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THE

Academy of Natural Sciences

OF

PHILADELPHIA

Williamson Libuau

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NOT TO B K LO A NK 1)

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I

Annals of Tropical Medicine
and Parasitology

CUW £Si

THE UNIVERSITY OF LIVERPOOL

Annals

OF

Tropical Medicine and
Parasitology

ISSUED BY THE

Liverpool School of Tropical Medicine

VOLUME I

(February ist, 1907, to February 29th, 1908)

With fifty Plain and. Coloured Plates and
seventy Figures in the text

LIVERPOOL: AT THE UNIVERSITY PRESS
LONDON: WILLIAMS & NORGATE, 14 HENRIETTA STKI I I
COVENT GARDEN, LONDON

RC u i R~
AL
V, |

32231

CONTENTS

No. i

Newstead, R. ; Dutton, J. E., and Todd, J. L.

Insects and other Arthropoda collected in the Congo Free State. Plates I-' 1

Neumann, G.

Description of two new species of African Ticks ......

Looss, A.

On some parasites in the Museum of the School of Tropical Medicine. Liverpool
Plates VII-IX .

Carter, R. Markham.

The presence of Spirochaeta duttoni in the ova of Ornithodoros mouluua. l’l.n< X

Moore, Benj. ; Nierenstein, M., and Todd, J. L.

A note on the Therapeutics of Trypanosomiasis ....

No. 2

Ross, E. H., and Ross, H. C.

An Automatic Oiler for the Destruction and Prevention of Mosquito Larvae
in Cesspools and other collections of water. Plate XI

Stephens, J. W. W., and Newstead, R.

The Anatomy of the Proboscis of Biting Flies, Part II, Stomoxys. Plate- XI I - \ I \

Dutton, J. E. ; Todd, J. L., and Hanington, J. W. B.

Trypanosome Transmission Experiments

Dutton, J. E. ; Todd, J. L., and Kinghorn, A.

Cattle Trypanosomiasis in the Congo Free State .

Moore, Benj.; Nierenstein, M., and Todd, J. L.

Concerning the Treatment of Experimental Trypanosomiasi-

PAGP

I

*»3

121

*55

i6t

*63

169

*99

*3*

• *73

CONTENTS

No. 3

Dutton, J. E. ; Todd. J. L., and 'Iobey, E. N.

PACE

Concerning certain Parasitic Protozoa observed in Africa. Plates XX-XXIl . 285

Branch, C. W.

Yaws

• 37*

Graham, W. M.

A description of some Gold Coast Entomostraca . . . . . 415

Brady, G. S.

Notes on Dr. Graham’s Collection of Cyclopidae from the African Gold Coast.

Plates XXIII-XXXVI . 423

Breinl, A.

On the Morphology and Life Plistory of Spirochactu duttoni. Plate XXXVII . 433

Salvin-Moore, J. E., and Breinl, A.

I he Cytology of the Trypanosomes. Plates XXXVI 1I-X LI I .

No. 4

Walker, C. E.

Observations on the so-called “ Canary Fever ”

Broden, A., et Rodhain, J.

Contribution a l'otade do Poroc'fhalm monilifor, ms.

Plate XLIIJ

• 493

Newstead, R.

iing plac« of the common House-fly {Mtuc a

• 507

Van Loghem, J. j.

HEM, J. J.

in the organs of rats.

Jones, W. H. S.

Malaria and History.

Stephens, J. \y. YV

TW° "ew Cestodes

!i anel a new Linguatulid. Plate I,

• 549

INSECTS AND OTHER ARTHROPODA
COLLECTED IN THE CONGO
FREE STATE

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INSECTS AND OTHER ARTHROPOD A
COLLECTED IN THE CONGO
FREE STATE

Being the Seventh Interim Report of the Expedition of the Lire>p> <>l
School of Tropical Medicine to the Congo. 1003-05

BY

ROBERT NEWSTEAD, A.L.S., F.E.S., &c.

( LECTURER ON ECONOMIC ENTOMOLOGY AND PARASITOLOGY. I III M IL Ol . I
I ROPICAI. MEDICINE. UNIVERSITY OF I IVKRPOOI i

The late J. EVERETT DUTTON, M.B. (Vict.)

(WALTER MYERS FELLOW. UNIVERSITY OF I IVKRPOOI i
AN I)

JOHN L. TODD, BA, M.D., C.M. McGill

(DIRECTOR OF THE RUNCORN RESEARCH LABORATORIES Of III!

LIVERPOOL SCHOOL OF TROPICAL MEDICINE

INTRODUCTORY

This expedition was sent to the Congo Free State, on the
invitation of His Majesty King Leopold II, by the Liverpool School
of Tropical Medicine to report upon the sanitary condition of the
more important posts and to study human trypanosomiasis,
collection of various biting insects formed, therefore, a principal part
of its work; but once the more important ones, as (j lossina and
Anophelina , were obtained in each district the systematic search i :
insects was discontinued. A large part ot this collection repi « •->«
then, merely the casual wayside gleanings made during a i"'irn<-\
through Central Africa.

From the point of view of a dipterologist the collection is therefore
regrettably incomplete, especially so, since a large number • ■'
specimens have been unavoidably ruined by moulds. Had time hern
spent in collecting, it is probable that many of those inserts rep- rt.-d
from only one or two localities would have been found to have a Dr
wider distribution.

Twenty-three months were actually spent in the Congo I n-'
State ; during that time a distance of some 2,000 miles was travelled
The course followed by the expedition is indicated on the map; while

4

uic vrfixuus puais

the dates when it was at, or passed through,
briefly mentioned below.

England was left on September 4, 1903. Banana,' l hr first point
of call in the Free State, was reached on September 23. From
September 24 to October 27 was spent at Boma, and from October _*S
to November 20 at Matadi. Matadi is connected with Leopoldville
by railroad. From I’umba, the half way point on the railway line,
two of the members of the expedition made a trip of three weeks’
duration (November 5 to 28) to Wathen and through the District of
the Cataracts. A stay of seven months, from November 22, it> to
June 23, 1904, was made at Leopoldville.

From Leopoldville the expedition went up the Congo to
Falls in a small steamer placed at their disposal b> t (

Vhen it seemed advisable, stops and short side joUTOC)
at posts along the river for the purpose of studying local conditions
Work was done at M'Swata from June to aft At Tshumbiri

nland2 V T Was Spent at a nativ(- «"»8« some ten miles

" ™ ,7 M“POieil?Ue visited on July 7 to ft and Irehu „n Jafy

O Ea L Jr C°qU,"‘aTtVille Gv,y 18 to 9) trips were made

to ft p 1 M T?' J°uvelIe Anvers was visited front Angus, ,4

Mongala to Bokanga ^ ^ "* "WH" "»*

(August 26 to *o) and nt U . S PS Were mad(* :,t l

of these places short i„t m ( gUSt 3 '.t0 SeP^mber 4) ; at both
made at Basoko (September’^ i ml*nd‘ Short staXs Nvrr<*

'5). On Septimus /tani^ Fa,," Yal<U5U '3 -

expedrtton remained until September af* rea' ' d' wbere thr

from Stanley Falls the journey no'the r
Kasongo by canoe. Ponthierville B° "'aS con,ln"'-H »°

°n‘°ber 9. From £^^7" October s to

expedition made a side trip Qf , 2 d J, *'V° n"‘l"bers of the
of the Congo, They paralleled the i ^ duratlon lnland to the east
once more took canoe. From Lok rA “ Kumba- where they
a J-rney of some l8 mi, ™ <0 November ,)
N Sendwe (November 4 tQ “ ^ * Stops "ere made at

Kasongo was reached on November

‘ 5,ay°f five month.

5

was made at this post, and it is there that Dr. Dutton died and is
buried.

Kasongo was left on April 27, 1905. The journey thence to Pania
Mutombo lay overland and was done on foot. Kalombe was reached
on May 3, Maomedi on May 5. No stop of more than a day was made
before reaching Tshofa (May 13 to 25). Four days, June 2 to 6,
were passed at Cabinda. At Pania Mutombo (June 12) canoes were
once more employed for the two days' journey down stream to
Lusambo (June 5), where the expedition remained until July 8, 1905.

The journey from Lusambo to Leopoldville was done as quickly
as possible by steamer, and no stops were made for work. At
Leopoldville (July 21 to 30) arrangements were made for leaving Afrit .1
The railroad between Leopoldville and Matadi was once more
traversed, and on August 8, 1905, the expedition left the Congo Free
State. On the way home Freetown, Sierra Leone, was visited for a
week (August 16 to 22). Liverpool was reached on September 5, 11/ 5 ,
the expedition had been absent from England for just two years

Other side-trips were made besides those mentioned ; and it must
be understood that when a stay of any duration was made in a post
short journeys were always made in its neighbourhood.

As is shown by the map, the route followed by the expedition
twice crossed the equator and traversed a territory King between
2° North and 6° South of that line. The climates of the posts visited
therefore varied somewhat. The succession of the seasons at the
most important places is briefly as follows :

At Boma the seasons are irregular. There is usually said to be .1
long and a short wet, and a long and a short dry, season. Practically,
we may say that from May to September there is but very little
or no — rain ; and that from October to the end of April there is a
good deal ; from January to April are the hottest months of the year
It is then, from October to May that conditions are the must
favourable for the breeding of mosquitoes. At Leopoldville rain
more likely to fall in every month of the year than at Boma. 1 o m
the 15th of June to the 1st of September is usually considered to 1 ><•
the dry season, and during this period there may be practically no rain
Coquilhatville is almost on the equator, and consequently has m
marked variation in its seasons. Rain falls in ever)- month u! tin*
year, perhaps the most in November and December The

6

temperature is constantly high, and during the first four months of the
year it may be very hot. The climate is then admirably suited for the
development of mosquitoes.

Bumba is one of the furthest north among the posts visited by the
expedition. The seasons there are not well marked, r.iin fall-, in
every month of the year ; perhaps the least in January and Februar\

At Kasongo and Cabinda the seasons are much the same .is at
Soma, save that the ‘dry season'' is shorter and rain often falls m
August and May.*

When the expedition arrived at Boma the rainy season had .dread)
set in. For the first half of the month spent there, rain fell almost
daily, the thermometer meanwhile varying between 85° anti >°I-

During the latter two weeks there was . . jcasional a . . 01 ' •

and the temperature was slightly higher.

While at Matadi there was but little rain, so that but few of those
places were seen where rain-water collects during the wet
orm breeding-places for mosquitoes.

was reached the season *» »c

were fitd t gr°Und WaS S°dden W'th Waler an<i strear,

dai y and tl ” 0‘" November April ran, fell almo

da,ly and there were frequent thunderstorms. The dally tempemtu

■ dryed st:;. wtoth89°tF' ,n May ^ re1,,
collenced lhhoTS thL^nr^ **' ^

™ - . . . . .

r - -tt; z:tz z . . t .

. . — °~ » '*• "■>“>' a

szr-x

are indebfed^/S™a,1°n concerning the clinuue o! >1 - n -

:

* . V*"''"’ ''

Commandant VeSic^fe”1 °f hla “«"»l°gual obaerlaUnn, ^“7," *'“

kasongo. a copy „f ,h„

7

were, however, one or two hot days (maximum g6°F), and while at
the Falls there were a couple of severe thunderstorms.

From Stanley Falls the route lay south through the equatorial
zone towards Kasongo where the rainy season had already set in
Frequent showers and thunderstorms were encountered on the way
up river. The weather was pleasantly cool, the temperature varying
usually between 82° and yi°F.

From November to April there was much rain at Kasongo
(December, 2 16*3 mm.). The temperature meanwhile varied between
68° and 85°F. The march overland from Kasongo to Lusambo in
May and June was made during the commencement of the dry season
Rain fell but rarely, the temperature became higher and its daily
variation greater. Streams and swamps dried up rapidly, and by the
end of the first week in May the prairie grass was often dry enough
to burn.

Rain fell only twice during the stay of the expedition at Lusambo.

By referring to this synopsis of the weather encountered by the
expedition the climatic conditions under which the insects, described
in this report, were collected may be easily ascertained.

In the case of a post at which a long stay was made the month in
which the specimen was obtained has been specified. To facilitate
the finding of places in the map, the names of localities at which
specimens were collected are always mentioned in the order in which
they were visited.

A certain number of insects were sent to the expedition from
places it left unvisited. In referring to these specimens the name of
the collector and, when possible, the date of capture are given.

A considerable number of flies, collected in the British Colon) • >t
the Gambia* and in French Senegal, were taken, together with certain
drawings and observations referring to them, to the British Museum
on the return of the expedition of the Liverpool School of I ropica
Medicine to Senegainbia in July, 1903.

At the end of 1905 these insects had not been described by the
British Museum authorities. It was therefore requested that they
should be returned to the Liverpool School. While at the Museum. ■>
considerable part of the collection had unfortunately been mislaid

* For a map showing the localities mentioned in the Gambia, see Men t« ' 1 ;

the Liverpool School of Tropical Medicine.

8

but a description of the remaining insects, which were returned to
Liverpool, has been incorporated in this publication, with one or two
exceptions, and these will be subsequently referred to as ocrasion
may arise.

We desire to express our indebtedness and thanks to Mr. 1 V
Theobald and Mr. E. E. Austen, for kind assistance rendered in
the identification of the more obscure insects; and to Profess r (i
Neumann for the identification of the Ixodidae, and also for his
valuable contribution to this group of the Arthropoda

FAMILY CULICID^E

SUB-FAMILY ANOPHELINyE

Pyretophorus costal is, Loew

Localities: Zambie ; Boma (Oct. 8 to 21); Prince's Island;
Matadi (Oct 29 to Nov. 25); Tumba; Wathen; Leopoldville (Dec
*o, .903, to June ,6, 1904) ; Telegraph post No. 4 ; Kitoto ; Yumbie
above Lukolelau. bush; Irebu ; CoquUhatville ; Bamamia ; Luton* •
NouveUe Anvers; Bokanga ; Lisala ; Bumba ; Ya,„b„,, X

MalVa ^ntrFnegbU ; L°kand"; Scndwe:

Lusambo; Lado enclave (Nov. Lentire) ^ ^ !

takend,n *■“ -

mosquito nets of servants sWa ' uroPeans and natives, in the
time, far in the forest itself aU | ' 1 ^ du"ng the da>

aquatic grasses growing atong river 'edge's Tn^ ‘°"nd “““S" lhC
puddles of rain-water Win? nn , . .g ’ ,n SWamPs- »n small clear

Pools used by the natives of the l^^r5011’ ***

r fuu by t rr ngo for .

Congo. It seems to feed f he “Mt common AnopheUne tn the

hefore dawn. m°St fiercely after sunset and again

— - .. -

p Fr°m - «-"• - ^ “

5 a.m. they fed fiercely, 15 or 20 insects attacked one at the same moment, .1
were able to feed successfully through clothing.

At Coquilhatville, early in the evening, these mosquitoes were seen Hying tut
the houses of Europeans where ordinarily none could be found during the da.
time. The parasites of malaria were seen to develope in this mosquito at Homa

In some of the mosquito breeding pools at Leopoldville the larvae
of P. cost alls were completely destroyed by larger, cannibalistic Cities
larvae.

At Leopoldville in December, 1903, and in July, 1905, as well as at
Ponthierville in October, 1904, it was observed that Culicida, were
not breeding in the water of puddles and small streams containing an
amorphous, brownish, apparently vegetable deposit, although ->imil.u
neighbouring collections of water contained many larvae

Pyretophorus marshallii, Theobald

Six specimens, all females, were captured in the localities
mentioned below. They were in all cases associated with P costa/ is.
but may readily be distinguished from the latter by the characteristic
banding of the palpi. It is important to note, however, that among
the long series of P. coslalis there are many intervening forms
between typical examples of the two species.

Localities: — Boma ; Leopoldville (May); Coquilhatville.

Yambinga.

Circumstances of capture: The specimens were taken in the
huts and mosquito nets of natives.

Myzomyia funesta, Giles

Localities: Zambie ; Prince’s Island; Matadi ; Wat hen ;
Kalombe ; Lusambo.

Circumstances of capture:- — Examples of this mosquito were
taken in the houses of both Europeans and Africans as well as in the
mosquito nets of native servants. It is curious to note that not a
single specimen of this mosquito was seen at Leopoldville where .1
careful mosquito survey was made.

Breeding places: Examples of this species were obtained from
larvae developed in the laboratory. It is probable that in the I-1 '
Congo it breeds amongst aquatic grasses along the banks of roer-,
and in pools used for steeping manioc.

♦ Possibly a species of Crcnothrix 011 whose gelatinous sheath iron bcO'im -
deposited as ferric oxide.

IO

Field notes:- Malaria parasites were seen to developc in these
mosquitoes at Lusambo.

Myzorhynchus pnludis, Theobald

Localities: — Leopoldville (Dec. lyoj to Feb. iq< })• Bamu
Island; Bamamia ; Eala ; Barumbu ; Kuniba ; Kasongo (Dec.)
Lusambo.

Circumstances of capture: — Imagines were caught m and

marshes and in natives’ huts. I hey also came on hoard the steamer
in the evening after the lamps were lighted, and at Lusambo were
caught about the lamps on three occasions.

Breeding places : — Adults were bred from larvae taken in marshes
and in the stagnant and overgrown but fairly clean water left in pits
from which clay had been taken for brick-making.

Myzorhynchus mcmritianus, Grandpre

Localities : Zambie ; Boma ; Leopoldville (Dec) ; Banm Island.
Kasongo (Dec.).

fo ^z:zicaf,u — lmagines were caught °n* - >he

graf aDd PUpae Were taken from among thc

fn puddl s o„ elf gCS ^ f™" w..ler roUec.cd

clean ta e l 7 TT ' “d ^ -ergrown, but fa.rly

Cellia pharoensis, Theobald

Localities: — This
mens were caught in
larvae.

mosquito was collected
dwelling-places and some

only at Boma Sped
were also reared from

P^ces: --Larvae were

sr *- “» «%, i. ,

amongst water-grasses
and in a dirty, muddy

seen to develope in this

II

SUB-FAMILY TOXORHYNCHITIN^, Theobald

Toxorhynchites marshallii, Theobald

FEMALE. Head. scales bronzy-blue at the sides and between tin-
eyes; bronzy-green on occiput, black along the nape. Palpi of four
segments; sub-apical and apical segments minute; clothed with
brilliant azure blue scales. Antennae black; first segment black it
the base dorsally, grey at the apex with a patch of dull white scales
pubescence grey ; hairs black. Proboscis azure blue. Thora i Pr<
thoracic lobes dull azure blue, with bronzy-green reflections; me-o
thorax black with brilliant bronzy-green scales; scutellar scales rich
bronzy-yellow ; and there are patches of the same coloured scales near
the base of the wings; pleurae almost covered with dull white scales
Abdomen azure blue with rich violet reflections ; penultimate segment
with brilliant ruby reflections ; anterior half of anal tuft black, the re -a
rich orange ; there are white lateral spots to the first, second, third and
fifth segments ; venter blue with white scales on the fifth segment
Legs dark violet ; coxae with some white scales ; mid and hind femora
dull bronzy-yellow beneath with white reflections, in some lights, and
in the hind tibiae this colour extends to the upper surface on the
inside; first segment of mid and hind tarsi with a dusky white band ,
second segment almost entirely white forming a distinct broad band ;
second tarsal segment to fore legs with an inconspicuous basal band
of dusky white scales ; fore, mid and hind ungues equal and simple

Length 9 to 10 mm., exclusive of the proboscis.

A description of the ? has been given as it has not,
apparently, been hitherto described.* Of the 22 specimens collected,
only one is a male, the rest are all females.

Localities: — Tshumbiri and Coquilhatville.

Circumstances of capture: — Our thanks are due to the Rev and
Mrs. Billington, members of the American Baptist Missionary Union
stationed at Tshumbiri, for a series of these mosquitoes collected in
March, April, June and November. They have also supplied the
following notes on the bionomics of these mosquitoes. “ The adults
fly with a characteristic loud humming; they were frequently caught
in a European's house. The larvae were found in a metal water tank

Mono. Culicid., vol. 3, p, 121, 1903.

12

SUB-FAMILY CULICIN./E

Erehuapodites inornatus, n. sp Xcwstcad
(PI. i, fig. 10)

Head clothed with brilliant silvery metallic scales with .1 patch
of black ones in the centre. Thorax rich brown with two median,
two short lateral, and a continuous marginal stripe of golden yellow
scales. Abdomen black; penultimate segment of the male with tw.
lateral silvery spots ; apical segment, in female only, almost entirely
covered by silvery metallic scales; venter with five white mctalli<
bands. Legs long, black with coppery or bronzy brown reflections,
no paddles to hind tarsi of 3 .

Female.- Large basal median area ol head clothed with flat
black scales and a few isolated metallic silvery ones ; upright forked

scales black, rigid, occupying the dark are; . . , nape with few

narrow-curved, golden-yellow scales; lower third at the sides with flat
black ones. Palpi densely clothed with black scales
lack with beautiful peacock-blue reflections.

obes clothed with flat silvery scales except at base where they are
b ack, there are also a few outstanding yellow scales and three long
black bristles ; mesothorax covered with dark brown

rco ™ tl,e laUcr forlnine . . . two lateral and

zrrrr1 stnpei u,e meiimn ***« «* - . « . . <

wX wlT tOUC nng the mar6,nal Stripe . nud-lul, of . . .

bfac" omn PT °f SilVery melall'C Adored by narrow

r1 ‘obe* with a . . J —

black chaetae and thr^ * ^ b*aCk scalcs ; raetanotum with three
(in type oTltoT °“T golden-yellow aedes

-'very ,ine. Abdemm J™' *ha7f I1’"01'!"'''’ a well-defined
some lights; penultimate ’ bronzy-brown and black in

terminal segment s ^ H3^™1 ""h ^ -K-n

brilliant ^ b venter black with five

terminal band is interrupted iT*! t®rminai silver)

scales, which are continued into 'the'*”*'* ^ " "arTO'‘ 'me °f BoWcn

°r” a rect“gular basal patch hTT^' "g Seenien' whfre ,he>'

green m some lights- tarsi n 1 ^ br°nz>r‘brown « peacock

posterior pan morp °^ous ; knee spots to

anterior and mid pairs

13

Wings clothed with dark brown scales, but in strong lights are nt .<
beautifully bronzy peacock-blue and green.

Length 4’ 50 to 6-50 min.

Male.- Head as in the ? but the narrow-curved golden scale-
are more numerous and extend over the whole of the black median
patch. Thorax: Prothoracic lobes and mesothorax as in the V ;
scutellum rubbed in both specimens, but there are traces of a siimlai
decoration to that which is seen in the ? : metanotum nude
Abdomen slender, cylindrical; black with peacock blue reflections;
antepenultimate segment enormously dilated, with two large, lateral,
silvery patches. Genitalia with the clasps of great length, and the
slender basal segment densely clothed with hairs and scales -croud
segment with a long apical spine.

Length about 4 mm.

Fig. i. Eretmapodites inornatus. Wing ok Female. x 4”-

Distinguished from E. quinquevittatus, Tlieob., by the presence
of narrow-curved golden-yellow head scales, and the absence ••!
paddles to the hind tarsi of the $ besides other important details
Theobald makes no reference to the presence of chaetae and scales <>n
the metanotum of E. quinquevittatus either in his monograph or in
the Genera Insectorum, but they are quite evident in this species and
also in E. austeniiA Theob. ; at the same time they are not traceable
in the males of either of these species, their absence may, however. b<
due to abrasions.

•There is no trace of scales or chaetae in either of the two main whjdl
collected.

t Mono. Culcidae, vol. 1, p. 283 ; this species awaits descriptiot

Localities: — Coquilhatville ; Lusainbo.

Circumstances of capture: Adults were caught only in the bush
near water. On one occasion they were found in large numbers near
a native village. They fed viciously at five in the afternoon ; none of
some 30 males and females caught at that hour appeared to have
previously fed on blood.

Eretmapodites austenii , Theobald

Localities: Coquilhatville; Stanley Falls; Kasongo.

Circumstances of capture :- Adults were twice caught in the
afternoon in European houses. They were also taken in the M
near a native village placed on a small forest -covered island ; thc\ fed
viciously at live in the afternoon.

jasciata, !• abricius

Localities:-.- Matadi ; Wathen; Leopoldville (Dec, ,qo3);
Sendwe ; Kasongo (April) ; Tshofa ; Lusambo

... ZZZZZZT' A"“ — «»

larvfr^d du PlaC‘S! ^du'ts were hatd'«> in the laboraton- from

i- . . -

Stegomyia argenteopunctata , Theobald

* . . *« « —

day time in the thick forest. At Kw' C*Ught onl> dun,« the

numerous about a forest snrino- A M'nout 1 theX werc exceedingly
caught i„ a similar spot at aboiT " . * day

fee^

- - - ■— - ::-r r •

W,,w. M Theobald

Kasongo. ‘ ; athen ' Leopoldvdle (Dec.) . Tshumb.r,

lamps h^d). m th- 'Vternoon and at „ight (,.3^”

T5

Breeding places : — Larvae were taken from water collected in ol«l
pots and tins and from a font containing holy water in a mission
building. Imagines were bred from them.

Stegomyia luteocephala, n. sp. (Newstead)

(PI. ii, fig. 5)

Head yellow. Palpi black with white tips. Thorax brown, with
two large, anterior, lateral silvery spots, a median yellow stripe, .uni
posterior lateral yellow spots ; scutellum white. Abdomen black with
pale narrow bands, terminal segments silvery. Legs black with
silvery spots and white-banded tarsi.

FEMALE.— Head with the large central area thickly clothed with
large, loose, flat yellow scales, gradually merging into smoky yellow in
front ; a narrow silvery-white line to the anterior half of the eves
formed of a single series of broad, flat, closely appressed scale .
between the marginal line and the central yellow patch is a broad
band of brownish-black scales, from which, anteriorly, arise several
upright forked scales ; nape with a few long, thick, straight or slight h
curved, pale golden scales, on either side of which is a group of upright
forked scales, intermixed black and yellow ; lower basal portion with ll.u,
dusky-white scales, the marginal ones forming two dull silvery spots,
sharply divided by a dense black spot. Antennae black, nodes white
hairs black; pubescence grey. Palpi black; tips with long silver)
white scales. Thorax : Prothoracic lobes with flat silvery white - >1« -
mesothorax with a well-defined median line and two lateral spots of
narrow-curved, golden-yellow scales; anteriorly there arc also two
large spots of flat silvery-white scales, and a few silvery-white sc. ties
on the lower margin of the posterior yellow spots; the rest of the
mesothorax with rich dark brown scales ; scutellum with flat silvery
white scales; pleurae dark brown with two large patches of silver)
scales. Abdomen rich bronzy-brown; segments i to 6 each with
well-defined, narrow, basal band of smoky-yellow scales; penult in at-
segment with a large lateral patch, and the terminal segment almost
covered with brilliant metallic-silvery scales; venter with well-defined
more or less triangular patches of metallic- silvery scales narrowing
towards the apex, where they appear as two divergent lines; tin
scales forming the outer lateral angles of the spots projecting .«t the
sides of the abdomen appearing as outstanding scales. Legs hron/v

1 6

blackish-brown ; coxae and trochanters, ochreous ; anterior and mid
femora with scattered metallic-silvery scales ; hind femora with
central anterior band and an apical group of silvery scales, anterior
and mid tarsi with narrow dull-white basal bands to the first three
segments, metatarsal band broadest; hind tarsi with a broad white
basal band to the first, a narrow one to the second, and the third
segment almost entirely white above, basal ly it is not s«. IVing*
uniformly pale brown rather densely scaled, first sub-marginal ell
much longer and slightly narrower than the second posterior

Locality:— A single specimen of this species was caught in the
bush at Kumba.

oiegomyni ai oomarginata ,

(P1- ». % 4)

Head Wack, anterior margin white. Thorax grey brown . plane
grey. Abdomen dark brown with lateral grey angular spots to the
h segment ; venter pure white. Legs pale bronry -brown ; femora
lute beneath ; hind femora entirely so.

witlfdulfbron ^ I** tHe fla‘ SCa'es lmiformly olivaceous-black
1111 dul1 bronzy reflections, with a well-defin^H - . ,

white ones, broadening towards the ba • T

of loose flat dusky-white scales and hel ’ ! * "apal patch

and very short unrio-ht f i- i " °" * ,em a few narrow-curved.

bronzy-black. Proboscis vrith^nzy^^ ***? ***** * nin*

basal half with a median • ^ own and ochreous scales.

with the basal segment ^ °f whitc sca,es A ntcnnac

remaining segments darker^ hajrs d^kT0'"3 Sl’Kment t>ale hrown-

white. Thorax: Prothoraci'c lob T , °W” °r h,ack- P“bcScrnce

mesothoracic scales in front „• ” C 0t led Wlth P«r* white scales .

‘he base of the win Z numero White ; a< «* sides from

whole area is sparsely clothed with n ' '

mtermixed at the sides and n , 7°W'curved dark-brown scales

-‘el.ar scales dul, wtoe^tirrt ?’ **>*

°[ ^ ,mid l°be. and also at the I T? b'ack ones at the base
ateral lobes. Abdomen with the! Tf °f the marSinaI hairs to the
a few dull white sca]es *h ho first four segments brown, s, , -black ;

- - and . venter uniformly

*7

white as far as the end of the fourth segment ; the remaining segments
wanting. Legs with the coxae dusky brown, paler apically, with
numerous flat white scales ; scales to anterior and mid femora smok>
brown above, ventrally the}/ are white ; hind femora clothed with
white scales, except a narrow dorsal and a broad apical anterior patch
of blackish scales; tibiae and tarsi of anterior and mid-tarsi blackish ,
posterior tibiae with a broad apical white band, tarsi wanting. IV mg
with dark bronzy-brown scales, those on the costa much the darkest,
almost black ; first submarginal cell much longer than the second
posterior; posterior cross-vein nearly three times its length from the
mid.

Length about 4 mm.

Locality Only two specimens of this species were caught, during
the day, in a European’s house at Kasongo (Dec.).

Scutomyia sugens, Wiedemann

Localities: Matadi ; Leopoldville (Dec. and March); Kisut

Circumstances of capture : — Adults were caught in the bush at
some distance from houses. They were seen to attempt to bite at
mid-day.

Breeding places : One adult was reared from a larva taken from
a puddle on clayey ground containing clean rain-water.

Cat-ageiomyia senegalemis, Theobald

A culicid with the habits of this mosquito was observed at Banana
No specimens were taken, but it is believed, from a hasty field
examination, that they were referable to this species.

DUTTONIA, nov. gen. (Newstead)

FEMALE. — Head clothed principally with flat scales, with narrow
curved ones behind and a row bordering the upper half of the eye'
upright forked scales on the dorsal area. Mesothorax with smal .
narrow-curved, and minute hair-like scales; scutellum with flat scale--.
the lateral lobes small , somewhat tuberculatc , and furnished
eight bristles. Palpi short. Fork cells long, almost equal; lateral
vein scales long and narrow, but broadening towards the apex 1 >f " '
wing.

MALE. — With the anterior tarsi subchelate.

B

1 8

This genus is apparently related to Aedimorphus (Theobald . but
differs in the presence of narrow-curved border scales to the eyes and
the character of the lateral lobes to the scutellum I lie number of
bristles to the small lateral lobes are also unusual ; and moreover therr
is a complete absence of flat scales* to the mesothorax

Duttonia tarsalis, n. sp. (Newstead)

(PI. ii, figs. 6-8)

Head grey. Thorax rich dull orange-brown with two median pah
lines. Abdomen dark brown with silvery -white basil band** . apical
segment almost entirely white. Legs pale ochreous yellow. ale*- n
tibiae and tarsi pale brown, femora faintly speckled

Male. Lateral and median flat scales of the head dull white the

Fig. a Duttonia tarsalis. Wing ok Walk.

narrow-curved scales pMe^ello * ^ ^ Pale yellow ; ,,aPal gr°up <
forked scales in W vP.l ’ " S°me Ughts al'"<*> « >>"<• • «prig!
PalPi of four segments black’ lalFral ones vollov

°chreous with dark brown 7;“' " furnis!’<-H ** fa* . rs. ,,

hand. Proboscis dark b SCa'e* and ln<lications of a fain, media
nodes dark brown ; hairs °f *”•<**•' dusky white

rich o range-brown, f with a few^auT 7\ ^ b,Wtt Titr* ' «*“

scales. andTfew goId^<^rrr . 7"*

, ’h, . g n-ye"ow ones m fron

Culicid., Vol in

t One of four raales hag th*S‘hose ™ the scute! h,,* * ^ °f *m*"

morax dark brown

i g

and towards the scutellum ; the yellow scales also form two lateral spot'
in front of the wings ; there are traces also of a lateral white line
extending from the wings to the front of the mesothorax ; scutellum
with flat white scales ; pleurae dull ochreous with patches of white
scales. Abdomen very pale ochreous, with dark brown and dull
ochreous scales intermixed ; basal bands of pearly- white scales ;
bands 3 to 5 broader than the rest; terminal segment almost covered
with pearly-white scales ; venter pale ochreous with broad indefinite*,
basal, dusky-white bands. Legs pale ochreous, speckled throughout
with ochreous-brown scales ; anterior tarsi with the terminal segment
subchelate; base considerably dilated, ventrally concave; base
with four apparently bilateral spines, apex with a short curved
spine, and immediately anterior to it a long plumose hair with a
bulbous base ; ungues subequal in length, the larger with a long
central tooth, the smaller with a short basal tooth; mid-ungues
unequal, the longest simple, the smallest with a central tooth ; hind
ungues approximately equal, simple. Genital armature with the basal
segment of the claspers furnished with numerous long bristles, two of
which, near the inner apical extremity, are narrowly lanceolate ; second
segment broadly dilated at the apex with an outer, curved, spine-1 ike
projection; terminal claw small, slender. Wings sparsely clothed
with small brown scales ; first forked vein narrower and slightly longer
than the second posterior.

