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?, A REVISION OF THE TERMIT 
SUBFAMILY NASUTITERMITINAE 
(ISOPTERA, TERMITIDAE) FROM THE 
ETHIOPIAN REGION 


oS 
y 
2, RS 


W. A. SANDS 


BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Supplement 4 
LONDON : 1965 


A REVISION OF THE TERMITE SUBFAMILY 
NASUTITERMITINAE (ISOPTERA, TERMITIDAE) 
FROM THE ETHIOPIAN REGION 


BY 
W. A. SANDS 


Termite Research Unit, 
Ministry of Overseas Development ° 


BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Supplement 4 
LONDON : 1965 


THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted im 1949, 1s 
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 1s Supplement No. 4 of the Entomolog- 
ical series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 


© Trustees of the British Museum (Natural History) 1965 


TRUSTEES OR 
THE BRITISH MUSEUM (NATURAL HISTORY) 


Issued October, 1965 Price £3 5s. 


A REVISION OF THE TERMITE SUBFAMILY 
NASUTITERMITINAE (ISOPTERA, TERMITIDAE) 
FROM THE ETHIOPIAN REGION 


By W. A. SANDS 


CONTENTS 


INTRODUCTION 4 
MATERIAL , : : : : ‘ - ; : ; 5 
METHODS : : : - : : : 7 
NASUTITERMITINAE Have: diagnosis : : : ; : 8 
Phylogeny . - : : ; : ‘ : : ; 10 
Distribution . 3 , ; . , d < ; : II 
Keys to genera. , , - : ‘ ; : : ; 12 
Nasutitermes Dudley . , ‘ ‘ : - : , : 15 
Leptomyxotermes gen. n. - , . : : : : : 51 
Grallatoteymes Holmgren. : : : : : : : 55 
Fulleritermes Coaton . ‘ : ‘ ‘ : : : ; 55 
Rhadinotermes gen. n. ; : . - : é : ; 71 
Mycterotermes gen. n. 2 2 . . , ; : : 75 
Trinervitermes Holmgren. : : 2 p : : : 76 
Baucaliotermes gen. n. : ; . : z - : 138 
Eutermellus Silvestri . : : “ - : : : : 141 
Afrosubulitermes Emerson . : : é : : - 3 153 
Postsubulitermes Emerson . : ; , 5 é : : 155 
Verrucositermes Emerson. : : ‘ ; : : : 156 
Mimeutermes Silvestri ; : : : , ; : , 158 
Tarditeyrmes Emerson . : ; : - : : : : 166 
ACKNOWLEDGMENTS . : ; ‘ : 4 - : : : 167 
BIBLIOGRAPHY : : , : : : : - ; : 168 
SPECIES INDEX. : F ' ; : E : : : ; 171 


SYNOPSIS 


The subfamily Nasutitermitinae is revised for the Ethiopian Zoogeographical Region. 47 
species are recognized, of which 7 are new, in 14 genera, of which 4 arenew. 77 names are found 
to be synonymous, of which 62 are newly established synonyms. 4 species are removed from 
the subfamily. The phylogeny of the subfamily is discussed, and keys are given to genera and 
species. The distribution of species and genera is related to the vegetation types. Notes are 
provided on the biology of the species. 


4 W. A. SANDS 


INTRODUCTION 


THIS paper provides a revision of the termite subfamily Nasutitermitinae Hare 
(1937) (Isoptera, Termitidae) as it occurs in the Ethiopian Zoogeographical Region. 
The subfamily is characterized by the adaptation of the head capsule of the soldier 
caste to a chemical defence mechanism: the frons and vertex are produced into a 
tubular or conical nose terminating in the opening of the frontal gland, which 
secretes a sticky fluid. In all of the genera included in this paper the soldier man- 
dibles are reduced to vestiges. The more primitive genera with mandibulate 
soldiers do not occur in Africa, being confined to the Neotropical Region. 

The Ethiopian Zoogeographical Region is taken to consist of the continent of 
Africa, south of the Sahara Desert, and those adjacent parts of the Arabian penin- 
sula in which the climate permits the survival of African species. There are no 
unquestionable records of the subfamily from Palaearctic Africa; the Malagasy fauna 
is separate and distinct and is therefore excluded from this revision. 

More material is available for study from the African continent than from any 
other, enabling a more detailed assessment of variation to be made than is possible 
for any other part of the world. All the species included are endemic to the Ethio- 
pian Region, as are all but 3 of the genera. Of the latter, Grallatotermes has only 
one African species; Tvinervitermes is primarily African with only 6 little-known 
species in the Indo-malayan Region; Nasutitermes is the only tropicopolitan genus. 
Monographic treatment of Nasutitermes must needs be unbalanced owing to dis- 
parities between collections from different regions: for example, a recent revision by 
Prashad and Sen-Sarma (1959) of 18 species of Nasutitermes in the Indian Region 
refers to only 39 nest series. For these reasons, this revision of a faunal section of 
the subfamily has been undertaken rather than separate monographs of component 
genera. 

The first African nasute species to be described was Termes arborum Smeathman 
(1781), followed by T. tvinervius Rambur (1842). Sjéstedt (1g00—1904) provided 
the first revisions and keys and recognized 18 species of African nasutes, all placed 
in the genus Eutermes. By the time Sjdstedt (1926) produced his second revision of 
the African termites, the numbers had increased to 88 species and 2 subspecies in 5 
genera. Emerson (1928) listed go species in 7 genera, and Snyder (1949) catalogued 
g2 species and 4 subspecies in 6 genera. Subsequent modifications and additions 
have brought this up to 102 species in II genera. 

In addition to these, species previously synonymized by other authors have been 
re-determined, with the result that a total of 120 previously described species and 
subspecies are involved in this revision. In all but 5 of these, the holotype, lecto- 
type, syntypes, paratypes or paralectotypes from the type colony have been 
examined. Where the type series consists of alate imagos collected at light, as much 
as possible of the original material has been studied to eliminate the risk of confusion 
resulting from a mixture of species. Of the 5 species in which types have not been 
seen, one, Nasutitermes arborum (Smeathman) lacks a type specimen and a neotype 
has been selected. The remaining 4 belong to other subfamilies. These are as 


ETHIOPIAN NASUTITERMITINAE 5 


follows: Trinervitermes bouviert Sjéstedt and T. somaliensis (Sjéstedt) which belong 
to Microtermes (Macrotermitinae), T. sudanicus (Sj6stedt) which is unidentified, and 
Nasutitermes profestus (Sjéstedt) which belongs to Noditermes (Termitinae). These 
determinations are extracted from unpublished notes made by Dr. A. E. Emerson, 
and the species are therefore omitted from further consideration. From the exami- 
nation of the remaining I15 type specimens and series, combined with a study of 
variation in the other material, 77 names have been found to be synonymous, of 
which 62 are newly established synonyms. 

In the subfamily Nasutitermitinae from the Ethiopian Region, I recognize as valid 
47 species, of which 7 are new to science and described in this paper. These belong 
to 14 genera, of which 4 are new, one being completely new to science and the others 
newly formed genera containing existing species. 


MATERIAL 

A total of 2401 nest series from all parts of Africa (Map 1), have been examined, 
each containing from one to several hundred specimens. Much of this material has 
been collected by the members of the Termite Research Unit (Ministry of Overseas 
Development) and is deposited at the British Museum (Natural History). A 
further large collection, made by Dr. A. E. Emerson, is now housed at the American 
Museum of Natural History. Types and other specimens have been studied on 
personal visits or as a result of loans of material, through the courtesy of specialists 
and others mentioned in the acknowledgments, from the following museum 
collections: 

Museum National d’Histoire Naturelle, Paris (Mus. Paris); Musée Royal de 
VAfrique Centrale, Tervuren (Mus. Tervuren); Institut Royale des Sciences 
Naturelles Belgiques, Brussels, Selys Longchamps Collection (Selys Longchamps 
coll., Inst. R. Sci. Nat., Brussels); Institut des Parcs Nationaux du Congo et du 
Rwanda, Brussels (Inst. des Parcs Nat. du Congo, Brussels); Institut Frangais 
d’Afrique Noire, Dakar (Inst. Frang. d’Afr. Noire, Dakar); Museo Civico di Storia 
Naturale, Genoa (Mus. Civ, di Stor. Nat., Genoa); Naturhistorisch Museum te 
Maastricht (Mus. Maastricht) ; Naturhistoriska Riksmuseum, Stockholm (Naturhist. 
Riksmus., Stockholm); Zoologisches Staatsinstitut und Zoologisches Museum, 
Hamburg (Mus. Hamburg); Silvestri Collection, Laboratorio di Zoologia della R. 
Scuola d’Agricoltura in Portici, Naples (Silvestri coll., Lab. Zool. Sc. Agr., Portici) ; 
American Museum of Natural History, New York (A.M.N.H.); National Collection 
of Isoptera, Plant Protection Research Institute, Pretoria (N.C.I., Pretoria). 

The abbreviations given in brackets are those used throughout the text in listing 
the material examined. The British Museum (Natural History), where abbreviated, 
is given as B.M.(N.H.). Where the museum of deposit is given in brackets in the 
text, the specimens concerned have not been examined by the author. Where 
lectotypes are designated, that of the senior synonym in each case is given separately, 
whilst those of junior synonyms are included in the lists of type material examined. 

The material examined constitutes virtually all of that known to exist in museum 
collections. In listing material in the text, co-ordinates of latitude and longitude 
are given for localities not included in the Times Atlas, vol. IV. 


6 W. AS SANDS 


Mari. Collecting localities of Nasutitermitinae. (The Republic of South Africa has been 
collected over so extensively that on this scale there would be little space between indi- 
vidually marked localities.) 


ETHIOPIAN NASUTITERMITINAE 7 


METHODS 


Nearly two-thirds of the specific names in existence have been found to be 
synonyms. Most of the descriptions of species published hitherto are based on 
characters now found to be of little taxonomic usefulness, and omit many which I 
have found of most value. In relatively few cases are they adequately illustrated. 
It has therefore been considered desirable to redescribe the majority of the species, 
and to provide a full set of figures to aid identification. All species originally 
described in languages other than English are redescribed, together with others in 
publications that are rare and likely to be inaccessible. Only in the case of fully and 
recently described species is redescription omitted from this revision, and in some of 
these, additional or modified characters are given for use in conjunction with the 
original descriptions. 

It should be noted that the degree of detail given in the descriptions which follow 
varies from genus to genus. This is intentional, because the characters used vary 
in number and importance; to standardize descriptions on the most detailed pattern 
throughout would be wasteful of space. The layout is however standardized for 
easier comparison. 

Characters used for the first time or found of greater importance than hitherto 
recognized are as follows :— 

Imago: pilosity of head capsule 

proportions of female cerci 
form of molar area of mandibles 
size of ventral scent gland on 5th abdominal sternite of 2 


Soldier: distribution of head and other setae 


The proportions of the 9 cerci and scent glands are used, but in the J, these features 
are less diagnostic since all male cerci are slightly longer and all 3 scent glands 
distinctly smaller than the 9. Marginal indentations of thoracic nota have not in 
general been found of value, nor has imago wing venation. In several genera a very 
wide range of variation has been encountered in all characters, both of measurement 
and shape. In others, the degree of variation has been found to be limited in spite 
of the availability of ample material. Each case of generic or specific status or 
synonymy has therefore had to be judged on its own merits, and not by the standards 
of even closely related taxa. These apparent inconsistencies of treatment are further 
discussed in the section of phylogeny, and under generic and specific headings in 
tie text. 

All figures have been drawn with the aid of a camera lucida, from ethanol preserved 
specimens immersed in ethanol. Only three scales are used, each of the two larger 
being twice the next smaller. All the illustrations of the entire head capsule of 
species on the “ Procornitermes branch’’ are on the smallest scale, and all other 
figures are on the middle scale apart from the imago mandibles of species on the 
“ Paracornitermes branch”’ and all the imago fontanelles, which are on the largest 
scale. Measurements are largely self-explanatory, but it should be noted that in the 
imago, head width is measured from the extremes of curvature of the eyes as seen in 


8 W. A. SANDS 


frontal view of the face, and the length of fore wing includes the stump; in the 
soldier, head width is the greatest width of the head capsule, and depth of head 
capsule is taken in profile, excluding the postmentum and measuring vertically from 
the lower margin to the highest point of curvature of the head capsule, wherever it 
occurs. The measurements are given in abbreviated form in the keys, as follows: 

Head width (across eyes in imago), W; Head length to tip of nose, L; Depth of 
head, D; Length of hind tibia, T,;; Greatest diameter of eye, E; Ocellus, O. 


In describing colouration, a limited number of terms are applied as consistently 
as possible according to an arbitrarily decided scale of intensity from white (un- 
sclerotized) to pitch black, as follows: 


white 
yellow-white 
pale yellow 


yellow 
| 

(with reddish tinge) (without reddish tinge) 
orange-yellow pale yellow-brown 
orange yellow-brown (or very pale brown 

if without yellow) 
ferruginous orange pale brown 
ferruginous brown 
chestnut brown sepia brown 
dark chestnut brown dark sepia brown 
very dark chestnut brown very dark sepia brown 


pitch black 


VEGETATION MAP 


A map is provided of the vegetation zones mentioned in the text. It is based on 
the vegetation map of Africa south of the Tropic of Cancer prepared for UNESCO 
by Keay e¢ al. (1959), differing in that only the boundaries that are known to have 
significance in termite distribution are indicated (Map 2). 


Subfamily NASUTITERMITINAE Hare 


Nasutitermitinae Hare, 1937 : 462. Type-genus, Nasutitermes Dudley, 1890. 


Imago. Labrum broader than long, without transverse sclerotized band, tip partly hyaline ; 
mandibles with apical tooth slightly shorter to distinctly longer than first marginal, left third 
marginal with anterior edge at most only slightly longer than posterior (up to 14 times) ; 
fontanelle commonly slit-like, often bifurcate, sometimes nearly obsolete. Tibial spurs 
2:2: 2 (Syntermes with 3 : 2 : 2, is not African). Styli absent. 

Soldier. Head capsule modified to form a proboscis terminating in opening of frontal gland ; 
mandibles reduced to vestiges, with or without “ points’’. Tibial spurs, 2:2: 2. Styli 
absent. 


ETHIOPIAN NASUTITERMITINAE 


Map 2. Vegetation of Africa showing divisions significant in termite distribution. (1-5), 
montane types ; (6), temperate and subtropical grassland ; (7), moist forest at low 
and medium altitudes ; (8-9), forest-savannah mosaic ; (16), moist woodlands ; 
(17-19), Isoberlinia and Brachystegia- Julbernardia woodlands and savannahs ; (20-21), 
relatively dry savannah-woodlands ; (22), woodland with abundant Colophospermum 
mopane ; (24, 26, and 27), grass steppes ; (25), wooded steppe with abundant Acacia 
and Commiphora ; (28-29), Karoo succulent and subdesert steppe ; (30-31), sub- 


desert steppe, tropical and transitional ; (15, 32), desert and Cape Macchia, and (14), 
in Ethiopia only, evergreen thickets. 


10 W. A. SANDS 


PHYLOGENY 


The subdivision of the Nasutitermitinae into two branches at an early stage in the 
evolution of the subfamily was postulated by Ahmad (1950). His phylogeny was 
based mainly on the “ imago-worker ”’ mandible dentition. A detailed study of the 
soldier mandibles (Sands, 1957) supported Ahmad’s fundamental conclusions, but 
indicated more frequent occurrence of parallel evolutionary trends. Each of the 
two branches was referred to by the name of its most primitive genus, these being 
Procornitermes and Paracornitermes. The genera of the Ethiopian Region are 
divided between the branches as follows: 


“ Procornitermes branch ”’ “ Paracormtermes branch ” 
Nasutitermes Dudley Eutermellus Silvestri 
Leptomyxotermes gen. n. A frosubulitermes Emerson 
Grallatotermes Holmgren Postsubulitermes Emerson 
Fulleritermes Coaton Verrucositermes Emerson 
Rhadinotermes gen. n. Mimeutermes Silvestri 
Mycterotermes gen. n. Tarditermes Emerson 


Trinervitermes Holmgren 
Baucaliotermes gen. n. 

The two branches are, according to Ahmad, characterized by the long straight 
cutting edge between first and third marginal teeth of the left “‘ imago-worker ” 
mandible, and the narrower angle between apical and first marginal in the “‘ Procorn- 
ttermes branch’”’, as contrasted with the shorter sinuate cutting edge, and the 
wider angle between apical and first marginals in the “‘ Paracornitermes branch ’’. 
While these distinctions are correct for the species actually chosen by Ahmad to 
represent the genera, there are many other species belonging to the same genera 
which do not fit this classification so exactly. 

The two branches appear to have arisen because one of them became adapted early 
to feeding on soil-humus, and has remained so, whilst the other was less specialised, 
and its members have diverged to fit many “ ecological niches ’’’. It has been stated 
by several authors including Ahmad that the “ imago-worker ’’ mandibles do not 
exhibit clearly identifiable adaptive characteristics related to the food supply. The 
grounds for disagreement with this statement will be discussed elsewhere; for the 
present purpose it is sufficient to note that the most characteristic adaptation of 
soil-humus feeding termites is the loss of transverse grinding ridges on the molar 
plates, particularly the right one, and their transformation to crushing cups by the 
development of rounded flanges on both sides. This occurs convergently in the sub- 
families Termitinae, Amitermitinae, and Nasutitermitinae, and in the latter is the 
distinguishing feature of the “‘ Paracornitermes’’ branch. In some genera the ridges 
are not entirely lost, remaining as differences in sclerotisation of an otherwise smooth 
surface (Text-figs. 431-450). 

The “‘ Procornitermes branch’ possesses variously modified mandibles, but these 
always have prominently ridged molar plates (Text-figs. 157-196). 

The detailed aspects of phylogeny can only be discussed tentatively in a study 
based on species from only one zoogeographical region. The genus Nasutitermes 


“é 


ETHIOPIAN NASUTITERMITINAE II 


includes species which differ markedly, and show resemblances to genera on different 
subsidiary divisions of the “‘ Procornitermes branch”’. This suggests a polyphyletic 
origin for the genus, which needs to be clarified by a study of all the species. In 
the Ethiopian Region, N. latifrons, N. schoutedent and N. lwjae appear to be derived 
from ancestors close to the genera Hirtitermes and Havilanditermes as suggested by 
Ahmad, whereas N. kempae and N. elegantulus show resemblances to the Neo- 
tropical genera Diversitermes and Parvitermes. Both N. latifrons and N. kempae 
exhibit traces of “‘ instar ’’ polymorphism in soldier and worker. 

Leptomyxotermes doriae from West Africa appears to be most closely related to the 
Hospitalitermes—Lacessititermes group of constricted-headed genera, as is its relative 
Grallatotermes africanus from East Africa. Leptomyxotermes also shows traces of 
soldier polymorphism which have been lost in Grallatotermes. 

The genera Fulleritermes and Rhadinotermes, although they have constricted- 
headed soldiers, are not closely related to Leptomyxotermes and Grallatotermes. 
Their affinities are instead to another subsidiary branch of mainly Neotropical 
genera, including Velocitermes, Tenuirostritermes, and Diversitermes. In Fuller- 
itermes, F’. coatont has distinctly dimorphic soldiers whereas in F. contractus and 
F. tenebricus there are only vestiges of polymorphism, and F. mally: has a single 
soldier form, as does Rhadinotermes. The importance of soldier polymorphism as 
a character varies greatly. In some genera it appears to be on the point of dis- 
appearance and varies between species, whereas in others its presence is well marked, 
and closely related monomorphic forms must be given separate generic status. Such 
is the case with Trinervitermes, which is always polymorphic though showing inter- 
gradations from true dimorphism to near “instar ’’ polymorphism, and Baucalto- 
termes which is truly monomorphic. These genera, and Mycterotermes, also appear 
to be most closely related to the Tenwirostritermes—Diversitermes branch, which 
would therefore seem to be the most important subdivision of the “‘ Procornitermes 
branch ”’ of the subfamily. 

The collection of new material of humus-feeding species on the ‘‘ Paracornitermes 
branch ” has permitted a more detailed assessment of their relationships. Euterm- 
ellus is the most primitive genus of the Ethiopian Region. 

In some respects, such as the distinctly trilobed soldier labrum, and the “ imago- 
worker ’’ mandibles with a sinuate cutting edge between left first and third mar- 
ginals, and the right molar plate with vestigial ridges, it appears to be more primitive 
than the Neotropical Angularitermes, in spite of the long points on the soldier 
mandibles of the latter genus. The relationship of the genera are discussed in detail 
under their own generic headings. 


DISTRIBUTION 


Representatives of the subfamily are to be found in every vegetation zone from 
the depths of the tropical rain forests and mangrove swamps to the fringes of the 
deserts both North and South of the Equator. 

In the humid rain forest areas of West Africa and the Congo, Nasutitermes is the 
dominant genus with ten species feeding mainly on dead wood of all kinds. 


12 W. A. SANDS 


Leptomyxotermes is found in rotting wood, and the humus-feeding genera of the 
“ Paracormtermes branch”’ are Afrosubulitermes Postsubulitermes, Verrucositermes, 
Tarditermes, and some species of Eutermellus and Mimeutermes. In the less moist 
coastal forests and woodlands of East Africa, the fauna is much reduced, with only 
two species of Nasutitermes and one of Grallatotermes. The feeding habits of the 
latter are not known, but it probably also feeds on wood. 

In the tall grass savannah zones adjacent both north and south to the Congo rain 
forest, the grass feeding “‘ Harvester’ genus Tvinervitermes becomes dominant, with 
Fulleritermes in a subsidiary wood-feeding role. Humus-feeders are restricted to 
one or two species of Mimeutermes and Eutermellus, though it is understood that a 
few related species or genera await description from the southern areas. In East 
Africa, Tvinervitermes is present, but Rhadinotermes appears to replace Fulleritermes, 
although their feeding habits are probably rather different. No humus-feeding 
members of the “ Paracormitermes branch’ are known from East Africa. 

As conditions become drier, both northwards and southwards towards the deserts, 
the faunae become more restricted. Fewer species of Tvinervitermes are found, and 
Fulleritermes persists, but the humus-feeding genera are not known beyond the 
limits of the Isoberlinia and Brachystegia-Julbernardia savannah woodlands, types 
17, 18, 19 of Keay e¢ al. (1959). 

The driest areas of all on the fringes of the deserts have relatively very few species ; 
one or two Tvinervitermes occur, and the small related genera Mvycterotermes and 
Baucaliotermes are found in these types in Arabia and South-West Africa respectively. 

More detailed accounts of individual distribution are given under generic and 
specific headings. 

In the distribution maps of individual species, the shaded portions show the full 
extent of the vegetation types from which each species has been recorded. These 
are not always floristically identical North and South of the Congo forest, but are 
considered ecologically equivalent. The shaded parts therefore indicate the prob- 
able potential distribution in addition to the actual distribution to which the species 
is confined by biotic or other ecological barriers. Records of specimens that I have 
examined are shown by black triangles ; those of other authors unsubstantiated by 
redetermination but thought reliable are shown by hollow triangles. 


Krys TO GENERA 

In the keys which follow, certain characters will be more readily appreciated if reference is 
made to the appropriate figures. The separation of constricted-headed from other soldier 
castes is such a case, as also are certain features of the imago mandibles. Although it has 
become customary to refer to “‘ imago-worker ”’ mandibles, and to use those of the worker to 
avoid damaging scarce imago material, they are in fact different. The “ left mandible index ” 
is the distance between left apical and first marginal teeth divided by the distance from first to 
third marginals and expressed in decimals. This index usually differs considerably between 
imago and worker castes, probably indicating the greater degree of food adaptation in the latter. 
In the humus-feeding species the worker index is greater than the imago, and in the others it is 
approximately equal to or smaller than the imago. In this revision, the imago mandibles have 
been used wherever possible, and where worker mandibles have to be used this is noted in the 
text. The key to imagos has however been arranged so that it can also to a large extent be used 
to identify workers in spite of their differences. Measurements are given in millimetres. 


ste) 


ETHIOPIAN NASUTITERMITINAE 13 


IMAGOS 


Left mandible with apical and first marginal teeth subequal, or apical shorter, left 
mandible index 0-2—0-6 ; molar plate of right mandible with prominent ridges 
(Text-figs. 157-196) : 

Left mandible with apical tooth longer than first siccipiaal, left mandible index 
0-9-3°2 ; molar plate of right mandible without ridges, or ridges vestigial in 


NN 


shining smooth surface (Text-figs. 431-450) . , 8 
Left mandible with cutting edge between first and third marginals entire, straight or 

at most slightly sinuate - 3 
Left mandible with cutting edge betw een first and third marginals indented i in — 

of third marginal. 6 


Right mandible with posterior cutting edge of first marginal equal 'to or shorter than 

posterior cutting edge of second marginal ; molar plate with 9-11 regular ridges 
NASUTITERMES (p. 15) 

Right mandible with posterior cutting edge of first marginal longer than posterior 
cutting edge of second marginal ; molar plate with 5-9 irregular ridges. 4 

Smaller, W, under 1-28 ; E, under 0°38 ; T;, under 1-63. Right molar plate ith 

indented margin at basal end. Colour very dark brown to pitch black 

RHADINOTERMES (p. 71) 

Larger, W, over 1-35; E, over 0-42; T;, over 1-63. Right molar plate without 

pronounced irregularity of outline in surface view. Colour paler, to sepia brown 


or dark chestnut brown - 5 
Pilosity of head capsule evenly fine, setae arising from small sockets of regular size 

with a few larger sockets scattered among them : TRINERVITERMES (p. 76) 
Pilosity of head capsule uneven, sockets of setae varying from small to large, the 

latter numerous and conspicuous . : . BAUCALIOTERMES (p. 138) 


Right mandible with molar plate bearing 13-14 ridges and without pronounced 
irregularity of outline in surface view. Postclypeus weakly inflated, width 3-0 
times length : : GRALLATOTERMES (p. 5 

Right molar plate with os 10 ridges, and with indented margin at basal end. Post- 
clypeus strongly inflated, width 1-7—2-1 times length : , 

Eyes large compared with head width, W/E index 2-4. Fontanelle slit-like, not 
distinctly bifurcate anteriorly. Left mandible with subsidiary tooth between 
third marginal and molar prominence not visible in the gap between them in 
surface view. Colour pale brown . ‘ LEPTOMYXOTERMES (p. 51) 

Eyes smaller compared with head width, W E index 3:2—4°5. Fontanelle slightly to 
distinctly bifurcate anteriorly. Left mandible with subsidiary tooth visible in gap 
between third marginal and molar prominence in surface view. Colour dark 


“ 


brown to pitch black . ‘ : FULLERITERMES (p. 55) 
Left mandible with cutting edge betw een first and third marginals indented in front 

of third, and sinuate ; left mandible index under 1-4 : 9 
Left mandible with third marginal absorbed in short cutting edge ‘behind first ; left 

mandible index over 2-0 : ? II 


Very small, W, 0-56 or less ; Ts, 0-71 or Jess. “Head shining, pilosity reduced to 
paired prominent setae on vertex, frons and postclypeus. Pronotum, length at 


least three-quarters width . : : AFROSUBULITERMES (p. 153) 
Larger, W, 0:82-0:99 ; Ty, over I-00. Head with scattered fine and longer paired 

setae. Pronotum, length less than three-quarters width . ; 10 
Smaller, W, 0-82 ; E, 0:27; T3, 1-00. Pilosity finer, more sparse, surface of head 

shining. Fontanelle a pale streak. ‘ ‘ . VERRUCOSITERMES (p. 156) 


Larger, W, 0:85-0:99 ; E, 0:29-0:38 ; T,, 1-00-1-20. Pilosity dense, surface of 
head almost obscured. Fontanelle triangular, dumb-bell-shaped, or nearly 
obsolete. : : é = : : : . EUTERMELLUS (p. 141) 


14 


irae 


WW. A. SANDS 


Posterior margin of head capsule narrowly rounded, width behind eyes distinctly less 

than length to front of postclypeus ; eyes not set out from sides of head. Smaller, 

W, 0-80 ; pronotum, width, 0-70, length, 0-42 POSTS UBULITERMES (p. 155) 
Posterior margin of head capsule broadly rounded, width behind eyes approximately 

equal to length to front of postclypeus ; eyes slightly to distinctly set out from 

sides of head. Larger, W, 0:89 or over ; pronotum, width 0-78 or over, length 

0:61 or Over : é 5 : : : ¢ 5 : : : : 12 
Larger, W, 1:28-1°35 ; E, 0-41-0-42 ; pronotum, width 1:20-1:23, length 0-68— 

0:75 ; I3,1-45-1°56. Pilosity sparse, surface of head shining. Colour, head and 

pronotum yellow 3 : c ; : : . TARDITERMES (p. 166) 
Smaller, W, 0:89-1:12; E, 0:29-0:35; pronotum, width, 0:78-1:35; length, 

0:61-1:15 ; T3, 0:82-1:06. Pilosity dense, surface of head almost obscured. 

Colour uniformly dark sepia brown 3 : F . MIMEUTERMES (p. 158) 


SOLDIERS 


Head capsule more or less constricted or contracted behind antennae (Text-figs. 


II9Q—156, 388-395, 414-430, 464-469) . : < ‘ : : 2 
Head capsule not constricted or contracted behind porate (Text-figs. 24-99, 

256-296, 338-378, 396-398, 470— ak 497-— cn : , 5 < : : iB 
Antennae 14 segmented . : : : : ; ; : 3 
Antennae 11-13 segmented : ; ; : c : ; : : 5 
Vestigial mandibles without points . . TRINERVITERMES minor soldiers (p. 76) 
Vestigial mandibles with distinct points. : : : : : : 4 


Nose slender, cylindrical. Smaller, T,, 1-23-1- a. Colour, yellow to orange 
LEPTOMYXOTERMES (p. 51) 
Nose thick, conical, sharply tapered and slightly turned down at tip. Larger, T;, 


1:96-2:05. Colour dark brown . : GRALLATOTERMES (p. 55) 
Head in side view raised behind Ganstriction, above line of nose profile including 

basal hump if present . : 6 
Head in side view with undulating or convex poate 40) sed bate Geseceoe 

above line of nose . a 


Vestigial mandibles without marae Vertex of head ene eee ce 
(sometimes a pair of minute setae present). Constriction of head weak 
TRINERVITERMES minor soldiers (p. 76) 
Vestigial mandibles with small to distinct points. Vertex with single pair of 
bilaterally placed setae. Constriction of head strong (Text-figs. 130-156) 
FULLERITERMES and RHADINOTERMES (p. 55) 
Nose shorter than rest of head capsule, measuring from hind margin of antennal 


socket. Soldiers dimorphic : : : MYCTEROTERMES (p. 75) 
Nose longer than rest of head capsule, eer from hind margin of antennal 
socket. Soldier monomorphic : 8 


Nose and anterior lobe of strongly sureties head eapsale eh numerous paper 
like outgrowths of cuticle, also present on basal antennal segments 
VERRUCOSITERMES (p. 156) 
Nose and head without such outgrowths . : ‘ 9 
Labrum with anterior corners extended as short oes maidclle a sani enn 
bowed outwards producing trilobed appearance. Legs with long prominent setae, 
I on each coxa, 3-6 on each femur and tibia, among shorter more typical setae. 
Nose somewhat rugose ; : . EUTERMELLUS (p. 141) 
Labrum with anterior margin rounded or at most sinuate. Legs without such setae, 
or, if one or two present then soldier of minute size, L, 0:98-1:07 ; T3, 0:47—0°53. 
Nose smooth . : . : : . 5 : : 5 : ; 10 


ETHIOPIAN NASUTITERMITINAE 15 


to Smaller, L, 0-98—-1:07 ; W, 0:37-0°41 ; T3,0°47—-0'53. Nose shorter in proportion to 


rest of head capsule (53 : 47) : : - AFROSUBULITERMES (p. 153) 
— Larger, L, 1-39-1°53 ; W, 0:55-0:60 ; T;,0-71-0-79. Nose longer in proportion to 
rest of head capsule (59 : 41) ’ : POSTSUBULITERMES (p. 155) 


11 Head capsule in plan view tapering continuously from near back to nose tip, top- 
shaped or tear-drop-shaped. Nose tip without prominent setae 
MIMEUTERMES (p. 158) 
— Head capsule in plan view not so, nose distinct from rest of head. Nose tip with 4 or 


more prominent setae . : ; : s 2 . 3 ° ; ’ 12 
12 Vestigial mandibles with points a : = : . NASUTITERMES (p. 15) 
— Vestigial mandibles without points . : : : : : : ; d 13 
13 Labrum with sinuate anterior margin. Head capsule in plan view a rounded 

rectangle wider than long. ; . TARDITERMES (p. 166) 
— Labrum with rounded anterior margin. ‘Head capsule in plan view, various, if 

wider than long, then not a rounded rectangle 3 14 


14 Head capsule without prominent setae except on nose and occasionally one pair of 
small setae on vertex. Where minute setae are present, sockets not visible at 
normal stereomicroscopic magnifications. Soldiers dimorphic or polymorphic 
TRINERVITERMES (p. 76) 
— Head capsule with one pair of setae on vertex, one or two pairs near base of nose, 
and four at nose tip with smaller setae behind them. Entire head capsule and nose 
with scattered minute setae with distinct pale sockets. Soldiers monomorphic 
BAUCALIOTERMES (p. 138) 


NASUTITERMES Dudley, 1890 


Nasutitermes Dudley, 1890 : 158. Type-species, by subsequent designation (Emerson, 1925 : 
379), Nasutitermes sanchezi (Holmgren, 1910) = Nasutitermes costalis (Holmgren, 1910). 


In a recent paper on this genus as it is represented in the Indian region, Prashad 
and Sen-Sarma (1959) attribute the authorship of the generic name to Banks (1920). 
They give as their reasons for this change the opinions that (i) the description 
“soldiers with beak ”’ as given by Dudley (1890) “ does not constitute an indication, 
a definition, or a description’, on the grounds that the description is applicable to 
most genera in the subfamily, and (ii) “‘ the indirect evidence from the termitophiles 
is untenable ’’.... Because the definition “ soldiers with beak ’’, though meagre, 
satisfies the requirement of Article 12 of the International Code, and because all the 
other nasute genera at present recognized came into being after the establishment of 
the genus Nasutitermes by Dudley, many of them having been included in it as 
subgenera or species when first described, these arguments are not acceptable. Not 
only is the definition of the genus valid as it stands, but it is supported by its author’s 
reference to his own earlier paper of 1889, in which a detailed description of nest 
structure and habits was given, together with illustrations of all castes (pp. 107, 
fig. 7, and 108, fig. 8). These fully satisfy the requirements of Article 16 (a), 
sections (i) and (viii). The evidence of the termitophiles is irrelevant to the dis- 
cussion of the authorship of the genus. 

Dudley neither designated a type-species, nor included any nominal species in the 
genus. Whilst there is little doubt from the evidence of the termitophile Tevmto- 
gaster insolens Casey, mentioned by Dudley, and now known to be host specific, 


16 WAL SAN DS 


that the species concerned was N. corniger (Motschulsky, 1855), this does not con- 
stitute a valid citation of the species under Article 69 (a). Prashad and Sen-Sarma 
are therefore correct in rejecting the designation of N. corniger as the type-species 
in Snyder (1949). 

The first use after Dudley of the generic name Nasutitermes was made by Banks 
(1919) when eight species were cited by name ; from among these Banks (1920) 
proceeded to designate as type-species Nasutitermes morio (Latreille, 1805). Emer- 
son (1925) stated that the species name Termes morio Latreille was a junior primary 
homonym of Termes morio Fabricius (1793) (now placed in synonymy with Copto- 
termes testaceus (L.)), and had to be replaced by the next available name Nasutitermes 
costalis (Holmgren). This he considered to be synonymous with N. morio (Latreille), 
and therefore was to be the type-species of the genus. The description in Latreille 
(1805), referred to by both Banks and Emerson is given under the heading 
“TERMES MORIO; Termes morio. Fab.’’. Clearly Latreille identified his 
specimens as Fabricius’ species, and in so doing did not create a new species. 
Though the identification has since proved erroneous, the description cannot be held 
to have designated a new species since no alternative name was proposed. ‘‘ Termes 
morio Latreille’’ has therefore never existed as a valid nominal species, and the 
specific name movio cannot be retained for the species which was before Latreille 
(Article 49). Banks’ designation of this as the type-species of Nasutitermes was 
thus in error and must be rejected in favour of the first subsequent designation. 
Nasutitermes costalis (Holmgren) was not among those cited by name by Banks (1919) 
and was therefore not directly available for designation, Article 69 (a) clauses (i) 
and (ii). 

Article 69 (a) clause (iv) states: “ (iv) If an author designates (or accepts 
another’s designation) as type-species a nominal species that was not originally 
included, and if, but only if, at the same time he synonymizes that species with one 
of the originally included species, his act constitutes designation of the latter as 
type-species of the genus’. 

Emerson (1925) synonymized a number of species described by Holmgren (1910) 
and chose from among them the name N. costalis for the species. One of those 
synonymized was N. sanchez, which was among those originally included in the 
genus by Banks (1919). Thus the type-species of Nasutitermes in accordance with 
Article 69 of the International Code must be taken to have been designated by 
Emerson (1925) as Eutermes sanchezt Holmgren, 1910 (=Eutermes costalis Holmgren, 
1910). Nasutitermes costalis (Holmgren) is therefore only type-species by virtue of 
its containing N. sanchezi (Holmgren) as a synonym. The Holotype of N. 
sanchezt was reported by Weidner (1955) to be a soldier caste, in the Zoologischen 
Staatsinstituts und Zoologischen Museums, Hamburg. 

The generic name Nasutitermes was first applied to species from the Ethiopian 
Region by Emerson (1928), who dealt with some species formally, and some in- 
formally, without references to their author’s names, in comparisons with the 
formally treated species. I have thought it best in making out specific synonymies 
to refer the first use of ‘‘ Nasutitermes’’ to Emerson (1928) only in those species 


ETHIOPIAN NASUTITERMITINAE 17 


dealt with formally in that paper, although he clearly regarded all such related 
species as belonging to the same genus. 

Imago. Left mandible with apical tooth as long as or slightly shorter than first marginal, 
distance from apical to first marginals less to slightly more than half distance between first and 
third marginals (left mandible index -37—-54), second marginal obsolete or only represented by 
slight undulation of cutting edge between first and third marginals ; right mandible with apical 
and first marginals subequal as left, posterior cutting edges of first and second marginals 
approximately equal in length, molar plate with 9-11 regular transverse ridges, and no marked 
irregularity of outline in surface view (Text-figs. 157-162). Postclypeus, length slightly less 
than half width, to a quarter or less of width. Fontanelle, usually pale coloured, slit-like, and 
forked anteriorly, but in a number of species virtually obsolete, and scarcely different in colour 
from rest of head capsule, sometimes visible only as an indistinct round or oval mark. 

Soldier. Monomorphic, though sometimes size differences appear to correspond to instars 
at which metamorphosis from worker caste took place. Nose a weak to strong cone, not 
cylindrical in species of Ethiopian Region. Vestigial mandibles usually with points. Head 
capsule without any pronounced constriction behind antennae. 

The characters given in the diagnosis are those which distinguish the genus from 
others in the Ethiopian Region, and are not intended to be diagnostic on a world 
scale, neither are the more generalized subfamily characteristics included. Only 
Grallatotermes among the Ethiopian genera has a right molar plate which resembles 
that of Nasutitermes, though certain Neotropical and Oriental genera are similar. 
Grallatotermes africanus Harris differs from Nasutitermes in having a long cutting 
edge to the right second marginal, compared to the space between right apical and 
first marginal, and a deep indentation of the cutting edge on the left mandible 
between first and third marginals. The soldier also has a constricted head capsule. 

The genus is confined in the Ethiopian Region to rain forest, riverain gallery forest, 
and forest outliers arising from locally moister conditions, or to moist woodland 
approaching forest conditions and usually situated near forest. This has resulted 
in a comparatively sharp division into two faunistic groups of species, one of which, 
the larger, consisting of ten species, is confined to the forests of the Congo and 
Guinean zones, including northern Uganda. The second consists of two species, 
mainly found in the coastal forest and woodlands of East Africa, but extending 
into Zululand in the South, and one of the two up the Rift Valley through Nyasaland 
and the western border of Tanganyika to the borders of the Congo and Uganda at 
Rwindi. The distribution of the species is shown on the maps (Maps 3-12). 


KEYS TO SPECIES 


The increased understanding of the variability of all castes has led to difficulties in constructing 
suitable keys to species. In the case of the imagos, it has been found necessary to resort to the 
use of numerical indices to express differences in proportions where actual measurements overlap. 
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. 

IMAGos 


1 Postclypeus width 2-3 times length (Text-figs. 1-5, 8-17) . ‘ : : 2 
— Postclypeus width more than 3 times length (Text-figs. 6, 18-2 3). 
2 Eyes relatively larger in proportion to head-width, W/E index under 2 es ocelli 

closer to eyes, O-E distance o-o1-o-o4mm. 2 : ; : : : 3 


aoln 


We. A. SANDS 


Eyes relatively smaller in proportion to head-width, ini index over 2°8, ocelli 
further from eyes, O—-E distance 0-04—0-:13mm. : ; : 
Smaller, W, 1:04-1:18 ; E, 0:41-0:51 ; O, 0-09-0-II X O-II-O: iA" ; T, 1:25-1-38 


(W. Afr., Congo, Uganda) : : elegantulus (p. 
Larger, W, 1:20-1:36; E, 0-48-o- oe ; O, oO: =e te SSE a 20; Ts, 1-38-1°64 

(BiCentralyS: Ai:) : : . kempae (p. 
T, shorter in proportion to W, T,/w facies I-10. . : : : sa (p. 


T, longer in proportion to W, T,/W index over 1-15 
Lateral extensions of “‘ fork’ of fontanelle wide, reaching half an to ocelli_ 


camerunensis (p. 
Lateral extensions of “‘ fork ’’ short . : - : ‘ : : : : 
Cerci of female shorter than width at base. : : : : arborum (p. 
Cerci of female as long as width at base c : : : infuscatus (p. 
Smaller, E under 0-42, W under 1-33 : : : : : ; lujae (p. 


Larger, E over 0-44, W over 1-33 


Smaller, W, 1:35-1:40 ; T, under 1-68 ; : : ; : schoutedeni (p. 


Larger, W over 1:42 ; T, over 1°75 
Head capsule width stig pai behind — apprixioately equal t to length te 


front of postclypeus . latifrons (p. 


Head capsule width behind Ss slightly Tess ten length to ont ae postclypeus 


chrysopleura (p. 


SOLDIERS 
Whole head capsule and nose with numerous scattered long yellow setae 


hirticeps (p. 


Head capsule with few (o—-8), more eee aie setae, nose with setae con- 
centrated at tip 

Tip of nose markedly ieee (ext: fee, 40, = oe. 70, a 80, 82, or, 96) 

Tip of nose with 4 prominent setae, and very few small inconspicuous setae behind 
them, or none (Text-figs. 26, 29, 35, 47, 52, 89) 

Postmentum with setae scattered over entire surface Gomeeme nn and smi. 


No long setae on vertex (Text-figs. 78-87) : : latifrons (p. 


Postmentum with setae confined to anterior margin. = bilaterally eee sind 
setae usually present on vertex 

Antennae 11-12 segmented 

Antennae 13-14 segmented : 

Head profile convex from change of epatour at Thee a nose peeeeed or come 
straight with slight over-all ee: numerous small setae on vertex (Text- 


figs. 39-45) : : : elegantulus (p. 


Head profile more or less concave, nseulby slightly pocaienle: so at base of nose ; 


vertex without such small setae (Text-figs. 51-57) . : fulleri (p. 


Nose, measuring from hind margin of antennal socket, shorter fae rest of head 
capsule : : 

Nose, measured as phone orca to or “longer than west of head eapsale ¢ 2 . 

Head profile concave ; nose seas be at tip (Text-figs. 95-99) (W. Africa, 


Congo, Uganda) . .  schoutedeni (p. 


Head profile more or ees. straight : nose more piendece ae evenly tapered to tip 


(Text-figs. 62-68) (E. and Central Africa) ; : infuscatus (p. 


Smaller, T, less than 0-90, head profile convex from Gramps of contour at base of nose 
backwards, or nearly straight with slight over-all convexity (Text-figs. 39-45) 


elegantulus (p. 


Larger, T,; over 0-90, head profile straight or more or less concave, or sinuate, or 
straight to well behind base of nose with raised vertex, rarely with very slight 
over-all convexity, and these much larger, L over 1-61 


SS oth SEATS) 
Bi og ed Sag D 


Ke) 


aN 
ae 


27) 


nun 


ETHIOPIAN NASUTITERMITINAE 


19 


Fics. 1-23 : 


fulleyi (unique morphotype, 12 reconstructed from 11) ; 


N. kempae ; 


Nasutitermes, front and side views of imago head capsule. 
4, 5, N. camerunensis ; 6, 7, N. chrysopleura ; 8-10, N. elegantulus ; 11-13, N. 


1-3, N. arborum; 


14, 15, N. infuscatus ; 16, 17, 


8, 19, N. latifrons ; 20, 21, N. lujae ; 22, 23, N. schoutedent. 


20 We ears AUN GDS 


9 Larger, L, 1-61-2-05 ; W, 0:88-1:29 ; D, 0-61-0-82 ; T;, 0-90-1-28. Head profile 
straight or slightly concave ; rarely with slight over-all Sec Aaa (Text-figs. 
69-77) (E. and Central Africa) : : ; : ‘ . kempae (p. 39) 

— Smaller, L, 1:25-1:63; W, 0:68-0:95; D, 0:49-0:63; Ts3, pee 1:18. Head 
profile straight to well behind base of nose, with vertex raised and rounded 


(Text-figs. 88-94) (W. Africa, eee aia é : : : lujae (p. 45) 
io Antennae 12 segmented . c : : ; : : fulleri (p. 34) 
— Antennae 13 segmented . : II 


11 Nose distinctly shorter than rest of head eens Seeeee fade hind meee aE 
antennal socket, strongly to very strongly conical in profile (cone angle 18-30 
degrees) ; 2-4 prominent setae on top of head at base of nose. Colour, head 
capsule ferruginous to chestnut brown, nose brownish black with red tip. (Text- 
figs. 46-50) . : . ferranti (p. 32) 

— Nose approximately cael to or eee fon rest of fase Sapsule, ely to strongly 
conical (cone angles 10-25 degrees) ; no such prominent setae present. Colour, 
yellow to brown, nose sometimes darker, to chestnut brown. ; < E EZ 

12 Head profile evenly concave ; nose weakly to moderately conical (angle 10-15 
degrees). Colour brown, often with darker nose, sometimes banded around its 
base (Text-figs, 34-38) . : : . diabolus (p. 29) 

— Head profile straight, slightly Soe or aed ay well behind base of nose ; 
nose moderately to strongly conical (angle 13-25 degrees). Colour orange yellow 
to ferruginous orange, nose ferruginous to chestnut brown : 13 
13 Vertex with two long bilaterally placed setae. Nose tip usually mae severe 
smaller setae in addition to the four prominent apical setae (Text-figs. 88-94) 
lujae (p. 45) 
— Vertex without setae. Nose tip with four prominent apical setae only, or with one 
or two very inconspicuous additional smaller setae (Text-figs. 24-33) arborum (p. 20) 


Nasutitermes arborum (Smeathman) 
(Text-figs. 1-3, 24-33, 157-159, 198-199 ; Map 4) 


Termes avborum Smeathman, 1781 : 141, 161. Type locality : SIERRA LEONE. Type material 
lost or destroyed. 

Eutermes maculiventris Sjostedt, 1904 : 104. Type locality : GABON. syn. n. 

Termes (Eutermes) maculiventris (SjOstedt) Desneux, 1904a : 43. 

Eutermes (Eutermes) maculiventris (SjOstedt) Holmgren, 1912 : 62. 

Euteymes infuscatus var. peyfusca Silvestri, 1914 : 35. Type locality : Danomry, Cotonou. 
syn. n. 

Nasutitermes arboreus (Smeathman) Snyder, 1949 : 266. 

Nasutitermes infuscatus var. perfusca (Silvestri) Snyder, 1949 : 280. 

Nasutitermes maculiventris (Sj6stedt) Snyder, 1949 : 285. 

Nasutitermes torquatus (Sjostedt) ; Sands, 1957 : 12 


Imago. Head capsule dark sepia brown, area between eye and base of mandibles, and post- 
clypeus paler, yellow-brown to brown, postclypeus with darker brown shading from mid-line 
outwards to a varying extent. Labrum yellow to yellow-brown. Antennae yellow-brown to 
brown. Pronotum dark sepia brown with slightly paler “‘ T ’’-shaped mark in middle of front 
half. Meso- and metanota, pleural and ventral thoracic sclerites yellow-brown to brown. 
Legs yellow to yellow-brown, slightly darker at ends of femora and bases of tibiae, yellow-brown 
to brown. Abdominal tergites and lateral parts of sternites sepia brown, middle of abdominal 
sternites, excepting posterior two or three, yellow to yellow-brown. Wings translucent brown, 
venation sepia brown. 


ETHIOPIAN NASUTITERMITINAE 21 


Posterior margin of head behind eyes distinctly less than semi-circular ; fontanelle variable, 
in g, from a short whitish streak, slightly bifurcate at anterior end, to an indistinct mark only 
slightly paler than rest of head capsule, and in 9, from a whitish slit to an indistinct pale patch, 
in both sexes flat to very slightly depressed posteriorly ; eyes prominent in proportion to own 
diameter, though not large relative to head width, W/E index 2-8—3-5 (only one specimen 
recorded under 3:0), nearly circular to angularly oval ; ocelli small to medium sized, separated 
from eyes by one-third to slightly more than own least diameter, oval ; postclypeus moderately 
inflated, width 2-5 to 3:0 times length, anterior margin nearly straight to distinctly sinuate, 
posterior margin convex, rounded, slightly sinuate towards outer corners ; antennae 15 seg- 
mented, II slightly longer than IV, III and V subequal, slightly shorter than IV, III slightly 
narrower than V. 


Cerci of 9 distinctly shorter than width across base, with somewhat mamilliform tip. 
Measurements (25 specimens from 7 localities) in millimetres. 


Range Mean 
Head width across eyes I-16-1-50 ; ; 1+32 
Greatest diameter of eye 0°35-0°48 3 ; o-41 
Ocellus : ; - 0:09-0°14 X 0°13-0°20_ . : O:II X 0-16 
Ocellus toeye . : 0:04.0°10 - : 0-08 
Width of pronotum 0-93-1°18 : : 1-04 
Length of pronotum . 0°58-0°78 : 4 0:66 
Length of hind tibia. 1-38-1-80 : : 1°57 
Length of fore wing. 9°40-14°70 ; ; 11-40 


Soldiers. Head capsule orange to ferruginous-orange, nose ferruginous to chestnut-brown. 
Pronotum and antennae as head capsule. Abdominal tergites and femora of legs yellow, 
abdominal sternites, tibiae and tarsi pale yellow. 

Head capsule in plan view nearly circular, slightly tapering towards front, nose conical. In 
profile very nearly straight, sometimes very slightly concave or sinuate, nose equal in length or 
slightly longer than rest of head capsule, measuring from hind margin of antennal socket, angle 
of nose cone 15-25 degrees ; postmentum slightly and evenly convex ; antennae 12-13 seg- 
mented, usually 13, proportions of segments somewhat variable depending on size, II and IV 
usually subequal, shorter than V, which may be shorter than or equal to III. Setae on head, 
excluding appendages, confined to four at tip of nose, with sometimes a very few small fine 
setae immediately behind them, and a few on anterior margin of postmentum. 

Abdominal tergites with scattered minute fine setae, and longer setae on posterior margins of 
last 3-4 segments ; sternites with shorter scattered setae, and long setae on posterior margins. 

Measurements (157 specimens from 32 localities) in millimetres. 


Range Mean 
Head length to tip of nose. I+33-1°89 : : ; } 1°58 
Head width - ? ; 0:75-1°13 : ; , F 0:93 
Depth of head capsule . : 0+50-0'75 : : F 4 0-62 
Width of pronotum =... : 0°40-0°55 : : : ; 0-48 
Length of pronotum . d 0°18-0:25 : : ; - 0:20 
Length of hind tibia. : 0:93-1°39 : d : ‘ I-13 


Variation. Imagos from the eastern end of the range tend to be slightly larger and to have 
the postclypeus less clouded with dark brown than those from the west coast of Africa, but the 
soldiers remain closely similar throughout the range. Soldier variation consists mainly of 
differences in the thickness of the nose cone, and of minor alterations in the angle of the nose and 
the degree of straightness of the profile. 


The material from which Smeathman (1781) described this species has never 
been traced, and was probably never sent to any museum. Griffin (1942) records 
that the specimens of nests sent to Drury by Smeathman arrived in fragments, and 


22 W. A. SANDS 


“mouldered to dust ’’, so that no type material either of the termite or its nests 
now exists. 

The commonest truly arboreal species in West Africa has been given various names 
which appear in the synonymy. It has been collected from a number of places in 
Sierra Leone, the type locality for N. arborum, and I consider it to be the species 
most likely to have been described by Smeathman under that name. 

The name “ Eutermes arborum (Smeathman) ’”’ has been used by several authors, 
but these references have all been checked back to specimens first identified by 
Sj6stedt (1900) and, following him, Silvestri (1912). These have been examined, 
and found to belong to WN. latifrons (Sjéstedt, 1896). It is most undesirable that the 


48. 


Fics. 24-57: Nasutitermes, soldier head capsule, side and plan views, nose tip and post- 
mentum. 24-33, N. arbovum ; 34-38, N. diabolus ; 39-45, N. elegantulus ; 46-50, 
N. fervantt ; 51-57, N. fullerz. 


Fics. 58-99 : 
mentum. 
pilosity) ; 


ETHIOPIAN NASUTITERMITINAE 


99. 

Nasutitermes, soldier head capsule, side and plan views, nose tip and post- 
58-61, N. hirticeps ; 62-68, N. infuscatus ; 69-77, N. kempae (72, vertex 
78-87, N. latifrons ; 88-94, N. lujae ; 95-99, N. schoutedeni. 


w 


24 W. A. SANDS 


name arborum Smeath. should be used to replace Jatifrons, for the following reasons : 
(i) The specimens identified by Sjostedt and Silvestri were not from the type locality 
of N. arborum. N. latifrons has only recently been collected from Sierra Leone for 
the first time. (11) NV. Jatifrons is not, in general, an arboreal-nesting species, and is 
unlikely to be that described by Smeathman. It most commonly occurs in dead 
logs, at the foot of tree trunks, and in the mounds of other species, such as Cubitermes 
spp. and Thoracotermes, on the forest floor. (i) The name WN. Jatifrons is well 
established and widely used for a species which is probably the most widely distributed 
in the Ethiopian Region. 

It would seem that in the interests of stability of nomenclature, it is essential to 
fix the identity of N. arvborum by the designation of a Neotype, particularly since 
another arboreal species, N. lujae closely resembles it in both imago and soldier, and 
is easily confused with it. It is distinguished in the imago by the more prominent 
postclypeus ; in the soldier by the lack of bilaterally placed long setae on the 
vertex, and the less hairy nose tip. In N. fulleri the single imago morphotype has a 
proportionately shorter hind tibia, and the soldier has a distinctly hairy nose tip, 
vertex setae, I2 segmented antennae, and an angularly concave profile. N. 
schoutedeni is separable in both castes by the same characters as N. lujae. The 
East African species N. infuscatus is closely similar and difficult to distinguish. 
The imago often has shallow diverging grooves on the frons anterior to the fontanelle, 
but these are not always well developed. They are absent in N. avborum, in which 
the 9 cerci are shorter than their width at the base, whereas in N. infuscatus they 
are as long. The soldier of N. infuscatus has usually a more slender nose, with a 
slightly more hairy tip, and sometimes has setae on the vertex. 

The original species-name arvborum of Smeathman was altered in Snyder (1949) 
to arboreus, presumably with the object of making it an adjective agreeing with the 
generic name. The name arborum is however the genitive plural of arbor, giving the 
meaning “‘ of the trees’’, and as such is a valid species name. I have therefore 
reverted to the use of the original form of the name. 


Neotype. SIERRA LEONE: Nijala (Lat. 8° 6’ N., Long. 12° 5’ W.), 27.1.1955 
(W. V. Harris Coll. No. gor). Neotype soldier and other material from same colony 
in British Museum (Natural History). Morphotype. NIGERIA: Western Region, 
Benin Province, Sobo Plain, Obanokoro, 9.11.1957 (W. Wilkinson Coll. No. WW571). 
Morphotype imagos in British Museum (Natural History). 

Type material. DAnomEy : Cotonou, 7.11.1913 (F. Silvestri), syntypes, female 
imago and soldier, type colony, N. infuscatus var. perfusca (Silvestri), A.M.N.H. 
(other syntypes in Silvestri Coll., Lab. Zool. Sc. Agr., Portici). 


GABON : No detailed locality (Aubry-Lecomte), syntype male imago and soldier, 
type colony, N. maculiventris (Sjéstedt), A.M.N.H., (other syntypes in Naturhist. 
Riksmus., Stockholm). 


Other Material. SIERRA LEONE : Kenema, 12-13.1.1958 (W. Wilkinson). 


GUINEA : Kakoulima (between Conakry and Kindia), 1912 (F. Szlvestrt), 
A.M.N.H. ; Nimba Mts., 1957 (M. Lamotte), Inst. Franc. d’Afr. Noire, Dakar. 


ETHIOPIAN NASUTITERMITINAE 25 


LIBERIA : No detailed locality, 1908 (Sherer), A.M.N.H. 

GHANA: Yeji, 9.1.1927, Accra, 13.v.1936 (A. W. J. Pomeroy) ; Bobiri Forest, 
23m. from Kumasi on Accra Road, 21.ii.1959, Kumasi, 22.11.1959, Ejura, 23.11.1959, 
6om. N. of Ejura on Tamale Road, 24.11.1959, Awura Forest, 4m. from Ejura on 
Kintampo Road, 28.iii.1959, 9m. from Nkoranza on Techiman Road, and 12m. N. 
of Techiman on Wenchi Road, 29.iii.1959 (W. A. Sands). 


NIGERIA: Western Region ; Sobo Plain, Obanokoro, 8.1.1957, Lagos, Light- 
house Beach, 10.iv.1957 (W. Wilkinson) : Olokemeji, 30m. W. of Ibadan, 8.xii.1957, 
Agodi, Ibadan, 10.xii.1957, between Shagamu and Ijebu-Ode, 14.xii.1957 (W. A. 
Sands). Eastern Region ; Port Harcourt, 14—16.iii., I-2.iv.1957, 29m. from Port 
Harcourt on Owerri Road, 19.iv.1957 (W. Wilkinson). Northern Region ; 6m. N. 
of Minna, 20.x11.1956, 18m. from Bida on Badeggi Road, 24.xii.1956, Idu Forest, 
2om. from Abuja on Keffi Road, and Gwagwa Forest, near Abuja, 28.x11.1956, 
Gwachipe Forest, 12m. from Abuja on Minna Road, 29.xii.1956, Dogon Forest, 
65m. from Jos on Kafanchan Road, 8.1i.1957, 6m. N. of Jebba on Mokwa Road, 
16.xii.1957, 42m. from Gboko on Oturkpo Road, 26.11.1958, 22m. from Lokoja on 
Kabba Road, 10.ili.1958, 23m. from Kabba on Ondo Road, 11.iii.1958 (W. A. Sands). 

CAMEROUN : Mbalmayo, 7.x.1962 (G. Becker), own collection. 

REPUBLIC OF CONGO: Brazzaville, 7.vi.1948 (A. E. Emerson), A.M.N.H. 


Conco: Basoko, 1909 (H. Lang and J. P. Chapin) ; 5km. from X-roads near 
Bilni (near Mambasa, Ituri Forest), 10.v.1948, Stanleyville, 25.v.1948, Yangambi, 
30.v.1948 (A. E. Emerson) (with Lang-Chapin coll., at A.M.N.H.) ; Garamba Nat. 
Park, 1950 (H. de Saeger and G. Demoulin), Inst. des Parcs Nat. du Congo, 
Brussels. 


UGANDA : Bwamba, 19.11.1949, Kigezi, 1947 (W. V. Harris); 16m. from 
Kampala on Entebbe Road, 1.ii1.1955 (W. Wilkinson). 

A total of 52 nest series were examined, and all material is in the British Museum 
(Natural History) unless otherwise stated. 


Though common from Sierra Leone to Uganda, this species appears to have a 
more limited range south-eastwards than some of its congeners. It is found mainly 
in rain forest or forest outliers in forest-savannah mosaic areas (Keay ef al., 1959, 
vegetation types 7 and 8), though occasionally recorded from true savannah-wood- 
land vegetation when growing under moist conditions near forest. The records of 
this species from Senegal and the Cape (referring to the Cape of Good Hope) by 
Hagen (1858), repeated in Snyder (1949), are erroneous. The Cape of Good Hope 
specimen of the soldier is Trinervitermes trinervoides (Sjéstedt), and the Senegal 
alate specimen does not belong to the subfamily Nasutitermitinae at all, but is 
probably Termes or Promirotermes. 

The nests of this species are of dark brown, almost black carton, usually situated 
above ground level in trees or on woody lianes. The carton foraging runways 
extend in all directions for considerable distances, and it is common to find only the 
foraging workers and soldiers and to be unable to trace the nest. 


26 W. A. SANDS 


Nasutitermes camerunensis (Sj6stedt) comb. n. 
(Text-figs. 4, 5, and 201 ; Map 3.) 


Eutermes camerunensis Sj6stedt, 1899 : 37. Type locality : CAMEROUN. 
Eutermes camerunensis Sjostedt : Snyder, 1949 : 349 [not classified]. 
Microcevotermes (?) camerunensis (SjOstedt) Weidner, 1955 : 249. 


Imago. ead capsule chestnut brown, postclypeus, labrum, and antennae bright yellow- 
brown, almost orange. Pronotum, meso- and metanota, and abdominal tergites brown, 
pronotum with paler irregular “‘ T’’-shaped mark in middle of front half. Legs yellow. 
Abdominal sternites pale yellow in middle, clouded with pale brown round stigmata. Wings, 
membrane very pale brown, almost hyaline, slightly iridescent, veins pale brown outlined with 
sepia brown. 

Posterior margin of head behind eyes distinctly less than semi-circular ; fontanelle small, 
pale, but widely bifurcate anteriorly, lateral arms of “ fork’’ reaching half-way to ocelli ; 
eyes prominent, medium-sized, angularly oval, W/E index 2-9 ; ocelli medium-sized, oval, 
separated from eyes by half own least diameter or slightly less ; postclypeus moderately inflated, 
width 2:5 times length, anterior margin nearly straight, posterior margin rounded ; antennae 
15 segmented, II and IV subequal, slightly longer than III and V. 

Cerci (both known specimens are 3) about as long as width across base, mamilliform. 

Measurements (2 specimens, holotype and one other) in millimetres. 


Range 

Head width across eyes é : ; I+34—-1°35 
Greatest diameter of eye . ; : 0-46 

@cellusy >: : : 3 : : 0-13-0°14 X O-16—-0-19 
Ocellus toeye . ; S : : 0:05-0:06 
Width of pronotum . : : : I-06—-I-10 
Length of pronotum . : : : 0-64—0-69 
Length of hind tibia . 3 : j 1-61 


This species is only known from the holotype imago and one other specimen. 
The soldier and worker castes are unknown. It was first recognized as belonging 
to Nasutitermes by Emerson, and my attention was drawn to it by his unpublished 
notes. Since then a further specimen has been found in the collection of the Termite 
Research Unit, confirming its existence as a separate species. It may be noted that 
only two species from the mainland of Africa are now known only from the soldier 
caste, namely N. diabolus and N. ferranti. Similarly only two are known from the 
imago alone, N. camerunensis and N. chrysopleura. In view of the widespread 
records of all the mainland species now known, it seems probable that these soldiers 
and imagos belong together in two species, although this cannot be settled until all 
castes are collected together. It is however possible to predict that if this is correct 
the most likely combinations would seem to be the soldier of N. dzabolus with the 
imago N. chrysopleura and the soldier of N. ferranti with the imago N. camerunensis. 

N. camerunensis differs from N. kempae and N. elegantulus in having relatively 
smaller eyes ; the hind tibia is proportionately longer than in N. fuller1, and the 
widely bifurcate fontanelle distinguishes it from N. arborum and N. infuscatus. In 
the remaining species the postclypeus is much shorter in proportion to its width. 

Material. CAMEROUN: 111.1892 (H. Brauns), unique holotype § imago, Mus. 


Hamburg. 
NIGERIA: Eastern Region ; Port Harcourt, 15.11.1957 (W. Wilkinson). 


ETHIOPIAN NASUTITERMITINAE 27 


Nasutitermes chrysopleura (Sjostedt) 
(Text-figs. 6, 7, and 200; Map 3) 


Eutermes chrysopleura Sj6stedt, 1897 : 126. Type locality : CAMEROUN, Victoria. 
Termes (Eutermes) chrysopleura (SjOstedt) Desneux, 1904a : 41. 

Eutermes chrysopleura Sjostedt ; Sjéstedt, 1926 : 324. 

Nasutitermes chrysopleura (Sj6stedt) Snyder, 1949 : 271. 


Imago. Head capsule brown, postclypeus, antennae yellow-brown. Pronotum pale brown, 
legs yellow-brown. Abdominal tergites brown, sternites pale brown round stigmata, yellow- 
brown in middle. Wings missing in cotype. Colours may be faded in old specimen. 

Posterior margin of head behind eyes nearly semi-circular ; fontanelle distinct, pale, tapering 
posteriorly, and concave rather than bifurcate at anterior end ; eyes medium sized, only 
moderately prominent in proportion to own diameter, nearly circular, W/E index 3-0 ; ocelli 
medium sized, separated from eyes by half own greatest diameter, oval ; postclypeus weakly 
inflated, width 3-8 times length, anterior margin nearly straight, posterior margin convex, 
widely arched, distinctly sinuate towards outer corners ; antennae damaged, II and V sub- 
equal, slightly shorter than IV, III longer than IV. Meso- and metanota with very wide- 
angled posterior margins. Cerci very short, only about two-thirds width at base, nearly 
conical. 

Measurements (single paratype specimen) in millimetres. 


Head width across eyes : : : ’ ; 1°53 
Greatest diameter of eye. 2 ‘ : ; 0-50 
Ocellus : ’ : ; * ; . 5 O14 x O19 
Ocellus toeye . : , ; ; , ; 0-09 
Width of pronotum . : 3 : : : 1°38 
Length of pronotum . : : : : : 0°83 
Length of hind tibia . ; 3 , : : 1°95 


This species is known only from the type material It closely resembles 
N. latifrons, which is common in the vicinity of the type locality, and it may 
ultimately prove to be the same species. There are however certain differences, 
even from the N. latifrons from the same area, and it may be that N. chrysopleura is 
the at present unknown imago of N. diabolus. The length of the head capsule to the 
front of the postclypeus is slightly but distinctly more than its width behind the 
eyes. In N. latifrons these dimensions are equal. The posterior margins of the 
meso- and metanota are more widely angled in N. chrysopleura. Though this is 
not usually a specific character in Nasutitermes, they appear to be consistently more 
narrowly angled in N. Jlatifrons. The posterior margin of the postclypeus in N. 
latifrons tends to be straighter, slightly angled in the middle instead of curved, and 
not sinuate towards the outer corners. The 9 cerci are also slightly longer and more 
mamilliform in N. latifrons. 

N. chrysopleura is therefore retained as a separate species until further specimens 
are found to clarify its identity. It is unlikely to be confused with any other species, 
being larger than the rest of those having a short wide postclypeus. 


Material. CAMEROUN: Victoria, ix.1873 (Bucholz), single paratype 2 imago, 
A.M.N.H. (Holotype, in Mus. Greifswald). 


28 Wi Az SAUNEDS 


NASUTITERMES CAMERUNENSIS 


NASUTITERMES CHRYSOPLEURA 


NASUTITERMES FERRANTI 


NASUTITERMES HIRTICEPS 


GRALLATOTERMES AFRICANUS 
MYCTEROTERMES MERINGOCEPHALUS 
TRINERVITERMES ARABIAE 


TRINERVITERMES SAUDIENSIS 


PEPI BOO 


BAUCALIOTERMES HAINESI 


Map 3. ‘“ Procornitermes branch’’; genera and species not mapped individually. 


ETHIOPIAN NASUTITERMITINAE 29 


Nasutitermes diabolus (Sjéstedt) 
(Text-figs. 34-38; Map 5) 


Eutermes diabolus Sjéstedt, 1907 : 250. Type locality : Conco, Mukimbungu. 

Eutermes (Eutermes) diabolus Sjéstedt ; Holmgren, 1912 : 62. 

Eutermes (Eutermes) torquatus Sjéstedt, 1924c : 494. Type locality : Conco, Stanleyville. 
syn. n. 

Eutermes diabolus Sj6stedt : Sjéstedt, 1926 : 318. 

Eutermes torquatus Sjostedt ; Sjéstedt, 1926 : 321. 

Nasutitermes (Nasutitermes) diabolus (Sj6stedt) Emerson, 1928 : 477. 

Nasutitermes diabolus (Sj6stedt) ; Snyder, 1949 : 274. 

Nasutitermes torquatus (SjOstedt) Snyder, 1949 : 299. 


Imago. Unknown, but see discussion on N. camerunensis and chrysopleura. 

Soldier. Head capsule ferruginous to chestnut brown, nose ferruginous, uniform with head 
capsule, to dark pitchy brown, almost black. Some specimens and colonies have the distal half 
of the nose ferruginous to chestnut brown, shading paler near the base, and then a distinct 
brown to dark chestnut brown ring around the base and extending on to the head capsule. All 
intermediates between this and the uniformly coloured form exist, sometimes in one colony. 
Pronotum, antennae, legs, and abdominal tergites yellow-brown to brown, abdominal sternites 
yellow to yellow-brown. 

Head capsule in plan view near circular or slightly oval, tapering towards front, nose weakly 
conical. In profile, evenly concave, sometimes very slightly sinuate at base of nose, nose 
slightly shorter to slightly longer than rest of head capsule, measuring from hind margin of 
antennal socket, angle of nose cone 10-15 degrees ; postmentum evenly moderately convex ; 
antennae 13 segmented, II, III, and V subequal, or III slightly longer, IV distinctly shorter than 
these. Vertex with a pair of bilaterally placed long setae in some but not all specimens, and a 
few very fine microscopic setae ; nose with four long setae at tip, and sometimes a very few 
minute fine setae immediately behind them ; postmentum with a few long setae on anterior 
margin only. 

Abdominal tergites with scattered very short setae, and longer setae on posterior margins of 
all except first or first and second, those on posterior tergites longer than rest ; sternites with 
numerous shorter setae and long setae on posterior margins. 

Measurements (25 specimens from 9 localities) in millimetres. 


Range Mean 
Head length to tip of nose. 1:50-1:80 : : - : 1-60 
Head width : : : 0-80-1-00 : ; : : 0-87 
Depth of head capsule . ; 0-58-—0-70 : : ; - 0-61 
Width of pronotum : ; 0-41-0°50 : J ; < 0°45 
Length of pronotum . : 0°15—0°20 : : : : o'18 
Length of hind tibia. 0:98-1:18 1:05 


Variation. The shape and size of the soldier caste is less Vv ariable than i in some other species, 
but the colour pattern is highly variable. The form with the banded nose was described 
separately by Sjéstedt (1924) as N. torquatus. 


References to N. torquatus by subsequent authors (Emerson, 1928, and Sands, 
1957) are not correct, and are listed in the appropriate synonymies. The darker 
coloured and longer-nosed specimens of N. /ujae have sometimes been confused with 
this species, but they are distinguishable by the more hairy nose tip and the less 
evenly concave profile. N. lwae also always has a pair of bilaterally placed setae 
on the vertex, and no smaller hairs have been detected in this position. JN. latifrons 
is sometimes similar in general shape, but is easily distinguished by the hairy 


30 W. A. SANDS 


postmentum, and the usually paler colouration. N. schoutedent has a sharply 
tapered and distinctly hairy nose tip. N. kempae is East African in distribution, 
and also has a hairy nose tip. 


Type Material. Conco: Mukimbungu, 1906 (K. E. Laman), syntype soldiers 
from type colony, No. 1o11, A.M.N.H. (other syntypes in Naturhist. Riksmus., 
Stockholm) ; Stanleyville, 1913 (H. Kohl), syntype soldier from type colony, 
No. 1037, of N. torquatus (Sjéstedt), A.M.N.H. (other syntypes in Naturhist. 
Riksmus., Stockholm). 


Other Material. NiGerIA: Western Region ; Sobo Plain, Obanokoro, 7.1iii. 
1957 (W. Wilkinson), B.M.(N.H.). Eastern Region ; 8m. from Port Harcourt on 
Owerri Road, 10.xii.1957, and Port Harcourt, 18.xii.1957 (W. Wilkinson), B.M.(N.H.) 

REPUBLIC OF CONGO: Brazzaville, 7.vi.1948 (A. E. Emerson), A.M.N.H. 

Conco: 20km. E. of Ndjili and 18km. S. of Leopoldville, 10.vi.1948, Yangambi, 
30.v.1948 and Epulu, 23.v.1948 (A. E. Emerson) ; Stanleyville, 21.vi.19g13 (H. 
Kohl), 18.11.1948 (N. A. Weber), all in A.M.N.H. 

A total of 11 nest series were examined. 


This species has not been collected further westwards than Western Nigeria, and 
it seems likely that this is near the limit of its range. It is thus confined to the main 
forest block of West and Central Africa. It was recorded by Emerson (1928) as 
being a tree-nesting species, but like the rest of those with this habit, it is most 
frequently collected from runways on trees, and the nest is not easily located. 


Nasutitermes elegantulus (Sj6stedt) 
(Text-figs. 8-10, 39-45, 160-162, 202, 203; Map 6) 


Eutermes elegantulus Sjéstedt, I911b : 13. Type locality : CAMEROUN, Victoria. 

Eutermes macyophthalmus Silvestri, 1912 : 240. Type locality : ISLAND OF PRINCIPE, Roca 
Infante. 

Eutermes (Subulitermes?) elegantulus SjOstedt ; Holmgren, 1912 : 64. 

Eutermes (Eutermes) macrophthalmus Silvestri ; Holmgren, 1912 : 62. 

Eutermes (Eutermes) minusculus Sjostedt, 1924 : 40. Type locality : Conco, Lukula. syn. n. 

Eutermes elegantulus SjOstedt ; Sjéstedt, 1926 : 309. 

Eutermes macrophthalmus Silvestri [= E. elegantulus Sjostedt] ; Sjostedt, 1926 : 309. 

Eutermes minusculus SjOstedt ; Sjdstedt, 1926 : 308. 

Nasutitermes (Nasutiteymes) kohli Emerson, 1928 : 484. Type locality : Conco, Kandolo, 
near Stanleyville. syn. n. 

Nasutitermes (?Nasutitermes) minusculus (Sj6stedt) Emerson, 1928 : 487. 

Nasutitermes elegantulus (SjOstedt) Snyder, 1949 : 275. 

Nasutitermes kohli Emerson ; Snyder, 1949 : 282. 

Nasutitermes minusculus (Sj6stedt) ; Snyder, 1949 : 288. 


Imago. Head capsule yellow-brown, with three indistinct streaks on vertex, one in mid-line 
and two diverging laterally, slightly paler, yellow to pale yellow-brown. Postclypeus, labrum, 
antennae, pronotum, meso- and metanota, and legs yellow to pale yellow-brown. Abdominal 
tergites yellow-brown to pale brown, slightly darker at sides than in middle ; sternites yellow 
to pale yellow-brown, darker round stigmata. Wings translucent, pale yellow-brown, subcosta 
yellow-brown, radius sector yellow to yellow-brown outlined with brown, other venation pale 
brown. 


ETHIOPIAN NASUTITERMITINAE 31 


Posterior margin of head behind eyes less than semi-circular, distinctly sinuate immediately 
behind eyes ; fontanelle distinct, pale ‘‘ Y ’’-shaped, sometimes slightly depressed posteriorly ; 
eyes large to very large relative to head width, W/E index 2-0 to 2:5, but only moderately 
prominent in proportion to own diameter, slightly angularly oval ; ocelli small, from almost 
touching eyes, to separated from them by less than half own least diameter ; postclypeus 
moderately inflated, width 2-4 to 3-1 times length, anterior margin nearly straight to slightly 
concave, posterior margin convex, rounded ; antennae 15 segmented, II, III, and IV subequal 
in length, but III slightly thinner. 

Cerci of 2 as long as width across base, conical. 

Measurements (14 specimens from 3 localities) in millimetres. 


Range Mean 
Head width across eyes 1°04-1°18 : ‘ rr2 
Greatest diameter of eye O-41-0'51 P ; 0°47 
Ocellus : P ; 0-09-O'II X O-II-O'1I4 . : 0-10 X O-12 
Ocellus toeye  . : 0-OI-0:04 : : 0-02 
Width of pronotum . 0-85-1-00 e ; 0-94 
Length of pronotum . 0°55-0°68 ; 0-61 
Length of hind tibia. 1-25-1°38 ‘ i I +33 
Length of fore wing. 9-40-11-80 3 : 10°62 


Soldier. Head capsule orange-yellow to orange, nose ferruginous orange to ferruginous. 
Pronotum and antennae as head capsule. Abdominal tergites and legs yellow to orange-yellow, 
abdominal sternites pale yellow. 

Head capsule in plan view oval, tapering towards front, often slightly ‘‘ shouldered ’’ behind 
antennae, nose very weakly conical, and slender. In profile, usually with distinct convex 
change of contour at base of nose, but sometimes nearly straight or somewhat sinuate, nose 
approximately equal in length or slightly longer than rest of head capsule measured from hind 
margin of antennal socket ; angle of nose cone 7-15 degrees ; postmentum prominent and 
distinctly convex ; antennae 11-13 segmented, 4 out of 5 specimens with 12, proportions 
of segments in the latter, If and V subequal, IV shorter than these or equal, III shorter than 
IV ; in 13 segmented forms, II and VI subequal longer than V, III and IV shorter than V, 
sometimes subequal, or III longer than IV. Setae on head consist of a bilaterally placed pair on 
vertex, two or four across base of nose, and rather hairy nose tip, sometimes with four setae 
more prominent than those behind them ; microscopic short bristle-like setae scattered on 
vertex, visible in profile ; postmentum setae confined to anterior margin. 

Abdominal tergites with very short scattered setae, and longer setae on posterior margins, 
more prominent towards the rear ; sternites with more numerous short setae and longer setae 
on posterior margins. 

Measurements (59 specimens from 17 localities) in millimetres. 


Range Mean 
Head length to tip of nose. 1-18-1-48 : : : . I-31 
Head width . : ; : 0°59-0°81 : : : : 0-70 
Depth of head capsule . - 0°39-0°55 . : ; : 0-46 
Width of pronotum . , 0°33-0°43 ‘ : , : 0°36 
Length of pronotum : : O-II-0-16 : : : ; 0-13 
Length of hind tibia : : 0:70-0:90 O77 


Variation. In the imago the main variation is in the wiative size of the eyes. Soldier 
variation is largely in the head profile, and in the development of the small scattered hairs on 
the vertex, which are sometimes not much in evidence. They are very prone to being rubbed 
off, and old or battered specimens are likely to have lost most of them. 


Only two species show any great resemblance to N. elegantulus. The East 
African species N. kempae is similar but larger, and these two would appear to be 
more closely related to each other and to certain species from other zoogeographical 


32 W. A. SANDS 


regions than to any other Ethiopian species. The only species which appears to be 
somewhat intermediate between these and the other Ethiopian forms is N. fulleri, 
separated from N. elegantulus in the soldier by the difference in profile and the lack 
of small setae on the vertex, and in the alate, by the relatively smaller eyes, with the 
ocelli more distant from them. 


Type Material. CAMEROUN: Victoria, 19.1x.1893 (Bucholz), syntype soldier, 
type colony, N. elegantulus (Sjostedt), A.M.N.H. 

ISLAND OF PRINCIPE : Roca Infante (L. Fea), syntype 2 imago and soldier, type 
colony, N. macrophthalmus (Silvestri), A.M.N.H. (other syntypes in Silvestri coll., 
Lab. Zool., Se: Agr., Portici). 

Conco: Lukula (H. Schouteden), syntype soldier, type colony, N. minusculus 
(Sjostedt), A.M.N.H. (other syntypes in Congo Mus., Brussels) ; Kandolo, near 
Stanleyville, 13.vii.1913 (H. Kohl), paratype soldiers, type colony, N. kohli Emerson, 
A.M.N.H. and B.M.(N.H.) (Holotype in A.M.N.H.). 


Other Material. GUINEA: Nimba Mountains, 1957, 3 vials (M. Lamotte), Inst. 
Frang. d’Afr. Noire, Dakar. 

GHANA : Bobiri Forest, 23m. from Kumasi on Accra Road, 21.11.1959, and gm. 
from Nkoranza on Techiman Road, 29.111.1959 (W. A. Sands). 

NIGERIA: Western Region; Sobo Plain, Obanokoro, 9.i, 7.111.1957 (W. 
Wilkinson) ; 30m. from Ijebu-Ode on Benin Road, 12.xii.1957 (W. A. Sands). 
Eastern Region ; 25m. from Enugu on Awgu Road, 2.11.1957, Nkpoku, 12m. from 
Port Harcourt on Owerri Road, 17.vili.1957, 8m. from Port Harcourt on Owerri 
Road, 10.xii.1957, and Port Harcourt, 19.xu1.1957 (W. Wilkinson). Northern 
Region ; 18m. from Bida on Badeggi Road, 24.x1i.1956, and Idu Forest, 20m. 
from Abuja on Keffi Road, 28.xii.1956 (W. A. Sands). 

CAMEROUN : Mbalmayo, 17.x.1962 (G. Becker), own collection. 

ISLAND OF PRINCIPE : Roca Sundi, 16, 23.1x.1949 (G. R. Gradwell and D. Snow). 

Conco : Luluabourg, 28.vi.1913 (Callewaert) ; Malela, 4.vili.1915 (J. Bequaert) ; 
Epulu, 17.v.1948, 4 vials (A. E. Emerson), all in A.M.N.H. 

A total of 27 nest series were examined, and all material is in the British Museum 
(Natural History) unless otherwise stated. 

This species is widely distributed throughout the West African and Congo forests, 
but is never common. It has been recorded as building soft carton nests, but is 
most frequently found in the nests of other genera, in rotting logs, or in narrow 
runways on trees. The workings have a reticulate shagreen patterned lining of 
black carton. 


Nasutitermes ferranti (Wasmann) 
(Text-figs. 46-50; Map 3) 


Eutermes feryvanti Wasmann, 1911 : 167. Type locality : Conco, Kasai, Sankuru. 
Eutermes (Eutermes) ferranti Wasmann ; Holmgren, 1912 : 62. 

Eutermes ferrantti Wasmann ; Sjéstedt, 1926 : 310. 

Nasutitermes ferranti (Wasmann) Snyder, 1949 : 276. 


ETHIOPIAN NASUTITERMITINAE 33 


Map 4, NASUTITERMES ARBORUM 


Map 5, NASUTITERMES DIABOLUS 


Map 6, NASUTITERMES ELEGANTULUS 


Maps 4-6. (4), Nasutitermes arborum, Vegetation types: 7, 8, and 16; (5), Nasuti- 
termes diabolus. Vegetation type: 7; (6), Nasutitermes elegantulus. Vegetation 
types: 7 and 8. 


34 WA. SANDS 


Imago. Unknown, but see discussion on N. camerunensis and N. chrysopleura. 

Soldier. Head capsule ferruginous orange to chestnut brown, nose dark chestnut brown to 
brownish black, with reddish tip. Pronotum, antennae, and abdominal tergites yellow-brown. 
Legs yellow, abdominal sternites pale yellow. 

Head capsule in plan view nearly circular, nose broadly conical. In profile, near straight or 
slightly sinuate to well behind base of nose, then raised and rounded to back of head, small hump 
at base of nose common. Nose distinctly shorter than rest of head capsule, measuring to hind 
margin of antennal socket, angle of nose cone 18-30 degrees ; postmentum weakly convex ; 
antennae 13 segmented, II and V subequal and longer than IV, but shorter than III. Head 
setae consist of bilaterally placed pair on vertex, two or four across base of nose, four at nose 
tip, and a few on anterior margin of postmentum ; there are also scattered microscopic setae 
on vertex, only visible at certain angles of illumination. 

Abdominal tergites with very sparsely scattered minute setae, and longer setae on posterior 
margins, four to each tergite arranged in longitudinal rows down abdomen ; sternites with 
sparse short setae, and longer setae on posterior margins, less regularly arranged than on 


tergites. 
Measurements (17 specimens from 4 localities) in millimetres. 

Range Mean 
Head length to tip of nose. I-39-1°58 : : : : I-49 
Head width . ; : : 0:79-1:00 : : ; ‘ 0:92 
Depth of head capsule . : 0°54-0°65 : : : : 0-61 
Width of pronotum : : 0-4I-0'50 : . : . 0°44 
Length of pronotum ; ; 0°15—0:20 ; , : : Oy 
Length of hind tibia : ; 0:95-1:08 , : : : I-02 


This species had not been recognized since it was first described, but new material 
has now been discovered. It is distinct from all other African species, with its 
short, thick nose tipped with only four setae, and the two or four setae across the 
base of the nose. 


Material. Conco: Kasai, Sankuru (EZ. Lwja), unique holotype soldier, Nasuti- 
termes ferrantt (Wasmann), Mus. Maastricht. 

NIGERIA : Eastern Region ; Bonny, i.xii.1957 (W. Wilkinson), B.M. (N.H.). 

REPUBLIC OF Conco: Brazzaville, 1.iv.1948, and Kipome, 13km. W. of Brazza- 
ville, 8.vi.1948 (A. E. Emerson), A.M.N.H. 


Though apparently widely distributed, this species must be very rare to be so 
poorly represented in collections. It has been recorded from soft runs on trees, 
and from a mound of Odontotermes scrutor (Sjostedt) shared with Amuitermes evunctfer 
Silvestri. 


Nasutitermes fulleri Emerson 
(Text-figs. 11-13, 51-57, 204; Map 7) 


Nasutitermes (Nasutitermes) fulleri Emerson, 1928 : 483. Type locality : Conco, Stanleyville. 
Nasutitermes fulleyri Emerson ; Snyder, 1949 : 276. 


Imago. Previously undescribed. Head capsule sepia brown, postclypeus paler, yellow- 
brown, labrum yellow. Pronotum, and abdominal tergites sepia brown, meso- and metanota, 
pleural and ventral thoracic sclerites and femora brown. Antennae, tibiae and tarsi pale 
yellow. Abdominal sternites brown laterally, shading to pale yellow-brown in middle. Wings 
translucent pale brown, subcosta and radius sector yellow-brown, outlined sepia brown, rest 
of venation brown. 


ETHIOPIAN NASUTITERMITINAE 35 


Posterior margin of head less than semi-circular behind eyes, distinctly sinuate immediately 
behind eyes ; fontanelle slightly paler than rest of head, Y-shaped ; eyes medium sized relative 
to head width, W/E index 3-0, not very prominent in proportion to own diameter, slightly 
angular oval ; ocelli rather small, separated from eyes by slightly less than half own least 
diameter ; postclypeus moderately inflated, width 3-0 times length, anterior margin straight, 
posterior margin convex, rounded ; antennae 15 segmented, II slightly longer than III, III 
than IV, and IV than V. 

Cerci of 2 as long as width across base, mamilliform. 

Measurements (unique morphotype Q) in millimetres. 


Head width across eyes. : é approx. 1-26 (damaged) 
Greatest diameter of eye . ; : 0-43 

Ocellus. . : . . : : O-II X O'15 
Ocellus to eye : : : : 0-05 

Width of pronotum . : . : 1-00 

Length of pronotum ; : : 0-60 

Length of hind tibia ‘ F . 1-38 


Length of fore wing . 10°40 


Morphotype. UGANDA: Bwamba, 19.11.1949 (W. V. Harris coll. no. 550). 
Morphotype 9 imago in British Museum (Natural History). 


Soldier. The new material now available necessitates some additions to the original descrip- 
tion, as follows ; nose slightly shorter than rest of head capsule, measured from hind margin 
of antennal socket, weakly conical, angle of nose cone 7—15 degrees, nose tip with four main 
setae, and sparsely hairy behind them. Postmentum setae confined to anterior margin. 

Measurements (36 specimens from 13 localities) in millimetres. 


Range Mean 
Head length to tip of nose. I*23-1°53 : : : 1°39 
Head width . : : : 0-65-0-90 ; ; : : 0:78 
Depth of head capsule . : 0-43-0-60 ; ; : F 0°53 
Width of pronotum ; : 0°35-0°45 : ‘ ; : 0-40 
Length of pronotum - : 0-13-0'18 : : ; : Or17 
Length of hind tibia 3 é 0°75-0°98 : F : 2 0°85 


The distinctions between this species and N. elegantulus have been given in the 
discussion on the latter. N. fullert also shows certain similarities to N. infuscatus, 
but is separated in the soldier by the consistently 12 segmented antennae and in the 
imago by the relatively shorter hind tibia. The unique morphotype female of 
N. fulleri is damaged, and more material is needed to establish fully its relationships, 
and distinctions from other species. 


Material. Conco: Stanleyville (H. Kohl), paratype soldiers, type colony, 
N. fulleri Emerson, A.M.N.H. and B.M.(N.H.) (Holotype in A.M.N.H.) ; Epulu, 
Ir and 23.v.1948, 7 vials (A. E. Emerson), A.M.(N.H.). 

SIERRA LEONE: Freetown, 8.i.1958 (W. Wilkinson), B.M.N.H. 

GuINEA : Nimba Mountains, 1957, 2 vials (M. Lamotte), Inst. Franc. d’Afr. 
Noire, Dakar. 

NIGERIA : Western Region ; Sobo Plain, Obanokoro, 6~7.iii.1957, 2 vials 
(W. Wilkinson), B.M.(N.H.). Northern Region ; Idu Forest, 20m. from Abuja 
on Keffi Road, 28.xii.1956, and Dogon Forest, 65m. from Jos on Kafanchan Road, 
8.11.1957 (W. A. Sands), B.M.(N.H.). 


36 W. A. SANDS 


CAMEROUN: Im. W. of Calabar Junction on Mamfe-Ikom Road, 27.v.1957 
(W. Wilkinson), B.M.(N.H.). 

CENTRAL AFRICAN REPUBLIC: Bengessa (Bangassou), Bas Mbomu, 12.iii.1948 
(iN: An Weber), ALINE. - 

UGANDA : Ruwenzori Mountains, tom. E.N.E. of Bundibugyo, 26.viii.1952, 
2 vials (G. O. Evans), B.M.(N.H.). 

A total of Io nest series were examined. 


This species occurs from Sierra Leone to Uganda. Its southward distribution 
appears to be more limited, but this may be a reflection of its scarcity and due to 
failure of collection rather than absence of the species. It has generally been found 
in woody debris on the forest floor or in small runways on trees. An arboreal nest 
has been recorded, but this may not have been of its own construction. 


Nasutitermes hirticeps sp. n. 
(Text-figs. 58-61; Map 3) 


Imago. Unknown. 

Soldier. Head capsule orange, nose ferruginous orange. Pronotum, antennae and abdominal 
tergites orange-yellow, abdominal sternites and legs yellow. 

Head capsule in plan view almost circular, nose conical. In profile sinuate with distinct 
hump at base of nose, then rather abruptly raised and rounded to back of head ; nose distinctly 
shorter than rest of head capsule, measuring from hind margin of antennal socket, angle of nose 
cone 11-15 degrees ; postmentum moderately and evenly convex ; antennae 13 segmented, 
III and V subequal, IV shorter than these, but slightly longer than IT. 

Entire body and head capsule including nose with long yellow setae, but confined on post- 
mentum to anterior margin ; those on head capsule and nose scattered irregularly, nose tip 
without any concentration of smaller setae ; anterior margin of pronotum with row of long 
conspicuous setae ; those on posterior margins of abdominal sternites longer than other 
abdominal setae. 

Measurements (10 specimens randomly selected from type colony) in millimetres. 


Range Mean 
Head length to tip of nose. I+53-1°63 : : : : 1°57 
Head width . é : : 0:95-1:03 ‘ : : ; 0:99 
Depth of head capsule . : 0:65—0-70 : : ; : 0-68 
Width of pronotum ‘ : 0:50-0°53 : 5 : : O-'51 
Length of pronotum é 4 0:20-0:23 é ¢ : : 0:21 
Length of hind tibia : . I+25-1-38 3 : 5 ; 1-30 


N. hirticeps is quite unlike any other species in the Ethiopian Region, in being 
conspicuously hairy on both head and body. Its restricted occurrence on a single 
island on a chain in which both this and other islands also share the mainland fauna, 
immediately raises the question of possible introduction from elsewhere. In view 
of the intense activity of the slave trade in that area in times past, such an intro- 
duction is not impossible, but a species of Nasutitermes would appear a somewhat 
unlikely candidate for a journey of this type ; the distributions of Coptotermes 
formosanus and Reticulitermes flavipes do however show that dispersal by human 
agencies is not confined to Kalotermitidae. 


ETHIOPIAN NASUTITERMITINAE 37 


N. hirticeps closely resembles several South American species. N. migriceps 
(Haldeman, 1858) is not readily distinguishable from it except by its darker coloura- 
tion ; in view of the age of the types it is possible there has been some fading in 
preservation, though the appearance of the specimens does not suggest this. N. 
macrocephalus (Silvestri, 1903) is larger, and the soldier postmentum bears scattered 
setae, but it is otherwise closely similar. 

I consider it desirable in the circumstances to name N. hirticeps as a new species 
in the list for the Ethiopian Zoogeographical Region, with the proviso that further 
material and information may lead to its inclusion in an already established species 
from elsewhere. 


Holotype soldier. ISLAND oF SAO THOME: Binda, 1918 (A. F. de Seabra, 
“ with Cephalotermes ’’), A.M.N.H. 


Paratypes, same data as holotype, A.M.N.H. & B.M.(N.H.). 


Nasutitermes infuscatus (Sj6stedt) 
(Text-figs. 14, 15, 62-68, 205, 206; Map 8) 


Eutermes infuscatus Sjostedt, 1902 : 40. Type locality : NyAsALAND, Zomba. 

Eutermes usambarensis Sjéstedt, 1904 : 103. Type locality : TANGANYIKA, Usambara Moun- 
tains. 

Termes (Eutermes) infuscatus (SjOstedt) Desneux, 1904a : 43. 

Termes (Eutermes) usambarensis (SjOstedt) Desneux, 1904a : 46. 

Eutermes (Eutermes) infuscatus Sj6stedt ; Holmgren, 1912 : 62. 

Eutermes infuscatus SjOstedt ; Sjéstedt, 1926 : 318. 

Eutermes usambarensis Sjostedt ; Sjéstedt, 1926 : 322. 

Nasutitermes infuscatus (Sj6stedt) Snyder, 1949 : 280. 

Nasutitermes usambarensis (SjOstedt) Snyder, 1949 : 300. 


Imago. Head capsule chestnut brown, postclypeus and antennae yellow-brown, labrum pale 
yellow-brown. Pronotum and abdominal tergites chestnut brown, meso- and metanota brown. 
Abdominal sternites pale brown laterally, narrowly paler, yellow-brown in mid-line. Legs pale 
yellow-brown. 

Posterior margin of head behind eyes nearly semi-circular, only slightly sinuate immediately 
behind eyes ; fontanelle distinct, paler than rest of head, Y-shaped ; eyes short oval, moderately 
prominent in proportion to own diameter but not large relative to head width, W/E index 
3:0-3'5 ; ocelli small, separated from eyes by slightly less than own least diameter ; post- 
clypeus moderately inflated, width 2-8—3-0 times length, anterior margin slightly concave or 
sinuate, posterior margin convex, rounded, distinctly sinuate towards outer corners ; antennae 
15 segmented, II longer than III and IV which are subequal and slightly longer than V. 

Cerci of 2 as long as width across base, slightly irregularly conical. 

Measurements (11 specimens from 4 localities) in millimetres. 


Range Mean 
Head width across eyes 1*20-1°38 : : I-31 
Greatest diameter of eye 0°37-0°44 : : O-4I 
Ocellus ; ‘ ‘ O-IO-O'II X O-I4-O'17 : O-IE X O'15 
Ocellus toeye . : 0:07—0'10 : : 0-08 
Width of pronotum I-O00-I'13 5 - 1-08 
Length of pronotum . 0+58-0-72 : é 0:68 
Length of hind tibia. 1°54-1°73 : = 1°63 


Length of fore wing. 11-60-11 ‘90 : : 77 


38 W. A. SANDS 


Soldier. Head capsule yellow to ferruginous orange, nose orange to ferruginous. Pronotum 
and antennae yellow to orange-yellow. Abdominal tergites and legs yellow, abdominal 
sternites pale yellow. 

Head capsule in plan view short oval, tapering towards front, nose weakly conical. In 
profile nearly straight, sometimes slightly concave or sinuate with weak hump at base of nose ; 
nose shorter than rest of head capsule, measuring from hind margin of antennal socket, angle of 
nose cone 12-15 degrees ; postmentum weakly convex ; antennae 12-14 segmented, usually 
13, I1 and IV subequal, shorter than V, III longer than V. Head setae confined to four at nose 
tip with shorter setae behind them, sometimes a bilaterally placed pair on vertex, and several 
on front margin of postmentum. 

Abdominal tergites with very sparse minute setae, and a few fine short setae on posterior 
margins of some or all segments ; sternites with scattered short setae, and longer setae towards 
posterior margins. 

Measurements (22 specimens from 16 localities) in millimetres. 


Range Mean 
Head length to tip of nose. I+33-1°70 : ‘ ‘ : 1°53 
Head width . : : ; 0:63-1:05 ‘ : : ‘ 0-89 
Depth of head capsule . : 0:50-0:68 : : s : 0-60 
Width of pronotum : : 0-42-0°64 : : : ; 0-48 
Length of pronotum : - 0*17—0°30 : - : : 0-21 
Length of hind tibia ; : 0:97-1°33 : : I-08 


Variation. The imago has not been found as acai as in some other species. In the 
soldier caste it is common to find larger soldiers which have slightly different form and propor- 
tions from the smaller members of the same colony, with distinct breaks in the range of variation. 
These are probably derived from a different instar of development, and represent a vestigial or 
rudimentary dimorphism ; they appear only to be found in the more populous colonies. 


This species is closely similar to N. arvborum in certain features of both imago and 
soldier, and differences have been given under that species. N. fulleri is also 
related, but is separated in the soldier by its 12 segmented antennae, and in the 
imago by its proportionately shorter hind tibia, and the ocellus being closer to the 
eye: 

Dr. A. E. Emerson has selected and labelled as lectotype one of the type series, 
but the designation has not been published hitherto :— 

Lectotype : NyASALAND: Zomba, iv.1899 (Cameron), lectotype imago and 
paralectotype soldier, N. infuscatus (Sjostedt), Naturhist. Riksmus., Stockholm. 
Other paralectotype imago and soldier, A.M.N.H. 

Type Material. TANGANYIKA: Usambara Mountains, Derema (Conradt), syn- 
type soldiers, N. usambarensis (Sj6stedt), Naturhist. Riksmus., Stockholm (others 
in Berlin Mus.). 


Other Material (additional to that recorded in Sands, 1957). Conco: 8km. N. 
of Rwindi Camp, 4.v.1948, and Rwindi Camp, 5.v.1948, 3 vials (A. E. Emerson), 
A.M.N.H. 

KenyA: Shimba Hills, 4.vi.1952 (W. A. Sands) and 13.v1.1952 (P. B. Kemp). 

TANGANYIKA : Ifakara, 30.ix.19g49 (M. Luscher), AM.N.H.; Amani, 7-9.11. 
1952, 3 vials, and Mwakijembe, 11.11.1952 (W. A. Sands). 

MaFIA IsLanD: 1937 (W. V. Harris). 

PEMBA ISLAND: Wete, ii.1952 (Packenham). 


ETHIOPIAN NASUTITERMITINAE 39 


NYASALAND : 15m. from Limbe on Cholo Road, 12.viii.1953, 9m. E. of Mlanje, 
18.viii.1953, 2 vials, 12m. E. of Chiradzulu, 21.vili.1953, 25m. from Kota-Kota on 
Kasungu Road, 17.ix.1953, and 7m. from Nkata Bay on Ekwendeni Road, 22.ix. 
1953 (W. A. Sands, W. Wilkinson). 

A total of 32 nest series were examined, and all material is in the British Museum 
(Natural History) unless otherwise stated. 


N. infuscatus (Sjéstedt) has been found in the coastal belt of East Africa, and 
inland up the Great Rift Valley to the borders of the Congo. It has not been found 
in central Kenya or Tanganyika, and there are no records from the Rift Valley 
between the northern shoreline of Lake Nyassa and Rwindi. It is thought that it 
probably occurs in patches of damp woodland on the fringes of all lakes in the Rift 
Valley, along the western border of Tanganyika. 

The nest is commonly arboreal, usually attached to the side of a tree trunk rather 
than taking the spherical form on a branch or liane seen in some other species. 


Nasutitermes kempae Harris 
(Text-figs. 16, 17, 69-77, 207, 208; Map 9) 


Nasutitermes kempae Harris, 1954 : 135. Type locality : TANGANYIKa, Handeni. 


Imago. Some additions to the original description (Sands, 1957) are necessary. W/E index 
2°4-2°6 ; postclypeus inflated, width 2-o—2-3 times length. 

Cerci of 9 slightly shorter to slightly longer than width across base. 

Further material has increased the known range of variation in size. 

Measurements (14 specimens from 2 localities) in millimetres. 


Range Mean 
Head width across eyes 1:20-1°36 : : i 27 
Greatest diameter of eye 0*48-0°55 2 - o'5I 
Ocellus - : r 0°13-0'16 X O:16-0'20_ . : 0-14 X O19 
Ocellus toeye . ; 0-01-0'03 - : 0:02 
Width of pronotum 0-94-1°18 : 3 1°05 
Length of pronotum . 0-56-0°72 ; . 0-66 
Length of hind tibia. 1-38-1-64 : - I-51 
Length of fore wing. 9°90-12:20 ; 11-18 


Soldier. The following additions to the original description (Harris, 1954) are necessary 
angle of nose cone, 10-17 degrees ; postmentum in profile weakly convex, almost meee ee 
Setae on head consist of a bilaterally placed pair on vertex, sometimes one or two small bristle- 
like setae at base of nose, and rather hairy nose tip, occasionally with four setae more prominent 
than those behind them ; microscopic small setae scattered sparsely over whole head. 

Abdominal tergites with scattered short setae and marginal setae so small and sparse as to 
appear absent except under high magnification ; sternites with sparse short setae, and longer 
setae on posterior margins. 

Measurements (29 specimens from 13 localities) in millimetres. 


Range Mean 
Head length to tip of nose. I ‘61-2:05 2 : : : 1:78 
Head width . P : : 0:88-1:29 ; . i 3 I'05 
Depth of head capsule . : 0:61-0:82 : ‘ ‘ : 0:66 
Width of pronotum , : 0°43-0°64 : : : - 0°53 
Length of pronotum : ; O+15-0°25 : é : : 0:22 
Length of hind tibia : : 0:89-1:28 1-04 


Variation. This is another species in which the soldier caste in a’ populous colonies appears to 
be derived from several instars, indicating a residual or possibly rudimentary polymorphism. 


40 WwW. A. SANDS 


| Map 7. NASUTITERMES FULLERI 


| Map 8. NASUTITERMES INFUSCATUS } Map 9, NASUTITERMES KEMPAE 
Maps 7-9. (7), Nasutitermes fulleri. Vegetation types: 7 and 8 ; (8), Nasutitermes 
infuscatus. Vegetation types: 7 and 9g, and small moist woodland and forest patches 
in Rift Valley ; (9), Nasutiteyrmes kempae. Vegetation types: 7 and 9, and small 
riverain moist woodland and forest patches inland. 


ETHIOPIAN NASUTITERMITINAE 41 


The close relationship of this species to N. elegantulus has been discussed under 
the latter, and its distinctions from other species in an earlier paper (Sands, 1957). 


Material (additional to that recorded in Harris, 1954, and Sands, 1957). 

MozaMBIQUE : Lourenzo Marques, 31.v.1953 (E. Giddy), N.C.I., Pretoria. 

REPUBLIC OF SouTH AFRICA: North Zululand, Ubombo, Lake Sibayi, and 
Sordwana Bay, 19.xi.1955 (W. G. H. Coaton), N.C.I., Pretoria, and B.M.(N.H.) ; 
Ingwavuma, 29.1.1959 (W. G. H. Coaton), N.C.I., Pretoria. 

A total of 21 nest series were examined. 


The new records from Zululand provide a great but not unexpected southward 
extension of the range of this species, and these must be very near the limit, since 
the subtropical climate gives place to the temperate zone only a little further south. 


Nasutitermes latifrons (SjOstedt) 
(Text-figs. 18, 19, 78-87, 209-212: Map 10) 


Eutermes latifrons Sjostedt, 1896 : 298. Type locality : CAMERouN, Ekundu. 

Eutermes arborum (Smeathman) ; Sjéstedt, 1900 : 204. 

Eutermes sjostedti Wwasmann, 1902 : 716. [n. n: for E. latifrons Sjéstedt| [Not valid, Desneux 
1904 : 151]. 

Termes latifrons (SjOstedt) Desneux, 1904 : 151. 

Eutermes latifrons Sjostedt ; Sjéstedt, 1905 : 19. 

Eutermes pius Sjostedt, 1911b : 16. Type locality : CAMEROUN, Victoria. syn. n. 

Eutermes (Eutermes) latifrons Sjéstedt ; Holmgren, 1912 : 62. 

Eutermes (Trinervitermes) pius SjOstedt ; Holmgren, 1912 : 65. 

Eutermes (Eutermes) incurvus Sjostedt ; 1924a : 41. Type locality : Conco, Kunungu. syn. 
n. 

Eutermes (Eutermes) ueleensis SjOstedt, 1924b : 494. Type locality : Conco, Haut Uélé, Moto. 
syn. n. 

Eutermes indoensis Sjéstedt, 1925 : 55. Type locality : RepusLic or Conco, Indo, Sibiti. 
syn. n. 

Eutermes latifrons Sj6stedt ; SjOstedt, 1926 : 311. 

Eutermes incurvus Sjostedt ; Sjéstedt, 1926 : 320. 

Eutermes ueleensis Sjostedt ; Sjéstedt, 1926 : 317. 

Trinervitermes pius (Sj6stedt) Sjéstedt, 1926 : 338. 

Eutermes expulsus Sjéstedt, 1926b : 154. Type locality : Conco, Bas Uélé, Djamba. syn. n. 

Nasutitermes (Nasutitermes) incurvus (Sj6stedt) Emerson, 1928, p. 478. 

Nasutitermes latifrons (Sjéstedt) Snyder, 1949 : 283. 

Nasutitermes incurvus (Sjostedt) ; Snyder, 1949 : 279. 

Nasutitermes ueleensis (SjOstedt) Snyder, 1949 : 300. 

Nasutitermes expulsus (Sjostedt) Snyder, 1949 : 276. 

Nasutitermes incurvus (Sjostedt) ; Sands, 1957 : 8. 


Imago. Head capsule dark chestnut brown to dark sepia brown, slightly paler in front of 
ocelli, postclypeus yellow-brown, labrum yellow, antennae brown. Pronotum dark sepia 
brown, with slightly paler patch in middle of front half ; meso- and metanota and other thoracic 
sclerites sepia brown. Legs yellow, darker at tips of femora and bases of tibiae, brown. 
Abdominal tergites dark sepia brown, lateral parts of sternites sepia brown, paler, yellow-brown 
in middle. Wing membrane translucent brown, venation sepia brown, radius sector with dark 
sepia brown line on inner side. 


42 W. A. SANDS 


Posterior margin of head behind eyes regularly rounded but less than semi-circular ; fon- 
tanelle not conspicuous, variable, from Y-shaped and slightly paler than head capsule, to almost 
obsolete, or even slightly darker than head ; eyes nearly circular, moderately prominent to 
prominent in proportion to own diameter, but not large relative to head width, W/E index 
2:9-3°4 ; ocelli medium sized, separated from eyes by two-thirds to distinctly more than own 
least diameter, oval ; postclypeus weakly inflated, width 3-3—4-2 times length, anterior margin 
slightly concave or weakly sinuate, posterior margin convex, obtusely angular in middle, near 
straight or slightly curved to outer corners ; antennae 15 segmented, II and IV subequal, 
slightly longer than III and V which are also subequal. 

Cerci of 2 somewhat mamilliform conical, shorter than broad at base. 

Measurements (25 specimens from 7 localities) in millimetres. 


Range Mean 
Head width across eyes I-43-1-70 , f 1°54 
Greatest diameter of eye 0°44-0'55 ‘ 3 0°49 
Ocellus : : : O-II-O-15 X O°13-O'2I . : 0-13 X 0-18 
Ocelliss:oreyeuue: : 0:08-0°13 : d 0-10 
Width of pronotum . I-18-1-43 3 : I-32 
Length of pronotum . 0-70-0°88 : : 0-79 
Length of hind tibia. 1°75-2'13 : : 1°93 
Length of fore wing . 11-40-1310 ; : £232 


Soldier. ead capsule orange-yellow to ferruginous orange, nose ferruginous orange to 
chestnut brown. Pronotum and antennae as head capsule. Abdominal tergites and femora of 
legs yellow, abdominal sternites, tibiae and tarsi pale yellow. 

Head capsule in plan view varies with size : smaller specimens oval, tapering towards front, 
medium sized, near circular, largest, wider than length to base of nose, often indented in mid 
line at back of head, and sometimes with small projection on each side of base of nose ; nose 
weakly to moderately conical. In profile distinctly concave, with back of head raised and 
rounded, except in smallest specimens, often somewhat sinuate with slight hump at base of nose ; 
nose distinctly shorter to distinctly longer than rest of head capsule, measuring from hind 
margin of antennal socket, often within one colony, angle of nose cone 5-17 degrees ; post- 
mentum slightly convex ; antennae 12-14 segmented, usually 13, proportions vary with size, 
III long, up to twice II, 1V and V subequal, slightly longer than II. Head setae confined to the 
hairy nose tip and scattered over entire surface of postmentum ; dorsal surface of head capsule 
without prominent setae. 

Abdominal tergites with scattered minute setae, and longer setae on posterior margins of last 
three or four segments ; sternites with scattered shorter setae, and long setae on posterior 


margins. 
Measurements (144 specimens from 35 localities) in millimetres. 

Range Mean 
Head length to tip of nose. 1°40-2°1I B : : - 1:93 
Head width . : : : 0°73-1°35 ; ; ; é 1:07 
Depth of head capsule . : 0°53-0°95 ; : : : 0:76 
Width of pronotum ; : 0-38-0-65 ‘ : : ; 0°50 
Length of pronotum : : 0-13-0°38 : é : : 0:21 
Length of hind tibia . : 0:93-1°73 1:26 


Variation. Imagos from the eastern end of the range od to Z slightly een and to have 
a slightly paler postclypeus than those from the west coast of Africa, as in certain other species. 

The soldiers are more variable than in any other species in size, and in the shape of both 
profile and plan views of the head. The smallest specimens included in the above range of 
measurements are somewhat nanitic, but even in the more mature colonies the smaller soldiers 
may be only 1-55mm. long. The nanitic soldiers have a much less concave profile, with the 
vertex rather abruptly raised. In mature colonies soldiers apparently derived from more than 
one instar occur, suggesting a residual or rudimentary polymorphism, as in some other species. 


ETHIOPIAN NASUTITERMITINAE 43 


The development of the postmentum setae in the soldier appears to form a cline. They are 
most numerous in specimens from West Africa, gradually decreasing southwards and eastwards, 
until in the South-East Congo specimens they are sometimes very sparse. It is necessary where 
they appear absent to examine numerous specimens, because in most samples, some will usually 
be found with more than the rest. 


The nanitic soldiers of N. latifrons have been confused with N. /ujae, from which 
they differ in having no vertex setae, II or 12 segmented antennae, and usually a 
few postmentum hairs. N. schoutedeni resembles the shorter nosed forms in shape, 
but also has vertex setae, a much more abruptly tapered nose tip, and usually, no 
postmentum hairs, though one or two may be present very rarely. N. kempae 
lacks postmentum hairs, often has vertex setae, and usually a straight or slightly 
convex profile. N. arborwm usually has a straight profile, but also only four main 
setae at the nose tip, and none on the postmentum surface. 

The imago is distinguishable from most species by its larger size. Differences 
from N. chrysopleura and N. schoutedeni are discussed under those species. 

Though he described N. Jatifrons in 1896, Sj6stedt (1900) referred further specimens 
to N. arborum, and was followed in this by Silvestri (1912) and subsequent authors. 
The result has been that all the references to N. arborum since its original description 
have in fact been based on specimens of N. latifrons. This has been discussed under 
N. arborum, and the position has been stabilized by the selection of a neotype of 
the latter. 


One of the syntype specimens of N. Jatifrons (Sjéstedt) in the Naturhistoriska 
Riksmuseum, Stockholm, has been selected and labelled as lectotype by Dr. A. E. 
Emerson. This is as follows :— 

Lectotype : CAMEROUN: Ekundu, r891 (Y. Sjéstedt, No. 964), lectotype 9 
and ¢ paralectotype (other paralectotype in A.M.N.H.). 

Type Material. CAMEROUN: Ekundu, r8gq1 (Y. Sjéstedt), paralectotype J imago 
N. latifrons (Sjéstedt), A.M.N.H. Victoria (Bucholtz), syntype soldier, Trinervitermes 
pius (Sj6stedt), A.M.N.H. (other syntypes in Naturhist. Riksmus., Stockholm). 

REPUBLIC OF Conco: Indo, Sibiti, 9.1i.1925 (A. Unsgaard), syntype soldiers, 
type colony, N. indoensis (Sjéstedt) A.M.N.H. (other syntypes in Naturhist. Riks- 
mus., Stockholm). 

Conco: Kunungu (H. Schouteden) syntype soldiers, type colony, N. incurvus 
(Sjéstedt), A.M.N.H. (other syntypes in Mus. Tervuren). Haut Uélé, Moto (Bur- 
geon), syntype soldier, type colony, N. weleensis (Sjéstedt) A.M.N.H. (other syntypes, 
in Naturhist. Riksmus., Stockholm) ; Bas Uélé, Djamba, 17.xii.1924 (H. Schoute- 
den), syntype soldier, type colony N. expulsus (Sjéstedt) A.M.N.H. (other syntypes 
in Naturhist. Riksmus., Stockholm). 


Other material. PoRTUGUESE GUINEA: Cassine, I.ii.1900 (L. Fea), A.M.N.H. 

GUINEA : Nimba Mountains, viii.1957, 4 vials (M. Lamotte), Inst. Franc. d’ Afr. 
Noire, Dakar. 

SIERRA LEONE: 60m. from Freetown, 24.i.1955 (W. V. Harris). 

GHANA: Aburi, 15.i.1913 (F. Silvestri), AMM.N.H.; Bobiri Forest, 23m. from 
Kumasi on Accra Road, 2.11.1955 (W. V. Harris), and 21.11.1959 (W. A. Sands) ; 


Ai W. A. SANDS 


Kumasi, 22.x1.1957 (W. Wilkinson) ; 63m. from Accra on Kumasi Road, 18.11.1959, 
and 54m. from Dunkwa on Obuasi Road, 4.iv.1959 (W. A. Sands). 

NIGERIA: Eastern Region; Bugeme, 5.1.1957, 25m. from Onitsha on Owerrl 
Road, 11.1.1957, 39m. from Port Harcourt on Owerri Road, 31.i.1957, 3m. from 
Ikanga on Calabar Road, 22.iv.1957, 29m. from Port Harcourt, on Owerri Road, 
19.vi.1957, 8m. from Port Harcourt on Owerri Road, 10.xii.1957, Port Harcourt, 
19.xll.1957 (W. Wilkinson). Western Region ; Agege, 1914 (A. D. Peacock) ; Sapobar 
and Sapele, 29.iv.1955 (EZ. A. J. Duffy) ; 34m. from Ibadan on Ife Road, 1955 (B. J. 
MacNulty) ; Obanokoro, 31.1.1956 (D. Kay) ; Sobo Plain, Obanokoro, 7.i11. and 
7.Vil. 1957, and Lagos, 10.iv.1957 (W. Wilkinson) ; 27m. S. of Ilorin on Oyo Road, 
4.X11.1957, 4 vials, 63m. S. of Ilorin on Oyo Road, 4.xii.1957, 2 vials, Ararome near 
Ibadan, 9.x11.1957, Balogun, 30m. E. of Ibadan on Ife Road, 11.x11.1957, and 30m. 
from Ijebu-Ode on Benin Road, 12.xil.1957, 2 vials (W. A. Sands) ; Ibadan, 
1959 (H. J. Sutton). Northern Region ; 11m. from Mokwa on Bida Road, 
22.x11.1956, 1om. N. of Bida, 23.xii.1956, Idu Forest, 20m. from Abuja on Keffi 
Road, 28.xii.1956, 2 vials, and Ayangba (Egume), 6.i111.1958 (W. A. Sands). 

CAMEROUN : Yaounde, 2 vials (Zenker), AM.N.H.; Longji, 3 vials (Blank) ; 
A.M.N.H.; Edea, A.M.N.H.; Debundscha (Linnel), A.M.N.H.; Victoria, 
6.1.1913, 2 vials (F. Silvestri), A.M.N.H.; Attogondame, 8.vi.1913 (Eschertch), 
A.M.N.H.; Nyong Forest, 30.xi.1949 and 24.11.1950 (J. Burket-Smith, J. Dahl) ; 
Mbalmayo, 17.x.1962 (G. Becker), own collection. 

ISLAND OF SAO THOME: 28.vii.1932 (W. H. T. Tams). 

GaBon : Libreville, ii.1913 (F. Szlvestri), A.M.N.H. 

REPUBLIC OF THE CoNGO: 13km. W. of Brazzaville, 8.vi.1948, 2 vials (A. E. 
Emerson), A.M.N.H. 

Conco : Stanleyville (H. Kohl), AMM.N.H. ; Kasai, Kondue (£. Luja), A.M.N.H. ; 
Sankuru, A.M.N.H. ; Ganda Sundi, Mayumbe, I0.vii.1g1I, 5 vials, and Limba, 
IO.vil.Igir (R. Mayné), A.M.N.H.; between Walikale and Lubutu, 25.1.1913 
(J. Bequaert) ; Lukula, vii.1g20, Kai Bumba, 10.x.1920, Eala, i.1921, Luebo, 
ix.1921, Kidada (Kitobola), 11.1922, and Kasai, Ngombe (H. Schouteden), 
A.M.N.H.; Stanleyville, 18.iii.1948 (N. A. Weber), A.M.N.H.; Leopoldville, 
3.1v.1948, 4 vials, Sona Mpungu, 12.iv.1948, 2 vials, 35km. E. of Mambasa, 
10.v.1948, Camp Putnam, Epulu, 12-23.v.1948, 13 vials, Tshopo River, on Epulu 
Stanleyville Road, 25.v.1948, Stanleyville, 25-27.v.1948, 5 vials, Yangambi, 
29.v.1948, 3 vials, Avakubi, on Ituri River. v.1948, 15km. S.W. of Leopoldville, 
6.vi.1948, 2 vials, and 2okm. E. of Ndjili and 18km. S. of Leopoldville, 10.vi.1948 
(A. E. Emerson), A.M.N.H. ; Garamba National Park, 26.v.1950 (G. Demoulin), 
Inst. des Parcs Nat. du Congo, Brussels. 

SuDAN : Aloma Plateau, Khor Aba, vii—vili.1939 (N. A. Weber), A.M.N.H. 

UGanDA: Entebbe (O. John), A.M.N.H.; Namanwe, iv.1939 (G. A. E. 
Hopkins) ; Budongo Forest ix.1939 (C. C. Gowdey) ; Kyagwe, Lugala Estate, 
29.vi.1949, and Budongo Forest, 8.vii.1962, 3 vials (W. V. Harris). 

A total of 126 nest series were examined, and all material is in the British Museum 
(Natural History) unless otherwise stated. 


ETHIOPIAN NASUTITERMITINAE 45 


This is the commonest and most widely distributed species of Nasutitermes in 
Africa. It is found throughout the forest areas in West Africa and the Congo, and 
in most forest outliers, or riverain gallery forests, in the adjacent savannah areas. 

In none of the material examined of N. latifrons was an arboreal nest recorded of 
the type found in N. arborum, N. lujae, or N. schoutedem. The great majority of 
records are from dead logs, hollow or rotten tree trunks, and the mounds of other 
genera such as Culitermes, Procubitermes, Thoracotermes, and Microcerotermes. 
Sjéstedt (1926) recorded a nest illustrated by Hegh (1922 : 485, fig. 343), as belong- 
ing to this species, but the figure is a photograph of a typical mound of Thoraco- 
termes. Runways up trees are frequently recorded, but the nest can only rarely, 
if ever, be arboreal. 


Nasutitermes lujae (Wasmann) sp. rev. 
(Text-figs. 20, 21, 88-94, 213, 214: Map 11) 


Eutermes lujae Wasmann, 1911 (August) : 168. Type locality : Conco, Kasai, Sankuru. 
sp. rev. [ex syn. N. maculiventris (Sjéstedt, 1926 : 320). 

Eutermes ekunduensis SjOstedt, 1911b (Decembér) : 16. Type locality : CAMEROUN, Ekundu. 
syn. n. 

Eutermes (Eutermes) lujae Wasmann ; Holmgren, 1912 : 62. 

Eutermes (Eutermes) ekunduensis Sjostedt ; Holmgren, 1912 : 62. 

Eutermes (Eutermes) nanus Sjostedt, 1924 : 41. Type locality : Conco, Barumbu. 

Eutermes (Eutermes) impetus Sj6stedt, 1924b : 493. Type locality : Rerpusiic or Conco, 
Brazzaville. syn. n. 

Nasutitermes (Nasutitermes) bequaerti Emerson, 1928 : 416. [New name proposed for E. nanus 
Sjéstedt}]. syn. n. 

Nasutitermes (Nasutitermes) santschii Emerson, 1928 : 484. Type locality : Conco, Stanley- 
ville. syn. n. 

Nasutitermes (Nasutitermes) torquatus (Sjéstedt) ; Emerson, 1928 : 481. 

Nasutitermes bequaerti Emerson ; Snyder, 1949 : 267. 

Nasutitermes ekunduensis (SjOstedt) Snyder, 1949 : 275. 

Nasutitermes impetus (SjOstedt) Snyder, 1949 : 279. 

Nasutitermes santschii Emerson ; Snyder, 1949 : 297. 


Imago. Head capsule dark sepia brown, postclypeus slightly paler, brown. Labrum yellow- 
brown. Antennae brown. Pronotum dark sepia brown, meso- and metanota, pleural and 
ventral thoracic sclerites brown. Legs yellow-brown, brown at ends of femora and bases of 
tibiae. Abdominal tergites dark sepia brown, sternites sepia brown laterally, shading to pale 
brown in middle. Wings translucent pale brown, venation sepia brown. 

Posterior margin of head behind eyes distinctly less than semi-circular ; fontanelle variable, 
generally inconspicuous, from a pale slit, sometimes weakly bifurcate in front, to a smooth flat 
patch without setae, of same colour as rest of head capsule, or sometimes slightly depressed 
posteriorly ; eyes moderately prominent in proportion to own diameter, but not large relative 
to head width, (W/E index 3-0-3-5) nearly circular, outline slightly angular ; ocelli small, 
separated from eyes by slightly less to distinctly more than own least diameter ; postclypeus 
rather weakly inflated, width 3-2 to 3-7 times length, anterior margin nearly straight, posterior 
margin convex, rounded, very slightly sinuate towards outer corners ; antennae 15 segmented, 
II longer than IV, IV slightly longer than V, V longer than III. 

Cerci of 9 distinctly shorter than width across base, mamilliform. 


46 W. A. SANDS 


Measurements (22 specimens from 5 localities), in millimetres. 


Range Mean 
Head width across eyes I-18-1°33 é é 1:26 
Greatest diameter of eye 0:36-0°42 : 5 0°39 
Ocellus : ; : 0:09-0-12 X 0-13-0'18 .. : O-II X O-14 
Ocellus toeye  . : 0:09-0'13 : : 0-10 
Width of pronotum . 0-95-1°18 : , 1-04 
Length of pronotum . 0:60-0:70 “ : 0-66 
Length of hind tibia. 1°38-1°65 5 . I-51 
Length of fore wing. 10-90-11:80 4 : II-40 


Soldier. Head capsule orange-yellow to ferruginous orange, nose ferruginous orange to 
chestnut brown. Pronotum and antennae, orange-yellow. Abdominal tergites and legs yellow, 
abdominal sternites pale yellow. 

Head capsule in plan view a short oval, tapering slightly towards front, nose conical. In 
profile straight or very weakly sinuate to well behind base of nose, then slightly but rather 
abruptly raised and rounded to back of head ; specimens with completely straight profile are 
rare. Nose equal in length or slightly longer than rest of head capsule, measuring from hind 
margin of antennal socket, angle of nose cone 13—20 degrees ; postmentum weakly and evenly 
convex ; antennae 12 or 13 segmented, only one in ten with 12, proportions of segments some- 
what variable, in 13 segmented forms, IIT longer than or equal to II and V, IV shorter than these. 
Setae on head confined to a bilaterally placed pair on vertex, four at tip of nose with smaller 
setae behind them, and a few on anterior margin of postmentum. 

Abdominal tergites with very short sparsely scattered setae, and longer setae on posterior 
margins of last three or four segments ; sternites with more numerous short setae, and longer 
setae on posterior margins. 

Measurements (59 specimens from 14 localities) in millimetres. 


Range Mean 
Head length to tip of nose. 1:25-1-63 : : : 5 1°43 
Head width . : : : 0-68—0°95 : : : é 0:80 
Depth of head capsule . : 0-49-0°63 - ; : : 0:56 
Width of pronotum , : 0:38-0°48 5 : : : 0°43 
Length of pronotum ; ; 0:15-0:20 5 ; ‘ : Onn 
Length of hind tibia ‘ ‘ 0:95-1:18 1-04 


Variation. The imago is less variable in respect of size oad bear fae N. eoruan (Smeath.). 
Soldier variation consists mainly of slight differences in the thickness and length of the nose and 
the straightness of the profile. Specimens occasionally occur with one or two setae at the base 
of the nose in addition to those on the vertex. 


The distinctions between this and the closely similar N. arboruwm have been 
discussed under that species. Darker, thicker-nosed specimens of the soldier have 
been confused with N. ferranti, but this has two or four setae in line across the top 
of the base of the nose, and only four setae at the nose tip. The smallest specimens 
of N. schoutedeni, with flatter profiles, are very closely similar, but generally have the 
nose distinctly shorter than the rest of the head capsule, and more abruptly tapered 
at the tip. The imago of that species is also closely similar, but larger, and has the 
wing membrane less hairy but with more distinct asters (this character cannot 
readily be used in a key or illustrated, but is useful if material is available for com- 
parison). The soldier of N. infuscatus is similar, but usually has a shorter nose ; 
the imago is quite distinct with a longer more inflated postclypeus. 

Sjéstedt (1926) placed the name Eutermes Iuwjae Wasmann in synonymy with 
E. maculiventris Sjéstedt. The syntypes from the type colony of both have been 


ETHIOPIAN NASUTITERMITINAE 47 


examined, and they are undoubtedly different. The former is the same as the 
species hitherto called N. ekunduensis, and the latter, N. arborum. Both N. lwae 
and N. ekunduensis were described in 1911, but there is no doubt of the priority of 
the name N. lwjae which is now removed from synonymy with N. arborum (= 
N. maculiventris). 

Type Material. CAMEROUN: Ekundu, 3.x1i.1890 (Y. S7dstedt), syntype soldier, 
type colony, N. ekunduensis (Sjéstedt), A.M.N.H. (other syntypes in Naturhist. 
Riksmus., Stockholm). 

REPUBLIC OF CONGO: Brazzaville (Roubaud), syntype soldier, type colony, 
N. impetus (Sjéstedt), A.M.N.H. (other syntypes in Mus. Paris and Naturhist. 
Riksmus., Stockholm). 

Conco: Kasai, Sankuru, 1907 (E. Luja), syntype soldier, type colony, Eutermes 
lwjae Wasmann, A.M.N.H.; Barumbu (J. Bequaert), syntype soldiers, type colony, 
Eutermes nanus Sjéstedt (N. bequaerti Emerson), A.M.N.H. (other syntypes in 
Naturhist. Riksmus., Stockholm) ; Stanleyville, 1913 (H. Kohl), paratype soldiers 
and g and 92 imagos, N. santschit Emerson, B.M.(N.H.) and A.M.N.H., including 
type colony, (Holotype in A.M.N.H.). 


Other Material. Ivory Coast: Abengouru, xi.1963, 3 vials (G. Becker), own 
collection. 

GHANA: 7m. S. of Akumadam on Wenchi-Kumasi Road, I.iv.1959 (W. A. 
Sands), B.M.(N.H.). 

CAMEROUN: Nyong Forest, 3.i.1950 (J. Birket-Smith, J. Dahl), B.M.(N.H.) ; 
Longji, undated (Blank), A.M.N.H.; Mbalmayo, 17.x.1962 (G. Becker), own 
collection. 

Conco: Kasai, Kondue, 1919 (EF. Lija) ; Stanleyville, 24.vii.1913, and others 
(H. Kohl) ; Avakubi, 6.i.1914 (J. Bequaert) ; Basongo, vii.1g21 (H. Schouteden) ; 
Stanleyville, 18.iii.1g48 (N. A. Weber) ; Camp Putnam, 15.v.1948, 2 vials, Epulu, 
15.v.1948, 4m. N. of Camp Putnam, 20.v.1948, 2 vials, Yangambi, 29.v.1948, 
3 vials (A. E. Emerson), all in A.M.N.H. 

UGANDA: Sese Islands, Lake Victoria, Bugalla Island, 1918 (C. C. Gowdey), 
B.M.(N.H.). 


A total of 29 nest series were examined. 


This species has been recorded from Ghana to Uganda, but is much less common 
in the north-western parts of its range, and appears to become more numerous in the 
central and southern Congo Forest. This contrasts with N. avborum, and although 
they have both been collected in the same locality in parts of their range, it would 
appear that they occupy a similar ecological niche, and that the conditions which 
suit one may be less favourable to the other. N. arborum has certainly been found 
in drier, more marginal forest outliers, than has N. /wjae, and a greater tolerance of 
drier forest conditions would account for its greater abundance in West Africa. 

The nest of N. lwjae is commonly arboreal, and similar to that of N. arborum 
though usually slightly smaller. 


48 WA SAUNIDS 


Nasutitermes schoutedeni (Sjostedt) 
(Text-figs. 22, 23, 95-99, 215, 216; Map 12) 


Eutermes (Eutermes) schoutedeni Sjéstedt, 1924 : 40. Type locality : Conco, Basongo. 

Eutermes (Eutermes) konduensis Sjostedt, 1924 : 40. Type locality : Conco, Kasai, Kondue. 
syn. n. 

Eutermes (Eutermes) putidus Sj6stedt, 1924: 40. Type locality : Conco, Kasai, Kondue. 
syn. n. 

Eutermes (Eutermes) aethiops Sjostedt, 1924 : 41. Type locality : Conco, Luebo. syn. n. 

Eutermes (Eutermes) dulcis Sjostedt, 1924b : 494. Type locality : Conco, Haut Uélé, Moto. 
syn. n. 

Eutermes schoutedeni Sjéstedt ; Sjéstedt, 1926 : 314. 

Eutermes konduensis SjOstedt ; Sj6stedt, 1926 : 307. 

Eutermes putidus Sj6stedt ; Sjdéstedt, 1926 : 314. 

Eutermes aethiops Sjostedt ; Sjdstedt, 1926 : 318. 

Eutermes dulcis Sj6stedt ; Sjdstedt, 1926 : 297. 

Nasutitermes chapint Emerson, 1928 : 480. Type locality : Conco, Ngayu. syn. n. 

Nasutitermes schoutedeni (Sjéstedt) Snyder, 1949 : 297. 

Nasutitermes konduensis (SjOstedt) Snyder, 1949 : 282. 

Nasutitermes putidus (Sj6stedt) Snyder, 1949 : 294. 

Nasutitermes aethiops (Sjéstedt) Snyder, 1949 : 265. 

Nasutitermes dulcis (Sj6stedt) Snyder, 1949 : 274. 

Nasutitermes chapini Emerson ; Sands, 1957 : 8. 


Imago. Previously undescribed. Head capsule dark sepia brown, postclypeus and antennae 
sepia brown, labrum brown. Pronotum and abdominal tergites dark sepia brown ; meso- and 
metanota and lateral parts of abdominal sternites sepia brown, legs and middle of abdominal 
sternites yellow-brown. Wings, membrane pale translucent sepia brown, subcosta and radial 
sector yellow-brown, radial sector lined posteriorly with dark brown ; other venation brown. 

Posterior margin of head capsule behind eyes evenly rounded, less than semi-circular ; fon- 
tanelle small, pale, weakly bifurcate in front ; eyes moderately prominent in proportion to own 
diameter, slightly angular oval, medium sized relative to head width, W/E index 2-9-3:1 ; 
ocelli medium sized, separated from eyes by a half to threequarters own least diameter ; post- 
clypeus weakly inflated, width 3-6 times length, anterior margin slightly sinuate, posterior 
convex, almost straight in middle, curved to outer corners ; antennae 15 segmented, II longer 
than IV, IV longer than V, V longer than III. 

Cerci of 2 about as long as width across base. 

Measurements (8 specimens from 2 localities) in millimetres. 


Range Mean 
Head width across eyes I-33-1°40 : ; ony 
Greatest diameter of eye 0:44-0°46 : : 0°45 
Ocellus : : : O-II-0-13 X O'15-O'18_ : O72 >< (O56) 
Ocellus toeye  . : 0:08-0:10 : : 0:08 
Width of pronotum I-IO—-I‘17 : ‘ 114 
Length of pronotum . 0:66-0°74 : ; o-71 
Length of hind tibia. 1:58-1:68 : é 1-63 
Length of fore wing. II-50—12°40 5 : II-go 


Morphotype. Ivory Coast: Abidjan, xi.1949 (R. Geigy). Morphotype Q in 
American Museum, Natural History. 


Soldier. Head capsule orange-yellow to ferruginous orange, nose ferruginous to chestnut 
brown with reddish tip. Pronotum and antennae as head capsule. Abdominal tergites and 
legs yellow, abdominal sternites pale yellow. 


ETHIOPIAN NASUTITERMITINAE 49 


Head capsule in plan view short oval to circular, nose conical with abruptly tapered tip. In 
profile distinctly concave, often with slight hump at base of nose, back of head raised and 
rounded. Nose shorter than rest of head capsule, measuring from hind margin of antennal 
socket, in all but smallest specimens ; angle of nose cone 12-15 degrees ; postmentum moder- 
ately and evenly convex ; antennae 12-14 segmented, commonly 13, II and IV subequal, 
shorter than V, III longer than V. Head setae consist of a bilaterally placed pair on vertex, 
not always present, a distinctly hairy nose tip, and a few on anterior margin of postmentum ; 
rarely one or two minute setae scattered on rest of postmentum, and usually a few on vertex. 

Abdominal tergites with very short scattered setae, and longer setae in four rows on posterior 
margins of last four segments ; sternites with scattered short setae, and six long setae on posterior 
margin of each segment. 

Measurements (39 specimens from 9 localities) in millimetres. 


Range Mean 
Head length to tip of nose. 1°43-1°75 2 P ; : 1°59 
Head width . : P : 0:85-1°13 . : : : 0-98 
Depth of head capsule . : 0°55-0°78 ; 2 é ; 0-68 
Width of pronotum : ¢ 0°40-0°57 , : ; - 0-49 
Length of pronotum - : 018-024 F ; : - o-19 
Length of hind tibia F : 1-05—-1°36 - : : ; I°I2 


Variation. Sjostedt (1924a) recorded specimens of N. putidus ranging in size up to head 
length 1:96, width 1-35, and depth 0-85 millimetres ; no such measurements have been encoun- 
tered from a wider range of specimens, including paratypes from type colony. The latter fall 
outside the range given by Sjéstedt, and I therefore consider that his measurements, though not 
impossible, are probably incorrect ; particularly since the measurement of head width for N. 
konduensis is given as 1:35mm., for a length of 1-65mm. and a depth of o-60mm., and would 
thus be of quite different proportions from all the other specimens if correct. Paratypes from 
type colony of N. konduensis measured 0-95mm. in width for a length of 1-58mm. 


N. schoutedent was described four times under different names in the same paper 
by Sjéstedt (1924), three times on the same page, and once again in another paper 
the same year (1924b). The name chosen has page priority, and is not otherwise 
inappropriate. 

It is most closely related to N. latifrons, which it closely resembles in the soldier 
caste, but is distinguishable by the presence of vertex setae, and the absence of 
postmentum setae except on the anterior margin (with rare exceptions). The imago 
is smaller, and the wings have setae distributed more or less evenly over membrane 
and veins, whereas in N. Jatifrons the veins carry a distinct row of setae with a bare 
border on each side. 

The smallest soldiers of N. schowtedeni have a straighter head profile, and closely 
resemble N. lujae ; some specimens are probably indistinguishable, but JN. 
schoutedent usually has a more abruptly tapered and closely hairy nose tip. The 
imago is also very like that of N. lwjae, but larger, with the ocelli closer to the eyes, 
and the wing membrane is less hairy with larger more distinct asters. 


Type Material. Conco: Basongo, vii.tg2t (H. Schouteden), syntype soldier, 
type colony, N. schoutedeni (Sjéstedt), A.M.N.H. (other syntypes in Naturhist. 
Riksmus., Stockholm) ; Kasai, Kondue (£. Luja), syntype soldier, type colony, 
N. putidus (Sjéstedt), A.M.N.H. (other syntypes in Congo Mus., Brussels) ; Luebo, 
ix.1g2I (H. Schouteden), syntype soldier, type colony, N. aethiops (Sjéstedt), 


50 W. A. SANDS 


| Map 10, NAsUuTITERMES LATIFRONS 


)} Mae 11, NASUTITERMES LUJAE 


Map 12, NASUTITERMES SCHOUTEDENI 


Maps 10-12. (10), Nasutitermes latifrons. Vegetation types: 7,8, and 16; (11), 
Nasutitermes lujae. Vegetationtype: 7; (12), Nasutiteymes schoutedeni. Vegetation 


type: 7: 


ETHIOPIAN NASUTITERMITINAE 51 


A.M.N.H. (other syntypes in Naturhist. Riksmus., Stockholm). Haut Uélé, Moto 
(L. Burgeon), syntype soldier, type colony, N. dulcis (Sjéstedt), A.M.N.H. (other 
syntypes in Mus. Tervuren) ; Ngayu, 22.xii.1g09 (H. Lang, J. P. Chapin), para- 
type soldiers, type colony, N. chapint Emerson, A.M.N.H. and B.M.(N.H.). (Holo- 
type soldier in A.M.N.H.). 

Other Material. Ivory Coast: Abengouru, xi.1963, 2 vials (G. Becker), own 
collection. 

NIGERIA : Western Region ; 30m. from Ijebu-Ode on Benin Road, 12.xii.1957 
(W. A. Sands). 

CAMEROUN : no data, from N. Holmgren, A.M.N.H.; Ekododo, 9.ix.1913 
(Escherich), A.M.N.H. 

Conco: Kasai, Kondue, 2 vials (E. Liya) ; Basongo, vii.ig21, and Luebo, 
ix.1g21 (H. Schouteden) ; Epulu, 17.v.1948 (A. E. Emerson), A.M.N.H. 

Ucanvda: Ankole Province, xi.1939 (H. B. Johnston). 

A total of 18 nest series were examined, in the British Museum (Natural History) 
unless otherwise stated. 


Although widely distributed from the Ivory Coast to Uganda, this species is not 
common. The nest is arboreal, more or less spherical, and approximately one foot 
in diameter. 


LEPTOMYXOTERMES gen. n. 


(Leptos, Gr. “‘ thin ’’, myxa, Gr., ‘‘ lamp-nozzle, spout ’’) 

Type-species, Eutermes doriae Silvestri, 1912 : 242. 

Imago. Left mandible with apical tooth as long as first marginal, distance from apical to 
first marginal half distance between first and third marginals (left mandible index -50), cutting 
edge between first and third marginals with distinct indentation in front of third marginal ; 
right mandible with apical and first marginal subequal, posterior cutting edge of first marginal 
about equal to that of second, this being distinctly concave ; right molar plate in surface 
view with indentation near basal end of anterior margin, posterior margin extending in rounded 
flange, first molar ridge forming tooth, remaining 6—7 ridges diminishing towards base (Text-figs. 
164, 165). Fontanelle slit-like, not clearly bifurcate anteriorly. Eyes and ocelli very large and 
prominent. 

Soldier. Monomorphic with traces of dimorphism. Nose cylindrical. Vestigial mandibles 
with small points. Head capsule sharply constricted behind antennae. Antennae 14 
segmented. 


This genus is proposed to accommodate the single African forest species doriae 
Silvestri, hitherto included with Rhadinotermes in Coarctotermes, which is now restric- 
ted to the Malagasy species. Leptomyxotermes is clearly widely different both 
ecologically and morphologically from the other African constricted-headed genera. 
Although it shares with Fulleritermes the indentation of the cutting edge between 
first and third marginals of the left mandible, their proportions are different, and the 
small tooth connecting the molar prominence to the third marginal is retracted 
below the level of the groove between them, whereas in Fulleritermes and Rhadino- 
termes it is clearly visible in surface view. The concavity of the cutting edge of the 
second marginal of the right mandible is more pronounced than in the other genera, 


52 Wr. TAC SAINIDIS 


and the first and second marginals are in line instead of overlapping slightly. The 
fontanelle and large eyes are also very distinctive. 


Leptomyxotermes doriae (Silvestri) comb. n. 
(Text-figs. 100, IOI, 119-126, 163-165; Map 13) 


Eutermes doriae Silvestri, 1912 : 242. Type locality : ISLAND oF FERNANDO Poo, Basilé. 
Eutermes (Hospitalitermes) doviae Silvestri ; Holmgren 1912 : 66. 
Coarctotermes doriae (Silvestri) Sj6stedt, 1926 : 358. 


Imago. WHead capsule orange-yellow, clouded with brown between eyes and on frons, darkest 
on upper rim of ocelli ; postclypeus and labrum, yellow, antennae yellow-brown. Pronotum, 
meso- and metanota, other thoracic sclerites and legs, yellow. Abdominal tergites brown, 
sternites pale yellow-brown, paler in mid-line. Wings translucent very pale brown, venation 
brown, darker at base ; radius sector with brown inner margin, and with adjacent yellow streak 
in distal half of wing. 

Posterior margin of head much less than semi-circular behind eyes, slightly sinuate im- 
mediately behind eyes ; eyes oval, very prominent, and large relative to head width, W/E 
index 2-4; ocelli large, oval, scarcely separated from eyes ; postclypeus strongly inflated, 
width 1-8 times length, anterior margin straight, posterior regularly convex ; antennae 15-16 
segmented, II and IV subequal, III longer, or II and III subequal, IV shorter. 

Cerci of 2 mamilliform, shorter than width across base. 

Pilosity of vertex confined to a few long paired setae. 

Measurements (4 specimens from 2 localities) in millimetres. 


Range Mean 
Head width across eyes I-15—1-20 : . 1-18 
Greatest diameter of eye 0-48-0'51 : ; 0-50 
Ocellus : ‘ : 0°15 X O*19Q—-0-22 E = § 0-15) 5< 0-20 
Ocellus toeye  . : 0-OI—0-02 : : 0-01 
Width of pronotum ; 0:97-1:00 : ’ 0-99 
Length of pronotum . 0:59-0:68 : : 0-61 
Length of hind tibia. 1:54-1:66 F : 1-62 
Length of fore wing . 10-80 


Soldier. Head capsule yellow to orange, nose orange to ferruginous. Pronotum, antennae 
and abdominal tergites orange-yellow, legs and abdominal sternites yellow. 

Head capsule in plan view with angularly rounded posterior margin. In profile depressed 
behind base of nose or straight in front of raised posterior lobe ; nose shorter to longer than rest 
of head measuring from hind margin of antennal socket, maximum angle of taper 5 degrees ; 
antennae 14 segmented, elongate, II short, IV, V and VI subequal, longer than II, III longer 
than these, twice length of II. Setae on head confined to four at nose tip and a bilaterally 
placed pair on vertex. 

Abdominal tergites almost glabrous except for 4-6 long yellow setae on posterior margins of 
each of last 4-5 segments ; sternites with scattered very sparse short setae and longer setae on 
posterior margins 3 or more times length of short. Remaining characters given in generic 


diagnosis. 
Measurements (26 specimens from 5 localities) in millimetres. 
Range Mean 
Head length to tip of nose. 135-1 :93 : : 5 é 1-67 
Head width . : : : 0°55-I1-00 : : ; : 0:77 


Depth of head-capsule . : 0°43-0°75 : : : : 0-58 


ETHIOPIAN NASUTITERMITINAE 53 


Range Mean 
Width of pronotum : F 0°43-0°50 ‘ : ° P 0°47 
Length of pronotum 2 : 0:18-0:25 : : : : 0-21 
Length of hind tibia a : I+23-1°73 : : : : 1°43 


Variation. The profile view of the soldier is the main feature of variation in this species. 


L. doriae is unlikely to be confused with any other African species, and its dis- 
tinguishing features are discussed under the generic heading. It appears to be 
related far more closely to the Hospitalitermes — Lacessititermes group of constricted- 
headed genera than to the Coarctotermes — Fulleritermes — Rhadinotermes group, and 
in assigning the species to Hospitalitermes, Holmgren (1912) was nearer to the truth 
than subsequent authors. Common features in the soldier are the distribution of 
setae and the long appendages, and in the imago, the large eyes, pale colour, and 
not distinctly bifurcate fontanelle, the retracted tooth between left third marginal 
and molar prominence, and the reduced overlap of right first and second marginals. 
The right molar plate is however distinctly different ; and this, together with the 
stronger notch in the left mandible, the more concave right second marginal, and 
the better developed apical teeth, may be an indication of a more primitive position 
for L. doriae in this group. 


Material. ISLAND OF FERNANDO Poo: Basilé, vili—ix.1gor (L. Fea), syntype 
soldiers, type colony, L. doriae (Silvestri), Silvestri coll., Lab. Zool. Sc. Agr., Portici, 
A.M.N.H., and B.M.(N.H.). 


GUINEA : Nimba Mountains, 1957, 4 vials (M. Lamotte), Inst. Frang. d’Afr. 
Noire, Dakar. 


SIERRA LEONE: Freetown, 7.1.1958 (W. Wilkinson). 
Ivory Coast: Banco Forest, 4.ii.1962 (C. Novrot). 


GHANA: Aburi 1912-13 (F. Silvestri), morphotype king, Silvestri coll., Lab. 
Zool. Sc. Agr., Portici ; 20m. S. of Techiman on Kumasi Road, 1.iv.1959 (W. A. 
Sands). 


NIGERIA : Western Region ; Benin Province, Obanokoro, Sobo Plain, 4.11. 
1957 ; Eastern Region ; 53m. from Port Harcourt on Owerri Road, 6.xii.1957, 
Port Harcourt, 18.xii.1959 (W. Wilkinson). 


CAMEROUN : Victoria, i.1913 (F. Silvestri), A.M.N.H. 
Conco : Ituri, Beni, Irumu, 24.11.1948 (N. A. Weber), A.M.N.H. 


A total of 14 nest series, in the British Museum (Natural History) except where 
otherwise stated. 


L. dorvae is confined to the rain forests of West Africa and the Congo. It appears 
to feed almost exclusively on damp, well rotted wood, and to nest in chambers in or 
under dead logs, or parts of the mounds of other species. It is never common, and 
the patchy distribution of records reflects the difficulty experienced by collectors 
rather than the occurrence of the species, which may extend to all parts of the 
Congo forest. 


Wik. SANDS 


54 


Te 


Vegetation type : 


Leptomyxotermes doriae. 


Map 13. 


ETHIOPIAN NASUTITERMITINAE 55 


GRALLATOTERMES Holmgren 


Eutermes (Grallatotermes) Holmgren 1912 : 65. [Subgenus of Euterymes Heer.] 
Grallatotermes Light, 1930: 16. Type-species by original designation, Termes (Eutermes) 
grallatoy Desneux, 1905. 


Imago. Left mandible with apical tooth slightly shorter than first marginal, distance between 
them one quarter of distance between first and third marginals (left mandible index -25), 
second marginal separated by deep notch in cutting edge in front of third marginal, small 
subsidiary tooth connecting third marginal and molar prominence retracted behind gap between 
them when viewed from above ; right mandible with apical and first marginal subequal, 
posterior cutting edge of first marginal shorter than that of second, not overlapping anterior 
edge of second (apparent overlap embedded in smooth shining fused surface), molar plate with 
no marked irregularity of outline in surface view, and 13-14 ridges, diminishing in size posterior- 
ly. Postclypeus short and lightly inflated, length approximately one-third width. Fontan- 
elle pale, roughly T-shaped. Pronotum almost without raised anterior margin. Pilosity of 
head rather sparse, fine, and uniform, with few more prominent setae. 

Soldier. Monomorphic. Nose thick but weakly conical, tip sharply tapered. Vestigial 
mandibles with points. Head capsule slightly constricted behind antennae, setae confined to 
four at nose tip, and bilaterally placed pairs on postclypeus and vertex. 

The imago mandibles of this genus are quite distinct from any others of the sub- 
family in Africa, in combining the notched cutting edge of the left mandible between 
first and third marginals with the many-ridged right molar plate. The latter feature 
is typical of the Hospitalitermes—Grallatotermes group of constricted-headed genera, 
as also is the retracted subsidiary tooth between left third marginal and molar 
prominence. Leptomyxotermes has a somewhat similar left mandible, but the molar 
plate and second marginal of the right mandible are very different. In Fudleritermes 
the subsidiary tooth between left third marginal and molar prominence is well 
developed and the right molar plate is distinct. 


