P’REPACE.

A PRACTICAL and detailed knowledge of Tsetse- is essential for the material progress of a large part of Africa. How large a part is affected by these formidable -pests may be estimated by a glance at the Map which appears in this “ Handbook.” In order to settle the grave questions which are connected with the transmission of certain insect-borne diseases, it is not sufficient to know that a giveu insect belongs to the * Glosaina-or, in other words, is a Tsetse--but, as indicated by the author in his Introduction, it may be of vital importance to know with certainty the species to which it belongs. ‘I. The study of the Tsetse-flies was greatly advanced by the ,+ appearance of Mr. Austen’s Cc Monograph of the Tsetse-Flies,” c: which was published by order of the Trustees of the British ii Museum in 1903. That work has been out of print for some time, and moreover our knowledge of the genus Glossina has considerably increased, during the interval. The present “ Hand- book,” by the same author, is an attempt to bring the subject up to date ; and it is hoped that it has been drawn up in such a way that, in addition to being a reliable contribution to systematic literature, it will prove useful to medical officers and others engaged in the struggle with disease in Africa.

SIDNEY F. H,4RMER, Keeper of Zoology. \

BRITISFE MUSEUM (NATURAL HISTORY), LONDON, S.W. March 18th, 1911. TABLE OF CONTENTS.

PREFACE ...... "'111

LIST OF PLATES ...... vi

LIST OB FIGURES IN THE TEXT ...... vii

INTRODUCTION ...... ix

CHAPTER I. THE GENERAL CHARACTERISTICS AND DISTRIBUTION OB TSETSR- FLIES (GENUS GLOSSINA) ...... 1

II. THE EXTERNAL CHARACTERS OF THE GENUS GLOSSINA . . 9

III. THE FOUR GROUPS INTO WHICH TSETSE-FLIES ARE DIVIDED. -TABLES FOR THE 'DETERMINATION OF GROUPS AXD SPECIES ...... 18

IV. THE GLOSSINA PALPALIS GROUP . . . . . 23

V. THE GLOSSINA MORSITANS GROUP . . . . . 48

VI. THE GLOSSINA FUSCA GROUP...... 69

VII. THE GLOSSINA BREVIPAI,PIS GROUP . . . . . 85 ' LIST OF PLATES. LI$

FIQ. PLATE 1. A Tsetse-f l%ontispiece attitu I. Glossina palpalis, Robineau-Desvoidy. ( d . x 6) . gram] II. Glossinu caliginea, Austen. ( 0 . x 6) . . . . At end 2. Sto?noxys of the III. Glossinapallicera, Bigot. ( 8 . x 6) . . . . ,, 3. HaeyhF;; IV. Gloss&a tacAinoides, Westwood. ( 2 . x 6) . . . ,, x 4) V. Gloss&a rvwrsitams, Westwood. (2 . x 6) . . . ,, 4. Pupa of G terior VI. Glossinapallidipes, Austen. ( 2. x 6) . . . . ,, %ifi VII. Glossina longipalpis, Wiedemann. ( 6. x 6) . . ,, 5. Pupae of A, G. VIII. Glossinafusca, Walker. ( 2. x 6) . . . . ,, extre Austt IX. Gloss&a brevipalpis, Newstead. ( 6. x 6) . . . ,, Desv X. Glossina longipen?zis, Corti. ( 0. x 6) . . . . ,, 6. Pupae of I speci New5 Map of Africa south of the Sahara, illustrating the distri- missi bution of Glossina ...... facing page 8 E, G. enlar 7. Diagram I used Norm-The crossed lines indicate the natural size (length and wing- expanse) ; the vertical line includes the palpi. 8. Left ante Students who are not trained entomologists sometimes find it difficult side to realise the natural size and appearance of an insect from a greatly not T enlarged figure. By viewing such a figure through a pocket lens, held at joint a distance from the object considerably greater than the focal length for magnification, and also at a distance from the eye, there can be obtained 9. Diagram a reduced image of the illustration, which can be made to approximate 10. Diagram very closely to the natural size of the insect. The desired result will be facilitated if the length of the cross-line indicating the wing expanse, 11. Antenna I and that of the line representing the length of the fly, be first compared with the diameter of the lens; before beiug examined in this way, a 12. Antenna I figure should, of course, be inverted. 13. Antenna 14. Distal PI (ven abov LIST OF FIGURES IN THE TEXT.

1. A Tsetse-fly, Gloss&a Zongipelznis, Corti (Somaliland), in resting ispiece attitude, showing the position of the wings. (Partly dia- grammatic. x 4) ...... 2 4t end 2. Stomoxys calcitrans, L., in resting attitude, showing the position of the wings. (Partly diagrammatic. x 4) _ . . 3 ,, 3. Haematopota &tata, Lw. (Tropical Africa), in resting attitude, showing the position of the wings. (Partly diagrammatic. I> x4) ...... 3

,, 4. Pupa of Glossina pallidipes, Austen, dorsal aspect (x 12) : a, pos- terior extremity, showing pit and (st) right larval stigma 0 (x 24) ; b, anterior extremity, showing longitudinal seam, which opens to permit the escape of the fly (x 8) . . 5 I, 5. Pupae of six species of Tsetse-flies, dorsal aspect, natural sise.- A, G. brevipalpis, Newst. ; B, G. fusca, Walk. (cephalic (lower) II extremity missing) ; C, G. morsitans, Westw.; D, G.palZidipes, Austen; E, G. tachinoides, We&w.; F, G. palpalis, Rob.- >, Desv. (From photographs of actual specimens) . . . 5

,, 6. Pupae of six species of Tsetse-flies, dorsal aspect (x 6) : the same specimens as those represented in Fig. 5.--A, G. brevipalpis, Newst. : B. G. fz~sca. Walk. (cenhalic (lower) extremitv We&w. ; D, G. pallidipes, Austen ; Iage S F, G.paZpaZis, Rob.-Desv. (From enlargements of photographs of actual specimens) . . 7. Diagram showing nomenclature of external characters of Gbssina, used in description ...... wing- 8. Left antenna of Gloss&a pallidipes, Austen, 6 , from the inner Bicu1t side (x 60) ; the very minute proximal joint of the arista is reatly not visible in this view ; p, aperture of sense-organ on third Bid at joint of antenna ...... 12 #h for ained 9. Diagram of thoracic chaetotaxy of Glossina, dorsal aspect (x 10) 16 imate fill be 10. Diagram of thoracic chaetotaxy of Glossina, pleural aspect (x 10) 17 mnse, 11. Antenna of Gloss&a caliginea, Austen (x 30) . . . 31 pared ‘a~, a 1 12. Antenna of Glossina palpalis, Rob.-Desv. (x 30) . . . 31 13. Antenna of Glossilza palliceva, Bigot. (x 30) . . . . 36 i ., ’ 14. Distal portion of abdomen of Glossina morsitans, We&w., 6 (ventral aspect), showing (h) hypopygium and the hectors :. aboveit (x 15)...... Vlll LIST OF FIGURES IN THE TEXT. FIQ. PdQE 15. Distal portion of abdomen of Glossina longipalpis, Wied., d (ventral aspect), showing (h) hypopygium and the hectors aboveit (x 15)...... 51 16. Antenna of Glossi%a fusca, Walk. (x 30) . . . . . 70 17. Distal extremity of & abdomen of (A) Gloss&a fusca, Walk., and (B) G. fuscipkuris, sp. nov. (ventral aspect), showing (h) hypopygia and (hect.) hectors (x 15) . . . . . 76 THE ( 18. Antenna of GZossi?za Izigvofusca, Newst. (x 30) . . . . 75 Of vital imp0 ii 19. Antenna of Glossina taba?zifoymis, Westw. (x 30) . . . 82 Tropical Afr best quahfiet 20. Heads of males of (A) Glossina breuipalpis, Newst., (B) G. fusca, Walk., and (C) G. l%edicorunz, sp. nov., dorsal aspect; disseminated showing differences in the size and shape of the head as a palpalis. 1% whole, in the shape of the eyes, and in the length of the palpi(x 10) ...... 86 cases of the c of the Luang 21. Heads of females of (A) Glossina b~e~ipalpis, Newst., (B) G. fusca, Walk., and (C) G. medico?um, sp. nov., dorsal aspect; localities GI( showing differences in the size and shape of the head as caused grave a whole, in the shape of the eyes, and in the length of the palpi(x 10) ...... 87 ( Glossina mar time been a~ 22. A Tsetse-fly (Glossina ??zovsitans, Westw., 9 ), before feeding (X 6) ‘33 somiases of , 23. A Tsetse-fly (Glossina morsitans, Westw., P), after feeding, several specie showing abdomen distended with blood (x 6). From a drawing from life, kindly lent by Colonel Sir David Bruce, possibility tk C.B., F.R.S., A.M.S...... 93 trypanosomia 24. Distal extremity of 6 abdomen of (A) Glossina bl-euipalpis, insect vector Newst., and (B) G. nzedicoru?n, sp. nov. (ventral aspect), discovery of showing (h) hypopygia and (hect.) hectors (x 15) . . 99 appeared to authorities bm from Trypanc point out tha agent of the than one spec be increased. Medical Office a ready mea Tsetse-flies fa present volun In the eig the author’s ‘ these insects, his collaborat Todd, the Iat6 Roubaud, and which is now ‘EXT. P‘IQE ris, Wied., d 1 the hectors . . . 51 INTRODUCTION. . . . 70 !z, Walk., and -c- showing (h) . . 76 TIZE insects described and illustrated in the followings pages are . . . 7s of vital importance to the prosperity and future development of . . 7 a2 Tropical Africa. Until quite lately it was considered by those best qualified to form an opinion that Sleeping Sickness is (B) G. fusea, lrsal aspect ; disseminated solely by the species of Tsetse-fly known as Glossilsa he head as a palpalis. The recent occurrence, however, of a number of isolated ength of the . . . 86 cases of the disease in the Nyasaland Protectorate and the valley of the Luangwrt River, North-Eastern Rhodesia, in both of which (B) G. fusca, rrsal aspect ; localities G’lossina palpalis is believed to be non-existent, has the head as caused grave suspicion to attach to two other species of Tsetse ength of the . . 87 (Glossina worsifans and G. brevipalpis). Experts have for some time been aware that Nagana or other closely similar trypano- feeding (x 6) Y3 somiases of domestic can be conveyed by the bites of fter feeding, several species of Tsetse-flies, in&din 6). From a g Glosuiua yalpalis, and the 3avid Bruce, possibility that Trypcmosonza gan&iense, t,he parasite of human . . . 93 trypanosomiasis, or Sleeping Sickness, may have more than one breviyalpis, insect vector naturally suggested itself ; yet, until the alarming ltral aspect), discovery of the cases referred to, the results of observation 5) . 99 appeared to negative any such conclusion. \Vhet#her, as cert’ain authorities believe, the R,hodesian parasite is specifically distinct from Trypmosoma ~a~6ienue or not, it is here only necessary to point out that should such a diff’erentiation in the actual causative agent of the disease be established, the probability that more than one species of Glossina is acting as a carrier would thereby be increased. In any case, it is obviously desirable that Colonial Medical Officers and other officials in Tropical Africa should have a ready means of recognising and accurately identifying the Tsetse-tiies found in their districts, and it is hoped that the present volume may serve as a means to this end. In the eight years that have elapsed since the publication of the author’s “ Monograph of the Tsetse-Flies ” our knowledge of these insects, thanks to t)he researches of Sir David Bruce and his collaborators, Professors Minchin and Newstead, Dr. J. L. Todd, the late Dr. Koch, Drs. Stuhlmann and Kleine, Mons. E. Roubaud, and others, has enormously increased. The 3Iono,rrraph, which is now out of print, contained descriptions and illustrations u

c X INTRODUCTION.

of seven species. In the following pages that number is increased to fifteen, including two species here described for the first time. At the time of the publication of the Monograph the con- nection of Glossina palpalis with Sleeping Sickness had not yet been established, and practically nothing was known as to the habits of that particular Tsetse, upon which a host of observers have since concentrated their attention. Thus, although much space in the Monograph was devoted to the subject of “ Bio- nomics,” the majority of the statements given referred to Gloss&a morsitans, the pest of the elephant-hunter and pioneer in South Africa, and the species to which the name “ Tsetse ” was originally applied. No observations worth mentioning on any other species had in fact been made, and what was known as to Gtossina morsitans had consequently to do duty for the genus Glossina as a whole. We now know that haunts and habits are not necessarily the same in all species, and in the present work a summary of the ascertained facts with regard to “ Bionomics ” is given under the heading of each species.* It will be seen that in the case of certain species much-in some instances everything-has yet to be learnt. There is thus a wide field for further observation, and it need hardly be said that information from those in a position to supply it will gladly be received. At the present time a sub-committee appointed by the Entomological Research Committee (Tropical Africa) is endeavouring, by means of series of carefully framed questions addressed to reliable observers on the spot, to obtain accurate information as to the habits, dependence or otherwise upon big game, etc., of Gloss&a morsitans, matters about which much uncertainty still exists. When all the replies that it is hoped to receive in this way are collated, a mass of useful knowledge should be acquired con- cerning a species which is one of the greatest enemies of the stock-breeder in Rhodesia and elsewhere. In conclusion, it need only be added that prophylactic measures in the fight against Tsetse-flies, to which attention is now being devoted, are not referred to, since they are beyond the scope of this work. ERNEST E. AUSTEN.

BRITISH MUSEUN (NATURAL HISTORY), r LONDON, S.W. MarcF, 16tk, 1911.

* Except 0. pulpalis--see page 27. number is increased I for the first time. lonograph the con- :kness had not yet 3 known as to the & host of observers us, although much 3 subject of “ Bio- referred to Glossina 1 pioneer in South ltse ” was originally n any other species wn as to Glossina gnus Glossina as a - are not necessarily .ork a summary of its ” is given under CHAPTER I. that in the case of rthing-has yet to THE GENERAL CHARACTERISTICS AND DISTRI- lrther observation, BUTION OF TSETSE-FLIES (GENUS GLOSSINA). 1 from those in a At the present The insects forming the subject of this volume General Characters nological Research ,,f Tsetse-fiies : how may be described as ordinary-looking dark 3y means of series to distinguish them brown, blackish, yellowish-brown, or yellowish reliable observers from other flies. flies, varying in length from about 6 or 8 as to the habits, millimetres, as in the case of Glossina ta&nodes, to as much as etc., of Glossinn 13 or 13.5 millimetres in that of G. brevipalpis or longipennis,* dnty still exists. with a prominent proboscis in all species. The hinder half of the ve in this way are body, or abdomen, in the species belonging to the Glossina be acquired con- morsitans group and in G. tachinoides is of a paler colour and marked st enemies of the with sharply defined dark brown bands, which are interrupted on the middle line ; the abdomen, however, is invisible when the phylactic measures insect is at rest, as it is then concealed by the wings. The sexes Ition is now being of Tsetse-flies can readily be distinguished when specimens can lyond the scope of be examined, since in the male the external genitalia form a conspicuous knob-like protuberance (hypopygium) beneath the : E. AUSTEN. end of the abdomen (see Figs. 14 and 15, pp. 50, 51), which is absent in the female. * These measurements are only from the head to the end of the body snd are exclusive of the proboscis and palpi, which project horizontally in front of the head. In bhe crossed lines on the coloured plates, however, the vertical line indicates the average length of the whole insect, inclz~ding tL proboscis, the transverse line showing the wing-expanse. n2 2 DISTINCTIVE CHARACTER3 OF TSETSE-FLIES. GLOSS13

It is probable that only those who have suffered from the can hardly I attacks of Tset,se-flies can recognise them when on the wing, but blood-sucker in the resting position their identification is easy. In this attitude animals vE they can be distinguished from all other blood-sucking Diptcra, Stomoxyys al with which confusion could possibly take place (especially from boscis, it it those belonging to the genera Stomoxys, Fig. 2, and Haematopotu, palpi, and Fig. 3, which are most likely to be mistaken for them), by the more slende fact that t,he wigs lie closed jlat over one another down the back, proboscis o’ of the fol greyish, gr flies, and s the resting one over t angle (see E domestica, : them from ’ on the othc horse-flies ( dantly reprl resembles ( the species

Fig. 1. -4 Tsetse-ily, Glossina Zon~ipe?rni~, Corti (Somaliland), in resting attitude, showing the position of the wings. (Partly diagrammatic. x 4.)

like the blades of a pair of scissors, while the proboscis (i.e. the proboscis ensheathed in the palpi) projects horizontally in front of the head (see Fig. 1). Apart from the prominent proboscis and the mode of carrying c the wings when at rest, there is nothing in any way remarkable

or striking about the appearance of a Tsetse, and the descriptions Haematopotl of most African travellers emphasise this fact. As already of the wir mentioned, species of Stomoxys and Haematopota are most likely as Tsetse-f to be mistaken for Glossinn, and apart from these confusion one over t p TSETSE-FLIES. h GLOSSINA, STOMOXYS, AND HAEMATOPOTA. 3 e suffered from the can hardly take place. The members of both genera are greedy en on the wing, but blood-suckers, and often torment human beings and domestic 99. In this attitude animals very greatly. Although od-sucking Diptera, Stomoxys also has a prominent pro- !ace (especially from boscis, it is not ensheathed in the 2, and Haematopotcr, palpi, and is consequently much \4 n for them), by the more slender in appearance than the dher down the back, proboscis of Glossina. The species I of the former genus are usually > greyish, greyish-black, or blackish flies, and since their wings when in / the resting position, instead of closing one over the other, diverge at an angle (see Fig. 2), like those of Musccc domestica, it is easy to distinguish them from Tsetse-flies. Haematopota, Pig. 2. on the other hand, a genus of small stomzys ~azcitm~s, L., ill rest+ attitude, showing th --‘L’ horse-flies (Family Tabanidae) abun- the xv++ (partly d dantly represented in Tropical Africa, x 4.) resembles Glossina somewhat closely when at rest. Although the species of this genus are of much the same elongate shape

resting attitude, natic. x 4.)

proboscis (i.e. the ontally in .front of

) mode of carrying r way remarkable d the descriptions act. As already a are most likely as Tsetse-flies, their wings in the resting position do not close 1 these confusion one over the other, but dix-erge slightly at the tips and are 4 USUAL HAUNTS OF TSETSE-FLIES.

also somewhat tect(form, i.e. they meet together at the base like the roof of a house (Fig. 3). The antennae, too, afford a further means of distinction. While the antennae of Tsetse-flies, as of all other , are drooping, those of Haematopota project horizontally in front of the head, and being of some length are readily seen. In those parts of Africa in which Tsetse-flies Where Tsetse-flies occur they are not found everywhere, but are are usually found. generally confined to particular tracts, which are known as “ Fly-belts,” and are often of very limited extent. We are still somewhat in the dark as to the factors that determine the limits of these GC belts,” but, although Tsetse are undoubtedly dependent upon the blood of vertebrates for their continued existence, all recent evidence goes to show that the most important element is the physical character of the locality, and, probably, its suita- bility as a breeding-ground. As a general rule it may be said that Tsetse-flies are usually met with in damp, hot localities, either on the borders of rivers or lakes, or at any rate not far from water ; this, however, applies more especially to the species belonging to the Glossina palpalis group; other Tsetse-flies, especially those of the Gloss&a morsitans group, are often found at a considerable distance from water. Cover in the shape of more or less thick bush or forest: shady trees, or reeds, is essential, and Tsetse-flies are absent from open grass plains. The mode of reproduction in Tsetse-flies-first Reproduction. discovered by Sir David Bruce, during his classical investigations in Zululand in 1895-96-is extremely remarkable, since, instead of laying eggs as do the majority of Diptera, the pregnant female produces at each birth a single full-grown larva, which, while retained within the oviduct of the mother, is nourished by the secretion of special glands, and on being born crawls away into some hiding place and immediately changes into a pupa. Observations show that, under normal conditions as regards temperature and humidity, the extrusion of larvae takes place at intervals of from about eight to eleven days. The Tsetse-fly larva is a yellowish, footless maggot, nearly as large as the abdomen of the mother, with a shagreen-like integument, and the anal extremity (twelfth segment) produced into a pair of large, black, granular prominences (the tumid lips), separated by a pit-shaped depression cont,aining the stigmata or respiratory apertures. X-FLIES. PUPAE OF TSETSE-FLIES.

?r at the base like 10, afford a further of Tsetse-flies, as I of Haematopota nd being of some

which Tsetse-flies rerywhere, but are ular tracts, which of very limited dark as to the 3e ‘( belts,” but, upon the blood 3nce, all recent rtant element is sobably, its suita- e it may be said up, hot localities, any rate not far lly to the species ther Tsetse-flies, are often found in the shape of ?es, or reeds, is Pass plains.

l!setse-flies-first Pupa of Glosr;incc pallidipes, An&n,, dorsal aspect (X 19 : a, pos,terior e$remity, showing pit and (st) right larval stigma (X 24) : b, anterior extremity, showmy longs- :uce, during his tudinal seam, which opens to permit the escirpe of the fly (x 8). G-is extremely the majority of t birth a single e oviduct of the glands, and on #nd immediately under normal ;he extrusion of i to eleven days. ;got, nearly as t shagreen-like nent) produced ihe tumid lips), E’ig. 5. alpae of Eix species of Tsetse-flies, dorsal aspect, [email protected], G. brevi@pis, New&. ; he stigmata or B G. fusca, Walk. (cephalic (lower) extremity missing) ; C, G. mmsz~a?K W&w. ; 6, G. pallidipes, Austen ; E, cf. tachinoides, We&w. ; F, G. palpalzs, Rob.-Des’. (From photographs of actual specimens.) 6 (:EOC:BAPHICAL I)ISTNl3VTIOXT.

The shape and appearance of the JM~CL * will, it is hoped, be readily understood from Figs. 4, 5, and G. Tsetse-fly pupae are dark brown or clove-brown in colour, with a slight sheen which produces somewhat the efYect of the bloom on a black plum. The tumid lips seen in t’he larva are equally conspicuous in the pupa, and the shape of the notch between them, in conjunction with the size and shape of the lips themselves, affords a valuable means of identifying and distinguishing pupae belon,+ng to different species (see Fig. 6). Althou&.0 I since the discoverv of Glos&a Geographical Distribution. tnc7~i~oides, We&w., in Southe& Arabia,,? the generalisation that existin. Tsetse-flies are con- fined to Africa no‘ionger holds good, it is nevertheless t’rue that the genus Glossi~rr at the present day occurs only in the Ethiopian Region, inasmuch as the southern portion of Arabia belongs zoo-geographically to the region in questi0n.J In the African continent Tsetse-flies have a very wide distribution in the tropical and sub-tropical zones. Roughly speaking, so far as our present knowledge goes, the northern boundary of the genus may be represented by a line drawn from the mouth of the across the middle of Lake Chad to the Nile, just south of the twelfth parallel of north lat’itude, and thence to the east coast at about 4O K. ; while its southern limit may similarly be shown by tracing a line from the mouth of the Cunene River, the southern boundary of Angola, to the north-eastern extremity of St. Lucia Lake, in Zululand. Within this area Tsetse-flies, as already explained, are not found continuously, but are restricted to “belts ” or “patches ” of forest, bush, bannnn- plantation, or eyen reeds. The accompanying map must not be regarded as affording anythin g more than an appom’mnte idea of the distribution of Glossi~. Since the necessary cover is mainly found in the valleys of rivers and watercourses, and on the shores of lakes, it is chiefly the rirer-systems that, are marked on the map. In a map on so small a scale it is, however, almost impossible to conrey any idea of the actual extent of fly-belts or fly-areas. Since it is practically certain that Glossi,ln pnlpdis

* What is seen is really thepqmCzm, or chitinous pups-case, consisting of the hardened and contracted skin of the adult larva; the true, soft- bodied 233q.m lies within this protecting envelope. Puparia of this character are invariably found in the Muscidae, and other families of Diptem belonging to the same division of the Order. t Cf. p. 41. l Cf. p. 41, note t. PUPAE OF SIX SPECIES OF TSETSE-FLIKS. 7

’ will, it is hoped, be Tsetse-fly pupae are a slight sheen which 1 a black plum. The spicuous in the pupa, in conjunction with lrds a valuable means tlonging to diff’erent

covey o f Gbossina buthern Arabia,7 the : Tsetse-flies are con- :vertheless true that nly in the Ethiopian of Arabia belongs ..z In the African ution in the tropical 3 far as our present the genus may be rth of the 8enegal 1 Nile, just sout)h of thence to the east it may similarly be the Cnnene River, h-eastern cxtremitr is area Tsetse-flies, tinuouxly, but are est, bush, banana- g map must not be 1 nppoziwnte idea necessary cover is ttercourses, and on OS that arc marked is, however, almost tent of fly-belts or j Glossiittr p+ctlis

pup&-case, consisting larva ; the true, soft- baris of this character families of Diptera, 8 GEOGRAPHICAL DISTRIBUTION. occurs, for instance, throughout the Congo-basin, the Congo and its affluents are coloured red ; but, for the reasons already stated--apart from the fact that a species of Glossina is not necessarily to be found all through the year at a spot at which it may be known to exist-it must not be imagined that G. pul- palis occurs continuously over the whole of the Congo River system. The present map should only be considered as indi- cating, very roughly, how large a portion of Africa is included within the area of distribution of the genus Glossina,, and con- sequently exposed to the contingent perils.

CHAPTER II.

THE EXTERNAL CHARACTERS OF THE GENUS GLOSSINA.

F “I. GENUS GLOSSINA.

Glossina, Wiedemann, AussereuropLische zweifliigelige Insekten, Zweiter Theil, pp. 253, 254 (1830). Nemorhina, Robineeu-Desvoidy, Essai sur les Myodaires (Memoires p&sent&s . . . & l’tlcademie Royale des Sciences de l’Institut de France . . . Sciences iUath6matiques et Physiques. Tome Deuxieme), pp. 389, 390 (1830).

Narron-bodied, elongate, dark brown, blackish, yellowish-brown, or yellowish flies belonging to the Family Muscidae, ranging in size from about 6 or 8 mm.* in the case of Glossina tachinoides, W&w., to as much. as 13 or 13.5 mm.* in that of a large female of G. brevi- palpis, Newst., or longipennis, Corti; recognisable when alive ancl at rest by the wings being closed flat one over the other above the abdomen (beyond zuhich they project considerably), instead of diva&ate (as in the case of Stomoxys) or tectiform (as in Haematopota), and by the proboscis (i.e. proboscis ensheathed in the palpij, projecting hori- zontally in front of the head ; palpi, as seen in the natural position, extending slightly beyond the proboscis, their inner sides grooved so as to form a sheath for the latter, to ml&h iu life they are applied so closely as ent&ely to conceal it; base of proboscis suddenly expanded beneath into a large onion-shaped bulb.

Head rather narrower than the thorax; eyes separate in both sexes; width of the front at the vertex in the 6 ranging from one-third to rather more than one-half of that of the eye at its widest part, and in the Q from one-half to rather more than

* Length measured from the face to the end of the abdomen, excluding the proboscis and wings. NOMENCLA 10 HEAD. one-half of the width of the eye ; vertex slightly depressed below the level of the eyes ; facial pit deep, undivided by a septum, extending to the oral margin, which is upturned and prominent ; curved suture (Bogenml~t of Brauer and von Bergenstamm) ending in a pit above the lower margin of the eye, the pit on each side being prolonged into a more or less marked, &allow depression (transuersc inzpwsGon of Hough) separating the face from the jowls ; ;X- the latter narrow in front, broad and rounded posteriorly, crossed by a line of black bristles which passes on to the occipital surface of the head, and is continued as the series of small bristles marking otT the posterior orbits; .jowls clothed with fine hair behind the line of black bristles just mentioned, in front, like the face, bare; vibrissal ridges sharp and narrow, not broadening out below as in S”tomox,~p, Hmnrcttobicc, and other genera of Xuscinae, ciliated to the level of half the length of the third .joint of the antennae, or higher : vibrissae small, fine, no elongated or conspicuous vibrissa even on the \-ibrissal angle, which is widely separated from the oral margin. Gnder side of the roof of the buccal cavity with two large, dark brown, strongly chitinised patches, very conspicuous in the majority of species when the head is viewed from below. Eyes bare, in bot’h sexes the facets in front towards the inner margin conspicuously larger than those behind. First two joints of the antennae small, the third long, nearly reaching the oral margin, concave in front, the tip being produced into a prominent forwardl+irected angle. On the inside of the third joint of the antenna near the base a minute though well-marked 1’ore can usually be distinguished, constituting the orifice of a sense-organ (see Fig. S).t Arista three-jointed, though first two joints very small, and first joint visible only when the antenna is removed and examined under the microscope ; terminal joint remarkably broad throughout its length, flat, tapering somewhat to the tip, but. far from being reduced distally to the proportions of a fine Ilair, as in the case of the arista crf Stonlozp; feathered on the upller side alone &ngitzUtwLl V, using the term jozvl for the part of the head beneath the eyes (&z&e, of III. Third ,, German authors), separated from the firm (German, TVCUUJC) by the curved IV. Fourth ,, V. Fifth ,, suture or its prolongation. In 1698 Hough (“ Proc. Acad. Nat. SC., Phila- VI. Sixth ,, delphia,” 1898, p. 1fX) proposed the terms ge~n and b~cca for Woizge and Backo respectively. t Dr. Stuhlmann, writing of G. b~c~&k~, Newst., calls this an IL auditory ” organ. If the antenna be mounted in glycerine and ex- NOTE.-The frOlEtcd1 amined under a low power, the sense-organ has the appearance of an of the front (the spr oval sac. I lye, the pit on trked, s&dlow ?TbbiS _,’ tting the face ,/’ 1 and rounded ThTrax h passes on to I as the series jowls clothed mentioned, in i narrow, not la, nnd other length of the mall, fine, no brissd angle, Iinder side of own, strongly -. ):.--- Scuielfum._-_____-- (part o/mesothoW___-__--. ty of species in both sexes uously larger ae small, t,he in front’, the rected angle. ar the bane a istinguished, 8).t Arista Id first joint mined under Iroughout its from being Fig. 7. 8 in the case Diagram showing nomenclature of external characters of Glossim, used in description. side alone VEISS *ND CELLS IN THE WINQ. Longitudimzl litim. Tmnsrerse Veins. C&S. Ia. Auxiliary vein. A. Anterior transverse vein. In, lb, lc, First, second, and third costal cells. etc.,“ 1901) in I. First longitudinal reill. U. Posterior ,, ,, II. Second ,, 1, C. Anteriorbasal ,, ,, 2. Marginal cell. yes (Backe, of III. Third D. Posterior ,, ,, ., 3. Sub-marginal cell. by tho curved :: :: 4. Discal cell. IV. Fourth 5, 6, 7, First, second, and at. SC., Phila- V. Fifth ,, ,, ,, third posterior cells. x Wmgc and VI.Sixth ,, S. Anterior basal cell. 9. Posterior ,, ,, calls this an 10. Anal cell. ?rine and BT- arance of an NOTE.-The front& stripe is the strip of coloured integument running down the centre of the &mt (the space between the eyes), from the ocelli to the base of the antennae. 12 ANTENNAE. PROBOSCI:

(including the tip) with from seventeen to twenty-nine fine, rod-like, of the sar curving, Branc7wd hairs (see Fig. S).* Proboscis projecting as tapering towards tl already described and, with the palpi, curving slightly down- minute bristles, a ro wards ; reddish-brown below, glistening yellowish-white above bristles running alo nearly to the tip, which is marked off by a slight constriction separated by sometb running along the o two somewhat longs and forwards on the tip. (For the bristl -Cephalic Bristles. Thorax quadrate to the naked eye aI with minute black number, arranged Transverse suture 1 flattened, with a pal on each side. Mark as shown in Plates caliginea, Austen), distinguishable at a: grey or brown blotc Abdomen also fla second segment wit1 grooves or impressec converging towards enclosing a more or in colour than the abdomen thinly cl appressed black ha segment become lon following segments, fine black bristle41 Fig. 8. Left ?nteUna Of, C%S$~%a Pallidipes, Austen, d, from the inner side (x 60) ; the very they are longest ant mmnute ~rOxma1 Jolnt’Df the arMa is not visit& in this view ; p. aperture of sense. . organ on third joint of antenlla. (The av&z is the process--really the distal and also that of the I extremity of the antenna-bearing the branched hairs.) of longer bristle-likl and*:exhibits beneath a tiny but conspicuous black spot; the especially conspicuo bulbous base of the proboscis is enwrapped posteriorly with a fine bristles on eacl fold of skin forming the hind wall of the buccal cavity. Palpi The male hypopygiu * The number of these hairs is usually about twenty-two or twenty- and 15, pp. 50,51), : three, but sometimes there are not more than seventeen br eighteen while * In dried specime in other cases there may be as many as twenty-eight or twenty-nin’e ; the indistinguishable. numbers on each arista may even be unequal in the same specimen. The t The modified eig last one or two hairs (at the tip of the arista) are usually bare. surface of which the L PROBOSCIS, THORAX, AND ABDOMEN. 13

-.. ____ - -__ Is the tips, clothed with short black hairs and minute bristles, a row of some eight to thirteen tiny erect black ; bristles running along the upper margin, the individual bristles ;- separated by something like regular intervals, and a similar row running along the outside on the basal two-thirds of the palpus ; ; two somewhat longer and stouter bristles * directed downwards

tip. (For the bristles of the head see below under (‘ Chaetotaxy -Cephalic Bristles.“) Thorax quadrate, flattened or only very slightly arched above, to the naked eye appearing bare, but in reality sparsely clothed with minute black hairs; the macrochaetae likewise few in number, arranged as described below under “ Chaetotaxy.” Transverse suture well marked, forming a groove. Scutellum flattened, with a pale impressed median line, and a darker patch on each side. Markings of the thorax, when completely developed, as shown in Plates I and II (Glossina palpalis, Rob.-Desv., and caliginea, Austen), but in some species very inconspicuous; if distinguishable at all from the ground colour, taking the shape of grey or brown blotches, streaks, or spots. Abdomen also flattened, tapering to the apex ; dorsum of the second segment with a pair of, more or less distinct, longitudinal grooves or impressed lines, one on each side of the middle line, converging towards the hind border of the segment and thus enclosing a more or less lanceolate area, which is sometimes paler in colour than the remainder of the segment; dorsum of the abdomen thinly clothed with minute and exceedingly short, appressed black hairs, which on the basal angles of the second segment become longer and erect, and on the hind margins of the following segments, from the third to the fifth, take the shape of fine black bristle-like hairs, especially on the apical angles, where 1(X 60); the very they are longest and stoutest ; hind margin of the sixth segment, aperture of sense- -really the distal and also that of the seventh in the female, with a complete row of longer bristle-like hairs ; seventh segment in the male with an 3ck spot ; the especially conspicuous transverse row of black bristle-like hairs or lriorly with a fine bristles on each side of the median line beyond the middle. evity. Palpi The male hypopygizm t highly characteristic in shape (see Figs. 14 iwo or twenty- and 15, pp. 50,51), nlore or less oval and tumid, its longer axis lying eighteen, while * In dried specimens these are often missing-one at least is frequently_ _ enty-nine ; the indistinguishable. Ipeoimen. The t The modified eighth abdominal segment, to the inner and concealed re. r c--1 2.1 L1~. . . ..I^ _^.^ LA-I .----a^-^” ^-^ ^‘L^^l.^2 14 LEGS BND WINGS.

in the antero-posterior direction, with a vdviform median groove course of the f (t,he anus) running from the anterior margin to beyond the middle. verse vein, at I Ventral surface of the sixth segment in the male, immediately in portion of the f front of the hypopygium, exhibiting on each side of the middle transverse vein line a chitinised plate, roughly oval in shape and bearing a patch narrow the bar of close-set, erect or recurved, minute black hairs ; * \-entral portion of the a surface of the remainder of the sixth segment and that of the the fourth vein remaining segments except the first in the male, and of all the with the ante] segments except the first in the female, membranous. what is practic Legs simple, rather long, claws longer and pulvilli somewhat obliquely dowr larger in the male than in the female. Front femora with a row exact point 0: of bristle-like hairs above and below, and (except in the case of ultimately reac (3. morsitnns t) on the posterior surface with a median row of the apex. Tk minute black spines. Middle and posterior femora with a row tion is narrow of bristle-like hairs on the basal half of the anterior surface, near vein is very cl the upper margin, and with a shorter row of similar hairs, or mation to the ( short bristles as the case may be, at t,he base beneath. Middle narrow ; the femora with a solitary bristle, usually conspicuous in the larger second, third, : species, on the upper side near the tip. Tibiae with a well-marked the tips, and tl ridge, edged with a row of closely-set minute black spines, longitudinal VI running down the outer surface, so that they, especially the though in clea front and hind pair, usually have the appearance of being to the margin somewhat flattened from side to side ; the middle tibiae hare a closing the an second row of still more minute spines in advance of that just angles of the ( mentioned, the space between, owing to the presence of t,he curved, bent ridge, having the appearance of being excavated. Tibiae also tudinal. Wir with a row of short fine hairs or bristle-like hairs running down light drab to s the outer side, and sometimes with one or two short bristles close to the apex on that side, otherwise, with the exception of the usual apical spines, entirely devoid of isolated bristles or bristle- like hairs. On the three basal joints of the front and middle Cephalic E tarsi three rows of minute black spines similar to those on the (the largest a tibiae ; on the basal joints of the hind tarsi they are less distinct,, occipital regior but apparently two rows are present. (Proc. Acad. 1 Wings with an absolutely unique venation (see Fig. ‘i), which, greater ocellar apart from all other characters, will at once serve to distinguish post-vertical pa a specimen of the genus. The most striking peculiarity is the the lesser ace.

