest. A miscellany of species exists in eastern Africa, from Lake Nyasa
to the Red Sea, and from this nucleus one species, H. serripes Fabricius, has spread
through the northern savannas to the Gambia, while another, H. nobilis Loew, goes
southwards to the Cape and S.-W. Africa.

The presence of a species of Hoplistomerus in the Pare National du Garamba is
therefore rather surprising, until one looks at the description of the localities con-
cerned : 483 — savane arborescente, sur les herbes ; 529 — galerie forestiere seche ;
3282 — savane arborescente ; 3298 — herbacees sur le pourtour d'une prairie. In
localities such as these a savanna fauna is able to establish itself within the confines
of the Pare.

Hoplistomerus garambensis sp. n.

(Text-figs 26, 27)

Most closely resembles H. erythropus Bezzi, from Somalia, from which species it
differs in having black stripes on the femora ; in the abdominal colouring, which is
more normal for the genus than it is in erythropus ; and in the male genitalia. (Text-
figs 26, 27).

$ Head. Black, obscured by thick, yellowish tomentum, except for a bare patch of variable
extent in centre of face. Hairs and bristles all yellow or whitish. Antennae blackish or dull
reddish, with yellowish hairs. Palpi pointed and rather swollen, with hairs mainly black, some

ASILIDAE OF THE CONGO BASIN 247

yellow ; proboscis black with silvery hairs at base (part of beard) and deep yellow ones at apex.
Beard silvery, occipital bristles yellow.

Thorax black. Mesonotum heavily punctured, with little or no tomentum, but covered with
golden yellow hairs, white at sides ; a row of 5-6 supra-alar and one postalar bristles yellow or
whitish, otherwise dorsum without bristles. Scutellum covered with yellow tomentum and
very short, golden yellow hairs ; a deep, transverse groove is bare, shining black ; no marginal
bristles. Pleura black with thick white tomentum ; meso- and sternopleura with a large shining
black area.

Abdomen black^ Each segment narrowly shining black basally, otherwise covered with long,
adpressed, curled, golden yellow hairs. Laterally a patch of these adpressed hairs is silvery
white. First five complete segments each has 1-4 strong bristles laterally. Venter rather bare,
black-brown, with thin grey tomentum and sparse pale hairs. Male genitalia as in Text-fig. 27.

Legs. Coxae like pleura, black with thick white tomentum and white hairs. Trochanters,
femora, tibiae and tarsi orange, only femora with dorsal black stripe over middle part of their
length. Short clothing hairs white, even on tarsi ; bristles yellow. Ventrally on hind femora
the tubercles as usual in Hoplistomerus.

Wings. Venation normal for genus. Marginal cell closed on margin, vein Rz+3 recurrent.
First and fourth posterior cells closed with short stalk. Dark microtrichiae form the usual two
cross-bands across base of discal cell, and from radial fork to apex of wing, the two united
posteriorly. Veins blackish brown in dark areas, orange in clear patches.

Length of body 13 mm ; of wing n mm.

Holotype £. P.N.G., 3282, Ppk. i4/g/2, 4.^.1952 (IPNC).

Paratypes. Same data as holotype, i $ ; 483. I/a/i. i ?, 5^.1950 ; 422,
I /a/3. i <j>, I7.iv.i95o ; 529 Akam, 19^.1950 (IPNC).

TRICHARDIS Hermann

Trichardis Hermann, 1906 : 137 ; 1920 : 177. Type-species : Laphria testacea Macquart,

1838 ( = Trichardis testacea Hermann, 1906), by designation of Hermann, 1923.3
Strobilothrix Becker, 1907 : 42. Type-species : Strobilothrix albipila Becker, 1907, monotypic.

Closely allied to Hoplistomerus Macquart, and not easy to separate clearly from
that genus. Hermann's first two species, testacea and picta, were distinguished by
the absence of the conspicuous wart-like excrescences beneath the hind femora, but
in other species which he later assigned to Trichardis the ventral spines arise from
smaller, but distinct protuberances. As I recorded in 1940 (p. 310, note), the late
Dr S. W. Bromley considered that Trichardis was only a subgenus of Hoplistomerus,
but I am still not sure that I agree with this view. In general facies the two genera
seem to be distinguishable, Hoplistomerus being more robust. Probably they can
be formally separated by the different relative length of the first segment of the
hind tarsus, as well as by the male genitalia, Hoplistomerus having the claspers
prominent and usually hooked.

Since Trichardis does not appear to penetrate into the Congo Basin proper, this
is hardly the place in which to attempt a detailed study of the genus, but one new
species from the south-eastern uplands may be noted.

3 Engel (1924 : 106) stated categorically that Laphria testacea Macquart was a Storthyngomems, and
was quite different from Trichardis testacea Hermann. This is not in agreement with Macquart's
original description, in which the antennae are described as 'assez courtes'. Dr. L. Tsacas has very
kindly examined Macquart's type in Paris and confirms: (a) that it is definitely a Trichardis and not a
Storthyngomerus ; and (b) that it agrees with the description of Trichardis testacea Hermann.

248

H. OLDROYD

Trichardis katangaensis sp. n.

(Text-fig. 29)

FIGS 26-32. 26-27, Hoplistomerus spp., <$ genitalia. 26, H. erythropus ; 27, H. garam-
bensis. 28-29, Trichardis spp., left mesopleuron. 28, T. cribrata ; 29, T. katangaensis.
30-32, Andrenosoma spp. 30, ^4. complexa, proboscis and palp ; 31, A. boranica,
proboscis and palp ; 32, A. complexa, male genitalia.

ASILIDAE OF THE CONGO BASIN 249

Very closely resembling T. cribrata Loew, but distinguished by the much larger
bare area of the mesopleuron (Text-figs 28, 29), and by the larger and more con-
spicuous white patches in the posterior angles of the abdominal tergites.

<J$ Head. Black with thin white tomentum. Four strong black ocellar bristles ; sides of
frons with one or two short black bristles, but also softer white ones. Moustache consisting of
a few black bristles on mouth-margin, and longer, silvery hairs and fine bristles rooflike above
these, extending at sides almost up to bases of antennae. Centre of face with some fine black
hairs. Antennae may be entirely black, or more or less reddish, especially first two segments.
Proboscis and palpi black, but with hairs mainly silvery. Fine, silvery hairs in beard and over
occiput, but postocular bristles strong and black.

Thorax. Mesonotum shining black, with little tomentum except for a white patch adjoining
humeri, and a narrow, transverse band immediately before scutellum. Covered uniformly with
short, adpressed, golden yellow hairs, with short, erect, fine, black hairs distributed among
them. Scutellum with very fine black marginal bristles. Pleura heavily covered with white
tomentum, only mesopleuron with a large, shining black area (Text-fig. 29).

Abdomen shining dark red-black, covered with rather sparse, silky silvery hairs, each hair
arising from a definite puncture. A broad posterior band of white tomentum on each segment
is broadly interrupted, or reduced to a pair of widely separated lateral patches. Venter dark
reddish, with white hind margins and whitish hairs.

Legs. Hind femora markedly swollen, and with considerable tubercles. Femora shining
black ; trochanters, tibiae and tarsi dull reddish. Legs covered with silvery hairs ; bristles
mostly yellow, but black ones more numerous on tarsi.

Wings almost uniformly pale greyish, no obvious pattern. Marginal cell closed on margin,
vein 7?2+3 recurrent ; first and fourth posterior cells closed, with short stalk.

Length of body 10 mm ; of wing 8 mm.

Holotype <$. LULU A : Kapanga, X.IQ32 (F. G. Overlaet) (MRAC).

Paratypes. KATANGA : Elisabeth ville, i $, xi.ign (Miss. Agric.) ; TANGAN-
IKA : Sunkutu, 1140 m, Km 96, Rte Pepa-Moliro, i $, xii.1953 (H. Romans)
(MRAC).

HYPERECHIA Schiner

Hyperechia Schiner, 1966 : 673 ; Marshall, 1902 : 287-584 ; Griinberg, 1907 : 515-524 ;
Thorpe, 1927 : 177-185 ; Lamborn, 1927 : 44-47 ; Engel, 1929 : 147-162 (larva) ; van
Bruggen, 1962 : 313-317. Type-species : Laphria xylocopiformis Walker (an Indian species),
by original designation.

Flies of this genus are well-known because of their close resemblance to Xylocopid
bees. They are short and broad, with strong legs fringed with long pile, and with
wings broad basally, pointed at apex. The head is broad, the third antennal
segment long and clavate. The flattened, leaf-like palpi are characteristic of those
genera which Hull (1962 : 349) unites into the tribe Andrenosomini. The genus
Hyperechia is characterized by general appearance rather than by any diagnostic
character. Systropalpus Hull (1962 : 355), mentioned in the key to genera, is based
on a single male from Ethiopia, which I have not seen.

The various species differ in the distribution of white, yellowish or reddish pile
on the mesonotum, scutellum, abdominal segments 1-3, and legs. Several species
bear a startlingly close resemblance to particular species of Xylocopa, and to this

250 H. OLDROYD

extent they may be said to mimic the bees, especially as the larva of the fly lives in
the burrows of the bee, and the adult flies are found in association with the adult
bees. Yet Hyperechia adults are often taken in association with Xylocopa other
than the species to which they bear a mimetic resemblance. An example of this
occurs in the collections from the Musee du Congo, where a specimen of Xylocopa
flavorufa Deg. from Abok in the Ituri is labelled as having been taken in association
with Hyperechia imitator Grim. ; but the species of Hyperechia which imitates this
species of Xylocopa is H. bomboides Loew, which occurs at Tchad and similar drier
regions.

The following key is presented as an interim guide to the species, more particularly
those of the Congo Basin, or which might occur there.

KEY TO SPECIES OF HYPERECHIA

1 Sides of abdomen with pale yellowish or white fringes ...... 2

- Sides of abdomen with black fringes, at least on posterior segments ... 4

2 Mesonotum and scutellum with black hairs only

consimilis Wood, 1874 : 158 (usambarae Lichtw., 1907 : 85)

- Pale hairs on, or just before, scutellum ........ 3

3 A conspicuous tuft of pale hairs on scutellum, with black marginal bristles

floccosa Bezzi, 1908 : 377 (p. 251)

- A transverse band of pale hairs just before scutellum, but scutellum itself has black

hairs, and very strong black marginal bristles. nigrita Griinberg, 1907 : 520 (p. 251)

4 Thorax and abdomen both uniformly black, with black hairs. Female with white

fringe on fore tibia, male with black fringe . imitator Griinberg 1907 : 522 (p. 251)
Thorax, or abdomen, or both with conspicuous white, yellow or red-brown hairs . 5

5 Thorax and abdomen both with some yellow hairs ...... 6

Either thorax or abdomen with yellow hairs, but not both (except in marshalli, where

the first abdominal segment has a few sparse yellow hairs) .... 8

6 Thorax with yellow hairs on posterior margin, and segments 1-3 of abdomen also

with yellow hairs. . nigripennis Wied., 1830 : 646 (albifasciata End., 1930 : 69)

- Thorax with yellow hairs, but only segments i or 1-2 of abdomen with yellow hairs 7

7 Mesonotum and scutellum entirely covered with yellow hairs. Wings dark brown,

with base conspicuously paler .... hirtipes Fabricius, 1805 : 158

- Only posterior third or quarter of mesonotum with yellow hairs. Wings uniformly

dark brown, not obviously paler basally

bifasciata Griinberg, 1907 : 579 (pellitiventris End., 1930 : 68)4 (p. 250)

8 Hairs of mesonotum uniformly pale ..... bomboides Loew, 1851 : 21

- Thorax black with some yellow or red-brown hairs

marshalli Austen, 1902 : 341 (fuelleborni Griinberg, 1907 : 521)

Hyperechia bifasciata Griinberg

Hyperechia bifasciata Griinberg, 1907 : 519.

LULUA : Luluabourg, i $ (P. Callewaert) ; Kapanga, 2 $, V.IQ33 (G. F. Overlaet) ;
Kanzenza, i $, 1932 (R. P. Lefebure). KATANGA : du lac Moero au lac Bengwelo,
i c£ (Dr Gheral) ; Kiambi, i §, 5-15^.1931 (G. F. de Witte) ; Elisabethville, i $,
25.4.27 (Dr M. Bequaert). SANKURU : Tschombe-Ste Marie, i $, 1948 (Rev. P.
Gustave) ; Komi, i $, vii.i928, Lodja (/. Ghesquiere) ; Mukabe-kesari, i $, 1939

4 bifasciata and pellitiventris may be distinguishable by small genital differences.

ASILIDAE OF THE CONGO BASIN 251

(R. P. de Donckere). TANGANIKA : Kiambi, i <j>, 23.^.1931 (G. F. de Witte).
ITURI : Bunia, 2 ?, vii.1937 (H. J. Bredo) (MRAC).
URUNDI : Kitega, i <j>, 18.1.52 ; i <$, 1.1953 (FJF).

Hyperechia floccosa Bezzi

Hyperechia floccosa Bezzi, 1908 : 377.

LULUA : Kapanga, i <£, v.1933 (G. F. Overlaet). BAS-CONGO : Kisantu, i <$,
29-30. xii. 1952 (P. Basilewsky} ; Matadi a Leopoldville, i $, n.xi.i936 (/. Vrydagh).
Kunkungum N'Kele, i $, i $, i6.viii.i944 (C. Schoutederi) (MRAC).

Hyperechia imitator Griinberg
Hyperechia imitator Griinberg, 1907 : 522.

BAS-CONGO : Moanda, i $, 1933 (R. P. Bittremieux) ; Kisantu, i 9, 29~3O.xii.i952
(P. Basilewsky) ; BOLOBO : Makamndelu, N'Kele, 2 $, 1938 (Dr Schoutederi) ;
Bumbuli, i <$, 1915 (R. Mayne). ITURI : Abok, Lundigi, i 9» T928 (Ch. Scops)
(MRAC).

Hyperechia nigrita Grunberg

Hyperechia nigrita Grunberg, 1907 : 520.

BAS-CONGO : Kimwenza, i <£, i.iv.i956 (R. P. van Eyen) ; Kisantu, i $ (Rev.
P. Regnier). KASAI : Luebo, i $, iii.i93i (/. P. Colin). UELE : Bambesa, i $
io.iv.i937 (/. Vrydagh) (MRAC).

PROAGONISTES Loew

Proagonistes Loew, 1857 : 362, 367 ; 1860 : 170 ; Bromley, 1930 : 209-224. Type-species :
Proagonistes validus Loew, monotypic.

These large, aggressive-looking robber-flies are distinguished by their powerful,
sickle-like proboscis, usually upturned at the tip, and by their bare, elongate appear-
ance, mimicking some of the bigger wasps, especially Psammocharid wasps of the
genus Salius.

Neave (quoted by Bromley, 1930 : 213) associated flies of this genus with forest,
or at least with woodland, and wrote : ' I have never found them except in associa-
tion with trees.' Specimens from the Congo Basin suggest that Proagonistes is not
typical of high forest, but occurs along the great rivers, on the forest fringe, and in
savanna woodland.

The genus is quite closely related to Andrenosoma Rondani, and it is not easy to
indicate constant differences between the two genera, though each has a recog-
nisable fades. Proagonistes is more elongate, with long, slender legs, and cylindrical
abdomen which is scarcely constricted basally. In the wing the first posterior cell
is wide open, with veins R$ and MI parallel, or even diverging slightly. Andreno-
soma is typically more bee-like, the abdomen broader, and often constricted basally ;
the hind legs stouter, with distinct long hairs on tibiae and tarsi in addition to stout

252

H. OLDROYD

bristles ; and in the wing the first posterior cell is narrowed, often closed and
stalked.

Bromley (1930 : 209-224) published a revision of the genus Proagonistes, to which
may be added two species of hitherto doubtful affinities : Nusa africana Ricardo,
and Proagonistes igniferum Engel and Cuthbertson, both of which have at some
time or other been assigned to Andrenosoma. One new species from the Congo
Basin is described in the present paper.

The following key is modified from Bromley (1930 : 214), and includes all the
African species.

KEY TO AFRICAN SPECIES OF PROAGONISTES

i Wings orange, apex broadly brownish . . apicalis (Curran), 1927 : 6 (p. 254)

- Wings without noticeably darker tip, usually completely brown, or with clearer

centres to some cells 2

FIGS 33-36. Proagonistes spp., $ genitalia. 33, P. lampyroides ; 34, P. neavei
35, P. athletes ; 36, P. rufibarbis.

ASILIDAE OF THE CONGO BASIN 253

2 Hind tarsi with fine blackish hairs, causing them to appear black dorsally. Facial

and pleural hairs deep yellow or orange. Abdomen deep purplish brown with
reddish hairs ...... leoninus Bromley, 1930 : 216 (p. 254)

Hind tarsi with orange hairs only ......... 3

3 Scutellum uniformly black .......... 4

Scutellum not uniformly black .......... 16

4 Hind femora reddish or yellowish ......... 5

Hind femora more or less black ......... n

5 Vestiture of head mostly black .......... 6

Vestiture of head reddish or yellowish ........ 7

6 Beard white. Antennae and facial knob black. Abdomen reddish

africana Ricardo, 1929 : 278
Beard black. Antennae and facial knob red. Abdomen black

pliomelas Speiser, 1907 : 358 (p. 255)

7 Moustache long and thick, straw-yellow. First two segments of abdomen with fine

white hairs at sides ...... mystaceus Bromley, 1930 : 218

Moustache reddish ............ 8

8 Thorax with bristles mostly reddish .... vulpinus Bromley, 1930 : 219
Thorax with bristles black .......... 9

9 Huge species, about 45 mm in length, hind legs extremely long (Madagascar)

gigantipes Bromley, 1930 : 219
Species about 30 mm in length (West Africa) . . . . . . . 10

10 Occipital bristles black. Thorax black, but with slight brassy tinge, and covered

with dark tawny down. ...... ufens Walker, 1849 : 372

Occipital bristles reddish ; thorax black . rufibarbis (Fabricius), 1805 : 157 (p. 255)

11 Fore coxae with black hairs, beard black, or with some reddish hairs intermingled 12
Fore coxae with whitish hairs .......... 13

12 Small species (about 27-28 mm long) ; beard black ; femora black with extreme

apex reddish ....... praeceps Walker, 1855 : 542

Huge species (over 40 mm long) ; beard black with some of the hairs reddish at
base ; hind femora black, apically reddish, more extensively reddish below than
above ........ saliodes Bromley, 1930 : 221

13 Thorax mostly black ........... 14

Thorax broadly reddish anteriorly and along sides above wings ; a reddish line on

each pleuron ........ austeni Bromley, 1930 : 221

14 Hind femora mostly black .......... 15

Hind femora mostly yellowish, but apically black, especially dorsally

praedo Austen, 1909 : 57

15 Large, robust species (29-43 mm) ; beard reddish . . athletes Speiser, 1907 : 356
Smaller, slenderer species (23-31 mm) ; beard straw-coloured

neavei Bromley, 1930 : 223 (p. 255)

1 6 Dorsum of thorax orange, unpatterned, contrasting sharply with black-brown

pleura and slightly shining, blue-black abdomen

redimiculurn Speiser, 1914 : 7 (p. 255)

- Dorsum of thorax orange with distinct black stripes and spots ; not contrasting

sharply with pleura, which are patchily black and orange, obscured by white
tomentum ............. 17

17 Abdomen orange with orange hairs and bristles. Black longitudinal stripes of

mesonotum reaching scutellum and extending laterally into connecting black spots

igniferum Engel & Cuthbertson, 1937 : I2

- First five abdominal segments black, rest bright orange. Black longitudinal stripes

of mesonotum not reaching scutellum, and not connected with any lateral spots

lampyroides sp. n. (p. 254)

254 H- OLDROYD

Proagonistes apicalis (Curran)

Lamyra apicalis Curran, 1927 : 6 ; 1928 : 331.
Proagonistes apicalis (Curran) Bromley, 1930 : 216.

LULUA : Kapanga, i $, 11.1934 (G. F. Overlaet). COQUILHATVILLE : Eala, i $,
vi.i932 (A. Corbisier). STANLEYVILLE : Yangambi, i °-> 3O.vii.i959 (/. Decelle).

Proagonistes lampyroides sp. n.

(Text-fig. 33)

Shares with P. apicalis (Curran) and P. redimiculum Speiser the orange, black-
banded mesonotum, but is distinguished from apicalis by the wholly dark wings,
and from redimiculum by the conspicuous red tip to the black abdomen. This
involves the sixth and following segments, and the ovipositor is distinctly shorter
than that of $ redimiculum.

$ Head. Red. Ocellarium has a small, black patch running longitudinally, not transverse
as in redimiculum. Frons otherwise rather bare and shining brown ; rest of head, including
buccae and occiput, orange with orange tomentum, bristles and hairs ; only a small tuft of
whitish hairs in beard. Palpi red with orange hairs. Proboscis red at base, shining black for
most of its length, but with orange hairs. Antennae orange with orange hairs, third segment
about twice as long as first two segments together.

Thorax. Orange with black pattern. On mesonotum a pair of longitudinal black stripes
from anterior margin almost to reach scutellum ; on each side a roundish black spot before
transverse suture and an elongate triangular black spot posteriorly. Scutellum entirely orange.
Sides of pronotum, propleuron, mesopleuron and sternopleuron with round black spots ;
otherwise pleura orange with tomentum that is mostly white, especially ventrally and pos-
teriorly. Postnotum (' metanotum ') orange with a pair of black spots.

Abdomen. First five segments dorsally and ventrally black with black-brown tomentum
and predominantly black short hairs ; some orange hairs laterally on tergites, where there are
one or two orange bristles on all five segments. Sixth segment and posteriorly bright orange
with orange hairs.

Legs. Coxae like pleura but with white hairs as well as white tomentum ; legs otherwise
entirely orange, with orange or yellow hairs ; only claws apically black.

Wings. Entirely black-brown, with orange hairs on base of costa. Halteres orange.

Length of body 23 mm ; of wing 20 mm.

Holotype <J>. GABON : Lambarene", 1921 (E. le Moult] (MHNP).
Paratypes. CONGO : Riv. de San Benito, i $ (Quivae}. EQUATEUR : Bokuma,
i ?, vii.i932 (R. P. Lootens) (MRAC).

Proagonistes leoninus Bromley

Proagonistes leoninus Bromley, 1930 : 216.

ILE DE SAO THOME : i $ (De Saeger ; S.A.R. Prince Leopold] (MRAC).
Sao Thome" is the type-locality of this species.

ASILIDAE OF THE CONGO BASIN 255

Proagonistes neavei Bromley

Proagonistes neavei Bromley, 1930 : 223.

UELE : Bambesa, i $, iii.1937 (/. Vrydagh) (MRAC).

Originally described from UGANDA, this is a robber-fly that may well occur in
the Pare National du Garamba.

Proagonistes pliomelas Speiser

Proagonistes pliomelas Speiser, 1907 : 358 ; Bromley, 1930 : 217.

LULUA : Kapanga, 3 <$, i $, xi.iQ32 (G. F. Overlaet). UELE : Bambesa, i <#,
30.x. 1933 (/. V. Leroy). LUALABA : Sandoa, i °., vi.i932 (G. F. Overlaet) (MRAC).

Described from the northern Cameroons, and known also from Ghana and
Mayumbe.

Proagonistes redimiculum Speiser

Proagonistes redicimiculum Speiser, 1914 : 7 ; Bromley, 1930 : 217.

COQUILHATVILLE : Eala, I9.x.i93i (H. J. Bredo) (MRAC).

Originally described from Tiko, near Victoria, in the Cameroons. The present
specimen agrees closely with Speiser's description, except that the scutellum is
entirely fire-red, instead of being black with a red tip. The rest of the colour-
pattern is so striking, with the bright red head, thorax and legs, coupled with black
pleura, abdomen and wings, that I think we must accept the two as conspecific.

Proagonistes rufibarbis (Fabricius)

Laphria rufibarbis Fabricius, 1805 : 157.
Lamyra rufibarbis (Fabricius) Curran, 1928 : 331.
Proagonistes rufibarbis (Fabricius) Bromley, 1930 : 220.

LULUA : Kapanga, 5 <£, 3 $, 1932-3 (G. F. Overlaet}. STANLEYVILLE : Basoko,
i <J, ^.1948 (P. L. G. Benoit).

Curran (loc. cit.) recorded this species from Malela.

ANDRENOSOMA Rondani

Andrenosoma Rondani, 1856 : 160 ; Hull, 1962 : 349. Type-species : Asilus ater Linnaeus,
1758, monotypic.

The affinities of this genus are discussed briefly under Proagonistes. Andrenosoma
occurs in all the major zoogeographical regions, but is an enigmatic genus every-
where ; the huge Andrenosoma of Australia and South America have little obvious
affinity with the three distinctive Palaearctic species. The species of tropical
Africa have been particularly obscure, since they have never numbered more than
one or two, and these have been questioned, and in some cases transferred to other

256 H. OLDROYD

genera ; e.g. africana Ricardo and igniferum Engel & Cuthbertson to Proagonistes
in the present work.

At present only two remain, separable as follows :

KEY TO SPECIES OF ANDRENOSOMA IN TROPICAL AFRICA

i Mesonotum tomented, with a pattern of dark brown stripes on an ashy grey ground.

Post-vertex entirely covered with grey tomentum. Proboscis and palpi as in

Text-fig. 31 boranica (Corti)

- Mesonotum with a broad, bare, median stripe, flanked by double spots, which are

also bare and shining. Post-vertex with a large, triangular, bare, shining area.

Proboscis and palpi as in Text-fig. 30 ; male genitalia as in Text-fig. 32

complexa Sp. n.

Andrenosoma boranica (Corti) was described from Ethiopia, and is also known
from the savannas of West Africa. (Text-fig. 31).

Andrenosoma complexa sp. n.

(Text-figs 30, 32)

Distinguished from boranica by the characters given in the key. The unique
specimen is somewhat teneral, but even so its mahogany-brown colour is probably
natural.

o* Head. A pronounced, rounded facial tubercle occupies half of height of face ; face and
frons with white tomentum and long white hairs at sides, but bare in middle line, both below
antennae and in a median frontal area which includes ocellar tubercle and a large triangular
area of post-vertex. Moustache consists of about a dozen long, black bristles, flanked by long
white hairs from sides of face. Ocellar tubercle with a pair of long, backwardly curved black
bristles, and numerous black bristles on upper occiput ; lower occiput and buccae with soft,
silvery hairs. Palpi and proboscis as in Text-fig. 31 : proboscis slender, more pointed than in
boranica, palpi smaller, but still flattened, scale-like as typical of genus ; hairs at tips of palpi
and bulbous base of proboscis longer than in boranica. Antennae black, with reddish joints,
black bristles and white hair ; clavate third segment about as long as sum of first two segments.

Thorax in ground colour mahogany-brown, with white tomentum, which leaves bare areas
dorsally ; on mesonotum a broad median stripes, complete from front to rear, flanked by broad
sublateral stripes extending from rear to just in front of transverse suture. Scutellum with
white tomentum basally and a bare, shining tip. Hairs and bristles fine, long, black ; scutellum
with six marginal bristles. Pleura entirely covered with white tomentum : all sclerites with
long, fine white hairs, mixed with a few black ones dorsally ; mesopleuron with a single strong
black bristle.

Abdomen also mahogany-brown in ground colour, with white tomentum thin dorsally, but
never quite absent ; dorsally with fine black hairs, laterally and ventrally with longer white
hairs. Male genitalia as in Text-fig. 32, long and complex, with long black hairs and bristles.

Legs. Coxae covered with white tomentum ; rest of legs mahogany-brown with abundant
long, silky hairs, which are black dorsally on tibiae and tarsi, and tips of femora, white elsewhere.

Wings. Uniformly faintly greyish, without darker colour ; veins dark brown, marginal and
fourth posterior cells closed, with long stalk, anal cell with short stalk ; first posterior cell fully
open, veins R$ and M\ running parallel to margin.

Length of body 19 mm ; of wing 14 mm.

Holotype^. BAS-CONGO : Moerbeke (P. Damage) (MRAC).

ASILIDAE OF THE CONGO BASIN 257

Tribe ATOMOSIINI

The tribe Atomosiini is a compact group of Asilidae, which look rather like small
solitary bees. They are united by a peculiarity of wing-venation, in which the
veins forming the posterior, apical angle of the discal cell form a cross +, or a dis-
placed crossing -y. Though apparently trivial, and occasionally to be found in
other tribes, this detail is a very constant point of recognition. Hull (1962 : 369)
points out that almost diagnostic of this tribe is the sclerotization of the area
immediately behind the hind coxae, but this detail is not so easily observed as the
wing- venation.

Atomosiini are divided into a very large number of genera, most of which are
confined to South America. Hermann (1912) established most of them. These
genera are justified by the number of structural variations that occur, especially in
the head and antennae, but it is doubtful how far these differences are of generic
value. The tribe is generally placed in the subfamily Laphriinae, but a much
fuller study is needed to establish its true relationships, (cf. Karl, 1959).

In the Ethiopian Region only a very few species of Atomosiini have been des-
cribed. Hermann created the genus Goneccalypsis for Atomosia argenteoviridis
Hermann from the Transvaal. Engel (1929) described Loewinella nigripes from
Southern Rhodesia as a variety of the Palaearctic L. virescens (Loew), but it seems
clear that this should be treated as a distinct species. Loewinella aphaea Se"guy
(1950) comes from the oasis of Air (Agades) in the Sahara, and Curran (1927)
described an Atractia arcuata from Stanleyville.

The last species is well represented in the collection of the Muse"e du Congo, and
is not "an Atractia (as, indeed, Curran suspected) ; it is clearly congeneric with
Loewinella nigripes Engel.

Loewinella arcuata (Curran) comb. n.

Atractia arcuata Curran, 1927 : 5.

COQUILHATVILLE : i ex., io.xi.i93i (Lt. Domari) ; Eala, 8 ex., xi.j_93i-v.i932
(H. J. Bredo) ; i ex., 22.vii.i9i4 (R. Mayne) ; i ex., xi.ig34 (/. Ghesquiere).
STANLEYVILLE : 22 ex., ii-iii.i928 (A. Collart) (MRAC).

Tribe SAROPOGONINI

Modern attempts to improve upon Loew's division of the Asilidae into four sub-
families find most difficulty in subdividing Loew's Dasypogoninae. Workers in
each region tend to give prominence to genera that are peculiar to their own region :
thus G. H. Hardy (1934), a pioneer of tribal classification in Asilidae, erected tribes
Phellini, Chrysopogonini and Brachyrrhopalini that are all restricted to Australasia ;
while Carrera (1952) recognized a tribe Megapodini for certain S. American genera.

The world classification of Hull (1962) goes further than any other, breaking the
old Dasypogonihae into twelve tribes. At the present stage of knowledge of the

258 H. OLDROYD

African Asilidae it is not profitable to comment critically upon this arrangement,
beyond saying that it appears to raise problems in placing certain genera. I shall
therefore continue to use the classification set out in my earlier paper (Oldroyd,
1963), and divide the old Dasypogoninae into three tribes : Saropogonini, Sticho-
pogonini and Xenomyzini. Of these, Saropogonini are recognized by having the
prosternum completely separated from the pronotum, and appearing as a small
sclerite surrounded by membrane. This is usually visible from the side, or below,
though it may be necessary first to brush off some of the obscuring hairs.

Out of the 26 genera included in my key to African Saropogonini (Oldroyd,
1963 : 6) the present collection contains only eight, together with one new genus,
Dogonia. There is therefore little point in reprinting the key, but a few notable
absences may be recorded. Sisyrnodytes and Acnephalum, Saropogon and Steno-
pogon are generally associated with somewhat arid terrain ; Habropogon, Holopogon
and Heteropogon are essentially subtropical or Mediterranean genera : Hypenetes,
Lycostomus, Teratopus, Hermanella and Spanurus are found in more southerly parts
of Africa, although any one of these may yet be found within the Congo Basin.

NEOLAPARUS Loew

Laparus Loew, 1851 14; [nee Laparus Billberg, 1820, Lepidoptera] . Type-species : Laparus

tabidus Loew, by original designation.
Neolaparus Williston, 1889 : 255 [new name].
Cenopogon Wulp, 1898 : 120. Type-species : Cenopogon bifidus Wulp, monotypic.

Specimens of Neolaparus are abundant in collections of African Asilidae, and are
extremely difficult to classify. Structurally they are remarkably uniform. The
face bulges a little below the antennae, and projects as a rim on the epistoma, but
has no clearly defined knob, and the moustache is very sparse, generally consisting
of two or four stiff bristles with at most a few fine hairs. Only rarely is there a
more extensive moustache. The third antennal segment is usually clubbed, and is
elongate only in one or two species. The male terminalia are fairly elaborate, and
show some inter-specific variation, but only rarely is this decisive enough to be
diagnostic. The wing-venation is primitive, and remarkably uniform.

The species therefore are mostly defined by colour and pattern, chiefly of the
thorax and abdomen. These are reasonably reliable in well preserved specimens,
but unfortunately are quickly ruined by bad preservation. A great majority of all
specimens available for study are badly greased, and one can only guess at their
natural appearance.

As long ago as 1860 Loew made very similar comments about the difficulty of
classifying species of this genus, which he divided into two sections, one with only
two strong bristles on the epistoma, and the other with four or even six. Bromley
(1956 : 140), while using this arrangement in his key, wrote : ' This is not a good
grouping, as the number of bristles may vary within a species '. There is, never-
theless, evidence of a division of the genus into two groups of species, the one con-
sisting of larger and darker flies, the other more fragile and paler in colour. The
first group have the alula of the wing larger, and with a distinct posterior angle,

ASILIDAE OF THE CONGO BASIN 259

whereas the flies of the second group have the alula smaller, and with its posterior
margin only slightly convex.

Hull (1962 : 256) says that : ' Both sexes of Lagodias may be separated from
the related Neolaparus and also from Pegesimallus by the absence of the ventral
hypopleural patch of dense pile '. This is true of the type-species of each genus,
but is not a valid generalization. This patch of pile is widespread among many
genera of this and other tribes, e.g. Laphria, but it appears sporadically, and may
be present or absent in species that are clearly congeneric.

The species of Neolaparus are in such confusion that it seemed better in the present
paper to mention only those few specimens that come from the Pare National du
Garamba, deferring a more general account of the Neolaparus for a more com-
prehensive revision. Curran's key (1934) is a useful guide if it is used with caution,
but no really satisfactory key to the species of Neolaparus has yet been produced.

Neolaparus ophion Speiser

Neolaparus ophion Speiser, 1910 : 86.

GARAMBA NATIONAL PARK : P.N.G., 527, I/o/i, i $, 17^.1950 ; 766, I/o/i, i <$,
2i.viii.i95o (G. Demoulin) ; 786, I/o/i, i °-, 25.viii.i95o (G. Demoulin) ; 3480,
Inimvua, i ex., i6.v.i952 ; 764, 1/0/7, I ?» I9.viii.i95o (G. Demoulin) ; 349, I/o/i,
i $, 28.iii.i95o ; riv. Abita, i $, i.iv.i954 (C. Nebay) (IPNC).

Neolaparus angusticornis Ricardo

Neolaparus angusticornis Ricardo, 1925 : 245.

GARAMBA NATIONAL PARK : P.N.G., 352, 1/0/3, * & 31.111.1950 ; 1588, II/hc/4,
i $, 2o.iv.i95i (/. Verschuren] ; 1849, H/hc/4, i <£, 31^.1951 (/. Verschureri) ;
3328, Pidigala, i $, 23^.1952; 1494, II/fd/17, i <£, 4.^.1951.

Neolaparus munroi Bromley

Neolaparus munroi Bromley, 1936 : 142.

GARAMBA NATIONAL PARK : P.N.G., 1464, II/fd/2, i $, 28.iii.i95i ; 3328,
Pidigala, 2 £, i °., 23.^.1952 ; 3476, Aka/2, 19^.1952 ; 3488, Inimvua, 2 <$, 20.v.
1952 ; riv. Abita, i $, i ?, i.kv.i952 (C. Nebay) (IPNC).

Neolaparus decoratus sp. n.

A well-marked species, of distinctive pattern. Resembles ophion Speiser in
having microtrichiae entirely covering the wing, and not confined to an apical area,
but differs in having a clear-cut abdominal pattern, and in the bicoloured, strongly
clubbed antennae.

$ Head. Pronto-facial area broad, not greatly narrowed at antennae, black in ground colour,
but obscured by thick tomentum. On frons this tomentum is brassy brown, more yellow along
eye-margins and just above antennae, where there is a short, vertical black stripe. Face quite
prominent at epistoma, but triangular in profile, with flat upper surface, tomentum yellow-

260 H. OLDROYD

brown, white laterally ; only two strong, pale bristles on epistoma, and a few short, pale yellow
hairs. Probosics and palpi black, with black hairs. Antennae with first two segments yellow-
brown, short, subequal, with yellow bristles ventrally, black ones dorsally ; third segment
strongly clavate, pale yellow at base, black on expanded portion. A pair of yellow postvertical
bristles. Occiput with yellow tomentum and a few yellow hairs.

Thorax olive-brown, with indistinct, divided, median stripe, outlined by very narrow dark
brown lines, as in ophion. Very short black hairs, each arising from a dark spot, run along lines
of dorsocentrals, and cover a large posthumeral area, as well as the rather paler lateral margins
of the mesonotum. Bristles yellow, very strong, restricted to one notopleural, one supra-alar,
and one postalar. Scutellar disc concolorous with mesonotum, but rim dark red-brown ;
short hairs in line with dorsocentrals, but no bristles.

Pleura with brassy yellow tomentum, through which the ground colour is visible : pronotum
and propleuron, mesopleuron, sternopleuron, hind coxae and a little of the other coxae show a
dark ground colour ; pteropleuron, mesopleuron and metanotal callosities have a yellow ground.

Abdomen black, with a dull sheen, and therefore contrasting strongly with thorax. Dorsally
first segment and base of second with bronze tomentum, leaving bare the bulla on the second
segment (a feature of Neolaparus and Lagodias). Segments 2-6 each has a pair of spots of
white tomentum : on second segment these are short, transverse lines, and occur at half-way
position ; on each succeeding segment they become further forward until on sixth they are small
spots on fore margin. These segments also have on extreme lateral margins a spot on anterior
border and a short band in posterior angle. Hairs very short, very scattered, black. Venter
shining black with grey hind margins. Segments 7, 8 shining black, forming an ovipositor.

Legs. Long, slender, all femora and tibiae slim on basal half, distinctly thickened towards
tip. Generally chestnut-brown in colour, but femora pale on narrow basal stalk and black-
brown over thicker apex. Clothing hairs black, long bristles yellow.

Wings. Venation normal, all cells open. Microtrichiae cover entire wing membrane, as in
ophion, and do not form a grey tip. Alula very shallow, gently curved.

Length of body 13 mm ; of wing n mm.

o* not yet known.

Holotype <j>. GARAMBA NATIONAL PARK : P.N.G., 3287, II/gc/6, 5.^.1952
(IPNC).

LAGODIAS Loew

Lagodias Loew, 1857 : 345 ; 1860 : 69. Type-species : Lagodias albidipennis Loew, 1857 ;
by monotypy.

The males of Lagodias are conspicuous in having some part of the legs, and
especially of the hind legs, with a spectacular fringe of flattened, scale-like hairs.
Apart from this feature they are typical Neolaparus, and the females are very
difficult to recognize as Lagodias. Hull (1962 : 256) claims that both sexes of
Lagodias can be distinguished from Neolaparus by not having the small, dense tuft
of short hairs on the hypopleuron, but, as we have already seen, this is not found in
all species of true Neolaparus.

The practical effect of this feathering of the legs is an interesting question.
Hermann (1906 : 144) stresses that this is a secondary sexual character, and that
the extent of the feathering varies in different species. It may be found on the
tarsus only ; on the tarsi and apically on the tibiae ; on tarsi and tibiae, but not on
femora ; or on all segments of the hind and middle legs. The shape of the individual
scales varies, too, from slightly flattened hairs to large scales, flattened like a paddle-
blade or a banana leaf.

ASILIDAE OF THE CONGO BASIN 261

Hermann also speculates about the possible function of the feathered legs :
' Then one can hardly avoid the conclusion that this monstrous feather-apparatus
must seriously handicap the insect, which is far from robust, and it can therefore
be imagined that the insect possibly hunts its prey over the smooth surface of
water, and that when the feathers are erected they may form a swimming paddle.'.
He wisely adds that this remains mere speculation until observations on the habits
of these flies can be recorded — observations that are still wanting, nearly sixty
years later. On the whole Hermann's guess seems rather unlikely, the more so
since both sexes seem to have the same hunting habits. Is it not more likely that
the ornamentation of the legs of the males contributes to sexual display, and that
the so-called genus Lagodias is no more than a collective name for those species of
Neolaparus in which the males have developed such sexual ornamentation : com-
pare, for example, the genus Dolichopus?

The collection from the P.N. Garamba contains only a single specimen, which
cannot be referred to any known species.

Lagodias griseus sp. n.

Distinguished from most of the known species of Lagodias by the fact that the
scales of the middle and hind legs completely cover the tarsi and the tibiae, but there
are none on the femora. There is at least one other unidentified species with this
type of ornamentation, but the present species is unique in having the thorax almost
entirely dusted with white, and only indistinctly darker paired longitudinal stripes
are visible. Wings nearly white, with pale veins.

c£ Head. Shaped as in those species of Neolaparus that have a broad front, and have facial
hairs in addition to the two strong bristles. Fronto-facial area almost as broad as one eye,
scarcely narrowed at antennae. Evenly covered with white tomentum, and with white hairs.
Two strong, white bristles on epistoma, and many white hairs forming a moustache, but not
extending more than half-way to antennae. Proboscis and palpi black with mainly black hairs.
Antennae blackish, with pale constrictions between the segments, and at base of third segment :
hairs long, white ventrally, shorter and black dorsally. Occiput uniformly covered with white
tomentum, and with numerous silvery white hairs, among which the pair of white postverticals
are almost lost.

Thorax black, with a thick covering of white tomentum, with no pattern except for a paired
longitudinal stripe which appears blackish because the tomentum is thinner there. Abundant
very fine white hairs curled upwards (compare Pegesimallus) . Strong bristles are i notopleural,
i supra-alar, and i postalar. Scutellum with erect, soft, white hairs all over disc, but no strong
bristles.

Abdomen. Dorsum shining black, with a purplish sheen. White tomentum covers first
tergite, and surrounds bulla on second, as well as forming a pair of prominent white spots of
segments 2-6. Venter shining black with yellow segmentations. Genitalia black, epandrium
reddish.

Legs. Elongate and slender, femora and tibiae swelling to a knob at apex. Femora and
tibiae yellow basally, otherwise legs red-brown or dark brown. Fore legs without ornamenta-
tion. Middle and hind legs each with no scales on femora, and with a broad, conspicuous,
double fringe of dense black-brown scales, which continue without diminution to apex of fifth
tarsal segment. Normal vestiture of short black clothing hairs and white stiff bristles present
on all legs.

262 H. OLDROYD

Wings. Hyaline, very pale yellowish and iridescent clear membrane, no microtrichiae
except at extreme tip of wing.

Length of body 15 mm ; of wing 13 mm.

Holotype $. GARAMBA NATIONAL PARK : P.N.G., 1514, II/gg/8, io.iv.i95i
(IPNC).

Paratype $. ITURI : Mongabi, pres Faradje, 4.^.1930 (Collart) (MRAC).

PEGESIMALLUS Loew

Pegesimallus Loew, 1857 : 344 '• 1860 : 69. Type-species : Pegesimallus ursinus Loew,
monotypic.

As stated above, Pegesimallus is closely related to Neolaparus, and only to be
distinguished by being covered with abundant, rather long, soft hairs. Hull's
claim that Pegesimallus, like Neolaparus, has the small, compact hair-tuft on the
hypopleuron, whereas Lagodias has not, is apparently not valid. The present
specimen has no such tuft, although in proportions of body and head, as well as in
general hairiness, its affinities clearly lie with Pegesimallus ursinus rather than with
the stark, bare flies that are typical of Neolaparus.

Three species have been described in Pegesimallus, all from southern Africa.
The present species differs from all of these in the much greater length and density
of the hairs clothing the hind leg (Text-fig. 37). Neolaparus morio Bezzi from West
Africa, and one or two undescribed allied species, ought perhaps to be transferred to
Pegesimallus, if this genus is to be maintained at all.

Pegesimallus saegeri sp. n.

(Text-fig. 37)

A dull black, or black-brown species, with brown antennae and legs. Middle and
hind legs exceptionally hairy, but not scaly as in Lagodias, hairs longer and denser
than in any other species.

o* Head. Fronto-facial area parallel-sided, about two-thirds as broad as one eye. Antennae
at upper third. Frons black, with thin brown tomentum. A row of black hairs on each side
runs from vertex and crosses diagonally to base of each antenna. Face produced at epistoma
about as far as length of palpi, but with a flat profile, no facial hump (compare Lycostomus).
Covered with dense brown-grey tomentum. Moustache black, composed of two strong bristles
and a mass of black hairs, which does not quite extend up to antennae. Palpi cylindrical, or
very slightly thickened at tip, first segment tiny and inconspicuous ; black, with very long
black hairs. Proboscis black. Antennae chestnut-brown ; first two segments subequal ;
third constricted basally, slightly clavate, microsegment at tip almost undetectable, with a
small apical spine ; hairs and bristles black, third segment with a number of short black bristles
dorsally and externally. Occiput with brown tomentum and a mass of fine black hairs, a pair
of strong black postvertical bristles, and one or two others concealed among the fine hairs.

Thorax. Mesonotum strongly convex, but in a uniform curve, not projecting anteriorly ;
covered with thick, black-brown tomentum, with only faint traces of a pair of median longi-
tudinal stripes, and more reddish on lateral margins. The mane of fine black hairs is less dense
than in Pegesimallus ursinus Walker, and is confined to the dorsocentral stripes, a large area
behind each humeral lobe, and broad lateral margins. Strong black bristles comprise one
notopleural, one supra-alar, and one postalar, with some bristly hairs nearby. Scutellum with

ASILIDAE OF THE CONGO BASIN

263

no bristles, and with a tuft of hairs on each side in line with supra-alars (as in many species of
Neolaparus). Pleura black-brown, with rather long black hairs on most sclerites, but no trace
of a dense little patch on hypopleuron.

Abdomen black, only faintly reddish at sides of first three tergites. Covered with black
tomentum, seventh tergite with a pair of large areas of shining black. Hairs both dorsally and
ventrally long, dense, and erect. Abdomen a little narrowed basally into a waist, but after
fifth segment it is laterally compressed. Male genitalia of similar construction to those of
Neolaparus, mostly shining black, but epandrium and aedeagus red.

Legs. Long and moderately slender, chestnut-brown. Hind femora a little stouter than in
typical Neolaparus, but made to appear more so by the long, dense, black hairs, as shown in
Text-fig. 37. Bristles few, and scarcely to be distinguished from the surrounding hairs. Middle
legs similar, fore legs rather less hairy. Coxae with tomentum and hairs like those of pleura.

Wings. Venation normal, only anal cell closed on wing margin. Alula small, rounded.
Membrane stained almost uniformly yellow-brown.

Length of body 15 mm ; of wing 12 mm.

Holotype^. GARAMBA NATIONAL PARK : P.N.G., 349, I/o/I, 28.111.1950 (IPNC).

MICROSTYLUM Macquart

Type-species : Dasypogon venosum Wiedemann, 1821, by

Type-species : Mimoscolia fafner Enderlein, 1914, by

26.

168.

Microstylum Macquart, 1838

designation of Back, 1909.
Mimoscolia Enderlein, 1914

original designation.

This is another very large and difficult genus, in this respect resembling Neolaparus,
and equally in need of a thorough revision, taking into consideration all the species
throughout the Ethiopian Region. The study of Microstylum is further complicated

FIG. 37. Pegesimallus saegeri. Left hind leg.

264 H. OLDROYD

by the existence of a large number of species in Madagascar, where they form a
striking part of the Asilid fauna. Some species are amongst the biggest flies in the
world.

The present collection from the Pare National du Garamba contains only three
specimens of Microstylum, belonging to two species, both of which have a spur-like
prolongation of the tip of the middle tibia (Text-fig. 39). This brings them into
Mimoscolia Enderlein, which Hull (1962 : 160) regards as a full genus, though other
authors have considered it to be at most a subgenus of Microstylum. Its status may
be even less than that : it is possible that those species with some form of prolon-
gation of the middle tibia are not even a species-group, but merely unrelated species
that happen to have this detail in common. This view receives support from the
fact that M. cilipes Macquart and M. mydas Engel, both from Madagascar, have a
process of a sort, not on the tibia, but on the basitarsus of the middle leg, perhaps
serving a similar practical purpose. Hull (1962) puts mydas in Microstylum and
cilipes in Mimoscolia, though Engel (1932) puts them into the same couplet of his
key.

Bezzi (1908) described two genera from the Congo Basin that are very closely
allied to Microstylum. Eclipsis, monotypic for maculiventris Bezzi, is distinguished
from Microstylum by having the first posterior cell of the wing closed, and the costal
vein not extending much beyond R± ; while EpibUpharis , monotypic for pedunculata
Bezzi, has the second posterior cell, of characteristic shape in Microstylum, pedun-
culate rather than sessile on the tip of the discal cell. Hull (1962 : 158, 159) rightly
suspects that these may be no more than anomalous species of Microstylum. One
badly preserved specimen from the Mayumbe, in the collection du Muse"e du Congo,
could be Epiblepharis pedunculata, but if this is correctly identified it supports Hull's
view, because the second posterior cell is sessile in one wing, and barely pedunculate
in the other.

The following key is extremely incomplete, especially in regard to species des-
cribed by the older authors from the Cape. It is given because it covers most of the
species described from the Congo Basin or areas immediately adjacent, and because
the existing keys deal mainly or partially with the many Microstylum from
Madagascar.

KEY TO CENTRAL AFRICAN MICROSTYLUM

1 Middle tibia with spur, the tibia itself being prolonged into an acute process (Text-

fig- 39) (Mimoscolia Enderlein) ......... 2

Middle tibia sometimes with a comb of spines, or a slightly extended rim, but never

with a spur (Microstylum s. str.) ......... 4

2 Abdomen black, with white hind margins ; last two or three segments red (?) or

with grey bands becoming more complete from third segment onwards (<$)

helenae Bezzi, 1914 : 3 (p. 266)
Abdomen without distinctive cross-bands, and without red tip .... 3

3 Mesotibial spur claw-like, with several strong spines. Femora black with red tips ;

tibiae and tarsi red. Abdomen mainly red, with black base and tip, and white
clothing hairs ....... braunsi Bromley, 1932 : 263

- Mesotibial spur conical, with small, globular tip. Legs black or only obscurely

reddish. Abdomen black, with ashy grey or brown tomentum pollex sp. n. (p. 266)

ASILIDAE OF THE CONGO BASIN 265

4 Abdominal tergites with distinct transverse bands ...... 5

Abdominal tergites not distinctly banded ........ 6

5 Abdominal tergites black, with grey hind margins

lituratum Loew, 1863 : 10 (elegans Ricardo), 1900 : 168
Abdominal tergites grey with narrow black segmentations . hirtipes Ricardo, 1925 : 259

6 Pteropleural bristle present .......... 7

Pteropleural bristle absent .......... 16

7 Proboscis abnormally long and slender, much longer than height of eye . . 8

- Proboscis sometimes a little longer than height of head, but not conspicuously

slender ............. 12

8 Occiput with yellow hairs and bristles, only isolated black ones .... 9
— Occiput with bristles predominantly black. . . . . . . . 10

9 Femora black at base and tip, with red ring in middle . . unnamed sp. (BMNH)
Fore and middle femora black at base, hind femora red . rufum Ricardo, 1925 : 254

10 Scutellum orange ....... nigrescens Ricardo, 1900 : 169

- Scutellum black ............ n

11 Legs distinctly red in part ..... acutirostre Loew, 1852 : 658
Legs black, or only indistinctly brownish . . nigrimystaceum Ricardo, 1925 : 258

12 Occipital bristles partly or wholly black . . . . . . . . 13

Occipital bristles all pale ........... 15

13 Wings milky white (? Q* only). Halteres reddish lacteipenne Wiedemann, 1821 : 371
Wings dark brown, not milky. Halteres black . . . . . . . 14

14 Hind femora as well as hind tibiae covered with white clothing hairs

ricardoae sp. n. (p. 268)

- Hind femora and most of hind tibiae with black clothing hairs ; tibiae with white

hairs ventrally on basal half ..... nigribarbatutn Bigot, 1878 : 408

15 Short hairs covering femora and tibiae uniformly white . validurn Loew, 1857 : 347
Short hairs covering femora and tibiae uniformly black, except for basal half of

anterior face of hind tibiae, where hairs are longer and white

capensis Fabricius, 1805 : 154

16 Face not swollen except on mouth-margin, to which moustache is confined. Coxae

with short, stiff bristles or spines, sometimes mixed with fine hairs . . . 17

- Face swollen into a conspicuous hump, with a dense moustache. Coxae clothed

partly or entirely with long, fine hairs, sometimes mixed with a few strong bristles 26

17 Base of wing, posteriorly, milky white, constrating with blackish tip

pica Macquart, 1846 : 59
Base of wing not conspicuously different from rest . . . . . . 18

18 Coxae hairy as well as bristly .......... 19

Coxae bristly, with few or no fine hairs . . . . . . . .22

19 Scutellar disc with only tiny spines. Legs mainly black ..... 20
Scutellar disc with fine hairs. Legs orange . . . . . . . 21

20 Proboscis distinctly longer than height of one eye. Male genitalia as in Text-fig. 38

sessile Bezzi, 1908 : 376 (p. 269)
Proboscis shorter than height of one eye. Male genitalia as in Text-fig. 40

attenuatum Ricardo, 1925 : 257

21 Abdomen clothed with short yellow hairs, as well as rust-brown tomentum

venosum Wiedemann, 1821 : 215

Abdomen clothed with rust-brown tomentum, at least on basal segments, but

without short hairs ...... capucinum Bigot, 1878 : 408

22 Abdomen predominantly orange . . . . . . . . .23

- Abdomen dark ............ 24

23 Thorax and legs chocolate-brown, contrasting with abdomen, which is orange

except at extreme base and tip ..... dispar Loew, 1858 : 348

266 H. OLDROYD

- Thorax ashy grey, heavily tomented. Legs mostly red, femora darker above.

Abdomen orange, a little darker at tip, but orange at base

unicolor Ricardo, 1925 : 256

24 Dorsum of abdomen shining black, with narrow yellow segmentations ; narrow strip

of tomentum only along each lateral margin . . spinipes Ricardo, 1925 : 255

- Dorsum of abdomen dull, tomented, at least on first few segments ... 25

25 Hind margins of anterior abdominal segments very clearly margined in yellow

spurinus Walker, 1849 : 323

- Hind margins of abdominal segments not, or very indistinctively marked

partitum Walker ; par cum Karsch, 1887 : 373

26 Legs partly reddish or orange ; femora usually with dark stripe . . . . 27
Legs entirely dark brown or black . . . . . . . . .31

27 Abdomen orange, only extreme base and genitalia black. Hind femora orange with

brown tip . . . . . . . fenestratutn Wiedemann, 1828 : 377

- Abdomen darker. All femora with dark stripe ....... 28

28 Abdomen clothed with rather long, pale hairs ....... 29

Abdomen with sparse and inconspicuous hairs ....... 30

29 Wings heavily stained dark brown. Mesonotum with shaggy hairs in addition to

bristles ........ hermanni Ricardo, 1925 : 249

— Wings quite clear, hyaline. Mesonotum with strong bristles, and very short hairs

only, no shaggy hairs ....... villosutn Bigot, 1878 : 406

30 Palpi orange ......... bloesus Walker, 1849 : 307

Palpi black, though with some long, pale hairs . . varipennatum Bigot, 1878 : 407

31 Halteres dark, and bristles before them black . . . . . . .32

- Halteres yellow, and bristles before them white . . ignobile Loew Q*. 1857 : 347

ruftneurum Macquart $, 1855 : 48

32 Wings extensively whitish basally, with dark tip. Abdomen black with white

lateral margins. Hairs of occiput, beard and coxae mainly black

ustulatum Engel & Cuthbertson, 1938 : 133

— Wings almost uniformly brown. Hairs of occiput, beard and coxae mainly white

glabrum Ricardo, 1900 : 168

Microstylum helenae Bezzi

Microstylum helenae Bezzi, 1914 : 3.

GARAMBA NATIONAL PARK : P.N.G., 331, I/o/i, i $, 27.111.1950 (IPNC).
KATANGA : Ngaye, i <£, 1932 (R. 0. Claquin] (MRAC).

Microstylum pollex sp. n.

(Text-fig. 39)

A species of moderate length (19 mm), rather small for a Microstylum ; black,
with dark wings, and heavily tomented abdomen. Belongs to the group Mimoscolia
Enderlein because of its spur to the middle tibiae. Nearly all the known members
of this group occur in Madagascar, and apart from the distinctly banded helenae (see
above), the only mainland species recorded is braunsi Engel from the Cape. This
differs from pollex in the shape of the mid-tibial process, the colour of the legs, and
the predominance of black bristles on the head.

<£ Head black, with cinereous grown tomentum and white hairs and bristles. Sides of frons
with multiple row of white bristly hairs. Face in profile almost straight, and only a little

ASILIDAE OF THE CONGO BASIN

44

FIGS 38-44. 38-40, Microstylum spp. 38, M. sessile, $ genitalia ; 39, M. pollex, mid-
tibial spur ; 40, M. attenuatum, 3 genitalia. 41-44, Dogonia spp. 41, D. nigra, $
genitalia ; 42, D. saegeri, head and detail of antenna ; 43, D. saegeri, $ genitalia ;
44, D. saegeri, head from in front.

268 H. OLDROYD

prominent on mouth-margin ; with brown tomentum and no hairs above moustache, which is
dense, entirely white, and confined to epistoma. Palpi black with black hairs ; proboscis black
with white hairs beneath base, merging with snow-white beard. Occiput covered with whitish
tomentum and white hairs and bristles.

Thorax black, with ashy grey tomentum. Mesonotum more brown in middle, and traces of
paired, rather broad, longitudinal stripes, and between them a groove in surface. Tiny clothing
hairs black over most of area, but white round margins. Scutellum grey, bare on disc, with
two strong marginal bristles, and one or more weaker ones laterad. Pleura ashy grey or blackish,
with sparse hairs, mostly black, but white tufts on propleura and all coxae.

Abdomen black. Dorsum with ashy grey tomentum, which is more brown in middle. Short
clothing hairs white. Venter grey with white hairs. Genitalia shining black : hairs and
bristles strong and white on hypandrium, mostly black on epandrium.

Legs black, knees dull reddish, tibiae and tarsi no more than obscurely reddish. Short
clothing hairs nearly all white, some black on femora ; strong bristles of tibiae and tarsi mixed
white and black.

Wings uniformly dark brown. Halteres with light brown stalk and darker knob.

Length of body 19 mm ; wing 14 mm.

Holotype^. GARAMBA NATIONAL PARK : P.N.G., 331, I/o/i, 27.111.1950 (IPNC).
Paratype $. 3267, Ndelele/Kii7, 27.111.1952 (IPNC).

Microstylum ricardoae sp. n.

Long ago, Miss Ricardo set aside two female Microstylum in the BMNH collection
from Kambove, in the Katanga, as ' $ of nigribarbatum Bigot, or sp. nov.' In the
collection of the Musee du Congo Beige is a series of females of the same species, and
one male, clearly conspecific, and all from the vicinity of Elisabeth ville. It thus
becomes apparent that Miss Ricardo's species is not the female of Bigot's, but
should be described as new. It is easily distinguished from nigribarbatum by
having the hind femora and hind tibiae conspicuously clothed with white hairs.

$ Head black, thickly covered with yellowish grey tomentum, more brownish on frons, paler
on face. Frons with bristly black hairs in multiple rows along each eye-margin. Face bare
except for a dense moustache of stiff bristles which may be either bright yellow or whitish, on a
tubercle that occupies lower half of face. Palpi and proboscis black with black hairs. Beard
silky, black ; occipital bristles and hairs nearer eye-margins, black ; inner part of occiput with
silky white hairs.

Thorax dull black, with rusty brown tomentum and black hairs and bristles. An abundant
covering of fine hairs as well as strong bristles at sides of mesonotum and on scutellum (2 strong,
2 sometimes weaker) ; no supra-alars. Pleura also with many fine black bristles, including
pteropleuron which, however, has no strong bristles.

A bdomen black, dorsally and ventrally sharply divided into a dull anterior half and a shining
posterior half. First four segments entirely obscured by black tomentum, but with short
white clothing hairs. Segments 5-8 shining black, bare.

Legs. Black : coxae with dense, long, black hairs, no obvious strong bristles ; rest of legs
with predominantly white clothing hairs, especially on hind femur, tibia and basitarsus ; else-
where with some black hairs and bristles.

Wings uniformly rusty brown. Halteres black.

Length of body 27 mm ; of wing 21 mm.

cJ very similar, except that all the abdominal segments are dull, without a shining black tip.

ASILIDAE OF THE CONGO BASIN 269

Holotype $. KATANGA : 150-200 m. W. of Kambove, 3500-4000 ft, ly.x-
i.xi.oy (Neave Coll : BM 1907-230) (BMNH).

Paratypes. Same data as holotype, i $ (BMNH) ; KATANGA : Elisabethville,
i (£, 2 $ (Miss. Agric.} ; Lumbashi, 3 $, n.x.28 (Ch. Seydel) ; Elisabethville, I $,
12. xi. 1923 (Ch. Seydel) (MRAC).

Microstylum sessile Bezzi

Microstylum sessile Bezzi, 1908 : 376.

BAS-CONGO : Kisantu, i <£, 1927 (R. P. Vanderyst) ; Congo da Lemba, i <£, i $,
1912 (R. Mayne) ; Lemfu, i $, xii.i945 (Rev. P. L. de Beier) ; Boma, i $ (R. F.
Achille) ; Banana, i $, xi.i934 (P. Henrard) (MRAC).

DOGONIA gen. n.

Type-species : Dogonia saegeri sp. n., by present designation.

Related to Scylaticus Loew and to Cyrtopogon africanus Ricardo, but easily dis-
tinguished from both by having the metanotal callosities hairy. Hull's key
(1962 : 119) to his tribe Stenopogonini recognizes only seven genera of this tribe
that have hairy metanotal callosities, and three of these belong to Microstylum
sensu lat. In Hull's key the new genus runs down to Neodysmachus Ricardo, but is
quite different in appearance from that elongate, bristly, Australian genus. In
general appearance Dogonia looks like a Saropogon, but without the fore tibial spurs.

Head (Text-figs 42, 44). In profile eyes shallow, occiput conical, exposing a large surface,
which has long, fine hairs but no stiff bristles. Seen from in front, face is parallel-sided, almost
one quarter of head-width ; eye-margins excavate above antennae (Text-fig. 44). Ocellar
tubercle large, but sunk below level of vertex. Antennae with two rather short, subequal
segments, then an awl-shaped third segment, ending in a short, conical, microsegment, and a
distinct apical style. Face gently convex in upper half, lower half expending smoothly into a
distinct knob ; moustache confined to this knob. Palpi two-segmented, little longer than
epistoma ; both segments subequal, second a little swollen basally, narrowed apically, with
apical pit and style. Proboscis as long as height of eye.

Thorax not structurally remarkable.

Abdomen. Depressed cylindrical, tapering a little apically, like that of Saropogon. Bare,
shining, with narrow band of tomentum on extreme lateral margins only. Male genitalia
compact, with pointed lobes, rotated through about 90° anticlockwise. Female with rather
long, slender acanthophorites.

Legs. No spurs on tibiae, and no other structural peculiarities. Femora and tibiae slightly
swollen in one species, less so in other.

Wings. Of primitive shape : i.e. broad, with costal cell broad, and all cells open on wing-
margin, including anal.

Dogonia saegeri sp. n.

(Text-figs 42-44)

A rather small species, with dark head, thorax and wings, red abdomen and legs.
It differs from the following species, D. nigra sp. n. in coloration and in the male
genitalia. (Text-figs 41, 43).

270 H. OLDROYD

(J Head. Black. Frons and face with dense yellowish tomentum. Frons with multiple
row of bristly hairs along eye-margins, and another row leading from each antenna to ocellar
tubercle : mostly pale yellowish, a few brownish or black ; ocellar tubercle with two or three
pairs, some black. Face similar, but with no hairs dorsal to facial tubercle ; all hairs and bristles
concentrated into a moustache, which is confined to facial tubercle, and almost entirely pale
yellowish or white, with only one or two black ones. Palpi black with black hairs. Proboscis
black or mahogany-red, with yellow hairs. Antennae black, with pale yellowish hairs. Occiput
black with whitish tomentum and silky whitish hairs, no strong bristles.

Thorax. Ground colour black with some reddish areas, notably humeri, postalar calli and
posterior parts of pleura. Covered with brown tomentum, yellow laterally. Pronotum with
long, fine yellowish hairs, and no strong bristles. Mesonotum clothed uniformly and fairly
densely with black hairs, which are slightly longer than first antennal segment. Scutellum
with no true apical bristles ; with fine discal bristles and a submarginal row of 6-8 longer, erect.
Pleura patchily black and red, covered with dense yellowish tomentum except for a large area of
shining black on mesopleuron. Pleural tufts of hairs pale yellowish, except those before
halteres, which may be partly black.

Abdomen bright orange : dorsum bare and shining, except for a narrow strip of whitish
tomentum along each side. Face black, rather long, clothing hairs distributed over entire
dorsum. Male genitalia (Text-fig. 43) red with black tips, and with long black hairs.

Legs. Coxae like pleura, with whitish hairs. Rest of legs mahogany-brown or irregularly
blackish, with black bristles, and clothing hairs that are uniformly whitish. Pulvilli light
brown ; claws black, reddish at base.

Wings dark brown, with purple reflections, and perhaps with clearer centres to some cells
Halteres yellow.

Length of body 12 mm ; of wing 9 mm.

$ closely similar.

Holotype^. GARAMBA NATIONAL PARK. P.N.G., 327, Akam, 24.111.1950 (IPNC).

Paratypes. Same data as holotype, i $ (IPNC) ; 305, Mt. Ndogo, i <£, 3 $,
15.111.1950 ; 3262, II/fc/i8, i £, 3i.iii.52 (IPNC).

Dogonia nigra sp. n.

(Text-fig. 41)

Distinguished from D. saegeri by its black colour, and by the different shape of
the male terminalia (Text-figs 41, 43).

<J Head black, with dense white tomentum, a little brownish on vertex. Bristles a little
stronger, perhaps, than in saegeri, in some rows on frons, along eye-margins, and from antennae
to ocelli ; mostly pale yellowish, but with isolated black ones. Moustache confined to facial
knob, composed of white bristles with a single black one at each side. Palpi and proboscis
black with yellow-brown and black hairs. First antennal segment black, second and third
dark reddish ; first two segments with yellowish hairs. Occiput with white tomentum and
silky white hairs.

Thorax. Pronotum speckled brown-grey, with fine, pale yellowish hairs. Mesonotum with
thick tomentum, black-brown in two longitudinal stripes and paired lateral spots, yellowish in
between ; entirely clothed with rather long black, bristly hairs, individually spaced, and hardly
to be distinguished from the thicker bristles. Scutellum ashy dark grey, with sparse, curved
clothing hairs, and with 4-6 submarginal bristles, curved upwards, and the middle pair even
crossed. Pleura black, with ashy yellowish grey tomentum, leaving bare a large, shining black
patch on mesopleuron. Hairs mostly pale, but with some black bristly hairs on mesopleuron.

Abdomen. Dorsum shining black-brown ; entire first segment, side-margins of rest, and a
narrow hind margin of seventh tergite with grey tomentum. Venter shining black-brown.

ASILIDAE OF THE CONGO BASIN 271

Male genitalia generally similar to those of saegeri, but differing in detail as shown in Text-figs

4i. 43-

Legs. Coxae like pleura, with ashy yellowish tomentum and pale hairs. Rest of legs shining
black-brown, with hairs and bristles partly white, partly black. Pulvilli brown, claws brown
at base, black at tips.

Wings dark brown with distinctly paler centres to most cells. Halteres yellow.

Length of body 10 mm ; of wing 7 mm.

Holotype $. GARAMBA NATIONAL PARK : P.N.G., I/a/i, 13.01.1950 (IPNC).

ANCYLORRHYNCHUS Berthold

Ancylorrhynchus Berthold in Latreille, 1827 : 498. Type-species : Asilus glaucius Rossi, 1790,
monotypic.

Xiphocera Macquart, 1834 : 279. Type-species : Xiphocera percheroni Macquart, 1834, original
designation.

Enchocera Blanchard, 1845 : 463. No included species ; recognizable because published as
synonym of Xiphocera Macquart.

Opegiocera Rondani, 1845 : 153. [Nomen nudum].

Elasmocera Rondani, 1845 : 153. Type-species : Elasmocera cingulata Rondani, 1845, mono-
typic.

This distinctive genus is easily recognized by the most peculiar shape of the
proboscis (Text-figs 44-48), short, pointed, backwardly curved, and when seen in
profile, resembling a comma, or a parrot's beak. The genus occurs in the Palaearctic,
Ethiopian and Oriental Regions, and is remarkably constant in general characters
which, except for the proboscis, are close to those of Scylaticus (q.v.).

It seems evident that the characteristic proboscis must be related to some
peculiarity of diet, but as far as I know no-one has attempted to speculate what it
might feed upon. Some observations on this proboscis will be the subject of a
separate small paper.

At least 22 species of Ancylorrhynchus exist in the Ethiopian Region, but only
A . crux Bezzi is recorded from the Congo Basin, and that from an unknown locality.
The collections from the P. N. du Garamba contain no Ancylorrhynchus ; M.
Fran9ois collected A . crux and an undescribed species from Urundi.