Grallatotermes africanus Harris 
(Text-figs. 102, 103, 127-129, 166-168, 217, 218; Map 3) 

Grallatotermes africanus Harris, 1954 : 135. Type locality : TANGANyIKA, Tanga District, 

Ngomeni. 

Material (additional to that recorded in Harris, 1954). TANGANYIKA: 25m. 
from Dar es Salaam on Morogoro Road, 21.vii.1954 (R. M. C. Williams). 

MOZAMBIQUE: Ribaué, 20.v.1932 (J. Vincent). 

A total of 7 nest series were examined. 


The extension of the recorded distribution of this species was not unexpected, and 
further collecting may show that its distribution is similar to that of Nasutitermes 
kempae Harris, with Zululand as its southernmost extremity. 


FULLERITERMES Coaton 


Fulleritermes Coaton, 1962 : 151. Type-species, by original designation, Eutermes contractus 
SjOstedt, 1913Aa. 
Imago. Left mandible with apical tooth as long as first marginal, distance from apical to first 
marginal more than one-third distance between first and third marginals (left mandible index 
-36—-43), cutting edge between first and third marginals with notch separating third marginal ; 


56 W. A. SANDS 


right mandible with apical and first marginal subequal as left, first marginal with small subsidiary 
tooth near basal junction with apical, relative lengths of cutting edge of first and second 
marginals variable ; right molar plate in surface view with very weak to strong indentation near 
basal end of anterior margin, posterior margin extending in rounded flange, first molar ridge 
forming stout tooth, remaining 7-9 ridges diminishing towards base (Text-figs. 169-180). 
Fontanelle slit-like, distinctly bifurcate anteriorly, usually paler than head. 

Soldier. Mono- or dimorphic. Nose cylindrical or conical. Vestigial mandibles with small 
to medium-sized points. Head capsule scarcely to sharply constricted behind antennae, 
posterior lobe commonly more than one-fourth wider than anterior. Antennae 12-13 segmented. 

This genus was separated by Coaton (1962) from Coarctotermes on the basis of the 
notched cutting edge in front of the third marginal tooth of the left imago mandible. 
It contains four species, contractus (Sjdstedt), coatont sp. n., mally: (Fuller) and 
tenebricus (Silvestri). 

This character is convenient for separating Fulleritermes from Coarctotermes and 
Rhadinotermes, but it is shared by two other African constricted-headed genera, 
Grallatotermes and Leptomyxotermes ; however both these have much larger eyes 
and ocelli in the imago, and in Grallatotermes the right molar plate has many ridges. 

Within Fulleritermes itself there are specific differences. In F. mallyi the left third 
marginal tooth of the imago is asymmetrical, the anterior edge being longer than the 
posterior and slightly concave, and the right molar plate scarcely indented in outline. 
In the other species the left third marginal is equilaterally triangular and the right 
molar plate deeply indented in outline, with fewer ridges. The soldier caste of 
F. mallyi is also distinct in shape and colour from the other species which closely 
resemble each other. 

I do not consider it desirable to separate Ff’. mallyi generically from the other species 
at present because further work is required to evaluate fully the importance of these 
rather small differences in imago mandible characteristics. 

The genus occurs in the savannah and other drier open vegetation zones, and only 
occasionally enters the forest edges, being absent from the depths of true rain forest. 
Of the four species recognized in this paper, one occurs north of the Congo forest 
block, one to the south, one in tropical South Africa, (Transvaal, Natal, northern 
Cape Province) and the last in the Montane and Karroo vegetation of temperate and 
subtropical South Africa. There is apparently no overlap in their distribution, each 
exploiting a separate and distinct vegetation type or set of types. The genus appears 
to occupy a subsidiary or minor ecological “ niche’ in the termite fauna of these 
areas, being never found abundantly and usually scarce and difficult to discover. It 
may however be close to the evolutionary stem from which more abundant specialized 
genera such as Tvinervitermes arose. It is absent from East Africa, where Rhadino- 
termes is found, but it is thought that R. coarctatus occupies a rather different 
ecological “‘ niche ’’ because of the differences in its imago and worker mandibles. 


Krys to FULLERITERMES anp RHADINOTERMES 
Imaco (FULLERITERMES only) 
1 Antennae 14 segmented, though with segment III long, sometimes partly subdivided. 
Wings under 1omm. in length ° ; : : : : ; : 4 2 
— Antennae 15 segmented. Wings over 1omm. in length : : 3 mallyi (p. 65) 


ETHIOPIAN NASUTITERMITINAE 57 


118. 


Fics. 100-113 : Imago head capsule, front and side views, 114-118, vertex pilosity. 
100, 101, Leptomyxotermes doriae ; 102, 103, Grallatotermes africanus ; 104, 105, 114, 
Fulleritermes coatont ; 106, 107, 116, F. mallyi ; 108, 109, 118, F. tenebricus ; 110, 
III, 115, F. contvactus ; 112, 113, 117, Rhadinotermes coarctatus. 


58 W.A. SANDS 


2 Vertex of head viewed diagonally upwards with numerous yellowish bristle-like setae 


(Text-fig. 115) : ‘ contractus (p. 61) 
— Vertex with one or two peeeies oad pease peateeeed stories hairs, a few more 
bristles immediately behind eyes (Text-figs. 114, 118) ; 3 


3. Right mandible with cutting posterior edge of first marginal ipo eel eyed 
anterior margin of second marginal, longer (10 : 8) than equivalent cutting edge of 


second marginal (Text-fig. 171) (S. Africa, S. of Limpopo) . coatonisp.n. (p. 58) 
— Right mandible with cutting edge of first marginal shorter (8 : 10) than equivalent 
edge of second marginal (Text-fig. 180) (West Africa to Sudan) . tenebricus (p. 68) 
SOLDIER 


1 Nose slender, cylindrical, head capsule in plan view with sides converging towards 


front and slightly sinuate, scarcely constricted behind antennae F. mallyi (p. 65) 
— Nose more or less conical, head capsule in plan slightly to distinctly constricted 

behind antennae . ; : 4 - 2 : : : : . : 2 
2 Antennae 12 segmented . : , 2 : ; : : - : : 3 
— Antennae 13 segmented . : 4 


3. Colour paler, head capsule orange to Sere. nose as cou or aoe i pitch pEee 
F. coatoni sp. n. (p. 58) 
— Colour darker, head capsule be ee @ to pitch black, nose pitch black with reddish 
tip : : : . R. coarctatus (p. 71) 
4 Soldiers comes. “ Points ”’ ve este maedibles al but distinct. Head 
capsule in plan smoothly and regularly rounded behind constriction 
F. coatoni sp. n. (p. 58) 


— Soldiers monomorphic. “ Points’’ of vestigial mandibles minute or absent. Head 
capsule in plan slightly angularly rounded behind constriction, often weakly 
sulcate at back . ; : : ; ; 4 ; 3 : : 5 

5 West Africa, N. Congo, Sadan : : - 5 : . F. tenebricus (p. 68) 

— S$. Congo, Angola, N. and S. medical : : : : . F. contractus (p. 61) 


Fulleritermes coatoni sp. n. 
(Text-figs. 104, 105, 114, 130-134, 169-171, 219 ; Map 14) 


Coarctotermes contractus (SjOstedt) ; Fuller, 1922 : 117. 
Fulleritermes contractus (Sj6stedt) ; Coaton, 1962 : 152, [part : specimens identified by Fuller 

(1922) only]. 

Imago. Wead capsule very dark brown, almost pitch black, postclypeus and antennae, apart 
from two yellow-brown basal segments, dark sepia brown, labrum brown posteriorly shading to 
yellow at tip. Pronotum as head capsule, meso- and metanota and all other thoracic sclerites 
and legs sepia-brown, except tarsi and tips of tibiae, which are yellow to yellow-brown. Abdom- 
inal sclerites sepia-brown except anterior abdominal sternites paler brown in mid-line. Wings 
translucent brown, darker at base, slightly greenish iridescent ; venation sepia brown or 
outlined sepia brown towards wing base. 

Posterior margin of head almost semi-circular behind eyes ; fontanelle very small, short 
Y-shaped, scarcely to distinctly paler than rest of head, almost obsolete in some males ; eyes 
angularly ovoid, not prominent, and small relative to head width, W/E index 3:5-3°8 ; ocelli 
small, oval, separated from eyes by 14 to 2 times own least diameter ; postclypeus strongly 
inflated, width 1-9—2-0 times length, anterior margin nearly straight, posterior margin convex, 
rounded ; antennae 14-15 segmented, usually 14, II, V and VI subequal, longer than IV, III 
twice IV, often partly subdivided ; right mandible with posterior cutting edge of first marginal 
exposed beyond second, longer than that of second marginal, molar plate moderately indented 
near base, with 8 ridges. 


ETHIOPIAN NASUTITERMITINAE 59 


126. 


149. 152. 153. 


Fics. 119-156: Soldier head capsule, side and plan views, and nose tip. 119-126, 
Leptomyxotermes doriae ; 127-129, Grallatotermes africanus ; 130-134, Fulleritermes 
coatont ; 135-139, F. contractus; 140-145, F. mallyi; 146-153, F. tenebricus ; 
154-156, Rhadinotermes coarctatus. 


60 Wr A SANDS 


Cerci of 2 as long as width across base, slightly mamilliform conical. 

Pilosity of vertex, one or two long bristle-like setae and sparsely scattered shorter hair-like 
setae, with a few more bristle-like setae around eyes and ocelli (Text-fig. 114). 

Other characters given in generic discussion. 

Measurements (11 specimens from 5 localities) in millimetres. 


Range Mean 
Head width across eyes I-I2-1-19 : : I-16 
Greatest diameter of eye 0-31-0°33 : ; 0:32 
Ocellus F ‘ ; 0:06-0:09 X O:0Q-O-II . ; 0:08 X O-IO 
Ocellus toeye . : O-13—O-15 : : O13 
Width of pronotum 1-02-I-17 ; : 1:08 
Length of pronotum . 0-71-0-79 : : 0°74 
Length of hind tibia. 1°38-1°45 2 : I-41 
Length of fore wing. 8:20-9:60 ; : 8-99 


Soldievs. Major soldier : head capsule orange to ferruginous, nose ferruginous orange to 
pitch black with paler tip. Pronotum, antennae and abdominal tergites brown to sepia-brown, 
femora of legs brown, tibiae and tarsi yellow. Abdominal sternites pale brown. 

Head capsule in plan view constricted behind antennae, posterior margin regularly rounded ; 
nose conical. In profile straight or slightly concave to behind antennal socket, then abruptly 
raised and rounded ; nose shorter than rest of head capsule, measuring from hind margin of 
antennal socket, angle of nose cone 15—20 degrees ; antennae 13 segmented, II and IV subequal, 
shorter than V, III and VI subequal and longer than V. Head setae consist of 6-8 at nose tip, 
and a bilaterally placed pair of small setae on vertex, with very few scattered minute setae on 
dorsal surface. 

Abdominal tergites with very sparse scattered minute setae, slightly longer on posterior 2-3 
segments ; sternites with sparse short setae and longer setae up to 3 times length of short on 
posterior margins. Minor soldier : colour and pilosity, exactly as in major soldier. Head 
capsule in plan view more sharply constricted behind antennae. In profile concave, slightly 
sinuate, or indented in front of smaller posterior lobe of head ; nose shorter to equal in length to 
rest of head capsule, measuring from hind margin of.antennal socket, angle of nose cone 10-17 
degrees ; antennae 12-13 segmented, usually 13. 

Measurements (Major and minor soldiers 14 specimens each, from 8 localities) in millimetres. 


Major soldiers Range Mean 
Head length to tip of nose. 1-43-1-61 : ; : : 1°52 
Head width . 4 : 4 0:78-0:89 . - é : 0-80 
Depth of head capsule . : 0+54—-0°66 : : : : 0:58 
Width of pronotum : : 0-41-0'49 : ° : : 0°43 
Length of pronotum : : 0:18-0:23 : : : : 0-19 
Length of hind tibia : : I-OO-I°17 : : 5 : I-08 

Minor soldiers Range Mean 
Head length to tip of nose. 1 -28-1-44 : 2 3 : r-33 
Head width . : j ‘ 0:56-0:66 5 ‘ : : 0:61 
Depth of head capsule . : 0:41-0°49 ‘ : : . 0°45 
Width of pronotum ; : 0-33-0°37 : F : j 0°35 
Length of pronotum : ¢ 0-13-0'18 2 : ; : O'15 
Length of hind tibia 5 3 0:82-1:02 : : c : 0-94 


Variation. Like F. mally1, this species has proved to have relatively little variation over its 
comparatively short geographical range. The most variable caste is the minor soldier. 


This species is retained in Fudlleritermes in spite of its distinct soldier dimorphism 
because of its very close similarity in all respects, morphological and ecological, to 
F.. tenebricus and F’. contractus, with which latter species it has in the past been 


ETHIOPIAN NASUTITERMITINAE 61 


confused. It is separable from both by the differences of the right imago mandible, 
and from F. contractus by the different pilosity of the vertex. The more rounded 
posterior lobe of the soldier head distinguishes F’. coatoni from I’. contractus, and the 
points of the vestigial soldier mandibles are often though not always more pronoun- 
ced. The distinguishing features of F. mally: are discussed under that species. 


Holotype 2 winged imago, allotype 3, REPUBLIC OF SOUTH AFRICA : Transvaal, 
Sibasa, 24.x.1959 (W. G. H. Coaton), Coll. No. T.M. 6716, N.C.I., Pretoria. 


Paratypes, soldiers and workers, same data as holotype, in N.C.I., Pretoria and 
B.M.(N.H.). 

Other paratype material. SOUTHERN RHODESIA : Bulawayo, 13.v.1915 (Arnold), 
N.C.I., Pretoria. 

REPUBLIC OF SOUTH AFRICA: Transvaal; Bronkorstspruit, 6—12.v.1938, 
23-24.1X.1957, 26.vil.1960, 10 vials, Warmbad, 12.iv.1938, 25.ix.1957, 3 vials, 
Groblersdahl, 21.11.1939, 26.ix.1957, 2 vials, Pietersburg, 25.v.1939, 5.x.1960, 
Witbank, 30.vili.1959, Schweizer Reneke, 3.x.1956, 2 vials, N.C.I., Pretoria, 
Potgietersrus, 26.1x.1957, 2 vials, Pretoria, 23—25.1x.1957, 3.x.1960, 5 vials, Sibasa, 
8.vil.1960, 3 vials, Waterberg, 3.x.1960, 2 vials, Rustenburg, 1 & 26.ix.1961, 2 vials, 
Marico, 1.ix.1g61 (W. G. H. Coaton) ; Pretoria, 12.xi.1957, Sibasa, 30.xi1.1959, 
Waterberg, 6.1.1960, Rustenburg, 6-8.i.1960 (P. C. Joubert) ; Sibasa, 15.x.1960, 
2 vials, Pietersberg, 2.vili.rg61 (J. L. Sheasby); Soutpansberg, 24.1v.1958 (J. H. 
Grobler) ; Cape Province ; Vryburg, 25.11.1961, Mafeking, 26.11.1962, N.C.I., Pretoria 
(W.G. H. Coaton) ; Kimberly, 7.xi.1960 (P. C. Joubert) ; Natal ; Entonjaneni, 
13.1.1962, 2 vials (J. L. Sheasby). 

A total of 56 nest series of this species were examined, and material is in the 
British Museum (Natural History) unless otherwise stated. 


F.. coatom has been recorded as inhabiting parts of the mounds of various other 
genera. It appears to be confined to the relatively drier woodlands and wooded 
steppes included in vegetation type 20 of Keay ef al. (1959), occurring in Southern 
Rhodesia and the northern parts of South Africa. Several localities in Cape Province 
are beyond the mapped limit of type 20 vegetation. Coaton (1962) considers that 
the species does not occur west of Longitude 24°E., and it therefore seems probable 
that F. coatoni does not extend far into the even drier Acacia-Commiphora wooded 
steppe of type 25. These localities are thus likely to be in the transition zone, or 
even outliers of type 20 too small to be recorded on the map. 


Fulleritermes contractus (Sj6stedt) 
(Text-figs. 110, III, I15, 135-139, 172-174, 220 ; Map 15) 
Eutermes contractus SjOstedt, 1913a : 384. Type locality : Conco, Katanga, Welgelegen. 
Coarctatotermes contractus (Sj6stedt) Fuller, 1922: 117. [unjustified emendation of generic 
name Coarctotermes Holmgren). 
Coarctotermes contractus (Sjostedt) ; Sjdstedt, 1926 : 356. 
Coarctotermes brunneus Noirot, 1955 : 150. Type locality : ANGoLa, Humpata, Leba. syn. n. 
Coarctotermes brunneus Noirot ; Sands, 1957 : 14. 
Fulleritermes contractus (Sj6stedt) Coaton, 1962 : 152. 
Fulleritermes brunneus (Noirot) Coaton, 1962 : 153. 


62 Wie eA ANID S 


Imago. Head capsule pitch black, postclypeus very dark sepia brown, labrum sepia brown 
with hyaline tip ; antennae sepia brown. Pronotum very dark sepia brown, meso- and meta- 
nota, other thoracic sclerites and legs (apart from tarsi) sepia brown, tarsi paler brown. Abdom- 
inal tergites sepia brown, sternites paler in mid-line. Wings brown, darker at base, green and 
purple iridescent, venation sepia brown, outlined near base. 

Posterior margin of head behind eyes semi-circular ; fontanelle small clear-cut Y-shaped, 
distinctly paler than head, in profile with small raised bump in front of fork ; eyes angular 


er ee, ees en ae i 
— Kors 


Map 14. Fulleritermes coaton. Vegetation type: 20. 


ETHIOPIAN NASUTITERMITINAE 63 


oval, not prominent, small relative to head width, W/E index 3-4-3-5 ; ocelli small, oval, 
separated from eyes by about twice own least diameter ; postclypeus strongly inflated, width 
1-8-2-1 times length, anterior margin slightly concave, posterior margin convex, rounded ; 
antennae 14 segmented, II, V and VI subequal, slightly longer than LV, III twice IV, sometimes 
partly subdivided ; right mandible with posterior cutting edge of first marginal tooth exposed 
beyond second, shorter than that of second marginal, molar plate deeply indented near base, 
with 8 ridges. 

Cerci of 2 shorter than width across base, slightly mamilliform conical. 

Pilosity of vertex of rather coarse long yellow setae scattered among shorter setae, slightly 
less dense on ‘“‘crown’’. (Text-fig. 115.) 

Other characters mentioned in generic diagnosis. 

Measurements (3 specimens from 2 localities) in millimetres. 


Range 

Head width across eyes , 3 - é 0-94-I°10 
Greatest diameter ofeye  . : : : 0:28-0°31 
Ocellus . P “ - , : : 0:06-0:08 * 0:09 
Ocellus toeye . é : : : : O-13-0'14 
Width of pronotum . : : : . 0:92-1:05 
Length of pronotum . , : : ; 0-64-0-79 
Length of hind tibia . : : ; : I-15—-1°43 
Length of fore wing . : ; : : 7:20 


Soldier. Head capsule ferruginous orange to sepia brown, nose ferruginous to pitch black 
with reddish tip. Pronotum, antennae, abdominal tergites and femora of legs sepia brown, 
abdominal sternites and tibiae and tarsi of legs paler, brown. 

Head capsule in plan view, strongly constricted behind antennae, posterior margin usually 
somewhat angularly rounded and often sulcate in the middle to a varying degree ; nose moder- 
ately conical. In profile, dorsal surface of nose straight to well behind antennal socket, or 
slightly sinuate with small hump at base of nose, with or without slight indentation in front of 
raised and rounded posterior lobe of head ; nose shorter, to equal in length to rest of head 
capsule measuring from hind margin of antennal socket, angle of nose cone 10-20 degrees ; 
antennae 12-13 segmented, usually 13, II and V subequal, slightly longer than IV and shorter 
than III. Head setae confined to 6-8 at nose tip, and one bilaterally placed pair on vertex. 

Abdominal tergites with very sparse, minute scattered setae, slightly longer upon last two ; 
sternites with sparse short setae and longer setae on posterior margins up to twice length of 
short. 

Measurements (30 specimens from 7 localities) in millimetres. 


Range Mean 
Head length to tip of nose. I-I14—I-50 - : : ; 1-39 
Head width . . : : 0:54-0°83 F - : : 0:73 
Depth of head capsule . : 0*40-0°55 : : é : 0-50 
Width of pronotum : ; 0+32-0°45 : : : ; o-4I 
Length of pronotum : : 0-15-0'18 ; F : : 0-16 
Length of hind tibia ‘ , 0:68-1:05 ; 0:93 


Variation. Imago variation cannot be assessed. The soldier varies considerably in colour 
and proportions, though slightly less than in F. tenebricus. Angularity of the head capsule and 
the sulcate posterior margin appear to be commoner at the eastern end of the range, but material 
from the west is meagre. 


The soldiers of F’. contractus are very difficult to separate from F. tenebricus, 
having a slightly thicker nose on average, slightly less pronounced posterior lobe 
of the head capsule, and proportionately shorter hind tibia. The ranges of all these 
characters overlap however. The imagos of F. contractus are distinguishable by 


64 W. A. SANDS 


the more hairy vertex, the small bump or tubercle in front of the fontanelle, and the 
wider separation of ocelli from eyes. The distinguishing features from F. coatont 
are discussed under that species. 


A lectotype has been selected from among the syntype series of F’. contractus, and 
labelled as such. 


a = pos ee 


SS 


? 


Map 15. Fulleritermes contractus. Vegetation types: §8, 16, 18, and 19. 


ETHIOPIAN NASUTITERMITINAE 65 


Lectotype : Conco: Katanga, Welgelegen, 14.vi.1912 (J. Bequaert), lectotype 
queen, paralectotype soldiers and workers, Mus. Tervuren. Other paralectotype 
soldiers, type colony, A.M.N.H. and B.M.(N.H.). 


Type Material. ANGoLA : Humpata, Leba, 17.1x.1949 (A. de Barros Machado), 
paratype soldier, type colony, F. brunneus (Noirot), A.M.N.H. 


Other Material. Conco: Leopoldville, Kitona, 9.viii.1g59 (J. Ruelle), N.C.L., 
Pretoria ; Katanga, Keyberg, 11° 47’S., 27 ° 25’E., 19-25.iv.1948, 2 vials (A. E. 
Emerson), A.M.N.H. 


ANGOLA : Munhango, 24.x.1927 (M. Burr). 

NORTHERN RHODESIA : Abercorn, iii.1947 (P. E. Glover) ; Lusaka, xi.1963 (E. N. 
Cooling). 

SOUTHERN RHODESIA : Salisbury, 1911 (G. A. K. Marshall). The last 4 nest 
series in British Museum (Natural History). 9 nest series were examined. 


The habits of this species seem to be similar to those of F’. tenebricus in the northern 
savannahs, sharing the mounds of other species and occasionally constructing its 
own small mound, or carton nest around a branch stump on a tree. It occurs in 
the savannah zones south of the Congo forest block, comprising vegetation types 
8, 16, 18, and 19 of Keay et al. (1959), thus having a somewhat more restricted 
distribution than its northern congener. 


Fulleritermes mallyi (Fuller) 
(Text-figs. 106, 107, 116, 140-145, 175-177, 221, 222 ; Map 16) 


Tenuirostritermes mallyi Fuller, 1922: 115. Type locality : REPUBLIC OF SouTH AFRICA, 
Cape Province, Malmesbury. 

Eutermes mallyi (Fuller) Sjéstedt, 1926 : 310. 

Coarctotermes mallyi (Fuller) Snyder, 1949 : 311. 

Fulleritermes maillyi (Fuller) Coaton, 1962 : 152. 


The following additions to Fuller’s descriptions are necessary : 

Imago. Posterior margin of head almost semi-circular behind eyes ; eyes small relative to 
head width, W/E index 3:8—4-4, moderately prominent in proportion to own diameter ; ocelli 
small, separated from eyes by slightly more than to nearly twice own least diameter ; post- 
clypeus very strongly inflated, width 1-7-1-8 times length, anterior margin almost straight, 
posterior regularly convex ; posterior cutting edge of first marginal tooth of right mandible 
exposed beyond second, longer than that of second marginal. 

Cerci of 9 shorter than width across base, mamilliform. 

Pilosity of vertex densely scattered rather long yellow bristle-like setae of varying length, rest 
of body with yellow hair- and bristle-like setae. Other characteristics described in generic 
diagnosis. 

Measurements (24 specimens from 7 localities) in millimetres. 


Range Mean 
Head width across eyes 1°37-1°43 : . I-40 
Greatest diameter of eye 0:31-0:36 : : 0°34 
Ocellus : : : 0:09-0'IO X O:10-O'16 : 0:09 X 0°13 


Ocellus toeye  . : 0-10-0'19 : : 0:16 


66 W. A. SANDS 


Range Mean 
Width of pronotum . 1-:22-1-40 ; ‘ I-32 
Length of pronotum . 0:71-0:84 ; 5 0:80 
Length of hind tibia. I -63-1-°79 : : 7 
Length of fore wing . I1-50—14°30 : : 12°71 


Soldiey. Setae on head consist of a bilaterally placed pair on vertex, four at nose tip with 
sparse shorter setae behind them, and one or two towards base of nose. 

Abdominal tergites with very short sparsely scattered setae and longer setae on posterior 
margins ; sternites with more numerous short setae and longer setae on posterior margins 
2-3 times length of short. 

Antennae, 12 (6 out of 10) to 13 segmented. 

Measurements (53 specimens from 16 localities) in millimetres. 


Range Mean 
Head length to tip of nose. 1-43-1-86 : : ; : 1°59 
Head width . : : : 0°64-0:94 : : : : 0-82 
Depth of head capsule . : 0-43-0°64 : : : : 0°57 
Width of pronotum ; : 0-38—-0'51 : : ; : 0-44 
Length of pronotum : E 0:18-0:28 : : : : 0:23 
Length of hind tibia : ; 0:89-1:30 Lk2 


Variation. The 5% size variation shown by the i “Sa ete over ae saue disedpacen range is 
extremely low, and other characters varied similarly little. In the soldier there was more 
variation in the proportional lengths of nose and head capsule and in antennal length, with slight 
indications of a cline in these characters from south-west to north-east. Colour varied little 
in both castes. 

Though placed in Fulleritermes, the relationships of F. mallyi are open to doubt. 
The long wings and 15 segmented antennae of the imago together with the prepon- 
derantly 12 segmented antennae of the soldier, show closer resemblance to R. 
coarctatus than to the other species of its own genus. 

These characters, the slender cylindrical nose and comparatively weak head 
constriction of the soldier, and the larger size of both imago and soldier castes, 
distinguish it from other Fulleritermes and R. coarctatus. It is unlikely to be con- 
fused with any other African species. 


Material. REPUBLIC OF SOUTH AFRICA: Cape Province, Malmesbury, x.1920 
(R. Birch), syntype female imagos and soldiers, type colony, F. mally: (Fuller), 
A.M.N.H. (other syntypes in N.C.I., Pretoria) ; Malmesbury, 19.ii.1956, Clan 
William, 21.i11.1956-20.11.1961, 7 vials, Calvinia, 6.x.1956—15.i.1958, 6 vials, 
Willowmore, 18.ix.1961, 4 vials, Beaufort West, 19.ix.1961, Aberdeen, 20.1x.1961, 
3 vials (W. G. H. Coaton) ; Laingsburg, 15.xi.1960, 4 vials, Ladismith, 16.xi.1960 
(P. C. Joubert, H. P. Nieman) ; Uniondale, 12.11.1961, Ceres, 19.11.1961, Clanwilliam 
and Calvinia, 20.11.1961, Namaqualand, I5-19.x.1961, 17 vials, Van Rhynsdorp, 
20.X.1961, 5 vials (J. L. Sheasby, G. F. Pretorius). 

A total of 54 nest series were examined, and material is in the British Museum 
(Natural History) unless otherwise stated. 


This species appears to be confined to the Karroo and related neighbouring vege- 
tation types of Cape Province, South Africa. It has commonly been collected from 
beneath stones and from mounds of other species such as Amutermes hastatus 
(Haviland). 


ETHIOPIAN NASUTITERMITINAE 67 


ALL OF Pm 


Map 16. Fulleritermes mallyi. Vegetation types : 28 and 29. 


68 W. A. SANDS 


Fulleritermes tenebricus (Silvestri) 
(Text-figs. 108, 109, 118, 146-153, 178-180, 223, 224 ; Map 17) 


Eutermes tenebricus Silvestri, 1914 : 44. Type locality : Guinea, Kakoulima. 

Eutermes (Coarctotermes) tenebricus Silvestri ; Hegh, 1922 : 469. 

Coarctotermes tenebricus (Silvestri) Sj6stedt, 1926 : 355. 

Nasutitermes (Coarctotermes) suffuscus Emerson, 1928: 496. Type locality : Conco, near 
Yakuluku (4° 20’ N., 28° 50’ E.) syn. n. 

Coarctotermes suffuscus (Emerson) Snyder, 1949 : 3IT. 


Imago. Head capsule dark sepia brown, slightly paler anteriorly ; pale patches in front of 
ocelli and paler coronal suture sometimes present ; postclypeus sepia brown, labrum yellow to 
brown, antennae sepia brown. Pronotum as head capsule, meso- and metanota and other 
thoracic sclerites sepia brown. Femora of legs sepia brown, tibiae brown with yellow tips, tarsi 
yellow. Abdominal tergites uniformly sepia brown, sternites paler in mid-line. Wing brown, 
darker at base, translucent when wet, bluish iridescent when dry, venation outlined sepia brown 
at base, becoming indistinct distally. 

Posterior margin of head behind eyes almost semi-circular, slightly sinuate immediately 
behind eyes ; fontanelle elongate Y-shaped, slightly to distinctly paler than head ; eyes 
angularly ovoid, not prominent, small relative to head width, W/E index 3-2-3-6 ; ocelli small, 
oval, separated from eyes by about 14 times own least diameter ; postclypeus strongly inflated, 
width 1-8—2-1 times length, anterior margin slightly concave, posterior, regularly convex ; 
antennae 14 segmented, II, V, and VI subequal, IV slightly shorter than these, III longer, up to 
twice IV, sometimes partially subdivided ; right mandible with posterior cutting edge of first 
marginal exposed beyond second, shorter than that of second marginal, molar plate deeply 
indented near base, with 8 ridges. 

Cerci of 2 slightly mamilliform conical shorter to equal in length to width across base. 

Pilosity of vertex, one or two long bristle-like setae and sparsely scattered shorter hair-like 
setae, with a few more bristle-like setae around eyes and ocelli. (Text-fig. 118.) 

Other characters given in generic diagnosis and discussion. 

Measurements (18 specimens from 6 localities) in millimetres. 


Range Mean 
Head width across eyes O-QI-1:22 : : 1-06 
Greatest diameter of eye 0:29-0°34 ‘ : 0:31 
Ocellus : j : 0:06-0:09 X O-IO-O'I2. : 0:08 X OvlI 
Ocellus toeye  . : O-10—-0'14 2 : O-12 
Width of pronotum . 0:88-1:23 . : I-09 
Length of pronotum . 0-60-0°88 c : 0°75 
Length of hind tibia . I+25-1°54 ; : I*41 
Length of fore wing . 6-10—9-00 : : 7°42 


Soldier. ead capsule ferruginous orange, slightly darker posteriorly, to dark chestnut 
brown, nearly black posteriorly, nose ferruginous to pitch black with reddish tip. Pronotum, 
antennae, and abdominal tergites brown, femora of legs pale brown, tibiae and tarsi yellow. 
Abdominal sternites yellow-brown. 

Head capsule in plan view strongly constricted behind antennae, posterior margin regularly 
to somewhat angularly rounded, often sulcate in the middle to varying degree ; nose near 
cylindrical to moderately conical. In profile dorsal surface of nose straight to well behind 
antennal socket, or slightly sinuate with small hump at base of nose, with or without slight 
indentation in front of raised and rounded posterior lobe of head ; nose shorter to slightly longer 
than rest of head capsule measuring from hind margin of antennal socket, angle of nose cone 
5-17 degrees ; antennae 13 segmented, II short, IV, V, and VI progressively slightly longer, 
III longer up to twice II. Head setae confined to 6-8 at nose tip, one bilaterally placed pair on 
vertex and sometimes another at base of nose. 


ETHIOPIAN NASUTITERMITINAE 69 


Abdominal tergites with very sparse scattered minute hair-like setae slightly longer on pos- 
terior margins ; sternites with denser short setae and longer setae up to 3 times length of short 
on posterior margins. 

Measurements (30 specimens from 19 localities) in millimetres. 


Range Mean 
Head length to tip of nose. I-I5-1°50 : : : ; 1-36 
Head width . : . : 0:59-0:86 > - 2 ‘ 0°75 
Depth of head capsule . : 0°43-0-61 ; ; ; ‘ 0°54 
Width of pronotum ; ; 0:38-0:47 ? ‘ : : 0°43 
Length of pronotum - : O-°15-0-21 ‘ - , : O'17 
Length of hind tibia : 0°94-1°23 1-06 


Variation. In the imago the vasiablen was chiefly in size with eraportionis fairly constant. 
The soldiers have proved highly variable in size, colour, and head profile, but the range has been 
found indivisible. Angularity of the posterior margin of the head is commoner at the western 
end of the distribution range. 

The relationship of this species to F’. coatoni and I’. contractus are mentioned under 
those species. 

Type Material. GuiINEA: Kakoulima (9° 40’N., 13° 25’ W.), 12.vill.1g12 (FP. 
Silvestri), syntype soldiers, type colony, F. tenebricus (Silvestri), Silvestri Coll., Lab. 
Zool. Sc. Agr., Portici, and A.M.N.H. 

Conco : near Yakuluku (4° 20’N., 28° 50’ E.), xii.rg11 (H. Lang, J. P. Chapin) 
paratype soldier, type colony, F. suffuscus (Emerson), A.M.N.H. 


Other material. GUINEA : Nimba Mountains, 1957, 10 vials (M. Lamotte), Inst. 
Franc. d’Afr. Noire, Dakar. 

GHANA : Yeji, 10.xii.1926 (A. W. J. Pomeroy) ; 60m. N. of Ejura on Tamale 
Road, 24.11., 12m. S. of Tamale on Kumasi Road, 26.ii., 4m. from Daboya on Tamale 
Road, 28.11., 55m. from Tamale on Larabanga Road, 3.111., 415m. from Gambaga on 
Walewale Road, 5-7.iii., 25m. S.E. of Gambaga on Yendi Road, 14m. from Gambaga 
on Bawku Road, 6.iii., 20m. from Bolgatanga on Tamale Road, 9.iii., tom. N. of 
Bawku, I1.iii., 11m. from Tumu on Navrongo Road, 15.iii., 4om. from Tumu on 
Lawra Road, 16.iii., Lawra and gm. N. of Lawra on Nandom Road 18.iii., 6m. N. of 
Wa on Lawra Road, 19.iii., 24-52m. S. of Wa on Bole Road, 20.iii., 3-23m. from 
Larabanga on Bole Road, 21-22.iii., 6m. N. of Wangasi-Turu on Tamale Road, 
27.111., 7m. from Nkoranza on Ejura Road, 28.iii., and 25m. N. of Wenchi on Bamboi 
Road, 30.iii.1959 (W. A. Sands). 

NIGERIA : Western Region ; Olokemeji, xi.1g12 (F. Silvestri), ALM.N.H. Eastern 
Region ; Onitsha, 20.i. and 5.ix.1957 (W. Wilkinson) ; 7-12m. from Enugu on 
Onitsha Road, 2-3.ili.1958 (W. A. Sands). Northern Region : 20m. from Zaria on 
Funtua Road and 30m. from Funtua on Yashi Road, 14.xi., 25m. from Katsina on 
Daura Road, 19.xi., 71m. from Gusau on Sokoto Road, 22.xi., 2 vials, 45m. from 
Sokoto on Argungu Road, 23.x1., Zaria, 4.xii., 33m. from Kaduna on Zungeru Road, 
20.x11., 16m. S. of Zungeru on Bida Road, 21.xii., Bida, 23.xii., 39m. from Abuja on 
Bida Road, 24.xii., and Gwagwa Forest near Abuja 28.xii.1956, 5m. from Jos on 
Bukuru Road, 6.ii., 42-7m. from Jos on Wamba Road, 8.ii., 40m. from Jos on 
Bauchi Road 11.ii., Bauchi, and 25m. from Bauchi on Jos Road, 12-13.1i., 18, 20, 
and 29m. from Jos on Pankshin Road, 14.ii., 3.iv., and 25.vii., 16m. from Kaduna 


70 W. A. SANDS 


on Zungeru Road, 15.ii., 39m. from Bauchi on Gombe Road 6m. from Gombe on 
Hinna Road, 9.v., Tula, 37m. S.E. of Gombe, Io.v., 2 vials, 1om. S. of Mayo Faran, 
14.v., 34-54m. from Yola on Jalingo Road, 14-16.v., 60m. S.W. of Jalingo on Takum 
Road 19.v., 15m. N.N.W. of Tibak, Schebschi Mountains, 24.v., 80m. from Yola on 
Biu Road, 28.v., 32m. from Damaturu on Potiskum Road, 5.vi., and 5m. S. of 
Yashi on Funtua Road, 3.x.1957, 24m. from Gboko on Makurdi Road, 25.ii., and 
22m. from Kabba on Ikole Road, 11.11.1958 (W. A. Sands). 


ae a = ea we 
ie VS 


Sg FZ. 


nay, 
° 
5 
a 


Map 17. Fulleritermes tenebricus. Vegetation types: 8, 16, 17, and 20. 


ETHIOPIAN NASUTITERMITINAE 71 


CENTRAL AFRICAN REPUBLIC: Haut Mbomu, Zemio, 11.1948 (N. A. Weber), 
A.M.N.H. 

Conco: Niangara, 20.v.1913 (H. Lang, J. P. Chapin) ; Dika,* 26.iii.19g25 (H. 
Schouteden), A.M.N.H.; Tora, 26.ix.1926, 2 vials (H. Burgeon) ; Garamba Nat. 
Park, 6.vii.1951 (Miss. de Seager), Inst. des Parcs Nat. du Congo, Brussels. 

SuDAN : Delami, 20.v.1927 (W. Ruttledge). 

A total of 86 nest series were examined and material is in the British Museum 
(Natural History) except where otherwise stated. 


F. tenebricus is a species occurring in the Guinean savannah vegetation zones from 
the forest edge to the relatively dry Sudan type (Keay et al., 1959, vegetation types 
8, 16, 17, and 20). In this respect its distribution north of the Congo forest is as 
wide as the combined distributions of its two congeners F. coatoni and F. contractus 
in similar vegetation zones to the south of the forest block. It sometimes constructs 
a small hard earth mound with dark carton interior but is more commonly found 
inhabiting parts of mounds constructed by other genera such as Cubitermes, Trinervt- 
termes and Macrotermes. It is also found under stones, in dead branches on trees, 
and in narrow earth runways which it constructs in foraging. 


RHADINOTERMES ¢en. n. 
(Rhadinos, Gr., “ tapering ’’) 
Type-species, Eutermes coarctatus Sjéstedt, 1902a : 304. 


Imago. Left mandible with apical tooth as long as first marginal or slightly shorter, distance 
from apical to first marginal about one-third distance between first and third marginals (left 
mandible index -32—:36), second marginal obsolete, cutting edge between first and third mar- 
ginals entire ; right mandible with apical and first marginals subequal as left, first marginal 
with small subsidiary ‘‘ tooth ’’ near basal junction with apical, and with posterior cutting edge 
exposed beyond anterior edge of second marginal longer than posterior cutting edge of latter ; 
right molar plate in surface view with weak indentation near basal end of anterior margin, 
posterior margin extending in rounded flange, first molar ridge forming a stout tooth, 6-7 
remaining ridges diminishing towards base (Text-figs. 182, 183). Fontanelle, very narrow slit- 
like, scarcely bifurcate anteriorly in g, more clearly in Q, slightly to distinctly paler than head. 

Soldier. Monomorphic. Nose conical. Vestigial mandibles with small points. Head 
capsule sharply constricted behind antennae, posterior lobe commonly only about one-fourth 
wider than anterior. Antennae 12 segmented. 

This genus is proposed to accommodate the single African mainland species 
coarctatus Sjostedt, hitherto included, because of similarities in the soldier castes, in 
the Malagasy genus Coarctotermes of which the type-species is Eutermes clepsydra 
Sjéstedt, 1904 : 108. The imago differs from Rhadinotermes coarctatus in a number 
of features. The left mandible index is -30, and the third marginal tooth in fresh 
a Pa a on the locality Dika, recorded by Emerson (1928) as 7° 50’ N., 18° 45’ E., i.e. near Ft. Crampel, 

° r. Kep. 
Other localities Otitbisimamerenstiat 42 25° IN. 20° an’ B.s5° sof N. 15° 13’ E.;.5° 33’ N., 20° oo E., 


in C. Afr. Rep., at 4° 36’ N., 28° 36’ E. in Sudan, and in Congo at 4° 17’ N., 27° 42’ E., and 4° o9’N., 
27° 40’ E., one of these last probably being that known to H. Schouteden. 


72 W.A. SANDS 


unworn specimens has been completely absorbed in the cutting edge joining it to the 
first marginal. The fontanelle is a rounded equilateral triangle with long side arms. 
It is a very much larger insect, having a head width across eyes of approximately 
2mm., and much more prominent eyes and ocelli. The posterior margin of the 
pronotum is deeply indented. In its general appearance and superficially in its 
ecology, R. coarctatus closely resembles the genus Fulleritermes Coaton, 1962 : 
151-4 from which it has recently been separated on the basis of the presence in the 
latter of a notch in the cutting edge in front of the third marginal tooth of the left 
mandible. This is discussed further under that genus. Other differences such as 
the greater proportionate length of this cutting edge and also that of the right first 
marginal in Rhadinotermes, and the much less prominent molar regions indicate that 
this genus is probably adapted to a somewhat different ecological niche, especially 
since it has been recorded from the same localities as members of Fulleritermes. 

It is convenient to include R. coarctatus in the keys to Fulleritermes. The account 
of distribution follows the specific description. 


Rhadinotermes coarctatus (Sj6stedt) comb. n. 
(Text-figs. 112, 113, 117, 154-156, 181-183, 225, 226 ; Map 18) 


Eutermes coarctatus Sjostedt, 1092a : 304. Type locality : NyasaLanp, Zomba. 

Eutermes (Coarctotermes) coarctatus Sjostedt ; Holmgren, 1912 : 65. 

Coarctatotermes coarctatus (Sjostedt) Fuller 1922: 118. [Unjustified emendation of generic 
name. | 

Coarctotermes coarctatus (SjOstedt) Sjostedt, 1926 : 355. 

Coarctotermes coarctatus (SjOstedt) ; Sands, 1957 : 14. 


Imago. Wead capsule very dark brown to pitch black, postclypeus and antennae dark sepia 
brown. Labrum sepia brown, shading to yellow at tip. Pronotum as head capsule, meso- and 
metanota, pleural and ventral thoracic sclerites dark sepia brown. Legs dark sepia brown apart 
from paler tips of tibiae and yellow tarsi. Abdominal tergites dark sepia brown, outer ends 
paler, sternites sepia brown, pale brown in mid-line. Wings translucent pale brown, venation 
yellow-brown, radius sector with brown streak along posterior margin. 

Posterior margin of head almost semi-circular behind eyes ; eyes not prominent, small relative 
to head width W/E index 3-4-3-8, somewhat angularly ovoid ; ocelli small, oval, separated 
from eyes by about 1} times own least diameter ; postclypeus strongly inflated, width 2-2 
times length, anterior margin slightly concave, posterior, indistinct, convex ; antennae 15 
segmented, II, IV, and V subequal in length, II thinner, III longer than these. 

Cerci of 2 shorter than width across base, mamilliform. 

Pilosity of vertex scattered rather long yellow bristle-like setae rest of body with pale yellow 
hair- and bristle-like setae. Other characteristics described in generic diagnosis. 

Measurements of 8 specimens from 3 localities were given in an earlier paper (Sands, 1957), 
these being all the material known apart from one nest series described by Fuller (1922) and 
falling within this size range. 

Soldier. ead capsule sepia brown to pitch black, nose pitch black with reddish tip. 
Pronotum, antennae and abdominal tergites sepia brown. Legs, femora brown, tibiae and tarsi 
yellow to pale brown. Abdominal sternites yellow-brown to brown. 

Head profile raised and rounded behind constriction, more or less straight in front. Nose 
equal in length or shorter than rest of head capsule, measuring from hind margin of antennal 


ETHIOPIAN NASUTITERMITINAE 73 


socket, angle of nose cone 10-20 degrees ; antennae 12 segmented, II short, IV and V longer, 
and III subequal or slightly longer than these. Setae on head confined to four at nose tip and 
bilaterally placed pair on vertex, with a few short curved setae scattered on dorsal surface of 
head including nose. 

Abdominal tergites with very short sparsely scattered setae, and longer setae on posterior 
margins ; sternites with more numerous short setae and longer setae on posterior margins 2-3 
times length of short. Remaining characteristics given in generic diagnosis. 

Measurements (25 specimens from 16 localities) in millimetres. 


Range Mean 
Head length to tip of nose. I-17-1°39 : : : 2 1-28 
Head width . : : F 054-066 , ; : 3 0:60 
Depth of head capsule . - 0-40-0°49 . : - ; 0°45 
Width of pronotum : : 0-33-0°38 P ; ; . 0:36 
Length of pronotum : : 0-15-0'18 : , ; ; O17 
Length of hind tibia : 0-92-I-10 I-OL 


Variation. Imago variation was very slight in the small amount of waterial known. In the 
soldier caste small variations in length and angle of nose occur. 


The relationships of this species are discussed under the generic heading. It is 
unlikely to be confused with any species other than small dark specimens of Fudlert- 
termes contractus, and then only then in the case of the soldier caste collected alone. 


Material (additional to that recorded in Sands, 1957). NYASALAND : Zomba, 
1899 (Cameron), syntype soldiers and workers, R. coarctatus (Sjéstedt), A.M.N.H. 
(other syntypes in Mus. Cambridge, and Naturhist. Riksmus., Stockholm). 

NORTHERN RHODESIA : Choma, 16.1.1957, 2 vials, Magoye, 17.1.1957, Lusaka, 
19.i.1957, Broken Hill, 20.i.1957 (W. G. H. Coaton). 

ConGo : Keyberg, 11° 47’S., 27° 25’ E., 21.1v.1948 (A. E. Emerson), A.M.N.H. 

REPUBLIC OF SOUTH AFRICA: Transvaal; Brits, no date (H. K. Munro) ; 
Sibasa, 5.viii.1959 to 13.x.1960, 8 vials, Barberton, 23-27.iv.1960, 3 vials, Pietersberg, 
Lydenberg, 5.x.1960, 3 vials, Letaba, 8-16.x.1960, 5 vials, Nelspruit, 21.x.1g960, 
3 vials, and Soutpansberg, 10.xii.1g60 (W. G. H. Coaton, P. C. Joubert, and J. L. 
Sheasby) ; Pilgrims Rest, 3.ix.1960, 2 vials (A. Boonzaaier) ; Barberton, 7.x.1960 
(H.P. Nieman) ; Potgietersrus, 21.xii.1961 (P. le S. Milstein) ; Natal ; Ingwavuma, 
13.x11.1959 (J. Nel) ; Ngotshe, Nongoma, 10.i.1962, 5 vials, Mahlabatini, 12.1.62, 
2 vials (J. L. Sheasby). 

SWAZILAND : Hlatikulu, 24.x.1960, 3 vials (W. G. H. Coaton). 

A total of 56 nest series were examined and material is in the British Museum 
(Natural History) unless otherwise stated. 


R. coarctatus has been found in the nests of other species, under stones, and 
occasionally building a small mound structure of its own. It is restricted to eastern 
Africa, and its distribution appears to approximate most closely to the rather drier 
vegetation types coming between the Brachystegia-Julbernardia woodland and the 
Acacia-Commiphora steppes, grouped in Keay et al. (1959) under vegetation type 20. 
It also occurs in the coastal savannah-forest mosaic, type 9, which has similar 
rainfall to type 20. These vegetation types are more widely distributed in fact 
than can be shown on the map, as small areas fringing the other types around 
mountain blocks and plateaux, and in valleys. 


74 w. A. SANDS 


°c 1. mm mm 


Map 18. Rhadinoteymes coarctatus. Vegetation types: 9 and 20. 


ETHIOPIAN NASUTITERMITINAE 75 


MYCTEROTERMES €en. n. 


(Mycteros, Gr., ‘‘ beak ’’) 
Type-species, Mycterotermes meringocephalus sp. n. 


Imago. Unknown. 

Soldier. Dimorphic. Major soldier, nose short, cylindrical, vestigial mandibles with small 
but distinct points showing trace of marginal as well as apical tooth ; head capsule very weakly 
constricted behind antennae, vertex and nose with short but distinct bristle-like setae. Minor 
soldier, as major soldier, except nose weakly conical, head capsule more definitely constricted 
behind antennae. Antennae probably 13 segmented in major, 12 segmented in minor soldier. 

There is no doubt that these specimens belong to a new genus of some phylo- 
genetic importance, being close to but more primitive than 7vinervitermes, and 
possibly even ancestral to it. The distinguishing features given in the diagnosis 
have much in common with the Neotropical genus Diversitermes and it appears 
most probable that Tvinervitermes arose from this branch of the hypothetical 
phylogenetic tree in which Ahmad (1950) placed Velocitermes, Tenutrostritermes, and 
Ceylonitermes, and to which I (Sands, 1957) added Coarctotermes sensu lat. (Fulleri- 
termes, Rhadinotermes, and Coarctotermes s. str.). At least 3 of these genera have 
become adapted to grass feeding and two to grass storage convergently with 
Trinervitermes, occupying equivalent ecological niches in their respective zoogeo- 
graphical areas (Neotropical, Ethiopian, and Malagasy). Other genera of similar 
habits and appearance from the Australian region have not been studied in detail 
but may also be related. 

Mycterotermes is unlikely to be confused with any genus other than 7vinervitermes 
in the area covered by this revision. Its occurrence in Western Aden brings it into 
the Ethiopian Zoogeographical Region. 


Mycterotermes meringocephalus sp. n. 
(Text-figs. 388-395 ; Map 3) 


Imago. Unknown. 

Soldiers. Major; Head capsule orange-yellow, nose ferruginous. Antennae yellow, 
pronotum and abdominal tergites yellow-brown. Abdominal sternites and legs pale yellow. 

Head capsule in plan view tapering slightly towards front, sides slightly sinuate behind 
antennae, posteriorly weakly sulcate in mid-line, nose cylindrical to near base. In profile level 
from tip of nose to downward curve at back of head, with slight hump at base of nose and very 
weak constriction behind antennae ; nose distinctly shorter than rest of head capsule, measuring 
from hind margin of antennal socket. Antennae, broken, II and IV subequal and shorter than 
V, III nearly twice as long as II. 

Abdominal tergites with sparse fine minute setae, sternites with sparse shorter setae and longer 
setae on posterior margins. 

Minor ; as major soldier except antennae, II shorter than III, IV, and V which are subequal. 

Remaining characters given in generic diagnosis. 

Measurements (Unique holotype major soldier, 2 minor soldiers). 


Major soldier Minor soldiers 
Head length to tip of nose 1°63 : : é : 1-22 
Head width . : : 1-00 : j : : 0°54 


Depth of head capsule. 0-59 é : F é 0:38 


76 W. A. SANDS 


Major soldier Minor soldiers 
Width of pronotum c O51 c : ; E 0-41 
Length of pronotum : 0:23 P : : : 0-18 
Length of hind tibia : both broken . : : ; 0:94-1:00 


Since this is a monotypic genus, its relationships have already been discussed 
under the generic heading. There is a slight possibility that the imagos described 
as Trinervitermes saudienstis sp. n. from Jiddah belong to this species, in view of their 
unusually wide fontanelle and small eyes, but this cannot be settled without further 
collecting. 


Holotype, major soldier, ADEN PROTECTORATE : Burum, W. of Mukalla, 12.xii. 
1951 (W. V. Harms, coll. no. 796), in B.M.(N.H.). 


Morphotype and one paratype minor soldiers, same data as holotype, also in 
B.M.(N.H.). 

It is recorded as found with Tvinervitermes avabiae Harris in small loose sand 
heaps over narrow exit holes. 


TRINERVITERMES Holmgren 


Eutermes (Tvinervitermes) Holmgren, 1912 : 59 [Subgenus of Eutermes Heer]. 
Trinervitermes Fuller, 1921 : 19 [full generic status]. Type species, by subsequent designation 
(Sjéstedt, 1926 : 325), Tvinervitermes trinervius (Rambur, 1842). 


Holmgren (1912) on first creating the subgenus, Eutermes (Tvinervitermes) included 
a total of sixteen species, but did not designate the type-species. Fuller (1921) 
raised Tvinervitermes to full generic status, consequent on the substitution by Banks 
(1919) of the name Nasutitermes for Eutermes s. str. as used by Holmgren (1912), but 
again, did not designate a type-species. Termes trinervius Rambur, 1842 was 
designated by Sjdéstedt (1926) as type-species of the genus Tvinervitermes. This 
choice was unfortunate because the unique holotype of 7. tvinmervius (Ramb.) is a 
pinned dried g, from which the head capsule has been removed or lost, and the 
species is the most confused in the genus as a result of subsequent misidentifications. 
There is however no doubt of its correct assignment to the genus, in spite of its 
somewhat uncertain specific identity. The usage of Tvinervitermes as a genus or 
subgenus was subject to some variation between I9g2I and 1951, but it is now 
generally accepted as having full generic status. 


Imago. Left mandible with apical tooth as long as or slightly shorter than first marginal, 
distance from apical to first marginal slightly less to slightly more than one-third distance 
between first and third marginals (left mandible index -29—-41), second marginal obsolete, cutting 
edge between first and third marginals straight ; right mandible with apical and first marginal 
subequal as left, cutting posterior edge of first marginal distinctly longer than that of second 
marginal, molar plate with no marked irregularity of outline in surface view, and 5—9 transverse 
ridges, first much larger than rest, forming a blunt tooth, the more posterior ridges less prominent 
(Text-figs. 185-195). Postclypeus length slightly less to slightly more than half width. Fon- 
tanelle pale coloured, slit-like, forked anteriorly, to approximately triangular, lateral arms 


ETHIOPIAN NASUTITERMITINAE 77 


dilated terminally or tapering. Pilosity of head capsule rather evenly fine, arising from small 
sockets of regular size with a few larger scattered sockets among them. 

Soldier. Dimorphic in most species, with traces of subdivision of ‘‘ major ’’ and “‘ minor ”’ or 
almost complete inter-gradation occurring in some colonies or local populations of some species. 
Nose cylindrical to strongly conical. Vestigial mandibles usually without points. Head 
capsule without any constriction behind antennae, and devoid of prominent setae except on 
nose (major soldier only, minor soldier often has constricted head capsule). 


The longer cutting edges of the imago mandibles, and the mght molar plate, 
distinguish this genus from Nasutitermes. The only genus closely resembling 
Trinervitermes in this respect is Baucaliotermes gen. nov., included in Trinervitermes 
by Emerson (1960) but differing in having monomorphic soldiers and certain other 
features. 

The genus is mainly grass and leaf feeding and occurs only in the savannah and 
steppe vegetation types, (Keay et al., 1959). Wherever agricultural practices have 
degraded the vegetation to grassland in the fringes of the forest zones, Tvinervitermes 
has been found, though it has not been recorded from similar areas deep in the forest. 
Conversely, in equatorial western Uganda, the recent degradation of a forest barrier 
to grassland has not yet led to the mingling of the West African species to the north 
of it, with the East and Central African species to the south. 

There are four groups of species which are recognizable as differing in their geo- 
graphical distribution. The “ West African’ group reaches northern Uganda and 
the Sudan. The “ East African’”’ group includes species both widespread (7. 
rapulum and T. dispar) and more restricted (7. gratiosus and T. bettonianus). There 
is a single “‘ Central African ’’ species, 7. rhodesiensis, and one in South Africa, 
T. trinervoides. The distributions of the East, Central, and South African species 
overlap to a considerable extent, as can be seen from a comparison of maps 20, 
25, 26, and 29. 

Trinervitermes is the most difficult genus in the Nasutitermitinae, and perhaps in 
the order Isoptera. In both imagos and soldiers there is wide variation, and few 
reliable taxonomic characters ; some are of value for one or two species, and value- 
less for the rest. The majority of those used by earlier workers in the genus have 
been abandoned as useless, and a number of new features are used for the first time. 
Unlike the genus Nasutitermes, the species are represented by numerous specimens. 

An interesting result is that the localised operation of distinctive genes or gene 
complexes has been observed in certain species, for example : the partial or com- 
plete suppression of the outer corners of the imago pronotum, or the development of 
unusually large ocelli. Both of these occurred in a number of colonies of 7. dispar 
from the same or nearby localities. 

In the case of the soldier caste, only the major soldiers are identifiable to species. 
It would be possible to key the minor soldiers to groups of species, but these would 
be too large to be of practical value ; even in the case of the major soldiers, the key 
is based on the characters of the majority, since there are specimens of several 
species which cannot be reliably identified in the absence of their imago. On the 
other hand, some species are more readily separated in the soldier caste than in the 
imago. 


78 WwW. A. SANDS 


Y 160. 4 162. 


iWASY 177. 


— 


178 £ 180 A A 
bie =i icy) 


183. 
> / 184 : 87 
qf niger eg N 
8 8 © 5 § 196. 197 
188. 189. 190 191. 192. 193. 194. 195. 


Fics. 157-197 : Imago mandibles of ‘‘ Procorniteymes branch ’’, showing details of right 
molar plate. 157-159, Nasutitermes avborum ; 160-162, N. elegantulus ; 163-165, 
Leptomyxotermes doriae (late nymph) ; 166-168, Grallatotermes africanus ; 169-171, 
Fulleviteyrmes coatoni ; 172-174, F. contvactus ; 175-177, F.mallyi ; 178-180, F. 
tenebricus ; 181-183, Rhadinotermes coarctatus ; 184-187, Tvinervitermes trinervoides ; 
188-190, T. bettonianus ; 191, T. dispar; 192, T. gratiosus; 193, T. vapulum ; 
194, 195, I. rhodesiensis ; 196, 197, Baucalhotermes hainest. 


ETHIOPIAN NASUTITERMITINAE 79 
KEY TO SPECIES 


IMAGosS 


Ocellus separated from compound eye by more than two-fifths of own width (i.e. 
shortest diameter) : Z : 2 
Ocellus separated from compound eye by ene fifths of own width or less ; : 4 
Frons usually with two shallow diverging depressions in front of fontanelle. Smaller, 
head width across eyes (W), 1-45-1-83 ; greatest diameter of compound eye (E), 


0:43-0°58 ; hind tibia length (T,), 1-81-2-24 : ; 2 : : ; 3 
Frons without such co ea ie W, 1:61-2:03; E, 0-42-0°56; Ts, 
2°02-2'47 . : . trinervoides (p. 132) 


Fontanelle with short tera arms. Compound eye larger in proportion to head 
width, W/E index 3-0-3-5. Ocellus separated from eye by up to half own width. 
Female cerci longer than basal breadth . , : rapulum (p. 110) 

Fontanelle lateral arms long, reaching almost half way "es pod. W/E index 3-5-3:9. 
Ocellus separated from eye by half own width or more. Female cerci as long as 


basal breadth ? : : ; ; : ‘ é ; saudiensis (p. 120) 
Antennae 15-16 segmented 5 
Antennae 17-18 segmented F . ; 14 
Scent gland on fifth abdominal sternite of female large, over 1-omm. w ide, over 

o-25mm. long ; that of male smaller, approximately as female of other species. 6 
Scent gland on fifth abdominal sternite of female smaller, under o-gmm. wide, under 

0:23mm. long ; that of male, smaller than female . - 7 


Molar plate of right mandible with 5-6 ridges, posterior ridges obsolescent, some- 
times unevenly shaped. Smaller, W, 1:55-1-82 ; E, 0:47-0:59 ; Ts, 1:98-2:43. 
Antennae 15 segmented, sometimes with a basal segment partially subdivided 
bettonianus (p. 84) 
Molar plate of right mandible with 8-9 ridges. Larger, W, 1:80-1:96 ; E, 0-58- 
0:60 ; T,, 2:32-2:50. Antennae, 63% of specimens 16 or more segmented 
gratiosus (p. 97) 
Antennae 16 segmented or if 15, with one or more basal segments partially sub- 


divided : : ¢ : ; : : : f - : q F 8 
Antennae 15 segmented . : 10 
Vertex with three diverging pale streaks or patches on ‘darker ground éolowsr. 

W. Africa, N. Congo, N. Uganda, Sudan : : togoensis (p. 122) 
Vertex without such streaks, though sometimes clouded or patched with aed colour 

in one area. East Africa, S. Congo, Central and S. Africa 3 9 
Frons usually with two shallow diverging depressions anterior to fontanelle: * Bose: 

clypeus, width 1-9—2-2 times length : 2 Z rapulum (p. 110) 
Frons without such depressions, though sometimes oath “single median shallow 

depression. Postclypeus, width 2-2—2-5 times length : . rhodesiensis (p. 114) 
Compound eye, greatest diameter under 0-50 ; : : : : é II 
Compound eye, greatest diameter 0-50 or more : : 12 


Frons usually with two shallow diverging depressions in front of fontanelle ; fon- 
tanelle short broad Y-shaped or U-shaped, margins often indistinct. Postclypeus, 
width 1-9-2-2 times length. Larger, W, 1:45-1:83 ; E, 0:-47—-0'58 ; T,, 1:81-2 
rapulum (p. 110) 
Frons without paired depressions, sometimes with single median shallow depression 
in front of fontanelle, sometimes flat ; fontanelle slender Y-shaped, sometimes 
with dark patch at posterior end. Postclypeus, width 2-1-2-5 times length. 
Smaller W, 1:35-1:58 ; E, 0:43-0:50 (one only recorded) ; T,, 1-63-1-99 
dispar (p. 87) 


80 W. A. SANDS 


209. 210. 


Fics. 198-242: Imago fontanelle. 198, 3, 199, 2, Nasutitermes arborum ; 200, Q, N. 
chrysopleuva ; 201, 3, N. camerunensis ; 202, g, 203, 2, N. elegantulus ; 204, 9, N. 
fullert ; 205, 3, 206, 9, N. infuscatus ; 207, 5, 208, 2, N. kempae ; 209, 210, g, 211, 
212, 9, N. latifrons ; 213, 6,214, 9, N.lwjae; 215,64, 216, 9, N. schoutedent ; 217,64, 
218, 2, Grallatotermes africanus ; 219, 3, 9, Fullevitermes coatoni ; 220, 3, 2, F. con- 
tvactus ; 221, 6, 222, 9, F. mallyi; 223, 35, 224, 9, F. tenebricus ; 225, 3, 226, 9, 
Rhadinotermes coarctatus ; 227, 3, 228, 9, Trvinervitermes bettonianus ; 229, 3, 230, 2, 
T. dispar ; 231, 232, 3, 233, 234, 9, T. geminatus ; 235, 3, 236, 9, T. gratiosus ; 237, 
238, 5, 239, 240, 9, T. occidentalis ; 241, 5, 242, 9, T. oeconomus. 


12 


13 


14 


16 


ETHIOPIAN NASUTITERMITINAE 81 


Frons usually with two shallow diverging depressions in front of fontanelle . : 13 
Frons without such depressions, though sometimes with single median depression 
(see also 9 above) : : . rhodesiensis (p. 114) 


Eye relatively smaller, W/E cuties 3 satis 5- Postclypeus more inflated, width 

1-9-2:2 times length. Ocelli separated from compound eyes by one-fifth to one- 

half own shortest diameter. East, Central, South Africa. : rapulum (p. 110) 
Eye relatively larger, W/E index 2-6—3-0. Postclypeus less inflated, width 2-2—2-6 

times length. Ocelli separated from compound eyes by one-tenth to one-fifth own 

shortest diameter. W. Africa, N. Congo, S. Sudan, N. Uganda occidentalis (p. 100) 
Pilosity of ‘‘ forehead ’’ consisting of a “‘ pelt ’’ or “‘ mat’’ of short, curved setae of 

even length, with scattered long setae 3—5 times longer than these (Text-figs. 383, 

385). Scent gland on fifth abdominal sternite of female smaller, width usually 

under o-gmm., length under o-2mm. (some T. ¢vinervius larger, approach the 

larger dimensions of T. oeconomus) : 15 
Pilosity of ‘‘ forehead ’’ uneven, setae generally straight or ‘slightly sunved, length 

variable from very short to long, not forming an even “ pelt ’’ (Text-fig. 384). 

Scent gland on fifth abdominal sternite of female larger, width over t-omm., 

length over 0-3mm. (some T. geminatus have more uneven pilosity, but these have 

the small scent gland on the fifth female abdominal sternite) 16 
Postclypeus, more strongly inflated, width 2-:2-2-4 times length. Eyes larger in 

proportion to head width, W/E index 2-3—2-9. Cerci of female usually longer than 

basal breadth, some equal. : : trinervius (p. 127) 
Postclypeus, less inflated, width 2-4-2:8 times length. Eyes smaller in proportion 

to head width, W/E index 2:7—3-0. Cerci of female shorter than basal breadth, 

rarely equal F : geminatus (p. 92) 
Pilosity of “‘ forehead ”’ sparse and fine, Postclypeus more inflated, width 2-0-2°3 

times length. Ocelli separated from eyes by one-sixth to two-fifths own shortest 

diameter. Cerci of female as long as basal breadth, or shorter. East Africa, 

S. Uganda, S. Congo, N. Angola . , gratiosus (p. 97) 
Pilosity of ‘“‘ forehead ’’ abundant and coarse ‘(Text- fig. 384). Postclypeus less 

inflated, width 2-3-2-6 times length. Ocelli separated from eyes by one-fifteenth 

to one-quarter own shortest diameter. Cerci of female longer than basal breadth. 

W. Africa across to N. Uganda and Sudan _. ; ‘ . oeconomus (p. 105) 


SOLDIERS (MAJOR SOLDIERS ONLY) 


Nose in profile cylindrical or weakly conical (angle of cone o-14 degrees) 

Nose in profile moderately to strongly conical (angle of cone 15-25 a : : II 
Nose tip distinctly hairy . 

Nose tip with four prominent setae at most, often few er or none, and w ith a few 


minute, pale setae behind them, or none 6 
Antennae 13 segmented 4 
Antennae 14 segmented 5 


Fontanelle large, 0-o5mm. to over o-Imm. across. Nose in profile more broadly 
conical towards base. Head capsule in plan view “‘ shouldered ’’ behind base of 
antennae (Text-figs. 282-289) : . occidentalis (p. 100) 

Fontanelle small 0-03—0-05mm. across. Meee in prodile Ereaty tapered to near 
base. Head capsule evenly rounded in plan view (Text-figs. 351-358, 376-378) 

T. arabiae (p. 82) and togoensis (p. 122) 

Nose usually as long as or longer than rest of head capsule measuring from hind 
margin of antennal socket. South Africa, parts of South-Central Africa 

trinervoides (p. 132) 

Nose usually shorter than rest of head capsule, measuring from hind margin of 

antennal socket. West Africa, N. Congo, across to Sudan, N. Uganda : : I2 


82 WW. A. SANDS 


6 Head capsule in plan view “ shouldered ”’ behind base of antennae : ‘ . 7 
— Head capsule outline various, not “shouldered ’’ in front . 8 
7 Head capsule in profile rising from base of nose above antennal eee in alone a 
straight line to near highest point, before downward curve to occiput. Nose 
always shorter than rest of head a measuring from hind margin of antennal 
socket (Text-figs. 338-343) . : : rapulum (p. Ifo) 
— Head capsule in profile scarcely rising in a curve ae base = nose to behind antennal 
socket, then commencing convex curve to highest point, continuous with down- 
ward curve to occiput. Nose often equal, sometimes longer than rest of head 
capsule, measuring as above, sometimes shorter (Text-figs. 344-350) 
rhodesiensis (p. 114) 
8 Hind tibia under 1-39mm. in length . : i : ; : : dispar (p. 87) 
— Hind tibia over 1:-39mm. in length . F 9 
g Paler coloured, head capsule yellow to feccupiions) nose denice, oxeaiee: to a. 
brown, with paler tip. West Africa, N. Congo, Sudan, N. Uganda 
oeconomus (p. 105) 
— Darker coloured, head capsule ferruginous to pitchy brownish black, nose darker than 
head, chestnut brown to jet black with red tip : 10 
10 Head capsule in profile rising from base of nose in front of antennal socket more or 
less straight to highest point of head, nose always shorter than rest of head 
capsule. East Africa, Southern Congo, Angola : : gratiosus (p. 97) 
— Head capsule in profile scarcely rising in a curve to behind antennal socket, then 
convex to highest point. Nose often equal to or longer than rest of head capsule. 


West Africa, N. Congo, Sudan, N. Uganda . : . : trinervius (p. 127) 
iz Antennae 12-13 segmented. FE. and Central Africa . : bettonianus (p. 84) 
— Antennae 14 segmented. W. Africa, N. Congo, Sudan, N. Uganda ‘ 3 ; 12 


12 Middle abdominal tergites, seen in profile, with distinct (though minute) setae over 
entire surface. Colour of head capsule usually darker, ferruginous orange, to 
dark sepia brown, nose darker with red tip (Text-figs. 270-275) geminatus (p. 92) 
— Middle abdominal tergites, seen in profile, with setae so minute and pale as to appear 
absent except at high magnifications. Colour of head capsule usually paler, yellow 
to ferruginous, nose darker, orange to chestnut brown, with paler tip (Text-figs. 
290-296) . ‘ : : : ‘ ; : : .  oeconomus (p. 105) 


Trinervitermes arabiae Harris 
(Text-figs. 376-378 ; Map 3) 
Trinervitermes avrabiae Harris, 1957 : 430. Type locality : WESTERN ADEN PROTECTORATE, 

Dhala. 

The soldier caste of this species is indistinguishable taxonomically from T. 
togoensis (Sjéstedt), but the nearest recorded locality of the latter is in Uganda, a 
distance of about 1,000 miles. In the absence of the imago caste and of examples 
of the soldier from intermediate localities, I consider it undesirable to reduce T. 
arabiae to a Synonym at present. It may be noted however, that evidence from 
other genera shows that species typically found in the Guinean to Sahel vegetation 
zones (Keay et al., 1959, types 8, 16, 17, 20, and 25) also occur in South-West Arabia. 


ETHIOPIAN NASUTITERMITINAE 


Fics. 243-255 : Trinervitermes, front and side views of imago head capsule. 
T. bettonianus ; 246, 247, T. dispar ; 248, 249, T. geminatus ; 250, 251, T. gratiosus ; 
252, 253, I. occidentalis ; 254, 255, T. oeconomus. 


243-245, 


84 W. A. SANDS 


Trinervitermes bettonianus (Sjostedt) 
(Text-figs. 188-190, 227, 228, 243-245, 256-261 ; Map 29) 


Eutermes bettonianus Sjostedt, 1905 : 19. Type locality : Kenya, Athi River. 

Eutermes segelli Sj6stedt, 1907a : 26. Type locality : Tancanyi1ka, Boma ngombe, Kiliman- 
jaro area. 

Eutermes (Tvinervitermes) bettonianus Sjostedt ; Holmgren 1912 : 64. 

Eutermes (Tvinervitermes) segelli Sjostedt ; Holmgren 1913 : 348. 

Eutermes (Trvinervitermes) ruficeps Holmgren, 1913 : 343. Type locality : TANGANYIKA, Amani. 
syn. n. 

Eutermes crassinasus SjOstedt, 1914 : 6. Type locality : MozamMBiqueE, Zambesi, Villa Fontes. 
syn. n. 

Trinervitermes bettonianus (SjOstedt) Sjostedt, 1926 : 332. 

Trinervvitermes segelli (SjOstedt) Sjéstedt, 1926 : 326. 

Trinervitermes cvassinasus (Sj6stedt) Sjéstedt, 1926 : 326. 

Trinervitermes ruficeps (Holmgren) Sjéstedt, 1926 : 327. 

Nasutitermes (Ivinervitermes) bettonianus (SjOstedt) Emerson, 1928 : 492. 

Trinervitermes bettonianus (SjOstedt) ; Snyder, 1949 : 323. 

Trinervitermes crassinasus (SjOstedt) ; Sands, 1957 : 21 


Imago. Some additions to the previous description (Sands, 1957) are necessary. Eyes 
medium sized relative to head width, W/E index 2-9—-3-4 ; ocelli separated from eye by one- 
eighth to one-third own least diameter ; postclypeus width 2:0—2-5 times length ; molar plate 
of right mandible with 5—6 ridges, posterior ridges somewhat atrophied. 

Pale coloured “‘ scent’ gland on anterior margin of fifth abdominal sternite of 9 very large, 
1-07—1 33mm. wide, 0:28—-o-51mm. long ; cerci of 2 slightly shorter to slightly longer than width 
across base, conical or slightly mamilliform. Pilosity of vertex uneven, not forming a 
 jovellic ip mavens © 

Measurements (34 specimens from 13 localities) in millimetres. 


Range Mean 
Head width across eyes 1°55-1°82 : - 1-68 
Greatest diameter of eye 0:47-0'59 : ; 0°53 
Ocellus F 3 ‘ 0:20-0:24 X 0:22-0'31  . : 0-21 X 0-26 
Ocellus toeye . ; 0:03-0:06 : : 0-04 
Width of pronotum . I+39-1°85 : 7 1°57 
Length of pronotum . 0:88-1:22 ; ; I-03 
Length of hind tibia. I -98—2°43 ; : 2°13 
Length of fore wing . 14:2-22°3 : 17°95 


Soldiers. The range of variation now includes the more momaded head capsule in plan view 
previously ascribed to T. crassinasus. Antennae 12-13 segmented, 4 out of 5 with 12. 

Measurements (Major soldier, 119 specimens from 35 localities ; minor soldier, 40 specimens 
from 25 localities) in millimetres. 


Major Soldiers Range Mean 
Head length to tip of nose. 1-93-2°49 ; : : : 2220 
Width of head 6 . 2 0-97-1°43 : - é c 1:20 
Depth of head : : é 0:72-1:07 : 4 ; : 0-90 
Width of pronotum : : 0:54-0°72 : : : : 0:63 
Length of pronotum : : 0-22—0°32 : ‘ : : 0:27 
Length of hind tibia : : I-II-1I‘57 : : : : 1°34 

Minor Soldiers Range Mean 
Head length to tip of nose. I -33-1-°82 : : : : 1°58 
Width of head : : ; 0°57—-0'93 2 : : : 0°75 


Depth of head F : : 0:43-0:70 : : : : 0°57 


ETHIOPIAN NASUTITERMITINAE 85 


Minor Soldiers Range Mean 
Width of pronotum F : 0°40-0°54 : : ; . 0°47 
Length of pronotum 2 : 0-18-0-25 : ‘ - ¥ 0:22 
Length of hind tibia - - 0-95-1°40 ; : : ; 1-18 


The relationships of this species have been discussed in an earlier paper, Sands 
(1957). Since then, specimens have been examined of the soldier caste of T. 
rhodesiensis and T. occidentalis which approach T. bettontanus in shape, but differ in 
the former by the less conical nose and the latter by the larger fontanelle ; both 
have 13-14 segmented antennae. The imagos of these species differ in having much 
smaller “‘ scent ’”’ glands on the 9 fifth sternites. 


Lectotypes have been designated from the existing syntype material of 7. 
bettoniranus (Sjostedt), and T. crassinasus (Sj6stedt), as indicated below, the desig- 
nation for the senior synonym being given separately. 

Lectotype : Kenya: Athi River, 8—19.v.1899 (C. S. Betton), lectotype 3 imago, 
T. bettonianus (Sjéstedt), in B.M.(N.H.). (Paralectotype imagos in Naturhist. 
Riksmus., Stockholm.) 


Type Material. TANGANYIKA : Amani, (Morstatt), syntype soldier, type colony, 
T. ruficeps (Holmgren), A.M.N.H. (Other syntypes in Mus. Stuttgart.) 

MozaMBIQUE : Villa Fontes, 1914 (H. Swale), lectotype soldier and paralectotype 
soldier, type colony, T. crassinasus (Sjéstedt), B.M.(N.H.). (Other paralectotype 
soldiers from type colony in Naturhist. Riksmus., Stockholm.) 


Other Material (additional to that recorded in Sands, 1957). KENYA : Machakos, 
17.11.1951 (W. V. Harris) ; Ruiru, 11.1ii.1951, Thika, Ngong (R. C. H. Sweeney) ; 
Thika, 15.v.1952, Ngong, 16.vi.1953 (W. A. Sands). 

TANGANYIKA : Ukerewe Is., 1938 (W. V. Harris) ; Mwanza, 4.ix.1948, Amani, 
4.1X.1949, II.x.1950, and 4.i.1952, Itigi, 8.ix.1949, Singida, 10.1x.1949, Ngomeni, 
1g.v.1951 (P. B. Kemp) ; Tanga, 1950, 4 vials, Pangani, 20.11.1951 (R. C. H. 
Sweeney), Hakara, 10.vii.1949, Kwiro, 20.vii.t949, and Madidi, 24.vii.1g49 (M. 
Luscher), A.M.N.H. 

NYASALAND : nr. Ft. Johnstone, 23-24.viii.1953, 2 vials, 2m. from Bilila on Ft. 
Johnstone Road, 27.vili.1953, 20m. from Ft. Johnstone on Zomba Road, 28.viii. 
1953, Im. W. of Lake Shirwa, 31.viii.1953, Namatalu Hill, 1.xi.1953, 2 vials (W. A. 
Sands, W. Wilkinson). 

NORTHERN RHODESIA : Choma, Siamambo 10.i.1957, 2 vials, near Kafue River 
bridge between Lusaka and Mazabuka, 18.i.1957, 2 vials, 26m. from Broken Hill on 
Lusaka Road, 20.1.1957, 4m. N. of Lusaka, 19.1.1957 (W. G. H. Coaton). 

SOUTHERN RHODESIA : Rekomitjie, 6.x.1964 (M. G. Bingham). 

A total of 79 nest series were examined, and all material is in the British Museum 
(Natural History) unless otherwise stated. 


The distribution and biology of this species were discussed in an earlier paper, 
Sands (1957). Further study and new records suggest that T. bettonianus does not 
extend far to the south of the valley of the River Zambesi. Areas of apparently 


86 W.-A. SANDS 


suitable vegetation types occur considerably further south, but it has not been 
collected, whereas its less conspicuous congener, T. dispar, has been recorded from 
Natal. 


Map 19. Trinervitermes bettonianus. Vegetation types: 9, 20, 22, and 25. 


ETHIOPIAN NASUTITERMITINAE 87 


Trinervitermes dispar (Sj6stedt) 
(Text-figs. 191, 229, 230, 246, 247, 262-269 ; Map 20) 


Eutermes dispar Sjéstedt, 1920a : 303. Type locality : NyasALANp, Zomba. 

Eutermes gemellus Sjostedt, 1920a : 303. Type locality: REPUBLIC OF SOUTH AFRICA. 
syn. n. 

Termes (Eutermes) dispar (Sjostedt) Desneux, 1904a : 41. 

Termes (Eutermes) gemellus (SjOstedt) Desneux 1904a : 42. 

Eutermes dispar SjOstedt ; Sjostedt, 1904 : 107. 

Eutermes gemellus Sj6stedt ; Sjéstedt, 1904 : 98. 

Eutermes kulloensis Sjéstedt, 1912 : 13. Type locality, Etuiopia, Eritrea, Mt. Kullo, Maggio. 
syn. n. 

Eutermes (Trinervitermes) dispar Sjostedt ; Holmgren, 1912 : 64. 

Eutermes (Trvinervitermes) gemellus SjOstedt ; Holmgren, 1912 : 65. 

Eutermes katangensis Sj6stedt, 1913 : 382. Type locality, Conco, Katanga, Elizabethville. 
syn. n. 

Eutermes (Trinervitermes) erythreae Holmgren, 1913 : 345. Type locality : Erniopta, Eritrea. 
syn. n. 

Eutermes grootfonteinsis Sjéstedt, 1914a : 86. Type locality : Soura West Arrica, Grootfon- 
tein. syn. n. 

Trinervitermes umzinduzit Fuller, 1922: 114. Type locality : REPUBLIC OF SoUTH AFRICA, 
Natal, Pietermaritzburg. syn. n. 

Trinervitermes dispar (SjOstedt) Fuller, 1922 : 108. 

Trinervitermes gemellus (Sj6stedt) Fuller, 1922 : 110. 

Trinervvitermes kulloensis (Sjostedt) Sjéstedt, 1926 : 328. 

Trinervitermes katangensis (SjOstedt) Sjéstedt, 1926 : 328. 

Trinervitermes evythreae (Holmgren) Sjéstedt, 1926 : 327. 

Trinervitermes grootfonteinsis (SjOstedt) Sjéstedt, 1926 : 330. 

Trinervitermes eldirensis Ghidini, 1941 : 33. Type locality : Eruiopia, El Dire. syn. n. 
Imago. Some additions to the previous description (Sands, 1957) are necessary. Eyes 

medium sized relative to head width, W/E index 2-90-3:3 ; ocelli separated from eye by slightly 

less than one-sixth to slightly less than two-fifths own shortest diameter ; postclypeus width 

2-1-2'5 times length ; molar plate of right mandible with 7-8 ridges. 
Pale coloured ‘“‘ scent ’’ gland on anterior margin of fifth abdominal sternite of 2 very small, 

0-45-0'54mm. wide, 0-15—o-18mm. long ; cerci of 2 distinctly longer than width across base, 

conical. 


Pilosity of vertex uneven, not forming a “ pelt ’’ or ‘‘ mat’’. 


Measurements (41 specimens from 17 localities) in millimetres. 


Range Mean 
Head width across eyes 1*35-1°58 : : 1°44 
Greatest diameter of eye 0°42-0°50 : 3 0-46 
Ocellus : : : 0:15-0:20 X 0:17-0:26. : 0-18 X 0:22 
Ocellus toeye . : 0:03-0:06 : 4 0-04 
Width of pronotum . 1-20-1°48 : P I +32 
Length of pronotum . 0°76—-0:97 ; : 0°84 
Length of hind tibia. 163-1 -99 ; : 1°97 
Length of fore wing. 14°00-18-90 : : 16-10 


Soldiers. The following additions to the previous description of the major soldier are neces- 
sary : nose cylindrical to moderately conical, angle of cone o—20 degrees, usually less than 14 
degrees. Head capsulein plan view rounded oval, rarely slightly shouldered behind antennae. 
Nose tip with four prominent setae, and a few minute pale setae behind them. 


88 WwW. A. SANDS 


Measurements (Major soldiers, 151 specimens from 39 localities, minor soldiers, 28 specimens 
from 13 localities). 


Major Soldier Range Mean 
Head length to tip of nose. I°71-2°23 5 : : : 2:02 
Width of head : : - 0:93-1 -36 : ; : ; I-05 
Depth of head 3 : ; 0:68-0:99 ; : , : 0°84 
Width of pronotum : : 0-54-0°68 : : : : 0:58 
Length of pronotum : : 0-18—0'25 E : ; : 0:23 
Length of hind tibia F : 104-1 °39 : : ; : I-19 

Minor Soldier Range Mean 
Head length to tip of nose. 118-1 -61 3 : ; : I-41 
Width of head : : : 0-43-0°64 ‘ : ; : 0°53 
Depth of head : , j 0:36-0'50 : ‘ , - 0°43 
Width of pronotum ; : 0-38-0°47 : ‘ ; ‘ 0-41 
Length of pronotum : : 0-13-0:20 : : ; : 0-16 
Length of hind tibia F . 0:82-1:18 5 ‘ : : I-O1 


A darker coloured form of the imago with dusky pronotum has been recorded from South 
Africa, but it is less variable in size and shape than most other species. It only overlaps in size 
range with T. bettonianus and T. vapulum among East African species, and T. occidentalis and 
T. togoensis among West African. 


T. dispar differs from T. bettonianus in the very much smaller “ scent ’’ gland on 
the 9 fifth abdominal sternite, in the narrower fontanelle, and in the 7-8 ridges on 
the right molar plate. 7. vapulum differs in the shape of the fontanelle, in the some- 
what more inflated postclypeus, and in most specimens, in having two small diverging 
depressions in front of the fontanelle, though these are not always present. T. 
togoensis has usually 16 segmented antennae, and three diverging pale coloured 
streaks on the vertex ; T. occidentalis has larger eyes. 

The range of variation in the soldier caste overlaps with T. rhodesiensis, T. rapulum, 
and JT. togoensis. Some specimens of T. rhodesiensis and T. rapulum are indistin- 
guishable from some T. dispar, but T. togoensis is usually separable by the more 
densely hairy nose tip, and the other two species by slight differences of profile and 
plan view of the head capsule, given in the key. 


Type Material. Eruiopia: Eritrea, Mt. Kullo 1871 (Antinom), T. kulloensis 
(Sj6stedt), J imago specimen labelled “‘ co-typus ’’, unique, Mus. Civ. Stor. Nat., 
Genoa. The additional material recorded by Sjéstedt as deposited at the Naturhist. 
Riksmus., Stockholm, was reported in September 1962, to be lost ; Eritrea 
(Escherich), syntype soldiers from type colony, T. erythreae (Holmgren) A.M.N.H. 
and B.M.(N.H.) ; El Dire, 18.v.1939 (G. M. Ghidint), paratype soldiers from type 
colony, 7. eldirvensis Ghidini B.M.(N.H.) and A.M.N.H. (Holotype in Mus. Civ. Stor. 
Nat., Genoa.) 

NYASALAND : Zomba (Cameron), syntype soldiers, 7. dispar (SjOstedt) Naturhist. 
Riksmus., Stockholm. 

Conco: Katanga, Elizabethville (J. Bequaert), syntype soldiers, type colony, T. 
katangensis (Sjostedt) A.M.N.H. (other syntypes listed by Sjéstedt (1926) as in 
Naturhist. Riksmus., Stockholm). 


ETHIOPIAN NASUTITERMITINAE 89 


269. 


275. 


Per 273 


Fics. 256-275 : Tvinervitermes, soldier head capsule, side and plan views, and nose tip. 
256-261, T. bettonianus ; 262-269, T. dispar ; 270-275, T. geminatus. 


90 W.-A. SANDS 


SOUTH-WEST AFRICA : Grootfontein 7.vi.1g11 (W. Michaelsen), syntype soldiers, 
type colony, T. grootfonteinsis (Sjéstedt) A.M.N.H. (other syntypes, in Mus. Hamburg 
and Naturhist. Riksmus., Stockholm). 

REPUBLIC OF SOUTH AFRICA : No other data. Syntype soldiers, type colony T. 
gemellus (Sj6stedt), A.M.N.H. and Naturhist. Riksmus., Stockholm ; Natal, 
Pietermaritzburg x1.1918 (E. Warren), syntype 2 imago and soldiers from type 
colony T. wmzinduzi Fuller, B.M.(N.H.) and A.M.N.H. (other syntypes at N.C.L., 
Pretoria) ; Transvaal, Pretoria, Meukleneuk Hill 22.x.1914 (C. Fuller), “‘ Paratype ”’ 
soldiers, T. pretoriensis Fuller, vial number F.491, not type colony, not same species 
as type colony, A.M.N.H. and N.C.I., Pretoria. 


Other Material (additional to that recorded in Sands, 1957). SOMALI REPUBLIC : 
Bender Kassim, 9.1.1952 (FE. J. Van Ingen). 

Kenya : Rumuruti, 8.x.1950 (W. V. Harris) ; Marsabit, 6.3.53 (W. A. Sands) ; 
Taveta (Kirby), A.M.N.H. 

UGanbDaA : Karamoja Distr., Toror Hills, near Kotido, 8.x.1952 (W. A. Sands) ; 
Ankole Distr., Luentobo, 1947 (W. V. Harris). 

TANGANYIKA : Mkweme, near Kahama, 13.viii.1948 and 18.iii.1949, Tabora, 
7.1x.1948, Tongoni, near Tanga, 25.x.1950 and Pare Mountains near Kihurio, 
21.v.1952 (P. B. Kemp) ; Igalula, 21.xii.1934, Kahama, v.1935 (W. V. Harris) ; 
Mkasu, 26.vii.1949 (M. Luscher), A.M.N.H. 

NYASALAND: 17m. W. of Blantyre, 20.vill.1953, 18m. from Ft. Johnstone on 
Farringdon Road, 24.vili.1953, 36m. N. of Mzimba 28.ix.1953 (W. A. Sands, W. 
Wilkinson). 

NORTHERN RHODESIA : Siamambo, Choma, 10, 11, and 14.1.1957, 3 vials, 15—1gm. 
from Kitwe on Ndola Hill Road, 23—24.1.1957, 3 vials, and Samfya, Lake Bangueulu, 
27.1.1957 (W. G. H. Coaton) ; Sikalongo, I1.vili.1959 (E. N. Cooling). 

SOUTHERN RHODESIA : Salisbury, 1g11 (G. A. K. Marshall), Mt. Darwin, 15.vi. 
1949 (G. H. Bunslt). 

SWAZILAND : 3m. S. of Komati River, Piggs Peak to Mbabane Road, 23.x.1960, 
and 6m. from Gollel on Hluti Road, 25.x.1960 (J. L. Sheasby) ; 4m. from Stegi on 
Gollel Road, 24.x.1960 (W. G. H. Coaton). 

REPUBLIC OF SOUTH AFRICA: Transvaal, Witbank, 3.1x.1939, Krugersdorp, 
1.x.1956, Groblersdahl, 15.xi.1956, Warmbad, 25.1x.1957, 2 vials, Waterberg, 
25.1x.1957, and 27.1x.1961, Potgietersrus, 26.1x.1957, 2 vials, Pilgrims Rest, 27—28.x. 
1959, 3 Vials, Soutpansberg, 10.x.1960, 3 vials, Nelspruit, 21.x.1960, 2 vials (W. G. H. 
Coaton) ; Barberton, 29.xi.1956, and Sibasa, 5—I0.vill.1959, 4 vials (J. H. Grobler) ; 
Marico, 4.x.1961 (G. F. Pretorius). Natal, Lower Umfolosi, 11.xi.1922 (R. H. 
Harris) ; Hlabisa, 7.1i1.1937 (C. Jacot) ; Ubombo, 21.xi.1955, Mtuzini, 28.x.1957, 
Ingwavuma, I4.xll.1959, 2 vials ; Pietermaritzburg, 5.x.1957, 7 vials (A. Hewett, 
D. Fletcher) ; Ndwedwe, 11.xii.1959 and Richmond, 14.x11.1959 (P. C. Joubert) ; 
Nongoma, 10.1.1962, Entonjaneni, 13.1.1962 and Nkandhla, 14.1.1962 (J. L. Sheasby), 
Cape Province, Mafeking, 6.x.1961 (G. F’. Pretorius). 

A total of 11I nest series were examined, and all material is in the British Museum 
(Natural History) unless otherwise stated. 


ETHIOPIAN NASUTITERMITINAE gi 


This species is distributed over a much greater area than was recorded in a 
previous paper (Sands, 1957), from Mount Kullo in northern Eritrea, to southern 
Natal. It appears to be found mainly in vegetation types ranging from savannah 
woodland with Brachystegia and Julbernardia to wooded steppe with Acacia and 
Commiphora (Keay et al., 1959, types 9, 18, 19, 20, 25, and 26). These areas are 
shaded on the map. It has only rarely been found in Colophospermum mopane 
woodland (Type 22). 


a 


Map 20. Trinervitermes dispar. Vegetation types: 18, 19, 20, and 22. 


92 W. A. SANDS 


It appears to fill an equivalent ecological niche to 7. togoensis with which its 
distribution only marginally overlaps, and its exclusion from the Guinean and Sudan 
vegetation zones may result partly from the biotic barrier caused by this similarity. 


Trinervitermes geminatus (Wasmann) 
(Text-figs. 231-234, 248, 249, 270-275, 385 ; Map 21) 


Eutermes geminatus Wasmann, 1897 : 170. Type locality : GHANA. 

Termes (Eutermes) geminatus (Wasmann) Desneux, 1904a : 42. 

Eutermes geminatus Wasmann ; Sj6stedt, 1904 : 40, 44. 

Eutermes (Trinervitermes) geminatus Wasmann ; Holmgren, 1912 : 64. 

Eutermes (Trinervitermes) gvossus Sjéstedt, 1924b: 493. Type locality: MAuRITANIA 
(probable), “‘ Cote Atlantique du Sahara’’. syn. n. 

Trinervitermes grossus (SjOstedt) Sjdstedt, 1926 : 342. 

Trinervitermes geminatus (Wasmann) Sjéstedt, 1926 : 343. 

Trinervitermes ebenervianus Sjostedt, 1925a : 73. Typelocality : SuDAN, “‘ Tanguru”’. syn.n. 

Trinervitermes tbidanicus Sjéstedt, 1926 : 347. Type locality : Nicreria, Ibadan. syn. n. 

Trinervitermes ebenerianus Sjostedt ; Sands, 1957 : 24. 


Imago. Head capsule, central part of frons and vertex brown to dark chestnut brown, 
vertex with three diverging yellow-brown streaks ; postclypeus, labrum, anterior part of frons 
behind postclypeus and in front of ocelli, area surrounding antennae and eyes and extending to 
a point behind eyes also paler, orange-yellow to yellow-brown. Antennae orange-yellow to 
orange. Pronotum orange-yellow to brown. Meso- and metanota, other thoracic sclerites and 
legs yellow to yellow-brown. Abdominal tergites brown to chestnut brown round stigmata, 
slightly paler in middle, distinctly paler towards lateral margins ; sternites yellow to yellow- 
brown, shaded brown round stigmata. Wings translucent very pale yellow-brown to brown, 
subcosta and radius sector yellow to yellow-brown, partially outlined brown at base, membrane 
behind radius sector with yellow to orange streak, rest of venation narrowly yellow-brown to 
brown. 

Posterior margin of head capsule behind eyes much less than semi-circular, slightly sinuate 
immediately behind eyes ; fontanelle distinct, paler than head, V- or inverted A-shaped, some- 
times slightly depressed posteriorly, lateral arms often but not always dilated terminally ; area 
of fontanelle usually partly occluded posteriorly by pigmented cuticle only slightly paler than 
head, length including occluded portion approximates to twice breadth at base of lateral arms ; 
eyes oval, moderately prominent in proportion to own diameter, and medium-sized relative to 
head width, W/E index 2-7—3:0 ; ocelli medium sized to large, oval, separated from eyes by one- 
sixth to nearly one-third own least diameter ; postclypeus moderately inflated, width 2-4—-2:8 
times length, anterior margin slightly concave or sinuate, rarely nearly straight, posterior 
margin convex, rounded ; molar plate of right mandible with 7-8 ridges, antennae usually 17 
segmented, II, IV, V, and VI subequal, VI slightly longer, III longer up to nearly twice these, 
sometimes divided to produce 18 segmented forms. 

Pale coloured “‘scent’”’ gland on anterior margin of fifth abdominal sternite of 2 medium 
sized, 0:64 to o-80mm. wide, 0-15 to 0-18mm. long ; cerci of 2 female shorter than or equal to 
width across base, conical. 

Pilosity on vertex consists of short curved setae of rather even length tending to form a 
“pelt ’’ or “mat ’”’, with scattered setae 3-5 times as long. (Text-fig. 385.) 

Measurements (31 specimens from 5 localities) in millimetres. 


ETHIOPIAN NASUTITERMITINAE 93 


Range Mean 
Head width across eyes 1-78—2-03 . ' 1-88 
Greatest diameter of eye 0:59-0'72 : : 0°65 
Ocellus ¢ ‘ , 0:20-0:26 X 0°24-0°34 . : 0:22 X 0:27 
Ocellus toeye. : 0:02-0:06 ; 3 0:05 
Width of pronotum . 1-60-1-89 . 5 1-76 
Length of pronotum . 1:00-1-18 F . L-rO 
Length of hind tibia. 2:43-2°80 - : 2°58 
Length of fore wing. 17 -QO-21-go 19°90 


Soldiervs. Major soldier : head capsule ferruginous orange to dark sepia brown, commonly 
ferruginous to chestnut brown, nose chestnut brown to pitch black, with red tip. Pronotum 
and antennae yellow-brown, meso- and metanota pale yellow-brown, legs yellow. Abdominal 
tergites pale brown to sepia brown, sternites pale yellow. 

Head capsule in plan view short oval, circular, or slightly wider than length, nose conical. In 
profile near straight to distinctly concave, with back of head rising from above antennal socket, 
and rounded ; nose equal in length to distinctly shorter than rest of head capsule, measuring 
from hind margin of antennal socket, angle of nose cone, 8-22 degrees ; antennae 14 segmented, 
proportions of basal segments somewhat variable with size, usually Il and V subequal, III and [IV 
subequal and longer than II and V._ Head setae confined to hairy nose tip ; dorsal surface of 
head capsule without prominent setae. 

Middle abdominal tergites with scattered minute setae, distinctly visible in profile, posterior 
tergites with longer setae. Sternites with short backwardly directed scattered setae, and longer 
downward or slightly forward directed setae on posterior margins, arranged approximately in 
longitudinal rows 3 on each side of mid-line of abdomen, longer setae about 2—3 times length of 
shorter. 

Minor soldier : Colour as darker specimens of major soldier, antennae 13-14 segmented, 
proportions of basal segments variable. 

Measurements (Major soldiers, 57 specimens from 13 localities, minor soldiers, 8 specimens 
from 5 localities). 


Major Soldiers Range Mean 
Head length to tip of nose. 2°17-2-64 : - d : 2°40 
Head width . : . : I‘I7-1°73 : : F : 1°45 
Depth of head capsule . 3 0:84-1:22 : : : 3 I-02 
Width of pronotum : : 0:56-0°75 : : 2 ; 0-68 
Length of pronotum : 5 0:25-0°36 : ; ‘ : 0°32 
Length of hind tibia F ‘ 1°57-2:19 P : ; ‘ 1°84 

Minor Soldiers Range Mean 
Head length to tip of nose. 1-51-2:04 : : . : 1-70 
Head width . ; . : 0+59-1-00 : , : : 0°75 
Depth of head capsule . s 0:49-0'77 , : ; : 0-60 
Width of pronotum : : 0-38-0°51 : x : : 0°47 
Length of pronotum 3 : 0:20-0-26 : - ; : 0-2 
Length of hind tibia : 2 1-40-1-76 : : : 1°59 


Variation. The full range of variation in colour and form in both imagos and soldiers is some- 
times found within a comparatively restricted locality. On the other hand, specimens from the 
extremes of the geographical range, over 3,000 miles apart, are clearly of the same species, and 
no character clines have been detected, though a tendency has been noticed for specimens from 
the moister parts of West Africa to be darker in colour. 


T. geminatus in some of its forms, very closely resembles its sympatric congeners, 
T. oeconomus and T. trinervius. It is readily distinguishable from T. oeconomus in 
the 2 imago by the much smaller abdominal “ scent ’”’ gland and the shorter cerci, 


94 WAS SAINIDIS 


also in most specimens by the differing pilosity of the vertex ; the ¢ is less readily 
separable because the ¢ cerci of all species are longer than in the 9, and the ventral 
“ scent’ glands are smaller. The soldiers are usually darker in colour, and have 
visible small setae on the middle abdominal tergites, but specimens which are 
virtually indistinguishable do occur. 

T. geminatus is more readily distinguished from T. tvinervius in the soldier caste, 
by the distinctly more hairy nose tip, and usually by the darker pigmentation of the 
abdominal tergites. The imagos are less easily separable, having usually propor- 
tionately smaller eyes and less inflated postclypeus ; in addition, the cerci of the 9 
T. tyinervius are usually longer, and somewhat irregular in outline, and the shorter 
vertex setae very regular in length. There is little likelihood of confusion with any 
other species apart from the larger, darker soldiers of T. togoensis, but some indivi- 
duals in any nest series of this species have 13 segmented antennae with the basal 
segments only partially or not subdivided. 


Dr. A. E. Emerson has selected and labelled a specimen from the type series as 
lectotype, but the designation has not been published hitherto: 

Lectotype : GHANA : no other data, lectotype pinned major soldier, 7. geminatus 
(Wasmann) (in Mus. Maastricht). Paralectotypes, Naturhist. Riksmus., Stockholm, 
and A.M.N.H. 


Type Material. MAuRITANIA (Probable) : “‘ Cote Atlantique du Sahara’”’ no 
date (A. Gruvel), syntype soldier, from type colony T. grossus (Sjéstedt), A.M.N.H. 
(other syntypes, Mus. Paris, and Naturhist. Riksmus., Stockholm). 

GHANA : no other data. Paralectotype soldiers from type colony, T. geminatus 
(Wasmann), A.M.N.H. 

NIGERIA : Ibadan, 5.viii.1g22 (A. W. J. Pomeroy), syntype imagos and soldiers 
from type colony T. zbidanicus Sjostedt, A.M.N.H. (other syntypes, Naturhist. 
Riksmus., Stockholm). 

SuDAN : “ Tanguru’”’ (= Tungaru, 10° 14’N, 30° 42’E), 7.1v.1914 (R. Ebner), 
syntype soldiers from type colony T. ebenerianus Sjostedt, A.M.N.H. (other syntypes 
Mus. Vienna and Naturhist. Riksmus., Stockholm). 


Other Material. SIERRA LEONE : Samaia, 29.v.1948 (Ff. A. Squires). 

GHANA: Accra, 29.vi.19g26 (A. W. J. Pomeroy) ; 1926 (W. H. Patterson) ; 
Achimota, 21.1.1955 (W. V. Harris) ; Prampram, 16.xi.1955, 54m. from Accra on 
Ada Road, 17.xi.1955, Afienya, and Nungoa, near Accra, I3—16.vi.1956, I0 vials 
(W. Belfield) ; 30m. N.E. of Accra, ii.1958 (R. Hamilton) ; 12m. from Accra on 
Kumasi Road, 18.11.1959, 12m. S. of Tamale on Kumasi Road, 27.11.1959, 5m. and 
16m. from Daboya on Tamale Road, 28.11.1959, 2 vials, 28m. from Tamale on Bolga- 
tanga Road, 1.111.1959, 55m. from Tamale on Larabanga Road, 3.il1.1959, 35m. 
from Tamale on Yendi Road, 4.11.1959, 4m. from Gambaga on Walewale Road, 
5.111.1959, 25m. S.E. of Gambaga on Yendi Road, and 17m. N.E. of Gambaga on 
Bawku Road, 6.111.1959, 20m. from Bolgatanga on Bawku Road, 8.11i.1959, 20m. 
from Bolgatanga on Tamale Road, and gm. from Bolgatanga on Navrongo Road, 
g.li.1959, 31m. from Navrongo on Tumu Road, 13.1ii.1959, 11m. from Tumu on 


ETHIOPIAN NASUTITERMITINAE 95 


Navrongo Road, 15.iii.1959, 9m. from Lawra on Nandom Road, 18.iii.1959, 6m. 
from Wa on Lawra Road, I9.iii.1959, and 52m. from Wa on Bole Road, 20.11.1959 
(W. A. Sands). 

Upper VoLta : 15m. N. of Bawku (Ghana), 11.11.1959 (W. A. Sands). 


NIGERIA : Eastern Region ; 14m. from Enugu on Abakaliki Road, 1.11.1957, and 
13m. from Awgu on Okigwe Road, 2.11.1957 (W. Wilkinson). Northern Region ; 
Katsina, 1933 (Anon) ; 60m., 1oom., and 140m. from Kaduna on Jos Road, 4 vials, 
125m. S. of Jos, 2 vials, Kano, 2 vials, 5m. S. of Funtua, 7m. S. of Zaria, 13m. S. 
of Makurdi, 1954 (B. J. McNulty) ; Kano, 21.11.1955, and Samaru, 23.11.1955 
(W. V. Harris) ; 20m. N.W. of Zaria on Funtua Road, 14.ii.1956, 6 vials, 30m. N. 
of Funtua on Yashi Road, 14.xi.1956, 3 vials, 43m. E. of Kano, Gaya River, 17.xi. 
1956, 21m. W. of Kano on Katsina Road, 18.xi.1956, 5 vials, 3-gm. N. of Kafin Soli 
on Katsina Road, 20.x1.1956, 3 vials, 41m. N.W. of Katsina on Jibva—Gusau Road, 
21.x1.1956, 4m. W. of Gusau on Sokoto Road, 22.xi.1956, 28--44m. from Sokoto on 
Argungu Road, 23.xi1.1956, 5 vials, 25m. from Sokoto on Gusau Road, 25.xi.1956, 
30-35m. N. of Sokoto on [lela Road, 26.xi.1956, 2 vials, 18m. from Kaduna on 
Zaria Road, 8.xii.1956, 16, 18, 33, 68, and 76m. from Kaduna on Zungeru Road, 
18.xii.1956, 5 vials, 27 and 45m. from Zungeru on Kaduna Road, 19.xii.1956, 2 vials, 
26m. from Minna on Zungeru Road, 20.xii.1956, 3 vials, 16m. S. of Zungeru on Bida 
Road, 21.x11.1956, 41m. from Mokwa on Bida Road, 22.xii.1956, Bida, 23.xi1.1956, 
4om. from Abuja on Bida Road, 24.xii.1956, Diko, tom. from Abuja, 27 and 31.xu. 
1956, 2 vials, Kudaru, on Zaria—Jos Road, 5.ii.1957, 2 vials, 4m. E. of Bukuru, 
7.11.1959, 42m. from Jos on Wamba Road, 8.11.1957, 16, 40, and 75m. from Jos on 
Bauchi Road, and Bauchi, 11—13.11.1957, 7 vials, 18m. from Jos, on Pankshin Road, 
14.11.1957, 32m. from Jos on Kaduna Road, 5.iv.1957, 39m. from Bauchi on Gombe 
Road, 7.v.1957, Gombe, and 6—-30m. E. of Gombe, 9.v.1957, 6 vials, 4om. S.E. of 
Gombe on Numan Road, I0.v.1957, 2 vials, 92m. from Gombe on Numan Road, 
II.v.1957, 6m. from Numan on Yola Road, I1.v.1957, Yola, 13.v.1957, near Mayo 
Faran, 14.V.1957, 2 vials, Faro River, E. of Yola, 15.v.1957, 2 vials, 54m. from Yola 
on Jalingo Road, 16.v.1957, 2 vials, 125m. S.W. of Jalingo on Takum Road via Bel, 
18.v.1957, 50m. S. of Jalingo, 19.v.1957, 35 and 80m. from Yola on Biu Road, 
28.V.1957, 3 vials, near Biu, 29.v.1957, 2 vials, 55m. N. of Biu on Damaturu Road, 
29.V.1957, Maiduguri, and 20—43m. from Maiduguri on Potiskum Road, 31.v.1957, 
and I.vi.1957, 4 vials, 43-50 and 77m. from Maiduguri on Fort Lamy Road, 3.vi. 
1957, 3 Vials, 3, 32, and 55m. from Damaturu on Potiskum Road, 5.vi.1957, 4 vials, 
30m. from Potiskum on Kano Road, 6.vi.1957, 4m. S. of Ilorin on Oyo Road, 
4.X11.1957, 45m. S. of Jos on Wamba Road, 23.11.1958, 85m. from Makurdi on 
Wamba Road, 24.11.1958, 24m. from Gboko on Makurdi Road, 25.11.1958, 30m. from 
Lokoja on Okene Road, 8.111.1958, 47m. from Lokoja on Kabba Road, 10.11.1958, 
Samaru, near Zaria, 3.1i.1956, 10.ix.1957, Vi, Vii.I959, 12 vials (W. A. Sands) ; 
Bussa, 1.xii.1g61 (J. McMahon). 

SUDAN: Kadugli, I.vii. to xii.1952, 7 vials (R. C. H. Sweeney) ; 100m. N. of 
Malakal, 1952 (R. H. Gunn) ; 20m. W. of Omdurman, 3.1.1963 (J. Cloudsley 
Thompson). 


96 W: Al SANDS 


UGANDA : 40m. from Moroto on Soroti Road, 12.x.1952 (W. A. Sands) ; 3m. from 
Moyo on Arua Road, 22.11.1952 (W. Wilkinson). 

A total of 187 nest series were examined and all material is in the British Museum 
(Natural History) unless otherwise stated. 


Map 21. Tvrinervitermes geminatus. Vegetation types : 


8, 16, 17, 20, 21, 25, and possibly 
also 31. 


ETHIOPIAN NASUTITERMITINAE 97 


T. geminatus (Wasmann) has previously been recorded as West African in distri- 
bution (Snyder, 1949). The amalgamation of the species listed in the synonymy, 
and much new material, has made it clear that it extends from the extreme west coast 
of Africa, across to the foothills of the Ethiopian Massif, in the Guinean, Sudan, and 
Sahel savannah-woodland vegetation zones (Keay ef al., 1959, types 8, 16, 17, 20, 
and 25). It is, in fact, the most abundant species in these areas, where it builds a 
small mound from a few inches to two or three feet in diameter and height. The 
biology of this species has been described in detail under its better-known synonym, 
T. ebenerianus Sjstedt, in earlier papers (Sands, 1961, 1961a, 1963). 


Trinervitermes gratiosus (Sj6stedt) 
(Text-figs. 192, 235, 236, 250, 251, 276-281; Map 22) 


Eutermes (Trvinervitermes) gratiosus SjOstedt, 1924 : 42. Type locality : Conco, Luluabourg. 

Eutermes (Trinervitermes) carbo Sjéstedt, 1924 : 42. Type locality : Conco, Luluabourg. 
syn. n. 

Trinervitermes gratiosus (SjOstedt) Sjéstedt, 1926 : 349. 

Tyinervitermes carbo (SjOstedt) Sjéstedt, 1926 : 351. 

Imago. Head capsule, central part of frons and vertex pale brown to brown ; vertex with 
three diverging yellow-brown streaks, middle one dilated terminally immediately behind fon- 
tanelle ; postclypeus, labrum, anterior margin of frons behind postclypeus, around antennal 
socket, and a triangular area behind eye paler, yellow brown. Antennae yellow-brown. Pro- 
notum yellow, meso- and metanota yellow with posterior margins yellow-brown, other thoracic 
sclerites pale yellow, legs yellow, bases of tibiae shaded darker, yellow-brown. Abdominal 
tergites yellow-brown ; sternites yellow, yellow-brown round stigmata. Wings, membrane 
translucent pale yellow-brown, subcosta yellow, and radius sector yellow, partially outlined 
yellow-brown, rest of venation narrowly yellow-brown. 

Posterior margin of head behind eyes much less than semi-circular, slightly sinuate immed- 
iately behind eyes ; fontanelle distinct, pale Y- or inverted A-shaped, lateral arms sometimes 
somewhat dilated terminally ; eyes oval, prominent in proportion to own width, and medium 
sized relative to head width, W/E index 2-8—3-1 ; ocelli medium sized, separated from eyes by 
one-fifth to two-fifths own diameter ; postclypeus inflated, width 2-o-2-3 times length, anterior 
margin slightly concave, posterior convex, rounded ; molar plate of right mandible with 8-9 
ridges ; antennae, among eleven specimens examined, four had 15 segments, five 16, and two 17, 
proportions of basal segments variable. 

Pale coloured “‘scent’’ gland on anterior margin of fifth abdominal sternite of 2 large, 
1-15mm. wide, 0-38mm. long ; cerci of 2 shorter than or equal to width across base, conical. 

Pilosity of vertex fine, silky, sparse, and of uneven length, not forming a “‘ pelt ’’ or ‘“‘ mat’”’, 
head capsule appears more shining than in many species. 

Measurements (11 specimens from 5 localities) in millimetres. 


Range Mean 
Head width across eyes 1-80-1-96 : : 1:89 
Greatest diameter of eye 0:58-0:68 : : 0-64 
Ocellus ; : : 0:20-0:22 X 0:23-0:29 . - 0-21 X0227) 
Ocellus toeye . : 0:04-0:08 : : 0-06 
Width of pronotum . 1-65-1°85 : : 1-79 
Length of pronotum . I-00-1-18 . : T6100 
Length of hind tibia. 2°32-2°50 é - 2°36 


Length of fore wing. 17°30—23:20 - : 20°79 


98 W. A. SANDS 


Soldiers. Major soldier : head capsule orange-yellow to chestnut brown, nose ferruginous 
orange to pitch black with very short red tip. Antennae orange-yellow to orange. Pronotum 
yellow-brown to brown, meso- and metanota yellow to yellow-brown, legs yellow. Abdominal 
tergites yellow-brown to brown, sternites pale yellow to yellow. 

Head capsule in plan view short oval to circular, nose weakly conical. In profile distinctly 
concave, with back of head rising from base of nose in front of antennal socket, and rounded ; 
nose distinctly shorter than rest of head capsule, measuring from hind margin of antennal 
socket, angle of nose cone, 5—12 degrees ; antennae 13-14 segmented, two out of three specimens 
with 14, many of those with 13 have basal segments partially subdivided, proportions variable. 
Head setae confined to nose tip, these few, short, and fine apart from 4 apical setae. 

Abdominal tergites apart from posterior one or two almost devoid of setae ; sternites with 
sparse scattered short backwardly directed setae about half length of longer setae on posterior 
margins, approximately in longitudinal rows, 2 or 3 on each side of mid-line, directed down- 
wards and slightly forwards. 

Minor soldiers : colour as major soldiers. Antennae 12-13 segmented, proportions of basal 
segments variable. 

Measurements (Major soldiers, 64 specimens from 21 localities, minor soldiers, 14 specimens 
from 8 localities) in millimetres. 


Mayor Soldier Range Mean 
Head length to tip of nose. 2-II-2°93 ‘ : : : 2°47 
Head capsule width : : 118-181 : : : : 1°49 
Depth of head capsule . ; 0:86-1:25 5 : : é I-07 
Width of pronotum : : 0-61-0°84 ; : : : 0-72 
Length of pronotum : : 0:29-0°43 : : : ; 0:36 
Length of hind tibia ; : 1-54-1-96 : , : ; 1°72 

Minor Soldier Range Mean 
Head length to tip of nose. I-61-1-94 E : : : 1-80 
Head capsule width : : 0-75-1:06 : ; : : 0-89 
Depth of head capsule . s 0:57-0:82 : : : : 0:66 
Width of pronotum : : 0:47-0°61 : 3 : ; 0°54 
Length of pronotum : : 0°22-0°25 : : : ‘ 0:24 
Length of hind tibia : : 1-22-1:68 ‘ I-41 


’ 


Variation. The soldier caste is not always clearly divided into “‘ major ’’ and “‘ minor ”’ 


categories, intermediates being found in the largest colonies. 


Sj6stedt (1924) in describing T. gvatiosus referred only to the major soldier, 
although the material from the type colony included the imago which he described 
subsequently (1926). The lectotype of this species therefore should be selected from 
among the soldier syntypes. 7. carbo was described immediately after T. gratiosus 
on the same page of the same paper, from specimens from the same locality. In this 
case the imago alone was described, and the soldiers stated to be very near 7. 
gratiosus. The description of the imago is clearly not a Tvinervitermes, and the 
specimens were identified by Dr. A. E. Emerson (in unpublished notes) as Cubitermes 
minitabundus (Sjéstedt). Sjéstedt placed the species in Tvinervitermes on the basis 
of the soldier caste, however, and the lectotype would therefore again be selected 
from among the soldier syntypes. 

The resemblance between T. gratiosus and its partially sympatric congener T. 
bettontanus was discussed in an earlier paper (Sands, 1957 : 19). It also resembles 
the ‘‘ West African ’’ T. oeconomus and shorter-nosed paler soldiers of T. trinervius. 
T. oeconomus is distinguishable in the imago by the more pilose head capsule, less 


ETHIOPIAN NASUTITERMITINAE 99 


Map 22. Trinervitermes gratiosus. Vegetation types: 20, 24, 25, 26, and 27. 


inflated postclypeus, longer 9 cerci, and ocelli nearer the eyes. The differing dis- 
tribution is the only consistent feature on which to separate the soldiers, though the 
majority of 7. gratiosus are darker than T. oeconomus and paler than T. trinervius, 
with a shorter nose than either. 


Type Material. Conco: Luluabourg, no date (P. Callewaert), syntype imagos 
and soldiers, type colony, T. gratiosus (Sjéstedt), Naturhist. Riksmus., Stockholm, 


oo W. A. SANDS 


and A.M.N.H. (others in Mus. Tervuren.) ; Luluabourg, no date (P. Callewaert), 
syntype soldiers, type colony, T. carbo (Sjéstedt), A.M.N.H. (others in Naturhist. 
Riksmus., Stockholm.) 


Other Material (additional to that listed in Sands, 1957). CoNnco: Gihinga, 
14.vil.1g41 (H. Bredo) ; Rwindi River, no date (H. Kirby), A.M.N.H. 

ANGOLA : Sao Paola de Loanda, 22.11.1913 (F. Silvestri), A.M.N.H. ; 4m. S. of 
Luanda, 23.vii.1949 (G. R. Gradwell, D. Snow), B.M.(N.H.). 

KENVA*: “1°25 Si¢g5° 10” E., to n° 38°S., 35° 27” BE, i948 (We. Wee 
A.M.N.H. 

A total of 24 nest series were examined. 

It has been noted (Sands, 1957) that this species tolerates drier conditions than 
many, and the additional records are from relatively dry areas also. 

The type locality, Luluabourg, is well within the forest-savannah mosaic zone 
(Keay et al., 1959, type 8) on the southern border of the rain forest, and thus appears 
to be in more moist conditions, but there are in that area many patches of dry grass 
steppe on Kalahari sand, and other drier savannah types (Keay et al., 1959, types 
20 and 24). T.gvatiosus appears to occur in these, though absent from the surround- 
ing moister savannah woodland, where T. rhodesiensis is the common species. 

Sj6stedt (1926) recorded T. carbo from Haut Uélé, but this must be a misidentifi- 
cation since T. gratiosus is an East and Central African species. 


Trinervitermes occidentalis (Sjdstedt) 
(Text-figs. 237-240, 252, 253, 282-289 ; Map 23) 


Eutermes occidentalis Sj6stedt, 1904 : 93. Type locality : PoRTUGUESE GUINEA, Bissau. 

Termes (Eutermes) occidentalis (Sj6stedt) Desneux, 1904a : 44. 

Eutermes (Trinervitermes) occidentalis SjOstedt ; Holmgren, 1912 : 64. 

Trinervitermes occidentalis (Sj6stedt) Sjéstedt, 1926 : 332. 

Trinervitermes auriterrvae SjOstedt, 1926a : 3. Type locality : Guana, Keta. syn. n. 

Trinervitermes maudanicus Sjostedt, 1926b : 156. Type locality : Conco, Haut Uélé, Mauda. 
syn. n. 

Nasutitermes (Tvinervitermes) bettonianus subsp. sulciceps Emerson, 1928 : 493. Type locality : 
Conco, Garamba. syn. n. 

Nasutitermes (Tvinervitermes) lutzi Emerson, 1928 : 494. Type locality : Conco, Niangara. 
syn. n. 

Trinervitermes bettonianus subsp. sulciceps (Emerson) Snyder, 1949 : 323. 

Trinerviterymes lutzi (Emerson) Snyder, 1949 : 328. 

Trinervitermes auritervae SjOstedt ; Sands, 1957 : 16. 


Imago. Head capsule brown to chestnut brown ; vertex with two or three indistinct diverg- 
ing yellow-brown patches or streaks, or sometimes one median pale patch, postclypeus, labrum, 
areas in front of ocelli and small area behind each eye paler yellow-brown to brown. Antennae 
yellow. Pronotum yellow to yellow-brown, meso- and metanota, other thoracic sclerites and 
legs yellow. Abdominal tergites brown, darker sepia-brown around stigmata ; sternites pale 
yellow to yellow-brown, brown around stigmata. Wings translucent pale brown, subcosta and 
radius sector yellow, the latter outlined sepia-brown at base, fading to yellow-brown distally ; 
bright orange streak in membrane behind radius sector ; rest of venation brown. 

Posterior margin of head capsule behind eyes much less than semi-circular, slightly sinuate 
immediately behind eyes ; fontanelle Y-shaped, often slightly depressed posteriorly, length 


ETHIOPIAN NASUTITERMITINAE IOI 


280. 


278. 
281. 


283. 


285. 287. 289. 


290. 


291. 293. 295. 296. 


Fics. 276-296 : Trinervitermes, soldier head capsule, side and plan views, and nose tip. 
276-281, T. gratiosus ; 282-289, T. occidentalis : 290-296, T. oeconomus. 


102 W. A. SANDS 


two to four times breadth, lateral arms short or very short, sometimes dilated terminally ; 
frons with two small diverging depressions in front of fontanelle ; eyes commonly oval, 
moderately to distinctly prominent in proportion to own diameter, and medium-sized relative 
to head width, W/E index 2-6-3-0 ; ocelli medium sized, near circular to short oval, separated 
from eyes by one-tenth to one-fifth own least diameter ; postclypeus moderately inflated, 
width 2-2—2-6 times length, anterior margin straight or weakly concave, posterior margin convex, 
often more rounded towards outer corners, sometimes slightly obtusely angular in middle ; 
molar plate of right mandible with 7 ridges ; antennae with 15 segments, only one specimen 
recorded with III partially subdivided, usually III longer up to twice II, IV longer than II and 
shorter than III, V, and VI subequal, about as long as II but stouter. 

Pale coloured “‘scent’’ gland on anterior margin of fifth abdominal sternite of 2 small, 
0:56-0-64mm. wide, o-11-0-13mm. long ; cerci of 2 as long as, or longer than basal width, 
conical or slightly mamilliform. 

Pilosity of vertex of uneven length, not forming a “ pelt’ or “‘mat”’. 

Measurements (22 specimens from 4 localities) in millimetres. 


Range Mean 
Head width across eyes 1-48-1°75 : : 1°57 
Greatest diameter of eye 0°53-0'63 : 3 0°57 
Ocellus 3 : ‘ 0:19-0:25 X 0°24-0°:29 . : O'2I X 0:26 
Ocellus toeye  . : 0:03-0:04 : : 0:03 
Width of pronotum . I -33-1 -68 ; P 1°48 
Length of pronotum . 0:80-1:08 . : 0:93 
Length of hind tibia. 1-98—2°38 : : 22M 
Length of fore wing. 14*10—20°30 : : 17:07 


Soldievs. Major soldier: head capsule orange-yellow to ferruginous orange, commonly 
orange, nose ferruginous-orange to chestnut brown with red tip, commonly ferruginous. Pro- 
notum, meso- and metanota orange yellow, antennae and legs yellow. Abdominal tergites 
yellow-brown to brown, sternites pale yellow. 

Head capsule in plan view slightly to distinctly “shouldered ’’ behind antennae, rounded 
posteriorly, length equal to or slightly more than width, posterior margin often slightly sulcate 
in middle ; nose weakly conical, often with curved taper increasing towards base. In profile 
variable from near straight to distinctly concave with back of head rising from behind antennal 
socket and rounded ; nose equal in length to shorter than rest of head capsule, usually with 
curved taper increasing from near cylindrical tip to distinctly conical base (resembling an 
inverted trumpet), fontanelle large, o-o5-o:lomm. in diameter ; antennae 12-14 segmented, 
87% with 13, one recorded with 14, proportions of basal segments variable, usually III longer 
up to twice II and IV, V and VI longer than these, shorter than III. Prominent head setae, 
confined to densely hairy nose tip, dorsal surface of head capsule only with very sparse micro- 
scopic short setae. Middle abdominal tergites almost devoid of setae, those on posterior two 
short and sparse. Sternites with short, sparse backwardly directed setae, one-sixth to one-third 
length of longer setae on posterior margins approximately in longitudinal rows, 2 or 3 on each 
side of mid-line, directed downwards or slightly forwards. Minor soldiers : colour as major 
soldiers except nose usually chestnut brown. Antennae 12 segmented. 

Measurements (Major soldiers, 92 specimens from 24 localities, minor soldiers 26 specimens 
from 20 localities) in millimetres. 


Mayor Soldiers : Range Mean 
Head length to tip of nose. I-Q0-2°50 ‘ 3 : ; 2°21 
Head width . : . P 105-1 °54 : , : : 1-26 
Depth of head capsule . : 0:78-1:07 , 2 2 : 0-92 
Width of pronotum : : 0:50-0°68 : : : ‘ 0°58 
Length of pronotum : : 0:23-0:29 : ‘ : : 0:25 


Length of hind tibia ; - I-20-1°55 ‘ : ; ‘ Te37, 


ETHIOPIAN NASUTITERMITINAE 103 


Minor Soldiers : Range Mean 
Head length to tip of nose. 1-35-1°79 2 ; - : I-51 
Head width . : : 2 0°55-0°80 . . ‘ ; 0-64 
Depth of head capsule . : 0-45-0°64 : : : - 0°52 
Width of pronotum . : 0°35-0°47 : 3 : : O-41 
Length of pronotum : - 0:15-0:22 - : ‘ F (ols te) 
Length of hind tibia : - 1-08-1-33 ; : - : I-19 


Variation. In the imago caste this is comparatively slight, whereas in the soldier there is 
considerable variation in profile view, both in degree of straightness, and in the shape of the nose. 
The latter arises mainly from differences in the point at which the curved taper begins, resulting 
in approximately conical, inverse trumpet-shaped, or almost cylindrical forms : there is some- 
times also a slight hump at the base of the nose. The vestigial soldier mandibles occasionally 
bear small points. This appears to be a species more tolerant of higher altitudes up to 5,000 feet, 
and these specimens are larger than those from lower down. 


The imago is readily distinguished from sympatric congeners in nearly all cases by 
the 15 segmented antennae, those few 7. togoenstzs which have only 15 segments 
usually having III partially subdivided on at least one side, and shorter female 
cerci. The East and Central African species T. rhodesiensis and T. rapulum are not 
easily separable from T. occidentalis. T.rhodesiensis usually lacks small depressions 
on the frons, 50°% of specimens have 16 segmented antennae, and the eyes are 
relatively smaller in many cases. The characters given in the key are sufficient in 
the majority of cases to distinguish T. rapulum. In the soldier caste, the large 
fontanelle and hairy nose tip distinguish 7. occidentalis from the other two species. 
The latter character was not recognized in an earlier paper (Sands, 1957) with 
the result that specimens of T. rhodesiensis were attributed in error to T. lutzi, now 
found to be a synonym of T. occidentalis. 


A lectotype has been selected for the junior synonym T. auriterrae as indicated 
below. 


Type Material. PoRTUGUESE GUINEA : Bissau, 4.x.1898 (H. Ehrhardt), syntype 
g and 2 imagos, T. occidentalis (Sjostedt), A.M.N.H. (Other syntypes in Mus. 
Hamburg and Naturhist. Riksmus., Stockholm.) 


GHANA: Keta, 1926 (W. H. Patterson), lectotype soldier and paralectotypes, 
T. auriterrae Sjéstedt, B.M.(N.H.) (other paralectotypes from type colony in the 
Naturhist. Riksmus., Stockholm). 


Conco: “Haut Uélé”’ Mauda, 1925 (H. Schouteden). From the dates of 
Schouteden’s travels in 1925, Mauda must be near to Arebi since he was in both 
localities on the same day. Arebi is in the Kibale—Ituri district, not the adjoining 
Haut Uélé as stated by Sjéstedt. Syntype soldier, type colony, 7. maudanicus 
Sjostedt, B.M.(N.H.) (other syntypes in Mus. Tervuren, Naturhist. Riksmus., 
Stockholm, and A.M.N.H.) ; Garamba, vii.1g12 (H. Lang and J. P. Chapin), para- 
type soldiers, type colony, T. bettonianus s. sp. sulcicepbs, Emerson, A.M.N.H. 
(Holotype in A.M.N.H.) ; Niangara, 3° 40’ N., 26° 50’ E., 20.v.1913 (H. Lang, 
J. P. Chapin), paratype soldiers, type colony, T. lutzi Emerson, B.M.(N.H.) and 
A.M.N.H. (Holotype soldier in A.M.N.H.) 


104 W. A. SANDS 


Other Material. Ivory Coast: Dabou, 50km. W. of Abidjan, 1962-4, 8 vials 
(P. Bodot). 

GHANA : 2m. W. of Adafon, 7.xi.1955 (W. Belfield) ; 10m. N. of Yeji on Tamale— 
Kumasi Road, 25.11.1959, 12m. S. of Tamale on Kumasi Road, 27.11.1959, 35m. from 
Tamale on Yendi Road, 4.iii.1959, 5m. from Nakpanduri on Gambaga Road, 
6.i11.1959, 20m. from Bolgatanga on Tamale Road, 9g.i11.1959, Navrongo, 12.111.1959, 


Map 23. Tvinervitermes occidentalis. Vegetation types: 8, 16, 17, and 20. 


ETHIOPIAN NASUTITERMITINAE 105 


2 vials, 31m. from Navrongo on Tumu Road, 13.iii.1959, 11m. from Tumu on 
Navrongo Road, 15.iii.1959, 6m. from Tumu on Lawra Road, 16.iii.1959, gm. N. of 
Lawra on Nandom Road, 18.iii.1959, 6m. N. of Wa on Lawra Road, 19. iii.1959, 24 
and 52m. S. of Wa on Bole Road, 20.iii.1959, 2 vials, 3m. from Larabanga on Bole 
Road, 22.iii.1959, 22m. from Larabanga on Tamale Road, 24.iii.1959 (W. A. Sands). 

NIGERIA : Western Region ; Ibadan, 1937 (F. Toovey), A.M.N.H.; Lagos, 
Lighthouse Beach, 9.iv.1957 (W. Wilkinson). Eastern Region ; Enugu, ii.1955 
(W. V. Harris) ; Onitsha, 5.iv.1957 (W. Wilkinson) ; 7 and 12m. from Enugu on 
Onitsha Road, 2-3.111.1958, 2 vials, Enugu, xii.1958 (W. A. Sands). Northern 
Region ; 28 and 44m. from Sokoto on Argungu Road, 23.xi.1956, 3 vials, 26m. from 
Minna on Zungeru Road, 20.xii.1956, Mokwa, 21.xii.1956, 2 vials, 39m. from Abuja 
on Bida Road, 24.xii.1956, Diko, 1om. W. of Abuja, 27.x11.1956, 22m. from Kaduna, 
on Kachia Road, 30.1.1957, 5m. from Jos on Bukuru Road, 7.ii.1957, 5m. from 
Bukuru on Vom Road, 7.1i.1957, 34m. from Yola on Jalingo Road, 14.v.1957, 
114-125m. S.W. of Jalingo on Takum Road, via Beli, 18.v.1957, 2 vials, Adamawa, 
Tiba, tom. N.W. of Tibak, 24.v.1957, 5 vials, and Gangoro 25.v.1957, 32m. from 
Damaturu on Potiskum Road, 5.vi.1957, Samaru near Zaria, 10.ix.1957, and vi-vil. 
1959, 3 Vials, 4m. from Gboko on Oturkpo Road, 26.ii.1958, 2 vials, and 47m. from 
Lokoja on Kabba Road, 10.1i1.1958 (W. A. Sands). 

CENTRAL AFRICAN REPUBLIC : Haut Mbomu, Djema, 6° 5’ N., 25° 12’ E., iii.1948 
(N. A. Weber), A.M.N.H.. 

Conco : Garamba, 25.i.1950 (H. de Saeger), Inst. des Parcs Nat. du Congo, 
Brussels. 

UGANDA : Mbale, 6.xi.1937, Serere, x.1948 (W. V. Harris). 

A total of 68 nest series were examined, and all material is in the British Museum 
(Natural History) unless otherwise stated. 


T. occidentalis is one of the typically ‘‘ Guinean ”’ species, occurring in savannah 
vegetation zones ranging from moist savannah patches in forest zones, to the 
moderately dry “‘ Sudan ”’ savannah from the west coast across to the Sudan and 
Uganda (Keay et al., 1959, types 8, 16, 17, and 20). Its nest and feeding habits 
have been described elsewhere (Sands, 1961, 1961a). Though never abundant in 
any habitat and usually very scarce compared with the other four sympatric species, 
it is thought to be one of the most primitive of the genus, sharing with T. rapulum 
the habit of feeding on woody material in addition to the more usual grass and leaves. 


Trinervitermes oeconomus (Tragardh) 
(Text-figs. 241, 242, 254, 255, 290-296, 384 ; Map 24) 


Eutermes oeconomus Tragardh, 1904 : 23. Type locality : SupaN, Kaka. 

Eutermes mobilis Sjostedt, 1904 : 91. Type locality : Marr, Alahina. syn. n. [ex syn. T. 
trinervius (Rambur) ; Snyder, 1949 : 332]. 

Termes (Eutermes) oeconomus (Tragardh) Desneux, 1904a : 44. 

Termes (Eutermes) mobilis (SjOstedt) Desneux, 1904a : 43. 

Eutermes tchadensis Sj6stedt, 1911b : 14. Type locality : Cuap, “ Tchari-Tchad ”’ [probably 
Ft. Lamy area]. syn. n. 


106 W. A. SANDS 


Eutermes (Tvinervitermes) oeconomus Tragardh ; Holmgren, 1912 : 64. 

Eutermes (Tvinervitermes) mobilis Sjostedt ; Holmgren, 1912 : 65. 

Eutermes (Trvinervitermes) tchadensis SjOstedt ; Holmgren, 1912 : 68. 

Eutermes schubotzianus Sjéstedt, 1914b : 93. Type locality : CHap, Ft. Crampel. syn. n. 
Eutermes (Trinervitermes) schubotzianus Sjostedt ; Hegh, 1922 : 693. 

Trinervitermes oeconomus (Tragardh) Sjostedt, 1925a : 73. 

Trinervitermes tchadensis (SjOstedt) Sjéstedt, 1926 : 353. 

Trinervitermes schubotzianus (Sjéstedt) Sjéstedt, 1926 : 383. 

Trinervitermes mobilis (SjOstedt) Sj6stedt, 1926 : 353. 


Imago. Head capsule, central part of frons and vertex brown to dark chestnut brown ; 
vertex with three diverging yellow-brown streaks ; postclypeus, and area surrounding eyes, 
extending to a point behind eyes and to mandible base in front, yellow-brown ; labrum and 
antennae yellow to yellow-brown. Pronotum yellow to yellow-brown, meso- and metanota, 
other thoracic sclerites and legs yellow. Abdominal tergites brown to dark chestnut brown, 
slightly darker round stigmata, with yellow to yellow-brown border, and slightly paler towards 
lateral corners of front margin ; sternites yellow to yellow-brown, brown round stigmata. 
Wings translucent pale brown, subcosta and radius sector yellow, paler distally, latter outlined 
brown at base, membrane between them shaded brown for short distance from suture ; mem- 
brane behind radius sector with or without orange-yellow streak ; rest of venation narrowly 
yellow-brown to brown. 

Posterior margin of head capsule behind eyes much less than semi-circular, slightly sinuate 
immediately behind eyes ; fontanelle distinct, usually Y-shaped with lateral arms dilated 
terminally, sometimes slightly depressed or occluded posteriorly, length approximately twice 
breadth at base of lateral arms ; eyes slightly angularly oval and moderately prominent in 
proportion to own diameter, medium-sized relative to head width, W/E index 2-6-3-0 ; ocelli 
medium sized to large, separated from eyes by one-fifteenth to one-fourth own least diameter ; 
postclypeus moderately inflated, width 2-3 to 2-5 times length, anterior margin nearly straight, 
posterior margin convex, rounded ; molar plate of right mandible with 7—8 ridges ; antennae 
17 segmented, II, IV, and V subequal, VI slightly and III distinctly longer than these. 

Pale coloured “‘scent’’ gland on anterior margin of fifth abdominal sternite of 2 large, 
0:97-1:07mm. wide, 0-26—0-41mm. long ; cerci of 2 longer than wide across base, conical. 

Pilosity of vertex of varying length, setae not forming a “‘ pelt ’’ or “‘ mat ”’ (Text-fig. 384). 

Measurements (40 specimens from 9 localities) in millimetres. 


Range Mean 
Head width across eyes I-72-1°95 : : 1-82 
Greatest diameter of eye 0:58-0°74 ¢ : 0-66 
Ocellus : : : O0°19-0'25 X 0:25-O'31  . : O22, X 0°28 
Ocellus toeye  . : 0:02-0:05 : : 0:03 
Width of pronotum . 1:58-1:95 . : 1-69 
Length of pronotum . 0-98-1'18 : : I-09 
Length of hind tibia. 2:30-2:68 ; : 2:48 
Length of fore wing. 16-10—20:80 : : 18-61 


Soldiervs. Major soldier ; head capsule yellow to ferruginous, commonly orange-yellow, nose 
orange to chestnut brown with reddish tip. Antennae and pronotum orange-yellow, meso- and 
metanota, and legs yellow. Abdominal tergites brown, sternites pale yellow. 

Head capsule in plan view short oval or circular, nose conical. In profile near straight to 
distinctly concave, with back of head rising from above antennal socket, and rounded ; nose 
equal in length to distinctly shorter than rest of head capsule, measuring from hind margin of 
antennal socket, angle of nose cone 5-18 degrees ; antennae 13-14 segmented, only one in 
twenty with 13, proportions of basal segments variable, in 13 segmented forms, II and IV sub- 
equal, shorter than V and VI which are also subequal, III twice II and IV, in 14 segmented, 
III, V and VI subequal, IV slightly longer and II slightly shorter. Head setae confined to four 


ETHIOPIAN NASUTITERMITINAE 107 


at nose tip with smaller setae behind them, sometimes very sparse ; dorsal surface of head 
without prominent setae. Middle abdominal tergites with very sparse scattered minute setae, 
scarcely visible in profile, only slightly longer on posterior tergites. Sternites with short back- 
ward directed scattered setae, and longer downward or slightly forward directed setae on 
posterior margins, arranged approximately in 6 longitudinal rows, 3 or more times length of 
shorter setae. 

Minor soldier ; head capsule orange-yellow to reddish chestnut brown, nose ferruginous 
orange to brownish black with red tip. Antennae 13-14 segmented, proportions of basal 
segments variable. 

Measurements (Major soldiers, 60 specimens from 20 localities, minor soldiers, 11 specimens 
from 9 localities) in millimetres. 


Major Soldiers : Range Mean 
Head length to tip of nose. 1:95-2°65 : ’ : ; 2°33 
Head width . - : - I-13-1°63 ; ; : é 1°39 
Depth of head capsule . ; 0:80-1:08 : 3 2 : 0:97 
Width of pronotum : : 0°55-0°73 : : ; : 0°65 
Length of pronotum ; ; 0:23-0°36 ; : : s 0-30 
Length of hind tibia : : 1-40-1 -82 : : : : 1°64 

Minor Soldiers : Range Mean 
Head length to tip of nose. I-50-1 -82 . : ‘ : 1:66 
Head width . P é ; 0:64-0°82 : ; P : 0:74 
Depth of head capsule . é 0+50-0°64 : ; : < 0°57 
Width of pronotum : ; 0-43-0°50 : : : ; 0-46 
Length of pronotum : : 0:18-0:25 ; d : ‘ 0:22 
Length of hind tibia : , I-20-1°50 : : ; - 1°40 


T. oeconomus is similarly variable to its sympatric congeners, with which it is 
easily confused. It is distinguishable in the 9 imago from 7. geminatus by the 
larger “‘ scent ’’ gland on the fifth sternite and the longer cerci, and from both this 
and T. trinervius by the uneven vertex pilosity. In the soldier caste, T. trinervius 
is much darker in colour and has few nose setae. Most 7. geminatus are also darker, 
with slightly more prominent abdominal setae, but indistinguishable specimens do 
occur. Larger, paler specimens of 7. togoensis have been found to occur at greater 
altitudes, and these sometimes have 14 segmented soldier antennae, thus also 
becoming indistinguishable from 7. oeconomus. Of the allopatric species, the 
closest is T. gratiosus, in which the imago has a more inflated postclypeus, ocelli 
usually further from the eyes, and shorter female cerci. The paler coloured soldiers 
of this species are almost indistinguishable from 7. oeconomus, except that the nose 
tip of the latter has more setae in the majority of specimens. 

The synonymy of this species is straight-forward apart from 7. mobilis, which 
was described from imagos collected at light. The syntype material consists of 
two species, the larger of which corresponds to the original description, the smaller 
being 7. togoensis. Snyder (1949) synonymised T. mobilis with T. trinervius, 
but the accepted usage of the latter is for the species also named T. posselensis and 
T. carbonarius. This I regard as correct and therefore use the next oldest name, 
T. oeconomus for the species to which the specimens named T. mobilis by Sjéstedt 
also belong. 


108 W. A. SANDS 


A specimen has been selected and labelled by Dr. A. E. Emerson as lectotype for 
the junior synonym, 7. tchadensis, but the designation has not been published 
hitherto. It is given under the type material heading. 

Type Material. Supan: Kaka, iii.rgor (J. Tvdgardh), syntype soldier from type 
colony, T. oeconomus (Tragardh), A.M.N.H. (other syntypes in Naturhist. Riksmus., 
Stockholm). 

Mati: Alahina, (near Toukoto?), vi.1g03 (Kermorgant), syntype imago from 
type series, JT. mobilis (Sjostedt), A.M.N.H. (other syntypes in Mus. Paris and 
Naturhist. Riksmus., Stockholm). 

CuaD : Tchari-Tchad, no date, (no collector), paralectotype soldier from type 
colony, T. tchadensis (Sjostedt), A.M.N.H. (lectotype soldier and other paralecto- 
types, in Naturhist. Riksmus., Stockholm). 

CENTRAL AFRICAN REPUBLIC: Fort Crampel, 6.1.1911 (A. Schubotz), paratype 
soldier from type colony, T. schubotzianus (Sjéstedt), A.LM.N.H. (Holotype soldier 
and other paratype, in Mus. Hamburg, other paratypes in Naturhist. Riksmus., 
Stockholm.) 


Other Material. SIERRA LEONE: Samaia, 24.v.1948 (Ff. A. Squires). 

Ivory Coast : Toumodi, 120km. N.W. of Abidjan, 1962-1964, 3 vials (P. Bodot). 

GHANA : Cape Coast Castle, no date (G. Williams) ; Accra, 26.v—29.v1.1926, 3 
vials (A. W. J. Pomeroy) ; Nungua, 16.1. and 21.vi.1956, 3 vials, Achimota, 19.1.1953 
and 6.1.1956, 3 vials (W. Belfield) ; Accra, 16.11.1959, 113m. from Accra on Kumasi 
Road, 18.11.1959, 2 vials, Ejura, 21.11.1959, 33-60m. N. of Ejura on Tamale Road, 
24.11.1959, 2 vials, 1om. N. of Yeji on Tamale Road, 25.11.1959, 12m. S. of Tamale 
on Kumasi Road, 26.11.1959, 5 and 16m. from Daboya on Tamale Road, 28.11.1959, 
2 vials, 30m. from Tamale on Larabanga Road, 3.11.1959, 35m. from Tamale on 
Yendi Road, 4.111.1959, 25m. S.E. of Gambaga on Yendi Road, 6.iii.1959, Volta 
Bridge on Bolgatanga—Bawku Road, 8.ii1.1959, Navrongo, 12.111.1959, 29m. from 
Navrongo on Tumu Road, 13.11.1959, 12m. N. of Navrongo, 14.11.1959, 11m. from 
Tumu on Navrongo Road, 15.11.1959, 40m. from Tumu on Lawra Road, 16.1ii.1959, 
om. N. of Lawra on Nandom Road, 18.iii.1959, 24 and 52m. from Wa on Bole Road, 
20.11.1959, 3 vials, 3m. from Larabanga on Bole Road, 22.i11.1959, 12m. from Accra 
on Takoradi Road, 7.iv.1959 (W. A. Sands). 

NIGERIA: Eastern Region, Enugu, 12.xi.1955 (W. V. Harms) ; Abakaliki, 
28.v.1957 (W. Wilkinson) ; Northern Region ; Yashi, 3.1x.1954, 2 vials, Kano, 
4.1x.1954 (B. J. McNulty) ; 43m. E. of Kano, Gaya River, 17.xi.1956, 31m. W. of 
Kano on Katsina Road, 18.xi.1956, Katsina, 18.xi.1956, Kaura Namoda, 21.xi.1956, 
71m. from Gusau on Sokoto Road, 22.x1.1956, 25m. from Sokoto on Gusau Road, 
25.X1.1956, 30m. N. of Sokoto on Ilela Road, 26.xi.1956, Samaru, near Zaria, 3.X1. 
1956, 2 vials, and 14.v.1959, 18m. from Kaduna on Zaria Road, 8.x11.1956, 2 vials, 
33m. from Kaduna on Zungeru Road, 18.x11.1956, 45m. from Zungeru on Kaduna 
Road, 19.xii.1956, 29m. from Minna on Zungeru Road, 20.xii.1956, 2 vials, 16m. 
from Zungeru on Bida Road, 21.xii.1956, 3 vials, 41m. from Mokwa on Bida Road, 
22.xi1.1956, 39m. from Abuja on Bida Road, 24.xii.1956, 22m. from Kaduna on 


ETHIOPIAN NASUTITERMITINAE 109 


Kachia Road, 30.i.1957, Kudaru on Zaria—Jos Road, 5.ii.1957, 4 vials, 5m. from Jos 
on Bukuru Road, 6.1i.1957, 2 vials, Naraguta, near Jos, 10.ii.1957, 16 and 40m. from 
Jos on Bauchi Road, 11.ii.1957, 4 vials, 25m. from Bauchi on Jos Road, 13.ii.1957, 
50m. from Kaduna on Jos Road, 15.ii.1957, Afaka, 16m. from Kaduna on Zungeru 
Road, 28.11.1957, 4 vials, 24-32m. from Jos on Kaduna Road, 3.iv.1957, 4 vials, 


a ht = aa ae eye ae es Ep PEL Pk 


e 


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Map 24. Trinervitermes oeconomus. Vegetation types: 8, 16, 17, 20, 21, and 25, and 
possibly also 31. 


I1O W.A. SANDS 


7-39m. from Bauchi on Gombe Road, 7.v.1957, 3 vials, Gombe, and 30m. E. of 
Gombe, 8—9.v.1957, 5 vials, 22m. from Gombe on Numan Road, 10.v.1957, 2 vials, 
40-52m. S.E. of Gombe on Numan Road, I0.v.1957, 5 vials, g2m. from Gombe on 
Numan Road, I1.v.1957, 2 vials, 6m. from Numan on Yola Road, 12.v.1957, 24—34m. 
from Yola on Jalingo Road, 14.v.1957, 4 vials, 54 and 75m. from Yola on Jalingo 
Road, 16.v.1957, 2 vials, 3m. from Jalingo on Yola Road, 17.v.1957, 125m. S.W. of 
Jalingo on Takum Road via Beli, 18.v.1957, 35 and 80m. from Yola on Biu Road, 
28.V.1957, 2 vials, 15 and 55m. N. of Biu on Damaturu Road, 29.v.1957, 2 vials, 
Maiduguri, I.vi.1957, 43m. from Maiduguri on Fort Lamy Road, 3.vi.1957, 32m. 
from Damaturu on Potiskum Road, 5.vi.1957, 2 vials, 4m. S. of Ilorin on Oyo Road, 
4.X11.1957, 45m. S. of Jos on Wamba Road ,23.11.1958, 2 vials, 12m. from Keffi on 
Makurdi Road, 24.11.1958, 30m. from Lokoja on Okene Road, 8.111.1958, Lokoja, 
g.11.1958 (W. A. Sands) ; Bussa, 1.xii.1961 (J. McMahon). 

Conco : Garamba Nat. Park, 29.ix.1951 (H. de Saeger), Inst. des Parcs Nat. du 
Congo, Brussels. 

SuDAN : Renk, 4.vii.1gog (H. H. King) ; Darfur Province, Kulme, 12° 36’ N., 
23° 27' B., 20:vi.19g21 (H. Lynes) ; Port Sudan, virx939 (7. M. Telford) > Aan 
Keilak, 30.vi.1952, Kadugli, 22.vii.1952 (R. C. H. Sweeney). 

UcanpDA : additional to those recorded in Sands (1957) ; Serere, 17.vii.1939 
(P.R.S.), Fort Portal, ii.1948 (N. A. Weber), A.M.N.H. ; 15m. from Kampala on 
Masindi Road, 7.x1.1955 (R. M. C. Williams). 

Lisya? : “ Tripoli’’, no date (Walker). 

A total of 146 nest series were examined and all material is in the British Museum 
(Natural History) unless otherwise stated. 


T.oeconomus was recognized by Sjostedt (1925a) as occurring from the Sudan across 
to Ghana, and further records have extended the range to include vegetation types 
ranging from forest-savannah mosaic, to the Sahel and even in the case of the Port 
Sudan record, subdesert steppe (Keay ef al., 1959, types 8, 16, 17, 20, 25, and 31). 
The specimen recorded as T. trinervius by Walker (1853) from “‘ Tripoli’’ has been 
examined, and is undoubtedly 7. oeconomus. There must be considerable doubt 
attached to the locality from which it was recorded. According to Capra (1938), no 
other Tvinervitermes has been found in North Africa, and it seems probable that this 
specimen originated in the Sudan. 

The wide distribution of this species may partly be explained by its habit of 
commonly building its low wide mound in shaded or sheltered locations, and often 
occupying parts of mounds of various species of Macrotermitinae. 


Trinervitermes rapulum (Sj6stedt) 
(Text-figs. 193, 297, 298, 308-313, 338-358 ; Map 25) 


Eutermes vapulum Sj6stedt, 1904 : 99. Type locality : TaNGANy1Ka, Tanga. 
Termes (Eutermes) vapulum (Sj6stedt) Desneux, 1904a : 44. 
Eutermes (Trinervitermes) rapulum Sj6stedt ; Holmgren, 1912 : 65. 


ETHIOPIAN NASUTITERMITINAE ILI 


Eutermes (Trinervitermes) dispar form zuluensis Holmgren, 1913d : 340. Type locality : 
ReEpuBtic oF SoutH Arrica, Zululand, Mfolosi Drift, Mkosi. syn. n. 

Eutermes (Trinervitermes) dispar form disparioides Holmgren, 1913 : 342. Type locality : 
ReEpuBLic oF SoutH Arrica, Zululand. Mbhlazinga. 

Trinervitermes zuluensis (Holmgren) Fuller, 1922 : 113 [including Tyinervitermes dispay form 
disparioides Holmgren as a synonym). 

