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PRESENTED
BY
Ghe Trustees

OF

THE BRITISH MUSEUM,

———

ane pl
on A MONOGRAPH
a 7 OF THE

Wien TSETSE-FLIES

[GENUS GLOSSINA, WESTWOOD]

BASED ON THE COLLECTION IN THE
BRITISH MUSEUM

BY

ERNEST EDWARD AUSTEN

WITH
A, CHAPTER ON MOUTH-PARTS
BY H. J. HANSEN, PHIL. DOC.

LONDON
PRINTED BY ORDER OF THE TRUSTEES
SOLD BY
Lonemans & Co., 39, PaTERNOSTER Row, E.C.

B. QuaritcH, 15, Piccapinty, W.; Dutau & Co., 37, Sono Squarn, W.
Keean Pau, Trencu, Trisner & Co., Cuarina Cross Roap, W.C.
AND AT THE
Britis Musrum (Narurat History), CromwExLL Roap, S.W.

1903
All rights reserved

LONDON:
PRINTED BY WILLIAM CLOWES AND SONS, LIMITED,-
DUKE STREEY, STAMFORD STRENT, S.E., AND GREAT WINDMILL STREET, W.

CARDED
DIV. OF INSECTS

‘

Hv. Ise
U. Se NATLe MUS»:

Ze institi tien

KS

aus?

National *

PREFACE.

a oe

Tue Trustees of the British Museum have authorised the
preparation and publication of the present work in view of
the great practical importance of an accurate knowledge of the
genus of flies to which Wiedemann gave the name Glossina. The
discovery by Colonel Bruce of the fact that the common Tsetse-fly
of South Africa produces the death of horses and cattle rightly
ascribed to its attacks by introducing into the blood of its
victims a minute parasite, the Trypanosoma Brucei, has been
followed by the discovery of similar parasites in the Indian
disease known as Surra, in the Mal de Caderas of South America,
in the Dourine of Algeria, and, lastly, in the blood of human
subjects. The last and most important discovery is that by
Castellani of the occurrence of a Trypanosoma in the cerebro-
spinal fluid of nearly seventy per cent. of the cases of sleeping
sickness examined for this purpose by him at Entebbe, Uganda,
in the early part of this year. It is estimated that thirty
thousand of the native inhabitants of the Uganda Province have
died of the sleeping sickness since its sudden appearance among
them two years ago.

We are not yet sufficiently acquainted with the facts as to
the distinguishing characteristics of the different species of
Trypanosoma concerned in causing these diseases. Nor do we
know in any case, excepting that of the Nagana disease, what
is the nature of the insect (if insect it be, as is probable) by
which the Trypanosoma is introduced into the blood of a pre-
viously healthy animal or man. In the case of the Nagana
disease, we know that a Tsetse-fly — Glossina ‘morsitans of
Westwood—is the habitual and specific carrier of thé parasite.
Even though other blood-sucking insects may occasionally act as
intermediaries and pass on the Nagana parasite from one animal
to another, it is to Glossina morsitans, and possibly also to

a 2

PREFACE.

G. pallidipes, that the disease owes its prevalence in special
localities, and it is apparently these species which Colonel Bruce
used in his experiments.

The question immediately arises as to whether the other
species of Glossina are carriers of disease germs; and whether
other species of Trypanosoma, especially that found infesting the
human blood and cavitary fluids, are carried by species of
Glossina or Tsetse-fly.

An accurate knowledge of Tsetse-flies is clearly indispensable
for further progress in this inquiry.

Inasmuch as the genus Glossina is found. only in Africa, it
is certain that in India and in South America other carriers of
the Trypanosoma must be at work. At present, though it
appears that Stomoays and some other “ biting flies,” not scienti-
tically determined, have been experimentally shown to be capable
of carrying the Trypanosoma of Surra and of Nagana from one
animal to another, no constant association of any genus or
species of fly with these Trypanosoma-diseases has been sug-
gested, excepting that of Glossina with the African Nagana
disease.

In view of the fact that the Trypanosoma parasite is not
observed to undergo any developmental changes within the
Tsetse-fly similar to the changes which the malaria-parasite
Laverania undergoes in the body of the gnats of the genus
Anopheles, it is not unreasonable to suppose that there is no
such exclusive bond of association between Glossina and Trypa-
nosoma as there is between Anopheles and Laverania. Other
carriers may serve for Trypanosoma, though Laverania must
have Anopheles and no other.

On these points we require further observation. Collections
from all parts of the world of flies (and other insects) which suck
the blood of human beings and other animals are needed, accom-
panied by careful notes as to habits, locality, and life-history.
Such collections will be received with special welcome, and at
once investigated if addressed to me, here. It is clear that an
accurate and comprehensive study of blood-sucking organisms,
in view of their possible action as carriers of disease, has become
a matter of the most urgent public importance.

As an illustration of the significance of the discrimination of
the different species of blood-sucking flies, 1 may mention that
whilst these lines are in the press, I have received from Colonel
Bruce a small box of biting flies taken at Entebbe, Uganda,

PREFACE.

where sleeping sickness is now rife. Among these Mr. Austen
has determined one horse-fly (Tabanus sp. incert.), and ten
Tsetse-flies (five males and five females) of the species Glossina
palpalis, Rob.-Desv.

This particular species of Tsetse-fly is essentially a West
African species, known from the Gambia to the Congo. It
certainly suggests the need for an inquiry into the possible con-
nection between this fly and the sleeping sickness, when we
remember that that disease has been established for years on
the West Coast of Africa, but was unknown in Uganda until
two years ago. Sir Henry Stanley met with “Tsetse-fly ” for a
long distance on the Upper Congo, and it is suggested by Mr.
Austen that Glossina palpalis reaches Uganda by way of the
valleys of the Congo and Aruwimi.

E. RAY LANKESTER.

British Musrum (Natrurau History),
CROMWELL Roap, Lonpon, S.W.,

May 15th, 1903.

N.B.—Parcels intended for the Museum should be addressed
to Tur Director, as above, and should be accompanied by a

separate letter of advice.

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TABLE OF CONTENTS.

epee
INTRODUCTION ° . 5 : : . .
CHAPTER '

I. Tur Bronomics oF TsEtsE-Fius (Genus Glossina) . . °

II. Historican Survey—

I, Bionomic
II, Systematic

III. Systematic Position oF THE GENUS

IV. Systematic DESCRIPTION OF THE GENUS Glossina AND ITS
SPECIES, WITH NOTES ON THEIR DISTRIBUTION AND HABitTs

V. Tur Movuts-Parts or Glossina AND Stompns. By Dr. H. J.
-Hansen (Copenhagen) . - : :

VI. BrIsLioGRAPHY—

Part I.—The Literature of Tsetse-flies .
Part II.—Tsetse-fly Disease and the Trypanosoma of
Nagana and Kindred Species

VII. APPENDICES—

Apprnpix A.—Abstract of Lieut.-Colonel Bruce’s ‘*‘ Further
Report on the Tsetse-fly Disease ”’

ApprnpIx B.—‘‘A Note on the ‘Tse-Tse’ Flies of the
British Central Africa Protectorate.” By copay
Richard Crawshay . F .

APPENDIX C.—Copies of Papeete on the Subject of she
Supposed Connection between the Tsetse and the
Buffalo (Bubalus caffer, Sparrman); transmitted by
the Foreign Office to the British Museum en
History) :

ApprrenDIxX D.—Native ‘Mothads of Panteating Aulraals
from the Attacks of Tsetse-flies. Native Prophylactics
against and Remedies for Tsetse-fly Disease. Native
Names for Tsetse-flies . : : .

APPENDIX E.—The Geographical Distribution of Ne agana,
and the Possible Transmission of the Parasite of the
Disease by Species of Glossina other than Gl.
morsitans, and by Blood- eee Flies other than
Tsetse

APPENDIX PA Note on — Origin of eis sei
‘Tse-Tse,’”? By Captain Richard Crawshay

ApPpENDIx G.—The Species of Tsetse found at Entebbe,
Lake Victoria, Uganda

APPENDIX H.—Trypanosomiasis on the Upper Niges. a
Note by Dr, Cuthbert Christy P

PAGE

105

121
253

268

287

290

297

LIST OF PLATES.

PLATE
I. Glossina palpalis, Robineau-Desvoidy. (¢.)

II. Glossina pallicera, Bigot. (6 .)
III. Glossina morsitans, Westwood. (?.)
IV. Glossina pallidipes, Austen. (9 .)
V. Glossina longipalpis, Wiedemann. ( ¢.)
VI. Glossina fusca, Walker. (9 .)
VII. Glossina longipennis, Corti. (?.)
VIII. The Mouth-Parts of Glossina and Stomozxys.

IX. The Mouth-Parts of Glossina and Stomoxys.

Map of Africa south of the Sahara, illustrating present
knowledge of the distribution of Glossina . : - facing page 28

Norr.—In the coloured plates, the vertical line indicating size includes
the proboscis.

FIG.

ae

LIST OF FIGURES IN THE TEXT.

Trypanosoma brucei, Plimmer and Bradford, the parasite of
Nagana, or Tsetse-fly disease, in blood of donkey (x about
2,500 diameters); from a stained preparation kindly lent by
the London School of Tropical Medicine . ‘ : =

. A Tsetse-fly (Glossina longipennis, Corti, from Somaliland), in

resting attitude, showing the position of the wings (x 4)

. Stomoxys sp., from Natal, in resting attitude, oF iphuiiag the position

of the wings(x 4) .

. Hematopota sp., from Zululand, in resting attitude, mais the

position of the wings (x 4)

5. A Tsetse-fly (Glossina morsitans, Westw., ?), before feeding (x 6)

. A Tsetse-fly (Glossina morsitans, Westw., 92), after feeding,

showing abdomen distended with blood (x 6), From a
drawing from life kindly lent by Lieut.-Colonel Bruce .

. Pupa of Zululand Tsetse-fly, dorsal aspect (x 12); a, posterior

extremity, showing pit and right larval stigma, st (x 24);
b, anterior extremity, showing bifurcated longitudinal seam,
which opens to permit the escape of the imago (x 8) .

. Diagram showing nomenclature of external characters of Glossina,

used in description . * : ; .

. Left antenna of Glossina pallidipes, Austen, g, from the inner

side (x 60); p, aperture of sense-organ on third joint . .

. Diagram of thoracic chetotaxy of Glossina, dorsal aspect (x 10).
. Diagram of thoracic chetotaxy of Glossina, pleural aspect (x 10)

. Distal portion of abdomen of Glossina morsitans, Westw. (ventral

aspect), showing (k) hypopygium (x 15)

. Distal portion of abdomen of Glossina longipalpis, Wied. (ventral

aspect), showing (i) hypopygium (x 15)

. Dog suffering from Tsetse-fly disease, or Nagana (after Biles .
. Horse suffering from Tsetse-fly disease, or Nagana (after Bruce) .

. Donkey suffering from Tsetse-fly disease, or Nagana (after Bruce)

PAGE

iv

281
282
284

Fig, 1.
Trypanosoma brucei, Plimmer and Bradford, the parasite of N agana, or Tsetse-fly disease,
in blood of donkey (x about 2,500); from a stained preparation kindly lent by the
London School of Tropical Medicine.

INTRODUCTION.

——_ +o ——

More than fifty years have elapsed since Gordon Cumming drew
the attention of all who were interested in African travel and
adventure to the fatal effects upon horses and cattle of the bite
of ‘‘the famous fly called ‘Tsetse.’” With the publication of
successive volumes containing the experiences of explorers, and
of elephant-hunters and sportsmen who gradually began to pene-
trate into what were then the happy hunting-grounds between
the Vaal River and the Zambesi, the tale of loss and disaster due
to the Tsetse was rapidly swelled ; and it was not long before
these insignificant-looking insects were recognised as constituting
a barrier more formidable to the explorer and the colonist than
almost any other. Thus the general interest taken in the genus
which forms the subject of this book has never slackened, and
‘the Tsetse-fly ”—for it is a common error to speak as though
there were but a single species—is known by name to thousands
of people who have no idea as to what a specimen is like.

For a long time the Tsetse was believed to be directly
responsible for the havoc caused by its bite, the prevailing
opinion being that it elaborated within itself a subtle poison
which when injected into domestic animals occasioned their more
or less speedy death. It was not until the year 1895 that the
brilliant researches of Lieut.-Colonel Bruce, in Zululand, showed
that this idea was mistaken,-and that the part played by the
Tsetse-fly in producing the disease which bears its name was in
the main analogous to the réle of certain mosquitoes in the dis-
semination of malarial fever among human beings. Bruce proved
that the Tsetse is merely the carrier of a hematozoon or blood-
parasite, now known as Trypanosoma brucei, Plimmer and
Bradford,* which appears to live normally in the blood of many

* «Proceedings of the Royal Society of London.” Vol. LXIV.
(August 31, 1899), p. 280.

V1 INTRODUCTION.

species of wild animals in Africa without doing them any harm,
but when taken therefrom and introduced by the proboscis of
the Tsetse into the blood of domestic animals multiplies with
enormous rapidity, and eventually causes death. Quite recently
a similar parasite has been discovered in the blood of man in
West Africa, where several species of Tsetse-flies are known to
occur. In other parts of the Continent all experience goes to
show that the bite of the Tsetse has no ill effect upon man ;
and though it is not yet known whether a Tsetse is the means
of conveying this new parasite which has been found in human
blood, the discovery has at least caused the various West African
species to be suspected, and has lent these insects a new
importance in the eyes of students of tropical medicine.

The present work has been prepared with a view not only to
supplying a résumé of our knowledge of the Tsetse-flies, but also
in order to enable those who may be engaged in Africa itself
upon the investigation of the maladies produced in different
animals by the various species of Trypanosoma, to determine the
species of Glossina responsible for the dissemination of the
hematozoon. The systematic portion of the volume has been
a matter of considerable difficulty, owing partly to the remark-
able dearth in the genus Glossina of structural characters such
as might be utilised for the distinction of species, and partly to
the faulty condition of the bulk of the material available for
examination, Tsetse-flies are generally rare in collections, and
so far as regards the number of specimens of these insects in its
possession the British Museum would probably compare favour-
ably with any other similar institution. But unfortunately
most of the material has been collected by sportsmen unprovided
with the necessary materials for its preservation, with the result
that owing to the rough and ready methods perforce adopted,
there are few specimens in our collection that are not more or
less damaged. It would not be too much to say that the
specimens representing Glossina in the British Museum collection
are in poorer condition than those of any other well-known genus
of Diptera. In consequence of this, and also of the limited
number of specimens from the various localities, no attempt has
been made in the systematic portion of this volume to do more
than describe the external anatomy of the genus and its
different species. The special attention of the reader is, how-
ever, directed to Chapter V., in which Dr. H. J. Hansen gives
the results of his investigation of the mouth-parts.

INTRODUCTION, vii

It is to be hoped that this Monograph will at least have
the result of inducing those who have the opportunity to
collect Tsetse-flies for the British Museum, pinning some
specimens on the spot and preserving others carefully in spirit ; *
this would enable very necessary dissections to be made of the
proboscis and other structures, including the male genitalia.
It would be particularly interesting to see whether the specific
distinctions described in Chapter IV. are supported by hidden
differences in the latter organs.

One new species is described in Chapter IV., and, with the
exception of the type of Gl. palpalis, Rob.-Desv., which, as ex-
plained in the proper place, is probably no longer in existence,
I have been so fortunate as to be able to examine the types of
all the other species, so that the correctness of my identifications
may consequently be relied upon. To Prof. Fr. Brauer and
Herr Josef Bischof, of the K. K. Naturhistorisches Hofmuseum,
Vienna; Dr. R. Gestro, of the Museo Civico di Storia Naturale,
Genoa; Prof. E. B. Poulton, F.R.S., of the Hope Museum,
Oxford; and Mr. G. H. Verrall, of Newmarket, all of whom
have with the greatest readiness allowed me to examine the
various types in their charge or possession, it is my pleasant duty
to express my grateful acknowledgments. My sincere thanks
are likewise due to Mr. L. R. Crawshay, who courteously allowed
me to examine a series of specimens of different species collected
by his brother, Captain Richard Crawshay, to whom I am
indebted for the interesting observations on Tsetse-flies in
British Central Africa, printed in Chapter VII., Appendix B. ;
to Prof. Poulton, who lent me certain specimens of Glossina
morsitans in the collection of the Oxford Museum, in addition to
the types of the three species described by Westwood ; and to
Dr. K. Grinberg, of the Kénigliches Zoologisches Museum,
Berlin, who most willingly sent for my examination the whole
of the Tsetse-material recently collected by Dr. Schilling in
Togo-land, West Africa. As a result of my examination of the
specimens from Togo, I have been able to show that the true
Glossina morsitans, Westw., occurs in that country, which was
hitherto unknown. Whether or not Trypanosoma brucei, the
parasite of Tsetse-fly disease, is capable of being conveyed from
animal to animal by species of Glossina other than this has yet

* For directions as to the way in which Tsetse-flies should be
collected, see the author’s pamphlet,—‘ How to Collect Diptera (T'wo-
Winged Flies),”—issued by the British Museum (Natural History).

Viii INTRODUCTION.

to be proved; but the importance of the occurrence of Gi.
morsitans (in addition to other specits of Tsetse) in Togo is
obvious in connection with the fact that Tsetse-fly disease (called
Surra by German writers) has also been shown to exist there.

Before closing my list of acknowledgments, a word of special
thanks is due to my artist, Mr. A. J. Engel Terzi, for the
unremitting care and attention bestowed by him upon the
coloured plates. Mr. Terzi’s beautiful drawings have been well
reproduced by Messrs. Carl Hentschel & Co., with the result
that the reader now has the advantage of possessing accurate
representations of Tsetse-flies, such as have not hitherto been
available. I have also to thank Mr. Terzi for the trouble that
he has taken over the drawings for the figures that illustrate
the text. To Dr. L. W. Sambon I am indebted for references
to certain papers dealing with Tsetse-fly disease, as well as for
kind assistance in other ways; while 1 have to thank my uncle,
Dr. T. P. Smith, for the preparation of the Index to this
volume. Last, but by no means least, I have to express my
gratitude to Dr. H. J. Hansen, of Copenhagen, who furnished
within a very brief period the valuable account of the mouth-
parts in Glossina and Stomoxys contained in Chapter V., in
addition to preparing the admirable drawings reproduced on
Plates VIII. and IX. I can but consider myself fortunate in
having secured the co-operation of so painstaking and accom- |
plished an investigator, who had previously paid special attention
to the mouth-parts of Diptera, and is already well-known owing
to his work on the subject.

In the first portion of the ‘ Bibliography” (Chapter VI.) it
was deemed advisable in most cases to quote the actual passages
in extenso in addition to giving references, since the majority
of the volumes referred to are out of print or otherwise inacces-
sible to ordinary students, not to speak of those who may use
this book in outlying portions of the Empire. The “ Biblio-
graphy” cannot, unfortunately, lay claim to anything like
completeness. It was thought necessary that this volume should
be published with as little delay as possible, and to have con-
sulted all the books on Africa south of the Sahara, not to speak
of the Proceedings of Foreign Geographical Societies, in which
no doubt many valuable notes le buried, would have entailed
many months’ more work. Such as it is, however, it is to be
hoped that a perusal of the ‘‘ Bibliography ” will give the reader
a fairly good idea of the bionomics and distribution of Tsetse-

INTRODUCTION. ie

flies, as well as of the practical effects of their attacks. If, as
seems likely, the Tsetse in Southern Africa, at any rate, is
destined to become extinct with the big game and the northward
spread of civilisation, the records of earlier struggles with the
pest of the pioneer and elephant-hunter will at least possess a
historic interest, when the railway-train and the traction-engine
have largely superseded the ox-waggon, and the building of the
promised hotel at the Victoria Falls has become an accomplished
fact.

ERNEST E. AUSTEN.

British MusEeum (Natura History),
Lonpon, 8.W.

March 16, 1903.

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A.

MONOGRAPH OF THE TSETSE-FLIES.

(Genus GLOSSINA, Wied.)

CHAPTER “I.

THE BIONOMICS' OF TSETSE-FLIES (GENUS
GLOSSINA).

Although a fairly good idea of the Bionomics of Tsetse-flies
may be gained from a perusal of the Bibliography and Appen-
dices to the present work (Chapters VI. and VII.), it will
probably be more useful and convenient to the reader to give a
short résumé of our knowledge of this subject in a single chapter.
At the outset, however, a word of explanation is necessary as
to the use of the term “Tsetse.”* Since the insects which
form the subject of this Monograph were first encountered by
Englishmen in the vicinity of the Limpopo, the word Tsetse was

* The form here given, which is employed throughout this book, is the
most usual one; but the following variants in spelling are also used by
different authors: Tséts3, Tse-tse, 'I'setze, '[se-tsi, Tzetse, Tzetze, Tzee-tzee,
Tetse, Sétsé (Major Vardon).. There can, of course, be no doubt that the
word is onomatopoetic, and, if used originally for Glossina, that it owes
its origin, to the peculiar buzzing sound made by the fly on the wing or
when commencing to suck blood. But, in spite of many endeavours, I
have so far failed to discover with certainty whether it belongs to the
language of one of the South African Bantu tribes, or was invented by the
Boer ‘‘ Voortrekkers”’ on coming in contact with the fly for the first time
after crossing the Vaal River between 1835 and 1837. Since the fly was
first met with (by English hunters, at any rate—see Chapter II.) in the -
vicinity of the Limpopo, it might be supposed that the word Tsetse is
Matabeli or Zulu, but I have been unable to find out whether this is so.
Gordon Cumming, the earliest English writer to use the word, although
not the first British sportsman to meet with the fly, merely speaks of “‘ the
fly called ‘Tsetse’”’; further on in the same volume he describes an encoun-
ter with ‘‘the famous fly called ‘ Tsetse.’”? Vardon and Oswell, who procured
the actual specimen described by Westwood under the well-known name
Glossina morsitams, are equally reticent as to the origin of its ordinary
designation, and all subsequent writers have been content to use the word

B

2 USE AND ORIGIN OF TERM “ TSETSE.”

originally used by them to mean Glossina morsitans, and that
species alone. Even at the present day it is still so used by the
majority of people interested in African problems, who are un-
aware of the existence of more than one species of Glossina. For
a long time, indeed, while “‘ the» Dark Continent” was no mere
figure of speech, and a trip to the Victoria Falls was looked upon
as a journey into “the Far Interior,” Glossina morsitans remained
the only species known by practical experience, or to any but
entomologists. With the opening up of Africa, however, people
have encountered other species of Glossina, and, recognising
their similarity to or perhaps not distinguishing them from
Gl. morsitans, have spoken of them as “the Tsetse,” or “the
Tsetse-fly.” Thus, although it is a fact that certain writers have
used the phrase ‘the true Tsetse-fly,” apparently with the
intention of distinguishing Gl. morsitans from its congeners, the
word. Tsetse has, in recent years undoubtedly come to be
employed in a generic rather than a specific sense, and it is in a
generic sense that it will be used in the present work. It is
true that we have yet to discover whether all the species of
Glossina are capable of conveying the hematozoon of Tsetse-fly
disease, or whether this baleful distinction belongs to GJ. mor-
sitans alone.* But the species of the genus differ so markedly
from other blood-sucking flies in various details of external
structure, as well as in their appearance (due to the mode #

in one or other of its forms without offering any explanation. Standard
dictionaries of the English language complacently label the word ‘‘ native
name,’’ and proceed to. give a short description of the fly. I am indebted
to my friend and colleague Mr. R. Campbell Thompson, of the Department
‘of Egyptian and Assyrian Antiquities, British Museum, for the nearest
approach to a solution of the question yet achieved. After searching
through a number of African vocabularies, Mr. Thompson writes as
follows:-—“ W. H. J. Bleek’s ‘ Languages of Mozambique ’ (1856) gives’a
comparative table of African dialects, and under ‘fiy’ are the following
‘(the name of the dialect in each case is given in brackets) :—(Sofala)
Tundsi; (Tette) Sénse ; (Sena and Quilimane) Tsénse. I think the last
probably solves the mystery, as the letter » is so constantly assimilated.”
lt is therefore possible that a native word, which perhaps originally
signified any fly, eventually, in the valleys of the Zambesi and the
Limpopo, came to be specially applied, by the natives themselves, to: the
genus Glossina ; just as among Englishmen in South Africa at the present
day the Tsetse is commonly spoken of as ‘‘the fly.’ On the other
-hand, the earliest English and Dutch hunters to penetrate into the
Waterberg and Zoutpansberg, on being told. by their Kafir boys that the
unfamiliar insect which was persecuting their trek-oxen and horses was
as Tsetse, ” may have merely received from them the native nk Fey sae for
Sea flysy 0.7
a Seb <e VII. % oe E., p. 300,

DISTINCTIVE CHARACTERS OF TSETSE-FLIES. 3

carrying the wings) when at rest, that it is impossible to speak
of-a particular species as ‘‘the Tsetse” or “‘the true Tsetse-fly ”:
in*contra-distinction to the others. It follows, therefore, that
the designation Tsetse, in default of a vernacular term for:
each species of Glossina, must be taken as the equivalent of:
the genus rather than of any one species, even the one whichis
best known. :

ae A technical description of the genus Glossina
Géneral Characters — : i is
of Tsetse: how nd of the species (seven in number) at present
to distinguish them known will be found in Chapter IV., so that
from other flies. ., . 3
it is here only necessary to characterise Tsetse-
flies in-simple language, in such a way as to enable a non-
entomological reader to recognise a specimen at sight. Tsetse,
then, may be described as ordinary-looking sombre brownish or
greyish-brown flies, varying in length from 3 to 42 lines (74 to
10 millimetres) in the case of Glossina morsitans, to about 53 lines
(11 millimetres) in that of GI. fusca or longipennis,* with a
prominent proboscis in all species. The hinder half of the body,
or abdomen, in the best-known species, though not in all, is of a
paler colour and marked with sharply defined dark brown hands,
which are interrupted on the middle line ; the abdomen, however,
is invisible when the insect is at rest, as it is then concealed
by the wings. The sexes of Tsetse-flies can readily be distin-
guished when specimens can be examined, since in the male the
external genitalia form a conspicuous knob-like protuberance
(hypopygium) beneath the end of the abdomen (see Figs. 12 and
13, p. 94), which is absent in the female.

It is probable that only those who have suffered from the
attacks of Tsetse can recognise them when on the wing, but in
the resting position their identification is easy. In this attitude
they can be distinguished from all other blood-sucking Diptera,t
especially. from those belonging to the genera Stomowys (Fig. 3)
and Hematopota (Fig. 4), which are most likely to be mistaken
for them, by the fact :that the brownish wings lie closed flat over
one another down the back, like the blades of a pair of scissors,

" * These meastirements are only from the head ‘to the end of the body,
and are exclusive of the proboscis, which projects horizontally in front of
the head, and has a length of about a line to a line and a half, according
to the species. In the crossed lines on the coloured plates, however, the
vertical line indicates the average length of the whole insect, including tiie
proboscis, the transverse line showing the wing-expanse.

t Except midges, which, however, need not be considered since they
cannot be contused with Tsetse-flies.

B 2

4 DISTINCTIVE CHARACTERS OF TSETSE-FLIES.

while the proboscis (i.e. the proboscis ensheathed in the palpi)
projects horizontally in front of the head (see Fig. 2). As
pointed out by Col. Bruce, the closed wings thus give the fly “an
elongated appearance” (Cf. Chapter ViI., Appendix A, p. 271;
see also 55, 76, 119, 152, 161).* Measured from the tip of the.
proboscis to the end of the closed wings the length of Glossina
morsitans is about half an inch; that of Gl. fusea about three-
quarters of an inch.

Most writers who have attempted a description of the Tsetse

Fig. 2.

A Tsetse-fly (Glossina longipennts, Corti, from Somaliland) in resting attitude,
showing the position of the wings. (xX 4.)
have stated that it is rather larger than the common house-fiy,
but the comparison is somewhat unfortunate, since few people,
other than trained Dipterists, are capable of recognising .the
true house-fly (Musca domestica, Linn.), Moreover, at least two
other species of Muscidae, Homalomyia canicularis, Linn., and
Cyrtoneura stabulans, Fln., are quite common in houses in Europe
at certain seasons; the latter of these is considerably larger than

* The numbers refer to the Bibliography (Chapter VI).

GLOSSINA AND STOMOXYS. 5

the true house-fly (for which both are no doubt often mistaken),
while the former is only about half its size.*

Apart from the prominent proboscis and the mode of carrying
the wings when at rest, there is nothing in any way remarkabie
or striking about the appearance of a Tsetse, and the descrip-
tions of most travellers emphasise this fact. As already
mentioned, species of Stomoxys and Hematopota are most likely
to be mistaken for Glossina, and apart from these confusion
can hardly take place. ‘The females of both genera are greedy —
blood-suckers, and often torment domestic animals very greatly.
Of Stomoxys, which, as is shown in Chapter ITI., is a
near ally of Glossina, several species (all closely resembling,
though apparently distinct from, the European Stomoays cal-
cttrans, Linn.) are found in Africa, and in the abdomens of
specimens of one of them in the British
East Africa Protectorate, the late
Captain A. J. Haslam found the
Trypanosoma of Nagana, or Tsetse-fly
disease. Although Stomoxys also has
a prominent proboscis, it is not en-
sheathed in the palpi, and is conse-
quently much more slender than the
proboscis of Glossina, The species of
the former genus are little greyish flies
with black markings; they are much
smaller than Tsetse-flies, and since
their wings when in the resting posi-
tion, instead of closing one over the | Som’
other, diverge at an angle (see Fig. 3), ete shoeing the pauline
like those of Musca domestica, it is easy of the wings. (x 4.)
to distinguish them. Hematopoia, on the other hand, which is
a genus of small horse-flies (Family Tabanide) often known as

Fig. 3.

* The average Musca domestica in Kurope measures about 3 lines (6 to
7 millimetres, or slightly more than a quarter of an inch) in length, and
has a wing-expanse of between 7 and 8 lines (from 15 to 16 millimetres) ;
therefore it is in reality decidedly smaller even than a small specimen
of Gl. morsitans. In the Tropics and hot countries generally Musca
domestica is usually somewhat smaller than in Europe. The common
house-fly justifies its name, for its distribution is now practically world-
wide, as it has been carried by ships to all quarters of the globe. As
regards South Africa, swarms of house-flies are mentioned by Lichtenstéin
(Hinrich Lichtenstein: ‘‘ Reisen im siidlichen Africa in den Jahren 1803,
1804, 1805, and 1806” (C. Salfeld: Berlin, 1811), pp. 192-193) as infesting
houses near the Karroo, in Cape Colony, in the early years of the 19th
century; while Musca domestica was at certain seasons one of the pests
of our troops in standing camps during the late campaign. :

6 GLOSSINA AND HAMATOPOTA.

“clegs” in various parts of Great Britain, resembles Glossina
somewhat closely when at rest. The species of this genus, of
which no less than twenty-two have been recorded from various
parts of Africa, are of much the same size as the larger Tsetse-
flies,* and are of the same brownish colour and elongate shape.
In no case, however, is the abdomen marked with dark bands on
a light ground, while the wings in the resting position do not
close one over the other, but diverge slightly at the tips and are
also somewhat tectiform, i.e. they meet together at the base like
the roof of a house (Fig. 4). The antennz, too, afford a further

Fig. 4.

Hematopota sp., from Zululand, in resting attitude, showing the position
of the wings. (x 4.) The wing markings are omitted. -

means of distinction. While the antenne of Tsetse-flies, as of all
‘Muscide, are drooping, those of Heematopota project horizontally
in front of the head, and being of some length are readily seen.
Where Tsetse-flies Vem in those parts of Africa in which the
are usually found: Tsetse occurs it is not found everywhere, but
their fondness for is confined to definite tracts which are known
river banks and : :
the neighbourhood as “ Fly-belts.” We are still somewhat in
of water, and the dark as to the factors that determine
avoidance of open P,. ¥
plains: swarms met the limits of these “belts,” but, although
Wath in, the Tsetse is undoubtedly dependent upon
certain places. P * i ,
the blood of wild animals for its continued

* The average length of an African Hematopota is 5 lines (11 milli-
metres); average wing-expanse 11 lines (24 millimetres).

USUAL HAUNTS OF TSETSE. 7

existence, all recent evidence goes to show that the most
important element is the physical character of the locality.
Thus, Mr. Alfred Sharpe writing from British Central Africa to
Lord Lansdowne in 1901, with reference to the special con-
nection which many people have supposed to exist between the
Tsetse and the buffalo, states that ‘‘ what regulates the presence
of Tsetse-fly is the description of the country almost as much as
the abundance or scarcity of game.” (See Chapter VII., Ap-
pendix C, p. 295.) As a general rule it may be said that the
Tsetse is confined to damp, hot, low-lying localities, either on the
borders of rivers or lakes, or at any rate not far from water.
Cover in the shape of more or less thick bush or forest is
essential, and the fly is not found on open grass plains. In the
days when South Africa was still a sportsman’s paradise, and the
veld in what is now the Orange River Colony was teeming with
game of every kind, the early hunters who rode and shot over
those plains never met with the Tsetse until, pushing farther
afield, they entered the hot, moist kloofs in the valley of the
Limpopo.

James Chapman [30], one of the earliest authorities, writing
in 1868, states that ‘“‘the Tsetse-fly is generally found within a
few miles of water, in rich sandy ridges near marshy spots, and
generally in mopani or mimosa forests.” In the Northern
Transvaal the Tsetse in some places is found on low hills, and
Vardon and Oswell having met with the original specimens of
Glossina morsitans on the Siloquana Hills (between the Maga-
laqueen or Nylstroom River and the Limpopo) in 1845, Vardon
was led to assert that it is “usually found on hills.” Later
knowledge, however, does not warrant such a generalisation,
although it is true that patches of Tsetse-fly are not always
situated in low-lying districts. Thus, while in British Central
Africa, according to Sir Harry Johnston [145], the Tsetse-fly is
‘roughly speaking . . . . absent from any district that is above
3,000 feet in altitude,” Captain A. St. H. Gibbons [152] in
Northern Rhodesia, in February, 1896, during the wet season,
found it very troublesome on high ground to the south of the
head-waters of the Nanyate River, at an altitude of 4,110 feet
above the sea-level. “This, however, must be somewhat excep-
tional, and it is in hot, moist river-valleys that fly-belts
generally occur. In the Jubaland Province of the British East
Africa Protectorate, according to Mr. J. W. P. McClellan, ‘“ The
worst places are damp, dark, and low-lying, shaded chiefly by

8 TSETSE AND RIVER-BANKS: LOCAL SWARMS.

the Ndoma Palm.” * Similarly in Sierra Leone Glossina palpalis
occurs in mangrove thickets where streams enter the sea.

It is curious that Sir Harry Johnston [125] should have
committed himself to the statement that the Tsetse-fly “appears
never to go near the edge of a river,” and has “ a great dislike
to water” [145], since the universal experience of travellers and
sportsmen is that precisely the reverse is the case. Thus Mr. F.
C. Selous [76], writing in 1881 of the plague of Tsetse-flies along
the southern bank of the Zambesi and Chobe, to the westward
of the Victoria Falls, states that they are “usually found in
great numbers near the river,” and that “along the water’s edge
they are an incredible pest. . . .”. Instances to the same effect
might be multiplied, but as they are all recorded in the Biblio-
graphy it will suffice to refer the reader to the testimony of,
among others, Thomas Baines [26] (Tsetse at the Victoria Falls);
Sir John Kirk [28], who states that the Tsetse extends along
the banks of the Rovuma River for 115 miles; E. C. Hore ['78]
(Tsetse-fly on the shores of Lake Tanganyika) ; and Dr. Holub
[110] (abundance of Tsetse at the water’s edge on the Mo-Njeko
River, August 7th, 1886).

Within the limits of a fiy-belt, however, Tsetse are not
always numerous, and it may happen that only a few specimens
are encountered in the course of a day. On the other hand they
are often met with in much greater numbers. In the passage
already referred to, Selous [76] speaks of the fly “attacking one
in a perfect swarm, from daylight to sunset .. . .”; David and
Charles Livingstone [27], writing in 1865 of their experience of
the Tsetse-fly in a valley to the west of Zumbo, state that the
flies “accompany us on the march, often buzzing round our
heads like a swarm of bees” ; Captain F. F. Carter [66] in 1880
found “ Tsetse-fly in thousands” on the site of a large deserted
village, near Karema, German East Africa; Selous [86],
writing of Mashunaland in 1882, speaks of the Tsetse as
“swarming” along the Panyame and Umsengaisi Rivers, and
as occurring in another locality “in millions.” The late W. M.
Kerr [98], describing his experience of Tsetse on the Mukumbra
River in 1884, states that ‘the stifling sultry air was literally
alive with the Tsetse-fly, against whose maddening attacks
clothes were no protection, our only safeguard being to beat
them off with twigs and small branches of shrubs... .”
M. Edouard Foi gives a similarly graphic account of an

* Chapter VII., Appendix C, p. 294.

“FLY-BELTS” AND THEIR EXTENT. 9

encounter with swarms of Tsetse on the Kapotche River, their
furious onslaughts rendering it impossible to take a meal, or
even to remain seated. Sir Harry Johnston [145] states that
in British Central Africa, “on the Mwanza River, an affluent of
the Shire nearly opposite to the Katunga, the Tsetse are so
numerous that the only domestic animals which can be kept by
the natives are fowls.” Lastly, to pass for a moment to Somali-
land, Mr. C. V. A. Peel [163] who met with Tsetse (Glossina
longipennis, Corti) in that country in 1895, mentions an occasion
when his camels on returning to camp in the evening “ were
followed by a perfect swarm of Tsetse-fly.”
4 . As stated at the commencement of the pre-
Aha lh ceding section a tract of country in which the
distribution and Tsetse occurs is termed a ‘‘Fly-belt.”* Al-
limits of Tsetse = though along the courses of rivers, or in the
within these areas. © =) ?
low country bordering a coast line, fly-belts
may extend for hundreds of miles, varying greatly in width
according to the nature of the country, it does not follow that
Tsetse-flies are to be met with at every point throughout the
extent of the belt. More usually they are confined to particular
patches of forest or bush, the area of which may be quite small.t
Thus, according to Baines [55], ‘‘the fly is extremely local, and
extensive districts in which it prevails may be passed through
by the aid of guides, who know the ‘patches’ of fly, just as a
pilot knows the shoals of an estuary.” Mr. J. W. P. McLellan,
writing of the Jubaland Province of the East Africa Protect-
orate, states that “in many places the areas in which fly
exist are quite small, possibly only a few hundred yards in
extent... .”’{ Again, Mr. F. J. Jackson in 1887 found Tsetse
‘*‘in great numbers in a small patch of thick bush, about a mile
and a half long and three-quarters of a mile wide,Ӥ near
Taveita ; later he met with it ‘in considerable numbers in a
narrow belt of forest, not more than a mile wide, between
Mkonumbi and Witu.Ӥ The limits of these patches or areas,
as of the fly-belts themselves, are doubtless defined more or less
by the physical characters already alluded to ; but, nevertheless,
it is often difficult to account for them. Various writers have

* Often spoken of shortly as ‘‘ the Fly,” just as the term ‘‘ Fly” is also
used in the special sense of ‘‘ Tsetse.”

+ These small fly-infested areas are themselves termed belts by some
writers.

t Chapter VII., Appendix C, p. 294,

§ Ibid, p. 296,

10 RESTRICTED OCCURRENCE OF TSETSE.

drawn attention to the fact that Tsetse may abound on ‘one
bank of a stream while there are none on the other. According
to Andersson [20], “cattle may be seen grazing securely on one
side of a river, whilst the opposite bank swarms with the insect.”
On one occasion, Livingstone [21], during a two months’ sojourn
on the Chobe River, preserved his cattle by keeping them on the
northern bank, where not a single Tsetse was found, although
the south bank, “only fifty yards distant,” was “infested” by
the fly. A graphic description is given by Mr. C. V. A, Peel
[163] of his experience in Somaliland in 1895. Mr. Peel writes:
“Tt is an extraordinary sensation coming into a belt of ‘fly.’
There may be but a tiny river-bed. On one side of it not a fly
will be encountered, but walk a dozen feet and they suddenly
come buzzing by one in hundreds.” The results of an incursion
into the “Fly Country” in Zululand are thus described by
Lieut.-Colonel Bruce *: “On entering ‘ Fly Country’ one is not
left long in ignorance of the presence of the Tsetse. The natives
may be seen slapping their naked legs, the dogs bite round, and
the horses kick. The Tsetse, however, may be said to be some-
what local in its distribution in the ‘Fly Country,’ being only
met with now and then and in few numbers, until you enter
some glade or clear space in the thorns, when suddenly the
slapping, biting, and kicking go on with tenfold energy, and you
ean catch thirty or forty flies in a few minutes.”

Just as the worst patches of fly within the limits of a Fly-
belt are often sharply defined, so in a large tract infested by
Tsetse there may be small areas which for some reason, perhaps
owing to their having been cleared of bush, the fly never enters.
These spots form veritable harbours of refuge for the traveller
who may be compelled to cross the Fly-belt with oxen and
horses, since, by travelling at night and taking care to keep the
animals within an asylum of this kind during the day, the
dangerous zone may be traversed in comparative safety. Thus
Selous [121] describes how the forty miles of Tsetse-infested
swamp on the western bank of the Loanja River can easily be
crossed in three nights, owing to the fact that “ there are two
islands in the swamp free from ‘fly,’ to which the oxen can be
driven to feed and rest during the day-time.” .

The reader who may wish to discover what is known of the
actual extent of Fly-belts in a particular region must consult the
Bibliography, but should be careful to note the dates to which

* Cf. Chapter VII., Appendix A, p. 271.

THE EXPLANATION. OF FLY-BELTS. 11

statements refer ; since, owing to the opening-up and colonisa-
tion of many parts of Africa and the enormous diminution in
the numbers of big game, as will be shown in the next section, it
by no means follows that the limits of a Fly-belt are the same
to-day as they were even a few years ago; while in certain cases
the belt itself may have ceased to exist.

The limitation of the Tsetse to “belts” is not so remarkable
as might at first sight appear. There can be little doubt that it
is due to a characteristic of the Order to which these insects
belong, which, though frequently overlooked, is exhibited by the
majority of species of Diptera, and has attracted special attention
in the case of the Tsetse owing to their blood-thirsty nature and
the fatal consequences of their bites when inflicted upon domestic
animals. Although not “social” insects in the ordinary sense of
the term as applied to animal communities, Diptera, as a whole,
show a marked social tendency on the part of individuals of the
same species. That is to say, there is a tendency which causes
individuals of the same species to assemble in particular places in
more or less close proximity to one another, although without
exhibiting anything in the shape of mutual aid. Thus, in spite
of the fact that the distribution of most species of Diptera is
remarkably wide, it will be found in the majority of cases
that individuals of the same species do not occur everywhere
throughout its area, but are confined to particular spots; and
wherever they are met with they are usually to be found in
some numbers. Of course there are many exceptions, but the
generalisation nevertheless holds good. Again, although remark-
able instances of migration have been recorded in the case of
certain species of Diptera, they must be regarded as exceptional,
and it will be found that Diptera, as a rule, do not travel far
from the spot in which their larval existence was passed. The
result is that we find a larger or smaller number of individuals
of the same species collected within, perhaps, quite a small area,
outside which it may be impossible to find a single example.
This is well-known in the case of the midges and other allied
forms, which attract attention owing to the habit of the males
of collecting together and dancing in the air in a swarm at
certain seasons; and similar phenomena are exhibited by the
males of certain species of Muscide (belonging to what is still
generally termed the Family Anthomyide). In England, again,
phenomena precisely analogous to “ Fly-belts” are displayed on
a limited scale by some of the smaller forms of ‘“ daddy-long-legs ”

12 ASSOCIATION OF TSETSE WITH BIG GAME.

belonging to the Family Limnobide, species of which have been
found to be confined to certain damp spots in woods, where
individuals are always to be met with in smaller or larger
numbers at the proper season, although none may be found in
what are apparently equally suitable localities elsewhere. Thus
the occurrence of Tsetse-flies in “belts” is not difficult of
explanation, although the alleged fact that animals may be in
perfect safety on one bank of a stream when bushes on the other
side are full of the fly is exceedingly hard to understand.
Although, as already explained in a previous
Association of Tsetse section, the Tsetse is particularly local in its
with Big Game. f _ H ° Z
distribution, and is only found in warm, moist
tracts in the neighbourhood of water, where cover occurs in the
shape of forest or bush, the only possible conclusion to be drawn
from the practically unanimous and very numerous statements
on the subject of the connection between the fly and game,
which will be found in the Bibliography, is that were it not for
the big game, on the blood of which it feeds, the Tsetse would
soon cease to exist, at least in numbers sufficient to be formid-
able. Mr. Alfred Sharpe, C.B., writing to Lord Lansdowne
from Zomba, British Central Africa Protectorate on September
30th, 1901, says: ‘‘ Tsetse-fly would appear to depend upon wild
game for their existence, as I have never found Tsetse in any
locality where game was totally non-existent” (Chapter VII.
Appendix C, p. 295). So far as I am aware the only evidence to
the contrary comes from East Africa, where, according to the
testimony of various observers, Tsetse would appear to be less
intimately connected with game than in the Central and
Southern portion of the Continent (127, 151]. Thus Mr. J. W.
P. McClellan, writing from Nairobi, East Africa Protectorate,
on September 4th, 1901, expresses the opinion that ‘“‘ Tsetse-fly
is to be found in certain dark, damp, low-lying localities, irres-
pective of big game of any kind” (Chapter VII., Appendix C,
p- 294). Again, Mr. F. J. Jackson [127] states that “it is
supposed by a good many people that the Tsétsé-fly only exists
where game beasts, especially buffaloes, are most plentiful, and
that the fly disappears as the game is killed off or driven away.
This may be so in South Africa, but it is certainly not the case
in East Africa, as the belts of fly country in East Africa are
almost devoid of game, with the exception of the river Tana.”
Mr. Neumann [151], writing of the Athi River above its
junction with the Tsavo, in May 1895, notes the remarkable

CONDITIONS IN EAST AFRICA. 13

scarcity of game “even along the banks of the river.” He
adds: “Here are great stretches of uninhabited bush country
with a perennial river running through it, and hardly any
animals, though plenty of birds and of ‘ fly’ (Tsetse).” More
recently, in a letter to the Marquis of Lansdowne, dated 27th
September, 1901 (see Chapter VII., Appendix C, p. 297), Mr.
Jackson has expressed the belief that “the Tsetse is, like the
mosquito, only a blood-sucker by predilection.” The meaning of
this phrase would appear to be that, in default of blood, which
it prefers, the Tsetse can continue to subsist on the juices of
plants. Be this as it may, it must, I think, be regarded as an
established fact that when game absolutely ceases to exist in a
given locality, the Tsetse-fly, if present, soon disappears. As an
argument in support of his belief, Mr. Jackson mentions that
near Kibwezi, in East Africa, in April 1892, “at a time when
the whole of the ‘fly-belt’ was parched and dried up—there
being no water between Msogoleni and Tsavo River, a distance
of fifty miles, and consequently no game of any kind—-the Tsetse
was more plentiful than at any other time, before or since,”
when he has traversed that district. He also states that though
a species of Tsetse is “plentiful” in the Botanical Gardens at
Entebbe, on Lake Victoria, there are no mammals there “ with
the exception of a few monkeys and squirrels, and certain small
nocturnal beasts, such as ichneumons, etc., and an occasional
hippopotamus.” Whether the proboscis of the Tsetse can pierce
the integument of plants as easily as that of mammals has yet to
be proved by actual observation ; at any rate no one has so far
ventured to make such an assertion. But in connection with
Mr. Jackson’s statements it must be remarked that even
assuming that the Tsetse in the Botanical Gardens at Entebbe
make up for the deficiency in the supply of mammalian blood
by imbibing the juices of flowers and plants, those in the fly-
belt near Kibwezi could hardly have done so, since we are
expressly told that at the time referred to the whole area was
“ parched and dried up.”

The true explanation of the Kibwezi phenomenon seems to
be that it is possible for one generation of adult Tsetse-flies, as
for other adult insects, to continue to live for some time or for
the whole period of existence without food, if the latter is not
forthcoming. Thus assuming a fly-belt to be deprived of all
mammalian life, the Tsetse-flies which thereafter emerged from
pupe would continue to exist even though fasting ; and they

r4 CONDITIONS IN SOUTH AFRICA.

would naturally avail themselves of the opportunity of obtaining:
a meal of blood afforded by the passage of men and other
animals through their haunts. Whether reproduction is im-
possible without such a meal, as appears to be the case as’
regards certain species of mosquitoes although apparently not:
the rule, remains to be proved. If, however, Tsetse which have
not fed on blood are incapable of reproduction, it follows that
they would become extinct in one generation after this food
supply ceased.

Our knowledge of the distribution of the various species of
Glossina is as yet very scanty, but it is possible that the species
concerned in Mr. Jackson’s statement with regard to the Fly-
belts in East Africa [127] referred to above, is Gl. pallidipes,.
Austen, rather than Gl. morsitans, Westw. If this is so, the:
discrepancy between Mr. Jackson’s testimony and the consensus
of opinion as to the close association between Gl. morsitans and:
big game might be due to a difference in the habits of the two
species. But the more probable explanation would seem to be:
that, as already mentioned in a previous section, the Tsetse will
not live on open grassy plains, even when teeming with game.*
It may be added that in the subsequent letter to Lord Lans-
downe, Mr. Jackson appears at the outset to admit the con-
nection, even in East Africa, between the Tsetse and. game;
although, as already shown, he afterwards advances the view
that ‘the Tsetse is . . . . only a blood-sucker by predilection.”

‘Three species of Tsetse are known to occur in Hast Africa,
and more precise observations are necessary to decide the
relationship between each of them and the game of that region,
With regard to South Africa, however, there can be no question
that the extermination of big game has been followed or aecom-
panied by the disappearance of the Tsetse from many localities
in which it formerly abounded. That this would ultimately be
the case was suggested long ago by Livingstone [21], and the
assertion has been repeated by many subsequent writers.[ Ample
proof of the correctness of the forecast is furnished by what has
taken place in the more southern portions of the Transvaal [ 105,
112]. In South Africa, according to the testimony of various

* In East Africa, according to Mr. Jackson [127], ‘‘ the open, undulating;
grassy plains of the Masai country, and other places of a like nature, are
the headquarters of by far the greatest quantity and variety of game, and
are entirely free from the Tsétsé-fly,....” "

+ Glossina morsitans, Westw., Gl. pallidipes, Austen, and: Gl. fusca,
Walker.
~~ Y Cf. 55, 57, 112, 128, 135, 153.

TSETSE AND BUFFALO. ° 15

writers, the Tsetse follows the movements of the big game, and
should the latter return to a locality from which it has been
driven out the fly may come back with it. This is stated to
have happened in the country between the Gwaai and Deka
rivers in Rhodesia, in consequence of Matabele and other raids
(77, 153]; and similarly in Zululand the Tsetse is said to have
increased in numbers, as an indirect result of the Zulu War of
1879, since the devastation of many inhabited tracts caused by
the campaign led to the multiplication of wild animals in these
localities.

Whether the Tsetse is more partial to and more dependent
upon one species of game than another is doubtful, although the
majority of writers, from Chapman [30] onwards, have stated
that it is especially associated with the buffalo. Most of these
statements naturally refer to South Africa, and a possible
explanation is that in that region, prior to the advent of the
rinderpest, the buffalo was more abundant than other game in
precisely those localities beloved by the Tsetse [¢/: Selous, 76].
In Central and East Africa, at any rate, as shown by the letters
from well-qualified observers printed in Chapter VII., Appendix
C, and elicited by the assertion that to protect the buffalo would
be tantamount to protecting the Tsetse as well, the latter does
not appear to be more dependent upon the buffalo than upon any.
other species of game. The association of the Tsetse with the
buffalo in South Africa has led to the belief that the fly breeds
in the dung of that animal, but thanks to the observations of
Colonel Bruce we now know that this idea is without foundation.
In addition to the buffalo, various writers have stated that they
have found the Tsetse associated with the elephant, rhinoceros,
and the larger antelopes, while Livingstone [49], who met with
the Tsetse on the Rovuma River in 1866, found that the only
game in that locality were hippopotamus and pig.

Although it has been stated by Colonel Bruce
that, so far as he is aware, “ Nagana is not
confined to certain months of the year as
Surra is,” * the Tsetse itself, where it occurs, would appear to be
more abundant at one season of the year than at another, while
temporarily it may even be absent altogether. Thus, according to
Buxton [41]: ‘“ The fly appears only at certain seasons, . . . The
head of a kraal, about thirty miles distant from the point where
we found the Tsetse most abundant, told us at that time the fly

* Cf. Chapter VII., Appendix A, p. 286.

_ Seasonal
‘Prevalence.

16 SEASONAL PREVALENCE.

was not in his district, and pointed to a heifer and some goats,
which he said he intended to send away before the fly season
came on.” Our information on this point, however, is as yet
very scanty, and to a certain extent conflicting. It was found
by Harris [5] that the fly infested the Mural Berge, in the north
of the Transvaal, especially during the rainy season ; and Mr.
and Mrs. Petherick [37] were informed that at the same period
of the year a certain kraal in the country of the Rhol tribe, to
the north-west of Gondokoro, was untenable owing to the pre-
sence of Tsetse. According to Chapman [30], the Tsetse ‘ often
moves about” with the buffalo in the rainy season ; but the same
writer subsequently states that “heavy rains are said to kill
the fly in the season when they prevail.” Kirk [28] tells us that
Tsetse are “most numerous and troublesome in the hot sultry
weather before rains ;’ while Mr. F. J. Jackson [119], speaking
of East Africa, says that, “ where it exists,” the fly “ appears in
much greater numbers on a dull or rainy day.” It should be
noted that in South Africa, at any rate, the rains fall in the
summer, the winter being the dry season. It is therefore curious,
especially in view of the evidence already adduced, to find that

Dr. Bradshaw ['77], writing of the fly-belt on the south bank of

the Zambesi and Chobe, states that the Tsetse “is much more

plentiful during the winter months, namely, from April to end of

September, than during the summer” ; while, on the other hand;

Selous, referring to the Batonga country between the Zongwe

River and the Zambesi in June (mid-winter), says that Tsetse

were “pretty numerous, and must be very much so later on

during the hot months.” Bradshaw, however, is supported by
an experience of Dr. Holub, who on the Mo-Njeko River, on

August 7th, 1886, found Tsetse more abundant than he had

ever seen them before.

The first of Colonel Bruce’s suggestions for future work on
Tsetse-fiy disease* is—‘ That all trustworthy information regard-
ing seasonal prevalence [of Nagana] be collected”: it will have
been seen from the foregoing statements that “trustworthy
information ” as to the seasonal prevalence of Tsetse-flies then
selves is equally necessary.

Period of the day at Lhe Tsetse, like other day-flying insects, does
which Tsetse-flies not begin to be active until the sun grows
are most active. . hot;.. Livingstone [21] states that it is ‘“re-

markably alert, avoiding most dexterously all attempts to capture

* Of. Chapter VII., Appendix A, p. 286,

FEEDING-TIME OF TSETSE. 17

it with the hand at common temperatires ;* in the cool of the
mornings and evénings it is less agile.” Again, according to
Colonel Bruce,t-—“ The Tsetse is not at all easy to catch with the
hand, especially during the day, being nimblé and quick of move-
ment, but at early morning or stinset they become more lethargic,
and are then more readily secured.” The following' remarks by
Sir John Kirk [28] are also worth quoting :—“ In the morning;
while the dew hangs on the grass, and before the heat of the
rising sun has warmed the air, the Tsetse is dull and sluggish,
resting on the under side of some leaf or blade of grass; when
forced to take wing they may then be easily caught. Even at nine
o'clock they aré not very active, and fly about with a peculiar
buzzing sound; with the heat of the day they become a real
annoyance to the traveller, constantly biting him on the hands,
face, or neck, dexterously evading a blow, and again alighting on
the very spot from which they have been driven.”

According to Captain Crawshay,{ the Tsetse bites “at any
time,” but is “ niost aggressive” “ during the hottest hours of the
day” ; Bradshaw [77] says, “It bites throughout the day, except
when it rains” ; and Selous ['76], writing of the plague of Tsetse-
flies on the south bank of the Zambesi and Chobe, to the west of
the Victoria Falls, in 1874, speaks of the insects as’ “ attacking
one in a perfect swarm, from daylight till sunset.” On the other
hand, Colonel Bruce, describing his experience of Tsetse-flies in
Zululand, says: “ About sunset seems a favourite feeding-time,
and ‘then the poor creatures [horses] would be surrounded by 4
perfect cloud of the flies, while some hundreds of them-would bé
settled on them at thé same moment.”§ Again, it may “be noted
that Dr. W.'S. Radford, referring to a species of “ gad-fly” (i.e.,
horse-fly) in Jubaland, East Afriea Protectorate, the bites of
which he asserts produce in camels symptoms “ identical in every
particular with those produced by Tsetse,” states that “This
fly, unlike the Tsetse, attacks animals during the day at all
hours.” || The inference from this, although Dr. Radford does not
mention the period. of the day at which Tsetse-flies in Jubaland
are most aggressive, is opposed to the testimony of Crawshay,
Bradshaw, and Selous, which has already been quoted... But the
explanation may very possibly be that the Jubaland Tsetse-fly

* Baines [55] states that on a cold day the fly is “‘dormant.”—E. E. A.
+ Cf. Chapter VII., Appendix A, p, 272.
¢ Chapter VII., Appendix B, p. 288.

§ Cf. Chapter VII., Appendix A, p. 272.
|| Chapter VIT., Appendix ©, p. 293.

18 BEHAVIOUR AT NIGHT.

belongs to a species * with habits somewhat different from those
ot Glossina morsitans and the other better-known forms.

The Tsetse appears to pass the night resting
either on the ground or on the smaller branches
of bushes and trees,t and although there is a wide-spread belief
that during the hours of darkness fly-belts may be crossed with
impunity, there is abundant evidence to show that this is not
invariably the case. A moonlight night during the South African
winter may be safe enough,{ but at other seasons and in more
tropical latitudes it is well to wait until the temperature of the
air has fallen sufficiently to render the insects sluggish. Brad-
shaw ['77] states that the Tsetse bites “ during part of the night,
if warm,” adding, “ I consider it dangerous to travel at night with
cattle and horses, until it begins to grow cold towards the middle
of the night, as I have been bitten often until past 11 P.M.”
Selous [76] describes a moonlight tramp along the Chobe River,
when the Tsetse ‘kept flying up from the ground on to my
naked legs, and bit as furiously as in the daytime”; and Captain
Crawshay, writing of his experience of Tsetse-flies in Nyasaland,
agrees that they bite at night as well as by day, when the moon
is bright.§ On the other hand, Kirk [28] writes: “ By night
I have never been bitten by Tsetse, nor do they fly about after
sunset” ; and Bruce|| says that he has ‘not noticed them
biting during the night....” These discrepancies, however, are
no doubt due to differences in locality, temperature, and other
conditions. Captain A. St. H. Gibbons [152], writing of his
experience of Tsetse on the Njoko River, Northern Rhodesia,
in August 1895, says: “At night the danger is very small,
though it is a mistake to imagine that the Tsetse keeps such
early hours as other flies. I have at times been worried by
them an hour after the sun has gone down, and have known
flies to buzz into my tent as late as 9 o’clock on a dark night

Behaviour at night.

* Perhaps Glossina longipennis, Corti.

+ Dr. Holub [rroj draws attention to the risk incurred by crossing at
night ‘‘certain thickly-wooded laterite knolls,” in the vicinity of the
Zambesi, which happen to be infested by Tsetse, since owing to the narrow-
ness of the track 1 is impossible to prevent the waggons from brushing
the flies from the twigs and carrying them along.

t Cf. C. J. Andersson, [20]. }

§ Cf. Chapter VII., Appendix B, p. 288, Sir Harry Johnston’s advice
[145] that, ‘‘1f a Tsetse-haunted district must be crossed, it should be
done at night-time—by moonlight if possible,” should therefore be received
with caution. According to Baines [55], a ‘‘dark and cloudy ” night is
safest.

|| Cf. Chapter VII., Appendix A, p, 272.

FLIGHT. AND BUZZING. 19

and make a bold dash for supper at my expense. At that time
of night they are easily caught, and almost invariably found
their way into spirits of wine.”
Authorities agree in describing the flight of
Mode of flight, and : . ”

noise produced the Tsetse as “rapid and straight” [30, 55).
by pi when on Baines [55] says that “its sight and smell
ee wae seem to be keen,” and Foa [135] asserts that it
‘comes to its prey from a long distance, whether it is that its
sight is keen or its sense of smell is exceptionally acute.” * In
the opinion of M. Foa the latter suggestion is more probably
correct, since he has observed that the Tsetse “ always comes up
wind, and generally bites by preference on the leeward side.”
Colonel Bruce writest:— “The Tsetse has a direct flight,
flopping, if I may use the term, suddenly on the animal attacked, ,
and is very pertinacious.” On the completion of its meal, ‘when /
its abdomen is full of blood, its flight becomes heavy and it at |

once hides itself in order to digest its meal in peace” [135].

That the Tsetse owes its name to the sound produced by it
when on the wing has already been pointed out in the note at
the commencement of this chapter,t and its characteristic buzz
has been referred to by various writers. Bradshaw [77] remarks
that, ‘Its buzz is peculiar and not easily forgotten”; while
according to Livingstone [21], “Its peculiar buzz, when once
heard, can never be forgotten.” Captain Gibbons writes [152] :—
“The fly is frequently to be heard giving vent to the high
pitched buzzing note which gives it a name . ”: and Colonel
Bruce says,§ ‘‘The fly makes a loud buzzing sound when flying,
but after its feed and at rest it emits a peculiar sharp, shrill
note, probably caused by an action of the wings.” || According to

* It may be remarked that a large sense-organ, possibly olfactory in
function, is situated in the third joint of the Tsetse’s antenna (See Fig. 9,
p. 63).

t+ Vide Chapter VII., Appendix A, p. 272.

} See page 1.

§ Vide Chapter VII., Appendix A, p. 272.

| The postprandial song of the Tsetse is more probably produced by
the thoracic stigmata or spiracles, or by the vibration of the thoracic
walls. A large number of Diptera, especially Muscide and Syrphide
(Hover-flies, &c.), possess the power of thus producing a shrill sound
while at rest, with the wings closed. Cf. “ Sericomyia borealis singing
while at rest”: Rev. E. N. Bloomfield, Entomologist’s Monthly Magazine,
Vol. XVIII. (1881), pp. 159-160: see also, A. H. Swinton, sbed. (January
1882), pp. 189-190, and J. Hellins, ibid., p. 190. G. C. Bignell (Zntomolo-
gist’s Monthly Magazine, 2nd Series, Vol. II. (1891), p. 225) states that
Microdon mutabilis (Linn.), 2 somewhat rare British Syrphid, produces a
humming noise when at rest and annoyed, by means of the vibration of the

halteres (the little knobbed organs found in all Diptera, except certain
c 2

20 MODE OF FEEDING. °

Captain Crawshay,* the flight of the Tsetse ‘“‘is powerful and
noisy : the buzz is full-toned and somewhat highly-pitched—not
dull and droning like the buzz of the ordinary horse-fly.”

Fig. 5.
A Tsetse-fly (Glossina morsitans, Westw., 2), before feeding. (x 6.)

When alighting for the purpose of making a.

Behaviour when - , :
sucking blood; meal, T'setse-flies, according to Captain Craw-
time taken by the shay,* “do not settle as slowly as the horse-fly

fly in sucking its ‘ ‘

fill: blood-sucking but land with a bump, standing well up on
ity to their legs.” The method of feeding is described
OStRriat by the same authority as follows: * “ When a
‘Tse-Tse’ settles with the intention of feeding—in the early
mornings they usually simply settle on men’s backs to sun
themselves, away from the ground and vegetation wet with the

Fig. 6.

A Tsetse-fly (Glossina morsitans, Westw., 9), aiter feeding, showing abdomen distended
with blood. (x 6.) From a drawing from life, kindly lent by Lt.-Colonel Bruce.

dew-——he inserts his proboscis, lowers his head, and raises his
abdomen until it is almost vertical: when doing this, and for
some little time after he has commenced sucking, he works his:

wingless forms, and projecting from the thorax behind the wings), which:

come into contact with ‘‘some short bristly hairs situated on the abdomen

and pointing towards the thorax.”
~* Chapter VIL, Appendix B, p. 288.

MODE OF FEEDING, 21

wings, buzzing in a minor key, rather like a bee when held
forcibly, though not so powerfully: when the keenness of his
appetite has been somewhat appeased, he stops working his
wings and sucks in silence. If left to himself, he will suck until
his originally skinny barred abdomen becomes a large crimson
bead [compare Figs. 5 and 6]. He is then almost. helpless: if
touched he will not fly, and if brushed off he will only go a yard
or two, to settle heavily on a bush, or gradually sink down to
the ground.” A similar account is given by Foa [135], who
writes :—‘‘ When it. settles, it does so so gently that one does
not feel it; if remains thus motionless for fifteen or twenty
seconds, its proboscis projecting in front, in a mistrustful
attitude, ready to take flight. When it believes itself safe, it
lowers its weapon, separates its feet. so as to flatten itself out
more and pierces the flesh without causing any pain to start
with, as does the mosquito. . . . While its proboscis, which is
at least a quarter of a centimetre in length, disappears com-
pletely in the flesh, it remains motionless sucking blood, its
abdomen swelling and becoming rose-coloured owing to its
transparency, and immediately afterwards appearing deep red
and plump. It is not until the moment when it has already
imbibed a large portion of its meal, that a slight pain or rather
itching betokens its presence. When its stomach is full it is
still very difficult to catch with the hand, for it does not fly off,
but quickly dodges to one side.”

It should be noted that M. Foa’s statement that the proboscis
of the Tsetse “ pierces the flesh without causing any pain to start
with,” is not altogether borne out by the experience of others.
Thus, according to Selous [76], the bite of the Tsetse is “far
more severe” than that of a flea, and “about one in every ten
bites (that perhaps touches a nerve) closely resembles the sting
of a wasp or bee, as it will cause one when seated to spring up as
if pricked with a needle ;” while Colonel Bruce writes * :—“ In
man the burying of the proboscis in the skin is accompanied by
a sharp prick, which draws one’s attention suddenly to the spot ;
but the pain is, as a rule, trivial, and the subsequent redness. and
irritation very slight indeed—in fact, less than that caused by
the mosquito.” Captain Gibbons [152] states that the tread of
the Tsetse “is so light that the sharp prick of the proboscis is
generally the first indication of his whereabouts.” | According
to Mr. F. J. Jackson [119] :—‘Its bite in man, though very

* Cf. Chapter VII., Appendix A, p. 272. t See also [27].

22 SENSATIONS PRODUCED BY TSETSE BITE.

sharp and needle-like, leaves no irritation, and it invariably
chooses those parts that are shaded from the rays of the sun—
such as under the brim of the hat, behind the ear, neck, and
under the throat if not protected by a beard. If allowed to
suck itself full, its body becomes so distended that it has the
appearance of a small unripe purple grape [see Fig. 6], and it is
unable to fly more than a few yards before settling. Its proboscis,
which is very slightly curved downwards, is attached to the head
horizontally, and is protected by a sheath divided longitudinally.
When it bites the proboscis is vertically lowered, whilst the double
sheath is retained in its horizonal position.” The “sheath” here
referred to is formed by the palpi, which in the normal position,
when the proboscis is not in use, are applied to the latter so
closely as to conceal it and give the appearance of a single organ.

According to Chapman [30] the Tsetse “assails different
animals in their most defenceless parts: a man behind the back
between the shoulders, and an ox on the back or under the
belly ; a horse in the same places, and inside the nostrils ; and a
dog on the forehead, &c.” Colonel Bruce writes : *—“ It affects
in horses the legs, especially in the region of the fetlock, and on
being detached from its position by the vigorous kick the horse
gives, will only rise a few inches and again settle near the same
spot. After the initial prick there is no irritation, and the
animal will stand quietly while the fly has its feed.” Stanley,
however [45], describes an occasion when, owing to the bites of
Tsetse, “‘ Horses and donkeys streamed with blood, and reared
and kicked through the pain”: and Rankin ['79], in his account
of the experiment made to test the value of elephants as baggage-
animals in East Africa, states that, “‘The fly swarmed on the
elephants till blood trickled down their flanks in a constant
stream.” The writer adds that although the elephants ‘‘ endured
this” for days, without subsequently showing any ‘“ prolonged
siens ” of Tsetse-fly disease, they ‘‘seemed pained and distressed
during the infliction” of the bites.

The time taken by the Tsetse in sucking its fill is remarkably
short. According to Andersson [20] it is but “a few seconds” ;
while Colonel Bruce sayst: “It is astonishing with what
rapidity the flies fill themselves; in as small a space of time as
twenty or thirty seconds a fly will become swollen out like a
balloon with bright red blood ” [see Fig. 6].

Contrary to what is the case among horse-flies (Tabanide), of

* Cf. Chapter VII., Appendix A, p. 272. ¢ Ibid, p. 271.

DISLIKE OF TSETSE TO FAECAL MATTER. 23

which the females alone suck blood, in the Tsetse both sexes are’
blood-sucking flies. This has at present been recorded only by
Colonel Bruce,* whose statement, however, is supported by the
fact that a male of Glossina longipennis, Corti, from Somaliland, in
the collection of the British Museum, has its abdomen swollen and
distorted, owing to containing decomposed blood, while a small
globule of dried blood may be seen adhering to the end of its
proboscis.

It is remarkable that ordure, which has
Dislike of Tsetse to 5 A
human habitations, 80 strong an attraction for many Diptera

and to animal (especially of the family Muscide, to which
excreta. . F

the genus Glossina belongs), has precisely the
opposite effect on the Tsetse. For this reason the fly appears to
avoid the presence of man, and is rarely found in the vicinity of
human habitations, or within the confines of a town or other
settlement.t Livingstone [21] alludes to “ the well-known disgust
which the Tsetse shows to animal excreta, as exhibited when a
village is placed in its habitat,” and states that the fact “has
been observed and turned to account by some of the [native]
doctors,” who protect animals about to pass through a fly-belt
by smearing them over with a mixture containing excrement
and other ingredients. Similar statements have also been made
by subsequent writers [cf. Kirk, 28, and Captain Gibbons, 152].
According to Fo& [135], on killing an antelope in a patch of
“Fly,” “in order to get rid of the Tsetse, which literally cover
game and hunters, it is only necessary to open the animal’s belly
and evacuate the entrails ; the insect at once ceases to torment
you.”
Until the publication of Colonel Bruce’s
“ Report,” the Tsetse was believed to lay eggs
and breed in the droppings of the buffalo, and various writers
have attempted to account in this way for the close association
that, in South Africa, at any rate, was alleged to exist between
the fly and that animal [cf. Bradshaw, 77, and Nicholls and
Eglington, 114]. But although the flies belonging to the genera
Stomoxys and Lyperosia, which are the nearest existing allies of
Glossina, undoubtedly breed in dung,f it will not surprise the

* Cf. Chapter VII., Appendix A, p. 273.

+ That this is not invariably the case is shown by the fact that the
British Museum collection contains specimens of Glossina pallidipes, Austen,
stated to have been ‘‘ caught in Witu town.”

{ Bouché (‘* Naturgeschichte der Insekten”’ (Berlin, 1834) pp. 55-56)
states that the larva of the common European Stomoxys calcitrans, Linn.,
is found in summer and autumn in horse-dung, in company with the larva

Reproduction.

24 MODE OF REPRODUCTION

reader, after what has been stated in the preceding paragraph,
to learn that the Tsetse has a totally different life-history, albeit
the details of the process are even more exceptional than could
have been anticipated.

According to Colonel Bruce: * ‘The Tsetse-fly does not lay
eggs as do the majority of the Diptera, but extrudes a yellow-
coloured larva nearly as large as the abdomen of the mother.
This larva is furnished with a black hood at one pole and two
minute spikes at the other. It is annulated and consists of ten
segments. Immediately on being born the larva creeps about
with a good deal of activity, evidently searching for some cover
or hole in which to hide. Having found a resting-place, it imme-
diately begins to change colour, and after a few hours has turned
into a jet-black hard pupa or nympha.

“Tf these pupal cases are placed in a perfectly dry place, as
in a wooden box, the perfect insect hatches out in about six
weeks. From this it would appear that the life-history of this
species of fly is very simple, it only being necessary for the female
insect to deposit the larva on the surface of the soil or in the

| grass, when the larva creeps into the nearest shelter, in a few
| hours becomes hard and black, and in five or. six weeks hatches
/ out into the fully-developed Tsetse-fly. It has often been sur-
“mised that this fly is bred in buffalo-dung, but from a consideration
of the foregoing facts it is evident that nothing is wanted except
any moderately dry place.”

Such are the details of the life-history of the Tsetse as pre-
sented by Colonel Bruce, and it will at once be perceived by
those who know anything of the metamorphosis of Diptera, in
general that they exhibit a remarkable similarity to what takes
place in the group of parasitic flies which, on account of their
peculiar mode of reproduction, have been termed Pupipara.t

of the house-fly (Musca domestica, Linn.), which it much resembles.
Lyperosia irritans, Linn., is said to breed in cow-dung. (Cf. ‘Insect
Life,” Vol. II. (1889), p. 97): in the paper referred to, the species is called
Hematobia serrata, Rob.-Desv.).

* Cf. Chapter VII., Appendix A, p. 271.

+ Of these the best known are the mammal- and bird-parasites belonging
to the family Hippoboscide (genera Hippobosca, Ornithomyia, Stenopteryx,
Oxypterum, Lipoptena, and Melophagus). Hippobosca equina, Linn., is a
well-known plague of horses and cattle in the New Forest; other species
occur in Africa and India, and are found on horses, cattle, camels, and
dogs; the genus Ornithomyia includes a number of species, all of which
are parasitic on various kinds of birds; Stenopteryx hirundinis, Linn.,
is found on the martin (Chelidon urbica, Linn.), Oxypterum pallidum,
Leach, on the swift (Cypselus apus, Linn.); Lipoptena cervi, Linn., is
parasitic. on the red- and roe-deer, while Melophagus ovinus, Linn., is the

POSSIBLY VARIABLE. 95

Further observations, however, on the reproduction of Tsetse-flies
are urgently needed, particularly with a view to discovering
whether the process is the same in all the species. From the
reproductions of photographs given by Colonel Bruce in his
‘Further Report,” one of which represents a fly in the act of
parturition, the species which formed the subject of his investi-
gations would appear to have been Glossina pallidipes, Austen,
rather than Gl. morsitans, Westw. It is therefore necessary
to point out that, from what we know of the life-history of other
Muscide, it must not be taken for granted that all species of
Glossina necessarily produce a full-grown larva, which imme-
diately assumes the pupal stage, since in some genera there is a
remarkable diversity in the mode of reproduction even in the
case of closely allied species. Thus, according to Portchinski,
the common European Mesembrina meridiana, Linn., is vivi-
parous, while Mesembrina resplendens, Wahlbg., and M. mystacea,
Linn., lay eggs; and whereas Musca domestica, Linn., lays
from 120 to 160 small eggs, Musca corvina, Fabr., either lays
about twenty-four large eggs, provided with peculiar appendages,
or, as was found by Portchinski to be the case in summer in the
Crimea, produces a single full-grown larva.*

so-called ‘‘sheep-tick.”” In all of these the female produces but a single
full-grown larva at a birth. While within the body of the mother the
larva is nourished by the secretion of special glands, and assumes the
pupal stage immediately after extrusion: unlike that of the Zululand
Tsetse, the Hippoboscid larva is incapable of movement. Bruce’s descrip-
tion shows that the larva of the Zululand Tsetse-fly must be very similar
in appearance to the newly-extruded larva of Hippobosca equina, which is
yellowish white in colour, with the terminal segment deep black. The
latter also exhibits two promimences, which, however, although distinct,
are not so pronounced as in the larva of the Zululand Tsetse. The mature
pupa of Hippobosca is dark brown, but shows no distinct trace of segmenta-
tion. In Hippobosca camelina, Savigny, which is parasitic on camels in
Northern Africa and in Asia, and is a much larger species than H. equina,
the protuberances at the posterior end of the pupa are scarcely distinguish-
able, although the dull black area is just discernible. In Oxypterwm
pallidum the pupa shows faint traces of the annuli belonging to the larval
integument, while the “black hood” takes the shape of a flat rosette-
shaped plate, devoid of prominences. rt

* Cf. Baron ©. R. Osten Sacken, ‘On Mr. Portchinski’s publications
on the larve of Muscide”: Berliner Entomologische Zeitschrift, Bd. xxxi.
(1887), pp. 17-28). According to Portchinski, Musca corvina in Northern
Europe deposits on dung about twenty-four large eggs with peculiar
appendages. In the Crimea in early spring its reproduction takes place 1n
the same way; but towards the end of spring, and in summer almost
exclusively, Portchinski found a single very large egg within a matrix-like
expansion in the body of the female. From this egg a larva was
developed which grew within the body of the mother, but passed directly
from the first to the third, or final stage (Cf. Osten Sacken, loc. cit., p. 26).
Another remarkable instance of larval development within the body of the

26 PUPA OF ZULULAND TSETSE.

From the examination of a series of Tsetse-fly pupe from
Zululand, kindly presented to the British Museum by Colonel
Bruce, I am enabled to give the following description of the pupal
stage (Fig. 7) in the only species of which the life-history has yet
been observed. As will be seen from remarks in Chapter IV.,*
the identity of the species is a little uncertain, though it must be
either Glossina morsitans, Westw., or Gl. pallidipes, Austen.

The extreme length of the pupa in this species varies from
64 to 7 millimetres (3 to 3} lines), while its greatest width is from
34 to 33 millimetres (13 to 1? lines). The colour is dark brown,
except in the case of the last segment which is deep black, repre-
senting what Bruce calls the “black hood” in the larva. In
shape the pupa is very remarkable, owing to the peculiar develop-
ment of the last segment. Instead of exhibiting the regular
contour seen in the Muscid pupa of the ordinary “barrel-
shaped” type (as in the house-fly, blow-fly, and green-bottle flies),
and although the main portion of the pupa is an elliptical oval
with the anterior end somewhat obtuse, similar to the pupe of
the types referred to, the posterior extremity (last segment) is

mother was discovered by Portchinski in the case of Dasyphora pratorum,
Mg., which also belongs to the Muscine, i.c., to the Muscidze in the most
restricted sense. The species in question deposits in the dung of cattle
a large larva, which is retained within a matrix-like receptacle in the
body of the mother until it reaches the third stage, After extrusion the
larva ‘‘very soon reaches its full growth and goes underground for its
further development” (Cf. Osten Sacken, loc. cit., p. 25). As to the
connection between the mode of development of the Pupipara and that
of the flies just mentioned, Osten Sacken writes as follows (loc. cit., pp.
26-27) :—‘‘ Hitherto the Pupipara had an isolated position among the
Diptera. The modes of larval evolution of Musca corvina and Dasyphora
pratorum, discovered by Mr. Portchinski, bridge over the interval. He
even ventures the hypothesis that the Pupipara began by being
coprophagous in their larval state, and laid an almost full-grown larva
like that of the two above-mentioned flies. Later, owing to the parasitic
mode of life of the imago and the diminution of the powers of flight
necessary for providing the proper environments for the larve, those
changes in the mode of evolution of the larvee were produced which
distinguish the Pupipara now.” It is not yet known whether in the
Tsetse and the other Muscide, which produce.a single more or less adult
larva at a birth, the oviduct of the female is provided with glands by the
secretion of which the larva is nourished, as in the case of the so-called
Pupipara; but it must be remarked that, whatever be the case with regard
to the latter, Portchinski’s hypothesis will not serve to explain the origin
of the mode of development of the Tsetse, as exhibited by the Zululand
species, which presents an even closer approximation to that of the
Pupipara, than does the development of either Musca corvina or Dasyphora
pratorum. For the Tsetse, although feeding on blood, is not parasitic in
the ordinary sense of the term, while its powers of flight have been in no
way affected by its mode of life,
* See page 89.

PUPA OF ZULULAND TSETSE. 27

produced on each side into a prominent tumid lip. These lips
enclose a deep pit, within which in the larval state were situated
the posterior stigmata or respiratory apertures (Fig. 7a).

Fig, 7,

Pupa of Zululand Tsetse-fly, dorsal ke (X 12); a, posterior extremity,
showing pit and right larval stigma, st (x 24); b, anterior extremity,
showing bifurcated longitudinal seam, which opens to permit the
escape of the imago (xX 8).

The pupa consists of twelve segments, divided by fine grooved
rings as if traced with the point of a needle. The first eleven
segments are smooth, but when examined under a low power
are seen to have a shagreen-like or tessellated surface. At
the anterior pole the seam which opens to permit the escape
of the fly is visible as a semicircular groove, running across
the first three segments in a lateral longitudinal direction,
and terminating on each side in a bifurcation on the fourth
segment (Fig. 7b). In the centre of the first segment (which is
much smaller than any of the others), at the anterior pole, a

28 PUPA AND LARVA.

slight depression is noticeable, representing the mouth of the
larva. When the pupa is viewed from above or below (see
Fig. 7) the lateral tumid lips, into which, as already stated,
the Jast or twelfth segment is produced, are very conspicuous.
They are connected by a dorsal and a ventral ridge, and lips and
ridges together enclose the above-mentioned pit, the depth of
which is from 1} to 2 millimetres. In this pit can be seen the
vestiges of the posterior stigmata of the larva, one on each side
at the base of the lateral lip. Under a platyscopic lens, magni-
fying about ten diameters, each stigma has the appearance of a
minute closed pore, surrounded by a slight elevation (see
Fig. 7a). Each lip is covered externally with minute tubercles
or granules, and its edge bears four more or less clearly marked
grooves or furrows. The anterior margin of the last segment is
slightly rugose, with small longitudinal furrows, as shown in
Fig. 7.

The examination of the pupa renders it possible to supplement
or amend, in one or two respects, Colonel Bruce’s description of
the larva. According to Bruce the latter consists of ten seg-
ments, but the examination of the pupa shows that in reality it
is composed of twelve. In a specimen which is practically a
dried larva, since pupation is evidently not complete, there are
strongly marked depressions between the segments producing a
conspicuously annulated appearance, which agrees with Bruce’s
description of the larva. Bruce states that the larva is provided
with “ two minute spikes ” at the end opposite the “black hood”;
the latter is represented in the pupa by the deep black twelfth
segment, with its prominent lips. The “minute spikes” of the
larva would therefore seem to be the mouth-hooks of the ordinary
Muscid maggot, which in the Tsetse are perhaps used by the
larva, to attach itself to the wall of the oviduct of the mother. In
Plate I. of Colonel Bruce’s “‘ Further Report,” the pupa is likewise
shown. equipped with a similar pair of hooks at the anterior pole.
But this must be a mistake; the pupze before me display no
trace of any such structures, and Bruce himself says nothing
about them.

While most other genera of blood-sucking

tell agg Diptera are practically world-wide in their
occurrence, Tsetse-flies are confined to Africa,

and a glance at the accompanying map will show the reader what
has been learnt of their distribution since Sir William Cornwallis
Harris [5] encountered Glossina morsitans to the south of the

A

AHARA.

medge of the

|

s included in
ork, and from
ecimens con-
sum and other

Rudolf. a

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MembaB. 4
i FmOZAM = MojangaBay, :

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North Latitude “ae

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a ri
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30°

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Startords Geog. Establishment

~ fe Br.

AFRICA | ep | pte ove,
SOUTH OF THE SAHARA. ell “hE OT meth mim BD Vue (SESS NG Carl ae

Illustrating present knowledge of the
distribution of

GLOSSINA,

as compiled from records included in

the Bibliography to this work, and from

local data derived from specimens con-

tained in the British Museum and other
collections.

Prandin &

| 4 uy 0ssessiOns ~ nf farce t4 2 Chapainy (i
pon : Pp. ee C diaper j 7
| Pas ¢
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so = = <4
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Showing distribution of { 4 i
GLOSSINA p II
Seale, 1:30.096,000, 475Enghsh Miles to Linch. NI I
Seale of English Miles é
= ft

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100-8 100 200 300 400 500 600
Tristan da Cunha , Telegraph Cables, us...
(British) Ee

20° ¢ “a a 10° WiGr. O° ‘E.Gr. 10 20 20

GEOGRAPHICAL DISTRIBUTION. 29

Limpopo in 1836. For reasons which have already been ex-
plained, however,* it does not necessarily follow that the Tsetse
is to be found to-day in the precise localities coloured red on the
map, and this applies more especially to regions south of the
Zambesi. In this portion of the Continent, owing to the advance
of civilisation and the retreat or disappearance of the big game,
many tracts in which the fly formerly abounded are now entirely
or to a large extent free from it; so that in order to learn what
is known of the extent of the fly-belts in South Africa at the pre-
sent time, the latter portion of the first part of the Bibliography
must be consulted, in addition to the records of the localities of
specimens given in Chapter IV.

With this reservation, therefore, it may be stated that so far
as can be ascertained at present, either from the examination of
actual specimens or from records in literature, the genus Glossina
ranges from the Gambia in the west (approximate latitude
13° 66’ N.) to Somaliland in the east, extending south from this
line until it reaches its southernmost limit in the northern
portion of Zululand, near St. Lucia Lake (approximate latitude
27° 60'S.). Quite recently it has been recorded from the region
of Lake Tchad [171], but as yet we do not know whether it
occurs in the Egyptian Sudan or in the river valleys and ravines
of Abyssinia ; although from what we are told as to the mortality
among horses and mules during the Abyssinian Expedition of
1867, it is probable that it is indigenous in the latter country,
at any rate [cf. 171]. There is, however, a doubtful record
from Sennaar [47], and, should this be confirmed, the northern
limit of the Tsetse would be represented by a line running from
that country through Lake Tchad to the Gambia, or roughly by
the 13th parallel of north latitude. But it is almost certain
that the genus Glossina will ultimately be found to extend
farther north than this, and whether it occurs or not on the
White and Blue Nile up to the latitude of Khartum, it can
searcely fail to do so on the Senegal River and throughout the
valley of the Niger.

It will be seen from the map that records of localities north of
the Equator are at present very scanty, but that they become far
more numerous in the eastern half of the southern portion of the
Continent. On the western side, although the Tsetse no doubt
occurs throughout Portuguese West Africa, there appear to be
no records whatever of its having been met with south of the

* See pp. 10, 11.

30 GEOGRAPHICAL DISTRIBUTION.

Congo. Turning to German East Africa, we find that records
are fairly numerous, but, as the map shows, they are far more
abundant than anywhere else in the valleys of the Zambesi and
Limpopo, which, as has been stated by many previous writers,
are the main strongholds of Glossina morsitans. Although still
or formerly occurring extensively in the northern and eastern
portions of the Transvaal, the Tsetse, as already explained,* does
not enter the Orange River Colony ; but Glossina morsitans, pro-
bably in company with Gl. pallidipes, abounds in Zululand, at
least in the northern portion of that country between the Lebombo
Mountains and St. Lucia Lake.

In conclusion, it may be stated that there can be little doubt
that species of Glossina will eventually be found to occur at
intervals throughout Tropical Africa, wherever the conditions are
suitablet and there is a chance of obtaining food.

* See page 7.

+ See page 7. While the Tsetse recedes before the advance of
civilisation, this is not the case with regard to other blood-sucking
flies, some of which may ultimately prove to be capable of conveying
Trypanosoma brucei (see Chapter VII., Appendix E, page 803). It would
therefore seem to be highly desirable that investigations should be set on
foot without delay with a view to determining the geographical distribution
of the latter; that is, that the endeavour should be made, on the lines of
Colonel Bruce’s inoculation experiments, to discover in what regions of
Africa the parasite of Tsetse-fly disease is naturally present in the blood
of wild animals. For this purpose, as pointed out by Colonel Bruce (ef.

Chapter VII., Appendix A, p. 286) ail wild animals should be examined,
and not merely those usually entitled ‘‘ game.”

3l

CHAPTER II.
HISTORICAL SURVEY.

I.—Bionomic.

In the minds of most people Tsetse-flies appear to be especially
associated with the name of David Livingstone, as though
the celebrated missionary-explorer had been the first to discover
and bring them to notice. This assumption, however, is not
borne out by facts. As may be seen from the “ Bibliography ”
(Chapter VI.), and as will be shown below, Livingstone’s works
contain numerous statements concerning the Tsetse, while at the
outset of the expedition which resulted in his death among the
swamps of Lake Bangweolo Livingstone made a particularly
interesting experiment with a view to ascertaining whether
Indian buffaloes and camels were proof against the disease
disseminated by the fly. But, as will be seen later on, references
to the Tsetse-fly and its habits had appeared in works published
before the year 1857 (the date of the publication of ‘ Missionary
Travels”), while for the earliest notices of the havoc wrought by
the fly * we must go back nearly three hundred years.

According to Sir Harry Johnston (145, 160]{ we read in
Portuguese records how the “earlier Portuguese expeditions of
five hundred or six hundred mounted men, which would set out
from Sena on the Lower Zambezi in the 16th and 17th centuries
to secure the gold mines to the north and south,” were brought
to ruin owing to the deaths of the horses from the bite of a fly,

* In a-chapter dealing with the history of our knowledge of the Tsetse-
flies it is impossibie to avoid constant reference to J’setse-fly disease,
although, since the epoch-making work of Lieut.-Col. Bruce (see
Chapter VII., Appendix A), the earlier theories as to the etiology of the
disease possess a purely historic interest. 3

+ The numbers in square brackets refer to the “ Bibliography ”
(Chapter VI.).

32 “TSALTSALYA” OR “ZIMB” OF JAMES BRUCE.

which there can be little doubt was the Tsetse. Sir Harry
Johnston proceeds to show [145] that—‘‘ But for the Tsetse-fly,
the whole history of South-Central Africa would be different.”

Returning to the nineteenth century, we find as early as the year
1813, in the well-known “Travels to Discover the Source of the
Nile,” of James Bruce [1], statements concerning the “ T'saltsalya
or Fly,” of which the modern Arabic name is said to be Zimb.
This insect, of which a rough wood-cut is given, is stated to be
destructive to cattle during the rainy season on the Upper
Atbara River, on the confines of the Sudan and Abyssinia. All
that can be said, after an examination of the description and
figure, is that the insect is either a Tsetse or a horse-fly (Family
Tabanide) belonging to the genus Pangonia. James Bruce con-
sidered that it was the “fly” referred to in Isaiah vii, 18 and 19
[see 1], and Westwood [10], writing thirty-seven years later, was
of the opinion that Bruce’s ‘“‘Zimb” was at any rate a species of
Glossina, if not actually GI. morsitans. If this is so, it is inte-
resting as showing that the range of the genus extends to the
Sudan on the Abyssinian frontier.

Robineau-Desvoidy [3], writing in 1830, expressed his belief
that the proboscis of Vemorhina (Glossina) palpalis (see below,
p. 71) was not used for sucking blood, an opinion that, as we
shall see, was subsequently shared by his compatriot Macquart.*

So far as it has been possible to discover, the earliest English
writer to refer to the Tsetse-fly in South Africa was Captain
(afterwards Sir) William Cornwallis Harris [5], who, in his “Wild
Sports of Southern Africa,” published in 1839, states that the
Mural Berge, a range of hills on the south side of the Limpopo,
in the Waterberg district of the present Transvaal, are, especially
during the rainy season, ‘infested by a large species of gad-fiy,
nearly the size of a honey-bee, the bite of which . . . . proves
fatal to cattle.” In spite of this misleading description, and
although the Tsetse is not mentioned by name, there can be no
doubt that it is the fly referred to.