KEY TO THE SPECIES OF ANCYLORRHYNCHUS IN THE ETHIOPIAN REGION

1 Fourth posterior cell closed, with or without stalk ...... 2

- Fourth posterior cell open, even if narrowed ....... 5

2 Wings entirely dark funebris Bromley, 1936 : 135

- Wings pale in part ............ 3

3 Legs entirely black ....... susurrus Karsch, 1879 : 380

Legs entirely reddish ........... 4

4 Mesonotum red with a black cross. Ground colour of face yellow

cruciger Loew, 1857 : 348

- Mesonotum almost entirely black, with red only on humeri, postalar calli and hind

margin of scutellum nyukinus Speiser, 1910 : 89

5 Wings entirely pale ............ 6

- Wings at least partly dark, or uniformly but irregularly smoky .... 7

6 Mesonotum red, with a thin black median line ; scutellum red with median black

spot. Halteres black pretoriensis Bromley, 1836 : 135

272 H. OLDROYD

— Mesonotum black, with red colour on humeri, and posteriorly ; scutellum red with

black base. Halteres tawny yellow . . . maculatus Bigot, 1878 : 428

7 Legs entirely black. Wings indistinctly bicoloured ...... 8

— Legs not entirely black. Wings distinctly bicoloured, or entirely dark, or irregularly

smoky without definite paler areas ........ 9

8 Moustache pale. Scutellum black .... nomada Wiedemann, 1828 : 297

- Moustache black. Scutellum black, reddish at sides . unifasciatus Loew, 1857 : 349

9 Wings entirely dark, or uniformly smoky grey . . . . . . . 10

- Wings bicoloured . . . . . . . . . . . . 13

10 Abdomen black with sharply defined yellow (not red) hind margins, sometimes

broken into a pair of spots ; second segment with a broad band of white tomen-

tum, interrupted in middle ...... zonalis Bromley, 1936 : 137

— Abdomen without sharply defined bone-yellow bands . . . . . . 1 1

it Moustache pale ............ 12

Moustache black ....... rnunroi Bromley, 1936 : 135

12 Antennae and humeri reddish. Abdomen shining black dorsally, reddish ventrally

humeralis Wiedemann, 1821 : 235

- Antennae and humeri black. Abdomen bright red, except for black first segment

(an unnamed sp. from S. Africa)

13 Legs bicoloured ............ 14

Legs entirely reddish ........... 16

14 Moustache entirely black . .... quadrimaculatus Loew, 1857 : 348

- Moustache entirely or predominantly pale . . . . . . . . 15

15 Scutellum black ....... variegatus Bigot, 1878 : 429

- Scutellum partly red. Thorax with brown tomentum, unusual in this genus.

Hind femora red with longitudinal black stripe . . striatus sp. n. (p. 274)

1 6 Scutellum entirely reddish or tawny, Wings light brown, with costal border yellow.

Second abdominal segment with a pair of large grey patches . . . . 17

- Scutellum partly black, usually at base . . . . . . . . 18

17 Lateral borders of mesonotum black from behind humeri to transverse suture.

Abdomen black, posterior segments with orange hind margins, expanded laterally.
Palpi with black or brown hairs .... braunsi Bromley, 1936 : 136

- Lateral borders of mesonotum broadly red. Abdominal segments 1-4 black, with

narrow red hind margin ; fifth segment much brighter red. Palpi with red hairs
(tricolor Loew — either braunsi or reynaudii) reynaudii Macquart, 1838 : 48

1 8 Yellow base and brown tip of wings sharply divided . . . . . . 19

— Yellow and brown indistinctly merging, or dark area of indefinite extent, even if

contrasting in colour with yellow base . . . . . . . .21

19 Abdomen entirely black, segments 2-5 grey dusted. Large, robust species (25 mm)

Pteropleuron red. Dark tip of wing including half of total area, and entire discal

cell ......... magnificus Bromley, 1936 : 136

- Abdomen dull orange, first segment and base of second black, with other black

markings. Smaller (15 mm) ......... 20

20 First abdominal segment entirely black. Pteropleuron black

upicalis Curran, 1934 : 7 "\ . , . , .

yjt i v no difference m descriptions

splendens Bromley, 1936 : 137 j

- First abdominal segment partly red. Pteropleuron reddish

Two species recognized by Hermann but apparently undescribed

21 Mesonotum mostly black, red colour confined to humeri, postalar calli, and apex of

scutellum. Abdomen black, all segments broadly margined with reddish colour

fulvicollis Bigot, 1878 : 429

- Mesonotum mainly red, with a black cross . . . . . . . 22

22 Hypopleuron distinctly red ..... insignis Bromley, 1936 : 137

- Hypopleuron black, or only indistinctly reddish ....... 23

ASILIDAE OF THE CONGO BASIN

273

23 Fourth posterior cell wide open. Lateral arms of mesonotal black cross triangular.
Abdominal segments 2-5 silky yellow, with narrow, interrupted median black
stripe ......... crux Bezzi, 1908 : 377 (p. 274)

Fourth posterior cell only narrowly open. Lateral arms of mesonotal cross quadrate

hylaeformis Speiser, 1910 : 88

FIGS 44a~48. Ancylorrhynchus sp., proboscis. 44a, labium, showing dorsal groove ;
45, maxilla, with palp ; 46, tip of maxilla ; 47, glandular organ of palp ; 48, hypo-
pharynx, with detail.

274 H. OLDROYD

Ancylorrhynchus crux Bezzi

Ancylorrhynchus crux Bezzi, 1908 : 377.

URUNDI : Terr, de Bururi, i $, 19^.1952, colline Rumonge, loc. Imbo., alt.
780 m ; Rumonga, sable et brousailles pres du rive du Lac Tanganika, alt. 790 m,
i $, I9~20.vi.i948 (FJF).

Ancylorrhynchus striatus sp. n.

One of the rather few species of Ancylorrhynchus to have bicoloured legs, this
species is distinct from all others in having a narrow black longitudinal stripe on all
the femora, which are otherwise red.

$ Head in ground colour black, covered with dense yellow tomentum, leaving only a small bare
spot beneath the antennae, and one on each side of mouth-margin. Hairs of frons and face pale
yellow, including a shallow moustache on mouth-margin, linked with tracts of longer hairs on
each side of face. Hairs and bristles of occiput and beard, in contrast, deep red, bristles strong
and dense. Antennae red, third segment with blackish stripes, first two segments with yellow
hairs and bristles. Palpi reddish with black bristles. Proboscis red.

Thorax. Pronotum red anteriorly, black posteriorly and laterally. Mesonotum red, with
usual cruciform black markings, its transepts triangular, and a distinct brown median line ;
behind each humerus a barrow black crescent ; scutellum red with a transverse black basal
band ; bristles yellow, 4 notopleurals, 3 postalars. Entire dorsum covered with a white,
powdery dusting, which greatly obscures the pattern. Pleura entirely black with brown
tomentum, except for a reddish area beneath hind spiracles and halteres.

Abdomen. First six segments black, each becoming more reddish posteriorly, and with a
narrow red hind margin. Thick brown or yellowish brown tomentum, and numerous pale
yellow clothing hairs, but no strong bristles. Last two segments and terminalia orange, with
orange spines.

Legs. Coxae like pleura. Legs otherwise red, femora each with a narrow black stripe along
dorsal surface. Bristles yellow, becoming black on fore tibiae.

Wing. All cells open on margin, including anal (narrowly). From anterior cross-vein to
wing- tip is brown, and a further brown band runs across base of discal cell. Halteres bright
yellow.

Length of body 16 mm ; of wing 12 mm.

Holotype $. URUNDI : Nyamibu, Lac Tanganika, 24.^.1949 (FJF).
Paratype $>. Same data as holotype (FJF).

SCYLATICUS Loew

Scylaticus Loew, 1858 : 346, 349. Type-species : Scylaticus zonatus Loew, by designation of
Engel, 1929 : 369.

A genus of distinctive flies, bristly, with rounded head and inflated occiput.
Obviously closely related to Ancylorrhynchus, but having a straight, acute proboscis
of normal type instead of the characteristic beak-like proboscis of Ancylorrhynchus.
The third antennal segment, too, is different.

Scylaticus is a genus of rather arid places, and in the Ethiopian Region the species
are known from South Africa as far north as Nyasaland. Other species occur in

ASILIDAE OF THE CONGO BASIN 275

the Sahara and the eastern Mediterranean, as well as in India and in S. America.
It seems likely that the two populations in North and South Africa may be linked
through Kenya and Tanzania.

There are no specimens of Scylaticus in the collections from the P. N. du Garamba,
and only three specimens, all different, from the Katanga. These will be left for
future consideration, along with the S. African species. Engel (1932 : 276) pub-
lished a key to those species known to him.

GONIOSCELIS Schiner

Gonioscelis Schiner, 1866 : 670. Type-species : Dasypogon hispidus Wiedemann, 1819, by
original designation.

A very distinctive genus, generally resembling a small Stenopogon but recognized
at once, in both sexes, by the characteristic development of the fore femora (Text-fig.
49). This is obviously a highly efficient apparatus for seizing and holding prey.
The proboscis, though short, is stiff, acutely pointed, and slightly curved. The
eyes are close together, as in Stenopogon, and there is the same dusty, bristly
appearance and sandy colouring, appropriate to hunting in scrubby, arid areas.

There are no Gonioscelis in the collection from the P. N. du Garamba, but species
occur in Urundi and in the Congo Basin.

Gonioscelis maculipennis Engel

Gonioscelis maculipennis Engel, 1925 : 169.

TANGANIKA : Albertville, i $, 1-20.^.1919 (R. Mayne) (MRAC). Provisionally
assigned to this species, as its sex and condition make it impossible to identify
with more certainty.

Gonioscelis occipitalis sp. n.

(Text-fig. 49)

A red-legged species, with dull ashy grey-brown abdomen, distinguished from
related species by the white-tomented occiput, with black triangles and entirely
black, proclinate bristles.

$ Head. Even more strongly compressed than is usual in this genus, irons and upper part of
face forming a parallel-sided strip scarcely broader than the two antennae together. Frons
black, with bronze tomentum, and with proclinate hairs on each side, just above antennae.
Upper part of face, above facial hump, similar, with a band of strong black bristles medially ;
facial hump only moderately prominent, covered with white tomentum except at extreme lower
angles, where there is a bare, black spot on each side. Moustache mainly of strong black
bristles, with white ones on mouth-margin. Occiput covered with white tomentum, except for
a pair of large, shining black, bare triangles just behind vertex. From these triangles arise very
strong, proclinate, black bristles, with finer black bristles surrounding them, and a single row of
straight black bristles parallel to eye-margin, but some distance behind it. Beard yellowish
white ; palpi and proboscis black with black and white hairs. Antennae with first two seg-
ments blackish with black hairs (rest missing).

276

H. OLDROYD

Thorax. Mesonotum ashy, more black-bronze anteriorly, more whitish posteriorly. Clothed
with very short black bristles and also with 3 pairs of dorsocentrals, 3-4 notopleurals, 3 supra-
alars, all black ; 2-3 strong yellow postalars. Marginal scutellars black, one strong pair and
one weaker. Pleura with no strong hairs or bristles.

Abdomen dully shining black-brown through thin ashy grey tomentum, and covered with
short, stiff black bristles, with longer tufts at sides of first and second segments. Segments 7, 8
shining reddish brown and forming part of ovipositor. Venter similar.

Legs. Coxae like pleura. Fore and middle coxae with strong white bristles, third with a few
black bristles. Fore femora black basally on anterior surface ; mid femora a little so ; legs
otherwise reddish yellow, even to tips of tarsi. Short clothing hairs white, bristles black.

Wings uniformly smoky brown. Halteres reddish.

Length of body 13 mm ; of wing 10 mm.

Holotype $. LULUA : Sandoa a Kapanga, ix.ig28 (Dr. Walker) (MRAC).

Gonioscelis tomentosus sp. n.

A black, dark-winged species, in general appearance resembling only G. nigripennh
Ricardo, but distinguished from that species by having the thorax not shining black,
but covered with tomentum, that of the pleura especially being dull bronze. From
G. lacertosus Engel, which also has dark wings, G. tomentosus is distinguished by the
leg-colour, the legs of lacertosus being almost entirely orange.

FIG. 49. Gonioscelis occipitalis, fore femur.

ASILIDAE OF THE CONGO BASIN 277

? Head black, except for conspicuously orange third segment of antenna. Frons and face
very narrow, touching bases of antennae and ocellar tubercle, covered with golden tomentum, a
little bare in centre of face and on buccae, with black hairs and bristles. No hairs or bristles on
frons except on ocellar tubercle. Moustache entirely black, or with one or two white bristles
ventrally, mounted on a slight facial tubercle, which merges gradually with upper face. Vertex
narrow, rather flat, with no bare areas, and with black and yellow bristles not in an orderly
arrangement. Antennae black basally, with black hairs, third segment and part of second red.

Thorax. Ground colour black, with orange humeri and dull reddish lateral margins, including
postalar calli. A dull bronze tomentum exposes a darker pattern of a divided broad median
stripe and lateral spots. Mesonotum clothed with short, spiky black bristles; longer bristles
may be black or dull red, latter especially laterally ; three pairs of strong postsutural dorso-
centrals with one or two feeble pairs anteriorly. Scutellum black, with one pair of black
marginal bristles. Pleura uniformly covered with bronze tomentum, without dark areas or
other pattern.

Abdomen red-black, or mahogany coloured, with base of first segment and hind margins of
others a lighter red ; dully shining through thin grey tomentum which becomes visible in certain
directions of the light. Clothing hairs short, erect, black, with longer tufts at sides of first and
second tergites.

Legs. Fore coxae with bare, shining black streak posteriorly, and a smaller spot antero-
basally. Trochanters black, legs otherwise red, with fore and middle femora extensively black
anteriorly on basal half ; hind femora and posterior face of middle femora obscurely darker.

Wings dark brown, with clearer centres in some cells. Halteres red.

Length of body 15 mm ; of wing 10 mm.

Holotype $. LULUA : Kapanga, viii.1932 (F. G. Overlaet) (MRAC).
Paratype $. KATANGA : Fokele, 2g.xi.i9ii (Dr. Bequaert) (MRAC).

Gonioscelis congoensis sp. n.

(Text-figs soa, 5ob)

This species is described, in spite of the poor condition of the available material,
because it is confusingly close to genitalis Ricardo, from South Africa. The two
species may be distinguished by the male genitalia (Text-figs 5oa, 5ob).

cJ Head. Frons and face very narrow, hardly diverging towards mouth-margin. Ground
colour black, covered with tawny tomentum and tawny hairs and bristles. Ocellar tubercle
with a few black bristles. Moustache entirely tawny. Antennae black basally, with tawny
hairs, becoming more reddish apically (third segment broken off in 2 <$ available). Proboscis
and palpi black with yellowish hairs. Occipital hairs and bristles entirely tawny.

Thorax mostly red, with broad, divided median black stripe and large lateral black spots ;
scutellum black. This pattern mostly obscured by brassy yellow tomentum. Bristles long,
tawny, dorsocentrals postsutural only. Pleura uniformly covered with dense, brown tomentum.

Abdomen dully shining through ashy tomentum, which leaves segmentations yellow : seventh
and eighth segment more shining. Clothing hairs short, tawny ; venter similar. Male
genitalia as in Text-fig. 5oa.

Legs predominantly red, but with femora black (specimens badly preserved). Fore and
middle femora not marked with a strong black stripe as in genitalis Ricardo.

Wings almost uniformly brown, a little darker in marginal cell.

Length of body 15 mm ; of wing 10 mm.

$ Similar. Third antennal segment reddish. Seventh and eighth segments incorporated in
ovipositor.

Holotype <£. LULUA : Luluaborg, 21^.1939 (/. /. Deheyn] (MRAC).

278

H. OLDROYD

Paratypes. LULUA : Luluaborg, i <$, i8.v.igi2 (P. Callewaert) ; Wombali, i $,
vi.igi3 (P. Vanderijst] ; Manyema, i $ (R. Mayne) ; Bolobo, Makamandulu, i $,
1938 (Dr. Schouteden) (MRAC).

Gonioscelis francoisi sp. n.

(Text-fig. 51)

Distinguished from the other species of Gonioscelis in the Congo Basin by the fact
that the dorsocentral bristles extend forwards in front of the transverse suture of
the thorax ; hairs of the head predominantly black, and hind femora slender and
black.

cJ Head. Narrow, frons about twice as broad as ocellar tubercle, covered with yellowish
tomentum and with a row of black hairs along each eye-margin. Face diverging little above

FIG. 5oa. Gonioscelis congoensis, <$ genitalia.

FIG. 5ob. Gonioscelis genitalis, $ genitalia.

ASILIDAE OF THE CONGO BASIN

279

facial knob, and then sharply down to mouth-margin ; facial knob slight, with moustache
usually entirely black, sometimes pale ; above it, reaching nearly up to bases of antennae, are
other black hairs ; lower ones shorter and more bristly, a tuft beneath antennae longer, prom-
inent, and rather silky. Occiput with yellow tomentum and two large, diamond -shaped,
shining black patches (cf. occipitalis sp. n.) ; occipital bristles mixed black and yellow, a multi-
serial row some distance behind eye-margin. Antennae, proboscis and palpi black, with black
hairs.

Thorax black with brassy yellow tomentum. Mesonotum without pattern, though with faint
indications of a median stripe. Covered with very short black bristles ; 5 pairs of long, strong
black supra-alars ; a notopleural supra-alar row of 4-5 black ; 2 postalars, yellow or black.
Scutellum with one pair of black bristles slightly before posterior margin. Pleura with brassy
tomentum and no long hairs.

Abdomen. Dorsum black, with yellowish grey tomentum, posterior margins of segments
darker, at least medially ; seventh and eighth tergites more shining blackish. Clothing hairs
short, yellow ; longer tufts on first and second segments black. Venter shining black, with
short yellow hairs. Male genitalia red, short, as in Text-fig. 51.

Legs. Fore and middle coxae with black hairs, hind coxae with yellow hairs. Femora
mainly black ; fore and middle femora red posteriorly and apically, hind femora long, slender,
and entirely black. Tibiae and tarsi reddish, obscurely darkened, especially on posterior faces
of tibiae. Clothing hairs yellow, bristles mostly black.

Wings. Dark brown with paler centres to many cells, especially to first basal cell.

Length of body 13 mm ; of wing 9 mm.

$. Quite similar. Ovipositor includes tergites 8, 7 and most of 6.

Holotype $. URUNDI : Kisenyi, Busoni, 1800 m, ly.xii.igso (F. J. Franfois)
(FJF). '

Paratypes. Same data as holotype, i $.

A <£ paratype from URUNDI, Kitega, 1700 m, 21. x. 1950 (F. J. Franfois) has the
fore and middle femora less heavily darkened. A $ paratype from Kitega, 1720 m,
i.xii.i95o has the colouring of the preceding female, but with the moustache entirely
white. There is evidently some degree of variation in this species.

FIG. 51. Gonioscelis francoisi, <$ genitalia.

28o H. OLDROYD

RHABDOGASTER Loew

Rhabdogaster Loew, 1858 : 346, 351 ; Engel, 1929 : 168. Type-species : Rhabdogaster nudus
Loew, monotypic.

Described both by Loew and by Engel as resembling a Leptogaster, but with
complete pulvilli, and with acanthophorites in the female. The male genitalia
are of a rather distinctive structure, with the epandrium divided into two large
lobes, and with long, curved aedeagus (Text-fig. 52). Posterior to the hind coxae,
beneath the base of the abdomen, is a heavily sclerotized arch instead of the usual
membranous area : according to Hull's key (1962 : 119) this character distinguishes
Rhabdogaster from related genera.

Two of the known species of Rhabdogaster — nudus Loew (1858) and maculipennis
Engel (1929) — are both small or very small (7-8 mm), delicate flies, pale yellow in
colour. The new species is larger than this (10 mm), and shining black, and so is
close to nitidus Hull (1967), from S.W. Africa, which I have not seen.

Rhabdogaster major sp. n.

(Text-fig. 52)

$ Head in front view relatively broad, but not as broad as that of Xenomyza : breadth of
frons : height of frons + face = 2:1 instead of 2-5 : i. Frons and face parallel-sided, more
than half as broad as an eye ; shining black in ground colour, with sparse yellow tomentum.
Face has no perceptible facial knob, but a moustache of sparse white hairs occupies basal half,
and is surmounted by very short, scaly yellow hairs which reach up to antennae. Hairs and
weak bristles on ocellar tubercle, vertex and occiput, all white or pale yellowish. Proboscis

FIG. 52. Rhabdogaster major, $ genitalia.

ASILIDAE OF THE CONGO BASIN 281

and palpi black, with whitish hairs. Antennae black, with yellow hairs, third segment slender,
pointed, as long as first two together ; style almost two-thirds as long as third segment.

Thorax. Mesonotum shining black, with only small areas of yellow tomentum, of which
humeral triangles are the most conspicuous ; a narrow line of tomentum in suture at base of
scutellum, and at extreme lateral margins of mesonotum. Extremely short, fine yellow hairs
along lines of acrostichals and dorsocentrals, and on transverse suture. Two strong noto-
pleurals, a cluster of four or five prescutellars, but no marginal scutellars. Scutellum shining
black, rather convex, with a deep submarginal groove. Pleura also shining black with very
thin white tomentum : tufts of white hairs on pronotum, propleuron, upper sternopleuron and
before halteres.

Abdomen elongate, cylindrical, much longer than wings, which reach only to tip of fifth
segment. First five segments bare, shining black ; sixth-eighth segments with dark brown
tomentum ; all tergites with short black hairs. Venter with pale yellowish tomentum and
whitish hairs. Genitalia as in Text-fig. 52, prominent.

Legs predominantly black, but knees and hind tibiae and metatarsi red or reddish, especially
posteriorly. Hairs and bristles of legs almost entirely white.

Wings. Venation simple : anal cell closed, with short stalk, but all other cells open on wing-
margin ; discal cell long and narrow, its origin almost as far back as fork of RI/RZ+S. Veins
brown or black. Membrane clear and apparently colourless, but with microtrichiae scattered
over most of area except basally. Halteres orange.

Length of body 10 mm ; of wing 7 mm.

9 Similar. Ovipositor simple, with acanthophorites and a small ventral keel.

Holotype <£. GARAMBA NATIONAL PARK : P.N.G., 3480, Inimvua, 16^.1952
(IPNC).
Paratype $. Same data as holotype.

Tribe STICHOPOGONINI

This is a small tribe of interesting Asilidae, set apart from Saropogonini by
having the prosternum complete, or nearly so, and firmly bridged to the propleuron.
The female terminalia are characteristic (Text-figs 54, 55), with the eighth sternite
produced like the hull of a boat, and often with a distinct keel, surmounted by a
crown of spines, and sometimes terminated with a dense brush of silky hairs.
Though some Saropogonini have a similar structure it is not so fully developed.

The most obvious characteristic of this tribe is the saddle-shaped excavation of
the vertex, as a result of which the eyes are much more widely separated at the
vertex than opposite the antennae (Text-figs 53, 56, 57, 60).

Hull (1962 : 104) recognizes thirteen genera in this tribe, including Eremodromus
Zimin and Turkmenomyia Paramonov, which were unknown to Hull, and omitted
from his key. Lasiopogon, though apparently correctly assigned to this tribe,
stands apart from the rest, and is a genus of Holarctic robber-flies that hunt in
grassy areas from shrubs and low herbage. The rest of the tribe are essentially
xerophilous, and are to be found sitting in full sunlight on sand, or on the stones of
a dry stream-bed. For concealment they rely on their cryptic grey colouration
and on the dazzling effect of the light. They sit motionless until potential prey
flies overhead, and it may be that the characteristic broadening of the frons and
separation of the eyes at the vertex is a device to improve stereoscopic vision directly
overhead.

282

H. OLDROYD

Apart from Lasiopogon, the other genera are basically very much alike, and should
perhaps be reduced to subgenera of Stichopogon. The differences are small details
of antennal structure, of the arrangement of the moustache, of the wing- venation,
and of the presence or absence of pulvilli. The last character is somewhat elusive
outside the tribe Leptogasterinae, occurring sporadically, and perhaps not even of
generic significance ; for instance in the genus Glyphotriclis.

Though especially characteristic of arid areas, Stichopogon occurs throughout all
the tropical and warm-temperate countries of the world, and is often found in
stream-beds in well-watered country. In such areas, however, the number of
species appears to be small, and individual species have a wide, if scattered, dis-
tribution.

FIG. 53. Stichopogon caffer, head and antenna.

FIG. 54. Stichopogon caffer, $ terminalia.

ASILIDAE OF THE CONGO BASIN

283

Hull (1962) records seven species from the Ethiopian Region. Of these Dasypogon
dilutus Walker was removed to Stenopogon by Miss Ricardo ; 5. grossus from the
Durban Bluffs is clearly a Clinopogon, and is possibly a synonym of C. nicobarensis
Schiner, widely distributed round the shores and islands of the Indian Ocean. The
remaining five species may be separated as follows:

KEY TO THE SPECIES OF STICHOPOGON OF THE ETHIOPIAN REGION

N.B. — Some of the species of the arid region of Egypt and the Near East may extend into
eastern Africa.

1 Frons and face very much constricted at antennae (Text-fig. 60). Legs entirely black

unicolor Ricardo, 1925 : 276

- Frons and face less heavily constricted. Legs not entirely black .... 2

2 Femora mainly or entirely red .......... 3

Femora mainly or entirely blackish ... 4

3 Wings with a diffuse brown patch stretching from 7?i to R±

maculipennis E. & C., 1939 : 188

- Wings clear ......... hermanni Bezzi, 1910 : 145

4 Grey bands of abdomen unequal, with prominent bands on segments i, 2, 4, 5.

Males with small, but very distinct spot on fork of #4 +5- Females with trace of
dark spot, and with eighth abdominal tergite almost entirely tomented ; ventrally
with conspicuous tuft of silky golden hairs (Text-fig. 55)

punctutn Loew, 1851 : 15 (p. 284)

- Grey bands of abdomen incomplete, but equal, without any segments especially

prominent. No dark spot on wings. Females with eighth tergite mostly bare and
shining, tomented only dorsally, and without prominent golden tuft (Text-fig. 54)

coffer Hermann, 1907 : 3 (p. 283)

The collection from the P.N. du Garamba contains only one species of Stichopogon,
the widespread S. pundum Loew, though 5. coffer Loew and S. hermanni Bezzi also
occur in the Congo Basin.

Stichopogon caffer Hermann
Stichopogon caffer Hermann, 1907 : 3.

LOMAMI : Luputa, i <£, 2 <j>, xii. 1934-1. 1935 (Dr. Bouvier) ; Kanjama, i $, 1931

FIG. 55. Stichopogon punctum, $ terminalia.

284

H. OLDROYD

(R. Massart) ; Coquilhatville, Basankusu, i <j>, 1949 (ten Bunderen) ; UBANGI :
Nouvelle, Anvers, i <$, g.xii.igsa (P. Basilewsky) ; STANLEYVILLE : i <j>, 4.111.1928
(A. Collart) (MRAC). URUNDI : Minago, i <£, 5.^.1949, 730 m (FJF).

Stichopogon punctum Loew
(Text-figs 56, 58)

Stichopogon punctum Loew, 1851 : 15.
Stichopogon punctatus Loew, 1852 : 658.

GARAMBA NATIONAL PARK : 1461, II/fc/i8, i $, 28.111.1951 ; 2910, Il/fd/iy, i <J,
I4.xii.i95i ; 3177, II/gb/i4, 2 ?, 8.111.1952 ; 5262, II/fe/i8, i ?, 31.111.1952 ; 214,
I/b/2, i <J, 22.11.1950 ; 1223, II/ed/i7, i & 6.11.1951 ; 1167, II/fc/5, i ?, 31.1.1951 ;
94, I/b/3, i $, 11.1.1950; 199, I/a/3, i $, 7.111.1950 (IPNC). The last two females
are distinctly larger than the others, but seem otherwise conspecific.

BAS-CONGO : Boma, 2 $, v.1913 (Dr. Bequaert] ; KWANGO : Popkabaka, 2 $,

FIGS 56-59. 56, Stichopogon punctum, head. 57. Stichopogon hermanni, head.
58, Stichopogon punctum, antenna. 59. Stichopogon hermanni, antenna.

ASILIDAE OF THE CONGO BASIN

285

iii.1952 (L. Pierquiri) ; Mayidi, i $, 1942 (P. van Eyen) ; TANGANIKA : Mpala,
780 m, 2 $, vii/viii. 1953 (H. Bomans) ; Dt. de BONGALA : i $, viii.i92o (L.
Burgeon) ; Stanleyville, i $, 9.111.1927 (A. Collart) (MRAC).

URUNDI : Nyanza Lac, alt. 780 m, 3 $, 5 $, 2o.ix.i948 ; Usumbura, Lac, 3 $

Stichopogon hermanni Bezzi

(Text-figs 57, 59)

Stichopogon hermanni Bezzi, 1910 : 145.

COQUILHATVILLE : Eala, i $, 111.1932 (H. J. Bredo}. (MRAC)

Tribe XENOMYZINI

These are the ' goggle-eyed ' flies referred to in the Introduction, distinguished to
the naked eye by the reduction of frons and face, and antero-posteral flattening of
the head into a disc in which the eyes are large and prominent.

These features differ only in degree from conditions in many other genera of
Asilidae, and are sometimes difficult to assess : it is generally easy to say when
flies are ' goggle-eyed ', but not so easy to be certain when they are not. In the
Ethiopian Region the prosternum is fully bridged to the pronotum in Xenomyzini,
but this is not necessarily true in other Regions.

My key to genera (Oldroyd, 1963 : 7) recognized eight genera in this tribe, but a
later revision of the genus Rhipidocephala (Oldroyd, 1966) rejected Holcocephala as
not African, and merged Margaritola and Paroxynoton into Rhipidocephala. This

FIG. 60. Stichopogon unicolor, head and antenna.

286 H. OLDROYD

reduced the Xenomyzini of the Ethiopian Region to four genera : Oxynoton
Janssens ; Rhipidocephala Hermann ; Xenomyza Wiedemann and Oligopogon Loew.

KEY TO GENERA OF AFRICAN XENOMYZINI

1 Third antennal segment small, with long arista. Anal cell closed and stalked ;

fifth posterior cell making contact with discal cell . XENOMYZA Wiedemann (p. 287)
Third antennal segment elongate, with apical style. Anal cell open, or barely

closed ; fifth posterior cell not making contact with discal cell. (Text-fig. 69) . 3

2 Mesonotum with an exaggerated hump, strongly projecting forwards. Ovipositor

with spines (acanthophorites) .... OXYNOTON Janssens (p. 286)

Mesonotum in side view symmetrically rounded ....... 3

3 Second segment of antennal style with long hairs. Abdomen elongate-cylindrical.

Ovipositor with spines (acanthophorites) . . . OLIGOPOGON Loew (p. 294)
- Both segments of antennal style with short hairs. Abdomen at most twice as long
as broad. Ovipositor without acanthophorites

RHIPIDOCEPHALA Hermann (p. 286)

OXYNOTON Janssens
Oxynoton Janssens, 1951 : i. Type-species : O. francoisi Janssens, monotypic.

Remarkable for the excessive development of the mesonotum into a strong hump.
The female, like the female of Oligopogon, has a crown of spines on the ovipositor.
Only one species of Oxynoton is so far known, originally described from Urundi.

Oxynoton francoisi Janssens

Oxynoton francoisi Janssens, 1951 : i.

TANGANIKA : N.E. Kondoa, i $, May 55 ( J.F. Lamerton) ; Gonja, May 58, 2 <$,
(/. D. Phipps) (BMNH).

RHIPIDOCEPHALA Hermann

Rhipidocephala Hermann, 1926 : 174 ; Oldroyd, 1966 : 149. Type-species : Rhipidocephala
angustior Oldroyd, 1966, by original designation as analis Macquart sensu Hermann, nee
Macquart.

Paroxynoton Janssens, 1953 : n. Type-species : P. tigrinum Janssens, by original designa-
tion.

Margaritola Hull, 1958 : 255. Type-species : Margaritola mirabilis Hull, 1958, by original
designation.

A revision of this genus was published by Oldroyd (1966), and the following
records relating to the Pare National du Garamba are extracted from it :

Rhipidocephala tigrina (Janssens)

Paroxynoton tigrinum Janssens, 1953 : 12.

URUNDI : Gihanga & Bubanza (IRSNR).

ASILIDAE OF THE CONGO BASIN 287

Rhipidocephala mono Hermann
Rhipidocephala morio Hermann, 1926 : 180.

KATANGA : Mulungivishi, i $, 1.1931 (G. F. de Witte) (MRAC).
Rhipidocephala congoiensis Oldroyd

Rhipidocephala congoiensis Oldroyd, 1966 : 164.

KASAI : 4^, 3 <j>, 1928 (D. Walker) ; Dolo, 4^, i $, xi.igia (F. Chaltin] ; LOMAMI :
Katompe, 2 ?, I2.xiii.i923 (M. Bequaert) (MRAC).

Rhipidocephala scutata Oldroyd

Rhipidocephala scutata Oldroyd, 1966 : 166.

GARAMBA NATIONAL PARK : P.N.G., 469, I/a/i, i $, i.v.igso (G. Demouliri) ;
1588, II/hc/4, 10 $, 4 0-, 20.iv.i96i (/. Verschuren] ; 1824, H/fd/27, * <$> 28^.1951 ;
1887, II/gd/7, i ?, 8.vi.i95i ; 3323, Pidigala, 2 3, 23.^.1952 ; 3447, II/gd/4, 2 <$,
i 9, 8.V.I952 ; 3678, Ndelele, i $, 2 9, 4.i8.vi.i952 (IPNC).

Kivu : Uviva, 3 J, 4 9, xi.i922 (CA. Seydel] ; 16-23.111.1953 (P. Basilewsky]
(MRAC).

XENOMYZA Wiedemann

Xenomyza Wiedemann, 1817 : 60. Type-species : Damalis planiceps Fabricius, 1805, by

designation of Coquillet, 1910.
Damalis auctt. nee Fabricius, 1805 : 147.

The name of this genus is usually Damalis, but the original concept of Damalis
confused Asilids and Empids, and Westwood, 1835 designated as type of Damalis
the first species, D. curvipes Fabr., which is an Empid. I therefore follow the
example of Carrera, and use Wiedemann's name Xenomyza for this genus in Asilidae.

The species that have been described from Africa are nearly all from Southern
Africa, and none of them appear to conform with any of the four species recognized
below, which are all therefore described as new. The four are easily separated on
genital characters as well as by colour differences, though, as I explain under the
specific description, it is probable that specimens I have included in taciturna sp. n.
should be separated into more than one species.

KEY TO SPECIES OF XENOMYZA IN THE CONGO BASIN

1 Hind femora noticeably swollen, and with strong bristles ventrally (Text-figs 65, 66) 2

- Hind femora not noticeably swollen, ventral bristles not noticeably strong. Wings

typically colourless, but in some included specimens they have a yellow tinge

taciturna sp. n. (p. 289)

2 Hind femora slender in basal half, expanding considerably in distal half (Text-fig. 65)

amphora sp. n. (p. 290)

- Hind femora expanded over entire length (Text-fig. 66) ..... 3

3 Mesonotum with three shining black stripes, broad, and touching each other ;

scutellum covered with white tomentum .... poseidon sp. n. (p. 291)

- Mesonotum tomented, but scutellum bare and shining . . scutellata sp. n. (p. 292)

288

H. OLDROYD

FIGS 61, 62. Xenomyza taciturna, <$ genitalia. two variants in shape.

ASILIDAE OF THE CONGO BASIN 289

Xenomyza taciturna sp. n.

(Text-figs 61, 62)

A variable species, by far the most numerous in the present collections, and
showing a considerable range of size and colour. It may be that more than one
species is confused here, but the characteristic male genitalia vary between the two
types shown in Text-figs 61, 62, without any constant differences.

o* Head. Ground colour brown, covered with brown tomentum ; frons entirely tomented,
except for ocellar tubercle ; face mostly bare and shining brown except for upper third. Vertex
with long and fairly dense black hairs. Face with very sparse brown hairs and a moustache
consisting of only 5-6 black bristles in a single row. Antennae black-brown with black hairs.
Proboscis and palpi black-brown but with pale hairs. Occiput with grey tomentum and white
bristles.

Thorax brown, entirely covered with tomentum except for edges of humeri and of postalar
calli. Tomentum of mesonotum whitish, with a pattern of three stripes in brown. Scutellum
covered with whitish tomentum. Pleura with grey tomentum and a dense tuft of yellow hairs
on metapleuron ; other pleura with only sparse yellowish hairs.

Abdomen black-brown in ground colour, covered uniformly with yellowish grey tomentum
except for a transverse band basally on second segment. Hairs sparse, yellowish, but not
noticeably longer laterally. Venter similar. <$ genitalia as in Text-figs 61, 62, shining red-
brown, contrasting with dull, blackish abdomen.

Legs clear mahogany-brown with rather long, yellowish clothing hairs, and a few fine black
bristles. Hind femora as in Text-fig. 66.

Wings clear, colourless. Halteres yellow, including knob.

Length of body 8 mm ; of wing 7 mm.

Holotype <$. GARAMBA NATIONAL PARK : P.N.G., 3461, Inimvua, 16^.1952

(IPNC).

Paratypes. 3488, same data as holotype, 5 $, 16 ? (IPNC) ; Garamba National
Park, 3387 Mt. Embe, 3 $, i7-2i-iv.i952 ; 3351, 3352 Pigidala, 3 $, 22-iv-i952 ;
3476» 3514. 35*5 Aka/2, 3 ?, 22-V-I952 ; 3481, 3499, Dedegwa, 2 $, 2i-v.i952 ;
3941 II/gc/6, i $, i4-viii-i952 ; unnumbered, 1949-52 ; 469 I/a/i, i $, i-v-igso
(G. Demoulin) (IPNC).

This species shows considerable variation in size and colour, partly arising from
differences in the age and state of preservation of the specimens, but with un-
usually large variations even after allowing for these factors. This is particularly
true of the specimens in the collection of the Muse'e R. de 1'Afrique centrale, many
of which are considerably bigger and more yellow than the specimens from the
P.N.G. For the present I provisionally identify all specimens with the male genitalia
notched as in Text-figs 61, 62 as belonging to taciturna, but I feel sure that eventually
it will be possible to set aside two, if not three species in this complex. The Congo
specimens include a good series from Kifumashi (most nearly resembling the typical
series from P.N.G.) ; Kapanga ; Mayumbe ; Stanleyville ; Elisabethville and
Mayidi. I do not list these doubtful specimens as paratypes.

2 go

H. OLDROYD

Xenomyza amphora sp. n.

(Text-figs 63, 65)

Distinguished from the other species by the characteristic shape of the hind femora,
which are narrow and tubular for the basal half, abruptly swollen and flask-shaped
apically.

3 Head. Frons and face dull brown, entirely tomented ; without obvious hairs except for a
fringe of short black hairs across vertex, and on ocellar tubercle ; very short black hairs on face
are visible only at an oblique angle. Moustache consists of four black bristles on epistoma.
Antennae with first segment shining brown, second tomented black-brown, both with black
hairs ; third segment shining black-brown, aristiform extension white apically. Proboscis and
palpi brown with black hairs. Occiput red in ground colour with thin brown tomentum and
fine black hairs.

Thorax. Mesonotum dark brown, yellow on humeri, postalar calli, scutellum and a pre-
scutellar area. Also with faint traces of two narrow black stripes, with brown tomentum but
with no obvious hairs. Pleura dull, yellow and brown, entirely tomented, hairs only on meta-
pleuron before halteres.

Abdomen strongly constricted basally, with whole of second segment and base of third
narrowed, third segment then broadening abruptly ; posterior abdomen has an oval outline.
Shining red-brown, more yellow laterally, and all the broader segments with close-lying short
brown and black hairs. Venter reddish with a few short black hairs. $ genitalia (Text-fig. 63)
downturned, with on each side a long process and a cluster of three strong black bristles.

Legs blackish brown, translucent. Basal half of hind femora and hind tibiae more yellowish,
slender, apically clavate and blacker. Hind femora with a few strong, short spines ventrally
near tip (Text-fig. 65), and other bristles on enlarged hind trochanters. Bristles and hairs of
legs black, sparse.

Wings uniformly dark brown, with dense microtrichiae on both surfaces. Halteres with red
stem and black knob.

FIG. 63. Xenomyza amphora, $ genitalia.

Length of body 9 mm
$ closely similar.

ASILIDAE OF THE CONGO BASIN

of wing 9 mm.

291

Holotype <$. UELE : Bambesa, ix-xii.i933 (H. J. Bredo) (MRAC).

Paratypes. Same data as holotype, 2 $ ; LEOPOLDVILLE : Kikwit, i $,
(P. Vanderijst) ; TERR. DE KASONGO : R. Lumami, i $, x-xii.igso (Benoit) (MRAC).

Xenomyza poseidon sp. n.

(Text-figs 64, 66)

A brilliant little species, with most of mesonotum and legs shining black. It has
hind femora like those of scutellata sp. n., but has the colour-differences shown in
the key.

c£ Head shining black in ground colour, with dense pale yellow tomentum, which leaves
certain clearly denned areas bare, shining black : a narrow triangle surrounding ocellar tubercle ;
a transverse line through bases of antennae ; lower third of face, underlying all swollen facial
tubercle. Moustache consisting of 4-6 isolated black bristles ; short hairs pale, present only
at vertex and on parafacial areas. Antennae black, third segment and apex of second more
mahogany-brown, apical half of arista white ; first two segments with black hairs, third
segment elongate, tapering. Palpi black with black bristles ; antennae black-brown with
yellow hairs. Occiput entirely tomented grey, with a single row of light yellowish occipital
bristles.

Thorax shining black, with dense yellowish white tomentum, which leaves certain areas bare
and shining black ; mesonotum with a broad, divided, median stripe, flanked by the usual
double spot on each side, but all stripes end abruptly opposite wing-bases. Scutellum covered
with tomentum except for a very narrow black rim. Numerous very short black bristles in
clusters, especially thick on upper supra-alar area. Pleura similarly covered with tomentum,

FIG. 64. Xenomyza poseidon, $ genitalia.

292 H. OLDROYD

but without short spines, leaving bare two large areas, one on mesopleuron and one immediately
behind it, on pteropleuron. Pronotum, propleuron and metapleuron with fine pale yellowish
hairs. No marginal scutellar bristles are visible.

Abdomen without a constriction (cf. amphora sp. n.), but second segment distinctly grooved
transversely. Abdomen, like thorax, black in ground colour, with areas of white tomentum
anteriorly and laterally on each segment (only laterally on second segment), and with areas of
short spines posteriorly and laterally on all segments. Venter shining black with posterior
tomentum on each segment, and with short yellowish hairs. Genitalia as in Text-fig. 64.

Legs black, middle and hind tibiae orange, darker at tip. Hind femora as in Text-fig. 66,
swollen along whole length, and with a double row of strong black spines ventrally. Other
femora less swollen, and with only isolated spines. Bristles and most hairs black, but many
clothing hairs yellow.

Wings almost uniformly brown, alula a little paler. Halteres orange including knob.

Length of body 10 mm ; of wing 9 mm.

? Similar, knob of halteres dusky.

Holotype $. KATANGA : Elisabeth ville, I2.xii.ig24 (MRAC).

Paratype $. Elisabeth ville, Lubumbashi, 20.xii.i92o (Dr. M. Bequaert] (MRAC).

Xenomyza scutellata sp. n.

(Text-fig. 67)

Shares with poseidon sp. n. the uniformly swollen hind femora, with double row
of ventral spines, but instead of being shining black, the entire legs are honey-
brown, with long, recumbent clothing hairs ; differs also in having the mesonotum
more tomented but the scutellum, in contrast, bare, shining brown.

<J Head black-brown, covered with dense brown tomentum, frons and face with no clearly
defined bare areas. Venter with a small number of long, black hairs : frons without long hairs ;
face covered with rather long, pale yellow hairs, which are reclinate, and grow upwards along
face. Moustache consists of a double row of yellow bristles on mouth-margin. Antennae
black : first segment with a long, black bristle ; second with a shorter brown one ; aristiform
extension long, white apically. Proboscis and palpi black with yellow hairs.

Thorax shining black in ground colour, entirely covered with tomentum, but in preserved

FIGS 65, 66. Xenomyza spp., hind femur of <$. 65, X. amphora ; 66, X. poseidon.

ASILIDAE OF THE CONGO BASIN

293

specimens this may be rubbed away in irregular areas. Tomentum is bronze, more yellow
towards sides and posteriorly : in contrast, humeri, posterior calli and scutellum are all bare,
shining brown. Yellow bristles, though fine, are relatively strong for the genus Xenomyza,
and follow lines of dorso-centrals and acrostichals, as well as clustering laterally. Scutellum
with 5-6 quite strong yellow marginals. Pleura covered with tomentum, yellowish, partly
brownish, and with unusually abundant pale yellow hairs on pro-, meso- and metapleura.

Abdomen shining black in ground colour, with bronze tomentum. First segment with well-
developed transverse membranous area, posteriorly bare and shining ; other segments only
irregularly shining where rubbed. Almost bare of hairs on disc, but tufts of longer yellowish
hairs laterally ; ventrally with whitish tomentum and pale yellowish hairs. Genitalia as in
Text-fig. 67.

Legs honey-brown, femora and tarsi rather darker than tibiae. Hind legs noticeably long and
thick compared with those of fore and middle legs ; hind femora strongly and uniformly
swollen, with a double row of short, black spines ventrally. Otherwise legs mostly covered
with rather long yellow bristles and long, recumbent yellow clothing hairs ; only tarsi with
substantial numbers of black bristles.

Wings smoky black-brown, more intensively so anteriorly. Halteres yellow with whitish
knob.

Length of body 7 mm ; of wing 7 mm.

$ Similar.

HolotypecJ. GARAMBA NATIONAL PARK : P.N.G., 3450, Aka, 14^.1952 (IPNC).

Paratypes. GARAMBA NATIONAL PARK : P.N.G., Aka, 2 $, 22^.1952 (IPNC) ;
Eala, i $, ix.igao (Dr. P. Staner) ; Wamba, i $, 1936 (Dr. Degotte) ; Bambesa, i $,
3O.viii.i933 (/. V. Leroy) (MRAC).

FIG. 67. Xenomyza scutellata, <$ genitalia.

294

H. OLDROYD

OLIGOPOGON Loew

Oligopogon Loew, 1847 : 497. Type-species: O. hybotinus Loew, monotypic.

This is a strange, small genus, which has attributes of both Stichopogonini and
Xenomyzini. The female genitalia closely resemble those of Stichopogon (Text-fig.
68), but the head is quite unlike the characteristic saddle-shaped vertex and scaly
moustache of the Stichopogonini. Instead, it is goggle-eyed, with a fringed antennal
style, and is much like Rhipidocephala of the Xenomyzini. In fact, Oligopogon
looks like a head and thorax of Xenomyza, with an abdomen of Stichopogon and legs
of a Saropogonine genus such as Holopogon. Engel (1929 : 372) says that Oligopogon
has strong discal bristles on the abdomen, and in his key he gives these as a means
of separating Oligopogon from Rhipidocephala, but there are no such bristles in any
specimen that I have seen.

One constant character of Oligopogon is the small size of the discal cell (Text-fig.
69), a feature which excludes 0. atrum Bigot from the genus.

The type-species, 0. hybotinus Loew, was described from the island of Rhodes,

69

FIGS 68-69. 68, Oligopogon hybotinus, $ genitalia ; 69, Oligopogon hybotinus, wing.

ASILIDAE OF THE CONGO BASIN 295

and this was the only Palaearctic record known to Engel (1929 : 372) ; the BMNH
has one female of hybotinus from Turkey. Efflatoun (1937 : 290) described a dis-
tinct species nitidus from the Red Sea coast (Gebel Elba), which forms a link be-
tween the Palaearctic hybotinus and the other species of Oligopogon in the Ethiopian
Region.

In 1858 Loew extended Oligopogon to the Ethiopian Region by describing 0.
penicillatus from Caffraria. Engel (1932) added 0. pollinosus from Rhodesia, and
Engel & Cuthbertson (1937) described 0. nigripennis from the Vumba Mountains of
S. Rhodesia. 0. nigripennis, with its dark wings and rather robust appearance
sounds as if it might, in fact, be a Rhipidocephala, but I have seen no specimen of it,
and include it in the key from the description, for the sake of completeness.

Finally, Rhipidocephala hyalipennis Oldroyd (1959), from Madagascar, is properly
an Oligopogon. It seems, therefore, that Oligopogon extends from the eastern
Mediterranean down to the Cape, and it is not surprising that the present collections
should include a new species from the Katanga, as well as a series of specimens from
Urundi that I am unable to separate from the Palaearctic hybotinus Loew.

The following key includes all the species of Oligopogon at present known to me,
but these small, obscure flies easily escape notice, and it is very likely that other
species will subsequently be discovered.

KEY TO THE SPECIES OF OLIGOPOGON IN THE ETHIOPIAN REGION

1 Wings brown ............. 2

- Wings clear ............. 3

2 Mesonotum with grey anteriorly ; posterior mesonotum and scutellum shining black

nigripennis Engel & Cuthbertson, 1937 : X4

- Mesonotum with yellow or white tomentum, and 3 shining black longitudinal stripes;

scutellum shining black ....... nitidus Engel, 1937 : 29°

3 Mesonotum with shining black stripes separated only by narrow bands of brassy

tomentum. Scutellum inflated, entirely shining black except for a narrow
tomented strip at base ; scutellar margin with some weak hairs but no strong
bristles ....... hybotinus Loew, 1847 : 498 (p. 296)

- Mesonotum without distinct longitudinal bare stripes, at least anteriorly, though it

may have bare spots. Scutellum sometimes with strong bristles ... 4

4 Scutellum entirely tomented, with two long, whitish, marginal bristles ... 5

- Scutellum partly shining, inflated, with black bristles or none at all . . 6

5 Frons relatively broader, not so deeply excavated at vertex, ocelli not falling below

level of eyes. Femora entirely yellow or yellow-brown pollinosus Engel, 1932 : 282

- Frons relatively narrower, ocelli falling well below level of eyes. Femora blackish

brown with brown tips . . . hyalipennis Oldroyd (Madagascar), 1959 : 277

6 Femora and antennae yellow with distinct dark band . . 6* harlequini sp. n. (p. 296)

- Femora and antennae not distinctly banded, though sometimes dark with base and

tip narrowly pale ............ 7

7 Femora shining black, only extreme base and tip red, contrasting strongly with

orange hind tibiae. Hairs of antennal style very long, as long as third segment

superciliatus sp. n. (p. 297)
Femora yellow or brown, but not contrasting strongly with tibiae. Hairs of antennal

style not as long as third segment ......... 8

8 Abdominal segments with basal bands of grey tomentum continuous. Legs bright

orange-yellow . . . . . . . . ? harlequini sp. n. (p. 296)

296 H. OLDROYD

- Abdominal segments with basal bands of tomentum broadly interrupted in middle.

Legs brown ........ penicillatus Loew, 1857 : 282

Oligopogon harlequini sp. n.

Distinguished from all the other species of Oligopogon by the conspicuous orange
and black bands of the antennae and legs.

o" Head. Eyes in dried specimens often rusty brown with distinct and symmetrical black
spots. Face and frons almost parallel-sided, about two-thirds as broad as one eye ; black-
brown, covered with dense brassy tomentum. Ocellar tubercle large, also covered with brassy
tomentum, but with one pair of long, black, ocellar bristles, and several smaller pairs. Frons
with fine, inconspicuous short brown hairs ; moustache covering lower half of face, but rather
sparse, with a few stronger bristles, and a number of fine bristles, all yellow. First two antennal
segments black, with brassy tomentum and yellow hairs ; third segment orange on basal two-
thirds, black on apical third ; first segments of style yellow, bare ; second segment of style
black with black hairs. Proboscis and palpi black with yellow hairs. Occipital bristles in a
single row, longer and black dorsally, shorter and yellow ventrally. No distinct beard.

Thorax densely covered with brassy tomentum, with an unusual arrangement of bare, shining
black spots, all in posterior half of mesonotum : a short median stripe, flanked by crescentic
spots extending into supra-alar area, and a small spot on each side just in front of suture ;
postalar calli partly bare ; two small spots of brown tomentum anteriorly ; scutellum bare
posteriorly, tomented anteriorly. Mesonotum clothed with short, erect, fine hairs, black
anteriorly and medially, yellow posteriorly and laterally ; two strong, yellow notopleural
bristles, otherwise no strong bristles, even on scutellum. Pleura, including postscutellum,
entirely covered with brassy yellow tomentum, with fine yellow hairs on mesonotum, and
bristly yellow hairs in mesopleural fringe.

Abdomen mostly shining. Dorsally each segment brown anteriorly, black posteriorly, the
proportion of black greatest on basal segments, and least towards tip of abdomen. Third to
sixth segments also with a narrow basal band of white tomentum, which extends along extreme
lateral margin. Hairs fine, yellow, semi-recumbent. Venter mostly light brown, with thin
white tomentum, and each segment with a pair of large, bare spots. Male terminalia black or
brown, with yellow hairs.

Legs yellow-brown, with distinct black bands : each femur has a broad preapical band, and
each tibia a prominent black tip.

Wings clear, hyaline, no grey patches. Halteres orange.

Length of body 5 mm ; of wing 4 mm.

$ Similar, but without dark bands on legs, which are entirely orange in the only female
specimen known to me.

Holotype <$. N. NIGERIA : Udubo, 28.viii.57 (? collector) (SAIMR).

Paratypes. Same data as holotype, 2 $ (SAIMR) ; GOLD COAST : N. Territories,
Yagaba, i <j>, n.viii.i9i4 (Capt. Armitage) (BMNH).

Oligopogon hybotinus Loew
(Text-figs 68, 69)

Oligopogon hybotinus Loew, 1847 : 498.

URUNDI : Bururi, 4^, 6 $, x.iQ48 (F. J. Francois) (MRAC)
GARAMBA NATIONAL PARK : P.N.G., 308, Mt. Ndogo, 2 $, 4 $, 15.111.1950, I/a/i,
i °-, i.v.5o (Demoulin) ; 3461, Inimvua, i $, i6.v.i952 (IPNC).

ASILIDAE OF THE CONGO BASIN 297

0. hybotinus was described from the island of Rhodes in the Eastern Mediterranean,
and has hitherto not been known from anywhere else. In the BMNH is a female
specimen from TURKEY : Antalya, Kaldivar nr Gasipazo, c. 750', 2O-22.vii.i963
(E. James). I have no doubt that the specimen from Turkey is correctly identified
as hybotinus, but I am not satisfied that the examples from Urundi are really con-
specific. It is not possible to decide finally until male specimens of hybotinus from
the Palaearctic Region are available to me.

Oligopogon superciliatus sp. n.

Somewhat resembling 0. penicillatus Loew, but distinguished by the excessively
long rays of the antennal style, which are as long as the entire third antennal seg-
ment, and by the strong contrast between the orange tibiae and tarsi and the black
femora, orange only at base and tip.

$ Head. Frons and face covered with dense yellow tomentum, rather long, and divided by a
vertical median line on the frons. Two long, strong, black ocellar bristles, other hairs of frons
weak and yellow. Moustache of sparse but strong black or blackish bristles. Antennae
notable for the excessive length of the cilia of the style : all segments brownish, covered with
yellowish tomentum, basal segments with black hairs. Proboscis and palpi black with yellow
hairs. Occipital hairs in a single row : black, longer and proclinate dorsally ; white, shorter
and straighter ventrally.

Thorax entirely covered with yellowish grey tomentum, in which there are bare, shining black
spots of irregular extent in posterior half of mesonotum. Postalar calli brownish, bare.
Scutellum with white tomentum anteriorly, shining black posteriorly ; hairs fine, erect, black,
no strong marginals. Pleura black-brown in ground colour, entirely covered with yellowish
grey tomentum ; metapleuron with a vertical row of long, yellowish bristles, and a few long
yellow hairs on sternopleuron, otherwise pleura without hairs.

Abdomen dorsally shining black, each segment with an anterior band of white tomentum,
broken in middle into two spots, which are extended laterally as narrow white margins ; hairs
of dorsum short, white on tomented areas, black elsewhere, longer and paler at sides. Venter
black-brown with thin white tomentum and yellowish anterior margin to each segment.

Legs. Femora black, red at base and apex ; tibiae and tarsi orange, only indistinctly darker
towards tips of tarsi. Many short yellow hairs, but longer hairs and bristles mainly black.

Wings clear, hyaline, veins yellow anteriorly, black posteriorly . Halteres pale yellow.

Length of body 6 mm ; of wing 5 mm.

o* not yet known.

Holotype $. KATANGA : Elisabeth ville, Miss. Agric. (MRAC).
Paratype <j>. Same data as holotype (MRAC).

Tribe ASILINI

This complex tribe of many genera is the most advanced in the family. It is
characterized by the universal closure of the marginal cell, and by the third antennal
segment ending not in a style, but in an arista, sometimes long, but always bare.
The members of the tribe Ommatiini are distinguished from Asilini by having the
antennal arista conspicuously feathered, but it is doubtful how much significance
should be attached to this character (see under tribal heading Ommatiini).

Most Asilinae are elongate flies, with slender abdomen, and a generally dusty,

298 H. OLDROYD

appearance. The genera fall into various groups, of which one of the most distinc-
tive is the Promachus-group.

4

The PROMACHUS-GROUP

Characterized by having three submarginal cells in the wing in place of the usual
two. The significance of this in the evolution of Asilidae has already been discussed
in the general part of this paper. The genera of this group have a rather curious
distribution : Promachus itself occurs in every zoogeographical region, with a
complexity of subgenera ; Apoclea is located in the middle eastern arid belt ;
Philodicus and Alcimus are Indo-ethiopian ; and in central and tropical South
America there is a small complex of bee-like genera centred round Mallophora.
Apart from the distinctly bee-like habitus of the Mallophora subgroup, the principal
distinctions are in the wing-venation. Although these differences seem trivial when
described, they are remarkably constant, and they are reinforced by other differences
in appearance which confirm that the genera of this group — or at least the prinicpal
genera already mentioned — are really distinct.

PHILODICUS Loew

Philodicus Loew, 1848 : 391. Type-species : Asilus javanus Wiedemann, by original designa-
tion.
Teretromyia Bigot, 1859 : 416. Type-species : Teretromyia cothurnata Bigot, by monotypy.

A typically asiline genus, which occurs in the Ethiopian and Oriental Regions,
and in the intervening areas of Iran and Pakistan, which are technically Palaearctic.
The African species were reviewed by Blasdale (1957) with a key, and comparative
figures of male and female genitalia, from which the following species of the Congo
Basin may easily be recognized.

Philodicus alcimoides Blasdale

Philodicus alcimoides Blasdale, 1957 : I37-

GARAMBA NATIONAL PARK : P.N.G., 214, I/b/2, i $, 22.^.1950 ; AKAM, i $,
24.0.1950 ; 529, AKAM, i $, 19. v. 1950 ; I/b/3, i $, 24.^.1950 (G. Demoulin) ;
585, I/a/M, i ?, 7.^.1950 (G. Demoulin) ; 730, AKAM, i $, 28.vii.i95o (IPNC).

Philodicus doris (Curran)

Alcimus doris Curran, 1927 : 18.

Philodicus doris (Curran) Blasdale, 1957 : J44-

GARAMBA NATIONAL PARK : P.N.G., 316, 1/a/i, i <$, i $, 20.iii.i95o ; 465, 1/b/2s,
i $, 26.iv.i95o ; 497, I/a/3, i <$, 8^.1950 ; 483, I/a/i, i ?, 5^.1950 ; 529, AKAM ;
1441, II/db/4, i 3, i ?, 23.iii.i952 ; 1444, II/bd/4, i ?, 23.^.1951 ; 1458, II/fc/5,
8 <$, 10 $, 27.iii.i95i ; 1461, II/fc/i8, i <$, 28.iii.i95i ; 1494, II/fd/i7, i $, 4-iv.
1951 ; 1537, II/gc/7, i $, I4.iv.i95i ; 1672, II/gd/4, 2 ^, i ?, S.V.IQSI ; 1506.
II/gf/io, 2 ?, 6.iv.i95i ; II/gd/4, 2 <$, i ?, 13.^.1951 ; 1798, II/fd/i5, i $, 24-v.

ASILIDAE OF THE CONGO BASIN 299

1951 ; 1803, II/fd/i7, i £, 25^.1951 ; 1824, H/fd/iy, i ?, 28.v.i95i ; 1911, II/fc/6,
1 <J. I3-vi.i95i ; 2910, II/fd/i7, i & i ?, I4.xii.i95i ; 3250, Ndelele, K 120/2, 5 £,
4 ?, 28.111.1952 ; 3298, Ppk, i4/g/7, 8 & 2 9, 4.iv.i952 (IPNC).

ITURI : Akini, N. Aru, i ?, v.1936 (Z)r. Pasteels) ; LOMAMI : Lusuku, i <j>, xii.i93O
(P. Quarre) ; KASAI : Poste II, i $, 23.^.1912 (Dr. Mouchet] ; Elisabethville, 2 3
(Dr. Bequaert) ; Kasai, i $ (Dr. Walker) (MRAC).

Philodicus nigrescens Ricardo

Philodicus nigrescens Ricardo, 1921 : 181 ; Blasdale, 1957 : J45-

GARAMBA NATIONAL PARK : P.N.G., 71, I/o/i, i <£, 28.xii.i949 ; 74, I/b/2, i $,
28.xii.i949 ; 75, I/b/3, 2 ?, 2.xii.i949 ; 146, I/a/2, i $, 2.1.1950 (G. Demoulin) ;
214, I/b/2, 2(J, 2 ?, 22.iii.95o ; 261, I/b/3, x ?. 3°5> Mt. Ndogo, 4^, 2 $, 15.111.1950 ;
327, Akam, i <J, 24.111.1950 ; 483, I/a/i, i <J, s.v.igso (G. Demoulin) ; 529, Akam,
4c?, 2 ?, I9.V.I950 ; 585, I/a/M, 3 $, 7.vi.i95o (G. Demoulin) ; 853, 1/0/3 arcl, 2 $,
i <j>, 29.ix.i95o (G. Demoulin) ; 1426, II/fd/i8, i <j>, 19.111.1951 ; 1458, II/fc/5, i <j>,
27.111.1951 ; 1461, II/fc/i8, i $, 28.111.1951 ; 1561, II/fb/i8, i <j>, i8.iv.i95i (/.
Verschuren) ; 1576, II/fb/4, i $, 19.^.1951 (J. Verschuren) ; 1538, II/hc/4, 2 ?,
20.iv.i95i (/. Verschuren) ; 1947, II/gd/8, i ?, 20.vi.i95i ; 2016, II/gc/6, i <^,
29.vi.i95i ; 2379, II/fd/i7, i $t 2 ?, S.ix.igsi ; 3500, Nagero, i $, 10^.1952 ;
2917, II/gc/i5, i cJ, I7.xii.i95i (IPNC).

MANYEMA : Nyangwe, 9 <$, 5 $, 4^.1918 (/?. Mayne) ; Niambi, i ^, 23.iv.i93i
(G. F. ^ Witte) ; TANGANIKA : i $ (Lemaire) ; KATANGA : i <^, 18^.1925 (G. F. de
Witte) ; UELE : Dunga, i $ (^ Gr^/) ; Bukama, 3 $, xi.ign (Dr. Bequaert} ;
LUALABA : Kabelwe, i $, 24^1.1947 (Dr. M. Po//) (MRAC).

Philodicus furunculus Blasdale

Philodicus furunculus Blasdale, 1957 : J4^-

GARAMBA NATIONAL PARK : P.N.G., 497, I/a/3, i $, 8^.1950 ; 509, Km 17, 2 $,
io.v.1950 ; 594, I/a/i, i $, I2.vi.i95o ; 1494, II/fd/i7, i <J, 4.^.1951 ; 1824,
II/fd/i7, i <?, 28^.1951 ; 1855, II/gc/4, i $, i.vi.igsi ; 2024, II/gd/i4, 2 $, 3o.vi.
1951 ; 3401, II/gc/io, i $, 20.iv.i952 ; 3410, II/gd/4, 2 $, 2^.1952 ; 3449, II/gd/4,
i $, 8^.1952 ; 3476, Aka/2, 2 $, 19^.1952 ; 3488, Inimvue, i <£, i $, 20^.1952 ;
3515, Aka/2, i $, 22^.1952 ; 3583, Garamba/2 (source), i <j>, 6.vi.i952 (IFNC).

KATANGA : Elisabethville, 21 ^, n $ (various collectors) ; Lomami, Kambaye,
3 <$, 6 $, vii.i93O (P. Quarre) ; LULUA: Kapanga & R. Kapelekese, i ^, 4 $, 1932/33
(F. G. Overlaet) (MRAC).

Philodicus temerarius (Walker)

Trupanea temerana Walker, 1851 : 121 ; Blasdale, 1957 : X45-

This species is not represented in the collections from the Pare National du
Garamba, but it is widespread in the Congo Basin.
The specimens in the collections of the Muse"e Royale de 1'Afrique centrale are

300 H. OLDROYD

too numerous to be listed in detail, but there are specimens from the following
localities : Bambesa ; Eala ; Mayidi ; Congo da Lemba ; Stanleyville ; Ituri ;
Coquilhatville ; Uele ; Bambesa ; Tshuapa ; Bokuma ; Mayumbe ; Kisantu ;
Sankuru ; Komi ; Equateur ; Abumombazi ; Leopold ville.

Philodicus swynnertoni Hobby

Philodicus swynnertoni Hobby, 1933 : 109 ; Blasdale, 1957 : J44-

URUNDI : Gihanga, Ruzizi, i $, 5 <j>, xi.igsi ; Terr, de Bubanza, 3 <£, 23.ix.iQ5i
(FJF).

LULUA : Kasai, 2 <$, 4 $, 1918 (Dr. Walker) ; Luashi, i $, xi.ig38 (F. Freyne) ;
LOMAMI : Lusuku, 2 $ (P. Quarre) ; UELE : Aba, 2 <$, 5 <j> (M. Hutereau) ;
LUKUGA : Niemba, i $, i <j>, xi.igiy (Dr. Pong) ; Beni Bendi, i <$, v.igi^ (R.
Mayne) ; Bogo, i <$, 7.111.1912 (A. Pilette) ; SANKURU : Pania, Natumbo, i $,
1947 (V. Lagae) ; ITURI : Akini (A. Aru), 2 $>, ¥.1936 (Dr. Pasteels) (MRAC).

Philodicus cinerascens (Ricardo)

Alcimus cinerascens Ricardo, 1900 : 139.
Philodicus umbripennis Ricardo, 1921 : 184.
Philodicus cinerascens (Ricardo) Blasdale, 1957 : J39-

URUNDI : Terr, de Bubanze, 2 $, 6.iii.i952 ; Terr, de Bururi, 3^, i $, 19^.1952 ;
Rumonge, 7 $, 4 $, 1948-49 (FJF).

TANGANIKA : Mpala, 3 $, vii/viii, 1953 (H. Bomans) ; E. TANGANIKA : Kigoma,
i <J, 3 ?, Ix.igiS (R. Mayne) ; Albertville, 2 <£ (^. Mayne) ; Kasenyi, 2 (J, 2 $ (J^. /.
Bredo) ; URUNDI : Rumonge, 6 <J, 4 $, 7.111.1953 (P. Basilewsky) (MRAC).

PROMACHUS Loew

Trupanea Macquart, 1838 : 91, nee Schrank, 1803. Type-species : Asilus maculatus Fabricius,

by original designation.
Promachus Loew, 1848 : 390. Type-species : Asilus maculatus Fabricius, by designation of

Coquillet, 1910.

A large and complex genus, which occurs in all the zoogeographical regions, and
is easily recognized by the particular shape of the three submarginal cells.

Although distinctive at generic level, Promachus includes a great diversity of
species. Hull (1962) recognizes six subgenera (not five as he says), but the status
of these is uncertain. The subgeneric differences are mostly in the antennal arista
and in the form of the ovipositor : the former is often variable between closely
related species, and the latter appears to be an adaptive character, related to the
site of oviposition, and thus not necessarily indicative of relationships.

The African species have never been revised in their entirety, as yet. Miss
Ricardo (1920) published partial keys, but besides being incomplete they were ill-
constructed and difficult to use. At the present time (1969) Mr. P. Blasdale, who
revised the African species of Philodicus in 1957, has work in progress on the species
of Promachus, and has kindly examined the species of Promachus taken in Garamba
by the Mission H. De Saeger.

ASILIDAE OF THE CONGO BASIN 301

Promachus sp. i

GARAMBA NATIONAL PARK : P.N.G., 1412, II/gd/4, 5 <$, i <j>, 17.111.1951 ; 1441,
II/cb/4, i ?, 23.111.1951 ; 3450, Aka, i <£, 2 $, 14^.1952 ; 1299, II/hc/io, 9 $, 7 ?,
28.11.1951 ; 1314, II/kc/io, 3 c?» 4 ?» 2.111.51 ; 1458, II/fc/5, 3 $, i $, 27.111.1951 ;
1444, II/hc/4, i ?, 23.111.1951 ; 1506, II/gf/io, i ?, 6.1v.i95i (IPNC).

Promachus sp. 2

GARAMBA NATIONAL PARK : P.N.G., 1299, II/hc/io, i <J, 2 $, 28.11.1951 ; 3298,
PpK i4/g/7, i & 4-iv-52 ; 3267, Ndelele, i <J, i ?, 27.111.1952 ; 1412, II/gd/4, 5 d>
5 ?, 17.111.1951 ; (IPNC).

Promachus sp. 3

GARAMBA NATIONAL PARK : P.N.G., 1314, II/ke/io, 3 <$, i £, 2.111.51 ; 3298,
PpK i4/g/7, 2 <J, 4-iv.i952 (IPNC).

This and the previous species are obviously closely related, and the existence of
four indeterminate females suggests that they might be conspecific.

Promachus sp. 5
GARAMBA NATIONAL PARK : P.N.G., 2615, PpK 529, i <£, i6.x.i95i (IPNC).

Promachus sp. 6

GARAMBA NATIONAL PARK : P.N.G., 903, I/a/i, i <$, 26.x.i95o ; 1273, Gongala,
i ?, Oct. 1950 (IPNC).

Promachus ugandiensis Ricardo

GARAMBA NATIONAL PARK : P.N.G., 176, I/a/2, i <$, 23.1.1950 ; 204, I/b/3, 2 9,
8.X.I950 ; 205, I/a/i, i <J, I3.xi.i95o ; 3134, Mabanga, 9'", 2 <J, i ?, 19.11.1952
(IPNC).

THE F^SC/^TC/5-GROUP

This is a striking group of species centred round Pro machus fasciatus (Fabricius),
1775, and recognized by having the first three tergites of the abdomen adorned with
double fringes of white or golden hairs.

The present collection contains four females of this species-group. The males
have distinctive genitalia, and may be identified by using an excellent paper by
Hobby (1936 : 182-199, 231-249, 274-278). Unfortunately the females cannot be
confidently identified, except by association with males, and so the females in this
collection can only be listed, as follows :

GARAMBA NATIONAL PARK : P.N.G., 1527, II/gd/4, J ?» 13-^.1952 ; 1872,
II/bc/8, i ?, 5.vi.i952 ; 2059, II/gc/i3s, i ?, I2.vii.i95i ; 2341, Il/fd/i;, i ?,
3i.viii.5i (IPNC).