Trinervitermes pretoriensis Fuller 1922: 114. Type locality : ReEPUBLIC OF SoUTH AFRICA, 
Pretoria. syn. n. 

Trinervitermes vapulum (Sjdstedt) Sjéstedt, 1926 : 348. 


Additions to the previous descriptions (Sands, 1957) of both imago and soldier castes are 
necessary. 

Imago. Head capsule, central part of frons and vertex yellow-brown to dark chestnut brown, 
vertex often with median pale patch, pale yellow to yellow-brown ; postclypeus, labrum, 
antennae, and area surrounding eye, ocellus and antennal socket paler, pale yellow to yellow- 
brown. Pronotum yellow to yellow-brown, other thoracic sclerites and legs yellow. Abdominal 
tergites yellow to brown, often darker round stigmata, sternites pale yellow to yellow, sometimes 
brown round stigmata. Wings translucent pale brown, subcosta and radius sector, outlined 
brown at base ; membrane behind radius sector with orange streak ; rest of venation outlined 
brown. 

Posterior margin of head capsule behind eyes much less than semi-circular, slightly sinuate 
immediately behind eyes ; fontanelle rather small, shape varies in f from U- to near Y-shaped, 
2 from Y-shaped to triangular with concave anterior margin, lateral arms short, not dilated 
terminally ; frons usually with two small diverging depressions in front of fontanelle ; eyes 
prominent in proportion to own diameter but not large relative to head width, W/E index 
3:0-3°5 ; ocelli separated from eyes by one-fifth to more than half own least diameter ; post- 
clypeus width 1-9-2-2 times length ; molar plate of right mandible with 7-8 ridges ; antennae, 
70%, of specimens with 15 segments, 30% with 16. 

Pale coloured ‘“‘ scent ’’ gland on anterior margin of 2 fifth abdominal sternite small, 0-56— 
o-71Imm. wide, 0-15—-o-20omm. long ; cerci of 2 as long as or longer than basal width, slightly 
mamilliform. 

Pilosity of vertex of uneven length, not forming a “ pelt ’’ or “‘ mat’. 

Measurements (38 specimens from rr localities) in millimetres. 


Range Mean 
Head width across eyes 1°45-1°83 ; : 1-64 
Greatest diameter of eye 0:47-0:58 ; ; 0°52 
Ocellus : : : 0'15-0°23 X 0:20-0-30_ . ; 0-19 X 0:24 
Ocellus toeye . : 0:03-0:10 ‘ : 0:06 
Width of pronotum I+25-I°70 : é 1°49 
Length of pronotum . 0:78-1:05 : : 0:93 
Length of hind tibia. 1-81—2-20 i : 2:02 
Length of fore wing. 14°30—22-10 . ‘ 17°31 


Soldiers. Major soldier : head capsule yellow to ferruginous orange, nose ferruginous to 
pitch black with red tip, antennae pale yellow to orange-yellow. Pronotum, meso- and meta- 
nota yellow to yellow-brown, legs pale yellow to yellow. Abdominal tergites yellow to brown, 
sternites pale yellow. 

Head capsule in plan view slightly “‘ shouldered ’’ behind antennae, rounded posteriorly, nose 
cylindrical to weakly conical. In profile distinctly concave, with lowest point above or slightly 
in front of antennal socket, rising in a weak curve or nearly straight line to highest point near 
back of head, before downward curve to occiput ; nose shorter than rest of head, measuring 
from hind margin of antennal socket, angle of nose cone o—14 degrees. 

Head setae consist of four at nose tip with a few smaller setae behind them, and occasionally 
scattered sparse minute setae on vertex. Middle abdominal tergites with scattered small setae, 


112 W. A. SANDS 


longer on posterior segments, sternites with short backward directed setae up to one-third 
length of longer downward or slightly forward directed setae on posterior margins, arranged in 
4-6 longitudinal rows. 

Minor soldier : colour as major soldier except nose always dark, antennae 12-13 segmented, 
usually 12. 

Measurements (Major soldiers, 74 specimens from 18 localities, minor soldiers, 9 specimens 
from 6 localities) in millimetres. 


Major Soldiers : Range Mean 
Head length to tip of nose. 1 -85-2°33 : : : i 2:10 
Head width . é : . I-07—-1°50 : : F : I-30 
Depth of head capsule . ‘ 0:75-1:07 : : : : O-9O1 
Width of pronotum : < 0°54—0°71 ; ; 3 , 0-61 
Length of pronotum ; : 0:22-0:29 . : - - 0:25 
Length of hind tibia 3 3 I-07—-1°53 : : : . 1:25 

Minor Soldiers : Range Mean 
Head length to tip of nose. 1°43-1-62 : ; : : I-49 
Head width . ; ; : 0:57-0:64 : : : : 0-60 
Depth of head capsule . : 0°45-0°54 : : : : 0-48 
Width of pronotum : : 0*40-0°45 : : : : 0-42 
Length of pronotum : - 0:18-0:22 : : : : 0-19 
Length of hind tibia : : I-04—1 25 : é 2 é Ie Wit 


T. rapulum is extremely difficult to separate from T. rhodesiensis in many samples 
of both imago and soldier castes, and these may be regarded as siblings, being 
partially sympatric. Most but not all imagos of T. rapulum have small diverging 
depressions on the frons, and are slightly smaller, with a slightly more inflated 
postclypeus than T. rhodesiensis ; the fontanelle is often smaller and slightly 
differently shaped (Text-figs. 308-313). Soldiers of T. vapulum have usually a 
shorter nose and a slightly different profile, but specimens occur which are virtually 
indistinguishable from some 7. rhodesiensis. Where the two species occur together 
in one locality, they are usually recognizably different, and the difficulty arises when 
attempting identification of meagre material of only one. Comparisons with T. 
dispar and T. occidentalis are made in the discussion of those species. 


Type Material. TANGANYIKA : Tanga, no date (v. Reden), syntype soldiers, type 
colony, T. vapulum (Sjéstedt), Naturhist. Riksmus., Stockholm (other syntypes in 
Mus. Berlin and A.M.N.H.). 

REPUBLIC OF SouTH AFRICA: Natal, Zululand, Mfolosi Drift, Mkosi, 1905 (J. 
Tragardh), syntype 3 imago, type colony, T. zulwensis (Holmgren), A.M.N.H. (other 
svntypes stated by Holmgren to be in Mus. Goteborg) ; Transvaal, Pretoria, 
Meukleneuk Hill, 3.x.1916 (C. Rudolph), paratype 2 imagos, type colony T. pretorien- 
sis Fuller, B.M.(N.H.). (Holotype in N.C.I., Pretoria.) 


Other Material (additional to that recorded in Sands (1957)). TANGANYIKA : 
Dar-es-salaam, no date (Regner) ; Iringa, 8.1.1935, 2 vials (H. Kirby), A.M.N.H. 

SOUTHERN RHODESIA : Salisbury, iv.1917 (R. W. Tucker) ; also 6.iv. and 3.vii. 
1949, and 5.xii.1950 (G. H. Buns) ; Guluene River, 21° 41’ S., 31° 47’ E., 13.1x. 
1962 (J. C. Felton). 

REPUBLIC OF SOUTH AFRICA: Transvaal; Letaba, 23.ix.1g1g (C. Fuller). 
Olifant River, x.1927 (H. Lang), A.M.N.H. ; Pietersburg, 8.xii.1936, Groblersdal, 


ETHIOPIAN NASUTITERMITINAE 113 


26.ix.1957, Marico, 2.ix.1961 (W. G. H. Coaton) ; Nelspruit, 15.xi.1958, Marico, 
5.-xii.1958, Sibasa, 29-30.1959, 2 vials, Waterberg, 6.1.1960 (P. C. Joubert) ; Sout- 
pansberg, 10.viii.1g60 (J. L. Sheasby) ; Lydenburg, 7.ix.1961 (L. J. de Weerdt) ; 
Warmbad, 2.x.1961 (G. F. Pretorius) ; Natal, Ubombo, i.1926 (Collins) ; also 
8.xii.1959, 2 vials (W. G. H. Coaton). 

A total of 42 nest series were examined, and all material is in the British Museum 
(Natural History) unless otherwise stated. 


Map 25. Tyvinervitermes vapulum. Vegetation types: 9, 20, and 22. 


114 W. A. SANDS 


T. vapulum shares with T. occidentalis the features of comparative rarity and 
primitive habit of feeding at least partly on woody material. It appears to have a 
restricted distribution along the east coast of Africa, where it is usually found in 
somewhat dry conditions, (Keay et al., 1959, vegetation types 20, 22), though there 
are also records from the moister coastal savannah (type 9). It has not often been 
recorded from the true Brachystegia—Julbernardia woodland (type 18), though its 
close similarity to T. rhodesiensis may have led to a few records being wrongly 
assigned to that species. T. vapulum and T. occidentalis may be relic members of 
the genus, less well adapted to grass feeding in the savannah zones than commoner 
congeners. 


Trinervitermes rhodesiensis (Sjéstedt) 
(Text-figs. 194, 195, 299-300, 314-319, 344-350 ; Map 26) 


Eutermes rhodesiensis Sjéstedt, 1911a : 186. Type locality : SOUTHERN RuopesiA, Salisbury. 

Eutermes brutus Sjéstedt, 1911b : 13. Type locality : Conco, Mukimbungu. syn. n. [ex 
syn. T. bettonianus (Sj6stedt) ; Emerson, 1928 : 492]. 

Eutermes (Trinervitermes) brutus Sjostedt ; Holmgren, 1912 : 65. 

Eutermes (Tvinervitermes) rhodesiensis Sjostedt ; Holmgren, 1912 : 65. 

Eutermes agricola Sjéstedt, 1913a : 380. Type locality : Conco, Katanga, Mufungwa. syn.n. 

Eutermes vufonasalis Sjostedt, 1913a : 383. Type locality: Conco, Katanga, Sankisia. 
syn. n. 

Eutermes (Tvinervitermes) vosent Holmgren, 1913 : 344. Type locality : Ruopesia [according 
to Holmgren], Conco, Banana [according to Sjéstedt, 1926 : 336]. syn. n. 

Eutermes (Tvinervitermes) kalahavicus Holmgren, 1913 : 346. Type locality : BECHUANALAND 
Prot., Kalahari, Kooa, Sekgoma-Khakea. syn. n. 

Trinervitermes gemellus form kalaharvicus (Holmgren) Fuller, 1922 : 110. 

Trinervitermes abassas Fuller, 1922: 114. Type locality: REPUBLIC OF SOUTH AFRICA, 
Namaqualand, Abassas on Orange River. syn. n. 

Trinervitermes thermarvum Fuller, 1922: 114. Type locality : REPUBLIC OF SouTH AFRICA, 
Transvaal, Warmbad. syn. n. 

Eutermes (Tvinervitermes) diplacodes Sjostedt, 1924: 41. Type locality : Conco, Boma, 
[syn. T. vosent (Holmgren) Emerson, 1928 : 491]. 

Eutermes (Trinervitermes) muneris SjOstedt, 1924 : 42. Type locality : Conco, Boma. [syn. 
T. voseni (Holmgren) Emerson, 1928 : 491]. 

Eutermes (Tvinervitermes) loubetsiensis SjOstedt, 1924c : 496. Type locality : REPUBLIC OF 
Conco, Loubetsi. syn. n. 

Trinervitermes vhodesiensis (SjOstedt) SjOstedt, 1926 : 346. 

Trinervitermes brutus (SjOstedt) Sjéstedt, 1926 : 331. 

Trinervitermes agricola (SjOstedt) Sjéstedt, 1926 : 329. 

Trinerviteymes vufonasalis (SjOstedt) Sjéstedt, 1926 : 334. 

Trinervitermes vosent (Holmgren) Sjéstedt, 1926 : 336. 

Trinervitermes loubetsiensis (SjOstedt) Sjostedt, 1926 : 333. 

Trinervitermes lutzi Emerson ; Sands, 1957 : 25. 


Imago. Head capsule, central part of frons and vertex brown to dark chestnut brown, 
vertex with median pale patch, yellow to brown, sometimes partly subdivided into indistinct 
streaks ; area surrounding eyes and antennal socket, in front of ocelli, and bordering posterior 
margin of postclypeus yellow to yellow-brown ; postclypeus and labrum yellow to yellow- 
brown, antennae yellow. Pronotum yellow to yellow-brown, sometimes darker at edges, meso- 
and metanota, other thoracic sclerites and legs yellow. Abdominal tergites yellow to yellow- 


ETHIOPIAN NASUTITERMITINAE 115 


brown, clouded brown around stigmata, middle tergites sometimes narrowly brown between 
stigmata ; sternites yellow, yellow-brown round stigmata. Wings translucent pale brown, 
subcosta and radius sector outlined brown at base, posterior outline of radius sector becoming 
orange distally ; membrane behind radius sector with orange streak. 

Posterior margin of head capsule behind eyes much less than semi-circular, slightly sinuate 
immediately behind eyes ; fontanelle distinct, usually rather long slender Y-shaped, with 
lateral arms often but not always dilated terminally, length often more than twice breadth at 
base of lateral arms ; eyes short oval, prominent in proportion to own diameter, medium sized 
relative to head width, W/E index 2-8—3-4 ; ocelli medium-sized, separated from eyes by one- 
seventh to two-fifths own least diameter ; postclypeus moderately inflated, width 2-2-2:5 
times length, anterior margin slightly concave, posterior margin convex, more strongly rounded 
laterally, more lightly arched in middle ; molar plate of right mandible with 7-8 ridges ; anten- 
nae 15-16 segmented, approximately half with each, basal segments variable, in 15 segmented, 
Il and V subequal, [V and VI subequal and slightly longer, III longer than these, in 16 segmented, 
II and IV subequal, III and VI subequal and slightly longer, V slightly longer still. 

Pale coloured ‘“‘ scent ’’ gland on anterior margin of fifth abdominal sternite of 2 medium- 
sized, 0-69-0-90mm. wide, 0-13—0:23mm. long ; cerci of female longer than width across base, 
conical. 

Pilosity of vertex of varying length, setae not forming a “ pelt ’’ or ‘‘ mat’’. 

Measurements (30 specimens from fo localities) in millimetres. 


Range Mean 
Head width across eyes 1:60-1:84 ; : 1:70 
Greatest diameter of eye 0°50-0°59 : : 0°54 
Ocellus : : : 0'16-0:23 X 0:2I-0-30_ .. 2 0-19 X 0°25 
Ocellus toeye  . : 0:03-0:08 é ; 0:06 
Width of pronotum . 1°43-1°73 : : 1°56 
Length of pronotum . 0:88-1:08 : : 0-98 
Length of hind tibia. I °93-2°43 : : 2°17 
Length of fore wing . 16-40-20°10 : : 18-33 


Soldiers. Major soldier: head capsule orange-yellow to ferruginous, nose ferruginous to 
chestnut brown with reddish tip. Antennae, legs, and abdominal sternites yellow, pronotum, 
meso- and metanota, and abdominal tergites orange-yellow to brown. 

Head capsule in plan view, slightly ‘‘ shouldered’’ behind antennae, nose cylindrical to 
weakly conical. In profile usually distinctly concave, but near straight, slightly humped, or 
only rising in slight curve, to behind antennal socket, then more strongly curved to highest 
point and evenly down to occiput ; nose slightly longer to distinctly shorter than rest of head 
capsule, measuring from hind margin of antennal socket, angle of nose cone o—14 degrees. 

Antennae 13-14 segmented, 7 out of 8 with 13, II and IV subequal and longer than V and VI, 
III longer up to twice II, when 14 segmented, II, III, IV and V subequal. 

Head setae consist of four at nose tip with smaller setae behind them, usually few but some- 
times more numerous. Middle abdominal tergites with scattered small setae, only longer on 
posterior margin of last tergite ; sternites with scattered short backward, and longer downward 
and forward directed setae, the latter on posterior margins and 3 or more times length of shorter. 
Minor soldier : colour as major soldier, antennae usually 12 segmented. 

Measurements (Major soldiers, 63 specimens from 17 localities, minor soldiers, 9 specimens 
from 7 localities) in millimetres. 


Major Soldiers : Range Mean 
Head length to tip of nose. 1:98—2°55 : : : : 2:28 
Head width . 3 : ‘ 1-08—1-50 E é : : 1-28 
Depth of head capsule . 2 0:78-1:14 ; : : - 0:94 
Width of pronotum : 5 0°53-0°74 ¢ g : : 0-62 
Length of pronotum ‘ : 0:20-0:32 : : : : 0:27 


Length of hind tibia : 2 I-16-1-68 A 5 : . I-41 


116 


W:. A. SANDS 


305. 


Fics. 297-307 : Tvinerviteymes, front and side views of imago head capsule. 


297, 298, T. 


vapulum ; 299, 300, T. rhodesiensis ; 301, 302, T. saudiensis ; 303, 304, T. togoensis ; 


305-307, IT. trinervius. 


ETHIOPIAN NASUTITERMITINAE 117 


Minor Soldiers : Range Mean 
Head length to tip of nose. 1°50-1-79 : 7 : ‘ 1-65 
Head width . : ; : o-61-0°78 . : : F 0:69 
Depth of head capsule . . 0*50-0°64 - - : : 0°57 
Width of pronotum 2 : 0-38-0'54 3 : - 2 0°45 
Length of pronotum : : 0:18-0:23 - : ; : 0:20 
Length of hind tibia ; ' 1:08-1:39 , 3 : 4 1°27 


Variation. In the imago, the fontanelle is sometimes partly occluded posteriorly by cuticle 
of slightly darker pigmentation than that surrounding it ; this has been observed in specimens 
from the lower Congo, and from the Transvaal. There may occasionally be small diverging 
depressions in front of the fontanelle, but these are not as common as in T. vapulum. In the 
soldier, specimens from the lower Congo tend to have slightly more conical noses than those 
from Rhodesia and S. Africa, but there is considerable overlap. 


The difficulty of separating this species from T. rapulum has been stated under 
that species. Imagos of T. rhodesiensis often have paler cuticle behind the entire 
width of the posterior margin of the postclypeus, whereas in 7. rapulum the darker 
pigmentation of the frons extends to this margin in the middle. 

The specimens attributed to T. /utzi in an earlier paper (Sands, 1957) were based 
on comparison of the soldier castes alone. These are closely similar, but in 7. 
rhodesiensis the fontanelle is smaller and the nose tip less hairy, a character not then 
recognized. The differences of the imagos of T. occidentalis and T. bettonianus have 
been discussed under those species. The large setae around the imago fontanelle 
noted in that paper are not a constant feature of the species. 


A lectotype has been selected for T. rhodesiensis and labelled accordingly : 

Lectotype soldier : SOUTHERN RHODESIA : Salisbury, 1911 (G. A. K. Marshall), 
and paralectotype from type colony, T. rhodesiensis (Sjéstedt), B.M.(N.H.) (other 
paralectotypes in A.M.N.H., and Naturhist. Riksmus., Stockholm). 


Type Material. Conco : Mukimbungu, 1904 (K. E. Laman), syntype soldiers, 
type colony, T. brutus (Sjéstedt), A.M.N.H. (other syntypes in Naturhist. Riksmus., 
Stockholm) ; Katanga, Mufungwa, 12.xii.1g11 (J. Bequaert), syntype soldiers, type 
colony, 7. agricola (Sjostedt), A.M.N.H. and Naturhist. Riksmus., Stockholm ; 
Sankisia, 16.vii.tg11 (J. Bequaert), syntype soldiers, type colony, 7. rufonasalis 
(Sjéstedt), A.M.N.H. (other syntypes in Naturhist. Riksmus., Stockholm) ; Boma, 
vili.1920, and Moanda, 28.viii.1g20 (H. Schouteden), syntype soldiers, T. diplacodes 
(Sjéstedt), A.M.N.H. (other syntypes in Naturhist. Riksmus., Stockholm) ; Boma 
4.xi.1913 (Styckzynski), syntype soldiers, type colony, T. muneris (Sjdstedt) 
A.M.N.H., and Naturhist. Riksmus., Stockholm. 

REPUBLIC OF CONGO : Loubetsi, approx. 3° 40’ S., 12° 15’ E., no date (A. Nillson), 
syntype soldiers, type colony, T. loubetsiensis Sjéstedt, A.M.N.H. (other syntypes in 
Naturhist. Riksmus., Stockholm). 

BECHUANALAND : Kooa, Sekgoma Khakea, no date (A. Schulze), syntype, T. 
kalaharicus Holmgren, A.M.N.H. 

REPUBLIC OF SouTH AFRICA : Namaqualand, Abassas on Orange River, iii.1921 
(J. H. C. Kraphal), paratype 3 imagos, T. abassas Fuller, B.M.(N.H.) (Holotype 
and other paratypes in N.C.I., Pretoria) ; Transvaal, Warmbad, xii.1g16 (C. 


118 W. A. SANDS 


Rudolph), paratype 2 imagos, T. thermarum Fuller, B.M.(N.H.) (Holotype and 
other paratypes in N.C.I., Pretoria.) 


Other Material. REPUBLIC oF Conco: 13km. W. of Brazzaville, 8.vi.1948, 2 
vials (A. E. Emerson), A.M.N.H. 


Conco: Luluabourg, no date (P. Callewaert) ; Mukimbungu, 1904 (K. E. 
Laman) ; Boma, 4.xi.1913 (Styckzynskt) ; Thysville, i.vi.tg15 (H. Lang, J. P. 
Chapin) ; also 3.vi.IgI5, 2 vials, and Matadi, 18.vi.1g15 (J. Bequaert) ; Boma, 
10-I2.vili.1g20, 4 vials, and Moanda, viii.1g20 (H. Schouteden) ; Gihinga, 14.viii. 
1941 (H. Bredo) ; Sona Mpangu, 10-12.iv.1948, 7 vials, and 18km. S. of Leopold- 
ville, 9.vi.1948 (A. E. Emerson), all in A.M.N.H. 


ANGOLA: St. Antonio, vili.1g15 (H. Lang, J. P. Chapin), A.M.N.H.; Porto 
Alexandre, 1954 (J. Balfour-Browne). 


NORTHERN RHODESIA: Lake Mweru, Masenka Flats, 8.vili.1g42 (H. Bredo), 
A.M.N.H. ; Abercorn, iii.1947, and xi.—xii.1948, 6 vials (P. E. Glover). 


SOUTHERN RHODESIA : Salisbury, no date (W. Barbrook) ; ditto, 1g11 (G. A. K. 
Marshall) ; Worldsview, 1920 (C. W. Mally) ; Concession, 20.v.1927 (R. Jack). 


BECHUANALAND : Topsi, 11.11.1921 (C. E. Godman) ; Tsukutsa Pan, 3.vili.1g24 
(C. W. Mally) ; Tsotsoroga Pan, viii.t930, and Makarikari Pan, 5.viii.1930 (H. 
Lang), A.M.N.H. ; Gantsena Pan, 1955, and 60m. S. of Ghanzi, vili.1955 (R. Story). 


REPUBLIC OF SOUTH AFRICA : Cape Province ; Taungs, 3.x.1956, Barkly West, 
4.x.1956 (W. G. H. Coaton). Transvaal ; Potgeitersrust, 17.1x.1918, and Letaba, 
20.1x.1918 (C. Fuller) ; Waterberg, 25.ix.1957, and Soutpansburg, I1.x.1960 
(W. G. H. Coaton) ; Soutpansburg, 1.1.1960, Waterberg, 6.i.1960, and Marico, 
g.i.1960, 2 vials (P. C. Joubert) ; Rustenburg, 4.x.1961 (G. F’. Pretorius). 


A total of 64 nest series were examined and all material is in the British Museum 
(Natural History) unless otherwise stated. 


The distribution range of 7. rhodesiensis overlaps to a limited extent with its 
sibling congener, T. rapulum, in Southern Rhodesia and the Transvaal. T. rhodestensts 
covers the Brachystegia—Julbernardia savannah woodland and the other savannah 
gradations to semi-desert, from the southern border of the Congo forest southwards 
to the beginnings of the Karroo, Macchia, and temperate grasslands of the Cape. 
The East-West barrier separating the species from the Zambesi northwards appears 
to approximate to the Great Rift Valley. A few of the most South-Western records 
are from areas mapped by Keay e¢ al., (1959) as desert, but these are near the Orange 
River and it is presumed that the conditions are locally subdesert, and that full 
desert would constitute an ecological barrier. The distribution of T. rhodesiensis to 
the south of the Congo Rain forest approximates in climatic and vegetation range 
to that of T. oeconomus (Trag.) to the north. 


ETHIOPIAN NASUTITERMITINAE 119 


Map 26. Trinervitermes rhodesiensis. Vegetation types: 8, 16, 18, 19, 20, 25, and 31. 


120 W. A. SANDS 


Trinervitermes saudiensis sp. n. 
(Text-figs. 301, 302, 320, 321 ; Map 3.) 

Imago. Colouration uncertain due to fading in old dried specimens, but thought to be similar 
to T. vapulum. 

Posterior margin of head capsule behind eyes less than semi-circular, slightly sinuate immed- 
iately behind eyes ; fontanelle distinct, Y-shaped, lateral arms long, reaching almost half-way 
to ocelli, slender but slightly dilated at point where termination occurs in other species ; frons 
with two diverging depressions in front of fontanelle, or these united ; eyes very short oval, 
moderately prominent in proportion to own diameter, small relative to head width, W/E index 
3°5-3°9 ; ocelli medium sized, separated from eyes by more than half to more than three- 
quarters own least diameter ; postclypeus moderately to strongly inflated, width 2-1-2-4 
times length, anterior margin slightly concave, posterior margin convex, rounded, slightly arched 
in middle ; molar plate of right mandible with 7-8 ridges ; antennae 15 segmented, II, IV, and 
V subequal, III and VI subequal and slightly longer than these. 

Pale coloured “‘ scent’ gland on anterior margin of fifth 2 abdominal sternite small ; cerci 
of 2 as long as width across base, slightly mamilliform conical. 

Pilosity of vertex of varying length, not forming a “ pelt’ or “‘ mat’. 

Measurements (10 specimens from 1 locality) in millimetres. 


Range Mean 
Head width across eyes 161-176 - : 1-66 
Greatest diameter of eye 0°43-0°48 : : 0-46 
Ocellus F ‘ : 0-16-0-18 X 0:20-0:25. ‘ 0-17 X 0:22 
Ocellus toeye  . : 0-I10-0'13 5 s o-ll 
Width of pronotum . 1-38-1-61 : : 1-48 
Length of pronotum . 0°84—0°94 : : 0:90 
Length of hind tibia. 2°04—2°24 : : 2°17 
Length of fore wing . 15°50-16:30 : : 0573 


Soldiers. Unknown. 

T. saudiensis is separable from T. dispar and T. rapulum by the proportionately 
smaller eyes, with the ocelli more distant from them, and the shorter 2 cerci. The 
same characters apart from the cerci distinguish it from T. togoensis and T. occident- 
alis. The wide spreading lateral arms of the fontanelle are unlike any other 
Trinervitermes except some T. trinervoides. 

T. saudiensis is described here because it is so distinct from the other species of the 
Ethiopian Zoogeographical Region and because the existence of a further Tvinervi- 
termes species in the Arabian Peninsula is an important link in the distribution of 
the genus in Africa and India. I consider it most improbable that this species is 
the imago of T. avabiae. The various primitive features such as the wide branched 
fontanelle, small eyes widely separated from ocelli, the long postclypeus, slightly 
mamilliform cerci and small ventral scent gland, are shared with several other species. 
but none shows such a combination of features common to the rest, suggesting that 
this species may be close to the ancestral form of the genus. There is a possibility 
that this is the at present unknown imago caste of Mycterotermes meringocephalus, 
which might be expected to show a similar combination of features. 

Two vials of this species were studied. 


Holotype g, SAUDI ARABIA: Jidda, 24.xi.1926 (H. St. J. B. Philby), B.M.(N.H.). 
Allotype 9, same data as holotype, B.M.(N.H.). 
Paratypes J, same data as above and also 3.ii.1934 (G. L. Bates), B.M.(N.H.). 


ETHIOPIAN NASUTITERMITINAE 121 


S22. 333. 336. 


Fics. 308-337: Imago fontanelle. 308-310, 3, 311-313, 2, Tvinervitermes vapulum ; 
314-316, 3, 317-319, 9, T. rhodesiensis ; 320, go, 321, 9, T. saudiensis ; 322, 323, 4, 


324, 325, 2, T. togoensis ; 326-328, g, 329-331, 9, T. trinervoides ; 332, 333, d, 334, 
335, 2, I. trinervius ; 336, 3, 337, 2, Baucaliotermes hainesi. 


122 Wi -FAs SANDS 


Trinervitermes togoensis (Sj6stedt) 


(Text-figs. 303, 304, 322-325, 351-358 ; Map 27) 
Eutermes togoensis Sjéstedt, 1899 : 37. Type locality : Guana, Trans-Volta Togoland, Ho. 
Eutermes tervicola Tragardh, 1904 : 31. Type locality : SupAN, Kaka. syn. n. 
Termes (Eutermes) togoensis (Sj6stedt) Desneux, 1904a : 45. 
Termes (Euteyvmes) tervestvis Desneux, 1904a : 45 [mew name for E. tervicola Tragardh, presum- 
ably (not stated) on grounds of homonymy with Termes (Termes) terricola Sjostedt, 1902]. 
Eutermes togoensis SjOstedt ; Sjostedt, 1904 : 27 and 87. 
Eutermes tervicola Tragardh ; Sjostedt, 1904 : 39 and 96. 
Eutermes (Trvinervitermes) togoensis SjOstedt ; Holmgren, 1912 : 62. 
Eutermes (Trinervitermes) terricola Tragardh ; Holmgren, 1912 : 64. 
Eutermes suspensus Silvestri, 1914 : 41. Type locality : Guinea, Kakoulima. syn. n. 
Trinevvitermes terricola (Tragardh) Sjdéstedt, 1926 : 326. 
Trinervitermes suspensus (Silvestri) Sjéstedt, 1926 : 329. 
Trinervitermes togoensis (SjOstedt) Sjdstedt, 1926 : 331. 
Trinervitermes nigeriensis SjOstedt, 1926 : 333. Type locality : Mati, Belia. syn. n. 


Imago. Head capsule, central part of frons and vertex brown to dark sepia brown ; vertex 
with two or three diverging paler streaks, middle one often indistinct or absent ; postclypeus, 
labrum, area between eye and mandible base, and tapering streak above each eye orange-yellow 
to brown, pale area in front of eye sometimes extending in front of ocelli and bordering posterior 
margin of postclypeus ; antennae orange-yellow to brown. Pronotum orange-yellow to 
yellow-brown, sometimes shaded brown at anterior corners, meso- and metanota yellow to 
orange-yellow ; other thoracic sclerites and legs yellow. Abdominal tergites brown to dark 
chestnut brown, darker round stigmata and paler on anterior corners, sternites pale yellow, 
sometimes brown round stigmata. Wings translucent pale brown, subcosta and radius sector 
yellow, partly outlined brown at base, membrane behind radius sector with yellow streak ; 
rest of venation narrowly brown or outlined brown. 

Posterior margin of head capsule much less than semi-circular behind eyes, slightly sinuate 
immediately behind eyes ; fontanelle small, paler than head, short Y- or inverted A-shaped, 
sometimes with small depression in front or behind, or partly occluded posteriorly by darker 
cuticle, lateral arms often dilated terminally ; eyes oval, weakly to moderately prominent in 
proportion to own diameter, medium sized relative to head width, W/E index 2-5-3-2 ; ocelli 
medium sized oval, touching eyes, to separated from them by one-third own least diameter ; 
postclypeus moderately inflated, width 2:3-2-6 times length, anterior margin somewhat concave, 
posterior margin convex, rounded, sometimes slightly indistinct in middle ; molar plate of 
right mandible with 6-8 ridges ; antennae 15-16 segmented, rarely 17, one in five with 15 but 
usually with basal segments partly subdivided, in 16 segmented form II, III, V, and VI are 
subequal and longer than IV. 

Pale coloured “‘scent’’ gland on anterior margin of fifth abdominal sternite of 2 small, 
0°35-0-64mm. wide, o-13mm. long ; cerci of 9 slightly shorter to slightly longer than width 
across base, conical. 

Pilosity on vertex of varying length, setae not forming a “ pelt’ or “ mat”. 

Measurements (36 specimens from 12 localities) in millimetres. 


Range Mean 
Head width across eyes I°43-1°76 5 : 1°56 
Greatest diameter of eye 0:50-0-63 : : 0°55 
Ocellus : : ; 0:15-0:22 X 0:20-0:28 . : 0-19 X 0:24 
Ocellus toeye  . : 0:00-0:05 ; : 0:03 
Width of pronotum . I*30-1°59 : : 1°43 
Length of pronotum . 0-83-1-05 : ; O-91 
Length of hind tibia. 1°95-2°43 ° ; 2°20 


Length of fore wing. 14°*20—21:00 : é 17-14 


ETHIOPIAN NASUTITERMITINAE 123 


\/¢ 


338. 


340. 
341. 


350 


358. 


3oi7. 


Fics. 338-358 : Tvinervitermes, soldier head capsule, side and plan views, and nose tip. 
338-343, T. vapulum ; 344-350, T. rhodesiensis ; 351-358, T. togoensis. 


124 W. A. SANDS 


Soldiers. Major soldier : head capsule orange to chestnut brown, nose ferruginous to pitch 
black with red tip. Antennae and pronotum orange-yellow, meso- and metanota and legs 
yellow. Abdominal tergites yellow to brown, sternites yellow. 

Head capsule in plan view short oval, circular, to slightly broader than long, nose near 
cylindrical to moderately conical. In profile near straight to distinctly concave with head 
rising from above or behind antennal socket in even curve to highest point and down to occiput ; 
nose much shorter than rest of head capsule, measuring from hind margin of antennal socket, 
up to equal in length, angle of nose cone o-16 degrees. Antennae 13-14 segmented, 9 out of 
10 with 13. IV and V subequal, longer than II and shorter than III. 

Head setae confined to distinctly to sparsely hairy nose tip. Abdominal tergites with 
scattered minute setae, slightly longer on posterior three. Sternites with short backward 
directed setae and longer downward or forward directed setae on posterior margins, arranged 
approximately in 6 longitudinal rows, 3 or more times length of shorter setae. 

Minor soldier : colour as the darker major soldiers, antennae 12-13 segmented, about equally 
divided between the two. 

Measurements (Major soldiers, 70 specimens from 18 localities, minor soldiers, 13 specimens 
from 13 localities) in millimetres. 


Major Soldiers : Range Mean 
Head length to tip of nose. I-70-2°43 5 ; * ; 2-01 
Head width . : : : 0°95-1°58 ‘ : : ; I°L7 
Depth of head capsule . : 0:68-1:07 : : ; : 0-84 
Width of pronotum : ‘ 0-48-0°77 : : ‘ : 0°55 
Length of pronotum 3 : 0:20-0:36 : : : : 0:24 
Length of hind tibia : ; 1-23-I°71 : é : : 1-39 

Minor Soldiers : Range Mean 
Head length to tip of nose. I°35-1°63 ; : ‘ : 1°46 
Head width . : : : 0°53-0°68 : : : : 0°59 
Depth of head capsule . : 0°43-0°53 : 3 : ‘ 0°47 
Width of pronotum : : 0:38-0°45 : : : : 0°39 
Length of pronotum : : 0:18-0:20 ; ; : 5 0-18 
Length of hind tibia : : I-03-1°28 : : : : 2a 


Variation. In the imago, there are sometimes small diverging depressions or a single median 
depression on the frons, but these are usually absent or indistinct. The soldier caste is especially 
variable in nose length, with a tendency for specimens from the western end of the range to be 
generally longer than those from the east. Particularly noticeable in this species is the tendency 
for specimens from higher altitudes to be larger than those from low. 

The imago of T. togoensis is not likely to be confused with other species except in 
a very small number of cases where the antennae are 15 segmented without appre- 
ciable subdivision of III, when it resembles T. occidentalis, and T. dispar. T. 
occidentalis has rather more prominent eyes, usually longer 9 cerci, and a differently 
shaped fontanelle ; 7. dispar has smaller eyes, lacks diverging pale streaks on the 
vertex, and usually has a shorter hind tibia and longer 9 cerci. 

In the soldier caste, some specimens of T. dispar, T. oeconomus and T. geminatus 
are almost indistinguishable from 7. togoensis, but the nose tip is usually less hairy 
in T. dispar, the middle abdominal tergites have fewer minute setae in T. oeconomus, 
and the antennae of T°. geminatus are always 14 segmented. 


Specimens have been selected and labelled as lectotypes of T. togoensis and T. 
nigeriensis by Dr. A. E. Emerson, but the designations have not been published 
hitherto ; that of the senior synonym Is given separately : 


ETHIOPIAN NASUTITERMITINAE 125 


Lectotype : GHANA: Trans-Volta Togoland, Ho, 1892 (Rossman), lectotype 2 
imago, T. togoensis (Sjéstedt), Mus. Hamburg, and paralectotype 2 imago, Natur- 
hist. Riksmus., Stockholm. 

Type Material. SupAN: Kaka, 1go1 (J. Tvdgardh), syntype soldier, type colony, 
T. terricola (Tragardh), A.M.N.H. (other syntypes in Naturhist. Riksmus., Stock- 
holm). 

GuINEA : Kakoulima, 9° 40’ N., 13° 25’ W., 1912-13 (F. Silvestri), holotype ° 
imago and paratype soldiers, 7. suspensus (Silvestri), Silvestri Coll., Lab. Zool., Sc. 
Agric., Portici. 

MALI: Belia, 15° 26’ N., 1° 45’ W., no date (R. Chudeau), lectotype 2 imago, 
T. mgeriensis (Sjéstedt), Mus. Paris. (Paralectotypes in Mus. Paris and Naturhist. 
Riksmus., Stockholm). 


Other Material. SIERRA LEONE: Cape S. Leone, 1858 (J. Foxcroft) ; Samaia, 
24.v.1948 (f°. A. Squires) ; Freetown, Brookfields, 27.v.1956 (P. F. Prevett) ; and 
9.1.1958 (W. Wilkinson). 

GHANA: 45m. from Accra on Ada Road, 16.11.1955 (W. Belfield) ; 114m. from 
Accra on Kumasi Road, 18.11.1959, 33m. N. of Ejura on Tamale Road, 24.11.1959, 
tom. N. of Yeji on Tamale Road, 25.11.1959, 16m. from Daboya on Tamale Road, 
28.11.1959, 28m. from Tamale on Bolgatanga Road, 1.i11.1959, 55m. from Tamale on 
Larabanga Road, 3.iii.1959, 4 and 15m. from Gambaga on Wale Wale Road, 5.111. 
1959, and 7.i1i1.1959, 4 vials, 25m. S.E. of Gambaga on Yendi Road, and 17m. N.E. 
of Gambaga on Bawku Road, 6.111.1959, 20m. from Bolgatanga on Bawku Road, 
8.i11.1959, 12m. N. of Navrongo, 14.iii.1959, 11m. from Tumu on Navrongo Road, 
15.lii.1959, 6 and 4om. from Tumu on Lawra Road, 16.11.1959, 2 vials, Lawra, and 
om. N. of Lawra on Nandom Road, 18.111.1959, 3 vials, 6m. N. of Wa on Lawra Road, 
19.i11.1959, 52m. S. of Wa on Bole Road, 20.iii.1959, 22m. from Larabanga on Tamale 
Road, 24.i11.1959, 41m. S. of Tamale on Kumasi Road, 27.ii1.1959, 25m. N. of Wenchi 
on Bamboi Road, 30.1ii.1959 (W. A. Sands). 

Upper Volta: 15m. N. of Bawku (Ghana), 11.11.1959 (W. A. Sands). 

NIGERIA : Western Region ; Lagos, Lighthouse beach, 9.iv.1957 (W. Wilkinson). 
Eastern Region ; 14m. from Enugu on Abakaliki Road, 1.11.1957, Abakaliki, and 
4om. from Abakaliki on Ikom Road, 27.v.1957 (W. Wilkinson) ; 12m. from Enugu 
on Onitsha Road, 2.iii.1958, 2 vials, and 20m. from Enugu on Oturkpo Road, 
Nsukka Hills, 5.iii.1958 (W. A.Sands). Northern Region ; Sokoto, ii.1955 (W. V. 
Harris) ; Kano, xi.1957 (R. A. Blease) ; 25m. from Katsina on Daura Road, 
1g.x1.1956, 28 and 42m. from Sokoto on Argungu Road, 23.xi.1956, 2 vials, 11-18m. 
from Kaduna on Zungeru Road, 8.xii.1956 and 28.ii.1957, 4 vials, 70m. from Kaduna 
on Zungeru Road, 18.xii.1956, 2 vials, 26m. from Minna on Zungeru Road, 20.xii. 
1956, 16m. S. of Zungeru on Bida Road, and Mokwa, 21.x11.1956, 4m. N. of Bida, 
23.X11.1956, 26m. from Abuja on Bida Road, 24.xii.1956, Abuja, 31.xii.1956, 22m. 
from Kaduna on Kachia Road, 30.1.1957, 2 vials, 4-16m. from Jos on Bauchi and 
Bukuru Roads, 6-11.11.1957, 5 vials, 4om. from Jos on Bauchi Road, 11.11.1957, 
Bauchi, and 25m. from Bauchi on Jos Road, 12—13.11.1957, 18-24m. from Jos on 
Pankshin Road, 14.ii.—3.1v.1957, 9 vials, 56m. from Kaduna on Jos Road, 15.11.1957, 


126 W.A. SANDS 


25-32m. from Jos on Kaduna Road, 5.iv.1957, 2 vials, 39m. from Bauchi on Gombe 
Road, 7.v.1957, Yola, 13.v.1957, 54m. from Yola on Jalingo Road, 16.v.1957, 7, 45, 
and 53m. S.W. of Jalingo on Takum Road, 17.v.1957, 3 vials, 125m. and 114m. 
S.W. of Jalingo on Takum Road via Beli, 18.v.1957, 3 vials, 55m. from Damaturu 
on Potiskum Road, 5.vi.1957, 4m. S. of Ilorin on Oyo Road, 4.xii.1957, 5m. S. of 
Kunya on Kano Road, 4.x.1957, 24m. from Gboko on Makurdi Road, 25.11.1958, 


\ 
a 


Map 27. Tyvinervitermes togoensis. Vegetation types: 8, 16, 17, 20, and 25. 


saa 


ETHIOPIAN NASUTITERMITINAE 127 


4m. from Gboko on Oturkpo Road, 26.11.1958, 30m. from Lokoja on Okene Road, 
8.iii.1958, Samaru, near Zaria, vi—vii.1959, 8 vials (W. A. Sands). 

SuDAN : Darfur Province, Kulme, 12° 30’ N., 23° 27’ E., 6 and 26.vii.1g21, 2 
vials (H. Lynes). 

Ucanpa : Karamoja Distr., Toror Hills, near Kotido, 8.x.1952 (W. A. Sands). 

A total of 108 nest series were examined, and all material is in the British Museum 
(Natural History) unless otherwise stated. 


T. togoensis is another of the species which extend from the extreme West Coast 
of Africa across the “ Guinean ”’ and “‘ Sudan ”’ vegetation zones (Keay et al., 1959 
types 8, 16, 17, and 20) to the Sudan and Uganda. 

Its biology and feeding habits have been described in detail in other papers (Sands, 
1961, 1961a). It seldom builds a mound structure of its own, being commonly 
found in the mounds of other Tvinervitermes species, or those of Cubitermes, Thoraco- 
termes (record of Silvestri, 1914), Macrotermes, Pseudacanthotermes, and Odonto- 
termes. In this respect it appears to occupy a closely similar ecological niche to that 
of T. dispar in the East and Central Africa fauna. 


Trinervitermes trinervius (Rambur) 
(Text-figs. 305-307, 332-335, 371-375, 383 ; Map 28) 


Termes trinervius Rambur, 1842 : 308. Type locality : SENEGAL. 

Termes (Eutermes) trinervius Rambur ; Hagen, 1858 : 216. 

Eutermes trinervius (Rambur) Sjéstedt, 1900 : 108. 

Eutermes holmgreni von Rosen, 1912: 224. Type locality: LrBeria. f[syn. T. trinervius 
(Rambur) Sjéstedt, 1926 : 340}. 

Eutermes (Trinervitermes) trinervius (Rambur) ; Holmgren, 1912 : 64. 

Eutermes (Trinervitermes) holmgreni von Rosen ; Holmgren, 1912 : 65. 

Eutermes posselensis Sj6stedt, 1914b : 94. Type locality : Cuan, Fort Possel. syn. n. 

Termes (Eutermes) trinervius Rambur ; Desneux, 1915 : 9. 

Trinervitermes trinervius (Rambur) Fuller, 1922 : 107. 

Trinervitermes posselensis (SjOstedt) Sjéstedt, 1926 : 348. 

Trinervitermes carbonarius SjOstedt, 1926b : 158. Type locality : Conco, Haut Uélé, Mauda. 
syn. n. 

Trinervitermes carbonarius Sjostedt ; Sands, 1957 : 20. 


Imago. Head capsule, frons and vertex dark chestnut to very dark sepia brown, area 
surrounding eye, extending to point behind and to mandible base in front paler, yellow-brown 
to brown ; vertex with three diverging paler streaks, and area in front of ocelli brown ; labrum 
yellow, postclypeus yellow-brown, clouded with brown ; antennae yellow-brown. Pronotum 
orange yellow to brown, meso- and metanota and other thoracic sclerites yellow, legs orange- 
yellow, slightly darker at base of tibia. Abdominal tergites dark sepia brown with yellow-brown 
margin of varying width, paler at anterior corners and sometimes in middle ; sternites yellow 
to orange-yellow, brown around stigmata. Wings translucent pale brown, subcosta and radius 
sector orange-yellow, outlined brown at base, posterior outline of radius sector continuing to 
near wing tip ; membrane behind radius sector with indistinct yellow-brown streak ; rest of 
venation narrowly brown or outlined brown. 


128 W.A. SANDS 


Posterior margin of head capsule much less than semi-circular behind eyes, slightly sinuate 
immediately behind eyes ; fontanelle pale, sharply contrasting with surrounding cuticle, 
Y-shaped, to short triangular owing to occlusion of posterior part by pigmented cuticle, lateral 
arms short, sometimes dilated terminally ; eyes oval, moderately prominent in proportion to 
own diameter, medium sized to large relative to head width, W/E index 2-4-2-9, ocelli medium 
sized to large, oval, separated from eyes by one-fourteenth to three-tenths own least diameter ; 
postclypeus moderately inflated, width 2-3—2-4 times length, anterior margin near straight to 
slightly concave, posterior margin convex, rounded, sometimes slightly indistinct or with slight 
bulge in middle ; molar plate of right mandible with 7-8 ridges ; antennae 17 segmented, 
II and III subequal, longer than VI, IV and V subequal and shorter than IV. 

Pale coloured “scent ’’ gland on anterior margin of fifth abdominal sternite of 2 medium- 
sized, 0:70-I-o2mm. wide, 0-20-0-:30mm. long ; cerci of 2 equal to or longer than width across 
base, conical with somewhat irregular outline. 

Pilosity on vertex consists of short curved setae of even length forming a “ pelt ’’ or “‘ mat’, 
and scattered longer setae among them. (Text-fig. 383). 

Measurements (22 specimens from 7 localities) in millimetres. 


Range Mean 
Head width across eyes 1 -78-2°17 : : 1°85 
Greatest diameter of eye 0:65-0°85 - , 0°74 
Ocellus ‘ : : 0:20-0:29 X 0:26-0°35 . ; 0:24 X 0:30 
Ocellus toeye  . : 0:02-0:06 : : 0:04 
Width of pronotum I -63—2-07 : 5 1-86 
Length of pronotum . 1-03-I°15 : : Ere 
Length of hind tibia . 2°33-2:°86 ; ‘ 2°49 
Length of fore wing . 16-40-21 -50 3 3 19:96 


Soldiers. Major soldier : head capsule, chestnut brown to nearly pitch black, nose dark 
chestnut brown to pitch black with reddish tip ; antennae yellow-brown. Pronotum, anterior 
lobe yellow-brown, posterior lobe, meso- and metanota, legs and abdominal sternites yellow ; 
abdominal tergites yellow to brown. 

Head capsule in plan view short oval to slightly wider than length, nose near cylindrical to 
weakly conical. In profile distinctly concave, with back of head rising from above or slightly 
behind antennal socket, and evenly rounded ; nose slightly shorter to distinctly longer than 
rest of head capsule, measuring from antennal socket, angle of nose cone 5—10 degrees ; antennae 
13-14 segmented, 6 out of 7 with 14, II, IV, V, and VI subequal and slightly shorter than III, in 
13 segmented forms III longer and usually partly subdivided. Head setae confined to 4 at nose 
tip and very few small pale setae behind these ; dorsal surface of head capsule without 
prominent setae, sometimes minute traces of bilaterally placed pair on vertex present. 

Middle abdominal tergites with scattered minute setae, visible in profile, longer on posterior 
two tergites ; sternites with short backward and longer downward and forward directed setae, 
the latter on posterior margins and 2 or more times length of shorter. 

Minor soldier : colour as major soldier, antennae 12-14 segmented, proportions of basal 
segments variable. 

Measurements (Major soldiers, 64 specimens from 1g localities, minor soldiers, 6 specimens 
from 4 localities) in millimetres. 


Major Soldiers : Range Mean 
Head length to tip of nose. 2°I1I-2-°70 : : A ‘ 2°47 
Head width . : 3 : 1:18-1:73 : : : c 1-48 
Depth of head capsule . ‘ 0:86-1:20 : : s , 0-88 
Width of pronotum : 6 0+53-0°74 . : . ; 0-68 
Length of pronotum : F 0:28-0:38 ‘ : : ‘ 0-32 


Length of hind tibia 3 : I -48-2-o1 : ; ‘ 3 1-78 


ETHIOPIAN NASUTITERMITINAE 129 


Minor Soldiers : Range Mean 
Head length to tip of nose. 1-73-2:04 : ; : : I-go 
Head width . , : : 0:78-0:94 ; 2 i ; 0-86 
Depth of head capsule . F 0-58-0-72 ; : : ; 0-65 
Width of head capsule . : 0°43-0°55 : - : * 0-50 
Length of pronotum 3 ; 0-20-0-29 : : ; : 0°24 
Length of hind tibia é : 1-39-1-68 1°54 


Variation. In the imago, there are sometimes small div exeing depressions on the frons. The 
large variation in eye size results in specimens of very different general appearance, and there is 
some indication of a cline in this character from moister to drier areas, the largest eyes occurring 
in the moister southern vegetation types. 

The “‘ major’ and “‘ minor ”’ soldier castes are not always distinctly separable. It appears 
to be characteristic of the largest colonies that a complete gradation of intermediate forms 
develops. 


The imago of 7. tvinervius closely resembles that of 7. geminatus, but is usually 
darker in colour, with larger eyes, a slightly more inflated postclypeus and longer 
9 cerci ; there is however an overlap in all these characters, and since the two 
species are sympatric, they constitute a further example of near-siblings among 
termites. The soldiers are usually distinguishable by the more hairy nose tip and 
darker coloured abdominal tergites of T. geminatus. T. oeconomus is also closely 
similar in the imago, but is distinguished by the vertex pilosity. 

Among allopatric species, only the darker soldiers of 7. gratvosus closely resemble 
T. trinervius, and are in some cases virtually indistinguishable, though usually 
much shorter-nosed. 

The close similarity of T. tvinervius to T. geminatus and T. oeconomus has in the 
past led to confusion which has arisen mainly because the unique holotype ¢ imago 
lacks the head. Rambur (1842) omitted mentioning the head or its absence, while 
including it in descriptions of other species ; it can thus be presumed that the 
holotype was headless when he described it. Hagen (1858) noted the deficiency. 

Walker (1853) described as T. trinervius a specimen of 7. oeconomus labelled 
“ Tripoli’, but this must have come from tropical Africa, probably the Sudan. 
The reference of Sjéstedt (1900) is included in the synonymy because of the change 
of genus involved ; the specimens described were probably again T. oeconomus, 
though some from South Africa were included. The specimens described by Silvestri 
have not been examined, but would appear to have been 7. geminatus ; some of 
the material collected by Silvestri in Senegal was however T. trinervius. 

I have found no reliable specific characters in the pronotum shape, colour, or 
pilosity, nor are the outlines of meso- and metanota of any value in Tyvinervitermes. 
The legs are not distinctive in closely related species, and the abdominal characters 
occur in the 9. I therefore think it unlikely that the use of Rambur’s name will 
ever be finally settled by examination of the holotype. The general appearance, 
the colouration, and the shape of the § cerci of this specimen, though not conclusive, 
appear to me to be most consistent with the species hitherto known as T. posselensis 
of which T. carbonarius is a junior synonym. In spite of usage by some authors 
subsequent to Rambur, I consider it inadvisable to use T. tvinervius as senior 
synonym to 7. oeconomus, since that species is well known and established under its 
present name, with adequate type material and several junior synonyms. 


130 WAS SANDS 


Type Material. SENEGAL: no detailed locality, no date, holotype, unique 
headless f imago, T. tvinervius (Rambur), Selys Longchamps coll., Inst. R. Sci. Nat., 
Brussels. 

CENTRAL AFRICAN REPUBLIC: Fort Possel, xi.1g10 (H. Schubotz), syntype 
soldiers, type colony, T. posselensis (Sjéstedt), A.M.N.H. (other syntypes in Mus. 
Hamburg and Naturhist. Riksmus., Stockholm). 

Conco: “ Haut Uélé’”’, Mauda, 27.11.1925 (H. Schouteden). [See discussion on 
Type locality of T. maudanicus Sjostedt.]| Syntype soldiers, type colony, 7. 
carbonarius Sjéstedt, Naturhist. Riksmus., Stockholm (other syntypes in Mus. 
Tervuren). 


Other Material. SENEGAL: Dakar, 2.vili.1g12 (F. Silvestri), A.M.N.H. 

GUINEA : Conakry, 8.viii.1gi2 (F. Silvestri), AMM.N.H. ; Mt. Nimba, 1957, 4 
vials (M. Lamotte), Inst. Franc. d’Afr. Noire, Dakar. 

SIERRA LEONE : Cape S. Leone, 1858 (J. Foxcroft). 

LIBERIA : 1908 (Scherer), A.M.N.H. [? paratype E. holmgreni von Rosen]. 

Ivory Coast : Dabou, 50km. W. of Abidjan, 1962-1964, 12 vials (P. Bodot). 

GHANA : 36m. from Accra on Ada Road, 18.ii.1955, Nungua, 16.11.1955, and 
I4—16.vi.1956, 5 vials (W. Belfield) ; Accra and 113m. from Accra on Kumasi Road, 
14-16.vi.1959, 60m. N. of Ejura on Tamale Road, 24.11.1959, 16m. from Daboya on 
Tamale Road, 28.11.1959, 55m. from Tamale on Larabanga Road, 3.iii.1959, Volta 
Bridge on Bolgatanga—Bawku Road, 8.111.1959, 12m. N. of Navrongo, 14.ili.1959, 
11m. from Tumu on Navrongo Road, 15.11.1959, 6m. from Tumu on Lawra Road, 
16.11.1959, 9m. N. of Lawra on Nandom Road, 18.ili.1959, 52m. from Wa on Bole 
Road, 20.11.1959, 22m. from Larabanga on Tamale Road, 24.11.1959, 16m. from 
Nkoranza on Kintampo Road, 28.1ii1.1959 (W. A. Sands). 

NIGERIA : Western Region ; Lagos, Ebute Metta, 7.v.1936 (T. Scott). Northern 
Region ; 13m. S. of Makurdi, 1954, 18m. from Kaduna on Zaria Road, 1954, 60m. 
from Kaduna on Jos Road, 1954, and 7m. S. of Zaria, 1954 (J. B. MacNulty) ; 17m. 
S. of Zungeru on Bida Road, 21.xii.1956, 40m. from Abuja on Bida Road, 24.xii. 
1956, 20m. from Abuja on Keffi Road, Idu, 28.xii.1956, 18-22m. from Kaduna on 
Kachia Road, 30.1.1957, 4 vials, Kudaru, on Zaria—Jos Road, 5.i11.1957, 4 vials, 5m. 
from Jos on Bukuru Road, and 6m. from Jos on Bauchi Road, 6.11.1957, 42m. from 
Jos on Wamba Road, 8.11.1957, 18m. from Jos on Pankshin Road, 14.11.1957, 50 
and 56m. from Kaduna on Jos Road, 15.ii.1957, 2 vials, 11m. from Kaduna on 
Zungeru Road, 28.11.1957, 24m. from Jos on Pankshin Road, 3.iv.1957, and 27.x. 
1957, 2 vials, 25-32m. from Jos on Kaduna Road, 5.iv.1957, 2 vials, 7m. from 
Bauchi on Gombe Road, 7.v.1957, 3m. from Jalingo on Yola Road, 17.v.1957, 114m. 
S.W. of Jalingo on Takum Road via Beli, 18.v.1957, 4-6m. S. of Ilorin on Oyo Road, 
4.X11.1957, 2 vials, 12m. from Keffi on Makurdi Road, 24.11.1958, 4m. from Gboko 
on Oturkpo Road, 26.11.1958, 22m. from Lokoja on Kabba Road, 10.iii.1958, and 
Samaru, near Zaria, 10.1x.1957, Vi-vii.1959, 4 vials (W. A. Sands). 

CAMEROUN : Near Mbalmayo, Presidency halt, 18.x.1962 (G. Becker), own collec- 
tion. 


ETHIOPIAN NASUTITERMITINAE 131 


CENTRAL AFRICAN REPUBLIC : Bangui, 1948 (A. E. Emerson), Bangassou, iii.1948 
(N. A. Weber), A.M.N.H. 

ConGo : Faradje, 1912 (H. Lang, J]. P. Chapin), ALM.N.H. ; Garamba Nat. Park, 
30.vil.1952 (H. de Saeger), Inst. des Parcs Nat. du Congo, Brussels. 

UcanDaA : Ngotokwe, Lango, iii.1g42, and Serere, x.1948 (W. V. Harris) ; Lale, 


b \ ) 

Dp : ) % ’ 

! ] 
i , : ye 5 Pf ‘ 
a Ts 6 
i rf 
t 
| 
; aa 

i Bn 


ae 


Map 28. Tvinervitermes trinervius. Vegetation types: 8, 16, and 17. 


132 W. A. SANDS 


Lake Kyoga near Soroti, 15.x.1952 (W. A. Sands) ; Kampala, iv.1955, 55 and 58m. 
from Kampala on Masindi Road, 2 vials, 7.x1.1955 (R. M. C. Williams). 

A total of 92 nest series were examined, and all material is in the British Museum 
(Natural History) unless otherwise stated. 


The biology of this species has been discussed under its junior synonym T. car- 
bonarius in earlier papers (Sands, 1961, 1961a). It is another of the species occurring 
in the Guinean savannah vegetation zones (Keay et al., 1959, types 8, 16, and 17). 
Unlike its sympatric congeners, its northward limit is the beginning of the drier 
Sudan savannah (Keay et al., type 20). It has therefore only been recorded from 
Uganda at the eastern end of its range, because the southern parts of the Sudan have 
been neglected by collectors. 


Trinervitermes trinervoides (Sjéstedt) 
(Text-figs. 184-187, 326-331, 359-370, 379-382 ; Map 29) 


Eutermes trinervoides Sjéstedt, 1911a : 186. [Type locality not stated]. 

Eutermes (Tvinervitermes) tyvinervoides Sjostedt ; Holmgren, 1912 : 65. 

Eutermes trinervoides SjOstedt ; Sjd6stedt, 1913 : 720. Type locality : REPUBLIC oF SOUTH 
AFrica, Cape Province, Laingsburg. 

Eutermes (Trinervitermes) trinerviformis Holmgren, 1913 : 336. Type locality : REPUBLIC OF 
SoutH Arrica, Natal, Pinetown. syn. n. 

Eutermes (Tvinervitermes) auriceps Holmgren, 1913 : 338. Type locality : REPUBLIC OF SOUTH 
AFrica, Natal, Pinetown. 

Eutermes (Ivinervitermes) bulbiceps Holmgren, 1913 : 339. Type locality : REPUBLIC OF 
SoutH Arrica, Natal, Pinetown. 

Eutermes (Trinervitermes) dubius Holmgren, 1913 : 339. Type locality : REPUBLIC OF SOUTH 
AFrica, Natal, Avoca. 

Eutermes hentschelianus Sjéstedt, t914a : 88. Type locality : SourmH West Arrica, “‘ Otjoson- 
dyombo ”’ [probably Otjosongombe, 20° 31’ S., 17° 18’ E.]. syn. n. 

Trinervitermes trinervoides (SjOstedt) Fuller, 1922 : 108, 110. 

Trinervitermes trinervifoymis Holmgren Fuller, 1922 : 111. 

Trinervitermes auriceps [= T. trinerviformis| Holmgren Fuller, 1922 : 111. 

Trinervitermes bulbiceps Holmgren [= T. tvinerviformis| Fuller, 1922 : 111. 

Trinervitermes dubius Holmgren [= T. trinerviformis] Fuller, 1922 : 111. 

Trinervitermes gemellus subsp. thomseni Fuller, 1922 : 111. Type locality : REPUBLIC OF SOUTH 
Arrica, Cape Province, Somerset West, Pearston. syn. n. 

Trinervitermes havilandi Fuller, 1922 : 113. Type locality : REPUBLIC OF SOUTH AFRICA, 
Natal, Haviland Rail. [‘‘ Teymes trineyvius Rambur’”’ Haviland, 1898 : 421]. syn. n. 

Trinervvitermes fuscus Fuller, 1922 : 113. Type locality : RepuBLic oF SouTH AFrRica, Trans- 
vaal, Warmbaths. syn. n. 

Trinervitermes kurumanensis Fuller, 1922 : 113. Type locality : REPUBLIC OF SOUTH AFRICA, 
Bechuanaland, Kuruman. syn. n. 

Eutermes (Trinervitermes) messoy SjOstedt, 1924a : 257. [n.n. for T. havilandi Fuller, reduced 
to synonymy, Snyder, 1949 : 326.] 

Trinervitermes bulbiceps Holmgren ; Sjéstedt, 1926 : 342. [ex syn. T. trinerviformis.] 

Trinervitermes dubius Holmgren ; Sjostedt, 1926 : 342. [ex syn. T. trinerviformis.] 

Trinervitermes auviceps Holmgren [= T. trinervoides]; Sjdéstedt, 1926: 346. [ex syn. T. 
tyinerutformis. | 


ETHIOPIAN NASUTITERMITINAE 133 


Trinervitermes hentschelianus (Sjostedt) Sjéstedt, 1926 : 344. 

Nasutitermes (Trinervitermes) messoy (SjOstedt) Emerson, 1928 : 416. 

Trinevvitermes hilli Snyder and Emerson ; in Snyder, 1949 : 327. [n.n. for Eutermes (Triner- 
vitermes) bulbiceps Holmgren, junior primary homonym of Euteymes (Eutermes) bulbiceps 
Holmgren.] syn. n. 


Imago. Head capsule, frons and vertex yellow-brown to dark sepia brown, sometimes paler 
around eyes, in front of ocelli, and bordering posterior margin of postclypeus ; vertex with or 
without one to three diverging paler streaks ; labrum yellow, postclypeus yellow to yellow- 
brown, strikingly paler than head in dark forms ; antennae yellow to yellow-brown. Pronotum 
yellow to sepia brown, meso- and metanota yellow to yellow-brown, other thoracic sclerites and 
legs yellow. Abdominal tergites brown, sepia brown around stigmata ; sternites pale yellow, 
brown around stigmata. Wings translucent yellow-brown to brown, subcosta and radius 
sector yellow to brown, outlined brown to dark brown at base ; membrane behind radius sector 
with a very weakly developed yellow streak or none ; rest of venation narrowly brown or 
outlined brown. 

Posterior margin of head capsule behind eyes varies from distinctly less than semi-circular to 
nearly semi-circular, very weakly or not sinuate immediately behind eyes ; fontanelle pale, 
contrasting with surrounding cuticle especially in dark forms, usually elongate Y-shaped, 
lateral arms short to long, often dilated terminally, middle of fontanelle sometimes slightly 
depressed ; eyes very short oval, prominent in proportion to own diameter, but small to medium- 
sized relative to head width, W/E index 3-3—3:9 , ocelli small to medium-sized, separated from 
eyes by from slightly over one-third to more than own least diameter ; postclypeus moderately 
to strongly inflated, width 1-8—2-4 times length, anterior margin straight or slightly concave, 
posterior margin convex, evenly rounded ; molar plate of right mandible with 8-10 ridges, 
usually 9 ; antennae 15—17 segmented, proportions of basal segments variable, partly subdivided 
in 15 segmented forms. 

Pale coloured ‘‘ scent ’’ gland on anterior margin of 2 fifth abdominal sternite small, 0-46— 
0:56mm. wide, 0-13—0:20mm. long ; cerci of 9 slightly shorter to slightly longer than width 
across base, conical or slightly mamilliform. 

Pilosity of vertex rather sparse, of varying length, setae not forming a “ pelt ”’ or ‘‘ mat’’. 

Measurements (43 specimens from 25 localities) in millimetres. 


Range Mean 
Head width across eyes I -61—2:03 : F 1-80 
Greatest diameter of eye 0°43-0'56 ; : 0°50 
Ocellus 3 : s O-10-0'23 X O-16-0'25 . : 0-18 X 0:24 
Ocellus toeye . ; 0:07—0'15 : 2 0-10 
Width of pronotum 1-48-1:94 : - 1-66 
Length of pronotum . 0-92-1'18 : F 0-99 
Length of hind tibia. 202-2 °47 : ‘ 2°26 
Length of fore wing. 1530-2260 : § 19°37 


Soldiers. Major soldier : head capsule orange-yellow to ferruginous, nose ferruginous orange 
to pitch black with red tip ; antennae yellow to yellow-brown. Pronotum, meso- and meta- 
nota, legs and abdominal sternites yellow, abdominal tergites yellow to brown. 

Head capsule in plan view, short oval to wider than long, often with posterior median indent- 
ation in larger specimens, nose cylindrical to weakly conical. In profile distinctly concave to 
well behind antennal socket, convex curve to occiput most pronounced at posterior extremity 
of head ; nose distinctly shorter to distinctly longer than rest of head capsule measuring from 
hind margin of antennal socket, angle of nose cone o-14 degrees ; antennae 14 segmented, II, 
III, IV, V, and VI subequal, rarely 13 segmented, when III nearly twice II. Head setae 
mainly confined to hairy nose tip ; bilaterally placed pair of minute setae sometimes present 
on vertex. 

Abdominal tergites with scattered minute setae and longer setae on hind margins, becoming 
more pronounced on posterior tergites ; sternites with scattered shorter backwardly directed 


134 


371. 


Fics. 359-378 : 


WrA. SANDS 


373. 375. 
Trinervitermes, soldier head capsule, side 


359-370, T. trinervoides ; 371-375, T. trinervius ; 


376. 378. 


and plan views, and nose tip. 
376-378, T. avabiae. 


ETHIOPIAN NASUTITERMITINAE 135 


setae, up to two-thirds length of long fine setae on posterior margins directed downwards and 
slightly forwards. 

Minor soldiers : as major soldiers, except head capsule in plan more oval, sometimes slightly 
constricted behind antennae. 

Measurements (Major soldiers, 56 specimens from 20 localities, minor soldiers, 15 specimens 
from 8 localities) in millimetres. 


Major Soldiers : Range Mean 
Head length to tip of nose. 2:04-2:96 ; : : ‘ 2°45 
Head width . : ‘ z 1-09—1°76 - - : : 1-32 
Depth of head capsule . : 0-78-I°17 : 2 : ; o-91 
Width of pronotum . : 0°55-0°84 : , - ; 0-65 
Length of pronotum : : 0:20-0:46 : : : : 0°34 
Length of hind tibia : ; 1°43-2°45 : : : ; 1°75 

Minor Soldiers : Range Mean 
Head length to tip of nose. 1:64-2:00 : : : ; 1-86 
Head width . - : ‘ 0:78-1:00 : : : F 0:82 
Depth of head capsule . : 0°59-0°75 : é ‘ : 0-65 
Width of pronotum ‘ - 0-48-0-61 - : : ; 0°52 
Length of pronotum : . 0:18-0:25 : : ; ; 0°24 
Length of hind tibia z : 1:38-1:89 : . . : 1°54 


Variation. The range of variation in this species is very great : specimens from different 
localities may differ from each other more than some genuine species from other parts of the 
African continent. 

In the imago, the extremes of colour have been given, but there is also variation in the dis- 
tribution of lighter and darker shades of pigmentation on head and body. The variation in 
shape of head capsule, size of eyes and ocelli, and degree of inflation of postclypeus may com- 
bine with colour patterns to produce a very different appearance. The soldier caste often 
provides more reliable identification in this species. 

There appear to be variant forms of imago and soldier castes occurring more frequently in 
certain ecological conditions, especially those associated with altitude. Several of these forms 
were given specific names by earlier authors, on the basis of the more limited samples of 
material available to them. These variants appear to be locally stable in limited populations, 
but it must be emphasized that in the large amount of material now examined, every gradation 
of intermediate between them has been found, and in addition a number of other minor forms of 
only very local importance have been observed. 

In the most distinctive of these variants the imago head is dark brown and the pronotum 
sepia brown, associated with a soldier caste in which the nose is rather short, slender, and dark 
in colour. This was described by Fuller (1922) as fuscus, and appears commonest at altitudes 
from 3,000-—5,000 feet. 

The name trinervoides was applied by Sjéstedt (1911a) to another form, most frequent in the 
southern part of Cape Province, at moderate to low altitudes. The imago head capsule is more 
broadly rounded posteriorly, and chestnut brown in colour ; the pronotum is yellow. The 
associated soldiers have a rather short, thicker nose, though a thinner-nosed form was described 
by Fuller (1922) as gemellus subsp. thomseni. In these southern specimens the postclypeus is 
often more inflated than in those from further north. 

The third important variant is that in which the soldier caste is large with the nose longer 
than the rest of the head ; the imago is similar in shape to fuscus although generally larger, 
with ¢vinervoides-type colouration, and often with larger eyes in proportion to its size than the 
southern specimens. This form has been named hentschelianus (Sj6stedt), trinerviformis 
(Holmgren), havilandi and kurumanensis Fuller, and hilli Snyder and Emerson, from various 
localities. It appears to be most common in the rather dry Acacia-Commiphora wooded steppe 
and the slightly less dry intermediate savannah (Keay ef al., 1959, vegetation types 25 and 20) 


136 W. A. SANDS 


of the more northern Cape Province and Transvaal at altitudes from 1,600 to 3,000—4,000 feet. 
It also occurs in parts of Natal and the Orange Free State. 

In the moist lowlands of Zululand the smallest and palest coloured specimens are found, and 
these if placed alongside the fuscus form appear completely dissimilar. The gradual change 
can be followed inland with specimens from increasing altitudes through Natal to the uplands 
of Transvaal. 


I have considered retaining the specific names of the synonyms to designate 
subspecies ; in view of the occurrence of every gradation of intermediate between 
the variants, and of apparent exceptions to the geographical distributions outlined 
above, I believe it is inadvisable to do so at present. 


The situation in T. tvinervoides would seem to be as follows : 

It is the only species of Tvinervitermes inhabiting the limits of subtropical and tem- 
perate South Africa. In so doing it has left behind congeners which might compete 
with it. South Africa has a very wide range of ecological conditions, which have 
given rise to correspondingly wide variation in this species. Whether or not the 
forms recognized constitute true allopatric subspecies, sections of one or more 
clines, or genetically heterogeneous variants of similar appearance induced by similar 
ecological conditions in different areas (‘‘ polytopic ’’ subspecies, Mayr, 1953) can 
only be shown by further studies. It may be noted that a “ fuscus ’’-like form of 
T. dispar has also been found at an altitude of 3,000—5,000 feet in Natal, perhaps lend- 
ing weight to the latter suggestion. It has also been observed that the nose-length of 
soldiers of all species and genera tends to be greater in the drier steppe vegetation 
of the interior. ; 

The smaller specimens of T. trinervoides resemble T. rapulwm in both imago and 
soldier castes. In addition to the characters given in the keys, the imago has usually 
a larger and differently shaped fontanelle and the head viewed from above is straight 
between the ocelli rather than indented in the middle around the fontanelle. The 
soldier has a much more hairy nose tip, and is distinguished from both T. rapulum 
and T. rhodesiensis by the rounder plan view of the head capsule. The imago 
differs from 7. rhodesiensis in having the ocelli further from the eye, and the ventral 
“scent ’’ gland distinctly smaller. 

The larger, more typical specimens of 7. trinervoides are unlikely to be confused 
with either of their partly sympatric congeners. The g or 10 ridges on the molar 
plate of the right imago mandible are a useful additional character, and the @ cerci 
are usually shorter. 


Type Material. REPUBLIC OF SOUTH AFRICA: Cape Province, Laingsburg, no 
date (Brunnthaler), syntype 2 imago and soldiers, type colony, T. trinervoides 
(Sjostedt), A.M.N.H., (other syntypes in Naturhist. Riksmus., Stockholm) ; Natal, 
Pinetown, 1905 (J. Tvrdgardh), syntype soldier, type colony, T. trinerviformis 
Holmgren, A.M.N.H. and B.M.(N.H.), (other syntypes in Naturhist. Riksmus., 
Stockholm) ; Cape Province, Somerset West, Pearston, xi.1g20 (Ff. Thomsen), 
syntype 2 imago and soldier, type colony, T. gemellus ssp. thomsent Fuller, A.M.N.H. 
(other syntypes in N.C.I., Pretoria) ; Natal, Haviland Rail, 1894 (G. D. Haviland), 
syntype soldiers from type colony, T. havilandi Fuller, A.M.N.H. and B.M.(N.H.), 


ETHIOPIAN NASUTITERMITINAE 137 


(other syntypes in N.C.I., Pretoria) ; Transvaal, Warmbaths, xi.1g20 and De 
Wildt, xi.1920 (C. Fuller), syntype g and 2 imagos and soldiers, 7. fuscus Fuller, 
A.M.N.H. and B.M.(N.H.), (other syntypes in N.C.I., Pretoria) ; Cape Province, 
Kuruman, xi.1g16 (Ff. Thomsen), syntype soldiers, type colony, T. kurwmanensis 
Fuller, A.M.N.H. and B.M.(N.H.), (other syntypes in N.C.I., Pretoria) ; Natal, 
Haviland Rail, 1894 (G. D. Haviland), syntype queen and soldiers, type colony, 


a i hl NEP 


Map 29. Tvrinervitermes trinervoides. Vegetation types: 6, 9, 15, 20, 25, 28, and 29. 


138 WA SANDS 


E.(T). messor Sjéstedt, A.M.N.H. and B.M.(N.H.) ; Natal, Pinetown, 1905 (J. 
Trdgardh), syntype, type colony, T. Milli Snyder and Emerson (T. bulbiceps 
Holmgren), A.M.N.H. (other syntypes in Naturhist. Riksmus., Stockholm). 

SoutH West AFrica: “‘ Otjosondyombo am Eiab”’, vi.tgog (O. Hentschel) 
[probably Otjosongombe, 20° 31’ S., 17° 18’ E.], syntype soldiers, type colony, T. 
hentschelianus (Sj6stedt), A.M.N.H., (other syntypes in Mus. Hamburg and Natur- 
hist. Riksmus., Stockholm). 


Other Material. SOUTHERN RHODESIA: Salisbury, 16.xii.1936, and Inyanga, 
1.1939 (A. Cuthbertson). 

MOZAMBIQUE : Chinavane, Lourenzo Marques, 18.x1i.1918 (C. Fuller) ; 17.x.1952 
(W. V. Harms). 

SWAZILAND : no data (J. B. Buchanan) ; N. of Umbuluzi R., vi.1g21 (C. Fuller) ; 
7 and 13m. from Mbabane on Bremersdorp Road, and 2m. from junction of Piggs 
Peak and Lochiel-Mbabane Roads, 8.vi.1956 (J. H. Grobler) ; Piggs Peak to Komati 
R., 22—3.x.1960, 4 vials, 4m. from Stegi on Gollel Road, 24.x.1960, 2 vials, 6m. from 
Gollel on Hluti Road, gm. from Goedgegund on Mankaiana Road, and 2m. from 
Mankaiana on Mbabane Road, 25.x.1g60 (W. G. H. Coaton and J. L. Sheasby). 

BASUTOLAND : no detailed locality, I.xi1.1939 (no collector). 

REPUBLIC OF SOUTH AFRIcA : In addition to that already at the B.M.(N.H.), all 
the material from the N.C.I., Pretoria has been received and examined. This 
covers every part of the Republic, and there is no point in giving details of localities 
or collectors, although a list is available at the B.M.(N.H.). The numbers examined 
have been as follows : Transvaal, 123 nest series ; Orange Free State, 83 nest series ; 
Cape Province, 406 nest series. A total of exactly 800 nest series of this species 
were examined, and all material is in the British Museum (Natural History) unless 
otherwise stated. 


T. trinervoides is the commonest species south of the River Limpopo in sub- 
tropical and temperate South Africa, and its most northerly records are from 
Salisbury, Southern Rhodesia. It covers a wide range of climatic and vegetation 
types, from the dry subdesert steppe of the interior of Cape Province to the moist 
tropical coastal savannah forest mosaic of Natal and Mozambique (Keay ef al., 
1959, types 6, 9, 15, 20, 25, 28, 29, and 30). 

The mounds of this species have been described as varying from hemispherical up 
to 6 feet in diameter, to columnar, 6 feet high. Quantities of cut grass are stored 
in the mounds as in many other species. 


BAUCALIOTERMES gen. n. 


(Baucalion, Gr. “‘ narrow-necked bottle ’’). 
Type-species, Subulitermes hainesi Fuller. 


Imago. Left mandible with apical and first marginal teeth subequal, distance from apical to 
first marginal (worn specimen) slightly less than one-third distance between first and third 
marginals (index -30), second marginal obsolete, cutting edge between first and third marginals 
entire, straight ; right mandible with apical and first marginal subequal as left, cutting posterior 
edge of first marginal longer than that of second marginal, molar plate with no marked irregu- 


ETHIOPIAN NASUTITERMITINAE 139 


larity in surface view, and 7-8 transverse ridges, first much larger than rest, forming a blunt 
tooth, the more posterior ridges less prominent. Postclypeus, length slightly less than half 
width. Fontanelle in g, small, pale, irregularly Y-shaped ; in 9, trapezoid with lateral arms 
extending from anterior corners and slightly dilated terminally ; both sexes with smooth 
shallowly depressed area behind fontanelle coloured as rest of head. Pilosity of head capsule 
uneven, sockets of varying size from small to large, the latter numerous and conspicuous. 

Soldier. Monomorphic. Nose very thin and cylindrical, at most slightly tapering. Vestigial 
mandibles without points. Head capsule not constricted. Antennae 12-14 segmented, usually 
14. Sockets of minute setae clearly visible as pale spots scattered over entire head capsule 
including nose. 


This genus is named to accommodate the single species B. hainest, known only 
from the arid areas of Cape Province, South Africa, which was first described in the 
genus Subulitermes and latterly transferred to Trinervitermes. However, the coarser 
pilosity with more conspicuous seta sockets distinguishes it from all known Tyvinervi- 
termes, and in addition the distribution of the lighter coloured areas of the imago 
head does not agree with that genus. In the soldier, the visible seta sockets are 
again the main distinguishing feature, being absent in Trinervitermes, which also 
has a dimorphic soldier caste. It is expected that when fresh winged imagos of 
Baucaliotermes are discovered, further generic differences will become clearer ; for 
example, in the queen secondary sclerotizations of the abdomen obscure the out- 
lines of the “ scent ’’ gland on the front margin of the 5th abdominal sternite, but 
it appears to be very small, and absent in the king. In Trinervitermes this gland 
is clearly visible even when not large. 

These two genera are closely related, as is shown by the similarity of imago and 
worker mandibles, which are distinct from other genera of the Ethiopian Zoogeo- 
graphical Region. 


Baucaliotermes hainesi (Fuller) comb. n. 
(Text-figs. 196, 197, 336, 337, 386, 387, 396-398 ; Map 3) 


Subulitermes hainesi Fuller, 1922: 115. Type locality : RepuBLic oF SoutH ArrRica, Cape 
Province, Steinkopf, Wolfdoon, Kenhardt. 

Eutermes hainesi (Fuller) Sjéstedt, 1926 : 310. 

Subulitermes hainesi Fuller ; Snyder, 1949 : 339. 

Trinervitermes hainesi (Fuller) Emerson, 1960 : 2. 


Imago. Previously undescribed. Head capsule chestnut brown, but probably faded in 
morphotype queen and king ; vertex with faint diverging streaks scarcely paler than rest; 
postclypeus yellow clouded brown posteriorly, labrum yellow. Area between eye, antennal 
socket, and lower margin of head, extending along latter a little behind eye in rounded patch, 
yellow-brown. Antennae pale yellow. Pronotum yellow, yellow-brown in front of pale 
Y-shaped median patch, meso- and metanota yellow ; other thoracic sclerites and legs pale 
yellow. Abdominal tergites brown ; sternites pale yellow-brown, brown around stigmata. 

Posterior margin of head capsule behind eyes much less than semi-circular, slightly sinuate 
immediately behind eyes ; eyes somewhat angularly oval, moderately prominent in proportion 
to own diameter, small relative to head width, W/E index 3:2-3-4 ; ocelli small, oval, separated 
from eyes by under one-third to over two-fifths own least diameter ; postclypeus moderately 
inflated, width 2-o-2-4 times length, anterior margin nearly straight, posterior margin convex, 
rounded ; antennae broken, II and V subequal, and longer than III which is longer than IV. 


140 


Ws AL SANDS 


394. 395) 
391. 396. 398. 


Fries. 379-398 : 379-382, Trinervitermes trinervoides, front and side views of imago head 
capsule ; 383-385, Tvinervitermes vertex pilosity (383, T. tvinervius, 384, T. oeconomus, 
385, IT. geminatus) ; 386, 387, and 396-398, Baucaliotermes hainesi, imago and soldier 
head capsules ; 388-395, Mycterotermes meringocephalus, head capsules of major and 
minor soldiers (391, vertex pilosity, 392, 393, major soldier mandibles). 


ETHIOPIAN NASUTITERMITINAE 141 


Cerci of 2 longer than width across base, conical. 
Other characters are given in the generic diagnosis. 
Measurements (2 specimens from 1 locality) in millimetres. 


Range 
Head width across eyes : ; - : : 1-61—1 -66 
Greatest diameter of eye. . é : : 0°47-0°53 
Ocellus. : : 2 , : ; , 0-16 XK 0:23 
Ocellus toeye . : ; ; : : ; 0°05-0-07 
Width of pronotum . F : ; d 2 1°45-1°53 
Length of pronotum . . . : : : 0:82-0-92 
Length of hind tibia 209-2 °12 


Morphotype : REPUBLIC OF SOUTH AFRICA: 70m. from Kliprand on Pofadder 
Road, 8.iv.1963 (A. Barnard and J. N. Diederichs), morphotype queen in N.C.L., 
Pretoria. 


Soldier. Head capsule brown to dark sepia brown, paler around antennal socket, nose sepia 
brown to pitch black with yellow-brown to ferruginous tip. Pronotum and antennae yellow, 
legs yellow-white. Abdominal tergites yellow-brown, sternites pale yellow. 

Head capsule in plan view short oval, tapering slightly towards front. In profile concave, 
back of head raised and rounded well behind antennal socket ; nose equal to or slightly longer 
than rest of head capsule, measuring from hind margin of antennal socket ; antennae, in 14 
segmented form, III and V subequal, II similar length but thicker, IV and VI subequal and 
longer than these. 

Head setae consist of a bilaterally placed pair on vertex, one or two pairs near base of nose, 
four at nose tip with smaller setae behind them, and minute setae scattered over entire head 
capsule and nose. Abdominal tergites with scattered very short setae becoming longer on hind 
margins and posterior segments ; sternites with short setae, and longer on hind margins. 

Measurements (18 specimens from 3 localities) in millimetres. 


Range Mean 
Head length to tip of nose. 1:63-2:-09 F _ : : 1:87 
Head width . : 2 ‘ O°79-1°15 . ‘ : ; 0:98 
Depth of head capsule . 2 0+59-0°79 ; ; . : 0-68 
Width of pronotum : : 0:46-0:56 : ; : : o-51 
Length of pronotum : : 0-23-0-28 : 7 ‘ . 0:25 
Length of hind tibia ; ; 1:25-1'79 : ‘ : : 1°55 


The relationships and distinctive features of B. hainest have been discussed under 
the generic heading. 

Material. REPUBLIC OF SOUTH AFRICA : Bushmanland, Wolfdoon, and Wortel, 
near Gella, r and 5.ix.1g17 (F. Thomsen), syntype soldiers, B. hainesi (Fuller), 
A.M.N.H. and B.M.(N.H.) ; 16m. from Pella on Springbok Road, 15.x.1g61 (J. L. 
Sheasby) ; 70m. from Kliprand on Pofadder Road, 8.iv.1963, further vial with king 
(A. Barnard, J. N. Diederichs). 


EUTERMELLUS Silvestri 


Eutermellus Silvestri, 1912 : 243. 

Eutermes (Eutermellus) Holmgren, 1912 : 64. 

Eutermellus SjOstedt, 1926 : 358. 

Type species, by original designation, Euteymellus convergens Silvestri, 1912. 


Imago. Left mandible with apical tooth distinctly longer than first marginal, but distance 
between them slightly shorter than that from first to third marginals (left mandible index 


142 W. A. SANDS 


-96—-98), cutting edge between the latter sinuate and indented in front of third marginal, which 
is approximately equilateral, small subsidiary tooth just visible in gap between third marginal 
and molar prominence ; right mandible, apical and first marginal as left, second marginal small 
but distinct with concave posterior edge, molar plate in surface view without basal indentation, 
asymmetrical with large posterior rounded flange and smaller anterior, weakly ridged. Post- 
clypeus inflated, width 1-8-2-5 times length. Fontanelle variable, Y-shaped, triangular, 
dumb-bell-shaped or near obsolete. Eyes scarcely to distinctly set out from sides of head. 
Pilosity of head not particularly regular in length, with numbers of conspicuous long setae 
tending to occur in symmetrical pairs. Pronotum, sides weakly convergent to rounded posterior 
margin, length approximately two-thirds width. 

Soldiey. Monomorphic. Nose conical to cylindrical, tapering to fine point, distinctly rugose. 
Vestigial mandibles with minute points or without, labrum with tri-lobed anterior margin. 
Head capsule very weakly to moderately constricted behind antennae. Antennae 13 segmented. 
Head and nose with scattered fine very short setae, longer at nose tip, and conspicuous longer 
setae arranged approximately in two rows, one in front and one behind the constriction, front 
row with 4-10 setae ; posterior row with 2-10. Rest of body with scattered setae of irregular 
length, longer on posterior margins of tergites and sternites, and towards rear of abdomen, legs 
with conspicuous long setae, 1 on each coxa, 3-6 on each femur and tibia, among the shorter 
more typical setae. 

Worker. Mandibles as imago except apical tooth longer in proportion (left mandible index 
1:24-1:62) and molar areas more pronounced. 


Eutermellus is in many respects the least specialized member of the “‘ Paracorni- 
termes-branch ”’ occurring in Africa, as shown by the imago and worker mandibles, 
the typically Y-shaped fontanelle of some species, the tri-lobed soldier labrum and 
the generally more numerous setae. The soldier mandibles were described in 
detail in another paper (Sands, 1956). It is distinguishable from Verrucositermes 
and Afrosubulitermes in the imago by its denser pilosity, and in the latter case 
by its greater size. The soldier is separated from these two genera both by its 
general shape and size, and by the numerous prominent setae on head, body, and 
legs. 

Eutermellus as used here includes three new species and is of interest because the 
inter-specific variation appears to indicate relationships to other genera. E. 
undulans has a slender nose in the soldier, combined with an imago in which the eyes 
are not noticeably set out from the sides of the head capsule, and the posterior margin 
of the postclypeus is indistinct, merging in profile with the line of the frons. This 
suggests affinities with Afrosubulitermes and Postsubulitermes. Other species such 
as EF. abruptus and E. bipartitus have the eyes set out from the sides of the head in a 
similar manner to Verrucositermes and Mimeutermes, whilst in E. convergens and E. 
aquilinus the more conical nose of the soldier and almost unconstricted head 
capsule also suggest a relationship to Mimeutermes. These distinctions between 
the species, though interesting, are not very marked, especially in the soldier castes. 
Though separable on the basis of the material known at present, from widely spaced 
localities, it is possible that some of the species named here might be found to be 
synonymous if further material were to be collected. 

This genus has been recorded from both the Congo and West African rain forests 
and from the adjacent savannah zones, but not from the savannah south of the 
Congo nor from East or South Africa. It is found in the mounds of other genera of 
termites and builds none of its own. 


ETHIOPIAN NASUTITERMITINAE 143 


Fics. 399-413 : Imago head capsule, front and side views. 399, 400, Euteymellus abyup- 
tus ; 401, 402, E. aquilinus ; 403, 404, E. bipartitus ; 405, 406, E. convergens ; 407, 
408, E. undulans ; 409-411, Afrosubulitermes congoensis (411, pronotum, plan view) ; 
412, 413, Postsubulitermes parviconstrictus. 


W. A. SANDS 


425. 427 


Fics. 414-430: Soldier head capsule, side and plan views, labrum, and mandibles. 414— 
416, Eutermellus abruptus ; 417-419, E. aquilinus ; 420, 421, E. bipartitus ; 422-424, 
E. convergens ; 425-427, E. undulans ; 428-430, Afrosubulitermes congoensis. 


ETHIOPIAN NASUTITERMITINAE 145 


KEYS TO SPECIES 


IMAGOS 


I Ocelli smaller, 0:07-0:09 xX 0-10-0-13, separated from eyes by half own least 


diameter or more 2 
— Ocelli larger, 0-09-0-1I X O-l nee ‘16, “separated from ‘eyes be less than half own 

least diameter ‘ 3 
2 Fontanelle a pale streak, slightly to distinctly bifurcate in front. Postclypeus less 

inflated, width 2-5 times length : : -  convergens (p. 149) 
— Fontanelle minute, triangular or 3 spots aeameed 3 in triangle, slightly paler than head 

capsule. Postclypeus more inflated, width 2-o-2-1 times length abruptus (p. 145) 


3 Larger; W, 0:93-0:99; E, 0-34-0-38. Fontanelle irregularly bilobed, sometimes 
divided into 3 spots, distinctly paler than head. Postclypeus more inflated, width 
1-8-1-9 times length, anterior margin straight, oe arched in middle, sinuate 
towards outer corners . F : : aquilinus (p. 147) 

— Smaller; W, 0:85-0:88 ; E, 0-31-0°33. Fontancile an elongate rounded triangle, 
slightly paler than head, or almost obsolete. Postclypeus less inflated, width 2-1 
times length, anterior margin very eed convex or sinuate, posterior regularly 


rounded indistinct. E : : : : 3 : ; 2 4 
amo ves, 023% ; Ocelli,o-09 x 0-13 ; Ts 1:20. Fontanelle an elongate rounded triangle 
(S. Congo) . ; , : : : bipartitus (p. 148) 
— Eyes, 0:33; Ocelli, O10, x 0-14 ; Ls; ¥00-T-00. Fontanelle almost obsolete 
(Guinean savannah, W. Africa) : ‘ : : : : undulans (p. 150) 
SOLDIERS 


1 Posterior lobe of head capsule behind constriction with 1-2 pairs of long conspicuous 
setae, nose thicker, conical, profile nearly straight 
aquilinus (W. Africa, p. 147) and bipartitus (S. Congo, p. 148) 
— Posterior lobe of head capsule with 3-5 pairs of long conspicuous setae. In two of 


three species, nose cylindrical or thinner conical, profile undulating ; remaining 

species, nose conical, profile nearly straight : ; ‘ : 2 
2 Nose thicker conical, profile nearly straight. T, shorter, aeoria (one only over 

0-70). Antennal socket almost without ridge on dorsal side .  convergens (p. 149) 
— Nose thinner, cylindrical or conical, profile undulating. T, longer, epics 

Antennal socket usually with ridge on dorsal side. 3 3 
3 Highest point of “‘hump”’ at nose base vertically above middle of postinentum in 

profile, and abruptly depressed behind (Congo forest species) . abruptus (p. 145) 
— Highest point of “hump ’”’ in front of middle of postmentum, profile more evenly 

undulating. (Guinean savannah species, W. Africa) : ; undulans (p. 150) 


Eutermellus abruptus sp. n. 
(Text-figs. 399, 400, 414-416, 431-433, 479-481; Map 30) 


Imago. Head capsule sepia brown with paler markings on frons, postclypeus and antennae 
yellow-brown, labrum yellow. Pronotum, meso- and metanota brown, other thoracic sclerites 
and legs yellow. Abdominal tergites brown, sternites yellow-brown. 

Posterior margin of head capsule behind eyes evenly rounded, slightly less than semi-circular, 
turned slightly outwards at eye rim ; fontanelle, paler than head capsule, minute triangular or 
three spots arranged in triangle ; eyes oval, slightly set out from sides of head, prominent in 
proportion to own diameter and large relative to head width, W/E index 2-6-2-9 ; ocelli small 


146 W. A. SANDS 


oval, separated from eyes by about half own least diameter or slightly more ; postclypeus 
moderately inflated, width about twice length, anterior margin sinuate, posterior convex, 
evenly rounded ; antennae 15 segmented, II, IV and V subequal, III shorter. 

Pilosity of head rather even and dense, but not sufficently so to form a “ pelt’ or ‘“‘ mat ”’ 
pronotum and abdominal pilosity similar, longer towards margins of sclerites. 

Measurements (3 specimens from 2 localities) in millimetres. 


Range 
Head width across eyes : : : 0-88-0-94 
Greatest diameter of eye . : : 0:33-0°35 
Ocellus . : : c : ‘ 0:08-0:09 X O-II-0'13 
Ocellus toeye . : : : : 0:04-0'05 
Width of pronotum . : ; : 0-73-0°79 
Length of pronotum . - : ‘ 0°49-0°54 
Length of hind tibia . ‘ : : I-10—-I-23 


Soldier. Head capsule pale yellow to yellow, nose orange-yellow to orange, rest of body 
cuticle pale yellow. 

Head capsule in plan view weakly constricted behind antennae, posterior margin evenly or 
slightly angularly rounded ; nose thin, weakly conical. In profile undulating with distinct 
hump at base of nose, to nearly straight with depression in front of evenly rounded posterior 
lobe, highest point of hump vertically above middle of postmentum, and somewhat abruptly 
depressed behind ; antennae, II, III and V subequal, IV shorter, antennal socket usually 
with small ridge over dorsal rim. 

Conspicuous setae of head consist of six in three pairs in line across base of nose in front of 
constriction, and six in three pairs similarly arranged on posterior lobe behind constriction. 

Measurements (7 specimens from 3 localities) in millimetres. 


Range Mean 
Head length totipof nose. I-25-1°40 : . ‘ : 1°34 
Head width . : 3 : 0-61-0°73 ‘ 2 : : 0-67 
Depth of head capsule . : 0:38-0:46 : : : : 0-42 
Width of pronotum : : 0:34-0:38 3 : : 4 0:36 
Length of pronotum : ; 0-13-0°14 3 ‘ ‘ E 0-13 
Length of hind tibia . : 0-71-0°83 c : : 0°74 


E. abruptus differs from E. bipartitus in the imago in eaten the ocelli slightly 
more distant from the eyes, and a differently shaped fontanelle ; the head capsule 
has a slightly more broadly rounded posterior margin. The soldier of E. abruptus 
has a thinner nose and more numerous head setae. Differences from E. aquilinus, 
E. convergens and E. undulans are discussed under those species. 


Holotype 2 imago, Conco: Njili, 18km. from Leopoldville, 11.vi.1948 (A. E. 
Emerson), in American Museum of Natural History. 


Morphotype soldier, same data, A.M.N.H. 


Paratype soldiers from type colony, same data, in A.M.N.H. and British Museum 
(Nat. Hist.). 


Other paratype material. Conco: Sona Mpangu, 10.iv.1948, soldiers and 
workers, A.M.N.H. and B.M.(N.H.), and Camp Putnam, 1° 24’ N., 28° 36’ E., 
18.v.1948, 2 vials, one containing soldiers and workers, and the other, two imagos 
that flew to light (A. E. Emerson), allotype $ imago in A.M.N.H. 


ETHIOPIAN NASUTITERMITINAE 147 


Eutermellus aquilinus sp. n. 
(Text-figs. 401, 402, 417-419, 482, 483 ; Map 30) 


Imago. Head capsule sepia brown with paler markings on frons, postclypeus and antennae 
yellow-brown, labrum yellow. Pronotum brown, meso- and metanota and femora yellow- 
brown, tibiae and tarsi yellow. Abdominal tergites brown, sternites yellow-brown. Wing 
membrane translucent very pale brown, venation brown. 

Posterior margin of head capsule behind eyes rounded, more strongly curved towards sides, 
and turned slightly outwards at eye rim ; fontanelle pale, irregularly bilobed with triangular 
middle section, lateral arms variable in length and shape, sometimes separated from middle 
section ; eyes scarcely set out from sides of head, oval, prominent in proportion to own dia- 
meter and large relative to head width, W/E index 2-5-2-8 ; ocelli large, oval, separated from 
eyes by less than one-third own least diameter ; postclypeus strongly inflated, width 1-8-1-9 
times length, anterior margin straight, posterior, convex, arched in middle, sinuate towards 
outer corners ; antennae 15 segmented, II, IV, and V subequal, III very small. 

Pilosity of head rather even and dense but not sufficiently so to form a “ pelt "’ or ‘‘ mat’, 
pronotum and abdominal pilosity similar, longer setae towards margins of sclerites. 

Measurements (8 specimens from 2 localities) in millimetres. 


Range Mean 
Head width across eyes 0:93-0°99 ‘ : 0:94 
Greatest diameter of eye 0°34—0°38 : , 0-36 
Ocellus : 2 : O‘I0-O'II X O-I4-O0'16 . : O-IO X O15 
Ocellus toeye . : 0:02-0'03 : ‘ 0-02 
Width of pronotum . 0:71-0:78 : : 0°74 
Length of pronotum . 0*50-0°56 F ; 0°53 
Length of hind tibia. 1:08-1:20 : : I-12 
Length of fore wing. 760-8 -60 ; 8-12 


Soldier. Head capsule pale yellow to yellow, nose orange-yellow to orange, rest of body cuticle 
pale yellow. 

Head capsule in plan view scarcely constricted behind antennae, posterior margin somewhat 
angularly rounded, nose rather thick, weakly conical. In profile almost straight or slightly 
convex, sometimes weakly humped at base of nose ; antennae, II, III, and V subequal, IV 
shorter. Conspicuous setae of head consist of six in three pairs arranged approximately in line 
across base of nose in front of ‘‘ constriction ’’ and one or two pairs on vertex behind constriction. 

Measurements (8 specimens from 2 localities) in millimetres. 


Range Mean 
Head length to tip of nose. 1-28-1°38 ; : : E 1°32 
Head width . : ; ; 0-60-0-65 : : : ; 0°62 
Depth of head capsule . ; 0°39-0°45 ‘ : : P o-4I 
Width of pronotum : ; 0°35 
Length of pronotum : : 0-13 
Length of hind tibia : : 0:68-0:70 : = : : 0:70 


E. aquilinus is separable from EF. bipartitus in the imago by the larger size, 
differently shaped fontanelle, more inflated postclypeus, and by the eyes being less 
distinctly set out from the sides of the head. The soldiers have, however, been 
found indistinguishable, and in this case the distribution is the only criterion. In 
the imago caste of the other species, E. convergens has the ocelli more distant from 
the eyes, a less inflated postclypeus, and a differently shaped fontanelle, FE. wndulans 
is smaller, with almost obsolete fontanelle and a differently shaped head capsule, 
and E. abruptus has smaller ocelli, more distant from the eyes, and a different 
fontanelle. The soldiers are all very much alike, but the remaining species all have 


148 We AS SANDS 


3-5 pairs of long setae on the vertex behind the constriction, and E. abruptus and 
E. undulans have thinner noses with different profile. 


Holotype 9 imago, allotype 3, morphotype soldier, and paratypes from type 
colony. GHANA: 20m. from Bolgatanga on Tamale Road, 9.i1.1959 (W. A. 
Sands, Coll. No. S.2364), in British Museum (Natural History). 


Other paratype material. GHANA: Awura, 4m. from Ejura on Kintampo Road, 
28.11.1959 (W. A. Sands), soldiers and workers. 

NIGERIA : Northern Region ; Samaru, 7m. from Zaria on Funtua Road, 17.Vv. 
1959 (W. A. Sands), imagos. Both in B.M.(N.H.). 

This species has been recorded from the edge of the forest-zone to the Northern 
Guinean savannah (Keay ef al., 1959, vegetation types 8, 16, and 17). 


Eutermellus bipartitus (Sjéstedt) 
(Text-figs. 403, 404, 420, 421, 484 ; Map 30) 
Eutermes bipartitus SjOstedt, 1911 : 168. Type locality : Conco, Mukimbungu. 


Eutermes (Coarctotermes) bipartitus Sjéstedt ; Holmgren, 1912 : 65. 
Eutermellus bipartitus (SjOstedt) Sjéstedt, 1926 : 358. 


Imago. Colour probably faded in old specimen. Described by Sj6stedt as having head 
capsule chestnut brown, dorsal sclerites pale red-brown. 

Posterior margin of head capsule evenly rounded, less than semi-circular, turned slightly 
outwards at eye rim ; fontanelle pale, elongate triangular, somewhat rounded, with indistinct 
oval mark in front ; eyes oval, slightly but distinctly set out from sides of head, prominent in 
proportion to own diameter and fairly large relative to head width, W/E index 2:9 ; ocelli 
medium-sized, oval, separated from eyes by slightly more than one-third own least diameter ; 
postclypeus moderately inflated, width 2-1 times length, anterior margin nearly straight, 
posterior convex, evenly rounded ; antennae probably 15 segmented, II, IV, and V subequal, 
III shorter. 

Pilosity of head dense, not sufficiently even to form a “ pelt’ or “ mat’’, pronotum and 
abdominal pilosity similar, longer towards margins of sclerites. 

Measurements (1 specimen) in millimetres. 


Gs 


Head width across eyes . ; : ; : 0:89 
Greatest diameter of eye. , : : : 0-31 
Ocellus. ; : ; : A : é 0:09 X 0°13 
Ocellus toeye . : : ‘ é ; ‘ 0:03 
Width of pronotum . : é : : : 0:68 
Length of pronotum . , , : , s 0-46 
Length of hind tibia . : : 1-20 


Soldiey. Head capsule pale yellow to yellow, nose orange-yellow to orange, rest of body 
cuticle pale yellow. 

Head capsule in plan view weakly constricted behind antennae, posterior margin evenly 
rounded ; nose rather thick, conical. In profile almost straight or slightly convex, with slight 
depression behind base of nose and in front of evenly rounded posterior lobe ; antennae 13 
segmented, II, III and V subequal, IV shorter. 

Conspicuous setae of head consist of four to six in two or three pairs in line across base of nose 
in front of constriction, and one or two pairs similarly arranged on posterior lobe behind con- 
striction. 


ETHIOPIAN NASUTITERMITINAE 149 


Measurements (4 specimens from 1 locality) in millimetres. 


Range Mean 
Head length to tip of nose. I-25-1°40 : : é ‘ E31 
Head width . : ‘ : 0:59-0:66 : - : . 0-62 
Depth of head capsule . - 0-33-0'41 ; : ; : 0:38 
Width of pronotum : : 0°29-0°35 ; : i ‘ 0°32 
Length of pronotum : : O-II-O'14 : : : : O-12 
Length of hind tibia ; : 0-61-0°74 ; j : : 0-68 


E. bipartitus differs from its earliest congener E. convergens in the imago in having 
the ocelli closer to the eyes, and the eyes more distinctly set out from the sides of the 
head ; the postclypeus is more inflated. The soldier of E. bipartitus, whilst similar 
in shape, appears to have fewer head setae. Differences from the new members of 
the genus, FE. abruptus, E. aqulinus, and E. undulans, are discussed under those 
species. 


Material. Conco: Mukimbungu, 5° S., 14° E., 9.x.1904, Coll. No. 1158 (K. E. 
Laman), syntype male imago and soldiers from type colony, EF. bipartitus (Sjéstedt) ; 
same locality, 19.x.1904, and 22.x.1904, 2 vials (K. E. Laman), the first of these 
containing further syntypes. All in A.M.N.H. 


This species is only known from the original material, but it appears possible that 
it may be a savannah species south of the Congo forest, equivalent in its ecology to 
E. aquilinus in the north. 


Eutermellus convergens Silvestri 
(Text-figs. 405, 406, 422-424, 485 ; Map 30) 


Eutermellus convergens Silvestri, 1912: 245. Type locality: ISLAND oF FERNANDO Poo, 

Basile. 

Eutermes (Eutermellus) convergens (Silvestri) Holmgren, 1912 : 64. 
Eutermellus convergens Silvestri ; Sjéstedt, 1926 : 358. [States convergens syn. of bipartitus 

(Sjostedt) ]. 

Eutermellus convergens Silvestri ; Snyder, 1949 : 344. [States convergens not. syn. of bipartitus 

(Sj6stedt)]. 

Imago. Head capsule dark sepia brown, with paler markings on frons, postclypeus sepia 
brown, labrum and antennae brown. Pronotum sepia brown, meso- and metanota brown, 
other thoracic sclerites and legs yellow-brown to brown. Abdominal tergites sepia brown, 
sternites brown. Wing membrane translucent very pale brown, venation brown. 

Posterior margin of head capsule behind eyes evenly rounded, less than semi-circular, turned 
slightly outwards at eye rim ; fontanelle pale, slit-like and slightly bifurcate in front to Y- 
shaped with lateral arms dilated or branching terminally ; eyes slightly set out from sides of 
head, oval, moderately prominent in proportion to own diameter and large or medium-sized 
relative to head width, W/E index 2-6-3-0 ; ocelli small, separated from eyes by three-quarters 
own least diameter ; postclypeus moderately inflated, width 2:5 times length, anterior margin 
slightly convex or sinuate, posterior convex rounded, somewhat indistinct ; antennae 15 
segmented, II, IV and V subequal, III shorter. 

Pilosity of head uneven, and dense not forming a “ pelt’’ or ‘‘mat’’. Pronotum and 
abdominal pilosity similar, longer setae towards margins of tergites. 


150 WW. A. SANDS 


Measurements (3 specimens from 2 localities) in millimetres. 


Range 

Head width across eyes : 5 : 0°88 
Greatest diameter of eye . : ; 0:29-0°33 
Ocellus. : ; : : : 0:07-0:08 X O-IO-O'II 
Ocellus toeye . : : : : 0:05-0:06 
Width of pronotum . ; : : 0:69-0:70 
Length of pronotum . : : : 0-44-0°48 
Length of hind tibia . : I-03-I-‘10 


Soldier. Head capsule pale yellow to ice, nose orange-yellow to orange, rest of body 
cuticle pale yellow. 

Head capsule in plan view constricted behind antennae, posterior margin evenly rounded ; 
nose rather thick, conical. In profile nearly straight, sometimes with slight depression in front 
of rounded posterior lobe ; antennae, II, III and V subequal, IV shorter. Conspicuous setae 
of head consist of six to eight in three or four pairs approximately in line across base of nose in 
front of constriction, and a similar number on posterior lobe behind constriction. 

Measurements (13 specimens from 5 localities) in millimetres. 


Range Mean 
Head length to tip of nose. I -23-1°39 : : . : I-30 
Head width . - : : 0-55-0°68 : : ; 7 0-61 
Depth of head capsule . : 038-040 : : : : 0-39 
Width of pronotum : ; 0:30-0:36 ; 3 ‘ : 0°34 
Length of pronotum ; : O-II-O'15 : : : : O'l3 
Length of hind tibia 3 : 0°57-0°72 : . 0-66 


E. convergens differs in the imago from E. bipartitus = the smaller ocelli, more 
distant from the eyes, slightly less inflated postclypeus, and shorter hind tibia. 
E. abruptus has a different fontanelle and more inflated postclypeus. The differences 
from E. aquilinus and E. uwndulans are discussed under those species. In the soldier, 
E. aquilinus and E. bipartitus have fewer head setae, and E. abruptus and E. undulans 
thinner noses. 


Material. ISLAND OF FERNANDO Poo: Basilé, date unknown (L. Fea), syntype 
imago and soldiers, type colony, E. convergens Silvestri, A.M.N.H. 

GHANA : Aburi, 1912-13 (F. Silvestri), Silvestri Coll., Lab. Zool. Sc. Agr., Portici. 

NIGERIA : Western Region ; 63m. S. of Ilorin on Oyo Road, 4.xii.1957, and 
Ibadan, Agodi, 10.xii.1957 (W. A. Sands) ; Benin Province, Obanokoro, Sobo 
Plain, 7.1.1957 (W. Wilkinson). 

CAMEROUN : Kumba, I9.x.1949 (H. Oldroyd). 

Material in British Museum (Natural History) unless otherwise stated. 


E. convergens appears to be a forest species, with only one record in riverain forest 
outside the main rain forest block of West Africa. 


Eutermellus undulans sp. n. 
(Text-figs. 407, 408, 425-427, 486, 487 ; Map 30) 


Imago. Head capsule sepia brown with paler markings on frons, postclypeus and antennae 
yellow-brown, labrum yellow. Pronotum, meso- and metanota brown, other thoracic sclerites 
and legs yellow. Abdominal tergites brown, sternites brown laterally, paler in mid-line. Wing 
membrane translucent very pale brown, venation brown, radius-sector outlined brown. 


ETHIOPIAN NASUTITERMITINAE 151 


Posterior margin of head capsule behind eyes rounded, less than semi-circular, turned slightly 
outwards at eye rim ; fontanelle almost obsolete, very indistinct, shape uncertain ; eyes oval, 
scarcely set out from sides of head, moderately prominent in proportion to own diameter, large 
relative to head width, W/E index 2-6-2-7 ; ocelli medium-sized, oval, separated from eye by 
less than one-fifth own least diameter ; postclypeus moderately inflated, width about twice 
length, anterior margin slightly convex, posterior convex, rounded, very indistinct ; antennae 
15 segmented, II, [LV and V subequal, III shorter. 

Pilosity of head rather even and dense, but not sufficiently so to form a “‘ pelt ’’ or “‘ mat’, 
pronotum and abdominal pilosity similar, longer towards margins of sclerites. 

Measurements (2 specimens from 1 locality) in millimetres. 


Range 

Head width across eyes : : 5 ; F 0:85-0°88 
Greatest diameter ofeye_ . F ‘ : ; 0°33 
Ocellus . d ; : : : : : O-10 X O-14 
Ocellus toeye . 5 : ; : “ d 0:02 
Width of pronotum . : : : - : 0-69-0°71 
Length of pronotum . - : : : : 0-50 
Length of hind tibia . - . : < : 1°00-I ‘09 
Length of fore wing . : : : ‘ ; 6:80-7:20 


Soldier. Head capsule pale yellow to yellow, nose orange-yellow to orange, rest of body 
cuticle pale yellow. 

Head capsule in plan view distinctly constricted behind antennae, posterior margin evenly 
rounded ; nose thin and cylindrical or slightly tapering. In profile undulating with distinct 
hump at base of nose, to nearly straight, with weak depression in front of evenly rounded pos- 
terior lobe ; antennae, II, III and V subequal, IV shorter, antennal socket usually with small 
ridge over dorsal rim. Conspicuous setae of head consist of six to ten in three to five pairs 
approximately in line across base of nose in front of constriction, and three to five pairs similarly 
arranged on posterior lobe behind constriction, the latter being less regular in position. 

Measurements (16 specimens from 4 localities) in millimetres. 


Range Mean 
Head length to tip of nose. 1:20-1:35 : : : : 1:29 
Head width . : : : 0+58—0-64 : - . : 0-62 
Depth of head capsule . - 038-043 : t é . o-41 
Width of pronotum : : 0°31-0°35 . : : : 0°34 
Length of pronotum ; : O-II-O'13 : : : = O12 
Length of hind tibia P ; 0-68-0-75 : : . : 0-69 


E. undulans differs in the imago from E. bipartitus in the almost obsolete fontanelle 
and shorter hind tibia. EE. convergens has a conspicuous fontanelle and smaller 
ocelli more distant from the eyes, as is also the case with E. abruptus. The differ- 
ences from E. aquilinus are discussed under that species. In the soldier, EF. abruptus 
is closest, differing only slightly in the more abrupt depression behind the base of the 
nose. £. convergens has a thicker nose, and the other two species fewer head setae. 


Holotype 2 imago, allotype g, morphotype soldier, and paratypes from type 
colony. GHANA: 25m. north of Wenchi on Bamboi Road, 30.iii.1959 (W. A. 
Sands Coll. No. S.2680). In British Museum (Natural History). 


Other paratype material. GHANA: 3m. from Larabanga on Bole Road, 22.1ii. 
1959, and 12m. N. of Techiman on Wenchi Road, 29.iii.1959 (W. A. Sands), soldiers 
and workers. 


152 Ww. A. SANDS 


NiGERIA: Northern Region ; 18m. from Kaduna on Zaria Road, 31.x.1957 
(W. A. Sands), soldiers and workers. All in B.M.(N.H.). 


This species has been recorded from the same vegetation zones as E. aquilinus, 
namely the Guinean savannah woodlands. 


E, CONVERGENS 


E. AQUILINUS 4 


E, ABRUPTUS 


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AF ROSUBULITERMES 
® CONGOENSIS ¢ 


VERRUCOSITERMES 
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PosTSUBULI TERMES 
A PARVICONSTRICTUS 


Vv TARDITERMES CONTRACOLG 


is: MBs = = ae = aN 
m7 } ~~ Neen ( 


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» EDENTATUS 


j 


Maps 30-32. (30), The genus Eutermellus ; (31), “‘ Paracornitermes branch ’’, monotypic 
genera ; (32), The genus Mimeutermes. 


ETHIOPIAN NASUTITERMITINAE 153 


AFROSUBULITERMES Emerson 


Afrosubulitermes Emerson, 1960: 7. Type-species by original designation, 4 frosubulitermes 
congoensis Emerson, 1960. 


Imago. Left mandible with apical tooth distinctly longer than first marginal, but distance 
between them slightly shorter than that from first to third marginals (left mandible index 
-89—-94), cutting edge between the latter deeply indented in front of third marginal and slightly 
or not sinuate, anterior edge of third marginal longer than posterior, gap between this and molar 
prominence very narrow ; right mandible, apical and first marginal as left, second marginal 
small but distinct, molar plate in surface view without basal indentation, asymmetrical, with 
large posterior rounded flange and smaller anterior, weakly ridged. Postclypeus scarcely 
inflated, following line of profile, width 2-3 times length. Fontanelle represented by elongate 
narrow ridge, visible only by side lighting on the highly polished head capsule, with pale oval 
spot in front. Eyes not set out from sides of head. Pilosity virtually absent from head, only 
prominent setae paired, on vertex, between ocelli, in middle of postclypeus and on its anterior 
margin. Pronotum, sides weakly convergent posteriorly, length three-quarters or more of width. 

Soldier. Monomorphic. Nose thin, cylindrical, tapering to fine point. Vestigial mandibles 
without points, labrum with sinuate anterior margin. Head capsule weakly constricted behind 
antennae. Antennae 12 segmented. Head and nose with numerous fine short setae, and six 
longer setae, one pair on vertex and two pairs at base of nose. Abdominal tergites with scattered 
short setae and longer on posterior margins, sternites similar but all setae longer, in both cases, 
marginal setae approximately twice length of rest. 

Worker. Mandibles as imago except apical tooth longer in proportion (left mandible index 
1-361 -39), cutting edge behind left first marginal more sinuate, and right molar plate with more 
pronounced flanges. 


The discovery of the imago of Afrosubulitermes has clarified its relationships to a 
considerable extent. Its closest connection among Ethiopian genera is Eutermellus, 
and the generic separation of the two is not as distinct as would appear from the 
original description of Afrosubulitermes. New material of Eutermellus has shown a 
greater range of variation than was previously known, and the main distinctive 
features of Afrosubulitermes are its extremely small size and much reduced pilosity 
in the imago, and its proportionately shorter nose in the soldier. It also has features 
of resemblance to Postsubulitermes and appears to be intermediate between this 
genus and Eutermellus. Afrosubulitermes was considered by Emerson (1960) to be 
closely related to Verrucositermes, but I believe that the latter was more probably 
separately derived from an ancestral form close to Eutermellus. 


Afrosubulitermes congoensis Emerson 
(Text-figs. 409-411, 428-430, 434-436, 488 ; Map 31) 
A frosubulitermes congoensis Emerson, 1960 : 9. Type locality : Conco, Stanleyville. 


Imago. Previously undescribed. Head capsule sepia brown, paler patches in front of ocelli, 
antennae and postclypeus brown, labrum pale brown. Pronotum and abdominal sclerites 
brown, meso- and metanota, other thoracic sclerites and legs pale brown. Wing membrane 
translucent pale brown, radius sector sepia brown, rest of venation brown, only subcosta and 
radius sector distinct. 

Posterior margin of head capsule behind eyes less than semi-circular, slightly sinuate imme- 
diately behind eyes ; eyes large relative to head width W/E index 2-3-2-4, but not prominent in 
proportion to own diameter, slightly angular oval ; ocelli medium-sized, oval, separated from 


154 WwW. A. SANDS 


440. 442. 


443. 


449. 


448. 450.~" 


Fics. 431-450 : Imago mandibles of “‘ Pavacornitermes branch ”’, showing details of right 
molar plate. 431-433, Eutermellus abruptus ; 434-436, Afrosubulitermes congoensis ; 
437-439, Postsubulitermes parviconstrictus (late nymph); 440-444, Verrucositermes 
tuberosus (440-442, worker, 443, 444, unique morphotype king) ; 445-447, Mimeutermes 
edentatus ; 448-450, Tarditermes contracolor. 


ETHIOPIAN NASUTITERMITINAE 155 


eyes by about one-fifth own least diameter ; anterior margin of postclypeus straight, posterior 
convex, evenly rounded ; antennae 13 segmented, II and V subequal, IV slightly shorter, 
III shorter still and narrower. 

Other characteristics given in generic diagnosis. 

Measurements (10 specimens from 1 locality) in millimetres. 


Range Mean 
Head width across eyes : 0°54-0°56 : : 0°55 
Greatest diameter ofeye . 0+23-0°24 ‘ : 0°24 
Ocellus . i : - 0:06 X 0:08-0:09 0:06 xX 0-08 
Ocellus toeye . : : 0-006—0-013 : ‘ O-OIl 
Width of pronotum . : 0°39-0°43 : : O-41 
Length of pronotum . - 0-29-0°31 : ; 0-31 
Length of hind tibia . 2 0-68-0-71 - : 0-70 
Length of fore wing . ; 4740-460 : - 4°48 


Morphotype 3 imago, NIGERIA : Eastern Region, 12m. from Aba on Ikot Ekpene 
Road, 12.ix.1957 (W. Wilkinson Coll. No. WW 830). In British Museum (Natural 
History). 


Soldier. The additional material has slightly increased the range of variation in size. 
Measurements (14 specimens from 4 localities) in millimetres. 


Range Mean 
Head length to tip of nose. 0:98-1:07 : : : : I‘02 
Head width . , : : 0°37-0'41 : : - ; 0-38 
Depth of head capsule . : 0:28-0:29 : ; : : 0:28 
Width of pronotum : : 0:24-0°32 ; : ; : 0:26 
Length of pronotum : : 0-09 
Length of hind tibia : : 0°47-0°53 : 0°50 


Variation. In the soldier caste the lack of any noticeable variation is significant, since the 
species has been recorded over a range of nearly 1,500 miles. 


The relationships of this species are discussed under the generic heading. 


Material (additional to morphotype colony). NIGERIA : Western Region, Ibadan, 
Agodi, 10.xii.1957 (W. A. Sands), one queen, B.M.(N.H.). 

ConGo : Stanleyville, 26.v.1948, I.vi.1948, paratype soldiers and workers, type 
and other colonies, Leopoldville, 5.vi.1948, paratype soldiers and workers (A. E. 
Emerson) ; Stanleyville, 19.iii.1948, paratype soldier (NV. A. Weber), A.M.N.H. 

REPUBLIC OF CONGO : 13km. W. of Brazzaville, 8.vi.1948, paratype soldiers and 
workers (A. E. Emerson), A.M.N.H. and B.M.(N.H.). 


This species appears widely distributed throughout the Congo and lower Guinean 
forest block, where it has been recorded from galleries within the mounds of other 
species. It has not been found west of the Dahomey-—Togoland gap in the Guinean 
forests. 


POSTSUBULITERMES Emerson 


Postsubulitermes Emerson, 1960: 10. Type-species by original designation, Postsubulitermes 
parviconstrictus Emerson, 1960. 


Imago. ‘The morphotype queen is unique and the mandibles have not been dissected. The 
worker mandibles are described below. Postclypeus moderately inflated, width twice length, 
anterior margin sinuate, convex in middle. Fontanelle elongate, Y-shaped. Eyes not set out 


156 W. A. SANDS 


from sides of head. Pilosity very fine and sparse with very few more prominent setae, three over 
each eye, two pairs on vertex and one pair on postclypeus. Pronotum, sides converging pos- 
teriorly, length less than two-thirds width. 

Soldier. Monomorphic. Nose thin, cylindrical, tapering to fine point. Vestigial mandibles 
without points, labrum with regular rounded anterior margin. Head capsule scarcely con- 
stricted behind antennae, weakly humped at base of nose. Antennae 12 segmented. Head 
setae confined to bilaterally placed pair on vertex, one or two pairs near base of nose, and few at 
nose tip. Abdominal tergites with scattered short setae one-third length of longer setae on 
posterior margins, sternites with scattered short setae longer than on tergites and half length 
of longer setae on posterior margins. 

Worker. Left mandible with apical tooth much longer than first marginal, distance between 
them, nearly three times that from first to third marginals (left mandible index 2-63), cutting 
edge between the latter entire, concave, gap between third marginal and molar prominence 
very narrow ; right mandible, apical and first marginal as left, second marginal absent, molar 
plate with weak basal indentation of outline, weak anterior and strong posterior rounded flanges, 
and no ridges. 


Emerson (1960) considered this genus to have been derived from close to Afro- 
subulitermes and I agree with this, particularly since the recently discovered imago of 
the latter shows close similarities of shape and appearance to Postsubulitermes. The 
imago and worker mandibles of Postsubulitermes are more specialized, and the soldier 
has an evenly rounded labrum, which isa further reduction of the tri-lobed shape seen 
in the more primitive members of the group, such as Eutermellus. The other genera 
are distinguishable by many characters, and are unlikely to be confused with 
Postsubulitermes. 


Postsubulitermes parviconstrictus Emerson 
(Text-figs. 412, 413, 437-439, 464-466, 489 ; Map 31) 
Postsubulitermes parviconstrictus Emerson, 1960 : 13. Type locality : Conco, Yangambi. 


Material. CoNnGo : Yangambi, 0° 47’ N., 24° 23’ E., 20.v.1948 (A. E. Emerson), 
holotype and paratype soldiers and morphotype queen ; Camp Putnam, 1° 24’ N., 
28° 36’ E., 13.v.1948, 4km. N. of Camp Putnam 19 and 20.v.1948 (A. E. Emerson), 
paratype soldiers, A.M.N.H. and B.M.(N.H.). 


VERRUCOSITERMES Emerson 


Verrucositermes Emerson, 1960: 3. Type-species by original designation, Vevrucositermes 
tuberosus Emerson, 1960. 


Imago. The morphotype king of V. tuberosus is unique and the mandibles have not been 
dissected. The worker mandibles in this group are particularly closely similar to the imago and 
are described below. Postclypeus moderately inflated, width 2-5 times length, anterior margin 
concave, sinuate. Fontanelle a pale streak, tapering and slightly depressed posteriorly. Eyes 
set slightly out from sides of head and undercut at rims. Pilosity of head fine and uniform with 
a few more prominent setae. Pronotum, sides converging posteriorly, length less than two- 
thirds width. 

Soldier. Monomorphic. Nose thin, cylindrical, tapering to fine point. Vestigial mandibles 
without points but angular in outline, labrum with sinuate anterior margin. Head capsule 


ETHIOPIAN NASUTITERMITINAE 157 


strongly inflated at base of nose and sharply constricted in middle, with numerous irregular 
small tubercle-like outgrowths of cuticle, denser on base of nose and anterior lobe of head. 
Antennae 12-13 segmented, III to IX or X with 3 tubercles each on inner side. Head and nose 
with numerous scattered small setae. Abdominal tergites with scattered short setae, longer on 
posterior margins, sternites with short setae more dense and longer than tergites, and longer 
on posterior margins. 

Worker. Left mandible with apical tooth longer than first marginal, distance between them 
longer than that from first to third marginals (left mandible index 1-36), notch present in cutting 
edge in front of third marginal, gap between third marginal and molar prominence little wider 
than this notch, subsidiary tooth between them just visible ; right mandible, apical and first 
marginal as left, second marginal almost absorbed in posterior edge of first, molar plate with weak 
basal indentation of outline, strong rounded posterior flange, “‘ ridges ’’ vestigial and embedded 
in smooth shining surface. 

The curious eyes of the imago of Verrucositermes are a feature occurring to a less 
marked degree in some Eutermellus species and in Mimeutermes. The worker 
mandible illustrated by Emerson (1960) appears to be that of Orthotermes depressi- 
frons (Termitinae), a common forest species frequently occurring in the nests of 
other genera. The paratype material deposited at the British Museum (Natural 
History) includes a worker of this species, and there is a notable resemblance in 
general appearance between this and the true paratype Verrucositermes worker in 
the same vial. The Orthotermes worker has a prominent comb of diverging bristle- 
like setae on the anterior coxae, and slight differences in the mandibles. Emerson 
notes in parenthesis that some workers of the type colony have the right second 
marginal tooth less conspicuous than that figured. This is characteristic of the true 
Verrucositermes worker, which also has a much narrower gap between left third 
marginal and molar prominence, and a more elongate right molar plate. 

The very characteristic soldier caste is the chief distinctive feature of the genus, 
though the rugosity of the nose and its base in some Eutermellus and Mimeutermes 
indicates a similar tendency in these genera ; indeed, the vertex and nose-base 
cuticle of Mimeutermes giffardit magnified < 500 shows very numerous outgrowths 
which are closely similar in appearance and may be homologous in spite of their 
much smaller size. 

The phylogenetic position of Verrucositeymes must remain in some doubt, but the 
most likely would appear to be derived from an ancestor of less specialized soldier 
caste, but otherwise similar, intermediate between the genera Eutermellus and 
Mimeutermes. 


Verrucositermes tuberosus Emerson 
(Text-figs. 440-444, 451, 452, 467-469, 490 ; Map 31) 
Verrucositermes tuberosus Emerson, 1960 : 6. Type locality : Conco, Leopoldville. 
Material. Conco : Leopoldville, Kalina Point, 4° 18’ S., 15° 18’ E., 5.vi.1948 
(A. E. Emerson), holotype soldier, paratypes, and morphotype king, A.M.N.H. ; 


Camp Putnam (1° 24’ N., 28° 36’ E.), 22.v.1948 (A. E. Emerson), paratype soldiers, 
A.M.N.H. and B.M.(N.H.). 


158 W. A. SANDS 


MIMEUTERMES Silvestri 


Mimeutermes Silvestri, 1914 : 47. Type-species by original designation, Mimeutermes giffardii 

Silvestri, 1914. 

Imago. Left mandible with long apical tooth, cutting edge from first to third marginals 
short, slightly concave (left mandible index 2-00—2-23), gap between third marginal and molar 
plate less than half as wide as this cutting edge ; right mandible, apical and first marginal as 
left, second marginal obsolete, posterior cutting edge of first marginal concave, molar plate with 
pronounced anterior and posterior rounded flanges, and tapering distally, ridges absent. Post- 
clypeus weakly inflated, width twice length or slightly less. Fontanelle pale, slit-like, slightly 
bifurcate with pale patch in front, to almost obsolete, little paler than head. Eyes slightly set 
out from sides of head. Pilosity of head variable. Pronotum, rounded posteriorly, anterior 
margin concave. 

Soldier. Monomorphic. Nose thickly conical, merging with taper of head capsule at base, 
fontanelle rather large. Vestigial mandibles with and without points, labrum with sinuate 
anterior margin. Head capsule in plan view not constricted, top-shaped including nose, with 
sides parallel or rounded towards back. Antennae 12 segmented. Head and nose with few 
conspicuous setae. Abdominal tergites with sparse scattered minute setae, longer on posterior 
margins and posterior 2—3 tergites ; sternites with sparse short setae and longer on posterior 
margins, approximately twice length of short. 

Worker. Mandibles as imago except apical tooth longer in proportion (left mandible index 
3°32-3°59), and right molar plate larger with more pronounced flanges. 


The genus Mimeutermes, like the other Ethiopian members of the “ Para- 
cornitermes-branch ’’, is very distinctive in form, and detailed discussion is not 
required. Its relationships to Tarditermes and Verrucositermes are discussed under 
those genera. It is unlikely to be confused with any other genus except in the 
worker caste. 

This is an unusual genus in that it appears to have evolved in the rain forest to 
the west of the Dahomey—Togoland gap. This forest limits the distribution of 
M. giffardiu, the most primitive species, but the other two appear progressively 
more adapted to savannah conditions, and have spread back to Nigeria, though un- 
known further eastwards. It seems probable that this reflects the genuine distri- 
bution, rather than the failure of collectors, in view of the frequent records of related 
genera from similar habitats in other localities. 


KEYS TO SPECIES 
IMAGO 


1 Larger, W, 1:08-1:12 ; pronotum very large, wider than head width across eyes, 
1:25-1:35 : ; ‘ : : : ; : : edentatus (p. 160) 
— Smaller, W, 0-89-0-97 ; pronotum narrower than head width across eyes, 0-78—0-89 2 
2 Eyes smaller, 0-29 ; T, longer, 1-02. Pilosity of head capsule and pronotum very 
even, forming a “ pelt’ with a few scattered longer setae (Text-fig. 463) 
sorex (p. 163) 
— Eyes larger, 0:31-0:33 ; T, shorter, 0-81-0-87. Pilosity of head capsule and prono- 
tum uneven, not forming a “ pelt ’’, with many scattered longer setae, (Text-fig. 
462) : . giffardii (p. 161) 


ETHIOPIAN NASUTITERMITINAE 159 


460 


Fics. 451-460 : Imago head capsule, front and side views. 461-463: Vertex pilosity. 
451, 452, Verrucositermes tuberosus ; 453, 454, 461, Mimeutermes edentatus ; 455, 456, 
462, M. giffardii ; 457, 458, 463, M. sovex ; 459, 460, Tarditermes contracolor. 


160 WwW. A. SANDS 


SOLDIERS 
— Vestigial mandibles with points . : : - - : : . giffardii (p. 161) 
— Vestigial mandibles without points . : 2 
2 Head capsule in plan view top-shaped, sides ene more or ie Pee towande 
front . : : sorex (p. 163) 
— Head capsule in ae view aes ican shaped, sides ra concave ape taper towards 
front . ‘ : : : : : ; : ‘ : edentatus (p. 160) 


Mimeutermes edentatus Sands 
(Text-figs. 445-447, 453, 454, 461, 470-473, 491 ; Map 32) 


Mimeutermes edentatus Sands, 1956 : 83. Type locality : Guana, Accra. 


Imago. Previously undescribed. Head capsule dark sepia brown, postclypeus sepia brown, 
antennae and labrum yellow-brown. Pronotum, meso- and metanota sepia brown, pronotum 
with distinct large pale T-shaped mark in middle of anterior half, other thoracic sclerites brown, 
legs yellow-brown. Abdominal tergites sepia-brown, sternites brown, paler in mid-line. Wing 
membrane translucent pale brown, venation sepia brown, distinct. 

Posterior margin of head capsule behind eyes widely rounded, slightly curved outwards at 
eye rim ; fontanelle a pale tapering slit ; eyes very slightly set out from sides of head, angularly 
short oval, moderately prominent in proportion to own diameter, and small to medium-sized 
relative to head width, W/E index 3-1-3-6 ; ocelli medium-sized, separated from eyes by half 
to four-fifths own least diameter ; anterior margin of postclypeus sinuate, convex in middle, 
posterior convex, rounded, width 1-8—1-9 times length ; antennae 15 segmented, IV and V 
nearly equal, II shorter and narrower, III much smaller. Pronotum very large, shield-like, 
wider than head across eyes. 

Pilosity of vertex brown, very dense, short and even, forming a “ pelt ’’ or “‘ mat’’, with a 
few longer emergent setae more or less symmetrically placed in pairs or groups, sometimes one 
very prominent seta between eye, ocellus and antennal socket on each side ; pronotum similar. 
Abdominal pilosity, tergites as pronotum, with few longer setae on posterior margins of last 6, 
more scattered and numerous on last 3 ; sternites with more longer setae, especially on posterior 
margins and last 4-5 segments. 

Measurements (10 specimens from 1 locality) in millimetres. 


“ 


Range Mean 
Head width across eyes I-08=1 -12 : 3 I-10 
Greatest diameter of eye 0°30-0°35 : c 0°33 
Ocellus : : 6 O'I0-O'I2 X O'13-O0'1I5 . ; O-II X O-14 
Ocellus toeye . : 0:06-0:08 , : 0:07 
Width of pronotum. I+25-1°35 : ‘ TZ 
Length of pronotum . I-O4—I°15 : ; I-10 
Length of hind tibia. I ‘00-1 -06 ¢ : I-04 
Length of fore wing . 10-60-11 -30 : ; 10°87 


Morphotype 2 imago. NIGERIA: Northern Region, 25m. from Jos on Kaduna 
Road, 5.i11.1957 (W. A. Sands Coll. No. S.1256). In British Museum (Natural 
History). 


Soldiey. The additional material has increased the known range of variation in size. 
Measurements (15 specimens from 3 localities) in millimetres. 


ETHIOPIAN NASUTITERMITINAE 161 


Range Mean 
Head length to tip of nose. I-22-1-61 : C . : 1°43 
Head width . : - : 0-82-1-02 : . - : 0-94 
Depth of head capsule . : 0°49-0°64 : - ‘ ; 0°58 
Width of pronotum . : 0°43-0°51 : 2 - : 0°47 
Length of pronotum : : 0-18—0-25 : : - : 0-21 
Length of hind tibia : . 0°59-0°77 : . F 0-67 


M. edentatus was described from a unique soldier, and additional material agrees 
well with the original description. The characteristics separating it from the other 
two species are discussed thereunder. 


Material (other than holotype, Sands, 1956). GHANA: 30m. from Navrongo on 
Tumu Road, 13.11.1959 (W. A. Sands). 


M. edentatus, like its congeners, has been recorded from the mounds of other 
genera, but appears, unlike them, to be adapted to life in shaded habitats in the 
savannah zones, and has not been found in rain forest. 


Mimeutermes giffardii Silvestri 
(Text-figs. 455, 456, 462, 476-478, 493, 494 ; Map 32) 


Mimeutermes giffardii Silvestri, 1914 : 48. Type locality : Guinea, Camayenne. 


Imago. Head capsule very dark sepia brown, antennae and postclypeus sepia brown, labrum 
yellow. Pronotum, meso- and metanota sepia brown, other thoracic sclerites brown, femora 
yellow, tibiae and tarsi yellow-brown. Abdominal tergites brown, sternites brown laterally, 
yellow-white in mid-line. Wing membrane translucent pale brown, venation sepia brown, all 
distinct. 

Posterior margin of head capsule behind eyes widely rounded, sharply curved outwards at 
eye rim ; fontanelle scarcely paler than head capsule, sometimes slightly bifurcate in front with 
paler spot ; eyes slightly but distinctly set out from sides of head, angularly short oval, promin- 
ent in proportion to own diameter and medium-sized relative to head width, W/E index 
2:9-3:0 ; ocelli small, separated from eyes by half to three-quarters own least diameter ; 
anterior margin of postclypeus sinuate, convex in middle, posterior convex, rounded, width 2:2 
times length ; antennae 15 segmented, IV and V nearly equal, II shorter and narrower, III 
much smaller. Pronotum narrower than head width across eyes. 

Pilosity of vertex and pronotum yellow-brown, uneven in length with many scattered larger 
setae, not forming a “‘pelt’’ or “‘mat’’. Abdominal pilosity longer and denser on posterior 
segments, sternites more so than tergites. 

Measurements (12 specimens from 3 localities) in millimetres. 


Range Mean 
Head width across eyes 0:89-0:97 : ‘ 0:94 
Greatest diameter of eye 0°31-0°33 : : 0°31 
Ocellus 3 : : 0:08-0:09 X O:09-O'II . : 0:09 X O-10 
Ocellus toeye  . : 0:04-0:07 2 : 0-06 
Width of pronotum . 0:78-0°89 : : 0°85 
Length of pronotum . 0-61-0:71 : : 0-66 
Length of hind tibia. 0:82-0:87 : : 0-84 
Length of fore wing . 8:20-8:60 8-38 


Soldier. Head capsule yellow to ferruginous orange, nose orange to ferruginous orange, rest 
of body cuticle yellow. 


162 W. A. SANDS 


464. A466. 
QD 
(> 
474 
470. 472. 
BS 
473. A75. 476. 478. 


Fics. 464-478 : Soldier head capsule, side and plan views, labrum, and mandibles. 464- 
466, Postsubulitermes parviconstrictus ; 467-469, Verrucositermes tuberosus ; 470-472, 
Mimeutermes edentatus ; 473-475, M. sovex ; 476-478, M. giffardiit. 


ETHIOPIAN NASUTITERMITINAE 163 


Head capsule in plan view top-shaped, sides parallel at back then sharply converging con- 
tinuously with nose to about half nose length, rest of nose tapering less sharply. In profile 
strongly humped above antennal socket, which is distinctly behind middle of head, nose slightly 
upturned ; vestigial mandibles with points ; antennae, proportions of basal segments some- 
what variable, usually II, [V and V subequal, longer up to twice III. Conspicuous head setae 
few, one bilaterally placed pair near back of head, and one pair near base of nose. Cuticle of 
vertex distinctly rastrate, at high magnifications ( x 500) small papillae visible. 

Measurements (14 specimens from 4 localities) in millimetres. 


Range Mean 
Head length to tip of nose. 1°25-1°48 : 3 , ‘ 1°39 
Head width . : : ‘ 0:72-0°87 : 2 : : 0°83 
Depth of head capsule . : 0-61-0°71 : : : : 0-66 
Width of pronotum : ; 0°43-0°46 : , : ; 0°44 
Length of pronotum : : 0-18—0-26 : : ‘ : 0:23 
Length of hind tibia ; : 0°54-0°64 : : : ‘ 0°59 


M. giffardii is easily distinguished from the other two species by the points on 
the soldier mandibles. Other features are the distinctly rastrate vertex and the 
differing profile in the soldier. In the imago, the coarser irregular vertex pilosity is 
distinctive ; in addition the larger eyes and shorter hind tibia separate it from 
M. sorex and the generally smaller size from M. edentatus. M. giffardii is probably 
the most primitive of the three species in possessing these characteristics. 


Material. GUINEA : Camayenne, 9° 25’ N., 13° 40’ W., 1912-13 (F. Silvestri), 
holotype 2 imago, paratype soldiers and workers, M. giffardii Silvestri, Silvestri 
Coll., Lab. Zool. Sc. Agr., Portici. 

GHANA : Aburi, 1912-13 (F. Silvestri), further paratype imago, soldiers, and 
workers, Silvestri Coll., as above ; Kumasi, Kwadaso, 19.ii.1959, and 53m. from 
Dunkwa on Obuasi Road, 4.iv.1959 (W. A. Sands), B.M.(N.H.). 


This species appears to be one of the very few which are confined to the rain 
forest areas to the west of the Dahomey—Togoland gap, presumably having evolved 
in isolation from the main Congo forest block. It has been recorded from the 
mounds of other genera of termites. 


Mimeutermes sorex Silvestri 
(Text-figs. 457, 458, 463, 473-475, 495 ; Map 32) 


Mimeutermes sovex Silvestri, 1914 : 50. Type locality : GuINEA, Mamou. 


Imago. Previously undescribed. Head capsule dark sepia brown, antennae and postclypeus 
brown, labrum yellow. Pronotum, meso- and metanota brown, other thoracic sclerites and 
legs yellow-brown. Anterior abdominal tergites yellow-brown, posterior brown ;_ sternites 
yellow-brown, brown around stigmata. Wings absent. 

Posterior margin of head capsule behind eyes widely rounded, sharply curved outwards at 
eye rim ; fontanelle a minute, pale, tapering slit with oval pale spot in front ; eyes slightly 
set out from sides of head, angularly short oval, prominent in proportion to own diameter and 
medium-sized relative to head width, W/E index 3-4 ; ocelli small, separated from eyes by 
approximately three-quarters own least diameter ; anterior margin of postclypeus sinuate, 
convex in middle, posterior convex, rounded, width 2-0 times length ; antennae 15 segmented, 


164 W. A. SANDS 


IV slightly larger than V, II smaller than these, III much smaller. Pronotum narrower than 
head width across eyes. 

Pilosity of vertex and pronotum brown, of medium length, very dense and even, forming a 
“pelt ”’ or “mat ”’, with a few longer emergent setae more or less symmetrically placed in pairs 
or groups. Abdominal pilosity, first 3-4 tergites as pronotum, posterior tergites and sternites 
with longer setae of uneven length. 

Measurements (Unique morphotype 9) in millimetres. 


Head width across eyes z : : : : 0:97 
Greatest diameter ofeye . : : : ; 0:29 
Ocellus. : : . : ? : 3 0:09 X O-IO 
Ocellus toeye . , ; - : : 4 0-06 
Width of pronotum . : . : : : 0-89 
Length of pronotum . ; : 3 : ; 0-66 
Length of hind tibia . : : : 2 ; 1:02 


Unique morphotype 2 imago. NIGERIA: Eastern Region: Port Harcourt, 
1.1V.1957 (W. Wilkinson, Coll. No. WW.607). In British Museum (Natural History). 


Soldiey. Head capsule yellow to ferruginous, nose orange to chestnut brown, rest of body 
cuticle pale yellow to yellow. 

Head capsule in plan view top-shaped, posterior corners rounded, sides sharply converging 
to near tip of nose in continuous line. In profile strongly humped above antennal socket, 
which is behind middle of head, nose more or less parallel to long axis of head ; vestigial man- 
dibles without points ; antennae, proportions of basal segments somewhat variable, usually II 
and V subequal, IV subequal or slightly larger, III smaller and narrower than these. Con- 
spicuous head setae few, usually one bilaterally placed pair towards back of head. Cuticle of 
vertex not markedly rastrate. 

Measurements (25 specimens from 8 localities) in millimetres. 


Range Mean 
Head length to tip of nose. 1°35-1°51 : 3 : : 1°43 
Head width . : : 3 0:75-0:89 : ; 3 , 0:82 
Depth of head capsule . : 0:61-0:69 : : : : 0°63 
Width of pronotum : : 0-41-0-46 : : : : 0°44 
Length of pronotum ; ‘ 0-18-0-22 - : : : 0-19 
Length of hind tibia : 0-64—0°71 0-66 


M. sorex differs from M. necnaee in the shape i dae! sicher ecia capsule, par- 
ticularly in profile, and the lack of points on the mandibles. M. edentatus is also 
different in shape. In the imago, the smaller eyes and very even dense head and 
pronotum pilosity separate M. sorex from M. giffardii. M. edentatus is larger, with 
a wider pronotum, and shorter but still dense and even pilosity. 

The description above differs from that given by Silvestri in several respects, of 
which the most important is the lack of points on the soldier mandibles. The reason 
for this is that the syntypes of M. sovex include specimens of M. giffardi1, presumably 
having been collected from a mound containing both species. This fact was not 
recognized by Silvestri, who described the general shape of M. sorex, clearly dis- 
tinguishing it from the paratype soldiers of M. giffardii. The soldier mandibles he 
must have illustrated and described from a specimen of M. giffardii in which the 
points were slightly smaller than usual. The true WM. sovex syntypes are without 
points, and no other specimen with points has been found. When the syntypes 
were examined in Portici in 1958, a lectotype was not selected, and this designation 
must await a further personal visit since the Silvestri collection is not accessible by 


ETHIOPIAN NASUTITERMITINAE 165 


484. 485 
i) 6 f pene 
U 4 id 
491. 492 493 494. 


497. 500 


Fics. 479-496 : Imago fontanelle. 497-500: Soldier head capsule, nose tip, labrum and 
mandibles. 479, 3, 480, 481, 2, Eutermellus abruptus ; 482, 3, 483, 92, E. aquilinus ; 
484, 5, E. bipartitus ; 485, 5, E. convergens ; 486, 3, 487, 2, E. undulans ; 488, 3, 2, 
A frosubulitermes congoensis ; 489, @, Postsubulitermes parviconstrictus ; 490, 4, 
Verrucositermes tuberosus ; 491, 3, 2, Mimeutermes edentatus ; 492, 3, 493, 494, 2, 
M. giffardii ; 495, 2, M. sorex ; 496-500, Tarditermes contracolor. 


166 W. A. SANDS 


post. However, there is no doubt as to the true identity of M. sovex and its genuine 
distinctness from M. giffardit. 


Material. GUINEA: Mamou, 10° 20’ N., 12° 15’ W., 1912-13 (F. Silvestri), 
syntype soldiers and workers, M. sorex Silvestri, Silvestri Coll., Lab. Zool. Sc. Agr., 
Portici, and A.M.N.H. 

GHANA: 17m. N.E. of Gambaga on Bawku Road, 6.ili.1959, and 12m. N. of 
Techiman on Wenchi Road, 29.111.1959 (W. A. Sands). 

NIGERIA : Western Region ; Ikeja, 10.11.1955 (W. V. Harris) ; Benin Prov., 
Sobo Plain, Obanokoro, 7.111.1957 (W. Wilkinson) ; between Shagamu and Jjebu- 
Ode, 14.xii.1957 (W. A. Sands). Eastern Region ; Port Harcourt, 18.iv. and 
8.xi.1957, and Onitsha, 6.x1.1957 (W. Wilkinson). 

Material in British Museum (Natural History) unless otherwise stated. 


M. sorex, like its congeners, has been recorded from the mounds of other genera. 
It is more widely distributed than M. giffardiu, having apparently become more 
tolerant of the marginal conditions of riverain and relict forest patches in the 
savannah zones, and has crossed the Dahomey-—Togoland gap to the Nigerian rain 
forest. It has not however been recorded from the Congo forest block proper. 


TARDITERMES Emerson 


Tarditermes Emerson, 1960: 15. Type-species by original designation, Tarditermes contra- 
coloy Emerson, 1960. 


Imago. Left mandible with apical tooth very long, distance from apical to first marginal 
over 3 times that from first to third marginals (left mandible index 3-17), cutting edge between 
the latter entire, slightly concave, gap between third marginal and molar prominence about 
half as wide as this cutting edge, small subsidiary tooth visible in gap in surface view ; right 
mandible, apical and first marginal as left, second marginal absent, molar plate with no marked 
irregularity of outline in surface view, and no ridges. Postclypeus, width 2-1 times length, 
moderately inflated, anterior margin convex, sinuate. Fontanelle a light round spot surrounded 
by an indefinite pale patch with lateral arms. Eyes very slightly set out from sides of head. 
Pilosity of head short and even with scattered longer setae. Pronotum almost semi-circular 
with rounded corners. 

Soldiey. Monomorphic. Nose long, thick, conical, tapering to large fontanelle. Vestigial 
mandibles without points, labrum with sinuate anterior margin. Head capsule not constricted, 
rounded rectangular in plan and wider than long to base of nose. Antennae 13 segmented. 
Head setae confined to nose tip apart from scattered microscopic hair-like setae on head and 
nose. Abdominal tergites with minute setae, longer towards posterior ; sternites with short 
setae and longer on posterior margins. 

Worker. Mandibles as imago, except right molar plate weakly indented near base and with 
more pronounced anterior and posterior flanges (left mandible index 3:23). 


According to Emerson (1960), the imago of Tarditermes is larger than all its nasute 
relatives except the neotropical Angularitermes. This is true but the recently 
discovered imago of Mimeutermes edentatus Sands to a large extent bridges the gap. 
Other features of the latter species appear to indicate a relationship with Tardt- 
termes, including the characteristic slow movement of the neuter castes in life. The 
imago of Tarditermes most closely resembles Mimeutermes in the widely curved 


ETHIOPIAN NASUTITERMITINAE 167 


posterior margin of the head capsule, the prominent eyes, the shape of the postclyp- 
eus, the rounded pronotum, and the imago mandibles. The soldier of Tarditermes 
is closely similar to Mimeutermes except for the longer nose. The distinguishing 
features of the imago are its striking colour pattern, reduced pilosity, and larger 
size compared with Mimeutermes. Tarditermes thus appears to be a more specialized 
genus, probably derived from an ancestor very close to Mimeutermes. 


Tarditermes contracolor Emerson 
(Text-figs. 448-450, 459, 460, 496-500 ; Map 31) 


Tarditermes contracoloy Emerson, 1960: 18. Type locality : Conco, near Camp Putnam, 
Epulu River. 
Material, Conco: 4km. N. of Camp Putnam, 1° 24’ N., 28° 36’ E., 19.v.1948, 
and Camp Putnam, 17.v.1948 (A. E. Emerson), holotype and paratype imagos, 
paratype soldiers and workers, A.M.N.H. and B.M.(N.H.). 


ACKNOWLEDGEMENTS 

I owe sincere gratitude to a number of persons and institutions from whom I have 
received generous assistance in carrying out this revision. 

Firstly I wish to thank Dr. A. E. Emerson, lately Professor at the University of 
Chicago, for all he did during my visit to assist and encourage my studies, and 
particularly for free access to his unique collections and card indices. Secondly, I 
wish to thank Dr. W. G. H. Coaton, of the Plant Protection Research Institute, 
Pretoria, for great generosity, as on previous occasions, in providing material of 
South African species, including one new to science and named in his honour. 