Female. — Head dark brown ; median flat scales smoky-brown,
lower laterals forming a relatively large white patch ; median narrow -
curved ones yellowish-white, frontals white, marginal series to eyes
pale yellow ; upright forked scales numerous, apparently all black
Antennae very dark brown, with numerous minute, loose, flat, white
scales, giving the segments a mealy appearance. Palpi pale brown
with darker scales; tips with dusky white hairs. Clypeus very dark
brown with minute, scattered, white scales like those on the antennae
Eyes yellowish-brown with two to three black spots. / fwrax as in
the $ , but with a well-marked series of white scales in tront of the
mesothorax. Legs with patches of white scales on the coxae ; underside
of hind tibiae clothed with white scales, with a broad apical white
band ; the legs are also more uniformly coloured and browner than m
the 3 , but the upper surface of the tibiae are speckled. Abdona-n
clothed with dark brown scales with paler ones at the articulat " 0

20

venter with lateral white spots ; median area greyish

Length 4-50 to 5 mm.

Taken in thick bush at noon, October 4th, 1 904

Locality : — Kisui.

Duttonia africana , n. sp. (Newstead)

Head with the palpi and proboscis black. Thorax dark hr >wn in
front, paler behind. Abdomen dark brown, paler at the articulations
Legs smoky-brown with pale knee spots; apex of hind tibiae with
a white band.

Female. — Head clothed principally with flat scales, median and
frontal ones blackish, those at the side with dull bronzy refledl OS
below these there is a narrow curved band of bright ochreous ones

aTr*cana.

numerous and scattered over h u , '***'• Uprieht forkcd “=
Antennae dark brown : bZ *lWh°e ot dorsal area, bl.
darker at ,he sides ' pJjV™ ^^-yellow a, the a,

int°SUS d°thed With bIack scaIes°rtL a"d cxtren,c|y 1,s

front, paler behind ; prothorari T*ora* lining, rich. dark br,

™ !ca,ea ™d black bristle, m ^ With la^. white, narr
hair-hke scales, and a few lamer n'a meS°n°tum «’>th minute, black

are alS° numerous black brisdes ye"OWish **** • >'

at the S,des at|d on the dorsum in

21

region of the wings ; scutellum trilobed ; mid-lobe with small, flat, dark
brown scales, and traces also of a few white ones ; lateral lobes small,
almost tuberculate, furnished with eight long bristles, and a patch of
flat dull white scales. Abdomen clothed with dark brown and almost
black scales ; venter with broad, pale, basal bands interrupted by a
median ochreous line which broadens towards the apex Legs
uniformly brownish-black ; femora pale beneath ; knees and tibio t.ir^.il
articulations pale; hind tibiae with a broad apical white band
Wings with the first submarginal and second posterior cells about
equal in length ; posterior cross-vein about its own length distant
from the mid.

Length about 5 mm.

Locality: — A single ? taken in a water-closet at Kasongo, April
7th, 1905.

Culex albitarsis, Theobald

Localities: — Boma ; Kasongo (Feb.).

Circumstances of capture: — Adults were taken during the day in
the open, and at night in a European’s house.

Breeding places: Larvae taken from swamps produced adults in
captivity. The pupa of this mosquito is particularly large.

Culex annulioris, Theobald

All four specimens of this species were obtained from larvae caught
in a grassy swamp near Leopoldville.

Culex dissimilis, Theobald

Specimens were caught in a tent near Tumba, October 26th, 1903

Culex fatigans, Weidemann

Localities: — Boma; Matadi ; Wathen ; Leopoldville (Dec);
Mswata ; Irebu; Coquilhatville ; Bamamia; Lisala ; Basoko ; Stanley
Falls; Kisui ; Kasongo (Feb.).

Circumstances of capture: — Imagines were taken by day and by
night in the dwelling-places and in the mosquito nets of both
Europeans and negroes. They were also caught at noon on ste imer^

22

and in thick forest at some distance from any village. At night they
were often seen on board the Expedition’s steamer.

Breeding places Larvae and pupae were found breeding m old
tins, in water reservoirs, amongst grass at the edges of rivers, in
swamps and in the foul-smelling pools used for steeping iiinni'ii

Field notes: — Enormous numbers of this mosquito, together with
Mansonia uniformis and its var. afr nanus, were found during tin-
day time in the prison at Bomba. Fliey rested thirkh dustt rod an
motionless on the whitewashed walls of the cells, • few
inches above a drain used as a urinal by the prisoners; scared)
an) of these mosquitoes were seen on the neighbouring tarred portions
of the wall. No breeding places existed within 200 yards of the
prison.

( -ulex luteol literal is, Theobald

Localities : Boma ; Bantu Island; Kasongo (March); Lusambo.
ircumstunces of capture. Specimens of this beautiful little cuiex
were caught in the early morning and afternoon, in the forest, in cofftt

P an ations and in the houses and mosquito nets of native servants
I hey attempted to feed but rarely.

at thztdlls ~'LarJaf Were -aken fr0m amongst tbi
obtaining irnagineXm them ””*** W* ******* ”

i neobald

Localities : Miambwe . Lusambo

JZ »::;{t;TTTAdults were takcn « •** bUsh.

and at a few hundred yards from the nearest village

Culex th*lassius, Theobald
from a tub on from a ^ tak

Localuies:- Bo

Yambinga; Kasongo ‘ (ApriHo“eb <Det); Tshorabir

6 J,cu">stances of capture ■ 1 ’• lanihwe 1 Lusambo.

rnarstms, during the day time on CaUght in fores,s »'

“ n,ght m the houses of Europeans" steamer a,

23

Breeding -places: — Larvae and pupae, allowed to develope in the
laboratory, were taken from among the grass at the edges of rivers,
from swamps, from the foul water of stagnant puddles, from collections
of rain-water in old tins, and from the foul-smelling pools used for
steeping manioc.

This is evidently a variable species. None of the females agree
with the typical form in the specific coloration of the head and thoracic
scales. In the majority of the females, which agree best with
Theobald’s* description, the narrow-curved head scales are creamy
white in colour with, in some lights, a trace of pale yellow,
especially in the frontal and large median ones ; and the thoracic
scales are smoky-brown with metallic reflections, almost golden in
some lights, with the distribution of the yellow spots and lines as in
the type. In addition to these variations, five of the females also
possess a distinct median band to the proboscis and other distinctive
characteristics. These forms are described below as a distinct variety

Culcx tigripes, var. consimilis, n. var. (Newstead)

FEMALE. — Head with the narrow-curved scales creamy-white, in
some lights with a faint trace of yellow ; upright forked scales black
with smoky-grey tips; median hairs golden yellow. I horacic scales
chiefly of a greyish colour, with dark brown ones intermixed
Proboscis brownish-black with a broad and well-defined median band
of pale ochreous scales; labella pale ochreous-brown. Palpi black
or brownish-black with a patch of pale bright ochreous scales at the
articulation of the first long segment ; apex with dusky white scales
Abdomen dark brownish-black with scattered pale brown scales and
narrow basal bands; basal segment with two median black apical
spots ; sixth and seventh segments with two lateral apical pale spots
Legs as in the typical forms.

Length 67 mm.

Localities: — Tshumbiri (July); Kasongo (Feb., April, May),
Yambinga (Sept); M iamb we ; Leopoldville. The example from
Leopoldville was bred from a larva procured in a grass swamp near
the terminus of the railway. It is important to note that three female^
and four males of the more typical forms were bred from larvae
obtained at the same time.

* Mono. Culicid, II, p. 34, 1901.

24

Culex viridis, Theobald

Localities: — Boma; Wathen ; Leopoldville (Dec., 1903) , Coquil
hatville ; Miambwe ; Kasongo (Jan.) ; Lusambo.

Circumstances of capture: — Imagines were caught during the d.
in the bush, in the mosquito nets covering the beds of " boys" in
houses of Europeans and Africans, and in the state rooms of a
steamer on the Upper Congo.

Breeding places: -Larvae and pupae were taken from among the
grass along the edges of rivers, in swamps, in dirty, foul smelling
collections of water, such as pools used for steeping manioc* and in
wa ei collected in old tins. Imagines were reared from these

have-T^^T'' ThiS mOSquit0 has U*‘'" «5peatedl> observed to
have a distinctly green colour during life.

Cnlex laurenti, n. sp. (Newstead)

brown ad Center XT if™*' ab,domen Pal« i 'eg* uniformly pale
unhanded. P ei.rae, pale, dull orange-brown;

■n rfr^::h trrr* . .

black. Antennae dark brown T 1 " m<? : upnKl" fotked one* all

with a faint yehow Unee " P,„ “ -'very-grey, so,nc lights
towards the apex a few scaX ” ? ^°thed W1,h bro«n scales, and
sb°rt ' -eh ochreou3 °nps «■>

golden-yellow; pleurae with a few * n°"n' 7 /torac“ scales dull
Whlte scales, some of which are l , T , °f narrow«v-rounded

uniformly pale brown. Legs umfo ' 1 U SpnU,k% sh',»,r,i ' •

-a.es, silvery-grey beneaX TZZZZ*

['C1 ro'vnish-yellow with dark brown l* * ’ COXi,e an<i trochantae

banf to tarsi. br°Wn SCaI^ ! no knee spots and no

Male. — Head sc 1

bZT/lth Pale bro^nSscales aptd‘ ***** *°ng* Pale ochreous
al the Sldes, posteriorly.' i ,l ''U<1 1111,1 femora wil

°»ien pale brown with

25

rather dark brown scales; segments 5, 6, 7 and 8 with a few lateral
white scales ; terminal segment with a broad apical band ol cream)'
white scales ; venter pale ochreous, scales pale brown and white
intermixed. Wing scales dark brown on the basal half of the veins ,
apical portion with pale ochreous scales.

Locality: — Leopoldville (Oct. and Dec., 1903).

Two males and one female only were taken.

This species is dedicated to the memory of the Botanist, Profcssoi
Laurent, who died while on an expedition to the Congo Free State

Culex par, n. sp. (Newstead)

(PI. i, tig. 11)

Head and thorax grey. Abdomen dark brown ; fifth and sixth
segments with lateral pale spots; venter greyish, dark brown bnsally
Legs brown ; femora and tibia freckled, the former much more so than
the latter; anterior and mid tarsi with three narrow pale bands at the
articulations. Proboscis with a broad median band.

FEMALE. — Head with the narrow-curved scales creamy-white .
upright-forked ones pale ochreous, with a few very dark brown ones
at the sides; flat scales few' in number, creamy white. Antennae
wanting. Palpi short ; apical segment minute, clothed with dark
brown scales. Proboscis dark bronzy-brown with a broad median
creamy -white band ; labella paler. Thorax dark brown ; scales chief!)
pale ochreous and creamy-white with a few' black ones intermixed
posteriorly and on the scutellum, the black ones predominate .
there are also indications of four small equidistant black spots
Abdomen clothed with very dark brown almost black scales; a few
isolated dull creamy-white ones on the first and second segment, a
small lateral apical spot on the third, a large spot on the fourth and
fifth ; remaining segments rubbed ; venter with a median pale
line ; last two segments with dull white basal bands, very
pronounced on the penultimate segment; the remaining basal
segments are not sufficiently clear to determine. Legs uniformly pale
browm or ochreous-brown ; knee spots faintly indicated; femora and
tibiae with pale freckles; articulations of front and middle tarsi with
three narrow pale bands ; hind tibiae somewhat paler than the rest
Wings with pale and somewhat metallic-brown scales; fork cells

26

with their bases almost opposite; posterior cross vein .ilx.ni . r mi
a fourth times its length from the mid.

Allied to C. thallasius (Theobald), but differs in the absrncr i
basal abdominal bands, the broader band to the proboscis, tin- pre
dominating pale upright forked scales of the head, ami also m tl.r
colour of the thoracic scales.

Locality: — A single ? taken at Tshumbiri on ful\ ;

Taeniorhynchus fiiscopetmaiiis , Theobald

This mosquito was caught only, near marshy ground, in the bush
at Lusambo.

i in nun

njnvmt* unneiiu , i iieobalcl

Localities: Wathen ; Coquilhatville ; Kalombe ; Lusambo

Circumstances of capture:- This mosquito was Only Ought in

bush, and as a rule in the neighbourhood of marshy ground Its
larvae were not seen. 3 k

Taeniorhynchus cm rites, Theobald

mCbu ;.Kalombe • Lusamb<

the bush, and as -> ml • +1 s mosquito was only caught in

°f —>■ *«•

Taeniorhynchus tenax, Theobald
oca Hies , Boraa ; Leopoldville (Dec ) ■ I k.

Circumstances of capture ■ } ' Ll,sa,nbo

marshy ground. ' Adults wcre caught in the bush near

Breeding places •— t

grasses at the edges of rZTs and CO“eC,ed fr°m amonE thc

m the laboratory. “d from ^amps, produced imagines

Localities :—.Qom • p'l'a “’"S0r""s’ rheobald

Ba7.IsIand; Lnlangl' ^ ° ^ ^ Leopoldv.lle (Dec,
Llrcunis/ances of

“J ™""d» ‘•Jrjz*.

27

Mansonia unifunnis, var. africanus, Theobald

Localities: — Zambie ; Boma; Prince’s Island ; Tuinba; Wathen ;
Leopoldville (Dec. and May); Bamu Island (Jan.); Telegraph Post
No. 4; Tshumbiri ; Bolobo ; frebu ; “ in the bush above Lukolela " ;
Yumbie; Bamamia ; Bumba ; Basoko ; Kama; Benaburungu ;
Nyangwe ; Kasongo; Kalombe ; Tshofa ; Lusambo.

Circumstances of capture : — This species is by far the commonest
mosquito in the Congo. It is seen almost everywhere in the bush,
and lives equally well in grass country or forest. It always bites
fiercely and feeds with equal readiness by day or by night. At Bamu
Island, near Leopoldville, it was actually observed to pierce through
canvas-seated chairs and even through the soft goat skin leather ol
Madeira boots. It was noted in the diary of the Expedition as \er>
unusual that no species of Mansonia were seen in the bush where
camp was pitched for the night on the banks of the Congo a short
distance above Lulonga.

Adults were caught in the houses of Europeans and Africans in
the day time and at night. They were taken from mosquito nets
covering the beds of natives and whites and were raptured during
the day on steamers, in the bush and on sand banks.

Breeding places: — We were not successful in rearing the imago
from the numerous larvae we had from time to time in the laboratory

Melanoconion rimns, Theobald

Localities: Boma and Kasongo (Dec.).

Circumstances of capture:- The specimens in the collection were
all bred from larvae.

Breeding places: Larvae were taken from among the aquatic
plants at the edges of a river and from a disused pit, in a brickyard,
filled with water and overgrown with weeds.

SUB-FAMILY AEDEOMYINjE

Aedeomyia squammipennis, Arribalzaga

Localities: Boma; Leopoldville (Oct.)* ; Yambinga.

Circumstances of capture : The specimens in the collection w< :
bred from larvae or caught in the evening on board a river steamci

* \'o specimen ; recorded from field notes.

28

Breeding places : — Larvae were taken from a grass grown pudd <
of clean water.

Uranotaenia balfoun, Theobald

A single ? of this beautiful species was bred at Kasongo.
December, 1904, from a larva taken in an “old brick pool 15 feet
across." Theobald* states that the mid lobe of his type was
ribbed, in this specimen it is clothed with scales. It may be important
to add that the scales at the base of the wing are of a delicate pale
blue ; but in certain lights appear white as described in the typ<

Mimomyia uniformis , Theobald

I his mosquito was observed only at Boma ; the specimens in the
collections were hatched in the laboratory from pupae collected in a
papyrus marsh and from among aquatic plants at tin- edge of a

bield notes:— The pupae are yellow and possess very 1- mg and
conspicuous respiratory siphons.

Mimomyia africana, n. sp. (Newstead)

(PI- i, %. 4)

Uniformly dark bronzy-brown; legs paler

H AT TS 7 r » _ 0 r

Female.— Head very dark br
paler brown s ’
ochreous, base
clothed with
brown. Thor,
bristles and n-,
cuticle ; scutel

very dark brown, completely covered with flat

unfortunately, has the

Apart from this.

P- *2. pi. vi. fig. 6.

29

however, the insect possesses well-marked specific characters and can
easily be recognised from any other known species by its uniform
dark brown colour.

Locality : — Taken at Nouvelle Anvers, on August 14th, 1904

Mimomyia malfeyti, n. sp. (Newstead)

(PI. i. figs. 1-3)

Head dark brown, with pale ochreous scales. Thorax dark brown,
with bright blue reflections and clothed with long backward -curved
bristles; pleurae pale ochreous. Abdomen brown; apical segment
paler ; venter ochreous. Legs dark brown, with pale ochreous femora
Wings with a pale spot at the base.

FEMALE. Head brown, covered with rather loose, flat, cream
coloured scales and a napal group of black, upright forked one
Antennae deep brown; basal segment paler. Proboscis swollen
apically, but much less so than in the 3 > dark brown ; labella, dull
ochreous. Thorax dark brown, shining, with blue reflection s in

Fig 4. — Mimomyia malfeyti. Wing ok Femalk. x 4°

certain lights , but this character is much less evident under the
microscope than under a pocket lens ; hairs dense and long, especially
at the sides above the pleurae ; there are also one dorsal and two,
more or less distinct, sub-dorsal rows of shorter hairs between which
is a double series of minute, narrow-curved, ochreous scales,
which are very difficult to see and are often wanting ; scutellum with
four or five narrow-curved, almost hair-like, black scales f,lten
wanting; metanotum pale brown, nude. Wings with brown scales
and a basal, pale, nude patch ; median vein scales in a double row mi
the subcostal and first longitudinal vein for two-thirds of the basal

30

portion ; in single rows on the remaining veins ; outstanding scales
present on all but the sixth vein, but arc most numerous on the apical
half of the wing ; first submarginal and second posterior cells about equal
in length, the former slightly the narrower ; posterior cross vein a little
more than its own length distant from the mid cross vein, llaltere-
creamy ; knobs clothed with flat brown scales. Abdomen unhanded,
pale brown or ochreous, with somewhat scattered, flat, brown scales
which give it a more or less mottled appearance ; apical segment
paler ; venter ochreous, with four or five dark marginal triangular spots
Legs uniformly brown with pale reflections, except the ventral and

basal half of the femora, which are pale ochreous ; all the articulations
are pale.

Length 2-50 111m.

5- —M imotnyia maljcyti.

'Vino ok Malk.

31

This species comes very near to Mimotnyia uni for mis (Theobald) ;
but differs in the colour of the head scales, in the presence of a double
row of vein scales to the costal and first longitudinal vein, and in the
thorax with its regular series of bristles and bright blue reflections,
and also the mottled appearance of the abdomen.

The peculiar proboscis and palpi of the 3 are characteristic also
of M imomyia nniformis.

Locality : — This mosquito was observed only at Boma ; the
specimens in the collection were hatched in the laboratory from larvae
taken in a grass-grown puddle of clean water.

This species is dedicated to Major Malfeyt, Ilaut Commissaire
Royal, in recognition of constant courtesy and assistance rendered
during the expedition.

POSITION UNCERTAIN

N eomclaniconion* pulpalc, n. sp. (Newstead)

(Rl. i, figs. 7- 9)

Uniformly brown. Antennae densely plumose ; hairs smoky
brown with the apical portions grey; segments clothed with minute
white scales ; nodes black.

Male. Palpi long, dark brown, with a few scattered blackish
brown scales; apex faintly clavate and densely clothed with long
brown hairs, more especially so at the sides, where the)' form a long
continuous fringe. Proboscis straight, pale brown, clothed with dark
brown scales; apex extending to base of apical segment ot antennae
Central area of head clothed with narrow-curved, pale golden-yellow
scales, intermixed at the sides with long, flat, dusky-brown scales ; sides
with flat dark brown scales intermixed with a few yellow and dull
cream-coloured ones ; nape with a few small, upright-forked, black
scales. Mesothorax (partly denuded) and scutellum with narrow
curved pale golden-yellow scales ; metanotum nude ; prothoracic lobes
denuded; pleurae with small dull cream-coloured scales. Ilaltercs
pale ochreous basally, knobs pale brown, with a few small, flat, brownish
scales. Abdomen , where denuded, almost black; scales pale brown
with faint, dull, greenish-blue and dull coppery reflections; two well
defined sub-median, triangular, black spots at the base ol the third

* Theobald. M.S.

32

segment, and there are traces of similar markings on the second and
fourth; fifth and sixth, with a basal band of pale yellow . \< nt. r. ,r
the base, with pale golden-yellow scales in the centre, dull ore.uny ones
intermixed at the sides; the remaining segments are rendered
invisible by the curved condition of the abdomen Wing scales
uniformly brown, darker on the CQSta ; posterior OOSS VI
and a halftimes its length distant from the mid cross-vein ; first sub
marginal cell scarcely longer than the second posterior, the 111
the veins widely divergent; outstanding scales tong. DUIOW, -.h -
parallel, apex convex ; frmge scales long, lanceolate. Legs uniformly
rown; postenor t,b,ae with a pale apical band; tarsi dark brawn

under ‘, OUt2'5° ‘° 3 The ha* the abdomen cemd

unde , so that * » ^possible to give the exact meuurament

island Mow n 7 7'* ^ {r0n‘ 8 W taken in a pool on »
island below Basoko, September 3. 1904.

'-'"'•tiamcotuon palpate.

**-*»«..

it (^1- i. figs. t;, (,)

n,argms Am—

black' a,bcll!1 bright “uhreous Th'o 5”“ered brigrlit ochre
second P°a the a"terior pair bJJ ”***?***■ ochreous with f

band “ dSma"er 1- * Sga:dUCs thC 7BC" = W’i"d

Legs SD" ;,men bI«<* With a dorsa? ’eWhat '" defined bit

.... ■** ,z * g;tr:

33

FEMALE. Head with both black and pale cream-coloured upright
forked scales, the latter predominating ; anterior portion with rather
large, narrow-curved, bright fulvous scales, and apparently a few long
flat ones, intermixed with the former are a few black ones tipped with
bright fulvous ; sides with flat, somewhat loose, cream-coloured scales.
Antennae dusky brown; nodes pale. Palpi densely scaled, black,
dorsally with white tips. Proboscis swollen apical ly ; scales black
intermixed with a few cream-coloured ones ; swollen portion entirely'
black ; labella bright ochreous. Legs with the anterior femora pale
ochreous with a black apical patch in the centre of which is a distinct
central crescentic band ; mid and hind femora black, the former with
two, the latter with one yellowish band ; tibiae black, each with a sub
apical but somewhat diffused band ; tarsi black with bronzy brown
reflections, each with five distinct yellowish bands; ungues equal,
simple. Wings with bronzy-black and cream-coloured scales; basal
half of costa with a long black bar and two irregular spots; all the
nervures with irregular groups and isolated black scales ; fringe pale .
costal scales bronzy-black, becoming paler towards the apex ; out
standing scales claviform ; a few of the black ones, especially those on
the sixth vein, heart-shaped. Halteres cream-coloured with dark-
brown scales at the tips. Thorax with narrow-curved, golden-yellow
scales ; four literal spots of black ones, the anterior pair arc
the largest, and there is a broad indefinite band behind
the second pair; scutellum with pale ochreous narrow-curved
scales ; metanotum black, with three dusky-yellow lines, and
a basal, linear, patch of minute, flat, cream-coloured or white scales
prothoracic lobes and pleurae with flat, dull, cream-coloured scale-
Abdomen clothed with black scales, having brownish coppery reflee
tions ; each segment with a long narrow median, basal, patch of dull
cream-coloured scales; apical segment only with two lateral pale
spots ; venter ochreous.

Length 375 mm.

Locality: Boma ; bred from a larva which was caught in .1
marshy pool.

BOYCIA, nov. gen. (Newstead)

Head with a median area of narrow-curved scales, flat, loose,
lateral ones, and numerous upright-forked ones. I borax and

c

34

scutellum with narrow-curved scales. Wings with the lork '•11-
relatively short; anterior forked vein slightly shorter than the first
posterior; scales resembling those in M imomyia. Palpi short in the
5 , long and clavate in the 3 . Proboscis swollen at the tip in both
sexes. The narrow curved scales in the mid region of the head are
fewer in the ? than in the 3 ; and iti the former the central "ties are
arranged in two distinct lines, but are almost completely hidden h\
the numerous upright-forked scales.

This genus is somewhat difficult to place, but agrees licst with the
group of Culcinae in which the palpi of the males are swollen .it the
tips.

noycui mitnomyiaformis,

(PI figs. 1-3)

Head greyish. Mesothorax grey-brown, with two anterior spots

?K,a roa transverse band opposite the insertion of the wings, bfau k

i onien ac ’ Wlll‘ P:ilr narrow basal bands; vein

certairTl^li f °chleous’ Legs brown; base of femora and tarsi (in
- tain lights) paler. Proboscis swollen towards U

white scales thn^^ a!Ctl °f Wltl' na,T"u rur'(',i

intermixed with thele Ire f°n”'nK lwo '°ngit..dinal series,

extending almost to L f TT “Pr«h‘

spatulate^ creamy scale! n" h“d 5 “<*“"*»* ** «*-

scales; all the fori i i ° ' numerous upright forked, black

ntarked dentil ^ ^ d"^d with, usua.ly. five well-

rale golden-yellow' scales narrow -curved

band of black ones • the 1 U'° tinter'or spots and a broad transverse
some specimens, the ^ COi«tricted in front giving it, in

scutellum with large ^ °f tWO confluent spots;

lateral lobes with golden reflet* ’ scales, those on th<

creamy-yellow. Abdomen clothed , metanotum nude* ; pleurae
appearing bronzy-purple Jn sq Wlth dark brown and black scales.

Pale ochreous scales ; venter Dal tf** an<l n;,r" ’" basal

rown. spots. Legs unbanded, bronJ^b^' ',n-'lLu

7ZT _ _ y rown ; femora pale ochreous-

l\vn i . - — — - —

from thrmSXixh°W a sin8,e curved sc^T^ th„e

are apparently loose ones

35

brown at the apices, forming pale knee spots ; tibio-tarsal articulations
pale ; last three segments of tarsi bronzy-ochreous.

Length 2-50 mm.

Male. Head with a well-defined median, long, angular patch of
narrow-curved creamy scales, no regular series as in the female ; sides
broadly clothed with large, rather loose, flat, cream-coloured scales, and
a rather large lateral patch of brown ones ; a few short upright forked
ones at the nape, black, and a few, very short, creamy ones towards
the front of the head. Antennae densely plumose, Iiairs grey with a
few black ones intermixed. Palpi strongly clavate, with a pale
cream-coloured band at the anterior end of the first long segment
the remaining portion of the segment and the basal half of the
succeeding one with very dark brown, almost black scales; apical

I'lG. 7 .—Boycia mimomyiaformis. WlNG OK FEMALE. x 40.

segment with short stiff yellow bristles. Proboscis swollen apically,
dark brown; labella paler, especially at the tips. Mesothorax with
the narrow-curved scales dusky yellow intermixed with grey and a
few black ones, and there are two ill-defined lateral black spots in
front of the wings ; space between the wings denuded, dark brown ;
pleurae and legs as in the $. Wings with pale brown scales;
anterior forked vein slightly shorter than the first posterior, the latter
much the widest; both are relatively short Abdomen as in the ?
but the bands are a little more pronounced.

Localities: -AH our specimens of this mosquito were bred from
larvae caught at Boma (Oct.) in a papyrus swamp and among the
aquatic plants of a small stream.

FAMILY CHIRONOMIDiC

Ccratopogoti sp. ?

This fly was only twice observed, once at Binza (I3cc). near
Leopoldville, and again near I umba. In these localities it was the
most common bloodsucker observed. Although it is almost invisible
its bite raises a large weal which itches for hours

Chironotniis sp. t

The specimens which we obtained <»f this insect were bred tr m
larvae, at first sight resembling mosquito larvae, caught in a swamp
at Boma. This insect was also seen at Leopoldville

FAMILY PSYCH ODIDiS*

Phlebotomus n. sp. (

d lie only specimens which we now possess are not sufficiently well
preserved for descriptive purposes.

Localities: Leopoldville (March) ; Wanie Numbu.
t ircumstances of capture: Specimens were taken in the daytime
in the bush and in rest houses for Europeans.

Oi ivi U nu/h

Sirnulium damnosum , Theobald
Matadi ; Lutete ; Leopoldville (Nov.) ; nen

Local ilies
Miambwe.

catvhtb ‘othnT6* °a ca*ture: Specimens of this species w«
near, and at some distance from, water. They bite free!

Simuliunt sp. ?

Bat^r Le°poldville (No*- >903. to Feb.. 1904); M'SwaU

and far from, water flles wcre cauBht both nei

often found in houses. hif'T5 ,°CCUr “ S"arms 30,1 3

attacks, and when crusbf»rt 'C ree y’ are persistent in thf
crushed emit a peculiar - bed-bug like " odour.

37

HABITS AND STRUCTURAL CHARACTERS OF THE LARVA
OF SIMULIUM*

As the characteristics of the larvae of the European species of
Simiilium are not likely to differ very materially from their tropical
relations the subjoined notes may prove of some guidance to students
in other parts of the world. The notes refer to a single specie-,
Simulium ornaturn , Mg.

Shallow rapid streams fully exposed lo the sun on hill sides,
moors and open spaces in woods are the favourite resorts of these
curious larvae. They are invariably found where the stream flows

Fig 8. — Larva of Simulium ornatum in semi-erect position with the fan-like
or fringed appendages expanded. x 6J4-

most rapidly ; never in the side pools or in comparatively still water ,
and very rarely in those portions of a stream which are overshadowed
by trees or shrubs. In streams where all the necessary conditions are
available the larvae sometimes occur in countless numbers, covering
the undersides of nearly every submerged leaf or blade of grass, and
also, where not too much overshadowed by aquatic plants, on the
stones and fragments of rock at the bottom of the stream.

* These notes were compiled from observations made in I-.ngland and Helgumi
in the summer of 1906. — R.N.

When massed together on the stones and rocks tlic\ look at a
short distance remarkably like a waving mass of dark brown or
blackish algae or moss; but this resemblance is much less marked in
colonies attached to the leaves and stems of plants.

The larva, when at rest, attaches itself to a fixed object by means
of its anal sucker, standing in a semi-erect position (fig 8) with its
head pointing down stream; but in very rapid rivulets the bods
lies almost prone with the object upon which it rests, or the
extremities may be brought together so that the hods forms a distinct

c

R. N. <M.

a

* J t>

9* apnendapes///p/vi'iaW fr,'atum 1 a- head of larva with the fringed
'• portions of x u4° ’ ** Ringed append >ed{

l ^ ^l^v*** uteral x SI,

characteristic nodes (enlarged). S,lke" threads showin* ,he

w,°th ispr°,vided with a of beautiful fan-1, ke process

Unlike the 1 6 ““"c*1 SWeepS particles of fo°d into Us . nil

s~i^ JTZ mOSquitoes they have little or no power c
progression stm^ m'enUns from Place to place, their method c
of the geometridgy,,i?5embleS *hat °f the larvae or looPer raterpiUai

^eadf”ranSdir± theyPr°grCS" f^m place places, Ik,

along which thev °nS’ 0rmm£ an irregular web or networ

N;!th.great — When forobly dh
distances varying- fr t Ct S immediately float down stream t
by a slender silken thread^0 ^UrtGen ,nches* suspending themselve

’ res 1 y sPun» or by the network of thread

39

which may have already existed. Many of them will immediate!)
commence to haul themselves back again by walking along the silken
thread or threads, and this they accomplish by forming the body inn
a succession of loops, in the same way as they progress along stones
and plants — the mouth or sucker feet grip the thread finnh
while the posterior sucker is brought forward until it almost touches
the anterior segments -the action being repeated until a place of
safety is reached. Many, however, float down stream supporting
themselves by their threads until some fixed object is reached. If .1
leaf or stem supporting a colony of larvae be pulled partly out of tin
water the larvae will all crawl back again to the stream and usual l\ fix
themselves to the submerged portions of the same leaf or stem

Fig 10. — Larva ok Simulium ornatum : — a. maxilla and palpus ; b. mandible .
c. labrum ; d. labial plate, x l25-

If placed in still water the larvae soon become sluggish and death
takes place in about 17 hours, but if completely removed from the
stream and kept in a thoroughly moist vessel they will survive for a,
much longer period.

When about to pupate the larva spins for itself a little elongated
cocoon, shaped somewhat like the toe of a slipper, with a large
opening at the broad end, and this is almost invariably placed with
the opening pointing down stream. From this the head of the pupa,
with its external respiratory filaments, projects and a sufficient suppb
of oxygen is thus obtained.

The period of pupation is only of a few days' duration : the
minimum being two, the maximum six clays. On bright sunny days
they begin to hatch as early as 9 a.m. and continue to emerge
during the heat of the day. A feu examples also hatched in tin-
laboratory during the night, but this was apparently an exception to
the rule.

In order to rear the imagines the cocoons should be removed from
the stream immediately they are formed ; and the leaves and stones
to which they are attached should be placed in glass jars Covered nth
ne muslm or chiffon. All that is necessary is to keep the material
perfectly moist ; however, many imagines were successfully reared

rom pupae which had become perfectly dry. (»„ . .

the pupae he left in standing water.

JZSnirr ,°f the imag° the sk,n of »- P«P» spins aloqg
forwa“de3Teilneat°tfhethe th°raX and t"C raPidly P-hes itself

greyish or silvery Sa“’e 8,,rro,,nded h>’ »« which Rives .t a
The moment the in^T ^rlS '^f °f "" ^

carried to the surface wiiPn n • " '

in contact in its rapid courf"? * ***** °^eCt W,t** w^ch it a
foothold instantly takes to win'o :< "o' and ll;,vinK gained a

completely immersed in the T ccas,ol™Hy the insects were seen
either when waling a W \? ^ « at perfect ease

legs and antennae. In such c- ^ ^ ^ P,antS or clea™ng their

the abdomen so that they tapered t & WingS Were fol,l"l partly round
"if1"6 SGen t0 a large air c ♦ * ^ behind’ and this way

observed in still Hus remarkable trait was

m still water only.
description of //te ^

Legs in two, coaiescent, pairs-' thi^ ';yl'.ndrira'. swollen posteriorly,
first thoracic segment and pro.ee r , *5°? *** are *tt»*ed to the
provided with little hooklets • thl "T beneath "le head and are

the a 1 . and fo«S'a^ -ond pair (% M) are p|acfd at

H " ° ,0°klets ^ranged in rP , SUcker whid> * surrounded
- f Head (fig- 9. «) relatively T r transverse senes.
-T^^T^frepresented by two ' lab‘U'n hroadl>' rounded

' I°T - pot! Antennae ^ ft <0

41

slender, almost filiform, of apparently three segments, of which the
first is extremely short, the second about half the length of the third
Immediately below the antennae is a large stout process furnished at
the end with a long fan-like fringe of hairs, with a much
smaller, but similar group of hairs arising from the centre of the lower-
lateral margins of the same process. The hairs forming the large
fans are provided with an inner lateral fringe of delicate short hairs
Mandibles furnished at the tip with two stout and slightly ' lined
spines and long silken hairs. Maxillae each with a long slender sub
terminal spine which is sometimes completely hidden in the dense tuft

K. iV. ,1/1.

f>

Fig II.— Larva of Simulium ornatuni: — a. anal segment; />. papillae, .7 . sinker r r
with its coronet of hooks ( x 45) ; b. anal appendages ( x 451.

of silken hairs. Antennae short, stout, of two segments, the terminal one
being about one-fifth the length of the first. Labial plate broad and
angular; the anterior margin emarginate and furnished with nin<-
strong teeth or spines, the outer teeth bidentate, the median tooth
slightly the longest ; anterior half of the lateral margins strongly
serrate ; and there is a submarginal row of long stiff bristles.