302 H. OLDROYD

HELIGMONEURA Bigot
Heligmoneura Bigot, 1858 : 357, 662. Type-species : H. modesta Bigot, monotypic.

Engel (1927 : 135) discussed the definition and relationships of this genus, and
especially its distinction from Neomochthems, for which he presented a table.
Heligmoneura is one of several genera in which the vein ^5 of the wing has a pro-
nounced angle at mid-length ; it has pilose metanotal callosities, and a low facial
hump, which becomes nasiform through the narrowing of the face. In the male the
upper forceps (epandrium) is forked, and often conspicuously so (Text-figs 70-72).
The ovipositor is short and downturned.

Seven species from Africa and Madagascar remain in Heligmoneura after making
allowances for past confusion with other genera such as Neomochtherus and
Neolophonotus. The present collection contains three species, only one of which is
from the P.M. Garamba. The three species are easily separated by the very dis-
tinctive male genitalia (Text-figs 70-72) ; a key to species using other characters,
and applicable also to females, awaits a fuller study.

Heligmoneura laevis Engel

(Text-fig. 70)
Heligmoneura laevis Engel, 1927 : 137.

This was one of the species recognized by Hermann, but not published before his

FIG. 70. Heligmoneura laevis, $ genitalia.

ASILIDAE OF THE CONGO BASIN 303

death ; Engel (1927) attributes the species to ' Hermann in litt.', but it appears that
Engel was the author of the published description, and hence of the name.

The form of the male genitalia (Text-fig. 70) appears distinctive, but small
variations occur, and it is possible that more than one species has male genitalia of
this type. A particular problem is N eomochtherus litoralis Lindner (1955), which
is annotated in the original paper ' Auch nach Oldroyd " not laevis Engel, probably
new ". .' I saw the specimen afterwards made type of litoralis, which was from
Dar-es-Salaam, but it is not now before me. From the description and figure it is
certainly a Heligmoneura, not N eomochtherus, and very close to H. laevis. It
appears to differ from laevis in having more pale hairs and bristles on the thorax,
but clearly this needs to be corroborated by evidence from more specimens.

H. laevis was described from the Waterburg District of the Transvaal, and is
known from Malawi. The present record of the species from the P.N. du
Garamba extends the range considerably, with litoralis in Dar-es-Salaam as a
possible link.

GARAMBA NATIONAL PARK : P.N.G., 1249, H/id/4, i ?, 14.^.1951 ; 1494, II/fd/i7,
7 & 4 ?, 4-iv.i95i ; 1527. II/gd/4, 2 ?, 13.^.1951 ; 1610, II/gd/4, i ?, 25.^.1951
(/. Verschuren) ; 1612, II/ee/14, i ?, 26.iv.i95i (/. Verschureri) ; 2917, II/gc/i5,
i ?, I7.xii.i95i ; 3401, II/gc/io, i <?, 29.^.1952 ; 3449, II/gd/4, i $, 8.^1952 ;
1458, II/fc/5, i <?, 27.iii.i95i ; 1855, II/gc/4, i ?, i.vi.5i ; Nagero, i ?, 2/29.ix.i954
(Nebay) ; 1537, II/gc/7, i <?, 14.^.51 ; 1684, II/gd/4, i cT, 7-V.I95* (IPNC).

Heligmoneura modesta Bigot
(Text-fig. 71)

Heligmoneura modesta Bigot, 1858 : 357.

A West African species, which spreads across into the Congo Basin, and so has a
distribution complementary to that of laevis above.

GARAMBA NATIONAL PARK : P.N.G., 786, ±/o/i, * ?> 25.viii.5o (Demouliri) ;
868, 1/0/2, i °., s.x.igso (Demouliri) ; 944, I/o/i, i <£, g.x.i.ig^o (IPNC).

UELE : Bambesa, ix-x.i933 (/. V. Leroy ; H. J. Bredo), 3 & 4 <j> ; Bambili
(Dr. Rodhain), i <£ ; Tukpwu, i $, ix.i937 (L. Leconte) ; Albertville, i <$, 1.1919
(R. Mayne) ; Libenge, 29.xi.i939, i ^ (Leontovitch) (MRAC).

Heligmoneura rodhaini sp. n.

(Text-fig. 72)

A single male, abundantly distinguished from the preceding two species by the
genitalia (Text-figs 70-72). The upper forceps are less deeply cleft than in those
species, and suggest the even less elaborate male genitalia of H. africana Ricardo,
from Nyasaland. This raises the question of the separation of Heligmoneura from
N eomochtherus, readily apparent between typical species of each, but less certain
when the genitalic differences are smaller.

304

H. OLDROYD

o* Head. Space between eyes broader than in H. modesta (cf. Engel, 1927 : 135), at antennae
greater than length of first antennal segment, but not as great as sum of first plus second seg-

FIG. 71. Heligmoneura modesta, <$ genitalia.

FIG. 72. Heligmoneura rodhaini, <$ genitalia.

ASILIDAE OF THE CONGO BASIN 305

ments ; slightly narrower at vertex, and broader at mouth margin. Tomentum of face and
frons brassy yellow ; frons with black hairs along each eye-margin ; facial hump moderate, of
same shape as usual in Heligmoneura, but not so nasiform ; moustache yellow centrally and
ventrally, black dorsally. Antennae rather elongate, almost as long as height of one eye, al
segments proportionately elongate, arista i£ times as long as third segment. Palpi and
proboscis black with yellow hairs ; beard and lower occipital bristles yellow, upper occipital
bristles black.

Thorax. Pronotum with yellow bristles and yellow hairs. Mesonotum entirely tomented
yellowish grey, with a rather indistinct pattern of brown ; two longitudinal stripes, and three
pairs of lateral spots ; covered with short, bristly black hairs, and with longer, strong black
bristles. Scutellum with short, black hairs and with two strong marginal bristles, which, in
type-specimen, are one yellow and one black. Pleura uniformly tomented, with a few sparse
yellow hairs, and with yellow bristles before halteres.

Abdomen dorsally dull black, with broad yellow hind margins, covered with ashy grey tomen-
tum ; clothed, except of yellow bands, with short, bristly black hairs, longer and stronger
posteriorly on each segment, and stronger still laterally ; strongest lateral hairs and bristles
are yellow. Genitalia (Text-fig. 72) shining black, hairs and bristles mainly yellow or brown.

Legs. Orange, with black trochanters, and small black tips to femora and to hind tibiae.
Hairs and bristles mainly black on hind legs and dorsally on others ; yellow elsewhere.

Wings infuscated at tip and along hind margin as far inwards as fork of M$ +4 ; membrane
also stained yellow.

Length of body 16 mm ; of wing 15 mm.

Holotype <j>. CONGO : Semio, 20.ix.igi3 (Dr. Rodhain) (MRAC).

CONGOMOCHTHERUS gen. n.

Type-species : Congomochtherus lobatus sp. n., by present designation.

The type-species bears a deceptive resemblance to Machimus hirsutus Ricardo,
being generally black and grey, with distinctive reddish bands basally on the tibiae.
M. hirsutus, however, is a true Machimus, with dorsocentral bristles extending
forward of the transverse suture, and ovipositor flattened, bare, with free anal
lamellae. Congomochtherus is distinguished by having the presutural dorsocentrals
weak or absent, and by the spiny anal lamellae of the female (Text-fig. 73).

The male of Congomochtherus has an enlargement of the ninth sternite (hypan-
drium), which is quite different from the sclerotized process of the eighth sternite
which is found in many Machimus (Text-fig. 74). In Congomochtherus the ninth
sternite bears a brush or pencil of stiff black hairs, and in the species lobatus the
sternite is extended posteriorly as a rounded lobe, which is yellow and of mem-
branous appearance.

Congomochtherus is clearly allied to Machimus in facial and general structure ;
the genitalia somewhat resemble those of Neomochtherus, which also has no pre-
sutural dorsocentrals, but the spiny lamellae of the ovipositor have no counterpart
except in genera such as Alcimus and Proctacanthus, which are not close related to
Neomochtherus and Machimus. Vein R$ is distinctly angled in the middle, and the
metanotal lobes bristly.

306 H. OLDROYD

KEY TO SPECIES OF CONGOMOCHTHERUS

i Legs entirely black, except for reddish rings at bases of tibiae only. Ninth sternite
of male with yellow median lobe, as well as black bristles (Text-fig. 74) ; discal
scutellar hairs black ......... lobatus sp. n.

- Legs black except for reddish rings at bases of tibiae and of femora. Ninth sternite
of male large, with black bristles, but with no median lobe. Discal scutellar hairs
white ......... penicillatus Speiser (p. 308).

Congomochtherus lobatus sp. n.

(Text-figs 73-75)

A predominantly black and grey species, the tibiae each with a distinct reddish
ring basally, not extending along the dorsal surface.

o* Head. Eyes close together, at vertex separated by little more than breadth of bases of
antennae. Frons covered with brassy tomentum, which also covers ocellar tubercle, and with
short, black, bristly hairs, including several on ocellar tubercle. Tomentum of face white,
yellowish only on eye-margins ; moustache black above, white in middle and below ; facial
tubercle half as high as face, well-defined. Upper occiput with strong black bristles ; bristles
and hairs of lower occiput and buccae white. Antennae black, with black hairs : third seg-
ment as long as first two together ; arista as long as third segment, with microsegment and
small apical style. Proboscis and palpi black with white hairs.

Thorax. Mesonotum entirely tomented : black, with dull yellowish patches behind humeri,
extending into sublateral stripes ; posteriorly these stripes unite with lateral stripes above
wing-bases. Scutellum dull black, inflated, with a recessed grey rim. Hairs and bristles
black : dorsocentrals fairly strong posteriorly, but becoming weak or absent before reaching
transverse suture : scutellum with two strong marginals. Pronotum with yellowish tomentum,
white hairs and black bristles. Pleura covered with thick, ashy grey tomentum ; mesopleuron
and sternopleuron showing areas of velvety grey, which shift according to the angle of the light.

10

FIG. 73. Congomochtherus lobatus, ? genitalia.

ASILIDAE OF THE CONGO BASIN

307

FIGS 74-75. 74, Congomochtherus lobatus, <$ genitalia. 75, Congomochtherus lobatus, head.

3o8 H. OLDROYD

Hairs white, but some black bristles before halteres and on hypopygium.

Abdomen. Each tergite grey anteriorly, posteriorly and laterally, with a large, quadrate,
dark brown spot. Hairs and bristles black on dark areas, white on grey areas. Venter grey,
with hairs mostly white ; no strong ventral bristles. Male genitalia distinctive (Text-fig. 74) :
ninth sternite has median, ovate, lobe, yellow, and lightly sclerotized ; immediately anterior
to this lobe is a cluster of stiff, backwardly-facing, black bristles.

Legs. Black, except for a narrow, reddish or orange ring at base of each tibia. Clothing
hairs white, bristles white and black.

Wings almost uniformly greyish, with microtrichiae in all cells ; marginal and first sub-
marginal cells distinctly wrinkled. Halteres yellowish.

Length of body 14 mm ; of wing 10 mm.

$ Closely similar ; ovipositor as in Text-fig. 73 ; eighth sternite extending backwards as a
firm support beneath very spiny anal lamellae (tenth tergite).

Holotype^. N. NIGERIA : i.v.igia (J. W. S. McFie) (BMNH).

Paratypes. CONGO : Kalembelembe — Baraka, 2 <£, 5 °., vii.igiS (R. Mayne) ;
Manyama, i <$ (R. Mayne) ; UELE : Aba, i <$, 20.iv.igi4 (Dr. Rodhain) ; Kapiri,
i $, X.IQI2 (Miss. Agric.) ; TANGANIKA : Mpata, 780 m, i g, vii-viii.i953 (H.
Bomans) ; KATANGA : Elisabethville, i <$, 11.1929 (Dr. Bequaert) (MRAC).

Congomochtherus penicillatus (Speiser)5

Machimus penicillatus Speiser, 1910 : 100.

Distinguished from lobatus by the male genitalia and the leg-colouring. Ninth
sternite well developed, and with a tuft of black bristles (as in Text-fig. 74), but
lacking the yellow median lobe. Legs black with reddish rings at bases of tibiae,
and at bases of femora as well.

This species bears an extraordinary resemblance to Machimus hirsutus (Ricardo),
the scanty and ill-preserved type-material of which was collected on the Mara
River in Kenya by Capt. A. O. Luckman ; yet hirsutus has neither the spiny
lamellae of the female nor the extended ninth sternite of the male.

Recorded by Speiser from the lowlands surrounding Mem, Ngare na nyuki, and
by Lindner (1955 : 40) from TANGANIKA : Ngaruka. KENYA : Narosswa R.,
6 (£, 2 °., 1912 (W. P. Lowe) ; Masai Reserve, between Guaso Nyeri and Narosswa,
i $, i °-» 23.11.1914 (Capt. A. 0. Luckman) ; Masai Reserve, Ngarenarok, 6000 ft,
i <j>, 3i.xii.i9i3 (Capt. A. 0. Luckman) (BMNH). KiRi-KiRi : Nioka, fin i <£, i <j>,
1913 (Re"gie des Mines). (MRAC).

HOPLOPHEROMERUS Becker

Hoplopheromerus Becker, 1925 : 241.

Tsacas and Oldroyd (1967) revised this genus, which has an interesting distribution
in Africa and in the Far East. They described two new species from the Congo
Basin, and discussed relationships with Heligmoneura and Neomochtherus.

5 While this paper was in the press, Dr. Tsacas has been kind enough to indicate to me certain
specific differences between this species and one of the syntypes of Machimus penicillatus .Speiser which
he has been able to examine. It is possible, therefore that future study of the syntypical series, with
selection of a lectotype, may require a renaming of the species recorded above as penicillatus Speiser.

ASILIDAE OF THE CONGO BASIN 309

MACHIMUS Loew

Machimus Loew, 1849 : i. Type-species : Asilus chrysitis Meigen, 1820, by designation of

Coquillet, 1910.
Tolmerus Loew, 1849 : 94. Type-species : Asilus pyragra Zeller, 1840, by designation of

Coquillet, 1910.
Conosiphon Becker, 1923 : 36. Type-species : Dysmachus pauper Becker, 1907, by original

designation.

Typical Machimus has a distinctive structure, though no item of this is exclusive
to this genus. Even the question whether or not the dorsocentral bristles extend
ahead of the transverse suture is debatable : in typical Machimus there are one or
two strong presutural dorsocentrals, but not as many as in Dysmachus.

The ovipositor of Machimus has the anal lamellae simple and free, not inserted
into the ninth tergite as in Dysmachus, and not spiny as in Congomochtherus. In the
male it is the eighth sternite that is prominent, not the ninth as in Congomocherus,
and this sternite is often drawn out into a conspicuous process (Text-fig. 76) : con-
versely, it may be without process (Text-fig. 77).

Machimus has the metanotal lobes of the thorax hairy or bristly, and vein R$
of the wing has a more or less conspicuous kink. These two characters separate
Machimus clearly from the Neolophonotus-group of genera, as well as from
Ommatius.

The species fall fairly easily into two groups : Machimus (sensu stricto) with a
ventral process in the males, and with four or more scutellar bristles ; and those
species that have no ventral process, and only a single pair of scutellar bristles.
The latter may conveniently be referred to the subgenus Tolmerus, though a formal
separation of the two subgenera is difficult.

The Machimus of the Ethiopian Region are mainly located in eastern tropical
Africa, perhaps as an offshoot from the Palaearctic Region.

KEY TO THE SPECIES OF MACHIMUS IN THE ETHIOPIAN REGION

Scutellum with 4-8 bristles .......... 2

Scutellum with only 2 bristles .......... 5

Tarsi distinctly reddish yellow ...... gytnnus Oldroyd, 1939 : 40

Tarsi black ............. 3

Thorax excessively bristly : long, strong dorsocentrals extend to anterior margin of

thorax ; four or five strong notopleurals .... comans Oldroyd, 1940 : 158

Thorax not excessively bristly ; dorsocentrals extending only a little ahead of
transverse suture, then becoming weaker. Only two strong notopleurals, with one
or two finer bristles ........... 4

Short hairs of scutellum black. Male with long central process (Text-fig. 76)

(? caudiculatus Speiser) ugandiensis Ricardo, 1919 : 56 (p. 310)

Short hairs of scutellum white (? pammelas Speiser, 1910 : 101) juxta Oldroyd, 1939 : 41
Legs entirely black, or with only a faint trace of red at bases of tibiae

nigripes (Ricardo), 1922 : 62 (p. 310)
Tibiae extensively red dorsally .......... 6

Tarsi reddish yellow ; tibiae black towards tip . . . rubripes Ricardo, 1920 : 63
Tarsi black : tibiae reddish only on dorsal surface . . hirsutus Ricardo, 1920 : 62

310 H. OLDROYD

Machimus ugandiensis Ricardo

(Text-fig. 76)
Machimus ugandiensis Ricardo, 1919 : 56.

ITURI : Beni a Lesse, 4^, i $, fin vi.ign (Dr. Martini) ; Beni, i $ (Lt. Bonnevie) ;
Beni, 2 $, x.1928 (A. Collart) ; Bunia, i 9. vi.i928 (P. Lefevre). URUNDI ; Bururi,
1800-2000 m, 23 <£, 17 <j>, 5/12.111.1953 (P. Basilewsky) ; RUANDA : Gitarama,
1850 m, terr, Nyanza, i <j>, 1.1953 (P. Basilewsky) ; Kisanyi, 2 $, xi.igsi (A. E.
Bertrand) ; Lac Mohasi, i <$, iv.i934 (H. Hegh). Kivu : Kisengni, i $ -f 2 spec.,
1953 (.R. Fan Saceghem) ; Kashusha, i <£, i $, 1937 (Vandelannoite) ; Ngoma, i $,
3.x. 1932 (L. Burgeon) ; Lulega, i ^, 8.xi.i925 (Df. /Z". Schouteden) ; USUMBURA :
i $ Bugarama, (Dr. Henrard) ; Rutschuru, i $t i $, 1.1928 (CA. Seydel) ; Bukima,
i $?, iv.i948 (/. F. Leroy) ; Escarpment Kabasha, Chambi, i $, x.1933 (Dy. ^e Wulf)
(MRAC).

URUNDI : Bururi, alt. 1950 m, 20^, 14 $, vi.-x.ig48 ; Bubanza, colline Kagunuzi
(Imbo), alt. 900 m, i #, i $, 2i.vi.ig55 ; Kitega, i $, iii.ig57 (FJF).

Machimus nigripes (Ricardo)
(Text-fig. 77)

Tolmerus nigripes Ricardo, 1922 : 62.

GARAMBA NATIONAL PARK : P.N.G., 1458, II/fc/5, 4 ^, 2 $, 27.iii.ig5i ; I4g4,
II/fd/i7, 5 <J, 8 $, 4.^.1951 ; 1506, Il/gf/io, i $, 6.iv.i95i ; 1525, H/gf/4, i $.

FIG. 76. Machimus ugandiensis, $ genitalia.

ASILIDAE OF THE CONGO BASIN 311

io.iv.i95i ; 1576, II/fb/4, i 9, ig.iv.igsi ; 3450, Aka, 2 9, 14^.1952 ; 3488,
Inimvua, i $, 20^.1952 ; 1671, II/fd/i7, i o, S.v.igso ; 3476, Aka/2, i 9» ig.v.
1952 ; 3480, Inimvua, i $, 16^.1952 ; 3583, Garamba/2 (source), i <£, 6.vi.i952 ;
3656, PPSK, 5/3, i 9, 20.vi.52 (IPNC).

KATANGA : Elisabethville, Kifumashi, sur herbes vivantes, 3 £, 2 9, 9-xii-55 (M.
L^s) ; Elisabethville (piege Harris), i <$, i 9> xi-1935 (P. Quarre] ; Elisabethville,
Lumbumbashi, 2 <J, I $, 11.12.1928 (Z)r. M. Bequaert] ; Elisabethville, i <£, 1953
(//. Bomans) ; Elisabethville, i 9 (Miss. Agric.) ; Lubudi, i 9> viii-ix.i936 (M.
Prinz) ; LUALABA : Kabada (Mutuka), i 9, xii.i953 (R. P. Th. de Caters] ; Kolo-
Kwilu-Madiata, i 9» ix.i9i3 (R. Verschuren] ; Escarpment Kabash-Chiambi, i 9»
x.1933 (Dr. de Wulf] ; LOMAMI : Kmaina, i 9, 1930 (R. M assart] ; UELE : Faradje,
i 9> 14^.1914 (Dr. Rodhain] ; Bondo, Yakoma, i <^, ix.i9i4 (Dr. Rodhain] ;
Bambesa, i 9> 25.viii.i933 (/. V. Leroy] ; La Kando, i <$, Nov. 1925 (Ch. Seydel]
(MRAC).

ASTOCHIA Becker

Astochia Becker, 1913 : 538. Type-species : Astochia metatarsata Becker, monotypic.

Typical specimens of Astochia are distinctive. The male genitalia are character-
istically pendulant, with the upper forceps rather widely set apart from the lower
forceps and claspers (Text -fig. 78). In the female the telescopic ovipositor incor-
porates the seventh to ninth segments, as in Neoitamus. Typically, in both sexes,
the anterior basitarsus is swollen and very bristly.

Astochia closely resembles Neoitamus which, typically, can be easily separated
from Astochia by the proclinate postoccipital bristles, the different male terminalia,
and the normally developed basitarsi. These differences are shown clearly by the

FIG. 77. Machimus nigripes, $ genitalia.

3I2

H. OLDROYD

type-species of the two genera, Astochia metatarsata Becker of the Oriental Region,
and Neoitamus cyanurus of the Palaearctic. The species of the Ethiopian Region
do not fall so easily into the two genera, and the following assignments are provisional.

Astochia armata (Becker)

Neoitamus armatus Becker, 1910 : 22. Originally described from Voi, in KENYA.

GARAMBA NATIONAL PARK : P.N.G., 1461, II/fc/i8, 2 <j>, 28.iii.i95i ; 415, I/a/i,
i $, 20.iii.i95o (IPNC).

Astochia strachani sp. n.

(Text-fig. 78)

Distinguished by the leg-pattern, and by the characteristic male genitalia (Text-
fig. 78). All femora are black on anterior face and dorsally, with a red ring basally,
which is absent in related species neavensis Ricardo and armatus Becker ; posterior
and ventral faces of femora reddish yellow.

cJ Head. Black, obscured by thick tomentum. Tomentum of frons brassy, including ocellar
tubercle ; 2 long ocellar bristles and 2-4 small ones, as well as vertical row along each eye-
margin, all black. Face with more brownish white tomentum ; facial tubercle low but long,
ending abruptly about as far below antennae as length of first segment. Moustache black
above, but mostly white ; no other hairs on face. Occipital bristles very strong, black, one
or two slightly proclinate at tip ; occipital hairs and beard white. Antennae black, with black
hairs. Palpi and proboscis black, with white hairs.

FIG. 78. Astochia strachani, $ genitalia.

ASILIDAE OF THE CONGO BASIN 313

Thorax. Mesonotum entirely tomented : brownish grey, with the usual partially divided
median stripes, reaching back to scutellum, and lateral stripes interrupted at transverse suture ;
clothed with short, bristly black hairs, and strong, black bristles. Scutellum entirely grey,
with white hairs, and two marginal black bristles. Pronotum brownish grey, with rather weak
bristles, some of which are black, and with white hairs. Pleura grey, a little more brownish
dorsally : bristles black in front of halteres, mostly yellowish elsewhere.

Abdomen. Dorsum grey, each segment with a median brown spot and a pair of lateral ones
which become larger posteriorly. Clothing hairs mostly black and bristly, white ones laterally ;
each segment laterally with one or more strong white discal bristles, and on first tergite with a
tuft of white hairs and a row or cluster of bristles, usually black. Venter brownish grey, with
whitish hairs and bristles.

Legs. Coxae like pleura, with yellow or whitish hairs and bristles. Femora reddish yellow,
posteriorly and ventrally, black anteriorly and dorsally, leaving a narrow reddish ring at base.
Fore and middle tibiae and basitarsi reddish yellow, their tarsi otherwise dusky ; hind tibiae
dull reddish, more dusky in apical half, all hind tarsal segments dusky. Close, velvety, bright
yellow fringe on interior surface of fore tibia extends on to two basal segments of tarsus.

Wings. Narrow, pointed. Marginal cell ($} broadened and ridged ; vein R^+5 sinous.
Infuscation of cells back to fifth posterior, and a little in discal cell. Halteres yellowish.

$ Closely similar, but marginal cell of wing not ridged.

Holotype $. NIGERIA : Lagos (G. Strachan) (BMNH).

Paratypes. Same data as holotype, 3 ^, 2 $ (BMNH) ; SIERRA LEONE : Nzala,
i $, 23.111.62 (M. F. Rushton) ; COSTERMANSVILLE : Kasongo, i $, 3 ?, viii-ix.igsg
(Dr. J. Claessens) ; Kapiti, 2^, i <j>, Iv.i9i2 (Miss. Agric.) ; LOMAMI : Luputa, i<$,
ix.i935 (Dr. Bomans) ; KATANGA : Elisabethville, route Sakania, i #, 2i.viii.iQ52
(L. Remy) (MRAC).

~NEOITAMUS Osten-Sacken

Itamus Loew, 1849 : 84 [Praeocc. Coleoptera]. Type-species : Asilus cyanurus Loew, 1849,

by designation of Coquillet, 1910.
Neoitamus Osten-Sacken, 1878 : 82, 235.

Comments have already been made, under Astochia, about the difficulty of
separating the two genera Astochia and Neoitamus. The following species seems
more likely than the others to be properly placed in Neoitamus.

Neoitamus africanus Ricardo

Neoitamus africanus Ricardo, 1919 : 73. Originally described from Mt Kenya.

STANLEYVILLE : Mahagi-Niarembe, i $, x.iQ35 (Ch. Scops) ; Kivu : Kisenyi,
i$, 11.1928 (Ch.Seydel) ; Muturak i $, 3.11.1922 (vanSacaghem) ; RUANDA : Kisenyi,
1800 m, i <£, i8.xi.ig6i (A. E. Bertrand) (MRAC).

Tribe OMMATIINI

Members of this tribe are characterized by the plumed antennae, which have a
ventral fringe of long hairs on the arista (Text-figs 79-81). It might be doubted
whether this single character is sufficient to define a tribe, but the few genera com-
prised in Ommatiini are distinctive even when the antennae are broken. Hull

314 H. OLDROYD

(1962 : 433) regards the chitinized postmetacoxal arch as a tribal character. These
genera have much in common with certain genera of Asilini, notably N ' eomochtherus
and Heligmoneura, but recognition of Ommatiini as a tribe is convenient.

In the genus Ommatius Wiedemann the antennae are similar to those of Asilini,
with short, seedlike third segment, but with the addition of a ventral fringe on the
arista. Cophinopoda Hull, 1958 segregates a small group of species that are distri-
buted from Madagascar eastwards to China and Queensland. Michotamia Macquart,
1838 (Allocotosia Schiner, 1866) comprises a number of Oriental and Australian
species which are characteristically either yellow or purple in colour, and in which
the third antennal segment itself is elongate, with correspondingly shorter, fringed
arista (Text-fig. 80). A single species from the Congo, described in the present
paper, seemed at first to be the only African representative of Michotamia, but it
later proved to be a new genus Thallosia, differing from Michotamia in the genitalia
(which resemble those of Cophinopoda), and in having antennae that are intermediate
between those of Michotamia and those of Ommatius (Text-fig. 79).

The genus Ommatius is one of the most difficult genera of Asilidae, in spite of the
complex genitalia of many of the species. About fifty species occur in Africa south
of the Sahara, and what is known of their distribution suggests that some species
at least may occur over a wide area. For these reasons the present report of this
genus has been confined to the species of Ommatius that actually occur within the
Pare National du Garamba, leaving the larger and more diversified collection of
Ommatius from the rest of the Congo Basin to form part of a projected revision of the
African species of Ommatius, to be made at a later date.

80

FIGS 79-81. Tribe Ommatiini, antennae. 79, Thallosia ; 80, Michotamia

81, Ommatius.

ASILIDAE OF THE CONGO BASIN 315

THALLOSIA gen. n.

Type-species : Thallosia congoicola sp. n., by present designation.

Differs from the varied assortment of species of Ommatius in the head structure as a whole.
Antennae as in Text-fig. 79, with the third segment and arista about equal in length to each
other, and to the first two segments combined. Face with a distinct but small tubercle, occupy-
ing less than half of face, and bearing a sparse moustache of only a few strong bristles (Text-fig.
82). Occipital bristles strong, but few in number ; beard unusually sparse. Proboscis long,
spade-like (i.e. dorsoventrally flattened) and arising from a stout base, which bears a small
clump of strong bristles ventrally. Palpi cylindrical, with fine hairs.

Thorax with a strongly developed pronotum, with distinct 'collar'. Scutellum rather small,
with a deep transverse furrow on its disc, and another immediately before scutellar suture.
Male genitalia of the unique species remarkably like those of Cophinopoda, distinguished by the
long, curved appendage to the upper forceps (epandrium) (Text-fig. 83).

Legs slender, without special features ; pulvilli square at tip as in Michotamia. Wings
rather broadly rounded at tip. Vein R$ ending at, or shortly behind, wing-tip.

Thallosia congoicola sp. n.

(Text-figs 82, 83)

A mainly yellow species with distinctly patterned thorax. Male genitalia as in
Text-fig. 83.

cj Head black in ground colour, vfrons with dark brown tomentum, including ocellar tubercle,
but a small yellow patch just in front of this tubercle. Two black postocellar bristles, and a
very few small frontal bristles, black. Face entirely covered with tomentum, which is whitish,
a little brownish beneath antennae. Moustache confined to facial tubercle, consisting of only
a few white hairs and bristles, and one or two black bristles dorsally. Extensive clypeal region
(Text-fig. 82) bare, shining brown. Palpi cylindrical, shining black-brown, with yellowish
hairs. Proboscis shining black, with a conspicuous clump of brown hairs ventrally near base.
Antennae entirely yellow-brown with black bristles. Upper occiput with strong, moderately
long, yellow bristles ; lower occiput with fine yellowish hairs, beard pale, unusually sparse.

Thorax entirely tomented. Mesonotum golden brown with a distinct pattern of darker brown,
consisting of two widely separated longitudinal stripes, which end in middle of scutum and are
succeeded by a single median stripe, and flanked by three well defined lateral spots. Scutellum
yellowish brown, with sparse, stout yellow hairs but no marginal bristles. A few very strong
black thoracic bristles : 2 notopleural, i supra-alar, i postalar, and one dorsocentral well ahead
of scutellar suture ; only a few, sparse very fine black hairs aligned as dorsocentrals and acro-
stichals. Pleura covered with yellow tomentum, but bare of hairs except for a strong ptero-
pleural bristle and weak bristles ahead of halteres.

Abdomen dorsally reddish brown, without pattern, but more yellow basally, especially on
second segment. Hairs black dorsally, yellow laterally ; hind margins of segments with longer,
slender bristles, and laterally with at least one, quite strong, yellow bristle. Venter reddish
yellow, with yellowish hairs. Male genitalia as in Text-fig. 83 ; upper forceps with long,
curved process, recalling that found in Cophinopoda (Oldroyd, 1964).

Legs slender, entirely reddish yellow, except for slight darkening of last tarsal segment.
Hind femora ventrally with a row of six short, black bristles.

Wings slightly and uniformly smoky. Marginal cell with a very short stalk ; stalk of fourth
posterior cell long. Halteres reddish yellow.

Length of body 8 mm ; of wing 8 mm.

H. OLDROYD

FIG. 82. Thallosia congoicola, head.

FIG. 83. Thallosia congoicola, $ genitalia.

ASILIDAE OF THE CONGO BASIN 317

Holotype <$. GARAMBA NATIONAL PARK : P.N.G., 54, I/o/i, IO.V.IQSO (IPNC).
Paratype $. UELE : van Kerkovenville (Degree/) (MR AC).

OMMATIUS Wiedemann

Ommatius Wiedemann, 1821 : 213. Type-species : Asilus marginellus Fabricius, 1781, by

designation of Coquillet, 1910.
Ommatinus Becker, 1925 : 84. Type-species : Ommatius pinguis Wulp, 1872, by original

designation.

This genus originally contained only three species, marginellus Fabricius, auratus
Fabricius and fulvidus Wiedemann. The two last have been removed to other
genera, auratus to Michotamia and fulvidus to Cophinopoda. The type-species,
marginellus, is one of a great range of tropical species of Ommatius which can only
be resolved by a more detailed study. The following are merely the species taken
by the Mission H. De Saeger in the P. N. du Garamba.

Ommatius vittatus Outran

(Text-figs 84, 85)
Ommatius vittatus Curran, 1927 : 13.

There is a complex of species related to 0. variabilis Engel, and notable for the
swollen hind femora of the males (Text-fig. 84). The exact number of species
involved is obscure, but the present material clearly belongs to vittatus Curran, and
should be known by that name until a study of the whole genus is possible.

GARAMBA NATIONAL PARK : P.N.G., 261, I/b/3, i $, i.iii.igso ; 469, I/a/i, i $,
i.v.iQSO (G. Demoulin) ; 457, 1/3/3, * ?, S.v.igso ; 585, I/a/M, i <£, 2 ?, y.xi.igso
(G. Demoulin) ; 789, Napukumweli, i $, 26.viii.i95o (G. Demoulin) ; 998, I/I/d,
i $, 2i.xii.i95o (/. Verschuren) ; 1458, II/fc/5, i <$, 27.^.1951 ; 1525, II/gf/4, i ?,
io.iv.i95i ; 1527, II/gd/4, i <$, 2 ?, 13.^.1951 ; 1576, II/CH/4, 2 <$, i ?, 19.^.1951
(/. Verschuren) ; 1588, II/hc/4, i $, 2O.iv.i95i (/. Verschuren) ; 1645, II/gc/ii,
i & 4-V.I95I ; I798. H/fd/15. 2 £, i ?, 24.^1951 ; 1824, II/fd/i7, i ?, 28^.1951 ;
1872, II/hc/8, i £, 5-vi.i95i ; 1886, II/gc/6, i & i ?, 8.vi.i95i ; 1920, II/gd/8, i ?,
i6.vi.i95i ; 2015, II/gc/6, 3 <$, 4 ?, 29^1.1951 ; 2024, II/gd/i4, i $, 30^.1951 ;
2056, II/fd/i7, i ?, 9.vii.i95i ; 2072, II/fd/6, i ?, 13.^.1951 ; 2158, II/gc/8, 4 ?,
27.vii.i95i ; 2225, II/fd/i5, 2 $, 7.viii.i95i ; 2243, II/gc/6, i ^, g.viii.igsi ; 2290,
II/fd/6, i (?, i ?, 23.viii.i95i ; 2361, II/gc/i3s, i £, ix.igsi ; 2395, II/fd/i8, i $,
8.ix.i95i ; 2448, II/gd/7, i ?, 2O.iv.i95i ; 2456, II/fd/i5, 8 ^, 2 ?, 2i.ix.i95i ;
2521, II/gc/ii, 2 c?, 3 ?, 5-X.I95I ; 2575, II/fc/6, i cJ, io.x.1951 ; 2653, II/fc/i8,
i & I2.x.i95i ; 2699, II/fc/6, 4 ?, so.x.igsi ; 2740, II/gd/9, i $, S.xi.igsi ; 2774,
II/fc/i35, i cJ, 2i.xi.i95i ; 2506, II/fc/i8, i $, i ?, 24.xi.i95i ; 2881, II/fc/i4, i ?,
io.xii.i95i ; 2935, II/fd/io, 2 $, 20.xii.i95i ; 2941, II/fc/6, 7 (J, 14 $, 26.xii.i95i ;
3287, II/gc/6, 2^, 9 ?, 5-iv.i952 ; 3399. H/gc/n, 3^, i ?, 29.^.1952 ; 3424, H/fd/7,
i $, 5-V.I952 ; 3429. II/M/iS, i ?, 6.V.I952 ; 3567, II/hd/6, 4 c?, 9 ?. 30^.1952 ;
3623, Iso II/2, i $, i8.iv.i952 ; 3656, PPSK 5/3, i <$, 20.vi.i952 ; 3678, Ndelele/4,
i #, i8.vi.i952 ; 3729, II/fc/7, i ^, 2 ?, 4.vii.i952 ; 3811, Utukura/4, i ?, 22.vii.

H. OLDROYD

FIG. 84. Ommatius vittatus, $ hind femur and tibia.

FIG. 85. Ommatius vittatus, $ hind femur and tibia.

ASILIDAE OF THE CONGO BASIN 319

1952 ; 3844, Mt. Moyo, i <J, 2g.vii.ig52 ; 3963, II/gc/6, i <J, 2i.viii.i952 ; 3964,
II/gd/4, i <?, 22.viii.52 ; 3878, II/gc/io, i $, 4.viii.i952 ; 3884, II/fd/i2, i <J, s.viii.
1952 ; 3940, II/gc/7, i & i ?, i4.vii.52 ; 3952, II/gd/6, 2 $, ig.viii.52 ; 4042,
II/gc/8, i $, 9-ix.i952 ; 4057, II/gc/7, J ?. i6.xi.ig52 (IPNC).

Ommatius macroscelis Bezzi

Ommatius macroscelis Bezzi, 1906 : 292 (Ditterei Eritrei) ; Oldroyd, 1939 : 433 ; Lindner,
1955 : 44-

This is a very distinctive little species, easily recognized by its black colour, with
white tomentum on pleura and dorsally on abdomen, especially in the male. The
most characteristic feature is the row of short, mostly black, spines ventrally on the
hind femora.

GARAMBA NATIONAL PARK : P.N.G., 63, 1/b/3, i $, 2i.xii.i949 ; 308, Mt. Ndogo,

1 & i ?, I5.iii.i95o ; 422, I/a/3, 6 <J, 4 ? 17.^.1950 (G. Demoulin) ; 441, Akam,

2 <?, 2i.iv.ip5o ; 456, I/b/i, i <?, i ?, i2.iv.ig5o ; 467, I/b/i, i <£, i $, 26.iv.ig5o
(G. Demoulin) ; 479, I/o/i, i ^, 4^.1950 (G. Demoulin) ; 483, I/a/i, 2 <J, 2 <j>, 5.v.
I95° (G. Demoulin) ; 529 Akam, i <£, ig.vii.ig5o ; 508, I/o/i, i <j>, 7.ix.i95o (G.
Demoulin) ; 999, II/c, i <J>, 2i.xii.i95o (/. Verschuren) ; 1494, Il/fd/iy, i & 4-iv.
i95i ; 1537. II/gc/7, 2 (?, i4.iv.iQ5i ; 1566, II/gd/4, i^, i7-iv-I95i (/• Verschuren) ;
1576, II/fb/4, (j$, ig.iv.i95i (/. Verschuren) ; 1588, II/fc/4, 3 <?, 2 <j>, 2o.iv.ig5i
(/. Verschuren) ; i5go, II/fc/4, 3 c?> 2 $, 2o.iv.ig5i (/. Verschuren) ; i5go, II/ c/4,
2^, 2 ?, 20.iv.ig5i (/. Verschuren) ; 1613, II/hc/4, i & 24.iv.ig5i (/. Verschuren) ;
1618, II/gd/4, 2 cJ, 2 ?, 25.iv.igsi (/. Verschuren) ; 1645, Il/gc/n/ 3 <J, 2 ?, 4.v.
ig5i ; 1671, II/fd/17, i <J, S.V.IQSI ; 1700, II/gd/8, i ^, i ?, g.v.igsi ; 1772,
II/fc/i7, ii ^, 5 $, 22.iv.igsi ; 1811, II/fb/ii, i ^, i $, 25.v.ig5i (/. Verschuren) ;
1824, II/fd/i7, 2 ?, 28^.1951 ; 1855, II/gc/4, i <J, 3 ?, i.vi.igsi ; 1866, II/hc/8,

1 9, 4.vi.ig5i ; 1867, II/gc/6, i ^, i ?, 4-vi.ig5i ; 1876, II/gd/i4, 2 $, 6.vi.ig5i ;
1886, II/gc/6, i ?, 8.vi.i95i ; 1887, H/gd/7, 2 $, 2 ?, S.vi.igsi ; igig, II/gd/8,

2 ^, 3 ?, i6.vi.igsi ; 2015, II/gc/6, 3 ?, 2g.vi.ig5i (/. Verschuren) ; 2052, II/gd/4,
2 & i ?, s.vii.igsi ; 2055, II/gd/4, * ?. 6.vii.ig5i ; 24g6, II/nd/4, i $, 2.x.ig5i ;
2615, PpK. 52g, 2 <$, i ?, i6.x.ig5i, Il/gc/g, 2 $, 2o.x.5i ; 2860, PpK. go/ii5,
i (J, 3.xii.ig5i ; 2653, II/fc/i8, i (J, I2.x.ig5i ; 2668, II/fd/4, 2 <J, i ?, 24.x.ig5i ;
2680, PpK/55, i (J, 3 ?, 26.x.ig5i ; 26g7, II/fd/i6, i $, i $, 2g.x.ig5i ; 26gg,
II/fc/6, i cJ, 30.x.ig5i ; 2708, II/id/8, 2 <J, 3i.x.ig5i ; 2724, PpK.8o.go., 12 ^,
ii ?, S-xi.igSi ; 2731, II/gd/4, 7-xi.ig5i '> 2757. H/gc/n, 2 <?, I3.xi.ig6i ; 2773,
PpK/55, 2 cJ, ig.xi.ig6i ; 2774, II/gc/i3s, i ^, 2i.x.ig5i ; 2780, II/gd/4, i ?,
23.xi.ig5i ; 2806, II/fc/i8, 8 ^, 10 ?, 24.xi.ig5i ; 2814, II/fd/i6, 2 ^, 28.xi.ig5i ;
2818, II/fc/17, 2 (J, i ?, 2g.xi.igsi ; 2831, II/gd/4, i ?. 30-xi.ig5i ; 2860, PpK,
90/115, i & 3.xii.i95i ; 2881, II/fc/14, i ^, 3 $, io.xii.ig5i ; 2882, II/gc/io, i ?,
ii.xi.ig5i ; 2910, II/fd/i7, 3 <J, 2 $, I4.xii.ig5i ; 2917, II/gc/i5, I <J, 3 $, i7-xii.
i95i ; 2935, II/fd/io, 3 (J, 2 $, 20.xii.i95i ; 2g3g, II/fd/i8, i ?, II/fd/i8 ; 2941,
II/fc/6, i #, i <j>, 26.xu'.ig5i (/. Verschuren} ; 3262, II/fc/i8, i ?, 3i.iii.ig52 ;
3311, PpK 73/d/g, 2 #, 2 ?, 8.iv.52 ; 33gg, Il/gc/n, 2 ^, i ?, 2g.iv.ig52 ; 3410,
II/gd/4, 2 & 2 ?, 2.v.ig62 ; 3424, II/fd/7, i #, 5.v.ig52 ; 342g, II/fc/i8, i ?, 8.v.

320

H. OLDROYD

1952 ; 3461, Inimvua, i ?, i6.v.ig52 ; 3476, Aka/2, 2 $, 19^.1952 ; 3480, Inimvua,
i <?, i6.v.i952 ; 3514, Aka/2, i ?, 22.V.I952 ; 3964, II/fd/4, i $, 3^.1952 ; 3701,
II/gd/i, i $, 24^1.1952 ; 4100, Iso III, i <£, 26.ix.ig52 (IPNC).

Ommatius macquarti Bezzi

(Text-fig. 86)
Ommatius macquarti Bezzi, 1908 : 379.

Easily recognized by the bulbous male genitalia (Text-fig. 86), but extremely
variable in size and general stature.

GARAMBA NATIONAL PARK : P.N.G., 213, 1/2/3, x <$> 20.^.1950 ; 414, I/b/3, i <£,
414, I/b/3, I <$> i4-iv.i95o ; 529, Akam, i $, 19^.1950 ; 1890, II/fd/i7, i $, n.vi.
1951 ; 1916, II/fd/i, !<£, I5.vi.i95i ; 1960, II/fd/i7, i $, 25.vi.i95i (IPNC).

The collection of the Muse"e R. de 1'Afrique centrale contains many examples of
this species, which will be considered later in a generic revision.

FIG. 86. Ommatius macquarti, $ genitalia.

321

distinguished in

ASILIDAE OF THE CONGO BASIN

Ommatius digit tat us sp. n.
(Text-figs 87-89)

A small, delicate species, with slender abdomen and yellow legs
the male by the characteristic genitalia (Text-fig. 87).

$ Head. Eyes rather widely separated. Frons entirely covered with dark chocolate-brown
tomentum, with several long, proclinate, black occipital bristles, and with long black hairs along
eye-margins. Face with dense yellow tomentum, only slightly rounded in profile, without any
distinct tubercle ; moustache with white or yellowish hairs and bristles near mouth-margin,
and continued up to bases of antennae by a double row of black bristles, flanked with slender
black hairs. Antennae black, with some black and some yellow hairs and bristles ; third
segment short, little longer than first. Occipital bristles black and strongly proclinate ; lower
occipital bristles and beard yellowish.

Thorax. Mesonotum entirely tomented, without definite pattern, though anteriorly and
medially it is darker than postero-laterally. Thoracic bristles longer than usual, mostly yellow ;
2 notopleural, i supra-alar, sometimes one or two dorsocentrals ; fine hairs in positions of
dorsocentrals, acrostichals, humeral and lateral areas are also exceptionally long and fine,
longer than total of three antennal segments. Scutellum grey, with long, curved yellow hairs
on disc, and several bristly yellow hairs, but no strong bristles, on margin. Pronotum and
pleura grey, with entirely yellow hairs and bristles ; bristles are present on pronotal collar, and
before halteres, and in a vertical row on posterior margin of mesopleuron.

FIG. 87. Ommatius digittatus, <$ genitalia.

322

H. OLDROYD

Abdomen. Tergites densely covered with velvety tomentum, which alters in tone according
to the direction of the light, and almost conceals dull yellow hind margins of segments. Hairs
entirely golden yellow, longer and more bristly laterally, with a clump of strong yellow bristles
on each side of first segment. Sternites yellowish grey, with a few fine hairs, but with an array
of strong yellow bristles in a characteristic pattern (Text-fig. 89). From seventh sternite
onwards abdomen is darker brown both dorsally and ventrally, and ends in very distinctive
male genitalia (Text-fig. 87).

Legs. Coxae and trochanters black with grey tomentum. Legs otherwise clear yellow down
to tip of basitarsus ; rest of tarsi black. Hairs and bristles long, slender, entirely yellow
except towards tips of tarsi, where short bristles are black.

Wings. Very narrow and elongate ; radial fork bell-mouthed, straddling wing-tip ; fourth
posterior cell with long petioles both basally and apically. Halteres pale yellow knob on a grey
stalk.

Length of body 9 mm ; of wing 5 mm.

$ Generally resembles male, except that sternites lack the pattern of strong bristles ; tergites
with one strong yellow bristle on each side, in posterior angle ; $ genitalia as in Text-fig. 88.

Holotype
(IPNC).

GARAMBA NATIONAL PARK : P.N.G., 4103, Mabanga, 29.ix.i952

FIGS 88-89. 88, Ommatius digittatus, $ genitalia ; 89, Ommatius digittatus
abdominal sternites with bristles (cf. fig. 93).

ASILIDAE OF THE CONGO BASIN

323

Paratypes. P.N.G., 1^,1$, 26.iv.ig5o (G. Demoulin) ; 497, 1 /a/3, i(£, 8.v.ig5o ;
657, I/O/i, i $, 30.111.1950 (G. Demoulin) ; 812, I/o/i, i $, ii.ix.i95o (G. Demoulin) ;
866, 1/0/2, 2 c?, i ?, 3.x. 1950 (G. Demoulin) ; 898, I/o/2, i $, 20. x. 1950 ; 2479,
II/gc/6, i ?, 27.ix.i95i ; 2554, II/fd/i8, i ?, 8.x.i95i ; 2680, PpK/35, i ?, 28.x.
1951 ; 2699, II/fc/6, i & i ?, 30.x.5i ; 3i97> Anie /9. * & i ?, 18.111.1952 ; 3399,
II/gc/ii, i (J, i ?, 2g.iv.i952 ; 3583, Garamba/2 (source), i $, 6.V.I952 ; 3589,
Ndelele/2, i $, 6.vi.ig52 ; 3678, Ndelele/4, 2 $, 2 ?, i8.vi.ig52 ; 3743, II/gd/4,
i $, 5.vii.i952 ; 3763, Il/fd/iy,. i & g.vii.ig52 ; 4101, Pali /8, 2 <?, 2 ?, 27.ix.ig52
(IPNC).

Ommatius garambensis sp. n.

(Text-fig, go)

Apparently one of the vittatus-variabilis group of species, but distinguished from
others by the male genitalia (Text-fig, go), with the square-tipped upper forceps,
and by the colour of the femora, which are black with a dorsal or postero-dorsal
reddish stripe.

(J Head. Frons and face relatively broad, narrowest at vertex, broadest at mouth-margin.
Frons with brassy brown tomentum over a black ground ; ocellar tubercle shining black
dorsally, with short black hairs on each side ; a single pair of black postocellar bristles on each
side ; a single pair of black ocellars. Tomentum of frons paler, more yellowish ; facial tubercle
occupying lower third of face, with a strong moustache of mainly black bristles, with white hairs
and bristles on mouth-margin. Antennae black, with black hairs and bristles ; proboscis and
palpi black with whitish hairs. Upper occipital bristles strong and black, lower occiput and
beard with white hairs.

FIG. 90. Ommatius garambensis, $ genitalia.

324 H. OLDROYD

Thorax. Mesonotum with ashy grey and brown tomentum, giving a moderately distinct
pattern of two longitudinal stripes and two lateral spots on each side. Bristles and hairs
black : 2 notopleurals, i supra-alar, 3-6 dorsocentrals ; fine black hairs along lines of dorso-
centrals and acrostichals, spreading into large lateral areas above wings. Scutellum uniformly
yellowish grey, with pale discal hairs, and two strong black marginal bristles. Pronotum and
pleura with grey tomentum. Pronotum with a collar of black bristles ; one strong black
pteropleural bristle, and a vertical row of black bristles before halteres ; mesonotum unusually
hairy, with long, fine, black hairs.

Abdomen. Tergites with dull yellow posterior margins ; clothed with short, black hairs,
which become yellowish and longer laterally ; no distinct strong bristles except on first tergite.
Sternites similar, hairs pale.

Legs. Coxae black in ground colour, with grey tomentum and white hairs ; middle and hind
coxae each with a single, strong, yellow bristle externally. Femora black, with a dorsal reddish
stripe, which may extend into an apical ring, and on to ventral surface, especially on fore and
middle femora. Clothing hairs of femora black anteriorly on fore and middle legs, white
posteriorly, and on entire hind femora. Fore femora without strong bristles ; middle and hind
femora with very strong black bristles on anterior face, and in a ventral row on hind femora.
Fore and middle tibiae and tarsi reddish yellow, with black tips to segments ; hind legs with
only basal half of tibiae reddish yellow ; rest, including entire tarsi, black. Bristles mixed
black and yellow.

Wings uniformly pale greyish ; marginal cell a little ridged, but not strongly dilated on
costal margin ; radial fork bell-mouthed, vein R^ ending distinctly before wing-tip. Halteres
brown.

Length of body 14 mm ; of wing n mm.

$ Generally similar, but larger areas of reddish colour on legs.

Holotype #. GARAMBA NATIONAL PARK : P.N.G., 3277, PpK, 5i/g/a, 2.iv.52
(IPNC).

Paratypes. Same data as holotype, 6<£, 3 $ ; 808, 1/o/i, 7.ix.ig5o (G. Demoulin) ;
2056, II/fd/i7, i $, 9.vii.i96i ; 2242, II/fd/i7, i $, I3.viii.i95i ; 2341, II/fd/i7,
i $, 3i.viii.i95i ; 2814, II/fd/i6, i $, 28.xi.i95i ; 2839, II/fd/i8, i $, 2i.xii.i95i ;
3262, II/fc/i8, i c?, 3i.iii.i952 (IPNC).

Ommatius caligula sp. n.

(Text-figs 91-93)

Superficially resembling digittatus sp. n., but distinguished in both sexes by the
genitalia, and in the males by the arrangement of strong bristles on sternites 3, 4, 5
of the abdomen (Text-fig. 93). From drusus sp. n., both digittatus and caligula are
distinguished by having the femora and tibiae entirely reddish yellow.

(J Head. Eyes rather widely separated. Frons dark velvety brown, with slender, black
hairs on ocellar tubercle, and along eye-margins. Face gently rounded, without definite
tubercle, densely covered with yellow tomentum. Moustache extensive, covering most of face,
and extending almost to bases of antennae ; rather sparse, composed of a mixture of black and
yellow bristles and hairs. Antennae black, with velvety brown tomentum and black hairs ;
third segment almost as short as second. Palpi and proboscis black with yellow hairs. Bristles
of upper occiput black, fine, strongly proclinate, with fine black hairs medially ; lower occiput
and beard with fine, sparse, white hairs.

Thorax. Black with rather thin, golden brown tomentum. Mesonotum more brown
anteriorly, becoming grey posteriorly and on scutellum, but without definite pattern ; a little
paler brown near humeri. Bristles mostly white, but occasionally black : 2 notopleural, i

ASILIDAE OF THE CONGO BASIN

325

supra-alar, i postalar ; a single pair of dorsocentrals conspicuous in a row of fine dorsocentral
hairs, black anteriorly, yellow posteriorly ; a few fine black acrostichals. Scutellum grey,

FIG. 91. Ommatius caligula, <$ genitalia.

FIGS 92-93. 92, Ommatius caligula, $ genitalia ; 93, Ommatius caligula,
abdominal sternites with bristles (cf. fig. 89).

326

H. OLDROYD

with fine, yellowish discal hairs, and two long, yellow, marginal bristles. Pleura with thin,
golden brown tomentum on dorsal half and whitish tomentum ventrally. Hairs and bristles
mostly yellowish ; a distinct pteropleural bristle, and a vertical row posteriorly on meso-
pleuron, as well as before halteres.

Abdomen dorsally with dense brown tomentum, more greyish anteriorly, more reddish brown
posteriorly, with fine yellow hairs becoming more bristly laterally, but without true bristles on
tergites. Sternites 4, 5 bare, shining, others covered with tomentum. A remarkable arrange-
ment of strong yellow bristles : a single pair on segment 2 and an arrangement on segments
3, 4, 5 as shown in Text-fig. 93.

Male genitalia very long, shining black, shaped as in Text-fig. 91.

Legs. Coxae like pleura, trochanters black. Femora entirely reddish yellow : fore femora
ventrally with a row of about 4 powerful yellow bristles. Tibiae and basitarsi also reddish
yellow, tarsi then becoming progressively darker.

Wings. Uniformly greyish, with a coating of microtrichiae. Halteres reddish brown.

Length of body 7 mm ; of wing 6 mm.

$ closely resembles male, except that ventral bristles on abdomen — evidently a secondary
sexual character — are much less strongly developed. Female genitalia as in Text-fig. 92,
eighth tergite strongly convex and overlapping sternite, giving this species a distinctive
appearance.

Holotype <£. GARAMBA NATIONAL PARK : P.N.G., 3678, Ndelele /4, i8.vi.i952

(IPNC).

Paratypes. Same data as holotype, 2 <$, 7 $ ; 2680, 2 $ ; 3488, PpK/55, 3 9.
20.^1952 (IPNC).

Ommatius canicoxa Speiser
(Text-fig. 94)

Ommatius canicoxa Speiser, 1913 : 142.

GARAMBA NATIONAL PARK : P.N.G., 199 I/a/3, i ?, 7.11.1950 ; 213, I /a/3, i ?,
20.11.1950 ; 395, I/o, i <£ 27.11.1950 ; 528, 529, Akam, i $, 2 $, 19.^1950 ; 832,
I/O/2, I <J, 2 ?, ix.igso ; 3476, Aka/2, i <J, 19^.1952 (IPNC).

FIG. 94. Ommatius canicoxa, <$ genitalia.

ASILIDAE OF THE CONGO BASIN

327

0. canicoxa Speiser was described from the Kamerun, near Duala, and as far as
can be decided from the description alone, it is represented in the BMNH by a small
series from the Cameroons and Nigeria. The material agrees with Speiser's descrip-
tion in the general colouring, and in particular in the wings, which are brownish
antero-apically, and have the costal margin dilated, [die Ausbuchtung des Vorder-
randes, welche vielen Ommatius- A.rten zukommt. . . .]

East African specimens resembling these were recorded by Oldroyd (1939 : 42)
and by Lindner (1955 : 44), with a note of interrogation. Although there is con-
siderable individual variation in chromatic characters as well as in size, it seems
possible to distinguish two species by the dilation or not of the costal margin in the
male, and in both sexes by the extent of the reddish base of the hind femora. Both
species occur in the present collection ; those listed immediately above are the
specimens believed to be true canicoxa, and the others, including Oldroyd's and
Lindner's specimens, are referred to a new species, Ommatius drusus sp. n.

Ommatius drusus sp. n.

(Text-figs 95, 96)

Ommatius sp. near canicoxa Speiser ; Oldroyd, 1939 : 41.
Ommatius aff. canicoxa Speiser ; Lindner, 1955 : 44.

Closely similar to canicoxa Speiser, and with almost identical male genitalia
(Text-figs 94, 95), but distinguished by not having the costal margin of the wing
noticeably dilated, and usually by having the hind femora dimidiate, i.e. divided
in the middle into a reddish yellow basal half, and a black apical half, with oblique
border between the two colours.

FIG. 95. Ommatius drusus, $ genitalia.

3*8

H. OLDROYD

cJ Head. Frons narrower than in canicoxa, vertex less deeply excavated ; tomentum of frons
light golden brown, ocellar tubercle shining black, prominent, and with two very strong black
bristles ; hairs along eye-margins extremely small and inconspicuous ; face high and narrow,
tomentum pale yellowish, moustache white on mouth-margin, extended towards antennae as
two rows of black bristles. Antennae black, with black bristles ; third segment short. Pro-
boscis and palpi black with yellow hairs. Upper occipital bristles black, proclinate ; lower
occipital bristles and scanty beard whitish.

Thorax. Mesonotum rust-brown, more yellowish on shoulders and posteriorly, but without
definite pattern. Strong black bristles : 2 notopleurals, i supra-alar, i postalar, 3 pairs of
weak dorsocentrals. Scutellum with brassy yellow tomentum, weak and indistinct pale discal
hairs, and one pair of black marginal bristles. Pronotum brownish medially, laterally pale
yellowish like pleura ; pronotal collar of weak black bristles ; pleura with fine, pale, yellow
hairs and stout black bristles, including a pteropleural bristle.

Abdomen. Dorsally cinereous, with dull reddish posterior margins to segments. Clothing
hairs black, longer on hind margins, replaced laterally by yellowish hairs and one or more long
marginals. Venter similar, with fine, yellow hairs. Male genitalia as in Text-fig. 95, black,
but sometimes reddish.

Legs. Coxae somewhat yellowish in ground colour, with yellow hairs and bristles. Tro-
chanters reddish. Femora reddish yellow with black markings : fore and middle femora black
anterodorsally ; hind femora black apico-dorsally, with an oblique line of division between
black and reddish areas (extent of colour varies, as well as distinctness of division between the
two areas). Fore and middle tibiae and basitarsi reddish, with black tips ; a dorsal black
stripe on tibiae and segments 2-5 of tarsi black. Hind tibiae reddish on basal half, blackish
apically, tarsi blackish.

Wings. Costal margin not appreciably dilated ; sometimes only dark brown over a small
stigmal area, otherwise mostly pale greyish as a result of microtrichiae which cover all cells
except second basal. Halteres with yellowish stalk and deep red knob.

Length of body 10 mm ; of wing 9 mm.

? Similar to male ; $ genitalia Text-fig. 96.

FIG. 96. Ommatius drusus, $ genitalia.

ASILIDAE OF THE CONGO BASIN 329

Holotype <J. UGANDA : Namwamba Valley, 6500 ft (F. W. Edwards) (BMNH).

Paratypes. Same data as holotype, 3 9 '< UGANDA : Budongo Forest, I 9 ;
Kilembe, 4500 ft, 4 <£, 4 $ ; Mobuka Valley, c. 4000 ft, i <$, 2 9 ; Mbarara, i 9 (all
coll. F. W. Edwards) Kyarumba, 4500' (D. R. Buxton), 3 9 ; Nyamgasani Valley,
6400' (D. R. Buxton), i 9 (BMNH).

GARAMBA NATIONAL PARK : P.N.G., 456, I/b/i, i <$, I2.iv.i95o (G. Demoulin) ;
527, ll)/i, i <?, i ?, 17^.1950 ; 3197, Anie/9, i $, 18.111.1952 ; 3229, BESK, 8/d/g,
i 9, 25.111.1952 ; PpK, 73/d/9, i ?, 8.iv.52 ; 3463, Aka, i 9, 15.^1952 ; 3476,
Aka/2, i (J, 19^.1952 ; II/fd/i7, i <£, 27.viii.52 (IPNC).

Ommatius nongipennis Lindner

Ommatius longipennis Lindner, 1955 : 45.

GARAMBA NATIONAL PARK : P.N.G., 3480, Inimvua, i <J, 16^.1952 ; 3844, Mt.
Moyo, i 9, 29.vii.52 (IPNC).

UELE : Aba, i $, 2o.iv.i9i4 (Dr. Rodhain) (MRAC).

This species is very variable in the colour of the hind femora. It is provisionally
identified with longipennis Lindner, though no information is given in Dr. Lindner's
description about the genitalia. Confirmation of this identification must await a
fuller study of the genus Ommatius in Africa.

Ommatius imperator Oldroyd

Ommatius imperator Oldroyd, 1939 : 45.

UELE : Bambesa, i <J, i 9. io.x.i933 (/. Leroy) (MRAC).

This species was described by me many years ago from specimens collected in
Uganda by Neave and by T. H. E. Jackson. As far as I am aware, this is the first
time that any other specimens have been noted, and so I record them in this paper.

0. imperator is perhaps the most striking species of the genus Ommatius, having
the black body and yellow, black-tipped wings that are common to a whole complex
of aposematic insects of various Orders.

ACKNOWLEDGEMENTS

I am deeply indebted to the late Dr. V. van Straelen for inviting me to study this
most interesting material, and for originally agreeing to publish it, with the addition
of other relevant material from the Congo Basin ; to Dr. P. Basilewsky for
courteously allowing me to incorporate large collections of specimens from the
collection of the Musee Royale de I'Afrique centrale, and to Monsieur F. J. Fran£ois
and the authorities of the Institut R. des Sciences naturelles de Belgique for similar
courtesies in respect of material collected in Urundi by Monsieur Fran£ois.

I am also indebted to Mr. Philip Blasdale for determining the specimens of
Philodicus and Promachus and to Dr. L. Tsacas for advice about specimens of
Neomochtherus, which he will incorporate into his own studies of that genus.

330 H. OLDROYD

REFERENCES

Oldroyd (1963) gives a fairly complete Bibliography of papers on African Asilidae up to that
date, but not including the classic references to Linnaeus, Fabricius, Meigen and Macquart.

BLASDALE, P. 1957. The Asilidae (Diptera) of the genus Philodicus Loew in the Ethiopian

Region. Trans. R. ent. Soc. Land. 109 : 135-148.
BROMLEY, S. W. 1935. New Asilidae from the Belgian Congo (Diptera). Revue Zool. Bot.

afr. 26 : 404-415.
CURRAN, C. H. 1927. Undescribed Asilidae from the Belgian Congo. Am. Mus. Novit.

272 : 1 8 pp.

- 1928. Diptera of the American Museum Congo Expedition II. Bull. Am. Mus. nat.
Hist. 53 : 328-338.

HULL, F. M. 1962. Robber Flies of The World 2 vols. : 908 pp.

JANSSENS, E. 1951. Contribution a 1'etude des dipteres de 1'Urundi. I. Un nou veaugenre
de diptere asilide : Oxynoton n.g. Bull. Inst. r. Sci. nat. Belg. 27 : (54) : 1-4.

— 1952. Revision du genre Lasiocnemus Loew. Bull. Inst. r. Sci. nat. Belg. 28 (24) : i-io.

— 1953. Une nouvelle espece de diptere asilide d'Afrique centrale : Laphria maynei n. sp.
Bull. Annls Soc. r. ent. Belg. 88 : 289-290.

- 1953. Contribution a l'6tude des dipteres de 1'Urundi. IV. Asilidae. Bull. Inst. r.
Sci. nat. Belg. 29 (42) : 15 pp.

!953D. Revision des especes africaines du genre Ammophilomima Enderlein (Diptera :

Asilidae). Bull. Inst. r. Sci. nat. Belg. 29 (12) : 44 pp.

- 1954. Leptogasterinae (Diptera : Asilidae) in Explor. Pare. natn. Upemba Miss. G. F
de Witte (1946-49), 25 : 113-134.

- 1954. Leptogasterinae du Congo Beige. Annls Mus. r. Congo Beige, Sci. Zool. 1 : 400-
401.

1955- Leptogasterinae du Mission P. Basilewsky 1953 en Ruanda-Urundi. Annls Mus.

r. Congo Beige, Sci. Zool. 36 : 303-305.

KARL, E. 1959. Vergleichend-morphologische Untersuchungen der mannlichen Kopulations-

organe bei Asiliden (Diptera). Beitr. Ent. 9 : 619-680.
LINDNER, E. 1955. Ostafrikanische Asiliden (Dipt.). Ergebnisse der Deutschen Zoologischen

Ostafrika-Expedition 1951-1952. Jh. Ver. vaterl. Naturk. Stuttgart 110 : 24-46.
OLDROYD, H. 1939. Asilidae in Ruwenzori Exped. 1934-35 2 : 25-46. London.

1940. The genus Hoplistomerus Macquart (Diptera : Asilidae). Trans. R. ent. Soc.

Lond. 90 : 307-318.

1963. The Tribes and Genera of the African Asilidae (Diptera). Stuttg. Beitr. Naturk.

107 : 16 pp.
— - 1964. Diptera from Nepal : Asilidae (inc. genus Cophinopoda). Bull. Br. Mus. nat.

Hist. (Ent.) 15 : 239-254.

1966. The genus Rhipidocephala (Diptera : Asilidae). Bull. Br. Mus. nat. Hist. (Ent.)

18 : 145-172.

— 1969. The family Leptogastridae (Diptera). Proc. R. ent. Soc. Lond. (B) 38 : 27-31.
TSACAS, L. & OLDROYD, H. 1967. Revision du genre Hoplopheromerus Becker (Dipt., Asilidae).

Annls Soc. ent. Fr. (N.S.) 3 (2) : 257-325.