* These plates, the function of which obviously is to assist in holding * Cf. Garry d the female during copulation, may be termed the kctors (Greek IKTW~, “ Biological Bul: holding fast) ; they sometimes furnish useful characters for the distinction fine grooves in 1 of species. ’ - manner towards denotes a highs t In the o&se of G. Zongipennis, too, the POW appears to be broken up into a double line and is consequently barely distinguishable. unrilled wing.” / CHAETOTAXY. 15

nzedinn pow course of the fourth longitudinal vein. The anterior basal trans- md the middle. verse vein, at the base of the discal cell, is very short, and the mmediately in portion of the fourth longitudinal vein before the anterior (small) of the middle transverse vein is bent downwards in such a way as greatly to Baring a patch narrow the basal half of the discal cell, the width of the distal irs ; * mmtra1 portion of the anterior basal cell being correspondingly increased ; d that of the the fourth vein bends abruptly upwards to the point of contact tnd of all the with the anterior transverse vein, where it forms with itself IS. what is practically a right angle ; from this point it again runs 4li somewhat obliquely downwards, and is once more bent upwards at the ra with a row exact point of contact with the posterior transverse vein, in the case of ultimately reaching the margin of the wing some distance before edian row of the apex. The first posterior cell is open, but its distal por- a with a row tion is narrow and much drawn out. The third longitudinal surface, near vein is very close to the second, and shows a marked approxi- lilar hairs, or mation to the Costa, the sub-marginal cell being correspondingly ath. Niddle narrow ; the third costal cell is considerably elongated ; the in the larger second, third, and fourth longitudinal veins all turn upwards at , well-marked the tips, and the anterior transverse vein is very oblique; the sixth black spines, . longitudinal vein becomes obsolete soon after the anal cell, specially the though in clear, unchitinised form it can still be traced nearly Lee of being to the margin of the wing. Posterior basal tranverse vein, tibiae hare a closing the anal cell, angulate in the middle, so that the distal of that just angles of the cell are acute. Posterior transverse vein slightly k, gence of the curved, bent up somewhat abruptly to meet the fourth longi- Tibiae also 8: tudinal. Wings varying in colour according to the species from Inning down PL light drab to sepia-brown ; membrane of the wing rilled.* bristles close ption of the CWAETOTAXY OF GLOSSINA. es or bristle- and middle Cephalic Bristles.-One pair of vertical bristles of large size those on the (the largest and most conspicuous bristles on the head) ; on the less distinct. occipital region no trace of the pair of bristles termed by Hough (Proc. Acad. Nat. SC. Philad. 1698, p. 166) the oeeipito-central; ;. 7), which, greater ocehr bristles (ocellar pair of Osten Sacken) small ; distinguish post-vertical pair not differentiated in size from the remainder of iarity is the the lesser ocellar bristles; frontal bristles largest below (trans-

3t in holding * Cf. Gamy de N. Hough, M.D., “ Some Muscinae of North America,” :Greek ~K~WP, “ Biological Bulletin,” Vol. I (1899), p. 20, note 1 :-“ These rills are very 1e distinction fine grooves in the surface of the wing, which run in a sort of radiate manner towards the border. They are very numerous. A rilled wing denotes a higher stage of development, a more recent form, than an )e broken up -.....:,,.a ?A-.,. ), CHAETOTAXY. frontal group of Hough, lot. cit.) ; orbital bristles absent in roughly i both sexes ; vibrissal angle (separated by a well-marked and Generally rather wide interval from the oral margin) without a large and the notopl conspicuous vibrissa ; vibrissal ridges ciliated to a point on a level row form wit’h or rather above the middle of the third joint of the marginal antenna. Pleura Thwack Bristles.--The general arrangement, as can be seen Becker, I from the accompanying diagrams, is, on each side, as follows :- Dorsal aspect (Fig. 9).-&nreraZ, from 1 to 3 (sometimes apparently 4), the lowest bristle the largest. Post-lmmeral (intra- humeral of O&n Sacken) absent,. Notopleural (of Girschner Humera/ &&/es -’

Nofopfeumf BriWes

Anterior .sm

vertical r( Fig. 9. bristles in Diagram of thorncic chaetotaxy of Glossina, dorsal aspect. (X 10.) with small and Hough, post-humeral of O&en Sacken), 2. Praesutwal, l.* and 2 post Supradar, 1. In&a-alar, 1, situated just in front of the post- the middle alar calhrs, on a level with the second dorso-central. Post-alar, 3, * Pterop the foremost somewhat smaller than the other two, and placed Haemutobia directly &ove the alar frenum. Dorso-central, 3, two near together, close to the hind margin of the dorsum, and one immediately in front of the suture ; the latter bristle may be termed the praesutural dorso-central. Inner dorso-central, 1, near the hind margin of the thorax. Scutellar, 2, one marginal (near the basal angle), the other apical; additional scutellar bristles are occasionally found. The relative size of the various bristles enumerated above is

* Often small and difficult to distinguish in G. palpalis. PLEURAL CHAETOTAXY. 17

absent in roughly indicated in the diagrams by the size of the dots. arked and Generally speaking the largest bristles are the lowest humeral, large and the notopleural, supra-alar, post-alar, the bristles of the transverse on a level row formed by the first and inner dorso-centrals, and the nt of the marginal and apical scutellar bristles. Pleural aspect (Fig. lo).-Prothoracic, 1. Stigma& (cf. m be seen Becker, Berl. Ent. Z. xxxix (1894), p. 83), 1. Mesopleural, a llows :-

sometimes Pfzsutural dorso-central Bristle,X Sq0ra-a far Bristle 2”ddo

Fig. 10. Diagram of thoracic chaetotaxy of Glossina, pleural aspect. (x 10.)

vertical row of about 6 along the posterior edge, with smaller bristles in between. Pteropleurat,* a vertical row of usually 3, 0.) with smaller bristles on either side. Sternopleural, 3,-l anterior, tural, 1.” and 2 posterior : below, a conspicuous row of bristles in front of the post- the middle coxae. Hypopleural, none. &alar, 3, td placed * Pteropleursl bristles seem to be entirely wanting in Stomoxys, Eatmatobia, and other genera of the Muscinae. iwo near and one 3 may be Z, 1, near oal (near r bristles

above is I. C2 CHAPTER III.

THE FOUR GROUPS INTO WHICH TSETSE-FLIES ARE DIVIDED.-TABLES FOR THE DETERMINATION OF GROUPS AND SPECIES.

IN his recent paper on the male genital appendages*-one of the most valuable contributions to our knowledge of the Tsetse- flies yet published-Mr. Newstead has divided the genus Glossina into three groups. In the present work this arrangement is adopted in a somewhat modified form, since Glossina brevipalpis, Newst., and its immediate allies are removed from the GZ&sina fusca group, in which they are placed by Mr. Newstead, and formed into a group by themselves. It is believed that this arrangement represents a natural grouping, which will aid the student in the identification of species. The following Tables are based solely upon external characters, in the ordinary sense of the term; though of course subordinate in value to those derived from the male genitalia, external characters have been found by experience to be sufficient for the determination of the species recognised in this volume, while they possess the additional advantage of being present in both sexes. The Tables here given must not be regarded as anything more than a series of “short cuts” to the determination of species; the characters mentioned are simply those that are most convenient and easily recognisable for this purpose, and it must on no account be imagined that these represent the only differences between species. The detailed descriptions in the following chapters are in all cases comparative, and at the end of the account of each species the differences between it and its nearest allies are specially emphasised under the heading “ Affinities and

* “ A Revision of the Tsetse-Flies (Glossina), based on a Study of the Male Genital Armature,” by Robert Newstead, XSc., A.L.S., etc. (Bulletin of Entomological Research, Vol. II, Part 1, pp. 9-35, Figs. l-17 (April, 1911).) DETERMINATION OF GROUPS. 19

Distinctive Characters.” An epitome of the diagnostic characters of each species will be found printed in italics at the commence- ment of the detailed description, which should always be consulted in cases of doubt. The dichotomic system is employed in the subjoined Tables, which it is hoped will prove readily intelligible after a little practice. In using the Tables, the Group to which a specimen or species belongs should of course be determined first.

TABLE FOR THE DETERMINATION OF GROUPS.

1. Hind tarsi entirely dark brown, black- ish-brown, or black above (or at least all joints more or less dark; in cf. tachinoides, Westw., the extreme tips of the first three joints and the ex- treme bases of the second and third joints are buff, and a larger or smaller area at base of first joint is also paler than remainder of joint, especially in 0 ) ...... I. Glossina palpalis Group. Hind tarsi not entirely dark brown, blackish-brown, or black above, but either last two joints alone so coloured, and thus forming a conspicuous contrast with remaining joints, which are entirely pale, or last two joints, viewed from above, con- spicuously darker than first two or first three joints ...... 2 2. Upper surface of abdomen distinctly banded, i.e. ground colour pale (drab- grey, buff, or ochraceous-buff), marked with very conspicuous, dark brown or clove-brown, transverse bands, inter. rupted in the median line * ...... *... II. Glossina morsitans Group.

* N.B.-The abdomen of G. tachinoidcs, Westw., is banded in this way, but the coloration of the hind tarsi places this species in Group I. Note that in individuals belonging to Group II. (Glossina morsilalzs Group) the inner extremities of the interrupted abdominal bands are often partially obliterated; in such oases, however, the outline of the obliterated portions is usually traceable. DETERMINATION OF SPECIES.

Upper surface of abdomen not so banded (i.e. abdominal markings not con- sisting of interrupted dark bands on a pale ground) ...... 3 1. La 3. Wings fairly dark (dull sepia-coloured), palpi (except in cf. tabaniformis, La We&w.) long and slender ...... III. Glossina fusca Group. Wings pale (pale is&bells-coloured oc light drab), palpi short ...... IV. Glossina brevipalpis Group.

TABLES FOR THE DETERMINATION OF SPECIES.

I. 2. Tl GLOSSINA PALPALIS GROUP.* 1. Docsum of abdomen ochcaceous-buff oc buff; third and following segments exhibitingsharply deEned, dark brown oc clove-brown, interrupted, tcans- verse bands ...... , ...... tad&wicks, Westw. Docsum of abdomen not so marked...... 2 2. Third joint of antennae pale (cceam- buff to ochraceous-buff), clothed with long and fine hair, forming a oon- spicuous fringe on front and hind margins (cp. Fig. 13, p. 36) ...... pallicera, Bigot. Third joint of antennae entirely dark (mouse-grey), except at extreme base on outer side, and without 5 con- spicuous fringe of long and fine hair 3 1. ? 3. Dorsal surface of abdomen dark sepia- brown ; median paler area on second segment broad, and more or less quadcate oc irregular in outline; hypopygiumt of d buff oc ochcaceous- buff ...... ,...... caligimea, Austen. Dorsal surface of abdomen blackish- brown ; median paler area cuneate (i.e. triangular in outline) ; hypopy- gium of d grey , ...... palpalis, Rob.-Desv.

* Gzossi?za fuscipes, Newst. (see p. 29), which belongs to this Group, is not inoluded in the above Table, since the author has not yet had the opportunity of studying this species. t See p. 13, note t. * The a. submov at any cf race of a DETEiMINATION OF SPECIES. 21

II.

GLOSSINA MORSITANS GROUP.

1. La s t two joints of front and middle tacsi with sharply defined, clove-brown or black tips.. . 2 Last two joints of front and middle tacsi with- out sharply defined, clove-brown or black tips (front and middle tacsi either entirely pale or, at most, last two joints of front tacsi faintly brownish at the tips, and last joint and distal half of penultimate joint of middle tacsi light brown-never so dark as to form a sharp contrast with the remaining ’ joints) ...... pall&pees, Austen. 2. Third joint of antennae with a distinct fringe of fine hair on front margin; dark brown or clove-brown bands on abdominal seg- ments extending close to hind margins (i.e. pale ground colour, apart from the median interspace, confined to a very narrow hind border)...... hgipalpis, Wied. Third joint of antennae without a distinct fringe of fine hair on front margin; dark brown or clove-brown bands on abdominal segments not extending close to hind macgms ...... wmsitans, JYestw.*

III.

GLOSSINA FUSCA GROW. 1. Third joint of antennae fringed with fine hair on anterior and posterior margins ; fringe on anterior margin conspicuous under a hand-lens magnifying 15 dia- meters (nominal), when head is viewed in profile ...... 2 Third joint of antennae with fringe of fine hair on anterior margin so short 5s to be scarcely noticeable under 5 hand-lens magnifying 15 diameters (nominal), when head is viewed in profile (longest hairs in fringe in length not exceeding one-sixth of width of third joint) ; palpi long and slender ...... 3

* The designation Glossina morsitalzs, as used in this book, includes 6’. submorsitalzs, Newst. As explained on pp. 51-52, the author, provisionally at any rate, is inclined to regard G. submorsitans, Newst., as a form oc race of cf. morsitans, Westw., rather than as a distinct species. 22 D.ETERMINATION OF SPECIES.

2. Longest hairs in fringe on front margin of third joint of antennae, in length equal to from one-fourth to one-third (not exceeding one-third) of width of third joint ; palpi of moderate length ...... tabanijorolmis, Westw. Longest hairs in fringe on front margin of third joint of antennae in length equal to from one-half to three-fourths of width of third joint ; palpi noticeably long and slender , ...... nigrojusca, Newst. 3. Pleurae drab-grey or isabella-coloured, hind coxae buff or greyish-buff...... _ . . jusca, Walk. Pleurae dark grey, hind coxae mouse-grey . jicscipleuris, sp. nov. ‘1

IV. 1. Dar; GLOSSIXA BREVIPALPIS GROUP. br 1. Dorsum of thorax with four sharply defined, ex dark brown, more or less oval or elongate 01 spots, arranged in a parallelogram, two in VC front of and two behind the transverse Dor suture; proboscis bulb with a sharply 2 Thil defined brown or dark brown tip ...... , longipennis, Corti. bl Dorsum of thorax without such spots ; pro- 10 boscis bulb not brown or dark brown at fr the tip ...... 2 (c 2. Wings with upper, thickened portion of Thi anterior transverse vein much darker in (1 colour than adjacent veins, and thus stand- 01 ing out conspicuously against the rest of “I the wing ., . . . ., ...... bmvipalpis, New& 3. DOI Wings with upper, thickened portion of b anterior transverse vein not much darker SI in colour than adjacent veins, and thus 9 not standing out conspicuously against II the rest of the wings (wings practically E unicolorous).. .,. ,...... , ...... 9nedicolu9n, sp. nov. D o t ( I

* Glossi not include opportunity t Seep. 23

CHAPTER IV.

THE GLOSSINA PALPALIS GRO.UP.

SYNOPSIS OF SPECIES.*

1. Dorsum of abdomen ochraceous-buff or buff; third and following segments exhibiting sharply defined, dark brown or clove-brown, interrupted, trans- verse bands ...... tachimides, Westw. Dorsum of abdomen not so marked . . . 2 2. Third joint of antennae pale (cream- buff to ochraceous-buff), clothed with long fine hair, forming a conspicuous fringe on front and hind margins (cp. Fig. 13, p. 36) ...... pallicera, Bigot. Third joint of antennae entirely dark (mouse-grey), except at extreme base on outer side, and without a con- spicuous fringe of long fine hair . . . . . 3 3. Dorsal surface of abdomen dark sepia- brown ; median paler area on second segment broad and more or less quadrate or irregular in outline ; hypopygium t of 6 buff or ochraceous- buff ...... ,.. caliginea, Austen. Dorsal surface of abdomen blackish- brown ; median paler area cuneate (i.e. triangular in outline) ; hypo- pygium of 6 grey ...... pulpalis, Rob.-Desv.

* Gloss&a juscipes Newst. (see p. 29), which belongs to this Group, is not included in the above Table, since the author has not yet had the opportunity of studying this species. t See p. 13, note t. 24 DESCRIPTION OF GLOSSINA PALPALIS. GLOSS

tarsi blad Glossina palpalis, Rob.-Desv. with sepia. (Plate I.) (G. caligin Nemorhina palpalis, Robineau-Desvoidy, Essai sur les Myodaires (Memoires Head.- Present& par Divers Savans it I’Academie Royale des Sciences de 1’Institut de France. Sciences Mathematiques et Physiques. Tome surface of Deuxieme), p. 390 (1830). from ochri Glossina longipalpis, Walker (net Wiedemann), Entomologist, VI, p. 328: or parafrc (1873). elongate a Glossina ventricosa, Bigot, Ann. Sot. Ent. France, 6’ Serie, T. 5, pp. 122, dark-brow 123 (1885). defined dt Glossina longipalpis, Bigot (net Wiedemann), ibid. p. 122. conspicuoi Glossina tabanifomis, Bigot (net Westwood), ibid. p. 123. alztennae Glossina longipalpis, Austen (net Wiedemsnn), Report of the Proceedings narrowly of the Expedition for the Study of the Causes of Malaria, etc., p. 18 mouse-gre; (1899). blackish o Glossina palpalis, Austen, “A Monograph of the Tsetse-Flies,” p. 71,. Thora: Plate I (1903); Newstead, Dutton and Todd, Annals of Tropical bluish-gre; Medicine and Parasitology, Series T. M., Vol. I, No. 1, pp. 57-71, in Plate 1 Plate III, figs. 7-9 (February 1, 1907). narrow St: Glossina maculata, Newstead, Annals of Tropical Medicine and Parasit- reaching ology, Series T. M., Vol. I, No. 1, p. 73, Plate III, figs. 5, 6 (February 1, 1907). hind mar; expanded 6, ? .-Length,* 8 (10 specimens, from different localities} immediate S to 9 mm., ? (10 specimens, from different localities) 8.6 to pair of n 10.2 mm.; width of head, 8 2.4 to 2.6 mm., ? 2.5 to just fluent wi under 3 mm.; width of front at vertex, 8 0.6 mm., $’ 1 mm. ; stripes on length of wing, 6 7 to 8.4 mm., ? S.4 to 9.2 mm. defined 03 more or 1 Abdomen clove-brown t or blackish-brown ; thora& usually paler, but in fro ,with dark brown markings on a greyish ground, as shown in and then Plate I ; abdomen generally with at least aA indication of a pale dorsum nc or slate-grey longitudi’nal median stripe, with pale lateral triangular a broad il markings, and usually the hiacl margins of the segments narrowly the latere pale. Femora in typical race more or less mouse-grey, greyish- inwards brown, or dark slate-coloured; tibiae, extreme tips of femora, and Humeral joints *first three qf *front and middle tarsi buff or ochraceous-buj; the curve hind (or middle and hind) tibiae sometimes infuscated in & ; hind the post-a * The length given is in the case of all species exclusive.,of the proboscis drab-grey and palpi, and’is measured from the epistoma (front margm of the lower part of the face) to the tip of the abdomen. in the ce t In the names employed for colours throughout this book, the author olive-grey follows Ridgway’s “ Nomenclature of Colors for Naturalists ” (Boston : of the m Little, Brown, and Company, 1886). x3. GLOSSINA ‘PALPALIS: HEAD AND THORAX. 25

tarsi blackish-brown or clove-brown above ; wings strongly tinged ywuo 0” u,w,n, (G. caliginea, Austen-Plate II).

I (MBmoires iciences de Head.-Face and jowls cream-coloured or cream-buff; posterior UN. Tome surface of head (occiput) entirely cinereous ; frontal stripe varying from ochraceous to dark chestnut ; frontal margins (sides of front VI, p. 328 or parafrontals) greyish, seen from the side with a dark brown I elongate area below; ocellar triangle smoke-grey, enclosing the 5, PP. 122, i dark-brown ocellar spot, which is joined posteriorly to a sharply ; deiined dark brown band, uniting the vertical bristles and very conspicuous except in the darkest specimens ; second joint of antennae more or less yellow at the apex in front, third joint roceedings narrowly buff at extreme base on outer side, otherwise entirely etc., p. 18 mouse-grey ; arista buff, dark brown beneath ; palpi mouse-grey, blackish on upper side ; proboscis bulb dark brown. :s,” p. 71, Thoraz.-Dorsum in the most clearly marked specimens f Tropical bluish-grey or greyish-olivaceous, with brown markings as shown pp. 57-71, in Plate I. These markings when fully visible are as follows : a d Parasit- narrow stripe on each side of the median line, interrupted before ebruary 1, reaching the transverse suture and again before reaching the hind margin ; the section of each stripe behind the suture is expanded posteriorly, and the terminal portion of the stripes ocalities) immediately in front of the hind margin takes the shape of a t) 8.6 to pair of more or less confluent ill-defined spots, sometimes con- i to just fluent with the stripes in front ; next to the two admedian 1 mm.; stripes on each side on the suture itself a more or less sharply defined oval spot ; on the outside of this a longitudinal stripe, more or less interrupted and sometimes obsolete in the middle, Ily paler, but in front curving round outwards behind the humeral callus rhown in and then running backwards along the lateral margin of the f a pule dorsum nearly to the post-alar callus ; in the area thus enclosed *iangular narrowly a broad ill-defined patch in front of and behind the suture, while greyisla- the lateral stripe itself sends off two prolongations, which run iora, and inwards for a certain distance on each side of the suture. ?OUS-bll& Humeral callus with a spot on its upper portion, confluent with 8; hind the curved stripe behind it; a more or less ill-defined spot on the post-alar callus also. Pleurae olive-grey or smoke-grey in d, proboscis drab-grey or olive-grey in $’ , a more or less distinct brown patch ;he lower in the centre of the mesopleura. Scutellum buff or cream-buff, 18 author olive-grey at the base, the usual dark brown patch on each side (Boston : of the median line more or less conspicuous ; apical scutellar 26 GLOSSINA PALPALIS VAR. WELLMANI. DIS bristles in the 0 variable in length,-sometimes (as in specimens humeral ca from the Congo Free State and Uganda) short, sometimes (as in especially specimens from Liberia, Sierra Leone, and the Gambia) nearly as slightly frl long as in the 6. The tl Bbdonzen.-Dorsum clove-brown or blackish-brown ; first seg- River, An ment and a median triangular area on the second (its base Creighton resting on the front and its apex on the hind margin of the in the I second segment) buff-coloured or cinereous, the pale triangle Benguella continued backwards as a narrow, more or less well-defined about eig. median stripe, usually reaching at least as far as the hind the banks margin of the fifth segment ; lateral margins of the segments the heighi from the second onwards grey, expanded on the apical angles Beside into triangular markings ; extreme hind margins of the segments rently al from the second to the sixth usually narrowly pale or grey ; Katanga seventh segment, as also the hypopygium in the 6, entirely Islands ir grey. that the Legs.-Last joint of front and middle tarsi dark brown, often specimem more or less buff at base, sometimes distal third alone dark Coast, an brown, remainder buff; penultimate joint of front and middle to be into tarsi dark brown, more or less buff at base. and G. pa Wings as described in diagnosis printed in italics above. Squnmae waxen-white, border of the antisquama darker, fringed with short, darker hairs. Halteres cream-buff.* The e includes Senegal Gloss& palpalis var. ,wellmani, Austen. welllnani Glossina palpalis wellmani, Austen, Annals and Dfagazine of Natural the soutl History, Ser. 7, Vol. XV, p. 390 (April, 1905). boundar: Glossina bocagei, Franqa, Jornal de Sciencias Mathematicas, Physicas e includes Naturaes, 2” Ser., T. VII, No. xxvii, p. 134 (June, 1905). Protect0 8, $’ .-Distinguishable from typical G. palpalis, Rob.-Desv., German by a peculiar reduction in the markings qf the dorsum of the end of t thorax. mately f and the Frontal stripe pale ochraceous; thoracic markings much district reduced, so that the thorax in a well-preserved specimen appears not to 01 spotted, the antero-lateral markings taking the form of spots or blotches ; the spot immediately behind the inner extremity of the

* For au account of the internal anatomy of Glossina palpalis, see owi Minchin, Proceedings of the Royal Society, Vol. 76n, pp. 531-547 (1905) : found a. for the structure of the proboscis, see Stephens and Newstead, Liverpool knowlec School of Tropical Medicine, Memoir XVIII, pp. 53-74, Six Plates (1906). NI. DISTRIRtiTION OF GLOSSINA PALPALIS. 27 specimens oes (as in nearly as -The types of this variety-which are from the Katumbela first seg- River, Angola, and were collected in November, 1904, by Dr. F. (its base Creighton Wellman, in whose honour the form was named-are n of the in the British Museum (Natural History). Writing triangle Benguella on February 5th, 1905, Dr. Wellman said :-“ I took :ll-defined about eighty specimens of these Tsetse-flies in four days along the hind the banks of the Katumbela River, two days from the coast, in segments the height of the rainy season.” al angles Besides occurring in Angola the variety wellmani is appa- segments rently also found in other localities, as in the Gambia, the or grey; Katanga District of the Congo Free State, in the Matondwi entirely Islands in Lake Tanganyika, and elsewhere. It may be added that the National Collection contains a considerable number of wn, often specimens from various localities, including the Gambia, Gold me dark Coast, and Northern and Southern Nigeria, which would appear 1 middle to be intermediate between typical Glossina pulpalis, Rob.-Desv., and G. pnlpalis var. welbmani, Austen. :s above. *, fringed DISTRIBUTION OF G. palpalis, Rob.-Desv. The area occupied by Glossina palpalis, as at present known, includes the whole of West Africa, from the mouth of the Senegal River (about 16” N.) to Angola (where the variety wellmani apparently predominates), and extends eastward into f Natural the southern Bahr-el-Ghazal. Proceeding southward, the eastern boundary of the species follows the valley of the Xle, and ‘hysicas e includes the eastern shores of Lakes Victoria (in the East Africa Protectorate and German East Africa) and Tanganyika (in b.-Desv., German East Africa), and their affluents ; from the southern ‘l of the end of the latter lake the boundary trends south-west, approxi- mately following the frontier between North-Eastern Rhodesia and the Congo Free State, and passing through the Katanga ;s much district of the latter country into Angola. G. palpalis appears appears not to occur on Lake Nyasa. spots or ‘y of the

Ipalis, SEC L7 (1905) : Liverpool IS (1906). knowledge of the bionomics of Glossina pulpalis, concerning 28 G: PALPALIS: SYNONYMY, AFFINJTIES, ETC. DES

which a mass of literature is already in existence, and is Prom G. pt increasing week by week. The latest information on the subject apart from will be found in the monthly “ Bulletins ” of the Sleeping the dusky ( Sickness Bureau.” its antenna there is no

SYNONYMY, AFFINITIES, AND DISTINCTIVE CHARACTERS. abdominal

As stated in the author’s “Monograph” (p. 78), Glossina ventricosa, Bigot, was described from two female examples of G. palpalis, Rob.-Desv., in which the abdomen is enormously distended with coagulated blood. Thanks to the generosity of Glossina fus Mr. G. H. Verrall, these specimens are now in the National ology, SG Collection. No spec Some few years ago, by the courtesy of Mr:Newstead, the in the posse author was enabled to examine the type of Glossina maculata, have yet ri Newst. (a 0 ), which proved, in his opinion at any rate, to be a following, specimen of G. palpalis, Rob.-Desv., curiously spotted by some description. foreign matter. Mr. Newstead is unfortunately unable to accept “ This t this conclusion, and it is therefore with especial reluctance that Rob.-Desv., the author of this volume is compelled to state his conviction, or dusky le and to regard Gloss&a maculata, Newst., as a synonym of G. tachinoic G. palpalis, Rob.-Desv.7 and altoget In external characters, at least, Glossina palpalis, Rob.-Desv., with infuse is more nearly related to G. fuscipes, Newst. (p. 29) and resembles t G. caliginea, Austen (p. 30, Plate II), than to any other species form ; but of the Glossina palpatis group yet described. According to claspers ar Newstead’s description of the former species, G. palpalis is insects. distinguished from by its much larger size, and by G. fuscipes ‘I Len@ the thorax not being “ dusky grey,” or the legs “uniformly “ MAL, infuscated or dusky.” From G. caliginea, Austen, G. palpalis is dusky whit distinguishable, by the head being distinctly narrower, inter alia, spot. Ante? t.he arista shorter and stouter, the abdomen blacker, the median brown, wit1 paler area on the second abdominal segment cuneate in outline castaneous. instead of more or less quadrate, and by the wings being paler. u Thora * A most useful summary (by Dr. A. G. Bagshawe, Director, Sleeping from the fr Sickness Bureau) of what was known as to the bionomios of C. pulpalis at the commencement of 1909 is to be found in the Sleeping Sickness Bureau straight an Bulletin, Vol. I, No. 3, pp. 89-106 (January, 1909). This has since been greyish-blat reprinted, and is to be obtained separately from the Sleeping Siokness Bureau, Royal Society, Burlington House, London, W. nants of th + It is perhsps permissible to mention that Mr. C. 0. Waterhouse, grey, with : Vice-President of the Entomological Society of London (late Assistant- Keeper, Insect Section, British Museum (Natural History) ), who made an of the late] entirely independent examination of the type of G. maculate, was emphati- pleurae dar cally of the opinion that the spotted appearance of the speclmeo was “ not cL Abdon natural.” :ES, ETC. DESCPYIPTION OF GLOSSINA FUSCIPES. 29 we, and is From G. pa&era, Bigot (p. 35, Plate *III), the present species, the subject apart from being much darker, may at once be distinguished by le Sleeping the dusky colour of and absence of long hair on the third joint of its antennae. With G. tachinoides, Westw. (p. 39, Plate IV), there is no possibility of confusion owing to the characteristic

ACTERS. abdominal markings of the former. B), Gloss&a xamples of enormously Glossina fuscipes, Newst. :nerosity of *Glossina fuscipes, Newstead, Annals of Tropical Medicine and Parasit- e National ology, Series T. M., Vol. IV, No. 3, p. 375 (December 20, 1910). No specimens of this species, of which the type is at present rstead, the in the possession of the Liverpool School o$ Tropical Medicine, : macthlata, have yet reached the British Museum (Natural History) ; the te, to be a following, however, is a verbatim transcript of the original d by some description. e to accept “ This tsetse-fly may be readily distinguished from Gl. palpalis, tance that Rob.-Desv., by its much smaller size, by the uniformly infuscated :onviction, or dusky legs, and the dusky-grey thorax. In size it resembles ‘nonym of G. tachinoides, West., but it is a relatively stouter built insect ; and altogether it is most like a dwarfed specimen of GI. pnlpnlis ,ob.-Desv., with infuscated legs and a dusky thorax. The genital armature 29) and resembles those of Cl. palpalis and Gl. tachinoides in its general er species form ; but the shape of the superior claspers and the inferior ording to claspers are markedly distinct from those of either of these alpalis is insects. e, and by “ Length.-74 mm. ; length of wing, 8 mm. uniformly “ MALE.--Elena : frontal stripe yellowish brown, margins jalpalis is dusky white ; ocellar triangle buff, enclosing a very dark brown narrower, spot. Antennae grey ; arista dark brown ventrally. Palpi smoky- Le median brown, with a paler median line. Proboscis with the bulb dark in outline castaneous. ng paler. ‘( Thorax with a median dark grey rectangular area extending r, Sleeping from the front to the scutellum, the sides of which are perfectly mzpazis at am Bureau straight and parallel, on either side of this grey area are two lime been greyish-black and somewhat triangular blotches, evidently rem- r Sickness nants of those found in other species of Glossina ; scutellum dark aterhouse, grey, with a very faint pale grey median line, margins buff; sides Assistsnt- 1 made an of the lateral thoracic depressions dark brown or almost black ; emphaei- pleurae dark grey. was “ not ((Abdomen resembling that of Gl. palpalis, but the bi-lateral 30 DESCRIPTION OF GLOSSINA CALIGINEA. ANTENNA banding faintly pronounced ; lateral margins with pale angula#r areas. ‘6 Legs strongly and almost uniformly infuscated (dark grey) ; femora with the basal and ventral portions ochreous-buff ; hind tarsi all dark brown or almost black. ‘6 Wings.-Anterior transverse vein strongly and suddenly incrassate at its junction with the third longitudinal vein. “ HABITAT.-The only example which we possess of this interesting tsetse-fly was taken by Dr. Shircore* at Nimule, Nile Province, Uganda. Fortunately it proved to be a male, otherwise it may have passed as a small dusky form of Blossina palpalis.”

Glossina caliginea, Austen. (Plate II.) Glossina caliginea, Austen, Bulletin of Entomological Research, Vol. I, Part 4, pp. 294-297, figs. l-3 (January, 1911).

6 , 0 .-Length, 6 (15 specimens) 8 to 8.5 mm. ,t ? (15 specimens) 9 to 10 mm. ; width of head 2.8 to 3 mm. ; width of front at vertex, 8 0.75 mm., 9 1 mm. ; length of wing, 8 8 to 8.75 mm., 0 9.4 to 10 mm.