I am also grateful to the following specialists and curators who have been kind 
enough to send me type specimens or other material on loan from their respective 
museums : Prof. Dr. H. Weidner, of Hamburg, Prof. Dr. L. Brundin and Dr. K. J. 
Hedgrist, of Stockholm, Dr. E. M. Kruytzer, of Maastricht, Dr. F. Capra and 
Dr. Delfa Guiglia, of Genoa, Dr. P. Basilewsky, of Tervuren, M. A. Descarpentries, of 
Paris, and M. le Directeur, Institute Royal de Sciences Naturelles, Brussels. 

My thanks are also due to Dr. W. V. Harris, Officer-in-Charge, Termite Research 
Unit, for help and advice throughout the progress of this revision. 

I further wish to express my gratitude to the Trustees of the Leverhulme Research 
Awards Foundation, for the generous grant which made possible a study visit to the 
University of Chicago in 1962. Finally, my thanks are due to the Trustees of the 
British Museum (Natural History) for permission to examine types and other 
material in the termite collection. : 


168 MW A SANDS 


REBERENCES 


AuMAD, M. 1950. The phylogeny of termite genera based on the imago-worker mandibles. 
Bull. Amer. Mus. nat. Hist. 95 : 37-86, 17 text-figs. 

Banks, N. 1919. Antillean Isoptera. Bull. Mus. comp. Zool. Harv. 62 : 475-489, 2 pls. 

Banks, N. & SNYDER, T. E. 1920. A revision of the Nearctic termites with notes on biology 
and geographic distribution. Bull. U.S. nat. Mus. 108 : 1-228, 70 text-figs., 35 pls. 

Capra, F. 1938. Sulla presenza in Libia di Tvinervitermes tripolitanus (Sjdstedt) e trinervius 
(Rambur) (Isoptera). Boll. Soc. ent. ital. 70 : 125. 

Coaton, W. G. H. 1962. Survey of the termites of the Kruger National Park (including 
diagnosis of Fulleriteymes gen. nov. : Nasutitermitinae). Koedoe (Sci. J. nat. Pks. Bd. S. 
Afr.) 5 : 144-156, 11 text-figs. 

DESNEUX, J. 1904. Notes termitologiques. Amn. Soc. ent. Belg. 48 : 146-151. 

1904a. Isoptera, family Termitidae. Geneva Insect. 25 : 52, 2 pls. 

1905. Isoptera of New Guinea collected by L. Bird. Ann. hist.-nat. Mus. hung. 3: 

367-377, 5 text-figs. 

—— 1915. Isoptera, in: Collections zoologiques du Baron Edm. de Selys Longchamps 3 : 1-10, 
Brussels. 

DuplLey, P. H. & Beaumont, J. 1889. Observations on the termites or white ants of the 

Isthmus of Panama. Tvans. N.Y. Acad. Sci. 8 : 85-114, 9 text-figs., 1 pl. 

1890. Termites of the Isthmus of Panama, part II. Tvans. N.Y. Acad. Sci. 9 : 157-180. 

Emerson, A. E. 1925. The termites of Kartabo, Bartica District, British Guiana. Zoolo- 
gica, N.Y. 6 : 291-459, 94 text-figs., 1 pl. 

1928. Termites of the Belgian Congo and the Cameroon. Bull. Amer. Mus. nat. Hist. 

57 : 401-574, 79 text-figs., 19 pls. 

1960. New genera on the Subulitermes branch of the Nasutitermitinae from the Ethiopian 

Region (Isoptera, Termitidae). Amer. Mus. Novit. 1987 : 1-21, 11 text-figs. 

Fasricius, J.C. 1793. Entomologia Systematica 2. viii + 519 pp. Copenhagen. 

FULLER, C. 1921. The termites of South Africa ; being a preliminary notice. S. Afr. J. nat. 

Hist. 3 : 14-52. 

1922. The termites of South Africa. S. Afr. J. nat. Hist. 3 : 70-131. 

GHIDINI, G. M. 1941. Nuove specie di termitidi. Boll. Soc. ent. ital. 73 : 30-34. 

GRIFFIN, F. 1942. Henry Smeathman (?-1789). Proc. R. ent. Soc. Lond. (A). 17 : 1-19. 

Hacen, H. A. 1858. Monographie der Termiten, part 2. Linn. ent. 12 : 1-342, 3 pls. 

HALDEMAN, S. S. 1853. Descriptions of some new species of insects, with observations on 
described species. Proc. Acad. nat. Sci. Philad. 6 : 361-365. 

Hare, L. 1937. Termite phylogeny as evidenced by soldier mandible development. Ann. 
ent. Soc. Amer. 30 : 459-473, 13 pls. 

Harris, W. V. 1954. Further records of East African termites—II. Proc. R. ent. Soc. 

Lond. (B). 23 : 127-137, 6 text-figs. 

1957. Isopterain: B.M.N.H. Rept. Exp. S.W. Arabia : 421-433, 5 text-figs. 

Hecu, E. 1922. Les Termites. 756 pp., 460 text-figs., 1 map. Brussels. 

HotmGREN, N. 1910. Versuch einer Monographie der amerikanischen Eutermes-Arten. 

Mitt. naturh. Mus. Hamb. 27 : 171-325, 78 text-figs. 

1912. Termitenstudien 3. Systematik der Termiten. Die Familie Metatermitinae. 

Kk. svenska VetenskAkad. Handl. 48 : 1-166, 4 pls. 

1913. Termiten aus Natal und dem Zululande. Gesammelt von Dr. Ivar Tragardh. 

Ent. Tidskry. 34 : 321-366. 

Kray, R. W. J. etal. 1959. Vegetation map of Africa South of the Tropic of Cancer. Explana- 
tory notes. 24 pp.,1 map. Oxford. 

LaTREILLE, P. A. 1805. Histoive naturelle, génévale et particuliére des Crustacés et des Insectes 
13. 432 pp. Paris. 

Licut, S. F. 1930. Notes on Philippine termites IV. Philipp. J. Sci. 42 : 13-58, 1 text-fig., 
8 pls. 


ETHIOPIAN NASUTITERMITINAE 169 


Mayr, E., Linstey, E. G. & Usincer, R. L. 1953. Methods and principles of systematic 
zoology. ix + 328 pp. New York. 

MotscHULsky, V. DE. 1855. Etudes Entomologiques 4 : 10. 

Norrort, C. 1955. Termites du Centre et du Sud-Ouest de |’Angola récoltés par A. de Barros 
Machado. Publ. Cult. Comp. Diam. Angola 27 : 141-150, 8 text-figs. 

PRASHAD, B. & SEN-SARMA, P. K. 1959. Revision of the termite genus Nasutitermes, Isoptera, 
Termitidae : Nasutitermitinae from the Indian Region. 66 pp., 25 text-figs. Delhi. 

RamBur, J. P. 1842. Histoire naturelle des Insectes. Névroptéves. 534 pp., 12 pls. Paris. 

Rosen, K. von. 1912. Neue Termiten aus der zoologischen Staatssammlung in Miinchen 
sowie einigen anderen Sammlungen. Zool. Anz. 39 : 221-232, 9 text-figs. 

Sanps, W. A. 1956. <A new species of Mimeutermes from the Gold Coast (Isoptera : Nasuti- 

termitinae). Pvoc. R. ent Soc. Lond. (B). 25 : 83-84, 1 text-fig. 

1957. The soldier mandibles of the Nasutitermitinae (Isoptera, Termitidae). Jmsectes soc. 

4 : 13-24, 5 text-figs. 

— 1957a. A revision of the East African Nasutitermitinae (Isoptera). Bull. Brit. Mus. 

(Nat. Hist.) Ent. 5 : 1-28, 6 text-figs. 

1961. Nest structure and size distribution in the genus Tvinervitermes (Isoptera, Termi- 

tidae, Nasutitermitinae) in West Africa. Insectes soc. 8 : 177-188, 2 text-figs., 2 pls. 

— 1961a. Foraging behaviour and feeding habits in five species of Tvinervitermes in West 
Africa. Entomologia exp. appl. 4 : 277-288, 2 text-figs. 

SILVEsTRI, F. 1903. Contribuzione alla conoscenza dei Termiti e Termitofili dell’America 

meridionale. Media 1 : 1-234, 6 pls. 

1912. Termiti raccolte da L. Fea alla Guinea Portoghese e alla isole S. Thomé, Annobon, 

Principe e Fernando Poo. Ann. Mus. civ. Stor. nat. Genova (3). 5 : 211-255, 20 text-figs. 

1914. Contribuzione alla conoscenza die Termitidi e Termitofili dell’ Africa occidentale. 

I, Termitidi. Boll. Lab. Zool. Portici 9 : 1-146, 84 text-figs., 1 pl. 

SJOSTEDT, Y. 1896. Termiten aus Kamerun. Ent. Tidskr. 17 : 297-298. 

1897. Neue Termiten aus West-Afrika (Vorlaufige Mitteilung). Ent. Tidsky. 18 : 123-126 

— 1899. Neue afrikanische Termiten (Vorlaufige Mitteilung). Ent. Nachr. 25 : 34-39. 

1900. Monographie der Termiten Afrikas. K. svenska VetenskAkad. Handl. 34 : 1-236, 

9 pls. 

1902. Termites novos ex Africa reportatos. Ent. Tidsky. 23 : 40. 

— 1go2a. Neue afrikanische Termiten. Ent. Tidskr. 23 : 302-304. 

1904. Monographie der Termiten Afrikas. K. svenska VetenskAkad. Handl. 38 : 1-120, 


4 pls. 
1905. Ueber eine Termitensammlung aus Kongo und anderen Teilen von Afrika. Ark. 
Zool. 2 : 1-20. 


1907. Ueber eine Termitensammlung aus Kongo. Ent. Tidskr. 28 : 233-250. 

1g07a. Termitidae in: Wiss. Evgebn. schwed. zool. Exped. Kilimandjaro-Meru, 1905-6. 
3: 1-24, 4 pls. Uppsala. 
1o1r. Zur Termitenfauna Kongos. Ent. Tidskr. 32 : 137-170, 8 text-figs. 
1g1ta. Neue Ost- und Westafrikanische Termiten. Ent. Tidsky. 32 : 173-188. 
to11b. Neue afrikanischen Termiten im Schwedischen Reichsmuseum. Ark. Zool. 
7: 1-18. 

1912. Neue Termiten aus Tripolis, Ober-Aegypten, Abessinien, Erithrea, dem Galla- und 
Somalilande. Ark. Zool. 7 : 1-17. 

1913. Ergebnisse einer botanischen Forschungsreise von J. Brunthaler nach Deutsch- 
Ostafrika und Siidafrika (Kapland, Natal und Rhodesien). Termiten. Denkschy. Akad. 
Wiss. Wien 88 : 720-721. 
1913a. Uber Termiten aus dem inneren Congo, Rhodesia, und Deutsch-Ostafrika. ev. 
Zool. Bot. afv. 2 : 354-391. 

1914. Termiten aus Zambezi, Rhodesia, Nyassa, und Siid-Nigeria. Avk. Zool. 8 : 1-9, 
3 text-figs. 


170 W. A. SANDS 


SJOSTEDT, Y. 19g14a. Isoptera in: W. Michaelsen. Betty. Land-u. Siiswasserfauna 
Dtsch-SiidwAfr. 1 : 73-92, 2 text-figs., 1 pl. 

—1914b. Termiten. Wiss. Evgebn. dtsch. zent.-Afr. Exped. 1 : 89-95, 4 pls. 

—— 1924. Neue Termiten im Kongo-Museum. fev. Zool. Bot. afr. 12 : 39-42. 

——  1924a. Neue Arten und Gattungen afrikanischer Termiten. ibid. 12 : 253-257. 

— 1924b. Neue afrikanische Termiten. ibid. 12 : 490-494, 495-497. 

—— 1925. Neue Termiten aus Afrika und Madagascar. Konowta 4 : 53-55. 

—  1925a. Wissenschaftliche Ergebnisse der mit Unterstiitzung der Akademie der Wissen- 

schaften in Wien aus der Erbschaft Treitl von F. Werner unternommenen Zoologischen 

Expedition nach dem Anglo-Agyptischen Sudan (Kordofan) 1914. XXI, Isoptera. 

Denkschr. Akad. Wiss. Wien. 100 : 71-76, 1 pl. 

1926. Revision der Termiten Afrikas 3. Monographie. K. svenska VetenskAkad. Handl. 

(3). 3: 1-410, 83 text-figs., 16 pls. 

1926a. Neue Termiten aus der Goldkiiste. Avk. Zool. 18 : 1-5. 

1926b. Kongo-Termiten aus dem Ituri und Uelegebiet. Rev. Zool. Bot. afr. 14 : 140-164, 

3 text-figs. 

SMEATHMAN, H. 1781. Some account of the termites which are found in Africa and other hot 
climates. Phil. Trans. 71 : 139-192, 4 pls. 

SNYDER, T. E. 1949. Catalog of the Termites (Isoptera) of the World. Smithson. misc. Coll. 
112 : 1-490. 

TRAGARDH, I. 1904. Termiten aus dem Sudan. Results Swed. Zool. Exped. Egypt and White 
Nile, 1901 12 : 1-47. 

WALKER, F. 1863. List of the specimens of Neuropterous insects in the British Museum. (Part 
3, Termitides). 658 pp. London. 

WasMaAnn, E. 1897. Termiten von Madagascar und Ostafrika (Wiss. Ergebn. der Reisen in 

Madagascar und Ostafr. 1889-95 von Dr. A. Voeltzkow). <Abh. senckenb. naturf. Ges. 

21 : 137-182, 2 pls. 

1902. Einige Bemerkungen zu Sjéstedt’s 

Zbl. 22 : 714-715. 

1911. Zur Kenntnis der Termiten und Termitengdste von Belgischen Kongo. Rev. Zool. 

Bot. afr. 1 : 91-117 and 145-176, 8 pls. 

WEIDNER, H. 1955. Die Typen der Isoptera-Sammlung des Zoologischen Staatsinstituts und 

Zoologischen Museums, Hamburg. Insectes soc., 2 : 247-254. 

1956. Beitrage zur Kenntnis der Termiten Angolas, hauptsachlich auf Grund der Samm- 

lungen und Beobachtungen von A. de Barros Machado. Publ. Cult. Comp. Diam. Angola 

29 : 55-106, 45 text-figs. 


« 


Monographie der Termiten Afrikas’’. Biol. 


171 


INDEX TO SPECIES AND SUBSPECIES 
(synonyms in italics, species of non-nasute genera in brackets). 


abassas, 114 fuscus, 132 
abruptus sp. n., 145 
aethiops, 48 gemellus, 87 
africanus, 55 geminatus, 92 
agricola, 114 giffardii, 161 
aquilinus sp. n., 147 gratiosus, 97 
arabiae, 82 grootfonteinsis, 87 
arborum, 20 £VvOSSUS, 92 
auriceps, 132 
auritervae, 100 hainesi, 139 
havilandi, 132 
bequaerti, 45 hentschelianus, 132 
bettonianus, 84 hill, 133 
bipartitus, 148 hirticeps sp. n., 36 
(bouvieri), 5 holmgreni, 127 
brunneus, 61 
brutus, 114 ibidanicus, 92 
bulbiceps, 132 impetus, 45 
incuYvUS, 41 
camerunensis, 26 indoensis, 41 
carbo, 97 infuscatus, 37 
carbonarius, 127 
chapini, 48 kalaharicus, 114 
chrysopleura, 27 katangensis, 87 
coarctatus, 72 kempae, 39 
coatoni sp. n., 58 kohhi, 30 
congoensis, 153 konduensis, 48 
contracolor, 167 kulloensis, 87 
contractus, 61 kurumanensis, 132 
convergens, 149 
crassinasus, 84 latifrons, 41 
loubetsiensis, 114 
diabolus, 29 lujae, 45 
diplacodes, 114 lutzi, 100 
dispar, 87 
disparioides (‘‘form’’ = ssp.), III macrophthalmus, 30 
doriae, 52 maculiventris, 20 
dubius, 132 mallyi, 65 
dulcis, 48 maudanicus, 100 
meringocephalus, sp. n., 75 
ebenerianus, 92 messorY, 132 
edentatus, 160 minusculus, 30 
ekunduensis, 45 mobilis, 105 
eldivensis, 87 muneris, 114 
elegantulus, 30 
evythreae, 87 nanus, 45 
expulsus, 41 nigeriensis, 122 
ferranti, 32 occidentalis, 100 


fulleri, 34 oeconomus, 105 


72 


parviconstrictus, 156 


ervfusca (‘‘var.’’ = ssp.), 20 
P 


pius, 41 
posselensis, 127 
pretoriensis, 111 
(profestus), 5 
putidus, 48 


rapulum, 110 
rhodesiensis, 114 
vosent, 114 
ruficeps, 84 
vufonasalis, 114 


santschit, 45 
saudiensis sp. n., 120 
schoutedeni, 48 
schubotzianus, 106 
segelli, 84 

sjostedti, 41 
(somaliensis), 5 
sorex, 163 
(sudanicus), 5 


INDEX 


suffuscus, 68 
sulciceps (ssp.), 100 
suspensus, 122 


tchadensis, 105 
tenebricus, 68 
terrestris, 122 
tervvicola, 122 
thermarum, 114 
thomsent (ssp.), 132 
togoensis, 122 
torquatus, 29 
trinerviformis, 132 
trinervius, 127 
trinervoides, 132 
tuberosus, 157 


ueleensis, 41 
umzinduzit, 87 
undulans sp. n., 150 
usambarensis, 37 


zuluensis, 111 


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is THE LEPTOCORISINAE \v, 
(HETEROPTERA : ALYDIDAE) ~~ 
OF THE WORLD 


I. AHMAD 


BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY SUPPLEMENT 5; 
LONDON: 10965 


las \ 
t | 


i wre) 


THE LEPPOCORISINAE 
(HETEROPTERA : ALYDIDAE) 
OF THE AWORED 


BY 


I. AHMAD \ | 


Dept. of Zool. & Entom., Univ. of Connecticut, Storrs, U.S.A. 


BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY SUPPLEMENT 5 
LONDON : 1965 


THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted im 1949, 1s 
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 Supplement No. 5 of the Entomological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 


© Trustees of the British Museum (Natural History) 1965 


TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 


Issued 19th November, 1965 Price {2 15s. 


THE. LEP TOCORISINAE 
(HETEROPTERA : ALYDIDAE) 


OF FHER: WORED 
By I. AHMAD 


CONTENTS 


Page 
SYNOPSIS 3 
INTRODUCTION ; - ‘ : : - : : : 3 
MATERIALS AND METHODS . : ; ; , - : ; : 4 
ABBREVIATIONS USED IN TEXT-FIGURES 5 
FaMiILy ALYDIDAE 6 
KEy TO SUBFAMILIES 6 
LEPTOCORISINAE 7 
Kry To GENERA : P 9 
ZOOGEOGRAPHY AND PHYLOGENY - : F : . ; : 146 
ACKNOWLEDGMENTS . . : : : : , : ; , 150 
REFERENCES . : . P : : ; : - : 2 I5I 
INDEX . : , 2 , : : : : : : : 155 


SYNOPSIS 


Prior to this revision, the subfamily Leptocorisinae Stal contained eight genera and thirty 
eight species, including major rice pests of the Far East. There was confusion over the identities 
of many species and since Stal (1873) no world-wide revisional work had been undertaken and in 
his work only fourteen species (including some synonyms) were covered. 

During the present study, three new tribes, one new genus, two new subgenera, nineteen new 
species have been described and one species, because of its homonymy, has been renamed. 
Stenocoris Burmeister (1839) (subgenus sensu Stal, 1873) has been given generic rank and has 
been separated from Leptocorisa Latreille, 1829. Rhabdocoris Kolenati, 1845 (subgenus of 
Leptocorisa Latreille sensu Stal, 1873) has been synonymized with Leplocorisa. The undescribed 
mis-identified type-species of the subgenus Evbula has been renamed and described. Nine 
previously described species have been considered synonyms, whilst four species regarded as 
synonyms by earlier workers have been reinstated. 


INTRODUCTION 


LEPTOCORISINAE Stal is a subfamily of the coreoid family Alydidae and contained, 
prior to the present revision, eight genera and thirty eight species, including a 
number of important rice pests of the Indochina, Malaya and Australian Sub- 
regions. 

In the whole subfamily, there has been confusion and uncertainty over the identi- 
fication of species. For example, China (1930), in “‘ Insects of Samoa and other 
Samoan terrestrial Arthropoda ’’, expressed his doubts on the specific identity of 
Noliphus discopterus Stal and stated that perhaps it was a local race of N. insularis 


4 I. AHMAD 


Stal. Villiers (1955), formulating keys for the species of French African Stenocoris 
Burmeister, then regarded as a subgenus of Leptocorisa Latreille, suggested the 
synonymy of S. (Erbula) distinguenda (Blote, 1937) with S. (E.) elegans (Bl6te, 1937). 

During the present work the forms of male and female genitalia (Ahmad & 
Southwood, 1964) have been found to be of great taxonomic value, clearly separating 
the species. The structure of female genitalia, and in particular the shape of 
spermatheca, have suggested that Oriental and Australian species of Leptocorisa 
Latreille, 1829, (previously considered under the subgenus Rhabdocoris Kolenati, 
1845) should be separated from Ethiopian, Nearctic and Neotropical species (pre- 
viously considered under the subgenera Stenocoris Burmeister, 1839, Evbula Stal, 
1873 and Leptocorisa s. str. Stal, 1873). Other characters found to be of value are 
as follows: 

(a) The form of head, its colouration and its sutures. 

(b) Length and colour of antennal segments. 

(c) Size of rostrum and length of the components. 

(d) The comparison of the interocellar distance with the distance between an 

ocellus and an eye (in both cases including the diameter). 

(e) Length, breadth and the colour of pronotum. 

“({) The posterior margin of seventh abdominal tergum in g and the seventh 
abdominal sternum in 9. 

(g) In, the shape of the pygophore, claspers, thecal appendages and the form 

of vesica when fully everted. 

(h) In @ the first gonocoxae, first gonopophyses, second gonopophyses and 

spermathecal shape. 

Three of Bergroth’s genera, Dicrorymbus, Xenoceraea (described within Lepto- 
corixaria near Mutusca Stal, 1866 in 1918) and Procamptus, also described under 
Leptocorixaria in 1925, have been excluded from the subfamily Leptocorisinae and 
have been transferred to the tribes Dasynini Brown, 1955 and Procamptini trib. n. 
of the subfamily Coreinae (sensu Stal, 1873) (Ahmad, 1964). 


MATERIAL AND METHODS 


A large quantity of material (about 10,000 specimens) of the group was borrowed 
from major museums and personal collections from all over the world. (A complete 
list of these museums and of those who lent their personal collections is given under 
Acknowledgments.) Freshly prepared material (in 70% alcohol) was also obtained 
by the courtesy of Dr. J. W. Evans (Australian Museum, Sydney) and Dr. G. 
Rothschild (Sarawak Agricultural Institute). 

Every effort was made to examine and dissect the genitalia of the holotypes of all 
species (including those believed to be synonyms). In the great majority of cases 
it has been possible to do this or obtain authoritative information about the loss or 
damage of the original holotype, so necessitating the designations of neotypes. 
Lectotypes have also been designated from syntypes in a few cases. A full account 
of the circumstances and action taken is given for each species. 

gd and @ genitalia were dissected after softening the terminalia in 10% potassium 


THE LEPTOCORISINAE OF THE WORLD 5 


hydroxide solution. It was found easiest to inflate the aedeagus in water just after 
it was softened. The form of the genitalia was considered to be a reliable character 
for distinguishing species, as little intergradation was found between its form in two 
allied species and within a species as many as twenty five examples from widely 
separated localities were dissected and found to have similar genitalia. Large 
structures were stained in 1% acid fuchsin in acetic acid and were examined in 
methyl benzoate, cedarwood oil or creosote. All the drawings were made by using 
a squared eyepiece under a binocular microscope, from the type specimen when 
available. Genital structures were preserved in glycerine in small tubes and were 
pinned with the insects. 

All measurements were taken by using micromillimetre eyepiece graticules. 
When available, five specimens of each sex were measured and extreme ranges have 
been given. The length of the head was measured from the apex of the clypeus up 
to the anterior margin of the collar of the pronotum and the width was taken across 
the compound eyes including their diameter. Where very large numbers of speci- 
mens (e.g. L. acuta) were available it could be seen that the measurements of even 
the exceptional specimens did not lie far outside the ranges obtained by the above 
method. There is little variation within a species in the size of the parts measured. 

The distributions given for each species are based largely on the material seen by 
the present author, since there has been so much confusion over the identification of 
the species in the past. In general, published records have been ignored except 
those given by the authors who originally described the species. 


ABBREVIATIONS USED IN TEXT-FIGURES 


Ap apodeme of the basal plate (pivot) Ls lateral stapes 
At attaching apodeme of pivot Or outer ramus 
At2 attachment of second gonocoxa P claspers (parameres) 
Bp basal plate PaGpr point of attachment of first gonapo- 
Cc conjunctiva physis 
Ca conjunctival membrane PaRr point of attachment of the first 
Cl clypeus ramus 
Cm connecting apodeme of the aedeagal Ph phallosome (theca) 
strut PE process of 9th segment 
Da dorsal thecal appendage BS pronotal spine 
Dea dorsal conjunctival appendage Pt paratergite 
Dla dorso-lateral conjunctival appendage R ramus 
Ex exosoma (theca) Ra right lateral conjunctival appendage 
F fold of the 9th segment Rb rib of 9th segment 
Fa front conjunctival appendage S stapes 
Fs femoral spine Sa supporting appendage 
G genae Sl spine-like appendage 
Gg gonangulum Sl2 second spine-like appendage 
Gp gonapophysis St sternum 
Gp2 second gonapophysis Ta terminal appendage 
Gx gonocoxa Ts oe sstretnre 
Ivs intervalvular sacs Tw tubular way to genital chamber 
Ju paraclypeae (jugum) We vesica 
La left lateral conjunctival appendage Va ventral thecal appendage 


6 I. AHMAD 


ALYDIDAE Amyot et Serville 


Alydides Amyot et Serville, 1843 : 221 

Alydidae Dallas, 1852 : 467 ; Douglas and Scott, 1865 : 18 

Alydida Stal, 1868 : 542 

Alydina Stal, 1873 : 34, 84 

Alydinae Distant, 1880 : 156 

Alydidae Reuter, 1912 : 49 

Coriscidae Stichel, 1925 : 45 ; Blatchley, 1926 : 256 

Alydidae Southwood, 1955a: 80; lLeston and Scudder, 1956: 231; Leston, 1958: gII ; 
Southwood and Leston, 1959 : 65 ; Scudder, 1959 : 418 

Alydinae China and Miller, 1959 : 7, 40 

Alydidae Wagner, 1961 : 141 


Macropterous or at least brachypterous species, body length usually more than 10 mm. ; 
head proportionally much larger ; the part between the eyes wider than base of scutellum ; 
eyes usually very prominent ; ocelli usually large, usually narrowly separated ; apical seg- 
ment of antennae as long as or usually longer than third ; bucculae short, not extending pos- 
teriorly beyond the insertion of antennae, sometimes entirely anterior to antenniferous 
tubercles ; metapleural scent gland ostioles usually distinct ; spiracles ventral in normal 
position, away from lateral margins ; fourth and fifth dorsal abdominal segments with basal 
margins lobately sinuate into preceding segments ; legs long, posterior femora usually thick- 
ened, in § sometimes with spines ; first joint of tarsi usually more than twice as long as second 
and third together ; usually narrow, elongate or cylindrical, dark coloured species. 

6 genitalia usually very complicated and variable ; in 2, second gonocoxae fused with each 
other and with second gonopophyses ; complicated sclerotized expansions present, supporting 
the genital chamber (Scudder, 1959). In eggs, pseudo-operculum absent (Southwood, 1956) ; 
usually four-lobed principal salivary glands present and accessory ducts with swelling midway 
along its length (Bugnion, 1910 and Southwood, 1955a) ; in larvae, tergum five usually medially 
with anterior and posterior margins deflected posteriorly and head pointed anteriorly, longer 
than broad (Leston and Scudder, 1956) ; chromosome number, 2x = 13 (xo) (Leston, 1958). 


Alydidae can easily be separated from other related families by the presence of 
very short bucculae, not extending posteriorly beyond the insertion of antennae. 


KEY TO SUBFAMILIES 


i Pronotum at widest point only slightly wider and longer than head. (Aedeagus 

usually without ventral pair of thecal appendages) . 2 
— Pronotum at widest point usually at least half as wide aed as a Beene Tees: 

than head. (Aedeagus always with a ventral pair of thecal appendages and usually 

very complex with a number of lateral, frontal and dorsal conjunctival appendages. 

In 9, second gonopophyses usually rounded at apices ; spermatheca usually flask- 

shaped, sometimes with a median flange) : : . LEPTOCORISINAE 
2 Second rostral segment usually distinctly longer ae posterior two together ; 

fourth rostral segment about twice as long as third; hind femora never with spines. 

(In 3, pygophore with a median posterior spine and aedeagus with a dorsal pair of 

thecal appendages. In 9, posterior margin of seventh abdominal sternum usually 

with a median split ; both pairs of rami fully developed ; spermatheca with a long 

coiled tube) . A : . MICRELYTRINAE 
— Second rostral segment aoa distinctly Shorter wee posterior two together ; 

fourth never twice as long as third ; hind femora swollen, always with a series of 

ventroposterior spines. (In 3, pygophore without a median posterior spine and 

aedeagus without thecal appendages. In 9, posterior margin of seventh abdominal 

sternum emarginate without a median split, both pairs of rami very short and 

thin ; spermatheca with a short connecting tube) . , : : ALYDINAE 


THE LEPTOCORISINAE: OF THE WORLD a 


LEPTOCORISINAE Stal 


Gerrida Stal, 1868 : 543 

Gerrina Stal, 1870 : 217 

Leptocorisaria Stal, 1872 : 54, 1873 : 85 ; Uhler, 1886 : 13 
Leptocorisidae Lethierry and Severin, 1894 : 102 

Myodochinae Kirkaldy, 1902 : 306 

Leptocorisaria Stal ; Distant, 1902 : 409 ; Oshanin, 1906 : 24 
Leptocorixaria Bergroth, 1913 : 159 

Leptocorisini Blatchley, 1926 : 257, 259 

Leptocorisinae Tillyard, 1926 : 148 


Head elongated ; epicranial suture (ecdysal cleavage line) usually present ; occipital suture 
usually complete ; ocelli minute to large, usually centrally placed behind the compound eyes. 
First rostral segment extending well beyond the compound eyes ; second rostral segment 
usually distinctly shorter than posterior two together, fourth usually almost equal to third. 
Pronotum at widest point usually at least half as wide again as and distinctly longer than head. 
Posterior angle of metasternum acutely produced. Posterior margin of second dorsal segment 
of abdomen truncate. Hamus (hook vein) of hind wing and vena decurrens not at all or little 
distant from base. Legs slender ; hind femora always without spines. 

6 genitalia, especially the aedeagus, highly specialized with a long coiled vesica and usually 
with a number of lateral, frontal and dorsal conjunctival appendages ; dorsal and ventral pairs 
of thecal appendages usually present (Text-fig. 1). 

Q genitalia highly specialized ; apices of second gonopophyses rounded ; gonangulum 
distinctly marked, with the first inner ramus attached ventrolaterally connecting first gonopo- 
physis with it, outer ramus of gonopophysis usually marked ; second gonocoxae elongated, 
always with a long, curved and very distinct second ramus connecting second gonopophyses 
with it ; spermatheca usually balloon-shaped, with a coiled tube, sometimes with a median 
flange. 


be ee 


“ Re TERMINAL APP 


GONOPORE ~-—--- -=--~--Ist | 


VESICA - ~ ---- "oa 


SUPPORTING “APP [S- ===> ive SPINE LIKE APP 


_ bt 
--- 4th 
a - ord 

RIGHT LATERAL 
---— 2nd [ CONJUNCTIVAL APP 


‘T’ APPARATUS - - ~~ -- 


FRONTAL APP- - --- 
DORSO -LATERAL _- Yd f | 
conuuncrival (7 ona) fat t 

APP We We 


VENTRAL THECAL” ~~ a 

AP heh 
DORSAL THECAL ._. Lt 
APP rs 

‘ot 

DORSAL KNOB LIKE _ 1 
PROCESS / 


EJACULATORY DUCT - --—- 


BASAL PLATE------ 


Fic. 1. Hypothetical 3 aedeagus, fully everted. 


8 I. AHMAD 


This subfamily can be distinguished from Micrelytrinae by the length of second 
and fourth rostral segments. In the latter, the second rostral segment is usually 
distinctly longer than posterior two together and the fourth is always at least twice 
as long as the third. In Alydinae the hind femur is always armed whereas in this 
subfamily it is always without spines. Other distinctive characters are given 
under ‘“‘ Key to the subfamilies ”’. 


0:5 mm. 
2 


Fic. 2. Stenocoris (Oryzocoris) filiformis, 2, dorsal view. 


THE LEPTOCORISINAE OF THE WORLD 9 


KrEy TO GENERA 


1 Paraclypeae produced and meeting in front of clypeus tienen 3), dorsal surface of 
pronotum more or less flat (Leptocorisini trib. n.) : ; 2 
— Paraclypeae shorter than clypeus and never enclosing its apex (Text- fig. 5) ; dorsal 
surface of pronotum usually convex and sloping down towards head (Text-figs. 4, 
6 and 7) (Noliphini trib. n.) . : : 2 : ; 5 
2 First antennal segment swollen Pvadhout (Text- fig. 3), slender only at base ; in 
lateral view wider beyond middle. (In g, posterior margin of seventh abdominal 
tergum roundly produced in middle ; in 9, posterior margin of seventh abdominal 


sternum with a median fold splitting it into two lobes) : . MUTUSCA (p. 110) 
— First antennal segment only swollen at apices ; in lateral view not wider beyond 
middle : - ‘ : 3 


3 Genae inflexed, exeathy produced in cone nf Giyrions (Text- fig. 8). (In 3, posterior 

margin of seventh abdominal tergum medially produced with a truncated apex, 

lobes of ninth segment greatly developed, usually produced like horn ; in Q, 

posterior margin of seventh abdominal sternum usually conically produced in 

middle, sometimes with a short median split. : . BLOETEOCORIS (p. 115) 
— Genae normal : é : : 4 
4 Claspers made up of two components; always crossed behind ; in 2, posterior 

margin of seventh abdominal sternum never distinctly medially produced, never 

with a median fold, bilobed or with a median split ; spermatheca always with a 

median flange ; genus only represented in Oriento-Australian regions. 

LEPTOCORISA (p. 69) 

— Claspers never made up of two components, never crossed behind ; in Q, posterior 

margin of seventh abdominal sternum either medially produced, with a median 

fold, bilobed or with a median split ; genus son pat in Ethiopian, Nearctic and 

Neotropical regions : 3 STENOCORIS (p. 9) 
5 Lateral angles of pronotum posteriorly armed rent. spines (Text-fig. 6). In g, 

pygophore normally more than half retracted within seventh ; in 9, posterior 

margin of seventh abdominal sternum with a median fold . . NOLIPHUS P. 131) 
— Lateral angles of pronotum posteriorly without spines . ; 6 
6 Anterior femora on undersurface below apex with a distinct spine (Text- fig. 9); 

ocelli very closely placed, almost touching each other ; disc of pronotum always 

black (Text-fig. 4) ; found in Oriento-Australian regions . COSMOLEPTUS (p. 126) 
— Anterior femora without a spine, ocelli not at all touching each other ; disc of prono- 

tum of pale colour (Text-fig. 7) ; found in Neotropical region . LYRNESSUS (p. 123) 


LEPTOCORISINI trib. n. 


Body linear. Head always longer than broad ; paraclypeae very well developed, always 
distinctly longer than and enclosing clypeus ; head behind the eyes not markedly narrower. 
Pronotum never broader than long, almost rectangular and flat. Aedeagus usually complicated. 
Spermatheca usually flask-shaped, never conical, sometimes with a median flange. 


This tribe is distinguished from Noliphini by the characters of the anterior region 
of the head and pronotum. 


STENOCORIS Burmeister. 


Myodocha Latreille, 1810 ; Burmeister, 1835 : 325 
Stenocovis Burmeister, 1839 : 1010 ; Stal, 1873 : 87 [subgenus] ; Bl6éte, 1937 : 289 [subgenus] ; 
Villiers, 1955 : 1142 [subgenus] ; Villiers, 1963 : 232 [subgenus] 


Type-species : Cimex tipuloides Degeer. 


10 I. AHMAD 


Body linear or comparatively robust. Head elongated. Antennae with apices of basal 
segments slightly swollen. Rostrum usually short, reaching second coxae. 
In g, posterior margin of seventh abdominal tergum usually truncate ; pygophore usually 


1:0 mm. 


Fics. 3-9. 3, Mutusca proliva, head, dorsal view ; 4, Cosmoleptus limbaticollis, pronotum, 
dorsal view ; 5, Noliphus erythrocephalus, head, dorsal view ; 6, Noliphus erythroce- 
phalus, pronotum, dorsal view ; 7. Lyrnessus geniculatus, pronotum, dorsal view ; 8, 
Bloeteocoris inflexigena, head, lateral view ; 9. Cosmoleptus limbaticollis, portion of femur 
and tibia, inner view showing ventro-lateral fore femoral spine. 


THE LEPTOCORISINAE OF THE WORLD II 


elongated ; claspers variable in form and position ; aedeagus usually with a ventral pair of 
thecal appendages ; frontal conjunctival appendages usually present, sclerotised ; mem- 
branous appendage short, usually without a terminal appendage but usually with a pair of 
dorsal appendages. 

In 8, posterior margin of seventh abdominal sternum either bilobed, medially produced, with 
a short median split or concave with a median fold ; first gonocoxae usually large, and tri- 
angular ; intervalvular sacs usually absent ; spermatheca usually flask-shaped, without a 
median flange and usually with a long coiled tube. 

This genus shows some affinities with all the other genera in the tribe Leptocorisini 
and indeed to Lyrnessus and Cosmoleptus sumatranus in Noliphini. It could possibly 
be polyphyletic or have retained many of the characters of the more generalized 
ancestral form. 

As Myodocha Burmeister nec Latreille was an erroneous identification as well as 
an emendation of Myodocha Latreille, it is not necessary that the replacement name 
Stenocoris Burmeister, 1839 should take the same type as that of Myodocha Latreille. 
The latter is M. serripes Olivier, 1811 (a Lygaeid) and has been so designated under 
the Plenary Powers by the International Commission on Zoological Nomenclature 
(1961, Bull. zool. Nomencl., 18 : 287). Stenocoris is not a new name but is in fact a 
new generic name for the taxon wrongly identified by Burmeister (1835). The type- 
species of Stenocoris is one of the two species cited by Burmeister in 1835, 1.e., 
varicorms Fabricius, 1803 and tipuloides Degeer, 1773. It has been fixed by 
Burmeister (1839) as Cimex tipuloides Degeer, 1773. 

Stenocoris was reduced to subgeneric rank within Leptocorisa Latreille, 1829 by 
Stal. In fact Stal misidentified Leptocorisa Laporte nec Latreille for Leptocorisa 
Latreille. The most senior subgenus Stenocoris is therefore hereby proposed as the 
generic name for the taxon. An application with Dr. W. E. China has been made 
to the International Commission on Zoological Nomenclature for its recognition (1963). 


Kry TO SUBGENERA 


al 


Claspers never crossed, when at rest parallel to one another (Text-fig. 19) ; posterior 
margin of 2 seventh abdominal sternum always medially produced (Text-fig. 22), 
sometimes notched in the middle, never bilobed, concave or with a median split. 

ERBULA (p. 14) 

— Claspers crossed (Text-fig. 75) ; posterior margin of 2 seventh abdominal sternum 

either bilobed teen 82), concave (Text-fig. 15) or with a short median split 
(Text-fig. 181) : ; : : : : : : : : 2 

2 Claspers curved like an “ ‘S ne apex thick and truncated (Text-fig. 11) ; apical 

segment of antennae with at least basal half pale ; posterior margin of 9 seventh 
abdominal sternum concave with a median fold PSEUDOLEPTOCORISA (p. 12) 

— Claspers never curved like an “S’’, apex always acutely pointed (Text-fig. 76) ; 

apical segments of antennae with small pale area at base, never extending to half 

the length of the segment ; posterior amuas of 2 seventh abdominal sternum 
never concave with a median fold . : : : ; p k : 3 

3 Posterior angles of pygophore acutely pointed ; claspers crossed over pygophore 

(Text-fig. 178) ; posterior margin of 9 seventh abdominal sternum always with a 
short median split 3 - ORYZOCORIS (p. 60) 

— Posterior angles of pygophore not noutaly pointed : ‘elaspers crossed in a socket 

(Text-fig. 75) ; posterior margin of 9 seventh abdominal sternum always medially 

bilobed, never witha split. : c : : : STENOCORIS (p. 30) 


12 I. AHMAD 


STENOCORIS (PSEUDOLEPTOCORISA) subégen. n. 


Type-species : Leptocorixa (Rhabdocoris) erratica Blote. 


Body comparatively robust. Head only a third longer than broad ; paraclypeae long, about 
two-fifths the length of head, with pointed apices ; ocelli very close together, inter-ocellar 
distance only slightly less than half the distance between an ocellus and an eye. Antennae 
with at least basal halves of apical segments pale. Rostrum short, reaching second coxae. 
Lateral black line strongly represented, extending from base of antenniferous tubercles to 
posterior of propleuron. Pronotum distinctly sloping down towards head, with latero-posterior 
tips light brownish black. Abdomen dorsally pale blackish ; abdomen ventrally dull 
ochraceous. In g, posterior margin of seventh abdominal tergum produced, with a slight 
median notch ; pygophore round ; claspers extending behind pygophore, asymmetrically 
crossed, ‘‘S ’’-shaped with broad truncated apices ; aedeagus with dorsal and ventral pairs of 
thecal appendages ; frontal conjunctival appendage absent ; membranous appendage present 
but supporting and dorsal appendages wanting ; vesica with very small gonophore. 

In 9, posterior margin of seventh abdominal sternum concave with median fold ; first 
gonocoxae very large with lobe-like apices ; two pairs of intervalvular sacs ; spermatheca 
flask-shaped, medially thick with long curved neck, and very short tube. 


This subgenus is isolated in Stenocoris but exhibits the characters of the genus in 
body shape, in lacking supporting appendages, in the form of membranous appendage 
of the aedeagus and in having curved “ S ’’-shaped claspers composed of one compo- 
nent and a plain flask-shaped spermatheca without a median flange. However it is 
close to Leptocorisa in having a rounded pygophore, dorsal and ventral pairs of thecal 
appendages of the aedeagus and in the concave posterior margin of the female seventh 
abdominal sternum. It differs from both Leptocorisa and Stenocoris by the absence 
of frontal conjunctival appendages and in having only two pairs of intervalvular sacs. 
Pale basal halves of apical segments of antennae, lateral black line extending up to 
the posterior of propleuron, claspers with thick and truncated apices and the median 
fold on the posterior margin of the seventh abdominal sternum in the female are 
unique characters of this subgenus. This subgenus also shows some relations with 
Lyrnessus geniculatus (Guérin) and Cosmoleptus sumatranus Blote of the tribe Noliphini 
in the characters of the hemielytra and apical antennal segment. 


Stenocoris (Pseudoleptocorisa) erratica (Bléte) comb. n. 
(Text-figs. 10-18) 
Leptocorixa (Rhabdocoris) ervatica Blote, 1937 : 286; Villiers, 1955 : 1143, 1963 : 234 


Characterized by the pale basal half of the apical segment of antennae, by the 
round pygophore and by ¢ and @ genitalia. 


3d. Body length, 13-0-14-0 mm., width, 2:0-2:2 mm. Head length, 1-7-1-8 mm., width, 
I:3-I-4 mm., (only paraclypeae, 0-7 mm.) Antennae, segment I, 3:7—4-7 mm., II, 2-1-2-6mm., 
III, 2-7-3:2 mm., IV, 4:2-4-9 mm. Rostrum, segment I, 1-2-1:3 mm., II, 1-3-1-4 mm., III, 
o-7-0:8 mm., IV, 0-6-0:77 mm. Pronotum, length, 2-2-2:5 mm., width 2-0-2:2 mm. Hind 
femora, 5-6-6-4 mm. ; hind tibia, 6-1-7-2 mm. Distance between ocelli, 0-4-0-5 mm., distance 
between an ocellus and an eye, 0-66—0-7 mm. 


THE LEPTOCORISINAE OF THE WORLD 


Fics. 10-18. Stenocoris (Pseudoleptocorisa) evvatica. to, dg terminalia, dorsal view ; 11,3 


claspers, outer and inner views ;_ 12, 9, first gonocoxae, ventral view ; 


13, spermatheca; 


14, d aedeagus (inflated), ventral view ; 15, 9 terminalia, ventral view ; 16, Q, first 
gonopophysis, ventral view ; 17, ovipositor, ventral view ; 18, same, dorsal view. 


13 


14 I. AHMAD 


Body comparatively robust, pale tinged black. Head only a third longer than broad with a 
pair of brownish black spots behind ocelli ; occipital suture medially concave ; paraclypeae 
thicker in middle and pointed at apices. Antennae dark brownish black with underside of 
first, articulations between first and second, second and third and about basal half of fourth 
pale. Rostrum short, hardly reaching second coxae. Lateral black line extending from 
antenniferous tubercles on to posterior of propleuron. Pronotal disc pale, posteriorly tinged 
black with two smooth black spots anteriorly on either side, black spots on latero-posterior tips 
absent. Legs pale with apices of femora, tibiae and tarsi tinged black. Hemielytra tinged 
brownish black with prominently pale veins. Posterior margin of seventh abdominal tergum 
slightly produced in middle ; pygophore round, posterior margin medially acutely produced 
(Text-fig. 10). 

6 GENITALIA : Claspers extending behind pygophore, strongly curved in middle, more or 
less ‘‘S’’-shaped, blunt at apices ; both dorsal and ventral pairs of thecal appendages present, 
ventral, sac-like, 2 pairs of asymmetrical spiculed lateral appendages ; membranous appendage 
short with a small round terminal appendage ; frontal, dorsal and supporting conjunctival 
appendages wanting ; vesica with a minute gonopore (Text-fig. 14). 

@ very similar to g in colour, general appearance and measurement. Posterior margin of 
seventh abdominal sternum concave in middle with a small median fold (Text-fig. 15). 

@ GENITALIA : First gonocoxae almost conical when viewed externally, lobe-like at apices ; 
two pairs of intervalvular sacs, anterior round and posterior pair oval, spermatheca flask-shaped, 
thicker in middle with a very short tube (Text-fig. 13). 


Type material. Holotype 9, UGANDA: Southern Toro, Mbarara, Fort Portal 
Road, 3,888—4,200 ft., 22-24.x. (S. A. Neave), B.M.(N.H.) ; allotype 9 and paratypes, 


43,5 9, data as holotype, B.M.(N.H.) ; Tero Forest, 23, 2 9, vii.1g12 (C. C. Gowdey), 
B.M.(N.H.). 


Other material. UGANDA: 10g,132 ; ConGo: 269,192 ; CAMEROONS : 164, 
16 Q. 


This species seems to be very common in Congo and its range extends into Uganda 
in the east and Cameroons in the west. The adults have been collected in every 
month of the year except January, August and September. 


This species is isolated in the genus Stenocoris with the characters of blunt “S ’’- 
shaped claspers, rounded pygophore, concave posterior margin of 2 seventh abdom- 
inal sternum and pale basal half of apical antennal segment. 

Blote described this species under the subgenus Rhabdocoris Kolenati of Leptocorixa 
Berthold. During the present study Rhabdocoris is synonymized with Leptocorisa 
and the latter has been separated from other subgenera with Oriento-Australian 
species, previously considered under the subgenus Rhabdocoris. This species, being 
different from Leptocorisa in generic characters, is placed under the monotypic 
subgenus Pseudoleptocorisa as described here. 


STENOCORIS (ERBULA) Stal. 
Erbula Stal, 1873 : 86; Blote, 1937 : 282 ; Villiers, 1955 : 1141, 1963 : 229 
Type-species : Stenocoris (Erbula) southwoodi sp. n. 


Body comparatively short. Head usually only slightly longer than broad ; paraclypeae 
short and rounded at apices ; ocelli close together but distance between an ocellus and an eye 
about half again as long as interocellar distance. Antennae with apical segment pale only at 


THE LEPTOCORISINAE OF THE WORLD 15 


base ; second usually distinctly shorter than third. Rostrum short, never extending beyond 
second coxae. Lateral black line usually present but never extending beyond middle of 
propleuron. Pronotum slightly sloping down towards head with latero-posterior tips usually 
black. Abdomen usually pale dorsally, ventrally usually dull ochraceous. 

In g, posterior margin of seventh abdominal tergum truncated ; pygophore usually rounded ; 
claspers short, curved medially with apices acutely pointed, never crossed ; aedeagus with 
dorsal and ventral pairs of thecal appendages ; dorsal pair united ; single frontal conjunctival 
appendage bifurcated at apices ; membranous appendage comparatively long with single dorsal 
appendage, terminal appendage wanting. 

In 9, posterior margin of seventh abdominal sternum medially produced, usually with a 
median notch ; first gonocoxae elongated ; intervalvular sacs absent ; spermatheca elongated 
with short tube coiling round neck. 


Males of Erbula can be distinguished from those of other subgenera by examining 
the claspers which are never crossed and the females in having the posterior margin 
of the seventh abdominal sternum with a median process which is sometimes 
notched in the middle, but never indented, concave, or with a short median split 
and in having the spermatheca with a short tube coiled round its neck. 


Mis-identification of Stenocoris (Erbula) southwoodi sp. n. for Stenocoris apicalis 
(Westwood, 1842), the type of the subgenus and the request for its recognition are the 
subjects of an application to the International Commission on Zoological Nomencla- 
ture (Ahmad, in press). 


KryY TO SPECIES 


1 Lateral black line very faintly represented, usually in the form of a black dot on the 
lateral side of collar ; pygophore elongated ; in Q, posterior margin of seventh 
abdominal sternum with a central median round a which has a distinct 
median notch ; ‘ : ; similis (p. 23) 

— Lateral black line usually strongly represented, extending. at least on to collar ; 
pygophore usually rounded ; in Q, posterior margin of seventh abdominal sternum 


either triangular, rounded or parallel-sided, if rounded never with a distinct notch 2 
2 Claspers short, nearly reaching apex of pygophore ; in 9, median projection of seventh 

abdominal sternum round, faintly striated (Text-fig. 59) . : southwoodi (p. 24) 
— Claspers long, always reaching apex of pygophore ; in 9, median anil of 

seventh abdominal sternum parallel-sided or triangular. ; 3 
3 Apex of pygophore pointed ; lateral black line never extending ioe ard on to senae ; 

rostrum usually short, reaching second coxae . : 4 
— Apex of pygophore medially notched ; lateral black line natally extending forward 

on to genae ; rostrum usually extending at least slightly beyond second coxae . 5 


4 Claspers broad in middle and knob-like at apices (Text-fig. 19) ; in 9, median projec- 

tion of seventh abdominal sternum a a ae with a distinct median notch 
(Text-fig. 22) ‘ . annulicornis (p. 16) 

— Claspers narrow in middle and acutely pointed at epices (Text- figs. 29 and 30) ; in 9, 

median projection of seventh abdominal sternum triangular (Text-fig. 31) 

distinguenda (p. 18) 

5 Claspers strongly outwardly curved (Text-figs. 37 and 38) ; in 9, median projection 

: of seventh abdominal sternum triangularly produced, with round apex (Text-fig. 41) 
elegans (p. 20) 

— Claspers only curved in middle (Text-figs. 65 and 66) ; in 9, median projection of 
seventh abdominal sternum rectangular (Text-fig. 70) ; : : stali (p. 28) 


16 I. AHMAD 


Stenocoris (Erbula) annulicornis (Signoret) comb. n. 
(Text-figs. 19-27) 


Leptocorisa annulicornis Signoret, 1860 : 941 
Leptocorixa (Stenocoris) annulicornis (Signoret) Bléte, 1937 : 289 
Leptocorixa (Evbula) annulicornis (Signoret); Villiers, 1963 : 230 


Characterized by the claspers which are broad towards the base and knob-like at 
apices, by the parallel-sided median projection of the 2 seventh abdominal sternum, 
with a distinct notch and by the form of 3 and @ genitalia. 


6. Body length, 12-0-12-5 mm., width, 1-6-1-83 mm. Head length, 1-5-1-6 mm., width, 
I-2-1-3 mm. Antennae, segment I, 3:0-3:3 mm., II, 1-7-2:0 mm., III, 2-0-2:3 mm., IV, 
3°8-4:0 mm. Rostrum, segment I, 1-1-1-2 mm., II, 1-2-1-3 mm., III, 0-65-0-7 mm., IV, 
o:6-0:'7mm. Pronotum, length, 1-9—2:0 mm., width, 1-6-1-°8 mm. Hind femora, 4-8-5-0 mm. 
Hind tibia, 5-1-5:3 mm. Distance between ocelli, 0-4 mm., distance between an ocellus and an 
eye, 0-6 mm. 

Body comparatively short, slim, tinged with brown. Head pale with area between the eyes 
and behind occipital suture brownish black ; occipital suture medially concave with sides 
convex ; paraclypeae short, thicker in middle and thinner at apices. Antennae brownish 
black, with underside of basal segment, median portion of second, articulation between first and 
second and that between second and third and basal portion of apical segment pale. Rostrum 
short, usually reaching base of second coxae. Lateral black line well represented on sides of 
antenniferous tubercles and extending to anterior of propleuron. Pronotal disc with brownish 
black punctures and black spots on posterior angles. Legs sometimes tinged with green. 
Hemielytra tinged with black. Abdomen with dorsum pale, tinged with brown, venter pale 
ochraceous, sometimes reddish anteriorly. Posterior margin of seventh abdominal tergum 
regularly curved, pygophore rounded with apex pointed. 

6 GENITALIA : Claspers broad towards base, external surface curved only in middle (Text- 
figs. 19 and 20) ; ventral thecal appendage as in Text-fig. 21 ; four left lateral conjunctival 
appendages as in Text-fig. 21 ; frontal appendage bifurcated in middle, sometimes left branch 
longer ; four right lateral conjunctival appendages as in Text-fig. 21 with spine-like appendage, 
thicker at base and apex ; dorsal appendage thick and curved at base as in Text-fig. 21. 

2. Body length, 13-0-14-0 mm., width, 2:0-2:2 mm. Head length, 1-6-1-73 mm., width, 
I-3-1°35 mm. Antennae, segment I, 3-0-3-1 mm., II, 1-8-1-9 mm., III, 2-0-2:2 mm., IV, 
3:°8-4-0mm. Rostrum, segment I, 1:33 mm., II, 1-33 mm., IIJ,o-7mm.,IV,0-7mm. _ Prono- 
tum, length, 2:25-2:35 mm., width 2:0-2:13 mm. Hind femora, 5-3—5-6 mm. ; hind tibia, 
5:8-6:0 mm. ; distance between ocelli 0-46 mm., distance between an ocellus and an eye, 
0-7 mm. 

Very similar in general appearance and colour but distinctly longer than g; seventh 
abdominal sternum with small median parallel-sided notched projection. 

Q GENITALIA : First gonocoxae thicker in middle and curved near apex with round apices ; 
intervalvular sacs absent, spermatheca flask-shaped with long neck (Text-fig. 25). 


Material studied. Mauritius: I g, 2 9; SEYCHELLES: I 9 ; MADAGASGAR : 
192; Conco : 184, 11 9. 


The range of this species extends in the west as far as the Congo (Ituri river) and 
in the east to Mauritius. It is also found in the Seychelles and Madagascar. It 
has mostly been collected during the early part of the year (from December until 
June), but it has also been taken in August. . 


This species is close to distinguenda (Blote) in having similar type of paraclypeae, 
lateral black line and the similar shape of the pygophore, but can easily be separated 


THE LEPTOCORISINAE OF THE WORLD 17 


by examining the g claspers, which are broad towards the bases and knob-like at 
apices in this species, whereas they are uniformly thin and acutely pointed in 
distinguenda. The small parallel-sided, medially notched projection of the 2 seventh 
abdominal sternum separates it from all the species of this subgenus. 


Fics. 19-27. Stenocoris (Evbula) annulicornis. 19-21, g, 22-27, 2. 19, terminalia, dorsal 
view ; 20, claspers, inner and outer views ; 21 aedeagus (inflated), ventral view ; 
22, terminalia, ventral view ; 23, first gonopophysis, ventral view ; 24, ovipositor, 
dorsal view ; 25, spermatheca ; 26, ovipositor, ventral view: 27, first gonocoxa, 
ventral view. 


18 I. AHMAD 


Stal (1873) synonymized this species with L. (E.) apicalis Westwood but in 1937 
Blote redescribed it from Mauritius and Seychelles, placing it in the subgenus 
Stenocoris. Signoret’s type material is in the Naturhistorisches Museum, Vienna, but 
Prof. Max Beier informed the author that it was on loan to Dr. A. Villiers and thus 
it was not possible to examine it. However, Dr. Villiers (personal communication) 
has selected a lectotype and has published figures of the terminalia (J, 9). These 
agree with the concept of annulicornis in this study. Villiers has pointed out that 
annulicorms of Blote was not the true annulicornis Signoret but another species, 
which he described as Leptocorixa blote: ; however the ¢ of this species is clearly 
S. phthisica (Gerstacker) (see page 70), but the 9 represents the true annulicorms and 
Villiers’ figures of the seventh abdominal sternum of the 9 of blotez and annulicornis 
are almost identical. 


Stenocoris (Erbula) distinguenda (Blote) comb. n. 
(Text-figs. 28-36) 


Leptocorixa (Erbula) distinguenda Blote, 1937 : 286; Villiers, 1955 : 1142 [as synonym of L. 
elegans] 


Characterized by the uniformly thin and smoothly tapering claspers, by the tri- 
angular median projection of the 2 seventh abdominal sternum and by the form of 
6g and genitalia. 


6. Body length, 12-0-12-5 mm., width, 1-7-1-8 mm. Head length, 1-6-1-7 mm., width, 
I‘I5-1:'25 mm. Antennae, segment I, 2-9-3-0 mm., II, 1:7-1-75 mm., III, 1-9-2: mm., IV, 
4:0-4:'2 mm. Rostrum, segment I, 1-1-1-3 mm., II, 1-2-1-3 mm., III, 0-6-0-7 mm., 
IV, 0-6-0:65 mm. Pronotum, length, 2:0-2:1 mm., width, 1-7-1-8 mm. Hind femora, 
4:7-4:9 mm. ; hind tibia, 5:2-5:-5 mm. Distance between ocelli, 0-4-0-43 mm., distance 
between an ocellus and an eye, 0-6—-0-65 mm. 

Body usually comparatively very short, pale, tinged with brownish black ; occipital suture 
concave in middle with lateral arms curving outwards towards sides of head ; paraclypeae 
short, thicker in middle, thinner and curved near apices. Antennae pale, with brownish black 
tinge, underside of basal segment, basal half of second, articulations of second and third and 
basal portion of apical segment pale. Rostrum short, approaching second coxae. Lateral 
black line well represented from sides of antenniferous tubercles to anterior of propleuron. 
Pronotal disc with brownish black punctures, posterior angles with black spots. Legs pale 
unicolourous, sometimes with apices of femora and bases of tibiae tinged with brown. Hemiely- 
tra with black punctures. Abdomen with dorsum except sides of connexiva yellowish red, 
venter pale ochraceous. Posterior margin of seventh abdominal tergum truncated, slightly 
convex in middle ; pygophore with pointed apex. 

6 GENITALIA: Claspers long, uniformly thin and distinctly reaching apex of pygophore, 
straight at apex ; ventral thecal appendages as in Text-fig. 28 ; four lateral conjunctival 
appendages as in Text-fig. 28 ; frontal appendage long, thinner in middle and bifid at apex with 
almost equal branches, four right lateral conjunctival appendages as in Text-fig. 28 with spine- 
like appendage thin in middle, apex blunt; dorsal appendage convex at base, extending 
concavely towards apex. 

2. Body length, 14-5-15-6 mm., width, 2-0-2: mm. Head length, 1-65-2:0 mm., width, 
I-3-I'4 mm. Antennae, segment I, 3-0-3:4 mm., II, 1-8-2-0 mm., III, 2:03-2:2 mm., IV, 
4:0-4:3 mm. Rostrum, segment I, 1-3-1-33 mm., II, 1-3-1-33 mm., III, 0-7-0-8 mm., IV, 
0:-6-0'7 mm. Pronotum, length, 2:3-2:5 mm., width, 2:0-2-1 mm. Hind femora, 5-o—-5-6mm. 


THE LEPTOCORISINAE OF THE WORLD 19 


Hind tibia, 5-5-5-6 mm. Distance between ocelli, 0-4-0-5 mm., distance between an ocellus 
and an eye, 0-6-0-7 mm. 

Very similar to f in general appearance and colour, but distinctly longer and slightly darker ; 
posterior margin of seventh abdominal sternum with triangular median projection following 
deep curve on either side. 


0:5 mm. 


Fics. 28-36. Stenocoris (Erbula) distinguenda. 28-30, 3, 31-36, 9. 28, aedeagus (inflated), 
ventral view ; 29, claspers, outer and inner views ; 30, terminalia, dorsal view ; 31, 
same, ventral view ; 32, first gonocoxa, ventral view ; 33, spermatheca; 34, first 
gonopophysis, ventral view ; 35 ovipositor, ventral view ; 36, same, dorsal view. 


20 I. AHMAD 


@ GENITALIA: First gonocoxae very narrow and curved at apices, outer margin regular ; 
intervalvular sacs absent ; spermatheca flask-shaped, with a short and curved neck as in 
Text-fig. 33. 


Type material. Holotype g, NYASALAND (head and hind tibia missing), B.M. 
(N.H.) ; allotype 9, with similar locality data (antennae missing), B.M.(N.H.). 


Other material. CONGO: 15, 18 9. 


This species seems to be represented in the south eastern region as far as Nyasa- 
land. Adults are found in the early part of the year up to late May. No specimens 
were collected between July and October. 


This species is close to S. (E.) elegans (Blote) and S. (£.) stali sp. n. on one hand 
and S. (£.) annulicornis (Signoret) on the other. From the former pair it can be 
separated by the short rostrum, the triangular median projection of the 2 seventh 
abdominal sternum and from the latter species by its uniformly thin and acutely 
pointed claspers. 

Villers doubted the validity of this species and suggested its synonymy with 
S. (E.) elegans (Blote). He remarked that the shape of the 9 seventh abdominal 
sternum was highly variable and this might have led Blote to describe this species. 
The form of 3 and 9 genitalia clearly demarcates these two species and the shape 
of the 2 seventh abdominal sternum has also been found to be fairly constant. 


Stenocoris (Erbula) elegans (Blote) comb. n. 
(Text-figs. 37-46) 
Leptocorixa (Evbula) elegans Blote, 1937 : 285 ; Villiers, 1955 : 1142, 1963 : 230 


Characterized by the claspers which are regularly curved, by the triangular median 
projection of 2 seventh abdominal sternum and by the form of § and 9 genitalia. 


6. Body length, 12-3-13-5 mm., width, 1-7-2-0 mm. Head length, 1-5-1-85 mm., width, 
1:25-1:45 mm. Antennae, segment I, 3:0-3-4 mm., II, 2-0-2:2 mm., III, 2:3-2-6 mm., IV, 
4°0-4:3 mm. Rostrum, segment I, 1-25-1-5 mm., II, 1-3-1:5 mm., III, 0-7-0-8 mm., IV, 
0-65-0-7mm. Pronotum, length, 2-o-2:5mm., width, 1-7-2-omm. Hind femora, 5-5-5-9 mm. 
Hind tibia, 5-8-6-:1 mm. Distance between ocelli, 0-4—0-5 mm., distance between an ocellus 
and an eye, 0:6—0-65 mm. 

Body short, comparatively slim, pale, with brownish black tinge. Head pale, slightly 
brownish in middle ; occipital suture “‘U’’-shaped in middle, side convex ; paraclypeae 
short, thick at bases, thin, rounded and divergent at apices. Antennae brownish black, some- 
times with underside of first, middle of second, articulations between first and second and 
between second and third, and basal portion of apical segment pale. Rostrum usually com- 
paratively long, extending at least slightly beyond second coxae. Lateral black line well 
represented, usually from genae to middle of propleuron sometimes faint to genae. Pronotal 
disc tinged brown, posterior angles with black dots. Legs pale, with apices of femora and 
bases of tibiae light brown, rarely with reddish tinge. Hemielytra with brownish black punc- 
tures. Abdomen with dorsum, except margins, red, venter pale ochraceous, sometimes tinged 
red. Posterior margin of seventh abdominal tergum usually at least slightly convex in middle ; 
pygophore with posterior margin medially notched. 

6 GENITALIA : Claspers long, reaching apex of pygophore, outer margin regularly curved and 
apices acutely pointed ; ventral thecal appendages as in Text-fig. 40 ; four left lateral con- 


THE LEPTOCORISINAE OF THE WORLD 21 


junctival appendages as in Text-fig. 40 ; frontal appendage bifurcated at apex, right branch 
shorter and thicker near apex ; four right lateral conjunctival appendages as in Text-fig. 40 
with spine-like appendage thicker in middle, thinner and pointed at apex ; dorsal appendage 
thick in middle and curved near base and apex. 

Q. Very similar to f in general appearance and measurements, usually longer (13-5—15-0 mm.) 
with a triangular median projection of 2? seventh abdominal sternum, this rounded at apex. 

Q GENITALIA: First gonocoxae with outer margin slightly sinuate, curved near apex ; 
intervalcular sacs absent ; spermatheca flask-shaped with thin and short neck as in Text- 


fig. 43. 


0-5 mm. 


Fics. 37-40. Stenocoris (Erbula) elegans, 3. 37, terminalia, dorsal view ; 38, claspers, 
outer and inner views ; 39, head, dorsal view ; 40, aedagus (inflated), ventral view. 


Type material. Holotype g, SIERRA LEONE: I1.x.1912 (J. J. Simpson), B.M. 
(N.H.) ; allotype 9, data as holotype, B.M.(N.H.) ; paratypes, SIERRA LEONE : 
Jahama, 3 4, 2 2 (J. J. Simpson), B.M.(N.H.). 


Other material. SENEGAL : 29,29; NIGERIA: 59,39; Conco: 259,159; 
TANGANYIKA : 3 Q, 2 9. 

I 9, 1 2 from SuMATRA, in Breddin’s collection, labelled ‘‘ Leptocorisa lepida 
Breddin det. Kiritshenko ” (Dtsch. Entom. Inst. Berlin) were also examined. It 
is felt that there has been some mistake in the locality data. 


The range of this species extends from Senegal in the west to Tanganyika in the 
east. This species is probably very common and adults have been collected through- 
out the year. 


This species is close to stali sp. n. and distinguenda (Blote). In stali the lateral 


22 I. AHMAD 


black line is faint, whereas in distinguenda (Blote) the rostrum is short and does not 
extend beyond the second coxae. Its triangular median projection with the round 
apex of the 9 seventh abdominal sternum and the claspers with regularly curved 
outer margins are its peculiarities. 


Alyy | 
yn 


/ 


Figs. 41-46. Stenocovis (Evbula) elegans, 2. 41, terminalia, ventral view ; 42, first 
gonocoxae, ventral view ; 43, Spermatheca; 44, first gonopophysis, ventral view ; 
45, Ovipositor, ventral view ; 46, same, dorsal view. 


THE LEPTOCORISINAE OF THE WORLD 


Stenocoris (Erbula) similis (Blote) comb. n. 
(Text-figs. 47-55) 
Leptocorixa (Evbula) similis Blote, 1937 : 283 


Fics. 47-55. Stenocorvis (Erbula) similis. 48-50, 3, 47, 51-55, 2. 47, spermatheca ; 
48, terminalia, dorsal view ; 49, claspers, outer and inner views ; 50, aedeagus; 51, 
first gonopophysis, ventral view ; 52, ovipositor, ventral view ; 53, first gonocoxae, 
ventral view ; 54, terminalia, ventral view ; 55, ovipositor, dorsal view. 


24 I. AHMAD 


Characterized by the faint lateral black line, which is usually represented by a 
lateral black dot on collar and by the form of § and 9 genitalia. 


6. Body length, 12-5-14-5 mm., width, 2-0-2: mm. Head length, 1-9-2:0 mm., width, 
I-35-1'5 mm. Antennae, segment I, 3:4-3:5 mm., II, 2:3-2-4 mm., III, 2:5-2-7 mm., IV, 
4°4-4:6 mm. Rostrum, segment I, 1-4-1-5 mm., II, 1-4-1-5 mm., III, 0-7-0-9 mm., IV, 
0:6-0-7 mm. Pronotum, length, 2-5-2-6mm., width, 2-0-2: mm. Hind femora, 5-4—5:6 mm. 
Hind tibia, 6-3-7-0 mm. ; distance between ocelli, 0-5 mm., distance between an ocellus and an 
eye, 0-7 mm. 

Body short, slightly comparatively robust, pale with a blackish tinge. Head with brownish 
markings ; occipital suture ‘“‘ U’’-shaped with arms curved backwards near sides of head ; 
paraclypeae short, thicker in middle. Antennae with brownish black tinge, apical segment red, 
pale at base. Rostrum short, approaching posterior limit of second coxae. Lateral black line 
very faintly represented, usually like a black dot on either side of collar. Pronotal disc with 
black latero-posterior spots. Legs pale, unicoloured, sometimes with blackish tinge. Hemiely- 
tra pale with black punctures on clavus, near membrane, on anal area and on cell. Abdomen 
pale, tinged red dorsally ; ventrally pale ochraceous. Posterior margin of seventh abdominal 
tergum truncated ; pygophore elongated, apex narrow and truncated. 

6 GENITALIA: Claspers long, distinctly reaching apex of pygophore, curved outwardly in 
middle and inwardly at apices ; ventral thecal appendages as in Text-fig. 50 ; four left lateral 
conjunctival appendages as in Text-fig. 50 ; frontal appendage bifid at apex, branches equal ; 
four right lateral conjunctival appendages with first spine-like appendage ; dorsal appendage 
thicker at base, inwardly curved and acutely pointed. 

9. Body length, 13-5-15:5 mm., width, 2:2-2:'4 mm. Head length, 2-0-2:1 mm., width, 
I-4-I-5 mm. Antennae, segment I, 3:3-3°5 mm., II, 2:1-2:3 mm., III, 2-4-2-6 mm., IV, 
4:0-4°5 mm. Rostrum, segment I, 1-4-1-6 mm., II, 1-4-1-5 mm., III, 0-7-0-9 mm., IV, 
o-6-0'8mm. Pronotum, length, 2-7-2:8 mm., width, 2:2-2-4mm. Hind femora, 5-4—5:6 mm. 
Hind tibia, 5-9-6: mm. Distance between ocelli, 0-5-o-7 mm. ; distance between an ocellus 
and an eye, 0-6—0-8 mm. 

Usually longer than ¢ and slightly darker in colour, seventh abdominal sternum with a 
rounded median projection, which has a distinct median notch. 

Q GENITALIA : First gonocoxae triangular with rounded apices ; intervalvular sacs absent ; 
spermatheca thicker at base with a thin long and rounded neck as in Text-fig. 47. 


Type material. Holotype g, ANGOLA: Quirimbo, v.1934 (K. Jordan), B.M. 
(N.H.) ; paratypes, 15 5, 10 9, with data as holotype, B.M.(N.H.). 


Other material. Conco: 344,159; TANGANYIKA: I<. 


This species is common in the Congo and its range extends south west into Angola. 
The adults have been collected from February until May and also in December. 


This species is closest to S. (E.) elegans and S. (E.) stali sp. n., but can be separated 
from them by the character of the lateral black line, which is in this species commonly 
reduced to a black dot on either side of the collar. The rounded and medially notched 
median projection of the 9 seventh abdominal sternum and the elongated pygophore 
with a truncated posterior margin are its peculiarities. 


Stenocoris (Erbula) southwoodi sp. n. 
(Text-figs. 56-64) 


Leptocorisa apicalis Westwood ; Stal, 1866 : 88, 1873 : 86 
Leptocorixa apicalis Westwood ; Blote, 1937 : 282 ; Villiers, 1955 : 1141, 1963 : 229 


THE LEPTOCORISINAE OF THE WORLD 25 


Characterized by very short claspers, which are thick and curved in the middle and 
do not reach the posterior margin of pygophore, by posterior median round projec- 
tion of 2 seventh abdominal sternum and by the form of 3 and 9 genitalia. 


3g. Body length, 12-2~-13-5 mm., width, 1-9-2: mm. Head length, 1-7-1-9 mm., width, 
I-4-I1°5 mm. Antennae, segment I, 3:2-3-6 mm., II, 2-0-2:2 mm., III, 2-4-2-6 mm., IV, 
4°4-4:7 mm. Rostrum, segment I, 1-25-1-5 mm., II, 1-4-1-5 mm., III, 0-7-0-9 mm., IV, 
0-5-0-7 mm. Pronotum, length 2-2-2-4 mm., width, 1-9-2-1 mm. Hind femora, 5-0—-5-5 mm. 
Hind tibia, 5-5-5-8 mm. Distance between ocelli, 0-5 mm., distance between an ocellus and an 
eye, 0°75 mm. 

Body short, darker in colour. Head pale with blackish tinge ; occipital suture “ V ’’- 
shaped in middle with arms curving backwards towards side of head ; paraclypeae short, 
slightly thicker in middle. Antennae brownish black, with underside of basal segment, articu- 
lations between first and second and between second and third and basal portion of apical seg- 
ment, pale. Rostrum short, reaching apex of second coxae. Lateral black line extending 
from genae up to lateral side of collar. Pronotal disc pale with brownish black tinge, posterior 
angles brownish black. Legs pale, usually tinged black. Hemielytra, except costal margin 
and membrane, covered with black punctures. Abdomen with dorsum pale tinged red, venter 
pale ochraceous with reddish tinge anteriorly. Posterior margin of seventh abdominal tergum 
truncated, only slightly convex in middle ; pygophore rounded with a pointed apex. 

6 GENITALIA : Claspers very short, never reaching posterior margin of pygophore, thicker 
and curved in middle ; ventral thecal appendages as in Text-fig. 58, four left lateral conjunc- 
tival appendages as in Text-fig. 58, frontal appendage bifurcated from middle, branches equal ; 
four right lateral conjunctival appendages as in Text-fig. 58, with spine-like appendage reduced 
to a very small bristle-like appendage ; dorsal appendage thicker at base and strongly 
curved. 

9%. Body length, 13:0-14-5 mm., width, 1-9-2: mm. Head length, 1-8-1-95 mm., width, 
1-4-1‘'6 mm. Antennae, segment I, 3-0-3:6 mm., II, 2-:0-2:3 mm., III, 2-4-2-6 mm., IV, 


0-5 mm. 


Fics. 56-58. Stenocoris (Erbula) southwoodi, 3. 56, terminalia, dorsal view ; 57, 
claspers, outer and inner views ; 58, aedeagus (inflated), ventral view. 


26 I. AHMAD 


4°5-4:7 mm. Rostrum, segment I, 1-4-1-6 mm., II, 1-4-1:6 mm., III, 0-7-0-9 mm., IV, 
o-6-0'°8 mm. Pronotum, length, 2:3-2:5 mm., width, 1-9-2: mm. Hind femora, 5-6-6-0 mm. 
Hind tibia, 6-o-6:1 mm. Distance between ocelli, 0-5 mm., distance between an ocellus and an 
eye, 0-75 mm. 

Very similar to g§ in general appearance, slightly darker and longer ; posterior margin of 
seventh abdominal sternum with a round, median, faintly striated projection. 


Fics. 59-64. Stenocoris (Evbula) southwoodi, 9. 59, terminalia, ventral view ; 60, first 
gonopophysis, ventral view ;_ 61, first gonocoxae, ventral view ; 62, ovipositor, ventral 
view ; 63, same, dorsal view ; 64, spermatheca. 


THE LEPTOCORISINAE OF THE WORLD 27 


© GENITALIA : First gonocoxae elongated and narrow, apices rounded ; intervalvular sacs 
absent ; spermatheca ovate without a distinct neck, as in Text-fig. 64. 


Holotype 3, SIERRA LEONE: Sembehun, at light, 14.11.1924 (E. Hargreaves), 
labelled ‘“‘ Leptocorixa apicalis Westwood, det. H. C. Blote ’’, B.M.(N.H.). 


Paratypes. NIGERIA: Sapolar, 1 g, 10.v.1955 (E. A. Duffy), Lagos, 6 3 (G. 
Strachan), B.M.(N.H.) ; S. NIGERIA : 1g, 1912 (A. D. Peacock), with label” Lepto- 
corixa apicalis Westwood, det. H. C. Blote’”’, B.M.(N.H.) ; SIERRA LEONE: I 9 
(Afzel), Stal’s collection, Riksmus. Stockholm ; GHANA: Ashanti Dist., 1 g, I Q, 
7.V.1913 (A. E. Evans), B.M.(N.H.) ; PRINCIPE ISLAND: I Q, 1.1.1933 (W. H. T. 
Tams), B.M.(N.H.) ; SAo THoME IsLAND : I 4, Ig1g—I921, B.M.(N.H.) ; Conco : 
Ituru, i g (Gyld), Riksmus. Stockholm, various localities, 250 3, 345 Q, ili—vi. 
vili—xi. (various collectors), Mus. R. Afr. centr. ; SOMALIA : Basso Giula Portrizi, 
I 9, vii.1934, B.M.(N.H.) ; UGAnpA: Kawanda, light trap in coffee plantation, 
6-12.1, 4-7.li, 8-11.xi.1959 (J. Bowden), Ahmad collection, Ruwenzori Range, 
Ibanda, 4,700 ft., I g, 4-6.vii.1952 (D. S. Fletcher), Bwamba Valley, I 3, vii.1945 
(van Someren), Kampala, Jinja Road, 4,000 ft., Mabira Forest, 3,500—3,800 ft., 
Chagwe, 1 9, 1 4, vil.1g1r (S. A. Neave), Nymgasair Valley, 6,400 ft., between 
Mitiana and Entebbe, 3,800 ft., Tero Forest, 5 3, 5 9, ix.I9II, 12.1V.1913, 2.Vi.1930, 
Xli.1934, 1.1935 (various collectors), B.M.(N.H.) ; TANGANYIKA : Ilonga, nr. Kilosa, 
vii.1962 (Robertson), B.M.(N.H.) ; ZANZIBAR : TI Q, 1.1i.1925 (H. J. Shell), B.M.(N.H.) ; 
?SUMATRA : 2 4, in series of Leptocorisa lepida Breddin, 1909, with label “ L. lepida 
det. Kiritshenko”’ (probably an incorrect locality label), Dtsch. Entom. Inst., 
Berlin. 


This is the most widely distributed species of Erbula. Leaving aside the doubtful 
record from Sumatra, its range extends from Sao Thomé Island and Principe Island 
in the west to Zanibar in the east and appears to comprise the whole of the African 
equatorial belt. The adults are very common in late May and early June, but they 
have been collected throughout the year. 


This species is close to S. (£.) annulicornis (Signoret) in general appearance and 
colour but comparatively isolated and can be easily separated from all other species 
of the subgenus Erbula by having very short, medially thick and curved claspers in 
the g, which never reach the apex of the pygophore. In the 9, the round median 
projection of the seventh abdominal sternum separates it from all other species of 
this group. 

An examination of the material that was seen by Stal and Blote (see above) has 
shown that Stal misidentified this species as L. apicalis Westwood, and Blote 
correctly followed Stal’s interpretation. However their specimens are not Stenocoris 
apicalis (Westwood) (see page 38) and thus the present species, which was the type 
of Evbula Stal, is hitherto undescribed. This species has also been recorded by 
Southwood (1961) in light trap catches at Kawanda, Uganda. I have pleasure in 
naming this species in honour of Dr. T. R. E. Southwood, who suggested the present 
problem and has supervised my studies on it. 


28 I. AHMAD 


Stenocoris (Erbula) stali sp. n. 
(Text-figs. 65-74) 


Characterized by the shape of the claspers, by the median posterior margin of the 
2 seventh abdominal sternum and by the form of Jf and 9 genitalia. 


3. Body length, 12-4-14-0 mm., width, 1:66-2:26 mm. Head length, 1-6—-1:86 mm., width, 
I:2-1:'25 mm. Antennae, segment I, 3-1-3:35 mm., II, 2-0-2:26 mm., III, 2:26-2-4 mm., IV, 
4:0-4:13 mm. Rostrum, segment I, 1-2-1-3 mm., II, 1-2-1-33 mm., III, 0-65-0-7 mm., IV, 
0-65-0'7 mm. Pronotum, length, 2-0-2:4 mm., width, 1:66-2:26 mm. Hind femora, 
5:2-5°6 mm. Hind tibia, 5-5-6:0 mm. Distance between ocelli, 0-4-0-46 mm. Distance 
between an ocellus and an eye, 0-65—0-7 mm. 

Body comparatively long, slim, pale, tinged with brown. Head medially and posteriorly 
brown ; occipital suture medially concave with sides curved ; paraclypeae short, usually flat, 
of uniform thickness. Antennae usually pale, tinged brown. Rostrum long, usually extending 
at least slightly beyond apices of second coxae. Lateral black line faint but extending from 
genae to anterior third of propleuron. Pronotal disc pale unicoloured with black spot on 
posterior angles. Legs pale, with apices of femora and bases of tibiae tinged with dark brown. 
Hemielytra with brown punctures on clavus and on anal area. Abdomen with dorsum, except 
connexiva, palish red, venter pale ochraceous. Posterior margin of seventh adbominal tergum 
truncated, pygophore elongated with bilobed apex. 

$ GENITALIA : Claspers curved only in middle, narrow towards apices, acutely pointed ; 
ventral thecal appendages as in Text-fig. 68 ; four left lateral conjunctival appendages as in 
Text-fig. 68 ; frontal appendage thinner in middle, bifid at apex, left branch longer and thicker ; 
four right lateral conjunctival appendages as in Text-fig. 68 ; dorsal appendage strongly 
inwardly curved with pointed apex. 


Fics. 65-68. Stenocoris (Evbula) stali, 3. 65, terminalia, dorsal view ; 66, claspers, 
inner and outer views ; 67, head, dorsal view ; 68, aedeagus, ventral view. 


THE LEPTOCORISINAE OF THE WORLD 29 
9. Body length, 15-0-15-8 mm., width, 2-0-2: mm. Head length, 1-8-1-9 mm., width, 
I-2-1'4 mm. Antennae, segment I, 3-0-3:35 mm., II, 1-9-2:1 mm., III, 2-2-2:35 mm., IV, 
4°0-4:'2 mm. Rostrum, segment I, 1-33 mm., II, 1-33 mm., III, 0-7 mm., IV, 0-7 mm. _Pro- 
notum, length, 2:3-2-4 mm., width, 2-0-2: mm. Hind femora, 5-8mm. Hind tibia, 6-0 mm. 
Distance between ocelli, 0-43 mm. ; distance between an ocellus and an eye, 0-68 mm. 
Very similar to $ in general appearance but usually distinctly longer ; posterior margin of 


Fics. 69-74. Stenocoris (Erbula) stali, 2. 69, first gonocoxa, ventral view ; 70, termina- 
lia, ventral view ; 71, first gonophysis, ventral view ; 72, ovipositor, ventral view ; 


73, same, dorsal view ; 74, spermatheca. 


30 I. AHMAD 


seventh abdominal sternum with a median rectangular projection as in Text-fig. 70. 
Q GENITALIA : First gonocoxae broad, apex rounded ; intervalvular sacs absent ; sper- 
matheca flask-shaped with a long curved neck as in Text-fig. 74. 


Holotype 3, Conco : Congo da lemba, 1.ii.1913 (R. Mayné), Mus. R. Afr. centr. 


Paratypes. 2, 29, with data as holotype, Mus. R. Afr. centr. ; Conco: Kat- 
anga, Kando, 2 9, 25.iii.1931 (G. F. de Witte), Sokole, Eala, Sankuru, and Ubangi, 
3.6, 2 9, 3I-XI1.IQII, X.1923, 5.1V.193I, V.1937, 1952 (various collectors), Mobwasa, 
I 9, 30.xi.1912 (R. Mayné), Mus. R. Afr. centr. ; SIERRA LEONE : Freetown, I 4, 
13.vii.1899 (E. E. Austen), B.M.(N.H.) ; Niceria ; Old Calabar, 1 3 (Schouteden), 
Mus. R. Afr. centr. 


The range of this species extends in the west into Sierra Leone and in the east into 
Congo. Adults have been collected in the early part of the year until May, but they 
have not been collected in June or July. 


This species is very close to elegans (Blote) in general appearance but can easily be 
separated by the form of the claspers of the g. In 9, the median projection of the 
seventh abdominal sternum separates it from all the species of this subgenus. 


STENOCORIS (STENOCORIS) Burmeister 
Type-species : Cimex tipuloides Degeer. 


Body robust. Head elongated, usually distinctly longer than broad ; paraclypeae usually 
short, straight and rounded at apices ; ocelli close together but distance between an ocellus and 
an eye about one-third as long again as interocellar distance. Antennae with apical segment 
pale only at base, second usually distinctly shorter than third. Rostrum long, usually reaching 
second coxae. Lateral black line usually present, never extending beyond middle of propleuron. 
Pronotum sloping down towards head with posterior angles usually brownish black. Abdomen 
with dorsum usually palish red ; venter usually pale unicoloured. 

In 3, posterior margin of seventh abdominal tergum usually truncate ; pygophore usually 
rounded ; claspers crossed in socket (Text-fig. 75), curved in middle with apices pointed; 
aedeagus with only ventral pair of thecal appendages ; dorsal pair wanting ; frontal conjunc- 
tival appendages paired, elongated and sclerotized ; membranous appendage with usually 
paired dorsal appendages, terminal appendage absent. 

In 9, posterior margin of seventh abdominal sternum medially bilobed ; first gonocoxae 
large, often mainly hidden under seventh abdominal sternum ; intervalvular sacs absent ; 
spermatheca flask-shaped with a long coiled tube. 


This subgenus can be distinguished from other subgenera in the § by the character 
of claspers which are crossed in a socket and in the 9 by the bilobed posterior margin 
of seventh abdominal sternum. 


KEY TO SPECIES 


t Rostrum reaching anterior base of hind coxae ; apices of femora tinged with red 
(frontal conjunctival appendages in g aedeagus very short (Text-fig. 167)) ; 
spermatheca cylindrical (Text-fig. 174) . F ; : : v-nigrum (p. 57 

— Rostrum never reaching hind coxae ; apices of femora never tinged with red : 2 

2 Posterior angles of pygophore produced backward (Text-fig. 148) ; in 9, posterior 
margin of seventh abdominal sternum deeply indented in middle (Text-fig. 93) . 3 


THE LEPTOCORISINAE OF THE WORLD 31 


Posterior angles never produced backward ; in 9, posterior margin of seventh 
abdominal sternum only slightly indented Lr wea 124) or concave in form of a 
5 hanes : - 7 
Pygophore large with peetiealad margin eee? more ov half Ww widths ‘ai panties 
margin of j seventh abdominal tergum (Text-fig. 88); processes of posterior margin 
of 9 seventh abdominal sternum large and very prominent, sometimes conical or 
“U-shaped (Text-fig. 93) . p 4 
Pygophore small, with narrow posterior states, a two- fifths width of posterior 
margin of seventh abdominal tergum (Text-fig. 107) ; processes of posterior 
margin of 2 seventh abdominal sternum small, never triangular, usually posterior 
margin truncated (Text-fig. 111) . : : ; : : : 5 
Rostrum long, extending beyond second coxae ; lateral black line, prominent on 
genae to middle of propleuron ; (claspers narrow in middle (Text-figs. 148 and 
149), second spine-like oe in aedeagus absent ; spermatheca flat (Text- 
fig. 156))  . .  sordida (p. 52) 
Rostrum short, reaching second coxae, lateral black line faint, usually absent on 
genae ; (claspers broad in middle (Text-figs. 88 and 89) ; second spine-like 
appendage in aedeagus present; spermatheca elongated (Text-fig. 95)) apicalis (p. 36) 
Head elongated, about half again longer than broad (Text-fig. 78) with paraclypeae 
long and cylindrical, about 2/5 length of entire head ; lateral black line usually 
faint or absent on genae (aedeagus with first dorsal conjunctival appendage very 
small and curved ; spermatheca with smoothly curved neck (Text-fig. 80)) 
africana (p. 32) 
Head broad, usually only slightly longer than broad ; paraclypeae short, usually 
one-quarter length of entire head ; lateral black line ames on genae and 
extending to anterior of propleuron ; : < : : 6 
Rostrum long, extending well beyond apices of second c coxae ; species comparatively 
robust, at least 14 mm. in length (dorsal thecal appendage present, rod-like in 
appearance (Text-fig. 112) ; right lateral conjunctival appendages three in 
number, fourth left lateral conjunctival appendage club-shaped ; in Q, first 
gonocoxae with truncate posterior margin). - ; claviformis (p. 41) 
Rostrum short, only just extending beyond apices of second coxae ; species com- 
paratively small, length about 13:5 mm. (dorsal thecal appendage absent ; right 
lateral conjunctival appendages four in number, fourth bifurcated at apex, 
fourth left lateral conjunctival appendage rod-like, elongate) . . ethiopis (p. 43) 
Rostrum long, extending well beyond apices of second coxae ; pygophore rounded 
with posterior margin medially distinctly emarginate (Text-fig. 119) ; a large 
basal portion of first gonocoxae concealed (Text-fig. 124). (Frontal conjunctival 
appendages very long nearly reaching anterior base of basal plate ; spermatheca 
with a narrow and a smoothly curved neck) . . : maculosa (p. 44) 
Rostrum short, never extending well beyond apices of second coxae, usually reaching 
second coxae ; pygophore usually elongated (Text-fig. 128), rounded only in 
American species ; a very small basal portion of first gonocoxae concealed (Text- 
He t32)) : ; ° : : : : : - 8 
Pygophore elongated (Text- fig. 128) ; first gonocoxae comparatively narrow at 
base, so that only a small basal portion of eighth paratergite concealed externally. 


African species. : é ; d : : : : ; 9 
Pygophore rounded (Text. -fig. 98), first gonocoxae broad at base so that a large basal 
portion of eighth paratergite concealed externally (Text-fig. 102). American species 10 


Lateral black line usually represented extending from either side of antenniferous 
tubercles to anterior of collar ; black spots on posterior angles of pronotum 
usually present, paraclypeae comparatively long and thinner at apices (frontal 
conjunctival appendages short and thick ; spermatheca elongated, with smoothly 


32 


Io 


za 6 


I. AHMAD 


curved neck) c - phthisica (p. 49) 
Lateral black line usually oe cemed by a lacks dot on Baier side of collar ; black 

spots on posterior angles of pronotum absent ; paraclypeae short and uniformly 

thick (frontal conjunctival appendages long and thin ; spermatheca almost 

pelican-shaped with a thick and coiled tube) . : . pallida (p. 46) 
Pale unicoloured species with thin paraclypeae, spots on ees apie: of pronotum 

absent, spermatheca round . . : : braziliensis (p. 48) 
Palish brown species with uniformly thick apeechovere ; spots on posterior angles 

of pronotum always present ; spermatheca flask-shaped . : : : : II 
Rostrum at least slightly extending beyond apices of second coxae ; lateral black 

line faintly represented ; in g, posterior pis of pygophore truncated (Text- 

fig. 85) : . : americana (p. 34) 
Rostrum reaching ee aera 4 ceagud coxae ; ed black line strongly 

represented ; in 3, posterior margin of pygophore medially emarginate (Text- 

fig. 158) é : : : c : : ¢ : ‘ tipuloides (p. 55) 


Stenocoris (Stenocoris) africana sp. n. 
(Text-figs. 75-84) 


Characterized by uniformly thin and long paraclypeae (Text-fig. 78), which are 


about two-fifths of entire head and by the form of g and @ genitalia. 


Fics. 75-78. Stenocoris africana, 3. 75, terminalia, dorsal view ; 76, claspers, inner 


and outer views ; 77, aedeagus (inflated), ventral view ; 78, head, dorsal view. 


THE LEPTOCORISINAE OF THE WORLD 33 


3g. Body length, 12-4-13:6 mm., width, 1-8-1-9 mm. Head length, 1-6-1-7 mm., width, 
1-2-1'3mm. Antennae, segment I, 3:2~-3:7 mm., only paraclypeae 0-66 mm., II, 2-0-2-4 mm., 
III, 2:1-2-7 mm., IV, 3-6-4: mm. Rostrum, segment I, 1-2-1-3 mm., II, 1-3—1-4 mm., III, 


Fics. 79-84. Stenocoris africana, 2. 79, first gonocoxa, ventral view ; 80, spermatheca ; 
81, first gonopophysis, ventral view ; 82, terminalia, ventral view ; 83, ovipositor, 
ventral view ; 84, same, dorsal view. 


34 I. AHMAD 


0-6-0:7 mm., IV, 0-7-0-8 mm. Pronotum length, 2-1-2:2 mm., width, 1-8-1-9g mm. Hind 
femora, 5:3-6:3 mm. Hind tibia, 5-6-6-5 mm. Distance between ocelli, 0-4 mm. ; distance 
between an ocellus and an eye, 0-7 mm. 

Body comparatively slim, palish brown. Head elongated, usually approximately half as long 
again as broad; occipital suture “‘U’’-shaped in middle, sides convex ; paraclypeae long, uniformly 
thin and round at apices, two-fifths of entire head. Antennae brownish black with underside 
of basal, usually basal halves of second and basal portions of apical segments pale. Rostrum 
short, usually reaching anterior margin of second coxae. Lateral black line well represented 
from lateral sides of antenniferous tubercles on to collars. Pronotal disc pale, posteriorly with 
brownish tinge, posterior angles with brownish black spots. Legs pale, with apices of femora 
and tibiae tinged with red. Costal and anal margins of hemielytra black, with black punctures 
on claval and anal areas. Abdomen dorsally, except connexival areas, tinged with red ; 
venter pale ochraceous. Posterior margin of seventh abdominal tergum truncated, slightly 
medially concave ; pygophore small with small posterior processes, posterior margin narrow, 
about one-third of seventh abdominal tergum. 

6 GENITALIA : Claspers small, medially thick and rounded at apices (Text-fig. 76) ; dorsal 
thecal appendages absent, ventral pair thin, medially curved and truncated at apices ; four 
left lateral conjunctival appendages as in Text-fig. 77 ; frontal appendages paired, thin, short 
and medially curved ; four right lateral conjunctival appendages as in Text-fig. 77, dorsal 
appendages as in Text-fig. 77. 

©. Body length, 15-0-15-5 mm., width, 2:2-2:-4 mm. Head length, 1-:7-1-9 mm., width, 
I-3-1:4 mm. Antennae, segment I, 3-3-3-6 mm., (only paraclypeae o-7-0-8 mm.), II, 2-3— 
2-4 mm., III, 2-4-2:5 mm., IV, 3-1-3:3 mm. Rostrum, segment I, 1:45-1:55 mm., II, 1-56— 
1-6 mm., III, 0-7-0-73 mm., IV, 0-8-0-83 mm. Pronotum, length, 2:4-2:6 mm., width, 
2:2-2'4 mm. Hind femora, 6-0-6:-4 mm. Hind tibia missing. Distance between ocelli, 
0-4 mm., distance between an ocellus and an eye, 0-7 mm. 

Very similar to g in general appearance ; posterior margin of seventh abdominal sternum 
indented in middle, forming broad and truncated lobes on either side (Text-fig. 82). 

Q GENITALIA: First gonocoxae largely concealed by seventh abdominal sternum, outer 
margin irregular with apices truncated ; intervalvular sacs absent ; spermatheca elongated, 
tapering towards apex with smoothly curved neck and thick coiled tube. 


Holotype 3, Conco: Kivu, Terr. Mwenna Kitutu, 650 m., (lumiére), vi.1958 
(N. Leleup), Mus. R. Afr. centr. 


Paratypes. Conco: Uelé, Gangalena Bodio, I 3, 14.v.1936 (L. Lippens), 
Nyangwe, I5am, 2 4, 30.11, iv.1918 (R. Mayné), Kivu-mulungo, Tshibinda, 1 3, 2 9, 
5.1V.1937, 22.1v.1938 (Hendrickx), Elizabethville, Kasongo, Kibambo, 4 g, vi.1930, 
ili.1932, Vlil.1959 (various collectors), Mus. R. Afr. centr.; SIERRA LEONE : Freetown, 
13.vii.1899 (E. E. Austen), B.M.(N.H.) ; ApyssiniA : Mulatos, Haxor, I 3, 22-25. 
x.1920, A.M.N.H. 


This species is very close to S. phtlisica (Gerstacker) and S. maculosa (Bléte), but 
can easily be separated from the latter by short rostrum and from the former by 
short lateral black line, never extending beyond the collar. 


Stenocoris (Stenocoris) americana sp. n. 
(Text-figs. 85-87) 


Characterized by pale body, long rostrum, reaching beyond the second coxae and 
by the form of 3 and 9 genitalia. 


THE LEPTOCORISINAE OF THE WORLD 35 


6. Body length, 13-0-14:0 mm., width, 2-0-2: mm. Head length, 1-8-1-9 mm., width, 
I-3-I'4 mm. Antennae, segment I, 3:0-3-:1 mm., II, 2-2-2-3 mm., III, 2-3 mm., IV, 3:3 mm. 
Rostrum, segment I, 1-3 mm., II, 1-4 mm., III, 0-7 mm., IV, 0-8 mm. Pronotum, length, 
2:2-2:4 mm., width, 2-0-2: mm. Hind femora, 5-9-6-:0 mm. Hind tibia, 6-1-6-3 mm. 
Distance between ocelli, 0-4 mm., distance between an ocellus and an eye, 0-66 mm. 

Body pale, comparatively slim. Head slightly tapering behind occipital suture, at least a 
third as long again as broad, pale with a brownish tinge ; occipital suture medially concave, 
arms bending forward ; paraclypeae short, about one-third of head ; uniformly thin and 
pointed at apices. Antennae with underside of basal segment, three-quarters of second and 
third and basal portions of apical segments pale. Rostrum comparatively long, at least 
extending slightly beyond apices of second coxae. Lateral black line faint, extending from 
genae on to anterior third of propleuron. Pronotal disc pale, tinged with light brown, black 
spots on posterior angles faint. Legs pale, apices of femur and tibia tinged with light brown. 
Hemielytra brownish black. Abdomen with dorsum, excluding connexiva, red, venter pale 
ochraceous. Posterior margin of seventh abdominal tergum truncate ; pygophore rounded 
with sides curved near apex, posterior margin truncate. 

6 GENITALIA : Claspers strongly curved in middle and at apices, acutely pointed ; dorsal 
thecal appendages absent, ventral pair long thicker in middle and truncated at apices ; four 
left lateral conjunctival appendages as in Text-fig. 87 ; frontal appendages paired, longest of 
all, strongly inwardly curved and acutely pointed at apices, four right lateral conjunctival 
appendages as in Text-fig. 87 ; dorsal appendages as in Text-fig. 87. 

No @ available for study. 


Holotype 3, Ecuapor : Puna, Riksmus. Stockholm. 


Paratypes, I 3, with same data as holotype. Riksmus. Stockholm; VENEZUELA: 
Dapure San Fernada de Apure, Mayeul, 1 g, 1923, Mus. Hist. nat. Paris. 


0-5 mm. 


Fics. 85-87. Stenocoris americana, §. 85, terminalia, dorsal view ; 86, claspers, 
inner and outer views ; 87, aedeagus (inflated), ventral view. 


36 I. AHMAD 


This species is very close to S. tipuloides (Degeer) and S. braziliensis sp. n. but can 
be separated by its comparatively long rostrum, which extends beyond apices of 
second coxae and by faint brownish black spots on the posterior angles of pronotum. 


Stenocoris (Stenocoris) apicalis (Westwood) comb. n. 
(Text-figs. 88-96) 
Leptocorisa apicalis Westwood, 1842 : 18 


Characterized by medially strongly indented 9 seventh abdominal sternum, giving 
triangular processes on either side, by large pygophore with strong posterior pro- 
cesses and by g and & genitalia. 


6. Body length, 12-8-13-5 mm., width, 2:1-2-3 mm. Head length, 1-6-1-9 mm., width, 
I1-3-1'46 mm. Antennae, segment I, 3-6—3:86 mm., II, 2-3-2-6 mm., III, 2-35-2-:7 mm., IV, 
4:0-4:2 mm. Rostrum, segment I, 1-4-1-5 mm., II, 1-4-1-5 mm., III, 0-7-0-73 mm., IV, 
0-7-0:73 mm. Pronotum, length, 2:-3—2-5 mm., width, 2-1-2-3mm. Hind femora, 5-6—6-0mm. 
Hind tibia, 6-0-6-4 mm. Distance between ocelli, 0-4—o-5 mm., distance between an ocellus 
and an eye, 0-7—-0:75 mm. 


0:5 mm. 


0:5 mm. 


Fics. 88-90. Stenocoris apicalis, 3. 88, terminalia, dorsal view; 89, claspers, outer 
and inner views ; 90, aedeagus, ventral view. 


THE LEPTOCORISINAE OF THE WORLD 37 


Body comparatively robust, pale, tinged with black. Head slightly longer than broad ; 
occipital suture medially “‘ V ’’-shaped, following convexly towards sides ; paraclypeae short, 
uniformly thick and rounded at apices. Antennae with underside of basal segment and basal 
portion of apical segments pale. Rostrum comparatively long, distinctly extending beyond 


Fics. 91-96. Stenocoris apicalis, 2. 91, ovipositor, ventral view; 92, first gonocoxa, 
ventral view ; 93, terminalia, ventral view ; 94, first gonopophysis, ventral view ; 
95, Spermatheca ; 96, ovipositor, dorsal view. 


38 I. AHMAD 


apices of second coxae. Lateral black line faint, extending from genae on to anterior of pro- 
pleuron. Pronotal disc pale, brownish black spots on posterior angles present. Legs pale, 
tibiae and tarsi tinged with black. Anal margins and M-Cu cell of hemielytra black. Abdomen 
with dorsum, excluding connexiva, red, venter pale ochraceous. Posterior margin of seventh 
abdominal tergum truncated, pygophore large, posterior processes prominent, acute ; posterior 
margin broad, more than half the width of posterior margin of seventh abdominal tergum. 

6 GENITALIA: Claspers broad in middle, not strongly curved at apices; dorsal thecal 
appendages absent, ventral pair thicker at base with round apices ; four left lateral conjunc- 
tival appendages as in Text-fig. 90 ; frontal conjunctival appendages paired, left long, thin and 
curved ; four right lateral conjunctival appendages as in Text-fig. 90 ; dorsal appendages as 
in Text-fig. 90, with second spine-like appendage at base. 

2. Body length, 13-3-14-6 mm., width, 2:2-2-3 mm. Head length, 1-6-1-9 mm., width, 
I-4-I1'5 mm. Antennae, segment I, 3-3-3-6 mm., II, 2-0-2-3 mm., III, 2-3-2:5 mm., IV, 
3°9-4:1 mm. Rostrum, segment I, I-4-1-5 mm., II, 1-4-1:5 mm., III, 0-7-0-8 mm., IV, 
0:7-0°8 mm. Pronotum, length, 2-3—-2-4 mm., width, 2-2-2:3 mm. Hind femora, 5-2-6-:0 mm. 
Hind tibia, 5-9-6:3 mm. ; distance between ocelli, 0-4-0-5 mm. ; distance between an ocellus 
and an eye, 0-7-0-8 mm. 

Very similar to gf in general appearance ; posterior margin of seventh abdominal sternum 
strongly indented medially, giving triangular processes on either side. 

@ GENITALIA : First gonocoxae large, concealing a large portion of ninth paratergite, broader 
at base and acutely pointed at apices ; intervalvular sacs absent ; spermatheca of thick 
crescent shape with a long and coiled tube. 


Type material. Holotype 2, without data but with type labels, Hope Dept., 
Univ. Mus. Oxford. 


Other material. CoNGo: 214, 3092; GHANA: 2¢; CAMEROON : 154, 13 9. 


This species seems to be restricted to the West African region. Adults have been 
collected from February until October. 


This species is the closest to S. sordida (Bléte) but can be separated by triangular 
processes of 9 seventh abdominal sternum and by broad posterior margin of the 
pygophore, which is always more than half the width of seventh abdominal tergum. 

Stal, by misidentifying a hitherto unnamed species (now described as S. (Evbula) 
southwoodi sp. n.) as Stenocoris apicalis Westwood, established a subgenus (Evbula) to 
contain it. 

The holotype 2 (see above) is a typical Stenocoris s. str. in having bilobed 
posterior margin of seventh abdominal sternum and in lacking short tube of sperma- 
theca, which is curled round its neck in Evbula. 


Stenocoris (Stenocoris) braziliensis sp. n. 
(Text-figs. 97-105) 


Characterized by parallel-sided head, pale unicoloured body and by the form of ¢ 
and 9 genitalia. 


6. Body length, 14:0-15:2 mm., width, 2:0-2:3 mm. Head length, 1-8-1-9 mm., width, 
I-3-I'4 mm. Antennae, segment I, 3:0-3-6 mm., (only paraclypeae, 0-66-0-7 mm.), II, 
2:1-2:3 mm., III, 2:3-2:-5 mm., IV, 3-9-3: mm. Rostrum, segment I, 1-:2-1-4 mm., II, 
I:2-1-'4 mm., III, 0-7-0:75 mm., IV, 0-8-0-85 mm. Pronotum, length, 2-4-2-5 mm., width, 
2:0-2:3 mm. Hind femora, 5-6-6:0 mm. Hind tibia, 6-:0-6-5 mm. Distance between ocelli, 
0-5 mm., distance between an ocellus and an eye, 0-6 mm. 


THE LEPTOCORISINAE OF THE WORLD 39 


Body comparatively robust, pale unicoloured. Head parallel-sided, long, at least a third 
again longer than broad ; occipital suture “‘ V ’’-shaped with arms curving outwards towards 
sides of head ; paraclypeae long, more than a third of head-length, uniformly thin and rounded 
at apices. Antennae pale, tinged black, with underside of basal segment, middle of second 
and third and basal portions of apical segments pale. Rostrum short, hardly reaching apices of 
second coxae. Lateral black line usually very faint, extending from sides of antenniferous 
tubercles on to anterior of propleuron. Pronotal disc, posteriorly with greenish tinge, posterior 
angles without black spots. Legs with bases of femora tinged with green and apices brownish. 
Anal margin and M-Cuw cell of hemielytra brownish black. Abdomen with dorsum pale, 
unicolourous, venter pale, sometimes brownish. Posterior margin of seventh abdominal 
tergum truncated ; pygophore rounded, tapering posteriorly with narrow, medially concave 
posterior margin. 

6 GENITALIA : Claspers comparatively long, thicker in middle, curved and pointed at apices ; 
dorsal thecal appendages absent, ventral pair long, thicker at bases and rounded at apices ; 
four left lateral conjunctival appendages as in Text-fig. 99 ; frontal conjunctival appendages 
paired, of equal sizes, long, thick at bases and acutely pointed at apices ; four right lateral 
conjunctival appendages as in Text-fig. 99 ; dorsal appendages as in Text-fig. 99. 


0-5 mm. 


Fics. 97-100. Stenocoris braziliensis. 97-99, 5, 100, 2. 97, claspers, inner and outer 
views ; 98, terminalia, dorsal view ; 99, aedeagus (inflated), ventral view ; 100, first 
gonocoxa, ventral view. 


40 I. AHMAD 


®. Body length, 14:5-15-5 mm., width, 2:2-2:3 mm. Head length, 1-9-2-0 mm., width, 
I:3-I1-4 mm. Antennae, segment I, 3-3-3-7 mm., (only paraclypeae o-7-0-8 mm.), II, 2-3- 
2:5 mm., III, 2-4-2-6 mm., IV, missing. Rostrum, segment I, 1:3-1-5 mm., II, 1-4-1-5 mm., 
III, 0-7-0-8 mm., IV, 0-8-0-85 mm. Pronotum, length, 2:50-2:66 mm., width, 2:2-2-3 mm. 
Hind femora, 5:7-6:4 mm. Hind tibia, 6-1-6:7 mm. Distance between ocelli, 0-5-0-6 mm., 
distance between an ocellus and an eye, 0-6—-0-7 mm. 

Very similar to gf in general appearance and colour ; posterior margin of seventh abdominal 
sternum with a “ U ’’-shaped median indentation. 

Q GENITALIA: First gonocoxae almost triangular in form, apices rounded ; intervalvular 
sacs absent ; spermatheca round, bladder-shaped with a long and thick coiled tube (Text- 
fig. 101). 

Holotype 3, Brazit: State of Amazonas, Rio Cairy Uaapes, 1906 (Schmidt), 
A.M.N.H. 


Paratypes. 1d, 1 9, with data as holotype, A.M.N.H. ; Mexico: Tapanatepec, 
3 miles north Oaxaca, 2 J, I 9, 22.xii.1955 (J. C. Schaffner), Ahmad Coll. ; 1 g with 
some data as above, in Coll. J. C. Schaffner ; PANAMA CANAL ZONE : Darien, 2 9 
(fF. Geay), Mus. Hist. nat. Paris. 


This species seems to be distributed as far as Oaxaca (Mexico) in the northern part 
of Neotropical region. 


This species is close to S. tipulotdes (Degeer) and S. americana sp. n. but can be 
separated by parallel-sided head, by uniformly thin, straight and long paraclypeae, 
which are rounded at the apices and by the lack of black spots on the posterior 
angles of pronotum. 


ji! 
ry) 


104 


Fics. 101-105. Stenocoris braziliensis, 2. 101, spermatheca ; 102, terminalia, ventral 
view ; 103, first gonopophysis, ventral view ; 104, ovipositor, ventral view ; 105, 
same, dorsal view. 


THE LEPTOCORISINAE OF THE WORLD 41 


Stenocoris (Stenocoris) claviformis sp. n. 
(Text-figs. 106-115) 


Characterized by club-shaped appendage in the aedeagus, by truncated processes 
of 9 seventh abdominal sternum and by ¢ and 9 genitalia. 


3. Body length, 13-5-14:2 mm., width, 2-0-2:15 mm. Head length, 1-6-1-7 mm., width, 
I-4-1'6 mm. Antennae, segment I, 3:3-3-6 mm., II, 2-:1-2:3 mm., III, 2-3-2-6 mm., IV, 
3:°6-4-0 mm. Rostrum, segment I, 1:5-1:-6 mm., II, 1-5-1:-6 mm., III, 0-8—o0-83 mm., IV, 
o-8-0-85 mm. Pronotum, length, 2-3-2-4 mm., width, 2-0-2:15 mm. Hind femora, 5-5— 
56mm. Hind tibia, 6-0-6-1 mm. Distance between ocelli, 0-4-0-45 mm., distance between 
an ocellus and an eye, 0-7—0-73 mm. 

Body robust, pale, tinged with black. Head comparatively short, slightly longer than broad ; 
occipital suture medially concave, sides curving outwards ; paraclypeae comparatively short, 
thinner at base and rounded at apices. Antennae brownish black, with underside of basal 
segments, bases and middle of second and bases of apical segments pale. Rostrum com- 
paratively long, extending well beyond apices of second coxae. Lateral black line well represen- 
ted, extending from genae on to anterior propleuron. Pronotal disc with blackish punctures, 
black spots on posterior angles present. Legs with apices of femora, tibiae and tarsi tinged 
with dark brown. Hemielytra with black punctures. Abdomen with dorsum, excluding 
connexiva, red, venter pale ochraceous. Posterior margin of seventh abdominal tergum 
truncated, usually slightly produced in middle ; pygophore broader at base, tapering pos- 
teriorly ; posterior margin about two fifths of width of posterior margin of seventh abdominal 
tergum. 

d GENITALIA: Claspers uniformly thin, strongly curved and acutely pointed at apices ; 
very small, fused, rod-like dorsal thecal appendages present, ventral appendages paired, thick 
at bases and very thin and curved near apices ; four left lateral conjunctival appendages as in 
Text-fig. 113 ; frontal appendages longest of all, left thick, longer and curved ; dorsal con- 
junctival appendages as in Text-fig. 113. 

9. Body length, 14-4-15-0 mm., width, 2:2-2:3 mm. Head length, 1-7-1-8 mm., width, 
I-4-1'6 mm. Antennae, segment I, 3:2-4-:1 mm., II, 2-0-2-:7 mm., III, 2:2-2:5 mm., IV, 
3:8-4:2 mm. Rostrum, segment I, 1-5-1-6 mm., II, 1-5-1-6 mm., III, 0-7-0-8 mm., IV, 
o-8-o-9 mm. Pronotum, length, 2:3—2-5 mm., width, 2:2-2:3 mm. Hind femora, 5:3—-5-6 mm. 
Hind tibia, 5-6-6-1 mm. Distance between ocelli, o-4-0-46 mm., distance between an ocellus 
and an eye, 0-7—-0-8 mm. 

Very similar to § but slightly longer, posterior margin of seventh abdominal venter medially 
indented, giving two truncated processes (Text-fig. 111). 

Q GENITALIA: First gonocoxae comparatively small, apices truncated (Text-fig. 110) ; 
intervalvular sacs absent ; spermatheca tubular, slightly thicker in middle with a long coiled 
tube as in Text-fig. 109. 


Holotype J, Conco : Kasongo, ix.1959 (P. L. G. Benott), Mus. R. Afr. centr. 


Paratypes. Id with same data as holotype, Mus. R. Afr. centr. ; Uelé, Gangalano 
Bodio, Lulua, Capsanga, Bongo da lemba, 5 g, 7 9, 15.v.1936, ix.1952, 1.1933, Ul. 
1913 (various collectors), Mus. R. Afr. centr. ; CAMEROON : Lolodorf, 8 J, 3 9 (A. I. 
Good), Carnegie Mus. 


This species is closest to S. africana sp. n. in having small posterior processes of 
pygophore and truncated processes of 9 seventh abdominal sternum. However it 
can easily be separated from S. africana by long rostrum, extending well beyond the 
apices of second coxae. 


42 


Fics. 106-115. Stenocoris claviformis. 


I. AHMAD 


107, 108, 112-113, g, 106, IogQ-III, 114-115, 9. 
107, terminalia, dorsal view ; 108, claspers, 


inner and outer views ; 109, spermatheca ; 110, first gonocoxa, ventral veiw ; III, 
terminalia, ventral view ; 112, theca, dorsal view ; 113, aedeagus (inflated), ventral 


view ; 114, Ovipositor, ventral view ; 115, same, dorsal view. 


106, first gonopophysis, ventral view ; 


THE LEPTOCORISINAE OF THE WORLD 43 


Stenocoris (Stenocoris) ethiopis sp. n. 
(Text-figs. 116-118) 


Characterized by small size of the body, divergent paraclypeae and by the form of 
the g and 9 genitalia. 


3. Body length, 12-0 mm., width, 1-8 mm. Head length, 1-6 mm., (only paraclypeae 
0-6 mm.), width, 1-3 mm. Antennae, segment I, 3:2 mm., II, 2-0 mm., III, 2-3 mm., IV, 
471mm. Rostrum, segment I, 1-3 mm., II, 1-3 mm., III, 0-65 mm., IV, 0-7 mm. Pronotum, 
length, 2:2 mm., width, 1-8 mm. Hind femora, 5-1 mm. Hind tibia, 5-6 mm. Distance 
between ocelli, 0-4 mm., distance between an ocellus and an eye, 0-7 mm. 


05 mm. 


Fics. 116-118. Stenocoris ethiopis, 3. 116, terminalia, dorsal view ; 117, claspers, inner 
and outer views ;_ 118, aedeagus (inflated), ventral view. 


Body comparatively very short and slim, pale, tinged with black. Head only about a 
quarter as long again as broad ; occipital suture medially concave, arms curving outwards ; 
paraclypeae about one-third of head, thicker in middle, very thin and divergent at apices. 
Antennae with underside of basal segments, articulations of first and second, second and third, 
middle of second and basal portions of apical segments, pale. Rostrum extending at least 
slightly beyond apices of second coxae. Lateral black line faint, extending from genae on to 
anterior of propleuron. Pronotal disc pale with black punctures, with smooth black spots on 
posterior angles. Legs with apices of femora slightly brownish. Hemielytra with black 
punctures on anal and claval areas. Abdomen with dorsum pale, unicolourous, venter pale 
ochraceous. Posterior margin of seventh abdominal tergum truncated, slightly laterally 
produced on either side ; pygophore with small and rounded posterior processes, posterior 
margin a third of width of posterior margin of seventh abdominal tergum. 


An I. AHMAD 


¢ GENITALIA: Claspers thicker in middle, strongly curved and pointed at apices ; dorsal 
thecal appendages absent, ventral paired, very thin, elongated and truncate at apices, four 
left lateral conjunctival appendages as in Text-fig. 118 ; frontal appendages paired, long, thin 
and curved at bases ; four right lateral conjunctival appendages with fourth bifurcated at 
apex (Text-fig. 118) ; dorsal appendages as in Text-fig. 118. 

No 9 available for study. 


Holotype g, GHANA : Tafo, 24.i1i-20.1v.1959 (R. Smith) (recorded by Southwood, 
1961, on light trap), B.M.(N.H.). 


Paratypes. 2d, same data as holotype, but without terminalia, B.M.(N.H.). 
This seems to be a rare species and is probably confined to West Africa. 


This species is close to S. claviformis sp. n. and S. africana sp. n. but can be sepa- 
rated from them in having medium sized rostrum only slightly extending beyond 
apices of second coxae (extending well beyond the apices of second coxae in clavi- 
formis and in africana never extending beyond second coxae). Its bifurcated fourth 
right lateral conjunctival appendage is its peculiarity. 


Stenocoris (Stenocoris) maculosa (Blote) comb. n. 


(Text-figs. 119-127) 


Leptocorixa (Stenocoris) maculosa Blote, 1937 : 290 ; Villiers, 1963 : 232 


Characterized by long rostrum extending well beyond the apices of second coxae 
and by the form of the ¢ and 9 genitalia. 


Figs. 119-121. Stenocorvis maculosa, g. 119, terminalia, dorsal view ; 120, claspers, 
inner and outer views ; 121, aedeagus (inflated), ventral view. 


THE LEPTOCORISINAE OF THE WORLD 45 


6. Body length, 13-0-15:5 mm., width, 1-9-2-1 mm. Head length, 1-7-1-9 mm., width, 
I-2—1-3 mm. (only paraclypeae 0-6 mm.). Antennae, segment I, 3-2—-3-5 mm., II, 2-1-2-4 mm., 
III, 2-3-2:5 mm., IV, 3:5-3:7 mm. Rostrum, segment I, 1-3-1-4 mm., II, 1-3-1-4 mm., III, 
0-7-0:73 mm., IV, 0:75-0:85 mm. Pronotum, length, 2:2-2-4 mm., width, 1-9-2: mm. Hind 
femora, 5-I-6-0 mm. Hind tibia, 5:3-6-5 mm. Distance between ocelli, 0-4 mm. ; distance 


between an ocellus and an eye, 0-75 mm. 


122, ovipositor, ventral view ; 123, first gonopo- 


Fics. 122-127. Stenocoris maculosa, 9. 
125, first gonocoxa, ventral view ; 


physis, ventral view ; 124, terminalia, ventral view ; 
126, spermatheca ; 127, ovipositor, dorsal view. 


46 I. AHMAD 


Body linear, pale, tinged with black. Head at least a third as long again as broad ; occipital 
suture medially concave, arms curving outwards towards sides ; paraclypeae long, uniformly 
thin and rounded at apices. Antennae with underside of basal segment, middle of second and 
third and bases of fourth, pale. Rostrum long, extending well beyond apices of second coxae. 
Lateral black line usually represented by two black dots on sides of antenniferous tubercles and 
collar. Pronotal disc tinged with hght brown, posterior angles with brownish black spots. 
Legs with apices of femora tinged with red. Hemielytra blackish, anal and claval areas with 
black punctures. Abdomen with dorsum, excluding connexiva, red, venter pale ochraceous. 
Posterior margin of seventh abdominal tergum truncate ; pygophore small, rounded, with 
emarginate posterior margin. 

dg GENITALIA : Claspers thicker in middle with a long thin neck, apices pointed ; dorsal 
thecal appendages absent, ventral pair curved in middle and rounded at apices ; four left 
lateral conjunctival appendages as in Text-fig. 121 ; frontal appendages paired, very long, 
uniformly thin and curved near bases and apices ; four right lateral conjunctival appendages 
as in Text-fig. 121 ; dorsal appendages as in Text-fig. 121. 

9. Body length, 14-5-16-0 mm., width, 2-1-2-4 mm. Head length, 1-8-2:0 mm., width, 
I-4-I'5 mm. Antennae, segment I, 3:1-3:2 mm., II, 2:2-2-4 mm., III, 2:3-2:5 mm., IV, 
3:0-3°6 mm. Rostrum, segment I, 1-5-1:6 mm., II, 1:5-1-6 mm., III, 0-7-0:75 mm., IV, 
0:75-0°85 mm. Pronotum, length, 2-4-2:8 mm., width, 2-1-2-4 mm. Hind femora, 5-9— 
62mm. Hind tibia, 6-3-6-5 mm. Distance between ocelli, 0-4—-0-5 mm., distance between an 
ocellus and an eye, 0-7—0-85 mm. 

Very similar to g in general appearance and colour, slightly larger ; posterior margin of 
seventh abdominal sternum almost inverted “‘ V ’’-shaped in middle as in Text-fig. 124. 

Q GENITALIA: First gonocoxae largely concealed by seventh abdominal sternum, apices 
rounded ; intervalvular sacs absent, spermatheca tapering anteriorly with lateral margins 
irregular and a small, thick and coiled tube (Text-fig. 126). 


Type material. Holotype gf, Sout Arrica : Pondoland, Port St. John, x.1923 
(R. E. Turner), B.M.(N.H.) ; allotype 9, data as holotvpe, B.M.(N.H.) ; paratypes, 
SouTtH ArFrica : Zululand, Natal, Durban, 4 g, 3 9, 20-21.11.1925 (C. N. Barker), 
B.M.(N.H.) ; Umkomaas River, 1 gf, 1847, B.M.(N.H.) ; UGANDA : Siroko R., near 
foot of Mt. Elgon, 3,600 ft., 1 g, I-14.viii.1g11 (S. A. Neave), B.M.(N.H.). 


Other material. Conco: 30 9, 36 2; UGANDA: 14,292; ANGOLA: 24; 
TANGANYIKA : 144,592; CAMEROONS: Ig; RHODESIA (N. and S.): 24. 


The range of this species seems to extend in the west into Cameroons and in the 
east into Tanganyika, but it is particularly common in southern Africa, Rhodesia and 
Angola. The adults have been collected from May until November. 


This species is close to S. phthisica (Gerstacker), S. pallida (Blote) and S. africana 
sp. n. in general appearance but can easily be separated from them by the long rost- 
rum, which extends well beyond the apices of second coxae. 


Stenocoris (Stenocoris) pallida (Blote) comb. n. 
(Text-figs. 128-137) 
Leptocorixa (Stenocoris) pallida Bléte, 1937 : 295 


Characterized by the pale unicolourous disc of pronotum, which is without black 
spots on posterior angles, by a black dot on sides of collar and by the form of fj and 9 
genitalia. 


THE LEPTOCORISINAE OF THE WORLD 47 


g. Body length, 13-5-14-5 mm., width, 1-9-2: mm. Head length, 1-8-2-0 mm., width, 
I-3-I-4 mm. (only paraclypeae 0-6-0-7 mm.). Antennae, segment I, 3-6-3:8 mm., II, 2-4— 
2-6mm., III, 2-5-2-:7 mm., IV, 3-8-4: mm. Rostrum, segment I, 1-3—1-5 mm., II, 1-3-1-5 mm., 
III, 0-65-0-75 mm., IV, 0:75-0:85 mm. Pronotum, segment I, 2-3-2:6mm., width, 1-9—2:1 mm. 
Hind femora, 6:0-6-4 mm. Hind tibia, 6-56-83 mm. Distance between ocelli, 0-4-0-45 mm., 
distance between an ocellus and an eye, 0-6—0-67 mm. 


‘ 
Vth vee, 
tat baty 77 


0:5 mm. 


Figs. 128-131. Stenocoris pallida, $. 128, terminalia, dorsal view ; 129, head, dorsal 
view ; 130, claspers, inner and outer views ; 131, aedeagus (inflated), ventral view. 


Body comparatively robust, pale, unicoloured. Head long, more than a third as long again 
as broad ; occipital suture medially ‘“‘ V’’-shaped, arms curving outwards towards sides ; 
paraclypeae long, more than a third as long as head, thicker at bases and rounded at apices. 
Antennae pale, with apices of second, third and apical three-quarters of fourth segments 
brownish black. Rostrum short, hardly reaching apices of second coxae. Lateral black line 
usually represented by a black dot on sides of collar. Pronotal disc pale unicolourous, black 
spots on posterior angles absent. Legs with apices of femora usually tinged with red. Hemiely- 
tra usually brownish black. Abdomen with dorsum, excluding connexiva, red ; venter pale 
ochraceous. Posterior margin of seventh abdominal tergum truncate ; pygophore elongate, 
thicker at base and tapering posteriorly. 

6 GENITALIA: Claspers strongly curved in middle, strongly curved and pointed at apices ; 
dorsal thecal appendages absent ; ventral pair long, almost straight and rounded at apices ; 
four left lateral conjunctival appendages as in Text-fig. 131 ; frontal conjunctival appendages 
paired, long, at least as long as or longer than first lateral conjunctival appendages ; four 
right lateral conjunctival appendages as in Text-fig. 131 ; a single dorsal appendage as in 
Text-fig. 131. 

®. Body length, 14-5-16-0 mm., width, 2:0-2:1 mm. Head length, 1:8-2:0 mm., width, 
I-3-1:5mm. (only paraclypeae 0-6-0-7mm.), Antennae,segment I, 3:1-3:8mm.,II,2-3-2:5mm., 
III, 2-4-2:7 mm., IV, 3:9-4:2 mm. Rostrum, segment I, 1-3-1-4 mm., II, 1-3-1-4 mm., III, 


48 I. AHMAD 


0:7-0°75 mm., IV, 0:75-0:85 mm. Disc of pronotum 2-3—2-6 mm., width, 2-0-2: mm. Hind 
femora, 6:0-6-2 mm. Hind tibia, 6-5-6-7 mm. Distance between ocelli, 0-4-0-45 mm., 
distance between an ocellus and an eye, 0-6-0-7 mm. 

Very similar but usually larger than g ; posterior margin of seventh abdominal sternum 
almost an inverted ‘‘ V’’-shape in middle as in Text-fig. 132, large basal portion of ninth 
paratergite visible. 


Fics. 132-137. Stenocoris pallida, 9. 132, terminalia, ventral view ; 133, ovipositor, 
ventral view ; 134, same, dorsal view ; 135, first gonopophysis, ventral view ; 136, 
first gonocoxa, ventral view ; 137, spermatheca. 


THE LEPTOCORISINAE OF THE WORLD 49 


Q GENITALIA: First gonocoxae largely concealed by seventh abdominal sternum, outer 
margin irregular ; intervalvular sacs absent ; spermatheca irregular, pelican-shaped with a 
long, thick and coiled tube. 


Type material. Holotype g, UGANDA : between Mitiana and Entebbe, 3,800 ft., 
g-I1.i. (S. A. Neave), B.M.(N.H.) ; allotype 2, and paratypes, 4 3, 1 2, with same 
data as holotype, B.M.(N.H.) ; Kenya : Shimba, 4 J, 3 9, vii.1g12 (A. D. Milne), 
B.M.(N.H.) ; NyASALAND : Zomba, Mlanje, Mimosa, 3 g, 3 9, iv.1962, 7.xi.1913, 
10.v.1919, B.N.(M.H.) ; Zomba, Mlanje, 2,300 ft., lower Shire valley, near Chikawa, 
600 ft., I5.viil., 5.ix., 7.xX1.I1913, I6.iv.IgI0, 13.vi., vil.IgI2 (various collectors), 
B.M.(N.H.) ; ZANZIBAR: I 4, 1.11.1925 (H. J. Shell), B.M.(N.H.) ; KENYA: 
Mazimosa, Tayari, I 4, 9.ii. (B. Elbon), B.M.(N.H.) ; UGanpba: S. E. Shore of L. 
Kiaja, 2,400 ft., Entebbe, 3 g, 2 9, II.vili.IgII, 14-25.v.1g12 (S. A. Neave, C. C. 
Gowdey), B.M.(N.H.). 


Other material. KENYA: 34,492; TANGANYIKA: 10g¢,132; UGANDA: 44, 
ZS. 

This is an East African species. Its range extends in the south into Nyasaland 
and in the north into Kenya. It has not been reported from central or western 
Africa. The adults have been collected mainly between April and November, but 
2 9 were collected from Zanzibar and Kenya in February. 


This species is closest to S. phiiisica (Gerstacker) but can be separated from it by 
the paraclypeae, which are thicker at base and thinner and rounded at apices in this 
species (Text-fig. 129), whilst they are thicker in the middle in phthisica (Text-fig 
140). The lateral black line in the form of a black dot on either side of collar and 
lack of black spots on the posterior angles of pronotum separate it from all the other 
species of the subgenus Stenocoris. 


Stenocoris (Stenocoris) phthisica (Gerstacker) comb. n. 
(Text-figs. 138-147) 


Leptocorisa phthisica Gerstacker, 1873 : 406 
Leptocorixa (Stenocoris) annulicornis Signoret ; Blote, 1937 : 289 syn. n. 
Leptocorixa (Stenocoris) blotei Villiers, 1963 : 232 Syn. n. 


Characterized by long paraclypeae, which are thicker only in the middle and are 
rounded at apices (Text-fig. 140) and by the form of j and @ genitalia. 


6. Body length, 13-5-15-0 mm., width, 1-8-2-0o mm. Head length, 1-7-1-9 mm., (head 
exposed 2-:0-2:1 mm.), width, 1-:2-1-3 mm. (only paraclypeae 0-66-0-:7 mm.). Antennae, 
segment I, 3-6-3-9 mm., II, 2-6—2-7 mm., III, 2-7—2-:75 mm., IV, 4-0-4: mm. Rostrum, segment 
I, r-1-1-4 mm., II, 1-1-1-4 mm., III, 0-6-0-7 mm., IV, 0:7-0:75 mm. Pronotum, length, 
2°I-2:4 mm., width, 1-8-2-:0 mm. Hind femora, 5-9-6:7 mm. Hind tibia, 6-3-6-9 mm. 
Distance between ocelli, 0-4—0-5 mm., distance between an ocellus and an eye, 0-66—-0-7 mm. 

Body linear, pale unicolourous. Head long, about half as long again as broad ; occipital 
suture medially concave, arms following straight towards sides ; paraclypeae long, about two- 
fifths of entire head, thicker in middle and thinner at bases and apices, apices rounded (Text- 
fig. 140). Antennae brownish black with underside of basal segment, middle of second and 
third and bases of apical segments pale. Rostrum short, reaching second coxae. Lateral black 
line faint, usually extending from sides of antenniferous tubercles on to anterior of propleuron. 


50 I. AHMAD 


Pronotal disc pale unicoloured, black spots on posterior angles sometimes present. Legs with 
apices of femora and bases of tibiae tinged with red. Hemielytra tinged brownish black. 
Abdomen with dorsum tinged with light brown, venter pale ochraceous. Posterior margin of 
seventh abdominal tergum truncate, pygophore elongated, thinner in middle, posterior margin 
emarginate (Text-fig. 138). 

6 GENITALIA: Claspers long, uniformly thin and rounded at apices (Text-fig. 139) ; dorsal 
thecal appendages absent ; ventral pair thinner at bases, thick and rounded at apices ; four 
left lateral conjunctival appendages as in Text-fig. 141; frontal conjunctival appendages 
paired, comparatively short and thick, left uniformly thin and curved ; four right lateral con- 
junctival appendages as in Text-fig. 141 ; dorsal appendage as in Text-fig. 141. 

2. Body length, 15-5-16-0 mm., width, 2:1-2:2 mm. Head length, 1-8-2:0 mm., width, 
I-3-1:35 mm. Antennae, segment I, 3:5-3-6 mm., II, 2-:3-2-4 mm., III, 2-4-2:5 mm., IV, 
3°9-4:1 mm. Rostrum, segment I, 1-:3-1-4 mm., II, 1-4-1:5 mm., III, 0-7-0-°8 mm., IV, 
0:75-0'8mm. Pronotum, length, 2:3-2-6mm., width, 2-1-2:2mm. Hind femora, 6-0-6:3 mm. 
Hind tibia, 6-3-6:6 mm. Distance between ocelli, 0-4-0:5 mm., distance between an ocellus 
and an eye, 0-66—-0-7 mm. 

Very similar to gf in general appearance and colour but usually larger ; posterior margin of 
seventh abdominal sternum with “ U ’’-shaped indentation in the middle and curved towards 
extremities as in Text-fig. 142. 

Q GENITALIA: First gonocoxae with outer margin irregular, apices rounded (Text-fig. 146), 
almost triangular in external view ; intervalvular sacs absent ; spermatheca irregular, thicker 
in middle, with a long and coiled tube of varying thickness (Text-fig. 147). 


Hl 


ait 
yyy 


0-5 mm. 


Fics. 138-141. Stenocoris phthisica, 3. 138, terminalia, dorsal view ; 139, claspers, inner 
and outer views ; 140 head, dorsal view ; 141, aedeagus (inflated), ventral view. 


THE LEPTOCORISINAE OF THE WORLD 51 


Type material. Holotype J, East Arrica : Mbaramu (V. I. Deck), Humbold. 
Univers. Berlin ; cotype g, without data, Humbold. Univers. Berlin. 

Other material. Conco : 124,892; TANGANYIKA : 69,39; KENYA: 64,69; 
SEYCHELLES : Ig; MAURITIUS: I. 


0:25 mm. 


Fics. 142-147. Stenocoris phthisica, 9. 142, terminalia, ventral view ; 143, Ovipositor, 
ventral view ; 144, same, dorsal view; 145, first gonopophysis, ventral view ; 146, 
first gonocoxa, ventral view ; 147, spermatheca. 


52 I. AHMAD 


This species is distributed in eastern and central Africa. Its range extends in the 
east into Mauritius and Seychelles and in the west into Congo, Central Africa. 
Adults have been collected during March to August. 


This species is close to S. maculosa (Blote), S. pallida (Blote), and S. africana sp. n. 
but can easily be separated from them by paraclypeae which are long, thicker in the 
middle, and very thin and rounded at apices. 

Blote redescribed Leptocorisa annulicormis Signoret, 1860 (now Stenocoris (Erbula) 
annulicornis) and placed it under the subgenus Stenocoris. Villiers showed that 
Blote had misidentified Signoret’s species and suggested that annulicornis Bléte nec 
Signoret from Seychelles was a new species and named it blotei. It has not been 
possible to borrow types of L. (S.) blote: for the present study, but g specimens (with 
Blote’s det. label) from Mauritius and Seychelles in the B.M.(N.H.) have been 
examined. The structure of the aedeagus shows that L. blotei is a synonym of S. 
phtlisica (Gerstacker). However 9 specimens from Seychelles : Mahe, 5.12.1905, 
leg. J. S. Gardiner (B.M.(N.H.)) also seen by Blate are in fact true annulicornis and 
this would appear to apply also to the supposed 9 of L. (S.) blote: figured by Villiers 
(1963 : 232). The posterior margin of seventh abdominal sternum is produced in 
the middle and the posterior margin of this projection is slightly concave in the 
middle, which is characteristic of the subgenus Evbula. 


Stenocoris (Stenocoris) sordida (Blote) comb. n. 
(Text-figs. 148-156) 
Leptocorixa (Stenocoris) sovdida Blote, 1937 : 292 ; Villiers, 1955 : 1143, 1144, 1963 : 232 


Characterized by prominent, pointed posterior processes of pygophore, rounded 
processes of posterior margin of 2 seventh abdominal sternum and by the form of ¢ 
and 9 genitalia. 


6. Body length, 12-5-15-0 mm., width, 2:1-2:5 mm. Head length, 1-6-2:1 mm., width, 
I-3-1-5mm. Antennae, segment I, 3:5-3-8 mm., II, 2:3-2:-5 mm., III, 2-31-2:5 mm., IV, 
4:0-4:2 mm. Rostrum, segment I, 1-4-1-6 mm., II, 1-4-1-6 mm., III, 0-7-0-9 mm., IV, 
0-7-0‘9mm. Pronotum, length, 2:3—2-7 mm., width, 2-1-2-5 mm. Hind femora, 5-8-6-0 mm. 
Hind tibia, 6-o-6:-5 mm. Distance between ocelli, 0-4-0-5 mm. ; distance between an ocellus 
and an eye, 0-65-0-8 mm. 

Body comparatively robust, pale, tinged with black. Head usually slightly longer than 
broad ; occipital suture medially concave, curving outwards towards sides ; paraclypeae 
comparatively short, thinner at base and rounded at apices. Antennae dark brown, with 
underside of basal segment and bases of the apical segments pale. Rostrum long, usually 
reaching halfway between apices of second and bases of third coxae. Lateral black line well 
represented from genae on to anterior of propleuron. Pronotal disc pale, posteriorly with black 
punctures, posterior angles with smooth black spots. Legs with apices of femora brown, tibiae 
and tarsi tinged with brownish black. Hemielytra with black punctures. Abdomen with 
dorsum, excluding connexiva, brown, venter pale ochraceous. Posterior margin of seventh 
abdominal tergum truncate ; pygophore large (Text-fig. 148), posterior processes prominent, 
acutely pointed, posterior margin less than half of posterior margin of seventh abdominal 
tergum. 

3d GENITALIA: Claspers strongly curved and acutely pointed at apices; dorsal thecal 


THE LEPTOCORISINAE OF THE WORLD 53 


appendages absent ; ventral pair thicker at base and very thin and rounded at apices ; four 
left conjunctival appendages as in Text-fig. 150; frontal conjunctival appendages paired, 
left slightly longer and straighter ; four right lateral conjunctival appendages as in Text-fig. 
150; a pair of dorsal conjunctival appendages as in Text-fig. 150 without second spine-like 
appendage. 

2. Very similar to f in general appearance, colour and measurements ; posterior margin of 
seventh abdominal sternum medially strongly indented, forming rounded processes on either 
side (Text-fig. 151). 

Q GENITALIA: First gonocoxae not concealing a large portion of ninth paratergite, rounded 
at apices ; intervalvular sacs absent ; spermatheca broad, curved, with a short coiled tube. 


Type material. Holotype g, UGANDA: Entebbe, xii.1g12 (C. C. Gowdey), 
B.M.(N.H.) ; allotype 2 and paratypes, 1 g, 2 9, with same data as holotype, 
B.M.(N.H.) ; shores of L. Isolt or Wanala, 3,800 ft., 1 9, 7-8.i.1912 (S. A. Neave), 
B.M.(N.H.) ; Bugma Forest, Ruwenzori Range, 6,500 ft., Kimbe, Tero Forest, 
Semliki Plain, near south shore of L. Albert, 2,200 ft., 4 3, 4 9, li, vil, ix, xii.1g12, 
1.1913 (various collectors), B.M.(N.H.). 


Other material. SIERRA LEONE: 44,19; GHANA: 44,19; NIGERIA: 19; 
CAMEROONS : Ig; CONGO: 1554, 1412; UGANDA: 24,1 9. 


This is a West African species. Its range extends in the east into central Africa, 
Congo and East Africa, Uganda and in the west into Sierra Leone. Adults have been 
collected in every month of the year but it seems that it is abundant in the months of 
April and May. 


149 


0-5 mm. 


Fics. 148-150. Stenocoris sordida, 3. 148, terminalia, dorsal view ; . 149, Claspers, 
inner and outer views ;_ 150, aedeagus (inflated), ventral view. 


54 I. AHMAD 


S.(S.) apicalis (Westwood) is its closest ally, from which it can be separated by the 
lateral black line, which is well represented on genae and extends beyond sides of 
collar, whereas in S. apicalis this line is faint on genae. The posterior margin of the 
pygophore is comparatively narrow and the processes formed by median indentation 
of 2 seventh abdominal sternum are rounded in this species, whereas they are tri- 
angular in S. apicalis. 


154 


0:5 mm. 


sav/ 155 


\\ 0:25 mm. 


Fics. 151-156. Stenocoris sordida, 9. 151, terminalia, ventral view ; 152, ovipositor, 
ventral view ; 153, same, dorsal view ; 154, first gonopophysis, ventral view ; 155, 
first gonocoxa, ventral view ; 156, spermatheca. 


THE LEPTOCORISINAE OF THE WORLD 55 


Stenocoris (Stenocoris) tipuloides (Degeer) comb. n. 
(Text-figs. 157-165) 


Cimex tipuloides Degeer, 1773 : 354, 1778 : 272 ; Gmelin, 1788 : 2194 

Myodocha tipuloides (Degeer) Latreille, 1807:126; Olivier, 1811 :106; Burmeister, 
1835 : 325 

Stenocoris tipuloides (Degeer) Burmeister, 1839 : 1010 

Leptocorisa tipuloides (Degeer) Amyot et Serville, 1843 : 229 

Gerris tipuloides (Degeer) Stal, 1870 : 218 

Leptocorixa tipuloides (Degeer) Hussey, 1951 : 152 

Leptocorisa crudelis Westwood, 1842 : 18 


Characterized by the round pygophore, the posterior margin of which is medially 
emarginate and by the form of $ and @ genitalia. 


6. Body length, 14-0-15°5 mm., width, 2-1-2:2 mm. Head length, 1-9-2-0 mm., width, 
I-3-1-'4 mm. (length of paraclypeae 0-70 mm.). Antennae, segment I, 3-3-3-7 mm., II, 2-3— 
2‘6mm., III, 2-4-2:7 mm., 1V, 4-:0-4:2mm. Rostrum, segment I, 1-3-1-5 mm., II, 1-4-1-5 mm., 
III, 0-7-0-8 mm., IV, 0-75-08 mm. Pronotum, length, 2-4-2-7 mm., width, 2-1-2-2 mm. 
Hind femora, 6-0-6:7 mm. Hind tibia, 6-4-6-9 mm. Distance between ocelli, 0-5-o-6 mm., 
distance between an ocellus and an eye, 0:66—0-7 mm. 

Body comparatively robust, pale, tinged with light brown. Head slightly tapering behind 
occipital suture, about half as long again as broad ; occipital suture medially ‘‘ U ’’-shaped, 
with arms bending outwards towards margins of head ; paraclypeae long, more than one-third 
length of head, uniformly thin and rounded at apices. Antennae pale, usually tinged with 
pink. Rostrum short, never extending beyond apices of second coxae. Lateral black line 


7 

Yya a 1 

bi 1 Ae Ee craigs yeu vie 
\ 

Vs Fper ny 


if 
> = . “ny Ny tiny 


0:5 mm. 


Fics. 157-159. Stenocoris tipuloides, 3. 157, aedeagus (inflated), ventral view ; 
158, terminalia, dorsal view ; 159, claspers, inner and outer views. 


56 I. AHMAD 


usually faintly represented, extending from sides of antenniferous tubercles on to collar. Pro- 
notal disc uniformly pale, black spots on posterior angles absent. Legs usually with apices of 
femora and bases of tibiae pink. Hemielytra tinged brownish black. Abdomen with dorsum, 
excluding connexiva, pink ; venter pale ochraceous. Posterior margin of seventh abdominal 
tergum truncate ; pygophore rounded, smoothly tapering posteriorly, posterior margin medially 
emarginate (Text-fig. 158). 


Fics. 160-165. Stenocoris tipuloides, 2. 160, terminalia, ventral view ; 161, ovipositor, 
ventral view ; 162, same, dorsal view ; 163, first gonopophysis, ventral view ; 164, 
first gonocoxa, ventral view ; 165, spermatheca. 


THE LEPTOCORISINAE OF THE WORLD 57 


$6 GENITALIA: Claspers thicker at bases, strongly curved in middle (Text-fig. 159) ; dorsal 
thecal appendages absent ; ventral pair long, thicker at bases, very thin and truncated at 
apices ; four pairs of lateral conjunctival appendages as in Text-fig. 157 ; frontal conjunctival 
appendages, paired, long, curved in middle, not at all curved at apices, dorsal appendages as in 
Text-fig. 157. 

2. Body length, 15-0-16-0 mm., width, 2-2-2-3 mm. Head length, 1-9-2-0 mm., width, 
I-3-I-4 mm. (only paraclypeae 0-7omm.). Antennae, segment I, 3-o-3-1 mm., II, 2-:1-2-3 mm., 
III, 2-3-2-4 mm., IV, 3:6-3-9 mm. Rostrum, segment I, 1-3-1-5 mm., II, 1-4-1-5 mm., III, 
0-7-0°8 mm., IV, 0:75-0:85 mm. Pronotum, length, 2:6-2:7 mm., width, 2:2-2:-3 mm. Hind 
femora, 5:9-6-3 mm. Hind tibia, 6-1-6-4 mm. Distance between ocelli, 0-5-0-56 mm., 
distance between an ocellus and an eye, 0-66—0-7 mm. 

Slightly larger than g ; posterior margin of seventh abdominal sternum slightly notched in 
middle, forming rounded lobes, large basal portion of eighth paratergite concealed (Text-fig. 160). 

Q GENITALIA: First gonocoxae comparatively small, outer margins sinuate and apices 
rounded (Text-fig. 164) ; intervalvular sacs absent ; spermatheca flask-shaped with a curved 
long neck and uniformly thick coiled tube (Text-fig. 165). 


Type material. Holotype without data, Riksmus. Stockholm. Holotype Q, of 
Leptocorisa crudelis Westwood, without data, Hope Dept., Univ. Mus., Oxford. 


Other material. U.S.A.: 29; Brazir: 5¢,59; CotomBia : 2¢; MExIco: 
346,59; BriTIsH GUIANA: Ig¢:; PANAMA: 84,592; Ecuapor: 24,292; CUBA: 
ie, PARAGUAY : Ig, I-9. 


This species is very common in the northern Neotropical region, its range extend- 
ing in the north into Texas and Florida, and in the south into Brazil. Adults have 
been collected throughout the year but it seems to be more common in April, May, 
and June. 


This species is closest to S. americana sp. n., from which it can be separated by a 
short rostrum, which never extends beyond apices of second coxae and by the frontal 
conjunctival appendages of the aedeagus. 


Stenocoris (Stenocoris) v-nigrum (Bléote) comb. n. 
(Text-figs. 166-174) 


Leptocorixa (Stenocoris) v-nigrum Blote, 1937 : 293 ; Villiers, 1955 : 1142, 1144, 1963 : 232 


Characterized by long rostrum, which approaches third coxae and by the form of 
3 and 9 genitalia. 


6. Body length, 14:0-15-5 mm., width, 2:1-2:25 mm. Head length, 1-9-2-:25 mm., width, 
I-5-1°6 mm. (only paraclypeae o-6-0-7 mm.). Antennae, segment I, 3-7-3:8 mm., II, 2-7— 
2-9 mm., III, 2-8-3-0 mm., IV, 3:7-3-9 mm. Rostrum, segment I, 1:65-1:75 mm., II, 1-7- 
2-0 mm., III, o-8-o-9 mm., IV, 0-8-1-0 mm. Pronotum length, 2-3-2:5 mm., width, 2-1- 
2:25 mm. Hind femora, 5-o-6-0 mm. Hind tibia, 6-4-6-6 mm. Distance between ocelli, 
0-4-0'5 mm., distance between an ocellus and an eye, 0-7—0-8 mm. 

Body comparatively robust, pale, tinged with black. Head about a third as long again as 
broad ; occipital suture ‘“‘ V’’-shaped in middle, arms continuing straight towards sides ; 
paraclypeae about one-third of head length, uniformly thick and rounded at apices. Antennae 
brownish black, tinged with red, underside of basal segment, articulation between first and 
second segment, middle of second and at least basal third of apical segment pale. Rostrum 
long, usually reaching anterior margin of third coxae. Lateral black line well represented on 


58 I. AHMAD 


genae, extending on to collar. Pronotal disc pale, posteriorly with brownish black punctures, 
black spots on posterior angles present. Legs pale, with apices of femora, tibiae and tarsi 
brownish. Hemielytra tinged with black. Abdomen with dorsum, excluding connexiva, red, 
venter pale unicolourous. Posterior margin of seventh abdominal tergum truncate ; pygo- 
phore large, tapering posteriorly, posterior margin truncate (Text-fig. 166). 


Fics. 166-171. Stenocoris v-nigyum, 166, g terminalia, dorsal view ; 167, aedeagus (in- 
flated), ventral view ; 168, ovipositor, ventral view ; 169, ¢ claspers, inner and outer 
views ; 170, § first gonopophysis, ventral view ; 171, ovipositor, dorsal view. 


THE LEPTOCORISINAE OF THE WORLD 59 


dg GENITALIA: Claspers thicker in middle with thin and curved apices (Text-fig. 169) ; 
dorsal thecal appendages absent ; ventral pair long, uniformly thick, slightly curved in middle 
and rounded at apices ; four left lateral conjunctival appendages as in Text-fig. 167 ; frontal 
conjunctival appendages paired, minute, reduced to spines, four right lateral conjunctival 
appendages as in Text-fig. 167 ; dorsal appendages as in Text-fig. 167. 

®. Body length, 15-2-16-6 mm., width, 2-2-2-6 mm. Head length, 2-0-2-4 mm., width, 
I-5-1-6 mm. (only paraclypeae o-7-0-°8 mm.). Antennae, segment I, 3-7-3:8 mm., II, 
2-7-2:9 mm., III, 2-75-30 mm., IV, 3-9-4: mm. Rostrum, segment I, 1-6-1:8 mm., II, 
I-7-1-:9 mm., III, o-g-1-0 mm., IV, 0-9-1: mm. Pronotum, length, 2:4-2:8 mm., width, 
2:2-2°6 mm. Hind femora, 5-6-6:1 mm. Hind tibia, 6-0-6-6 mm. Distance between ocelli, 
0-4-0-45 mm., distance between an ocellus and an eye, 0-7—-0-75 mm. 

Distinctly longer than g ; posterior margin of seventh abdominal sternum almost of inverted 
“VY ’’-shape in middle, forming small rounded lobes (Text-fig. 172). 

Q GENITALIA: A large portion of first gonocoxae covered by seventh abdominal sternum, 
apices rounded ; intervalvular sacs absent, spermatheca tubular with a thick, curved and 
short neck (Text-fig. 174). 