INDEX

(Synonyms in italics)

abdominalis, Laxenecera, 236, 238 albicincta, Laxenecera, 236, 237, 240

Acurana, 235 albipila, Strobilothrix , 257

acutirostre, Microstylum, 265 alcimoides, Philodicus, 298

aequinoctialis, Ammophilomima, 219 Ammophilomima, 216, 219

africana, Proagonistes, 253 amphora, Xenomyza, 287, 290

africanus, Neoitamus, 313 Ancylorrhynchus, 271

Afromelittodes, 225 Andrenosoma, 226, 251, 255

INDEX

331

angusticornis, Neolaparus, 259

anthracinus, Lasiocnemus, 224

Anypodetus, 225

aphaea, Loewinella, 257

apicalis, Ancylorrhynchus, 272

apicalis, Leptogaster, 218

apicalis, Proagonistes, 252, 254

apiformis, Laxenecera, 240

argenteoviridis, Atomosia, 257

aristalis, Leptogaster, 217

armata, Astochia, 312

Asilini, 297

Astochia, 311

athletes, Proagonistes, 253

Atomosiini 257

attenuatum, Microstylum, 265

aureopilosa, Laphria

auricomata, Laxenecera, 236, 237, 238

auricorpus, Laphria, 227

aurifer, Laphria, 227, 228

auripennis, Ammophilomima, 219

austeni, Proagonistes, 253

basilewskyi, Ammophilomima, 219
basilewskyi, Leptogaster, 218
bella, Laphria, 226
bequaerti, Laphria 226
bicingulata, Leptogaster, 217
bicolor, Euscelidia, 220
bipenicillata, Laphria, 227
bloesus, Microstylum, 266
boranica, Andrenosoma, 256
braunsi, Ancylorrhynchus, 272
braunsi, Microstylum, 264

caffer, Stichopogon, 283
caligula, Ommatius, 324
canicoxa, Ommatius, 326
capensis, Microstylum, 265
capucinum, Microstylum, 265
carbonaria, Laphria, 227, 228
carpenteri, Perasis, 245
caudiculatus, Machimus, 309
Cenochromyia, 226, 235
chapini, Laxenecera, 238, 243
chrysonema, Laxenecera, 236, 242
cinerascens, Philodicus, 300
comans, Machimus, 309
complexa, Andrenosoma, 256
congoensis, Gonioscelis, 277
congoicola, Thallosia, 315
congoiensis, Rhipidocephala, 287
Congomochtherus, 305
Conosiphon, 309

consimilis, Hyperechia, 250
consimilis, Laphria, 228
contristans, Laphria, 227, 228
cribrata, Trichardis, 249
cruciger, Ancylorrhynchus, 271
crux, Ancylorrhynchus, 273, 274
Ctenota, 225
ctenoventris, Laphria, 227, 228

Damalis, 287
Dassylina, 231
Dasyllina, 224, 231
dasypoda, Laxenecera, 238
Dasythrix, 225 243
decoratus, Neolaparus, 259
digittatus, Ommatius, 321
dimidiata, Laxenecera, 236, 240
dispar, Microstylum, 265
Dogonia, 269
Dolichoscius, 216, 223
dorata, Euscelidia, 220
doris, Philodicus, 298
drusus, Ommatius, 327
dymes, Laphria, 231
Dyseris, 235

ealensis, Leptogaster, 217

Eclipsis, 264

Elasmocera, 271

elegans, Microstylum, 265

Enchocera, 271

Epiblepharis, 264

erythropus, Hoplistomerus, 246

eumenoides, Ammophilomima, 219

Euscelidia, 216, 219

evanescens, Ammophilomima. 219

evanescens, Leptogaster, 217

fascipennis, Lasiocnemus, 223
fenestratum, Microstylum, 266
festiva, Euscelidia, 220
flavipes, Laphria, 227
floccosa, Hyperechia, 250
fortipes, Laphria, 227
francoisi, Dolichoscius, 223
francoisi, Gonioscelis, 278
francoisi, Laxenecera, 236, 240
francoisi, Oxynoton, 286
fulvicollis, Ancylorrhynchus, 272
fulvithorax, Dasyllina, 231
fumosa, Leptogaster, 217
funditor, Laxenecera, 236
funebris, Ancylorrhynchus, 271
furunculus, Philodicus, 299

332

INDEX

garambaensis, Hoplistomerus, 246

garambaensis, Ommatius, 323

genitalis, Gonioscelis, 278

Gerrolasius, 225

ghesquierei, Ammophilomima, 219

gigantipes, Proagonistes, 253

glabrum, Microstylum, 266

Glyphotriclis 225

Goneccalypsis, 257

Gonioscelis, 275

Gonypes, 216

griseicinctipes, Lasiocnemus, 224

griseus, Lagodias, 261

gymna, Laxenecera, 237, 242

gymnus, Machimus, 309

Halictosoma, 243
harlequini, Oligopon, 295, 296
helenae, Microstylum, 264, 266
Heligmoneura, 302
hera, Laphria, 227
hermelina, Leptogaster, 217
hermanni, Lasiocnemus, 223
hermanni, Microstylum, 266
hermanni, Stichopogon, 283, 285
hirsutus, Machimus, 309
hirtipes, Microstylum, 265
Hoplistomerus, 225, 246
horizontalis, Laxenecera, 236
humeralis, Ancylorrhynchus, 272
hyalipennis, Lasiocnemus, 223
hyalipennis, Oligopogon, 295
hybotinus, Olipogon, 296
hylaeformis, Ancylorrhynchus, 273
Hyperechia, 226, 249

igniferum, Proagonistes, 253
ignobile, Microstylum, 266
imitatrix, Ammophilomima, 219
imperator, Ommatius, 329
insignis, Ancylorrhynchus, 272
iola, Laphria, 227, 230

juxta, Machimus, 309

Katharma, 226
Key to tribes, 215

lacteipenne, Microstylum, 265

Lagodias, 260

Lagynogaster, 219

lampyroides, Proagonistes, 253, 254

Lamyra, 225

Laphria, 224, 226

Laphriini, 224
Laphystia, 225
Lasiocnemus, 223
laevis, Heligmoneura, 302
lateralis, Laphria, 226, 227
Laxenecera, 225, 226, 235
leoninus, Proagonistes, 253, 254
Leptogaster, 216
Leptogasterini, 215
lituratum, Microstylum, 265
lobatus, Congomochtherus, 306
Loewinella, 257
longipennis, Ommatius, 329
longipes, Dolichoscius, 223
luctuosa, Laphria, 227
ludens, Leptogaster, 217
lugens, Lasiocnemus, 223

Machimus, 309
macquarti, Ommatius, 320
macroscelis, Ommatius, 319
maculatus, Ancylorrhynchus, 272
maculipennis, Gonioscelis, 275
maculipennis, Stichopogon, 283
maculiventris, Eclipsis, 264
magnificus, Ancylorrynchus, 272
major, Rhabdogaster, 281
Margaritola, 286
maynei, Laphria, 227, 230
melanomystax, Leptogaster, 217
metalli, Laphria, 227
Microstylum, 263
Mimoscolia, 263

minor, Storthyngomerus, 231, 232, 234
misema, Laxenecera, 236, 239
modesta, Heligmoneura, 303
montana, Ammophilomima, 219
morio, Rhipidocephala, 287
moyoensis, Euscelidia, 222
multipunctata, Laphria, 227
munroi, Ancylorrhynchus, 272
munroi, Neolaparus, 259
mystaceus, Proagonistes, 253

neavei, Proagonistes, 253
Neoitamus, 313
Neolaparus, 258
nigra, Dogonia, 270
nigrescens, Laphria, 227
nigrescens, Microstylum, 265
nigrescens, Philodicus, 299
nigribarbatum, Microstylum, 265
nigribimba, Laphria, 228, 230
nigrimystaceum, Microstylum, 265

INDEX

333

nigripennis, Oligopogon, 295
nigripes, Loewinella, 257
nigripes, Machimus, 309, 310
nigrociliata, Laxenecera, 238, 243
nigrocuprea, Laxenecera, 241
nitidus, Oligopogon, 295
nomada, Ancylorrhynchus, 272
Nusa, 225, 243
Nusina, 231
nyukinus, Ancylorrhynchus, 271

obscuripennis, Lasiocnemus, 224
occipitalis, Gonioscelis, 275
Oligopogon, 286, 294
Ommatiini, 313
Ommatinus, 317
Ommatius, 317
Opegiocera, 271
ophion, Neolaparus, 259
Orthogonis, 225
Oxynoton, 286

pallida, Smeryngolaphria, 235
pallipes, Leptogaster, 217, 218
pammelas, Machimus, 309
Paractenota, 225
parcum, Microstylum, 266
Paroxynoton, 286
partitum, Microstylum, 266
pedunculata, Epiblepharis, 264
Pegesimallus, 262
pellucida, Leptogaster, 217
penicillata, Leptogaster, 218
penicillatus, Congomochtherus, 306, 308
penicillatus, Oligopogon, 296
Perasis, 225, 244
Philodicus, 298
pica, Microstylum, 265
pictipennis, Leptogaster, 217
pilicnemis, Leptogaster, 218
plebeja, Leptogaster, 217
pliomelas, Proagonistes, 253, 255
pollex, Microstylum, 266
pollinosus, Oligopogon, 295
poseidon, Xenomyza, 287, 291
praeceps, Proagonistes, 253
praedo, Proagonistes, 253
pretoriensis, Ancylorrhynchus, 271
Proagonistes, 226, 251
Promachus, 300
puella, Leptogaster, 217
pulchella, Laxenecera, 234, 239
punctum, Stichopogon, 283, 284
pyragra, Leptogaster, 218

quadrimaculatus, Ancylorrhynchus, 272

redimiculum, Proagonistes, 253, 255
reynaudii, Ancylorrhynchus, 272
Rhabdogaster, 280
Rhipidocephala, 286
ricardoae, Microstylum, 265, 268
ricardoi, Laphria, 228, 230
rodhaini, Heligmoneura, 303
rubripes, Machimus, 309
rufa, Leptogaster, 217,
rufescens, Leptogaster, 217, 218
rufibarbis, Proagonistes, 253, 255
rufineurum, Microstylum, 266
rufitarsis, Laxenecera. 236, 239
rufum, Microstylum, 265

saegeri, Dogonia, 269
saegeri, Pegesimallus, 262
saliodes, Proagonistes, 253
Saropogonini, 257
Saucropogon, 225, 244
schoutedeni, Laphria, 227
schoutedeni, Leptogaster, 217
scopifera, Laxenecera, 238
scutata, Rhipidocephala, 287
scutellata, Xenomyza, 287, 292
Scylaticus, 274
sericea, Leptogaster, 217
serpentina, Laphria, 227
sessile, Microstylum, 265, 269
Smeryngolaphria, 226, 234
sororcula, Laxenecera, 236
spinipes, Microstylum, 266
splendens, Ancylorrhynchus, 272
splendida, Laxenecera, 241
spurinus, Microstylum, 266
Stichopogonini, 281
stichosoma, Leptogaster, 218
Stiphrolamyra, 225
Storthyngomerus, 224, 231
strachani, Astochia, 313
straeleni, Ammophilomima, 219
striatus, Ancylorrhynchus, 272, 274
Strobilothrix, 247
stuhlmanni, Laxenecera, 237, 241
superciliatus, Oligopogon, 295, 297
sussurus, Ancylorrhynchus, 271
swynnertoni, Philodicus, 306
Systropalpus, 225, 249

taciturna, Xenomyza, 287, 289
tarsalis, Leptogaster, 218
temerarius, Philodicus, 299

334

Teretromyia, 298

testacea, Trichardis, 231, 247

Thallosia, 315

tigrina, Rhipidocephala, 286

Tolmerus, 309

tomentosus, Gonioscelis, 276

toroensis, Storthyngomerus, 232, 233

transvaalensis, Perasis, 244, 245

Tribes, key to, 215

Trichardis, 225, 247

tridentatus, Storthyngomerus, 232

Trupanea, 300

ufens, Proagonistes, 253,
ugandiensis, Machimus, 309, 210
ugandiensis, Promachus, 301
unicolor, Microstylum, 266
unicolor, Stichopogon, 282
unifasciatus, Ancylorrhynchus, 272
upembana, Leptogaster, 218
urundiana, Leptogaster, 217

INDEX

usambarae, Hyperechia, 250
ustulatum, Microstylum, 266

validum, Microstylum, 265
variabilis, Laphria, 227
variegatus, Ancylorrhynchus, 272
varipennatum, Microstylum, 266
velutina, Leptogaster, 217
venosum, Microstylum, 265
villosum, Microstylum, 266
vindex, Leptogaster, 217
vittatus, Ommatius, 317
vulpinus, Proagonistes, 253

Xenomyza, 286, 287
Xenomyzini, 285
Xiphocera, 271

zonalis, Ancylorrhynchus, 272
zonata, Dyseris, 241

HAROLD OLDROYD

Department of Entomology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON, S.W.7

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A LIST OF THE TYPE-SPECIMENS OF

PLECOPTERA AND MEGALOPTERA

IN THE BRITISH MUSEUM

(NATURAL HISTORY)

D. E. KIMMINS

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 24 No. 8

LONDON : 1970

A LIST OF THE TYPE-SPECIMENS OF

PLECOPTERA AND MEGALOPTERA

IN THE BRITISH MUSEUM

(NATURAL HISTORY)

BY

DOUGLAS ERIC KIMMINS

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A LIST OF THE TYPE-SPECIMENS

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IN THE BRITISH MUSEUM

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By D. E. KIMMINS

CONTENTS

Page

SYNOPSIS ............ 337

INTRODUCTION ........... 337

NOTES ............ 337

PLECOPTERA ........... 338

MEGALOPTERA ........... 352

REFERENCES ........... 358

SYNOPSIS

In these lists, 171 taxa of Plecoptera and 74 taxa of Megaloptera are considered. Lectotypes
are designated for 43 taxa of Plecoptera and 21 taxa of Megaloptera. The types of 8 other
taxa of Plecoptera are either unlabelled and not recognizable from the descriptions, or have
not been traced, being possibly in other collections.

INTRODUCTION

THIS list follows the general pattern of my previous Odonata Type Lists, but as the
Plecoptera and Megaloptera are relatively small orders, I have not considered it
necessary to present separate lists for the various families, all the taxa being dealt
with alphabetically under the respective orders, and a combined list of references
is given.

NOTES

1. The recognition of the individual types of the Plecoptera and Megaloptera
described by Stephens presents some difficulties, since at some time since its
presentation to the BMNH, the Stephens Collection was re-arranged by what
McLachlan has described as a 'non-expert'. The specimens in most cases bore only a
Stephens Collection label and were arranged over a small handwritten determination
label. In few cases was there any indication of locality. During this re-arrange-
ment, the various synonyms were placed together and the determination label
placed on one of the series, with the result that it is almost impossible to differentiate
the various type-series. In labelling types, the example bearing a determination
label has been accepted as type, unless there is definite evidence to the contrary.

2. In the case of some of the Leptoperlid types from Australia, reference is
made to the wings being mounted in de Faure's medium, a water-soluble mountant.
This was recommended to me by the late Dr F. W. Edwards in the early j_94o's.
It works quite well for wings and other very thin objects, particularly if the mounts
are ringed with varnish. I found it however quite unsuitable for use with thicker
mounts, such as whole Leptoperlids. Shrinkage of the medium in drying caused

338 D. E. KIMMINS

air to be drawn in and these preparations had eventually to be dissolved in warm
water, dehydrated and cleared and remounted in Canada balsam.

3. The Walker Type label is circular, bearing the word 'Type' and ringed with a
green line. McLachlan's type label is small, rectangular, pink, red or purplish,
bearing the word 'Type'.

PLECOPTERA

abdominalis Stephens (Leuctra), 1836 : 145; Mosely, 1932 : 33, pi. 2, fig. ja,. LECTOTYPE
9. Stephens Coll. / abdominalis / klapaleki $ (not albida), det. K. G. Blair / Leuctra
fusciventris Steph. (klapaleki Kny) M. E. Mosely det. / Leuctra abdominalis Steph., $ Lecto-
type, D. E. Kimmins det. 1969.

The $ type is in fragments. Mosely (1932 : 33) restricted the type-series to the 9 with
abdomen mounted in Canada balsam, and figured on pi. 2, fig. ja. Currently placed as a
synonym of L. fusciventris Stephens.

abnormis Newman (Perla), 1838 : 177-178. Holotype 9- Abnormis Newman, Ent. Mag.
V.i 77, Trenton Falls, N. A. (R. Foster) / Perla abnormis Newm., 9 Holotype.

Newman's label is much blackened and has been transcribed. Currently placed in
Acroneuria.

acicularis Despax (Chloroperla), 1934 : 3^7~37°> pis x> ng- 4> 4. ng- T5I IT. ng- 3°.' text-figs
22-24. Holotype <§ (in 2% formaldehyde solution and on two microscope preparations).
[France, Cantal], Le Lioran, 9-i9.vii.[i9]24 (M. E. Mosely), Chi. acicularis Despax, <$ Type.
Currently placed in Isoperla.

aethiops Walker (Perla), 1852 : 154. Holotype $. Walker type-label / Mexico / 45. PERLA
^ETHIOPS [printed label from Walker's catalogue] / Anacroneuria [C. G. Froelich, 1968].
Currently placed in Anacroneuria.

affinis Stephens (Nemoura), 1836 : 141. LECTOTYPE [sex unknown]. Stephens Coll. /
Nemoura affinis Steph., Lectotype, D. E. Kimmins det. 1969.

The lectotype lacks its abdomen, sex unknown. Currently placed as a synonym of Nemoura
cinerea (Retzius).

albidipennis Walker (Nemoura), 1852 : 191; Ricker, 1938 : 133, fig. 8. Holotype 9-
Walker type-label / R [on green paper] / Nova Scotia (Redman) / Nemoura albidipennis Walk.,
9 Holotype, D. E. Kimmins det. 1959.

Hind wings mounted dry between celluloid, metathorax and abdomen cleared and mounted
in Canada balsam. Currently placed in Nemoura (Ostrocerca) .

albomacula Kimmins (Udamocercia), 1951 : 90-91, figs 39, 4oa-c. Holotype^. Tas[mania],
Cradle Mt, i6.i.[i9]i7 (R. ] . Tillyard). / Udamocercia albomacula Kimm., £ Type.

One pair of wings mounted dry, genitalia in Canada balsam, attached to specimen pin.
Currently placed in Kimminsoperla.

anglica Aubert (ssp. Capnia vidua), 1950 : 315-316, fig. 35. Holotype^ (in 2% formaldehyde
solution). [England], Cheshire, Woodhead, i.iv.i933 (H. Britten) / Capnia anglica Aub.,
cJ Holotype, $ allotype, designated by D. E. Kimmins (i.l.), 1950.

anglica Kimmins (Rhabdiopteryx), 1943 : 42-44, 8 text-figs. Holotype $. [England],
Yorkshire], Pickering Beck, 28.iii.i942 (H. Whitehead) / Rhabdiopteryx anglica Kim.,
$ Holotype.

bifasciata Kimmins (Udamocercia), 1951 : 91-92, figs 4od-f. Holotype <^. Tasmania

(/. W ' . Evans) / Udamocercia bifasciata Kim., $ Type.

Mounted as two preparations, in Canada balsam. Currently placed in the genus
Kimminsoperla.

TYPE-SPECIMENS OF PLECOPTERA & MEGALOPTERA IN BMNH 339

biloba Newman (Pteronarcys), 1898 : 176. Holotype $. Pteronarcys Newman. Biloba
Newman, Ent. Mag. V. 176, Trenton Falls, N.A. (R. Foster).

The label is blackened but still legible. Currently placed as Pteronarcys (Allonarcys).

bituberculata Kimmins (Nemoura), 19500 : 208-209, fig- J7- Holotype $. Assam, Delai
valley, Taphlogam, 4000 ft, ii.xi.i936 (M. Steele) / Nemoura bituberculata Kimm. $ H.T.
Type.

Mounted as two preparations, in Canada balsam.

brevipennis Kimmins (Dinotoperla), 1951 : 67-68, figs i8a-e. Holotype <$. [Australia],

New South Wales, Bolaro, 22.xii.i935 (R. J. Tillyard) / Dinotoperla brevipennis Kim., $ TYPE.

Mounted as two preparations, wings in de Faure's medium, remainder in Canada balsam.

bullata Kimmins (Spaniocerca), 1951 : 88-90, figs 38a-e. Holotype <$. Tasmania (/. W.
Evans) / Spaniocerca bullata Kimmins, $ TYPE.
Mounted as two preparations in canada balsam.

californicus Newport (Pteronarcys), 1848 : 388; 1851 : 450. Holotype <$. California
(Hartweg) / Pteronarcys californica Newport, Proc. Linn. Soc. 1848 / Pteronarcys californicus
Newp., <$ Holotype, D. E. Kimmins det. 1969.

The first description of this species is in Proc. Linn. Soc. Lond. for 20. vi. 1848, and it should
be noted that the actual date of publication of p. 388 was 1848, but p. 389 was not published
until 1849. The paper subsequently appeared in the Trans. Linn. Soc. Lond. 20 (3) : 444-452,
in 1851.

[cambrica Stephens (Nemoura), 1836 : 143. I have been unable to recognize the type-
specimen of cambrica in the Stephens Collection, the three possible examples (females) being
smaller than the dimensions given by Stephens.]

[carpenteri Tillyard (Dinotoperla), 19216 : 270-274, 4 text-figs.

The type of this species did not come to the BMNH with the Tillyard Bequest. It may
still be in the Cawthron Institute, Nelson, N.Z.]

chrysostoma Klapalek (Brahmana), 1916 : 63. LECTOTYPE $. Mungphu (Atkinson) /
chrysostoma, Klapalek / Brahmana chrysostoma Klap., $ Lectotype, D. E. Kimmins det.
1969.

citronella Newport (Perla), 1848 : 388; Walker, 1851 : 169-170 (as citrinella); Ricker, 1948 :
144-145, figs 28-30 ($ lectotype designated). Lectotype $. [Canada], Hudson's Bay,
[Albany River, St. Martin's Falls] / 269 or 668, Perla citronella / Perla citronella Newp.,
<$ Lectotype, design. W. E. Ricker, 1938.

Currently placed in Isoperla. For note on date of publication, see californicus (Pteron-
arcys), p. oo.

clio Newman (Isogenus), 1839 : 415. LECTOTYPE <£. Georgia / Perla Clio / Isogenus clio
Newm., ^ Lectotype, D. E. Kimmins det. 1969.

The lectotype has the abdomen cleared and mounted in canada balsam. There is one
<£ paralectotype.

clymene Newman (Chloroperla), 1839 : 87. Holotype $. Georgia / Clymene Newm. /
clymene, Klapalek.

Currently placed in Neoperla.

compacta McLachlan (Dictyopteryx), 1872 : 53-54, pi. i, figs 6, 7~7b. LECTOTYPE $.
Type [McL. label] / Sibir. Or. (Maack) / compacta McL. / Dictyopteryx compacta McL.,
<£ Lectotype, D. E. Kimmins det. 1969.

The BMNH also possesses the $ paralectotype of this species. Currently placed in the
genus Arcynopteryx.

completa Walker (Nemoura), 1881 : 191-192; Ricker, 1938 : 133. Holotype <$. Walker
type-label / R [on green paper] / N[ova] Scotia (Redman) / 52. NEMOURA COMPLETA [label
cut from Walker's catalogue] / Nemoura. W. R[icker].

340 D. E. KIMMINS

The abdomen of the type has been cleared and mounted in canada balsam. Currently
placed in Nemoura (Prostoia), of which it is the type-species.

concolor Banks (Peltoperla) , 19310, : 411-412. LECTOTYPE £. B[ritish] N[orth] Borneo,
Mt Kinabalu, Kamborangah, 7000 ft, 27.^.1929 (H. M. Pendlebury) [ex F.M.S. Museum] /
Peltoperla concolor Bks, type / Peltoperla concolor Bks, £ Lectotype, D. E. Kimmins det. 1969.

cordata Kimmins (Capnia), 1947 : 733-734, figs 9, A-D. Holotype <$. Tibet, Gyantse,
13000 ft, 28. vi. 1928 (F. M. Bailey] / Capnia cordata Kimm., $ Type, 1946, D. E. Kimmins
det.

Type pinned (in poor condition), abdomen cleared and mounted in canada balsam.

cruciata Stephens (Nemoura), 1836 : 141. LECTOTYPE $. Stephens Coll. / cruciata /
Nemoura cruciata Steph., $ Lectotype, D. E. Kimmins det. 1969.
Currently placed as a synonym of Nemoura cinerea (Retzius).

corsicana Morton (Netnura (Protonemura)), 1930 : 80-8 1, pi. 2, figs 3-5. LECTOTYPE^.
Corsica, Corte, 2i.v.-8.vi.i928 (M. E. Mosely) / Nemoura corsicana Morton, <$ Lectotype,
D. E. Kimmins det. 1969.

Morton did not specify a type nor its location, but Mosely had labelled the preparations,
from which Morton's genitalia figures were drawn, as Type. To validate his action, I have
designated these preparations as Lectotype. Currently placed in the genus Protonemura.

cydippe Newman (Chloroperla), 1839 : 88. Lectotype $ (Ricker, 1938 : 148, fig. 38).
Georgia, i8a / 88. PERLA CYDIPPE (label cut from Walker's catalogue) / Chloroperla cydippe
Newman, $ Lectotype, W. E. Ricker, 1938.

One pair of wings mounted dry, abdomen in canada balsam. Currently placed in
Hastaperla.

[cymodoce Newman (Per/a), 1839 : 37. No types of this species found in BMNH.]

cyrene Newman (Chloroperla), 1845 : 853; Kimmins, 1938 : 564-566, figs 3-4. Holotype^.
Saunders [Collection] / Chloroperla Cyrene, New Zealand.

One pair of wings mounted dry, abdomen in canada balsam. Currently placed in
Austroperla.

decisa Walker (Per la (Chloroperla)), 1852 : 170. Holotype^. Walker type-label / Hudson's
Bay, [Albany River, St Martin's Falls] / Perla (Chloroperla) decisa Walker, $ Holotype, D. E.
Kimmins det. 1969.

The holotype has one pair of wings mounted dry between celluloid and the abdomen
cleared and mounted in canada balsam. Currently placed in Isogenus (Cultus).

decolorata Walker (Perla (Chloroperla)), 1852 : 170; Ricker, 1938 : 145-146, fig. 31.
LECTOTYPE $. Walker type-label / N[orth] Amer[ica], Gt Bear L[ake] / 98. PERLA
DECOLORATA [label cut from Walker catalogue] / Perla decolorata Walker, $ Lectotype, D. E.
Kimmins det. 1969.

Abdomen of lectotype cleared and mounted in canada balsam, for study by Ricker.
Currently placed in Isoperla. There is also one incomplete paralectotype.

despaxi Mosely (Leuctra), 1930 : 249-250, figs 27-28. Holotype <J. France, Pyrenees
Orientales, R. Tet, Mont Louis, 22.vi-2.vii.i923 (M. E. Mosely) / Leuctra despaxi Mosely^
Type.

The holotype is a whole insect, cleared and mounted as a microscope preparation in euparal.

dichroa McLachlan (Dictyopteryx), 1872 : 52-53, pi. i, figs 4-43, 5~5b. LECTOTYPE $.
Type [McL. label] / Sibir. orient. (Maa\c]k) / dichroa McL. / Dictyopteryx dichroa McL.,
<$ Lectotype, D. E. Kimmins det. 1969.

The type-series in BMNH includes i <$ and i $ paralectotypes. Currently placed in the
genus Arcynopteryx.

TYPE-SPECIMENS OF PLECOPTERA & MEGALOPTERA IN BMNH 341

diminuta Kimmins (Megaleptoperla), 1938 : 568-570, figs 6, 7. Holotype £. New Zealand,
Ohakune, 1922-1923 (T. R. Harris) / Megaleptoperla diminuta Kimmins, $ Type, det. D. E.
Kimmins.

The abdomen of the type has been cleared and mounted in Canada balsam. Fig. 2 shows
the ? wings, not those of the <$; the anterior branch of 3 A is erroneously shown as forked.
[diversipes Tillyard (Tasmanoperla), 1931 : 41.

The type of this species did not come to the BMNH with the Tillyard Bequest collection.]
divisa Klapalek (var. of Togoperla perpicta Klapalek), 1921 : 64. LECTOTYPE <J. N.
China [Shanghai, Fortune Coll.] / infuscata Klapalek / var. divisa / Togoperla perpicta var.
divisa Klap., <$ Lectotype, D. E. Kimmins det. 1969.

This specimen has been labelled 'Type' for many years. The second syntype, from China,
is Walker's spec, 'b' of infuscata and is a $, not a $ as stated by Klapalek.

dorsalis Kimmins (Kyphopteryx) , 1947 : 725-727, figs iB, 3A-G. Holotype <£. Tibet,
Chumbi Valley, nooo ft, 2.iv.i924 (R. W. G. Hingston), Everest Exped. / Kyphopteryx
dorsalis Kim., <$ Type.

One pair of wings mounted dry, the other pair and the remainder of the type in Canada
balsam (3 slides).

drytno Newman (Isogenus), 1839 : 86. LECTOTYPE ?. Georgia / Perla drymo [with
addition 'OK WR'] / Isogenus drymo Newman, $ Lectotype, D. E. Kimmins det. 1969.
There is also a $ paralectotype. Currently placed in Perlinella.

ephyre Newman (Chloroperla), 1839 : 87-88. Holotype $. Georgia / 17. Ephyre Newman,
Georgia, 17 / Having only 2 ocelli this species belongs (according to Klapalek) to genus
Atoperla, C.J.G.

ettlegnica Tillyard (ssp. of Eusthenia spectabilis), 19210 : 230, pi. 13, fig. 4. Holotype $.
Tasmania, Tyenna, 29.xii.[ig]i6 (C. E. Cole) / Eusthenia spectabilis eulegnica n. subsp.,
$ Holotype, R.J.T.

evansi Kimmins (Dinotoperla), 1951 : 70-71, figs 2oa-c, 2ia. Holotype <J. S. Australia,
Mt Lofty, x.i 93 1 (J. W. Evans) / Dinotoperla evansi Kim., <$ TYPE.

Mounted as three preparations, wings dry, remainder in canada balsam.

exigua Kimmins (Leptoperla) , 1951 : 53-54, figs 6a-d. Holotype $. W[est] Australia,
Kelmscott, 22.xi.[i9]32 (R. J. Tillyard) / Leptoperla exigua Kim., $ TYPE.

Mounted as two preparations, wings in de Faure's medium, remainder in canada balsam.

exilis McLachlan (Perla), 1872 : 54-55, pi. i, figs 8, 8a. Holotype $. Type [McL. label] /
Sibir. orient. (Maa\c]k) / exilis McL. / Perla exilis McL., <$ Holotype, D. E. Kimmins det. 1969.
lilies (1966, Tierreich, 32 : 501) treats this as a nomen oblitum.

extensa Tillyard (ssp. Eusthenia purpurescens), 19210 : 230. Holotype $. Tasmania,
Russell, 26.xii.i9i6 (C. E. Cole) / Eusthenia purpurescens extensa n. subsp., 9 Holotype,
R.J.T.

falcula Kimmins (Protonemura), 19500 : 196-198, figs 1-3. Holotype <$. Assam, Mishmi
Hills, Chhaglon, 5350 ft, 26. ii. 1935 (M. Steele) / Protonemura falcata Kim., $ TYPE.
Mounted as two preparations in canada balsam.

fascipennis Banks (Javanita), 1931 : 378. LECTOTYPE <$. Peninsular Siam, Nakon
Sri Tamarat, Khao Ram, 750 ft, 24.11.1922 (H. M . Pendlebury) / Ex F.M.S. Museums / Javanita
fascipennis Bks type / Javanita fascipennis Bks, <$ Lectotype, D. E. Kimmins det. 1969.
Currently placed as Neoperla.

fasciata Tillyard (Dinotoperla), 1924 : 193. Holotype $. Queensland], National Park,
1500-2000 ft, 2O.ii.[i9]2i [A. J. Turner] / Dinotoperla fasciata Till., Holotype $, R.J.T.
One pair of wings mounted dry, in celluloid.

ferruginea Walker (Nemoura (Leuctra)), 1851 : 183; Ricker, 1938 : 134, fig. (Lectotype $
designated). Lectotype $. Walker type-label / R on green label [Redman] / Nova Scotia

342 D. E. KIMMINS

(Redman) / 18. NEMOURA FERRUGINEA [label cut from Walker catalogue] / Nemoura ferruginea
Walker, $ Lectotype, design. W. E. Ricker, 1938.

Currently placed in the genus Leuctra. Ricker's designation of lectotype appears a little
vague, but as he states that the lectotype was used as the basis of the figure, and only one
example was cleared for drawing, that specimen only can be the lectotype. There are two
paralectotypes.

filigera Kimmins (Amphinemura), 1947 : 730-731, figs 6A-C. Holotype ^ [India], Bengal,
Darjeeling distr., xi.i945 (D. E. Kimmins) / Amphinemura filigera Kim., <$ Type.
Type with wings mounted dry, remainder in Canada balsam.

flavescens Kimmins (Nesoperla), 1938 : 570-571, figs 8A-C. Holotype $. New Zealand,
[S. Island], L. Wakatipu, ii.ign (G. V. Hudson) / Nesoperla flavescens Kimm., $ Type, det.
D. E. Kimmins.

One pair of wings mounted dry, apex of abdomen in Canada balsam.

flavomaculata Mosely (Leuctra), 1935 : 560-561, figs 4-5. Holotype <$. France, Puy-de-
Dome, Le Mont-Dore, 24.vi-6.viii.i934 (M. E. Mosely) / Leuctra flavomaculata Mosely,

c? TyPe-

The £ type was taken coupled, and the two specimens have been cleared and mounted as a
preparation in euparal. Mosely specified the $ as Type, the $ as paratype.

flavotincta McLachlan (Perla), 1872 : 54, pi. i, figs g-ga. Holotype $. Type [McL. label] /
Sibir. Or. (Maack) / flavotincta McL. / Perla flavotincta McL., $ Holotype, D. E. Kimmins
det. 1969.

Currently placed in the genus Paragnetina.

fontana Kimmins (Dinotoperla), 1951 : 64-66, figs i6a-e. Holotype $. Australia /
F[ederal] C[apital] Territory], Lee's Springs, xi.i932 (R. J. Tillyard) / Dinotoperla fontana
Kim., <J TYPE.

Mounted as three preparations, wings in de Faure's medium, remainder in Canada balsam.

foveolata Klapalek (Neoperla), 1921 : 321. LECTOTYPE $. Hong Kong / foveolata
Klapalek / triangle of black paper / Neoperla foveolata Klap., $ Lectotype, D. E. Kimmins
det. 1969.

There are also i $, i $ paralectotypes from N. China.

fraterna Banks (Nogiperla), 1938 : 223, figs 7, 14. Holotype $. Federated] M[alay]
S [tales], Pahang, Cameron's Highlands, 4800-5500 ft, 8.vi.ig35 (H. M. Pendlebury) / E.
F.M.S. Museums / Nogiperla fraterna Banks, Type.

fraterna Morton (Leuctra), 1930 : 79, pi. 2, figs 6-7; Mosely, 1932 : 20. LECTOTYPE ,$.
Corsica, Corte, 2i.v.-8.vi.i928 (M. E. Mosely) / Leuctra fraterna Morton [M.E.M. label] /
Leuctra fraterna Morton, $ Lectotype, D. E. Kimmins det. 1969.

Mosely (1932) stated that the type was a preparation in his collection and he placed a
BM type-label on it. This has now been replaced by my Lectotype label.

frontalis Newman (Isogenus), 1838 : 178. LECTOTYPE $ [incomplete]. Isogenus Newman.
Frontalis Newman, Ent. Mag. V, 178, Trenton Falls, N. A. (R. Foster) / Isogenus frontalis
Newman, $ Lectotype, D. E. Kimmins det. 1969.

Currently placed as Isogenus (I sogenoides) . There is one 9 paralectotype, labelled R.
Foster, New York.

fuliginosa Stephens (Nemoura), 1836 : 141. LECTOTYPE $. Stephens Coll. / fuliginosa /
Nemoura fuliginosa Steph., $ Lectotype, D. E. Kimmins det. 1969.
Currently placed as a synonym of Nemoura cinerea (Retzius).

fumosa Stephens (Nemoura), 1836 : 143. LECTOTYPE [?sex]. Stephens Coll. / fumosus /
Nemoura fumosa Steph., Lectotype, D. E. Kimmins det. 1969.

Morton (1894 : 571) states that this is Nemoura cinerea Olivier [= Amphinemura sulcicollis
(Stephens)]. The type now lacks its abdomen.

TYPE-SPECIMENS OF PLECOPTERA & MEGALOPTERA IN BMNH 343

fusca Kimmins (Dinotoperla), 1951 : 71-72, figs 21 b, c. Holotype $. Tasmania (/. W.
Evans] / Dinotoperla fusca Kim., $ TYPE.

Mounted as two preparations in Canada balsam.

\Juscipennis Stephens (Chloroperla), 1836 : 138. Type unlabelled and not recognizable
from Stephens' description.]

fusciventris Stephens (Leuctra), 1836 : 145; Mosely, 1932 : 32-33, text-figs 50-51, pi. 2,
fig. 7. Lectotype <$. Stephens Coll. / fusciventris / Leuctra fusciventris Steph., <$ Lectotype,
D. E. Kimmins det. 1969.

Mosely (1932 : 33) restricted the type-series to the <$, of which the abdomen had been
cleared, mounted in Canada balsam and figured on pi. 2, and in effect designated this specimen
as a lectotype.

geniculata Stephens (Leuctra), 1836 : 145. LECTOTYPE $. Stephens Coll. / geniculata /
Leuctra geniculata Steph., $ Lectotype, D. E. Kimmins det. 1969.

One other $ example remains in the Stephens Coll., and has been labelled as a paralectotype.

glacialis Newport (Netnoura (Brachyptera)), 1849 : 389; 1851 : 451; Walker, 1851 : 192;
Ricker, 1938 : 131, 132, figs 1-7 (lectotype designation). Lectotype $. Hudson's Bay,
[Albany River, St Martins Falls] / Nemoura glacialis Newport Holotype (= Taeniopteryx,
s. 1.) W. E. Ricker.

The lectotype has one pair of wings mounted dry and the abdomen in canada balsam.
For notes on date of publication, see Pteronarcys californicus Newport.

hamulata Morton (Isopteryx), 1930 : 78-79, pi. 2, figs 8-9. LECTOTYPE $. Corsica,
Corte, 2i.v-8.vi.i928 (M. E. Mosely) / Isopteryx hamulata Morton, $ Type [M.E.M. label] /
Isopteryx hamulata Morton, <$ Lectotype, D. E. Kimmins det. 1969.

Type mounted as two preparations, one in canada balsam, one pair of wings mounted dry.
I consider this to be a synonym of Chloroperla apicalis (Newman). Currently placed in the
genus Chloroperla.

hingstoni Kimmins (Capnia), 1947 : 732-733, figs 8A-F. Holotype <$. Sikkim, Gnatang,
12000 ft, 3i.iii.ig24 (R. W. G. Hingston) Everest Exped. Capnia hingstoni Kim., <$ Type.
Type on microscope slides, wings dry, remainder in canada balsam.

hudsoni Kimmins (Spaniocercoides), 1938 : 577-579, figs 13-15. Holotype $. New
Zealand (G. V. Hudson) / Korokoro, 12. x. 1923 / Spaniocercoides hudsoni Kimmins, ^ Type,
det. D. E. Kimmins.

indica Kimmins (Protonetnura ?) 1947 : 727~72^. ngs 4A-C. Holotype <?. [India], Bengal,
Darjeeling district, xi.i945 (D. E. Kimmins).
Type preserved in 2% formaldehyde solution.

insularis Morton (Chloroperla), 1930 : 77-78, pi. 2, figs 1-2. LECTOTYPE <?. Corsica,
Corte, 2i.v-8.vi.i928 (M. E. Mosely) / Chloroperla insularis Morton $ Type / Chloroperla
insularis Morton, <$ Lectotype, D. E. Kimmins det. 1969 (see note to Nemura corsicana,
p. 340).

Lectotype mounted as preparations in canada balsam, one pair of wings mounted dry.
Currently placed in the genus Isoperla.

internata Walker (Perla), 1852 : 152-153; Ricker, 1938 : 139, figs 14, 15. Holotype $.
Walker type label / North America? / Entomological] Club, [:8]44-i2 / 41. PERLA INTERNATA
[label cut from Walker's Catalogue]. /

The type is now much faded. Currently placed in Acroneuria.

intermixta Walker (Perla), 1852 : 153. Holotype $. Walker type label / Venezuela / 42.
PERLA INTERMIXTA [label cut from Walker's Catalogue] / Anacroneuria intermixta (W.) $
[label by C. G. Froehlich, vii.i968].

lilies, 1968 : 503 places this species as a nomen oblitum.

344 D- E. KIMMINS

irrorata Tillyard (Trinotoperla), 1924 : 194, fig. i. Holotype $. N[ew] S[outh] W[ales],
[Mt] Kosciusko, [5000-5500 ft], 24. xi. 1921 (R. /. Tillyard) / Trinotoperla irrorata Till.,
Holotype ?, R.J.T.

Holotype body much damaged by insect pests when received in Tillyard Bequest, one pair
of wings mounted dry between celluloid, <$ allotype also with body much damaged by pests.

ketnpnyi Mosely (Leuctra), 1932 : 14, pi. 3, figs 14, i4a; text-figs 13-14. Holotype $.
France, B[asses] Pyrenees, Eaux-Bonnes, 4-2 i.vii. 1924 (M. E. Mosely} / Leuctra kempnyi
Mosely <$ $. Taken coupled. Type $.

Of the pair, mounted as microscope preparation in euparal, Mosely specifies the $ as type,
the $ as paratype.

[lacrimosa Klapalek (Paragnetina), 1921 : 62. Type not traced in BMNH.]

lacustris Tillyard (Eusthenia), ig2ia : 231-232, pi. 12, fig. 5. Holotype $. Tas [mania],
Cradle Mt., [Lake Lilla], I2.i.[i9]i7 (R. J. Tillyard) / Eusthenia lacustris n. sp., $ Holotype,
R.J.T.

Allotype <J and i incomplete $ paratype also in BMNH.

larvata Klapalek (Tetropina), 1909 : 223 (in key). LECTOTYPE <J. N.W. Borneo, Baram /
larvata, Klapalek / Tetropina larvata Klap., <J Lectotype, D. E. Kimmins det. 1969.
I have selected the BMNH example as lectotype, as it is the example which is figured.

lateralis Stephens (Chloroperla), 1836 : 138. LECTOTYPE <J. Stephens Coll. /Chloroperla
lateralis Steph., <$ Lectotype, D. E. Kimmins det. 1969.

Abdomen mounted in canada balsam. Currently placed as a synonym of Isoperla gram-
matica (Scopoli).

lepida Klapalek (Anacroneuria), 1922 : 91. Holotype (?sex). Cent[ral] Brazil, Chapada,
2600 ft, xi.i9O2 (A. Robert) / lepida, Klapalek / Anacroneuria lepida Klap., Holotype.
Most of abdomen missing.

lugubris McLachlan (Perla), 1875 : 172-173. Holotype $, Japan, [Kobe (Lewis)] / Perla
lugubris, McL. (Type).

Currently placed in the genus Kiotina.

lunata Kimmins (Rhabdiopteryx), 1947 : 722-724, figs lA, 2A-F. Holotype $. Tibet,
Rongbuk, 15500 ft, 22^.1924 (R. W. G. Hingston) / Everest Exped. / Rhabdiopteryx lunata
Kim., cj Type, det. D. E. Kimmins, 1946.

The figures were made from a paratype taken at the same time as the holotype.

lunulata Tillyard (Eusthenia), 19210! : 230-231, pi. 12, fig. 3. Holotype $. Cradle Mountain,
[3000 ft], 21. i. [19117 (R. J. Tillyard) / Eusthenia lunulata n. sp. $ Holotype, R.J.T.

luteicollis Walker (Perla), 1852 : 154-155. Holotype $. Walker type-label / Venezuela /

Perla luteicollis [pencil written] / 46. PERLA LUTEICOLLIS [label cut from Walker's Catalogue].

One pair of wings has been removed and mounted between celluloid, and the abdomen has

been cleared and placed in a small vial of glycerine. The wings are not entirely dark brown

as indicated by Walker but have paler areas. Currently placed in Macrogynoplax.

[luteicornis Stephens (Netnoura), 1836 : 142. I have been unable to recognize the type of
this species. The specimen bearing the label 'luteicornis' is the example with the dissimilar
antennae ; as it is aberrant and does not appear to be an artefact, I am not considering it as
a 'type'. It is Nemoura cinerea (Retzius).]

luteipes Kimmins (Amphinemura), 1947 : 728-730, figs 5A-D. Holotype $. [India],
Bengal, Darjeeling distr., xi.i945 (D. E. Kimmins) / Amphinemura luteipes Kim., <J Type.
Type with wings mounted dry, remainder in Canada balsam.

[lycorias Newman (Perla), 1839 : 35. Newman quotes 'In the cabinets of the British Museum
and the Rev. F. W. Hope.' The BMNH example has not been traced, unless it is the example
described by Newport as Acroneuria sonans]

TYPE-SPECIMENS OF PLECOPTERA & MEGALOPTERA IN BMNH 345

magellanica Klapalek (Kempnyia), 1916 : 69. Holotype <$. Straits of Magellan, Trinidad
Channel, 24.ii.[i8]79, flying over a mountain stream, H.M.S. 'Alert' / magellanica Klapalek.

Currently placed as a synonym of Pictetoperla gayi (Pictet) .

manevali Kimmins (Isopteryx), 1935 : 645-650, figs 1-8. Holotype <£. France, Mt. Mezenc,
5.vi[i9J33 (H. Maneval) / Beaten out of grass, on banks of stream, at foot of mountain, c.
1600 m / Chloroperla manevali (Kim.), <$ TYPE.

Type mounted as two microscope preparations. Currently placed as a synonym of
Chloroperla (Siphonoperla) torrentium (Pictet).

mclachlani Kimmins (Protonemura?), 195001 : 200-203, ngs 5~8. Holotype <$. Assam,
Khasi Hills, McLachlan Collection / Protonemura ? mclachlani Kim., <$ Holotype.

Mounted as two preparations, wings dry, remainder in Canada balsam. Placed provisionally
in Protonemura, in spite of the apparent absence of prosternal gill-vestiges.

media Stephens (Chloroperla), 1836 : 138-139. LECTOTYPE ?. Stephens Coll. / media /
Chloroperla media Steph., $ Lectotype, D. E. Kimmins det. 1969.

Another example, with a small label 'D' is possibly one of the series from Darenth Wood in
June. Currently placed as a synonym of Isoperla grammatical (Scopoli).

media Walker (Perla (Isogenus)), 1852 : 145; Ricker, 1938 : 141. Holotype [? sex]. Walker
type-label / Hudson's Bay, [Albany River, St Martins Falls] / 266 or 665, Perla media [Barn-
ston Mss] / The abdomen does not belong to this specimen, P. W. Claassen / Perla media
Walker, D. E. Kimmins det. 1969.

The abdomen has been glued on, and according to Claassen, does not belong to this
specimen. Currently placed in Paragnetina.

minima Newport (Perla), 1848 : 388; 1851 : 430; Walker, 1851 : 183; Ricker, 1938 : 136.
LECTOTYPE <J. Hudson's Bay [Albany River, St Martins Falls] / 268 or 667, Perla
minima / Allocapnia W. R[icker]. /

Type mounted on two microscope slides. Currently placed in Allocapnia. For notes on
date of original publication, see Pteronarcys californicus Newport, (p. 339).
minor Klapalek (Arcynopteryx), 1912 : 22. Lectotype $. (Ricker, 1938 : 144, figs 25-26).
Arctic America / Klapalek, minor / Perlodes (Arcynopteryx) minor Klap. $ 'Type' selected
by Ricker, 1935-36 [label, D.E.K.].

Klapalek lists a pair in the BMNH collection, and I consider Ricker's statement (1938 : 144)
'Figured here are the <$ genitalia, from the type' as equivalent to a selection of the <$ as
Lectotype. The $ genitalia were cleared by Kimmins and mounted in Canada balsam for
the purpose of Ricker's study. Currently placed as a synonym of Arcynopteryx compacta
McL. The female has been labelled paralectotype.

minor Kimmins (Spaniocerca), 1938 : 575-576, figs n, I2A-C. Holotype^. New Zealand,
Arthur's Pass, 3000 ft, 16.4.1935 (G- V • Hudson) / Spaniocerca minor Kimmins, <$ Type, det.
D. E. Kimmins.

The figures in the original description are from a paratype.

minor Kimmins (Trinotoperla), 1951 : 78-79, figs 2ga-e. Holotype <$. [Australia], N[ew]
S[outh] W[ales], Bolaro, xii.i935 (R. J. Tillyard] / Trinotoperla minor Kim., $ TYPE.

Mounted as three preparations, wings dry, body in de Faure's medium.

minuta Klapalek (Anacroneuria), 1922 : 89. Lectotype <J (Froehlich des. (i.l.) 1968).
Brazil, Santarem / minuta Klapalek / Anacroneuria minuta Klap., <J Lectotype, des. Froehlich
1968.

Wings mounted dry, abdomen cleared and in glycerine, i <$ paralectotype, Brazil, Tapayos.

mishmica Kimmins (Protonemura), 19500 : 203-204, figs 9-11. Holotype £. Assam,

Mishmi Hills, Chhaglon, 5350 ft, 26. ii. 1935 (M. Steele) / Protonemura mishmica Kim., <$ TYPE.

Mounted as two microscope preparations, in Canada balsam.

montana Kimmins (Capnia), 1944 : 735-736, fig. nA. Holotype $. Sikkim, Tangu,
11500 ft, 26.iv. 1924 (R. W. G. Hingston) Everest Exped.
Type in poor condition, cleared and mounted in canada balsam, on two slides.

346 D. E. KIMMINS

tnontana Kimmins (Protonemura), 1941 : 89-93, figs J3> J4- Holotype $. Westmorland,
Rydal Beck, 1700 ft, 4.viii.i94O (Noel Hynes) / Protonemura montana Kim., $ Holotype.
Holotype mounted on two microscope slides.

montivaga Kimmins (Capnid), 1947 : 736-737, fig. u B. Holotype $. Tibet, Lamna La,
15000 ft, 17. vi. 1924 (R. W. G. Hingston) Everest Exped. / Capnia montivaga Kim., <$ Type.
1946. D. E. Kimmins det.

Type in poor condition, abdomen mounted in canada balsam.

mosellae McLachlan (Perld), 1895 : 1 11-112. LECTOTYPE $. Type [McL. label] / Alf,
1894, Moselle / P. selysii Pict. v. mosellae McL. / Perla selysii Pict. var. mosellae McL. Q*
Holotype, D. E. Kimmins det. 1969.

There are i <$ and 10 $ paralectotypes. Currently placed as a synonym of Marthamea
selysi Pictet

moselyi Despax (Chloroperla), 1934 : 3?6-379, pis 5, fig. 16, 10, fig. 29, text-figs 30-32.
Holotype <$ (in 2% formaldehyde solution and on two microscope preparations). [France,
Pyrenees-Ori6ntales], Mont-Louis, R. Tet, 22.vi-2.vii.i923 (M. E. Mosely), Chi. Moselyi
Despax $ Type.

Currently placed in Isoperla.

moselyi Despax (Nemura), 1934 : 265-267, figs 5-8. Holotype Q* (in 2% formaldehyde
solution). France, B[asses] Pyrenees, Eaux-Bonnes, 4-2 i.vi. 1927 (M. E. Mosely), Nemura
moselyi Despax, Type.

The generic name is now spelt Nemoura.

nebulosa Stephens [nee Latreille] (Nemoura), 1836 : 140. LECTOTYPE $. Stephens Coll.
/ nebulosa / Nemoura nebulosa Steph., $ Lectotype, D. E. Kimmins det. 1969.
Currently placed as a synonym of Nemoura cinerea (Retzius).

nigricoxa Kimmins (Dinotoperla), 1951 : 72, fig. 2od. Holotype $. New South Wales,
Mt Kosciusko, io.xii.i934 (^- /• Tillyard) / Dinotoperla nigricoxa Kim., $ TYPE.
Mounted as three preparations, wings dry, remainder in Canada balsam.

nigrifrons Kimmins (Leptoperla), 1951 : 58-59, figs loa-d. Holotype $. Tasmania (/. W.
Evans] / Leptoperla nigrifrons Kim., <$ TYPE.

Mounted as two preparations, wings dry, remainder in canada balsam.

ni mbore lla Mosely (Protonemoura), 1930 : 250-252, figs 29-31. Holotype ^ Switzerland,
Klosters, 3o.viii.i927 (M. E. Mosely) / Nemoura nimborella Mosely, <$ Type.

Mounted as preparation in canada balsam. Of the two abdomens in this preparation, that
in dorsal view should be considered the TYPE.

niponensis McLachlan (Perla), 1875 : 172. LECTOTYPE $. Type [McL. label] / Japan
(Pryer) / Perla niponensis McL. / Perla niponensis McL., $ Lectotype, D. E. Kimmins det.
1969.

Of the two females referred to in the original description (in Wormald's collection), one is
in McLachlan's collection and is designated Lectotype.

nitida Kimmins (Neoperla), 1950 : 184-185, figs 9, 10. Holotype <$. S. India, Tinnevelly
Dt., Naraikadu, 2500-3000 ft, 3-8.x.[i9J38 / Neoperla nitida Kim., <$ Type.

Mounted as two preparations, wings dry, remainder cleared and in canada balsam.

nitida Stephens (Nemoura), 1836 : 143. LECTOTYPE $. Stephens coll. / nitida / in-
conspicua Pict., picteti Mort., det. K. G. Blair / Nemoura nitida Steph., $ Lectotype, D. E.
Kimmins det. 1969.

Placed as a synonym of Nemurella picteti Klapalek. There are also two paralectotype $$.

nivata Kimmins (Trinotoperla), 1951 : 78, figs 27b, 28c. Holotype $. [Australia], Victoria,
Snowy River, 3. i. 193 3 (R. J. Tillyard) / Trinotoperla nivata Kim., $ TYPE.
Mounted as three preparations, wings dry, body in canada balsam.

TYPE-SPECIMENS OF PLECOPTERA & MEGALOPTERA IN BMNH 347

nubecula Newman (Isogenus), 1833 : 415. Holotype $. Isogenus Newman, Nubecula
Newman, Ent. Mag. i. 415, Worcester (P. Burlingham).
The label is much blackened and has been transcribed.

nubila Kimmins (Amphinemura), 1950 : 191-192, fig. 26. Holotype 9- S. India, Coonoor,
6000 ft, 22-23. iv. 1937 (G- M. Henry).

Type mounted as two preparations, body in Canada balsam, wings dry. There is also a $
paratype, pinned.

olivacea Walker (Perla (Isogenus), 1852 : 144-145; Ricker, 1938 : 143, figs 21-23. Holo-
type o"- Walker Type label / Hudson's Bay [Albany River, St. Martin's Falls] / 267 or 666,
Perla olivacea [Barnston Mss] / Isogenus olivacea Walk. Type [DEK writing].

The abdomen was removed and cleared, for study by W. E. Ricker, and has been mounted
in canada balsam.

opposita Walker (Perla (Chloroperla)), 1852 : 171; Kimmins, 1951 : 63-64, fig. 15 (lectotype
designation.) Lectotype <$. Walker Type label / V[an] D[iemens] L[and] / Dinotoperla
opposita Walk., Type, det. D. E. Kimmins, 1.1940.

The abdomen is mounted in de Faure's medium, attached to pin, and one pair of wings
mounted on a microscope slide. I consider my statement (1951 : 'I have selected as type the
female from Mr Smith's collection, labelled V. D. L., 51-153' as equivalent to a designation
of the lectotype. Currently placed in the genus Dinotoperla.

oxylepis Despax (ssp. of Chloroperla grammatica), 1936 : 357, pi. 3, fig. 12, pi. 7, fig. 23,
text-figs 10-12. Holotype <$ (in 2% formaldehyde solution and on two preparations).
[France], Le Lioran, 9-i9.vi.[ig]24 (M. E. Mosely), Chi. gr. oxylepis Despax, Type.

Currently placed in the genus Isoperla. Despax quotes Gerardemer and Le Lioran in the
type-series, but has labelled the Le Lioran example as Type.

[pallicornis Stephens (Nemoura), 1836 : 143. Type not recognized. The specimen bearing
this label agrees in wing expanse with the description but the antennae, far from being pale,
are dark reddish brown. The specimen bears a label 'H' [PHertford] but I am inclined to the
view that the label 'pallicornis' has been wrongly placed upon it, and do not consider it to
be a type. It is a $ Nemurella picteti Klapalek. Morton doubtfully synonymized this
species with cambrica or inconspicua [picteti}. The labelled specimen is certainly the latter.]
pallida Stephens (Chloroperla), 1836 : 139-140. LECTOTYPE 9. Stephens coll. / pallida /
Chloroperla pallida Steph., 9 Lectotype, D. E. Kimmins det.

Currently placed as a synonym of Chloroperla torrentium (Pictet).

pallida Stephens (Nemoura), 1836 : 141. LECTOTYPE $. Stephens Coll. / pallida /
Nemoura pallida Steph., 9 Lectotype, D. E. Kimmins det. 1969.
Currently placed as a synonym of Nemoura cinerea (Retzius).

pallipes Stephens (Nemoura), 1826 : 142. Holotype <$. Stephens Coll. / pallipes / standfussi
(not inconspicua Pict.), det. K. G. Blair. / Amphinemura standfussi RiScJ, D. E. Kimmins det.
This dried example was determined as A . standfussi Ris by Blair, and while I had little
doubt as to the accuracy of his identification, I have cleared the abdomen and can confirm
his identification. lilies (1966, Tierreich 82 : 47) treats Nemoura pallipes Stephens as a
nomen oblitum, thus avoiding the suppression of the well-known name Amphinemura stand-
fussi Ris.

pedestris Kimmins (Capnia), 1947 : 731-732, figs 7A-C. Holotype $. Tibet, Everest base
camp, Rongbuk Glacier, 16500 ft, v.ig22 (T. G. Longstaff] / Capnia pedestris Kim., <$ Type.

The type has been cleared and mounted in canada balsam. Currently placed in the genus
Allocapnia.

perfecta Walker (Nemoura), 1852 : 191; Ricker, 1938 : 133. Holotype cJ (mounted on three
microscope slides). Walker type-label / R [on green label] / Nova Scotia (Redman) / 51.
NEMOURA PERFECTA [label cut from Walker catalogue] / Nemoura WR. (punctipennis Claassen
syn) [W. E. Ricker's writing, 1935-36].

Currently placed in Nemoura (Paranemoura) .

348 D. E. KIMMINS

perpicta Klapalek (Togoperla), 1921 : 63, 64. LECTOTYPE £. Hong Kong, Feb.-May
(/• /• Walker) / infuscata Klapalek / Togoperla perpicta Klap., <$ Lectotype, D. E. Kimmins
det. 1969.

There are 4 <$, i ? paralectotypes in BMNH. The explanation of the absence of correct
Klapalek det. labels appears to be due to a change of mind between labelling the specimens
and the posthumous publication of the description.

perspicillata Klapalek (Gibosid), 1916 : 61. LECTOTYPE ?. N[orth] China / triangle of
black paper / perspicillata Klapalek / Gibosia perspicillata Klap., 9 Lectotype, D. E. Kimmins
det. 1969.

There are also 2 9 paralectotypes labelled Hong Kong.

postica Walker (Perla (Isogenus)), 1852 : 144; Ricker, 1938 : 143-144. Holotype 9-
Walker Type label / [Mackenzie River], Arctic Amer[ica] / Perla (Isogenus) postica Walk.,
Holotype 9, det. D. E. Kimmins, 1969.

The holotype lacks part of its abdomen, but the presence of eggs in the remainder of the
ahdomen leaves no doubt as to its sex. Currently placed as a synonym of Diura bicaudata
(L.).

[praecox Morton (Nemoura), 1894 : 566-567, pi. 13, Nemoura praecox, figs 1-2.

The BMNH has two syn types (<$, 9) given to McLachlan, labelled Cleghorn, I3.iii / PRAECOX
/ Morton's type, 1894 [McL. writing] / Nemoura praecox Mort., syntype, D. E. Kimmins det.
1969.

The lectotype should be chosen from Morton's collection, Royal Scottish Museum, Edin-
burgh.]

prasina Newman (Chloroperla), 1845 : 853. Holotype $. N[ew] Zea[land] / Saunders
[collection] / Chloroperla prasina, New Zealand.

One pair of wings mounted dry, abdomen in Canada balsam. Currently placed in
Sienoperla.

proteus Newman (Pteronarcys), 1838 : 177. Lectotype $ (Smith, 1917 : 453). E. Double-
day, Trenton Falls, New York, Pteronarcys Proteus Newm. / Pteronarcys proteus Newm. <J.
This (J had at some time acquired an incorrect register number applying to a Trichopteron ;
this has been corrected. The 9 type of proteus has been shown by Smith (1917) to be in-
correctly associated with the <J and is in fact the 9 ofPt. comstocki Smith.

pseudocingulata Mendl (Leuctra), 1968 : 311-314, figs 5-7. Holotype <$. France, Vosges,
Retournemer, 7-3 i.vii. 1930 (M. E. Mosely) / Leuctra carinthiaca Kempny, det. M. E. Mosely
/ Leuctra pseudocingulata Mendl, g Holotype, D. E. Kimmins det. 1968.

Holotype mounted as a preparation in canada balsam. The holotype and paratypes were
labelled by Kimmins at Herr Mendl's request.

purpurescens Tillyard (Eusthenia), 1921 : 230, pi. 13, fig. 6. Holotype 9- [Tasmania],
Hobart, 6.xii.[i9]i3 (G. H. Hardy) / Eusthenia purpurescens sp. n. 9 Holotype, R.J.T.
<J Allotype also in BMNH.

pusilla Stephens (Nemoura), 1836 : 142. LECTOTYPE <J. Stephens Coll. / pusilla /
Nemoura pusilla Steph. $ Lectotype, D. E. Kimmins det. 1969.
Currently placed as synonym of Nemoura cinerea (Retzius).

pyrenaica Mosely (Protonemoura), 1930 : 250, figs 29-31. Holotype (J. France, Basses-
Pyren6es, Eaux Chaudes, io.vii.ig29 (M. E. Mosely) / Protonemoura pyrenaica Mosely
A Type.

Type mounted as preparation in canada balsam.

quadridentata Kimmins (Protonemura), 19500 : 205-206, fig. 12. Holotype 9- Khasis,
Nat[ive] Collector] / Protonemura quadridentata Kim., 9 Type, D. E. Kimmins det. 1948.
Type pinned, with abdomen mounted in canada balsam.

TYPE-SPECIMENS OF PLECOPTERA & MEGALOPTERA IN BMNH 349

regalis Newman (Pteronarcys), 1838 : 176. Holotype ?. [The label is much blackened,
but by oblique lighting the following data can be read] Pteronarcys Newman, Regalis Newman,
Ent. Mag. V. 176, Canada, E. Newman.
The abdomen is now missing.

renata Kimmins (Amphinemura), 19500. : 206-208, figs 13-16. Holotype £. Assam,
Mishmi Hills, Minutang, 3900 ft, 17.11.1935 (M. Steele) / Amphinemura renata Kim., ^ Holo-
type.

Holotype mounted as two preparations, wings dry, remainder in Canada balsam.

reticulata Kimmins (Leptoperla), 1951 : 57-58, figs ga-b. Holotype $. [Australia], New
S[outh] Wales, Mt Kosciusko, Spencers Creek, xii.i932 (R. J. Tillyard] / Leptoperla reticulata
Kim., ? TYPE.

Mounted as three preparations, wings dry, remainder in Canada balsam.

reticulata (Klapalek Mss) Tillyard (Eustheniopsis), 19210 : 233. Holotype $. Tasmania /
reticulata Klapalek / Eustheniopsis reticulata Till., $ Holotype, D. E. Kimmins det. 1969.

[rufescens Stephens (Chloroperla), 1836 : 139. Type not recognizable from description.]

ruficosta Tillyard (Tastnanoperla), 1924 : 193. Holotype $. N[ew] S[outh] W[ales],
[Mount] Kosciusko, [5500 ft], 24. xi. 1921 (R. J. Tillyard) / Tasmanoperla ruficosta Till.,
Holotype $, R.J.T.

rugosa Kimmins (Leptoperla), 1951 : 56-57, figs 8a-e. Holotype <$. Australia, F[ederal]
C[apital] Territory], Lee's Springs, xi.[ig]32 (R. J. Tillyard) / Leptoperla rugosa Kim., <$
TYPE.

Mounted as one preparation, in canada balsam.

scutata Barnard (Aphanicer cello), 1934 : 54O-54T. ngs i6a-f. Holotype cJ. [South Africa],
Wellington Mts, [Witte River], ix.i933 (H- G- W\pod\) / Aphanicercella scutata Brnd, Holo-
type cJ, $ [Barnard's writing].

Holotype 3, paratype $ in 2% formaldehyde solution.

scutigera Kimmins (Protonemura), 19500 : 199-200, fig. 4. Holotype $. Assam, Mishmi
Hills, Delai Valley, Chanliang, Alt. 4840 ft, 25. xi. 1936 (M. Steele) / Protonemura scutigera
Kim., $ TYPE.

Mounted as two preparations in canada balsam.

serricauda Kimmins (Dinotoperla), 1951 : 62-63, ngs J4 a~e- Holotype <J. Tasmania,
R. Ouse, 4.11.1933 (R. J. Tillyard) / Dinotoperla serricauda Kim., <$ Type.

Mounted as three preparations, wings in de Faure's medium, remainder in canada balsam.

signata Walker (Perla), 1852 : 157. Lectotype $. Walker type label / Venezuela / 54.
PERLA SIGNATA [label cut from Walker catalogue] / Anacroneuria signata Walker, Lectotype,
det. Froehlich, 1968.

Currently placed in Anacroneuria.

sonans Newport (Barnston Mss) (Perla), 1851 : 447-449. Holotype $. 265 or 664, Perla
sonnans / abnormis, Klapalek / Perla sonans Newport.

Perla sonans (Barnston Mss) was published in synonymy with Perla abnormis Newman by
Newport and would not normally be available, but Ricker (1938 : 140) treats it as an avail-
able name and thus validates it (Art. u, d, Intern. Rules, 2nd edition). He gives St Martin's
Falls, Albany River, Ontario as the type-locality, but there is no such indication on the
specimen. Barnston's label gives the specific name as 'sonnans' , but the published name is
'sonans' ' .

Currently placed in Acroneuria, probably a synonym of abnormis Newman.

spectabilis Westwood (in Griffiths et al.) (Eusthenia), 1832 : 348; Newman (1839 : 33).
LECTOTYPE $. 41.9.25.2 [New Holland, V. D. L., In exchange with Westwood] / Eus-
thenia spectabilis Westw., in Griff. An. K., N. Holl., (V.D.L.). / Eusthenia spectabilis West-
wood, $ Lectotype, D. E. Kimmins det. 1969.

350 D. E. KIMMINS

There is one paralectotype $.

The selection of a type-specimen for this species has been something of a problem, which
has been made more difficult by the fact that Westwood himself did not publish a description
of the species, Griffith's account being based upon notes supplied by Westwood.

The first description (1832 : 348) is very brief 'The body is dark brown, the upper wings
pale, with a brown spot in each cell; the anterior margin purplish brown, and with a whitish
fasciae rather beyond the centre: the hinder wings red, with a very broad bluish-black
margin.'

The second description, by Newman (1839 : 33) is more detailed and agrees well with the
two examples in BMNH determined by Westwood (and probably seen by Newman before
they reached the BMNH). One was received from Westwood in exchange in 1841 and the
other from the Entomological Club in 1844. The chief differences in the two descriptions
are that Griffiths (1832) does not mention the conspicuous reddish streak along the radius in
the fore wing, and the specimens do not show a pale fore wing with a brown spot in each cell.
They are in fact brownish with a purple tinge, the cross-veins finely bordered with white.
It is possible that Griffiths meant that the cells were almost entirely filled with brownish.

Tillyard based his interpretation of the species on these two examples, which were long
ago labelled as syntypes, and I have therefore made the above lectotype selection, which is
in conformity with current usage.

spio Newman (Chloroperla), 1839 : 86. Holotype <$. Sierra Leone / Chloroperla Spio
Newman, Mag. Nat. Hist. n.s. Ill p. 86 [with on reverse], presented by the Revd. D. F.
Morgan.

Currently placed in the genus Neoperla.

subarmata Despax (ssp. of Chloroperla grammatica), 1924 : 354-357, pi. 8, fig. 22, text-
figs 7-9. Holotype <J (in 2% formaldehyde solution and as microscope preparation).
Angl[eterre], Capel Curig [vi.[ig]ig] (M. E. Mosely).

The locality 'Capel Curig' is in fact in the Welsh county of Caernarvonshire. Currently
placed in the genus Isoperla.

suffusa Walker (Perla), 1852 : 154. LECTOTYPE $. Walker type label / Nepal, with on
reverse 'Hardwicke Bequest' / Perla suffusa Walker, $ Lectotype, D. E. Kimmins det. 1969.

The type-series consists of two $ syntypes, both from Nepal. Walker's description agrees
fairly well, although one venational character agrees only in one out of four fore wings. This
is the number of cross-veins in the terminal areolet (space between costa and RI beyond the
termination of Sc), which is said to be seven. The numbers are actually 4, 5, 5 and 7. I
have selected as Lectotype the example which has seven cross-veins in the terminal areolet
of one fore wing. The second example becomes a paralectotype.

Currently placed as the type-species of Brahmana Klapalek, 1916.

sulcicollis Stephens (Nemoura), 1836 : 143. Syntype [?sex]. Stephens Coll. / Nemoura
sulcicollis Steph., D. E. Kimmins det. 1969.

In the Stephens Collection there is one example (now lacking abdomen) which agrees
reasonably well with the original description, and although it does not carry a specific label
from the Stephens Collection, I am prepared to accept it as at least one of the syntypes.
Stephens' collection of Nemouridae was studied by Morton (1894 : 571) and sulcicollis was
placed by him as a synonym of Nemoura cinerea Olivier, 1811. The latter is a homonym of
Nemoura cinerea (Retzius, 1784) and Brinck (1949 : 37) has used the name Nemoura sulcicollis
Stephens, 1836 as the first available synonym for Nemoura cinerea Olivier, 1811. Currently
placed in the genus Amphinemura.

tasmanica Kimmins (Leptoperla), 1951 : 51-53, figs 5a-e. Holotype <$. Tasmania, Gouldt
County, io.ii[i9]33 (R. J. Tillyard) / Leptoperla tasmanica Kim., $ TYPE.

Holotype mounted as three preparations, two of wings (in de Faure's medium), remainder
in Canada balsam.

TYPE-SPECIMENS OF PLECOPTERA & MEGALOPTERA IN BMNH 351

tasmanica Tillyard (Spaniocercd), 1924 : 195, fig. 3. Holotype $. Tas[mania], Mount
Wellington, 3i.i.[i9]i7 (R. J. Tillyard) / Spaniocerca tasmanica Till., Holotype $, R.J.T.

Holotype (seriously damaged by pests in Tillyard collection) is now represented by two
anterior and one posterior wings.

tenebrosa Klapalek (Kempnyia), 1916 : 69-70. LECTOTYPE $. Theresopolis / 22. ix.
[i8]8y / triangle of black paper / tenebrosa Klapalek / Kempnyia tenebrosa Klap., $ Lecto-
type, D. E. Kimmins det. 1969.

One paralectotype (without abdomen). Currently placed as synonym of Kempnyia klugi
(Pictet).

terminalis Walker (Perla), 1852 : 155-156. LECTOTYPE $. Walker type label / East
Indies / India / 49. PERLA TERMINALIS. / Perla terminalis Walker, $ Lectotype, D. E. Kimmins
det. 1969.

The second example is in less good condition and has been marked paralectotype $.
Currently placed in Marthamea and doubtfully synonymized with M . vitripennis by Klapalek,
1923 : 98.

tibetana Kimmins (Capnia), 1947 : 734-735, figs loA-D. Holotype <$. Tibet, Gaotsa,
12300 ft, 2i.iii.i933 (Raymond Greene), Mt. Everest Exped. / Capnia tibetana Kim., <$ Type.
Type mounted on two microscope slides, in Canada balsam.

tillyardi Kimmins (Spaniocerca), 1951 : 87-88, figs 37a-e. Holotype <J. [Australia],
F[ederal] C[apital] Territory], Lee's Springs, xi.i932 (R. J. Tillyard) / Spaniocerca tillyardi
Kim., <$ TYPE.

Mounted as three preparations, wings dry, head, mouth parts and abdomen, thorax and
legs in canada balsam.

tinctipennis McLachlan (Perla), 1875 : 71. Holotype $. Type [McL. label] / Japan,
Pryer / Perla tinctipennis McL.
Currently placed in Paragnelina.

tragula Kimmins (Amphinemura), 1950 : 189-191, figs 23-25. Holotype <$. Turkestan /
i / Amphinemura tragula Kim., £ TYPE.

Type cleared and mounted in canada balsam.

transmarina Newman (Chloroperla), 1838 : 499-500. Lectotype $ (Ricker, 1938 : 146,
figs 32, 33). Ent. Club / Transmarina Newman, Ent. Mag. ¥.499, N. America, R. Foster /
Chloroperla transmarina Newman, $ Lectotype, W. E. Ricker det. 1935-36.