* In a letter written in 1901, Mr. F. J. Jackson, C.B., the well-known
African eportsman, gives it as his opinion that the Tsetse is, “like the
mosquito, only a blood-sucker by predilection” (see Chapter VII.,
Appendix C, p. 297), by which he doubtless means that in default of blood
itis able to exist on the juices of plants. In support of his view he
mentions that in the fly-belt near Kibwezi, British East Africa, in April,
1892, when the whole district was ‘“‘ parched and dried up,”.and ¢onse-
quently destitute of game of any kind, he found the Tsetse more plentiful
than he has ever known them before or since; but he does not say that
he has ever seen a Tsetse-fly imbibing vegetable juices.

GORDON CUMMING MENTIONS TSETSE. 33

Writing in 1843 of the genus Glossina, of which the only
species then known was Gil. longipalpis, Wied., the French
dipterist Macquart was led [6], owing to the fineness of the
mouth-parts, to infer that the flies were not blood-suckers, but
subsisted upon the nectar of flowers. Macquart expressed his
opinion in the following terms: ‘‘I] est probable qu’il ne vit pas
du sang des animaux comme les Stomoxes, mais du suc des fleurs.
Les deux soies que renferme la trompe, et qui constituent le
sucoir, sont d’une finesse telle qu'il est difficile de concevoir
qu’elles puissent percer la peau, et la faiblesse de cet organe
parait encore démontrée par la modification des palpes, qui
s'alongent et se creusent pour lui former un fourreau” (op. cit.
p. 113). Macquart gave some exceedingly poor figures of what
he called Glossina longipalpis in the work in question, and in the
“Supplément” [8] published in 1850.

In the latter year R. Gordon Cumming [9], who had hunted
in the districts previously visited by Captain Cornwallis Harris,
first mentioned by name “the fly called ‘Tsetse,’” which he
referred to a little later as “the famous fly called ‘tsetse,’ whose
bite is certain death to oxen and horses.” Gordon Cumming
described the “ hunter’s scourge” as “similar to a fly in Scotland
called ‘kleg,’ * but a little smaller,” and gave the earliest pub-
lished details of the effects of its bite on horses. From his use
of the word Tsetse it is evident that it was the name given to
the fly either by Boer hunters or by the natives who at that
time inhabited what is now the Zoutpansberg district of the
Transvaal. In the same year (1850) Westwood [10] published
his well-known paper on the genus Glossina, in which Glossina
morsitans and two other species (though these prove to have
been previously described) were named and described (vide
infra). Westwood quoted a letter from Major Vardon (a well-
known sportsman and contemporary of Gordon Cumming), from
whom he had received his specimens of Gl. morsitans, and
endeavoured to identify the “ Tsetse” of South Africa with the
“ Zimb” or “ Tsaltsalya” of James Bruce.

Notes on the Tsetse-fly were published in 1852 and 1853 by
William Cotton Oswell [11, 15, 18], who had hunted with
Major Vardon, while a French writer, M. Arnaud [12], having
examined a specimen of Glossina morsitans brought home by
Oswell, expressed the opinion that it was identical with a fly met
with by himself in Sennaar, where its effects on cattle appear to

* Hematopota or Tabanus spp. (Fam. Tabanide).—H. FE. A.
D

34 ANDERSON, LIVINGSTONE, BURTON, KIRK.

be similar to those mentioned by James Bruce in the case of
the “Zimb” on the Upper Atbara, and, during the months from
January to May, the season at which it is most troublesome,
cause the natives who keep cattle to behave in a similar manner
to those referred to by Bruce.

In 1856 C. J. Andersson [20] gave ‘details as to the habits
of Gl. morsiians, with particulars of losses among domestic animals
caused by it,—details which were considerably amplified in Living-
stone’s “ Missionary Travels” [21] of the following year. Bracy
Clark [22], writing in 1857, and doubtless carried away by his
mterest in his favourite Gistride (Bot- and Warble-flies), foolishly
endeavoured to show that the Tsetse is identical with MWstrus
(Hypoderma) bovis, one of the two common cattle Warble-flies of
Europe. On October 5th of the same year, Westwood [24]
exhibited a specimen of Glossina morsitans, brought home by
Major Vardon, at a meeting of the Entomological Society of
London, and took occasion to refute Bracy Clark’s contention ;
but ‘himself repeated the ‘curious mistake first made in his paper
of 1850, by ‘stating that Gl. morsitans was from Lake Tchad.
In the following year L. de Castelnau [23] described the habits
of Glossina morsitans, ‘and endeavoured to define its geographical
limits; he also gave a curious account, apparently on the
authority of Mr. Edwards, “the companion of Mr. Chapman,”
of an experiment by Bushmen to prove their assertion that the
Tsetse-fly is viviparous.

Between 1860 and 1865 statements as to the occurrence
and habits of the Tsetse in different parts of Africa appeared in
records of travel by various writers, including Captain R. F.
Burton [25], Thomas Baines [26], and David ‘and Charles
Livingstone [27]; Captain Burton, who met with the fly in the
course of his ‘expedition from Bagamoyo to Lake Tanganyika,
being the first to record its occurrence in East Africa,
Dr. (now Sir) John Kirk’s* paper, “On the ‘Tsetse’ fly of
Tropical Africa” [28] is the best réswmé of our knowledge of the
habits of the fly yet published. Sharing in the opinion then
prevalent, that the fatal results of the bite of the Tsetse in the
case of domestic animals were due to ‘a poison, elaborated by the
fly itself ‘and injected into the wound at the moment of biting,
Dr. Kirk dissected the bulb at the base of the proboscis to see
whether it contained a poison gland. Those who know anything
of the internal anatomy of Diptera will scarcely need to be told

* Dr, Kirk had-been botanist to Livingstone’s Zambesi expedition.

SCEPTICISM AS TO POWERS OF TSETSE. 35

that he failed to find any such structure, since the salivary
glands, the secretion from which produces the irritation which
is experienced after a stab from a biting fly, are always situated
in the thoraa.

Sir Samuel Baker [29], writing in 1866, mentioned the
Tsetse-fly as occurring in Obbo, to the south-east of Gondokoro,
a new locality. James Chapman, in his “ Travels in the Interior
of South Africa” [30], published in 1868, gave copious details of
the occurrence, habits, and effects of the Tsetse in the country
to the south of the Zambesi, mentioning among other things an
instance of acquired immunity to Tsetse-fiy disease in the case of
Makoba dogs, which “from time immemorial” have been bred
by natives in the Fly country.

An early letter from Karl Mauch [36], the German pioneer-
explorer of the gold-fields of the northern Transvaal and
Mashuna Land, and subsequent discoverer of the ruins of
Zimbabye, published in 1869, is interesting from the fact that
it mentions an external application of Asafcetida, or internal
application of ammonia as possible remedies for Tsetse bite.
In the same year Mr. and Mrs. Petherick [37] encountered
the Tsetse in the country of the Rhol Tribe, north-west of
Gondokoro.

We now meet with the earliest expression of a certain
scepticism as to the reputed effects of the bite of the Tsetse
upon domestic animals, which, appearing first in the year 1870,
continued side by side with the orthodox belief until well into
the eighties. Mr. St. Vincent Erskine [38], an African traveller
well known at the time, addressing a meeting of the Natural
History Association of Natal on May 380, 1870, is reported to
have “‘combated the popular idea that the bite of the Tsetse-fly
was destructive to the life of certain animals, especially the ox,
horse, and dog.” Mr. Erskine’s theory “was that the deaths of
the animals were to be attributed more to change of grass or
climate than to the bite of such a small fly as the Tsetse.” No
arguments are recorded in support of this belief, which is
evidently due to a revolt against the popular idea already
referred to, that the deaths of domestic animals bitten by the
Tsetse were caused by a poison secreted by the fly itself. The
belief in the existence of a specific Tsetse poison may be said to
have prevailed until the publication of the results of Col. David
Bruce’s investigations in 1897 (see Chapter VIT., Appendix A),
though, as we shall shortly see, the true state of the case was

D2

36 ALLEGED IMMUNITY OF DONKEY.

hinted at less than ten years after the date of Erskine’s paper.*
St. Vincent Erskine’s theory, although attacked shortly after-
wards by E. C. Buxton [41], was warmly espoused by the well-
known English entomologist, Edward Newman [39, 44], who
considered the Tsetse to be a ‘‘ myth.”

A second letter from Karl Mauch [40], published in 1870,
contained an early expression of the commonly accepted idea
that the donkey is immune to Tsetse-fly disease. Writing of the
behaviour of a donkey in the Fly country in the Northern
Transvaal, Mauch says:—“ The main advantage, however, con-
sisted in the fact that the Tsetse could not do her any harm,
whether it be that the donkey finds in certain leaves or in the
bark of certain trees an antidote against the poison, or that the
long hair or the effluvium from the beast repels the insect.” | In
this letter Mauch also recorded the uselessness of sal ammoniac,
administered internally, as an antidote to Tsetse-fly disease in
cattle. A map by A. Petermann to illustrate Mauch’s travels,
published in the same volume as the letter, shows the “ Limit of
the Tsetse-fly” in the vicinity of the Limpopo, and also to the
south of the Zambesi.

Eduard Mohr [42] met with the Tsetse in 1870 to the south
of the Victoria Falls, but Otto Kersten [43], who took part in
Baron C. C. von der Decken’s travels in East Africa from
1859 to 1865, did not encounter it between the coast and
Dschagga (the region of Mt. Kilima Njaro), though later on he
mentions its occurrence on the Lower Juba River as the reason
why the Wabuni, a scattered tribe of Galla, are unable to keep
cattle.t Stanley [45], in ‘‘ How I Found Livingstone,” published
in 1872, recorded his experience with Tsetse-flies, and mentioned
native names under which they are known in different regions.

* Livingstone himself [21] appears to have partially realised the truth.
After describing the effects of the bite of the Tsetse-fly in cattle, he
proceeds to say (‘‘ Missionary Travels,” p. 82):—‘‘These symptoms seem
to indicate what is probably the case, a poison in the blood; the germ of
which enters when the proboscis is inserted to draw blood. The poison-
germ, contained in a bulb at the root of the proboscis, seems capable,
although very minute in quantity, of reproducing itself,...” It is
evident, however, that Livingstone supposed the fly to possess a poison-
gland, situated in the bulb at the base of the proboscis, the secretion of
which was fatal to cattle.

+ Col. Bruce’s experiments showed that Tsetse-fly disease is invariably
fatal to the donkey in Zululand (see Chapter VII., Appendix A).

+ Probably this is the earliest definite reference to the Somaliland
Tsetse-fly, described long afterwards by Corti as Glossina longipennis :
if so it is interesting as evidence that a species of Glossina distinct from
Gl. morsitans is able to carry the hematozoon, (Cf. Chapter VIL,
Appendix E.)

EXPERIMENT WITH INDIAN BUFFALO. 37

E. Marno [47], writing in 1873, stated that Tsetse-flies had
been brought to him in Sennaar as being included among the
flies known there by the name “Surréta.”* The occurrence of
a species of Glossina in Sennaar, however, needs confirmation
[ef. 12]. Marno adopted much the same view as St. Vincent
Erskine [38—vide supra], expressing the opinion that the bites
of flies, whether called Tsetse or Surréta, are only one, and
‘‘perhaps even a subordinate factor,” in causing the mortality
among imported domestic animals, which occurs in certain parts
of Africa either throughout the year, or only during the season
called Charif in Sennaar, and is actually due to adverse climatic
conditions.

“The Last Journals of David Livingstone” [49], edited by
Horace Waller, and published in 1874, are of especial interest
from our present point of view, since they contain the account
of the buffalo and camel experiments already referred to.
Unfortunately the experiments were to a large extent vitiated
by the ill treatment of the animals by sepoys, whom Livingstone
had brought with them from Bombay. The “Journals” contain
many references to the Tsetse-fly on the Rovuma River and
elsewhere. In the same year Karl Mauch [50] repeated the
statement that donkeys and goats are immune to Tsetse-fly
disease, and went on to say :—‘‘Only one remedy appears to be
effective, and that is based upon homeopathic principles ; the
fly itself, taken internally, makes the punctures innocuous, as I
have seen in the case of a dog,...” The use by natives of
the Tsetse-fly itself as a means of making animals immune to
the disease, has been referred to by various writers [Cp. 27, 30,
55, 97].

In 1876, E. Mohr [52] recorded an instance in his own
experience of the apparently curative effect of an internal
application of ‘‘strong doses of eau de l’huis, or extract of
ammoniac,” in the case of horses suffering from Fly disease,
and also stated that goats are immune.

Dr. Hartmann [53], in the course of remarks on the Tsetse-
fly addressed to the “ Gesellschaft naturforschender Freunde ”
of Berlin, on July 17, 1877, mentioned the occurrence of a species
of Glossina (considered by the lecturer to be identical with Gi.
morsitans, Westw., but from his description almost certainly either
Gl. palpalis, Rob.-Desv., or GI. longipalpis, Wied.) in Loango,

* Blood-sucking horse-flies (Tabanidx) are commonly spoken of as
“Seroot”’ flies by Englishmen on the White Nile.

38 DR. DRYSDALE’S HYPOTHESIS.

where he stated that Tsetse-fly disease was apparently unknown.
The lecturer’s views as to what he considered to be the exagge-
rated statements concerning the harmful effects of the bite of the
Tsetse-fly, as also of the Diptera known under the name Surrita
(and other variants of the same word), were identical with those
of Marno (vide supra).

In 1877 Dr. Joseph Mullens [54] gave an account of an
interesting experiment with draught oxen on the road between
Saadani and Mpwapwa, which seemed to have proved satisfactorily
that the Tsetse-fly did not occur on that section of one of the
possible routes to Lake Tanganyika. The same year saw the
publication of Thomas Baines’ “ Gold Regions of South-Eastern
Africa” [55], which gave numerous details as to the occurrence of
Glossina morsitans in the Transvaal, with notes on the way in which
it may be distinguished from other flies, the effects of its bite in
cattle, as summarized by Henry Hartley, a well-known elephant-
hunter and pioneer at that time, on the effects of the external
application of ammonia as a remedy in the case of fly-bitten
horses, etc. Baines included mules, donkeys, sheep and goats
among the animals “believed to be unaffected by the virus,”
though he proceeds to add that mules are not always safe, and
that “Mr. St. Vincent W. Erskine doubts the safety of the
donkey on the south-east coast.” From the latter remark it
would appear that by this time Erskine had seen fit to change
his ideas with regard to the effects of the Tsetse’s bite.

_ In “Through the Dark Continent” [56], published in 1878,
Stanley mentions having encountered the Tsetse during his journey
down the Congo, on the islands below Rubunga. In a paper
by F, B. Fynney [57], which appeared in the same year, the
author stated that during the preceeding six years the Tsetse had
disappeared from many parts of the Transvaal, formerly known
as Fly Country, and remarked that it was hardly necessary to
attach so much importance to the presence of the Tsetse as was
usually done, ‘because the fly is merely a temporary and
ephemeral scourge, and always disappears with the large game.”

The first definite suggestion of the true part played by the
Tsetse-fly in connection with the disease which bears its name
appeared in 1879, in the report of an address delivered by
Dr. J. J. Drysdale [58] to the Literary and Philosophical Society
of Liverpool, where it is stated that the Tsetse-fly “may be the
intermediate host of some ... . blood-parasite; or it may be
the carrier of some infective poison. It is highly improbable

STATEMENTS BY BRADSHAW AND SELOUS. 39

that any mere poison or venom should exist so powerful as to
cause the death of a large animal in such small dose.” It will
be seen that in his first hypothesis Dr. Drysdale came very near
the truth, though Col. Bruce’s investigations, with the subsequent
papers by Laveran and Mesnil [XXI.], and Plimmer and Brad-
ford [XX.] on the life-history of Trypanosoma brucei, would seem
to show that the Tsetse-fly is in no sense its host: recent German
writers on Tsetse-fly disease, however, incline to the opposite
view. As we shall find, between this date and that of Col.
Bruce’s memoir, various writers published notes expressing practi-
cally the same view as Dr. Drysdale concerning the etiology of
Tsetse-fly disease.

Captain J. F. Elton [59], writing in 1879, mentioned the
Tsetse as abounding near Livingstonia, at the south end of Lake
Nyasa;* and in the same year Sir Rutherford Alcock [60]
stated that the Tsetse-fly “on the road to Mpwapwa” had
destroyed the draught cattle belonging to a party sent by the
London Missionary Society to Lake Tanganyika [Cf. 54]. <A
note printed in the Proceedings of the Royal Geographical
Society [62] about the same time records the fact that bullocks
were working satisfactorily on the first section of the road then
being made from Dar-es-Salaam to the interior, and adds, “ It is
therefore clear that there is no Tsetse-fly on the forty miles
already traced of the route... .”f <A letter published in
“The Times,” of February 25th, 1879, by Lewis Hornor [64]
recommends the Boer method of crossing fly-belts at night—a
mode of avoiding the attacks of the fly that, from the experience
of subsequent travellers, is not invariably effective [Of. 77, 110].

Various books and papers [66-75] published during the next
two years (1880-1881), and dealing mainly with South Africa,
contain statements of the occurrence of the Tsetse-fly in different
localities.

Selous’ “Hunter’s Wanderings in Africa” [76], one of the
classics of South African sport, and a paper on “ The Tsetse Fly,”
by Dr. B. F. Bradshaw [77], both of which appeared in 1881,
include many interesting details. The experience of both writers
had been gained in the same country, viz., the vicinity of the

* Sir Harry Johnston [145], writing in 1897, states that the south
coast of Lake Nyasa is almost entirely free from Tsetse. ,

+ Cf., however [71], where it is stated that the Tsetse-fly occurs on this
road forty miles from the coast, rendering the employment of horses and
bullocks out of the question: “the belt of fly-country is, in fact, here
wider and more continuous than it is further to the north.”

40 EXPERIMENT WITH INDIAN ELEPHANTS.

Chobe and Zambesi, and therefore the species to which the
statements of both refer is probably Glossina morsitans.

The failure, owing to Tsetse-fly, of the London Missionary
Society’s endeavour to use bullock waggons on the road to Lake
Tanganyika in 1877 was recorded by E. C. Hore [78] in 1882,
and the same writer also gave details of the occurrence of the
fly on the shores of Lake Tanganyika itself. In the same year
L. K. Rankin [79] described the results of a highly interesting
attempt, which was considered to have resulted in success, to
discover whether Indian elephants, employed as baggage-animals
on the march from Dar-es-Salaam to Mpwapwa, would withstand
the attacks of the Tsetse-fly—in this case probably Glossina
pallidipes, Austen.

Dr. E. Pechuél-Loesche [80], writing in “Die Loango Expedi-
tion” (1882), stated that a Tsetse-fly had been collected near
Tschintschotscho, about a hundred miles north of the mouth of
the Congo. The species is not mentioned, but possibly it was
either Gl. palpalis, Rob.-Desv., or Gl. longipalpis, Wied. Since,
however, cattle are kept both at Landana and Boma in the same
district, and not only roam about freely, but also thrive toler-
ably well, either the fly does not occur near those towns or else
the Trypanosome of Tsetse-fly disease is absent ; or, if present, is
not carried by the local Tsetse-flies.

In ‘The Wild Tribes of the Soudan,” by F. L. James [81],
published in 1883, mention was made of a disease called guffer,
which attacks camels in the district to the east of Kassala, and
was stated by some of the natives to be “caused by the bite of
the Tsetse-fly during the rainy season.” There is no record,
however, of a Tsetse-fly actually having been captured in this
region. In the same year a Swiss entomologist named Schoch [83],
in a well-reasoned paper, advanced arguments for considering
that the Tsetse-fly is not in itself poisonous, but “at most the
carrier of a bacterium-like poisonous matter.” Nearly every
previous writer had referred to the Tsetse-fly as though but a
single species existed, but Schoch speaks of ‘“‘ two long-winged
Muscide of similar appearance, one as large as our house-fly, the
other somewhat larger, and less dreaded.” Apparently, there-
fore, the writer in question was acquainted with Glossina fusca,
Walk., though he gives no authority for stating it to be “less
dreaded” than GI. morsitans. Various earlier writers had hinted
at the probable disappearance of the disease, as the Tsetse either
followed or became extinct with the big game, but Schoch

FLY-DISEASE IN SOMALILAND. 41

attained a higher and more scientific standpoint by his conclusion
that, “with more intimate knowledge of the active principle, the
disease can be successfully opposed, and will gradually disappear.”
Although this consummation has not yet been reached, and
though no antitoxin for the effects of Trypanosoma brucei in
domestic animals has so far been discovered, the lines of practical
research were thus foreshadowed by Schoch a dozen years before
it was actually attempted, and his name will deserve to be
remembered when the investigations that are now being under-
taken are at length crowned with success.

Passing over one or two records of the occurrence of the
Tsetse in different localities, as well as another attempt to show
that the belief in the harmfulness of the fly is mere prejudice
[84], the next publication of importance appeared in the year
1884 in the shape of a paper on the Tsetse-fly, by the late Dutch
entomologist, F. M. van der Wulp [88], which also included a
short bibliography. From a report of further remarks by the
same writer [91], published in the following year, it is evident
that he sided with those who doubted the stories of the effects of
the bite of the Tsetse-fly in domestic animals. In the same year
the Austrian dipterist, Josef Mik, also recently deceased, in an
abstract [90] of Schoch’s paper pointed out the existence of what
were supposed to be three additional species of Glossina.

Among records of the occurrence of Tsetse-flies in different
localities, published during the year 1885, a paragraph in a
paper by F. L. James [94], entitled, “A Journey through the
Somali Country to the Webbe Shebeyli,” is worthy of notice
from three points of view. The writer states that the Adone,
the people living on the Shebeyli, “have large herds of cattle
and flocks of sheep, but all these animals are poor and suffer
from the fly in the rain and from the ticks in the dry season ;
neither camels nor horses are used, for they will only liye in the
dry season... .” In this passage the Tsetse is not referred to
by name, but we now know that Glossina longipennis, Corti, the
Somaliland Tsetse-fly, occurs on the Webbe Shebeyli, and it
would seem, therefore, that we have here a definite indication
that the species in question is a carrier of the hematozoon.
Secondly, the statement that sheep suffer from the attacks of the
fly is of interest, since various writers have mentioned native
sheep and native goats as animals immune to Tsetse-fly disease.*

* Col. Bruce was unable to obtain native sheep and goats for his
experiments in Zululand,

42 GEOGRAPHICAL RECORDS.

The mention of ticks is of interest in connection with the know-
ledge that the parasite of Texas fever among cattle is dissemin-
ated by these animals; although it is scarcely likely that ticks
can act as carriers of Tsetse-fly disease, it would perhaps be worth
while to settle the point by actual experiment.

“De Angola A Contra-Costa,” by H. Capello and R. Ivens
[96], published in 1886, contains a chapter (illustrated with
rough wood-cuts of the head and foot enlarged) on the Tsetse-
fly, entitled ‘A Tzé-Tzé”; but, since it is in Portuguese, the
present writer has unfortunately been unable to examine its
contents. In the same year interesting details of the occurrence
of the Tsetse in Mashuna Land and the region between the
Zambesi and Lake Nyasa were given by W. M. Kerr [97, 98}
who also described the means adopted by the native women in
Mashuna Land for making dogs, goats, and sheep—‘‘ of which
they have very few, seemingly only kept as pets”—immune to
Tsetse-fly disease when young.

In 1887 Josef Chavanne [99] declared that the Lower Congo
must be free from Tsetse-fly, since cattle throve fairly well in
that region. The same year saw the publication of the second
edition of Justus Perthes’ special large-scale map of Africa, in
which the limits of the Tsetse-fly in the vicinity of the Limpopo
are indicated by dotted lines. In 1887, too, Dr. C. W. Schmidt
[ror] recorded the fact that the Tsetse-fly did not occur in the
districts of Usambara and Bondei, near the Pangani River,
German East Africa, and that consequently it was possible to
keep cattle on a large scale in Usambara.*

The ranks of those who considered that the disease produced
by the bite of the Tsetse-fly in domestic animals was not due to
a poison secreted by the fly itself received a further accession in
1888, when A. Laboulbéne [102] expressed the opinion that the
Tsetse “carries with its proboscis septic matter drawn from
diseased animals, and communicates it to others that are
healthy, the result of which is illness and possible death. It
remains to be explained why the bite, which is said to be deadly
to cattle, is not so to human beings.” In the same year, in a
communication to the Royal Geographical Society from Dr. Holub
[104], allusion was made to the abundance of Tsetse-fly in
forests, consisting of small trees, in the Batoka country;
F. Jeppe [105] recorded the partial disappearance of the fly,

* A recent paper by Stuhlmann [XXVI.] would seem to show that nowa-
days, at any rate as regards Usambara, this is a mistake,

LUGARD ON TSETSE IN EAST AFRICA. 43

together with the big game, from the Kaap Valley in the
Transvaal, as a result of gold-mining operations; and the
ninth edition of the ‘‘ Encyclopedia Britannica” gave a short
résumé of existing knowledge of Glossina morsitans, and its
distribution.

In 1889 the occurrence of ‘a yellowish-brown Glossina”
(doubtless Gl. fusca, Walk.) in the forest at Barombi Station,
Cameroons, was recorded by Dr. Preuss [107]; and in 1890
Dr. Holub [110] gave a number of details of the occurrence
and effects of the Tsetse in the region of the Zambesi. Among
other statements, Dr. Holub gave his reasons for thinking that,
under certain circumstances, trekking at night with waggons
through a fly-belt may not be such a safeguard as is generally
imagined, and also expressed the opinion that the diminutive
size of the cattle kept by the Matoka is due to the influence of
the fly.

“Ten Years in Equatoria,” by Major Gaetano Casati, pub-
lished in 1891, contains the statement that the raising of cattle
by the Mambettu, a tribe living to the west of Lake Albert
Nyanza, is impossible owing to the presence in their country ‘of
a fly called Tsetse, the stings of which cause death... .” In
the same year E. A. Maund [112], writing ““On Matabele and
Mashona Lands,” repeated the oft-made assertion that the Tsetse
is destined to disappear with the game, and also mentioned the
fact, previously recorded by Kerr (vide supra), that the
Mashonas administer the dried and pounded fly to their dogs as
a prophylactic against the disease. According to Maund the
medicine is given on the extremely restricted scale of “a fly a
day”!

Captain (now Sir) F. D. Lugard [115] writing in 1893,
emphasised the superior advantages possessed by British Hast
Africa over South Africa, owing to the fact that the former
country is “practically free from the Tsetsefly.” He also
showed that German East Africa is much worse off, since there,
according to Dr. Pruen, ‘the Tsetse-fly holds undisputed sway ”
throughout the whole of a broad zone bordering the coast : the
fly-belt known to exist near Taita is then discussed. It is
perhaps worth while to remark that the knowledge accumulated
since the publication of Sir Frederick Lugard’s book shows that,
especially in the north-eastern portion of British East Africa, the
Tsetse-fly is more prevalent than the distinguished administrator

supposed,

44 GEOGRAPHICAL RECORDS: OSWELL.

During the same year (1893) there appeared a number of
other records of the occurrence of Tsetse-fly in various localities,
chiefly in East and South-east Africa. The late Mr. Theodore
Bent [116], in a newspaper interview, mentioned the importance
of the Beira Railway, in rendering it possible to cross without
difficulty the Tsetse belt through which it passes ; while Major
J. J. Leverson [118] gave his experience of the fly in the same
region, on and near the Pungwe River. Mr. F. J. Jackson,
C.B. [119], also in a newspaper interview, discussed the merits
of alternative routes to Uganda, with special reference to the
presence or absence of the Tsetse-fly, which, as was also stated
by Captain Foster [120], infests some sixty miles of the Sabaki
River route, near the Tsavo. Mr. F. C. Selous [12r], in his
volume entitled “ Travel and Adventure in South-East Africa,”
gave numerous details of the occurrence of the fly in Southern
Rhodesia ; and the existence of two species of Glossina in the
hinterland of the German Protectorate of Togo, West Africa,
was recorded by Dr. Karsch [122].

In a letter from Dr. Donaldson Smith [124], published in
1894, mention is made of “the dreaded camel-fly of the Webi
Shebeli ”—doubtless the same species as that referred to nine
years earlier by Mr. F. L. James [94—vide supra]. In the same
year Commissioner (now Sir Harry) Johnston [125] dealt with
problems raised by the presence of Tsetse-fly in British Central
Africa, while its occurrence in British East Africa was referred
to by Sir Gerald Portal [126] and Mr. F. J. Jackson [127].
The latter writer mentioned the interesting fact that, although
it is supposed by many people that the Tsetse exists only
“where game beasts, especially buffaloes, are most plentiful,”
and though this may be the case in South Africa, in East
Africa at any rate the fly-belts, “with the exception of the
River Tana,” “are almost devoid of game.”* The late William
Cotton Oswell [128], writing in the same volume as Mr. Jackson
on “South Africa Fifty Years Ago,” mentioned his experience
of Glossina morsitans in that region in the middle of last century,
when he hunted with Vardon and was the companion of
Livingstone on his earlier travels.

In 1894, too, Jules de Guerne [129] allied himself with
Schoch [83--vide supra] by stating that :—“ Everything leads

* For further information as to the ability of Tsetse in Hast Africa
to exist independently of game, the reader is referred to the letter
from Mr, Jackson printed in Chapter VII., Appendix ©, p. 295.

FLY-BELT ON BEIRA RAILWAY. £5

us to believe that Glossina inoculates domestic animals with the
germs of some virulent malady analogous to anthrax.”

The year 1895 witnessed the publication of Surgeon-Major
(now Lieut.-Col.) Bruce’s “ Preliminary Report on the Tsetse-fly
Disease” [131], which afterwards was practically embodied in his
“Further Report” of two years later. In 1895, too, one or two
interesting notes on different species of Tsetse-flies appeared in the
Geographical Journal. What appears to have been the Somali-
land Tsetse-fly [vide supra, and 94 and 124], Glossina longzpennis,
Corti, is recorded to have been met with in the previous year in
the Korayo Valley, Somaliland, where its bite proved fatal to
baggage animals belonging to Major Mainwaring [133]. The
second note, by W. A. Eckersley [134], refers to Tsetse-fly in
connection with the Beira Railway, then in course of con-
struction. . The fly is stated to have made its appearance at
Chimoio, in Portuguese East Africa, as a result of the presence
there ‘(of a large number of horses, oxen, and other animals

., attracted thither by the facilities of transport offered by
the railway ....” Another interesting statement by Mr.
Kekersley is that a couple of ponies, ‘purchased in Natal for the
use of the survey party, passed through Beira, Fontesvilla, and
the intervening ‘fiy-belt, to Chimoio without suffering any ill
effects; they served the party until the conclusion of the work,
and, were finally sold at a profit. No particular precautions
against the ‘fly’ were adopted, except occasional brushing witli
green boughs. It is quite certain that the Tsetse-flies settled
on the horses in considerable numbers, and remained quite long
enough to allow of their biting.” Since it is stated that the
country traversed by the first twenty miles of the railway was,
at the period referred to, “teeming with big game, including
lions, buffaloes, most of the South African species of antelope,
wart hogs and wild boars,” it would. almost seem as though the
hzematozoon of Nagana must, for some reason or other, have
been absent from the blood of the wild animals near the course
of the line.

M. Edouard Fod’s volume [135] on his experiences of big
game shooting, also published in 1895, contains many observa-
tions on the Tsetse-fly, including an interesting description of its
method of feeding.

In the following year a review of Bruce’s “ Preliminary
Report” was published by W. F. H. Blandford [136], who laid

stress on the importance of studying the bionomics of the para-

46 COLONEL BRUCE'S FURTHER REPORT.

site rather than continuing investigations upon purely clinical
lines. Other publications of the year 1896 worthy of notice from
our present point of view are a further Report by Sir Harry
Johnston [138] dealing with the British Central Africa Protec-
torate, and “The Great Rift Valley,” by Dr. J. W. Gregory [139].
According to Sir Harry Johnston the Tsetse-fly is already fast
disappearing from British Central Africa, and ‘‘the unchecked
increase of the negro population” should be encouraged as a
means of hastening its “entire extinction.” Dr. Gregory described
how, in the Tsetse-haunted belt of forest between Witu and the
coast, he endeavoured to protect his camels by the Galla method
of raising clouds of smoke.

With the publication in 1897 of Col. David Bruce’s ‘‘ Further
Report on the Tsetse-Fly Disease or Nagana, in Zululand” [142],
not only was the disease conclusively proved to be due to a
hematozoon, but also, by means of a brilliant series of pre-
liminary experiments, the part played by the Tsetse in trans-
mitting the malady was finally established. Since the Report
will be found fully analysed elsewhere (see Chapter VII,
Appendix A), there is no need to say more about it here.

From this time onward the literature of the Tsetse-flies con-
sists mainly of statements in books of travel and Government
Reports. These statements contain much additional information
as to the geographical range of the genus Glossina, but do not
add greatly to our knowledge in other respects. Of the further
contributions furnished by the year 1897, the most important is
that contained in Sir Harry Johnston’s work entitled “ British
Central Africa” [145], wherein several pages are devoted to the
Tsetse-fly. The problem raised in West Africa by the occurrence
of several species of Glossina was alluded to in the same
year by the late Miss Mary Kingsley [148], who pointed out
that it would be useless to make a practicable road for draught
animals from the Gold Coast to its hinterland, “because of the
horse-sickness and Tsetse-fly which oceur as soon as you get into
the forest behind the littoral region.” . In the course of remarks
on the Tsetse-fly in “The New Africa,” by Messrs. Schulz and
Hammar [149], also published in 1897, the interesting statement
is made that in 1874 donkeys safely performed the journey from
Delagoa Bay to Lydenburg, in the Transvaal, “when that
country was thickly infested with flies that killed cattle, horses,
and even the few camels that were imported as an experiment.”
Tt is admitted, however, that donkeys succumb like other animals

RINDERPEST AND TSETSE. 47

when the Tsetse is present in overwhelming numbers. Further
on it is stated that the goats which the authors had with them
showed no signs of being affected by the fly, and their immunity
to the disease is considered to be established.*

The deaths of cattle and horses from the bites of Tsetse-flies
in the Aulihan country, Somaliland, in January, 1896, was
recorded by Prince Nicholas Ghika [150] in 1898. In the same
year Mr. A. H. Neumann [151], in his volume on “ Elephant-
Hunting in East Equatorial Africa,” stated his experience of
Tsetse-flies in British East Africa and on the shores of Lake
Rudolph ; in the latter locality he encountered one of the larger
species, doubtless Glossina fusca, Walk. Captain Gibbons [152]
gave many details as to the occurrence of the Tsetse in Rhodesia,
and both this writer and Captain the Hon. Arthur Lawley [153]
—the latter in a magazine article referring to the same country —
dwelt on the close association of the Tsetse with the big game.
According to Captain Gibbons, “ where the wild buffalo is to be
found in large numbers the Tsetse invariably teems”; while
Captain Lawley states that in the neighbourhood of the River
Deka, since the game has been nearly exterminated by rinder-
pest, “the fly also has left the country, excepting a few belts of
thick bush, and apparently it is gradually disappearing.” As we
have seen, according to Mr. F. J. Jackson [127, vide supra], the
fly in East Africa is not so intimately associated with big game.
Other contributions in 1898 included notes by Mr. W. W. A.
Fitzgerald [154] on the Tsetse-fly in the coast lands of British
East Africa ; and some interesting observations in a Government
Report on the progress of the Uganda Railway [155], in which
it is stated that, of the transport animals engaged in supplying
the advanced parties with food and water mules were found less
susceptible to Tsetse bite than either camels, donkeys, or bullocks.
In the same year Dr. Garry de N. Hough [156] identified two
specimens collected by Dr. Donaldson Smith in Somaliland in
August, 1894, as Glossina longipennis, Corti.; while a short
article in the “Globe” newspaper [157], of July llth, 1898,
stated that a malady, which had ‘played havoc” with the
horses and transport animals of the Royal Niger Constabulary
at Lokoja, had been identified as Tsetse-fly disease. It was also

* The question of the immunity of the goat to Tsetse-fly disease has
Tong been’a moot point, though more than twenty years ago it was stated
by Selous [76] that goats, other than animals whose ancestors had been

bred in the Fly-country for generations, enjoy no special immunity
whatever.

48 NATIVE THEORY: TSETSE IN XVI. CENTURY.

stated that “the Lokoja natives, knowing nothing of microbes
and bacteria, assert that the fly extracts from a certain small red
monkey the virus with which it inoculates the bush cow or dwarf
buffalo.” Though this is very possibly a simple folk-tale, it
perhaps suggests the advisability of examining the blood of the
South African baboon (Papio porcarius, Bodd.) to see whether
Trypanosoma brucei can be detected in it.

Among contributions published in 1899, we may notice a
statement contained in Sir Harry Johnston’s “History of the
Colonization of Africa by Alien Races” [160], with reference to
the destruction, apparently by the Tsetse-fly, of the horses and
camels of an early Portuguese expedition which proceeded from
Quelimane to Sena in 1569. This has already been alluded to
at the commencement of this chapter (see p. 31). In 1899, too,
W. Harvey Brown [161] wrote of the difficulty caused by the
Tsetse-fly in the importation of machinery into many of the
mining districts of Rhodesia, before the construction of the
Buluwayo Railway ; while the present writer reported the occur-
rence of Glossina longipalpis, Wied. (really Gl. palpalis, Rob.-
Desv.), in the vicinity of Freetown, Sierra Leone.

In 1900 Mr. C. V. A. Peel [163] narrated his experience of
Glossina longipennis in Somaliland, and gave an interesting
description of the sensation of suddenly entering a “ fly-belt.”
In the same year Mr. H. A. Bryden [165] repeated the statement
of so many previous writers on sport in South Africa, by assert-
ing that “wherever the African buffalo is plentiful, there you
will almost certainly find the Tsetse-fly,” adding that when the
buffalo was killed off or driven out the fly disappeared. The
Bushman story that the fly breeds in buffalo droppings is referred
to as possibly accounting for its special association with the
animal, but since Bruce’s description of the mode of reproduction
we know that belief to be a myth. According to Bryden, a line
of coaches from the Pungwe River towards Mashonaland, estab-
lished “when Rhodesia was first being opened up,” had to be
abandoned owing to the deaths of the horses and mules from the
bites of the Tsetse-fly.

In 1901 Sir Harry Johnston [166], in a Report on the Pro-
tectorate of Uganda, referred to the length of time formerly
taken on the journey to that country from the coast before the
construction of the Uganda Railway, since it “had to be per-
formed mainly on foot owing to the difficulty of conveying riding
animals through the belt of country near the coast infested with the

RESULTS FROM UGANDA RAILWAY. 49

Tsetse-fly.” In the same year Dr. Max Schoeller [167], writing
on German East Africa, suggested that should the domestication
of zebras for transport purposes prove feasible, those animals
would be of much more use than horses or donkeys owing to
their immunity from Tsetse-fly disease. Dr. Schoeller also dis-
cussed the question of the possible identity with the Tsetse of
the “‘ Wandorobo-fly ” of the Wadschagga, the bite of which was
stated by the latter to have been responsible for the death of his
donkeys, though attempts to obtain specimens of the fly proved
unsuccessful, The author in question seems inclined to believe
that his donkeys really died from a form of the disease known to
the Boers as “ Dikkopziekte”—the South African horse-sickness.
Among other publications of 1901 to be noticed, is a “ Report by
His Majesty’s Commissioner on the East Africa Protectorate”
[168], in which it was stated that since the advent of the Uganda
Railway horses can be brought up from the coast to Ukamba for
travelling purposes, while formerly it was the custom in that
country “to make long marches on foot,” owing to the impossi-
bility of conveying horses from the coast through the intervening
Fly-belts. ‘The Commissioner supports Dr. Schoeller [167, vide
supra| by advocating the domestication of the zebra and the
wildebeeste, and for the same reason, adding that “ the prosperity
of the country is at present sadly hampered by want of efficient
transport.” In the same year Mr. and Mrs. Hinde [169], in
“The Last of the Masai,” expressed the fear that the establish-
ment of the Uganda Railway might contribute towards the final
extinction of big game “in the last stronghold left in Africa,”
since now that the barrier formed by the Tsetse-fly was no longer
an obstacle it was possible for “sportsmen” to enlist the aid
of the horse. Towards the end of 1901 Dr. Schilling {170|
recorded the presence of a disease which is fatal to horses and is
called by Schilling “Surra” (no doubt in agreement with Koch,
who pronounced T'setse-fly disease in East Africa to be identical
with Surra in India), in the German Protectorate of Togo, on
the Slave Coast, West Africa, where, as is shown in Chapter IV.
of the present work, Glossina morsitans occurs, as well as GI.
palpalis and GI. fusca. Schilling was unable to make experiments
with the fly, but described the Trypanosome which causes the
disease.

Turning to the year 1902, an important work by MM.
Laveran and Mesnil [171] must be noticed, since besides treating
of the hematozoa of Nagana and kindred diseases, it contains

E

50 PROBLEMS FOR INVESTIGATION.

various statements on the distribution of Tsetse-flies and of
the eponymic malady disseminated by them. From Sir Harry
Johnston’s great work on “The Uganda Protectorate” [172],
published a few months ago, it appears that although “ flies of
the genus Glossina” * are found in that country, Tsetse-fly
disease is absent, a statement that suggests the advisability of
prosecuting a careful enquiry into the geographical distribution
of Trypanosoma brucei without delay.| Reference may also be
made to a recent statement by Mr. J. N. Justice [173], who, in
a paper on “ Prospecting in Northern Rhodesia,” mentions that,
according to the natives, “since the rinderpest exterminated the
buffalo, the Tsetse has lost much of its venom... .” This is
merely a reassertion in another form of the old stories of the
specially close connection between the Tsetse and the buffalo,
which,.as may be seen from the letters printed in Chapter VIL,
Appendix C, is not admitted as regards East Africa by some
of those best qualified to express an opinion on the subject.
Finally, we may notice two papers [KXVI., 174], by Dr. Franz
Stuhlmann on Tsetse-flies (Glossina morsitans and Gl. fusca) and
Tsetse-fly disease. (which the author, like other recent German
writers, calls “ Surrah”) in German East Africa ; in the former
paper Dr. Stuhlmann gives a map to ‘iden the occurrence
of the disease in the region in question.

Tn conclusion, then, it will have been seen from the foregoing
résumé, that while in the space of sixty odd years our know-
ledge of the Tsetse-flies has gradually increased, we have still
much to learn. It is only within the last few years that the
-remarkable mode of reproduction has been made known, and it
has yet to be discovered whether this is the same in all species.
Very little is known as. to the distribution of certain species ;
and, most important of all, we are still in almost complete
ignorance as to whether all the. species are equally capable of
conveying the parasite of Tsetse-fly disease (see Chapter VIL.,
Appendix EK, p. 300).

II,—Systematic.

Although the name “Tsetse” did not become generally
known in England until the year 1850, when the habits and
ravages of “the fly called ‘Tsetse’” were mentioned by R.

* Probably Gl. fusca, Walk., and perhaps Gl. pallidipes, Austen.
+ Cf. Chapter I., p. 30, note +.

HISTORY OF GENUS GLOSSINA. 51

Gordon Cumming [9], while the species to which the name was
first applied was soon afterwards described by Westwood [10]
under the well-known designation Glossina morsitans, the genus
Glossina dates from twenty years earlier. In the year 1830,
therefore, the genus Glossina was founded by Wiedemann [2]
for his species Gl. longipalpis, the type of which was stated to
have been obtained at Sierra Leone by Adam Afzelius.*

In the same year, 1830, a second species of Tsetse-fly was
described by Robineau-Desvoidy [3] from a specimen from the
Congo, under the name Nemorhina palpalis. It has since been
customary among Dipterological writers to regard Nemorhina
palpalis, Rob.-Desv., as a synonym of Glossina longipalpis, Wied. ;
but, as is shown in Chapter IV., the two species are really quite
distinct. This raises the question of priority between Glossina
and Nemorhina, since both names date from the same year: the
question, however, in default of anything in the shape of internal
evidence, is now incapable of solution, and in the present work
no attempt has been made to upset existing nomenclature so
far as the genus is concerned, since to abolish a name so well
established in literature as Glossina would be for every reason
eminently undesirable.

After 1830 no addition was made to the genus Glossina for a
space of nineteen years, until in 1849 Glossina fusca (Stomoays
fuscus) was described by Francis Walker [7], from a single
female from an unknown locality. In 1873 this species was
sunk by its author [46] as a synonym of Gi. longipalpis, Wied.,
though, as is shown in the systematic portion of this work
(Chapter IV., p. 100), it is in reality perfectly distinct and must
be maintained.

The next species to be described were Glossina morsitans,
tachinoides, and tabanzformis, all of which date from Westwood’s
paper in the Proceedings of the Zoological Society for 1850 [10].
Of these three species, as is shown in Chapter IV., GJ. tachinoides,
from West Africa, is nothing more than a variety of GI. palpalis,
Rob.-Desv., while Gl. tabaniformis, from the Gold Coast, is a
synonym of Gl. fusca, Walk.

After the appearance of Westweod’s paper came an interval

* A Swedish botanist, who visited what was at that time the newly-
founded settlernent of Freetown, Sierra Leone, so long ago as 1792 and
1794, and made general collections of objects of Natural History. The
existence of a Tsetse-fly in Sierra Leone was not recognised, however,
until the present writer found Gl. palpalis, Rob.-Desv. (at the time
suppesed to be Gl. longipalpis, Wied.) there in 1899,—C/. [162].

BE 2

62 SPECIES ADDED IN LAST HALF-CENTURY.

of five-and-thirty years, during which no addition was made to
our systematic knowledge of the genus Glossina, though in the
Appendix to Oates’ “ Matabele Land and the Victoria Falls”
[73] a coloured figure of Gl. morsitans, and details of the antenna,
mouth-parts, and tarsus were given by Westwood[74]. In 1885
a supposed new species of Glossina was described by the late
Mons. J. M. F. Bigot [89], under the name Gl. ventricosa, and
the description was preceded by a table (in French) for the
determination of this and five other previously described species
of Tsetse-flies, The writings of this author, however, were
seldom distinguished by critical acumen, and the table in
question, being based merely upon descriptions, is of little value.
As to Glossina ventricosa, an examination of the typical speci-
mens shows that they are nothing more than examples of Gi.
palpalis, Rob.-Desv., with their abdomens distended — with.
coagulated blood. Bigot’s suggestion that his supposed new
species hails from Australia will be found discussed below under
GL. PALPALIS—“ Synonymy and Affinities” (Chapter IV., p. 78) :
as will be seen, there are in reality no grounds whatever for
supposing that Glossina occurs outside Africa.

The next two species of Glossina—Gl. grossa and Gi. palli-
cera—were described, likewise by Bigot [113], in 1891. The
types of both of these are from Assinie, Ivory Coast, and their
author suggested that both might be merely varieties of
Glossina longipalpis, Wied., by which he meant Gl. palpalis,
Rob.-Desv. ; but, as will subsequently appear (Chapter IV.),
Gl. grossa is actually a synonym of Gil. fusca, Walk., while
Gl. pallicera is a perfectly good species.

Of the remaining species, Glossina longipennis, Corti, the
Somaliland Tsetse-fly, was described [130] in 1895 from a
specimen from the Boran Galla country, on the confines of
Abyssinia and Somaliland ; while the last on the list, Glossina
pallidipes, is now described in Chapter IV. for the first time,
having hitherto been confounded with GI. morsitans, Westw.,
with which it is apparently associated to some extent in

geographical range.

CHAPTER Tr:

SYSTEMATIC POSITION OF THE GENUS.

THE object of the present treatise being essentially practical, it
would be out of place to enter at length into questions of classi-
fication ; but enough must be stated to enable the reader to
understand what are the nearest allies of the Tsetse-flies among
existing insects. Suffice it then to say at once that the genus
Glossina belongs to the calyptrate division of the Family Muscipz%
(Muscaria Scuizomeropa, of Brauer and von Bergenstamm) of
the Order Diptera, or Two-winged Flies. Diptera may be
characterised shortly as insects with only one pair of wings, a
thorax fused into a single mass, and mouth-parts adapted for
piercing and sucking, or for suction alone. The metamorphosis
is complete; the larve are totally different in appearance from
the perfect insects, and, though varying greatly in outward form,
are usually footless grubs ; those of the Muscide are generally
known as maggots. The pupa either shows the appendages of
the perfect insect, although these are enclosed in a sheath and
adherent to the body, or else it is entirely concealed within the
hardened and contracted larval integument, which forms a pro-
tecting case. Diptera have been divided by Professor Brauer
into two great groups, termed Orthorrhapha and Cyclorrhapha,
according tothe manner in which the pupa-case splits to admit
of the escape of the perfect insect. In the Diptera Orthor-
rhapha, which include the less specialised families, such as the
Cecidomyide (Gall-Midges), Mycetophilide (Fungus - Midges),
Culicide (Gnats or Mosquitoes), Tipulide (Daddy Long-Legs),
Tabanidz (Horse-flies), Asilidee (Robber-flies), etc., the pupa
case splits longitudinally down the median dorsal line. In the
Diptera Cyclorrhapha, on the other hand, which comprise the
most highly specialised families, such as the Syrphide (Hover-
flies), istridee (Bot- and Warble-flies), and Muscide (sensi latiore
—House-flies, Tsetse-flies, Blue- and Green-bottle-flies, Flesh-

54 FAMILY” MUSCID.

flies, Flies of which the larve are internal parasites of cater-
pillars, etc.), the imago is enabled to escape by the splitting off
of a cap from the cephalic end of the puparium.

The family Muscide in its widest sense is by far the most
numerous in genera and species of the whole order. It is
divided into two main divisions, Calyptratz and Acalyptrate ;
in the former of these the sguama, or membranous scale, situated
on the thorax behind and beneath the base of the wing, is
sufficiently large to conceal the halter, the little club-shaped
organ that projects beneath it ; in the Acalyptratie, on the other
hand, the squama is so reduced in size that the halter is not
concealed when the insect is looked at from above. The
Muscide Acalyptratz include a very large number of small flies,
in which there is a great diversity of habits, especially in the
larval state, though the majority of the perfect insects resemble
the Common House-fly more or less closely in outward form,
though not necessarily in colour or markings. The group is split
up into a number of sub-divisions, which are regarded by some
writers as sub-families, while others assign to them at all events
nominal family rank. The Muscide Calyptrate are generally
speaking flies of larger size and stouter build than the Acalyptrate.
The group is divided into a smaller number of sub-divisions than
the Acalyptrate, and in its widest sense (= Muscaria Schizo-
metopa, of Brauer and von Bergenstamm), includes the nominal
families Tachinide (bristly flies, the larvee of which are parasitic
in caterpillars and certain other insects), Dexide, Sarcophagidie
(marbled grey Flesh-flies), Muscide (House-flies and their blood-
sucking allies belonging to the genera Stomoxys, Hematobia,
Lyperosia and Glossina, Blue-bottles, Green-bottles, etc.), Gistridie
(Bot- and Warble-flies), and Anthomyidz (non-bristly Muscide,
in which the first posterior cell of the wing is wide open).

While there can be no doubt that it is impossible to find
hard and fast characters, which would warrant the maintenance
of the independence of the six so-called families that have just
been enumerated, there is fortunately no occasion to enter here
into the vexed question of Muscid taxonomy, which has engaged
the attention of various well-known dipterists during the last
few years. As has already been pointed out by the late F, M. van
der Wulp,* the genus Glossina was placed by its founder, Wiede-
mann, in the immediate neighbourhood of Stomowys, Geoff., and
if we add to the group Hematobia, Rob.-Desv., Lyperosia, Rond.,

* “Tijdschrift voor Entomologie,” xxvii. (1884), p, 144.

NEAREST EXISTING ALLIES OF TSETSE. 55

and Beccarimyia, Rond., the nearest allies of the Tsetse-flies
have been designated. The genera Stomoxys and Hzematobia
include several species of small brownish or mottled grey and
brown flies, with a prominent though short chitinous proboscis,
which are greedy blood-suckers, and plague both men and cattle.
Stomoxys calcitrans, Linn., which is the only European species
of its genus, though others occur in Africa, is very common: in
England in August and September, sitting about in numbers on
rails and gates in fields. Hzematobia stimulans, Mg., and Lyperosia
(Heematobia) irritans, Linn., are common summer plagues of cattle
in England; and the latter, known to American writers’ as
Hematobia serrata, Rob.-Desv., has been introduced into the
United States, where, owing to its habit of clustering about the
bases of the horns of cattle, it is commonly called the “ Horn-
fly.” Another species of Lyperosia, closely allied to L. irritans,
but smaller and with narrower palpi, was found by Mr. W. R.
Ogilvie Grant swarming on his camels in the’ Dimichiro Valley,
Sokotra, in January, 1899.* The genus Beccarimyia, founded by
Rondani,t for the new species B. glossina, from Keren, about
seventy miles from Massowah, differs from Hmatobia and ‘the
other genera in having a very prominent epistoma, and the first
posterior cell of the wing closed before the margin.

The genus Glossina is placed by Brauer and von Bergenstamm {
in the “Subsectio Sromoxys,” of the “ Sectio Muscina.”§ The
other genera of which the ‘“Subsectio Sromoxys” is formed are :
Beccarimyia, Rond., Stomoxys, Geoffr., Hematobia, Rob.-Desv.,
and Lyperosia, Rond.; the latter, which is sometimes regarded
as a sub-genus of Hematobia, was founded|| for Conops (Stomoxys,
Heematobia) irritans, Linn. There is no doubt that this grouping
represents the most commonly accepted ideas of the’ affinities of
Glossina. In respect of certain structural details (e.g. the venation

* Lyperosia minuta, Bezzi (Ann. Mus. Civ. Genov.,” .xxxii. (1892),
p. 192), occurs in Somaliland, and two other species are found in Ceylon.
One of the latter, which closely resembles L. minuta,; Bezzi, but is probably
new, has recently been stated to be very troublesome to ponies at the
Ceylon breeding station.

+ ‘Annali del Museo Civico di Storia Naturale di Genova,” iv. (1873),

. 287.

fi t “‘Vorarbeiten zu einer Monographie der Muscaria Schizometopa
(exclusive Anthomyide), Pars III.’’: “ Denkschriften der mathematisch-
naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissen-
schaften,” lx. Band, Wien, 1893, pp. 177-178. 5

-§ Subsequently termed by Prof. Brauer (‘‘ Verhandlungen der k. k.
zoologisch-botanischen Gesellschaft in Wien,” Jahrgang 1893, p. 516), the
“ Sectio Musca.” a

|| Rondani, ‘‘ Dipterologiz Italicze Prodromus,” Vol. I. (1856); ‘p. 93.

56 STOMOXYS, AND OTHER ALLIES.

of the wings, the branched hairs on the arista, the bulb at the
base of the proboscis, and the remarkable male genitalia), Glossina
occupies a position altogether unique.* On the other hand, if
we consider the characters afforded by the proboscis and the
feathering of the arista, the affinity between Glossina and the
genera with which it is associated in the system of Brauer and_
von Bergenstamm appears to be incontestable. The blood-
sucking habit is common to all, and correlated with this we find
a rigid, horny proboscis, which is carried in a horizontal position.
The palpi, however, which in Glossina are so remarkably modified
to form, in the living insect, a perfect sheath for the proboscis,
are in Stomoxys of the ordinary Muscid type, being exceedingly
slender, short (not projecting beyond the buccal cavity), and in
no way affording any protection to the proboscis. In Hxmatobia,
as represented by H. stimulans, Mg., the palpi, while apparently
capable of forming a partial sheath for the proboscis, are dis-
tinctly shorter than that organ and expanded at the tips, while
the tip of the proboscis itself bears a few short hairs, and the
modified fleshy labella are plainly visible—as, though on a
smaller scale, they also are in the proboscis of Siomoxys. So far
as can be judged from Rondani’s description and figure,f the
palpi and proboscis of Beccarimyia resemble those of Heematobia
stimulans, though the palpi do not appear to be expanded at the
tips. Lastly, in Lyperosia, Rond., as represented by Lyperosia
(Hematobia) irritans, Linn., of Europe and North America, the
palpi, though flattened from side to side, as in Glossina, are not
expanded at the tips, while they form a complete sheath for the
proboscis, which they equal in length. As to the proboscis itself,
though the tip still bears a few minute hairs, the labella are
so reduced as to be practically invisible, their position, when
examined under a low power, being merely indicated by a slight
constriction, as in the case of the proboscis of Glossina.

So far as regards proboscis and palpi, therefore, Glossina
approaches Lyperosia more nearly than any other genus yet
mentioned. Moreover, in Hematobia the arista bears three or four
short hairs on the under side as well as those on the upper, but
Lyperosia $ agrees with Glossina in having the arista feathered

* The aberrant mode of reproduction described by Col. Bruce (ef.
Chapter I., p. 24) forms a further proof of divergence, and constitutes an
approximation, so far as it goes, to the group of Diptera known as the
Purpipara (Hippobosca, Ornithomyia, etc.).

+ “Ann. Mus, Civ. Genoy.’’ iy. (1873), p. 287.
} As also Stomoxys.

GIRSCHNER’S CLASSIFICATION. 57

only on the upper side ; while in the closeness of the feathering
and the curvature of the hairs on the arista Lyperosia also
approaches more nearly to Glossina than does Hxmatobia.

In the general shape of the head, on the other hand, and in
the relative width of the front in the male, Glossina agrees better
with Stomoays than with any of the other genera already referred
to, while differing from all of them in having the vibrissal ridges
ciliated nearly to the middle, and in being destitute of post-
humeral bristles.*

According to Girschner’s most recent exposition of his system
for the classification of the Muscide,f Glcssina and its allies
belong to the ‘Section Muscine aricieformes” of the Family
Anthomyide (Girschner, nee Avuct.). In an earlier paper by the
author in question,{ it is stated that the primary object of
the macrochetz (large bristles) on the pleure of Muscids is
the protection of the stigmata or respiratory apertures. The
Calyptrate Muscide may be divided into two great groups
according to the presence or absence of macrochete on the
hypopleure, protecting the metathoracic stigma ;§ and Glossina
and its allies belong to the group, which also includes the
Anthomyine of authors, in which the hypopleure are bare.
From the generalisation that, “We may rightly regard those
forms as the more perfect (evolved at a later date), that, while
having a similar mode of life, are provided with the best protective
arrangements,” || Girschner proceeds to argue that the allies of
Cenosia, Aricia, Anthomyia and Cyrtoneura (the group, therefore,
that in his system would include Glossina) are on a lower
developmental plane than, e.g., the allies of Sarcophaga, Dexia

* The term post-hwmeral is here employed in the sense in which it was
originally used by Mik (‘ Verh. z.-b. Ges. Wien,” xxx. Bd., Jahrg. 1880
(1881), p. 350, fourth line from the top), and afterwards by Girschner
(‘‘ Berl. Ent. Z.” xxxviii. Bd. (1893), p. 298), and Hough (‘‘ Proc. Acad.
Nat. Sci. Philadelphia,” 1898, p. 176); not as used by Osten Sacken
(‘‘Essay of Comparative Chetotaxy,’ ‘Trans. Ent. Soc. Lond.” 1884,
p. 508): cf. Mik, ‘‘ Verh. z.-b, Ges. Wien,” xxxii. Bd. (Jahrg. 1882), p. 13.
Osten Sacken’s post-humeral bristles are in the present work termed
notoplewral, in accordance with Girschner (loc. cit.).

+ E. Girschner, “ Hin neues Musciden-System”: “ Illustrierte
Wochenschrift fiir Entomologie,” I. Bd. (1896), pp. 12-16, 30-32, etc.
(Classification of Muscide according to sete, etc., characters of families).—
Unfortunately I have not been able to consult this paper at first hand,
and so have had to rely upon the partial résumé furnished by Hough
(‘‘ Biological Bulletin,” Vol. I. (1899), pp. 19-20).

¢ “Beitrag zur Systematik der Musciden”’: ‘‘ Berliner Entomologische
Zeitschrift,” Bd. xxxviii. (1893), p. 299.

§ Loc. ctt., pp. 297, 298.

|| Loc. cit. p. 299.

58 SPECIALISATION IN WING OF GLOSSINA.

and Echinomyia, since the former have an exposed, the latter a
protected metathoracic stigma. In venation, however, Glossina
is more specialised than the Anthomyine that in Girschner’s
arrangement are grouped with it, since it is provided with a
* Spitzenquerader” (bent-up terminal portion of the fourth vein),
which, joining the margin of the wing before the apex, leaves
the latter entirely membranous ; this condition is in Girschner’s
opinion a protection to the veins in most intimate connection
with the respiratory system.* In respect of venation, too, as
also in having the membrane of the wings rilled instead of smooth,
Glossina is more specialised than its immediate allies, Stomoxys,
Hematobia and Lyperosia, in which the third and fourth veins
both reach the apex of the wing, the fourth vein being only
slightly bent up at its extreme tip, well beyond the posterior
transverse vein, while the membrane of the wings is not
rilled.

It is stated by Girschnert that the chetotaxy of the sterno-
pleure in his family “ Anthomyide” (which includes the groups
“ Coenosiine ” and “ Muscinee”) differs very much, and is even
variable in the species of the same genus, so that it is of only
subordinate value for systematic purposes. The author in
question goes on to say :—‘‘It is, however, characteristic of all
Anthomyids that, in the cases in which the sternopleura shows
only three macrochete, the order of arrangement is invariably
1:2, i.e. the larger number of bristles always occurs on the hind
border of the sternopleura.” As will be seen from Fig. 11, p. 69, this
is precisely what we find in the case of Glossina. In Girschner’s
family “ Tachinide,” however, when three sternopleural bristles
are present, the characteristic arrangement is always 2:1, or
the three bristles stand in a row, as in Sarcophaga and
Brachycoma.§

In conclusion, since Girschner has introduced an element of
confusion by employing old family names in new senses, it is
sufficient for present purposes to remember that, in terms of the
simplest taxonomy, the genus Glossina belongs to the sub-family

* Cp. Girschner, loc. cit. pp. 299-300.

+ Cf. Hough, ‘‘ Biological Bulletin,” Vol. I. (1899), p. 20, note 1:—
“These rills are very fine grooves in the surface of the wing which run in
a sort of radiate manner toward the border. They are very numerous. A
rilled wing denotes a higher stage of development, a more recent form,
than an unrilled wing.”

t Loc. cit. p. 302.

§ Girschner, Joc. cit. p. 303.

AFFINITY TO LYPEROSIA AND STOMOXYS. 59

Muscin&, of the family Muscip#, and that its nearest existing
allies are found in the genera Stomoays, Hematobia, Lyperosia
and Beccarimyia ; but that it is a highly specialised form,
exhibiting several unique structural features, in addition to a
very peculiar mode of reproduction. In some respects Glossina
approaches most nearly to Lyperosia, while as regards other
structural characters it comes nearer to Stomoxys. If, however,
we rely upon the character and mutual relations of the proboscis
and palpi, the Tsetse-flies are certainly more closely allied to
Lyperosia than to any other genus at present known.

60

CHAPTER “iy:

SYSTEMATIC DESCRIPTION OF THE GENUS GLOSSINA
AND ITS SPECIES, WITH NOTES ON THEIR
DISTRIBUTION AND HABITS.

Genus GLOSSINA.

Glossina, Wiedemann, Aussereuropiische zweifliigelige Insekten, Zweiter
Theil, pp. 253, 254 (1830). Nemorhina, Robineau-Desvoidy, Essai
sur les Myodaires (Mémoires présentés . . . a l’Académie Royale des
Sciences de l'Institut de France . . . Sciences Mathématiques et
Physiques. Tome Deuxiéme), pp. 389, 390 (1830).

Narrow-bodied, elongate, dark greyish-brown or yellowish-brown
dull-coloured flies, ranging in size from 74 millim.* (33 lin.) in
the case of a small specimen of Glossina morsitans, Westw., to
12 millim.* (52 lin.) in that of a large female of Gl. fusca,
Walk. ; recognisable when alive and at rest by the wings being
closed flat one over the other above the abdomen (beyond which they
project considerably), instead of divaricate (as in the case of
Stomoxys) or tectiform (as in Hematopotat), and by the proboscis
(i.e., proboscis ensheathed in the palpi), which in length is equal to

* Length measured from the face to the end of the abdomen, excluding
the proboscis and wings.

+ In the case of the genus Hematopota (family Tabanide), the species
of which (in England often known as ‘“‘clegs’’) are greedy blood-suckers,
and, though readily distinguishable by the prominent antenne, are not
unlike those of Glossina in shape, size, and general colouration, the wings,
when the insect is at rest, are somewhat tectiform (i.e. their anal angles
meet together like the roof of a house), as well as slightly divaricate.
Dr. Kertész’s Catalogue of Tabanidw (‘ Catalogus Tabanidarum Orbis
Terrarum Universi: conscripsit Dr. Colomannus Kertész. Budapestini,
1900”’) contains the names of some twenty species of Hematopota described
from various parts of Africa, and from specimens brought home by
collectors it is clear that Hematopota is sometimes mistaken for Glossina
when alive (Cf. Fig. 4, p. 6).

HEAD. 61

the thorax without the scutellum,* projecting horizontally 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 which in life they are applied so closely as entirely
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, though the front in the 9? is distinctly narrower
than in the 9 of Stomoxys; width of the front at the vertex in
the ¢ ranging from one-third to rather more than one-half of
that of the eye at its widest part, and in the ? from one-half to
rather more than 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
up-turned and prominent ; curved suture (Bogennaht 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 shallow
depression (transverse impression of Hough) separating the face
from the jowls ¢ ; the latter narrow in front, broad and rounded
posteriorly, crossed by a line of black priate: which passes on to
the occipital surface of the head, and is continued as the series
of small bristles marking off the posterior orbits ; jowls thickly
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 Stomoxys, Hematobia,
and other genera of Muscine, ciliated to the level of half the
length of the third joint of the antenne, or higher ; vibrisse
small, fine, no elongated or conspicuous vibrissa even on the
vibrissal angle, which is widely separated from the oral margin.
Underside of the roof of the buccal cavity with two large dark
brown strongly chitinised patches, very conspicuous when the
head is viewed from below in every species except GI. fusca and
Gil. longipennis. _ Eyes bare, in both sexes the facets in front

* The length of the visible portion of the palpi, i.e. the portion
projecting beyond the buccal cavity, in Glossina morsitans, Westw., is
just under 3 millim. (14 lin.) ; the length of the entire proboscis, measured
from the base of the bulb, is 34 millim. (12 lin.),

+ I follow Mr. G. H. Verrall («British Flies, Syrphide, etc.,’’ 1901) in
using the term jowl for the part of the head beneath the ey es, which in
former papers I have called the cheek (Backe, of German authors),
separated from the face (German, Wange) by the curved suture or its
prolongation. In 1898 Hough (Proc. Acad. Nat. Sc., Philadelphia,” 1998,
p. 168) proposed the terms gena and bucca for Wange and Backe respee-
tively,

62 NOMENCLATURE OF EXTERNAL STRUCTURE.

towards the inner margin conspicuously larger than those
behind. First two joints of the antennz small, the third long,
nearly reaching the oral margin, concave in front, the tip being

Soper (ensheathed in the palpi).

/ ~Ffroné /

| Ii] Fropé lag -
ent i

ate a

/

Cositi/
for

| Arista PY

.

ra

«
\

ao)

Py es Scutellum (part of mesothorar)

eee oe

i
:
eS
aS
iS
cS

ee

farsus |.
(5 jointed) \-—

ow oe ee ; eee 0 -—
‘
’
U
‘
i
‘
b)
1

=,

Fig. 8.
Diagram showing nomenclature of external characters of Glossina, sed in description.
VEINS AND CELLS IN THE WING.

Longitudinal Veins, Transverse Veins, Cells.

Te, Auxiliary vein. A. Anterior transverse vein. 1a, 1), le, First, second,

J. First longitudinal vein. B. Posterior 4 ne and third costal cells.
II. Second is 5 ©. Anterior basal ,, 8 2, Marginal cell.
Ill. Third y 7 D. Posterior ,,_ ,, AS 3. Sub-marginal cell.
IV. Fourth <i - 4, Discal cell.

V. Fifth P- a 5, 6, 7, First, second, and
VI. Sixth F i third posterior cells,

‘ 8. Anterior basal cell.
9. Posterior ,,_ ,,
10, Anal cell,

ANTENN i. 63

produced into a prominent forwardly-directed angle. On the
inside of the third joint of the antenna near the base a minute
though well-marked pore can usually be distinguished, constituting
the orifice of a sense-organ (see Fig. 9).* Arista distinctly two-

Fig. 9.

Left antenna of Glossina pallidipes, Austen, 6°, from the inner side (x 60) ;
p, aperture of sense-organ on third joint.

jointed, though basal joint very small; second joint remarkably
broad throughout its length, flat, tapering somewhat to the tip, but
far from being reduced distally to the proportions of a fine hair,
as in the case of the arista of Stomorys; feathered on the upper
side alone (including the tip) with some twenty-two fine, curving,

* Dr. Stuhlmann [174], writing of Gl. fusca, Walk., calls this an
“auditory” organ. If the antenna be mounted in glycerine and ex-
amined under a low power, the sense-organ has the appearance of an
oval sac.

64 PROBOSCIS, THORAX, AND ABDOMEN.

branched hairs (see Figure 9).* Proboscis projecting as already
described and, with the palpi, curving slightly downwards ; red-
dish-brown below, glistening yellowish-white above nearly to the
tip, which is marked off by a slight constriction and exhibits
beneath a tiny but conspicuous black spot ; the bulbous base
of the proboscis is enwrapped posteriorly with a fold of skin
forming the hind wall of the buccal cavity. Palpi rod-like,
of the same width throughout, neither swollen nor tapering
towards 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 and
forwards on the under and outer side immediately before the tip.
(For the bristles of the head see below under ‘“ Chetotaxy.—
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 macrochete likewise few in
number, arranged as described below under “Chetotaxy.”
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 very inconspicuous, and,
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, 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 they are longest and
stoutest ; hind margin of the sixth segment, 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 especially conspicuous
transverse row of black bristle-like hairs or fine bristles on each
side of the median line beyond the middle. The male hypopygium

* The number of these hairs is usually about twenty-two or twenty-
three, but sometimes there are not more than eighteen, while in other
cases there may be as many as twenty-eight; the numbers on each arista
may even be unequal in the same specimen.

+ In dried specimens these are often missing—one at least is frequently
indistinguishable.

LEGS AND WINGS. 65

highly characteristic in shape (see Figures 12 and 13, p. 94),
oval and tumid, its longer axis lying in the antero-posterior
direction, with a vulviform median groove (the anus) running from
the anterior margin to beyond the middle. Ventral plate of the
sixth segment in the male, immediately in front of the hypo-
pygium, bearing a patch of close-set erect minute black hairs
no each side of the median line: ventral plates of the remaining
segments except the first in the male, and of all the segments
except the first in the female, membranous.

Legs simple, rather long, claws longer and pulvilli somewhat
larger in the male than in the female. Front femora with a row
of bristle-like hairs above and below, and (except in the case of
Gl. morsitans *) on the posterior surface with a median row of
minute black spines. Middle and posterior femora with a row
of bristle-like hairs on the basal half of the anterior surface, near
the upper margin, and with a shorter row of similar hairs, or
short bristles as the case may be, at the base beneath. Middle
femora with a solitary bristle, usually conspicuous in the larger
species, on the upper side near the tip. Tibiz with a well-marked
ridge, edged with a row of closely-set minute black spines,
running down the outer surface, so that they, especially the
front and hind pair, usually have the appearance of being
somewhat flattened from side to side; the middle tibize have a
second row of still more minute spines in advance of that just
mentioned, the space between, owing to the presence of the
ridge, having the appearance of being excavated. Tibi also
with a row of short fine hairs or bristle-like hairs running down
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
tarsi three rows of minute black spines similar to those on the
tibize ; on the basal joints of the hind tarsi they are less distinct,
but apparently two rows are present. —

Wings with an absolutely unique venation (see Figure 8), which,
apart from all other characters, will at once serve to distinguish
a specimen of the genus. The most striking peculiarity is the
course of the fourth longitudinal vein. The anterior basal trans-
verse vein, at the base of the discal cell, is very short, and the
portion of the fourth longitudinal vein before the anterior (small)

* In the case of Gl. longipennis, too, the row appears to be broken up

into a double line and is consequently barely distinguishable.
. F

66 VENATION.

transverse vein is bent downwards in such a way as greatly to
narrow the basal half of the discal cell, the width of the distal
portion of the anterior basal cell being correspondingly increased ;
the fourth vein bends abruptly upwards to the point of contact
with the anterior transverse vein, where it forms with itself
what is practically a right angle; from this point it again runs
obliquely downwards, and is once more bent upwards at the
exact point of contact with the posterior transverse vein,
ultimately reaching the margin of the wing some distance before
the apex. The first posterior cell is open, but its distal por-
tion is narrow and much drawn out. The third longitudinal
vein is very close to the second, and shows a marked approxi-
mation to the costa, the sub-marginal cell being correspondingly
narrow; the third costal cell is considerably elongated; the
second, third, and fourth longitudinal veins all turn upwards at
the tips, and the anterior transverse vein is very oblique ; the sixth
longitudinal vein becomes obsolete soon after the anal cell,
though in clear, unchitinised form it can still be traced nearly
to the margin of the wing. Posterior basal transverse vein,
closing the anal cell, angulate in the middle, so that the distal
angles of the cell are acute. Posterior transverse vein slightly
curved, bent up somewhat abruptly to meet the fourth longi-
tudinal. Wings of an uniformly brownish hue, varying in
intensity in the different species, but apparently palest in
GI. morsitans ; membrane of the wing rilled.*

[Resemblance in venation between Glossina and Hypoderma.—
While there are many noticeable differences, the venation of
the wing of Glossina resembles that of the strid Hypoderma
(Warble-flies) more closely than that of any other genus, especially
as regards the shape of the first posterior cell. In both cases the
fourth vein is bent up to form’ what German authors call a
« Spitzenquerader” precisely at its junction with the posterior
transverse vein. But in the wing of Hypoderma the anterior
transverse vein is not oblique, and the fourth longitudinal vein
before reaching it is not bent down in the manner so characteristic

* Cf. Garry de N. Hough, M.D., ‘Some Muscine of North America,”
“ Biological Bulletin,” Vol. I. (1899), p. 20, note 1 :—‘' These rills are very
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
unrilled wing.” "

CHATOTAXY. 67

of Glossina ; besides this, the first and second longitudinal veins
pursue a different course, and the shape of the posterior basal
and of the anal cell is very different. The shape of the wing
itself is also different, and the wing is much smaller in proportion
in Hypoderma than in Glossina. ]

CHETOTAXY OF GLOSSINA.

Cephalic Bristles—One pair of vertical bristles of large size
(the largest and most conspicuous bristles on the head) ; on the
occipital region no trace of the pair of bristles termed by Hough
(Proc. Acad. Nat, Sc. Philad. 1898, p. 166) the occipito-central ;
greater ocellar bristles (ocellar pair of Osten Sacken) small ;

| Preesutural Bristle ,, tesa Huimeral Bristles
pei -----, Notopleural Bristles
Presuiural (of Girschner and pecan.
: 4 post-humeral of Osten
Oorso-central Bristie_______ . or a
2” dorso-central _-Supra-alar Bristle
Bristle. =i

se a f
a sa Cito Manip to o:-{-s>,---------- /ntra-étar Gristle

“~ Inner dorso-central
' Bristle

ee
pao on

Fig. 10. ;
Diagram of thoracic chetotaxy of Glossina, dorsal aspect. (xX 10.)

post-vertical pair not differentiated in size from the remainder of
the lesser ocellar bristles ; frontal bristles largest below (trans-
frontal group of Hough, loc. cit.); orbital bristles absent in
both sexes; vibrissal angle (separated by a well-marked and
rather wide interval from the oral margin) without a large and
conspicuous vibrissa ; vibrissal ridges ciliated to a point on a level
with or rather above the middle of the third joint of the antenna.
Thoracic bristles.—The general arrangement, as can be seen
from the accompanying diagrams, is, on each side, as follows :—
Dorsal aspect (Fig. 10).—Humeral, from 1 to 3 (sometimes
apparently 4), the lowest bristle the largest. Post-humeral
¥F 2

68 / CH ETOTAXY.

(intra-humeral of Osten Sacken) absent. Notopleural (of
Girschner and Hough, post-humeral of Osten Sacken), 2. Pre-
sutural, 1.* Supra-alar, 1. Intra-alar, 1, situated just in front
of the post-alar callus, on a level with the second dorso-central.f
Post-alar, 3, the foremost somewhat smaller than the other two,
and placed directly above 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 presutural dorso-central.§ Inner dorso-ceniral, || 1,
near the hind margin of the thorax. Scutellar, 2, one marginal
(near the basal angle), the other apical ; in certain specimens of
Glossina fusca, Walk., and Gil. longipennis, Corti, there is a
second marginal bristle in front of the ordinary one, and also, in

* Often small and difficult to distinguish in Gl. palpalis.

+ I follow Hough (loc. cit, p. 180) in numbering the dorso-centrals
from rear to front, except that, for reasons explained in the note below, I
term the comparatively large dorso-central bristle immediately in front of
the transverse suture the preswtwral dorso-central.

t Cf. Osten Sacken, Trans. Ent. Soc. Lond. 1884, p. 504,

§ In certain cases (e.g. individual specimens of Gl. morsitans and
Gi. longipalpis) there are one or more small additional bristles (little larger
than the small bristle-like hairs clothing the general surface of the
dorsum, but nevertheless recognisable as bristles belonging to the dorso-
central series) between the second dorso-central and the suture. It is
therefore obviously impossible to give the dorso-central bristle in front of
the suture a numerical designation.

N.B.—In Fig. 10 the extra dorso-central bristles, as well as certain
others which sometimes occur, are not shown, the object being to represent
so far as possible the normal chetotactic arrangement in the genus.
Aberrations in cheetotaxy noticeable in the material available for examina-
tion will be referred to under the individual species.

| Cf. Mik, ‘‘ Ueber Acrostichalbérstchen (setulx acrostichales)” : Wien.
Ent. Z. xix. Jahrg. 1900, pp. 151-152.—As pointed out by Mik (loc. cit. p. 152),
receut writers on Muscidve, such as Girschner and Stein (as also Hough),
following an original slip on the part of Osten Sacken (cf. Trans. Ent.
Sos. Lond. 1884, p. 509), have misapplied the term acrostichal in using it
for the inner dorso-central series of bristles. The small acrostichal
bristles, however, cannot be interpreted as inner dorso-central bristles.
Mik adds: ‘“ The dorso-central bristles are always symmetrically
(bilaterally) arranged macrochetw, whereas the little acrostichal bristles
(Acrostichallérstchen) belong to the ordinary original covering of the
dorsum of the thorax and occur by no means seldom in a single row along
the sagittal line, or may form even more than two longitudinal rows. It
may also happen that little acrostichal bristles occur even when the two
rows of the inner dorso-central bristles are present. _ Whoever, for instance,
examines a Dolichopodid provided with little acrostichal bristles will at
once recognise that these little bristles have nothing to do with the
macrochete, but that they belong to the same category of covering
(Bekleidung) as the tiny bristles that occur on the anterior portion of the
mesothorax, especially behind the humeral calli.”

Iam indebted to Mr. J. E, Collin for directing my attention to Mik’s
note, which otherwise I should have overlooked, as it is concealed among
the last instalment of the lamented author’s “‘ Dipterologische Miscellen.”

ADDITIONAL BRISTLES. 69

some specimens of Gl. fusca, an extra bristle between the
ordinary marginal and the apical.

Out of the material examined certain specimens, belonging to
GI. morsitans, Westw., Gl. pallidipes, Austen, and Gil. palpalis,
R.-Desv., show an additional small bristle in advance and a little
to the outside of the intra-alar: whether this bristle should be
regarded as belonging to the intra-alar or to the supra-alar series
T am unable to determine. In Gi. longipennis, Gl. fusca, and
(less clearly) in Gl. pallidipes an incomplete circlet of short
stoutish bristles (not shown in the figure) can be seen on the
anterior margin of the dorsum of the thorax between the

Presutural dorso-central Bristle. — Supra-alar Bristle 2*dorsocentrai Bristle

Preesutural Bristle. 5 \ Intra-alar Bristle’ / i 7“dorso-central
ass : , — ae Bristle

Aeuimeral
Bristles a
Notopleural _ Scutellar Gristles
&ristles . hs ¢ A, . : HR. (marginal and apical.)
; ra eG evap llieiie *s %
sige tl re ot (2 / fh s Tbe Rac ‘.. Aost-alar
9 eee Sable ae se Sristles
Prothoracie “1-7 ° a or Pe ,
Bristle <1" OF a a Ee ie a ah Base of wing
: sae = \ “ > rey, = Le z= att
Stigmatic Bristle i i Fi 4 eS cenee
(Becker. Ber!.Ent.Z, The ge ee a SHU
XXHIK (1894), 0.83.) fay oe Hage PS
5 ae fi 7 ; A ;
Mesopleural Sristles “ Ps fy Posterior Sternopleural! Bristles

5 ie
,

“ee
Anterior Sternopleural Bristle /Pteropleural Bristles

Fig. 11.
Diagram of thoracic chetotaxy of Glossina, pleural aspect (x 10).

N.B. There are no bristles on the hypopleura (the region above the last two pairs of
cox) in Glossina, though Osten Sacken writes (Trans. Ent. Soc, Lond., 1884, p. 513) :—
“‘ Most of the Calyptrata, except the Anthomyide, have a tuft or row of bristles on the
hypopleura, a region which is destitute of them in the other families of Diptera.’”’—
See also Girschner, ‘‘ Beitrag zur Systematik der Musciden:” Berliner Entomolo-
gische Zeitschrift, Bd. xxxviii. (1893), pp. 297-298. According to Girschner’s arrange-
ment, Glossina belongs to the group Muscine of his ‘‘ Family Anthomyide.”

humeral callosities. These bristles are frequently difficult to
distinguish owing to the shadow between the head and the
thorax. There are usually three bristles on each side, close to
the humeral callosity, while a bristle on each side of the median
line would appear to belong to the inner dorso-central series.

The relative size of the various bristles enumerated above is
roughly indicated in the diagrams by the size of the dots.
Generally speaking the largest bristles are the lowest humeral,
the notopleural, supra-alar, post-alar, the bristles of the transverse

70 PLEURALCHATOTAXY. SYNOPSIS OF SPECIES.

row formed by the first and inner dorso-centrals, and the marginal
and apical scutellar bristles. /

Pleural aspect (Fig. 11).—Prothoracic, 1. Stigmatic (ef. Becker,
Berl. Ent. Z. xxxix (1894), p. 83), 1. Mesopleural, a vertical row of
about 6 along the posterior edge, with smaller bristles in between.
Pteropleural,* a vertical row of usually 3 (stout and conspicuous
only in Gl. longipennis, Corti, and Gil. fusca, Walk., more
numerous in the latter than in the former), with smaller bristles
on either side. Sternopleural, 3,—1 anterior, and 2 posterior :
below, a conspicuous row of bristles in front of the middle coxe.
Hypopleural, none.

SYNOPSIS OF SPECIES.

1. Hind tarsi entirely dark ......... Tyce cnechenssacas 7 2
Hind tarsi not entirely dark: last two joints
alone dark, remainder pale .............scossese 3
2. Third joint of antennae dusky-brown to
CIMETEGUA=DIACK 4, ...\-carTesssOusededeccowes dered ot palpalis, Rob.-Desv.

Third joint of antennae pale (orange-buff) ... pallicera, Bigot.
3. Large species: length at least 103 millim.
(5 lin.), wing-expanse (measured from tip
to tip, when wings are set at right angles
to body) at least 25 millim, (112 lin.) ...... 6
Smaller species: length not exceeding 104
millim. (5 lin.), often considerably less ;
wing-expanse not exceeding 224 millim.)

(MOD. siciin ccs cdnieamansides wale aveviieintaacheetieea ae 4
4, Last two joints of front and middle tarsi with
sharply defined dark brown or black tips... 5

Last two joints of front and middle tarsi with-
out sharply defined dark brown or black
tips—front and middle tarsientirely yellow,
or last two joints of former faintly tipped
WAH palo PEO WII! sassccscteaces taeiecsoetee sss «55. pallidipes, sp. nov.
5. Generally distinctly larger; head wider; front
darker and narrower in both sexes, sides
parallel in g ; abdominal bands deeper,
leaving hind margins of segments only
narrowly pale; hypopygium in ¢ smaller,
darker, and more hairy; tip of ¢ abdomen
more thickly clothed laterally with short
black hair, bristles on 6th segment finer
and Tess promMINent \eersacescrcess-odsissscseaweos longipalpis, Wied.

* Pteropleural bristles seem to be entirely wanting in Stomozys,
“Hematobia, and other genera of Muscine.

SYNOPSIS OF SPECIES. 71

Usually smaller; head narrower ; front paler
and wider; eyes in ¢ as well as in ?
distinctly converging towards vertex ; ab-
dominal bands less deep, pale hind margins
of segments therefore deeper ; hypopygium
in ¢ larger, paler, somewhat more oval in
outline, and clothed with fewer fine hairs ;
tip of g abdomen less hairy laterally ;
bristles on 6th segment in ¢ stouter and
MOLE CONSPICUOUS —.... es sensecsescncrencseeoes morsitans, Westw.
6. Dorsum of thorax with four sharply defined
small dark brown oval spots, arranged in a
parallelogram, two in front of and two
behind transverse suture; bulb at base of
proboscis brown at the tip ...........scsceeeees longipennis, Corti.
Dorsum of thorax without such spots, though
with more or less distinct longitudinal
stripes; bulb at base of proboscis not brown
Siti HG il aoe fe Meared ctiainsiaitguiat’s piehidcin aisha <as cosines’ fusca, Walk.

Glossina palpalis, Rob.-Desv.
(Plate I.)

Nemorhina palpalis, Robineau-Desvoidy, Essai sur les Myodaires
(Mémoires Présentés par Divers Savans a l’Académie Royale des
Sciences de l'Institut de France. Sciences Mathématiques et
Physiques. Tome Deuxiéme), p. 390 (1830).

Stomoxys longipalpis ?, Walker (nec Wiedemann), List Dipt. Ins. in Coll.
Brit. Mus., Pt. III., p. 682 (1849).

Glossina tachinoides, Westwood, Proc. Zool. Soc. Lond., Pt. XVIIL.,
p. 267, Pl. XIX., fig. 2 (1850): Ann. Mag. Nat, Hist. Ser. 2 Vol. X.,
p. 147 (1852).

Glossina longipalpis, Walker (nec Wiedemann), Entomologist, VI., p. 328
(1873).

Glossina ventricosa, Bigot, Ann. Soc. Ent. France, 6° Série, T. 5,
pp. 122, 123 (1885).

Glossina longipalpis, Bigot (nec Wiedemann), ibid, p. 122.'

Glossina tabanifornis, Bigot (nec Westwood), ibid. p. 123.

Glossina longipalpis, Austen (nec Wiedemann), Report of the Proceedings
of the Expedition for the Study of the Causes of Malaria, etc., p. 18
(1899).

3, ?.—Length* 8 to 94 millim. (33 to 44 lin.) ; length of wing
8 to 94 millim. (32 to 4} lin.) ; width of head 2} to 2} millim.
(14 to 12 lin.).

* The length given is in all cases exclusive of the proboscis and palpy,
and is measured trom the epistoma to the tip of the abdomen.

72 DESCRIPTION OF GLOSSINA PALPALIS.

Dark brown ; thorax usually paler, with dark brown markings on
a grayish ground ; abdomen generally with at least an indication of
a pale longitudinal median stripe, with pale lateral triangular
markings, and usually the hind margins of the segments narrowly
pale. Legs (except the hind tarsi and last two joints of the front and
middle pairs) sometimes entirely buff-coloured (var. tachinoides,
Westw.) ; usually the femora for the most part or entirely dark brown,
in well preserved specimens clothed with grayish dust, and the tibize
yellowish.

Head :—face yellowish, clothed below with grayish dust ;
posterior surface of head entirely cinereous ; frontal stripe varying
from ochraceous to dark chestnut ; frontal margins grayish, seen
from the side with a dark brown elongated area below ; ocellar
triangle cinereous, enclosing the dark-brown ocellar spot, which
is joined posteriorly to a sharply defined dark-brown band, uniting
the vertical bristles and very conspicuous except in the darkest
specimens. Antennze: 2nd joint more or less yellow at the apex
in front, 5rd joint narrowly yellow at the extreme base on the
outer side, otherwise entirely dark, with grayish shimmer ; arista
yellowish, dark brown on the under side. Palpi cinereous, blackish
on upper side. Bulb at base of proboscis dark brown. Thorax
in the most clearly marked specimens bluish-gray or cinereous,
with brown markings as shown in Plate I. These markings when
fully visible are as follows : a narrow stripe on each side of the
median line, interrupted before 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 immediately in front of the hind margin
takes the shape of a pair of more or less confluent ill-defined spots,
sometimes confluent with the stripes in front; next to the two
admedian 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, but in front curving round outwards behind the humeral
callus and then running backwards along the lateral margin of
the dorsum nearly to the post-alar callus; in the area thus
enclosed a broad ill-defined patch in front of and behind the
suture, while the lateral stripe itself sends off two prolongations,
which run inwards for a certain distance on each side of the
suture. Humeral callus with a spot on its upper portion,
confluent with the curved stripe behind it; a more or less ill-

GLOSSINA PALPALIS: ABDOMEN AND LEGS. 73

defined spot on the post-alar callus also. Pleurz cinereous, a
more or less distinct brown patch in the centre of the mesopleura.
Scutellum cinereous at the base, yellowish towards the tip and
along the margin, the usual brown patch on each side of the
median line more or less conspicuous.

In dark specimens the markings on the dorsum of the thorax
become confluent to such an extent that the dorsum appears
almost wholly brown, the bluish-gray or cinereous ground colour
being scarcely visible except in front on either side of the
confluent ad-median stripes.

Abdomen dark brown ; first segment and a median triangular
area on the second (its base resting on the front and its apex on
the hind margin of the second segment) buff-coloured or cinereous,
the pale triangle continued backwards as a narrow, more or less
well-defined median stripe, usually reaching at least as far as the
hind margin of the fifth segment ; lateral margins of the segments
from the second onwards cinereous or buff-coloured, expanded on
the apical angles into triangular markings; extreme hind
margins of the segments from the second to the sixth usually
narrowly pale or cinereous ; seventh segment, as also the hypo-
pygium in the ¢, entirely cinereous.

In a very dark ¢ from Asaba, R. Niger (Dr. W. H. Crosse),
the pale triangle on the second segment is but faintly indicated,
and the remainder of the upper surface (with the exception of
the cinereous lateral triangles which, since the lateral margins
are turned down, cannot be seen from above) is wholly dark
brown, the pale median stripe and hind margins to the segments
being wanting.