Length of fully matured larva 7-50 to 8‘50 mm.

FAMILY TAB AN ID JE

Heematopota duttoni, n. sp. (Newstead)

(PI. iv, fig. 3)

Face dark brown, clothed with grey pubescence, with two black
spots. Palpi dull yellowish -brown, sparsely clothed with black

4-2

pubescence above, grey at the sides Antennae pale brown* tips Made;
first segment almost cylindrical, the third slightly longer than the fat
and second together. Frontal callus pole yell, wish brown
the eyes, slightly produced in from ; a broad redan*
rich chestnut between the eyes and .1 paler but ,11 d. lm.-.l aU.v,
it in the centre of the callus; paired spots large, black, and slightly
inform; unpaired spot distinct. No markings on the vertex,
/w dark brown with three equidistant tnereoos
Abdomen brown, with golden pubescem

U,th jrather lar?e bla<l< Spots; tlu-

are sp° e ony at the sides with the exception of the two h •

bear traces of small spots on the dorsal areas; scutefem
l ereous. Legs pale brown; anterior tarsi and apical half cf

.p eal t l ird5d PrP,C, brOWn' WingS ' < ■'••-I'.-r.-l ..pprarancr,

apical third darker than the test; upper . .

between the rv -> ,1 ] *lt s<1"1 1,1 •' "

Paired spots not ,‘l . .

Length «rSo mm. ; wing :Tmm °f dypeal SP°P

ocaltties : Nyangwe ; Kasongo ■ Tshof . \l l
Circumstances of capture • Tiv . « ’ M“,nbwt

about cattle. Ils -v was caught along rivers and

Head iv- %• 2)

black, tibiae dusky^hte ^1^ ' abdomen smoky-brown;
e^mdistant, oblique dusky smoky-brown -

Female -Uead. v^Z^ eIon8ated >P«s

black" 1 b>*<=k, shining pJeeP ^

the rest- ^Verse suture at the sides akd ^

.

• anlerior and hind tibiae or I, too

4.5

white, apices dark brown; mid-tibiae entirely ochreous- while ; tarsi
black; first segment of posterior tarsi ochreous, the remaining
segments brown-black ; first segment of mid-tarsi pale at the basal
half, the remainder brown, the other segments are wanting. Wings
narrow, uniformly smoky-brown ; with three costal dusky-white spots ;
the first, about midway between the base and apex of the wing, is
placed below the costa and immediately above the first transverse
vein ; the second about midway between the first and third spots,
forms a continuous oblique band extending from the costa to the base
of the first forked vein ; the third is slightly narrower and extend
from the costa to the hind margin, leaving the extreme 4 ip of smoky
brown.

Length 9 nun. ; length of wing 8 mm.; greatest width of wing
275 mm.

A clearly distinct species, easily recognised by the curious colora¬
tion of the wings.

Locality : — Two specimens were caught in a European’s house at
Yakusu.

H&matopota brunnipennis, Ric.

Localities: Coquilhatville ; Nyangwe ; Kasongo.

Circumstances of capture: -These flies were only caught on the
river.

H czmatopota spp.

Several rather worn specimens belonging to this genus were taken
on cattle and horses at Matadi. Just to the north of the Limposo
river one flew into the carriages of a moving train.

Chrysops dimidiatus, v. d. Wulp.

(PI. iv, fig. 1)

Localities: -Banana (Dr. Etienne); Matadi; N’Kussu ; Wat hen ;
fshumbiri ; Lisala (reported by Rev. K. Smith); Yakusu.

C ircumstances of capture: Specimens were taken both near and
at short distances from water. One was also caught in a European’s
house.

44

A ty lotus iiigroniaculatus, Kic.

This fly was caught in a European house placed on high ground
at the mouth of the Gambia River.

Tabanus canus, Karsch.

(PI. iv, fig. 9)

Localities: Lutete ; Matadi ; Lukolfla; IkdeDlba and Lopori
rivers (March and Nov., Major Malfeyt).

Circumstances of capture: This fly was only taken by , arsons
travelling in canoes. It is reported to fly very rapidly and its bite
is said to be very severe.

I' aba nns dorsivitta. Walk.

(PI. iv, fig. 4)

Mr.Tne„°nly Specimen we obtained was collected by the Rev and
Mrs. Billmgton at Tshumbiri.

(PI. iv, fig. 14)

ITnhieverateof ?bgiVen ‘° S°mewhat marked variation.

Ereen ^ ^ ^ “ “ -t.re absence of the ap

yellow than is seen in the Zf Qthe ^ ^ dCC'd<'d ‘

as transitional between T r • examples may be considc

Austen. foseta/us and the sub-species niloli

Localities : _ . t

(March, Major MalfeytV ^P° d^lle (N°v., Dec.); Lopori R:
Basongo ; and at mam^plar ^°uve e Anvers ; Tubila ; Kaso„f

Chances of It ^ Ga"’b-

Most of the specimens were taken'"5 seen °nly near wa

E,"0eS- Two specimens we e u """ by or

Eur°Pea’«- h°Wev"' caught in the houses

Lie Id notes; _ Wi,,i .

The eyes are a bright| meet^™g this Hy 15 * very beautiful obj<

is oft"7 m“Ch bri^hter than in dr and the Colours of ,he 1,0

" Seen bo-d Steamers over " ““ *"”*'>• a

>ards from the river’s bai

45

Tabanus gabonensis, Macq.

(PI. iv, fig. 15)

Localities: — Lukolela (July, Rev. Whitehead); Ikelemba and
Lopori rivers (March, Nov., Major Malfeyt) ; Baringa (July, Dr.
Angela) ; Nouvelle Anvers (April, Dr. Miiller).

C ircumstances of capture: — Most of the specimens were taken by
persons travelling on the rivers by steamer or in canoes.

Tabanus gratus, Loew

This species was caught in March, 1903, on the Kunchau ( reek,
about 1 75 miles up the Gambia River.

Tabanus par, Walk.

(PI. iv, fig. 6)

A single specimen of this insect was caught on board a steamer
plying on the Gambia River.

'Tabanus pluto, Walk.

(PI. iv, fig. ;)

This Tabanid was caught in the Congo near the river at Matadi
and Kisantu.

Tabanus nijipes, Macq.

(PI. iv, fig. 8)

Localities: Matadi (?); Tshumbiri (April, Nov., Rev. Billing-
ton); Bolengi ; Lulanga ; Lisala ; Stanley Falls; Wanie Numbu ;
Kasuku ; Tshofa; Pania Mutombo; Lusambo.

C ircumstances of capture: — This fly was caught only near water,
usually while attempting to bite passengers in canoes or steamers.

Field notes: — They fly very rapidly and strongly, and have been
seen at over 100 yards from the banks of the river. They seem to be
very local. For example, near Pania Mutombo ten or a dozen were
seen at the same moment darting about a canoe. A little further
down the river towards Lusambo none were seen for miles.

Tabanus tarsalis, Adams
(PI. iv, fig. 13)

1 his insect was only seen once, at Lutete.

Tabanus mimaculatus, Macq.

(PI- iv, fig. 5)

One specimen of this fly was collected in November, 1903, ai
Matadi by Dr. Bourguignon.

Tabanus albovcntralis, n. sp. (Newstead)

Thorax dark brown with three narrow lines; abdomen darker
rown, with a b.Iateral series of oblique white spots. The whole of

IT surface, including the head, white. Legs pale ochreous,
with the tarsi and apices of the tibiae dark brown.

EMALE—Heaaf.. Space between the eve. .

lru bands> ^ . -

pul'L AtZ ** "°r with dense

Thorax- dull erev hr ** ^ C I*™”1* Palpi pore white and wax like

lines; margins greyZ' leTus^h”^ ^ MBdl“

Abdomen dark brown w;th * Posterior margin of I

to the fifth segment with • ' . ,M,r *PicaI 1
greyish-white, the spots grad, J.*": ?bKque* rl' '
extremity, so that the la « ^ ' ' t*|n^,n*sk toward

segment extends right arro S<‘"','l-V vimI)1<'' while that on the SCO
band; there is also 1 hint Z scZn,rt,t- '<rmnui„
yellow hairs; vem« ^ T ft>WMd b> -

pubescence,

oclneous border. Legs wif. , ‘ gments with a distinct, narrow,

f biae pale ochreous with the w T^’ tr°chantae and femora white;

transparent, withoutTarkhigs ' br°Wn * a" dark h"”™

-X r s 1 1

blw, 'w

:lr; ™*2i“ >—■ *,!«, .«

W ap.ca! band. Wings ;lTd fourth segments with a

3 narr™ median dark dgtag

47

line and a broad sub-apical band ; apex with a clear triangular space ;
the anterior basal, and succeeding cell clearer than the rest. Male
Thorax red-brown, covered with grey dust. Abdomen black, with
narrow grey bands to all the segments. Nervures of wing all
margined with brown. Fore tibiae in both sexes white; the

remaining segments blackish.

MALE. Head: Eyes dull bronzy-brown (in dry specimen); ocelli
ochreous ; space between the eyes black, shining, brown below .
clypeus and cheeks grey, the latter with long, silken, white pubescence ,
posterior surface of head dull grey. Antennae black, basal segment
clothed with grey dust, the rest with ochreous-brown pubescence
Proboscis and labium black. Thorax in front of transverse suture,
dull castaneous, with a narrow, faintly-indicated median line, and two
narrower sub-median lines, which terminate in front as two black-
depressions ; margins dusky greyish-brown; suture opposite the sub-
median lines orange-brown, continued downwards as a short narrow
streak ; posterior half of thorax slightly darker than the anterior ;
the whole is covered with a greyish powder, and scanty black
pubescence ; scutellum and pleurae grey, but the latter are paler than
the former. Abdomen smoky-black with narrow grey apical bands to
the segments, these gradually diminish towards the apex, until the)
entirely disappear on the last one ; venter of the same colour and
banded as on the dorsum. Legs black ; mid femora dark piceous .
upper two-thirds of anterior tibiae dull white, the remaining third
black. Wings with the nervures dark brown, all with a broad diffuse
band of orange-brown surrotmding them.

Length 16111m.; length of wing 15 mm.

T EMALE.— Head as in the male, but the face is pale ochreous
Thorax rich dark brown, with two, narrow, sub-median pale fulvus
lines which gradually darken and entirely disappear at the transverse-
suture ; margins with a well-defined band of pale fulvus pubescence
which is continued round the margin of the scutellum, and in front
extends over the pleurae. Abdomen black, piceous in some lights,
basal segment pale fulvus with the fringe of the pubescence paler ;
second, third and fourth segments with a conspicuous narrow' gre\
apical band ; venter with similar bands on the second to the fifth
segments. Legs as in the male, but the anterior tibiae have the basal
half only of a pale ochreous or dull white colour.

4«

Length 17 to 20 mm.; length of wing !<> nun.; expanse of wing
34 to 35 mm.

Localities:— Several females of this species were first H ail t» M
by Rev. and Mrs. Billington, from Tshtimbiri. It has received its
specific name as a mark of our appreciation of the interest shown in
the work of the Expedition by the collectors A female was also
caught, near the river, at Bolengi. The only male wc |hissc'n was sent
to us by the Rev. M. Ave, from near Matadi.

It is an extremely well-marked insect, and is easily recognised from
the other African species of Tabanus both by the banding of the
abdomen and the markings of the wings.

Tabanus spp. incert.

Six additional species of this genus were taken in the localities
named below; but the specimens have all suffered from mould and
other injuries, so that it is impossible either to identify them with any
certainty or to give adequate descriptions of them. One can onl)
add that two of these specimens arc referable to the Social ts group.

Localities: I.uano on the Kwilu River (Major Malfeyt, June;
1905); Matadi; Wathen, Leopoldville (Dec.) ; Lukolcla (Rcn White
head, Aug.); Kuzu, in the region of the Lower Congo

a EuropelnChouse^ TW° °f ^ Specimens werc caughl “

family

SARCOPHAGI DAE

Sarcophaga spp.

Leopoldville (Dec ^ ^re^e"tatlves of th>s genus were collects

Malfeyt); and * Rasongo '(Apll) “‘t' ^ “

d.stnbuted one throughout ^ ‘ ^ *”

Circumstances of capture - A ^

and in privies. Adult* 1 P c,mens were caught in the c

brought by natives hfhed laboratory front la,

Some species are vivin * 01 larvae of Auchnterontyia lute

deposited. Within a very Tew ^ ^ Ver>’ rap

0 ten almost covered with them m,nutes ^reshly excreted faeces

40

FAMILY MUSCIDyE

Pycnosoma marginale, Wied.

Specimens of this beautiful species were collected only at Leopold
ville.

Pycnosoma elara, Walk.

This fly was only observed at Leopoldville. The specimens were
hatched from larvae brought by natives in mistake for the maggots of
Auchmeromyia luteola.

Pycnosoma putorium, Wied.

Specimens of this insect were collected at Wathen. It was not
seen in any other locality.

Pycnosoma sp. ?

The only specimen we possess is not in a sufficiently perfect
condition for verification. It was captured, in company with others,
at Tshumbiri.

Lucilia fuscina, Walk. ?

Localities: — Tshumbiri; Kutu ; Basoko ; Nya Lukemba (Jan.,
Dr. De Maria).

Circumstances of capture: — These flies were caught in the bush
and were hatched from larvae found in the mud floor of a native hut
They are said to follow cattle.

Lucilia spp. ?

Several additional specimens of this genus were caught at Wathen
and at Kutu (Feb.).

Auchmeromyia luteola, Fabr.

(Figs. 12-14)

Localities: — San Salvador; Noki ; (Portuguese Congo, reported) ;
Lukungu ; Matadi ; Wathen; Kimfuti ; Leopoldville (Oct. -June) ;
M’Swata ; Tshumbiri; Lukolela; Irebu ; Bikoro (Rev. Clark);
Bamamia ; Lisla ; Upoto; Bongandanga (Rev. Gamman) ; Basoko;
Loeka (reported) ; Yalembe ; Yandongi ; Romee ; Yakusu ; Stanley
Falls ; WanieNumbu ; Ponthierville ; Kirundu ; Utikakadjia ; Kuinh.i ,

I)

50

Lokandu ; Sendwe ; Nyangwe ; Kasongo (Nov. to April) ; Molemba ;
Tshofa ; Miambwe ; Kabinda ; Lusambo ; Portuguese Angola
(Bastian, interpreter at Lusambo); Lake Tchad (Monsieur ( tievalier
? Lagos (native reports) ; Sierra Leone (reported by Captain Grattan
Circumstances of capture: Most of the specimens were taken
during the day time in or near the houses of natives and Europeans
One or two were caught at night as they buzzed loudly about a
European’s house.

Bionomics : — We have little to add to the description of the habits
of this fly published in Memoir XIII of this School The flies were
again noticed to be attracted by the sleeping mats of natives As
before, the pupa usually took from a fortnight to three weeks t(

develop. The shortest period observed was ten d iperature

not unusually high).

The larva of this fly has already been described and figured* ; but
the mouth parts and stigmata need further elucidation to enable
students to distinguish it from other allied species.

The 8reat ™<>*th hooks. The anterior half of iW

SCh°°' MeSic"'ethc l:Xped' *"

to. Memoir XIII of the Liverpool

Fig i2. — Auchmeromyia luteola (“ Congo Floor Maggot Mouth parts, &c.,
of larva: — a. tip of antenna: md i and 2. the great mouth hooks;
cs. cephalo-pharyngeal sclerites ; hs. hypostomal sclerite ; ct. central
or median teeth ; ps 1. anterior-lateral palmate teeth ; fsj. posterior-
lateral teeth ; st /, 2, 3. anterior or thoracic stigmata ; ds. dermal
spines. (All greatly enlarged.)

FlG 13. — Auchmeromyia luteola. Anal segment of larva : — a. anus ; at. anal
papillae; mf. marginal papillae; dp. dorsal papillae: </ postnior
stigmen. (All greatly enlarged.)

52

and the individual teeth or spines are distinctly curved upwards and
inwards, and none of them arc bidentate.

The anterior stigmata or spiracles {si. /, 3, j ) are branched ; each
with ten to twelve relatively large orifices ( st . j). The posterior
stigmata (fig. 13, st.) are large obconical projections of brown chitinr .
each presenting three darker transverse slits, partially closed by a
fine transverse grating.

The posterior segment (fig. 13) bears along its margin four
conspicuous bilateral papillae {m.p.)t of which the median and third
pairs are much the largest; the second and fourth pairs arc modi
smaller; all are sharply attenuated and hear faint traces of

segmentation.

On the method of the escape of the imago from the pupamum
ie anterior pole of the puparium is invariably broken away by the

imago on its escape, as is normally the case with Other insects ; but is
tour out of five specimens new rw • »l. w r .«.•

R- N. d<!

' ‘4 — A uchmerom

53

a propupa; though why it should do this is not clear. Neither is it
possible with the limited material at hand to say whether the reversed
position of the imago is abnormal or not. It is certain, however, that
the pupa may also lie in its normal position with its head at the
anterior pole of the puparium, as in dissecting out a number of puparia
one imago was found in this position.

Thus we have a very remarkable anomaly ; in the first instance the
true larval stage is continued until after the formation of the
puparium. and secondly, a large percentage of the flies escape back
wards from the puparium.

M usca spp.

One species was taken on cattle at St. Louis, Senegal (May, 1903) ;
another on cattle at Kasongo, and a third at Wanie Numbu. The
specimens have, unfortunately, suffered somewhat from mould and art-
therefore unsuitable for descriptive purposes. Two of the species are
apparently new to science.

A new Genus unci Species of Bloodsucking Fly, allied to Muse a

Several specimens of these interesting flies were caught on cattle
and donkeys at St. Louis, Senegal, May, 1903, and also at Zambie on
the Congo in September of the same year. All the specimens
captured were forwarded to the British Museum in 1903 and 1904.
and a series of five were returned to this Institute. Mr. Austen lias
given us to understand that he wishes to publish a description of this
insect, and we await his publication* with interest.

Lyperosia minutii, Bezzi

Many specimens were caught on horses in a stable at Salikaine
Creek, Gambia River.

Mr. E. E. Austen considers these as a new sub-species, having
darker legs than the typical forms.

Lyperosia ! sp.

Specimens of a species allied to the above were caught on camels
at St. Louis, Senegal, in May, 1903. We have not been able to
identify these ; but they are probably new.

. * ^r' ,Austen has now ijan. iX. 1907) very kindly promised to publish the dugn< -i
this insect 111 an earlv number of this publication.

54

GENUS GLOSSINA (TSETSE FLIES

Native Names. — Only occasionally did the names given below
accurately specify the insects for which they were used For example,
the word designating “ tsetse fly " meant often only " a fly of medium
size which bites,’ and therefore the same term was sometimes
employed without distinction for Gl

and even Slomoxys. In the same way the words given for “floor
maggot” sometimes signified almost any maggot or even “a crawling
thmg which bites.” Glossina pa! pal is is by far the commonest tsetse

in the Congo. The names given for “ tsetse fly " were obtained free
natives by showing specimens of it.

tribeTsh!Ja?US !fng:Ua8e5 are mentioned in the order in which the

Piaces off “TV6" VisilCd'. . . . . - Pven

mentioned. 6 r°Ute ° ** exPed,t,on tllc name of the informant is

nsagTeho7reCo0n *** “d — Relied according to the

are Wri7n L accTd T P“b'ications. The nantes of ^rashes

for the of iztT:z°rd by ,hc Royal

District of the Lower Congo.

Bacongo language.

Tsetse fly
Maggot

Maggot fly
1 SHUMBlRi.

Bateke language
f setse

mavekwa ( vekua )

ul‘ (at Matadi) ; n'tungwa (at Watfcen,

Is a 6eneral name for all maggots)
kuluyanzi

j Vibi (S)
[beyibi (p)

Bobangi language
ey'yi (S)

Note these vvn ^ • " ' ' 09

fUrther d*st'nSuished ^abanus or tsetse : they ate

Maggot . Sma"- °r/"- = large.

UezeufP) ' b"“u (S)

•nabinzn (P)

55

IREBU.

Kundu language

Tsetse - cntune

Maggot - n’kisu

Tabanus - ivovoko

Lulonga.

Lusakani language
vii
kisu

chelifer (also bed bug) — silibongu

Elehu language

Tsetse - yiyi

Maggots - mabinzu

Maggot fly ( Auchmeromyia luteola) - mnboiu
Upoto.

N’gombe language
Tsetse
Maggot
Tabanus
Chrysops
Mosquito

ipokupoku

lutu

ehololwlu
mo tun a
ngungu

Bum ba.

Bangala language

Tsetse - etuna

Maggot - kiso

Language (?) men from

Loeka (Itembiri river)
motoke
n’kusu

Basoko.

Language (0

Tsetse - difu or lifugu (seems to be special name)

YAKUSU.

Keli language

Tsetse - makuku (also signifles Tabanus)

Stanley Falls.
Bagenia language
Tsetse
Maggot

Tabanus

Bakumu language

kowowo

kalombo or bio (?) - kusu

(At Taritubu general name
for all maggots is kiyo)

dikuku

Wanie Numbu.

Language (?)

Tsetse - bungu ( ngt — any fly)

Mosquito - tti'bu

People just below Wanie Numbu called Glossina palpalis 4 kabnb* ■

Swahili language (local dialect)

1 setse - chafua

To the East of the Congo River at Lowa.

Bujero language Bakumu (?) or Babe, no people

- otjapara, said - sumo (S)

to be specific basumo (P)

Tsetse

Kumba.
Language (?)
Tsetse

karboiboi

Kasongo.

Swahili language (current trade language
mjhe eastern part of the Congo Free

Tietse - kibou kidogo, i.e., a small

Tabanus biti"R %

ktb°u makubwa , i.e.. a

Stomoxys , .‘arge bitinK fly

Maggot" ' ”gt’ ■ a Ay

fuiiza or funga

Bang° BanS° language

Ma^°‘ - kivinyc

Katanga dIStrict

7--—,

- kKembe (kasembe) ka%cmM'

Bakusu langt
mazanga

lllfcuSU

57

Cabinda.

Basonge language

Tsetse - kiboua ( l undo at M iamb we) ; same name is
used for Haematopota
Maggot - kifinya

LUSAMBO.

Baluba and Bakuba people present Basongomeno language

Maggot - kikusu (S) Tsetse - bohembc

bukusu (P)

Angola district.

Language (?) (Bastian, native interpreter at Lusambo)

Tsetse - dibubulu (S)

mabubiU-u (P)

Maggot - dindwe (S)

inandwe (P)

Lad o enclave.

Miza language (Commandant Charles Lemaire)

Tsetse - ewe

In the North of the Free State.

Mangbetu Azande (natives of these tribes)

Maggot - ngousu - agbiti

At Koussour on the Schari River near Lake Tchad.

Maggot - tadi

Maggot fly - acou

1 his information was given by an intelligent lad in the service of
the expedition led by Monsieur Chevalier.

Glossina palpalis, Rob.— Desv.

(PI. iii, figs. ;-q)

Larva. Orange-yellow ; hood or anal segment black. Form
somewhat cylindrical. Segment^ very pronounced ; transversely
wrinkled, and finely tesselated, the individual tesserae being flat and

shiny. Mouth hooks minute, contiguous and black. Anal segment or
“ hood ” with the tumid lips deeply divided and widely separated
When fully matured the larva almost completely fills the abdominal
cavity of the parent, and lies with its posterior extremity ("black
hood ”) towards the vaginal opening. A female which died while in
the act of parturition has been preserved with the anterior extremity
of the larva still attached to the vaginal Orifice. Plus interesting
phase in the bionomics of this tsetse fly is illustrated on PL in. fig ;
Length 4-50 mm. ; width 175 mm.

These measurements were taken from larvae laid by captive flit'
among whom abortion is frequent These larvae may, the rcfon 1
be quite fully matured.

Pupcir This agrees so well with the description given by Austen*
that it is necessary only to call attention to the folk whiff details
Colour, m a bright light, dull steel 1,1.,,. but traceable only in

widH specimen^' r,ie tumid Kps ;>>v divided to the - and

widely separated.

Length 5 to 575 mm. ; width 3 nun.

prefenTlf the w *“***** ‘5 «■“■«» «° be certain*

wh"rLeciln, l me"ti0ned tUe foIlowin8 bst. Places at

mentioned. If th^Tped*"1611 ^ ^ expedition arc' 3S ““"I 5,mPl.v
and the date are gtven d°naled by friends """""

obtained, appear onV<the hst^c' f*°'/ ’"'I*.*1? "" sI>cc,mcns have been
present at them because nf ,1 ' palpalts ,s Relieved to have been
been seen and because f *C prox’ni'ty places in which it has
nature of this evidence’ • eV‘denCe assert'ng its presence. The
indicated whether oer *** every instance mentioned. It i>
were shown specimens Z “^“d with the name of the fly
11 was sported as present /eC°gntsed u as be»ng present, or whether
evidence cannot be accented ose -knowing its name. As a rule socfc
a small tsetse (not G. fuse* m .Cat*nS lnore than the presence of
tary llst llas therefore been r, n£lPL>l’iis). A second and supplemen
,100d sPecimens have not hPrepared of Places from whose neighlx -ur
been ported lo exisnt0t b7; received, at which “ tsetse flies " have

rM^api - —^.11 thC n'arki,,&s Indicating the

ne rsetse Mies. Hrj, XI

M"' X “- "-•! . . ,

„„t,

T”<” ,,,,,

H (see fiages )

59

distribution of Glossina pal pal is are based upon the first list ; those
showing the distribution of undifferentiated species of “ tsetse flies,”
partially depend upon the second list.

Banana (Oct., Dr. Etienne) ; Shiloango, Lukula and Lubengi
Rivers (reported, June, 1905, de Laval); Banza Manteka (reported);
Lufu River (reported, Rev. Morgan); Tumba; Wathen ; Kuzu ;
Kisantu; Sabuka ; Leopoldville (Nov., 1905, to July, 1905) ; Brazza¬
ville; Bamu Island (Jan. -Feb.); Lisha (April, June, July, 1905);
MSwata; Kwamouth ; Kitoto ; Tshumbiri (seen ten miles inland,
said to exist along all small streams of neighbourhood); Bolobo
(recognised as present); Yumbi ; Lukolela ; Irebu ; Bikoro

(reported by natives and Rev. Clark); Bolengi ; Coquilhatville ;
“ AH alon£ Lukmga River to Baringa ” (July, Dr. Angela) ; Ikelemba
River (Nov., Major Malfeyt) ; Eala ; Bamamia ; “ All along overland
route from Boyembe to Lulonga ” (July, reported, Rev. Gilchrist);
Lulonga ; Monsembe ; Nouvelle Anvers ; Bosesera (reported, Jan.,
Chef de Poste) ; Mobeka; Bokanga ; Lisala, Bwela, Bosogodo (May,
reported, Rev. K. Smith) ; “ Along Congo from Lisala to Bumba and
up the Itimbiri River" (reported, Dr. De Valkeneer); Bumba;
Botsali; Yambinga ; Bopamba ; Basoko ; Yalembe ; Isangi j
Yarbumbo (recognised by natives); Yandonge (recognised by
natives); Yakusu (recognised by natives and Rev. Milman) ; Stanley
I alls (recognised by natives) ; Katanga (recognised by natives) ;
Batikalela ; Wanie Bakula ; Kewe ; Wanie Numbu ; Kisui ; Ponthier-
ville; Kirundu ; Lalowa River ; Utikakadjia ; Lulindi River ; Kumba ;
Kasuku ; Maboka ; Lokandu ; Ukungvva ; Kamimbi ; Sendwe ;
Makula ; Mfunkiva ; Kibombo ; Kundu River (Dec., reported, Com¬
mandant Verdi ck) ; Nyangwe ; Salt Springs (Oct., reported, Monsieur
happen); Kasongo (Oct. to May); East Bank of Lake Albert (April,
Commandant Lngh) ; Lado Enclave, 6° 33' N., 290 58' E. (March,
ommandant Lemairc) ; “ Glossina pal palis is everywhere present
m Lele even in the smallest rivers” (Feb., 1906, reported, Dr.

osselet) ; Kalombe (May, recognised by natives); Muadi River;
v 1; Tshofa; M iamb we (recognised as present by natives);
cminda; kiambi ; Katanga (Aug., Major Malfeyt) ; Pania Mutombo ;
atampas ; Lusambo (none at post itself, many along Lubi River, just
opposrte, and up and down Sankuru River) ; Lubefu ; Basongo ; Luano
(kwilu River, July, Major Malfeyt). There were but few G. palpalis

6o

among the grass-covered islands of the Lower Kasai They were,
however, numerous in every patch of forest. In the Gambia (Sept
1902, to April, 1903) Glossina pal pal is was everywhere present along
the river and its tributaries from its mouth to a point some fifty mife
above Fatotenda.

Circumstances of capture: On reviewing this list and cumpanng
it with Maps I and II, it is apparent that 6\ pal pal i\ is probably
present in almost every part of the Congo Free State

Tsetses were usually only found near water. If there be a fringe
o! forest or brush, perhaps 200 yards in breadth, along the water’,
edge, more flies will be seen by gently paddling in a canoe along the
ank or by walking on the land side just .n the edge of the bdt :
brush than by even a prolonged stay in the forest itself. The collec
t.on of water need not be large. 6'. palp,,/,. ,

caught along very small forest streams.

ForTe!fr nrbr Were f0Und t0 varr ereatly in different locate
Sendwe ”?’ I ^ SabuU ne" Coquilhatville. near

BatiunpasandruberoTth:V‘;:i“'f7 * ? *'

outskjrfc ^of ^ o n ! a *** S at Z*

Falls, all places at which ^ *7”' ^

n°t a single fly couid be founj Ca ^al,s P“**

stay in each of these places both hv T W3S made dunng 0Ur
b°ys ; it may be that ^ ourselves and by our fly-catching

days when the Aieas ,n vis"ing — °f *** -

nottced that the number of tset e h“ *>«" repeatedly

vary considerably from dav , f H,es Presen> «> any locality may

on October ^ ^ ^ "**“'* W«-t cause,

dming^the halVoMhe^ °" UpP“ C<**

Again the ^ ^

■tamed because they were ^ of the places

water fronts. «>™Parat,vely free from brush ;lIon/thcir

It seems certain

neighbourhood of - tsetses in the nnmed,

any S12e may be

6x

all thick underbrush and rank grass, either near the water or a few
hundred yards inland, be carefully removed.*

The station of Irebu, a military training post on the middle Congo, is beautifully
bept. We remained there for three days and saw no tsetses, although there were mam
only a few hundred yards further up or down the river. A little over 1,000 yards
outside the station was a trading factory placed in the middle of a patch of rank grass
Glossina falfalis existed there. Resides searching for tsetses on the station ourselves
we sent out five boys trained to catch them, together with 60 soldiers. Between them
all only a dozen specimens were taken. All were caught in a patch of rank grass,
about 350 yards from the river, that had not been cut for two or three months. There
were many scores of acres of cultivated land on the station, mostly consisting of well
kept, but shady, cocoa and coffee plantations. In them not one tsetse was found.

As Leopoldville developes and the surrounding brush is kept down tsetses become
fewer, and we are told they are not nearly so numerous within the Protestant mission
station as they formerly were. During our stay in Leopoldville Glossina palpatis was
much more often caught in the house of the Rev. Mr. Morgan than in the building
occupied by our expedition. The mission house was surrounded by a grove of palm
trees which extended uninterruptedly to the river’s edge, while the house of the
expedition stood in an open piece of ground.

The experimental farm of the Government of the Congo Free
State at Eala possibly may be considered to be an exception to this
rule.

This farm is comparatively free from undergrowth. It is extensive and well kept,
yet Glossina falpalis is quite frequently seen on it. There is a herd of 43 cattle
and a fair amount of other live stock on the farm. It is believed that the flies are
attracted by them.

It often happened, however, that no,, or extremely few, tsetses were
found, even after a careful search, in places where everything, shade,
shelter and water, apparently favoured their presence.

Near Yambinga a search (of several hours), made on a bright day, along the
thickly wooded banks of the main river of a small stream failed to find Glossina
fal falls, although they were present in the near neighbourhood.

At Yakusu several hours in the middle of a fine dry day were spent in a canoe
paddling along the banks of the river and about wooded islands. Tsetse flies were
not seen.

The late Rev. W. Holman Rently told us that at a place called Vela in the Lower
Longo G. falpalis was absent. The natives noticed this and decided that they were
-.offering from a want of the beneficent blood-letting enjoyed by their neighbours wh' .
ived in places where the fly existed. Men were therefore sent out to catch tsetses,
bring them back and let them loose at Vela !

Rehind the soldiers’ lines at T.usambo runs a brush-covered stream which never

u*1 ^fiar La ma'ad’e du sommeil et la tse-tse a Novo Redondo. Reports
o t e 1 teenth International Congress of Medicine at Lisbon, Fasicule 2.
page 294. It is stated in this publication that Glossina falpalis is not
>een in w ell-kept plantations nor at a higher altitude than 400 metres above the sea
I his latter assertion is negatived by an observation of Commandant Charles Lemaire.
who sent us specimens of the fly from the banks of the Ialo River (6° 33' North :
29° 58 Last) caught at an altitude of 530 metres, and also by Lieutenant Rrohez.
who caught Glossina at an altitude of 1,600 metres, in places where the moraine
temperature went down to o° C. (loc. cit., vide infra).