The data label is now black and the data discernable only with difficulty. Ricker, in effect,
selected this example by his statement 'The $ type, in the British Museum'. The other
examples (implied in Newman's description) have probably been transferred to other species.
Currently placed in the genus Isoperla.

transversa Klapalek (Perlodes), 1912 : 40. Holotype 9- Rheinwald (W. Bennett) / Ent-
[omological] Club / transversa, Klapalek. /
Synonym of Perlodes intricata (Pictet).

tricolor Klapalek (Togoperla), 1921 : 65. Holotype $. Kiu-Kiang / tricolor Klapalek /
Togoperla tricolor Klapalek, $ Holotype, D. E. Kimmins det. 1969.

trijuncta Walker (Perla), 1852 : 153; Ricker, 1938 : 140. Holotype $. Walker type label /
Georgia / 43. PERLA TRIJUNCTA [label cut from Walker Catalogue].
Currently placed as a synonym of Acroneuria arenosa Pictet.

turkestanica Kimmins (Capnia), 1950 : 187-189, figs 14-18. Holotype^. 7 / 517 / Turkestan
/ Capnia turkestanica Kim., D. E. Kimmins det. 1948.
Wings mounted dry, abdomen in canada balsam.

uncata Kimmins (Filchneria), 1947 : 737-739, figs 12, I3A-E. Holotype^. Tibet, Yatung,
10000 ft, 3.^.1924 (R. W. G. Hingston) Everest Exped.

Type (J cleared and mounted in canada balsam, on two slides.

352 D. E. KIMMINS

uniformis Kimmins (Dinotoperla), 1951 : 68-69, figs i9a-g. Holotype <$. Australia, N[ew]
S[outh] W[ales], Rule's Point, 4450 ft, 3o.xii.[i9]34 [R. J. Tillyard] / Dinotoperla uniformis
Kim., <$ TYPE.

Mounted as three preparations, wings in de Faure's medium, remainder in canada balsam.

varia Kimmins (Leptoperla), 1951 : 54-56, figs ya-d. Holotype <$. Tasmania, L[ake] St
Clair, 6.ii.i933 (R. J. Tillyard) / Leptoperla varia Kim., $ TYPE.

Mounted as two preparations, wings in de Faure's medium, remainder in canada balsam.

variegata Stephens (Nemoura), 1836 : 144. Holotype $. Stephens Coll. / variegatus /
Nemoura variegata Steph., $ Holotype, D. E. Kimmins det. 1969. / Brachyptera risi Morton,
$, D. E. Kimmins det. 1969.

Nemoura variegata Stephens is a homonym of Nemoura variegata (Olivier) and takes the
name of the first available synonym, Brachyptera risi Morton.

venosa Kimmins (Neoperla), 1950 : 183-184, figs 7-8. Holotype <$. S. India, Kodaikanal,
7000 ft, 3i.iii.[i9]36, BM-CM Exped. to S. India, 1936 / Neoperla venosa Kim., det. D. E.
Kimmins.

Wings mounted dry, <J genitalia in canada balsam.

[venosa Stephens (Chloroperla), 1836 : 139. Type not recognizable from description.]

vernalis Newport (Capnia), 1848 : 388; 1851 :45i; Walker, 1851 : 176; Ricker, 1938 :
135-136, figs 10-12. Lectotype <J (Ricker, 1938). Hudson's Bay, [Albany River, St
Martin's Falls] / Originally mounted with female / Capnia vernalis Newp., 5" Lectotype,
W. E. Ricker, 1938.

The lectotype has wings mounted dry and abdomen in canada balsam.

xanthenes Newman (Perla), 1838 : 478. Lectotype $ (Needham & Claassen, 1925 : 194).
Georgia / Perla xanthenes Newman $ Type, det. D. E. Kimmins.

Currently placed in Acroneuria. The <J of xanthenes Newman is placed as a synonym of
Perla kansensis Banks.

MEGALOPTERA

affinis Stephens (Raphidia), 1836 : 131. LECTOTYPE $. Stephens Coll. / affinis / Raphidia
afnnis Steph., $ Lectotype, D. E. Kimmins det. 1969.

Currently placed as a synonym of Raphidia (Atlantoraphidia] maculicollis Stephens. A
second labelled example appears to be conspecific and has been labelled 9 paralectotype.

albipennis Walker (Hermes), 1853 : 207-208; Kimmins, 1949 : 766-768, text-fig, i. LECTO-
TYPE $. Walker type label / Nepal, [with on reverse] Hardwicke Bequest / H. albipennis,
Nepaul / Hermes albipennis Walker, $ Lectotype, D. E. Kimmins det. 1969.

This is the example referred to as '$ holotype' in Kimmins, 1949 : 767. Currently placed
in Protohermes.

anticus Walker (Hermes), 1853 : 205. Holotype $. Walker type label / China / H. anticus,
China / Hermes anticus Walker, Holotype, D. E. Kimmins det. 1969.

Weele (1909 : 36) states that the type is a mature $; the abdomen is now missing.
Currently placed as a synonym of Neochauliodes sinensis (Walker).

assamensis Kimmins (Protohermes), 1948 : 773-775, text-figs 11-13. Holotype $. Assam
Protohermes assamensis Kim., $ Type, det D. E. Kimmins.
Abdomen cleared and mounted in canada balsam.

assimilis Albarda (Raphidia), 1891 : 144-146, pi. 8, fig. 23. Holotype $. Type [McL.

label] / 47 / 61 / Vancouver I[sland] (Mat hew) / assimilis Alb. / H. & U. Aspock vid. 1968.

Abdomen cleared and preserved in glycerine. Currently placed in the genus Agulla.

auritus Kimmins (Platyneuromus), 1928 : 369-370, figs 5-7. Holotype <$. Honduras /
Platyneuromus auritus Kimmins, <$ Type, det. D. E. Kimmins / Doeringia auritus Kimmins,
det. D. E. Kimmins, 1932.

TYPE-SPECIMENS OF PLECOPTERA & MEGALOPTERA IN BMNH 353

australica Kimmins (Archichauliodes), 1954 : 424~425. fig- 6- Holotype $. N[ew] S[outh]

W[ales], Upper Murrumbidgee R., Rule's Point, 4500 ft, 2O.xii.i934 (R. J. Tillyard) / Abdomen

in slide cabinet / Archichauliodes australica ^ Kim., D. E. Kimmins det.

It may be pointed out that in the published data of the holotype, the sex is incorrectly

given as $.

australiensis Tillyard (Stenosialis), 1919 : 824. Holotype $. Queensland], Mt Tam-
bourine, I7.xii.[i9]i6 (W. H. Davis) / Stenosialis tambourinensis Till., TYPE, R.J.T. / Steno-

sialis australiensis Till., <$ TYPE, det. D. E. Kimmins.

The type is very much compressed laterally. Tillyard evidently changed his mind about

the specific name, but omitted to replace the determination label. Currently placed in

Austrosialis.
batesi McLachlan (Cordyalis), 1868 : 232-233, pi. 8, fig. i. Holotype ?. Ega (Bates) /

Corydalis batesii McL. (Type).

Generic name now spelt Corydalus.
bellulus Banks (Protohermes), 1931 : 412-413, fig. 18. LECTOTYPE <$. B[ritish] N[orth]

Borneo, Mt Kinabalu, Lumu Lumu, 5500 ft, 8.iv.i929 (H. M. Pendlebury) / Ex F.M.S.

Museums / Protohermes bellulus Bks, type / Protohermes bellulus Banks, <$ Lectotype, D. E.

Kimmins det. 1969.
bicolor Albarda (Raphidia), 1891 : 152-154, pi. 9, fig. 22. LECTOTYPE <$. Type [McL.

label] / Colorado / 42 / bicolor Alb. / Agulla bicolor Alb. <$ Type, det. D. E. Kimmins / Raphidia

bicolor Alb., $ Lectotype, D. E. Kimmins det. 1969.

Apex of abdomen cleared and mounted in canada balsam. There are 3 <$ and 6 $ para-

lectotypes in BMNH. Currently placed in Agulla.
biconicus Kimmins (Protochauliodes), 1954 : 443-444, fig. 22. Holotype £. [Australia],

N[ew] S[outh] Wales, Nowra, 8.x. 1928 (F. A. Rodway) / Abdomen in slide cabinet / Proto-
chauliodes biconicus Kim., £, D. E. Kimmins det. 1953.
bowringi McLachlan (Chauliodes), 1867 : 260 (= Hermes sinensis Walker, 1852 : 203, nee

1852 : 199). Holotype $. Hongkong / H. sinensis, China / Chauliodes bowringi McL.,

Type, D. E. Kimmins det.

Currently placed in Neochauliodes.
burmana Aspock (Inocellia), 1968 : 63; 19680 : 188, fig. 3. Holotype $. N. E. Burma,

Hpungan, iy.iii.i934 (R- Malaise) / Inocellia burmana Aspock & Aspock, 1968, Holotypus.

Abdomen cleared and preserved in glycerine. The first reference is a nomen nudum.
californicus Walker (Chauliodes), 1853 : 199. Holotype ?. Walker type label / California

[From M. Hartweg's collection] / C. californicus, California / Abdomen in slide cabinet /

Chauliodes californicus Walker, <$ Holotype, D. E. Kimmins det. 1969.

Currently placed in the genus Neohermes.
chilensis Kimmins (Archichauliodes), 1954 : 425~427. ngs 7> 8. Holotype <J. Chili

(Culvert) I Abdomen in slide cabinet / Archichauliodes chilensis Kim., <$, D. E. Kimmins det.

1953-
chilensis McLachlan (Sialis), 1870 : 145-146. Holotype <J. Type [McL. label] / Chili /

Sialis chilensis McL.

Currently placed in the genus Protosialis. Weele (1910 : 77), from a photograph of

McLachlan's type, suggests it is a female, because the antennae did not appear to be hairy.

They are in fact pilose and the type is definitely a male.
continentalis Weele (Parachauliodes), 1909 : 259; 1910 : 60, fig. 47, pi. 4, fig. 30; Kimmins,

1954 : 432. text-fig. 12. LECTOTYPE <J. [Korea], Ssu-shima, 27^.1891, H.t. / Meta-

chauliodes continentalis Weele, type / Parachauliodes continentalis Weele, <$ Lectotype,

D. E. Kimmins det. 1969.

The generic name Metachauliodes is a nomen nudum and was evidently replaced at the last

minute by Weele with Parachauliodes, the same name occurring twice (Weele, 1910 : 61) in

error for Parachauliodes. There is one $ paralectotype in BMNH.

354 D- E. KIMMINS

corripiens Walker (Hermes), 1860 : 180. LECTOTYPE ?. [No locality label] / Walker
type label / Saunders [Coll.] / corripiens / Hermes corripiens Walker, $ Lectotype, D. E.
Kimmins det. 1969.

Walker's measurements suggest that he had more than one example; and there is a second
example from Brazil, also from the Saunders collection, which may be one of the type-series.
It lacks the anterior spots on pronotum, which Walker says may be obsolete. Both speci-
mens are a little larger than the dimensions given by Walker.

costalis Walker (Hermes), 1853 : 207; Kimmins, 1949 : 775-778, figs 14-18. LECTOTYPE
cj. Walker type label / North China / H. costalis, N. China / Hermes costalis Walker, 3
Lectotype, D. E. Kimmins det. 1969.

Lectotype with abdomen cleared and mounted in canada balsam. The second example
has been labelled paralectotype $. Currently placed in the genus Protohermes.
crassicornis McLachlan (Corydalis), 1867 : 233-235, pi. 8, fig. 2. Holotype (J. Texas /
Corydalis crassicornis McL. (Type).

Generic name now spelt Corydalus.

deceptor Kimmins (Archichauliodes), 1954 : 423-424, fig. 5. Holotype <$. Queensland,
Toowoomba, 2000 ft, lo.xii, 1884 / Abdomen in slide cabinet Chauliodes guttifer McL.
[McL. det.] /Archichauliodes deceptor Kim., <$ D. E. Kimmins det. 1953.

decimmaculatus Walker (Hermes), 1860 : 180; Kimmins, 1949 : 768-770, figs 2-4. Holotype
cj. [No locality data] / Saunders [coll.] / lo-maculatus W. / Hermes decimmaculatus Walker,
$ Holotype, D. E. Kimmins det. 1969.
Abdomen mounted in canada balsam.

disjunctus Walker (Chauliodes), 1866 : 334. Holotype $. Walker type label / British]
Columbia] / Brit[ish] Columbia], Chalukweyuh Lake, ix.i859 / Chauliodes disjuncta.
Currently placed in the genus Dysmicohermes.

diversus Walker (Hermes), 1853 : 205; Kimmins, 19380 : 354-358, pi. I3A, text-figs i, 2, 5,
6, 9, 10 [designation of type]. Lectotype <$ (Kimmins, 1938). Walker type label / N[ew]
Zeal[an]d / Hermes diversus Walker <J, selected as type by D. E. Kimmins, Feb. 1938.

The type has the abdomen cleared and mounted in Canada balsam. Currently placed in
the genus Archichauliodes.

dubitatus Walker (Hermes), 1854 : 204; Kimmins, 19380 : 354-358, pi. 13, text-figs 3, 4, 7, 8.
Holotype $. Walker type label / 26 / H. dubitatus [no locality] / Abdomen in slide cabinet /
Protochauliodes dubitatus Walker, $ Type, det. D. E. Kimmins.

Locality possibly South American. Currently placed in the genus Protochauliodes.

esbenpeterseni Kimmins (Leptochauliodes), 1930 : 663-665, 2 figs. Holotype <J. C[ape]
C[olony], Hott[entots] Holl[and] Mts, 4000 ft, Caledon, 1916 ([K. H. Barnard) / Abdomen in
slide cabinet / Leptochauliodes esbenpeterseni Kimmins <$, det. D. E. Kimmins, 1930 /
Taeniochauliodes ochraceopennis Esb.-Pet. <$, det. D. E. Kimmins.

fasciatus Walker (Chauliodes), 1853 : 201. LECTOTYPE. Walker type label / 'N[ew]
Holl[and]' / C. fasciatus, Australia / Chauliodes fasciatus Walker, $ Lectotype, D. E. Kimmins
det. 1969.

Of the two original examples, the above mentioned specimen is the only one bearing
locality and determination labels. There is a smaller example, without any data whatever,
but I feel that it would be unwise to mark it as a paralectotype.

fenestralis McLachlan (Neuromus), 1869 : 42-43. LECTOTYPE <$. Dhargeeling / Neuro-
mus fenestralis McL. / Neuromus fenestralis fenestralis McL., <J Lectotype, D. E. Kimmins
det. 1969.

The second $ syntype has been labelled paralectotype. Currently placed in the genus

Neoneuromus.

ferrugineus Walker (Sialis), 1853 : 195. Holotype <$. Walker type label / Georgia [From
Mr Abbot's collection] / S. ferrugineus N. Amer[ica] / Sialis ferrugineus Walk., $ Holotype,

TYPE-SPECIMENS OF PLECOPTERA & MEGALOPTERA IN BMNH 355

D. E. Kimmins det. 1969.

Currently placed as a synonym of Sialis americana (Rambur).
fletcheri Kimmins (Neochauliodes), 1954 : 434» fig- :3- Holotype <$. India, Assam,

Shillong, 5000 ft, 26.vi.-io.vii.i928 (T. Bainbrigge Fletcher) / Abdomen in slide cabinet /

Neochauliodes fletcheri Kim., <$, D. E. Kimmins det. 1953.
forcipatus Kimmins (Ctenochauliod.es), 1954 : 429~43°. fig- I][- Holotype $. China,

[Szechwan?], Kwanshien, vii.i93o (G. M. Franck) / Abdomen in slide cabinet / Ctenochauliodes

forcipatus Kim., <J, D. E. Kimmins det. 1953.

fraternus McLachlan (Chauliodes), 1869 : 37. Holotype 9- N. China / Chauliodes fraternus
McL.

Currently placed as ssp. of Neochauliodes sinensis (Walker).

fulvostigmata Aspock (Inocellia), 1968 : 63; 19680 : 184-187, figs 1-2. Holotype Q\ n.viii.
/ Kashmir, Gulmarg, n.viii.[i9]3i (T. B. Fletcher) / Inocellia fulvostigmata Aspock et
Aspock, 1968, Holotypus.

The first reference is a nomen nudum.

Juscinata Aspock (Inocellia), 1964 : 62; 1965 : 352-353, fig. 22. Holotype <$. Turkey,
Amasya, 9.vi.iQ59 (K. M. Guichard) / Raphidia fuscinata H. et U. Aspock, Holotypus <J.

Currently placed in the subgenus Turcoraphidia. Abdomen of holotype cleared and pre-
served in glycerine.

guttiferus Walker (Hermes), 1853 : 204; Kimmins, 1954 : 419-421, figs 2, 3. Holotype
[?sex]. Walker type label / [locality unknown] / guttifera / Hermes guttifera Walker,
holotype, D. E. Kimmins det. 1969.

Currently placed in the genus Archichauliodes.

hageni Albarda (Inocellia), 1891 : 171-172, pi. n, fig. 33. Holotype 9- Type [McL. label] /
57 / [California], S[an] Francisco (Edwards) / Hageni, Alb.

Hecate McLachlan (Corydalis), 1866 : 1-2, pi. 20. LECTOTYPE <$. Brazil / Corydalis
hecate McL., Type / Corydalis cephalotes Rbr [McL. writing] / Corydalis hecate McL., <$
Lectotype, D. E. Kimmins det. 1969.

Currently placed as a synonym of C. cephalotes Rambur.

ignicollis Tillyard (Austrosialls), 1919 : 823. Holotype <$. [Tasmania], Maria Isl[and],
29xii.[i9]i3 / Austrosialis ignicollis Till. Type, R.J.T.

inamabilis McLachlan (Corydalis), 1867 : 235-236, pi. 8, fig. 3. Holotype <J. Texas /
Corydalis inamabilis McL. (Type).

Generic name currently spelt Corydalus.

indecisus Walker (Hermes), 1953 : 203-204. Holotype 9- Walker type label / indecisa /
H. indecisus / Hermes indecisus Walker, 9 Holotype, D. E. Kimmins det. 1969.

Currently placed as a synonym of Chauliodes rastricornis (Rambur), a North American
species.

infectus McLachlan (Neuromus), 1869 : 41; Weele, 1910 : 37 [type restricted to BMNH
example]. Lectotype <?. [India], Dhargeling / Neuromus infectus McL.

Weele, 1910, restricted the type-series to the <J from Darjeeling, in BMNH and in effect
designated it as lectotype.

infumata Newman (Sialis), 1838 : 500. Holotype 9- [Locality and determination label
blackened and almost illegible], Sialis Latr., infumata Newm., [North America, Trenton Falls
(Doubleday)] Ent. Club / Sialis infumata Newm., 9 Holotype, D. E. Kimmins det. 1969.
Left hind wing missing, part of left fore wing mounted dry, abdomen in Canada balsam.

intimus McLachlan (Neuromus), 1869 : 44-45. LECTOTYPE <J. Ind[ia] / Saunders
[Coll.] / intimus McL. type [W. F. Kirby's writing] / Neuromus intimus McL., <$ Lectotype,
D. E. Kimmins det. 1969.

McLachlan stated that the type locality was 'India orientali', but this appears to be an

356 D. E. KIMMINS

error, as the two syn types are labelled 'Ind[ia]' and 'Saunders [Coll.]'. Weele (1910 : 29) is
also wrong in stating that the 'type was labelled "East Indies" ', although he correctly quotes
the other syntype as 'India (Collection Saunders)'. Had he quoted the type-data correctly,
I would have accepted his statement as a lectotype designation.

japonicus McLachlan (Chauliodes), 1867 : 232. Holotype $. Japan / Chauliodes japonicus
McL. (Type).

Currently placed in Parachauliodes.

kitntninsi Aspock (Raphidia), 1964 : 62; 1965 : 315-317, fig. 2. Holotype <$. Turkey,
Amasya, 1400 ft, 9.vi.i959 (K. M. Guichard) / Raphidia kimminsi H. & U. Aspock, 1964,
Holotypus (J.

koreanus Weele (Neochauliodes), 1909 : 261; Kimmins, 1954 : 437~438, fig. 16 [Lectotype
designation, as 'holotype'j. Lectotype <$. Korea, Seoul, viii (E. Scarlett) / Neochauliodes
koreanus Weele, type / Abdomen in slide cabinet / Neochauliodes koreanus Weele, <£ Lecto-
type, D. E. Kimmins det. 1969.

I regard my designation of '<$ holotype' as equivalent to lectotype.

latratus McLachlan (Neuromus), 1869 : 43-44. Holotype <J. E[ast] Ind[ies] / latratus,
McL. type [W. F. Kirby's writing].

Although labelled 'East Indies', our museum register states that most of this collection was
from India. Currently placed in Neoneuromus.

londinensis Stephens (Raphidia), 1836 : 130. LECTOTYPE ?. Stephens Coll. / Raphidia
londinensis Steph., Lectotype $, D. E. Kimmins det. 1969.

Currently placed as a synonym of Raphidia (Raphidilla) xanthostigma Schummel.

longicornis Albarda (Inocellia), 1891 : 169-170, pi. n, fig. 32. Holotype <$. Type [McL.
label] / 58 / California (Walsingham) / Longicornis Alb. / Inocellia longicornis Alb., <$ Type,
D. E. Kimmins det. 1969.

maclachlani Albarda (Inocellia), 1891 : 162-164, pi- IO. fig- 29- LECTOTYPE <J.
[Corsica], Porto Vecchio / 100 / Inocellia mclachlani Alb., det. H. Albarda / Inocellia
maclachlani Alb., <$ Lectotype, D. E. Kimmins det. 1969.

Abdomen of lectotype cleared and preserved in glycerine. There are i $, 2 $ paralectotypes
in BMNH, the females being from Sardinia. Currently placed in the genus Fibla.

maculicollis Stephens (Raphidia), 1836 : 131. LECTOTYPE $. Stephens Coll. / maculi-
collis / Raphidia maculicollis Steph., $ Lectotype, D. E. Kimmins det. 1969.

There is a second example $, also labelled maculicollis. Currently placed as Raphidia
(Atlantoraphidia) maculicollis Stephens.

tnaculifera Walker (Hermes), 1853 : 203. Holotype $. Walker type label / Malabar,
[with on reverse] 'Ent. Club' / H. maculifera, Malabar / Hermes maculifera Walker, $ Holotype,
D. E. Kimmins det. 1969.

maculipennis Gray (Hermes), in Griffiths et al., 1832 : 331. Holotype (J. H. maculipennis
(ruficollis Ramb.), Java.

According to the BMNH Register, this specimen was purchased in 1840, at Mr Children's
sale.

megacephala Stephens (Raphidia), 1836 : 130. LECTOTYPE $. Stephens Coll. / mega-
cephala / Raphidia megacephala Stephens, $ Lectotype, D. E. Kimmins det. 1969.

The head and prothorax of this example had become detached and wrongly gummed on
to the metathorax of a specimen labelled 'Londinensis', with which it was an obvious misfit.
Currently placed as a synonym of Raphidia (Navasana) notata Fabricius. A second example,
labelled 'megacephala', is R. cognata.

montanus McLachlan (Neuromus), 1869 : 42; Kimmins, 1949 : 780-781, fig. 23. Holotype
$. Sikkim, Lacken, 9000 ft, / N. Ind[ia] / Neuromus montanus McL.

The type has badly battered wings; abdomen cleared and mounted in canada balsam.

TYPE-SPECIMENS OF PLECOPTERA & MEGALOPTERA IN BMNH 357

nigricollis Albarda (Raphidia), 1891 : 142-144, pi. 8, fig. 22. LECTOTYPE <J. Frankf[urt]
a[m] M[ain] / 58 / Raphidia nigricollis Alb., det. H. Albarda / Raphidia nigricollis Albarda,
<$ Lectotype, D. E. Kimmins det. 1969.

In addition, there are 2 <J and 3 § paralectotypes in BMNH. Currently placed in Raphidia

( V enustoraphidia) .

obscurus Weele (Neochauliodes), 1909 : 262-263. Holotype ?. Presented] by Dr Watts,
Manipur / Neochauliodes obscurus Weele, 1908, type.

occidentalis Weele (ssp. Neochauliodes sinensis), 1909 : 266; Kimmins, 1954 : 438-439,
figs lya-c [Lectotype designation (as 'holotype)]. Lectotype $. [W. China], Omei-Shan /
Neochauliodes sinensis Wlk., occidentalis Weele Type / Abdomen in slide cabinet / Neochau-
liodes sinensis occidentalis Weele, <J Lectotype, D. E. Kimmins det. 1969.

I consider that my designation of a '<$ holotype' in 1954 as equivalent to lectotype.

orientalis McLachlan (Corydalis), 1899 : 281-283, pi- 9- Holotype <$. W. China, Chia-ting-
fu, [1000 ft, May] / Corydalis orientalis McL., Type. /
Currently placed in Acanthacorydalis.

plomleyi Kimmins (Archichauliodes), 1954 : 422-423, fig- 4- Holotype $. [Australia,
New South Wales], Mt Irvine, [2300 ft], 25.xii.i934 (N. J. Plomley) / Abdomen in slide
cabinet / Archichauliodes plomleyi Kim., D. E. Kimmins det. 1953.
One pair of wings mounted dry, abdomen in Canada balsam.

pontica Albarda (Raphidia), 1891 : 102-104, P1- 4. fig- 6- LECTOTYPE £. 41 / [Asia
Minor, Siwas], Amasia / Raphidia pontica Alb. / Raphidia pontica <J Lectotype, D. E. Kimmins
det. 1969.

cj abdomen cleared and preserved in glycerine. Currently placed in the genus Agulla.
The $ paralectotype lacks head and pronotum. Albarda also lists an example without
abdomen in the Berlin Museum and two females in Geneva.

pusillus McLachlan (Chauliodes), 1867 : 231-232. Holotype $. [locality unknown to
McLachlan, S. Africa]. Chauliodes pusillus McL. (Type).
Currently placed in Platychauliodes.

reedi Kimmins (Protochauliodes), 1954 : 44°~443> n§s 20> 2I- Holotype $. Chili (Reed) /
Abdomen in slide cabinet / Protochauliodes reedi Kim., <J, D. E. Kimmins det. 1953.
One pair of wings mounted dry, abdomen in Canada balsam.

[selysi Weele (Hermes), 1909 : 256; 1910 : 89.

The BMNH has one example (lacking abdomen), which is labelled 'Type' by Weele, but in
1910 : 89 he indicates (by an asterisk in the specific list) that the 'type' [strictly lectotype] is
in the Selys Collection. This is the damaged <$ from Sylhet. Our example is therefore a
paralectotype.]

sibirica McLachlan (Sialis), 1872 : 55-56, pi. i, figs 10, loa. LECTOTYPE 3. Type [McL.
label] / Siber[ie] Orient. (Maack) / Sialis sibirica McL. / Sialis sibirica McL., <$ Lectotype,
D. E. Kimmins det. 1969.

There are two <$ and three $ paralectotypes.

simplex Walker (Chauliodes), 1853 : 200. LECTOTYPE #. Walker type label / Silhet
[Mr Argent's collection] / simplex / Chauliodes simplex Walker, <$ Lectotype, D. E. Kimmins
det. 1969.

One of the two paralectotypes, which carries Weele's determination label, was the specimen
figured by him in 1909 and by Kimmins, 1954 : 433> figs I3a-c.

sinensis Walker (Chauliodes), 1853 : 199. LECTOTYPE $. North China / C. Sinensis,
China / Abdomen in slide cabinet / Chauliodes sinensis Walker, <J Lectotype, D. E. Kimmins
det. 1969.

The second example, from China, has not been traced in BMNH.

sinensis Walker (Hermes), 1853 : 203. See bowringi McLachlan (Chauliodes).

358 D. E. KIMMINS

subfasciatus Westwood (Chauliodes), 1848 : 70, pi. 34, fig. 5. LECTOTYPE <?. Sylhet /
Saunders [Coll.] / Chauliodes subfasciatus, West. Cab. or. Ent., pi. 34, fig. 5 / Chauliodes
subfasciatus Westwood, <J Lectotype, D. E. Kimmins det. 1969.

Both the lectotype and paralectotype males lack abdomens. Currently placed in
Neochauliodes .

subnubilus Kimmins (Protohertnes), 1949 : 770-771, figs 5-7. Holotype <$. Birmah,
Ruby Mines / Protohermes subnubilus Kim., $ TYPE, det. D. E. Kimmins.

Apex of abdomen mounted in Canada balsam.

tennis McLachlan (Chauliodes), 1869 : 38. Holotype $. S[outh] Afr[ica], Knysna, / C.
tenuis type [W. F. Kirby's writing] / Chauliodes tenuis McL., S. Afr.

The holotype now lacks the right fore wing and the left hind wing. Currently placed in
Platy chauliodes.

tonkinicus Kimmins (Anachauliodes), 1954 : 428-429, figs 9, 10. Holotype <$. Tonkin,
Ngai-Tio, 22. iv. 1924 (H. Stevens) / Abdomen in slide cabinet / Anachauliodes tonkinicus Kim.,
cj, D. E. Kimmins det. 1953.

One pair of wings mounted dry, abdomen of holotype <$ in Canada balsam.
truncatus Kimmins (ssp. of Neochauliodes sinensis), 1954 : 440, fig. 19. Holotype <$.
[Assam], Khasis, Shillong, 5000 ft, 15. vi. 1928 (7". Bainbrigge Fletcher) / Abdomen in slide
cabinet / Neochauliodes sinensis truncatus <$ Kim., D. E. Kimmins det. 1953.

umbratus Kimmins (Neochauliodes), 1954 : 436> ng- T5- Holotype <$. Indo-China (A.

Vuillet) I Abdomen in slide cabinet / Neochauliodes umbratus Kim., <$, D. E. Kimmins det.

1953-
unifortnis Banks (Protohermes), 1931 : 412, figs 19, 20. Holotype $. B[ritish] N[orth]

Borneo, Mt Kinabalu, Lumu Lumu, 5500 ft, 9.^.1929 (H. M. Pendlebury) / Ex F.M.S.

Museums / Protohermes uniformis Bks, type.

REFERENCES

ALBARDA, H. 1891. Revision des Rhaphidides. Tijdschr. Ent. 34 : 65-184, pis. 2-11.
ASPOCK, H. & U. 1964. Zwei weitere neue Arten des Genus Raphidia L. (Neuroptera) aus

Kleinasien (Vorlaufige Beschreibung) . Ent. Nachrbl., Wien 11 (8) : 62.

— 1965. Zur Kenntnis der Raphidiiden von Siidosteuropa und Kleinasien (Mit kritischen
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- 1968. Vorlaufige Mitteilung zur generischen Klassifizierung der Raphidiodea (Insecta,
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— 19680. Zwei neue Inocelliiden-Spezies aus Asien. Ent. Ber., Amst. 28 : 184-189, 3 figs.
AUBERT, J. 1950. Note sur les Plecopteres europeens du genre Taeniopteryx Pictet

(Nephelopteryx Klapalek) et sur Capnia vidua Klapalek. Mitt, schweiz. ent. Ges. 23 :
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BANKS, N. 1931. Neuropteroid Insects from the Malay Peninsula. /. fed. Malay St. Mus.

16 : 377-409, 40 figs.

- 19310. Neuropteroid Insects from North Borneo, particularly from Mt Kinabalu.
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— 1938. Further Neuropteroid Insects from Malaya. /. fed. Malay St. Mus. 18 : 220-235,

17 figs.

BARNARD, K. H. 1934. South African Stone-flies (Perlaria), with descriptions of new species.

Ann. S. Afr. Mus. 30 : 511-548, 21 text-figs.

BRINCK, P. 1949. Studies on Swedish Stoneflies. Opusc. ent. Suppl. 11 : 1-250, 61 text-figs.
DESPAX, R. 1934. Plecopteres pyrendens. VIII. Etude et description de quelques formes

de Nemoures apparentees a Nemura marginata (Pict.) Klap. Bull. Soc. Hist. nat. Toulouse

66 : 255-270, 12 text-figs.

TYPE-SPECIMENS OF PLECOPTERA & MEGALOPTERA IN BMNH 359

GRIFFITH, E. et al. 1832. The Animal Kingdom arranged in conformity with its organization,
by the Baron Cuvier, with supplementary additions to each order by Edward Griffith, F.L.S.,
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London.

KIMMINS, D. E. 1928. New and little known Neuroptera of Central America. Eos, Madr.
4 : 363-370, 7 text-figs.

— 1935. A new brachypterous Isopteryx from France. Ann. Mag. nat. Hist. (10) 15 : 645-
650, 8 text-figs.

- 1938. Notes on the Plecoptera of New Zealand, with descriptions of new species. Ann.
Mag. nat. Hist, (n) 2 : 561-580, 15 text-figs.

- 19380. Hermes dubitatus and Hermes diver sus Walker (Sialidae, Megaloptera) . Ann.
Mag. nat. Hist, (n) 2 : 354-358, pi. 13, 10 text-figs.

- 1941. A new species of Nemouridae (Plecoptera). /. Soc. Br. Ent. 2 (3) : 89-93, ngs

I3-I4-

- 1943. Rhabdiopteryx anglica, a new British species of Plecoptera. Proc. R. ent. Soc.
Lond. (B) 12 : 42-44, 8 figs.

1947. New species of Himalayan Plecoptera. Ann. Mag. nat. Hist, (n) 13 : 721-740,

14 text-figs. [Issue dated November 1946, but actual date of publication 13 Oct. 1947,
due to post-war publication delays.]

1948. A new species of Anacroneuria (Plecoptera Perlidae) from Trinidad. Proc. R. ent.

Soc. Lond. (B) 17 : 105-106, 5 text-figs.

— 1949. Notes on the genus Protohermes Weele (Megaloptera), with descriptions of two new
species. Ann. Mag. nat. Hist. (12) 1 : 765-781, 23 text-figs. [This is the Nov. 1948 issue,
which was actually published in March 1949.]

- 1950. Some new species of Asiatic Plecoptera. Ann. Mag. nat. Hist. (12) 3 : 177-192,
26 text-figs.

- 19500. Some Assamese Plecoptera, with descriptions of new species. Ann. Mag. nat.
Hist. (12) 3 : 194-209, 17 text-figs.

- 1951. A revision of the Australian & Tasmanian Gripopterygidae and Nemouridae
(Plecoptera). Bull. Br. Mus. nat. Hist. (Ent.) 2 (2) : 43-93, 40 text-figs.

- 1954. A new genus and some new species of the Chauliodini (Megaloptera). Bull. Br.
Mus. nat. Hist. (Ent.) 3 (u) : 415-444, 22 text-figs.

KLAPALEK, F. 1909. Vorlaufiger Bericht liber exotische Plecoptera. Wien. ent. Z. 28 : 215-
232.

— 1912. Perlodidae. Collns zool. Selys-Longchamps, IV (i) : 1-66, 58 text-figs.

— 19120. Plecopterum Genus : Kamimuria Kip. Cas. ceske Spol. ent. 9 (2) : 103-110.

— 1916. Subfamilia Acroneurinae Kip. Cas. ceske Spol. ent. 13 : 45-84.

- 1921. Plecopteres Nouveaux. I. Sousfamille des Perlinae et Neoperlinae (Travail
posthume). Annls Soc. ent. Belg. 61 : 57-67, 320-327.

- 1922. Plecopteres Nouveaux (Travail posthume). Quatrieme partie. Annls. Soc. ent.
Belg 62 : 89-95.

— 1923. Fam. Perlidae. Collns zool. Selys-Longchamps IV (2) : 1-193, 61 text-figs.
LORD, J. K. 1866. The Naturalist in Vancouver Island and British Columbia 2. London.
MCLACHLAN, R. 1866. Descriptions of a new neuropterous insect belonging to the genus

Corydalis Latreille. /. ent. London 14 : 1—2, pi. 20.

- 1867. New Genera and species, etc., of Neuropterous insects; and a revision of Mr F.
Walker's British Museum Catalogue, part ii (1853), as far as the end of the genus Myrmeleon.
J. Linn. Soc. (Zool.) 9 : 230-281, pi. viii.

— 1869. Considerations on the neuropterous genus Chauliodes and its allies; with notes and
descriptions. Ann. Mag. nat. Hist. (4) 4 : 35-46.

— 1870. On the occurrence of the neuropterous genus Sialis in Chili. Entomologist's mon.
Mag. 7 : 145-146.

- 1872. MateYiaux pour une faune neVropterologique de 1'Asie septentrionale. Seconde
partie, Non-Odonates. Annls Soc. ent. Belg. 15 : 47-77, i pi.

360 D. E. KIMMINS

McLACHLAN, R. 1875. A sketch of our present knowledge of the neuropterous fauna of Japan
(excluding Odonata and Trichoptera) . Trans, ent. Soc. Lond. 1875 : 167-190.

— 1895. A small contribution to a knowledge of the neuropterous fauna of Rhenish Prussia.
Entomologist's mon. Mag. (2) 6 : 109-112.

— 1899. A second asiatic species of Corydalis. Trans, ent. Soc. Lond. 1899 : 281-283, pi. 9.
MENDL, H. 1968. Zur Unterscheidung von zwei Leuctra-A.rten. (L. cingulata Kempny und

L. pseudocingulata nov. nom.). Mitt, schweiz. ent. Ges. 41 : 305-319, 7 text-figs.
MORTON, K. J. 1894. Palaearctic Nemoiirae. Trans, ent. Soc. Lond. 1894 : 557-574, pis.

13-14-

- 1930. Plecoptera collected in Corsica by Mr Martin E. Mosely. Entomologist's mon.
Mag. 66 : 75-81, pi. 2.

MOSELY, M. E. 1930. New European Trichoptera and Plecoptera. Trans, ent. Soc. Lond.
78 : 237-253, pi. 14, 33 text-figs.

— 1932. A revision of the european species of the genus Leuctra (Plecoptera). Ann. Mag.
nat. Hist. (10) 10 : 1-41, 5 pis., 57 text-figs.

— 1935. New Trichoptera and Plecoptera in France : some Auvergne Plecoptera. Trans.
R. ent. Soc. Lond. 83 : 557-562, 8 text-figs.

NEEDHAM, J. G. & CLAASSEN, P. W. 1925. A monograph of the Plecoptera, or Stoneflies of
America north of Mexico. Entomological Society of America, Thomas Say Foundation
2, 397 PP-. 5° Pls-. 29 text-figs.

NEWMAN, E. 1833. Entomological Notes. Ent. Mag. 1 : 413-416.

- 1838. Entomological Notes. Ent. Mag. 5 : 168-181, (372-402), 469-483.

- 1839. On the synonymy of the Perlites, together with brief characteristics of the old,
and a few new species. Mag. nat. Hist. (N. S.) 3 : 32-37, 84-90.

1845. Addendum to 'Synonymy of the Perlites, by Edward Newman', published in the

Mag. Nat. Hist, for 1839. Zoologist 3 : 852-854.

NEWPORT, G. 1848-49. Proc. Linn. Soc. Lond. i : 388-389 (20. vi. 1848) [p. 388 published in
1848, 389 in 1849].

1851. Descriptions of some American Per lidae, together with notes on their habits.

Trans. Linn. Soc. Lond. 20 (3) : 447-452.

RICKER, W. E. 1938. Notes on specimens of American Plecoptera in European collections.
Trans. R. canad. Inst. 22 (i) : 129-156, 38 figs.

SMITH, L. W. 1917. Studies of North American Plecoptera (Pteronarcinae and Perlodini).
Trans, am. ent. Soc. 43 : 433-489, pis. 29-34.

STEPHENS, J. F. 1836-37. Illustrations of British Entomology ; or a Synopsis of Indigenous
Insects; containing their generic and specific distinctions; with an account of their meta-
morphoses, times of appearance, localities, food and economy, as far as practical. Vol. 6.
[Date of publication of Plecoptera and Megaloptera, I5.viii.i836.] London.

TILL YARD, R. J. 1919. Australian Megaloptera or alder-flies. Proc. Linn. Soc. N.S.W.
43 : 819-825.

— 1921. A new classification of the Order Perlaria. Can. Ent. 53 : 35-43, figs 1-4.

- 19210. Revision of the family Eustheniidae (Order Perlaria) with descriptions of new
genera and species. Proc. Linn. Soc. N.S.W. 46 : 221-236, pis xi-xv, 4 text-figs.

- 19216. The wing- venation of the Leptoperlidae (Order Perlaria), with description of a
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4 text-figs.

— 1924. New genera and species of Australian Stone-flies (Order Perlaria). Trans. R. Soc.
S. Austr. 48 : 192-195, 3 text-figs.

WALKER, F. 1852. Catalogue of the specimens of neuropterous insects in the collection of the
British Museum. Part I. 192 pp. London.

- 1860. Characters of undescribed Neuroptera in the Collection of W. W. Saunders, Esq.,
F.R.S., &c. Trans, ent. Soc. Lond. (2) 5 : 176-199.

WEELE, H. W. VAN DER. 1909. New genera and species of Megaloptera. Notes Leyden Mus.
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TYPE-SPECIMENS OF PLECOPTERA & MEGALOPTERA IN BMNH 361

— 1910. Megaloptera. Collns zool. Selys Longchamps 5 (i) : 93 pp., 4 pis., 70 text-figs.
WESTWOOD, J. O. 1848. The Cabinet of Oriental Entomology : being a selection of the rarer and
more beautiful species of insects, natives of India and the adjacent islands, the greater portion
of which are now for the first time described and figured. 88 pp., 42 pis. London.

D. E. KlMMINS

Dept. of Entomology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON, S.W.y

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3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177 :
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9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species
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10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rho-
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A REVISION OF THE
TERMITES OF THE GENUS

MACROTERMES

FROM THE ETHIOPIAN REGION
(ISOPTERA : TERMITIDAE)

J. E. RUELLE

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 24 No. 9

LONDON: 1970

A REVISION OF THE
TERMITES OF THE GENUS

MACROTERMES

FROM THE ETHIOPIAN REGION
(ISOPTERA : TERMITIDAE)

BY

JEAN-EMILE RUELLE

Universite Lovanium

Kinshasa
R6publique D6mocratique du Congo

Pp. 363-444 ; 12 Maps, 152 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol.24 No. 9

LONDON : 1970

THE BULLETIN OF THE BRITISH MUSEUM

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

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

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

This paper is Vol. 24, No. 9, of the Entomological
series. The abbreviated titles of periodicals cited follow
those of the World List of Scientific Periodicals.

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

Trustees of the British Museum (Natural History) 1970

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 24 July 1970 Price £2 I2s.

(£2-60)

A REVISION OF THE
TERMITES OF THE GENUS

MACROTERMES

FROM THE ETHIOPIAN REGION
(ISOPTERA : TERMITIDAE)

By J. E. RUELLE

CONTENTS

Page

INTRODUCTION .......... 365

MATERIAL ........... 366

METHODS ........... 367

MACROTERMES HOLMGREN : DIAGNOSIS AND DISCUSSION. . . 368

DISTRIBUTION .......... 369

KEYS TO SPECIES .......... 370

M. bellicosus (Smeathman) ........ 374

M. falciger (Gerstacker) sp. rev., comb. n. ..... 381

M. herus (Sjostedt) 388

M. ivorensis Grass6 and Noirot . . . . . . . 391

M. lilljeborgi (Sjostedt) 394

M. mossambicus (Hagen) stat. n. . . . . . . . 398

M. muelleri (Sjostedt) ......... 405

M. natalensis (Haviland) ........ 410

M. nobilis (Sjostedt) ......... 415

M. subhyalinus (Rambur) sp. rev. ...... 419

M. ukuzii Fuller stat. n. ........ 428

M. vitrialatus (Sjostedt) ........ 432

ACKNOWLEDGEMENTS ......... 440

REFERENCES .......... 441

SYNOPSIS

The genus Macrotermes, including the genus Bellicositermes of some authors, is revised for the
Ethiopian Zoogeographical Region. Of 23 species and n forms or varieties found in the
literature, the varieties have not been retained ; ten specific names are found to be synonymous
and three more are dubious. Since one variety has been given specific status and one species
has been split into two, this leaves 12 recognized and redescribed species. M. ukuzii and
M. mossambicus have been given specific status (the latter being a senior synonym of M.
michaelseni) , M. falciger is another senior synonym (of M. goliath), M. natalensis has been split
into M. bellicosus and M. natalensis, M. subhyalinus replaces M. bellicosus. Keys are given
to the species ; notes are provided on their distribution and biology.

INTRODUCTION

AMONG the ten genera of the subfamily Macrotermitinae Kemner (1934), Macro-
termes, like Microtermes and Odontotermes, occurs both in the Indo-malayan and the
Ethiopian Zoogeographical Regions, the other genera being endemic to the Ethiopian

366 J. E. RUELLE

Region. One of the latter, Ancistrotermes, was recently revised (Harris, 1966) ;
from the same region, 82 species of Odontotermes and 34 of Microtermes were recorded
(Bouillon & Mathot, 1966). This paper deals with the African Macrotermes, which
are represented by a very abundant material.

Smeathman (1781) described the first known species of this genus (Termes belli-
cosus). Holmgren (1909, 1910) introduced the name Macrotermes as a subgenus of
Termes. Various generic names were attributed to the Ethiopian Macrotermes
from 1910 to 1926 (year of Sjostedt's last revision). The three genera recognized by
Sjostedt, with 22 species altogether, were reduced in Snyder's catalogue (1949) to
one (Macrotermes}, with 23 species. After 1949, two new species were described
(M. angolensis, M. ivorensis) and two synonymized (M. convexus, M. jeanneli] .

Type-specimens, i.e. theholotype, lectotype, syntypes, paratypes or paralectotypes,
where available, have been studied. One type-collection (M. ivorensis) has not been
accessible ; three neotypes have been selected : M. bellicosus, M. subhyalinus and
M. kibonotensis, the latter a junior synonym of M.falciger. In three cases (M. niger,
M. gratus, M. ituriensis ; Sjostedt, 1898, 1900, 19240) the loss of the type-material
and the impossibility of selecting a neotype leave the concerned species as nomina
dubia. The words 'LECTOTYPE' and 'NEOTYPE' in capitals indicate designation
in this paper.

MATERIAL

A total of 2,123 nest series from all parts of Africa have been examined, con-
taining each from one to sometimes more than one hundred insects. The study
started with the material of the British Museum (Natural History) (B.M.(N.H.)),
much of which had been collected by the members of the Termite Research Unit.
The following institutions have also been visited :

Muse'e Royal de 1'Afrique Centrale, Tervuren (Terv.) ; American Museum of
Natural History, New York (A.M.N.H.) ; Institut Royal des Sciences Naturelles,
Bruxelles (I.R.S.N.) ; Easier Museum, Abteilung Entomologie ; Museum National
d'Histoire Naturelle, Paris.

Through the courtesy of specialists and others mentioned in the acknowledgements,
types and other specimens have been borrowed from :

Naturhistoriska Riksmuseum, Stockholm (Stockh.) ; Zoologisches Museum,
Berlin ; Zoologisches Staatsinstitut und Zoologisches Museum, Hamburg (Hamb.) ;
Museum of Zoology, Cambridge ; National Collection of Isoptera, Plant Protection
Research Institute, Pretoria (N.C.I.) ; Museu do Dundo, Lunda, Angola (Dundo) ;
Lamotte collection, Museum de Paris (on loan at the A.M.N.H.).

Private collections (M. G. Bingham, J. Deligne, D. H. Kistner, C. Noirot, J. M.
Pasteels) and unidentified specimens both from the N.C.I, and from the collections
of the Lovanium University, Kinshasa (U.Lov.), where this work has been com-
pleted, must also be mentioned here, as totalizing nearly 900 of all the nest series
examined.

The abbreviations given in brackets indicate the museum of deposit, when listing
material in the text ; if the specimen concerned has not been examined by the
author, the museum of deposit is given in brackets. Localities not indexed in the

TERMITES OF THE GENUS MACROTERMES 367

Times Atlas, vol. IV, or homonymous within the same country (e.g. Katanda,
Congo) are specified by their co-ordinates of latitude and longitude.

METHODS

The species involved in this revision have been redescribed and a full set of
illustrations has been prepared. Previous descriptions have been translated or
quoted whenever the characters involved have been found both useful and clearly
expressed ; in such cases they are integrated into the standardized layout that has
been adopted throughout this work.

Many characters utilized in earlier descriptions of species have been found un-
necessary for this generic revision ; others have proved more variable within one
species than was previously thought ; the following have been newly used or found
of greater importance than hitherto recognized :

Imago : density of setae on head capsule

Soldiers : head profile

shape of metanotum
pilosity of gula

The figures have been drawn with the aid of a camera lucida, from ethanol-
preserved specimens immersed in ethanol ; three scales are used, each of the two
larger being twice the next smaller. The largest scale is used only to illustrate the
inner margin of the left mandible in the minor soldiers (Text-figs 18, 19, 90) ; the
smallest, for the major soldiers throughout, the major workers (Text-figs 20-22)
and the minor soldiers, with the exception of M. ivorensis (Text-figs 46, 47) which
is on the intermediate scale, as are the imagos. In some cases (e.g. Text-figs 70,
123) the drawings include details of the light marks on the cuticle. Antennal
segments have not been illustrated and the setae are usually omitted, except when
conspicuously dense.

With regard to the measurements, it should be noted that the length of the
pronotum has been taken on the median line in the imagos, on the longest lateral
lobe (left or right as the case may be) in the soldiers. The head-width across eyes
in the imagos is measured in dorsal view, from the extremes of curvature of the
eyes ; in the soldiers it is the greatest width, wherever it occurs. Similarly the
depth of the head capsule in the soldiers is taken as the distance in profile view from
the postmentum (or the lowest point of curvature of the ventral genae, whichever
protrudes most) to the highest point of curvature of the vertex, in a line perpen-
dicular to the longitudinal axis of the head. The length of the fore wing is measured
from the suture line to the apical point ; the diameter of the eye is its greatest
one ; the length of the ocellus includes its posterior, non-translucent part ; and,
when measuring the length of the hind tibia (usually the left one), care was taken to
put it at an angle with the femur in order to get the full length of the chitinized
sheath. For the abbreviations used and for other specifications (concerning the
head length and the length of left mandible in the soldiers), see the keys to species.
Finally, as experience has shown that various people using various optical devices
may find slightly different values in measuring the same object, the upper and
lower ranges of dimensions incorporated in the keys have been checked for possible

368 J. E. RUELLE

discrepancies of the kind.

The colour scale adopted in this work ranges from pale yellow to pitch-black, with
(reddish yellow, amber, orange-brown, chestnut-brown) or without (brownish
yellow, brown, smoky brown, sepia-brown) a red shade.

MACROTERMES Holmgren

Macrotermes Holmgren, 1909 : 193. Type-species by monotypy : Termes lilljeborgi (Sjostedt,
1896).

Termes (Macrotermes) (Holmgren) Holmgren, 1910 : 286.

Macrotermes Fuller, 1921 : 17 [return to full generic status].

Bellicositermes Emerson, 1925 : 298. Type-species by original designation : M. bellicosus
(Smeathman) [as subg., syn. Macrotermes, Snyder, 1949 : 208].

Hepilitermes Sjostedt, 1926 : 79, nom. nov. for Tumulitermes Sjostedt, 19240 : 253. Type-
species : Macrotermes schoutedeni Sjostedt [syn. Macrotermes, Snyder, loc. cit.].

Amplitermes Sjostedt, 1926 : 81, nom. nov. for Termes Holmgren [as subgen.] [syn. Bellicosi-
termes Sjostedt, 1926 : 366].

Bellicositermes Emerson ; Grasse, 19370 : 34 [full genusl> [sYn- Macrotermes, Snyder, 1949 : 208].

Imago. Large, width of head across eyes 2-8 to 4-5 mm, length of fore wing 24 to 43 mm ;
head oval, postclypeus same colour as, or lighter than, head capsule, barely to distinctly inflated ;
fontanelle inconspicuous to distinctly protruding ; distance ocellus ex eye 1/4 to 4/3 of ocellus
major diameter ; a subsidiary tooth on the first marginal of the right mandible ; antennae
ig-jointed ; pilosity variable, but without microsetae distinct from the longer bristles.

Pronotum trapezoidal, index length/width 0-4 to 0-6.

Wings translucent to smoky brown ; mediana of the anterior wing isolated from the scale
onwards ; intersegmental membrane of the abdomen in queens without setae.

(The characters of right mandible, pilosity, and mediana of fore wing are useful in separating
Macrotermes from Odontotermes.)

Soldier. Dimorphic. Head length (without mandibles) of the major soldier 4-0 to 7-7 mm,
of the minor 2-05 to 4-77 mm ; labrum widest at or before its middle, with a well-developed
hyaline tip (acute to obtuse, sometimes lanceolate, faintly suggesting a trilobed one) ; mandibles
swordlike, more or less slender and incurved, but without any conspicuous tooth near the middle
of inner margin ; antennae 17 to i8-jointed ; gula narrow, at least twice, often three times
longer than wide, sides subparallel to concave.

Pronotum sellate, distinctly bilobed, middle region of front margin more or less deeply
incurved, but never with forward projecting spines.

Head capsule in major soldiers rectangular to pear-shaped, the length exceeding the maximum
width by 15 to 50% of the latter. Pilosity variable. (Again, no macro- and microsetae.)

Minor soldier : size about half that of major soldier, similar in general appearance, but with
mandibles more slender, antennae and legs often proportionately longer. Colour usually
lighter.

(The overall size, the ubiquitous dimorphism of the caste with a general similarity between
major and minor soldiers, the mandibles, the pronotum and the hyaline tip of the labrum are
among the most important of the generic characters of Macrotermes soldiers.)

Worker. Dimorphic. Mandibles similar to those of the imagos. Head capsule rounded,
width in the major caste 2-05 to 3-50, in the minor 1-33 to 2-41 mm.

Holmgren (1909) first created the genus Macrotermes with Termes lilljeborgi as
type-species by monotypy ; subsequently (1910, 1911) he reduced it to subgeneric
status and erroneously designated another species, M. carbonarius, as the type-
species. In 1912 he listed all Indo-malayan species under this subgenus, while

TERMITES OF THE GENUS MACROTERMES 369

stating that the soldier of M. gilvus does not have its head constricted in front like
the other Macrotermes ; his other subgenus, Termes, included T. bellicosus, T.
goliath, T. michelli, T. natalensis and T. nigeriensis.

Fuller (1921) pointed out that the name Termes was invalid for this particular
genus, and proposed instead the name Macrotermes.

Sjostedt (1926) created the genera Hepilitermes and Amplitermes in addition to
Macrotermes. In the same work, however, he acknowledged (p. 366) the priority of
the subgenus Bellicositermes, created by Emerson (1925).

Emerson (1928), while maintaining Macrotermes and Bellicositermes as subgenera
of Macrotermes, expressed his doubts about the value of this subgeneric division.
In his later publications as well as in his private communications, he dropped the
name Bellicositermes altogether.

Grasse" (from 1937) and Noirot have maintained a generic division between
Macrotermes and Bellicositermes up to the present day, although they noted (Grasse" &
Noirot, 1951 : 330) that the Asiatic species did not fit very well in this distinction.
In describing M . angolensis, Noirot (1955) remarked that the species had some mixed
characters.

These authors have not taken into consideration M. vitrialatus, a widely variable
form including several formerly recognized species as synonyms. This species
finally invalidates any attempt to divide the genus into two, because it spans the
inter-specific gap postulated by Grass6 & Noirot in both its ecology and morphology.
It confirms the impression gained from the Asiatic and Indo-malayan species that
this is one genus.

Some kind of infra-generic grouping may be valid and useful for descriptive
purposes, but it is thought that this should be of an informal kind without nomen-
clatural status.

DISTRIBUTION

This study being restricted to the Ethiopian species of one genus, the phylogeny
of the group is not discussed here. The density of individual species throughout
their geographical distribution is another point about which no definite, let alone
quantitative, conclusions can be drawn, since a few countries only have been
systematically searched. With these restrictions, the following may be noted :

Macrotermes has been collected over the whole of the Ethiopian Region, in a
variety of habitats from rain forests to subarid zones bordering either the Sahara
or the Kalahari deserts. It extends north-east to Southern Arabia, but records
from Madagascar, already considered as doubtful by Emerson (1928 : 445), have not
been confirmed.

The forest species : M. ivorensis, M. lilljeborgi, M. muelleri, M. nobilis (Emerson,
1928 : 441 ; Grass6 & Noirot, 1951 : 309), seem indeed restricted to sheltered
habitats, although they are not strictly confined to the lowland coastal or equatorial
zones. M. muelleri does occur in many a riverine or secondary, low-canopied forest
of minor area in the Congo basin, as illustrated by Weidner (1961 : 34) from Dundo,
Angola. (Incidentally, no forest Macrotermes have been recorded south of this latter
locality.)

370 J. E. RUELLE

The most widely distributed species, after the splitting of M. natalensis, is M.
subhyalinus (formerly M. bellicosus). This species tolerates drier conditions than
does M . bellicosus, as Grasse" & Noirot (1961 : 325) have already stated. The same
authors, however, make it clear (loc. cit. and PI. XV) that M. subhyalinus can
thrive in the equatorial forest.

M. bellicosus tolerates relatively dry conditions in West Africa, being found from
the forest edge to the Sahel savanna (800-400 mm annual rainfall). Its abundance
from Senegal to Uganda and in the Congo basin makes it hard to explain its absence
from further south, although it should be noted that specimens from Congo forest
clearings are larger than those from Northern Nigeria, suggesting a more favourable
environment in the sub-climatic zones with abundant rains.

M . falciger, which ranges mainly in eastern and southern Africa, seems to have
somewhat higher humidity requirements than M. mossambicus and M. natalensis.
From the extensive collections made by the Plant Protection Research Institute of
Pretoria, it appears that the two latter species, together with M. subhyalinus,
are living in many places in South -West Africa where no M. falciger has been
recorded.

Little can be said here of M. herus and M. ukuzii, in view of their limited dis-
tribution. M. vitrialatus, through the range of climates under which it has been
collected, compares most closely with M. falciger but for the fact that it extends
farther to the north-west than the latter, while being apparently absent from
Tanzania.

Additional notes, together with the distribution maps, will be found under the
description ol each species, but more data are needed about the nutritional habits of
Macrotermes before deciding whether and to what extent the presence or absence of
some vegetation type regulates the geographical distribution of the individual species.
Similarly, their soil requirements, if any, have failed to show up clearly in the
course of this revision.

KEYS TO SPECIES

' The increased understanding of the variability of all castes has led to difficulties
in constructing suitable keys to species. ( . . ) Some of the species are represented
by very small numbers of specimens, and it is anticipated that discovery of new
material may necessitate the reassessment of the validity of key characters in these
cases.' (Sands, 1965 : 17). This holds true with Macrotermes. However, in
species where numerous specimens could be examined, it has been found that mean
numerical values are worth mentioning in the following keys. The text-figures
should also help in appreciating some qualitative characters.

IMAGOS

1 Head width across eyes (H) more than 3-8 mm ....... 2

- H less than 3-8 ............ 6

2 Median length of pronotum (PL) more than half its maximum width (PW) ; fon-

tanellar area depressed, fontanelle protruding ....... 3

- PL less than half of PW ; fontanelle not protruding ...... 4

TERMITES OF THE GENUS MACROTERMES 371

3 Depressed area horseshoe -shaped ; ocellus ex eye (O-E) not less than f of ocellus

major diameter (O) ; larger, H, 3-76 to 4-27, mean 3-97 (41 specimens out of 42
measured are wider than 3-8 mm) ; hind tibia (Ts), 4-99-5-97, mean 5-58 falciger (p. 381)
— Depressed area semicircular ; O-E usually | of O ; smaller, H, 3-12-3-96, mean 3-48
(9 specimens out of 145 measured are beyond 3-8 mm) ; TS, 4-18-5-29, mean 4-69

subhyalinus (p. 419)

4 Head capsule dark sepia-brown to black, wings smoky ; postclypeus not inflated

(Text-figs 67, 68, 92), same colour as head ; (forest species, West and Central
Africa) ............. 5

- Head capsule chestnut-brown, wings translucent ; postclypeus inflated (Text-figs

136-138), lighter than head ; (savannah and woodland species, Central and South
Africa) .......... vitrialatus (p. 432)

5 Head and postclypeus proportionately longer (Text-figs 65, 66), third antennal

segment longer than first ........ muelleri (p. 405)

- Head and postclypeus proportionately wider (Text-fig. 91), third antennal segment

not longer than first ......... n obi Us (p. 415)

6 O-E more than 0-4 mm, and equal to O ; fontanelle small, not protruding ; (forest

species, West Africa) ........ ivorensis (p. 391)

- O-E less than 0-4 mm and inferior to O ; fontanelle, whether or not in a depression,

protruding ............. 7

7 Tibiae not darker than femora and/or postclypeus same colour as head ... 8
Tibiae darker than femora, postclypeus lighter than head ..... 9

8 Dark-coloured species ; head capsule sparsely pilose ; eye small, major diameter

(E) less than 1-13 ; O-E averaging 0-3 mm ..... herus (p. 388)

- Chestnut-coloured species ; setae abundant on head capsule ; E, 1-14-1 -33,

mean 1-20 ; O-E averaging 0-15 ...... mossambicus (p. 398)

9 Area around fontanelle not depressed ; head pilosity conspicuous ; postclypeus

inflated ; eye proportionately large (H/E index usually 2-9) ; O-E about ^ of O ;
pronotum distinctly narrower than head capsule ; small, H, 2-84-3-47, mean 3-18 ;
(West and Central Africa, Uganda) ..... bellicosus (p. 374)

- Area around fontanelle depressed ; fontanelle followed anteriorly by a median

ridge ; head pilosity variable, rather sparse ; postclypeus less inflated (Text-figs

78, 99, 124 versus 4-5) ; H, any value from 2-93 to 3-96. . . . . 10

10 Eyes proportionately small, H/E index usually 3-3 ; a few inconspicuous setae on

head capsule ; H, 3-03-3-68, mean 3-43 mm .... natalensis (p. 410)

- Eyes proportionately larger, H/E index usually 3 -o or less ; pilosity variable ; H in

most cases either below 3-2 or above 3-35 mm . . . . . . n

11 Smaller, H, 2-93-3-32, mean 3-14 ; T3, 3-48-4-58, mean 3-98 ; (South-East Africa)

ukuzii (p. 428)

- Larger, H, 3-12-3-96, mean 3-48 ; TS, 4-18-5-29, mean 4-69 ; (whole Ethiopian

Region) subhyalinus (p. 419)

MAJOR SOLDIERS

HL : length of head from base to external socket of mandible (in a line parallel to longitudinal
axis of body) ; HW : head maximum width ; M : length of left mandible — always measured
ventrally — from tip to lower external condyle.

1 M always more than 4-3 mm ....... lilljeborgi (p. 394)

M less than 4-3 ............ 2

2 Sides of metanotum straight or slightly convex, perpendicular to hind margin

(Text-figs 69, 72, 93, 140, 146) ......... 3

Sides of metanotum distinctly convex, acute or rounded (Text-figs 9, 27, 45, 53,

79, 101, 126) .........••• 5

372 J. E. RUELLE

3 Head yellow-brown to reddish brown ; eyespots not very conspicuous ; base of gula

not much wider than middle, sides nearly parallel ; margins of pro- and mesonotum
not much darker than centre ; (savannah and woodland species, Central and
South Africa) ......... vitrialatus (p. 432)

Head darker ; eyespots conspicuous ; base of gula distinctly wider than middle,
sides concave ; margins of pro- and mesonotum conspicuously darker than centre
(forest species, Angola, Central and West Africa) ...... 4

4 Median light mark on mesonotum merging with paired light spots near middle of hind

margin, or the whole median region of mesonotum lighter than the edges ; head
rather pear-shaped ; eyespots small ; M, 3-0-3-6; TS, 4-3-5-6 . tnuelleri (p. 405)

Median light mark on mesonotum separated by a dark transversal band from paired
light spots near hind margin ; head rather trapezoidal, sides straight ; eyespots
bigger, diameter about 0-2 mm ; M, 2-6-3-06 ; TS, 3-9-4-51 . . nobilis (p. 415)

5 Head distinctly narrowed in front ; HW more than PW by about i -4 ; T$ less than

HW by about 0-3 mm ; (forest species, West Africa) . . ivorensis (p. 391)

Head scarcely narrowed in front ; HW more than PW by less than i -4 ; T$ less than

HW by at least 0-6 mm ; (any region) ........ 6

6 Fontanelle inconspicuous and no ridge or grooves in front of it ; mandibles not

upcurved, left one only slightly more incurved than right (Text-figs 8, 1 1) ; thoracic
nota much narrower than head, poorly chitinized, corners acute. bellicosus (p. 374)

Anterior part of head more or less deeply sculptured and/or thoracic nota large,
distinctly more chitinized than abdominal nota, sides generally rounded ;
mandibles upcurved, at tip or throughout (Text-figs 57, 84), or left one strikingly
more incurved than right (Text-figs 28, 56) ....... 7

7 Head conspicuously pilose (Text-figs 31, 32, 129) ...... 8

- Cephalic setae rather sparse (e.g. not more than 20 to 30 setae on gula) ... 9

8 HW more than 3-7, M more than 2-7 ; (Text-figs 27-30) . . falciger (p. 381)
HW less than 3-7, M below 2-7 ; (Text-figs 125-127) .... ukuzii (p. 428)

9 Head in plan view rather ovate or horseshoe-shaped, sides convex, moderately

converging towards anterior end ; in profile evenly flattened (Text-figs 83, 84) ;
fontanelle conspicuous but not followed anteriorly by elevated median ridge ; left
mandible evenly incurved and longer than HD ( = head depth, incl. gula) ; HL,
4-13-6-21, mean 5-15 ; (South, South -West Africa) . . natalensis (p. 410)

- No such combination of characters ......... 10

10 Sides of metanotum angularly convex ; head rectangular, in profile abruptly

flattened from fontanelle to base of labrum, with deep grooves diverging forwards
and conspicuous median ridge ; tip of left mandible upcurved and incurved
enough to point at right angle with main axis ; M less than HD ; HL, 4-26-5-68,
mean 5-02 ; (Text-figs 56, 60) ; (South-West and South Africa) tnossambicus (p. 398)
Sides of metanotum rounded ; left mandible evenly incurved and/or HL more than

5-8 ; (Text-figs 40, 104, 105) ......... n

11 Head dark brown, mandibles black throughout ; smaller, HL, 4-93-6-10, mean

5-44; (North-East Africa) ........ herus (p. 388)

Head reddish yellow to chestnut-brown, mandibles usually with lighter base ;
larger, HL, 4-84-7-70, mean 6-49 (in 435 specimens out of 456 measured, HL
is more than 5-60 mm) ; (whole Ethiopian Region) . . . subhyalinus (p. 419)

MINOR SOLDIERS

Minor soldiers are less characteristic. The key therefore will identify the majority of
specimens but cannot be completely reliable owing to variation.

1 TS more than 3-0 mm ........... 2

TZ less than 3-0 mm ........... 9

2 Ts/HW index less than i -7 .......... 3

TERMITES OF THE GENUS MACROTERMES 373

T/3/HW index more than i -7 ....... . 5

3 Maximum width of gula less than 0-8 mm ; HL, 2-20-3-25, mean 2-62 ; mandibles

straight, tip slightly curvate (Text-figs 147-152) ; (savannah and woodland

species, Central and South Africa) vitrialatus (p. 432)

Maximum width of gula 0-8-1-04 mm ........ 4

4 Left mandible evenly incurved, curvature equal to or greater than 90° ; larger,

HL, 3-04-4-77, mean 3-79 ; T3, 2-57-3-87, mean 3-21 ; (Text-fig. 37)

falciger (p. 381)

- Only tip of left mandible distinctly incurved, total curvature inferior to 90° ;

smaller, HL, 2-67-4-45, mean 3-57 ; Ta, 2-19-3-35, mean 2-78 ; (Text-fig. 116)

subhyalinus (p. 419)

5 M less than 2-2 ..... . -7

M more than 2-2 ........... 6

6 M, 2-22-2-99, mean 2-47 ; distinctly wider than right mandible ; front margin of

pronotum deeply emarginate ; HL, 2-83-3-82, mean 3-11 ; (Text-fig. 49) ;
(West and Central Africa) lilljeborgi (p. 394)

- M, 1-64-2-57, mean 1-82 ; not wider than right mandible ; front margin of pronotum

shallowly emarginate ; HL, 2-32-3-23, mean 2-53 ; (Text-fig. 94) ; (Central
Africa) nobilis (p. 415)

7 Quite a few bristles on head capsule ; gula in lateral view protruding, and more so

in the anterior third (Text-fig. 47) ; general colour reddish brown ; gula L/W

index 2-3 ivorensis (p. 391)

Head smooth ; gula in lateral view less protruding and evenly convex (Text-figs

76- 95) • 8

8 General colour light to dark reddish brown ; thorax not lighter than head ; gula

narrow (L/W index, 2-9) muelleri (p. 405)

General colour sepia-brown ; thorax lighter than head ; gula proportionately

wider (L/W index, 2-45) nobilis (p. 415)

9 Fontanelle inconspicuous, i.e. opening small, level with head capsule, circular or

slit-like 10

Fontanelle conspicuous, i.e. protruding, wart-like, crateriform, wide open or any

similar appearance .......... 1 1

10 TS less than HL ; metanotum not wider than mesonotum ; left mandible stout,

width about ^ of length, serrate when viewed from beneath (Text-figs 15-17,

18-19) bellicosus (p. 374)

TS more than HL ; metanotum distinctly wider than mesonotum ; left mandible
slender, width about | of length (Text-figs 147-152) ; T3, 2-52-3-42, mean
3-00 . . . . . . . . . . . vitrialatus (p. 432)

11 Large, HL usually 3-6-3-8, PW 2-4-2-6 . . falciger or subhyalinus, see 4
Medium, HL usually 2-8-3-1, PW more or less 2-0 .. . 12

- Small, HL, 2-21-2-63, mean 2-38 ; PW, 1-43-1-66, mean 1-56 ; mandibles almost

straight ; (South Africa) ukuzii (p. 428)

12 Head uniformly orange-brown ; margins of thoracic nota darker than middle region ;

in plan view, head shortly ovate, metanotum distinctly wider than mesonotum

(Text-figs 61-64) . mossambicus (p. 398)

Head chestnut-brown to smoky brown, paler beneath, more elongate and more

distinctly narrowed in front ....... • X3

13 Colour generally dark (smoky to dark sepia-brown) ; setae on labrum, thoracic and

abdominal tergites with conspicuously lighter area around base ; postero -lateral
sides of pronotum rather straight (Text-fig. 44) .. . herus (p. 388)

Colour generally lighter (yellow to light chestnut-brown) ; setae without con-
spicuously lighter area at base ; postero-lateral sides of pronotum convex
(Text-figs 87-89) natalensis (p. 410)

374 J- E. RUELLE

Macrotermes bellicosus (Smeathman)
(Text-figs 1-20 ; Map i)

Termes bellicosus Smeathman, 1781 : 141. Type-locality : SIERRA LEONE. [Type materia
lost.]