Allied to and resembling Glossina palpalis, Rob.-Desv., but browner, and usually somewhat larger; head distinctly broader, front if anything slightly narrower, at least in 8 ; dark portion of dorsal surface of abdomen browner (dark sepia-brown, instead of clove-brown or blackish brown), paler area on second segment, instead of being con$ned to a, usually narrow, median triangle, broad and more or less quadrate or irregular in outline, its lateral margins being generally ill-dejned; hypopygium of 6 buff or ochraceous- buff, instead of grey as in typical G. palpalis; pollinose dorsum of seventh abdominal segment in both sexes often cream-b& and consequently yellower than in G. palpalis; extreme hind wargins of preceding segments (except posterior angles) never ligkter.

Head.-Face, jowls, and occiput coloured as in G. pulp&is, but sides of face (parafacials) slightly narrower, and jowls not

* (‘ The data here given are taken from Dr. Shircore’s letter; the actual light mouse: locality is not stated by him--R. N.” t A specimen in the Museum collection with its abdomen distended by coagulated blood measures 9.4 mm. in length. :INEA. ANTENNAE IN G. CALIGINEA AND G. I’ALPALIS. 31

pale anguhx descending so much posteriorly (i.e. lower margin of eye nearer to lower margin of head on each side) ; frontal stripe chestnut ; (dark grey) ; us-buff ; hind

md suddenly I vein. XSHSS of this ’ at Nimule, ) be a male, n of Glossina

Isearch, Vol. I,

nm.,t 0 (15 Fig. 11. mm. ; width Antenna of Glossina caligima, Austeu. (x 3O.P ;h of wing,

‘6:Dew., but :tly broader, 4 portion of I, instead of ment, invtead e, broad and :ral margins ’ ochraceous- 8e dorsnna of ln-buf, and I margins of r. Fig. I?. Antenna of Glossinn palpalis, Rob:Dew. (x 30.) oalpalis, but jowls not frontal margins (sides of front or parafrontals) varying from V; the actual light mouse-grey to olive, often darker than in G. pulpalis, distended by * The author desires to acknowledge his indebtedness to the Entomo- logical Research Committee (Tropical Africa) for the loan of the blocks for this and the two following figures. D 3 2 G. CALIGINEA: THORAX AND ABDOMEN. DISTR: showing, as in the latter species, a dark brown elongate area shape, but’sc ‘below when viewed from the side ; ocellar triangle drab-grey or from the ven mouse-grey, ocellar spot and dark brown band uniting vertical not quite so bristles as in G. pulp&s ; antennae mouse-grey, narrowly cream- Legs ochr buff at apex of second joint and extreme base of third joint on blotched wit1 outer side, anterior and posterior edges of third joint in both or streaked 7 sexes without a long and conspicuous fringe of tine pale hair * almost entirt (cf. Figs. 11 and 13), and its distal extremity only moderately middle tarsi t prominent, a&ta distinctly longer and more slender (tapering off brown above less abruptly) than in G. palpalis (cp. Figs. 11 and IS) ; palpi Wing8 UI mouse-grey, darker above and towards distal extremity. cream-colours !Z%oraz.-Dorsum with dark brown markings of same type Halteres 4 as in typical form of G. palpalis, but often somewhat more The typis extensive ; pleurae and interspaces between markings on dorsum Southern Ni light mouse-grey ; scutellzcm, at least in Q , somewhat more elon- both taken 1 gate than in G. pulpalis, apical scutellar bristles in Q unusually the 8 on a ( short, reduced to small spines, hind margin of scutellum and on a creek be interspace between distal extremities of dark blotches on dorsum cream-buff. AbdorrLen.-First segment cream-buff or buf?‘, with a dusky (dark grey) patch on each side; dorsum of second to sixth Up to the from Southe segments inclusive dark sepia-brown ; median paler region on dorsum of second segment cream-buff to ochraceous-buff or pale and Q , the f8 cinnamon, the central lanceolate area enclosed within the usual collection : -: admedian grooves exhibited by this segment often paler than the 1910, and 1 ( region outside the grooves ; one or more of the four segments by the Enton following the second usually with a paler (buff, ochraceous-buff, July, 1904, a: or cinnamon-coloured) median area on the dorsal surface ; in River, Lagos some specimens the third and fourth, third, fourth, and fifth, or acTif, 0, even the third to the sixth segments each show a narrow paler w. H. TV. St median longitudinal streak,, which usually fails to reach the hind east of Lagos margin of each segment ; in others the paler area on t.he third sented by the segment, or on the third and fourth segments is as broad as that between Aro on the second, while on subsequent segments the paler area is 16. iii. 1909, reduced to a streak ; lateral margins and posterior angles of “ caught in ca second to sixth segments inclusive smoke-grey, scarcely visible on creek nea from above ; pollinose dorsum of seventh segment varying from Calabar, 14. smoke-grey to cream-buff or buff, often infuscated at base in Q ; mens, in the 1 hypopygium of 8 resembling that of G. palpalis, Rob.-Desv., in (Tropical Afr Oni River, 2, * There is a fringe on the third joint of ‘the antennae, but it is so short Calabar, 16. v ond the hairs composing it are so minute as to be scarcely noticeable when the joint is examined in profile under an ordinary platysoopic lens. IMEN. [ DISTRIBUTION OF GLOSSINA CALIGINEA. 33 i ogate area I shape, but’somewhat larger, and, as seen by careful examination rab-grey or k from the ventral side, somewhat squarer (i.e. its posterior margin ig vertical not quite so much rounded off). vly cream- c Legs ochraceous-buff; coxae buff, outer surface of front coxae *d joint on blotched with dark grey; femora, especially front pair, blotched It in both or streaked with dark grey or greyish-brown, front femora often pale hair * almost entirely dark mouse-grey; last two joints of front and noderately middle tarsi tipped with dark brown ; hind tarsi entirely blackish- apering of brown above. 12) ; pulpi Wilzgs uniformly sepia-brown, fairly dark. Squanaae light ‘. cream-coloured, antisquama hyaline, its fringe brownish. same type Hulteres cream-buff. rhat more The typical specimens of this species are a 8 and Q from on dorsum Southern Nigeria, in the British Museum (Natural History), nore elon- both taken by iKr. G. Garden, Veterinary Officer, 8. Nigeria, unusually the $ on a creek near Akitipupu, 18. v. 1909, the Q in a canoe ellum and on a creek between Aro Chuku and Itu, 16. iii. 1909. on dorsum DISTHIBUTIOR OF G. caliginea, Austen. 1 a dusky Tip to the present time, this species has been received only I to sixth from Southern Nigeria. In addition to the types of the $ region on and Q , the following specimens are contained in the Museum off or pale collection :-1 8, Benin (A. Xllson) ; 1 d, Benin City, 18. iii. the usual * than the 1910, and 1 &, Forcados, 30. iv. 1910 (J. J. Simpon ; presented segments by the Entomological Research Committee) ; 1 S, Sobo Plains, :eous-but?‘, July, 1904, and 1 Q , Sapele (Dr. G. F. Darker) ; 1 S, 1 Q , Owa... rface ; in River, Lagos, August, 1904 (Dr. W. H. W. Strachan, C.N.G.) ; d fifth, or 2 d ,3, 1 Q, Bende, May, 1908 (Dr. Clay: presented by Dr. row paler W. H. W. Strachan, C.M.G.) ; 1 &, 2 Q Q , Oni River, 70 miles 1 the hind east of Lagos, 2, 9, and 29. v. 1910 (Dr. W. A. Lamborn: pre- t,he third sented by the Entomological Research Committ.ee) ; 1 6, 3 Q Q , td as that between Aro Chuku and Itu, “caught in canoe on creek,” :r area is 16. iii. 1909 ; 2 & 8, 1 Q , between Agbabu and Akitipupu, angles of “ caught in canoe on creek,” 17 and 19. v. 1909 ; and 2 8 ,-3,l Q , dy visible on creek near Akitipupu, 18. v. 1909 (G. Garden) ; 1 Q, Old ying from Calabar, 14. v. 1900 (Dr. H. E. Annett). The following speci- )ase in Q ; mens, in the possession of the Entomological Research Committee -Desv., in (Tropical Africa), have also been examined :--3 6 6, 4 Q Q , Oni River, 2, 4, and 9. v. 1910 (Dr. W. A. Lamborn) ; 2 8 $ , is so short Calabar, 16. vi. 1910 (J. J. Simpson). cable when ,ns. D :’ EIONOMICS OF GLOSSINA CALIGINEA. 34 DE

BIONOMICS. Gloss&a caliginea, which, from the data printed above, would appear to be prevalent in parts of Southern Nigeria in the month of May, is evidently prone to attack travellers in canoes. Glossina 1 This has been observed both by Mr. Garden and Dr. Lamborn, Ann& Flies, the former of whom, with reference to the specimens taken by him in a canoe on a creek between Agbabu and Akitipupu, on 8, 0 May 17 and 19, 1909, contributes the interesting note that the mens) 8 flies “ attacked during heavy rain ” ; Mr. Garden also states that of front at the spot at which the flies were encountered there was “ dense (Jstoa cover,” consisting of “ long grass and water-plants.” One of Dr. Lamborn’s specimens (a male, taken on the Oni River, on Grou’ May 9, 1910) bears the following field-note :--(‘ As I was going well-deve, up river in a canoe at 3.0 P.M., this insect bit me on the arm, type ; do and became so distended with my blood that it was unable to segment c tly away.” A female in the possession of the Entomological extending Research Committee, taken by the same collector on the same immediatinanaediat river and date, bears the label :-“ From European’s back and correspon distended with his blood, while canoeing up river.” A male and segments, female from Dr. Lamborn, caught on May 4, 1910, are labelled usually c as being “ off European’s clothes, while journeying up river in a these SE: canoe,” and the remaining specimens of this species received ochraceoi from the same source by the Entomological Research Committee formingfornaing (t are also stated to have been caught on a European’s clothes. hind tars

Head l A FFINITIES AND DISTINCTIVE CIIARL~TERS. posterior Among the Glossinn palpalis group of species, with dark hind ocellar tarsi, constituted by Glossin,a palpalis, Rob.-Desv., G. pallicera, upper p Bigot, G. fuscipes, Newst., G. caliginea, Austen, and G. tachinoides, brown, c We&w., the new species can only be confused with t,he two first bases of mentioned. Care is necwzary at the outset in order to distinguish or dark G. caliginea from G. pnlpalis, but the very real differences between brown&l the two species have been pointed out in the course of the is prom. foregoing description. The usually darker hue and larger size of long am G. caliginea, combined with the characters supplied by the third twenty-f joint of the antennae (dusky coloration, stouter shape, less pro- and darl minent distal extremity, and especially the absence of a conspicuous brown OI fringe of pale hair on the anterior and posterior edges (see Thor Figs. 11 and 13) ), will suffice to differentiate the new species humeral from G. pallicera. times WL a more ‘NEA. DESCRIPTION OF GLOSSINA PALLICEBA. 35

Glossina pallicera, Bigot. above, would (Plate III.) ;eria in the trs in canoes. Glossina pallicera, Bigot, Annales de la SocibtB Entomologique de France, )r. Lamborn, Ann& 1891, Vol. LX, p. 378 ; Austen, “A Monograph of the Tsetse- Flies,” p. 79, Plate It (1903). ns taken by .kitipupu, on 8, 0 .-Length, 8 (8 specimens) 8 to 8.75 mm., 0 (4 speci- ote that the mens) 8 to 9 mm. ; width of head, & ? 2% to 2% mm. ; width o states that of front at vertex, 8 0.6 mm.,0 0.8 mm. ; length of wing, was ‘< dense 8 8 to 8.4 mm., ? 8.6 to S.8 mm. ts.” One of ri River, on Ground colour qf dorsum of thorax olivaceous or olive-grey, with I was going well-developed, dark sepia-brown, longitudinal naarkings of usual on the arm, We ; dorsum of abdomen mgmnay-brown or dark sepia-brown, Jirst BS unable to segnaent and a narrow, median, cuneate area on second (not always ntomological extending to hind margin) buf; in 0, portion of second segment m the same inamediately adjacent to and on each side of cuneate area, and a .‘s back and corresponding region on third and fourth, or third, fourth alad Jiftth A male and segments, sometimes with exception qf hind border in each case, are labelled usually cinnamon-coloured, therefore paler than lateral portions of 3 river in a these segments ; third johat of antennae pale (cream-bu$ to res received ochraceous-buf)), clothed with long and $-fine, pale yellowish hair, i Committee forvaing a conspicuous fringe on front and hind margins (cp. Fig. 13) ; clothes. hind tarsi dark brolun above, last two joints dote-browrz.

Head buff or cream-buff, face and jowls yellowish pollinose, 3. posterior surface smoke-grey, sides of front (parafrontals) and h dark hind ocellar triangle mouse-grey ; frontal stripe cinnamon-rufous, a. pa&w-a, upper portion more or less mummy-brown; oceZZar spot dark . tachinoides, brown, connected with usual dark brown transverse band joining ;he two first bases of vertical bristles ; second joint of antennae buff (brownish ) distinguish or dark brown above), third joint usually long and slender, often ices between brownish or mouse-grey at and towards distal extremity, which Nurse of the is prominent, arista buff, dark brown beneath, terminal joint arger size of long and slender, branched hairs eighteen to twenty-three or by the third twenty-four in number ; palpi buff or cream-buff, greyish above pe, less pro- and dark brown at tips; proboscis bulb raw-umber-coloured, dark conspicuous brown on upper edges. - edges (see Thorax.--Humeral calli, pleurae, and pectus smoke-grey, new species numeralL .__^^.. ., tiam--,,: -.-ILLw~lru a_ uarlcJ.-L uruwnL._- ___. sp--.L-.--A’ob next upper - ^... margin, -.--.-f.. some...- times wanting or indistinct ; scutellum raw-umber-coloured, with a more or less sharply defined dark brown blotch, roughly 36 G. PALLICERA: ABDOMEN AND WIKGS. DIST. triangular in outline, on upper surface on each side of middle line ; apical scutellar bristles in $! sometimes very short, but varying in length in different individuals from one-sixth to one- Glossin half of that of those of 6. apparently Abdomen.-Posterior angles of third to sixth segments and of whi inclusive, seventh segment entirely, and hypopygium of 8 the Sierra isabella-coloured or drab, pollinose ; interspace between hectors are the dei of 6 very narrow or non-existent. Collection’ Legs yellowish clay-coloured ; front femora infuscated on (Major A. inside (an especially dark blotch on distal third, not extending 1 0, Suji, R.AXC.) 1 $J , Assir from the Kumasi, 1 (forwarded Graham, 1 on N’Kaw path on t 3.0 P.M.” (. 1907, “ cal Southern (J. J . Six Committee As reg: by Profess (Mayo-Kal, Dr. Ducar

Fig. 13. species app Antenna of Glossha pallicera, Bigot. (x 30.) to tip), middle and hind femora more or less tinged with mouse- grey on outer side, and on under side with an elongate clove- For tht brown blotch near distal extremity, usually connected with a are indebt smaller mark of similar colour on outer side just before tip ; Dr. W. M distal margin of penultimate joint of front and middle tarsi found the brown or dark brown, rather less than distal half of last joint of ground, bu front and middle tarsi dark brown, extreme tips of remaining Tsetse as joints of front and middle tarsi often tinged with mummy-brown. Dr. Graha Wings with a strong brownish tinge. Squamae cream-coloured. path, bord Halteres buff. This fly a

* Cf. A. p. 254 (1908) WIKGS. DISTRIBUTIOX OF GLOSSINA PALLICERA. 37 side of middle DISTRIBUTION OF G. pallicera, Bigot. ry short, but csixth to one- Glossina pal&era is a West African species, which is apparently one of the rarest of the Tsetse-flies at present known, rth segments and of which the area of distribution extends, at any rate, from pygium of 8 the Sierra Leone Protectorate to French Congo. The following tween hectors are the details with reference to t,he specimens in the National Collection’:-Sierra Leone Protectorate : 1 6, Firo, 1’7. iv. 1909 .nfuscated on (Major A. Pearse, R.A.M. C.). Liberia : 1 Q , Tappoima, 11. v., lot extending 1 Q , Suji, 21. v., 1 8 , Bonnatown, 23. v. 1909 (Major A. Pearse, R.A.N.C.). French Ivory Coast : 1 8 (the type of the species), 1 Q , Assinie (Ch. Alluaud; presented by Mr. G. H. Verrall, from the collection of the late M. Bigot). Ashanti : 1 Q , Kumasi, 19. vii. 1907, “ caught on horse on polo ground ” (forwarded by Dr. T. E. Rice, S.M.O. ; presented by Dr. W. M. Graham, W.A.M.S.) ; 3 & $, Kumasi, 26, 27. x. 1907, “caught on N’Kawi Road, 11.0 AM,” “ caught on palm stump in bush path on the N’Kawi Road,” and ‘(caught on N’Kawi Road, 3.0 P.M.” (Dr. W. M. Graham, W.A.M.S.); 1 8, Prahsu, 16. xii. 1907, “ caught, behind kitchen ” (Dr. W. M. Graham, W.A.M.S.). Southern Nigeria : 1 8, Ojogbo, Benin City, 16. iii. 1910 (J. J. Simpson : received from the Entomological Research Committee). As regards French Congo, a specimen of G. pallicera is stated by Professor Laveran to have been taken on the Benue River (Mayo-Kabbi region), in January or February, 1907, by Dr. Ducasse.* In Ashanti, according to Dr. Graham, the species appears to be “ extremely local.”

BIONOMICS. with mouse- mgate clove- For the few details that can be given under this heading we cted with a are indebted to the observations made in Ashanti, in 1907, by ; before tip; Dr. W. M. Graham, who informed the author that he always niddle tarsi found the fly in the “ bush,” where it ‘( does not settle on the last joint of ground, but sits on the upper side of leaves.” Writing of this f remaining Tsetse as met with by him on the N’Kawi Road, Kumasi, nmy-brown. Dr. Graham says :---‘I G. pallicera was found in a narrow bush Lm-coloured. path, bordered by short bushes, but free from tall trees. . . . This fly appeared strictly localised. I found it only on this

* Cf. A. Laveran, Bulletin do la SociBtb de Pathologie Exotique, T. i, 38 GLOSSINB PBLLICERA : AFFINITIES, ETC. narrow path. It did not associate with G. palpalis upon this path, nor was it to be found associated with G. palpalis upon the main road into which the path opens. The insects were found sunning themselves upon the leaves of the tall plants Gloss: bordering the path, or playing in sunshine about the stumps of P a fallen palm (Elaiis guineensis). I caught a number of males, P but t,hough I spent some hours t’here, and returned to the path P: c on a second occasion, when I again caught males, I failed to N catch a female on either occasion. Though I sat among the ,Gloss insects for some hours no attack was made upon me by them.” * F With reference to the last remark by Dr. Graham, it may be Gloss noted that t’he abdomen of the female fly caught on a horse on B the polo ground, Kumasi, as recorded above, is distended by coagulated blood. In Ashanti it appeared to Dr. Graham that d in the case of G. pulpalis the females were more aggressive than spew the males, and possibly this is also true of G. pallicera. 0 2 ,0*75

A FFINITIES AND DISTINCTIVE CHARACTERS . 7.8 n

With Glossina palpalis, Rob.-Desv. (p. 24, Plate I), G. 0 fuscipes, Newst. (p. 29), G. caliginea, Austen (p. 30, Plate II), ,of th and G. tachinoides, West-w. (p. 39, Plate IV), G. pallicera mark’ forms the Glossinrr palpalis group of Tsetse-flies, the members .abdor of which are characterised by all the joints of the hind brows tarsi being wholly or for the most part blackish-brown, clove- with brown, or dark brown above. With G. tachinoides, owing to ‘wV its abdominal markings and much smaller size, there is no risk or lea of confusion (compare Plates III and IV) ; from all other secant species of the Glossina palpalis group at present known, joint G. pallicera, although exhibiting an exceedingly close resemblance in general appearance to an unusually light-coloured G. caliginea, b can at once be distinguished by the coloration, shape, and of frc hairiness of the third joint of the antenna, as described above. blot& betwc * From “ Report by Dr. W. 11. Graham upon Entomological Observa- ,ochra tions made in Southern and Central Ashanti, 1907,” p. 3 (printed for private circulation by the Colonial Office, 1909). trian; ,dark anten brow: side,

* 1 the M blood, [ES, ETC. DESCRIPTION OF GLOSSINA TACHINOIDES. 39 lalis upon this Glossina tachinoides, Westw. palpalis upon e insects were (Plate IV.) le tall plants Glossina tachinoides, Westwood, Proc. Zool. Sot. Lond., Pt. xviii, p. 267, the stumps of Plate XIX, fig. 2 (1850) ; Ann. Mag. Nat. Hist., Ser. 2, Vol. X, nber of males, p. 147 (1652); Austen, Ann. Mag. Net. Hist., Ser. 7, Vol. XIV, ?d to the path pp. 152-154 (1904) ; British Medical Journal, September 17, 1904: 3s, I failed to Captain R. Markham Carter, I.M.S., British Medical Journal, November 17, 1906, p. 1393. tt among the ,Glossina palpalis var. tachinoides, Austen, (‘ A Monograph of the Tsetse- ae by them.” * Flies,” p. 74 (1903). ,m, it may be Glossina Decorsei. Brumnt. Corn&es rendus des seances de la Soci& de on a horse on Biologie (SQance du 1s kvril, i904), T. lvi, p. 628 (1904). distended by Graham that $, 0 .-Length, 8 (9 specimens) 6 to 6.75 mm.,” 0 (9 gressive than .spe;unensj 63 to S.4 mm. ; width of head, & 2 to 2.2 mm., era. 0 2.3 to 2.25 mm.; width of front at vertex, & 0.5 mm., 0 ,0.75 to 0.S mm. ; length of wing, & 6 to 6.5 mm., $’ 7 to

RS. 7.8 mm.

Plate I), G. ONE OF THE SXALLEST OF THE KXOWN TSETSE-FLIES.-DOTSUT ;O, Plate II), of thorns olive-grey or smoke-grey, with dark brown longitudinal G. pallicera markings qf usual type, though qften indistinct; ground colour of the members .abdomeu ochsaceous-buff or buff, dorsum with sharply dejned cloue- of the hind brown, or dark brown markings as shown AZ Plate IV, second segment brown, clove- with CL aery conspicuous, square or oblo2lg, pale (ochraceous-buff or les, owing to bu$) area in centre; hind tarsi dark brown, i.e. all the joints more re is no risk or less dark, extreme tip.3 of @at three joints and extreme bases of lrn all other second and third joints buf, a larger or smaller aren at base of first * sent known, joint also paler thall remainder of joint, especially in 0 . I resemblance G. caliginecc, Head cream-buff pollinose, posterior surface smoke-grey, sides shape, and of front (parafrontals) and ocellar triangle light gre.y, a darker )ed above. blotch s&etimes visible on each pa&frontal about midway between ocellar spot and base of antennae ; frontal stripe gical Observa- ,ochraceous or tawny-ochraceous, ramus on each side of ocellar ) (printed for triangle brownish ; ocellar spot dark brown, connected with usual dark brown transverse band joining bases of vertical bristles ; antennae partly brown or brownish, partly buff, second joint brown or dark brown on inner surface and usually buff on outer .side, sometimes brownish on outer side also and buff only at tip,

* A 6 from the Benue River, Northern Nigeria (Dr. K. Mackay), in +hr. Tld..nn..- ,,I,,,+:,.- +I.,? nl.,lnm..., ,%a- ..rL:nL :” A:..en”A,4 L-7 ,,~“nlrl”+,4 40 G. TACHINOIDES: THORAX, ABDOMEN, ETC. DISTRIBUTIC

third joint greyish-brown or mouse-grey, more or less tinged with recorded from th buff at base, arista buff, second joint and proximal half of t’erminal Africa Protectorr joint dark brown below, branched hairs fourteen to sixteen in Africa, since G. number ; pal+ buff-yellow, greyish-brown at tips ; proboscis bulb German East Afr mummy-brown or raw-umber-coloured. Arabia,? thus bei Thorax.-Humeral calli and pleurae drab-grey, former with of Glossina found a dark brown spot or blotch above (sometimes wanting or the islands off the indistinct); pectus smoke-grey; scutellum cream-buff, with a dark As regards th brown blotch, roughly triangular in outline, on upper surface on noides has been id each side of middle line; apical scutellar bristles in 0 about Marigot de Bay one-third of length of those of 8. Lower Rio Nun Abdomen.-Median quadrate pale area on dorsual of second ; the ban1 segment cery conspicuous, and (regarded in conjunction with the Sudan ; and the 1 dark hind tarsi) distinctive, owing to its shape ; dark brown or Of this specie clove-brown interrupted bands on subsequent segments, as shown present possesses in Plate IV, not reaching either lateral or posterior margins ; to which are as fc seventh segment cream-buff or smoke-grey pollinose, in 9 some- Gambia: 1 6 times with a trace of dark markings like those on preceding Gold Coast : 1 d segment ; hypopygium of 8 oval, pale isabella-coloured. 1908, and 1 8, i. liegs buff, hind tarsi as described in diagnosis printed in (,G. C. Dudgeon) ; italics above ; front or front and middle femora more or less grey Territories, “ cauj on outer side, hind femora also sometimes with a greyish infusca- caravans), by a w tion on outer side towards distal extremity, middle femora with dry bed of a sl a conspicuous dark brown elongate blotch on under side near tip,. Northern Nigeri hind femora with a trace of a similar mark ; tips of last two 3 & 8, Pakim, I joints of front and middle tarsi mummy-brown, tips of remaining Benue R., and 2 joints of middle and sometimes also of front tarsi faintly 3&&,l?,Kan brownish. Benue R., 26. v. Wings tinged with drab. Squanzae waxen-white, fringe short, v. 1904, 1 &,3 ’ yellowish. Halteres cream-coloured. two days below 1 The foregoing re-description is mainly based on a 8 and ? in * Apud Stuhlmr the British Museum (Natural History), taken at a spot six miles Heft 3, p. 301 (1907) south of Mulgue, Bornu Province, Northern Nigeria, on July vicinity of Amani, a t Cj. Captain R 22nd, 1910, by Dr. B. Moiser, W.A.M.S. British Jledical Jc invalidate the gener Ethiopian Region, I DISTRIBUTION OF G. tackinoides, We&w. graphically to the 16 b other African insecl G. tachinoides has a very wide range in West Africa, where, Mandeb. 1 Cf. A. Laveran as for instance in Northern Nigeria, it is often locally abundant. t&s6 dans 1’Ouest a Besides being found from Senegal to French Congo, the species (Comptes rendus dt 932 (1905)) ; and also occurs in the French Sudan (on the shores of Lake Chad and l’ilfrique intertropic the banks of the Bani and Shari Rivers) ; and, although not yet T. i, pp. 252-255 (19 :EN, ETC. DISTRIBU’l?ION OF GLOSSINA TACHINOIDES. 41 tinged with recorded from the Congo Free State or the. Uganda or East E of terminal Africa Protectorates, its range evidently extends right across 0 sixteen in Africa, since G. tac7inoides is not only reported to exist in proboscis bulb German East Africa,* but has also been met with in Southern Arabia,t thus being so far as is known the only existing species former with of Glossina found outside the African continent and certain of want’ing or the islands off the coast. with a dark As regards the French possessions in West Africa, G. tachi- r surface on noides has been identified by Professor Laveran from Casamance, in p about Marigot de Bayla, Bignona, and Carabane in Senegal; the Lower Rio Nunez, Bake, and the Tinkisso River in French nl of second Guinea ; the banks of the Bani and Shari Rivers in the French m with the Sudan ; and the Mayo-Kabbi region in French Congo.$ k brown or Of this species the Brit&h Museum (Natural History) at is, as shown present possesses 44 8 6 and 69 0 0, the details with reference )r margins ; to which are as follows :- in ? some- Gambia: 1 8 , 2 ? 9 , Gambia River, 1907 (G. C. Dudqeon). a preceding Gold Coast: 1 8, Lorha R., Northern Territories, February, :d. 1908, and 1 8, 2 0 0 , Volta R., north of Krachi, March, 1908 printed in (,G. C. Dudgeon) ; 1 0, between Salaga and Makongo, Northern or less grey Territories, (( caught at Tonga (temporary resting-place for native ish infusca- caravans), by a water-hole, empty though still damp, dug in the ‘emora with dry bed of a stream,” 14. ii. 1910 (Dr. T. J. A. Beringer). de near tip, Northern Nigeria : 2 ? 0 , Yola, Benue R., 12. iii. 1904, of last two 3 8 6, Pakim, Kilengi R., 24. iv. 1905, 2 0 0 , Mutum Biu, E remaining Benue R., and 2 9 $‘, near Mairanewra, Benue R., 24. v. 1904, trsi faintly 3 6 6, 1 ? , Kamberi, Benue R., 25. v. 1904, 1 0, near Amara, Benue R., 26. v. 1904, 2 8 8, 8 ? 0, Jibu, Benue R., 28, 29. tinge short, v. 1904, 1 6, 3 Q 0, near Ibi, Benue R., 29. v. 1904, 18, 10, two days below Ibi, Benue R., ‘I biting donor in canoe,” 2. vi. f and $’ in * Apd Stuhlmann (Arbeiten a. d. Kais. Gesundheitsamte, Bd. xxvi, ot six miles Heft 3, p. 301 (1907) ), G. tachinoides occurs in German East Africa, in the a, on July vicinity of Amani, at the foot of the Usambara Mountains. t Cf. Captain R. Markham Carter, I.M.S., “ Tsetse-Fly in Arabia” : British Medical Journal,, November 17, 1906, p. 1393. This does not invalidate the generalisation that existing Tsetse-flies are confined to the Ethiopian Region, since the southern portion of Arabia belongs zoo-geo- . graphically to the region in question. In addition to G. tacizinoides, many other African insects occur on the eastern shore of the Strait of Bab-el- ica, where, Mandeb. 1 Cf. A. Laveran, “ Contribution I l’etude de la repartition des mouches abundant. t&s6 dans 1’Ouest africain franpais et dans l&at independant du Congo ” the species (Comptes rendus des seances de l’bcademie des Sciences, t. cxli, pp, 929- 932 (1905)) ; and “Contribution a l’etude des mouches piquantes de D Chad and 1’Afrique intertropicale ” (Bulletin de la Societe de Pathologie Exotique, igh not yet T. i, pp. 252-255 (1908). 42 GLbSSINA TACHINOIDES IN NIGERIA. GLOSS1 (Major C. B. S 1904, 1 8, 1 Q , three days below Ibi, Benue It., 3. vi. 1904, larva, 3 pupa-1 2 Q Q , four or five miles below Albins, Benue R., 4. vi. 1904, Arabia : 1 Q , 1 1 Q , ten miles below Amagede, Benue R., 7. vi. 1904, 1 Q; three Al Hay section miles above Loko, Benue R., 5. vi. 1904, 1 8, 2 Q Q , near The species Dakmon, Kaduna R., 21, 22. vi. 1906, 1 d, 1 Q , near Mureji, distributed alor Kaduna R., 22, 24. vi. 1906, 1 8, l- Q ,- near Egga, Niger R., In Katagum P 24. vi. 1906, 1 6, 1 Q , near Muye, Niger R., 3 Q Q near Baro, found G. ta& Niger R., 25. vi. 1906 (W. F. Gozcers) ; 2 8 6, 3 Q Q , Benue R., be common in 1. x. 1905 (Dr. K. Mackn~) ; 1 6, 1 Q , Pateji, Niger R., Ilorin miles to the e: Province, November, 1904, “ caught in the bush” (received from Province, on J Sir F. D. Lugard, K.C.M.G.) ; 1 8 , Adiale, Bassa Province, found G. ta&n, November, 1906 (Dr. G. J. Pirie, m7.AX.S.) ; 16, Niger R., at at that time ‘( 1 junction of Menai R., 19. xi. 1906,*” taken on horse’s fetlock, in surrounded by I shade of leafy trees close to water’s edge, where there is a stone trees grew on tl cliff,” 2 Q Q ,* Niger R., right bank, between Yantalah and Borgu, high . . . The 29. xi. 1906, “ on canoe-men” (Major C. B. Simonds, R.G.A.) ; about thirty mi 1 Q , Kaoge, 1. vi. 19Oi, 1 &, Garran Gabbus, July, 1907, 2 Q Q , grazed.” t Aida and Dongwan, Ka R., 18. vii. 1907, 2 Q Q , Loko, In Nigeria i Benue R., and 1 Q , Katsena Allah, August, 1907 (J. Bran,d) ; that the areas ( 3 Q Q , South Bornu, September, 1907 (Dr. H. A. Fey, W.A.M.S.) ; nate with thoa 2 8 &, Benue It., 2. vii. 1907 (Dr. J. i&F. Pollard, W.A.M.S.); , the author refl 4 Q Q , Dugurie, October, 1907, ‘