62

goes dry, and in the wet season measures from ten to twenty yard' in breadth On?
could not imagine a place better-suited for tsetses ; yet three competent obsentn
(Dr. Broden, Dr. Polidori and J. (.. T.i duri ieral months mw now

there, although they were looked for many times. The observation might b?
explained by the fact that the stream is used as a latrine, and in hot weather reel*
with effluvia. If this explanation be accepted it seenx strange that G falftlis bn
twice been seen in latrine sheds.

Tsetse flies will sometimes follow persons or animals for a
considerable distance.

In the Lower Congo the belief that (7. falfa
have not confirmed the observation. We are told that in the neighbour!., d U
Leopoldville G. falfalis was called by m .

(Dr. Broden).

tsetses we!°bfala’ “ ,Catt!f St3tion in the Low“ ( aaaerted tttf
tsetses?were formerly absent. Pigs were introduce,., and *„», ,, • •

brackishhwa?erniof ^ 'm ^ * 'Ime a hm,se P,ac#d a mile ft t, the

KHe!,! f * mangrove -swamp where were a , d - ;

through Seew^rSr^These^ h.ouse by per* vt ' ' ! J" '

. ■*- — . — ^

used as a laboratory ^Two tan|f '’T't' "" had ' '' r’

ff. taltalis followed them btOU«ht ; two or three

support the idea^thfd tsetTe'fl^T^Ln150^31’0" "as «athered »'» th' Cong, tn
many (7. jalfaUs were seen in W iSP "" 1

of any sort. ** “ ,OCa,'tles " •> gJe

seen atP places'* where* thev ^ tsetse n'es were occasionally not

When the exp^on Z T” P^sent

Iron' to Pania Mutombo emT"" > ' " *"* ,h,‘ 11 overland

several of that ,setse dies were not found at

and Lukola during the ove I ^ ne3r K:,lo,nbc' Oibwe, Miambwe
was usually made ^er,and J°«ney of the expedition Search

number of porters had “d bridges after a considerable

fl'es were not found in ** Sli" p— The

Ae'r presence and although human'T’ although ,he natives asserted
earch of one or trypanosomiasis existed localh

fjre ** bud at first sZ/dto7 u ^ at a ™ple of streams

-;"ke

i, , . .

63

percentage of resident natives having enlarged cervical glands without
apparent cause is indicated — Kalombe (i; per cent.), Dibwe (t8 per
cent.), Miambwe (; per cent.), Lokula (6 per cent.).

A proportion of the infected persons probably contracted their disease elsewhere.
These villages have a considerable immigrant population ; they supply many carrier'-
and a large proportion of their people are continually absent, collecting rubber, in
the forest near rivers where tsetse flies are said to exist.

Had the flies been present in as large numbers as, for instance, at
Kamimbi, it is quite inconceivable that the passage of even 300 men
should have enticed them all away from their usual haunts.

The occasional presence of G. palpalis in places at a considerable
distance from any collection of water may perhaps be explained by
their habit of following animals.

In the Gambia, and again in the Congo, a single G. -palpalis (?) ha- been
seen on a dry plateau a couple of hundred feet above and about a mile distant from
the nearest stream.

As would be surmised from what has already been said, Gloss inn
palpalis is able to fly quickly and for comparatively long distances.

They are said to occasionally fly into the railway carriages as they pass near
Palabala in the Lower Congo. They are frequently seen on steamers or canoes in
mid-river, and distant 300 to 500 yards from either bank. On entering a moving
vehicle the flies seek sheltered spots, and it is under the sunshade of the canoe or in
corners of a cabin sheltered from the wind that they take refuge.

On one occasion in the Gambia a G. palpalis (?) persistently followed one of us
(J. E. D.), who was riding rapidly on a bicycle, for several hundred yards.

At Tshofa, G. palpalis was frequently seen on the verandah of our house,
although it was placed on high ground and at least two hundred feet above the river,
which was about half a mile away.

Although G. palpalis has such extensive powers of flight, it seems
to be very local in its habits. As has often been observed, not a
single fly may be seen at 100 yards from a river although its banks
swarm with them.

At M’Swata, however, G. palpalis was regularly seen along a little-frequented
path on high ground and in open park country at a distance of 500 yards
from the water and of 250 yards from the edge of the fringe of brush bordering the
river. In partial explanation it should be stated that there was a good deal of game
in the neighbourhood.

It has been often noticed that G. palpalis is most numerous at,
and seems to lie in wait by, fords or frequented paths. Its habits
have seemed to be more or less regular.

On one occasion, while resting by the side of a path, a G. palpalis was noticed to
return on three occasions to the same spot on the sheltered under surface of a gra"
stalk after having been driven off by a native porter whom it was trying to bite.

Tsetse flies were practically never seen during rain and wind.
They were most conspicuous and seemed most vivacious on bright.

64

warm days. In the cool of the early morning or on chilly days thev
were sluggish and were certainly much more easily captured; undo
these conditions tsetse flies have often been caught with the hand
ordinarily almost an impossibility'.

Glossina palpalis is often seen in native village l0«s

ot Luropeans living within a few yards ol the rivers edge

^\Xnnl\nl'‘^ganaSSTedthi,' 'U' ' * •' • how asm

- » ** ,he;\v0e"e\TiT^w * ITT" ^ ^ * *++

Of the resident European. " K * d thumb, were wen ta the boon

on lbld ”TS °n /h‘ Bionomics °f ,sf,s‘ /?<« These note, are bwd

.0 traZrr5 made on the cap,,v" •>

sersns: y ,heir bi,es- and - - - ee

thefer^^n? PrehPareS l° feed* " to kw, prim*

it were, "seithL T ^ wholc borl> “W to host; .has*
they form an ansi/T 1° US“,eS*-' 1 l"’ P'lll» an

There " occaZa,. ** '35 <l.r hared

proboscis is inserted X O ? *W° °f Natation before Hr I

"* in »

behae tri“ °r du<* ^^thes.^Theywer! T' ""

m Je "'<rab]e to I-netrate 52 ^ ‘ .

"If 3 Paten‘ed cloth. t,ghtl'V WOVen ‘exture of a pair of riding breech,

1 he Penetration of the «d-* i

conSi^:st;a°ccasi™^ somci,iws ause

rhe: ■ T-' hJW a: i

The probos PPeared' D,,e’ The Wra,s- evidence that l

r„°f feed,n^ -Z"thr,eZ inserted at commence

partialK- \ d operations are com Proboscis is withdraw

The flW raWn to be re-TnserteHenCed 3 P'-e. or ,, .

- — ^ving observations will perh apparcml>’- a new direction

*r;- €3Cpiain l,u' u ch this

"ith * pocket-, ens while

65

may be done. It seems that G. palpalis flexes its proboscis in two
distinct ways. The first, the more usual, is a sharp bending of the
labella. Flies who had just fed were often seen, while preening
themselves, to sharply flex their labella so that they stood out,
almost at right angles, from the palps. Fhe second movement was
only seen in flies from whom the palps had been removed preparatory
to dissection. In these the whole proboscis bowed until it described
a well-marked curve directed ventrally. The second movement is
probably due to the contraction of the powerful “ tendons " lying in
the labium *

G. palpalis does not fill with blood so quickly as is usually
imagined. The length of time between the insertion of the proboscis
at the commencement of feeding and its withdrawal was noted in
flies carefully fed on an experimental animal. The greatest care was
taken to avoid disturbing them. The longest fed for eleven minutes,
the shortest for minutes, while the average was slightly over three
minutes.

The extraordinary way in which tsetses distend themselves with
blood is well known.

A female G. fusca was watched steadily filling itself with blood for six minutes
It was stopped feeding then lest it should burst itself. Its abdomen was so distended
that there was a space of almost a millimetre between each of the dark coloured
chi tin plates on the dorsal surface of its abdomen.

In about a minute after feeding commences a drop of yellowish or
brownish, opaque, but liquid faeces is extruded and a minute later
while still feeding — clear serum rolls, drop by drop, from the anus.

Sometimes tsetses who were plainly in need of food absolutely
refused to attempt to feed. Others frantically probed animal after
animal and failed to get blood. It has occasionally been thought that
in captivity male flies did not seem to be so eager for blood as were
the females.

In the cool of the early morning or evening G. palpalis is not
nearly so voracious as at mid-day.

One is, however, occasionally surprised, as at Lubefu on the Kasai, in finding
them active and biting freely early on a cold damp morning, or as at Kasongo and
I.ukolela in seeing them feed until dusk and fly actively about after dark
G. palpalis has twice been caught about the lamps at night.

I hese flies sometimes seem to prefer natives to Europeans, and dark to light
coloured clothes. This first point is frequently commented on by European-,
travelling in canoes. They see that the native paddlers are worried by the flies while
they are themselves often comparatively unmolested.

* Memoir XVIII of the Liverpool School of Tropical Medicine, page 53.

li

66

One day while coming down the Kasai the Captain, finding it c Id, put > V •
cloth jacket. Many more tsetses were not- than on the vfeik

duck suits of the European passengers and of the negro steersman

The flies seem to find their hosts rather by sight than by Mnnll, sin. e in i gne
locality a moving person is usually more troubled by them than i » one wh- .u% *•:'.!

The persistency with which Glossitta /'<>//'■ attacks

despite slapping and brushing is very characteristic As has alrrah
been stated, its flight is ordinarily rapid and darting It is usually
accompanied by a peculiar and charSU ' i m fly can

nevertheless, if it wishes, fly as noiselessly as an Anophelt A fuD\
gorged fly is exceedingly heavy on the wing, and Utoall] 1 t only a
yard or two before taking refuge in some quiet corner, where it
quickly lightens itself by the rapid extrusion per tt n urn of large
quantities of colourless serum. Immediately . goes

through an elaborate toilet and carefully deans .is proboscis. W
times just after feeding a drop of blood has been seen adher.ng to Ihr
outside of the proboscis at about its middle.

If a tsetse (G. fusca or G. pal pahs) be held by its « ingl. or other-

Z d t C3n bC "lade *° drop Tclear Hod

from the ,p of tts proboscis. Koch has made a sin, 1

association w 7 tha‘ ■» * . unly found .

plants were pointed™'^ One Z ConR" ,W°

other a variety of mimosa Rotl * *°n °f ^ ^

river bank, and therefore were ' fZ '°'V lanH alonS
Needless to say, the flies h . * Ways Seen whcrr tsetses existed
both were absent. ^ °en ' 'l'"’"' l\ seen in n!

was found to be ab^mhomZ'1'0" 3S WeD “ U‘e fact ,hal
tatsed the question whether th, ''PP'>rcntly well suited to it hare
given locality may not varv T nun,bers of ‘wise flies present m a
influences. No satisfactory^ *** ®rea,ly *hrou8h seasonal or other
observations are, however 7,e 7 ^ hCe" obtained The following
ts AtflMatadi and elsewhere h ^ th'S c°nnection

Br-h” tnSsSs^2>“/“l in '*>• t-ambi*. « J the

b“e more ««c.ly i„ ?h",T"y:ana «*•« that i„ Kit, Ini' ]t‘.

more numeroui
f-t/fa/ii pnwHil

.in., Forw-hnngmir
• ^7- 'ov. aj, iqoj.

nark

6 7

In April, just at the end of the dry season, a camp was made on the banks of the
Upper Gambia river. Glossina palfalis was very numerous, but, strange to say,
caused but little trouble ; there was a great deal of game in the neighbourhood.

It has been reported that certain localities were infested with tsetses which at an
earlier period were free from them. These statements have not been substantiated,
and they seem sometimes to have resulted from insufficient observation. The possi¬
bility of their accuracy must not, however, be forgotten, especially since it is
frequently asserted that the area over which Glossina exists is widening.

Observations on captive Glossina pa l pal is.- During our attempts,
in the Gambia," to transmit trypanosomes by the bites of tsetse flies,
we found that they would not feed well in small cages and died in
the course of two to four weeks. It was thought that this might have
been one of the reasons why our Gambian experiments had been
unsuccessful, and when we recommenced our experiments in the
Congo we determined to reproduce, as nearly as possible, the natural
surroundings of the fly by keeping them out of doors in larger cages
containing water and growing grass. Cubical cages of wire gauze
measuring 18 inches along each side were employed. In each cage
was placed from 20 to 70 flies. These larger cages were kept in a
large mosquito house, measuring 13 by 8 by 9 feet, made of wire
gauze. It was placed in a thick clump of bamboos, distant about
100 yards from the river. The flies were sheltered by the bamboos
from the mid-day sun and, on two sides, from strong winds. Food
was supplied to them by experimental animals, guinea-pigs and rat-,
which were immobilised and placed in the cages. Glossina palpalis
certainly lived better under these conditions than they had done in
smaller cages in the Gambia. They frequently lived for over 30 days ;
43 and 59 days were the longest lives noticed. In addition they
seemed not to require food so often as had been necessary in the
Gambia, and they were frequently left for 48 hours without an
opportunity of feeding. This may be explained in part by their being
out of doors and more exposed to cold and damp. Under such
circumstances tsetses are certainly more sluggish in their movements.

The disadvantages of this method of keeping flies were however
great. It was almost impossible in such large cages to count exactly
the number of flies which had fed or to estimate the number present
They seemed to seek concealment and hid themselves in the smallest
crevices. It was difficult to feed flies kept in this way on monkeys
and the cages were too large to be easily transported.

* Memoir XI, Liverpool School of Tropical Medicine.

7 o

If they were left without an opportunity to feed for much more
than 24 hours they died very quickly, about Qi per cent in 37 hours
Neither sex seemed particularly resistent.

To feed the flies the cage was simply pressed against an infected
person or animal, and so long as the victim kept fairly still the die-

, . . ” A*tt,HOD OK Fkr, Tskt.sk Kues

ted without difficult v Tl , -

and practically the same ^ .S n° Preference for human

(man* monkey, guinea-pip- JCenUj?c ot n,es fed on whatever a

0 . .

;111 aVera^e 80 per cent, of ouTffie^f thousand fl,e5

" cases, when great care ' "" When ,he °PP°rtunity ol

Perfectly stiU and the L! L Z ' ° k«P an.mal and

Ifrl P"Cen‘ages "ere obtam^ P7S‘Sted in for some hours,
roh:f had partially Z 7ted frrent'-V - per cent,
at once resume their mea on ^ could often be pers,

°n a Se“nd animal, „ was prac

7i

impossible to get flies which had been allowed to gorge themselves to
again feed within from three to five hours’ time, although they would
do so readily eight hours after a first feed.

It has already been stated that G. palpalis may buzz loudly
while flying. When at rest, often just after feeding, it sometimes
produces a shrill, high-pitched note. A wingless fly was watched with
a pocket-lens while sounding in this way. The tips of its halteres
were seen to be depressed downwards and forwards, but the
mechanism by which the note was produced was not ascertained.

Glossina palpalis, var. welhnani, Austen

Localities: — On the Congo River, between Nyangwe and
Kasongo ; on the Gambia River, near its mouth, at Oyster Creek.

Circumstances of capture : — So far as was observed the habits of
these flies were quite similar to those of Glossina palpalis.

Glossina morsitans, Westwood

Localities: Specimens have been received, through Monsieur
Brohez, only from the Katanga district. The exact locality and the
circumstances under which they were caught are not certain, but they
were probably taken along the Luvua River near Pueto. Dr. Hosselet
definitely reports that this fly is everywhere present in the zone of
Uele, but we have seen no specimens.

Glossina fusca, Walker

Larva ( dissected from body of parent but fully matured). — This
consists of twelve strongly defined segments ; surface finely tesselated,
the individual tesserae being circular, flat, and shiny. Form, rather
short ovate. Mouth-hooks relatively large, black, rather widely
separated, but not porrected. Anal segment or “ black hood ” very
coarsely rugose, the space between the converged points of the tumid
lips one-third the diameter of one lip. Colour white, with a trace of
lemon-yellow ; hood or anal segment black.

Length 5-50 to 6 mm. ; width 3 to 4111m.

Pupa much more tumid and ovate than that of G. palpalis,
especially so posteriorly. The tumid lips are also more depressed,
and being continuous at their bases and only slightly divided
posteriorly form a deep siphon-like tube or crateriform process, so

72

deep that it is impossible to see the stigmata which he in the interior.
Colour, dull steel-black, without an y trace of the dull steel blue seen
in the puparium of G. pal pahs.

Length 7 50 to 8 mm. ; width 4 to 4*50 mm.

Localities: — Leopoldville (Dec.; reported, all the year round)
Tshumbm; Yumbie (Dr. Rodham); LukoleJ cad);

Bolengi (Dr. Dye); Coquilhatville (Dr. Mordighlia) ; N’ouvcllc
Anvers; Bwela (reported, Rev. Kenred Smith {tongipenuis ?));
Aruwimi River; Lulu River (reported, ( '< unm.111. 1.mt Lund ( longi ■

pennis ?)); near Lalowa ; near Utikakadja; Sendwe; Mftmlrin
Kasongo (Oct.- May).

Circumstances of capture:— This fly occurs in very much smaller
numbers than G. pal pahs and it is the rarest thing for more than one
to be seen at a time. Most of our specimens were caught in the bush
or from canoes paddled slowly along the edge of a wooded stream
one were seen to fly on board the steamer used by the expedition
on their journey from Leopoldville to Stanly l ulls, altht ■

t M ^ f°Und " °nC °f the “bins. Gloss,,,, fuse, has

inT tabk.een erVed th£ C°ng° ‘° be morc or lcss noctunuI

A buffalo was shot Part .u

arrived about an hour after nightfall “ 1^11^1 "h* b>' Ca,"P’ wher* "

trying to feed— there was no hlnnH ■’ ■, 3 ' a ,aml' ,l aught

two later a pair of G. fusca were * 1 i! St0mach — on buffalo'* neck

On two occasions specimens have w8 CO,tu’ about ‘he lighted lamp in a tent,

of the expedition. ha'e been cau«ht a*‘er nightfall in the illuminated tents

fly have beea

Glossina pallidipes, Austen

were seen, although ,05^^ ^ °btamed at KasonS° No others
ments, were carefully examined6 CaUght for transmission expen-

. Localities;- ThtlTh Wiedemann

district. Two lots of soeL ^ y been sent to us from the Katanga
^ough Major Mai feyt, WW -“ived One reached as

PPer Luapula. 'Cr V'as bent b>’ a local official from the

73

Field notes: — It is asserted locally that these flies are more
persistent and feed more viciously than Glossina palpalis.

Glossina maculata, n. sp. (Newstead)

(PI. in, figs. 5, 6)

General appearance- Very dark brown ; posterior surface of head
cinerous, spotted with black ; thorax dark brown and faintly cinerous
in places, with elongated transverse black spots ; pleurae, coaxe and
femora cinerous with conspicuous black spots ; first and second
segments of hind tarsi dark. (The remaining segments wanting.)

FEMALE. Head with the frontal stripe rich ochreous brown ;
frontal margins and occellar spot bright yellowish-white to pale
ochreous ; ocelli black ; posterior surface of head cinerous with large
well-defined irregular black spots. Antennae dull red-brown in front,
sides with a cinerous surface ; arista dull reddish-brown. Palpi blackish
above, paler beneath. Bulb of proboscis dark castaneous, shining.
L'horax dark brown and cinerous with numerous irregular elongated
black spots placed transversely ; anterior angles pale brown. The
scutellum spotted like the thorax, margin pale brown ; pleurae
cinerous and pale brown, with numerous, irregular, and more or less
confluent, black spots. Abdomen very dark brown ; the narrow basal
segment cinerous, with numerous black spots ; the broad second
segment with the large median area, the anterior angle, and broad
hind margin, cinerous, with black spots; the remaining segments,
with the exception of the last, with a narrow well-defined median
stripe, and hind margins narrowly cinerous ; lateral margins of
segments three to five with a cinerous triangular patch ; the hind
margins with an occasional, more or less obscure, black spot, with the
exception of the sixth, which has a regular series of ten or eleven on
the grey cinerous band ; last segment darker than the rest with a
lateral greyish spot in the centre of which is a small black one. Legs :
All the coxae cinerous and pale reddish-brown, with numerous more
or less confluent black spots ; anterior and mid femora cinerous
basally and apically bright red-brown, cinerous area with numerous
confluent black spots ; hind femora with the basal third and apex pale
red-brown, the rest faintly cinerous with a few large, faint, dull brown
blotches; anterior and hind tibiae pale brown, mid tibiae slightly
darker with, in some lights, faint indications of dark spots ; anterior

74

tarsi and first segment of the mid pale brown ; first and second
segment of hind tarsi dark brown. (The remaining segments of the
nnd and hind tarsi are unfortunately wanting.) Wings uniformly
brown.

Length, w exclusive of proboscis, 9 mm. ; length of wing 9 nun

A single specimen of this fly was sent to us by the Rev and
Mrs. Billington. It was collected by them at I siiuinbin in 1905.

I his interesting species may be readily recognised from .ill oilier
members of the genus, by the curious spotted or mottled apjicaran t
of the thorax, pleurae, and femora I*o the unaided eye it lot Its
like a dark specimen of Clossina pal pal h% R I) ; but a pocket km
immediately reveals the peculiar markings.

e u iv 1 ri k

TSETSE FLIES

Trustw

• t t "°rthy persons frequently stated that tsetse flies of sm;
2 2 °r existed in various localities Th,

^;,revwurble ,o further ***& ^

1 st 0 0 r theief°re c°,npiled ^Mowing supplements

havebelnl1: **“ flles ~ The names .. ’ ,rrmg Z

distribution Of fteC/JVj* prePa™‘i°n of the map showing «

Makunga, M^nfema (I'porL^An Dr. BignamO

Kamba,ontheLomami R 1 ; ' ' '?°S' Chef <k' P°«') ■ Bo

road from Kasongo to ^ na,ives) : " A11 alonS tl

confirmatory reports h; V ' * (reported 1905. Bishop Rodens
along this route Piani T 3een received from the following place
yamba; Kibonga 2*™^’ Kabambare. Kalembelembe, Vamba
Klvu (reported, Command^f l ^eportcd> Chcf de Poste); Lak
Bishop Rodens) ; “ pj. , lunt‘) : f PPer Luama River (reports
district are so few as to h t.Setses do not exist in the Katang

Chef de^.Secto ^11* 1 (reP°rted Feb. 4th. 19°!
- - - Places in Katanga at whirl

t This statement i • ,s has not been allowed for in th

vSrl* report, received bv u,

“wera ** c- * -

75

tsetses are definitely said to exist are : Ankoro ; along the Lubili
River ; on Lake Tanganyika at Mpala and Vua ; * Lusaka ; Kiambi ;
Mpweto ; Lukonzolwa ; Kilwa ; Lukafu ; Kasenga ; Kambove ;
Tenke ; Shiniama ; Kalonga and along the Lualaba, Lufira and
Luapula Rivers ; Upper Kasai and Luebo Rivers (reported, Mr.
Vemer) ; Angola (reported, Bastian, Interpreter at Lu sambo) ; Popo
kabaka (reported, Dr. Hollebeke).

Map II lias been prepared to show the distribution of Glossinae in
the Congo Free State. For the sake of uniformity we have used the
same signs, as Mr. Austen to indicate the various species of Glossinae.
As will be noted, our observations confirm his map in many instances
and amplify it in others. In it has been incorporated the information
contained in the article on tsetse flies by Austen, published in Volume
VI of the Reports of the Sleeping Sickness Commission of the Royal
Society and in a communication, on the same subject, made by
Laveran to the Academic des Sciences.!

Remarks. An important deduction to be made from the above
notes on the Glossinae is that Glossina pal pal is, at least, cannot be
said to be absent from a given spot until several careful searches have
been made at considerable intervals of time.

A careful study of the bionomics of tsetse flies is still urgent ly
needed. Comparatively little is known at present concerning their
habits ; especially lacking is accurate information on the variation of
their numbers in various localities (“ fly belts ”) and on the causes
governing their appearance or disappearance. On only one occasion
did any hint of a natural enemy of these flies reach us.

The Rev. Mr. Grenfell had slightly wounded a bird (a darter? — native name,
idling a). It was taken living into the canoe in which he was travelling, and it was
seen to eagerly catch the tsetse flies flying about the native paddlers.

GENUS STOMOXYS

Stomoxys calcitrant , Linn.

(PI. iii.fig. 4; P!v, figs. 1-8)

Localities: Lulango ; Lusambo ; in the Gambia at Bathurst and
McCarthy Island.

Dr. Xoble has recently identified the tsetse flies “plentiful at Vua ” as G. palpali -

f “ Comptes rendus des stances de 1’ Academic des Sciences, t. cxli. p. 920.

4 ddeembre, 1905.”

Circumstances of capture: This fly was caught in the open and
in the houses of Europeans. It feeds on all mammals, but seems to
be especially fond of feeding at the tips of dogs’ ears Both on the
Gambia and on the Congo dogs were often seen with their ears rav
and bleeding from the attacks of this pest.

*It is, I believe, over 60 years since Bouche discovered the larvae of
Stomoxys calcitrans in warm stable manure. Since that time no
additional information on the habits of this insect has apparently been
givent ; and the characteristics of the earlier stages and also the meta¬
morphoses have remained practically unknown. It was resolved
therefore to : -

i. i race out

insect from the egg ; and

*be natural habitat for the eggs, larvae and pupae,
he first was accomplished with comparatively little trouble; but!
n ing of the natural habitat lor the earlier stages of this insect prov
no light task, and involved a considerable amount of time and diligl
eaicimg. In the end, however, l succeeded in solvira

intere-TV ^ ll0ped tliat t*iese *»Otes will HOt Otll> piOVe

engaeed inU| l a S<J °a practical value to those students whoa

disease by hZllucZg7eT “ CO,H K' ' *"»*** :

for thriarvae anT'pupae^f ' Tv “ UnremittinK search WaS
all the domesticated l ? , [ HS. insecl in the facccs of near
but without success- m ^ I?* ^ *** l;nu. y.ml- and the fid
searched, but nnnP ’ lla“Ure heaps both old and recent were al:

- - were discoverable. A number of females wei

. ~ " - — - - , _

Pl' TS*"' UePrin,ed ,r°m ,he J°“rn' K“»- »W.. vot. i. No. <. pp. 15
insect fauna of Hmn6 excellent paper on “ a . .

P- 578) I had oveIwtnJexcremen‘’’ (Proc \v» .c.ontribul,on l<> the study c
this fly was in l8i?oke^» says Our first \V“sh,.ngton Acad, of Science, Vc
specimens of sCZf’ when studying the horn fl"16?0* w,,h ,he breeding hat
Washington, a£‘ reared,?/ Cai!,e Au*‘* >»*

observed by Mr m",.? '9th of the *roP1 horse manure collect!

and May ,4> iSqo^oth*11’ attracted to freshlv T*h *ir?e nu™bers of adults
nianure being collect^ sPec>mens were re-,£? r°Ppet horse mam,re. Jaiiua
e«ther the larval or ^ °? November 27^'" £0rse ”'a»ure. the last I
manure. °r Pupal condition (probaSfr .1° t,hat ,ht* hibemat

. This species hnc ' * Iatter) in or iust undei

feSS ^Virginia ;° an

77

then captured and placed in a large cage well supplied with light and
air, and fresh faeces of the horse, sheep, and rabbit. A small
percentage of the insects laid their eggs under these conditions ; but
the eggs were invariably removed to receptacles which were more
convenient for the study of the metamorphosis. It was found that
two important conditions were necessary for the development of the
larvae, viz., an almost complete absence of light and an abundance •>/
moisture. Such conditions as these could only obtain in a state of
nature in large faeces in shady or damp situations, or in heaps of
manure.

Towards the end of September the earlier stages of this insect
were found under natural conditions, and the facts relating to them
are given under the heading of additional field notes.

HABITAT.— Farm-yards and stables are evidently the favourite
haunts of this fly ; it occurs also in the fields, parks, and open woods,
especially where cattle are grazing, but is much less numerous in such
places. It is evidently also by no means uncommon in some of our
large towns, and numbers were seen at rest on the shop-fronts in the
main streets of both Liverpool and Chester. It is fond of resting on
surfaces fully exposed to the sun, such as doors, gates, and rails, and
to a less extent also on stone and brick walls. Painted surfaces are
also attractive to it, and the greatest number seen congregated
together were disporting themselves on the sunny side of a red
painted iron tank at the old Chateau de Goumont, Waterloo, Belgium.
They are very active ; but their flight is quite inaudible at a short
distance, the noise produced being very feeble. When disturbed they
frequently return to the same spot, but more especially so in favourite
resting-places. At night they retire to some sheltered spot, and
numbers may be found at rest on the beams and rafters in open sheds
in farm-yards, where they remain, almost inert, till the morning sun
tempts them out again. They will also occasionally enter stables in
the day time, and they were seen to enter such places through a
narrow opening or a crack in the door.

They frequently clean their wings when in captivity, and this is
accomplished with great precision, the hind legs being used for this
purpose. The under surface of the wings are first combed, then the
upper, the legs are then rubbed together, and the process is again
repeated in exactly the same order.

Position when at Rest. The front part of the body is
slightly raised, but not invariably so ; and the wings, which invarab:
touch at their bases, are widely divergent and carried in a horuonti!
position, lying practically in the same plane as the abdomen
Ratio of Sexes.— During the heat of the day the male* p?
ponderated ;* but towards evening the sexes occurred in about equal
numbers; the captured females W(

emerged ones, and a large proportion of the eggs which thev Wir
captivity proved infertile.

Food of the Adult.— During a period of 14 days a axtfc
,a ^ was eP* 011 both cattIe and horses m various farm yards

u « 1 c < onuiK hi, but no flies were sen < 1 1

”"u.i s or to suck blood from them ; game time sneal

pno- P ° • ?0t*' ln,llt's il1"1 females pttued m

blood More°ver. a freshly emerged mat

fT ,lK' write^s own hand Then- IS 1,

" " <x" .a8to ,,u' Wood sucking habits of this •!% which
need not be repeated in this communication

apparent^feeTupoTTl ^ -t0 °" fre5h CO" dun& **

proboscis rapidly over the « on "• P*«“«

tional or rarely seen in . ^ ; snch *»bits were apparently •
fed on the fresh faeces of H ° !’ natUrc ; b,,t in captivity they readily
the latter. A female ^ lorst' and sheep, more especially so or

thorax of a dead ron aS ^S° seen to drive its proboscis into the
body. Three specimeTTd apparentl>r *uck up the juices of its
also sucked up the f ° r Upon sornc sugar and water, and some

Movements of -m t> *°m a decayed ,n'1 fetid ;

frequently depressed tu ROBoscis- Die base of the proboscis »
of the labium remain* 11 * PaJp* become fully exposed, but
» etevated. same position, or is very

eX,ble’ and can be instmH^ J*°int in front of ,he palp* is
?n be COniPletely straiVrhm mflated 50 that the entire proboscis
jessed vertically. Tfle , ,ed* and either extended horizontally or

"• Straightened. takin ^ 3,50 be :

- “ the labmm pr0P"

i

79

2. Curved upwards and outwards, with a quick alternate move

ment to either the right or left, or repeatedly and rapidly
curved to one side only. As the labella curve upwards the
anterior portion of the labium also gives a distinct lateral
twist, so that the dorsal groove is presented laterally.

3. The teeth of the labella can be curved outwards and ventral

wards, giving them a bilobed appearance ; the teeth also
apparently move rapidly backwards and forwards, and a
clear fluid was seen to pass down the tube when the insect
was slightly pinched between the fingers.

In sucking blood from the writer’s hand the insect sat high upon
its legs, but the anterior pair were much elbowed, and all the joints
of the tarsi generally rested upon the skin of the host. The whole
of the proboscis was straightened and held vertically, and the anterior
third was driven into the flesh.® During the process, which lasted
altogether for a period of 15 minutes, the proboscis was constantly, but
somewhat slowly, moved up and down, and also with an occasional
semi-rotary movement, reminding one somewhat of the action of a
quarryman’s hand drill. This action was continued until the fly had
pumped its body full of blood. The initial pain was trifling compared
with that of a mosquito ; but there were two subsequent pricks which
were quite as irritating as the first. A small drop of blood was left
over the puncture, and when this was washed away a small roseala
was revealed ; but there was no subsequent irritation or soreness of
any kind. A clear fluid was passed from the anus four times during
the process, and on several occasions subsequently, and judging from
the size of the abdomen the food was rapidly assimilated. This fly
died twelve hours after feeding.

Duration OF Life. — In captivity they lived for several days ;
but they were supplied with abundance of fresh air and some moist
faeces. The females died either immediately or shortly after laying
their eggs.

EGG-LAYING. — When the female is about to lay its eggs the
ovipositor becomes fully extended, and nearly equals the length of the
abdomen proper. The eggs are passed rapidly down the ovipositor
at intervals of a few seconds, t and were usually laid in an irregular

*In thick-skinned animals the proboscis would in all probability be driven mi
still further.

t 5*3° seconds.

* 8o

heap. In some instances the female was seen to separate the eggs by
carefully passing her proboscis between them, and then drag them
away or scatter them with her legs. In cases where the eggs were laid
during extremis the female generally died on tin* spot, and made no
attempt to scatter them. Counts were made of seven batches, the
maximum being 71, the minimum 48; the actual counts were 48, 48,
54* 5 7. 59. 62, 7 1. The incubation period, at an average temperature
of 720 F. in the day and 65° F. at night, was two to three days
Time of Appearance. They are ibundant during

August and September ; but gradually diminish in numbers during
the early part of October ;* and few examples are seen after a spell of
cold wet weather.

The Egg. Coriaceous; white at first, but changing to creamy
white. Those which were laid on faeces, fully exposed to the sun
had the exposed portions tinged with pinkish -brown, but this colour
eventually disappeared. Form very elongate, shaped somewhat like
a banana, being curved on one side and almost straight on the other
du straight side with a broad deep groove which widens at the
anterior end giving it a spatuloid form. Surface with faint polygonal
reticulations. The larva effects its escape by Splitting the broad end
of the groove, leaving it slightly raised (see PI. v. fig. 4). and apparent!)
intact on the opposite side.