Termes nigeriensis Sjostedt, 191 ia : 183. Type-locality : NIGERIA, Yaba, syn. n.

Termes carboniceps Sjostedt, 1924 : 39. Type-locality : CONGO, Yambata, syn. n.

Bellicositermes convexus Grasse, 19370 : 35 [svn- M. bellicosus (as Bellicositermes natalensis)
Grasse & Noirot, 1961].

Macrotermes bellicosus (Smeathman) Snyder, 1949 : 209 [first use of combination ; mis-
identification].

The above synonymy contains only entries of nomenclatural significance. Since
Hagen (1858) made the first misidentification, Smeathman's name has been wrongly
applied to the species now recognized as M. subhyalinus and M. mossambicus.
To correct all the subsequent misidentifications in the synonymy would obscure
the nomenclatural entries. It is therefore to be noted that citations of the species-
name natalensis under the generic headings of Amplitermes, Macrotermes or Bellicosi-
termes in the literature (e.g. Sjostedt, 1926 ; Emerson, 1928 ; Grasse & Noirot,
1951, 1955, 1958, 1961) and not included in the material examined in this revision,
should henceforward be referred to M. bellicosus, but only if recorded from the
geographical range of this species.

FIGS 1-5. Macrotermes bellicosus, imago. 1-3, head capsule and pronotum, plan view

4-5, head capsule, side view.

TERMITES OF THE GENUS MACROTERMES

375

Neotype, major soldier. Head capsule yellowish brown, mandibles black, with reddish base.
Gula darker than ventral genae. Pronotum lighter than head ; mesonotum and metanotum
lighter still, and similar to the abdominal tergites. Legs yellow, tibiae not darker than femora.

Head in plan view rectangular, sides almost straight, only slightly converging in front.
Mandibles not upturned, left one broader and slightly more incurved than right, inner margin
viewed from beneath faintly serrate. In profile, head capsule rather flat (opp. subhyalinus) ,
upper and lower surfaces parallel, gula not prominent. Antennae ly-jointed, third segment
almost twice as long as the second ; fourth and subsequent segments shorter than second ;
first and third equal. A few scattered setae on the head capsule, more setae on the gula.
Labrum oval ; hyaline tip triangular with almost concave sides.

Pronotum relatively small (opp. subhyalinus,) front margin more emarginate than hind
margin ; antero-lateral corners more acute than in subhyalinus, sides nearly straight. Meso-
notum distinctly narrower than pronotum and only slightly narrower than metanotum ; side
of mesonotum with obtuse tip. Hind margin of metanotum convex. Thoracic nota less
sclerotized than in subhyalinus.

Measurements in millimetres : holotype — range and mean of 671 specimens from 131 localities :

Head length .....

Head width .....

Head depth inc. gula

Length of gula .....

Maximum width of gula

Minimum width of gula

Length of left mandible

Length of hind tibia

Maximum length of pronotum .

Maximum width of pronotum

Minor soldier. Head somewhat paler than that of major soldier. Mandibles dark reddish
brown. Thoracic tergites paler than the head, and similar to the abdominal tergites. Head
more elongate, mandibles more slender and more symmetrical than in the major soldier. Left
mandible distinctly serrate (Text-figs 18-19). Hyaline tip of labrum with convex sides.

Thoracic tergites : pronotum the widest, mesonotum the narrowest ; metanotum, however,
not much wider than mesonotum.

Measurements (427 specimens from 106 localities) in millimetres.

Head length ....
Head width ....
Head depth inc. gula
Length of gula
Maximum width of gula
Length of left mandible
Length of hind tibia .
Maximum length of pronotum .
Maximum width of pronotum

Major worker. Head brownish, lacking the reddish tinge of subhyalinus ; posterior margin
less rounded than in either natalensis or subhyalinus (Text-figs 20-22).

Head width (204 specimens from 94 localities) : range 2-04-2-91, mean 2-47 mm.

Morphotype imago. Head dark brown. Postclypeus lighter than head, with dark median
line. Thoracic tergites similar in colour to head, pronotum with a lighter T-shaped mark
(as in subhyalinus). Abdominal tergites dark brown. Wings translucent, smoky.

Holotype

Range

Mean

5-3i

4-13-6-30

5-3i

4-40

3-I4-5-I7

4-16

2-58

i -96-2 -96

2-48

3-6i

2 -99-4 -08

3-54

1-30

0-92-1-33

I-I2

0-83

0-70-1-13

0-85

2-72

2-10-2-93

2-6O

3-47

2-68-4-01

3'4°

1-47

I-I8-I-69

I -46

3-13

2-33-3-56

2-99

Range

Mean

2-13-3-42

2-77

i -77-2 '79

2-22

1-09-1-72

i-37

i -50-2 -02

i-75

0-56-0-75

0-65

I-43-2-I5

1-77

1-77-2-63

2-25

0-79-1-23

0-94

1-43-2-16

1-77

376

J. E. RUELLE

Head rounded, numerous setae on head and pronotum, with light dots around base. Frons
flat, not depressed around fontanelle ; fontanelle conspicuous, on top of a small hump, without
any anterior median ridge (opp. natalensis). Eyes broadly oval, large, but less rounded than in
subhyalinus and less inflated than in mossambicus. Ocelli nearly circular, not more than about
J of their diameter distant from the eyes (£ in natalensis). Postclypeus wider and more inflated
than in subhyalinus. Antennae ig-jointed, third segment longer than second and fourth.

Pronotum with anterior margin flat, posterior broadly indentated ; proportionately wider
than in subhyalinus.

Measurements in millimetres : morphotype — range and mean of 160 specimens from 56
localities :

Head width across eyes
Eye major diameter .
Ocellus major diameter
Ocellus minor diameter
Ocellus ex eye .
Median length of pronotum
Maximum width of pronotum
Length of hind tibia
Length of fore wing (tip-scale)
Maximum width of fore wing

Morphotype Range Mean

3-07

2-84-3-47

3-18

1-07

0-95-1-25

i-n

0-42

0-34-0-50

0-41

0-36

0-20-0-40

o-33

0-14

0-05-0-27

O-II

1-68

1-36-1-76

i -60

2-86

2-57-3-39

3-04

4-50

3-96-5-21

4-64

27-00

24-00-34-00

29-40

7-10

6-00-9-00

7-30

Variation. Lateral expansions of the mesonotum may make it occasionally wider than the
metanotum, in the major soldiers. In a sample of 50 specimens from 19 localities, this was the
case with 27 specimens. In point of size, the West African specimens are smaller than those
from Central Africa, the average difference being 0-2 mm for head width of imago and head
length of minor soldier, 0-5 for head length of major soldier, 6 • o for length of fore wing in
alates. But the only non-overlapping figures are those of the wing length and this character
alone does not seem to warrant a subspecific distinction.

The termites from the oldest collections of the Linnean Society (London) include
one unlabelled major soldier of M. bellicosus and two minor soldiers, labelled :
' Termes bellicosus milites Smeathman, Termes fatalis L. milites '. These two
minor soldiers belong to one species of Macrotermes, the specific determination of
which could not be ascertained. Since there is neither an indication of locality nor
any resemblance of the writing on the labels to Solander's writing, it has been
concluded that Smeathman's types are lost. A neotype from Sierra Leone has been
selected from the B.M.(N.H.) collections. Although Freetown is nearer to the
place where Smeathman had made his observations, a sample from a mature colony
at Njala has been chosen, since the material available from Freetown is less satis-
factory (immature colonies).

Evidence obtained in the course of this revision has revealed both a morphological
and a geographical gap between what used to be called M. natalensis in West and
Central Africa and the species described by Haviland ; hence the present splitting.

The type imago of Termes nigeriensis Sjostedt could not be found in the Stockholm
collections ; the ' topotype ', as labelled by Dr A. E. Emerson (unpublished),
and any other samples of ' M. nigeriensis ' from the same region are indiscernible
from M. bellicosus.

Termes carboniceps Sjostedt, known by the soldier caste only, is a larger than

TERMITES OF THE GENUS MACROTERMES

377

FIGS 6-22. Macrotermes, soldiers, major worker. 6-19. M. bellicosus. Major soldier ;
6-10, head capsule and thoracic nota, plan view ; 11-14, head capsule, side view.
Minor soldier ; 15-17, head capsule and thoracic nota, plan view ; 18-19, left mandible,
inner margin. 20-22. Major worker, head capsule, plan view : 20, M. bellicosus ;
21, M. natalensis ; 22, M. subhvalinus.

378 J. E. RUELLE

average and accidentally blackened sample of M. bellicosus. This conclusion was
reached after a collecting trip through the type-locality (n.ix. to 2.x. 1966).

NEOTYPE. SIERRA LEONE : Njala (Lat. 8°6' N., Long. 12 °5' W.), 24.viii.iQ30
(E. Hargreaves, Coll. No. 407). Neotype major soldier and other material from
same colony in B.M.(N.H.). Morphotype imago. GHANA : Accra, i.i955 (W. V.
Harris, Coll. No. 887), B.M.(N.H.).

Type Material. Paratypes, type colony, A.M.N.H. (previously determined by
Dr A. E. Emerson as M. natalensis var. tumulicola, with the comment : ' comp.
cotype natalensis from Natal & believe different variety at least ') : other paratype,
N.C.I.

Other Material. PORTUGUESE GUINEA : Bamboya (12° N., 16° W.), I.R.S.N. ;
Bissao 1898 (H. Ehrhardt], Stockh. £ Hamb., 1901 (S otter), Hamb. ; Bula Bulamelo,
(i2°io' N., I5°4o' W.), I2.iii.i953 (/. P. P. Amara) ; no locality, 1897 (Schrecken-
bach), Hamb.

GUINEA : Conakry, io.xi.i892 (Brauns), Hamb. ; Fouta Djalon (Pobeguin),
Stockh. ; Kindia, 2i.viii.i9i2 (F. Silvestri), A.M.N.H. ; Kissidougou (Chevalier),
Stockh. ; Nimba Mts, 1941 to 1956, 37 vials (M. Lamotte), Lamotte collection.

SIERRA LEONE : Bo, 11.1.1958 and Freetown, 7.1.1958 (W. Wilkinson) ; Pujehun,
I5.iii.i947 (F. A. Squire], A.M.N.H.

LIBERIA : no locality, no date, ' cotype T. tumulicola ' (Benson), 4.viii.i926 (/.
Bequaert}, Bendija (co-ordinates?), 1940, 4 vials (W. M. Mann), A.M.N.H. ; Cape
Mount (6°4i' N., n°2o' W.), no date (Buttikofer] , Stockh. ; Charlesville (6°i5' N.,
io°2o' W.), io.vi.i962 (D. H. Kistner), private collection ; Loffa River, I7.viii.i937
(H. Schomburgk], Hamb. ; Monrovia, vii.i926 (/. Bequaert), A.M.N.H. ; Zeanschve
(6°5o' N., 9°5o' W.), ix.i926 (/. Bequaert), Terv. & A.M.N.H.

IVORY COAST : Adiopodoume* (5°2o' N., 4°o8' W.), 7.viii.i953 (M. Lilscher),
Touba, ' cotype B. convexus ', A.M.N.H. ; no locality, 1908 (Bouet), Stockh.

GHANA : Accra, ? i., Kumasi, 3.^.1955 (W. V. Harris) ; 10 m. N. of Yeji on
Kumasi-Tamale Rd, 25., n m. fr. R. Volta on Nyankpala (Tamale) Rd, 28. ii.,
35 m. fr. Tamale on Yendi Rd, 4., 29 m. fr. Navrongo on Tumu Rd, 13., 40 m. tr.
Tumu on Lawra Rd, 16., 9 m. fr. Lawra on Nandom Rd, 18., 52 m. fr. Wa on Sawla
Rd, 20., 19 m. fr. Sawla on Damongo Rd, 21., Ejura, 27., 12 m. fr. Techiman on
Wenchi Rd, 29.^.1959 (W. A. Sands).

TOGO : Bismarckburg (8°n' N., o°4o' E.), no date (Buttner), Stockh. ; Lome",
i4.iii.i9O2 (Ed. Burdis], Hamb.

NIGERIA : Western Region : Yaba nr Lagos, 4.^.1933, collector unknown,
topotype collection of Termes nigeriensis, A.M.N.H. Other Material : Ijora
(Lagos), no data, Ibadan, 2.^.1953 (G. C. Webb), A.M.N.H. ; Bennin (PBenin City),
1907 (C. Manger), Hamb. ; 27 m. S. of Ilorin on Oyo Rd, 4., Olokemeje, 8., Agodi
(Ibadan), io.xii.i957 (W. A. Sands) ; 4 m. fr. Asaba on Benin Rd, 7., Obanokoro,
8.i., 4.iii., Lagos, n.iv.i957 (W. Wilkinson) ; 17 m. fr. Lagos on short Ibadan Rd,
1955 (B. J. MacNulty). — Eastern Region : Niger delta, 1907 (C. Manger), Hamb. ;
Onitsha, 1954, 27 m. fr. Onitsha on Owerri Rd, 1955 (B. J. MacNulty) ; 47 m. fr.
Enugu on Onitsha Rd, 2., 20 m. fr. Enugu on Oturkpo Rd, 5.1^.1958 (W. A. Sands) :

TERMITES OF THE GENUS MACROTERMES 379

Aba, 27.xii.i956, Abonema and Degema, 5.1., Calabar, 24.ii., Ahoada, so.iv., 40
m. fr. Port Harcourt on Owerri Rd, ig.vi.i957 (W. Wilkinson] ; Port Harcourt,
1954, 1957, 3 vials (W. V. Harris, W. Wilkinson}. — Northern Region : Abuja
(H. Mills], Vom (9°4o' N., 8°4o' E.), 3.^.1950 (G. C. Webb], A.M.N.H. ; Zaria, 4.,
Katsina, 19. xi., 33 m. fr. Kaduna on Zungeru Rd, 18., 16 m. fr. Zungeru on Bida Rd,
21., 5 m. fr. R. Kaduna on Mokwa-Bida Rd, 22., 21 and 41 m. fr. Mokwa on Bida Rd,
22., Bida, 2 vials, 23., Diko, 27.xii.i956, 65 m. fr. Jos on Karachan Rd, 8., 16 m. fr.
Jos on Bauchi Rd, n., 25 m. fr. Bauchi on Jos Rd, ig.ii., Kwal nr Miango, i.iv.,
2 m. fr. R. Gongola on Gombe-Dadinkowa Rd, 9., Tula, 10., 22 m. fr. Gombe on
Ture Rd, 2 vials, 10., 34 m. fr. Yola on Jalingo Rd, 14., 27 m. fr. Beli on Jalingo Rd,
ig.v., 43 m. fr. Maiduguri on Ft Lamy Rd, 3., 30 m. fr. Maiduguri on Bama Rd,
4.vi.i957, 42 m. fr. Gboko on Oturkpo Rd, 26.ii., Lokoja, 8., 22 m. fr. Lokoja on
Kabba Rd, io.iii.i958 (W. A. Sands).

CAMEROON : Mamfe, 27^.1957 (W. Wilkinson).

CENTRAL AFRICAN REPUBLIC : Bania, and Nola (Eriksson), Stockh. ; Lobaye
(3°4o' N., i8°io' E.), 1966 (L. Bouquiaux], own collection.

CONGO (Brazzaville) ; Mate"le" (o°3i' N., i6°38' E.) (Karlsson), Ngoko (Gravot),
Stockh.

CONGO : Yambata, ii.-iii.i9i4 (de Giorgi), type collection of Termes carboniceps,
Terv. (other syntypes in Stockh. and A.M.N.H.). Other Material : Umangi (de la
Kethulle, Wilverth,J. E. Ruelle], Terv., I.R.S.N., Stockh., U.Lov. ; Duma (Montchal,
Schubotz], Terv., Hamb., Stockh. ; Kisantu, xii.i92O, Luebo, viii.i92i (H. Schou-
tederi), Terv., A.M.N.H., Stock. ; Bumba (2°io' N., 22°30; E.) (Lootens, A. Anberg),
Kabambare, 18, 2O.X.I954 (N. Leleup), Tshikapa, 25.xi.i92i (H. Schouteden}, Terv.,
A.M.N.H. ; Bikoro, i.iii.igai (H. Schouteden], Dungu (Hutereau), Moto (3°O2' N.,
29°3o' E.), 1920-1923 (L. Burgeon], Nyangwe, 7.1.1911 (/. Bequaert], Yakoma
(Bomstein), Terv., Stockh. ; Kimwenza (4°3o' S., I5°i3' E.), 30.ix.i96o, Lisala,
I9.ix.i966 (/. E. Ruelle}, U. Lov., N.C.I. ; Alberta (2°i2' N., 22°26' E.), xi.i92i
(Tinant), Bambesa (3°23' N., 25°47' E.), 29. xi. 1930 (Vrydagh], Banzyville, ix.ig26
(Mestdagh), Bengamisa (A. Henrion], Doruma, 1927 (Walkiers], Egbunda (2°4o'
N., 27°io' E.), in. 1935 (Johnen), Irumu, 2.vii.i9i4 (/. Bequaert), Karawa, 1936-1937
(Wallin), Katako-Kombe, 19.1.1908 (G. Gustin), Kiambi (7°i4' S., 27°52' E.), 1.1931
(P. Quarre], Kindu, 1935 (Rossignol), Kondue (4°59' S., 23°2o' E.) (E. Luja], Kotili
(2°5i' N., 24°34' E.), 14.1., Mauda (4°o5' N., 27°4i' E.), i.iii., Mahagi, 18^.1925
(H. Schouteden), Lakulu (3°28' N., 23°42' E.), 1930 (van den Branden], Lukolela
(i°io' S., 17°!!' E.), xi.i934 (Ledoux), Mpa (3°io' S., 18° E.) (/. Maes), Mukishi
(8°3i' S., 24°4i' E.), 25.xi.ig27 (A. Becquet), Pawa (2°3i' N., 27^0' E.) (Claessens),
Tora, 1926, 3 vials (L. Burgeon), Van Kerkhovenville (3°2i' N., 29°32' E.) (De Greeff],
Witshi-Tadi (3°3o' S., 23°3o' E.), 1.1930 (/. Allaer], Terv. ; Kabinda (Mutter],
Kangu (5°i7' S., i2°57' E.), 1932 (de Schaetzen), Kinshasa (Lamarche), Popokabaka
(/. Mertens), I.R.S.N. ; Faradje, 1912 and 1948, Niangara, 1913 and 1948 (Lang-
Chapin, N. A. Weber), Kwamouth, 14^^.1909 (Lang-Chapin), Rwindi Camp (o°47'
S., 29°i8' E.), 6.v., Rifflart (4025' S., I5°2i' E.), 9^.1948 (A. E. Emerson), A.M.N.H.;
Tsunza (5^8' S., i6°28' E.), Kimvula, 9., Mabaka-Nzadi (6°o6' S., i6°42' E.), 10.,
Kitenda (7°oi' S., I7°i8' E.), 12., Suka (7°34' S., I9°i4' E.), 13., Wamba R. (7°i4'

38o J. E. RUELLE

S., I7°43' E.), Panzi, 14., Kisandji, 15., Kikwit, 18., Inzia R. (5°2g' S., I7°29' E.),
19., Imbela (5°54' S., I7°o8' E.), 20., Swa-Kilamba (4°53' S., I7°o8' E.), 22., Motor-
ensiene (3°5o' S., 18° E.), 25., Beno, 25., Gwafumu R. (3°28' S., i7°35' E.), 26.,
Kutumpai (3°45' S., I7°23' E.), 28.viii.i959, Katanda (6°23' S., 23°55' E.), 23.vii.,
Mbuji-Mayi (= Bakwanga), 3i.vii.i963, Mondongo (2°io' N., 2i°io' E.), I3,i5,29.ix.,
Yalosemba (2°35' N., 2i°5o' E.), 2i.ix.i966 (/. E. Ruelle), Kutu, iS.ig.viii., Kin, 3.,
Pendjua (i°o6' S., I9°o5' E.), 5.ix.i966 (P. Cappelle), U. Lov.

MAP i. Collecting localities of Macrotermes bellicosus.

TERMITES OF THE GENUS MACROTERMES 381

SUDAN : Mt Bangenze, 17. ¥.1937 (J. G. Myers), Imatong Mts, 24.vii.i939, Shambe,
vii.-viii.i939 (N. A. Weber], A.M.N.H. ; Singi (5° N., 29° E.), zo.vi., Magwe,
4.viii.i962, Wadupe (4° N., 31° E.), 4.1.1960 (H. Schmutterer], Hamb.

UGANDA : Southern Buruli, Coll. No. 457, Entebbe, 2O.xi.i94i, Kazinga Flats
(o°4o' S., 30° E.), 1937, Kichwamba, I5.xi.i949 (W. V. Harris], Karamoja 40 m. fr.
Moroto, I2.X.I952 (W. A. Sands], Salalira (i°i4' N., 34°i7' E.), 24.1.1930 (H.
Hargreaves], B.M.(N.H.) ; Kibuji (Lango), Coll. No. i, West Nile Region, Coll.
No. 6, Bunyoro (Butiaba), Coll. No. 7,14, Katwe, Coll. No. 9, Wasa, Coll. No. n
(D. R. Buxtori), Kampala, 9,n.xii., Kinyanga Estate (i°46' N., 3i°33' E.), i8.xii.i934
(H. Kirby], Munyonyi (o°i6' S., 30°i4' E.), ix.i940 (collector unknown], Serere,
xii.i929 (P. Chandler], Teso, xii.1937 (S. M. Watson], A.M.N.H.

KENYA : West Suk Distr., 4.X.I952 (W. A. Sands] ; Busnia (? Busia), 17.11.1948
(N. A. Weber), A.M.N.H.

TANZANIA : Amani, 25.ix.ig34 (H. Kirby), A.M.N.H.

ANGOLA : Dundo, v.-viii.i948, 4 vials (A. de Barros Machado], (Dundo).

A total of 314 nest series were examined, and all material is in the B.M.(N.H.)
unless otherwise indicated.

Neither Dundo nor a few other, not listed above, records from the Ivory Coast
(Abidjan, Bingerville, Kouibli, Man) or former French Equatorial Africa (Bossem-
bele1, Makoua) have been substantiated by redetermination and the localities have
not been plotted on the distribution map. The determinations, however, are
thought reliable (Weidner, 1956 ; Grasse", Grasse" & Noirot passim], the name
' natalensis ' meaning ' bellicosus '.

The biology and the ecology of M. bellicosus have already been extensively studied
(Smeathman, 1781 ; Grasse", 19370, 1944 ; Kutter, 1943 ; Grass6 & Noirot, 1951,
J955. I958, 1961 ; Liischer, 1955, 1956, 1961 ; Ruelle, in Bouillon, 1964 : 231,
327 ; Bodot, 1967, 1 967*2). As far as is known, their architecture is the most
elaborate of all the Macrotermes.

Macrotermes falciger (Gerstacker) sp. rev., comb. n.

(Text-figs 23-37 '• Map 2)

Termes falciger Gerstacker, 1891 : 186. Type-locality : TANZANIA, Mbusine.

Termes goliath Sjostedt, 18990 : 156. Type-locality : TANZANIA, Kilimandjaro, syn. n.

Termes michelli von Rosen, 1912 : 223. Type-locality : CONGO, North Katanga, [syn. M.

falciger (as Amplitermes goliath), Sjostedt, 1926 : 85.]
Termes swaziae Fuller, 1915 : 462. Type-locality : REPUBLIC OF SOUTH AFRICA, Tzaneen,

syn. n.

Macrotermes usutu Fuller, 1922 : 79. Type-locality : SWAZILAND, Usutu R., syn. n.
Tumulitermes kibonotensis Sjostedt, 19240 : 253. Type-locality : TANZANIA, Kilimandjaro,

Kibonoto, syn. n.
(Here again, citations of the species-name goliath, or misidentifications without nomenclatural

significance, such as : Sjostedt, 19000, 1904, 1907 ; Fuller, 1922 ; Harris, 1958, have been

omitted from the synonymy.)

Imago. Head dark chestnut-brown, lighter around fontanelle. Postclypeus same colour as,
or slightly lighter than, head. Pronotum darker than head, with T-shaped yellowish mark in

382 J. E. RUELLE

the middle, two yellow spots near antero-lateral corners. Tibiae darker than femora. Abdominal
tergites darker than sternites. Wings light smoky.

Shape of head capsule in plan view not distinctive ; setae sparse. Fontanellar area depressed,
depression in dorsal view horseshoe- or U-shaped ; fontanelle conspicuous, surrounded by
short setae, followed anteriorly by a distinct median carina. Eyes proportionately small
(H/E index superior to 3-1), not very prominent. Ocelli small, oval, about f of their major
diameter distant from the eye. Postclypeus relatively narrow, but somewhat inflated.
Antennae ig-jointed, third segment equal to fourth plus fifth.

FIGS 23-26, 35-37. Macrotermes falciger, imago, minor soldiers. 23-26. Imago : 23-24,
head capsule and pronotum, plan view ; 25-26, head capsule, side view. 35-37.
Minor soldier : head capsule and thoracic nota, plan view.

TERMITES OF THE GENUS MACROTERMES 383

Pronotum wider than head across eyes, at the same time long, with front margin shallowly
emarginate, posterior flatly incurvate ; antero-lateral corners rounded. Setae more numerous
than on head capsule.

Hind tibia long, usually well beyond 5-0 mm.

Measurements (42 specimens from 19 localities) in millimetres :

Range Mean

Head width across eyes .... 3-76-4-27 3-97

Eye major diameter . .... 1-11-1-34 I-23

Ocellus major diameter .... 0-37-0-48 0-41

Ocellus minor diameter .... 0-27-0-38 0-32

Ocellus ex eye ...... 0-25-0-45 0-32

Median length of pronotum . . . 1-97-2-27 2-12

Maximum width of pronotum . . . 3-61-4-41 4-01

Length of hind tibia. .... 4 -99-5 -97 5-58

Length of fore wing ..... 33-00-43-00 38-30

Maximum width of fore wing . . . 8-00-10-20 9-10

Major soldier. Head capsule dark chestnut-brown ; mandibles black throughout. Antennal
segments darker than head. Pronotum at times darker than head, meso-, metanotum and
abdominal tergites same colour as pronotum ; abdominal sternites lighter. Legs yellow-brown,
tarsi not darker than tibiae.

Head in plan view rectangular, without distinctive shape, although usually bigger and wider
than in subhyalinus ; in profile showing numerous setae on postmentum, frontal area moder-
ately depressed with ' a median, elongate, triangular and transversely wrinkled mound ' (Fuller,
1915 : 463). Mandibles very stout, left one broader and more incurved than right, the latter
being just hooked at the tip. Antennae 17-jointed, third segment twice as long as the second,
fourth intermediate between third and second. Pilosity of head capsule and postmentum
conspicuous. Hyaline tip of labrum with sides straight or even concave.

Thoracic nota very large, strongly chitinized ; pronotum with lateral lobes broadly rounded ;
pronotum the widest, mesonotum in most cases wider than metanotum ; hind margin of
metanotum not infrequently concave.

Measurements (142 specimens from 64 localities), in millimetres :

Range Mean

Head length ...... 5-20-7-43 6-64

Head width ...... 4-00-6-14 5-29

Head depth inc. gula .... 2-65-3-84 3-28

Length of gula. 4-05-4-93 4-45

Maximum width of gula i -44-1 -83 i -60

Minimum width of gula .... 0-94-1-26 1-09

Length of left mandible .... 2-85-4-01 3-57

Length of hind tibia . .... 3-10-5-16 4-18

Maximum length of pronotum . . . 1-70-2-61 2-21

Maximum width of pronotum . . . 3'3Q-5-i8 4-35

Minor soldier. Head smoky brown, paler beneath ; mandibles black, with dark sepia-brown
base. Antennal segments distinctly darker than head capsule. Thoracic nota and abdomlina
tergites same colour as antennae.

Dorsal outline of head capsule trapezoidal, sides convex ; fontanelle slightly protruding,
wart-like. Mandibles more slender than in major soldier, left one very evenly incurved, right
one hooked, tip of both at 90° or more inwards. Length of hind tibia usually superior to head
width.

Sides of thoracic tergites rounded ; mesonotum nearly as wide as metanotum.

Measurements (176 specimens from 65 localities) in millimetres :

384 J. E. RUELLE

Range Mean

Head length ...... 3-04-4-77 3-79

Head width ...... 2-59-3-87 3-11

Head depth inc. gu la .... 1-64-2-47 2-03

Length of gula . ..... 1-91-3-15 2-40

Maximum width of gula .... 0-84-1-25 0-93

Length of left mandible .... 2-23-3-19 2-65

Length of hind tibia. .... 2-57-3-87 3-21

Maximum length of pronotum . . . 1-20-2-10 1-45

Maximum width of pronotum . . . 2-15-3-42 2-61

Major worker. Head dark chestnut-brown, posterior margin rounded ; width (138 speci-
mens from 36 localities) : range 2-64-3-50, mean 2-99 mm.

Variation. This species is generally darker in colour than M. subhyalinus, but not always.
Neither can one rely on the index mesonotum width/metanotum width or on the concavity of
the hind margin of metanotum in major soldiers to separate M. falciger from M. subhyalinus.
Although the evidence is as yet inconclusive, paler specimens and shorter wings seem more
frequent in South Africa than elsewhere.

The type-collection of M. falciger in the Hamburg Museum had been considered
in 1898 by Sjostedt as Termes bellicosus (date of hand-written label found with the
sample). The re-determination by the present author was first communicated to
Dr H. Weidner and published by him (1966 : 224) as follows : Nach J. E. Ruelle in
litt. : Macrotermes goliath (Sjostedt). According to the law of priority, the name
' goliath ' can not be maintained.

Sjostedt's note (1926 : 85) on T. michelli von Rosen has proved right. T. swaziae
and M. usutu Fuller were already regarded as possible synonyms of M. falciger by
Dr A. E. Emerson (in litt.) ; and, indeed, the samples from South Africa are even
closer to those from East Africa than the samples of M. bellicosus from Nigeria are
to those from the Congo. Tumulitermes kibonotensis had entirely disappeared
from Sjostedt's work between 1924 and 1926. No type material of this species
has been found and it must be presumed lost. A neotype has thus been selected to
establish this synonymy.

Type Material. TANZANIA : Mbusine (6 "15' S., 38° E.), 29.viii.i888 (Fr. Stuhl-
mann), minor soldiers, major and minor workers, Hamb. ; other syntypes in
Stockh.

Other Material. CONGO : North Katanga, 1911 (Michell), type collection of
T. michelli von Rosen (Mus. Zool. Staatssammlung, Munchen), paratypes in Stockh.
and A.M.N.H. — Lubumbashi (ex-Elisabethville) : 1912, 3 vials (/. Bequaert),
1915-1916, 3 vials (Rimgoet), 1930 (Lamoral), 1948, 5 vials (A. E. Emerson), Terv.,
Stockh., N.C.I., A.M.N.H.— Karibwe R. (9° S., 27° E.), 8-io.iii.i947 (G. F. de
Witte), (Institut des Pares Nationaux du Congo et du Rwanda, Bruxelles) and
B.M.(N.H.). — Bukama, iv., Kundelungu Mts (9-10° S., 26-27° E.), ig.xii.ign
(/. Bequaert), Terv.

UGANDA : Mityana, x.i926 (H. Hargreaves), B.M.(N.H.) and A.M.N.H. ; Entebbe,
no date (0. John).

KENYA : Kwale, 15. vi. 1952 (P. B. Kemp}.

TANZANIA : Kilimandjaro (T. Paesler), holotype major soldier of T. goliath Sj. ;

TERMITES OF THE GENUS MACROTERMES

385

paratypes and paramorphotypes, A.M.N.H., (Berlin), Hamb., (Museum de Paris). —
Kibonoto (— Kibongoto), 27.iv.i9o6 (Y. Sjostedt}, NEOTYPE major soldier of
Tumulitermes kibonotensis Sj., Stockh. — m'Karamo, Massai (? Massai tribe), no date
(Pagani), Dar-es-Salaam, no date (Stuhlmann), Usambara, no dates (Brunnthaler ,
Sjostedt}, Stockh. ; Katesh, 18 & 31^.1957, 2 vials (Basilewsky & Leleup), Hamb. and
Terv. ; Kisarawe, 1903 (F. Eichelbaum}, Hamb. ; Tanga, 9^.1966 (D. H. Kistner),

FIGS 27-34. Macrotermes falciger, major soldier ; 27-30, head capsule and thoracic nota,
plan view ; 31-34, head capsule, side view.

386 J. E. RUELLE

own collection ; Tanga, 19. ix. 1924, between Ngare, Nairobi and Longido, 8.xi.i934,
Mburu (? Mbulu), 6.1.1935, 45 m. fr. Mbeya tow. Iringa, 12.1.1935 (H. Kirby],
A.M.N.H. ; Iringa, 11.1937, Arusha, 3i.viii.i95o (W. V. Harris], Arusha, 27.^.1961
(B. Hocking), Babati, Pienaars Heights, 20.11.1958 (collector unknown).

ZAMBIA : Mufulira, x.1949 (collector unknown), Broken Hill-Lusaka, 20.!.,
Kitwe-Ndola, 21,23,24,25.!., Ndola, 21.30.!., Lusaka, 19.1.1957, Choma, 13,14.1.1957
(W. G. H. Coatori), 26.1.1957 (G. M. Calvert), 6,9.viii., io.ix.i959 (E. N. Cooling),
N.C.I, and U. Lov. ; Lusaka, 22.xii.i966 (M. G. Bingham), U. Lov.

MALAWI : Somba (? Zomba), no date (Cameron), Stockh. ; Mlanje, i.viii.i929
(R. Boulton), A.M.N.H. ; Shire Highlands (B.M.(N.H.) Ace. No. 1914-291), Cholo,
I2.viii., Dedza, 7-ix., Fort Lister Gap (i5°49' S., 35°5o' E.), 2i.viii., 23 m. fr.
Kasungu on Lilongwe Rd, iS.ix., 10 m. fr. Mzimba on the Rumpi road, 28. ix.,
Songwe, i.x. 1953 (Sands 6- Wilkinson), Zomba, io.viii.i953 (Sands 6- Wilkinson),
no date (H. H. Johnston) .

MOZAMBIQUE : Boane (near Louren90 Marques), no date (P. C. Joubert), N.C.I. ;
Palma, I2.iv.i9i4 (A. Loveridge).

RHODESIA : Salisbury, no date, 2 vials (Marshall), Stockh., iv.i9i7 (R. W.
Tucker), 2o.vii., i.viii.i95o (G. C. Martin), N.C.I., I9.xii.i949 (G. H. Bunzli),
B.M.(N.H.) ; Selinde Mt., I3,i8.xii.i929 (R. Boulton), A.M.N.H. ; Bulawayo, 1903
(C. P. Lounsbury), I.R.S.N. ; Concession, 2O.V.I922 (R. Jack), N.C.I. ; Angwa, x.,
Matsikite, x., Matopos, 29.xi.i965 (M. G. Bingham), U. Lov. ; Mazoe, I4.xi.i9i7
(collector unknown), Gokwe Distr., 20. xi. 1962, 2.111.1964 (M. G. Bingham), Rusape,
3.111., Hartley, 2O.v., Trelawney, 3.viii.i949, Norton, 4.vii., 2i.viii., Marandellas,
26.x. 1950, Melfort, 17^.1951 (G. H. Bunzli).

REPUBLIC OF SOUTH AFRICA — (From now on, mention of the material kept in
the N.C.I., Pretoria, will for brevity's sake be limited to locality and accession No.)—
Type Material of Termes swaziae Fuller : several samples of the N.C.I, bear the label
' cotype ', viz. : Tzaneen F-35O, also in A.M.N.H., F-35I, F-353, also in B.M.(N.H.),
F-354 ; Barberton F-352 ; Ledzee F-85O ; Mokeetsi F-853, F-948. — Other Material.
Transvaal : Johannesburg, no date (R. Schwab], Hamb. ; Acornhoek Rail F-i,538,
Bushbuck Ridge TM-i,576, 3,500, 7,597, Duiwelskloof TM-n,340, Gravelotte
TM-n,775, Hectorspruit TM-ng, Komatipoort TM-5,i85, 6,899, Louis Trichardt
TM-7,i43, Nelspruit TM-5,i53, 5,168, 5,173, Ofcalaco TM-i3,346, Punda Milia
TM-6,764, Satara TM-6,758, Steelpoort F-i,6o7, The Downs TM-i3,349, White
River TM-5,i5i, Mokeetsi F-i,074, TM-ii5,n6,i2O,i99, 13,328, N.C.I. ; Chirinda
Mt, 25°23' S., 32°49' E., xii.i9o6 (D. Odendaal).

SWAZILAND : Type Material of M. usutu Fuller : Hlatikulu, 1920 and vii.i92i
(Pierce), type colony, N.C.I., syntypes in B.M.(N.H.). Other Material : 39 m. fr.
Bremeisdorp tow. Gollel, 22.^.1935 (H. Kirby), Hamb. and A.M.N.H. ; Stegi-
Bremersdorp TM-6,942, N.C.I.

A total of 177 nest series were examined, and all material is in the B.M.(N.H.)
unless otherwise indicated.

M. falciger is predominantly a woodland species. Its typical habitats in Zambia
and Rhodesia are the Brachystegia woodlands on hillsides and it seems to require

TERMITES OF THE GENUS MACROTERMES

387

very well-drained soils (M. G. Bingham, in litt.}. The workers occasionally leave
their mud galleries to forage in the open, even in broad daylight (Fuller, 1915 : 466 ;
Bingham and W. G. H. Coaton, in litt.},

This species is not the only Macrotermes that can be collected from huge, massive
mounds. Specific behaviour (Harris, 1956 ; 1961 : 35 ; Coaton, 19620) is not
involved here and the large termite hills, found to contain M. subhyalinus (Grasse &
Noirot, 1957, 1961), M. bellicosus, Pseudacanthotermes and other genera (own

MAP 2. Collecting localities of Macrotermes falciger.

388 J. E. RUELLE

observation) , in various parts of Africa, are, like those of M. falciger, the result of a
long story of successive recolonizations at the same site. The name ' goliath '
has been misleading.

It is still not clear whether the architectural potential of M. falciger, under similar
circumstances, differs from that of the partially sympatric M. subhyalinus. P. B.
Kemp (1955 : 130) found no differences between their nests in North-Eastern
Tanzania. Pictures published by Coaton (1962 : 68, 69) suggest that M. falciger
in Zambia builds low hummocks whereas M . subhyalinus erects tall, spired mounds.
This, however, was negated by the determination of the insects concerned.

More data are also wanted about the regular presence of a royal chamber. The
published evidence is inconclusive, for M. falciger (Sjostedt, 19074:, pi. 3 ; opp.
Fuller, 1915 : 466) as well as for M. subhyalinus (Grassd & Noirot, 1961 : fig. 14 ;
opp. Weidner, 1961 : 38). In comparison, M. bellicosus regularly builds such a cell ;
M. muelleri does not.

Finally, the swarming behaviour of M . falciger has already been described from
Tanzania, Kilimandjaro and Malawi, Somba (? Zomba) by Sjostedt (19000 : 95 ;
1926 : 84). In Zambia, this species emerges just after dark, M. subhyalinus around
midnight (M. G. Bingham, in litt.}.

Macrotermes herus (Sjostedt)
(Text-figs 38-44 ; Map 3)

Termes herus Sjostedt, 19140, : 79. Type-locality : ETHIOPIA, Dire Daua.
Macrotermes herus (Sjostedt) Snyder, 1949 : 213.

Imago. Head capsule dark reddish brown to black, no lighter area around fontanelle.
Postclypeus same colour as head, the median line not darker. Mandibles : base lighter than
postclypeus, tip black. Pronotum black, T-shaped dark red mark faintly visible in the middle.
Tibiae same colour as femora, abdominal sternites as dark as tergites. Wings smoky brown.

Posterior margin of head capsule nearly semi-circular ; scattered bristles between eyes and
ocelli, between ocelli and clypeus. Fontanelle protruding, surrounded by short setae, followed
anteriorly by small, rounded longitudinal ridge. Eyes flattened, small (H/E index about 3-3) ;
ocelli very small, distant from the eye by at least f of their major diameter. Postclypeus
relatively long, not prominent. Antennae ig-jointed, second segment longer than third by
about half the latter.

Pronotum not wider than head across eyes ; trapezoidal, front margin straight, sides convex
to straight, hind margin barely emarginate. Setae on margins, a few on disc also.

Hind tibia shorter than 4-95 mm.

Measurements (13 specimens from 4 localities) in millimetres.

Range Mean

Head width across eyes .... 3 -35-3 -77 3-48

Eye major diameter . .... 0-98-1-12 1-04

Ocellus major diameter .... 0-32-0-46 0-37

Ocellus minor diameter .... 0-20-0-37 0-28

Ocellus ex eye ...... 0-17-0-37 0-28

Median length of pronotum . . . i -76-1 -87 i -82

Maximum width of pronotum . . . 3 -34-3 -50 3-43

Length of hind tibia . .... 4-26-4-95 4-51

Length of fore wing ..... 31-00-35-00 33 -50

Maximum width of fore wing . . .•• 7-20-8-50 8-00

TERMITES OF THE GENUS MACROTERMES 389

Major soldier. Head capsule dark chestnut-brown ; labrum concolorous with head. Gula
darker than ventral genae. Mandibles black throughout. Antennae: first and second segment
as head capsule, 3rd to xyth dark brown. Thoracic nota dark brown, lighter in the middle.
Abdominal sternites lighter than tergites. Dorsal side of femora light brown ; ventral side
and tibiae dark brown.

Head rectangular, sides sub-parallel, faintly converging anteriorly. Fontanelle not very
prominent, but wide and relatively far behind anterior end of head capsule (i.e. situated at
about 35% of HL), followed anteriorly by a median ridge. Mandibles slightly incurved and
upcurved, length of left one about the same value as head depth. Antennae ly-jointed, third
segment 50% longer than second. A few scattered bristles on head capsule and postmentum,
practically none on thorax.

Pronotum with a transverse as well as a longitudinal axis of symmetry ; mesonotum narrower
than pronotum, sides rounded ; metanotum width between those of pro- and mesonotum, hind
margin flatly incurvate.

Measurements (50 specimens from 17 localities) in millimetres.

Range Mean
Head length ...... 4-93-6-10 5-44

Head width ...... 3-59-4-84 4-24

Head depth inc. gu la .... 2-38-3-22 2-78

Length of gula. . . 3-56-3-87 3 '75

Maximum width of gula i -30-1 -53 i -40

Minimum width of gula .... 0-86-1-18 i-oo

Length of left mandible .... 2-54-3-06 2-81

Length of hind tibia . .... 2-86-3-64 3-29

Maximum length of pronotum . . . 1-54-2-02 1-78
Maximum width of pronotum . . . 2-77-3-81 3-34

Minor soldier. Head smoky brown, paler beneath ; gula, however, darker than ventral
genae. Mandibles black, base dark brown. Antennae dark sepia-brown. Thoracic nota and
abdominal tergites sepia-brown. Abdominal sternites lighter than tergites. Tibiae and tarsi
same colour as femora.

Head elongate, sub-trapezoidal, sides straight to slightly convex ; fontanelle conspicuous,
followed anteriorly by a faint ridge. Mandibles long, slender, apex incurved. Thoracic nota
rather long, sclerotized, sides straight. Hind margins of pro- and mesonotum emarginate,
of metanotum straight.

Measurements (56 specimens from 17 localities) in millimetres.

Range Mean
Head length ...... 2-47-3-68 3-08

Head width ...... 2-07-3-15 2-56

Head depth inc. gula .... 1-42-2-11 1-69

Length of gula . ..... 1-89-2-45 2-11

Maximum width of gula .... 0-78-0-99 0-86

Length of left mandible .... 1-83-2-37 2-09

Length of hind tibia . .... 2 -07-2 -96 2 -48

Maximum length of pronotum . . . 1-00-1-40 1-16
Maximum width of pronotum . . . 1-62-2-55 1-98

Major worker. Similar to that of M. subhyalinus, but smaller ; head width (47 specimens
from ii localities) : range 2-02-2-75, mean 2-48 mm.

Variation. The darker specimens of this species have been collected at higher altitudes
(up to 7,800 ft) in Ethiopia. The backward location of the fontanelle is in fact too variable to
provide a specific distinction from the closely similar M. subhyalinus major soldiers, but it does
suggest a transition between the Ethiopian and the Indo-Malayan Macrotermes (of which six
species, cursorily examined at the B.M.(N.H.), had their fontanelle at between 38 and 53% of
HL ; the highest value found in M ' . herus — and in any Ethiopian Macrotermes — being 42).

390

J. E. RUELLE

The specimens listed below, when previously identified at all, had been labelled
in the collections examined as ' bellicosus ' (7 samples), ' natalensis ' (4), even ' herus '
(2). This suggests that M. herus is very close to M. subhyalinus and M. bellicosus.
However, to say nothing of the dark coloration (because it may simply be correlated
with altitude), the eyes and ocelli are definitely smaller and farther apart in M. herus
images than in those of M. subhyalinus, and the hind tibiae are also shorter. The
difference between soldiers is more a matter of size than of proportions, although
the head is somewhat more flattened and the mandibles less incurved in M, herus
major soldiers than in M. subhyalinus. With respect to M. bellicosus, the head-
depth, the ridge and grooves in front of the fontanelle and the sclerotization of the
thoracic nota usually provide a good distinction. While realizing that further
research may reduce M. herus to a local variant of M. subhyalinus, the evidence at

FIGS 38-44. Macrotermes herus, imago, soldiers. 38-39. Imago ; 38. head capsule
and pronotum, plan view ; 39, head capsule, side view. 40-43, Major soldier ; 40-41,
head capsule and thoracic nota, plan view ; 42-43, head capsule, side view. 44.
Minor soldier, head capsule and thoracic nota, plan view.

TERMITES OF THE GENUS MACROTERMES 391

present available is insufficient and M. herus is therefore considered as a distinct
species.

Since the disappearance of the type-specimen (Sjostedt Ace. No. 303) from the
Stockholm collections, another imago from the type-locality had been selected and
labelled as lectotype by Dr A. E. Emerson, but the designation has not been pub-
lished hitherto :

Type Material. ETHIOPIA : Dire Daua, no date (Hagenbeck), LECTOTYPE
winged female and paralectotypes in Stockholm (Sj. Ace. No. 273), other paralecto-
type in A.M.N.H. ; 1919 (A. Marchand), morphotype major soldier and para-
morphotypes in Stockholm (Sj. Ace. No. 282), other paramorphotype in A.M.N.H.

Other Material. NIGERIA : Northern Region : 4 mi. N. of Bida, 23.xii., 26 m.
fr. Abuja on Bida Rd, 24.xii., Diko (9°io' N., 7°o6' E.), 3i.xii.i956, 22 m. fr. Gombe
on Ture Rd, io.v.i957, 20 m. fr. Yandev on Makurdi Rd, 25.^.1958 (W. A. Sands).

SUDAN : Tozi (i2°40' N., 33°5o' E.), x.i96o, 7 vials (H. Schmntterer), Hamb. ;
Kaka, no date (/. Trdgdrdh), Stockh. ; Barakat, 15.^.1927 (F. G. S. Whitfield}.

ETHIOPIA : Dire Daua (other than type material), 2o.vi.i9ii (E. Wache), Hamb.,
no date (M. Rothschild), Stockholm, 1961 (B. G. Hill], B.M.(N.H.) ; Moulou (co-
ordinates ?), 1901 (de Zeltner), I.R.S.N. ; Nazareth (8°34' N., 39°i9' E.), no date
recorded, Bishoftu, 5, 6 & g.iv. (7 vials), Debrezed (8°5o' N., 38°5o' E.), 6.iv., 100
km S. E. of Addis Ababa, 9.^.1963, 2 vials (A E. and El. Emerson), A.M.N.H.

SOMALI REPUBLIC : Shimba Berris (9°io' N., 46°o8' E.) (or io°45' N., 47°i5' E.),
i6.iv.i957 (E. J. van Ingen).

KENYA : Nairobi, g.xi.igso (W. V. Harris).

PEMBA ISLAND : no date, 2 vials (Pakenham).

A total of 33 nest series were examined, and all material is in the B.M.(N.H.)
unless otherwise indicated.

The biology of M. herus is practically unknown. Alates have been collected
inside the nest, not far from Addis Ababa, in April.

Macrotermes ivorensis Grasse* & Noirot
(Text-figs 45-47 ; Map 4)

Macrotermes ivorensis Grass6 & Noirot, 1951 : 333. Type-locality : IVORY COAST, Adiopo-
doume (5°2o' N., 4°o8' W.).

Imago. The following notes are translated from the original description (illustrated, op.
cit. : 336). Head capsule dark reddish brown ; anteclypeus very pale. Head elongate,
sides tapering towards anterior end. Postclypeus small, inflated. Fontanelle small but
distinct. Eyes rather large ; ocelli circular and prominent, O-E equal to ocellus diameter.
Pronotum moderately sellate, shape almost triangular, anterior margin flat with a very shallow
median indentation.

Measurements in millimetres : Head width across eyes, 3-7 ; Eye diameter, i-o ; Ocellus
diameter, 0-45 ; width of pronotum, 3-52 ; length of hind tibia, 4-4. (The corresponding
figures for a male alate from Port Harcourt, Nigeria, are : 3-41, 0-98, 0-41, 3-18 and 4-0 mm.
It has not been possible to study the type-queen.)

Major soldier. Again translated from Grass6 & Noirot, op. cit. : 333. Head capsule reddish.

392

J. E. RUELLE

Mandibles black with reddish base. Labrum reddish, tip hyaline, without setae. Thoracic
tergites dark brown, lighter in the middle.

Head in plan view with sides distinctly converging towards anterior end ; posterior angles
rounded. Median region of gula considerably narrowed. Mandibles stout, left one thicker
than right. Antennae ly-jointed, first and third segment twice the second.

Pronotum trapezoidal, slightly indentated in middle of front and hind margin.

The following measurements (9 specimens from 4 localities ; in millimetres) have been taken
in the course of this revision :

MAP 3. Collecting localities of Macrotermes herus.

TERMITES OF THE GENUS MACROTERMES 393

Range Mean

Head length ...... 5 -09-5 -68 5 -39

Head width ...... 4-34-4-89 4-63

Head depth inc. gula .... 2-68-3-12 2-90

Length of left mandible .... 2-99-3-25 3-12

Length of hind tibia . .... 4-05-4-64 4-33

Maximum length of pronotum . . . i -64-1 -95 i -77

Maximum width of pronotum . . . 2-95-3-55 3-21

Minor soldier. Grass6 & Noirot's description : General coloration reddish brown, head
more reddish ; thoracic and abdominal tergites brownish ; legs and abdominal sternites
lighter.

Head narrowly oval ; fontanelle inconspicuous. Mandibles long and slender, almost straight.
Median region of gula not much narrowed. Labrum triangular, elongate, with slightly narrowed
base, hyaline tip conspicuous. Antennae i7-jointed, very long, extending backwards beyond
hind margin of metanotum by at least three segments.

Pronotum almost hexagonal, lateral angles barely rounded. Legs long and slender.

Measurements (own figures ; n specimens from 5 localities) in millimetres :

Range Mean

Head length ...... 2-05-2-50 2-21

Head width ...... i -46-1 -86 i -65

Head depth inc. gula .... 1-18-1-32 1*25

Length of gula (2 specimens) . . . 1-36 & 1-41

Maximum width of gula (id.) . . . 0-59 & 0-62

Length of left mandible i -59-1 -84 i -68

Length of hind tibia . .... 3 -2 1-3 -55 3-35

Maximum length of pronotum . . . 0-91-1-03 0-96

Maximum width of pronotum i -32-1 -59 i -43

The following notes should be added to the original description :

Soldiers. Head capsule definitely more setose than in M. muelleri ; major soldiers usually
lighter than those of M. nobilis ; minor soldiers with a head thicker and less elongate than that
of M. muelleri, the anterior third of the gula in side view diverging more from the underside
of the head capsule, the gula itself being proportionately wider (being also wider than the gula
of M. nobilis minor soldier).

Major worker. Similar to, but smaller than, those of M. muelleri and M. nobilis ; head
width (17 specimens from 4 localities) : range 2-21-2-67, mean 2-47 mm.

With regard to the figures published by Grass£ & Noirot, it will be noted that they
extend the upper range of the head length in major soldiers to 6 mm, of the head
width in the same to 5-15, and of the hind tibia in minor soldiers to 3-92. As
compared with the figures found in the course of this revision, this last result is not
too surprising, since another forest Macrotermes has yielded precisely the same range
for the similar segment in samples from the same locality (M. muelleri, mature
colonies from Mondongo, Congo : 3-26 to 3-97 mm ; the overall range for this
latter species being still larger).

Type Material. IVORY COAST : Adiopodoume", no date (Grasse 6- Noirot), paratype
from type colony, A.M.N.H. (Unpublished evidence indicates that the holotype and
morphotype are kept at the " Laboratoire d'Evolution des Etres Organises, 105,
boulevard Raspail, Paris 6eme ".)

Other Material : Yapo Forest Reserve, 24.viii.i953 (M. Luscher), own collection

394

J. E. RUELLE

and A.M.N.H. ; Labbe" Forest, 30 km N. of Abidjan, 30.111.1961 (C. Noirot), Noirot
Coll. No. 675 and B.M.(N.H.).

GUINEA : Nimba Mts, viii.-x.i946, 4 vials (M. Lamotte), Lamotte collection.

SIERRA LEONE : Kabui Hills, 7°57' N., ii°n' W., 13.1.1958 (W. Wilkinson).

NIGERIA : Eastern Region : Port Harcourt, 7-8^1.1957 (W. Wilkinson] \ 12 m.
fr. Enugu on Onitsha Rd, 2.111.1958 (W. A. Sands).

A total of 10 nest series were examined, and all material is in the B.M.(N.H.)
unless otherwise indicated.

M. ivorensis is a forest species ; a distinct queen cell has been found inside its
nest (Grasse" & Noirot, 1951 : 317). Swarming in Eastern Nigeria can occur in
early June, according to the collecting date of the sample from Port Harcourt.

Macrotermes lilljeborgi (Sjostedt)
(Text-figs 48-50 ; Map 5)

Termes Lilljeborgi Sjostedt, 1896 : 269. Type locality : CAMEROON, Kitta (4°3o' N., 9°o' E.).
Imago. Unknown.

Major soldier. Head reddish brown to black. Labrum same colour as head. Gula darker
than ventral genae. Mandibles black. Antennae dark chestnut brown, apical segments
yellow. Pronotum lighter than head, margins dark. Abdominal tergites darker than the

FIGS 45-47. Macrotermes ivorensis, soldiers. 45. Major soldier, head capsule and
thoracic nota, plan view. 46-47. Minor soldier : 46, head capsule and thoracic nota,
plan view ; 47, head capsule, side view.

TERMITES OF THE GENUS MACROTERMES

395

sternites, but lighter than thoracic nota. Legs pale to brownish yellow ; tibiae and tarsi not
darker than femora.

Head capsule conspicuously narrowed in front, in plan view with two shallow grooves diverging
from fontanelle to base of clypeus. Mandibles very long, evenly and moderately upcurved,
left one wider than right (Emerson's statement to the contrary (1928 : 442) must have been a
misprint). Antennae i7-jointed, third segment the longest, more than twice the second.
Fontanelle inconspicuous.

Pronotum with front margin deeply, hind margin shallowly emarginate. Legs rather thick.

Measurements (8 specimens from 3 localities) in millimetres.

MAP 4. Collecting localities of Macrotermes ivorensis.

396

J. E. RUELLE

Head length

Head width

Head depth inc. gula

Length of gula.

Maximum width of gula

Minimum width of gula

Length of left mandible

Range
6-26-7-10
5-45-6-17
3-10-3-72
4-09-4-55

i -34-i '59
o-86-i-oo
4-32-4-54

Mean
6-66
5-78
3-36
4-28
1-42
o-93
4'43

MAP 5. Collecting localities of Macrotermes lilljeborgi.

TERMITES OF THE GENUS MACROTERMES

397

Length of hind tibia .
Maximum length of pronotum
Maximum width of pronotum

5-31-6-45
1-91-2-13

5-95
1-99

3-77

Minor soldier. Head sepia-brown to black, ventral side usually not lighter than dorsal side.
Labrum same colour as head, with light dots around base of setae. Mandibles dark reddish
brown to black. Thoracic nota lighter than head capsule, darker than abdomen, the latter
brownish yellow.

Head in plan view sub-rectangular, sides faintly convex. Mandibles : left one wider than
right, median part already incurved (only tip in the right one). Antennae 17-jointed, third
segment equal to fourth, longer than fifth to seventh, again equal to 8th-i2th.

Front and hind margin of pronotum equally indentated in middle ; lateral angles acute,
antero-lateral sides distinctly more concave than postero-lateral. Mesonotum narrower than
pronotum, hind margin emarginate ; metanotum as wide as pronotum, hind margin flat.

Measurements (14 specimens from 5 localities) in millimetres.

Head length ......

Head width ......

Head depth inc. gula ....

Length of gula

Maximum width of gula .

Length of left mandible ....

Length of hind tibia. .

Maximum length of pronotum .
Maximum width of pronotum

Major worker. According to Sjostedt (igooa : 85), the head width of the major worker
ranges from 3 to 3-4 mm. The type collection yields a mean value of 3-22 mm ; other speci-
mens, however, are slightly below 3-0 mm.

Range

Mean

2-83-3-82

3-n

2-07-2-82

2-30

i -49-2 -oo

i -60

1-94-2-53

2-10

0-64-0-85

0-74

2-22-2-99

2-47

3-86-5-04

4-38

1-11-1-32

I-I5

1-62-2-25

1-77

48

FIGS 48-50. Macrotermes lilljeborgi, soldiers. 48. Major soldier, head capsule and
thoracic nota, plan view. 49-50. Minor soldier : 49, head capsule and thoracic nota,
plan view ; 50, head capsule, side view.

3Q8 J. E. RUELLE

The major soldier of M. lilljeborgi has very long mandibles. Even with the 4-14
mm measured by Emerson (1928 : 442), the whole range is still beyond that of any
other Macrotermes.

Sjostedt (19000 : 84, 91) mentions the deep indentation of the front margin of the
pronotum in the major worker. This has also been observed in the soldier caste ;
the segment, however, remains quite different from that of Pseudacanthotermes.

Type Material. CAMEROON : Kitta, ly.iii.iSgi (Y. Sjostedt), syntype soldier in
Stockholm (Sj. Coll. No. 212). Other syntypes in A.M.N.H., B.M.(N.H.) and
I.R.S.N.

Other Material. CAMEROON : Kunabembe (2°3o'-3°2o' N., I4°3o'-i5°3o' E.),
3.iii.i9ii (Arn. Schultze), Stockh. ; Nyong R., I3.xii.i949 (Birket-Smith 6- Dahl),
(Universitetets Zoologiske Museum, K0benhavn), and B.M.(N.H.) ; Sangmelima,
no date recorded (B. C. Z. Evans), A.M.N.H. ; Victoria, 8.1.1913 (F. Silvestri),
(Silvestri collection, Portici), and A.M.N.H.

NIGERIA : Eastern Region ; Old Calabar, c. 1900, collector unknown, B.M.(N.H.).

CONGO : Bamania, o°oi' N., i8°24' E., 1.1921 (H. Schouteden), Terv., Stockh.
and A.M.N.H. ; Eala, no date (P. Staner), Terv.

A total of 9 nest series were examined. Two samples, from Kribi (Mus. Berlin ;
Sjostedt, 19000 : 87) and San Benito River (Museum Paris ; Sjostedt, 1904 : 49),
could not be examined ; the determinations being thought reliable, the localities
have accordingly been reported on Map 5, above.

Neither Sjostedt (1896) nor Silvestri (1914) could find the nest, and the imago of
M. lilljeborgi is therefore unknown. Since this caste has not been collected, the
minor workers do not seem to accompany the foraging expeditions.

The geographical range of the species is thought to extend from the Cameroon to
primary forest regions of the Congo, near the Equator. The scanty material
suggests that other forest Macrotermes, esp. M. muelleri, are much more abundant
than M. lilljeborgi, or at least much easier to find.

Macrotermes mossambicus (Hagen) stat. n.
(Text-figs 51-64 ; Map 6)

Termes bellicosus var. mossambica Hagen, 1858 : 59-83 ; 118-121. Type-locality : MOZAM-
BIQUE.

Termes (Termes) Michaelseni Sjostedt, 19140 : 77. Type-locality : SOUTH-WEST AFRICA,
Okahandja, syn. n.

Macrotermes bellicosus (Smeathman) form kunenensis Fuller, 1922 : 73. Type-locality :
SOUTH-WEST AFRICA, Omango, syn. n.

Macrotermes bellicosus (Smeathman) form limpopoensis Fuller, 1922 : 73. Type-locality :
SOUTH AFRICA, Messina, syn. n.

Macrotermes bellicosus-tonga Fuller, 1927. Type-locality : SOUTH AFRICA, Upper Mkuzi
Drift, syn. n.
Imago. Head capsule dark chestnut-brown, lighter around fontanelle. Postclypeus same

colour as head, with dark median line. Mandibles with base slightly paler than postclypeus,

tip black. Pronotum same colour as head, with fairly conspicuous moth-shaped light mark

in the middle (other light marks rather indistinct). Abdominal tergites slightly, sternites

TERMITES OF THE GENUS MACROTERMES

399

distinctly lighter than thoracic nota. Tibiae darker than femora. Wings light yellowish
brown.

Vertex of head capsule moderately depressed, with small median longitudinal ridge, pilose
with light area around base of setae. Fontanelle prominent, surrounded by numerous setae.
Eyes large, sub-circular and prominent ; ocelli very large, short oval, distant from the eye by J
to J (exceptionally A) of their major diameter. Postclypeus moderately inflated, anterior
margin straight. Antennae ig-jointed, second segment the shortest, third about twice the
second, and longer than fourth.

Pronotum with conspicuous setae on margins and disc, antero-lateral corners acute, sides
straight, hind margin short.

Measurements in millimetres : $ holotype (dried specimen) — range and mean of 25 specimens
from 1 8 localities :

Head width across eyes
Eye major diameter ..
Ocellus major diameter
Ocellus minor diameter
Ocellus ex eye ...
Median length of pronotum
Maximum width of pronotum
Length of hind tibia
Length of fore wing ..
Maximum width of fore wing

Holotype Range Mean

3-41 3-29-3-74 3-50

1-23 1-14-1-33 1-20

0-50 0-45-0-60 0-52

0-40 0-32-0-47 0-39

o-io 0-10-0-23 0-15

1-68 1-63-1-87 1-77

3-32 3-09-3-70 3'43

4-40 4-26-4-99 4-54

33-oo 33-00-37-00 35-1°

8-50 8-20-10-00 9-30

Major soldier. Head capsule reddish brown to dark brown, ventral side usually same colour
as dorsal and gula not darker than ventral genae. Mandibles, although not uniformly coloured,
darker than head. Thoracic nota darker than head capsule, abdominal tergites paler than
thoracic nota.

Head capsule in plan view parallel-sided, hind margin flattened ; in side view thick, with
characteristically truncated forehead. Fontanelle protruding, followed anteriorly by two
conspicuous diverging grooves. Mandibles incurved and upcurved, left one hooked or sickle-
shaped, short, its length inferior to head depth. Hyaline tip of labrum with sides convex to
straight. Antennae i7-jointed, third segment shorter than twice the second. Lighter area
around base of setae, otherwise pilosity not distinctive.

Pronotum almost twice as wide as long, with antero-lateral sides evenly concave, front and
hind margin conspicuously emarginate in the middle ; sides of metanotum angular.

Hind tibia short, not exceeding 3-5 mm.

Measurements (171 specimens from 104 localities) in millimetres :

Head length .....

Head width .....

Head depth inc. gula

Length of gula .....

Maximum width of gula

Minimum width of gula

Length of left mandible

Length of hind tibia ....

Maximum length of pronotum .

Maximum width of pronotum

Minor soldier. Head capsule uniformly orange-brown, only sides of gula darker. Mandibles
black with reddish base. Antennae usually darker, without the reddish tinge of head capsule.
Thoracic nota brownish, margins darker than middle. Abdominal tergites lighter than, or

Range
4-26-5-68

Mean
5-°2

3-22-4-32

3-74

2-15-2-91
3-04-3-68

2-53
3-36

i -07-1 -37
0-78-1-04

1-23
0-88

2-15-2-91

2-40

2-47-3-45
1-33-1-81
2-50-3-68

2-97

1-58
3-10

400

J. E. RUELLE
sternites lighter than tergites.

Femora and tibiae whitish,

same colour as, thoracic nota
tarsi darker.

Head in plan view short ovate, rather wide. Fontanelle conspicuous, followed anteriorly
by a shallow ridge extending towards the clypeus. Mandibles broad, left one more incurved
than right, apical part only slightly more incurved than basal part.

Metanotum distinctly wider than mesonotum, sides rather angular than rounded.

Measurements (180 specimens from 76 localities) in millimetres :

63

64

FIGS 51-64. Macrotermes mossambicus, imago, soldiers. 51-52. Imago : 51, head
capsule and pronotum, plan view ; 52, head capsule, side view. 53-60. Major soldier :
53—56, head capsule and thoracic nota, plan view : 57-60, head capsule, side view.
61-64. Minor soldier, head capsule and thoracic nota, plan view.

TERMITES OF THE GENUS MACROTERMES 401

Range Mean

Head length ...... 2-22-3-35 2'79

Head width ...... 1-84-2-80 2-33

Head depth inc. gula i -23-1 -85 i -53

Length of gula . ..... 1-44-1-94 1-67

Maximum width of gula .... 0-68-0-87 0-77

Length of left mandible .... 1-64-2-27 1-92

Length of hind tibia. .... 1-85-2-57 2-21

Maximum length of pronotum . . . 0-91-1-34 i-n

Maximum width of pronotum . . . 1-55-2-36 1-95

Major worker. Head capsule reddish brown, in plan view posteriorly rounded ; head width
(126 specimens from 35 localities) : range 2-11-2-91, mean 2-54 mm.

Variation. In the images, Sjostedt (i9i4«) had noticed that the pronotum is wider than the
head across eyes ; this does not constitute a specific character. Neither is the fore wing always
wider in M. mossambicus than in M. subhyalinus. Otherwise, in this and in the soldier caste,
no consistent geographic variation has been found worth mentioning.

This species is similar to M. subhyalinus ; the ranges of quantitative characters
overlap broadly. However, in the images the pilosity of the head capsule and the
large size of the eyes and ocelli, in the major soldier the overall size, the curvature
of the left mandible and the profile of the head capsule, in the minor soldiers the
width and the angular shape of the metanotum usually allow a correct identification
at first glance. (The difference between mesonotum and metanotum width in
major soldiers is not a reliable character, although the metanotum is, more often
than not, wider than the mesonotum in M. mossambicus.}

In addition to specimens of M. michaelseni, the junior synonym, the material
listed below includes quite a few samples (from Malawi, Rhodesia and South Africa)
that had been identified by Fuller as ' M. bellicosus ' or a variety of it.

Type Material. MOZAMBIQUE : no locality, no date (Peters), $ holotype imago
Zoologisches Museum der Humboldt-Universitat zu Berlin.

Other Material. TANZANIA : Ruponda, 22. xi. 1950 (W. V. Harris}.

ANGOLA : Pembe, i6°53' S., i4°57' E., 8.ix.ig56 (A. Matos), Dundo and Hamb. ;
Caxiaxia, 8°56' S., 2O°38' E., iS.x.igdi (Champion), Dundo.

ZAMBIA : Kafue Township, 28.x. 1966 (M. G. Bingham), U. Lov. ; Kafue R.,
N. of Mazabuka, 17,18^.1957 (W. G. H. Coaton), Magoye, 4 vials, 17.1.1957 (E. N.
Cooling, W. G. H. Coaton), N.C.I.

MALAWI : Louwangwa Valley nr Tuchila, 4.ix.i956 (A. W. R. McCrae) ; 4 and
14 mi. N. of Nchalo Expt Station (i6°io' S., 34°53' E.), 3 vials, 14., Chikwawa,
14., 5 and 24 mi. fr. Ngabu on Chiromo Rd, 15., i and 17 mi. N. of Port Herald,
16., 8 and 12 mi. N. of Chiradzulu, 3 vials, 21., n mi. N. of Domasi on Namwera
Rd, 22., 3 mi. fr. Monkey Bay on Ft Johnston Rd, 24., 5 mi. fr. Ft Jonhston on
Monkey Bay Rd, 2 vials, 25., 2 mi. fr. Bilila on Ft Johnston Rd, 2 vials, 27., 20 mi.
fr. Ft Johnston on Zomba Rd, 28., i mi. E. of Lake Shirwa, 3i.viii., Namatulu Hill
(Zomba Distr.), i., 23 mi. fr. Benga on Salima Rd, 10., 7 mi. fr. Nkata Bay on
Ekwendeni Rd, 22.ix.i953 (W. A. Sands and W. Wilkinson).

MOZAMBIQUE : Porto Amelia, Hamb. (ace. No. 85, 1927) ; Zimbiti (Beira),
(P. A. Sheppard), Luabo, 24.viii.i957 (P. /. Cohen), Buzi R. (Beira), 7 vials,

402 J. E. RUELLE

1915, Xinavane and Umbuluzi R. (Lourerupo Marques), n, 14,18. xii. 1918 (Cl.
Fuller], N.C.I. ; Beira, I4.X.I952 (W. V. Harris), Vila Fontes, no date (H. Swale).

SOUTH WEST AFRICA : Okahandja, 1910 (G. Fock), 1911 (W. Michaelsen), type-
collection of Termes michaelseni, Hamb., Stockh., A.M.N.H., N.C.I. Omango
(co-ordinates unknown), G.xii.igig (Hartig), type collection of M. bellicosus kunenen-
sis, N.C.I., B.M.(N.H.) and A.M.N.H. Other Material : Onjatu (2i°2o' S.,
I7°2o' E.), viii.1909 (D. Hentschel), Stockh., Hamb., A.M.N.H. and N.C.I. ;
Outjo-Okaukuejo, i6.ix.i965 (/. L. Sheasby), N.C.I, and A.M.N.H. ; Okapchuri
(2i°53' S., i6°3o' E.), viii.-ix.ign (M. v. Rudno-Rudzinski) , Hamb. and Stockh. ;
Otjiwaronga, 1.^.1939 and 8.vi.i940, 3 vials (collector unknown), i6.xi.ig67 (W. G. H.
Coaton), Outjo, xii.i949 (v. d. Berg), Baynes Mts nr Otjinungwa, (i7°i7' S., 12 "27' E.),
viii.1956 (R. Story), Kalidona 100 mi. E. of Okahandja, 2 vials, 28.viii., Ongoro-
sengo (2i°i6' S., i8°o7' E.), 3i.viii., Etemba (2i°2o' S., 18° E.), 6., Okavango R.
10 mi. E. of Andara, I3.ix.i962 (W. G. H. Coaton), Katima Mulilo, 2 vials, 29^.1965
(A. Barnard), N.C.I.