I’ BIONOMICS. after dark. net, and hi Writing in the Sleeping Sickness Bureau Bulletin No. 6 than mosqu (May 7th, 1909), p. 210, Dr. A. G. Bagshawe remarks that synonym G Glossina tachinoides “ at least _ in Nigeria and on the Lake Chad the shores ( waterways, is confined to river banks as is palpalis and occurs in the ily wit1 large numbers, biting man. It is therefore allied in its habits to the wate to palpalis.” That the species with which we are dealing is are more I found only in close proximity to water certainly seems to be the bite of generally true with regard to the region referred to, though in species of ( Southern Arabia Captain R. M. Carter did not always meet after the bi with it near the edge of water, and no information is at present In Cent available as to the haunts of G. facl~inoides in German East. is a migrat Africa. Since G. tachinoidcs belongs to the G. pulpalis group of present in L the genus Glossina, it is natural that its habits should be similar from May to those of the recognised carrier of sleeping sickness. Recently considerabl in South Bornu, Northern Nigeria, Dr. Alexander found is the same G. tachinoides in a large marsh consisting of long elephant grass for more op mixed with occasional clumps of palm trees and thick under- wild palms growth, though he failed to discover the fly on the banks of a Dahomey d river some three quarters of a mile away.” Dr. Dalziel, writing moisture, C from Yola Province, Northern Nigeria, states that “ Glossina beyond the tachinoides seems to be found only close to the river bank, in may be met sight of water almost, though it may follow along the path. On the The longest distance to which it was found to follow away from Dr. G. J. water was 1,200 yards (one fly). Glossina morsitans on the “ restricted other hand, has not been seen directly at the river bank, being other. ‘ Tl always out of sight of water, in bush with thick grass and shrubs, stream cam trees, and climbers. Both species are found together on paths was coverer through the bush near water.” t In the Western and Northern or two pla Provinces of Ashanti, on the Tain and ‘Black Volta Rivers, Dr. were very c A. Kinghorn finds that in habitat and habits G. tachin,oides fly was seer closely resembles G. yalpalis, in company with which it always could be o occurs. According to Kinghorn, the present species “ will feed Nigeria, wl- on man with avidity,” and is “ quite active in dull weather and found G. ta in the very early hours of the morning, both times when Gloss&a on the Bad palpalis is usually quiescent.” $ According to Dr. Alexander, on a river near Amar, in South Bornu, ” Glossina tachinoides bites * Sleepiq (February 15 * Cf. Sleeping Sickness Bureau Bulletin, Vol. III, NO. 24, p. 93 t Brumpt (February 15,1911). $ Compte: t Sleeping Sickness Bureau Bulletin, Zoc. cit. (FBvrier 13? 1 1 Cf. Sleeping Sickness Bureau Bulletin, Vol. III, No, 25, p. 136 Q Sleepmg (March 14, 1911). (April 11, 191 :INOIDES. BIOXOMICS OF GLOSSINA TACHINOIDES.. 45

after dark. At 7.0 p.m. he had to take refuge in his mosquito net, and his boys remarked that the flies were more troublesome Bulletin No. 6 than mosquitoes.” Se Brumpt,t writing of this species under its e remarks that synonym G. decorsei, says that in the Shari River basin and on the Lake Chad the shores of Lake Chad, where the natives associate the bite of lis and occurs in the fly with disease among their cattle, it appears to be confined ed in its habits to the water’s edge, and that in the month of March the males 3 are dealing is are more numerous than the females. According to Brumpt , oly seems to be the bite of G. tachinoides “ is disagreeable, as is that of all the :d to, though in species of Glossina, but not very painful ; it causes some time ot always meet after the bite a rather acute itching.” m is at present In Central Dahomey, according to Roubaud,$ G. tachinoides 1 German East. is a migratory species, which comes from the north, and “ is zlpalis group of present in abundance on the banks of the large watercourses only .ould be similar from ‘May to August, when the rains are at their height and less. Recently considerable changes in the river-systems take place. Its habitat 3xander found is the same as that of 0. palpalis, though it displays a preference elephant grass for more open wooded tracts, and is not found in the groves of 1 thick under- wild palms along the small streams.” Roubaud states that in ;he banks of a Dahomey during the wet season, when the air is saturated with ialziel, writing moisture, G. tachinoides, palpalis, and longipa&s are all found ;hat (’ Glossina beyond the limits of their usual haunts, and isolated specimens river bank, in may be met with in the Savannah zone, far from any watercourse. ong the path. On the Wobe River, North Bornu, Northern Nigeria, 1ow away from Dr. G. J. Pirie, in November, 1909, found G. tachinoides rsitcots on the “ restricted to patches sometimes on one bank, sometimes on the er bank, being other. ‘ The fly patches always occurred where the main river ss and shrubs, stream came right up to its bank, and the bank at the patches ther on paths was covered with river bush with thick undergrowth. At one and Northern or two places the ‘ fly ’ came off to the barge in numbers, but #a Rivers, Dr. were very difficult to catch. Where the river bush was thin, no G. ta&nodes fly was seen.’ No evidence of trypanosomiasis in man or animals ich it always could be obtained.” S At Bebua, Bauchi Province, Northern ies ‘( will feed Nigeria, where, as already mentioned, Dr. J. McFarlane Pollard 1 weather and found G. tachinoides ‘(in far greater numbers than G. palpalis vhen Gloss& on the Bade River (a small stream) and its little tributaries,” r. Alexander, mhinoides bites * Sleeping Sickness Bureau Bulletin, Vol. III, No. 24, pp. 92, 93 (February 15,191l). No. 24, p. 93 t Brumot. ZQC. cit. 1). 629. i Cornpies Rendu;de l’Aca&mie des Sciences, T. 152, No. 7, p. 406 (FQvrier 13! 1911). ?o. 25, p. 136 § Sleepmg Sickness Bureau Bulletin, Vol. II, No. 16, pp. 146-147 (April 11, 1910). 46 G. TACHINOIDES : REPRODUCTION, ETC. G. TACHINOI: the cover consisted of “ long thick grass, and trees overhanging than in the cast the stream.” In a note attached to his specimens, Dr. Pollard p. 7j, though t adds that there was “ plenty of shade, but the trees were by no much bigger fly 1 means continuous.” tumid lips are In Southern Arabia, G. tachinoides was found by Captain between them is R. M. Carter sparsely and locally in thick belts of euphorbia, pupa of G. morsii babal thorn, and tamarisk, and also in cactus belts. It was and 6 c, p. 7). never seen “in the date groves or along patches of cultivation,” the characterist. and was not always met with near the edge of water. Captain G. pulp&s (corn1 Carter writes :-“ The Arabian Glossina tachinoides does not depend for its existence on big game, for, excepting gazelle, S YNONYMY , nothing else frequents the belts of bush which it haunts. Only once was the fly recognised by Arabs, and then by a band of The identity ( wild Bedouins from North-Western Subaihi. . . . They stated this Westw., was pro tly used to be more frequent in Subaihi, that it bit goats, to the original donkeys, horses, dogs, and men, but did not attack camels or author by Dr. B sheep. They further stated that it sucked blood only after the G. tachinoide, spring rains, that some years it disappeared from a district also one of the entirely, that rarely it became abundant, and then they moved readily distinguie their settlement and cattle by night marches to a fly-free group, to which i district.” * of its abdomen, i to G. pallidipes, REPRODUCTION AND PRELIMINARY STAGES. species of its own Roubaud, who in Dahomey has observed the behaviour of pale area on the G. ta&nodes in captivity, states that, as in the case of G. has already been pulpal& under similar conditions, copulation takes place directly to be dist,inguishe after the flies are hatched. At 25” C. (77” F.) larvae are even though the deposited on the average at intervals of eight days ; the duration * Cf. E. E. Awl of the pupal stage at a mean temperature of 24” to 25’ C. Glossina, Wiedemar at the Annual Mee (75.2” to 77” F.) was found to be from twenty-eight to thirty-five 1904 : British Medic days.? A dried and apparently not quite mature larva of this species, from Amani, German East Africa (Dr. F. Vosseler), measures 6 mm. in length, but, since the specimen is unfor- tunately somewhat crushed, its maximum breadth cannot be stated. Ap~ppa (Figs. 5 E and 6 E, pp. 5 and 7) from Amani, also presented by Dr. Vosseler, is 5.5 mm. long, by 3 mm. in greatest width. These specimens show that the most striking characteristic of the larva and pupa of G. tachinoides is the relatively large size of the tumid lips, which are actually larger * Captain R. 11. Carter, I.iKS., Zoc. cit. t Cf. Roubaud, lot. cit., p. 407. X, ETC. G. TACHItiOIDES : SYNONYMY AND AFFINITIES. 47 s overhanging than in the case of G. pctllidi]~es (compare Figs. 6 JZ and 6 D, s, Dr. Pollard p. 7), though the perfect insect of the latter is of course a es were by no much bigger fly than G. tnchinoides. In the present species the tumid lips are just under 1 mm. in length, and the notch d by Captain between them is similar in shape to that between the lips of the of euphorbia, pupa of G. morsitans, although not so narrow (compare Figs. 6 E belts. It was and 6 C, p. 7). The notch is much wider than, and has not cultivation,” the characteristic shape of that exhibited by the pupa of ter. Captain G. pdpnlis (compare Figs. 6 E and 6 F, p. 7). ides does not pting gazelle, SYNONYMY, AFFINITIES, AND DISTINCTIVE CHARACTERS. launts. Only by a band of The identity of Glossinn decorsei, Brumpt, with G. tachinoides, ey stated this Westw., was proved by the examination of specimens belonging it bit goats, to the original series of G. decorsei, kindly submitted to the author by Dr. Brumpt.* (ck camels or G. tnch’noides, which, besides being one of the smallest, is mly after the also one of the most easily recognised members of it’s genus, is m a district readily distinguishable from the other species of the G. pu,lpalis a they moved group, to which it belongs, owing to t,he colouration and markings to a fly-free of its abdomen, in which respects it presents a closer resemblance to G. pallidipes, Austen, or G. lom@l~~is, \Vied., than to any ES. . species of its own group. The characteristic shape of the median behaviour of pale area on the second abdominal segment, to which attention e case of G. has already been drawn, will however at once enable G. tnchinoides place directly to be dist,inguished from any species of the 6. worsitnns group, ) larvae are even though the hind tarsi be missing. the duration * Cj. E. E. Busten, “ Supplementary Notes on the Tsetse-Flies (Genus 4” to 25O c. Glossina, Wiedemann) ” : a Paper read in the Section of Tropical Diseases at the Annual hleeting of the British aledical Association, Oxford, July, to thirty-five 1904 : British 1Iedicsl Journnl, September 17, 1904.

arva of this F. Vosseler), Ien is unfor- i cannot be I Amani, also 7 3 mm. in lost st’riking co&s is the :tually larger 48 DESCR!

9.6 mm. ; wid 3 mm. ; width length of wine CHAPTER V.

THE GLOSSINA MORSITANS GROUP. Dorsum oj in 6, drab-gre SYNOPSIS OF SPECIES. pletely developt 1. Last two joints of front and middle tarsi with streaks ; . dorsa sharply defined, clove-brown or black tips.. . 2 specimens son Last two joints of front and middle tarsi with- out sharply defined, clove-brown or black mortem chaq tips (front and middle tarsi either entirely (sometimes ind pale or, at most, last two joints of front the second seg tarsi faintly brownish at the tips, and last each with a cef joint and distal half of penultimate joint of in the mediar middle tarsi light brown-never so dark as to form a sharp contrast with the remaining extending beyol joints) ...... _...... pallidipes, Austen. legs bu$, last t 2. Third joint of antennae with a distinct fringe joints of from of fine hair on front margin; dark brown spicuously tip! or clove-brown bands on abdominal seg- middle tarsi t ments extending close to hind margins (i.e. pale ground colour, apart from the median distal hay or interspace, confined to a very narrow hind or clove-brown, border)...... lo?zgipalpis, Wied. pale. Third joint of antennae without a distinct fringe of fine hair on front margin; dark Head brown or clove-brown bands on abdominal crew segments not extending close to hind posterior surfa marg~ns...... morsitans, Westw. grey, sides of blotch on up] a Glossina morsitans, We&w.” smaller and 1~ (Plate V.) or posterior er Glossina morsitans, Westwood, Proc. Zool. Sot. Lond., Part xviii, p. 261, spot dark gre Plate IX, figs. 1, la-lf (1850) ; Kirk, “ On the ‘ Tsetse ’ Fly of Tropical sometimes COI Africa” (Glossi~~a nao~itans, Westwood), Journ. Linn. SOC., Zoology, first and secon Vol. VIII, pp. 149-156 (1865) ; Austen, ‘( A Monograph of the Tsetse- of second joint Flies,” p. 81, Plate III (1903), and British Medical Journal, basal angle bl September 17th, 1904 ; Sander, “ Die Tsetsen,” pp. 60-63 (Leipzig : J. 9. Barth, 1905); Newstead, Bulletin of Entomological Research, mate joint at Vol. II, Part 1, p. 32, fig. 16 (male genitalia) (April, 1911). terminal joint 6, $! .-Length, 8 (15 specimens, from various localities) number ; palp 7.2 to 9 mm., ? (11 specimens, from various localities) 8.6 to dark grey; pr, Thorax.--l * The designation Gloss&a morsitans, as used in this book, includes incomplete ve G. submorsitans,~Newst. As explained on pp. 51,52, thi: author, provisionally at any rate, is mclined to regard G. .slLbmoysitans, Newst:, as a form or Rob.-Desv. (CC race of G. morsitans, Westw., rather than as a distinct species. drab-grey in $ llES&~IPTION OF GLOSSINA MORSITANS. 49

9.6 mm. ; width of head, & 2.4 to 2.75 mm., Q 3.5 to just under 3 mm. ; width of front at vertex, & 0.6 mm., Q 0.8 to 1 mm. ; length of wing, 6 7.6 to 8.2 mm., Q 8.3 to 9.5 mm.

UP. Dot-sum of thorax light grey, otivaceous-grey, or smoke-grey in 6, drab-grey in Q , the thorucic markings in both sexes incom- pletely developed, am2 reduced to brownish or mouse-grey longitudinal streaks ; . dorsum of abdomen bu$ to ochraceous-bu@’ (in pinned 2 specimens sometimes mouse-grey or olivaceous owing to post mortem changes), with a larger or smaller clove-brown blotch (sometimes indistinct or almost zctanting) near each basal angle of the second segment, and the third to the sixth segments inclusive each with a nery conspicuous clove-brown transverse band, interrupted in the median line, not rea,ehing the laternl margins, and not extending beyond the basal three-fourths of the segment, if so far ; legs buff, last two joints qf hind tarsi clove-brown or black, last two joints of front tarsi and penultimate joint qf middle tarsi con- spicuously tipped with clove-brown or dark brown, last joint of middle farsi entirely da& brown above in typical race, other&e distal half or third qf last joint of middle tarsi alone dark brown or clove-brown, remainder of joint merely brozonish or even entirely pa&s, Wied. pale.

Head cream-buff, face and ,jowls cream-coloured pollinose, posterior surface olive-grey or smoke-grey, ocellar triangle smoke- itans, We&w. grey, sides of front (parafrontals) in Q each with a dark brown blotch on upper part of lower half, wanting or at any rate smaller and less conspicuous in $ ; frontal stripe buff, its upper or posterior extremity often more or less mummy-brown ; ocellar ,t xviii, p. 261, spot dark grey, spots at base of vertical bristles dark brown, Ply of Tropical sometimes connected by a transverse band of similar colour ; h., Zoology, first and second joints of antennae dark brown, distal extremity ,f the Tsetse- of second joint buff, third joint mouse-grey or drab-grey, posterior ical Journal, -63 (Leipzig : basal angle buff, front margin not fringed, arista buff, penulti- cd Research, mate joint at base and lower margin of proximal extremity of terminal joint dark brown, branched hairs about twenty-one in 3 localities) number; palpi buff, yellowish-grey above, tips mouse-grey or ties) 8.6 to dark grey ; proboscis bulb clove-brown or dark brown. Thorax.-The markings on the dorsum are more or less look, includes provisionelly incomplete vestiges of those exhibited by Glossina palpalis, hs a form or Rob.-Desv. (compare Plates V and I) ; pleurae smoke-grey in $ , drab-grey in Q , pectus smoke-grey ; scutellum cream-buff’, upper I%2 50 G. MORSITAKS : ABDOMEN AND LEGS. G. M 1 surface with a dark brown or brownish patch, triangular in dark patch 1 outline, on each side of median line ; apical scutellar bristles in under side o ? extremely short, reduced to mere stumps. middle and hi Abdomen.-On dorsal surface, basal angles of second segment greyish, seventh segment, posterior margin of sixth and sometimes also that of fifth segment, and lateral margins and posterior angles of second to sixth segments inclusive light grey pollinose ; the clove-brown, interrupted, transverse bands taper more or less abruptly towards the basal angles of the segments on which they are present, and at the median interruption, though sometimes straight-cut, are, in the more typical forms, generally more or less rounded off or even partially obliterated ; the obliteration of

Distal po aspect), FI side ; penultin base. lVing8 fain cream-coloured Halteres crean

Fig. 14. Distal portion of al~lomen of Glossina morsitnns, Wesrm., g (ventral Glos, aspect), showing (IL) hypopyginm and the hectors * above it. (X 15.1 Glossim submon bands, when it occurs, usually affects the lower inner angles and ology, Serie: mediad portion of the hind margins of the two halves of the Bulletin of bands affected, or, at any rate, the inner extremities of the p. 34 (male ( two halves of the bands, so that, even when the bands are Relying ch much obliterated, their outer extremities are always visible ; appendages of seventh segment in 0 with a more or less distinct indication of this form as a an interrupted clove-brown band at base ; hypopygium of 8 but after mak (Fig. 14) greyish ochraceous-buff, large as compared with that of large number G. longipalpis, Wied. (compare Figs. 14 and 15). Westw., receil Legs.-Front femora on inside, middle and hind femora on Museum (Nat outside more or less infuscated (mouse-grey or greyish clove- Africa, and COI brown), outside of front femora also in some cases with a small species, which * See p. 14 (first paragraph and note *). G. MORSITANS, FORM SCBMORSITANS. 51 liar in dark patch beyond middle ; a dark brown elongate blotch on ItIes in under side of middle and hind femora near distal extremity; middle and hind tibiae sometimes more or less infuscated on outer 3gment retimes sterior linose ; or less h they tetimes Lore or tion of

FiS. 15. Distal portion of abdomen of Glossina longipdpis, Wied., ,3 (ventral aspect), showing (h) hypopygium and the hectors * above it. (X 15.)

side; penultimate joint of hind tarsi narrowly pale at extreme base. Wings faintly tinged with drab. Squamae waxen-white, with cream-coloured border and fringe (fringe of antisquama brownish). Halteres cream-coloured.

Glossina morsitans, form submorsitnns, Newst. Glossbaa submorsitans, Newstead, Annals of Tropical Medicine and Parasit- 3s and ology, Series T. M., Vol. IV, No. 3, p. 371 (December 20, 1910) ; of the Bulletin of Entomological Research, Vol. II, Part 1, p. 33, fig. 17, of the p. 34 (male genitalia) (April, 1911). ds are Relying chiefly on certain differences in t,he internal genital isible ; appendages of the male, Nr. Sewstead has recently described ;ion of this form as a species allied to t’he true G. morsitans, Westw. ; I of 8 but after making a minute external examination of the very ;hat, of large number of typical and other specimens of G. morsitans, Westw., received within the last few years by the British ora on Museum (Natural History) from many different localities in clove- Africa, and comparing with them the types of the supposed new small species, which have been kindly presented to the Kational - * See p. 14 (first paragraph end note *). 52 DISTRIBUTION OF GLOSSINA MORSITANS. BION

Collection by its author, I am satisfied that it is impossible to (Lake Ngan regard G. subm,orsitans, Newst., as specifically distinct from Zululand. P G. nlorsitans, Westw., and that it is in reality nothing more enable us to than a form or race of the latter. As will shortly be shown, and Protect, G. morsitans is the most widely distributed of all the Tsetse-flies, (besides OCCUI and it is, therefore, not surprising to find that even morphological found in : the characters are not absolutely identical in the case of specimens Territories), from localities situated at the opposite extremities of its area, Free State ( and separated from one another by some thousands of miles. the Anglo-E) The form submorsitans has hitherto been found in West Africa East Africa, and the Anglo-Egyptian Sudan, and it is of course possible that, and S.), and e.g. a male from Northern Nigeria belonging to this form might space, it is u be incapable of mating with a female of the typical race of dates of Capt1 G. morsitans from the Bechuanaland Protectorate. Still, if the received by 1 area of G. nlorsitans could be traversed from end to end, starting last eight yea from the Bechuanaland Protectorate or the North-Eastern Transvaal and proceeding to Northern Nigeria or the Anglo- Egyptian Sudan, it is not unreasonable to suppose that it would Until ael be found that the form submorsitans constitutes, as it were, one the only Tset, end of a chain of groups of interbreeding Tsetse-flies, of which and the very the other extremity would be represented by typical Gloss&a of elephant al morsitans, Westw. in South and As indicated in the description printed above, the dark bands Cumming ( 18 on the abdomen of G. morsitans are subject to much individual to this species variation as regards the extent to which they are developed. was originally The external characters mentioned by Mr. Newstead as serving applied, while to distinguish his G. submorsitacs from G. morsitans, Westw., used in its ori which are entirely derived from the abdominal bands, have been account of th carefully tested by examining the material in the National lies ready to Collection, and comparing it both with the types of the form year 1903 3 submorsitans and with typical specimens of G. morsitans, Westw., * In this SI with the result that in each case the character selected has G. morsitans, W sooner or later proved unreliable and ultimately invalid. intermediate fo (for the separa author, as poini from N. Nigeria DISTRIBUTIOX OF G. morsitans, Westw. Tsetse-flies hit1 examination ” t Glossina morsitans, Westw., once regarded as an exclusively Trop. Med. i~nc South African species, is unquestionably the most widely dis- December 20, 1E specimens of G. tributed of existing Tsetse-flies, since its range extends from be regarded as Senegambia (about 16” N.) in the north-west, to Southern Collection ~150 Territories, Gold Kordofan (about 12’ N.) and Southern Abyssinia in the north- characters, exe east, and thence southwards to the Bechuanaland Protectorate clove-brown onl, BIONOJIICS OF GLOSSINA MORSITANS. 53

(Lake Ngami district), the North-Eastern Transvaal, and Zululand. Although present knowledge is not sufficient to enable us to state whether G. morsitans exists in every country and Protectorate within the limits indicated, the species (besides occurring in the countries already mentioned) is certainly found in : the Gambia, French Guinea, the Gold Coast (Northern Territories), Togoland, Dahomey, Northern Nigeria,* the Congo Free State (Katanga District), the Bahr-el-Ghazal Province of the Anglo-Egyptian Sudan, the Uganda Protectorate, German East Africa, the Nyasaland Protectorate, Rhodesia (N.-E., N.-W. and S.), and Portuguese East Africa. Owing to limitations of space, it is unfortunately impossible to give details (localities, dates of capture, etc.) of the hundreds of specimens of this species received by the British Museum (Natural History) within the last eight years. BIOXOXICS.

Until well within the last decade, G. morsitans was almost the only Tsetse-fly of which the habits had ever been observed, and the very numerous references to ‘( Tsetse ” in the writings of elephant and other big game hunters, explorers, and pioneers in South and South Central Africa, from the time of R. Gordon Cumming (1850) down to the present day, refer almost exclusively to this species. It was to G. morsitaws that the name ‘( Tsetse ” was originally, and indeed until a few years ago exclusively applied, while even now the popular designation is still largely used in its ori,ginal and restricted sense. For the writing of an account of the habits of Glossiacc naomitans a mass of material lies ready to hand, and the statements available down to the year 1903 were duly summarised by the author in his

* In this survey no distinction is of course made between typical G. morsitans, Westw., and G. morsitans form submorsitans, Newst., or any intermediate forms or varieties. The types of G. submorsitans, Newst. (for the separation of which from G. mm&am, Westw., the present author, as pointed out above, is unable to find sufficient justification), are from N. Nigeria, and Newstead considers it probable that all West African Tsetse-flies hitherto regarded as G. nlorsitans will prove “on further examination” to be referable to his G. szLbmorsitans (cJ Newstead, Ann. lusively Trop. Med. and Parasitol., Series T. &I., Vol. IV., No. 3, pp. 369-370, December 20, 1910). While it is true that the malority of West African :ly dis- specimens of G. rnolaitans in the British Museum (Natural History) may !s from be regarded as belonging to the form submorsitans, Newst., the National juthern Collection also possesses specimens from Senegambia, the Northern Territories, Gold Coast, and N. Nigeria (Kontagora), which in all esternal north- characters, except for the fact that the last joint of the middle tarsi is ctorate clove-brown only at the distal extremity, are typical G. ??>orsitans, Westw. 54 BIO~OMIC!S O F GLOSSINA JIOXSITANS. BIONC

‘ Monograph.‘* In the present volume, however, the space Glossina morsi, available renders it out of the question to attempt a re’sunae’ of districts. Son the published observations on this subject ; while, in view of the time District fact that the Entomological Research Committee (Tropical Museum a nu Africa) is now endeavouring to collect and collate from reliable January, 190’ observers, resident on the spot, accurate statemen& concerning District, Nort the bionomics of G. nrorsitans,~ there is the less need for a to 5,500 ft. summary based largely on the result,s of necessarily limited Sharpe,* G. rnt observation, and referring in some cases to an epoch since passed The idea, away. A few points may, however, be noted. “appears to a Ss indicated by the term “ fly-belt,” originally applied to the vicinity oi G. morsitans, this species is usually confined to quite definite town or other tracts, often of very limited extent. Within these “ belts ” the villages in arei fly sometimes attacks human intruders in such numbers as to the fly.1 Thr have been compared to a swarm of bees. Cover, usually in the writing on ‘ sha,pe of large trees with thick undergrowth, open thickets, or Lundazi Distr scattered, shady trees is essential to the existence of G. morsitans, villages (Naw; which, like other species of Tsetse, is never found on the open, present time, c sun-scorched veld. Authorities like Mr. F. C. Selous, Sir John Kirk, or seven year and others (whose experiences, it should be pointed out, relate to Free State, G the period before the buffalo was driven northward, or almost Neave to ente exterminated by rinderpest), write of G. morsituns occurring in between the L swarms on the banks and “ along the water’s edge ” of the Glossina m Zambesi, Chobe, Rovuma and other rivers. Recent observers, on any large man the other hand, such as Dr. L. Sander (in German East Africa), to the special c Sir Alfred Sharpe (in Nyasaland), and Mr. S. A. Neave (on the cu$er), which sout#h-west shore of Lake Nyasa, and on the Luangwa River in view of the en, North-Eastern Rhodesia), lay stress on the fact that, in their Research Comr experience, G. morsitans evinces a dislike to, rather than a the question WI preference for, the immediate vicinity of water, whether river or that the availa lake, and is not infrequently met with at a considerable distance (half a mile or so) from any water.$ In this connection it is not * Bulletin of I 1910). uninteresting to remember that the original specimens of t See Austen, morsitans, found by Vardon and Oswell in 1845, were $ According 1 G. in the case of vi1 obtained on the Siloquana Hills in the Northern Transvaal, surrounding bus1 between the Magalaqueen or Nylstroom River and the Limpopo.§ 5 Bulletin of sketch-map (Octc * Cf. E. E. Austen, “A Monograph of t.he Tsetse-Flies,” pp. 6-23 (1903). I[ Cf. Sheffield t See Bulletin of Entomological Research, Vol. I, Part 2, pp. 147-149, April 25, 1908, p. and 152-153 (July, 1910). 7 Like G. bre f It may be noted that in Pola Province, Northern Nigeria, Dr. Dal&l blood of wild pi; has always found G. morsitam “ out of sight of water,” although near to Nyasaland Protec it (see p. 44). “literally in him 5 Cf. E. E. Austen, “A Monograph of the Tsetse-Flies,” pp. 7, 83-84 Journal, May 15, (1903). ** Vido szqwa ANS. BION6MICS OF GLOSSlNd MOBSITANS. 5.5

the space Glossina morsitanns, indeed, is by no means confined to low-lying a re’sunze’ of districts. Some three years ago, Mr. E. A. Copeman, at that view of the time District Commissioner at Kasempa, forwarded to the s (Tropical Museum a number of specimens of this species taken by him in Born reliable January, 1908, on the Congo-Zambesi watershed, Kasempa concerning District, North-Western Rhodesia, at an altitude of from 5,000 need for a to 5,500 ft. In Nyasaland, however, according to Sir Alfred ilg limited Sharpe,” G. morsitam is ‘( seldom found above 3,000 feet.” iince passed The idea, formerly generally entertained, that G. vnorsitans “ appears to avoid the presence of man, and is rarely found in applied to the vicinity of human habitations, or within the confines of a ite definite town or other settlement,” 7 is only partially true, since native belts ” the villages in areas in which this species occurs are often infested by lbers as to the fly.$ Thus Mr. P. E. Hall, Native Commissioner, Lundazi, ially in the writing on Cc The Movements of Glossina rnorsitans in the thickets, or Lundazi District, North-Eastern Rhodesia,” S speaks of native I. morsitans, villages (Nawalia and vicinity) which are “full of fly ” at the i the open, present time, or in which G. morsitans was “ a plague ” some six John Kirk, or seven years ago. In the Katanga District of the Congo t, relate to Free State, G. morsitnns has been observed by Dr. Sheffield or almost Neave to enter villages on a caravan-path infested with the fly, :curring in between the Lufira and Lualaba Rivers. (j ;e” of the Glossina morsitans feeds with avidity on the blood of almost ‘servers, on any large mammal that comes in its way.9 The old dispute as st Africa), to the special dependence of this species upon the buffalo (Bubahs Lve (on t,he ca$er), which has so often been alleged, still continues, and, in a River in view of the enquiry now being prosecuted by the Entomological t, in their Research Committee,** anything like a final pronouncement upon er than a the question would be premature. All that need here be said is 3r river or that the available evidence upon the point is to a certain extent le distance * Bulletin of Entomological Research, Vol. I, Part 3, p. 174 (October, m it is not 1910). cimens of t See Austen, “A Monograph of the Tsetse-Flies,” p. 23 (1903). 1 According to Sir Alfred Sharpe (Zoc. cit.), freedom from G. morsitu?ts 845, were in the case of villages situated in fly-areas is due to the clearing of the Transvaal, surrounding bush in order to make food-gardens. Limpopo.$ 5 Bulletin of Entomological Research, Vol. I, Part 3, pp. 183-184, and sketch-map (October, 1910). 6-23 (1903). I( Cf. Sheffield Neave, M.R.C.P., X.R.C.S., British Medical Journal, pp. 147-149, April 25, 1908, p. 988. T Like G. brevipalpis (see p. 94) the present species readily sucks the Dr. Dalziel blood of wild pigs; in Nyasaland, according to Dr. H. Hearsey, P.M.O., lgh near to Nyasaland Protectorate, G. morsitans was on one occasion seen to settle “literally in hundreds ” on the carcass of a wart hog (cf. British Medical pp. 7, 83-84 Journal, May 15, 1909, p. 1211). ** Vide sqwa, p. 54. 56 G. MORSITANS : PRELIMINARY STAGES. G. 1

contradictory. Whereas it is quite possible that formerly, in “I am sent Africa south of the Zumhesi, this particular Tsetse-fly was specially that of the con associated with and supported by the buffalo, since both were between the r found in the same places, and the buffalo in the region in question or seventy yar some five and thirty years ago far outnumbered all other game, escarpment in the evidence of many reliable observers shows that, in other parts east of the Hu of Africa at the present day, G. morsitans is in no way speci& ascertain the 1 dependent upon the buffalo or any other species of game. tree under whit Although by some observers stated to be most aggressive well populated during the hottest hours, according to other authorities Glossina tree was of an nzorsitans attacks human beings throughout the day, in addition Numerous pupz to biting, like other Tsetse-flies, during part of the night, present, but of especially when there is a bright moon and the weather is warm. nor was I able tion SufXicient ’ fly ’ country. REPRODUCTIOX AND PRELIMINARY STAGES. in that part is 7 Accurate observations on the reproduction and preliminary had fallen for s8 stages of Glossina nrwsitnns are urgently needed. That this may be commor should be so is not a little surprising when it is considered that kindly enclosed this Tsetse-fly may be said to have been a familiar insect ever and the exact s; since the species was described more than sixty years ago. In a As will be s letter published in “ The Field,” of September 28, 1907, Mr. R. 5 c, 5 F, 6 c, 6 L. Harger, late of North-Eastern Rhodesia, stated that he had than that of G often watched G. naorsitans depositing its “ eggs [i.e. larvae] in lips are also con the damp soil thrown up by the digging of a trench ” round his lacks the charac tent. The species of Tsetse used by Sir David Bruce in Zululand, notch between t in 1895-96, in the course of his epoch-making researches into the tumid lips are s etiology of Nagana, now proves to have been Glossina pallidipes, at present kno\ Austen, to which the larvae and pupae described and figured 5.4 mm. in lengt in the “ Further Report ” * by the distinguished investigator are much small referred to consequently belong. Harger’s brief statement there- p. 62), while the fore represents all that has so far been published on the life- intervening note history of Glossina morsitans. No larvae of this Tsetse-fly have as yet reached the Museum, but during the preparation of this AFFl volume the author received from Mr. R. W. Jack, Government Entomologist, Salisbury, Rhodesia, a pupa-case which undoubtedly Glossina mor belongs to G. morsitans (see Figs. 5 c and 6 c, pp. 5 and 7), Plate VI), and accompanied by the following letter, dated “ Department of the Glossina n&or Agriculture, Salisbury (Rhodesia), 20th October, 1910.” conspicuously b2 tarsi are pale. * “ Further Report on the Tsetse Fly Disease, or Nagana, in Zululend.” By Surgeon-Major David Bruce, 8.M.S. Ubombo, Zululand, 29th May, .sufficiently distil 1896 (London : Harrison and Sons. 1897). Pp. 2-3, Plates I and II. group by the abs BGES. G. MORSITANS : AFFINITIES, ETC. 57 formerly, in “ I am sending you an empty pupa-case, which I believe to be was specially that of the common Tsetse-fly (Glossinn morsitans). It was taken e both were between the roots of a tree (F&s sp. 1) on the banks of (sixty 3 in question or seventy yards from) a dry stream in the low veld below the other game, escarpment in the Lomagundi district, Southern Rhodesia, and I other parts east of the Hunyani River. I was at the time endeavouring to vay specinll~ ascertain the breeding grounds of the common Tsetse, and the *me. tree under which the pupa-case was found was in the middle of a t aggressive well populated belt of ‘ fly.’ The earth between the roots of the ;ies Glossina tree was of an easily-worked sandy nature, and rich in humus. ) in addition Numerous pupa-cases of other Diptera and of Lepidoptera were the night, present, but of living Dipterous pupae I was able to find none, ler is warm. nor was I able to discover any more cases, in a state of preserva- tion sufficient for recognition, during twelve days’ work in the ‘ fly ’ country. I may mention that the season on the low veld in that part is very late, and when I was there (October) no rain preliminary had fallen for several months. It is possible that living pupae That this may be common after the first rains.” With his letter Mr. Jack sidered that kindly enclosed a photograph showing the roots of the Ficus tree, insect ever and the exact spot where the pupa-case was found. ago. In a As will be seen from the photographs on pages 5 and ‘i (Figs. )OT, Mr. R. 5 c, 5 F, 6 c, 6 F), the pupa-case of Glossina morsitans is smaller hat he had than that of G. palpalis and neater in appearance; the tumid . larvae] in lips are also conspicuously smaller, and the notch between them ” round his lacks the characteristic 7cey-hole-like constriction exhibited by the n Zululand, notch between the lips of the puparium of the latter species. The ies into the tumid lips are smaller than those of any other Glossina-puparium G pallidipes, at present known. The specimen found by Mr. Jack measures tnd figured 5.4 mm. in length, by 3 mm. in greatest breadth; these dimensions nvestigator are much smaller than those of the pupa of G. pallidipes (see nent there- p. 62), while the size and shape of the tumid lips and the size of the m the life- intervening notch are very different (cp. Figs. 6 c and 6 D, p. 7). he-fly have ;ion of this AFFINITIES AND DISTINCTIVE CHARACTERS. .overnment odoubtedly Glossina morsiians, Westw., with G. pallidipes, Austen (p. 58, 5 and i), Plate VI), and G. longipalpis, Wed. (p. 63, Plate VII), forms trtment of the Glossina morsitans group of species, in which the abdomen is I.” ,conspicuously banded, while the first three joints of the hind tarsi are pale. Apart from all other characters, G. morsitan,s is L Zululmd.” 29th May, sufficiently distinguished from both the other members of its bnd II. group by the absence of a distinct fringe of tine hair on the front 58 DESCRIPTION OF GLOSSINA PALLIDIPES. DISTRIEl margin of the third joint of the antennae, and the greater depth extremity of t of the pale hind margins to the banded abdominal segments. and tapering, Further distinctive characters from G. pallidipes are pointed out branched hairs on page 63. palpi cream-b{ greyish-brown clove-brown, ri Glossina pallidipes, Austen. distal extremil (Plate VI.) Thorax.-1 Glossina pallidipes, Austen, “A Monograph of the Tsetse-Flies,” p. 87, in 6, paler (E Plate IV (1903). buff, upper so triangular in c 6, $! .-Length, 8 (11 specimensj 8.5 to 10.4 mm., ? (9 speci- bristles long ir mens) 9.75 to 11.25 mm. ; width of head, 8 just under 3 to Abdomen.- 3.2 mm., ? 3 to 3.25 mm. ; width of front at vertex, & 0.5 to, blotch on ea( 0.6 mm., ? 6.S to 1 mm. ; length of wing, 8 S.6 to 9.4 mm., extending to 1 0 9.5 to 10.25 mm. sixth segments Medium-sized or rather large species, clcsely resembling the band, starting Test African 0. longipalpis, H&d. (p. 63, Plate VII), but di s t i n- thus confining guishable owing to the coloration, of the last two joints of the front space, to a ve and middle tarsi.-Front in & very narrow; dorsum yf thorax band, when fl olive-grey, smoke-grey, or drab-grey, with the usual dark brown, interruption, b longitudinal ntarkings generally nluck reduced, broken up, rtnd but bands on the t little conspicuous ; abdomen bu$, dorsum with dark brown or clove- posterior angle brown bands a8 shown in Plate VI ; legs huff, last two joints oj- kind coloured ; seve tarsi clove-brown, front and middle tarsi either entirely pale or, at of 6 ochraceol most, last iwo joints of front tarsi faintly brownish at fhe ti$s, and Legs.--Inn last joint and distal half of penultimate joint oj’ middle tarsi light surfaces of mi brown-never so dark as to form a sharp contrast with the remaining more or less @ joints. mouse-grey stri Head cream-buff, face and jowls bright cream-coloured pol- hind femora ea linose, posterior surface light grey, ocellar triangle smoke-grey or under side nea drab-grey, sides of front (parafrontals) each with a dark brown TVings ligl elongate blotch, sometimes indistinct, a little below half way white, with crf between level of ocellar spot and that of base of antennae ;. frontal stripe raw-sienna-coloured, upper fourth mummy-brown or D brownish ; ocellar spot slate-coloured, connected with clore-brown The area of transverse band joining bases of vertical bristles; first and representative second joints of antennue (cf. Fig. 8, p. 12) dark brown, third Plate VII), inc joint light or dark mouse-grey, more or less buff at base, elongate African contin and with a prominent distal extremity, front margin with a northern boun conspicuous fringe of tine hair of moderate length, arista cream- at present kr buff or buff, base of penultimate and lower margin of proximal ‘IDIPES. DISTRIBUTION OF GLOSSINA PSLLIDIPES. 59