Length, 1 mm.

on a bright sunny day during the

8i

a small dentate sclerite ; the hypostomal sclerite articulates with two
large bifurcated cephalo-pharyngeal sclerites, and in front of the upper
arms of these pates is a small perforated sclerite. In a freshly-
prepared specimen both the retractor and extensor muscles to these
sclerites can be distinctly traced. Ventral surface of the last seven
segments furnished with raised bands of tactile tubercles. Posterior
stigmata two in number, circular ; thoracic stigmata placed sub-Iaterally
on the third segment, each consisting of apparently- five circular
orifices, these are connected posteriorly' with a large bilateral air sac
which extends along the fourth segment.

Length of adult, 1 1 mm.

Young larvae are much more transparent and glass-like, and the
large anterior mouth-hook is not developed, a blunter process taking
its place.

Habits of the Larvae. — They move rapidly along a smooth
surface, pulling themselves along chiefly by means of the large mouth
hook ; and proceed practically in a straight line, moving the head
rapidly but irregularly from side to side or up and down. There is,
however, no regular alternate movement of the head during progres¬
sion as in some muscid larvae. Their progress through the burrows
in their food is much more rapid than on a smooth surface, and when
disturbed they disappear with extraordinary rapidity. The larval
stage lasted, under favourable conditions, from 14 to 21 days; but the
absence of excessive moisture and the admission of a little light
materially retarded their development, which then extended over a
period of from 31 to 78 days.* The larvae exposed to such conditions
produced much smaller pupae and correspondingly small imagines.

Method of Pupation. — This is completed in about two hours
At first the larva rapidly shortens itself, chiefly by contracting the
anterior segments, and becomes barrel-shaped. At this period it is of
a creamy-white colour, and the mouth parts of the larva are still visible
through the soft integument. The colour rapidly changes to bright
ochreous, and in the space of two hours or even less, the integument
hardens and the puparium assumes its normal colour. In cases where
soil was placed below the faeces the larvae generally burrowed into
the former to a depth of about half an inch, but a few also pupated

* A few specimens still remain in the breeding cage, and may nossihlv nas- th<
winter in this stage.

F

8 1

in the dryer portions of the dung-. Where no soil was provided the
larvae generally' pupated at the bottom of the breeding cage.

The Puparium or Pupa. Colour bright terra cotta red.
changing to dark chestnut-brown a few days before the emergence of
the fly. Form barrel-shaped, slightly narrowed in front, broadly
rounded behind. Eleven segments only are visible, the anterior oik
bearing the minute bilateral thoracic stigmata of the larva; posterior
segment with two large disc-like stigmata ; all the segments with fine
transverse striae, the striae, under the microscope, producing a slight
irridescence ; dorsally the articulations have a double series of minute
papillae, one series being more minute than the other; the posterior
segment has also a median longitudinal series which terminates
between the stigmata; ventrallv the papillae are replat ed by a regular
series of fine ridges, forming a distinct and relatively broadband; the
ast segment also bears a somewhat lunula* or angular shaped fat d
m°re °' 1(SS rounded papillae. This stage lasted from q to 13 days.

Length, 5 to 5-50 mm.

comparatively dry faeces produced m

en~e°oPfTT OF ™E NYMPH A before I

SI ' mSeCt the CUtlcle °f ‘he puparium darkens a

transversely alOT^thTf ''rth'0”6 ^ k1"*1 a”d niedian lines iwd a
% escapes Prior ,o d S®""* ’ ^ fa"s "* 1

pushing its effete skin rfTi 1 ? nymph undergoes its final ecdyt
Puparium. On im Awards into the posterior end of t

thick rudimentary wingTtf " ,ai^ars as a sma11 dark-grey fly, wi

m the mid costa, below Which are” t leaden r0lOl,r Hnd a decp n0t
15 much larger and wid i S °ng convoluted folds. The he;

attenuated. Its subseauent J ^ 1,10 thorax» ^d the abdomen
mto the following stages •- °prnent ina>' he conveniently dividi

an hour. Durino- Pe.r*0d* vvhicli lasts for approximately ha
whole of its attenf ^ Pen°d the insect devotes nearly tl
If Placed in a g-lassTV0 ^ CSCapc from its environmen
ln 't, the fly imrn„jU 6 witl1 a barrier of loose cotton wo
through it, ancj tL • . la e y endeavours to effect a passag
accomphshes with marvellous rapidit;

83

making headway by constantly inflating the frontal sac, at
the same time pushing itself forward with the legs. When
liberated, a great deal of time is devoted to combing out the
hairs on the arista of the antennae, this being accomplished
in the following way : the head is turned either to the right
or left, as desired, and the frontal sac is then inflated on that
side farthest from the thorax ; this process lifts the antennae
into a prominent position, and the long hairs of the arista
are then rapidly and carefully combed out with the under
surface of the anterior tibiae. The frontal sac also receive^
a share of attention, and so also does the abdomen, and
occasionally the rudimentary wings. This stage is remark
able, in that nature so provides that, under normal conditions,
the insect may successfully escape from its larval habitat
before the wings develop, and so impede its progress or
render its escape impossible.

2. In this stage the frontal sac is usually contracted, and the head

presents a more normal condition ; the fly also becomes
quiescent, and remains as a rule in a fixed attitude, with the
legs well displayed, and the head extended forwards, so that
the narrow neck is stretched to its fullest extent. Air is then
pumped into the body by repeated and alternate contractions
and extensions of the abdomen. The body increases in size,
and the integument becomes extremely pallid in colour.
At this stage the wings are apparently filled with air, which
passes rapidly along the costal region, then across to the hind
margin, and finally the tip unfolds, sometimes aided by the
use of the hind legs. The first portion of this stage some
times occupies over twenty minutes ; but the wings develop
as a rule in about three minutes.

3. 1 he fly still remains more or less quiescent, but gives some

attention to cleaning itself, and when the integument and the
wings are sufficiently hardened the proboscis is raised from
the ventral to its normal horizontal position ; when this is
accomplished the insect takes flight.

Defaecation lakes place shortly after the imago is perfected ; the
faeces being milk-like both in substance and colour.

Summary of Life Cycle.

Larvae fed on moist sheeps' dung.

Eggs procured from captive females.

Average day temperature - . j2° F

Average night temperature o<?° F

Month - August.

Ova— Incubation period - 2 ,

Larval stage - _

Pupal stage -

Complete cycle

-21 ..
9 -13 «
25—37 h

Summary of Life Cycle

Fo°d allowed to partly dry and some light admitted,
temperatures as above, during the month of August.

Ova— Incubation period -

Larval stage - . - - 2 3 days

Pupal stage approximately as above
Complete cycle

■Some larvae of this brood may hibernate through the winter

Additional Field Notes.

search for the larvae ITJriou^ . the Press 1 continued

Chester, and on 21st SeDtemh 5 ^ aces a few miles radiu;

finding the insect in all sta»e my eff°rtS Were at lasl rewarded
on the outside of a cucumber b H “?)tamorPh°sis The habitat 1
males and females were djsnorf ' fu R°SSett' Flintshire. Here b
“d also on some matting usedT themSe,ves on ‘lie cucumber fra
was first g.ven to an exfm“ej cov“ing the glass. My attend

ho‘-bed. but this was quite “Id r ““ S‘abIe — forming .
7"ld b/' to the ratheTd v n ?Ve' “ 1 Ve"tIlred >° ‘l™k that
a^gs,de the cucumber bed waf a h ^ ^ " a,ls But **

recem,Ultd dU™^ *e sealn ?? °f «"■»»»**, which In
this tl y bCen deposited, and this w 6 uppermost layer had on

tered a\graSS Was <Wte rotten LT ‘LT h0t (9°° to 9S° F ), belo
temperature of ;o° F n '"th moisture, but reg>

F- It was ,» this layer that 1 found ?h

5

first lot of larvae. They were chiefly full-fed, and some of them
pupated during the next two or three days. Continuing my search
further I also found numbers of larvae and pupae, in the still older and
quite cold deposits, some of them were only a few inches below the
surface, others were deeper down ; they sometimes occurred singly,
in other cases several were found together, some were mature, others
only partly developed. On disturbing the newer deposits they
naturally gave off the strong smelling fumes characteristic of heated
grass, and this produced a result which I had neither hoped for nor
anticipated. A female Stomoxys was seen to alight on the hot and
freshly disturbed grass and to quickly disappear among the interstices,
there she remained for a minute or so and then flew away. The grass
was carefully examined, and amongst it, at a depth of nearly three
inches, were found a number of her eggs. A regular succession of
females then followed, and very soon three of them were engaged in
laying their eggs in a small area which could have been covered with a
crown piece ; the first-comer being not in the least disturbed by a
companion running completely over her body. In all cases the
abdomen was depressed, and pushed into the material as far as
possible, and in two instances the wings were partly extended, in
order, apparently, to secure a firmer support. One female remained
in the same spot for five minutes, and then changed her position to
another a few inches away. A second female was occupied for twelve
minutes in laying her eggs, but she did not change her position during
the time. All three females flew away immediately afterwards, and
did not seem in any way weakened by the process. For how long
they survive in a state of nature it is impossible to say, but some
females that were caught immediately after egg-laying died on the
fourth or fifth succeeding days. Some females which were caught in
glass tubes as they alighted on the grass laid their eggs immediately
afterwards, and these also survived until the fourth and fifth days
Reference has already been made to the almost noiseless flight of
these flies when disporting themselves over and about their favourite
haunts during the heat of the day. But 1 found that when the females
were negotiating the habitat previous to laying their eggs the noise
was distinctly audible, and resembled the characteristic hum produced
by other muscids.

8f>

The day on which the foregoing observations were made was a
delightfully bright and sunny one, and the hour from 3 to 3 45 in tin
afternoon ; later, when the sun had lost its power, the flies
disappeared, in their usual way, to find some sheltered spot in which
to pass the night.

The eggs which were procured on this occasion were kept at a
temperature varying between 64° and 67° F. ; under these conditions
the larvae did not begin to hatch until the c ighth day, thus the
incubation period was greatly prolonged ; had they been left in the
warm grass, where the temperature near the surface was 700 F., they
would in all probability have hatched, as they .lid in til- summer
months, on the second or third day.

Whether the larvae of these autumn broods will pupate before
»e w.nter is at present impossible to say, but judging from the higi

foS Ti • habitati wh,ch w,n i'c,,'iml> •«

we shall a. V** reasonable to asaw»e that they will do so, and

stage * Am ^ ^ th* winter ** chiefly in the papal

tesTltinf, iTT disturbance of the habitat in question would

T °f 1116 artificial heat ■*» • . . * to toe

hibernate throuo-lTT™*" ^ SUCh a Case tlle ,arvae would probably
early summer. Fortuna'tdv'tl and PUpate in tlie following spring or
as the owner has verv Id Ji • 1S 001 I,keIy to ”> this instance^

whole of the material '^l ^ glVen mf un disturbed possession of the
tinue the observations’ thro ^ ?, WlI1\ lt ,S ,lopedl be possible to con-
economic importance can ^ 7 ' W,nter and spring, though little of
this country. n°V' e addcd to the habits of the insect in

Localities ^tomoxys sitiens , Rond.

Circumstances of Lulon&° J Nyangwe ; Kasongo.

and ln a European house Ti f pecimens were caught about cattle

U teeds vigorously.

A large ser' S*omoxys sitiens ?

J he>’ Were P^served in alcohol "T' C3Ught on catt*e at Zambie.

_ 1C°h01 and canr,ot therefore be definitely

87

Stomoxys omega, n. sp. (Newstead)

(PI. iii, figs. 2, 3)

Wings dusky, strongly' irridescent in a bright light. Thorax
blackish, shining, anterior half in front of transverse suture pale
greyish-blue, with a distinct JVshaped black design in the male; in
the female the submedian black lines blend with the black colour in
front of the suture so that the up-curved terminals of the /V shaped
markings become indistinct. Abdomen, smoky-brown.

Male. Head: Eyes almost meeting, the narrow space between
them, above, velvety black ; frontal margins and clypeus silvery -white.
Basal segment of antenna blackish; second segment brown; apical
segment silvery-grey ; arista brown ; posterior surface of head velvet)
black ; lower angles silvery- white with black hair. Proboscis black,
shining ; labella with many, very long, fine outstanding hairs.
t horax in front of the median transverse suture grey-blue, with a
large /^-shaped black design, consisting of two distinct broad sub
median longitudinal stripes which curve upwards before reaching the
suture and form two conspicuous rounded dilations ; thorax below
/he transverse suture black, shining, with a rather broad band of dusky
grey-blue in front of the scutellum, which is continuous to the base of
the wings ; scutellum smoky-brown, shining ; with one central and
two lateral black bristles ; the thorax is also clothed with fine blackish
hairs, and there are three bilateral black bristles at the sides in front
of the wings; pleurae in front delicate grey-blue, a small area at base
of wing pale brown. Abdomen hairy; smoky-brown, the segments
gradually darkening towards the apical margins, especially so at t In¬
sides ; in perfect specimens there is a broad basal band on the second,
third and fourth segments of faintly greenish-grey dust, each band is
interrupted in the middle ; the last segment almost entirely clothed
with a paler grey dust ; venter pale dusky-brown. Legs relatively
long, smoky-brown, blackish in some lights ; knees pale brown

Length, exclusive of proboscis, 6 mm. ; length of wing y mm.

Female. Head with the space between the eyes relatively
narrow; vertex, blackish, shining; margins and clypeus pale bluish
grey, bright but scarcely silvery ; posterior surface of the head similar
to that in the male, but the sides below have a broader margin of
blue-grey. Apical segment of antenna dusky-grey. Thorax in Iron!

x.s

of the median transverse suture grey-blue, with two short, wdl
defined submedian black lines in front, these gradually merge
into large and somewhat ill-defined black areas, leaving the blue-grey
ground visible only at the sides and between the black longitudinal
bands; thorax behind the suture as in the ; scutellum black, paler
in the centre, shining ; pleurae and legs as in the g . Abdomen
similar to that in the male but without the grey , dust-/ ike margins.

Length, exclusive of proboscis, 6 mm. ; length of wing 6*50 mm.

Easily distinguished by the beautiful pale blue 0< th<

Of the thorax, with, in the , , the distinct black ^-shaped pattern.

The relatively narrow space between the eyes in both sexes is
also noteworthy, these characters being much more pronounces! than
in the species of the ca/citrans type.

Localities : — Ukungwa ; Send we.

°f Cat‘Ure:~ 'Th,S Ry was **" only near water

Which ,hevCaUg VnjCan0eS' °thers °n a buffal° shot in a marsh, on

S half co /eedlng' Wh6n th£ buffalu "as fi-. seen it was

w e n m Water' n° d°”bt ‘0 avoid the S,omoxys. which

were present ,n almost incredible numbers.

CUTANEOUS MYIASIS

? Ockrornyia anthropophaga (E. Blanch.), in man.

specimens of a laLT^^b ^ Grenade of Leopoldville for three
taken from a coloured ° °Se^ resembhng that of an oestrid
of them occurred ln the h°Spital al Leopoldville Two

These larvae aPTee' ,, °ther in the neck-
anthropophaga, E. Blanch ^ I ^ cIescnption given of Ochromyia
°r a closely related one. ^ 1 1C> Posslt>ly belong to this species

The fo„o • * ‘ ^ F'y «» * **

- t * - — taben fn

eventually lead to the idem fi ^S°ng°’ Feb™aIy '6th, ,90.
been unable to fix it. “‘““‘“‘“a of the insect; so far w

Description _ Le

rather suddenly taperfng ” Znt but slightly flat.

front, rounded behind but no, ,n

8<)

Segmentation very pronounced and regular above but convoluted
ventrally ; all the segments, with the exception of the antepenulti
mate which carries the anterior stigmata, almost covered with ven
short and rather blunt spines, the majority of which have dark brown
or piceous tips ; these spines are arranged in short straight transverse
lines, each group consisting usually of from three to five spines, in
some cases there are but two and in others more than five. The great
mouth hooks are black ; and there is a lateral palmate group of
external teeth as in the larva of Auchmeromyia luieola (q.v.)
Posterior stigmata on the last segment, subdorsally placed, and very
close together.

Further details cannot be given, as it has been thought well to
preserve the specimen intact.

Larvae of a Muscid Fly in a Mule

The mule illustrated in fig. 16 was brought to us at Kasongo. In
the centre of a large and very conspicuous oedematous swelling on its
abdomen was a deep-seated ulcer, some three inches in diameter. On
inspection, the whole ulcer, below the level of the epidermis, was found
to be so closely set with the larvae, described below, that it was
impossible to see any part of its base.

The mouth parts of the larva were invariably directed inwards and
the bodies of the parasites were so deeply buried in the tissues of the
host that their posterior extremities were alone visible. It was found
that there were two layers, one above the other, of larvae, packed side
by side closely together. It seems probable that they were able to
burrow in the tissues of their host, since a few larvae were found in
the sloughing tissues at a depth of some five centimetres from the
apparent surface of the ulcer. None of the larvae penetrated into the
abdominal muscles. About 160 to 180 larvae were taken from this
lesion, which healed quickly after a thorough curetting and dressing
with iodoform. A description of the larva is herewith appended :

Length n mm.; greatest width 2*25 mm.

Colour, in formol, dull pink, with the anterior segments paler ; some
examples were almost white. Form spindle-shaped, tapering from
the mid region almost to a point in front, and slightly so posteriorly
Great mouth hooks prominent, unidentate, and black. Segmentation
very pronounced, each with a strongly marked ridge forming

a complete ring round the body ; all the ridges furnished with short
brownish-coloured spines, which to the naked eye appear as dutinr.
equidistant rings; anal segment somewhat truncate, hut deeply
wrinkled, and furnished dorsally with four very short papillae and
ventrally with two much longer ones \nal stigm.it

castaneous, almost touching ; irregularly ovate, with three transvei*
slits.

Attempts to raise the fly from these larvae were unsuccessful

-rauscid Fly.

family hippoboscim

ff’PPnbacsa equina, Linn.

T A exampies Qf . (Figs' V. 18)

5 Pa'maS' C— y Island's, ThiLTr fOUnd °n ^ *

°n board sh'P on their way to the

9i

Congo Free State (September 15th, J903). This is an important
record, and furnishes a further proof of the ready means by which
these parasites may be introduced into a new and uninfected region.

Flies of this genus were also caught on camels and cattle at
St. Louis, Senegal, May, 1903 ; their freshly deposited larvae were
marked with very delicate tracings. Unfortunately the specimens
and drawings were lost while in the hands of the Authorities of the
British Museum and cannot now be traced.

Fig. 18. — Hippobosca equiriii
Puparium about twelve
hours after extrusion.
At this stage the anal
tubercles are black; the
rest of the integument
terra-cotta red. x 6.

Lipoptena* paradoxa, n. sp. (Newstead).

(Figs. 19, 20)

FEMALE. Specimens preserved in Canada balsam and alcohol are
bright red-brown inclining to orange-brown at the sides of the
abdomen ; claws black ; base of abdomen with a bilateral patch of
darker chitin, the median area of the remaining segments also with
darker markings, but these are both irregular and inconstant in the
preserved examples. Head as wide as the anterior part of the

Lipoptera of Siebold and Loew.

thorax; ocelli absent. Mouth parts rudimentary Outer
of eyes with a double series of spinose hair* Thorax nank¬
in front than behind, with a submedian series of ate

nine long spinose hairs forming a curved line, and a ste
submarginal series of usually four similar ones terminating oppts
the insertion of the mid legs; posterior margins with four loe;
spinose hairs on either side of the scutellum ; the last wire
organ ,s also furnished with four similar hairs Abdomen short oral
almost sub circular, with numerous spinose hairs arranged

* poradoxa.

hi the figure Vent • * ~~‘ ~ •*« **■

7nVfX with numerous mUnerOUS ^ Spinose hairS; '
s ender spinose hairs the n,inute equidistant tubercles t
strongly rugose. Le ,^Paces between the tubercles fine

l™ymg len^hs and vfryino- d’ StOUt, Sparsel>' Allied with hi
not extending beyond of thickness; the postenc

pui •,1,11Ia Ie&s stout ; tibia/3 • k° abdomen tibial spine to ai

fea h lbr°ad1^ da ted ' ‘° P°Ster,°r »***»* *

Serots"S,le Str°n^ ^ose tTlddle °UtWard5'

irrep-i l Pmes* tbe inner with U lIPPer surface with onl>
7 toothed on the inner targ^^ ' ungues very fa inti;

93

Length 4 mm. ; width of abdomen 2 mm.

Habitat : Taken from an antelope at Kasongo, January 28th, 1 905.

All four specimens are females. The same host also harboured a
number of ticks, (q.v.)

The absence of ocelli in the female is rather remarkable. There
is also an almost entire absence of external mouth parts, including the
labial sheath ; the only indication of these organs being a minute
truncated cone, the exact nature of which could not be determined in
the limited supply of material.

FAMILY PULICIDiE

Dertnatophilus ( Sarcopsylla ) penetrans, Linn.

(PI. vi, figs. 1, 2)

Larva long, cylindrical, of fourteen almost equal segments. Head
slightly longer than the second segment, narrowest in front, but rather
widely rounded. Antennae short, apparently of three segments, the
basal and apical segments very short ; apex with a central long slender
spine and two to three minute ones. Surrounding the base of
the antenna ventrally is a semicircular group of blunt spines, the two

nearest the base of the antenna much the largest Mandibles Urge
slightly curved, unidentate. Cuticle below the buccal cavity with •
curved row of short, stout, and backwnrdly curved spines ; on either
side of this series of spines are two divergent Sub Utai
the ventral one extending almost to the articulation of the head Vid
the thorax ; the upper sclerite about half the length of the voW
one ; both are narrow and rod-like ; the base of the ventral sdmt

15 a'SO much d,lated’ and ‘he point where it meets the upper®
IS unequally bifurcated and spine-like. Cuticle of all the ague
presenting a stiongly marked scale-like appearance; each segment s

; '7 u a Sing,e transver- Of long ... pud, sun, has
f i r" ,, art,CU'ati0n5: "™' -gmen. bilobed. each *
lobes are ^ St°U‘ spinc and numerous hairs fa

.mmttufela^0 u ’ ^ TheSe mems are from appart:

small (i mmHt 'is'h-^r* U’at th<? SeXUally mature insects are very
much greater' dimensions^ Pr°bab^e - ‘he larva docs no, *

This pesfisrfoInd6rnientS g‘Ven 'S inclusive of 'he head
Theywl slrtoPbrtlCal,>; everywhere in the Congo,
that at Rutshuru and Re P entlful at Nyangwe, and we were tok
The larvae are ^ a P^Rue.

a prolonged search dlfficu,t to ,ocate ; and it was only after

1 hey occurred ainonp- n, "T SU^ceeded in finding three specimens

was literally swanning with chigg^ ^ fl°°r °f * nat,Ve hut WhiCh

family phdiculid^

This louse was Q“r‘

Leopoldville and KasoLo^T.,0" C3SeS °f human trypanosomiasis*
wider. s°"g° Their distribution is mo^Lbably W*

These vermin '**"*• Leach,

distribution is very^^noticed^ Bom, and LeopoIdvi|fc ; ,h„

95

ORDER HEMIPTERA

FAMILY CIMICIDiE

Cimex lectularius (Common Bed-bug)

Localities : — Banana (Dr. Etienne) ; Tshumbiri (Rev. Billington) ;
Nouvelle Anvers (Dr. Miiller) ; Tshofa; Kabinda; Lusambo.

Circumstances of capture : — Specimens were taken from the cre¬
vices in the cane beds and grass-cloth pillows of natives.

ORDER HYMENOPTERA

Melipona, sp.

One or more species referable to this genus were seen in several
places in the Congo Free State. They were particularly numerous at
Leopoldville and Dibwe. They caused intense annoyance by the
persistent manner in which they swarmed about one and crawled into
one’s mouth, eyes, nose and ears. When crushed they emitted a
peculiar and characteristic odour.

At Leopoldville a colony of these stingless bees had taken up their
quarters in a large lock and rendered it quite useless by the great
accumulation of wax which they had formed. In this instance it was
interesting to note a waxen tubular opening, about 4 cm. in length
and about the thickness of a pencil, projecting from the keyhole. A
species of Melipona has been noticed to have very similar habits in
the Soudan.*

INSECTS OF NON-ECONOMIC IMPORTANCE

In addition to the insects dealt with in the foregoing chapters of
this Report, a large number of insects, representative of various orders,
were also collected ; but as they are of no economic importance it has
been thought desirable to deal with these elsewhere. We would add,
however, that among the more remarkable species is a new Dejeania
which was discovered among the Glossinac in the fly cages at
Kasongo. It had evidently been caught by one of the boys in
mistake for an engorged tsetse, which it very closely resembles. We
also obtained several specimens of a rather remarkable species of

* 2nd Report Wellcome Research Laboratory, Department of Education, Sondan
Government, 1906, page 89.

96

Notocanlhis of a uniformly blackish colour, with numerous white spots
on the wings. These flies were only observed in one locality in the
bush about a native village situated on high, dry, sandy soil, near
Lisala (July, 1904). We possess no particulars concerning its habits.
Whilst at Boma (Oct. 27, 1904) we also found a number of larvae or
rat-tailed maggots apparently belonging to the genus Eristalis.

ORDER ACARINA

FAMILY HYDRACHNIDAi

Ectoparasites of M osquitoes

ie larvae of four distinct species of Acari belonging to this
family were observed on several species of mosquito. The site
prefei red by these ectoparasites was invari
ably the dorsal area of the abdomen, and
generally, on the first few segments. Most
of them, though not all, were fixed to the
articulations of the segments, to which they
seem firmly attached. The mosquitoes most

were

attached to them In 6 Parasites

as five were a T ““ 38 m*"y

on the articulation "of g tOS:ether

The species met with ,!”S * Segmem-

dry) is of a bright °n h* ^ (When
with a dull oranv/ . °chreous colour

large black eye-spots a"d rather

Kumba, Kasonga and ^sambo0031'^ '°re

^.ed^tefr^ti:^ percent^ _ — -

one of a n«;f — , leie were three wpII i j

ree well-marked species

. /' \

I’lG 21. — Ectoparasites oj
Mosquitoes, x 20, about
From a sketch by the late
Di. J. E. Dutton.

of these insects als

. ’ vvuicn t lip 1-0 . ” * “‘MV. 1113CC-L3 a 1

e o a uniformly orange-red colo ^ WeI1_marked specie;

or pale ochreous. In llff another of a creamy-whi

' 'eS: r'eoP°ldville( Tshumbiri/K^. aCands Wer<? pink °r yeI1°'

pink _
asongo and Lusambo.

97

FAMILY SARCOPTIDA*

A case of Symbiotic mange occurred on a rabbit which was
imported from Europe for experimental purposes. The condition was
se\eie and the external ear was entirely filled with the parasites and
the very profuse epithelial desquamation they had excited. On
macerating the dead specimens in potash, one finds that they are a
species of Symbioles (now referred to the genus Chorioptes) and agree
best with the descriptions given of Symbiotes communis var. cuniculi

FAMILY DERMANYSSIDvE

Pnemnonyssus duttoni, Newstead and Todd

Memoir xviii, Liverpool Sell, 'l'rop. Med. p. 4 (with plate), 11)05.

Since the publication (l.c.) of the description of the adult mite
Dr. J. W. B. Hanington has been successful in discovering the earlier
stages of this acarid, and a description of them is appended below :

Ovum. Rather narrowly ovoid ; faintly yellowish-white and, as
seen by transmitted light, rather granular in appearance.

Length -32 mm.

Hexapod larva. Short ovate, narrowed in front, widely rounded
behind. Capitulum prominent. Palpi of five segments furnished
with short slender hairs with the exception of the apical one, which
has several long stiff hairs ; basal segment broadest, but scarcely
longer than the apical one; second, third and fourth shortest and
nearly equal in length ; formula (1,5) (234) ; tip of the apical segment
reaching to the middle of the third segment of the anterior legs.
Legs directed forwards; all the segments with whorls of hairs near
the articulations, those on the basal segments are mostly short and
slender, while those on the remaining segments are long and stiff .
tarsi longer than the two preceding segments; all with very Ion-
hairs, and a few short slender spines; claws simple, placed on a long
pedicel which is much narrower at the articulation ; pulvillus scared)
wider than the dilated claws, but the end extends beyond them.

Length '50 mm.; width *30 mm.

c;

9s

FAMILY TROMBIDID^E

The larvae of an acarid belonging to the genus T rombidium were
found in considerable numbers attached to the naked skin on the
muzzles of horses at Kasongo. They appeared to the unaided eye as
minute crimson specks, and were evidently the cause of an intense
eruption. I he skin was pink, rough, and doited here and there with
tiny' spots of dried blood ; still attached to the skin or on slightly
raised hairs were numerous profuse crusts of desquamated epithelium
and dried serum. Although a careful search was made the parasites
were not found elsewhere on these animals nor on any other host
in that locality.

1 he larval forms of these acarids are well known for the intense
itching and soreness which they cause by burrowing under the skin
of man and other animals. It is generally an unnatural position for
utes, and .is a rule they soon die ; though one species which is

° Cn °Un °n tllc ears of cats in this country (Great Britain) lives for
many days causing great annoyance to the infected animals.

a ,U \ lnite IS not Parasitic but predaceous, living on small

rCc^me ,eoc:rwa?er:nIy, wult form coi,cr,ed in ,,w

S^^:.c2Stocrd with ■ bn,"an*

as to appear quite velvety in texture.

FAMILY IXODI DJE

new and undescr^ed^^ ^ ***** catalo&ued include two that are

I he only member of the Arrm -j
Ornithodoros mouba/a • tl • g dae which we met with was
described and figured in lls imPortant species has already been
fever. t We therefore onl ^ a fP°rt °n the nature of Human Tick
bionomics. 11 y a t here a few additional notes on its

A few hundreds of the

•September, 1905; they }) & . 1C S Were brought to Liverpool in

temperature of between 190° been keP! in an incubator at a

f'eely, hut under the^ 2 ^ wkere they have reproduced

Editions the eggs take fwo or three day<

1 nis —

* This col, - ^

1 Liverpool School of TropfcalM«If ?deQtified l,v G. Neumann.

opical Medicine Memoir XV, I (Plfttes arul luap.

99

longer to hatch-out than was noted in the Congo. Many of the
original ticks are still alive ; they have of course been well and
frequently fed. One female raised in the laboratory from an egg has
now been under observation for 25 months; during this period she
has moulted six times. This seems to be rather less often than is
usual, since ten out of fourteen ticks raised from the eggs moulted
six to nine times during the year. Not infrequently ticks do not
seem to be able to shed their old skin, and die while moulting. As
a rule the ticks would not feed properly more often than every seven
to ten days. I icks remained alive on several occasions without food
for four months, and in one instance for over six months. Adult
ticks caught before April, 1905, or their progeny, which have never
been fed upon an infected animal, are still (November 22, 1906) able
to infect susceptible animals by their bites.

A mblyomwa hebraeum (Koch), var. splendidum , Gieb.

i' orty-three males and eight females were taken from a Buffalo
(Bos nanus ) which was shot in a marsh near Tshumbiri. In life the
ground colour of this beautiful tick is of a soft green, with brownish
ornamentation ; there is little or none of the irridescence so
conspicuous in specimens preserved in alcohol.

Amblyumma variegaium, Fab.

On cattle, Kasongo (Feb.). Associated with Hyalomma aegyptium,
Rhipicephalus sanguineus and Margaropus annulaius.

Amblyomma variegatum ?, Fab.

A single male was caught on a bovine at Kasongo (Feb.).

Hyalomma aegyptium, L.

I liree females were caught on cattle at Kasongo (Feb ). They
were associated with the preceding species.

Haemaphysalis l each i. And.

I his tick is the recognised carrier of Malignant Jaundice
(P iroplasnia canis) in dogs.

Thirteen males and two females were taken from a leopard at
1 shumbiri by the Rev. Billington (June); and Dr. Etienne sent a
single specimen (host not mentioned) from Banana (Oct.).

I OO

M argu r opus annnlatus , Say.

Common on cattle at Kasongo (Feb.). Xainbir and CoquilhatvjOt
1 lie specimens collected were all females.

M argaropus annnlatus , var. calcaratus, Bir.

In all the active stages, on cattle at t'oquilliatvillc.

Rhipicephalus bursa. Can. and Fan/.

Collected by the ( lief de Poste on cattle at Nya Lukemba to the
North of Lake 1 anganyika. A single female was also caught on a
bovine at Kasongo.

1 liis species is the carrier of Carceag or Malignant Jaundice it
sheep.

Rhipicephalus capcnsis, Koch

Li^lit females and four males were taken on cattle at Nyangwe.
Rhipicephalus nitens, Neumann

Locality. One male and three females were taken from an
antelope (L ragelaphus script us) shot at Kasongo (Feb.).

Rhipicephalus shuns , Koch

- t0 °ne 0f the carr'ers of Pin, plasma par-, urn, Theto

■■.ales at BananaTorDrEt! “T ^ ’ ^ **

obtained in either case’ ’ e)' 1 he na",c of the <«** *'

L RhiPlcephalus sanguineus, Latr.

named; at Kasono- ^Ct^’ co^ected by Dr. Etienne, host no’

preceding species and on catt,e ; occurs in association "'ith

In addition to the ih

were also discovered UV° Spec*es °f Ixodidae new to scieno’

succeeding paper. CSC m ^escr>bed by Prof. Neumann in tin’

Plate iv. . r . ERRATA

M«P «• Delete ^ Tabanid^-

pages!,"

102

EXPLANATION OF PLATE l

Mi mom via MALKEYTI. (Page 29.)

Fig. I, Head of female showing the chi nbutionof

the scales, x 25, about.

Fig. 2. Tibial spines of male. ><75.

Fig. 3. Genital armature of male (left half). * 75

MlMOMYIA AFRICANA. (Page 28.1
Fig. 4. Head of female showing the charactd and distribution of
the scales, x 25 about.

Anisogheleomyia QUADRIMACULATA (Page 32.1
Fig. 5. -Head of female showing the character and distribution 0:

the scales. x 25, about.

Fig. 6. Proboscis of the female. Enlarged.

NEOMELANICONION PALPAI.E. (Page 31 )

Pig. 7. Head of male showing the character and distribution 0:
the scales, x 25, about.