Type material. Holotype g, UGANDA: Southern Toro, Mbarara, Fort Portal 
Road, 3,800—4,200 ft., 22-24.x.1911I (S. A. Neave), B.M.(N.H.) ; allotype 9, 1 para- 
type 3, same data as holotype, B.M.(N.H.) ; paratypes, Toro or Doro Forest, 4,000- 
4,500 ft., I g, 25-29.x.1g1I (S. A. Neave), B.M.(N.H.) ; Western Ankola, 4,500- 
5,000 ft., r 9, 10-14.x.1911 (S. A. Neave), B.M.(N.H.). 


a 025 mm. 


Fics. 172-174. Stenocoris v-nigrum, 9. 172, terminalia, ventral view ; 173, first 
gonocoxa, ventral view ; 174, spermatheca. 


60 I. AHMAD 


Other material. Conco: 104,99; GHANA: 24,19; TANGANYIKA: I04, 
aes 

The range of this species extends in the west into Ghana and in the east into 
Tanganyika, but this seems to be more common in eastern Africa. Adults have been 
collected mainly in April and May. 


This species is isolated in the subgenus Stenocoris s. str. by the characters of long 
rostrum, which approaches third coxae, greatly reduced frontal conjunctival append- 
ages in the aedeagus and by tubular spermatheca. 


STENOCORIS (ORYZOCORIS) subgen. n. 


Leptocorisa Latreille 1829 ; Laporte, 1833 : 25 ; Stal, 1873 : 87 
Leptocorixa Laporte ; Hussey, 1951 : 152 


Type-species : Cimex filiformis Fabricius by monotypy. 


Body linear. Head elongated, almost half as long again as broad ; paraclypeae short, 
straight and rounded at apices ; ocelli wide apart, interocellar distance only slightly shorter 
than distance between an ocellus and an eye (in both cases including the diameter of both). 
Antennae with apical segment pale only at base, second subequal to third. Rostrum very short, 
hardly approaching second coxae. Lateral black line usually entirely absent, sides of head and 
thorax pale. Pronotum flat with latero-posterior tips usually pale but sometimes brownish 
black. Abdomen dorsally yellowish red, abdomen ventrally usually pale unicoloured. 

In g, posterior margin of seventh abdominal tergum truncated ; pygophore elongated, 
laterally posterior tips acutely pointed ; claspers asymmetrically crossed over pygophore, 
curved in the middle and strongly pointed at apices ; aedeagus with only ventral pair of thecal 
appendages, dorsal pair wanting ; frontal conjunctival appendages paired, very small ; mem- 
branous appendage rounded with paired dorsal appendages, terminal appendage absent. In 
9, posterior margin of seventh abdominal sternum with short median split ; first gonocoxae 
small, almost truncated at apices ; intervalvular sacs absent ; spermatheca very large oval, 
with a short coiled tube of varying thicknesses. 


This subgenus can be separated from other subgenera, in the g by the claspers 
which, at rest, are always crossed over the pygophore and are not in a socket (Text-fig. 
178) and in the 2 by the posterior margin of seventh abdominal sternum, which is 
always with a short median split (Text-fig. 181). 


Laporte described Leptocorisa and included linearis (Nearctic and Neotropical) (= 
filiformis Fabricius) but in 1830 Guérin had already described his flavida (Oriental) 
( = acuta Thunberg syn. n.) under Lepftocorisa Latreille. This was the first redescrip- 
tion of Leptocorisa Latreille, 1829 and flavida, the first and only species ever included 
in the genus, became the type-species of Leptocorisa Latreille by monotypy. Stal 
divided Leptocorisa into four subgenera as follows : (1) Rhabdocoris Kolenati, 1845 ; 
(2) Stenocoris Burmeister, 1839 ; (3) Evbula Stal, 1873 ; (4) Leptocorisa Laporte s. 
str. During the present study Rhabdocoris Kolenati (type-species arcuata Kolenati, 
1845) has been synonymized with Leptocorisa Latreille and all the Oriento-Australian 
species (previously described under subgenus Rhabdocoris) have been separated from 
other Ethiopian, Nearctic and Neotropical species. Stenocoris being the senior 
subgenus has been given generic rank and Leptocorisa Laporte (a hitherto undes- 
cribed subgenus) has been described here as Oryzocoris. 


THE LEPTOCORISINAE OF THE WORLD 61 


KEY TO SPECIES 
1 Median longitudinal red line on ventral abdominal segments present, sometimes faint ; 
pygophore with posterior processes rounded and pointing posteriorly (Text-fig. 
195) ; basal portions of eighth paratergites largely visible (Text-fig. 192) 
furcifera (p. 67) 


0:25 mm. 


: 175 


176 177 


Fics. 175-177. Stenocovis (Oryzocorvis) spp. 175, filiformis; 176, furcifera; 177, 
fabricti. A, claspers, outer and inner views ;_ B, first gonacoxa, ventral view; C, 
spermatheca. 


62 I. AHMAD 


— Median longitudinal red line absent ; pygophore with posterior processes pointed and 
facing each other (Text-fig. 185) ; basal portions of eighth paratergites largely 
concealed by first pair of gonocoxae (Text-fig. 181) . ; 2 
2 Pygophore greatly elongated, posterior margin very narrow, about one-sixth of 
posterior margin of seventh abdominal tergum (Text-fig. 178) ; lobes of 2 seventh 
abdominal sternum with posterior margin medially concave (Text-fig. 181). (Second 
right lateral conjunctival appendage of aedeagus sickle-shaped (Text-fig. 180)) 
fabricii (p. 62) 
— Pygophore almost rectangular, posterior margin broad, more than half of posterior 
margin of seventh abdominal tergum (Text-fig. 185) ; lobes of 2 seventh abdominal 
sternum with convex posterior margin (Text-fig. 187). (Second right lateral con- 
junctival appendage of aedeagus almost straight (Text-fig. 186)). filiformis (p. 64) 


Stenocoris (Oryzocoris) fabricii sp. n. 
(Text-figs. 177-184) 


Characterized by extremely narrow posterior margin of pygophore, by the tri- 
angular form of first gonocoxae and by other ¢ and 9 genital characters. 


6. Body length, 14-5-15:06 mm., width, 2-0-2: mm. Head length, 1-8-1-9 mm. (para- 
clypeae only, 0-5-0-55 mm.), width, 1-4 mm. Antennae, segment I, 3-6-3-73 mm., II, 
2:0-2:26mm., III, 2:1-2-4 mm., IV, 4:0-4:4 mm. Rostrum, segment I, 1-33 mm., II, 1:33 mm., 
III, 0-66 mm., IV, 0-66 mm. Pronotum, length, 2-4-2-5 mm., width, 2-0-2: mm. Hind 
femora, 5:46-5-6mm. Hind tibia, 5-73-6-:0mm. Distance between ocelli, 0-46 mm., distance 
between an ocellus and an eye, 0-7 mm. 


Fics. 178-180. Stenocoris (Oryzocoris) fabricu, g. 178, terminalia, dorsal view ; 
179, theca, dorsal view ; 180, aedeagus (inflated), ventral view. 


THE LEPTOCORISINAE OF THE WORLD 63 


Body comparatively robust, pale. Head never half as long again as broad ; occipital suture 
medially concave, arms bending inwards towards sides ; paraclypeae short, about one-third of 
head, straight and of uniform thickness. Antennae with underside of basal segment pale, 
above black ; second and third grey, second slightly shorter than third, fourth tinged with red, 
pale only at base. Rostrum comparatively long, reaching apices of second coxae. Lateral 
black line faint, extending from sides of antenniferous tubercles on to collar. Pronotal disc 
pale, posteriorly with greenish tinge, black spots on posterior angles absent. Legs pale, with 


Fics. 181-184. Stenocoris (Ovyzocoris) fabricii, 2. 181, terminalia, ventral view ; 182, 
first gonopophysis, ventral view ; 183, ovipositor, ventral view ; 184, same, dorsal 
view. 


64 I. AHMAD 


femora greenish, tibiae and tarsi tinged with black. Hemielytra with anal and medial veins 
brownish black. Abdomen with dorsum, excluding connexiva, red, venter pale, tinged with 
red ; median longitudinal red line absent. Posterior margin of seventh abdominal tergum 
sinuate, medially concave ; pygophore greatly elongated, tapering posteriorly, posterior margin 
narrow, about one-sixth of width of posterior margin of seventh abdominal tergum ; posterior 
processes pointed, facing each other (Text-fig. 178). 

¢ GENITALIA : Claspers strongly curved in middle, acutely pointed and curved at apices 
(Text-fig. 177, A) ; dorsal thecal appendages absent ; ventral pair as in Text-fig. 180 ; four 
left lateral conjunctival appendages as in Text-fig. 180; frontal conjunctival appendages 
paired, minute, right slightly longer ; four right lateral conjunctival appendages with second 
sickle-shaped as in Text-fig. 180 ; dorsal appendages as in Text-fig. 180. 

9. Body length, 15:2-15:6 mm., width, 1:9-2-0 mm. Head length, 1-77-1-9 mm. (only 
paraclypeae 0:53—-0°55 mm.), width, 1-33 mm. Antennae, segment I, 2-93-3-:1 mm., II, 1-8— 
1-96 mm., III, 1-8-2-0 mm., IV, 4-0-4:2 mm. Rostrum, segment I, 1-33 mm., II, 1:33 mm., 
III, 0-66 mm., IV, 0-6 mm. Pronotum, length, 2-36-2-4 mm., width, 1-9-2-0 mm. Hind 
femora, 5:2-5:34 mm. Hind tibia, 5-32-5-6mm. Distance between ocelli, 0-46 mm., distance 
between an ocellus and an eye, 0-7 mm. 

Very similar to g in general appearance ; seventh abdominal sternum with a short median 
split, forming a lobe on either side ; lobes with concave posterior margins (Text-fig. 181). 

Q GENITALIA: First gonocoxae large, concealing eighth paratergites, triangular, apices 
facing each other (Text-fig. 181) ; intervalvular sacs absent ; spermatheca oval, with a smoothly 
curved neck and a short coiled tube (Text-fig. 177, C). 


Holotype 3, PERu: Tingo Maria, Huan, 2,200 ft., 11.x.1946 (J. C. Pallister), 
A.M.N.H. 


Paratypes. I 3, I 9, with same data as holotype, A.M.N.H., 1 3, without any 
data, from a series of filiformis Fabricius in coll. Fabricius, Zool. Mus. Copenhagen ; 
VENEZUELA : Estate of Guasdualito (Guarico), Camagua, I 9, 1923, (S. Mayeul 
Crisol), Mus. Hist. nat. Paris ; ARGENTINA: Misiones, Erv. de San Ignacio Villa, 
Lutecia, I gf, 1910 (E. R. Wagner), Chaco de Santa Fe las Garzas, near Figlas Garzas, 
20 km. to Docampo, I g, 1903 (EF. R. Wagner), Mus. Hist. nat. Paris ; BRaziL : I 9, 
1898 (Noualler), Pernambuco, Pery, I g, 1898 (Nouwalhier), Mus. Hist. nat. Paris ; 
BRITISH GUIANA : Georgetown, I 9, 6.vii.1g11, A.M.N.H. ; Honpuras: Teguci- 
galpa, 28.11.1917 (7. JaDre7) aN. 


This species is found at high altitudes and seems to be common in Peru. Its range 
extends in the north into Venezuela. Adults have been collected in April and 
October. 


This species is close to S. (O.) furcifera (Westwood) in general appearance but can 
easily be separated by the absence of red median longitudinal line on venter of abdo- 
men. 


Stenocoris (Oryzocoris) filiformis (Fabricius) comb. n. 
(Text-figs. 175, 185-190) 


Cimex filiformis Fabricius, 1775 : 727, 1781 : 375, 1787 : 308 ; Gmelin, 1789 : 2191 
Gerris filiformis (Fabricius) Fabricius, 1794 : 191, 1803 : 260 ; Stal, 1866 : 66 
Leptocorisa (s. str.) filiforymis (Fabricius) Stal, 1873 : 87 

Leptocorisa linearis Laporte, 1833 : 26 

Leptocorisa tipuloides Dallas, 1852 : 484 


65 


THE LEPTOCORISINAE OF THE WORLD 


Characterized by the long cylindrical paraclypeae, by almost rectangular pygo- 
phore, by convex posterior margin of the lobes of the 2 seventh abdominal sternum 


and by the form of 3 and @ genitalia. 
3. Body length, 14-0-15-0 mm., width, 1-8-1-9 mm. Head length, 1-9-1-3 mm. (only 
paraclypeae, 0:66 mm.), width, 1-3-1-4 mm. Antennae, segment I, 3:0-3:26 mm., II, 1-8- 


Wey FU aye 


Realy 

Vy SMV 
RAE hee 
N YOY aww) 

AS 


Ze 
- 


190 
185-186, g, 187-190, 9. 185, termin- 
187, terminalia, ventral 
; 190, 


Fics. 185-190. Stenocoris (Oryzocoris) filiformis. 
alia, dorsal view ; 186, aedeagus (inflated), ventral view ; 
188, first gonopophysis, ventral view ; 189, ovipositor, ventral view ; 


view ; 
same, dorsal view. 


66 I. AHMAD 


2-0 mm., III, 2:2-2-26 mm., IV, 3:7-4-:0 mm. Rostrum, segment I, 1:2-1-:3 mm., II, 1-2— 
1-3 mm., III, 0-6-0-66 mm., IV, 0-6-0:-66 mm. Pronotum, length, 2:3-2:4 mm., width, 
1-8-1-9 mm. Hind femora, 5-5-5-9 mm. Hind tibia, 5-9-6-:0 mm. Distance between ocelli, 
0-5-0°53 mm., distance between an ocellus and an eye, 0-66—0-7 mm. 

Body medium sized, linear, usually yellowish with greenish or sometimes reddish tinge. 
Head comparatively broad, breadth usually less than two-thirds of length ; occipital suture 
“V-shaped in middle ; paraclypeae long and cylindrical, more than one-third of head. 
Antennae usually palish brown, sometimes tinged red, second distinctly shorter than third. 
Rostrum, only reaching anterior margins of second coxae. Lateral black line well represented 
extending from sides of antenniferous tubercles on to collar. Pronotal disc pale, tinged with 
green, black spots on posterior angles absent. Legs with apices of femora sometimes reddish. 
Hemielytra with anal vein usually more prominently black than medial. Abdomen with 
dorsum pale unicoloured, sometimes with reddish tinge, venter pale ochraceous, red median longi- 
tudinal line absent. Posterior margin of seventh abdominal tergum truncate, pygophore 
curved near apex, posterior processes acutely pointed and facing each other, posterior margin 
broad, more than half of width of seventh abdominal tergum (Text-fig. 185). 

¢ GENITALIA: Claspers irregularly crossed over pygophore, curved near apex (Text-fig. 
175, A) ; dorsal thecal appendages absent ; ventral pair as in Text-fig. 186 ; four left lateral 
conjunctival appendages as in Text-fig. 186 ; frontal conjunctival appendages paired, minute, 
right slightly thicker and longer ; four right lateral conjunctival appendages, with second 
almost straight as in Text-fig. 186 ; dorsal appendages as in Text-fig. 186. 

©. Body length, 15-0-16-5 mm., width, 1:9-2-0 mm. Head length, 1-8-1-9 mm., width, 
I:3-1°4 mm., (length of only paraclypeae, 0-66-0-7 mm.). Antennae, segment I, 2-56-3-I mm., 
II, 1:86-2:06 mm., III, 2-2-2:26 mm., IV, 3:7-4-0 mm. Rostrum, 1-:16—-1:2 mm., II, 1-16— 
2:0 mm., III, 0-6-0-63 mm., IV, 0-6-0-63 mm. Pronotum, length 2:3-2:4 mm., width, 
I-9-2:0 mm. Hind femora, 5:2-5:-3 mm. Hind tibia, 5-6-5:8 mm. Distance between ocelli, 
0:5-0:56 mm., distance between an ocellus and an eye, 0-6—0-63 mm. 

Distinctly longer than 3, posterior margin of seventh abdominal sternum with a short median 
split, forming convex lobe on either side. 

@ GENITALIA : First gonocoxae broad, concealing a large basal portion of eighth paratergite 
(Text-fig. 187) ; outer margin regular, apices rounded ; intervalvular sacs absent ; sperma- 
theca flask-shaped with a distinct, smoothly curved neck as in Text-fig. 175C. 


Type material. Lectotype g, without data (“ filiformis ”’ Fabricius’ handwriting), 
Zool. Mus. Copenhagen ; syntypes, I g, I 2, with no data, Zool. Mus. Copenhagen. 


Other material. Mrxico: 19; U.S.A.: 64,109; CuBA: 14; COLOMBIA : 
26,19; BritisH GUIANA: 24,29; LEEWARD ISLANDS: 24,29; BRAZIL: 
216, 84 2: 

The range of this species extends in the east into the Leeward Islands and in the 
west into Mexico and Gorgona Island, Colombia. It is the commonest Central 
American species and in the north is found in Texas and Leesburg, Florida. Adults 
seem to be very common in August. 


This is close to S. (O.) furcifera (Westwood) and can be separated from it by the 
absence of red median longitudinal line on abdominal venter. 

Four syntypes from Fabricius’ collection have been examined ; all except one ¢ 
represent the present concept of S. (0.) filiformis (Fabricius). One $ with Fabricius’ 
own label “‘ filiformis”’ has been designated lectotype of Stenocoris (Oryzocoris) 
filiformis (Fabricius) and has been given a lectotype label. The exceptional 
represents S. (O.) fabrici sp. n. 


THE LEPTOCORISINAE OF THE WORLD 67 


Stenocoris (Oryzocoris) furcifera (Westwood) comb. n. 
(Text-figs. 176, IgI—-196) 


Leptocorisa furcifera Westwood, 1842 : 18 
Gerris nigricornis Stal, 1870 : 218 syn. n. 
Leptocorisa nigricornis (Stal) Stal, 1873 : 87 


Characterized by red median longitudinal line on abdominal venter and by the 
form of gf and 9 genitalia. 


6. Body length, 14-0-16:12 mm., width, 1:-8-2-o mm. Head length, 1-9-2-:0 mm., (length 
of paraclypeae 0-5-0'56 mm.), width, 1-3-1-4 mm. Antennae, segment I, 3-3-3-6 mm., II, 
2:20-2:26 mm., III, 2:20-2:26 mm., IV, 3:7-4:0 mm. Rostrum, segment I, 1-3-1-:33 mm., II, 
I-3-1-33 mm., III, 0-6—-0-63 mm., IV, 0:-6-0-63 mm. Pronotum, length, 2-4-2-6 mm., width, 
1-8-2-0 mm. Hind femora, 5-73-6:0 mm. Hind tibia, 6:-5-7:1 mm. Distance between 
ocelli, 0-46—0-5 mm., distance between an ocellus and an eye, 0:6—0-66 mm. 

Body usually greatly elongated, linear and usually pale unicolourous. Head about half as 
long again as broad ; occipital suture medially concave, arms curving outwards towards 
sides ; paraclypeae short, about one-third length of head, thicker at bases and thinner at apices. 
Antennae pale, with apices of each segment (except fourth) black, fourth light red, second and 
third usually subequal. Rostrum very short, usually not reaching second coxae. Lateral 
black line faint, extending from sides of antenniferous tubercles on to collar. Pronotal disc 
usually pale unicolourous, sometimes with greenish tinge posteriorly ; posterior angles without 
black spots, although sometimes trace of brownish spot present. Legs pale unicolourous. 
Hemielytra with both anal and medial veins prominently black. Abdomen with dorsum, 
excluding connexiva, red, venter palish red with a median longitudinal red line, this line some- 
times faint. Posterior margin of seventh abdominal tergum slightly medially concave ; pygo- 
phore smoothly tapering posteriorly ; posterior processes rounded, directed backward (Text-fig. 
195). 

6 GENITALIA : Claspers crossed regularly over pygophore, not curved at apices (Text-fig. 
176, A) ; dorsal thecal appendages absent, ventral pair as in Text-fig. 191 ; four left lateral 
conjunctival appendages as in Text-fig. 191 ; frontal conjunctival appendages paired, minute, 
almost of equal length ; four right lateral conjunctival appendages, with second almost straight 
as in Text-fig. 191 ; dorsal appendages as in Text-fig. ror. 

@. Body length, 17-66-18-66 mm., width, 2-0-2:26 mm. Head length, 1-86-2:0 mm., 
width, 1:4-1:-5 mm. Antennae, segment I, 3:3-3:73 mm., II, 2-2-2-:26 mm., III, 2:2-2:26 mm., 
IV, 3:3-3:8 mm. Rostrum, segment I, 1-3—1-33 mm., II, 1-3-1-33 mm., III, 0-63—-0-7 mm., IV, 
0:63-0-7 mm. Pronotum length, 2:5—2:66 mm., width, 2:0-2:26 mm. Hind femora, 5-33- 
577mm. Hind tibia, 6-0o-6:1 mm. Distance between ocelli, 0-46-0-5 mm., distance between 
an ocellus and an eye, 0-66—0-7 mm. 

Distinctly longer than g ; posterior margin of seventh abdominal sternum with a median 
split, forming concave lobe on either side (Text-fig. 192). 

Q GENITALIA : First gonocoxae comparatively narrow, leaving a portion of eighth paratergite 
unconcealed, outer margin irregular, apices truncated ; intervalvular sacs absent ; spermatheca 
large, bladder-shaped, without a distinct neck. 


Type material. Holotype 2, without data but with a label “ Br/c”’ and other 
type labels, Hope Dept., Univ. Mus. Oxford ; holotype 2 of Leptocorisa migricornis 
Stal, Brazit : Minas, Riksmus. Stockholm. 


Other material. BRraAziL: 21 gf, 16 9; BRITISH GUIANA: I ¢ ; VENEZUELA: 
i? =. Mexmco: 24,22; PANAMA (CANAL ZONE): 192; U.S.A., Florida 27 3. 


68 I. AHMAD 


This species seems to be very common in Brazil. Adults have been collected 
throughout the year. Its range extends in the north into Florida. 


This species is very close to S. (O.) filiformis (Fabricius) but can be separated by 


Fics. 191-196. Stenocoris furcifera. 191, 195, gd, 192-194, 196, 9. 191, aedeagus (in- 
flated), ventral view ; 192, terminalia, ventral view ; 1093, first gonopophysis, ventral 
view ; 194, ovipositor, dorsal view ; 195, terminalia, dorsal view ; 196, ovipositor, 
ventral view. 


THE LEPTOCORISINAE OF THE WORLD 69 


the median longitudinal red line which runs on the ventral side of adbominal seg- 
ments (but is sometimes only faintly represented) by the black anal and medial vein 
of hemielytra and by the short paraclypeae, which are thicker at base and thinner at 
apices. 


LEPTOCORISA Latreille 


Leptocorise Latreille, 1825 : 421 

Leptocorisa Latreille, 1829: 197; Guérin, 1830: 178; Westwood, 1842:18; Dallas, 
1852 : 481 ; Costa, 1863: 260; Montrousier, 1865 :227; Walker, 1871 :172; Stal, 
1872 : 54, 1873 : 86; Lethierry and Severin, 1894 : 102 ; Distant, 1901 : 335, 1902 : 409 ; 
Breddin, 1909 : 293 ; Oshanin, 1912 : 24; Breddin, 1912: 104 ; Van Duzee, 1914 : 378 ; 
China, 1924 : 237; Tillyard, 1926: 148; Miller, 1956:55; Southwood and Leston, 
1959 : 65 

Leptocorixa Berthold, 1827: 418; Kirkaldy, 1908: 123; Reuter, 1913: 78; Bergroth, 
1913 : 12, I913a : 159 

Rhabdocoris Kolenati, 1845 : 67 Syn. n. 


Type-species : Cimex acutus Thunberg, 1783. 


Body robust. Head elongated but shorter than pronotum, ocelli placed close together. 
Antennae with apices of basal segment slightly thicker. Rostrum usually short, touching 
second coxae. 

In g, posterior margin of seventh abdominal tergum truncated ; pygophore rounded, claspers 
crossed behind, made up of two components, shaft (or main stem) and blade (or bent portion) ; 
aedeagus with both (dorsal and ventral) pairs of thecal appendages ; frontal conjunctival 
appendage membranous with usually two or three sclerotized pieces ; membranous appendage 
large, usually with a terminal and a dorsal appendage. 

In 2, posterior margin of seventh abdominal sternum emarginate, entire, without a fold or 
split ; first gonocoxae large, conical in form ; intervalvular sacs at least four pairs ; sperma- 
theca usually irregular, always with a median flange. 


This genus is close to Stenocoris Burmeister but can readily be distinguished by the 
form of the claspers, which in Stenocoris consist of a single component and by emargin- 
ate posterior margin of seventh abdominal sternum (which in Stenocoris is usually 
with a median projection, indentation or split). These genera can also be separated 
by the structure of aedeagus, for in Stenocoris the frontal conjunctival appendage is 
usually sclerotized and supporting and terminal appendages are wanting. The 
median flange of spermatheca of Leptocorisa is unique. The ranges of Leptocorisa 
and Stenocoris are distinct, for Leptocorisa is confined to Oriento-Australian region 
and Stenocoris is found in Ethiopian, Nearctic and Neotropical regions. From 
Bloeteocoris this genus can be separated by its normal genae and by other characters 
in the genitalia (see page II5). 

The preservation of the name Leptocorisa Latreille and the suppression of Lepto- 
corixa Berthold are the subjects of an application to the International Commission 
on Zoological Nomenclature (China and Ahmad, 1963). 


KeEy TO SPECIES 


t Rostrum long, always extending beyond the apices of second coxae, sometimes 
reaching third coxae . : : : : : : 
— Rostrum short, never extending beyond second coxae. é : : : : 5 


7° 


I 


I. AHMAD 


Lateral black line very prominent, extending from the bases of the antenniferous 
tubercles for at least three-quarters of the length of the propleuron ; posterior of 


disc of pronotum with black punctures . . : : - 3 
Lateral black line faint, never extending beyond the prone eer ; disc of prono- 
tum never with black punctures, sometimes with greenish tinge : 4 


Central black line on venter of abdomen present, extending from the first = oath 
segments ; rostrum usually extending halfway between second and third coxae, 
never touching third, basal quarters of apical segments of antennae never pale 
(pygophore almost of square shape with thin lobes of ninth segment (Text-fig. 342), 
blades of the claspers long (Text-fig. 345), ventral thecal appendages asymmetrical 
(Text-fig. 344) ; first gonocoxae large with truncated broad apices (Text-fig. 251) ; 
spermatheca large, fusiform with a rounded smooth neck and a short tube (Text- 
fig2 237) : : . solomonensis (p. 104) 
Central ees line on vente of aeaomen Spear: ee usually reaching third 
coxae ; basal quarters of apical segments of antennae pale. (Pygophore rounded, 
with thick and rounded lobes of ninth segments (Text-fig. 291) ; blades of claspers 
short (Text-fig. 200) ; ventral thecal appendages symmetrical (Text-fig. 293) ; 
first gonocoxae small, with rounded lobe-like apices (Text-fig. 242) ; spermatheca 
elongated and thin, with a long curved neck, giving a small knob near its apex 
(Text-fig. 228)) . : : ; discoidalis (p. 87) 
Paraclypeae remarkably siodeated. ticsel aad peut (Text-fig. 322) ; lateral 
black line only faintly represented by two brownish black dots, one on sides of 
antenniferous tubercles and the other on collar. (Ventral thecal appendages 
symmetrical (Text-fig. 325) ; right lateral conjunctival appendages three in 
number (Text-fig. 324) ; dorsal membranous appendage with a pointed hooked 
terminal and a short curved dorsal appendage (Text-fig. 324) ; spermatheca witha 
broad median flange and a tube which is twisted at the base (Text-fig. 229)) 
palawanensis (p. 98) 
Paraclypeae short, thick at base ; lateral black line represented on sides of antenni- 
ferous tubercles, extending back to the sides of collar. (Ventral thecal appendages 
asymmetrical (Text-fig. 337) ; right lateral conjunctival appendages two in 
number (Text-fig. 338) dorsal membranous appendage with a bulb-like terminal 
and a bilobed dorsal appendage (Text-fig. 338) ; spermatheca short, flask-shaped 
with a transverse neck (Text-fig. 238)) . : : sakdapolrakae (p. 102) 
Posterior third of dorsum of abdomen black, apical Seements of antennae pale both 
at bases and apices; paraclypeae acutely pointed, claspers narrow at base, 
strongly and abliquely truncate (Text-fig. 203). Ventral thecal appendages 
spiny (Text-fig. 287) ; spermatheca short, twisted at apex (Text-fig. 231) costalis (p. 85) 
Dorsum of abdomen uniformly pale, brown or ochraceous ; apical antennal seg- 
ments never pale at both bases and apices ; paraclypeae never acutely pointed 6 
Hemielytra, except costal margins above the membrane, dark black, claspers 
extending well beyond pygophore, with broad truncated apices (Text-fig. 273). 
(Second right lateral conjunctival appendage bifurcated oe 216,F) ; sperm- 


atheca linear and thin (Text-fig. 230)) . : : : biguttata (p. 80) 
Hemielytra usually unicolourous, pale or ochraceous, never dark black as above, 

claspers never extending beyond pygophore, apex never broadly truncated as above 7 
Body short, length not exceeding 12 mm. . : - ‘ : : : : 8 
Body long, length exceeding 14 mm. : : : nfo) 


Paraclypeae long, somewhat aoe sacle repair (fee fig. 296) ; claspers 
usually extending slightly beyond pygophore, apices pointed. (Second right 
lateral conjunctival appendage short, thicker in middle and pointed at apex 
(Text-fig. 220) ; spermatheca below flange elongated and pointed (Text-fig. 234)) 

lepida (p. 88) 


THE LEPTOCORISINAE OF THE WORLD 


— Paraclypeae short, usually blunt, either thicker at base or in middle (Text-fig. 309 
or 329) ; claspers never extending posterior margin of pygophore, curved and 
rounded 7 : : - : : : : . : ; : 

9 Paraclypeae thin at base and thicker in middle (Text-fig. 309) ; in g, posterior 
margin of seventh abdominal tergum with a slight truncated median projection, 


Fics. 197-205. Leptocorisa spp., claspers, outer and inner views. 197, ovatorius ; 
acuta ; 199, chinensis ; 200, discoidalis ; 201, palawanensis ; 202, biguttata ; 
costalis ; 204, tagalica ; 205, luzonensis. 


198, 
203, 


iF | 


72 


Io 


Il 


I2 


I. AHMAD 


pygophore rounded (Text-fig. 308). (Dorsal membranous appendage of aedeagus 
with a flat pointed (first terminal appendage) appendage at base of terminal 
appendage (Text-fig. 311) ; first gonocoxae Sees and almost pointed (Text- 
fig. 250)) . : . luzonica (p. 92) 
Paraclypeae eee at Eee Fad uaeeier in aicille 7 in 3, posterior margin of 
seventh abdominal tergum truncated, not produced as above; pygophore 
elongated (Text-fig. 332) ; (dorsal membranous appendage with an unsclerotized 
triangular appendage (first terminal appendage) at the base of terminal appendage 
(Text-fig. 331) ; first gonocoxae broad with rounded apices (Text-fig. 249)) 
pseudolepida (p. 100) 


Apices of femora and bases of tibiae black, two black spots behind ocelli \ = rat 
Legs usually pale unicolourous, sometimes with blackish tinge but specially apices 
of femora and bases of tibiae never black ; no black spots behind ocelli_ . 12 


Body pale ochraceous ; claspers with very long cylindrical and pointed blades 
(Text-fig. 204). (Right lateral conjunctival appendages three in number (Text- 
fig. 350) ; terminal appendage flat (Text-fig. 350) ; in Q, lateral posterior margin 
of seventh abdominal sternum giving tiny backward projections (Text-fig. 351) ; 
spermatheca elongated and pointed (Text-fig. 232)) . : . tagalica (p. 106) 
Body light brown, claspers with a short and pointed blade (Text. fig. 205) (only one 
bilobed right lateral conjunctival appendage (Text-fig. 305) ; terminal appendage 
triangular (Text-fig. 305) ; in 9, lateral posterior margin of seventh abdominal 
sternum without tiny backward projections (Text-fig. 306) ; spermatheca balloon- 
shaped, blunt, with a tube of irregular thickness (Text-fig. 233)) luzonensis (p. 90) 
Body robust, usually with brownish black ventro-lateral spots on abdomen ; 
claspers wide at base with extremities curved and tapering to a point (Text-fig. 


“WwW ¢.C0 


Fics. 206-210. Leptocorisa spp., claspers, outer and inner views. 206, lepida ; 
207, pseudolepida ; 208, luzonica ; 209, solomonensis ; 210, sakdapolrakae. 


THE LEPTOCORISINAE OF THE WORLD 


0:25 mm. 


OV 2 2187/7214 Y215 26 2h, 218 


Fics. 211-218. Leptocorisa spp., aedeagal appendages, outer view; A, frontal append- 


age ;_ B, dorsal conjunctival appendage ; C, terminal appendage ;_ D, ventral thecal 
appendage ; E, left lateral conjunctival appendage ; F, second right lateral con- 
junctival appendage. 211, ovatovius ; 212, acuta; 213, chinensis ; 214, discoidalis ; 
215, palawanensis ; 216, biguttata ; 217, costalis ; 218, tagalica. 


73 


74 I. AHMAD 


197). (Second right lateral conjunctival appendage bilobed (Text-fig. 211, F) ; 

in 9, posterior margin of seventh abdominal sternum deeply emarginate (Text- 

fig. 315), first gonocoxae elongated, giving a tiny lobe at apices (Text-fig. 239)) 
oratorius 


2AD 220) 22) DDD 2223 i204 


Fics. 219-224. Leptocorisa spp., aedeagal appendages, lettering as in figs. 211-218. 


(p. 94) 


219, 


luzonensis ; 220, lepida; 221, pseudolepida ; 222, luzonica; 223, solomonensis ; 


224, sakdapolrakae. 


THE LEPTOCORISINAE OF THE WORLD 75 
— Body more cylindrical, without ventro-lateral spots on abdomen ; claspers never 
pointed ; second right lateral conjunctival appendage never bilobed. : : 13 
13 Basal joint of antennae usually pale unicoloured, lateral black line represented by 
only a black dot on the sides of collar ; claspers bifid at apices (Text-fig. 198). 
(Second right lateral conjunctival appendage trilobed (Text-fig. 212, F) ; in 9, 
posterior margin of seventh abdominal sternum with a pointed median projection 
(Text-fig. 269) ; spermatheca irregular (Text-fig. 226)) . ‘ ‘ acuta (p. 78) 
0-5 mm. 
Fics. 225-238. Leptocovisa spp., spermatheca, ventral view. 225, ovatorius ; 226, 
acuta ; 227, chinensis ; 228, discoidalis ; 229, palawanensis ; 230, biguttata; 231, 
costalis ; 232, tagalica; 233, luzonensis; 234, lepida; 235, pseudolepida ; 236 


luzonica ; 237, solomonensis ; 238, sakdapolrakae. 


76 I. AHMAD 


— Basal joint of antennae dark brown or black ; lateral black line extending from 
bases of antenniferous tubercles to sides of collar, apices of claspers truncated 
(Text-fig. 199). (Second right lateral conjunctival appendage with a single lobe 
(Text-fig. 213, F) ; in 9, posterior margin of seventh abdominal sternum with a 
rounded median projection (Text-fig. 280) ; spermatheca thinner in middle, 
almost divided into two compartments (Text-fig. 227)) . ; chinensis (p. 82) 


Fics. 239-248. Leptocovisa spp., first gonocoxa, ventral view. 239, ovatovius ; 240, 
acuta ; 241, chinensis ; 242, discoidalis ; 243, palawanensis ; 244, biguttata; 245, 
costalis ; 246, tagalica ; 247, luzonensis ; 248, lepida. 


THE LEPTOCORISINAE OF THE WORLD 


Fics. 249-266. Leptocorisa spp., 249-252, first gonocoxa, ventral view ; 


77 


253-266, first 


gonopophysis, ventral view. 249, pseudolepida ; 250, luzonica ; 251, solomonensis ; 
252, sakdapolrakae ; 253, ovatovius ; 254, acuta; 255, chinensis ; 256, discoidalis ; 
257, palawanensis ; 258, biguttata; 259, costalis ; 260, tagalica; 261, luzonensis ; 


262, lepida; 263, pseudolepida ; 264, luzonica; 265, solomonensis ; 
polrakae. 


266, sakda- 


78 I. AHMAD 


Leptocorisa acuta (Thunberg) 
(Text-figs. 198, 212, 226, 240, 254, 267-272) 


Cimex acutus Thunberg, 1783 : 34 

Leptocorisa (Rhabdocoris) acuta (Thunberg) Stal, 1873 : 86 
Cimex angustata Fabricius, 1787 : 308 

Cimex angustus Fabricius ; Gmelin, 1789 : 2193 

Gerris angustata (Fabricius) Fabricius, 1803 : 262 

Gerris varicornis Fabricius, 1803 : 260 syn. n. 

Leptocorisa flavida Guérin, 1830 : 178 syn. n. 


\ { ’ 
-f ' 
repeal eielesaaae: SU 
Nye Patel eV 


yo Wes 
We 7 F 


0:25 mm. 


Fics. 267-272. Leptocorisa acuta, 267, 268, 270, g, 269, 271, 272 9. 


267, terminalia, 


dorsal view ; 268, aedeagus (inflated), ventral view ; 269, terminalia, ventral view ; 
270, theca, dorsal view ; 271, ovipositor, ventral view ; 272, same, dorsal view. 


THE LEPTOCORISINAE OF THE WORLD 79 


Coreus varicornis (Fabricius) Burmeister, 1834 : 298 

Myodocha varicornis (Fabricius) Burmeister, 1835 : 325 

Leptocorisa varicornis (Fabricius) Dallas, 1852 : 484 

Leptocorisa (Rhabdocoris) varicornis (Fabricius) Stal, 1873 : 86 ; China, 1924 : 237 


Characterized by claspers which are bifurcated at their apices and by the form of 
the ¢ and @ genitalia. 


6. Body length, 15-0-16-0 mm., width, 2-3-2-4 mm. Head length, 1-8-2-0 mm., width, 
I-4-I'5 mm. Antennae, segment I, 4-0-5-0 mm., II, 2:7-2-38 mm., III, 3-0-3-4 mm., IV, 
5°7-5°9 mm. Rostrum, segment I, 1-4-1-6 mm., II, 1-4-1:6 mm., III, 0-7-0o-9 mm., IV, 
0-7-0:'9mm. Pronotum, length, 2-4—2-5 mm., width, 2-3-2-4 mm. Hind femora, 6-0o—-6:2 mm. 
Hind tibia, 6-8-7-0 mm. Distance between ocelli, 0-45-0-5 mm., distance between an ocellus 
and an eye, 0:7-0'75 mm. 

Body cylindrical, palish brown. Head elongated, pale ; occipital suture medially deeply 
concave ; paraclypeae long and pointed. Antennae uniformly pale, sometimes tinged with 
brownish black. Rostrum short, hardly reaching apices of second coxae. Lateral black line 
represented by only a black dot on either side of collar. Pronotal disc with whitish margins, 
with black spots on posterior angles. Legs uniformly pale, sometimes slightly darker. Pos- 
terior margin of seventh abdominal tergum truncate ; pygophore rounded and small (Text- 
fig. 267). 

6 GENITALIA: Claspers bifid at apices (Text-fig. 198) ; dorsal and ventral pairs of thecal 
appendages present ; ventral thecal appendages symmetrical as in Text-fig. 268 ; left lateral 
conjunctival appendage reduced, spine-like ; frontal conjunctival appendage with a distinct 
posterior sclerite ; two right lateral conjunctival appendages as in Text-fig. 268, second tri- 
lobed ; dorsal membranous appendage with a pointed terminal and bilobed dorsal conjunctival 
appendages as in Text-fig. 268. 

2. Very similar to J in colour and measurements ; seventh abdominal sternum emarginate, 
with a very small median triangular projection (Text-fig. 269). 

Q GENITALIA : First gonocoxae small, outer margins sinuate, apices rounded (Text-fig. 240) ; 
six pairs of intervalvular sacs (Text-fig. 272) ; spermatheca irregular, with a median flange and 
a long coiled tube (Text-fig. 226). 


Type material. Holotype of Cimex acutus Thunberg, without any data (abdomen 
and antennal segments missing), Zool. Inst., Uppsala ; neotypeg, MALAYA : “ Cimex 
acutus Thunb. ”’ “ with Stal’s label’’ “‘ sec. Mus. Thunb. ’’, Riksmus. Stockholm ; 
holotype g of Gerris varicornis Fabricius, without any data (Jrassquebana, Dalldorff. 
Mus. Jondlund), Zool. Mus. Copenhagen ; holotype ¢ of Gerris angustata Fabricius, 
without data, labelled “‘ angustata ’’, Zool. Mus. Copenhagen. 


Other material. INDIA: 414, 239; PAKISTAN: 24,19; BuRMA: 34,29; 
BHUTAN : 21 ¢, 1592; VIETNAM: 113 6, 114 2; Formosa: 1 9; HONGKONG : 
ie © 9 >.. BAILAND :..317-4,.E19 2 ; Manaya : .12.6,.8.9; SUMATRA: 16; 
SARAWAK : 17 g, 11 2; NORTH BorRNEO: 187 g, 207 2; PHILIPPINE ISLANDS : 
46,42; NEw GUINEA: 218 gf, 110 9 ; NEW BriTAIN: I ¢ ; NEW IRELAND : 
18g, 15 2 ; AUSTRALIA : 554,569; SoLomon ISLANDs : 544, 25 2 ; NEW CALE- 
DONIA : 26,292; NEw HEBRIDES: 44,392; Fij1 ISLANDS: 118 fg, 137 9; Sa- 
MOAN ISLANDS : 194 4, I15 9. 


This species is a major pest of paddy crops and is found in all rice growing regions 
of the Far East countries. It is very common in India (U.P.) (and here it is known as 
“ Gandhi bug ”’), Sarawak, and New Guinea. Its range extends in the west into 


80 I. AHMAD 


India and in the east right into Samoa, New Caledonia and the Fiji Islands and is 
found throughout the year. No specimens have been recorded from Java and Ceylon. 
Its biology and life history have been studied in India by Lefroy (1908) and Akbar 
(1958) under the name of Leptocorisa varicornis (Fabricius). 


This species, with L. orvatorius (Fabricius) and L. chinensis Dallas, forms a group 
of widely distributed pests of paddy crops in the Far East, but it can easily be 
separated from them and in fact from all other species of Leptocorisa in the § by the 
claspers, which are bifurcated at apices and in the 2 by a very small median triangular 
projection of the seventh abdominal sternum. 

Stal redescribed acutus Thunberg and varicornis Fabricius as distinct species. 
During the present study the types of both the species have been examined, and 
were found to represent the same species. In view of the damaged condition of the 
supposed type of acutus Thunberg in the Zoological Institute, Uppsala, and with the 
agreement of Dr. Kullenberg (Mr. Hedstrom), a male specimen in the Riksmus., 
Stockholm from the type locality (Malacca) and bearing Stal’s label “‘ Cimex acutus 
Thunb. Mus. Thunb.” has been designated neotype and has been given a neotype 
label. 


Leptocorisa biguttata Walker 
(Text-figs. 202, 216, 230, 244, 258 and 273-277) 


Leptocorisa biguttata Walker, 1871 : 174 
Leptocorisa biguttata Walker; Distant, 1901 : 372 [as synonym of L. costalis Herrich Schaffer] 


Characterized by hemielytra which, excluding the costal margins, are black, by 
lateral black line extending from sides of antenniferous tubercles on to pronotal 
collar, by short paraclypeae, which are thick at bases and pointed at apices, and 
by the form of 3 and 9 genitalia. 


3g. Body length, 16-0-17-0 mm., width, 2-2-2-4 mm. Head length, 2-2-2:3 mm., width, 
1-7-1-8 mm. Antennae, segment I, 4-8-4:9 mm., rest missing. Rostrum, segment I, 1-6— 
1-7 mm., II, 1-7-1-8 mm., III, 0-9-1-0 mm., IV, 0-8 mm. Pronotum, length, 2-7-2-8 mm., 
width, 2:2-2-4 mm. Hind femora, 7-0 mm. Hind tibia, 7-:0-7-2 mm. Distance between 
ocelli, 0-5—0-55 mm., distance between an ocellus and an eye, 0:95 mm. 

Body elongated, robust and brownish black. Head elongated, yellowish brown with occipital 
suture medially smoothly concave ; paraclypeae short, very thick at bases and pointed at 
apices. Rostrum short, reaching apices of second coxae. Lateral black line extending from 
sides of antenniferous tubercles on to collar. Pronotal disc yellowish brown, posterior angle 
with black spots. Hemielytra, excluding costal margin, dark black. Legs brownish black, 
hind femora darker. Posterior margin of seventh abdominal tergum truncated, medially very 
slightly convex ; pygophore almost rectangular (Text-fig. 273). 

6 GENITALIA: Claspers extending well beyond the posterior margin of pygophore, apices 
broadly truncated (Text-fig. 202) ; both dorsal and ventral thecal appendages present as in 
Text-fig. 274, symmetrical ; left lateral conjunctival appendage short, pointed and curved like 
an arch ; frontal conjunctival appendages with three sclerotized pieces as in Text-fig. 274 ; two 
right lateral conjunctival appendages, second thick at base and bifurcated at apex (Text-fig. 
216, F) ; dorsal membranous appendage with a laterally pointed terminal, and a bilobed dorsal 
appendage (Text-fig. 274). 

9. Body length, 17:-5-17°8 mm., width, 2-4-2-5 mm. Head length, 2-:2-2:4 mm., width, 
1:8-1-9mm. Antennae, segment I, 5-omm., II, 3-omm., III, 3:3 mm., IV, missing. Rostrum, 


THE LEPTOCORISINAE OF THE WORLD 81 


segment I, 1-5-1-6 mm., II, 1-7-1-3 mm., III, 0-8-o-9 mm., IV, 0-7-0-8 mm. Pronotum, 
length, 2:8-2-9 mm., width, 2-4-2-5 mm. Hind femora, 6-9-7-0mm. Hind tibia, 7-0-7: mm. 
Distance between ocelli, 0-5 mm., distance between an ocellus and an eye, 0-9 mm. 

Slightly longer than 3 with usually less dark hemielytra, posterior margin of seventh abdominal 
sternum emarginate, with a very small rounded projection in the middle (Text-fig. 275). 


0:25 mm. 


Fics. 273-277. Leptocorisa biguttata, 273, 274, 3, 275-277, @- 273, terminalia, dorsal 
view ; 274, aedeagus (inflated), ventral view ; 275, terminalia, ventral view ; 276, 
ovipositor, dorsal view ; 277, same, ventral view. 


82 I. AHMAD 


Q GENITALIA: First gonocoxae broad at bases, apices pointed (Text-fig. 244) ; fifteen 
pairs of intervalvular sacs (Text-fig. 276) ; spermatheca elongated and thin, with a coiled tube 
and a median flange as in Text-fig. 230. 


Type material. Holotype 2, CELEBEs : Gilolo (Wallace Coll.), B.M.(N.H.). 


Other material. MaALaya: 134,14 9; PHILIPPINE ISLANDS: 44,29; SARA- 
WAK : 254,109; CELEBES: 69,69; HALMAHERA ISLAND: 24,1 9. 


The range of this species extends in the west into Malaya and in the east as far 
as Halmahera Island east of the Celebes. This species has also been reported on 
paddy and on grasses, Panicum barbatum. It seems to be very common in the early 
part of the year, December, January and February. 


This species is the closest ally of L. costalis (Herrich Schaffer) but can easily be 
separated from this species by the apical antennal segments, which are pale only 
at bases (pale both at bases and at apices in costalis), by the dorsum of abdomen which 
is yellowish brown, unicolourous (posterior one-third black in costalis) and by the 
short paraclypeae, which are thicker at bases and pointed at apices (cylindrical, 
pointed and knob-like at apices in costalis). Its broadly truncated claspers extending 
well beyond the posterior margin of pygophore and linear spermatheca are its 
peculiarities. 

Distant synonymized this species with L. costalis (Herrich Schaffer) and conse- 
quently this species was not included by China (1924) in his key for the Oriental 
species of Leptocorisa ; but the apical antennal segments, abdominal colour and gf and 
2 genitalia show it to be distinct (see above). 


Leptocorisa chinensis Dallas 
(Text-figs. 199, 213, 227, 241, 255 and 278-283) 


Leptocorisa chinensis Dallas, 1852 : 483 

Leptocorisa chinensis Dallas ; Stal, 1873 : 86 [as synonym of Leptocorisa (Rhabdocoris) varicornis 
Fabricius] 

Leptocorisa nitidula Breddin, 1912 : 107 Syn. n. 

Leptocorisa corbetti China, 1924 : 237 Syn. n. 


Characterized by the lateral black line, which is represented by two dots on the 
posterior sides of head and on the pronotal collar, by the basal antennal segments 
which are black at apices and by the form of 3 and 9 genitalia. 


6. Body length, 17-:3-17:3 mm., width, 2:7-2-9 mm. Head length, 2-3 mm., width, 
1-6 mm. Antennae, segment I, 4-:2-4-3 mm., II, 2-8-2-9 mm., III, 3-0-3-3 mm., IV, 4-4— 
46mm. Rostrum, segment I, 1-6-1-8 mm., II, 1-7-1-8 mm., III, 1-o-1-:1 mm., IV, 1-o—-1-1 mm. 
Pronotum, length, 3-0-3-:2 mm., width, 2-7-2-9 mm. Hind femora, 6-6-6-9 mm. Hind tibia, 
7-0-7:'I mm. Distance between ocelli, 0-6 mm., distance between an ocellus and an eye, 
o-8 mm. 

Body robust, pale. Head elongated, occipital suture medially concave ; paraclypeae 
elongated, and rounded at apices. Antennae pale, tinged with black. Rostrum short, reaching 
apices of second coxae. Lateral black line usually represented by two black dots on posterior 
sides of head and on pronotal collar. Pronotal disc pale unicolorous, black spots on posterior 
angles absent or faint. Legs pale, sometimes tinged with black. Posterior margin of seventh 
abdominal tergum truncated, pygophore large, rounded, (Text-fig. 278). 


THE LEPTOCORISINAE OF THE WORLD 83 


6 GENITALIA : Claspers truncated at apices (Text-fig. 199) ; both dorsal and ventral thecal 
appendages present, symmetrical (Text-fig. 281) ; left lateral conjunctival appendage elongated, 
pointed and curved in middle ; frontal conjunctival appendage with two sclerotized pieces 
(Text-fig. 281) ; two right lateral conjunctival appendages, second with one lobe (Text-fig. 


213,F) ; dorsal membranous appendage with a cap-shaped terminal appendage and a dorsal 
appendage as in Text-fig. 281. 


ee AD 
ryt 
a darnitty 
As) A Oe Se 


ui 


Fics. 278-283. Leptocorisa chinensis, 278, 279, 281, 3, 280, 282, 283, 2. 
dorsal view ; 279, claspers, outer and inner view ; 
281, aedeagus (inflated), ventral view ; 
view. 


278, terminalia, 
280, terminalia, ventral view ; 
282, ovipositor, ventral view ; 283,same, dorsal 


84 I. AHMAD 


9. Body length, 16-8-18- mm., width, 2-7-2-9 mm. Head length, 2-7 mm., width, 
1-7 mm. Antennae, segment I, 4-1-4:8 mm., II, 2-:7-2-9 mm., III, 3-1-3-7 mm., IV, 4:3- 
4°5 mm. Rostrum, segment I, 1-7-1-8 mm., IJ, 1-5-1-9 mm., III, 0o-8-1-0 mm., IV, o0-7— 
I-o mm. Pronotum, length, 2-9-3:1 mm., width, 2:7-2-9 mm. Hind femora, 6-7-6-8 mm. 
Hind tibia, 7-0-7: mm. Distance between ocelli, 0-5-0-6 mm., distance between an ocellus 
and an eye, 0-8—o-9 mm. 

Very similar to g in general appearance ; posterior margin of seventh abdominal sternum 
with a minute median rounded projection (Text-fig. 280). 

Q GENITALIA: First gonocoxae large, rounded at apices (Text-fig. 241) ; seven pairs of 
intervalvular sacs (Text-fig. 283) ; spermatheca elongated, almost divided in middle, with a 
median flange (Text-fig. 227). 

Type material. Neotype g, CuinA : Foochow (Fuchou), 10—-14.vii.1936 (M. S. 
Yang), B.M.(N.H.) ; holotype 2 of Leptocorisa mitidula Breddin, CHINA: Pingh’ 
siang, near Kwangsi (Coll. Breddin, labelled ‘‘ Leptocorisa nitidula Bredd.’’), 
Dtsch. Entom. Inst., Berlin ; holotype 3 of Leptocorisa corbetti China, MALAYA : 
Kuala Lumpur, 1933 (G. H. Corbett), B.M.(N.H.) ; paratypes, 1g, 1 9, of L. mtidula, 
with data as holotype, Dtsch. Entom. Inst., Berlin; paratypes, 1 g, 6 Q, of 
L. corbettt, with data as holotype, B.M.(N.H.). 


Other material. CHINA: 244, 23 9; Bonin ISLAND: 1 ¢; FORMOSA: 14; 
THAILAND : 246; VIETNAM: 16; PHILIPPINE ISLANDS: 10 4, 6 2; BHUTAN: 
406,352; Mataya: 104, 102; SUMATRA: 546,29; PALAU ISLAND: 1 Q. 


The range of this species extends in the north into Bhutan and China, in the south 
into Sumatra and in the east into Malaya. This species has not been recorded from 
Australian region. The adults have been collected throughout the year ; this 
species has also been recorded on grasses and four specimens were found in maize 
fields. 


This species is very close to L. ovatorius (Fabricius) and L. acuta (Thunberg) but 
can easily be separated from them by the lateral black line, which is represented by 
two brownish black dots on the sides of the head and pronotal collar. The outer 
side of basal antennal segment is usually black and the claspers are truncated at 
apices, whereas in acuta the lateral black line is represented by a black dot on the 
sides of collar and in oratorius lateral black line is absent. In ovatoriws claspers 
taper to a point posteriorly and in acuta these are bifurcated at apices. 

In the original description, Dallas (1852) mentions ‘‘ Head impunctate with a black 
line on each side. Antennae, with the basal joint fulvous black externally at the 
apex, legs pale fulvous, tibia sometimes black at the base and apex, tarsi black at 
the apex’. The two specimens recorded by Dallas and presented by G. T. Lay can 
not now be found in the British Museum (Natural History) and are presumably lost. 
This species was consequently unknown to Dr. W. E. China and was not discussed 
in his 1924 revision. A number of specimens agreeing with Dallas’s description 
have been found in the unnamed accessions in the British Museum. It is proposed to 
designate one of these (¢ B.M. 1948-548) from the type locality as neotype and this 
has been given a neotype label. 

Stal (1873) synonymized this species with varicornis Fabricius. The synonymy 
has been found incorrect, the two species being easily separable (see above). 


THE LEPTOCORISINAE OF THE WORLD 85 


Leptocorisa costalis (Herrich Schaffer) 
(Text-figs. 203, 217, 231, 245, 259 and 284-280) 


Myodochus costalis Herrich Schaffer, 1848 : 96 
Leptocorisa (Rhabdocoris) costalis (Herrich Schaffer) Stal, 1873 : 86 ; China, 1924 : 238 


Characterized by the apical segments of antennae, which are pale, both at bases 
and apices, by the basal third of the dorsum of the abdomen, which is black, by the 
paraclypeae which are cylindrical, and pointed, knob-like at apices, and by g and 2 
genitalia. 


0:25 mm. 


Fics. 284-289. Leptocorisa costalis, 284, 285, 287, dg, 286, 288, 289, 9. 284, terminalia, 
dorsal view; 285, claspers, inner view; 286, terminalia, ventral view; 287, 
aedeagus (inflated), ventral view ; 288, ovipositor, ventral view ; 289, same, dorsal 
view. 


86 I. AHMAD 


6. Body length, 14-7-15-:2 mm., width, 2-1-2-6mm. Head length, 2-0-2-3 mm., (length of 
paraclypeae o-7-0-°8 mm.), width, 1-5-1-7 mm. Antennae, segment I, 4-0-4-7 mm., II, 2-5— 
2-7mm., III, 2-8-2-9mm.,1V,5-0-5-4mm. Rostrum, segment I, 1-2-1-4mm., II, 1-2-1-4mm., 
III, 0-7-0-9 mm., IV, 0-7-0:-8 mm. Pronotum, length, 2-4—-2-:7 mm., width, 2-1-2-6 mm. 
Hind femora, 6-0-6:-4 mm. Hind tibia, 6-4-6-7 mm. Distance between ocelli, 0-4-0-5 mm., 
distance between an ocellus and an eye, 0-7—0-8 mm. 

Body cylindrical, light brown, tinged with black. Head elongated, occipital suture almost 
straight, paraclypeae remarkably elongated and pointed knob-like at apices. Antennae 
brownish black, with apical segment pale, both at bases and at apices. Rostrum short, usually 
reaching apices of second coxae. Lateral black line prominent, extending from sides of antenni- 
ferous tubercles to anterior third of propleuron. Pronotal disc palish brown, sometimes 
anteriorly blackish, posterior angles with black spots. Legs brownish black, with basal half of 
femora pale. Hemielytra, excluding costal margins, black. Posterior margin of seventh 
abdominal tergum almost truncated, slightly sinuate ; pygophore rounded (Text-fig. 284). 

6 GENITALIA : Claspers curved and acutely pointed, blades obliquely truncated (Text-fig. 
203) ; both dorsal and ventral pairs of thecal appendages present, with ventral appendages, 
asymmetrical, spiny and sac-like (Text-fig. 287) ; three left lateral conjunctival appendages 
as in Text-fig. 287 ; frontal conjunctival appendages with three sclerotized pieces (Text-fig. 
217,A) ; only one thin, curved and blunt right lateral conjunctival appendage present, (Text-fig. 
217,F), membranous appendage without a terminal or a dorsal appendage (Text-fig. 287). 

@. Body length, 15-0-16-0 mm., width, 2:2-2-4 mm. Head length, 2-1-2:-5 mm., width, 
I-6-1°8 mm. Antennal segment I, 4:5-4-7 mm., II, 2-4-2:-6 mm., III, 2-7-2:3 mm., IV, 
5:2-5'4 mm. Rostrum, segment I, 1:5-1:6 mm., II, 1-4-1-6 mm., III, 0-7-0-9 mm., IV, 
o:7-0-8 mm. Pronotum, length, 2:5-2-8 mm., width, 2-4-2:7 mm. Hind femora, 6-26— 
6-4 mm. Hind tibia, 6-4-6-6 mm. Distance between ocelli, 0-5 mm., distance between an 
ocellus and an eye, 0:8-0-85 mm. 

Very similar to gf in general appearance ; posterior margin of seventh abdominal sternum 
sinuately emarginate (Text-fig. 286). 

@ GENITALIA : First gonocoxae triangular, apices pointed (Text-fig. 245) ; sixteen pairs of 
intervalvular sacs (Text-fig. 289) ; spermatheca twisted at apices, with a median flange and a 
long coiled tube of variable thickness. 


Type material. Neotype 3, JAvA : Carnegie Mus. 


Other material. VIETNAM: 13 6, 17 9; SINGAPORE: 2 4, I 9; THAILAND: 
2g; Matava: 104,89; JAVA: 2g; SUMATRA: 104,129 ; SARAWAK: 264, 
16 9 ; NorTtH BorNEO: 18 g, 202; BORNEO: I 4; PHILIPPINE ISLANDS: 26. 


The range of this species extends in the west into Sumatra, Thailand and Malaya 
and in the east into Borneo and the Philippine Islands. It seems to be very common 
in Borneo. Adults have been collected almost throughout the year. 


It is a completely isolated species in the genus on the basis of the aedeagus. 
However it is close to L. biguwttata Walker in colouration of the hemielytra, but the 
two can easily be separated from each other by the characters of apical antennal 
segment, colouration of abdomen and shape of claspers (see above). 

In the original description Herrich Schaffer (1848) stated “‘ M. virescens antennis 
(articulo 4 basi albo) linea lateralis capitis, angulis posticis thoracis, scutello et 
elytris intus nigris, tibiis tarsique fuscis (Fig. 864)’. Dr. W. Forster (Zoologische 
Samlung des Bayerischen Staates) has kindly informed the author that in 1944 the 
type of Myodochus costalis Herrich Schaffer was destroyed as the result of an air 
raid, but a number of specimens agreeing with Herrich Schiaffer’s description and 


THE LEPTOCORISINAE OF THE WORLD 87 


figure were identified. A male specimen from Java (the type locality) from the 
Carnegie Museum (No. 8147) has been designated neotype of Leptocorisa costalis 
(Herrich Schaffer) and has been given a neotype label. 


Leptocorisa discoidalis Walker 
(Text-figs. 200, 214, 228, 242, 256 and 290—295) 


Leptocorisa discoidalis Walker, 1871 : 173 
Leptocorisa (Rhabdocoris) discoidalis (Walker) China, 1924 : 239 


’ i 
the way time! 
a) 


0:25 mm. 


Fics. 290-295. Leptocorisa discoidalis, 290, 291, 293, dg, 292, 294, 295, 9. 290, head, 
dorsal view; 291, terminalia, dorsal view ; 292, terminalia, ventral view ; 293, 
aedeagus (inflated), ventral view ; 294, ovipositor, dorsal view ; 295, same, ventral 
view. 


88 I. AHMAD 


Characterized by long rostrum, which reaches third coxae and by the form of ¢ 
and © genitalia. 


¢. Body length, 15-5-16-5 mm., width, 2:2-3-0 mm. Head length, 1:7-1-9 mm., width, 
I-5-I'7 mm. Antennae, segment I, 4:2-4:3 mm., II, 2-9-3:0 mm., III, 3:5-3-6 mm., IV, 
5:2-5:'3mm. Rostrum, segmentI,1-9mm.,II,1-9mm.,III,o-9mm.,IV,1-omm. Pronotum, 
length, 2:4-2:5 mm., width, 2-2-2-3 mm. Hind femora, 6-5-6:7mm. Hind tibia, 6-8-6-9 mm. 
Distance between ocelli, 0:44-0:45 mm., distance between an ocellus and an eye, 0:7-0-75 mm. 

Body elongate, testaceous, head short and broad (Text-fig. 290) ; occipital suture deeply 
concave in middle ; paraclypeae short, medially thick and rounded at apices. Antennae, basal 
segment with inner side pale outer side and bases of second and third black, apical segment with 
at least basal quarter pale. Rostrum long, usually reaching apices of third coxae. Lateral 
black line extending from sides of antenniferous tubercles on to anterior three-quarters of 
propleuron. Pronotal disc pale, posteriorly with black punctures, black spots on posterior 
angles present. Legs yellowish, with apices of femora, tibiae and tarsi blackish. Hemielytra, 
clavus with black punctures. Posterior margin of seventh abdominal tergum medially slightly 
convex ; lobes of ninth segment comparatively larger (Text-fig. 291). 

6 GENITALIA: Claspers very thick and truncated at apices (Text-fig. 200) ; dorsal and 
ventral thecal appendages present, with symmetrical ventral appendages (Text-fig. 293) ; 
lateral conjunctival appendage strongly inwardly curved (Text-fig. 214, E) ; frontal con- 
junctival appendage with two small sclerotized pieces (Text-fig. 214, A) ; two right lateral 
conjunctival appendages, with second short, thick and pointed at apex (Text-fig. 214, F) ; 
membranous appendage with a sclerotized cap-like terminal and a bilobed dorsal appendage 
(Text-fig. 293). 

9. Very similar to g in colour and measurements, posterior margin of seventh abdominal 
sternum emarginate with a small convex median projection and tiny backward projections at 
sides (Text-fig. 292). 

Q GENITALIA : First gonocoxae small, outer margin convex, apices rounded (Text-fig. 292) ; 
only four pairs of intervalvular sacs present (Text-fig. 294) ; spermatheca having a knob at 
apex, with a median flange and a long coiled tube (Text-fig. 228). 


Type material. Holotype 2, NEw GuINEA: B.M.(N.H.). 


Other material. CERAM ISLAND: I ¢; HALMAHERA ISLAND: I ¢; NEW 
GUINEA : 114 gf, 65 9; NEW IRELAND: 156, 13 9; PHILIPPINE ISLANDS: I 9; 
SOLOMON ISLANDS: 34,19; NEW HEBRIDES: 64, 3 9. 


The range of this species extends in the west into Indonesia (north of Ceram Island) 
and in the east into New Hebrides. It is very common in New Guinea and New 
Hebrides. The adults have been collected throughout the year. 


This species is closest to L. solomonensts sp. n. but can easily be separated by rost- 
rum which reaches third coxae, but not in solomonensis. In the aedeagus of 
solomonensis the ventral thecal appendages are asymmetrical and the terminal 
appendage is missing. 


Leptocorisa lepida Breddin 


(Text-figs. 206, 220, 234, 248, 262, and 296-302) 


Leptocorisa lepida Breddin, 1909 : 293 
Leptocorisa (Rhabdocoris) lepida (Breddin) China, 1924 : 238 


Characterized by very short and slim body, by long, cylindrical and pointed 
paraclypeae and by the form of § and 9 genitalia. 


THE LEPTOCORISINAE OF THE WORLD 89 
6. Body length, 11-5-11-8 mm., width, 1-6-1-8 mm. Head length, 1-6-1-7 mm., width, 
I-I-I'2 mm. Antennae, segment I, 3:5-3:7 mm., II, 2-0-2:2 mm., III, 2-5-2:6 mm., IV, 
missing. Rostrum, segment I, 1-o mm., II, t-o mm., III, 0-6 mm., IV, 0-6 mm. Pronotum, 
length, 1-7-1-9 mm., width, 1-6-1-°8 mm. Hind femora, 6-0 mm. Hind tibia, 6-0 mm. Dis- 
tance between ocelli, 0-33—0-35 mm., distance between an ocellus and an eye, 0:56—0-6 mm. 


296, head, dorsal 


Leptocorisa lepida, 296-298, 300, 3, 299, 301, 302, 9. 
298, terminalia, dorsal view ; 299, 


301, ovipositor, 


Fics. 296-302. 
view ; 297, claspers, inner and outer views ; 
terminalia, ventral view ; 300, aedeagus (inflated), ventral view ; 


ventral view ; 302, same, dorsal view. 


go I. AHMAD 


Body short and slim, pale unicolourous. Head elongate (Text-fig. 296) ; occipital suture 
complete ; paraclypeae elongated and cylindrical with pointed apices. Antennae pale, tinged 
with black, underside of basal segments and bases of each segment pale. Rostrum short, 
reaching apices of second coxae. Lateral black line faint on sides of antenniferous tubercles, 
extending on to collar like a black dot. Pronotal disc punctate, pale, sometimes tinged with 
green ; brownish black spots on posterior angles present. Legs uniformly pale, with articula- 
tions between femora, tibiae and tarsi black. Hemielytra pale unicolourous. Posterior margin 
of seventh abdominal tergum with a very small rounded projection ; pygophore round (Text- 
fig. 298). 

3 GENITALIA: Claspers with long blades and with curved and pointed apices (Text-fig. 
206) ; both dorsal and ventral pairs of thecal appendages present, with symmetrical sac-like, 
broad ventral appendages (Text-fig. 220) ; four lateral conjunctival appendages as in Text-fig. 
300 ; frontal conjunctival appendage without any sclerotized piece (Text-fig. 220,A) ; three 
right lateral conjunctival appendages, with second short, straight and thick only in middle 
(Text-fig. 220,F) ; membranous appendage with a spiculed, cap-shaped terminal and a short, 
curved and acutely pointed dorsal appendage. 

2. Very similar to f in general appearance and measurements ; posterior margin of seventh 
abdominal sternum sinuately emarginate (Text-fig. 299). 

Q GENITALIA : First gonocoxae short, broad, rounded at apices (Text-fig. 299) ; six pairs 
of intervalvular sacs (Text-fig. 302) ; spermatheca short, pointed at apex, with a median flange 
and a short coiled tube (Text-fig. 234). 


Type material. Holotype J, without data, labelled ‘‘ Leptocorisa lepida Bredd.”’, 
“ Type! ’’, Coll. Breddin, Dtsch. Entom. Inst., Berlin. 


Other material: InpiIa: 2 ¢, 2 2; BuHuTAN: 45 6,.35 9 7 BURMA + Ge 
THAILAND : 24, 2 Q. 


This species is common in India (U.P.) and its range extends in the east as far as 
Thailand. The adults have been collected in April, September and December. 


This species is very close to L. pseudolepida sp. n. and L. luzonica sp. n. and shares 
with them the character of small size of the body (less than 12 mm. in length). It 
can be separated from them by the long and cylindrical paraclypeae, which are 
pointed at their apices and by other genital characters (see above). 


Leptocorisa luzonensis sp. n. 
(Text-figs. 205, 219, 233, 247, 261 and 303-307) 


Characterized by apical antennal segments, which are equal to or longer than 
second and third segments together and by the form of § and @ genitalia. 


6. Body length, 17-0-17-5 mm., width, 2-6-2-°8 mm. Head length, 1-9-2-0 mm., width, 
1-6-1'°3 mm. Antennae, segment I, 4-8-5:1 mm., II, 2:8-3-0 mm., III, 3-5-3:7 mm., IV, 
6:5-6:7 mm. Rostrum, segment I, 1-5-1-7 mm., IJ, 1-5-1:7 mm., III, 0-8-o-9 mm., IV, 
0-7-0'75 mm. Pronotum, length, 2:6-2:8 mm., width, 2-6-2:7 mm. Hind femora, 7:0- 
7:2mm. Hind tibia, 7-5-7-6mm. Distance between the ocelli, 0-5 mm., distance between an 
ocellus and an eye, 0-7-0-75 mm. 

Species robust, yellowish brown. Head elongated, with two brownish black spots behind 
ocelli ; occipital suture medially concave ; paraclypeae short, with rounded apices. Antennae 
brownish black, with basal three-quarters of first segments, apices of second and third and bases 
of apical segments, pale. Rostrum short, reaching apices of second coxae. Lateral black 


THE LEPTOCORISINAE OF THE WORLD QI 


line extending from sides of antenniferous tubercles on to collar. Pronotal disc pale, with black 
spots on posterior angles. Legs pale, with apices of femora and bases of tibiae and tarsi black. 
Hemielytra pale unicolourous. Posterior margin of seventh abdominal tergum medially 
convex, pygophore large with thin lobes of ninth segment (Text-fig. 303). 

6 GENITALIA : Claspers with very short blades, apices pointed (Text-fig. 205) ; both dorsal 
and ventral pairs of thecal appendages present, ventral pair symmetrical as in Text-fig. 305 ; 


Fics. 303-307. Leptocorisa luzonensis, 303, 305, 5, 304, 306-307, 2. 303, terminalia, 
dorsal view ; 304, ovipositor, dorsal view ; 305, aedeagus (inflated), ventral view ; 
306, terminalia, ventral view ; 307, ovipositor, ventral view. 


92 I. AHMAD 


left lateral conjunctival appendage curved near apex (Text-fig. 219, E) ; frontal conjunctival 
appendage with two sclerotized pieces (Text-fig. 219, A) ; the only right lateral conjunctival 
appendage bilobed with a very thick main stem (Text-fig. 219, F) ; membranous appendage 
long, with a short triangular terminal and a bow-shaped, pointed dorsal appendage (Text-fig. 
305). 

2. Body length, 17:5-18:0 mm., width, 2:65-2:°8 mm. MHead length, 2-0-2-2 mm., width, 
1:8-1'°9 mm. Antennae, segment I, 4:8-5:0 mm., II, 2-:8-3-0 mm., III, 3:5-3:7 mm., IV, 
6-5-6:7 mm. Rostrum, segment I, 1-6-1:°8 mm., II, 1-6-1-8 mm., III, 0-9-1-0 mm., IV, 
o:7-0'8 mm. Pronotum, length, 2-6—2-8 mm., width, 2:65-2:80 mm. Hind femora, 7-2— 
7-4mm. Hind tibia, 7-8-8-0 mm. Distance between ocelli, 0:45-0:55 mm., distance between 
an ocellus and an eye, 0:75-0:85 mm. 

Usually slightly more robust than ¢ ; posterior margin of seventh abdominal sternum emar- 
ginate, with a very small median projection (Text-fig. 306). 

Q GENITALIA : First gonocoxae broad, outer margin sinuate (Text-fig. 247) ; eleven pairs of 
intervalvular sacs (Text-fig. 304) ; spermatheca elongated with a median flange and a short 
coiled tube of different thicknesses (Text-fig. 233). 


Holotype 3, PHILIPPINE ISLANDs : North Luzon, Los Bafios, 21.1.1914 (G. Bottcher) 
B.M.(N.H.). 


Paratypes. 1, 4 9, with some data as holotype, B.M.(N.H.), North Luzon, Los 
Bafios, 1 g, (P. I. Baker), Mus. Hels., 1 3, without any date, Riksmus. Stockholm, 
Luzon, Mt. Maquiling, Elev. 50 m., 3.ix.1949 (Boscos), Moravian Mus., Berne. 


This species is probably localized in Philippine Islands and has not been found in 
any other part of Oriental regions. Adults have been collected in the months of 
January and September. 


This species is closely allied to another, slightly more widely distributed species, 
L. tagalica, and shares the characters of black apices of femora, and bases of tibiae 
and tarsi ; however the § can easily be separated by very short and pointed blades of 
claspers (very long and cylindrical in tagalica) and the 9 by small, convex median 
projection of seventh abdominal sternum (a large median projection in tagalica) and 
by first gonocoxae curved near apices. This species can be separated from all other 
species of this genus by long apical antennal segments, which are at least as long as or 
longer than second and third together. 


Leptocorisa luzonica sp. n. 
(Text-figs. 208, 222, 236, 250, 264 and 308-314) 


Characterized by small body, straight and blunt paraclypeae and by the form of J 
and 9 genitalia. 


6. Body length, 11-3-11-5 mm., width, 1-5-1-6 mm. Head length, 1:5-1-7 mm., width, 
o:8-I-o mm. Antennae, segment I, 2-9-3-0 mm., II, 1-7-1°8 mm., III, 2-0-2-4 mm., IV, 
4:0-4'2 mm. Rostrum, segment I, 1-o-1-r mm., II, r-o-1:t mm., III, 0-5-0-6 mm., IV, 
0-6-0'7 mm. Pronotum, length, 1-7-1:8 mm., width, 1-5-1-6mm. MHind femora, 5-0—5:2 mm. 
Hind tibia, 5:2-5:3 mm. Distance between ocelli, 0-4 mm., distance between an ocellus and an 
eye, 0-7 mm. 

Species of small size, pale, unicolourous. Head comparatively short, occipital suture rather 
invisible at sides (Text-fig. 309) ; paraclypeae straight, thick at base, with rounded apices. 


THE LEPTOCORISINAE OF THE WORLD 93 


Antennae sometimes with brownish tinge, apices of second segments and apical segments 
brownish only pale at bases. Rostrum short, hardly reaching apices of second coxae. Lateral 
black line usually faint at sides of antenniferous tubercles and like a black dot on sides of 
collar. Pronotal disc pale, unicolourous, black spots on posterior angles rather faint, legs pale, 
unicolourous. Hemielytra pale. Posterior margin of seventh abdominal tergum truncated, 
very slightly medially produced ; pygophore rounded with pointed apices (Text-fig. 308). 

6 GENITALIA : Claspers laterally with short blades and curved sharp apices (Text-fig. 208) ; 
both dorsal and ventral pairs of thecal appendages present, ventral appendages symmetrical, 
sac-like with sclerotized linings (Text-fig. 222, D) ; four left lateral conjunctival appendages as 
in Text-fig. 311 ; frontal conjunctival appendage with two sclerotized pieces (Text-fig. 222, A) ; 
only two right lateral conjunctival appendages, second long and strongly curved in middle 
(Text-fig. 222, F) ; membranous appendage with two terminal appendages and a flat dorsal 
conjunctival appendage as in Text-fig. 311. 


0-25 mm. 


Fics. 308-314. Leptocorisa luzonica, 308-311, 3, 312-314, 2. 308, terminalia, dorsal 
view ; 309, head, dorsal view ; 310, claspers, inner view ; 311, aedeagus (inflated), 
ventral view ; 312, ovipositor, dorsal view ; 313, terminalia, ventral view ; 314, 
ovipositor, ventral view. 


94 I. AHMAD 


2. Body length, 11-5-12-0mm., width, 1-7-1-9mm. Head, length, 1-7 mm., width, 1-2 mm. 
Antennae, segment I, 3-5-3-7 mm., II, 2-o-2:2 mm., III, 2-6-2-8 mm., IV, missing. Rostrum, 
segment I, 1-0 mm., II, 1-0 mm., III, 0-6 mm., IV, 0-6 mm. Pronotum, length, 1-7-2-0 mm., 
width, 1:7-1-9 mm. Hind femora, 6-0 mm. Hind tibia, 6-0 mm. Distance between ocelli, 
0-3-0-4 mm., distance between an ocellus and an eye, 0-5—0-7 mm. 

Slightly robust, posterior margin of seventh abdominal sternum deeply emarginate medially, 
slightly convex (Text-fig. 313). 

Q GENITALIA : First gonocoxae elongated, curved in middle (Text-fig. 250) ; seven pairs 
of intervalvular sacs (Text-fig. 312) ; spermatheca long and cylindrical, thinner in middle with 
a long coiled tube (Text-fig. 236). 


Holotype g, PHILIPPINE ISLANDS: Luzon, Manginin, 17.1x (G. Bottcher), B.M. 
(N.H.). 


Paratypes. PHILIPPINE ISLANDS : 3 4, 2 9, same data as holotype, B.M.(N.H.), 
Mindanao, Sulu, near base of Mt. Dahao Jolo 1, 150 m., II g, 12 Q, 2.1x.1958 
(Milliron), Bern. Bishop Mus.; VietNAmM: Tonkin, Choganh, Hoa-Binh, 5 3, I 9, 1919 
(J. De Cooman, L. Duporte, R. Oberthur) Mus. Hist. nat Paris ; SARAWAK : Kuch- 
ing, Santubong, 797-1,500 m., 41 3g, 45 9, 18—30.vi.1958 (T. C. Maa), Bern. P. 
Bishop Mus., K. S. Road, 194 miles, Tarat Station, 1oth mile Bau Rd., 18th mile 
K. S. Rd. on grasses near wet paddy, 105 3, 95 9, 13.i11.1963 (S. K. Hueh, R. Lav), 
Dept. Agric., Sarawak. 


This species is found in Vietnam in the west, Philippine Islands in the east and 
Borneo in the south. The adults have been collected in March, June and September 
but is seems to be most abundant in March and September. This species has also 
been reported feeding on grasses near paddy fields, but probably does not commonly 
feed on paddy or this would have been recorded. 


This species is close to L. lepida Breddin and L. pseudolepida sp. n., its allopatric 
allies, in having small size of the body (less than 12 mm.) but can easily be separated 
from them by the shape of the paraclypeae, which are straight, thicker in the middle 
and rounded at the apices. The conjunctival appendages of the aedeagus, the first 
pair of gonocoxae in 9 and the shape of the spermatheca confirm its identity. 


Leptocorisa oratorius (Fabricius) 
(Text-figs. 197, 211, 225, 239, 253 and 315-320) 


Gerris ovatorvius Fabricius, 1794 : 191, 1803 : 261 

Leptocorisa bengalensis Westwood, 1842 : 18 

Rhabdocoris arcuata Kolenati, 1845 : 67 

Myodochus trinotata Herrich Schaffer, 1848 : 95 

Leptocorisa maculiventris Dallas, 1852 : 484 

Leptocorisa (Rhabdiocoris) acuta (Thunberg) ; Distant, 1902 : 410, China, 1924 : 237 [Mis- 
identifications] 


Characterized by the robust body, series of brownish black dots on the ventro- 
lateral side of abdomen and by the form of 3 and @ genitalia. 
3d. Body length, 18-0-18:5 mm., width, 2:9-3-0 mm. Head length, 2-4—2-6 mm., width, 


1-8-2-0 mm. Antennae, segment I, 5-0-5:2 mm., II, 3-4-3:5 mm., III, 3-9-4-:0 mm., IV, 
5:6-6:0mm. Rostrum, segment I, 1-9mm., II,1-9mm., III, 1-omm.,IV,o-9mm. Pronotum, 


THE LEPTOCORISINAE OF THE WORLD 95 


length, 3-1-3:5 mm., width, 2-9-3-0 mm. Hind femora, 6-7 mm. Hind tibia, 7-6 mm. _ Dis- 
tance between ocelli, o-6-o-7 mm., distance between ocellus and outer margin of eye, 0-8 mm. 

Body robust, pale unicolourous. Head elongated ; occipital suture medially concave ; 
paraclypeae short and pointed. Antennae usually pale, unicoloured, rarely with brownish 
black tinge. Rostrum short, reaching only the apices of second coxae. Lateral black line on 
the sides of head and propleuron absent. Pronotal disc pale unicolourous, posterior angles 
without black spots. Legs usually pale unicoloured, rarely tinged with black. Hemielytra 
pale. A series of ventro-lateral brownish black spots on abdomen usually prominently present, 
rarely faint. Posterior margin of seventh abdominal tergum medially convex ; pygophore 
very large and almost rectangular in form (Text-fig. 317). 

6 GENITALIA: Claspers curved and tapering to a point at apices (Text-fig. 197) ; both 
dorsal and ventral pairs of thecal appendages present, ventral pair symmetrical as in Text-fig. 
316 ; left lateral conjunctival appendage elongated, curved at base and apex (Text-fig. 211, E) ; 
frontal conjunctival appendage with two sclerotized pieces as in Text-fig. 211, A ; two right 
lateral conjunctival appendages, second bifid at apex (Text-fig. 211, F) ; membranous append- 
age usually with a bulbous or pointed terminal appendage and a hooked bilobed dorsal con- 
junctival appendage (Text-fig. 316). 


Fics. 315-317. Leptocovisa ovatorius. 315, 9 terminalia, ventral view ; 316, aedeagus 
(inflated), ventral view ; 317, d terminalia, dorsal view. 


96 I. AHMAD 


9. Very similar to j in general appearance and measurements ; posterior margin of seventh 
abdominal sternum emarginate, very slightly convex in middle (Text-fig. 315). 

Q GENITALIA : First gonocoxae elongated, about three times as long as broad, apices pointed 
(Text-fig. 239) ; six pairs of intervalvular sacs (Text-fig. 320) ; spermatheca thick in middle 
with a median flange, round and coiled tube (Text-fig. 225). 


Type material. Holotype ¢ of Gerris oratorius Fabricius, SUMATRA : (Daldorff), 
with label “‘ Mus. de Johestedt’’, “‘ Gerrits oratorius Fabricius, s.: G. varicornis 
Fabr. ”’ “‘ Teste Stal’, Zool. Mus. Copenhagen ; cotype 2 of G. angustata Fabricius 
1787, without data, Zool. Mus. Copenhagen ; holotype ¢ of Leptocorisa bengalensis 
Westwood, 1842, INDIA: Bengal, Hope Dept., Univ. Mus., Oxford ; holotype g, 


Or---- 5 


_0:25 mm. 


Fics. 318-320. Leptocorisa ovatorius, 9. 318, ovipositor, with 9th paratergite, ventral 
view ; 320, same, dorsal view. 


THE LEPTOCORISINAE OF THE WORLD 97 


Rhabdocoris arcuata Kolenati, 1845, without data (R. A. Kywakeeuya), with label 
“ Rhabdocoris arcuatus Klti’’, ‘‘ Mel II, 67”’, Zool. Mus. Leningrad ; 1 g, CHINA: 
Komai (Sanganchu) in Tibet, with label “ trinotatus ”’ “‘ Herrich Schaffer, fig. 863 ”’, 
Riksmus. Stockholm. 


Other material. CEYLON: 54,19; INDIA: 23g, 18 2; NICOBAR ISLAND : 
46; PAKISTAN: 14,29; BHUTAN: 404, 259; BuRMA: 19; THAILAND : 
19 g, 13 2; VIETNAM : 32 4, 282; MALAYA: 54,3 9; SINGAPORE: Ig; Su- 
Megea: ig; Java: 38d, 18 9; SARAWAK : 404, 37 2; CELEBES: 24,29; 
PHILIPPINE ISLANDS : 157 gd, 128 9 ; AUSTRALIA: 44, 42; SOLOMON ISLANDs : 


Be, 


The range of this species extends in the west into Ceylon, in the north to Malabar, 
Sylhet (East Pakistan), Bhutan, China (Komai in Tibet) in the south into North 
Queensland (Australia) and in the east as far as Solomon Islands. 


Uichanco (1921) has stated that its common name in India, China and Java is 
““Waleng sangeet ’’, but in the Philippines, it is called by various common names 
such as Tagalog, Bicol, Visayun, Pangasinan, Panpanga, Ilacano and Moro. It is 
a serious rice pest in the Far East countries and, to quote Uichancho, “ of the insect 
enemies of rice there is probably none that is so well known among the Filippino 
planters as the rice bug Leptocorisa acuta Thunberg’’. The figure he gives of the 
pygophore makes it clear that he was referring to this species and not L. acuta s. str. 
He has estimated that in abundance, it destroys at times over one-half of the season’s 
crop. It causes the greatest damage in the months of November and December, when 
the rainy season rice is in the milk stage. Its life history has also been studied by 
Koningsberger (1903) and Zehntner under the name of Leptocorisa acuta (Thunberg). 
It has also been collected at light in Java. 


This species is close to L. acuta (Thunberg) and L. chinensis Dallas but can easily 
be separated by a series of ventro-lateral black dots on abdomen. In the claspers 
which posteriorly taper into a point and in the the first pair of elongated and pointed 
gonocoxae are its peculiarities. 

Stal (1873) synonymized the following species : 

Cimex acuta Thunberg, Cimex angustata Fabricius 1787, Gerris oratorius Fabricius 
1794, Leptocorisa bengalensis Westwood 1842, Rhabdocoris arcuata Kolenati 1845, 
Myodochus trinotata Herrich Schaffer 1848 and Leptocorisa maculiventris Dallas 1852. 
During the present study the holotype of acuta Thunberg has been examined and 
because of its damaged abdomen, another male specimen (from type locality) and 
with Stal’s label ‘‘ Comex acutus Thunb. ! Mus. Thunb. ”’ has been designated neotype 
of C. acuta Thunberg (see page 80). The type of C. angustata Fabricius has been 
found to be identical with C. acuta Thunberg. The types of Gerris oratorius Fabri- 
cius, Leptocorisa bengalensis Westwood and Rhabdocoris arcuata Kolenati all differ 
from the type of acuta (J. S. varicornis) and represent the modern concept of L. acuta 
China nec Thunberg 1783. This acuta auctt., which is now without a name, should 
take the name of the most senior synonym, i.e. Gerris oratorius Fabricius 1794. The 
types of M. trinotata Herrich Schaffer 1848 and L. maculiventris Dallas could not 


98 I. AHMAD 


be examined. Dr. Forster (Zoologische Sammlung des Bayerischen Staates) has 
informed me that the type of M. trinotata Herrich Schaffer was destroyed during an 
air raid in 1944, and Dr. W. E. China (British Museum) has told me that the type of 
L. maculiventris Dallas must be presumed lost. However the original description 
and diagram (Fig. 863) of Herrich Schaffer for ¢tvinotata and the original description 
of L. maculiventris Dallas agree with the holotype of Gerris oratorius Fabricius. 


Leptocorisa palawanensis sp. n. 
(Text-figs. 201, 215, 229, 243, 257 and 321-328) 


Characterized by very long and pointed paraclypeae which are only very slightly 
less than half of entire head length, by rostrum, which extends beyond the apices 
of second coxae and by the form of genitalia. 


6. Body length, 14:9 mm., width, 2-30 mm. Head length, 2-2 mm. (only paraclypeae 
0:83 mm.), width, 1-5 mm. Antennae, segment I, 4:7 mm., II, 3-0 mm., III, 3:3 mm., IV, 
5-7mm. Rostrum, segment I, 1-7 mm., IJ, 1-5 mm., III, 0-8 mm., IV, 0-8 mm. Pronotum, 
length, 2-6 mm., width, 2-3 mm. Hind femora, 6-9 mm. Hind tibia, 7-3 mm. Distance 
between ocelli, 0-5 mm., distance between an ocellus and an eye, 0-8 mm. 


323 


0:25 mm. 324 


Fics. 321-325. Leptocorisa palawanensis, 3. 321, terminalia, dorsal view ; 322, head, 
dorsal view ; 323, claspers, inner and outer views ; 324, aedeagus (inflated), ventral 
view ; 325, theca, dorsal view. 


THE LEPTOCORISINAE OF THE WORLD 99 


Body cylindrical, pale. Head elongate ; occipital suture smoothly medially concave ; 
paraclypeae very long, cylindrical and pointed. Antennae with basal segments pale, only black 
at apices, second, third and fourth segments black, only pale at bases. Rostrum long, extending 
beyond apices of second coxae. Lateral black line faint on the sides of head, represented only 
by a black dot on either side of collar. Pronotal disc light brown, sometimes with greenish 
tinge ; brownish black spots on posterior angles present. Legs pale, with usually apices of 
femora, tibiae and tarsi black. Hemielytra pale. Posterior margin of seventh abdominal 
tergum sinuate, with a very small pointed median projection ; pygophore round (Text-fig. 321). 

6 GENITALIA: Claspers truncated and thick at apices with very short blades (Text-fig. 201) ; 
both dorsal and ventral pairs of thecal appendages present, ventral appendages bilobed and 
symmetrical (Text-fig. 325) ; left lateral conjunctival appendage strongly curved and pointed 
(Text-fig. 215, E) ; frontal conjunctival appendage with only one sclerotized piece (Text-fig. 
215, A) ; three right lateral conjunctival appendages as in Text-fig. 324, second smaller, curved 
in middle and pointed (Text-fig. 215, F) ; membranous appendage short, with a hooked terminal 
and a curved dorsal conjunctival appendage (Text-fig. 324). 

Q. Very similar to f in general appearance and measurements, posterior margin of seventh 
abdominal sternum emarginate with a small pointed median projection (Text-fig. 326). 


0-5 mm. 


Fics. 326-328. Leptocorisa palawanensis, 9. 326, terminalia, ventral view ; 327, 
ovipositor, ventral view ; 328, same, dorsal view. 


100 I. AHMAD 


Q GENITALIA : First gonocoxae small, outer margin convex, apices rounded (Text-fig. 243) ; 
seven pairs of intervalvular sacs (Text-fig. 328) ; spermatheca with a long coiled tube and a 
median flange (Text-fig. 229). 


Holotype 3, PHILIPPINE IsLANDs : N. Palawan, Binaluan, xi—xii.1913 (G. Roett- 
cher), B.M.(N.H.). 


Paratypes. PHILIPPINE ISLANDs : I g (Semper), Riksmus. Stockholm, Mindanao, 
Iligan, 1 g (Baker), Mus. Hels., Mindanao, Misamis, Sulu, Palawan, 3:2 km. 
south of Tarumpitao, 2.vi., 20.vil., 7.ix., I9-20.x.1958 (various collectors), 
Bern. P. Bishop Mus. ; NEw GuINEA: Papua, West District Orimo, Govt. 
Sta., 26-28.x.1960 (J. L. Gressitt), Bern. P. Bishop Mus. ; AUSTRALIA : Queensland, 
I g (Mjéberg), Riksmus. Stockholm. 


The range of this species extends in the north west into Philippine Islands and in 
the south east into northern Australia (Queensland). The adults have been collected 
in the late part of the year from June to December. 


This species is very close to L. acuta (Thunberg) and L. discoitdalis Walker but can 
easily be separated from them by very long (about half the length of the head), 
cylindrical and pointed paraclypeae. In the aedeagus, the bilobed symmetrical 
ventral thecal appendages and in the spermatheca the tube, which is twisted at is 
base, are its peculiarities. 


Leptocorisa pseudolepida sp. n. 
(Text-figs. 207, 221, 235, 249, 263 and 329-335) 


Characterized by very small size of the body, by short, thick and pointed para- 
clypeae and by the form of 3 and Q genitalia. 


6. Body length, 11-5-12:0 mm., width, 1-6-1-9 mm. Head length, 1:5-1-7 mm., width, 
I:2-1°3 mm. Antennae, segment I, 3-6-4-0 mm., II, 2:2-2:4 mm., III, 2-4-2:8 mm., IV, 
4:0-4:2 mm. Rostrum, segment I, 1-0-1: mm., II, 1-1-1-2 mm., III, 0-6-0-7 mm., IV, 
0:6-0:7mm. Pronotum, length, 1-7—-2-0 mm., width, 1-7-1-9mm. Hind femora, 5-o—6-2 mm. 
(specimen from Malacca or Borneo with smaller hind femora (5-0-5-2 mm.) from Selangor, 
Kuala Lumpur (6:2 mm.)). Hind tibia, 5-6-6-5 mm. (specimens from Borneo or Malacca 
5°6-5-8 mm. and from Kuala Lumpur 6-5 mm.). Distance between ocelli, 0o-3-0-4 mm., distance 
between an ocellus and an eye, 0-5—0:6 mm. 

Body short, pale, tinged with black. Head short, occipital suture complete (Text-fig. 329) ; 
paraclypeae short, thick at base, curved near apices and pointed. Antennae usually brownish 
black, with basal segments pale, only black at apices, second, third and fourth segments brownish 
black, only pale at their bases. Rostrum short, reaching apices of second coxae. Lateral black 
line faint on sides of head, only represented on sides of head by a black dot. Pronotal disc pale, 
unicoloured, sometimes with brownish tinge. Legs usually black, only femora pale. Hemiely- 
tra pale. Abdomen dorsally palish brown, connexiva and venter pale, ochraceous. Posterior 
margin of seventh abdominal tergum truncated, pygophore rounded, apex pointed (Text-fig. 
332). 

d GENITALIA: Claspers never extending beyond posterior margin of pygophore, dorsally 
rounded (Text-fig. 332), apices truncated (Text-fig. 207) ; both dorsal and ventral thecal 
appendages present as in Text-fig. 331, ventral, symmetrical ; four lateral conjunctival append- 
ages as in Text-fig. 331 ; frontal conjunctival appendage with two elongated sclerotized pieces 
(Text-fig. 221, A) ; three right lateral conjunctival appendages (Text-fig. 331) ; second straight, 


THE LEPTOCORISINAE OF THE WORLD 101 


thick at base and pointed at apex (Text-fig. 221, F) ; membranous appendage with a cap-like 
terminal and a thin curved dorsal appendage (Text-fig. 331). 


9. Very similar to J in general appearance and measurements ; posterior margin of seventh 
abdominal sternum deeply emarginate (Text-fig. 333). 


Q GENITALIA : First gonocoxae small and with outer margin sinuate (Text-fig. 249) ; five 


pairs of intervalvular sacs (Text-fig. 334) ; spermatheca broad at apex, with a median flange 
and a long coiled tube. 


Holotype 3, MALaya : Selangor, Kuala Lumper, 17th mile Chesar Rd., at light, 
17.1.1924 (E. Seimond), B.M.(N.H.). 


Paratypes. CEYLON: 2 3, 1 9, B.M.(N.H.) ; Maraya: Malacca (Keranton), 
with labels “‘ Leptocorisa lepida Bredd.’’, “‘ Coll. Breddin’’, Dtsch. Entom. Inst., 
Berlin ; Inp1A : S. Malabar, 4 3, 3 2 (various collectors), B.M.(N.H.) ; BORNEO : 
(south east), 1 gf, 1 9, 1895 (Atkinson coll.), Hope Dept., Univ. Mus., Oxford. 


0:25 mm. 


Fics. 329-335. Leptocorisa pseudolepida, 329-332, 3, 333-335) 2- 329, head, dorsal view ; 
330, claspers, inner and outer views ; 331, aedeagus (inflated), ventral view ; 332, 
terminalia, dorsal view ; 333, terminalia, ventral view ; 334, ovipositor, dorsal view ; 
335, Same, ventral view. 


102 I. AHMAD 


The range of this, possibly rather rare, species extends in the west into Ceylon 
and in the east into south east Borneo. The adults have been collected in January. 
This species has also been collected at light in Malaya (Selangor State). 


This species is close to L. lepida Breddin and L. luzonica sp. n., but can easily be 
separated from them by its short, thick, curved and pointed paraclypeae (Text-fig. 
329). In aedeagus, its elongated, ventral thecal appendages (Text-fig. 221, D), long 
and straight second right lateral conjunctival appendage (Text-fig. 221, F) and in the 
? the deep emarginate posterior margin of seventh abdominal sternum (Text-fig. 333) 
are its peculiarities. 


Leptocorisa sakdapolrakae sp. n. 
(Text-figs. 210, 224, 238, 252, 266 and 336-341) 


Characterized by short but cylindrical and pointed paraclypeae, long rostrum, 
which extends well beyond the apices of second coxae and by the form of the ¢ and 
2 genitalia. 


3g. Body length, 13-:06—-15-:2 mm., width, 1-8-2: mm. Head length, 1-73-2-0 mm., width, 
I-3-1-6 mm. Antennae, segment I, 3-6-4:26 mm., II, 2-0-2:8 mm., III, 2:5-3-:2 mm., IV, 
5°2-5'73 mm. Rostrum, segment I, 1-3-1-6 mm., II, 1-3-1-6 mm., III, 0-6-0-8 mm., IV, 
o-6-0-8mm. Pronotum, length, 2-1-2:3 mm., width, 1-8-2: mm. Hind femora, 5-3—6:5 mm. 
Hind tibia, 6-o-7-3 mm. Distance between ocelli, 0-4-0-46 mm., distance between an ocellus 
and an eye, 0-73-0°8 mm. 

Body slim, light brown, tinged with black. Head short ; occipital suture “ V ’’-shaped in 
middle ; paraclypeae short, cylindrical, thicker at base and pointed at apices. Antennae 
brownish black with basal segments palish brown, apices black, second, third and fourth seg- 
ments only pale at their bases. Rostrum long, extending well beyond apices of second coxae. 
Lateral black line, extending from sides of antenniferous tubercles on to collar. Pronotal disc 
pale, sometimes with greenish tinge posteriorly. Legs usually black, pale near articulations. 
Hemielytra pale, slightly dark. Posterior margin of seventh abdominal tergum medially 
convex, pygophore elongated, with thin lobes of ninth segments (Text-fig. 336). 

d GENITALIA : Claspers in dorsal view, extending beyond posterior margin of pygophore 
(Text-fig. 336) ; apices truncated (Text-fig. 210) ; both dorsal and ventral pairs of conjunctival 
appendages present, ventral asymmetrical (Text-fig. 337) ; left lateral conjunctival appendage 
strongly curved in middle and acutely pointed at apex (Text-fig. 224, E) ; frontal conjunctival 
appendage with two sclerotized pieces (Text-fig. 224, A) ; two right lateral conjunctival append- 
ages as in Text-fig. 338 ; second long, strongly curved in middle and acutely pointed (Text-fig. 
224, F) ; membranous appendage with a trinagular terminal and a bilobed dorsal appendage 
(Text-fig. 338). 

Q. Very similar to g in general appearance and measurements, highly variable in size ; 
posterior margin of seventh abdominal sternum emarginate, with a small convex median 
projection (Text-fig. 339). 

@ GENITALIA : First gonocoxae elongated, outer margin sinuate, apices rounded (Text-fig. 
252) ; fourteen pairs of intervalvular sacs (Text-fig. 340) ; spermatheca comparatively very 
short, elongated with a transverse neck, a median flange and a long coiled tube (Text-fig. 238). 


Holotype 3, JAvA : Carnegie Mus. 


Paratypes. JAVA: Sindanglava, 1 g, 27.xi.1920, A.M.N.H.; THAILAND: 
Kaochong, Trang, I g, I 9, 18.iv.1960 (Sakdapolrak), 1 3, I 2, with same data as 
above, “terminalia missing’’, Coll. Sakdapolrak ; Maraya: Phang, Kuala 


THE LEPTOCORISINAE OF THE WORLD 103 


Terenggan, 220 m., 25 g, 18 9, 15.xii.1958 (J. L. Gressitt), Bern. P. Bishop Mus. ; 
SuMATRA : Labuan, Bilik, 48 3, 42 2 (Palm), Mus. Hels. 

The range of this species extends in the north west into Thailand and in the south 
east into Java. Adults are probably in greatest abundance in the late part of the 
year, November and December. 


“WW S.Q 


Fics. 336-341. Leptocorisa sakdapolrakae, 336-338, 3, 399-341, 2. 336, terminalia, dorsal 
view ; 337, theca, dorsal view ; 338, aedeagus (inflated), ventral view ; 339, termin- 
alia, ventral view ; 340, ovipositor, dorsal view ; 341, same, ventral view. 


104 I. AHMAD 


This species is close to L. discoidalis Walker and L. acuta (Thunberg). From the 
former species it can be very easily separated by rostrum, which does not reach third 
coxae and from both by the lateral black line, which never extends beyond pronotal 
collar. Its asymmetrical ventral conjunctival appendages in the aedeagus show its 
relationship with L. solomonensis sp. n. and its very short spermatheca is its 
peculiarity. 

This species is named after Miss N. Sakdapolrak, who discovered it independently 
and generously made her material available to me. 


Leptocorisa solomonensis sp. n. 
Text-figs. 209, 223, 237, 251, 265 and 342-345) 
§ 9 


0-25 mm. 025 mm. 


Fics. 342-345. Leptocorisa solomonensis, §. 342, terminalia, dorsal view ; 343, aedeagus 
(inflated), ventral view ; 344, theca, dorsal view ; 345, claspers, inner and outer views. 


THE LEPTOCORISINAE OF THE WORLD 105 


Characterized by the dark black line whch extends from the sides of the antenni- 
ferous tubercles on to posterior of propleuron, by ventral black median longitudinal 
line which extends from first to sixth abdominal segment and by the form of the 3 
and 9 genitalia. 


3. Body length, 16-0-16-5 mm., width, 2-5-2-7 mm. Head length, 2-0-2-4 mm., width, 
1-6-1:7 mm. Antennae, segment I, 5:2-5-:3 mm., II, 2:5-3:3 mm., III, 3-8-4-0 mm., IV, 
5°0-5:'2mm. Rostrum, segment I, 1-7mm., II, 1-7mm., III, 1-omm.,1V,o-gmm. Pronotum, 
length, 2-6-2:9 mm., width, 2-5-2-7 mm. Hind femora, 7-0-7:3 mm. _ Hind tibia, 8-o-8-2 mm. 
Distance between ocelli, 0-5 mm., distance between an ocellus and an eye, 0-8 mm. 

Species of medium size, brownish black. Head short, occipital suture medially deeply 
concave, a black spot visible behind it ; paraclypeae short, thick at base and pointed at apices. 
Antennae brownish black, only bases of each segment pale. Rostrum long, extending well 
beyond apices of second coxae but never reaching third coxae. Lateral black line extending 


Fics. 346-348. Leptocorisa solomonensis, 9. 346, terminalia, ventral view ; 347, 
ovipositor, ventral view ; 348, same, dorsal view. 


106 I. AHMAD 


from sides of antenniferous tubercles on to posterior of propleuron. Pronotal disc pale, pos- 
terior three-quarters with black punctures, black spots on posterior angles present. Legs light 
brown, tinged black. Hemielytra pale, tinged with black. Abdomen with dorsum and venter 
red, a median longitudinal black line extending from first to sixth abdominal venter. Posterior 
margin of seventh abdominal tergum medially convex ; pygophore large, rectangular, with very 
thin lobes of ninth segment (Text-fig. 342). 

6 GENITALIA : Claspers usually extending beyond posterior margin of pygophore (Text-fig. 
342) ; blades very long with truncated apices (Text-fig. 209) ; both dorsal and ventral pairs of 
thecal appendages present, ventral pair asymmetrical (Text-fig. 344) ; left lateral conjunctival 
appendage reduced to a spine-like appendage (Text-fig. 223, E) ; frontal conjunctival appendage 
with two flat sclerotized pieces (Text-fig. 223, A) ; two right lateral conjunctival appendages 
(Text-fig. 343), second curved in middle like an arch (Text-fig. 223, F) ; membranous appendage 
with a curved and pointed dorsal appendage, but terminal appendage absent (Text-fig. 343). 

2. Very similar to g in general appearance and measurement ; posterior margin of seventh 
abdominal sternum emarginate, with a small triangular projection in middle (Text-fig. 346). 

Q GENITALIA: First gonocoxae with convex outer margin (Text-fig. 251) ; intervalvular 
sacs eight in number, three anterior pairs larger (Text-fig. 348) ; spermatheca large, flask-shaped 
with a thick, smoothly curved neck, median flange and a small coiled tube. 


Holotype 3, SoOLoMoN IsLANDs : Buin, 21.vii.1922 (E. A. Armytage), B.M.(N.H.). 


Paratypes. SOLOMON ISLANDS : Guadalanal, Lavoro, 2 3, I 9, 18.11.1934 (H. T. 
Pagden), Isles of Savu, I g, 28.iv.1922 (E. A. Armytage), B.M.(N.H.) ; Bougainville, 
I 9, (A. A. Voyce), labelled “‘ Leptocorisa discoidalis Walker ’’, det. N. C. E. Miller, 
S. A. Mus. ; NEw GUINEA: Papua, Port Moresby, Bisia tabu, Mt. Lamington, 
1,300-1,500 ft., 6g, 5 9, 1954 (W. N. Lock), S. A. Mus.; Brown River, 5 m., c. District 
of Otama Plant’n, 1 m. E. of Moresby, 5 3, 13 9, x.1959, 23-X, 2.Xi.1960 (various 
collectors), Bern. P. Bishop Mus. ; NEw IRELAND : Schleinitz Lelet Plateau, 5 3, 
Bern. P. Bishop Mus. 


The range of this species seems very restricted to eastern New Guinea (Papua) and 
the adjacent Solomon Islands. The adults have been collected at most times of the 
year. 


This species is closest to L. discoidalis Walker but can be easily separated by black 
median longitudinal line on the first to the sixth abdominal venter, the rostrum never 
reaches the third coxae and the pygophore (in the g) is comparatively very large and 
rectangular. Its asymmetrical ventral thecal appendages, the small spine-like 
second lateral conjunctival appendage, loss of terminal appendage and the elongated 
large flask-shaped spermatheca are other diagnostic features. 


Leptocorisa tagalica nom. n. 
(Text-figs. 204, 218, 232, 246, 260 and 349-353) 
Leptocorisa (Rhabdocoris) geniculata China, 1924 : 238 


Characterized by pale body, very long cylindrical and pointed claspers and other g 
and 9 genital characters. 
3g. Body, length, 17-0-17-5 mm., width, 2-5-2-°8 mm. Head length, 2-0-2:2 mm., width, 


1-6-1:'8 mm. Antennae, segment I, 5:2-5-4 mm., II, 3-4-3:6 mm., III, 3:8-3-9 mm., IV, 
6-0-6:3 mm. Rostrum, segment I, 1-:3-1-4 mm., II, 1-3-1-4 mm., III, 0-7-0-9 mm., IV, 


THE LEPTOCORISINAE OF THE WORLD 107 


o-7-0°8 mm. Pronotal length, 2-6—2:-9 mm., width, 2-5-2-8 mm. Hind femora, 7-0-7-7 mm. 
Hind tibia, 8-o-8-5 mm. Distance between ocelli, 0-5 mm., distance between an ocellus and 
an eye, 0-8—o-9 mm. 

Body robust, pale ochraceous, tinged with green. Head elongated, occipital suture medially 
concave, usually two brownish black spots behind it ; paraclypeae short, rounded at apices. 
Antennae with basal segments pale, black only at bases and apices, second, third and fourth 
black, only pale at bases. Rostrum short, reaching second coxae. Lateral black line extending 
from sides of antenniferous tubercles on to pronotal collar. Pronotal disc pale, densely punctate, 
with brownish black spots on posterior angles. Legs pale, with apices of femora and bases of 
tibiae prominently black. Hemielytra pale ochraceous. Abdomen with venter of dusty grey 
colour. Posterior margin of seventh abdominal tergum with a small convex median projection ; 
pygophore large, lobes of ninth segment thick (Text-fig. 349). 

6 GENITALIA : Claspers with short main stems and very long, cylindrical and pointed blades, 
(Text-fig. 204) ; both dorsal and ventral pairs of thecal appendages present (Text-fig. 350) ; 


Fics. 349-351. Leptocorisa tagalica. 349, d terminalia, dorsal view ; 350, aedeagus 
(inflated), ventral view ; 351, 9 terminalia, ventral view. 


108 I. AHMAD 


ventral pair symmetrical ; left lateral conjunctival appendage curved near apex, pointed 
(Text-fig. 218, E); frontal conjunctival appendage with two sclerotized pieces (Text-fig. 
218, A) ; three right lateral conjunctival appendages as in Text-fig. 350, second short and thick 
(Text-fig. 218, F) ; membranous appendage with a sclerotized cap-like and a dorsal appendage 
(Text-fig. 350). 

2. Very similar to gj in general appearance and measurement ; posterior margin of seventh 
abdominal sternum sinuately emarginate, with a small median rounded projection (Text-fig. 
351). 

@ GENITALIA: First gonocoxae broad, outer margin posteriorly concave (Text-fig. 246) ; 
thirteen pairs of intervalvular sacs (Text-fig. 353) ; spermatheca flask-shaped with a median 
flange and a long coiled tube (Text-fig. 232). 


Type material. Holotype 3, Leptocorisa gemiculata China, PHILIPPINE ISLANDS : 
(J. J. P. Mounsey), B.M.(N.H.) ; paratypes, 5 3, 3 9 of L. geniculata China, data as 
holotype, B.N.(M.H.). 


352 


353 


0-25 mm. 


Fics. 352-353. Leptocorisa tagalica. 2. 352, ovipositor, ventral view ; 
353, same, dorsal view. 


THE LEPTOCORISINAE OF THE WORLD 109 


Other material. SARAWAK : Ig, 19; HALMAHERA ISLAND : I 9 ; PHILIPPINE 
IseANDs : 13 9, 10 9: 


The range of this species extends in the north into Philippine Islands and in the 
south into Indonesia (Halmahera Islands). This species has also been found on 
paddy in Sarawak. The adults have been collected from March to November. 


This species is very close to L. /uwzonensis sp. n. and shares the characters of the 
black colour of the apices of femora and bases of tibiae and the two brownish black 
spots behind ocelli ; but can easily be separated from this species by pale ochraceous 
colour of the body, dusty grey colour of the venter of abdomen and short apical 
antennal segments. The long and cylindrical blades of claspers and other aedeagal 
appendages are of value in confirming identification. 

Leptocorisa geniculata China, 1924 is a primary homonym of Leptocorisa geniculata 
Guérin, 1857. In spite of the fact that Guérin’s species had been transferred to 
Lyrnessus by Stal (1866) before China described his species, China’s species must be 
renamed. Dr. W. E. China has kindly suggested that ¢agalica would be an appropri- 
ate name, representing its Philippine origin. 


Leptocorisa bipunctata Costa 
Leptocorisa bipunctata Costa, 1863 : 260 ; Stal, 1873 : 87 


Original description : ‘‘ Luteo—testacea, antennis brunneis, articulis 1z et 2z basi dilutioribu 
quarto basi pallido ; capitis linea utrinque laterali, pronoti punctis duobis humera-libus, 
elytrorum que cario margine laterali expecti fusco nigris, femoribus apice, tibiis tarsique fuscis 
6 long. mill 14, lat. mill 2 1/3 long, antenn. mill. 17. Patria ?”’. 


Every effort was made to find the type of this species but without success. 
Approaches to the University Museum, Naples were unsuccessful. None of the 
specimens examined had been labelled as Leptocorisa bipunctata Costa by earlier 
workers. The original description is so vague that it might apply to almost any 
member of the subfamily and without any knowledge of the type locality, its identity 
cannot be guessed. 


Leptocorisa burmeisteri Montrousier 
Leptocorisa burmeisteri Montrousier, 1865 : 277 ; Stal, 1873 : 87 


Original description (translated from French) : ‘ (length including antennae, 0-030 m.). 
Animal dark green above and yellowish below. Legs green, antennae brown. The upper 
surface of the head is light yellow, beneath it is yellowish green with the anterior portion apple 
green. The antennae which are as long as the body are brown with the extremity of the first 
segment dark green and the base of the fourth light green. The eyes are black, ocelli red. 
The rostrum, which is green, does not extend past the insertion of the middle legs. The pro- 
thorax is shagreened with gibbosity at its posterior angles, green marks on the rest of the border, 
green shades being paler at the sides, yellow below with a black spot at each side under the 
anterior angles. Scutellum finely punctate and almost metallic green. The elytra have the 
coriaceous part punctate, pink at the base and brown at the corners and bright green at the 
margins, especially the exterior, the membranous part obscurely smoky. Sternum is yellow 
and pale green in front of the insertion of the front legs. The abdomen is yellow, a little reddish 
on the sides with the tip green. Femora are green, the hairy tibiae are brownish and so are the 
tarsi. It is found at Kanala in the fields, on grasses and in vegetation.” 


110 I. AHMAD 


The type of this species could not be examined. According to Horn (1926-28), 
the Montrousier collection is supposed to be present in Montpellier Museum (Herault) 
France. M. Jean Claparede, the curator of this museum, wrote saying that there 
was no insect collection there and he would convey my request to the University of 
Montpellier, but no further information has been received. 

However two specimens labelled “ Leptocorisa burmeisteri Montrousier’’ from 
New Caledonia and New Hebrides in the collection of the Paris Museum were 
examined. These represented Leptocorisa acuta (Thunberg). As the title of 
Montrousier’s paper shows, Kanala is in New Caledonia, from where a number of 
specimens have been identified as Leptocorisa acuta. In the light of original descrip- 
tion and type locality it becomes certain that this species is a synonym of L. 
acuta (Thunberg). 


MUTUSCA Stal 
Mutusca Stal, 1866 : 160, 1873 : 86 
Type-species : Leptocorisa prolixa Stal by monotypy. 


Body linear. Head elongated. Antennae with basal segments swollen throughout their 
length, in side view wider beyond middle. Rostrum short, nearly reaching second coxae. 
Pronotum at least twice as long as broad. 

In g, posterior margin of seventh abdominal tergum produced into a medium rounded process 
(Text-fig. 355) ; pygophore rounded, claspers curved at apices, curved portion triangular (Text- 
fig. 357) ; aedeagus with both (dorsal and ventral) pairs of thecal appendages, with a pair of 
lateral conjunctival appendages, a dorsal membranous with a terminal appendage (Text-fig. 
356) and a pair of dorso-lateral conjunctival appendages. Frontal or dorsal conjunctival 
appendages wanting. 

In 9, posterior margin of seventh abdominal sternum emarginate, with a short median split 
and a fold (Text-fig. 368) ; intervalvular sacs absent ; spermatheca rounded, with a small 
twisted tube but without a median flange (Text-fig. 370). 


This genus can easily be separated from other genera of the tribe by the basal 
antennal segments, which are swollen throughout their length. 


KEY TO SPECIES 


1 Basal antennal segment long, at least a third as long again as third segment ; second 
at least slightly longer than third ; paraclypeae thin and straight (Text-fig. 3) ; 
usually a faint black median longitudinal line present on abdominal venter. (Ing, 
pygophore with a concave median posterior margin (Text-fig. 364) ; aedeagus with 
second left lateral conjunctival appendage represented by scattered spines and 
first lateral appendage asymmetrical (Text-fig. 366). In 9, ninth paratergite 
narrow and elongated, at least three times as long as broad (Text-fig. 368)) 
prolixa (p. 113) 
— Basal antennal segment comparatively short, only slightly longer than third ; 
second at least slightly shorter than third ; paraclypeae thick, medially curved 
and converging at apices (Text-fig. 354) ; faint black line on abdominal venter 
usually absent. (In g, pygophore with a pointed apex (Text-fig. 355) ; aedeagus 
with second left lateral conjunctival appendages represented by a bunch of spines, 
all fused together and first lateral appendage symmetrical (Text-fig. 356) ; in 9, 
ninth paratergite broad and comparatively short, about twice as long as broad)) 
brevicornis (p. 111) 


THE LEPTOCORISINAE OF THE WORLD It 


Mutusca brevicornis (Dallas) 
(Text-figs. 354-363) 


Leptocorisa brevicornis Dallas, 1852 : 483 
Mutusca brevicornis (Dallas) Stal, 1866 : 160, 1873 : 86 


Characterized by curved, thick and converging paraclypeae, by short basal an- 
tennal segment, which is only slightly longer than third and by the form of g and 9 
genitalia. 


6. Body length, 14-7-15:2 mm., width, 1-5-1-6 mm. Head length, 2:0-2:1 mm., width, 
I-3-1:33 mm. Antennae, segment I, 2:8-3-:1 mm., II, 2-3-2-4 mm., III, 2-4-2-45 mm., IV, 
2°3-2'4 mm. Rostrum, segment I, 1-3-1-35 mm., II, 1-3-1-35 mm., III, 0-4-0-5 mm., IV, 
0-7 mm. Pronotum, length, 2-4—2-5 mm., width, 1-3-1-5 mm. Hind femora, 4-7-4:8 mm, 
Hind tibia, 4-8-5:2 mm. Distance between ocelli, 0-4-0-5 mm., distance between an ocellus 
and an eye, 0:6—-0-7 mm. 

Body linear, comparatively more robust, usually pale unicolourous. Head with paraclypeae 
thick and medially curved, converging at apices (Text-fig. 354). Antennae with short basal 
segments, at most a quarter as long again as third segment, second always shorter than third. 
Rostrum comparatively short, nearly approaching anterior margins of second coxae. Pronotal 
disc with posterior angles brownish black. Legs pale, with basal half of femora usually tinged 
with green. Hemielytra at base of membrane brownish. Abdomen with dorsum, excluding 
connexiva, brownish black with three black lines ; venter pale, usually with two lateral black 
lines, median longitudinal black line usually absent. Posterior margin of seventh abdominal 
tergum with a median round projection, less punctate and with transverse lines ; pygophore 
rounded with pointed apex (Text-fig. 355). 

6 GENITALIA : Claspers comparatively with a long neck and a small median process (Text- 
fig. 357) ; both dorsal and ventral pairs of thecal appendages present ; second left as a bunch 
of spines, all fused together, first symmetrical ; membranous appendage with a smaller terminal 
appendage (Text-fig. 356), and a pair of dorso-lateral membranous appendages. 

@. Body length, 15-0-16-5 mm., width, 1-5-1:7 mm. Head length, 2-1-2-2 mm., (only 
paraclypeae 0-8 mm.), width, 1-3-1-5mm. Antennae, segment I, 3-0—3:2 mm., II, 2-4-2:5 mm., 
III, 2-43-2:6mm., IV, 2-5-2-6mm. Rostrum, segment I, 1-33 mm., II, 1-33 mm., III, 0-5 mm., 
IV,0-7mm. Pronotum, length, 2-6—2-7 mm., width, 1-5-1-7 mm. Hind femora, 4-7—4-8 mm. 
Hind tibia, 4-9-5:2 mm. Distance between ocelli, 0-4—0-5 mm., distance between an ocellus 
and an eye, 0:6—-0:7 mm. 

Longer than g, otherwise similar, posterior margin of seventh abdominal sternum emarginate, 
with a long median split and a median fold ; ninth paratergites comparatively broad, about 
twice as long as broad (Text-fig. 360). 

Q GENITALIA: First gonocoxae comparatively broad, breadth less than half of length, 
apices rounded (Text-fig. 360) ; intervalvular sacs absent ; spermatheca oval, with a short 
twisted tube. 


Type material. Lectotype g, AUSTRALIA: “‘ New Holland” (J. Byrne), B.M. 
(N.H.) ; paratype, 1 9, AUSTRALIA: “ New Holland ’’, Swan River (Richardson), 
BM.(N-H.). 


Other material. AUSTRALIA: 83 4,532; SoLomon Is.: 14. 


The range of this species extends from the type localities in Western Australia to 
Melville Islands in the north, to New South Wales and Tasmania in the south and 


112 I. AHMAD 

Solomon Islands in the east. The adults have been found on Daviesia, an Australian 
leguminous plant and they have also been collected at light. Adults have been taken at 
most times of the year, but they seem to be in abundance during the summer months 
of November to February. No specimen has been collected in July or August. 


Fics. 354-363. Mutusca brevicornis, 354-357, 3d, 358-363, 2. 354, head, dorsal view ; 
355, terminalia, dorsal view ; 356, aedeagus (inflated), ventral view ; 357, claspers, 
inner and outer views ; 358, first gonopophysis, ventral view ; 359, spermatheca ; 
360, terminalia, ventral view ; 361, first gonocoxa, ventral view ; 362, ovipositor, 


ventral view ; 363, same, dorsal view. 


THE LEPTOCORISINAE OF THE WORLD 113 


This species can easily be separated from M. prolixa (Stal) by examining the median 
posterior margin of pygophore in g, which is pointed in this species and concave in 
prolixa. In 9Q, the ninth paratergite is comparatively short and broad, breadth about 
half of the length in this species, whereas in prolixa it is elongate and breadth is one- 
third of length. 

Of the three specimens recorded by Dallas only two remain in the British Museum 
Collection. One is a § (Brit. Mus. 1844-4) collected by Surgeon J. Byrne, R.N. 
during the voyage of H.M.S. Beagle in North West Australian waters and presented 
to the British Museum by Haslar Naval Hospital in 1844. The other is a 9 collected 
on the Swan River, North West Australia and presented by Sir John Richardson to 
the British Museum in 1843 (Brit. Mus. 1843-14). Both are labelled New Holland. 

The ¢ syntype is here selected as lectotype of this species and has been given a 
blue museum lectotype label and labelled lectotype of Leptocorisa brevicornis Dallas. 


Mutusca prolixa (Stal) 
(Text-figs. 364-372) 


Leptocorisa prolixa Stal, 1859 : 235 
Mutusca prolixa (Stal) Stal, 1873 : 86 


Characterized by thin and straight paraclypeae, by long basal antennal segments, 
which are about a third longer than third segment and by the form of g and 9 
genitalia. 


3. Body length, 14:5-16-0 mm., width, 1-4-1‘6 mm. Head length, 1-9-2-2 mm., (only 
paraclypeae o-8-o-9 mm.), width, 1-2-1:3 mm. Antennae, segment I, 3-4—-3-7 mm., II, 2-4— 
2:°8mm., III, 2:3-2:7 mm., IV, 2-5-2:7mm. Rostrum, segment I, 1-1-1:2 mm., II, 1-1-1-2 mm., 
III, 0-5-0-6 mm., IV, 0-7-0-8 mm. Pronotum, length, 2-3—2-6mm., width, 1-4-1-6mm. Hind 
femora, 4-7-5:'2 mm. Hind tibia, 5:2-5-3 mm. Distance between ocelli, 0-4-0-43 mm., 
distance between an ocellus and an eye, 0:55-0:66 mm. 

Body linear, usually pale, tinged light brown. Head with paraclypeae uniformly thin, 
straight and diverging at apices (Text-fig. 3). Antennae, usually tinged with pink, with long 
basal segment, at least a third as long again as third segment ; second at least slightly longer 
than third. Rostrum usually reaching anterior margin of second coxae. Pronotal disc usually 
without brownish black spots on posterior angles. Legs pale, with femora tinged light green. 
Hemielytra with red subcostal and radial veins. Abdomen, with dorsum brownish black, 
venter pale with a black median longitudinal line. Posterior margin of seventh abdominal 
tergum with a median rounded projection, projection densely punctate ; pygophore rounded 
with median posterior margin concave (Text-fig. 364). 

6 GENITALIA : Claspers comparatively with a short neck and greatly developed median 
process (Text-fig. 365) ; both (dorsal and ventral) pairs of thecal appendages present ; second 
left lateral conjunctival appendage, represented by scattered spines, first lateral asymmetrical ; 
membranous appendage with a large terminal appendage (Text-fig. 366) and a pair of dorso- 
lateral membranous appendages. 

9. Body length, 17-0-18:1 mm., width, 1-6-1-7 mm. Head length, 2-0-2:2 mm., (only 
paraclypeae o-7-0-9 mm.), width, 1-3-1-33 mm. Antennae, segment I, 3-4-4-0 mm., II, 
2-4—2°8 mm., III, 2-3-2:7 mm., IV, 2:5-2:7 mm. Rostrum, segment I, 1:2-1-3 mm., II, 1-2— 
1-4 mm., III, 0-5-0-55 mm., IV, 0-7-0:8 mm. Pronotum, length, 2:5—2-9 mm., width, 1-6— 
1-7 mm. Hind femora, 4:6-5-6 mm. Hind tibia, 4-9-6-:0 mm. Distance between ocelli, 
0-4—0-43 mm., distance between an ocellus and an eye, 0-6—0:66 mm. 

Very similar to g in general appearance but usually longer, posterior margin of seventh 


114 I. AHMAD 


abdominal sternum emarginate, with a short median split and fold (Text-fig. 368) ; ninth 


paratergite elongated, breadth only a third of length. 
2 GENITALIA : First gonocoxae narrow, at least twice as long as broad, with rounded apices 


(Text-fig. 367) ; intervalvular sacs absent (Text-fig. 372) ; spermatheca bladder-shaped, with 
a long twisted tube. 


Fics. 364-372. Mutusca prolixa, 364-366, 3, 367-372, 9. 364, terminalia, dorsal view ; 
365, claspers, inner and outer views ; 366, aedeagus (inflated), ventral view ; 367, first 
gonocoxa, ventral view ; 368,terminalia, ventralview ; 369, first gonopophysis, ventral 
view ; 370, spermatheca ; 371, ovipositor, ventral view ; 372, same, dorsal view. 


THE LEPTOCORISINAE OF THE WORLD 115 


Type material. Holotype, CHINA: (Kzinb), labelled “typus’’, “ prolixus”’ 
abdomen missing, Riksmus. Stockholm. 


Other material. INDIA: Ig; JAVA: 19; MALAYA: 14; AUSTRALIA: 44, 
62; NEW CALEDONIA: I<. 


The range of this species is wide, extending in the west into India, in the south into 
Australia, in the north into China and in the east into New Caledonia. It has been 
collected on grasses, but not many individuals seem to have been found. 


BLOETEOCORIS gen. n. 


Type-species : Leptocorixa (Rhabdocorts) inflexigena Blote 1934. 


Body robust. Head elongated ; genae remarkably developed, with pointed and inflexed 
tips (Text-fig. 8) ; ocelli usually placed very close together ; interocellar distance usually 
about half the distance between an ocellus and an eye (in both the cases including the diameter 
of both). Antennae with apices of basal segments slightly thicker. Rostrum long, reaching 
hind coxae. 

In J, posterior margin of seventh abdominal tergum produced into a median truncated process 
(Text-fig. 376) ; pygophore rounded, lobes of ninth segments prominently developed, usually 
produced into horn-like processes ; median posterior margin pointed ; claspers branched at 
apices into a rectangular and a conical leaf-like portion (Text-fig. 374, A) ; aedeagus with both 
(dorsal and ventral) pairs of thecal appendages and two pairs of dorso-lateral conjunctival 
appendages ; supporting appendages always present but frontal and membranous conjunctival 
appendages absent (Text-fig. 377). 

In 9, posterior margin of seventh abdominal sternum usually with medially pointed apex, 
very small portion of first gonocoxae visible externally (Text-fig. 378) ; intervalvular sacs 
absent ; spermatheca usually flask-shaped, with a long coiled tube but without a median flange. 


This genus resembles Stenocoris and Leptocorisa but can easily be separated from 
them by the pointed and inflexed genae, developed processes of the lobes of the 
ninth segments in the g and in the 2 by very small externally visible first gonocoxae. 


It is named in honour of Dr. H. C. Blote. 

Hsiao (1963) has described a monotypic new genus, Grypocephalus for pallipectus 
new species. His generic description is based on the character of remarkably 
developed juga, which he has shown inflexed and pointed in his diagram of pallipec- 
tus. It is suspected that he has erred in recognizing the developed structure as 
“juga ’’ rather than genae. If this is so, my new genus Bloeteocoris will fall into 
synonymy of Grypocephalus Hsiao. Until the type specimen of G. pallipectus is 


studied, this cannot be ascertained. 


KEY TO SPECIES 


1 Larger species, body length exceeding 14 mm., head dorsally without a black mark 
or spot ; in 4, lobes of ninth segments produced into horn-like processes (Text-fig. 
382) ; in 9, posterior margin of seventh abdominal sternum eens with a small 
projection, without split (Text-fig. 378) . : : 5 : 2 
— Smaller species, body length in g 11-11:5 mm., and in 12, I2-13'2 mm. ; a pair of 
symmetrical trifurcate marks present on the dorsum of the head ; in 3, lobes 
of ninth segments rounded (Text-fig. 386) ; in 9, posterior margin of seventh 
abdominal sternum with a median split (Text-fig. 388) : ; minutus (p. 121) 


I16 I. AHMAD 


2 Ocelli larger, interocellar distance always greater than half the distance between an 
ocellus and an eye ; second antennal segments distinctly longer than third ; in J, 
processes of the lobes of the ninth segments greatly developed, inwardly curved and 

pointed (Text-fig. 376) ; in 9, distal margin of first gonocoxae regular (Text-fig. 

378 : : : : : ; : : inflexigena (p. 117) 


Fics. 373-375. Bloeteocorisspp. A, claspers, inner and outer views ;_ B, first gonocoxae, 
ventral view ; CC, first gonopophysis, ventral view ; D,spermatheca. 373, inflexigena ; 
374, meridianus ; 375, minutus. 


THE LEPTOCORISINAE OF THE WORLD 117 


— Ocelli minute, interocellar distance distinctly less than half the distance between an 
ocellus and an eye; second antennal segments subequal to third ; in dg, the 
processes of the lobes of ninth segment not greatly developed, straight, and rounded 

at apices (Text-fig. 381) ; in 9, distal margin of first gonocoxae irregular (Text-fig. 

383) ‘ meridianus (p. 119) 


Bloeteocoris inflexigena (Blote) comb. n. 
(Text-figs. 373, 376-380) 


Leptocorixa (Rhabdocoris) inflexigena Blote, 1934 : 283 


Characterized by acutely pointed and curved processes of the lobes of the ninth 
segments in the g and in the 2 by pointed median posterior margin of the seventh 
abdominal sternum and by the forms of 3 and 9 genitalia. 


6. Body length, 16-0 mm., width, 2:4mm. Head length, 2:2 mm., width,1-8mm. Anten- 
nae, segment I, 5-3 mm., II, 3:8 mm., III, 3:3 mm., 1V,4-9mm. Rostrum, segment I, 2-0 mm., 
II, 2:1 mm., III, 1:1 mm., IV, 1:6 mm. Pronotum, length, 2-8 mm., width, 2-4 mm. Hind 
femora, 6-4mm. Hind tibia, 7-6mm. Distance between ocelli, 0-5 mm., distance between an 
ocellus and an eye, 0-86 mm. 

Body robust, brown with a greenish tinge. Head large, slightly longer than broad, brownish 
green ; occipital suture medially concave, arms curving outwards towards sides ; paraclypeae 
converging at apices, genae with strongly developed tips. Antennae reddish black, with the 
bases of the apical segments pale. Rostrum long, reaching hind coxae. Disc of pronotum 
brownish green, behind the collar dorsally convex, a fork-shaped brownish black mark present 
anteriorly in the middle ; posterior angles with brownish black spots. Legs brownish black, 
femora tinged with green. The subcostal and radial veins of hemielytra brownish black. 
Abdomen dorsally pale, tinged with brown, a portion of seventh segment and pygophore black, 
venter pale unicolourous. Posterior margin of seventh abdominal tergum produced in middle, 
posterior margin of this projection medially concave ; lobes of ninth segments horn-shaped, 
inwardly curved and acutely pointed at apiecs (Text-fig. 376). 

6 GENITALIA : Claspers with leaf-like portions narrow and elongated (Text-fig. 373, A) ; 
both dorsal and ventral pairs of thecal appendages present, ventral triangular (Text-fig. 377) ; 
a pair of symmetrical and small lateral appendages (Text-fig. 377) ; vesica, long coiled tube 
with “ T”’ structures and a pair of supporting appendages (Text-fig. 377). 

. Very similar to g in general appearance, colour and measurements, only slightly larger ; 
posterior margin of seventh abdominal sternum medially produced with a pointed apex (Text- 
fig. 378). 

Q GENITALIA: First gonocoxae largely covered externally by seventh abdominal sternum, 
slightly longer than broad, posterior margin broader than anterior margin, outer margin smooth, 
convex in middle (Text-fig. 373, B) ; intervalvular sacs absent (Text-fig. 380) ; spermatheca 
with an uniformly thick and smoothly curve neck (Text-fig. 373, D). 


Type material. Holotype g, Burma : Moulmein, 12.v.1929, Mus. Leiden ; allo- 
type 9, data as holotype, Mus. Leiden. 


This species seems to be comparatively rare. 

The 3 can be separated from the closely allied species meridianus by curved and 
pointed processes of the lobes of ninth segments and the 2 by pointed median 
posterior margin of seventh abdominal sternum. 


118 


I. AHMAD 


Fics. 376-380. Bloeteocoris inflexigena, 376-377, 3, 378-380, 2. 376, terminalia, dorsal 
view ; 377, aedeagus (inflated), ventral view ; 


ovipositor, ventral view ; 


380, same, dorsal view. 


378, terminalia, ventral view ; 379, 


THE LEPTOCORISINAE OF THE WORLD 119 


Bloeteocoris meridianus sp. n. 
(Text-figs. 374, 381-385) 


6 characterized by the straight processes of the lobes of the ninth segment and 2 
by the small and rounded projection of the seventh abdominal sternum and by the 
form of genitalia. 


3. Body length, 15:0mm., width, 2-3mm. MHead length, 2-omm., width, 1-7mm. Anten- 
nae, segment I, 5:2 mm., II, 3:5 mm., III, 3-46 mm., IV, 4:3 mm. Rostrum, segment I, 
1-9 mm., II, 1-9 mm., III, t:omm., IV, 1-45 mm. Pronotum, length, 2-9 mm., width, 2:3 mm. 
Hind femora, 6:0 mm. Hind tibia, 7-6 mm. Distance between ocelli, 0-4 mm., distance 
between an ocellus and an eye 0-9 mm. 

Body robust, pale with a brownish tinge. Head large, slightly longer than broad ; occipital 
suture medially concave, slightly “‘ V ’’-shaped behind ocelli, arms curving outwards towards 
sides, paraclypeae diverging at apices ; genae with inflexed tips. Antennae reddish brown, 
with second and third segments tinged black and the bases of the apical segments pale. 
Rostrum long, reaching hind coxae. Pronotal disc palish brown, not remarkably convex 
behind collar, fork-shaped, brownish black mark absent ; posterior angles with brownish black 
spots. Legs palish brown, with tibiae and tarsi darker. Subcostal and radial veins of hemiely- 
tra brownish black. Abdomen with dorsum pale, tinged with brown; venter pale uni- 
colourous ; a portion of seventh segment and pygophore black. Posterior margin of seventh 
abdominal tergum medially produced, projection with a truncated margin ; pygophore with 
lobes of the ninth segments produced into straight processes, rounded at apices (Text-fig. 381). 

6 GENITALIA: Claspers with leaf-like portion broad and short (Text-fig. 374, A) ; both 
(dorsal and ventral) pairs of thecal appendages present, ventral elongated (Text-fig. 382) ; a 
pair of symmetrical lateral larger conjunctival appendages (Text-fig. 382) ; two pairs of dorso- 
lateral conjunctival appendages as in Text-fig. 382 ; vesica thin, long coiled tube with ‘“‘ T”’ 
structures and a pair of supporting appendages (Text-fig. 382). 

9. Body length, 16-5 mm., width, 2-5-2-6mm. Head, length, 2-1-2-:2 mm., width, 1-8 mm. 
Antennae, segment I, 5:1 mm., II, 3:5 mm., III, missing, IV, missing. Rostrum, segment I, 
2:0 mm., II, 2-0 mm., III, 1:0 mm., IV, 1-5 mm. Pronotum, length, 3-0-3-1 mm., width, 
2°5-2°6mm. Hind femora, 6-4 mm. Hind tibia, 7-7mm. Distance between ocelli, 0-4 mm., 
distance between an ocellus and an eye, 0-9 mm. 

Slightly larger than g, otherwise very similar ; posterior margin of seventh abdominal 
sternum with a rounded median projection (Text-fig. 383). 

@ GENITALIA: First gonocoxae externally almost entirely covered by seventh sternum, 
about as long as broad, posterior margin broader than anterior margin, outer margin sinuate 
(Text-fig. 374, B) ; intervalvular sacs absent (Text-fig. 384) ; spermatheca with a thin tapering 
neck (Text-fig. 374, D). 


Holotype 3, THAILAND: Trang. Banchong “ at light’, 26.iv.1924 (I. H. Evans), 
Pix F. M. S. Museum ”’, B.M.(N-H.). 


Paratypes. Maraya: Perak, Gunong Kledang, 2 ¢ xi.1916, “Ex F. M. S. 
Museum ”’, B.M.(N.H.). 


This species has most southernly range in the genus Bloeteocoris. All three 
Bloeteocoris species would appear to be allopatric. B. minutus in south China, north 
India and north east Burma, B. inflexigena in south Burma and B. meridianus in 
Thailand and Malaya. 


This species is very close to inflexigena, but can easily be separated by smaller 


120 I. AHMAD 


interocellar distance (less than half of the distance between an ocellus and an eye),and 
by the straight processes of the lobes of the ninth segment. 


Fics. 381-385. Bloeteocoris meridianus, 381-382, 5, 383-385, 2. 381, terminalia, dorsal 
view ; 382, aedeagus (inflated), ventral view ; 383, terminalia, ventral view ; 384, 
ovipositor, dorsal view ; 385, same, ventral view. 


THE LEPTOCORISINAE OF THE WORLD 121 


Bloeteocoris minutus sp. n. 
(Text-figs. 375, 386-390) 


Characterized by small size of the body, by a pair of trifurcate dorsal black marks 
on the middle of head and by the form of 3 and 9 genitalia. 


6. Body length, 11-0-11-5 mm., width, 1:7-1:-9 mm. Head, length, 1-5-1-6 mm., width, 
I-3-I'4 mm. Antennae, segment I, 2:6-2:7 mm., II, 1-9-2:1 mm., III, 1-9-2:1 mm., IV, 
2°4-2:°8 mm. Rostrum, segment I, 1-4-1-6 mm., II, 1-4-1-6 mm., III, 0-7-0-9 mm., IV, 
I-o-I'2mm. Pronotum, length, 1-9-2:1 mm., width, 1:7-1-9 mm. Hind femora, 3-5—3-:7 mm. 
Hind tibia, 4-3-4-6 mm. Distance between ocelli, 0-3-0-4 mm., distance between an ocellus 
and an eye, 0:6-0-7 mm. 

Body short, pale, tinged black. Head large, slightly longer than broad, with a pair of 
symmetrical trifurcate dorsal black marks between compound eyes ; postoccipital region 
black ; occipital suture concave in middle, arms curving outwards towards sides ; paraclypeae 
converging at apices; genae with inflexed tips. Antennae brownish black, unicolourous. 
Rostrum very long, extending beyond the apices of third coxae. Pronotal disc with brownish 
black punctures, margins and an oval portion in middle pale ; posterior angles with brownish 
black spots. Legs pale unicolourous, tarsi slightly darker. R-M vein of the hemielytra 
brownish black. Abdomen with dorsum pale, tinged with red, venter palish brown. Posterior 
margin of seventh abdominal tergum with a median rounded projection ; pygophore with lobes 
of ninth segments rounded (Text-fig. 386). 

6 GENITALIA: Apices of claspers slightly concave in middle with small leaf-like portion 
(Text-fig. 375, A) ; both dorsal and ventral pairs of thecal appendages present, ventral pair 
triangular (Text-fig. 387) ; a pair of symmetrical lateral conjunctival and two asymmetrical 
pairs of dorso-lateral appendages present (Text-fig. 388) ; vesica thin and coiled tube with “ T”’ 
structures and a pair of supporting appendages. 

9. Body length, 12-0-13:2 mm., width, 2-o-2:2 mm. Head length, 1-7-1-9 mm., width, 
I-4-I-5 mm. Antennae, segment I, 2:9-3:2 mm., II, 2-3-2-4 mm., III, 2-2-2:3 mm., IV, 
2:6-2:'8 mm. Rostrum, segment I, 1-:7-1:8 mm., II, 1:7-1-9 mm., III, 0-7-1:0 mm., IV, 
I-2-1:3mm. Pronotum, length, 2:2-2:7 mm., width, 2-0-2-2 mm. Hind femora, 4-3—4:7 mm. 
Hind tibia, 4-8-5-6 mm. Distance between ocelli, 0-4 mm., distance between an ocellus and an 
eye, 0-7—-0'8 mm. 

Very similar to f in general appearance but distinctly larger ; posterior margin of seventh 
abdominal sternum with a short median split and a conspicuous median fold (Text-fig. 388). 

Q GENITALIA : First gonocoxae largely covered by seventh abdominal sternum, triangular 
with rounded apices (Text-fig. 375, B) ; intervalvular sacs absent (Text-fig. 390) ; spermatheca 
kidney-shaped (Text-fig. 375, D). 


Holotype gj, N.E. Burma : Kambaiti, 7,000 ft., 12.1v.1934 (R. Malatse), labelled 
“ Leptocorisa costalis H. S., det. Kiritshenko ’’, Riksmus. Stockholm. 


Paratypes. 4 3, 4 Q, locality as holotype, iv—v.1934, Riksmus. Stockholm ; 
Cuina : Kunming, Yunnan, I f 24.vii.1945 (Cazier), AAM.N.H. ; Inp1A : Daryjiling, 
I g, Mus. Hist. nat. Paris, 2 g, 1 9, B.M.(N.H.). 


The range of this species extends from N.E. India in the west into southern China 
(Kunming, Yunnan) and N.E. Burma. Specimens have been collected in April, 
May and July. 


This is an isolated species in the genus and can easily be separated by its small size 
and characters of genitalia (see above). 


122 I. AHMAD 


The diagram of the lateral side of the head of G. pallipectus Hsiao (1963) resembles 
B. minutus and if he has erred in diagnosing juga for genae, B. minutus will fall as a 
synonym of G. pallipectus. I have written to Dr. Hsiao for the loan of his type 


material but so far have had no reply. 


Fics. 386-390. Bloeteocoris minutus, 386-387, g, 388-390, 2. 386, terminalia, dorsal 
view ; 387, aedeagus (inflated), ventral view ; 388, terminalia, ventral view ; 389, 
Ovipositor, ventral view ; 390, same, dorsal view. 


THE LEPTOCORISINAE OF THE WORLD 123 


NOLIPHINI trib. n. 


Body large and robust. Head broader than long, paraclypeae poorly developed, distinctly 
shorter than and never enclosing clypeus ; head behind the eyes very strongly narrowed and 
turned up towards pronotum. Pronotum broader than long, trapeziform and convex. 
Aedeagus with both dorsal and ventral pairs of thecal appendages, simple. Spermatheca 
usually tubular, sometimes triangular, never with a median flange. 


The characters of the anterior region of the head and the form of the pronotum 
clearly separate this tribe from Leptocorisini. 


LYRNESSUS Stal 


Lyrnessus Stal, 1862 : 59, 1865 : 185, 1866 : 159, 1868 : 54, 1873 : 86 
Coristenia Costa, 1863 : 261 


Type-species : Lyrnessus geniculatus (Guerin, 1857) by monotypy. 


Body robust. Head triangular. Antennae with apices of basal segments slightly thicker. 
Rostrum long, reaching third coxae. Pronotal disc trapeziform, without latero-posterior 
spines (Text-fig. 7) and without black sculpturation. Hemielytra brownish black with costal 
margins and veins prominently pale. 

In 4, posterior margin of seventh abdominal tergum truncated, medially only slightly concave 
(Text-fig. 392) ; claspers strongly curved, apically acutely pointed, with a median spine (Text- 
fig. 393) ; aedeagus with both (dorsal and ventral) pairs of thecal appendages, asymmetrical 
pair of lateral conjunctival appendages and a pair of fused supporting appendages (Text-fig. 
393). 

In 9, posterior margin of seventh abdominal sternum emarginate, with a median fold (Text- 
fig. 394) ; first gonocoxae large with round apices (Text-fig. 399) ; three pairs of intervalvular 
sacs, spermatheca tubular, curved in middle, with a long coiled tube. 


The only species of this genus, L. geniculatus (Guérin), is found in Neotropical region, 
whereas all the other species of the tribe Noliphini are confined to Oriento-Australian 
region, thus it is not surprising that it shows isolation in many characters. It lacks 
the fore femoral spines of Cosmoleptus and latero-posterior spines on pronotum of 
Noliphus. It also resembles Stenocoris (Pseudoleptocorisa) erratica Blote in the 
characters of the colour of the hemielytra. 


Lyrnessus geniculatus (Guérin) 
(Text-figs. 391-399) 


Leptocorisa geniculata Guérin, 1857 : 392 
Lyrnessus geniculatus (Guérin) Stal, 1866 : 159 
Paryphes tibialis Stal, 1860 : 31 

Coristenia flavicosta, Costa, 1863 : 261 


Characterized by pale basal quarter of apical antennal segments, by dark brownish 
black apices of femora, entire tibia and tarsi and by the form of 3 and 9 genitalia. 


6. Body length, 12-6-14-6 mm., width, 2:2-2:7 mm. Head length, 1-3-1-4 mm., width, 
I-4-1'7 mm. Antennae, segment I, 3-0-3-7 mm., II, 2:1-2:8 mm., III, 2:4-3-0 mm., IV, 
3:°9-411 mm. Rostrum, segment I, 1-4-1-6 mm., II, 1-4-1-7 mm., III, 0-6-0-7 mm., IV, 
0-9-I1'Imm. Pronotum, length, 2-1-2-6 mm., width, 2:2-2-7 mm. Hind femora, 4:2—5-5 mm. 


124 


I. AHMAD 


FIGS. 391-399. 


view ; 392, claspers, inner and outer views ; 


Lyrnessus geniculatus, 391-392, 3, 394-399, Q. 


394, terminalia, ventral view ; 395, ovipositor, dorsal view ; 


ventral view ; 
spermatheca. 


391, terminalia, dorsal 


393, aedeagus (inflated), ventral view ; 


396, first gonopophysis, 


397, Ovipositor, ventral view ; 3098, first gonocoxa, ventral view ; 399, 


THE LEPTOCORISINAE OF THE WORLD 125 


Hind tibiae, 4-9-6-6 mm., interocellar distance, 0-4—0-5 mm., distance between an ocellus and 
an eye, 0-7—-0-9 mm. 

Body robust. Head pale, with a black spot behind ocelli and one on clypeus at labral base ; 
distinctly broader than long ; ocelli large, interocellar distance slightly more than half the 
distance between an ocellus and an eye. Antennae brownish black, with bases of second, third 
and at least a quarter basal portions of apical segments pale. Rostrum long, reaching hind 
coxae with third segment shortest. Lateral black line extending from sides of antenniferous 
tubercle on to anterior one-fourth of propleuron. Pronotal disc posteriorly tinged green, 
sometimes pale unicoloured, with backward triangular processes on either side, brownish black 
spots on posterior angles present (Text-fig. 7). Legs with femora pale, tinged with green, 
apices of femora, entire tibiae and tarsi brownish black. Hemielytra brownish black with pale 
veins. Abdomen with dorsum yellowish brown, venter pale, unicoloured. Posterior margin of 
seventh abdominal tergum truncated (Text-fig. 391), medially slightly concave ; pygophore 
round, with narrow emarginated posterior margin. 

6 GENITALIA : Claspers strongly curved in middle, apically acutely pointed, with a smaller 
spine near apices and a larger in middle (Text-fig. 392) ; aedeagus with both (dorsal and ventral) 
pairs of thecal appendages as in Text-fig. 393 ; lateral appendages asymmetrical (Text-fig. 
393) ; vesica with a pair of fused supporting appendages. 

9. Very similar to f in general appearance and measurements ; posterior margin of seventh 
abdominal sternum emarginate, with a median fold (Text-fig. 394). 

Q GENITALIA : First gonocoxae large, round at apices with regular external margin (Text- 
fig. 398) ; three pairs of intervalvular sacs (Text-fig. 395) ; spermatheca tubular (Text-fig. 399). 


Type material. Holotype 2 of Paryphus tibialis Stal, BRAziL : (F. Schilb), Riks- 
mus. Stockholm ; MExIco : Oaxaca, 1 g (Sallé), with a label “‘ Leptocorisa geniculata 
Guérin (M. M. Type) ’’, Riksmus. Stockholm. 


Other material. CuBA : 24,29; Brazit : 3g,792; Peru: 34,19; PANAMA: 
ie, 1 25 “PARAGUAY : 1 9. 


The range of this species extends in the north into Mexico and in the south into 
Paraguay. The adults have been collected in March, September, October and 
November. 

In spite of enquiries at various Italian museums, it has not been possible to trace 
the type of Leptocorisa geniculata Guérin or Coristenia flavicosta Costa. It is possible 
that the specimens from the Riksmuseum, Stockholm with labels “‘ Typ” and 
“ Leptocorisa geniculata Guerin’’ could have been the type. However, although 
Guérin’s original description was based entirely on colour characters, this and the 
figure make it obvious that the present concept of the species (which is also Stal’s) is 
correct. This view is strengthened by the identification of a few specimens from the 
type locality (Cuba) as this species. In 1868 Stal synonymized Coristenia Costa, 1863 
with Lyrnessus Stal, 1860. Costa in his original description states ‘“‘ Luteo testacea, 
punctata, antennis nigris articulis 2 et 3 summa basi, 4 annulo latiore prope basi 
un albidis. Capitis vitta utrinque laterali, maculaque pone ocellos nigris ; 
pronote disc postico virescente, punctis, humeralibus nigris, elytrorum corio fusco 
virescenti venis discoidalibus fulvis, vena juxta marginata conspicua flava ; tibiis 
tarsique fuscis, illis annulo minuto basi pallido, long. mill. 13; lat. mill. 2}. Patria 
?”’. This description agrees in all details with that of L. geniculata and as neither 
the type specimen nor the type locality is known, it is proposed to accept Stal’s 
synonymy. 