greater depth :xtremity of terminal joint dark brown, terminal joint elongate inal segments. and tapering, with a prominent ridge on upper side at base, ne pointed out branched hairs numerous, twenty-two to twenty-eight in number ; pal+ cream-buff, yellowish smoke-grey above, slate-grey or dark greyish-brown at tips ; proboscis bulb dark mummy-brown or clove-brown, raw-umber-coloured on each side, except as a rule at distal extremity. Thorax.-Pleurae and pectus grey, mouse-grey, or dark grey se-Flies,” p. 87, in 8) paler (smoke-grey or drab-grey) in $’ ; scutellum cream- buff, upper surface with a well marked dark brown blotch, triangular in outline, on each side of middle line ; apical scutellar Im., 0 (9 speci- bristles long in both sexes. It under 3 to Abdomen.-Dorsum of second segment with a dark brown rtex, 6 0.5 to. j to 9.4 mm., blotch on each side (sometimes indistinct or wanting), not extending to lateral or hind margins ; dark bands on third to sixth segments inclusive cery deep (i.e. on each segment the dark resenlbling the band, starting from front border, extends close to hind margin, II), bzcf distin- thus confining the ground colour, apart from the median inter- nts of the front space, to a very narrow hind border) ; each half of each dark vlln1 qf’ thorax band, when fully developed, is cut ?ff squarely at the median I da& brown, interruption, but in many specimens the inner extremities of the m np, md bnt bands on the third and fourth segments are partially obliCerated ; brown or clave- posterior angles of second to sixth segments inclusive cream- P join fs of hind coloured ; seventh segment smoke-grey or drab-grey ; hypopygium *ely pole or, at of 8 ochraceous-buff, broadly oval in outline as seen from below. ct the tills, und Legs.-Inner surfaces of front femora, and usually also outer !dle tctrsi light surfaces of middle and hind femora towards distal extremities 1 the wmnining more or less grey or greyish; middle femora sometimes with a . ’ ” outer side, and middle and -coloured pol- hind femora each with the usual elongate dark brown mark on smoke-grey or under side near distal extremity. L dark brown Win~/s light drab-coloured, unicolorous. Squaniae waxen low half way white, with cream-coloured border. Halteres cream-coloured. of antennae ;. nmy-brown or DISTRIBUTION OF G. pallidipes, Austen. h clove-brown The area of distribution of G. pallidipes, which is the eastern es; first and representative of the West African G. longipalpis, Wied. (p. 6.3, brown, third Plate VII), includes the south-eastern and eastern portions of the base, elongate African continent, and extends from Zululand at any rate to the largin with a northern boundary of the East Africa Protectorate. So far as arista cream- at present known, in addition to the countries mentioned, z of proximal BIONO 4 60 DISTRIBUTION OF GLOSSINd YALLIDIPES.

Pope-Hennesay G. pallidipes occurs in Portuguese East Africa, the Nyasaland 1 ? , Kilindi, 3 Protectorate, North-Eastern Rhodesia, German East Africa, and The occurr the Uganda Protectorate. Portuguese Et The particulars with reference to the specimens of this species states, on the at present contained in the British Museum (Natural History) question G. 2: are as follows :- naorsitans.* II Zululand : 1 6 , 3 ? 0 ,* precise locality unknown, 1910 is said to be r (forwarded by Sir C. J. R. Saunders, K.C.M.G. ; presented by G. pallidipea h; Colonel Sir David Bruce, C.B., F.R.S., A.M.S.); 1 d, 1 0, 5,000 feet.7 I Mtunzini, 1910 (presented by Colonel Sir David Bruce). Portu- writing from F guese East Africa : 3 $ $, 2 9 0, Maganja da Costa, present species Quelimane, February, 1911 (Dr. J. F. Sant’ Anna--forwarded several 1Ocalitie for determination by the Director, Sleeping Sickness Bureau). Nyasaland Protectorate : 1 $ , Shire Highlands, caught in a locality where Tsetse-flies were supposed to be non-existent, 1893 (Sir H. H. Johnston, G.C.M.G., K.C.B.) ; 2 8 $ , precise locality The late D 24th, 1906, wit uncertain, but probably Lunyina River, Henga, 1895 (Captain R. Crawshay). North-Eastern Rhodesia : 1 6, Lake Mweru, collected by hi 18-22. iii. 1892 (Captain R. Craloshay). German East Africa: said : “ Of thir 1 6, 3 Q 0 (including the types of the species), Kilima Njaro, females. The f forest at Kiber 1888 (F. J. Jackson, C.B., CXG.). Uganda Protectorate : 1 8, 2 0 0 , Busoga, 1903, and 1 $ , 1 0, precise locality unknown, They were num scattered trees, June, 1909 (Colonel Sir David Bruce, C.B., F.R.S., A.M.,%‘); 4 8 6, 3 0 ? , Kibero, Nile Province, between Nimule and They attacked 1 caught with the Wadelai, June, 1906 (the late Dr. W. A. Densham). East Africa Protectorate : 1 $’ , Upper Sabaki River, 1890, and 18, ‘I caught blue service put in Witu town,” 1891 (presented by the late Eritish East Africa path after enter of the belt, whi Company) ; 1 $? , Witu Forest,, 1895 (the Sultan of Wtu; pre- flies were seen u. sented by the late W. S. Godfrey) ; 18 & 6, 2 $! 9, Machakos, arrival two were 1897 (received from the Tsetse-fly Committee of the Royal me. On return Society) ; 3 6 6, 1 0, Uganda Railway, 1897-1898 (the late u$il nearing th Vet.-Captain A. J. Hasianz) ; 1 ? , Uganda Railway (C. X. above may all 1 Bettotr) ; 1 &, 2 ,$ 8, Tsavo River, 1902 (received from Colonel much further ths Sir David Bruce, C.B., F.R.S., A.M.S.) ; 35 &, 20 0, Kibwezi, buffalo in the for Simba, and near Nairobi, (( in railway carriage ” (Dr. Mofat, are no natives n per Colonel Sir David Bruce) ; 11 3 6, 9 ? ? , Kibwezi, 1903 connects two vill; (received from the Sleeping Sickness Commission, per Dr. D. Nabarro) ; 6 & 8, Gosha, Jubaland, October, 1904, “ said by the * Cf. A. Laverr Sciences, t. cxliv, p (Bt. Mnjor L. H. R. natives to be deadly to cattle and camels” littoral region of PO of the species near G * On0 of these is grk3vid and contains an apparently full-grown larva, t Cj. Sleeping the tumid lips of which are visible through the delicate integument of (January 10, 1910). the posterior extremity of the fly. BIONOMICS OF GLOSSINA PALLIDIPES. 61 LIDIPES. Pope-Hennessy, D.S.O.) ; I 8, Voi, 19. iii. 1906 (W. L. Sclnter); ;he Nyasaland 1 0, Kilindi, January, 1909 (received from R. J. Stordy). Ist Africa, and The occurrence of G. pallidipes in the Mozambique district of Portuguese East Africa is recorded by Professor Laveran, who of this species states, on the authority of M. G. Vasse, that in the region in tural History} question G. pallidipes is found at higher altitudes than G. morsitans.* In the East Africa Protectorate, however, where it nknown, 19 10 is said to be more widely distributed than any of its congeners, presented by G. pallidipes has not been observed at an altitude of more than * 18, 1 0, 5,000 feet.t In the Uganda Protectorate, Dr. R. van Someren, iuce). Portu- writing from Fvt Portal on January lOth, 1911, states that the da da Costa, present species occurs, in company with G. morsitans, We&w., in au--forwarded several localities in Toro. ness Bureau). , caught in a BIONOMICS. existent, 1893 The late Dr. W. A. Densham, writing from Nimule on July lrecise locality 2&h, 1906, with reference to Glossina pallidipes as observed and 1895 (Captain collected by him in the Nile Province, Uganda Protectorate, Lake Mweru, said : “ Of thirteen specimens taken, nine were males and four East Africa : females. The flies were found near and in a narrow belt of true Kilima Njaro, forest at Kibero, between Nimule and Wadelai, in June, 1906. ctorate: 1 3, They were numerous along the native path, in long grass with Iity unknown, scatter&d trees, for a quarter of a mile before reaching the forest. 2.S., A.M.,??.‘) ; They attacked freely at 8.30 on a sunny morning, were easily L Nimule and caught with the hand, and were observed to bite through a worn East Africa blue service puttee. They were seen and caught along the native ‘( caught 16, path after entering the forest, and for two miles the other side h East Africa of the belt, which is only about 300 yards in width, occasional )f Witu ; pre- flies were seen until the village was reached. Half-an-hour after 0, Machakos, arrival two were captured by my boys in camp and brought to )f the Royal me. On returning from the village next day no flies were seen 1898 (the late until nearing the forest, SO that the ‘ occasional ’ flies mentioned ilway (C. X. above may all have been ‘ following ’ flies. If so, they follow from Colonel much further than G. palpalis. There are many rhino, pig, and ! 0 , Kibwezi, buffalo in the forest, and Colobus monkeys‘ are numerous. There (Dr. Moffcd, are no natives nearer than two miles, but a well-trodden path iibwezi, 1903 connects two villages, and passes through the Fly. The natives , per Dr. D. Lc said by the * Cj. A. Laveran, Comptes rendus des s&mces de l’Aca&mie des Sciences, t. cxliv, p. 551 (1907). That G. pallidipes also occurs in the ajar 1;. H. R. littoral region of Portuguese East Africa is proved by the recent discovery of the species near Quelimsne, by Dr. J. F. Sant’ Anna (ui& supra). ill-grown larva, t Cf. Sleeping Sickness Bureau Bulletin, Vol. II, No. 13, p. 37 integument of (January 10, 1910). 62 YUPA OF GLOSSINA PALLIDIPES. DESC:

know of the fly, but do not consider it dangerous to animals. abdominal They keep sheep and goats, and drive them along this path. No of the frc cattle are kept for many miles round.” printed in In the East Africa Protectorate, where, according to the narrow Dr. P. H. Ross, the species is most numerous in August and be separat September, Dr. A. D. Milne, P.M.O., states t’hat G. pallidipes, of the ant in company with G. hevipalpis and longipennis, attracted by with the d lamplight frequently enters railway carriages when a train is conspicuou waiting during the night at one of the stations in the fly-belt on of the antt the Uganda Railway, and thus may be carried for a distance of again as ir 150 miles or more.* blunter an

PRELIMINARY STAGES. shorter in From 6 A comparison of the pupae obtained by Colonel Sir David distinctly 1 Bruce in Zululand, and described and figured by the author in absence of his I‘ Monograph of the Tsetse-Flies,” pp. 26-28, Fig. 7 (1803 j, with front and the pupa-case of Glossina morsitans, Westw., recently discovered front and n by Mr. R. W. Jack at Lomagundi, Southern Rhodesia (see p. 57, distinguish1 and compare Figs. 6 D and 6 C, p. 7), conclusively shows that there distinctly n can be no doubt that the former belong to G. 2)nllidil,es,.Austen. margins of The pupa of G. pallidipes then (Figs. 4, 5 D, 6 nj, as represented the front is by Sir David Bruce’s examples (four specimens), is from 6.4 to 7 mm. in length, by 3.4 to 3.75 mm. in greatest width. The tumid lips, which are from 0.8 to 1 mm. long, and separated by a U-shaped notch (somewhat resemblin,r but with straighter sides and fuller at the bottom than the notch in the case of the pupa of G. fusca, Walk.), are much larger than the lips of the pupa of G. morsitans, while the notch between them is about twice as Glossina lor wide as in the latter. The notch is quite different in shape from Insekten, Maoquart that exhibited by the pupa of G. brecipalpis, Newst. (compare Exot,., T. Figs. 6 D and 6 A), and the inner edges of the distal portions of Kolonialp the lips are more rounded and less sharp than in Dhe case of the figs. 1,9, pupa of either G. brevipalpis or G. f~sca. p. 90, Pla

it, o.-- AFFINITIES AND DISTINCT~E CHARACTERS. mens) 9 to 1 + Among the species of the Glossina lllorsitccns group to which * To f&Ii it belongs, G. pallidipes is distinguished from G. morsitans, needle or the We&w., inter alin by the presence of a distinct fringe of fine under a lens. t Mr. N+ewr hair on the front margin of the third joint of the antennae, Austen, is me, by the narrowness of the p%le hind margins to the banded of Tropical MI -December 2~ * Cj. Sleeping Sickness Bureau Bulletin, Vol. II, No. 13, p. 37 specimen used (January 10,191O). study by Mr. N ES. DESCRIPTION OF GLOSSINA LONGIPALPIS. 6:: $ ; /II ’ I 8 to animals. abdominal segments, and by the coloration of the last two joints lis path. NO of the front and middle tarsi, as described in the diagnosis printed in italics above. In the male sex it is distinguished by according to the narrower front, and in both sexes G. pallidipes may easily August and be separated from G. morsitans by the shape of the third joint G. pallidipes, of the antennae, which in the former species is long and narrow, attracted by with the distal extremity pointed and (especially in the female) en a train is conspicuously turned forwards; in 8. morsitans the third joint he fly-belt on of the antennae is shorter and much broader (about half as broad a distance of again as in G. pallidipes), and darker towards the tip, which is blunter and not so much turned forwards; the arista is also shorter in the other species mentioned than in G. pallidipes.* From G. longipalpis, Wied., G. pallidipes, which is as a rule a lel Sir David distinctly larger species, is distinguishable in both sexes by the ;he author in absence of sharply defined dark tips to the last two joints of the (1903), with front and middle tarsi; as regards the males, even though the ly discovered front and middle tarsi be wanting, a male of G. pallidipes can be iia (see p. 57, distinguished from a male of G. longipalpis by the front being NS that there distinctly narrower at the vertex than anteriorly, since the inner il)~s, Austen. margins of the eyes converge above, whereas in G. longipalpis represented the front is of the same width or practically so throughout.7 from 6.4 to width. The parated by a aighter sides Glossina longipalpis, Wied. ; of the pupa (Plate VII.) the pupa of :ii mt twice as Glossina longipalpis, Wiedemann, “ BussereuropLische zweifliigelige t Insekten,” Zweiter Theil, p. 254, Taf. ix, figs. 10, a, b, c (1830); 1 shape from Macquart, Hist. Nat. des Ins. Dipteres, T. ii, p. 245 (1835), and Dipt. st. (compare Exot., T. ii, 3, p. 113, Tab. 14, fig. 1 (1843) ; Lichtwardt, Beitrage zur I portions of Kolonialpolitik und Kolonialwirtschaft, iv Jahrg., Heft 9, p. 264, et seq., e case of the figs. 1,9, 10, etc. (1903) ; Austen, “ A Monograph of the Tsetse-Flies,” p. 90, Plate V (1903). &, 0 .--Lengt,h, 8 (10 specimens) 8.4 to 9 mm.? 0 (6 speci- ii 3. width of head in both sexes 2.75 to 3 mm. ; mens) 9 to 10 mm.; :; ‘up to which * To facilitate comparison, the antennae should be removed pith a !. morsitans, needle or the point of a fine scalpel, mounted in glycerine and examined inge of fine under a lens. t Mr. Newstead’s recently published conclusion that Glossi~ pallidipes, 3 antennae, Austen, is merely a “colour variety” of G. longipal~is, Wed. (cf. Annals the banded of Tropical Medicine and Parasitology, Series TM., Vol. IV, No. 3, p. 371 -December 20, 1910), has since proved to be erroneous, and due to the lo. 13, p. 37 specimen used for examination having been incorrectly determined. The study by Mr. Newstead of further material submitted to him by the author F 64 Dl3SCRII’TION OF GLOSSLNA LONGIPALPIS. DISTRIB width of front at vertex, 8 0.6 mm., 9 0.75 mm. ; length of palpis is foul wing, 8 8 to 8.6 mm., ? 8.75 to 9.25 mm. district of tl Belgian Con Precisely similar to Glossina pallidipes, Austen (p. 58, Plate VI), Museum colic in general appearance, but distinguished by the last two joinfs of may be trust the front and middle tarsi having sharply dejined and conspicuous Kirk, G.C.M clove-brown tips, and in the male sex by the front not being distinctly Glossina pall narrower at the vertex. possession of Head cream-buff, face and jowls bright cream-coloured to a recently pollinose, posterior surface smoke-grey, ocellar triangle smoke- all the Tsetse- grey or drab-grey, sides of front (parafrontals) each with a dark ago as 1864, brown blotch, as in G. pallidipes ; frontal stripe tawny ochraceous Tete and the or ochraceous anteriorly, mummy-brown posteriorly ; ocellar spot has been mad and band joining bases of vertical bristles as in G. pallidipes ; follows that ( first and second joints of antennae buff, dark brown on inner since we knc side, third joint mouse-g&y, buff at extreme base, front margin has disappear fringed as in G. pallidipes, arista coloured as in G. pallidipes, abounded. A but terminal joint without so prominent a ridge on upper side solitary specin at base and consequently not so tapering, branched hairs twenty- or otherwise, one to twenty-three in number; palpi as in G. pnllidipes; Congo Free f proboscis bulb dark sepia. contrary is r Thoraz.-Dorsum olive-grey or dark olive-grey, with markings l forming the so as in G. pallidipes ; pleurae, pectus and scutellum as in G. as is at presen pallidipes ; apical scutellar bristles long in both sexes. of Glosvina lc Abdomen.-Coloration and markings precisely the same as in African specie G. pallidipes, inner extremities of interrupted bands on third and district of the following segments often more or less obliterated. The partici Legs.-Coloration, except in case of last two joints of front palpis (other tl and middle tarsi, the same as in G. pallidipes. Museum (Natu Wings, quamae, and halteres as in G. pallidipes. 4 Q Q, Dodow the break of t

DISTRIBUTION OF G. longipnlpis, Wied. River, March, 3 Q Q, Akwat Although essentially a West African species, the range of (Dr. G. J. Pir wGch extends to the north-west as far as Senegal,“> Glossina longi- Jebba, October, R. Niger, Nupe shows that males of 0. pallidi~es and longi$yis are readily distinguish- Southern Niger able by means of their genital appendages (cf. Bulletin of Entomological * G. longipalpi Research, Vol. II, No. 1, pp. 30-32, Fig. 15, April, 1911). and Todd (Annals * For records of the occurrence of G. longi@@s in Senegal, see Vol. I, No. 1, p. 72 Laveran, Cemptes Rendus des SBances de l’ilcadbmie des Sciences, Laveran, Compte T. cxliv, p. 547 (1907). For records from French Guinea and French T. cxxxix, p. 662 (I Congo, see Laveran, ibid., T. cxxxix, p. 659 (1904), and T. cxli, p. 931 t E. E. Austen (1905), respectively. $ Cj. Sleeping 1 GIPALPIS. DISTRIBUTION OF GLOSSINA LONGIPALPIS. 65 urn. ; length of palpis is found at least as far to the south-east as the Katanga district of the Congo Free State,* i.e. the south-eastern corner of Belgian Congo. As stated in the author’s “ Monograph,“? the p. 58, Plate VI), Museum collection contains a male of this species, ivhich, if its label st two joinfs oj- may be trusted, was collected by Dr. John Kirk (now Sir John and conspicuou,s Kirk, G.C.M.G., K.C.B., F.R.S.) on the Zambesi. Specimens of ! being distinctly Glossina palpalis, Rob-Desv., and G. brevipalpis, New&., in the possession of the Museum, also bear similar labels, and according to a recently published extract from a letter from Sir John Kirk $ cream-coloured all the Tsetse-fliesbrought home by him, whichwere collected so long triangle smoke- ago as 1864, were obtained in the valley of the Zambesi, between :h with a dark Tete and the Victoria Falls. Even assuming that no mistake wny ochraceous has been made in the labelling of this specimen, it by no means ly ; ocellar spot follows that G. longipalpis is still to be found on the Zambesi, L 0. pallidipes ; since we know, for instance, that Glossina naorsitans, Wesew., )rown on inner has disappeared from the Victoria Falls, where at one time it 3, front margin abounded. Apart from statements relating to Sir John Kirk’s 1 G. pallidipes, solitary specimen, there are no records or other evidence, recent I on upper side or otherwise, of the occurrence of G. longipalpis south of the jd hairs twenty- Congo Free State ; therefore, until indisputable proof to the G. pallidipes ; contrary is received, it will be safer to regard Katanga as forming the southern limit of the range of 0. longipalpis, so far , with markings as is at present known. The authenticated area of distribution dlum as in G. of Glossina longipalpis, which, as already stated, is a West ?XfX. African species, extends then from Senegal to the Katanga the same as in district of the Congo Free State. ds on third and The particulars with reference to the specimens of G. longi- palpis (other than Sir John Kirk’s example) now in the British joints of front Museum (Natural History) are as follows :-Gold Coast : 2 & 6, 4 0 0, Dodowah (30 miles from Accra), <‘very numerous after es. the break of the rains,” May, 1906 (6’. A. L&g) ; 1 6, Volta River, March, 1908 (G. C. Dudgeo%). Northern Nigeria : 1 d, ed. 3 $) ? , Akwatcha, Bassa Province, July and August, 1906 s, the range of (Dr. G. J. Pirie, W. A.M.S.) ; R. Niger, Ilorin Province, near +% Glossinn longi- Jebba, October, 1909 (Dr. C. W. Mcleay, W.A.M.S.); 1 $,&J-O, R. Niger, Nupe Province, 1909 (Dr. E. A. Chartres, W.A.M.S.). eadily distinguish- Southern Nigeria : 5 8 6, 2 ? 9, Igbassa, Ijami, between Ijami of Entomological * G. Zong~~alpis is recorded from the Upper Luapula River by Newstead and Todd (Annals of Tropical Medicine and Parasitology, Series T. AI., is in Senegal, see Vol. I, No. 1, p. 72 (February 1. 1907)) ; also from Lukafu, July, 1903, by nie des Sciences, Laveran, Comptes Rendus des SQances de l’Acad8mie des Sciences, uinea and French T. cxxxix, p. 662 (1904), and T. cxli, p. 932 (1905). ,nd T. csli, p. 931 t E. E. Austen, “ A Monograph of the Tsetse-Flies,” p. 92 (1903). $ Cf. Sleeping Sickness Bureau Bulletm, No. 3, p. 125 (January, 1909). F2 66 EIONOMICS O F c:T,OSSlK-,4 LONGIPALI’IS. G. LO1

and Meko, and between Meko and Igbassa (all localities in In Central Western Province), April, 1908 (G. C. Dudgeon) ; 1 8, 1 0, show a well-1 Sebe, 1 6, between Olebakin and Tfon, May, 1908 (G. C. Dudgeon). occurs in the T It will have’ been noticed that the Museum has as yet are found on15 received no specimens of G. longipalpis from Sierra Leone, where of the forest 1 the type of the species (and therefore also of the genus Glossina) in open clearin was obtained by the Swedish botanist Afzelius in the closing states that in years of the eighteenth century. rains, but set season, especia restricted thal Like other members of the Glossina morsitans group, of which are a to which it belongs, G. longipalpis would appear not to be so its oongeners 1 closely restricted to the immediate vicinity of water as the Like G. palpc species belonging to the Glossilaa palpalis group often, although according to not invariably, are. A field-note attached by Mr. G. C. Dudgeon mammals, and to a 8 and 9 taken by him near Ijami, Southern Nigeria, in present specie April, 1908, as mentioned above, states that they were caught in hippopotamus ; bush, where there was no water and also no game, with the In the Wet exception of small antelope; at Sebe, Southern Nigeria, where G. longipalpis Mr. Dudgeon found G. longipalpis in May, 1908, game was repre- found in the fc sented by the West African buffalo (Bubalus nanuu). Writing longipalpis to the author on April Sth, 1907; with reference to G. REI as found by him in Northern Nigeria, Dr. G. J. Pirie said : “ During the dry season at Akwatcha (twenty-five miles inland Glossina lc from the River Benue), Bassa Province,. Glossina Zongipalpis are Roubaud to coj verv few, while during the rains they are rather numerous, referred to at especially in the ‘desner bush-patches along the small water- about ten days courses.” Dr. Pirie added that “ trypanosomiasis both of animals of from 24” to twenty-six to t (h or,3 es and dogs) and of man exists at Akwatcha, especially in the rainy season from May to October,” and that he had searched carefully for Glosaina palpcrlis, but, so far as he could AFE judge, without success. At a certain spot on the O&m6 River, in Dahomey, where In the (71 both G. lotc!/ipalpis and palpcrlis occur, the latter species, accord- stated, G. lon ing to Roubaud, is for the most part confined to the immediate distinguished f vicinity of the water and to the belt of forest fringing the banks ; the same charac G. lou!gi’,alpis, on the other hand, which is scarcely to be found Austen, from in the actual palpalis zone, predominates in the bush bounding * Cf. E. Rou the forest on the outer side, and is there preyed upon by a D&homey” : Con Hymenopteran belonging to a new species of the genus Bmlbpr.i” T. 152, No. 7, p. 4 t Cj. Sleepiq * C-f. E. Roubaud, Comptes Rendus des SBances de 1’AcadGmie des (March 14, 1911). Scicnccs, T. 151, No. 8, p. 506 (August 22, 1910). $ Roubaud, lo PALI’I S . G. LONGIPdLPIS : REPROl~UCTION, ETC. 67

.l localities in In Central Dahomey, where, according to Roubaud, the sexes b); l&,1?, show a well-marked separation or localigation, G. lon&xlpi~ 7. C. Dudgem). occurs in the vicinity of streams and large rivers. “ The males II has as yet are found only in the clumps of brushwood along the inner edge t Leone, where of the forest belt near streams ; the females are to be met with genus Glossina) in open clearings where there are acacias and mimosas.” Roubaud in the closing states that in Dahomey G. longipalpis is abundant during the rains, but seems to disappear almost completely in the dry season, especially after the bush is burnt; its habitat is more restricted than that of either G. tachinoides or G. pcl&is, both r&tans group, of which are also found in the same district, and it mingles with not to be so its congeners less than does either of the two species mentioned. water as the Like G. palpalis and tachinoides, G. longipalpis in Dahomey, sften, although according to Roubaud, lives chiefly at the expense of wild G. C. Dudgeon mammals, and accompanies big game in its movements ; the rn Nigeria, in present species especially frecprents paths newly trodden by vere caught in hippopotamus and elephant.* ame, with the In the Western Province of Ashanti, according to Kinghorn, Nigeria, where G. longipalpis “is essentially an open country fly and is not me was repre- found in the forest belt.” t UY). Writing

G. longipalpis REPRODUCTION AND PRELIMINARY STAGES. J. Pirie said : I miles inland Glossinn lorqiya@iu in Dahomey was never observed by longipalpis are Roubaud to copulate in captivity. Females kept by the observer er numerous, referred to at 25” C. (‘ii” F.) deposited a larva at intervals of small water- about ten days, and the pupal stage, at an average temperature ,th of animals of from 24” to %Y C. (75.2’ to 77” F.), was found to last from ha, especially twenty-six to thirty-five days.$ that he had as he could AFFINITIES AND DISTIBCTIVR CHARACTERS.

homey, where In the Glossincc mo mi f n n s group, to which, as already lecies, accord- stated, G. longipalpiu belongs, the present species can be he immediate distinguished from G. mwsitanS, Westw., inter din by some of lg the banks ; the same characters as those serving to distinguish G. pallitli~~ea, y to be found Austen, from the species mentioned’ (see p. 6J), i.r. by the rsh bounding * Cf. E. Roubaud, “!&udes hiolo@ques sur les Glossines du moyen 1 up011 by a Dahomey ” : Comptes Rendus des SBances de l’bcadbmie des Sciences, nus &1 t r b e s . " T. 152, No. 7, p. 406 (FBvrier 13, 1911). t Cf. Sleeping Sickness Bureau Bulletin, Vol. 3, No. 25, p. 136 1’Acadtmie des @larch 14, 1911). $ Roubaud, lot. cit., p. 407. 68 G. LONGIPALPIS : DISTINCTIVE CHAIXACTERS. presence of a distinct fringe of fine hair on the front margin of the third joint & the antennae, and by the narrowness of the pale hind margins to the banded abdominal segments. The similarity in general appearance between Glossina longipalpis and its eastern representative G. pallidipes has been referred to in the diagnosis printed in italics above, where the external characters by which the two species may be distinguished are also pointed out ; it may be added that Glossina 1ongipalpi.s js usually a smaller insect than G. pallidipes, and that the specific 1. Third hair differences presented by the male genital appendages have quite friq recently been elucidated by Mr. Newstead. undc mete in p Third, hair scan mw head fring of w slen~ 2. Longes third to f em3 joint Longer third to frc of th slend 3. Pleurae coxac Pleurae

Stomoxys fuse p. 682 (1849 Glossina fusca Austen, MO Newstead, 1 Vol. IV, No

6, ?.--I (24 speciment IHARACTERS. 69

he front margin the narrowness minal segments. twina longipalpis CHAPTER VI. Een referred to re the external THE GLOSSINA FUSCA GROUP. istinguished are SYNOPSIS OF SPECIES. ta longipalpis i s hat the specific 1. Third joint of antennae fringed with fine tges have quite hair on anterior and posterior margins ; fringe on anterior margin conspicuous under a hand-lens magnifying 15 dia- meters (nominal), when head is viewed in profile ...... 2 Third joint of antennae with fringe of fine hair on anterior margin so short as to be scarcely noticeable under a hand-lens magnifying 15 diameters (nominal), when head is viewed in profile (longest hairs in fringe in length not exceeding one-sixth of width of third joint) ; palpi long and slender ...... 3 2. Longest hairs in fringe on front margin of third joint of antennae, in length equal to from one-fourth to one-third (not exceeding one-third) of width of third joint ; palpi of moderate length ...... taban~~ofor&s, Westw. Longest hairs in fringe on front margin of third joint of antennae in length equal to from one-half to three-fourths of width of third joint ; palpi noticeably long and slender . , ...... nigrofusca, Newst. 3. Pleurae drab-grey or isabella-coloured, hind coxae buff or greyish-buff ...... fusca, Walk. Pleurae dark grey, hind coxae mouse-grey. fuscipZe&s, sp. nov.