Pig. 8. A few segments of the antennae of the male. Enlarged,
f ig. 9. Male palpus. Enlarged.

Eretmapodites INORNATUS. (Page 12.)

1 ig. to. Ferminal segments of male abdomen. Enlarged.

CULEX PAR. (Page 25.)

h's' 11. End of Abdomen of female (ventral). Enlarged.

All the figures are reproduced the same si/e as the origin3'
drawings from which they are taken.

PLATE I.

it.

#’ fv»*. /« rf

Xewsfetnl, ml mil. del .•

EXPLANATION OK PLATE II

Boycia mimomyiaformis. (Page 34.)

l ig. 1. Head of female showing the character and distribution of
the scales. x 25 about.

I ig. 2. — Head of male showing the characters and distribution of
the scales. x 25, about.

Fig- 3- Palpus of male. Enlarged.

Stegomyia ALBOMARGINATA. (Page 16.)

1 ig. 4. Head of female showing the character and distribution of
the scales. x 25, about.

Stegomyia luteocephala. (Page 15.)

I ig. 5. Head of female showing the character and distribution of
the scales. x 25, about.

I 1UINIA 1ARSAL

hig. 6.— Anterior tarsus of male. x 75.

1' 'g- 7- Palpus of male. Enlarged.

big. 8.— Genital armature of male. x 25.

drawings T ‘"C reProduced the same size as the original

drawings from which they are taken.

ml mil.

/

del.

r

A’. Xewsteml,

/• l». /*»*.. /«*

PLATE II.

.

EXPLANATION OE PLATE III

Tabanus billingtoni. (Page 46.)
Fig. 1. Female with wings displayed. x 2$.

Stomoxys OMEGA. (Page 87.)
big. 2. Female with wings displayed. x 4.
hig. 3.— Male with the wings omitted. x 4.

Stomoxys calcitrans. (Page 75.)

I'ig. 4. —Female with wings omitted. x 4.

(In cutting away the background, the block maker has
inadvertently removed a portion from the left
side of the abdomen. R.N.)

Glossina MACULATA. (Page 7;.)

Fig. 5 -Female with the wings displayed, x 4.
ig. 6.— Female with the wings at rest (profile), x 4.

Fig.

Fig.

Fig.

Glossina palpalis. (Page 57.)

7— Female in the act of parturition, x 4

8- Puparium before the escape of the imago.
9. Pupanum after the escape of the imago

All the %ures are reproduced ^
drawings from which they are taken.

same size

x 4.
x 4.

as the

original

ft Wtt 'stead, ad imt. del.

—

•

' ci

-

_

Fig. i.
Fig. 2.
Fig- 3-
Fig. 4.
Fig. 5.
Fig. 6,
Fig-
Fig. 8,

Fig. 9,

Fig. 10.
Fig. 11,
Fig. 12.
Fig- 13-
Fig- 14-
Fig. 15.

All

direct.

inS

EXPLANATION OF PLATE TV.

Chrysops dimidiatus. (Page 44.)

FIae.matopota trimaculata. (Page 4^.)
Haematopota DUTTONI. (Page 41.)

Tabanus DORS1VITTA ? (Page 44.)

Tabanus UNIMACULATUS. (Page 46.)

Tabanus par. (Page 45.)

Tabanus pluto. (Page 45.)

Tabanus rufipes. (Page 45.)

Tabanus canus. (Page 44.)

Tabanus billingtoni. (Page 46.)

Iabanus billingtoni. (Page 46.)

Pabanus billingtoni. (Wings displayed.) (Page 46.)
Iabanus tarsalis. (Page 45.)

Iabanus fasciatus. (Page 44.)

1 ABANUS GABONENSIS. (Page 45 )

g res nit the actual size of the originals, photographed

PLATE IV.

II. N. Photo.

14

A FRICA N TA BA NIDE.

I IO

EXPLANATION OF PLATE V.

Illustrating Mr. Newstead s paper on “ 1 he Life-history of Stomotys

calcitrans , Linn.” p. 75.

J - Eggs twice natural size.

l'ig. 2. — View of the curved side of the egg. x 65.

* E ^8g in semi-profile showing the deep spatulate groove
x 65.

g 4 Empty egg as seen in profile, with the semi-detached capsule
at the anterior end. x 6s.

g 5 Dorsal view of larva showing the intestinal tract and course
of the main tracheae. x y.

Fig- 6- Thr!e te™linal see™ents of the larva in profile, with the
n erna mouth armature: an , antennae; m, muscles;
, per orated sclerite ; mdt mandible or great hook;

’ ypostomal sclerite ; cs, cephalo-pharyngeal 9derites;
v*> ventral tooth, x 60. 7 *

» » ‘he rred out Reference

^ " nthoeraclrmstnaI SegmemS °f ,arva' dorsa1' con,

Other refe trachea fonning internal a

" referCnce ,etters as in fig. 6. x fho.

lir\^ w’ .

F '8 ^ Puparium or pupa. x 7.

rn

ews^ad,d«]

d-1- r.at.

n.

Plate V

CALCITRANS.

Huth Lithr Lard

oun.' ...

771/

I I 2

EXPLANATION OK PLATE VI

Dermatophilus (Sarcopsylla) penetrans (Chiggoe Flea}

(Page 93.)

l ig. i. Plantar stirface of human foot with chiggers in situ. Acte
size.

(a a) Group of eleven females.

( bbbb ) Isolated females.

(cc) Pits or cavities left after the removal of ti*
females.

(d) Section of epidermis with two females in si
(Lateral view.)

Fig. 2.

(а) Larva showing the squamose character of th-
epidermis. x 60.

(б) Outline of a younger larva. x 60.

(0 Antenna of larva with its accompanying group
blunt spines. x 250.

00 Mandibles of larva, x 250.

(') ? Lephalo-pharyngeal plate of larva. x 250.
(/) Buccal spines of larva, x 250.

(g) Empty cuticles of ova. x 60.

PLATE Vt.

Fix. I. Hitman Foot with “ Chiggers " in si/ii.

/>. S'eu'sfeml, ml mil. del.

Fix ■ -• Derinatophilns ( Sarcopsylla ) pant ran
Details of La nut, etc.

DESCRIPTION OF TWO NEW SPECIES
OF AFRICAN TICKS

H

”5

DESCRIPTION OF TWO NEW SPECIES
OF AFRICAN TICKS

By

G. NEUMANN

HROFliSSKl'K A l.’KCOLE NATIONAI.K VflTfiRINATKE L)K TOULOUSE

Rhipicephalus duttoni , n. sp. (Neumann)

Male.— Body, narrow in front, broadest (1*85 mm.) a little
posterior to the middle, length with rostrum 3-55 mm. Scutum
slightly convex, chestnut-brown without spots, abdomen does not
extend beyond its margins ; cervical grooves are very broad, shallow,
and form elongated depressions, they are not punctated and are
continued posteriorly by a narrow superficial groove which extends
beyond the middle point of the length ; marginal grooves broad,
shallow, slightly and finely punctated, commencing almost imme
d lately behind the eyes and terminating in the groove which separates

Fig. 22.— Rhipicephalus duttoni, i . Rostrum, x 45

the two last from, the following festoon; punctations irregular,
coarsest in iront, fine and superficial over the remainder of the
surface; behind are three wide, shallow, unpunctated longitudinal
grooves, the middle one being the longest; festoons longer than they
are broad, slightly punctated, normal. Eyes flat, yellowish, large,
marginal. Ventral surface reddish-brown, covered by rather long
and abundant whitish hairs. Anus anterior to the middle of the

length of the adanal shields ; ad anal shields have the shape of a scalene

triangle and so form a long internal posterior spine (the internal e<%e
is longest, it is rectilinear in its anterior half hut is concave behind
the external edge is slightly convex ; the posterior edge is concavr
and bordered by punctations) ; the outer shields are replaced bp I
prominent, non-chitinous fold ; no caudal prolongation but a chibnosJ
thickening on the median festoon Peritrcmcs narrow, whitish. comma
shaped with the point bent back towards the dorsal surface Rostm
o' 6 mm. long, dorsal base almost twice as broad (o*6 mm.) as long [
lateral angles at about the middle of the length, posterior angles quite
prominent. I lypostome very slightly spatulated, has six rows of teeth
Palps as broad as long, flattened dorsally ; second segment scared;
longer than the third and retracted into a blunt point dorsally at it-

J. Ventral surface of posterior extremity. .»

posterior border T one 1 ■ •

white hairs • ant • ^ le allvely strong. Coxae covered with long

nairs , anterior summit mnrh m _ j , . .u.

Ventral surface of posterior extremity. * #
strong. Coxae covered with long

Ti • *

ih!s species
Everett Dutton.

ii;

Rhipicephalus lougus, n. sp. (Neumann)

Male. — Body narrow in front, sides subrectilinear, length with
rostrum 4- 1 mm. ; broadest (2' 15 mm.) towards the posterior third
Scutum almost flat, dark chestnut-brown, without spots; along the
posterior festoons the abdomen extends beyond its margin ; cervical
grooves, very short and deep ; marginal grooves deep, narrow, each
occupied by a row of punctations, commencing a little behind the
eyes, ending at the posterior border of the penultimate festoon,
and are continued forward, and especially inwards, by a row of n.ar -<\
scattered punctations; punctations irregular, abundant in the spur*
enclosed between the marginal grooves and the punctated antemn
prolongation; they are regular in size and distribution anteriorly mi

Eli'.. 24. — Rhipicephalus lougus, 3 . Rostrum, x 4 li¬
the space corresponding to the female scutum, and they exist vuth
the same appearances in the posterior part of the scutum, but
they become scanty and finer in the neighbourhood of the marginal
grooves and their punctated anterior prolongations ; the marginal
border is smooth save in the scapular angles which are hollowed In
six to eight larger punctations; posterior festoons longer than thin
are wide, almost smooth, followed by short abdominal festoons Iiy> •
Hat, yellowish, of medium size, marginal with a coarse tangential pum
tation at the internal edge. Ventral surface reddish brown, with a
lew short hairs. Anus at about the middle of the length of the adan.il
shields ; adanal shields long, semilunar (the inner edge concave, tin
external and the posterior convex) with fairly heavy punctations on
their surface ; external shields replaced by a prominent non-chitinous
fold , festoons subrectangular, sharply defined ; no caudal prolong.!

1 1 8

tion. Peritr ernes whitish, broad, comma shaped, with point curve

towards the dorsal surface. Rostrum , length 07 mm., dorsal baa

more than twice as broad as it is long, lateral angles very promme:
at about the anterior third of the length, posterior angles qnr.t
prominent. Hypostome slightly spatulated, with six rows of tot
Palps hardly longer than they are broad, flattened dorsally with’
second segment a little longer than the third and shortened 11
blunt point posteriorly at its inner border. Legs relatively strcc.
Coxae with long scattered hairs ; first with anterior summit ;
prolonged nor visible*on the dorsal surface, has two very long spine-

short spines the ' °U- 1 postenor border incurved to form t«
and sharp. Tarsi f ^ ,.e'n^ NVlc*e’ fiat and blunt, the outer nam
caruncules medium size 111 ^ " two terminal successive spur?

Female unknown.

at Kasongo (April) * estabhshed from a male taken on a

Khipicephczlus duttonz and R 1
manner in the table of Hiff . . fon*"* are placed in the fol

nf Rhipice.phatus. eient,ation of males of the various :

TT9

(Eyes flat. — 2.

1 ' ( Eyes prominent.

(Marginal groove well marked. 3.

(Marginal groove absent.

(Dorsal scutum uniformly brown. 4.

1 Dorsal scutum black and white. K. pulchdlus.

, Adanal shields not prolonged to a point (posterior border
straight or convex).- 5.

* Adanal shields prolonged in one or two points (posterior
border concave). 12.

I Posterior border of the body not prolonged nor furnished with
a caudal prolongation. 6.

V Posterior border of the body furnished with three
prolongations. R. super tritus.

/Adanal shields triangular or subtriangular (internal edge
straight or slightly concave). — 7.

b- Adanal shields sickle-shaped (their inner border very concave,
the two others forming a single regular curve).-
- R. haeniaphysnloid.es.

(Marginal groove deep, long, commencing near the eyes. S.

7. \ Marginal groove superficial, short, commencing at the middle
• of the length of the body. R. ziemanni.

8.

, Dorsal scutum with scattered punctations. 9.

I Dorsal scutum with numerous, serried, punctations.

.Dorsal scutum with unequal, very evident, irregularly distri
buted punctations. — 10.

Q- Dorsal scutum with large, equal punctations arranged in
longitudinal lines with or without additional fine and
hardly visible punctations.- R. simus.

Coxae I, with or without a short prolongation in front,
not visible from the dorsal surface. — 11.

Coxae I, with a long anterior prolongation, visible from
the dorsal surface. -R. appendtculatiis.

120

Dorsal scutum with mixed punctation, regularly distributed
— R. sanguineus.

Dorsal scutum with medium-sized uniform punctations between
the marginal grooves, rare in their neighbourhood, none
on the edge, large ones on the scapular angles
R. longus.

(Anal shields with a single (or principal) internal point. 13.
I2' 1 Anal shields with an external point. R. lunulatus.

/Coxae I, without visible prolongations from the dorsal
J surface ; adanal shields with two points. R.armatus.

| Coxae I, with prolongations visible from the dorsal surface;
adanal shields with one point. R. duttoni.

ON SOME PARASITES IN THE MUSEUM
OF THE SCHOOL OF TROPICAL
MEDICINE, LIVERPOOL

ON

SOME PARASITES IN THE MUSEUM OF
THE SCHOOL OF TROPICAL MEDICINE,
LIVERPOOL

BV

Dr. A. LOOSS,

PROFESSOR 11K PARA.SITOI.OGY, SCHOOI. Oh MEDICINE. ■ \IK«*

WITH A CONTRIBUTION ON

A CASE OF DISTOM1 ASIS OF THE LIVER AND
THE RECTUM

BY

EDWARD CUFFEY,

PHYSICIAN TO TUI BRITISH HOSPITAI. PORI SAII>

During a visit i recently paid to the School of Tropical Medicine
in Liverpool Dr. .Stephens showed me a number of parasites which
had been sent to the Museum of the School from various places
1 nfortunately, some of the bottles bore no indication whatever as to
the locality from which the specimens were obtained ; others were
labelled, but an inspection of their contents with the naked eye or
pocket lens raised within me strong doubts as to the correctness of
the respective identifications. I o make things sure Dr. Stephens
kindly agreed that 1 should take with me. of those forms of which
several were present, a few specimens for a more careful examination
and possible identification of the species. The results of this invest i
gation are given in the following pages.

Fasciolopsis buski (Lank.), 1857.

About a dozen specimens from Hong-Kong in a jar labelled
Distoma crassum.” Rather large and fleshy, apparently sonu
what contracted worms of about 30 mm. in length, 13 to 16 inn. in
breadth and nearly 4 mm. in maximum thickness. Bodv mode rat el\
elongated, rather regularly oval, its surface marked with numerous fine
transverse ridges brought about by contraction. Both suckers vcr>
dose to the anterior end of the body ; opening of the anterior sucker

124

minute and completely ventral, opening of the posterior sucker very
wide, transversely oval, about 2 mm. in its largest diameter The
same is the average distance between the centres of the two sucken
In some cases the part of the body containing the anterior sucker was
found separated from the rest in the shape of a minute but quite
distinct cone resembling that which characterises the members of the
genus hasciola. Immediately in front of the ventral sucker there b
a transverse slit indicating the position of the genital aperture All
round the margin of the body, the colour of the worms is considerably
duller than in the middle and in most cases this duller tint
projects twice on each side in an angular fashion into the lighter
1 entral area, this latter thereby assuming the shape of three
successive but incompletely separated discs. All these charaders
discernible by naked eye inspection suggested Fasciolofsis bush
(Lank ). A comparison of the internal organisation so far as this
could be made out from cleared entire individuals did not reveal any
noticeable difference from the description recently given of the species
\\ Odhner (1902). The determination Distoma crassum is therefore
coirect, for the name Distoma crassum (COBB), 1877. is only a
synonym of Fasciolopsis buski (Lank). 1857.

poncrea/icum (Jau^,,.

A bottle from Hong-Kong. without label, contained a nu,

recall T"S,Zed Whidl '***

Asia o "i pancreat,c of ruminating animals in Ea:

that there \ °SC1 'nSpC< l‘on il cou^ be seen even with the naked

with very wide I„d I"”61163 Present in material, a larger !
which the <?, 1 projecting suckers and a somewhat smalle

'ess prominent"8 71 aIs°' hut '»»'■ »»""

completely mat ° ' vanet*es were represented by evide

isrjirr —

'lirri.lii,, r j ™ f>f ,h‘\ae™'1 Uan"r Il'"

and four adults of «-i ' 1 wo sPec|mens of the first var

f°™ the basis If 11 1 Sma" kind>y given to me by Dr. STEPH
After dear in PreSCnt descriPtio»-
t airly well be made P°’nls °f tbeir internal organisation cc

were in full ap-reem T* ent're sPecil»ens ; the details obser
greement with the descriptions given of the Distc

pancreaticum by the earlier writers. There were, however, between
the two varieties in addition to the differences mentioned above in the
size of the suckers several other slight internal discrepancies which
leave no doubt that the two forms must be considered as two distinct
species. Both are closely allied to each other and members of the
same natural genus. The generic name hitherto usuallv adopted f< ir
Distoma pancreaticum is Dicrocoelium ; indeed, the principal features
of the anatomical structure are the same in Distoma pancreaticum
and Dicrocoelium lanceatum (St. and Hass.), the type species of the
genus. But besides this similarity in the main structure there are
also certain differences which to my mind forbid placing Distoma
pancreaticum in the genus Dicrocoelium itself , that is to say as it is
represented by its type lanceatum. One of the most conspicuous of
these differences lies in the thickness and the considerable
breadth of the body which latter allows the testicles to take a
distinctly lateral position at about the same level, whereas in
Dicrocoelium sensu strictiori, owing to the narrow spindle-shaped
outline of the body, the testicles are placed obliquely one behind the
other. In Lyperosomum which is also closely related to Dicrocodium
the two cross diameters are still more reduced, the members of this
genus presenting to the observer an almost filiform body of about
equal breadth and thickness in which the genital glands are found in
a straight line one behind the other. In addition to the peculiar
shape of their body the two Distoma pancreaticum differ from
Dicrocoelium by the more complicated structure of their excretorj
vesicle and the stronger development of their copulatory organs On
llie whole, therefore, there is sufficient reason for creating a new genus
for the two forms under discussion.

I he question as to the correct denomination of the species was
somewhat difficult to decide. Distoma pancreaticum was discovered
in Japan, and became first known in Europe during the Paris Exhibi
tion in 1 889, where the Agricultural School of Komaba exhibited a series
of parasites among which there was a “ Distoma pancreaticum from
the pancreatic duct of sheep,” and a “ Distoma pancreaticum var
from the pancreas of sheep.” The new parasite was referred to in
subsequent years by RAILLIET (1890) and JANSON (1893 and 1895)
the respective papers deal with the anatomical structure and the
pathological significance ol the parasite, they emphasise its

126

resemblance to Dicrocoelium lanceatum but do not so far at least
as I can personally consult them at present— contain any decisive
statement from which one or the other of the two Hong-Kong species
mentioned above might be recognised. In 1897. GlARD and BlLLE"
apparently ignorant of the existence of the Di stoma pancreatine
described a Distoma coelomat icum which the latter author had found
plentifully in the pancreas of an ox killed at the slaughter -house of
Cao-Ban g (1 onkin) ; there had at first been some mistake as to the
habitat ol the parasite which led to the denomination coelomaticuw
but this mistake was corrected afterwards upon a suggestion of
Railliet. The details given in GlARD and BILLET'S description
are sufficient to show that Distoma coelom at icum, also structural!)
resembles Dicrocoelium Invocation and Distoma paucreaticum , but
theie is again no statement which would be indicative of one of our

two species in contradistinction from the other.

The latest description of Distoma pane real icum is given t
AILLIE1 and Marotel in 1898. The specimens upon which the
I apei is based were collected by Dr. GOMY, a French veterinai
smgeon, from the pancreatic ducts of cattle and Indo-Chines
- U r.a °jS *n ^a’£oon (Cochin-China), a locality therefore which is nc
S° a' lslant from the place at which GlARD and BILLET’S materi;
a<en. This time, the description is more complete; the worm

h^toreadth SU^C froni 7 to 10 mm. in length and from 4 to 4-5 mu
. ... . ’ ,Clr suc^ers are °f about equal size, the posterior 0

anterior wWch" ‘p^esentf TT ^ S'ight'y ^ tb'

between the two suckers raricT " ‘° °'9 The di5tanC'

amounts to abom - roni x*3 to 2*4 mm., that is to say, r

be more fully seen her^fr' i!'* Iength °f the body- As wil
to the smaller of th^ ^ * theSC nieasurenients apply very well
antomical details IIo”ff'Konff species and among the

would not quite well ^ ^ .°rs tbere ‘s none moreover which

Perfectly safe to assume h!” ^ Nation. 1 thus think it

same as the Distoma y, ^ >S. snialIer Hong-Kong species is the

Marotel in 1898 fron, *nfrea/*cum described by RAILLIET and
paper (p. J2) aIso states . ,Cattle' . But Railliet in the same
specific identity Qf thk r ^ satisfied himself dc visu of the
parasite found in Tonki, ' ““‘"'Chinese pancreatic fluke with the
k,n by B,LI-et- This latter parasite had first

127

been described under the name of Distoma coelomaticum and tin-
question now arises: is this Distoma coelomaticum of Indo-Chma
really the same form as the Distoma pancreaticum of Japan? It tin-.
were so, then the smaller IIong-Kong form would have to be identified
as the genuine Distoma pancreaticum, the name coelomaticum would
have to be definitely dropped as a synonym and a new specific name
would have to be given to the larger IIong-Kong species with tin*
strongly developed suckers.

As I have already pointed out, the data given in the printed
description of the original Japanese pancreatic fluke are not sufficient
to decide the question. I possess, however, in my collection a number
of specimens of Distoma pancreaticum which I owe to the kindncs-
of Professor JANSON, of Tokio. They were collected from the
pancreatic ducts of cattle in Japan and I think there can not be an)
reasonable objection to the assumption that they are specifically tin
original Japanese Distoma pancreaticum. It is true that the fir -t
specimens of this species exhibited in Paris were labelled as collected
from sheep, but 1 am not inclined to attribute a great importance to
this difference in the host, because it is a well-known fact that many
parasites of cattle and buffaloes occur not only in sheep but in otliei
ruminating animals also. A comparison of this Japanese material lias
now shown, first, that there is only one species present in it and.
second, that this species is identical with the larger species of the
Hong-Kong material. On the ground of these facts I feel justified
in considering this latter species as the genuine Distoma pancreaticum
of JANSON ; for the smaller species I readopt the name coelomaticum of
CSiard and Billet, the suppression of which was, in the sense of the
nomenclatural rules, a mistake based upon an erroneous identification
of the species.

Eurytretna, nov. gen., Dicrocoeliidarum,

Differs from Dicrocoeliuni especially by the following particular-
body rather thick and considerably broadened, with the exception of
the hindmost part which retains its original shape and appears as ,1
small triangular appendage sharply set off from the rest of the body .
it is repeatedly referred to in the descriptions of the earlier authors a-
resembling in outline the cephalic part of Fasciola hepatica. Suckers
very large and prominent, the oral sucker completely ventral and

128

surmounted by the anterior margin of the body Excretory vesicle
composed of a median stem which divides into two at about the
middle of the total length, the transverse branches, in the neighbour¬
hood of the intestinal caeca, dividing again each into an anterior and
a posterior branch which run for the greater part outside of the
intestinal caeca, and terminate in the neighbourhood of the antenor
and posterior extremities of the body Testicles lateral, at about the
same level but far apart from one another Copulatory organs well
developed ; cirrus pouch thick, almost cylindrical, containing a long
but rather thin seminal vesicle and an equally long and thin ejacu
latory duct which both describe a number of coils within the pouch.
Vagina corresponds in length and arrangement to the ejaculatory
duct. Structure of the remaining organs as in Dierocotlium
Type: Eurytrema pancreaticuw (J ANSON) 1889.

b.urytrema pancreaticuw (JANSON),

(nec Dicrocoelium pancreaticuw , RaILLIET and MAROTEL, 1898;

I he two I long-Kong specimens at my disposal are fairly w<

extended and measure 13 and 14111m. in length; their maximu

breadth ol (1-5 and 7 mm. is reached immediately behind the ventr

sucker. 1 hence the body narrows more quickly towards the anterii

extremity than posteriorly but ends rather tnincatedly owing to tl

large size of the oral sucker above which the margin of the body pn

I cts in t ie shape ol a thick lip. I he small tongue-like appendage of tl

I lor end is very conspicuous. The thickness amounts to almo:

- mm m t ie median line of the anterior half, but gradually decrease
towards the erlo-p i 6 J

mntr a , 1 ne JaPanese specimens are somewhat moi

16 mm" 0 K1 t.lncker throughout, varying in length between 9-5 an

lo6“™' " breadth bet— 5-5 and *S n,n, -in Tuch a way that th

however aUnT^ ^ generaI1>' ,ess broad and vice versa. There art

but onlv'.cm^™^ matUre individuals 9-5 and 10 mm. long

lari Id br°ad' SWn' without armature. Suckers ver

r? - ** - -* «— •

of I-45 to 1-65 mm n 1 ie ventral which presents a diamete

everted reaching therebv ^ S°me specimens found more or les:
specimens mention^ u & °f In the two smallesi

"3».u °f the 5UCk- - - ‘--4-

y- I he} are, therefore, even in these

<

129

smallest specimens considerably larger than in the Dis.'owa
maticum of the same size. The ventral sucker is always found near
the end of the anterior half of the body; the distance between tin-
centres of the two organs varying in the larger specimens from p t.
57 mm. and being in the two small specimens 3*9 and 3-3 nun, that
is to say it amounts to at least a third and is usually but little less tli m
half of the total length. The genital opening has its position about
halfway between the two suckers and is sometimes found slight l\
raised above the level of its surrounding. In one of the specimen'. tin
penis is seen everted, apparently to its full length, for the larger part
of the seminal vesicle is located in it ; it represents an organ ; • j m ,
in length and at its base 0*37, at the free end 0-24 mm widt It
surface is smooth.

The internal anatomy agrees with the descriptions given In thr¬
eadier writers. The small pharynx of C47 mm. length and 1 mi
width is usually so much displaced dorsally that it is not visible fn
the ventral aspect; behind it there follows a short oesophagus v,1 1 1
as a rule, runs straight down towards the ventral side and may tlm-
more or less entirely disappear from view. The intestinal cae< a of
proportionally insignificant calibre quickly diverge and follow , , v g
course which becomes especially pronounced behind the testicles
Their blind terminations lie about as far as the length of the tongue
like end-portion of the body in front of the base of that portion and
usually not quite on the same level, the left branch being ns a rule
(but not always) the shorter. The full extent of the excretory v- ' •
could not be made out in any of the specimens at my disposal. -uving
to the thickness of the body and the strong development of the uterine
coils. The main parts described in the generic diagnosis are usualh
seen and followed without much difficulty; but, in addition to them,
one may observe in some places isolated portions of similar tu’<

\\ no seem to be ramifications of the anterior and post eric- r lat--r.il
ranc es. There are two running forwards on either side of the
ntra sucker, and two running backwards and towards the m-dun
me amidst the uterine coils in the posterior half of the h dv
t lough they appear as if they were parts of the excretory w I-
cannot tell with certainty whether or not they really belong to b ‘
organ. The terminations of the main anterior branches of th«- v-
ie c ose to the oral sucker at about the level of its centre, those of the

r

1

posterior branches at, or slightly in front of, the two notches which
separate the terminal triangular tongue from the rest of the body.

The position of the genital aperture has been given above; it lies
always a little behind the bifurcation of the intestine The thick
muscular cirrus pouch, usually somewhat curved in the shape of a
sausage, presents a length of 2*2 to 3 mm. and an average width of
o-6 mm. Its posterior end rests upon the anterior slope of the ventnl
sucker, but there are some variations in its length in connection with
the contraction of the body and the fulness of the seminal vesicle
This latter occupies the posterior half of the pouch, describing, within
it, a proportionally large number of small coils. The pars prostatica
is short and the number of glandular cells surrounding it limited
The ejaculatory duct is at first rather narrow, but widens somewhat
towards its free end ; its coils are proportionally numerous also. The
dimensions of the evaginated penis have been given above. The
air) laige testicles are normally found at the same level somewhat
1 nr the \entral sucker, but may in strongly contracted specimens
more or less completely shifted to its sides. They' have in the

g er number of my specimens a distinctly lobed shape, but

im?P T3 y t,1C IObeS may be SO sbort and l)road ‘hat the glands

n Z,OPOSSeSS U mereIy notched outline. The number of lobes (or

for the nth^t ^ f COnstantIy four for one (usually the left) and live

‘ sexuL ,mn, f " ^ COnditi°- are inverted in the cases of

well as in tlW V/'1 • W*llch are not infrequently met with in this as
35 ,n the following species.

testicles, on the^eft^ ' S USUal'y found a short distance behind the
of the body jt j 'n C<)ses sexual amphitypia on the right — side
distinctly lobed ok ^ T ^ tbe sPecimens with lobed testicles a

those specimens, ondL* Th ^7 munber of ,obes bein£ tl,ree ln
notched outline onl h ° ^ ,t,nd’ ,n wb,cb the testicles show the
less and appears coi^' * & °'ary’ to°’ ^oses its lobed outline more or
The oviduct starts from^' S°n.le irre^u^ar bulgings of its surface.

into a well-developed shentSinCnr S'^e and a^most immediately enters

an opaque sharply out WrM ^ 'V,llcb ln cleared specimens appears as
'he ventral surface Th ° y y tbe Slde °f the ovary and close to

average diameter of ^ ^ 3 fair,y ,ong LAURER’s canal of an

ovary. It carries a small m '^1'rb °Pens to the outside in front of the
I emmal recePtacle which I have never been able

to discover in entire specimens, but clearly saw in a series of sections
The receptacle represents in the sectioned specimen a globular bmi\
of about the same size as the ovary (0*4 mm. in diam.) and lying close
to its dorsal surface. It communicates with the LAURKR S canal by a
distinct thin duct, but does not contain many spermatozoa, a fact
which renders the seminal receptacle very transparent in unstained
specimens. This insufficient filling in combination with the p •rticiilar
position of the seminal receptacle is obviously the reason why it
escapes observation in entire worms.

The yolk-glands are only little developed; they are composed 1
rather numerous acini of a slender club shape collected into in<<n- r
less distinct groups, the number of which seems to van- between ten
and twelve. In extended specimens the groups lie in a nearly straight
line one behind the other and are fairly well discernible ; in contrai ><d
individuals they become more closely packed together and s<vin- >'{
them — especially those in the middle — may be shifted lateral!', in sm b
a way that for a certain distance there appear to be two irregularly
alternating rows of groups. In this latter case some of the follicle',
are pushed below the intestinal caeca, otherwise, the yolk -gland'; le¬
as a whole between these and the body margin, usually the gland on
the side of the ovary somewhat farther in front than that of t hr
opposite side. Their extent varies a little individually and ' '
the contraction of the body. In the greater number of specimens thrv
commence somewhere at the outer border of the testicles and thru-
posterior ends lie halfway between the posterior border of the ventr d
sucker and the base of the tongue-like end-portion of the body, but
may in strongly contracted specimens also be found further lop.'. a d
The transverse yolk-ducts originating, on each side, by the union of
an anterior and a posterior tributary arise from the glands r
about their middle; they join near the ovary (the left transverse yolk
duct being thus shorter than the right) and then enter the con plrv
of the shell-gland from behind. There is no distinct w idening of tin
unpaired part of the yolk-duct into a yolk reservoir The coils oi tin-
uterus are very numerous but do not, as a rule, appear to the ob-erver
m the shape of regular loops, but rather as loose tubes branching and
sometimes anastomising in an irregular manner. Occasionally only,
i.e. in individuals in a certain state of contraction, one may pern - 1
the picture which is so characteristic of the uterus of Diet (oclvr

132

lanceatum and in which the descending and ascending steins of the [
uterus cover each other in such a way as to appear as a straight maic
stem from which lateral ramifications are branching off As far as
the branches of the intestine extend backwards, it is only exceptional
to see one or the other uterine coil cross them towards the tnarp:
of the body ; behind their blind ends the coils extend much nearer to
the margin and the terminal small triangular appendage is. in perfectly
mature specimens, almost completely filled with them. Anteriorly the I
coils are limited by the testicles and the ventral sucker; the utens
then passes between the latter and the right testicle (i.e. the testiefc I
opposite the side of the ovary) and in front of this still forms sow I
small convolutions before entering into the long and thin, but rathe I
muscular vagina which accompanies the cirrus pouch while desorbing (
a number of narrow coils.

I he eggs have a striking likeness to those of Dicrocoeliu*

lanceatum ; they are slightly larger than these showing, on an

average, a length of 0-05 mm. and a width of 0*034 mm.- but apart I

from that, they present the same oval shape and the same thick shell

of deep brown colour as these and contain, in addition, a miracidium

" when fully developed, shows the same two enigmatical strongly ,

reraclive bodies which are peculiar to miracidia of Dicrocoeli**
lanceatum.

Eurytrema coeloniaticum (Giard and BILLET) 1892.