Legs.—Coxe yellowish cinereous ; femora cinereous to cine-
reous-brown (the middle and hind femora on the outside towards
the tips sometimes darker than elsewhere), the knees, extreme
base of front femora, basal half of the middle femora on the
inner side, more or less of the basal half of the hind femora on
both inner and outer sides, and usually an ill-defined area at the
tips of the hind femora also, yellow ; trochanters likewise yellow ;
front and middle tibie, and first three joints of front and middle
tarsi yellow; hind tibie yellow to yellowish-brown ; hind tarsi
entirely black above ; last two joints of front and middle tarsi
blackish-brown, except their bases, which are more or less
distinctly yellow.

The proportion of yellow on the femora is variable, and they
are sometimes more yellow than cinereous; on the other hand,

74 GLOSSINA PALPALIS, VAR. TACHINOIDES.

the very dark ¢ from Asaba, already alluded to, has the femora,
with the exception of the knees at the extreme tips and the
extreme base of the hind pair on the inside, entirely dark
cinereous-brown.

Wings uniformly brownish. Squame white, border of the
antisquama darker, fringed with short darker hairs. Halteres
yellowish white.

Var. tachinoides, Westw.—Legs, except hind tarsi and last two
joints of front and middle pairs, wholly pale yellow ; median stripe
and the other pale markings on the abdomen very conspicuous.

Bulb at base of proboscis ochraceous instead of dark brown,
but with a dark brown patch on each side towards the tip.
Pleurz and humeral calli more or less ochraceous-buff ; the same
colour may also appear on the dorsum of the thorax between the
curved stripes and the enclosed dark patches.

Abdomen.—In two ? 2 from the Gambia (Dr. Dutton) the
median stripe is unusually broad and well-marked, and extends
outwards along the front and hind margin of each segment from
the third to the sixth ; the lateral triangles are also exceptionally
conspicuous, more clearly buff-coloured than is usually the case in
the typical form, and are prolonged conspicuously on to the upper
side along the hind margins of the segments, the pale hind margins
themselves being broader than in the typical form and well-marked.*

The var. tachinoides, therefore, clearly shows the connection
between the typical Gl. palpalis, with uniformly dark-brown
abdomen, only relieved by a pale median stripe and more or less
conspicuous lateral triangles, and Gl. morsitans or Gl. longipalpis ;
for the abdomen of var. tachinoides, at any rate as represented by
the two 2 9 referred to, might well be described as cinereous-
buff, with dark brown interrupted transverse bands.

Legs, with the exception of the hind tarsi and last two joints
of the front and middle tarsi, entirely pale yellow, though there
is sometimes a small dark patch on the outside of the middle and
hind femora towards the tip.

* In addition to the two ¢ ? from the Gambia, the Museum collection
includes four other specimens (two ¢ ¢ and two ¢? @) which must be
regarded as belonging to the var. tachinoides. According to their labels,
they are from the “ Zambesi” (Dr. Kirk). All have the legs pale, with
the exception of the hind tarsi, which are wholly dark, and the last two
joints of the front and middle pairs, which are partially infuscated, but

the median abdominal stripe and the pale hind margins to the abdominal
segments are not especially conspicuous except in the case of one female.

DISTRIBUTION OF GLOSSINA PALPALIS. 75

Distripution oF Gil. palpalis, Rob.-Desv.

The present species, which is the Glossina longipalpis of
authors (nec Wiedemann), has a wide range in West Africa,
and, as is shown by the localities of the specimens before
me, in its distribution from north to south extends at least
from the Gambia (var. tachinoides) to the Congo, though from
our knowledge of the extremely local occurrence of Glossina
morsitans in South Africa, as indicated by the familiar term “ fly-
belt,” it is not at all likely that GJ. palpalis will be found to
occur everywhere on the West Coast. As to the range of the
species inland all that can be stated at present is that var.
tachinoides was found by Dr. (afterwards Sir) John Kirk on the
Zambesi—the precise locality is unfortunately unknown. Gl.
palpalis, however, is not the only species found in West Africa,
since, as will be shown later, Gl. pallicera, Big., Gl. fusca, Walk.,
Gl. longipalpis, Wied., and Gil. morsitans, Westw., also occur
there.

The above description has been drawn up from an examination
of forty-nine (29 ¢ g, 20 2 2) specimens from various localities,
as indicated by the following list * :—

4 $ $, mouth of Kissy River, near Free Town, Sierra Leone,
16. viii. 1899, ‘among mangroves, on boats” (EH. EH. Austen); 2g 2,
Wilberforce, near Free Town, Sierra Leone, 10. ix. 1899, “ settling
on stones in hillside stream at garrison watering-place” (H. E.
Austen); 1g, “Gold Coast” (A. Swanzy); 1 9, “Gold Coast ”
(Capts. Burton and Cameron) ;5 g 6,3 9 9, Assinie, Ivory Coast
(Ch. Alluaud) [V.]; 5 66,6 2 2, Togo (Dr. Schilling), [Royal
Zoological Museum, Berlin]; 1 ¢, Benin (A. Millson); 1 9,
Degema, Nigeria, 8. viii. 1900 (Dr. A. H. Hanley); 1 g,1 9,
Asaba, R. Niger (Dr. W. H. Crosse); 1 g, Fernand Vaz River,
French Congo (Du Chaillu) ; 3 g g, Congo (presented by Miss
EK. M. Bowdler Sharpe)’; 1 ? , Congo (the specimen referred to by
Walker, List Dipt. loc. cit., as Stomoxys longipalpis) ; 1 g , Congo
[O.—Ex Coll. Saunders]; 2 ¢ ¢, 1 9, locality unknown [V.];

* Throughout the present chapter, specimens not in the collection of the
British Museum are distinguished as follows: those belonging to the Oxford
Museum by the letter [O.]; those in the collection of Mr. G. H. Verrall
(formerly in that of the late M. Bigot) by the letter [V.]; and those in the
collection of Mr. L. R. Crawshay by the letter [C.] Wherever possible the
name of the Collector is given in italics, thus (f. J. Jackson).

+ Glossina palpalis was described from a specimen from the Congo;
Robineau-Desvoidy writes (loc. cit.): ‘‘ Cette curieuse espéce, originaire du
Congo, fait partie de Ja collection du comté Dejean.”

76 HABITS OF GLOSSINA PALPALIS.

2 99, locality unknown [V.—The types of Gl. ventricosa,
Big. }.

Var. tachinoides, Westw.—l ¢, ‘West Africa” [O.—The
type of GI. tachinoides, Westw.]; 2 9 9, R. Gambia, between
Nov. 1901 and Jan. 1902 (Dr. J. HE. Dutton); 2 6 6,2 2 9,
“ Zambesi” (Dr. (afterwards Sir) John Kirk).

Hasits, ETC.

Glossina palpalis was quite common in the vicinity of Free
Town, Sierra Leone, during the months of August and September,
1899. It always seems to occur along the beds of streams, where
it is fond of sitting on stones projecting from the water ; it also
abounds in mangrove thickets fringing the mouths of streams,
close to the sea-shore. On one occasion I found it among bushes
on the hillside below Wilberforce, settling on stones in a small
stream barely a yard in width. The fly was also met with on
the way to Regent, where the road crosses a little brook, and it
probably occurs along all the streams in the neighbourhood of
Free Town. It bites and sucks human blood freely, and the
writer and two companions were once severely bitten by it while
resting undressed on the sea-shore after bathing, close to a
mangrove thicket at the mouth of a stream. The fly is remark-
ably active and exceedingly difficult to catch, but persistently
returns to the same spot. English-speaking natives, when
questioned, call it a ‘“ mangrove-fly,"—a name that, in West
Africa, is also indiscriminately applied to various species of the
blood-sucking Tabanide (Horse-flies).

As to the var. tachinoides, Dr. J. Everett Dutton, of the
Liverpool School of Tropical Medicine, in a letter to Mr. F.
V. Theobald, dated “Liverpool, Feb. 21st, 1902,” writes as
follows: “The small mangrove-fly is very prevalent up the river
Gambia, where it comes on board the launches and bites
viciously.—Now the case of Trypanosoma I found in Bathurst
was in an Englishman who was master of the Government
launch, living on board, and frequently bitten by this fly—It
is also interesting to note that the Colonial Surgeon informs me
that the cases amongst the natives which he has seen, with
similar symptoms to those in the white man, occurred in boatmen
plying up and down the river.”

SYNONYMY AND AFFINITIES. 17

SyNONYMY AND AFFINITIES.

Although Robineau-Desvoidy’s description of Nemorhina
palpalis is very brief, it is nevertheless recognisable ; and though
in this case I have not been able to examine the type, which
there is only too much reason to fear is no longer in existence,*
there need, I think, be no hesitation in applying the name
palpalis to the present species. On the other hand, subsequent
writers, relying on Wiedemann’s description, and without having
seen his type, may well be excused for considering it to be
Wiedemann’s longipalpis, especially since the type of the latter
was stated to be from Sierra Leone. Judging merely from the
description,t Glossina longipalpis, Wied., would appear to have
been described from a specimen of the present species with pale
legs, and well-marked pale median stripe and hind margins to
the abdominal segments (i.e. var. tachinoides). As will be shown
below, however, an examination of Wiedemann’s type, which by
the courtesy of the authorities of the Vienna Museum I have
been enabled to make, proves conclusively that Glossina longi-
palpis, Wied., is a distinct species nearly allied to Gl. morsitans,
Westw. .

The type of Glossina tachinoides, Westw., is a mere fragment,
but fortunately sufficient remains to establish its identity, and to
show that it cannot be regarded as anything more than a variety
of palpalis.

* As already stated in the previous note, according to the author
Nemorhina palpalis was described from a specimen in the collection of
Comte Dejean. In the hope of obtaining some information as to the type,
and perhaps eventually the loan of it, I applied to the Muséum d’Histoire
Naturelle, Paris, and received the following courteous reply (dated
“17 avril 1902”) from M. Joanny Martin, Assistant in charge of the
collections of Diptera and Hemiptera in the Muséum :—‘‘J’ai le regret
de vous apprendre que nous ne possédons malheureusement pas la collec-
tion de Robineau-Desvoidy. Cet auteur avait légué ses collections au
Musée d’Auxerre (Yonne). Depuis plusieurs années, il ne reste plus rien,
absolument rien. Cependant comme ce Muscide appartenait & la collection
Dejean il est bien possible que le lot de Diptéres fut acheté par Bigot, qui
a cette époque déja étudiait les Diptéres. Dans ce cas vous trouveriez le
type qui vous préoccupe chez Mr. Verrall qui a acheté la collection Bigot.
C’est la derniére chance qu'il vous reste pour retrouver ce précieux Diptére.”
Unfortunately there is no trace of Robineau-Desvoidy’s type among the
specimens from the Bigot Collection which Mr. Verrall has kindly lent
me—unless indeed it is one of those without a label alluded to above, in
which case there is now no means of identifying it.

+ Wiedemann’s figure (Auss. Zw. Ins. II., Pl. IX., fig. 10a) is worthless,
but at the same time supports an erroneous conclusion by conveying the
impression of a species with very dark abdomen and well-marked pale hind
margins to the abdominal segments.

78 GL. VENTRICOSA = GL. PALPALIS.

Glossina ventricosa, Bigot, proves on examination of the typical
specimens, which, as already stated, are both females, to be
nothing but Gl. palpalis with the abdomen enormously distended
with coagulated blood. One of Sir John Kirk’s specimens of
Gl. palpalis from the Zambesi has the abdomen similarly distended,
though to a smaller extent, and one of Dr. Dutton’s examples of
var. tachinoides from the Gambia also has the abdomen full of
blood ; a male Glossina longipennis, Corti, from Somaliland,
belonging to the Museum collection, likewise has the abdomen
swollen and distorted owing to contained blood. According to
Bigot’s statement (loc. cit. p. 121), he discovered the specimens
described by him as Gl. ventricosa among a number of Diptera
from Australia, which he had acquired afew years before the date
of his paper (1885). In consequence of this Bigot gave the
locality of his specimens as “ Australie?” (loc. cit. p. 123), having
previously (p. 121) stated that it was improbable that there had
been any confusion as to their origin, and having drawn attention
to the interest attaching thereto, since no representative of the
genus had been met with outside Africa. Nevertheless there can
be no doubt that confusion had taken place here. As is well
known, the late M. Bigot was in the habit of removing the original
labels from the specimens he acquired, and copying the data as to
the locality on to the label bearing the specific name, which he
pinned into the box containing them. Thus Bigot may well
have removed the labels from these specimens, and then have
put them down inadvertently among the Australian Diptera and
forgotten them. Be this as it may, Glossina ventricosa, Bigot, is
absolutely identical with Gl. palpalis, Rob.-Desv., and there is
no evidence whatever that Glossina occurs outside Africa.*

* It is true that it had been previously suggested by Van der Wulp
(Tijdschrift voor Entomologie, XX VII. (1884), p. 150), that the ‘‘ Mouche
charbonneuse”’ of New Caledonia was ‘“ probably a species of Glossina,”
but the suggestion is superfluous, since, given the presence of anthrax
among cattle in the vicinity and an abraded surface of human skin, any
carrion-haunting Muscid might quite possibly, as is the opinion of Macleay,
produce the results (deaths from malignant pustule) supposed to be due to
the bite of a particular fly in New Caledonia. See ‘‘ Note on a Reputed
Poisonous Fly of New Caledonia”, by William Macleay, F.L.S., etc.,
Proc. Linn. Soc., New South Wales, Vol. VII. (1883), pp. 202-205. A fly
caught in the act of biting by Mr. E. L. Layard, C.M.G., H.B.M. Consul,
New Caledonia, and at first supposed to be the dreaded ‘‘ Mouche char-
bonneuse,’’ was determined by Macleay to be ‘‘a Stomoxys, an insect not
uncommon in this country [Australia], and very probably intreduced, as
Mr. Layard suggests, into New Caledonia and the Isle of Pines from
Australia, as the maggot of the fly lives in horse-dung”’ (cf. Macleay,
loc. cit. pp. 204-205). i

DESCRIPTION OF GLOSSINA PALLICERA. 79

Glossina palpalis is the darkest of all the species of Glossina,
and cannot well be confused with any other, with the exception
of Gl. pallicera, Bigot (q.v. for distinctive characters).

As compared with Glossina morsitans, Westw., the inner
margins of the eyes in both sexes are more parallel, and the eyes
are usually wider apart at the vertex. Moreover, from this
species, as from GI. longipalpis, Wied., it is distinguished at once
by the entirely black hind tarsi.

Glossina pallicera, Bigot.
(Plate II.)

Glossina pallicera, Bigot, Annales de la Société Entomologique de France,
Année 1891, Vol. LX., p. 378.

g, 9. Length, 8 millim. (32 lin.); length of wing, 8 millim.

Brown; thorax mouse-gray* with brown, sometimes confluent
markings ; antenne orange-buff,t ; abdomen with no conspicuous
markings, except a narrow pale median triangle on the second
segment, and yellowish-cinerous triangular lateral markings (scarcely
visible from above) on the segments from the third to the siath; legs
pale ochraceous ; hind tarsi, and tips of last two joints of front and
middle pairs, dark brown ; wings brown.

This species can only be confused with Gl. palpalis, R.-Desv.,
from which it is at once distinguished by the colour of the
antenne.

Head.—Frontal stripe ochraceous ; frontal margins yellowish-
cinereous ; face buff, with a whitish shimmer ; palpi ochraceous-
buff, infuscated at the tips; bulb at base of proboscis pale
ochraceous ; arista buff-coloured, dark brown on the under side
at the base.

Thorax.—The pattern of the dark-brown markings is the
same as in Gl. palpalis ; in the single 2 before me the markings
are more or less confluent, so that the dorsum of the thorax
appears of a mottled brown colour; the impressed suture and
remains of the cinereous stripe on each side of the confluent

* Ridgway, “Nomenclature of Colors” (1886), Pl. IT.
+ Lbid.sPl, Vis ,

80 GLOSSINA PALLICERA.—AFFINITIES.

ad-median brown stripes in front, paler; humeral calli and
pleure cinereous; scutellum cinereous at the base, ochraceous
buff on the distal half, with the usual dark-brown patches, one
on each side of the longitudinal median impression, separated by
a rather wide interval in the typical specimens.

Abdomen.— With the exceptions mentioned above, unicolorous ;
the typical g¢ has the hind margins of the second, third and
fourth segments, and a median stripe on the two latter darker
than the remainder of the upper side of these segments, so that
the third and fourth segments appear to be marked with an
interrupted transverse band, somewhat paler than the ground
colour; this, however, is probably an effect of drying ; seventh
segment and hypopygium in the g, and seventh segment, at least
at the sides, in the 9 clothed with yellowish cinereous dust.

Squamez yellowish white; fringe of antisquama brown ;
halteres yellowish.

Distrisution oF Gl. pallicera, Bigot.

Nothing is known as to the geographical range of this species.
The above description has been drawn up from the examination
of Bigot’s actual type (which isa ¢ and nota 2, as stated by
him) and a second specimen, a 9, which had been placed by
Bigot above the label “ @. longipalpis” (= Gil. palpalis, Rob.-Desv.),
but certainly belongs to the present species. Both of these
specimens, the only ones that I have seen, are from Assinie,

Ivory Coast (Ch. Alluaud) [V.]

Hasits, ETC.
Unknown.

Synonymy AND AFFINITIES.

Although it was suggested by Bigot (loc. cit.) that Gl. pallicera
is perhaps merely a variety of ‘“ G. longipalpis” (= Gl. palpalis,
Rob.-Desv.), it appears to me to be quite distinct and fully
entitled to specific rank. I express this opinion with the greater
confidence, since I have been enabled to examine a specimen of
each sex. Gil. palpalis is the only species with which Gl. pallicera
can be confused, but, although closely allied, it can readily be dis-
tinguished by the characters given above. In addition to these,
the front in both sexes of Gl. pallicera is conspicuously narrower
than in Gl. palpalis. In the ¢ the arista of Gl. pallicera is

DESCRIPTION OF GLOSSINA MORSITANS. 81

stouter and longer, but in the 9 the citennes if any, is less
noticeable. ie

Except for the more or less distinct pale triangle on the
second segment, there is no trace of a pale median hacgindian!
stripe on the abdomen of Gi. pallicera, and the narrow pale hind
margins to the abdominal segments, usually distinguishable in
the case of Gl. palpalis, are likewise absent. Thus the abdomen
of Gl. pallicera is unicolorous, though the presence of the narrow
lateral cinereous triangles on the segments from the third to
the sixth shows the connection with Gl. palpalis, in which the
banded type is clearly traceable, though far less distinct than in
Gl. morsitans, longipalpis, or pallidipes.

Glossina morsitans, Westw.
(Plate III.)

Glossina morsitans, Westwood, Proc. Zool. Soc. Lond., Pt. XVIII,
p. 261, Pl. IX., figs. 1, 1*-1' (1850); Kirk, “On the ‘Tsetse’ Fly of
Tropical Africa”? (Glossina morsitans, Westwood), Journ. Linn. Soe.,
Zoology, Vol. VIII., pp. 149-156 (1865); Westwood, Oates’ ‘‘ Matabele
Land and the Victoria Falls,” Appendix, pp. 363-364, Pls. G, fig. 2,
and H, figs. 5, 5*, 5° (1881): Bigot, Ann. Soc, Ent. France, 6° Série,
JT. 5, p. 123 (1885): V. von Réder, Jahrb. der Hamb. Wissensch.
Anstalten, X. 2, p. 205 (1893).*

Glossina longipalpis, Macquart (nec Wiedemann), Diptéres Exotiques,
Suppl. IV., p. 239, Tab, 22, fig. 4 (1850); ? Schiner, Reise der Oster-
reichischen Fregatte Novara, Zoologischer Theil. Bd. II., Diptera,
p. 311 (1866).

6, 9%. Length, 73 to 93 millim. (32 to 4} lin.) ; length of wing,
71 to 9 millim. (3) to 4 lin.); width of head, 2! to 25 millim.
(iy to 1} lin.); width of front at vertex in ¢ 1, in 2 1 of total
width of head measured across middle of eyes,

~~ “Thorax mouse-grey}, often somewhat paler in front, with more or
less distinct brownish longitudinal markings ; abdomen varying from
drab-greyt to ochraceous-buff,t the segments from the third to the
sixth inclusive with very, conspicuous dark-brown bands, interrupted

* Record of the finding of the species by Dr. Fr, Stuhlmann at. Mbu-
sini, in Usegua, German East Africa, 27. viii. 1888.

} Ridgway: ‘“ Nomenclature of bin tall Side BA

t Ridgway, op. cit. Pl. V. tad it3

82 DESCRIPTION OF GLOSSINA. MORSITANS.

in the median line, not reaching the lateral margins, and not ex-
tending beyond the basal three-fourths of each segment, if so far ;
tips of the last two joints of the front and middle tarsi dark brown
or black, sharply contrasting with the rest of the leg.

Head.—Pale yellow, occipital surface entirely clothed with
cinereous dust ; frontal stripe varying from buff to ochraceous,
face paler, clothed at the sides and below with yellowish silvery
dust ; sides of front and ocellar triangle also clothed with yellowish
or greyish-yellow dust ; a more or less distinct dark brown spot
on each side of the front below, and the upper portion of the
frontal stripe also sometimes infuscated; base of each vertical
bristle marked with a,dark-brown spot, but not connected by a
dark-brown band ; bristles on the vibrissal ridges not extending
higher than the middle. Chitinised patches in roof of buccal
cavity brown. Antenne greyish-buff to cinereous, third joint
often strongly infuscated, except at the extreme base; second
joint usually more or less brown in front, except at the tip ;
arista buff, dark-brown at the base beneath. Bulb at base of
proboscis dark-brown to reddish-brown. Palpi buff, more or less
cinereous above, the tips infuscated.

Thorax.—The markings on the dorsum are vestiges of those
described in the case of Gl. palpalis ; they are usually more or
less indistinct ; pleure cinereous ; in rubbed specimens the dorsum
may appear reddish-brown. Scutellum buff, with the usual patch
on each side of the median line greyish-brown.

Abdomen.—In addition to the interrupted dark-brown bands,
an ill-defined brown patch on the second segment on each side,
not extending to the lateral or hind margins; the dark-brown
interrupted bands taper towards the lateral margins, and at
the median interruption, though sometimes straight-cut, are
generally more or less rounded off; on the posterior angles of
the segments the pale hind margins are continuous with what in
Gl. palpalis and Gi. pallicera are the lateral triangles ; seventh
segment and hypopygium in the g cinereous buff; seventh
segment in the 9 with a more or less distinct indication of
the interrupted dark-brown band at the base.

Legs buff-coloured, the front femora on the inside, the middle
and hind femora on the outside more or less infuscated ; some-
times a smal] dark patch on the outside of the front femora also
beyond the middle; hind tarsi with the last two jomts (except
the extreme base of the penultimate) entirety black.

ORIGINAL FIGURE OF GL. MORSITANS. 83

Wings faintly brownish. Squamez white, fringe of antisquama
darker, as usual. Halteres pale yellow.

With reference to Westwood’s original figure of Gl. morsitans
(P. Z. S. 1850, Pl. xix, Fig. 2), it is perhaps worthy of note that
the head of the typical specimen, which is there represented, has
at some time or other become detached from the body, and has
subsequently been glued on in an unnatural position, the vertex
being attached to the front of the thorax’in such a way that the
long axis of the head is almost horizontal. Curiously enough this
defect was faithfully reproduced by Westwood in his drawing,
which thus gives an altogether distorted idea of the appearance
of the head of the insect when seen from above.* Westwood’s
type is in very poor condition, especially as regards the thorax,
which is denuded and almost entirely destitute of the greyish
dust with which it is normally clothed, so that the ground colour
appears ochraceous-buff, with four rather broad black stripes,
interrupted at the suture, the lateral stripes abbreviated in front,
the middle pair behind. These stripes, which are shown in
Westwood’s figure, are not nearly so distinct in well-preserved
specimens, since they are normally to a large extent concealed by
the greyish dust with which the ground colour of the thorax is
obscured.

DistRIBUTION OF GI. morsitans, Westw.

The above description has been drawn up from an examination
of seventy-one specimens (50 ¢ ¢, 21 9 9), as follows :—
1 g, the type of the species, locality uncertaint (Major Frank

* Westwood’s figure was reproduced on the title-page of Livingstone’s
“Missionary Travels and Researches in South Africa,” and also on p. 571
of that work, where it is accompanied by a copy of Westwood’s figure of
the head and mouth parts, and by a rough original wood-cut representing
the fly nearly natural size. In the Appendix to Oates’ ‘‘ Matabele Land and
the Victoria Falls,” p. 364, Westwood mentions that his figure was copied
upon the title-page of Livingstone’s work ‘‘ without acknowledgment,” but
from Livingstone’s own statement (op. cif. p. 571) it is clear that he sinned
in ignorance, since the drawing was supplied to him by Mr. J. E. Gray,
of the British Museum.

+ The specimen bears the following label, in Westwood’s handwriting :—
‘Glossina morsitans, Westwood (Setse), fm. Lake Tchad, Central Africa,
Captn. Frank Vardon”; but this locality must be wrong. So far as I can
discover, Major Vardon never visited Lake Tchad, and the specimen is
almost certainly from the Siloquana Hills, which are in the north of the
Transvaal, in the angle formed by the Nylstroom River with the Limpopo.
The late Wm. Cotton Oswell, who was a companion of Vardon on his
shooting expeditions (cf. “‘ Big Game Shooting,” Vol. I., pp. 88, 89), writing
in “Big Game Shooting,” Vol. IL, p. 113 (The Badminton Library,
London, Longmans, Green & Co., 1894), says:——‘‘ On the low Siloquana
Hilla near this we mado our acquaintance with the Tsétsé fly, which we

qa 2

84 GLOSSINA MORSITANS.—LOCALITIES.

Vardon) [0.]; 1 g, Grahamstown,* Cape Colony (purchased
from Higgins, 1869) [0.]; 1 $, 1? Zululand [V.];.2 ¢ 6,
Little Crocodile R., two days’ march from Barberton, Transvaal,
28. x. 1888 (R. Crawshay); 2 g g,1 9, same locality and date
as the last (R. Crawshay) [C.]; 1 $, Loewe’s Creek, Barberton,
1893 (Dr. Percy Rendall) [W. L. Distant]; 9 ¢ g, Tzende R.,f
Zoutpansberg District, N.E. Transvaal, 2, 3. vii. 1896 (C. Hesel-
tine); 3 & $,1 9, between Tati and Gwailo R., Matabeleland,
1873 (F. Oates) [0.]; 2 gg, Beira Railway, Lower Pungwe
- River, Portuguese East Africa, Oct. 1897: “ Flew into railway
carriage at night, attracted by the light” (@. A. K. Marshall) ;
4 8,2 99, Umfuli R., Rhodesia, Sept. 1895 (G. A. K:
Marshall); 1 g, Northern Zambesia, between 15° and 18° §.;

were the first to bring to notice; Vardon taking or sending to England
some he caught on his favourite horse,” The year to which this refers
was apparently 1845 (cf. Oswell, op. cit., pp. 36, 87, 88). In a letter to
Westwood, dated May, 1850 (published by Westwood, P.Z.S. 1850, p. 260),
Vardon speaks of ‘‘the fly of South Africa so destructive to. cattle,” and
further on he writes: “The specimens you saw cost me one of the best
[horses] in my stud.” Kirk (Journ. Linn. Soc., Zoology, Vol. VII., p, 150)
speaks of Vardon as having ‘ hunted about the same time and in the same
region as Gordon Cumming,” and a little later Kirk goes on to say:
‘‘Major Vardon further performed a most interesting experiment. Aware
of the existence in certain districts of the Bechuana country of plants
poisonous to cattle (such as the ‘ Kofiwhane,’ a species of Lasiosiphon), and
suspecting that some such herb might be the eause of the mischief
ascribed by the natives to the fly, he put the matter to the test by riding
his horse into a ‘‘ Tsetse”’-infested part, without dismounting or allowing
the animal to feed; the result was the death of the horse.” It can scarcely
be doubted that this incident is the same as that referred to by Vardon
himself in his letter to Westwood quoted above, and also alluded to as we
have seen by his companion Oswell. If so, the true locality of the type of
Gl. morsitans may be regarded as established with sufficient certainty.

* This locality also requires confirmation: the specimen is labelled, in
Westwood’s handwriting—* Grahams Town, South Africa. Higgins,
1869”; but though Vardon was wrong in supposing that Glossina morsitans
‘is not met with south of the Tropic of Capricorn,” further evidence is
needed before we can believe that it oecurs in the South of Cape Colony,
within eighty miles of Port Elizabeth.

+ The total number of specimens of Gl.‘ morsitans brought back by
Mr. Heseltine from this locality was fifteen, all of which were males.
From maps kindly lent me by Mr. Heseltine, with the localities marked,
it appears that he fouiid the fly on the Singwedsi River, where it crosses
the boundary between the Transvaal and Portuguése’E. Africa, just south
of Lat. 23° S., and west of Long, 32° EK. ‘This district:is within. thirty
miles of the valley: of the Limpopo,.and’the Singwedsi runs S.E: to join
Olifant’s River, a tributary of the Limpopo. Mr. Heseltine also met with
the fly on the Great Letaba River, where it turns N.E.: and crosses
Long, 31° E., while the specimens captured were taken on the Tzende
River where it crosses the Tropic of Capricorn, a few minutes west of
‘Long. 82° E, The Tzende River runs nearly due south to join the Great
‘Letaba River, which falls into Olifant’s River. : All. these ‘localities are sin
‘the N.E. corner of the Transvaal, 8's 6 Ge 5 Cow aid? sang ails

ae

DISTRIBUTION OF GL. MORSITANS. 85

and 24° and 28° E. (Wm. Oswell)*; 1 g, same locality as the
preceding (Wm. Oswell) [0.]; 2 66,2 99, Lake Mweru
(S.W. of Lake Tanganyika), 31. vii. 1892 (RB. Orawshay) {C.| ;
I 9%, Henga, circa 3,300 ft. (south-west from northern end of
L. Nyasa, between 10° and 11° S. lat.), between July and Dec.,
1893 (R. Crawshay) [C.]; 19, Msongozi’s, Loangwa Valley,
Senga, British Central Africa, 2,300 ft., Sept. 3, 1895
(RB. Crawshay); 2 6g, 2 99, Kilima Njaro (F. J. Jackson);
266, “E. Africa” (Capt. Speke); 3 ¢ g (fragments gummed
on card), ? Locality (Dr. Livingstone) [O.]; 10 gg, 10 9? 9,
Togo, W. Africa (Dr. Schilling) [Royal Zoological Museum,
Berlin]; 1 ¢, ?Locality and collector [O.]; 2 ¢ 6, ? Locality
and collector [V.].

Since Glossina morsitans has hitherto undoubtedly been
confounded with Gl. pallidipes (not to speak of Gl. longipalpis)
under the comprehensive term “ Tsetse-fly,” it is impossible to
draw any trustworthy conclusions as to the precise geographical
range of this species from statements in records of African travel
and sport. <A study of the foregoing list of localities of actual
specimens, however, will show that Gl. morsitans certainly occurs
—not, of course, continuously, but in “ fly-belts ”"—from Zululand
and the south-eastern portion of the Transvaal in the south to
Kilima Njaro in the north, while the fact that the species has
recently been collected by Dr. Schilling in Togoland on the
Slave Coast, shows that it can no longer be regarded as confined
to South or even Central Africa, but that its limits are very
much wider than has hitherto been supposed, As will be shown
below, it is probable that the species which was the subject
of Lt.-Col. Bruce’s memorable investigations in Zululand was
Glossina pallidipes and not Gl. morsitans, and if this is the case
the eastern halves of the areas of distribution of the two species
practically coincide. For while Gl. morsitans also occurs in
Zululand (two specimens from this region are in the collection
of Mr. G. H. Verrall), as has already been seen, the Museum
possesses specimens from Kilima Njaro, which is less than one
hundred miles south of Witu Forest, the most northerly locality
from’ which specimens of Gl. pallidipes have at present been
received. Until a large amount of material has been systemati-
cally collected, any more precise statement as to the relative
distribution of Gi. morsitans and Gi. pallidipes 1 is impossible.

oe Cf. Trans, Ent, Soe. Lond, New’ Ser., Vol, He p. 96 (1852, 1853)
Ann, Mag. Nat. Hist., Ser. 2, Vol. X. (1863), “OM

86 GL. MORSITANS.—RATIO OF SEXES.

Hapsirts, etc. (See Chapter I.).

It has already been pointed out at the commencement of
Chapter I. that the name Glossina morsitans is usually quoted as
if it were that of the only known species of the genus. Although
this, of course, is a mistake, nevertheless, if it is permissible
to draw any conclusion from the material at present available
for examination, Gl. morsitans is. one of the two commonest
Tsetse-flies of South and Central Africa. But since its range in
this region is practically co-extensive with that of Gl. pallidipes,
all statements in accounts of South African travel and sport as
to the occurrence and habits of “the Tsetse-fly ” or “the South
African Tsetse-fly ” must for the present be regarded as referring
to both species indiscriminately : while owing to the fact that
both Glossina longipalpis and Gl. palpalis would appear also to
occur on the Zambesi, some of the phenomena hitherto attributed
to Gl. morsitans should perhaps really be ascribed to one or both
of the latter species. With this explanation, readers who desire
to learn the habits of Glossina morsitans may be referred to
Chapter I., where all that is known on this subject will be found
fully ateascesl:

It will have been noticed that, in the foregoing list of speci-
mens examined, out of seventy-one examples only twenty-one are
females. ‘The whole of the fifteen specimens, too, brought back
by Mr. Christopher Heseltine from the Tzende River were males,
as were also the six specimens of Glossina palpalis taken by the
author in Sierra Leone; while out of fifty specimens of Gl.
pallidipes from Machakos, only three are females. This would
suggest the idea that in some at least of the species of Glossina
the two sexes may have slightly different habits (although they
both suck blood *), in consequence of which the female is more
easily able to escape observation or avoid capture: it should be
noted, however, that in the twenty specimens of Gl. morsitans
collected by Dr. Schilling, in Togo, the sexes are equally divided.
In view of the above facts there can be little doubt that Sir
John Kirk’s statement [28] that only on one occasion did he
“obtain two of what may be the male insect,” was due to his
having confused the sexes, unless indeed he happened to have
collected at a season when the usual ratio of the sexes was reversed.

* Among the Tabanidw (Horse- flies), in which the females alone suck

blood, while the males frequent flowers, the females are much the more
common,

DESCRIPTION OF GLOSSINA PALLIDIPES. 87

SyNoNYMY AND AFFINITIES.

There can be no question of the correctness of the identifica-
tion of Glossina morsitans, Westw., as defined above, and although
the specific name has no doubt often been used loosely in what
may be termed a generic manner, while Continental writers have
occasionally referred to the species as if it were Wiedemann’s
longipalpis, the species. itself has never been re-described under
any synonym,

Glossina morsitans, Westw., Gl. longipalpis, Wied., and Gl.
pallidipes, sp. nov., form a group of closely allied species, of which
Gl. palpalis, Rob.-Desv., is a somewhat more distant connection.
A discussion of the distinctive characters will be found on
pp- 93-95, under Gl. longipalpis.

Glossina pallidipes, sp. nov.
. (Plate IV.)

6, ?. Length, 8 to 10 millim. (33 to 43 lin.) ; length of wing,
82 to 94 millim. (41 to 43 lin.) ; width of head in male, 3 millim.
(1} lin.), in 9 23 millim. (14 lin.). Width of front at-vertex in
dé +, in 2 between + and } of total width of head measured

across middle of eyes.

Precisely similar to Gl. morsitans in colouration and general
appearance, but distinguishable at once by the front and middle
tarsi being entirely yellow *; the interrupted bands on the abdomen
generally darker, and approaching closer to the hind margins of the
segments (the pale hind margin usually confined to the posterior siath
or fifth of each segment).

As compared with Gl. morsitans, this species, in addition to
the differences already mentioned, is often distinctly larger ; the
eyes are larger and more prominent, while the front in the ¢ is
(actually and not merely relatively) narrower, and also as a rule
somewhat darker towards the vertex; the ground-colour of the
abdomen is ochraceous-buff to ochraceous ; the pale hind margins
of the segments from the 3rd to the 6th inclusive are narrower
than in Gl. morsitans, and sometimes exceedingly narrow, so that

* In specimens preserved in spirit the last two joints of the front tarsi
sometimes appear faintly tipped with pale brown.

88 VARIETIES OF GLOSSINA PALLIDIPES..

the interrupted dark-brown bands occupy more of the upper
surface of the abdomen.

Legs buff-coloured ; middle tarsi, like the front pair, entirely
pale* ; the last two joints of the hind tarsi (except the penultimate
at the extreme base) dark-brown.

The typical specimens are a ¢ and @ from Kilima Njaro
(F. J. Jackson), in the British Museum collection.

Varieties A ¢ and 9 from Witu, “ caught in Witu town”
(British East Africa Company), are much below the normal size,
since they do not exceed 7} millim. (34 lin.) in length; they
are further distinguished by the pale longitudinal stripe on the
median line of the abdomen being wider than usual, especially
in the 9, in which also it is less sharply defined, owing to the
interrupted dark transverse bands dying away towards the
median line. In the shape and size of the eyes, and especially
in the narrowness of the front in the male, however, these
specimens agree with the typica! form.

Several specimens from British East Africa (Vet.-Capt.
A, J. Haslam) and Kilima Njaro (F. J. Jackson), though other-
wise agreeing with the typical form, have the median pale
longitudinal stripe on the abdomen wider and less sharply
detined, since the interrupted dark transverse bands die away
towards the median line. Possibly this appearance is in part at
least due to immaturity.

Distrisution or Gil. pallidipes, sp. nov.

Of this species, which would appear to be the East African
representative of Gl. longipalpis, Wied., I have examined a series
of 81 specimens (63 g¢ g and 18 ? @), from various localities, as
indicated below :—

1 ¢, locality unknown (forwarded for identification by
Mr. J. F. Quekett, Durban Museum) ; 1 9, “Country south of
Mashonaland ” (Mr. Hooke: presented by the late J. C. Mansel-
Pleydell); 1 ¢, Shire Highlands, 1893 (H. H, Johnston.—
“Caught in a part of the Shire Highlands where we had thought
there was no Tsetse”: extract from letter from Mr. Johnston
(now Sir H. H. Johnston, K.C.B.), dated October 22nd, 1893, to
the late Sir Wm. Flower, K.C.B., enclosing specimen) ; 2 ¢ 6,
British Central Africa—exact locality uncertain, but probably
Lunyina River, Henga—1895 (R. Crawshay); 1 6, Lake

* See note on page 87.

LOCALITIES.—SPECIES STUDIED BY BRUCE, 89

Mweru, 18-22, m1. 1892 (R. Crawshay); 3 6 6, 32 @ (in
cluding the types of the species), Kilima Njaro (F. J. Jackson) ;
1 9, Kiboko River, Uganda Railway, British East Africa,
Nov., 1898 (R. Crawshay :—‘‘settled on back of collector’s neck
and bit him”: letter from Mr. L. R. Crawshay, April 13, 1902)
[C.]; 19, Upper Sabaki River, 1890 (presented by the late British
East Africa Company); 3 ¢ ¢,1 9, Uganda Railway, British
East Africa, 1897-98 (Vet.-Capt. A. J. Haslam); 1 9, Uganda
Railway (C. S. Betton); 47 6 6, 3 9 9, Machakos, 1897, and
246 46,2 99, Tsavo River (received from the Tsetse-fly Com-
mittee of the Royal Society, per Lt.-Col. Bruce); 1 6,2 2 9;
“KE. Africa” (Capt. Speke); 2 6 6, 1 9 (including the two
specimens of the small variety mentioned above), “caught in
Witu Town,” 1891 (Officer in charge of British East Africa
Company’s Station at Lamu: presented by the late British East
Africa Company); 2 ? 9, Witu Forest, 1895 (the Sultan of
Witu : presented by the late W. 8. Godfrey).

It has already been pointed out in dealing with the area of
distribution of Gl. morsitans that the eastern half of it practically
coincides with that of the present species, and reference has been
made to the probability that the species that formed the subject
of Lt.-Col. Bruce’s experiments in Zulu Land was Gl. pallidipes
and not Gl. morsitans. Such at any rate is the natural deduction
from a comparison of the reproductions of photographs of speci-
mens on Plate II. of Col. Bruce’s memoir [142]; on comparing
fig. 2 and the right-hand figure of fig. 7, it will be seen that the
spas pale streak on the ps aa is very narrow, and that the
dark interrupted bands extend close to the posterior borders of
the segments. The coloured figures on Plate I. are evidently less
reliable, and no importance can be attached to the obvious
discrepancies between them and the photographic reproductions
on the following plate. In the coloured figures the angles of the
interrupted dark abdominal bands are shown rounded off, while
the pale hind margins are deeper and the median stripe wider, so
that to judge from the abdomen these figures would appear to
have been taken from specimens of GI. morsitans. On the other
hand, the front tarsi are entirely pale ; but, since the hind tarsi
are likewise shown without dark tips, no conclusion can be
drawn from this, except that the artist certainly did not examine
the legs of his specimens closely. It is quite possible that in his
experiments Bruce may have made use of both Gl. morsitans and
Gl. pallidipes, af

90 DESCRIPTION OF GLOSSINA LONGIPALPIS.

Hasits, ETC.

See Chapter I.

The fact that, as recorded above, three specimens were
actually caught “in Witu Town” * is of special interest, since
it is commonly supposed that the Tsetse avoids the neighbourhood
of human habitations. Thus, Livingstone [21] writes of the
“well-known disgust which the Tsetse shows to animal excreta,
as exhibited when a village is placed in its habitat ;” and Sir
H. H. Johnston [145] says: “This insect has a... still
stronger dislike to a congeries of human habitations. . . . They
[horses and cattle] are also quite safe in the middle of any collec-
tion of huts or in any town.”

SYNONYMY AND AFFINITIES.

As has already been mentioned, Gl. pallidipes has not
hitherto been distinguished from Gl. morsitans, Westw., which
forms with it and Gl. longipalpis, Wied., a closely allied group.
Its distinctive characters will be discussed further in dealing
with the latter species (see p. 93), to which it presents an
exceedingly close resemblance.

Glossina longipalpis, Wied.
(Plate V.)

Glossina longipalpis, Wiedemann, ‘‘ Aussereuropiische zweifligelige
Insekten,” Zweiter Theil, p. 254, Taf. IX. fig. 10, a, b, ¢ (1830) ; Mac-
quart, Hist. Nat. des Ins. Diptéres., T. ii., p. 245 (1835); Dipt. Exot.,
T. ii., 3, p. 113, Tab. 14, fig. 1 (1843).

3, 2.—Length, 9 to 10 millim. (4} to 43 lin.); length of
wing, 8 to 94 millim. (33 to 4} lin.); width of head, in both

* The following is an extract from the ‘‘ Report of the Officer in Charge
of the British East Africa Company’s Station at Lamu” to the late British
East Africa Company, sending the specimens, and dated May 26, 1891 :—‘‘T
beg to forward specimens of two species of flies. The three smaller ones,
which I believe to be the true ‘‘ Tsetze,” or a species of it, I caught in Witu
Town, and caught others between Funga Sormbe and Witu. This fly is
also plentiful along the road between Mombasa—Gulu-Gulu—Teita and
Kibwezi in Ukambani. Its bite, though very sharp and needle-like, causes
little or no irritation.”” The other flies sent were specimens of Tabanus
latipes, Macq., one of the commonest of the African Tabanide (horse-flies),
bee ranges from Senegal to British Hast Africa and southwards ‘to

atal.

DESCRIPTION OF GLOSSINA LONGIPALPIS. 91

sexes, 3 millim. (1} lin.) ; width of front at vertex in ¢ |, in
? between } and } of total width of head measured. across
middle of eyes.

Thorax olivaceous-gray, with the usual dark-brown longitudinal
markings ; abdomen ochraceous-buff,* with interrupted dark bands,
which stand out conspicuously from ground-colour, and leave hind
margins of segments only narrowly pale; sides of front in g
parallel, in 9 converging slightly towards vertex ; legs ochraceous-
buff, last two joints of front and middle tarsi with sharply defined
black tips as in Gl. morsitans.

Very close to Gl. pallidipes, Austen, but distinguished at
at once by the black tips to the last two joints of the front and
middle tarsi. Eyes in ¢ large and tumid, resembling the eyes
of Gl. pallidipes more than those of Gl. morsitans. Front of ¢
not quite so narrow as that of ¢ pallidipes; frontal stripe
tawny, darker above, frontal margins and vertical triangle oli-
vaceous-gray ; elongate dark-brown spot on each side of frontal
stripe below generally very conspicuous, as is also the narrow
dark-brown band connecting the bases of the vertical bristles.
Dorsum of thorax with the brown markings described in the
case of Gl. palpalis fairly distinct. Scutellum buff, grey at the
base, with a strongly marked dark-brown triangular spot on
each side of the median line.

Abdomen, with the usual interrupted dark-brown bands on
the third to the sixth segments inclusive, strongly marked and
deep, precisely as in Gil. pallidipes, leaving the hind margins of
the segments only narrowly pale.

Legs buff, the front femora strongly infuscated on the inner
side, and generally more or less infuscated on the outer side also.
Middle femora usually more or less infuscated on the outer and
inner side; hind femora slightly infuscated on the outside
towards the distal extremity. Middle and hind femora often
with a strongly marked elongate black spot on the under side near
the end,

Distripution oF Gil. longipalpis, Wied.

The above description is based on an examination of twenty-
three specimens (14 ¢ ¢,9 9 9) from the following localities :—
1 g, the type of the species, ‘‘Guinea ” [Imperial and Royal

* Ridgway, ‘Nomenclature of Colors,” Pl. V. fig. 10.

92 TYPE OF GLOSSINA LONGIPALPIS.

Natural History Museum, Vienna]; 1 9, locality uncertain,
labelled “ Glossina longipalpis g, Wied.” in what is probably
Macquart’s handwriting [V.]; 12 $64, 8 99, Togo (Dr.
Schilling) [Royal Zoological Museum, Berlin]; 1 ¢, “ Zambesi”
(Dr. (now Sir) John Kirk).

Hasirs, ETc.

See p. 86. We have as yet no special information concerning
the habits of Gl. longipalpis, Wied.

SYNONYMY AND AFFINITIES.

A glance at the synonymy already given under Glossina
palpalis, Rob.-Desv., and Gl. morsitans, Westw., will show that
both of these species have at one time or another been mis-
taken for Gl. longipalpis, Wied. As pointed out on p. 77,
to judge merely from the description, the latter species would
seem to have been described from a specimen of Gl. palpalis
“with pale legs, and well-marked pale median stripe and hind
margins to the abdominal segments (i.e. var. tachinoides)” ;
moreover, as is further pointed out in Note (ft) on the same
page, Wiedemann’s figure, valueless as it is, would support this
conclusion “by conveying the impression of a species with very
dark abdomen and well-marked pale hind margins to: the
abdominal segments.” With only the description to rely on,
the present writer had previously identified the Tsetse-flies found
by him at Sierra Leone in 1899 as GI. longipalpis, Wied.,*—a
determination made with the greater confidence, since Wiedemann
states in his description that the type of the species is from
Sierra Leone, where it had been discovered by Adam Afzelius.t
On examining the specimen received from the Vienna Museum
as the type, however, it was at once seen that this belongs to a
species quite distinct from that described above as Gl. palpalis,
Rob.-Desv., and a doubt naturally arose as to whether the so-called
type was the true one, especially as the label attached to’ the
specimen bears the locality “Guinea” (instead of Sierra Leone),
in addition to the words “Glossina longipalpis, Wied.” TI there-
fore applied again to Herr Josef Bischof, of the Vienna Museum,
from whom I had received the insect, and obtained the following

* Cf. Austen [162]. th tegen

+ A well-known Swedish botanist, who visited Free. Town, Sierra

Leone, in the years 1792 and 1794, and made general collections of
zoological and botanical specimens. ~

GLOSSINA MORSITANS AND ALLIES. 93

reply, dated “ Vienna, 18. iii. 1902.”—*‘ The specimen sent is the
original type. The label placed beside it is Wiedemann’s original
one, and bears his handwriting.” In view of this statement it is
useless to dilate further upon apparent discrepancies between
description and type, and the species described above must be
taken as the true Glossina longipalpis, Wied.

As has already been stated in speaking of the affinities of
Gl. morsitans (p. 87) :—*“ Glossina. morsitans, Westw., Gl. longi-
palpis, Wied., and GI. pallidipes, sp. nov., form a group of closely
allied species, of which Gl. palpalis, Rob.-Desy. is a somewhat
more distant connection.” Through all of these species there can
be traced with greater or less distinctness the fundamental
system of abdominal markings, consisting essentially of a series
of dark transverse bands, interrupted on the median line, and
occupying a larger or smaller area of the upper surface of the
segments. Gl. palpalis, however, is distinguished at once by its
entirely dark hind tarsi.

Of the remaining species Gl. pallidipes is distinguished from
the other two by the front and middle tarsi being entirely pale,
and in the ¢ by the narrower front.. The awe: of the dark
tips to the last two joints of the front and middle tarsi seems
to be a reliable character, and one that cannot be due to possible
bleaching or immaturity, since the last two joints of the hind
tarsi are invariably dark. Altogether there need, I think, be
no hesitation in accepting Glossina pallidipes as a good species.
Apart from the tarsi, females of GJ. pallidipes are not easy to
distinguish from females of Gl. longipalpis, though from females
of Gl. morsitans they are readily “idiseniiable by the depth of
the abdominal bands. In both sexes, too, Gi. pallidipes may
easily be distinguished from Gl. morsitans by the shape of the
third joint of sn antenna, which in the former species is long
and narrow, with the apex pointed and (especially in the female)
conspicuously turned forwards; in GI. morsitans the third joint
of the antenna is shorter and much broader (about half as broad
again as in pallidipes), and darker towards the tip, which is
blunter and not so much turned forwards; the arista is also
shorter in morsitans than in pallidipes. To Laanilid ate comparison,
the antennz should be removed with the point of a fine scalpel,
mounted in glycerine and examined under a lens. As regards
the males, even though the front and middle tarsi be wanting,
a male of pallidipes can ‘be distinguished from a male of longi-
palpis, by the front. being narrower, especially at: the yertex,

94 HYPOPYGIUM.

towards which its sides distinctly converge; and a male of Gl.
longipalpis can be distinguished from males of both morsitans and
pallidipes by the inner margins of the eyes, forming the sides

Fig. 12.
Distal portion of abdomen of Glossina morsitans, Westw. (ventral aspect),
showing (hk) hypopygium. (x 15.)

Fig. 13.

Distal portion of abdomen of Glossina longipalpis, Wied. (ventral aspect),
showing (hk) hypopygium. (x 15.)

of the front, being parallel instead of converging towards the
vertex. Similarly, the shape and size of the hypopygium, as
already described in the “Synopsis of Species” (see pp. 70-71),

DESCRIPTION OF GLOSSINA FUSCA. 95

will serve to distinguish a male of longipalpis from a specimen
of morsitans belonging to the same sex (see Figs. 12 and 13).*

We have as yet no information as to the western limit of
Gl. pallidipes, so that at present it may be regarded as the
eastern representative of Gl. longipalpis.

Glossina fusca, Walk.
(Plate VI.)

Stomoxys fuscus, Walker, List Dipt. Ins. in Coll. Brit. Mus. Pt, ITI.
p. 682 (1849).— ¢.

Glossina, tabaniformis, Westwood, Proc. Zool. Soc. Lond. Pt. XVIII.
p. 268, Pl. XIX. fig. 3 (1850).—¢.

Glossina fusca, Walker, “ Entomologist,” VI. p. 328 (1873).

Glossina grossa, Bigot, Ann, Soc. Ent. Fr. Vol. LX. p, 377 (1891).— 9.

Glossina tabaniformis, Stuhlmann, Ber. iib. Land- u. Forstwirtsch. in
Deutsch-Ostafrika, 1 Bd., Heft. 2, p. 173 (1902).

3, %.—Length 11 to 12 millim. (5} to 53 lin.) ; length of
wing 102 to 13 millim. (5 to 6} lin.); wing expanse of largest
3g 26 millim. (124 lin.); wing expanse of largest ? 29 millim.
(13% lin.); width of head in g 33 millim. (1# lin.); in 9
31 millim. (Walker’s sy a) to 33 millim. (1} to 1} lin.); width
of front at vertex, in g 3 millim. (4 lin.), in 9 3 to 1 millim.
(just under } lin.).

Thorax drab-gray ¢ to grayish brown, with narrow darker longi-
tudinal stripes ; abdomen russet brown,t segments after the second
mummy brown,§ hind margins towards posterior angles sometimes
paler ; proboscis bulb pale yellow ; legs buff, middle and hind tibie
sometimes with a well marked dark ring, more or less incomplete,
round middle ; wings yellowish brown to brownish, sometimes larger
and broader in Q thanin 2, anterior and posterior transverse veins
often infuscated.

Head.—Buff, posterior surface grayish ; frontal stripe usually
somewhat darker, sometimes ochraceous ; face, including facial

* The shape and size of the hypopygium in Gl. pallidipes are the same
as in Gl. longipalpis.

+ Ridgway, “‘ Nomenclature of Colors,” Pl. i. fig. 13,

t Zbid. Pl. III. fig. 16.

§ Ibid. Pl. III. fig. 10.

96 GLOSSINA FUSCA.—DESCRIPTION.

pit, and frontal margins narrowly, shimmering whitish yellow ;
ocellar spot not, or scarcely darker than frontal stripe ; ocelli
large, anterior ocellus the largest. Bases of the vertical bristles
united by a more or less conspicuous brown band. Antenne buff ;
second joint on inside, and distal half or two-thirds of third joint
brown; second joint sometimes entirely brown. Arista buff,
second joint dark-brown at base beneath. Palpi buff, upper
surface grayish brown, and rather more than apical quarter
brown ; the row of short erect black bristles along upper margin
very conspicuous. *

Thorax.—Series of atic on dorsum, as described for
Gl. palpalis, much reduced and but little conspicuous ; most
distinct are the two narrow ad-median stripes in front of the
suture, the narrow curved stripe outside these on each side,
sometimes with prolongation to anterior margin of thorax, and
two faint blotches enclosed by the curved stripe, one in front
of and one behind the transverse suture; the small spot on the
suture, between the ad-median and the curved stripe on each side
more or less distinct ; spot on inner margin of humeral callus
usually very faint. Scutellum grayish-brown; margin and
impressed median line buff.

Abdomen.—The longer hair at the base of the second segment
golden-yellow in the middle, dark-brown or black at the sides ;
third, fourth, and fifth segments very short ; a pale median stripe
wanting, or scarcely visible if at all, but posterior angles and
(usually) more or less of extreme hind margins of the third to
the sixth segments inclusive yellowish, so that the colour of these
segments is obviously due to dark transverse bands, as in the
case of species previously described; seventh segment and
hypopygium of ¢ clothed with grayish-yellow dust..

- Legs.—Front femora sometimes more or less infuscated on
inner side ; middle femora with a brownish blotch on under side
near tip ; solitary bristle on upper side of middle femora near the
tip, stout and conspicuous; front tibie sometimes with. dark
brown blotch on inside in middle, representing ring sometimes
seen on middle and hind tibie; extreme tips of front and hind
tibie and extreme ‘base of first joint:of front and hind tarsi
reddish-brown on under side; front and hind tibie towards the
tips and first three joints of front and hind tarsi clothed beneath

* In Walker’s type (a ?) the bristles on the margin of the buccal

cavity below the vibrissal angle are golden-yellow ey of are. a trace
of this is sometimes seen in other specimens also.’s. 3. - :

WINGS IN GLOSSINA FUSCA. 97

with short golden pile ; last two joints of front tarsi more or less
infuscated at the tips ; last two joints of middle and hind tarsi,
except basal half of penultimate joint of middle tarsi, dark brown ;
sometimes extreme tips of remaining tarsal joints infuscated ;
claws black, russet-brown at base.

_ Wings.— Veins pale tawny ; upper portion of anterior trans-
verse vein often strongly incrassated, and, together with adjacent
portion of fourth vein lying towards base of wing, as well as
posterior transverse vein and portion of fourth vein immediately
beyond, conspicuously infuscated ; basal portion of the veins from
the second to the sixth also darker. Squamz not infuscated,
fringed with rather long pale-brown fine silky hair. Halteres
yellowish.

Since Walker’s type is a 9, and not in the best of condition,
the above description has been drawn up from a g from Kaporo,
1670 ft., near Songwi River, N.W. of Lake Nyasa, British
Central Africa Protectorate, 28. ii. 1895 (R. Crawshay), “Taken
on the path at night,” and a ? from Witu Forest, 1895 (Sulian
of Witu, presented by the late W. 8S. Godfrey); both of these
specimens are in the collection of the British Museum, and have
been specially labelled.

In certain specimens, including, as it happens, both the type
of the species and also that of Gl. tabaniformis, Westw., the
anterior transverse vein is less incrassated above than usual, and,
like the adjacent portion of the fourth vein lying towards the
base of the wing and the posterior transverse vein, is also not
infuscated. In these specimens the wing is usually somewhat
darker than in those in which the anterior transverse vein is
more strongly developed, while, owing to the absence of the
infuscations alluded to, it appears of a more uniform brown.
There can, however, I think, be no question of a specific difference.
The type of GI. grossa, Bigot, is an unusually dark specimen, with
the second joint of the antenne and the abdominal segments after
the second dark brown; the markings on the dorsum of the
thorax are also unusually distinct, and the apex of the third
joint of the antennz is very prominent.

It has recently been stated by Stuhlmann [174] that in Gl.
fusca, which he calls Gl. tabaniformis, the ocellar bristles on the
crown of the head are wanting, and also that in the resting
position the wings are slightly tectiform. Neither of these state-

i

98 GLOSSINA FUSCA.—LOCALITIES.

ments, however, is in accordance with fact. An examination of
well-preserved’ specimens of Gl. fusta shows that’ greater’ ocellar
bristles ‘are’ present in this as in ‘all other species of! Glossina ;
whilé’ a specimen from’ Witu Forest in. the British Museum
Collection, which has died and been pinned in the: resting: posi-
tion, has its wings closed flat over one another just ‘like the
blades of ‘a pair of scissors. \ Moreover, having also examined
specimens of Glossina longipennis,, Corti, palpalis, Rob.-Desv., and
pallidipes, Austen, all pinned in the resting attitude, and burving
had the opportunity of observing Gil. palpalis when-alive, I have
no hesitation in confirming the statement, as to the resting posi-
tion of the wings in Glossina, in the diagnosis of the genus at the
commencement of the present Chapter. .

DistRiBuTion oF Gi. feet Wallk

Of this Bode I have been able to examine a series of 2
examples (8. 6g and 14 99), from various - Tocalities, as
follows :— .

Pt); poe type of the species, locality unknown ; I 9, the
type of Glossina tabaniformis, Westw., Gold Coast* [0]; I é,
299, Gold Coast (received from the Tsetse- fly Committee of
the Royal Society, per Lt.-Col. Bruce); 1 ¢, 1 vs Togo
(Dr. Schilling) [Royal , Zoological Museum, Berlin]; 1 9, the
type of Glossina grossa, Big. , Assinie, Ivory Coast (Ch. Alluaud)
hes a 9, Asaba, R. ives (Dr. W. H. Crosse) ; et
“country south of Mashonaland 0 (Mr. Hooke: presented by the
late J. C. Mansel- Pleydell) ; 1 9, Zambesi (Dr. (afterwards Sir)
John Kirk) ; : 2 66 (including the specimen described in the
above re-description), Kapor o, 1670 ft., near Songwi River, north-
west of Lake Nyasa, 28. ii. 1895, taken on Hie” path at night ”
(R. Crawshay y); 1 g,, same locality, 25. ii, 1895 (R. Crawshay) [C.];
1 2, Kilima Njaro Ki A J. Jackson) ; 9: Upper Sabaki River,
British East Africa (presented by the late British East: Africa
Company) ; oe Uganda Railw ay, British East Africa, 1 Kiboko

* Westwood [10] writes: “‘ Hab. apud littus aureum Afric tropicalis
occidentalis (Mus. D. Hope).” The. specimen itself bears. onthe label
“« Raddon,, Africa,” which probably means that it was at one time in the
collection of W. Raddon, an English entomologist of the earlier part of
last century, well known: for his discovery ofthe caterpillars of the Spurge
Hawk-Moth in numbers in: North Devon i in the year 1814: cf. W. Raddon,
‘“Notes on Detlephila, Mal bie” : 1 piorolagiaat Pais spree Il.
pp. 535+586 (1835). ¥ Se : a

DISTRIBUTION AND HABITS OF GL. FUSCA. 99

tiver or Sabaki River, near its junction with the Tsavo River,*
1898 (Vet.-Capt. A. J. Haslam); 2 6 6, 2 9 2 (including the
? described in the above re-description), Witu Forest, British
East Africa (the Sultan of Witu; presented by the late W. 8.
Godfrey) ; 1 9, locality unknown.

The above list of localities shows that the present species
occurs both in West and East Africa, so that it doubtless
extends right across the continent, but as to whether it exists
on the headwaters of the Congo, or at other intervening points,
we have as yet no information. In West Africa the most
northerly locality from which the species has been recorded is
Bismarckburg, in the Hinterland of Togo, the approximate lati-
tude of which is 8° 12’ N., about 10° 32’ further north than Witu,
the most northerly locality on the east coast. The most southerly
locality is somewhat vaguely represented by Mr. Hooke’s specimen
from “country south of Mashonaland.”

‘Hasrrs, BTC.

It al be seen later on that, sidan to the doeisliAhtale
paragraph of Captain Crawshay’s ‘‘ Note on the ‘Tse-Tse’ Flies
of the British Central Africa Protectorate” (see Chapter VII.
Appendix B, p. 289), the specimens of this species taken at
Kaporo did not bite; but this was doubtless due to the fact
that they were ‘captured at eee and not during the heat of
the day.
ele ee letter dated 16.4.1898, forwarding the four’ specimens
of Glossina pallidipes, and the single example of Gl. fusca
already referred to, as’ well as a ae specimen of Gl. longi-
pennis, Corti, as mentioned below, the late Capt. A. J. Haslam,
at that time Transport, Officer to the Uganda Railway, wrote
as follows about what he termed the ‘‘larger” Tsetse, a designa-
tion that under the circumstances must be taken to include
both Gl. fusca and Gi. longipennis. —‘The larger one .. . is
the one found abundantly at Kiboko River (a very deadly fly
district), and on the Sabaki River near its junction with the
Tsavo River. ‘This large kind is larger than the South African
Tsetse. Its flight is typical, Endl like the other kinds it
flies low—going for the bare legs of porters and the legs of
animals. . . . I am not yct in a position to state whether the
wings of the larger kind actually cross, as do those of the

* See alow @icer iB Hila, i Be

100. GL. FUSCA.—SYNONYMY AND AFFINITIES.

smaller kind, when sucking blood, but this I will soon be in a
position to report upon. . . . I am as yet unable to say whether
a different organism is harboured by these flies.” *

SYNONYMY AND AFFINITIES.

In the year 1873 (“ Entomologist,” Vol. VI., p. 328) Walker
sunk Gl. fusca as a synonym of Gl. longipalpis, Wied., with
which of course it has nothing whatever to do. It is a perfectly
good species, and with Gl. longipennis, Corti, to which it is very
near akin, it forms a group which, by reason of the considerably
larger size of the individuals, contrasts strongly with the other
species of the genus at present known. The differences between
Glossina fusca and Gl. longipennis will be dealt with below under
the latter species.

* Captain Haslam’s work was unfortunately not destined to be com-
pleted, as he lost his life three months after the above-quoted letter was
written, while pursuing his investigations upon Tsetse-fly disease in British
East Africa. I am indebted to the courtesy of his friend Dr. H. C. L.
Scofield, of North Dulwich, for information relating to Captain Haslam’s
end. Dr. Scofield kindly forwarded to me for perusal letters received by
him from Captain Cooper, 7th Dragoon Guards, and Dr. A. T. White,
Medical Officer, Uganda Protectorate, who found and buried Captain
Haslam’s body, on July 21, 1898.

Captain Haslam was murdered by Mulukas (a tribe of the Wakikuyu)
on July 17, 1898, near the Theka Theka River, Kenia district, British East
Africa, while on an expedition in connection with his investigations upon
Tsetse-fly disease. He was killed by a spear-wound through the back, and
his body (much mutilated) was found four days later by Captain Cooper
and Dr. White, who were returning from an expedition to punish the
natives for an attack on Captain Cooper’s camp. Captain Haslam’s
remains were interred, with military honours, close to the spot where
they were found, but were afterwards taken into Kikuyu and buried by
the side of the grave of Captain Nelson, who had travelled with Stanley.
Of the results already achieved by Captain Haslam before his untimely
death, Dr. White writes as follows: ‘‘I only hope that some means will be
found of publishing his notes on the Fly disease, at which he had done so
much valuable work. I met him for some time at Machakos, where we
did a good deal of work together at the microscopic examination of the
blood of the local Horse-flies, not the Tsetse, and one could not help being
struck by the amount of work he had already done,—work of the very
greatest value for the future of this country.”

That Captain Haslam fully appreciated the possibility that the parasite
of Nagana may be conveyed by blood-sucking flies other than Glossina is
evident, and his discovery of Trypanosoma in a species of Stomoxys
is alluded to in Chapter VII., Appendix EK (p. 304). It is therefore
with keen regret that I learn that Dr. White’s hopes are vain, and
that yet another name must be added to the roll of those martyrs to
science who have perished and ‘‘have no memorial.” In a letter to the
author on June 4, 1902, Dr. Scofield wrote: ‘‘I have received further
communications stating that the papers containing Captain Haslam’s
notes of his researches were hopelessly lost in the looting following his

murder.”

DESCRIPTION OF GLOSSINA LONGIPENNIS. 101

Glossina longipennis, Corti.
(Plate VIL.)

Glossina longipennis, Corti, Ann. Mus. Civ. Genov, Serie 2*, Vol. XV,
(XXXYV.), p. 1388 (1895); Hough, Proc. Acad. Nat. Sci. Philadelphia,
1898, p. 172; Austen, Proc. Zool. Soc. Lond. 1900, p. 10.

6, 9.—Length, 102 to 11} millim. (5 to 53 lin.) ; length of
wing, 113 to 12 millim. (53 to 52 lin.) ; width of head in ¢,
3} millim. (13 lin.), in 2 33 to 32 millim. (1? to 1} lin.); width
of front at vertex, in ¢ 3 millim. (2 lin.), in 2 just over
1 millim. (} lin.) ; length of proboscis (palpi) beyond margin of
buccal cavity, 23 millim. (1} lin.).

Thorax isabella colour,* with a narrow faint longitudinal stripe
on each side of the median line, dying away behind the transverse
suture before reaching the hind margin, and four sharply defined
small dark brown oval spots, arranged in a parallelogram, two in
front of and two behind the transverse suture ; abdomen ochraceous-
buff,t the longer hair at the base of the second segment entirely
golden yellow, the third to the sixth segments inclusive each with a
dark brown lunate mark on each side at the base, widely distant
from the median line, but not extending quite into the basal angle ;
ocellar spot dark brown ; proboscis bulb chrome yellow, with a sharply
defined dark brown or reddish brown tip.

Head.—Buff, posterior surface grayish ; the spots at the base
of the vertical bristles brown, but not united by a brown band ;
frontal stripe scarcely darker than the margins, the latter, as
well as the face and facial pit, faintly shimmering whitish-yellow.
Ocellar spot small, but conspicuous owing to its dark brown
colour ; ocelli small, the anterior ocellus of the same size as the
other two ocelli. Antennz: first two joints buff; second joint
sometimes more or less brown on the inside; third joint
brownish, the tip broader and with more prominent apex than
is usually the case in Gl. fusca; arista buff, shorter than in
Gl. fusca, second joint dark brown on under side only at extreme
base. Palpi buff-yellow, not darker above, the tips brown.
Bristles fringing the margin of the buccal cavity, below the
facial angle fine, numerous, and golden yellow.

* Ridgway, ‘‘ Nomenclature of Colors,” Pl. III. Fig. 23.
+ Ridgway, Ibid. Pl. V. Fig. 10.

102 DESCRIPTION OF GLOSSINA LONGIPENNIS:

Thorax.—Clothed with grayish dust, but when denuded some-
times appearing reddish on median area of dorsum; the two
dark brown spots in front of the suture slightly further apart
than the two behind it ; an additional pair of fainter and some-
what more reddish- bedien oval spots usually visible on the suture
itself, one on each side of the ad-median stripes, and between
them and the spots already mentioned ; in addition to these
marks there is usually a faint trace of a longitudinal stripe on
each side (representing the inner arm of the curved stripe, as
described for Gl. palpalis), between the, two dark, brown, spots
and the spot on the suture ; of indications of this stripe are ‘usually
confined to the neighbourhood, of the humeral callus and the hind
margin ; eed is also usually a tiny reddish-brown fleck i in front
of the suture, and between the two dark ‘brown oval spots on
each side; humeral callus with only a faint trace of a» brownish
blotch on its upper portion, Scutellum pale brownish, the
margin and, impressed median line buff.* |

Abdomen.—The dark brown: interrupted transverse bands, of
the type seen in Gil..morsitans, are here much _ reduced (see
Plate VII.); posterior angles of the third to the sixth segments
inclusive clothed with whitish-gray dust.

Legs.—Buff, femora with faint dark. patches on the outer
side, and middle femora with a brownish ,blotch on under side
near, tip (as in GI. fusca) ; only a yery faint, trace (often entirely
wanting) of a darker ring on the hind tibie near the base ;, last
joint and tip of penultimate joint of middle tarsi infuscated ; last
two joints of hind tarsi black. Row of bristles at base of ‘front
cox ochraceous ; hair clothing front coxze below golden-yellow.

Wings.—Brownish ; veins pale tawny, with local infuscations,
as in Gl. fusca. Squame not infuscated; fringe pale yellow,
silky. Halteres yellowish-white. ,

The type of Gil. longipennis, Corti, is a g in poor condition,
which has evidently been kept: in spirit prior to being pinned,
The above description has, therefore, been prepared from a ¢
from Somaliland (Th.:Greenfield), in the collection of the British
Museum, and a ? from West Somaliland, 23-25. vi. 1895 (C. V.
~ * Tn two specimens (one of which is the type of the species) out of the
seven examples of Gl. longipennis that I bave been able to examine there
is possibly a second dorso-central bristle, but if so it is so small as to be

practically indistinguishable ‘from the surrounding hairs; in none > of the
specimens can an intra-alar bristle be distinguished.

GLOSSINA LONGIPENNIS.—LOCALITIES. 103

A. Peel), in the collection of the Oxford Museum ; both of these
specimens have, been specially labelled. ;

i J) we )

Disratneirot oF Gi. longipennia, Corti.

Of this species I have been able to examine a ser ies ‘of seven
examples as follows :— ,

1 g (type of the species), River, Uelmal, Dane Galla
Country, N.E. Africa, June, 1893 (Capt. Vittorio Bottego), lent
by the Museo Civico di Storia Naturale di Genova, through the
courtesy of Dr. R. Gestro; 1 f°) (type, of the above ‘dexeription
of the. 2), West Somaliland, ‘93-25. vi. 1895 (C, V. A. Peel) (O.];
3 6.6, (including type of above re-description. of the By ighatts
Somaliland, precise. locality unknown (7h. Greenfield) ; 1 6)
Uganda Railway, British East Africa—? Kiboko River, or
Sabaki River near its junction with the Tsavo River, 1898
(Vet.-Capt. A. di Haslam).

The above list of | localities shows that the present species is
the Tsetse- -fly of Somaliland and the adjacent regions, but that
its range overlaps that of Gl. fusca, Walk., somewhere in the
vicinity of the Sabaki River. A field-note by Mr. Peel attached
to his specimen mentioned above says: ‘ Fly-belt sharply defined
from Biermuddo to Boholo Deno.”

Vai phd ms ~-t HABITS; ETC:

_. See the remarks quoted | above (pp. 99- 100) under Gl. _fusea,
from Capt. Haslam’s letter of 16. 4. 1898.

Tt will have been observed that, of the seven specimens of
Gl. ongipennis examined, all but two are males. One of the
male specimens collected and presented by Mr. Greenfield was
evidently taken in the act of sucking, since its abdomen is
swollen and distorted owing to its containing decomposed blood,
while there is a globule. of coagulated blood at the tip of the
proboscis. .

SYNONYMY AND AFFINITIES.

Glossina longipennis, Corti, is perhaps the best and most
easily characterised species of the genus. A year before the
species was actually described I had noted the specimens pre-
sented to the British Museum by Mr. Greenfield as a new species
allied to Gl. fusca, Walk. It is unfortunate that at the end of

104 GL. LONGIPENNIS: DISTINCTIVE CHARACTERS.

his original description Corti (loc. cit., p. 189) states that the
species is allied to Gl. tachinoides, Westw. (= Gl. palpalis, R.-
Desv., var. tachinoides, Westw.), to which of course it is in reality
by no means closely related. As already stated in dealing with
Gl. fusca, Walk., the latter species, with the one under dis-
cussion, “forms a group which, by reason of the considerably
larger size of the individuals, contrasts strongly with the other
species of the genus at present known.”

Apart from all other characters, Glossina longipennis is
readily distinguished from Gil. fusca by the ocellar spot being
dark brown and therefore very conspicuous, and by the presence
of the four dark brown oval spots on the dorsum of the thorax.
These dark brown spots are perhaps remnants of the dark
patches enclosed in the curved stripe in the complete scheme of
thoracic markings in the genus Glossina, as described above in
the case of Gl. palpalis ; but they are so much darker than the
remaining very faint and vestigial markings on the dorsum of
the thorax in Gl. longipennis that they appear as altogether
independent markings.

It must be observed that the wings in the present species are
not noticeably longer in proportion to the size of the body than
are those of any other species of Glossina. On the other hand,
the proboscis (i.e., palpi and proboscis) is remarkably short—
actually considerably shorter than in Gil. fusca, and relatively
shorter than in any other species of the genus,* so that brevirostris
would have been a much more appropriate designation. |

Lastly, it should be noted that in both sexes of Gl. longipennis
the front is considerably broader than in Gi. fusca,

* It says much for the discernment of the late Captain A. J. Haslam,
whose untimely death cannot be too greatly deplored, that he should have
noticed this in the field. As has already been mentioned, Captain Haslam,
while acting as Transport Officer to the Uganda Railway in 1898, collected
and forwarded to the British Museum specimens of Glossina pallidipes
fusca, and longipennis, and it is therefore to Gl. pallidipes and Gl.
longipennis that the following observation, taken from his letter of
16.4.1898, must be held to apply: “It is seen that the proboscis of the
smaller ones is larger in proportion to the body than the proboscis of the
larger flies is to their bodies.”

CHAPTER V.

THE MOUTH-PARTS OF GLOSSINA AND STOMOXYS.
(Plates VIII. and IX.).

By Dr. H. J. Hansen (Copenhagen).

The Glossina material investigated by me consisted of four
specimens belonging to three species, viz., a male and female of
Gl. pallidipes, Austen, and males of Gl. morsitans, Westw., and
Gl. fusca, Walk.* All the specimens were dried ; the heads I
boiled in water and examined as well as possible, after which
three of them were cleaned in a cold solution of caustic potash,
so that the chitin could be better studied. For comparison I
have also examined some spirit specimens of the common
European blood-sucking form Stomoxys calcitrans, L., and one
dried specimen of an African species of the same genus.t After
the examination of St. calcitrans I was able to find nearly all
the same muscles in the boiled specimens of Glossina, but it is
of course impossible to discover and study such organs as small
glands, nerves, etc., in specimens which have been preserved in a
dried condition. In the following pages I furnish a description
of the essential features of the external and internal mouth-parts,
and attempt to show how the animals bite and pump the blood
into the alimentary canal. Neither the very short time (nine
days) available for my investigation, nor the quality of the
material of Glossina, permitted me to make a more special study

* The labels attached to the specimens were as follows :—Gl. pallidipes,
Austen, 6: ‘‘Kilima-Njaro, F. J. Jackson”; Gl. pallidipes, Austen, ? :
“‘Witu Forest, 1895, Sultan of Witu, per W. S. Godfrey”; GJ. morsitans,
Westw. 6: ‘‘Beira Railway, Lower Pungwe River, Portuguese East
Africa, Oct. 1897, G. A. K. Marshall,—‘ Flew into railway carriage at
night attracted by light’”; Gl. fusca, Walk.: ‘‘ Witu Forest, 1895, Sultan
Witu, per W. S. Godfrey.”

+ This specimen bears the label; ‘‘Pemba Island, Hast Africa,
26, viii. 1899, D. R. O’Sullivan Beare.”

106 GLOSSINA.—PALPI AND PROBOSCIS.

of minute structural features, and a very detailed description
would scarcely be suitable for the present work.*

A. Externat Movuru-parts 1N Glossina.

I was unable to discover the slightest difference between
these parts in the male and female of Gl. pallidipes, and the two
other species did not present any difference from the first-named
form worth mentioning. |

The mawillary palpi (Fig. r , p)t are somewhat longer than the
height of the. head, auceemaalla projecting, with a deep longi-
tudinal groove on the side turned towards the plane of symmetry ; ;
they enclose the major part of the proboscis, of which only an
inflated basal portion (2) is seen on the under side of the head.
The proboscis is rather thick at the base, but nearly three-
fourths of its length are proportionately exceedingly slender ; it
reaches almost to. the end of the palpi, and is feebly curved, with
the coneayity beneath. It consists of, three parts (Fig. 2); the
upper lip or labrum (a), the hypopharyna (b), cae the lower lip or
labium (e).

-* So far as T am aware, the mouth-parts of Glossina have never been
investigated before. Both Glossina and Stomoxys are fairly near allies of
the European blow-flies (Cailiphora erythrocephala, Mg. and C. vomitoria,
L.), and the mouth-parts of Oalliphora have been carefully studied by
K. Kraepelin (Zeitschr. f. wissensch. Zool. 39. Ba. 1883, pp. 683-719,
Taf. XL.XLL) and B. T. Lowne (‘‘The Anatomy and Physiology of the
Blow-Fly.” . London, 1870); a second and greatly enlarged edition of
Lowne’s work was published in 1890-95: *The Anatomy, Physiology,
Morphology, and Development of the Blow-Fly (Calliphora erythro-
cephala).” By B. T. Lowne. 2 Vols. (London: R. H. Porter), 1890-95.
My description of the mouth-parts of Glossina and Stomorys is not so
detailed as the accounts by Lowne and Kraepelin of those of Calliphora,
and I disagree with these authors as to some of their interpretations of
muscles, etce.; but I have deemed it advisable to omit special references to
their statements as well as critical remarks on some of their opinions.
From earlier investigations I am well acquainted with the mouth-parts in
several families of Diptera (H. J. Hansen ; ‘Fabrica Oris Diptérorum, I.
(Tabanide, Bombylide, Asilid#, Thereva, Mydas, Apiocera);” Naturhist.
Tidsskrift, 3... Rekke, B. 14, 1884, pp. 1-220, Tab. 1.-Y,); ;—otherwise I
should not have ventured to examine ‘nothing’ but “dried specimens of
Glossina.—I may add that F. Meinert, in ‘his’ work entitled ‘ Trophi
Dipterorum” (4to, Kjgbenhavn, 1881), devotes two-thirds of a page and
three. figures, to. the mouth-parts of Stomorys;. this isthe only modern
note on the proboscis in that genus., Meinert ‘Observes ( (p. 68) that the
labella of Stomozys. have “‘ rows of teeth and chitinous blades,” which form
“‘a saw with several rows of strong, sharp teeth, with which this Dipteron
is able to saw a hole in the skin of Mammals;” but his drawing (Tab. V.
fig.,'24) of the structure in question is very incomplete (the strong teeth
are wanting) and incorrect.
orhithe sae ee to figures in ‘this ca Semin all. relate to.Plates VIII.
and IX.—H. E.

LABRUM,, HYPOPHARYNX,..AND.LABIUM. 107

The labrum (Fig. 2, a) is somewhat shorter than the labium,.
reaching to a point situated at the beginning of: the distal ninth
part of the last-named organ, It is narrow, though a little
broader towards, the base; in the natural condition the lateral
margins can be: seen to turn downwards, and .even a little.
inwards, so that the labrum forms about two-thirds of .a tube
(compare the diagrammatic transverse section, Fig. 6); the
lateral walls are flexible, the dorsal wall is a little thicker, In
Fig. 3 the labrum is shown from below and rather flattened ;. the
median part (p), of a darker tintin the figure, is the median
portion of the wall of the concavity on the ower side—the.
so-called. epipharyngeal plate—which is moderately thick ; no
muscle is found in the cavity between this plate and the dorsal
wall. Fig. 4 shows that this: thickening in the lower wall
terminates before the distal end of the labrum, which is in the
shape of a triangular and rather feebly chitinised plate. | In
Fig. 5, the basal part’is seen from below; the lateral margins (m)
are bent a little inwards; the small rings.(s) are apertures in
the epipharyngeal plate (p), and from each aperture proceeds a
hair which is exceedingly fine (so much so. that it was impossible
to show it in the figure) and proportionately long; somewhat
similar structures have been discovered in other Diptera, and
considered as organs of taste (compare the labrum of Stomoays).
On. the basal part these structures are numerous, but on the
remainder of the labrum there are very few of them (two. are
shown in Fig. 4)... The inner processes (Fig. 2, 7) from the base
of the labrum, and the muscles to its base and to the tall
are as in Stomowys (see eo

The hypopharynx (Fig. 2, b) is somewhat jonger mea ie
labrum, and a little shorter pte the labium, reaching slightly
beyond the base of the labella. - As is usually the case in Diptera,
it is: an exceedingly slender cylindrical tube, being: the outlet
from the thoracic salivary glands ; ; its’ distal eer is mem-
branous (Fig. 7).

The dabium (Fig. 2, c, and Fig. 12, e) is very interesting.
The basal fourth is thick and inflated; considerably broader ‘doch
deep, with a median, rather broad dorsal groove ; its distal
three-fourths are very slender, nearly equal in depth throughout,
and somewhat compressed. This slender part has a very deep
median dorsal groove ; in reality it is strongly depressed, and the
‘broad lateral thin parts are bent upwards anda little inwards
above, thus' forming. the:groove between them, or, more correctly,

108 TEETH ON LABELLA IN GLOSSINA.

a long tube is formed, with a slit on the upper side (compare
Fig. 8, which is a diagrammatic transverse section of the labium
near the middle). The labium consists of two segments and the
labella, but without a comparison with Stomoxys, Musca, and
Asilus, it would be impossible to discern these elements in
Glossina. The second segment and the labella (Fig. 2, e and /)
are together only about one-fifteenth of the length of the labium.
The sides of the labella, of the second segment, and of the most
distal part of the first segment are brownish and rather firmly
chitinised, so that the different sections are not easily distin-
guished one from another ; a narrow band at the upper margin
and an exceedingly short terminal portion of the labella are more
or less membranous ; the labella (Fig. 2, f) have a dark brown or
black area below just behind the cleft between them, and the
second segment has a pair of small, rather dark, oblique plates
on the under side (Fig. 2, e, and Fig. 9, b). Seen from the side
the labella are rounded at the end; seen from below (Fig. 9, c)
the cleft between them is sometimes narrow and triangular, and
sometimes the labella lie close together.

The inflated basal part of the first segment is somewhat
strongly chitinised on the sides and below, while the slender part
of the same segment is firmly chitinised below, with moderately
thin lateral walls. The bottom of the median groove is a rather
thick plate, which begins just below the base of the labrum, and
continues nearly to the end of the labium; in the most distal
portion of the first segment, in the second segment, and on the
proximal part of the inner side of the labella this plate is very
strong, and covers not only the bottom, but the sides of the deep
groove, almost to its upper margin. Fig. 10 represents the right
labellum seen from the inner side; a is the outer ventral wall
(the black spot) divided in the median line; c the plate in the
bottom of the groove divided in the same way; d is the- lateral
part of this plate, and e its incurved upper margin. The same
tigure shows that the inner wall of the free part of the labellum
is sheathed with three plates (q), which are strongly chitinised,
and over the major portion of their surface adorned with a
considerable number of transverse rows of exceedingly small
teeth, thus presenting a certain resemblance toarasp. All these
plates are prolongations of the plate in the groove, but the upper
and lower ones are narrowed basally and marked off at the
base by a suture allowing a slight degree of mobility. The
lateral margins of all three plates are finely serrated (the number

LABELLA IN GLOSSINA. 109

of the teeth being identical with that of the transverse rows of
teeth), and lie close together. The upper distal angle of the
upper plate and the lower distal angle of the lower plate are
each produced into a triangular tooth (hk and i); at the distal
end of the plates are firmly inserted four very large teeth (k),
two on the median and one on each of the other plates, and these
six teeth are directed towards the distal end of the labella. The
skin between the insertion of the submedian teeth and the end
of the labium is membranous and scaly (/), and at the upper
margin bears a number of short, stiff setee (m) instead of scales.

I am inclined to think that a portion of membranous skin
has been invaginated from the end of the labella, and that in
material preserved in spirit, or at any rate in fresh specimens,
the labella would sometimes be found with the membrane
alluded to evaginated and easily visible. This assumption is
based on the fact that with transmitted light four very long
chitinous rods are easily seen between the outer and the inner
walls of the terminal portion of the labellum; in Fig. 11 this
portion of the right labellum has been drawn as seen from the
outer side, and the rods are indicated by dotted lines (d). Each
rod is very long, and neariy cylindrical, with the distal end
rounded, and is articulated to a somewhat short and rather
thicker cylindrical foot. I am unable to decide as to what the
function of these rods may be; they are hollow, but I could not
discover any aperture at the end, and if they were ducts from
glands a distal aperture must exist. I am therefore most
inclined to believe that these rods are sense organs, They must
be inserted in membranous skin, which is invaginated in the
dried specimens seen by me.

The outer side of the distal half of the labella exhibits a
small number of holes in which sete have been inserted ; in two
of my species these sete had been broken off near the base, but
in Gl. fusca they were present as moderately short stiff spines,
directed outwards and partly backwards. In the last-named
species I have also found four very small, clear rods arranged
along the distal end of the labellum ; these rods are certainly
sense organs.

In Tabanus, Asilus, and Calliphora the labium contains three
pairs of muscles: in Glcssina I could distinguish two pairs with
absolute certainty, and they fill up the basal inflated portion of
the organ. One pair proceeds from the sides of the labium in
an oblique direction and forwards to the margin of the plate

110 STOMOXYS.—PALPI AND PROBOSCIS.

situated in the bottom of the groove. The'second pair is very
large ; each of its muscles terminates in a tendon, which is very
strong and runs from the distal end of the inflated portion to
the outer wall of the labellum. I am unable to decide whether
the slender portion of the first-segment of the labium contains a
third pair of muscles, like those in Stomoays, described later on.

“The hypopharynx is firmly united, or one might say fused,
with the proximal end of the plate in the groove of the labium,
and ‘a little more distally it is connected sails the. base of the
lower wall of the labrum (compare. Fig. 22 of Stomoxys). The
hypopharynx being a direct prolongation of the posterior wall,
and the epipharyngeal plate (the lower side of the labrum) the
direct prolongation of the anterior wall of the pharynx, the
opening of the mouth may be said to be situated just at the
connection between the hy popharynx and the labrum.

' B, Exrernat Movrn-parts 1n Stomoxys.

)

There is no difference between these organs in the male and
female ; furthermore, the differences observed between the
mouth-parts in St. calcitrans and the African species are slight.
The following description is based essentially on St. calcitrans.
The proboscis is somewhat longer than the height of the head,
horizontally projecting’ (Fig. 15), but not covered by the palpi,
which are very slender, and cylindrical, and in the species
examined by me are scarcely one-third or not 4 quarter as long
as the proboscis. This consists of the same parts as in Glossina.
The labrum (Fig. 16, a) reaches nearly to the base of the
labella. It is more strongly chitinised than in Glossina and
almost of thé same shape (Fig. 17), but the lateral walls are more
incurved below, so that the labrum constitutes a tube with a
moderately broad slit along the lower side. . The apical portion
(Fig. 18) is oblong triangular, subacute, and rather firmly chiti-
nised,. Not far from the end some sense organs (Fig. 18, s) are
observed ; in the sub-basal portion of the labrum such organs are
rather numerous, and some are found in the intermediate portion.
Each organ bears a moderately short and clear hair, and the
ganglion at its base is often easily observed. To. each of the
lateral basal angles is firmly attached a strong chitinous rod or
apodeme (Fig. 17, b), which is rather long and proceeds upwards
in the head (Fig. 21, h); to its free end is attached a fairly
strong mus¢le, which runs.to the distal solid end of, the pharynx

TEETH ON LABELLA. Lil

(Fig. 21, m?; Fig. 17, c), and by the contraction of this pair of
muscles the distal end of the labrum is raised. From the base
of the upper wall of the labrum a muscle (Fig. 17, e; Fig. 21, m)
runs upwards to the*frontal part of the lateral wings of the
pharynx ; the contraction of this muscle will press the distal part
of the labrum downwards. For the secondary results of the
contraction of these muscles, and as to a third pair of muscles
(Fig. 17, d; Fig. 21, m?) attached near the base of the above-
mentioned apodemes the reader is referred to the section on the
internal mouth-organs.

The hypopharyna (Fig. 16, 6) is as long as the labrum ; it is a
thin cylindrical. tube, with the apical part membranous (Fig. 19),
as in Glossina.

The labium (Fig. 16, c) differs conaldordblly from that’ of
Glossina. Nearly the whole of the basal half is somewhat
swollen, but to a much lesser degree than in the other genus,
and its distal portion is considerably less slender ; the dorsal
groove is in the distal half proportionately less deep than in
Glossina. The labella are slightly higher than the second and
than the distal part of the first segment; the labella and the
second segment together occupy about one-sixth of the length
of the whole organ. The labella have on their outer side near
the end, and at the lower margin, a row of partly rather long,
partly short sete. The inner side of the labella is exceedingly
interesting (Fig. 20). As. in Glossina, ‘the plate (c) in the
groove is distally very strong, and occupies not only the bottom
but the major part of the sided: ; its end is obliquely truncated,
and on each labellum a very high plate (d) is attached to the
end by a well-developed articulation. From the basal part of
this labellar plate five enormous teeth (e) proceed forwards and
downwards. These teeth, which are arranged in a curved row,
are firmly anchylosed to the plate, very stout, a little more to a
little less than two and a half times longer than broad, distally
triangular and subacute. The three middle ones are distally
irregularly serrated along both margins, and possess in addition
small secondary processes. near one margin; the upper tooth is
serrated distally along the upper and the lower tooth along the
lower margin. The area between these teeth and the membranous
distal: margin of the labellum is occupied by two rows of large,
elongate, rather thin and very sharp chitinous blades (/), of
about the same shape as the blade of a knife, and directed the
same” way as: ‘the: large teeth.- A little-way inside the’ distal

112 ACTION OF MOUTH-PARTS.

margin of the labellum a number of hairs are inserted, some of
which at least are branched. Below the terminal portion of
each of the big teeth one or two dark-coloured, rather long
chitinous rods (g) are inserted ; they are sub-cylindrical, slightly
curved, and hollow, and seem to possess an aperture on the
distal obtuse end.