126 I. AHMAD 


COSMOLEPTUS Stal 
Cosmoleptus Stal, 1873 : 86-87 


Type-species : Lyrnessus limbaticollis Stal by monotypy. 


Body usually robust. Head triangular. Antennae with apices of basal segments slightly 
thicker. Rostrum long, approaching hind coxae. Pronotal disc trapeziform with black 
sculpturation, lateral and anterior margins pale, impunctate (Text-fig. 4). Posterior angles 
without spines. Legs with fore femora armed with ventro-posterior spines (Text-fig. 9). 
Hemielytra, cuneus with a white spot. 

In 3, posterior margin of seventh abdominal tergum truncated, medially slightly concave 
(Text-fig. 403) ; pygophore rounded ; claspers with pointed apices (Text-fig. 405) ; aedeagus 
with dorsal and ventral! pairs of thecal appendages, a pair of fused supporting appendages and 
membranous appendage with usually a terminal appendage (Text-fig. 406) ; frontal conjunc- 
tival and dorsal appendages wanting. 

In 9, posterior margin of seventh abdominal sternum emarginate, with a median fold (Text- 
fig. 408) ; first gonocoxae large, usually two pairs of intervalvular sacs present (Text-fig. 411). 
Spermatheca tubular (Text-fig. 407). 


This genus is close to Noliphus but can be separated by the absence of spines on 
posterior angles of pronotum. Its fore femoral spines are unique. 


KEY TO SPECIES 


1 Abdomen ventrally pale, unicoloured, with a series of brownish black spots on either 
side ; disc of pronotum dark brown with smooth brownish black dots posteriorly 
on either side ; hemielytra brownish black with prominently pale veins, fore 
femoral spines pale, only brownish at apices ; hind femora pale unicolourous 
sumatranus (p. 129) 
— Abdomen ventrally reddish brown, brownish black spots absent ; disc of pronotum 
dark brownish black, without distinct brownish black dots ; hemielytra palish 
brown, unicoloured ; entire fore femoral spines brownish black ; hind femora 
usually brownish black at apices c : : : ; 3 : : 
2 Smaller species (13-7-14-0 mm.) ; pronotum short, about two-thirds of basal antennal 
segments, with anterior quarter pale ; in g, posterior margin of seventh abdominal 
tergum truncate (Text-fig. 400) : ‘ : : bakeri (p. 126) 
— Larger and robust species (16-2-17-2 mm.) ; pronotum long, about three-quarters of 
basal antennal segments, with entire disc behind collar except sides dark brownish 
black ; in g, posterior margin of seventh abdominal tergum emarginate (Text-fig. 
403) . : : : : . : ; , : . limbaticollis (p. 128) 


Cosmoleptus bakeri sp. n. 
(Text-figs. 400-402) 


Characterized by small size, short pronotum, about two-thirds of basal antennal 
segment, with anterior fourth pale and by ¢ genitalia. 


3. Body length, 13-7-14-0 mm., width, 2:7-2-9 mm. Head length, 1:45-1:55 mm., width, 
1-7-1:8 mm. Antennae, segment I, 3-6-3-9 mm., II, 2:9-3-1 mm., III, 2-9-3-0 mm., IV, 
missing. Rostrum, segment I, 1-8-1:85 mm., II, 1-8-1-85 mm., III, 0-6-0-7 mm., IV, 1-o— 
1:05 mm. Pronotum, length, 2-4-2-5 mm., width, 2-7-2-9 mm. Hind femora, 6-0—-6:2 mm. 
Hind tibia, 6-9-7-0 mm. Distance between ocelli, 0:45—-0-5 mm., distance between an ocellus 
and an eye, 0-8—o-85 mm. 


THE LEPTOCORISINAE OF THE WORLD 127 


Body short. Head with ocelli pale, tinged pink at base ; distance between an ocellus and an 
eye usually only slightly less than twice the distance between two ocelli. Antennae brownish 
black, with basal portions of first segments pale. Rostrum approaching hind coxae. Lateral 
black line extending from collar of propleuron up to metapleuron. Disc of pronotum brownish 
black, with about anterior quarter pale, brownish black dots on posterior angles indistinct. 
Legs with fore femoral spines entirely black, all femora pale, but usually brownish black at 
apices, tibiae and tarsi dark brown. NHemielytra palish brown, with veins of similar colour, 
with pale spots pointing posteriorly. Abdomen ventrally palish brown, ventro-lateral spots 
absent. Posterior margin of seventh abdominal tergum truncate ; posterior margin of pygo- 
phore truncate (Text-fig. 400). 

$ GENITALIA: Claspers pointed at apices, points sharp, smoothly tapering (Text-fig. 401). 
Aedeagus with both dorsal and ventral pairs of thecal appendages ; a single left lateral conjunc- 
tival appendage long and thin (Text-fig. 402) ; a pair of asymmetrical supporting appendages, 
left longer and thinner ; a dorsal membranous appendage with a very small rounded terminal 
appendage. 

No 9 available for study. 


Holotype 3, PHILIPPINE ISLANDS : Mindanao-Davao (Baker), Mus. Hels. 
Paratype, I g, same data as holotype, B.M.(N.H.). 


This species can easily be separated from limbaticollis (Stal) and swmatranus Blote 
by its small size and short pronotum. 


Fics. 400-402. Cosmoleptus bakeri, 3. 400, terminalia, dorsal view ; 401, 
claspers, outer view ; 402, aedeagus (inflated), ventral view. 


128 I. AHMAD 


Cosmoleptus limbaticollis (Stal) 
(Text-figs. 403-411) 


Lyrnessus limbaticollis Stal, 1865 : 185 
Cosmoleptus limbaticollis (Stal) Stal, 1873 : 87 


Characterized by large and robust body, long pronotum, about three-fourths of 
basal antennal segments, with entire disc behind collar except the sides brownish 
black and by g and @ genitalia. 


THE LEPTOCORISINAE OF THE WORLD 129 


6. Body length, 16-2-17:2 mm., width, 3-1-3-3 mm. Head length, 1-9-2-0 mm., width, 
2°0-2:1 mm. Antennae, segment I, 3:8-4:2 mm., II, 2-9-3:2 mm., III, 2-9-3:1 mm., IV, 
5°4-6-2 mm., (pale portion, 2-5-3-0 mm.). Rostrum, segment I, 1-9-2-0 mm., II, 1-9-2: mm., 
III, 0-8-o-9 mm., IV, 1-1-1:2 mm. Pronotum, length, 3:0-3:2 mm., width, 3-I-3:3 mm. 
Hind femora, 6-7-7-2 mm. Hind tibiae, 7-8-8:2 mm. Distance between ocelli, 0-5-0-55 mm., 
distance between an ocellus and an eye, 1:0—-1-I mm. 

Body large and robust. Head with ocelli usually pink with brownish black rim, distance 
between an ocellus and an eye about twice the distance between two ocelli. Antennae dark 
brown, with inner sides of basal segments and at least three-sevenths of basal portion of apical 
segments pale. Rostrum hardly approaching hind coxae. Entire pro-, meso-, and meta- 
pleuron and sternum, except coxae, dark brownish black. Entire disc of pronotum behind 
collar, except sides, dark brownish black, posterior angles without brownish black dots. Legs 
with fore femoral spines entirely dark brown, all femora pale, with brownish black apices, 
tibiae and tarsi palish brown. NHemielytra dark brown with palish veins, pale spots at base of 
membrane almost quadrilateral. Abdomen ventrally palish brown, brownish black ventro- 
lateral spots absent. Posterior margin of seventh abdominal tergum medially emarginate and 
laterally convex, posterior margin of pygophore truncated (Text-fig. 403). 

6 GENITALIA : Claspers pointed at apices, points uniformly thick (Text-fig. 405). Aedeagus 
with both (dorsal and ventral) pairs of thecal appendages ; a single left lateral conjunctival 
appendage present, small and thin (Text-fig. 406) ; a pair of asymmetrical supporting append- 
ages, left large, curved and cup-shaped and a dorsal membranous appendage, terminal appendage 
wanting (Text-fig. 406). 

2. Very similar to g but usually slightly smaller in size. Posterior margin of seventh 
abdominal sternum truncated, with a median fold (Text-fig. 408). 

Q GENITALIA: First gonocoxae pointed at apices, broad, covering a larger portion of eighth 
paratergite (Text-fig. 409) ; three pairs of intervalvular sacs (Text-fig. 411) ; spermatheca like 
an elongated tube with a small, uniformly thick, coiled tube (Text-fig. 407). 


Type material. Holotype g, NEw Guinea: “Stal’’, “ limbaticots’’, Riksmus. 
Stockholm, allotype 9, data as holotype, Riksmus. Stockholm ; paratypes, NEW 
GuINEA : Ins. Aru, 1 g, Mysol (Stevens), Riksmus. Stockholm. 


Other material. NEW GUINEA: 34, 4 9. 


This species is restricted to New Guinea. Its range is the most easterly in the 
genus. The two other allopatric species are similarly restricted, swmatranus to 
Sumatra and baker: to the Philippine Islands. 


Cosmoleptus sumatranus Blote 
(Text-figs. 412-420) 
Cosmoleptus sumatranus Blote, 1934 : 285 


Characterized by uniformly pale venter of abdomen with a series of brownish black 
spots on either side, by the latero-posterior smooth brownish black dots on prono- 
tum and by ¢ and @ genitalia. 


Fics. 403-411. Cosmoleptus limbaticollis, 403, 405-406, 3, 404, 407-411, 9. 403, term- 
inalia, dorsal view ; 404, first gonocoxa, ventral view ; 405, claspers, outer view ; 
406, aedeagus (inflated), ventral view ; 407, spermatheca ; 408, terminalia, ventral 
view ; 409, Ovipositor, ventral view ; 410, first gonopophysis, ventral view ; 4II, 
ovipositor, dorsal view. 


130 I. AHMAD 


6: Body length, 15-7 mm., width,3:2mm. Head length, 1-7mm., width, 1-9 mm. Anten- 
nae, segment I, 4-0 mm., II, 3:3 mm., III, 3:2 mm., IV, 4-9 mm. (pale, 2-0 mm.). Rostrum, 
segment I, 1-7 mm., II, 1-8 mm., III, 0-8 mm., IV, 1: mm. Pronotum, length, 3-0 mm., 


FIGs. 412-420. Cosmoleptus sumatvranus, 412, 413, 415, 3, 414, 416-420, 9. 412, term- 
inalia, dorsal view ;_ 413, claspers, inner and outer views; 414, spermatheca; 415, 
aedeagus (inflated), ventral view ; 416, first gonocoxa, ventral view ; 417, terminalia, 
ventral view ; 418, ovipositor, ventral view ; 419, first gonopophysis, ventral view ; 
420, Ovipositor, dorsal view. 


THE LEPTOCORISINAE OF THE WORLD 131 


width, 3:2mm. Hind femora,6-4mm. Hind tibia, 7-4mm. Distance between ocelli, 0-5 mm 
distance between an ocellus and an eye, 0-9 mm. 

Body linear. Head with ocelli pale with reddish rim, distance between an ocellus and an eye 
usually less than twice the distance between two ocelli. Antennae dark brown, with basal 
portions of first and about two-fifths of apical segment pale. Rostrum approaching hind coxae. 
Lateral black line extending from posterior margin of eyes up to metapleuron. Disc of pronotum 
dark brown, with about anterior third pale, smooth brownish black dots on posterior angles 
present. Legs with fore femoral spines pale, only brownish black at apices, all femora pale, 
tibiae and tarsi brown. Hemielytra brownish black with prominently pale veins, pale spots at 
base of membrane pointing anteriorly. Abdomen ventrally pale, unicolourous, with a series of 
brownish black spots on either side. Posterior margin of seventh abdominal tergum truncate ; 
posterior margin of pygophore emarginate (Text-fig. 412). 

6 GENITALIA : Claspers pointed at apices, points knob-like, thicker at base (Text-fig. 413) ; 
aedeagus with both (dorsal and ventral) pairs of thecal appendages, a single left lateral conjunc- 
tival appendage flat ; a pair of asymmetrical supporting appendages, left larger and broader, 
and a dorsal membranous appendage with a terminal pyramid-shaped appendage (Text-fig. 415). 

2. Very similar to male in general appearance and measurements. Posterior margin of 
seventh abdominal sternum sinuately emarginate (Text-fig. 417). 

@ GENITALIA: First gonocoxae round at apices, narrow, not covering entire eighth para- 
tergite (Text-fig. 417) ; two pairs of intervalvular sacs (Text-fig. 420) ; spermatheca more or 
less tubular, with a very long tube of variable thickness (Text-fig. 414). 


Type material. Holotype jg, SUMATRA : Tandjong, Morawa, Serdang (B. Hagen), 
labelled ‘‘ Cosmoleptus sumatranus det Bléte’”’, Mus. Leiden ; allotype 9, data as 
holotype, Mus. Leiden. 


This species can easily be separated from limbaticollis (Stal) and bakeri sp. n. by 
the characters of latero-posterior smooth brownish black dots on protoum and by a 
series of ventro-lateral brownish black spots on uniformly pale venter of abdomen. 
This species also resembles Lyrnessus geniculatus (Guérin) and Stenocoris (Pseudo- 
leptocorisa) erratica (Blote) in having brownish black hemielytra with prominently 
pale veins. 


NOLIPHUS Stal 
Noliphus Stal, 1858 : 440, 1873 : 86-87 
Type-species : Noliphus erythrocephalus Stal by monotypy. 


Body usually robust. Head triangular. Antennae with apices of basal segments slightly 
thicker. Rostrum long, reaching hind coxae. Pronotal disc trapeziform, posterior angles with 
strong and curved spines (Text-fig. 6). 

In 3, posterior margin of seventh abdominal tergum truncated, sometimes slightly medially 
produced (Text-fig. 439), always concealing most of pygophore ; claspers with pointed apices ; 
aedeagus with both (dorsal and ventral) pairs of thecal appendages, membranous appendage 
with a long and curved dorsal appendage and usually two pairs of lateral conjunctival append- 
ages (Text-fig. 441) ; frontal conjunctival and terminal appendages wanting. 

In 9, posterior margin of seventh abdominal sternum emarginate, with a median fold (Text- 
fig. 444) ; first gonocoxae usually large ; intervalvular sacs absent ; spermatheca conical, with 
a short tube (Text-fig. 443). 


This genus can be separated from other genera of the tribe Noliphini by the strong 
and curved spines on the posterior angles of the pronotum. 


132 


I. AHMAD 


Key TO SPECIES 


Entire propleuron and pronotum shining black ; apical antennal segments long, 
usually half as long again as third é ‘ é 
Propleuron and pronotum light brown, sometimes amped and arte apical antennal 
segments short, never half as long again as third 5 : . 
Venter of abdomen pale brown, unicolourous ; ocelli pale Pac unicoloured, 
minute, interocellar distance less than half the distance between an ocellus and an 
eye. (Posterior margin of seventh abdominal tergum truncated, slightly medially 
produced (Text-fig. 448) ; in aedeagus, second right lateral conjunctival appendage 


partly membranous (Text-fig. 450)) : : : insularis (p. 


Venter of abdomen always with black spots or ae ree ; cell with conspicuous 
brownish black margins, large interocellar distance, usually more than half the 
distance between an ocellus and an eye : - : : : - 

Large and robust species ($f 13-5-15:2 mm., 2 15-2-16-0 mm.) ; venter of abdomen 
with four black transverse lines ; usually basal halves of apical antennal segments 
pale. (Posterior margin of seventh abdominal tergum truncate (Text-fig. 451) ; 
in aedeagus, frontal conjunctiva with bunches of spines (Text-fig. 452) ; in the 2 


first gonocoxae triangular, with rounded apices (Text-fig. 457)) . papuensis (p. 


Small species (g 12-4-13-2 mm., 9 12-8-13-6 mm.) ; black transverse lines on venter 
of abdomen absent ; only small basal portions of apical antennal segments pale 
A pair of black dots present anteriorly on venter of abdomen ; basal antennal seg- 
ments slightly shorter than third ; ocelli comparatively distant from each other, 
interocellar distance about two-thirds of the distance between an ocellus and an eye. 
(Claspers symmetrically placed (Text-fig. 461) ; in the aedeagus, second right lateral 
conjunctival appendage only slightly notched near apex (Text-fig. 466) ; sperma- 


theca elongated, with smooth curved neck (Text-fig. 465)) . : spinosus (p. 


Round black dots absent, but two black marks present on either side, being a con- 
tinuation of the connexival colour on to venter of abdomen ; basal antennal seg- 
ments slightly longer than third ; ocelli comparatively close together, interocellar 
distance about half the distance between an ocellus and an eye. (Claspers asym- 
metrically placed (Text-fig. 440) ; second right lateral conjunctival appendage 
distinctly bifid at apex (Text-fig. 44) ; spermatheca curved, with a transverse 


neck (Text-fig. 443)) ; : .  erythrocephalus (p. 


Species of very small size (8-4—-10 mm. in teaethy = pagal antennai segments very 
short, subequal to second ; third rostral segment comparatively longer, more than 
half of second. (In g, backward lateral projections of seventh abdominal tergum 
inconspicuous ; aedeagus with long and curved dorsal appendage, rounded at 
apex ; in 9, spermatheca almost tube-shaped, with thumb-like base (Text-fig. 434)) 


discopterus (p. 


Species larger (12:0-13°5 mm.) ; basal antennal segments distinctly longer than 
second ; third rostral segment Oa et short, paooee half the length of 
the second 

A pair of black dots present saterioey on eck ade of venter of abdomene Prcmiclynes 
usually pale, unicolourous. (Posterior margin of g seventh abdominal tergum with 
backward spine-like projections on either side (Text-fig. 421) ; in aedeagus, frontal 
conjunctiva with usually two or three spines (Text-fig. 425) ; posterior margin of ? 
seventh abdominal sternum deeply emarginate (Text-fig. 427)) . annulipes (p 

Without black dots on venter of abdomen ; hemielytra tinged with black. (Back- 
ward spine-like processes of the posterior margin of the ¢ seventh abdominal 
tergum absent (Text-fig. 469) ; in the aedeagus, frontal conjunctiva with a bunch 


of spines (Text-fig. 472)) : : 3 : : : : . timoris (p. 


138) 


140) 


142) 


137) 


135) 


. 133) 


144) 


THE LEPTOCORISINAE OF THE WORLD 133 


Noliphus annulipes Walker 
(Text-figs. 421-429) 
Noliphus annulipes Walker, 1871 : 176 
Characterized by the light brown pronotum and propleuron with two smooth black 
dots (in linear order) anteriorly on either side of the venter of abdomen and by the 
form of the g and the 9 genitalia. 


6. Body length, 12-0-13-2 mm., width, 2:3-2:5 mm. Head length, 1-4-1-5 mm., width, 
I-6-1'7 mm. Antennae, segment I, 2:3-2:5 mm., II, 2:0-2:2 mm., III, 2-3-2-4 mm., IV, 
3°I-3:2 mm. Rostrum, segment I, 1-4-1-5 mm., II, 1-4-1-6 mm., III, 0-7-0-75 mm., IV, 
I-o-1:'2mm. Pronotum, length, 2-2-2-4 mm., width, 2-3-2:-5 mm. MHind femora, 5-6—6-0 mm. 
Hind tibia, 6-0-6:-4 mm. Distance between ocelli, 0-4—0-5 mm., distance between an ocellus 
and an eye, 0:85—0:9 mm. 

Body of medium size, light brown. Head, underside pale. Antennae pale, with apices of 
first, second and third segments black and apical segments brownish black, only pale at bases. 
Rostrum comparatively short, reaching anterior margin of third coxae. Propleuron pale, some- 
times tinged brown, usually with a brownish black line extending from sides of posterior margin 
of eyes on to anterior third of propleuron. Pronotal disc light brown, posteriorly with brownish 
black tinge ; spines on posterior angles thicker in middle, apices pointed. Legs usually pale, 
with apices of femora, bases of tibiae and tarsi brownish black. Hemielytra pale, with brownish 
black tinge and punctures. Abdomen, dorsum palish brown, third, fourth, fifth and sixth 
connexiva on either side with black spots, venter pale, sometimes tinged brown, with black 
triangular marks pointing posteriorly in middle of fifth and sixth segments, seventh medially 
black. Posterior margin of seventh abdominal tergum medially convex, laterally on either side 
with spine-like backward projections, concealing about half of pygophore ; pygophore with 
emarginate posterior margin (Text-fig. 421). 

6 GENITALIA: Claspers asymmetrically placed, (Text-fig. 421) ; both (dorsal and ventral) 
pairs of thecal appendages present ; two pairs of left lateral conjunctival appendages as in 
Text-fig. 425 ; two pairs of lateral conjunctival appendages, second right, bifid at apex as in 
Text-fig. 425 ; dorsal conjunctival appendage thicker at base and curved near apex (Text-fig. 
425) ; frontal conjunctiva usually with two or three spines. 

9. Body length, 12-8-13:6 mm., width, 2-5-2-7 mm. Head length, 1-4-1-6 mm., width, 
I-7-1°8 mm. Antennae, segment I, 2:2-2:6 mm., II, 2-0-2-:3 mm., III, 2-:2-2-5 mm., IV, 
3:0-3:3 mm. Rostrum, segment I, 1-5—1-6 mm., II, 1:56-1:65 mm., III, 0-75-0-85 mm., IV, 
I-2-1-3mm. Pronotum, length, 2:3-2-5 mm., width, 2-5-2:7 mm. Hind femora, 5:6—6-2 mm. 
Hind tibia, 6-0-6-6 mm. Distance between ocellim 0-5 mm., distance between an ocellus and 
an eye, 0-9 mm. 

Very similar to § but slightly larger, with external genitalia usually pale unicolourous ; 
posterior margin of seventh abdominal sternum greatly emarginate, with a short median split 
and a fold and backward spine-like projections on either side (Text-fig. 427). 

© GENITALIA: First gonocoxae large, seen from above externally broader than long, with 
irregular outer margin and round apices (Text-fig. 423) ; intervalvular sacs absent (Text-fig. 
429) ; spermatheca with a broad neck (Text-fig. 424). 


Type material. Holotype g, CELEBES : presented by Saunders (pygophore miss- 
ing), B.M.(N.H.). 


Other material. CELEBES: I 9; AMBOINA ISLANDS: 3g, I 9; AUSTRALIA : 
eo: 9 &- 


The range of this species extends in the west into Java and in the south east into 
the Northern Territory and Queensland (Australia). 


134 I. AHMAD 


This species is very close to discopterus Stal, from which it can easily be separated 
by the larger size of the body (12-0-13-6 mm.) (discopterus being smaller—8-4— 
10-5 mm.). 


Fics. 421-429. Noliphus annultpes, 421, 422, 425, 5, 423, 424, 426-429, 9. 421, term- 
inalia, dorsal view ; 422, claspers, inner and outer views ; 423, first gonocoxa, ventral 
view ; 424,spermatheca; 425, aedeagus (inflated), ventral view ; 426, first gonopophy- 
sis, ventral view; 427, terminalia, ventral view ; 428, ovipositor, ventral view ; 
429, same, dorsal view. 


THE LEPTOCORISINAE OF THE WORLD 135 


Noliphus discopterus Stal 
(Text-figs. 430-438) 
Noliphus discopterus Stal, 1873 : 87 


\a aeeey, 


“05 mm. _ 


Fics. 430-438. Noliphus discopterus, 430-432, 3, 433-438, 2. 430, terminalia, dorsal 
view ; 431, claspers, inner and outer views; 432, aedeagus (inflated), ventral view ; 
433, terminalia, ventral view ; 434, spermatheca; 435, ovipositor, ventral view ; 
436, same, dorsal view ; 437, first gonocoxa, ventral view ; 438, first gonopophysis, 
ventral view. 


136 I. AHMAD 


Characterized by small body ranging from 8-4-10-5 mm. ; by the short basal 
antennal segment, which is subequal to the second and by the form of the f and 2 
genitalia. 


3. Body length, 8-4-10-5 mm., width, 1-7-2-0 mm. Head length, 1-3-1-5 mm., width, 
I'5-1-6 mm. Antennae, segment I, 2-0-2:1 mm., II, 1-9-2-:0 mm., III, 2-0-2:2 mm., IV, 
2-7-2-°9 mm. Rostrum, segment I, 1-3-1-4 mm., II, 1-3-1-4 mm., III, 0-66-0-76 mm., IV, 
I-o-I1'2mm. Pronotum, length, 1-7—2-0 mm., width, 1-7-2-0mm. Hind femora, 4-8—5-2 mm. 
Hind tibia, 5:3-5-6 mm. Distance between ocelli, 0-4-0-5 mm., distance between an ocellus 
and an eye, 0-7—-0-8 mm. 

Body of small size, usually pale, tinged brownish black. Head behind ocelli brownish black, 
underside, except brownish black bucculae and rostrum, pale unicoloured. Antennae with 
apices of first, second and third segments brownish black, apical segments brownish black with 
small basal portions pale. Rostrum comparatively long, extending beyond third coxae. 
Propleuron pale, sometimes tinged with brown, with a black lateral line extending from posterior 
margin of eye on to posterior of propleuron. Pronotal disc pale, tinged with light brown with 
brownish black punctures, spines on posterior angles slanting posteriorly, acutely pointed. 
Legs pale, with apices of femora, tibiae and tarsi brownish black. Hemielytra pale, with dark 
brownish black costal margins, densely punctate. Abdomen, dorsum pale, with black marks 
on third, fourth, fifth and sixth connexiva, seventh black, venter yellowish brown, third, fourth 
and fifth with black marks in continuation with connexival marks on margins, round black dots 
on third and fourth venter present on either side, median portion of seventh, visible portion of 
eighth and pygophore brownish black. Posterior margin of seventh abdominal tergum sinuate 
(without backward spine-like projections), medially convex, pygophore curved near apex, 
posterior margin truncate (Text-fig. 430). 

6 GENITALIA : Claspers asymmetrically placed (Text-fig. 430) ; both (dorsal and ventral) 
pairs of thecal appendages present, two pairs of lateral conjunctival appendages as in Text-fig. 
432 ; second right bifurcated from the middle ; dorsal conjunctival appendage bow-shaped 
(Text-fig. 432) ; frontal conjunctiva usually with three spines, close together. 

2. Very similar to g in general appearance and measurements, seventh abdominal venter 
and genital structures pale ; posterior margin of seventh abdominal sternum greatly emarginate, 
with a large median split and a fold ; backward spine-like projections on either side present 
(Text-fig. 433). 

@ GENITALIA : First gonocoxae of medium size, externally as long as broad (Text-fig. 433) ; 
triangular in form and rounded at apices (Text-fig. 438) ; intervalvular sacs absent (Text-fig. 
436) ; spermatheca almost tubular, thicker in middle and thumb-like at apices (Text-fig. 434). 


Type material. Holotype g, SAMOAN IsLANDS : (Schmeliz), labelled “‘ discopte- 
vus’’, Riksmus. Stockholm. 


Other material. SAMOAN ISLANDS: 24,192; NEW HEBRIDES: 14,1 9. 


The range of this species extends in the west into New Hebrides and in the east 
into Samoa Islands. Adults have been found in November and December. 


Noliphus erythrocephalus Stal 
(Text-figs. 439-447) 
Noliphus evythrocephalus Stal, 1858 : 440, 1873 : 87 


Characterized by dark black pronotum, propleuron and underside of the head, by 
two black marks on the ventro-lateral of abdomen and by the form of the g and 9 
genitalia. 


THE LEPTOCORISINAE OF THE WORLD 137 


6. Body length, 12-4-12-8 mm., width, 2-3-2-5 mm. Head length, 1-5-1-6 mm., width, 
1-6-1'8 mm. Antennae, segment I, 2:5-2-6 mm., II, 2:2-2-4 mm., III, 2-4-2:5 mm., IV, 
3°8-4-0 mm. Rostrum, segment I, 1-4-1-6 mm., II, 1-6-1-75 mm., III, 0-7-0o-9 mm., IV, 
I-2-1:-3mm. Pronotum, length, 2:3—-2-5 mm., width, 2-3-2-5 mm. Hind femora, 5-6—6-:0 mm. 


Fics. 439-447. Noliphus erythrocephalus, 439-441, 3, 442-447, 2. 439, terminalia, dorsal 
view ; 440, claspers, inner and outer views ; 441, aedeagus (inflated), ventral view ; 
442, first gonocoxa, ventral view ; 443, spermatheca ; 444, terminalia, ventral view ; 
445, first gonopophysis, ventral view ; 446, ovipositor, ventral view ; 447, same, dorsal 
view. 


138 I. AHMAD 


Hind tibia, 6:4-6-7 mm. Distance between ocelli, 0-5 mm., distance between an ocellus and 
an eye, 0-9 mm. 

Body of medium size, brownish black. Head yellowish brown, post-occipital region black, 
underside including apex of clypeus, bucculae and rostrum shining black. Antennae with 
apices of first, second and third brownish black, apical segment brownish black, pale only at base. 
Rostrum long, usually extending beyond posterior margin of third coxae. Propleuron shiny 
black. Pronotal disc usually shining black, densely punctate. Spines on posterior angles, 
thicker and curved at bases, tapering smoothly towards apices, pointed. Legs brownish black, 
with bases of femora, tibiae and tarsi pale. Hemielytra palish brown, with costal margins and 
parts of corium dark black. Abdomen, dorsum yellowish brown, seventh black ; third to sixth 
connexiva with black marks continuous on sides of venter. Venter, palish brown with black 
triangular mark pointing posteriorly in middle of fifth and sixth segments, seventh, visible 
portions of eighth and pygophore dark black. Posterior margin of seventh abdominal tergum 
sinuate and medially convex, on either side with spine-like backward projections, concealing 
about three-quarters of pygophore ; pygophore tapering posteriorly with emarginate posterior 
margin (Text-fig. 439). 

3 GENITALIA: Claspers asymmetrically placed (Text-fig. 439) ; both dorsal and ventral 
pairs of thecal appendages present, two pairs of lateral appendages, second right bifurcated at 
apex, with distinct branches ; dorsal appendage usually “‘L’’-shaped ; frontal conjunctiva 
usually without spines (Text-fig. 441). 

2. Body length, 13-0-13-6 mm., width, 2-4-2:7 mm. Head length, 1:5-1-65 mm., width, 
1:8-1-9 mm. Antennae, segment I, 2:5-2:-6 mm., II, 2:3-2:4 mm., III, 2:5-2-6 mm., IV, 
missing. Rostrum, segment I, 1:-6-1-7 mm., II, 1-6-1-66 mm., III, 0-8-o-9 mm., IV, 1-3— 
1:35 mm. Pronotum, length, 2-4-2:6 mm., width, 2-4-2:7 mm. Hind femora, 6-0—-6-5 mm. 
Hind tibia, 6:9-7:2 mm. Distance between ocelli, 0-5 mm., distance between an ocellus and 
an eye, 0-9 mm. 

Very similar to g in general appearance but usually distinctly larger ; seventh venter of 
abdomen and genitalia externally black ; posterior margin of seventh abdominal sternum 
sinuately emarginate, with a short median split and a median fold ; spine-like backward projec- 
tions on either side present (Text-fig. 444). 

@ GENITALIA: First gonocoxae small, externally longer than broad, outer margin regular, 
apices pointed and inner margin irregular (Text-fig. 442) ; intervalvular sacs absent (Text-fig 
447) ; spermatheca with a hump-shaped neck (Text-fig. 443). 


Type material. Lectotype gj, AUSTRALIA: Queensland, Cape York (Jhorey), 
Riksmus. Stockholm ; allotype 9, data as holotype, Riksmus. Stockholm. 


Other material. AUSTRALIA: 54, 4 9. 


This species has restricted range in north east Australia (Queensland). 

This species is close to spinosus sp. n. but can be separated by the absence of 
ventro-lateral smooth black dots on anterior of abdominal venter. 

Six specimens (3 g, 39) were obtained from the Riksmuseum, Stockholm. None 
was clearly labelled as the type and in a personal communication Dr. Kjellander 
informed me that Stal often failed to fix a definite type-specimen. Therefore one 3 
specimen from Cape York (Queensland) has been selected as the lectotype and has 
been given a lectotype label. 


Noliphus insularis Stal 
(Text-figs. 448-450) 
Noliphus insularis Stal, 1865 : 185, 1873 : 87 


THE LEPTOCORISINAE OF THE WORLD 139 


Characterized by yellowish brown unicoloured abdominal venter, palish brown 
unicoloured minute ocelli, about a third of basal portion of apical antennal segment 
pale and by the form of the ¢ and 9 genitalia. 


6. Body length, 13:5-15:2 mm., width, 2-1-2-4 mm. Head length, 1-5-1-7 mm., width, 
1-7-1°85. Antennae, segment I, 3:1-3-6 mm., II, 2-7-3:1 mm., III, 2-8-3:2 mm., IV, 5:1 mm. 
Rostrum, segment I, 1-66-1-7 mm., II, 1-7-1-8 mm., III, 0-8-o-96 mm., IV, 1-3-1-4 mm. 
Pronotum, length, 2-:1-2:4 mm., width, 2:1-2-4 mm. Hind femora, 5:9-6-4 mm. Hind tibia, 
6-8-8:0 mm. Distance between ocelli, 0-4 mm., distance between an ocellus and an eye, 
0:86-0:96 mm. 

Body elongated and slim, tinged brown. Head pale, unicolourous, dorsal and ventral ; 
antennae brownish black, at least a third basal portion of apical segment pale. Rostrum long, 
extending well beyond third coxae. Propleuron including procoxae, mesopleuron and underside 
of collar brownish black. Pronotal disc brownish black with at least anterior quarter pale ; 
latero-posterior spines long, thin, curved near apices and acutely pointed. Legs brownish 
black, with at least basal half of femora pale. Entire hemielytra, except pale costal margin, 
brownish black, densely punctate. Abdomen, dorsum including connexiva pale, unicoloured ; 
abdomen ventrally pale unicolourous, including seventh venter and visible portion of eighth 
segment and pygophore. Posterior margin of seventh abdominal tergum truncated, slightly 
medially convex, with backward spine-like projection on either side, concealing two-thirds of 
pygophore ; pygophore curved and tapering at apex, posterior margin concave (Text-fig. 448). 

6 GENITALIA : Claspers symmetrically placed (Text-fig. 448) ; both (dorsal and ventral) 
pairs of thecal appendages present, two pairs of lateral conjunctival appendages (Text-fig. 450), 
second right partly sclerotized (Text-fig. 451) ; frontal conjunctiva with two sclerotized rounded 
pieces in middle of basiconjunctiva and a spiny appendage on right, top of first right lateral 
conjunctival appendage ; membranous appendage with a bow-shaped dorsal appendage, 
strongly curved in middle (Text-fig. 450). 

No 9 available for study. 


Fics. 448-450. Noliphus insularis, 3. 448, terminalia, dorsal view ; 449, 
claspers, inner and outer views ; 450, aedeagus (inflated), ventral view. 


140 I. AHMAD 


Type material. Holotype g, Ins. Fiy1 (Fidschi) (Jhorey), Riksmus. Stockholm. 
Other material. NEW HEBRIDES: 2. 


This species is close to papuensis Stal but can easily be separated by examining 
pale uncoloured abdomen. 


Noliphus papuensis Stal 
(Text-figs. 451-459) 
Noliphus papuensis Stal, 1865 : 185, 1873 : 87 


Characterized by its large size, by at least basal half of apical segments of antennae 
being pale, by the four black transverse lines on the underside of abdomen and by the 
structure of the g and 2 genitalia. 


3g. Body length, 13:5-15:2 mm., width, 2-3-2-6 mm. Head length, 1-66-1-8 mm., width, 
1-8-2-:0 mm. Antennae, segment I, 3:5-4-4 mm., II, 3:2-3:3 mm., III, 3:3-3-6 mm., IV, 
4°8-5-0 mm. (2:4-2-6 mm. pale). Rostrum, segment I, 1-7-2-0 mm., II, 1-7—-2:1 mm., III, 
o-g-1°0 mm., IV, 1-4-1-6 mm. Pronotum, length, 2-:3-2:7 mm., width, 2-3-2-6 mm. Hind 
femora, 6:5-8-0 mm. Hind tibia, 7-7-9:1 mm. Distance between ocelli, 0-5 mm., distance 
between an ocellus and an eye, 0-9 mm. 

Large and robust body, shining black. Head pale unicolourous. Antennae brownish black, 
with bases of first, second, third and at least basal half of apical segment pale. Rostrum long, 
usually extending well beyond third coxae. Propleuron shining brownish black. Pronotal 
disc shining brownish black, collar dorsally pale with small punctures, spines on posterior angles 
short, curved near base and smoothly tapering apically. Legs yellowish brown, with apices of 


0-5 mm. 


Fics. 451-453. Noliphus papuensis, 3. 451, terminalia, dorsal view ; 452, 
claspers, inner and outer views ; 453, aedeagus (inflated), ventral view. 


THE LEPTOCORISINAE OF THE WORLD 141 


femora, bases of tibiae, tarsi brownish black. Entire hemielytra brownish black, clavus and 
corium densely punctate. Abdomen, dorsum including connexival areas and pygophore 
brownish black ; abdomen venter, except two anterior brownish black segments, pale with 
four brownish black transverse lines, lateral margins brownish black, including visible portions 
of eighth and ninth (pygophore) segments. Posterior margin of seventh abdominal tergum 


Fics. 454-459. Noliphus papuensis, 2. 454, terminalia, ventral view ; 455, sperma- 
theca ; 456, ovipositor, ventral view ; 457, first gonocoxa, ventral view ; 458, first 
gonopophysis, ventral view ; 459, ovipositor, dorsal view. 


142 I. AHMAD 


slightly convex, with a backward spiny process on either side on lateral extremities, concealing 
about two-thirds of pygophore ; pygophore rounded, curved near apex, posterior margin 
slightly concave (Text-fig. 451). 

S$ GENITALIA : Claspers symmetrically placed (Text-fig. 451), dorsal and ventral thecal 
appendages present ; two pairs of lateral conjunctival appendages (Text-fig. 453) asymmetrical ; 
frontal conjunctiva with bunches of spines, usually about fifteen, second right lateral conjunc- 
tival appendage thicker at base and near apex, curved in middle and pointed at apex ; mem- 
branous appendage with an elongated irregular dorsal appendage (Text-fig. 453). 

2. Body length, 15-:2-16-0 mm., width, 2-67-2-7 mm. Head length, 1-7-1-9 mm., width, 
I-9-2:0 mm. Antennae, segment I, 3-6-3-9 mm., II, 2-:9-3-4 mm., III, 3-3-3-6 mm., IV, 
5-2-5:33 mm. Rostrum, segment I, 1-8-2:2 mm., II, 2-0-2:2 mm., III, 1-o-1:1 mm., IV, 
I-5-1-6mm. Pronotum, length, 2-6—2-7 mm., width, 2-6-2-7 mm. Hind femora, 7-5—7-6 mm. 
Hind tibia, 8-5-8-7 mm. Distance between ocelli, 0-5-0o-6 mm., distance between an ocellus 
and an eye, 0:85-0:95 mm. 

Very similar to g in general appearance but distinctly larger ; seventh abdominal sternum 
and genital structures usually pale, tinged brownish black. Posterior margin of seventh 
abdominal sternum emarginate, with a median split and a large median fold making two smaller 
lobes, one on each side at extremities, with a backward spiny process on each side (Text-fig. 
454)- 

Q GENITALIA: First gonocoxae large, seen from outside, broader than long, almost conical 
in form, with lobe-like apices and medially curved and concave external margin (Text-fig. 458). 
Intervalvular sacs absent (Text-fig. 459) ; spermatheca elongated, flat and hump-shaped near 
apex (Text-fig. 455). 


Type material. Holotype Jj, NEw GuINEA: “Stal”, “ papuensis’”’, Riksmus. 
Stockholm. 


Other material. NEW GUINEA: 21 ¢, 8 2; Misoor ISLANDS: I ¢; CERAM 
ISLANDS : I. 


This species is completely isolated in the genus and can be easily separated from 
other species by its entirely brownish black hemielytra, by brownish black dorsum 
of abdomen, black transverse lines on the ventral side of abdomen, long rostrum 
extending well beyond hind coxae and by pale colouration of basal half of apical 
segment of antennae. This species shows some relationship with Cosmoleptus 
limbaticollis (Stal) in having similar shape and colouration of pronotum and almost 
similar form of the apical antennal segment. 


Noliphus spinosus sp. n. 
(Text-figs. 460-468) 


Characterized by the black propleuron, pronotum and usally black underside of 
head and two round and smooth black dots in succession on either side of venter of 
abdomen, by the short basal segment of antennae, and by the form of g and 2 
genitalia. 


6. Body length, 12:5-13-2 mm., width, 2-3-2-4 mm. Head length, 1-4-1:5 mm., width, 
1-6-1'7 mm. Antennae, segment I, 2:3-2-5 mm., II, 2:2-2-4 mm., III, 2-4-2:6 mm., IV, 
missing. Rostrum, segment I, 1-4-1-7 mm., II, 1-6-1-7 mm., III, 0-75-0-8 mm., IV, 1-2— 
I:3mm. Pronotum, length, 2:3-2-4 mm., width, 2:3-2-4 mm. Hind femora, 6-0mm. Hind 
tibia, 7-2 mm. Distance between ocelli, 0-5-o-6 mm., distance between an ocellus and an eye, 
0:83-0°85 mm. 

Body of medium size, brownish black. Head usually yellowish brown, median post-occipital 


THE LEPTOCORISINAE OF THE WORLD 143 


region brownish black, underside of head, including apex of clypeus, bucculae and second, third 
and fourth segment of rostrum black, but underside of head sometimes yellowish brown. 
Antennae pale, with apices of first, second and third segments black, apical segment brownish 
black with a small basal portion pale. Rostrum usually long, extending slightly beyond third 


Fics. 460-468. Noliphus spinosus, 460, 461, 463, 5, 462, 464-468, 2. 460, terminalia, 
dorsal view ; 461, claspers, inner and outer views ; 462, first gonocoxa, ventral view ; 
463, aedeagus (inflated), ventral view ; 464, terminalia, ventral view ; 465, sperma- 
theca ; 466, first gonopophysis, ventral view ; 467, ovipositor, ventral view ; 468, 
same, dorsal view. 


144 I. AHMAD 


coxae. Propleuron shining brownish black. Pronotal disc usually shining brownish black, 
densely punctate posteriorly, spines on posterior angles comparatively short, thicker in middle 
and acutely pointed at apices. Legs usually brownish black, with basal half of femora entire, 
middle of tibiae and bases of tarsi pale. Hemielytra brownish black, with clavus and a portion 
of corium yellowish brown, with brownish black punctures. Abdomen, dorsum yellowish brown, 
seventh segment black, third, fourth, fifth and sixth connexiva with black spots ; abdomen 
venter yellowish brown, with black semicircular spots on fifth and sixth venter, visible portion 
of eighth and (pygophore) ninth usually black. Posterior margin of seventh abdominal tergum 
truncated, only slightly convex medially, concealing about three-quarters of pygophore, with 
very small backward spiny processes on either side ; pygophore round, tapering posteriorly 
with emarginate posterior margin (Text-fig. 460). 

6 GENITALIA : Claspers symmetrically placed (Text-fig. 460), both (dorsal and ventral) pairs 
of thecal appendages present ; two pairs of lateral conjunctival appendages, right thicker at 
base and bifid at apex, frontal conjunctiva usually with four spiny appendages (three on one 
side and one on the other) (Text-fig. 463) ; membranous appendage with a “‘ V ’’-shaped dorsal 
appendage, thicker at base and curved near base and apex (Text-fig. 463). 

2. Body length, 12-8-13-5 mm., width, 2-5-2-6 mm. Head length, 1-45-1-7 mm., width, 
1-7-1:°8 mm. Antennae, segment I, 2-4-2-6 mm., II, 2:2-2:5 mm., III, 2:4-2:7 mm., IV, 
3:6-3:7 mm. Rostrum, segment I, 1-5-1-7 mm., II, 1-6-1-:7 mm., III, 0-7-0-9 mm., IV, 
I-I-1-3 mm. Pronotum, length, 2:2-2-6 mm., width, 2-1-2-5 mm. Hind femora, 5-6—6:4 mm. 
Hind tibia, 6-7-7:2 mm. Distance between ocelli, 0-5—o-6 mm., distance between an ocellus 
and an eye, 0-8—o-9 mm. 

Very similar to g in general shape and colour but usually longer, seventh abdominal sternum 
and genital structures seen from outside usually brownish black, sometimes palish brown. 
Posterior margin of seventh abdominal sternum sinuately emarginate, with a very short median 
split and a short median fold, large lateral lobes on either side with a backward spiny process on 
either side (Text-fig. 464). 

Q GENITALIA: First gonocoxae large, seen from above broader than long, outer margin 
regular with lobe-like but truncated apices ; inner margin regular, convex in middle (Text-fig. 
462) ; intervalvular sacs absent (Text-fig. 468) ; spermatheca elongated, curved in middle and 
thicker at apex (Text-fig. 465). 


Type material. Holotype 3, PHILIPPINE ISLANDS: with a label “‘ Noliphus 
erythrocephalus Stal’’, Riksmus. Stockholm ; paratypes, PHILIPPINE ISLANDS : 
2 9 (Semper), Riksmus. Stockholm. 


Other material. PHILIPPINE ISLANDS: I 2; CELEBES: I g; KeEy ISLAND: 
Ig; SoLomMon ISLANDS: 19; JAvA: 16, 2 9. 


This species is very close to annulipes Walker and erythrocephalus Stal but can 
easily be separated from the former by the dark black pronotum and propleuron (in 
annulipes these are paler) and from the latter by the two pairs of black dots on the 
underside of the abdomen (in ervythrocephalus the black marks are continuous with the 
connexiva). 


Noliphus timoris sp. n. 
(Text-figs. 469-471) 
Characterized by pale, tinged brownish black pronotum and propleuron with antero- 


ventrolateral of abdomen without black marks or dots and by the form of ¢ and @ 
genitalia. 


THE LEPTOCORISINAE OF THE WORLD 145 


3. Body length, 13-3 mm., width,2-8mm. Head length, 1-6mm., width, 1-8 mm. Anten- 
nae, segment I, 2:7 mm., II, 2-4 mm., III and IV missing. Rostrum, segment I, 1-6 mm., II, 
1°7 mm., III, 0-8 mm., IV, 1:3 mm. Pronotum, length, 2-6 mm., width, 2-8 mm. Hind 
femora and tibiae missing. Distance between ocelli, 0-5 mm., distance between an ocellus and 
an eye, 0-9 mm. 

Body of medium size, brownish black. Head yellowish brown, tinged black, underside 
yellowish brown, only clypeus, labrum and rostral segments brownish black. Antennae pale, 
tinged brownish black. Rostrum long, extending slightly beyond third coxae. Propleuron 
pale, tinged brown, a black line extending from collar on to anterior third of propleuron. Prono- 
tal disc pale, tinged brown, with anterior part behind collar and posterior part from latero- 
posterior spines brownish black ; spines on posterior angles long and sharply pointed at apices. 
Legs pale, with apices of femora, apices of tibiae and bases of tibiae and tarsi brownish black. 
Entire hemielytra brownish black, densely punctate. Scutellum brownish black. Abdomen, 
dorsum pale, tinged brown, third, fourth, fifth and sixth connexiva with brownish black marks ; 
seventh abdominal segment brownish black, abdomen veatrally pale, tinged brown, fifth and 
sixth venter with median black triangular marks pointing posteriorly, seventh venter, visible 
portion of eighth segment and pygophore brownish black ; no black dot or mark on ventro- 
lateral margins of third and fourth abdominal venter. Posterior margin of seventh abdominal 
tergum sinuately truncated, without a spine-like backward process on either side of lateral 
extremities, concealing about two-thirds of pygophore ; pygophore rounded, curved and 
tapering at apex, posterior margin slightly concave. (Text-fig. 469). 

6 GENITALIA : Claspers symmetrically placed (Text-fig. 469) ; both (dorsal and ventral) 
thecal appendages present ; two pairs of lateral conjunctival paired appendages, second right 
curved and bifid at apex (Text-fig. 471) ; frontal conjunctiva with a bunch of spines (about 11 
usually) at base of first right conjunctival appendage ; aedeagus with ‘“‘L”’ shaped dorsal 
appendage (Text-fig. 471). 

No @ available for study. 


Holotype g, Timor : Riksmus. Stockholm. 


0-5 mm. 


Fics. 469-471. Noliphus timoris, §. 469, terminalia, dorsal view ; 470, 
claspers, inner and outer views ; 471, aedeagus (inflated), ventral view. 


146 I. AHMAD 


This species can be separated from all the species of this genus except imsularis 
by absence of black dots or marks on ventro-lateral of third and fourth abdominal 
segments. From imsularis Stal it can be separated by the presence of black tri- 
angular marks on the underside of the abdomen in the middle of fifth and sixth seg- 
ments. 


ZOOGEOGRAPHY AND PHYLOGENY 


The subfamily Leptocorisinae is represented in all zoogeographical regions, but 
in the Nearctic and Palearctic it is only found in the extreme southern areas and it is 
not recorded from New Zealand. 

An exact phylogeny of this group cannot be attempted, for there is no fossil 
material, little knowledge of the biology of many members of the group and none of 
the biochemistry. Further more the phylogeny and even the classification of the 
family Alydidae, indeed of the whole Coreoidea is uncertain (Stys, 1961 ; Scudder, 
1963). However, a study of the zoogeography and the comparative morphology 
of the group throw at least some light on the relationships and a very tentative phylo- 
genetic scheme is shown on Text-fig. 472. 


ORYZOCORIS 


Median groove of 7th abd. 
sternum with deep split REN GCORS 


car aoe BLOETEOCORIS MUTUSCA 


Median groove developing in pro- 
jection on 7th abd. sternum; 
paired dorsal thecal appendages 
very reduced or absent 


ERBULA 
Inflexed genae. Basal atennal segment swollen; 
dorso-lateral conj dorso-lateral conj. app. mem- 
app. sclerotised braneous, 
NOLIPHUS 


Development of projection on median 
of 7th abdominal sternum. Loss of 
intervalvular sacs, reduction of dorsal 
thecal appendage 


Loss of intervalvular sacs, 
development of spines on 
LEPTOCORISA posterior of pronotum COSMOLEPTUS 


PSEUDOLEPTOCORISA 


Spine on 
Loss of intervalvular sacs, develop- front femora 


ment of dorso-lateral conjunctival 
STENOCORIS S. Lat. appendages 


Development of 
spermathecal flange. 


Loss of supporting 
conjunctival appendage 


LYRNESSUS 


LEPTOCORISINI NOLIPHINI 
Paraclypeae Clypeus 
clongaced clongated 


ee 


HYPOTHETICAL ANCESTOR 
Normal head, fore femora unarmed, both dorsal and ventral checal 
appendages (in pair), supporting conjunctival appendages present; 
absence of frontal and dorso-lateral conjunctival appendages in the 
male aedeagus, Intervalvular sacs in the female ovipositor, emar- 
ginate posterior margin of the seventh abdominal sternum and 
spermatheca without a median flange 


Fic. 472. Phylogenetic tree of the Leptocorisinae. 


THE LEPTOCORISINAE OF THE WORLD 147 


----=Boundary line 
of species 


Leptocorisa lepida 
luzo nica 
pseudo lepida 


Fic. 473. Distribution map of Leptocorisa spp. 


The two tribes Leptocorisini and Noliphini are clearly distinct on the form of 
head and pronotum, whereas in the Noliphini there are distinctions in the shape and 
form of the body in the different genera, the body form in the Leptocorisini is 
remarkably uniform and the taxonomic distinctions are mainly based on characters 
of the genitalia. It may be that evolutionary pressures due to the habitat (various 
Gramineae) of the Leptocorisini has restricted the modification of the form of the 
body ; the other large groups of Heteroptera that share this habitat, Stenodemini 
(Miridae) and Blissinae (Lygaeidae), are also both relatively uniform in body form. 
The habits of the Noliphini are unknown. Alternatively it may be that the 
Leptocorisini are more recently evolved and so less differentiated than the Noliphini. 

The Noliphini have an interesting distribution, similar to, but rather more exten- 
sive than that of the Marsupials : Neotropical region (Lyrnessus) and Australian 
region and Malayan and Philippine sub-regions of the Oriental region (Cosmoleptus 
and Noliphus). The centre of evolution of the tribe was clearly in the Australian 
region, perhaps in the Papuan sub-region ; the farthest west that the tribe has 
spread is north east Sumatra (C. swmatranus) but it does not seem to have crossed 
the Strait of Malacca or the South China Sea. C. bakeri is found in the Philippine 
Islands. Noliphus spinosus has the widest distribution of any member of that 
genus ; Java, Phillippine Isands, Solomon Islands and north Australia, but the 
remaining species are, so far as present records show, restricted to the Australian 
region (i.e. east of Wallace’s Line). 

In contrast the Leptocorisini are almost world-wide and some of the species have 
very wide ranges, e.g. Leptocorisa acuta, the ‘‘ Gandhi rice bug ”’, is found throughout 
the Oriental and Australian regions (except New Zealand). It is considered that 
Stenocoris (Pseudoleptocorisa) erratica retains more primitive characters than any 


148 I. AHMAD 


other species of the tribe and this, together with zoogeographical evidence, leads to 
the tentative suggestion that in contrast to Noliphini, the centre of evolution of this 
tribe was in the Ethiopian-Oriental regions. The subgenera of Stenocoris show a 
series of changes in the form of the posterior margin of the 9 seventh abdominal sternum 
running parallel with which is the reduction of the dorsal thecal appendages in the 
aedeagus. The dorsal thecal appendages are paired in Pseudoleptocorisa, they are 
fused in Evbula and very small and reduced in Stenocoris claviformis and absent in 
all the other species of Stenocoris and in all the species of Orvyzocoris. The subgenus 
Eybula is entirely Ethiopian, Stenocoris is distributed in Ethiopian, southern Nearctic 
and Neotropical regions and Ovyzocorts is limited to the southern Nearctic and Neo- 
tropical (occurring as far north as Florida and Texas). The Neotropical species of 
Stenocoris stand apart from the rest of the subgenus, more especially in the broad 
bases of the first gonocoxae that extend to conceal most of the eighth paratergite 
proximally. 


Stenocoris pallida. 
v maculosa 


“ll 


474 


Fic. 474. Distribution map of Stenocoris spp. 


THE LEPTOCORISINAE OF THE WORLD 149 


Sao Tome 


fe. = Stenocoris 
southwoodi 


also present on- 
Principé 
Sao Tome 
Zanzibar 


Fic. 475. Distribution map of Stenocoris (Erbula) southwoodt. 


The remaining genera of the Leptocorisini, Leptocorisa, Bloeteocoris and Mutusca, 
have Oriental and Australian distributions but Leptocorisa is closer to Stenocoris than 
are the other two. Mutusca and Bloeteocoris might be considered more specialized 
in having lost the ancestral intervalvular sacs in the ovipositor and developed dorso- 
lateral conjunctival appendages in the aedeagus. In Mutusca one pair of these 
appendages are present and are membranous in form ; running parallel with this 
character is the entirely swollen basal antennal segment, whereas in Bloeteocoris there 
are two sclerotized pairs of dorso-lateral conjunctival appendages in the aedeagus 
and this genus also shows remarkable development of genae, the tips of which are 
acutely pointed and inflexed. 

The species of Leptocorisa fall into a number of groups : 

L. costalis (range Indo-China, Philippines and East Indies) is in many ways the 
more generalized, showing more characters in common with Stenocoris erratica, 
especially the small membranous appendage in the male aedeagus, whilst L. 
biguttata (Indo-China, Sarawak, Celebes and East Indies) and L. palawanensis 


150 I. AHMAD 


(Philippine Islands and New Guinea) occupy a central position in the genus, around 
which are the four main species groups: 

1. luzonensts (restricted to Philippine Islands) and tagalica extending from the 
Philippine Islands to Sarawak, Halmahera Island, with two black dots behind the 
ocelli and brownish black tips to hind femora. 

2. sakdapolrakae (Thailand, Java, and Sumatra), discoidalis (Thailand, New 
Guinea, Australia, Solomon Islands and New Hebrides) and solomonensis (New 
Guinea and Solomon Islands) with a large rostrum extending beyond second coxae 
and the black line on the side of the head and prothorax lengthening in the order 
given above (which also, perhaps coincidently, corresponds with successive pro- 
gression of the ranges towards east). 

3. lepida (North and Central India, Bhutan, Burma and Thailand), pseudolepida 
(South India, Ceylon and Malaya) and /uzonica (Philippine Islands, Sarawak and 
Vietnam) (Text-fig. 473) which are smaller than the other species and possess the 
second right lateral conjunctival appendage, which is short in lepida, elongated in 
pseudolepida and elongated and curved in luzonica ; these appear to form a group 
of allopatric species, the range of lepida being north of that of pseudolepida and of 
luzonica north east of that of lepida. 

4. chinensis, acuta and oratorius, the most widely distributed species of all in 
the Oriental and Australian regions and major rice pests, show progressive reduction 
of the lateral black line (extending from antenniferous tubercles up to the lateral 
side of collar, sometimes like two dots in chinensis, represented by one dot at collar 
in acuta and completely absent in ovatorius) and parallel with this a trend for the apex 
of the claspers to become united, for whereas they are truncated in chinensis, one 
lobe is produced in acuta and finally in ovatorius, this lobe is enlarged and acutely 
pointed and the other lobe is lost. 


ACKNOWLEDGMENTS 


It is indeed a great pleasure for me to thank the following : Professor O. W. 
Richards who granted me facilities in his department, Dr. T. R. E. Southwood, my 
supervisor who suggested the problem, for his constant encouragement and valuable 
criticisms. Dr. W. E. China whose suggestions, criticisms and comments helped 
greatly. Iam glad to acknowledge my indebtness to other members of the Imperial 
College Field Station and the British Museum (N.H.), namely Messrs. R. G. Davies, 
M. Luff, J. W. Siddorn, W. O. Steel and Dr. M. S. K. Ghauri, Messrs. R. J. Izzard 
and J. A. Grant respectively, for kind help in various ways. 

I am also grateful to the authorities of the following museums and to the members 
of their staff mentioned for the generous loan of or access to specimens. 

I. British Museum (Natural History), London. 

2. Hope Collections, Oxford University Museum (Professor Varley, Mr. I. 
Lansbury). 

3. Australian Museum, Sydney, Australia (Dr. J. W. Evans). 

4. South Australian Museum, Adelaide, Australia (Dr. G. F. Gross). 

5. Carnegie Museum, Pennsylvania, U.S.A. (Dr. G. Wallace). 


THE LEPTOCORISINAE OF THE WORLD I51 


6. American Museum of Natural History, New York (Dr. P. Wygodzinsky). 
7. Bernice P. Bishop Museum, Honolulu, Hawaii (Dr. J. L. Gressitt and Miss S. 
Nakata). 
8. Museum do Dundo, Dundo, Lunda, Angola (Dr. A. de Barros Machado). 
9. Museum National d’Histoire naturelle, Paris, France (Dr. A. Villiers). 
o. Musée Royale de l'Afrique centrale, Tervuren, Belgium (Dr. P. Basilewsky). 
Ir. Riksmuseum, Stockholm (Dr. E. Kjellander). 
12. Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands (Dr. H. C. 
Blote). 
13. Zoological Institute, Helsinki (Dr. Hackman, Professor Lindberg, Dr. P. 
Naurteva and M. Meinander). 
14. Deutsches Entomologisches Institut, Berlin (Professor Dr. Hans Sachtleben). 
15. Zoologisches Institut Freie Universitat, Konigin Luise-Strasse 13, Berlin 
(Professor Dr. K. K. Gunther). 

16. National Museum, Prague, Czechoslovakia (Dr. L. Hoberlandt). 

17. Universitetets zoologiske Museum, Copenhagen (Dr. S. L. Tuxen). 

18. Zoological Institute, Uppsala University Museum (Dr. Kullenberg, L. Hed- 

strom, Professor Holms). 

19. Moravske Museum Biologicky ustav, Brno, Czechoslovakia (Dr. J. Stehlik). 

20. Zoological Institute, Academy of Sciences, Leningrad, V.164, U.S.S.R. (Dr. I. 

M. Kerzhner). 

The following have kindly sent me information about certain specimens : 

Dr. W. Forster, Zoologische Staatsammlungen, Miinchen, Germany. 

Dr. Livio Tamanini, Rovereto, Italy. 

Professor Dr. M. Beier, Naturhistorisches Museum, Vienna. 

Professor Dr. Woodward (Dept. of Entomology, University of Queensland, 
Brisbane, Australia), Dr. G. Rothschild (Agricultural Institute, Sarawak, Borneo), 
Miss N. Sakdapolrak (Department of Entomology and Parasitology, University of 
California) and Mr. J. C. Schaffner (Department of Zoology and Entomology, Iowa 
State University, Ames, Iowa) generously made their personal collections available 
to me. 

The University of London Central Research Fund made a grant to cover the cost 
of my travelling expenses to examine collections in Uppsala, Stockholm and Helsinki. 


REFERENCES 


AuMAD,I. 1964. Systematic position of Dicrorymbus, Xenocervaea and Procamptus (Hemiptera, 
Coreidae), three of Bergroth’s genera from the Philippine Islands, with descriptions of two 
new species. Amn. ent. fenn. 30 : 17-34. 

—— Erbula Stal, 1873 (Class Insecta, Order Hemiptera) : Proposed designation under the 
Plenary Powers of Stenocoris (Evbula) southwoodi Ahmad, 1965 as type. Bull. zool. Nom. 
(S.). (In press). 

Auman, I. & SoutHwoop, T. R. E. 1964. The Morphology of the Alydid abdomen with special 
reference to the genitalia and its bearing on classification. Tzjdsch. Ent. 107 : 365-378. 

AKBAR,S.S. 1958. The morphology and life history of Leptocorisa varicornis Fabr. (Coreidae, 
Hemiptera)—A Pest of Paddy Crop in India. Publ. Aligarh Univ. (Zool. Ser.) Ind. Ins. 


Typ. 5(2) : 1-50, 7 pls. 


152 I. AHMAD 


Amyot, C. J. B. & AUDINET-SERVILLE, J. G. 1843. Histoive naturelle des Insectes. Hémi- 
pteres. Ixxv + 675 + 6pp.- Paris. 

BERGROTH, E. 1913. Supplementum Catalogi Heteropterorum Bruxellensis, I]. Mém. Soc. 

ent. Belg. 22 : 125-183. 

1913. Bibliographisches tiber Hemipteren. Ent. Mitt. 2 : 10-12. 

1918. Studies in Philippine Heteroptera. I. Philipp. J. Sci. D, 13 : 43-126. 

1925. Note on some Coreidae (Hem. Het.). Konowia 4 : 82-88. 

BERTHOLD, A. A. 1827. Latyreille’s. . . Natiivlich Familien des Thierveichs mit Anmerkungen 
und Zusdtzen von A. A. Berthold. 602 pp. Weimar. 

BLaTcHLEY, W.S. 1926. Heteroptera or true bugs of Eastern North America. With especial 
vefevence to the Faunas of Indiana and Florida. 1116 pp. Indianopolis. 

Biore, H.C. 1934. Catalogue of the Coreidae in the Rijksmuseum van Natuurlijke Historie, 

Part I. Corizinae, Alydinae. Zodél. Meded., 17 : 253-285. 

1937. On African species of Leptocovixa Berthold. Temminckia 2 : 281-296. 

BREDDIN,G. 1909. Khynchoten von Ceylon gesammelt von Dr. Walter Horn. Avan. Soc. ent. 
Belg. 53 : 250-309. 

1912. Neue indo-malayische Hemiptera. Avch. Naturgesch. 78A : 95-107. 

Buenion, E. (with N. Poporr). t1g1o. L’appareil salivaire des Hémiptéres. II, 4. Arch. 
Anat. micr. 11 : 435-456. 

BuRMEISTER, C. H.C. 1834. Rhyngota seu Hemiptera. Nova Acta Leop. Carol. (Suppl. 1) 

16 : 235-306. 

1835-39. Handbuch dev Entomologie 2 : 1-400. Berlin. 

CASTELNAU, F. L. DE L. DE. 1832. Essai d’une classification systématique de l’ordre des 
Hémipteéres. Mag. Zool. 2. 88 pp., pls. 51-55. 

Cuina, W. E. 1924. <A preliminary revision of the Oriental species of Leptocorisa Latr. 

(Hemiptera, Coreidae). Bull. ent. Res. 14 : 235-2309. 

1930. Insects of Samoa and other Samoan Terrestrial Arthropoda. 2(3) : 81-162. London. 

1934. Anew species of Coreidae (Heteroptera) injurious to coconut in the Solomon Islands. 
Bull. ent. Res. 25 : 187-189. 

Cuina, W. E. & MILLER, N.C. E. 1959. Check list and keys to the families and subfamilies 
of Hemiptera-Heteroptera. Bull. Brit. Mus. (nat. Hist.) Ent. 8 : 1-45. 

Cuina, W. E. & Aumap, I. 1963. Leptocorixa Berthold, 1827 (Insecta, Hemiptera) : Pro- 
posed suppression under the Plenary Powers in favour of Leptocovisa Latreille, 1829. 
Bull. zool. Nom. 20 : 435-437. 

Costa, A. 1863. Illustrazione di alcuni Emitteri stranieri all’Europa. Nota seconda nuovi 
generei e nuovi specie di Coeridei. R.C. Accad. Napoli 2 : 250-261. 

Datras, W. S. 1852. List of the specimens of Hemipterous insects in the collection of the 
British Museum 3 : 369-352. London. 

DEGEER, C. 1773. Mémoires pour sevviy a Vhistoive des Insectes. 3. vili + 697. Stockholm. 

in GorzE, J. A. E. 1778. Entomologische Beytrage zu des Ritter Linné zwolften Ausgabe 

des Natursystems. 2. \xxii + 352 pp. Leipzig. 

Distant, W. L. 1880-1893. Biologia Centyvah-Americana, Zoology, Rhyncota Heteroptera, 1. 

xx + 642 pp. London. 

tgo1. Rhynchotal notes. IX. Heteroptera: Fam. Coreidae. Ann. Mag. nat. Hist. 

(7)7 : 416-432. 

1901. Revision of the Rhynchota belonging to the family Coreidae in the Hope Collection 

at Oxford. Pyvoc. zool. Soc. 1 : 325-335. 

1902. The Fauna of British India including Ceylon and Burma. WRhynchota. 1. xxxvili 

+ 438 pp. London. 

Dovuetas, J. W. & Scott, J. 1865. The British Hemiptera. 1, Heteroptera. xii + 628 pp. 
London. 

Fapsricius, J. C. 1775. Systema Entomologiae, sistens Insectorum classes, ovdines, genera, 
species, &c. xxxii + 832 pp. Flensburgi & Lipsiae. 


THE LEPTOCORISINAE OF THE WORLD : 153 


1781. Species Insectorum exhibentes eorum differentias specificas . . . adjectis observationibus, 
descriptionibus. 2: 495-517. Hamburgi & Kilonii. 

1787. Mantissa Insectorum, sistens eorum species nuper detectas : adjectis chavacteribus 
genericis differentiis specificis emendationibus, observationibus. 2. 1-382. Hafniae. 


1794. Entomologia systematica emendata et aucta ... adjectis synonymis, locis, observa- 
tionibus, descriptionibus. 4. viii + 477 pp. Hafniae. 
—— 1803. Systema Rhyngotorum ... adjectis synonymis, locis, observationibus, descriptionibus. 


X, 21, 314 pp. Brunsvigae. 

GERSTAECKER, C. E. A. 1873. Insecta, Arachnida, Myriapoda and Isopoda zm Baron von der 
Decken, Reisen in Ost Afrika in 1859-61. 3(2). xvi-+ 542 pp., 18 pls. Leipzig. 

GMELIN, J. F. 1789 (1790). in LINNAEUS, C.... Systema Naturae, Ed. XII, Insecta II, 1(4) : 
2041-2224. Lipsiae. 

GUERIN-MENEVILLE, F. E. 1830. Voyage autour du Monde ... sur la Coquille, pendant ... 

1822-25. Crustacea, Arachnida, Insecta. Zoologie 2(2) : 9-302. Paris. 

1857. Animaux articulés a pieds articulés. In Sacra, R. DE LA Histoire physique, 
politique et naturelle de I'fie de Cuba. Il. |xxxvii + 868 pp. Paris. 

HERRICH-SCHAFFER, G. A. W. 1848. In Haun, C. W. Die Wanzenartigen Insecten. Getreu 
dey Natur abgebildet und beschreiben. Fortsetzung. 8 : 1-124. Niirnberg. 

Horn, W. 1926-28. Ueber den Verbleib der entomologischen Sammlungen der Welt. Supp. 
ent. Berl. 12 : 1-133 ; 17 : 72-120. 

Hsiao, T. Y. 1963. Results of the Zoologico-Botanical Expedition to Southwest China, 
1955-57. Acta ent. sinica 12 : 332-344. 

KirRKALDY, G. W. t1g00. On the nomenclature of the genera of the Rhynchota Heteroptera and 

Auchenorrhynchous Homoptera. Ent. mon. Mag. 33 : 25-28, 262-265. 

1902. Memoirs on Oriental Rhynchota. |. Bombay nat. Hist. Soc. 14 : 46-58, 262-265. 

1908. A catalogue of the Hemiptera of Fiji. Proc. Linn. Soc. N. S. W. 33(2) : 345-391. 

1908. Bibliographical and nomenclatorial notes on the Hemiptera. No. 8. Ent. mon. 

Mag. 41 : 123-124. 

Kovenati, F. A. 1845. Meletemata Entomologica. 2 : 1—132. Petropoli. 

KONINGSBERGER, J.C. 1903. Ziekten van Rijst, Tabak, Thee en andere cultuurgewassen die 
door insectend worden veroorzakt. Meded. PlTuin, Batavia. pp. 6-10. 

~ LATREILLE, P. A. 1807. Geneva Crustaceorum et Insectorum secundum orvdinem natuvalem in 

familias disposita, &c.3. 126 pp. Parisiis & Argentorati. 

1810. Considérations générales sur l’ordve naturel des animaux composant les classes des 

Crustacés, des Avachnides et des Insectes. 444 pp. Paris. 

1825. Familles naturelles du Régne Animal, exposées succinctement et dansun ordre analytique 

avex, Vindication de leurs genres, &c. 570 pp. Paris. 

1829. in Cuvier, G.L.C.D.D. Le Régne Animal distribué d’aprés son organisation. 

5. xxiv -+ 556 pp. Paris. 

Leacu, W. E. 1815. Hemiptera. In Brewster, D. The Edinburgh Encyclopaedia 9 : 120- 
124. Edinburgh. 

Lerroy, H.M. 1908. The Rice bug (Leptocorisa varicornis Fabr.) Mem. Dept. Agric. India Ent. 
2 : I-T4. 

Leston, D. 1958. Chromosome number and the systematics of Pentatomorphs (Hemiptera). 
X Int. Congr. ent. Proc. 2 : 911-918. 

Leston, D. & ScuppER, G. C. E. 1956. A key to the larvae of the families of the British 
Hemiptera-Heteroptera. Ent. mon. Mag. 89 : 223-231. 

LeETHIERRY, L. F. & SEVERIN, G. 1894. Catalogue général des Hémiptéves 2. 277 pp. Brux- 
elles. 

Mitter, N.C. E. 1956. The biology of the Heteroptera. x + 162 pp. London. 

MONTROUSIER, P. & OTHERS. 1865. Essai sur la faune entomologique de Kanala (Nouvelle 
Calédonie) et déscription de quelques espéces nouvelles ou peu connues. Ann. Soc. Linn. 
Lyon. 11: 46-257. 


154 I. AHMAD 


OxiviER, G. A. 1811. In Encyclopédie Méthodique. Histoive Naturelle, Insectes. 8. 722 pp. 
Paris. 

OsHANIN, B. 1912. Katalog der paldarktischen Hemipteren (Heteroptera, Homoptera-A uchenor- 
vhyncha und Psylloidea). xvi-+ 187 pp. Berlin. 

RevuTER, O. M. 1910. Neue Beitrage zur Phylogenie und Systematik der Miriden, nebst 
einleitenden Bemerkungen iiber die Phylogenie der Heteropteren-Familien. Acta Soc. Sci. 
Fenn. 37(3) : 1-167. 

1912. Bemerkungen uber mein neues Heteropterensystem. Ofvers. finska vetensk. Soc. 

Foérh. 54(6) : 1-62. 

1913. Ausfihrliche Beschreibungen einiger palaarktischen Hemipteren. Ibid. 55(14) : 
I-I1l. 

SCUDDER, G. G. E. 1963. Adult abdominal characters in the Lygaeidae-Coreoid complex of 
the Heteroptera, and the classification of the group. Canad. J]. Zool. 52 : 1-14. 

SIGNORET, V. 1860. Faune des Hémipteres de Madagascar. 2° partie, Hétéropteéres, I. 
Ann. Soc. ent. Fr. (3)8 : 917-972. 

SouTHWooD, T. R. E. 1955. The morphology of the salivary glands of terrestrial Heteroptera 

(Geocorisae) and its bearing on classification. Tijdsch. Ent, 98 : 77-84. 

1956. The structure of the eggs of the terrestrial Heteroptera and its relationship to the 

classification of the group. Tvans. R. ent. Soc. Lond. 108 : 163-221. 

SouTHWoOOD, T. R. E.& Leston, D. 1959. Land and water bugs ofthe British Isles. x + 436 


pp. London. 
STAL, C. 1858 (1859). Hemipterologiska bidrag. Ofvers. VetenskAkad. Férh. Stockh. 1858. 
10 : 433-454. 


1859. Till kannedomen om Coreida. Jbid. 10 : 449-475. 

1859. Hemiptera species novas descriptit. In Fvregatten Eugenies Resa. Ins. Hemipt. 
7 : 219—208. 

1860—1862. Bidrag till Rio Janeiro-traktens Hemipter-Fauna, &c. K. svenska VetensA kad. 
Handl. 2(7), 184 pp., 3(6), 75 pp. 

—— 1865. Hemiptera nova vel minus cognita. Amn. Soc. ent. Fr. (4)5 : 163-188. 

— 1865. Hemiptera Africana, 2: 1-181. Holmiae. 

—— 1866. Analecta Hemipterologica. Berl. ent. Z. 10 : 151-172. 

—— 1868. Bidrag till Hemipternas systematik. Ofvers. VetensAkad. Férh. Stockh. 7 : 491— 
560. 

1870. Enumeratio Hemipterorum (1). Bidrag till en férteckning 6fver alla hittills 
kanda Hemiptera jemte systematiska meddelanden, &c. K. svenska VetenskAkad. Handl. 


OF 232° 
1872(1873). Genera coreidarum Europae disposuit. Ofvers. VetensAkad. Févh. Stockh. 
1872, 6 : 49-58. 


1873. Enumeratio Hemipterorum. Jbid. (3) 2 : 1-63. 

STICHEL, W. 1925. Iilustrierte Bestimmungstabellen dey Deutschen Wanzen. 499 pp. Berlin. 

Stys, P. 1961. Morphology of the abdomen and female ectodermal genitalia of the trichopho- 
rous Heteroptera and bearing on their classification. XI Int. Congr. Ent. Proc. 1 : 37-43. 

THUNBERG, C. P. 1783. Dissertatio Entomologica novas Insectorum species sistens. Uppsala. 

TILLYARD, R. J. 1926. The Insects of Australia and New Zealand. 560 pp. Sydney. 

UHLER, P.R. 1886. Check list of the Hemiptera Heteroptera of North America. 34pp. Brook- 
lyn. 

UicHanxko, L. 1921. The rice bug Leptocovisa acuta Thunberg in the Philippines. Philipp. 
agric. Rev. 14 : 87-125. 

Van Duzee, E. P. 1914. Nomenclatural and critical notes on Hemiptera. Canad. Ent. 
46 : 377-3809. 

VILLIERS, A. 1955. Les Leptocoriva (Hem. Coreidae) de la collection de VIFAN. Bull. Inst. 
franc. Afr. Noive, 17 : 1141-1144. 

1963. Les Leptocovixa africaines et malgaches du Muséum de Paris (Hem. Coreidae). 

Ibid. 25 : 229-234. 


THE LEPTOCORISINAE OF THE WORLD 155 


WAGNER, E. 1961. Heteroptera-Hemiptera. Tierwelt Mutteleuvopas, 4(3), 10a : 1-174. 
Leipsig. 

WALKER, F. 1871. Catalogue of the specimens of Heteropterous Hemiptera in the collection of 
the British Museum, 4. 211 pp. London. 

Westwoop, J. O. 1842. A catalogue of Hemiptera in the collection of F. W. Hope, with latin 
descriptions of the new species. 2. 26 pp. London. 

ZEHNTNER, D. De levensgeschiedenis van dan walang sangit. De indische Natuur 1: 77 
(cited by UIcHANCO, 1921), original not seen. 


INDEX TO GENERA AND SPECIES 


(Synonyms in italics) 


acuta Thunberg, 78 
acuta Distant, 94 
africana sp. n., 32 
americana sp. n., 34 
angustata, 78 
annulicornis Blote, 49 
annulicornis Signoret, 16 
annulipes, 133 
apicalis Stal, 24 
apicalis Westwood, 36 
arcuata, 94 


bakeri sp. n., 126 
bengalensis, 94 
biguttata, 80 
bipunctata, 109 


BLOETEOCORIS gen. n., 115 


blotei, 49 

braziliensis sp. n., 38 
brevicornis, III 
burmeisteri, 109 


chinensis, 82 
claviformis sp. n., 41 
corbetti, 82 
CORISTENIA, 123 
COSMOLEPTUS, 126 
costalis, 85 

crudelis, 55 


DICRORYMBUS, 4 
discoidalis, 87 
discopterus, 135 
distinguenda, 18 
elegans, 20 
ERBULA, 14 
erratica, 12 


erythrocephalus, 137 
ethiopis sp. n., 43 


fabricii, sp. n., 62 
filiformis, 64 
flavicosta, 123 
flavida, 78 
furcifera, 67 


geniculata China, 106 
geniculata Guérin, 123 
GRY POCEPHALUS, 115 


inflexigena, 117 
insularis, 138 


lepida, 88 

LEPTOCORISA Laporte, 60 
LEPTOCORISA Latr., 69 
LEPTOCORIXA, 69 
limbaticollis, 128 

linearis, 64 

luzonensis sp. n., 90 
luzonica sp. n., 92 
LYRNESSUS, 123 


maculiventris, 94 
maculosa, 44 
meridianus sp. n., 119 
minutus sp. n., 121 
MUTUSCA, t110 
MYODOCHA, 9 


nigvicornis, 68 
nitidula, 82 
NOLIPHUS, 131 


156 INDEX 


oratorius, 94 sordida, 52 
ORYZOCORIS, 60 southwoodi sp. n., 24 
spinosus sp. n., 142 
stali sp. n., 28 
STENOCORIS, 9, 30 
sumatranus, 129 


palawanensis sp. n., 98 
pallida, 46 

papuensis, 140 
phthisica, 49 
PROCAMPTUS, 4 tagalica n. n., 106 
prolixa, 113 tibialis, 123 
pseudolepida sp. n., 100 timoris sp. n., 144 


PSEUDOLEPTOCORISA, 12 tipuloides Dallas, 64 
tipuloides De Geer, 55 


trinotat 
RHABDOCORIS, 69 ie ne a a 
varicornis, 78 
sakdapolrakae sp. n., 102 v-nigrum, 57 
similis, 23 
solomonensis sp. n., 104 XENOCERAEA, 4 


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A LIST -OF SOUPPLEMENTS 
TO THE ENTOMOLOGICAL SERIES 
OF THE BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 


. 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. 

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

. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 
18 plates, 270 Text-figures. August, 1965. {4 4s. 

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


1965. £3558. 


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


yr. OTL, Fa 


DIPTERA FROM NEPAL 


CRYPTOCHAETIDAE, DIASTATIDAE 
& DROSOPHILIDAE 


T. OKADA 


BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Supplement 6 
LONDON : 1966 


DIPTERA FROM NEPAL 


ChyrrocuAcrivre, DIAsTATIDAE 
& DROSOPHILIDAE 


BY 
T. OKADA , 


Dept. of Biology, Faculty of Science 
Tokyo Metropolitan University 


BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Supplement 6 
LONDON: 1966 


THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), wnstituted im 1949, 1s 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

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

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

This paper is Supplement No. 6 of the Entomolog- 
ical series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 


© Trustees of the British Museum (Natural History) 1966 


TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 


Issued 6th May, 1966 Price £3 


DIPTERA FROM NEPAL 


GRYPTOCHAEBTIDAE;: DIASTATIDAE 
& DROSOPHILIDAE 


By T. OKADA 
CONTENTS 
Page 
SYNOPSIS 
INTRODUCTION 4 
ABBREVIATIONS USED IN THE TExt- FIGURES ; ; : , 5 
ACKNOWLEDGEMENTS : : ; : ; ; : : : 6 
Family CRYPTOCHAETIDAE : : : : . : 6 
Family DIASTATIDAE . c : 5 : : 5 : 8 
Family DROSOPHILIDAE : . : : : : g : 14 
Subfamily STEGANINAE ; 3 3 : : : : 15 
Subfamily DROSOPHILINAE . : : : P : F 37 
GEOGRAPHICAL DISTRIBUTION. ; ; : : , P F 112 
EcoLoGicaL DISTRIBUTION : : : : : 117 
NOTES ON SOME SPECIAL SYSTEMATIC CHARACTERS : : : , 119 
A. Archestinic characters : : : : : I1g 
B. Homeotic expression of the hooked scaly bristles ; ; 120 
C. Homology in the components of phallic organs and the systematic 
analysis of phallic formulae. : : : : : : 121 
REFERENCES . : : : : : ; é : : ; 124 
INDEX . : é ‘ : : : : : : : ; 127 
SYNOPSIS 


The Cryptochaetidae, Diastatidae and Drosophilidae collected by Mr. Ralph L. Coe, British 
Museum (Natural History), during the British Museum East Nepal Expedition, 1961-1962 
amounted to twelve hundred and eighty-seven well-preserved dried flies and a puparial case. 
They represent seventy-six species belonging to fifteen genera, including Lissodrosophila gen. n. 
and thirty-seven new species. In addition to describing new forms, insufficiently known species 
have been re-described according to current taxonomy. Brief notes of the geographical and 
ecological distributions of the genera and species concerned are given, and some systematically 
important problems concerning the occurrence of “‘ archestinic ’’ characters in a taxon, the 
homeotic and compensatory expression of characters, and homology in the components of phallic 
organs are analysed and discussed. 


4 T. OKADA 


INTRODUCTION 


THROUGH the recommendation of Dr. E. B. Basden, Institute of Animal Genetics, 
Edinburgh, the author was enabled to examine a large collection of Cryptochaetidae, 
Diastatidae and Drosophilidae from East Nepal, collected by Mr. R. L. Coe while a 
member of the British Museum East Nepal Expedition, Sept. 1961-Febr. 1962. 
The collection is made up of the following fifteen genera : Cryptochaetum (1 sp.) of 
Cryptochaetidae, Diastata (2), Campichoeta (1) and Apsinota (2) of Diastatidae, and 
Stegana (1), Leucophenga (16), Microdrosophila (6), Lissodrosophila (1), Hypselothyrea 
(x), Liodrosophila (4), Paramycodrosophila (1), Chaetodrosophilella (1), Scaptomyza (4), 
Diathoneura (1) and Drosophila (34) of Drosophilidae. 

The genus Lissodrosophila and thirty-seven species belonging to ten genera are 
described as new : Diastata (I sp. n.), Apsinota (1), Leucophenga (4), Microdrosophila 
(5), Lissodrosophila (1), Liodrosophila (3), Chaetodrosophilella (1), Scaptomyza (3), 
Diathoneura (1) and Drosophila (17). 

These genera have apparently not been recorded from Nepal, apart from Dyoso- 
phula, of which the following six species were recorded earlier by the present author 
(Okada, 1955) : D. bipectinata Duda, D. kikkawat Burla, D. melanogaster Meigen, 
D. takahashii Sturtevant, D. nepalensis Okada and D. immigrans Sturtevant. Of 
these, D. bipectinata alone is not recorded in the present collection, which has 
brought the number of Nepalese species of the three families involved to seventy- 
seven. 

The drosophilid fauna of Nepal is most interesting biogeographically and ecologi- 
cally, as it comprises Palaearctic and Oriental elements ranging from low, subtropical 
lands to the highest mountains in the world. In this respect, it closely resembles the 
pattern of the Japanese fauna, which is, though essentially Palaearctic, rich in 
Oriental elements, as for example with the mosquitoes (Edwards, 1921 : 264). In 
addition to describing new forms, some of the poorly-known species are re-described 
in the light of current taxonomic knowledge. 

Some taxonomic characters are explained below. 

Width of front at its posterior margin : measured across the ocellar triangle. 

Width of cheek : the greatest distance from eye margin measured vertically to the 
tangent drawn upon eye margin. 

Wing indices : C-index (costal index), a ratio in length of the second and the 
third costal sections ; 4V-index (4th vein index), a ratio in length of the distal and 
the proximal sections of the vein M (media) ; 4C-index, a ratio in length of the 
third costal section and the proximal section of M ; 5x-index, a ratio in length of the 
distal section of the vein Cw (cubitus) and the posterior crossvein ; Ac-index (acro- 
costal index, Burla, 1956 : 193), a ratio in length of the third and fourth costal 
sections. 

Cr-bristles : strong bristles at the end of the first costal section. 

C3-fringe : the portion of the third costal section bearing the strong costal fringe 
(Wheeler and Takada, 1964 : 168). 

Sterno-index : a ratio in length of the anterior and the ventral (posterior, Kikkawa 


DIPTERA FROM NEPAL 5 


and Peng, 1938) sterno-pleurals (Kikkawa and Peng, 1938 : 552). 

Mesonotum : prescutum and scutum of the morphological mesonotum (Sturtevant, 
EO2T : 28). 

Abdominal tergite : sometimes abbreviated as T, e.g., 1 means the first abdominal 


tergite. 


The terminology of the male and female genitalia is chiefly based on Hsu (1949), 
Nater (1953), and Okada (1954, 1956). The phallic formula (p.f.) is a series of 
alphabetical signs indicating various characters of the phallic organs of each species 
according to the following rules. 


Aedeagus apparently bifid, or (a) fused. 

Aedeagus non-pectinated, or (b) pectinated. 

Aedeagus bare, or (c) pubescent. 

Anterior parameres large, or (d) minute. 

Anterior parameres articulated to the aedeagus, or (e) non-articulated. 

Anterior parameres with sensilla mesad to or scattered evenly on the para- 
meres, or (f) distal, or absent (fo). 

Posterior parameres (penis mantle sheath, Nater, 1953) non-contiguous, or 
(g) contiguous to each other to compose a “‘distal bow”’ (Fota-Pessoa, 1954), 
or (g») absent. 

Posterior parameres non-branched or (h) branched. 

Novasternum with paired submedian spines (hypandrial bristles, Nater, 1953) 
on its caudal margin, or (i) without such spines. 

Novasternum without or (k) with a median notch on its caudal margin. 

Aedeagus vertically or (1) horizontally flattened. 

Aedeagus without or (m) with a prominent basal vertical rod or ventral 
recurved process. 

Basal apodeme of aedeagus as long as or longer than aedeagus, or (n) less than 
half the length of aedeagus. 


A small alphabetical letter with attached dash (’) indicates a feature intermediate 
between those of the corresponding large and small letters. 
Phallosomal index (PI) : a ratio in length of aedeagus and its apodeme. 


i? Oo S Af boo Cee 2 


ABBREVIATIONS USED IN THE TEXT-FIGURES 


. anterior paramere or anterior gonapophysis (phallic organs). 
. decasternum or bridge connecting claspers (periphallic organs). 
aedeagus or penis (phallic organs). 
. genital arch or novatergum (periphallic organs). 
. subterminal hair (egg-guide). 
basal isthmus (egg-guide). 
. Clasper (periphallic organs). 
lobe (egg-guide). 
. novasternum or hypandrial plate (phallic organs). 
. basal apodeme of aedeagus (phallic organs). 
. posterior paramere or posterior gonapophysis (periphallic organs). 


6 Lf OKADA 


vertical rod of aedeagus (phallic organs). 

submedian spine of hypandrium (phallic organs). 
anal plate or cercus (periphallic organs). 

ventral recurved process of aedeagus (phallic organs). 
. ventral fragma of novasternum (phallic organs). 


The scales of the text-figures I-327 : solid line, 0-1 mm. ; broken line, 1-0 mm. 


Se ia AGO 


ACKNOWLEDGEMENTS 


The author wishes sincerely to thank Dr. E. B. Basden, Institute of Animal Gene- 
tics, Edinburgh, for recommending that this material should be offered to the 
author to study. His hearty thanks are due to Mr. R. L. Coe, British Museum 
(Natural History), for entrusting the author with this precious material and reading 
the manuscript, and to Mr. J. P. Doncaster, British Museum (Natural History), for 
sanctioning the loan of the specimens and for accepting this paper for the Museum’s 
Bulletin. 

He is much obliged to Dr. M. R. Wheeler, University of Texas, for his valuable 
advice on taxonomic accounts, and also Dr. F. A. Turk, Shang-ri-la, Reskadinnick, 
Camborne, Cornwall, and Dr. R. H. Cobben, Landourwhogeschool, Wageningen, 
Netherlands, for their kind help in obtaining literature. He acknowledges grants 
from the Japan Society of the Promotion of Science, through the courtesy of Dr. K. 
Dan, Tokyo Metropolitan University, and from the Japan U.S. Co-operative Science 
Program through the courtesy of Dr. D. Moriwaki, Tokyo Metropolitan University. 


Family CRYPTOCHAETIDAE 


CRYPTOCHAETUM Rondani 
Cryptochaetum Rondani, 1875 : 16 
Type-species : Cryptochaetum grandicorne Rondani, 1875 : 167 (Europe). 


Cryptochaetum nipponense (Tokunaga) 
(Text-figs. 1-5) 


Hippelates npponensis Tokunaga, 1943 : 1075 (Japan). 
Cryptochaetum nipponense (Tokunaga) Foote and Arnaud, 1958 : 241 (Japan). 
Cryptochaetum grandicorne Rondani ; Coquillett, 1898 : 340 (Japan) ; Kuwana, 1922 1 : 28 ; 

2 : 37 (Japan) ; Okada, 1956: 9 (Japan). ([Mis-identification.] 

6. Body about3mm. Front large, frontal triangle not sharply pointed anteriorly. Anten- 
nae slightly shorter than face, apically angular, ending in a minute tubercle. Fore tarsi normal, 
not dilated. Posterior crossvein sinuate, anterior crossvein level with the second costal break. 
R, angulate at middle. Costa extending slightly beyond the end of R,1;. 


Taplejung District, damp evergreen oak forest above Sangu, c. 9,200’, 2-26.xi.1961, 
12 § (2 ¢ hovering in front of eyes) (R.L.C.) ; below Sangu, by stream in shady 
ravine, c. 6,000’, I g, 30.x.1961 (R.L.C.). 

Distribution : Nepal (new locality), Japan. 


DIPTERA FROM NEPAL ii 


All the features mentioned above and the detailed structures of male genitalia 
(Text-figs. 2, 5) fully agree with those of the Japanese form examined by the author 
(1956) and confirmed and re-described by Foote and Arnaud (1958). 


Fics. 1-5. Cryptochaetum nipponense (Tokunaga), g. 1, head, lateral aspect ; 2, peri- 
phallic organs, lateral aspect ; 3, head, dorsal aspect ; 4, wing; 5, phallic organs, 
ventral aspect. 


8 tf) OKADA 


Family DIASTATIDAE 


Kry To GENERA 


1 Mesopleura with bristles. Orbitals above the middle of front * : i ; 2 
— Mesopleura bare. Orbitals at the middle of front. Arista pubescent. Anterior 
reclinate orbitals slender. Swubcosta short, not reaching costa 
CAMPICHOETA Macquart (p. 10) 
2 Three orbitals present, anterior reclinate orbitals strong, posterior reclinate orbitals 
slender. Arista with short branches. Swubcosta short, not reaching costa 
DIASTATA Meigen (p. 8) 
— Only one orbital present, strong and reclinate. Arista with long branches both 
dorsally and ventrally. Subcosta complete, reaching costa 
APSINOTA van der Wulp (p. 11) 


DIASTATA Meigen 
Diastata Meigen, 1830 : 94. 


Type-species : Diastata vagans Loew, 1864 : 362 (Europe). 
KEY TO SPECIES 


1 Wing with two large transverse black bands surrounding crossveins. C-index about 


See - . 5 : : . : : : : ussurica (p.8) 
— Wing black only at its base and along posterior crossvein, fuscous along costa. 
C-index about 6:0 . ; 2 : : : ; : : . basdeni (p. 8) 


Diastata ussurica Duda 


Diastata ussurica Duda, 1934 : 8 (Ussuri) ; Okada, 1956: 13 (Japan) ; Okada, 1960b : 165 

(Japan). 

6. Wing patterns somewhat intermediate between those of Ussurian (Duda, 1934) 
and Japanese (Okada, 1956) forms, the black patch covering posterior crossvein 
distinctly narrowing below M as in the Ussurian form, but extending outside 
posterior crossvein as in the Japanese. 

Taplejung District, below Sangu, mixed vegetation on sheltered slopes above river, 
C. 4,000’, 3.1.1962, r ¢ (R.L.C.). 

Distribution : Nepal (new locality), Japan, Ussuri. 


Diastata basdeni sp. n. 
(Text-figs. 12-15) 


2. Body about 3-5 mm., black, slender. Head much broader than thorax. Eyes dark 
purplish red, bare. Antennae orange-yellow ; third segment broad, slightly more than twice 
as long as broad, apical half fuscous, anteroventrally narrowing. Arista black, shortly plumose 
on both sides. Palpus slender, curved, yellow, basally brown, with about three black setae near 
apex below. Ocellar triangle fuscous. Periorbit fuscous, strongly curved inward anteriorly, 
ending before middle of front. Front pale yellow, anterior margin concave, anteriorly slightly 
broader than length down middle, posteriorly half as broad as head width. Face pale yellow, 
shining silvery white in dorsal view. Carina undeveloped, represented by a weak median ridge. 
Cheeks yellowish grey, about two-ninths the greatest diameter ofeye. Postgenablack. Clypeus 
reddish brown. Occiput black, concave, grey pollinose. Anterior reclinate orbital longer than 


DIPTERA FROM NEPAL 


Fics. 6-15. 


6-11, Campichoeta obscuripennis (Meigen), d. 
head, dorsal aspect ; 


organs, ventral aspect ; 
Q. 12, head, lateral aspect ; 
15, wing. 


6, head, lateral aspect ; 7, 
8, wing; 9, periphallic organs, lateral aspect ; 10, phallic 
11, sixth abdominal tergite. 12-15, Diastata basdeni sp. n., 
13, egg-guides, ventral aspect ; 14, ventral receptacle ; 


10 T. OKADA 


proclinate, before and inside proclinate ; proclinate half as long as inner vertical ; posterior 
reclinate minute, behind anterior reclinate ; a minute seta before anterior reclinate. Only one 
strong oral, second oral about one-third length of vibrissa, third as long as second. 

Mesonotum, scutellum, and thoracic pleura black, bluish grey pollinose. One long humeral. 
Prescutellars long, somewhat divergent. Acrostichal hairs in about ten irregular rows. Ante- 
rior dorsocentrals about six-sevenths as long as posteriors ; distance between anterior and 
posterior dorsocentrals about four-fifths of distance between anterior pair. Anterior scutellars 
long, parallel ; posterior scutellars three-fifths as long as anteriors, upright and crossed. Sterno- 
index about 0-6. Legs yellowish brown or dark brown, femora especially dark. All metatarsi 
as long as next three segments together ; fore femur with a few stout but short setae inside near 
apex. Wing fuscous along costa, black at base below first costal section. Posterior crossveins 
narrowly clouded. ,;, straight where it reaches costa. R,,; and M slightly divergent 
distally. C-index about 6-0 ; 4V-index about 1:2 ; 4C-index about 0-3 ; 5x-index about 0-7 ; 
Ac-index about 1-1. Cr1-bristle one, long ; C3-fringe on basal two-thirds. Halteres white. 
Abdominal tergites black, somewhat shining blue. 

Egg-guide lobes pale brown, horizontally flattened, fused to each other, setigerous and 
pubescent, apically narrowing and curved inward. Ventral receptacle large, dark brown, 
proximally narrowing and curved rectangularly, apically with a U-shaped invagination. 


Holotype 9. East NepaL: Taplejung District, above Sangu, damp evergreen 
oak forest, c. 9,200’, 2-26.xi.1961 (R.L.C.), BMNH. 


Paratypes : 2 9, collected together with the holotype (R.L.C.), BMNH and Tokyo 
Metropolitan University. 


Allied to D. vagans Loew, 1864 (Europe), in having wings basally black and ante- 
riorly fuscous, but differs in the less distinct dark cloud on posterior crossvein and in 
the shape of egg-guide (distally not narrowing in vagans, Okada, 1960b) and ventral 
receptacle (apical invagination shallow triangular in vagans, Okada, 1g6o0b). 


CAMPICHOETA Macquart 
Campichoeta Macquart, 1833 : 547. 


Type-species : Diastata obscuripennis Meigen, 1830 : 97 (Europe). 


Campichoeta obscuripennis (Meigen) 
(Text-figs. 6-11) 


Diastata obscuripennis Meigen, 1830 : 97 (Europe). 
Campichoeta obscuripennis (Meigen) Duda, 1934 : 14 (Europe). 
Campichaeta obscuripennis (Meigen) ; Basden and Collin, 1958 : 139 (Europe). 


6. Body about 2:5 mm., matt black. Head as broad as thorax. Eyes dark red, large, 
bare. Antennae matt velvety black, second segment somewhat grey pollinose, third elongate 
and more than twice as long as broad, apically blunt. Palpus black, curved, apically rounded, 
with about two strong black ventral setae. Ocellar triangle matt black, grey pollinose. Peri- 
orbit black, grey pollinose, anteriorly rounded, about two-thirds as long as front, posteriorly 
narrowing. Front matt black, grey pollinose, anteriorly nearly as broad as median length, 
posteriorly about half as broad as the head width. Face black, marginally yellowish white. 
Carina flat, broad, black. Cheeks narrow, yellowish white, marginally black, about one-twelfth 
as broad as the greatest diameter of eye. Anterior reclinate orbital much before and inside 
proclinate, about one-fourth as long as posterior reclinate ; proclinate shortly before and 
outside posterior reclinate, about two-thirds as long as posterior reclinate. Vibrissae short, 
other orals fine. 


DIPTERA FROM NEPAL II 


Mesonotum matt black, somewhat bluish grey pollinose. Scutellum matt black, unicolorous. 
Thoracic pleura black, bluish grey pollinose. Only one prominent humeral. Prescutellars 
long. Acrostichal hairsin eight rows. Anterior dorsocentrals three-fourths length of posteriors, 
distance between anterior and posterior dorsocentrals slightly shorter than distance between 
anterior pair. Anterior scutellars divergent ; posterior scutellars three-fifths length of anteriors. 
Sterno-index about 0-7; median sternopleural slender. Legs yellowish brown ; femora 
darker ; fore femur with a row of about ten short black stout setae anteroventrally on distal 
two-thirds, ventrally with a stout seta near apex. Preapicals well developed on all legs ; apicals 
prominent on mid leg. Wings narrow, deeply fuscous especially along costa. Costa with 
sparse long black setae besides the usual strong fringe. C-index about 4-5 ; 4V-index about 
1-5; 4C-index about 0-6; 5x-index about 2-7 ; Ac-index about 1-5. Cr-bristles two, well 
developed ; C3-fringe on basal one-fifth. Halteres yellowish white. Abdomen black, slender ; 
caudal tergite with about three pairs of long stout black bristles. 

Periphallic organs : Genital arch black, broad, much narrowing above, with microtrichia and 
scattered macrotrichia, toe broad and triangular. Clasper absent. Cercus fusiform, brownish 
black, with thick microtrichia, which are especially long at lower apex. Phallic organs : 
Aedeagus orange-brown, rod-like, straight, apically curved ventrally, basally swollen laterally ; 
apodeme thick, straight. PI about 1-3. Anterior paramere slender, straight, tapering, orange- 
brown, with a few sensilla in a longitudinal row on distal half. Posterior parameres fused to 
form a broad triangular medially pointed black lobe. Ventral fragma short. p.f.—=ABCDEF 
gHikLMN. MHypandrial plate setigerous. 


Taplejung District, between Sangu and Tamrang, mixed shrubs in deep gorge, 
Ga5,200, 10.x1. 1961, 1 fr ex. (R.L.C.): 

Distribution : Nepal (new locality), Europe. 

The § genitalia agree with those figured by Basden and Collin (1958 : Text-fig. 5) 
for the European form. 


APSINOTA van der Wulp 
Apsinota van der Wulp, 1887 : 178. 


Type-species : Apsinota pictiventris van der Wulp, 1887 : 178 (Java). 


KEY TO SPECIES 
1 Front with a pair of dark longitudinal stripes. Mesonotum uniformly greyish white, 
somewhat bluish pollinose. Acrostichal hairs in about four rows. Anterior 
scutellar about twice as long as the posteriors. Legs entirely yellow ; mid tibia 
with row of long setae anteriorly : : : : ‘ : rufipes (p. 11) 
— Body larger and darker. Front with two pairs of dark longitudinal stripes. Mesono- 
tum dark greyish black, with two pairs of bluish white longitudinal stripes. 
Acrostichal hairs in six rows. Anterior scutellar about one and a half times the 
length of posteriors. Legs yellow ; tarsi fuscous ; mid tibia without a row of long 
setae anteriorly : : pictiventris (p. 13) 


Apsinota rufipes sp. n. 
(Text-figs. 16-22) 


3, 2. Body about 2:7 mm., greyish white, blue pollinose. Head round, slightly broader 
than thorax. Eyes large, dark purplish red, bare. Antennae with second segment orange-grey, 
shortly setose ; third elliptical, bright orange-grey, pubescent. Arista with about seven dorsal 
and three ventral long branches and a small terminal fork. Palpus black, elongate, with short 
setae below. Ocellar triangle greyish white. Periorbit short, narrow, one-third as long as front. 
Male front narrow ; anteriorly wider, two-ninths as broad as head width, greyish white, pollin- 


I2 


Fics. 16-26. 


16-22, Apsinota rufipes sp. n. 
dorso-anterior aspect ; 
lateral aspect ; 
Apsinota pictiventris van der Wulp, 9. 


T. OKADA 


16, g head, lateral aspect ; 
18, phallic organs, lateral aspect ; 
20, 9 egg-guides and cerci ; 


19, periphallic organs, 
22, 6 scutellum. 
24, antenna ; 


21, 6 wing ; 
23, head, dorsal aspect ; 


DIPTERA FROM NEPAL 13 


ose, darker above ; with a pair of greyish brown submedian longitudinal stripes, which are 
broadened anteriorly. Female front wider than in g. Face greyish white, pollinose. Clypeus 
black. Carina low. Mouth-parts black. Occiput greyish white, pollinose. Cheek very 
narrow, about one-fiftieth as broad as the greatest diameter of eye. Only one long reclinate 
orbital bristle, situated at one-third distance from the vertex to anterior margin of front. Vib- 
rissae small, second oral minute. 

Mesonotum strongly convex, black, bluish white pollinose. Scutellum greyish white pollinose, 
apically rounded and slightly yellowish grey, with about twelve discal and four marginal setae 
besides the scutellars. Thoracic pleura with two stout mesopleurals. Two subequal humerals. 
Prescutellars longer than anterior dorsocentrals. Acrostichal hairs in four irregular rows. 
Posterior dorsocentrals about two and a half times as long as anteriors ; distance between 
anterior pair of dorsocentrals slightly more than twice distance between anterior and posterior 
pairs. Anterior scutellars divergent, long, about twice the length of scutellum ; posterior 
scutellars convergent, closely approximated, as long as scutellum. Only one prominent sterno- 
pleural (posteroventral), the anterior one minute. Legs yellowish grey, slender ; fore femur 
with a row of about ten short thick black bristles anteroventrally on distal half. Preapicals on 
all legs, apicals prominent on mid leg. Wings hyaline, slender, iridescent ; R,+, straight ; 
costa reaching M ; R,,, ending at wing tip ; R,,; and M apically divergent. C-index about 
5:9; 4V-index about 0-7 ; 4C-index about 0-4 ; 5x-index about 0-4 ; Ac-index about 0:8. 
Cr-bristles two, C3-fringe on basal one-third. Halteres yellowish white. Abdominal tergites 
black, bluish grey pollinose, slender. In 9 each abdominal tergite with an anteromedian small 
longitudinal, and a lateral triangular apically broadened, dark spot. 

Periphallic organs : Genital arch black, broad, narrowing below, lower distal margin with 
short setae. Anal plate conical, black, hairy, separated from genitalarch. Clasper (?) elongate, 
black, apically bifurcated, without teeth. Phallic organs : Aedeagus brown, elongate, large, 
C-shaped in lateral aspect, apically somewhat swollen and forked. Basal apodeme recurved 
and pointed at tip, short. PI about 6-0. Ventral fragma pale brown, quadrate, proximally 
semicircular, distally closely fused to the tip of genital arch. Anterior paramere apparently 
fused to the lateral arm of ventral fragma, with a few long sensilla. Posterior paramere absent. 
p.f.=aBCdefg,HikIMN. 


Holotype g. East NepaLt: Arun Valley, below Tumlingtar, River Sabhaya, 
west shore, evergreen shrubs on sandy shore, c. 1,800’, Q-17.xii.1961 (R.L.C.), 
BMNH. 

Allotype 9. Same data as holotype male. BMNH. 

Paratypes : 44, 4 2, same data as holotype ; 12, Tumlingtar, bare rocky slope 
above River Sabhaya, west bank, c. 1,900’, 8—24.xi1.1961 (R.L.C.), BMNH. 1 6, 
I 9, same data as holotype, deposited at the Tokyo Metropolitan University. 


Resembles A. obscuripes de Meijere, 1911, (Java and Formosa) in wing venation, 
but differs from it in its pale hind tibia and tarsi and unicolorous mesonotum. 


Apsinota pictiventris van der Wulp 
(Text-figs. 23-26) 
Apsinota pictiventris van der Wulp, 1887 : 178 (Java) ; de Meijere, 1908 : 149 (Java) ; Duda, 
1924a : 177 ; 1924b: 224 (Java) ; Duda, 1929 : 415 (Moluccas, New Guinea, Sumatra). 

2. Body about 3:7 mm. Head slightly broader than thorax. Eyes reddish brown, bare, 
large. Antennae with second segment reddish brown ; third thrice as long as broad, somewhat 
pointed apically, greyish brown, basally reddish brown. Arista with about nine dorsal and three 
ventral branches and a moderate terminal fork. Palpus black, slender, without long setae. 
Ocellar triangle greyish brown. Periorbit greyish brown, short, about one-third length of front. 


14 T. OKADA 


Clypeus velvety black. Front anteriorly somewhat swollen above, parallel-sided, matt greyish 
brown, with two pairs of deep brownish black longitudinal stripes, lateral pair on anterior half 
of front, median pair posteriorly divergent and reaching occiput ; eye margin narrowly white. 
Occiput black, grey pollinose. Face black, silvery grey pollinose. Carina large but low. 
Cheeks very narrow, white. Vibrissae fine, no succeeding orals. 

Mesonotum dark greyish black, with two pairs of bluish white posteriorly narrowing longitudi- 
nal stripes, median pair occupying anterior half of mesonotum. Scutellum greyish brown, 
setigerous. Thoracic pleura with two long bristles and about ten minute hairs. Two long 
humerals, upper slightly longer. Prescutellars well developed, two-thirds length of posterior 
dorsocentrals. Acrostichal hairs in six rows. Anterior dorsocentrals about one-third length 
of posteriors, distance between anterior pair of dorsocentrals about one and a half times distance 
between anterior and posterior pairs. Anterior scutellars long, divergent ; posteriors diver- 
gent, two-thirds length of anteriors. Only one long sternopleural. Legs yellow, tarsi and 
apices of femora and tibiae somewhat fuscous. Fore femur with a row of about ten black 
hooklets anteroventrally on distal half. Coxa greyish white. All metatarsi as long as two 
succeeding tarsal segments. Mid tibia without long setae anteriorly. Preapicals on all tibiae ; 
apicals prominent on midleg. Wing hyaline, costa reaching M ; R,1, gently upcurved, straight 
where it meets costa. C-index about 5:0 ; 4V-index about 0-6 ; 4C-index about 0-3 ; 5x-index 
about 0-5 ; Ac-index about o-9. C1-bristles two ; C3-fringe on basal three-fifths. Halteres 
white, basally yellow. Abdominal tergites greyish black, laterally paler ; 1-2T white, with 
yellowish grey laterally broadened anterior cross bands ; 3-6T with a pair of bluish grey 
triangular spots on anterior borders. Cercus elongate. 


Arun Valley, east shore of River Arun below Tumlingtar, evergreen shrubs, 
bordering dry stream-beds, c. 1,800’, 14—23.xi1.1961, I 9 (R.L.C.). 

Distribution : Nepal (new locality), Java, Sumatra, Moluccas, New Guinea. 

Slightly different from the original form as re-described by de Meijere (1908), in 
having median pair of pale mesonotal stripes anteriorly almost equally distant from 
each other and from lateral pair (nearer to each other than to lateral pair in the 
original form). 


Family DROSOPHILIDAE 
KEY TO GENERA 


t Posterior reclinate orbital nearer to inner vertical than to proclinate. Anterior 
reclinate orbital nearly as long as proclinate. Discal and second basal cells 
usually separated (if confluent, costa with several thorn-like warts on the lower 
surface of the third section). Prescutellars present : (STEGANINAE) 2 

— Posterior reclinate orbital nearer to proclinate than to inner vertical. Anterior 
reclinate orbital usually shorter than proclinate. Discal and second basal cells 
confluent. Costa without thorn-like warts on the third section. Prescutellars 


usually absent . , ¢ (DROSO PHILINAE) 3 
2  Discal and second basal cells sone Se Mid ea with a row of long stout bristles 
dorsally. R,,; and M strongly convergent distally. ; ‘ STEGANA (p. 15) 


— Discal and second basal cells confluent. Mid tibia without a row of long stout 
bristles dorsally. R,,; and M not strongly convergent distally 
LEUCOPHENGA (p. 17) 
3. Face flat, without true carina. Postverticals minute. Anal vein abbreviated 
DIATHONEURA (p. 65) 
— Face withcarina . : : 4 
4 Periorbit much prosdencd sae? oss: reels ‘orbital fine. Anterior 
dorsocentrals situated near suture : c . MICRODROSOPHILA (p. 37) 


DIPTERA FROM NEPAL 15 


— Periorbit not greatly broadened anteriorly. Anterior dorsocentrals situated much 
behind suture : : : : : : J : : : 
5 Front broad, glossy, and flat. Postverticals minute or absent. Anal vein and cell 


abbreviated ; : 5 : : : 6 
— Front not exceedingly broad, nor glossy, nor fie. Postverticals usually long. 

Anal vein and cell usually well developed . ; ; : : : = 8 
6 Orbitals two, both proclinate, no reclinate. Antenna exceedingly large. Fore 

femur without a row of anteroventral spinules Z LISSODROSOPHILA (p. 45) 
— Reclinate orbital present. Antenna normal. Fore femur with a row of antero- 

ventral spinules . : ; : : : : : 7 


7 Postvertical and anterior reclinate orbital absent. Only one sternopleural 
HYPSELOTHYREA (p. 47) 
- Postvertical and anterior reclinate orbital present, though minute. Two sterno- 
pleurals. / , . LIODROSOPHILA (p. 48) 
8 Postvertical minute. roe qechincte orbital oe as long as proclinate. Second 
costal section somewhat swollen at tip, with deep incision 
PARAMYCODROSOPHILA (p. 55) 
— Postverticals usually not minute. Anterior reclinate orbital shorter than proclinate. 


Second costal section not swollen apically, with incision shallow : 9 
9 More than two pairs of dorsocentrals Seidl : CHAETODROSOPHILELLA (p. 57) 
— Dorsocentrals in two pairs : ‘ : : ; : : ; : 10 


to Acrostichal hairs in two or four rows. Body and wing slender SCAPTOMYZA (p. 58) 
— Acrostichal hairs usually in six or more rows. Body and wings usually not slender 
DROSOPHILA (p. 66) 


Subfamily STEGANINAE 


STEGANA Meigen 
Stegana Meigen, 1830 : 79. 


Type-species : Drosophila curvipennis Fallén, 1823 :1 (Europe). [teste West- 
wood, 1843 ; according to Wheeler, 1960}. 


Subgenus Steganina Wheeler 
Steganina Wheeler, 1960 : 110 
Type-species: Musca coleoptrata Scopoli, 1763 : 338 (Europe). 


Stegana (Steganina) nigrifrons de Meijere 
(Text-figs. 27-28) 


Stegana nigrifrons de Meijere, 1911 : 418 (Java) ; Duda, 1923: 37 (Formosa) ; Duda, 1924a: 182 
(Java) ; Duda, 1926 : 45 (Sumatra) ; Duda, 1939 : 13 (S. Africa). 
Stegana (Steganina) nigrifrons de Meijere ; Wheeler, 1960 : 110. 


9. Body about 2:2mm. Head slightly narrower than thorax. Eye dark purplish red, bare. 
Antenna with second segment yellowish orange, anteriorly brown ; third velvety black, pube- 
scent, anteroventral corner rectangular, posterior margin rounded, basally orange-yellow. 
Palpus slender, pale yellowish orange, with about three black ventral setae. Ocellar triangle 
black. Periorbit pale yellowish grey, half as long as front, straight, posteriorly somewhat 
broadened. Front deep glossy brown, concave, anterior width nearly two-thirds as broad as 
length down middle, posteriorly two-fifths as broad as head width. Clypeus yellowish grey. 
Face brownish black, lower margin broadly white. Carina short, half as long as face, high, 


16 T. OKADA 


narrow. Cheek white, glossy, black above, about one-fourth as broad as the greatest diameter of 
eye. Occiput brownish black, concave. Orbitals equally distant from eye margin ; anterior 
reclinate orbital two-thirds length of proclinate, twice as distant from posterior reclinate as from 
proclinate. Proclinate slightly longer than posterior reclinate. Vibrissae stout and long, 


succeeding orals fine. 


FIGS. 27-34. 27-28, Stegana (Steganina) nigrifrons de Meijere, 2. 27, head, lateral aspect ; 
28, wing. 29-34, Leucophenga abbreviata Duda, g. 29, head, lateral aspect; 30, 
wing ; 31,Scutellum ; 32, abdominal tergites ; 33, periphallic organs, lateral aspect ; 


34, phallic organs, ventral aspect. 


DIPTERA FROM NEPAL 17 


Mesonotum and scutellum dark brownish black, somewhat glossy. Scutellum triangular, 
somewhat pollinose. Thoracic pleura white, with a broad black longitudinal stripe above. 
One prominent humeral. Humeral callus somewhat pale. Acrostichal hairs in about ten 
irregular rows. Anterior dorsocentrals apparently absent. Anterior scutellars long, divergent ; 
posteriors half as long as anteriors, much nearer to each other than to anteriors. Sterno-index 
about 0-8. Legs greyish white ; knees dark brown ; fore and hind femora brown apically, 
mid femur dark on distal half ; mid and hind tibiae proximally dark. Front metatarsus as long 
as three succeeding tarsal segments together, subequal to the rest of tarsal joints in mid and hind 
legs. Wing brown, anteriorly darker. R,,,strongly convex ; R4,,and M strongly convergent 
apically. Crossveins clouded, posterior crossvein somewhat sinuated. Cu distally curved 
downward. C-index about 1-8 ; 4V-index about 1-9 ; 4C-index about 1-3 ; 5x-index about 
1‘°8 ; Ac-index about 20-0. Cr-bristle one, short ; C3-fringe on basal five-sixths. Abdominal 
tergites dark brownish black, darker than mesonotum. 

Taplejung District, between Sangu and Tamrang, mixed plants by damp cliff in 
deep river gorge, c. 5,200’, 22.xi.1961, 19 (R.L.C.). 

Distribution : Nepal (new locality), Java, Sumatra, Formosa, S. Africa. 


LEUCOPHENGA Mik 
Leucophenga Mik, 1886 : 317. 


Type-species: Drosophila maculata Dufour, 1839 : 50 (Europe). 


In homologizing components of the phallic organs of this genus, the author (1956) 
made some wrong interpretations, which are now amended. The main part of 
aedeagus (misidentified as the main part of posterior paramere) is elongate, usually 
partially bifurcated longitudinally, and distally with a hooked piece, which is 
contiguous with the posterior parameres, and proximally with a stout ventral 
recurved process (u, misidentified as the main part of aedeagus) (cf. pp. 121-122). 


KrEy TO SPECIES 


1 M distally abbreviated, not reaching wing margin : : : abbreviata (p. 18) 
— M distally not abbreviated, reaching wing margin 2 
2 Wing with more or less distinct dark markings . 3 
— Wing clear, without distinct dark markings 6 
3 Wing markings distinctly demarcated 4 
— Wing markings diffuse : : : 2 3 - 5 
4 Wing markings along costa and interrupted near middle : - interrupta (p. 21) 
— Wing markings along costa and on the crossveins , : : .  ornata (p. 21) 
5 Wing markings along costa and below R, . , : i ‘ .  costata (p. 19) 
— Wing markings only below R, . P ‘ : : : . subpollinosa (p. 21) 
6 Third abdominal tergite very short c : 7 
— Third abdominal tergite normal ; ! ‘ : ; : : - : 8 
7 Scutellum laterally black, apically white . 2 : 3 . leucozona (3) (p. 21) 
— Scutellum dark brown, unicolorous . ; , : albifascia (8) (p. 22) 
8 MHalteres with black knob 9 
— Halteres yellow or white . ¢ ; : ‘ II 
9 Mesonotum and scutellum white, Silver pollinose : ; argentata (p. 24) 
— Mesonotum yellowish brown, scutellum dark brownish black and apically white Io 
10 Thoracic pleura without black patches : 4 ‘ : halteropunctata (p. 25) 
— Thoracic pleura with black longitudinal stripe . . leucozona (9) (p. 21) 
1m Thoracic pleura yellow, without dark patches or stripes : : 12 


18 T. OKADA 


— Thoracic pleura dark brownish black or with dark longitudinal stripes . ; z 16 
12 Palpus black or dark brown . é : : : . 13 
— Palpus yellow or orange-yellow or at Test apically Peller : : : : 4 15 
13 Scutellum laterally black, apically pale yellow . ‘ ‘ : . salatigae (p. 27) 
—  Scutellum yellowish grey, basally somewhat fuscous . : : : : . 14 
14 Palpus slender, with long setae : : ‘ ; F : nigripalpis (p. 28) 
-— Palpus large, without setae : : guttiventris (p. 30) 
15 Palpus apically black. Abdominal tergites banded, not spotted . nigriceps (p. 31) 
—  Palpus pale yellowish white. Abdominal tergites spotted : rectinervis (p. 33) 
16 Palpus pale brown. Abdominal tergites mostly black, 1T yellow nigroscutellata (p. 34) 
—  Palpus pale yellow. Abdominal tergites spotted : : : . albiceps (p. 36) 


Leucophenga abbreviata (de Meijere) 
(Text-figs. 29-34) 


Drosophila abbreviata de Meijere, 1911 : 400 (Java). 
Drosomyiella abbreviata (de Meijere) Hendel, 1914 : 114 (Java, Formosa). 
Leucophenga abbreviata (de Meijere) Duda, 1924a : 185 (Java). 

6, 2. Body about 2:2-3:0 mm. Head as broad as thorax. Eyes dark red, bare ; ocelli 
bright red. Antennae with second segment yellowish brown ; third greyish brown, oblong, 
apically rounded. Arista with about seven dorsal and three ventral branches and a small 
terminal fork. Palpus yellowish orange, slender, with only one prominent apical seta, ventrally 
with along fine hair. Ocellar triangle black, pollinose. Periorbit greyish brown, short, medially 
constricted. Front somewhat glossy dirty yellow, darker in caudal aspect, parallel-sided, 
anterior width three-fourths as broad as length down middle, posteriorly two-fifths as broad as 
head width. Clypeus pale yellowish orange, somewhat protruded. Face yellowish grey, 
ventral margin somewhat convex. Postverticals small. Cheeks narrow, about one-thirteenth 
as broad as the greatest diameter of eye. Occiput dark brown, pale above. Posterior reclinate 
orbital slightly longer than others ; anterior reclinate thrice as distant from posterior reclinate 
as from proclinate. Second oral fine, half as long as vibrissa, succeeding orals shorter. 

Mesonotum yellowish brown, matt without prominent patches. Scutellum yellowish brown, 
grey pollinose, tip whitish. Thoracic pleura pale yellowish white, unicolorous. Humerals two, 
strong, upper the longer. Prescutellar longer than anterior dorsocentral, almost equally distant 
from each other and from dorsocentral. Acrostichal hairs in about ten irregular rows. Anterior 
dorsocentrals about one-third as long as posteriors ; distance between anterior and posterior 
dorsocentrals about one-fourth distance between anterior pair. Anterior scutellars divergent, 
long, one and a half times as long as posteriors, which are nearer to each other than to anteriors. 
Sterno-index about 0-5. Legs yellowish white ; last tarsal segment orange ; front metatarsi 
subequal to the remaining segments together, mid metatarsi twice next two segments to- 
gether, and hind metatarsi subequal to next three segments together. Preapicals on mid 
and hind legs ; apicals on mid and hind legs. Knees somewhat fuscous. Wings hyaline, 
somewhat fuscous ; veins yellowish brown, crossveins not clouded. R,;, slightly curved 
to costa at tip; R,4,; slightly convex anteriorly ; M apically abbreviated. C-index about 
2:3; 4V-index about 2-0; 4C-index about 1-3; 5x-index about 1-0 ; Ac-index about 
1-7. Cr1-bristles two, C3-fringe on basal two-thirds. Halteres yellow, apically black. 
Abdominal tergites yellowish or reddish brown ; 2T medially and laterally with black spots ; 
3-51 with caudal black bands projected forward at middle ; 6T mostly black, often with an 
obscure caudal band. 

Periphallic organs brownish black. Genital arch narrow above, truncate below, without 
microtrichia, posterior margin with several long setae ; toe somewhat projected posteriorly ; 
heel low, rectangular. Anal plate vertically elongate, fusiform, setigerous. Clasper nearly 
quadrate, pubescent, and setigerous. Phallic organs : Aedeagus black, basally pale, elongate, 
bifurcated on distal half, basally with a distally broadening apically pointed pale yellowish 


DIPTERA FROM NEPAL 19 


brown recurved process ; hooked piece yellowish orange. Anterior paramere fusiform, with 
long stalk, subapically with several sensilla. Ventral fragma anteriorly pointed, lateromedially 
somewhat swollen. PI =o. p.f. = ABCDeFgHikLmN. 


Taplejung District, Sangu mixed vegetation by stream in gully, c. 6,200’, 9.xi.1961, 
3 9; below Sangu, by stream in shady ravine, c. 6,000’, 30.x.1g61, 1 gf ; Arun 
Valley : east shore of River Arun below Tumlingtar, evergreen shrubs bordering 
dry stream-beds, c. 1,800’, 14-23.xil.1961, 24 J, 26 9 (R.L.C.). 

Distribution : Nepal (new locality), Java, Formosa. 


Leucophenga costata sp. n. 
(Text-figs. 35-37) 


9. Body about 2-3 mm. Head as broad as thorax. Eyes dark red, bare. Antennae with 
second segment yellowish grey ; third grey, basally white, apically dark orange. Palpus 
comparatively large, yellowish brown, apically rounded, with a subapical and a ventral black 
setae. Ocellar triangle black, grey pollinose. Periorbit black, grey pollinose, broad, two-thirds 
as long as front. Front yellowish grey, somewhat glossy, whitish and anterior width about 
seven-eighths as broad as length down middle, posteriorly one-third as broad as head width. 
Clypeus yellowish grey. Face greyish white, matt, concave. Carina low. Cheeks white, 
narrow, about one-twelfth as broad as the greatest diameter of eye. Occiput dark brown. 
Posterior reclinate orbital slightly longer than others ; anterior reclinate thrice as far from 
posteriors as from proclinate. One prominent oral ; second fine, one-third length of vibrissa. 
Postvertical rather long. 

Mesonotum dark yellowish brown, black behind, grey pollinose ; humeral callus grey polli- 
nose, dorsally bordered by a somewhat fuscous patch. Scutellum dark brownish black, grey 
pollinose. Thoracic pleura dark yellowish orange, with two black broad longitudinal stripes, 
upper one extending to postnotum ; lower short, limited to sternepisternum. One long 
humeral, a few shorter below it. Prescutellar longer than anterior dorsocentral. Acrostichal 
hairs in about eight irregular rows. Anterior dorsocentrals one-third length of posteriors ; 
distance between anterior and posterior dorsocentrals about one-third distance between anterior 
pair. Anterior scutellars divergent, longer than posteriors ; posterior scutellars four-fifths as 
long as anteriors, nearer to each other than to anteriors. Sterno-index about 0-7. Legs 
yellowish white, knees of mid and hind legs black ; all metatarsi much longer than remaining 
tarsal segments together. Wings hyaline, fuscous on and below costa ; costa reaching R445. 
Crossveins clear. R,,, weakly curved to costa at apex ; &,,; and M nearly parallel. C-index 
about 1-9 ; 4V-index about 2-4 ; 4C-index about 1-4 ; 5x-index about 1-5 ; Ac-index about 
2-0. Cr-bristles two, subequal ; C3-fringe on basal five-sevenths. Halteres yellowish white. 
Abdominal tergites glossy black ; 2T posteriorly silvery white pollinose, ultimate T white, 
3-51 latero-anteriorly with white patches. 


Holotype 2. East NEPAL: Arun Valley, east shore of River Arun below Tum- 


lingtar, evergreen shrubs bordering dry stream-beds, c. 1,800’, 14-23.xii. 1961 
@-L.C.), BMNH. 


Paratypes : 2 9, same data as holotype ; 1 9, Taplejung District, between Sangu 
and Tamrang, mixed shrubs in deep gorge, c. 5,200’, 10.xi.1961. (R.L.C.), BMNH 
and at Tokyo Metropolitan University (1 9 from Arun Valley). 

Similar to L. mgroscutellata Duda, 1924, (Formosa) in having black palpus, 
striped thoracic pleura, yellow halteres, black costa, and clear crossveins, but dif- 
fers in having entirely black scutellum (apically yellow in mgroscutellata) and palpus 
with only a moderately strong apical seta. 


20 


Tt, OKADA 


FIGS. 35-43. 35-37, Leucophenga costata sp.n., 3. 35, head, lateral aspect ; 36, head, 
dorsal aspect ; 37, wing. 38-39, Leucophenga subpollinosa de Meijere, 3. 38, peri- 
phallic organs, lateral aspect ; 39, phallic organs, ventral aspect. 40-43, Leucophenga 
leucoxona Duda, 3. 40, head, lateral aspect ; 41, wing; 42, phallic organs, ventral 
aspect ; 43, periphallic organs, lateral aspect. 


DIPTERA FROM NEPAL ZY. 


Leucophenga subpollinosa de Meijere 
(Text-figs. 38-39) 

Leucophenga subpollinosa de Meijere, 1914 : 263 (Java) ; Duda, 1923 : 27 (Formosa) ; Duda, 
1924a : 186 (Java, Annam, Formosa) ; 1924b: 237 (Formosa) ; Duda, 1926 : 52 (Sumatra) ; 
Duda, 1939 : 42 (Africa) ; Okada, 1956 : 30 (Japan) ; Wheeler and Takada, 1964 : 229 (Palau). 
Periphallic organs : Genital arch black, broadly truncate below, heel and toe rectangular. 

Anal plate black, elongate, fusiform. Clasper oval, setigerous tip, rounded. Decasternum 

laterally oblong, yellowish brown. Phallic organs : Aedeagus slender, distal half black and 

bifid, basally with pale brown, rod-shaped, ventral recurved process. Anterior paramere elon- 
gate, triangular, medially with a few sensilla. Ventral fragma narrow, proximal half rod- 
shaped. Posterior parameres triangular. PI = oo. p.f. = ABCDeFgHikLmN. 

Taplejung District, river banks below Tamrang Bridge, c. 5,500’, x—xi.1961, I J ; 
below Sangu, mixed vegetation on sheltered slopes above river, c. 4,000’, 3.1.1962, 
Ig ; Dobhan, small pockets of plants on arid slopes above River Maewa, c. 3,500’, 
2.1.1962, 12 J, 15 2; ibid., shady places on shrubby slope above River Tamur, 
C. 3,500’, 21-27.1.1962, 9 J, 9 2; Arun Valley, east shore of River Arun below 
Tumlingtar, evergreen shrubs bordering dry stream-beds, c. 1,800’, 14—23.xil.1961, 
ie ro 2 (R.L.C.). 

Distribution : Nepal (new locality), Java, Sumatra, Viet-Nam, Formosa, Japan, 
Africa, Micronesia. 

Slightly different from the Japanese form (Okada, 1956) in g genital organs ; 
clasper apically rounded (pointed and clawed in Japanese form). 


Leucophenga ornata Wheeler 


Leucophenga ornata Wheeler, 1959 : 184 ; Okada, 1964a : 105 (Japan : Amami). 
Drosophila ornatipennis Williston ; de Meijere, 1914 : 256 (Java). 
Leucophenga ornatipennis (Williston) ; Duda, 1924a : 186 (Java) ; Okada, 1956 : 37 (Japan) ; 
Paik, 1958 : 211 (Korea). 
Arun Valley, east shore of River Arun below Tumlingtar, evergreen shrubs 
bordering dry stream-beds, c. 1,800’, 14—23.xii.1961, I 2 (R.L.C.). 
Distribution : Nepal (new locality), Java, Japan, Korea. 


Leucophenga interrupta Duda 
Leucophenga interrupta Duda, 1924a : 187 ; 1924b : 237 (Formosa) ; Okada, 1956 : 36 (Japan); 
Okada, 1964a : 106 (Amami : Japan). 
Arun Valley : above River Sabhaya, east shore, swept from dwarf bamboos in 
deep ravine, c. 2,000’, 12.xii.1961, I g (R.L.C.). 
Distribution : Nepal (new locality), Formosa, Japan. 


Leucophenga leucozona Duda 
(Text-figs. 40-43, 96) 
Leucophenga leucozona Duda, 1924a : 188 (New Guinea). 


3, 2. Body about 2:2-2:8 mm. Head as broad as thorax. Eyes deep red, large, bare. 
Antennae with second segment yellow ; third grey, oblong, twice as long as broad. Arista with 


22 T. OKADA 


five to eight dorsal and two or three ventral branches besides a small fork. Palpus yellowish 
orange, with short black stout setae below. Ocellar triangle black. Periorbit yellowish grey, 
medially narrowing, two-thirds as long as front. Front yellowish grey, parallel-sided, anterior 
width two-thirds or three-fourths as broad as length down middle, posteriorly two-fifths as 
broad as head width, medially slightly fuscous. Face greyish white, laterally concave. Carina 
flat. Cheeks yellow, very narrow, about one-fourteenth as broad as the greatest diameter of eye. 
Orbitals equal sized ; anterior reclinate four times as distant from posterior reclinate as from 
proclinate. Only one prominent oral. Postverticals rather long. 

Mesonotum deep yellowish orange, unicolorous. Scutellum deep yellowish orange, laterally 
black, apically yellowish white. Thoracic pleura yellow, with a black longitudinal obscurely 
defined stripe. Prescutellar long, as long as anterior dorsocentral. Acrostichal hairs in about 
ten rows. Anterior dorsocentrals two-thirds length of posteriors ; distance between anterior 
and posterior dorsocentrals about two-sevenths distance between anterior pair. Anterior 
scutellars divergent, slightly longer than scutellum ; posteriors slightly shorter than anteriors. 
Sterno-index about 0:8. Legs yellow, thick; mid and hind knees slightly fuscous. Mid 
metatarsi sub-equal in length to the other segments together, fore and hind metatarsi slightly 
longer than the other segments together. Preapicals on all legs ; those on mid leg large. 
Wing hyaline, crossveins clear. R,;, gently curved to costa at tip; R,;; and M somewhat 
divergent apically. C-index about 2:2 ; 4V-index about 2:0; 4C-index about 1-1 ; 5x-index 
about 1-4 ; Ac-index about 2:0. Cr1-bristles two ; C3-fringe on basal seven-tenths. Halteres 
yellow, slightly black at tip. Abdominal tergites: In g, 1T yellow; 2T anteriorly black, 
posterior half silvery white ; 3T very short, silvery white, without macrotrichia ; 5—6T black, 
dorsolaterally with small yellowish spots ; anal cerci yellow. In 9, 2T with lateral black 
spot ; 3I not short ; 3-5T black, lateroanteriorly yellow ; 6T yellow ; anal cercus yellow. 

Periphallic organs : Genital arch white, narrow above, broadly truncate below, heel and toe 
nearly rectangular, with evenly distributed microtrichia and several setae on posterior and ven- 
tral margins. Anal cercus white, small, fusiform, setigerous, anteriorly convex. Clasper white, 
oval, pubescent, setigerous on lower half. Phallic organs : Aedeagus black, entirely bifurcated, 
apically with an orange-yellow hooked piece, basally with a pale rod-shaped recurved process. 
Anterior paramere pale, elongate, fusiform, distally with a few sensilla. Posterior parameres 
triangular, orange-brown. Ventral fragma proximally pointed, laterally somewhat dilated. 
PI=o. pf. = ABCDEFgHikLmN. 


Taplejung District, between Sangu and Tamrang, mixed shrubs in deep gorge, 
c. 5,200’, X-x1.1961, I ¢ ; Arun Valley, east shore of River Arun below Tumlingtar, 
evergreen shrubs bordering dry stream-beds, c. 1,800’, 14-23.xli.I961, 15 g, 5 @ 
(RE-C). 

Distribution : Nepal (new locality), New Guinea. 

Agrees with the original form (New Guinea, Duda, 1924a) in having yellow palpus, 
yellow and apically black halteres, and striped thoracic pleura, but differs in having 
no prominent macrotrichia on g 3T. 


Leucophenga albifascia sp. n. 
(Text-figs. 44-48, 95) 


3. Body about 2-5 mm. Head slightly broader than thorax. Eyes dark red, large, bare. 
Ocelli bright deep red. Antennae with second segment yellow, third grey. Arista with about 
six dorsal and three ventral branches and a small terminal fork. Palpus orange-yellow, slender, 
ventrally with a few strong black setae. Ocellar triangle black, grey pollinose. Periorbit 
brown, obscurely demarcated from front, anteriorly broadened, two-thirds as long as front. 
Front narrow, nearly parallel-sided, dark yellowish grey, anteriorly paler, anterior width as 


DIPTERA FROM NEPAL 


———___ __, 


Fics. 44-52. 44-48, Leucophenga albifascia sp. n., 3. 44, head, lateral aspect; 45, 
head, dorsal aspect ; 46, wing; 47, periphallic organs, lateral aspect ; 48, phallic 
organs, ventral aspect. 49-52, Leucophenga argentata de Meijere, 3. 49, head, lateral 
aspect ; 50, phallic organs, ventral aspect ; 51, periphallic organs, lateral aspect ; 
52, wing. 


23 


24 T. OKADA 


broad as length down middle, posteriorly about two-sevenths as broad as the head width. Face 
greyish white, flat, laterally concave. Carina undeveloped. Cheeks white, very narrow, about 
one-twentieth as broad as the greatest diameter of eye. Occiput brownish black. Anterior 
reclinate orbital thrice as distant from posterior reclinate as from proclinate. Postverticals 
slender. Inner vertical much shorter than outer vertical. Vibrissae long, succeeding orals about 
one-third as long as vibrissa. 

Mesonotum dark orange-brown, silver pollinose, mediocaudally and laterally more fuscous. 
Scutellum dark brown, grey pollinose. Thoracic pleura pale yellow, orange below, with an 
anteriorly narrowing dark brownish black median longitudinal stripe. Humerals two, upper 
longer. Prescutellar as long as anterior dorsocentral. Anterior dorsocentrals about half length 
of posteriors ; distance between anterior and posterior dorsocentrals about two-thirds distance 
between anterior pair. Anterior scutellars divergent, slightly longer than posteriors, which are 
nearer to each other than to anteriors. Sterno-index about 0-9. Legs yellow ; knees of hind 
leg somewhat fuscous ; last tarsal segment brownish. Front and mid metatarsi slightly longer 
than the other segments together, hind metatarsi as long as the other segments together. Pre- 
apicals on mid and hind legs ; apicals prominent on mid leg. Wings hyaline, crossveins clear. 
R,+4, nearly straight, gently curved to costa apically ; R.i, and M nearly parallel. C-index 
about 2:2 ; 4V-index about 2:1 ; 4C-index about 2:2 ; 5x-index about 1-3 ; Ac-index about 
2°5. Cr-bristles two ; C3-fringe on basal two-thirds. Halteres yellow, tip glossy black. 
Abdominal tergites black, matt; 1T yellow; 2T medioanteriorly yellow, caudal margin 
narrowly white ; 3T very short, white, without setae. Cerci black. 

Periphallic organs : Genital arch black, pubescent, broad ; heel low, triangularly pointed, 
with a few stout long setae ; toe nearly rectangular ; caudal margin with several long setae. 
Anal plate rather small, fusiform, setigerous, pubescent, ventral tip pointed. Clasper oval, 
pubescent and setigerous, proximally somewhat narrowing, distally rounded. Decasternum 
a narrow transverse bar. Phallic organs: Aedeagus black, flat, oblong ; ventral recurved 
process pale yellow, rod-shaped, parallel-sided, twice as long as aedeagus. Anterior paramere 
oblong, pale, with a row of several sensilla on nearly entire length. Ventral fragma appearing 
pointed proximally, laterally pale, fusiformly dilated. PI = oo. pf. = ABCDeFgHikLmN. 


Holotype g. East NepaL: Arun Valley, west shore of River Arun, below 


Tumlingtar, evergreen shrubs bordering dry stream-beds, c. 1,800’, I14—23.xil. 1961 
(K.£.C.); BMINE: 


Closely resembles L. leucozona Duda in having very narrow white 3T in 3, halteres 
with black knob, striped thoracic pleura, and yellowish orange palpus, but differs 
from it in having unicolorously dark brownish black scutellum, 2T only yellow 
anteriorly and cercus black. 


Leucophenga argentata (de Meijjere) 
(Text-figs. 49-52, 73) 


Drosophila argentata de Meijere : 258 (Java). 
Leucophenga argentata (de Meijere) Duda, 1924a : 188 ; 1924b : 238 (Java, Formosa) ; Sturte- 
vant, 1927 : 364 (Philippine) ; Wheeler and Takada, 1964 : 226 (Palau). 


3,9. Body about 2-5mm. Head slightly broader than thorax. Eyes reddish brown, large, 
bare. Antennae milky white ; third segment twice as long as broad, tip somewhat angular 
anteroventrally. Arista with about seven dorsal and three ventral branches and a small 
terminal fork. Palpus pale brown, darker in female, small, apically fuscous, ventrally with a 
few long black setae. Ocellar triangle black, grey pollinose. Periorbit milky white, obscurely 
demarcated from front. Front milky white, flat, parallel-sided, anterior width four-sevenths as 


DIPTERA FROM NEPAL 25 


broad as length down middle, posteriorly two-sevenths as broad as head width. Clypeus greyish 
white, medially white. Face white, narrow. Carina undeveloped. Cheeks white, narrow, 
about one-twentieth as broad as head width. Occiput dark brownish black, white above. 
Anterior reclinate orbital four times as distant from posterior reclinate as from proclinate, which 
is slightly shorter than reclinates ; posterior reclinate slightly nearer to proclinate than to inner 
vertical. Inner vertical one-third as long as outer vertical. Postverticals short, half as long as 
inner verticals. Only one prominent oral. 

Male mesonotum white, densely silver pollinose ; scutellum silver pollinose, unicolorous. 
Female mesonotum yellowish white ; scutellum yellowish white, apically white. Thoracic 
pleura white, matt, unicolorous. Only one long median humeral. Prescutellar as long as 
anterior dorsocentral. Acrostichal hairs in about eight rows. Anterior dorsocentrals two- 
thirds length of posteriors ; distance between anterior and posterior dorsocentrals about two- 
thirds distance between anterior pair. Anterior scutellars divergent, posteriors slightly nearer 
to each other than to anteriors. Sterno-index about 0-8. Legs white ; all metatarsi subequal 
in length to the other segments together. Preapicals on mid and hind legs ; apicals prominent 
on mid leg. Wings hyaline, slightly pointed at tip ; crossveins clear ; costa ending at the tip 
of Ry, ; R, and R,,, apically black ; R,,, gently curved to costa apically ; Ry, and M nearly 
parallel. C-index about 1-8 ; 4V-index about 2-9 ; 4C-index about 1-9 ; 5x-index about 1:9 ; 
Ac-index about 2:4. Cr-bristles two ; (C3-fringe on basal five-sevenths. Halteres white, 
knob black. Abdominal tergites yellowish white. In male, 2T laterally with a small black 
spot ; 4-5T medially with a long black stripe, laterally with two large black spots. In female, 
2T laterally with a large black spot ; 3—4T with five black spots ; 5T with seven black spots ; 
6T with two black spots. 

Periphallic organs yellowish white. Genital arch broad, gently narrowing below, pubescent, 
with a few setae on upper caudal margin, lower tip, and at middle ; heel rectangular ; toe 
rounded. Anal plate semicircular, setigerous and pubescent. Clasper nearly quadrate, 
broader than long, with long setae on caudal margin and inferior surface. Phallic organs : 
Aedeagus black, elongate, distal half bifurcated, basally with a pale yellowish brown elongate 
ventral recurved process, which is longer than aedeagus and apically slightly pointed. Anterior 
paramere long, narrow, slightly swollen medially, apically with a few sensilla. Posterior 
paramere brown, attached to a dark brown hooked piece of aedeagus. Ventral fragma black, 
proximally pointed. PI = oo. pf. = ABCDefgHikLmN. 


Taplejung District, Dobhan, evergreen trees overhanging stream in deep gully, 
c. 3,500’, 30.1.1962, 1g ; Arun Valley, east shore of River Arun below Tumlingtar, 
evergreen shrubs bordering dry stream-beds, c. 1,800’, I14—23.xul.1961, 6 3, 5 92; 
above River Sabhaya, east shore, swept from dwarf bamboos in deep ravine, c. 2,000’, 
ie ,Ri. FOOT, t ¢. (R.L-C.): 

Distribution : Nepal (new locality), Java, Formosa, Philippines, Palau. 


Leucophenga halteropunctata Duda 
(Text-figs. 53-55, 71) 


Leucophenga halteropunctata Duda, 1923 : 28 (Formosa) [nom. nud.]; Duda, 1924a: 188 ; 
1924b : 239 (Formosa) ; Sturtevant, 1927 : 364 (Philippine) ; Burla, 1954 : 29 (Africa) ; 
Wheeler and Takada, 1964 : 228 (Palau). 


2. Body about 2mm. Head as broad as thorax. Eyes dark red, bare. Antennae with 
second segment orange-brown ; third greyish yellow. Arista with about five dorsal and two 
or three ventral branches and a moderate terminal fork. Palpus dark brownish black, slender, 
with an apical and two ventral long setae. Ocellar triangle black, grey pollinose, Periorbit 
greyish yellow, not sharply demarcated. Front yellowish grey, parallel-sided, somewhat glossy, 


26 


T. OKADA 


Fics. 53-61. 


head, dorsal aspect ; 55, wing. 56-61, Leucophenga salatigae de Meijere. 


53-55, Leucophenga halteropunctata Duda, 9. 53, head, lateral aspect ; 


lateral aspect ; 57, ¢ head, dorsal aspect ; 


aspect ; 


60, phallic organs, lateral aspect ; 


54) 


56, 3d head, 


58, d wing ; 59, periphallic organs, lateral 


61, 2 egg-guide. 


DIPTERA FROM NEPAL 27 


anterior width three-fourths as broad as length down middle, posteriorly slightly less than one- 
third as broad as head width. Carina flat, grey. Buccal margin somewhat convex. Cheek 
narrow, yellowish white, about one-twentieth as broad as the greatest diameter of eye. Occiput 
dark brown, paler above. Proclinate orbital slightly shorter than others ; anterior reclinate 
just outside proclinate. Inner vertical much shorter than outer vertical. Postvertical minute. 
One prominent oral. 

Mesonotum orange-yellow, anterolaterally and caudomedially yellowish brown ; neck region 
brownish black above. Scutellum dark brown, laterally with a black patch at the insertion of 
anterior scutellar. Thoracic pleura white, without dark patches. Only one humeral. Pre- 
scutellar slightly longer than anterior dorsocentral. Acrostichal hairs in about nine irregular 
rows. Anterior dorsocentrals about half length of posteriors ; distance between anterior and 
posterior dorsocentrals slightly longer than one-third distance between anterior pair. 
Anterior scutellars divergent ; posteriors as long as anteriors, equally distant from each 
other and from anteriors. Sterno-index about 0-8. Legs yellowish white ; femora white ; 
tibiae fuscous apically and tarsi slightly so. Preapicals on mid and hind legs ; apicals 
on mid leg. Wing hyaline ; veins yellowish brown ; crossveins not clouded, posterior 
crossveins somewhat sinuated. R,,;, nearly straight, apically curved to costa; R44, and M 
nearly parallel, considerably convergent basally. C-index about 1:7; 4V-index about 3:1 ; 
4C-index about 1-9 ; 5x-index about 1-3 ; Ac-index about 2-8. C1-bristles two ; C3-fringe on 
basal three-sevenths. Halteres yellow, base orange, knob glossy black. Abdominal tergites 
yellowish brown, with black spots as follows : two small ones on 1T (usually absent), four outer 


larger, on 2T, three on 3T, median one large, five large ones on 4T, five small ones on 5T, and 
two small ones on 6T. 


Arun Valley, east shore of River Arun below Tumlingtar, evergreen shrubs 
bordering dry stream-beds, c. 1,800’, 14-23.xii.1961, 10 2 (R.L.C.). 
Distribution : Nepal (new locality), Formosa, Philippines, Palau, Africa. 


Leucophenga salatigae de Meijere 
(Text-figs. 56-61, 72) 


Leucophenga salatigae de Meijere, 1914 : 260 (Java). 
Leucophenga albicincta de Meijere var. Salatigae de Meijere ; Duda, 1939 : 47 (Africa). 


3,. Body about 3mm. Head as broad as thorax. Eyes dark red, large, bare. Antennae 
with second segment yellowish grey ; third grey, twice as long as broad, pale at base of arista, 
which has about six dorsal and two ventral branches and a small terminal fork. Palpus yellowish 
orange, slender, curved upward, apical half dark brown, with short setae below. Ocellar tri- 
angle black. Periorbit yellowish grey, not reaching anterior extremity of front. Front yellow- 
ish grey, matt, anteriorly pale, anterior width narrower than length down middle, posteriorly 
about two-fifths as broad as head width. Clypeus yellowish grey. Face yellowish white. 
Carina flat. Cheek very narrow, about one-twelfth as broad as the greatest diameter of eye. 
Occiput black. Postverticals moderate in size. Posterior reclinate orbital nearly twice as 
long as others ; anterior reclinate slightly behind proclinate and thrice as distant from posterior 
reclinate as from proclinate. Only one prominent oral, others slender. 

Mesonotum orange-brown, convex, unicolorous. Scutellum yellowish orange, apically pale 
yellow, laterally black. Thoracic pleura yellow, with a broad black longitudinal stripe for 
entire length. Humeral callus yellow. Humerals three, uppermost longest, lowermost shortest. 
Acrostichal hairs in about eight rows. Prescutellar nearly as long as anterior dorsocentral. 
Posterior dorsocentrals nearly twice as long as anteriors ; distance between anterior and posterior 
dorsocentrals about one-third distance between anterior pair. Anterior scutellars long, diver- 
gent ; posteriors two-thirds length of anteriors and as long as scutellum. Sterno-index about 


28 T. OKADA 


0-6. Legs yellow ; knees not absolutely black ; front metatarsi slightly longer than the other 
segments together, mid and hind metatarsi much longer than the other segments together. 
Preapicals on mid and hind legs, apicals prominent on mid leg. Wing slightly fuscous especially 
on anterior half, with black patch below R, ; crossveins slightly clouded. C-index about 2:4 ; 
4V-index about 1:2; 4C-index about 0-9; 5x-index about 1:6; Ac-index about 2:1. Cr1- 
bristles two ; C3-fringe on basal three-fourths. Halteres whitish yellow, not fuscous. Abdom- 
inal tergites black ; 1T yellow, 2T anteromedially yellow, 3T yellow on anterolateral margin. 
Abdominal sternites pale yellow. 

Periphallic organs : Genital arch black, broad, ventrally broader and truncate, pubescent and 
caudally with setae ; toe triangularly pointed caudally ; heel low, rectangular. Clasper oval 
pale brown, darker anteriorly, setigerous and pubescent. Anal plate fusiform, pale brown, 
setigerous but without microtrichia, ventral tip black. Phallic organs : Aedeagus pale brown, 
slender, bifid on distal two-thirds ; ventral recurved process slender, nearly as long as aedeagus ; 
apical hooked piece yellowish orange. Anterior paramere elongate, blade-like, pale yellowish 
brown, medially with a few sensilla. Posterior parameres dark brown, fused to be a quadrate 
lobe. Ventral fragma with distinct submedian spines. PI = oo. pf. = ABCDeFgHIkLmN. 


Taplejung District, Sangu, mixed vegetation by stream in gully, c. 6,200’, ix-x: 
1961, 1 9 ; between Sangu and Tamrang, shrubs by path, c. 5,800’, 6.xi.1961, 3d ; 
ibid., mixed shrubs in deep gorge, c. 5,200’, x—-xi.1961, 19g, 3 2 ; ibid., mixed plants 
by damp cliff in deep river gorge, c. 5,200’, 22.xi.1961, 3 gd, 1 9. Arun Valley, below 
Tumlingtar, River Sabhaya, west shore, evergreen shrubs on sandy shore, c. 1,800’, 
Q=17. MIL ROOK, 16, ORC.) 

Distribution : Nepal (new locality), Java, Africa. 