(Synonym: Dicrocoelium pancreaticum Railliet and MaROT:

1898).

anH?em^tm5breldtehSoftt7 diSP°Sa' “ 'enB'h °f '0'”

measure 7-15 by 55 mm* : younger (immature) specim*

almost regularly ' r \ ^ ^ mm » &c- The outline of the body

anterior extremit ^ ^ l*iere ’s 110 marked narrowing towards t

triangulaire ” Gf £» tr’angfular terminal portion (“ langue

thickness of the bod* ^ *S ^,llr^ conspicuous. The rnaximt
large (matured snp • ^ oes not seem to exceed 1 mm. In the k
Present diameters Wh,ch 1 have for examination the oral suck*
diameters of 0-q8 ° ° ^ anr* °’83 mm., the ventral suck<

91. 085 and 0^83 mm. respectively. T he

133

measurements agree fairly well with those given by RAILLIET who,
in the Saigoon specimens, found the oral sucker to vary in diameter
from 075 to 0'93 mm., the ventral sucker from 07 to 0*9 nun. The
distance between the centres of the two organs which RAILLIET give
as i‘3 to 2‘4mm. is in the four above mentioned specimens, 27, 2-97,
27 and 272 mm. ; the suckers are therefore, in Eurytrcma coelo-
maticum, proportionately much nearer to each other than
they are in E. pancreaticum , for the ventral sucker lies in the former
species with the greater part of its mass still in the anterior third of
the body and the distance between the two always remains than
a third of the total length. The size and relative position of the
suckers represent, apart from the general shape of the body, the mu-u.
conspicuous of the specific characters of Eurylrema coelomaticum.

The internal anatomy has been so fully described by RaILLIEI
and MAROTEL that I have nothing of importance to add ; it is more
over essentially the same as that of E. pancreaticum , as may be seen
by a comparison of the data given above with the description of the
French authors. There are nevertheless the following differences 111
detail which 1 am inclined to consider as of specific value, although
some of them are not absolutely constant. In E. coelomaticum, the
genital glands never present the lobed appearance of those ol
E. pancreaticum , but always show a compact shape with irregular
bulgings of their surface instead of lobes; the apparent rule that one
testicle has four the other five bulgings seems however to be preserved
here too. The yolk-glands which, in E. pancreaticum, were composed
of ten to twelve groups of follicles, show in E. coelomaticum , only >i.\
to eight; the glands are therefore as a whole shorter than in the
preceding species and they also begin somewhat farther backward-,
viz. at the same level with the posterior margin of the testicles In
spite of that, their posterior terminations appear much more distant
from the end of the body than in E. pancreaticum, a fact due to t In¬
considerable difference in the relative position of ventral sucker and
genital glands in the two species. In E. coelomaticum, finally, it . -
not so uncommon to see one or the other uterine coil pass beyond the
intestinal branches laterally.

I he eggs, in shape, aspect and structure of the miracidium, closely
resemble those of E. pancreaticum , but are somewhat smaller, namely

*34

0 042 to 0-046 mm. long and 0*023 tc> 0027 mm. wide. This size, by
the way, is the same as that of the eggs of Dicroc . lanceatum .

Gastrodiscus secundus n. sp.

A small jar containing about a dozen specimens and a label on
which there was written in pencil : “Gastrodiscus Sonsmonis Cobbifrom
mule. ' They were collected by GILES in Assam, who states that they
are also found in equines throughout India. In their external appear¬
ance they absolutely resembled the African Gastrodiscus aeg) ptiacui
(Cobbold) 1877 (= Dipl os tom a aigypriacum Cobbold 1877,
- Gastrodiscus polymasios LEUCKAKT 187;. ■ Gastrodism

sonsinoi Cobuold 1877), with the sole difference that all of them

were noticeably smaller . Three specimens at present at my disposal
si iow a length of 7, 8 and 8 mm., and a maximum breadth of 4-5, 5 and
5 nun. ; the corresponding measurements in Gastrodiscus acgyptiacus
being 10 to 13 nun, and 7 to 9 mm.

I he body shows the characteristic division into two parts: the
c) indrical 01 slightly dejjressed cephalic portion, and the broad oval
c isc, at the end of which the second sucker is placed. In the
specimens under discussion the former is about 1*35 mm. long and at
ise 1 6 mm bi oad. Near its top, there appears on the ventral
, C moudl °Pemng, about 0 37 nun. wide. In its neighbour-

... , 16 CUtjc^e shows a large number of small circular or oval

Tliev a 'V ProJect externally in the shape of small tubercles
mouth n 3pparentIy tactile papillae similar to those found round the
of the ce27 m a,lied f0rniS‘ Beh»ld mouth the ventral surface
abdominal dis/iT^n excavated along the median line. The

rolled inwards to * PParently niuch contracted, for its free edge is
terminal disc are vi ^1 30 extent dlat the ventral surface and the

it lies open to the b °ldy‘ * he ventral surface is, so far as

‘‘papillae,’’ or “ pseud DSC1Ver’ densely covered with tlie well-known
like openings. Some ^Ucgers Scaring on their summits small slit-
represent small knobs ° BleSC 01£ans are everted and, in this state,
hours. On inspectino^t?601^8’ somew^iat over their retracted neigh-
circular hole surrounded m W°rms Wlth a magnifying glass, a small
ln dle midventral line 1 ^ & S°lnevv^iat raised brim becomes visible
llle cephalic cone. t? °Ut 1 ^ to 1 "5 mm. distant from the base of

"as present at the same place in all the

135

specimens and therefore a normal structure, probably the genital
aperture. If the latter was true then the worms could not belong to
the species aegypliacus, for in this, the genital aperture is placed
immediately behind the cephalic cone, at the very base of the
projecting edge of the disc.

A close microscopical examination of the three specimens kindly
given to me by Dr. STEPHENS has confirmed this presumption ; the
parasites represent a new and at the same time a true member of the
genus Gastrodiscus. It appears superfluous to give a description ot
its internal structure here, for this would mean nothing but a second
description of the structure of Gastrodiscus aegypliacus. It may
therefore suffice to point out the differences between the two species
They are chiefly given in the position of the genital aperture and the
extension of the yolk-glands. The former has already been men
tioned ; it is a circular or transversely oval opening about 03 nun
wide, and leads into a genital atrium of 03 mm. in cross diameter, the
muscular wall of which is of considerable thickness and much more
conspicuous than in the African species. 1 he floor of the atrium is
again raised in the shape of a conical papilla on the top of which the
male and female genital ducts open close by one another in minute
pores. The yolk-glands show a richer development than in Gastrod.
aegypliacus. In this species, it will be remembered, they are located
in the ventral half of the body, between the intestinal caeca and the
free margin of the disc, and it is rather exceptional to find some
folliculi extending below the caeca into the space between these and
the median line of the body. In Gaslrodiscus secundus, the yolk-gland-
trespass on the intestinal caeca in such a manner that their loosely
grouped follicles occupy, in the ventral half of the body, all the space
left free by the other organs. A minor difference is afforded by the
size of the suckers. The oral sucker which, in Gastrod. acgyptiacu >,
presents a cross diameter of o-8 mm shows, in G. secundus , a diameter
varying, according to the size of the specimens, from 035 to 0‘65 mm. ;
the posterior sucker, in G . aegypliacus about 2 mm. wide, in
G. secundus scarcely reaches 1*2 mm. in size. The uterus, in the body
of the new species, principally takes the same course between the
organs as in G. aegypliacus , but, although it is, in the two larger
individuals at my disposal, thickly filled with ova, it seems that it does
not form so many secondary loops as in the older species. The ova

136

which, in the latter, show a length of 017 to eng mm. and a width c: |
O'liram., are, in G. secundus, somewhat smaller namely 0*1 5 to
01 6 mm. long and 0 09 to O'l mm. wide.

Opisthorchis sinensis (COBB.) 1875.

Several worms of a rusty yellow colour contained in a jar of
unknown origin and labelled “ Distoma stnenst COBBOLD " struck me
at once by the fact that they were considerably larger and stouter
than any others I had seen of the species up to that time. I therefore
had no hesitation in expressing the opinion that these worms could
not be the true Opisthorchis sinensis. Since the number of specimen;
was limited, I took two specimens only with me, an unhurt one and
another which was broken into two pieces. About a fortnight after
<ny visit to Liverpool I paid a visit to the “Institut fur Schiffs-und
ropenkrankheiten ” in Hamburg. Dr. hULLEBORN very kindly
showed me round and, during an inspection of the collection of human
parasites, 1 was rather surprised to recognise under the designation
. Plsth°rchis sinensis the same large form which I had seen a short
c pre\ iously in Liverpool. 1 here were about a dozen individuals,
t ° -r!ein *n sP’r*t> others mounted as microscopical prepara
1 jSj • . *1<1C! ^een co^ected from one case, a Chinaman who

ia ie in the Seamen’s Hospital in Hamburg. Besides these large
^ • 1S’ 1 lere was one niicrosopical preparation of an ordinary small

not n Tc 3S ^lose * ^iac^ a^°ne seen before. This specimen was
obtain d n case from which the large worms had been

material to rJ c ULLEBORN was kind enough to confide this whole
return to E^nt^0™1^15011 Wlth the LiverPool specimens. On my
viously a preai ’ °iUr Weeks later> 1 learned that a short time pre-
mortem examirw m Cr °f ^ver flul<es ” had been found at the post-
Hospital of Port a ^ ll’naman who had died in the British

a case of Opisthorchiosis I wrote
Port Said to ask f U^'C ^®cer °f the Town and Port of

Dr- cw«v, wh0 z °f.the worms. Thereupon,

the post-mortem e ' • ^ ^ Pat*ent under his care and had made
all the preserved m°St °bli&ingly placed at my disposa1

write a short tlle case- P*e was also kind enough to

abstract of it which x insert m full |,e«

137

because of some interesting particulars it contains.* The preserved
material sent to me consisted of about 200 loose worms, and many
others were still seen densely packed in the bile-ducts of some
preserved pieces of liver. Among the loose worms there was again
not a single specimen of the ordinary small variety, but all were the
large form. I had thus, within a very short lapse of time, thrice come
across a variety of the Chinese liver-fluke which I had never seen
before, and in all these three cases this variety had been present
alone, quite unmixed with individuals of the usual smaller variety
I had not the least doubt that this new form would prove to
be an independent species and that it would be easy to delimit it
anatomically from its smaller congener. I was therefore soniewhat
disappointed, when after a most minute and thorough comparison of
the specimens with those 1 possessed of the small “ Disloma sincusi,

1 could not, apart from the size of the body and the ratio of tin
suckers, find any constant internal differential character between them.
Even of the characters mentioned the last-named is by no means
conspicuous, and I had almost made up my mind that the two varieties
must be considered as one and the same species, when, on
reconsidering all the factors involved, 1 came to the conclusion that

* A CASE OF DISTOMIASIS OF THE LIVER AND THE DUODEN UM.

Too-ting, a Chinaman, aged .25, was admitted to the British Hospital, Fort
Said, on May ioth, 1906, suffering from Lumbar Abscess. He was landed from 1
ship proceeding from China to England. The abscess was pointing in the lumbai
region, posteriorly, 2^ inches from both the middle line of the back and the cre>t
of the ileum. It was opened and about 300 grins, of bile-stained pus were
evacuated. The walls of the abscess were explored and the transverse proce.-s of
one of the lumbar vertebrae was found to be necrosed. But no connection could
be found between the abscess and the liver. The cavity was washed out and
stitched up. Subsequently a drainage tube had to be inserted, for the pus collected
again ; but the man’s condition rapidly improved, and the wound over the site of
the abscess healed completely.

On June 21st, the man suddenly developed pneumothorax, affecting the lower
part of the right side of the chest. As he was thought to be suffering from an
abscess of the liver, this organ was explored again, but no pus was found. Oedema
occurred over the lower part of the right side of the chest and still no pus could be
found on exploration. The patient rapidly became worse, and died on June 25th.

I ost Mortem. 1 he abscess in the lumbar region had healed completely. V
reason for it could be found beyond the necrosed vertebra.

Liver : 1 he liver was enlarged and contained numerous abscesses in both lobes
I hey varied in size from that of a small pea to a tangerine orange. On sect.' :,
the liver presented hundreds of liver flukes in the bile-ducts. The hepatic duct-,
common bile duct, and gall-bladder all contained numbers of these flukes.

1 hey were also found free in the duodenum, but not in the ileum.

1 here were no flukes in the abscesses.

No cause for the pneumothorax could be found, and no connection between it
and the liver was apparent. All the other organs were normal.

( Signed) F.dward Cvfkey,

Medical Olficer, British Hospital,

Port Said.

such a ruling would be less defensible than the other alternative, viz
to declare them as separate species.

I am pretty certain that many of my fellow-helminthologists will
think I am pushing things too far, and that what I am about to do
here is nothing but “species-manufacturing" In order to meet this
reproach as far as I can, I may be permitted to give my reason;
somewhat in detail.

To begin with, it is a fact acknowledged by all observers who have
dealt with this particular group of parasites (BRAUN, LUEHE
Muehling, LOOSS) that the members of the genus O pisthorchis w
in many instances by no means easily to be distinguished. As one
out of several examples I will mention the case of Opisthorckis
/ el incus and O pisthorchis geminus , a case observed by myself. The
former species is in Europe a rather common parasite in the liver of
certain beasts ol prey, and especially in cats, and is in eastern Europe
even a not uncommon occasional parasite of man. 0 pisthorchis gemim
inhabits in Egypt the liver of certain birds ( Milvus aegyptius, the
common Egyptian kite, Circus aeruginosus, a glede, and Ante
bonhas fera , the wild duck). It resembles 0 pisthorchis felinW
anatomically to such an extent that the two forms are hardly
distinguishable from one another without knowledge of host and
oca lty. Their internal organisation, therefore, would not furnish

jectionable reasons for separating them specifically; nevertheless,
y must be considered as specifically different on biological grounds

imnn i i VC mUSt ac*n”t as true certain circumstances which are almost
.possible to comprehend. Among the hosts of 0 pisthorchis grndm

and thus f* P/lus >s a true Egyptian who never leaves the country
.hcoILm Tqmre hiS paiasites '« B»‘ if * does, it would b

actually Sha "" u d°gS’ C3tS *“* even Man— ll0sts that in EgJP1
also tV":;; °therf SPGCleS °f Parasites — should not p,ck then, «

the ^Ptian WrdranTthaTonheE EUr°Pe' *f **“

°ne and the same - ot * le European mammals were indeed

any trace of an OpiThorchi, however' have I in Egypt found

domestic; nor in Man - 1 °rm 111 can^ne or feline animals, wilder
ever there was an occ' ' 10U^' * *lave purposely looked for them when-
within the last ten ve^0!! ’ n°r haVC the Professors of pathology who
Kasr-el-Aini Hosnitnl S av<~ mac*e t,le post-mortem examinations at

° 1CGd anything of the sort. It can therefore

139

be considered as certain that no Opisthorchis species occurs in

mammals and Man in Egypt and the inference therefrom must 1 think

be that the Avian Opisthorchis which is indigenous in this country
cannot be the same species which inhabits mammals and Man in
Europe — all the resemblance they show in their internal organisation
notwithstanding.

The case of Opisthorchis felineus and Opisthorchis gem mu s is
not the only one known ; but this as well as the corresponding
instances tend to show that, in the case of the two Opisthorchis
sinensis- forms also, the internal similarity must not necessanl) be a
proof of their specific identity — the less so, as in the size of their
bodies and the ratio of their suckers characters are given which
practically allow a differentiation of the two forms.

A second point which seems to me very important is the iact that
the two varieties of Opisthorchis sinensis not unfrequently occur quite
separately. It has already been pointed out that the large vanct)
was present alone in the three cases mentioned above ; but the same
fact may be inferred from some earlier cases on record. The parasite
was discovered in 1874 by McCoNNELL; the measurements given of
the worms by the author are x7tj in. for the length and ) in. for the
breadth, this would correspond to 17-8 and 3-03111111. respectively
and is, as may be mentioned here in anticipation, exactly the size of
the specimens 1 at present possess of the large variety. The author

*It is perhaps not out of place here to draw attention to the fact that the sire
of preserved specimens of parasitic worms is within certain limits influenced b) the
metnods of preservation on the one hand, and by their own condition at the moment
of preservation on the other. If specimens are collected from a living or recent 1>
killed host, they are still alive ; if they remain in the organs after the death of their
host they will die, too, after a shorter or longer time; the same thing happens when
living parasites are brought into water, which is very harmful to them and destroys
certain species very quickly. Before death they stretch themselves to their full
size, and the commencing decomposition even makes them swell somew hat. Living
parasites, when preserved immediately after removal from their natural habitat,
always react by a contraction, which is the stronger the slower the preserving
leagent acts upon them. Specimens which have died — in water or in the organs
°‘ tlie*r hosts (as is usually the case in post-mortem examinations made some time
alter death and in warm climates) — do no longer contract, but may shrink more or
ess according to the dehydrating power of the preserving fluid. All these factors,
eretore, have their influence upon the dimensions the specimens exhibit alter
preservation, and it is necessary to remember this fact when the size of a 1 ertain
Pecies IS. used as one of its distinctive characters. However, the changes due to
preservation always keep within certain narrow limits, and can even be fixed b\
ac ual observation. The parasites of the Port Said case were obviouslv dead when
tney came into contact with the preserving fluid, for they are not contracted. 1 h.
nuid used had been formaline, which does not dehydrate: thus, the snecimen* «h,.u

1 - » - UrtW icauicu 00

shrunk to a length of 14 to 16 mm., but could be made to
termer length again by placing them hack into formaline.

adds that the breadth is very constant in all the specimens, wher®
the length presents slight variations from in. to in. (=abra
1 5*3 to 20 mm.). In 1877, the parasite was recorded by MACGREGOi
as the possible cause of a peculiar form of paralytic disease, of wind
he had seen three fatal out of eight cases. The observations werenui
in Port Louis, Mauritius, but all the patients were again Chinaae:
I have at present no access to MACGREGOR'S original article, but tin
parasites found and described by him were subsequently examined by
COBBOLD and recognised as identical with those of McCoNKELL
In 1878 McConnell himself describes a second case observed ;
him in Calcutta ; “ the subject was, curiously enough, again a China
man, and the distomata found " were quite identical with those
originally found.” Comparing these earliest observations we see that
m all cases the bearers had been Chinamen, and that the parasite
must have been of about the same size for no allusion is made to the
existence among them of any noticeably smaller individuals.

hi 1883, die presence of flukes in the liver of Man was reports:
from Japan, first in a paper written in Japanese (“ Observe
Hons on Distomes in the Lung and Liver ") by KlYONO, NAKAHAHA
Suc.a and Yamagata, acting physicians in the hospital of Okayama
and a short time later by Baelz in a paper “ Ueber einige neue
arasiten des Menschen. I nfortunately, 1 have at present noacces
is paper which is one of great importance for the question la0
low iscussing , thus I can here refer to its contents so far only#
cy are repioduced by later writers. According to these, BaELZ
recognises two dist.net species of liver-flukes, a Distoma hifi»

TuZnCUmJWe pernici0 s** , and a Distoma hepatis innocu**-
d. istinguished from one another (see LEUCKART,

attaining^ X* f°remost b>^ their quite different size, Dist. innocitu *
exceerl s «.„engt 1 UP tC> 20 ram-» whereas Dist. endemicum does not
mni. Of the other differential characters given by

««on,*lhehaf°l[°r”g seem t0 me.to be wortl' mentioning:
slightly 1-irn- ^ ^ ‘ lCrus of bghter colour but larger volume, (-)

-d (3) possesses **

The exact IL body-parenchyma numerous black granule

innocuum °f the OVa are’ according to BAELZ, in D*

in Dist. endemic u ° °3<3 mm- len£th by o-oiS to 0‘002 mm. bread*
°'°2 t0 °'°3 mm. length by 0-015 to 0*01/ **

breadth. From the names given to the species it may be inferred
that the small variety is very common in the country ("endemic'")
and harmful to its host (“pernicious”), presumably because of its
occurrence in large numbers, whilst the larger is rarer, i.e. not present
in large numbers and therefore comparatively harmless (“ innocuous " V
In subsequent years the Japanese liver-flukes were the object
of repeated scientific investigation. It was soon recognised that they
bore a close anatomical resemblance to the Chinese liver-fluke
discovered by McCoNNELL ; as a matter of fact, all later authors arc
agreed in declaring them more or less definitely as identical with mu
another as well as with the Distoma sinense. A comparative study ol
the papers published by these authors, however, reveals an interesting
fact, namely that so far as specimens of Japanese origin are concerned
these authors, with one exception, do not seem to have ever again seen
the large variety, i.e. the Dist. innocuuni of BAELZ. Thus in 1SS1
IjIMA, when describing Dist. endcmicum , gives the aver.ige length of
the body as ilfmm., whilst the largest specimens measure i t, tin-
smallest 8 mm. in length ; the breadth is 2 to 2f mm. LEUCKART
who named the Distoma sinense of COBBOLD Distomum
spathulatum , examined part of BAELZ’S original material md
specimens sent to him by IjIMA ; he found the worms to vary in
length from 10 to 13 mm. and in breadth from 2 to 3 mm. I saw tln-
material myself at the time and do not remember having noticed any
remarkable difference in size of the specimens. In iqoo, KATSURAPA
published a very exhaustive paper based upon 76 post-mortem
examinations in which altogether an enormous number of parasites
were found ; in 21 cases indeed their number exceeded r.ooo and in
five there were between 2,000 and 5,000. It may be mentioned hen
in passing that KATSURAPA for the first time states that in r;v-es < !
strong infection the parasites also occur in the pancreatic duct
causing there sometimes no perceptible changes, but generally .*
dilatation of the ducts combined with a thickening of their walls
occasionally, one may even observe inflammatory infiltration and
hypertrophy of the connective tissue in the neighbourhood which
seems to lead to atrophy and degeneration of the glandular substam <-
With regard to the question which interests us here, it is to l>r
regretted that KATSURAPA does not give exact measures of individu.il
specimens, but averages only derived from measurements of a certain

M*

number of worms found in separate cases. I lius, five individuals
measured of one case gave an average of IPOmm length by 2*50
breadth ; 21 individuals of another case ga\

length by 2*40 mm. breadth; and 35 individuals of a third case give
14-06 mm. by 3*88 mm. Nothing need be said concerning the two
first cases ; in the third, however, it is almost certain that large
individuals of 16 to 1 8 mm. length have been present, although the
author does not mention it especially. KATSURADA explains the
variations in the size of the worms by circumstances such as size of
the host, number of parasites actually present at the same place, &c,
and says that “in cases where few worms only inhabit a full-grown
man many of more considerable size arc found among them ; whilst,
on the contrary, in children, dogs and cats the worms are generally
smaller.”

This view, by the way, which was still very generally adopted not
so many years ago, is not supported by actual observations. I have
for several years paid special attention to this question, but all my
experiences tend to show that every species of parasite has a size of
its own. There may occasionally be individual specimens which
tiespass the normal size of their race more or less markedly (just as
this occurs in Man and higher animals), but, as a rule, their size is
very much the same, whether they live in a large or in a small host.
A full-grown Ascaris lumbricoid.es is as large in a child of one years
age as it is in an old man of seventy ; and the common liver-fluke is
not smaller in a goat or a sheep than it is in a cow, nor is it, when
lull -grown, smaller in a narrow bile-duct than in a wide one. In the
a ove mentioned case of Distoma sinense observed by Dr. CUFFEY.

C 77s thlckly crowded the bile-ducts of medium calibre;

I C 6SS’ not a s*n&Ie small specimen could be detected among the

osc ones. The small Distoma endemicum, on the other hand.

indivirt 1 SPeCi7 S'/G cIuke *rrespcctive of the number in which the
mdmduals may be present in a given host

collected in rVatlons far spoken of had all been made on material
al o been H Pan' a"""8 ^ however, the parasite had

Extreme 0*17 , ” SUbj6CtS other parts of the

I do n”:pors the s aS Tea' TOnk‘"' island Of Formosa,

refer the reader L 1,‘erature on earlier cases, but may

r O an article by R. Blanchard, “Note sur quelques

143

vers parasites de l’homme (1891) ” in which the necessary references are
given. In this same article BLANCHARD also records three new cases
from which the parasites obtained had been submitted to him for
examination. The material consisted of two bottles containing each
several hundreds of Distoma sinense which had been collected at the
post-mortems made of two Annamite soldiers in the military hospital
of Hanoi'. A third bottle from Hanoi contained, in addition to a piece
of liver, an extraordinary number of parasites ; so many indeed that
the author says that without the positive affirmation given it would
have been hard for him to believe that they had actually been derived
from one and the same autopsy. At another place (1901, p. 581), he
says that one might estimate the number of these worms as being
rather above ten thousand. The statements BLANCHARD makes with
regard to the external aspect of the parasites are so interesting that
I reproduce them in the original language (1891, p. 5). “ Dans l'un

des deux flacons requs de Mr. le Dr. TREILLE (i.e. those mentioned
above in the first place), tous les Distomes sont de grande taille et
semblables entre eux. Dans le second flacon, ainsi que dans celui que
j’ai re<ju du Dr. LOYE (the third bottle), on remarque au contraire des
Vers de deux sortes: les uns, d’aspect tres fonce, longs de 14 milli
metres, larges de 3 mm. 2 ; les autres, beaucoup plus clairs et mesurant
seulement 8 millimetres sur 2 millimetres.”

Thus, there was again one case in which the large form was present
alone, whereas in the other two both were represented, but so clearly
recognisable from each other that the author immediately goes on
saying : *' A premiere vue, on dirait qu’il s’agit de deux especes
distinctes ...” The conclusions drawn by BLANCHARD from these
observations will be discussed later on ; on the present occasion, 1
wish to draw attention to the fact that the large specimens were " d un
aspect tres fonce,” or, as I interpret it, strongly pigmented, whilst the
smaller specimens were “ beaucoup plus clairs,” that is to say without
pigmentation. The size of the worms recorded by BLANCHARD is
somewhat below the average for both varieties, very probably, there
fore, they are somewhat contracted owing to the action oi some
stronger reagent.

Reviewing now what has been recorded on the preceding pag< s
we have the following facts: The original Distoma sinense of
McConnell and COBBOLD has up to now chiefly been found in

M4

Chinese and is at present the only species of liver fluke known from
this race. It readies an average size of 18 by 3 5 mm. and has a more
or less opaque body owing to the presence within the body
parenchyma of granules of a dark pigment. In Japan, similar flukes
are also very common in the liver of Man and certain animals; but
here they vary, especially in size, to such an extent that one of the
first observers (Baelz) felt justified in establishing two distinct
species. One of them coincides in size with the Distoma Stnase and
also possesses the pigment granules in its body parenchyma It is
in Japan, apparently not very common, for some observers
subsequent to BAELZ have not come SCTOSS ,1 at all during their
investigations, whereas Baelz himself considered it as harmless-
presumably because he never saw it in large numbers -and
ATSURADa explains the large size of the specimens he saw in
certain of his cases by the fact of few individuals only being present
at tie same place. Hie second species is markedly smaller, attaining
ai average size of n to 12 by 2 mm. only; it is not pigmented and
^ I p ais t° be very common in Japan as indicated by the name
en emicum ’ and, furthermore, by the fact that several observers
ns form alone during their researches. There are finally the
three cases recorded by BLANCHARD ; in one of then, onlv 'large

;r0;;;err' in the tw° °th^ * ^ ^ «»»

,ighter ’ r remarkab,e by its dull colour, the latter ranch

thelwo^r °bsetVations 1 <*™ot but draw the conclusion that in
halet d^ whhyt red;arietieS °f sinense we actually

is indigenous on the ' As^tic^ f™*' ^ °' 'hem ^ ^

occasionally also found f r c°nt'nent. especially China, but is
Eastern Asia) the otl ” • ^an<^ probably the other islands of

Annam and Tonkin r ^ -T Inr,l"enous in Japan and common also in
This view Ss rS ^ *" al°n& the “«»•« ™ the Far East).

As quoted above, his ^ ^ C°nclus,on arrive<l at by BLANCHARD,
was that he had two disti ,n?PreSS!°n 0,1 seeing the two different sizes
examination, however C Spcc‘es before him ; a close comparative
t,on anatom ique est In m,n((fI ^im of the contrary. “La disposi-
dimensions de deux cas : la structure e, les

implement que les nIu~ " ^alement indent, ques. On constate

Plus Petits mdividus ont les vitel.ogines moins

developpes et l’uterus charge d’oeufs en moins grand nombre. <
observation demontre done que les petits individus sont des jeunes,
non encore parvenus a toute leur croissance ; elle explique en menu-
temps leur aspect plus clair.” I may confess that for a certain time
I held a similar view; but it cannot be upheld any longer lor the
following reason: Supposing that the small individuals were indeed
young, i.e. in the beginning of the period of their sexual activity,
one ought to expect that the large individuals would be without
exception perfectly sexually mature and full of ova. But this is
actually not the case. In the Hamburg material I have for examina
tion there are two immature specimens, both with a uterus developed
to its full length but containing, in the first specimen, only spcrmat< -/■ *
(the seminal receptacle containing nothing but a number of egg 1 '
and in the second specimen a number of loosely grouped ova wh
are still far from filling the coils as they do in ripe individual
Nevertheless, these worms are unmistakeable representatives of the
large species, for they measure, the first 1 3‘5 by 2-5 mm., the second
12*5 by 275 mm. ; they show, in addition, the size of the suckers and
the brown pigmentation distinctive of that form. One similar
specimen was found in the material from the Port Said case. I- or all
of them, the explanation alluded to above does not hold good . they
prove, on the contrary, that the large species when the period of in
sexual maturity begins has already reached, or even exceeded, tin-
maximum size of the small form. Unfortunately, the scant material
I possess of the latter does not contain any young individuals; hut
the fact just recorded suffices I think to show that the two forms
cannot be one and the same species.

If they are different the question arises, whether they are to be
considered as full species or as mere varieties. I suppose that many
of my fellow-helminthologists, regarding, in the first place, the aim m
peifect similarity of the internal organisation, as compared to which
the differences in size, &c., appear insignificant, would find tin latter
alternative quite sufficient to cover the facts, assuming perhaps
so-called “ local varieties.” I unhesitatingly proclaim the former
as being more logical and more consistent. The chief character of t In-
species, in contradistinction from the variety, lies in the constancy with
which it preserves its specific peculiarities. The variety is inconstant n
a larger or lesser degree and alternates with typical specimens of the

J

146

species. A similarity in the internal organisat ion, even if it is very
close, is not decisive for a variety, for otherwise it would be a simple
logical necessity to consider for instance the cat as a mere small
variety of, let us say, the leopard or the rat as a large variety of the
mouse. Both cat and leopard, and rat and mouse, are internally
almost absolutely alike and yet they are no varieties but genuine
species, for cat and mouse preserve their specific peculiarities through
generations, and never will one of their offspring suddenly be a
leopard or a rat, nor will the offspring of a leopard or a rat ever
become cat or mouse, although, hypothetically, a simple enlargement
(or reduction in size) of their bodies and internal organs combined
with some changes in colour, &c. would suffice to transform the one
into the other. With regard to these animals, therefore, (and to
almost all others as well) the size and some external features, like
colour, &c. are not considered as determinative of varieties, but of
species, and it is only the higher classified tory units such as genera,
sub-families, &c. which become determined by the greater or lesser
similarity in the internal organisation. Such principles of classification
are, so far as I can see, universally followed by systematists with
regard to the larger animals, and I do not see any reason, why they
should not be applied to parasitic worms also ; so much the more so,
as careful observation and comparison reveal among them quite a
similar gradual increase (or decrease) of structural differences as exists
among the larger animals. I thus hold that it is simply logical and
consistent to consider the two Opisthorchis sinensis- forms as two
distinct species rather than as varieties of one species.

1 , a,Ve *n a'JOVe explained the reasons for my opinion at some

ecause the question what in helminthology is to be considered
as a species still needs being definitely settled. It appears that

suhif* nU*Mi writeis> zoological and medical, on helminthological
, • J C,S bl C lng to the old assumption that there exist compara-
of varviiT ?*eC.ieS Parasdes only, but that these species are capable
to host 1IJ.Slze and organisation to a considerable degree according

Iwthirt1 \ U iS my “™ction that th,s view must be

based f * ^ °nCC* ^ helminthology is to become an exact science

fathers. Neither is'V' °n Prec0nceptions inherited from our fore¬
forms us iden justifiable any longer to declare all at once two

as , dent, cal simply because they agree in their general structure.

T47

nor is it sufficient to-day to give of new forms of parasites a superficial
account of some general characters, if such descriptions are to be of
real scientific value. It would not be a difficult thing to enumerate
dozens of modern papers of this sort ; they look and read quite nicely,
but nothing would have been lost had they never been published at
all. I will not mention any names, nor is it my intention to blame or
to offend those who do not share my scientific convictions, but 1 desire
to convince them, in the interest of our particular branch of science,
that by this time certain views considered as almost axioms b) our
predecessors can no longer be upheld.

The question as to the denomination of the two species of
0 pisthorchis sinensis in accordance with the modern rules of scientific
nomenclature is easy to decide. There is no doubt that the large form
is the same as originally found and described by McCONNEI.L ; it
must therefore retain the specific name sinensis COBBOLD. The
smaller species is, unmistakeably again, the same as first recognised
by Baelz and distinguished under the designation Distowa
endemicum ; I therefore re-establish this specific name for it. Both
species are members of a natural genus which might have been created
before this on the ground of the peculiar configuration of the testicles,
but becomes now more distinctly separated from 0 pisthorchis by the
fact that two different forms exhibit the same character.

6 lonorchis, n. g., O pisthorchiidarum.

Differs from 0 pisthorchis sensu strictiori by the fact that the
testicles are not notched or lobate, but distinctly ramified, the
branches, crossing the intestinal caeca on their ventral side and
extending very near to the body margin. Another difference seems
to he in the shape of the excretory vesicle. In O pisthorchis
s- str. this is \ shaped with a very long stem running in the form of
an S through the space between the testicles and bifurcating at the
posterior border of the seminal receptacle into two rather short
branches from the tops of which the main excretory vessels arise. In
C lonorchis y the Y shape of the vesicle is no longer clearly recog¬
nisable ; the unpaired tube simply becomes somewhat widened at its
anterior end assuming sometimes, but not always, the shape of an
nregular triangle the two upper angles of which might be considered

as the homologues of the branches in O pis/horchis s. str. I he main
excretory vessels do not arise from these angles, but somewhat
farther backward, at about the lower angle of the triangular widening
of the tube (see fig. 4).

Type: Clonorchis sinensis (COBBOLD) 1875.

Clonorchis sinensis (COBBOLl)) 1875.
(Synonym : Distomum innocuum BAELZ 1883.)