The labium contains three pairs of muscles. The first pair
is lamellar, with short fibres, and proceeds from the outer lateral
wall obliquely to the margin of the plate in the bottom of the
groove ; these muscles occupy the greater part of the length of
the labium. The second pair is very large, filling most of the
lumen of the basal third of the labium; each muscle terminates
in a very strong tendon, which is attached to the outer wall of
the labellum. The third pair is, so far as I was able to see,
very different from that in Calliphora and Tabanus ; the fibres
are very short, proceeding from the ventral side of the plate in
the groove obliquely forwards and downwards to the ventral
wall of the first segment, at least in its distal half.

The hypopharynx, labrum, and labium are attached to each
other at their base much as in Glossina ; but the attachment of
the hypopharynx to the labium seems to be less solid than in
that genus.

C. Mope or Action oF THE ExtERNAL MOUTH-PARTS IN
Glossina AND Stomoxys.

In both genera, the tube through which the blood of the
animal on which the fly may be feeding passes to the pharynx,
is constituted by the labium and the labrum together. The
labrum lies within the deep dorsal groove of the labium, closing
it above and also forming the inner lateral walls of the tube ;
beyond the end of the labrum the sub-membranous margins of
the labium touch or perhaps cover each other above; in specimens
of Stomoxys preserved in spirit, I have seen the margins of the
labium in contact for half the length of the proboscis. The
hypopharynx is completely enclosed in the tube mentioned. In
both genera a short terminal portion of the hypopharynx is
entirely membranous, and this organ is consequently useless for
the stabbing of victims; the labrum is somewhat shorter than
the labium, and in Glossina at any rate its extremity is not
strong enough for piercing. The insects belonging to both
genera perform this operation by means of the armature of the

PIERCING ORGANS. 113

inner side of the labella, and the extremely strong muscles, the
tendons of which are attached to the outer wall of the labella,
must be the chief factors in the necessary movements. By the
contraction of these muscles the inner sides of the labella are
turned so much outwards that their teeth are able to act upon
the skin of the victim; I am unable to decide whether the
muscles described above as the third pair in the labium of
Stomowys can assist by pushing the plate in the groove outwards,
and thereby affecting the plates on the inner side of the labella
ina similar manner. In Glossina the teeth are inserted rather
near the end of the labella, and it must be these six pairs of
teeth which are at least the chief instruments in piercing a hole.
But I am bound to confess that I can form no opinion as to the
significance of the three pairs of rasp-shaped plates—whether
they assist during the piercing or are useful in another respect.
In Stomoxys the five strong. teeth are situated near the base of
the labella ; but in this insect the labella are more movable, and
can be turned outwards so that the teeth can act on the skin of
a victim.* The blades in the two rows are very sharp, but
rather thin ; they probably have an accessory function in the act
of piercing.

It is, in my opinion, highly probable that the hole produced
by Stomoxys is not very deep, the labella being proportionately
rather thick, and the essential teeth being placed so far from the
end that the labella must be greatly shortened and much turned
outwards when the fly bites. But in Glossina the labella are
very slender, and more strongly chitinised on the outer side, with
their teeth near the end, and it is therefore probable that the
proboscis can be sunk rather deeply in thick skin. The saliva is
introduced into the wound from the end of the hypopharynx,
and—as in the case of mosquitoes (Culex, Anopheles)—is certainly
mixed with the blood which is to be pumped through the
proboscis.

D. Lower Portion oF THE HEAD In Botu GENERA.

As in the blow-fly and the common house-fly, the lower part of
the head, between the outer mouth-parts and the firmly chitinised
portion of the head, is completely membranous and can be forced

* In a specimen of Stomoxys preserved in spirit I found the labella
turned outwards to such an extent that the big teeth occupied their distal
extremity, and were directed almost vertically to the. sides, and only
slightly downwards.

I

114 INTERNAL MOUTH-PARTS.

downwards so as to constitute a cone, with the outer mouth-parts
on the end (Fig. 16): this cone is evidently somewhat longer in
Stomoxys than in Glossina. Fig. 12 represents the median part
of the posterior side of the head and the labium (e) in Glossina,
with the long membrane (d) between both; ¢ is the occipital
foramen, a the firmly chitinised part. It will be seen that there
is a deep median incision, which is membranous, and surrounded
at the sides by the firm chitin which borders the eyes below ; the
labium can be pulled upwards, so that its basal part occupies this
median area, the membrane being invaginated and folded. On
the front side of the head the epistoma, which is the median area
below the transverse prominence under the antenn, is mem-
branous in Glossina, with the exception of an upper transverse
area, which is coalesced with the wings of the pharynx (see
below). In Stomoxys the epistoma shows essentially the same
structure, but the sub-basal transverse solid chitin is short, and
from it two parallel dark strips run downwards (Fig. 16, 7) nearly
to the base of the maxillary palpi. The muscles which effect the
protrusion and retraction of the membranous cone are mentioned
below.. The maxillary palpi (Fig. 16, g) are inserted on the
anterior side of the cone, a little below its upper margin. No
other vestige of the maxille and no rudiment of mandibles can
be discovered ; but in Stomoxys I found at the base of each palpus
a very small curved chitinous strip (Fig. 16, kh), movably united
with the palpus. Whether this strip is a rudiment of the maxilla
I cannot say.

E. Tre Internat Movru-parts oF Glossina anv Stomoxys, AND
THEIR Mopr or Acrion.

From the tube constituted by the external mouth-parts the
blood is conducted into the pharynx, which is a very large and
complicated organ, In the natural position and seen from the
side (Fig. 21) it is nearly vertical, but slopes slightly forwards.
It is obliquely triangular, with the two upper posterior lateral
angles produced into a moderately short (Glossina, Fig. 2, m) or a
very long process (Stomoxys, Fig. 21); in Stomoays the two upper
anterior angles are also produced. Fig. 13 represents the pharynx
of Glossina seen from behind, together with the basal part of the
cesophagus (d), and a small anterior separate part (c) between
the pharynx itself and the external mouth-parts. Fig. 14 isa
diagrammatic section of the same pharynx at its thickest point :

SUCTORIAL ACTION. 115

it will be seen that posteriorly it has two walls, with a narrow—
or in the resting position with no—space between them, and the
anterior of these walls is the thinner. The lateral wall (c) is
thick, and anteriorly coalesced with the thickened part of the
epistoma (d). In reality the pharynx is a part of the cesophagus,
which has been firmly chitinised, widened, and strongly depressed,
with the anterior wall slightly concave and the lateral margins
produced into very broad wings (b), which turn forward and
coalesce with the epistoma. The large space between the lateral
wings, the anterior of the two posterior walls and the firm part
of the epistoma is nearly filled with enormous muscles (m) ; in
the distal part of the pharynx the fibres run very obliquely
upwards and forwards to the epistoma (Fig. 21); in the upper
part of the pharynx they run horizontally, or in front even a
little downwards (Fig. 21). The contraction of these muscles
produces a space between the two posterior walls of the pharynx,
and the whole organ is a pumping-engine. It must be supposed
that the distal fibres of the muscle contract, raising the corre-
sponding part of the front wall, and thus producing a space void
of air into which the fluid is sucked from the tube formed by the
external mouth-parts. Next, the more proximal fibres contract,
while the most distal ones relax, and the result of a successive
contraction of all the fibres from below upwards, the lower oncs
relaxing simultaneously with the contraction of the upper ones,
and so on, will force the blood of the victim upwards through
the pharynx into the cesophagus itself, and also produce a suction
from the wound at the end of the labella. The cesophagus itself
(Fig. 21,7) is a membranous tube (having in Glossina a small
sub-basal dorsal plate, Fig. 13, e), which is at first as broad as
the pharynx (Fig. 13), then narrower, and is bent backwards in
a curve to the occipital foramen (Fig. 21). Between the lower
end of the solid pharynx and the attachment of the labrum to
the hypopharynx is found a tube, which in Glossina is rather
short and wide, firmly chitinised in the posterior half (Fig. 13, ¢)
and membranous in front. In Stomowys it is longer and narrower,
with irregular sub-circular linear thickenings in the wall
(Fig. 22, 9).

From the postero-superior process of the pharynx a strong
muscle (Fig. 21, m7) proceeds downwards to the cheek at the side
of the epistoma. The thickened part of the epistoma, with which
the wings of the pharynx are coalesced, is united with the solid
chitin below the antennz by a narrow transverse membranous

12

116 PROBOSCIS AND PHARYNX—MUSCULATURE.,

strip. ,When the pair of muscles referred to (m") are contracted,
this strip functions as a hinge around which the pharynx moves
in a downward direction, and the result is that the lower end of
the pharynx, which by its apical tube (Fig. 21, &). is united with
the base of the external mouth-parts, is forced downwards, pulling
the external mouth-parts in the same.direction and extending the
membranous cone between these mouth-parts and. the firm chitin
of the head.. The cone is retracted by at least three pairs. of
muscles. One pair (Fig. 21, m°) is inserted. on its posterior wall
below the firm chitin, and proceeds upwards to .a spot outside
the occipital foramen ; the second pair of muscles (m*) proceeds
upwards from the proximal end of the lower wall of the labium
to the firm chitin on the posterior side of the head; the third
pair (m) runs from the sub-apical part of the apodemes of the
labium upwards and backwards to the posterior side.of the head
outside the occipital foramen. Probably a fourth muscle
(Fig. 21, m’), proceeding from the proximal end of the upper
wall of the labrum to the anterior, margin of the pharyngeal
wings, may also act as a retractor of the membranous cone, and
also as a depressor of the labrum, as mentioned above. That
the three first-mentioned pairs can act as retractors of the cone
is certain, but I must confess that I do not. know whether they,
or at least some of them, have some other function in addition.
The muscles (Fig. 21, m*) from the free end of the apodemes of
the labrum to the lower end of the pharynx can certainly not
only act as mentioned above, viz. raise the distal end. of, the
labrum, but also assist (m") by the evagination of the membranous
cone. iti ) ; Us esas sad
It has been mentioned above that the hypopharynx forms -the
outlet for the saliva from the glands in the thorax. The hypo-
pharynx (Fig. 22,.c) is perforated near its base on the under
side, and from this aperture begins the free salivary duct (h),
which runs upwards along the posterior side of the most distal
part of the pharynx, and then (¢) turns in a more backward
direction to the occipital foramen. Just behind the sub-apical
solid part of the pharynx the duct is dilated, and in the normal
condition depressed into a sort of oval disk, with the anterior
wall a little concave and the posterior convex. (In the specimen
drawn in Fig. 22 this organ, k, was unusually thick, the anterior
wall being raised.) From the anterior wall proceeds a pair of
very thin and long muscles (Fig. 22, m; Fig. 21, m%), the other
extremity of which is attached to the posterior surface of the

EXPLANATION OF PLATES VIII. AND IX. 117

pharynx near its upper end. The function of the dilatation of
the salivary duct would seem to be to retain the salivary secretion.
When the muscles referred to contract the anterior and posterior
walls are separated (as in Fig. 22), and the passage is open. I
‘can scarcely think that this organ acts as a pumping mechanism
or syringe. , |

In this chapter all essential chitinous structural features and
all the muscles belonging to the mouth-parts have been mentioned,
so that the main points of the functions of the different organs
can be understood. As already pointed out, the labella of Glossina
especially need further study (with high magnifying power),
which should be carried out on fresh material, or at least on
well-preserved spirit specimens. Moreover, the depth to which
these flies bury the proboscis in the skin of their victims should
be observed, as also the time occupied in biting, etc.*

EXPLANATION OF THE Figures (Puares VIII. ano IX.).

The camera lucida was used for all the drawings, with the
exception of the three diagrammatic sections of mouth-parts in
Glossina. Some of the figures have been drawn as seen by
transmitted light, and in these the thin-skinned parts are light-
greyish, the solid parts dark ; when a figure has been drawn in
this way, the fact’ is specially mentioned below. The other
figures (the diagrams excepted) have been shaded in accordance
with the shape of the parts: the membranous parts are light
greyish, while on the firm chitin dark shading is opposed to
white.

Fia. 1.—Head of Glossina morsitans, Westw., 3, X11: J, basal inflated
part of the labium; p, palpus.

Fie. 2.—External mouth-parts and pharynx of Glossina morsitans,
Westw., ¢,seen from the side, x rather less than 21; a, labrum; b, hypo-
pharynx; c, labium; d, its first segment; e, its second segment; f,
labellum ; g, basal part of the left maxillary palpus; h, anterior wall of
the membranous cone; i, left apodeme of the labrum: k, posterior wall
of the membranous cone; 1, pharynx; m, its upper process; 7, solid part

* According to M. Edouard Foa [135], when the Tsetse bites the
proboscis ‘disappears completely in the flesh,” while Lieut.-Col. Bruce
writes (vide Chapter VII., Appendix A, p. 274) :—‘‘The act of feeding is
remarkably quick. From the moment of settling on the animal until
the <: vi fully blown out with blood is often as little as 20 to 30 seconds,”

118 EXPLANATION OF. PLATES VIII. AND IX.

of the epistoma coalesced with the pharynx; 0, tube between the pharynx
itself and the labrum-hypopharynx; p, dilatation on the salivary duct;
q, muscle attached to this dilatation; 7, upper part of the salivary duct.

Fia. 8.—Labrum of Glossina pallidipes, Austen, ?, seen from below
and drawn by transmitted light, in a rather flattened condition; x 14:
p, the plate in the lower wall.

Fia. 4.—Distal part of the labrum shown in fig. 3, seen from below
and drawn by transmitted light; x 46: p, the plate in the lower wall;
8, sensory organ,

Fia. 5.—Basal part of the labrum shown in fig. 3, seen from below ;
x 46: m, marginal part turned a little inwards; p, plate in the lower
wall; s, sensory organ,

Fic, 6,—Diagrammatic section through the middle of the labrum of
Glossina ; p, plate in the lower wall,

Fic. 7.—Distal part of the hypopharynx of Glossina pallidipes,
Austen, 9; x 100: a, membranous apical portion,

Fia, 8,—Diagrammatic section through the middle of the labium of
Glossina ; p, plate in the wall of the groove.

Fia. 9.—Distal part of the labium of Glossina pallidipes, Austen, ?,
seen from below; x 100; a, terminal part of first segment; b, plate of the
second segment; c, labella,

Fia. 10,—Right labellum of Glossina pallidipes, Austen, 9, from the
inner side; x 890; a, ventral outer wall, divided longitudinally; b, space
between the outer ventral wall and the plate in the groove; c, the plate in
the groove divided longitudinally; d, lateral part of this plate; e, upper
part of the same plate bent a little inwards; f, upper sub-membranous
margin of the labellum; g, the rasp-like plates; h, tooth on the distal
upper margin of the upper rasp-like plate; 7, tooth on the distal lower
margin of the lower rasp-like plate; k, the large teeth, anchylosed to the
end of the three plates; 1, membranous scaly skin; m, setee on the mem-
branous skin ; 2, terminal portion of two of the long chitinous rods (? sense-
organs),

Fia. 11.—Distal part of the labellum shown in the preceding figure,
and seen from the outer side; x 890: a, lower margin; b, upper mem-
branous margin, with rows of exceedingly small spinules ; c, membranous
skin; d, long chitinous rods, seen through the outer wall; e, insertions of
hairs or spines, which have been broken off.

Fia, 12.Lower median part of the posterior side of the head of
Glossina pallidipes, Austen, ?, with the labium in an exserted condition,
seen from behind; x not quite 12: a, solid chitin on the posterior side of
the head ; b, solid chitin of the cheek; c, occipital foramen; d, membrane
between the head and the labium; e, labium.

Fig, 18.—Pharynx of Glossina pallidipes, Austen, ¢, from behind ;
x 19: a, posterior wall of the pharynx; b, one of its lateral wings; c, plate
in the portion between the pharynx itself and the hypopharynx; d, basal
part of the esophagus; e, small plate in the upper wall of the cesophagus.

EXPLANATION OF PLATES VIII. AND IX. 119

Fia. 14.—Diagrammatic transverse section through the pharynx of
Glossina ; a, posterior wall of the pharynx; b, its anterior wall; c, its
lateral wing; d, solid plate in the epistoma; e, pumping muscles,

Fira, 15.—Hoad of Stomowys sp., ?, from Pemba Island, Bast Africa;
x 12: J, labium; p, maxillary palpus.

Fig. 16,—Outer mouth-parts, in an exserted condition, of Stomoxys
calcitrans, Li, 6 (from Denmark); x 19: a, labrum; 6, hypopharynx ;
c, labium; d, exserted membranous cone between the outer mouth-parts
and the solid chitin of the head; e, apodeme of the labrum seen through
the skin; f, firm chitin of the head; g, maxillary palpus; h, very small
solid plate at the base of the palpus (? rudiment of the maxilla); 7, solid
vertical sub-median strip in the epistoma,

Fria, 17,—Labrum (with its apodemes) of Slomoxys caleitrans, L., ¢,
from below; x 19: a, median longitudinal groove; 6, strong chitinous
rod, or apodeme; c, muscle from the free end of this apodeme, to the
lower end of the solid pharynx; d, muscle from the apodeme to the
posterior side of the head; e, muscle from the base of the upper wall of
the pharynx to the anterior margin of the pharyngeal wings. (‘The entire
length of the two last-named pairs of muscles is not drawn.)

Fia. 18.—Distal part of the labrum shown in the preceding figure,
from below; x 120: a, lower margin of the labrum turned inwards;
b, the median groove; ¢, sensory organ,

Fia, 19.—Distal part of the hypopharynx of Stomowys caleitrans, L.,
é, from the side; x 120: a, membranous apical portion.

Fia. 20,—Right labellum of Stomoays caleitrans, L., 6, from the inner
side: x 280: a, lower wall divided longitudinally; 6, sub-membranous
upper margin; c, the plate in the bottom and on the sides of the median
groove; d, plate on the inner side of the labellum; e, very strong teeth
anchylosed to the plate of the labellum; /, elongate chitinous blades ;
g, chitinous rods; h, sete on the outer side near the terminal margin,

Fira, 21.—Vertical section, a little to one side of the plane of symmetry,
through the lower part of the head of Stomowys calcitrans, L., 6 5 x 27;
slightly diagrammatic: a, basal part of the proboscis, from the outer side ;
b, anterior wall of the membranous cone; c, maxillary palpus; d, oblique
membranous part of the epistoma; ¢, section through the solid chitin just
outside the epistoma; /, posterior wall of the membranous cone ; 7, section
through the solid chitin outside the occipital foramen: h, left apodeme of
the labrum; i, pharynx; /, tube from the lower ond of the solid pharynx
to the labrum-hypopharynx; /, esophagus; m', muscle from the upper
wall of the labrum to the anterior margin of the pharyngeal wing;
m?, muscle from the apodeme of the labrum to the posterior side of the
head; m*, muscle from the free end of the apodeme of the labrum to the
lower ond of the pharynx itself; m*, muscle from the proximal end of the
labium to the posterior side of the head (the upper part of this muscle
omitted); m*, muscle from the hind wall of the membranous cone to the
posterior side of the head outside the occipital foramen; m", pumping
muscle; m’, muscle from the posterior process of the pharynx to the solid
chitin just outside the epistoma; m*, muscle from the dilatation of the

120 EXPLANATION OF PLATES VIII. AND IX.

salivary duct to the upper part of the hind wall of the pharynx; », lower
part of the salivary duct; 0, upper part of the same.

Fia. 22.Basal portion of external and distal portion of some of the
internal mouth-parts of Stomoxys calcitrans, L., 6, drawn by transmitted
light; x 54: a, labrum; 3, basal part of its left apodeme; c, basal part of
the hypopharynx; d, base of hypopharynx and of the plate in the groove
of the labrum; e, small part of the base of the plate in the labial groove ;
f, lower part of the pharynx itself; g, tube between the pharynx itself and
labrum-hypopharynx; h, lower part of the salivary duct (the wall is thick);
4, upper part of the same duct (the wall is thin, with linear circular
thickenings); %, dilatation of the same duct; m, terminal part of its left
muscle.

COPENHAGEN,
August 26, 1902.

121

CHAPTER VI.

BIBLIOGRAPHY.—PART I.
THE LITERATURE OF TSETSE-FLIES.

[In order to render the following Bibliography more useful
to readers who happen to be specially interested in a particular
region of Africa, the records of observations and occurrences are
here shown classified under four main geographical headings :—

A. West Africa.
53, 56, 65, 654, 80, 95, 99, 107, 122, 146, 148, 157, 162, 170, 171.
See also Chapter VII., Appendix H, p. 310.

B. East Africa (including Somaliland, and German and British Hast
Africa).

I, 12, 25, 28, 43, 45, 47, 49, 54, 60, 62, 66, 71, 78, 79, 81, 94, IOI,
102, 115, II7, I19, 120, 124, 126, 127, 133, 139, 144, I50, 151,

154, 155, 163, 164, 166, 167, 168, 169, 171, 174, XXVi.
See also Chapter VII., Appendix ©, pp, 291-294, 296-297.

C. Central Africa (including the Valley of the Upper Nile, Uganda, and
Nyasaland).
254, 29, 37, 59, 61, 98, III, 125, 138, 141, 145, 172.
‘See also Chapter VII., Appendix B, and Appendix C, p. 295.

D. South Africa (including the Zambesi Valley, and the country to the
south),
5 9, Il, 20, 2i, 23, 26, 27; 28, 30, 36, 38, 40, 4i, 42, 50, 52, 53, 55) ‘

57, 64, 67, 69, 70, 72, 73, 74, 75, 79, 77, 85, 86, 93, 97, 98, 100,
104, 105, 105A, 106, 108, 110, 112, 114, 116, 118, 121, 128, 134,

135, 143, 149, 152, 153, 159, 160, 161, 165, 173.
See also Chapter VII., Appendix A, and Appendix C, p. 290.]

1. 1813. James Bruce.
“ TRAVELS TO DiscoveR THE SouRcE oF THE NILE”
3rd ed., 8vo., Vol. II., pp. 305-307 ; Vol. VII., pp. 300-—
303, 305 ; figure of the “ Tsalisalya or Fly,” 4to ed. (1813),
Vol. VII. (VIII.), No. 39.

1

2

“TSALTSALYA” OR “ZIMB” OF J. BRUCE.

“ Tsaltsalya or Fly.”—The fly, of which “'Tsaltsalya ”
is the “ Ethiopic” name, and “the true name of this par-
ticular fly in Geez,” is stated to be confined to a district
of “ black fat earth,” called Mazaga, on the Upper Atbara
River, on the confines of the Sudan and Abyssinia.
During the rains the cattle had to be taken down to the
sandy plains of the Lower Atbara, to escape the fly
(8vo ed. (1813), Vol. VII., pp. 300-303).

The figure of the Tsaltsalya is poor, and represents an
insect with hairy body and legs, but it is apparently
meant either for a Pangonia (Family Tabanide) or a
Tsetse.

“The insect is called Zimb, in modern or vulgar
Arabic ; it has not been described by any naturalist. It
is in size very little larger than a bee, of a thicker pro-
portion, and the wings, which are broader than those of a
bee, are placed separate like those of a fly; they are of
pure gauze, without colour or spot upon them ; the head
is large, the upper jaw or lip is sharp, and has at the end
of it a strong pointed hair of about a quarter of an inch
long ; the lower jaw has two of these pointed hairs, and
this pencil of hairs, when joined together, makes a resist-
ance to the finger nearly equal to that of a strong hog’s
bristle. Its legs are serrated in the inside; and the
whole covered with brown hair, or down. As soon as this
plague appears, and its buzzing is heard, all the cattle
forsake their food, and run wildly about the plain, till
they die, worn out with fatigue, fright, and hunger. No
remedy remains, but to leave the black earth, and to
hasten down to the sands of Atbara; and there they
remain while the rains last, this cruel enemy never daring
to pursue them farther ” (8vo ed., Vol. VII., p. 305).

[The description of. the size of the insect—“ very
little larger than a bee, of a thicker proportion,” with
wings “ broader than those of a bee,” and placed separate
like those of a fly”—is suggestive of Pangonia rather
than Glossina. |

“All the inhabitants of the sea-coast of Melinda,
down to Cape Gardefan [sic], Saba, and the south coast
of the Red Sea, are obliged to put themselves in motion,
and change their habitation to the next sand in the
beginning of the rainy season, to prevent all their stock

EARLY DESCRIPTIVE AUTHORS. 123

of cattle from being destroyed. This is not a partial
emigration ; the inhabitants of all the countries from the
mountains of Abyssinia northward, to the confluence of
the Nile and Astaboras [Atbara], are once a year obliged
to change their abode, and seek protection in the sands of
Beja; nor is there any alternative, or means of avoiding
it, though a hostile band was in their way, capable of
spoiling them of half their substance. . . .” (8vo ed.,
Vol. II., pp. 306-307).

[Bruce considers that this is the fly referred to in the
following passage in Isaiah, vii. 18 and 19:—‘ And it
shall come to pass, in that day, that the Lord shall hiss
for the fly that is in the uttermost part of the rivers of
Egypt, and for the bee that is in the land of Assyria.
And they shall come, and shall rest all of them in the
desolate vallies, and in the holes of the rocks, and upon
all thorns, and upon all bushes.”

Westwood believes Bruce’s “‘Zimb ” to be the Tsetse,
“or at least that that insect is a larger species of Glossina,
to whose real habits Bruce has added those of a species
of Cstrus” (P. Z. S., 1850, p. 262). If Westwood is
right it follows that the range of the Tsetse extends to the
Sudan, on the confines of Abyssinia. |

2. 1830. C. R. W. Wiedemann.
“ AUSSEREUROPAISCHE ZWEIFLUGELIGE INSEKTEN,”
Zweiter Theil: pp. 253-254, Taf. IX., Figs. 10*, 10°, 10°.
Original description of the genus Glossina, and of Gl.
longipalpis.

3. 1830. J. B. Robineau-Desvoidy.
“Essa sur Les Myopaires” (Mémoires présentés par
divers Savans « Académie Royale des Sciences de 0 Institut
de France. Tome Deuxiéme), pp. 389-390.
Original description of the genus Nemorhina, and of

N. (Glossina) palpalis.

4. 1835. J. Macquart.
‘‘Historre NavrureLLe pes Insectes. Dipréres” :
Tome Deuxiéme, pp. 244-245, Pl. 16, Fig. 8.
Description of the genus Glossina, and of Gl. longipalpis
(apparently the true Gl. longipalpis, Wied.). The figure
is a caricature.

‘124 TSETSE ENCOUNTERED BY HARRIS.

5. 1839. Captain (afterwards Sir) William Cornwallis Harris.
‘Tue Witp Sports oF SourHEerN Arrica ” (London :
John Murray), p. 231.

A passage referring to the occurrence of Tsetse in the
Mural Berge—a range of hills in the Waterberg District
of the present Transvaal, on the south bank of the Lim-
popo, close to the intersection of the 27th parallel E.
longitude and the Tropic of Capricorn—in November, 1836.
Although not mentioned by name, it is very evident that
the Tsetse is the fly alluded to, The author writes :—“ Here
the country again assumes a more. level character, but is
broken to the eastward by detached hills and low ridges,
imperceptibly increasing in importance, until they grow
into a great range of mountains, known to the natives as
the Mural. . . . During the rainy season especially, they are
infested by a large species of gad-fly, nearly the size of a
honey-bee, the bite of which, like that of a similar pest in
Abyssinia; proves fatal to cattle. A desire to escape the
officious visits of ‘these destructive insects, whose persecu-
tions relieved us of two of our oxen, soon obliged us to
abandon the willow-fringed river, which threads the
mountains for a considerable distance. . . .”

In the map at the end of the volume, entitled “ Africa
North-East of the Cape Colony, exhibiting the relative
positions of the Emigrant Farmers and the Native Tribes,
May, 1837,” the position of the Mural Berge is not quite
correctly indicated. They are there called the “ Murat
[sic] Mount»s,” and are marked as “abounding in flies
destructive to cattle.”

6. 1843. J. Macquart.

“Diprkres Exotiqurs Nouveaux ou Pru Connus”:
Tome Deuxieme. 3° Partie, pp. 112-114, Tab. 14,
Fig. 1, 1*.

Description of the genus Glossina, and notes as to its
affinities and probable habits: description of Gl. longi-
palpis 2 (t= Gl. morsitans, Westw.). The figure of the
insect is very poor.

7. 11849. F. Walker.
“List OF THE SPECIMENS OF Diprerous INSECTS IN
THE COLLECTION OF THE British Museum.” Part III.
p. 682.

GORDON CUMMING’S ACCOUNT. 125

Original description of Stomoays fuscus (Glossina fusca),
and description of “ Stomowys longipalpis? Glossina longi-
palpis” (= Glossina palpalis, Rob.-Desv.).

8. i850. J. Macquart.
‘‘Drprires Exorrqgues Nouveaux ou Pru Connvs.”
4¢e Supplément, p. 239, Tab. 22, fig. 4.
Description of Glossina longipalpis g (?) (GI. morsitans,
Westw.). The figure is an unrecognisable profile of the
abdomen.

9. 1850. R. Gordon Cumming.

“Five Years or A Hunter’s Lire iN THE Far INTERIOR
or Sourn Arrica” (London: John Murray), Vol. IT,
pp. 210, 219-220, 227, 270.

“They [natives] also told me that I should lose all
my cattle by the fly called ‘Tsetse’” (p. 210). “ When
under the mountains [on the south bank of the Limpopo
river, in the Zoutpansberg district of what is now the
Transvaal], I met with the famous fly called ‘Tsetse,’
whose bite is certain death to oxen and horses. This
‘hunter’s scourge’ is similar to a fly in Scotland called
‘kleg,’ but a little smaller; they are very quick and
active, and storm a horse like a swarm of bees, alighting
on him in hundreds and drinking his blood. The animal
thus bitten pines away and dies at periods varying from
a week to three months, according to the extent to which
he has been bitten” (pp. 219-220). ‘The next day
one of my steeds died of ‘tsetse. He had been bitten
under the mountain range lying to the south of this
fountain, The head and body of the poor animal swelled
up in a most distressing manner before he died. His
eyes were so swollen that he could not see, and in darkness
he neighed for his comrades who stood feeding beside
him” (p. 227). Death of a pony from “Tsetse ” (p. 270).

10. 1850. J. O. Westwood.

“OBSERVATIONS ON THE DESTRUCTIVE SPECIES OF
Diprerous Insects KNOWN IN AFRICA UNDER THE NAMES
oF THE TsETSE, ZIMB, AND TSALTSALYA, AND ON THEIR
SUPPOSED CONNEXION WITH THE FourtTH PLAGUE OF
Eayrpr” (Proceedings of the Zoological Society of London.
Part XVIIL., pp. 258-270, Pl. XIX., figs. 1, 1-1‘, 2, 3).

126 DOUBTFUL RECORD FROM SENNAR.

Includes the original description and figures of Glossina
morsitans, tachinoides (= palpalis, Rob.-Desv.—var.), and
tabaniformis (= fusca, Walk.).

11. 1852. W. Cotton Oswell.
«Sur UNE MoucHE VENIMEUSE DE L’AFRIQUE MERIDI-
ONALE” (Comptes Rendus Hebdomadaires des Séances de
V Académie des Sciences, Tome Trente-Cinquiéme, pp. 560-
561).
A note, translated in “ Ann, Mag. Nat. Hist.,” Ser. 2,
Vol. X., p. 463.

12. 1852. — Arnaud.
Comptes Rendus Hebdomadaires des Séances de l Académie
des Sciences, Tome Trente-Cinquiéme, p. 603.

A note to the effect that the fly brought home and
presented to the Société de Géographie by Mr. Oswell
seemed to the author to be the same as that met with in
the Isle of Sennar (translated in “ Ann. Mag. Nat. Hist.,”
Ser. 2, X., p. 464).

The following is a translation of M. Arnaud’s note:
«From the examination that I was able to make of this
fly in the office of the Société de Géographie, it seemed
to me to be identical with the one met with in the Isle
of Sennar, between 15° and 11° north latitude, where its
repeated punctures likewise kill animals, obliging shepherds,
especially those who keep cattle, to abandon the country
during the season at which it is most troublesome, that is
to say, in the months from January to May, in order to
take refuge on the banks of the Nile, where the fly is
only very rarely found.

“JT have myself been bitten by one of these flies, and
the sore that resulted lasted more than four months, with
insupportable itchings, which sometimes recur even to-day.”

[Whether the fly here referred to is a species of Tsetse
or not, it seems highly probable that it is identical with
the “Zimb” or “Tsaltsalya” of Bruce. The statement
that the Sennar fly avoids the banks of the Nile certainly
suggests the idea that it is not a Tsetse, though the same
objection might be urged against its being what is known
in the Sudan as a “Seroot” fly, i.e., one of the Tabanide,
either a species of Tabanus near T. dorsivitta, Walk., or
else a Pangonia. In the paper by E. Marno, published in

OSWELL AND ARNAUD. 127

Petermann’s “ Mittheilungen” for 1873 (see 47), the fly is
called Surréta, under which name, however, it is stated
that a large number of species is included. |

13. 1852. De la Roquette.
“ MoUCHE VENIMEUSE DE L’AFRIQUE MERIDIONALE ”
(L’Institut, Tome XX, p. 342).
Reprints of the notes by Oswell and Arnaud in the
Comptes Rendus.

14. 1852. J. O. Westwood.

“OBSERVATIONS ON THE Destructive Species oF
Diprerous Insects KNOWN IN AFRICA UNDER THE NAMES
oF THE TseTsr, ZimB, AND TSALTSALYA, AND ON THEIR
SUPPOSED CONNEXION WITH THE FourTH PLacux or Eaypr”
(Annals and Magazine of Natural History, Vol. X., Second
Series, pp. 138-150).

Reprint of the paper in the P. Z. 8. of 1850, without
the figures.

15. 1852. W. C. Oswell and — Arnaud.

“On A Venomous Fy or Soutnern Arrica” (Annals
and Magazine of Natural History, Vol. X., Second Series,
pp. 463-464).

Translations of notes by the authors in Comptes Rendus,
Tome 35 (1852), pp. 560 and 603. Mr. Oswell’s note was
afterwards printed in somewhat greater detail in the
Proceedings of the Entomological Society of London, 1852,
p. 96 (1853) [see 18]. M. Arnaud, after examining one of
Mr. Oswell’s specimens, considered it identical with a cattle-
destroying fly “‘found in the Isle of Sennar, between
15° and 11° N. latitude ” [see 12].

16. 1852. H. Schaum.
Archiv fiir Naturgeschichte, 18. Jahrgang, IT. Band,
pp. 236-237.
Résumé of Westwood’s paper in the P. Z. 8. of 1850.

17. 1853. H. Schaum.
Archiv fiir Naturgeschichte, 19. Jahrgang, II. Band,
pp. 268-269.
Summary of the notes by W. C. Oswell (Comptes
Rendus, T. 35 (1852), p. 560), and Mons. Arnaud
(op. cit., p. 603), translated in Ann. Mag. Nat. Hist.,
Ser. 2, X., pp. 463-464.

128 ANDERSSON’S STATEMENTS.

18. 1853. W. Cotton Oswell.
Proceedings of the Entomological Society of London,
1853, pp. 96-97.
Observations on the Tsetse-fly of South Africa, com-
municated by Mr. W. B. Spence.

19. 1853. J. O. Westwood.
Proceedings of the Entomological Society of London,
1852, p. 85.
Bibliographical notes.

90. 1856. C. J. Andersson.

“Take Neamt, or, Explorationsand Discoveries, During
Four Years’ Wanderings in the Wilds of South-Western
Africa” (London: Hurst and Blackett). i XXXVIL.,
pp. 488-491.

“ During my hunting excursions along the Teoge, I
encountered, for the first time, that most extraordinary
of insects, the Tsetse (Glossina morsitans, Westw.). Among
the several scourges to which the traveller is subjected in
the South African wilderness, one of the greatest is this
insect ; not, it is true, as to the wayfarer’s own person,
for he himself escapes almost unscathed, but as regards
the horses and cattle.

“The Tsetse is found chiefly in the bush, or amongst
the reeds, but rarely in the open country. It is confined
to particular spots, and is never: known to shift its
haunts. Thus, cattle may be seen grazing securely on one
side of a river, whilst the opposite bank swarms with the
insect. Should the natives, who are well acquainted with
localities frequented by the fly, have occasion to change
their cattle-posts, and are obliged to pass through tracts
of country where it exists, they choose, I am told, a moon-
light winter’s night, as, during the hours of rest in the
cold season, it does not bite” (pp. 488-489).

“ Very lately, indeed, a party of Griquas, about twenty
in number, who were elephant-hunting to the north-west
of the Ngami, and who were provided with three waggons
and a Jarge number of trek or draught oxen, lost, prior to
their return to the Lake, all their cattle by the bite of
the Tsetse. Some horses brought with them to further
their sport shared a similar fate.

“The very same year that this disaster happened to

LIVINGSTONE’S “MISSIONARY TRAVELS.” 129

the Griquas, a party of Englishmen, amongst whom was
my friend, Mr. Frederick Green, attempted to reach
Libébé ; but they had only proceeded seven or eight days’
journey to the north of the Ngami, when both horses and
cattle were bitten by the fly in question, and the party
were in consequence compelled to make a hasty retreat.
One of the number, I am told, was thus deprived of as
many as thirty-six horses, excellent hunters, and all
sustained heavy losses in cattle” (pp. 489-490).

‘** When allowed to settle on the hand of a man, all it is
observed to do is to insert its proboscis a little farther than
seems necessary to draw blood. It then partially with-
draws the dart, which assumes a crimson hue. The
mandibles now appear to be agitated ; the shrunken body
swells, and, in a few seconds, the insect becomes quite
full, and quietly abandons its prey” (p. 490, note).

‘‘A dog, reared on the meat of game, may be hunted
in Tsetse districts in safety ” (p. 491, note 2).

21. 1857. David Livingstone.

“ MisstoNARY TRAVELS AND ReEseARCHES IN SourH
Arrica” (London: John Murray), pp. 79, 80-83, 487—
488, 571.

“The cattle, in rushing along to the water in the
Mababe, probably crossed a small patch of trees contain-
ing Tsétsé, an insect which was shortly to become a
perfect pest to us” (p. 79).

“ A few remarks on the Tsetse, or Glossina morsitans,
may here be appropriate. It is not much larger than the
common house-fly, and is nearly of the same brown colour
as the common honey-bee. The after part of the body has
three or four yellow bars across it ; the wings project beyond
this part considerably, and it is remarkably alert, avoiding
most dextrously all attempts to capture it with the hand
at common temperatures ; in the cool of the mornings and
evenings it is less agile. Its peculiar buzz, when once
heard, can never be forgotten by the traveller whose
means of locomotion are domestic animals ; for it is well
known that the bite of this poisonous insect is certain
death to the ox, horse and dog. In this journey, though
we were not aware of any great number having at any
tmme lighted on our cattle, we lost forty-three oxen. by its

K

1380 LIVINGSTONE’S THEORY OF TSETSE-DISEASE.

bite. We watched the animals carefully, and believe
that not a score of flies were ever upon them.

«“ A most remarkable feature in the bite of the Tsetse
is its perfect harmlessness in man and wild animals, and
even calves, so long as they continue to suck the cows. We
never experienced the slightest injury from them ourselves,
personally, although we lived two months in their habitat,
which was in this case as sharply defined as in many
others, for the south bank of the Chobe was infested by
them, and the northern bank, where our cattle were placed,
only fifty yards distant, contained not a single specimen.
This was the more remarkable, as we often saw natives
carrying over raw meat to the opposite bank with many
Tsetse settled upon it” (pp. 80-81).

Notes on the bite and its effects on cattle (pp. 81-
82).

“These symptoms seem to indicate what is probably
the case, a poison in the blood, the germ of which enters
when the proboscis is inserted to draw blood. The poison-
germ, contained in a bulb at the root of the proboscis,
seems capable, although very minute in quantity, of
reproducing itself, for the blood after death by Tsetse is
very small in quantity, and scarcely stains the hands in
dissection” (p. 82). Animals that suffer from the bite,
or possess immunity (p. 82). “The mule, ass, and goat
enjoy the same immunity from the Tsetse as man and the
game.”

“The curious feature in the case, that dogs perish
though fed on milk, whereas the calves escape so long as
they continue sucking, made us imagine that the mischief
might be produced by some plant in the locality, and not
by the Tsetse ; but Major Vardon, of the Madras Army,
settled that pomt by riding a horse up to a small hill
infested by the insect without allowing him time to graze,
and though he only remained long enough to take a view
of the country and catch some specimens of Tsetse on the
animal, in ten days afterwards the horse was dead.’

“The well-known disgust which the Tsetse shows to
animal excreta, as exhibited when a village is placed in its
habitat, has been observed and turned to account by some
of the doctors. They mix the droppings of animals,
human milk, and some medicines together, and ‘smear the

TSETSE MAY PERISH WITH BIG GAME. i131

animals that are about to pass through a Tsetse district ;
but this, through it proves a preventive at the time, is
not permanent. There is no cure yet known for the
disease. A careless herdsman, allowing a large number
of cattle to wander into a Tsetse district, loses all except
the calves; and Sebituane once lost nearly the entire
cattle of his tribe—very many thousands—by unwittingly
coming under its influence. Inoculation does not insure
immunity, as animals which have been slightly bitten in
one year may perish by a greater number of bites in the
next ; but it is probable that with the increase of guns
the game will perish, as has happened in the south, and
the Tsetse, deprived of food, may become extinct simul-
taneously with the larger animals” (pp, 82-83).

“ Before reaching the Makondo rivulet, latitude
13° 23' 12” S., we came upon the Tsetse in such numbers,
that many bites were inflicted on my poor ox, in spite of
aman with a branch warding them off. The bite of this
insect does not affect the donkey as it does cattle. The
next morning, the spots on which my ox had been bitten
were marked by patches of hair, about half-an-inch broad,
being wetted by exudation (pp. 487-488),

Tsetse met with on the banks of the Chiponga, which
joms the Kafue a few miles above its confluence with the
Zambesi.

Figures of Glossina morsitans (enlarged) and mouth parts
(copied from Westwood’s figures in P. Z. S., 1850), also
small rough woodcut of Tsetse, about natural size (from
figures supplied by Mr. J. E. Gray) (p. 571).

The enlarged figure of the fly is also reproduced on the
title-page of the volume.

22. 1857. Bracy Clark.
“Tue Tzerze or AFRICA IDENTIFIED witH Csrrus
Bovis” (The Zoclogist, Vol. XV., pp. 5720-5721).
A mistaken attempt to show that the Tsetse is identical
with the cattle Warble-fly (Gistrus (Huypoderma) bovis).

28. 1858. L. de Castelnau.
“Sur La Tsirsk DE L’AFRIQUE AUSTRALE” (Compiea
Fiendus hebdomadaires des Séances de VAcadémie des
Sciences, Tome Quarante-Sixiéme, pp. 984-986). ..-
The Tsetse “is generally found on bushes and reeds
Be

132 STATEMENTS BY L. DE CASTELNAU.

bordering marshes, while the plains and other open places
seem less favourable to it” (p. 984).
“Tt occurs in great numbers between—

22° and 26° KE. long. (Greenwich) and 18° and 21° §. lat.

On anneal |. GY, Ki 5 LO ad 2OP" joes
27° and 29°, _,, aii’ 5, 22° and’25?”,, "i;
26° and 28°, _,, n », 24° and 25° ,, ,,. (p. 984).

“A little time ago, some Griquas having with them
eight waggons, attempted to cross the country in which
this insect lives to the north-west of the Transvaal
Republic. They lost all their animals, and were obliged
to abandon their waggons and return on foot” (p. 984).

“This fly seems to live only in localities in which
game abounds. . . . It appears neither to increase nor to
diminish in number, according to the statements of traders,
though, as in all parts of South Africa, the natives are in
the habit of setting fire to the pasture every year... .
The Tsetse most usually attacks the region between the
thighs and the belly of animals. . . .” (p. 985).

‘The Tsetse has no uncertain flight, like most of the
other Diptera ; quick as an arrow, it darts from the top
of a bush to the point that it wishes to attack; it seems
also to possess very keen sight. Mr. Chapman, one of the
travellers who has penetrated farthest into the interior of
South Africa, narrates that while out shooting, having an
almost imperceptible hole in his clothing, made by a thorn,
he often saw the Tsetse, which appeared to know that it
could not penetrate the cloth that he was wearing, make
a dart and succeed, without ever missing its aim, in biting
himn in the small space that was not protected.

“The Bushmen assert that this fly is viviparous, and
Mr. Edwards, the companion of Mr. Chapman, and 4,
highly intelligent man, having one day expressed to them
his disbelief as to this, they brought him a pregnant
female, and having in his presence opened it along the
middle fine of the abdomen, he states that he saw three
little flies ready to take flight emerge from it ” [!!] (p. 986).

94, 1858. J. O. Westwood. -
EXHIBITION. BY Westwoop at A MEETING oF THE
EntomoLoeicaL Society of Lonpon, HELD Ocroser 5TH,
1857, oF A SPECIMEN OF THE TSETSE BROUGHT HOME BY

BURTON’S OBSERVATIONS IN EAST AFRICA. 133

Masor Varpon (again erroneously said to be from Lake
Tchad). (Transactions of the Entomological Society of
London, New Series, Vol. IV., pp. 89-90). Westwood
refutes Bracy’s Clark’s contention (vide 22) that Tsetse is
merely another name for “ Cistrus bovis.”

25. 1860. Captain R. F. Burton.

“THe Lake Recions or CentraL Arrica” (London :
Longman, Green, Longman, and Roberts), Vol. I., pp.
187, 276, 289; Vol. II., pp. 18-19.

“In this foul jungle [inland from Bagamoyo, bordering
the path leading from Mbumi along the right bank of the
Mukondokwa River to its ford], our men also suffered
severely from the Tzetze. This fly, the torment of Cape
travellers, was limited, by Dr. Livingstone, to the regions
south of the Zambezi River. A specimen, brought home
by me and submitted to Mr. Adam White, of the British
Museum, was pronounced by him to be a true Glossina
morsitans, and Mr. Petherick has fixed its limits about
eight degrees north of the equator. On the line followed
by the Expedition, the Tzetze was found extending from
Usagara westward as far as the Central Lakes ; its usual
habitat is the jungle-strip which encloses each patch of
cultivated ground, and in the latter it is rarely seen. It
has more persistency of purpose even than the Egyptian
fly, and when beaten off it will return half a dozen times
to the charge; it cannot be killed except by a smart
blow, and its long, sharp proboscis draws blood even
through a canvas hammock. It is not feared by the
naked traveller; the sting is as painful as that of an
English horse-fly, and leaves a lasting trace, but this hard-
skinned people expect no evil consequences from it. In
the vicinity of Kilwa it was heard of under the name of
‘kipanga,’ the ‘little sword’” (Vol. I., p. 187).

Tsetse abundant at K’hok’ho, in Usenga (now German
East Africa) (Vol. I., p. 276).

Tsetse in a thin forest of thorns and gums, bare of bush
and underwood, near Jiwe la Mkoa, in Mgunda Mi’hali,—
“On the next day the road led through a thin forest of
thorns and gums, which, bare of bush and underwood,
afforded a broad path and pleasant, easy travelling. Sign
of elephant and rhinoceros, giraffe and antelope, crossed

134 BUSHMAN REMEDY FOR TSETSE-BITE.

the path, and as usual in’ such places, the asses were tor-
‘mented by thé Tzetze” (Vol. I., p:' 289).

Tsetse-fly (apparently) in Unyamwezi (to the east of Lake
Tanganyika).—‘‘ The Arabs describe a fly which infests
the forest-patches of Unyamwezi ; it is about the. size of
a small wasp, and is so fatal that cattle attacked by it are
at once killed and eaten before they become carrion from
its venomous effects” (Vol. II., pp. 18-19).

For localities, see Map at the end of Vol. II.

25a. 1862. Dr. (now Sir) John Kirk.

Report oN THE NaturaL Propucts AnD CAPABILITIES
OF THE SHIRE AND Lower ZamBest Vatiteys. By John
Kirk, Botanist to the Livingstone Expedition. Dated
Senna, Dec. 28th, 1860 (Proceedings of the Royal Geo-
graphical Society of London, Vol. VI., p. 29).

“ The Tsetse-fly is unknown among the hills, and very
rare in the Upper Shiré Valley on the eastern side. In
the lower valley, however, it is the natural accompaniment
of the large herds of elephants which inhabit the grass
plains and marshes.”

26. 1864. Thomas Baines.*

“ Explorations IN SourH-Wesr Arrica” (London :
Longman, Green, Longman, Roberts and Green), pp. 255,
351, 469, 470-471, 511.

Bushman remedy for Tsetse-bite—“ On a former occa-
sion a Bushman gave Chapman a remedy for the bite of
the Tsetse ; it seemed to be a parasitic plant growing
either in the hollow of a tree or between the wood and
the bark. Chapman thought it acted by restoring to the
blood the iron of which perhaps the bite of the fly deprived
it, but he sent it home with other specimens by a friend,
and has not yet heard any report on it” (p. 255).

Tsetse on the Teoge River, a tributary of Lake Ngami
from the north (1862).—* This is not unlikely to be the
Teoge River, and if so, it would not do to take the cattle
near it on account of the fly ” (p. 351).

Tsetse on the Zimboya Liver, a tributary of the
Matietsie River, south-west of Victoria Falls (July,
1862).—‘‘. . . we were riding on the track by which

* Baines accompanied Chapman in his travels.

BAINES’ EXPERIENCE AT VICTORIA FALLS. 135

I had returned, when Chapman came galloping back for
life (not his own but his horse’s). A fly had settled on
the rump of his steed, and though he had driven it off, he
could be by no means sure that others had not sucked
blood, and left the subtle venom with which they dilute
it. We altered our course, and held more north along
the rivulet in the open places, galloping the oxen when-
ever we were forced to enter the bush. A herd of water-
boks appeared ; Chapman wounded one and gave chase,
but again the fly attacked him and forced him to a pre-
cip.tate retreat” (pp. 469-470).

[Zimboya R.| “I saw a dozen of the dreaded little
pests hovering with that rapid motion of the wing that
keeps the insect stationary over the devoted cattle [trek-
oxen]; but near an hour passed before we were again
ready to move, and a few minutes of that time has most
likely served to inject the poison which dooms twelve
working oxen, two horses, and the cows, to a painful and
lingering death. No one can be assured of this, till in
three weeks or so the staring eye and roughened coat
begin to tell the tale of gradual waste. I heartily wish
(hope, I can hardly say) we may be mistaken, but our
guide, who had seen no fly, when asked whether he was
certain on the point, replied with an air of astonishment,
‘Is then the Tsetse a thing that a man forgets when he
has once seen it ?’” (pp. 470-71).

Tsetse at ihe Victoria Falls.—‘‘ Another hindrance is
the annoyance caused to the painter [of the Falls] by the
incessant persecutions of the Tsetse. At the moment,
perhaps, when one requires the utmost steadiness and
delicacy of hand, a dozen of these little pests take advan-
tage of his stillness, and simultaneously plunge their pre-
paratory lancets into the neck, wrists, and the tenderest
parts of the body ; one or more cunning fellows actually
selecting the places where the lines of fortune radiate or
cross, with a skill in palmistry that would do honour to
an experienced gipsy ” (p. 511).

27. 1865. David and Charles Livingstone.
“ NARRATIVE OF AN EXPEDITION TO THE ZAMBESI AND

irs TripuTaRies”” (London: John Murray), pp. 206-207,
232-933, 424,

136

BATOKA REMEDY FOR TSETSE-BITE.

Tsetse near the Valley of the Zambesi, to the west of
Zumbo.—“ We left the river here, 2nd proceeded up the
valley which leads to the Mburuma or Mohango pass. . . .
Tsetse-flies are more numerous and troublesome than we
have ever before found them. They accompany us on the
march, often buzzing round our heads like a swarm of
bees. They are very cunning, and when intending to
bite, alight so gently that their presence is not perceived
till they thrust in their lance-like proboscis. The bite is
acute, but the pain is over in a moment ; it is followed by
a little of the disagreeable itching of the mosquito’s bite.
This fly invariably kills all domestic animals except goats
and donkeys; man and the wild animals escape. We
ourselves were severely bitten on this pass, and so were
our donkeys, but neither suffered from any after effects”
(pp. 206-207).

Batoka remedy for Tsetse-bite.——‘‘ Another [Batoka]
Chief, who died a number of years ago, believed that he
had discovered a remedy for Tsetse-bitten cattle ; his son
Moyara showed us a plant, which was new to our botanist,
and likewise told us how the medicine was prepared ; the
bark of the root, and, what might please our homeopathic
friends, a dozen of the Tsetse are dried and ground
together into a fine powder. This mixture is administered
internally, and the cattle are fumigated by burning under
them the rest of the plant collected. ‘The treatment must
be continued for weeks, whenever the symptoms of poison
appear. This medicine, he frankly admitted, would not
cure all the bitten cattle. ‘For,’ said he, ‘cattle, and
men too, die in spite of medicine; but should a herd by
accident stray into a Tsetse district and be bitten, by this
medicine of my father’s, Kampakampa, some of them could
be saved, while, without it, all would inevitably die.’
He stipulated that we were not to show the medicine to
other people, and if ever we needed it in this region we
must employ him; but if we were far off we might make
it ourselves ; and when we saw it cure the cattle think of
him, and send him a present” (pp. 232-233).

Native ideas of mode of reproduction of the Tsetse.—
*‘Our distinguished countryman, Professor Owen, recom-
mended our attention to be directed to the genesis of the
Tsetse, in order to discover a means for the extirpation

SIR JOHN KIRK’S OBSERVATIONS. 137

of this pest. We frequently inquired of the different
tribes if they could help us in our inquiries ; and one of
the Makololo remembered that this very question was
once under public discussion at Linyanti; and, as usual, a
bet was laid that no one could tell. After a number of
days had elapsed, an old man claimed the prize, asserting
that the Tsetse laid its eggs, which were of a red colour,
on the leaves of the mopane tree. These were probably
only the eggs of an insect, described in the ‘ Missionary
Travels,’ as depositing over its eggs a sweet gum, which
is collected and eaten. Some denied that he had seen
them ; others affirmed that the red eggs were laid on the
twigs of trees, and not on the leaves ; and others insisted
that the eggs were placed in the droppings of buffaloes,
and these last were probably in the right. The destruc-
tion of all game by the advance of civilization is the only
chance of getting rid of the Tsetse” (p. 424).

28. 1865. Dr. (now Sir) John Kirk.

* On THE ‘ Tsetse’ Fry or Tropicat Arrica ” (Glossina
morsitans, Werstwoop) (The Journal of the Linnean
Society, Vol. VIII., pp. 149-156).

“During the Zambezi Expedition, the ‘ Tsetse’ Fly
has been met with by us on many occasions and at distant
parts. It was first seen on the Zambezi at Lupata, a
hilly and well-wooded district 150 miles from the coast.
To the north-east, between the Zambezi and Shire, it is
very abundant. Insome parts of the Batoka country, near
the Victoria Falls, it was again found; also at the
junction of the Chobe ; and in immense numbers on the
south bank, not far from the confluence of the Kafue.

‘On the Rovuma river in 10° S. Lat. it is met with
eight miles from the coast, and extends along its banks
for 115 miles, the furthest point explored ; here we found
it named ‘ Chipinga’; ‘ Kipanga’ being that given to it
at Kilwa, according to Captain Burton. Wherever met
with over this wide area of Tropical Africa south of the
equator, the habits of this fly as recorded are the same.

“Tt frequents open forest and well-wooded country,
being absent from extensive grass plains. In the morning
while the dew hangs on the grass, and before the heat of
the rising sun has warmed the air, the ‘Tsetse’ is dull

138

EFFECTS OF TSETSE-BITE ON MAN.

and sluggish, resting on the under side of some leaf or
blade of grass ; when forced to take wing they may then
be easily caught.

“ Even at nine o'clock they are not very active, and
fly about with a peculiar buzzing sound ; with the heat of
the day they become a real annoyance to the traveller,
constantly biting him on the hands, face, or neck,
dextrously evading a blow, and again alighting on the
very spot from which they have been driven. If per-
mitted, they will gorge themselves with blood and become
unable to fly to a distance. On man, the effects are not
more than follow an ordinary mosquito bite, redness,
swelling, and local irritation remaining for about an hour,
varying in amount according to the state of the individual.
In itself the bite is not so severe as that of the larger
Tabanidae.

«By night I have never been bitten by ‘ Tsetse,’ nor
do they fly about after sunset. They are most numerous
and troublesome in the hot, sultry weather before rains.

‘When once attention has been directed to the fly, it
cannot again be overlooked, although, from its common
and insignificant appearance, others might easily be
confounded with it by those who have not before expe-
rienced it. There is, therefore, more danger of its
geographical range being exaggerated than under-esti-
mated. Wherever I have found this fly, it has been
accompanied by the buffalo or elephant ; the native dog
and goat are the only domestic mammals I have found in
a country infested by it.”

* x * ‘* *

“ Although always found in company with large game,
the fly does not follow it everywhere ; other circumstances,
still unknown, check its universal distribution.

«“ Between Sesheke and Linyanti there is plenty of
game, yet the fly is limited to certain narrow spots, and
in like manner between Sesheke and the Victoria Falls.
So are these falls shut out from the south by only a
narrow belt of ‘ fly’ land, which may be crossed at night.

‘As much of what we know on these points rests on
native information, I would remark that where the person
obtaining it enjoys the contidence of the people and can
speak with them ina common language, without depending

MAKALOLO METHODS OF PROTECTION. 139

on interpreters, native testimony on matters of fact is
quite as good as European.

“The Makalolo are a people from infancy accustomed
to tend cattle, possessing a thorough knowledge of the
most fattening pastures to be sought, and noxious herbs to
be avoided. Their only wealth consists in cattle, which
they number by thousands. All affirm that on entering
certain localities by day the oxen die shortly afterwards ;
this they have proved, not in the small numbers of twenty
or forty noticed by Europeans, but in herds consisting of
hundreds ; whether in great or small numbers they have
found the result alike. They have further learned that
these deadly places may be crossed with safety by night
if sufficiently narrow to allow of the cattle being driven
through before sunrise. This has been tested by
Europeans and found also correct ; further, that goats
remain unaflected, and sheep suffer in a less degree than
oxen.”

* * * * +

“ The fly avoids human excrement, so the natives told
us, and we have found it true, and they say that cattle
have been passed by day through fly country when
smeared with a composition containing this. Native
doctors have an herb to which they attribute a similar
effect, but even they never assert that it will save all ;
only a small per cent. of the cattle exposed is the most
they claim ” (pp. 152-154).

Lung-sickness, African distemper, etc., “ differ mani-
festly from the ‘Tsetse’ disease in being contagious and
spreading from one place to another and from one animal
to another, whereas only those bitten by the fly die ; and
no danger has been apprehended or experienced ri eugp
cattle mixing with others” (p. 154).

“The first symptoms [of Tsetse-fly disease] appear
commonly within four days, but this varies with the
number of flies and the season of the year. Natives
report that cattle bitten die in greatest numbers before
the rains, or when they set in, and that some animals will
linger on until then ; that having passed a fly country you
do not know the full amount of loss until the rainy season
has begun ” (p. 155).

140 SWARM OF TSETSE ON ZAMBESI.

‘In most Tsetse countries the traveller is not usually
beset with more than two or three at a time ; in the course
of the heat of the day these might produce a number of
bites, besides falling in with fresh flies as one advanced.
But they are sometimes found in much greater numbers.
On the south side of the Zambezi, near the confluence of
the Kafue, while walking along the river-bank in search of
game, under flat-topped acacias, I heard a buzzing sound,
and saw a cloud of insects coming towards me. Supposing
them a swarm of bees, I ran off, while they followed. On
looking back I found it was only ‘Tsetse’; so, arming
myself with a leafy branch, I kept them off and continued
my journey; they accompanied me for some distance
however. I have never again seen them congregate in
this manner ; and, curiously enough, on this occasion, and
on this only, did I obtain two of what may be the male
insect ; these bear the proportion to the females of 1 : 30,*
judging by the numbers then caught” (p. 155).

General description of the structure and mode of action
of the proboscis (pp. 155-156).

The fatal effect of the bite of the Tsetse in the case of
domestic animals ascribed to ‘‘irritant matter” injected by
the fly, in order to “ facilitate the sucking of blood”

“ The irritation which follows the bite in man shows
that some irritant matter is at the same time injected
(although no organ for its secretion has yet been detected),
the object of which is no doubt to cause a local congestion,
and thus facilitate the sucking of blood. The accidental
effects of this, which in animals among whom the ‘Tsetse’
naturally lives produces no after result, in the domestic
animals before-named proves fatal” (p. 156).

29. 1866. Sir Samuel W. Baker.

“THe ALBERT N’yAnzA, GREAT Basin OF THE NILE,
AND EXPLORATIONS OF THE Nite Sources” (London:
Macmillan & Co.): Vol. I., p. 376.

Tsetse-fly in Obbo (S8.E. of Gondokoro, just north of
4° N. lat., and between 32° and 33° E. long.).—* The
wet herbage disagreed with my baggage animals.
Innumerable flies appeared, including the Tsetse, and in
a few weeks the donkeys had no hair left, either on their

* (Cf. p. 86.]

TSETSE FIRST ENCOUNTERED ON LIMPOPO. 141

ears or legs ; they drooped and died one by one. It was
in vain that I erected sheds and lighted fires; nothing
would protect them from the flies.”

30. 1868. James Chapman.

“TRAVELS IN THE INTERIOR oF SoutH AFRica” (London :
Bell and Daldy ; Edward Stanford): Vol. I., pp. 174-
177, 180; Vol. II., pp. 214, 225, 249-250.

Makololo remedy for calves suffering from Tsetse bite.—
“Their [the Makololo’s] cattle are carefully kept in the
plains, or in parts known to be perfectly free of fly. The
only remedy they use, they say, is to administer the fly
in milk to a fly-bitten calf, but they do not seem to be
very sanguine about the cure” (Vol. I., p. 174).

“ Now a word with regard to that insignificant-looking
insect, the Tsetse, or poison-fly. This great barrier to
African travelling was first met by the Boers and other
travellers on the Limpopo; and though most people on
their first encounters felt doubts regarding its repute of
the sting being fatal to horses and cattle, too painful
experience of its ravages has left no doubt on the
subject” (Vol. I., pp. 174-175).

“The Tsetse is, in extreme length, half an inch, or
very little more, and has very much the appearance of a
young bee just escaped from its cell, or a bee half-drowned
in honey, the wings being always closed when stationary.
. . . It is extremely quick of sight and keen of scent ; its
flight is rapid and straight.

“The bite of the Tsetse is something like that of the
mosquito, but the pain is not so lasting. It assails
different animals in their most defenceless parts: a man
behind the back between the shoulders, and an ox on the
back or under the belly; a horse in the same places, and
inside the nostrils ; and a dog on the forehead, ete. With
the proboscis they penetrate a pilot cloth coat and whole
suit of under-clothes. The bite of this insect is fatal to
cattle, horses, sheep, and dogs; but there is a peculiar
breed of the latter known as Makoba dogs, which are
exempt from the effects of its poison, the breed having
from time immemorial been reared in the “fly” country,
and escaped a cow milk diet, as the natives say. It has
no ill-effects whatever on game or upon men, except that

14

9

_

CHAPMAN ON FLY-DISEASE.

the being bitten by numbers is likely to induce headache,
as with the irritation of mosquitoes. The symptoms, as I
have observed them, are, first, in the ox, a swelling under
the throat, which, if lanced, emits a yellowish fluid.
The hair stands on end, or is reversed. The animals
become debilitated ; and, though the herbage be ever
so luxuriant, refuse to eat their fill, and become thin.
The eyes water, and at length, when the end is
approaching, a continual rattling in the throat may be
heard at a few paces’ distance. It sometimes happens
that a fly-bitten ox will live, but very rarely, and only
when it has no work to perform. Work and rain are
great precipitators of their end. In horses the symptoms
are swelling about the eyes, nostrils, testes, the hair is
reversed, and, though they have the best of food, they
become thin, sleepy, and, pining gradually, at length die.
“Both cattle and horses live from fourteen days to
six months after having been bitten by Tsetse, but they
generally die after the first rain has fallen. A dog dies
in ten or twelve days, or two or three weeks at. latest.
It is perceptible in the eyes, which are swollen and
protruding. After death the heart of an ox is generally
incased in a yellowish glutinous substance, which might
be mistaken for fat. The flesh is full of little bladders
of fluid, and the blood also is half fiuid, which becomes
congealed on cooling. The vitals are of a livid colour.
“The Tsetse-fly is generally found within a few miles
of water, in rich sandy ridges near marshy spots, and
generally in mopani or mimosa forests. I have known
them to shift their positions, or encroach on new ground,
or leave parts where fire-arms have driven the game out
of a district. They are mostly only found within a
certain range from water. To the buffalo m particular
the insect is more attached, and often moves about with
them in the rainy season” (Vol. I., pp. 175-177).
Author’s oxen and horses discovered to have been
bitten by the fly, ia the Makololo Country, on the. Chobé
River, W. of the Victoria Falls: two or three cattle die ;
the “speedy death” of the remaining animals considered
inevitable (Vol. I., p. 180).
Tsetse on the south bank of the Zambesi,:close to.and a
little to the west of the junction of the Gwai or Quagga R.—

TSETSE ON 8. BANK OF ZAMBESI. 143

“Ist November [1862].—The Tsetse are very troublesome
this morning. We did not notice them here in going,
nor at the last halting-place. I think there must be
“fly” from near the west bank of the Gwai all the way
to the Victoria Falls, and beyond to the junction of the
Tamalukan or Lingalo’s river, near Lake Ngami. Beyond
the Gwai to Sinamani’s [on the Zambesi, about thirty
miles to the east of the junction with the Gwai river] I
feel pretty confident there are none. There is no access
to the Zambesi with bullock-wagons, nor hunting on
horseback, nor with dogs, in these parts, unless with
great sacrifices, nor can I find a wagon-road, owing to the
broken nature of the ground—vrocks, ravines, and moun-
tains—even where there is no ‘fly.’ . . . Down the Daka
river [according to the author’s map this is otherwise
known as the Luisi, or Luluisi R.: it flows N.E., and
joins the Zambesi about 20 miles W. of the Gwai R.], too,
I think a road might be found as far as the river [Zambesi|,
though a district infested by ‘fly’” (Vol. IL., p. 214).

No ‘‘fly” on the south bank of the Zambesi, between the
junction of the Gwai R. and Sinamani’s, a little to the E. of
the junction of the Sebungo R., and about thirty miles E. of
the junction of the Gwai; “ but this may be owing to the
density of the population, and the scarcity of buffaloes,
which are so much hunted by the Batonga, on account of
the ravages they commit in the gardens” (Vol. IL,
p- 214).

* T think, with a little more knowledge of the country,
I could even find a way by which to take a wagon to the
Zambesi, but it would probably be impossible wholly to
escape the ‘fly’ in doing so. ‘These pests make their
appearance when and where you least expect them. I[
have not noticed any since leaving the Luluesie, but think
they must extend 10 or 15 miles this side of the river [i.e.
north of the Luluesie] along the mountains” (Vol. IT.,
p-. 225).

Tsetse-fly near the Luluesie River, Dec. 1862.—“ In the
evening John fell in with four buffaloes close by, and
shot one of them ; but he brought the terrible news that
there were thousands of the ‘fly’ on the one he had
killed. I gave orders that the cattle should go out early
to graze on the open to the south, while he returns to

144 CHAPMAN ON TSETSE AND BUFFALO.

the buffalo, and endeavours to catch some Tsetse for my
inspection.

“31st December.—John went off early, and soon sent
me a fly, which, happily, proved not to be the dreaded
Tsetse. The Makalakas say the ‘fly’ was brought into
these parts by the elands from the north, after they had
been denuded of their former inhabitants. Heavy rains
are said to kill the fly in the season when they prevail. I
have learned from the natives that the Tsetse deposit
their larve in the dung of the buffalo. Snyman tells me,
that when he was living at Sekelétu’s [another name for
Linyanti, on the Chobe R.] a report was brought in that
Tsetse had crossed the river to a cattle post. The people
in charge fled with the cattle, and Sekelétu sent off a party
to burn them back again, firing the grass, and afterwards
sent the cattle there again. I do not quite understand
this, for I know fields burnt down regularly every year
which are still infested with Tsetse. JI think the only
chance of exterminating them is to keep up constant
warfare with the buffaloes until they are driven out. Gene-
rally, when they have entered a new country with game,
they soon increase and extend themselves farther every
year, if the country is suitable” (Vol. IT., pp. 249-250).

$1. 1868. J. R. Schiner.
“ Reise der Osterreichischen Fregatte Novara um die
Erde:” Diptera, p. 311.
Note on a specimen of “ Glossina longipalpis W.”
(probably = Gl. morsttans, Westw.) from “Central Africa.”

$2. 1868.

“Din GrrTFLiece TzETze opEe TZALTZALA IN ABES-
stntEN” (Das Ausland, 41. Jahrgang, p. 192).

A short article in which the identity of the Tsetse
with the “Tzaltzala” (Tsaltsalya) of James Bruce* is
accepted as proved : it is suggested that since the name
“ Tzaltzala” has priority, it should be adopted instead of
Tsetse. Further references are given to allusions to the
“ Tgaltsal ” in the Bible.. In Job xli. 7, the word is used in
the sense of a fish-spear, therefore it is considered that its
transference to the fly is due to the sharpness of the
latter’s proboscis.

* Cf. (t]

33.

34.

35.

36.

37.

* See
Siid-Afrika” (Petermann’s Mittheilungen, 16. Band, 1870).

USE OF AMMONIA BY MAUCH. 145

1868.

“La Tsrtst, Diersre pe L’Apysstnie ” (Recueil de
Médecine Vétérinaire, Vol. 45 (5. Sér. Vol. V), pp. 148-
149).—From L’ Opinion Nationale, Dec. 3, 1867.

[I have not seen this paper. ]

1868.

“ Diz Tsetse-FuiEceE ” (Aus der Natur, 46. Band (Neue
Folge, 34. Band), pp. 783-784).—See also 47. Band
(Neue Folge, 45. Band), 1871, p. 447.

[I have not seen either of these papers. ]

1868. F. Jaennicke.
An article in Ergtéinzungsblitter zur Kenntniss der
Gegenwart, Band IIL., p. 680.
[I have not seen this paper. |

1869. Karl Mauch.

Mittheilungen aus Justus Perthes’ Geographischer Anstalt
tiber Wichtige Neue Erforschungen auf dem Gesammtgebiete
der Geographie, von Dr. A. Petermann. 15. Band, 1869,
pp. 190-191.

In a letter written in the course of his third journey
of exploration from the Transvaal towards Matabele Land,
and dated from the Olifant’s River, 24° 20’ 8. Lat.,
31° 37' E. Long., July 20, 1868,* Mauch says :—“ For
two days past we have been in the fly (Tsetse) country ;
neither in the case of the dog nor in that of the ox was it
possible to make an external application of Asa foetida,
while an internal application of ammonia was similarly
impossible owing to want of fat and meat” (p. 190),

In a further letter from Inyati, dated October 19,
1868, Mauch writes :—“ The pack-ox I was obliged to
shoot and eat on the Letsobo, although it was seriously
injured by Tsetse ; the dog died from want of flesh food,
or indeed of food of any kind, after I had secured him
against the Tsetse by the aid of ammonia. . . .” (p. 191).

1869. John and Mrs. Petherick.
“TRAVELS IN CENTRAL AFRICA AND EXPLORATIONS

or THE WesTERN Nite Trisutartes” (London: Tinsley
Brothers), Vol. I., pp. 220-221, 222.

map: ‘Originalkarte von C. Mauch’s Reisen im Innern von

L

146

38.

TSETSE NORTH-WEST OF GONDOKORO.

Tsetse-fly at a deserted kraal called Augur, on a low
eminence in a clearing in dense forest in which the ground
was slightly inundated and covered with rank grass as
high as a horseman’s shoulders,—near Adael, in the
country of the Rhol Tribe, north-west of Gondokoro.—
“ Atchwack tells me that the kraal is now, during the
rainy season, untenable from the presence of the destructive
Tsetse-fly, here called the mau. I had as yet seen none,
and hoped our horses would escape, but Atchwack said
to-morrow’s march might prove troublesome, as the fly
was known to exist in the bush in advance of us. The
chief, a more intelligent man than ordinary, states that
the sting of the mau, if in the head or spine of a bullock,
causes speedy death, but, if in the body, the animal might
linger a week or ten days ; but death is inevitable unless
a part of the root of a tree called T'shol goote is administered

_ internally, and rubbed well into the hide; the animal

might then recover. The fly occupies a certain bush, well
known to the negroes; at no great distance cattle may
graze with impunity ; but if they enter the limits occupied
by the fly, the consequences are fatal to numbers of the
herd, if not to the whole” (pp. 220-221).

[The approximate geographical position of the above
locality is 30° 8’ E. Long., 6° 35’ N. Lat., west of the
Nile, and north-west of Gondokoro. |

The march was continued on the following day :—“ Off
at seven A.M., and in ten minutes were following a narrow
track through a continuation of yesterday’s thick forest.

** Two halts of a few moments’ duration, when an eager

‘look-out was kept for the Tsetse or mau, several specimens

of which were netted ; they were precisely similar to the
description and sketch given by Dr. Livingstone, but
perhaps a trifle smaller. When marching not a fly was
seen” (p. 222).

1870. St. Vincent Ersicine.

The Natal Mercury, May 31, 1870 :—“ The meeting of
members of the Natural History Association of Natal,
held last evening, 30th May, 1870, was most successful.
... Mr. St. Vincent Erskine’s paper on the Tsetse-fly
was very interesting, and combated the popular idea that
the bite of the Tsetse-fly was destructive to the life of

OPPOSITION TO BELIEF AS TO TSETSE. 147

certain animals, especially the ox, horse, and dog.* Dr.
Livingstone’s statements on this point were severely criti-
cised by Mr. Erskine, whose theory was that the deaths
of the animals were to be attributed more to change
of grass or climate than to the bite of such a small fly as
the Tsetse. Referring to Dr. Livingstone’s assertion that
the natives of the Matabele country held the belief that the
Tsetse-fly destroyed their cattle, he said that natives of the
west of Africa, in whose country the fly was to be found,
and who could not keep cattle, did not attribute the
destruction of their cattle to the fiyidt

[Copied from The Entomologist, Vol. V., Dec, 1870,
p- 217, where the above report is reproduced. ]

39. 1870. Edward Newman.

Norte on Mr. St. Vincent Ersxkrnz’s PAPER REFERRED
TO ABOVE (The Entomologist, Vol. V., Dec. 1870, p. 218).
“Having invariably maintained that the word Tsetse
implied a disease rather than an insect, and was applied
indifferently to all flies that settled on diseased cattle, or
indeed on any cattle, and having incurred an oyerwhelm-
ing amount of ridicule for holding so heterodox an Opinion,
Tam delighted to find the opinion held also by a resident,
who has every means of obtaining the best information.
T have always protested against the importation of a
myth like the Tsetse into the domains of science; the
mixture of truth and fable in matters of science is always
to be deprecated. The Cholera-fly and the Aphis vastator
are banished from the domain of science 3 it is abundant
time to banish their African congener.”

40. 1870. Karl Mauch.

“Karu Mavcn’s Reisen 1m INNEREN von SUp-AFrika ”
(Mittheilungen aus Justus Perthes’ Geographischer Anstalt
tiber Wichtige Neue Exrforschungen auf dem Gesammtgebiete
der Geographie von Dr. A. Petermann, 16. Band. P. 1),

Letter written from Potchefstroom, on June 30, 1869,
describing the author’s journey from Lydenburg to
Inyati, July 10 to October 17, 1868 :—[Translation. ]
“The pack-ox, that I took with me from Lydenburg,
proved himself the most incapable beast of burden for the
long journey... . In spite of every attention, he soon

* Cf. [52.]

148 IMMUNITY OF DONKEY.

lost flesh, and the consequences of the innumerable bites
received day and night from the Tsetse-fly completely
finished him. The poison possesses the property of decom-
posing the blood, and thereby hinders nourishment and
relaxes all muscles. When a short day’s march south of
the Limpopo I found myself compelled to turn him finally
to account by eating him. Sal ammoniac—each week a
piece the size of a walnut dissolved in water and adminis-
tered internally——had no visible effect and is consequently
no antidote.

“The female donkey, that I bought at Lydenburg for
£9, held out better. .. . The main advantage, however,
consisted in the fact that the Tsetse could not do her any
harm, whether it be that the donkey finds in certain
leaves or in the bark of certain trees an antidote against’
the poison—or that the long hair or the effluvium from
the beast keeps the insect off.”

At the end of the volume of Mittheilungen in which
the above remarks appear is a map entitled : “ Originalkarte
von C. Mauch’s Reisen Im Innern von Siid-Afrika zwischen
Potchefstroom und Zambesi 1865-1869. Nebst Ubersicht
aller anderen Forschungen. Von A. Petermann.” This
map shows by means of a coloured band the ‘‘ Limit of the
Tsetse-fly ” (“‘ Grenze der Tsetse-Fliege”’) in the vicinity of
the Limpopo, and also farther north, to the south of the
Zambesi, in Mashonaland.

41... 1873. E.-C. Buxton.

The Entomologist, Vol. V., April, 1871, pp. 283-284 :—
«The plains on the south side of the Lobombo Mountains,
towards Delagoa Bay, was the only district where I met
with the Tsetse-fly, and immediately below the mountain
they seemed more numerous than at a greater distance.
The belief of the natives in the dangerous character of the
fly is universal ; and I never heard any doubt expressed
about it among the white hunters, many of whom have
come to this district for many years. We were told that
if we took our dogs over the mountains they would be
bitten by the fly, would go blind in a few days, and die in
ten days or a fortnight. The fly, which was pointed out
‘to us as the Tsetse, was very like a small Horse-fly (cleg,
as they are called in Lancashire) ; it was very common,

E. C. BUXTON’S OBSERVATIONS. 149

and answered the description and picture in Living-
stone’s first book accurately. We were frequently bitten
by them: the bite was very sharp, and felt like a red-hot
iron run into the flesh, but it did not leave any mark or
inflammation. I caught several, but mislaid them some-
where, as I have been unable to find them. The dogs
were frequently bitten, and one of them went blind
within a week, and died in about a fortnight. The other
did not show illness for some time later ; and, as we left
him with some of our party, knew nothing more about
him than that he died. The fly appears only at certain
seasons, and in limited localities. The head of a kraal,
about thirty miles distant from the point where we found
the Tsetse most abundant, told us at that time the fly was
not in his district, and pointed to a heifer and some goats,
which he said he intended to send away before the fly
season came on. ‘There is a general opinion that the fly
is connected, in some way, with the larger game, elephants,
rhinoceros, etc., and some think that it breeds in their
dung; but I never heard of any proof of this. Mr.
Erskine was the only person that I met with who
expressed any doubt about the Tsetse ; but, as I have not
seen his paper, I do not know why he doubts it. The fly
district nearest to Natal is about twenty days’ journey
distant. Our horses and oxen we did not take into the
fly country at all.”

42. 1871. E. Mohr.

“EpuarD Mour’s. Reisen 1m INNEREN von Sitp-
AFRIKA, VON DEN TatTse-GOLDFELDERN BIS ZUM ZAMBESI
UND zURUCK NACH Narat, 20. MArz—d), DezemBer 1870”
(Mittheilungen aus Justus Perthes’ Geographischer Anstalt
tiber Wichtige Neue Erforschungen auf dem Gesammtqebiete
der Geographie, von Dr. A. Petermann. 17. Band,
p. 164).

[Translation.] “ Finally at two o'clock in the after-
noon of May 25th my waggon reached its most northerly
point at 19° 10’ 51” 8. lat., and the manner and method
of my journeying thence to the Victoria Falls of the
Zambesi now underwent a change. Five nautical miles
further to the north the Tsetse-fly was encountered, not
in large swarms it is true, but nevertheless, singly, and it

150 CONDITIONS ON JUBA RIVER.’

would consequently have been an act of the greatest
imprudence to drive the indispensable oxen and the useful
horses still further” [ep. 52].

43. 1871. Otto Kersten.

‘Baron CarL CiLaus von DER Deckern’s Reisen IN
Ost-AFRIKA IN DEN JAHREN 1862 Bis 1865” (Leipzig und
Heidelberg: C. F. Winter’sche Verlagshandlung), Band
IT., pp. 83-84, 303, 304.

[Translation.] ‘ Nevertheless the Tsetse-fly (Glossina
morsitans, Westw.) which is so common in South and
North Africa does not occur in the precise territories
traversed by us in our journeys to Dschagga, but rather in
the Galla and Somali countries (we shall deal later on
with the fly and with the devastation brought about by it
there) ; instead of it, however, a representative exists
here in the shape of the Donderobo-fly, which is dangerous
to donkeys (see Band I., p. 249). Of species of horse-
flies (Tabanus), which in the North of Africa plague cattle
in an often incredible manner, one hears nothing here,
probably for the reason that the herds do not pass their
time in the sunny plain, but for the most part on shady
and cooler hills, whither these bloodthirsty flies rarely
stray ” (Band IT., pp. 83-84).

[Dschagga is the district on the southern slope of
Kilima Njaro. The territories passed through between
Mombasa and this region, according to Map VIII. at the
end of the volume, were the country of the Wateita, the
coast region from Mombasa to Wanga, and a line from
the latter to Dschagga running along the northern
boundary of the mountain-region Usambara, Pare, and
Ugono. The years in which these journeys took place
were 1861-62. ]

Tsetse-like, cattle-destroying fly in the vicinity of Manam-
sunde, a town on the Juba River, not far from its mouth,
and just north of the equator. The Juba River enters
the sea a little north of Kismayu.

[Translation.] ‘They [the inhabitants of Manam-
‘sunde, chiefly Wasegua] occupy themselves almost exclu-
sively with agriculture ; for owing to the occurrence in
numbers of a poisonous, Tsetse-like fly, they can keep only

‘ goats and sheep, but no cattle” (Band IT., p. 303).

BLOOD-SUCKING FLIES IN UKWERE. 151

Further on (Band II., p. 304), in writing of the
Wabuni, a scattered tribe of Galla, of whom there are
several settlements on the north shore of the Lower
Juba, while they extend as far as 3°8., the author
says :—

“Tike the Galla and Somali they live chiefly by
hunting. Moreover, they keep small domestic animals,
but no cattle, since in the territories occupied by them
the Tsetse-fly makes this impossible.”

44, 1871. E. Newman.

“Tue Tserse” (The Entomologist, Vol. V., May, 1871,
pp. 289-290).

A note criticising E. C. Buxton’s remarks printed
above [41], which are considered by Newman to refer to
Hematopota (Family Tabanide),—“ All that Mr. Buxton’s
note proves is that ‘clegs’ in South Africa are numerous,
and that their bite is innocuous to human beings.”

45. 1872. H. M. Stanley.

“How I Founp Livinestonn. Travels, Adventures,
and Discoveries in Central Africa; Including Four
Months’ Residence with Dr. Livingstone” (London:
Sampson Low, Marston, Low, and Searle), pp. 87-91,
213, 330, 354.

Tsetse-fly near Rosako, in Ukwere (about 20 miles
inland from Bagamoyo). Three different species of biting
flies were met with, and even found in the author’s tent,
The first, called Mabunga by the natives, is about an
inch long and evidently a species of horse-fly (Tabanus).
Stanley’s men “ unanimously stated that its bite was fatal
to horses as well as to donkeys.” “This fly, along with a
score of others, attacked my grey horse, and bit it so
severely in the legs that they appeared as if bathed in
blood ” (pp. 88-90).

Of the second species of biting fly only one specimen
was seen during the author’s stay at that particular camp:
the description is too vague to admit of its identity being
determined (p. 90).

“The third fly, called ‘chufwa,’ pitched a weak alto-
crescendo note, was a third larger than the house-fiy, and
had long wings. If this insect sang the feeblest note, it
certainly did the most work, and inflicted the most injury.

152 STANLEY’S EXPERIENCE IN EAST AFRICA.

Horses and donkeys streamed with blood, and reared and
kicked through the pain. So determined was it not to
be driven off before it obtained its fill, that it was easily
despatched ; but this dreadful enemy to cattle constantly
increased in numbers. The three species above named
are, according to natives, fatal to cattle; and this may
perhaps be the reason why such a vast expanse of first-
class pasture is without domestic cattle of any kind, a
few goats only being kept by the villagers. This fly I
subsequently found to be the ‘Tsetse’” (pp. 90-91).

“On the 16th [June, 1871] we arrived at Madedita,*
so called from a village which was, but is now no more.
Madedita is twelve and a half miles from the Nghwhalah
Mtoni. A pool of good water a few hundred yards from
the roadside is the only supply caravans can obtain nearer
than Tura in Unyamwezi. The Tsetse or chufwa-fly, as
called by the Wasawahili, stung us dreadfully, which is a
sign that large game visit the pool sometimes, but must
not be mistaken for an indication that there is any in the
immediate neighbourhood of the water. A single pool so
often frequented by passing caravans, which must of
necessity halt here, could not be often visited by the
animals of the forest, who are shy in this part of Africa of
the haunts of man” (p. 213).

“Monday, the 2nd of October [1871], found us tra-
versing the forest and plain that extends from the Ziwani
to Manyara}, which occupied us six and a half hours.
The sun was intensely hot; but the mtundu and miombo
trees grew at intervals, just enough to admit free growth
to each tree, while the blended foliage formed a grateful
shade. The path was clear and easy, the tamped and
firm red soil offered no obstructions. The only provocation
we suffered was from the attacks of the Tsetse, or Panga
(sword) fly, which swarmed here. We knew we were
approaching an extensive habitat of game, and we were
constantly on the alert for any specimens that might be
inhabiting these forests” (p. 330).

“ Buffalo gnats and Tsetse were very troublesome on

* In Unyanzi: approximate position, according to author’s map,
34° 1’ E. long., 5° 80' S. lat.
+ In Unyamwezi: approximate position of Manyara, according to the

author’s map, 6° S. lat., 32° 25’ E. long.; the Ziwani (or pool) is about
fifteen miles N.H.

MARNO ON “SURRETA” IN SENNAAR. 153

this march * [Oct. 8, 1871], owing to the numerous herds
of game in the vicinity ” (p. 354).

46. 1873. F. Walker.

“CENTRAL AFRICAN BLoop-sucKING Fixes” (The Ento-
mologist, Vol. VI., pp. 327-328).

Notes on Stanley’s statements about blood-sucking
flies, quoted above [45].

The fly called by the native name “ chufwa” by
H. M. Stanley is considered by Walker to be Glossina
longipalpis, Wied.

Glossina fusca, Walk., is erroneously stated to be
identical with Gl. longipalpis, Wied.

47, 1873. E. Marno.

“User pen Ernr uss DER FiecEn (TUBAN) UND INSBE-
SONDERE DER SuRRETA AUF DIE HausTHiIERE SENNAAR’S”
(Mittheilungen aus Justus Perthes’ Geographischer Anstalt
tiber Wichtige Neue Erforschungen auf dem Gesammtgebiete
der Geographie, von Dr. A. Petermann. 19. Band, pp.
246-249).

Tsetse-flies stated to have been brought to the author
under the name “Surréta,” with three other species of
flies, two of which were true Tabanidee—by natives in
Sennaar, who consider the “Surréta” to be the cause of
the mortality among their cattle in the rainy season.

[The occurrence of a species of Glossina in Sennaar
needs confirmation : ¢ the fly identified as such by Marno
was perhaps a Stomoxys, or Hzematopota. |

According to Marno (p. 249), the statement that “in
certain parts of Africa at certain seasons domestic animals
are killed by the poisonous bites of flies, which in some
countries even make the keeping of particular domestic
animals impossible” has been constantly repeated since
the time of Agatharchides.

Marno speaks (pp. 246-247) of “the Baiida, a small
gnat [eine kleine Miicke|, which is found the whole year
through, in damp, swampy lowlands, but occurs in the
Charif in myriads, and gives human beings fever by its
bite.” He proceeds wrongly to identify it with the

* In Unyamwezi: approximate locality, from author’s map, 6° 10'S.
lat., 32° 12’ Hi. long.
TUG At line]:

154 LIVINGSTONE’S “LAST JOURNALS.”

Tsetse: “This Dipteron, named in South Africa Tsetse,
in Sennaar Surréta, is commonly looked upon as the cause
of the perishing of domestic animals in many regions, in
consequence of which its evil reputation has since the
earliest times extended even to Europe. It is a fact,
that in Sennaar in and shortly after the wet season the
mortality among human beings and domestic animals is
much greater than at other times... .” Possibly the
insect in question is a malaria-carrying gnat (Anopheles).

Marno considers that the bites of flies, whether called
Tsetse or Surréta, under which names the natives include
a large number of species, are only one, and “ perhaps
even a subordinate factor” in causing the mortality among
imported domestic animals, which occurs in certain parts of
Africa either throughout the year, or only during the Charif,
and is actually due to adverse climatic conditions.

48. 1874. Dr. Grube.

“User pie TsetserLiece ” (Hin-und-fiinfzigster Jahres-
Bericht der Schlesischen Gesellschaft fiir vaterléndische
Cultur (Breslau: G. P. Aderholz’ Buchhandlung), pp.
50-51).

Report of a lecture (with exhibition of a specimen
of a Tsetse-fly provided by Prof. Loew) delivered by
Dr. Grube at a meeting of the “naturwissenschaftliche
Section” of the Gesellschaft, held October 29, 1872.—
A résumé of Livingstone’s observations.

49. 1874. David Livingstone.

“THe Last Journats oF Davip LivINGsToneE, in
Central Africa, From 1865 to his Death.” Edited by
Horace Waller, F.R.G.S. (London: John Murray).

Livingstone’s experiment with camels and Indian buffaloes
introduced from Bombay.

Mikindany Harbour, north of the mouth of the
Rovuma River, March 29-30, 1866.—‘ The people have
no cattle, but say there are no Tsetse-flies. .. . The
adjacent country has large game at different water pools,
and as the whole country is somewhat elevated it probably
is healthy ” (Vol, I., p. 15).

“8th April [1866] We spent the Sunday at a village
called NYangedi [about three miles inland from Mikindany
Bay]. Here on the evening of the 7th April our buffaloes

CAMELS AND INDIAN BUFFALOES. 135

and camels were first bitten by the Tsetse-fly ” (Vol. L.,
py 47):

“17th April [1866].—I fear that my experiment with
the Tsetse will be vitiated [it seemed likely that the camels
and buffaloes would die owing to their being overloaded
by Livingstone’s sepoys], but no symptoms yet occur in
any of the camels except weariness ”* (Vol. I., p. 23).

On the north bank of the Rovuma River, 20th April,
1866.—“ Tsetse are biting the buffaloes again. Elephants,
hippopotami, and pigs are the only game here, but we see
none: the Tsetse feed on them” (Vol. I., pp. 24~25).

23rd April, 1866.— Bufialoes bitten by Tsetse again
show no bad effects from it: one mule is, however, dull
and out of health; I thought that this might be the
effect of the bite till I found that his back was so strained
that he could not stoop to drink, and could only eat the
tops of the grasses. An ox would have been ill in two
days after the biting on the 7th” (Vol. L., p. 26).