Differs from the original form (Java, de Meijere, 1914) in having pale yellow 
halteres (fuscous in original form) and much smaller 4V-index (2.3 in original form). 
Occurrence of submedian spines on ventral fragma of phallic organ seems to be unique 
among the genus Leucophenga, although it is prevalent in the subfamily Drosophi- 
linae. 


Leucophenga nigripalpis Duda 
(Text-figs. 62-66) 
Leucophenga nigripalpis Duda, 1924a : 188 (Formosa). 


¢. Body about2mm. Headas broad as thorax. Eyes reddish brown, large, bare. Anten- 
nae with second segment pale yellowish brown, third greyish yellow. Arista with about five to 
six dorsal and two to three ventral branches and a moderate terminal fork. Palpus black, 
slender, with an apical and a few ventral long black setae. Ocellar triangle black within the 
bright red ocelli. Periorbit greyish yellow, short, medially constricted. Front pale yellowish 
grey, somewhat glossy, flat, parallel-sided, anteriorly yellowish white and two-thirds as broad 
as length down middle, posteriorly slightly narrower than one-third head width. Face flat, 
yellowish white ; buccal margin slightly convex. Cheek white, narrow, about one-fifteenth as 
broad as the greatest diameter of eye. Orbitals subequal in size ; anterior reclinate orbital 
just outside and behind proclinate, four times as distant from posterior reclinate as from proclin- 
ate. Postverticals small. Vibrissae rather short, second oral much shorter than vibrissa. 
Clypeus pale brown. 

Mesonotum pale yellowish orange, laterally and caudally fuscous. Scutellum yellowish brown, 
anterolaterally fuscous. Thoracic pleura pale yellowish orange, without dark patches. Two 
humerals. Prescutellar longer than anterior dorsocentral. Acrostichal hairs in about ten 
irregular rows. Anterior dorsocentrals short, about two thirds length of posteriors ; distance 


DIPTERA FROM NEPAL 


Fics. 62-75. 62-66, Leucophenga nigripalpis Duda, 3. 62, head, lateral aspect ; 63, 
head, dorsal aspect ; 64, wing; 65, periphallic organs, lateral aspect ; 66, phallic 
organs, ventral aspect. 67-70, Leucophenga guttiventris Duda, 2. 67, head, lateral 
aspect ; 68, head, dorsal aspect ; 69, wing ; 70, abdominal tergites. 71-75, abdominal 
tergites of 71, Leucophenga halteropunctata Duda, 2 ; 72, L. salatigae de Meijere, 3; 73, 
L, avgentata de Meijere,3 ; 74, L. nigroscutellata Duda, 2; 75, L.albiceps de Meijere ¢. 


29 


30 T. OKADA 


between anterior and posterior dorsocentrals about two-sevenths distance between anterior pair. 
Anterior scutellars divergent, slightly longer than posteriors, which are a little nearer to each 
other than to anteriors. Sterno-index about 0-7. Legs yellowish white ; mid and hind knees 
somewhat fuscous. All metatarsi longer than the other segments together. Preapicals on all 
legs ; apicals prominent on mid leg. Wings hyaline, slightly pointed at tip ; crossveins clear. 
R43 nearly straight ; R,,; and M somewhat convergent distally ; Cw black, other veins pale. 
C-index about 2:0 ; 4V-index about 2-2 ; 4C-index about 1-4 ; 5x-index about 1-1 ; Ac-index 
about 2:6. Cz1-bristles two ; C3-fringe on basal about three-fifths. Halteres yellowish orange, 
stalk paler. Abdominal tergites mostly black ; 1T and 2T except posterior margin yellowish 
brown. 

Periphallic organs : Genital arch brownish black, broad, pubescent ; anterior fragma extremely 
protruded forward ; heel lower than toe, with a few long hairs. Anal plate oblong, brown, 
hairy and pubescent. Clasper brownish black, roughly pentagonal, lower tip somewhat pointed, 
without teeth, pubescent on surface. Phallic organs: Aedeagus black, slender, basal half 
bifurcated ; ventral recurved process dark orange, elongate, distally much swollen, apically 
curved ventrally and pointed, nearly as long as aedeagus. Anterior paramere pale, broad, 
subapically with a few sensilla. Posterior paramere brown. Ventral fragma proximally 
pointed, distally narrowing. PI = oo. p.f. = ABCDefgHikLMN. 


Arun Valley, east shore of River Arun, Tumlingtar, evergreen shrubs bordering 
dry stream-beds, c. 1,800’, 14-23.xi1.1961, 3 J (R.L.C.). 

Distribution : Nepal (new locality), Formosa. 

Differs from the original form (Formosa) in having 3—5T largely black, without 
diffuse black lateral spots. 


Leucophenga guttiventris (de Meijere) 
(Text-figs. 67-70) 

Drosophila guttiventris de Meijere, 1911 : 414 (Java). 

Leucophenga guttiventris (de Meijere) Sturtevant, 1921 : 131 (Java) ; Duda, 1924a : 187, 188 ; 
1924b : 239 (Java, Formosa) ; Duda, 1923 : 28 (Java, Formosa) ; Bezzi, 1928 : 155 (Fiji) ; 
Duda, 1939 : 32; 51 (Africa) ; Okada, 1956 : 27 (Japan). 

Leucophenga guttiventris var. curvipila Duda, 1939 : 32 (Africa). 

Leucophenga guttiventris var. fuscipennis Duda, 1939 : 32 (Formosa). 

Leucophenga fuscipennis Duda, 1923 : 28 (Formosa). 

Drosophila maculiventris van der Wulp ; de Meijere, 1908 : 155 (Java). 


9. Body about 2:5 mm. Head as broad as thorax. Eyes pale reddish brown, large, bare. 
Antennae with second segment pale yellowish brown ; third greyish white, elongate. Arista 
with about six dorsal and four ventral branches and a small terminal fork. Palpus large, dark 
brownish black, basally paler, curved upward, subapically with a fine short seta. Ocellar tri- 
angle black, pollinose. Periorbit greyish white, short, flat, not well demarcated from front. 
Front white, somewhat glossy, posteriorly slightly yellowish brown, anteriorly about five- 
sevenths as broad as length down middle, posteriorly about two-fifths as broad as head width. 
Clypeus pale brown, narrow. Face white, flat. Carina undeveloped. Cheeks white, narrow, 
about one-twentieth as broad as the greatest diameter of eye. Occiput black, upper margin 
yellow. Anterior reclinate orbital slightly shorter than other orbitals, outside proclinate, thrice 
as distant from posterior reclinate as from proclinate. Only one prominent rather short oral. 

Mesonotum orange-yellow, caudomedially somewhat brownish, posteriorly slightly silver 
pollinose ; humeral callus pale. Scutellum orange-yellow, slightly glossy, laterally somewhat 
fuscous, pale at tip. Thoracic pleura pale yellowish white, without dark markings. Humerals 
two. Prescutellar as long as anterior dorsocentral. Acrostichal hairs in about eight rows. 
Anterior dorsocentrals about five-eighths as long as posteriors ; distance between anterior and 


DIPTERA FROM NEPAL 31 


posterior dorsocentrals about one-fourth distance between anterior pair. Anterior scutellars 
long, divergent ; posteriors shorter, nearly equally distant from each other as from anteriors. 
Sterno-index about 0-7. Legs yellowish white ; knees of mid and hind legs narrowly black ; 
all metatarsi nearly as long as the other segments together. Preapicals on all legs ; apicals on 
fore and mid legs. Wing hyaline, veins dark yellow, crossveins clear. R,;, somewhat convex 
anteriorly at middle, curved to costa at tip; R,,, strongly convex anteriorly before middle. 
Costa reaching tip of R,,,. C-index about 2:3 ; 4V-index about 3:3; 4C-index about 1:7 ; 
5x-index about 1-0; Ac-index about 2-7. Cr1-bristles two ; C3-fringe on basal five-eighths. 
Halteres yellow, stalk paler. Abdominal tergites orange-yellow ; 1T entirely yellow ; 2-6T 
with black spots as follows : two on 2T, 3 small on 3T, three large on 4T, three small on 5T, two 
small on 6T. 

Arun Valley, east shore of River Arun below Tumlingtar, evergreen shrubs 
bordering dry stream-beds, c. 1,800’, I4—23.xii.19g61, 5 9. Taplejung District, 
Dobhan, mixed vegetation in dry gully on wooded slope, c. 3,500’, 29.1.1962, I 9 
(R.L.C.). 

Distribution : Nepal (new locality), Java, Formosa, Japan, Fiji, Africa. 


Leucophenga nigriceps sp. n. 
(Text-figs. 76-80) 

3. 2. Body about 3mm. Head slightly broader than thorax. Eyes dark red, bare, large. 
Antennae with second segment yellow ; third yellowish grey, oblong, twice as long as broad, 
pale at the base of arista. Arista with about five dorsal and one ventral branches and a small 
terminal fork, branches rather short. Palpus slender, curved, yellow, apically black. Ocellar 
triangle black. Periorbit slightly paler than front, short, a little more than half as long as front. 
Front flat, narrow, parallel-sided, pale yellowish grey, matt, anteriorly five-sevenths as broad 
as length down middle, posteriorly one-third as broad as head width. Clypeus pale yellowish 
brown. Face yellowish white, concave. Carina undeveloped. Cheeks yellow. Very narrow, 
about one-sixteenth as broad as the greatest diameter of eye. Occiput black. Orbitals 
subequal in size ; anterior reclinate orbital close to and outside proclinate, more than four times 
as distant from posterior reclinate as from proclinate. Only one prominent oral. 

Mesonotum dark orange-brown, medioanteriorly paler, medioposteriorly fuscous. Scutellum 
black, apically pale. Thoracic pleura yellowish brown, with irregular longitudinal patches ; 
sternepisternum largely black. Humerals two, lower longer. Acrostichal hairs in about eight 
irregular rows. Anterior dorsocentrals two-thirds as long as posteriors ; distance between 
anterior and posterior pair nearly one-third distance between anterior pair. Anterior scutellars 
long, divergent ; posteriors half as long as anteriors. Sterno-index about 0-6. Legs yellow ; 
mid and hind knees dark ; front and hind metatarsi subequal to next three segments together, 
mid metatarsi longer than the other segments together. Preapicals on mid and hind legs ; 
apicals prominent on mid leg. Wing hyaline, veins black, crossveins clear. MW apically pale ; 
Roi, gently curved to costa at tip. C-index about 2:5 ; 4V-index about 1-8 ; 4C-index about 
I:I ; 5x-index about 1-3 ; Ac-indexabout1-9. Cr1-bristles two ; C3-fringe on basal three-fifths. 
Halteres entirely pale yellow. Abdominal tergites : 1T yellow, 2T black, medioanteriorly and 
laterally yellow, 3T shining silvery blue especially in male, 4T to caudal end largely black. 

Periphallic organs : Genital arch broad, black, paler below, much narrowed above, broadly 
truncate below, caudal margin with a few setae, anteroventral corner with a few shorter setae ; 
heel low, rounded. Anal plate small, oblong, pale. Clasper large, quadrate, pubescent inferior- 
ly, setigerous exteriorly, pale yellowish brown. Decasternum in shape of narrow cross bar. 
Phallic organs : Aedeagus black, bifid, sinuate in lateral aspect, with short basal apodeme ; 
ventral recurved process short, rod-like, pale, apically pointed. Anterior paramere fusiform, 
pale, with a longitudinal row of a few sensilla at middle. Posterior parameres yellow. PI = 
12:0. p.f. = ABCDeFgHikLmN. 


32 T. OKADA 


Fics. 76-85. 76-80, Leucophenga nigriceps sp. n., 3. 76, head, lateral aspect; 77, 
head, dorsal aspect ; 78, wing ; 79, periphallic organs, lateral aspect ; 80, phallic 
organs, ventral aspect. 81-85, Leucophenga rectinervis sp. n., 3. 81, wing; 82, 
head, lateral aspect ; 83, head, dorsal aspect ; 84, phallic organs, ventral aspect ; 
85, periphallic organs, lateral aspect. 


DIPTERA FROM NEPAL 33 


Holotype g. East NepaL: Taplejung District, Sangu, mixed vegetation by 
stream in gully, c. 6,200’, ix—x.1961 (R.L.C.), BMNH. 


Paratypes : 4 4, same data as holotype ; 9 dg, 2 2, Taplejung District, between 
Sangu and Tamrang, mixed shrubs in deep gorge, c. 5,200’, x—xi.I9g61 ; I g, Sangu, 
mixed vegetation by stream in gully, c. 6,200’, ix.1g61-1.1962 ; 1g, north of Sangu, 
dry grass above river bank, c. 5,000’, 5.i.1962 ; 1 g, Dobhan, evergreen trees over- 
hanging stream in deep gully, c. 3,500’, 30.i.1962 ; 2 9, between Sangu and Tamrang, 
mixed plants by damp cliff in deep river gorge, c. 5,200’, i-i1.1962 ; Ig, ibid., mossy 
ground under bushes by hill stream, c. 5,500’, 20.x.1g61 (R.L.C.), BMNH, 1 3 from 
between Sangu and Tamrang, at the Tokyo Metropolitan University. 


Allied to L. albiceps de Meijere, 1914, (Java and Formosa) but differs in having 
palpus black apically, 4-5T unspotted, and 3T silvery shining. 


Leucophenga rectinervis sp. n. 
(Text-figs. 81-85, 94) 


6,2. Body about 3mm. Head slightly broader than thorax. Eyes bright reddish brown ; 
ocelli bright red. Antennae pale yellowish white, third segment relatively slender. Arista 
with about seven dorsal and three ventral long branches and a small terminal fork. Palpus 
pale yellowish white, slender, gently curved upward, rounded at tip, with a strong apical anda 
few weaker ventral setae. Ocellar triangle black. Periorbit pale yellowish white, slightly 
glossy, obscurely demarcated from front. Front flat, pale yellowish white, matt, parallel-sided, 
anteriorly somewhat narrower than length down middle, posteriorly about one-third as broad as 
head width. Face white, laterally concave. Clypeus pale yellowish white. Cheeks pale, 
narrow, about one-twentieth as broad as the greatest diameter of eye. Occiput black. Pro- 
clinate orbital slightly shorter than other orbitals ; anterior reclinate outside and just posterior 
to proclinate. Only one prominent oral. 

Mesonotum yellowish brown, silver pollinose. Scutellum dark yellowish brown, matt, 
apically white. Thoracic pleura pale yellowish grey, with an obscurely demarcated dark cloud 
on pteropleuron. Humeral callus pale, with two humerals, lower shorter. Prescutellars long, 
parallel with each other, slightly longer than anterior dorsocentral. Acrostichal hairs in about 
ten irregular rows. Anterior dorsocentrals shorter than posteriors ; distance between 
anterior and posterior dorsocentrals about one-third distance between anterior pair. Anterior 
scutellars divergent ; posterior scutellars longer than anteriors and nearer to each other than to 
anteriors. Sterno-index about o-6. Legs yellowish white ; all metatarsi much longer than the 
other segments together. Preapicals on all legs ; apicals prominent on mid leg. Wing hyaline, 
veins brown ; crossveins clear. ,;, nearly straight, not curved to costa at tip ; costa weak 
beyond R,;; ; Ry; and M nearly parallel. Wing tip slightly pointed. C-index about 3:0 ; 
4V-index about 2:0 ; 4C-index about 1-o ; 5x-index about 1-3 ; Ac-index about 2:0. C1-bristles 
two ; C3-fringe on basal two-thirds. Halteres white. Abdominal tergites yellowish white, 
with black spots as follows : two small on 2T, three large on 4T, five large on 5T (median one 
small and longitudinal), two on 6T. 3T largely black. Caudal sternites black. 

Periphallic organs : Genital arch black, broad, not narrowing above, pubescent and setigerous, 
ventrocaudally obliquely truncate, heel low. Anal plate narrow, pale yellow, pubescent and 
setigerous. Clasper large, black, oval, pubescent and with short setae. Phallic organs: 
Aedeagus entirely bifid, black, medially somewhat broadened ; ventral recurved process pale 
brown, long, blade-like, apically pointed. Anterior paramere pale, fusiform, with a longitudinal 
row of a few sensilla on distal half. Posterior paramere pale yellow, contiguous with orange 
hooked piece of aedeagus. PI =oo. p.f. = ABCDeFgHikLmN. 


34 T. OKADA 


Holotype g. East NepaLt: Arun Valley, east shore of River Arun, below 
Tumlingtar, evergreen shrubs bordering dry stream-beds, c. 1,800’, 14-23.xii.1961 
(eL.C.),, BMENE. 


Allotype 2. Taplejung District, between Sangu and Tamrang, dense vegetation 
in tree shade by hill stream, c. 5,500’, 23.x.1961 (R.L.C.), BMNH. 


Paratypes : Ig, same data as holotype ; 14, below Sangu, mixed vegetation on 
sheltered slopes above river, c. 4,000’, 3.1.1962 ; I 9, below Sangu, edge of small 
mixed wood, c. 6,000’, 4.xi.1961 ; I 9, between Sangu and Tamrang, mixed shrubs 
in deep gorge, c. 5,200’, x-xi.Ig61 ; I 9, old mixed forest above Sangu, c. 6,200’, 
25-28.x.1961 (R.L.C.), BMNH ; 1 3g, below Sangu, deposited at the Tokyo Metro- 
politan University. 


Resembles L. flavipalpis Duda, 1939, (Uganda, Africa) in having pale palpus, 
clear wings, pale halteres, spotted abdominal tergites, and silvery pollinose mesono- 
tum, but differs from it in having nearly straight R,,, (hence the specific name) and 
larger C-index (2:5 in flavipalpis). 


Leucophenga nigroscutellata Duda 
(Text-figs. 74, 86—go) 
Leucophenga nigroscutellata Duda, 1924a : 186 ; 1924b : 237 (Formosa). 


3, 2. Body about 3 mm. Head slightly broader than thorax. Eyes reddish brown, bare. 
Antennae with second segment orange-yellow ; third greyish brown, oblong. Arista with about 
five dorsal and one ventral sparsely arranged branches and a large terminal fork. Palpus 
yellowish brown, oblong, with an apical and a few ventral setae. Ocellar triangle black, grey 
pollinose. Periorbit pale greyish brown, somewhat glossy, short, half as long as front, not 
constricted medially. Front yellowish white, parallel-sided, anteriorly paler and two-thirds as 
broad as length down middle ; posteriorly about one-third as broad as head width. Face 
whitish grey, somewhat concave. Clypeus brown. Cheeks yellowish white, narrow, about one- 
twentieth as broad as the greatest diameter of eye. Occiput dark brownish black. Anterior 
reclinate orbital four times as distant from posterior reclinate as from proclinate. Postverticals 
short. One prominent oral, others fine. 

Mesonotum dark brown, slightly glossy, not distinctly pollinose ; humeral callus pale. 
Scutellum brownish black, matt, apically pale. Thoracic pleura mostly dark brown. Three 
long humerals, median one longest, ventral one shortest. Acrostichal hairs in about eight 
irregular rows. Anterior dorsocentrals one-third as long as posteriors ; distance between 
anterior and posterior dorsocentrals about one-third distance between anterior pair. Anterior 
scutellars divergent ; posterior scutellars about two-fifths as long as anteriors, nearer to each 
other than to anteriors. Sterno-index about 0:6. Legs yellowish white ; mid and hind knees 
brown ; front metatarsi as long as next three segments together, mid metatarsi subequal to the 
succeeding segments, hind metatarsi longer than the succeeding segments. Preapicals on mid 
and hind legs ; apicalson midleg. Wing hyaline, veins pale. ,,, nearly straight, not curved 
to costa at tip ; R44, and M nearly parallel. C-index about 2:6; 4V-index about 1-8 ; 4C- 
index about I:I ; 5x-index about 1-9 ; Ac-index about 2:3. Cr1-bristles two ; C3-fringe on 
basal about two-thirds. Halteres yellowish white. Abdominal tergites mostly black, somewhat 
glossy ; IT yellow, often laterally fuscous ; 2T medially witha yellow caudally narrowing patch ; 
ultimate T white. 


DIPTERA FROM NEPAL 


Fics. 86-96. 86-90, Leucophenga nigroscutellata Duda, 3. 86, head, lateral aspect ; 87, 
head, dorsal aspect ; 88, wing; 89, periphallic organs, lateral aspect ; 90, phallic 
organs, ventral aspect. 91-93, Leucophenga albiceps de Meijere, 2. 91, head, lateral 
aspect; 92,abdominaltergites; 93, wing. 94-96, abdominal tergites of 94, Leucophenga 
vectinervis sp.n.,¢; 95, L. albifasciasp.n.,g; 96, L. leucozona Duda, g. 


35 


36 T. OKADA 


Periphallic organs : Genital arch broad, ventrally somewhat narrowing, brownish black, 
setigerous and pubescent ; heel low, rectangular. Anal plate pale yellowish brown, oblong, 
thickly setigerous. Clasper ovoid, distally rounded, brownish black, pubescent and with short, 
setae. Phallic organs : Aedeagus black, entirely bifid, sub-basally strongly sinuate ; ventral 
recurved process rod-shaped, orange ; apodeme unusually developed, long, and stout. Anterior 
paramere pale, short, fusiform, medially with a row of a few sensilla. Posterior parameres orange. 
Ventral fragma laterally pale yellow, broad, fusiform. PI = 5:0. p.f. = ABCDeFgHikLmN. 


Taplejung District, between Sangu and Tamrang, mixed shrubs in deep gorge, 
c. 5,200’, Xx-xi.1961, Ig, 1 9 ; above Sangu, evergreen scrub, c. 6,500’, 5—13.xX.1961, 
1 9. Arun Valley, east shore of River Arun below Tumlingtar, evergreen shrubs 
bordering dry stream-beds, c. 1,800’, 14—23.xil.1961, I 2 (R.L.C.). 

Distribution : Nepal (new locality), Formosa. 

Differs slightly from the original form (Formosa, Duda, 1924) in having larger 
body and sometimes entirely yellow palpus. 


Leucophenga albiceps de Meijere 


(Text-figs. 75, 91-93) 


Leucophenga albiceps de Meijere, 1914 : 258 (Java); Duda, 1924a : 190 (Java, Formosa). 
Leucophenga albiceps var. meijerea Duda, 1924a : 190 (Formosa). 
Leucophenga mevjerei Duda, 1924b : 240 [in citation]. 


2. Body about 3:2 mm. Head as broad as thorax. Eyes dark red, bare. Antennae with 
second segment yellow ; third yellowish white, quadrate, broad, apically truncate. Arista 
with about four dorsal and two ventral branches and a moderate terminal fork. Palpus yellow- 
ish white, large, medially broadened, with a few prominent ventral setae. Ocellar triangle 
black, bluish grey pollinose. Periorbit short, greyish white, well demarcated, apically curved 
inward. Front pale yellowish grey, parallel-sided, anteriorly slightly broader than median 
length down middle. Clypeus pale yellowish grey, protruded. VPostverticals short. Face 
greyish white, flat, laterally depressed, lower margin somewhat protruded. Cheeks narrow, 
whitish, about one-twelfth as broad as the greatest diameter of an eye. Posterior reclinate 
orbital longer than other orbitals ; anterior reclinate orbital outside and posterior to proclinate, 
four times as distant from posterior reclinate as from proclinate. Only one prominent oral. 

Mesonotum dark yellowish brown, somewhat silver pollinose ; neck region brown above. 
Scutellum dark yellowish brown, silver grey pollinose, apically white and broadly rounded. 
Thoracic pleura pale yellowish grey, somewhat fuscous below, with dark patches on pteropleuron. 
Humerals two, long, stout. Prescutellar longer than anterior dorsocentral. Acrostichal hairs 
in about eight rows. Anterior dorsocentrals half as long as posteriors ; distance between 
anterior and posterior dorsocentrals about one-fourth distance between anterior pair. Anterior 
scutellars divergent, longer than posteriors, which are nearer to each other than to anteriors. 
Sterno-index about 0-9. Legs yellowish white, knees pale. Front metatarsi slightly shorter 
than the succeeding segments, mid and hind metatarsi slightly longer than the succeeding 
segments. Preapicals on mid and hind legs ; apicals on all legs. Wings hyaline, veins yellow- 
ish brown. ,;, nearly straight, apically not curved to costa ; R,,,and M parallel ; R,,; only 
weakly convex near base. Posterior crossveins weakly curved inward. C-index about 2:3 ; 
4V-index about 1:8 ; 4C-index about o-g9 ; 5x-index about 1-2 ; Ac-indexabout1-6. Cr1-bristle 
two ; C3-fringe on basal five-sevenths. Halteres yellow. Abdominal tergites pale yellow, 
with black spots as follows : two small on 2T ; three large somewhat confluent on 3T ; three 
large on 4T ; five on 51 ; twoon6-7T. Anal cercus elongate, pale yellow, setigerous. 


DIPTERA FROM NEPAL 37 


Taplejung District, Sangu, mixed vegetation by stream in gully, c. 6,200’, 
ix—x.1961, I 9 ; old mixed forest above Sangu, c. 6,200’, 25-28.x.1961, 3 9 ; below 
Sangu, edge of small mixed wood, c. 6,000’, 4.xi.1961, I 9 ; below Sangu, mixed 
vegetation on sheltered slopes above river, c. 4,000’, 2.1.1962, I 9 (R.L.C.). 

Distribution : Nepal (new locality), Java, Formosa. 

Differs slightly from the original form in having less distinct dark band on the 
thoracic pleura and lower 4V-index. 


Subfamily DROSOPHILINAE 


MICRODROSOPHILA Malloch 


Microdrosophila Malloch, 1921 : 312. 
Type-species : Drosophila quadrata Sturtevant, 1916 : 341 (N. America). 


KEY TO SPECIES 


1 Third costal section with strong bristles on almost entire length. Second costal break 
deep, more than one-fourth as deep as the length of first costal section between two 
breaks. Wing somewhat pointed at tip . : pectinata (p. 37) 
Third costal section with strong bristles ending well before the apex 
2 Second costal break more than one-fourth as deep as the length of first costal section 


between two breaks. First costal section apically much swollen and black. sp. (p. 39) 
— Second costal break less than one-fifth as deep as the length of first costal section 
between two breaks. First costal section apically not much swollen and not black . 3 


3. Third costal section with strong bristles on its basal two-thirds. Arista with only three 
dorsal and two ventral branches beside terminal fork. Thoracic pleura dark only on 
humeral callus , . pauciramosa (p. 40) 

— Third costal section with strong bristles on its basal five. sixths ormore. Arista with 
about six dorsal and three ventral branches beside fork. Thoracic pleura with 


more or less distinct black longitudinal stripe. 4 
4 Palpus orange-yellow. Thoracic aor with diffuse dark brow longitudinal stripe. 

Halteres yellow . ; marginata (p. 42) 
— Palpus black. Thoracic pleura with a wel demarcated black longitudinal stripe. 

Halteres black or brownish black . : : 5 


5 Mesonotum with three distinct black longitudinal stripes. Ocellar triangle black 
nigripalpis (p. 41) 
— Mesonotum with three diffuse black stripes or uniformly dark brown. Ocellar 
triangle yellow . : é ‘ : : : ‘ nigrohalterata (p. 41) 


Microdrosophila pectinata sp. n. 
(Text-figs. 97-100) 


3. Body about 1-5 mm. Head slightly broader than thorax. Eyes dark red, pilose. 
Antennae with second segment dark yellowish brown, large, conical ; third narrower, yellowish 
brown, with long pubescence. Arista with about five dorsal and one ventral branches and a 
moderate terminal fork. Palpus pale yellowish brown, with a few stout black apical setae. 
Ocellar triangle pale yellowish grey, black within ocelli. Periorbit rather narrow, somewhat 
convex, anteriorly broadened, extending near anterior margin of front. Front pale yellowish 
grey, slightly glossy, mediolaterally fuscous, anteriorly two and a half times as broad as length 
down middle, posteriorly about three-fifths as broad as head width. Clypeus yellowish white. 


38 T. OKADA 


107 


103 106 


Fics. 97-107. 97-100, Microdrosophila pectinata sp. n., 3. 97, head, lateral aspect ; 98, 
head, dorsal aspect ; 99, wing; 100, periphallic organs, lateral aspect. 101-103, 
Microdrosophila paucivamosa sp. n., 9. 101, head, lateral aspect ; 102, head, dorsal 
aspect ; 103, wing. 104-107, Microdrosophila nigripalpis sp. n., 2. 104, head, 
lateral aspect ; 105, head, dorsal aspect ; 106, wing; 107, egg-guide, lateral aspect. 


DIPTERA FROM NEPAL 39 


Face pale yellowish white, somewhat narrowing below. Carina high, long, yellowish white, 
reaching buccal margin. Cheeks pale yellowish white, slightly less than one-fourth as broad as 
the greatest diameter of an eye. Occiput pale yellowish grey. Proclinate orbital slightly 
inside posterior reclinate ; anterior reclinate minute, not distinguishable from adjacent micro- 
trichia, outside other orbitals, almost equally distant from proclinate and posterior reclinate. 
Vibrissae strong, second oral slender. 

Mesonotum yellowish brown, matt. Scutellum pale yellowish grey, matt, slightly pollinose. 
Thoracic pleura pale yellowish white, somewhat fuscous above. Only one humeral. Acro- 
stichal hairs in about six rows. Distance between anterior and posterior dorsocentrals about 
five-sevenths distance between anterior pair. Anterior scutellars divergent, small, one-third 
length of posteriors, which are somewhat divergent and nearly equidistant from each other and 
from anteriors. Posterior sternopleural long, others missing. Legs yellow, anterior femur 
with a few long hairs below ; claws long ; all metatarsi subequal to the next three segments 
together ; last tarsal segment large. Preapicals on all legs ; apicals prominent on mid legs. 
Wing hyaline, second costal incision very deep ; tip pointed. R,,, apically curved to costa ; 
Ry; and M medially somewhat convergent. C-index about 1-1 ; 4V-index about 3:6; 4C- 
index about 3:3 ; 5x-index about 5:7 ; Ac-index about 3-6. Cr1-bristles two ; C3-fringe on 
nearly entire length. Halteres yellowish brown. Abdominal tergites matt black ; 1T pale 
yellowish grey, with dark patches ; 2T yellowish white laterally. Abdominal sternites pale. 

Periphallic organs: Pregenital lobe long, club-shaped, pale orange. Genital arch pale 
yellowish brown, nearly bare, ventrally pointed ; heel very high. Clasper apparently fused to 
genital arch, crescentic, pale brown. Anal plate pale brown, setigerous, narrow but long, 
ventrally curved anteriorly. Phallic organs not examined. 


Holotype g. East NepaL: Taplejung District, Dobhan, east bank of River 
Tamur, mixed vegetation by stream in deep gully, c. 3,500’, i-ti.1962 (R.L.C.), 
BMNH. 


Resembling M. tectifrons (de Meijere, 1914) (Java) in having pointed wings and 
only one ventral branch of arista, but differs in the paler body, R,,, not straight, 
and carina well developed. Belonging to the subgenus Oxystyloptera Duda, 1924. 


Microdrosophila sp. 


rex. Head much broader than thorax. Eyes green-black, thickly grey pilose. Antennae 
with second segment globular, yellowish brown, paler below ; third dark brown, oval, paler 
caudally. Arista with four dorsal and three ventral branches and a large terminal fork. Palpus 
yellowish brown, subapically with two strong black setae. Ocellar triangle glossy black, convex. 
Periorbit glossy grey, broad, medially somewhat constricted, anteriorly truncate, four-fifths as 
long as front. Front greyish yellow, anteriorly paler, black inside periorbits, with a pair of 
large triangular anteriorly pointed black patches. Clypeus pale brown, medially with a dark 
narrow cross stripe. Face yellow, very broad, parallel-sided. Carina large, high, pale. Cheek 
greyish white, three-eighths as broad as the greatest diameter of eye. Occiput yellowish grey. 
Anterior reclinate orbital minute, just outside proclinate, nearer to proclinate than to posterior 
reclinate, which is longer than anterior reclinate. Second oral two-thirds as long as vibrissa, 
succeeding orals slender. 

Mesonotum greyish yellow, matt, with five narrow dark brown longitudinal stripes, median 
one broadest. Scutellum greyish yellow, with a pair of broad dark longitudinal stripes. Thoracic 
pleura pale yellowish grey, with two dark longitudinal stripes. One long humeral. Acro- 
stichal hairs in six rows. Distance between anterior pair of dorsocentrals equal to distance 
between anterior and posterior pairs. Anterior dorsocentrals two-thirds as long as posteriors. 
Anterior scutellars divergent, as long as posteriors, which are nearer to each other than to 
anteriors. Sterno-index about 0-8. Legs yellow, fore and hind metatarsi subequal to next 


40 T. OKADA 


three segments together, mid metatarsi subequal to the following segments together. Pre- 
apicals on all legs ; apicals prominent on mid leg. Wing hyaline, 2-1 mm. in length ; first 
costal section apically much swollen and black ; second costal break deep. R,+, straight. 
Wing-tip somewhat pointed. R,,, and M parallel. C-index about 1:8 ; 4V-index about 1:9 ; 
4C-index about 1-3 ; 5x-index about 2:0 ; Ac-index about 3:3. Cr1-bristles two ; C3-fringe 
on basal half. Halteres missing. Abdomen missing. 


Taplejung District, between Sangu and Tamrang, mixed shrubs in deep gorge, 
C. 5,200", X xi. 4Q01, T ex (i 12-C,)- 

Characters intermediate between Styloptera Duda, 1924 and Miucrodrosophila 
Malloch ; first costal section apically swollen and black as in Styloptera, acrostichal 
hairs in six rows as in Muicrodrosopmla. A deep incision at the second costal 
break suggests that this species belongs to the subgenus Oxystyloptera Duda. 


Microdrosophila pauciramosa sp. n. 
(Text-figs. 10I—103) 


2. Body about 1-5 mm., yellowish brown. Head broader than thorax. Eyes deep purple- 
red, oval, pilose, with caudal margin straight. Antennae dark yellowish brown, third segment 
quadrate, slightly longer than broad. Arista with only three dorsal and two ventral sparsely 
arranged branches and a large terminal fork. Palpus pale yellow, broad, with two stout bristles 
below near apex. Ocellar triangle black ; ocelli pale. Periorbit large, somewhat convex, 
pale yellowish brown, anteriorly broadened and reaching anterior margin of front. Front pale 
yellowish grey, medio-anteriorly orange, anteriorly twice as broad as length down middle, 
posteriorly somewhat wider and about five-ninths as broad as head width. Clypeus dark brown. 
Face yellowish white, slightly narrowing below, buccal margin white. Carina high, long, 
reaching near buccal margin. Cheek pale yellowish white, darker above, about one-third 
as broad as the greatest diameter of eye. Occiput pale yellow, darker below. Proclinate 
orbital slightly shorter than posterior reclinate, close to and outside posterior reclinate ; anterior 
reclinate minute, just outside posterior reclinate. 

Mesonotum dirty yellowish brown, slightly glossy, with three indistinct longitudinal dark 
patches. Scutellum dirty yellowish brown. Thoracic pleura pale yellow, darker above. 
Humeral callus dark brown. Only one humeral. Acrostichal hairs in about eight rows. 
Distance between anterior pair of dorsocentrals about one and a half times distance between 
anterior and posterior pairs. Anterior scutellars slightly shorter than posteriors, divergent ; 
posterior scutellars as long as posterior dorsocentrals. Sterno-index about 0-4. Legs pale 
yellowish brown ; front metatarsi subequal in length to next three segments together, mid and 
hind metatarsi subequal to the rest of segments. Fore femur and mid coxa with long hairs 
below. Preapicals on all legs ; apicals prominent on mid leg. Wings hyaline ; crossveins 
clear ; first costal section distally curved posteriorly and swollen ; costa reaching M; Roi, 
subapically gently curved to costa ; R,,; and M parallel. C-index about 1-9 ; 4V-index about 
5:2; 4C-index about 2:7 ; 5x-index about 7:5 ; Ac-index about 4:3. Cz1-bristles two ; C3- 
fringe on basal two-thirds. Halteres greyish brown. Abdominal tergites dirty yellowish brown, 
laterally darker, with irregular dark brown clouds, posterior margin narrowly grey. Abdominal 
sternites pale. Cerci broader than long. 


Holotype 9. East NepaL: Taplejung District, Sangu, mixed vegetation by 
stream in gully, c. 6,200’, xi.1g61 — 1.1962 (R.L.C.), BMNH. 

Somewhat resembles M. cristata Okada, 1960 in having obscurely demarcated 
mesopleural dark stripe, gently curved R,,5, and orbitals close to each other, but 


differs from it in having fewer aristal branches, posterior reclinate orbital not inside 
proclinate and lesser range of C3-fringe. 


DIPTERA FROM NEPAL 41 


Microdrosophila nigripalpis sp. n. 
(Text-figs. 104-107) 


9. Body about 2 mm., dark yellowish brown. Head slightly broader than thorax. Eyes 
deep purplish black, thickly pilose. Antennae dark greyish brown ; second segment large, 
oval; third triangular, short, with long pubescence. Arista with seven dorsal and three 
ventral long branches and a moderate terminal fork. Palpus black, slender, apically rounded, 
with a few stout black setae below near apex. Ocellar triangle black ; ocelli pale yellow. 
Periorbits quite large, anteriorly broadened, pale yellowish white, convex, reaching anterior 
margin of front. Front short, anteriorly nearly four times as broad as length down middle, 
posteriorly more than half as broad as head width, dark orange-brown, caudally black. Clypeus 
pale yellow. Face pale, laterally concave. Carina pale yellow, large, long, evenly convex, 
extending near buccal margin. Cheeks yellowish white, black above, about one-fourth as broad 
as the greatest diameter of eye, narrow below. Occiput pale yellow, black below. Three 
orbitals close to each other ; proclinate orbital about half as long as posterior reclinate, just 
before and outside posterior reclinate ; anterior reclinate minute, slightly outside proclinate. 
Only one stout oral. 

Mesonotum yellowish brown, with three broad black longitudinal stripes ; median stripe 
diffuse, reaching apex of scutellum ; lateral stripes well demarcated, not extending in front of 
suture. Scutellum laterally yellowish brown, medially dark. Thoracic pleura pale yellow 
below, with broad black longitudinal stripe above. One humeral. Acrostichal hairs in about 
eight rows. Distance between anterior and posterior dorsocentrals about three-fourths distance 
between anterior pair ; anterior dorsocentrals slightly shorter than posteriors, situated near 
thesuture. Anterior scutellars missing ; posteriors slightly divergent, twice length of scutellum. 
Sterno-index about 0-3. Legs yellowish grey ; mid and hind pairs with distal half of femur and 
proximal part of tibia brownish black ; all metatarsi nearly as long as the other segments 
together. Preapicals on all legs ; apicals prominent on mid leg. Wings hyaline, tip pointed ; 
veins brown, crossveins clear ; costa reaching M ; second costal break deep. R.4, nearly 
straight, weakly sinuated ; R,,;, and M parallel. C-index about 1-3 ; 4V-index about 4:1 ; 
4C-index about 2-7 ; 5x-index about 6-0 ; Ac-index about 4-0. Cr1-bristles two ; C3-fringe on 
basal six-sevenths. Halteres greyish black. Abdominal tergites matt black, posterior margins 
narrowly pale ; xT laterally pale. Abdominal sternites pale yellow. Egg-guide triangular, 
dark brown, apically with a very long and two shorter setae. 


Holotype 9. East Nepat ; Taplejung District, old mixed forest above Sangu, 
c. 6,200’, 25-28.x.1961 (R.L.C.), BMNH. 


Allied to M. purpurata Okada, 1956, (Japan) in having black mesonotal and meso- 
pleural longitudinal stripes and triangular egg-guides, but differs from it principally 
in having palpus and halteres black and pointed wings. 


Microdrosophila nigrohalterata sp. n. 
(Text-figs. 114-118) 


3,9. Body about 2:5 mm., dark yellowish brown. Head broader than thorax. Eyes dark 
purplish red, somewhat quadrate, weakly sinuous at inner margin, pilose. Antennae thick, 
second segment dark yellowish brown, with two long stout setae ; third rounded at tip, greyish 
brown, with long pubescence. Arista with about five dorsal and three ventral long branches and 
a moderate terminal fork. Palpus dark brown or black, slender, with a few stout spical setae. 
Ocellar triangle highly convex, pale yellowish brown ; mouth-parts yellowish white. Periorbit 
large, highly convex, glossy greyish brown, anteriorly broadened, reaching near the anterior 
margin of front. Clypeus yellowish white. Front glossy greyish brown, highly convex, 


42 T. OKADA 


anteriorly and laterally orange-brown ; anteriorly two and a half times as broad as length 
down middle ; posteriorly about half as broad as head width. Face pale yellowish white, 
laterally depressed. Carina high, slender, two-thirds as long as face. Occiput yellowish grey, 
black below. Cheek pale yellowish black near neck region, about one-third as broad as the 
greatest diameter of eye, narrow below. Proclinate orbital two-thirds length of posterior 
reclinate, outside and slightly anterior to posterior reclinate ; anterior reclinate minute, just 
outside posteriors, twice as distant from proclinate as from posterior reclinate. Only one very 
stout and long oral. 

Mesonotum glossy yellowish brown, mediocaudally and laterally dark brown, or uniformly 
dark brown. Scutellum dark yellowish brown, glossy, often medially and marginally pale. 
Thoracic pleura yellowish white, with broad black longitudinal stripe above. Humerals two, 
very long, stout, subequal in length. Acrostrichal hairs in six rows. Anterior dorsocentrals 
near the level of suture ; distance between anterior and posterior dorsocentrals subequal to 
distance between anterior pair. Anterior scutellars short ; posteriors long, divergent, thrice as 
long as anteriors. Sterno-index about o:5. Legs fuscous yellow, femora pale on proximal half. 
Preapicals on all legs ; apicals prominent on mid leg. Wings hyaline, crossveins clear, tip 
slightly pointed, costa reaching M. R,4, apically curved to costa; R,,;; and M parallel. 
C-index about 1-3 ; 4V-index about 3-6 ; 4C-index about 2:2 ; 5x-index about 5:0 ; Ac-index 
about 4:4. Cz1-bristles two, long ; C3-fringe on basal ten-elevenths. Halteres greyish black. 
Abdominal tergites mostly black, glossy ; abdominal sternites yellowish white. 

Periphallic organs : Genital arch yellow, broad, narrowing below, pubescent, setigerous on 
upper caudal margin, ventral apex pointed anteriorly, caudal lower margin orange, rounded. 
Clasper and anal plate obscure. Decasternum triangular, yellowish white. Phallic organs : 
Aedeagus robust, orange, with several complicated triangular processes, basally narrowing and 
with a stout recurved dark orange ventral process, which is slightly shorter than aedeagus (cf. 
Text-fig. 117). Apodeme of aedeagus thick but short, about one-third as long as aedeagus. 
PI = 3:0. Anterior paramere seemingly fused to hypandrium and with about two sensilla. 
Posterior paramere elongate, connecting hypandrium and subbasal part of aedeagus (cf. Text- 
fig. 117). Ventral fragma triangularly pointed proximally ; hypandrial plate large, blade- 
like. p.f. = abCdeFgHiKlmn. Egg-guide yellow, oval, distally fuscous and with four stout 
brownish setae and a small hair. 


Holotype gj. East NEpaL: Taplejung District, between Sangu and Tamrang. 
mixed plants by damp cliff in deep river gorge, c. 5,200’, i-ti.1962 (R.L.C.), BMNH, 

Allotype 9. Taplejung District, above Sangu, mixed vegetation in dried-up 
ravine, C..0,800), 17.11.1962 (K.0.-C.) BINH. 

Paratypes : Taplejung District, Sangu, mixed vegetation by stream in gully, 
c. 6,200’, xi.1961-1.1962, 1 9 (R.L.C.), BMNH. Arun Valley, east shore of River 
Arun below Tumlingtar, evergreen shrubs bordering dry stream-beds, c. 1,800’, 
14—23.x11.1961, I 9 (R.L.C.), deposited at the Tokyo Metropolitan University. 

Closely resembles M. nigripalpis Okada, in having distinct broad black meso- 
pleural longitudinal stripe, black palpus, and black halteres, but differs from its 
allies in having less distinct mesonotal black stripes, pale ocellar triangle, and oval 
egg-guide. 


Microdrosophila marginata sp. n. 
(Text-figs. 108-113) 


36, °. Body about 1-3 mm., pale yellowish brown. Head very large, much broader than 
thorax. Eyes deep purplish red, large, thickly pilose. Antennae with second segment dark 


DIPTERA FROM NEPAL 43 


yellowish brown, large, globular ; third as long as second, much narrower and paler. Arista 
with about seven dorsal and three ventral very long branches and a small terminal fork. Palpus 
orange-yellow, small, apically rounded and with a stout black and a few thinner ventral setae. 
Ocellar triangle medially black. Frontal triangle large, convex. Periorbit large, convex, 
anteriorly much broadened, pale yellowish grey, somewhat glossy, reaching anterior margin of 
front. Front pale yellowish grey, somewhat glossy, anteriorly two-thirds as broad as length 
down middle, posteriorly slightly more than half as broad as head width. Face pale yellow, 
laterally concave. Carina high, long, pale yellowish white, wider below, reaching near buccal 
margin. Clypeus pale yellow. Cheeks narrow below, yellowish white, about one-fifth as broad 
as the greatest diameter of eye. Occiput pale yellow. Anterior reclinate orbital minute, just 
behind proclinate ; proclinate about two-thirds as long as posterior reclinate, before and out- 
side posterior reclinate. Only one long stout oral, others fine. 


Mesonotum yellowish brown, caudomedially and caudolaterally with indistinct dark brown 
patches. Scutellum yellowish brown, slightly darker medially. Thoracic pleura pale yellowish 
white, diffuse dark brown above. Only one stout humeral. Acrostichal hairs in six somewhat 
irregular rows. Anterior dorsocentrals at the level of suture ; distance between anterior pair 
of dorsocentrals about four-fifths distance between anterior and posterior pairs. Anterior 
scutellars small, parallel, less than half as long as posteriors, which are divergent. Sterno-index 
about 0-5. Legs yellowish grey ; all metatarsi subequal to the next three segments together. 
Preapicals on all legs ; apicals prominent on mid leg. Wings hyaline, pointed at tip (slightly 
below end of R,4,;) ; crossveins clear. R,;, gently curved to costa at tip ; R,,, and M slightly 
divergent. C-index about 1-7; 4V-index about 5:0 ; 4C-index about 3-0 ; 5x-index about 
5°3; Ac-index about 3:0. Cr1-bristles two ; C3-fringe on basal five-sixths. Halteres yellow. 
Abdominal tergites yellowish grey, laterally and caudally dark brown. Abdominal sternites 
pale. 


Periphallic organs : Genital arch pale yellow, broad, fusiform, pubescent, abruptly narrowing 
above, ventroanteriorly pointed downward, caudoventral margin setigerous. Anal plate pale 
yellow, oblong, setigerous, lower tip rounded. Clasper apparently absent. Phallic organs : 
Aedeagus large, pale orange-brown, fusiform in lateral view, with a narrow anterior projection 
subbasally, basal half rod-shaped, contiguous with a large distally truncate ventral recurved 
process, which is nearly half as long as aedeagus. Apodeme of aedeagus short, rod-like. PI = 
6-0. Anterior paramere elongate, pale yellow, proximally narrowing, subapically with a row 
of a few sensilla. Posterior parameres fused to each other and to ventral fragma. Ventral 
fragma like a narrow bow. p.f. = aBCdeFgHikLmn. Egg-guide pale yellowish brown, short, 
triangular, apically pointed, with a long apical and two shorter submedian stout setae and 
a fine subapical hair. 


Holotype g. East NepaL: Taplejung District, Dobhan, small pockets of plants 
on arid slopes above River Maewa, c. 3,500’, 2.1.1962 (R.L.C.), BMNH. 


Allotype 9. Taplejung District, Sangu, mixed vegetation by stream in gully, 
c. 6,200’, xi.1961-1.1962 (R.L.C.), BMNH. 


Paratype : I gf, same data as allotype, BMNH. 


Resembles M. maculata Okada, 1960, (Japan) in having dark second antennal 
segment, yellowish abdomen laterally dark, pale palpus, and large fusiform aedeagus, 
but differs from it in having pointed wings, entirely black ocellar triangle, entirely 
yellow halteres (knob darker in maculata), and triangular egg-guide (quadrate in 
maculata). 


44 


T. OKADA 


115 


Fics. 108-118. 108-113, Micvodvosophila marginata sp. n. 108, g head, lateral aspect ; 


109, g head, dorsalaspect ; 110,f wing; 111, periphallic organs, lateralaspect ; 112, 
phallic organs, ventral aspect; 113, 2 egg-guide, lateral aspect. 114-118, Muicro- 
drosophila nigrohaltevata sp.n. 114, head, lateralaspect ; 115,g wing ; 116, periphallic 
organs, lateral aspect ; 117, phallic organs, ventral aspect ; left upper figure, tip of 
aedeagus, lateral aspect ; 118, 9 egg-guide, lateral aspect. 


DIPTERA FROM NEPAL 45 


LISSODROSOPHILA gen. n. 
Type-species, Lissodrosophila longicorms sp.n. (Nepal). 


Eyes vertical. Arista without ventral branches below fork, dorsal branches long. Front 
broad, glossy, smooth. Periorbit very broad, reaching anterior margin of front. Orbitals two, 
both proclinate, without reclinate orbitals. Postverticals minute. Mesonotum very convex, 
glossy. Scutellum matt, short. Two long sternopleurals, one humeral, one supra-alar. Acro- 
stichal hairs in two rows. Only one pair of dorsocentrals. No prescutellars. Alula and anal 
vein abbreviated. 


Related to the genus Lissocephala Malloch, 1929, in having a smooth, glossy, and 
flat front, small postverticals, shining convex mesonotum, dull scutellum, no anal 
vein, and no row of spinules on fore femur. Differs from Lissocephala in having no 
reclinate orbitals, no ventral branches of arista, only one pair of supra-alars, two rows 
of acrostichal hairs, and only one pair of dorsocentrals. 


Lissodrosophila longicornis sp. n. 
(Text-figs. 119-122) 


6. Body about 1:5 mm. Head dorsoventrally elongated, much broader than thorax, 
anteriorly protruded at antennal base. Eyes vertically elongate, ventrally narrowing, deep 
purplish brown, bare. Antennae very large and long ; first segment black, longer than broad ; 
second black, distally broadened, rectangularly articulated to the first ; third matt black, with 
grey pubescence, about four times as long as broad, distally tapering. Arista with about seven 
dorsal long branches and a small terminal fork. Palpus club-shaped, yellow, subapically with a 
few black setae. Ocellar triangle dark brown, glossy. Periorbit very broad, reaching anterior 
margin of front, glossy deep brown. Postverticals minute. Front glossy deep brown, smooth, 
anteriorly one-third broader than length down middle, posteriorly more than half as broad as 
head width. A few frontal hairs present. Clypeus dark brown. Face narrow, deep glossy 
brown, buccal margin pale and much protruded upward. Carina absent. Cheeks narrow, 
yellowish brown, pale at eye margin, about one-ninth as broad as the greatest diameter of eye, 
black above. Occiput deep brownish black. Mouth-parts deep brown, robust and long. 
Anterior proclinate orbital long and upright, posteriors minute. A few short orals ; vibrissa 
somewhat longer than other orals. 

Mesonotum black, shining blue-green, convex, with thick grey pubescence. Scutellum velvety 
black, much shorter than broad, posteriorly grey pollinose. Thoracic pleura dark brownish 
black, glossy, somewhat blue-green. Dorsocentrals near the caudal end of scutum. Anterior 
scutellars divergent, about two-thirds as long as posteriors ; posteriors longer than dorsocentral, 
parallel, twice as far from each other as from anteriors. A small marginal hair between anterior 
and posterior scutellars. Sterno-index about 0-7. Legs mostly brownish black ; femora 
darker ; all metatarsi much longer than the other segments together. Preapicals on all legs ; 
apicals prominent on mid leg. Wings hyaline, tip rounded, veins yellow. Alula black, small, 
without fringe. ,,, strongly curved to costa at tip ; R,,,and M parallel ; posterior crossvein 
curved outward ; anal vein absent. C-index about 1:8 ; 4V-index about 2.0 ; 4C-index about 
I*3; 5x-index about 1:2; Ac-index about 3:1. Cz1-bristles two ; C3-fringe on basal half, 
sparse. Halteres white, stalk black. Abdominal tergites glossy black, shining metallic green, 
broader and shorter than thorax. Genital arch narrow, setigerous ; clasper oval, setigerous 
and pubescent ; anal plate elliptical, setigerous and pubescent. Phallic organs not examined. 


Holotype g. East NepaL: Taplejung District, below Sangu, by stream in 
shady ravine, c. 6,000’, 30.x.19g61 (R.L.C.), BMNH. 


46 


T. OKADA 


Fics. 119-126. 119-122, Lissodrosophila longicornis gen. & sp. n., 2. 119, head, lateral 
I2I, wing ; 122, abdomen, caudal aspect. 123- 


aspect ; 120, head, dorsal aspect ; 
126, Hypselothyrea guttata Duda, 9. 
125, wing; 126, egg-guide. 


123, head, lateral aspect ; 


124, head, dorsal aspect ; 


DIPTERA FROM NEPAL 47 


HYPSELOTHYREA de Meijere 
Hypselothyrea de Meijere, 1906 : 193. 
Type-species : H. dimidiata de Meijere, 1906 : 193 (New Guinea). 


Hypselothyrea guttata Duda 
(Text-figs. 123-126) 
Hypselothyvea guttata Duda, 1926 : 56 (Sumatra) ; Duda, 1928 : 86 (Formosa). 


9. Body about 1-6 mm., glossy purplish brown, with ornamented wing. Head somewhat 
broader than thorax. Eyes dark red, widely separated, with fine pilosity. Antennae with 
second segment dark brown, conical ; third broader, tip rounded, yellowish brown and anteriorly 
black. Palpus black, slender, basally narrowing, with about two stout apical and subapical 
setae. Ocellar triangle black. Periorbit glossy dark brown, anteriorly narrowing, posteriorly 
much broadened, two-thirds as long as front. Postverticals absent. Front brownish black, 
glossy, somewhat purplish, quadrate, as broad as long. Clypeus black or dark brown. Face 
yellowish brown, highly convex below. Carina yellowish orange or dark brown, high, short, less 
than half as long as face. Cheeks dark brown, paler above, about two-ninths as broad as the 
greatest diameter of eye, narrow below. Two long orbitals, no posterior reclinate ; proclinate 
orbital slightly shorter than reclinate ; both orbitals equidistant from eye margin. Vibrissae 
thin, long ; succeeding orals short. 

Mesonotum convex, yellowish or dark reddish brown, glossy, posteriorly darker. Scutellum 
triangular, tip protruded dorsoposteriorly, black, marginally white. Thoracic pleura pale 
yellowish grey, with a black longitudinal stripe above. One humeral. Acrostichal hairs in six 
rows. Posterior dorsocentrals slightly longer than anteriors ; distance between anterior and 
posterior dorsocentrals four-sevenths distance between anterior pair. Anterior scutellars 
parallel, half as long as posteriors, which are twice as distant from anteriors as from each other. 
Only one stenopleural. Legs pale yellowish grey ; fore femur basally black ; fore and mid 
coxae black ; hind coxa white ; mid and hind femora subapically black. Fore femur with 
anteroventrally a row of about ten small black teeth. Front metatarsi somewhat shorter than 
next three segments together, mid and hind metatarsi as long as the rest of segments together. 
Preapicals on all legs ; apicals prominent on mid leg. Wings broad, with scattered large black 
patches roughly in six transverse rows. Costa irregularly sinuous, alternatively white and black. 
R.+, gently curved to costa apically ; R,,, and M slightly divergent distally ; posterior cross- 
vein curved outward. C-index about 1-3 ; 4V-index about 4:0; 4C-index about 4-1 ; 5x- 
index about 1-3 ; Ac-index about 3-8. Cr1-bristles one, small ; C3-fringe on basal two-thirds. 
Halteres black. Abdomen narrowing anteriorly ; abdominal tergites mostly glossy black, 
mediolongitudinally yellowish brown ; abdominal sternites reddish brown. 

Egg-guide lobe slender, pale yellow, basally brownish, tip gently narrowing, with about fifteen 
marginal thick yellow teeth and four discal longer but thinner yellow teeth. Basal isthmus 
narrow, brown. 


Taplejung District, Sangu, rotting fruits of Bhor tree on ground, c. 6,200’, 
7-31.x.1961, 1 9 ; below Sangu, by stream in shady ravine, c. 6,000’, 30.x.1961, I 9 
WeseC.): 


This species has some features characteristic of the genus Liodrosophila Duda : 
an anteroventral row of small bristles on fore femur, smooth glossy front, and narrow 
triangular egg-guide. It has, however, no postverticals, no anterior reclinate 
orbital, and only one sternopleural, characteristics of the genus Hypselothyrea. 


Distribution : Nepal (new locality), Sumatra, Formosa. 


48 TOGA yx 


LIODROSOPHILA Duda 
Liodrosophila Duda, 1922 : 153. 
Type-species : Camilla coeruleifrons de Meijere, Ig1I : 421 (Java). 


Key TO SPECIES 
1 Wings with black markings. Front and mesonotum granulose, moderately shining 


black. Legs yellow, fore tarsi black : : ; bimaculata (p. 48) 
— Wings without black markings. Front and mesonotum emoeeh, glossy, not granulose. 
Fore tarsi yellow or yellowish grey, not black . ; : 2 


2 Acrostichal hairs in two rows. Mesonotum somewhat Shae bles, puiicbaced 
onchopyga (p. 50) 
— Acrostichal hairs in six rows. Mesonotum glossy, not punctured . 3 
3 Mesonotum usually brownish black. Clasper with a concave row of eee teeth. 
Wings somewhat fuscous. Periorbit relatively broad, not narrowing anteriorly 
Suscipennis (p. 51) 
— Mesonotum enamel-blue-black. Clasper with a convex row of marginal teeth. Wings 
hyaline. Periorbit much narrowing anteriorly : : : : nitida (p. 54) 


Liodrosophila bimaculata sp. n. 
(Text-figs. 127-133) 


3,2. Body about 2-5 mm. Head slightly broader than thorax. Eyes dark reddish brown 
with thick long grey pilosity. Antennae with second segment black ; third orange-brown, 
twice as long as broad. Arista with about six dorsal and two ventral long branches and a small 
terminalfork. Palpus black, slender, with a very long apical black seta and a few shorter ventral 
setae. Ocellar triangle bluish black, flat, not sharply demarcated. Mouth-parts black, thick. 
Periorbit bluish black, moderately shining, granulose, somewhat convex, three-fourths as long as 
front. Front moderately shining black, slightly bluish, flat, quadrate, granulose. Face 
moderately shining black, slightly bluish, not depressed ; buccal margin with a few transverse 
fine linear grooves. Carina high, black, broader below, ventrally reaching three-fifths length of 
face, flat above. Cheek moderately shining black, one-fifth as broad as the greatest diameter of 
eye. Occiput black. Postverticals slender. Anterior reclinate orbital minute, outside and 
behind proclinate, thrice as distant from posterior reclinate as from proclinate ; proclinate two- 
thirds as long as posterior reclinate. Vibrissae long, stout, succeeding orals short. 

Mesonotum moderately shining black, granulose. Scutellum velvety black, somewhat 
pollinose at anterolateral corners. Thoracic pleura moderately shining black, slightly bluish. 
Two long humerals, lower longer. Acrostichal hairs in six sparse and somewhat irregular rows. 
Anterior dorsocentrals about three-fourths as long as posteriors ; distance between anterior and 
posterior dorsocentrals four-sevenths distance between anterior pair. Anterior scutellars diver- 
gent, four-ninths as long as posteriors, which are nearly parallel and subequally distant from 
each other and from anteriors. Sterno-index about 0-3. Legs yellow, fore tarsi and basal half 
of fore femur black ; tips of mid and hind femora somewhat dark. Fore femur with a row of 
spinules anteroventrally on distal half and a row of long hairs ventrally on distal half. Front 
metatarsi as long as next three segments together, mid and hind metatarsi subequal to the other 
segments together. Preapicals on all legs ; apicals on mid leg. Wings with two large black 
spots, one triangular, basal to posterior crossvein, and the other round, large, outside posterior 
crossvein ; wing tip slightly pointed. #,,3; weakly curved to costa at tip ; R,,, and M nearly 
parallel. C-index about 1-8 ; 4V-index about 3-3 ; 4C-index about 2-3 ; 5x-index about 2:1 ; 
Ac-index about 3:8. Ci1-bristle one ; C3-fringe on basal five-sevenths. Halteres greyish brown, 
basally white, outer margin black. Abdominal tergites elongate, glossy black, somewhat 
purplish, slightly grey pollinose. 

Periphallic organs dark brownish black. Genital arch slightly broader below, ventrally 
truncate obliquely, sparsely setigerous ; heel low, rounded. Anal plate fusiform, setigerous, 


DIPTERA FROM NEPAL 


136 


Fics. 127-139. 127-133, Liodvosophila bimaculatasp.n. 127, g head, lateralaspect ; 128, 
6g head, dorsal aspect ; 129, wing; 130, Qegg-guide ; 131, periphallic organs, lateral 
aspect ; 132, ejaculatory apodeme ; 133, phallic organs, ventral aspect. 134-139, 
Liodrosophila onchopyga sp.n. 134, 3 head, lateral aspect ; 135, d head, dorsal aspect ; 
136, d wing ; 137, 9 egg-guide ; 138, periphallic organs, lateral aspect ; 139, phallic 
organs, ventral aspect. 


49 


50 T. OKADA 


caudal tip somewhat elongated below. Clasper elliptical, with about three rows of black teeth 
along outer margin. Decasternum orange, small, oval. Phallic organs: Aedeagus black, 
medially paler, rhombic in ventral aspect, basally with a pair of short triangular vertical rods ; 
apodeme slender, black, nearly twice as long as aedeagus. Anterior paramere fused to hypan- 
drium, without prominent sensilla. Hypandrial plate crescentic, brown, inferiorly with a stout 
short submedian spine. Ventral fragma large, ovoid, dark brown. Ejaculatory apodeme with 
black stalk and dark brown triangular plate. PI = 0-8. p.f. = aBCdef,g,HIklmN. Egg- 
guide lobe pale orange, triangularly pointed apically, basally tapering and black, with about 
eighteen marginal and five discal orange teeth, subapically with a rather stout yellow hair 
above and below. Basal isthmus short, black. 


Holotype g. East NEpaL: Taplejung District, below Sangu, by stream in 
shady ravine, c. 6,000’, 30.x.19g61 (R.L.C.), BMNH. 


Allotype 2. Same data as holotype (R.L.C.), BMNH. 


Paratypes : 2 2, same data as holotype ; 1 9, Taplejung District, Sangu, mixed 
vegetation by stream in gully, c. 6,200’, xi.1961-i.1962 (R.L.C.), BMNH (2 9), and 
the Tokyo Metropolitan University (1 9, from Sangu). 


Easily distinguished from other members of the genus Liodrosopluila in having two 
large black markings on wings, front and mesonotum granulose. 


Liodrosophila onchopyga sp. n. 
(Text-figs. 134-139) 


3, . Body about 1:o-1-5 mm., glossy black. Head broader than thorax. Eyes dark red, 
very sparsely pubescent. Antennae with second segment dark orange, third narrower, dark 
yellow, anteriorly and apically black. Arista with about four dorsal and two ventral long 
branches and a large terminal fork. Palpus black, with a fine long terminal seta and a few short 
ventral setae. Ocellar triangle glossy brownish black, convex. Periorbit glossy brownish black, 
convex, anteriorly narrowing, four-fifths as long as front. Front flat, glossy brownish black, 
shining blue, anterior one-fourth sometimes orange-brown, anteriorly one-third broader than 
length down middle ; posteriorly two-thirds as broad as head width. Face yellowish grey, 
concave. Carina high, narrow, yellowish grey, half as long as face. Clypeus black. Cheek 
yellowish grey, anterior corner black, one-sixth as broad as the greatest diameter ofeye. Occiput 
black. Postverticals minute, upright. Anterior reclinate orbital minute, outside and slightly 
behind proclinate, a little nearer to posterior reclinate than to proclinate ; proclinate slightly 
shorter than posterior reclinate. Two well developed orals, second a little shorter than vibrissa. 

Mesonotum black, glossy, slightly bluish, with fine punctures, without pubescence. Scutellum 
velvety black, tip grey pollinose. Thoracic pleura greyish brown, glossy. Three long humerals, 
uppermost one somewhat shorter than others. Acrostichal hairs in two rows. Anterior 
dorsocentrals about half as long as posteriors ; distance between anterior and posterior dorso- 
centrals half distance between anterior pair. Anterior scutellars nearly parallel, less than half 
as long as posteriors ; posteriors longer than scutellum, slightly more distant from each other 
than from anteriors. Legs yellow ; mid and hind metatarsi longer than next three segments 
together, front metatarsi slightly shorter than next three segments together. Preapicals promin- 
ent on hind leg ; apicals on mid leg. Wing hyaline, crossveins clear ; R,;, gently curved to 
costa apically ; R,;and M somewhat convergent apically. C-index about 1-6 ; 4V-index about 
2:0; 4C-index about 1:5 ; 5x-index about 1-7 ; Ac-index about 3:2. Cr-bristle one, short ; 


DIPTERA FROM NEPAL 51 


C3-fringe on basal half. Halteres yellowish orange. Abdominal tergites glossy dark brown, 
slightly shining purple. 

Periphallic organs : Genital arch brownish black, pubescent, dorsally narrowing, ventrally 
triangularly pointed ; heel high, rounded. Anal plate very narrow, ventrally broadened, pale 
yellowish brown, pubescent and setigerous. Clasper yellowish brown, elongate below, broader 
and darker basally, distally with a concave row of about fourteen marginal black teeth, discally 
with about three long stout black spines. Phallic organs : Aedeagus robust, brown, rod-shaped, 
subapically with a large dorsal triangular black projection (thus the specific name), gently sin- 
uous in S-form in lateral aspect, basally with a black narrow vertical rod and a long apically 
broadened black apodeme. PI = 1-5. Anterior paramere brown, minute, triangular, with a 
few sensilla, attached to vertical rod of aedeagus. Posterior parameres apparently absent. 
Hypandrial plate triangular, brown, apically with a large black hooked process. Ventral fragma 
quadrate, large, caudally with a pair of small black triangular submedian processes. p.f. = 
ab’CdEfg,HIKLmn. Egg-guide lobe brownish black, basally darker, broad, apically rounded, 
projecting mediodorsally, with about thirteen brown marginal teeth and a discal dorsal one. 
Basal isthmus rather long, dark brown. 


Holotype 3. East NrepaL: Taplejung District, Sangu, mixed vegetation by 
stream in gully, c. 6,200’, ix—x.1961 (R.L.C.), BMNH. 


Allotype 9. Same data as holotype, BMNH. 


Paratypes : Taplejung District, 6 3, same data as holotype ; 2 g, 1 9, Sangu, 
rotting fruits of Bhor tree on ground, c. 6,000’, 7-31.xi.19g61 ; 3g, 1 9, below Sangu, 
by stream in shady ravine, c. 6,000’, 30.x.1961 ; 1g, 1 9, between Sangu and Tam- 
rang, mixed shrubs in deepgorge, c. 5,200’, x—xi.1961. Arun Valley, 19, evergreen shrubs 
on sandy shore, below Tumlingtar, River Sabhaya, west shore, c. 1,800’. g—17.xil. 
1961 ; 44,7 9, above River Sabhaya, east shore, swept from dwarf bamboos in deep 
ravine, C. 2,000’, I2.xil.Ig61I ; 13 9, east shore of River Arun, below Tumlingtar, 
evergreen shrubs bordering dry stream-beds, c. 1,800’, I4-23.xil.Ig61 ; 2 3, 4 9, 
below Tumlingtar, west shore, fermented millet seeds on shore, c. 1,800’, 24.xii. 1961 
(R.L.C.), BMNH : 1 3, from rotting fruits, Sangu, deposited in the Tokyo Metro- 
politan University. 


Similar to L. rugulosa (de Meijere, 1915) (Java) in having brownish black finely 
punctured mesonotum, velvety black scutellum, and dark anterior corners of face, 
but differs from it in having only two rows of acrostichal hairs (six in rugulosa), 
yellow halteres (black in 7.), and yellow fore and mid femora (black in 7.). 


Liodrosophila fuscipennis sp. n. 
(Text-figs. 140-145) 


3,2. Body about1-:7-2:2mm. Head broaderthanthorax. Eyesdarkred, bare. Antennae 
with second segment conical, orange-brown ; third mainly black, as broad aslong. Arista with 
about five dorsal and two ventral long branches and a large terminal fork. Palpus distally 
broadened, with a slender apical and two shorter ventral setae. Ocellar triangle somewhat 
convex, shining black. Periorbit black, anteriorly narrowing, three-fifths as long as front. 
Front shining brownish black, anteriorly broader than length down middle ; posteriorly about 
two-thirds as broad as head width. Clypeusblack. Postverticals minute, upright. Face glossy 
brownish black, greyish white along eye margin. Carina high, brownish black, narrow, short, 


52 


T. OKADA 


FIGs. 140-151. 140-145, Liodrosophila fuscipennis sp.n. 140,35 head, lateral aspect ; 141, 


$ head, dorsal aspect ; 142, d wing; 143, periphallic organs, lateral aspect ; 144, 
phallic organs, ventral aspect ; 145, 9 egg-guide. 146-151, Liodvosophila mtida Duda. 
146, dg head, lateral aspect ; 147, d head, dorsal aspect ; 148, wing; 149, 9 egg- 
guide ; 150, periphallic organs, lateral aspect ; 151, phallic organs, ventral aspect. 


DIPTERA FROM NEPAL 53 


about half as long as face. Cheeks brownish black, grey along eye margin, two-sevenths as 
broad as the greatest diameter of eye. Occiput black. Anterior reclinate orbital minute, 
equally distant from posterior reclinate and from proclinate ; proclinate slightly inside others 
and shorter than posterior reclinate. Second oral about two-thirds as long as vibrissa. 

Mesonotum usually brownish black, glossy, without punctures. Scutellum velvety black. 
Thoracic pleura dark brown, darker below and caudally. Three humerals. Acrostichal hairs 
in about six rows. Distance between anterior and posterior dorsocentrals about three-fifths 
distance between anterior pair ; anterior dorsocentrals about two-thirds as long as posteriors. 
Anterior scutellars divergent ; posteriors twice as long as anteriors, equidistant from each other 
and from anteriors. Sterno-index about 0-5. Legs dark yellow ; fore and mid coxae and 
basal two-thirds of fore femur black ; hind coxa and trochanter yellowish white. Front and 
hind metatarsi subequal to the next three segments together, mid metatarsi subequal to the 
remaining segments together. Preapicals on mid and hind legs ; apicals strong on mid leg. 
Wings comparatively large, somewhat fuscous, crossveins clear. F,,, gently curved to costa 
at tip ; R,,,and M nearly parallel. C-index about 1-1 ; 4V-index about 1-9 ; 4C-index about 
1-6; 5x-index about 2:0; Ac-index about 4:5. Cr1-bristles two, lower weaker. C3-fringe 
sparse, on basal two-fifths. Halteres yellow. Abdominal tergites glossy black, somewhat 
purplish. 

Periphallic organs : Genital arch dark brown, rather narrow, posterior margin setigerous, 
dorsally and ventrally only moderately narrowed, ventral tip rounded ; heel low, angular, 
somewhat curved upward ; toe only slightly swollen. Anal plate yellowish brown, elongate, 
setigerous, basal margin dark. Clasper yellow, crescentric, projecting below genital arch, 
outer margin with a concave row of about nine black pointed teeth, the row not reaching 
ventral tip of clasper, and with a stout black submedian bristle. Decasternum large, yellowish 
brown, triangular. Phallic organs : Aedeagus dark brown, medially yellowish brown, fusiform 
in ventral aspect ; basal apodeme slender, slightly longer than aedeagus. PI = 1-2. Anterior 
paramere small, conical, subapically with a few sensilla. Posterior parameres apparently fused 
to decasternum. Ventral fragma V-shaped, hypandrial plate yellowish brown, apically with a 
stout yellow submedian spine. p.f. = aBCdEfgHIklm’N. Egg-guide lobe yellowish orange, 
proximally darker, tip obtusely pointed, mediodorsally much swollen, with about twenty 
marginal and four discal yellowish brown teeth. 


Holotype g. East NrepaL: Taplejung District, Sangu, mixed vegetation by 
stream in gully, c. 6.200’, xi.1961-i.1962 (R.L.C.), BMNH. 


Allotype 9. Same data as holotype (R.L.C.), BMNH. 


Paratypes: 4, 4 9, Taplejung District, between Sangu and Tamrang, mixed 
shrubs in deep gorge, c. 5,200’, x—xi.1961 ; 2 9, collected with holotype ; 3 9, below 
Sangu, by stream in shady ravine, c. 6,000’, 30.x.1g61 ; I 4, river banks below 
Tamrang Bridge, c. 5,500’, x—xi.19g61 ; 3 4g, below Sangu, mixed vegetation on 
sheltered slopes above river, c. 4,000’, 3.1.1962 ; 1 9, Sangu, mixed vegetation in deep 
gully, c. 6,200’, 2.i-13.11.1962 ; I 9, between Sangu and Tamrang, mixed plants by 
damp cliff in deep river gorge, c. 5,200’, i-li.1962 ; I g, edge of mixed forest above 
Sangu, c. 6,200’, 17.x-I.x1.1961. 1 9, Arun Valley, east shore of River Arun below 
Tumlingtar, evergreen shrubs bordering dry stream-beds, c. 1,800’, 14—23.xii.1961 
(R.L.C.), BMNH ; 1 9, at the Tokyo Metropolitan University. 


Resembles L. aerea Okada, 1956 (Japan) especially in the shape of phallic and 
periphallic organs, and somewhat fuscous wings, but differs from it in having front 
mesonotum, and abdominal tergites not shining metallic blue, and clasper teeth less 
numerous, the row of teeth not extending to the lower tip of the clasper. 


54 T. OKADA 
Liodrosophila nitida Duda 
(Text-figs. 146-151) 


Liodvosphila mitida Duda, 1922 : 157 (Annam, Formosa) ; Duda, 1924b : 241 (Formosa). 
Liodrosophila nitida var. marginifrons Duda, 1926 : 53 (Sumatra). 


36, 2. Body about 1-6mm. Head as broad as thorax. Eyes reddish brown, bare, oval in 
lateral aspect. Antennae with second segment large, rounded ; third black, basally brown, 
apically swollen and rounded, one and a half times as long as broad. Arista with about five 
dorsal and two ventral long branches and a large terminal fork. Palpus black, with a long 
apical and a few shorter ventralsetae. Ocellar triangle black, glossy. Periorbit glossy chestnut- 
black, anteriorly narrowing, four-fifths as long as front. Front glossy blue-black, anterior one- 
fourth orange-brown ; posteriorly broadened and about two-thirds as broad as head width ; 
anteriorly broader than length down middle. Cheeks dark brown, pale along eye margin, about 
one-fourth as broad as the greatest diameter of eye, narrow below. Occiput black. Anterior 
reclinate orbital minute, twice as distant from posterior reclinate as from proclinate, outside 
latter. Posterior reclinate stout, one and a half times as long as proclinate. Second oral 
slightly shorter than vibrissa, which is relatively weak. 

Mesonotum glossy bluish black, strongly convex, without punctures nor pubescence. Scutel- 
lum short, velvety black, apically with grey pubescence. Thoracic pleura glossy black, some- 
what bluish. Two subequal humerals. Acrostichal hairs in six rows. Anterior scutellars 
divergent, two-thirds as long as posteriors, which are nearer to anteriors than to each other. 
Distance between anterior and posterior dorsocentrals two-thirds distance between anterior 
pair ; anterior dorsocentrals three-fourths as long as posteriors. Sterno-index abouto-g. Legs 
yellowish brown ; femora black, glossy, apically paler. Mid metatarsi as long as the other 
segments together, fore and hind metatarsi subequal to next three segments together. Pre- 
apicals prominent on fore and mid legs ; apicals on mid leg. Wings hyaline, veins brown, 
crossveins clear. Costa reaching M, weak beyond R,,;; R.,, curved to costa at tip ; R,,,and M 
slightly convergent at tip. C-index about 1:2 ; 4V-index about 1:8 ; 4C-index about 1-7 ; 
5x-index about 1-3 ; Ac-index about 4-0. Cr1-bristle one, short ; C3-fringe on basal two-thirds. 
Halteres black, stalk yellow, knob large and triangular. Abdominal tergites glossy black, 
somewhat shining blue. Abdominal sternites small, quadrate, surrounded by pale membrane. 

Periphallic organs brownish black. Genital arch dorsally abruptly narrowed, ventrally 
slightly broadened and truncate below, apicoventrally black, projected caudally. Clasper oval, 
with a strongly convex row of about thirteen large black teeth, occupying entire outer margin, 
subapically with a large black spine on inferior surface. Anal plate elliptical, large, setigerous, 
ventrocaudal end with a tuft of short hairs. Phallic organs dark brown. Aedeagus elliptical 
in ventral aspect, medially pale ; basal apodeme slightly shorter than aedeagus, slender, straight, 
PI = 1-6. Anterior paramere small, elongate, subapically with a few sensilla. Hypandrial 
plate large, crescentic, apico-inferiorly with numerous minute tubercles. Ventral fragma 
quadrate, lateral arm pointed. p.f. = aBCdEFgHIklm’N. Egg-guide lobe brown, basally 
darker, fusiform, with both ends pointed, with about thirteen marginal and four long discal dark 
brown teeth, ultimate marginal tooth largest. Basal isthmus brown, straight, one-fifth as 
long as lobe. 


Taplejung District, old mixed forest above Sangu, c. 6,200’, ix—x.Ig61, I 3; 
Sangu, mixed vegetation by stream in gully, c. 6,200’, ix-x.1961, I 9 ; below Sangu, 
edge of small mixed wood, c. 6,000’, 4.x1.1961, I f ; below Sangu, by stream in shady 
ravine, c. 6,000’, 30.x.1961, I 9 ; Dobhan, east bank of River Tamur, mixed vegeta- 
tion by stream in deep gully, c. 3,500’, i-1.1962, 4 gd, 1 9. Arun Valley, above 
River Sabhaya, east shore, swept from dwarf bamboos in deep ravine, c. 2,000’, 
12.xli.1g61, 14 J, 15 9; below Tumlingtar, River Sabhaya, west shore, fermented 


DIPTERA FROM NEPAL 55 


millet seeds on shore, c. 1,800’, 24.xii.1g61, I 2 ; east shore of River Arun below 
Tumlingtar, evergreen shrubs bordering dry stream-beds, c. 1,800’, 14—23.xii.1g61, 
a4 (RL.C.). 

Distribution : Nepal (new locality), Sumatra, Viet-Nam, Formosa. 

Apart from its darker body, identical with the original form (Formosa), both 
having glossy blue metallic mesonotum without punctures, front anteriorly orange, 
six rows of acrostichal hairs, all femora black, somewhat convergent R,,; and M, 
and anteriorly narrowing relatively short periorbit. 


PARAMYCODROSOPHILA Duda 


Paramycodrosophila Duda, 1924a : 191. 
Type-species : Drosophila pictula de Meijere, Ig1I : 412 (Java). 


Paramycodrosophila pictula (de Meijere) 
(Text-figs. 152-157) 


Drosophila pictula de Meijere, 1911 : 412 (Java). 
Paramycodrosophila pictula (de Meijere) Duda, 1924a : 191 ; 1924b: 241 (Java, Formosa) ; 
Duda, 1926 : 58 (Sumatra) ; Wheeler and Takada, 1964 : 206 (Palau). 


6. Body about 1:5 mm., greyish white with black patches. Head slightly broader than 
thorax. Eyes large, dark red, slightly purplish, sparsely pilose. Antennae with second seg- 
ment yellow, conical ; third black, long, with long pubescence, anterior margin somewhat con- 
cave. Arista with six dorsal and one long subapicoventral branches and a large terminal fork. 
Palpus black, basally paler, slender, with a long apical seta. Ocellar triangle black. Periorbits 
two-thirds as long as front, straight, greyish white, medially narrowing, anterior end black. 
Front greyish brown, anteriorly pale yellow, posterolateral corner dark, pale around ocellar 
triangle ; anteriorly slightly narrower than length down middle ; posteriorly wider, half as 
broad as head. Face greyish white, parallel-sided. Carina high, narrow, short, pale grey, white 
at lower tip. Cheeks white, about one-sixth as broad as the greatest diameter of eye. Occiput 
brown. Postverticals minute. Anterior reclinate orbital outside and slightly before proclinate, 
as long as posterior reclinate ; posterior reclinate slightly shorter than proclinate, equidistant 
from proclinate and inner vertical. Vibrissae long, stout ; succeeding orals fine. 

Mesonotum greyish white, pollinose, with black clouds as following : a large median one 
anteriorly bilobed to reach anterior margin, a pair of large lateral ones above suture, a pair of 
smaller lateral ones above humeral callus, a large caudomedian one anteriorly narrowing and 
mediolongitudinally pale. Scutellum flat, black, grey pollinose, apically pale yellow. Thoracic 
pleura greyish white, pollinose, with two long broad black stripes. Humeral callus whitish grey. 
Two long subequal humerals. Acrostichal hairs in eight rows. Posterior dorsocentrals twice 
as long as anteriors ; distance between anterior and posterior dorsocentrals slightly more than 
one-third distance between anterior pair. Anterior scutellars convergent, as long as posteriors 
and as scutellum ; posteriors nearer to each other than to anteriors. Sterno-index about 0°5 ; 
median sternopleural fine but long. Legs yellowish grey, with black rings, two on fore femur, 
one on mid and hind femora, and two on fore tibia. Mid and hind metatarsi as long as next 
three segments together, front metatarsi subequal to next two segments together. Preapicals 
on all legs ; apicals stout on mid leg. Wings broad, hyaline, rounded at tip, with dark cloud 
below second costal break and below it between R,;, and M. Second costal section apically 
much swollen and black. Second costal break deep. &,+, nearly straight ; R,,,and M nearly 
parallel. C-index about 3:0 ; 4V-index about 2:3 ; 4C-index about 1:7 ; 5x-index about 1-9 ; 


56 


tT. OKADA 


160 


——— 


Fics. 152-163. 152-157, Pavamycodrosophila pictula de Meijere, 3. 152, head, lateral 


aspect ; 153, head, dorsal aspect; 154, wing; 155, ejaculatory apodeme; 156, 
phallic organs, ventral aspect ; 157, periphallic organs, lateral aspect. 158-163, 
Chaetodvrosophilella coei sp. n. 158, g head, lateral aspect ; 159, g head, dorsal aspect ; 
160, g wing; 161, 9 egg-guide ; 162, periphallic organs, lateral aspect ; 163, phallic 
organs, dorsal aspect. 


DIPTERA FROM NEPAL 57 


Ac-index about 3-8. Cu1-bristle one, short ; C3-fringe on basal half. Halteres white. Abdominal 
tergites mainly black ; 2~-5T each with a pair of white spots on anterior margin ; 6T white. 

Periphallic organs pale yellow. Genital arch not narrowing above, pubescent and rather 
weakly setigerous, subapically dilated ; heel high. Clasper proximally narrowing, with about 
seven large black teeth in a weakly concave row on entire distal margin and two large subapical 
black teeth. Anal plate separated from genital arch, oblong, large, setigerous and pubescent. 
Phallic organs pale yellow. Aedeagus elliptical in ventral aspect ; basal apodeme rod-shaped, 
as long as aedeagus. PI = 1-o. Anterior paramere fused to oval hypandrial plate, apically 
with about two sensilla ; hypandrial plate with a long submedian hair inserted outside sensilla. 
Posterior parameres fused to become a fan-shaped oval flap, lateral margin serrated. Ventral 
fragma large, oval. p.f. = abCdEfgHIKlm’N. Ejaculatory apodeme pale yellow with 
narrow plate and distally broadened stalk, which is slightly longer than the plate. 


Arun Valley, above River Sabhaya, east shore, swept from dwarf bamboos in deep 
ravine, C. 2,000’, 12.xii.1961, I ¢ (R.L.C.). 
Distribution : Nepal (new locality), Java, Sumatra, Formosa, Palau. 


CHAETODROSOPHILELLA Duda 


Chaetodrosophilella Duda, 1923 : 40. 
Chaetodrosophila Duda, 1924a : 180. 


Type-species : Drosophila quadrilineata de Meijere, 1911 : 396 (Java). 


Chaetodrosophilella coei sp. n. 
(Text-figs. 158-163) 


36, 2. Body about 1-6 mm., slender. Head somewhat broader than thorax. Eyes oval, 
dark red, relatively small, finely pilose. Antennae with second segment greyish brown ; third 
oval, grey, pubescent, broader than second joint. Arista with about four dorsal and two ventral 
branches and a moderate terminal fork. Palpus yellowish grey, slender, curved upward, with a 
long subapical and a few shorter ventral setae. Ocellar triangle black. Periorbit grey, large, 
anteriorly rounded, medially narrowing, three-fourths as long as front. Front yellowish grey, 
convex, anteriorly paler, depressed along margin of frontal triangle, anteriorly one and a half 
times as broad as length down middle ; posteriorly two-thirds as broad as head width. Clypeus 
yellowish grey. Cheeks yellowish grey, broad, nearly half as broad as the greatest diameter of 
eye. Occiput yellowish grey. Postverticals long. Anterior reclinate orbital minute, just 
outside proclinate, which is slightly shorter than posterior reclinate. Mouth-parts slender. 
Only one long oral. Face broad, yellowish grey, anteriorly rounded, flat, parallel-sided. Carina 
low, short, pale, half as long as front. 

Mesonotum greyish brown, somewhat pollinose, caudolateral side fuscous. Scutellum 
greyish brown. Thoracic pleura pale, darker above ; humeral callus dark. Thorax rather 
elongate. Two long humerals. Acrostichal hairs in six rows. Three pairs of dorsocentrals ; 
presutural pair slightly outside others ; distance between two posterior pairs four-fifths distance 
between individuals of each pair. Sterno-index about 0-6. Legs yellowish grey, tarsi slightly 
darker ; fore femur with a row of a few long black setae below, without a row of ventral spinules. 
All metatarsi as long as next three segments together. Preapicals on all legs ; apicals on fore 
and mid legs. Wings slightly fuscous, relatively narrow, weakly pointed at tip. R,,; and M@ 
distally somewhat divergent, medially narrowing. Crossveins clear. C-index about 2'5 ; 
4V-index about 2:7; 4C-index about 1-4 ; 5x-index abont 2:6 ; Ac-index about 2:5. C1- 
bristles two, subequal ; C3-fringe on basal half. Halteres greyish brown, knob pale. Abdominal 
tergites yellowish grey, each with a broad black medially contiguous caudal transverse band. 
Female cercus yellow, longer than broad. 


58 fy OKADA 


Periphallic organs yellowish grey. Genital arch pubescent, ventrally narrowing, pointed 
caudoventrally ; caudal margin with about twelve setae. Clasper quadrate, large, pubescent, 
with a row of about ten large pointed black teeth in a straight row on nearly entire distal margin ; 
inner surface with about eight short setae. Anal plate, oval, separated from genital arch, 
setigerous and pubescent, ventral tip with a tuft of short setae. Phallic organs : Aedeagus pale 
brown, rod-shaped, slightly curved ventrally, apically triangularly pointed. Basal apodeme 
two-thirds length of aedeagus, straight. PI = 1-3. Anterior paramere elongate, orange- 
yellow, basally attached to aedeagus, apically partially fused to hypandrium, rounded, and with 
about two long sensilla. Hypandrial plate conical, with a stout submedian spine outside 
sensilla. Ventral fragma narrow, triangular, lateral arms apically rounded. p.f. = aBCdEfg, 
HIKIMN. Decasternum triangular, orange-brown, lateral sides concave. Egg-guide lobe 
yellowish brown, darker below, elongate elliptical, rounded to tip, basally narrowing, with about 
thirteen dark brown marginal and four discal teeth, distal marginal teeth being larger. Basal 
isthmus dark, long, moderately swollen, medially about half as long as the lobe. 


Holotype g. East NepaL: Taplejung District, Sangu, mixed vegetation by 
stream in gully, c. 6,200’, xi.1961-1.1962 (R.L.C.), BMNH. 

Allotype 2. Same data as holotype (R.L.C.), BMNH. 

Paratypes : I g, same data as holotype ; I 9, same locality as holotype, ix—x. 
1961 ; Ig, between Sangu and Tamrang, mixed plants by damp cliff in deep river 
gorge, C. 5,200’, i-i1.1962 ; 1 9, ibid., mixed shrubs in deep gorge, c. 6,200’, x—x1.1961 ; 
I 9, above Sangu, evergreen scrub, c. 6,500’, 5-13.x.1961 ; I 9, ibid., mixed vegeta- 
tion in dried-up ravine, c. 6,800’, 17.11.1962 ; I g, I 9, Sangu, mixed vegetation in 
deep gully, c. 6,200’, 2.1-13.11.1962 (R.L.C.). 

Allied to C. quadrilineata (de Meijere, 1911) (Java, etc.), but differs in having 
mesonotal dark stripes obscurely demarcated (distinct in guadrilineata), six rows of 
acrostichal hairs (four in g.), two long sternopleurals (one long and two short in q.), 
and no row of spinules on fore femur. 


SCAPTOMYZA Hardy 
Scaptomyza Hardy, 1849 : 361. 
Type-species : Drosophila graminum Fallén, 1823 : 8 (Europe). 


KEY TO SPECIES 


1 Only one prominent, median, humeral. Acrostichal hairs in two rows. 4d genitalia 
with paralobes. 2 egg-guide lobe weakly sclerotized, with a few subapical long 


setae . F . SS. (Parascaptomyza) pallida (p. 59) 
— Two or three essinon Preise 3 eeceaira without paralobes. Egg-guide lobe 

well sclerotized, with numerous short stout bristles : S. (Scaptomyza) 2 
2 Two rows of acrostichal hairs. Legs mostly yellowish grey. Genital arch narrowly 

pointed below : : parasplendens (p. 59) 
— Four rows of acrostichal hae enital arch pleut Below : : 3 


3. Arista without branches below terminal fork. Clasper with a row a laxpe Stout 
pointed teeth occupying entire outer margin. Legs yellowish grey, ultimate two 
tarsal segments dark. Ocellar triangle black, pollinose’ . . melanissima (p. 62) 
— Arista with one branch below fork. Clasper with a row of marginal teeth on basal 
two-thirds on outer margin, with numerous discal teeth on lower one-third. Legs 
mostly black, fore coxa yellow. Ocellar triangle matt velvety-black . silvata (p. 63) 


DIPTERA FROM NEPAL 59 


Subgenus PARASCAPTOMYZA Duda 


Parascaptomyza Duda, 1924 : 203. 


Type-species : Drosophila pallida Zetterstedt, 1847 : 2571 (Europe). 


Scaptomyza (Parascaptomyza) pallida (Zetterstedt) 


Drosophila pallida Zetterstedt, 1847 : 2571 (Europe). 

Scaptomyza (Parascaptomyza) pallida (Zetterstedt) Hackman, 1959 : 41 (Asia, Australia, Hawaii, 
N. America, S. America, Europe, N. Aftrica, etc.) ; Wheeler and Takada, 1964 : 196 (Bonin 
Is.). 

i disticha Duda, 1921 : 64 (N. Africa). 

Scaptomyza (Parascaptomyza) disticha Duda, 1935 : 58 (Orbis Terrarum) : Okada, 1956 : 68 

apan). 

ee hovyms disticha (Duda) ; Basden, 1952 : 201 (Europe) ; Collin, 1953 : 149 (Europe) ; 
Basden, 1954 : 648 (Scotland) ; Hackman, 1955 : 80 (Finland). 

Taplejung District, Sangu, yellow blooms of cultivated Composite (Guizotia 
abyssinica Cassini), c. 6,200’, 16-29.x.1961, I f ; ibid., mixed vegetation by stream 
in gully, c. 6,200’, ix—-x.1961, I 9; ibid., xi.1g61-i.1962, 1 9 ; below Sangu, mixed 
vegetation on sheltered slope above River Tamur, c. 4,000’, 3.i.1962, I gf, 2 9; 
Dobhan, shady places on shrubby slope above River Tamur, c. 3,500’, 2I—27.i.1962, 
Ig; ex Lycopodium sp. on slope above Sangu, c. 7,800’, II-14.i.1962, 2 gd, 5 @; 
above Sangu, mixed vegetation in dried-up ravine, c. 6,800’, 16.11.1962, 1 9. Arun 
Valley, east shore of River Arun below Tumlingtar, swept from Ricinus communis L., 
c. 1,800’, 14—23.xil.1961, 7g, 7 2 ; ibid., evergreen shrubs bordering dry stream-beds, 
c. 1,800’, 14—23.xii.1961, I 9, (R.L.C.). 

Distribution : Nepal (new locality), Viet-Nam, Java, China, Formosa, Japan, 
Korea, Siberia, Iraq, Turkestan, Bonin Is., Hawaii, N. America, S. America, Europe, 
Canary Is., Azores, Madeira, Cape Verde Is., N. Africa. 

All of the Nepalese specimens examined belong to the type with dark body 
coloration. 


Subgenus SCAPTOMZYA Hardy 
Scaptomyza Hardy, 1849 : 361. 
Type-species : Drosophila graminum Fallén, 1823 : 8 (Europe). 


Scaptomyza (Scaptomyza) parasplendens sp. n. 
(Text-figs. 164-169) 


36,2. Body about 2:2 mm., dark greyish brown. Head slightly broader than thorax. Eyes 
dark red, with thick long grey pilosity. Antennae with second segment black ; third yellowish 
orange. Arista with about five dorsal branches, a long ventral one, and a small terminal fork. 
Palpus yellow, slender, apically dark. Ocellar triangle black, large, somewhat grey pollinose. 
Periorbit grey, somewhat pollinose, three-fourths as long as front, anteriorly slightly broadened. 
Front mostly yellowish grey, dark orange between periorbits and ocellar triangle, dark around 
ocellar triangle, anteriorly somewhat narrowing and narrower than median length, posteriorly 
nearly half as broad as head width. Clypeus pale brown, narrow. Face white, flat, parallel- 


60 T. OKADA 


172 


174 


Fics. 164-175. 164-169, Scaptomyza parasplendens sp. n. 164, 3 head, lateral aspect ; 
165, g head, dorsal aspect ; 166, g wing; 167, periphallic organs, lateral aspect ; 
168, phallic organs, ventralaspect ; 169, Qegg-guide. 170-175, Scaptomyza melanissima 
sp. n. 170, d head, lateral aspect ; 171, ¢ head, dorsal aspect ; 172, gd wing; 173, 
Qegg-guide ; 174, phallicorgans, ventralaspect ; 175, periphallic organs, lateral aspect. 


DIPTERA FROM NEPAL 61 


sided, broad, mediodistally with a small black spot. Carina very low, white. Cheeks yellowish 
white, black above, about one-fifth as broad as the greatest diameter of eye. Occiput black, 
medially yellow. Anterior reclinate orbital about half as long as proclinate, just outside pro- 
clinate, much nearer to proclinate than to posterior reclinate. Posterior reclinate slightly 
longer than proclinate. Vibrissa long, stout ; second oral about one-third as long as vibrissa ; 
succeeding orals shorter. 

Mesonotum greyish brown, bluish grey pollinose, anteriorly pale, dark along dorsocentral lines 
and caudomedially. Scutellum grey pollinose, grey-brown, laterally pale. Thoracic pleura 
greyish brown, pollinose. Two long humerals, lower slightly longer. Acrostichal hairs in two 
rows. Anterior scutellar long, divergent ; posterior scutellar three-fifths as long as anteriors, 
one and a half times as distant from anterior as from each other. Sterno-index 0-6. Legs 
mostly yellowish grey ; coxae yellow ; fore tibia and fore tarsi brownish black ; last two tarsal 
segments of mid and hind legs black. Pulvilli white, large. Front metatarsi as long as next 
two segments together, mid and hind metatarsi as long as next three segments together. Pre- 
apicals on all legs ; apicals prominent on fore leg. Wings narrow, hyaline ; crossveins clear. 
Ro+, gently curved to costa at tip ; R,,;and M parallel ; costareaching M. C-index about 2°6 ; 
4V-index about 1-4 ; 4C-index about 0-7 ; 5x-index about 1-8 ; Ac-index about 3:4. Cr1-bristle 
two ; C3-fringe on basal one-third. Halteres yellow, elongate, dark at tip. Abdominal 
tergites greyish black, bluish grey pollinose. Abdominal sternites pale grey. 

Periphallic organs : Genital arch brownish black, pubescent, dorsally narrowing, anterior 
margin with narrow projection below. Clasper brownish black, large, triangular, dorsocaudally 
with an acute black projection, distally with a convex row of about fourteen pointed black teeth 
on almost entire margin, and with a few short marginal setae below. Anal plate brownish black, 
very large, separated from genital arch, ventrally broadened, rounded below, setigerous and 
pubescent. Phallic organs : Aedeagus yellowish orange, oblong, apically rounded in lateral 
aspect, basally narrowing. Basal apodeme slender, rod-like, half as long as aedeagus. PI = 
2:0. Anterior paramere minute, conical, distally with a few sensilla. Ventral fragma tri- 
angular, brownish ; lateral arm broad and pointed apically ; median notch deep, acute ; 
submedian process narrow, without submedian spine. p.f. = aBCdEfg,HikLmn. Egg-guide 
lobe dark brownish black, robust, distal margin obliquely truncate, with about twenty-four 
marginal and twenty scattered discal strong black teeth. Basal isthmus short. 


Holotype 3. East NepaL : Taplejung District, Sangu, mixed vegetation in deep 
gully, c. 6,200’, 2.i-13.ii.1962 (R.L.C.), BMNH. 


Allotype 9. Same data as holotype (R.L.C.), BMNH. 


Paratypes : Taplejung District, 7 J, 4 9, same data as holotype ; 11 4, 8 9, 
Sangu, mixed vegetation by stream in gully, c. 6,200’, xi.1g61-i.1962 ; I 4g, 2 8, 
Sangu, mixed vegetation by stream in gully, c. 6,200’, ix—x.1961 ; 1 4, ibid., clearing 
in small wood, c. 6,200’, 25.xi.1961 ; I 9, ibid., spray-splashed rocks in shallow ravine 
c. 6,200’, 25-28.x.1g61 ; I 9, ibid., damp evergreen oak forest, c. 9,200’, 2—26.xi. 
1961 ; 7 dg, 6 Q, ibid., mixed vegetation in dried-up ravine, c. 6,800’, 16.11.1962 ; 
2 3, below Tamrang Bridge, river banks, c. 5,500’, x—xi.1g61 ; I g, below Sangu, 
edge of small mixed wood, c. 6,000’, 2—26.xi.1961 ; Ig, ibid., 4.xi.19g61 ; 2 Q, ibid., 
mixed vegetation on sheltered slope above river, c. 4,000,’ 3.1.1962 ; 2 9, ibid., by 
stream in shady ravine, c. 6,300’, 30.x.1961 ; 3 9, Dobhan, east bank of River 
Tamur, mixed vegetation by stream in deep gully, c. 3,000’, i-i.1962 (R.L.C.), 
BMNH, 1 3g, 1 2 at the Tokyo Metropolitan University. 


Closely allied to S. subsplendens Duda, 1935, (East Siberia) especially in having 
two rows of acrostichal hairs, medially pale occiput, and ¢ with large anal plate, but 


62 T. OKADA 


differs from it in having paler periorbit (black in subsplendens,) mostly yellow front 
(mostly dark in s.), f anal plate as large as in S. graminum (Fallén) (smaller in s.), 
and black fore tibia. 


Scaptomyza (Scaptomyza) melanissima sp. n. 
(Text-figs. 170-175) 


3,2. Body about1:8mm. Head slightly broader than thorax. Eyes dark purplish brown, 
thickly pilose, relatively small and oval. Antennae with second segment yellowish orange, 
dorso-anteriorly darker ; third pale yellowish orange, triangular. Arista with five short 
branches including a moderate fork, without ventral branches. Palpus yellow, apically swollen, 
with a few stout black ventral setae. Ocellar triangle black, large, grey pollinose, convex. 
Periorbit broad, black, grey pollinose, anterior tip rounded, about three-fourths as long as front. 
Front mostly black, grey pollinose, anteriorly orange, especially dark between periorbits and 
ocellar triangle, with a few frontal hairs, anteriorly narrowing and narrower than length down 
middle, posteriorly two-thirds as broad as head width. Clypeus small, yellowish grey. Face grey- 
ish white matt, concave, wider below. Carina low, narrow, slightly wider below. Cheeks white, 
posteriorly black, about one-third as broad as the greatest diameter of eye. Occiput black. 
Postverticals long. Anterior reclinate orbital two-thirds as long as proclinate, just outside 
proclinate, thrice as distant from posterior reclinate as from proclinate. Posterior reclinate 
slightly longer than proclinate. Only one long stout black oral. 

Mesonotum matt black, bluish grey pollinose, with diffuse median and lateral black longitu- 
dinal stripes, lateral pair extending to scutellum. Scutellum flat, bluish black pollinose. 
Thoracic pleura black, grey pollinose, somewhat bluish. Three long humerals, lowermost 
longest. Acrostichal hairs in six rows, relatively long. Posterior dorsocentrals more than twice 
as long as anteriors ; distance between anterior and posterior dorsocentrals three-fifths distance 
between anterior pair. Anterior scutellars very long, divergent, twice as long as scutellum ; 
posteriors about three-fifths as long as anteriors, twice as distant from anteriors as from each 
other. Sterno-index about o-7. Legs yellowish grey, last tarsal segment large, black ; penulti- 
mate black ; front and hind metatarsi longer than the other segments together, hind metatarsi 
subequal to next three segments together. Preapicals on all legs, apicals prominent on mid leg. 
Wings hyaline, narrow, slightly pointed at tip ; crossveins clear. R,,, medially convex 
anteriorly, apically gently curved to costa ; R,;; and MJ somewhat convergent distally. C-index 
about 4:0 ; 4V-index about 1-1 ; 4C-index about 0-5 ; 5x-index about 1-5 ; Ac-index about 
2:6. C1-bristle one ; C3-fringe on basal one-third. Halteres white, knob rounded and large. 
Abdominal tergites slender, black, matt, somewhat bluish grey pollinose. 

Periphallic organs : Genital arch brownish black, pubescent, dorsally not narrowing, ventrally 
broadly truncate and with several setae, subapically pale on caudal half. Clasper brownish 
black, crescentic, marginally with a concave row of about twelve long stout black pointed teeth, 
the row covering entire distal margin of clasper. Anal plate pale brownish black, elliptical, 
setigerous, pubescent, lower tip narrowly pointed and with a few short setae. Decasternum 
large, quadrate, tripartite. Phallic organs: Aedeagus yellowish grey, elliptical, basally 
tapering, ventral margin serrated and with a yellow vertical rod ; apodeme brown, straight. 
PI = 1-5. Anterior paramere minute, sensilla obscure, ventral fragma greyish brown, deeply 
notched, lateral arms distally broadened. p.f. = abCdEf)g,HikLmn. Egg-guide lobe large, 
oval, dark brown, distally broadly rounded, with about fifteen marginal and twenty scattered 
discal strong black teeth. Basal isthmus narrow, black, about one-fifth as long as egg-guide 
lobe. 


Holotype g. East NepaL : Taplejung District, damp evergreen oak forest above 
Sangu, c. 9,200’, 2—26.xi.1961 (R.L.C.), BMNH. 


DIPTERA FROM NEPAL 63 


Allotype 9. Taplejung District, Sangu, clearing in small wood, c. 6,200’, 25.x. 1961 
(R.L.C.), BMNH. 


Paratype g: Taplejung District, ex Lycopodium sp. on slope above Sangu, 
c. 7,800’, II—14.1.1962 (Head missing) (R.L.C.), BMNH. 


Closely allied to S. grahami Hackman, (China) but ocellar triangle not darker than 
any part of front, arista with three dorsal branches (four in grahamz), thoracic 
pleura not dark reddish brown, scutellum not brown medially, and male clasper with 
fewer and stronger teeth on entire outer margin (on lower half and without stout 
teeth in g.). 


Scaptomyza (Scaptomyza) silvata sp. n. 
(Text-figs. 176-182) 


3,9. Body about 2:2mm. Head very slightly broader than thorax. Eyes oval, relatively 
small, dark purplish brown, thickly pilose. Antennae orange-yellow ; second segment brown 
at base ; third broad, triangular. Arista with about three dorsal and a ventral short branches 
and a small terminal fork. Palpus orange in 3, black in 9, apically with a few rather short stout 
black setae. Ocellar triangle velvety black, large, longer than broad. Ocelli greyish brown. 
Periorbit grey, pollinose, without frontal hairs, apically rounded and not reaching anterior 
margin of front. Front grey, pollinose, anteriorly orange, somewhat narrowing, as broad as 
length down middle ; posteriorly about half as broad as head width. Face yellowish white, 
slightly concave, nearly parallel-sided. Carina low, black, narrow, broadened below, reaching 
buccal margin. Cheek yellowish white, black posteriorly, about one-fourth as broad as the 
greatest diameter of eye. Occiput black, medially white, pollinose. Anterior reclinate orbital 
two-thirds as long as proclinate, outside proclinate, and thrice as distant from posterior reclinate 
as from proclinate. Proclinate slightly shorter than posterior reclinate. Vibrissae strong, second 
and third orals one-fourth as long as vibrissae. 

Mesonotum black, bluish grey pollinose, with a median and lateral black longitudinal stripe, 
median one reaching end of scutellum. Scutellum black, bluish grey pollinose. Thoracic 
pleura black, somewhat glossy. Two humerals, upper longer. Acrostichal hairs in four rows. 
Anterior dorsocentrals four-fifths as long as posteriors ; distance between anterior and posterior 
dorsocentrals about five-eighths distance between anterior pair. Anterior scutellars long, 
divergent, twice length of scutellum ; posteriors two-thirds as long as anteriors, one and a half 
times as distant from anteriors as from each other. Sterno-index about 0-5. Legs greyish 
brown, trochanters and fore coxa yellow, femora black, last two tarsal segments black ; all 
metatarsi as long as next three segments together. Preapicals on all legs ; apicals on mid leg. 
Wings hyaline, veins brown, tip somewhat pointed. #,,, medially convex anteriorly, gently 
curved to costa apically ; costa reaching end of M ; R,;; and M parallel. C-index about 3:1 ; 
4V-index about 1-4 ; C4-index about 0-7 ; 5x-index about o-9 ; Ac-index about 2:5. Cr- 
bristle one, long ; C3-fringe on basal one-third. Halteres yellow. Abdominal tergites black, 
somewhat glossy. Abdominal sternites black, large quadrate. 

Periphallic organs : Genital arch dark brownish black, pubescent, dorsally much narrowing, 
ventrally broadly truncate and with a few long setae, subapically with a deep caudal incision. 
Clasper oval, dark brownish black, marginally with a row of about ten black teeth, discally with 
numerous black pointed teeth in about three rows above and six rows below. Anal plate dark 
brownish black, very large, fusiform, separated from genital arch, setigerous and pubescent. 
Decasternum rhombic, dark brownish black. Phallic organs: Aedeagus yellowish brown, 
somewhat sinuous, elliptical, tip rounded in lateral aspect. Apodeme black, narrow, short and 
straight, about one-fourth as long as aedeagus. PI = 4:0. Vertical rod as long as apodeme, 
black, slender. Anterior paramere small, oval, pubescent, brownish black, attached to the 


64 T. OKADA 


vertical rod, with a few sensilla subapically. _Hypandrial plate elongate, basally broad. Ventral 
fragma V-shaped. p.f. = aBCdEfg,HikLmn. Egg-guide lobe robust, brownish black, 
abruptly narrowing proximally, broadly truncate distally, with about twenty-four marginal and 
seventeen discal strong black pointed teeth. Basal isthmus black, short, one-sixth as long as 
egg-guide lobe. 


Fics. 176-182. Scaptomyza silvatasp.n. 176, 3 head, lateralaspect ; 177, ¢ head, dorsal 
aspect ; 178, g wing; 179, 2 egg-guide; 180, aedeagus, lateral aspect; 181, 
phallic organs, ventral aspect ; 182, periphallic organs, lateral aspect. 


DIPTERA FROM NEPAL 65 


Holotype g. East Nepat : Taplejung District, damp evergreen oak forest above 
Sangu, c. 9,200’, 2—26.xi.1961 (R.L.C.), BMNH. 


Allotype 9. Same data as holotype (R.L.C.), BMNH. 


Paratypes : 3 2, same data as holotype (R.L.C.), BMNH, 1 9 at the Tokyo 
Metropolitan University. 


Closely resembles S. migrita Wheeler, 1952, (N. America) but differs slightly in 
having deep velvety black, not pollinose, large ocellar triangle and front only 
orange anteriorly. 


DIATHONEURA Duda 


Diathoneuva Duda, 1924a : 180. 


Type-species : Drosophila taematipennis Duda, 1925: 168; designated by 
Malloch, 1934 (S. America). 


Diathoneura ripa sp. n. 


(Text-figs. 274-279, p. 94) 


3, §. Body about 2 mm., small, glossy black. Head slightly broader than thorax. Eyes 
large, dark reddish brown, with thick pilosity. Antennae black, third segment oval and with 
grey pubescence, apically widely apart from face. Palpus black, with a prominent apical bristle. 
Ocellar triangle black, convex, straight somewhat glossy. Periorbit black, slightly glossy, 
posteriorly somewhat narrowing, anteriorly blunt, not reaching anterior margin of front. Front 
evenly black, matt, grey pollinose, anteriorly grey, slightly broader than length down middle, 
posteriorly broader and slightly narrower than head width. Clypeus black. Face black, flat, 
parallel-sided, whitish between second segments of antennae. Carina entirely absent. Cheeks 
black, narrow, about one-ninth as broad as the greatest diameter ofeye. Occiput black. Anter- 
ior reclinate orbital slender, half as long as proclinate, one-fourth as long as posterior reclinate, 
about twice as distant from posterior reclinate as from proclinate. Proclinate inside other 
orbitals. Vibrissae long, second oral one-third as long as vibrissae ; other orals shorter. Post- 
verticals fine, very short. 

Mesonotum black, glossy, with fine punctures. Scutellum black, somewhat grey pollinose. 
Thoracic pleura glossy black. Two long humerals. No prescutellars. No propleurals. 
Acrostichal hairs in six to eight rows. Anterior dorsocentrals two-thirds as long as posteriors ; 
distance between anterior and posterior dorsocentrals half distance between anterior pair. 
Anterior scutellars divergent, slightly longer than posteriors, which are equally distant from 
each other and from anteriors. Sterno-index about 0-7. No prominent median sterno- 
pleural. Legs short, thick, dark brown, femora black. All metatarsi as long as the other segments 
together. Preapicals on all legs ; apicals on fore and mid legs. Wings hyaline, relatively 
slender, veins pale yellow, crossveins not clouded, tip rounded. , , straight, curved to costa 
at tip. R,,, and M distally somewhat convergent. C-index about 2:3 ; 4V-index about 3-1 ; 
4C-index about 2-6; 5x-index about 3:1 ; Ac-index about 5-3. Cz1-bristles only one ; C3- 
fringe on basal five-eighths. Halteres white. Abdominal tergites entirely black, somewhat 
glossy. Abdominal sternites pale brown, small, elongate. 

Periphallic organs: Genital arch black, dorsally slightly narrowing, setigerous ventro- 
posteriorly ; toe deep black, rounded ; heel somewhat high. Clasper black, crescentic, 
dorsally narrowly pointed, with a slightly concave row of about thirteen black teeth on entire 
distal margin. Anal plate fusiform, greyish black, setigerous, narrowly pointed ventroposterior- 
ly. Phallic organs: Aedeagus black, short, apparently bifurcated and hooked apically, 


66 Dy OFCLAD A 


medially broadened. Anterior paramere bifid, black, slender, basally recurved ventrocaudally 
and attached to hypandrium. Posterior parameres fused to form a large dorsally folded quad- 
rate flap. Ventral fragma quadrate, finely pubescent medially, and distally with a pair of short 
submedian processes. PI =0-3. p.f. = aBCdEf,gHikKLmN. Egg-guide lobe brown, dist- 
ally orange, fusiform, apically pointed, with about fifteen sharply pointed black marginal teeth, 
apical tooth largest, and with a prominent pale subterminal hair. Basal isthmus thick, black, 
half as long as lobe. 

Holotype g. East NrEpaL: Taplejung District, river banks below Tamrang 


Bridge, c. 5,500’, x—xi.1961 (R.L.C.), BMNH. 
Allotype 2. Same data as holotype (R.L.C.), BMNH. 


Paratypes : 6 3, 3 2, I ex. (abdomen missing), same data as holotype ; I 9, 
Taplejung District, below Sangu, mixed vegetation on sheltered slopes above river, 
c. 4,000’, 3.1.1962 (R.L.C.), BMNH, 1 f and 1 9 (below Tamrang Bridge) deposited 
at the Tokyo Metropolitan University. 


Small body size, minute postverticals, non-carinate face and relatively short 
anterior reclinate orbital are the features characteristic of the Neotropical genus, 
Diathoneura Duda. Differing from the American species of the genus, however, the 
present species has eyes with thick pilosity. Frota-Pessoa (1947) treats this genus 
as a synonym of Clastopteromyia Malloch, 1924 (in Malloch and McAtee, Proc. Biol. 
Soc. Wash. 37 : 27), but Wheeler (1952) considers them to be distinct genera. 


DROSOPHILA Fallén 
Drosophila Fallén, 1823 : 4. 
Type-species : Musca funebris Fabricius, 1787 : 345 (Sweden). 


KryY TO SUBGENERA 


1 Preapicals prominent only on hind tibiae. : ; é : ; : ; 2 
Preapicals prominent on all three tibiae : : : ; : 
2 The longest axis of eye nearly rectangular to body axis. 4 with submedian spines of 
novasternum short and usually paired. Egg-guide teeth usually yellowish brown 
HIRTODROSOPHILA Duda (p. 77) 
— The longest axis of eye considerably oblique to body axis. g novasternum with 
submedian spines rather long and in two pairs. Egg-guide teeth black 
DORSILOPHA Sturtevant (p. 81) 
3 Prescutellars present. ¢ with submedian spines of novasternum exceedingly long and 


stout. Egg-guide teeth yellowish brown . SCAPTODROSOPHILA Duda (p. 67) 
— Prescutellars absent. g with submedian spines of novasternum usually short and 
weak or frequently absent . : ; : : : 4 


4 The longest axis of eye very oblique = the pate a axis. Oceans inserted well outside 
ocellar triangle. Clasper teeth and egg-guide teeth relatively large in size and few 
innumber . : . : DICHAETOPHORA Duda (p. 73) 

— The longest axis of eye Pearly pooner to the body axis. Ocellars inserted inside 
ocellar triangle. Clasper teeth and egg-guide teeth aa Gee small in size and 


large in number . : 2 : : 5 
5 Abdominal tergites with realy: eonteneae fiaete cross- “banda: Ege guide teeth 
usually black : 2 : SOPHOPHORA Sturtevant (p. 81) 


— Abdominal tergites usually rae medially fereeeeeee black cross-bands or spots. 
Egg-guide teeth usually yellowish brown DROSOPHILA Fallén, s. str. (p. 96) 


DIPTERA FROM NEPAL 67 


Subgenus SCAPTODROSOPHILA Duda 
Scaptodrosophila Duda, 1923 : 37. 
Pholadoris Sturtevant, 1942 : 28 [teste Wheeler and Takada, 1964 : 185]. 
? Pavadrosophila Duda, 1923 : 43. 
Type-species : Scaptodrosophila scaptomyzoidea Duda, 1923 : 37 (New Guinea). 


As pointed out by Wheeler (1957) and Wheeler and Takada (1964), re-examination 
of Drosophila pictipennis Kertész, the type-species of Paradrosophila designated by 
Sturtevant, 1927, is required in determining relationships between Paradrosophila 
and Scaptodrosophila. The characteristics of this subgenus are most clearly expressed 
in male genital organs, and Drosophila brunnea Duda and D. pilicrus Duda are 
tentatively included in this subgenus as they have been included by Duda in his Para- 
drosophila, although the females alone have been examined by the present author. 


KEY TO SPECIES 


1 Scutellum yellowish brown, laterally black, apically white. Eyes bare. Arista with 
three branches below fork. Abdominal tergites with medially contiguous, laterally 
not broadened, black cross-bands_ . : . brunnea (p. 67) 
— Scutellum unicolorously yellowish brown, dark brown, or black. Arista with two 
branches below fork. Eyes pubescent. Abdominal tergites each with medially 
interrupted, laterally broadened, black cross-bands , 4 : E 2 
2 Wings black at base : : " ; : f . j . pilicrus (p. 67) 
— Wings not black at base : : . : 
3 Mesonotum deep black. Front velvety black. Wings hyaline Minute species with 


body length about 1 mm. ; - . minima (p. 69) 
— Mesonotum yellowish or reddish brown. Wings fuscous : : : : 4 
4 Front reddish brown, somewhat velvety. Scutellum dark brown. Two long 

humerals. R,,3 apically not curved to costa . : : marginata (p. 70) 
— Front golden brown. Scutellum yellowish brown, pollinose. Three long humerals. 

Ry,3 apically curved to costa F ; : é 5 . alternata (p. 72) 


Drosophila (Scaptodrosophila) brunnea de Meijere subgen. comb. n. 


Drosophila brunnea de Meijere, 1911 : 401 (Java). 

Drosophila (Paradrosophila) brunnea de Meijere ; Duda, 1924a : 206 (Java) ; Duda, 1926 : 79 
(Sumatra) ; Okada, 1964b : 445 (Borneo). 

Drosophila (Paradrosophila) scutellimargo Duda, 1923 : 43 {mom. nud.] ; Duda, 1924a : 206 
243 (Formosa). 


Taplejung District, Sangu, mixed vegetation by stream in gully, c. 6,200’, ix—x. 
moor, 1:2 °(R.L.C.). 
Distribution : Nepal (new locality), Java, Sumatra, Borneo, Formosa. 


, 


Drosophila (Scaptodrosophila) pilicrus Duda subgen. comb. n. 
(Text-figs. 183-185) 
Drosophila (Paradrosophila) pilicrus Duda, 1926 : 74 (Sumatra). 


9. Body about 2 mm. Head slightly broader than thorax. Eyes dark red, with fine 
pilosity. Antennae yellowish brown, third segment one and a half times as long as broad. 
Arista with about seven or eight dorsal and two or three ventral branches and a moderate 
terminal fork. Palpus brown, with an apical and a few ventral setae. Ocellar triangle matt 


68 


T. OKADA 


187 


190 


Fics. 183-195. 183-185, Drosophila (Scaptodrosophila) pilicrus Duda, 9. 183, 


head, lateral aspect ; 184, head, dorsal aspect ; 185, egg-guide. 186-191, Dyoso- 
phila (Scaptodrosophila) minima sp. n. 186, g head, lateral aspect ; 187, ¢ head, dorsal 
aspect ; 188, 2 egg-guide; 189, § wing; 190, periphallic organs, lateral aspect ; 
191, phallic organs, ventral aspect. 192-195, Drosophila (Drosophila) pentastriata sp. n. 
3. 192, periphallic organs, lateral aspect ; 193, phallic organs, ventral aspect ; 194, 
aedeagus, lateral aspect, sinistral side ventral ; 195, head, lateral aspect. 


DIPTERA FROM NEPAL 69 


black, somewhat pollinose, narrowing posteriorly, anteriorly rounded and ending shortly before 
anterior margin of front, laterally entirely separated from eye margin. Front dark brown, 
somewhat grey pollinose, nearly parallel-sided, black around ocellar triangle, anteriorly deeply 
depressed downward and twice as broad as length down middle, posteriorly five-eighths as broad 
as head width. Frontal hairs absent. Clypeus black. Face greyish brown, laterally black, 
parallel-sided. Carina high, long, ending before buccal margin, mediolongitudinally pale grey, 
laterally black. Cheeks greyish white, one-fifth as broad as the greatest diameter of eye. 
Occiput black. Postverticals very slender. Anterior reclinate orbital just outside proclinate, 
two-thirds as long as proclinate, more than twice as distant from posterior reclinate as from 
proclinate ; proclinate slightly shorter than posterior reclinate. Second oral slightly shorter 
than vibrissa ; succeeding orals gradually shorter. 

Mesonotum matt black. Scutellum matt black, grey pollinose. Thoracic pleura dark grey, 
lower tip of sternepisternum darker. Two long humerals. Sterno-index about 0-8. Pre- 
scutellar as long as anterior dorsocentral. Posterior dorsocentrals much longer than anteriors ; 
distance between anterior pair of dorsocentrals twice distance between anterior and posterior 
pair. Legs yellowish grey ; femora and fore and mid tibiae brownish black ; fore femur 
ventrally and apicodorsally with a few long black setae. Preapicals on all legs ; apicals on mid 
leg. Front metatarsi slightly shorter than next two segments together, mid and hind metatarsi 
subequal to next two segments together. Wings hyaline, veins yellow ; wing base and tip of 
second costal section black. R,,, nearly straight ; R,,, and M parallel. C-index about 1°5 ; 
4V-index about 2:0; 4C-index about 1°5 ; 5x-index about 1°5 ; Ac-index about 3:0. Cr1- 
bristle one, long ; C3-fringe of basal two-thirds. Halteres black, grey pollinose. Abdominal 
tergites yellow ; 1T-dark brown. Anal cerciconical. Egg-guide lobe orange brown, elongate, 
triangular, apically darker, narrowly pointed, basally much broadened, subterminally with a long 
marginal hair, without teeth. 


Taplejung District, Sangu, mixed vegetation by stream in gully, c. 6,200’, ix—x. 
goer, FD (R.E.C.). 

Distribution : Nepal (new locality), Sumatra. 

It is noticeable that the egg-guide resembles that found in the genus Micro- 
drosophila. 


Drosophila (Scaptodrosophila) minima sp. n. 
(Text-figs. 186-191) 


3, 2. Body about 1 mm., velvety black. Head black, as broad as thorax. Eyes dark red, 
finely pilose. Antennae with second segment dark brown ; third black, one and a half 
times as long as broad, rounded at tip. Arista with about four dorsal and two ventral long 
branches and a large terminal fork. Palpus black, with a strong apical and a weaker ventral 
setae. Ocellar triangle convex, large, black, somewhat grey pollinose. Periorbit dark greyish 
black, pollinose, anteriorly somewhat broadened and ending near the anterior margin of front. 
Front velvety black, anterior margin deeply concave, anteriorly narrowing and slightly narrower 
than length down middle, posteriorly half as broad as head width. Clypeus black, somewhat 
glossy blue. Face greyish black, medially somewhat broadened. Carina dark brown, long, 
high, almost reaching buccal margin. Cheeks narrow, black, grey pollinose, about one-seventh 
as broad as the greatest diameter of eye. Occiput black. Postverticals moderate in size. 
Mouth-parts greyish brown. Anterior reclinate orbital just outside proclinate, twice as distant 
from posterior reclinate as from proclinate, slightly shorter than proclinate, which is about two- 
fifths as long as posterior reclinate. Vibrissae strong ; second oral weak, half as long as vibrissae; 
other orals shorter. 

Mesonotum deep black, somewhat glossy, setigerous. Scutellum velvety black. Thoracic 
pleura black. Twohumerals. Prescutellars five times as long as anterior dorsocentrals. Acro- 


70 T. OKADA 


stichal hairs in eight rows. Anterior dorsocentrals half as long as posteriors ; distance between 
anterior and posterior dorsocentrals about one-third distance between anterior pair. Anterior 
scutellars parallel, long, as long as posteriors or as scutellum itself ; posteriors equally distant 
from anteriors and from each other. Sterno-index about 0-7. Median sternopleural long, 
intermediate in size between anterior and posterior ones. Legs black, femora darker. All 
metatarsi as long as rest of tarsal segments together. Preapicals on all legs ; apicals strong on 
mid leg. Wings hyaline, tip rounded, veins brown. Crossveins clear. Costa reaching end of 
R415 ; Ro13 medially straight, apically slightly curved to costa ; R,,, and M parallel. C-index 
about 1:3 ; 4V-index about 2:2 ; 4C-index about 2:3 ; 5x-index about 2-8 ; Ac-index about 
5:0. Cr1-bristles two, subequal ; C3-fringe on basal five-sixths. Halteres black. Abdominal 
tergites entirely black, somewhat glossy. 

Periphallic organs : Genital arch greyish brown, pubescent, not narrowing dorsally, broadly 
truncate ventrally, ventral tip black and somewhat swollen, posterior margin with a few long 
setae in a row, lateromedially somewhat swollen caudally. Clasper black, quadrate, lower tip 
pointed ; distal margin broad and with a concave row of about eight strong pointed black teeth, 
the row occupying entire length of distal margin. Anal plate greyish brown, separated from 
genital arch, oblong, setigerous and pubescent, caudal tip truncate and with a few strong short 
setae. Phallic organs: Aedeagus apparently bilobed, elongate, black, fusiform in ventral 
aspect. Apodeme straight, black, slightly shorter than aedagus. Anterior paramere large, 
dark brown, basally black, apically rounded, basally contiguous with aedeagus, with a row of 
about four sensilla on distal half. Posterior parameres dark brown, fused to be a broad distally 
flattened medially pale lobe. Hypandrial plate large, crescentric, dark brown, with a few 
submedian spines. Ventral fragma broad, brown, V-shaped, marginally black. PI = 1°5. 
p.f. = ABCDeFgHIKLMN. Egg-guide lobe dark brown, elliptical, apically narrowly rounded, 
with about fourteen marginal and three discal stout black teeth, ventral margin with a pro- 
minent subterminal hair. Basal isthmus black, one-third as long as lobe, medially broadened. 


Holotype g. East NepaLt: Arun Valley, east shore of River Arun below Tum- 


lingtar, evergreen shrubs bordering dry stream-beds, c. 1,800’, 14-23.xi1.1961 
(R.EC.), BMINE, 


Allotype 9. Same data as holotype (R.L.C.), BMNH. 


Paratypes : Ig, 2 9, same data as holotype (R.L.C.), BMNH, 1 9 at the Tokyo 
Metropolitan University. 


Distinguished from other members of the subgenus by extremely small body. 
Somewhat resembles D. coracina Kikkawa and Peng, 1938 (Japan and China), in 
the shape of genital arch of male periphallic organs, but differs in having much smal- 
ler and less glossy body and spiny clasper teeth. 


Drosophila (Scaptodrosophila) marginata Duda subgen. comb. n. 
(Text-figs. 201-206) 


Drosophila (Paradrosophila) marginata Duda, 1924a : 209 (Formosa) ; Duda, 1923 : 46 [nom. 
nud.] (Formosa). 


36, 2. Body about 2.4-2:-8 mm. Head as broad as thorax. Eyes dark reddish brown, 
finely pilose. Antennae with second segment brown ; third dark brown, posterior margin pale. 
Arista with about four dorsal and two ventral branches and a fine terminal fork. Palpus yellow- 
ish orange, with an apical and a ventral black setae. Ocellar triangle black. Periorbit greyish 
brown, somewhat glossy, short, half as long as front, anteriorly blunt, posteriorly pale and 
broadened. Front yellowish brown, somewhat velvety, parallel-sided, anteriorly somewhat 
darker and as broad as length down middle, posteriorly about half as broad as head width. 


DIPTERA FROM NEPAL 7 


206 


Fics. 196-206. 196-200, Drosophila (Scaptodrosophila) alternata de Meijere, 3. 196, head, 
lateral aspect ; 197, head, dorsal aspect ; 198, wing; 199, periphallic organs, lateral 
aspect ; 200, phallic organs, ventral aspect. 201-206, Drosophila (Scaptodrosophila) 
marginata Duda. 201, head, lateralaspect ; 202,$head,dorsalaspect ; 203, wing ; 
204, 2 egg-guide ; 205, periphallic organs, lateral aspect ; 206, phallic organs, ventral 
aspect. 


72 T. OKADA 


Face yellowish grey. Carina yellowish grey, high, broad, long. Clypeus yellowish brown. 
Cheek yellowish brown, very narrow, about one-fifteenth as broad as the greatest diameter of 
eye. Occiput dark brownish black. Postverticals long, slightly shorter than inner verticals. 
Anterior reclinate orbital short, just outside and shortly behind proclinate, one-third as long as 
proclinate, one-fourth as long as posterior reclinate, thrice as distant from posterior reclinate as 
from proclinate. Second oral half as long as vibrissae ; succeeding orals as long as second. 

Mesonotum dark brown, matt, humeral callus paler, along dorsocentral line pale. Scutellum 
flat, dark brown, somewhat velvety, elongate. Thoracic pleura dark brown, paler below. Two 
humerals, lower shorter. Prescutellars as long as anterior dorsocentrals. Acrostichal hairs in 
six rows. Anterior dorsocentrals one-third as long as posteriors ; distance between anterior 
and posterior dorsocentrals about half distance between anterior pair. Anterior scutellars 
divergent, slightly longer than posteriors, which are nearer to each other than to anteriors. 
Sterno-index about 0-7 ; median sternopleural as long as posteriors. Legs yellow, including 
coxa. Preapicals on all legs ; apicals prominent on mid leg. All metatarsi much longer than 
the other tarsal segments together. Wing very fuscous especially along costal margin, relatively 
large ; veins dark ; crossveins not clouded but distinct. R,,, nearly straight, not curved to 
costa apically ; R,,;and M slightly divergent ; costareaching M. C-index about 4:5 ; 4V-index 
about 1:5 ; 4C-index about 0°5 ; 5x-index about 1:3 ; Ac-index about 1-4. Cr1-bristles two ; 
C3-fringe on basal one-third. Abdomen elongate ; abdominal tergites yellowish brown, each 
with a caudal, medially contiguous, latero-anteriorly curved, black cross-band. Abdominal 
sternites pale yellow. 

Periphallic organs : Genital arch pale brownish black, densely pubescent and setigerous, 
dorsally not distinctly narrowing, ventral half broader, apically sharply truncate. Clasper pale 
brownish black, hemispherical, not extending below genital arch, outer margin with about 
thirteen strong black pointed teeth in a slightly concave row, occupying entire length of outer 
margin. Anal plate small, elliptical, separated from genital arch, setigerous and pubescent, 
ventrally obtusely pointed. Phallic organs : Aedeagus bifid, black, crescentic in lateral aspect ; 
basal apodeme one and a half times as long as aedeagus, brownish black, straight. Anterior 
paramere small, slender, pale yellowish brown, pubescent, attached to hypandrium. Posterior 
parameres (?) fusedtoform a large pale brown flat oval process, finely serrated along distal margin. 
Hypandrium brown, large, triangular, distally narrowly pointed and with a stout long submedian 
spine. Ventral fragma quadrate, anterolateral corners shortly projected forward, with deep 
median notch. Pl=o-5. p.f. = ABCdefgHIkKLMN. Egg-guide elongate elliptical, swollen 
dorsally, pale yellowish brown, distally and ventrally darker, apically narrowly rounded, with 
about twenty two marginal and two apical discal pointed brown teeth, upper two teeth being 
very long, and with a subterminal hair near apex. Basal isthmus very short and pale. 


Taplejung District, between Sangu and Tamrang, mixed plants by damp cliff 
in deep river gorge, c. 5,200’, 22.x1.1961, 2g, 1 29; between Sangu and Tamrang, 
mixed shrubs in deep gorge, c. 5,200’, Ig, I 9, x—x1.1961, (R.L.C.). 

Distribution : Nepal (new locality), Formosa. 


Drosophila (Scaptodrosophila) alternata de Meijere subgen. comb. n. 
(Text-figs. 196-200) 


Drosophila alteynata de Meijere, 1911 : 402 (Java). 
Drosophila (Pavadrosophila) alternata (de Meijere) Duda, 1924a : 207 (Java) ; Duda ; 1931 : 194 
(Sumatra) ; Mayer, 1935 : 467 (Sumatra). 


6. Body about 2 mm. Head as broad as thorax. Eyes dark red, large, finely pilose. 
Antennae with second segment orange-brown, conical ; third greyish black, basally orange- 
brown, oblong, twice as long as broad. Arista with about four dorsal and two ventral branches 
and a small fork. Palpus yellowish brown, with a few setae. Ocellar triangle medially black, 


DIPTERA FROM NEPAL 73 


marginally pale brown, well demarcated. Periorbit narrow, black, extending for upper half of 
front. Front flat, golden yellowish orange, anterior one-third paler and slightly narrower than 
length down middle, posteriorly half as broad as head width, with several frontal hairs and 
anterior periorbital microtrichia. Face grey, narrow. Carina high, long, greyish brown. 
Cheeks narrow, greyish yellow, about one-twelfth as broad as the greatest diameter of eye. 
Occiput yellowish brown, dark below. Anterior reclinate orbital small, one-fourth as long as 
proclinate. Vibrissae rather short ; second oral one-third as long as vibrissa ; succeeding orals 
shorter. 

Mesonotum greyish brown, with indistinct dark brownish patches. Scutellum yellowish brown, 
pollinose. Thoracic pleura dark yellowish brown, sternepisternum pale yellow. Humerals 
three, median longest. Prescutellar long, as long as anterior dorsocentrals. Anterior dorso- 
centrals one-third as long as posteriors ; distance between anterior and posterior dorsocentrals 
about one-fourth distance between anterior pair. Anterior scutellars divergent ; posteriors as 
long as anteriors, nearer to each other than to anteriors. Sterno-index about 0-3. Median 
sternopleural as long as posteriors. Legs yellow ; fore and mid metatarsi as long as the other 
segments together, hind metatarsi longer than rest of segments together. Preapicals on all legs ; 
apicals prominent on mid leg. Wings deeply fuscous, relatively narrow ; costa reaching end of 
M ; R43 nearly straight, apically curved to costa ; R,;, and M slightly divergent. C-index 
about 3:5 ; 4V-index about 1°5 ; 4C-index about 0-7 ; 5x-index about 2-0 ; Ac-index about 
1-8. Cr-bristles two, subequal ; C3-fringe on basal two-ninths. Halteres yellowish brown. 
Abdominal tergites yellowish grey, caudolaterally with a black medially interrupted laterally 
broadened cross-band. Abdominal sternites pale. 

Periphallic organs : Genital arch pale yellowish brown, setigerous and pubescent, ventrally 
gradually broadened, with high heel and rounded toe. Clasper low, semicircular, pale yellowish 
brown, relatively small, distal margin with a straight row of about eleven large long black teeth 
for entire length. Anal plate elliptical, yellowish brown, separated from genital arch, setigerous 
and pubescent; ventral corner with a tuft of short setae. Phallic organs : Aedeagus apparently 
bifid, oval in ventral aspect, pale yellowish orange, distally narrowing, basally rounded. Basal 
apodeme pale yellowish grey, straight, longer than aedeagus. PI = 0-8. Anterior paramere 
blade-like, pale yellowish orange, basally contiguous with aedeagus and hypandrium, apically 
pointed, laterally with two small pale depressions, which are probably sensory pits. Posterior 
paramere obscure. Ventral fragma pale yellowish grey, broad semicircular, distal margin 
straight, without median notch, with a pair of stout long yellowish brown submedian spines. 
p.f. = ABCDEFg,HIKLMN. 


Taplejung District, Sangu, yellowish blooms of cultivated Compositae (Guizotia 
abyssinica Cassini), c. 6,200’, 16-29.x.1961, I f (R.L.C.). 

Distribution : Nepal (new locality), Java, Sumatra. 

Strikingly like the original form except for larger 5x-index (slightly over I-o in 
original form) and distinctly interrupted abdominal cross-bands (obscurely inter- 
rupted in original form). Mayer (1935) described larva and puparium of this 
species from Sumatra. 


Subgenus DICHAETOPHORA Duda 
Dichaetophora Duda, 1940 : 19. 


Type-species : Drosophila aberrans Lamb, 1914 : 334 (Seychelles). 


Duda (1940) established this subgenus for one species, aberrans Lamb. Burla 
(1954) added two species, agbo Burla and suruk Burla from Africa, and two other 
species, magnidentata Lee, 1964, from Korea and raridentata Okada and Chung, 


74 T. OKADA 


were suggested by Lee (1964) to belong to this subgenus. Although this subgenus 
was originally characterized by only one pair of dorsocentrals, anterior pair being 
absent, some species have two pairs with anterior pair fine, and yet other species 
have both anterior and posterior pairs well developed. These species are character- 
ized in having very oblique eyes and ocellars inserted outside ocellar triangle, by 
which they can be assigned to Dichaetophora. Consequently, the occurrence of only 
one pair of dorsocentrals is an “‘archestinic’”’ character (Stenzel, 1963) which means a 
diagnostic character of a taxon occurring only in some individual units of that 
taxon (cf. p. 119). 
KEY TO SPECIES 


1 Antennae vertically erected from head, with the third segment remarkably swollen 
distally. Arista with only one branch below fork. Abdominal tergites almost 
entirely black ; 3 : . rotundicornis (p. 74) 

— Antennae recumbent on the fea ete fhe third See not remarkably swollen 
distally. Arista with two branches below fork. Abdominal tergites with distinct 
black cross-bands . : : : : : : : ‘ raridentata (p. 76) 


Drosophila (Dichaetophora) rotundicornis sp. n. 
(Text-figs. 207-213) 


3, 2. Body about 1-°8mm. Head slightly broader than thorax. Eyes dark red, triangular 
in lateral aspect, finely pilose and faint green iridescence. Antennae yellowish brown, large, 
upright ; second segment somewhat longer than broad ; third with thick long pubescence, 
distally dark brown and much swollen. Arista with about four dorsal and one ventral branches 
and a small terminal fork. Palpus yellow, tip rounded, with a prominent apical seta. Ocellar 
triangle black, marginally pale. Periorbit yellowish grey, not sharply demarcated from front, 
about two-thirds length of front. Front matt yellowish grey, flat, anterior one-third yellow, 
anteriorly narrower than median length ; posteriorly twice as broad as head width. A few 
frontal hairs present. Carina low, narrow, half of face, distally broadened. Face broad, grey- 
brown, much broadened below, buccal margin with a broad white transverse band, narrowly pale 
on carina and along eye margin. Clypeus dark brown. Cheeks yellowish white, somewhat 
pollinose, about one-third as broad as the greatest diameter of eye. Occiput greyish brown. 
Postverticals very long, longer than outer verticals. Ocellars long, inserted well outside ocellar 
triangle. Anterior reclinate orbital slightly before and outside proclinate, three-fifths as long as 
proclinate, two-thirds as long as posterior reclinate, thrice as distant from posterior reclinate as 
from proclinate. Second oral about half as long as vibrissa ; succeeding orals shorter. 

Mesonotum deep greyish brown, matt. Scutellum deep greyish brown, marginally and 
centrally somewhat pale. Thoracic pleura greyish orange, pollinose, with a dark brown longi- 
tudinal stripe on upper half. Three humerals, median longest. Acrostichal hairs in six rows. 
Anterior dorsocentrals two-thirds as long as posteriors ; distance between anterior and posterior 
dorsocentrals two-thirds distance between anterior pair. Anterior scutellars divergent, as long 
as posteriors, which are twice as distant from anteriors as from each other. Sterno-index about 
0:3. Legs dusty yellow, coxae paler, all metatarsi as long as the other segments together. Pre- 
apicals on all legs ; apicals prominent on mid leg. Wings hyaline ; veins yellowish brown ; 
crossveins not clouded ; costa paler and reaches the end of R,4;, ; R.,, gently curved to costa 
at tip; &,,, and M slightly divergent distally. C-index about 3:5; 4V-index about 2-4 ; 
4C-index about I-o ; 5x-index about 2-3 ; Ac-index about 2:0. Cr1-bristles two ; C3-fringe 
on basal three-sevenths. Halteres yellowish white. Abdominal tergites mostly black, some- 
what glossy ; 1T pale yellow at anterior corner. 


DIPTERA FROM NEPAL 


219 


Fics. 207-220. 207-213, Drosophila (Dichaetophora) votundicornis sp. nov. 207, ¢ head, 
lateral aspect ; 208, g head, dorsal aspect ; 209, g wing; 210, periphallic organs, 
lateral aspect ; 211, phallic organs, ventral aspect ; 212, aedeagus, lateral aspect, 
dextral side dorsal ; 213, Qegg-guide. 214-218, Drosophila (Dichaetophora) varidentata 
Okada and Chung, g. 214, head, lateralaspect ; 215, head, dorsal aspect ; 216, wing ; 
217, periphallic organs, lateral aspect ; 218, phallic organs, ventral aspect. 219-220, 
Drosophila (Drosophila) pentastriata sp.n. 3. 219, wing ; 220, head, dorsal aspect. 


73) 


76 T. OKADA 


Periphallic organs : Genital arch black, pubescent, medially and apically bare, with a few 
setae, ventrally conically projected, caudal margin rectangular above insertion of clasper. 
Clasper black, ovoid, with a row of about eleven black teeth along entire distal margin, median 
teeth smaller than upper and lower teeth. Anal plate white, pubescent and setigerous, narrow 
and long, dorsally somewhat narrowing. Phallic organs: Aedeagus robust, dark brown, 
apparently bifid laterally, quadrate in lateral aspect, caudodorsally round, upper margin truncate 
and with a few serrations. Basal apodeme shorter than aedeagus. PI = 1-5. Anterior 
paramere small, conical, contiguous to hypandrium, medially with a few sensilla. Ventral 
fragma black, semicircular, with a very deep and wide median notch and a median process 
projected from the bottom of the notch, laterally with submedian spines. p.f. = ab’CdeFg,HI 
kLMN. Egg-guide lobe deep brown, fusiform, with about ten marginal and four discal dark 
brown teeth, apical one largest and directed ventrally. Anal cercus of female yellow, elongate. 


Holotype g. East NepaL: Taplejung District, between Sangu and Tamrang, 
mixed plants by damp cliff in deep river gorge, c. 5,200’, 22.xi.1961 (R.L.C.), BMNH. 


Allotype 9. Same data as holotype (R.L.C.), BMNH. 


Paratypes : Taplejung District, 1 J, same data as holotype ; I g, 2 g, Sangu, 
mixed vegetation by stream in gully, c. 6,200’, xi.1961-1.1962 ; I g, 2 9, between 
Sangu and Tamrang, mixed plants by damp cliff in deep river gorge, c. 5,200’, 
1-11.1962 ; 4 9, I ex. (abdomen missing), river banks below Tamrang Bridge, c. 
5,500’, X-xi.1961. 1 9, Arun Valley, east shore of River Arun below Tumlingtar, 
c. 1,800’, I4—23.xil.1961 (R.L.C.), BMNH ; 1 gd and i 9 from between Sangu and 
Tamrang, deposited at the Tokyo Metropolitan University. 


Resembles D. (Dichaetophora) suruk Burla, 1954 (Ivory Coast, Africa), in having 
rather well developed anterior dorsocentrals, which are widely distant from posterior 
dorsocentrals, but differs from the latter in having only one ventral branch of arista 
(three in suruk), front narrower than length down middle (broader in s.), six rows of 
acrostichal hairs (four in s.), R,,; and M not sinuated (much sinuated in s.), and male 
anal plate narrow and long (broad in s.). 


Drosophila (Dichaetophora) raridentata Okada and Chung 
(Text-figs. 214-218) 


Drosophila (Drosophila) vavidentata Okada and Chung, 1960 : 28 (S. Korea). 
Drosophila varidentata Okada and Chung ; Wakahama, Kaneko, and Tokumitsu, 1963 : 94 

(Japan) ; Wakahama, 1964 : 56 (Japan). 

Drosophila (Dichaetophora?) vavidentata Okada and Chung ; Lee, 1964 : 108. 

3. Body about1:2mm. Head somewhat broader than thorax. Eyes oval, dark red, pilose. 
Arista with about five dorsal and two ventral branches and asmall fork. Palpus yellowish grey, 
slender, with a prominent apical seta. Ocellar triangle medially black. Periorbit yellowish 
grey, anteriorly somewhat swollen inward and one-third as long as front. Front dark grey, 
pale along median longitudinal line, anteriorly slightly narrowing and orange-yellow, with 
scattered frontal hairs, narrower than length down middle, posteriorly half as broad as head 
width. Clypeus grey. Face flat, grey, parallel-sided, buccal margin straight and narrowly 
white. Carina very small and low, yellowish grey, with a pair of small black spots apically. 
Occiput yellowish grey. Cheeks pale yellow, darker above, about one-fifth as broad as the 
greatest diameter of eye. Postverticals moderate in size, shorter than outer verticals. Anterior 
reclinate orbital two-thirds as long as proclinate, one-third as long as posterior reclinate, 


DIPTERA FROM NEPAL We 


twice as distant from posterior reclinate as from proclinate. Proclinate inside others. Second 
oral half as long as vibrissa. 

Mesonotum yellowish grey, matt. Scutellum yellowish grey, pollinose. Thoracic pleura 
paler, with two or three dark longitudinal stripes. Humeral callus paler. Humerals two, 
subequal. Acrostichal hairs in six rows. Anterior dorsocentrals two-thirds as long as posteriors ; 
distance between anterior and posterior dorsocentrals two-thirds distance between 
anterior pair. Anterior scutellars longer than posteriors, divergent ; posteriors thrice as distant 
from anteriors as from each other. Sterno-index about 0-4. Legs yellowish grey, coxa paler ; 
all metatarsi as long as three following segments together. Last tarsal segment large. Preapi- 
cals on all legs ; apicals on mid leg. Wings hyaline, slightly fuscous. R,;, gently curved to 
costa at tip ; R4,and M nearly parallel, costa reaches M, but weak beyond R,4;,;.  C-index about 
2°1; 4V-index about 2:5 ; 4C-index about 1-7 ; 5x-index about 4-5 ; Ac-index about 2:7. 
Cr-bristles two ; C3-fringe on basal half. Halteres yellowish grey, tip somewhat dark. Ab- 
dominal tergites yellowish brown, each with a distinct not medially interrupted, broad median 
black transverse band. 

Periphallic organs : Genital arch greyish brown, setigerous and pubescent, truncate below. 
Clasper large, quadrate, greyish brown, submedially with a row of about four very large pointed 
black teeth. Anal plate setigerous and pubescent, pale yellowish grey, oval, lower tip narrowing. 
Phallic organs : Aedeagus rod-shaped, yellowish grey, apically truncate ; apodeme one-third 
as long as aedeagus ; vertical rod short but thick. Anterior paramere large, oval, pale grey, 
basally contiguous with vertical rod, apically with a few sensilla. Ventral fragma pale grey, 
pubescent, with a deep median notch and a pair of finger-like submedian processes, apically with 
a pair of long submedian spines, lateral arms triangular. Decasternum triangular, large. 
PI = 3:0. p.f. = aBCDEfg,HIkimn. 


Taplejung District, Sangu, mixed vegetation by stream in gully, c. 6,200’, ix—x. 
for, to (k.L.C.). 

Distribution : Nepal (new locality), Japan, Korea. 

Differs somewhat from the original form (Korea) in having smaller body, longer 
anterior dorsocentral, and two ventral branches of arista (one in the original form). 
Originally described from g, the 2 was recorded by Wakahama, Kaneko, and 
Tokumitsu (1963) from Japan. 


Subgenus HIRTODROSOPHILA Duda 


Hirtodrosophila Duda, 1924a : 203. 


Type-species : Drosophila longecrinita Duda, 1924a: 204; I924b: 242 
(Formosa, New Guinea). 


Kry TO SPECIES 


t Arista with two branches below fork. Palpus with only one prominent apical seta. 
Cheeks broad, about two-fifths as broad as the greatest diameter of eye. Meso- 
notum glossy dark reddish brown. Acrostichal hairs in six rows. Preapicals on 
all legs. Third costal section with strong bristles. Postverticals longer than 
verticals : : ‘ ; i F : : ¢ ; tripartita (p. 78) 

— Arista with only one branch below fork. Palpus with a few prominent ventral setae. 
Cheeks narrow, about one-eighth as broad as the greatest diameter of eye. Meso- 
notum dark chestnut-brown. Acrostichal hairs in eight rows. Preapicals on mid 
and hind legs. Third costal section without strong bristles. Postverticals much 
shorter than inner verticals . : ‘ ‘ : : . limbicostata (p. 79) 


78 T. OKADA 


Drosophila (Hirtodrosophila) tripartita sp. n. 
(Text-figs. 262-267, p. go) 


3, 2. Body about 2-2 mm. Head as broad as thorax. Eyes deep red, fusiform in lateral 
aspect, with thick pilosity. Antennae orange-yellow ; third segment fuscous at tip, oval, much 
broader than second segment, slightly longer than broad. Arista with about four dorsal and two 
ventral branches and a large terminal fork. Palpus yellowish orange, with an apical seta. 
Ocellar triangle small, deep reddish brown. Periorbit glossy grey-brown, anteriorly somewhat 
curved inward, posteriorly slightly broadened and paler. Front orange-grey, paler anteriorly, 
nearly parallel-sided, frontal triangle fuscous, anteriorly broader than length down middle, 
posteriorly about four-sevenths as broad as head width. A few frontals present. Clypeus dark 
brown. Carina long, low, broadened below to reach buccal margin. Cheeks orange-yellow, 
broad, about two-fifths as broad as the greatest diameter of eye. Occiput yellowish brown, 
ventromedially fuscous. Postvertical very long, much longer than outer vertical. Anterior 
reclinate orbital about one-third as long as posterior reclinate, slightly nearer to proclinate than 
to posterior reclinate. Proclinate slightly shorter than posterior reclinate, inside others. Only 
one pair of long vibrissae, second oral one-fourth as long as vibrissa. 

Mesonotum glossy dark reddish brown. Scutellum dark reddish brown, unicolorous. Thor- 
acic pleura paler than mesonotum. Two humerals, lower longer. Acrostichal hairs in six rows. 
No prescutellars. Anterior dorsocentrals nearly half as long as posteriors ; distance between 
anterior and posterior dorsocentrals about five-ninths distance between anterior pair. Anterior 
scutellars divergent, longer than posteriors, which are nearer to each other than to anteriors. 
Sterno-index about 0-5 ; sternopleurals situated close to each other. Legs yellow ; all meta- 
tarsi as long as other segments together. Male fore leg without sex-combs ; mid leg without 
hooked scaly bristles. Preapicals on all legs ; apicals prominent on mid leg. Wings hyaline, 
crossveins clear. Costa reaching M. R,;, gently convex at middle, gently curved to costa at 
tip. R,,; and M parallel. C-index about 3:8; 4V-index about 1:6; 4C-index about 0°6 ; 
5x-index about 1:7 ; Ac-index about 2:1. C1-bristles two ; C3-fringe on basal three-eighths. 
Halteres yellowish orange, stalk paler. Abdominal tergites mostly dark brownish black, some- 
what glossy ; 1T and 2T anteriorly yellowish orange ; terminal T greyish white. 