Glossina fusca, Walk. (Plate VIII.) Stomoxys fuscus, Walker, List Dipt. Ins. in COB. Brit. Mus., Part III, p. 682 (1849). Glossina fusca, Walker, “Entomologist,” Vol. VI., p. 328 (1873) (net Austen, Monograph of the Tsetse-Flies, pp. 95-97, Plate VI (1903)) ; Newstead, Annals of Tropical Medicine and Parasitology, Series T. M., Vol. IV, No. 3, pp. 370, 373 (December 20, 1910). &, 0 .-Length, 6 (12 specimens) 9.6 to 11.6 mm., 0 (24 specimens) 10.5 to 11.8 mm. ; width of head, & 3 to 70 D%XXIl’TIOS OF GLOSSINA FUSCA. G. FUSCA

3.25 mm., 0 3.2 to 3.4 mm. ; width of front at vertex, d buff, terminr 0.6 mm., ? 0.6 to just under 1 mm. ; length of portion of hairs seventc palpi projecting beyond margin of buccal cavity, 6 3.2 to upper portioi 3.6 mm., !i! 3.6 to 4.5 mm. ; length of wing, rJ 10 to 11 mm., bulb buff-yell ? 10% to 12.6 mm. on its upper Thorax.- Large, dark brown, dark-winged species, with long and slender the usual tyl palpi, and without distinct dark bands on the abdomelz, which is grey or isabe lighter at the base ; in general appearance aery closely resembling clove-brown (Glossina nigrofusca, Newst. (p. 77), but distinguishable in both pollen) on e2 sexes by thr third joint qf the antennae not being clothed with long middle coxae median line : long or fairly Abdomen. brown or dl brown ; basa segment usu with a paler segment) in brown ; post segments inc yellowish-gre Fig. 17, A, p coloured. Legs buf more or less 7 inner side, m on outer side andjne pale hair, forming IC conspicuous .fringe down the anterior usual dark 1 crnd liosterior cages. under side r Head buff or cream-buff, posterior surface smoke-grey, face, more or less sides of front, and posterior orbits below yellowish pollinose ; middle, and .frontal stripe dark brown or mummy-brown, its anterior extremity sponding rin; mummy-brown or russet-brown ; ocellar spot dark brown, con- of first three netted with dark brown transverse band joining bases of vertical hair clothing bristles ; antennae partly buff, partly ‘brown or mouse-grey, under side a second joint dark brown at least on inner surface, third joint ochraceous ; I (tip of which is prominent) buff at base and on posterior margin, brown, last t elsewhere dark brown or mouse-grey, viewed in profile fairly Wings du broad, not clothed with long and tine pale hair, fringe of minute hyaline) ; ber hairs on front and hind margins in length not exceeding one- each end, pa sixth of breadth of third joint (qf. Figs. 16, 19 and 18), arista its extremity MCA. G. FUSCA: THORAX, ABDOMEN, LEGS, WINGS. 71

at vertex, 8 buff, terminal joint stout, dark brown at base beneath, branched of portion of hairs seventeen to twenty in number ; palpi buff, mouse-grey on ty, fj 3.2 to upper portion of outer surface, and dark brown at tips ; proboscis 0 to 11 mm., bulb buff-yellow or orange-buff, often with a dark brown blotch on its upper lateral margin on each side. Thorax.-Dorsum mouse-grey, with dark brown markings of the usual type, as shown in Plate VIII ; pleurae and pectus drab- grey or isabella-coloured pollinose, sternopleurae with a black or sely resembling clove-brown blotch (covered and partially obscured by yellowish &able in both pollen) on each side of median ventral line, between front and ‘bed with long middle coxae ; upper surface of scutellum dark brown, impressed median line and under surface buff, apical scutellar bristles of 0 long or fairly long. Abdomen.-Dorsum of first and second segments light mummy- brown or dull cinnamon, dorsum of remaining segments dark brown ; basal angles, lateral margins, and hind border of second segment usually infuscated ; third and fourth segments often with a paler area (of same colour as median portion of second segment) in centre, though their hind borders are still dark brown ; posterior angles and lateral margins of third to sixth segments inclusive yellowish-grey pollinose ; seventh segment yellowish-grey or brownish-grey pollinose ; hypopygium of 8 (cf. Fig. 17, A, p. 76) isabella-coloured, greyish-buff, or greyish fawn- coloured. Legs buff or ochraceous-buff, front femora usually with a more or less well-marked mouse-grey or greyish-brown streak on inner side, middle and hind femora often with a similar streak on outer side of their distal halves, and frequently also with the usual dark brown or mummy-brown elongate blotch on their under side near distal extremity ; hind tibiae usually with a ezrey, .face, more or less distinct dark brown or brown partial ring round h pollinose ; middle, and middle tibiae generally with a vestige of a corre- sponding ring ; tips of hind tibiae on outer side and upper side of first three joints of hind tarsi mummy-brown; short, felt-like hair clothing tips of hind tibiae on inner side, and also clothing under side of first three joints of hind tarsi, shining tawny third joint ochraceous ; tips of last two joints of front and middle tarsi dark rior margin, brown, last two joints of hind tarsi entirely clove-brown. hrofile fairly Wilzgs dull sepia-coloured, fairly dark ; alula paler (almost :e of minute hyaline) ; bent-up portion of fourth longitudinal vein, except at seeding one- each end, paler ; anterior transverse vein, although thicker at 18), nrista its extremity in contact with the third longitudinal vein, not so ‘ia DISTRIBUTION OF GLOSSINA FUSCA. G. FUSC noticeably incrassate as is the anterior extremity of the same and in bus transverse vein in the wing of G. brevipalpis, Newst. ; posterior Southern 1 transverse vein and portion of fourth longitudinal vein immedi- 1 8, Ikom, ately beyond it sometimes slightly infuscated. Squanaae semi- W. S. Cla hyaline, fringe long, yellowish. Halteres cream-buff. River, Eas Since the type of the present species is damaged, and of Protectorat unknown provenance, the re-description printed above has been close to she prepared from a 6 from Masaku, Ronietta District, Sierra Leone (Captain A. Protectorate, 16. ix. 1909 (Dr. J. J. Moore, W.A.M.S.), and a Q Edward, 19 from Boje, Liberia, 21. v. 1909 (Major A. Peame, R.A.M.C.). (with its pu .of Lake All

DISTRIBUTION OF G. j%ca, Walk. The specimens available for examination show that the range Dr. A. of the true G. fusca, Walk., as described above, extends at any states that : rate from the Sierra Leone to the Uganda Protectorates, although palpis muck no examples of the species have as yet been received at the found just a British Museum (Natural History) from French Guinea, the catch more French Ivory Coast, Togoland, Dahomey, Cameroon, French uncommon$ Congo, or the Congo Free State. There is at present no weather.” evidence that the species occurs in the East Africa Protectorate, Dr. Kingho German East Africa, the Nyasaland Protectorate, Portuguese East Africa, or Rhodesia. The following are the data with reference to the specimens examined, other than those already mentioned :- All that Sierra Leone Protectorate : 1 6, Warima, 9. ix. 1905, and Walk., rela 2 8 &, 3 Q Q , Robarri, 9, 10. ix. 1905 (Ca@ain-now Major-- Dr. A. G. H. W. Grattan, R.A.M.C.) ; 1 Q , Rotofunk, September, 1906 Bureau) f ou: (F. A. G. Fide) ; 1 6, Dodo, 15. iv. 1909, 1 Q , Gongo, near the sou 11. iv. 1909 (Major A. Pearse, R.A.M.C.); 1 8, Myopo, a number o 15. ix. 1909, 2 Q Q, Yelli and Moyamba, 13, 23., ix. 1909, together wit 1 Q , doctor’s quarters, Moyamba, 1. x. 1909, 4 Q Q , Monpeh, subsequently Matotoka, Maboon, and Connabai, 22-29. i. 1910-all localities presented to in Ronietta District (Dr. J. J. Moore, W.A.M.S.). Liberia : P.M.O., Uga 1 8, Dombolo, 23. iii. 1909, “ taken on an elephant,” 1 Q , near measures sol Da, 20. iv. 1909, 1 &, Da, 25. iv. 1909, 1 Q , on road between breadth, and Heye and Gondo, 2. v. 1909, 1 Q , near Bukei, 13. v. 1909, of G. brevipc 1s , Simbek, 16. v. 1909, ‘(caught buzzing round a candle, however, at about S.30 P.M.,” 1 &, Simbek, 18. v. 1909 (Major A. Pearse, between the t R.A.M.C.). Gold Coast Protectorate: 1 Q, Fumsu, Ashanti, * Sleeping 14. xii. 1907, “ caught in tent, 5.45 P.N.” (Dr. W. M. Graham, 1911). W.A.M.S.). Northern Nigeria : 2 Q Q , near rest-house, Ife, t See p. 96. ‘USCA. G. FUSCA: BIONOMICS AND PUPAL STAGE. ‘73 ty of the same and in bush between Poinia and Allu, 9. xi. 1910 (- Walfon). 1wst. ; posterior Southern Nigeria: 1 8, Okigwi, 30. v. 1910 (J. J. Simpson) ; t1 vein immedi- 1 d , Ikom, Cross River, Eastern Province, 6. vii. 1910 (Dr. Squamae semi- W. S. Clark) ; 1 6, Oban, 17. viii. 1910, and 1 Q, Cross lff. River, Eastern Province, 1910 (J. H. J. Farqukar). Uganda maged, and of Protectorate : 1 Q , Bugoma Forest, 35 miles west of Hoima, ,bove has been close to shore of Lake Albert, 1905, “caught biting native” t, Sierra Leone (Captain A. G’. H. Gmy, R.A.M.C.) ; 1 Q , north-east side of Lake MS.), and a Q Edward, 1906 (received from Dr. A. D. P. Hodges, P.M.O.) ; 1 Q R.9.M.C.). *(with its puparium), bred from pupa found near south-east corner ,of Lake Albert, 5. xii. 1906 (Dr. A. G. Bayskawe).

BIONOMICS. that the range lxtends at any Dr. A. Kinghorn, who has observed G. fusca in Ashanti, rates, although states that as regards its habitat this species resembles G. longi- Eeived at the pal+ much more closely than G. palpalis. (‘ It was usually h Guinea, the found just at the fringe of patches of bush, and it was rare to sroon, French catch more than one specimen or at most a couple. It feeds not tt present no uncommonly at night, and may be met with in dull or rainy t Protectorate, weather.” Specimens of G. fusca once or twice settled on e, Portuguese Dr. Kinghorn “ on a pitch-black, rainy night.“”

the specimens PRELIMINARY STAGES. All that can at present be said as to the life-history of 0. $ usca, ix. 1905, and Walk., relates to the pupal stage. On December 5th, 1906, -now Moior-- Dr. A. G. Bagshawe (now Director of the Sleeping Sickness stember, 1906 Bureau) found in a dry gully in the vicinity of the Mizizi River, 1 Q , Gongo, near the south-east corner of Lake Albert, Uganda Protectorate, 1 8, $-PO, a number of empty puparia belonging to the present species, 23: ix. 1909, together with one pupa from which the fly referred to above 7 0, Monpeh, subsequently emerged. These specimens, which were afterwards -all localities presented to the National Collection by Dr. A. D. P. Hodges, Z). Liberia : P.M.O., Uganda Protectorate, show that the pupa of G. fusca tt,” 1 Q , near measures some 8 mm. in length, by about 4.5 mm. in greatest road between breadth, and consequently agrees in its dimensions with the pupa 13. v. 1909, of G. brevipalpis, Newst.? The pupae of these two species can, od a candle, however, at once be distinguished by the shape of the notch >r A. Pearse, between the tumid lips or anal protuberances (cu. Figs. 6 B and 6 A). ISU, Ashanti, M. Graham, * Sleeping Sickness Bureau Bulletin, Vol. 3, No. 25, p. 136 (March 14, 1911). St-house, Ife, t See p. 96. 74 G. FUSCR : DISTINCTIVE CHARACTERS. DESCRI

If a pupa of each species be placed side by side in the position posterior e shown in the figures, that is to say with the tumid lips directed from G. .f upwards, and the more convex surface of what is now the upper characters extremity of the pupa towards the observer, in such a way that the tumid lips are viewed exactly in profile, it will be seen that whereas the notch between the lips of the pupa of G. brevi- paZ&s is relatively wide and shallow, looking something like a wide V,* the corresponding notch in the case of the pupa of G. fusca is narrower, deeper, and distinctly U-shaped. The Glossina fu! edges of the tumid lips are also sharper in the latter than in the Part 1, former species, in which the lips have a decidedly blunter and leas cleanly-cut appearance. The breadth of the posterior 6 .-L extremity of the pupa (as measured by a horizontal line drawn width of fr through the tumid lips when the pupa is in the position indicated) palpi projet is greater in G. fusca than in G. brevipalpis, and in the same of wing 10 position the outer markins of the lips in the former are more Dusky convex. distinguishe

AFFINITIES AND DISTINCTIVE CHARACTERS. isabella-colt or greyish-b With Glossina nigrofusca, Newst. (see below, p. 77), G. fu,.sci- qf the hind $euris, Austen (p. 75), and G. faban$ormis, Westw. (p. 81), 0. fusca forms an easily recognisable group (which may be termed Head ic the Glossirha fu.& group) of Tsetse-flies, characterised by large yellowish I size, a general dark brown coloration of t.he body without con- dark browr spicuous bands on the abdomen, and dusky wings. verse band nigrofusca, and .fusci$euris the palpi (and proboscis) are notice- clove-browr ably longer than in any other species of the genus ; in G. fabani- conspicuous fomis .they are of approximately the same length as in G. breci- branched I pa&is, New& From G. nigrqfusca and tnbaniformis, G.+fuscrc can dark mous at once be distinguished by the fringe of minute hairs on the front buff; probo margin of the third joint of the antenna, instead of being easily upper lateri visible when the antenna is examined under a platyscopic lens Thorax.- magnifying 15 diameters (nominal), and in length at least equal usual longit to one-fourth of the width of the third joint, being so short as to defined; hL be scarcely noticeable under a lens of the kind mentioned : t the admedian length of the palpi in G. fusca will also assist in distinguishing it longitudina: from. G. tabnn~forvais. The coloration of the pleurae (which are same colour drab-grey or isabella-coloured instead of dark grey) and of the Abdomen hind coxae (which are buff or greyish-buff instead of mouse-grey), umber-colou as well as the greater thickness of the black hairs fringing the of third to L * See p. 96. t Cf. figs. 18, 19, 16. ITERS. DESCRIPTION OF GLOSSINA FUSCIPLEURIS. 75 in the position posterior edge of the hind coxae, will serve to distinguish G..fusca d lips directed from G. f~sciyleuris, with regard to which further distinctive mow the upper characters will be found under the heading of the latter. ch a way that 11 be seen that I of G. brevi- kething like a Glossina fuscipleuris, sp. mv. the pupa of (Fig. 17, B.) shaped. The Glossina fusca, Austen (net Walker), Trans. Zool. Sot. London, Vol. XIX, SP than in the . Part 1, p. 100 (October, 1909). r blunter and the posterior 6 .-Length (1 specimen) 10.25 mm. ; width of head 3 mm. ; al line drawn width of front at vertex just over 0.5 mm. ; length of portion of ion indicated) palpi projecting beyond margin of buccal cavity 3.6 mm. ; length in the same of wing 10.4 mm. ner are more Dusky species, allied to and rewmbling G. fusca, Walk.,* but distinguished by the pleurae being dark grey instead qf drab-grey or

BS. isabella-coloured, by the hi& comae being mouse-grey instead of buf or greyish-bu.f, and by the black hair fringing the posterior margin 771, G. fwci- q,f the hind coxae below being distinctly longer and jiner. stw. (p. 81), lay be termed Head isabella-coloured, face and sides of front shimmering lsed by large yellowish pollinose, posterior surface mouse-gr& ; frontal stripe without con- dark brown; ocellar spot of same colour as frontal stripe, trans- In 6. .firsca, verse band connecting bases of vertical bristles darker ; antennae 1) are notice- clove-brown, third joint dull cream-buff at base, without a in G. taba.ni- conspicuous fringe of fine hair, arista buff, dark brown beneath, s in G. breci- branched hairs nineteen .in number in typical specimen ; palpi G. ~fuscrc can dark mouse-grey on outer side, darker at tips, extreme base I on the front buff; proboscis-bulb yellowish, with a dark brown blotch on its being easily upper lateral margin on each side. tyscopic lens Z’horaz.-General coloration of dorsum clove-brown, owing to It least equal usual longitudinal markings being well developed but not sharply :o short as to defined ; humeral calli and space on front margin on each side of iioned : t the admedian stripes olive-grey, interspaces between clove-brown inguishing it longitudinal markings elsewhere light mouse-grey ; pectus of : (which are same colour as pleurae ; scutellum as in G. fusca. ) and of the Abdomen.-First segment buff; dorsum of second segment raw- mouse-grey), umber-coloured, brownish at sides and on hind margin ; dorsum fringing the of third to sixth segments inclusive clove-brown, lateral margins

i. * Cj. p 69 and Plate VIII. 76 HYPOPYGIUM OF GLOSSINA FUSCIPLEURIS. DESCRIE

and posterior angles smoke-grey ; seventh segment smoke-grey D pollinose, central area mouse-grey ; hypopygium drab, viewed Since the from below more oval in outline (less flattened at the sides) than that of G. ~~uaca, Walk. (cJ Fig. 17). received, all f the species oc( Legs raw-sienna-coloured ; upper surface of femora dusky (dark grey or greyish-brown) ; middle and hind tibiae with an Congo Free S1 incomplete dark brown ring round centre ; distal halves of last two joints of front and middle tarsi dark brown, remainder of Unknown. last joint of middle tarsi brownish; last two joints of hind tarsi clove-brown, third joint of hind tarsi dark brown (less dark than AF Mention h belongs to th which is its n the characters as noted in t G. nigrofusca, pleuris, apart the ‘antennae from G. tabans by the (at leaf

PIQ. 17. Distal extremity of 6 abdomen of (A) GZossinafusca, Walk., and (B) G. .fuusci~Zeuris, Glossina nigrof sp. nav. (ventral aspect), showing (h) hypopygia and (hect.) hectors.* (x 15.) ology, Seriel last two joints), second joint of hind tarsi brown, first joint of Glossina gross: hind tarsi brownish above. Wings dull sepia-coloured, fairly dark ; extremity of anterior 1 3 q&6; s--L >‘A .; w transverse vein in contact with third longitudinal vein con- width of fron spicuously incrassate ; alula paler than remainder of wing, length of por nearly hyaline. Syunmae du sky, of same colour as membrane of cavity, 8 4 tc wing; fringe of squamae brownish. Halteres cream-buff. 11.75 mm., 0 The typical specimen of this species is a & in the British Museum (Natural History) from the Eturi Forest, north-eastern Resembling Congo Free State, between Irumu and Avakubi, alt. 2,000 ft., especially in t: Oct. 1906 (Hon. Gerald Legge and A. F. R. Wo7laston),-collected joint of the a during the Ruwenzori Expedition, 1905-1906. * Cf. p. * SEC p. 14 (first paragraph and riots *). FUSCIPLEURIS. DESCRIPTIOK OF GLOSSINA NIGROFUSCA. 77 segment smoke-grey DISTRIBUTION OF Cl. fuscipleuris, sp. nav. gium drab, viewed cl at the sides) than Since the type is the only example of G. juscipleuris yet received, all that can at present be said as to distribution is that e of femora dusky the species occurs in the forest region in the north-east of the lind tibiae with an Congo Free State. listal halves of last BIONOMICS. rown, remainder of joints of hind tarsi Unknown. )wn (less dark than AFFINITIES AND DISTINCTIVE CHARACTERS. Mention has already been made of the fact that this species belongs to the Glossina fusca group.* From G. fuxa itself, which is its nearest ally, G. fuscipleuris may be distinguished by thecharacters att’orded by the pleurae, hind coxae, and hypopygium, as noted in the diagnosis and description printed above. From G. nigrofusca, Newst.,? and G. tnbaniformis, Westw.,$, G. ~%sci- pleuris, apart from its smaller size, differs in the third joint of the ‘antennae not being conspicuously fringed with fine hair ; from G. tabaniformis the present species may also be distinguished by the (at least relatively) greater length of its palpi.

Glossina nigrofusca, Newt. (Fig. 18.)

and (B) G. fusci&znris, Glossina nigrofusca., Newstead, Annals of Tropical Medicine and Parasit- I hectom.* (x 15.) ology, Series T. M., Vol. IV, No. 3, p. 370 (December 20, 1910). own, first joint of Glossina grossa, Newstead, ibid., p. 373 (met Bigot).

memity of anterior 8, 0 .--Length, 6 (2 specimens) 12.4 mm., ? (2 specimens) iudinal vein con- 14.2 mm. ; width of head, 4 3.4 to 3.75 mm., 0 3.75 mm. ; nainder of wing, width of front at vertex, $ 0.6 to just under 1 mm., 0 1 mm. ; P as membrane of length of portion of palpi projecting beyond margin of buccal ream-buff. cavity, 6 4 to 4.4 mm., ? 4.8 mm. ; length of wing, $ 11 to 8 in the British 11.75 mm., ? 12.75 mm. est, north-eastern Resembling Glossina fusca, Walk. (see p. 69 and Plate VIII), bi, alt. 2,000 ft., especially in the ? sex, but distinguishable inter alia by the third aston),-Collected joint of the antennae being clothed with long and $ne pale huir,

$ P. 81. 7. * Cf. p. 74. t See below. $8 G. NIGIiOFUSCA: HEAD AND THORAX. DISTRIBl fornaiaq a conspicuous fringe down the anterior and posterior edge8 smoke-grey p (cf. Figs. 18 and 16), the length of the fringe on the anterior edge blotch on ea being equal to from one-half to three-fourths of the width of the G.&sea, but third joiiat. be judged frc so long as thl Head greyish-buff, posterior surface smoke-grey or mouse- Abdomen. grey ; .f’rontnl stripe mummy-brown, paler anteriorly ; ocellnr spot second segmc and dark brown transverse band joining bases of vertical bristles . each side, ar as in G. fusca; antennae partly buff, partly brown or mouse-grey, fourth segme second joint brown or dark brown, third joint mouse-grey or angles and ( greyish-brown, more or less buff at base, longer and narrower inclusive sm than in G. fusca (cp. Figs. 18 and 16), distal extremity prominent, brownish-gre hypopygium Legs buf inner and ou in case of h dark brown femora near hind tibiae conspicuous) often with a tips of hind of last two j( front tarsi d: dark than ti entirely Clov mummy-bra\ sometimes d: Bw. 18. Antenna of Glossina ~~i~rofuscn, Newst. (x 30.) of tips of hl tarsi as in G sometimes very prominent, arista buff, terminal joint as usual Wings, s4 dark brown at base beneath, ridge on upper side of proximal The abov portion of terminal joint very prominent, branched hairs more from Ashant numerous and closer together than in G. .fusca, varying in number from twenty two to twenty-eight or twenty-nine ; palpi slender, lighter or darker mouse-grey on outer side above, dark brown at tips, buff’ or greyish below on proximal three-fourths ; The few prol~oscis bulb as in G. ~&KXL the present 1 Y%oram.-Dorsum mouse-grey or drab-grey, with dark (olive- any rate frc brown or dark brown) markings of the usual type; the latter, appears to 1 though usually not clearly defined, are often so extensive that * One of th little of the grey interspaces is visible; pleurae and pectus scutellar bristl THORAX. DIi3TRIBUTlOx OF GLOSSINA NIGROFUSCA. $9

,nd posterior edges smoke-grey pollinose, sternopleurae, as in G. .fusca, uTith a dark the anterior edge blotch on each side of median ventral line ; scutellum as in ’ the width of the G. fuscu, but apical scutellar bristles of ? (so far at least as may be judged from a single specimen *j shorter, little more t’han half :-grey or mouse- so long as those of d. Abdonzen.-Dorsum clove-brown or dark brown, first segment, orly ; ocellnr spot t vertical bristles second segment except hind border and an ill-defined area on n or mouse-grey, . each side, and often also an ill-defined median area on third and It mouse-grey or fourth segments (not reaching hind borders), cinnamon ; posterior 3r and narrower angles and extreme lateral edges of third to sixth segments inclusive smoke-grey or drab-grey pollinose ; sevent’h segment :mity prominent, brownish-grey or mouse-grey pollinose, lighter grey on sides; hypopygium of & isabella-coloured or greyish fawn-coloured. Legs buff or ochraceous-buff, upper portion of femora on inner and outer sides, except at base and also at distal extremity in case of hind pair, more or less greyish-brown or mouse-grey ; dark brown elongate blotch on under side of mid_dle and hind femora near distal extremity often strongly marked ; middle and hind tibiae with a more or l&s conspicuous (sometimes very conspicuous) dark brown ring round middle, front tibiae also often with a more or less distinct trace of a corresponding mark, tips of hind tibiae with a faint brown blotch on outer side ; tips of last two joints of middle tarsi and tip of penultimate joint of front tarsi dark brown, tip of last joint of front tarsi brown (less dark than tip of preceding joint), last two joints of hind tarsi entirely clove-brown, upper side of remaining joints of hind tarsi mummy-brown or paler (cinnamon), third joint of hind tarsi sometimes dark brown at tip; short, felt-like hair on inner side of tips of hind tibiae and under side of first three joints of hind tarsi as in G. fusca. joint as usual Wings, squamae, and halteres as in G .fusca. de of proximal The above re-description has been drawn up from a 8 and $’ led hairs more from Ashanti (Dr. W. N. Graham), in the Museum collection. ‘a, varying in Ity-nine ; palpi le above, dark DISTRIBUTION OF G. niyrofusca, Newst. three-fourths ; The few specimens of this species received or recorded up to the present time show that the range of G. nigrofusca extends at th dark (olive- any rate from Ashanti (where, according to Dr. Graham, it le; the latter, appears to be of local rather than general occurrence) to the extensive that e and pectus * One of the two females available for examination has lost its apical scutellar bristles. SO G. NIiX0FUSCA : DISTINCTIVE CHARACTERS. DESCRIP’I

Congo Free State. The data with regard to the four examples segment in in the possession of the British Museum (Natural History) are latter. In as follows :- blance betwl Gold Coast Protectorate : 1 0, Dunkwa, Ashanti, 23. v. the former I 1907, ‘( caught at 2.30 p.m. on collector’s leg,” and 1 8, Obuasi, palpi, as we1 Ashanti, 31. vii. 1907, “in bush path” (Dr. W. M. Graham, of the third W.A.M.s.). Southern Nigeria : 1 &, Oni River, 70 miles east of Lagos, 16. v. 1910, “ caught in house, biting leg of European seated at table,” and 1 0, same locality, i. xii. 1910, (‘ caught in house ” (Dr. W. A. Lamborn : presented by the Entomological Research Committee). Newstead records * 2 8 8 and 1 0, taken in Ashanti, at Sunyani, Odumase, and Atroni, on March 5, April 27, and August 16, 1910, respectively (Dr. d. Kinghorn), Glossina taba and 1 6, caught at Kasongo, Congo Free State, on February 6, p. 268, Pk und Forst 1904 (the late Dr. J. E. Dutton and Dr. J. L. Todd). Glossina gros

BIONOMICS. & 0 .--I 10.25 to 1O.f All that can as yet be said under this heading may be width of fro] gathered from the brief notes printed in the previous paragraph, length of pa which show that G. nigrofusca sometimes enters houses and will cavity, 6 3 even attack the inmates, and that it appears to settle by ? 11.55 to 1 preference on the legs. Large, a Symmwy, AFFINITIES, AND DISTINCTIVE CHARACTERS. G. nigrofusc the palpi (ar The present species was actually described by Newstead the fringe of under the name Glossina grossn, Bigot, which, as will be seen antennae not below,t now proves to be a synonym of G. tabaniformis, Westw. half to h-et The alternative name nigrofusca, suggested by Newstead for and 1s). what he called G. grossa in the event of his species being found to be distinct from Bigot’s, must therefore be adopted. Head crf As stated above,$ in dealing with the atlinities of Glossina of head mo *fusea, Walk., Glossina nigrqfusca belongs to the G. fusca group, riorly ; 0celL from the three other members of which at present known transverse b it may be distinguished by the length of the fine hair on the antennae par third joint of the antennae (compare Figs. 18, 19 and 16). mainly dark Although the females of G. nigrofusca and fusca closely resemble brown (a lar each other in general appearance, the former is on the whole a * See belo7 larger and bulkier insect; the head in both sexes is distinctly t In the c: wider in G. niyofusca than in G. fusca, and the second abdominal kindness of iV History), the * Lot. cit., p, 374. t Cf. p. 81. ; P. 74. buccal cavity : DESCRIPTION OF GLOSSINA TABANIFORMIS. 81 segment in the male is also broader in the former than in the latter. In size and general appearance there is a close resem- blance between G. nigroficsca and G. tabanifornis, We&w.,* but the former may be distinguished by the greater length of the palpi, as well as of the hair on the anterior and posterior edges of the third antenna1 joint.

Glossina tabaniformis, We&v. (Fig. 19.) Gloss&a tabaniformis, Westwood, Proc. Zool. Sot. Lord., Part XVIII, p. 268, Plate XIX, fig. 3 (1850) :-r&x Stuhlmann, Bericht iiber Lend- und Forstwirtschaft in Deutsch-Ostafrika, Bd. i, Heft 2, p. 173 (1902). Glossina grossa, Bigot, Ann. Sot. Ent. Fr., Vol. LX, p. 377 (1891). 8 0 .--Length, & (1 specimen) 10.4 mm., 0 (4 specimens) 10.25 to 1O.S mm. ; width of head, 6 3.25 mm., ? 3.4 to 3.5 mm.; width of front at vertex, $ 0.6 mm., $’ 0.6 to just under 1 mm. ; length of portion of palpi projecting beyond margin of buccal cavity, d 3 mm., $’ 3 to 3.25 mm. ; 7 length of wing, 8 10 mm., ? 11.55 to 12.25 mm.

Large, dark brown, dusky-winged species, closely resembling G. nigrofusca, Newst., in general appearance, but distinguished by the palpi (and proboscis) being much shorter, and by t7Le length of the fringe of jne hair on the anterior edge of tile third joint of the antennae not exceeding one-third (instead of being equal to from one- hay to t7zree-fourths) qf the width of the third joint (cf. Figs. 19 and IS).

Head cream-buff, antenna1 cavity pearl-grey, posterior surface of head mouse-grey; frontal stripe mummy-brown, paler ante- riorly ; ocellar spot inconspicuous, of same colour as frontal stripe, transverse band joining bases of vertical bristles dark brown ; antennae partly buff, partly dark brown or brownish, second joint mainly dark brown or more or less brownish, third joint dark brown (a larger or smaller area at base buff or cream-buff), often * See below. t In the case of the type of G. gvossa, Big. (a 0 ), which, through the kindness of Mr. G. H. Verrall, is now in the British Museum (Natural History), the portion of the palpi projecting beyond the margin of the buccal cavity is somewhat longer, and measures 3.5 mm. G2 S2 DESCRIPTION OF GLOSSINA TABANIFORMIS. DISTRII broader and in Q at any rate usually shorter than in G. niqro- wanting, Y .fusca, Newst. (cf. Figs. 19 and lS), distal extremity prominent, side at dis fringe of hair on front margin in length equal to from one-fourth side of firs to one-third of width of third joint, fringe of minute hairs on than the 1 hind margin also much or distinctly shorter than corresponding Wings fringe in case of 6’. nigrofusca, arista buff, first and second joints Altho and third joint at base dark brown beneath, branched hairs from F.R.S., I eighteen’ to twenty-three in number ; palpi dark mouse-grey on (a Q fron outer side above, dark brown at tips, proximal three-fourths buff insect is below ; the palpi usually appear less slender-i.e. broader (viewed drawn ul in Augue the Briti Research

All t‘ the area and Gold The folk examined Ivory Cc Alluaud : 1 Q , Ikoi Clark : t Congo F J. E. Du Fig. 1% Antenna of Glossina tnbaniformis, W&w. (X 30.) from above) and deeper (viewed from the side)-than t’hose of G. nigrofttsca ; proboscis bulb as in latter species. It w T?Lorax.-General coloration as in G. nigrofuscn, usual dark Christy, markings of dorsum more or less distinct; apical scutellar Journal 1 bristles of Q not short (about four-fifths so long as those of 6 ). related 1 Abdomen as in G. nigroj’usca. ments sl Legs buff, femora less infuscated than those of G. nigrofusca; be fount middle femora sometimes with a dark greyish-brown elongate it bites 1 blotch on outer side beyond middle, hind femora occasionally * c with a faint indication of a similar infuscation ; middle and hind tibiae with a vestige of a more or less incomplete dark brown ring round middle, which, however, may be very faint or entirely 4NIFORiWIS. DISTRIBUTION OF GLOSSINA TABANIFORMIS. 83

tn in G. IZZ@~- wanting, hind tibiae with no trace of a brown blotch on outer ity prominent, side at distal extremity; tarsi as in G. nigrqfusca, though upper rom one-fourth side of first three joints of hind tarsi is sometimes scarcely darker inute hairs on than the hind tibia. corresponding Things, quamae, and hnlteres as in G. nigrofusca. d second joints Although, throu& the kindness of Professor E. B. Poulton, hed hairs from F.R.S., I have been enabled to re-examine the type of this species mouse-grey on (a Q from the Gold Coast, in the Oxford Museum’), since the bee-fourths buff insect is somewhat faded, the above re-description has been roader (viewed drawn up from a 6 and Q taken at Oban, Southern Nigeria, in August, 1910, by Mr. J. H. J. Farquhar, and presented to the British Museum (Natural History) by the Entomological Research Committee.

DISTRIBUTION OF G. tabanifomis, Westw.

All that can at present be said under this heading is that the area of distribution of G. tabaniformis includes the Ivory and Gold Coasts, Southern Nigeria, and the Congo Free State. The following are the data with reference to the specimens examined, in addition to those already mentioned :-French Ivory Coast : 1 Q (the type of G. grossa, Bigot), Assinie (C/l. A&ad: presented by Mr. G. H. Verrall). Southern Nigeria : 1 Q , Ikom, Cross River, Eastern Province, 6. vi. 1910 (Dr. W. 5’. Clark : the property of the Entomological Research Committee). Congo Free State : 1 Q , Leopoldville, 26. xii. 1903 (the late Dr . J. E. Dutton, and Drs. J. L. Todd and C. Christy).

-than those of It would seem probable that the observations by Dr. C. cn, usual dark Christy, published by the present writer in the British Medicd ?ical scutellar Journal of September 17, 1904, as referring to G. fusca, actually 3 those of 8). related to G. tabanifornais. If this be so, Dr. Christy’s state- ments show that in the Congo Free State G. tabanifornais may G. ni~rofusca; be found round houses and on board river steamers, and that ‘own dongrate it bites by night-at any rate, until about 11.0 P.M. a occasionally iddle and hind * Cf. Austen, “ A Blonogrsph of the Tsetse-Flies,” p. 98 (1903). e dark brown int or entirely 84 G. TABANIFORMIS : DISTINCTIVE CHARACTERS.

AFFINITIES AND DISTINCTIVE CHARACTERS. So far, at least, as regards external characters, Clossina tnbaniformis may be distinguished from G. niqrofusca, New&. (p. 77), G. fusca, Walk. (p. 69), and G. fuscipleuris, Austen (p. 75) -the three other species of the Glossina fusca group at present known, by the comparative shortness of its palpi ; the length of 1. I the fringe of hair on the front border of the third joint of the antennae will also serve to distinguish G. tabaniformk from G. nigrqfusca on the one hand, and from G. &sea and fuscipleuris on the other. The general resemblance of both G. nigrofusca and fusca to G. tabaniformis is certainly very strong, and it is possible I that either the first or last mentioned species may ultimately 4 prove to be a form of 6. fusca, Walk. ; in order to decide this question, however, a comparison of the male genital appendages is 2. ’ essential, and for this purpose the material at present available is insufficient in the case of G. tabaniformis.

Glossina om Glossina Fli@ (190: (Nov Med Kais D&W (ad: logic Vol. pp. I Glossina ma ERS. S5

68&a mst. CHAPTER VII. .75) THE GLOSSINA BREVIPALPIS GROUP. Bent h of SYNOPSIS OF SPECIES. the 1. Dorsum of thorax with four sharply defined, iom dark brown, more or less oval or elongate wris spots, arranged in a parallelogram, two in front of and two behind the transverse and suture; proboscis bulb with a sharply lible defined brown or dark brown tip ...... Zongipewnis, Corti. Dorsum of thorax without such spots; pro- WY ‘ this boscis bulb not brown or dark brown at the tip ...... 2 )s is 2. Wings with upper, thickened portion of able anterior transverse vein much darker in colour than adjacent veins, and thus stand- ing out conspicuously against the rest of the wing ...... brevipalpis, Newst. Wings with upper, thickened portion of anterior transverse vein not much darker in colour than adjacent veins, and thus not standing out conspicuously against the rest of the wing (wings practically unicolorous)...... ,. . . medicorum, sp. nov.