30th April, 1866.—On the north bank of the Rovuma
River, near Nachuchu: approximate position according to
Dr. Livingstone’s map, 11° 2’ §. lat., 39° 28’ EB. long.—
‘ Buffaloes again bitten by Tsetse, and by another fly
exactly like the house-fly, but having a straight hard pro-
boscis instead of a soft one t; other large flies make the
blood run. The Tsetse does not disturb the buffaloes, but
these others and the smaller flies do. The Tsetse seem to
like the camels best; from these they are gorged with
blood—they do not seem to care for the mules and
donkeys” (Vol. I., p. 30).

4th May, 1866. On the north bank of ihe Rovuma, west
of the N’konya River.—‘‘ The buffaloes were bitten again
by Tsetse on 2nd, and also to-day, from the bites of other
flies{ (which look much more formidable than Tsetse),
blood of arterial colour flows down ; this symptom I never
saw before, but when we slaughtered an ox which had been
'T'setse-bitten, we observed that the blood had the arterial
hue. The cow has inflammation of one eye, and a swelling

* “Dr. Livingstone was anxious to try camels and Indian buffaloes in
a Tsetse country to see the effect upon them.”—Epitor’s Norte.

+ Obviously a species of Stomoxys: the “other large flies” which
caused the buffaloes to bleed must have been horse-fiies (Tabanidie).
—H. E. A.

} Tabanide.—E. E. A.

156 LIVINGSTONE’S EXPERIMENT:

on the right lumbar portion of the pelvis; the grey
buffalo has been sick, but this I attribute to unmerciful
loading, for his back is hurt; the camels do not seem to
feel the fly, though they get weaker from the horrid
running sores upon them and hard work, There are no
symptoms of Tsetse in mules or donkeys, but one mule has
had his shoulder sprained, and he cannot stoop to eat or
drink” (Vol. I., pp. 33-34).

6th May, 1866. On the north bank of the Rovwma.—
“ Tsetse again, The animals look drowsy. The cow’s eye
is dimmed ; when punctured, the skin emits a stream of
scarlet blood.” .. .

“7th May.—A camel died during the night, and the
grey buffalo is in convulsions this morning. The cruelty
of these sepoys vitiates my experiment, and I quite expect
many camels, one buffalo, and one mule to die yet; they
sit down, and smoke and eat, leaving the animals loaded
in the sun.”

“7th May.—We are now opposite a mountain called
Nabungala,* which resembles from the north-east an
elephant lying down. Another camel, a very good one,
died on the way; its shiverings and convulsions are not
at all like what we observed in horses and oxen killed by
Tsetse, but such may be the cause, however. The only
symptom pointing to the Tsetse is the arterial-looking
blood, but we never saw it ooze from the skin after the
bite of the gad-fly as we do now.

“8th May.—We arrived at a village called Jpondé, or
Lipondé, which lies opposite a granitic hill on the other
side of the river (where we spent a night on our boat
trip), called Nakapuri; . . . One mule is very ill ; one
buffalo drowsy and exhausted ; one camel a mere skeleton
from bad sores ; and another has an enormous hole at the
point of the pelvis, which sticks out at the side. I suspect
that this was made maliciously. . . .” (Vol. I., pp. 35-36).

20th May, 1866. On the Loendi R., just above its
confluence with the Rovuma.—‘ The black buffalo is dead ;
one camel ditto, and one mule left behind ill. Were I not
aware of the existence of the Tsetse, I should say that

* From the map, the approximate position of Dr. Livingstone on this

day would appear to have been 11° 8’ §, lat., 38° 52’ HE. long.—on the N.
bank of the Rovuma.—H. E. A,

DOUBTFUL RESULT. 157

they died from sheer bad treatment and hard work”
(Vol. I., p. 42).

2nd June, 1866. Locality as above.—‘‘From the
appearance of the cow-buffalo, I fear the Tsetse is its
chief enemy, but there is a place like a bayonet-wound on
its shoulder, and many of the wounds or bruises on the
camels were so probed that I suspect the sepoys.” .. .

ca * * * *

. . . “the European house-fly chases away the blue-
bottle-fly in New Zealand. Settlers have carried the
house-fly in bottles and boxes for their new locations, but
what European insect will follow us and extirpate the
Tsetse? The Arabs have given the Makondé bugs, but
we have the house-fly wherever we go, the blue-bottle and
another like the house-fly, but with a sharp proboscis* ;
and several enormous gad-flies. Here there is so much
room for everything... .

. . . the wild hogs abound and do much damage, besides
affording food for the Tsetse: . . .”

3rd June, 1866. Same locality.“ The cow-buffalo fell
down foaming at the mouth, and expired. The meat
looks fat and nice, and is relished by the people, a little
glariness seems to be present on the fore leg, and I some-
times think that, notwithstanding the dissimilarity of the
symptoms observed in the camels and buffaloes now, and
those we saw in the oxen and horses, the evil may be
the Tsetse after all, but they have been badly used,
without a doubt. The calf has a cut half an inch deep,
and the camels have had large ulcers, and at last a
peculiar smell, which portends death. I feel perplexed,
and not at all certain as to the real causes of death” ft
(Vol. I., pp. 44-45).

* Stomoxys.—H. E. A.

+ According to Laveran and Mesnil (XXI., p. 47, note 7), Lingard
found Surra to be fatal to the buffalo in India (duration of the malady
125 and 51 days in two experiments), while Penning found it likewise
fatal to buffaloes in the Dutch E. Indies. The prolonged course of the
malady in these animals, as evidenced by Lingard’s experiments, would
seem to show that Livingstone’s buffaloes may really have been suffering
from Nagana at the time of their deaths,

As to camels, Laveran and Mesnil (op. cit., p. 48) write: “In the
dromedary Nagana develops pretty rapidly; in the Asiatic camel the
course of Surra is sometimes pretty rapid, and at other times very slow;

it may even last three years (whence the name Zei-barsa, signifying three
years, given to the malady of camels in certain districts of India).”
H, A.

158 OBSERVATIONS ON FLY-DISEASE IN DONKEYS.

26th June, 1866. Further west, along the south bank
of the Rovuma.—* My last mule died” (Vol. I., p. 61).

10th December, 1870. In the Manyuema country, to
the east of the Lualaba R.—<“Lion’s fat is regarded as
a sure preventive of Tsetse or bungo.* This was noted
before, but I add now that it is smeared on the ox’s tail,
and preserves hundreds of the Banyamwesi cattle in safety
while going to the coast ; it is also used to keep pigs and
hippopotami away from gardens: the smell is probably the
efficacious part in ‘ Heresi,’ as they call it” (Vol. IL., p. 87).

10th August, 1871. Near Mamohela, Manywema coun-
try.— Lion’s fat smeared on the tails of oxen taken
through a country abounding in Tsetse, or buiigo, is a
sure preventive ; when I heard of this, I thought that
lion’s fat would be.as difficult of collection as gnat’s
brains or mosquito tongues, but I was assured that many
lions are killed on the Basango highland, and they, in
common with all beasts there, are extremely fat ; so it is
not at all difficult to buy a calabash of the preventive,
and Banyamwezi, desirous of taking cattle to the coast for
sale, know the substance, and use it successfully (?)”
(Vol. II., pp. 149-150).

10th November, 1872. Near the Kalambo R. at the
south-east end of Lake Tanganyika.—< The donkey is
recovering ; it was distinctly the effects of Tsetse, for the
eyes and all the mouth and nostrils swelled. Another
died at Kwihara with every symptom of Tsetse poison
fully developed ” (Vol. II., p. 247).

With reference to the foregoing passage the editor
writes as follows :—‘‘ The above remarks on the suscepti-
bility of the donkey to the bite of the Tsetse-fly are
exceedingly important. Hitherto Dr. Livingstone had
always maintained, as the result of his own observations,
that this animal, at all events, could be taken through
districts in which horses, mules, dogs, and oxen would
perish to a certainty. With the keen perception and
perseverance of one who was exploring Africa with a view
to open it up for Europeans, he laid great stress on these
experiments, and there is no doubt that the distinct result
which he here arrived at must have a very significant

* This statement appears to have been obtained by Dr. Livingstone
from the Arabs..—E. EK. A

HOM@OPATHIC REMEDY FOR FLY-DISEASE. 159

bearing on the question of travel and transport” (Vol. IL.,
p- 247).

16th November, 1872. Near the Aeezy R., at the south
end of Lake Tanganyika.— After waiting some time for
the men I sent back yesterday to look after the sick donkey,
they arrived, but the donkey died this morning. Its
death was evidently caused by Tsetse bite and bad usage
by one of the men, who kept it forty-eight hours without
water. The rain, no doubt, helped to a fatal end ; it is a
great loss to me” (Vol. IT., p. 249).

50. 1874. Carl Mauch.

“Cart Mavcn’s Reisen 1m INNEREN von Stp-AFRIKA,
1865-1872. IV. Das Gesier zwiscnen Limpopo uND
ZAMBESI UND DIE RUINEN VON ZIMBABYE” (Mittheilungen
aus Justus Perthes’ Geographischer Anstalt iiber Wichtige
Neue Erforschungen auf dem Gesammtgebiete der Geographic,
von Dr. A. Petermann. Ergdénzungsband VIII, pp. 48-49).

[Translation.] ‘The Tsetse Fly.—A great drawback
to those regions is a small fly, in size and shape approaching
our house-fly, but somewhat paler in colour, of which the
natives assert that a single puncture is sufficient to kill
a horse, cow, or dog, while donkeys and goats suffer no
injury from it. Only one remedy appears to be effective,
and that is based upon homeopathic principles: the fly
itself, taken internally, makes the punctures innocuous,
as I have seen in the case of a dog, which after admin-
istering this remedy I took with me as far as the Lower
Zambesi and sent back again perfectly well with those
who had accompanied me. In the year 1868, when I had
an ox, a she-ass, and a dog with me and made experi-
ments with a solution of muriate of ammonia, the ox and
the dog perished, while the she-ass, to which I did not
administer any of the solution of this salt, after a few
days of rest suddenly attached itself to a troop of zebras
that were charging by, and ran off, without my ever
being able to catch it again” (p. 49).

51. 1875. J. P. Mégnin.

‘*MEMOIRE SUR LA QUESTION DU TRANSPORT ET DE
LINOCULATION DU VIRUS PAR LES MOUCHES” (1 planche)
(Journal de Panatomie et de Physiologie, &c. (Paris), XI,
pp. 121-133. Also in Journal de médecine vétérinaire
militaire, Paris, 1875, XII, pp. 461-475).

160 IMMUNITY OF GOATS ASSERTED BY MOHR.

52.

1876. E. Mohr.

“To THE VICTORIA Fats oF THE ZAMBEsI.” (Trans-
lated from the German by N. D’Anvers. London: Sampson
Low, Marston, Searle and Rivington.) Pp. 192, 244,
286-287, 290, 352-354.

Beneficial effects of ammonia, administered internally,
upon horses apparently suffering from Tsetse-fly disease.

“ Jennings’ party came to see me every evening... .
They had just come from the banks of the Ganyana
River, near the Zambesi, and three of their horses
showed symptons of the Tsetse sickness, the result of the
bite of the venomous insect of that name. I therefore
administered strong doses of eau de l’huis, or extract of
ammoniac; and all I can say as to the results of this
treatment is, that none of the horses died” (p. 192).

Goats siated not to suffer from Tsetse-fly disease.—
“Goats are especially suited for long journeys ; on account
of their elastic natures, they can accommodate themselves
to any circumstances, live upon any and the scantiest
food, walk immense distances, and above all, they never
suffer from the bite of the Tsetse-fly”” (p. 244).

In Matabeleland, south of the Shangani, May, 1870.—* I
marched with the greatest caution, as we might now at any
moment enter the district rendered dangerous to domestic
animals by the presence of the Tsetse-fly. I generally rode
on half a mile in advance of the rest of the party, or some
natives reconnoitred in front, so as to announce the appear-
ance of the poisonous insect at once” (pp. 286-287).

May 25th, 1870.—As above: from the author’s map
the precise locality is nearly due south of Wankies.—
“ Bokhis, who had ridden forward on Roland at twelve
o'clock, came back at three in the afternoon with the
news that the Tsetse were close upon us, and brought
half-a-dozen of these poisonous flies, which had settled on
his horse’s neck, as a positive proof of his assertion. We
could not therefore think of advancing any farther with
the horses and oxen. Latitude 19° 11' 8. was to be the
most northerly point reached by our waggon; and as
there was plenty of good grass and water here, we set
up a permanent encampment for the animals and their
attendants, resolving shortly to press on on foot for the
Zambesi, accompanied only by our baggage carriers ”

(p. 290). [Cp. 42.]

EDUARD MOHR’S OBSERVATIONS. 161

“Tn the districts recently traversed we had often met
with the Tsetse-fly, and in some places swarms had flown
out of the bushes, inflicting their sharp burning stings
upon every member of the party; but we were now *
passing out of its haunts, and its occurrence was rare,
although a short distance off there might still be thousands.
The insect flies rapidly, and stings as it settles on its
victim. The sharp sting penetrates easily through a
cotton shirt and flannel vest, but a severe momentary
itching is the only effect felt by a man; no evil results
ensue. I managed to catch several, which I put into a
hollow bird’s bone, closing the ends with resin, and subse-
quently gave them to my friend Dr. Hartmann,} of
Berlin, the African traveller.

“Some travellers, Vincent Erskine { amongst others,
have recently called in question the fatal results to
domestic animals of the bite of the Tsetse; but all the
natives of whatever race who have accompanied me on
my wanderings were agreed in accounting it poisonous,
and not one of them would have driven his own oxen or
horses into districts frequented by the Glossina morsitans.

“Moffat the missionary, who, accompanied by the
chief Mosilikatze, had wished to make his way to the
Zambesi in a north-westerly direction from the chief
kraal of the Matabeles, had to abandon his design
through losing all his oxen in the Tsetse district, and only
saved his waggons through the intervention of the chief,
who made hundreds of the natives yoke themselves to
them, and draw them out of the wilderness ” (pp. 352-353).

58. 1877. Dr. Hartmann.

Sitzungs- Bericht der Gesellschaft naturforschender
Freunde zu Berlin vom 17. Juli 1877, pp. 205-206.

Report of a lecture by Herr Hartmann.—{Transla-
tion.| ‘ Herr Hartmann further made some observations
on tite Tsetse-fly. At his request the late traveller
Eduard Mohr had brought home and handed over to
him to work out a number of specimens of the true
Glossina morsitans.§ They had been caught, it was
stated, with the hand, by Mohr’s people in the neighbour-

* July 6, 1870, near the Denzue R., south of the Shangani: approxi-
mate latitude, 18° 55’ 27” S.—H.H..A.

+ Cf. (53). t Cf. [38]. § Cf. [52].
M

162 ALLEGED HARMLESS TSETSE IN LOANGO.

hood of the Mési watunja, or Victoria Falls, of the
Zambezi, and had been excellently preserved in a piece
of hollow bone, coated with wax. On being taken out
they proved to be dried up, it is true, but still supple.
The lecturer found among the specimens of Glossina in
the Royal Entomological Museum here, two of the
Diptera collected by the German Loango Expedition,
which were likewise determined as belonging to Glossina.
Besides these there proved to be among the Loango
insects forwarded by Herr Falkenstein two more flies
preserved in alcohol, which in all respects presented the
characteristic features of the true Tsetse (Glossina
morsitans). On comparison with the Zambezi form all
that was noticeable was a slight difference in size.
While, for instance, the specimen from the first-mentioned
locality was 9 mm. in length, with a wing 10 mm. long,
the same measurements in the case of the Tsetse from
Loango were 10 and 11 mm. respectively. Moreover, in
the case of the former the abdominal bands were not so
very dark and yet appeared sharply differentiated one
from another, while in that of the latter these markings
looked darker and less clearly defined. For the rest, the
two specimens agreed in the structure of the proboscis,
and in the doubly-feathered arista (characteristic of
Glossina). The lecturer endeavoured to show this by
means of coloured drawings, magnified 100 times, of
the heads of the Zambezi and Loango Tsetse. According
to information supplied by Herr Falkenstein, the Tsetse
on the Loango Coast appears to be innocuous. No single
case has come to light there of the death of a domestic
animal caused by fiy-bites; while on the other hand
oxen and such-like animals are seen to perish there from
other diseases, which admit of ready diagnosis. This
agrees with the views first expressed by the lecturer
(Reise des Fretherrn v. Barnim in Nord-, Ost-Afrika, &c.,
Anhang XLI) as to the, if not absolute harmlessness,
still only slightly harmful nature of the Diptera known
under the name Surrita (Sorréta, Surréta, Serott, etc.),
as also of the Tsetse-fly in general. Subsequently the
traveller E. Marno expressed himself on the question in
a precisely similar manner (Reisen im Gebiete des blauen
und weissen Nil, Wien, 1874, p. 283).

EXPERIMENT WITH TREK-OXEN IN E. AFRICA. 163

“Herr Hartmann then stated his intention of making
further communications on this subject at one of the
autumn meetings.”

54. 1877. Rev. Joseph Mullens, D.D.

A New Rovte anp New Mone or TRAVELLING INTO
CrenTRAL AFRICA ADOPTED BY THE Rev. Rocer Price
IN 1876, described by Rev. Joseph Mullens, D.D.—Pro-
ceedings of the Royal Geographical Society, Vol. X XI.

“ Reflecting on these things, the Directors of the
London Missionary Society, when planning their expe-
dition to Lake Tanganyika, thought it worth while
specially to enquire into two points: (1) Could a route
be found to the north of the Wami River, on higher
ground, and free from the swampy levels found here and
there on the road from Bagamoyo? and (2) Was it
possible to employ on the entire line the waggon drawn
by bullocks, so common in the colonies of South Africa,
and that without risk from the Tsetse-fly 1” (pp. 235-236).

A trial expedition under Mr. Price, with a riding-donkey,
and a cart drawn by four oxen, left Saadani for Mpwapwa
(a distance cf 200 miles) on June 10, 1876, and arrived
and returned safely: no Tsetse-fly were met with.*

'“ As to the difficulty most feared of all, the Tsetse-fly,
which seems to be a trouble on the Bagamoyo road,
he [Mr. Price] says: ‘I regard the absence of Tsetse
between Saadéni and Mpwapwa as settled. I took the
four bullocks with me the whole way, and left thein at
Saadani on my return, apparently in perfect health.
Cattle, sheep and goats are to be met with here and
there along the whole route.’ Dr. Moffat gives it as his
experience that it is the inerease of population which
destroys Tsetse” (p. 241).

55. 1877. Thomas Baines.

“THe Gotp Reaions or SourH Eastern Arrica.”

(London: Edward Stanford. Port Elizabeth: J. W. C.
_ Mackay.)

Ammonia as a remedy for horses bitten by Tsetse.—
“T once used one pound of carbonate of ammonia, dis-
solved in a bucket of warm water, to wash four horses
during our passage through the Tsetse-fly. I cannot

* Cf. however [60].
m 2

164 AMMONIA AS A REMEDY FOR FLY-DISEASE.

assert that this saved them, but I have every reason to
believe so. Nine or more of our oxen died, but not a
single horse ” (p. 10).

Some of the hills near the Shasha River, Matabeleland,
known to be infested by Tsetse (p. 56).

“ The Tsetse is easily known by the manner in which
he folds his wings one over the other like a pair of
scissors, as we have before observed, giving him an
appearance of narrowness and length, differing from other
flies which settle with wings half expanded ; there are
some which close their wings at an angle of forty-five like
the roof of a house, but none of these are the Tsetse ”
(p. 57).

Tsetse-fly on the Macloutsie River, Matabeleland.*—“ On
Monday [October] 23rd [1871], we reached the wagons at
5 p.mM., and sent word to have the oxen kept away till
after dark, we set fire to the grass and to heaps of rubbish
to drive away the Tsetse, a few of which we saw. Gee
had seen a fly upon one of the horses; he touched the
place with. ammonia and the animal started with pain—a
sign that there was a puncture and that the ammonia had
entered it. I had about a pound of carbonate of ammonia,
and, dissolving the greater part in warm water, I had all
four of the horses washed. We observed some of them
flinch as if the remedy had found its way into punctures,
and it speaks well for our experiment, that up to the
present time, so far as I am aware, not one of those horses
has died. I had not enough to wash the oxen, but I
sprinkled them with tar-water in hope of keeping the fly
off, but without effect ” (p. 61).

Tsetse-fly on the Limpopo.—* We crossed [the Limpopo]
and outspanned [on the south side] on a place t reported
clear of fly, but after Mr. Biles had killed and brought
home a pullah, we saw a “fly ” on Jewell’s hat, but failed
to catch it. We brought up all the cattle and horses
immediately and rigidly examined them, tied up the
horses, posted a kafir to watch every insect that approached
them, and fired several shots to recall Gee; in a short
time he came followed by a kafir bringing another pullah,

* “T,ast outspan on Macloutsie River, lat. 21° 59’ 5”, long. 28° 44’ E.”
+ “Outspan on Limpopo, lat, 22° 35’ 31”, long. 28° 41’ 10” E., height
1,935 feet.”

BAINES’ OBSERVATIONS IN N. TRANSVAAL. 165

and I went out to meet him at a distance and make sure
that he brought no ‘fly’ upon his game. We saw one on
Plait’s fore leg, but could not kill it ; we washed the place
with ammonia, and saw probably the same fly on Mr.
Biles’s horse ; we tried to catch him with the edge of a
knife so held as to cast the narrowest possible shadow
a little way from the insect, but the Tsetse was too quick
even for this ; at last Biles struck him to the ground, and
T secured him in an envelope to be sent to London”
(pp. 63-64),

A “granite hill covered with bush and infested by the
dreaded fly,” Nov. 2, 1871, on the south bank of the
Limpopo, nearly opposite the “ Tslagool Hills (the Silika
or Siloquam of the maps)”: approximate position, lat.
22° 50! 4”, long. 28° 22' 40" (p. 65).

Tsetse in the Northern Transvaal.—* Friday, Nov. 3rd
[1871]... four or five men and boys joined us, they
told us the pan in front was called Madlala, and a day’s
journey beyond it was ‘Schimmel Paard Pan,’ there was
fly between them, but no water, and there is fly also
beyond Schimmel Paard Pan. But we must leave it
before dark so as to be able to get into a definite track,
and yet not so soon as to rush into the fly until it has
retired for the night. It is a day’s journey to Maghali-
quain River, through which we must not cross as there is fly
between it and Madzalana River, but after this we are
past all danger, and reach Matlalas in two days and

’ Makapans in four. From Maghaliquain there is fly to the
west, between us and the Limpopo” (pp. 65-66).

“The Matchopong mountains were visible to the
north-west, and as they are also seen from the Ba
Mangwata road they formed a landmark to test the
correctness of our longitude. Our last night’s track
through the fly country had been about 13 miles, and
during the afternoon and evening we made about 10 more.
The night was dark and cloudy, preventing any observa-
tion for latitude, but affording us additional security
against the insect pest” (p. 66).

“On Sunday, November 5th [1871], we inspanned
about half-past 5, so as to get into the definite road before
dark, and about seven we descended into a sandy tract
with dense bush haunted by the Tsetse. . . . At dawn on

166 “PATCHES OF FLY.”

Monday we trekked again, a steady shower protecting us
as we laboured on through the infested district, and in
10 miles and 3 furlongs we reached the west bank of
the Maghaliquain or Fierce Crocodile River, called by
the Boers the Nyl. . .. Our cattle would not eat the
poor and scanty grass,* but crossed the river to the better
pasturage they saw on the other side, but unfortunately
the ‘ fly’ was there and we had to drive them hack again.
. . . After sunset on the 9th we crossed the river and
trekked up its eastern bank for some distance, and passing
through the last patch of ‘fly,’ outspanned upon an open
plain 9} miles from our last camp. . . .” (pp. 66-67).

“We had now cleared the infested parts between the
Blauw Berg and the Hang Klip mountains, . . .” (p. 67).

[Near Nylstroom].—*TI halted also at the farm of
Theunis de Klerk, who told me they now knew where to
ride their horses with safety hetween the patches of fly ;
they also have safe or inoculated oxen and even ride their
horses in; they will not tell their medicine, but charge an
ox for making a horse safe; they told me they thought
the fly was a curse that was being removed from the
land ” (p. 68).

. . . I was obliged [end of January, 1871] to pro-
ceed without delay to Matabililand, where the King Lo
Bengula confirmed most fully the concession he had
already made me in the Northern Gold Fields, and gave
me liberty to come out of his country by a more direct
road, southward through the Tsetse-fly country into the
Transvaal. This might be considered no great boon, as
cattle once ‘ stuck’ by the fly are doomed to almost certain
death. But the Tsetse, though occupying large tracts of
country, does not completely overspread it, but leaves
parts which are known to various hunters, and which
serve as channels by which a course may be steered with
some chance of escape from the deadly pest. Unfortu-
nately, I could not obtain a skilful pilot, and came in
contact with one patch of fly, by whose stings I lost nine
oxen, but having a pound of carbonate of ammonia I dis-
solved it in a bucket of warm water and washed all the
horses. I am not prepared to affirm that this was the

* “Drift of Maghaliquain, lat. 23° 27' 20”, long. 28° 54’ 40”, ninety
miles, one furlong, sixty-four yards from the Limpopo.”

ROAD FROM PRETORIA TO DELAGOA BAY. 167

actual saving remedy ; but none of them died, and I think
the presumption is in favour of the ammonia ” (pp. 79-80).

‘We heard from a farmer we met during the day that
Andreas Duvenage (commonly called Devenaar) knows
of a safe road through the fly, between Blauwberg and
Zoutpansberg” (pp. 80-81).

“ Duvenage lives 18 miles to the north [of Mara-
bastad] and has the best known road through the Tsetse ;
he crosses the Limpopo at Commando Drift, meeting only
one patch of fly, which he rides through in the night”
(p. 84).

‘“‘ The fly leaves a country if the game is driven out or
the bush cut away, but returns if the conditions again
become favourable to its existence” (p. 89).

Road from Pretoria to Delagoa Bay.—‘ Here it may
be considered that the descent of the Drakensberg proper
has been completed, and the lower ranges of the Makond-
shwa and Lobomba have to be crossed. The road, however,
passing through by a tolerably level poort or valley south
of the Umbolosi or Dundar River, and immediately after
passing the last range, the Tsetse-fly, fatal to domestic
cattle, and the fever, no less deadly to man, await the
traveller. Fortunately there is only about 40 miles of
this unhealthy country ; but the fact should be known, in
order that the risk may be guarded against by pushing
through it as rapidly as possible, and at night or during
a cold day, when the fly is dormant ” (p. 108).

“ Tt must be remembered, however, on the other hand,
that from the Lebomba (sic) mountains to the port, a
distance of 30 or 40 miles, lies the tract of low country
from which Delagoa Bay derived its not undeserved
reputation for unhealthiness. A considerable portion of
this strip is infested by the Tetse-fly, and a point to be yet
proved is whether this belt of fly is sufficiently narrow
to be passed through in one night. . . .” (p. 109).

Appearance of Tsetse-flies ; symptoms of the disease in
cattle and horses ; remedies.

‘“‘ The Tsetse is little more than half an inch long, and
rather more slender than a common house-fly. The abdo-
men is marked with transverse stripes of yellow and
dark chestnut fading toward the centre of the back, so
as to give the idea of a yellow stripe along it; the belly

168

RESTING-POSITION OF TSETSE.

livid white, the eyes are purplish-brown, and the wings, of
dusky glassy-brown colour, slip one over the other, just as
do the blades of a pair of scissors when closed—so that
the Tsetse at rest on man or animal may infallibly be
known by this one token,

“No fly which rests with its wings half-expanded, like
the house-fly, or closed together Jike a pent-house roof, can
be the Tsetse ; but if one is seen in which the wings
exactly overlap—one lying flat upon the other—that is
‘the fly’. . . its sight and smell seem to be keen; its
flight straight and rapid. To speak of either its sting or
its bite would convey an erroneous idea. The Dutch
colonists say it ‘sticks, and this is certainly more correct,
as it first pierces the skin with its lancet, and then injects
a fluid (poisonous to oxen, horses, and dogs) to thin the
blood before drinking it. Men, mules, donkeys, sheep,
goats, and wild game are believed to be unaffected by the
virus. I, in common with other travellers, have been
stuck time after time with impunity. Mules, partaking
of the equine nature, are not always secure from dangers
to which the horse_is liable, and Mr. St. Vincent W.
Erskine doubts the safety of the donkey on the south-east
coast.

““My friend, Mr. Henry Hartley, the well-known
hunter and pioneer of the gold-fields, has kindly sum-
marised the symptoms exhibited by a fly-stuck ox as
under :—

“1st. The hanging of the ears, general languid appear-
ance, sometimes watering at the eyes. 2nd. Roughness
of the coat, the hair rising on end. 3rd. Feeding vora-
ciously, even to repletion, without improving the condition,
and standing panting in the heat of the day. 4th.
Occasional swelling at the gullet. 5th. Continual wasting
and pining away (but sometimes an ox may improve in
condition, and show no symptom of having been stuck for
two or three months, or till the first cold rain falls). 6th.
An ox slightly stuck goes on wasting till the skin sits fast
on the backbone. After this there is no hope; but if
severely stuck he dies before it can take place. Mr.
Hartley did not notice running at the nose or other
unusual discharge. If the ox is worked he will show
weakness in the loins. 7th. When the beast is skinned

FLY-DISEASE IN CATTLE AND HORSES. 169

after death the puncture of every fly can be seen on the
inside of the skin, and on the flesh is a ring of yellow
mucus, nearly as large as the palm of the hand, similar
to the mark that surrounds the bite of a snake, but
smaller.

“ A bullock belonging to Christian Harmoe was stuck
in May; he worked for 500 miles, then began to show
symptoms, and died in September. Some of my own—
stuck during the passage through the fly country in
September, 1871—worked about 300 miles, and died at
Mr. Hartley’s in January of the next year.”

* # “ * #*

“Mr. Hartley adds :—‘ Horses swell about the eyes,
nostrils, and testicles, where generally the wounds are
most numerous, they pine away, and their hair stands on
end, or is reversed ; cold rain also hastens their death.’ My
friend, the late Joseph Macabe, being incredulous, deli-
berately rode a valuable hunter right into an infested
tract, and returned to the outspan, where his steed died in
a few hours, and the pool is now called Schimmel Paard’s
Pan, or the pool of the dapple grey horse. Mr. Hartley’s
splendid grey elephant charger, ‘Camelbuck,’ died twenty
days after he was stuck ; ammonia was applied, and he
was led to stand in cold water, which is said to be some-
times effectual, but in vain. My own horse, ‘ Vegtmann,’
stuck, I believe, in October, 1869, travelled more than
1,100 miles, and died early next year in Pieter Maritz-
burg. Dogs pine and waste as oxen do. We lost some,
but one, a rough hairy bitch, seemed recovering; the
new hair that grew on places where she had apparently
been stuck was coarser and greyer than before.

“ The fly is extremely local, and extensive districts in
which it prevails may be passed through by the aid of
guides, who know the ‘patches’ of fly, just as a pilot
knows the shoals of an estuary ; but it shifts with the
migration of game, and, therefore, the knowledge of the
guide ought to be recent.

“ The hunters endeavour to, keep.it from .their road, in
Matabililand, by burning the dry grass, as they come out
at the end of the season. And when the Boers made their
celebrated ‘ Commando path,’ they destroyed the bush for
several hundred yards on either side as they approached

170 BAINES ON PROPHYLACTICS AND REMEDIES.

the Limpopo. If a belt of fly cannot be avoided, it may
be passed through, if not too wide, in the night, or on a
cold rainy morning; but the last is a dangerous experi-
ment, for should the sun break through the clouds it
rouses the insects with increased vigour and activity from
their torpor ; and it is well known that on a hot day all
poisonous creatures are more virulent and deadly.”

* # * * *

“Tam not aware that any certain remedy is known.
The native doctors inoculate oxen by giving them the fly
itself, mixed with herbs. The poor beast suffers dread-
fully, and is brought almost to the point of death, but
when it recovers is believed to be Tsetse proof.

** All young animals, while living on milk, are safe.
Some of the tribes living on the borders drive the calves
into the fly during the day, and bring them out to be
suckled morning and night. This is supposed also to
render them secure during the rest of their lives.

“Tt would be a great boon if any composition capable
of being sprinkled or syringed over the animals—innocuous
to them and disgusting to the fly—could be discovered ;
tar, ox dung mixed with milk, the kidneys of the meer
kat, etc., have been recommended, but carbolic acid would
perhaps be more effectual, diluted with water, and applied
by syringe or the rose of a watering-pot.

“Mr. Hartley tried a decoction of the bark of the
roots of the wittegaat boom, or motlopre, I believe, with
some success ; and there are Boers who profess to be able
to cure an animal recently stuck. Their fee is one good
ox for saving a horse. .

“While passing through the fly, in 1871, I mixed
about a pound of ammonia with a bucket of warm water,
and washed all four of our horses. We noticed that they
flinched, probably as the liquid entered the punctures.
None of them died ; and though I would not affirm on one
experiment that the ammonia saved them, I think it
highly probable such was the case.* Unfortunately, I
had not enough to wash the oxen with. We tried to

* “June 27,1874. I hear that during last season a hunter, on entering
the fly country, dosed or washed his horses continually with ammonia,
and has brought them all out safe, but very weak, probably from the
combined effects of the poison and its antidote.”

TSETSE AND TRANSVAAL GOLD-FIELDS. 171

restore tone to the blood with muriate of iron when I
reached Mr. Hartley’s farm, but it was then too late.

“‘Mr. Saunders, of Maghaliesberg, saved a horse with
Croft’s tincture of life, which contained ammonia, and two
oxen with Perry Davis’s pain killer. They stood for three
or four days with foam issuing from their mouths, as if the
poisonous matter were being thus ejected. After this
they began to eat voraciously and recovered their condi-
tion. Perhaps if milk could be given to animals as soon
as they are stuck they might recover ” (pp. 151-154).

“ The dangers, however, of the Delagoa Bay route [to
the Transvaal gold-fields] would be more efficiently met
by the construction of a good road, with the bush cleared
away as far as possible on either side, in the fly country,
leaving the unhealthy lowlands as soon as possible, and
climbing any elevated ridge, the course of which (even at
the cost of considerably increased distance) might be
followed to the highlands, so that man and beast might be
as little as possible exposed to fly or fever” (p, 154).

“In the Colonist of June 19th we are informed that
Mr. Isidore Alexandre has brought an ox wagon down
from the gold-fields to Delagoa Bay in nine days, and
others are to follow. There has been no loss among the
oxen. I would fain hope the best for Mr. Alexandre,
but I shall watch with great interest the future history
of these animals. We know that in cold weather the
fly is less virulent, but it never ceases to be dangerous ”
(p. 155).

“THE Gotp Recions or Sours-EasteRN AFRICA.”
Distances and Routes.

‘Tables of Distances by Trocheameter—Latitudes and
Longitudes (astronomical or computed)—Heights above
the sea-level, and other observations, from Port Natal to
the Ganyana River, vid Hartley’s, or the Potchefstroom,
Rustenberg, and Tati route, free from Tsetse-fly ; and
return route through the Fly country, vid Magholiquain
River and Makapan’s Poort. During the years 1869,
1870, 1871, and 1872. By Thomas Baines, F.R.G.8.”
(p. 164).

“The Tsetse-fly extends from south-east of Maghali-
quain [River] to Shasha” (p. 171).

172 TSETSE IN TRANSVAAL AND ON R. CONGO.

“Limpopo River, south side . . . Lat. 22° 37’, long.
28° 38’. Height in feet, 1,935. Granite quartz, palms,
thorns, castor-oil plant, Tsetse-fly” (p. 171).

“ Pass between Blaaweberg and Zoutpansberg through
Fly country, probably to Commando Drift, Limpopo River,
enquire for safe road of Der Venage or other hunters. . . .
Lat. 22° 20’, long. 29° 10’. Height, 1,700 or 1,800”
(p. 173).

Tsetse-fly (and fever) stated to be prevalent to the
south-east of Pretoriaskop, Transvaal : approximate posi-
tion, lat. 25° 12’, long. 31° 31' (p. 182). [Period to
which this refers is probably the early ’seventies—up to
1874. ]

“The Tsetse-fly abounds in this low country nearly to
Pretoriaskop. Mr. Macdonald’s expedition lost, I think,
114 oxen, and Mr. Arrowsmith reports a loss of fourteen.
Donkeys are generally safe, but not quite so here” (p. 183).
[This refers to a route from Pilgrim’s Rest to Delagoa
Bay. |

Patch of “fly” referred to in the author’s itinerary
of his route from Walvisch Bay to Victoria Falls and
Logier Hill, Zambesi River, near Matietsie River ; lati-
tude 18° 20’ (p. 187).

Broad red sand hill, with mopani trees, and infested
by fly ; latitude 18° 30’ (near Victoria Falls) (p. 187).

56. 1878 H. M. Stanley.

‘THROUGH THE Dark Continent, or The Sources of
the Nile, Around the Great Lakes of Equatorial Africa,
and Down the Livingstone River to the Atlantic Ocean”
(London : Sampson Low, Marston, Searle and Rivington).

Gad-(horse-)flies and Tsetse on the numerous low
islands below Rubunga, on the Congo, and attacking the
members of the author’s expedition descending the river
in their canoes and boat, February, 1877 (Vol. II.,
p. 292). Position of Rubunga, according to the author
(ibid. p. 281), 1° 40’ 44” N. lat., 21° 4’ E. long.

From the author’s subsequent remarks, the expedition
seems to have been attacked by horse-flies and Tsetse
during its further course among the islands in this
portion of the Congo, at least as far as 1° 22’ 15" S,. lat. :
February, 1877 (Vol. IT., p. 310).

57.

58.

59.

DR. DRYSDALE’S HYPOTHESIS. 173

1878. F. B. Fynney.

** THE GEOGRAPHICAL AND Economic FEATURES OF THE
TRANSVAAL, THE New British DEPENDENCY IN SoutH
Arrica” (Proceedings of the Royal Geographical Society,

Vol. XXIT.).
*‘Zoutpansberg forms with Waterberg the northern
boundary of the State, lying to the N.E.... Many

parts of this district, as well as Waterberg, are infested
with the Tsetse-fly; but there is scarcely need to
attach so much importance to this fact as is commonly
done, because the fly is merely a temporary and
ephemeral scourge, and always disappears with the
large game.

‘Many parts, which six years ago were known as Fly
country, are now entirely free, and therefore it may be
fairly hoped that the extinction of the pest is only a
matter of time” (p. 120).

1879. J. J. Drysdale, M.D.

“On THE GERM THEORIES OF INFECTIOUS DISEASES”
(Proceedings of the Literary and Philosophical Society of
Liverpool, No. XX XIII., p. 13, note).

An early suggestion pointing to the true part played
by the Tsetse-fly in connection with fly disease. After
referring to the then recent discovery by Dr. Manson of
the transference of Filaria sanguinis by mosquitoes, the
author adds the following footnote :—

“Tt is possible that we have here an explanation of
the destructive power of the Tsetse-fly, for it may be the
intermediate host of some similar blood-parasite ; or it
may be the carrier of some infective poison. It is highly
improbable that any mere poison or venom should exist
so powerful as to cause the death of a large animal in
such small dose.”

1879. Captain J. F. Elton.

“'TRAVELS AND RESEARCHES AMONG THE LAKES AND
Mountains OF Eastern AND CENTRAL ArFrica” (London :
John Murray), pp. 278, 404.

August, 1877.—Tsetse abounding at Livingstonia, at
the south end of Lake Nyasa (p. 278).

Still a question whether the deaths and sickness of
cattle on the Unyamyembe route from the coast to Lake

174 TSETSE-FLY ON ROAD TO MPWAPWA.

Tanganyika is “due to Tsetse or to the rank wet vegeta-
tion of the lower country” (H. B. Cotterill, in above
volume, p. 404).

60. 1879. Sir Rutherford Alcock, K.C.B.

ADDRESS ON THE OPENING OF THE 48TH SESSION OF
THE RoyAL GEOGRAPHICAL Society, November 11, 1878
(Proceedings of the Royal Geographical Society and
Monthly Record of Geography. New Monthly Series.
Volz]. acum

Destruction by the Tsetse-fly “on the road to Mpwapwa”
of the draught cattle belonging to a parity sent by the London
Missionary Society to Lake Tanganyika. [Cf. 78.]

61. 1879. Dr. Laws.

“ JOURNEY ALONG Part OF THE WESTERN SIDE OF LAKE
Nyassa, In 1878” (Proceedings of the Royal Geographical
Society and Monthly Record of Geography. New Monthly
Series. Vol. I.).

Cattle-destroying fly (? Tsetse) in the valley of the
Limpassa (an affluent of the River Lucia, which flows into
the Lake at Makambira).

“One place here, called Mudaye, presents what is
probably the best place we have seen in the district, with
an elevation of 800 or 900 feet above the lake. There is
the great drawback, however, that a fly is reported to
exist in the lower part of the plain, which if not the
Tsetse, resembles it by its bite killing cattle. The speci-
men of the fly shown to us was not the Tsetse, but belongs
to the blood-sucking tribe. The Mangone had at one
time lived in the hollows, but removed further up the
hills to save their cattle” (p. 317).

62. 1879.
“THe Dar-es-SaALaaM Roap” (Proceedings of the Royal
Geographical Society and Monthly Record of Geography.
New Monthly Series. Vol. I. (Geographical Notes), p. 129).
Tsetse-fly absent from the first forty miles of the road

inland from Dar-es-Salaam.
‘“Mr. Beardall, formerly of the Universities’ Rovuma
Mission, left England, on the 28th of November [1878],
to take charge of the works in connection with the road
‘now making from Dar-es-Salaam to the interior of Eastern

BOER MODE OF CROSSING FLY-BELTS. 175

Africa. . . . Eight out of the ten bullocks brought from
Madagascar are doing good service on the road, and it is
therefore clear there is no Tsetse-fly on the forty miles
already traced of the route; ... .”

63. 1879.

“THe ArFricAN Insect Scource. Tue Tsrrsz-Fiy”
(The Journal of Applied Science, Vol. X., May, 1879,
pp. 74-75).

[I have not seen this paper. ]

64. 1879. Lewis Hornor.

The Times, Feb. 25, 1879.—A letter recommending
the Boer method of crossing fly-belts at night (quoted by
Westwood in Oates’ “ Matabele Land and the Victoria
Falls,” First Edition (1881), Appendix, p. 364; Second
Edition (1889), p. 388).

“ Having hunted in the African fly country and seen
many horses and oxen die of the bite, against which no
external application is, I firmly believe, any safeguard, I
venture to call attention to the precautions adopted by
the Boer elephant hunters in the interior. The Tsetse
inhabits narrow and clearly defined strips of country,
familiar to all natives, and readily evident to strangers.
On approaching one of these ‘fly belts’ (so called) a halt
is made, and inspanning again at sundown the Boer treks
through at night in safety. I only remember one case of
mishap, when, in crossing a belt near the confluence of
the Chobé and Zambesi, two or three oxen out of nearly
forty were bitten, and that, if my memory serves me, on
a bright moonlight night.”

65. 1879. F. Karsch.
Zeitschrift fiir die gesammten Naturwissenschafien, LIT.
Band, p. 381.
Glossina longipalpis, Wied. taken at Chinchoxo, in
Loango, by Dr. Falkenstein.* [Chinchoxo, or Tschin-
tschdtscho, is a town on the coast about 100 miles north
of the mouth of the Congo. |

€5a. 1879. Dr. J. Falkenstein.

‘*Dr— Loanco - Exprpirion.” Zweite Abtheilung,
(Leipzig: Paul Frohberg), p. 84.

* Cf. [65a and 8o].

176 ABUNDANCE OF TSETSE NEAR KAREMA.

Eleven oxen, purchased in Loanda.and taken to
Tschintschitscho to be trained as baggage-animals, soon
died (1875). But although several specimens of a species
of Tsetse-fly were taken there, the deaths of the animals
are not considered to be due to the insect, but rather to
change of climate and food, since the symptoms were
carefully watched throughout the course of the disease.*

66. 1880. Capt. F. F. Carter.

“Captain Carter’s rast Marcu tn Centra AFRICA”
(Proceedings of the Royal Geographical Society and Monthly
Record of Geography. New Monthly Series. Vol. IL,
1880, p. 762).

[Extracts from the diary of the late Capt. F. F. Carter,
who conducted the Belgian Elephant Expedition from
Dar-es-Salaam to Mpwapwa [ef. 79]. Capt. Carter was
killed at Kasogera, on June 23rd or 24th, 1880, soon
after starting from Karema, on Lake Tanganyika, on his
return march to the coast. |

Tsetse-fly near Karema (German E. Africa).

“On 13th June [1880] started together from Karema.

.. On 15th ... reached place called Marimba,
where ground is marshy, lots of bad water; food for
elephants in dry season; once a large village, now de-
serted ; Tsetse-fly in thousands.”

* ‘ * * #

“Thursday, 17th.—Bitterly cold morning ; passed
through a sort of gorge in the mountains; very heavy
dews at night, and grass high and very wet, showering
dew on us; ... Thousands of Tsetse, nearly driving us
mad. On passing River Fuma we entered Mongway
district, this river dividing Fipa from it.”

67. 1880. Dr. Emil Holub.+

“JOURNEY THROUGH CENTRAL SouTH AFRICA, FROM THE
Diamonp Fietps to THE Upper Zampesi” (Proceedings
of the Royal Geographical Society and Monthly Record of
Geography. New Monthly Series. Vol. II., p. 174).

(See author’s map, inserted after p. 400, op. cit.)

Tsetse-fly in Matabele Land (1875), to the south of the
Victoria Falls, east of “Sandy Pool Plateau” (approxi-
mate position 19° 10' 8. lat., 26° E. long). The author

* Cf. [65 and 80). + Cf. [75]:

SIR BARTLE FRERE ON FLY-BELTS. 177

is proceeding north-west, to Panda ma Tenka and the
Chobe.

“T thus entered a high sandy plateau, a thickly
wooded forest where there are no rivers, but thousands of
pools, the greater number of which only contain water
after rain, and which I named the ‘Sandy Pool Plateau.’
Where I crossed, it is 102 miles in width. . .°. Here in
coming from the south we met with buffaloes, elephants,
and rhinoceroses for the first time. ... I found the
limits of the Tsetse were from ten to fifteen miles east of
the direction which I took.”

* * * * *

“[August, 1875.|—Eight miles from Pandama [?.e.
Panda ma Tenka] I crossed the first boundary of the
Tsetse, coming afterwards to a part free from Tsetse, and
the second time entering a part infested by it about
twenty-one miles south of the Chobe Junction” (p. 174).

68. 1880. G. Macloskie.

“THe Prososcis or THE Hovusr-Fiy” (The American
Naturalist. Vol. XIV., pp. 153-161, figs. 1-3).

69. 1881. The Right Hon. Sir Bartle Frere, Bart., G.C.B.,
G.C.S.1., &c.

“On Temperate Sourn Arnica” (Proceedings of the
Royal Geographical Society and Monthly Record of
Geography. New Monthly Series. Vol. IIT.).

Mode of crossing the Fly-belt south of the Zambesi, near
the Victoria Falls.

“Mr. Humphery, a young traveller now present, who
visited the Victoria Falls a few seasons ago, gives the
following account of the mode of passing the Tsetse-fly
belt on the south bank of the Zambesi about seventy
miles above the Falls.

“ Leaving the last halting-place free from Tsetse in the
evening, they travelled all night to avoid the insect, and
before morning reached a narrow strip of country free
from fly but without water, though there was grass for
the oxen. The next night a shorter march brought them
to the river in time for the oxen to drink and return back
to the spot free from fly before daylight.

“Tt must be remembered that the Tsetse-infected
tract varies from year to year with the movements of the

N

178

TSETSE DISAPPEARING WITH BIG GAME.

large game, and according to the nature of the season ;
and that the area where the fly is fatal appears to be
diminishing rather than increasing” (p. 14).

In the discussion on Sir Bartle Frere’s paper (read
November 22, 1880)—‘Sir Fowett Buxton observed
that the subject of Sir Bartle’s paper could not fail to be
of great interest to all who felt anxious about the con-
tinuance of the work of exploration in Central. Africa.
. .. One matter of some hopefulness was the fact that
the Tsetse-fly seemed to be gradually receding before the
advance of civilisation. Efforts had been made to con-
struct a road from the coast into the interior, and the
grand attempt made by the King of the Belgians to
introduce the elephant as a beast of burden should not be
forgotten. Some attempts had been made also to intro-
duce wheeled traffic, though the fly had hitherto stood in
the way. He could not, however, but hope that better
success was in store in the future, since it appeared that
the Tsetse scourge disappeared wherever the large game
was driven back.

“Sir Bartte Frere said with regard to the possi-
bility of getting across the fly-belt, the testimony of the
chief hunters and traders went to show that the Tsetse
certainly did, to a great extent, follow the movements of
the great game ; and also that as the country became better
known, particular points were discovered by the guides
where even during the day-time, in the immediate vicinity
of the bush infested by the fly, there were cleared spots
where the fly never touched a horse, and that those who
had been much in the fly districts were able to discover
routes through the infested belts by which horses could
with great care be conveyed; and of course if that was
the case on a small scale, it might gradually be that the
means of carrying animals subject to be bitten by the
Tsetse through the infested belt would increase. Mr.
Humphery had told him that traders had conveyed a
wagon across the river [Zambesi], and it had been
purchased by a chief on the other side, but unfortunately
he was not beyond the region so fatal to oxen. However,
there appeared to be very little doubt that there was a
region perfectly free from the fly at no great distance on
the northern bank, and within practicable reach” (p. 19).

TSETSE ON TIOGE RIVER. 179

70. 1881. Pére Duparquet. ’

Proceedings of the Royal Geographical Society and
Monthly Record of Geography. New Monthly Series,
Vol. IIL., p. 43 (Geographical Notes.—The River
Okavango).

Tsetse-fly on the Okavango, or Tioge or Tonke River, at its
junction with Lake Ngami.—* He [Pére Duparquet] would
have preferred, he says, to begin his description from Lake
Ngami, instead of devoting his attention to the portion of
the river between Libébé and the country of the Amboellas
(in about 16° 8. lat.), but few particulars were procurable,
as, Owing to the presence there of the Tsetse-fly, this part
of the river is but little frequented.”

71. 188r.

“Dr. Krrx’s Vistr to tHe Dar-us-Sataam Districr
tN East Arrica” (Proceedings of the Royal Geographical
Society and Monthly Record of Geography, Vol. III., pp.
308-309 : Geographical Notes).

Belt of Tsetse-fly wider and more continuous inland
from Dar-es-Salaam than farther to the north; fly found
40 miles from the coast.—“ Early in the present year Dr.
Kirk made an interesting journey, in company with
Captain Foote, R.N., along the road recently made at
the private cost of Mr. Mackinnon and Sir Fowell Buxton
towards the interior of East Africa from Dar-es-Salaam.
- .. The presence of the Tsetse-fly in the country 40
miles from the coast further renders the employment of
horses or bullocks as beasts of burden out of the question ;
the belt of fly-country is, in fact, here wider and more
continuous than it is further to the north.”

72. 1881. ‘Major Serpa Pinto.

‘How I Crossep Arrica: From the Atlantie to the
Indian Ocean, through Unknown Countries ; Discovery
of the Great Zambesi Affluents, ete.” Translated from
the author’s manuscripts by Alfred Elwes (London :
Sampson Low, Marston, Searle and Rivington), Vol. IT.,
p. 81.

October, 1878.—On the right bank of the Zambesi,
between Itufa and Sioma, Barotseland (approximate geo-
graphical position, 16° 30'S. lat., 23° 30' E. long) :—

“TI started whole flocks of heathcocks, quails, and

a

180

73.

74.

TSETSE AT VICTORIA FALLS.

Guinea fowl* (Numida meleagris), which were in greater
numbers than I had hitherto seen in Africa. I also found,
to my sorrow, that the Tzee-tzee-fly was quite as abundant
as the birds ; it troubled me excessively in the forest with
its sharp sting, which, however painful, is not dangerous
to man; and these insects were so numerous and pursued
me so inveterately that, after I had again got into the boat,
I had for some time to do battle with them.”

1881. F. Oates.

« MATABELE LAND AND THE VicTorIA Fatis” (London:
C. Kegan Paul & Co.), Ist edition; Plate G, fig. 2;
Plate H, figs. 5, 5a, 5b.

Tsetse-fly at the Victoria Fails (1873).—“ He [the Hon.
G. C. Dawnay] showed me some little sketches he had
made fof the Victoria Falls], but said it was almost
impossible to draw on account of the flies. The Tsetse-fly,
which kills everything, except men, wild beasts, and
donkeys, swarms there, and bites so furiously that your
hands and face are puffed up in no time” (p. 38).

“ He [a Transvaal Boer, named Lee] has tried donkeys
in the Tsetse-fly country, but the fly has always killed
them” (p. 48).

Notes by J. O. Westwood (pp. 363-365). See 74.

1881. J. O. Westwood.

Oates’ Matabele Land and the Victoria Falls, 1st ed.,
Appendix, pp. 363-365, Plate G, fig. 2; Plate H, figs. 5,
5a, 5b (2nd ed., 1889, pp. 388-389, Plate VILI., fig. 2;
Plate IX., figs. 5, 5a, 5b).

Copy of the letter by Lewis Hornor, in the Times, of
Feb. 25, 1879 (see 64) (1st ed., p. 364).

“The African traveller Hildebrandt recommends
strongly, in the Korrespondenzblatt der afrik. Gesellschaft,
the use of petroleum for those travelling in the tropics, as a
protection against insects. Occasional applications to the
face and hands ensured entire freedom from mosquitoes, and
the same method sufficed to preserve horses and cattle against
the deadly attacks of the Dondorobo gad-fly, which so often
cripples the movements of the explorer” (1st ed., p. 364).

Coloured figure (enlarged) of Gl. morsitans, and details
of antenna, mouth-parts, and last joint of tarsus.

* By an obvious slip the word here in the text is “ partridges.”—E. E. A.

SELOUS’ EXPERIENCE ON ZAMBESI. 181

75. 1881. Dr. Emil Holub.*

“Seven Years in Soutw Arrica: Travets, Re-
SEARCHES, AND HuntTING ADVENTURES, BETWEEN THE
DiamMonD-Fieips AND THE ZamBESI (1872-79).” Trans-
lated by Ellen E. Frewer (London: Sampson Low,
Marston, Searle and Rivington), Vol. II.

[August, 1875.] “In the evening we halted facing a
wooded ridge, which would have to be crossed at night, on
account of the Tsetse-fly with which it was infested... .
The night was dark, and we could scarcely see ten yards
in front of us, but shortly after two o’clock we ventured to
start, and got safely through the wood without any
inconvenience from the Tsetse-fly, finding ourselves at
dawn. on the the plain called the Gashuma Flat” [near the
Panda ma Tenka River] (p. 105).

Portion of route to the Victoria Falls near the Gashuma
Flat known to be infested with Tsetse (p. 183).

Tsetse-fly near Panda ma Tenka: havoc caused by it
amongst bullocks (pp. 373-374).

76. 1881. F. C. Selous.

“ A Hunrer’s Wanpertnes In Arrica” (London:
Richard Bentley & Son).

Umziligazi’s cattle decimated by the Tsetse-fly (in
Matabele Land, north of Buluwayo), p. 31.

“ The ‘Tsetse’ fly has now come up to the Jomani
[a little river in Mashuna Land], so that hunters of late
years have had to travel by another road more to the
south” (p. 34, note).

“In the day-time, too, ‘Tse-tse’ flies, whose numbers
increased daily as the season advanced, were very trouble-
some [on the Lower Chobe River]. Nowhere does this
virulent insect exist in such numbers as to the westward
of the Victoria Falls, along the southern bank of the
Zambesi and Chobe. It is usually found in great numbers
near the river, becoming scarcer and scarcer as one
advances inland, till at a distance of a few miles it dis-
appears, except in some particular patches of forest.
Along the water’s edge they are an incredible pest,
attacking one in a perfect swarm, from daylight till
sunset, and, without a buffalo or giraffe tail to swish them

* Cf. [67].

182 DISEASE FATAL TO AZZ DOMESTIC ANIMALS.

off, life would be unendurable. The well-known African
traveller, Andersson, says their bite has been not inaptly
likened to that of a flea. My experience is that it is far
more severe, and that about one in every ten bites (that
perhaps touches a nerve) closely resembles the sting of a
wasp or bee, as it will cause one when seated to spring up
as if pricked with a needle. As they are possessed of a
long probe, a thick flannel shirt offers no protection
against these most abominable of all created insects—
direct descendants, no doubt, of the flies that plagued
Kgypt. Though, during 1872-73, I had hunted elephants
on foot in fly-infested countries, yet never had I met with
them in sufficient numbers to cause much annoyance ; but
along the Chobe river, during the months of September and
October, hunger, thirst, fatigue, and all the other hard-
ships that must of necessity be endured by the elephant-
hunter, sank into insignificance as compared with the
continuous, unceasing irritation caused by the bites of the
‘Tse-tse’ flies by day, and three or four varieties of
mosquitoes by night. What a glorious field lies open
there for an enthusiastic entomologist ! I think that this
plague of ‘ Tse-tse’ flies, along the Chobe and Zambesi, is
due to the enormous numbers of buffaloes that frequent
their banks, as they always seem very partial to those
animals. The bite of this remarkable insect, as is well-
known, though fatal to all kinds of domestic animals, is
innocuous to every species of game, and to man. A
general belief exists, that amongst domestic animals, the
donkey, dog and goat are exceptions to this rule, but this
is a mistake, for I have seen all three die from the effect
of its bites. That all the natives living in the ‘fly’
country possess both dogs and goats I admit, but these
have been bred there from generation to generation, and
have become acclimatised, whereas, if you take either a
goat or a dog that has been bred outside the ‘fly’ country,
into a district where the ‘ Tse-tse’ is found, it will die
in nine cases out of ten, and the original progenitors of
the animals the natives now possess were no doubt such
exceptions to the general rule. Even now, the. natives
told me, out of a litter of pups, born in the country and
of acclimatised parents, some always die of ‘fly’ symptoms.
The ‘Tse-tse’ fly is about the same size as a common

ELEPHANT-HUNTERS AND THEIR HORSES, 183

horse-fly, of a dull grayish colour, with bars of a pinky
tinge across the body ; its wings, however, do not lie in
the form of a pent-house, but are like those of an English
house-fly, only longer. Animals, such as horses and oxen,
that have been bitten by the ‘fly’ during the dry season,
usually live on until the commencement of the rains, but
seldom survive long after the first shower has fallen. It
often happens that when hunting with horses outside
but close to the ‘ fly’ country, one is led in the ardour
of the chase into an infested district ; if such is the case,
and it is uncertain whether the horse has been bitten or
not, the truth can be ascertained by pouring a few
buckets of water over him, when, if he has been ‘stuck’
(as hunters call it), his coat will all stand on end, like
that of a lung-sick ox. On several occasions horses haye
been purposely taken into parts of the ‘fly’ country,
where elephants were known to be plentiful, in the hope
that, by their aid, their owners would be able to shoot
enough ivory to compensate for the loss entailed by their
inevitable death ; for, of course, in tolerably open country
a man ought to be able to kill very many more elephants
on horseback than on foot. My comrade, W., once made
an experiment of this sort, and he informed me that at
the end of two weeks his horse grew too weak to hunt
with, and at the end of three could not carry him at
all, though it did not die for some time afterwards”
(pp. 130-132).

“The forest being pretty open, and the moon about
full, we got along well enough, and at last, about midnight,
we struck the river at a point at least twelve miles from
our camp, which we did not reach much before daylight.
I think it is usually believed Tsetse-fly will not bite at
night; but along the Chobe river (where they swarm),
and by moonlight, I can feelingly say that this is a
mistake. They kept flying up from the ground on to my
naked legs, and bit as furiously as in the day-time ; and,
judging from the deep curses and loud slaps behind me,
I had no doubt they were paying similar attentions to my
Kaffirs” (p. 154).

Tsetse-fly found on the Umbila River, Mashuna Land :
three flies caught in cattle kraal belonging to the author’s
camp, and “fly” seen not a mile from the wagons.—

~~

184 FLY-BELTS ON ZAMBESI ROAD.

“Of course we thought that all our oxen and the two
horses we had left at the wagons were ‘fly-stuck,’ and
cursed our luck accordingly ; but we were eventually
very agreeably disappointed, for we did not lose a single
ox, though two of mine and one of Wood’s showed evident
signs of having been bitten, becoming very thin and
running at the eyes. They were all young animals,
however, and at last pulled through, though one of mine
did not commence to make flesh again for more than a
year. These facts convinced me that it takes more than
one fly-bite to kill an ox or any other animal, and that
recovery from Tsetse bite is possible when the blood has
not been too strongly impregnated with the poison” (p. 349).

77. 188x. Dr. B. F. Bradshaw.

“THe Tsrrse-Fiy” (The Transactions of the South
African Philosophical Society, Vol. I1., Part I., pp. 51-55).

The Tsetse-fly “‘is very tough, and bears a good squeeze
without being any the worse for it.”

“Tt inhabits the forest country on the north bank of
the Zambesi in great numbers, and is found in three belts
along the Zambesi road. Commencing at the first, is a
belt along the Zambesi and Chobe rivers, varying in width
from eight to twelve miles, and is more numerous in some
localities than in others, according to the amount of game,
being fewer where game is scarce and vice-versd ; is much
more plentiful during the winter months, namely, from
April to end of September, than during the summer. The
next belt of fly is about fifty miles from the river, along
the wagon road, in a southerly direction, between it and
Panta ma Tenka, is about six miles wide and eight miles
from the trading station at Panta ma Tenka. Sometimes
you may pass in the day-time and see no fly, at other
times you will notice many. This part of the country is
one of those sandy ridges covered with forest so common
in the country. About six inspans, or forty-five miles from
Panta ma Tenka, is the third belt of Tsetse; the wagons
generally inspan after sunset, and trek for about three
hours, then tie up the oxen and let them sleep for a
couple of hours, inspan again for four hours, and we con-
sider ourselves past the region of the fly, which must be
twelve or thirteen miles wide; it is also very uncertain

INCREASE OF TSETSE OWING TO ZULU RAIDS. 185

here, one time very numerous, at another none to be seen.
The game also is generally very scarce here. The country
consists of heavy sand and forest, being a portion of the
sandy pool plateau. The Bushmen have told me that the
fly breeds in the buffalo droppings, and it seems as if there
was some truth init, because where the buffaloes have
been driven away in certain tracts, the fly has almost
disappeared. It is not found on the north bank of either
the Zambesi or the Chobe rivers, close to the water where
there are flats. ‘ Wankie’s’ people, the Mashapatans,
have informed me that before ‘ Mosilikatze’ drove them
across the Zambesi, the hilly tract of country between
the ‘Gwai’ and Daka rivers had no Tsetse, was thickly
inhabited by three tribes, namely Mashapatans, Matongas,
and Batokas, and that they had plenty of cattle, but
since the country became uninhabited and overrun with
game, it has become one of the great strongholds of
Tsetse, extending from the Zambesi river for at least
sixty or seventy miles in a southerly direction—one effect
produced by the Zulus’ love of rapine and slaughter.

“Tt bites throughout the day, except when it rains,
and during part of the night if warm ; in fact, I consider
it dangerous to travel at night with cattle or horses, until
it begins to grow cold towards the middle of the night,
as I have been bitten often until past 11 p.m. It is very
cunning, always darting, if watched, behind one’s back,
and if much persecuted with a fly-duster, darts to the
grass or bushes, but soon returns to the attack, It will
follow, for miles, fresh meat, or natives, but soon returns
within its bounds. Tsetse will not stay long in a camp,
after fires are lighted, although numerous all round. The
Kaffirs have said to me, when much troubled by the fly
in camp, ‘ Make a fire and they will go away.’ I found
the experiment to succeed. Its buzz is peculiar and not
easily forgotten” (pp. 51-52).

‘All domestic animals perish if bitten, sooner or later,
with the exception of the goat; the donkey seems to
resist it longest. Dogs are taken into the fly district by
the Bushmen when pups, and are allowed to suckle the
mother, and at the same time given as much fly to eat as
can be captured for them. The mother in this case dies
and the pups live and grow up, but are the most miserable

186 IMMUNITY OF GOATS.

looking objects I ever saw belonging to the canine race—
emaciated, small, and the hair standing straight up all
over them. (Goats, like all game, get fat and do not seem
to mind it. In fact, the Kaffirs keep large flocks in the
fly districts near the ‘ Victoria Falls,’ on the north bank
of the river.

“Unless bitten by several Tsetse, animals do not show
any symptoms in particular until the rains fall. Then
oxen especially get emaciated and weak rapidly, much
weaker than they would be if in the same condition from
poverty alone. The hair stands, the eyes water and have
a dull fishy look, the nose also waters, occasionally the
feet swell, also the jaw and sometimes under the belly,
and the skin has a wrinkled appearance about the hind
quarters. If inspanned they soon knock up and lie down
in the yoke, and even if, as seldom happens, an ox recovers,
he is never good for anything for at least three years, and
then cannot do much work in a hot sun” (p. 53).

‘“*Many traders have told me they believe one fly is
sufficient to kill an ox, and natives say that if an animal
is bitten slightly at one time, it is not a preventative
against death if bitten at a future period” (p. 55).

[Some of the foregoing statements are reproduced in
the paper by the same author entitled “ Notes on the
Chobe River, South Central Africa”: Proc. Roy. Geogr.
Soc., Vol. III., April, 1881, p. 212.]

78. 1882. E. C. Hore.

“Dake TAanGANyIKA” (Proceedings of the Royal
Geographical Society and Monthly Record of Geography.
New Monthly Series. Vol. IV., pp. 1-2, 14-15).

Transport by bullock wagons, tried for the first time in
Central Africa in connection with the London Missionary
Society's pioneer expedition in 1877, a failure owing to
the Tsetse-fly.

“ Road to the Lake.—The road we were to take, though
parallel to some extent with what might be called the old
road used by Cameron and Stanley, was in fact almost
entirely new. The portions from Saadani to Mpwapwa,
avoiding the Makata swamp, were taken on the recom-
mendation of Mr. Roger Price,* who had made a pre-

* Cf. [54].

EXPERIMENT WITH INDIAN ELEPHANTS. 187

liminary journey so far;...” “More than this, we
were to introduce a new (for Central Africa, at least)
system of transport—bullock waggons. Landing at
Saadani in June [1877], we at once commenced training
bullocks for this service, and at the latter end of July
we really started for the interior with our bullock train.
Succeeding eventually in reaching Kirasa [in the Mukon-
dokwa valley], the bullock transport came to an end by
the death of these animals by the Tsetse-fly. Every other
difficulty had been overcome for 150 miles of the most
difficult part of the road, and only in respect of the fatal
Tsetse can this experiment be called a failure” (pp. 1—2).

The Tsetse-fly on the shores of Lake Tanganyika.

“The Tsetse-fly abounds on the lake shores from
Ujiji round the south end and so up the west coast as far
as Ubwari” (p. 2).

[The author met Capt. Carter, of the elephant expe-
dition,* at the station of the African International
Association at Karema, on March 26, 1880 (p. 14).]

“T was impressed very favourably with the accounts,
and with what I saw of the elephant work. The surviving
animal was one which for many years in India had done
no work, and Carter was about to leave Karema for the
coast to receive some more elephants from India with
which he was to start the work of catching and taming
the African ones. Why this work has been abandoned
I cannot tell. It has been proved that waggons can be
got through, but we cannot use bullock waggons on
account of the Tsetse. I think it has been proved that
elephants can be got through, but that they would not
answer because of the immense Jabour and the number of
men required to load and unload them daily, and because
of the great weight concentrated on four points on shaky
ground. By using elephant waggons both difficulties
would be done away with, and both successes usefully
combined ” (pp. 14-15).

79, 1882. L. K. Rankin.
“Tur ELEPHANT EXPERIMENT IN AFRICA: A BRIEF
ACCOUNT OF THE BELGIAN ELEPHANT EXPEDITION ON THE
MARCH FROM Dar-ns-SaLaam To Mpwapwa” (Proceedings

* Cf. (66, 79].

188

ELEPHANTS ATTACKED BY TSETSE.

of the Royal Geographical Society and Monthly Record of
Geography. New Monthly Series. Vol. IV., pp. 277,
278, 283, 285-286, 288-289).

(Four Indian elephants were purchased from the
authorities at Bombay by the Belgians belonging to the
African International Association. They were landed at
Msesani Bay, whence they marched to Dar-es-Salaam,
early in June, 1879. The expedition was commanded by
Captain F. F, Carter, who was joined by Mr. Rankin at
Zanzibar before starting. The Kingani River was crossed
at Gungu (lat. 7° 2' N., long. 38° 37' E.) on July 14,
1879.)

The elephants severely attacked by Tsetse beyond the
Kingani, near the Lungwa river at Charinzi; they did not
contract the disease, although donkeys attacked at the same
time did so.

“On July 17th [1879] we first saw the Tsetse-fly, in a
belt of country infested by it, through which we had been
marching since crossing the Kingani. We were now face
toface with one of the three problems the expedition had
specially to solve, viz., could the Indian elephant, being
removed by long captivity and by its artificial treatment
from the safeguards of the wiid state, resist the attacks
of Tsetse, or would he, along with the ox, the horse and
the donkey, succumb to them? The problem was solved,
and that in the hoped-for manner. The fly swarmed on
the elephants till blood trickled down their flanks in a
constant stream. For days they endured this; and yet
they showed no prolonged signs of Tsetse poisoning—
lassitude, melancholy, running at the eyes—either at the
prescribed time, viz., eight days, or afterwards, though they
seemed pained and distressed during the infliction. The
donkeys, on the other hand, sickened more and more after
this, and at Mpwapwa were in a dying condition”
(p. 277).

“Our next camp was at Tumundu in the district
Kikunguri, chief Aceda Murumu; it was on the line of
the lower road to Bagamoyo. The temperature was
894° F.; height of position between 800 and 900 feet.

# = * *

‘<¥For the first time we now began to see giant creepers.

After passing some conical mountains we came to a tract

INDIAN ELEPHANTS IMMUNE TO DISEASE, 189

of forest where every tree was hung, festooned, smothered,
shrouded, enlaced, draped with convolvulus; tree was
linked to tree in the most extraordinary manner, and the
ground was carpeted with the plants. We here saw
Tsetse again” (p. 278).

Tsetse-fly at Rumuma.—* At Rumuma I again saw
Tsetse-fly, an observation which M. Broyon’s experience
confirms, he having lost some oxen through the fly after
bringing them safely all the way to Mpwapwa” (p. 283).

On arrival at Mpwapwa, Aug. 3, 1879, the elephants
did not appear to be suffering from Tsetse-fly disease after
twenty-three days’ exposure to the fly—“ At Mpwapwa our
first task was to draw up the ‘ Report’ for the King of
the Belgians, of which, unfortunately, I have no copy ;
but I remember it contained the words: ‘The elephant
experiment has now been proved a complete success.’
An assertion which Carter justified on the three counts of
(1) Their immunity against Tsetse after twenty-three
days’ exposure to that insect; (2) their maintenance
during one month mostly upon the uncultivated food of
the country, and therefore at little cost ; (3) their ability
to march over all styles of ground, soft, stony, sandy,
boggy ; to conquer all eccentricities of topography—hill
and dale, river and jungle—while Jabouring under double
their due weight of baggage, some 1,500 instead of 700 lbs. ;
and this in a style that no other beast of burden could
hope to emulate. At this distance of time, and notwith-
standing the subsequent death of three elephants and the
discontinuance of the experiment, I see no reason to
withdraw a word of Carter’s claim to success” (pp.
285-286).

“‘T have not the shadow of a doubt that there is yet a
great future in Africa for the elephant, especially when
the stage of capturing and taming the native species has
been reached” (p. 288).

“ Our troubles came in a cluster at and after Mpwa-
pwa.* There one of the donkeys died, on August 30th.
On stepping forth from my tent one morning, I saw the

* [Two elephants died at Mpwapwa. In the opinion of Mr. Rankin
their deaths were due, at least in part, to overloading, excessive work, and
insufficient and unsuitable food: the result in no way vitiated the success

of the experiment as regards the ability of the Indian elephant to with-
stand Tsetse-fly disease. ]

190 CATTLE ON LOWER CONGO.

poor beast just outside, his neck over the medicine-chest.
In his dire need he had crawled from his own quarters
all the way to my tent. I could do nothing for him ;
the Tsetse had done their work ” (p. 289).

80. 1882. Dr. E. Pechuél-Loesche.

‘Diz Loanco-Exprpition.” Dritte Abtheilung,
Erste Hilfte (Leipzig: Paul Frohberg), p. 299.

Tsetse-fly (?species) collected near Tschintschiétscho
(a town on the coast about 100 miles north of the mouth
of the Congo).* But a small herd of cattle is kept at
Landana (a short distance south of Tschintschitscho) and
a larger one at Boma (on the north bank of the Congo,
near its mouth), and the animals roam about freely and
thrive tolerably well, so that the fly cannot occur in those
districts [or else the hematozoon of Tsetse-fly disease is
absent. |f

Reference to the failure of the attempt on the part of
the Expedition to introduce cattle as beasts of burden
[Cp. 654].

Dr. Pechuél-Loesche mentions the general absence of
cattle throughout by far the greater part of Lower
Guinea ; only to the south of the Kuansa River do they
become domestic animals in the hands of the natives.

81. 1883. F. L. James.

“THe Witp TRIBES OF THE SouDAN. An account of
Travel and Sport chiefly in the Basé Country, being
Personal Experiences and Adventures during Three
Winters spent in the Soudan” (London: John Murray),
pp. 128-129.

A disease of camels called by the natives guffer, said by
some of them to be caused by the Tsetse-fly. (The district
referred to is that to the east of Kassala. )

“There is a disease, very common amongst them
[i.e, camels], which the natives call the guffer. We were
never able to clearly make out what this disease was.
Some of the Arabs declared it was catching ; others that
it was not ; but all said that a number of the camels we
had bought on the Atbara were suffering from it when
they were bought. Whenever we had to complain of any
of the camel-drivers having, through negligence, allowed

* Cp. [65, 654.] t Cp. [99].

SCHOCH’S THEORY OF TSETSE-FLY DISEASE. 191

a camel to run down, the excuse was that it was suffering
from this mysterious disease, the guffer. We were once
asked to look at a camel said to be suffering from this
complaint. It was certainly in miserably poor condition,
and at the time appeared to have a fit or convulsion
of some kind. It rolled on the ground, apparently in
great agony, and was only induced to get up after much
difficulty. Somehow or other it got through the day’s
march, but was never afterwards good for much. Some
of the natives said this disease was caused by the bite of
the Tsetse-fly during the rainy season” (p. 129).

82. 1883. Karl Kraepelin.
© ZuR ANATOMIE UND PHyYSIOLOGIE DES RUSSELS VON
Musca” (Zeitschrift fiir Wissenschaftliche Zoologie, 39.
Band, pp. 683-719, Tafeln xl. und xli.)

83. 1883. G. Schoch.

“Dig Tserse FLIEGE AFRIKAS” (Mittheilungen der
Schweizerischen entomologischen Gesellschaft, Band 6.
Heft 10, October, 1883, pp. 685-686).

“Tt is usual to designate by the name Tsetse-fly two
long-winged Muscide of similar appearance, one as large
as our house-fly, the other somewhat larger, and less
dreaded. Their larve are said to live in the dung of big
game” (p. 685).

Reasons advanced (p. 686) for considering that the fly
is not poisonous, but “at most the carrier of a bacterium-
like poisonous matter.”

“We conclude, therefore, that the fly is not the
producer of the poison, but at most the carrier of a
miasma which arises here and there, and further that,
with more intimate knowledge of the active principle, the
disease can be successfully opposed, and will gradually
disappear ” (p. 686).

84, 1883.

“ JOURNAL DE GENEVE,” December 3.

An article on the Tsetse-fly, with statements by a
certain H. F. Gros, with a view to showing that the
belief in the harmfulness of the Tsetse is mere prejudice.
[Apud F. M. Van der Wulp, Tijdschrift voor Entomologie,
1885, p. cv. I have not seen this paper myself. ]

192. TSETSE IN MASHONALAND IN 1882.

85. 1883.
“THE DELTA AND LOWER COURSE OF THE Sasr River,
ACCORDING TO THE SURVEY OF THE LATE Caprain T. L.
Puipson-Wysrants” (Proceedings of the Royal Geo-
graphical Society and Monthly Record of Geography. New
Monthly Series, Vol. V., p. 274).
Tsetse-fly met with by Captain Phipson-Wybrants to
the north-west of Mapeia’s Kraal, on the Sabi, and
between it and Macoupi’s, November, 1880.

86, 1883. F. C. Selous.

“ FuRTHER EXPLORATIONS IN THE MasHuna Covun-
try” [1882] (Proceedings of the Royal Geographical
Society and Monthly Record of Geography. New Monthly
Series, Vol. V., pp. 269, 270).

Tsetse-fly on the Panyame and Umsengaisi Rivers, at about
16° S. lat., 1882.—On p. 269 is a map entitled—‘ Routes
between the Umfule and the Zambesi, by F. C. Selous.”
On this the interval between the Panyame and Umsengaisi
Rivers, on the 16th parallel S. lat., is marked—*“ A vast
plain, covered with mopani forests. Very dry and swarming
with Tsetse along the rivers, where game is also abundant.”

“ Below the mountains [the range to the south of the
above plain, running due west from the Umvukwe Mts.]
the Tsetse-fly are in millions, and we are very much
annoyed by their incessant bites” (p. 270).

87. 1884. G. Macloskie.

‘* KRAEPELIN’S Proposcis oF Musca” (The American
Naturalist, Volume XVIII., pp. 1234-1244, Figs. 1-12).

An abstract of Kraepelin’s paper, “ Zur Anatomie und
Physiologie des Riissels von Musca” [Cp. 82].

The author writes :—“ Kraepelin’s paper gives the
most complete account extant of the structure of an organ
which has excited interest since the time of Aristotle. His
investigations were chiefly on the proboscis of the Blow-fly
(M. vomitoria), and exclusively on its adult anatomy. The
embryology of these parts has not been attempted by
Kraepelin, baffled Weismann, and remains yet to be worked
out. The following is an abstract of Kraepelin’s paper
with pen-and-ink copies of the more important of his
thirty-eight fine illustrations. I venture to add some
criticisms in the form of foot-notes,”

88.

89.

90.

91.

F. M. VAN DER WULP ON TSETSE. 193

1884. F. M. van der Wulp.

“Trts OVER DE Tsetse VuiEG (Guossina) ” (Tijdschrift
voor Entomologie, Zeven en Twintigste Deel, Jaargang
1883-84, pp. 143-150).

Includes a short bibliography. See also op. cit., pp.
xci-xcil.

1885. J. M. F. Bigot.
“Genre Guossina ” (Annales de la Société Entomologique
de France, 6° Série, Tome Cinquiéme, pp. 121-124).
Synoptic table of six supposed species, in French ; also
the original description of Glossina ventricosa.

1885. J. Mik.

Wiener Entomologische Zeitung, 1V. Jahrgang, pp. 60-—
61.

An abstract of G. Schoch’s paper, ‘‘ Die Tsetse Fliege
Afrikas” [Cp. 83]. The writer draws attention to the
fact that there are three additional species of Glossina in
Africa, while a fifth species, Gl. ventricosa, Big., is said to
come from Australia. [As has been shown in the systematic
portion of the present work, this is certainly a mistake. ]

1885. F. M. van der Wulp.

Tijdschrift voor Entomologie, Acht en Twintigste Deel,
Jaargang 1883-85, pp. cili—cvi.

Report of remarks by Heer van der Wulp, with subse-
quent discussion, at a meeting of the Nederlandsche
Entomologische Vereeniging, held Jan. 25, 1885.—Certain
previously published papers are quoted with a view to
showing that Tsetse bite is not the cause of the deaths of
cattle commonly attributed to it, or that, at any rate, the
Tsetse merely disseminates noxious influences. Van der
Wulp’s own opinion, which he says he shares with Baron
Osten Sacken, the well-known Dipterist, is that, although
the cause of the mortality in cattle in Africa is not alto-
gether clear, it is not, or at least not exclusively, due to the
bites of Tsetse-flies, which in all probability are not more
poisonous than European blood-sucking gnats and flies.
A subsequent speaker (Heer Veth) considered it not
impossible that by means of the Tsetse, as by other blood-
sucking insects, highly-poisonous contagious matters are
conveyed into the blood.

)

194 STOCK-KEEPING ON WEBBE SHEBEYLI.

92. 1885. W. Marshall.

« Unser pie Tsersn-Fiincs ” (Biologisches Centralblatt,
V. Band, pp. 183-184). »

A résumé of F. M. van der Wulp’s remarks reported in
the Tijdschrift voor Entomologie, Vol. 28, pp. ciii—cvi [vide
gr].

93. 1885. E. H. Richards.

‘‘ An AMERICAN MISSIONARY’S J OURNEY IN East AFRICA,
West or InnAMBANE” (Proceedings of the Royal Geo-
graphical Society and Monthly Record of Geography. New
Monthly Series, Vol. VII., p. 381).

Mr. Richards made a journey from Inhambane to the
Limpopo,* in October, 1884: the T'setse-fly was first met with
in Makwakwatand, to the west of the Makwakwa ridge.

“ The Ama-kwakwa tribe was encountered on the third
day. ... Many kraals were deserted, and a tract of

. country séventy-five miles wide by a greater distance in
length lying west of the Makwakwa ridge was nearly
desolate. It was in this semi-deserted region that the
Tsetse-fly was first seen. The route lay to the W.N.W.
along the northern border of Makwakwa-land.”

94. 1885. F. L. James. é,
“ A JouRNEY THROUGH THE SomALI CounTRY TO THE
Wesse Suepeyii” (Proceedings of the Royal Geographical
Society and Monthly Record of Geography. New Monthly
Series, Vol. VII., p. 633).
Fly (? Tsetse) on the Webbe Shebeyli in the wet season.—
“Like the Somal, the Adone [the people on the Shebeyli]
have large herds of cattle and flocks of sheep, but all these
animals are poor and suffer from the fly in the rain and
from the ticks in the dry season; neither camels nor
horses are used, for they will only live in the dry season ;
but the Rer Hamer, who leave the river valley for the
plateau in the wet season, bring numbers to graze there in
the winter,”
95. 1885. H. M. Stanley.
© THe Conco and ror Founpine or its Frer Srare :
A Story of Work and Exploration ” (London: Sampson
Low, Marston, Searle and Rivington), Vol. T., p. 419.

* [On a subsequent page (459) the river reached is stated to have been
the Luizi, and not the Limpopo.]

TSETSE AS REMEDY IN MASHONALAND. 195

Author bitten by Tsetse (and horse-tlies) on board a
river steamer at-Kemeh I.,-in -the Kwa R., an affluent of
the Congo, above Stanley Pool, May 21,1882. (Approxi-
‘mate position of Kemeh I., from aebhors map, 2° 45’ 8.
lat., 17° 2’ E, long.)

96. 1886. H. Capello and R. Ivens.
“De Ancota 4 Contra-Costa,” Vol. II. (Lisboa:
Imprensa Nacional), Capitulo XVIII,, “ A Tzé-Tzé,” pp.
21-39.
___. A chapter on the Tsetse (in Portuguese): rough wood-
_cuts of head and foot enlarged, on p. 23.

97. 1886. W. M. Kerr.
“A Journey From Cape Town overtAND To Lake
Nyassa” (Proceedings of the Royal Geographical Society
and Monthly Record of Geography. New Monthly Series.
Vol. VITI., p. 74, Map, p. 136). .
Tsetse-fly near Chibinga, Mashonaland, July, 1884.—
“The Tsetse-fly abounds in these parts, consequently
cattle and horses are unknown. Poultry of small size
seem to do well. The native women dry quantities of the
‘ Tsetse-fly and pulverise it with the bark of a root, and
mixing it with water give it to the young animals, such as
dogs and goats or sheep, of which they have very few,
aang ie only kept as pets.”

sia cea W. M. Kerr.
“Tue Far Interior: A Narrative of Travel and
Adventure from the Cape of Good Hope across the
. Zambesi to the Lake Regions of Central Africa” (London:
» Sampson Low, Marston, Searle and Rivington), Vol. L.,
pp- 282, 293, 306, 307; Vol. IL., PP- 24,.33, 48, 54, 63,
65, 75, 118, 193.
1884.—Tseise-fly at Suru, to the north of the Makomwe
Mts., on the north bank of the Msingua River, a tribu-
tary of the Umkumbura, which is an affluent of the
Zambesi (Vol. I., p. 282). Approximate geographical
position of Suru, from the author's map, 16° 20' S.,
31° 55’ E
Tsetse fly at lend near Chibinga, on tigh, north “a of
the Msingua, about 10 miles north-west of Suru(1884).—
.: “Flies > were . literally . ia. myriads... Fortunately the

Oo. 2

196

SWARMS OF TSETSE IN DRY RIVER-BED.

Tsetse-fly—although swarming in the neighbourhood, and,
in fact, throughout all this belt of country between the
foot-hills of the Makomwe mountains and the Zambesi—
did not trouble us. In the town it was by no means
constant in its attendance; in one place they might be
innumerable, but a mile further east, few could be seen”
(Vol. I., p. 293).

Tsetse-fly on the Mukumbra (Umkumbura) River (called
Ukumbura on author’s map): 1884.—“Soon we arrived
at the Mukumbra River, which may be described as a long
winding stretch of sand. It forms a very wide road
(broadening in some places to three, four, or six hundred
yards) through the vast forest, its shifting sands extending
to the union with the Umzengaizi River about 12 miles
south of the Zambesi. . . . Upon these low and dry sands
we saw the spoor of almost every description of wild animal.
. . . It was now late and we had had a very hard day ;
the first on which I had got a real ‘ benefit’ from the
Tsetse-fly. The condition of torment which I thought
tiresome before, I would now have welcomed as a state of
comparative bliss. The heat had been intense. Bathed
in perspiration we walked through the shifting sand, which
yielded like soft snow under the feet, while the stifling
sultry air was literally alive with the Tsetse-fly, against
whose maddening attacks clothes were no protection, our
only safeguard being to beat them off with twigs and
small branches of shrubs, giving a by no means pleasant
exercise under a torrid heat. For the same purpose the
natives generally use the tail of a buffalo mounted on a
wooden handle, an implement with which they swish their
naked bodies while travelling through the fly-infested
country... . Fortunately at night the flies take a rest ; but
T have felt them ‘stick’ more than once during nocturnal
hours. General experience, however, shows that they do
not give much trouble after sundown” (Vol. I., pp. 305—
307).

Tsetse-fly between Chibinga and Tette: 1884.—“ Tsetse-
fly varied in numbers. Sometimes they were swarming,
at other times few were to be seen ” (Vol. IT., p. 24).

Tsetse-fly and goats in the Zambesi valley, to the west
of Tette: 1884.—“The Tsetse abounds throughout the
widespread valley. It has been remarked that the goat

TSETSE IN NORTHERN ZAMBESIA. 197

will live in the ‘fly country.’ But to me it was singular
that wherever goats were found in any considerable numbers
the fly was not observed. Where the fly was prevalent,
goats were sometimes kept as pets which had been born
in the ‘fly country’ (their mothers having died), and
physicked, when very young, by the women” (Vol. II.,
p. 33).

Tsetse-fly not seen by the author in the centre of the town
of Tette: 1884.—‘ In the heart of the town I never saw
the Tsetse-fly. Therefore a few poor-looking cows were
kept, but were not allowed to wander far. With reference
to the Tsetse, I should mention that when game, such as
the buffalo, elephant, etc., become scarce, as was the case
recently in the neighbourhood of Delagoa Bay, the fly in
a great measure disappears, it being said, with good
authority, that the little pest breeds upon the buffalo-dung.
Doubtless when the game, at present plentiful, disappears
from the Zambesi valley, a like result will follow” (Vol. II.,
p. 48).

Tsetse-fly on the Zambesi, at Tette: 1884.—‘‘ Tsetse-fly
abounds on the southern side, and there is also a belt of it
on the northern lands contiguous to the river” (Vol. I1.,
p. 54).

Tsetse-fly on the Revuqwe River, about six miles above
its confluence with the Zambesi: 1884.—‘‘ A sharp prick
at this time made me renew my acquaintance with the
Tsetse ; after sundown, too, which was rather alarming”
(Vol. IT., p. 63).

Tsetse-fly swarming in places during a few days’ march
in a north-easterly direction from the above locality on the
Revuque: 1884,—*‘ Forests of varying density were pene-
trated in our route during the next few days. Mopani,
thorn jungle, long coarse grasses, and clusters of bamboo
were the principal growths. River-beds of dry sand, such
as the Matizi, Nyamtara, and Nyabzigo, ran through the
country.

“The Tsetse swarmed in certain localities, but alto-
gether they were not so bad as we had found them here
and there farther south” (Vol. II., p. 63).

Tsetse-fly between the Shikambe River and the south-
western extremity of the Salumbidwa Mountains: 1884.—
Approximate geographical position, from the author’s map,

198 . RECORDS BY W. M. KERR,

_ 15° 56' §., 33° 55’ E.—*“ Pursuing our course for two days
through an uninteresting country, covered with tropical
vegetation, we crossed the Shikambe and Mbjova rivers,
both of which give their waters to the Zambesi above
3 Lupata gorge, and pitched a camp at a place which I have
designated the Palm Wells, 800 feet above the sea. . .
Every now and then the Tsetse would alight upon
our thinly-clad bodies, making us jump again as though
twitched with the fine lash of a — ” (Vol. IL, pp.
64-65).

Tsetse-fly' near the southern — of the Kapirizange
Mountains: 1884. Approximate geographical position,
from the author’s map, 15° 49’ 8., 34° 5’ E.—* Just before
beginning the ascent of the Kapirizange range, I had seen
Tsetse-fly ; but in the mountains none wére to be found ”
(Vol. II., p. 75).

Donkeys, though “ more tenacious of life than horses

or oxen when in the fly country,” eventually “pine away
and die like other victims” (Vol. IT., pp. 118~119).

Tsetse-fly not seen in the vicinity of Livingstonia, at the
southern end of Lake Nyasa: 1884.—‘“ No Tsetse-fly was
seen here, but it is more than probable that the deadly
insect is as migratory in its habits as the game on whose
dung it breeds ”* (Vol. II., p. 193).

[In the map at the end of Vol. II.—*“ Map showing
route from the Cape of Good Hope across the Zambesi
River at Tette to Lake Nyassa in 1884. From the
Survey by W. Montagu Kerr, C.E. Scale of English
miles—1 inch = 38 miles ”—the occurrence of the Tsetse-
fly in different regions is shown in red. A dotted curved
line running from about 19° 10'S. lat., and the 29th
parallel East longitude to about 17° 50' S., 30° 35' E., is
marked “ Approximate Southern Limit of Tsetse-Fly.”
The “Southern Limit of Tsetse-Fly,” south of the Zam-
bési, is: again shown by a line running east and west
from about 16° 20’ S., 31° 8’ E. to 16° 20'8., 32° 20’ E. ;
the country between this line and the Zambesi is marked
‘*Tsetse-Fly Belt.” North of the Zambesi a line running
W.N.W.—E.S.E., across the northern extremity of the
Salumbidwa Mountains, from about 15° 45’ §., 33° 40’ E.,

* Tdsnialn J. F. Elton found Tsetse abundant at Livingstonia in
August 1877.—Cf. 59.)