Further records from S. W. Africa include 230 samples collected north of the line
Usakos-vSteinhausen, i.e. 22°3o' S., by a N.C.I, team (W. G. H. Coaton, G. F.
Pretorius, T. L. Sheasby) from September 1965 to May 1967. The localities have
been plotted on the distribution map (Map 6) and a detailed list is available at the
B.M.(N.H.).

BOTSWANA : Kabulabula, i7°5o' S., 24°58' E., 2O.vii.i93o (H. Lang), A.M.N.H. ;
Serowe, ix. & xi. 1922 (collector unknown), Mahalapye, n., 60 mi. N.W. of Serowe,
20., Tsukutsa (? Chukudu) Pan, 30^1.1924 (C. W. Mally), N.C.I.

RHODESIA : Gwaai R., 3 vials, 22., Shangani R., 28.vi., Kennedy (i8°5o' S.,
27°o8' E.), ig.viii., I7.ix.i922 (R.Jack), N.C.I. ; Rekomitjie (i6°07' S., 29°23' E.),
20.x., 22.xi.i964, Mzarabani (i6°i9' S., 3i°io' E.), 2.iv., Chibara (i6°35' S., 3i°38'
E.), ?.iv., Mashumbi (i6°09' S., 3O°33' E.), P.viii., Rimuka Siding (i8°23' S., 29°5o'
E.), 23.x. 1965 (M. G. Bingham), U. Lov. ; Zhombe (i8°3o' S., 29°2o' E.), 9.xii.i962
(M. G. Bingham).

REPUBLIC OF SOUTH AFRICA : Messina, 24.xi.igi6 (Cl. Fuller), type collection
of M. bellicosus limpopoensis, N.C.I., B.M.(N.H.) and A.M.N.H. ; Upper Mkuzi
Drift, I7.ix.ig22 (Cl. Fuller), type collection of M. bellicosus tonga, N.C.I. (2 vials),
B.M.(N.H.) and A.M.N.H. Other Material. Natal : Junction Inyomite, Pongola
and Usutu Rivers, 20.ix, nr Pongola R. (4 mi. S. of Ndumu Store), 22.ix.i922 (Cl.
Fuller), Lower Mkuzi Drift, 1.1924 (Collins), Mkuzi Game Reserve, 19.^.1947
(D. v. V. Webb], N.C.I. — Transvaal : Johannesburg, no date (R. Schwab), Hamb. ;
Middelwit-Ganskuil, 4.x. 1961 (G. F. Pretorius), N.C.I, and A.M.N.H. ; Omloop nr
Vetfontein (22°45' S., 29°io' E.), 26.^.1946 (P. J. de Lange), nr Balangani tow.
Komatipoort, I2.i., Punda Milia-Louis Trichardt, 3O.xii.i959, Pontdrift (22°i3'
S., 29°o8' E.), 3.i., Maasstroom (22°4o' S., 28°i5' E.), 5.i., Vleeschfontein-Zeerust,
8.i., Nietverdiend-Zeerust, 9.1.1960 (P. C. Joubert), Zeerust-Derdepoort, 2.,
Kiesel-Cumberland, 6., Buffelsdrif (23°4o' S., 26°55' E.), 2 vials, 5., Buffelsdrif-
Stockpoort and Stockpoort-Beauty, 6., Tom Burke-S wart water, 8., Pont Drift-
Alldays, n., Martinsdrift (23° S., 27°55' E.), 19., Tom Burke-Baltimore, 20.,
Buffelsdrif-Maraheki, 26.^.1964 (W. G. H. Coaton), Maraheki-Kranskop, 3.x. 1961,

TERMITES OF THE GENUS MACROTERMES 403

Vivo (23° S., 29°O3' E.), 16., Bochum-Tonash, 17., Tolwe-Maasstroom, 19.^.1964,
Distr. Soutpansberg, 4 vials, 19-22. viii. 1968 (/. L. Sheasby), Nietverdiend-Derde-
poort, 2 vials, 2., Waterpoort-Louis Trichardt, 2 vials, 16., Steenbokpan-Hound-
slow, 3 vials, 25.iv.i964 (Coaton 6- Sheasby), N.C.I.

SWAZILAND : Hlatikulu, 1920 (/. H. Pierce), Lebombo Flats, 5.vi.i92i (CL Fuller],
Ranches Ltd (co-ordinates unknown), no date (Bartle), N.C.I.

A total of 386 nest series were examined. Where no museum of deposit is
indicated, the material is in the B.M.(N.H.).

MAP 6. Collecting localities of Macrotermes mossambicus.

4o4

J. E. RUELLE

Climatic data (Coaton, private communication) indicate that M. mossambicus
can live in fairly dry sub-regions : Okahandja (annual rainfall range, 350-400 mm),
Usakos (150-200 mm). In Zambia and Rhodesia, this species is common only in
the Zambezi valley area, has not been found at altitudes higher than 1200 m, and
tolerates poorly drained, alluvial soils (M. G. Bingham, in litt.). In South Africa,
it has not been found at higher latitudes than the north-eastern corner of Natal.

The mounds have been described as tall, hard, conical by Michaelsen (in Sjostedt,
1926 : 88) ; Weidner (1961 : 42) records a height of 3-5 m from Angola ; Bingham
(in litt.) has measured in Zambia a mound 3 m high and 3 x 2 m at the base. No
queen cell has hitherto been described, but Fuller's collecting data (1922 : 73)
seem to imply its presence.

Alates have been caught near Rekomitjie, swarming soon after dark in the last
days of November. Little else is known on the behaviour and ecology of M.
mossambicus.

67

FIGS 65-68. Macrotermes muelleri, imago
(with details of light marks)

65-66, head capsule and pronotum, plan view
67-68, head capsule, side view.

TERMITES OF THE GENUS MACROTERMES 405

Macrotermes muelleri (Sjostedt)
(Text-figs 65-76 ; Map 7)

Termes Mulleri Sjostedt, 18980 : 205. Type-locality : GABON, Agoncho.

Termes amplus Sjostedt, 1899 : 35. Type-locality : CONGO, Umangi [syn. Termes gabonensis

Sjostedt, 1907 : 238 ; ssp. rev. Macrotermes Mullen var. ampla, Sjostedt, 1926 : 78 ; syn.

Macrotermes mulleri, Snyder, 1949 : 215].
Macrotermes Mulleri (Sjostedt) Sjostedt, 1926 : 76.
Macrotermes muelleri (Sjostedt) ; Weidner, 1961 : 32.

(Sjostedt first changed from Mulleri to gabonensis (19000 : 92) because of Termes Mulleri von
Ihering (1887). After shifting the species to Macrotermes, he took the species-name Miilleri
again (1926). He was entitled to do this, since Termes Mulleri von Ihering lacks a description
and therefore is not valid.)

Imago. Head capsule dark sepia-brown to black, lighter beneath. Postclypeus same colour as
head, with inconspicuous lighter median line. Mandibles lighter than head, only inner margin
black. Pronotum with four light marks on the limit between anterior and posterior lobes.
Abdominal sternites lighter than tergites and femora lighter than tibiae. Wings smoky brown,
with darker veins.

Head in plan view oval, strongly tapering towards anterior end, with a very few, scattered
bristles. Fontanelle depressed, inconspicuous and not surrounded by small hairs. Eyes
relatively small, not prominent ; ocelli broadly oval, distant from the eyes by a little less than
their major diameter. Postclypeus almost flat. Antennae ig-jointed, third segment distinctly
longer than others.

Pronotum more than twice as wide as long, but usually narrower than head across eyes ;
front margin straight to concave, barely emarginate in the middle, hind margin emarginate,
a few short hairs on margins.

Measurements (32 specimens from 13 localities) in millimetres :

Range Mean

Head width across eyes . . . 3-80-4-50 4 -09 (°)

Eye major diameter .... 1-09-1-38 1-21

Ocellus major diameter . . . 0-41-0-50 0-47

Ocellus minor diameter . . . °'33-°'43 °'38

Ocellus ex eye ..... 0-28-0-52 0-43

Median length of pronotum . . . 1-58-1-87 1-71

Maximum width of pronotum . . 3-48-4-52 3-96

Length of hind tibia .... 4-45-5-61 4-88

Length of fore wing .... 29-20-38-00 33'3o

Maximum width of fore wing . . 7-50-9-00 8-40

(°) After Wasmann (1911) : 5 mm. The specimen could not be

re-determined and has probably been lost.

Major soldier. Head capsule dark reddish brown, lighter beneath, gula darker than ventral
genae. Mandibles uniformly black. Thoracic nota about as dark as head capsule, but without
reddish tinge ; pronotum with a light cross-shaped mark on the middle and two fairly distinct
light dots on each side, mesonotum with light mark in the middle extending to the hind margin.
Abdominal tergites not lighter, sternites distinctly lighter than thoracic nota.

Head capsule in plan view pear-shaped ; fontanelle slightly protruding, followed anteriorly
by a faint median ridge. Mandibles slightly longer than half the length of head capsule ;
rather evenly incurved, left one more incurved and wider than right one. Sides of hyaline tip
of labrum concave. Antennae i7-jointed, 3rd segment about twice as long as 4th. Pilosity
sparse.

Pronotum distinctly narrower than head capsule and relatively long, lateral corners acute.

406 J. E. RUELLE

Sides of mesonotum angular ; of metanotum parallel, but more or less truncated towards hind
margin. Metanotum wider than mesonotum.

Hind tibia longer than 4-5 mm in most cases.

Measurements (224 specimens from 38 localities) in millimetres :

Range Mean

Head length 5.35-6-60 6-01

Head width ...... 4-52-5-90 5-21

Head depth inc. gula .... 2-76-3-50 3-10

Length of gula 3 -45-4 -32 3-81

Maximum width of gula .... 1-07-1-33 1-19

Minimum width of gula .... 0-65-0-86 0-73

Length of left mandible .... 3 -00-3 -55 3-29

Length of hind tibia . . . . . 4-26-5-55 4-89

Maximum length of pronotum . . . 1-72-2-00 1-86

Maximum width of pronotum . . . 2-91-3-61 3-25

Minor soldier. Head capsule uniformly dark red-brown, gula not darker. Mandibles dark
brown to black, antennae lighter than head. Pronotum same colour as head capsule, other
thoracic nota and abdominal tergites a little lighter. Sternites distinctly lighter than tergites,
legs (without any conspicuous difference between femora, tibiae, tarsi) about the same colour
as the sternites.

Head in plan view quadrate — elongate, angles rounded ; fontanelle slit-like, inconspicuous.
Labrum large, reaching about half the length of the mandibles ; the latter long and slender.
Gula in side view evenly convex.

Thoracic nota : pronotum diamond -shaped, front margin indentated, hind margin incurved ;
mesonotum trapezoidal, hind margin incurved ; metanotum rectangular, about ^ wider than
mesonotum, hind margin straight.

Length of hind tibia nearly twice the head width.

Measurements (138 specimens from 31 localities) in millimetres :

Range Mean

Head length ...... 2-11-3-15 2-64

Head width ...... 1-53-2-31 1-90

Head depth inc. gula .... 1-15-1-49 1-33

Length of gula ..... 1-58-2-11 1-81

Maximum width of gula ; 0-59-0-72 0-62

Length of left mandible .... 1-63-2-18 1-91

Length of hind tibia . .... 3-26-4-35 3-70

Maximum length of pronotum . . . 0-86-1-18 1-04

Maximum width of pronotum i -20-1 -85 i -53

Major worker. Head capsule reddish brown, darker above antennal sockets ; in plan view
paraboloid, maximum width near external articulation of mandibles ; this width (89 specimens
from 17 localities) : range 2-50-2-93, mean 2-73 mm.

Variation. In spite of Text-fig. 70, which is given as an example, the light marks on the
pronotum are not species-specific : they are simply more conspicuous in dark-coloured speci-
mens throughout the genus. The major soldiers of this species may be darker or lighter ;
in the minor soldiers, the difference of coloration between thoracic and abdominal tergites
may be nil to distinct. Some imagos (Emerson, 1928) have their pronotum wider than the head,
but this is not the rule. The antennae may be i8-jointed in the soldier caste, both the minor
(Weidner, 1961 : 33) and the major (Emerson, 1928 : 443). In the latter, the metanotum has
always been found wider than the mesonotum.

Lastly, some samples from the Western part of the Congo (Text-fig. 72) are smaller than
average ; no distinct subspecies, however, could be recognized.

The de-alated imagos of M. muelleri resemble those of M. vitrialatus ; their eyes, however,

TERMITES OF THE GENUS MACROTERMES

407

are usually bigger, their ocelli more elongate and their pronotum proportionately wider. The
mean index pronotum length/pronotum width in the major soldier is the highest of the genus.
The index hind tibia length/head width in minor soldiers, as studied by Grass6 & Noirot (1951 :
332), has been checked with the figures now available, and found slightly lower than previously
thought : 1-95 (2-0). For comparison, M. bellicosus yields a value of i-oi, M. vitrialatus 1-44.

Type Material. GABON : Agoncho (probably 10° E. on the Equator), ix.-x.i874
(R. Buchholz), major and minor soldiers, workers (Museum Greifswald,) syntypes in
Stockh. and A.M.N.H. CONGO : Mukimbungu (inc. Boko, Madzia), 1904-1906
(K. E. Laman), morphotype and paramorphotype images, Stockh., Terv., A.M.N.H.

Other Material. CAMEROON : Batouri, 1935 (F. G. Merfield], B.M.(N.H.).

FIGS 69-76. Macrotermes muelleri, soldiers. 69-72. Major soldier, head capsule and
thoracic nota, plan view (70, with details of light marks). 73-76. Minor soldier : 74,
head capsule and thoracic nota, plan view ; 75-76, head capsule, side view.

4o8 J. E. RUELLE

CENTRAL AFRICAN REPUBLIC : Boukoko nr M'Baiki, vii.i948 (C. Noirot), own
collection and U. Lov.

CONGO (Brazzaville) : Ngoko (Gravot), Mate'le' (o°3i' N., i6°38' E.), 1924 (Karls-
son), Stockh.

CONGO : Umangi, ix., xi.i8g6 (E. Wilverth), type collection of T. amplus Sj.,
major soldiers in I.R.S.N., syntypes in Stockh. Other Material : Lisala, ig.ix.i966
(/. E. Ruelle), U. Lov. and B.M.(N.H.) ; Basongo, vii.igai (H. Schoutederi), Boma
Sundi (5°2o' S., I2°5o' E.), 27-29. viii. 1913 (/. Bequaert), Kisangani (ex : Stanley-
ville), no date, 3 vials (Arrhenius, H. Kohl), 29^.1933 (Vrijdagh), 1948 (A. E.
Emerson), Terv., Stockh., A.M.N.H. ; Yalosemba (2°35' N., 2i°5o' E.), 2i,23.ix.i966
(/. E. Ruelle), U. Lov., Terv., N.C.I. ; Yambata, ii.-iii.i9i4, 3 vials (de Giorgi),
Oshwe [not ' Obswe '], xii.i9i3 (/. Maes [not ' Mais ']), Mayumbe (c. 5° S., 13° E.),
no date (De Briey [not ' De Bricy ']), Bikoro [not ' Bikaro '], iii.i92i, Mongende
[not ' Mongenge '] (2°o6' S., i6°2o' E.), 22.iv.i92i (H. Schouteden), Terv., Stockh. ;
Medje, ig.ix.igio (Lang-Chapin), Terv., A.M.N.H. ; Mpese, no date (Cooreman),
8.viii.i959 (J. E. Ruelle), I.R.S.N., U. Lov. ; Inga (5 "30' S., J-4°3o' E.), 2Q.iv.,
Imbela (5°54' S., I7°o8' E.), 2i.viii.iQ59 (/. E. Ruelle), U. Lov., A.M.N.H. ; Banalia,
7.xii.i9i3 (J. Bequaert), Barumbu-Simba (i°i5' N., 23°29' E.), v.1927 (/. Ghes-
quiere), Boende, 1934 (Dubois), Buta, 1912 (Hutereau), Karawa, 1936 (Wallin),
Lokandu (2°34' S., 25°44' E.), V.IQ39 (Maree), Lusambo, 1898 (E. Luja), Moerbeke
(5°3o' S., I4°55' E.), no date (collector unknown), Mongbwalu, vii.1938 (Scheitz)
and iii.i939 (R. Lepersonne), Walungu (2°4o' S., 28°5o' E.), 1938 (Hautmann),
Moto (3°O2' N., 29°3o' E.), io.xi.i923, Tora, 1920, I3.vi., 12. x. 1926, 4 vials (L.
Burgeon), Ganda Sundi (4°47' S., I2°54' E.), ix.i92O, Mushie, vi.i92i, Mauda
(4°05' N., 27°4i' E.), iii.i925, Kunungu (2°o6' S., i6°26' E.), 1937 (H. Schouteden),
Terv. ; Bambesa (3°23' N., 25047' E.), no date (Vrijdagh}, I.R.S.N. ; Lukolela
(i°io' S., I7°n' E.), 2.vi.i948 (A. Anberg), Camp Putnam (i°23' N., 28°3o' E.),
ii.v., Yangambi, 28-30^.1948 (A. E. Emerson), Batama, i7.ix.i9O9, Akenge
(2°55' N., 26°5o' E.), x.1913 (Lang-Chapin), A.M.N.H. ; Takundi, 19.^.1964
(G. Mathot), Kitona (5°28' S., i7°42' E.), ig.viii.igsg, Mbata Kiela (5°i5' S., i2°55'
E.), 1.1.1965, Mondongo (2°io' N., 2i°io' E.), 13, 29-ix., Boumbu (2°4o' N., 2i°5o'
E.), 25. ix. 1966 (J. E. Ruelle), U. Lov.

SUDAN : Lado (? Lado Mt), no date (Hanolet), I.R.S.N.

ANGOLA : Dundo, 3.1.1961 (A. de Barros Machado), Dundo.

A total of 104 nest series were examined.

This species is the most abundant of the forest Macrotermes ; its distribution
has already been discussed above. In view both of its habitat and of the extensive
collections carried out in Nigeria and in southern regions of Africa, it is unlikely
that further research will reveal significant differences from the present geographical
range.

The nests have already been described (Grasse" & Noirot, 1951 ; Weidner, 1961
after de Barros Machado) ; but neither a queen cell nor even a physogastric queen
could be found. An earlier account (E. Luja, quoted in Wasmann, 1911) is un-
reliable, since the collector appears to have mixed M. bellicosus and M. muelleri
in his field notes. Recently (1966), reproductive imagos were collected in the

TERMITES OF THE GENUS MACROTERMES

409

Congo (Mondongo) from two mounds, 2 -5 and 3 -o m high ; in both cases the queen
was found in an ordinary nest chamber (well above ground level) and the male
escaped detection. This, with previous fruitless efforts, and the literature cited,
support the conclusion that M. muelleri usually does not enclose its royal pair in a
distinctive chamber.

The circular or semi-circular leaf cuttings, brought home from the foraging
expeditions and illustrated among others by Grass6 & Noirot (1951 : 315), are found

MAP 7. Collecting localities of Macrotermes muelleri.

410

J. E. RUELLE

not only with the forest Macrotermes, but with M. vitrialatus as well. The swarming
habits of M. muelleri are still unknown.

Macrotermes natalensis (Haviland)
(Text-figs 21, 77-90 ; Map 8)

Termes natalensis Haviland, 1898 : 383. Type locality : REPUBLIC OF SOUTH AFRICA, Natal

[Estcourt, cf. Harris, 19660].

Macrotermes natalensis (Haviland) Fuller, 1922 : 73.
Macrotermes natalensis var. durbanensis Fuller, 1927 : 135. Type locality : REPUBLIC OF

SOUTH AFRICA, Durban, syn. n.

What has been said about M. bellicosus could be repeated here to help sorting out
the misidentifications in the literature, such as those of Tragardh (1904 : 6), Was-
mann (1911 : 92), Sjostedt (1913 : 363) (1915 : 12), Emerson (1928 : 447), Grasse"
(19370, 1944), etc.

Imago. Head capsule chestnut to smoky brown, with lighter area around fontanelle ; post-
clypeus lighter, with dark median line ; base of mandibles intermediate between head capsule
and postclypeus, tip black. Pronotum slightly lighter than head, with a sometimes con-
spicuous moth-shaped mark. Abdominal tergites and sternites not distinctly lighter than
thorax. Tibiae darker than femora. Wings smoky.

Head in plan view rather short (i.e. the width across eyes is great in relation to the distance
between the anterior margin of the postclypeus and the posterior margin of the head capsule),
its hind margin regularly convex. Fontanelle in a depression, not very conspicuous, but
surrounded by short setae and followed anteriorly by a median carina ; the head otherwise
sparsely pilose. Eyes sub-pentagonal, neither large nor particularly prominent. Ocelli sub-
circular to elongate, distant from the eye by about £ of their major diameter. Postclypeus
wide (slightly wider and less inflated than in bellicosus}. Antennae ig-jointed, 3rd segment
twice as long as 2nd.

Pronotum ' subsemilunar ' (Haviland), front and hind margins not or barely emarginate,
lateral corners rounded, sides convex.

Hind tibia short.

Measurements in millimetres : $ lectotype — range and mean of 89 specimens from 47
localities :

Head width across eyes
Eye major diameter .
Ocellus major diameter
Ocellus minor diameter
Ocellus ex eye .
Median length of pronotum
Maximum width of pronotum
Length of hind tibia .
Length of fore wing .
Maximum width of fore wing

Lectotype Range
3-61 3-03-3-68
i-io
0-42

o-33
0-24
i -80

3'59
4-27

34*0
9-0

0-88-1-16

0-33-0-53
0-23-0-37
0-18-0-32
i -61-1 -94

3-H-3-59
3-81-4-77
33-0-38-0
8-0-10-0

Mean

3-43
1-03
0-41
0-30
0-24
1-79

3-34
4-n

34-6
9-0

Major soldier. Head capsule yellow to dark ferruginous brown, apical part darker than
base, ventral side as often as not same colour as dorsal side ; gula, however, darker than ventral
genae. Mandibles usually black throughout. Thorax lighter than head, margins of nota
smoky brown. Tibiae barely, if at all, darker than femora ; tarsi darker than tibiae.

Head capsule in plan view ' ovate-quadrate ' (Haviland) to ' horseshoe-shaped ' (Fuller),
sides moderately convex ; fontanelle more conspicuous than in M. bellicosus, i.e. wider and
slightly protruding ; frontal grooves shallow, but sometimes similar to those of M. subhyalinus.

TERMITES OF THE GENUS MACROTERMES

411

Sides of hyaline tip of labrum straight to convex. Mandibles long, upturned from the base
(not really upcurved, as is the case with M. mossambicus) , moderately and fairly regularly
incurved, left one not much wider than right. Antennae 17- to i8-jointed, ist segment longer
than 3rd, 3rd longer than 4th, 2nd the shortest. Pilosity variable, rather sparse on head and
thorax, often dense on abdomen ; lighter area around base of setae rarely conspicuous.

Anterior and posterior margins of pronotum equally indented, lobes rounded, antero-lateral
sides notched ; meso- and metanotum nearly as wide as pronotum, sides rounded, hind margin
of metanotum concave to straight ; the thoracic nota, on the whole, distinctly more chitinized
than in M. bellicosus.

Measurements in millimetres
128 localities :

morpholectotype — range and mean of 203 specimens from

Head length.

Head width ....
Head depth inc. gula
Length of gula
Maximum width of gula .
Minimum width of gula .
Length of left mandible .
Length of hind tibia
Maximum length of pronotum
Maximum width of pronotum .

Morpholectotype

4-97
4-00
2-65
3-12
1-30
0-94
2-89
3-oi
1-67

3-25

Range Mean
4-13-6-21 5-15

3-26-4-89

4-13

2-16-3-15

2-65

3-12-3-56

3-35

1-17-1-32

1-22

0-91-1-07

0-98

2-52-3-14

2-85

2-60-3-65

3-07

I -43-1 -90

1-69

2-55-3-74

3-19

Minor soldier. Head capsule yellow to light chestnut-brown, lighter beneath, gula darker
than ventral genae ; mandibles very dark red, base lighter. Thorax yellowish brown, often
with outline darker. Abdominal sclerites slightly paler than thorax ; femora lighter than
abdomen, tibiae same colour, tarsi darker.

Outline of head capsule in plan view similar to that of major soldier, although shorter and
more rounded. Fontanelle conspicuous, but not protruding. Left mandible more incurved
than right, and a little more near the tip than in basal part ; in ventral view without any distinct
serration of the inner edge.

Sides of thoracic nota rounded, metanotum in most cases not distinctly wider than mesonotum.

Measurements (171 specimens from 99 localities) in millimetres :

Head length ......

Head width ......

Head depth inc. gula ....

Length of gula ......

Maximum width of gula ....

Length of left mandible ....

Length of hind tibia .....

Maximum length of pronotum .
Maximum width of pronotum .

Major worker. 'Head castaneous, epistoma convex ' (Haviland). Posterior margin in plan
view nearly semi-circular (text-fig. 21). Head width (63 specimens from 23 localities) : range
2-05-2-65, mean 2-31 mm.

Variation. Statistics on the variability of some quantitative characters can be found in
Warren (1909, 1916). The ' natalensis — durbanensis ' variety, although somewhat larger than
average, is not really different from the ' natalensis s. str.', ' nat. transvaalensis ' and other forms
described by Fuller ; even specimens from South West Africa cannot be distinguished from
those collected on the other side of the Kalahari desert.

Range

Mean

2 -49-3 -68

3-02

2 -03-2 -99

2-49

i -39-1 -97

1-69

1-69-2-47

1-94

0-75-0-94

0-82

i -68-2 -42

2-IO

1-86-2-67

2-28

0-94-1-40

1-16

i -62-2 -49

2-00

4I2

J. E. RUELLE

85

FIGS 77-90. Macroiermes natalensis, imago, soldiers. 77-78. Imago : 77, head
capsule and pronotum, plan view ; 78, head capsule, side view. 79-86. Major soldier :
79-83, head capsule and thoracic nota, plan view ; 84-86, head capsule, side view.
87-90. Minor soldier : 87-89, head capsule and thoracic nota, plan view ; 90, left
mandible, inner margin.

TERMITES OF THE GENUS MACROTERMES 413

This species has long been confused with M. bellicosus. The major soldiers
nevertheless present a distinctive pattern in the dimensions of the left mandible,
the hind tibia and the thoracic nota ; such differences were implied in Dr Emerson's
unpublished comment on a sample of M. bellicosus from Sierra Leone (above).
Also, the size of the minor soldiers is larger in M. natalensis. The imagos are
usually separated by the pilosity and the width of the head capsule, the diameter
of the eye (and the distance between eye and ocellus) and the length of the hind
tibia ; the index length/width of the pronotum is less reliable.

A certain similarity with M. subhyalinus prompted Fuller's opinion (1915 : 458)
on the probability of M. natalensis being a synonym of ' M. bellicosus ' ; more
recently, Noirot (1960) identified as ' Bellicositermes bellicosus ? ' a sample of Macro-
termes from the Outeniqua Mts (Cape Province). This tentative determination
could not be checked, but in view of the geographic distribution found, and of the
measurements taken, in the course of this revision, the doubtful sample most
probably belongs to M. natalensis. As a matter of fact, this species has rarely been
collected together with M. subhyalinus, but can easily be recognized from the
latter in such cases by the mandibles of the major soldiers.

A lectotype imago and a morpholectotype major soldier have been selected from
Haviland's type material ; they are listed below with some of the collecting localities.

Type Material. REPUBLIC OF SOUTH AFRICA : Natal, Est court, 19.^.1894
(G. D. Haviland), $ LECTOTYPE imago, morpholectotype major soldier from type
colony No. 37, Museum of Zoology, Cambridge ; topotypes, various dates (Haviland),
A.M.N.H., B.M.(N.H.), I.R.S.N., N.C.I., 1895 (Trdgdrdh), Stockh.

Other Material. ZAMBIA : Kitwe, 1.1956 (E. N. Cooling), Choma, 10-13, 16.1.1957,
8.viii.i959, n vials (W. G. H. Coaton, E. N. Cooling), N.C.I, and U. Lov.

MALAWI : Chiromo, xi.i9o6 (K. Fricke), Hamb.

MOZAMBIQUE : Lourengo Marques, i6.xii.i9i8 (Cl. Fuller) and I4.xi.i945 (E. H.
Botha), n.ii.i935 (H. Kirby), ij.x.igfr (W. V. Harris], N.C.I., A.M.N.H.,
B.M.(N.H.) ; Inhambane, 1917 (W. Keyes), N.C.I. ; Porto Amelia, 1928 (collector
unknown), Hamb.

SOUTH WEST AFRICA : Ombahowe (co-ordinates unknown), I4.x.i9i9 (Hartig),
Kalidona, 100 mi. E. of Okahandja, 25,26,29,30^1. and 3,4,5.ix., Etemba distr.
Okahandja, 4 vials, 6.ix.i962 (W. G. H. Coaton), Rietfontein, 24.^.1963 (F. Gaerdes),
Windhoek, 12.^.1964 (/. E. Beveridge), Rehoboth to Kalkrand, 4.x. 1964 (/. L.
Sheasby), N.C.I.

Further records from the same territory : 45 localities, 120 vials, ^.1965 to
xi.i967 (W. G. H. Coaton, G. F. Pretorius, ]. L. Sheasby), N.C.I. — These localities are
all between 18° and 25° S., 16° and 2i°30' E. ; a detailed list is available at the
British Museum (Natural History).

RHODESIA : Selinde Mt, io,i5.xii.i929 (R. Boulton), A.M.N.H. ; Melfort and
Salisbury, 1950, Norton, 1951 (G. H. Bunzli), B.M.(N.H.) ; Bulawayo, xii.iSgS
(McDonald), Salisbury, 2i.iii.i95o (Bunzli), N.C.I. ; Matopos, 23.xi.ig65 (M. G.
Bingham), U. Lov.

REPUBLIC OF SOUTH AFRICA : Durban, xi.i9i6 (Cl. Fuller, C. P. v. d. Merwe),

4i4 J. E. RUELLE

type collection of M. natalensis var. durbanensis Fuller, N.C.I. ; syntypes in
B.M.(N.H.) and A.M.N.H. — Other Material. Cape Province : Lusikisiki, 23.x. 1956,
Cathcart, 28.1.1958, Kei Mouth, I3.ix.i96i (W. G. H. Coaton), Harding, ly.xii.igsg,
Vryburg, 1.1960 (P. C. Joubert), Mafeking, 5., Stella, 6.xii.i958 (H. Niemari), Kim-
berley, iv.i9i3 (H. J. Power), Kuruman, 7. x. 1961 (G. F. Pretorius), Mount Frere,
I5.X.I962 (/. L. Sheasby), N.C.I. — Natal : Port Richmond, Port Shepstone, 3.111.1935
(H. Kirby], A.M.N.H. ; Weenen, no data, B.M.(N.H.) ; Pietermaritzburg, i

MAP 8. Collecting localities of Macrotermes natalensis.

TERMITES OF THE GENUS MACROTERMES 415

vi.igiS (R. Braun), Port Natal, 23.^.1893 (collector unknown) , Hamb. ; Appelbosch
(29°23' S., 3Q°35' E.), no date (Ljungqvist), Howick Falls, no date (Trdgdrdh),
Stockh. — Orange Free State : Vierfontein, 22,23.11.1915 (F. Thomsen), Hertzog-
ville, 4.vlii.i96i (/. J. C. Nel), Bloemfontein, 28., 29., Bothaville, 29., Vredefort,
29. x. 1962 (/. L. Sheasby), N.C.I. — Transvaal : Pretoria, 1914, 1915 (Cl. Fuller),
N.C.I., B.M.(N.H.), vStockh. (29 more nest series in the N.C.I.) ; Johannesburg,
no date (R. Schwab), Hamb. (2 other samples in the N.C.I.) ; Pietersburg, 15-
I7.iv.i935, 5 vials (H. Kirby), A.M.N.H. (9 other samples in the N.C.I.) ; Louis
Trichardt, 25, 26. xi. 1916 (Cl. Fuller), Komatipoort, n., 13. xi. 1958, i6.xi.ig59
(P. C. Joubert), Rustenburg, 26.ix.i96i, Maraheki (24°37' S., 2y°32' E.), 26.^.1964
(W. G. H. Coaton), N.C.I.

In addition to this material, 249 nest series from 143 localities (Cape Province,
20 localities ; Natal, 53 ; Orange Free State, 7 ; Transvaal, 63) have been examined.
A detailed list is available at the B.M.(N.H.) ; the samples are kept in the N.C.I.

SWAZILAND : Ezulweni (26°26' S., 3i°io' E.), 22. ix., I7.X.I92O (C. Buchanan),
Mbabane (26°2o' S., 3i°o8' E.), 8.vi.i956, 2 vials (/. H. Grobler), Stegi, 23.1.1957
(P. C. Joubert), Balegane (26°io' S., 3i°2o' E.), 22., Piggs Peak, 23.x.i96o (W. G. H.
Coaton), N.C.I.

A total of 551 nest series were examined.

This species is tolerant both of dry climates (some collecting localities in S.W.
Africa have an annual rainfall range of 550 to 150 mm) and of cold seasons ; no
other Macrotermes has been found in the Cape Province and the Orange Free State.

The nests have been described by Fuller (1921) and Coaton (1947) among other
authors ; the reproductive images are usually enclosed in a distinct chamber.
Swarming takes place in the second half of November, the alates emerging immedia-
tely after dark (Fuller, 1915) .

Macrotermes nobilis (Sjostedt)
(Text-figs 91-95 ; Map 9)

Termes nobilis Sjostedt, igooa : 90 ; nom. n. for T. speciosus Sjostedt, 1899 : 35 [preoccupied by

T. speciosus Haviland, 1898]. Type-locality : CAMEROON, Johann-Albrechtshohe.
Termes (Macrotermes) nobilis Sjostedt ; Holmgren, 1912 : 29.
Macrotermes speciosus (Sjostedt) Sjostedt, 1926 : 78 [unjustified restoration of preoccupied

name].

Macrotermes nobilis (Sjostedt) Snyder, 1949 : 216.

(In his last revision, Sjostedt applied the same procedure to M. muelleri and to M. speciosus ;
it was invalid for the latter, since T. speciosus had been well described by Haviland.)
Imago (previously undescribed). Head capsule very dark chestnut-brown. Postclypeus
same colour as head, median line not conspicuous. Mandibles with reddish base, tip and inner
margin black. Labrum yellowish brown. Pronotum dark sepia-brown, margins black, four
light marks near anterior margin. Abdominal tergites same colour as thoracic nota, sternites
lighter. Legs sepia-brown, with femora lighter than coxae and tibiae.

Head in plan view wide oval, pilosity insignificant. Fontanelle not protruding, incon-
spicuous, without hairs around, but with a small light dot in front of it. Eyes sub-circular,
ocelli elongate, their distance from the eye equal to their major diameter. Postclypeus more
than three times wider than long, not very prominent. Third antennal segment not longer
than first.

416 J. E. RUELLE

Pronotum width nearly i\ times its median length, hind margin broadly emarginate.

Measurements of the morphotype queen in millimetres :

Head width across eyes . . . . . 4-13

Eye major diameter . . . . . 1-18

Ocellus major diameter . . . , . 0-48

Ocellus minor diameter ..... 0-38 '

Ocellus ex eye ...... 0-48

Median length of pronotum i -70

Maximum width of pronotum . . . . 4-10

Length of hind tibia ..... 4'?6

Major soldier. Head capsule dark brown to black, with a small lighter area around fontanelle ;
slightly lighter beneath, gula darker than ventral genae. Mandibles black throughout ;
antennae lighter than head capsule. Thoracic nota same colour as head ; T-shaped light mark
and two lateral light dots on pronotum ; mesonotum with a median light mark restricted to
anterior half. Abdominal tergites sepia-brown, sternites distinctly lighter. Legs inter-
mediate in colour between abdominal sternites and tergites.

Head in plan view trapezoidal, with very low median ridge expanding from fontanelle towards
clypeus ; hyaline tip of labrum lancet-shaped ; mandibles incurved and upturned, left one
barely wider than right. Antennae i8-jointed, third segment the longest. Sides of pro- and
mesonotum rounded, of metanotum at right angles with hind margin, the latter faintly convex.
A few scattered setae on head capsule.

Hind tibia relatively short, about 4-25 mm.

Measurements (21 specimens from 7 localities) in millimetres :

Range Mean

Head length ...... 5-58-6-26 5-98

Head width ...... 4-63-5-29 4-95

Head depth inc. gula .... 2-83-3-25 3-07

Length of gula . ..... 3-58-4-07 3-86

Maximum width of gula i -27-1 -43 i -33

Minimum width of gula .... 0-75-0-88 0-80

Length of left mandible .... 2 -64-3 -06 2 -87

Length of hind tibia . .... 3-92-4-51 4-26

Maximum length of pronotum . . . i -69-1 -82 i -77

Maximum width of pronotum . . . 3'°4-3'35 3'23

Minor soldier. Head sepia-brown, slightly lighter beneath. Mandibles black, base same
colour as head ; antennae brown, barely lighter than head. Thoracic and abdominal tergites
same colour as antennae, abdominal sternites and legs lighter.

Head in plan view ovate-quadrate, in side view with convexity of gula nearly parallel to
that of ventral genae ; fontanelle slightly prominent ; mandibles long and slender ; antennae
17-jointed, 3rd and 4th segment equal.

Front margin of pronotum shallowly emarginate ; metanotum c. ^ wider than mesonotum.

Measurements (20 specimens from 8 localities) in millimetres :

Range Mean

Head length . .... 2-32-3-23 2-53

Head width ...... 1-68-2-32 1-83

Head depth inc. gula i -27-1 -59 i -34

Length of gula. ..... 1-50-1-80 1-62

Maximum width of gula .... 0-63-0-72 0-66

Length of left mandible .... 1-64-2-57 1-82

Length of hind tibia . .... 3-41-4-58 3-67

Maximum length of pronotum . . . 0-95—1-14 1-04

Maximum width of pronotum . . . 1-32-1-71 1-49

417

Major worker. Head same colour as minor soldier, with a conspicuous white dot around the
fontanelle ; in plan view nearly semi-circular, the maximum width (19 specimens from 5
localities) : range 2-52-2-88, mean 2-65 mm.

This species is closest to M. muelleri. The head capsule of the imago is less
elongate, the ocelli more elongate, the postclypeus proportionately wider, the third
antennal joint shorter, the pronotum proportionately shorter and the index hind
tibia length/head width smaller. It should be noted, however, that the truly
specific differences can not be asserted before the discovery of new material. In
the major soldiers, the most obvious differences between the two species are the
head shape and the lengths of the left mandible and of the hind tibia. The head of
M. nobilis is narrower and less convergent, its mandible and hind tibia are definitely
shorter. In addition, the pattern of light marks on the mesonotum seems different
from that found in M. muelleri. The minor soldiers cannot be safely recognized ;
the gula in M. nobilis is proportionately wider than in M. muelleri and the difference
between the widths of meso- and metanotum is smaller ; but the left mandible and
the hind tibia are here unreliable as taxonomic characters.

Type Material. CAMEROON : Johann-Albrechtshohe (4°4o' N., g°2o' E.), 1895
(L. Conradt), (syntype soldiers in Berlin Museum), other syntypes in Stockh. and

FIGS 91-95. Macrotermes nobilis, imago, soldiers. 91-92. Imago : 91, head capsule
and pronotum, plan view (with details of light marks) ; 92, head capsule, side view.

93. Major soldier, head capsule and thoracic nota, plan view. 94-95- Minor soldier :

94, head capsule and thoracic nota, plan view ; 95, head capsule, side view.

418 J. E. RUELLE

A.M.N.H. — Morphotype imago. CONGO : Yamangi (2°25' N., 2i°5o' E.), 22.ix.i966
(/. E. Ruelle), U. Lov. (coll. No. VI 61) ; soldiers from same colony in B.M.(N.H.).

Other Material. CAMEROON : Lolodorf, I2.vii.-2.xii.i895 (L. Conradt),
B.M.(N.H.), A.M.N.H. ; Bipindi, no data, A.M.N.H.

GABON : Belinga Forest nr Makokou, 29^.1962 (/. Deligne), B.M.(N.H.).
CONGO : Yalosemba (2°35' N., 2i°5o' E.), 2i.ix.ig66 (/. E. Ruelle), U. Lov.,

MAP 9. Collecting localities of Macrotermes nobilis ; including Ayos (3°53' N., I2°3i' E.),

after Grasse and Noirot (1951).

TERMITES OF THE GENUS MACROTERMES 419

Terv. ; Mondongo (2°io' N., 2i°io' E.), 28.ix.i966 (/. E. Ruelle], U. Lov. ; Camp
Putnam (i°23' N., 28°3o' E.), 19^.1948, 3 vials (A. E. Emerson), A.M.N.H.
A total of ii nest series were examined.

The nest structure and fungus combs of M. nobilis have been described from the
Cameroon (Grasse & Noirot, 1951). A note by J. Deligne with his sample from
Belinga Forest (Gabon) states : ' clay hummock '. Observations made while
collecting the imago in a swampy area of primary forest confirm that the nests are
low, dome-shaped, barely 50 cm high with a diameter exceeding 2 m ; which is
strikingly different from the tall, mamillated mounds of M. muelleri in the same
forest. (Each species has, incidentally, its own vernacular name). No distinctive
queen cell was observed and the king could not be caught. The fungus combs
are as figured by Grasse & Noirot (1951, PI. II, fig. 15 [not 14]).

Macrotermes subhyalinus (Rambur) sp. rev.
(Text-figs 96-122 ; Map 10)

Termes subhyalinus Rambur, 1842 : 307. Type-locality : SENEGAL.

Termes bellicosus Smeathman ; Hagen, 1858 : 109 [misidentification].

Termes bellicosus subsp. sansibarita Wasmann, 1897 : 158. Type-locality : ZANZIBAR [syn.
M. subhyalinus (as T. bellicosus), Sjostedt, 1900 : 100].

Termes tumulicola Sjostedt, 1899 : 34. Type-locality : DAHOMEY, Gross Popo, syn. n.

Bellicositermes jeanneli Grass6, 1937 : 71. Type-locality : KENYA, Kalodeka [syn. M. sub-
hyalinus (as B. bellicosus) Grass6 & Noirot, 1961 : 323].

As with M. bellicosus, the above synonymy contains only entries of nomenclatural
significance. To list all the misidentifications that have followed Hagen's publica-
tion would be beyond the scope of this revision. Suffice it here to quote some
taxonomic papers (Silvestri, 1914 : 12, under the name of goliath ; Grasse, 1937^,
1944 ; Harris, 1936, 1948, 1957, 1963 ; Weidner, 1956, 1961), and to state that any
citation of the species-name bellicosus under the generic headings of Amplitermes ,
Bellicositermes or Macrotermes in the literature and not included in the material
examined or in the references cited in this revision, should be referred to M. sub-
hyalinus.

Neotype, $ imago. Head capsule chestnut-brown ; postclypeus lighter than head, with a
thin, dark, not very conspicuous median line. Pronotum same colour as head, a light T-shaped
mark in the middle and two light dots near the antero-lateral corners ; abdominal tergites
same colour as pronotum, sternites lighter. Legs less reddish-tinged than head capsule, tibiae
slightly darker than femora. Wings yellowish brown.

Head capsule in plan view showing a distinctly depressed, semi-circular area around fontanelle;
fontanelle itself protruding, surrounded by short setae, followed anteriorly by a narrow median
ridge towards clypeus ; in side view vertex of head capsule evenly convex, rather than hog-backed
(opp. bellicosus). Eyes big, not very prominent, rounded. Ocelli slightly elongate, distant
from the eye by about ^ of their major diameter. Postclypeus moderately inflated. Antennae
(in other specimens than the neotype) ig-jointed, 3rd segment the longest, and and 4th equal.

Front margin of pronotum convex, hind margin weakly concave, antero-lateral corners
acute.

Measurements in millimetres : neotype — range and mean of 145 specimens from 63 localities :

420

J. E. RUELLE

102

FIGS 96-102. Macrotermes subhyalinus, imago, major soldier. 96-100. Imago : 96-98,
head capsule and pronotum, plan view ; 99-100, head capsule, side view. 101-102.
Major soldier : head capsule and thoracic nota, plan view.

TERMITES OF THE GENUS MACROTERMES

421

Neotype

Range

Mean

3-42

3-12-3-96

3-48

1-22

I -OO-I -36

1-17

0-49

0-36-0-67

0-50

o-33

0-27-0-46

o-35

0-16

0-08-0-35

0-19

i-75

i -64-2 -02

1-83

3-15

2-95-4-I3

3-40

4-80

4-18-5-29

4-69

32-00

29-00-43-00

36-40

g-oo

8-00-10-50

9-30

Head width across eyes

Eye major diameter ....

Ocellus major diameter

Ocellus minor diameter

Ocellus ex eve .....

Median length of pronotum

Maximum width of pronotum

Length of hind tibia.

Length of fore wing ....

Maximum width of fore wing

Major soldier. Head red-yellow to reddish brown, usually lighter beneath, sides of gula
darker than ventral genae. Antennae darker than head capsule. Mandibles black with
reddish base. Thoracic nota : margins dark brown, centre yellowish brown. Rest of body
same colour as thorax, but abdominal sternites lighter than tergites and tibiae darker than
femora.

Head capsule in plan view rectangular, slightly narrowed in front, with two grooves from
the fontanelle to the articulations of the mandibles ; in side view, rather thick, forehead more
or less truncated. Mandibles stout, left one slightly wider and more incurved than right,
its length very nearly equal to head depth. Pilosity scattered, a little denser on gula. Sides
of hyaline tip of labrum usually convex. Antennae ly-jointed, 3rd segment about twice the
and.

Pronotum large, its length more than half its width, front and hind margin equally emarginate,
lateral corners rounded, setae longer than on head capsule ; mesonotum a little narrower than
pronotum ; metanotum more often than not narrower than mesonotum, its hind margin flat
to concave. The three nota well sclerotized.

Measurements in millimetres : morphotype (only a head capsule) — range and mean of 456
specimens from 163 localities :

Head length .....

Head width .....

Head depth inc. gula

Length of gula ....

Maximum width of gula

Minimum width of gula

Length of left mandible

Length of hind tibia.

Maximum length of pronotum .

Maximum width of pronotum

Minor soldier. Head capsule yellowish brown to chestnut-brown, lighter than that of major
soldier, anterior outline darker, especially between antennal sockets and upper mandibular
condyles ; gula darker than genae. Mandibles black, base dark red. Antennae smoky brown,
darker than head. Thorax darker or lighter than head capsule, the outline of the nota being
usually darker than the middle. Abdominal sternites same colour as tergites and lighter than
the thorax. Femora and tibiae same colour as abdomen (tibiae often darker), tarsi darker.

Head in plan view quadrate-elongate, sides nearly straight. Fontanelle usually conspicuous,
sometimes protruding and followed anteriorly by a faint median ridge. Mandibles slender,
incurved and a little upcurved. Pilosity very sparse on head capsule, somewhat more abundant
on labrum and body, setae darker than cuticle. Inner margin of left mandible not serrate.

Pronotum with sides fairly straight from rounded antero-lateral corners to hind margin ;
both front and hind margins emarginate. Sides of meso- and metanotum rounded, the latter
slightly wider than the former.

Morphotype Range

Mean

6-36 4-84-7-70

6-49

4-90 3-48-6-45

4-98

3-15 2-41-4-13

3'20

3-23-4-90

4-27

1-21-1-68

1-48

0-89-1-36

I'll

3-10 2-37-3-72

3-23

2-69-4-58

3-75

1-50-2-52

1-97

2-9I-4-57

3-83

J. E. RUELLE

107

108

FIGS 103-114. Macrotermes subhyalinus, major soldier. 103-108, head capsule and
thoracic nota, plan view ; 109-114, head capsule, side view.

TERMITES OF THE GENUS MACROTERMES 423

Measurements (439 specimens from 146 localities) in millimetres :

Range Mean

Head length ...... 2-67-4-45 3-57

Head width ...... 2-31-3-55 2-87

Head depth inc. gula .... 1-52-2-24 1-87

Length of gula. ..... 1-81-2-66 2-32

Maximum width of gula .... 0-81-1-01 0-92

Length of left mandible .... 1-85-2-90 2-41

Length of hind tibia . .... 2-19-3-35 2-78

Maximum length of pronotum i -04-1 -55 i -33

Maximum width of pronotum . . . 1-86-2-75 2-36

Major worker. Head capsule rich chestnut-brown ; in plan view, sides slightly convex,
posterior margin rounded (Text-fig. 22). Head width (335 specimens from 114 localities) :
range 2-18-3-27, mean 2-75 mm.

Variation. The imagos are smaller in West Africa (head width, 3-15-3-45 mm) than in East
Africa (3-30-3-87), the specimens from Central Africa being intermediate ; the wings, in this
last sub-region, are longer : 35-0-40-0 mm (East : 30-0-43-0, this upper limit being excep-
tional ; West : 29-0-35-0). No variation clines in coloration or pilosity could be determined.
The sculpturing of the head capsule as it appears in plan or oblique view is one of the most
constant characters. No imagos were available from South-West Africa.

The largest major soldiers have been found in Central Africa, the smallest in South-West
Africa (mean head length of 28 specimens from this sub-region, 5-55 mm). The index head
depth/length of left mandible is more often inferior to i in Central Africa (80% of the specimens)
than elsewhere (West and East Africa, 40%) : in other words, the head capsule appears flatter
and/or the left mandible longer in Central Africa. In 182 randomly selected specimens from

118

120

122

FIGS 115-122. Macrotermes subhyalinus, minor soldier, head capsule and thoracic nota,

plan view.

424 J. E. RUELLE

82 localities, the index mesonotum width/metanotum width has been found greater than i in
115 cases : in other words, the mesonotum is often, not ' nearly always ' wider than the metano-
tum (Grasse & Noirot, 1961). There are likewise many exceptions to the character ' hind margin
of metanotum not concave '.

The largest minor soldiers have also been found in Central Africa, but they are only slightly
larger than those of West Africa.

The images of M. subhyalinus usually have longer but fewer setae on the head
capsule than those of M. bellicosus ; the T-shaped mark on the pronotum is often
more conspicuous and the ocelli are bigger and more elongate ; the major soldiers
of both species can be recognized by the overall size, the sculpturing of the forehead
and the thoracic nota ; the minor soldiers mainly by the size and the left mandible ;
the major workers by head shape and colour. The images of M. falciger, usually
larger than those of M. subhyalinus, have proportionately smaller eyes and longer
tibiae, the depressed area around the fontanelle also differs in shape ; major soldiers
of falciger are usually darker and are furthermore characterized by their huge
thoracic nota, their wide head, thick antennae and conspicuous pilosity of the gula ;
the minor soldiers have longer tibiae and more incurved mandibles. The case of
M. mossambicus has already been discussed.

The identification of specimens from some collecting areas : Ubangi-Uele-
Uganda (bellicosus), Katanga-Zambia (falciger), South- West Africa (mossambicus),
where M. subhyalinus has been found with the species given in brackets, can still
be difficult. In the absence of any reliable evidence of inter-specific hybridization,
this is attributed to the specific variability.

While meaning to complete Smeathman's description, Hagen (1855-1858) mistook
specimens of M . subhyalinus for M. bellicostis. Most of the material examined by
him — notably the Imhoff collection — has not been found in the European institu-
tions ; but some of Rambur's and Latreille's specimens that had been passed to
the Selys Longchamps collection (Desneux, 1915) were studied at the I.R.S.N.,
Bruxelles. The next available name for the species described by Hagen was T.
subhyalinus Rambur (not T. capensis Latreille, 1804, since the latter author, mis-
takenly quoted by Snyder (1949 : 209), was not describing a new species). The
holotype has been lost, but the only other Macrotermes from the same country in
the same collection belong to that species : hence the neotype and morphotype that
have been selected and are listed below.

Types or syntypes of T. tumulicola Sjostedt and Bellicositermes bellicosus var. rex
Grasse* & Noirot have also been examined.

Type Material. NEOTYPE. SENEGAL : locality illegible, coll. No. 5, male
imago. Morphotype : coll. No. 27, head capsule of major soldier. Both at the
Institut Royal des Sciences Naturelles, Bruxelles.

Other Material. SENEGAL : Cayor (i5°o5' N., i6°3o' W.), v.i895 (Chaper),
Stockh. ; Thies, 26.ix.i9i2 (F. Silvestri), (Silvestri collection, Portici) and A.M.N.H.

GAMBIA : Abuko (i3°24' N., i6°39' W.), io.ix.i966 (W. A. Sands).

PORTUGUESE GUINEA : Bissao, 1896 (H. Traun), Hamb. and Stockh.

GUINEA : Conakry, 3o.xi.i892 (H. Brauns), no date (Chevalier), 8.viii.i9i2 (F.
Silvestri), Hamb., Stockh., (Silvestri coll., Portici) and A.M.N.H. ; Kindia, date not

TERMITES OF THE GENUS MACROTERMES 425

recorded (F. Silvestri), (Silv. coll., Port.) and A.M.N.H. ; Los Islands, no date
(Serand), Stockh.

SIERRA LEONE : Njala (8°o6' N., I2°O5' W.), I4.iv.ig28 (H. Hargreaves), 24.viii.
1930 (E. Hargreaves}, 24.11.1947 (F. A. Squire], 25.-29.1.I955 (W. V. Harris],
A.M.N.H., B.M.(N.H.), N.C.I.

MALI : Kayes, no date (Mineur], Stockh.

IVORY COAST : Adiopodoume" (5°2o' N., 4°o8' W.), 8.vi.i952 (collector unknown],
45 km N. of Se'gue'la, no date (M. Luscher], A.M.N.H.

GHANA : Peki (6°33' N., i°4i' W.), 1.1927 (G. S. Cotterell], A.M.N.H. ; Ho, 1892
(Rossmann], Sekondi, 6.iv.i9ii (C. Manger], Hamb. ; Accra, no date and 8.vi.i926
(A. W. J. Pomeroy], 30 mi. N.E. of Accra, 111.1958 (R. Hamilton], 28 m. fr. Tamale
on Bolgatanga Rd, I., 35 m. fr. Tamale on Yendi Rd, 4., 20 m. fr. Bolgatanga on
Bawku Rd, 8., 9 m. fr. Bolgatanga on Navrongo Rd, 9., 29 m. fr. Navrongo on
Tumu Rd, 13., Paga, 14., 9 mi. N. of Lawra on Nandom Rd, 18., 6 mi. N. of Wa on
Lawra Rd, 19.111.1959 (W. A. Sands}.

TOGO : Ane"cho, 21. x. 1893 (H. 0. Schmidt], Hamb. and Stockh. ; Noepe, 15. xi. 1905
(M. Otto], Misahohe, 29. xi. 1910 (collector unknown] , Hamb.

DAHOMEY : Gross Popo (6°ig' N., i°57' E.), 23.11.1897 (F. Martinsen], type
collection of T. tumulicola Sj., Hamb.

NIGERIA : Western Region : Akure, 11.111.1933 (D. P. W.), A.M.N.H. ; Agega
(? 6°4o' N., 3°2o' E.), 9.1.1912 (E. Ballard], Ikoyi, ? 1934 (collector unknown],
Olokemeje, 8., Agodi (Ibadan), io.xii.i957 (W. A. Sands], Akure, 20. x. 1956, Lagos,
25. xi. 1957 (W. Wilkinson}. Eastern Region : Okigwi, 1955 (B. J. MacNulty],
47 m. fr. Enugu on Onitsha Rd, 31.1.1957 (W. Wilkinson}. Northern Region :
Birnin Kebbi, I2.vii.i959 (Kuhlow], Hamb. ; Sokoto, no date (Gaillard), Stockh. ;
43 mi. E. of Kano, Gaya R., 17., 25 m. fr. Sokoto on Gusau Rd, 25., 35 mi. N. of
Sokoto on Illela Rd, 26.xi., 27 m. fr. Zungeru, 19., 15 m. fr. Kaduna R. on Mokwa-
Bida Rd, 22., 12 m. fr. Abuja on Minna Rd, 29.xii.i956, 5 m. fr. Jos on Bykuru Rd,
6., Bauchi, 12., Afaka Forest Reserve (io°3o' N., 7°o' E.), 28. ii., Kwei 4 mi. W. of
Jos-Panshin Rd near Heipang, 29.!!!., Gombe, 9., 18 m. fr. Yola on Numan Rd, 14.,
34 m. fr. Yola on Jalingo Rd, 14., Yola-Faro R. 0-5 m. fr. Faro R., 15., Tiba-
Gangoro, 25., 35 m. fr. Yola on Biu Rd, 28., 15 mi. N. of Biu on Damaturu Rd,
29.v., Wulgo on L. Chad, 2., 50 m. fr. Maiduguri on Ft Lamy Rd, 3^1.1957, Gboko,
26.11.1958 (W. A. Sands}.

CENTRAL AFRICAN REPUBLIC : Bossembele", 10^1.1948 (Grasse <§• Noirot], type
collection of B. bellicosus var. rex Gr.-N., (own collection) and A.M.N.H. ; Lobaye
(3°4o' N., i8°o' E.), 1965 (/. M. Pasteels], own collection ; Bania, Nola, no date
(G. Eriksson], Stockh. ; Zemio, 5 and 7-iii., Bombo (4°5o' N., 22°3o' E.), lo.iii.,
Bangassou, 12.111.1948 (TV. A. Weber], A.M.N.H.

CONGO : Luebo, ix. 1921 (H.Schouteden],^Qiv., Stockh. and A.M.N.H. ; Niangara,
xi.i92o (Lang-Chapin), 1931 (coll. unknown), Terv., A.M.N.H., N.C.I. ; Luluabourg,
20.xii., 3 vials, 2i.xii.i9i3, 2 vials, 22. xi. 1921, 6 vials (P. Callewaert], lo.iii.-
2O.iv.j_939, 7 vials (/. /. Deheyn], Lubumbashi (ex : Elisabeth ville), 1921 (Poppe],
ix.i92i (Devroye), Luanza, xi.i9i2 (de Paoli], Terv., Stockh. ; L. Nyanza, no date,
Mauda (4°o5' N., 27°4i' E.), 2.1v., Mahagi, 10 and 23^.1925 (H. Schouteden],

426 J. E. RUELLE

Ishwa (2°i2' N., 3i°io' E.), 2O.ix.i934, Kasenye, 7^.1935 (H. J. Bredo), Watsa,
vii.igao, Tora, i6.v.i926 (L. Burgeon), Abok (2°i6' N., 3O°59' E.), no date, Niarembe
(2°i4' N., 3i°07' E.), 7-V.I935 (Ch. Scops), Albertville, 1928 (Lejeune), Bambesa
(3°23' N., 25°47' E.j, 14.^.1937 (Vrijdagh), Doruma, 1927 (/. Walkiers), Irumu,
vii.i937 (J. Ghesquiere), Kanda-Kanda, 1935 (Luxen), Kanzenze, viii.i93i (G. F.
De Witte), Kasongo, no date (Van Diest), Kiabukwa, 3.^.1932 (P. Quarre), Luashi,
iii.1936 (Freyne), Nyangwe, ^.1924 (J. Henrard), Ukesere (i°55' N., 3O°32' E.),
21.11.1929 (A. Collart), Van Kerkhovenville (3°2i' N., 29^2' E.), no date, 2 vials
(De Greeff}, Terv. ; Kinda, no date (Miiller), I.R.S.N. ; Garamba (4°io' N., 29 "40'
E.), no date (Lang-Chapin), Uele-Bomokandi Junction (= Bambili), 2.11. , Bunia,
ii.-iii.i948 (N. A. Weber), A.M.N.H. ; Kamina, 2o.xii.i959 (A. Bouillon), Katanda
(6°23' S., 23°55' E.), 27.vii.i963 (/. E. Ruelle), U. Lov.

SUDAN : Torit, 5.viii.i962 (H. Schmutterer) , no date (Weber Coll. No. 1461), Hamb.,
A.M.N.H. ; Tozi (i2°4o' N., 33°5o' E.), 5^11.1959 and i., 2.x.i96o, Dumo (10° N.,
25°30' E.), 29.xii.i959, Yei, 6.vii., Tash (n°3o' N., 30° E.), i.xi.i962 (H. Schmutterer),
Hamb. ; Khor el Affin, no date, 3 vials (Ebner), Stockh. ; Imatong Mts, 24^11.1939
(N. A. Weber), Kagelu, i4.iv., Azza Forest (12° N., 37° E.), Bangenze Mt, ? v.,
Hokwa (5°09' N., 28 "41' E.), 14., 2 vials, Ibba, 23^.1937 (/. G. Myers), A.M.N.H. ;
Geneina, 26.111.1927 (Evans), Li Yubu (Zande Distr.), vi.i948 (G. M. Culwick), L.
Keilak, 30^1.1952 (C. Siveeney}.

CHAD : Ennedi Mts, x.ig57 (Brit. Ennedi Expedit.}.

ETHIOPIA : Boutta (? Butta Mts, 7° N., 37° E.), no date (Rothschild], Stockh. ;
Gula R. (i5°35' N., 38°25' E.), no date, Moyale, and N. of Mega, xi.i943 (Buxton).

SOMALI REPUBLIC : Abruzzi, no date (G. Paoli), 1930 (G. Russo), A.M.N.H.,
N.C.I. ; Halaya (9°34' N., 440o8' E.), 2O.iii., Lanmulaho (8°36' N., 45°i3' E.),
I3.vii.i946 (P. E. Glover).

WESTERN ADEN PROTECTORATE : Dhala, i4.ix., Jebel Jihaf, 2O.ix., Jebel Harir,
3O.X.I937 (H. Scott, E. B. Britton), Habban-Yeshbum Rd Junction, i.xii.igsi
(W. V. Harris}.

UGANDA : Bugala Isl., no date (Bayon), Stockh. ; Bugiri, 5.-8.viii.i957 (P.
Basilewsky and N. Leleup), Terv. ; Masaka, 1925 (L. R. Hancock), 20 m. fr. Kampala
tow. Entebbe, n.xii.i934 (H. Kirby), Bunyoro (Butiaba), Gulu, Bweramule (Toro
tribe), West Nile Region, no dates, 4 vials (D. R. Buxton), A.M.N.H. ; Katwe, no
date recorded, Kikundwa (i°25' N., 3i°22' E.), 1946, Kawanda (o°2o' N., 32°3o'
E.), 21^.1949, Serere, x.1949, Mbarara, 14. xi. 1949 (W. V. Harris}.

KENYA : Kalodeka (2°2o' N., 35°5o' E.), 20.1.1933 (R. Jeannel), type-colony of
B. jeanneli Grasse, (own collection), syntypes in A.M.N.H. Other Material :
Lumbwa, no date (Sandberg), Guaso Nyiro (? Waso), 19.11.1911 (E. Lonnberg),
Stockh. ; Turkana Distr., no date (D. R. Buxton), Ruiru, 20. xi., 26 mi. W. of Nakuru,
5.xii.i934, Namanga R., 4., Taveta Forest, 25., Mombasa, 30.1.1935 (H. Kirby),
A.M.N.H. ; Rumuruti, 8.x. 1950 (W. V. Harris}, Kalodeka (as above), iv.igsi
(B. F. F. Harrison), Mem, x.1952 (W. A. Sands), Kitui District, 1954 (Heisch).

RWANDA : Kigali, 1933 (Van Assche), Terv. ; Bugesera (2 "05' S., 3O°2o' E.),
1 3. -i 7. xi. 1 963, 8 vials (A. Bouillon), U. Lov.

BURUNDI : Kitega, x.1935 (P. Lefevre), Muhinga, 1938 (Smets), Terv.

TERMITES OF THE GENUS MACROTERMES 427

TANZANIA : Kokotoni (Zanzibar), I2.viii.i889 (4. Voeltzkow}, type collection of
T. bellicosus subsp. sansibarita Wasmann, Stockh., syntypes in A.M.N.H. ; Zanzibar,
no dates (Stuhlmann, Aders, collector unknown), Stockh., I.R.S.N., B.M.(N.H.) ;
Tanga, no date (v. Rederi), 8^.1935 (W. Dethleffsen}, Stockh., Hamb. ; Himo,
i., 5.xi.i934 (H. Kirby), i8.vii.i957 (P. Basilewsky and N. Leleup), A.M.N.H.,
Terv. ; Amani, 1908 (Oberleutnant von Puttkamer), i.x.1934 (H. Kirby), Hamb.,
A.M.N.H. ; Shinyanga, io.vi.ig32 (collector unknown), N.C.I, and A.M.N.H. ;
Funzi Isl., 20.iii.i9O3 (^4. Voeltzkow}, Moshi, 23.x., i.-4.xi.ig34, 6 vials, Dodoma,
22., Babati, 23.1.1935 (H. Kirby}, A.M.N.H. ; Kisarawe, 1903 (F. Eichelbaum),
Hamb. ; Karema, no date (Storms), I.R.S.N. ; Kibongoto, no date (Sjostedt),
Stockh. ; Longido, 17., 20.^.1957 (P. Basilewsky and N. Leleup), Terv. ; Dar-es-
Salaam, 15. vi. 1916 (A. Loveridge), Rift Valley, 1934 (coll. unknown), Same, 24.11.1951
(H. M.}, Tabora, no date (A. Lindeman), 23.viii.i936 (W. V. Harris}, vi.i952 (P. R.
Hesse), Bukoba-L. Ikimba, no date recorded, Kalambo R., 23.x. 1933, Tengeru,
3i.viii.i95o (W. V. Harris}.

ANGOLA : Cazombo, 10.11.1955 (A de Barros Machado), (Dundo) and Hamb. ;
Dundo, no date recorded (A. de B. M.}, (Dundo) and U. Lov. ; Luisavo Falls (n°5o'
S., 23°35' E.), 23.11.1955 (Luna de Carvalho}, Dundo; Perdiva, 30 mi. E. of Pt
Alexandre, no data, B.M.(N.H.).

ZAMBIA : N'Changa, no date (C. T. Macnamara), Lusaka, no data, and I7.viii.ig49
(Hope-Simpson), B.M.(N.H.) ; Choma, 10., 12., 14.1.1957 (W. G. H. Coaton), N.C.I. ;
Lusaka, g.viii., I2.xii.i966 (M. G. Bingham), U. Lov.

MALAWI : Zomba, no date (H. H. Johnston), 24 m. fr. Ngabu on Chiromo Rd,
15., 9 m. fr. Ft Johnston on Farringdon Rd, 25.viii., Bua R., I7.ix.ig53 (W. A.
Sands and W. Wilkinson).

MOZAMBIQUE : Guengire (ig°oi' S., 34°ig' E.), no date (Vasse), Stockh.

SOUTH WEST AFRICA : 40 mi. N. W. of Okahandja, 24.!!., Swartbooisdrift (i7°2i'
S., I3°5i' E.), i8.x.ig63 (F. Gaerdes), N.C.I.

In addition to those, exactly go nest series collected by W. G. H. Coaton, G. F.
Pretorius and /. L. Sheasby between September ig65 and October ig66 have been
identified as M. subhyalinus. They are kept in the N.C.I., Pretoria. A detailed
list of the 24 localities involved is available at the B.M.(N.H.) ; the following,
however, should be cited here : Epupa Falls (17° S., I3°2o' E.), Ohopoho, Ombombo,
Fransfontein, Omaruru, Andara.

RHODESIA : Rekomitjie (i6°o7' S., 2g°23' E.), I4.x.ig64 (M. G. Bingham),
B.M.(N.H.) and U. Lov. ; Mkwichi (i6°i7' S., 30° E.), Chenanga (i6°2g' S., 30°i4'
E.), x.ig65 (M. G. B.), U. Lov.

A total of 412 nest series were examined and, when no other indication is given,
the material is in the B.M.(N.H.).

Under the name of ' bellicosus ', the nest structure of this species has been studied
and its mounds figured from a variety of places : West and Central Africa (Grasse",
ig37«, ig44 ; Grasse" & Noirot, ig6i), East Africa (Harris, igs6, ig6i), Angola
(Weidner, ig6i), to quote only recent literature and reliable determinations.

From the collecting data accompanying the samples, some dates of swarming
may be mentioned : Adiopodoume (Ivory Coast), 8.vi.ig52 ; Luluabourg (Congo)

428

J. E. RUELLE

22.xi.ig2i, two days after rain and 20 minutes after moonrise ; Kawanda (Uganda),
21^.1949, night ; Lusaka (Zambia), I2.xii.i966, at midnight.

Macrotermes ukuzii Fuller
(Text-figs 123-132 ; Map n)

Termes (Termes) parvus Holmgren, 1913 : 325. Type-locality : REPUBLIC OF SOUTH AFRICA,
Zulnland [junior primary homonym of T. parvus Haviland, 1898].

MAP 10. Collecting localities of Macrotermes subhyalinus.

TERMITES OF THE GENUS MACROTERMES

429

Macrotermes ukuzii Fuller, 1922 : 80. Type-locality : REPUBLIC OF SOUTH AFRICA, Ukuzi

[= Mkuzi] R.
Macrotermes bellicosus var. ukuzii Fuller, 1927 : 139 [erroneous reduction to subspecies].

Imago (previously undescribed) . Head capsule light chestnut-brown, with four light dots
between fontanelle, ocelli and postclypeus ; postclypeus lighter than head, with conspicuous
dark median line. Margins of pronotum darker than head, central area lighter without sharply
denned pattern of marks or dots. Abdominal tergites same colour as thorax, sternites lighter.
Tibiae darker than femora. Wings smoky yellow.

Head capsule in plan view slightly elongate ; fontanelle protruding from a depression,
followed anteriorly by a median ridge which is lower than the fontanellar hump. Eyes broadly
oval and proportionately large ; ocelli wide, distant from the eye by J of their major diameter.
Postclypeus narrow, moderately inflated. Antennae ig-jointed, 3rd segment barely longer
than 2nd. Quite a few bristles on head capsule, shorter setae around fontanelle.

Pronotum long, numerous setae on margins and disc, front margin almost straight, hind
margin more emarginate, sides convex.