Glossina brevipalpis, Newt. (Plate IX.) GIossina brevipafpis, Newstead, Annals of Tropical Medicine and Parasit- ology, Series T. M., Vol. IV, No. 3, p. 372 (December 20, 1910). Glossina fusca, Austen (net Walker) ; Austen, Monograph of the Tsetse- Flies, pp. 95-97, Plate VI (1903); Sander, Die Tsetsen, pp. 66-68 (1905) ; Koch, Deutsche Medisinische Wochenschrift, Bd. xxxi (November 23, 1905),-translated by P. Falcke in Journal of Tropical Medicine, Vol. IX, pp. 137-138 (1906) ; Stuhlmann, Arbeiten aus dem Kaiserlichen Gesundheitsamte, Bd. xxvi, Heft 3, p. 301 et seq. (1907) ; Davey, Bulletin of Entomological Research, Vol. I, Part 2, pp. 143-145 (July, 1910) ; Pask, ibid., pp. 145146 ; Sanderson, Bulletin of Entomo- logical Research, Vol. I, Part 3, pp. 225-226 (October, 1910), and Vol. I, Part 4, pp. 299-302 (January, 1911) ; S. A. Neave, ibid., pp. 306-310. Glossina tabaniformis, Stuhlmann (ncc Westwood), Bericht iiber Land- und Forstwirtschaft in Deutsch-Ostafrika, Bd. i, Heft 2, p. 173 (1902). 6, 0 .-Length, 6 (317 specimensj 102 to 13.45 mm., 0 (42 specimens) 11 to 13.5 mm. ; width of head, 8 395 to 81; DESkRIPTION O F GLOSSINA RHEVIPALPIS. DESCI

3.6 mm., ? 3.25 to 0.75 mm. ; width of front at vertex, S 0.5 bulb, and ( to 0.6 mm., $’ just under 1 mm. to 1 mm. ; length of portion of the thorax palpi projecting beyond margin of buccal cavity, 6 2.25 to proboscis a (3 ) distinc~i

Heads of males of (A) Glossina brev+al>is, Kew&, (8) G. fuscn, Walk., and (c) G. medi- co,u?n, sp. nav. (see p. 98), dorsal aspect ; showing differences in the size and shape of the hend 8s & whole, in the shape of the eyes, and in the length of the palpi. (X 10.) Heads of fern: - medicorun 3 mm., $! 3 t’o 3.25 mm. : length of wing, & IO.2 to 12 mm., shape of t ? 11.5 to 14 mm. palpi. (x

Large species, somewhat reselnbling Cr. longipennis, C’orti, but in the & &sting&shed inter alia by the front being narrower, and by the coloration ( absence of a dnrk brown ocellm spot, of cc brown tip to the proboscis usual longi WIPALPI S . DliSCRIPTION OF GLOSSINA BREVIPALPIS. 87 at vertex, & 0.5 bulb, and qf the chnracteristic dark brow spots on the dorsum of th of portion of the thorax; distinguished from Q. fusca, Walk., inter alia by the vity, & 2.25 to proboscis and palpi being ~zuclr shorter, the head (especially in tlr(J 6 ) distindy wider and in both seam closer to thr ilmax, the front

:., and (0 G. wedi- the size and shape c he length of the Fig. 21. Eeads of females of (A) Glossina breui@&, New&, (8) G. fusca, Walk., and (C) G. 2 to 12 mm., mdacorum, sp. nav. (see p. 98), dorsal aspect ; showing differences in the size and -- shape of the head as a whole, in the shape of t,he eyes, and in the length of the palpi. (x 10.)

Lis, Corti, but in the 8 relaticlely narrower, the frontal stripe and general *, and bg the coloration qf the dorsuvn qf the thorax ad of the wings paler, the ) the proboscis usual longitudinal markings on the dorsum qf the thorax generally SS G. BREVIPALPIS : HEAD, THORAX, ABDOMEN. G. B less strongly developtd, the size being often larger, and the general segment in b appearance usually bulkier. hypopygiun~ ol roughly oval Head cream-buff, posterior surface greyish ; frontal stripe longer hair a buff, or light raw-sienna coloured ; oceltar spot light brownish, dark brown ( inconspicuous, ocelli relatively large ; face (including facial pit), very short (sl frontal margins narrowly, and posterior orbits below shimmering Legs buff, whitish-yellow pollinose ; bases of vertical bristles connected by an indication a more or less distinct brown or brownish band ; eyes with their darkest, and transverse diameters longer than in G. fusca, SO that,, when the incomplete, 81 head is viewed from above, its lateral portions appear somewhat front tibiae flattened in an antero-posterior direction ; antennae buff or cream- generally red buff, second joint brown on inside, third joint brown or dark middle ; fron brown except extreme base or proximal third, anterior distal middle and 1 extremity of third joint often prominent, arista buff, terminal usually slighl joint fairly stout, dark brown at base beneath, branched hairs brownish blot eighteen to twenty-two in number ; palpi buff, lighter in colour towards the 1 as well as shorter than in G. fusca and also stouter, mouse-grey clothed benea, on upper portion of outer surface and darker at tips; proboscis tarsi more 01 bulb pale yellow, often with a brown spot on its upper margin on middle and hi each side. of middle tart Thorax light mouse-grey or isabella-coloured, dorsum in sometimes infl rubbed specimens sometimes tinged with fawn colour, series of Wings pa markings on dorsum, as described for G. patpalis, much reduced darker than and often very inconspicuous, mouse-grey or light brown; most, of anterior tr distinct are the two narrow admedian stripes, the narrow curved verse vein tog stripe outside these on each side, two faint blotches enclosed by vein lying toT each curved stripe, one in front of and one behind the transverse vein and portic suture, and the spot on the suture, between the admedian and infuscated ; b: the curved stripe on each side ; spot on inner margin of humeral sixth also da callus usually very faint ; dark blotches on pectus (st.ernopleurae), brownish or : which are often conspicuous in G. fusca, scarcely visible, or cream-buff. brownish in colour and faintly indicated; scutellum greyish- The specin brown, margin and impressed median line buff or cream-buff, drawn up are apical scutellar bristles in ? of medium length. (Dr. J. B. Dc Abdomen.-Dorsum dark brown, first segment, and also Committee (!I second segment to a greater or less extent, paler, ochraceous-buff tion- the 8 or cinnamon-coloured, extreme hind margins of second to sixth on the Karol segments inclusive usually pale (cream-buff), posterior angles of 5. viii. 1909 these segments drab-grey or smoke-grey, dark brown area, 33. i. 1910. occupying third to sixth segments inclusive, sometimes distinctly broken up into a series of interrupted transverse bands ; seventh X, ABDOMEN, G. BREVIPALPIS: LEGS AND WINGS. 89

3 and the general segment in both sexes yellowish-grey or fawn-coloured pollinose ; hypopygium of 8 (Fig. 84 A, p. 99) buff pollinose, seen from below roughly oval in outline, with its anterior margin flattened; h ; frontal stripe longer hair at base of second segment golden-yellow in middle, )t light brownish, dark brown or black at sides; third, fourth and fifth segments uding facial pit), very short (shorter than corresponding segments in G. &WLC). glow shimmering Legs buff, middle and hind tibiae as a rule with (at any rate ;les connected by an indication of) a brown ring round middle, most conspicuous, * eyes with their darkest, and often complete on hind tibiae, fainter, generally ;’ that, when the incomplete, and often reduced to a mere trace on middle tibiae, appear somewhat front tibiae usually with a vestige of a corresponding ring, bae buff or cream- generally reduced to a faint brownish blotch on inside in ; brown or dark middle ; front femora on inside (sometimes also on outside), 1, anterior distal middle and hind femora on outside towards distal extremit,y ta buff, terminal usually slightly infuscated, middle femora generally with a I, branched hairs brownish blotch on under side near tip ; front and hind tibiae lighter in colour towards the tips and first three joints of front and hind tarsi suter, mouse-grey clothed beneath with short golden pile ; last two joints of front at tips ; proboscis tarsi more or less infuscated at the tips ; last two joints of upper margin on middle and hind tarsi, except proximal half of penultimate joint of middle tarsi, dark brown ; extreme tips of other tarsal joints ued, dorsum in sometimes infuscated. colour, series of Wings pale isabella-coloured, distal. extremity somewhat is, much reduced darker than proximal third; veins pale tawny, upper portion ght brown ; most of anterior transverse vein strongly incrassate, anterior trans- re narrow curved verse vein together with adjacent portion of fourth longitudinal tches enclosed by vein lying towards base of wing, as well as posterior transverse id the transverse vein and portion of fourth vein immediately beyond, conspicuously le admedian and infuscated ; basal portion of the veins from the second to the zrgin of humeral sixth also darker. Squamae not infuscated, fringe of pale 8 (sCernopleurae), brownish or yellowish, fine, silky hair rather long. Halteres brcely visible, or cream-buff. utellum greyish- The specimens from which the above description has been ff or cream-buff, drawn up are a 8 and Q from the Nyasaland ProteE torate (Dr. J. B. Davey : presented by the Entomological Research ment, and also Committee (Tropical Africa)), in the British Museum Collec- p, ochraceous-buff tion- the 6 from North Nyasa, taken in the evening second to sixth on the Karonga-Songwe road, two miles south of Kaporo, Isterior angles of 5. viii. 1909, the Q caught at Nkomo, Marimba district, rk brown area 23. i. 1910. ,etimes distinctly I bands; seventh 90 I)ISTlrtTBUTTOS O F GLOSSINA BREJWALPIS. BION

DISTRIBUTIOX OF G. bwvipulpis, Newst. (Dr. F. -K Sabaki Riv’ Glossinn b~~i~+i~, to which the name G. ,fuscn has hitherto Company) ; been applied in error, is the common large Tsetse-fly in many River, near parts of South-Central and East Africa, but, although found in vet.- c’rpt. . the Congo Free State, it does not, so far as our present know- (the Sultan ledge goes, occur in West Africa proper. Reliable records and .2 Q Q, Ki the data attached to the specimens available for examination Mofat, CA show that G. brevipalpis exists in Portuguese East Africa, the A.M.S.) ; 11 Nyasaland Protectorate (where it has a wide distribution, being Sleeping Si abundant at certain seasons in and near the Songwe Valley, at Mwatate d the north end of Lake Nyasa, besides occurring in and on the R. J. Stord edge of the Elephant Marsh, near the Lower Shire River, to the near Char07 south of the lake *), North-Eastern Rhodesia,? German East Lufira Riv Africa (where it is found in large numbers in the vicinity of (8. Neave, Amani-at the foot of the Usambara Mountains-and also 2. viii. 19( elsewhere $), the (British) East Africa Protectorate, the Katanga 17. viii. 191 district of the Congo Free State, and in Angola. It has not been thought necessary to give precise details concerning the localities and dates of capture of the whole of the 359 Specimens examined, the majority of which were In view collected in the Nyasaland Protectorate by Drs. Sanderson proved to 1 and Davey, to whose papers (quoted above) reference should that in tht be made for further information. The data as to the speci- stated to b mens received from countries other than Nyasaland are as domestic ar follows :- of the bio Portuguese East Africa : 2 8 8, 1 0 , Bamboo Creek, Beira important Railway, 190s (received from Ll. E. W. Bevan, M.R.C.V.S.). to the obs Sorth-Eastern Rhodesia : 1 & , Kanga Stream, eight miles east J. B. Dave: of Hargreaves, Luangwa Valley, “just after sunset,” 16. ix. 1910, concerning and 1 ? , within a few hundred yards of Molilo’s, fourteen miles other Tsets east of Hargreaves, “ about 10 a.m.” 17. ix. 1910 (5’. A. Neave : Stuhlmann presented by the Entomological Research Committee). German species as East Africa : 1 ? ; Kilima Njaro, 1888 (F. J. Jadcson, C.B., description 6”X.G.) ; 11 & 8, 7 $’ 0 (all bred in captivity), 2 dried has also prc larvae and 11 pupae and puparia, Amani, January, 1907 as observed conditions ; * Cf. Sanderson, lot. cit., p. 299, and Davey, 10~. cit.,, p. 143. t For notes on the occurrence of G. breripalpis m N.-E. Rhodesia, upon his s whew it has hitherto been met with only in the Luangwa Valley, in the tioned abol vicinity of Hargreaves (Chutika’s), and in very small numbers, see Protectorat S. A. Neave, Bulletin of Entomological Research, Vol. I, Part 4, pp. 306.-310 (January, 1911). $ Cf. Stuhlmann, Arb. a. d. I&is. Gesundheitsamte, Bd. xxvi, Heft 3, pp. 301-302 (1907). REVIPALPIS. BIONOMICS OF GLOSSlNd BREVIPALPIS. 91

3wst. (Dr. l? Voaseler). East Africa Protectorate : 1 Q , Upper Sabaki River, 1690 (presented by the late British East Africa &a has hitherto Company) ; 1 Q , Uganda Railway (‘1 Kiboko River or Sabaki se&fly in many River, near its junction with the Tsavo River), 1898 (the late lthough found in Vet.-Cgpt. A . J. Haslam) ; 1 ,j, 2 Q Q, Witu Forest, 1895 or present know- (the Sultan of Wita-presented by the late W. S. Godfrey) ; able records and .2 Q Q , Kibwezi, 1903, “ in railway carriage” (Dr. R. U. for examination Mqfat, CY.X.G.--per Colonel Sir David Bruce, C.B., F.R.S., East Africa, the A.M.S.) ; 10 6 8, 10 Q Q, Kibwezi, 1903 (received from t’he istribution, being Sleeping Sickness Commission, per Dr. D. Nabarro) ; 1 Q , ongwe Valley, at Mwatate district, near Voi, January, 1909 (received from g in and on the R. J. Stordy). Congo Free State (Katanga district) : 4 8 8 , lire River, to the near Charowi, 21. vi. 1907, and 12 & 6 , 3 Q Q , Kiubo Falls, ,t German East Lufira River (Lat. 9” 30’ S., Long. 27" E.), 23. vi. 19Oi L the vicinity of (5’. Neaue, M.R.C.P., M.R.C.S.) ; 1 6, 1 Q , Kiubo Falls, rtains-and also 2. viii. 1909 (Dr. F. 0. Xtoehr). Angola : 1 Q , Canhoca, ate, the Katanga 17. viii. 1910 (Dr. W. J. Ansorgej. e precise details BIOXOMICS. of the whole of of which were In view of the possibility that G. brevipalpis may ere long be Drs. Sanderson proved to be a carrier of human trypanosomiasis, and the fact reference should that in the German East African littoral, at any rate, it is as to the speci- stated to be the chief disseminator of Tsetse-fly disease among rasaland are as domestic animals,* special importance attaches to a knowledge of the bionomics of the present species. Although many boo Creek, Beira important details have doubtless still to be recorded, thanks n, M.R.C.V.S.). to the observations and writings of Drs. Franz Stuhlmann, eight miles east J. B. Davey, and Meredith Sandersoqt more is already known et,” 16. ix. 1910, concerning the habits of G. breaipalpis than about those of any 6, fourteen miles other Tsetse, with the exception of G. pal&is and morsitans. 10 (S. A. Neave: Stuhlmann (who, like the other authors, of course writes of the .ittee). German species as Glossinn “f USCCL “), besides furnishing a detailed . Jackson, C.B., description of the anatomy and histology of the internal organs,+, Yivity), 2 dried has also provided us with an interesting account of G. lwempalpis January, 19Oi as observed by him in German East Africa, both under natural conditions and in captivity, and the following r&me is based p. 143. 1 N.-E. Rhodesia, upon his statements and those of the English writers men- gwa Valley, in the tioned above, whose observations were made in the Kyasaland la11 numbers, see Part 4,pp. 306.310 Protectorate. * Stuhlmann, Zoc. cit., p. 30% Bd. xxvi, Heft 3, t Cf. references on p. 85. $ Lot. cit. 92 BIONOMICS OF GLOSSINA BREVIPALPIS. BION

Gloss& brevi$alpis, which is often met with in very large at about 5. numbers, is found, at altitudes varying from sea-level to some digging rot 1,700 feet, as a rule only where there is abundant shade and successful. protection from wind, its favourite haunts consisting of bush to emerge h mixed with creepers and young forest trees, either close to or dust on 1 watercourses (dry or otherwise), or at any rate within a few writer adds hundred yards of water. Stuhlmann has encountered the fly in the even close to the sea, near Dar es Salam, and Davey records the occurrence of a single specimen amongst “ bango ” reeds a few yards from the edge of Lake Nyasa. Sanderson has been informed by natives that during the rains (i.e. about January), ‘( at which time practically the whole country is under water,” the species is very prevalent in North Nyasa all over the grassy plain lying between the shore of the lake and a line of foot-hills some ten miles away, but this statement requires confirmation. Like G. palpalis and other species of Tsetse, G. hrevipalpis will occasionally follow cattle and other animals to some distance A l-set from its usual haunts; Stuhlmann states that in this way, especially during the hot weather, isolated specimens were forest on to frequently found for a time among the mountains of East for feeding. Usambara, at an altitude of from 800 to 1,000 metres (2,600 to “ seems read 3,250 feet), while, during the period from December to April, not set to w others have often been met with in settlements such as often exhib: Kwamkoro and Amani. Davey records the capture of a single individual (‘ on the finger of a native standing under a tree in the middle of a small village,” and in another village Sanderson once caught two specimens in a hut occupied by natives, and also containing a cow infected with trypanosomiasis. The present species of Tsetse is active and desirous of feeding only at two periods of the day--early in the morning before 8.0 o’clock, and in the evening, from about 4.0 P.M. onwards. The intervening hours, from 8.0 A.X. to 4.0 P.M., are passed by 6. brevipalpis in concealment, “ under leaves of bushes or in the A Tsetse-fly (Glo. grass, always near the ground ” (Sanderson), or low down on the with blood. trunks of trees, two or three feet from the base. While thus Bruce, C.B., resting motionless the Aies are difficult to discover and capture, and “ their presence would be entirely unsuspect’ed ” (Sanderson) ; that the ins at Kaporo, near the north end of Lake Nyasa, Davey found that suck ; they they preferred to rest on “ trees surrounded by creepers and quickness, ar undergrowth, and hid away in crevices in the bark or under the balloon with origin of branches.” Both Davey and Sanderson observed a pair flies of this in coit’(i on a tree-trunk, in the one case at midday, in the other hot and sul EVIPALPIS. BIONOMICS OF GLOSSINA BREVIPALPIS. 93 with in very large at about 5.0 P.M. ; attempts to find the pupae of this fly by 1 sea-level to some digging round the bases of various trees were however un- )undant shade and successful. Davey states that about 4.0 P.M. G. brevipalpis “ seems consisting of bush to emerge from its hiding place and settle on dried leaves, sticks es, either close to or dust on paths, apparently lying in wait for a meal ” ; the same rate within a few writer adds that he has noticed that “ game, on its way to water ocountered the fly in the evening, often stands for a time on emerging from the Davey records the ango ” reeds a few nderson has been ,e. about January), y is under water,” all over the grassy a line of foot-hills . lires confirmation. G. hwipalpis will Fig. 22. to some distance A Tsetse-fly (GZo8sina wwsitans, W&w., O), before feeding. (X 6.) hat in this way, specimens were forest on to a path, and would thus afford the tly an opportunity 3untains of East for feeding.” According to Davey, G. brevipalpis as a rule D metres (2,600 to “ seems ready to bite human beings in the evening, but it does scember to April, not set to work with the rapidity and voracity that G. morsitans tlements such as often exhibits.” When feeding captive flies, Stuhlmann finds tpture of a single g under a tree in village Sanderson by natives, and iasis. desirous of feeding e morning before 1.0 P.M. onwards. .&I., are passed by bushes or in the Pig. 23. A Tsetse-fly (Glossina morsitans, Westx;., o), after feeding, showing abdomen distended low down on the with blood. (x 6.) From a drawmg from life, kindly lent by Colonel Sir David ase. While thus Bruce, C.B., F.R.S., A.M.S. )ver and capture, ed” (Sanderson) ; that the insects generally wait for a while before deciding to >avey found that suck ; they then, however, drive in the proboscis with great by creepers and quickness, and the abdomen usually becomes swollen out like a lark or under the balloon with blood in from thirty to forty seconds. In captivity 1 observed a pair flies of this species show very little inclination to feed in very day, in the other hot and sultry weather, and this is also the case when the 94 BIONOMICS OF GLOSSINA l3REVIPALPIS. LARVA1

temperature is abnormally low.* Although continuing to bite and fly about (‘for some time after dark” (Sander-son), G. breci- pulpis appears not to bite during the actual night hours, and Accorc specimens kept in captivity in glass cylinders were found in of G. breve complete darkness to be perfectly still (Stuhlmann). Like G. pal- East Afri pnlis and blood-sucking flies in general, the present species quarter of displays a predilection for dark-coloured animals and surfaces. The larvae Stuhlmann states that he has frequently noticed that, when a 23” to 25” light- and a dark-coloured mule were moving side by side, only varied wit the latter was attacked, and Sanderson writes :-“ The flies bite mean intei through dark clothes, but have never been seen to sett’le on white and a hal: surfaces.” which how As to the relation of G. brevipalpis to game, Davey observes which fern that wherever he met with this fly in Nyasaland “ game the beginn (antelope, warthog, etc.) was abundant.” Once during the wet the extrue season, having shot two bush-pigs about sunset, on going up to mately reg the animals he found several G. brevipalpis settling upon the the hot th: carcases and “ apparently trying to suck blood ” from them, no reason although for some time previously he had searched for the flies continue t without success. height of It will have been noticed that, out of the 359 specimens of the develol this species examined by the present writer, only 4% were The ne females, and the enormous preponderance of males over females type and usually seen among captured specimens has been noted both by 9to 10 In Stuhlmann and Davey.t It is pointed out, however, by the exception former author that, as in the case of other flies, it is shown by colour, wit breeding G. brevipnlpis in captivity that the two sexes are segments 0 produced in approximately equal numbers. In order to catch the last or females, Stuhlmann states that it is necessary to use animals protuberan such as cattle and mules as decoys, but even so he observes that anterior po they are much more difficult to capture t’han males. He thinks tudinal rid; it probable that “ the ,gravid females are much more wary and is, as usual move about less than the males, and are consequently more rarely coarse or sl caught.” two more 0 divide it in * In order to keep captive flias alive, a meal of blood at intervals of from four to six days is necessary ; by feeding every fourth or fifth da;y, than in G Stuhlmann has kept G. brcvipalpis at Amani for considerable periods-m rounded). the case of individual females, for upwards of four months. t In German East Africa, fly-boys sent out by Stuhlmann into the Vosseler), n 3Ikulumusi Valley below Amani, brought back in one week over 1,200 width, shov G. bret+aZpis, of which only ‘JO were females. At Ksporo, near the north end of Lake Nyasa, out of 11’7 G. brczip@is caught in the evenings in being close August (dry season), all were males (Davey). On the other hand : “ Seven as in G. pc were captured at the same place and time of year about midday, and of these, four were males and three were females.” space, some WIPALPIS. LARVAL STAGE OF GLOSSINA Y,REVIPALI’IS. 95 continuing to bite REPHOIKJCTIOS AXIJ PKELIMNAHY STAGES. mderson), G. b~~ai- night hours, and According to Stuhhnann’s observations on female specimens :rs were found in of G. brevipalpis, kept by him in captivity at Amani, German tnn). Like G. pal- East Africa, the act of birth, though occasionally last’ing for a le present species quarter of an hour or more, generally takes place very quickly. mals and surfaces. The larvae produced by a female kept at a temperature of from ;iced that, when a 23” to 26” C. (73 * 4” to 77” I?.) were extruded at intervals, which side by side, only varied with the temperature fro& ten to twenty-two days, the 1 :-(( The flies bite mean interval being about twelve days ; thus in three months 1 t’o settle on white and a half a single female gave birth to eight larvae, two of which however were not viable. Throughout the period during ne, Davey observes which females of G. brevipalpis were kept at Amani (;.e. from Nyasaland “ game the beginning of September, 19q5, until the middle of May, 1906), ce during the wet the extrusion of larvae proceeded uninterruptedly at approxi- get, on going up to mately regular intervals, though these were somewhat shorter in settling upon the the hot than in the cold weather. Stuhlmann consequently sees tlood” from them, no reason why in nature the production of larvae should not erched for the flies continue throughout the year, though he suggests that the height of both the wet and dry seasons may be unfavourable to 0 359 specimens of the development of the pupae. er, only 42 were The newly extruded ~WVU, which is of the usual Glosvina males over females type and shape, and, as stated by Stuhlmann, measures from een noted both by 9 to 10 mm. in length by 2 to 3 mm. in width, is, with the however, by the exception of the hindmost segment, pale yellowish-white in kes, it is shown by colour, with the surface of the integument of the first eleven he two sexes are segments of the usual finely granular and shagreen-like texture ; In order to catch the last or twelfth segment, which bears the tumid lips or anal try to use animals protuberances, is, as in other Glossinrr larvae, deep black. The 30 he observes that anterior portion of this segment is encircled by a band of longi- males. He thinks tudinal ridges, and the outer surface of each lip or protuberance tch more wary and is, as usual, granular (though the granulations are not nearly so luently more rarely coarse or sharply defined as in the case of G. pnlpalis), and bears two more or less clearly marked grooves or furrows, which thus divide it into three prominences. The lips are actually smaller blood st intervals of , fourth or fifth day, than in G. palpdis and different in shape (not so full and miderable periods-in rounded). A dried larva of G. brevipalpis from Amani (Dr. F. onths. Stuhlmann into the Vosseler), measuring 9.5 mm. in length, by 4.45 mm. in greatest one week over 1,200 width, shows that the inner surfaces of the tumid lips, instead of :aporo, near the north lt in the evenings in being close together and separated by a deep and narrow notch, ) other hand : (‘ Seven as in G. pulpal&, are divided by a relatively wide and shallow aboht midday, and of space, something like a wide V. SSINA BREVIPALPIS: LARVA AND PUPA. G. BREVI

LL If,” writes Stuhlmann, (( the new-born larva be placed in a its tracheae glass dish or on blotting-paper, it crawls about for a time wings, whit exactly like an ordinary fly-maggot, after which it becomes ” - -- and the al stationary and soon contracts, its chitinous integument thickens The probos and darkens, and in about three-quarters of an hour it has is now str hardens. i to tell fron burrows into it, making a straight tunnel ; thus in one case a young, and, larva penetrated to a depth of-8 :5 cm. (31. inches). Under such blood greedi conditions, from an ho& and a quarter to an hour and a half It may elapsed before the change to the pupal stage was completed. In genesis may dry sand a larva did not burrow so deeply, since, as it burrowed, Stuhlmann f the sand continuallv fell in, but nevertheless it reached a depth of G. brev4 of from 2 to 3 cm (2 to l& inch). We may assume that- in larvae. nature the larvae behave in a similar way ; the fly will deposit its offspring on a spot which is sheltered and slightly damp, and the larva will at ‘once burrow beneath the surface.” As will The pupa (Figs. 5 A, 6 A, pp. 5, 7), which is of the usual black given above or clove-brown colour, measures about i.5 to 8 mm. in length, author in hi by 4 to 4.5 mm. in greatest breadth ; * the shape and dimensions recently cha of the tumid lips and the shape of the notch between them are of brevipalpis, 2 course the same as in the larva. At Amani, the duration of error has r the pupal stage was found by Stuhlmann to be from thirty to writers, all ( sixty-five days, according to the temperature ; when pupae were they have kept in the breeding-cage at 30” C. (86O F.) the flies emerged on Africa, have the average in about thirty-six days: The pupal stage appeared At the time to Stuhlmann to last somewhat longer in this species than in material at G. tachinoides. most part in Although a Gloss&a escapes from its pupa-case in precisely as to the chc the same manner as any other Muscid fly, it may be of interest however, th to quote Stuhlmann’s account of the emergence of t’he image of complain of 1 G. brevipalpk. (L The fly,” writes Stuhlmann, “bursts off the determinatio cap of the pupa-case, and protruding from the opening its As a matter cephalic vesicle,? forces its body-fluid into the latter, so that its that under body becomes sufficiently thin to be able to creep out of the confounded, puparium. Immediately after the insect has made its escape the name) the ; cephalic vesicle is retracted into the interior of the head and Mr. Newstea never used again. In a short time the young fly draws air into Together * So far as may be judged from an examination of seven specimens Austen (see 1 from Amani (Dr. Vosscler); since these are from larvae deposited in group (whicl 4 captivity, it is possible that in certain ca-es the dimensions are slightly subnormal. t Termed by Dipterists the piiZinw~z.-E. E. A. l * Austen, ‘I t CJ Auste VA AND PUPA. G. BREVIPALPIS: SYNONYMY AND AFFINITIES. 97 larva be placed in a its tracheae, and, as a result of so doing, expands greatly ; the ; about for a time wings, which until now have been folded, become spread out, * which it becomes and the abdomen swells to twice or thrice its original size. integument thickens The proboscis, which in the pupa was directed backwards, of an hour it has is now stretched out in front, and the chitin everywhere pupa. If, however, hardens. After from three to five hours it would be impossible np sand, it at once to tell from the appearance of the fly that it is still quite thus in one case a young, and, although on the first day reluctant to feed, it sucks mhes). Under such blood greedily on the second.” an hour and a half It may be noted that there is some evidence that paheno- was completed. In genesis may in exceptional circumstances occur in Glossina, since mce, as it burrowed, Stuhlmann states that at Amani on two occasions virgin females it reached a depth of G. brevipalpis, bred in captivity, produced fully developed lay assume that in larvae. the fly will deposit SYNONYMY AND AFFINITIES. 1 slightly damp, and rface.” As will have been seen from the references to literature is of the usual black given above, the species of Tsetse described and figured by the o 8 mm. in length, author in his “ Monograph ” as Glossina &sea* was in reality that iape and dimensions recently characterised by Newstead under the name Glossinn. xtween them are of brevipalpis, and not the true G. fusea, Walk.. This regrettable mi, the duration of error has naturally had the effect of misleading subsequent ) be from thirty to writers, all of whom with the exception of Newstead, in so far as ; when pupae were they have dealt with the Tsetse-flies of Nyasaland or East he flies emerged on Africa, have referred to the present species as Glossina. fussa. upal stage appeared At the time when the Monograph was written the Glossinn- his species than in material at the author’s command was both scanty and for the most part in poor condition,t so that to form correct conclusions la-case in precisely as to the characters of species was by no means easy ; since then, may be of interest however, the National Collection has had little reason to ice of t’he imago of complain of lack of specimens, and it is now possible to revise the n, “bursts off the determinations of eight years ago with a fair degree of confidence. a the opening its As a matter of fact, the author has for some time been convinced I latter, so that its that under the designation Glossinn fusca several species were ) creep out of the confounded, and he had actually drawn up (under another made its escape the name) the above description of G. breripnlpis in MS. before r of the head and Mr. Newstead’s paper appeared. : fly draws air into Together with Glossina lonigipennis, Corti, and G. medicorum, n of seven specimens Austen (see below, pp. 102, 98), G. brevipalpis forms a well-defined larvae deposited in group (which may be called the Glossina brevipalpis group) of mer)sions are slightly l * Austen, “ Monograph of the Tsetse-Flies,” pp. 95-97, Plate VI (1903). t CJ Austen, op. cit., “ Introduction,” p. vi. II 2 08 DESCRIPTION OF GLOSSINA MEDICORUM. G. MEDI generally large-sized species, the members of which may, as a second joil rule, be distinguished from those belonging to the Glossina fusca prominent: group by their much shorter proboscis and palpi; the wings too pollinose 8 are generally paler in the former than in the latter group. The brown at 1 most conspicuous differences exhibited by the present species as number ; p compared with G. longipennis, which it closely resembles in size, and darke have already been given in the diagnosis which precedes the yellow. description printed above. The characters by which G. medicorum Thorax may be distinguished from G. brevipalpis are point,ed out below seal-brown in the diagnosis and description of the former, whose area of grey ; pect distribution, so far at any rate as at present known, is quite Westiges tE different from that of G. brevipalpis.

Glossina medicorum, sp. nav. (Figs. 20, C, and 21, C, pages 86, 87, and Fig. 24, B.) S, ? .-Length, & (3 specimens) 9 to 10 mm., $’ (3 specimens) 10 to 10.6 mm. ; width of head, 6 2.8 to 3 mm., 9 3.2 mm. ; width of front at vertex, & 0.5 to 0.75 mm., ? just under 1 mm. ; length of portion of palpi projecting beyond margin of buccal cavity, & 2.5 to 2.75, 0 2.8 to 3 mm.; length of wing, 6 9.75 to 10.25, $’ 11.5 to 12 mm.

Medium sized, or in the $’ sex (which, 80 far as may be Budged from the three specimens of eac?L sex available for examination, considerably exceeds the 6 in size), fairly large species, allied and presenting a superficial resemblance to G. brevipalpis, NeLust., but Distal extremi distin,guished in both sexes by the wings having the appearance of sp. “0”. ( bein!! uniformly coloul;ed (the upper portion of the anterior trans- surface of verse vein, the posterior transverse vein, and the portions qf the surface bul jourth longitudinal vein, that in G. brevipalpis are conspicuously Abdomc infuscated, are not or scarcely at all infuscated in the present median art species), and in the 8 by well-marked diferences in the hypopygium a similar and hectors (cf. B and A, Fig. 24, p. 99). ochraceous pollinose ; Head buff, posterior surface greyish ; frontal stripe cinnamon hypopygiul or light mummy-brown ; ocellar spot inconspicuous; face and relatively lower half of posterior orbits shimmering whitish-yellow pollinose ; hectors bei bases of vertical bristles connected by a well marked dark brown spines, plal band ; antennae partly buff, partly brown or mouse-grey, outer surface of second and base of third joint buff, inner surface of MEDICORUM. G. MEDICORUM : HY POPYGIUM AND HECTORS. 99 of which may, as a second joint dark brown, third joint (,tip of which is moderately $0 the Glossina j&en prominent) greyish brown or dark mousegrey with a light grey mlpi; the wings too pollinose shimmer, arista buff, terminal joint fairly stout, dark I latter group. The brown at base beneath, branched hairs seventeen to nineteen in 3 present species as number ; palpi buff, mouse-grey on upper portion of outer surface, y resembles in size, and darker or even dark brown at tips ; proboscis bulb pale which precedes the yellow. which G. medicorum !Z’horaz.-Dorsum drab-grey, with the usual dark markings pointed out below seal-brown or light seal-brown and fairly distinct ; pleurae smoke- timer, whose area of grey ; pectus fawn-coloured, no dark blotches (or merely faint ent known, is quite ‘vestiges thereof, much reduced in size) on sternopleurae ; upper

nov.

Fig. 24, Ii.) to 10 mm., ? (3 I, 6 2.8 to 3 mm., 0.5 to 0.75 mm., of palpi projecting to 2.75, ? 2.8 to 1.5 to 12 mm. zr a8 may be judged ble for examination, re species, allied and nipalpis, Newst., but FIG. 24. Distal extremity of 8 abdomen of(a) Gloshabretipalpis, Newt., and (B) G. medicorum, lg the appearance of sp. nav. (ventral aspect), showing(h) hypopygia and (hect.) hect@x.* (X 15.) If the anterior trans- surface of scutellum brownish, impressed median line and under the portions qf the surface buff, apical scutellar bristles of ? of medium length. Bs are conspicuously Abdomen.-Dorsum dark brown, not banded, first segment, :ated in the present median area of second segment, except hind border, and sometimes es in the hypopygium a similar area on following two or three segments buff or ochraceous-buff, seventh segment in both sexes yellowish-grey pollinose ; hypopygium and ll&ors * of 8 as shown in Fig. 24 B ; da1 stripe cinnamon hypopygium buff pollinose, viewed from below broader (therefore spicuous; face and relatively if not actually bulkier) than in G. brevipalpis, Newst. ; ish-yellow pollinose ; hectors beset with relatively very coarse short bristles or minute marked dark brown spines, placed close together ; the group of minute spines forming r mouse-grey, outer luff, inner surface of * See p. 14 (first paragraph and note *).