99.

100.

101.

102.

PERTHES MAP OF AFRICA. 199

to about 15° 53'S., 34° 30’ E., is indicated as the “ Northern
Limit of Tsetse-Fly.” Besides the above, “ Tsetse-fly ” is
shown as occurring to the N.E. of Tette, 8.W. of the
intersection of the 16th parallel 8. lat., and 34th meridian
E. long., and also at the southern end of Lake Nyasa, on
the western side of the southern extremity of the bay to
the west of the promontory on which Livingstonia is
situated. Two localities bear the words * No Tsetse-Fly ” ;
the first a valley among the foot-hills to the east of the
Umvukwe Mountains, in the Makorikori country : approxi-
mate geographical position, 16° 55'S., 31° 40' E.; the
second a portion of the valley of the Vilange River, an
affluent of the Revuqwe, approximate geographical posi-
tion, 14° 57'S., 34° 14' E.]

1887, Josef Chavanne,

“‘ REISEN UND ForRSCHUNGEN IM ALTEN UND NEUEN
KONGOSTAATE IN DEN JAHREN 1884 unp 1885” (Jena:
Hermann, Costenoble), p. 365.

The fact that cattle thrive fairly well, in spite of an
only moderate supply of unsatisfactory fodder, shows
that the region of the Lower Congo is free from
Tsetse-fly.*

1887. Hermann Habenicht.

Justus Perthes’ SprEZzIALKARTE VON AFRIKA, ent-
worfen von Hermann Habenicht (Gotha, April, 1887).
Scale 1 : 4,000,000,

Sections 9 (“Capland ”) and 10 (‘ Delagoa Bai”) show
by means of dotted lines the limits of the Tsetse-fly north
and south of the Limpopo, and south of the Zambesi.

1887. Dr. C. W. Schmidt.
Berliner Entomologische Zeitschrift, XXXI. Band,
p- 368.
Tsetse-fly stated at to occur in Usambara and Bondei
[districts near the Pangani River, German East Africa],
Cattle can therefore be kept on a large scale in Usambara.}

1888. A, Laboulbéne.
Annales de la Société Entomologique de France, 6° Série,
Tome Huitiéme, Bulletin, pp. lxxxviii, clvii.

* (Cf. 80.]
+ Cf., however, Dr. Stuhlmann’s statements [XXVI.].

200 LABOULBENE’S THEORY OF TSETSE-DISEASE.

103.

104.

Opinion expressed that the Tsetse is not poisonous
in itself. The ravages of the Tsetse or of the species of
Glossina appear to the author to be due to septic matter
derived by the insect from unhealthy animals or from
carcases, and inoculated from one animal to the other
(p. lxxxviii).

A communication read from R. P. Leroy, according
to which the Tsetse is very troublesome in the dry and
sterile plain between Ukami and Nguru (inland from
Bagamoyo), on which buffaloes, zebras, giraffes, and
antelopes abound, while there are even a few elephants.
Leroy states that a Masai army with a herd of 300 cattle,
camped on the plain in order to attack Mrogoro, was
forced to beat a precipitate retreat in consequence of their
cattle having been put to flight by the Tsetse. Laboul-
béne explains that he did not attribute the effects of the
bite of the Tsetse to the fly having previously settled on
putrefying carcases. He adds: ‘I think that it carries
with its proboscis septic matter drawn from diseased
animals, and communicates it to others that are healthy,
the result of which is illness and possible death. It
remains to be explained why the bite, which is said to
be deadly to cattle, is not so to human beings”
(p. elviii).

1888. A. Laboulbéne.

‘Une Movucue Tst-Tst pr L’ AFRIQUE TROPICALE ”
(Revue Scientifique, Troisiéme Série, Tome XV, p. 700).

Contains no fresh facts: according to a foot-note, the
article is an extract from a communication made by the
author to the Académie de Médecine, at a meeting held
May 29, 1888. [The title of this communication is,
‘Sur une mouche tsé-tsé de l'Afrique Australe” : Bulletin
de ? Académie de Médecine, Paris, 1888, 2° Série, XIX,
pp. 721-724.]

1888.

“Dr. Hoius’s JouRNEY 1N THE Batoka Country ”
(Proceedings of the Royal Geographical Society and Monthly
Record of Geography. New Monthly Series. Vol. X.,
p. 647.)

Tsetse-fly in the Batoka country, 1886.

“ He [Dr. Holub] says the country of the Batoka is

105.

105.

106.

TSETSE IN TRANSVAAL. 201

wooded, but the forests are only composed of small trees,
in which the Tsetse-fly abounds.”

1888. F. Jeppe.

“THE Kaap GoLp-FIELDS OF THE TRANSVAAL” (Pro-
ceedings of the Royal Geographical Society and Monthly
Record of Geography. New Monthly Series. Vol. X.,
p. 441).

Partial disappearance of the Tsetse-fly together with
game from the Kaap Valley.—<Game has almost dis-
appeared, frightened by the reports of the dynamite and
the prospector’s gun, and the Tsetse-fly has gone away
with the game, save in some parts of the valley. The
valley, formerly teeming with all kinds of game, from the
elephant to the steinbuck, is now deserted except by a
stray leopard in the kloofs, or the lion on the Lebombo
flats, where more game is found.”

1888. Prof. A. H. Keane.

Articte “TRrANsvAAL” 1N Encyclopedia Britannica,
9th edition, Vol. XXIII., p. 518.

« A tsetse belt 40 miles wide along the whole course
of the Limpopo still bars the spread of European settle-
ments beyond Transvaal in the direction of the Zambesi.”

1888.

“TsErsE-FLY (Glossina morsitans)” (Encyclopedia
Britannica, 9th edition, Vol. XXIII., p. 601).

A short article——‘ At present no cure is known for
the bite, nor does inoculation seem to afford any pro-
tection. The fly is said to avoid animal excreta, and in
some parts a paste composed of milk and manure is
smeared on cattle which are about to pass through the
‘ fly-belts.’ This affords a certain amount of protection.
Lion’s fat is used in the same way, and is said to be
efficacious.

“The fly is found as a rule in the neighbourhood of
water, and its habitat is usually sharply defined. Often
it occurs on one side of a stream, but not on the other.
The limits of the ‘fly-belts’ are well known to the
natives, and travellers can ensure comparative safety to
their cattle by passing through these districts after
sundown. The northern limits of the area inhabited by

bo
S
bo

107.

108.

109.

TSETSE IN CAMEROONS.

the Tsetse are not known. It is found throughout the
valley of the Limpopo River, but does not come much
south of this, except in the eastern borders of the Trans-
vaal. Here it extends far south of Delagoa Bay, and
infests the Lobombo’ Mountains and the Amatonga
country, reaching to the confines of Santa Lucia Bay.
It appears to be gradually retreating northwards following
the big game.” :

1889. Dr. Preuss,

*‘Bertcut von Dr. PREUss UBER BOTANISCHE UND
ENTOMOLOGISCHE BEOBACHTUNGEN AUF DER BaRomsI-
sTATION ” (Mittheilungen von Forschungsreisenden und Gelehr-
ten aus den Deutschen Schutzgebieten. Mit Benutzung
amtlicher Quellen herausgegeben von Dr. Freiherr von
Danckelman (Berlin: Kommissions-Verlag von A. Asher
& Co.), II. Band, 2. Heft, p. 60),

A species of Glossina at Barombi Station, Cameroons.

[Translation.] ‘On the other hand, a yellowish-
brown Glossina is distinguished by its particularly painful
bite (Stich). This fly is found chiefly in the primeval
forest, and pursues and persecutes human beings with
downright revolting cunning and pertinacity. "When one
is bitten a small drop of blood at once exudes and the
place becomes much swollen.”

[Barombi Station is situated to the south-east of
Barombi, or Elephant Lake, a little to the south of
5° N, lat., and just E. of 9° 30’ E. long.]

1889. F. S. Arnot,

“ JouRNEY FROM Naran To Bint anp BENGUELLA,
AND THENCE ACROSS THE CENTRAL PLATEAU OF AFRICA
TO THE Sources OF THE ZAMBESI AND Conco” (Pro-
ceedings of the Royal Geographical Society and Monthly
Record of Geography. New Monthly Series. Vol, XT.

Tsetse-fly prevalent “in the country between the Mababi
flat and the Zambesi” [1882] (p. 67).

1890. John B. Smith.

*‘ Norges ON THE STRUCTURE AND History or Hama-
TOBIA SerRatTA” [= Lyperosia (Heematobia) irritans, L.|:
Psyche, Vol. V., pp. 343-347, figs. 1 to 6 (woodcuts of
details of proboscis and other organs).

TSETSE NEAR PANDA-MA-TENKA. 203

110. 1890. Dr. E. Holub. :

“Von per Capstapt ins LAND DER MascHUKULUMBE.”
Reisen im siidlichen Afrika in den Jahren 1883-1887.
(Wien: Alfred Hélder), II. Band, pp. 10-12, 191, 398,
483, et seq.

From Panda-ma-Tenka to Gazungula, June, 1886.

[Translation.] ‘In the night between June Ist and 2nd
we set out in order by the aid of the waggon to transport
the greater part of my belongings to the mouth of the
Chobe—.e., to the southern bank of the Zambesi, which,
on account of a shady Gazungula tree which stands
directly on the shore, is called Gazungula—there to
remain until we crossed the Zambesi. ... We found
ourselves obliged, as already mentioned, to cover the
intervening space by night, since on part of the road
to be traversed the Tsetse-fly was still to be found
during the heat of the day, so that this spot could
only be passed at night. If, as it has hitherto been
customary to believe, the Tsetse-fly is really the destructive
agent that kills our domestic animals in certain districts
of Africa by means of its poisonous bite, it is, in my
opinion, no protection to pass certain thickly wooded
laterite knolls, which usually cross our path at right
angles and are infested by Tsetse, in the night, that is at

_a time when this dangerous insect is resting on twigs
asleep. The path is so narrow and the trees are so close
to it, that it is quite impossible to pass without the trek-
oxen and the waggon scraping against the twigs and so
shaking off the Tsetse and carrying them along with them.
‘I found that at the time of my first visit, on the section
from Panda-ma-Tenka to the mouth of the Chobe, the
Tsetse-fly extended from Schneemann’s Pan (12 English
miles from Leschumo Station) to the Chobe mouth,
Since then, in consequence of periodical forest fires,
the Tsetse-fly in this district has gradually been destroyed,
until it has been restricted to one laterite knoll close to
the spot where the Leschumo Valley opens into that of
the Zambesi.

“As, however, a waggon passed this spot at the
most twice or three times a year, no opportunity was
afforded for transporting the poisonous fly to any con-
siderable extent, and it remained confined to the narrow

204 TSETSE BROUGHT INTO LESCHUMO VALLEY.

tract of woodland already referred to. But when,
in the year 1885, the missionary Coillard repeatedly
drove from the Leschumo to the Zambesi in order to
migrate to Sesheke, and when besides in the year 1886
the trading-store was transferred from Panda-ma-Tenka
to Gazungula, and the spot was thus frequently passed
with waggons and draught animals, it was naturally quite
possible for the fly to be transported to the Leschumo
Valley. I already expressed this apprehension on this
my first visit to the Leschumo Valley (February, 1886),
and was glad when I was at last able to take my depar-
ture and had brought my trek-oxen safely out of the
Leschumo Valley to Panda-ma-Tenka again. It was
indeed high time that this should be done, for when, after
I had accomplished my journey, Messrs. Westbeach and
Coillard travelled more frequently to and fro between the
Leschumo Valley and the Zambesi, so many of the flies
were transported to the former valley, that the insect was
able to establish itself there, and moreover the conse-
quences of this soon showed themselves. The draught
animals belonging to other travellers, who came from the
south into the Leschumo Valley, fell sick with the
symptoms of Tsetse poison, and perished all but two.
This much is certain, that this dangerous insect, which
will cause great difficulties in the colonisation of South
Africa, recedes and disappears before the advance of
civilisation.

“The Leschumo Valley from the mission station
occupied by us as far as its opening into the Zambesi
Valley exhibits a slightly mountainous character; it is
only a few hundred metres in width, covered with high
grass and wooded like a park, and is bordered on each
side by the high laterite ridge. Now at the mouth of the
Leschumo Valley the laterite ridge on the left sends a
spur towards that on the right, and this wooded spur is
said to have constituted the fiy-belt that still remained
here, while the level, meadow-like, bushy and wooded
tract extending from the spurs as far as the Chobe and
Zambesi Rivers is said to be free from Tsetse. The latter
I have never believed, and I think that those also who
maintained it did not believe it to start with, for other-
wise, why did Messrs. Westbeach and Coillard, as often

DIMINUTIVE SIZE OF CATTLE DUE TO TSETSE. 205

as they had brought their goods to Gazungula by night,
drive their teams the same night back to the Leschumo
Valley? This precaution on their part seemed, however,
gradually to be neglected ; and, in spite of my remon-
strance, Coillard left his teams in Gazungula, since Mr.
Westbeach’s remark that there was ‘no Tsetse-fly in
Gazungula’ carried more weight than my advice”
(p. 10-12).

Tsetse-fly along the Mo-Njeko River, and in the elevated
plain to the north of Masosa Village, on the southern confines
of Mashukulumbe Land. The Matoka in this district possess
cattle, sheep, goats and dogs, but they are dwarfed, and the
author thinks that this is due to the effects of fly-bite.—
Approximate geographical position of Masosa, according
to the author’s map, 16° 12’ S., 28° 7’ E.

“On this day [July 16, 1886] we had emerged from
the northern Matoka Highlands, and now stood on a
plain, which once more towards the north appeared to
descend to the Luenge, and was bounded only towards
the east and in the distance by a tract of highland, and
in the west by the elevation of laterite rock. This plain,
which extended in front of us far towards the east, north-
east and north, is stated by the Matoka to be a region
free from ‘I'setse-fly. Here were situated a few cattle.
posts belonging to King Mo-Panza and certain of his
chiefs, who prided themselves not a little upon their
small stock of cattle. . . . The news about the Tsetse-fly
seemed to me to be absolutely incredible, for I satisfied
myself that down below on the river, but especially on the
further bank, even then, though it was winter, the Tsetse-
fly occurred in much greater numbers than at any of the
other places north of the Zambesi, though, unfortunately,
we had found these localities everywhere infested by this
destructive insect. The Tsetse also existed on this plain,
and I think that the cattle that occur there have indeed
accommodated themselves to the poison of this fly, but
have remained so diminutive [the northern, or Maschu-
kulumbe race of the cattle belonging to the Matoka,
‘small cows and oxen not above middle size’] owing to
the influence of the poisonous insect, that is, they have
degenerated, just as the Matoka in the Tsetse country are
able to keep only stunted sheep, goats and dogs” (p. 191).

206 - TSETSE ON MO-NJEKO RIVER.

Abundance of Tsetse on the Mo-Njeko, near. Masosa
Village, August 7th, 1886.—‘‘I had started. about half-
past five in the morning, and it must have been about
two o'clock when I reached the Mo-Njeko, and soon after-
wards the little village on the south (right) bank.
Nowhere had I yet beheld so many Tsetse-flies as here.
On sitting down by the margin of the Mo-Njeko river—
the current was only very slight—and bathing my feet in
the clear water, I scarcely had time to keep the Tsetse-
flies off poor Daisy, who this time had accompanied me.
That Daisy did not succumb to the bites of the Tsetse is
doubtless due to the fact that previously on the Limpopo
he had thrice eaten meat poisoned with strychnine”
(p. 398).

Gazungula to Shoshong, 29th November, 1886, to 11th
February, 1887.—Sickening and subsequent death of
27 oxen from Tsetse-fly disease; serious illness of re-
mainder (pp. 483, et seq.).

111. 1891. Major Gaetano Casati.

“Tren Years In Equaroria” (London and New York:
Frederick Warne & Co.), Vol. I., p. 234.

Tsetse-fly in the Mambettu Country, west of Lake
Albert Nyanza, south of the Niam Niam Country, and
north of the Nowelle or Aruwimi River.

“ Cattle cannot be successfully reared by the Mambettu
on account of a fly called Tsetse, the stings of which cause
death. . . .” (p. 234). Reduced copies of Westwood’s
figures of fly and mouth parts.

112, 1891. E. A. Maund.

“On MaraBeLe AND Masnona Lanps” (Proceedings
of the Royal Geographical Society and Monthly Record of
Geography. New Monthly Series. Vol. XIIL.).

‘Three scourges farmers have to combat: lung-sickness
among cattle, horse-sickness, and the Tsetse-fly. ...
The Tsetse-fly, whose bite is so deadly to domestic
cattle, will disappear with the game. The Transvaal,
since the game has been so shot out, is now nearly free
from this pest. The Mashonas dry and pound the fly,
and give it to their dogs, a fly a day, as a safeguard
against the effects of it” (pp. 11-12)...

SIR F. D. LUGARD ON BRITISH EAST AFRICA. 207

113.' 1891. J. M. F. Bigot
Annales de la Société Entomologique de France, Année
1891, Vol. LX., pp. 377-378.
Original descriptions of Glossina grossa and Gl.
pallicera,

114. 1892. j. A. Nicolls and W. Egilington.

‘Tue Sportsman in Sourn Arrica” (London: The
British and Colonial Publications Company, and Simpkin,
Marshall, Hamilton, Kent & Company, Limited), pp.17,73.

“Some Remarks on the Shooting Horse and the
Hunting of Large Game.—If we except those portions of
the low-lying and most unhealthy country on the East
Coast between the mouths of the Zambesi and Crocodile
Rivers which are impenetrable to horses on account of
the presence of the Tsetse-fly pest, all the ordinary larger
game (if the Koodoo and different varieties of water-
loving antelopes be excluded) nowadays roam so far from
the permanent waters as to render hunting on foot
almost a sport of the past” (p. 17).

“In South-Central Africa, or, indeed, wherever the
Buffalo is met with in abundance, the Tsetse-fly (Glossina
morsitans) is everywhere prevalent.” Symptoms of fly-bite
in horses and oxen. “Some authorities are willing to
account for this pest being always found in attendance
on the Buffalo by the fact that the insect hatches its
eggs in the skin or the dung of that animal” (p. 73).

115, 1893. —~—. : bey my cm D.S.O, (now Sir F. D. Lugard,

“Tae Risk or Our East Arrican Empire” (Edin-
burgh and London: William Blackwood & Sons), Vol. I.,
pp. 389-390, 390-391.

“On the other hand, East Africa has many great
advantages. It is practically free from the Tsetse-fly
(Glossina morsitans) which arrests progress in South
Africa” (pp. 389-390).

“South Africa, as I have said, is at a disadvantage
by reason of the Tsetse-fly, which precludes all forms of
animal transport, and all agricultaral methods which
depend on the use of the horse, bullock, or donkey, as
well as all stock-rearing farms. Hides, ghi, milk, and
beef are products which cannot be procured in the

208 IMPORTANCE OF BEIRA RATLWAY,

infested districts. In German East Africa throughout
the broad zone, which he describes as the coast area,
Dr. Pruen says ‘the Tsetse-fly holds undisputed sway.’
In British East Africa the fly has been found in an
extremely narrow strip of country near Taita, but so far
it has only been caught as a natural history curiosity.
Indeed, by diverging either to the right or the left, it
can, I think, be avoided, for it is doubtful if it exists on
the Sabakhi River, and Mr. Jackson* says that, pro-
ceeding from Mombasa vid Taveta to Machako’s, ‘ Through-
out the whole of the way both good water and fodder are
plentiful ; there is no thick bush, and there are no Tsetse.’
Mr. Jackson is a competent authority, who has travelled
in this district (Taveta), and it was by him that the
Tsetse-fly in the direct route was first discovered and sent
home for identification to the British Museum. Thus
transport animals could be used throughout East Africa
from the coast to the far interior. Even across the strip
of country in which it has been found the Arabs and
Swahilis have for years driven trains of pack bullocks and
donkeys in their slow-moving caravans, and from this it
would appear that the fly can hardly be said to exist in
sufficient quantities to be a real obstacle, for these people
could not afford to risk the loss of even a few animals.
The Company have sent up caravans of donkeys, but I
greatly doubt if the mortality amongst them was due to
Tsetse. Three horses were safely imported to Uganda ;
and the tribes breed and rear cattle continuously from
the coast to the far interior” (pp. 390-391).

116. 1893. Theodore Bent.

The Standard, October 11, 1893.—Interview with a
representative of Reuter’s Agency.

Importance of the Beira Railway, in that by its aid
it is possible to cross the Tsetse belt without difficulty.
Numbers of waggons seen rotting on the roadside near
Chimioyo, “all the oxen having succumbed to the attacks
of the fly.”

117. 1893. V. von Réder.
Jahrbuch der Hamburgischen Wissenschaftlichen An-
stalten, X. Band, p. 205.

* Pall Mall Gazette, Feb. 4th, 1898 [Cf. 119].

TSETSE NEAR PUNGWE RIVER. 209

Glossina morsitans, Westw., taken at Mbuzini, in
Uzeguha, German East Africa, 27. viii. 1888, by Dr.
Fr. Stuhlmann.

118. 1893. Major J. J. Leverson.

“GEOGRAPHICAL RESULTS OF THE ANGLO-PoRTUGUESE
De.imiration Commission In SoutH-East Arrica” (The
Geographical Journal, Vol. II. London: The Royal Geo-
graphical Society).

Tsetse-fly on and near the Pungwe River, 1892.

“When proceeding up-country from the Pungwe we
did not get clear of the Tsetse-fly till we reached Shimoya’s.
Thence to Massi-Kessi ox-waggons brought down from
Mashonaland were able to work along the road. South
of Massi-Kessi, the only place in which we came across
the fly was in the neighbourhood of the Sabi. Five
ponies which we took with us from Natal all died, or had
to be shot within six weeks of their landing at Beira.
We also lost from fly-bite every one of twelve oxen which
made the journey from Shimoya’s to Mandigo’s and _ back,
a section of the Beira Massi-Kessi road which till the
commencement of 1892 was believed to be free from fly ”
(pp. 517-518).

119. 1893. F. J. Jackson.

Pall Mall Budget, February 23, 1893, p. 295.

Notes, in an interview entitled ‘“‘ Uganda Once More,”
on the Tsetse-jly on the Teita-Kibwezi route to Uganda, in
1892-93.

“Nearly the whole way is bush between Teita and
Kibwezi, in which there is very little fodder to be found,
and that of a coarse and very inferior kind. Further, as
you approach the Tsavo, after leaving Teita, you have to
pass through the Tsetse-fly country, and that continues
until you reach Kibwezi. The Tsetse is a small fly, rather
larger than the common house-fly, of a brown colour, with
a yellowish line down the centre of the body. This fly
attacks men and beasts both by day and night. It can
easily bite through khaki breeches or flannel pyjamas.
Where it exists—and it is extremely local—it appears in
much greater numbers on a dull or rainy day. Its bite in
man, though very sharp and needle-like, leaves no irrita-
tien, and it invariably chooses those parts that are shaded

P

~

10

MR. F. J. JACKSON ON TSETSE.

from the rays of the sun—such as under the brim of the
hat, behind the ear, neck, and under the throat if not
protected by a beard. If allowed to suck itself full, its
body becomes so distended that it has the appearance
of a small unripe purple grape, and it is unable to fly more
than a few yards before settling. Its proboscis, which
is very slightly curved downwards, is attached to the
head horizontally, and is protected by a sheath divided
longitudinally. When it bites, the proboscis is vertically
lowered, whilst the double sheath is retained in its hori-
zontal position. This fly is perfectly harmless to human
beings, but its effect on domestic animals is of the most
disastrous nature—in fact, as far as my knowledge goes,
all domestic animals, with the exception of goats, and
perhaps of the grey donkey, when struck by this fly are
bound, sooner or later, to die. I do not believe that
donkeys are impervious to its bite, and I think that the
wasting disease which kills off so many of them comes from
the effect of being bitten by these flies. JI have seen a
bullock cut up, after dying from these wasting symptoms,
which I knew to have been bitten by the Tsetse; the
whole of the juices of the body were turned into greenish-
yellow slime. I sent home a number of these Tsetse
which I caught myself, and which were certified as the
genuine fiy by the authorities at the British Museum.
. .. The route which appears to me to be far more
practicable for transport by means of animals is that by
Taveta, which lies at the foot of Mount Kilima-Njaro, and
from there runs almost direct northwards to Machakos.
Throughout the whole of that way both good fodder and
water are plentiful ; there is no thick bush, and there are
no Tsetse. The fact of the immense herds of native cattle
which the Masai have roaming over all these plains is
sufficient to prove this; if these herds are able to live, the
Tsetse cannot be there. . . . The depot at Taveta would
have to be built outside the forest, on the banks of the
River Lumi, as there is a fly, not the Tsetse, but exactly
like a house-fly with a long proboscis, that is very plentiful
inside, and worries beasts to such an extent that they are
unable to eat, and die.” ;

[The fly alluded to in this last paragraph is obviously
a species of Stomoxys, and since Trypanosoma brucei was

120.

121.

TSETSE IN RHODESTA. 211

found by the late Captain Haslam in specimens of
Stomoxys on the line of the Uganda Railway, it is quite
possible that the animals that Mr. Jackson considers to
have been worried to death by Stomoxys at Taveta really
perished from Nagana. In this connection it is worthy
of note that Dr. J. W. Gregory attributed the death of his
camels at Ngatana to the bites of a Stomoxys.

The above article is illustrated by a small sketch-map
of alternative routes from the coast to Uganda, showing
“Tsetse-fly country” near Kibwezi, and also by a rough
wood-cut (three times natural size) of a Tsetse-fly. ]

1893. Captain Foster,

“ Hanppook or Britiso East Arrica” (prepared in
the Intelligence Division, War Office, 1893), pp. 93, 111.

Tsetse-fly during the wet season infests some 60 miles
of road up country near the Tsavo; it is believed that
donkeys withstand the attacks of the fly better than other
animals (p. 93).

Tsetse-fly common on the Sabaki River route (which
follows the river) between Mombasa and Tsavo; on the
Taru desert route the fly does not appear till the valley of
the Tsavo is reached (p. 111).

1893. F. C. Selous.

“TRAVEL AND ADVENTURE IN Sourn-East AFRICA”
(London: Rowland Ward & Co.), pp. 54, 55, 63, 99, 202-
203, 208, 215, 245, 246, 250, 274, 410-411, 426.

Tsetse-fly said to be on the Savakaranga, a small stream
between the Manyame and Umsengesi Rivers, tributaries
of the Zambesi, in northern Mashonaland (p. 54).

Tsetse-fly making its presence disagreeably felt in the
Zambesi Valley, between the Manyame and Umsengesi
Rivers (p. 54).

“The Tse-tse-fly swarmed along the River Kadzi [near
the Umsengesi], and was a great pest, keeping one in a
perpetual state of irritation all day long” (p. 55).

“The Tse-tse-fly swarms on both sides of the river in
the immediate vicinity of Zumbo [a town on the northern
shore of the Zambesi, close to the confluence of the River
Loangwa], so that cattle cannot be kept there... .”
(p. 63).

Lsetse-fly on the Sabi River, Matabeleland.—“I was

P 2

212 DONKEYS AND TSETSE-FLY DISEASE.

now close to the edge of the ‘ fly’ country, which at this
time [1883] extended from here [on the Upper Sabi,
where the river crosses 32° E. long.] southwards along
both banks of the Sabi” (p. 99).

Resisting power of the donkey to Tsetse poison.— In
countries where the Tse-tse-fly is in excessive numbers, the
donkey, strong as he is, will not live long; but he resists
the poison far better than any other domestic animal, and
will pass through belts of ‘fly’ country without taking
much harm, his constitution being so strong that he will
recover from the effects of the poison if not kept too long
in the infected district, whereas an ox or a horse, if once
impregnated, will almost to a certainty pine away and
die. I say almost to a certainty, because I have heard of
cases of oxen and horses recovering after having been
bitten ; but such cases are very exceptional. The younger
the animal, whether horse, ox, or donkey, the better will
he resist the ‘ fly’ poison ” (pp. 202-203).

Tsetse-fly numerous between the Zongwe River and
Zambesi, in the Batonga country to the north of the latter
river.— Tse-tse-fly were also pretty numerous, and must
be very much so later on during the hot months” (p. 208).

Tsetse-fly on the Loanja River, “or rather swamp,”
which joins the Zambesi north of the confluence of the
Chobe.—“ The road from here follows the western bank of
the Loanja for about forty miles, the whole of which dis-
tance is infested with ‘fly,’ and has therefore to be traversed
at night by both oxen and horses. It can be done in two
nights by ox-waggons, but is usually done in three, and
there are two islands in the swamp free from ‘fly,’ to which
the oxen can be driven to feed and rest during the day-
time. These islands are, too, conveniently situated so as to
divide the journey into three easy nights’ work ” (p. 250).

Cattle at Tete, on the Zambesi, ‘originally brought
from Mashonaland; though how they passed the ‘fly’
infested district between Tete and that country is a
mystery to me” (p. 274).

Tsetse-fly in 1891 on the west bank of the Revui River,
a tributary of the Buzi, which flows into the sea just
below the Pungwi, a little south of Beira ; the flies settled
on two horses in a little clearing on a hippopotamus foot-
path “leading through the dense bush on the river's

FLY-BELTS IN RHODESIA. 213

bank.” Sixteen flies were caught on the horses by “ pin-
ning their feet with a knife-blade (the best way of catching
them, as they are as quick as lightning), and the Kaffirs
securing some with their hands.” The greater number of
the flies were caught before doing any damage, and the
rest were frightened away. The horses did not contract
the disease. The native name for the fly on this river is
“impugan” (pp. 410-411).

Low country between the Buzi and Pungwi Rivers
infested with Tsetse-fly (1891).—< After this I was sent by
the Company to try to find a route to the coast along the
watershed between the Pungwi and Buzi rivers, free of
‘fly’ and therefore suitable for a waggon-road. But in
this I was unsuccessful, as I found the whole of the low
country between the Buzi and Pungwi rivers to be
infested with the Tse-tse-fly ” (p. 426).

Tsetse-fly near the Kasaia River [1888], an affluent of
the Majili, which flows into the Zambesi about 20 miles
above its junction with the Chobe.—“ In the afternoon I
continued my journey, and that night passed through an
extensive mopani forest, to the east of the Kasaia River,
in which the Tse-tse-fly still lingers, though in no great
numbers ” (p. 245).

Horses stampeded into the fly-infested mopani forest
by a herd of zebras.—‘“The horses had twice passed
through the fly-infested forest beyond the Kasaia, but as
it was a cloudy day and a high wind was blowing, and the
flies there are few and far between at this time of year, I
do not think they were “stuck,” and, at any rate, they
never showed the slightest sign of being in any way
affected ” (p. 246).

Tsetse numerous in a strip of forest between the
Magoi-ee (or Makoe) and Ungwesi Rivers, Batonga
country (p. 215). [The Magoi-ee River “rises a little
to the south of U-kesa-kesa, and runs north into the
Kafukwi.” The Ungwesi runs into the Magoi-ee. |

122....1893... Dr. F...\Kaxseh.
“Die BEI BismarckBuRG puRCH Dr. R. Burrner
UND (zUM THEIL) von Hauprmann KLING GESAMMELTEN,
BISHER BESTIMMTEN HEXAPODEN.—D1IPTEREN”: Ergebnisse
der Forschungsreisen im Hinterlande von Togo 1890 bis

214 CAMEL-KILLING FLY ON WEBI SHEBELI.

1892 von Hauptmann E. Kling und Dr. R. Bittner
(Mittheilungen von Forschungsreisenden und Gelehrien aus
den Deutschen Schutzgebieten. Mit Benutzung amtlicher
Quellen herausgegeben von Dr. Freiherr von Danckelmann.
VI. Band (Berlin, 1893. Ernst Siegfried Mittler und
Sohn), p. 225).

“Glossina longipalpis (Wied.)” and “Glossina tabani-
formis (Westw.)” [= Gl. fusca, Walk.] are included in the
list of Diptera collected in the neighbourhood of Bismarck-
burg, in the Hinterland of the Togo Protectorate, W. Africa.
The approximate geographical position of Bismarckburg is
8° 12’ N. lat., 0° 52’ EB. long.

123. 1894.

La Nature (1), p. 286.

Note on Glossina morsitans. [I have not seen this. ]

124. 1894. Dr. Donaldson Smith.

«Dr. Donatpson Suitn’s EXPEDITION IN SOMALILAND”
(The Geographical Journal, Vol. IV., p. 530. London :
The Royal Geographical Society). ;

A letter written by Dr. Donaldson Smith during his
expedition to Lake Rudolf, and dated ‘“ Webi Shebeli
(or Webi Erer?), lat. 7° 11’ N., long. 42° 11’ 23” E.,
September 1-3 [1894].”

Camel-killing fly on the Webi Shebeli.

‘‘T have seen one or two flies that I believe to be the
dreaded Camel-fly of the Webi Shebeli, but they are very
rare.”

125. 1894. eB) Johnston (now Sir H. H. Johnston,
K.C.B.).

*“ Report oF THE First THREE YEARS’ ADMINISTRA-
TION OF THE HasteRN Portion or British CENTRAL
Arrica.” Dated March 31, 1894. [With five maps.]
Presented to both Houses of Parliament by command of
Her Majesty, August, 1894. (London: Harrison & Sons.
Printed for Her Majesty’s Stationery Office.) Pp. 18-19.

“Let us now consider the very important question of
horses and cattle, and the possibilities of stock-rearing.
Perhaps the greatest curse which nature has laid on this
and other parts of South-Central Africa is the existence
of the Tsetse-fly. Were it not for this the value of the
country would be centupled. In the first place, the diffi-

TSETSE IN BRITISH CENTRAL AFRICA. 215

culties of colonization would be solved, and also the
difficulties of that conquest which must, sooner or later,
take place if the slave-raiding tribes are to be kept under
control. At present it is only over small and widely
separated patches that horses and cattle can be kept free
from the attacks of the Tsetse-fly. For instance, there is
no Tsetse-fly on the road from Katunga to Zomba, a
distance of over sixty miles. The convenience of this
may be estimated from the fact that I can ride from
Katunga to Zomba with a change of horses in two days,
whereas it would take me three days hard and very
uncomfortable travelling to perform this journey in a
machilla.* On the other hand, I cannot ride over to
Mlanje, which is only some forty miles distant, because
there is a belt of Tsetse-haunted country in between.
Neither can I ride the short distance of twenty-one miles
to the Upper Shiré, for the same reason. Were it not for
this, we could keep the whole Shiré province under control
by a small troop of fifty mounted police. Nevertheless,
we may be thankful that when proper precautions are
taken it is quite possible, and not even very difficult, to
introduce horses into the Shiré highlands. The Tsetse-
fly appears never to go near the edge of a river or to fly
over a river. Consequently, when horses and cattle are
being brought up in steamers they are not attacked by
the Tsetse as long as they do not leave the river ; and,
inasmuch as the piece of shore between Katunga and
Chikwawa is free from Tsetse, we are able to land horses
there and send them up into the highlands, where the fly
is absent. By taking the horses down at night through
the fly-belt between Zomba and the Upper Shiré to
Mpimbi and embarking the beasts at once, they could
also be got up to Lake Nyasa without being bitten.
There is a considerable area round Fort Johnston and
the south end of Lake Nyasa where there is no Tsetse-fly,
and nearly all the country along the eastern shore of the
lake is without this pest. On the contrary, the Tsetse
occurs in patches along the west coast, but is quite absent
from the north end of the lake. The southern shore of
Lake Tanganyika is infested with the fly, which kills even
dogs, goats and donkeys, the two latter beasts generally

* “ \ hammock carried by native porters.—H. H. J.”

216

126.

MEANS OF EXTERMINATING TSETSE.

escaping. Yet over almost the whole of the region coloured
brown on my sketch-map* the Tsetse-fly is absent. It
is scarcely ever found above 3,000 feet. The Luangwa
Valley, on the other hand, is full of Tsetse, so are certain
portions of the Zambezi banks.

‘There is one hope as regards the extermination of
this fly, and that is, that it always dislikes the presence
of human beings and avoids human settlements. It also
seems to depend for its existence on large herds of game.
Therefore, by the increased settlement and population of
the country and the reducing the herds of wild animals,
we may hope in time to exterminate the Tsetse to the
same extent as it has by the past action of man been got
rid of throughout the Soudan, where one never hears of
it, and likewise in the regions of the Niger. It would be
very interesting if the authorities at the British Museum
would compile for use, from the information they possess,
a map showing the distribution of the Tsetse-fly in
Africa ; I mean of course of that species of Glossina, the
bite of which is fatal to horses, cattle and dogs. There
are other species of Glossina whose bite is apparently
innocuous. . . . Still, putting aside these occasional
epidemics or endemic sicknesses, cattle do very well in
such parts of South Central Africa which are free from
the Tsetse-fly, and which are provided with good pasture ”

(p. 19).

1894. Sir Gerald Portal.

“Reports ReLatTiInG TO UGanpA BY Sir GERALD
Porta.” Presented to both Houses of Parliament by
command of Her Majesty, March, 1894. (London:
printed for Her Majesty’s Stationery Office by Harrison
& Sons, St. Martin’s Lane, Printers in Ordinary to Her
Majesty.)

Page 6.—Report by Col. Francis Rhodes to Sir G.
Portal (dated “Kampala, April 2, 1893”) “on the route
between Mombasa and Uganda”: “ We were told on no
account to bring ponies with us, as there was Tsetse-fly
between Zavo and Kibwesi. We saw no sign of it, and
the two ponies we brought with us did admirably, and
improved in condition on the road.”

* Whe Orography of British Central Africa.

GAME SCARCITY IN E. AFRICAN FLY-BELTS. 217

127. 1894. F. J. Jackson.

“Bie Game Suoortine,” Vol. I., pp. 185-186 (The
Badminton Library of Sports and Pastimes. London :
Longmans, Green & Co.).

No reason why horses should not be more generally
employed in East Africa, “provided the belts of ‘fly’
country are avoided” (p. 185).

Tsetse-fly in East Africa not closely associated with big
game.—* It is supposed by a good many people that the
Tsétsé-fly only exists where game beasts, especially
buffaloes, are most plentiful, and that the fly disappears
as the game is killed off or driven away. This may be so
in South Africa, but it is certainly not the case in Hast
Africa, as the belts of fly country in East Africa are
almost devoid of game, with the exception of the river
Tana. As, however, the open, undulating, grassy plains
of the Masai country, and other places of a like nature,
are the headquarters of by far the greatest quantity and
variety of game, and are entirely free from the Tsétsé-fly,
and as they are also well adapted to hunting on horse-
back, the game would very soon be exterminated if
pursuit on horseback were permitted ...” (pp. 185-186).

128. 1894. W. Cotton Oswell.

“ Bia Game Suootine,” Vol. I., pp. 113-115, 147, 150
(The Badminton Library of Sports and Pastimes. London :
Longmans, Green & Co.).

“On the low Siloquana hills [in the north of the
Transvaal, between the Magalaqueen River and the
Limpopo] near this we made our acquaintance with the
Tsétsé-fly, which we were the first to bring to notice
[the year referred to is 1845]; Vardon taking or sending
to England some he caught on his favourite horse.”—
Symptoms and effects of the bite of the fly on cattle
(p. 113).

* Wild animals are not affected, but all domestic ones
are, save the ass and the goat, and the calf as long as it
sucks. Man escapes scot free. The flies settle on him
and bite him sharply, but no results follow” (pp. 113-114).

“This pest, like all others, is held in check by an
antagonist, one of the ichneumons—a rakish-looking crea-
ture which catches and sucks it out on the wing, dropping

218 OBSERVATIONS BY W. C. OSWELL.

the empty cases much as the locust bird does the locusts ”

(p. 114).

[The insect here referred to as attacking the Tsetse
is probably one of the Asilidee—a family of Diptera, the
members of which prey on other insects in the manner
indicated. The Ichneumonide are Hymenoptera, which
lay their eggs in living caterpillars, but do not themselves
hawk after insects.—H. EH. A.|

Cattle take their death-bites quite calmly—‘“ with a
whisk of their tails, as is their custom with other flies”
(p. 114).

Tsetse cn the Chobe River, in the bush, not in the reeds:
the flies begin to buzz about as the sun rises (p. 147).

“We had travelled all night through the sleeping
flies” (pp. 146-147).

“Livingstone woke up, smoothed down my visitor,
and inquired what we could do with the cattle. We
could not leave them where they were; they would find
nothing to eat, and besides, when the sun got hot the
flies would find their way to them. We must drive them
across the river, as there were no Tsétsé there, the man
told us ; and we found that it was so, the narrowest lines
frequently defining the limits of safety and danger”
(pp. 147-148).

“The Tsétsé, that great enemy to the cattle-breeder,
will disappear before the approach of civilisation, and the
killing off of the game, especially the buffalo, its standing
dish, as it has done many times already in African lore.
I am speaking of the tracts south of the Zambesi”

(p. 150).

129. 1894. Jules de Guerne.

Compte-Rendu Sommaire de Séance de la Société Philo-
mathique de Paris.—Compte-Rendu Sommaire de la Séance
du 14 Avril 1894, No. 12, p. 2.

Report of the exhibition of two specimens of Glossina
morsitans, Westw., collected in Central Africa by M.
Edouard Foa.

[Translation.] ‘“ Everything leads us to believe that
Glossina inoculates domestic animals with the germs of
some virulent malady analogous to anthrax. ... No
doubt it would be possible to cultivate the bacteria

DOUBTFUL RECORD FROM SOMALILAND. 219

130.

131.

132.

133.

disseminated by the insects, attenuate their virus, and
perhaps also find a new vaccine capable of saving the
animals necessary for the prompt colonisation of Central
Africa,”

1895. E. Corti.

Annali del Museo. Civico di Storia Naturale di Genova,
Serie 2.*, Vol. XV. (XXXYV.), pp. 138-139.

Original description of Glossina longipennis, from
Somaliland.

1895. le Major David Bruce, A.M.S. (now Lt.-Col.
David Bruce, R.A.M.C.).

“ PRELIMINARY REPORT ON THE TSETSE-FLY DISEASE,
or Nacana, In ZULULAND” (Durban: Bennett & Davis,
Field Street), 28 pp.

[I have not seen this Preliminary Report: it was
reviewed by W. F. H. Blandford, in Nature, April 16,
1896, For the “Further Report,” see below [142], and
also Chapter VII., Appendix A.]

1895. A. Whyte.

“Report oN THE BoranicaL Aspects OF Britis
CentraL Arrica.” Foreign Office, 1895. Miscellaneous
Series, No. 373. Reports on Subjects of General and
Commercial Interest. Africa (Central). — (London :
printed for Her Majesty’s Stationery Office, by Harrison
& Sons, St. Martin’s Lane, Printers in Ordinary to Her
Majesty.)

Tsetse-fly (pp. 16-18).

“This is a pest which will seriously interfere (for
some time at least) with the introduction and rearing of
domestic stock, and thus form a drawback to agricultural
enterprise” (pp. 16-17).

Notes on specimens in the British Museum, with
information furnished by E. E. Austen.

1895. Major H. S. Mainwaring.

“THe Korayo Vauury, Somauianp” (The Geo-
graphical Journal, Vol. VI., p. 474. London: The Geo-
graphical Society).

Tsetse-fly (?) in the Korayo Valley, 1894.

“The Korayo valley, placed by Major Mainwaring
north-west of the Tug Turfa or Turfo, would seem to be

134.

APPEARANCE OF TSETSE AT CHIMOIO.

that of the Erer, visited lower down by Dr. Smith. . . .”

‘The valley itself is described by the latter [Major
Mainwaring] as quite the Eden of Somaliland, being
thickly wooded, and the tops of the smaller hills covered
with fresh green grass. At the bottom of the valley were
found a running stream, luxuriant tropical vegetation,
brilliant-plumaged birds, and bright flowers. It is un-
fortunately infested by a fly, apparently a species of
Tse-tsi, which proved fatal to the animals.”

1895. W. A. Eckersley.

* Notes iv Eastern MasHonatanb” (The Geographical
Journal, Vol. V. London: The Royal Geographical
Society).

Appearance of the Tsetse-fly at Chimoio, Portuguese
East Africa, on the line of the Beira Railway in 1894:
two ponies taken by the survey party in 1893 through the
Jly-belt to Chimoio without suffering any ill effects.

“The temporary terminus of the railway, usually
known as the ‘Seventy-five mile peg, is 43} miles
from Chimoio by the line recently surveyed for the
proposed extension to that point; the distance measured
in a straight line is very considerably shorter. The
Seventy-mile terminus is connected with Chimoio by a
waggon road lately cut through the forest, thus bringing
the railway into direct communication with the ‘Selous’
road to Massi-Kessi, Umtali, and Salisbury. The terminus
is unfortunately situated well within the limits of the district
infested by the ‘ Tsetse-fly,’ thus rendering imperative the
early extension of the line to Chimoio, or even further to
the west. Chimoio, up to the present, has been entirely
free from the fly. The doubt has been expressed that
the presence of a large number of horses, oxen, and other
animals at Chimoio, attracted thither by the facilities of
transport offered by the railway, might in its turn be the
means of attracting the destructive fly to that place.
This doubt has now become a certainty, recent letters
containing the news that the dreaded fly had actually
made its appearance in Chimoio.

“Tt is interesting to note that two ponies, purchased
in Natal for the use of the survey party, passed through
Beira, Fontesvilla, and the intervening ‘ fly-belt,’ to

ESCAPE OF PONIES FROM FLY-DISEASE, 221

Chimoio without suffering any ill effects; they served the
party until the conclusion of the work, and were finally
sold at a profit. No particular precautions against the
‘fly’ were adopted, except occasional brushing with green
boughs. It is quite certain that the Tsetse-flies settled
on the horses in considerable numbers, and remained
quite long enough to allow of their biting. It was only
during one day, however, that the flies were present in
large numbers ; this was on the course of the railway,
between 38 and 45 miles from Fontesvilla.*

“For the first 20 miles the line of railway traverses a
perfectly flat, nearly treeless, alluvial plain, covered with
long grass and teeming with big game, including lions,
buffaloes, most of the South African species of antelope,
wart hogs, and wild boars” (p. 29).

Tsetse-fly in Mashonaland.—* White ants and borers
are the worst enemies of the settlers in the high country,
while the Tsetse-fly causes incalculable mischief in the
low-lying districts” (p. 38).

135. 1895. Edouard Foa.

“ Mus GRANDES CHAssmes DANS L’?ArrIQqUE CENTRALE ”
(Paris: Librairie de Firmin-Didot et Ci), pp. 22, 28-33,
220-221.

[Translation.] ‘The day of our arrival on the banks
of the Crocodile River, we were on the eve of our troubles.
. . . There was something more, that we learnt next day :
we were entering the region infested by the terrible
Tsetse-fly, and we must expect to meet with it at any
moment. It would be the death of our oxen; there was
no remedy in our power, we could only resign ourselves to
our fate” (p. 22).

“There is nothing repulsive or remarkable in its [the
Tsetse’s] appearance to anyone who does not know it; its
flight is exceedingly quick, and it is impossible to distinguish
it in the air before it has fed ; when its abdomen is full of
blood, its flight becomes heavy and it at once hides itself
in order to digest its meal in peace. Owing to its quick-
ness, it is impossible to catch it like an ordinary fly.
When it settles, it does so so gently that one does not feel
it ; it remains thus motionless for fifteen or twenty seconds,

* The starting-point of the Beira railway,

222 FOA’S ACCOUNT OF TSETSE SUCKING BLOOD.

its proboscis projecting in front, in a mistrustful attitude,
ready to take flight. When it believes itself safe, it
lowers its weapon, separates its feet so as to flatten itself
out more and pierces the flesh without causing any pain to
start with, as does the mosquito. . . . While its proboscis,
which is at Jeast a quarter of a centimetre in length, dis-
appears completely in the flesh, it remains motionless
sucking blood, its abdomen swelling and becoming rose-
coloured owing to its transparency, and immediately
afterwards appearing deep red and plump. It is not until
the moment when it has already imbibed a large portion
of its meal, that a slight pain or rather itching betokens
its presence. When its stomach is full it is still very
difficult to catch with the hand, for it does not fly off, but
quickly dodges to one side. The natives and I myself,
when they showed me the way, used to catch it in a
different fashion : the blade of a knife is laid at a distance
of 30 centimetres from the fly flat on the arm or on
whatever part of the body it has settled, and is then
slowly slid along until it meets and squeezes the proboscis
of the fly still buried in the flesh and thus makes the
insect a prisoner * ; then, without relaxing the pressure,
you raise the blade and turn it over, and so kill the fly, or
else seize it with the fingers. Of course you have already
been bitten by the fly, but you console yourself by thinking
that, at any rate, there is one enemy less among the
surrounding thousands.

“‘T have now to speak of the sensations occasioned in
domestic animals by its bite. Of these animals, I shall
mention those that one has to possess in Africa: the ox,
dog, donkey, mule, sheep, pig, and goat. Livingstone
states that the latter and sometimes the donkey are
exempt from the consequences of the bite, while all the
other animals die from it. Iam able to state, after having
made several experiments, that none of the animals [
have mentioned survives it ; it depends simply and solely
on the number of the bites. The local fauna is inoculated
with the poison of the fly from its youth up; it is more-
over from it that the insect obtains its food; but when

* In Sierra Leone my Timminy hammock-boys used to catch Tsetse-
flies (Glossina palpalis) in the same way.—E. E. A.

+ “This way of catching it with a knife seems to prove that the Tsetse
does not see in front of and below it,”

OLFACTORY POWERS OF TSETSE. 223

the Tsetse accidentally meets with domestic animals, it
pursues them with peculiar pertinacity ; the animal has
an instinctive sense of the danger that threatens it: it
jumps, shies, and after the first bite, the noise alone
of the fly literally deprives it of its senses; it loses
its head and bolts, hoping in this way to outstrip the
murderous insect buzzing round it. The venomous fly
comes to its prey from a long distance, whether it is
that its sight is keen or its sense of smell exceptionally
acute ; I would incline rather to the latter hypothesis,
since I have noticed that the fly always comes up wind,
and generally bites by preference on the leeward side.
The Tsetse keeps on the under, not on the upper side of
leaves, as one never sees it, and it prefers shade to sun.
Jt especially dislikes the odour of excrement ; as soon as
an antelope is killed, for instance, in order to get rid of
the Tsetse which literally cover game and hunters, it is
only necessary to open the animal’s belly and evacuate
the entrails ; the insect at once ceases to torment you”
(pp. 29-31).

Symptoms of Tsetse-fly disease in cattle (pp. 31-32).

The interval that elapses before death depends upon
the number of bites (p. 32).

The Tsetse follows the big game, of the existence of
which in a district its presence is a proof. “It is parti-
cularly partial to buffaloes and the large antelopes ; that
is why one finds districts containing small game devoid of
Tsetse.” The fly will disappear simultaneously with the
destruction of the big game (pp. 32-33).

“Thave still to speak of the sensations experienced by
a human being after being bitten by the poisonous insect.
The itching feeling changes in a few seconds to a painful
pruritus which lasts about a quarter of an hour ; the bitten
part reddens, swells slightly, and continues to be uncom-
fortable for a short time (pendant un moment). A large
number of bites may undeniably have a disturbing effect
upon the organism; above all, they have the effect of
exciting one excessively, and a feeling of rage is added to
the suffering. The parts affected by the fly in the case of
man are generally those that are uncovered : hands, arms,
neck, cheeks, and legs, always on the under side as much

as possible ” (p. 33).

224 ENCOUNTER WITH SWARMS OF TSETSE.

Swarms of Tsetse on the Kapotché River.—* The momen-
tary sense of comfort experienced in this wood was very
quickly forgotten ; never had I seen round us such swarms
of Tsetse ; it was simple torment and it was impossible to
remain seated. The accursed insects seemed to be parti-
cularly infuriated with us, and each one of the 200 men
had a large detachment to himself.

“T found it impossible to breakfast; one was bitten
on the arms, legs, neck—in twenty different places at
once. I had with me a tame goat, who was very amusing.
... The poor beast died under a swarm of Tsetse, in
the space of an hour: maddened by the bites, he had
begun by roljing on the ground, then foam gathered on
his lips, his belly became swollen, and, when I noticed
what was happening, he was already dying.

“ After a few hasty mouthfuls, I caused the camp to
be struck and everyone was very glad to be off. Never
had I seen so many Tsetse together in the same spot. This
must be their village, as Maonda said. We left there the
body of my goat, whose sufferings I had terminated by a
knife-thrust ” (pp. 220-221).

136. 1896. W. F. H. Blandford.
“ THE TsErse-FLy Disease” (Nature, No. 1381, Vol. 53,
April 16, 1896, pp. 566-568, Figs. 1 and 2: wood-cuts of
Glossina sp., larva, puparium, and hematozoa of Nagana
in the blood of a horse).
A review of Surgeon-Major Bruce’s Preliminary Report
on the Tsetse-fly Disease, or Nagana, in Zululand.—{ Vide

supra, 131.]
137. 1896.

The Field, April 11, 1896. An article on the

Tsetse-fly.
[I have not seen this. ]

188, 1896. Sir Harry Johnston, K.C.B.
Report By Commissioner Srr Harry Jonnston,
K.C.B., on THE TRADE AND GENERAL CONDITION OF THE
British Centra ArricA ProrectoraTr. April 1, 1895,
to March 31, 1896. [With a Map.] Presented to both
Houses of Parliament by command of Her Majesty.
August, 1896. (London: Printed for Her Majesty’s

139.

GALLA MODE OF CROSSING FLY-BELTS, 225

Stationery Office by Harrison & Sons, St. Martin’s Lane,
Printers in Ordinary to Her Majesty.)

“We must encourage the unchecked increase of the
negro population in British Central Africa, for the pre-
sence of millions of men with strong arms, and relatively
proof against malarial fever, or, at any rate, amongst
whom life is very cheap, will soon dispel malaria, the
Tsetse-fly, and other drawbacks to the utility of British
Central Africa, . . . The Tsetse-fly loathes the presence
of man, and is already fast disappearing from the Pro-
tectorate. It is never, in any case, found in the hills, and
we have only to get those great riverain plains and lake
shores densely peopled to be able to chronicle its entire
extinction” (p. 12).

Disappearance of the Tsetse-fly from the country to the
south of Lake Nyasa.—The bringing down of cattle from
Lake Nyasa to Blantyre ‘‘has shown in many cases how
completely the Tsetse-fly has disappeared from regions
where formerly its presence in large numbers would have
killed almost all the cattle that crossed the infested
territories ” (p. 15).

1896. J. W. Gregory.

“ Tue Great Rirr Vautey” (London : John Murray),
pp. 21-22.

Tsetse-fly in belt of forest between Witu and the coast.—
“‘ There were no preparations for firing the belt of forest
at Pangani through which the road passes, and which is
the haunt of the Tsetse-fly (Glossina morsitans, Westw.).
On the journey through it on our way up, I had found
this dreaded fly. To expose our camels to an attack from
it was to risk the whole safety of the caravan; I had
therefore sent back a note to say that before the baggage
animals entered this belt of forest, fires ought to be
lighted in it and the animals driven through the smoke.
This is the method commonly used to protect animals
when passing through a fly-haunted district. I found out
afterwards that the Galla in the Tana valley know all the
fly-infested patches, and always adopt this precaution
when driving their cattle through them. As Dr. Rae
seemed better, he went on with two men, and I stopped
there to prepare the fires, sending on a note urging that

Q

226

140.

141.

142,

148.

TSETSE ON ZAMBESI ABOVE SESHEKE.

these should be lighted half an hour before the camels
were allowed to enter the forest. The rest of the day was
spent in building up stacks of wood and dead leaves ;
these were arranged as far as possible on the model of the
‘smudge-fires’ used in Manitoba to protect the wheat
from the early frosts of August, by raising a cloud of
smoke above the fields.”

1896. W. Donitz.

Berliner Entomologische Zeitschrift, 41. Band, pp.
(17)-(18).

Report of a paper on Tsetse-flies, containing a summary
of previous statements,—No new facts.

1896. Alfred Sharpe.

‘‘THEe GEoGRAPHY AND Resources oF British CENTRAL
Arnica” (The Geographical Journal, Vol. VII.). (London:
The Royal Geographical Society).

Tsetse-fly absent from the highlands of British Central
Africa.—< In the high parts of B.C.A. cattle and horses
thrive, and there is no Tsetse-fly ” (p. 384).

1897. Surgeon-Major David Bruce, A.M.S. (now Lt.-Col.
David Bruce, R.A.M.C.).

‘“FurtHer Report on THE TsetsE-FLty DISEASE OR
Na@ana IN ZULULAND.” Ubombo, Zululand, 29th May,
1896. (London: Harrison & Sons, St. Martin’s Lane,
Printers in Ordinary to Her Majesty. 4th February
1897.) 69 pp., Plates I-VI.

[See Chapter VIT., Appendix A.}

1897. Captain A. St. H. Gibbons.*
« A JOURNEY IN THE Marotsrk AND MASHIKOLUMBWE
Counrrizs” (The Geographical Journal, Vol. IX).
Tsetse-fly swarming on the banks of the Zambesi, 35 miles
above Sesheke, July, 1895.—‘ Some 35 miles from Sesheke
brought me to high banks covered with forest and teeming
with Tsetse-fly. Such is the general character of the river
as far as the southern extremity of Borotse” (p. 123).
Tsetse-fly absent from the Borotse “Valley,” which extends
along both banks of the Zambesi from 16° 15' S. lat. to con-
siderably beyond Lialui.—* On the second day after this I
reached the southern extremity of what is frequently

* Cf. [x52].

TSETSE IN EAST AFRICA PROTECTORATE. 227

144,

described as the Great Borotse ‘ Valley’ (16° 15’ 8. lat.).
Livingstone first spoke of the Borotse as a ‘valley,’
though ‘plain’ or ‘flats’ would convey a more correct
idea of what in reality is a huge treeless alluvial plain in
places 50 or 60 miles wide, and extending a very con-
siderable distance to the north of Lialui, which stands
70 miles as the crow flies from the southern boundary of
the plain. In the winter season the Borotse yields an
excellent cattle pasture, and, being free from the ‘Tsetse ’-
fly, supports many thousand head of cattle” (p. 126).

Tsetse-fly absent from the grassy valley through which
Jlows the Njoko River, above and below its confluence with
the Rampungu (p. 134).

1897. Sir A. Hardinge.

“Report By Sir A. HarpDINGE ON THE CoNDITION
AND Progress oF THE East Arrica PRoTEcTORATE FROM
its EsTABLISHMENT TO THE 20TH JuLy, 1897.” [With
Map.] Presented to both Houses of Parliament by com-
mand of Her Majesty. December, 1897. (London: Printed
for Her Majesty’s Stationery Office by Harrison & Sons,
St. Martin’s Lane, Printers in Ordinary to Her Majesty.)

“It may appear somewhat remarkable that the Pro-
vince of Ukamba, the most uncivilised division of the
territory, should be the best supplied with roads, but it
must be remembered that the coast provinces carry on
their communciations with one another to a great extent
by sea, and secondly, that over a large portion of them
the climate conditions and the presence of Tsetse-fly pre-
clude the animal transport by carts, which in the interior
is the great incentive to road-making. In Witu, for
instance, where the Administration built a good carriage-
road from the capital to the Port of Mkunumbi, the
bullocks employed for the waggons on it all died, and the
old wretched system of human porterage has still to be
resorted to for transport, even donkeys not being pro-
curable in sufficient numbers” (p. 51).

“ Along the Uganda Road transport by carts drawn by
bullocks has been successfully initiated. Between Mazeras
and Kibwezi there are, however, several tracts infested by
Tsetse. 2°...” (p. 52).

Cattle flourish in Ukamba (p. 52).

Q 2

228
145.

INFLUENCE OF TSETSE ON HISTORY.

1897. Sir H. H. Johnston, K.C.B.

“British Centrat Arrica” (London: Methuen & Co.),
pp. 54, 64, 367, 377-380.

In Chapter ITI., dealing with the “ History ” of British
Central Africa, the author writes (p. 56):—‘ At first
Sena, on the Lower Zambezi, was the headquarters of the
Portuguese Administration, and from hence various expe-
ditions, during the sixteenth century, were sent southwards
to discover the gold mines of Manika—expeditions which
were mostly unsuccessful, owing to the unhealthiness of
the climate and the presence of the Tsetse-fly.”

Indian buffaloes, introduced into Africa by Livingstone,
not affected by the bites of the Tsetse-fly.—‘‘ Livingstone also
tried [in 1866] to introduce the Indian buffalo,* an experi-
ment not repeated until my reintroduction of this animal
from India in 1895. It is interesting to note that Living-
stone’s buffaloes passed through the Tsetse-fly country,
and, seemingly, were not affected by the bites of that
insect, though they all subsequently died as the result of
maltreatment at the hands of the sepoys” (p. 64).

*“ And now we come to the Tsetse, perhaps the most
serious of all the many insect pests of Africa in its check
to European enterprise. It is difficult to overestimate the
importance of the part played by this noxious little insect
in preventing the opening up of Central Africa.

“This was first experienced by the earlier Portuguese
expeditions of five hundred or six hundred mounted men
which would set out from Sena on the Lower Zambezi in the
sixteenth and seventeenth centuries to secure the gold
mines to the north and south. We read in Portuguese
records how their horses soon succumbed to the attacks of
a fly. The riders were left without steeds and the expe-
ditions came to an abortive termination, many of the
Europeans dying of fever or succumbing to the attacks of
the natives through having to make their way on foot.
But for the Tsetse-fly the whole history of South-Central
Africa would be different. It would have been rapidly
traversed by mounted men, not nearly so much ill-health
would have pursued explorers and pioneers forced to travel
on foot, and the whole question of transport would be
rendered infinitely more easy, as coaches and waggons

* Cf. [49].

TSETSE IN BRITISH CENTRAL AFRICA. 229

could run and huge numbers of pack-animals—horses,
mules and oxen—might convey goods which at present
are carried on men’s heads. Undoubtedly the Tsetse-fly
has checked the southward range of the Muhammadan
raiders from the north. But for the presence of this
insect in the Congo Basin and in Equatorial East Africa,
the Muhammadanised negroes and Arabs of the Sudan
would have spread much farther south than they have done
already on their sturdy little ponies” (p. 377).

“Fortunately the Tsetse-fly is not present in all parts
of British Central Africa. Roughly speaking, it may be
said that it is absent from any district that is above
3000 feet in altitude, and is not found in many of the low-
lying lands for some hitherto unexplained reason, no
doubt connected with human settlement. It is present
throughout the whole valley of the great Luangwa River
from the Zambezi to the verge of the Nyasa-Tanganyika
plateau. It is found on part of the upper course of the
River Luapula and on the shores of Lake Mweru, but is
absent from the greater part of the country round
Bangweolo. It is most abundant on the south coast of
Tanganyika, disappearing, however, as soon as the slopes
of the Nyasa-Tanganyika plateau are reached. On Lake
Nyasa it is absent from almost the entirety of the east
coast. On the west coast it is met with between Deep
Bay on the north and the River Bua on the south, some
patches in between, however, being free from it. From
about Kotakota and the Marimba district it is absent. It
reappears again south of the Marimba in the northern
part of the coastlands of the Central Angoniland district.
From the south coast of Lake Nyasa it is almost entirely
absent, but it is found again on a small portion of the
Upper Shire.

“In the low-lying country round Lake Chilwa up to
the slopes of Mlanje and the hills near Zomba it is present.
On the Central Shire at Chikwawa and Katunga there is
no Tsetse, but in the Elephant Marsh below it abounds,
as also in much of the Ruo district and in the district of
the Lower Shire. Always, however, when the land rises
to 3000 feet and beyond the Tsetse disappears. This
insect has a great dislike to water and a still stronger
dislike to a congeries of human habitations, In conse-

230 CONVEYANCE OF ANIMALS BY WATER.

quence, it is possible to convey horses and cattle up the
rivers without the least danger of their being bitten, as
long as they remain on the boat anchored in mid-stream.
They are also quite safe in the middle of any collection of
huts or in any town. Itis a fortunate thing that there is no
Tsetse at Katunga or Chikwawa on the Central Shire, as
live stock can be brought the whole way by water to this
place from the mouth of the Zambezi,* landed there and
sent up to Blantyre, and can thence be conveyed by various
routes which are free from Tsetse-fly to the Upper Shire
and so on to Lake Nyasa. Another important fact to be
borne in mind is that the Tsetse-fly does not bite at night,
therefore if a Tsetse-haunted district must be crossed it
should be done at night-time—by moonlight if possible.
It is said also that smearing the bodies of the animals with
cow-dung will repel the insect ” (pp. 377-378).

[Five sketches of Tsetse-flies, 14 times life size, are
given on p. 378, but are not sufficiently accurate to be
useful. |

‘Donkeys are far less subject to the poisonous cha-
racter of its bite than horses or mules ; indeed, it is said
that the domestic donkey of East Africa, which is only
one degree removed from the Abyssinian wild ass, is
impervious to its attacks, and certainly none of those
animals have died from Tsetse bite in British Central
Africa. Major Lugard, I believe, has found on his expe-
dition to Lake Ngami that his donkeys were the only
animals that survived the attacks of the Tsetse. Dogs
are killed by it, and even cats will not resist its attacks
when too frequent. On the Mwanza River, an affluent of
the Shire nearly opposite to Katunga, the Tsetse are so
numerous that the only domestic animals which can be
kept by the natives are fowls” (p. 379).

Notes on the distribution of Tsetse-flies in Africa
generally, from specimens in the British Museum (p. 379).

“In the greater part of the Nigerian, the Central and
the Egyptian Sudan the Tsetse is absent, thus permitting
a far more rapid and healthy development and conquest
of these countries, as horses are abundant and can be
employed to mount cavalry and transport travellers, while

* “The Tsetse is apparently absent from Chinde and Quelimane and
much of the Zambezi Delta.”

CONDITIONS IN CONGO FREE STATE. 231

for trade purposes mules and oxen can be employed, and
an unlimited number of cattle might be reared” (p. 379).

Theories as to the nature of the “Tsetse poison”
discussed (p. 379).

“Certainly the Tsetse tends to disappear before the
presence of man, and the one certain cure for it would
seem to be the placing of all the low-lying parts of British
Central Africa under cultivation, and the settlement of
innumerable negroes. Fortunately the fly does not much
trouble our political economy, for the further reasons that
so much of the country lies above its habitat. In those
districts where it is healthy for Europeans to settle, the
altitude is already too great to permit of the existence of
the Tsetse-fly ” (p. 380).

146. 1897. Lieut. Th. Masui and — Seeldrayers.

“@uIDE DE LA Section pe L’Erar InpiPENDANT DU
Conco A Exposition DE BRUXELLES-TERVUEREN EN 1897”
(Bruxelles: Imprimerie Veuve Monnom), p. 300.

The common house-fly [Musca domestica, L.] said to
cause the Tsetse to disappear !

[Translation.] “ We will not speak of the FLY oF
Europe, which is already acclimatised everywhere, and,
it seems, should have the advantage of bringing about the
disappearance of the Tsetse, that enemy of stock. . . .”

Further on, pp. 306-310, horses, donkeys, and mules,
all three of which have been introduced, and cattle, goats,
sheep, and dogs are mentioned as existing in the Congo
Free State, and nothing is said as to their suffering from
the attacks of the Tsetse-fly.

147, 1897. Edouard Foa.

“Du Cap av tac Nyassa” (Paris), p. 148.

[Apud Laveran & Mesnil, “ Recherches morphologiques
et expérimentales sur le Trypanosome du Nagana,” etc.,
p- 5, note 2.]

“There cannot be any doubt as to the harmlessness
of the bite of the Tsetse in the case of wild animals,
especially buffalo and the large antelopes” (p. 148).

According to Laveran and Mesnil the author states
that he was bitten thousands of times by Tsetse-flies,
without experiencing anything beyond very slight local
effects, and a feeling of irritation against the fly.

232
148.

149.

DONKEYS AND TSETSE-FLY DISEASE.

1897. Mary H. Kingsley.

“TRAVELS IN West Arrica” (London: Macmillan
& Co.).

Tsetse-fly on the Gold Coast.

A made road running up country on the Gold Coast
“cannot be of use for draught animals, because of the
horse-sickness and Tsetse-fly which occur as soon as you
get into the forest behind the littoral region” (p. 637).

1897. Aurel Schulz, M.D., and August Hammar.

“THe New Arrica: A Journey up the Chobe and
down the Okovanga Rivers. A Record of Exploration
and Sport” (London: William Heinemann), pp. 67-68.

Tsetse-fly on the swampy off-shoots from the Chobe River,
July, 1884.

“Oh, those horrid swamps, worse for the reason that
our donkeys were badly Tsetse-fly bitten, and the immer-
sion of their hides in the water would inevitably hasten
their death, although donkeys show more resistance to
‘fly-bite’ than any other domestic animals. But, given
such swarms of ‘fly’ as we encountered, even this hardy
brute will succumb, as ours eventually did, to the virulence
of this pest. It has often been asserted, not without
mild proof, that the donkey of all domestic animals is the
only one that will survive the Tsetse-fly bite. I myself
have seen them traverse safely the distance from Delagoa
Bay to Lydenburg in the Z. A. R. in 1874, when that
country was thickly infested with flies that killed cattle,
horses, and even the few camels that were imported as
an experiment. But when bitten by such overwhelming
swarms as we experienced, poor Jack and Jenny succumb
in the usual way. ‘The most sensitive animal to fly-bite
is the horse, then the dog, next the ox, and last the
donkey. ‘The bites of five Tsetse-flies have been known
to kill a horse, while it takes a much larger number to
kill a dog or ox. Experiments have been made by
travellers to counteract the effect of fly-bite by a wash of
ammonia or sheep dip. Whatever effect these may have
for the moment, the subsequent and continuous bites
generally are victorious in the end. My esteemed friend,
Mr. Reuben Beningfield, of Durban, however, declares
that he has cured animals only slightly bitten, and

ALLEGED IMMUNITY OF GOATS. 233

hastily isolated from further molestation, by a wash com-
posed of ingredients only known to himself,

“The ‘fly’ poison is injected into an animal in the
same manner as the mosquito bites, and leaves a small
ditfuse lump on the bitten spot, causing much local
irritation, especially as the fly attacks the least exposed
parts of the animal, where the skin is thin, and it cannot
be driven away by the tail. The bite is much aggravated
by contact with water, and it is a well-known fact that
the early rains carry off all fly-bitten cattle. The bitten
animal shows signs of great lassitude, its head swells,
and the joints and limbs become stiff. At this stage a
merciful bullet is advisable to forestall a death of general
debility and asphyxia. On cutting the beast open, one
finds the subcutaneous tissue injected with a yellow serous
fluid not unlike the result of some snake bites, and also
the lungs injected. Beyond this I have not been able to
investigate, as these episodes usually occur on a march
when neither time nor opportunity is offered for scientific
observation.

“The goats we had with us showed no signs of being
affected by fly-bite, and I feel safe in the assertion that
they are impervious to this poison. To mankind and all
wild beasts the bite is innocuous, although surprisingly
unpleasant in its sting, equal in quality to that of a good
healthy wasp or bee. To illustrate the violence of the
bite, let me relate that on one occasion I had already
sighted the rifle at a buffalo standing looking at me some
thirty yards off, and was just in the act of pulling the
trigger, when a Tsetse-fly settled on my hand, and the
sting was so acute that I had not the nerve to pull
the trigger, but had first to brush the fly off, and by the
movement scared the buffalo away.

“The fly on the Pungwe River is, I think, slightly
smaller and darker in hue than those on the Upper
Zambesi or Chobe rivers” (pp. 67-68).

On the Chobe, August, 1884.—Death of a donkey from
“fly-bite”: ‘the other two were almost unable to walk
from stiffness brought about by the same cause” (p. 163).

“ Jumbo the big donkey showed severe symptoms of
fly-bite on the 13th August. In the morning he was
quite stiff and unable to rise, but after being set on his

234 TSETSE-FLY DISEASE IN SOMALILAND.

feet, struggled about until the stiffness went off a little
and he was able to get about and graze. In the evening
he was able to walk a little. A diffuse swelling extended
from between the hind legs, reaching along the belly up
to the forelegs. Otherwise there appeared to be no dis-
tress in breathing, or any increased temperature judged
by feeling the ears” (p. 167).

150. 1898. Prince Nicholas D. Ghika.

“Cinq Mois au Pays pes Somauis” (Georg & Co.:
Genéve et Bale), pp. 131-132.

Tsetse-fly destroying cattle and horses in the Aulihan
Country, Somaliland (on the north bank of the Prince
Ferdinand River, and south of the Webi Shebeyli), in
January, 1896.

[Translation.|—“ Another consideration inclined us to
listen to them ; that is to say the presence, in the Aulihan
Country, of the Tsetse-fly: one after another several
camels have succumbed to its bites, and this very morning
one of our horses collapsed suddenly (est tombé foudroyé)
at the moment that it was being bridled; another, also
bitten, is so ill that it cannot be long before it has to be
left by the wayside.”

151. 1898. A. H. Neumann.

“ KLEPHANT-HuNTING IN East EquaroriAL AFRICA.
Being an Account of Three Years’ Ivory-Hunting under
Mount Kenia and among the Ndorobo Savages of the
Lorogi Mountains, including a trip to the north end of
Lake Rudolph” (London: Rowland Ward, Limited),
pp. 8, 28, 142, 145, 160, 161, 162, 163, 282.

December, 1893.—Mombasa to Laiju (“a district on
the north side of the Tana, and close to the foot of the
Njambeni or Jambeni range, which is a little east of
Mount Kenia”)—‘“I ventured to disregard advice to
take the Tana River route—involving a sea voyage, a
fresh organisation, and a journey through difficult and
unhealthy ‘fly ’-infested bush all the way, with little
useful help from canoes (which could not take animals)
against the stream—and elected for the overland one
through northern Ukambani” (p. 8).

Return to Laiju, April 14, 1894.—“The news that
awaited me was far from cheering. Although before I left

TSETSE AND LACK OF GAME ON ATHI RIVER. 235

many of my donkeys were already dead, I had hoped that
when those that had suffered most from ‘fly’ in passing
through the Tana valley * had succumbed, the remainder
would keep healthy ; for I did not then know that Laiju
was one of the most deadly places in Africa for domestic
animals, not excepting the generally hardy ass. But on
my return now I found that all were either dead, dying,
or sick” (p. 28).

On the south bank of the Athi River, above its junction
with the Tsavo, May, 1895.—‘It is pleasant to get on
the banks of this river (for which I always retain an
affection), with its green willows and fresh scent, and
its wide, though shallow and rather muddy, stream, and
to ‘have an unstinted supply of good sweet water... .
The river runs through the same barren, scrub-covered
country—in some parts gravelly ridges, in others sandy
flats bordering its banks... . One thing that strikes
me about this country is the scarcity of game; for
even along the banks of the river there is marvellously
little—a very few waterbuck at intervals, an occasional
lesser koodoo or two, a little lot of impala, sometimes,
at one particular point, and rarely an odd bushbuck, are
all that is seen, without counting the diminutive paa, and
now and again a rhino, giraffe, or hippo spoor. Formerly
there was a sprinkling of buffalo in small herds ; but these
the cattle-plague swept off. The other game, however,
was always scarce. It is easy to say that the conditions
are not suited to its requirements—too much scrub and too
little grass—but one cannot help wondering why some
species have not become adapted to the land. Here are
great stretches of uninhabited bush country with a peren-
nial river running through it, and hardly any animals,
though plenty of birds and of ‘ fly’ (Tetse) ” (pp. 141-142).

July, 1895.—Tsetse “‘ very numerous” in thick bush to
the north of the Tana (approximate geographical position,
from the author’s map, 0° 10’ S., 38° 10’ E.) (p. 161).

July, 1895. Laiju to northern extremity of Jambeni
hills—“ The number of my donkeys was reduced some-
what, as one by one they succumbed to the effects of the
‘fly’ we had come through; the numerous streams, too,
caused much delay and trouble with the pack-animals ;

* [A “wide shallow valley full of monotonous dense scrub” (p. 10).]

236 TSETSE ON LAKE RUDOLPH.

but, when once the point of the Jambeni hills is weathered,
the country becomes more open and the grass less rank,
and from there onward it is healthy for them” (pp. 161-
162).

tay dt 1895. On the eastern shore of Lake Rudolph,
near the northern end.— Another frequent and disagree-
able accompaniment of abundant game was a fly, two
specimens of which I caught, that I have always taken
for the female of the ‘ Tetse,’ though in this I may be in
error. It is larger than the typical one and of a uniform
brownish colour with no markings, but in other respects
exactly similar both as to appearance and habits,* and I
have always before noticed it associated with the smaller
striped variety, though I saw none of the latter kind here.
Whether this kind is poisonous or not I am not sure. My
donkeys never suffered from having passed here; but then
they can stand a few ‘fly,’ though where the ‘Tetse’ is
numerous they soon succumb” (p. 282).

152. 1898. Captain A. St. H. Gibbons.+

“EXPLORATION AND* Huntinc in CENTRAL AFRICA
1895-96 ” (London: Methuen & Co.), pp. 61-65, 225, 228,
326.

On the left bank of the Njoko River, near its confluence
with the Zambesi, August, 1895.—‘ Many of the northern
tributaries of the Zambesi, unlike the dry sand rivers
of South Africa, flow through wide alluvial valleys, occa-
sionally quite 1000 yards in width. These valleys, though
dry in winter, become swampy in the rainy season. The
rich soil produces excellent cattle pasture, capable of
sustaining vast herds in those districts which are not
infested by the Tsetse-fly. This cruel little pest is parti-
cularly numerous on the lower reaches of the Njoko
River.{ The excessive attention they paid to the back of
my neck resulted in boil-like lumps, which at one time
threatened to give much pain and inconvenience ; but zinc
ointment and a protecting handkerchief proved a rapid
and efficacious remedy.

* [The fly here alluded to is obviously Glossina fusca, Walker, one of
the two large species of Tsetse-flies: the other large species is Glossina
longipennis, Corti, which is found in Somaliland.—E. E. A.]

+ Cf. [143].

} (Latitude, according to author’s map, 17° 8’ 4” to 17° 4’ 8” S.]

CAPTAIN GIBBONS’ OBSERVATIONS, 237

“The Tsetse is in reality very little bigger than the
English house-fly, though his wings being longer he appears
to be much larger. The fore part of the body is so hard
that more than an ordinary pinch is necessary to deprive
this insect of life. I have frequently thrown flies away
for dead after giving them a vigorous squeeze, only to
see them fly away before reaching the ground... .
The proboscis of the Tsetse protrudes in a horizontal
direction and does not point downwards, as is the case
with other flies. It is about one-eighth of an inch long,
and penetrates the skin through a thick flannel shirt
without an effort. The fly is frequently to be heard giving
vent to the high-pitched buzzing note which gives it a
name, but when advancing to attack he noiselessly makes
straight for his mark without all the preparatory fuss
employed by others of his genus. His tread is so light
that the sharp prick of the proboscis is generally the first
indication of his whereabouts. The Tsetse avoids open
plains and is only to be found in forest or bush, and even
there the limits of his habitat are so clearly defined, and
the fly-belts so permanently established, as to give rise to
much speculation as to the reason why one of two con-
tiguous districts of a similar character should teem with
‘fly,’ while the other is quite free from the pest. Certainly
where buffalo is thick the Tsetse is numerous—generally,
at least—but this rule does not necessarily apply to most
game. Districts occur in which game abounds, which,
though within measurable reach of fly-belts, are perfectly
free of their presence. There is much mystery and conse-
quent speculation about the nature and peculiarities of the
Tsetse. Hard facts are known well enough, but the
scientist has not yet arrived on the scene who can explain
its raison d’étre and the paradoxes of its nature. It is
commonly supposed in South Africa that the fly lays its
ova in the skin of the wild buffalo, but this is not so, as
experiments by Mr. Trimen, formerly curator of the Cape
Town Museum, have proved ; still, where the wild buffalo
is to be found in large numbers the Tsetse invariably
teems, and yet the domestic ox succumbs more readily to
the bite than any other animal, except perhaps the horse,
whose first cousin the zebra wanders through belts unhurt,
So, too, the wild dog and jackal are impervious, but few

238 INHERITED IMMUNITY TO FLY DISEASE.

domestic dogs survive the bite many months. On the
other hand, native dogs whose ancestors have been bred
in the fly country for many generations do not succumb
to the poison. The same rule applies to goats reared
under similar conditions ; though it would seem it must
not be applied to sheep or cattle. Of all domestic animals
the lowly donkey alone makes a good fight of it. As high
a proportion as four donkeys out of five have spent a
whole season in the fly country without signs of the poison
taking effect ; though donkeys will, it is believed, at times
die of fly bites in the second season after being bitten. As
a rule, animals bitten by flies in the dry season will live
till the first rains fall, when they die within a few days.
In the same way a horse if bitten will generally die within
twenty-four hours of being driven through a river... .
Like the ‘horse sickness’ and malarial fever, this curse
to travel and transport undoubtedly recedes before the
‘advance of civilisation, so that the far future may yet see
the extinction of the Tsetse.

‘Oswell reported the existence of the fly some 600
miles south of the Zambezi, when he hunted there fifty
years ago. Now waggons can be taken from Bechuanaland
to the Zambezi without any danger of the oxen being
‘stuck.’ Several flies are necessary to produce a fatal
effect, but in passing through a belt in the day-time
several are forthcoming. At night the danger is very
small, though it is a mistake to imagine that the Tsetse
keeps such early hours as other flies. I have at times
been worried by them an hour after the sun has gone
down, and have known flies to buzz into my tent as late
as 9 o'clock on a dark night and make a bold dash for
supper at my expense. At that time of night they are
easily caught, and almost np shaper found their way into
spirits of wine.

“ With all their faults, these destructive little creatures
have the merit of being clean feeders. The natives, in
taking an animal through a fly-belt, plaster it with cow-
dung, which effectively keeps the fly at a safe distance. . . .”
(pp. 61-65).

From the head-waters of the Sejlefula River to the
Nanyate: Latitude of evening camp, as given on author's
map, 16° 47’ 9" 8.; approximate longitude, 25° 48' E.—

TSETSE ON HIGH GROUND. 239

Tsetse-fly numerous.—“ In the morning of the following
day [Dec. 26, 1895] camp was struck, and I travelled
through a country still pleasant, but not quite so useful-
looking as that through which I had passed during the
preceding few days. The Tsetse-fly became much more
numerous, and the native population very sparse. In the
evening, after travelling seventeen miles and crossing two
Machili tributaries, I camped on a third, the Nanyate, at
some distance from its source—probably ten or twelve
miles” (p. 225).

Camp on rising ground in mopani swamps near the
Nanzela River (an affluent of the Kafukwe) ; approximate
position, from the author's map, 16° 29'8., 26° 5 E.—‘*On
turning out next morning [Dec. 30, 1895] a large herd of
wildebeest was to be seen standing about 200 yards from
the tent, gazing intently at what to them must have been
quite phenomenal. .. . The Tsetse here were very trouble-
some, and for some little time after sunset went for us so
boldly that I was able to consign many of them to my
collecting-tube” (p. 228).

Tsetse-fly very troublesome on high ground to the south
of the head-waters of the Nanyate: approximate position,
JSrom the author's map, 16° 57' 1" §., 26° 4’ E.—* The next
day—the 21st of February [1896|—I camped on the
highest point of the plateau traversed ; my observations
made it 4110 feet above the sea-level. The Tsetse-fly had
been very troublesome on this high ground, which at this,
the wet season, is as a rule teeming with game; but so
deadly had the rinderpest proved itself that only once had
game been seen since leaving Bisi. Deprived of the blood
of beasts, they collected themselves and attacked the boys
and myself to some tune” (p. 326).

153. 1898. Captain the Hon. Arthur Lawley.

“From Butuwayo To THE Vicroria Fauus” (Black-
wood's Magazine, Dec. 1898, p. 748).

‘‘ Before the rinderpest, this neighbourhood [the banks
of the Deka, an affluent of the Zambesi] was wonderfully
rich in all sorts of game, which seemed to have suffered
more here from the plague than in any other part of
Matabele-land. The buffalo has entirely disappeared, the
koodoo (formerly very plentiful) are almost extinct, and

240 TSETSE AND BIG GAME.

the other varieties have suffered severely, especially the
wild pig.

“TJ afterwards learnt from the Barotse that before the
Basuto invasion this district was thickly inhabited and
rich in cattle. The Tsetse-fly was not then known between
Deka and Gwaai to the east and Deka and the Zambesi
to the north. After their invasion the natives were either
killed by alternate raids from the Barotse and Matabele,
or else they moved elsewhere, taking their cattle with
them. Then the game returned, and with the game came
the Tsetse-fly. Two years ago, the fly were very numerous,
and the country east of the Pandamatenka road was on
this account impassable for cattle and mules. Now that
the game has been almost exterminated by rinderpest, the
fly also has left the country, excepting a few belts of thick
bush, and apparently it is gradually disappearing. All of
which goes, in my opinion, to prove that the Tsetse-fly
moves with the big game of the country, and that, with
the latter, it is bound to disappear before the advance of
civilisation.”

154, 1808. W. W. A. Fitzgerald.

“ TRAVELS IN THE CoASTLANDS OF BritisH East AFRICA
AND THE IsLANDS OF ZANZIBAR AND PremBa” (London:
Chapman & Hall, Ltd.), pp. 355-357, 358, 419, 432, 434.

“One impediment to the future agricultural develop-
ment of this part of the country (existing in East Africa,
however, to a much less extent than in any other part of
the Continent) is the presence of the Tseftse-fly. A variety
of it, and another species, a gad-fly, are found in Witu.
In the lower coast region, between Mombasa and Uganda,
the Tsetse-fly is said to be localised in a comparatively
narrow belt, easily passed during the night, and there is
no doubt that the destruction of cattle and game during
the epidemic considerably lessened the numbers of this
insect pest. In Witu, too, the variety known there
appears to be confined to a very limited area, chiefly forest,
and in which Utwani is included. The Wa-Galla are
accustomed to guard against the fly (called locally
“ Ganda”), when driving their cattle through the forest,
by doing so at night, and by lighting large tires, the smoke
of which acts as a protection to the cattle” (p. 355, et seq.).

TRANSPORT ANIMALS ON UGANDA RAILWAY. 241

[A fairly good enlarged figure of Glossina morsitans
(more probably Gl. pallidipes) is given on p. 356, and a
figure of the antenna (called by mistake the proboscis)
on p. 357.]

“ Ganda” -fly reported to exist from Wange (near Lamu)
up to Itembe, in consequence of which neither cattle nor
donkeys thrive (p. 419).

People of Simambaya Sendeni (a large pipe about
three miles beyond Vundene) possess “neither cattle nor
donkeys, owing to the prevalence of the Ganda-fly” (p.
432).

“The Ganda-fly is prevalent here also [in the neighbour-
hood of Rubu, a large village on the sea-coast, with the
island of Simambaya lying opposite to it], consequently
the people possess no cattle, but there appeared to be
numbers of sheep and goats” (pp. 434-435).

[Wange is at the west end of the Mongoni Channel,
which runs inland from Manda Bay, to the north of
Manda and Lamu Is. Jtembe is close to the coast, north-
east of Patta I. Vundene and Rubu are close to the
coast, five to eight miles east of Itembe. |

155. 1898.

‘* REPORT ON THE PRoGRESS OF THE MomBasa- VICTORIA
(UcanpA) Ratbway, 1897-98.” Presented to both Houses
of Parliament by command of Her Majesty, July, 1898.
London: Printed for Her Majesty’s Stationery Office by
Harrison & Sons, St. Martin’s Lane, Printers in Ordinary
to Her Majesty.

‘*The country in parts being infested with Tsetse-fly
as far as mile 220 [a little beyond Kiboko], the mortality
amongst transport animals would have been very heavy
had larger transport arrangements been made for expedi- -
ting works far ahead of the rails, and an enormous outlay
would have been entailed which circumstances hardly
seemed to justify ” (p. 3).

“ Transport.—It has been found necessary to keep up
a transport department for the purpose of supplying the
advanced earthwork and survey parties with food and
water. Camels, donkeys, bullocks, and mules have been
tried, but mules alone have been found effective, owing to

' the ravages caused: by the Tsetse-fly amongst the other
R

242 CONDITIONS ON UGANDA RAILWAY.

‘classes of animals. It has been found that by taking
special precautions and making applications of sheep-dip
the mortality was to a certain extent reduced, and in the
cases of mules it was only 10 per cent. during the last six
months of the year” (p. 3).

With reference to the above, and especially the state-
ment in the first-quoted paragraph as to ‘‘the country in
parts being infested with Tsetse-fly as far as mile 220,” the
following extract from p. 2 of the Report will be of interest
as describing the country referred to :—

“Of the entire length (263 miles) of line surveyed, it
may be said that, at least, 200 are through difficult
country. Except for small rivers met with at long inter-
vals, the country is waterless and covered with dense
impenetrable thorny scrub, while the surface’ is much
broken and intersected with ravines.

“This belt of country, extending inland 200 miles, is,
therefore, one through which survey operations could only
be carried out under circumstances of considerable diffi-
culty and hardship. Beyond the belt the country is
more open.

“ Mr. Hodson, attached to the Indian Public Works
Department, who made an inspection of the railway as

far as mile 100, states in his report as follows: ‘The
country it traverses is almost continuous thorny jungle
with but few inhabitants and no cultivation, much of it
practically waterless; the jungle is thorny scrub, high
enough and thick enough to make surveying, and even
exploring, off the regular track of the caravans, a very
arduous undertaking, but devoid of all trees with foliage
sufficient to give any appreciable shade.’ ”

156. 1898. G. de N. Hough.
Proceedings of the Academy of Natural Sciences of
Philadelphia, 1898, p. 172.
Two females of Glossina longipennis, Corti, collected in
Somali Land by Dr. A. Donaldson Smith, August 19 and
24, 1894.

157. 1898.

“Tue Secret oF THE Tsetse” (The Globe, July 11,

1898).
“ A mysterious illness which has played havoc with

BELIEF OF NATIVES AT LOKOJA. 243

the mounts and transport animals of the Royal Niger
Constabulary at the Lokoja headquarters has now been
identified as the baneful work of the Tsetse-fly.”

“+++ the Lokoja natives, knowing nothing of microbes
and bacteria, assert that the fly extracts from a certain
small red monkey the virus with which it inoculates the
bush cow or dwarf buffalo.”

158. 1899. D. Sharp.

The Cambridge Natural History. ‘‘Insxcts,” Part IT.
(London: Macmillan & Co., Limited), pp. 512-513.

Notes on Glossina morsitans, with woodcuts of perfect
insect, larva, and pupa (the two latter after Bruce).

“‘ Although it has been supposed that the Tse-tse-fly
is a formidable obstacle to the occupation of Africa by
civilised men, there is reason to suppose that this will not
ultimately prove to be the case. It only produces disease
when this pre-exists in animals in the neighbourhood ;
only certain species are liable to it; and there is some
evidence to the effect that even these may in the course
of a succession of generations become capable of resisting
the disease inoculated by the fly. As long ago as 1878,
Dr. Drysdale [vide supra, 58] suggested that this fly only
produces disease by inoculating a blood-parasite, and all
the evidence that has since been received tends to show
that his idea is correct” (p. 513),

159. 1899. F. V. Kirby.
“Sport in East Cenrrat Arrica, Being an Account
of Hunting Trips in Portuguese and other Districts of
Kast Central Africa” (London: Rowland Ward, Ltd.),
p. bits
On the right bank of the Inkambedsi River, near Chinde
(1894).—*The heat was stifling and oppressive in the
dense acacia thickets, where all the trees wore the apparel
of approaching summer ; and also in the bushy hollows
between the rough and stony ridges, where, to add to our
discomforts, Tsetse-flies swarmed in such numbers that
we had to use small branches to keep them off” (p. 17).