Measurements in millimetres : morphotype— range and mean of 26 specimens from 7
localities :

Head width across eyes
Eye major diameter ..
Ocellus major diameter
Ocellus minor diameter
Ocellus ex eye ...
Median length of pronotum
Maximum width of pronotum
Length of hind tibia ..
Length of fore wing ..
Maximum width of fore wing

Morphotype Range

3-08

2-93-3-32

1-09

0-93-1-15

0-46

o-37-o-53

o-35

0-28-0-40

0-14

O-I2-O-2O

i-59

i -53-i '74

2-91

273-3-48

3-79

3-48-4-58

—

32-00-37-50

—

8-00-9-50

Mean

3-14
1-05

o-45
o-35
0-15
1-66
3-09
3-98
33-50
8-90

Major soldier. In addition to the original description (Fuller) : head capsule sometimes
very dark ; rather flattened in side view ; frontal depression shallow. Mandibles usually
stout, only tip hooked. Pilosity of head capsule conspicuous.

Pronotum not much narrower than head capsule, its front and hind margins distinctly
emarginate, sides of the anterior lobes also emarginate. Shape of meso- and metanotum
variable, metanotum wider than mesonotum.

Hind tibia short.

Measurements in millimetres : holotype — range and mean of 26 specimens from 13 localities :

Head length .....

Head width .....

Head depth inc. gula

Length of gula .....

Maximum width of gula .

Minimum width of gula

Length of left mandible

Length of hind tibia.

Maximum length of pronotum .

Maximum width of pronotum

Minor soldier. Additional notes to Fuller's description : Gula same colour as head, sides
darker. Antennae sepia brown. Thoracic nota yellowish brown, lighter than head, outline
darker. Abdominal sternites lighter than tergites, the latter slightly lighter than thoracic
nota.

Holotype

Range

Mean

4-19

4-19-5-48

479

3-10

2 -96-3 -5°

3-28

2-06

1-92-2-36

2-20

2-96

2-96-3-74

3-34

I-OO

1-00-1-33

i-i3

0-68

0-62-0-85

0-76

2-31

2 -O8-2 -57

2-37

2-52

2-23-2-86

2-59

i-33

I-33-I-63

1-49

2-44

2 -44-3 -O6

279

430

J. E. RUELLE

Head capsule elongate, barely narrowed in front, sides straight.

Measurements (27 specimens from 13 localities) in millimetres :

Range Mean

Head length ...... 2-21-3-12 2-48

Head width ...... 1-79-2-34 1-99

Head depth inc. gula .... 1-17-1-66 1-36

Length of gula 1-38-2-15 1-63

Maximum width of gula .... 0-62-0-78 0-68

Length of left mandible .... 1-59-2-08 1-76

Length of hind tibia. .... 1-62-2-31 1-87

Maximum length of pronotum . . . o-88-i-oi 0-94

Maximum width of pronotum . . . i -43-1 -66 i -56

Major worker. Head width (7 specimens from 4 localities) : range 2-09-2-51, mean 2-36 mm.

129

13O

FIGS 123-132. Macrotermes ukuzii, imago, soldiers. 123-124. Imago : 123, head
capsule and pronotum, plan view (with details of light marks) ; 124, head capsule, side
view. 125-130. Major soldier : 125-127, head capsule and thoracic nota, plan view ;
128-130, head capsule, side view. 131-132. Minor soldier, head capsule and thoracic
nota, plan view.

TERMITES OF THE GENUS MACROTERMES 431

Variation. The major soldiers have a strikingly pilose head capsule ; specimens from
southern Tanzania, however, do not (see discussion below).

This species is closely related to M. natalensis. In the images, the postclypeus
is more inflated and definitely longer in proportion to its width ; the ocelli are closer
to the eyes and these are proportionately bigger ; the median ridge in front of the
fontanelle is lower, not higher, than the fontanellar hump. The head capsule of

MAP ii. Collecting localities of Macrotermes ukuzii.

432 J. E. RUELLE

the major soldier, apart from its small size, is narrow (index head length/head width
close to 1-5, the highest value in the genus) and pilose.

The holotype major soldier of T. parvus Holmgren is unusually small, even for
M. ukuzii ; it agrees, however, with Fuller's specimens and the additional material
examined. The probability that all of those nest series would be aberrant samples
of a widely distributed species like M. mossambiciis or M. natalensis is not excluded,
but is considered too low to prevent recognizing M. ukuzii as a distinct species.
Hybrids, as Fuller (1927) seems to imply, are also possible, but this is a matter for
field study.

Type Material. REPUBLIC OF SOUTH AFRICA : Zululand, no date (Trdgdrdh),
holotype major soldier of T. parvus Holmgren, A.M.N.H. Morphotype : Natal :
Ndumo, 2o.ix.ig22 (Cl. Fuller), $ imago, N.C.I.

Other Material from Natal : Mkuzi R. near Mtanto Drift, 26.vi.i92O (E. Collins),
type collection of M. ukuzii Fuller, N.C.I, and B.M.(N.H.) ; Ndumo, 20. ix. 1922,
Lower Mkuzi Drift, u.ix.i923, Somkele (28°2o' S., 32°O3' E.), 4.xi.ig24 (Cl. Fuller),
Hluhluwe, I5.ii.i937 (C. J acot-Guillarmod) , Ingwavuma (27°io' S., 32° E.), i8.xi.i939
(W. G. H. Coaton), 28.1.1959 (/. H. Grobler), N.C.I. Transvaal : Komatipoort,
io.xi.i958 and i6.xi.i959 (P. C. Joubert), Lower Sabie (25°io' S., 3i°54' E.), 9.,
Skukuza, io.xi.i949, Evangelina-Alldays (22°4o' S., 28°55' E.), 10., Waterpoort-
Louis Trichardt, 16., Stolzenfels-Tonash (22° 43' S., 28°35' E.), 19.^.1964 (W. G. H.
Coaton), N.C.I. ; Eastern Lowveld, no locality, iii.1966 (D. H. Kistner), own collec-
tion.

SWAZILAND : Mankaiana, 26.x. 1960, 3 vials (W. G. H. C.), N.C.I.

TANZANIA : Shinyanga, 1931 (S. N. Bax), A.M.N.H. ; Kakoma, no date, Itigi,
io.ix.iQ49 (-P- B. Kemp), B.M.(N.H.).

A total of 27 nest series were examined.

The identity of the 3 samples from Tanzania is less certain, for reasons of geo-
graphical gap, scarce pilosity of head capsule in major soldiers and large size of
minor soldiers. Pending further evidence, however, they are assigned to M.
ukuzii.

From the collecting dates, this species swarms in Transvaal at night, around
the middle of November. (M. natalensis swarms at the same time, at least in
Komatipoort.)

Macrotermes vitrialatus (Sjostedt)
(Text-figs 133-154 ; Map 12)

Termes vitrialatus Sjostedt, 1899 : 34. Type-locality : CONGO.

Termes (Macrotermes) vitrialatus Sjostedt ; Holmgren, 1912 : 29.

Termes imperator Sjostedt, 1913 : 359. Type-locality : CONGO, Sankisia, syn. n.

Termes waterbergi Fuller, 1915 : 466. Type-locality : REPUBLIC OF SOUTH AFRICA, Warm bad,

syn. n.
Macrotermes Schoutedeni Sjostedt, 1924 : 39. Type-locality : CONGO, Banana [not Boma],

syn. n.
Amplitermes mozambicanus Sjostedt, 1926 : 89. Type-locality : MOZAMBIQUE, Andrada

[neither ' Anodrada ' — Sjostedt — nor ' Anodraba ' — Snyder], syn. n.

TERMITES OF THE GENUS MACROTERMES

433

135

138

FIGS 133-138. Macrotermes vitrialatus, imago : 133-135. head capsule and pronotum,
plan view (with details of light marks) ; 136-138, head capsule, side view.

434

J. E. RUELLE

Holotype
3-92

Range
3-61-4-35

Mean
4-05

1-07

0-90-1 -17

1-03

o-43
o-37
0-40
1-90

o-33-o-49
0-22-0-42
0-34-0-60
1-61-2-02

0-41
0-36
0-50
1-89

3-90
4-84

3-25-4-45
4 -00-5 -oo

4-04
4-57

33-50

31-50-40-0

35-50

9-00

7-00-9-50

8-40

Macrotermes angolensis Noirot, 1955 : 142. Type-locality : ANGOLA, Coemba, syn. n.

Imago. Head brown to dark chestnut-brown ; postclypeus lighter than head, with dark
median line ; base of mandibles as postclypeus, apical third black. Pronotum same colour
as head, with two large yellow spots towards the sides in concave depression near front margin,
two light dots in the middle on small humps near hind margin. Abdominal tergites brown,
sternites more reddish than tergites. Legs yellowish brown, tibiae slightly darker than femora.
Wings transparent, veins yellowish to brownish yellow.

Fontanelle an inconspicuous light dot in a very shallow depression, followed anteriorly by a
faint median carina. A few scattered bristles on head capsule. Eyes sub-circular, prominent,
but small ; ocelli rounded, separated from the eyes by more than their major diameter. Post-
clypeus in side view moderately prominent. Antennae ig-jointed, 2nd segment equal to 4th,
3rd longer than both.

Width of pronotum twice or more its median length and often superior to head width ;
antero-lateral corners broadly rounded, short setae conspicuous on front edge only.

Measurements in millimetres : holotype — range and mean of 39 specimens from 15 localities :

Head width across eyes

Eye major diameter ....

Ocellus major diameter

Ocellus minor diameter

Ocellus ex eye .....

Median length of pronotum

Maximum width of pronotum

Length of hind tibia ....

Length of fore wing ....

Maximum width of fore wing

Major soldier. Head reddish yellow to reddish brown, ventral side lighter, gula darker than
genae. Mandibles usually black throughout, base sometimes dark reddish brown. Thorax
darker than head, especially the outline ; abdominal tergites more often than not lighter than
thorax, sternites lighter than tergites ; tibiae usually not darker than femora ; setae darker
than cuticle, with light area around base.

Head capsule in plan view trapezoidal to pear-shaped, more or less constricted in front ;
frontal area not deeply depressed, but low median ridge following anteriorly the fontanelle,
the latter more or less indistinct. Mandibles long, rather evenly incurved, left one a little
more than right. Hyaline tip of labrum acute, sides straight to concave. Antennae 17- to
i8-jointed ; when 17-, 3rd segment constricted across middle, and longer than twice the 2nd.
Pilosity variable, usually sparse.

Pronotum relatively wide, lateral corners rounded, front and hind margin about equally
incurved ; mesonotum at times (i in 3) wider than metanotum, the latter with a typical outline
of parallel sides and straight hind margin.

Measurements (138 specimens from 37 localities) in millimetres :

Head length
Head width
Head depth inc. gula
Length of gula .
Maximum width of gula
Minimum width of gula
Length of left mandible
Length of hind tibia .
Maximum length of pronotum
Maximum width of pronotum

Range

Mean

4-00-6-86

5-73

3-26-5-58

4-73

2-24-3-68

3-00

2-9I-4-39

3-58

1-10-1-52

1-27

0-76-1-07

0-94

2-58-4-14

3-37

3-27-5-26

4-12

1-46-2-21

1-82

2-70-4-14

3-37

TERMITES OF THE GENUS MACROTERMES

435

143

144

145

146

FIGS 139-146. Macrotermes vitrialatus, major soldier, head capsule and thoracic nota,

plan view.

436

J. E. RUELLE

Minor soldier. Head capsule yellowish brown to reddish brown ; gula, at least along the
sides, darker than ventral genae ; mandibles dark red ; antennae lighter than head. Thorax
as often as not lighter than head, front and hind margins darker. Abdominal tergites same
colour as head capsule but without reddish tinge, sternites lighter ; legs same colour as sternites.

Head capsule in plan view ovate-quadrate, fontanelle inconspicuous. Labrum large. Man-
dibles long and slender, tip incurved. Antennae ij-jointed, 4th segment longer than 3rd.
Pilosity sparse, even on abdominal nota.

Pronotum with sinuate anterior margin, posterior emarginate ; mesonotum distinctly
narrower than both pro- and metanotum.

Measurements (106 specimens from 33 localities) in millimetres :

Head length ......

Head width ......

Head depth inc. gula ....

Length of gula. .....

Maximum width of gula ....

Length of left mandible .

Length of hind tibia .

Maximum length of pronotum .

Maximum width of pronotum
Major worker. Head capsule drab reddish brown, paler between fontanelle and hind margin
of postclypeus ; in plan view evenly rounded, widest behind articulation of mandibles. Head
width (207 specimens from 24 localities) : range 2-23-3-38, mean 2-69 mm.

Variation. The outline of the head capsule in the major soldier is variable (Text-figs 139-
146). Besides a denser pilosity found in some South African specimens, other samples from
Western Congo are smaller and more rounded. Still others, from Kwango and adjacent Angola,
have a trapezoidal outline with sides nearly straight ; the lateral corners of the pronotum are
also more acute and the mandibles less incurved than usual. Imagos are not distinctive to the
same extent. However, it is worth noting that native people from northern Angola use different
vernacular names for the specimens from Dundo (the usual form, called ' lungo ') and for those
from Camissombo-Capaia and Caluango (this variant, called ' Ikhala '). These observations
have been communicated, with some doubts, by a well-known field naturalist (A. de Barros
Machado, in lift.). It is thus anticipated that a new subspecies may be created when more,

Range

Mean

2-20-3-25

2-62

1-73-2-73

2-09

i -08-1 -85

1-47

1-39-2-07

i-59

0-63-0-77

0-69

1-74-2-36

2-OO

2-52-3-42

3-00

0-92-1-35

I -06

1-51-2-28

1-74

148

15O

151

152

FIGS 147-152. Macrotermes vitrialatus, minor soldier, head capsule and thoracic nota,

plan view.

TERMITES OF THE GENUS MACROTERMES 437

and more complete, samples, including all castes and collected in the same locality, will be
available. For the time being, the evidence is still considered insufficient.

The following notes are added here to the comparison already made between this
species and M. muelleri (above) : The imago of M. vitrialatus has the smallest eyes
in the whole genus : index H/E 3-93 (in M. muelleri 3*38, in M. bellicosus 2-87) ;
the postclypeus also looks relatively longer and slightly more prominent than in
M. muelleri. The major soldier's head capsule in M. vitrialatus is less narrowed in
front, the pronotum is proportionately wider and its sides are more rounded than is
the case with M. muelleri. The gula in the minor soldier is shorter and wider than
in M. nobilis and M. muelleri ; the 3rd antennal joint is longer than the 4th in
muelleri. (Incidentally, it may be noted that seven of the species redescribed in
this work have in the major soldiers a left mandible that averages more than half
of the head length : falciger, herus, ivorensis, lilljeborgi, muelleri, natalensis, vitria-
latus.}

Type Material. CONGO : no locality, ? 1893 (H. Freyschmidt} , holotype <$ imago
of T. vitrialatus Sjostedt, Hamb. ; Sankisia (9*22' S., 2^46' E.), ig.xi.ign
(Bequaert), type collection of T. imperator Sj., Terv. ; other syntypes in Stockh.,
A.M.N.H., U. Lov. ; Banana (the label ' Boma ' is a mistake : H. Schouteden,
priv. comm.), vii.i92O (H. Schouteden}, type collection of M. schoutedeni Sj., Terv. ;
other syntypes in Stockh. and A.M.N.H.

Other Material. CONGO (Brazzaville) : 3 m. fr. Brazzaville on N'Gabe Rd,
2.vii.i959 (/. E. Ruelle}, U. Lov. and A.M.N.H.

CONGO : Kimwenza (4°3o' S., I5°i3' E.), G.xi.igsg, Mayidi (5°n' S., i5°o9' E.),
3.vii.i963, 22.1.1964, 4^.1968, 14 m. fr. Thysville on Matadi Rd, 9.^.1964 (/. E.
Ruelle}, U. Lov. and B.M.(N.H.) ; Boma, 3.iv., 20^1.1913 (Styczynski], Terv. and
Stockh. ; Mukimbungu, i4.xi.i9O4 (K. E. Laman}, Stockh. and A.M.N.H. ; Dilolo,
no data, Kapanga, ix.i932 (Overlaet}, Tumbwe, xi.i92i (M. Bequaert}, Luashi,
iii.i936 (Freyne}, Terv. ; Kinda, no date (Muller}, Mpese, no date (Cooreman),
Vista (5°5i' S., I2°i7' E.), no date (V. Moerenhout}, I.R.S.N. ; Kimbao (5°3o' S.,
I7°28' E.), 20.viii.i959 (/. E. Ruelle}, U. Lov.

ANGOLA : Coemba (i2°io' S., i8°i5' E.), Dundo Ace. No. Ang. 1606, type
collection of M. angolensis Noirot, (Dundo) ; other syntypes in A.M.N.H., Terv.,
U. Lov. Other Material : 5 and 9 mi. E. of Dundo, 1961, 2 vials (Ed. Luna de
Carvalho and A. de Barros Machado}, Camissombo-Capaia (8°io' S., 2O°4o' E.),
15.^.1961 (L. de C.}, Caluango (8°i9' S., I9°52' E.), 3 and s.ix.igGi, 4 vials (L. de C.
and de B. M.), Dundo.

ZAMBIA : Lusaka, no date (W. V. Harris}, 2.vii. and 2i.xi.i966 (M. G. Bingham},
B.M.(N.H.), U. Lov., A.M.N.H. ; Chinsi Hills, i6.viii.io,49 (Hope-Simpson),
B.M.(N.H.) ; Choma, 15.1.1957 (W. G. H. Coaton}, Magoye, 17.1.1957 (E- N- Cooling),
N.C.I.

MALAWI : 25 m. fr. Kota-Kota on Kasungu Rd, 17., 7 m. fr. Loweya R. on
Nkata Bay Rd, 2i.ix.i953 (W. A. Sands and W. Wilkinson), B.M.(N.H.).

SOUTH WEST AFRICA : Otjitambe (Outjo), 14.1.1951 (F. Gaerdes}, 20 mi. ex
Gobabis-Windhoek, 12.^.1965 (W. G. H. Coaton}, Katima Mulilo, 30^.1965

438 J. E. RUELLE

(A. Barnard), Ohopoho-Rua Cana Falls, i2.-i4.iv., 3 vials, Ohopoho-Ombombo,
i.v., 2 vials, Ombombo-Gauko Otavi, 2.v.ig66 (W. G. H. Coaton, J. L. Sheasby).
Further material from same territory includes 7 nest series from 4 places between
i8°-20°i5' S. and i9°45'-20°45' E., I2.iv. to 1^.1967 (W. G. H. C., J. L. S. and
G. F. Pretorius), N.C.I.

RHODESIA : Selinde Mt, ig.xii.i928 (R. Boulton), B.M.(N.H.) and A.M.N.H. ;
Gokwe District, i8.xii.i962 (M. G. Bingham), B.M.(N.H.) ; Salisbury, no dates,
2 vials (Marshall), Stockh. ; Matsikite (i6°2o' S., 29°47' E.), Chenanga (i6°29' S.,
3O°i4' E.), x.igGs, Matopos, 23. xi. and 5.xii.i965 (M. G. Bingham), U. Lov.

MOZAMBIQUE : Andrada, 1905 (G. Vasse), type collection of Amplitermes mozam-
bicanus Sj., (Museum Paris), syntypes in Stockh. and A.M.N.H.

REPUBLIC OF SOUTH AFRICA : Transvaal : Warmbad, I4.x.i9i4 (Simmons),
type collection of M. waterbergi Fuller, N.C.I. ; paratypes from De Wildt (25°39'
S., 27°57' E.), I4.X.IQI4 to iv.igig, 4 vials (Cl. Fuller), N.C.I, and B.M.(N.H.).
Other Material : Rus De Winter (25°i2' S., 28°28' E.), 12. and 24.ix.i957 (P. C.
Joubert), 13.^.1963 (/. L. Sheasby), N.C.I., U. Lov., A.M.N.H. ; Rustenburg-
Boshoek, i., Vaalwater, 27.ix.i96i (W. G. H. Coaton), Northam-Middelwit,
4.x.i96i, Maraheki-Warmbad, 26.^.1964 (/. L. Sheasby), N.C.I, and U. Lov. ;
Nylstroom, i6.ix.i9i8 (Cl. Fuller), Hammanskraal, I2.viii., 20.ix.i963, 6., 28.,
30.1.1964 (/. L. Sheasby), N.C.I.

A total of 89 nest series were examined.

This species was described under six different names from 1899 to 1955 ; in the
collections, some samples had been identified as M. natalensis and one other mis-
identification should be quoted here (Sjostedt, 1911 : 143 ; ' Termes bellicosus ').

The case of T. imperator parallels that of T. carboniceps with samples of unusually
large soldiers. (At the lower range we have here M. angolensis.) T. vitrialatus and
A. mozambicanus were first represented by alates only and collected far apart.
It may be noted that M. schoutedeni had already been considered as a synonym of
M. vitrialatus before this revision was undertaken (A. E. Emerson, private com-
munication). As regards M. waterbergi, sufficient evidence has been gathered
from Transvaal and Rhodesia since Fuller's time to leave little doubt about its
status, in spite of its somewhat more conspicuous pilosity. The specimens from
some parts of Angola have been discussed above.

M. vitrialatus may be more abundant than it looks from the distribution map ;
its nests indeed are often inconspicuous, even completely subterranean. They are
aptly described by Fuller (1921, 26) as mounds ' of loose soil, not unlike that of
new grave ' : this has been confirmed since in South Africa and Angola (Coaton,
de Barros Machado, in litt.) as well as in the Congo, near Kinshasa. Harder
hummocks do occur, however, on more clayey soils, e.g. the region around Thysville.
(The fig. 3 in Noirot, 1955 : 144, which looks like a nest of M. bellicosus, could
unfortunately not be checked by redetermination of the sample.)

Although this is not a forest species, its major workers cut leaf fragments in a
fashion similar to that observed with M. muelleri, as has already been noted.
Reports from South Africa and Rhodesia (Coaton, Bingham, in litt.} agree with the
observations made in the Congo.

TERMITES OF THE GENUS MACROTERMES

439

The foraging behaviour in broad daylight, as observed with the type collection of
T. imperator, has also been observed near Kinshasa ; according to Bingham and
Coaton, it further occurs with M. falciger in Rhodesia and South Africa. This
ethological trait is not, therefore, a characteristic of the forest Macrotermes.

The evidence available suggests that M. vitrialatus, unlike other Macrotermes,
does not swarm at the onset of the rainy season (alates have been collected towards
the end of it, in the Congo) ; as far as is known, the images would take off immedi-
ately after dark rather than in the morning.

MAP 12. Collecting localities of Macrotermes vitrialatus.

440 J. E. RUELLE

ADDENDUM : DOUBTFUL PLACES

It has proved impossible to trace some localities on the maps available. Those
which have not been cited in this work as : X. (co-ordinates unknown) are listed
below with the species concerned and the place of deposit. Vague indications such
as ' Kafferlandet ', ' Deutsch Ost-Afrika ' and the like — including 33 nest series-
have been omitted altogether (save for the type specimens).

ETHIOPIA. M. subhyalinus : Lake Hora Harsadi, ? 1927 (collector unknown),
B.M.(N.H.) ; Katchinoa, Yaba, no dates (Rothschild), Stockh.

CONGO. M. subhyalinus : Tarambo (Lang-Chapin coll. No. 1528), A.M.N.H.

MALAWI. M. vitrialatus : Kasu, iii.i9i6 (/. B. Davey), B.M.(N.H.).

ACKNOWLEDGEMENTS

First of all I thank Prof. Albert Bouillon, of the Lovanium University, Kinshasa,
for starting my interest in Macrotermes early in 1959 and for providing all kinds of
facilities — including financial support — throughout this study.

Next I wish to express my gratitude to Dr W. V. Harris, Officer-in-Charge,
Termite Research Unit, British Museum (Natural History), for access to the London
and Cambridge collections since 1962, not less than for helpful advice and encourage-
ment ; to Mr W. A. Sands, of the same unit, I owe much of the enlightening remarks
and constructive criticism in the preparation of this revision.

My special thanks go to Dr A. E. Emerson for liberal communication of his long
experience and unique collection of data on termites as well as for his hospitality
at home ; in the same year (1967) I enjoyed staying at the American Museum of
Natural History, New York, with all the facilities generously provided by Jerome G.
Rozen Jr., Curator of the Department of Entomology, and especially with the warm
helpfulness of Dr Kumar Krishna.

Among other curators, Dr P. Basilewsky, Department of Entomology, Musee
Royal de 1'Afrique Centrale, Tervuren, should have been mentioned first ; for he
not only welcomed me to work on his rich termite collections in 1965, but at the
same time enabled me to borrow, thanks to the kindness of Dr E. Kjellander,
Stockholm and of Dr H. Weidner, Hamburg, the all-important collections kept in
the latter institutions. Special circumstances (my usual residence in the Congo)
made the sojourn at Tervuren the more valuable.

In spite of the insecurity attributed to the same Congo, Dr W. G. H. Coaton,
of the Plant Protection Research Institute, Pretoria, did not hesitate to send to
Kinshasa the bulk of the N.C.I, collections, of which many specimens have been
donated by him to Lovanium University ; this, and the useful collecting notes he
added, well deserve a detailed acknowledgement.

Dr Cooreman, Institut Royal des Sciences Naturelles, Bruxelles, and Prof. C.
Noirot, Faculte des Sciences, Universite de Dijon, have helped me in providing
access to, and in sending samples from, their invaluable accessions. To Prof.
Noirot I will add Messrs A. de Barros Machado and M. G. Bingham, when it comes to
acknowledging not only the liberal communication of samples, but also the very
interesting field notes gathered by experienced observers in parts of Africa that I was

TERMITES OF THE GENUS MACROTERMES 441

unable to visit myself. I am also grateful to Drs J. Deligne, D. H. Kistner, and
J. M. Pasteels for their collections ; not to forget the help received at Basel from
Dr Eb. Ernst.

Finally, my thanks are due to Miss M. Van Hove for help with the drawings and
the distribution maps, and to other people working in the laboratory of Prof. A.
Bouillon.

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444 J- E. RUELLE

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Professor Dr JEAN-EMILE RUELLE

FACULTE DES SCIENCES, B. P. 220

UNIVERSITE LOVANIUM

KINSHASA XI

DEMOCRATIC REPUBLIC OF THE CONGO

INDEX TO VOLUME XXIV

New taxonomic names, and names involved in nomenclatural changes, are in bold type

abdominalis, Laxenecera . . . 238

abdominalis, Leuctra .... 338
abdominalis, Microcephalothrips . . 89
abercornenis, Aciagrion steelae . . 181
aberrans, Carcelia ..... 36
abnorme, Anomisma . . . .193
abnormis, Perla ..... 338
acicularis, Chloroperla .... 338
actaeon, Drepanosticta . . . 174

acuta, Elattoneura . . . 175

adamsi, Argia . . . . .182
adjuncta, Polythore derivata . . .199
adytum, Agrion . . . . .182
aequatoriale, Heteragrion . . .196
aequatorialis, Thore picta . . .199
aestivalis, Echinomyia praeceps . . 40
aethiops, Perla . . . . -338
affinis, Nemoura ..... 338
affinis, Raphidia ..... 352
afra, Catoptropteryx 141 (fig.), 142 (fig.), 156-157
africanus, Neoitamus . . . -313
alatipes, Psilocnemis . . . .179
alatus, Thyridanthrax .... 76
albescens, Argiolestes griseus . . .196
albicauda, Lestes . . . . .194
albicauda, Synlestes . . . 193

albicincta, Laxenecera . 237 (figs), 240-241
albidipennis, Hermes .... 352
albidipennis, Nemoura .... 338
albistigma, Ischnura . . . .182
albomacula, Udamocercia . . . 338
albopunctata, Alophora .... 35
alcimoides, Philodicus .... 298
alcyone, Agriocnemis . . . .182
alcyone, Cora ..... 199

alleni, Austrolestes .... 194

alpinus, Argiolestes . . . .196
aluensis, Teinobasis . . . .182
amaurodytum, Agrion .... 182
ambigua, Catoptroptreryx 132 (fig.), 143 (figs),
144 (fig.), 149 (fig.). 15° (fig-). 159-160
ambigua, Polythore derivata . . .199
amelia, Neoneura . . . . .176
amphora, Xenomyza . 290 (fig.), 292 (fig.)
anascephala, Drepanosticta . . .174
Ancylorrhynchus . . . . 271-274
Andrenosoma .... 255-257
anglica, Capnia vidua .... 338
anglica, Rhabdiopteryx .... 338

angolense, Agriocnemis . . . .182
angusticornis, Neolaparus . . . 259
angustipennis, Heteragrion aequatoriale . 196
annandalei, Caconeura . . . .176
anomala, Phorocerosoma ... 49

antelopoides, Protosticta . . .174

antennatus, Baphikothrips . 92 (figs), 93
anticus, Hermes ..... 352
apicale, Acanthagrion . . . .182
apicalis, Catoptropteryx 132 (fig.), 140 (figs), 141
(fig.), 142 (fig.), 144 (fig.), 149 (fig.), 150 (fig.),

I55-I56. PL i

apicalis, Chloroneura . . . .176
apicalis, Platysticta . . . .174
apicalis, Proagonistes .... 254
arachnoides, Pseudocopera . . .179
arcuata, Loewinella . . . -257
argentea, Agriocnemis . . . .182
argenteostriata, Eutachina ... 42
argentifrons, Thyridanthrax . . -75-76
aridus, Lestes ..... 194
armata, Astochia . . . . .312
armstrongi, Amorphostigma . . .182
arthuri, Mortonagrion . . . .182
artocarpi, Machatothrips . . .119

aruanus, Lestes . . . . .194
assamensis, Protohermes . . . 352

assimilis, Rhaphidia .... 352
asteliae, Agrion . . . . .182
Astochia ..... 311-313
ater, Dolicholon ..... 56
atkinsoni, Gymnodexia .... 45
atomaria, Psilocnemis . . . .179
atrocyana, Notoneura . . . .176
attenuatum, Aciagrion . . . .182
aureifrons, Eutachina aureifrons . . 42
aureisquamosa, Eutachina . . .42
aureofrons, Pseudagrion . . .182

aureus, Argiolestes .... 196

aurichalcea, Eutachina .... 42
auricolor, Amorphostigma . . .182

auricolor, Caconeura dorsalis . . .176

auricomata, Laxenecera . . 237 (fig.), 238
aurifer, Laphria . . . 228, 229 (fig.)
aurita, Catoptropteryx 132 (fig.), 139 (fig-).
141 (figs), 143 (figs), 145 (figs), 149 (figs), 167-

168, PI. i

auritus, Platyneuromus . . . -352
australe, Dolichocolon . . . .40

446

australica, Archichauliodes
australiensis, Stenosialis
australis, Agriocnemis
autumnalis, Caconeura .
aygulus, Anthrax .
azureum, Aciagrion

bagnalli, Ecanthothrips

balli, Elattoneura .
banksi, Isosticta
Baphikothrips
Baphothrips .
basalis, Eutachina
batesi, Corydalis .
beadlei, Pseudagrion
beata, Thore
bellicosus, Macrotermes .

374 (figs)

bellicosus -tonga, Macrotermes .

bellona, Ceriagrion

bellulus, Protohermes

bezzi, Erycia .

bezziana, Myiobia .

bezziana, Myiofijia

bezziana, Stomatomyia .

bicolor, Euscelidia

bicolor, Raphidia .

biconicus, Protochauliodes

bidens, Nesobasis .

bidentatum, Ceriagrion .

bifasciata, Hyperechia .

bifasciata, Udamocercia .

bilineata, Melanoneura .

bilineata, Trichocnemis .

biloba, Pteronarcys

bisetosa, Sturmia .

bispinus, Perilestes

bituberculata, Nemoura .

blackburni, Megalagrion

boliviana, Thore .

borneensis, Trichocnemis

botti, Caconeura . . . ,

bowringi, Chauliodes

bradleyi, Teinobasis

brevipennis, Dinotoperla

brisbanense, Agrion

buckleyi, Mecistogaster .

buitenzorgiensis, Carcelia

bullata, Spaniocerca

burmana, Inocellia

burmanensis, Caconeura verticalis ,

burmanica, Chaetoptiliopsis

buxtoni, Ischnura .

cacharensis, Disparoneura campioni
caflfer, Stichopogon
calcipennis, Microstigma
caldwelli, Monoleptophaga
caldwelli, Zenillia . . . ,
californica, Archilestes .

INDEX

353 californicus, Chauliodes .

353 californicus, Pteronarcys

183 caligula, Ommatius

176 calliphya, Agrion .

72 cambrica, Nemoura

183 campioni, Coeliccia

campioni, Disparoneura .

96 canescens, Neosticta

176 canicoxa, Ommatius

176 canningi, Caconeura

.90-91 capreola, Catoptropteryx

94 142 (figs), 144 (figs), 146

• 42

353 cara, Protoneura .

183 carbonaria, Laphria

199 carboniceps, Termes .

377 (figs), cardinalis, Ischnura

381 carpenteri, Dinotoperla .

. 398 carpenteri, Perasis

183 carrillica, Philogenia

353 caspica, Eucarcelia

41 castellani, Coenagrion

47 Catoptropteryx

47 caudata, Carcelia .

49 caudatella, Carcelia
220 cervula, Ischnura .
353 ceylanica, Archibasis
353 championi, Philogenia
183 chapini, Laxenecera
183 chaseni, Calicnemis

250 chatterjeeana, Sturmia .

338 cheesmanae, Austrolestes
176 cheesmanae, Ischnura
179 chelata, Argia

339 chilensis, Archichauliodes

50 chilensis, Sialis
193 chirripa, Cora

339 chrysoides, Argiolestes .

183 chrysonema, Laxenecera

199 chrysostoma, Brahmana

179 cinerascens, Philodicus .

176 cinnamonea, Atriadops .

• 353 circulatum, Enallagma .
183 citrinum, Ceriagrion

339 citronella, Perla

183 civiloides, Eutachina

. 193 claratibia, Phlaeothrips

37 clauseni, Pseudagrion

339 clio, Isogenus

353 clymene, Chloroperla

176 coartans, Argiolestes

39 coelestina, Alloneura

183 coelestis, Pseudagrion

coeruleiceps, Pseudagrion

176 coeruleum, Pseudagrion .

283 colensonis, Agrion .

193 collopistes, Telebasis

47 coloratus, Baphikothrips

54 compacta, Dictyopteryx

194 completa, Nemoura

• 353

• 339
324. 325 (ng.)

. 183

• 339

• i?9
. 176
. 176

• 326 (fig.)
. 176

132 (fig.), 141 (fig-),

(fig.), 147 (fig.), 150

(figs), 151

. 176

. 228

• 374
. 183

• 339

• 245
196

41

• 183
127-170

37
37

• 183
. 183

196

• 243
. 179

50

. 194
. 184
. 184

• 353

• 353

• 199
196
242

• 339
. 300

81

. 184
. 184

• 339

42

. . .98
. 184

• 339

• 339
196

. 176

i . 184

. . .184

• . . -, . 184

. 194

. • - . 184

91, 92 (fig.)

- » . -339

• 339

INDEX

complexa, Andrenosoma
compressa, Platerycia
concinna, Thore
concolor, Peltoperla
congoensis, Gonioscelis
congoicola, Thallosia
congoiensis, Rhipidocephala
Congomochtherus
conjuncta, Lestoidea
continentalis, Parachauliodes .
contristans, Laphria
corallinum, Ceriagrion
cordata, Capnia
corripiens, Hermes,
corsicana, Nemura (Protonemura)
costalis, Hermes
crassicornis, Corydalis
crassipes, Euoplothrips .
crassiseta, Bactromyia .
crocops, Pseudagrion
cruciata, Nemoura
crux, Ancylorrhynchus .
ctenoventris, Laphria .
cupraurea, Argia .
curtum, Megapodagrion .
cyane, Cora ....
cyaneothorax, Coeliccia .
cyaneovittata, Esme
cyanops, Trichocnemis .
cydippe, Chloroperla
cymodoce, Perla

dammermani, Eucarcelia
Dasyllina ....
davenporti, Ceylonolestes
davenporti, Protosticta .
deceptor, Agrion .
deceptor, Archichauliodes
decimmaculatus, Hermes
decipiens, Lestes .
decisa, Perla (Chloroperla)
decolorata, Perla (Chloroperla)
decoloratum, Libyagrion
decoratus, Neolaparus
dentifera, Drepanosticta
derivata, Thore

descendens, Acanthagrion apicale
despaxi, Leuctra .
dichroa, Dictyopteryx
digiticollis, Telebasis
digittatus, Ommatius
dilatus, Mystrothrips
dimidiata, Laxenecera
diminuta, Megaleptoperla
dinoceras, Coeliccia
dioscoridae, Villa
disarmatus, Lestes
disjunctus, Chauliodes
distincta, Carcelia .
distincta, Exorista

248

277
315

228

(figs), 256
49

• 199

• 340

288 (figs)
316 (figs)

. 287

305-308
. 198

• 353
229 (figs)

. 184

• 34°

• 354

• 34°

• 354

• 354

97

35

. 184

• 340

• 274
228, 230

. 184
196

• 199

• 179
. 176
. 179

• 340

• 340

41

. 231

• 194

• 174
. 184

• 354

• 354

• 194

• 340

• 340
. 184

• 259

• 174

• 199
. 184

• 340

• 340
. 184

• 32i (fig.)

95 (figs), 100

240

• 34i

• 179
73. 74 (fig-)

• 195

• 354

• 37

• 43

220,

diversa, Togoperla perpicta

diversipes, Tasmanoperla

diversoides, Winthemia

Dogonia

Dolochoscius

dolorosa, Pacificagrion .

dorata, Euscelidia

doris, Philodicus ....

dorothea, Coeliccia

dorothea, Coenagrion

dorothea, Lestes ....

dorrigoensis, Neosticta canescens

dorsalis, Elattoneura

dorsalis, Kyphopteryx

draconis, Pseudagrion kersteni

drusus, Ommatius . 327 (fig.),

drymo, Isogenus ....

dualis, Cora .....

dubiosus, Goniophthalmus

dubitatus, Hermes

dubium, Pseudagrion

dulitensis, Drepanosticta

dundoense, Pseudagrion

duodecima, Paraphlebia

durbanensis, Macrotermes natalensis

eboracus, Argiolestes griseus .
ecornutum, Agrion
edentata, Voria
ephippiatus, Argiolestes .
ephyre, Chloroperla
erythrogastrum, Heteragrion .
esbenpeterseni, Leptochauliodes
eudytum, Agrion .
eulegnica, Eusthenia spectabilis
eumenoides, Ammophilomima
eumorphophaga, Arrhinodexia
euphorbia, Argia extranea
eutachinoides, Blepharipoda .
evansi, Dinotoperla
evelynae, Notoneura
excelsa, Orolestes .
exclamationis, Austroagrion
exigua, Leptoperla
exilis, Perla ....
extensa, Eusthenia purpurescens
extensipes, Catoptropteryx

falciger, Macrotermes 381,
falcula, Protonemura
fallax, Agrion amaurodytum .
fasciata, Dinotoperla
fasciatus, Chauliodes
fascipennis, Javanita
femorata, Phoriniophylax
fenestralis, Neuromus
ferruginea, Nemoura (Leuctra)
ferruginous, Sialis
festiva, Euscelidia
fieldi, Austrosticta

447

• 34i

• 341

54

269-271
. 223
. 184

221 (fig.)
. 298

• 179
. 184

• 195
176
176

• 341

• I85
328 (fig.)

• 341

• 199

45

• 354
. 185
. 174
. 185

• 197
410

. 197

. 185

53

. 197

• 34i
. 197

• 354
. 185

• 34i
219

35
. 185

• 36

• 34i

• 177

• 195
. 185

. i68

382 (figs), 388
'. '• 185

• 354

57

• 354

• 354
220

• 177

448

filicicola, Oristicta
filigera, Amphinemura .
fimhrii, Solomonthrips

INDEX

105

flavescens, Neoperla
flaviceps, Burmargiolestes
flavipes, Hoplandrothrips
flavomaculata, Leuctra .
flavostriata, Coeliccia
flavotincta, Perla .
fletcheri, Disparoneura .
fletcheri, Neochauliodes .
floccosa, Hyperechia
floridense, Acanthagrion gracile
fontana, Dinotoperla
fontanus, Argiolestes
forcipatus, Ctenochauliodes
forficula, Drepanosticta .
foveolata, Neoperla
fragilis, Ischnura .
francoisi, Gonioscelis .
francoisi, Laxenecera .
francoisi, Oxynoton
fransseni, Bactromyia
fraseri, Coeliccia
fraterna, Leuctra .
fraterna, Nogiperla
fraternus, Chauliodes
frontalis, Carcelia .
frontalis, Isogenus
fuliginosa, Nemoura
fulvithorax, Dasyllina
fulvostigmata, Inocellia .
fumosa, Nemoura .
furunculus, Philodicus
fusca, Dinotoperla
fuscinata, Inocellia
fuscipennis, Anthrax
fuscipennis, Eutachina .
fuscipennis, Leuctra

gangetica, Ischnura
garambensis, Hoplistomerus

342
(fig.), 108 (figs),

IIO, 112 (fig.)

• 342

• 197
.98-99

• 342
. I79

• 342

• 177

• 355

• 251

• 185

• 342

• 197

• 355

• 174

• 342

• 185
278, 279 (fig.)

240
. 286

35

. 180

• 342

• 342

• 355

37

• 342

• 342
. 231

• 355

• 342

• 299

• 343

• 355

72
42

• 343

246
258
323

garambiensis, Ommatius .

gautama, Indagrion

geniculata, Leuctra

glacialis, Nemoura (Brachyptera)

goliath, Termes .

Gonioscelis . . . .

gowdeyi, Haplothrips ..

gracilicornis, Phloeothrips

gracilis, Protosticta ..

grandis, Psalidothrips .

granti, Ischnura ...

granti, Tolmetothrips . 114-116

185

-247.

(fig.)

(fig.)

185

343
343

275-279

. 98

• 174

102 (figs), 103
. 185
117

(fig.)

greeni, Platysticta . . . . .174

greensladei, Solomonthrips 105 (figs), 107-

108 (fig.), 109, 112 (fig.)

griseus, Lagodias

grossa, Eucarcelia .
guichardi, Pseudagrion .
guineaensis, Ecanthothrips
guttatipes, Catoptropteryx

. 261
. 41

. 186
... 96

132 (fig.), 141 (fig.),

J43 (fig-). M5 (fig-). M9 (fig-), 15° (fig-).

163-164

guttiferus, Hermes .... 355

gymna, Laxenecera .... 242

haemastigma, Ischnura .

hageni, Inocellia

hainanense, Coeliccia scutellum

hamadryas, Drepanosticta

hamulata, Argia

hamulata, Isopteryx

harlequini, Oligopon

hawaiiense, Agrion

hearseyi, Protosticta

hecate, Corydalis .

helena, Indolestes .

helenae, Microstylum

Heligmoneura

hemimacquartioides, Isocarceliposis

herberti, Argia

hermanni, Stichopogon .

herus, Macrotermes

hesperus, Bradythrips

heterogamias, Agrion

hingstoni, Capnia .

hirsuta, Carcelia

hokkaidensis, Vibrissina

hokkaidensis, Winthemia

hudsoni, Spaniocercoides

hutsoni, Sturmia . .

hyalipennis, Eutachina .

hybotinus, Oligopogon

Hyperechia .

hyperythra, Alloneura .

. 186

• 355
. 180

• 174
. 186

• 343
. 296
. 186

• 174

• 355

• 195
. 266

302, 303

45
. 186

• 285
388-390 (figs), 391

. 96
. 186

• 343

• 37
53

• 54

• 343

50

. 42
(figs), 296
249-251

• 177

294

ignicollis, Austrosialis .... 355
ignitum, Ceriagrion . . . .186
imitator, Hyperechia . . . -251
immaculipennis, Catoptropteryx 151-153
immsi, Enallagma . . . . .186
imperator, Ommatius . . . -329
imperator, Phorocera .... 48
imperator, Termes .... 432

inamabilis, Corydalis .... 355
inauratus, Hemipenthes • • • 73

incisa, Risiocnemis (Prionocnemis) . .180
incisuralis, Sumatrodexia ... 52
inconspicuella, Sturmia .... 50
inconspicuoides, Sturmia ... 50

indecisus, Hermes ..... 355
indica, Eucarcelia ..... 42
indica, Protonemura .... 343
indicatrix, Argia . . . . .186
indicum, Pseudagrion . . . .186
indicus, Phlaeothrips .... 98

INDEX

449

inermis, Ormothrips .

infectus, Neuromus
infrequentula, Argia
infumata, Sialis
inopinata, Thaumatoneura
institutiimperialis, Leverella .
insularis, Austrolestes
insularis, Chloroperla
insularis, Exoprosopa
intermedia, Erycia
intermedius, Argiolestes griseus
intermedius, Solomonthrips
intermixta, Perla .
internata, Perla
interrupta, Agriocnemis .
interruptum, Acanthagrion
intimus, Neuromus
iola, Laphria
irrorata, Trinotoperla
isabeli, Phorocera .
issikii, Arrhinomyia
ivorensis, Macrotermes .

jacksoni, Pseudagrion
japanica, Protonemoraea
japanica, Tamanukia
japonicus, Chauliodes
jocosa, Cora
jugorum, Agrion .

katangensis, Trichardis

kauaiense, Agrion
kempnyi, Leuctra
kibalense, Pseudagrion .
kibonotensis, Tumulitermes
kimminsi, Idiocnemis
kimminsi, Raphidia
kinabaluensis, Protosticta
klugi, Protoneura .
koreanus, Neochauliodes
kunenensis, Macrotermes bellicosus

lachrymosa, Pacificagrion
lacrimosa, Paragnetina .
lacustris, Eusthenia
ladelli, Eutachina .
laevis, Heligmoneura
Lagodius ....
laidlawi, Teinobasis
lampyroides, Proagonistes
lanceolatum, Agrion
lankanensis, Ceylonosticta
Laphria

larvata, Tetropina
Lasiocnemis .
laterale, Acanthagrion
lateralis, Chloroperla
latiforceps, Sturmia
latistylata, Parexorista .
latistylata, Sturmia

96 lativentris, Rhaebothrips . . .123

355 latratus, Neuromus .... 356
1 86 lawrencei, Ommatidothrips 121 (fig.), 122-123

. 355 Laxenecera 235-243

198 leefmansi, Cadurcia .... 36
46 leios, Baphothrips . . 94, 95 (figs)

195 leoninum, Argiagrion .... 187

343 leoninus, Proagonistes .... 245

79, 90 (figs) lepida, Anacroneuria . . . . 344

41 leptodemas, Agrion . . . .187
197 leucorrhinum, Heteropodagrion Mesagrion 197

105 (figs), no leveri, Nesobasis . . . . .187

343 leveri, Prosopodes ..... 49

343 leveriana, Eurystaea .... 42
1 86 lieftincki, Coeliccia . . . .180
1 86 lilljeborgi, Macrotermes 394~397 (figs), 398

. 355 limpopoensis, Macrotermes bellicosus 398

230 lindgreni, Protosticta .... 194

. 344 lobatus, Congomochtherus 306, 307 (figs)

48 londinensis, Raphidia .... 356
35 longfieldae, Enallagma . . . .187

39I-394 (figs) longicornis, Inocellia .... 356
longipennis, Ommatius .... 329

1 86 longistyla, Esme . . . . .177

49 loogali, Coeliccia . . . . .180
53 louisae, Amphicnemis . . . .187

356 lugubris, Perla ..... 344

199 lunata, Rhabdiopteryx .... 344

187 lunulata, Eusthenia .... 344
luteicollis, Perla ..... 344

248 (fig.), 249 luteicornis, Nemoura .... 344

187 luteipes, Amphinemura .... 344

. 344 lycorias, Perla ..... 344

187 lyelli, Agrion . . . . .187

. 381 lymetta, Drepanosticta . . . 174

. 180

. 356 machaeralis, Hapalioloemus ... 45

174 Machatothrips .... 118-119

177 Machimus ..... 309-311

356 mackwoodi, Caconeura . . . .177

398 maclachlani, Agriocnemis . . .187
maclachlani, Allolestes . . . .197

187 maclachlani, Inocellia .... 356

344 macquarti, Ommatius . . . 320 (fig.)
344 macrophallus, Sturmia . . . .51

42 macroscelis, Ommatius . . . -319
302-303 macrostigma, Coeliccia . . . .180
260-262 Macrotermes .... 368-444

187 maculibasis, Leiosiopsis . . . .46

252 (fig.), 254 maculicollis, Raphidia .... 356

187 maculifera, Hermes .... 356

174 maculipennis, Catoptropteryx . . 161

226-231 maculipennis, Gonioscelis . . -275

. 344 maculipennis, Hermes .... 356

223-224 madelenae, Amphicnemis . . .187

187 magellanica, Kempnyia .... 345

344 magna, Myiostoma .... 48

50 magna, Phorocera ..... 48
48 magnanimum, Pseudagrion . . .188
50 major, Rhabdogaster . . . 280 (fig.)

45°

makoka, Argia
malabaricum, Pseudagrion
malaitae, Sophikothrips
malayana, Carcelia
maldivense, Enallagma .
malekulana, Nesobasis .
malgassica, Tatocnemis .
mallochi, Winthemia
mallochiana, Actia
manevali, Isopteryx
marshalli, Chlorocnemis .
marsyas, Drepanosticta .
maxima, Phorocera magna
maynei, Laphria
mclachlani, Protonemura
media, Chloroperla
media, Perla (Isogenus)
megacephala, Raphidia .
megametta, Drepanosticta
melanicterum, Pseudagrion
melidora, Palaiargia
michaelseni, Termes (Termes)
microdemas, Agrion calliphya
microstigma, Argiolestes
Microstylum
mildredae, Ischnura
minima, Perla
minimus, Argiolestes
miniopsis, Oxyagrion
minor, Arcynopteryx
minor, Psalidothrips .
minor, Spaniocerca
minor, Storthyngomerus
minor, Trinotoperla
minuta, Anacroneuria
mirabilis, Euthore .
mirabilis, Ilia
mirabilis, Pseudolestes .
misema, Laxenecera .
mishmica, Protonemura .
mishuyaca, Argia .
modesta, Helimoneura .
mollis, Bombylius .
mollusca, Argia . . .
molokaiense, Agrion
monardi, Pseudagrion
montana, Capnia .
montana, Coeliccia
montana, Protonemura .
montana, Sumatrodexia
montanus, Neuromus
montivaga, Capnia
moorei, Ceriagrion
morio, Rhipidocephala .
mortoni, Protosticta
mosella.e, Perla
moselyi, Chloroperla
moselyi, Nemura .
mossambicus, Macrotermes

INDEX

. 188

. 188

102 (fig.), 114

37

. 188
. 188
. 197

• 54
54

• 345

• 177

• 174

49

. 230

• 345

• 345

• 345

• 356

• 175
. 188
. 1 88

• 398
. 188

• 197
263-269

. 188

• 345

• 197
. 188

• 345
102 (fig.), 103

• 345

232 (fig.), 234

• 345

• 345

• 199

45
. 198

• 239

• 345
. 188

303, 304 (fig.)
. 68
. 188
. 188
. 188

• 345

. 180

• 346

52

• 356

• 346
. 188
. 287

• 175

• 346

• 346

• 346
398-400 (figs),

401-404

moultoni, Disparoneura . . . .177
moyoensls, Euscelidia . . 222 (fig.)

mozambicanus, Amplitermes . . . 432
mudiensis, Esme . . . . .177
muelleri, Macrotermes . 404 (figs), 406 (figs),

408-410

multinervosa, Devadatta . . .174

munda, Cora . . . . .199

mungomeryi, Eutachina ... 43

munroi, Neolaparus .... 259
mutata, Disparoneura . . . .177
mutata, Thore . . . ._• . 200
mylitta, Drepanosticta . . . 175

Mystrothrips ..... 99

naevia, Gatoptropteryx 132 (fig.), 142 (figs),
144 (fig.), 147 (fig.), 149 (fig.), 150 (fig.), 153-155
nahuana, Argia agriodes . . .188

naia, Agriocnemis . . . .188

nanus, Catoptropteryx 132 (fig.), 142 (fig.),
147, 149 (fig.), 150 (fig.)
natalensis, Macrotermes 377 (fig.), 410-412 (fig.),

413-415

neavei, Proagonistes .... 255
nebulosa, Nemoura .... 346

nemoricola, Coeliccia (Trichocnemis) . 180

Neolaparus ..... 258-260
nesiotes, Agrion . . . . .188
neutralipennis, Catoptropteryx 132 (fig.), 142
(fig.), 144 (fig.), 149 (fig.), 150 (fig.), 157-159

nietneri, Ceylonosticta .
niger, Plagioderophagus .
nigeriensis, Termes .
nigra, Bornargiolestes
nigra, Caconeura .
nigra, Dogonia .
nigra, Elattoneura
nigra, Teinobasis .
nigrescens, Philodicus
nigrescens, Synlestes weyersi
nigribarbis, Sturmia
nigribimba, Laphria
nigriceps, Lestes
nigricollis, Raphidia
nigricosta, Erycia .
nigricoxa, Dinotoperla .
nigrifrons, Leptoperla
nigripes, Bezziomyiobia .
nigripes, Chlorocnemis .
nigripes, Machimus
nigrita, Hyperechia
nigrociliata, Laxenecera .
nigroflavum, Ceriagrion .
nigrohamata, Coeliccia .

267 (fig.)

175
49
374
197
177
270

177
189
299
194

229 (figs), 230

• 195

• 357

41

• 346

• 346

• 36

• 177
310, 311 (fig.)

. 251

• • 243

. 189

181

nigrolineatum, Agrion nigrohamatum

189

nigrospinosa, Catoptropteryx

nigrotibialis, Argyrophylax
nimborella, Protonemoura
nipalica, Calicnemia

143 (fig.), 145
(fig.), 160-161

35

. 346
181

INDEX

niponensis, Perla .

nitida, Nemoura

nitida, Neoperla

nivata, Trinotoperla

nobilis, Agiolestes icteromelas

346
346
346
346
197

nobilis, Macrotermes 415-417 (figs), 418-419
nobilis, Meigenia mutabilis ... 47
novaeguineae, Leverella .... 46

novaehispaniae, Enallagma coecum . .189
nubecula, Isogenus .... 357

Nusa ...... 243-244

nymphalidophaga, Erycia . . .41

obscurus, Lestes . . . . .195
obscurus, Neochauliodes . . -357

obsoletum, Leptagrion . . . .189
occidentalis, Catoptropteryx 132 (fig.), 143
(fig.), 145 (fig.), 149 (fig.), 150 (fig.), 164-165
occidentalis, Neochauliodes sinensis . 357

occipitalis, Gonioscelis . 275-276 (fig.)
oceanica, Sturmia bella .... 50
oceanicum, Megalagrion . . . .189
octava, Carcelia . . . . -37
oculata, Disparoneura . . . -177
oculata, Masicera ..... 46
oculata, Sturmia . . . . .51
Oligopogon ..... 294-297
olivacea, Perla (Isogenus) . . . 347
olivaceum, Ceriagrion . . . .189
Ommatidothrips . . . .120

ophion, Neolaparus . . • . . 259

opposita, Perla (Chloroperla) . . . 347
orbitale, Dolichocolon . . . .40
oresitrophum, Agrion . . . .189
orientalis, Calotheresia (Calotheresiopsis) . 36
orientalis, Corydalis . . . -357
orientalis, Eurospivora .... 43
originata, Polythore derivata . . . 200
orobates, Agrion . . . . .189
oxylepis, Chloroperla grammatica . . 347
Oxynoton . . . . . .286

pachystigma, Allocnemis . . .181

pacificum, Agrion . . . . .189
painei, Sturmia . . . . .51
pallens, Argia violacea . . . .189
pallicornis, Nemoura .... 347
pallida, Chloroperla .... 347
pallida, Nemoura ..... 347
pallida, Smeryngolaphria . . -235
pallidistigma, Calilestes . . . .197
pallidum, Ceriagrion . . . .189
pallipes, Leptogaster . . . .218
pallipes, Nemoura ..... 347
palpata, Erycia . . . . .41
paludensis, Aciagrion . . . .189
paludosus, Lestes . . . . .195
pan, Drepanosticta . . . .175

pandellei, Fabriciella .... 44
paradoxalis, Sturmia . . . .51

parvicornis, Takanoella . . . -53
pauli, Chlorocnemis . . . .177
paulina, Libellula . . . . 175

paulitoyaca, Palaemnema . . .175
pedestris, Capnia ..... 347
peles, Agrion amaurodytum . . .189
pellucida, Thaumatoneura . . .198
pendleburyi, Ceriagrion . . . .190
penicillatus, Congomochtherus . . 308

peramans, Protoneura . . . .177
peramoena, Disparoneura . . .178
Perasis ..... 244-246

percellulata, Argia . . . .190

perfecta, Nemoura .... 347

perparva, Ischnura . . . .190

perpicta, Togoperla .... 348
perplexus, Plesiothrips .... 89
perspicillata, Gibosia .... 348
peruviana, Aeschnosoma . . .174

peruviensis, Protoneura . . . .178
Philodicus, ..... 298-300
phrynoides, Exorista .... 43
picta, Exorista ..... 44
picta, Sturmia . . . . .51
pilosa, Carcelia ..... 37
pilosa, Meigenia mutabilis . . .47
pilosella, Carcelia ..... 37
plagiata, Euthore ..... 200
plana, Argia vivida . . . .190
platystigma, Ceriagrion . . . .190
pliomelas, Proagonistes .... 255
plomleyi, Archichauliodes . . . 357
pollex, Microstyllum . . 266, 267 (fig.)
pontica, Raphidia . . . . -357
popoluca, Argia . . . . .190
poseidon, Xenomyza . 291 (fig.), 292 (fig.)
postica, Perla (Isogenus) . . . 348

poungyi, Coeliccia . . . . .181
praeclarum, Pseudagrion . . . 190
prasina, Chloroperla .... 348
priesnerianus, Haplothrips ... 98
prima, Carcelia . . . . -37
Proagonistes . . . . -255

Promachus ..... 300-301
proteus, Pteronarcys .... 348
prothoracalis, Argiolestes . . .197
prothoracicum, Telagrion . . . 190
protostictoides, Protoneura . . .178
pruinescens, Agriocnemsis . . .190
pruinosa, Metacnemis . . . .181
Psalidothrips .... 100-104

pseudelongatum, Enallagma . . . 190
pseudocaudata, Asiocarcelia 35

pseudocingulata, Leuctra . . . 348
psyche, Lestes . . . . .198
psychidarum, Tricholyga • • • 53

pulchella, Laxenecera . . . 239

pulchella, Sapho ..... 200
pulcherrima, Ceylonicolestes . . . 195
pulchra, Zenilliana .... 58

452

INDEX

pulex, Actia ...... 35

punctipennis, Exoprosopa . . 77, 78 (fig.)
punctulata, Catoptropteryx 132 (fig.), 141 (fig.).
143 (figs), 145 (fig.), 146 (fig.), 149 (fig-). 150 (fig-).

160-163

punctum, Stichopogon . . . 284 (figs)
purpurescens, Euthenia .... 348

pusilla, Nemoura ..... 348

pusillus, Chauliodes .... 357

pyrenaica, Protonemoura . . . 348
pyriformis, Coeliccia . . . .181

quadridentata, Protonemura . . . 348
quadrigerum, Agrion . . . .190
quadrimaculata, Exorista ... 44
quadriseta, Eutachina .... 43
quadrisetosum, Dolichocolon ... 40
quarto, Carcelia ..... 38
quercifolia, Lestes . . . . .195
quinta, Carcelia ..... 38
qninta, Paraphlebia . . . .198

rajah, Teinobasis . . . . .190

ramburi, Indoneura . . . .178

ramulosa, Catoptropteryx 142 (fig.), 144

(fig.), 146 (figs), 148

ranauense, Pseudagrion pruinosum . .190
raphaella, Philogenia . . . .198
rarum, Ischnuragrion .... 191
rasella, Carcelia ..... 38
rasoides, Carcelia ..... 39
rectangulata, Calicnemia . . .181

redimiculum, Proagonistes . . . 255
reedi, Protochauliodes . . . -357
reflexa, Risiocnemis . . . .181
regalis, Pteronarcys .... 349
remiger, Amphicnemis . . . .191
remissa, Protoneura . . . .178
renata, Amphinemura .... 349
reticulata, Eustheniopsis . . . 349

Rhabdogaster ..... 280
Rhipidocephala .... 286-287
rhoadsi, Argia . . . . .191
ricardoae, Microstylum .... 268
ricardoi, Laphria ..... 230
risi, Chorismagrion . . . .193

rodhaini, Heligmoneura . 303-304 (fig.)
rogersi, Argia . . . . .191

rondaniella, Catacarcelia 39

roseanella, Zenillia .... 54

roseonotata, Podopteryx . . .198

rubricauda, Agriocnemis . . .191

rubrocinctus, Selenothrips ... 89
rubroviridis, Pseudagrion . . . 191
rufa, Carcelia ..... 39

rufescens, Chloroperla .... 349
rufescens, Dolichocolon .... 40
rufescens, Leptogaster . . . .218
rufibarbis, Proagonistes .... 255
ruficosta, Tasmanoperla .... 349

rufipes, Dexiomimops .... 40
rufipes, Metacnemis . . . .181
rufitarsis, Laxenecera .... 239
rufiventris, Kosempomyiella ... 46
rufocinctum, Pseudagrion . . . 191
rufofemorata, Erycia . . . .41
rufostigma, Protosticta . . . .175
rufostigma, Pseudagrion . . . 191

rugosa, Leptoperla .... 349

rusticella, Eutachina . . . .43
rutilloides, Carcelia .... 39

saegeri, Dogonia . 267 (figs), 269-270

saegeri, Pegesimallus . 262, 263 (fig.)

salomonica, Euvespivora . 43

salomonis, Alloneura . . . .178
salomonis, Lieftinckia . . . .181
samoensis, Pseudagrion .... 191
sanguinostigma, Protosticta . . 175

saundersi, Thore ..... 200
sauteri, Trichoformosomyia 53

scintillans, Chalcopteryx . . . 200

scutata, Aphanicercella .... 349
scutata, Rhipidocephala . . . 289

scutellata, Xenomyza . . 292, 293 (fig.)
scutellum, Coeliccia . . . .181
scutigera, Protonemura .... 349
Scylaticus . . . . . -274
secunda, Carcelia ..... 39
selysi, Hermes . . . . -357
selysi, Orolestes . . . . 195

selysi, Synlestes . . . . .194
senionvhitei, Exorista . . . .44
septima, Carcelia ..... 39
serricauda, Dinotoperla .... 349
serrifera, Gatoptopteryx 132 (fig.), 143 (fig.),
145 (fig.), 149 (fig.), 150 (figs), 165-167
serva, Trithemis . . . . .174
sessile, Microstylum .... 269
setlfer, Solomonthrips 105 (fig.), 108 (fig.),

Ill, 112 (fig.)

setigerum, Megapodagrion . . . 198
setosella, Carcelia ..... 39
sexta, Carcelia ..... 39
siamense, Euthelairosoma ... 43
sibirica, Sialis . . . . -357
sica, Chiasmella ..... 69
signata, Perla ..... 349
signatipennis, Catoptropteryx . 155

sikkima, Lestes . . . . .195
silenus, Drepanosticta . . . . 175
sillemi, Cnephalia ..... 39
simplex, Chauliodes .... 357
simulatrix, Lestes . . . . 195

sincerus, Mecistogaster jocaste . . 193

sinensis, Chauliodes . . . -357
sinensis, Hermes . . . . -357
sita, Disparoneura . . . .178

skinneri, Cora ..... 200
Smeryngolaphria .... 234-235

snodgrassi, Mecynothrips
sobrina, Telebasis .
socotrae, Bombylius
sokotrae, Petrorossia .
solitaris, Alloneura
solomonensis, Chaetexorista
solomonensis, Doleschalla
solomoni, Atractothrips

. igi
68

70 (fig.)
. 179
39
40
116, 117 (fig.), 121

(fig.)

solomonica, Bactromyia fransseni . . 36
solomonica, Winthemia di versa . . 54
solomonicola, Formosia mirabilis . . 44
Solomonthrips .... 104-113
somali, Systoechus . . . .69

somalicum, Enallagma . . . .191
sonans, Perla ..... 349
Sophikothrips .... 113-114
souteri (Disparoneura) . . . .178
spatiata, Machatothrips . . .118
spectabilis, Eusthenia .... 349
spencei, Pseudagrion . . . .190
spiniceps, Chirothrips .... 89
spinithrips, Ecanthothrips . 92 (figs), 96

spio, Chloroperla ..... 350
steelae, Aciagrion . . . . .190
stellatum, Pseudagrion . . . .190
stevensi, Protosticta . . . 175

Stichopogon 282-285

Storthynogomerus . . . 231-234

strachani, Astochia . . .312 (fig.)
striatus, Ancylorrhynchus . . -274
striatus, Solomonthrips 105 (fig.), 112 (figs),

sturmioides, Ctenophorocera 55

subaequistyla, Copera . . . .181
subarmata, Chloroperla grammatica . 350

subfasciatus, Chauliodes . . . 358

subhyalinus, Macrotermes . 419, 420 (fig.),
421-422 (figs), 423 (figs), 428
subnodalis, Disparoneura . . .178
subnubilis, Protohermes . . . 358

suffusa, Perla ..... 350
sulcicollis, Nemoura .... 350
sumatrana, Eutachina aureifrons . . 42
Sumatrana, Sturmia . . . 51

superciliatus, Oligopogon . . . 297
superplatypes, Copera . . . .181
swynnertoni, Philodicus .... 300

tabaci, Thrips ..... 90
taciturna, Xenomyza . . 288 (figs), 289
takanoi, Actia • • • • • 35
takanoi, Erycia . . . . .41
takanoi, Protonemoraea ... 49

talamanca, Argia . . . . .192
tarascana, Argia . . . . .192
tarsalis, Leptogaster . . . .218
tasmanica, Ischnura heterosticta . .192
tasmanica, Leptoperla .... 350
tasmanica, Spaniocerca . . . 351

INDEX 453

:o temerarius, Philodicus .... 299

tenebrosa, Kempnyia . . . 351

tenuiforceps, Eutachina .... 43
tenuis, Argiolestes griseus . . .198
tenuis, Chauliodes ..... 358
tenuissimus, Austrolestes . . .195

terminalis, Cora ..... 200
terminalis, Perla . . . . 351

terminata, Polythore derivata . . 200

terminatum, Microstigma . . . 193
tertia, Carcelia ..... 39

Thallosia 315

theebawi, Caconeura . . . .178
tibetana, Capnia . . . . 351

tigrina, Rhipidocephala .... 286
tillyardi, Spaniocerca . . . 351

tinctipennis, Erythromma . . .192
tinctipennis, Perla . . . 351

tomentosus, Gonioscelis . . 276-277
tonkinicus, Anachauliodes . . . 358
toroensis, Storthyngomerus 232 (figs), 233
torresiana, Erythromma . . .192

townsendi, Macrozenillia ... 46

tragula, Amphinemura . . . 351

transmarina, Chloroperla . . 351

transversa, Perlodes . . . 351

triangularis, Pericnemis .... 192
tricellulare, Heteragrion . . .198

Trichardis ..... 247-249
tricolor, Orthetrum . . . .174
tricolor, Togoperla . . . 351

tridens, Lestes . . . . .196
tridentatus, Storthyngomerus . . 232 (figs)
trifoliata, Argia . . . . .192
trijuncta, Perla . . . . 351

trinervis, Neurolestes . . . .198
trisetosa, Sturmia . . . . .51
trisetosoides, Sturmia .... 52
tropicus, Synlestes . . . .194

truncatipenne, Anisagion . . .192
truncatus, Neochauliodes sinensis . . 358
tumuli cola, Termes . . . .419
turkestanica, Capnia . . . 351

ugandiensis, Machimus . . .310 (fig.)
ugandiensis, Promachus . . .301

ukuzii, Macrotermes 428-430 (figs), 431-432

ulmeca, Argia . . . . .192
umbratus, Neochauliodes . . . 358
umbriaca, Argia . . . . .192
uncata, Filchneria . . . . • 351
underwoodi, Argia . . . .192

unguicularis, Sturmia .... 52
unicolor, Lestes . . . . .196
uniformis, Dinotoperla .... 352
uniformis, Protohermes .... 358
unisetosa, Sturmia .... 52

unispinus, Thrips ..... 90
usutu, Macrotermes . . . . 381

454

INDEX

vagabondum, Agrion . . . .192
vanderwulpi, Cadurcia .... 36
vansomereni, Enallagma . . . 192

vansomereni, Pseudagrion . . .192
varia, Leptoperla ..... 352
variegata, Nemoura .... 352
varralli, Mortonagrion . . . .192
venosa, Neoperla . . . -352

ventrale, Spogostylum .... 72
vernalis, Echinomyia praeceps . . 41

veronica, Indolestes . . . .196
versicolor, Protosticta . . . .175
versicolor, Telebasis . . . .192
vicinalis, Exorista ..... 44
vicinella, Sturmia . . . . .52
victoria, Agriocnemis . . . .192
victoria, Thore ..... 200
villeneuvei, Hemidegeeria ... 45
violacea, Coenagrion . . . .193
viridis, Drepanosticta . . . .175
vitrialatus, Macrotermes 432-433 (figs), 434-
435 (figs), 436 (figs), 437-440
vittata, Alloneura (Disparoneura) . .178
vittata, Sumatrodexia 53

vittatus, Ommatius . . . -317
vulgaris, Meigenia mutabilis ... 47
vulpinoides, Micropalpus . . -47

waianeanum, Agrion amaurodytum . 193

wainwrighti, Sturmia .... 52

wallacei, Alloneura . . . .178

wallacei, Lestes . .... .196

wallacei, Teinobasis . . . .193

walli, Ceylonosticta . . . .175

waterbergi, Termes .... 432

wheeled, Drepanosticta . , . ,i- .175

whellani, Ceriagrion . . . .193

whellani, Pseudagrion . . . .193

williamsoni, Pseudagrion . . .193

winthemioides, Nemosturmia ... 57

xanthenes, Perla ..... 352
Xenomyza ..... 287-293

yerburyi, Ctenophoroceropsis ... 39
zelandica, Telebasis . . . .193

/ Ci r

A LIST OF SUPPLEMENTS
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3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177 :
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5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World.
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6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso-
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7. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family
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8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the
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9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species
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10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rho-
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11. MOUND, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 172 :
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12. WATSON, A. The Taxonomy of the Drepaninae represented in China, with
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13. AFIFI, S. A. Morphology and Taxonomy of Adult Males of the families
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14. CROSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa
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15. ELIOT, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera :
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