MEDICOPLUM. G. MEDICORUM : DISTINCTIVE CHABACTEKS. 101 posite group by a bungalow at night” (Dr. W. M. Graham, W.A.M.S.); 1 8, tegument, whereas Odumase, Gold Coast, March, 1908 (G. C. Dudgeon); 1 9 , istles forming the Asaba, Southern Nigeria, 1895 (the late Dr. W. H. Crosse). erally intermingle, I as a rule partly BIONOMICS. lally more or less Nothing is yet known as to the habits of this species, although nora often with a from the note quoted above, attached to the specimen taken by L blotch on under Dr. Graham at Sekondi, it would seem that it sometimes moves ly with a more or about after dark, at any rate when attracted by light. nd middle, a trace on middle tibiae ; AFFINITIES AND DISTINCTIVE CHARACTERS. arsi, and last two As stated above, in the section dealing with the affinities of penultimate joint, the foregoing species, Glossina medicorum belongs to what may be termed the G. brevipalpis group, the third member of which micolorous, distal is G. longipcnnis, Corti. Attention has already been drawn to third, and upper, the distinctive characters of G. nredicorunz as compared with vein not (as in G. brraipalpis, Newst., and the former may at once be distin- tjacent veins, and, guished from G. longipennis, Co&i, by its smaller and narrower st the rest of the head, and by the absence of a dark brown ocellar spot, of a !d ; veins tawny- brown tip to the proboscis-bulb, and of the dark brown spots on the dorsum of the thorax characteristic of Corti’s species. ~evipalp’is. From Glossina fusca, Walk., G. medicorum differs in both ’ in the British sexes (apart from the wings, which are usually paler) in the n River, Liberia, front, viewed vertically from above, being (at least relatively) nt’ed by Surgeon wider, in the proboscis and palpi being shorter, and the latter (as seen from above) being broader and less slender, and in the above as typical, absence of a conspicuous dark blotch on each sternopleura; as ‘ies in which the regards characters confined to one sex, the 8 of G. medicorum is e specimens from distinguished from that of G. fusca by the minute spines or igeria, mentioned bristles forming the hectors being coarser, while in the 0 the tason to regard as apical scutellar bristles in the former species are stouter and shorter (about half the length of the corresponding bristles in G. fuscn 0 ). G. medicorum is distinguishable from G. tabani- . nav. *formis, Westw., which occurs in the same region, inter alia by the third joint of the antenna not being clothed with long and ge has been found fine pale hair, by the general coloration of the body being as a outhern Nigeria). rule less dusky, by the wings being paler or at any rate not so lowing specimens, brown, and by the anterior transverse vein being less coarse. been examined :-- . . McCloy, R.N.) ; .ght in P. W. D. 102 DMKXII’TIOS OF GLOSSINA LONGII’ENNIS. DESCR

Glossina longipennis, Corti. tips dark brown or 1 (Plate X.) Thoraz Glossina longipennis, Gorti, Ann. Mus. Civ. Genov., Serie 2”, Vol. XV faint, mou (XXXV), p. 138 (1695); Hough, Proc. $cad. Nat. Sci. Philadelphia, median lin 1898, p. 172; gusten, Proc. Zool. Sot. Land., 1900, p. 10, and “A Monograph of the Tsetse-Flies,” p. 101, Plate VII (1903); Brumpt, hind marg Comptes rendus des Seances dc la Socibte de Biologie, T. Iv, p. 1497 the diagnc (1903), and T. lvi, p. G73 (1904). markings brownish &, $! .---Length cJ (6 specimens) 10.2 to 11.6 mm., ? and a sim (3 specimens) 11.4 to 13 mm. ; width of head, 6 3.4 to 3.6 mm., admedian I 0 3.5 to 3.75 mm. ; width of front at vertex, 8 just under slightly f u1 1 mm., 9 1.2 mm. ; length of portion of palpi projecting beyond on the sutr margin of buccal cavity, 8 2.4 t’o 2.3 mm., 0 3 mm. ; length cream-buff of wing, (3 10.3 to 11.25 mm., ? 12 mm. blotch, tri, Resembling G. brevipalpis, Newst. (p. 85, Plate IX), in size scutellar 1 and general appearance, but disting2rishetl by the greater luidth of as those oj the.front in both sexes, by the ocellar spot being dark brown or clove- Abdom brown, and conspicuous, instead of merely light brwcunish and in- ochraceom conspicuous, by the pro6o;vcis bulb having a Sharply deJined brown or yellow, th dark brown. tip instead of’ beincr uniformly vale uellolv Cat anv rate Plate X, 1 except on the upper margins), nnd b!/ the dorsum of the thoraz side at b; exhibiting, in addition to a pair of admedian spots on the suture extending itself, four 8harply defined, dark brown, more or less oval or elon,gate angles of spots, arranged in a 2’arclllrlogrtrIrl: two i?l *front qf rtnrl two behind pollinose. the transverse xuturcj. Legs Ir hair cloth Head buff, posterior surface smoke-grey, yellowish-grey, or less blotcl drab-grey ; frontal stripe scarcely deeper in colour than frontal outer side margins (parafrontals), the latter, as well as the face and facial and hind pit, faintly shimmering whitish-yellow pollinose ; ocellar spot at distal conspicuous owing to its colour, ocelli small, of equal size ; brownish vertical bristles with a mummy-brown or light brown spot at with a da base of each, but not connected by a brown band ; first and corresponc second joints of anknnae cream-buff, second joint more or less tibiae, an (entirely 1 base ; las L D mummy-b of G. brevipaliis, Newst.), distal extremity broad ind prominent, Wings especially in 0, arista buff, terminal joint fairly stout and in 9 darker in somewhat short, dark brown at base beneath, branched hairs incrassatf about twenty-one in number ; palpi buff-yellow, not darker above, of fourth I 3II’ENNIS. DESCRIPTION OF GLOSSINA LONGIPENNIS. 103

tips dark brown ; proboxcis 6~111~ buff-yellow, distal third dark brown or brown. Thorax: isabella-coloured pollinose, dorsum with a narrow, erie 24, Vol. XV faint, mouse-grey or brownish, longitudinal stripe on each side of ci. Philadelphia, median line, dying away behind transverse suture before reaching 1, p. 10, and “A (1903) ; Brumpt, hind margin ; besides these stripes and the spots mentioned in gie, T. Iv, p. 149’i the diagnosis printed in italics above, the principal noticeable markings on the dorsum are a more or less distinct brown or 11.6 mm., $J brownish fleck on each humeral callus, near its upper margin, 94 to 3.6 mm., and a similarly coloured, elongate mark between this and the 6 just under admedian stripe ; the two dark brown spots in front of the suture jetting beyond slightly further apart than the two behind it ; the admedian spots 1 mm. ; length on the suture itself paler in colour than the other four ; scutellum cream-buff, upper surface with a mouse-grey or greyish-brown blotch, triangular in outline, on each side of middle line ; apical te IX), in maize scutellar bristles in ? short and stout (about one-third so long rester rdth of as those of 6 j. brown or clow- Abdonzera.--Dorsum ochraceous-buff, ochraceous, or tawny- omish and in- ochraceous, longer hair at base of second segment entirely golden- ejned brown or yellow, third to sixth segments inclusive each, as shown in )2u (at any rate Plate X, with a dark brown more or less lunate mark on each 2 of tile t1wm:tJ side at base, widely distant from median line but not always B on the suture extending quite into basal angle, seventh segment and posterior ma1 or dongate angles of third to sixth segments inclusive yellowish-grey marl trco behind pollinose. Legs buff; row of bristles at base of front coxae ochraceous, hair clothing front coxae below golden-yellow ; femora more or lowish-grey, or less blotched or streaked with lighter or darker mouse-grey on r than frontal outer side, front femora similarly infuscated on inside, middle face and facial and hind femora flecked with mouse-grey or brownish on outside ; oeellar spot at distal extremity, middle femora with usual dark brown or )f equal size ; brownish blotch on under side near tip; middle tibiae sometimes brown spot at with a dark blotch on outer side beyond middle, and a trace of a md ; first and corresponding blotch sometimes visible on hind tibiae also ; hind t more or less tibiae, and sometimes middle tibiae also, with a faint trace wnish-grey, or (entirely wanting in some specimens) of a dark brown ring near Se of fine and base ; last joint and tip of penultimate joint of middle tarsi 1 fringe in case mummy-brown, last two joints of hind tarsi clove-brown. md prominent, Wings light drab or pale isabella-coloured ; veins pale tawny, tout and in ? darker in places, upper portion of anterior t’ransverse vein strongly branched hairs incrassate, anterior transverse vein together with adjacent portion ; darker above, of fourth longitudinal vein lying towards base of wing, as well as 104 DISTRIBUTION OF GLOSSINA LONGTPENNIS. G. LONG1

posterior transverse vein and portion of fourth vein immediately disseminate beyond, dark brown. Sqzcamae semi-hyaline, border buff, fringe d‘ aino,” Gl( pale yellow. Halter-es cream-buff. him from ; specimen 0‘ DISTRIBUTIOX OF G. lonyipen&, Corti. as mention Biermuddo Glossina longipennis is an East African species, the range of which is apparently somewhat restricted, since up to the present time this Tsetse-fly has been recorded only from Somaliland and the East Africa Protectorate. Since, however, it is found on No obs the line of the Uganda Railway, it is possible that its area also have yet be ext’ends into German East Africa, while to the north it perhaps that the nc occurs, as Dr. Andrew Balfour has pointed out,* in Southern Woosman i Abyssinia. Towards its southern boundary the range of supra), shov G. longipennis overlaps that of G. brevipalpis, Newst. Whether and G. palli G. longipennis in places where it occurs is found only in small Railway.” numbers the author is unable to say; in any case, very few specimens of it have so far reached the British Museum (Natural History). The following are the particulars with reference to Not yet the nine specimens of this species at present contained in the National Collection. Somaliland : 3 6 6, 1 Q , precise locality unknown, 1894 (Tlr. Greu$eZd). East Africa Protectorate : 1 8, ? Kiboko Glossina RiT-er, or Sabaki River near the contluence of the Tsavo River, brevipalpis 1898 (the lute Vet.-Capf. A. J. Haslum); 1 6, 1 Q, Uganda known Tset Railway (the ‘6 taken at Kibwezi Station, ‘( in thorny bush “), brown spot 1903 (Captain E. D. 157 G&g, IX.I.X.,-presented by Colonel defined bro Sir David Bruce) ; 1 6, Lake Baring0 (Captain R. Ford) ; prevent it f 1 Q , Uganda Railway, Tsavo River, 6,000 feet, 12. x. 1910, yet describe “ came into railway carriage at night ” (R. B. Woosnam). Newst., the In addition to the foregoing, the author has also had the be confusec opportunity of examining :-1 & (the type of the species), from above. the River Uelmal, Boran Galla Country, N.-E. Africa, June, 1893 (Captain Pittorio Boffego), lent by the Museo Civic0 di Storia Naturale di Genova, through the courtesy of Dr. R. Gestro; 1 Q , from West Somaliland, 23-25. vi. 1895 (C. V. A. Peel), lent by the University Museum, Oxford, through the c kindness of Professor E. B. Poulton, F.R,.S. Brumpt 1 states that in the Ogaden country, Somaliland, where both the flv and a trvnanosomiasis of camels and mules d 4I * Second Report of the Wellcome Research Laboratories at the Gordon 1\Iernorial College, Khartoum, p. 31 (1906). t LOG cit. LONGIPENNIS. G. LONGIPENNIS : DISTINCTIT’E CHARACTEIiS. 105 ath vein immediately disseminated by it are known to the natives by the name i, border buff, fringe <‘ aino,” Gloss&a longipennis was the only Tsetse encountered by him from July to October, 1901. A field-note attached to the specimen obtained in West Somaliland by Mr. C. V. A. Peel, , Corti. as mentioned above, says :-(< Fly-belt sharply defined from L Biermuddo to Boholo Deno.” species, the range of ce up to the present ram Somaliland and BIONOMICS. Iver, it is found on No observations on the bionomics of Glossina longipennis de that its area also have yet been placed on record. It may be pointed out, however, ;he north it perhaps that the note attached to the specimen taken by Mr. R. B. 1 out,” in Southern Woosman at the Tsavo River, East Africa Protectorate (vi& lary the range of .qra), shows that the present species, like G. brevipalpis, Newst., I, Newst. Whether and G. pallidipes, Austen, enters railway carriages on the Uganda found only in small Railway.* any case, very few #h Museum (Natural PRELIMINARY STAGES. es with reference to Not yet observed. ’ at contained in the

AFFINITIES ANU DISTINCTIVE CHARACTERS. .ity unknown, 1894 B: 1 8, ? Kiboko Glossina longipennis, Corti, which belongs to the Glossina of the Tsavo River, brevipalpis group, is one of the most easily recognisable of the 1 6, 1 9, Uganda known Tsetse-flies, since the characteristic and distinctive dark ” in thorny bush “), brown spots on the dorsum of the thorax, and the sharply mesented by Colonel defined brown or dark brown tip to the proboscis bulb suffice to ICaptain R. Ford) ; prevent it from being mistaken for any other species of its genus . 1 feet, 12. x. 1910, yet described. Further points of difference from G. brevipalpis 1. WoosnanL). Newst., the only species with which G. longipennis can possibly lr has also had the be confused, are mentioned in the diagnosis printed in italics If the species), from above. Y.-E. Africa, June, * See also p. 62. le Museo Civic0 di courtesy of Dr. R. t 1. vi. 1895 (C. V. A. mxford, through the ountry, Somaliland, f camels and mules )ratories at the Gordon Glossilza, nor bounds1 INDEX. - bocagei, & palis VI __ brevipalpl desorip head c Abdomen in Tsetse-flies, concealed diagram showing nomenclature of ferns by wings in resting position, 1; of, 11 and ab distension of, after a meal of Coloration of Tsetse-flies, 1 gium ( blood (figure), 93 Committee, Entomological Re- wings, Xino, a native name in the Ogaden search (Tropical Africa), en- localitil country, Somaliland, for Glos- quiry by, concerning habits, in Mm sina longipennis and a trypano- etc., of Glossina morsitans, x than 1 somiasis of camels and mules Cover essential to Tsetse-flies ; bionom disseminated by it, 105 nature of, 4 duction Alexander. Dr., observations by, stages on Glos.&a tachinoides in S. Dalziel, Dr. J. McE., on Glossina vations Bornu! N. Nigeria, 44 tachinoides and G. nwrsitans in depth Anatomy~(mterns1) of Glossirza pal- Yola Province, N. Nigeria, 44 rows, 9 p&s, reference to paper by Davey, J. B., observations by, on of, 5, 7 E. A. Minohin on. 26 (note) Gloss&a breviaalvis in the fly wht $ntennse in Tsetse-flies, description Nyasaland Pro&&rate, 91-94 emergh of, 10; figures of, in various Densham,. the late Dr. W. A., on case (S species, 12, 31, 36, 70, 76, 62 Glossznu pallidipes in the Nile tiq& Arabia (Southern) occurrence of Province, Uganda Protectorate, genesis Gloss&a tachinoides in, 6,41,46 61 affinitit Arista, explanation of term, 12 (at Division of Tsetse-flies into four from end of description of Fig. 8) ; Groups, 18 from c figures of, in various species, Dudgeon, G. C., on Glossina tachi- (figure) 12, 31, 36, 70, 73, 32 noides and G. palpalis in - brevipalpi Nigeria, 43 grave SI Bsgshawe, Dr. A. G., discovery by, in conr of pupa and puparia of Gloss&a Entomological Research Committee Sickner fusca in Uganda Protectorate, (Tropical Africa), enquiry by, Luangl in 1906, 73 concerning habits, etc., of GZos- Rhodes Bionomics, account of in author’s sina morsitans, x __ brevipalpi “Monograph of the Tsetse- External characters of Glossina, maun i flies” related chiefly to Gloss&a diagram showing nomenclature Germal mowitans, x of, 11 of four Blood (vertebrate) essential for con- a meal tinued existence of Tsetse-flies, “ Fly-belts,” explanation of term, or fifth 4 4 ; factors determining limits - brevipalpi Branched hairs (on arista),, 12 of, 4 determ Brumpt, Dr. E., on Glosszna tachi- E’ront, explanation of term as used Table i noides in the Shari River basin in Dipterology, 11 (note) species and on the shores of Lake Frontal stripe, explanation of term, descrip Chad, 45 11 (note) longing Buffalo (Bubalzs ccL$L’r), alleged - caliginea, saecial deoendence of Glossina Geographical distribution of Tsetse- head, 3 &or&am *on ; evidence to B flies, 6 31; the certain extent contradictorv. Glossina, external characters of hww genus, 9 (diagram showing no- wings, menclature of, 11); head, 9; bionom antennae, 10; proboscis and distinct palpi, 12; thorax, abdomen, distingl and hypopygium, 13 ; hectors, palis, 3 14 (and note * ); legs and cera, 34 wings, 14 ; venation, 15 ; chae- _ decor&, totaxy (diagrams of thoracic, tachino 16, l-l), 15-17 - f uscn, desl INDEX. 107

Glossina, northern and Isouthern and figure of antenna, 70; boundaries of genus, 6 figure of head of male, 86, __ bocagei, a synonym of G. pal- of head of female, 87; thorax, palis var. tucllmani. 26 abdomen (figure of hypo- __ brevipalpis, synonymy and pygium of male,, 76), legs, descriptron of, 85 ; figure of and wings, 71; dmtribution, sm showing nomenclature head of male, 86, of head 72 ; localities, etc., of speci- of female, 87 ; head, thorax, mens in Museum, 72; bio- n of Tsetse-flies, 1 and abdomen, 38; hypopy- nomics, 73; pupa (figures Entomological Re- gium (figure, 99), legs and of, 5, 7), 73, distinguished 2’ (Tropical Africa) wings, 69 ; distribution, 90 ; from that of G. brevipalpis, by, concerning habF;i localities, etc., of specimens 73 ; affinities and distinctive Df Glossina morsitans, x in Museum from elsewhere characters, 74 ; distinguished ssential to Tsetse-flies ; than Nyasaland, 90, 91 ; from G. nigrofusca, 70, 74, :e of, 4 bionomics, 91-94 ; repro- from G. tabaniformis, 74, duction and preliminary from G. fuscipleuris, 74, 75, 3.. J. McE., on Glossina stages (Stuhlmann’s obser- 76 (figure) noides and G. mowitans in vations), 95; larva, 95; Gloss&a fusca Group, Table for de- Province, N. Nigeria, 44 depth to which larva bur- termination of? 19,20; Table B., observations by, on rows, 96; pupa, 96, figures for determinatron of species ‘ina brevipalpis in the of, 5, 7; behaviour of the belonging to, 21: 69; de- aland Protectorate, 91-94 fly when and shortly after scriptions of speoles belong- emerging from the pupa- the late Dr. W. A on I ing to, 69-84 &a pallidipes I the’&le case (Stuhlmann’s observa- , _ fascipes, not included in Table, me, Uganda Protectorate, tion& 96, 97; partheno- 20 (note), 23 (note) ; copy genesm, 97; synonymy and of original description of, of Tsetse-flies into four affinities, 97 ; distinguished 29 : habitat. 30 P, 18 from G. Zongipennis, 86, - fusci&uris, description of, 75 ; G. C., on Glossina tachi- from G. medicorum,, 98, 99 head, thorax, and abdomen, r and G. palpalis in (figure) ia, 43 - brevipalpis (and G. morsitans), grave suspicion attaching to, gical Research Committee in connection with Sleeping distinctive characters: 77 ; ical Africa), enquiry by, Sickness in Nyasaland and distinguished from G. fusca, rning habits, etc., of Glos- Luangwa R. Valley, N.-E. 75, 76 (figure), 77, from G. no&tans, x Rhodesia, ix nagrofusca and G. tabani- characters of Gloss&a, ~ brevipalpis kept alive by Stuhl- formis, 77 brn showing nomenclature mann in captivity at Amani, - longipalpis, description of, 63 ; German E. Africa, upwards head, thorax, abdomen, legs, of four months, by supplying and wings, 64; figure of ” explanation of term a meal of blood every fourth hypopygium of male,.51 ; dis- ?tors determining limit; or fifth day, 94 (note) tribution, 64; localitms, etc., __ byevipalpis Group, Table for of specimens in Museum, 65 ; llanation of term as used determination of 19 20. bionomics, 66 ; reproduction nerology, 11 (note) Table for determInatibn 0; and preliminary stages, 67 ; ripe, explanation of term, species belonging to, 22, 85 ; affinities and distinctive Ite) descriptions of species be- characters,67; distinguished longing to, 85-105 from G. pallidipes, 64, 68, cal distribution of Tsetse- - caliginea, description of, 30 ; from G. morsitans, 67, 68 5 head, 30 ; figure of antenna, --- longipennis, figure of, in rest- external characters of 31; thorax, abdomen, and ing attitude (partly dia- 9 (diagram showing no- hypopygium, 32; legs and grammatic), 2 ; description Lture of, 11); head, 9 ; wings, 33 ; distribution, 33 ; of, 102; head, 102; thorax, oae, 10 ; proboscis and bionomics, 34; affinities and abdomen, legs, and wings, 12 ; thorax, abdomen, distinctive characters, 34 ; 103 ; distributron (including ypopygium, 13 ; hectors, distinguished from G. pal- localities, etc., of specimens palis, 30, 34, from G. palli- in Museum), 104 ; enters cera,, 34 carriages on Uganda Rail- __ decorsea, a synonym of G. way, attracted by lamplight, tachinoides, 39 62, 105; affinities and dis- - fusca, description of, 69 ; head tinctive characters, 105 ; 10s INDEX.

distinguished from G. bye&- determination of, 19 ; Table tee palpis, 102, from all other for determination of species Va species of Glossina yet de- belonging to? 21,48 ; detcrip- Glossina 2 scribed, 105 tions of species belonging to, lie1 Glossina mac&zta, a synonym of 48-68 G. palpalis, 24, 28 Glossina nigrofusca, synonymy and - palpa - medicorum, description of, 98 ; description, 77; head (with tin, head, 98 ; thorax and abdo- figure of antenna) and pal men (figure of hypopygium thorax, 78; abdomen, legs, 26; and hectors of male), 99 ; and wines. 79 : distribution. - palpa legs and wings, 100; pale 79 ; loc&ies,‘eto., of speci- wit form,, 100 ; distribution (in- mens in Museum, 80; syn- ing cludmg localities, .etc., of onymy, affinities, and dis- mir speOimens in Museum), 100 ; tinctive characters, 80 ; dis- det affinities and distinctive cha- tinguished from G. fusca, lon racters, 101; distinguished 77, 80, from G. tabaniformis, tior from G. breoipalpis, 98, 99 81 24- (figures of hypopygia and - submc hectors),from G. longipennis, - pallkern, description of, 35 ; head and thorax, 35; abdo- of ( G. fusca, and G. tabani- tint formis, 101 men, legs, wings, and figure of antenna, 36 ; distribution 52 ~ morsitans, description of, 48; - taban head and thorax, 49; sbdo- and bionomics, 37 ; affinities and distinctive characters, des men,legs, and figure of hypo- fi&T pygium, 50; wings, 51; dis- 38 ; distinguished from rest of Glossina paZgaZis Group, rax tribution, 52 ,; bionomics, wir 53-56 ; alteration in appear- 38 - pallidipes, the species used clu ance of abdomen (as in other w species of Glossina) produced by Sir David Bruce in Zulu- bio by a meal of blood, 93 land, in 1895-96, 56; de- dis (figures) ; reproduction and scription of, 58; head, 58 ; disi preliminary stages, 56; pupa- figureof antenna, 12; thorax, fus case (description of), 57, abdomen, legs, and wings, ani (figures of), 5, 7; affinities 59 ; distribution, 59 ; locali- ties, etc., of specimens in - tachi? and distinctive characters, des 57 ; distinguished from G. Museum, 60 ; bionomics, 61; tho pallidzpes and G. longipalpis, pupa (figures of), 5, 7, dis- wir 57 tinguished from that of G. IOCl - morsitam, enquiry by Ento- mowitans, G. brevipalpis, in mological Research Com- and G. fusca, 62; affinities mittee (Tropical Africa) con- and distinctive characters, cerning habits, etc., of, x 62; distinguished from G. anti - morsitans (and G. brevipalpis), nzorsitans, 62, 63, from G. figr grave suspicion attaching to, longipalpzs, 58, 63 and in connection with Sleeping - pulpalis? synonymy and de- affi: Sickness in Nyasaland and scriptron of, 24; head and cha Luangwa R. valley, N.-E. thorax, 25; abdomen, legs, fro] Rhodesia, ix and wings, 26 ; figure of an- Grc - nzowitam, name “ Tsetse ” tenna, 31; distribution and G. : originally applied to this bionomics, 27; figures of Graham, 1 species,.x pupa, 5,7 ; synonymy, affini- by. on - morsitans, occurs at an altitude ties, and distinctive charac- Ashanl of 5,000-5,500 ft. in Kasempa ters of, 28; distinguished Groups (fo District,N.-W.Rhodesia,55; from G. ftmcipes and G. cali- flies ar original specimens of, found ginea, 2S, from G. pal&era determ by Vardon and Oswell in N. and G. tachinoides, 29; in- Transvaal (Siloquana Hills), ternal anatomy of (reference Haematopo, in 1845, 54 to paper by E. A. Minchin), most 1 - morsitam Group, non-associa- 26 (note) Glossil; tion with water of species- pulpalis, believed to be non- when a belonging to, 4; Table for existent in Nyasaland Pro- - distin INDEX. 109 termination of, 19 ; Table tectorate and Lusngwa R. (Glossinn), 3 ; figure of, in rest- * determination of species Valley, N.-E. Rhodesia, ix ing attitude. 3 longing to? 21,48 ; derorip- Glossina palpalis until lately be- Hall, P. E., observations by, on ns of specres belonging to, lieved to be sole dissemina- Gbssina morsitms in the Lun- -68 tor of Sleeping Sickness, ix dszi District, N.-E. Rhodesia, kigrofusca, synonymy and - palpalis var. zuellmani, dis- 55 gcription, 77; head (with tinguished from typical G. Harger, R. L., observation by, on ure of antenna) and palpalis, 26 ; description of, Gloss&a morsitalzs (extrusion max, 76; abdomen, legs, 26 ; occurrence of, 27 of larvae), 56 d wings, ‘79 ; distribution, - palpalis Group, association Haunts of Tsetse-flies, 4 ; localities, eta., of speci- with water of species belong- Hectors, explanation of term, 14 ing to, 4; Table for deter- (first paragraph, and note *) sns in Museum, 80; syn- < gmy, affinities, and dis- mination of, 19; Table for Hypopygium (a knob-like protuber- otive characters, 80 ; dis- determination of species be- ance beneath end of male abdo- guished from G. fibsca, longing to, 20, 23 ; descrip- men, affording means of dis- 80, from G. taba~z@~ormis, tions of species belonging to, tinguishing sexes of Tsetse- 24-47 flies), 1; characteristic shape - zm, description of, 35; submorsitans, a form or race of, 13 ; explanation of term, 13, td and thorax, 35; abdo- of G. morsitans--not a dis- notet; figures of, in different n, legs, wings, and figure tinct species, 21. (note), 51, species, 50, 51, 76, 99 antenna, 36 ; distribution 52 Jack, R. W.,, note by, on discovery I. bionomics, 37 ; affinities - tabaniformis, synonymy and of pupa-caseb-case of of Glossina Glossina morsi- d distinctive characters, description of, 81 ; head, 81; tans in S. Rhodesia, 57 ; distinguished from rest figure of antenna, 82; tho- rax, abdomen, and legs, 82 ; Kinghorn, Dr.>r. A.,A., on Glossina tachi- Glossina pal&is Group, wings, 83; drstribution (in- wides in Ashanti (Western and cluding localities. etc.. of Northern Provinces), 43, 44; iipcs, the species used specimins examined), ‘and on G. Zor,gL&@s in Ashanti Sir David Bruce in Zulu- bionomics, 83 ; affinities and ~ (Western Province), 67 ; on .d, in 1895-96, 56; de- distinctive characters, 84 ; ~ G. fusca in Ashanti, 73 iption of, 58; head, 58 ; distinguished from G. nigl-o- lreof antenna,lZ; thorax, Lake Nyasa, Glossi?ra palpalis fusca, 81, 84, from G. fzLsca ~ apparently absent from, 27 lomen, legs, and wings, and G. fusciplezrris, 84 ; distribution, 59 ; locali- Larva of a Tsetse-fly, general de- - tachinoides, synonymy and scription of, 4; of Glossinn 3, etc., of specimens in description of, 39 ; head, 39 ; Iseum, 60 ; bionomics, 61; brez;ip&is (Stuhlmann’s obser- thorax, abdomen, legs, and vations), 95, 96 pa (figures of), 5, 7, dis- wings, 40 ; distribution, 40; Length of Tsetse-flies, 1 guished from that of G. localities, etc., of specimens rsitans, G . bwvipalpis, in IIuseum, 41-43; in Ni- Map, explanation of, 6, 8 !. G. fuscn, 62; sfhnities geria, 43; in S. Arabia, 46 ; Milne, Dr. A. D. (P.M.O., E. Africa l distinctive characters, bionomics, 44 ; reproduction Protectorate), statement by, distinguished from G. and preliminary stages, 46 ; that Glossina pallidipes, G. rsztans, 62, 63, from G. figures of pupa, 5, 7 ; larva brevipalpis, and G. longipennis, gipalpzs, 58, 63 and pupa, 46,47 ; synonymy, attracted by lamplight, fre- :lis~ synonymy and de- affinities, and distinctive quently enter carriages on the iptlon of, 24; he&d and charaoters,47; distinguished Uganda Railway, 62 Irax, 25; abdomen, legs, from Gloss&a morsitans Monograph of the Tsetse-Flies 1 wings, 26 ; figure of an- Group, and from rest of ; (E. E. Austen), ix; number of .na, 31 ; distribution and G. palpalis Group, 47 speciesdescribed and illustrated nomics, 27; figures of Graham, Dr. W. RI., observations ~ in, ix, x ?a, 5,7 ; synonymy, affini- by. on Glossina pallicera in Neave, Dr. Sheffield, observation I, and distinctive charac- Ashanti, 37 by, on Glossina morsitam en- 3 of, 28; distinguished Groups (four) into which Tsetse- ~ tering villages in Kstanga Dis- m G. fuscipes and G. cali- flies are divided, 18; Table for trict, Congo Free State, 55 ea, 28, from Q. @Zicera determination of, 19 Neave, S. A., observation by, on 1 G. tachinoidcs, 29; in- apparent drslike-. to immediate nal anatomy of (reference Hanzatopota and Stomoxys, genera vicinity or water evmced by paper by E. A. Minchin), most likely to be mistaken for Glossina morsitans on south- (note) Gloss&u, 2,3 ; position of wings west shore of Lake Nyasa, and Jis, believed to be non- when at rest, 3 on Luangwa R., N.-E. Rho- stent in Nyasaland Pro- - distinguished from Tsetse-flies desia, 54 INDEX.

Newstead, R., description by, of Sanderson, Dr. M., observations by, Glossina fuscipes, 29 on Gloss&a brevipalpis in the Number of species of Tsetse-flies Nyasaland Protectorate, 91, 92, recognised in present volume, x QA Nyasa, Lake, Glossina yalpalis Sexes-of Tsetse-flies, how distin- apparently absent from, 27 guished, 1 Sleeping Sickness, until lately be- Parthenogenesis, twice observed by lieved to be conveyed solely by Stuhlmann in Glossina brevi- Glossina nalnalis : reasons for palpis, at Amani, German E. suspecting G. m&sitans and Africa, 97 G. brevipalpis of acting as Pigs (wild), blood of, sucked by carriers in Nyasaland and Glossina morsitans, 55 (note), Luangwa R. Valley, N.-E. and G. brevipalpis, 94 Rhodesia, ix Pirie, Dr. G. J., observations by, Stomoxys and Haematopota, genera on Gloss& ta&no&s in N. most likely to be mistaken for Bornu, N. Nigeria, 45; on G. Glossina, 2,3; position of wings lonyipulpis in Bassa Province, when at rest, 3 N. Nigeria, 66 StomoxysT distinguished from Tse- Pollard, Dr. J. McF., observations tse-fires (Glossina), 3 ; figure of, by, on Glossina tachinoides in in resting attitude, 3 Bauchi Province, N. Nigeria, Stuhlmann, Dr. F., observations 45, 46 by, on Glossina brevipalpis in Preponderanceof males over females German E. Africa, 91-9’7 among captured specimens of Glossina brevipalpis, 94 (and Tsetse, name originally applied to note) Glossina nwrsitans, x Proboscis of Tsetse-flies, reference Tsetse-flies (see also Gloss&z) : to paper by Stephens and New- - abdomen of, concealed by wings stead on structure of, 26 (note) in resting position, 1 Prophylactic measures (in fight - ooloration and length of, 1 against Tsetse-flies) not referred - general characters of, and dis- to in present volume, x tinctions from other flies, 1 Ptilinum, 96 (note) - number of species recognised Pupa (puparium) of a Tsetse-fly, in present volume, x general description of, 6 ; - haunts of, 4 figures of in various species, GJossina brevipalpis, G. fusca, Venation, distinctive characters of, . G. morsitans, G. pallidipes, G. in Glossina, 14; figure showing palpalis, G. tachinoides, 5, 7; nomenclature of, 11; descrip- pupa (puparium) of Glossina tion of, 15 tachinoides, 46, of G. morsitans, Villages (native), often infested by 56, 57, of G. pallidipes, 62, of Glossina morsitans, 55 G. fusea, 73, of G. brevipalpis, 96 Water, association with, of species Puparium, explanation of term, 6 belonging to Glossina pal- (note) palis Group, 4 ; dislike to im- mediate vicinity of, evinced Railway carriages on Uganda Rail- by G. morsitans in certain way, E. Africa Protectorate, parts of Africa, 54 entered by Glossina pallzdipes, Wellman, Dr. F. Greighton, note on G. breuipalpis, and G. longipen- Glossinn palpalis var. wcll- nis (statement by Dr. A. D. mani, in Angola, 27 Milne, P.M.O.), 62 Wings of Tsetse-flies, distinctive Reproduction, mode of, in Tsetse- venation of, 14 ; position of, flies, 4 when at rest (enahling specs- Roubaud, E., on Glossina tachi- mens of these insects to be noides in Central Dahomey, 45, distingnished from other 46; on G. longipalpis in Da- blood-sucking flies liable to homey, 66, 67 be mistaken for them), 2