160. 1899. Sir H. H. Johnston, K.C.B.
“A History oF THE COLONIZATION OF AFRICA RY
Auien Races” (Cambridge: The University Press).
RB 2

244 TSETSE AND MINING IN RHODESIA.

The Arabs at Sena accused of having poisoned the
horses and camels of Captain-General Francisco Barreto’s
expedition from Quelimane to Sena in 1569. “ What
really took place, however, seems to have been that the
horses and camels were exposed to the bite of the Tsetse-
fly, and died in consequence of the ato of this
venomous insect” (p. 48).

161. 1899. William Harvey Brown.

‘On THE SoutH ArricaNn Frontier. The Adventures
and Observations of an American in Mashonaland and
Matabeleland ” (London: Sampson Low, Marston & Co.),
pp. 151, 314-315.

Tsetse-fly numerous in the Concession Hill district, west
of Hartley Hill, Mashonaland, Jan. 1, 1891.

“The sky was overcast with heavy clouds ; the atmos-
phere was damp, and the day warm. ... Much of the
country through which we travelled was covered with
enormous ant-heaps and thick jungle.

“In some places the Tsetse-flies were exceedingly
troublesome, even biting through our clothing” (p. 151).

“Tn addition to pleuro-pneumonia among cattle, and
horse-sickness among horses, there is in Rhodesia that
worst of all scourges to stock, the Tsetse-fly. This is a
great drawback to the importation of machinery to many
of the mining districts, which lie in the fly-infested
sections. Traction-engines and railways will be necessary
to overcome the obstacles created by this insect pest.
Fortunately, the Tsetse-fly does not occur on the plateau.
It is met with mainly in the low-lying country bordering
the Zambesi River. This insect is somewhat similar in
appearance to a house-fly, but about three times as large.
Its bite is fatal to all domestic animals, but it does not
seem to affect wild game or man. When badly bitten,
horses and cattle usually succumb in two or three weeks.
Donkeys last longer. Few animals, however, ever recover
from its bite. Until recently the supposition has been
entertained that a direct poison is introduced into the
system by the bite of the fly. Surgeon-Major David
Bruce, of the British Army, after a long series of
experiments in Zululand, has advanced the theory that
‘the Tsetse acts as a carrier of a living virus, an infinitely

TSETSE IN SOMALILAND. 245

small parasite, from one animal to another, which enters
the blood-stream of the animal bitten or pricked, there
propagates, and thus gives rise to the disease.’ This theory
is more in harmony with modern science” (pp. 314-315).

162. 1899. E. E. Austen.

“Report OF THE PROCEEDINGS OF THE EXPEDITION
FoR THE Stupy oF THE Causes or Manaria, despatched
to Sierra Leone, West Africa, under the leadership of
Major Ronald Ross (late Indian Medical Service), by
the Liverpool School of Tropical Diseases, July 29th,
1899.” Printed by Order of the Trustees of the British
Museum. (London: Printed for Her Majesty’s Stationery
Office by Darling & Son, Ltd., 1-3, Great St. Thomas
Apostle, E.C.). Pp. 18-19.

Notes on “‘ Glossina longipalpis, Wied.” (really Gl. pal-
palis, Rob.-Desv.), in the vicinity of Free Town, Sierra
Leone.

168. 1900. C. V. A. Peel.

“ SomaLinanD” (London: F. E. Robinson & Co.),
pp. 116-117 :—see also map at end of volume.

Tsetse-fly to the north of Mount Kuldush, between Bun
Feroli and Biermuddo, east of the Daghato River, a
tributary of the Webbi Shebeyli.

“ At night, when the camels came in, they were
followed by a perfect swarm of Tsetse-fly. The sting of
this fly, though harmless to human beings, is very painful,
and made me jump every time I was bitten, as if a needle
had been stuck half an inch into my flesh. One of my
ponies was badly bitten on the ‘billy,’ his tail not being
long enough to whisk the pests off. The Somalis said the
pony would live until the next rain. We rubbed sheep’s
fat on the ponies and the camels every day.

“Tt is an extraordinary sensation, coming into a belt
of ‘fly.’ There may be but a tiny river-bed. On one side
of it not a fly will be encountered, but walk a dozen
feet and they suddenly come buzzing by one in hundreds.’
[The year referred to is 1895.]

164. 1900. E. E. Austen,
Proceedings of the Zoological Society of London for the
Year 1900;.p. 10... a

246 COACHING ENTERPRISE RUINED BY TSETSE.

165.

166.

A female of Glossina longipennis, Corti, taken in West
Somali Land, between June 23 and 25, 1895, by Mr.
GC. V. A. Peel.

“Mr. Peel’s note on this specimen says :—‘ Fly-belt
sharply defined from Biermuddo to Boholo Deno’’’
[vide supra, 163].

1900. H, A. Bryden.

“ ANIMALS OF ArFrica” (London: Sands & Co.), pp.
228-230.

A line of coaches, established some years ago to run from
the Pungwe River towards Mashonaland, abandoned owing
to the Tsetse-fly—It is a rather curious fact that
wherever the African buffalo is plentiful, there you will
almost certainly find the Tse-tse-fly. As the buffaloes are
killed out or driven away the Tse-tse disappears, and
horses and oxen can be introduced. It is supposed by
many that the Tse-tse is bred in the dung of the wild
buffalo, which would, of course, account for its presence in
districts where buffaloes are abundant. This is a very
troublesome and a very costly insect for African explorers,
hunters, and settlers. Many thousands of pounds must
have been lost by its ravages, and vast districts remain
unopened owing to the presence of this dangerous pest.
Some few years since, when Rhodesia was first being
opened up, an attempt was made to run a line of coaches
from the Pungwe River, on the East Coast, towards
Mashonaland. Splendid American coaches were imported
and plenty of fine horses and mules got out. But the
Tse-tse-fly beat this spirited enterprise. The horses and
mules died, the attempt had to be abandoned, and the
coaches still lie rotting in the wilderness. In its turn,
however, I am glad to say, the deadly Tse-tse has been
overcome by the railway ” (pp. 229-230).

Igor. Sir H. H. Johnston, K.C.B.

“Report By His Magzsry’s Specta, ComMIssIoNER
ON THE ProrecToRATE oF Ucanpa,.” With Map. Pre-
sented to both Houses of Parliament by command of His
Majesty. July, 1901. London: Printed for His Majesty’s
Stationery Office by Harrison & Sons, St. Martin’s Lane,
Printers in Ordinary to Her late Majesty.

“The journey from Mombasa on the east coast to the

DONKEY DISEASE IN EAST AFRICA. 247

administrative centre of Uganda occupied [in 1894] under
favourable circumstances from three to four months, and
had to be performed mainly on foot owing to the difficulty
of conveying riding animals through the belt of country
near the coast infested with the Tsetse-fly ” (p. 3).

167, 1901. Dr. Max Schoeller.

“ MITTEILUNGEN UBER MEINE REISE NACH AQUATORIAL-
Ost-AFRIKA UND Uaanpa, 1896-1897,” Band I. (Verlag von
Dietrich Reimer (Ernst Vohsen), Berlin, 1901), pp. 118-120.

In spite of careful search no Tsetse-fly was found: the
deaths of the author’s baggage-donkeys were apparently
due to a disease allied to the horse-sickness of South
Africa (“‘ Dikkopziekte” of the Boers).

The author, writing of an attempt, which was being
made by a Berlin society at the period to which this work
refers, to catch zebras, and cross them with horses and
donkeys, at Mbuguni, between Mt. Meru and, Kilima
Njaro, says :—

[Translation.] ‘‘ Zebra teams would certainly have
had a great advantage over all other material which comes
into consideration for East Africa, in the first place
because the Tsetse-fly would probably here be powerless,
while horse and donkey succumb to it. Whether, how-
ever, the horse-sickness, that causes such great devastation
in South Africa, will keep aloof from the zebra, or zebra-
hybrids as the case may be, remains an open question,
though I believe indeed that it will do so. According to
the observations that we had the opportunity of making
in connection with the successive deaths of our donkeys,
we might almost believe that the disease called Dikkop-
ziekte (thick-head sickness) by the Boers likewise occurs
in the steppe regions of East Africa. The donkeys that
we lost in the Natron Valley, and also later, all exhibited
those symptoms which are mentioned as characteristic
of thick-head sickness. The animals died either in the
course of a-few hours, or else, as was usually the case, in
that of several days. Our Wadschagga believed the sick-
ness to be the consequence of the bite of an insect (die
Folge eines Insektenstiches); they even designated the
producer by the name: Wandorobo. We have searched in
vain for such an insect in the places where the disease

248

WANDOROBO-FLY OF WADSCHAGGA.

occurred. In vain have we also looked for the puncture
of the insect on the genitals of the donkeys, in which
region according to the declarations of the natives it
ought to be found ; all that we are able to see, and this
invariably, was a sometimes slower, sometimes quicker
swelling-up of the mucous membranes, of the upper orbits,
lips, nostrils, and jaws accompanied by greater swelling
of the genitalia. The animals finally had a very high
body temperature, with quickened breathing, and died of
suffocation owing to difficulties in respiration. At that
time we also were of the opinion that we had only lost our
donkeys in the territories of the Tsetse-fly, and that the
latter had occasioned the phenomena of poisoning. To-day,
on the contrary, we would incline to the view, that we are
confronted with a phenomenon analogous to what is found
in the case of horse-sickness in South Africa, and that the
sickness represents a special form of infection with the
greater probability, since the South African disease already
appears to be recognised as such. Definite grounds for
believing in insects (bestimmte Anhaltspunkte fiir Insekten)
have not been discovered, and the imported Muscat
donkeys were attacked soonest and in the largest per-
centage. According to the statement of the natives,
moreover, similar symptoms of disease sometimes appear
in human beings. At the coast it had already been pro-
phesied that we should lose our donkeys on the steppe,
and we therefore searched for the Tsetse-fly so zealously,
that it would scarcely have escaped us if we had really
entered its territory. That the Wadschagga designate
a small fly by the name Wandorobo appears to me to be
probable, but the question is whether we are here dealing
with the Tsetse-fly. The districts in which we lost our
baggage-donkeys cannot be called free from fever ; the
Natron Valley, where the sickness first appeared, is in
many places swampy, and the neighbourhood of Kibwesi,
which is characterised as especially unfavourable for the
escape of the animals, would doubtless exhibit malaria.
As Professor Koch has shown, punctures by insects have
an importance in connection with malarial infection in so
far as blood-sucking insects carry contagious matter into
the blood. It may well be doubted whether it is the
Wandorobo-fiy alone that is able to transport the malaria

RESULTS FROM UGANDA RAILWAY. 249

poison to certain animals, or to inoculate the horse-sickness
that is caused by plasmodia—rather may the most widely
different biting-flies, midges, etc., provide for an occa-
sional transference of fever bacteria. Just as the latter
under definite conditions of life develop into definite
forms, so also will the poison occasion special sickness
symptoms in the case of the different animals. In South
Africa it is supposed that the donkey and the Quagga
[sic] escape the horse-sickness, but that observation in no
way guarantees that the same is also the case in East
Africa. It is also not impossible in the case of other
forms of bacteria, that at certain times, in certain districts,
and under all kinds of apparently unimportant collateral
circumstances, they fall into a condition of harmless
sterility, to attack their victim later on or elsewhere under
more favourable conditions with devastating vehemence.”

168. 1901. Sir C. N. E. Eliot, K.C.M.G.

«Report py His Masesry’s ComMMIssionER ON THE
East Arrica Prorscrorate (Africa, No. 9 (1901)).” Pre-
sented to both Houses of Parliament by command of His
Majesty. August, 1901. London: Printed for His
Majesty’s Stationery Office, by Harrison & Sons, St.
Martin’s Lane, Printers in Ordinary to His Majesty.

“ Horses and mules are now used for travelling pur-
poses in Ukamba, and it is no longer the custom to make
long marches on foot. This is one of the many advan-
tageous changes wrought by the [Uganda] railway.
Horses do not thrive in the coast districts, and it was
formerly impossible to take them up-country through
regions where they were exposed to Tsetse-fly and various
sicknesses. Now they can be transported rapidly through
the dangerous zone to the interior” (p. 20).

“T have hoped that in course of time some practical
steps may be taken towards domesticating the zebra and
the wildebeeste. These animals appear to be immune to
the deadly Tsetse-fly which abounds in some districts of
the Protectorate, and, if either of them or mules bred from
them could be employed as pack or draught animals, it
would in all probability have an incalculably good effect
upon the country, the prosperity of which is at present
sadly hampered by want of efficient transport.” (p. 25).

250

169.

170.

TSETSE IN TOGO.

igor. S. L. Hinde and H. Hinde.

“Tus Last or THE Masar” (London: William
Heinemann), p. 167.

Field Notes on the Game of East Africa (i.e. The
British East Africa Protectorate).—‘‘ Owing. to the
difficulty of getting horses through the Tsetse-fly belt,
which extends over 200 miles from the sea-coast, all
hunting and shooting has until now been done on foot.
Since the advent of the railway it is feared that
luxurious ‘ sportsmen’ will bring their horses with them,
and another factor will be brought to bear upon the final
extinction of big game in the last stronghold left in Africa.”

1901. Dr. Schilling.

“ BERICHT UBER DIE SURRA-KRANKHEIT DER PFERDE”
(Centralblatt fiir Bakteriologie, Parasitenkunde und In-
Jektions - Krankheiten. Erste Abteilung: Medicinisch-
hygienische Bakteriologie und tierische Parasitenkunde.
xxx. Band. No. 15. Jena, 30 Oct. 1901, pp. 545-551).

The author, who writes from Kleinpopo, Togo, gives
an account of a disease which is fatal to horses in Togo,
Slave Coast, W. Africa. He calls the disease Surra, and
describes the Trypanosome which causes it.

Tsetse-fly in the Togo Protectorate-—({Translation. |
“The Tsetse-fly is accused of being the carrier of Surra.
So far as I am aware, this fly does not occur in the Togo
Protectorate littoral, which lies between sea and lagoon.
On the other shore of the lagoon, consequently at a
distance of some three kilometres from the sea-coast, it is
very common. The natives know the fly by the name
*‘adjoé.” Experiments intended to elucidate the réle of
the Tsetse-fly in infecting animals with the disease are
only in the initial stage, since hitherto a sufficient supply
of flies has been wanting. At any rate, the fly bites very
readily, and sucks up large quantities of blood with its
extremely fine, tubular proboscis. In the case of dogs
and horses no perceptible swelling appears after the bite,
and therefore it seems to be questionable whether a
“saliva,” as in the mosquito, is produced at.all. The
dissection of the insects is very difficult, owing to the
great development of the musculature and the firmness of
the tracheal network.” (p. 551). -.

LOCALITIES OF TSETSE DISEASE. 251

171. 1902. A. Laveran et F. Mesnil.

‘‘ RECHERCHES MORPHOLOGIQUES ET EXPERIMENTALES
sUR LE TRYPANOSOME DU NaGaNA ou MALADIE DE LA
Movucue Tstrst (Annales de L’Institut Pasteur, Jan. 25,
1902, pp. 1-55).

Three figures of Glossina (probably Gl. pallidipes,
Austen) on p. 8 (after Bruce).

According to the authors, the Tsetse-fly occurs in the
vicinity of Lake Tchad: in an enumeration on p. 2 of the
localities in which Tsetse-fly disease is known to exist,
they state that it is particularly prevalent in the region
of Lake Tchad, on the banks of the Chari and_ its
affluents ; adding in a foot-note that the statement is
based upon information supplied by Inspector-General
Kermorgant, who received it from Surgeon-Major Morel,
of the French Colonial Army. Specimens of the Tsetse-
fly were enclosed in General Kermorgant’s letter.

It is stated on the authority of Blanchard (Bulletin de
LV Académie de médecine, 3° série, xlvi, 29 Oct. 1901), who
quotes a writer named Brumpt, that at Imi, on the Webi
Shebeli, Somaliland, all the camels of a mission, as well
as the mules and donkeys, succumbed to a Trypanosoma.
The epizooty observed by Brumpt did not appear to be
propagated by the ordinary Tsetse-fly, but by a closely
allied species of Glossina [doubtless Gl. longipennis, Corti].

During the Abyssinian campaign of 1867 there was
great mortality among the transport animals, and a
veterinary surgeon named MHallen,* proceeding from
Abyssinia to India, was struck by the resemblance
between Surra and the Abyssinian disease.

172. 1902. Sir Harry Johnston, G.C.M.G., K.C.B.

“Tux Ucanpa Prorecrorate” (London: Hutchin-
son & Co.), Vol. I.

‘‘Hither there is no true Tsetse-fly in the Uganda
Protectorate or it is not able to obtain and introduce
into the bodies of domestic animals the malarial germs
which cause Tsetse fever. Therefore, theoretically there
is no part of the Uganda Protectorate in which cattle,
sheep, goats, and horses cannot be kept” (pp. 288-289).

* The late Lieut.-Colonel J. H. B. Hallen, A.V.D., who accompanied

Lord Napier’s expeditionary force to Magdala, and was afterwards for some
time Chief of the Veterinary School at Bombay. »

bo
IU

2 TSETSE IN UGANDA PROTECTORATE.

“Flies of the genus Glossina (a genus of which the
Tse-tse is a member) exist in the Uganda Protectorate.
They have been caught there by Mr. Jackson and myself,
not to mention many other collectors. But either the
true T'se-tse is absent from all parts of the Protectorate
or it is unable to obtain there the germs of fever which
it is the agent in introducing to the blood of horses,
cattle, and other beasts. This is a very fortunate cir-
cumstance, as it removes a serious hindrance in the way
of rearing live-stock and developing transport” (pp. 413-
414).

173. 1902. J. N. Justice.

‘PROSPECTING IN NorTHERN Ruopesia” (Travel,
Vol. VII., July, 1902. London: Horace Marshall &
Son, p. 102).

Tsetse-fly near the Kafue River, 1900.—-Natives state
that the virulence of the bite of the fly has greatly diminished
since the extermination of the buffalo by rinderpest.

“Tt was not far from here [near the Kafue River,
about ninety miles above its confluence with the Zambesi|
that we sustained our first losses from the Tsetse-fly ;
sixteen donkeys and two horses fell victims to this scourge
of the country in the nine months of the expedition.
Livingstone thought that donkeys were immune from the
evil effects of the fly’s bite, but this has long since been
disproved ; the natives, however, state that since the
rinderpest exterminated the buffalo the tsetse has lost
much of its venom, and it does indeed seem to be less
virulent than formerly, as a horse lives now for three
weeks after a bite, whereas in former times it died a few
days afterwards.”

174. 1902. Dr. Franz Stuhlmann.

“ VORKOMMEN VON GLOSSINA TABANIFORMIS (WESTW.)
BEI Dar-Es-SatAm” (Berichte tiber Land- und Forstwirt-
schaft in Deutsch-Ostafrika. Herausgegeben vom Kaiser-
lichen Gouvernement von Deutsch-Ostafrika, Dar-es-Salam.
1 Bd. Heft 2, pp. 173-175, with figure of the antenna,
magnified thirty-eight times, in text. Heidelberg, June,
1902: Carl Winter’s Universitiitsbuchhandlung).

A few specimens of a biting fly, which, on examina-
tion, proved to be ‘ Glossina . tabaniformis, Westw.,” (Gl.

TSETSE NEAR DAR-ES-SALAAM. 253

fusca, Walk.), “ were caught in January, 1902, at Manga-
mara, a little way to the north of Dar-es Salam, beyond
Msimbasi Creek,”

The following is a translation of the concluding
paragraph of this paper (p. 175).

‘‘ Now the most interesting question is whether this
species of Glossina also conveys Surra-disease. Reasoning
from general conclusions derived from analogy in con-
nection with the diseases due to infection of the blood,
in which each parasite has its own particular host and
intermediate host, I should be inclined a priori to answer
this question in the negative. This biting fly is evidently
not rare near Dar-es-Salim, and no case of Surrahas yet
been observed originating from there. At the place
where the flies were caught the same cattle and goats
have grazed for a long time, without disease appearing
among them. The examination of the blood of the goats
has so far not revealed any Trypanosomes, but this
examination must be continued and extended to cattle
and donkeys. I think, however, it may be assumed that
Surra-disease is not conveyed by Glossina tabaniformis.”

BIBLIOGRAPHY.—PART II.

TSETSE-FLY DISEASE AND THE TRYPANOSOMA OF
NAGANA AND KINDRED SPECIES.
I. 1897. Surgeon-Major David Bruce, A.M.S. (now Lieut.-Col.
David Bruce, R.A.M.C.).
“FurTHER Report ON THE TsETSE-FLY DISEASE OR
Naaana IN ZuLuLanp.” Ubombo, Zululand, 29th May,
1896 (London: Harrison & Sons, St. Martin’s Lane,
Printers in Ordinary to Her Majesty, 4th February,
1897), 69 pp., Plates I-VI.
[See Chapter VII., Appendix A.]

II, 1898 G. H. F. Nuttall.
‘“ NEUERE UNTERSUCHUNGEN UBER Mataria, TExas-
FIEBER UND ‘TSETSEFLIEGENKRANKHEIT” (Hygienische
Rundschau, Berlin, 15th Nov., 1898, VIII, pp. 1084-1103).

254 TRYPANOSOME IN BLOOD OF MAN.

III. 1898. R. Koch,

‘ REISEBERICHTE UBER RINDERPEST, BUBONENPEST IN
INDIEN UND AFRIKA, TSETSE- ODER SURRAKRANKHEIT,
Texas Fiseser, TROPISCHE MaLaria, SCHWARZWASSER-
FIEBER ” (Berlin: J. Springer : 136 pp.). Pp. 65-72, 87-88.

According to Laveran and Mesnil, [171] p. 2, notes 1
and 2, the author records the occurrence of Nagana in
German East Africa, and states (p. 66) that in 1895 he
observed Trypanosomes in blood-films relating to Tsetse-
fly disease, which had been forwarded to him from Togo
(Slave Coast).

IV. 1898. Dr. G. Nepveu.

“Sur uN TRYPANOSOME DANS LE SANG DE L’HOMME”
(Comptes Rendus Hebdomadaires des Séances et Mémoires
de la Société de Biologie. Tome cinquiéme—Dixiéme Série.
Séance du 24 Décembre 1898, pp. 1172-1174).

The parasite was found in 1890 fairly common in the
blood of a patient in Algeria, together with Laverania.
It is not described in detail or named, but exhibited all
the characters of Trypanosoma. The organism was first
mentioned by the author in his paper entitled “ Ftude
sur les Parasites du Sang chez les Paludiques” (Compt.
Rend. Hebd. des Séances et Mém. Soc. Biol. Tome
Troisisme—Neuviéme Série. Année 1891, pp. 39-50).

V. 1898. A. A. Kanthack, H. E. Durham, and W. F. H. Blandford.
“On Nacana, or Tserse-Fiy Diskasez.” Report,
made to the Tsetse-fly Committee of the Royal Society,
of Observations and Experiments carried out from
November, 1896, to August, 1898 (Proceedings of the
Royal Society of London, Vol. LXIV., Nov. 19, 1898,

pp. 100-118).

VI. 1899. G. H. F. Nuttall, M.D.
“On THE Rote or Insects, ARACHNIDS AND
MyYRIAPODS, AS CARRIERS IN THE SPREAD OF BACTERIAL
AND Parasitic DisgeAsES OF Man and ANIMALS. A
CriticaL AND Hisrorican Srupy (Johns Hopkins
Hospital Reports, Vol. VIII., pp. 1-155, Plates I.-III.).
Tsetse-fly Disease, pp. 68-71.—Résumé, chiefly of
* Bruce’s results.
Bibliography, pp. 139-140.

REPRODUCTION IN TRYPANOSOMES. 2!

Or
or

VII. 1899. H. G. Plimmer and J. Rose Bradford.
“A PretiminAry Note on THE MoRPHOLOGY AND
DisTRIBUTION OF THE ORGANISM FOUND IN THE TSETSE-
FLY Disrasn” (Proceedings of the Royal Society of London,

Vol. LXV., Aug. 31, 1899, pp. 274-281).

VIII. 1900, A. Laveran et F. Mesnil.

“Dr tA Loncurt CONSERVATION A LA GLACIERE DES
TRYPANOSOMES DU Rav ET DE L’ AGGLOMERATION DE CES
Parasites” (Comptes Rendus Hebdomadaires des Séances
et Mémoires de la Société de Biologie. Séance du
6 Octobre 1900, pp. 816-819).

IX. 1900. A. Laveran et F. Mesnil.
‘Sur L’ AGGLUTINATION DES TRYPANOSOMES DU Rat
PAR DIVERS Strums” (Comptes Rendus Hebdomadaires des
Séances et Mémoires de la Société de Biologie. Séance du

10 Novembre, pp. 939-942).

X. r900. A. Laveran et F. Mesnil.
“‘SuR LE MODE DE MULTIPLICATION DU TRYPANOSOME
pu Rat” (Comptes Rendus Hebdomadaires des Séances
et Mémoires de la Société de Biologie. Séance du 17
Novembre, 1900, pp. 976-980. With figures in text.)

XI. xrg0r. A. Laveran et F. Mesnil.

“SuR LE MODE DE MurtipiicaTion DU TRYPANOSOME
pu Nacana” (Compies Rendus Hebdomadaires des Séances
de la Société de Biologie. Tome LIII. Séance du
23 Mars 1901, pp. 326-329).

[With five figures showing the multiplication of the
Trypanosome by longitudinal division. |

XII. 1901. A. Laveran et F. Mesnil.

“Sur ta Nature CENTROSOMIQUE DU CoRPUSCULE
CHROMATIQUE PosTERIEUR DES TRYPANOSOMES ” (Comptes
Rendus Hebdomadaires des Séances de la Société de Biologie.
Tome LITI. Séance du 23 Mars 1901, pp. 329-331).

XIII. rg901. E. Nocard.
“Sur Les RAPPORTS QUI EXISTENT ENTRE LA Dourine
ET LE Surra ou LE Nagana” (Comptes Rendus Hebdomadaires
des Séances de la Société de Biologie. Tome LIII, Séance
du 4 Mai 1901, pp. 464-466).

256

ATV,

XIVa.

XV.

XVAa.

XVI.

MORPHOLOGY OF TRYPANOSOMES.

1901. A. Laveran et F. Mesnil.

“SurR LA STRUCTURE DU TRYPANOSOME DES GRE-
NOUILLES ET SUR L’EXTENSION DU GENRE Trypanosoma,
Grusy” (Comptes Rendus Hebdomadaires des Séances de la
Société de Biologie. Tome LIII. Séance du 22 Juin
1901, pp. 678-680 (with figures in text) ).

The following is a translation of the final paragraph
of this paper :—‘‘In conclusion, our observations show
that the Trypanosome of the green frog exhibits all the
peculiarities regarded by Wasielewsky and Senn as
characteristic of the genus Herpetomonas. Under these
conditions this genus must disappear, and the generic
name ZJ'rypanosoma alone will serve to designate all the
flagellated parasites in the blood of Vertebrates, at least
all those at present known.”

1901. A, Laveran et F. Mesnil.

“Sur LA MoRPHOLOGIE ET LA SYSTEMATIQUE DES
FLAGELLES A MEMBRANE ONDULANTE (GENRES Trypanosoma
Gruby et Trichomonas Donné)” (Comptes Rendus Hebdom-
adaires des Séances de V Académie des Sciences, Tome
CXXXITI., No. 3 (15 Juillet 1901), pp. 131-137, with
five figures in text).

1901. A. Laveran et F. Mesnil.

“Sur tA Nature BactTkRIgENNE DU PRETENDU
TRYPANOSOME DES Huvirres (Tryp. Balbiani, Certes) ”
(Comptes Rendus Hebdomadaires des Séances de la Société
de Biologie. Tome LIII. Séance du 19 Octobre, pp.
883-885).

rigor. A. Laveran et F. Mesnil.

‘Sur LES FLAGELLES A MEMBRANE ONDULANTE DES
Poissons (GENRES. Trypanosoma Gruby et Trypanoplasma
n. gen.) ” (Comptes Rendus Hebdomadaires des Séances de
LAcadémie des Sciences, Tome CXXXIII., No. 18
(28 Octobre 1901), pp. 670-675, with four figures in
text).

1go1. A. Laveran et F. Mesnil.

“Sur Le TRYPANOSOME DES Rats” (T. lewisi, Kent)
(Annales de l'Institut Pasteur, September 25, 1901, pp.
679-684). :

XVII.

XVIII.

XIX.

EXPERIMENTS WITH HORSE-FLIES. 257

1901. Dr. Schilling.

“ BERICHT UBER DIE SuRRA-KRANKHEIT DER PFERDE”
(Centralblatt fiir Bakteriologie, Parasitenkunde und In-
Sektionskrankheiten. Erste Abteilung: Medicinish-hygie-
nische Bakteriologie und tierische Parasitenkunde. XXX.
Band. No. 15. Jena, 30 Oct. 1901, pp. 545-551).

The author, who writes from Kleinpopo, Togo, gives
an account of a disease which is fatal to horses in the
Togo Protectorate, Slave Coast, W. Africa. He calls the
disease Surra, and describes the Trypanosome which
causes it.

igor. A. Theiler.
“Diz Tserse-Krankueit” (Schweizer Archiv fiir
Thierheilkunde, XLII. Band, p. 97).

1901. L. Rogers, M.D.

“Tur TRANSMISSION OF THE Trypanosoma Evansi BY
Horsrz Fuies, AND OTHER EXPERIMENTS POINTING TO
THE PropaBLe Ipentity oF Surra OF INDIA AND
Nacana or TseErse-FLY Disease or Arrica (Pro-
ceedings of the Royal Society of London, Vol. LX VIII,
pp. 163-170).

“ The close resemblance between Surra of India and
Tsetse-fly disease of Africa has long been known, while
Koch, after having seen the living Trypanosoma Hvansi at
Muktesar in India, and soon after studied the parallel
disease in German East Africa, pronounces them to be
the same, and in his ‘ Reiseberichte’ calls the disease
seen in the latter place ‘ Suryakrankheit’ (p. 163).

“In every case in which the flies had been kept from
one to four or more days after biting the infected animals,
no disease ensued in the healthy ones. Many such flies
were dissected and microscopically examined, but in no case
was anything which might be taken for a development of
the Trypanosoma in the tissues of the insect detected... .
When, however, flies which had just sucked infected
blood were immediately allowed to bite another healthy
animal, positive results were obtained after an incubation
period corresponding with that of the disease produced
when a minimal dose of infected blood is inoculated into
an animal of the same species. The result was uncertain
if only one or two flies were allowed to bite, and especially

3

258

XX.

XXII.

TRYPANOSOMA BRUCII AND NEW. SPECIES.

if they were allowed to suck as much blood as they
wished without being disturbed. If, on the other hand,
several flies, which had just sucked an infected animal,
were induced to bite a healthy one, and especially if they
were disturbed and allowed to bite again several times,
infection was always readily produced in both rabbits and
dogs, the fur of the latter having been carefully cut,
without abrading the skin, at the site over which the
flies were applied ” (p. 164).

“In every point, then, that I have so far investigated,
the results obtained in the case of Surra closely agree
with those of the Royal Society’s Committee in Tsetse-
fly disease, and so far as they go they support the view
that the two diseases are probably identical ” (p. 169).

1902, J. R. Bradford and H. G. Plimmer.

‘THE TRYPANOSOMA BRUCII, THE ORGANISM FOUND IN
Nagana, or Tst-Tsr-rty Disease” (Quarterly Journal of
Microscopical Science, Vol. 45. New Series, pp. 449-471, —
Plates 24 and 25).

1902, A. Laveran et F. Mesnil.
‘RECHERCHES MoORPHOLOGIQUES ET Runisrandtiites

- SUR LE TRYPANOSOME DU NaGaNna OU MALADIE DE LA

XXII.

XIII.

XXIV.

Movcue Tsitsé” (Annales de Institut Pasteur, Jan. 25,
1902, pp. 1-55).
1902, A. Laveran.

“SuR UN NOUVEAU TRYPANOSOME DES Bovipts”
(Comptes Rendus Hebdomadaires des Séances de ? Académie
des Sciences, Tome CXXXIV, No. 9 (3 Mars 1902),
pp. 512-514).

1902, A. Laveran.

“Dr action Du Strum HumaAIN suR LE TRYPANOSOME
pu Nacana (Jr. Brucei)” (Comptes Rendus Hebdoma-
daires des Séances de [Académie des Sciences, Tome

CXXXIV, No. 13 (1* Avril 1902), pp. 735-739).

1902. Lieut.-Col. D. Bruce, R.A.M.C.

* Nore ON THE Discovery on A New Trypanosoma”
(Proceedings of the Ro vibe Society of London, Vol. LXIX.,
p. 496).

“ This new species. can be at once distinguished from
the Trypanosomas of Surra, Tse-tse-fly disease, or Rat by

XXV.

XXVI.

NAGANA IN GERMAN EAST AFRICA. 259

its larger size, it being almost twice as large as any of the
others. In general appearance it conforms closely to the
others in possessing an oval protoplasmic body, a longitu-
dinal fin-like membrane, and a single flagellum.

“This new Trypanosoma was lately discovered by
Dr. A. Theiler, who is in charge of the Bacteriological
Laboratory of the Medical Officer of Health, Pretoria,
Transvaal.”

Dr. Theiler “found the parasite for the first time in
the blood of a young ox which had just recovered from
an attack of rinderpest. ...” ‘He found that the new
Trypanosoma only infects cattle. Horses, dogs, goats,
rabbits, and guinea-pigs are all immune, neither showing
symptoms nor the presence of the parasites in the blood.”

1902. J. E. Dutton, M.B.

“ PRELIMINARY NoTE UPON A TRYPANOSOME OccuR-
RING IN THE Bioop or Man” (Thompson Yates Laboratory
Reports, Liverpool, Vol. IV., Part II., pp. 455-468,
Plates V., VI., Charts 1-4).

1902. Dr. Franz Stuhlmann.

“ NOTIZEN UBER DIE TSETSEFLIEGE (GLOSSINA MORSITANS,
WESTW.) UND DIE DURCH SIE UBERTRAGENE SURRAHKRANK-
HEIT IN Deurscu-Ostarrika” (Berichte tiber Land- und
Forstwirtschaft in Deutsch-Ostafrika. Herausgegeben vom
Kaiserlichen Gouvernement von Deutsch-Ostafrika, Dar-
esSalim. 1 Bd. Heft 2, pp. 137-153, Tafel I and IT,
and four figures in the text. Heidelberg, June, 1902:
Carl Winter’s Universititsbuchhandlung). Tafel I gives
rough figures of Glossina morsitans, and details, and
also figures of several species of Trypanosoma. Tafel II
consists of a sketch-map of German East Africa, on which
places known to be, or suspected of being, centres of
‘‘Surra,” and roads on which bullock-teams have been
attacked by the disease, are marked in red.

The author commences by stating that a portion of
the mortality among cattle in German East Africa was
determined by Koch, in 1897, to be due to Surra. He
proceeds to discuss the true part played by the Tsetse-fly,
and rejects the idea that it is merely a passive carrier of the
heematozoon.

[Translation.] “The investigations upon Texas fever

8s 2

260

RELATION OF TSETSE TO PARASITE.

which is carried by ticks, and especially upon the different
forms of malaria lend the greatest probability to the
conclusion, that each of these blood-diseases can be carried
only by a perfectly definite kind of blood-sucking insect
(human malaria by Anopheles, bird malaria by Culea,
Texas fever by the tick, recurrent fever by bugs, and so
on), and that in this intermediate host the organism that
occasions the disease passes through a special stage of
development, which appears generally to be a sexual
reproduction.*

‘Now since in South Africa experiments have
furnished incontrovertible proof that the Trypanosoma-
disease is carried by the Tsetse-fly, the bite of which had
long been dreaded as being fatal ; since on the other hand
the same disease occurs here ; and finally seeing that the
true Tsetse-fly has been observed here, it is justifiable to
conclude that it is highly probable that in our colony
also Surra is carried, and, moreover, exclusively carried
by this one fly, the Tsetse. The fact is that, in the case
of all these blood diseases, the host and intermediate host
are animals of perfectly definite kinds. It is, moreover,
improbable that the proboscist of the Tsetse merely
operates as an inoculating-needle ; rather must we assume,
from analogy with malaria, that within the fly the Trypa-
nosoma passes through a special stage of development,
which it is true is at present unknown” (pp. 137-138).

Numbers of Tsetse-flies, which were identified as speci-
mens of Glossina morsitans, Westw., were found on Feb. 18
and 19, 1900, on the road from Kilwa to Barikiwa: the
weather at the time was thundery and rainy (p. 138).

Notes on the genus Glossina, and on Stomoxys calei-
trans, with woodcuts of the latter and its transformations
(after Howard) (pp. 138-141). At the bottom of p. 139
the development of the Tsetse is stated in error to be still
entirely unknown. Pages 141 to 146 are devoted to a
description of Glossina morsitans, based upon an examina-
tion of three specimens taken near Kilwa, as, mentioned

* It should be noted, however, that the host in which a parasite
becomes sexually mature is the definitive and not the intermediate one.
Thus in the case of estivo-autumnal fever Anopheles is the definitive and
man the intermediate host of Laverania precox.—H. E. A.

.t The word in the original is Stich, meaning literally ‘‘stab” o
“puncture.”—E. E. A.

NAGANA-FOCI IN GERMAN EAST AFRICA. 261

above. On p. 145, there is a comparative diagnosis of
Gl. morsitans, “ tachinoides,” and “ tabaniformis.”

Notes on the distribution of “Surra” and Tsetse-flies in
German East Africa.

[Translation.] “Since the labours of Professor Koch
innumerable cases of Surra have been observed here,
above all by Veterinary-Surgeon Schmidt; in addition to
this small herds of cattle have been driven, by way of
experiment, along certain roads, in order to discover by
what roads cattle can be transported without danger of
contracting the disease. The results are shown on the
appended map (Tafel II), on which the red spots indicate
the places where Surra infection has been positively
demonstrated or where it is strongly suspected to exist,
while the red streaks show the roads on which animals
were infected. It looks quite distressing to find the very
foot of the Usambara Mountains, in the neighbourhood
of the coffee plantations, so thickly beset with Surra-foci,
but we must consider that that was the place where many
cases came under observation, and that consequently it is
probable that in future many a centre of infection will
also be found in the remainder of the territory. On the
whole, river-lowlands with high reed-grass seem to be
exceptionally dangerous, just as in South Africa the
plains of the Limpopo and Zambesi are especially dreaded.

“ Particularly at the foot of East Usambara have
cases of infection frequently been shown to have occurred ;
many a head of cattle kept on the plantations, which had
formerly been driven through these districts, died from
Surra. The following localities are especially suspicious :
Bondei Land, Kwa Maromo, the neighbourhood of Lewa,
and also Mombo, Masinde, the eastern foot of the Pare
Mountains and the borders of Lake Jipe. The mountains
of West Usambara, the localities Korogwe and Massigi,
as well as Lake Manga are certainly free from the disease.
The Masai, who are well-known as the best herdsmen,
always ascribe the deaths of cattle to the fact of their
having eaten reed-grass ; but it is to be supposed that it
is precisely this grass that harbours the Tsetse-fly, which
occasions the infection. The fly dreaded by the Masai
under the name ‘ Ndorobbo’ is still unknown ; it appears,
however, from the description to be more probably a

262

DANGEROUS ROADS.

horse-fly. Tsetse-flies have also been observed quite close
to Mombasa (Koch, Reiseberichte, p. 70).

“In the Hinterland of Dar-es-Salém the low ground
bordering the Gerengere River is under suspicion ;
Veterinary-Surgeon Schmidt himself observed the fly
between Mt. Kikundi and the Uluguru Mountains
(January 1901). The whole of the road as far as
Mpwapwa is for the present suspected, while further on
as far as Tabora it appears to be free from Tsetse-flies.
On the other hand cases of infection have been observed
on the Tabora-Ujiji road, probably on the upper Mlagarasi
brook. The road from Iringa to Kilossa is very risky,
and especially on the Great Ruaha River (called Mpagali
from Tayeta to beyond the point where the Lukose
debouches into it) is it impossible to keep any cattle.
Similarly no cattle can be kept in Marore. The Tsetse
belt begins probably at the Great Ruaha and extends as
far as Kirengawana. In the same way the road from
Iringa, by Dwangire-Ngahoma, to Kungulio is risky.
Inland from Kilwa the Tsetse-fly has definitely been
shown to occur on the Donde road between the 29th and
52nd kilometres (Knochenhauer), and also near Gerengere
village (Veterinary-Surgeon Schmidt). For this reason the
choice of this place as agricultural experiment station for
the commune of Kilwa is not a happy one. On the road
from Dar-es-Salim to Songea two mules were attacked by
Surra, as also were cattle on the journey from Songea
to Lukuledi. At that time (July 1899), according to
reports received from Lukuledi Mission, the Tsetse-fly
was observed at the camping-places, Kokoma, Chamba,
Darhammam, and Chitwanga, and especially in Chamba.
In the latter place it has been experimentally proved by
the Songea Station that cattle kept for only two days
become infected by Surra.

“The great cattle-breeding countries of the inter-
lacustrine region, Ugogo, Unyamwesi, the region devoid
of watercourses, Masailand, and so forth appear to be free
from Surra.

‘‘ Nevertheless, it appears from the facts ascertained
up till now, that on account of the risk of infection the
transporting of loads by means of draught animals is only
possible in certain places in the colony, and that, there-

IMMUNITY TO TSETSE-DISEASE. 263

fore, we must begin to build railways, if we would replace
the system of carrying loads on the heads of niggers”
(pp. 146-147). iil wi

Discussion of the question of immunity, or “ facultative’
immunity to Tsetse-fly disease (p. 148). The following is a
translation of the concluding sentences of this section:
“Tt was thought, hitherto, that the gray, or so-called
Masai donkey is immune; but in Mombo at the foot of
the Usambara Mountains, the parasites of Surra have
been detected in the blood of Masai donkeys which fell
sick there. Perhaps, however, the greater number of
Masai donkeys are facultatively immune, At any rate,
the infection experiments of Koch had a negative result
(Reiseberichte, p. 70). It is probable that immunity
may also be produced artificially. Professor Koch
and Veterinary-Surgeon Schmidt have already made a
beginning here with experiments in this direction, while
experiments with a serum-treatment were recently carried
out by Dr. Schilling, in Togo, apparently with success.
(Sitzungsber. des kolonialwirtschaftlichen Komitees, 1901).”

Veterinary-Surgeon Schmidt's views as to the nature of
Tsetse-fly disease (pp. 148-150).

[Translation.] “ Veterinary-Surgeon Schmidt has re-
corded his opinions on Surra in the following memorandum,
addressed to the Songea Station and here reproduced
verbatim :—

“61. In a district free from Tsetse Surra cannot be
conveyed from one animal to another either directly or
indirectly. Flies and horse-flies other than the Tsetse
are of no importance as regards the occurrence of Surra,
since, analogously to all parasitic diseases, a correlation
exists between host, parasite, and intermediate host, so
that the disease is not developed if only a single one of
the three conditions is not fulfilled.

‘<¢(a) The parasite lives only in the blood of animals
susceptible to the disease (single- and double-hoofed
Ungulates),* but can be transmitted to almost all animals
by artificial inoculations.

* The limitation of Tsetse-fly disease to Ungulates is not in accordance
with the result of Lieut.-Col. Bruce’s experiment, in which the disease
was produced in a dog by inoculation with blood from a hy#na (see
Chapter VII., Appendix A, p. 280).—E. EH. A.

264

MEMORANDUM ON TSETSE-DISEASE.

“*(b) The only intermediate * host is the Tsetse-fly, in
which the parasite passes through a process of development
which has not yet been observed.

*«¢(c¢) After attaining its full development the parasite
once more enters the blood of the animals susceptible to
Surra, by means of the bite of the Tsetse-fly, and after an
interval of about four to six weeks produces noticeable
symptoms of disease.

«2. In inoculation experiments the interval which
elapses until symptoms of the disease manifest themselves
is usually from four to six weeks.

***(a) Consequently at the same period parasites can

also be demonstrated in the blood. Analogously to malaria,

fresh attacks of fever always occur simultaneously with
the development of a new generation of the parasites,
while in the interval the temperature is normal, and it
is also impossible to demonstrate the presence of the
parasites by means of the microscope. It is therefore .
necessary to take samples of blood every three or four
days, corresponding with the process of development,
shortly before a fresh attack of fever supervenes.

‘**«The number of the parasites, the condition of the
animal, and the amount of work demanded from it all
affect the course of the malady. When carefully tended,
animals suffering from Surra live for years, while great
bodily exertion renders the course of the disease acute.

“<“(b) Simultaneously with the appearance of the
parasites in the blood careful observation alse shows
symptoms of the disease (intermittent fever, cessation of
appetite, sluggishness of movements). Noticeable symptoms
occur only after the disease has lasted some time, in the
shape of emaciation and anemia of the mucous membranes.
The only positive proof for the expert is and remains the
demonstration of the parasites in the blood. Cattle are
more susceptible to Surra than mules.

«3. It is not permissible to regard districts rich in
game as being infected with Surra, since all known
single- and double-hoofed Ungulates succumb to the
disease ; neither, for the same reason, can the occurrence
of Tsetse-flies be associated with that of the buffalo.’

“ According to this, Veterinary-Surgeon Schmidt

* Cf. p. 260, note *.—H. H. A.

TSETSE AND TRYPANOSOME. 264

appears to be of the opinion that wild animals are also
susceptible to Surra.” *

Notes on the habits of the Tsetse, and on the Relations
between it and the TryPANosoma (p. 150).

[Translation.] ‘So far as I am aware, not many
observations upon the bionomics of the Tsetse exist. The
fly appears to live in reed-beds bordering streams, and in
damp bush. It flies at its victim with a buzzing noise,
bites it in the flanks, on the belly, and elsewhere, and,
according to an observation by Veterinary-Surgeon
Schmidt, disappears again with equal rapidity. It is
said not to pursue its yictim. The fly bites only during
the day, or, at most, on bright moonlight nights in
addition. Consequently in most cases Tsetse-districts can
be passed through with animals without harm in the
night.

“The mode of development of the Tsetse is still, so
far as I am aware, entirely unknown.t A yet richer field
for research, however, is offered by the investigation of
the development and life-history of the blood-parasite
(Trypanosoma), which is the actual cause of Surra disease.
Probably, just as the plasmodium of malaria does in
Anopheles, the parasite passes through a special generation
in the body of the fly. Undoubtedly, just as in the case
of malaria, it is possible for one ox to be directly infected
with Surra from another by means of inoculation of
blood ; but the normal course of development will be, as
in malaria and Texas fever, that in the biting insects
the Trypanosomes pass through a special developmental
cycle (sexual reproduction). In malaria, according to
the investigations of Ross and Grassi, the parasites
(Plasmodium) pass into the stomach, thence into the
stomach-wall, and finally into the salivary glands.”

On pp. 151-152 a diagnosis of the genus Trypanosoma
is given, derived from Biitschli’s article on Protozoa in
Bronn’s “ Klassen und Ordnungen des Tierreichs.” On

* In other words, if Schmidt’s view is correct, the parasite of Tsetse-
fly disease will only exceptionally be found in the blood of an antelope,
buffalo, or zebra; since its presence in the blood would entail the death of
a wild animal equally with that of a domesticated one. This, however, is
contrary to the experience of other observers.—E. EH, A.

+ Dr. Stuhlmann is evidently ignorant of Col. Bruce’s discovery on
this point.—H. E. A.

6

bo
for)

POINTS FOR FUTURE INVESTIGATION.

p- 152 the author suggests that, among others, the
following points should be aimed at in future investigation
of Tsetse-fly disease :—

“1, To bring together for examination specimens of
biting flies from as many different points as possible.

“2. To observe the habits and life-history of the
Tsetse.

“3. To find out the places in which, according to the
experience of the natives, cattle can never be kept.

“4, To make a careful examination of the blood of
many diseased animals, especially in cases in which it
is possible to state definitely where the infection took
place.

“5. To ascertain whether, in the case of the different
kinds of animals, the disease is conveyed by one and the
same species of biting fly.

“6. To investigate the fate of the Trypanosomes in
the Tsetse, and the manner in which they pass from it
into the mammal (salivary glands ?)

“7. To investigate the susceptibility of big game,
facultative immunity, especially in young animals, and
artificial immunity owing to treatment with serum.”

XXVII. 1902, Dr. Schilling.

“‘ BeRIicHT UBER DIE SuRRA-KRANKHEIT DER PFERDE
unD Rinper im Scuurzcesrer Toco” (Centralblatt fiir
Bakteriologie, Parasitenkunde und Infektionskrankheiten.
Erste Abteilung: Medicinish-hygienische Bakteriologie
und tierische Parasitenkunde. XXXI. Band. No. 10).

XXVIII. 1902. Dr. Hans Ziemann.

XXIX,

“ TsE-TSE-KRANKHEIT IN Toco (West Arrika) ” (Ber-
liner klinische Wochenschrift, 1902, No. 40. Separate
pagination pp. 1-18).—Translated by P. Falcke, in “ The
Journal of Tropical Medicine,” Vol. V., No. 23, Dec. 1,
1902, p. 367, et seq.

1902, R. M. Forde, L.R.C.S., &c.

“Some Cuinican Norges on A European Patient IN
wHOosE Bioop a Trypanosoma was OsserveD” (The
Journal of Tropical Medicine, No. 17, Vol. V., September 1,
1902, pp. 261-263 : with temperature charts, and a plate
showing the parasite in a stained preparation of blood).

REFERENCES TO LATEST LITERATURE. 267

XXX. 1902. R. Boyce, M.B., F.R.S.; Ronald Ross, C.B., F.R.S.,
F.R.C.S.; C. S. Sherrington, M.D., F.R.S.

“ Note oN THE Discovery oF THE Human TRYPANO-
some” (The British Medical Journal and The Lancet,
Nov. 22, 1902).

XXXI. 1902. R. M. Forde.

“Tur Discovery oF THE. Human TRYPANOSOMA ”
(The British Medical Journal, Nov. 29, 1902).

XXXII. 1902. L. W. Sambon, M.D.

“ Nore on THE Discovery oF THE HUMAN TRYPANO-
some ” (The Lancet, Dec., 5, 1902).

XXXIII. 1902. A. Laveran.

“ RECHERCHES SUR LE TRAITEMENT ET LA PREVENTION
pu Nacana” (Annales de l'Institut, Tome XVI., No. 11
(25 Nov. 1902), pp. 785-818).

XXXIV. 1903. Dr. Schilling.
“On NAGANA AND OTHER TryPANOSOMES” (The Journal
of Tropical Medicine, No. 3, Vol. VI., February 2, 1903,
pp. 45-47).

268

‘CHAPTER VII.

A..P.P END LC TS.

APPENDIX A.

Apstract oF Lieur.-CotoneL Bruce’s “ Furtuer Report on
THE TsETsE-FLty Disease” (‘‘FurTHER REPORT ON THE
Tsetse-Firy Disrase or NaGana IN ZULULAND. By
Surgeon-Major Davin Bruce, A.M.S.—Ubombo, Zululand,
29th May, 1896.”) (London: Harrison & Sons, St. Martin’s
Lane, Printers in Ordinary to Her Majesty. 4th February,
1897. 69 pp., sm. folio, Plates I.-VI.)

Since this is by far the most important work that has yet
appeared on Tsetse-fly Disease as affecting animals in Africa
itself, besides being, in conjunction with the no longer accessible
*‘ Preliminary Report,” of epoch-making interest as the memoir
in which the disease was first proved to be due to a hematozoon,
it seems advisable to give a detailed analysis of its contents.

The Report is prefaced by a covering letter—“To His
Excellency The Hon. Sir Walter Hely-Hutchinson, K.C.M.G.,
Governor of Natal and Zululand, etc., etc.,” in which the author,
after stating that,in accordance with His Excellency’s instructions,
he “left Pietermaritzburg on the 21st August, 1895, and arrived
at Ubombo, Zululand, on the 8th September, 1895, for the purpose
of investigating the Tsetse-Fly Disease, or Nagana, as it occurs
in Zululand ”—gives the following statement of its contents :—

“1.—A description of the Tsetse-Fly, with experiments
designed to show the part this Fly takes in the causation of
the disease.

“‘2.—A description of the Hematozoon or Blood Parasite,
which is the cause of the disease.

“‘3.—The results of experiments having for their object proof
of the connection which exists between the Big Game and the
spread of the disease.

CONTENTS OF BRUCE'S FURTHER REPORT. 269

“4,—A description of the disease as it affects domestic
animals, with illustrative cases.

“‘5.—Inoculation and feeding experiments to show the
communicability of the disease from affected to healthy animals.

“‘6,—Treatments of the disease, prophylactic and curative.”

The “Report” is illustrated with six plates (in addition to
temperature charts) as follows :—

Plates I. and II. are devoted to the “ Tsetse-Fly.”

Plate I. contains the following coloured figures :—

1.—Fly at rest (natural size). 4.—Fly at rest (x 2).
2.—Larva (natural size). 5.—F ly with expanded wings (x 2).
3.—Puparium (natural size), 6.—F ly before feeding (x 2).

7.—Fly after feeding (x 2).

Plate II. is devoted to reproductions from photographs, as
follows :—

1, 2.—Fly with expanded wings. 5.—F ly during parturition.
3.—F ly at rest. 6.—Pupa.
4,—F ly pregnant with larva. 7, 8.—Flies in various positions.

As has already been remarked in the systematic portion of
this work (p. 89), the photographs appear to have been taken
from specimens of Glossina pallidipes, Austen, while the coloured
figures seem to represent Gl. morsitans, Westw.: the apparent
discrepancy may be due to minor inaccuracies in the coloured
figures.

Plate III. shows hematozoa in the blood of the dog, horse,
and cow, “all magnified 1,000 times.”

Plate IV.—‘“ Horse suffering from Fly Disease or Nagana.”

Plate V.— Donkey suffering from Fly Disease or Nagana.”

Plate VI.—* Dog suffering from Fly Disease or Nagana.”

ConTENTS.—

1.—Definition
2.—Nomenclature 5
rt —Distribution in Zululand..
4,.—Historical Account
5.—Htiology ..
A,—The Tsetse- -fly A ‘ Be 55
1.—Description of .. Po ad A
2.—Experiments with
3.—The Hematozoon or Blood Parasite of the
Fly Disease a 17
B.—Relation of the Big Game to the Fly Disease.. 19
6.—Description of the Fly Disease or Nagana as it occurs

PAGE

FP phd NNR er

in the Domestic Animals .. = i at san 2
A.—In the Horse xe ae 3s a we 25
B.—In the Donkey na ar 8 an a» £30
C.—In Cattle a3 ae ‘F 3. ie Soe

D.—In the Dog .. te oe cs “sf ce Oe

270 DEFINITION OF TSETSE-FLY DISEASE.

PAGE
eels eso oes of Blood from Affected to Healthy Animals 41
A.—Inoculation of Blood in which the Hematozoa

can be demonstrated by the Microscope .. 41
B.—Inoculation of Blood in which the Hematozoa
cannot be demonstrated by the Microscope on 0
8.—Feeding Experiments... ee a 4 te ae
9.—Treatment, Medicinal 4c Ris Saye
A.—Arsenic as a Curative Agent és $s he AS;
B.—Arsenic as a Prophylactic pean * aa Oe
10.—Concluding Remarks sa on Br <5 Ob

The following is the author’s “ Definition” of Tsetse-Fly
disease :—

“The Fly Disease or Nagana is a specific disease which
occurs in the horse, mule, donkey, ox, dog, cat, and many other
animals, and varies in duration from a few days or weeks to
many months. It is invariably fatal in the horse, donkey, and
dog, but a small percentage of cattle recover. It is characterised
by fever, infiltration of coagulable lymph into the subcutaneous
tissue of the neck, abdomen or extremities, giving rise to swelling
in these regions, by a more or less rapid destruction of the red
blood corpuscles, extreme emaciation, often blindness, and the
constant occurrence in the blood of an infusorial parasite, either
identical with or closely resembling the Trypanosoma Evansi
found in Surra, a disease of India and Burma. On post-mortem
examination the following changes are noticed ; deposition of a
yellow jelly-like material in the subcutaneous tissue, inter-
muscular layers, and under the serous covering of the heart ;
with purplish stains or ecchymoses in various regions, as on the
inner aspect of the skin, the serous membrane covering the
lungs, and outer and inner surfaces of the heart; enlargement
and softening of the spleen and congestion and fatty degenera-
tion of the various organs ” (p. 1).

Meaning of the term ‘‘ Nagana.”—“ Nagana” is a Zulu word
meaning “ to be low or depressed in spirits,” and has reference
to “ the symptoms We by the animals suffering from the
disease” (p. 1).

In the Congo Free State the disease is known as “la mouche ”

1).
ie Distribution of the disease in Zululand.— Broadly it may be
stated that: the disease is limited to certain tracts, the physical
conditions of which imply heat and moisture. These tracts in
Zululand are situated in the level coast plain which extends some

fifty miles inland, and in the river valleys which enter or debuuch
on this plain” (p. 1).

‘‘ ErioLocy.—The opinions of the Europeans settled in Zulu-
land and of the natives themselves are so conflicting that little or
no good will be gained by entering fully into their evidence.

REPRODUCTION IN ZULULAND TSETSE. 271

“‘ Two theories are held.

«“ A.—That the disease is caused by the bite of the Tsetse-fly.
This is the European theory, and, as everyone knows, has been
popularly prevalent ever since white men first landed in South
Africa.

‘“‘ B.—That the disease is caused by the presence of large
game, the wild animals in some way contaminating the grass or
drinking-water by their saliva or excretions. This may be called
the native theory ” (p. 2).

A.—Tue Tserse-Fry.—The Tsetse “acts as a carrier of a
living virus, an infinitely small parasite, from one animal to
another, which entering into the blood stream of the animal
bitten or pricked, there propagates and so gives rise to the
disease” (p. 2).

Position of the wings when the fly is at rest.—* When the
Tsetse is at rest the wings fold over one another like the blades
of a pair of scissors, mC c the fly an elongated appearance
(Plate I., figs. 1 and 4)”

Mode of reproduction, we the Tsetse-fly does not lay eggs as
do the majority of the Diptera, but extrudes a yellow- coloured
larva nearly as large as the abdomen of the mother. This larva
is furnished with a black hood at one pole and two minute spikes
at the other. It is annulated and consists of ten segments.
Immediately on being born the larva creeps about with a good
deal of activity, evidently searching for some cover or hole in
which to hide. Having found a resting-place, it immediately
begins to change colour, and after a few hours has turned into
a jet-black hard pupa or nympha (Plate I., figs. 2 and 3).

“Tf these pupal cases are placed in a perfectly dry place, as
in a wooden box, the perfect insect hatches out in about six
weeks. From this it would appear that the life-history of this
species of fly is very simple, it only being necessary for the
female insect to deposit the larva on the surface of the soil or in
the grass, when the larva creeps into the nearest shelter, in a
few hours becomes hard and black, and in five or six weeks
hatches out into the fully-developed Tsetse-fly.

“Tt has often been surmised that this fly is bred in buffalo

dung, but from a consideration of the foregoing facts it is evident
that nothing is wanted except any moderately dry place”
pp. 2-3).
; Habits of Tsetse-flies—“ It is astonishing with what rapidity
the flies fill themselves ; in as small a space of time as twenty or
thirty seconds a fly will become swollen out like a balloon with
bright red blood.

“On entering ‘ Fly Country,’ one is not left long in ignorance
of the presence of the Tsetse. The natives may be seen slapping
their naked legs, the dogs bite round, and the horses kick. The
Tsetse, however, may be said to be somewhat local in its distribu-

272 HABITS OF TSETSE-FLIES.

tion in the Fly Country, being only met with now and then and
in few numbers, until you enter some glade or clear space in the
thorns, when suddenly the slapping, biting, and kicking go on
with tenfold energy, and you can catch thirty or forty flies in a
few minutes,

“The Tsetse has a direct flight, flopping, if I may use the
term, suddenly on the animal attacked, and is very pertinacious.
It affects in horses the legs, especially in the region of the fetlock,
and on being detached from its position by the vigorous kick the
horse gives, will only rise a few inches and again settle near the
same spot. After the initial prick there is no irritation, and the
animal will stand quietly while the fly has its feed.

“In man the burying of the proboscis in the skin is accom-
panied by a sharp prick, which draws one’s attention suddenly to
the spot ; but the pain is, as a rule, trivial, and the subsequent
redness and irritation very slight indeed—in fact, less than that
caused by the mosquito.

“But under some circumstances the Tsetse may become
almost unbearable. In January, when camping in the Fly
Country close to the River Mkusi, these flies were very numerous,
and in conjunction with the intense heat—the thermometer often
registering 100° F. to 106° F. in the shade of a double tent—
almost drove one mad, One can easily imagine the effect of the
constant worrying of these pests by day, followed by mosquitoes
at night, and minute ticks at all times, on any nervous system
except the most stable. The horses fared no better, and could often
be seen trotting about trying to escape from their tormentors.

* About sunset seemed a favourite feeding-time, and then the
poor creatures would be surrounded by a perfect cloud of the
flies, while some hundreds of them would be settled on them at
the same moment.

“JT have not noticed them biting during the night, but that
excellent sportman and observer Selous states that by moonlight
in some places they bite just as furiously as in the daytime.

“The fly makes a loud buzzing sound when flying, but after
its feed and at rest it emits a peculiar sharp, shrill note, probably
caused by an action of the wings.

“In horses, dogs, and cattle I have usually failed to find any
swelling or symptoms of irritation following the bite, but on
several occasions a well-marked soft swelling under the skin of
the part bitten, as large as a hazel nut, was observed, and this
persisted for some time.

“The Tsetse is not at all easy to catch with the hand,
especially during the day, being nimble and quick of movement,
but at early morning or sunset they become more lethargic, and
are then more readily secured. When I wanted living specimens
for any purpose I provided myself with a butterfly-net, with
which it is easy enough to catch them unhurt.

EXPERIMENTS WITH TSETSE. 273

“The Tsetse differs from the mosquito in that both sexes are
blood-sucking flies.

“Tt is said that the fly follows the large game, but I never
found them in any number in places in which a few minutes
before large herds of buffalo or wildebéeste had been standing.
Of course, the fly may have literally followed the game. One
day, after killing a wildebeeste, I certainly found a large number
of Tsetse on the dead animal, and again on a dead bufialo I
counted as many as forty flies feeding on it at the same time”

(p. 3).

Experiments with the Tsetse-fly (pp. 4-16).—These experiments
are divided into seven series, and were made with a view to the
elucidation of the following questions :—

“(a.) Is the Tsetse-fly capable of giving rise per se to any
local or general disease in susceptible animals ?

“(b.) Can the. Tsetse-fly convey the disease from an affected
animal to a healthy one?

“(c.) How long does the Tsetse-fly retain this infective power ?

*“(d.) How long does the blood of an affected animal remain
capable of giving rise to the disease in a dried condition ?

*“(e.) How long does the blood of an affected animal remain
capable of giving rise to the disease if kept in a natural con-
dition ?

“(f.) Does the Tsetse-fly under natural conditions convey the
disease from animal to animal ?

“(q.) Is the Tsetse-fly capable of giving rise to the disease if
taken out of the Fly Country into a healthy locality?”

(a.) Details are given of three experiments in which various
numbers of Tsetse-flies, brought from the Fly Country and then
kept in captivity at Ubombo for some days, in order to give time
for the hematozoon to disappear, in case it had been present
in the bodies of the flies when they were caught, were sub-
sequently fed daily on dogs during periods varying from ten days
to two months. All the dogs remained perfectly healthy, and
continued so after the experiments were stopped. The author
adds: “The above experiments then prove, I trust satisfactorily,
that the Tsetse-fly is not capable of giving rise per se to any local
or general disease in susceptible animals” (p. 4).

(b.) In connection with the question: “Can the Tsetse-fly
convey the disease from an affected to a healthy animal!” the
author writes as follows (p. 5) :—

“That all blood-sucking flies are not capable of transferring
the Fly Disease from affected to healthy animals is, I think,

T

274 OTHER FLIES SEEM NOT TO CARRY PARASITE.

shown by the fact that up here at Ubombo, where we have
several species of these pests, no single instance of the disease
arising spontaneously has occurred, although healthy horses,
cattle, and dogs have been constantly and closely associated with
those suffering from the disease. Why this should be so is at
present a mystery, and it is to be hoped that some point may be
discovered which will throw light on the subject. There may be
some anatomical peculiarity in the Tsetse which enables it to act as
carrier, or there may be some undiscovered fact in the life-history
of the parasite associating it with this particular species of fly.

“One fact, however, must be borne in mind, and that is the
enormously greater number of Tsetse-flies in the Fly Country than
any other species of blood-sucking fly met with there. I have
seen as many as 200 Tsetse-flies on a horse at the same time ;
and when it is stated that I have often caught and put into a
cage singly some fifty Tsetses in half-an-hour, some conception may
be gained of the enormous number of Tsetse-tlies which may
visit a horse during its sojourn for one day in the ‘ Fly.’

“The act of feeding is remarkably quick. From the moment
of settling on the animal until the fly is fully blown out with
blood is often as little as twenty to thirty seconds. But still this
does not explain why none of the healthy animals up here on the
top of the Ubombo have become infected by their diseased neigh-
bours, because we have many small stinging flies—some species
probably of Stomoxys—which cause drops of blood to exude from
the legs and ears of our animals. But this I can assert, and it is
an important point, that in no single case, as far as I am aware,
has any case of the disease occurred up here due to infection from
the diseased to the healthy.

“That the ‘Tsetse-fly can act as a carrier of the parasite is, I
think, shown by the following experiments.”

Details, with charts, are given of two experiments in which
eight flies were allowed to feed on a healthy dog immediately
after having fed for a short time on a dog suffering from the
disease. ‘The procedure was repeated on several days, and in
each case on the sixth day after the flies were last fed the dog
was found, by microscopical examination of his blood, to be
suffering from the disease. The author adds: ‘ Up to the present
then from these two series of experiments it is seen in the first
place that the fly per se does not give rise to any local or general
disease, and this is further borne out by other experiments in
which I placed minced up flies under the skin of dogs without any
results, and secondly it proves that the fly can act readily as a
carrier of the Fly Disease from affected to healthy animals” (p. 6).

““(c.) How long does the Tsetse-fly retain this infective
power ? !
“ Now as one of the main objects of this investigation is to

DURATION OF INFECTIVE POWER OF TSETSE, 275

establish the part, if any, that the wild animals play in the
dissemination of this disease, and as my working hypothesis at
present is that some species of animals living in the Fly Country
harbour the Nagana parasite in their blood, and that the Tsetse-
fly carries the infection from affected to healthy animals, much in
the same way as the vaccinating needle carries the infection of
vaccinia from child to child, it will evidently further this object,
if it can be discovered how long the Tsetse-fly retains its infective
power. In passing through Fly Country of course it is possible
that a Tsetse-fly may be feeding on a wild animal one minute
and the next have transferred itself to the horse, ox, or dog ; but
if it can be proved to retain the power of infection say for
twenty-four hours, the fly’s power to do harm is evidently much
widened” (p. 6).

Details, with one chart, are then given of three experiments
on native dogs, in which various numbers of Tsetse-flies, from six
to twelve, were repeatedly fed on the subject of the experiment,
twelve, twenty-four, and forty-eight hours respectively after
feeding on an affected animal. The results were that hematozoa
were found in the blood of the dog on the 37th day, with the
twelve-hour interval; on the 38th day, with the twenty-four
hour interval; and on the 32nd day, with the forty-eight hour
interval ; reckoning i in each case from the date of the commence-
ment. of the experiment,

“‘(d.) How long does the blood of an affected animal remain
capable of giving rise to the disease in a dried condition ?

“This point may be approached from another direction and
some control placed on the results. Instead of using the proboscis
of the Tsetse as the conveying instrument, let us use short pieces
of thread dipped in blood containing the hematozoa, dried, and
at varying periods of time threaded, by means of a needle, under
the skin of healthy dogs” (p. 8).

Details, with charts, are given of seven experiments in which
this procedure was carried out with varying results, as to which
the author writes (p. 10): ‘‘These experiments would go to show
that the blood of animals affected by Fly Disease retains its
capability of transmitting the disease in a dried condition for
twenty-four hours, but that this is exceptional, and that at the
end of forty-eight hours the blood is inert.

«But as the proboscis of the Tsetse may retain blood ina
moist condition, it will be interesting to show how long the blood
itself, taken aseptically and preserved by suitable means from
putrefaction, will retain this infective power.

“(e.) How long does the blood of an affected animal remain
capable of giving rise to the disease if kept in its naturally moist
condition ?”

y 2

276 EXPERIMENTS UNDER NATURAL CONDITIONS.

Here follows a statement (with one chart) of the results of
eight experiments, as to which the author remarks: ‘“ These
experiments point to the fact that the blood of animals suffering
from Fly Disease can retain its infective power for four days but
not for seven” (p. 11).

“(f.) Does the Tsetse-fly under natural conditions convey the
disease from animal to animal?

“ Having seen that the Tsetse-fly cannot of itself give rise to
disease; that, on the other hand, it can readily act as a carrier,
and that it is able to retain its infective power for at least
twenty-fcur hours, let us now consider whether in a state of
nature it really does convey the disease” (p. 12).

Details and charts are given of five experiments in which
perfectly healthy horses were taken down into the Fly Country
for a few hours, and were there allowed to be bitten freely by
Tsetse-flies, but were not permitted to eat or drink until their
return to the top of the hill. All the horses contracted the
disease, and the author writes :—‘“ These five experiments show
abundantly that horses cannot be taken with impunity for a few
hours into the Fly Country, even although they are not allowed to
eat or drink there, and they afford a strong presumptive proof
that the disease is carried to them by the Tsetse-fly.

“This last series of experiments, however, proves less than at
first sight it might be supposed. It only proves that susceptible
, animals cannot be taken into the Fly Country, even although
they are not allowed to eat or drink there. It does not absolutely
prove that the disease is carried to them by the Tsetse-fly.
There may be other ways of taking the disease, for example, by
inhalation. ... This is a hard thing to understand, and in
order to make it possible the parasite of the Fly Disease must be
able to exist in some other form than that in which it is found
in the blood. The supposition would be that it forms a resting-
stage or spore form in which it can exist as a dry, impalpable
dust. For my part I have much ditliculty in believing that
animals are infected as a rule with Fly Disease by inhaling the
materies morbi, and until I find animals still susceptible to the
disease which are protected in some way or other both from
feeding and the fly, I shall continue to be sceptical.

“On account then of the lack of absolute proof furnished by
the last experiments, I set myself on the 22nd November to try
to infect susceptible animals with the disease by having them
bitten by flies brought: up daily from the low country and
straightway placed on the animals,

“(g.) Is the Tsetse-fly capable of giving rise to the disease if
taken out of the Fly Country into a healthy locality ?
“The method of carrying out this experiment was to go déwn

FATE OF HAXMATOZOA INGESTED BY TSETSE. 277

to the Fly Country in the early morning, catch the flies, return
to the top of the Ubombo, and straightway place them on the
animal under experiment.

“The greatest care was taken that the flies were placed on a
perfectly healthy animal, as to have allowed them to puncture
one already affected by the disease would naturally vitiate the
experiment,

“The time which elapsed between catching the flies and
placing them on the animal under experiment varied from four
to seven hours” (p. 15).

Details and charts are given of two experiments, one on a
horse, the other on a pointer dog. The animals were allowed to
be bitten in the manner indicated by a varying number of flies
on different dates, with the result that both contracted the
disease.

With reference to the outcome of the experiments the author
remarks :—‘‘ As these experiments have had a successful result,
I now consider it proved that the Tsetse-fly does commonly, in
a state of nature, convey the disease from animal to animal, and
that on the other hand there is no proof that the drinking of
water or the eating of soiled herbage plays any réle in the
process” (p. 16).

Pages 17 to 19 are devoted to “ Tor Ha#marozoon or BLoop
PARASITE OF THE FLY DIsEAsE.”

With reference to the possibility of the existence of a resting-
stage, the author writes:—“We have seen by the thread
experiment that the hematozoa retained their vitality only on
one occasion for twenty-four hours in a dried condition, and for
less than seven days in a moist condition, and that after this they
were no longer capable of giving rise to the disease. This proves
that under these conditions there had been no spore or resting-
' stage formation” (p. 18).

As to “ The fate of the Hematozoa when ingested by the Tsetse-
Sly,” Col. Bruce remarks :—

“ A priori, one would think that the heematozoa on being taken
into the stomach of the Tsetse-fly would soon perish on account
of the processes of digestion which one imagines going on. It
will therefore be interesting and bear directly on the subject if
we trace the history of the hematozoon afterit has been removed
from the blood-vessels of an affected animal and swallowed by
the Tsetse-fly. I therefore caused the flies to feed on an animal
suffering from Nagana, whose blood contained numerous hema-
tozoa, and at hourly intervals subjected the proboscis, the contents
of the stomach, and the contents of the lower end of the intestine
to microscopical examination.

* * + * *

278 RELATION OF BIG GAME TO FLY-DISEASE:

“The result of my examination up to the present of the
Tsetse-fly is briefly this :—

“Tmmediately after feeding, the tube of the proboscis can be
seen to be crammed full of red blood corpuscles, among which the
hrematozoa can be seen actively wriggling. Up to 46 hours
after feeding I have seen living hematozoa and red blood
corpuscles in “the proboscis, After “)18 hours the hematozoa are
still very numerous and vigorously active in what remains of the
blood in the stomach: .After 140 hours the stomach is empty.
After 25 and 70 hours IT have seen many motionless hematozoa
in the faces, but I have never seen at any time any appearance
of life in the hematozoa in the feeces or contents of the lower
part of the intestine. The parasites appeared, however, un-
changed in form, and I intend making some injection experiments
with the feces to find out if these motionless hematozoa have
any vitality left in them” (p. 19).

Pages 19 to 24 are devoted to the second part of the account
of the etiology of Nagana, viz. :—

“ B.—ReELaTIoN oF Bic Game To Tue Fry Disease.

“As we have found that the Tsetse-fly in a state of nature
does act as a carrier of the hematozoa of Fly Disease, it is evident
that it must procure the parasite somewhere, and what more
natural than it should procure it from the blood of warm-blooded
animals living in the Fly Country ?

“Tt is not necessary to suppose that in these animals Nagana:
is a fatal disease, but only that the big game harbour the
parasite which causes the disease for a longer or shorter time
with little or no disturbance to health. We have seen, as in the
case of the heifer mentioned on page 33, that cattle may have
the parasite in their blood for at least eighteen months without
causing death, and it is possible that some of the big game die of.
this disease. When in the Fly Country quite lately, I chanced
on a dead wildebeeste surrounded by vultures. The skin of this_
animal was quite intact and showed no signs of its having come
to a violent death. It was somewhat emaciated and had the
appearance of an animal dead of Nagana. I did not inject any
blood from it into a susceptible animal, as it was in a state of
decomposition ” (p. 19).

ae # % *: *

“Tn regard to this relation of big game to the Fly Disease,’
there can be no question as to the very widely spread opinion
which exists regarding the wild animals as in some way or other
responsible for its propagation, and in this connection the buffalo,
wildebeeste, quagga, waterbuck and koodoo are usually named.
Europeans and natives in all parts of the country are found who

EXPERIMENTS WITH BLOOD FROM BIG GAME. 279

state most emphatically that where there is no game there is no
Nagana, and example after example could be cited of the state-
ment that the game having been driven out, the disease dis-
appeared, and on the. reappearance of the game that the cattle
again became sickly. Those who hold the Tsetse-fly theory
explain this by affirming that where the big game is there also is
the Fly. Others, as mentioned above, are of opinion that the
game contaminate the grass and drinking-water and so infect
the cattle.

“That the presence of the wild animals in the vicinity of
horses and oxen is not the only factor in the problem is shown
by the fact that in the old days when the big game was numerous
and roamed over the whole country, hunters and travellers never
complained of the Fly until they encountered the disease in low-
lying tracts of country, or along the large river valleys ; and at
the present day it is stated, on the authority of Mr. L. Peringuey,
that in the Hermansdorp District of Cape Colony, where herds of
buffalo are still found, the Fly Disease is unknown, nor has the
Tsetse-fly itself been ascertained to occur there. Now we know
the disease to be caused by a blood parasite, and that the. disease
can be carried by the Tsetse from affected to healthy animals, we
can see how such theories may have some foundation in fact.

«“ At first I thought that by a careful microscopical examination
of the blood of the big game some light could be thrown on the
subject, and to this end I submitted the blood of buffalo, wilde-
beeste, koodoo, impala, bush-buck, reed-buck, and the smaller
varieties shot in the heart of the Fly Country to a lengthened
microscopical examination, but with no result.

«Thinking, however, that the parasite might exist in’ too
small numbers, as we have seen is frequently the case in cattle,
to be readily discovered by direct examination, I next instituted
a series of experiments by injecting a moderately large quantity »
of blood, taken immediately after death, into dogs, as it has been
proved by experiment that in cases in which the parasite could
not be demonstrated by the microscope, an inoculation experi-
ment of this kind readily discovered them. | .

“Of course, it was necessary that the dogs used for experi-
ment should be kept at the top of the Ubombo, as the results
would have been useless if they had been exposed to infection in
any other way than by the injection of the blood.

«This meant a delay of several hours between the procuring
of the blood and the injection of it into the dog, as the wild
animal might be shot five or ten miles from the foot of the hills,
and the Ubombo itself is some 2,000 feet high.

‘Again, it would evidently facilitate matters if the blood
could be kept liquid, as the blood coagulum might entangle in
itself all the parasites contained in the blood.

“J therefore made a preliminary experiment to find out if

280 RESULTS OF INOCULATION EXPERIMENTS.

blood kept liquid by some suitable means, and kept in a bottle at
the temperature of the air for several hours, would still retain its
infective power” (p. 20).

In the preliminary experiment 10 c.cm. of blood were taken
from a dog suffering from Nagana, mixed with “a twentieth of
its bulk of a 0°5 per cent. citrate of potash solution,” then kept
“in a bottle at the temperature of the air (80° F.) for seven hours,”
and finally injected into a healthy native dog. Hematozoa of
Fly Disease were found in the blood of the dog after five days,
“and he died of the disease twelve days later.”

“ Tt was evident then that we might expect successful results
by injecting blood procured in the Fly Country and kept liquid
by the above-mentioned means into native an kept on the top
of the Ubombo” (p. 21).

On pages 21 to 23 is given a statement, with results, of
thirty-four experiments on native dogs, carried out on the above
lines, with blood from various species of wild animals. An
analysis of the results shows that in five cases the dog died from
an unknown cause; in twenty-one cases the dog remained healthy ;
in eight cases the dog contracted fly disease.

On page 24, there is printed a tabulated statement showing
“the results of the above and other experiments, the various
species of wild animal being tabulated together.”

The following is an analysis of the results :—

Out of eight dogs inoculated with blood from buffaloes, one
dog contracted the disease.

Out of thirteen dogs inoculated with blood from wildebeeste,
three contracted the disease.

Out of four dogs inoculated with blood from four koodoos,
three contracted the disease.

Two dogs were inoculated with blood from water-buel:, and
both remained healthy.

Two dogs were inoculated with blood from two impala, and
both remained healthy.

One dog inoculated with blood from a Burchell’s zebra re-
mained healthy.

One dog inoculated with blood from a bush-buck contracted
the disease.

Three dogs inoculated with blood from a stein-buck, wart-hog
and wild-pig, respectively, all remained healthy.

One dog inoculated with blood from a hyxna contracted the
disease.

The author concludes this section of the “ Report” with the
following remarks :—
“These experiments I think prove that several species of wild

DOGS FEEDING ON INFECTED TISSUES. 281

animals inhabiting the Fly Country harbour the Nagana parasite,
and the links in the chain of the causation of this disease con-
necting the game through the Tsetse-fly with the domestic
animals are complete.

“In regard to other points relating to the etiology of this
disease, such as seasonal prevalence, sex, age, breed, etc., I may
dismiss them in this Interim Report by the remark that, according
to my present knowledge, neither season, sex, age nor breed has
any predisposing or protective influence in this disease” (p. 24).

Pages 25 to 41 are devoted to a—“ Drscriprion OF THE FLY
Disrask orn NAGANA AS Iv OCCURS IN THE Domestic ANIMALS ”
(horses, donkeys, cattle, and dogs), with temperature charts,
showing “ fluctuations in the number of the blood corpuscles and
parasites,” and a “short description of the principal changes
found on post mortem examination.”

Fig, 14.
Dog suffering from Tsetse-fly disease, or Nagana
' (after Bruce).

Of the remaining portion of the “Report,” pp. 41 to 45 are
devoted to experiments on — “INOCULATION oF BLoop FROM
AFFECTED TO Hrauruy ANIMALS.” —

A.—“ INOCULATION OF BLoop IN WHICH THE Ha@matozoa
CAN BE DEMONSTRATED BY THE Microscope” (pp. 41-45).

B.—“ InocuLaTION oF BLOopD IN WHICH THE HaMATOZOA
CANNOT BE DEMONSTRATED BY THE Microscope” (p. 45).

On p. 46 is given a statement of an experiment on—
“Freping HeattHy ANIMALS ON ‘TissUES FROM ANIMALS
AFFECTED BY Nagana.” A native dog, after eating a piece of
coagulated blood from the heart of a heifer, Exp. 216, was found
to be suffering from the disease on the sixth day.

In connection with this experiment the author remarks :—

“Tn the Segane Valley, many of the dogs belonging to one of
the kraals fed on the raw flesh of a quagga, and I am informed
that several of these dogs subsequently died of Nagana.

“Mr. B. G, Lloyd also states that in the winter of 1892,
Mr. Saunderson, of Spitz Kop, Z.A.R., took some oxen into the
“ Fly” hunting. and lost them all, though one of them died after
returning to the farm. Some dogs got at the dead bullock and

282 ~ MEDICINAL. TREATMENT.

ate a lot of the meat raw. They all died within three months,
showing all the signs of “Fly,” whereas some pups which were
kept shut up were fed on the same meat, but cooked, for several
days, yet none of them showed any sign of being affected.”

Pp. 46 to 66 are devoted to “ Mupicinat TREATMENT.”

| © A.— ARSENIC AS A Curative AcEnt ” (pp. 47-64).

In connection with the experiments charts are given, showing
fluctuations in temperature, and in the number of blood
corpuscles and hematozoa, and the number of grains of arsenic
administered daily. ~

“oGi) Treatment in Horses” (pp. 47-56).

The author writes (p. 47) :—“Since I have been here at
Ubombo I have tried the arsenical treatment on several horses,
donkeys, and dogs, but in this Interim Report I would rather not
make any general statement concerning the usefulness or other-
wise of the drug, leaving rather the “yeader to draw his own
conclusions from the exper riments themselves. . . «

Fig. 15.
Horse suffering from Tsetse-fly disease, or Nagana
(after Bruce).

«The horses were given the arsenic in solution. The arsenic
was dissolved by heating in water with an equal quantity of
carbonate of soda, I made the solution in the following strength :—
Arsenic 6 grs., carbonate of soda 6 grs., and water 1 ounce.
The ounce of fluid was merely sprinkled night and morning over
his feed of crushed mealies, Xa) tat there was no trouble in the
administration of the drug.”

Since the subject of Tsetise-fly disease in horses is of general
interest, it may be useful, without going into the details of the
experiments, but as an indication of the results obtained, to
quote a few lines from the author’s “ Remarks” at the conclusion
of certain of the cases.

Exp. 205a.—(This horse eventually died of Horse Sickness.)

EFFECTS. OF ARSENIC IN HORSES. 283

“This horse remained alive sixty-five days from the com-
mencement of the disease, and then only succumbed to Horse
Sickness. While under arsenic he very fairly retained his
strength, as may be proved by the fact that he very often carried
me (riding over fourteen stone) down to the Fly Country and
back, when I went down to collect flies. This means being
on the march from 5 a.m. until 5 p.m., with a climb in the
afternoon of the Ubombo, which is some 2,000 feet above the

lain.”
, Exp. 224.—(This horse likewise eventually succumbed to
Horse Sickness.)

“Jt is evident that the arsenic had much less effect than in
the previous case (Exp. 205a).”

Exp. 264.—“ Horse, Bay.—‘ M.D.’”

** Remarks.—This case illustrates the uselessness of arsenic
as a means of preventing “ Fly Disease.” Although “M.D.” was
well saturated with arsenic and smelt strongly of it, two short
exposures to the “ Fly” were sufficient to set up the disease. The
first exposure was at Mkusi Drift, at the western end of the
Mkusi poort, a drift which has usually been considered free
from “Fly.” This year many Tsetse-flies have been seen there,
and several cases of Nagana have occurred among the animals
crossing the Segane Valley to Ubombo. At the eastern end of
the poort, and within a couple of miles of the drift, a herd of
buffaloes have their headquarters, so that it can readily be
understood how the flies find their way to the drift. As will be
seen from the temperature chart, the arsenic has held the disease
in check for some two months, but not sufficiently to prevent the
right eye from becoming affected by an opacity causing blindness.
The hematozoa are now beginning to reassert themselves, and
the case will probably end fatally in a few weeks.” [The horse
eventually became unable to stand, or to rise, and was therefore
shot.

ites. 278.—“ Another case showing the futility of arsenic as
a prophylactic agent.”

Exp. 283.—“In this case arsenic is seen to be practically
useless, both as a prophylactic and a curative agent.”

“ (2.) Treatment in Donkeys” (pp. 56-64).

“The donkeys were given arsenic in the dry condition, the
mouth being opened by an assistant and the powder placed on
the back of the tongue. There is no difficulty in administering
it in this way” (p. 57).

In the five cases described all the donkeys, except one,
eventually succumbed to the disease.

In “ Remarks” on Exp. 257 the author writes :—“ This case
shows in a marked manner the effect of arsenic in prolonging life
in this disease” (p. 61).

284 ARSENIC AS PROPHYLACTIC.

On Exp. 251 (the non-fatal case), in which the donkey,
having been placed on 12 grains of arsenic on February 13th,
after hematozoa had been discovered in its blood three days
earlier, was still alive on July 19th, in spite of several visits to
the “Fly Country” in the interval, the author’s “Remarks”
are as follows :—

“The preceding chart gives the principal facts of what is up
to the present the most successful case I have had of treatment
by arsenic. After five days of treatment the hematozoa are
seen to disappear from the blood and not to return for a period
of almost five months. The red blood corpuscles, which in
February lie between three and four millions per cubic milli-

Fig. 16.

Donkey suffering from Tsetse-fly disease, or Nagana
(after Bruce).

metre, are seen to gradually rise in numbers to five millions.
Corresponding with this there has been a gradual increase in
fitness, and at present the donkey has all the appearance of a
perfectly healthy animal. The arsenic was stopped on the
17th May, but the hematozoa reappeared on July 10th, when
the animal was again placed on arsenic” (p. 64).

“ B.— ARSENIC AS A PropuyLactic AGcrnt” (pp. 64-66).

The author commences this section with the following para-
graph :— .

«« As has been seen in the cases in which arsenic was given as
a curative agent, the drug undoubtedly markedly modified the
course of the disease. It, therefore, seemed probable that the
disease would be prevented altogether if the arsenic were given
for some time previous to the animal being exposed to the
infection. If twelve grains of arsenic given daily can cause the
hematozoa to disappear rapidly out of the blood, it is difficult
to imagine how they could appear in the first place if the animal
before entering the Fly Country had been given this quantity of
the drug for some time previously. But twelve grains daily is a

QUESTIONS NEEDING ELUCIDATION. = 285

very large dose of this powerful drug to give to a healthy horse,
and probably cannot be given in every case without setting up
inflammation of some of the internal organs, with falling away in
condition, and general debility ” (p. 64).

After giving details of five experiments made to determine
the point, the author sums them up with the following observa-
tion :— From these experiments. it may be concluded that
arsenic is quite useless as a prophylactic agent, but that it is
useful in prolonging life and usefulness in the Fly Country after
the disease has begun” (p. 66).

“10.—Concituping Remarks” (pp. 66-69).

After discussing the various ways in which an infective
disease may be attacked, the author goes on to say :-—

“To return to Nagana, what do we know of the disease ?
We know that it is caused by the entrance into the blood of a
minute parasite which multiplies there and causes death; that
this parasite exists in the blood of many animals and that it is
conveyed from animal to animal by the Tsetse-fly, or by the
eating of the raw flesh of animals affected by the disease. We
also know that the disease is limited to certain tracts of country
having certain physical features, but that its distribution in
these tracts is very variable.

“We do not know how the parasite causes death, but surmise
that it may do so by the poisonous action of some substance or
substances elaborated or secreted by it, or by producing a pro-
gressive hemolysis and anemia, leading to a form of auto-
intoxication. We do not know all the animals in which this
parasite may exist, but judging from the number of domestic
animals in which it is fatal, we may consider that it has a wide
range. It may exist not only in the koodoo, wildebeeste, and
other big game, but it may also exist in the wild cats, rats, birds,
and even fish of the Fly Country.

“We do not know whether there are other species of fly which
convey the disease besides the Tsetse, but this perhaps is a minor
consideration. We do not know of any method at present of
isolating the hypothetical poisonous substance elaborated by the
hematozoon so as to be able to study it or to render animals
immune to it by gradually increasing doses. No experiments
have, as far as I am aware, been made in this country in the
direction of studying the effect of the serum of animals in whose
blood the hematozoa have been numerous. Dr. Lingard has,
however, made a few with a negative result in Surra” (p. 67).

Remarking that treatment of disease by means of serum “is
very popular at present,” the author proceeds to quote in extenso
the results of Dr. Lingard’s experiments on :—‘‘ The subcutaneous
inoculation of blood and its derivatives in its different forms,

286 SUGGESTIONS FOR FUTURE. WORK,

obtained from animals suffering from, or which had lately
succumbed to, Surra” (pp. 67-68).

After this extract from Lingard, Col. Bruce observes :—

“Up to the present then the results of experiments with
serum have been negative, but I purpose if time permits to
continue this line of investigation.

* Another important point we are still ignorant of is whether
animals can become affected by this disease without the agency
of the Tsetse-fly or by eating raw: flesh, as, for example, by
drinking contaminated water or eating soiled herbage. This
point I am engaged in investigating. And, lastly, up to the
present we do not know of any drug which will prevent the
disease or cure it in every instance when incurred” (p. 68.)

The author concludes with the following “ suggestions ” as to
the lines on which future work in connection with Tsetse-fly
disease should be conducted :—

“J, That all trustworthy information regarding seasonal
prevalence be collected.

* * * * *

« According to the ‘ Natal Almanack,’ the dry season includes
the months of April, May, June, July and August; the wet
season September, October, November, December, January,
February and March. As far as I am aware, Nagana is not
confined to certain months of the year as Surra is, but I should
like to have more definite information on this point.