Periphallic organs : Genital arch pale yellowish orange, setigerous and pubescent, dorsally 
gently narrowing, medially swollen, ventrally tapering triangularly. Clasper dark brown, 
oblong, with a sinuous row of about thirteen black pointed teeth occupying nearly entire length 
of outer margin, and with a few short fine setae on inferior surface of distal half. Anal plate 
pale yellowish brown, fusiform, separated from genital arch, pubescent and shortly setigerous, 
lower tip pointed ventrally and with about five strong black bristles ina clump. Phallic organs 
yellowish brown. Aedeagus rod-shaped, hairy, apically somewhat broadened, proximally 
much swollen. Anterior paramere slender, curved, tapering, orange-brown, basally contiguous 
with hypandrium. Posterior parameres fused to form a large quadrate flap, distal and proximal 
margins medially concave. Ventral fragma triangular, pale brown, with deep median notch. 
PI = 2:3. p.f. = aBcDEf,gHiKIMN. Egg-guide lobe triangular, proximally narrowing, 
yellowish orange, with about seventeen marginal and twenty-two discal short scattered yellowish 
orange bristles on distal half. Subterminal hair short but distinct. Basal isthmus short. 


Holotype g. East NepaL: Taplejung District, Sangu, mixed vegetation by 
stream in gully, c. 6,200’, xi.1961-1.1962 (R.L.C.), BMNH. 


Allotype 2. Same locality as holotype, ix—x.1961 (R.L.C.), BMNH. 
Paratypes : Taplejung District, I J, same data as holotype ; 1 g, between Sangu 
and Tamrang, mixed shrubs in deep gorge, c. 5,200’, x—xi.1961 ; I g, above Sangu, 


mixed vegetation in dried-up ravine, c. 6,800’, 16.11.1962 (R.L.C.), BMNH, (2 3), 
and at the Tokyo Metropolitan University (1 g, between Sangu and Tamrang). 


DIPTERA FROM NEPAL 79 


Closely similar to D. (H.) denticeps Okada and Sasakawa, 1956 (Japan), especially 
in male periphallic organs and female egg-guide, but differing from it and also in 
general from species of the subgenus in having two ventral branches of arista and 
well developed preapicals on all legs. This species is, consequently, only tentatively 
included in the subgenus Hirtodrosophila. 


Drosophila (Hirtodrosophila) limbicostata sp. n. 
(Text-figs. 221-225) 


9. Body about 2 mm. Head slightly broader than thorax. Eyes dark purplish brown, 
large, with thick pilosity. Antennae with second segment conical, yellowish brown, medio- 
laterally dark ; third black, large, and rounded. Arista with about two dorsal and one ventral 
short branches and a fine terminal fork. Palpus yellowish orange, thick, tip rounded, with a 
few short black apical and ventral setae. Ocellar triangle large, black, grey pollinose, convex. 
Periorbit posteriorly somewhat narrowing, black, grey pollinose, anteriorly nearly reaching 
anterior margin of front. Front velvety black, nearly parallel-sided, anterior half deep orange- 
brown anteriorly as broad as length down middle, posteriorly two-fifths as broad as head width. 
Clypeus brown. Face grey, concave. Carina grey, narrow, low, less than half length of face. 
Cheeks narrow, yellow, about one-eighth as broad as the greatest diameter of eye ; postgena 
black. Occiput black. Postverticals short. Anterior reclinate orbital slightly shorter than 
proclinate, which is as long as posterior reclinate. Three orbitals nearly equally distant from 
eye margin, anterior reclinate equidistant from proclinate and from posterior reclinate. 

Mesonotum dark chestnut-brown, somewhat glossy, with fine yellow punctures. Humeral 
callus pale yellowish grey. Scutellum dark yellowish brown, medially and anterolaterally 
blackish brown. Thoracic pleura pale brown. Three humerals, uppermost shortest. Acro- 
stichal hairs in eight rows. Prescutellars fine. Anterior scutellars divergent, slightly longer 
than posteriors, which are one and a half times as distant from anteriors as from each other. 
Postscutellum chestnut-black. Sterno-index about 1-o ; median sternopleural as long as 
others. Anterior dorsocentrals half as long as posteriors ; distance between anterior and 
posterior pair less than one-third distance between anterior pair. Legs yellow, last tarsal seg- 
ment dark, claw large. All metatarsi subequal to next segments together. Preapicals on 
mid and hind legs ; apicals prominent on midleg. Wingssomewhat fuscous, broad, tip rounded. 
Ro+3 apically gently curved to costa; R,,;, and M parallel. C-index about 2-6; 4V-index 
about 1-5 ; 4C-index about 0-9 ; 5x-index about 1:6 ; Ac-index 2-3. Cr-bristles two, fine, 
subequal ; C3-fringe absent, strong costal bristles ending before tip of R,,,. Halteres yellow. 
Abdominal tergites : 1T yellowish grey, others brownish black, each medioanteriorly with a 
small triangular yellow spot. Abdominal sternites paler. 

Egg-guide lobe fusiform, apically pointed and somewhat curved downward, with about 
eighteen marginal and five apicodiscal pale brown teeth, nine upper marginal ones arranged in a 
compact straight row, apical tooth largest. Subterminal hair distinct. Basal isthmus yellow, 
narrow, straight, about one-fourth length of lobe. 


Holotype 9. East NEpAL: Taplejung District, between Sangu and Tamrang, 
mixed plants by damp cliff in deep river gorge, c. 5,200’, 22.x1.1961 (R.L.C.), BMNH. 


Peculiar among the family Drosophilidae in having strong costal bristles ending 
before end of R,,3;. Somewhat similar to D. (H.) Mirticornis de Meijere, 1914, 
(Java) in having very narrow cheeks and fuscous wings, but differs from it in short 
branches of arista, pubescent eyes, and brownish black abdominal tergites. 


80 


T. OKADA 


FIGS. 221-231. 221-225, Drosophila (Hirtodrosophila) limbicostata sp. n., 9. 221, head, 


lateral aspect ; -222, head, dorsal aspect ; 223, scutellum; 224, wing; 225, egg- 
guide. 226-231, Drosophila (Sophophora) coei sp. n. 226, 3 head, lateral aspect ; 


227, 6 head, dorsalaspect ; 228,f wing; 229, periphallic organs, lateral aspect ; 230, 
phallic organs, ventral aspect ; 231, 2 egg-guide. 


DIPTERA FROM NEPAL 81 


Subgenus DORSILOPHA Sturtevant 
Dorsilopha Sturtevant, 1942 : 28. 


Type-species : Drosophila buscku Coquillett, Igor : 16 (N. America). 


Drosophila (Dorsilopha) busckii Coquillett 
Drosophila busckwi Coquillett, rg01 : 16 (N. America). 


Taplejung District, Sangu, from rotten Naspati fruits in traps, c. 6,200’, 8-31.x. 
1961, 15 g, 12 Y; ibid., from rotten bananas in traps, c. 6,200’, x—xil.I96I, 2 @ ; 
below Sangu, by stream in shady ravine, c. 6,000’, 30.x.19g61, Ig ; river banks below 
Tamrang Bridge, c. 5,500’, x—xi.1961, I d (R.L.C.). 

Distribution : Nepal (new locality), India, Sumatra, Formosa, China, Japan, 
Korea, Bonin Is., Australia, Africa, Europe, N. America, S. America. 


Subgenus SOPHOPHORA Sturtevant 
Sophophora Sturtevant, 1939 : 139. 


Type-species : Drosophila melanogaster Meigen, 1830 : 85 (Europe). 


KrEy TO SPECIES 


1 Anterior reclinate orbital minute, one-fifth or less as long as posteriors. ¢ fore legs 
without sex-combs. Mesonotum strongly convex . : mommai-group 2 
— Anterior reclinate orbital usually large, about one-third or more as long as posteriors. 
3 fore legs with sex-combs. Mesonotum usually not very convex 


melanogaster-group 4 

2 Palpus with only one prominent apical seta. Crossveins clear. Head much broader 
than thorax . . coei (p. 82) 

— Palpus with a few prominent ventral setae. Crossveins clouded. " Head as broad as 
thorax c : ; ; , : : ‘ 3 

3. Palpus black. Front glossy Eeowa. " Median sternopleural fine. Third abdominal 
tergite entirely black . - . xzonaria (p. 83) 

— Palpus yellow. Front matt orange. Median sternopleural as long as anteriors. 
Third abdominal tergite not entirely black . : r . serriflabella (p. 84) 
4 Palpus with a few prominent ventral setae : Z : . melanogaster (p. 92) 
— Palpus with a prominent apical seta : : - - - : : : 5 
5 6 wings distally with a large black patch . : : - . : : ; 6 
— 6 wings without black patch distally 8 

6 Large distal black patch of $ wing not extending ie the wing tip. Small black 
patches at wing base . é pulchrella (p. 89) 

— Large distal black patch of 3 me extending to the mie ie No black patches at 
wing base : 7 

7 Large distal black patch of 3} wane Grescentic, extending below M. ‘Anterior reclinate 
orbital about five-eighths as long as posteriors é - : tristipennis (p. 89) 

— Large distal black patch of J wing oval, not extending below M. Anterior reclinate 
orbital about half as long as posteriors . : : : : nepalensis (p. 91) 


8 Two humerals subequal in length c : : : : : : 
Upper humeral longer than lower. ¢ sex-combs large, longitudinal. Second oral 
nearly as long as vibrissae  . : : 5 : . c : : c II 


82 T. OKADA 


9 6 sex-combs larger, in two eee sets on metatarsus. Second oral nearly as long as 
vibrissae  . é : bipectinata (p. 95) 
-— 6 sex-comb eroled HOSTSSE. Gerond oral Sash Bhortee than vibrissae  . ; Io 
10 Acrostichal hairs in six rows. Anterior reclinate orbital about half as long as 
posteriors, one-third as long as proclinate. Distance between anterior and post- 
erior dorsocentrals about two-fifths distance between anterior pair immacularis (p. 87) 
- Acrostichal hairs in eight rows. Anterior reclinate orbital one-third as long as 
other orbitals. Distance between anterior and posterior dorsocentrals about half 
distance between anterior pair : takahashii (p. 92) 
11 Front nearly parallel-sided. Periorbit Beret Pea past erioely narrowing, 
anteriorly curved inward. Anterior scutellars convergent. : . kikkawai (p. 92) 
— Front considerably narrowed anteriorly. Periorbit black, narrow, anteriorly not 
curved inward. Anterior scutellars divergent ; : .  trapezifrons (p. 93) 


THe MOMMAT Group 


3 foreleg without sex-comb. Mesonotum highly convex. Anterior reclinates 
orbital minute. Only one prominent oral. Anterior paramere of phallic organs 
large, hairy, usually with a longitudinal row of sensilla. Aedeagus hairy, often 
with a pair of basal elongate processes. 


Drosophila (Sophophora) coei sp. n. 
(Text-figs. 226-231) 


3, 2. Body about 1:8 mm. Head shghtly broader than thorax. Eyes deep purplish red, 
fusiform, relatively small, pilose. Antennae with second segment yellowish orange, third 
greyish orange and triangularly pointed. Palpus black, slender, with a long terminal seta. 
Ocellar triangle deep reddish brown.  Periorbit glossy greyish white, somewhat convex, broad, 
three-fourths as long as front. Front greyish white, glossy. Carina high, broadened below. 
Clypeus brown. Cheeks yellowish brown, about two-sevenths as broad as the greatest diameter 
of eye. Occiput yellowish brown, centrally somewhat fuscous. Postverticals rather short. 
Anterior reclinate orbital minute, outside and slightly posterior to proclinate, somewhat nearer 
to proclinate than to posterior reclinate. Proclinate about three-fourths as long as posterior 
reclinate. Vibrissae long ; second oral half as long as vibrissa, others shorter. 

Mesonotum glossy brown or yellowish brown, sometimes deep brown, strongly convex, 
laterally with a dark longitudinal stripe. Scutellum glossy brown or deep brown, unicolorous. 
Thoracic pleura pale, glossy, dark brown below. Humeral callus mostly dark brown. Only 
one long lower humeral, upper one very fine. Acrostichal hairs in six rows. Anterior dorso- 
centrals three-fifths as long as posteriors ; distance between anterior and posterior dorsocentrals 
about half distance between anterior pair. Anterior scutellars parallel or convergent, as long as 
posteriors, which are nearer to each other than to anteriors. Sterno-index about 0-4. Legs 
yellow, glossy, relatively slender ; tarsi fuscous ; coxae white, fore coxa sometimes black (in 
darker specimens) ; all metatarsi subequal to the next segments together. Preapicals on all 
legs ; apicals prominent on fore and mid legs. Wings relatively large, somewhat fuscous, veins 
yellow, crossveins clear. Costa reaching M, R,,, nearly straight and apically curved to costa ; 
R,,, and M somewhat convergent distally. C-index about 2-7 ; 4V-index about 1:8; 4C- 
index about I:o0 ; 5x-index about 1:5 ; Ac-index about 2-5. Cz1-bristles two ; C3-fringe on 
basal one-fourth. Halteres yellow. Abdomen slender and dorsally convex, tergites mostly 
glossy black. Female cercus yellow or yellowish brown, triangular in dorsal aspect. 

Periphallic organs : Genital arch black, pubescent and setigerous, broad, ventrally triangular, 
caudal margin with a large acute triangular projection below anal plate, deeply incised below 
the projection. Anal plate black, relatively small, narrow fusiform, setigerous and pubescent, 
separated from genital arch, ventrally exceedingly narrowing and with a few stout short black 


DIPTERA FROM NEPAL 83 


setae. Clasper large, black, triangular, with a straight row of about ten pointed black teeth 
occupying dorsal two-thirds of distal margin. Phallic organs mostly deep brownish black. 
Aedeagus elongate fusiform, hairy, pale yellow, ventrally with a pair of elongate apically bifid 
gently curved projections. Apodeme straight, about twice as long as aedeagus. Anterior 
paramere elongate, medially curved, proximally broadened, distally narrowing and thickly 
hairy, with a longitudinal row of several sensilla on median two-thirds, and basally with a large 
pointed ventrally directed processes. Posterior parameres fused to form a pale large triangular 
flap, laterally attached to the lateral processes of ventral fragma. PI =o0.5. p.f. = Ab’CDE 
fgHiKLMN. Egg-guide lobe orange-yellow or dark orange, elongate oval, distally narrowly 
rounded, proximally tapering, with about thirteen marginal and a discal pointed orange-yellow 
teeth, ultimate marginal tooth longest. Subterminal hair distinct. Basal isthmus yellow, 
straight, about one-fifth as long as lobe. 

Holotype g. East NepaL: Taplejung District, between Sangu and Tamrang, 
mixed plants by damp cliff in river gorge, c. 5,200’, i-1.1962 (R.L.C.), BMNH. 

Allotype 2. Taplejung District, Sangu, mixed vegetation in deep gully, c. 6,200’, 
2.1-13.1i1.1962 (R.L.C.), BMNH. 

Paratypes : 64, same data as holotype ; 10 2, same data as allotype ; Taplejung 
District, 1 J, 2 2, Sangu, by stream in gully, c. 6,200’, ix-x.1961 ; 1 4, ibid., clearing 
in small wood, c. 6,200’, 25.x1.1961 ; 7 3, 20 9, ibid., mixed vegetation by stream in 
gully, c. 6,200’, xi.1g61-i.1962 ; 2 g, between Sangu and Tamrang, collected with 
holotype ; 1 g, 1 9, ibid., dense vegetation in tree shade by hill stream, c. 5,500’, 
23.X.1961 ; I, ibid., mossy ground under bushes by hill stream, c. 5,500’, 20.x.1961; 
I g, 5 2, above Sangu, old mixed forest, c. 6,200’, 25-28.x.19g61 ; I 9, ibid., damp 
evergreen oak forest, c. 9,200’, 2.26.x1.1961 ; 4, 17 9, ibid., mixed vegetation in 
dried-up ravine, c. 6,800’, 16.11.1962 ; I g, 2 9, below Sangu, by stream in shady 
ravine, c. 6,000’, 30.x.1961 ; 2 9, ibid., mixed vegetation on sheltered slopes above 
river, C. 4,000’, 3.i.1962 ; I g, I 9, north of Sangu, dry grass above river bank, 
c. 5,000’, 5.1.1962 ; Ig, I 9, river banks below Tamrang Bridge, c. 5,500’, x—xi. 1961 
ih...C.), BMNEH. 

Closely resembles D. (S.) mommai Takada and Okada, 1960 (Japan), in having 
highly convex mesonotum and abdominal tergites, and in the general features of 
phallic and periphallic organs, but distinguished from it in having genital arch less 
prominently narrowing below, lateral lobes of aedeagus apically bifid (simple in 
mommat), discal teeth of egg-guide only one (several in m.), and posterior scutellars 
as long as anteriors and not upright (short and upright in m.). 


Drosophila (Sophophora) zonaria sp. n. 
(Text-figs. 232-237) 


3, 9. Body about 1:7 mm. Head as broad as thorax. Eyes dark purplish red, thickly 
pilose, somewhat fusiform in lateral aspect. Antennae with second segment orange-yellow ; 
third yellowish grey, basally pale, and nearly twice as long as broad. Arista with about six 
dorsal and three ventral branches and a small terminal fork. Palpus black, slender, with a few 
strong ventral black setae. Ocellar triangle glossy black. Periorbit glossy pale brown, anteri- 
orly broader and curved inward, not reaching anterior margin. Front glossy greyish brown, 
parallel-sided, anterior one-third yellowish, anteriorly one and a half times as broad as length 


down middle, posteriorly half as broad as head width. Face white, glossy. Carina white, high, 


84 i OKADA 


broad, broader below. Clypeus white, laterally black. Cheeks broad, white, about one-fourth 
as broad as the greatest diameter of eye. Occiput yellowish brown, ventromedially black. 
Anterior reclinate orbital very fine, equidistant from proclinate and from posterior reclinate, 
slightly outside proclinate ; proclinate about two-thirds as long as posterior reclinate. Vibrissae 
long ; second oral fine, two-fifths as long as vibrissa ; succeeding orals finer. Postverticals 
moderate in size. 

Mesonotum and scutellum glossy grey-brown, unicolrous. Thoracic pleura pale yellowish 
white. Two humerals, upper half as long as lowers. Acrostichal hairs in six rows. Anterior 
dorsocentrals half as long as posteriors ; distance between anterior and posterior dorsocentrals 
one-third distance between anterior pair. Anterior scutellars divergent, longer than posteriors, 
which are nearer to each other than to anteriors. Sterno-index 0-4. Legs yellow, tibiae and 
tarsi slightly fuscous ; all metatarsi subequal to next three segments together. Preapicals on 
all legs ; apicals prominent on mid leg. Wings somewhat fuscous ; crossveins clouded ; costa 
reaching F,;,, strong bristles on costa relatively fine. R,,,and M distally somewhat convergent. 
C-index about 2-7 ; 4V-index about 2-1 ; 4C-index about 1-1 ; 5x-index about 0:8 ; Ac-index 
about 2:6. Cr1-bristles two ; C3-fringe on basal three-tenths. Halteres yellowish brown. 
Abdominal tergites yellowish-brown, glossy, each with a caudal black medially contiguous 
cross band ; 3T nearly entirely black (thus the specific name). 

Periphallic organs pale brown. Genital arch pubescent and setigerous, much narrowed 
dorsally, gently projected at middle caudally, ventrally narrowing and rectangularly curved 
caudally at tip. Clasper large, triangular, distally broadened, with a nearly straight row of 
about twelve pointed black teeth, occupying almost entire distal margin. Anal plate separated 
from genital arch, elliptical, pubescent and setigerous, with short hairs at ventral tip. Phallic 
organs yellowish orange. Aedeagus short, thickly hairy, apically pointed, proximally much 
broadened ; apodeme slender, straight, nearly twice as long as aedeagus. Anterior paramere 
orange, hairy, slender. Posterior paramere absent? Ventral fragma quadrate, large, with 
lateral arms conically projected ventrally, without median notch. PI = 0-5. p.f. = ABcDef, 
g ,HiKLMN. Egg-guide lobe orange-brown, well sclerotized, fusiform, with about thirteen 
marginal and three large brownish discal teeth ; subterminal hair distinct. Basal isthmus 
about one-fourth as long as lobe. 


Holotype g. East NEPAL : between Sangu and Tamrang, mixed shrubs in deep 
gorge, C. 5,200’, x-xi.1961 (R.L.C.), BMNH. 


Allotype 2. Same data as holotype (R.L.C.), BMNH. 


Paratypes : I g, I 9, Sangu, mixed vegetation by stream in gully, c. 6,200’, 
Xi.I9Q61-1.1962 ; Ig, 3 2, same data as holotype ; 1 Q, river banks below Tamrang 
Bridge, c. 5,500’, x-xi.1961 ; I g, between Sangu and Tamrang, mixed plants by 
damp cliff in deep river gorge, c. 5,200’, 22.xi.1961 (R.L.C.), BMNH. 

Somewhat resembles D. (S.) coed, in having broad front, minute anterior reclinate 
orbitals, broad cheeks, hairy aedeagus, and hairy anterior paramere, but differs 
from it in having paler body, white face, unicolorous mesonotum, clouded wings, 
and clouded crossveins. It differs also in the details of genital organs. 


Drosophila (Sophophora) serriflabella sp. n. 
(Text-figs. 238-243) 


3,2. Body about 2-5 mm. Head as broad as thorax. Eyes deep purplish red, fusiform at 
lateral aspect, relatively small, pilose. Antennae with second segment dark yellowish orange ; 
third greyish brown, oval, as broad as long, basally paler. Arista with about five dorsal and 
three ventral branches and a small terminal fork. Palpus narrow, yellowish brown, with a 


DIPTERA FROM NEPAL 


Fics. 232-243. 232-237, Drosophila (Sophophora) zonaria sp. n. 232, ¢ head, lateral 
aspect ; 233,¢head,dorsalaspect ; 234,g wing; 235,9egg-guide; 236, periphallic 
organs, lateral aspect ; 237, phallic organs, ventral aspect. 238-243, Drosophila 
(Sophophora) serrifiabella sp. n. 238, g head, lateral aspect; 239, dg head, dorsal 
aspect ; 240, g wing; 241, periphallic organs, lateral aspect ; 242, phallic organs, 
ventral aspect ; 243, 9 egg-guide. 


85 


86 T. OKADA 


long apical and two ventral black setae. Ocellar triangle small, glossy brown. Periorbit glossy 
grey-brown, broad, posteriorly pale, anteriorly rounded, three-fourths as long as front. Front 
matt orange-brown, broad, nearly parallel-sided, anteriorly one and a half times as broad as 
length down middle, posteriorly three-fifths as broad as head width. Face greyish yellow, con- 
cave, whitish along eye margin. Carina high, short, half as long as face. Clypeus grey-brown. 
Cheeks yellowish brown, broad, about two-sevenths as broad as the greatest diameter of eye. 
Occiput yellowish brown, ventromedially orange. Postverticals moderate in size. Anterior 
reclinate orbital very fine, outside proclinate, twice as distant from proclinate as from posterior 
reclinate, which is one and a half times as long as proclinate. Vibrissae long, second oral one- 
third as long as vibrissa. 

Mesonotum glossy yellowish brown, strongly convex. Scutellum glossy yellowish brown. 
Thoracic pleura paler. Two humerals, upper much shorter, two-fifths as long as lower. Acro- 
stichal hairs in six rows. Anterior dorsocentrals half as long as posteriors ; distance between 
anterior and posterior dorsocentrals about half distance between anterior pair. Anterior 
scutellars divergent, slightly longer than posteriors ; posteriors nearer to each other than to 
anteriors. Sterno-index about 0:5 ; median sternopleural as long as anteriors. Legs fuscous 
yellow ; femora paler above. All metatarsi as long as other segments together. Preapicals on 
all legs ; apicals on fore and mid legs. Wings relatively large, tip slightly pointed, somewhat 
fuscous along costa ; crossveins clouded. #,;, medially gently convex anteriorly, apically 
strongly curved to costa; R,,;; and M parallel. C-index about 2-4; 4V-index about 1°5 ; 
4C-index about I-o ; 5x-index about 1-7 ; Ac-index about 3:0. Cr1-bristles two ; C3-fringe on 
basal three-eighths. Halteres dark yellowish brown. Abdominal tergites glossy yellowish brown 
with sharply demarcated caudal black bands, which medially jut out triangularly in front. 
The triangular patches increasingly larger on the more caudal tergites ; 1T yellowish brown ; 
6T mostly black. 

Periphallic organs yellowish brown. Genital arch large, distally wider, setigerous and pubes- 
cent, ventrally curved posteriorly, subapically with deep incision on caudal margin, with a low 
triangular process above insertion of clasper. Clasper small, triangular, distal margin broad and 
straight, with a row of about ten long black teeth, occupying nearly entire distal margin. Anal 
plate relatively small, pubescent and setigerous, fusiform. Phallic organs : Aedeagus short, 
thickly hairy. Apodeme straight, twice as long as aedeagus. PI =—o-4. Anterior paramere 
small, conical, medially with a few sensilla. Posterior parameres fused to from a large fan- 
shaped marginally finely serrated yellowish orange lobe. Hypandrial plate large, conically 
projected posteriorly. Ventral fragma triangular. p.f. = ABCdefgHiKLMN. Egg-guide 
lobe brownish black, proximally tapering, distal margin broadly truncate, with about twelve 
marginal and two discal large brown teeth. Subterminal hair prominent. Basal isthmus black, 
about one-fifth as long as lobe. 


Holotype g. East NEepaL: Taplejung District, between Sangu and Tamrang, 
mixed plants by damp cliff in deep river gorge, c. 5,200’, x—xi.1961 (R.L.C.), BMNH. 

Allotype 9. Taplejung District, Sangu, mixed vegetation by stream in gully, 
c. 6,200’, x1.1961-1.1962 (R.L.C.), BMNH. 

Paratypes : Taplejung District, 1 g, same data as holotype ; I 9, between 
Sangu and Tamrang, mixed shrubs in deep gorge, c. 5,200’, x—xi.1961 ; 24, between 
Sangu and Tamrang, mixed plants by damp cliff in deep river gorge, c. 5,200’, 
i-li.1962 (R.L.C.), BMNH, and the Tokyo Metropolitan University (1 f from between 
Sangu and Tamrang). 

Closely resembles D. fruhstorferi Duda, 1924 (India), especially in having broad 
front, only one long oral, clouded crossveins, and medially triangularly projected 
black caudal bands of abdominal tergites, but differs from it in having three ventral 
branches of arista (only one in fruhstorfert). 


DIPTERA FROM NEPAL 87 


THe MELANOGASTER Group 
THe SUZUKII SupcGroup 
Drosophila (Sophophora) immacularis sp. n. 
(Text-figs. 244-250) 

3. Body about 2mm. Head as broad as thorax. Eyes dark red. Antennae with second 
segment yellowish orange, third yellowish grey, one and a half times as long as broad. Arista 
with about four dorsal and three ventral branches and a large terminal fork. Palpus yellow, 
with an apicalseta. Ocellar triangle dark reddish brown. Periorbit greyish brown, posteriorly 
narrowing, anteriorly rounded, not reaching anterior margin of front. Front somewhat glossy 
greyish brown, anteriorly orange and as broad as length down middle, posteriorly broadened and 
about four-ninths as broad as head width. Clypeus pale brown. Face greyish white, parallel 
sided. Carina high, long, anteriorly somewhat broadened. Cheeks yellowish orange, one-sixth 
as broad as the greatest diameter of eye. Occiput yellowish brown, fuscous below. Anterior 
reclinate orbital about half posterior reclinate and two-thirds as long as proclinate, twice as far 
from posterior reclinate as from proclinate. Proclinate inside others. Postverticals long. 
Second oral half as long as vibrissa, succeeding orals shorter. 

Mesonotum dark yellowish brown, somewhat glossy, medially with a darker longitudinal 
stripe. Scutellum greyish brown, slightly glossy. Thoracic pleura paler, unicolorous. Two 
long subequal humerals. Acrostichal hairs in six rows. Anterior dorsocentrals half as long as 
posteriors ; distance between anterior and posterior dorsocentrals about two-fifths distance 
between anterior pair. Anterior scutellars convergent, as long as posteriors, which are nearer to 
each other than to anteriors. Sterno-index about 0-7. Median sternopleural slightly shorter 
than anteriors. Legs yellow ; ¢ fore tarsus with a terminal transverse sex-comb composed of 
five black bristles ; second tarsal segment with a similar sex-comb of two bristles. All meta- 
tarsi subequal to the next three segments together. Male mid leg with black hooked scaly 
bristles, four at the apex of tibia and one at the proximal end of metatarsus. Preapicals on all 
legs ; apicals on fore and mid legs. Wings hyaline, crossveins clear ; costa weak beyond F,4,;; 
R,4, apically weakly curved to costa ; R,,;, and M parallel. C-index about 2-9 ; 4V-index 
about 2:2 ; 4C-index about 2:5 ; 5x-index about 2:2 ; Ac-index about 2:5. C1-bristles two, 
long ; C3-fringe on basal four-ninths. Halteres yellowish orange. Abdominal tergites yellow- 
ish brown, each with a broad black medially entire cross-band ; 1T pale. 

Periphallic organs : Genital arch elongate, fusiform, medially somewhat constricted, dorsal 
half black, dorsal end narrowing, ventral half yellowish grey, posteriorly triangularly projected, 
setigerous on posterior margin and ventral tip. Claspers two. Primary clasper large, brown 
lower tip recurved, with a row of about ten long black bristles on distal margin ; secondary 
clasper quadrate, situated above primary clasper, with a row of about eight large black teeth 
along distal margin. Anal plate setigerous, fusiform, ventrocaudally narrowly projected. 
Phallic organs : Aedeagus pale yellowish grey, rod-shaped, subapically pubescent, apically 
curved and pointed dorsally ; apodeme black, straight, curved and pointed dorsally ; apodeme 
black, straight, slightly longer than aedeagus. PI = 1-0. Anterior paramere black, rod-shaped, 
basally curved to attach the base of aedeagus, apically narrowly pointed and curved, subapically 
with a few sensilla. Posterior paramere pale yellowish grey, nearly as long as anterior paramere, 
apically pointed, basally bilobed to attach aedeagus by one arm and to ventral fragma by 
another. Ventral fragma quadrate, distal margin nearly straight, with a pair of short submedian 
spines, which are close to each other. p.f. = ABcDEfGHIKLMN. 


Holotype g. East NepaL: Taplejung District, between Sangu and Tamrang, 
shrubs by path, c. 5,800’, 6.xi1.1961 (R.L.C.), BMNH. 

Closely allied to D. (S.) suzuki (Matsumura, 1931) (China and Japan), especially 
similar in phallic and periphallic organs, but differs in having $ wings without black 
patch and aedeagus distally pubescent. 


88 


T. OKADA 


cee 


-——— ——___ ______, 


Fics. 244-254. 244-250, Drosophila (Sophophora) immacularis sp.n., 3. 244, head, lateral 


aspect ; 245, head, dorsal aspect ; 246, sex-combs on proximal two tarsal joints of 
fore leg ; 247, hooked scaly bristles on tibia and metatarsus of mid leg ; 248, wing ; 
249, phallic organs, ventral aspects ; 250, periphallic organs, lateral aspect. 251-254, 
Drosophila (Sophophora) pulchrella Tan, Hsu, and Sheng, g. 251, periphallic organs, 
lateral aspect ; 252, phallic organs, ventral aspect ; 253, left : hooked scaly bristles 
on tibia and metatarsus of mid leg ;_ right : sex-combs on proximal two tarsal joints of 
fore leg ; 254, wing. 


DIPTERA FROM NEPAL 89 


Drosophila (Sophophora) pulchrella Tan, Hsu, and Sheng 
(Text-figs. 251-254) 


Drosophila (Sophophora) pulchrella Tan, Hsu, and Sheng, 1949 : 198 (China) ; Okada, 1956 : 106 

(Japan). 

3, @. General features as described by Tan, Hsu, and Sheng (1949) and redescribed by Okada 
(1956). Head as broad as thorax. Ocellar triangle grey. Periorbit grey, glossy, constricted 
near caudal end. Front yellowish grey, slightly narrowing anteriorly, where it is as broad as 
length down middle, posteriorly four-ninths as broad as head width. Face white. Anterior 
dorsocentral half as long as posteriors. Anterior scutellars convergent, as long as posteriors, 
which are slightly nearer to each other than to anteriors. All metatarsi subequal to next three 
segments together. 4 mid leg with hooked scaly bristles, four on tibia, one on metatarsus. 
Wings with a large distal costal patch and small basal black spots, one below R,4;, one on 
anterior crossvein. PI =o0-7. p.f. = ABCDEfGHIKLMN. 

Taplejung District. Sangu, yellow blooms of cultivated composite (Guizotia 
abyssinica Cassini), c. 6,200’, 16-20.x.19g61, I g ; ibid., clearing in small wood, 
c. 6,200’, 25.xi.1961, Ig ; ibid., mixed vegetation by stream in gully, c. 6,200’, 
x1.1961-1.1962, 7 J, 4 2 ; ibid., mixed vegetation in deep gully, c. 6,200’, 2.1-13. 
11.1962, 2 2; between Sangu and Tamrang, in tree shade by hill stream, mossy 
ground under bushes, c. 5,500’, 20.x.19g61, I g, I 9, 23.x.1961, I 2 ; ibid., mixed 
plants by damp cliff in deep river gorge, c. 5,200’, i-ii.1962, 2 2 ; ibid., mixed shrubs 
in deep gorge, c. 5,200’, x—xi.Ig61, I fg ; below Sangu, edge of small mixed wood, 
c. 6,000’, 4.xi.Ig61, I 2 ; above Sangu, ex Lycopodium sp. on slope, c. 7,800’, 1I-14.1. 
1962, 1¢ ; ibid., mixed vegetation in dried-up ravine, c. 6,800’, 16.11.1962, I 3, 4 9 ; 
ibid., old mixed forest, c. 6,200’, 25-28.x.1961, 1 9 ; Dobhan, evergreen trees over- 
hanging stream in gully, c. 3,500’, 30.1.1962, 1 g ; ibid., east bank of River Tamur, 
mixed vegetation by stream in deep gully, c. 3,500’, i-ii.1962, I d (R.L.C.). 

Distribution : Nepal (new locality), China, Japan. 

The § has two small black markings near the wing base besides a large distal 
costal black marking. This type of wing pattern is the same as in the original form 
(China) (Tan, Hsu, and Sheng, 1949). In the Japanese form, however, the basal 
black markings are absent or very weak except in a form from the southern island, 
Yakushima, Kyushu, collected by the author in summer, 1962. 


Drosophila (Sophophora) tristipennis Duda 
(Text-figs. 255-261) 


Drosophila tristipennis Duda, 1924a : 215 ; 1924b : 247 (Formosa) ; Sturtevant, 1927 : 371 

(India). 

6, 2. Body about 1:7 mm. Head as broad as thorax. Eyes dark red, pilose. Antennae 
with second segment yellow and third grey in female, second white and third pale yellowish 
white in gj ; thirdoval. Arista with three dorsal and two ventral branches and a large terminal 
fork. Palpus yellow, apically broadened, with an apical prominent and a few ventral shorter 
setae. Ocellar triangle grey, posteriorly black. Periorbit grey, glossy in 9, milky white in 3. 
Front yellowish grey, frontal triangle with brown margin in 9, milky white and posteriorly 
somewhat orange in ¢, anteriorly one and one-third times as broad as length down middle, 
posteriorly slightly broader than half head width. Face matt greyish white. Carina long, 


go 


T. OKADA 


Fics. 255-267. 255-261, Drosophila (Sophophora) tristipennis Duda. 255, 3 head, lateral 
aspect ; 256, dg head, dorsal aspect ; 257, g wing; 258, periphallic organs, lateral 
aspect ; 259, phallic organs, ventral aspect ; 260, 9 egg-guide ; 261, hooked scaly bristles 
on g mid tibia (upper), sex-combs on proximal two tarsal joints of 3 fore leg (lower). 
262-267, Drosophila (Hirtodrosophila) tripartita sp. n. 262, g head, lateral aspect ; 263, 
3 head, dorsal aspect ; 264, g wing ; 265, periphallic organs, lateral aspect ; 266 phallic 
organs, ventral aspect ; 267, 2 egg-guide. 


DIPTERA FROM NEPAL 91 


narrow, low. Clypeus greyish yellow. Cheeks white in 3, yellowish grey in 9, about one-sixth 
as broad as the greatest diameter of eye. Occiput yellow, medially black. Anterior reclinate 
orbital five-eighths as long as others, twice as distant from posterior reclinate as from proclinate. 
Vibrissae long ; second oral half as long as vibrissa ; succeeding orals shorter. 

Mesonotum yellowish grey, somewhat glossy, nearly unicolorous. Scutellum pale yellowish 
brown, somewhat glossy. Thoracic pleura pale yellowish brown ; humeral callus pale. Two 
long humerals. Acrostichal hairs in six rows. Anterior dorsocentrals about half as long as 
posteriors ; distance between anterior and posterior dorsocentrals about two-fifths distance 
between anterior pair. Anterior scutellars convergent, slightly shorter than posteriors, which 
are nearer to each other than to anteriors. Sterno-index about 0-7. Legs yellowish white ; 
fore femur ventrally with several long hairs ; mid tibia apically with four hooked scaly bristles ; 
metatarsus without such bristles. Front and hind metatarsi as long as next three segments 
together, mid metatarsi as long as the other segments together. Preapicals on all legs ; apicals 
prominent on mid leg. Wings hyaline, broad, in g with a crescentic black patch along outer 
margin and a black stripe below F,, in 2 with a black spot at the apex of R,,,. Costa reaching 
M, R.+, straight, not curved to costa at tip ; Ry; and M parallel. C-index about 2:0; 4V- 
index about 2:7 ; 4C-index about 1:6 ; 5x-index about 4-0 ; Ac-index about 3:1. Cr1-bristles 
two ; C3-fringe on basal one-fourth. Halteres yellow. Abdominal tergites mostly black ; 1T 
yellow, 2T yellow on anterior half. Female cercus dark brown. 

Periphallic organs : Genital arch black, medially widely pale, mediocaudally gently pro- 
jected, dorsally exceedingly narrowing, ventrally truncate, pubescent at dorsal tip, setigerous 
along caudal margin. Clasper pale yellow, elongate, ventrally narrowing, subapically with a 
row of about five large teeth on distal margin. Secondary clasper present, above the primary 
clasper, with a straight row of about seven large black teeth on entire distal margin. Anal 
plate black, setigerous, rounded above, ventrally narrowing and with a few thick black bristles. 
Phallic organs : Aedeagus slender, bifid, orange, marginally with long serration ; basal apodeme 
slightly longer than aedeagus. P.I. = 0-7. Anterior paramere short, rod-like, black, attached 
to aedeagus basally, apically somewhat broadened and with a few sensilla. Posterior paramere 
pale, broad, fusiform in lateral aspect, proximally bilobed, attached to aedeagus by one arm 
and to the lateral corner of ventral fragma by the other. Ventral fragma semicircular, with a 
pair of short submedian spines ; hypandria pubescent. p.f. = AbCd’EfGHIKIMN. Egg- 
guide lobe pale yellow, oblong, subapically swollen dorsally, with about eight marginal black 
slender setae, ventrosubapical one longest. Basal isthmus yellow, one-sixth as long as lobe. 


Taplejung District, Sangu, mixed vegetation by stream in gully, c. 6,200’, 
ix-x.196I, I 9, xi.19g61-i.1962, I g ; edge of mixed forest above Sangu, c. 6,500’, 
I7.X-I.xi.1961, 11 g, 8 2 ; between Sangu and Tamrang, dense vegetation in tree 
shade by hill stream, c. 5,500’, 23.x.1961, I g, 2 9; ibid., mixed plants by damp 
cliff in deep river gorge, c. 5,200’, 22.xi.1961, I 9 ; Dobhan, evergreen trees over- 
hanging stream in deep gully, c. 3,500’, 30.1.1962, I g (R.L.C.). 

Distribution : Nepal (new locality), India, Formosa. 

Differs somewhat from the original form (Formosa) in having six rows of acrosti- 
chal hairs (eight in original form), and posteriorly black ocellar triangle (yellowish 
white in original form). The pectinate aedeagus shows a close relationship to the 
members of the melanogaster-subgroup. 


Drosophila (Sophophora) nepalensis Okada 
Drosophila nepalensis Okada, 1955 : 388 (Nepal). 


3, 2. Body dark brown, especially dark in g. Head as broad as thorax. Arista with about 
four dorsal and three ventral long branches and a moderate terminal fork. Front matt yellowish 
grey, with silver-white pollinosity, anteriorly narrowing and somewhat broader than length 


92 T. OKADA 


down middle. Ocellar triangle black, outer margin also black. Periorbit black, two-thirds as 
long as front, posteriorly narrowing, anterior end rounded. Anterior scutellars nearly as long 
as posteriors, which are equidistant from each other and from anteriors. All metatarsi subequal 
to next three segments together. Face deep black above, white below. Egg-guide lobe yellow, 
broadly truncate apically, with about twenty pointed black marginal teeth, ultimate three teeth 
widely apart from each other. Basal isthmus brown, long, nearly half as long as lobe. Sub- 
terminal hair long. Other features as described by Okada (1955). 

Taplejung District, Sangu, mixed vegetation by stream in gully, c. 6,200’, ix—x. 
1961, 1 2 ; ibid., rotten fruits of Bhor trees on ground, c. 6,200’, 7-31.x.1961, 10d, 
3 2. Arun Valley, below Tumlingtar, River Sabhaya, west shore, fermented millet 
seeds on shore, ¢. 1,800’, 24.xi1.1961, 4, 1 9 (R-L.C.). 

Distribution : Nepal, India. 


THE TAKAHASHII SuBcGRoupP 


Drosophila (Sophophora) takahashii Sturtevant 


Drosophila takahashi Sturtevant, 1927 : 371 (Formosa) ; Chino, 1927 : 473 (Japan) ; Peng 
1937 : 27 (China) ; Kikkawa and Peng, 1938 : 534 (Formosa, Manchuria) ; Okada, 1955: 
388 (Nepal). 

Drosophila (Sophophora) takahashu Sturtevant ; Sturtevant, 1942 : 29; Okada, 1956: I09 
(Japan) ; Okada, 1964a, : 111 (Japan: Amami) : Okada, 1964b : 449 (Borneo) ; Wheeler 
and Takada, 1964 : 174 (Mariana Is.). 

Taplejung District, Sangu, rotten fruits of Bhor tree on ground, c. 6,200’, 7—31.x. 

TOOL, 15.4, 0) 2 (e.C:). 

Distribution : Nepal, Formosa, China, Ryukyu, Manchuria, Japan, Mariana Is. 


THE MELANOGASTER SusBGrovup 
Drosophila (Sophophora) melanogaster Meigen 


Drosophila melanogaster Meigen, 1830 : 85 (Europe). 


Taplejung District, Sangu, from rotten Naspati fruits in traps, c. 6,200’, 8-31.x. 
1961, 13 6, 6 Y; ibid., mixed vegetation by stream in gully, c. 6,200’, xi.1961— 
i.1962, I 9; ibid., from rotten bananas in traps, c. 6,200’, x-xii.1g61, 20 9, 16 @ ; 
between Sangu and Tamrang, mixed plants by damp cliff in deep river gorge, 
€, 5,200, 225xt. TOO TL, prt? (ee): 

Distribution : Nepal, India, China, Manchuria, Korea, Formosa, Japan, Micro- 
nesia, Hawaii, Australia, Africa, Europe, N. America, S. America. 


THE MONTIUM Svupcroup 
Drosophila (Sophophora) kikkawai Burla 


Drosophila (Sophophora) kikkawai Burla, 1954a : 47 (Japan, China, Brazil, Hawaii?, Samoa?) ; 
Okada, 1955 : 387 (Nepal) ; Okada, 1956: 118 (Japan) : Okada, 1964b : 457 (Borneo) ; 
Kang, Chung, and Lee, 1959 : 62 (Korea) ; Wheeler and Takada, 1964 : 176 (Mariana Is., 
Palau, Ponape). 

Drosophila montium de Meijere, 1917 ; Duda, 1923 : 53 (Formosa, India) ; Duda, 1924 : 215 ; 
247 (Java, Formosa, India) ; Duda, 1926: 99 (Sumatra) ; Chino, 1927 : 473 (Japan) ; 


DIPTERA FROM NEPAL 93 


Kikkawa and Peng, 1938 : 530 (Japan, Korea, Manchuria, Saipan) ; Pavan and da Cunha, 

1947 : 20; 23 (Brazil) ; Tan, Hsu, and Sheng, 1949 : 197 (China). [Mis-identification]. 

Taplejung District, Sangu, mixed vegetation by stream in gully, c. 6,200’, ix—x. 
1961, 1g, 1 9; ibid., rotting fruits of Bhor tree on ground, c. 6,200’, 7-31.x.1961, 
23,32; ibid., from rotten Naspati fruits in traps, c. 6,200’, 8—31.x.1961, 18 g, I0 9, 
I puparium ; ibid., from rotten bananas in traps, c. 6,200’, x—xil.19g61, 33 J, 4 9; 
ibid., mixed vegetation by stream in gully, c. 6,200’, xi.1g61-1.1962, I 2 ; between 
Sangu and Tamrang, mixed shrubs in deep gorge, c. 5,200’, x-xi.1g61, 1 9 ; Arun 
Valley, below Tumlingtar, River Sabhaya, west shore, fermented millet seeds on 
shore, c. 1,800’, 24.xi1.1961, 14g, 11 9 ; east shore of River Arun below Tumlingtar, 
evergreen shrubs bordering dry stream-beds, c. 1,800’, 14—23.xil.1961, I ¢ (R.L.C.). 

Distribution : Nepal, India, Java, Borneo, Sumatra, Viet-Nam, Formosa, 
China, Japan, Korea, Micronesia, Hawaii, Samoa. 

The males and some females from Arun Valley are evidently smaller and darker 
than those from Taplejung, the former having sex-combs composed of lesser number 
of teeth (17 and 16 in the first and second tarsal joints, respectively) than in the 
latter (33 and 23), and the egg-guide teeth also few in number (12 to 16) than in the 
latter (20). These two forms agree however in the general features of phallic and 
periphallic organs. 


Drosophila (Sophophora) trapezifrons sp. n. 
(Text-figs. 268-273) 


3, 2. Body about 2 mm. Head slightly broader than thorax. Eyes oval, dark reddish 
brown, thickly pilose. Antennae with second segment yellowish orange ; third greyish brown, 
slightly longer than broad. Arista with about four dorsal and two ventral long branches and a 
small terminal fork. Palpus yellow, slender, with only one apical seta. Ocellar triangle black, 
somewhat glossy ; frontal triangle black. Periorbit black, narrow, straight, five-sevenths as 
long as front. Front dark orange-yellow, anteriorly paler and much convergent, anteriorly 
about two-thirds as broad as length down middle, posteriorly about half as long as head width. 
Face black, pollinose greyish white, paler along eye margin. Clypeus pale brown. Carina high, 
white, anteriorly broadened and fuscous, laterally fuscous. Cheek whitish grey, fuscous above, 
about one-seventh as broad as the greatest diameter of eye. Occiput brownish black, medially 
darker. Postverticals long. Anterior reclinate orbital about one-third as long as posterior 
reclinate, one and a half times as distant from posterior reclinate as from proclinate ; three 
orbitals nearly equidistant from eye margin, proclinate slightly inside others. Two long orals, 
succeeding orals finer. 

Mesonotum dark yellowish brown, glossy, posteriorly darker. Scutellum dark yellowish 
brown, slightly pollinose. Thoracic pleura yellowish grey, paler below. Two humerals, upper 
longer. Acrostichal hairs in six rows. Anterior dorsocentrals half as long as posteriors ; 
distance between anterior and posterior dorsocentrals two-fifths distance between anterior 
pair. Anterior scutellars divergent, slightly shorter than posteriors, which are slightly nearer 
to each other than to anteriors. Sterno-index about 0-6 ; median sternopleural one-third as 
long as posteriors. Legs yellow ; all metatarsi as long as next three segments together. Male 
fore leg with large longitudinal sex-combs, composed of about twenty-one black teeth on meta- 
tarsi and eighteen similar teeth on the second segment. Preapicals on all legs ; apicals on mid 
and hind legs. Wing hyaline, veins brown, crossveins clear. Costa reaching M ; R,,, and M 
nearly parallel ; R,+, straight, not curved to costa at tip. C-index about 2-8 ; 4V-index about 


94 


T. OKADA 


Fics. 268-279. 268-273, Drosophila (Sophophora) trapezifrons sp. n. 268, 3 head, lateral 


aspect ; 269, 3 head, dorsal aspect ; 270, dS wing ; 271, periphallic organs, lateral 
aspect ; 272, phallic organs, ventral aspect ; 273, 9 egg-guide. 274-279, Diathoneura 
vipa sp.n. 274, $ head, lateral aspect ; 275, 3 head, dorsal aspect ; 276, 3 wing ; 
277, periphallic organs, lateral aspect ; 278, phallic organs, ventral aspect ; 279, 2 
egg-guide. 


DIPTERA FROM NEPAL 95 


3:4; 4C-index about 1-4; 5x-index about 3-7 ; Ac-index about 2-2. Cr1-bristles two, sub- 
equal ; C3-fringe on basal two-sevenths. Halteres yellow. Tergites yellowish brown, with broad 
black cross-band towards tip of abdomen. 

Periphallic organs : Genital arch broad, black, pale grey at lower one third, caudal margin 
setigerous and gently convex below middle. Clasper in two sets : primary clasper ventrally 
narrowing, triangular, grey, with about ten small black teeth in a straight row on lower two- 
thirds of distal margin ; secondary clasper quadrate, above the primary one, partly attached to 
the lower tip of anal plate, with two large and one small black ventrally curved stout bristles on 
distal margin and a strong spine on inner margin, lower margin hairy. Anal plate separated 
from genital arch, oval, setigerous, with about three strong black bristles below. Phallic 
organs : Aedeagus compact, hairy, distally somewhat narrowing, subbasally with paired lateral 
black acute projections ; basal apodeme slightly shorter than aedeagus, straight, pale yellowish 
orange. PI = 1-4. Anterior paramere conical, black, rounded and with a few sensilla apically, 
with two acute projections medially, basally prolonged to join the base of aedeagus. Posterior 
paramere elongate, pale orange, gently curved outward, apically tapering. Ventral fragma 
quadrate, pale, with a large broad caudomedian process, which is apically shallowly concave and 
subapically with a pair of short submedian spines. p.f. = aBcDefGHIKIMN. Egg-guide lobe 
yellowish brown, oblong, gently narrowing apically and broadly truncate, with about fourteen 
black pointed marginal teeth, last tooth well removed from penultimate. Subterminal hair 
distinct. Basal isthmus black, one fourth as long as lobe. 

Holotype g. East NepaL : Taplejung District, Sangu, rotten fruits of Bhor tree 


on ground, c. 6,200’, 7—31.x.1961 (R.L.C.), BMNH. 
Allotype 9. Same data as holotype (R.L.C.), BMNH. 


Paratypes : Taplejung District, 1 3, 1 2, same data as holotype ; I g, between 
Sangu and Tamrang, mossy ground under bushes by hill stream, c. 5,500’, 20.x.1g61 ; 
4 3, ibid., mixed shrubs in deep gorge, c. 5,200’, x—xi.1961 ; I g, old mixed forest 
above Sangu, c. 6,200’, 25—28.x.1961 ; I 2, Sangu, from rotten Naspati fruits in traps, 
c. 6,200’, 8-31.x.1961 ; 2 9, ibid., from rotten bananas in traps, c. 6,200’, x-xil.1g6I ; 
I 9, ibid., mixed vegetation by stream in gully, c. 6,200’, ix—x.1961 ; I 9, below 
Sangu, mixed vegetation on sheltered slopes above river, c. 4,000’, 3.1.1962 ; I 3, 
between Sangu and Tamrang, deep river gorge, c. 5,200’, x—xi.I961I ; I 4, ibid. 
mixed plants by damp cliff in deep river gorge, c. 5,200’, 22.xi.1961 ; 1 9, Dobhan, 
shady places on shrubby slope above River Tamar, c. 3,500’, 21-27.1.1962 ; I 9, 
above Sangu, mixed vegetation in dried-up ravine, c. 6,800’, 16.11.1962 (R.L.C.), 
BMNH ; 24 (from Sangu and between Sangu and Tamrang) deposited at the Tokyo 
Metropolitan University. 


Resembles D. (S.) awraria Peng, 1937 (China and adjacent countries) in having 
trapezoid front and laterally clawed aedeagus, but differs in having ¢ face not 
whitish, periorbit black (grey in auraria), aedeagus hairy (bare in a.), anterior par- 
mere conical and hairy (elongate and not hairy in a.), and caudal margin of genital 
arch without prominent conical projection. 


THE ANANASSAE SuBGRovuP 
Drosophila (Sophophora) bipectinata Duda 
Drosophila bipectinata Duda, 1923 : 52 (India). 


96 T. OKADA 


No specimens were found in the present collection. The author (1955) recorded it 
from Central Nepal. 


Distribution : Nepal, India, Borneo, Formosa, Ryukyu, Japan (Amami Is.). 


Subgenus DROSOPHILA Fallen 
Drosophila Fallén, 1823 : 4. 


Type-species : Musca funebris Fabricius, 1787 : 345. 


KEY TO SPECIES 
I Body and legs dark brownish black. g anal plate contiguous with genital arch 


virilis section, robusta group . . lacertosa (p. 97) 
— Body and legs usually yellowish or yellowish brown, if dark brown, fore femur with a 
row of spinules. 4 anal plate separated from genital arch quinaria section 2 
2 Fore femur antero-ventrally with a longitudinal row of short stout spinules 
immigrans group 7 
— Fore femur antero-ventrally without a row of spinules 2 : : 9 : 3 


3 Second oral as long as or only slightly shorter than vibrissa. Palpus with a few 
prominent ventral setae 


: : : : 4 
— Second oral small or minute, at most half as s long as vitrciccel Palpus with only one 
long apical seta 2 grandis group 5 
4 Two long sternopleurals. Posterior parameres fused to form a large distal bow 
bizonata group . . trizonata (p. 97) 
— Three long sternopleurals. Posterior parameres apparently absent 
histrio group, part . . trisetosa (p. 99) 
5 Mesonotum with six dark longitudinal stripes . : : ; editinares (p. 103) 
— Mesonotum without distinct dark longitudinal stripes : é : : - 9 
6 Thoracic pleura with three distinct dark longitudinal stripes : acutissima (p. 101) 
— Thoracic pleura without distinct dark longitudinal stripes, with only one faint small 
dark patch above - : flexicauda (p. tot) 
7 Front and thoracic pleura ae distinct dake onetime Seas : : 
— Front and thoracic pleura without distinct dark longitudinal stripes . II 
8 Mesonotum and scutellum brownish black ; humeral callus and lateral sides of 
scutellum yellow. Wing slightly fuscous along costa : cubicivittata (p. 105) 
Mesonotum and scutellum pale yellowish brown, with dark longitudinal stripes. 9 
9 Mesonotum with six dark longitudinal stripes. Scutellum with two dark longitudi- 
nal stripes. Arista with only five branches including a ventral branch and a 
terminal fork é é notostriata (p. 107) 
— Mesonotum with five or seven dcr foeeteaeliacll Heres, Searels with or without 
distinct dark longitudinal stripes . : : , : : : 10 
10 Mesonotum with seven dark longitudinal stripes. Seaceligen with two dark longi- 
tudinal stripes. Legs with darkring . ; annulipes (p. 110) 
— Mesonotum with five dark longitudinal aries Sqelea without distinct dark 
longitudinal stripes. Legs without dark ring : . pentastriata (p. 109) 
11 Wing intensively clouded along costa. Posterior half of siescnee aa and scutellum 
largely brownish black. Femora largely black : : Suscicostata (p. 111) 
— Wing without dark clouds along costa. Mesonotum absoas yellowish brown or 
yellowish orange. Femora yellow : 12 
12 Tips of longitudinal veins R,,3, Ry; and M lacie 3 oe! ae vies eeates. 
¢$ fore metatarsi thick and with a tuft of long hairs : .  immigrans (p. 112) 


— Tips of longitudinal veins not black. ¢ front silvery shining. ¢ fore metatarsi not 
thick and without a tuft of long hairs . : d ; : . nasuta (p. 112) 


DIPTERA FROM NEPAL 97 


Tue VIRILIS SEcTION 
THE ROBUSTA Group 


Drosophila (Drosophila) lacertosa Okada 


Drosophila (Drosophila) lacertosa Okada, 1956 : 158 (Japan). 
Drosophila lacertosa Okada ; Takada and Lee, 1958 : 114 (Korea). 


3, 2. Body about 3:5 mm., brownish black. Head as broad as thorax. Ocellar triangle 
glossy greyish black, strongly convex. Periorbit glossy greyish black, anteriorly rounded, 
posteriorly narrowing. Front anteriorly broader than length down middle. Anterior reclinate 
orbital slightly nearer to proclinate than to posterior reclinate. Anterior dorsocentral three 
fourths as long as posteriors. Front metatarsi as long as next three segments together, mid and 
hind metatarsi much longer than the other segments. ,;, nearly straight. p.f. = aBCdefog, 
HIikLmn. PI = 3:5. Other features as described by Okada (1956) for the Japanese form. 


Taplejung District, Sangu, from rotten Naspati fruits in traps, c. 6,200’, 8-31.x. 
1961, 1g ; ibid., mixed vegetation by stream in gully, c. 6,200’, xi.1g61-1.1962, I 9 ; 
above Sangu, old mixed forest, c. 6,200’, 25-28.x.19g61, I 2 ; between Sangu and 
Tamrang, spray-splashed rocks in deep gorge, c. 5,200’, 6-28.x1.1961, 3g; Dobhan, 
spray-splashed rocks in River Maewa, c. 3,500’, 25.1.1962, I 2 (R.L.C.). 

Distribution : Nepal (new locality), Japan, Korea. 


THE QUINARIA SECTION 
Tue BIZONATA Group 


Anterior reclinate orbital minute. Second oral long. Palpus with a few promi- 
nent ventral setae. Posterior parameres present, fused to form a broad distal bow. 


Drosophila (Drosophila) trizonata sp. n. 
(Text-figs. 280-285) 


3, 9. Body about 2 mm. Head slightly broader than thorax. Eyes bright red, large, 
pilose. Antennae with second segment orange-yellow ; third greyish orange, apically 
rounded, about twice as long as broad. Arista with about five dorsal and three ventral branches 
and a small terminal fork. Palpus yellow, distally broadened, ventrally with a few long black 
setae. Ocellar triangle reddish brown ; inner margin of ocelli black. Periorbit orange-grey, 
somewhat glossy, posteriorly narrowing, anteriorly rounded and ending before the anterior 
margin of front. Front matt velvety orange, flat, laterally somewhat fuscous, anteriorly slightly 
narrowing and one and one-third times as broad as length down middle, posteriorly slightly 
more than twice as broad as head width. A few frontalhairs present. Clypeus yellowish brown. 
Face yellowish grey. Cheeks yellowish brown, about one-fourth as broad as the greatest dia- 
meter of eye. Occiput yellowish brown. Orbitals nearly equally distant from eye margin ; 
anterior reclinate orbital minute, not distinguished from neighbouring hairs, slightly nearer to 
proclinate than to posterior reclinate. Proclinate orbital slightly shorter than posterior reclin- 
ate. Postverticals long. Two long subequal orals, succeeding orals much shorter. 

Mesonotum glossy pale yellowish brown, medially and laterally with obscure fuscous patches. 
Scutellum yellowish brown, mediolaterally fuscous. Thoracic pleura yellowish brown, paler 
than mesonotum, with obscure fuscous patches on mesepisternum and sternepisternum. Two 
long humerals, lower one slightly longer. Acrostichal hairs in six rows. Anterior dorsocentrals 
half as long as posteriors ; distance between anterior and posterior dorsocentrals half distance 


98 


T. OKADA 


Fics. 280-292. 280-285, Drosophila (Drosophila) trizonata sp. n. 280, $ head, lateral 


aspect ; 281, g head, dorsal aspect; 282, wing; 283, periphallic organs, lateral 
aspect ; 284, phallic organs, ventral aspect ; 285, 2 egg-guide. 286-292, Drosophila 
(Drosophila) trisetosasp.n. 286, 3 head, lateralaspect ; 287, § head, dorsal aspect ; 
288, d wing ; 289, 9 egg-guide ; 290, periphallic organs, lateral aspect ; 291, phallic 
organs, ventral aspect, with aedeagus laterally placed ; 292, aedeagus, ventral aspect. 


DIPTERA FROM NEPAL 99 


between anterior pair. Anterior scutellars convergent, as long as posteriors, which are upright 
and nearer to each other than to anteriors. Sterno-index about 0-5 ; median sternopleural 
short. Legs yellow, all metatarsi as long as the next segments together. Preapicals on all 
legs ; apicals on mid leg. Wings hyaline, crossveins clear. Costa reaching M, weak beyond 
Ry+5- Ry+,and M parallel ; R.,, gently curved to costa at tip. C-index about 3-2 ; 4V-index 
about 1-8 ; 4C-index about 0-8 ; 5x-index about 1-4 ; Ac-index about 21. Cr1-bristles two ; 
C3-fringe on basal two-sevenths. MHalteres dark yellowish brown. Abdominal tergites glossy 
yellowish brown, each with a caudal black transverse band which is broadly interrupted at 
middle and narrowing laterally and is especially large on 2-4T, somewhat triangular on 2T. 

Periphallic organs : Genital arch pale yellowish brown, dorsally pubescent and not distinctly 
narrowing, ventrally narrowing abruptly and directed posteroventrally like a finger, apically 
with about four strong black setae. Clasper basally brown, short and broad, with a straight row 
of about thirteen long black teeth on entire outer margin, medially with about two strong black 
bristles. Anal plate large, pale yellowish brown, setigerous, dorsally pointed, ventrally broadly 
truncate, ventrocaudally with a short projection on which are inserted a few short stout setae. 
Phallic organs : Aedeagus dark orange, straight, rod-like, apically rounded. Anterior 
paramere brown, conical, partly fused to hypandrium, distally with a few sensilla. 
Posterior parameres fused to make a brownish, horizontally flattened, laterally deeply concave 
flap. Hypandrial plate oval, terminally with a long submedian spine. Ventral fragma tri- 
angular, pale brown, with deep acute median notch. p.f. = aBCdefgHIkIMN. PI = 2:0. 
Egg-guide lobe yellow, elliptical, broadly rounded apically, with about twenty marginal and 
three discal yellowish orange short thick teeth, ventrally with a short subterminal hair. Basal 
isthmus orange-brown, narrow, about one-fifth as long as lobe. 


Holotype g. East Nepat: Taplejung District, old mixed forest above Sangu, 
c. 6,200’, 25-29.x.1961 (R.L.C.), BMNH. 


Allotype 9. Same data as holotype (R.L.C.), BMNH. 


Paratypes : Taplejung District, 2 g, same data as holotype (R.L.C.), one at 
BMNH, the other at the Tokyo Metropolitan University. 


Closely resembles D. (D.) bizonata Kikkawa and Peng, 1938 (Japan and neigh- 
bouring countries), in having several long ventral setae on palpus, two long orals, 
glossy yellowish brown mesonotum and in general similar features of periphallic and 
phallic organs, but distinguished from it by having flat front, upright posterior 
scutellars, aedeagus less dilated distally, ventrally narrowing genital arch, somewhat 
triangular black bands on tergites, and clear crossveins. 


Tue HISTRIO Group 


Anterior reclinate orbital minute or small, about one-third to one-fifth as long as 
proclinate. Second oral nearly as long as vibrissa. Palpus with a few prominent 
ventral setae. Posterior parameres apparently absent. Including /istrio Meigen, 
1830, sternopleuralis Okada and Kurokawa, 1957, and trisetosa Okada in this paper. 


Drosophila (Drosophila) trisetosa sp. n. 
(Text-figs. 286-292) 


6,2. Body about 1-8mm. Head slightly broader than thorax. Eyes red, pilose. Anten- 
nae with second segment orange-brown, third greyish brown with rounded tip. Arista with 
about seven dorsal and four ventral branches and a small terminal fork. Palpus orange-brown, 
rounded at tip, with a long terminal and a few ventral black setae. Ocellar triangle black, 


100 T. OKADA 


marginally grey pollinose. Periorbit greyish brown, somewhat glossy, anteriorly blunt and not 
reaching anterior margin of front. Front matt orange-grey, anteriorly orange, and one and one- 
third times as broad as long, posteriorly half as broad as head width. Clypeus brown. Face 
greyish yellow, laterally concave. Carina high, long, laterally fuscous. Cheeks greyish yellow, 
about one-eighth as broad as the greatest diameter of eye. Occiput yellowish brown. Anterior 
reclinate orbital outside other orbitals, about one-third as long as posterior reclinate, half as long 
as proclinate, twice as distant from posterior reclinate as from proclinate. Postverticals 
moderate in size. Second oral nearly as long as vibrissa, succeeding orals short. 

Mesonotum and scutellum matt yellowish brown, somewhat pollinose ; humeral callus paler. 
Thoracic pleura greyish brown, sometimes with two broad obscurely demarcated dark brown 
longitudinal patches. Two humerals, lower longer. Acrostichal hairs in six rows. Anterior 
dorsocentrals less than half as long as posteriors ; distance between anterior and posterior 
dorsocentrals half distance between anterior pair. Anterior scutellars parallel, longer than 
posteriors, which are slightly nearer to each other than to anteriors. Sterno-index about 0:5. 
Median sternopleural strong, longer than anteriors, but shorter than posteriors. Legs fuscous 
yellow. Front metatarsi as long as next three segments together, mid and hind metatarsi as 
long as the other segments together. Preapicals on all legs ; apicals on mid leg. Wings 
hyaline, crossveins clouded. Costa reaching WM. R,+, gently convex at middle, apically nearly 
straight. R,,, and M parallel. C-index about 3:3; 4V-index about 1:8 ; 4C-index about 
0-8 ; 5x-index about 1-6 ; Ac-index about 1-7. Cr1-bristles two, subequal ; C3-fringe on basal 
two-thirds. Halteres yellow. Abdominal tergites matt yellowish grey, each with a medially 
interrupted clearly indicated straight black caudal band ; caudal tergites mostly black. 

Periphallic organs : Genital arch brownish black, caudal margin medially protruded tri- 
angularly and with a few long hairs, deeply and broadly concave at the base of clasper, narrow- 
ing below. Clasper brownish black, quadrate, large, with a slightly concave row of about eight 
black teeth on nearly entire length of distal margin. Anal plate oval, setigerous and pubescent, 
ventrally somewhat pointed and with a tuft of short hairs. Phallic organs : Aedeagus robust, 
yellowish orange, rod-like, apicodorsally with a short acute projection, which is directed caudally, 
apically somewhat dilated, basally with a large verticalrod. Apodeme of aedeagus thick, short, 
about one-third as long as aedeagus. PI = 3-0. Anterior paramere pale, narrow, apically 
with a long sensilla. Ventral fragma long, narrow, triangular. NHypandrial plate slender, 
laterally serrated, apically curved, basally attached on lateral arm of ventral fragma. A stout 
long submedian spine inserted below hypandrial plate. Median notch of ventral fragma very 
deepand wide. p.f. = ab’Cdefg,HIklmn. Egg-guide lobe yellowish brown, apically narrowing, 
with about sixteen marginal yellow teeth and three slender dorsal and discal setae. 


Holotype g. East NepaL: Arun Valley, east shore of River Arun below Tum- 


lingtar, evergreen shrubs bordering dry stream-beds, c. 1,800’, 14-23.xi1.1961 
(ROL.C.); MEN EL 


Allotype 9. Same data as holotype (R.L.C.), BMNH. 


Paratypes : 33, 20 2, same data as holotype ; 1 g, river banks below Tamrang 
Bridge, c. 5,500’, X—xi.I96I ; 1 9, between Sangu and Tamrang, mixed shrubs in 
deep gorge, c. 5,200’, x—xi.1961 ; 1 9, Arun Valley, above River Sabhaya, east shore, 
swept from dwarf bamboos in deep ravine, c. 2,000’, 12.xii.1961 ; 2g, 6 9, below 
Tumlingtar, River Sabhaya, west shore, evergreen shrubs on sandy shore, c. 1,800’, 
Q-17.xii.1961 ; 4, 4 9, ibid., west shore, fermented millet seeds on shore, c. 1,800’, 
24.xil.1961 ; 2 9, Dobhan, east bank of River Tamur, mixed vegetation by stream 
in deep gully, c. 3,500’, i-11.1962 (R.L.C.), BMNH, and 1 J, 1 9 at the Tokyo Metro- 
politan University. 


Closely allied to D. (D.) sternopleuralis Okada and Kurokawa, 1957 (Japan and 


DIPTERA FROM NEPAL 101 


Korea) in general features of § and @ genitalia, in having three long sternopleurals, 
and dark patches of thoracic pleura, but differs from it in the details of phallic and 
periphallic organs : lower tip of genital arch pointed (rounded in sternopleuralis), 
aedeagus more robust, and hypandrial plate serrated (non-serrated in s.). 


THE GRANDIS Group 


Yellowish brown or dark brown species, with black patches or stripes on thoracic 
pleura. Anterior reclinate orbital small, less than one-third as long as posterior 
reclinate. Second oral much shorter than vibrissa. Phallic organs without poster- 
ior parameres. Including D. (D.) grandis Kikkawa and Peng, 1938, tenuicauda 
Okada, 1956, acutissima Okada, flexicauda Okada, and editinares Okada, the 
latter three species included in this paper. 


Drosophila (Drosophila) acutissima Okada 
(Text-figs. 293-298) 
Drosophila (Drosophila) acutissima Okada, 1956 : 139 (Japan). 

Taplejung District, between Sangu and Tamrang, dense vegetation in tree shade 
by hill stream, c. 5,500’, 23.x.1961, I g ; ibid., mixed shrubs in deep gorge, c. 5,200’, 
X-X1.1961, 4.5, 3 2 ; ibid., mixed plants by damp cliff in deep river gorge, c. 5,200’, 
22.X1.1961, 4g, 3 2; below Sangu, edge of small mixed wood, c. 6,000’, 4.xi.1g961, 
I 2; Sangu, mixed vegetation by stream in gulley, c. 6,200’, xi.1961-1.1962, 7 3, 
8 2; above Sangu, mixed vegetation in dried-up ravine, c. 6,800’, 16.11.1962, 3 5 
preLC.). 

Distribution : Nepal (new locality), Japan. 

Slightly differing from original form (Japan) in having medioventrally swollen 
aedeagus and no distinct lateral lobe of aedeagus. Other features strictly agree 
with those of the Japanese form. 


Drosophila (Drosophila) flexicauda sp. n. 
(Text-figs. 299-305) 


36,2. Body 1-5-2 mm. Head somewhat narrower than thorax. Eyes oval, small, purplish 
red, pilose. Antennae with second segment orange-grey, third yellowish grey, apically some- 
what narrowing. Arista with about six dorsal and two ventral branches and a small terminal 
fork. Palpus yellowish grey, slender, rounded at tip, with a long apical seta. Ocellar triangle 
black, small. Ocelli pale yellow. Periorbit pale greyish yellow, broad, anteriorly extending 
near the anterior margin of front. Front broad, yellowish grey, anteriorly orange-yellow and 
broader than length down middle, nearly parallel-sided, posteriorly about three-fifths as broad 
as head width. A few frontal hairs present. Clypeus orange-grey. Face yellowish white. 
Carina broad, high. Cheeks yellowish white, dark at the insertion of orals, more than one-third 
as broad as the greatest diameter of eye. Occiput yellowish grey. Postverticals moderate 
in size. One strong oral, much longer than posterior reclinate orbital ; succeeding orals very 
slender. 

Mesonotum and scutellum uniformly yellowish brown, somewhat glossy. Thoracic pleura 
paler, with diffuse dark patch below humerus. Two humerals, upper longer. Acrostichal hairs 


102 T. OKADA 


Fics. 293-298. Drosophila (Drosophila) acutissima Okada. 293, 3 head, lateral aspect ; 
294, d head, dorsal aspect ; 295, wing; 296, 9 egg-guide ; 297, periphallic organs, 
lateral aspect ; 298, phallic organs, ventral aspect, with aedeagus laterally placed. 


DIPTERA FROM NEPAL 103 


in six rows. Anterior dorsocentrals two-thirds as long as posteriors ; distance between anterior 
and posterior dorsocentrals three-fourths distance between anterior pair. Anterior scutellars 
long, divergent ; posterior scutellars shorter than anteriors, as long as posterior dorsocentral. 
Sterno-index about 0:6. Legs yellow ; fore femur ventrally with about three very long setae ; 
all metatarsi nearly as long as the other segments together. Preapicals on all legs ; apicals on 
mid leg. Wings hyaline ; veins yellowish brown ; crossveins clear. Costa reaching slightly 
beyond R,,;. +, only weakly curved to costa apically ; Ry, and M nearly parallel. C-index 
about 2:7 ; 4V-index about 2:2 ; 4C-index about 1-1 ; 5x-index about 2:1 ; Ac-index about 
2-5. Cr-bristles two, subequal ; C3-fringe on basal one-third. MHalteres yellowish white. 
Abdominal tergites pale yellow, each with a broad black entire caudal band. 

Periphallic organs pale yellowish grey. Genital arch narrowly pointed ventrally, setigerous 
along caudal margin and on ventral tip, caudal margin acutely projected caudally above inser- 
tion of clasper, with a straight row of about eight long black teeth occupying ventral three-fifths 
of distal margin. Anal plate oval, large, with short hairs, ventrally broadened and truncate. 
Decasternum flat, triangular. Phallic organs pale brown. Aedeagus rod-shaped, straight, 
apically tapering and recurved dorso-anteriorly, two basally inserted elongate distally tapering 
appendages ; apodeme shorter than aedeagus, apically swollen. PI = 2-5. Anterior para- 
mere slender, as long as lateral appendages of aedeagus, proximally tapering and attached to 
the base of aedeagus, apically with a few sensilla. Hypandrial plate large, oval, basally with a 
stout submedian spine. Ventral fragma V-shaped, with deep median notch. Posterior para- 
mere apparently absent. p.f. = aBCdEfg,HIklmn. Egg-guide lobe yellowish orange, 
ventrally dark, elongate fusiform, with about fifteen marginal and four orange-brown discal 
teeth, distal marginal teeth large. Basal isthmus yellowish brown, medially much swollen, 
more than one-third as long as lobe. 


Holotype g. East NEPAL: Taplejung District, between Sangu and Tamrang, 
mixed plants by damp cliff in deep river gorge, c. 5,200’, 22.xi.1961 (R.L.C.), BMNH. 


Allotype 9. Taplejung District, above Sangu, mixed vegetation in dried-up 
ravine, c. 6,800’, 16.ii.1962 (R.L.C.), BMNH. 


Paratypes : Taplejung District, 2 J, 2 9, 1 ex., between Sangu and Tamrang, 
deep river forge, c. 5,200’, x—-xi.1961 ; I gf, Sangu, mixed vegetation in deep gully, 
c. 6,200’, 2.i-13.11.1962. 1, Arun Valley, east shore of River Arun below Tumling- 
tar, evergreen shrubs bordering dry stream-beds, c. 1,800’, 14-23.xii.1961 (R.L.C.), 
BMNH, 1 4, 1 9, from between Sangu and Tamrang deposited at the Tokyo Metro- 
politan University. 


Closely resembles D. (D) tenuicauda Okada, 1956 (Japan), especially in general 
structure of g and 2 genital apparata, and in having anterior reclinate orbital 
minute, only one strong oral, and a long apical seta of palpus, but distinguished from 
it by having non-striped mesopleura and apically recurved aedeagus. 


Drosophila (Drosophila) editinares sp. n. 
(Text-figs. 306-300) 


2. Body abouti-4mm. Head greyish yellow, as broad as thorax. Eyes dark purplish red, 
pilose. Antennae with second segment pale brown, third dark brown, oval, tiprounded. Arista 
with about four dorsal and two ventral branches and a small terminal fork. Palpus slender, 
dark brown, with a long apical seta. Ocellar triangle black, convex. Periorbit somewhat 
glossy grey, convex, relatively broad, medially narrowing, anteriorly almost reaching the 
anterior margin of front. Front broad, yellowish grey, matt, darker between ocellar triangle 


104 


T. OKADA 


Fics. 299-309. 299-305, Drosophila (Drosophila) flexicauda sp. n. 299, & head, lateral 


aspect ; 300, g head, dorsal aspect ; 301, 3 wing ; 302, periphallic organs, lateral 
aspect ; 303, phallic organs, ventral aspect ; 304, aedeagus, lateral aspect, dextral 
side dorsal; 305, 9 egg-guide. 306-300, Drosophila (Drosophila) editinares sp. n., 9. 
306, head, lateral aspect ; 307, head, dorsal aspect ; 308, wing ; 300, egg-guide. 


DIPTERA FROM NEPAL 105 


and periorbits, parallel-sided, anteriorly paler and one and a half times as broad as length down 
middle, posteriorly three-fifths as broad as head width. Clypeus greyish yellow, medially witha 
narrow dark transverse stripe. Face broad, pale yellowish grey, paler along eye margin, 
parallel-sided. Carina very large, broad, high, and long. Cheeks glossy yellowish grey, black 
at the insertion of vibrissa, nearly half as broad as the greatest diameter of eye. Anterior 
reclinate orbital half as long as proclinate, one-fourth as long as posterior reclinate, situated out- 
side proclinate, twice as distant from posterior reclinate as from proclinate. Vibrissae very long 
and black, succeeding orals slender. Postverticals relatively short, half as broad as inner verti- 
cals. 

Mesonotum yellowish grey, with three pairs of brownish longitudinal stripes, median pair 
broadest and reaching to the tip of scutellum, which is yellowish in ground colour. Thoracic 
pleura yellowish grey, with three longitudinal brownish stripes. Two humerals, lower weaker. 
Acrostichal hairs in six rows. Anterior dorsocentrals three-fourths as long as posteriors ; 
distance between anterior and posterior dorsocentrals equal to distance between anterior pair 
or slightly shorter. Anterior scutellars divergent, posteriors four-fifths as long as anteriors and 
somewhat nearer to each other than to anteriors. Legs yellow ; fore femur with several long 
setae below ; all metatarsi subequal in length to next three segments together. Preapicals on 
all legs ; apicals on mid leg. Wings broad, hyaline ; crossveins clear, tip of second costal 
section somewhat swollen and black. ,;, not prominently curved to costa at tip ; R4,, and M 
parallel ; costa weak beyond F,,;. C-index about 1-7 ; 4V-index about 2-3 ; 4C-index about 
1-5; 5x-index about 1-6 ; Ac-index about 3:1. Cr1-bristles two, subequal ; C3-fringe on basal 
half. Halteres yellowish white. Abdominal tergites yellowish grey, each with a broad entire 
caudal black band. Egg-guide lobe orange-yellow, apically narrowing and rounded, with about 
sixteen orange-brown thick marginal teeth and four slender discal teeth. Basal isthmus black, 
short, about one-eighth as long as lobe. 


Holotype 9. East NepaL: Taplejung District, between Sangu and Tamrang, 
mixed shrubs in deep gorge, c. 5,200’, x—xi.1961. (RL.C.), BMNH. 


Resembles D. grandis Kikkawa and Peng, 1938 (Japan), in having striped meso- 
notum and thoracic pleura, but differs from it in the swollen tip of second costal 
section and rounded tip of egg-guide. 


THe IMMIGRANS Group 


Drosophila (Drosophila) cubicivittata sp. n. 
(Text-figs. 310-315) 


3g. Body about 2:8 mm. Head as broad as thorax. Eyes dark brownish red, with grey 
pollinosity and pilose. Antennae with second segment greyish brown, third segment and arista 
missing. Palpus black, slender, with a long apical seta and a ventral one. Ocellar triangle 
glossy black, the black spot extending caudally to vertex. Periorbit yellowish white, obscurely 
limited, dark along either side between orbitals. Front matt yellowish white, with a pair of 
broad dark anteriorly convergent longitudinal stripes, parallel-sided, anteriorly broader than 
length down middle. posteriorly slightly broader than half head width. Clypeus pale brown, 
small. Face yellowish grey, somewhat glossy. Carina high, long, yellowish white, with a small 
brownish spot at lower tip. Postverticals moderate in size. Occiput yellowish grey, laterally 
black. Cheeks yellowish white, somewhat glossy, about one-fourth as broad as the greatest 
diameter of eye. Anterior reclinate orbital about half as long as proclinate, one-third as long as 
posterior reclinate, slightly nearer to proclinate than to posterior reclinate. Proclinate slightly 
inside other orbitals. Vibrissae long, succeeding orals less than half as long as vibrissa. 

Mesonotum dark chestnut-black, grey pollinose. Humeral callus yellow. Scutellum dark 
chestnut-black, pollinose, laterally yellow. Thoracic pleura pale yellowish grey, with three 


106 


T. OKADA 


Fiacs. 310-319. 310-315, Drosophila (Drosophila) cubicivittata sp.n., 5. 310, head, lateral 


aspect ; 311, head, dorsal aspect ; 312, wing ; 313, phallic organs, ventral aspect ; 
314, periphallic organs, lateralaspect ; 315, aedeagus, lateral aspect, dextral side dorsal. 
316-319, Drosophila (Drosophila) notostriata sp.n.,3. 316, phallic organs, ventral aspect ; 
317, aedeagus, lateral aspect, dextral side dorsal ; 318, fore femur; 319, periphallic 
organs, lateral aspect. 


DIPTERA FROM NEPAL 107 


dark brown longitudinal stripes, upper two anteriorly convergent and fused. Two humerals, 
upper longer. Acrostichal hairs in six rows. Anterior dorsocentrals half as long as posteriors ; 
distance between anterior and posterior dorsocentrals half distance between anterior pair. 
Anterior scutellars divergent, posteriors missing. Sterno-index about 0-6; median sterno- 
pleural as long as anteriors. Legs yellow ; all metatarsi subequal in length to the other seg- 
ments together. Fore femur anteroventrally with a row of about ten short stout black spinules 
on distal two-thirds. Preapicals on all legs ; apicals on mid leg. Wings hyaline, somewhat 
fuscous along costa. C reaching M ; R,,, medially convex forward, gently curved to costa 
apically ; Ry; and M nearly parallel. C-index about 3:9; 4V-index about 1:6; 4C-index 
about 0-6 ; 5x-index about 1-4 ; Ac-index about 2:1. Cr-bristles two ; C3-fringe on basal half. 
Halteres white. Abdominal tergites matt yellowish white, each with four large quadrate caudal 
black patches, 1T yellowish white. Abdominal sternites yellowish grey. 

Periphallic organs : Genital arch pale grey, pubescent and setigerous, anterior and posterior 
margins nearly straight and parallel, ventrally rounded and darker. Clasper pale grey, nearly 
quadrate, distal margin convex caudally on upper half and with a straight row of about eight 
black teeth on lower half. Anal plate brown, oval, setigerous and pubescent, narrowly projected 
caudoventrally, apically with about four stout black pointed bristles. Phallicorgans : Aedeagus 
yellowish orange, elliptical at lateral aspect, laterally flattened, ventro-anteriorly projected 
roundly ; basal apodeme rod-like, half as long as aedeagus. Anterior paramere small, conical, 
pubescent, attached to aedeagus basally, apically with a few sensilla. Ventral fragma quadrate, 
pale grey ; hypandrial plate large, quadrate, pale grey ; median notch deep and broad. PI = 
2:2. p.f. = aBCdEfg,HIklmn. 


Holotype g. East NepaL: Taplejung District, Sangu, mixed vegetation by 
stream in gully, c. 6,200’, xi.1961-i.1962 (R.L.C.), BMNH. 

Peculiar among the smmigrans-group in having black mesonotum, striped front, and 
anal plate ventrally narrowly projected with stout apical bristles. D. (D.) annu- 
lipes Duda, 1924 and D. (D). circumdata Duda, 1924, as well as the two succeeding 
species in this paper, have also striped fronts, but their mesonota are similarly 
striped, thus differing from the present species. 


Drosophila (Drosophila) notostriata sp. n. 
(Text-figs. 316-322) 


6. Body about 2.6 mm. Head slightly narrower than thorax. Eyes dark brownish red, 
pilose. Antennae with second segment dark brown, especially anteriorly ; third yellowish 
orange, anteriorly black. Arista with only two dorsal and one ventral branches and a large 
terminal fork. Palpus yellowish brown, distally broadened, apically darker, with a few stout 
ventral and an apical setae. Ocellar triangle black. Periorbit dull yellow, medially slightly 
narrowing, anteriorly somewhat curved inward and blunt, about two-thirds as long as front. 
Front yellowish orange, matt, with a pair of broad anteriorly convergent dark brown stripes, 
anteriorly somewhat narrowing, as broad as length down middle, posteriorly slightly narrower 
than half head width. Clypeus reddish brown, with a black median spot on buccal margin. 
Face yellowish grey. Carina high, grey, relatively short. Cheeks yellowish brown, about one-fifth 
as broad as the greatest diameter of eye. Occiput yellowish brown, laterally dark. Postverti- 
cals relatively short. Anterior reclinate orbital one-third as long as posterior reclinate and of 
proclinate, slightly nearer to proclinate than to posterior reclinate. Proclinate inside others. 
Second oral about one-third as long as vibrissa ; succeeding orals shorter. 

Mesonotum dull yellow, matt, with six narrow brownish black longitudinal stripes, outermost 
pair interrupted at sutures. Scutellum dull yellow, matt, with two broad brown longitudinal 
stripes. Thoracic pleura brownish black, grey pollinose, paler at sutures. Two long humerals, 


108 T. OKADA 


Fics. 320-327. 320-322, Drosophila (Drosophila) notostriata sp. n., $. 320, head, lateral 
aspect ; 321, head, dorsal aspect ; 322, wing. 323-327, Drosophila (Drosophila) 
fuscicostata sp.n.,3. 323, head, dorsal aspect ; 324, head, lateralaspect ; 325, wing ; 
326, periphallic organs, lateral aspect ; 327, phallic organs. 


DIPTERA FROM NEPAL 109 


subequal. Acrostichal hairs in six rows. Anterior dorsocentrals half as long as posteriors ; 
distance between anterior and posterior dorsocentrals half distance between anterior pair. 
Anterior scutellars divergent, as long as posteriors, which are somewhat nearer to anteriors than 
to each other. Sterno-index about 0-6; median sternopleural slender. Legs yellow ; fore 
femur anteroventrally with a row of about ten stout black spinules on distal two-thirds, larger 
and smaller teeth alternating. Preapicals and apicals on all legs. All metatarsi subequal in 
length to the next three segments together. Wings hyaline, vein Cu and posterior crossvein 
darker but not clouded, tipslightly pointed. R,;, nearlystraight ; R,,,and M slightly conver- 
gent apically. C-index about 3:1 ; 4V-index about 1-6 ; 4C-index about 0-8 ; 5x-index about 
1-5 ; Ac-index about 2:3. C1-bristles two ; C3-fringe on basal two-sevenths. Halteres yellow. 
Abdominal tergites matt yellow, each with a medially widely interrupted, and laterally somewhat 
broadened, posterior black transverse band, which is sometimes divided into four spots on 
posterior tergites. 

Periphallic organs greyish brown. Genital arch tapering above, swollen subventrally, some- 
what caudally curved apically, setigerous and pubescent, apically with several stout setae. 
Clasper triangular, somewhat caudally projected on upper half, with a straight row of about six 
stout black teeth along lower half of distal margin. Anal plate separated from genital arch as 
usual, fusiform, setigerous and pubescent, with about thirty-five short stout black pointed 
bristles in several rows ventro-apically. Phallic organs pale yellowish brown. Aedeagus 
straight, pubescent, distally swollen and club-shaped, laterally angulated, with a few short conical 
processes apically. Anterior paramere oval, half as long as aedeagus, with a few prominent 
sensilla apically, basally attached to aedeagus. Posterior parameres apparently fused to form 
a broad quadrate lobe. Ventral fragma quadrate ; hypandrial plate large and fusiform. 
Apodeme of aedeagus slender, straight, as long as aedeagus. PI = 1:2. p.f. = ab’CdEfgHik 
ImN. 


Holotype g. East NrepaL: Taplejung District, Sangu, mixed vegetation by 
stream in gully, c. 6,200’ xi.1g61-i.1962 (R.L.C.), BMNH. 


Allied to D. hexastriata Tan, Hsu, and Sheng, 1949 (China), but differs from it in 
having dark longitudinal stripes on front, branches of arista much fewer, second oral 
shorter, and acrostichal hairs in six rows (eight in hexastriata). The presence of a 
“sinous bow’”’ or “distal bow’”’ of fused posterior parameres in this species suggests 
a connection with the tvipunctata-group. Although some species of this group have 
the row of spinules on fore femur, like the members of the zmmigrans-group, this 
resemblance was considered by Frota-Pessoa (1954 : 265) to be superficial, since the 
latter group lacks the distal bow. 


Drosophila (Drosophila) pentastriata sp. n. 
(Text-figs. 192-195, p. 68, 219-220, p. 75) 


6. Body about 2:4 mm., relatively slender. Head slightly broader than thorax. Eyes deep 
red, thickly pilose. Antennae with second segment dark yellow ; third broader, oval, yellow 
with anterior half black. Arista with about three dorsal and one ventral very long branches and 
a very large terminal fork. Palpus and mouth-parts unfortunately missing. Ocellar triangle 
black. Periorbit yellowish grey, somewhat glossy, somewhat narrowing anteriorly, four-fifths 
as long as front. Front orange-yellow, with two broad dark brown anteriorly convergent 
longitudinal stripes, proximal half of these diffusely dilated inferiorly. Front anteriorly narrow- 
ing and as broad as length down middle, posteriorly slightly narrower than half head width. 
Clypeus yellow, medially with a narrow brownish transverse band. Face wide, greyish white, 
parallel-sided, white along eye margin. Carina high, long, narrow, yellowish white. No frontal 
hairs. Cheeks yellowish white, about two-sevenths as broad as the greatest diameter of eye. 


IIo T. OKADA 


Occiput yellowish orange. Postverticals moderate in size. Anterior reclinate orbital about 
half as long as others, nearly equally spaced from posterior reclinate and from proclinate, which 
is inside others. Only one long oral, others much shorter. 

Mesonotum orange-brown, with five narrow dark brown longitudinal stripes, median one faint 
and ending before posterior extremity, lateral one interrupted at suture, becoming more dorsal 
behind suture. Scutellum yellowish brown, pollinose. Thoracic pleura paler than mesonotum, 
with two narrow obscurely dark longitudinal stripes, upper one longer and reaching propleura 
anteriorly. Two humerals, upper much shorter. Acrostichal hairs in six rows. Anterior 
dorsocentrals half as long as posteriors ; distance between anterior and posterior dorsocentrals 
subequal to distance between anterior pair. Anterior scutellars divergent, as long as posteriors, 
which are slightly nearer to each other than to anteriors. Sterno-index about 0:5 ; anterior 
sternopleural much thinner ; median one as long as anteriors. Legs yellow ; front and mid 
metatarsi subequal in length to next three segments together, mid metatarsi slightly longer than 
next three segments together. Preapicals on all legs ; apicals on fore and mid legs. Fore 
femur anteroventrally with a row of small black equal-sized spinules on median half. Wings 
hyaline, costa reaching M ; R,4, straight ; R,,; and WM gently convergent distally. C-index 
about 3:0 ; 4V-index about 1-5 ; 4C-index about 0-8 ; 5x-index about 1-3 ; Ac-index about 2:7. 
C1-bristles two ; 3C-fringe on basal one-fourth. Halteres yellow. Abdominal tergites dark 
yellowish orange ; 1T pale, others each with a broad black, medially narrowly interrupted, 
posterior transverse band. 

Periphallic organs : Genital arch pale yellowish grey, relatively narrow, swollen and oval 
ventrally, pubescent and setigerous. Clasper pale brown, triangular, proximally narrowing, 
narrowly projected upward dorsally to reach the lower tip of anal plate, with a straight row of 
about eight stout blunt black teeth on entire distal margin. Anal plate large, fusiform, setige- 
rous and pubescent, ventrally with a tuft of a few short stout black setae directed ventrally. 
Decasternum pale, broad, triangular. Phallic organs pale yellow. Aedeagus robust, apically 
blunt, rhombic and basally swollen upward in lateral aspect. Basal apodeme of aedeagus 
straight, half as long as aedeagus. Anterior parameres oval, small, close to each other, finely 
pubescent, basally attached to aedeagus, subapically with about two sensilla. Hypandrial plate 
quadrate, finely pubescent, medially with a long submedian spine. Ventral fragma elongate, 
quadrate ; lateral corners sharply pointed apically. PI = 2-0. p.f. = aBCdefg,HIklmn. 


Holotype §. East NepaL: Taplejung District, Sangu, mixed vegetation by 
stream in gully, c. 6,200’, xi.1961-1.1962 (R.L.C.), BMNH. 


Allied to notostriata in having striped front and mesonotum as well as thoracic 
pleura, but differing from it in having an odd number of mesonotal stripes, larger 
fork of arista, rounded wing tip, equal-sized short spinules on fore femur, non- 
pubescent aedeagus, and in the details of ¢ genital armatures. 


Drosophila (Drosophila) annulipes Duda 


Drosophila annulipes Duda, 1924a : 209; 221 (Formosa) ; Duda, 1923: 58 [nom. nud.], 

(Formosa) ; Wheeler and Takada, 1964 : 183 (Bonin Is.). 

Drosophila (Drosophila) virgata Tan, Hsu, and Sheng, 1949 : 203 (China) ; Okada, 1956 : 149 

(Japan); Kim, 1963 : 73 (Korea). 

Taplejung District, Sangu, mixed vegetation by stream in gully, c. 6,200’, 1x—x. 
1961, I g ; ibid., xi.1961-1.1962, 1 g ; below Sangu, by stream in shady ravine, 
c. 6,000’, 30.x.1g61, I 2 ; above Sangu, old mixed forest, c. 6,200’, 25-28.x.1961, 
IQ; ibid., mixed vegetation in dried-up ravine, c. 6,800’, 16.11.1962, I 2 (R.L.C.). 

Distribution : Nepal (new locality), Formosa, China, Korea, Japan, Bonin Is. 


DIPTERA FROM NEPAL | I1I 
Drosophila (Drosophila) fuscicostata sp. n. 
(Text-figs. 323-327) 


3. Body about 3 mm. Head as broad as thorax. Antennae with second segment brown ; 
third greyish brown, slightly longer than broad. Arista with about four dorsal and one ventral 
branches and a large terminal fork. Palpus yellowish brown, ventrally with about two black 
setae. Ocellar triangle black, marginally grey pollinose. Periorbit glossy greyish brown, 
posteriorly broadened, two-thirds as long as front. Front orange, anteriorly somewhat narrow- 
ing and slightly broader than length down middle, posteriorly fuscous and broader than half 
head width, caudolaterally black and grey pollinose outside insertion of postverticals. Face 
glossy brown ; buccal margin strongly convex. Carina high, short, medially whitish grey, 
broader below. Cheeks yellowish grey, dark at bases of orals, broad, about two-ninths as broad 
as the greatest diameter of eye. Occiput yellowish grey, black below. Anterior reclinate 
orbital slender, about one-third as long as posterior reclinate, half proclinate, slightly nearer to 
proclinate than to posterior reclinate ; proclinate inside others. Postvertical moderate in size. 
Second oral fine, about two-fifths as long as vibrissa ; succeeding orals finer. 

Mesonotum orange-brown, matt, posteriorly and laterally dark glossy brown. Scutellum 
brown, anterolaterally and apically yellowish orange, with two diffuse black stripes. Thoracic 
pleura black, grey pollinose, pale along suture. Only one long humeral. Acrostichal hairs in 
six rows. Distance between anterior and posterior dorsocentrals about half distance between 
anterior pairs. Scutellars missing. Posterior scutellars nearer to each other than to anteriors. 
Sterno-index about 0-6 ; median sternopleural half as long as anteriors. Legs yellowish grey ; fore 
femur anteroventrally with a row of about twelve black spinules on distal two thirds ; femora 
black except at ends ; tibiae fuscous, with indistinct dark rings subbasally and subapically. 
Front and mid metatarsi subequal in length to next three segments together, hind metatarsi 
longer than the rest of segments together. Wings extensively fuscous from costal margin up 
to below F,+3, in the cell R,;, before anterior crossvein, along /, and along crossveins. 5+, 
straight, not curved to costa apically ; R,;, and 17 somewhat convergent distally. C-index 
about 3-8 ; 4V-index about 1-7 ; 4C-index about 0-7 ; 5x-index about 1-1 ; Ac-index about 
2:3. C1-bristles two ; C3-fringe on basal two-thirds. MHalteres yellow. Abdominal tergites 
matt yellowish grey, each with a glossy black caudal transverse band, which is laterally curved 
anteriorly, medially interrupted on 2-3T, medially contiguous and projected anteriorly on 
4-51 ; 11 mostly yellow ; 6T black. 

Periphallic organs brownish black. Genital arch not narrowing above, ventrally rounded, 
setigerous and pubescent. Clasper projected below genital arch, ventrally prolonged, with a 
slightly concave row of about ten stout black teeth occupying nearly entire distal margin. Anal 
plate setigerous and pubescent, dorsally tapering, ventrally broadened and with numerous short 
stout black bristles. Phallic organs : Aedeagus orange-brown, straight, subapically broadened 
and dentated on lateral sides, proximally tapering, subbasally with an elongate distally black 
and caudally pointed dorsal appendage. Anterior paramere small, conical, distally with about 
two sensilla. Hypandrial plate pale yellow, marginally black, broad, elliptical, with a stout 
submedian spine. p.f. = ab’Cdefg,HIkIMn. 


Holotype g. East NepaL: Taplejung District, Sangu, mixed vegetation by 
stream in gully, c. 6,200’, xi.1g61-1.1962 (R.L.C.), BMNH. 


Somewhat resembles D. hypocausta Osten Sacken, 1882 (Philippines, Java, 
Sumatra, and Caroline Islands) in having black femora, black thoracic pleura, and 
apically pale scutellum, but differs from it in having six rows of acrostichal hairs 
(eight in hypocausta), only one strong oral (two strong in h.), anteriorly fuscous 
wings, only one ventral branch of arista (four in h.), ventrally setigerous anal plate 
and long dorsal projection of aedeagus. 


II2 T. OKADA 


Drosophila (Drosophila) immigrans Sturtevant 


Drosophila immigrans Sturtevant, 1921 : 83 (N. America). 


Taplejung District, Sangu, rotting fruits of Bhor tree on ground, c. 6,200’, 7—31.x. 
1961, 19 g, 15 9; ibid., from rotten bananas in traps, c. 6,200’, x—xii.1961, 2 Q ; 
ibid., mixed vegetation by stream in gully, c. 6,200’, xi.1g61-1.1962, I 9 ; between 
Sangu and Tamrang, mixed shrubs in deep gorge, c. 5,200’, x—xi.1961, 1 9. Arun 
Valley, below Tumlingtar, River Sabhaya, west shore, from rotten bananas in traps, 
c. 1,800’, x—xil.1961, 13 J, 27 9; ibid., evergreen shrubs on sandy shore, 9-17.xii. 
1961, I 9 ; ibid., fermented millet seeds on shore, c. 1,800’, 24.xii.1961, 2 2 ; ibid., 
east shore of River Arun, evergreen shrubs bordering dry stream-beds, c. 1,800’, 
14—-23.x1.1961, I 9 (R.L.C.). 

Distribution : Nepal, India, Formosa, China, Korea, Japan, Philippines, Micro- 
nesia, Hawaii, Australia, Africa, Europe, N. America, S. America. 


Drosophila (Drosophila) nasuta Lamb 


Drosophila nasuta Lamb, 1914 : 346 (Seychelles). 

Drosophila (Drosophila) nasuta Lamb : Okada, 1964b : 463 (Borneo) [synonymy] ; Wheeler 
and Takada, 1964 : 180 (Mariana Is., Truk, Ponape, Kusaie, Wake Is.) [synonymy]. 
Taplejung District, Sangu, rotting fruits of Bhor tree on ground, c. 6,200’, 7-31.x. 

1961, 16g, 112 ; between Sangu and Tamrang, mixed shrubs in deep gorge, c. 5,200’, 

X-x1.1961, 1 9. Arun Valley, below Tumlingtar, River Sabhaya, west shore, from 

rotten bananas in traps, c. 1,800’, x—xii.1961, I 9 (R.L.C.). 

Distribution : Nepal (new locality), Borneo, Sumatra, Seychelles, Formosa, 

Japan, Moluccas, New Guinea, Micronesia, Samoa, Fiji, Hawaii. 


GEOGRAPHICAL DISTRIBUTION OF THE NEPALESE SPECIES 


Table 1 shows that the drosophilid fauna of Nepal seems to be most closely related 
to that of Formosa, nearly one-third of the known Nepalese species being in common 
(twenty-five species). Much fewer Nepalese species have been found in common with 
the adjacent areas, India, Viet-Nam or China, and this is certainly due mainly to the 
poor knowledge of the fauna in these areas as compared with the relatively rich 
records from Formosa. Similar relationships between the number of species in com- 
mon and the degree of survey can be recognized among the islands of Java, Sumatra, 
and Borneo. Of these, Java has been best surveyed and is richest in the number of 
species in common (fourteen species), while Borneo is poorest in the degree of survey 
as well as in the number of species in common (three). 

Japan comes next to Formosa in the number of species in common (nineteen 
species), which is about one-fourth of the total Nepalese species. This is undoubt- 
edly due partly to the extensive surveys made in Japan and partly to the biogeo- 
graphical similarity between the two countries. The latter has frequently been 
proved concerning various groups of insects and plants, e.g., Epiophlebia (Aniso- 
zygoptera, Odonata) and Deuterophlebia (Deuterophlebiidae, Diptera) in insects and 
Balanophora, Burmannia, and Hydrobryum in Angiosperms, referring only to special 


DIPTERA FROM NEPAL 113 


cases. In regard to the drosophilid fauna, Cryptochaetum nipponense, Drosophila 
acutissima and D. lacertosa, which are common to both countries and are so-called 
“wild species”, and D. trizonaria and D. trisetosa, which are closely related to the 
Japanese species, D. bizonata and D. sternopleuralis respectively, should be the 
examples proving connection between fauna of the two countries. Table 2 shows 
that one hundred and twenty-seven species, so far known from Japan, belong to 
sixteen genera, of which ten also occur in Nepal. The remaining six genera, Curto- 
notum, Amiota, Mycodrosophila, Dettopsomyia, Lissocephala, and Chymomyza have 
not been found in Nepal, while five genera, Campichoeta, Apsinota, Lissodrosophila, 
Diathoneura and Paramycodrosophila, recorded from Nepal in this paper, have not 
been found in Japan. Six subgenera of the genus Drosophila (Scaptodrosophila, 
Dichaetophora, Hirtodrosophila, Dorsilopha, Sophophora, and Drosophila s. str.) 
occur in both countries. The subgenus Lordiphosa is absent from Nepal, while 
occurring in Japan. The occurrence of an originally African subgenus, Dichaeto- 
phora, in both Nepal and Japan is rather easily interpreted biogeographically, but the 
discovery of an originally South American genus, Diathoneura, is harder to explain. 
This might be a phenomenon of “‘trans-palaeo-equatorial distribution” (Maekawa, 
1963). 

The Nepalese species which also occur in Europe (five species), Australia (eight), 
Africa (nine), N. America (four), and S. America (five) are mostly cosmopolitan or 
circumtropical, except Campichoeta obscuripennis and Leucophenga leucozona, 
which, outside Nepal, are found only in Europe and Australia, respectively. The 
genus Leucophenga and the melanogaster-group are the taxa flourishing especially 
in the Oriental Region and seem to have spread simultaneously to Nepal and Japan, 
which have several species in common. 


OKADA 


an 


114 


TABLE I 


v's 


PoIOUW 


BoJeWYy *N 


adomg 


Bolly 


BTeajsny 


eisauhjog 


EISAUOIOI 


aurdditud 


oaul0g 


Pate 


eajyeuns 


WeUjaTA 


esoul1o.7 


ueder 


vaI0y 


Buty 


erpul 


Tedon 


‘ds "Ww 
ByeUI}0ed ETLYdoso pos 
sdasiqie *y 
EJETTANOSoAsIu “ry 
STAJOUT}OAI "7 
sdao1a31u 
S1.1]U9A1})NS 
sidyedia31u 


aeaijeles 


aT 

al 

int 

at 
eye}Oundosa;TeYy *T 
eByEjuesie *y 
PIOsenq(e *T 
Buozooney] *T 
Bydniz1aju *7 
ByeUuIO “TT 
Bsouttjodqns *7 

Bye}S09 "7 

P}BIAVIQge eduaydoonayT 
SUOIPIISIU vuese}S 
SIIJUSATIOId *y 

sadin4a ejoursdy 
Stuuadtanosqo ejaoyotduieg 
luepseq *q 

BOLINssn eye}SeIG 


esuauoddiu wnjaeyooidh19 


115 


DIPTERA FROM NEPAL 


TABLE 1 (Continued) 


roawy *S 


pollawy *N 


adoing 


eoldyy 


eiyeajsny 


eisaudjog 


PISIUOIOIW 


audditiud 


oaus0g 


eaee 


eajewns 


weujat, 


rsow soy 


ueder 


ra10y 


eulyD 


PIpuy 


jedan 


1300 


Tryosnq 


PIEISOIIQUITT 


vjsed1 41} 


PyeWsplies 


stusooipunjo.1 


v}eUusaye 


a jw VA GE 4 a e e 


pyeulsreW 


PUNUTUL 


snazoiid 

vauuniq epiydosoig 

edia einauoyieiq 

PRATIS *¢ 

PUIISSIULTAU *¢ 
suapualdsered “sg 

epitred ezAuojdeog 

1909 PT[aT1udosoazpoyaryD 
PIN}o1d eptydosoaposAweseg 
eprytu "Ty 

stuuadtosny "Ty 

edAdoyouo "7 

ByE[NoewI1q e[Iydosozpor'y 
e}e}1NF vaIAyJOTasdAH 
s1uroo1du0] e[tydoso.zpossr'] 
BIVUISZIEU "A 
BeIITBYOISIu “PL 
sidjediisiu “Ww 


esowresoned “WW 


T. OKADA 


I16 


TABLE I (Continued) 


eollawy °S 


eoIauly *N 


adoing 


BOY 


eITeIsny 


visaudjog 


or 


eISQUOIOIN, 


autddiqryd 


oaul0g 


bl 


rare 


It 


eayeuns 


WEI A 


SZ 


BSOUIOJ 


61 


ueder 


ealoy 


euryd 


erpul 


LL 


Tedon 


Te}OL 


eynseu *q 


supas1wwt 


BIB SOOTOSNy 


sadijnuue 


eeLAWsSeUed 


PyeLISOJOU 


ByeIWATOIQND 


SaieUulyIpe 


epneoixeayy 


BWISST}NOe 


BSOJISI 1} 


PyeuoZII} 


BSO}IIIOPT 


eyeuljoad1q 


suoajizode.1y 


TeME PTY 


Jajsesourjaw 


TIyseyexEy 


sisueledou 


suuedijs117 


. 


RTeazyotnd 


Slivpnoeurunt 


. 


A AA Alla a ia wal va (Al a a wa ta a ee re eo roa 


V[PARTPl41as 


BIIBUOZ 


DIPTERA FROM NEPAL 117 


TABLE 2 
Comparison of Nepalese and Japanese drosophilid faunae 


Genus and Nepalese Japanese Species 

Subgenus species species in common 
Cryptochaetum I I 
Diastata 
Campichoeta 
A psinota 
Stegana 
Leucophenga 
Microdrosophila 
Lissodrosophila 
Hypselothyrea 
Liodrosophila 
Paramycodrosophila 
Chaetodrosophilella 
Diathoneura 
Scaptomyza 
Drosophila 
Other six genera 


Lal 
_ 


ne eR eR HH OOH NY HN 


| Ww 
° 
nD 
AN WOH ONHOANWOO RN 


Bla ol 
oO OoNH OOO OOO Of OO OF 


Total 


“ 
“ 


127 


Scaptodrosophila 
Dichaetophora 
Hirtodrosophila 
Dorsilopha 
Sophophora 
Drosophila s. str. 
Lordiphosa 


HoH 
OWNRNARNNWN 


~ 
“I 
ONNnNH OHO 


ECOLOGICAL DISTRIBUTION OF THE NEPALESE SPECIES 


Following is a summarized list of the collection data recorded by Mr. R. L. Coe for 
the present material. The numbers following the generic and subgeneric names are 
those given to the species in Table 1. 


Taplejung Districts 
Sangu, c. 6,200’. 

Mixed vegetation by streamin gully. Leucophenga (8, 17, 20, 23), Microdrosophila 
(26, 28, 29), Liodrosophila (32, 33, 34, 35), Chaetodrosophillela (37), Scaptomyza 
(38, 39), Drosophila (43, 44, 48, 49, 50, 53, 54, 55, 57, 58, 59, 61, 62, 63, 65, 68, 71, 72, 
73 74, 75, 76, 77)- 

Rotting fruits of Bhor tree on ground. Hypselothyrea (31), Liodrosophila (33), 
Drosophila (59, 60, 62, 63, 76, 77). 

Mixed vegetation in deep gully. Liodrosophila (34), Chaetodrosophillela (37), 
Scaptomyza (39), Drosophila (53, 69). 

Clearing in small wood. Scaptomyza (39, 40), Drosophila (53, 57). 


118 T. OKADA 


Yellow blooms of cultivated Composite (Guizotia abyssinica Cassini). Scaptomyza 
(38), Drosophila (47, 57). 

From rotten Naspati fruits in traps. Dvrosophila (52, 61, 62, 63, 65). 

From rotten bananas in traps. Drosophila (52, 61, 62, 63, 76). 

By stream in gully. Dvosophila (53). 
Above Sangu. 

Old mixed forest, c. 6,200’. Leucophenga (21, 23), Microdrosophila (27), Liodroso- 
pula (35), Scaptomyza (39), Drosophila (53, 57, 63, 65, 66, 74). 

Mixed vegetation in dried-up ravine, c. 6,800’. Microdrosophila (28), Chaeto- 
drosophilella (37), Scaptomyza (38, 39), Drosophila (50, 53, 57, 63, 68, 69, 74). 

Damp evergreen oak forest, c. 9,200’. Cryptochaetum (1), Diastata (3), Scapto- 
myza (39, 40, 41), Drosophila (53). 

Ex Lycopodium sp. on slope, c. 7,800’. Scaptomyza (38, 40), Drosophila (57). 

Evergreen scrub, c. 6,500’. Leucophenga (22), Chaetodrosophilella (37). 

Edge of mixed forest, c. 6,500’. Liodrosophila (34), Drosophila (58). 


Below Sangu. 

By stream in shady ravine, c. 6,000’. Cryptochaetum (1), Leucophenga (8), 
Lissodrosophila (30), Hypselothyrea (31), Liodrosophila (32, 33, 34, 35), Drosophila 
(52, 53, 74). 

Mixed vegetation on sheltered slopes above river, c. 4,000’. Diastata (2), Leuco- 
phenga (10, 21, 23), Liodrosophila (34), Scaptomyza (38, 39), Diathoneura (42), 
Drosophila (53, 63). 

Edge of small mixed wood, c. 6,000’. Leucophenga (21, 23), Liodrosophila (35), 
Scaptomyza (39), Drosophila (57, 68). 

Below stream in shady ravine, c. 6,200’. Scaptomyza (39). 


North of Sangu. 
Dry grass above river bank, c. 5,000’. Leucophenga (20), Drosophila (53). 
Mossy ground under bushes by hill stream, c. 5,500’. Leucophenga (20), Droso- 


phila (53, 63). 


Between Sangu and Tamrang. 

Mixed shrubs in deep gorge, c. 5,200’. Campichoeta (4), Leucophenga (9, 13, 17, 
20, 21, 22), Liodrosophila (33, 34), Chaetodrosophilella (37), Drosophila (46, 50, 53, 54, 
55, 57, 62, 63, 67, 68, 70, 76, 77). 

Mixed plants by damp cliff in deep river gorge, c. 5,200’. Stegana (7), Leucophenga 
(17, 20), Microdrosophila (28), Liodrosophila (34), Chaetodrosophilella (37), Drosophila 
(46, 48, 51, 53, 54, 55, 57, 58, 61, 63, 68, 69). 

Dense vegetation in tree shade by hill stream, c. 5,500’. Leucophenga (21), Droso- 
phila (53, 58, 68). 

Shrubs by path, 5,800’. Leucophenga (17), Drosophila (56). 

In tree shade by hill stream, c. 5,500’. Drosophila (57). 

Spray-splashed rocks in deep gorge, c. 5,200’. Drosophila (65). 

Deep river gorge, c. 5,200’. Drosophila (63, 60). 


DIPTERA FROM NEPAL 119 


Below Tamrang Bridge. 
River banks, c. 5,500’. Leucophenga (10), Liodrosophila (34), Scaptomyza (39), 
Diathoneura (42), Drosophila (48, 52, 53, 54, 67). 


Dobhan, c. 3,500’. 

East bank of River Tamur, mixed vegetation by stream in deep gully. Mzucro- 
drosophila (24), Liodrosophila (35), Scaptomyza (39), Drosophila (57, 67). 

Evergreen trees overhanging stream in deep gully. Leucophenga (15, 20), Droso- 
phula (57, 58). 

Shady places on shrubby slope above River Tamur. Leucophenga (10), Scaptomyza 
(38), Drosophila (63). 

Small pockets of plants on arid slopes above River Maewa. Leucophenga (10), 
Microdrosophila (20). 

Spray-splashed rocks in River Maewa. Drosophila (65). 

Mixed vegetation in dry gully on wooded slope. Leucophenga (19). 


Arun Valley 
Below Tumlingtar. 

East shore of River Arun, evergreen shrubs bordering dry stream-beds, c. 1,800’. 
Apsinota (6), Leucophenga (8, 9, 10, II, 13, 15, 16, 18, 19, 21,22), Microdrosophila (28). 
Liodrosophila (33, 34, 35), Scaptomyza (38), Drosophila (45, 48, 62, 67, 69, 76). 

East shore of River Arun, swept from Ricinus communis L, c. 1,800’. Scaptomyza 
(38). 

West shore of River Arun, evergreen shrubs bordering dry stream-beds, c. 1,800’. 
Leucophenga (14). 

River Sabhaya, west shore, fermented millet seeds on shore, c. 1,800’. Liodroso- 
phila (33, 35), Drosophila (59, 62, 67, 76). 

River Sabhaya, west shore, evergreen shrubs on sandy shore, c. 1,800’. Apsinota 
(5), Leucophenga (17), Liodrosophila (33), Drosophila (76). 

River Sabhaya, west shore, from rotten bananas in traps, c. 1,800’. Drosophila 
(70, 77). 


Tumlingtar 
Bare rocky slopes above River Sabhaya, west bank, c. 1,900’. Apsinota (5). 


Above River Sabhaya. 
East shore, swept from dwarf bamboos in deep ravine, c. 2,000’. Leucophenga 
(12, 15), Liodrosophila (33, 35), Diathoneura (36), Drosophila (67). 


NOTES ON SOME SPECIAL DIAGNOSTIC CHARACTERS 
A. ARCHESTINIC CHARACTERS 


‘ 


Stenzel (1963) proposed the terms “ archestinic”’ and “ archekastic’’ for the 
diagnostic generic characters which occur in some and all of the member taxa, 
respectively. The idea “ archestinic’”’ can be ascribed to the polythetic principle 


ce 


120 T. OKADA 


of the Neo-Adansonian school (Sneath, 1961) or of numerical taxonomy (Sokal and 
Sneath, 1963). Although much conflict and dispute has been raised between the 
orthodox or phylogenetic taxonomists and the Neo-Adansonians, the principle 
claiming overall similarity or using the “greatest possible number of characters’ in 
classification is unequivocally accepted by most of the current phylogenetic as well 
as numerical taxonomists. The archestinic character should be of taxonomic 
importance in this context (Kiriakoff, 1965 : 64 ; Mayr, 1965 : 73). 

Some examples of the archestinic characters can be cited from the present investi- 
gation. The subgenus Dichaetophora of the genus Drosophila has necessarily been 
characterized by the absence or abbreviation of the anterior dorsocentrals, while 
D. rotundicornis possesses rather well developed anterior dorsocentrals, which can be 
explained as archestinic. Likewise, the occurrence of six rows of acrostichal 
hairs in ChaetodrosopMilella coei should be archestinic, because the genus Chaetodroso- 
piulella shows essentially two or four rows of acrostichal hairs (Wheeler and Takada 
1964). Incidentally, accepting the overall similarity principle, the author agrees 
with L. H. Throckmorton (personal communication) to treat Drosophila subtilis 
Kikkawa and Peng, 1938, from Japan as a member of the subgenus Scaptodrosophila, 
although this species differs from ordinary species of Scaptodrosophila in having no 
prescutellars. 


B. HOMEOTIC EXPRESSION OF THE HOOKED SCALY BRISTLES 


The “ hooked scaly bristles ’’ (Okada, 1963) are special thick bristles of a secondary 
sexual character found on the male tibia and metatarsus of the mid leg in some 
melanogaster group species belonging to the suzuki, takahashii, and ficusphila 
subgroups and in Drosophila (Tanigastrella) gracilis (Duda, 1924) (Okada, 1964b). 
It was proved that amongst present material, D. (Sophophora) tristipennis and D. (S.) 
ammacularis, both belonging to the suzuki subgroup, also have such bristles. So 
far as examined by the author, the species possessing the hooked scaly bristles have, 
without exception, also sex-combs on the male fore legs, but the species having sex- 
combs do not always possess the hooked scaly bristles. This fact makes it plausible 
that the occurrence of the hooked scaly bristles pre-requires the possession of the sex- 
combs and that the development of the sex-combs on the fore leg homeotically 
induce the hooked scaly bristles on the homonymial organ, the mid leg. The excessive 
development of the sex-combs, however, seems to diminish the homeotic expression 
of the hooked scaly bristles through “‘ enantiomorphosis ”’ (Okada, 1960c) or through 
“material compensation ”’ (Rensch, 1954). The compensation would be manifested 
in two such structures with the same function as these two kinds of bristles of similar 
secondary sexual character. The same is true for the reversal correlation found by 
Prevosti (1954) between the number of sex-comb teeth and of clasper teeth among 
the species of the obscura group as well as among the natural populations of Droso- 
phila subobscura Collin, 1936 in Europe. 

The homeotic and compensatory developments of the hooked scaly bristles and 
sex-combs can be divided into five stages (Text-fig. 328). Stage 1. Both sex- 
combs and hooked scaly bristles are absent (willistont and mommat groups). Stage 


DIPTERA FROM NEPAL 121 


2. Sex-combs are present but the hooked scaly bristles absent (ananassae and 
melanogaster subgroups, obscura group). Stage 3. Sex-combs are better developed, 
inducing homeotic expression of the hooked scaly bristles (swzwkit and takahashit 
subgroups, D. (T.) gracilis). Stage 4. Sex-combs are further well developed, the 
hooked scaly bristles being compensatorily weakened in expression (/ficusphila 
subgroup). Stage 5. Further compensation is pronounced, resulting in disappear- 
ance or non-development of the hooked scaly bristles (montiwm and nipponica 
subgroups). 


Fig. 328 


Homeosis 


Compensation 


¢ HSB 


eee eee 


—< Character development— + 


(ita © Bled 5 | Stage 
Will anan suzu _- mont 
Momm (mela taka ‘4S nipp Taxon 


Obsc Mela 


Fic. 328. Homeotic and compensatory relationships between the hooked scaly bristles 
(HSB) of male mid leg and the sex-combs (SC) of male fore leg. Mela., melanogaster 
group ; Obsc., obscura group; Momm., mommai group; Wiill., willistoni group ; 
anan., ananassae subgroup ; /ficus., ficusphilasubgroup ; mela., melanogaster subgroup ; 
mont., montium subgroup; nipp., nipponica subgroup; suzu., suzuki subgroup ; 
taka., takahashu subgroup. For further explanations see text. 


C. HoMOLOGY IN THE COMPONENTS OF PHALLIC ORGANS AND THE SYSTEMATIC 
ANALYSIS OF PHALLIC FORMULAE 

As is frequently stressed, the phallic organs of insects are systematically and 

diagnostically essential key characters. Their extensive diversification, however, 

makes it often difficult to establish homology of their components even between 

closely related taxa. Those of Drosophilidae are not exceptional, e.g., the present 

author (1956) considered the ventral recurved process of aedeagus (u) of the genus 


122 T. OKADA 


Leucophenga (p. 17) to be the main part of the aedeagus. The present research 
revealed the real homology in the components of aedeagus between the genera 
Leucophenga and Mucrodrosophila. Microdrosophila migrohalterata (p. 41) and 
M. marginata (p. 42) have the ventral recurved process well developed as in Leuco- 
phenga, and the former species shows the posterior parameres contiguous to the 
subapical part of aedeagus (Text-fig. 117) as in Leucophenga. The facts indicate 
that these two genera are closely related, and this is confirmed by the analysis of 
phallic formulae. 

As a means to analyse phallic formulae the ideas of “‘ divergency index ”’ (d.i.) and 
“ difference value ”’ (d.v.) (Okada, 1954, 1956) have been taken into consideration. 
The d.i. is a sum of values 0, 0:5, 1-0, and 2:0 afforded to a large letter, small letter 
with dash, small letter, and small letter with attached y respectively, in the phallic 
formula. The phallic formula of a higher taxon is obtained by calculating mean value 
for each letter of the member species. The d.v. means a sum of differences in value 
for each of corresponding letters between two taxa. 

The phallic formula and d.i. thus determined for each genus or subgenus are as listed 
below. The d.v. between every two genera or subgenera is shown in Tables 3 and 4. 


G Number of 

enus Or : : E 
Sabeeons species phallic formula di. 

examined 

Cryptochaetum I ae BC id) Ete, Ee kL amen 9°5 
Diastata I al BiGe DEE eimec El ike ame Ni 6:0 
Campichoeta I iWs) (CG IDI Des Tek ie Ve Ibe WE IN, 3:0 
Apsinota I a Be@ des i gi bie kl SMa; 9:0 
Leucophenga se) ABC Die Ee Haske emis 5:0 
Microdrosophila 2 abaGrd ge hve. Eon eke ieee 8:5 
Liodrosophila 4 aBiCad ie faien iotnkasleem NN) 8-0 
Pavamycodrosophila I ab Cid Bei ye rio ke oe ms Ny 7°5 
Chaetodrosophilella I ab Ce eh co Ei Kola Ninn 5:0 
Scaptomyza 3 aD Caw aieron tl tke e mm 9°5 
Diathoneura I 2 Bi Cide Eye) Heike Lym iN 7:0 
Drosophila 21 ae cud weaterewih 12 kgelvsmieN 6°5 
Scaptodrosophila 3 AV BiGrdseh te orhielivk: ein 3°55 
Dichaetophora 2 ae Crdyemic egitiinle em n? 7:0 
Hirtodrosophila I AmB aC DEE t or il torte dee ving, 7:0 
Sophophora 7 SEBECeDr iE ft geht Mer vir Ni 3:0 
Drosophila s. str. 8 amin Crdime teen i ke iesmen? 8:5 


The genus Leucophenga shows d.v. with Microdrosophila 4:5, which is the lowest 
among the values shown between Leucophenga and any genus of Drosophilinae 
(Table 3). Although the d.v. between Leucophenga and Drosophila is also 4-5, this 
low value is due to high diversification of the phallic organs among the species of 
Drosophila. Indeed, except with the subgenus Scaptodrosophila, Leucophenga shows 
d.v. higher than 4:5 with each subgenus of Drosophila. 

The analysis of phallic formulae also indicates that among the genera of Diasta- 
tidae, Diastata is nearer to Campichoeta than to Apsinota, the d.v. between Dvastata 
and Campichoeta (3-0) being smaller than that between Campichoeta and Apsinota 


DIPTERA FROM NEPAL 123 


(6-0) and between Diastata and Apsinota (5-0), and that among the genera of 
Drosophilinae, Liodrosophila, Paramycodrosophila, and Chaetodrosophilella are 
mutually closely related and the subgenus Dichaetophora is closer to Drosophila s. str. 
than to any other subgenus of the genus Drosophila (Tables 3 and 4). 


TABLE 3 
The difference value (d.v.) of phallic formulae between every two genera. 
Cryptochaetum 0 
Diastata 6.5 0 
Campichoeta 8.5 3.0 0 
Apsinota 7.5,--5.0 6.0 0 
Leucophenga 8.5 3.0 2.0 6.0 0 


Microdrosophila 10.0 5.9 6.5 5.9 4.5 0 


Liodrosophila 4.5 4.0 7.0 3.0 T.0° 635 0 
Paramycodro- 
sophila 7.0 4.5 6.5 4.5 7S ~~ TO 2.5 0 
Chaetodroso- 
philella 7.5 5.0 6.0 4.0 8.0 6.5 3.0 2.5 0 
Scaptomyza 6.0 3.5 6.5 4.5 6.5 4.0 3.5 5.0 6.5 0 
Diathoneura 5.5 5.0 6.0 6.0 6.0 5.5 5.0 5.9 4.0 4.5 0 
Drosophila Tod 3.5 5,5 5.5 4.5 5.0 4.5 5.0 4.5 6.0 4.5 0 
4 | 
4 e 3 
=o | n <= 
5 2 ee ae 
a 3 = 8 a 3 3 N 5 S 
£ - 2 3 2 = 2 = 3 E 2 s 
8 = S i) a 3 ° E 3 i} 6 = 
8 [7 = ° ° fe = 2 = = 
a n = ° & so ot o a. = 2 
ae ee ee ee ee 
6 a oO < Sibel a a o a a a 
TABLE 4 


The difference value (d.v.) of phallic formulae between every two subgenera of the 
genus Drosophila, and between each subgenus and Leucophenga. 


Leucophenga 0 
Scaptodrosophila 3.5 0 
Dichaetophora 6.0 4.5 0 
Hirtodrosophila 8.0 7.5 7.9 0 
Sophophora 5.0 S.5 7.0 7.5 0 
Drosophila 7.5 5.5 1.5 7.5 7.5 0 
a 
a a % 
Aen eae a 
§ ° i) a 
oe © wy Oca ke 4s 
zB = 2 2 2 = 
os 8 a & o 
2 a = = = 4 
a.6hCUS 3 & a 5 
oO le} 
4 an Qa = a a 


124 Tt: OKADA 


REFERENCES 


BaspEN, E. B. 1952. Some Drosophilidae (Diptera) of the British Isles. Entomologist’s mon. 
Mag. 88 : 200-201. 
1954. The distribution and biology of Drosophilidae (Diptera) in Scotland, including a 
new species of ‘“‘ Dvosophila’’. Tvans. R. Soc. Edinb. 72 : 603-654, 2 pls., 9 text-figs. 
BaspENn, E. B. & Coin, J. E. 1958. On the examination of some Meigen and Zetterstedt 
specimens of Campichaeta Macquart (Diptera : Diastatidae). Proc. R. ent. Soc. Lond. (B) 
27 : 133-144, 5 text-figs. 

Bezz1,M. 1928. Diptera, Brachycera and Athericera of the Fiji Islands. viii + 220 pp., 54 figs. 
Brit. Mus. (Nat. Hist.) London. 

Burva, H. 1954. Zur Kenntnis der Drosophiliden der Elfenbeinkiiste (Franzdsisch West- 

Afrika). Rev. Suis. Zool. 61 (Suppl.) : 1-218, 224 text-figs. 

1954a. Distinction between four species of the melanogaster group, Drosophila seguyi 
D. montium, D. kikkawai n.n. and D. auraria (Drosophilidae, Diptera). Revta bras. Biol., 
Rio de Janeiro, 14 : 41-54, 19 figs. 

1956. Die Drosophilidengattung Zygothrica und ihre Beziehung zur Drosophila-Unter- 
gattung Hirtodrosophila. Mutt. zool. Mus. Berl. 32 : 189-321, 359 text-figs. 

Cuno, M. 1927. Genetic study of Japanese Drosophila (in Japanese). Zool. Mag., Tokyo, 
39 : 472-476. 

CoLiin, J. E. 1953. On the British species of Scaptomyza Hardy and Parascaptomyza Duda. 
Entomologist 86 : 148-151. 

CoguILLETT, D. W. 1898. Report on a collection of Japanese Diptera, presented to the U.S. 

National Museum by the Imperial University of Tokyo. Pyoc. U.S. natn. Mus., 21 : 301- 

340: 

1901. Three new species of Diptera. Ent. News 12 : 16-18. 

Dupa, O. 1921. Kritische Bemerkungen zur Gattung Scaptomyza Hardy (Diptera). Jh. 

Ver. schles. Insektenk. 13 : 57-69. 

1922. Liodvosophila und Sphaerogastrella, zwei neue, zu den Drosophiliden und nicht zu 
den Camilliden gehérige Dipteren-Gattungen aus Siidostasien. Avch. Naturgesch. 88 A (4) : 
150-160. 

1923. Die orientalischen und australischen Drosophiliden-Arten (Dipteren) des Ungari- 
schen National Museums zu Budapest. Avnls hist.-nat. Mus. natn. hung. 20 : 24-59. 

1924a. Beitrag zur Systematik der Drosophiliden unter besonderer Beriicksichtigung der 
palaarktischen u. orientalischen Arten (Dipteren). Avch. Naturgesch. 90 A (3) : 172-234, 
7 pis. 

1924b. Die Drosophiliden (Dipteren) des Deutschen Entomologischen Institutes der 
Kaiser Wilhelm-Gesellschaft (friih. Deutsches Entomolog. Mus.) aus H. Sauter’s Formosa 
Ausbeute. Ayvch. Naturgesch. 90 A (3) : 235-259. 

1925. Die Costaricanischen Drosophiliden des Ungarischen National Museums zu Buda- 
pest. Avnls. hist.-nat. Mus. natn. hung. 22 : 149-229. 

1926. Fauna Sumatrensis, Beitrag N. 26. Drosophilidae (Dipt.), Supplia ent. 14 : 42-116, 
23 text-figs. 

1928. Beitrag zur Kenntnis der siidostasiatischen Drosophilidengattung Hypselothyvea de 
Meijere (Dipt.). Anmnls hist.-nat. Mus. natn. hung. 25 : 79-90, 5 text-figs. 

1929. Fauna Buruana, Diptera, Acalyptrata (Overgedruktuit). Tveubia, 7 : 415-422. 

1931. Chloropidae, Drosophilidae. Dipterologischer Beitrag zu der von H. J. Feuerborn, 
F. Ruttner und A. Thienemann im Jahre 1928 und 1929 nach Java, Sumatra, und Bali 
untergenommenen limnologischen Forschungsreise. Avch. Hydrobiol., Suppl. 9 : 192-198. 

1934. Diastatidae, in Lindner, Die Fliegen dev Paldarkt. Reg. 58 e : 1-18, 2 pls., 10 text- 
figs. 

1935. Drosophilidae, in Lindner, Die Fliegen dev Paldarkt. Reg. 58 g : 1-118, 5 pls., 30 
text-figs. 


DIPTERA FROM NEPAL 125 


1939. Revision der afrikanischen Drosophiliden (Diptera) I. Avnmnls hist.-nat. Mus. natn. 
hung. 32 : 1-57, 7 text-figs. 

1940. Revision der afrikanischen Drosophiliden (Diptera) II. Avnmnls hist.-nat. Mus. 
natn. hung. 33 : 19-53, 6 text-figs. 

Durour,L. 1839. Mémoire sur les métamorphoses de plusieurs Larves fongivores appartenant 
a des Diptéres. Ann. Sci. Nat. Paris, 12 : 5-60. 

Epwarps, F. W. 1921. A revision of the mosquitoes of the Palaearctic Region. Bull. ent. 
Res. 12 : 263-351, 18 text-figs. 

Fapricius, J.C. 1787. Mantissa Insectorum, 2, 382 pp. 

FaLien, C. F. 1823. Diptera Sueciae, Geomyzides : 4, I. 

Foore, R. H. & Arnaup, P.H., Jr. 1958. Notes on the taxonomy and habits of Cryptochaetum 
nipponense (Tokunaga) in Japan. Proc. ent. Soc. Wash. 60 : 241-245, 5 text-figs. 

Frota-Pessoa, O. 1947. Revisao do género Clastopteromyia (em cuja sinonimia é colocada 

Diathoneura), com descrigao de 9 espécies novas. Summa bras. Biol. 1 : 181-221, 9 pls. 

1954. Revision of the tripunctata group of Drosophila with description of fifteen new 
species (Drosophilidae, Diptera). Avchos Mus. parana. 10 : 253-330, 11 pls. 

Hackman, W. 1955. On the genera Scaptomyza Hardy and Parascaptomyza Duda (Dipt., 

Drosophilidae). Notul. ent. 35 : 74-91, 32 text-figs. 

1959. On the genus Scaptomyza Hardy (Dipt., Drosophilidae) with descriptions of new 
species from various parts of the world. Acta zool. fenn. 97 : 1-73, 72 text-figs. 

Harpy, J. 1849. Note on Remedies for the Turnip-fly amongst the Ancients and on the 
Turnip-fly of New Holland, with Notice of a New Genus and Species of Diptera. Proc. 
Berwickshire Nat. Club, 2 : 359-362. 

HENDEL, F. 1914. Acalyptrate Musciden (Dipt.), 3. Suppl. ent. 3 : 90-117. 

Hsu, T.C. 1949. The external genital apparatus of male Drosophilidae in relation to system- 
atics. Univ. Texs Publs 4920 : 80-142, 18 pls., 1 text-fig. 

Kana, Y. S., Counc, O. K. & LEE, H. Y. 1959. Studies on the classification and the living 
conditions of Drosophilidae in Korea (2). ‘‘ On the unrecorded species of Drosophila flies 
in Korea’’. Korean J. Zool. 2 : 61-65. 

Kixkawa, H. & PENG, F. T. 1938. Drosophila species of Japan and adjacent localities. 
Jap. J. Zool. 7 : 507-552, 6 pls., 29 text-figs. 

Kim, K. W. 1963. Drosophilidae of Tol-San island, Korea. Dyvosoph. Inf. Serv. 38 : 73. 

KrriaAkorFr, S. G. 1956. Some remarks on Sokal and Smeath’s Principles of Numerical 
Taxonomy. Syst. Zool. 14 : 61-64. 

Kuwana,I. 1922. Studies on Japanese Monophlebinae. Bull. Dept. Pl. Quarant. Stn 1 : 1-58; 
2 : 1-43, 4 pls. 

Lams, C. B. 1914. Reports of the Percy Sladen Trust Expedition, Diptera, No. 15. Hetero- 
neuridae, Ortalidae, Trypetidae, Sepsidae, Micropezidae, Drosophilidae, Geomyzidae, 
Milichidae. Tvans. Linn. Soc. Lond., (2) 16 : 307-372, 3 pls., 48 text-figs. 

Ler, T. J. 1964. New and unrecorded species of Drosophilidae in Korea. Korean J. Zool. 
7 : 37-40, 3 text-figs. 

Lorw, H. 1864. Ueber die europadischen Arten der Gattung Diastata. Berl. ent. Z. 8 : 357- 
368. 

Macguart, M. 1833. AHistoive naturelle des Insectes (Diptéves), 2 : 710 pp. 

MarExkawa, F. 1963. Trans-palaeo-equatorial and bipolar distribution. Sci. Rep. Tohoku 
Univ. ser. IV, 29 : 355-363, 9 text-figs. 

Mattocu, J. R. 1921. Some notes on Drosophilidae (Diptera). Ent. News. 32 : 311-312. 

1934. Diptera of Patagonia and South Chile based mainly on material in the British Museum 

(Natural History). Part 6 (5), Acalypterata (concluded) : 393—489, rt pl. 

MaTSuMURA, S. 1931. Six thousand illustrated insects of Japan Empire. \xii + 1694 pp., 
to pls., ca. 6000 text-figs. Tokyo. 

Mayer, K. 1935. Die Metamorphose einiger Drosophiliden aus Niederlandisch-Indien. 

Arch. Hydrobiol., Suppl. 13 (5) : 462-473, 19 text-figs. 


126 T. OKADA 


Mayr, E. 1965. Numerical phenetics and taxonomic theory. Syst. Zool. 14 : 73-79. 

MEIGEN, J. W. 1830. Systematische Beschreibung dev bekannten Euvopdischen zweifliigeligen 
Insekten, 6: 401 pp. Hamm. 

MEIJERE, J.C.H.pDrE. 1906. Uber einige indo-australische Dipteren des Ungarischen National 
Museums, bez. des Naturhistorischen Museums zu Genua. Annls hist.-nat. Mus. natn. 
hung. 4 : 193-194. 

— 1908. Studien iiber Siidostasiatische Dipteren, II. Tijdschr. Ent. 51 : 106-180, 4 pls. 

—g11I. Studien tiber Siidostasiatische Dipteren, VI. Tijdschr. Ent. 54 : 258-432, 5 pls. 

1914. Studien iiber Siidostasiatische Dipteren, IX. Tijdschr. Ent. 57 : 137-275, 7 pls. 

Mik, J. 1886. Dipterologische Miscellen. Ween. ent. Ztg. 5 : 317. 

Nater, H. 1953. Vergleichend-morphologische Untersuchung des ausseren Geschlechtappa- 
rates innerhalb der Gattung Drosophila. Zool. Jb. (Syst.), 81 : 437-486, 33 text-figs. 
Oxapa,T. 1954. Comparative morphology of the drosophilid flies. I. Phallic organs of the 

melanogaster species group. Kontyt, 22 : 36-46, 2 pls. 

1955. Fauna and Flora of Nepal Himalaya. Drosophila. Scient. Results Jap. Exped. Nepal 

Himalaya, 1952-1953, 1 : 387-390, 1 pl. 

1956. Systematic Study of Drosophilidae and allied families of Japan. 183 pp., 83 text- 
figs., Gihddo, Tokyo. 

1960a. Thegenus Microdrosophila Malloch from Japan (Diptera, Drosophilidae). Kontyi, 
28 : 211-222, 1 pl., 3 text-figs. 

1960b. Notes on Diastatidae from Japan (Diptera), Kontyi, 28, : 165-168, 2 text-figs. 
1960c. Allomorphosis of the wing-veins vs. wing-membranes in insects. Nat. Sci. Rep 

Ochanomizu Univ., 11 : 35-50, 24 text-figs. 

1963. Hooked scaly bristles found on the male mid legs in some drosophilid flies. Dvrosoph. 

Inf. Serv. 38 : 38-39, 4 text-figs. 

1964a. New and unrecorded species of Drosophilidae in the Amami Islands, Japan. 
Kontyt, 32 : 105-115, 13 text-figs. 
1964b. Drosophilidae (Diptera) of South-east Asia, collected by the Thai-Japanese 

Biological Expedition 1961-62. Nature and Life in South-east Asia. 3 : 439-466, 8 text- 

figs. 

Oxaba, T. & CHuNG, Y. J. 1960. Three species of Drosophilidae from South Korea. Akitu, 
9 : 25-30, 3 text-figs. 

Oxapa, T. & Kurokawa, H. 1957. New or little known species of Drosophilidae of Japan 
(Diptera). Kontyi, 25 : 2-12, 2 text-figs. 

Oxapba, T. & Sasakawa, M. 1956. Leaf-mining species of Drosophilidae (Diptera). <Azkztu, 
5 : 25-28, 7 text-figs. 

Paik, Y. K. 1958. Seasonal changes in Dvosophila populations at the two adjacent areas in 
Korea. Commem. Theses sixtieth Birthday Dr. C. C. Woo. Apr. 1958 : 209-227. 

PenG, F. T. 1937. On some species of Drosophila from China. Annotnes zool. jap. 16 : 20-27, 
5 text-figs. 

Prevosti, A. 1954. Genetical variability in natural population of Drosophila subobscura. 
Atti IX Congr. internatn. genet. Bellagio (Como) 660-663. 

RenscuH, B. 1954. Neuere Probleme dey Abstammungslehre, die transspezifische Evolution. 
2nd ed., xi + 436 pp., 113 text-figs., Ferdinand Enke, Stuttgart. 

RoONDANI, C. 1875. Species Italicae ordinis Dipterorum. Boll. Soc. ent. ital. 7 : 166-191. 

Scoporl, G. A. 1763. Entomologia Carniolica, 420 pp. Vindobonae. 

Sokal, R. R. & SneatH, P.H. A. 1963. Principles of numerical Taxonomy. xvi + 359 pp., 
38 text-figs. Freeman & Co., San Francisco and London. 

SNEATH, P. H. A. 1961. Recent developments in theoretical and quantitative taxonomy. 
Syst. Zool. 10 : 118-139, 5 text-figs. 

STENZEL, H. B. 1963. A generic character, can it be lacking in a given Genus? Syst. Zool. 
12 : 118-121, 2 text-figs. 


DIPTERA FROM NEPAL 127 


StuRTEVANT, A. H. 1916. Notes on North American Drosophilidae with descriptions of 23 

new species. Ann. ent. Soc. Am. 9 : 323-343. 

1921. The North American species of Drosophila. Publs Carnegie Instn. 301 : 1-150, 

3 pls., 49 text-figs. 

1927. Philippine and other oriental Drosophilidae. Philipp. J. Sci. 32 : 361-374. 
1939. Onthe subdivision of the genus Drosophila. Proc. natn. Acad. Sci. U.S.A.25 : 137- 

141. 

1942. The classification of the genus Drosophila, with descriptions of nine new species. 

Univ. Texas Publs, 4213 : 5-51. 

Taxapa, H. & Lee, J. J. 1958. <A preliminary survey of the Drosophilidae from Kongju and 
its adjacent localities, South Korea. Annotnes zool. jap. 31 : 113-116, 3 text-figs. 

Taxapa, H. & Oxapa, T. 1960. Drosophila survey of Hokkaido XI. A new species of 
Drosophila (Sophophora) from Japan. Annotnes zool. jap. 33 : 142-145, 5 text-figs. 

Tan, C.C., Hsu, T.C. & SHENG, T.C. 1949. Known Drosophila species in China with descrip- 
tions of twelve new species. Univ. Texas Publs, 4920 : 196-206. 

Toxunaca, M. 1943. Medical Entomology (Iyé Konchtigaku) Vol. 2, ii + 727 pp., 250 text- 
figs. Kanehara Co. Ltd., Tokyo. 

WakKAHAMA, K. 1964. Some ecological studies in Drosophila. Bull. Shimane Univ. (Nat. Sci.), 
No. 13 : 50-122, 64 text-figs. 

WaAKAHAMA, K., KANEKO, A. & Tokumitsu, T. 1963. Dyrosophila survey of Hokkaido XIX. 
Collection records in three localities of Hokkaido, with a note on the diurnal behaviour of 
certain species. Amnnotnes zool. jap. 36 : 92-96, 2 text-figs. 

WHEELER, M. R. 1952. The Drosophilidae of the Nearctic Region, exclusive of the genus 

Drosophila. Univ. Texas Publs, 5204 : 162-218, 1 text-fig. 

1957. Taxonomic and distributional studies of Nearctic and Neotropical Drosophilidae. 

Univ. Texas Publs, 5721 : 97-114, 25 text-figs. 

1959. Anomenclatural study of the genus Drosophila. Univ. Texas Publs, 5914 : 181-205, 

I text-fig. 

1960. A new subgenus and species of Stegana Meigen (Diptera : Drosophilidae). Proc. 
ent. Soc. Wash. 62 : tog-111, 1 text-fig. 

WHEELER, M. R. & Takapa, H. 1964. Diptera : Drosophilidae. Insects of Micronesia, 14, 
No. 6 : 163-242, 28 text-figs, 1 map. 

Wutp, F. M. vAN DER. 1887. Aanteekeningen betreffende Javaansche Diptera. Tijdschr. 
Ent. 30 : 175-180. 

ZETTERSTEDT, J. W. 1847. Diptera Scandinaviae, 6 : 2163-2580. 


INDEX 


Synonyms printed in italics 


aberrans, 73 Apsinota, 8, I1 
abbreviata, 17, 18 argentata, 17, 24 
acutissima, 96, IOI auraria, 95 

aerea, 53 

agbo, 73 basdeni, 8 

albiceps, 18, 33, 36 bimaculata, 48 
albicincta 27 bipectinata, 82, 95 
albifascia 17, 22 bizonata, 99, I13 
alternata, 67, 72 bizonata-group, 96, 97 
ananassae-subgroup, 95 brunnea, 67 


annulipes, 96, 107 busckii, 81 


128 INDEX 


Campichoeta, 8, 10 Hirtodrosophila, 66, 77 
Chaetodrosophila 57 histrio, 99 
Chaetodrosophilella, 15, 57 histrio-group, 96, 99 
circumdata, 107 hypocausta, III 
Clastopteromyia, 66 Hypselothyrea, 15, 47 
coei (Chaetodrosophilella) 57 
coei (Sophophora), 81, 82, 84 immacularis, 82, 87 
coeruleifrons, 48 immigrans, 96, 112 
coleoptrata, 15 immigrans-group, 96, 105 
coracina, 70 interrupta, 17, 21 
costata, 17, 19 
cristata, 40 kikkawai, 82, 92 
Cryptochaetidae, 6 
Cryptochaetum, 6 lacertosa, 96, 97 
cubicivittata, 96, 105 Leucophenga, 14, 17 
curvipennis, 15 leucozona, 17, 21 
cuvvipila, 30 limbicostata, 77, 97 
Liodrosophila, 15, 47, 48 
denticeps, 79 Lissocephala, 45 
Diastata, 8 Lissodrosophila, 15, 45 
Diastatidae, 8 longecrinita, 77 
Diathoneura, 14, 65 longicornis, 45 
Dichaetophora, 66, 73 
dimidiata, 47 maculata (Drosophila),17 
disticha, 59 maculata (Microdrosophila), 43 
Dorsilopha, 81 maculiventris, 30 
Dorsophila, 66 magnidentata, 73 
Drosophila, 15, 66 marginata (Microdrosophila), 37, 42 
Drosophila s. str., 66, 96 marginata (Scaptodrosophila), 67, 70 
Drosophilidae, I4 marginifrons, 54 
Drosophilinae, 04, 37 meijerea, 30 
meieret, 36 
editinares, 96, IOI, 103 melanissima, 58, 62 
melanogaster, 81, 92 
flavipalpis, 34 melanogaster-group, 81, 87 
flexicauda, 96, IoI melanogaster-subgroup, 92 
fruhstorferi, 86 Microdrosophila, 14, 37, 40, 69 
funebris, 66, 96 minima, 67, 69 
fuscicostata, 96, III mommai, 83 
fuscipennis (Leucophenga), 30 mommai-group, 81, 82 
fuscipennis (Liodrosophila), 48, 51 montium, 92 
montium-subgroup, 92 
grahami, 63 
graminum, 58, 59, 62 nasuta, 96, I12 
grandicorne, 6 nepalensis, 81, 91 
grandis, 101, 105 nigriceps, 18, 31 
grandis-group, 96, IOI nigrifrons, 15 
guttata, 47 nigripalpis (Microdrosophila), 37, 41, 42 
guttiventris, 18, 30 nigripalpis (Leucophenga), 18, 28 
nigrita, 65 
halteropunctata, 17, 25 nigrohalterata, 37, 41 
hexastriata, 109 nigroscutellata, 18, 19, 34 


hirticornis, 79 nipponense, 6 


‘ie 


nitida, 48, 54 
notostriata, 96, 107, I10 


obscuripennis, 10 
obscuripes, 13 
onychopyga, 48, 50 
ornata, 17, 21 
ornatipennis, 21 
Oxystyloptera, 39, 40 


pallida, 58, 59 
Pavadrosophila, 67 
Paramycodrosophila, 15, 55 
Parascaptomyza, 58, 59 
parasplendens, 58, 59 
pauciramosa, 37, 40 
pectinata, 37 
pentastriata, 96, 109 
Pholadoris, 67 
pictipennis, 67 
pictiventris, II, 13 
pictula, 55 

pilicrus, 67 

pulchrella, 81, 89 
purpurata, 41 


quadrata, 37 
quadrilineata, 58 
quinaria-section, 96, 97 


raridentata, 73, 74, 76 
rectinervis, 18, 33 
Tipa, 65 
robusta-group, 96, 97 
rotundicornis, 74 
rufipes, 11 

rugulosa, 51 


INDEX 129 


salatigae, 18, 27 
Scaptomyza 15, 58, 59 
Scaptodrosophila, 66, 67 
scaptomyzoidea, 67 
scutellimargo, 67 
serriflabella, 81, 84 
silvata, 58, 63 
Sophophora, 66, 81 

sp., Microdrosophila, 39 
Stegana, 14, 15 
Steganina, 15 
Steganinae, 14, 15 
sternopleuralis, 99, 100, IOI 
Styloptera, 40 
subpollinosa, 17, 21 
subsplendens, 61 

suruk, 73, 76 

suzuki, 87 
suzukii-subgroup, 87 


takahashii, 82, 92 
takahashii-subgroup, 92 
taeniatipennis, 65 
tectifrons, 39 
tenuicauda, IOI, 103 
trapezifrons, 82, 93 
tripartita, 77, 79 
tripunctata-group, 109 
trisetosa, 96, 99 
tristipennis, 81, 89 
trizonata, 96, 97 


ussurica, 8 
vagans, Io 
virgata, 110 


virilis-section, 96, 97 


zonaria, 81, 83 


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A YLIST, OF) SURPEEMENTS 
TO... THE, ENTOMOLEOGICAL- SERIES 
OF ‘THE? BULLEGIN’ OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 


. 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. 

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

. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177 : 
18 plates, 270 Text-figures. August, 1965. £4 4s. 

. SanpDs, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172: 500 Text-figures. September, 
1965. £3 5S. 

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


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