“9. Towards the end of the dry season or when the water
pools are nearly dried up, ‘can the Trypanosoma be discovered in
this concentrated water by the microscope or by feeding or
injection experiments ?

~“3, The question of herbage.—Dr. Lingard states that in
India after the rains have commenced the resting forms of the
organism which have been clinging to the grass bordering upon
the water-holes begin to soften, and when the animals feed on
such vegetation, repeated doses of the contagium are taken into
the stomach. It is probable that the small discs gradually gain
an entrance into the blood stream and-their development takes
place in the liver, spleen, and glands generally. . All grass-eating
animals, therefore, may be the subjects of Surra, so that the
opinions held with regard to the big game by the natives are
possibly founded on fact.

“In Africa we are, as yet, in dense ignorance respecting the
resting-stage of the Fly hematozoon; this would be a most
important point to work up. In regard to the herbage, will
such grass and water brought up from the Fly Country give
rise to the disease in healthy animals kept in a healthy locality ?

“4. It would be important to find out as far as possible what
species of wild animals, vultures, barbel, etc., harbour the parasites.

SUGGESTIONS FOR FUTURE WORK. 287

“5, As many examinations [as possible] of the blood and fluids
in the body-cavity of the Tsetse-flies should be made and the
result noted. Injections of fluids in which several flies have
been mashed up might also give results.

“6. How long do the hematozoa retain their vitality in the
body. of a dead animal ?

““7, How long do the hematozoa retain their vitality in
ordinary pond water 1

‘*8,. The Fly Disease in the native sheep and goat has to be
worked up. I have up to the present been unable to procure
these animals from the natives here.

“9, Further information is wanted regarding the breeding of
the Tsetse-fly.

“10. Are the young of animals affected by the ‘ Fly’ in any
way immune to the disease ?

“11. Sir Walter Hely-Hutchinson informs me that he has
heard it stated that animals can be rendered immune to Tsetse
by taking them into the Fly when young, at intervals. Is there
any truth in this statement ?

“12, Will the passage of the parasite through a series of
animals influence its morbific power ?

“13. Would large quantities of the serum of wild animals;
say the wildebeeste, have any influence on the course of the
disease if injected under the skin or into the peritoneal cavity of
the animal affected by the disease ?

“14, Can anyone study the disease in tamed wild animals,
for example the koodoo or any of the smaller buck? . . .

“15. Why should the wild animals be spared whilst tame
animals suffer? What is it in domesticity that removes immuhity !
Can the domestic animal be supplied with what is present in the
wild? Can this something be discovered !

** These, then, are a few of the problems to be solved in the
‘Fly’ question . ...” (pp. 68-69).

APPENDIX B.

ae

“A Nore on THE ‘Tse-Tse’ Furies oF THE British Crenrran
AFRICA Prorecrorate.” By Caprain RichHarp CrRAWSHAY.

The following ‘‘ Note,” which refers mainly to Glossina morsitans, Westw.,,
and Gl. pallidipes, Austen, was kindly written at the author’s request
by Captain Richard Crawshay, during a sojourn in London in June,
1896, on his return from Nyasaland.

“«Tse-Tse’ were formerly met with almost everywhere through-
out what is now known as the B. C. A. Protectorate ; but of late

288 CAPT. CRAWSHAY ON TSETSE IN NYASALAND.

years have decreased very much in the Eastern portion, or have even
become extinct in some localities, owing probably to the disappear-
ance of the larger game, such as elephants, rhinoceros, and buffalo.

“ During my earlier visits to this part of Africa, between 1883
and 1890, I encountered them practically wherever I went :—e.q,,
on the Lower Shiri River, numerous in the Elephant Marsh so
lately only as in 1891—the last time I was there ; on the Blantyre-
Matope Road, on the Lirangwi flats at Sambamwaliand Nachimvu;
very numerous opposite Matope on the Upper Shiri, where buffalo
were very plentiful; and wherever, almost, one touched on the
W. Coast of Lake Nyasa—notably at and around Deep Bay.

“ Nowhere in the eastern portion of the Protectorate do I
recollect having seen them very numerous, unless perhaps opposite
Matope ; nor have I experienced any great annoyance from their
bites, as compared with the torture one has undergone in the
western region. There, in 1891 and 1892, I found them a
terrible ordeal in the country lying between the 8.W. portion
of Lake Tanganyika (‘ Liemba’ of the natives) and Lake Mweru.
In Kabwiri they simply swarmed: I shall not soon forget the
Mofwi Valley and the Kalongwizi Flats !

‘“‘Many people are of opinion that ‘ Tse-Tse’ do not bite man,
or that if they do their bite is not painful, This theory is soon
dispelled in the Mweru country. There, marching at the head of
my caravan—as one does, to be ready for game—I have had as
many as forty or fifty ‘Tse-Tse’ at me at the same time. Life
was not worth living on the march: there was no getting a
moment’s peace: a large leafy branch had to be kept going
incessantly round one’s head, neck, and arms, and even then one
received many bites. ‘Tse-Tse’ are most active and vicious
assailants, and there is no limit to their perseverance ; they care
nothing for the flip of a fly-switch or a leafy branch, neither are
they easily felled to the ground with a slap of the hand, but fly
off apparently none the worse: a blow sufficient to fell an
ordinary blue-bottle will not fell the ‘ Tse-Tse.’

“Tt will convey some idea of how bold they are when I say
that they will creep in under one’s shirt, when open at the chest,
and are not disturbed by the garment touching them.

“ During the hottest hours of the day they are most aggressive,
but they bite at any time, even at night, when it is bright moon-
light. Their flight is powerful and noisy: the buzz is full-toned
and somewhat highly pitched :—not dull and droning like the
buzz of the ordinary horse-fly: they do not settle as slowly as
the horse-fly, but land with a bump, standing well up on their
legs. When a ‘Tse-Tse’ settles with the intention of feeding—
in the early mornings they usually simply settle on men’s backs
to sun themselves, away from the ground and vegetation wet
with the dew—he inserts his proboscis, lowers his head, and
raises his abdomen until it is almost vertical: when doing this,

TSETSE NEAR LAKE NYASA. 289

and for some little time after he has commenced sucking, he
works his wings, buzzing in a minor key, rather like a bee when
held forcibly, though not so powerfully : when the keenness of his
appetite has been somewhat appeased, he stops working his wings
and sucks in silence. If left to himself, he will suck until his
originally skinny barred abdomen becomes a large crimson bead.
He is then almost helpless: if touched he will not fly, and if
brushed off he will only go a yard or two, to settle heavily on a
bush, or gradually sink down to the ground.

“A common habit of ‘Tse-Tse’ is to sit on paths and game-
tracks, by day and on moonlight nights. They then fly up and
attack the first person who comes along, and follow him up in
an ever-increasing swarm, so that the leader of a caravan will
soon have forty or fifty about him; whereas the men behind will
hardly have one.

“Tn 1893-1895, I took ‘ Tse-Tse’ [Glossina morsitans, Westw.
—E. E. A.] in Henga—the highlands to the 8.W. of Deep Bay,
the altitude of which is some 3,300 feet. There are not many
there now: I doubt if one could take a dozen ina day. Those
I collected were most of them taken at the foot of Jakwa
Mountain, close to the junction of the Rukuru River with the
Lunyina; but they were to be found along the entire course of
the Lunyina River, even in the gorge through which it flows
into the lake, where there are still buffalo.

“The Ahenga, Ankamanga and Atumbuka know this insect
as ‘ Kasyembi.’ The Awemba, of Itawa and Kabwiri, know it as
‘Chisemberi’: pl. ‘ Visemberi.’

“The Arabs and Coastmen had a wholesome dread of the
‘Tse-Tse’ of the Mweru Country: I was warned what to expect
in Kabwiri by Abdallah bin Suleiman, of Kavuta, two or three
days before I encountered them in any numbers. In the Upper
Loangwa River Valley, some seven days’ journey south-west from
the north end of Lake Nyasa, ‘ Tse-Tse’ were plentiful in August
and September last year. I collected specimens which Mr. Austen
has determined to be Glossina morsitans, Westw. The altitude
of this part of the Loangwa Valley is 2,250 feet; there is a great
deal of game there, including elephants in the rains.

“In February, 1895, when visiting the recently established
Administration post at Kaporo, at the north end of Lake Nyasa,
T caught some very large ‘'Tse-Tse’* belonging to a species new
to me, much larger and darker than Gl. morsitans. These I
found sitting on the path at sunset: they did not bite. Two
specimens of these I sent home, a third I have by me.”

RicHARD CRAWSHAY.

Lonpbon,
June 17th, 1896.

* Glossina fusca, Walk., g.v.—lh. E. A.

290 TSETSE AND BUFFALO.

APPENDIX C.

Copizs:or LETTERS ON THE SUBJECT OF THE SUPPOSED CON-
NECTION BETWEEN THE TSETSE-AND THE Burrato (Bubalus
caffer, Sparrman): transmitted by the Foreign Office to the
British Museum (Natural History).

Mwenaa R.,
‘‘Hook of the Kafue,”
N.E. Rhodesia,
' 24th February, 1901.

Mr. Vat GteLtaup to-the SrcRETARY OF THE BrivisH Soutrn
AFRICA COMPANY.

Sir,—I see from the papers that the international con-
ference for the preservation of big game has decided to recom-
mend that Buffalo be placed on the list of game to be absolutely
protected. I, therefore, wish to bring to your notice my obser-
vations on the Tsetse-fly and the Buffalo, the correctness of
which are, I believe, borne out by the experience of Mr. George
Grey when travelling through a fly country in 1899 and also by
information obtained from native sources.

The Tsetse-fly has always abounded in districts where Buffalo
were numerous, and since the almost total destruction of Buttalo
by rinderpest these flies have not disappeared, although, perhaps,
not so numerous as formerly. The bite of the insect, however,
appears to have become much less deadly to domestic animals,
and stock and dogs not only survive the bites for a much longer
period than formerly, but in many cases suffer no ill effects
at all.

This has been my personal experience, and I think I am
correct in saying that Mr. Grey’s is similar.

The natives say that now the Buffalo are dead the Tsetse-fly
no longer kills stock, but of course natives are notoriously
careless and loose in their statements. It is a fact also that in
many places where Buffalo have become extinct the Tsetse-fly
has also vanished ; this is the case in parts of Sebungwi and
Zankie districts. As there are large fly districts in northern
Rhodesia I call your attention to these facts, as it appears to me
that the protection of the Buffalo and the Tsetse-fly are identical,
and I would doubt if the benefit accruing from the preservation
of the former will compensate for the disadvantages arising from
the existence of the latter.

I am, etce.,
VAL GIELGUD.

TSETSE-BELT IN BRITISH EAST AFRICA. 291

MomBASA,
6th September, 1901.

Str Caarues Extot, K.C.M.G., C.B., to the Marqugss or
LanspowngE, K.G., etc., etc.

My Lorp,—In:reply to your Lordship’s despatch, No. 259
of July 20th, respecting the connection between the Tsetse-fly
and the Buffalo, I have the honour to transmit letters from
Messrs. Stordy and McClellan and Doctor Radford.

After reading this correspondence and discussing the question
with other persons, my own opinion is that where there are
Buffaloes, Tsetse-flies are usually (but not always) found, but that
the flies also occur in districts where there are no Buffaloes.
Hence it would appear that the Buffalo cannot be the only host
of the parasite which the Tsetse-fly introduces into the blood of
domestic animals with fatal results.

I have, ete.
(Signed) C. Exior.

NAIROBI,
Hast Africa Protectorate,
3rd September, 1901.

R. J. Storpy, M.R.C.V.S., to Sir CHartes Enior, K.C.M.G.,
C.B., His Majesty’s Commissioner and Consul General,
Mombasa.

Sir,—I beg to acknowledge receipt of a copy of the
correspondence re Tsetse-fly and the preservation of the Buffalo
which you forwarded to me for my opinion, and I have the
honour to transmit to you herewith my observations on the
subject. ;

The Tsetse-fly belt of British East Africa, exclusive of Juba-
land, may be said to extend from Mtoto Andei to Simha, a
distance of roughly ninety miles; it is situated in a densely-
wooded, low-lying part of the country, about 3000 feet to 3400
feet above sea level.

Driajani, an old camping ground, within this area, was
considered by the late Captain Haslam and myself to be the
most dangerous place for fly on the old transport route, but
strange to say it was practically devoid of game of any kind.

In my opinion, Buffalo and other big game are not the only
factors in the Tsetse-fly theory, and we must first consider the
question of climate and humidity before we condemn the Bos
caffer as the true and only source of the Tsetse-fly disease.

I believe that the distribution of the fly is entirely influenced

U 2

292 TSETSE AND BUFFALO IN EAST AFRICA

by the physical ‘aspects of the country and that for its existence
it must have a humid, low-lying position.

Major Bruce in his excellent report (‘‘ Further Report on
Tsetse-fly Disease and Nagana in Zululand,” 1897, p. 20), says:
“That the presence of wild animals in the vicinity of horses and
oxen is not the only factor in the problem is shown by the fact
that in the old days when big game was numerous and roamed
over the whole country, hunters and travellers never complained
of fly until they encountered the disease in low-lying tracts of
country or along the large river valleys.”

As in the Hermansdorp district of Cape Colony herds of
Buffalo are still to be found, yet Tsetse-fly with its concomitant
disease is unknown, so in the high altitude of the Kedong
6000 feet), in this Protectorate, herds of Buffalo are to be met
with, greatly reduced in numbers by rinderpest within recent
years it is true, yet neither Tsetse-fly nor Tsetse-fly disease have
ever been known to occur, nor has the fly or its disease been
heard of in the Baringo district of the Uganda Protectorate,
where herds of Buffalo and other big game exist.

When studying the causes which rendered the Island of
Mombasa uninhabitable for horses, I ascertained that an
organism, the morphology of which was identical with that
found in animals suffering from Tsetse-fly disease, was found in
donkeys which had never left the island.

I expressed an opinion then (vide ‘‘ Preliminary Report as to
the causes which rendered the Island of Mombasa uninhabitable
for horses in 1899”) with regard to African Nagana and Indian
Surra being one and the same disease, and as the occurrence of
Surra cannot be attributed to the presence of wild animals or
Tsetse-fly, we must explain, ere we destroy the buffalo in an
attempt to stamp out Nagana, why a disease identical with that
caused by the bite of the Glossina morsitans occurs in places such
as Mombasa, where Tsetse-fly and buffalo are non-existent.

I have, etc., °
(Signed) Rosert J. Srorpy, M.R.C.V.S.
(Vety. Officer, E.A.P.).

NAIROBI,
East Africa Protectorate,

3rd September, 1901.
Dr. Raprorp to R. J. Srorpy, Esq., M.R.C.V.S.

Dear Mr. Storpy,—Many thanks for sending me the
communications you have received from Sir Charles Eliot, ve the
Buffalo and Tsetse-fly.

That the two should be associated is not extraordinary when

NOT CLOSELY ASSOCIATED. 293

one remembers that both inhabit densely-wooded, damp, secluded
districts, but the arguments advanced to prove that the Bos caffer
alone is the host of the Tsetse, and that the extermination of
the former leads to the disappearance of the latter do not appear
to me to be convincing. From my own observation I am inclined
to the opinion that hosts other than the one species mentioned
(or some other factors) are necessary for the propagation of the
fly, and that the hematozoon is in all probability to be found in
many species of diptera.

During my residence in Jubaland, East Africa Protectorate,
which extended over two years, I had ample opportunity of
studying the habits and distribution of the Tsetse-fly and the
effect it produced on domestic animals.

Speaking generally, the fly belt is confined to two distinct
areas in that province.

(1) The valley of the Juba River within the forest belt.
Here fly abounds for a distance of upwards of 400 miles—in
fact, so general is it in places that it is a source of annoyance to
Europeans and natives. Yet within the whole of that great
tract of country Buffalo are few and far between. The late
Mr. Jenner and I came across them in one place only (Lake
Galey and district). Within this belt is the district of Gosha,
80 to 100 miles in length, where the fly is peculiarly abundant,
yet Buffalo are not found ; notwithstanding this, the hematozoon
is very virulent, and on the occasion of the late Mr. Jenner’s
expedition to Lugh in 1899, he lost every camel and pack-ox
that went through (vide my Report, May 16th, 1899, forwarded
to the Secretary of State for Foreign Affairs, No. 53, May 31st,
1899). ‘

(2) The region of Lake Kumbi to the north-east of Desek
Wama (Lake Hardinge), where the distribution of the fly is
restricted to the dense forest belt in the neighbourhood ; yet
Buffalo are not known to frequent this district. It appears to
me reasonable to assume that where fly is abundantly found
extending over a large tract of country that its natural host (if
one only) should be in large numbers also; or else the fly must
be possessed of extraordinary migratory powers.

If that host be the Buffalo, it is strange that it is particularly
conspicuous by its absence in the extensive districts mentioned,
while waterbuck and bushbuck are common in most parts, and
yet again in others no animals of any sort are found save
monkeys and rodents. Amongst other pests that affect camels
very seriously in Jubaland is a species of gad-fly which is
restricted in its distribution to the open plains and sparsely-
bushed country in the districts of Desek Wama and Derib.
This fly, unlike the Tsetse, attacks animals during the day at all
hours, and the symptoms produced by it (in camels) are identical
in every particular with those produced by Tsetse, but I was

294 TSETSE AND BUFFALO ON JUBA RIVER.

unable at the time to substantiate this theory microscopically
owing to lack of the necessary materials, etc., for carrying on
investigations.
Yours truly,
Wiuiam 8. Raprorp,
Medical Officer, East Africa Protectorate. —

NAIROBI,
4th September, 1901.

Mr. J. W. P. McCLeian to Str Cuartes Exror, K.C.M.G., C.B.

Sir,—Referring to our conversation on the subject of
Tsetse-fly in the Jubaland Province, I would state that along
the Juba River where low-lying forest exists (my observations
cover a distance of some hundred and twenty miles from
Bulbula to Gele) Tsetse-tly abounds. The worst places are
damp, dark, and low-lying, shaded chiefly by the Ndoma Palm.
In many such localities there is little or no game and certainly
no Buffalo.

On the other hand, from Mtudo northwards Buffalo are
found, especially in rainy weather and in the heat of the day
frequent the thickest depth of the forest; here, too, Tsetse-fly
is found in large numbers, as also in the dry lake beds adjacent
to the river. At the same time half a mile or so away from the
river and lake beds in the dense dry bush, where Buffalo feed in
the early mornings and evenings, the fly does not appear. In
many places the areas in which fly exist are quite small, possibly
only a few hundred yards in extent, and I have seen a watering
place made entirely free of fly for the time being by cutting
down forest and undergrowth and burning the grass in the
immediate vicinity. Again, at Lake Hardinge (now dry), where
little or no forest exists, where Buffalo are frequently seen, and
always large herds of waterbuck, and Somalis graze their cattle
at all times of the year without ill effects. Passing on, however,
some thirty miles to the Rumbi forest on the Affmadu-road,
Tsetse-fly abounds, especially in wet weather.

My opinion, therefore, is that Tsetse-fly is to be found in
certain dark, damp, low-lying localities, irrespective of big game
of any kind.

T have, etc.,

(Signed) J. W. P. McCuettan.

FLY NOT DEPENDENT ON BUFFALO ALONE. 295

THE RESIDENCY, ZomBA,
British Central Africa Protectorate,

30th September, 1901.

ComMISSIONER SHARPE, C.B., to His Maszsty’s PrincrPan
SECRETARY OF STATE FOR FoREIGN AFFAIRS.

My Lorp,—With reference to your Lordship’s despatches
Nos. 141 and 155, enclosing copies of correspondence on the
subject of the connection between the existence of Tsetse-fly and
the preservation of Buffalo, I have the honour to append a few
notes which give my own experience during the past fourteen
years in Africa on this subject.

1. Tsetse-fly would appear to depend upon wild game for
their existence, as I have never found Tsetse in any locality
— game was totally non-existent.

. Tsetse does not in any way appear to be spedillly dependent
pene Buffalo. On the plains at the-north end of Nyasa,. before
rinderpest made its appearance, there were vast herds of Buffalo,
but no Tsetse. The natives at the north end at that time owned
large quantities of cattle, which could be seen grazing in close
proximity to Buffalo. When rinderpest came it killed practically
all the cattle and all the Buffalo. In other districts of British
Central Africa Tsetse are’ found in large quantities where Buffalo,
at the present day at any rate, do not exist.

3. Tsetse are not found (in British Central Africa) in open
plains, although such plains may have large quantities of game on
them, and in spite of the fact that at the edges of the plains, where
forest abounds, Tsetse are found. It would appear therefore that
what regulates the presence of Tsetse-fly is the deseription of the
country, almost as much as the abundance or npg ae! of game.

f have, ete.;
(Signed) Adamnep SHARPE,
His Majest y's Commissioner and Consul General.

Bivtuians UGANDA, -
27th September, 1901.

Deputy Commissioner Jackson, C.B., to Tue Marquess oF
Lanspowne, KG, ote etc.

My Lorp,—I have the ae to acknowledge the receipt
of your Lordship’s despatch No. 190 of July 20th, with enclosures
regarding the question of the existence of the Tsetse-fly in
connection with the preservation of the-Buffalo, and in reply to
submit the following remarks, in so far as my own experience
has taught me, on this vexed question.

296 CONDITIONS IN EAST AFRICA

I may say at once that I am firmly of the opinion that, in
East Africa, the existence of the Tsetse-fly was never in any way
connected with the presence of the Buffalo more than any other
species of game.

I first met with the true Tsetse in very great numbers, and
consequently suffered much from their needle-like bite, in German
East Africa, about 80 miles inland from Saadani in Feb. 1886.
At that time impala, hartebeest, zebras, and wart-hogs were
found in large numbers, also a few sable antelopes, but there
were no Buffaloes anywhere in the vicinity of my shooting
grounds. In 1887 I again found this fly in great numbers in a
small patch of thick bush, about a mile and a half long and
three-quarters of a mile wide, some ten miles west of Taveita.
In this bush, which projected from the forest, I certainly found
Buffaloes occasionally, but as a rule they preferred to lie up for
the day in the thick and cooler forest, in which there were no
Tsetse. The bush in question was a favourite resort of impala
and a small dik-dik (Madoqua)—the latter in great numbers—
and also a few bush-buck and water-buck. At that time (1887)
Buffaloes may be said to have swarmed in the vicinity of Taveita,
but I never saw a Tsetse-fly except in this one particular patch
of bush.

Later on, in 1889 and 1890, the fly was met with, also in
great numbers, along the old caravan road from about two miles
south of the Tsavo River as far as Kibwezi. Between these two
points there were practically no Buffalo, but a great number of
dik-dik and a few impala. The fly and the small game are still
there, but there are certainly no Buffaloes.

In 1891-92, after rinderpest had carried off nearly all of the
Buffaloes (at least 90 per cent.) throughout East Africa,
Mr. Rogers, the present Sub-Commissioner of the Tanaland
Province, and myself found the Tsetse-fly existing in considerable
numbers in a narrow belt of forest not more than a mile wide,
between Mkonumbi and Witu, and we were told by the natives
that the Gallas, when driving cattle to Lamu for sale, always
drove them through the forest by night, and that the herdsmen
carried smoking firebrands to keep the flies off. With the
exception of a few bush-buck and duyker there was no game in
the vicinity of this belt of forest.

These four places are the only areas, the first and third ones
alone being of any considerable extent, in which I have myself
met with the true Tsetse-fly, and yet, until they were decimated
by rinderpest, Buffaloes were more or less common throughout
East Africa; and perhaps in no part of the Continent were
they ever more plentiful than in the Masai Country between
Kilima Njaro and Lake Baringo, Mau Plateau and Turkwell.
Throughout the whole of this vast area the Tsetse was, and is,
non-existent.

AND ON SHORE OF LAKE VICTORIA. 297

I may add that there is a species of Tsetse-fly found along
the wooded portion of the lake shore here at Entebbe. A speci-
men of this fly I gave to Sir Harry Johnston, and I believe he
sent it home.* It is plentiful in the Botanical Gardens. In
these Gardens, with the exception of a few monkeys and squirrels,
and certain small nocturnal beasts, such as the ichneumons, etc.,
and an occasional hippopotamus, there are no mammals, and if,
as is supposed, the fly is necessarily dependent on the presence
of suitable mammals on which to feed, the blood of these animals,
and occasionally man, must necessarily form its food supply.

In conclusion, I may add that I have ventured to hold the
opinion that the Tsetse is, like the mosquito, only a blood-sucker
by predilection ; and, in support of this view, I may state that on
my return from Kibwezi in April, 1892, at a time when the whole
of the “fly-belt” was parched and dried up—there being no water
between Msogoleni and Tsavo River, a distance of 50 miles, and
consequently no game of any kind—the Tsetse was more plentiful
than at any other time, before or since, that I have passed
through that area. Between Mtoto-Ndai and Kinani I caught
on my own person thirteen of these flies, and my half-naked
porters suffered even more than I did from their bites. I can,
therefore, not readily believe that all these flies could exist in
such a dried-up, and, at that time, intensely hot locality, if solely
dependent upon the blood of a very infrequent passer-by or a
stray dik-dik.

I have, etc.,
(Signed) F. J. Jackson.

APPENDIX D.

Native Metuops or Prorecrinc ANIMALS FROM THE ATTACKS
or ‘TsETsE-FLIES. NATIVE PROPHYLACTICS AGAINST AND
RemMeEDIES FOR TsETSE-FLY Disease. Native NAMES FOR
TsETSE-FLIES.

Native Methods of The practice of crossing Fly-belts at night,
Protecting Animals With a view to the protection of oxen and
from the Attacks of horses, has been referred to in Chapter I,
Tsetse-fles. where it was shown not to be an infallible
safeguard although usually free from any great risk. According
to Livingstone [21] and Kirk [28], natives sometimes take
advantage of the remarkable dislike evinced by the Tsetse to

* As I have not seen this specimen, the particular species of Tsetse-fly
founds Entebbe, on Lake Victoria, must be a matter of conjecture.

298 NATIVE PROPHYLACTICS AND REMEDIES.

animal excreta, by plastering animals which have to pass through
a Fly-infested tract with a compound containing ordure and other
matters, which is stated to keep the Fly at a distance for the
time being. Subsequently, Livingstone was told [49] that the
Banyamwezi use lion’s fat, smeared on the tails of the oxen, for
a similar purpose with equal effect. In East Africa the Galla
herdsmen, when obliged to pass through a belt of forest infested
by Tsetse, light large fires and drive their cattle through the
smoke, which keeps off the flies [139]: according to Mr.
W. W. A. Fitzgerald [154], they also adopt the “additional
precaution of moving the animals at night, and Mr. F. J. Jackson
(Appendix C, p. 296) says that he was “ told by the natives that
the Gallas, when driving cattle to Lamu for sale, always drove
them through the forest by night, and that the herdsmen carried
smoking firebrands to keep the flies off.)

The specifics which come alia this heading,
eer a aaa as recorded by Various authors quoted in the
for Tsetse-fly Disease, Bibliography, are apparently nothing more

than the nostrums of witch-doctors. and
other impostors, and are probably unworthy: of serious notice.
In some cases plants or roots, which have not been identified, are
said to be used as a remedy [26, 28, 37]; in others the Fly itself
in varying quantities, and sometimes. with the addition of. the
bark of a certain root, is administered internally |27, 30]. Thus
W. M. Kerr [97], writing of the country near Chibinga, Mashona-
land, where Tsetse abounded in July, 1884, says: ‘The native
women dry quantities of the Tsetse-fly and pulverise it with the
bark of a root, and mixing it with water give it to the young
animals, such as dogs and goats or sheep, of which they have
very few, seemingly only kept as pets.” This treatment seems
at any rate to have been believed in by the pioneer-explorer
Carl Mauch [50], who writes: ‘Only one remedy appears to be
effective, and that is based upon homeopathic principles : the Fly
itself, taken internally, makes the. punctures innocuous, as I have
seen in the case of a dog, which after administering this remedy
I took with me as far as the Lower Zambesi and sent back. again
perfectly well with those who had accompanied me.” It may be
mentioned that, according to Chapman [30], a peculiar breed of
dogs known as’ Makoba, bred inthe Fly-country ‘from time
immemorial, ” escapes Tsetse-fly disease ; while Andersson [20]
asserts that, “a dog reared on the meat of game, may be hunted
in Tsetse districts in safety. me

be? Colonel Bruce, however, found that a native dog contracted Tsetse-
fly disease after eating a piece of coagulated blood from the heart of a
heifer which had died from ‘it. He also mentions ‘an instance in which
dogs contracted and died from Nagana after eating the flesh of a zebra;
as also another case in which a similar fate was stated to have overtaken
dogs which had fed on the raw flesh of a bullock, which had perished from
the disease (Cf. Appendix A, pp. 281-282).

NATIVE NAMES FOR TSETSE-FLIES. 299

Livingstone’s assertion that calves do not contract Tsetse-fly

disease so long as they are being suckled by their mothers was
doubtless based on the statements of natives ; while the native
belief in “homeopathic” prophylactics or remedies is further
illustrated by Baines [55], who writes: “All young animals
while living on milk are safe. Some of the tribes living on the
borders drive the calves into the fly during the day, and bring
them out to be suckled morning and night. This is supposed
also to render them secure during the rest of their lives.” Similar
precautions are said to be adopted by the Bushmen in the case
of dogs. According to Bradshaw [77]: “Dogs are taken into
the fiy district by the Bushmen when pups, and are allowed to
suckle the mother, and at the same time given as much fly to
eat as can be captured for them. The mother in this case dies,
and the pups live and grow up, but are the most miserable
looking objects I ever saw belonging to the canine race—
emaciated, small, and the hair standing up all over them.”
The following list of native names is of course
very far from complete, but it is printed here
in the hope that it may prove of some slight
use to investigators.

Native Names for
Tsetse-flies.

The Tsetse is called kipanga (meaning “little sword’) near Kilwa.—

CaptTain R. F. Burron [25].

> » panga (or ‘‘sword’’-fly) in Unyamwezi—H. M.
STANLEY [45].

‘ss 55 chipinga on the Rovuma River.—Dr. Krrx [28].

HS [3 chufwa by the Wasawahili, near Rosako, in
Ukwere, about 20 miles inland from Bagamoyo.
—H. M. Sranuey [45].

= Pe burgo in the Manyuema country.—Dr. Livinc-
STONE [49].

the af ganda by the Wa-Galla.—W. W. A. FirzGERaLp
[154].

3 “9 mau in the country of the Rhol Tribe, north-west
of Gondokoro.—MRr. and Mrs. PerHerick [37].

a a kasyembi, by the Ahenga, Ankamanga, and Atum-
buka in British Central Africa,—Caprain CrAw-
sHay (Appendix B).

os 8 chisemberi (pl. visemberi) by the Awemba of Itawa
and Kabwiri, British Central Africa—Caprain
CRAwsHAY (Appendix B).

ae rf impugan on the Revyui River, Mashonaland.—F. C.
SELOUS [121].

5 se adjoé in Togo.—Dr. ScHILLING [170].

300 DISTRIBUTION OF NAGANA.

APPENDIX E.

THE GuoGRAPHICAL DisTRIBUTION OF NAGANA, AND THE PossIBLE
TRANSMISSION OF THE PARASITE OF THE DisEASE BY SPECIES OF
Glossina OTHER THAN Gl. morsitans, AND BY BLoop-SucKING
FLIES OTHER THAN TSETSE.*

In spite of the apparently over-hasty conclusions of Koch and
other German investigators, the question as to the identity of
Nagana with the disease known in India as Surra is not yet by
any means finally decided. This, however, is beyond the scope
of the present work, which is confined to Africa.

Apart from the region to the south of Lakes Tanganyika
and Nyasa, the chief foci of Tsetse-fly disease at present definitely
known are situated in German East Africa, British East Africa,
and Somaliland on the east, and in the German Protectorate of
Togo on the west. In the opinion of MM. Laveran and Mesnil
[XXI.], Nagana probably occurs wherever Glossina morsitans or
one of its congeners exists, but information at present available
does not altogether warrant such a conclusion. Thus Stuhl-
mann [174] records the capture in January, 1902, of specimens
of “Glossina tabaniformis, Westw.” (Gl. fusca, Walk.), near
Dar-es-Salim, at a place where the same cattle and goats had
grazed for a long time without the disease having appeared
among them. On this account, and—“ Reasoning from general
conclusions derived from analogy in connection with the diseases
due to infection of the blood, in which each parasite has its own
particular host and intermediate host ”—-Dr. Stuhlmann thinks
that it may be assumed that Tsetse-fly disease “is not conveyed
by Glossina tabaniformis.” In this case we are given no informa-
tion as to the presence or absence of game in the vicinity, but
what appears to have been the same species of Tsetse was met
with in December, 1895, by Mr. A. H. Neumann [152], on the
eastern shore of Lake Rudolph, where game was abundant.
Mr. Neumann writes: ‘“ Whether this kind is poisonous or not I
am not sure. My donkeys never suffered from having passed
here ; but then they can stand a few ‘fly,’ though where the
‘Tetse’ is numerous they soon succumb.” Turning to another
species of Tsetse-fly, in September, 1899, at Lumley, a few miles
from Free Town, Sierra Leone, I myself saw a herd of cattle
being pastured on grass-land in close proximity to a mangrove
swamp where Glossina palpalis, Rob.-Desv., was present in some

* The statements in this section are by no means to be taken as
constituting the last word on the subjects referred to; they are intended
rather to suggest lines for future investigation, by drawing attention to
the paucity of our present knowledge.

CONVEYANCE OF. PARASITE. 301

numbers.* The flies were not actually observed attacking the
cattle, but they probably did so since they bit my two com-
panions and myself when seated on the seashore hard by. It
is but fair to mention that, with the exception of a few small
buck, there are no wild animals of any size near Free Town
which might afford sustenance to the Tsetse and at the same
time harbour the Trypanosome. But this is not the case in
Uganda, where, according to Sir Harry Johnston [172], “ nearly
every known type of African antelope is represented,” not to
mention buffalo, giraffes, and zebras, while Glossina (species as
yet undetermined) also occurs there, although Tsetse-fly disease
is absent. With regard to Uganda, Sir Harry Johnston suggests
that “either the true Tse-tse is absent from all parts of the
Protectorate, or it is unable to obtain there the germs of fever
which it is the agent in introducing to the blood of horses,
cattle, and other beasts.” By “the true Tse-tse,” Sir Harry
Johnston doubtless means Glossina morsitans, Westw., and if so
the first of his suggestions is in accordance with the view of
Dr. Stuhlmann, as expressed in a recent paper [XXVI.], which
shortly stated is that GJ. morsitans is the only species capable of
conveying Tsetse-fly disease. From a quotation already given
the reader will understand that Dr. Stuhlmann bases his conclu-
sion upon the analogy between Tsetse-fly disease and malaria
and Texas fever (cf. p. 300). Elsewhere he writes [XXVI.]:
“The fact is that, in the case of all these blood diseases, the host
and intermediate host are animals of perfectly definite kinds.
It is, moreover, improbable that the proboscis of the Tsetse
merely operates as an inoculating needle; rather must we
assume, from analogy with malaria, that within the fly the
Trypanosoma passes through a special stage of development,
which it is true is at present unknown.” In this conclusion
Stuhlmann is supported by Veterinary-Surgeon Schmidt, in a
memorandum which is quoted on p. 263. So far as I am aware,
the only modes of reproduction which have yet been observed in
any Trypanosome are either binary longitudinal fission, or the
breaking off from a plasmodium, formed by the fusion of a
number of adults, of amceboid forms which subsequently assume
the well-known adult stage. Nevertheless, although nothing
has yet been observed in the case of Trypanosoma in any way
comparable to the gametocytes in that of the parasites of
malarial fevers in man, it is of course quite possible that on
entering the stomach of the Tsetse the parasites become sexually
mature, and that it is their progeny and not they themselves
that subsequently find their way into the blood of another

* Somewhat similar conditions would appear to exist in Loango,
or at least to have existed there at the time of the German Loango
Expedition about thirty years ago (Cf. 65, 65a, 80, and Chapter IL1.,
p. 40).

302 CONVEYANCE OF .PARASITE.,

mammal.* In this case the Tsetse would be a true host + of the
parasite, and- not the mere “inoculating needle” which Colonel
Bruce’s investigations would lead us to suppose. Colonel Bruce
found that the Tsetse is capable of conveying the disease forty-
eight hours after feeding on an animal suffering from Nagana,t}
which would certainly allow plenty of time for the Trypanosome
to make its way back from the fly’s stomach into the proboscis,
assuming no process of reproduction to take place. If this is
what happens, there would seem to be no reason whatever why
one species of Glossina should not be capable of conveying the
parasite of Nagana as well as another, since Dr. Hansen’s
investigations § tend to show that the proboscis of each species is
identical in structure. On the other hand, if the Tsetse is a
true host of the parasite, the latter may still be conveyed, if not
by all the species of Glossina, at any rate by more than one
species, just’ as the parasite of zstivo-autumnal fever in man is
stated to be conveyed by several species of mosquitoes of the
genus Anopheles. According to Laveran and Mesnil [171], who
quote from Blanchard, the deaths of the camels, mules, and
donkeys, of an expedition at Imi, on the Webi Shebeli, Somali-
land, as observed by Brumpt, did not appear to be due to
Glossina morsitans, but to a closely allied species. In view of
the locality, this was doubtless Gl. longipennis, Corti. Never-
theless, it must be admitted that there is yet no definite
evidence of the occurrence of Tsetse-fly disease in a locality
where Glossina morsitans is not found, because our know-
ledge of the distribution of this species of Tsetse, as of that of
all the other species, is still far from complete; and the
unexpected discovery that Gl. morsitans occurs in Togoland,]||
where Tsetse-fly disease is known to exist, is sufficient to show
that the species extends much further from the Limpopo and the
Zambesi than has hitherto been supposed. We must, therefore,
be careful not to assume too hastily that GJ. morsitans is absent
from a country where Nagana is known to occur, simply because
no specimens have yet been collected there. At the same time,
without attempting to draw deductions from it, it seems advis-

* Colonel Bruce’s observations, however, so far as they go, would seem
to negative this, since they show that the hematozoa are still active and
apparently unaltered in what remains of the blood in the stomach of the
fly nearly five days after its meal. Bruce writes: “ Immediately after
feeding, the tube of the proboscis can be seen to be crammed full of red
blood corpuscles, among which the hematozoa can be seen actively
wriggling. Up to 46 hours after feeding I have seen living hematozoa
and red blood corpuscles in the proboscis. After 118 hours the hematozoa
are still very numerous and vigorously active in what remains of the
blood in the stomach. After 140 hours the stomach is empty.” (Cf.
Appendix A, p. 278.)

+ The definitive, not the intermediate host as stated by German writers.
Cf. p. 260, note *,

+ See p. 275. § Cf. Chapter V. || See Chapter TV., p. 85.

BELIEF .IN..NOXIOUS..FLIES \IN..AFRICA. 303

able once more:to call ‘attention to the fact that, while a disease
that appears to be Nagana undoubtedly exists in Somaliland
(on the Webi Shebeli,:in. the; Korayo - Valley, the, Aulihan
country and elsewhere),*. with fatal effects to horses, camels,
donkeys, and. mules, all the: Tsetse-flies yet received from that
country prove to belong to Glossina longipennis, Corti. On the
other hand, the experience of Mr- W. A. Eckersley [134] near
Beira, in 1893,t lends force to the second of the above quoted
hypotheses of Sir Harry Johnston to account. for the absence of
Tsetse-fly disease from the Uganda Protectorate, and goes to
support the suggestion at the end of Chapter I. that systematic
efforts should be made to study the geographical. distribution of
the parasite of Nagana, as existing in the blood of wild animals
in Africa, by means of Colonel Bruce’s method of injecting blood
from the latter into domestic animals.}

As to the question whether the parasite of Nagana can be
conveyed by blood-sucking flies other than the Tsetse, it may be
remarked at the outset that were this the case to any large extent
the disease would probably be of much more general occurrence.
Nevertheless, a certain amount of evidence undoubtedly exists,
which; if reliable, can only be interpreted in one of three ways :
either the Tsetse occurs in localities where it has not yet been
identified, and the deaths of domestic animals ascribed by natives
to the agency of other and unknown flies are really due to it ; or
blood-sucking flies other than Glossina may in certain localities,
and perhaps only at certain seasons, convey a hematozoon which
is either that of Nagana or an organism closely related to it ; or,
lastly, the disease in these cases, whatever it may be, is quite dis-
tinct from Nagana, and is possibly, like Texas fever, disseminated
by means of ticks. According to Marno [47], the assertion that
‘“‘in certain parts of Africa at certain seasons domestic animals
are killed by the poisonous bites of flies, which in some.countries
even make the keeping of particular domestic animals impossible,”
has been constantly repeated since the time of Agatharchides.§

* Cf. [94, 124, 133, 150, 171).

+ A survey party to which Mr. Eckersley was attached took two Natal
ponies through the Fly-belt between Beira and Chimoio without any
ill-effects to the animals, although Tsetse-flies (in this case doubtless
Gl. morsitans) settled on them in considerable numbers.

+ Cf. Chapter I., p. 80, note ¢, and Appendix A, p. 280. According to
Mr. R. J. Stordy (see Appendix C, p. 292), a parasite ‘‘the morphology of
which was identical with that found in animals suffering from Tsetse-fly
disease ’’ occurs in Mombasa in donkeys, which have never left the island,
although the Tsetse is said not to be found there.

§ ‘‘Acatharchides (Ayafapxiins) or Agartharcus (’Ayd@apxos), a Greek
grammarian, born at Cnidos, lived at Alexandria, probably about B.c. 130.
He wrote a considerable number of geographical and historical works; but
we have only an epitome of a portion of his work on the Erythraean Sea,
which was made by Photius. It is printed in Hudson’s Geogr. Script. Gr.
Minores.’—Smith's Classical Dictionary.

304 STOMOXYS AND NAGANA.

Possibly some of these statements refer to the Fayfim, in Upper
Egypt, where, as I am informed by my colleague, Dr. C. W.
Andrews, the Arab camel-drivers declare there is a fly in the
months of May and June which kills camels ;* so convinced are
the drivers of this, that they will not allow their camels to visit
the district during the dangerous season. If this fly has any
existence in fact, it seems scarcely possible that it can be a
Tsetse, since the Fayiim, which lies approximately between 29°
and 29° 30’ N. latitude, is far removed from the nearest known
habitat of Glossina. It is more probable that the fly, if it
exists at ail, is either a Stomoxys or Lyperosia, or else a horse-fly
(Tabanus, Hematopota, or Pangonia) ; and in connection with the
first-mentioned genus, it may be remarked that the British
Museum collection contains a series of specimens of a species
of Stomoxys, which were brought home by Dr. J. W. Gregory
from Ngatana, British East Africa, where he stated that they
killed his camels. Moreover, a fly that is obviously a species of
Stomoxys was declared by Mr, F. J. Jackson [119] to be ‘“ very
plentiful” in the forest on the banks of the River Lumi, at Taveta,
British East Africa, where it “ worries beasts to such an extent
that they are unable to eat, and die.” In this case, as has
already been pointed out (see Chapter VI., Bibliography, p. 211),
the mortality may really be due to a Trypanosome. ‘The late
Captain A. G. Haslam, at Machakos, in June, 1898, found the
living Trypanosome of Tsetse-fly disease in the posterior part of
the abdomen of specimens of two species of Stomoxys caught
sucking the blood of mules suffering from the disease ; and in a
letter dated July 3, 1898, which he sent to the author with some
of the insects, he stated that he had ‘caught these flies on every
kind of animal, including gazelles, wildebeeste, and all domestic
animals, and also on meat exposed for a few seconds.” Captain
Haslam went on to say: “ Animals do not object to them much
after the preliminary stab with the proboscis. They remain
sucking for several minutes, and bulge out their abdomens with
blood. I am now engaged in experiments to show whether these
flies I send you actually inoculate other animals with the Tsetse
disease.” The lamented death of the writer, which took place
but a fortnight after the date of this letter,t prevented these
experiments from being completed ; but it may be pointed out
that any blood-sucking fly feeding upon the blood of an animal
suffering from Nagana, in which the parasites were numerous,

* This insect, whatever it may be, must not be confused with the
camel bot-fly (Cephalomyia maculata, Wied.—Family CMistride), the
larvee of which are very frequently met with in the nostrils of camels,
but do not appear to be injurious to the animals. When mature
they are sneezed out by the camels and pupate in the sand. (For a
figure of the fly, see Sharp, Cambridge Natural History, Insects, Part II.,
p. 515). :

+ See page 100, note.

WANDOROBO- OR DONDEROBO-FLY. 305

would naturally suck up some of the hematozoa with the blood,
and that the presence of living parasites in the stomach of the
fly by no means proves that the latter is capable of communi-
cating the disease. On the other hand, it the parasites remain
alive in the alimentary canal of Stomoxys sufficiently long, it is
possible that some of them might make their way back into the
proboscis, as Colonel Bruce supposes them to do in the case of
the Tsetse, and so be subsequently introduced into the blood of
another animal. Or one might suppose Stomoxys to be capable
of conveying the parasites directly from an infected to a healthy
animal, by flying to and biting the latter immediately after
sucking blood from the former, just as Rogers [XIX.] in India
found that the Trypanosome of Surra could be conveyed by horse-
flies. In view of Rogers’ results, it is certainly curious that no
instance of a spontaneous outbreak of Nagana, due to the direct
transference of the parasite from diseased to healthy animals by
flies other than Tsetse, occurred in the course of Colonel Bruce’s
experiments on the Ubombo Mountain in Zululand.* This, of
course, does not prove that such cases may not sometimes occur
under natural conditions, when in the same locality blood-sucking
flies other than Tsetse and the parasites of the disease in the
blood of wild animals are both numerous.

Turning to the horse-flies (Tabanide), we find a certain
amount of evidence from East Africa tending to show that if
present in sufficient numbers their bites may be fatal to domestic
animals. The so-called “ Wandorobo ”- ‘ Donderobo ”- or ‘* Ndo-
robo ”-flyt of the Wadschagga and Masai is as yet unidentified,

_ ™ Cf. Appendix A, pp. 273-274. Colonel Bruce’s experience is in
accordance with Kirk’s statement [28] that ‘‘no danger” has been found
to result from cattle bitten by Tsetse ‘‘ mixing with others.”

+ Cf. Oscar Baumann, ‘‘Durch Massailand Zur Nilquelle” (Berlin:
Dietrich Reimer, 1894), p. 28—['Translation] ‘‘The loss in personnel in
Umbugwe made itself felt, besides which our baggage-donkeys had
suffered from the sting (Stich) of the Ndorobo-fly. This insect frequents
water-courses and is dangerous to donkeys, since it stings them in the
anus, which results in swellings and death.’ (This refers to the region
near Lake Manyara, W.S.W. of Kilima-Njaro.) Cf. also Otto Kersten,
‘‘ Baron Carl Claus von der Decken’s Reisen in Ost-Afrika in den Jahren
1859 bis 1861” (Leipzig und Heidelberg: C. F. Winter’sche Verlagshand-
lung), Band I. (1869), p. 249, Band II. (1871), p. 36. The region referred to
in the following passage (Bd. I., p. 249) is the Wateita Country, inland
from Mombasa. [Translation] ‘‘ We had rescued the donkeys from the
robbers’ hands, but only to see them die after all the same day: in the
case of three of them, blood and matter escaped from the nose, the head
and genital organs were swollen, and the air-passages were so narrowed by
ulcers that their breathing resembled a rattling—they were stung by
Donderobo flies! I myself have never caught sight of the poisonous fly,
and have not been able to get possession of any in spite of the offer of a
reward; but the natives declared that it was only the sting of the
Donderobo, which was frequently dangerous to their flocks of goats, that
produced such effects. Just as Tsetse-flies attack cattle, so do the
Donderobo flies chiefly attack donkeys and goats, more rarely sheep, but
never cows. On the third day the animal that has been stung is already

x

206 HORSE-FLIES AND CAMELS IN JUBALAND.

and if such an insect really exists, may eventually prove to
be a Tsetse ; but it is considered by Stuhlmann [XXVI.] to be
‘“‘more probably a horse-fly.”* Schoeller [167], however, whose
remarks on the question are worth referring to, considers that
the disease which seems chiefly to attack baggage-donkeys “ in the
steppe regions of East Africa,” and is attributed by the Wadschagga
to the bite of an unknown fly which they call “‘ Wandorobo,” is
really a malady sui generis, allied to the horse-sickess of South
Africa (‘‘ Dikkopziekte” of the Boers). Be this as it may, Dr.
W.S. Radford, writing from Nairobi, East Africa Protectorate,
on September 3rd, 1901, states that in Jubaland symptoms
identical with those of Tsetse-fly disease are produced in camels
by the bite of a horse-fly. Dr. Radford says: ‘“ Amongst
other pests that affect camels very seriously in Jubaland is a
species of gad-fly [i.e. horse-fly] which is restricted in its distribu-
tion to the open plains and sparsely-bushed country in the
districts of Desek Wama and Derib. This fly, unlike the Tsetse,
attacks animals during the day at all hours, and the symptoms
produced by it (in camels) are identical in every particular
with those produced by Tsetse, but I was unable at the time to
substantiate this theory microscopically, owing to lack of the
necessary materials, etc., for carrying on investigations.”
Although this is a mere statement, unsupported by any evidence
of scientific value, the fact that the fly in question is confined
‘to the open plains and sparsely-bushed country ” would seem to
preclude the possibility of its being a Tsetse, even one of the
large species (Glossina fusca, or Gl. longipennis), and the effort
should be made to identify and experiment with the fly without
delay, in addition to making a microscopical investigation of the

incapable of movement, and it is only seldom that it survives the fifth
‘day.” The country referred to in the next passage (Bd. II., p. 36), is that
just south of Kilima-Njaro.—[Translation] ‘‘Even before the start we
lost one of the three donkeys that remained to us, without doubt in
consequence of the sting of the Donderobo fly (see Band I, p. 249). The
genital organs and the inguinal glands first showed a considerable
swelling; this then spread further and finally made any movement
impossible; blood and matter flowed from the nose; nay, even from the
orbital cavities, and an entire loss of strength indicated the near approach
of the end of the animal.” The statement that ‘‘the Donderobo flies
chiefly attack donkeys and goats, more rarely sheep, but never cows,” is
curious. Some of the symptoms mentioned (swollen head and genitalia)
would seem to suggest that the ‘‘ Donderobo fly” is none other than the
Tsetse; but the blocking of the air-passages by ulcers is noteworthy, and
death on the fifth day appears too speedy to be the result of Tsetse bite.
On the whole, Schoeller’s view [167] that the disease is not due to the bites
of flies at all, but is a malady allied to the South African horse-sickness,
seoms reasonable.

* According to Otto Kersten [43], on the other hand, nothing is heard
of the attacks of horse-flies in the region between Mombasa and Dschagga,
although ‘“ the Donderobo-fly, which is dangerous to donkeys,” is alleged
by natives to occur there.

+ Cf. Appendix C, p. 293.

DOOG AND BALAAD ON WEBI SHEBELI. 307

blood of the diseased camels. Dr. Radford expresses the opinion
that the hematozoon of Tsetse-fly disease ‘‘is in all probability to
be found in many species of Diptera,” but he advances no further
arguments in support of this view, which, in so far as it refers to
the normal conveyance of the parasite, is, as already pointed out,
@ priort improbable. With regard to the alleged effects of the
bites of Tabanide upon camels, however, there is some further
evidence, though in this case it is quite possible that the real
culprit is Glossina longipennis, which is of about the same length
as the smaller fly referred to. Colonel Swayne, writing to the
Secretary of the Zoological Society in 1894, said: “I send you
three specimens of ‘ Doog’ (a large fly) and three specimens of
‘Balaad’ (a small fly), . . . I was very much pestered by ‘ Doog’
on my way through Ogaden to the Webbe Shebeyli, in Somaliland.
They swarmed on my camels, constantly drawing blood. The
other fly, ‘ Balaad,’ which looks not unlike the common house-fly,
is far the worst fly on the Webbe; a valuable camel, on which I
eaught three or four two months. ago is now dying, and the
Somalis say that this is due to the bites of ‘ Balaad.’ If there are
many of them they kill horses and camels, and the Somalis will
not have their live stock grazing where ‘ Doog’ and ‘ Balaad ’ are
found.” * On the other hand, the conclusion which might be
drawn from Colonel Swayne’s statements is traversed by others
contained in a Report (dated May 26, 1891) from the officer in
charge of what was then the Imperial British East Africa Com-
pany’s Station at Lamu. According to this document, which
accompanied specimens of Tsetse-flies and Tabanide sent to the
British Museum for determination, the bites of horse-flies inflicted
upon domestic animals do not cause ‘any kind of blood-poisoning
like those of the Tsetse,” although it is stated that some of the
natives declare that they do. The Report observes that the bite
of the Tabanide “is extremely sharp, and, though causing
comparatively little irritation, produces much swelling of the
surrounding parts.” The writer adds: “ Both cattle and donkeys
bleed a good deal from the puncture, and the Germans tell me
that in some cases their cattle bled to death, it being impossible
to stop the bleeding, although they tried acids, etc.” The latter
statement suggests the possibility that, in some cases at any
rate, animals said to have been killed by the bites of flies other
than Tsetse, may really have died from hemorrhage caused by

* Two species seem here to be distinguished by the Somalis, but the
specimens sent by Colonel Swayne included examples of three: of the
three flies representing ‘‘Doog,” two proved to belong to a new species
subsequently described by the author under the name Pangonia tricolor
(nom. preocc. = P. beckeri, Bezzi), while the third specimen belonged to
another species of Pangonia, but was too much damaged for. determination.
The three specimens representing ‘‘Balaad’’ were examples of a small
Tabanid, somewhat resembling Hematopota in form, but with clear wings ;
these specimens were also too much damaged to determine.

a 4

308 SEROOT FLIES IN SUDAN.

the bites—such bleeding as, for instance, according to Sir H. M.
Stanley [45], results from the bites of a large horse-fly, called
Mabunga by the natives, near Rosako, in Ukwere, about twenty
miles inland from Bagamoyo. “This fly,’ writes Stanley,
“along with a score of others, attacked my grey horse, and
bit it so severely in the legs that they appeared as if bathed
in blood.” * The explorer’s men “unanimously stated that its
bite was fatal to horses as well as to donkeys.” In “ The Nile
Tributaries of Abyssinia” (London: Macmillan and Co., 1867),
the late Sir Samuel W. Baker, writing of the country between
the Settite and Atbara Rivers, describes and figures a ‘“ Seroot
fly ” (pp. 184-185), which he states (p. 184) is the species “ that
drives the camels from the country”; from the figure and
description it is evident that this insect is a horse-fly belonging
to the genus Pangonia, and not a Glossina. Further on (p. 196),
a remarkable description is given of an instance in which swarms
of Seroot flies rendered it impossible to take possession of a
giraffe which had been shot. Sir Samuel Baker writes: ‘The
Seroot-fly was in swarms about the carcase, thousands were
buzzing about our ears and biting like bull-dogs ; the blood was
streaming from our necks, and, as I wore no sleeves, my naked
arms suffered terribly. I never saw such an extraordinary
sight ; although we had killed our giraffe, we could not take
possession ; it was no wonder that camels and all domestic
animals were killed by this horrible plague, the only wonder was
the possibility of wild animals resisting the attack. The long
tails of the giraffes are admirable fly-whippers, but they would
be of little service against such a determined and blood-thirsty
enemy as the Seroot.” Subsequently (p. 210) the author men-
tions herds of game as retreating from the south before the
attacks of the Seroot. On the Blue and White Niles several
species of horse-flies, belonging to the genera Tabanus and
Pangonia, are known to Europeans under the comprehensive
name Seroot ; all are greedy blood-suckers, but it would lead us
too far from the question at issue to pursue the subject further.
It is hoped, however, that enough has been said to demonstrate
the advisability of determining once for all, by a series of ‘practi-
cal experiments, whether the various African species of Tabanide,
as also those near allies of the Tsetse belonging to Siomowys or
Lyperosia can, under any circumstances, become the carriers of
the Trypanosome of Nagana, or possibly of some other closely
related hematozoon.

* There igs no need, however, to go to Central Africa in order to see
horses’ legs streaming with blood from the attacks of Tabanide, since
this can be observed in Switzerland and Italy. That African cattle can
withstand the loss of a considerable quantity of blood without succumbing
is proved by the well-known habit of the Masai of bleeding their cattle to
the point of faintness for the sake of drinking the blood; in this case the
cattle recover after bleeding has terminated.

ORIGIN OF WORD “TSE-TSE.” 309

APPENDIX F.

‘© A Nove ON THE ORIGIN OF THE WorRD ‘TsE-Tsz.’” By Caprain
RicHarD CRAWSHAY.*

“The word ‘Tse-Tse’ at once indicates Bantu origin. It is
a good example of the remarkable onomatopeia abounding in
Bantu languages, especially in the names of mammals, birds,
reptiles and insects. It also illustrates that affinity in root
characteristic of this group of languages, traceable even in the
case of tribes separated from one another by vast distances.

«“ According to Messrs. Mabille and Jacottet, ‘Tse-Tse’ is
the name by which the Basuto know the insect, but whether it
originated with them or has been introduced from another tribe
I do not know.

«This word is not to be confounded with ‘ Ntsintsi,’ the Sesuto
name for the common fly, which in Central Africa with the
Anyanja is ‘ Nchenchi,’ with the Ahenga ‘Lumembi, and with
the Awemba of the head waters of the Congo River ‘ Runzi.’
In Equatorial Africa the Waswahili of the coast know the
common fly as ‘ Inzi,’ while the Akikuyu of the forest highlands
call it ‘Ngi.’”

RIcHARD CRAWSHAY.

LEIGHTON BuzzARD,

March 3rd, 1903.

* Received too late for inclusion in the body of the work: the reader
should compare the note on pp. 1-2.—H. EH. A.

APPENDIX G.

Tun Species oF TSETSE FOUND AT ENTEBBE, Lake VICTORIA,
Uaanpa (see page 297).

A series of specimens forwarded by Lt.-Col. Bruce which I
have just had the opportunity of examining, prove that this
species is none other than the West African Glossina palpalis,
Rob.-Desv. This unexpected discovery becomes less surprising
than it might otherwise appear when we consider the extremely
wide range of certain other species of Tsetse-flies: ¢.g., Glossina
fusca, Walk., occurs in East as well as in West Africa, while the
South African Glossina morsitans is also found in Togoland, on
the Slave Coast (see page 85).

E. E. A.
May 15th, 1903.

310 TRYPANOSOMIASIS ON UPPER NIGER.

APPENDIX H.

TRYPANOSOMIASIS ON THE Upper Nicer. A Note BY
Dr. Curusert CHRISTY.

Just before publication, I have received the following com-
munication from Dr. Cuthbert Christy, who was recently sent
to Uganda, with other observers, by a Committee of the Royal
Society in order to investigate the causes of the Sleeping Sickness.

Dr. Christy says :—“In 1898-99, at Jebba, on the Upper
Niger, there was and had been for several seasons a great
mortality among the horses of the Royal Niger Company’s
Constabulary. I remember that the mortality was so serious
that orders were at one time given to kill all the remaining
horses, healthy or otherwise, and to burn down the whole of the
extensive grass-sheds used as stables. This was done, but
without having the slightest effect in staying the disease among
the fresh supply of horses.

“T was at that time stationed at Jebba, as Senior Medical
Officer of the 2nd Battalion West African Frontier Force, and
in examining a fresh preparation of the blood of one of the sick
horses, I discovered that it contained Trypanosomes. After
this it became necessary for me to make a microscopical examina-
tion of the blood of every horse before it was purchased for the
Imperial forces, and I found that fully 50 per cent. of the horses
sent in at certain times of the year were infected with Try-
panosome disease.

‘“‘ Kach rainy season our horses had to be sent to some high
ground on the right bank of the river, or they invariably died.
A species of Tsetse was extremely common in the neighbourhood
of Jebba, although not universally distributed. On certain
reaches, often not more than a mile or so in extent, the fly made
canoe-travelling without a net most uncomfortable: the reach
that I remember best was one just north of Bussa. Again, the
left bank of a small stream only two or three miles above Jebba
was a favourite ‘belt,’ and a remarkable one to us, since we
knew little of the subject at that date, owing to the fact that
it was not more than four or five hundred yards in width. To
remain in the belt was torment, but five hundred yards from the
stream not a fly was to be seen for many miles.”

311

INDEX.

—_—_— So

A.

Alcock, Sir R. (1879), Tsetse-fly,
destruction of draught cattle
by, 39, 174

Allies of Tsetse, nearest existing, 53

Andersson, C. J. (1856), Tsetse-fly
near Lake Ngami, 10, 20, 34,
128

Antenne of Glossina, 62; figure, 63

Arnaud, M. (1852), note on flv
brought home by Mr. Oswell,
33, 126

Arnot, F. 8. (1889), Tsetse-fly pre-
valent between Mababi-flat and
the Zambesi, 202

Arsenie as a remedy for Tsetse-fly
disease, 282, 284

Association of Tsetse with big game,
12, 173, 287, 290

Austen, EK. HE. (1899), Proceedings
of expedition for study of causes
of malaria. Notes on Glossina
longipalpis (really Gl. palpalis)
in vicinity of Free Town, Sierra
Leone, 48, 245

(1900), Proceedings of Zoological

Society of London. A female
of Gl. longipennis, taken in
West Somaliland, 245

B.

Baines, Thomas (1864), Tsetse-fly
in S. Africa; at the Victoria
Falls, 34, 134

(1877), Tsetse-fly in Matabele-
land, on the Limpopo, in the
Northern Transvaal; appear-
ance of fly, symptoms caused,
remedies, 19, 38, 163

Baker, Sir S. W. (1866), Tsetse-fly
in Obbo, 35, 140

Beira Railway, fly-belt on, 45

Belts, limitation of Tsetse-fly to, 11 |

Bent, T. (1893), importance of Beira
Railway for crossing the Tsetse-
belt, 44, 208

Bibliography; literature of Tsetse-
flies, 121; of Tsetse-fly discase,
253

Bigot, J. M. F. (1885), original de-
scription of Glossina ventricosa
(= GU. palpalis), 113, 193

(1891), Glossina grossa and pal-
lacera, 207

Bionomics of the Tsetse-flies, 1

Blandford, W. F. H. (1896), the
Tsetse-fly disease, 45, 224

Blood-sucking, behaviour of Tsetse
during, 20

Boyce, Ross and Sherrington (1902),
note on discovery of the human
Trypanosoma, 267

Bradford and Plimmer (1902), Try-
panosoma brucei, the organism
found in Nagana, or Tsetse-fly
disease, 258

Bradshaw, Dr. B. F. (1881), the
Tsetse-fly about the Zambesi,
16, 19, 39, 184

Brown, W. H. (1899), Tsetse-fly in
Mashonaland, 48, 244

Bruce, James (1813), Tsetse-fly per-
haps described under name of
Tsaltsalya, 32, 121

Bruce, Surgeon-Major (1895), pre-
liminary report on Tsetse-fly
disease, 45, 219

(1897), Further Report on Tsetse-
fly disease in Zululand, 46, 226

definition of the disease, 270

distribution in Zululand, 270

nomenclature, 270

etiology, 270

description and habits of fly, 271

experiments with, 273

hematozoon (the), or blood-para-
site of fly-disease, 277

feeding experiments, 281

fly-disease or Nagana, as it occurs
in domestic animals, 270

312

relation of big game to the fly-
disease, 278
treatment, medicinal; arsenic as
a curative agent, 282; as a pro-
phylactic, 284
(1902), note on the discovery of a
new Trypanosoma, 258
Bryden, H. A. (1900), Tsetse-fly
causing abandonment of a line
of coaches from Pungwe river
towards Mashonaland, 48, 246
Buffaloes, connection between and
prevalence of fiy, 278, 290
Indian, experiments with, as to re-
sistance to effects of fly, 37, 154
Burton, Capt. R. F. (1860), Tsetse-
fly in Central Africa, 34, 133
Buxton, E. C. (1871), experiences of
the Tsetse-fly on south side of
Lebombo Mountains, 15, 36,
148

C.

Camels, Livingstone’s experiments
with, 37, 154

Capello, H. and R. Ivens (1886), a
chapter on the Tsetse-fly (in
Portuguese), 42, 195

Carter, Capt. F. F. (1880), Tsetse-fly
in Central Africa, 8, 176

Casati, Major G. (1891), the Tsetse-
fly west of Lake Albert Nyanza,
43, 206

Castlenau, L. de (1858), the Tsetse-
fly in South Africa, 34, 131

Chapman, James (1868), symptoms
of bite of Tsetse-fly; native
remedies; prevalence of fiy in
various parts, 7, 22, 35, 141

Characters, general, of Tsetse-fly,

Chavanne, J. (1887), region of Lower
Congo free from Tsetse-fly, 42,
199

Christy, Dr. C., note on Trypano-
somiasis in horses on the Upper
Niger, 310

Clark, Bracy (1857), attempts to
show that Tsetse is identical

with Wstrus (Hypoderma)
bovis; refuted by Westwood,
34, 131

Coaches, line of, abandoned, owing
to Tsetse-fiy, 48

Corti, E. (1895), original description
of G. longipenmis from Somaili-
land, 52, 219

Crawshay, Capt. R., note on the
“ Tse-Tse ” fiies of British Cen-
tral Africa Protectorate, refers

INDEX.

mainly to Glossina morsitans
and pallidipes, 20, 287
note on the origin of the word
“ Tse-Tse,”? 309
Cumming, R. Gordon (1850), the
Tsetse-fly in the Transvaal, 33,
125

D.

Day, time of, at which Tsetse most
active, 16

Dogs, liable to suffer from Tsetse-fly
disease, 280

Donitz, W. (1896), report of paper
on Tsetse-flies, 226

Donkey, immunity of, asserted by
Mauch, 36; immunity denied,
46

Drysdale, Dr. J. J. (1879), an early
suggestion as to the true part
played by the Tsetse-fly in con-
nection with the disease, 38,
173

Duparquet, Pére (1881), Tsetse-fly
on the Okavango river, at its
junction with Lake Ngami, 179

Duration of illness in animals bitten
by Tsetse, 142

Dutton, Dr. J. B. (1802), preliminary
note on a Trypanosome occur-
ring in the blood of man, 259

E.

Eckersley, W. A. (1895), Tsetse-fly
on line of Beira Railway; two
ponies taken through fly-belt
without ill results, 45, 220

Elephants, Indian, experiments
with, as to immunity from
effects of bite of fly, 40, 187

Eliot, Sir C., letter from, regarding
connection of fly with buffaloes
ard other big game, 291

Elton, Capt. J. F. (1879), question
whether disease and death of
cattle on Unyamyembe route
from coast be due to Tsetse or
to rank vegetation, 39, 173

Encyclopedia Britannica (1888),
article on Tsetse-fly referred to,
43, 201

Erskine, St. V. (1870), paper ques-
tioning idea that bite of Tsetse-
fly is destructive of animal life,
35, 146

Explanation of Figures (Plates VIII.
and IX.), 117

INDEX.

F,
Fitzgerald, W. W. A. (1898). The
Tsetse-fly on the coast of

British Hast Africa, 47, 240

Flight of Tsetse-fly, mode of, 19

‘EF ly-belts,” 9; Boer method of
crossing at night, 39, 184, 203

Foa, E. A. (1895), the Tsetse-fly in
Central Africa, 19, 45, 221

(1897), bite of the fly harmless to

wild animals, especially buffalo
and large antelope, 231

Forde, R. M. (1902), clinical notes
on a Huropean patient in whose
blood a Trypanosoma was ob-
served, 266

(1902), the discovery of the human

Trypanosoma, 267

Foster, Capt. (1893), Handbook of
British East Africa; localities
affected by Tsetse-fly, 44, 211

Free Town, Sierra Leone, Gl. pal-
palis found there by author,
76, 92

Frere, Sir B. (1881), method of
crossing the fly-belt south of
the Zambesi, 177

Fynney, F. B. (1878), Tsetse-fly in
Zoutpansberg ; predicts its dis-
appearance with extinction of
big game, 38, 173

G.
Galla method of crossing fly-belts,
225

Game, large, prevalence of Tsetse-
fly dependent on existence of,
and extinction with disappear-
ance of, 12, 173, 287, 290, et seq.

Geographical distribution of Tsetse-
flies, 28

Ghika, Prince N. D. (1898), des-
truction by Tsetse-fly of cattle
and horses in the Aulihan
country, Somaliland, 47, 234

Gibbons, Capt. A. St. H. (1897),
Tsetse-fly on the banks of the
Zambesi, 47, 226

(1898), Exploration and hunting
in Central Africa, 7, 18, 236

Gielgud, Mr. van, 1901, letter on
connection between Tsetse-fly
and buffalo, 290

Glossina and Stomoxys, the mouth
parts of, 105

—— antenne of, 62

— chetotazy of, 67

313

Glossinaand Stomoxys, distinguished
from specimens of Stomoxys
and Hxmatopota, 5

—— distinguishing characters of
genus, 60

— external mouth-parts and mode
of action, 106

—— genus containing seven known
species, 3

—— head, description of,61; thorax,
abdomen, 64; legs and wings,
65; antenne, 62

—— internal mouth-parts and mode
of action, 114

—— resemblance between and Lype-
rosia, 56

—— —— in venation of wings be-

tween, and Hypoderma,
6

—— synopsis of species of, 70

— fusca, description of, 95; head,
thorax, abdomen and legs,
96; veins in wings, 97; dis-
tinguishing characters of, 97;
distribution, 98; habits, etc.,
99; synonymy and affinities,
100

— longipalpis, description of, 90;
distinguished from Gl. palli-
dipes, 91, 93; males distin-
guished from those of Gi.
morsitans and pallidipes, 94 ;
head, thorax, abdomen and
legs, 91; distribution of, 91;
synonymy and affinities, 92;
hypopygium in, figure of, 94

—— longipennis, description of,
head, 101; thorax, abdomen,
legs and wings, 102; distri-
bution of, 103; habits, etc.,
103; synonymy and affini-
ties, 103

— morsitans, description of, 81;
distinguished from Gl. palli-
dipes and longipalpis, 93-94 ;
head, thorax, abdomen, legs,
82; hypopygium in, figure
of, 94; wings, 83; distri-
bution of, 83; habits of, 86;
synonymy and affinities of,
87

—— pallicera, description of, head,
thorax, 79; abdomen, 80;
distribution and habits, 80;
synonymy and affinities of, 80

— pallidipes, description of, 87;
distribution of, 88; habits
of, 90; synonymy and affini-
ties of, 90; distinguished
from Gl. morsitans and longi-
palpis, 93

314

Glossina palpalis, description of, 71 ;
head, 72; abdomen and legs,
73; distribution of, 75, 309;
habits of, 76; synonymy and
affinities of, 77; var. tachin-
oides, Westw., 77
— and Stomoxys, the mouth-
parts of, by Dr. H. J. Hansen,
105 (see also mouth-~parts)
Goats, doubtful immunity of, from
effects of Tsetse, 47
Gregory, J. W. (1896), Tsetse-fly in
belt of forest between Witu
and coast, 46, 225
Grube, Dr. (1874), lecture on Tsetse-
fly, with résumé of Living-
stone’s observations, 154
Guerne, Jules de (1894), report of
exhibition of two specimens of
Gl. morsitans, collected in Cen-
tral Africa by E. Foa, 44, 218

H.

Habenicht, H. (1887), limits of
Tsetse-fly north and south of
Limpopoand south of Zambesi,
42, 199

Hzmatopoita, distinguished from
Glossina, 6

—— figure of, in resting attitude, 6

Hematozoon of Tsetse-fly disease,
see Trypanosoma

Hansen, Dr. H. J., mouth-parts of
Glossina and Stomoxys, 105

Hardinge, Sir A. (1897), Tsetse-fly in |
East Africa; cattle flourish in —

Ukamba, 227

Harris, Sir W. C. (1839), Tsetse-fly _
in Mural hills, Waterberg dis- |

trict of Transvaal, 16, 32, 124
Hartmann, Dr. (1877), lecture on
specimensiof Tsetse-flies, 53, 161
Haslam, Capt. A. G., possible con-
veyance of hematozoon of
Tsetse-fly disease by Stomoxys,
304; death of, 100 (note)
Hinde, 8. L. and H. (1901),
Tsetse-fly belt in Hast Africa,
49, 250
Historical survey of literature of
Tsctse-flies, 31
Holub, Dr. EK. (1880), Tsetse-fly in
Matabeleland, 176
(1881), near Panda ma Tenka
river, 181
(1888), in the Batoka country, 42,
200

(1890), on the Zambesi, 43, 203

the ©

INDEX.

Hore,E. C, (1882), Tsetse-fly on shores
of Lake Tanganyika, 40, 186

Hornor, Lewis (1879), recommend-
ing Boer method of crossing
fly-belts at night, 39, 175

Horse-flies (Tabanidz) bites may be
fatal to domestic animals, 305

Hough, G. de N. (1898), on two
females of Gil. longipennis, col-
lected in Somaliland, 47, 242

House-fly, differences between, and
Tsetse-fly, 64

Humidity of soil, and low-lying
positions generally necessary
for existence of Tsetse-fly, 294
(Cf., however, 7)

Hypopygium, 64; figures of, in Gi.
morsitans and longipalpis, 94

J.

Jackson, F'. J. (1893), Tsetse-fly on
Teita-Kibwezi route to Uganda,
16, 21, 44, 209; (1894), T'setse-
fly in Hast Africa not closely
associated with big game, 12,
44,217

— letter on non-dependence of

Tsetse-fly on presence of big
game, 295

James, F. L. (1883), ‘‘The Wild
Tribes of the Soudan,’ supposed
presence of 'T'setse- fly, 40, 190;
(1885), fly (?Tsetse) on the
Webbe Shebeyli (Somaliland)
in the wet season, 41, 44, 194

Jeppe, F. (1888), partial ‘disappear-
ance of the fly together with
game from the Kaap Valley,
Transvaal, 14, 42, 201

Johnston, Sir H. H. (1894), regions
of eastern portion of British
Central Africa affected by
Tsetse-fly, 44, 125, 214; (1896),
disappearance of fly from eoun-
try to south of Lake Nyasa, 31,
46, 224; (1897), British Central
Africa, Indian buffaloes not
affected by bites of Tsetse-fly,
46, 228; parts in which fly is
present, 7, 8,9; (1899), History
of Colonization of Africa by
alien Races: Arabs accused in
1569 of having poisoned horses
and camels, 31, 48, 243; (1901),
journey from Mombasa _ to
Uganda, difficulties owing to
fly, 48, 246; (1902), suggested
absence of the “‘true Tsetse-
fly” from the Uganda a sepa
rate, 50, 251, 301

INDEX. 315

Justice, J.N.(1902), virulence of bite ©
of fly said to have diminished —
since extermination of buffalo |

in Northern Rhodesia, 50, 252

K,

Kanthack, Durham and Blandford
(1898), on Tsetse-fly disease,
254

Karsch, Dr, F. (1893), Tsetse-fly in
hinterland of Togo protecto-
rate, 44, 213

Keane, A. H. (1888), article “‘ Trans-
vaal” in Encyclopedia Britan-
nica, 201

Kerr, W. M. (1886), the Tsetse-fly in
Mashonaland; travels from the
Cape across the Zambesi, ex-
periences of fly, 42, 195

Kersten, Otto (1871), Tsetse-like fly
on the Juba River, 150

Kingsley, Mary H. (1897), Tsetse-fly
on the Gold Coast, 46, 232

Kirby, F. V. (1899), the T'setse-fly
in Hast Central Africa, 243

Kirk, Sir J. (1862), the Tsetse-fly in
the Shiré, and lower Zambesi
valleys, 134; (1865), the Tsetse-
fly of tropical Africa, 16, 18, 34,
137; (1881), visit to Dar-es-
Salaam district, the Tsetse-fly,
179

Koch, R. (1898), fly-disease in
German East Africa, and Try-
panosomes in blood-films re-
lating to Tsetse-fly disease, 254

L.

Laboulbéne, A. (1888), Tsetse-fly
not poisonous in itself; very
troublesome in districts inland
from Bagamoyo, 42, 199

Larva of Tsetse-fly, Col. Bruce’s
description, 24

Lavyeran, A., and F. Mesnil, mode
of multiplication, morphology,
etc., of Trypanosoma, 49, 255,
256, 258

Laveran, A., onthe action of human
blood-serum on the Trypano-
some of Nagana, 258

— Researches on the Treatment
and Prevention of Nagana, 267

Lawley, Capt. Hon. A. (1898), move-
ments of the Tsetse-fly with

the big game of the country,

47, 239

Laws, Dr. (1879), cattle-destroying
fly (? Tsetse) in valley of Lim-
passa; fly absent on portion of
road (40 miles) from Dar-es-
Salaam, 174

| Leverson, Major J. J. (1893), Tsetse-

fly on and near Pungwe river,
44, 209

Limitation of Tsetse-fly to belts,
explanation of, 11

Livingstone, David (1857), remarks
on the Tsetse-fly (Glossina mor-
sitams), 10, 14, 23, 34, 129;
David and Charles; (1865),
Tsetse-fly mear the valley of
the Zambesi, 8, 135; (1874),
experiments with camels and
Indian buffaloes introduced
from Bombay, 37, 154

Lugard, Sir F. D. (1893), British
East Africa practically free
from Tsetse-fly, 43, 207

M.

Macloskie, G. (1884), abstract of
Kraepelin’s paper on the pro-
boscis of Musca, 192

Macquart, J. (1835 and 1843), de-
scription of the genus Glossina,
and of Gl. longipalpis, 33, 123,
124; (1850), description of Gl.
longipalpis, 33, 125; owing to
the fineness of the mouth-parts,
believed that the flies did not
suck blood, 33

McClellan, J. W. P., on Tsetse-fly
in Jubaland province; its pre-
sence not depending upon big
game, 294

Mainwaring, Major H. S. (1895),
Tsetse-fly in Somaliland, 45, 219

Map (1887), Justus Perthes’ Special
Map of Africa, showing limits
of fly-belts on Limpopo and
Zambesi, 42, 199

Marno, EH. (1873), specimens of
Tsetse-fly, called ‘‘ Surréta” by
natives, 37, 153

Masui, Lieut., and — Seeldrayers
(1897), the common house-fly
said to cause Tsetse to dis-
appear, 231

Mauch, Karl (1869), the Tsetse-fly
near Olifant’s River, 35, 145;
(1870), on a journey from
Lydenburg to Inyati, 36, 147;
(1874), Tsetse-fly between the
Limpopo and the Zambesi, 37,
159

516 INDEX.

Maund, E. A. (1891), Tsetse-fly in
Mashonaland, 14, 43, 206

Mik, J. (1885), reference to three
additional species of Glossina,

?
Mohr, E. (1871), Tsetse-fly on road
to Victoria Falls, 36, 149
(1876), Tsetse-fly near the Zam-
besi, 37, 160
Mombasa-Victoria (Uganda) Rail-
way, Report on Progress of;
mules more resistant to Tsetse-
disease than other animals, 241
Mouth-parts of Glossina and Stom-
oxys (Dr. Hansen), 105
— Glossina, external mouth-parts,
maxillary palpi, 106 ; labrum,
107; hypopharynx, 107;
labium, 107
— Stomoxys, external mouth-
parts, labrum, 110; hypo-
pharynx and labium, 111
— external, of Glossina and Stom-
oxys, mode of action of, 112;
internal, mode of action of,
114
— actual piercing organs in bath
genera, 113
Mullens, Rev. Dr. J. (1877), absence
of Tsetse-fly on a new route to
Lake Tanganyika, 38, 163
Muscide, family to which Tsetse-
flies belong, 53

N.

Nagana, or ‘Tsetse-fly disease,

Bruce’s definition of, 270

foci in German East Africa,

261

Native names for Tsetse-flies, 299

Nepveu, Dr. G. (1898) on a Try-
panosoma in human blood, 254

Neumann, A. H. (1898), ‘‘ Elephant
Hunting in East Equatorial
Africa”; mentions “plenty of
birds and of fly’ on Athi River,
where other animals are scarce,

, 47, 234

Newman, Edward (1870), supports
Eirskine’s view that the Tsetse-
fly’s bite is not destructive of
animal life, 36, 147; (1871) cri-
ticises Buxton’s remarks to the
contrary, 151

Nicholls, J. A. and W. Eglington
(1892) ‘‘ The Sportsman in South
Africa’’; the Tsetse-fly preva-
lent where buffaloes are abun-
dant, 114, 207

Niger, R., Trypanosomiasis on
Upper, note on, by Dr, GC.
Christy, 310

Night, Tsetse-fly sometimes active
during, 18

Nocard, HE. (1901), the relations
between Dourine, and Swrra or
Nagana, 255

Nuttall, G. H. F. (1898), recent
investigations on malaria, Texas
fever, and Tsetse-fly disease,
253; (1899), the réle of insects
as carriers in the spread of
bacterial and parasitic diseases,
254

O.

Oates, (1881), Tsetse-fly near
Victoria Falls, 52, 180
Cistrus (Hypoderma) bovis, sup-
posed identity of, with Tsetse-
fly, 34, 131

Olfactory powers of Tsetse, 19, 223

Origin of name Tsetse, 1 (note), 309

Oswell, W. C. (1894), Big Game
Shooting: insect attacking the
Tsetse-fly ; districts affected by
fly, 44, 217

Oswell and Arnaud (1852), on a
venomous fly in Southern
Africa, 383, 127; observations
on the Tsetse-fly, 128

Ps

Parasite of Tsetse-fly disease, ques-
tion whether conveyed by blood-
sucking flies other than Tsetse,
803

Pechuél-Loesche, Dr. (1882), the
Tsetse-fly on the Congo, 40,
190

Peel, C. V. A. (1900), Somaliland,
prevalence of Tsetse-fly, 9, 48,
245

Perthes, J. (See Habenicht).

Petherick, John and Mrs. (1869),
experiences of Tsetse-fly north-
west of Gondokoro, 16, 35, 146

Pinto, Major Serpa (1881), Tsetse-
fly on right bank of the Zam-
besi, 179

Plimmer and Rose Bradford (1899),
the morphology and distribution
of the organism found in the
Tsetse-fly disease, 255

Portal, Sir G. (1894), absence of
Tsetse-fly between Zavo and
Kibwesi, 44, 216

INDEX.

Preuss, Dr. (1889), a species of
Glossina at Barombi, Came-
roons, 43, 202

Problems awaiting solution in con-
nection with Tsetse-fly, 286

Proboscis of different species of
Tsetse practically identical in
structure, 106, 302

Prophylactics against and remedies
for Tsetse-fly disease, 282, 297

Pupa of Zululand Tsetse-fly, 26,
et seq.; figure of, 27

R.

Radford, Dr., letter on Tsetse-fly in
East Africa Protectorate; con-
nection between it and buffalo
and other big game, 292

Rankin, L. K. (1882), the elephant
experiment in Africa stated to
be a complete success, 22, 40,
187

Reproduction of Tsetse-fly, 23

Resting-position, identification of
Tsetse-fly in, 3, 168

Rhodesia, northern, Tsetse-fly in,
50, 252

Richards, E. H. (1885), Journey in
East Africa; the Tsetse-fly on
the Luizi River, 194

Robineau-Desvoidy (1830) original
description of genus Nemorluna
and of N. (Glossina) palpalis,
82, 51, 123

Réder, V. von (1893), Glossina mor-
sitans taken at Mbuzini,German
East Africa, 208

Rogers, Dr. L. (1901), probable iden-
tity of Swrra of India, and
Nagana, or Tsetse-fly disease,
of Africa, 257

s.

Sambon, Dr. L. W. (1902), Note on
discovery of the human Try-
panosome, 267

Scepticism as to Tsetse-fly’s powers
35, 146, 191

Schilling, Dr. (1901), Tsetse-fly in
the Togo Protectorate, absent
from the littoral, 49, 250;
‘‘surra,”’ a disease fatal to horses
in the Togo Protectorate, 257 ;
description of the trypanosome
causing the disease, 250 ; (1902)
the ‘‘surra”’ disease of horses

317

and cattle in the Togo Protec-
torate, 266; (1903) Nagana and
other trypanosomes, 267

Schoch, G. (1883), Tsetse-fly of
South Africa; a carrier of
bacterium-like poisonous mat-
ter, 40, 191

Schoeller, Dr. M. (1901), reports
death of baggage donkeys in
Hast Africa, but Tsetse-fly not
found, 49, 247

Schmidt, Dr. C. W. (1887), non-
occurrence of Tsetse-fly in
Usambara [Cf., however, 261],
42, 199

Schulz, Dr. A., and A. Hammar
(1897), Tsetse-fly on the swampy
offshoots from the Chobe River,
46, 232

Seasonal prevalence of Tsetse-fly, 15

Selous, F'. C. (1881), Tsetse-fly in
Mashonaland, description and
details, 8,17, 18, 39,181; (1883),
further experiences of fly, 8,
192; (1893), Travel and Adven-
ture in South Hast Africa, the
Tsetse-fly, 10, 44, 211

Sensations produced by bite of
Tsetse-fly, 21, 245

Seroot flies in Sudan, 308

Serum treatment of Tsetse-fly dis-
ease, 285

Sharp, D. (1899), Cambridge Natural
History, notes on Glossina mor-
sitans, 243

Sharpe, A. (1896), Tsetse-fly absent
from highlands of British Cen-
tral Africa, 226; (1901), letter
on presence of Tsetse-fly not
dependent solely on that of wild
animals, 295

Smith, Dr. D. (1894), experiences in
Somaliland; camel-killing fly
in the Webi Shebeli, 44, 214

Smith, J. B. (1890), structure and
history of Hematobia serrata,
202

Somaliland, fly-disease in, 41, 194,
214, 251

Stanley, H. M. (1872), experiences
of Tsetse-fly (three species of
biting flies), 22, 36, 151; (1878),
gad-(horse-)flies and Tsetse-fly,
38, 172; (1885), bitten by Tsetse-
fly on board steamer on an
atfluent of Congo, 194

Stordy, R. J., M.R.C.V.S. (1901),
letter stating that buffalo and
other big game are not the only
factors connected with the
Tsetse-fly, 291

318

Stomoxys, figure of, in resting atti-
tude, 5; perhaps capable of
conveying hematozoon of
Tsetse-fly disease, 304; distin-
guished from Tsetse, 5, 56;
external mouth-parts and their
mode of action, 110; internal
mouth-partsand mode of action,
114; lower portion of head, 114

Stuhlmann, Dr. F. (1902), Surra-
disease assumed not to be con-
veyed by Glossina tabantformis
(fusca), 50, 253; notes on the
Tsetse-fly and surra-disease
conveyed by it in German East
Africa, 259

*Surra,”’ a disease fatal to horses,
question whether conveyed by
Tsetse-fly, 49, 153, 250, 253, 257,
259.

Symptoms in animals bitten by
Tsetse, 142, 168, 186, 247

pie

Tabanidx (horse-flies) effects pro-
duced by bites of, whether
similar to fly-disease, 305

Theiler, A. (1901), the Tsetse-disease,
257

— discovery of a new Trypano-
soma, peculiar to cattle, 259

Transvaal, Tsetse in, 201

Trypanosoma brucei, the hemato-
zoon of Tsetse-fly disease, v, 253—
267, 277, 301

Trypanosomiasis in horses on the
Upper Niger, note on, by Dr.
C. Christy, 310

“Tsaltsalya” or “ Zimb” of J. Bruce,
82, 121

Tsetse-fly (see also Glossina)

—— association of, with big game,
questions as to species most
closely connected with fly,
12,15, 293, 295, 296

— behaviour of, at night; sluggish
when cool, 18

— —— when sucking blood, 20

bionomics of, 1

—— blood-sucking habit common
to both sexes, 23

time occupied in 22

-—— chief foci of disease caused by,

300

— dislike of, to human habita-
tions and to animal excreta,
23

—— distinguishing characters of
genus, 3, 60

—— discovery of species of, 2

INDEX.

Tsetse-fly, distinguished from Sto-
moxys and Hxmatopota, 3
— doubtful whether all species
are capable of conveying
hematozoon, 2
—— earliest accounts of, 31
effects of bites, 21, 22
—— existence of, in districts desti-
tute of big game, 13, 291,
293-296
—— fly-belts, extent and distribu-
tion, 6, 11
—— fondness of, for river-banks and
neighbourhood of water, 7
formerly supposed to secrete
a poison; in 1895 discovered
to be the carrier of a hema-
tozoon, v., 219, 226
—— genera most closely allied to, 59
—— general characters of, and
distinction from other flies, 3
—— geographical distribution of, 28
—— historical survey of bionomics
of, 31, and of systematic
literature of, 50
— larva of, 24
—— limitation of, to belts, ex-
plained, 11
localities in which found, fly-
belts, 9, 11, 28, 300
—— mentioned as such first by
Gordon Cumming (1850), 33,

125
—— mode of feeding, 21
—— —— flight, and noise thus

produced, peculiarities
of, 19
—— name, origin of, probably first
applied by natives to flies
generally 1 (note); note on
origin of, by Capt. R. Craw-
shay, 309
—— native names for, 290
olfactory powers of, 19, 223
pain caused by bites of, 21,
245, 288
parts of animals usually at-
tacked by, 22
—— period of day in which most
active, 16, 288
—— probability of extinction, fol-
lowing destruction of big
game, 288
-—— proboscis of different species
practically identical in struc-
ture, 106, 302
—— pupal stage of in Zululand
species, 26
—— question as to manner in which
effects of bite are produced, v
—— reproduction of, 23

INDEX.

Tsetse-fly, seasonal prevalence of, 15
— systematic position of genus,
51, 53
technical description of genus
and species, 60
term as used, generally equiva-
lent to the genus, 2; onoma-
topoetic in origin, 1 (note)
—— three species known to exist in
East Africa, 14
—— time taken in sucking its fill,
20
—— types of species examined by
author, vii

U.
Uganda, absence of fly-disease in,
301

railway, results from, in extinc-
tion of big game, 49, 250;
enables horses and mules to
be transported through fly-
belt for use in Ukamba, 249
Ungulates, Tsetse-fly disease not
limited to, 263

W.

Walker, F. (1849), original descrip-
tion of Stomoxys fuscus, 51, 124 ;
(1873), Central African blood-
sucking flies, 153

319

Wandorobo or Donderobo fly, 305
Westwood, J. O. (1850), original
description and figures of Glos-
sina morsitans, tachinoides
= palpalis) and tabaniformis
aon 32, 33, 34, 51, 125, 127,
128

Whyte, A. (1895), Report on botanical
aspects of British Central
Africa; the Tsetse an obstacle
to agricultural enterprise, 219

Wiedemann, ©. R. W. (1830),
original description of genus
Glossina, and of Gil. longipalpis,
51, 123

Wulp, F. M. van der (1884), an
account of the Tsetse-fly, 41,
193 ; (1885), remarks at meeting
of Nederlandsche Entomolo-
gische Vereeniging, 41, 193

Z.

Ziemann, Dr. H. (1902), the Tsetse-
fly disease in Togo, 266

‘“‘Zimb,” term mentioned by J.
Bruce, perhaps designating
Tsetse, 32, 121

Zululand, Tsetse-fly abundant in
parts of, 30

LONDON:
PRINTED BY WILLIAM CLOWES AND SONS, LIMITED,
DUKE STREET, STAMFORD STREET, S.E., AND GREAT WINDMILL STREET, W.

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