Full grown specimens vary in length from 1 3 to 19111111. and in breadth
from 3 to 4 mm. At the commencement of the period of sexual
activity the body has attained 12 to 13 mm. in length and 2*5 to 3 mm
in breadth ; such specimens are recognisable by their uteri in which
the ova are comparatively scarce in number and loosely grouped, i.e
with more or less wide empty spaces between the individual ova
The oral sucker averages 06mm. in transverse diameter; the lowest
figure 1 have found in one specimen (of about 60 measured) was
0 52 mm., the largest which is not uncommon 0*63 mm. ; usual
measurements between 0*58 and 0-62 mm. The ventral sucker
averages 0-47 mm. in transverse diameter ; lowest figure found in one
case 0-39111111., highest figure observed once also 0-5211101.; usual

measurements between 0*45 and 0*49 mm. The ratio of the suckers
thus varies in the main between 15: 12 and 16: 12. The pigmentation,
i.e. the deposit within the body parenchyma of numerous fine granules
of a yellowish or brown substance, is very characteristic of this species
• ometimes it is so little developed that the preserved worms appear
white and the pigment granules are only detected in cleared specimens
it tie microscope, yet may in other specimens assume such an
extent that the parasites become perfectly opaque and of a rusty
) ow, greyish, or brownish tint throughout. In cases of little
Jfi . . tl0n’ tbe granules accumulate more especially behind the
of thThoH m thC S,ldeS °f the Pharynx and at the posterior extremity

the outhnes of^esticllT'-md^6 alTg ^ intestinal CaCCa ^

the testicles f 1 d Semmal recePtacle. The ramifications of

the bodv • "f aU ^ l°n^’ *n accordance with the greater breadth ot
testicle the bn nH Unfrequently be recognised that in the anterior

main stems. All f°Ur’ the Postenor testicle from five

em ,e 111 about one plane which runs parallel to

r49

the ventral surface of the body ; there are however very often, in one
testicle or in both, other branches or, more precisely, bulging* of one
or two main stems which project towards the ventral surface and
appear as thick, compact bodies with irregular outlines. They were
already seen by McConnell, but interpreted as the whole testicles,
whereas in truth they are only parts of them, and are not even
constant in their presence. The ovary is usually trilobate, but very
often shows, in addition to the large lobes, three to six
smaller ones which are sharply separated from the former. The yolk
glands vary somewhat in their extent. They normally reach from
the level of the ovary to the ventral sucker, but it is not at all
uncommon to see them, on one side or on both, commencing and
ending somewhat farther forward or somewhat farther backward of
the usual place. I hey present in Cl. sinensis, a striking peculiars
inasmuch as almost constantly a certain number of the groups ot
follicles of which they are composed remain undeveloped ; sometime
one only on one side, sometimes more, on one side or on both; tlu
bands represented by the glands as a whole showing thus one or
several interruptions by empty spaces. In perfectly mature specimens
the seminal vesicle extends about as far back as the middle of the
uterus. 1 he remaining organs do not show any peculiarities.

The average dimensions of the eggs are 0*029 mm. length and
0*oi6mm. width; the limits of the former being 0*026 and 0*030110,
and of the latter 0*015 and 0*017 mm.* la many specimens of tlu
species the eggs show a distinct narrowing towards the anterior
extremity, and their rather high lid is marked off by a sharply
projecting brim. I have however also seen specimens in whose ov
these peculiarities were but little pronounced (see fig. 7).

An interesting variation chiefly observable among the specimens
of the Port Said case concerns the colour of the uterus as a whole In
some individuals, this is only a light yellowish-brown, even in tin*
most anterior coils of the organ, whereas in other individuals the whole

lhere exist, of course, among the immense number of ova in an individual
. orm always some which are either larger or smaller than the rest, or even
K 1 ently misshapen. In my opinion it is of no use to carefully record thr
measurements of these eggs also. For the description and definition of a spe< ;•

15 much more important to select for measurement those ova which appear to be
normal and to present the size and shape typical for the species. It may be addrd
ln Passtng that young worms with few ova in their uterus usually do not afford
normally-shaped and normally-sized ova.

I5°

uterus, with the exception of a few of the hindmost loops, appears
almost black. As there is otherwise not the slightest difference
between the examples of these two extremes neither in the size of
the body, nor in the size of the suckers, &c. 1 do not quite know how

to account for these variations. To a certain degree, they are
evidently due to the number of ova contained in the roils, hut to
another degree just as evidently to a darker shade of the egg- sheik
themselves. Since the two extremes described above are not sharply
separated, but connected with each other hy numerous intermediary
tints it is perhaps not unjustifiable to assume that the shell-substance
elaborated by the shell-glands becomes darker with the growing age
of the parasites, and it may be noted in this connection that the latter
live in, and feed on, a coloured substance, the bile.

Clonorchis sinensis is chiefly a parasite of Chinese, but occurs
rarely also in Japan— i.e. according to what we know at present. For
it is not at all improbable that the worm will he oftener found as soon as
attention is paid to its existence ; it appears also not unlikely that it
may be restricted to certain localities. Up to now it does not seem to
have been found in animals.

C lonorchis cndcmicus (BAELZ) 1883.

This species practically takes the place of the true Cion, sinensis
in the literature published after 1883 and is at present the one usually
described m text books, &c. as “ Opisthorchis sinensis." Its measure-
are given by the authors as 10 to 13 mm. length and 2 to
3 mm. breadth. The smallest specimen actually at my disposal (it
comes rom the Annamite case described by BLANCHARD) is sonie-
C!ntrfC‘ed and measures 6 by 1-8 mm. ; my largest and fairly
i-he ~ en e sPec*men (°f Japanese origin) measures 13 by 2'6 mm ;
oraUt!!JIning lndlvuluals ranSe in size between these extremes. The
diameter dVera§es 0 43 to 0 45 mm. in transverse diameter ; smallest
another ^ the smalIest specimen mentioned above and

,ong Md ■■6m-

0-5 mm. Ventral * fPeClmen 10 8 nun- loI>E and 2-3 mm. broad)
diameters observed o-J, “ TT* °'37 ‘° °'4° ' S,"alleS,
(in another long 8 mm h ' ^ SmalIest 5Pec™en) and o'34"®j

°’45 mm. fin tl, Ji ’ b d 17 mm)i largest diameter observed

argest specimen above mentioned). The ratio of the

suckers varying between about 13 : 12 and 15: 12 is therefore fairly
similar to that in Cion, sinensis, but in actual size the suckers are
quite different. Among the specimens at my disposal there is none
with pigment-granules in its parenchyma. The yolk-glands normally
have the same extent as in Cion, sinensis, but present similar
variations in regard to the relative position of their anterior and
posterior terminations as in that species. It may also happen that ..I
the groups of follicles of which they are composed some remain
undeveloped, but this occurs, so far as may be inferred from my
limited material, much more seldom and the actual state of things is
perhaps best described by saying that the absence of one . ir sewn I
groups of follicles is the rule in Cion, sinensis, and the exception in
Cion, endemicus. The seminal vesicle ends (so far as visible) at
about the limit between the first and second thirds of the length -a
the uterus; this character, however, had better not be considered a
distinctive because it may vary.

The eggs have about the same length as those of the preceding
species (0*026 mm.), but their width is decidedly somewhat smaller,
amounting on an average to o-oi5 mm. with the lower limit at 0-013.
the upper at o-oi6mm. The narrowing towards the anterior end is
in the main not so marked and the margin of the rather flat lid nm
so sharply projecting as in Cion, sinensis (see fig. 8); but these
differences are on the whole very slight and not recognisable in r.i
specimen. With regard to the colour of the uterus as a whole, van.i
tions may be found similar to those described in the preceding spe* -

Clonorchis endemicus, according to existing records, is vm
common in Japan, and presumably so in Annam and Tonkin ton. In
Japan the localities especially stricken with the parasite are situated
on the sea-shore; it is possible therefore that its geographical i.mgr
chiefly extends along the shore in the other localities also.

Apart from Man, the species is reported to have been found in
certain animals of Japan, such as cats, dogs and hogs. Several
specimens taken from the latter host, which I owe to the kindness of
rofessor J ANSON, of Tokyo, are indeed Cion, endemicus, for tin v
present all the peculiarities of this species. I am, however, somewhat
doubtful with regard to the statement that it inhabits the liver
ot the cat also- at least as a normal parasite of this animal, in 1 880.
iJIMA after an examination of specimens derived from this source

writes that the worms “from the liver of cat showed slight differences
inasmuch as they possessed very fine spines in the skin and were of
smaller size than the former (viz ( the human live:

Perhaps this cat-distome from Okayama is to be considered as a
distinct species, but I firmly believe that in Tokyo, Dtst endmian
does sometimes inhabit the liver of cats" (1886, p 49). This latter
possibility can certainly not be denied and is even most probable, but
in cases of this sort Lion, end appears to be

occasional parasite of the cat, whilst the smaller form menti ned
IjlMA probably represents one of its normal parasites I have
examined several cat-specimens which were sent from Japan to
Professor Leuckart in Leipzig years ago (by whom I do not km
and of which Professor CHUN kindly allowed me to take some in
exchange for other parasites. They arc unfortunately in such a poor
state of preservation that not very much is left to be seen in them;
t e general impression they make upon the observer is certainly rather
1 erent from that of the genuine Cion, cndcmicus. If one considers,
moreover, their small size (those in my possession do not
exceed 4-5 mm. in length and 0-9 mm. in breadth, but are as it seems
not yet quite full grown) the presence of fine spines in their skm

as reported by Ijima there is no doubt left that thev cannot be
Leon, endemic us.

I , ^ worth being borne in mind in the examination of

cannn?50" ° 1“man>°Pisthorchiosis is that this Feline Opis/horchis( it
may non ^ ^ Whether il is ai1 Opistkorckis or a t lonorchis )

sionallv o'Th ^ “ *** also* ■» 1 *°* « is occa'

certain state . T y CatS and hogS' 1 a,n °ven inclined to think that
three wonns^ound Tn KATSURADA’s PaPer OW p. 481) alluding to
Saga, which presented ““ °f sevcnteen from the province ot

breadth of 0 06 m avera^e le“gth of 5-16 and an average

infection with the FeiineV* ^ ^ eXp,ained by such 00035101131

I53

LITERATURE

Baei.z, E., 1883. Leber einige neue Parasiten des Menschen, in Berl klin.
Wochenschr. p. 234-238. (Not accessible.)

Blanchard, R., 1891. Note sur quelques vers parasites de l'homme, in C. R. Sor.
Biol. Paris. Stance du 18 juillet.

Cobbold, Th. Sp., 1875. The new human fluke, in The Lancet, Vol. 2, Sept 18,
P- 423-

Giard, A., and Billet, A., 1892. Sur quelques Trdmatodes parasites des boeuf*
du Tonkin, in C. R. Soc. Biol. Paris, Vol. 94, p. 613. (Not available.)
Ijima, I., 1886. Notes on Distoma Kndemicum, Baelz, in Tourn. Sc. Coll Tokyo
Vol. 1.

Janson, J., 1893. Die Ilausthiere in Japan. IV, Die Krankheiten der llausthi.rr
in Japan— (2) Parasitare Krankheiten, in Arch. wiss. u. prakt. Thierheilk,
Vol. 19. (Not available.)

Janson, J., 1895. Die tbierischen Parasiten bei Japanischen Wiederkauern, in
Mitth. Ges. Nat. u. Volkerk. Ostasiens, Tokio, Vol. 6, p. 272.

Kaisurada, I'., 1900. Beitrag zur Kenntniss des Distomum spathulatum. in
Ziegler, Beitr. z. path. Anat. u. allg. Path., Vol. 28, p. 479-505.

Leuckart, R., 1876. Die Parasiten des Menschen, etc., T.eipzig 11. Heidelberg,
Vo!. 1, p. 336-355.

MACGREGOR, W., 1877. A new form of Paralytic Disease, associated with the
presence of a new species of Liver Parasite (Distoma sinense), in Glasgow
Med. Journ., January, 1877. (Not available, quoted after Cobbold, Parasite- .
London, 1S79, P- 3°-) Abstract in The Lancet, 1877, May 26th, p 775.
McConnell, 1. 1*. P., 1875. Remarks on the Anatomy and Pathological Relations
of a new species of Liver-fluke, in The Lancet, August 21st, p. 271-274.

McConnell, I. F. P., ,878. “Distoma sinense” (McConnell), in The Lancet.
March 16th, p. 406.

IC)02‘. Tasciolopsis Buski (Lank.) (= Distomum crassum Cobb.), ein

Ori ’ vV®mg hekannter Parasit des Menschen in Ostasien, in Ctrbl. Bakt
ung., Vol. 31, p. 574.58,.

Railliet, A 1800 t bo

Naturaliste, (2)Nr'-oPnTteS deS. animaux domes‘i(lu^ an Japon. in Le
1898, p. 30.) /9’ P' 43’ 15 Jmn' (Not avai,ab,e' quoted after Railliet,

Railliet, A., and Marotel, G ,8n8 t n , .

e. de, bl,mes « Cochiac^in bM“"

*54

EXPLANATION OF FIGURES

Reference Letters.

App. — Lateral pouches of the
oral sucker.

C .ex. — Lateral excretory vessels.

C. P. — Cirrus pouch.

D. ej. — Ejaculatory duct.

Disc. — Edge of the ventral disc.
Dr- Retracted grooves (“pseudo¬
suckers ”).

Dr. — The same organs, everted.
hit.-- Intestine.

L.C. — Laurer’s canal.

/. 7 est. — Left testicle.

Met. — Metraterm.

Nerv. — Nervous system.

Oes- -Oesophagus.

0v.~ Ovary.

P.ex. — Excretory pore.

P.Gett. — Genital aperture.
Phar. — Pharynx.

P.P. - Pars prostatica.
R.sent.— Seminal receptacle
r. T est. Right testicle.

Sh.Gl. — Shell gland.

Ut. — Uterus.
lag. — Vagina.

V .ex. — Excretory vesicle.

V. ex./at. — Lateral branches of
the excretory vesicle.

V it. Yolk glands.

V it.D. — Unpaired vitelloduct.

V .sew. Seminal vesicle.

Fig. i.
Fig. 2.
Fig. 3-

Fig- 4-

Fig- 5 -
Fig. 6.-

Fig.

Fig. 8. -

Rurytrema patter ealicum. Specimen from Japan seen from
the ventral aspect. Enlargement about it.

Ilurytrema coeloniaticum. Specimen from Hong Kong seen
iom the ventral aspect. Enlargement about II.

G astrodtscus secundus seen from the ventral side. Enlarge-
men about n. 1 he opening in the anterior part of the
ven i a surface leads into the genital atrium from the
ottom of which the papilla bearing on its top the male
and female genital openings arises.

GaaZltiSCUS. ■ secun<fus ■ Median sagittal section; the

boiiriiicr CUt f” U iaVe been conipleted from the neigh

intern if SCC 10nS so as to give a general picture of the
■nternal organisation. Enlargement about 16.

Ventral , Specimen from the Port Said case.
Ventral aspect. Enlargement about 9.

vTT endemuus- Specimen from Okayama, Japan,
the” preceding fi EnlarSement exactly the same as that of

comparison ^Thf^ S° t0 alIow of an aPPr0Pnate

visible in ti>,v . unequal length of the intestinal caeca
other individuals^01”160 1S accidental and not Present ,n

eggs, each* errer^t' L- c^,e^ forms presented by the

worm EnlinS *' vCn ^rom a separate specimen of the
Clo \ ■ gement ca- 990.

and under^exaedv^th EggS fi^ured in the same manner

ceding drawino- J . f saJFe enlargement as in the pre-
& as to allow of comparison.

F‘g /.

”«*■ del.

1

PLATE VI/.

Fit!

Pax

PLATE VIII

_

A Looss, ad nut. del.

P P Imp

PLATE IX

Fig. 5:

Fig. 7.

Fseoi.

Vsenc

POen.

Sh.Ct,

R.Serro

C.ex

LC

L7!‘sl

A. Looss, ad nut. dal.

the PRESENCE OF SP1R0CHAE TA
DUTTONI IN THE OVA OF ORNI-
THODOROS MOUBATA

L_

157

THE PRESENCE OF SPIROCHAETA
D UTTONI IN THE OVA OF ORNI-
THOD OR OS MO UBA TA

BY

Capt. R. MARKHAM CARTER, I.M.S.

From the Runcorn Research Laboratories of the Liverpool School »f
Tropical Medicine

Koch, while discussing the question of the transmission of
Spirochaeta duttoni , the organism of African relapsing fever1 states
that he finds on the surface of the ovaries of infected ticks large
numbers of spirochaetes. He considers that these organisms multiply
in this position in the tick. He further states that the eggs, when first
laid, show scanty infection, whereas in those examined later lie
describes the occurrence of spirochaetes in vast numbers arranged in
densely-tangled masses. This he considers an undoubted proof of
the rapid multiplication of the spirochaetes in the eggs. Lastly, he
again draws attention to the fact that ticks bred from batches of
infected eggs transmit the disease.2 The following observations were
made with the object of confirming the second of these important
statements.

Forty-two, naturally-infected, adult specimens of Ornithodoro >
moubala, collected in the Congo Free State, were fed on January 2;th,
I9°6, upon an animal infected with Spirochaeta duttoni. These tick
were again fed on an uninfected rat on July 26th, 1906. This rat
showed infection on August 1st, innumerable spirochaetes to the field
were seen on August nth. As no eggs were laid, the ticks were
again fed ; ovulation then commenced, but it was very scanty'. Some
twenty-six adults were therefore fed once more on an uninfected rat
ten day's later. Large numbers of eggs were now found in batches
about the dish ; 32 ova were taken from different collections of the--
eggs and microscopically examined to determine the presence or
otherwise of the spirochaetes. Six of them were found to be infected

The method used was as follows-. — The egg, having been placed

I5«

in the centre of the slide, was pricked with a very fine capillary tube,
and the contents were quickly withdrawn until the egg capsule became
transparent and collapsed. The egg contents were then emptied,
forming a tiny drop, on t lie slide, which drop was quickly smeared into
a thin film with a fine-pointed, cold, glass rod. The more rapidly
these stages succeeded each other, the greater the perfection of the
difficultly^ made film. The preparation was quickly dried at 450 C.
and was then fixed in absolute alcohol before being stained by 1
modification of Romanowsky’s method;3 finally' the smear was
examined with a Zeiss apochromatic 2 mm. objective and a No. 18
compensating eyepiece, fitted to a tube length of 250mm. (x 2250)

I he number of the spirochaetcs found varied from but three in the
whole contents of an egg, to as high as 45 counted parasites to a field,
with, in addition, one or more spindle-shaped collections of
innumerable organisms. As was observed by Koch it was found that
most spirochaetes occurred in those eggs which had been laid longest

Morphology of Parasites occurring in the liggs (see plate X).

As well as the typical forms of the spirochaete, occurring as th<
did in large numbers, several peculiar varieties were also noted ai
will be discussed in detail below.

By far the majority of the spirochaetes were of the typical simp
spin ar form. These varied in length from 5^ to about 20y (sing
parasites) and were seen to occur singly and in groups of three to foi
^ llowever> anc* this was seen in the most highly infecte
wer_ * ,r°chaetes were arranged in remarkable collections an

are ocnsT C!' Spiral f°r spira1’ with each other. These grouf
those sniropr Spindle'sIlaPed by reason of the decreasing length <

iatera,iy as ""th *■* ^ -

UtnhboundPb“ndfe of °LhkTSe tHeSe gr°UPed SPir0Chae“

nucleus blepharo jP'Cd^ SI11^e spirochaete presents no evidence of
lateralflager^^ no indications c

shaped on sectionS & ° seems to be a spiral, possibly flat and ribbon

deeply stained, but h termin£d P°>nt of the spirochaete is oftei
to a point, possibly to f °r may stain faintly and tape

i°ng forms of thp °rm * terminal flagellum. In, especially. th<

ganism, transverse, faintly-staining section:

159

occasionally occur. These lightly-stained areas nrnir ai about tbr
middle of the parasite and occasionally excentriealh n * nr <>r n* ’r
places; this is also particularly noticeable in the ex ; i* n.»IK V <
forms. It seems as though the deeply- staining, - *• rhrxnj*

coreof the parasite were absent and the more light!) t rung [-■ ; ' ■ ■
alone remains. The same appearance is seen at tin ' me . , <
some parasites ; in many the condition is exaggerated m ■ hr
chromatin of the spirochaete exists only as a fragment. 1 «an rr*» <> f
granules (figs. 8, 9, 24).

In figs. 19a and 1 9^ are illustrated peculiar f nr: i I In , .

apparently single organisms, in which longitudinal :wr, •! .

seems to be commencing. The undivided portion of • «, l%4tr,
is normal to within a short distance of its bifujr.it • 1 r •

chromatin becomes collected into one or two large granule • ]] .

divided portions lie side-by-side in parallel curves, and ’ rigs**
tation occurs in them it is noticeable that their corn s;, :
lie directly opposite each other.

Swellings, usually occurring one to a parasite, are fro; efl-l, T
in both normal and fragmented spirochaetes. The-. . r, *-i
terminally or in the length of the parasite and are of thro • . ; . * hr

rSl 'S the smaller insists of an oval, homo o ,* ,

chromatimc mass placed either centrally or laterally o„ ,

( gS' a"d 30)- The second type is slightly larger. ,* , , ,

irvr-r vesic,e m which tw° eh™1 *• . - 4 -

r 1CS at.the Slde of the P^nsite and is attar!,, d ■ ,t b 4

very fine connection (fig. 29). This form is by far the r

seen but once. The j 1

of a vesicle contain,! a dLt ^ P™',,

.

confirming these suggestive° observations1}! the " '

specimens and of more carefully PrePared rnaten.!1' ° f *

i6o

From these observations we conclude :

1. That ticks infected by spirochaetes lay infected eggs.

2. That multiplication of the spirochaetes probably takes place

in the eggs.

3. That morphological changes in the spirochaetes also occur in

the eggs.

LITERATURE

1. Koch, 1905. Vorlaufige Mittheilungen iiber die Krgebnisse einen Forschungs-

reise nach Ostafrika. Deutsche Medizinischc Wochenschrift. 23 Nov.

2. Dutton and Todd, 1905. Memoir XVII, Liverpool School of Tropical

Medicine.

3. Stephens and Christophers. The practical Study of Malaria, and edition,

Williams and Norgate, London.

4- JlREiNL AND Kinghorn, 1906. Memoir XXI, Liverpool School of Tropical
Medicine.

Plate X

IN EGGS OF ORNITHODORUS MOUBATA.

A NOTE ON THE

THERAPEUTICS OF TRYPANOSOMIASIS

BV

Professor BENJAMIN MOORE, M.A., D.Sc.

M. NIERENSTEIN, Ph.D.

AND

JOHN L. TODD, B.A., M.D., C.M. McGill

Thomas and Breinl* found that atoxyl would almost invariably
drive the trypanosomes from the peripheral circulation of an infected
animal, but, since the parasites frequently recurred, it seemed possible
that they might exist somewhere else in the body of their host in a
form uninfluenced by this drug. It was therefore determined to
commence a series of experiments in which animals first treated by
atoxyl were subsequently treated by some other drug. A variety of
combinations have been tried. That which has been by far the most
successful in the treatment of rats infected with Trypanosoma brucei
is atoxyl, followed by small doses of bi-chloride of mercury after the
trypanosomes have left the peripheral blood. Animal experiments
are still in progress.

This communication is made in the hope of inducing those who
have opportunities of treating patients to try the effect of atoxyl
followed by bi-chloride of mercury on their patients.

Thomas and Breinl; Memoir XVI of the l.ivetpool School of Tropical Medicine.

V

AN AUTOMATIC OILER FOR THE
DESTRUCTION AND PREVENTION
OF MOSQUITO LARVAE IN CESSPOOLS
AND OTHER COLLECTIONS OF WATER

K

i65

AN AUTOMATIC OILER FOR THE
DESTRUCTION AND PREVENTION
OF MOSQUITO LARVAE IN CESSPOOLS
AND OTHER COLLECTIONS OF WATER

BY

E. H. ROSS, and H. C. ROSS,

LATE SURGEONS R.N. AND MEDICAL OFFICERS’ DEPARTMENT OF
PUBLIC HEALTH IN EGYPT

( Received January 16th, 1907)

In the large towns of Egypt the breeding of mosquitoes is
practically confined to the cesspools. With the exception of
Alexandria there are at present no schemes for the disposal of
sewage on modern lines, and therefore it passes into cesspools under
or near the houses. These cesspools are practically the only
collections of water where mosquitoes can breed, and it is to them
that most attention must be paid in anti-mosquito campaigns.

Besides mosquitoes, all sorts of vermin live in the cesspools;
and in plague-infected towns, the knowledge of the fact that rats
have to depend on these collections of water for drinking purposes,
is of the utmost importance.

In Poit Said and Ismailia, where there have been successful
campaigns, all cesspools are treated with a mixture of crude and
refined petroleum once a week. This has up to now involved very
considerable labour, a constant supervision, and a large expenditure
of oil ; and it was in the endeavour to overcome these difficulties that
the following automatic oiler was designed.

Our aim was to produce an apparatus which would give a constant
supply of oil to the surface of the water in the cesspools ; which
could be easily applied by unskilled labour (native) ; which could be
cheaply made in any country; and lastly, one whose application
would, if possible, reduce the cost of a campaign. We have found by
experiment that this apparatus has fulfilled the requirements.

1 he apparatus is made of metal. We use the empty kerosene

tins in which the crude oii is supplied. The shape is that of ate
with two spouts, one at each end, and is shown in the drawing
It will be noticed that the spouts do not arise from the centre of the
ends of the drum, but from one edge of them ; are directly oppose
to one another, and are numbered A and 13. On the spout B a
weight is suspended. All joints arc soldered.

In A a sand filter is placed. This consists of a glass tube, with
an internal diameter of one centimetre, containing sand. The tube
passes through a cork which is pressed home in the spout. To mate
the sand filter : — Draw out some tubing to a point and cut off so that
the lumen is about half a centimetre in diameter. Place a small plug
of glass wool in the neck so formed and fill up two inches of the tube
with fine sand. Then place another small plug of asbestos wool arc
again draw out the tube round it and cut off.

I he whole apparatus should now be tested to see if it is tighi
To complete it, a piece of galvanized iron wire is fastened to each
spout, each piece being about two feet in length. The two pieces
are joined together above, and from their junction a wire passe:
upwards which must be long enough to reach from nearly the bottom
of the cesspool to the manhole door. This appliance when filled with
water will immediately sink, assuming a horizontal position under
'vater ; but when filled with oil, would float were it not for the weight
on the spout B. Consequently, when filled with oil and dropped into
'va er, it will sink to the bottom, but the end A will be pointing
o?P anc| t^e °if will pass out through the sand filter at the rate

Qf 0jj CU *c cem,nietres a day. The oilers hold nearly 3j l'tre:

A mil ic a.- ^ °^cr 111 a cesspool the following procedure is adopted
oiler is held' ^ ^ Wa^ of t*ie manhole just under the lid The

means of a f " ** t*le sP0llt B upwards and oil is poured into it b)
ground with rh ^ ^ ^en it is allowed to rest horizontally on the

The anna t SP°UtS ^ t0P S° that little or no oil is sPilled (F«* ^
cesspool until ifUf ^ ra'Sec^ ^le w*re and lowered gently into the
water is reached °uC ^ Water It: remains horizontal until the
it sinks under th W ^ ,mme^ateIy assumes its vertical position a-
air-bubbles comin *" 2) No oil will pass out until the

of oil begins. ^ °Ut ° the end A have ceased, when the exudation

167

The wire is payed out until the weight is felt to rest on the bottom
of the cesspool ; it is then pulled up for about one foot and fastened to
the nail. In this way the oiler will rest about eighteen inches from
the bottom of the cesspool. (Fig. 3.) Two litres of oil are then
thrown on to the surface of the water to kill the existing larvae, and
the manhole door is closed. In the oil thrown on to the surface it
has been found better to mix five per cent, of powdered resin, as this
materially prevents evaporation. But resin must not be mixed with
the oil inside the oiler, as it may clog the sand filter.

The exudation of oil goes on at the rate of about 50 cubic
centimetres a day for nearly two months. As oil comes out of
spout A water enters the oiler through the spout B, and therefore it
becomes heavier daily and consequently assumes a more horizontal
position, until in about two months, when there is about three-quarters
of a litre of oil left, the oil comes out of the spout B and the oiler is
suddenly emptied of the mixture. (Fig. 4.) So that in the eighth
week of the oiler’s existence in the cesspool there is a sudden and
large ejection of oil on to the surface of the water.

The reason we have designed this is that if the cesspool is a large
one and the exudation of oil hardly sufficient, any larvae which may
have evaded the oil towards the end of the two months will be
destroyed.

In ten weeks the oiler is removed and replaced by a new one. In
Egypt the cost of each oiler complete without oil is sixpence. The
apparatus has been given a fair trial, and has been found to be
successful. It has saved a large amount of labour and oil, and will
shortly be used extensively in Cairo and its suburbs.

automatic oiler

— — TOR CESSPOOLS -

PLATE XI

SECTION OF GL&Vj fiLT£H

THE ANATOMY OF THE PROBOSCIS
OF BITING FLIES

THE ANATOMY OF THE PROBOSCIS
OF BITING FLIES

BY

J. W. W. STEPHENS, M.D. Cantab.

AND

R. NEWSTEAD, A.L.S, F.E.S.

( Received May nth , 1907)

PART II.* STOMOXYS (STABLE FLIES)

As far as we are aware, there have been only two descriptions, in
recent years at least, of the anatomy of the mouth parts of Stomoxys.
I he first, by Hansen,1 is, as far as it goes, a good and accurate
account, but it is incomplete in many respects, even in regard to
important structures, which we should have thought it impossible to
overlook. Moreover, sections were not cut by this author, and without
these, it is, in our opinion, impossible to understand the relationship of
the parts. The second description is by Giles.2. We feel bound to
point out that this account is misleading and inaccurate ; and that
the same criticism is equally applicable to his description of
Glossina (/. c.)

Methods. — For section cutting we find that a combined celloidin-
paraffin method gives the best result. In this way it is possible to get
intact sections of the chitin, otherwise a difficult matter. Owing to
the opacity of the proboscis, examination with high powers, of
specimens mounted in toto, is almost impossible ; but the whole
proboscis can be rapidly rendered transparent to any desired extent
by bleaching in chlorine gas : the chitin not being at all affected.

* Glossina (Tsetse-flies), was published in Memoir XVIII, 1906, Liverpool

School of lropical Medicine.

172

The material at our disposal having been fixed in alcohol has not
been in a sufficiently good condition for an accurate study of the soft
cellular structures of the labella, e.g., the nerves. We hope eventually
to examine these structures in detail.

The species examined by us has been Stomoxys calcitrans, Linn,
(the common “ stable-fly ”). caught in England.

The proboscis of Stomoxys (fig. i), as in Glossina, consists of three
parts: the lower lip or labium (r), the hypopharynx (6), and the upper
lip or labrum-epipharynx ( a ). The actual length from the base of
the bulb to the tip of the labella measures 24 1 min.; the dimensions
just behind the fork are 130 /x vertically and 150/i horizontally.

The Labium (fig. 1, c) is relatively stout and thick, with the basal
portion swollen and gradually tapering towards both the proximal and
distal extremities, but less so basally. Externally in its entire length,
as far as the labella. the labium is transversely and regularly striated
or latticed, so that the edges of preparations mounted entire appear
faintly crenulated. Scattered over t he labium art* also a number of short
fine hairs, set on transparent spherical areas ; more numerous,
however, on either side of the dorsal groove. Immediately behind
the fork of the labella (fig. 5, e), and articulating with it. are two
conspicuous ventral sclerities (fig. 5, /).

Labella (General account). While the arrangements of the parts
in the labium is easily understood, that of the labella presents, on the
contrary, considerable difficulty, and we must consider the structures
in detail. I he labella are bilaterally symmetrical. They consist of
•in external and internal wall on either side (fig. 20), separated by
soft tissues. The two halves are at first separate, near the base the
sides join ventrally (figs. 21, 22), but remain open dorsally. When the
labella are in the resting position, the internal walls consist of two
strong chitinous plates (fig. 20), which we may divide into a dorsal
and ventral area. The dorsal half (k in all the figures) consists of a
flat plate, which is the true plate of the labella. while the ventral half
is highly specialized to form a series of teeth (j in all the figures) and
other structures, which we shall consider in detail later. If we place
our two hands back to back, the fingers will represent the ventral half
tie teeth, etc.), while the palms back to back represent the dorsal
ia . Further in the process of eversion the ventral half is capable

*73

of being bent outwards and backwards. If we now flex the fingers
this gives approximately the position that the ventral half of the plate
(the teeth) takes when the proboscis is everted.

The two bilateral inner walls are at first approximately parallel.
If we now trace them backwards, we find that at first the inner
walls approach one another ventrally so that they now enclose a
somewhat V-shaped space (fig. 22). The dorsal half of the
plate (£) grows downwards at the expense of the ventral, so that
in fig. 21 we have only a few remnants of the ventral half left
(and here the two sides are united ventrally), and eventually these
disappear, so that we have on either side what was the dorsal half
now occupying the whole inner side (fig. 22, k). These two sides now
become fused ventrally by a median structure (figs. 23. 24), the
axial apophysis (if), which serves to bind them together.

Dorsally the inner sides now become replaced by new structures
appearing, viz., by two lateral chitinous rod-like sclerites, seen in cross-
section in figs. 21-24, r, and medially a structure (r1) appears which
will form eventually the floor of the labial gutter when these two rods
and the median piece fuse to form a single trough.

Labial gutter (r, r1 in all the figs.). I he commencement of the
labium can be seen in a cleared proboscis as a transverse line stretching
across (above) the middle of the axial apophysis. In sections it appears
at about the level of the tips of the fork as a slightly hollowed-out free
chitinous piece. About the same time appear, dorsal to it, two
chitinous rods (r) seen in cross section in figs. 21-28. These rods
are hooked at their edges, and to these edges are attached ligamentous
tissues continuous with the chitin externally. At first there is only a
fibrous junction with the ventral piece, which soon expands laterally
and dorsally, and it is not till some way down that the fusion takes
place and the canal assumes its complete shape. In the region of the
base of the fork the canal has developed a ventral projection (fig. 28),
and this goes on developing until the appearance of a row -lock is
obtained, as seen in fig. 29, which again disappears in the bulb.

The axial apophysis expands posteriorly into a rounded mass seen
in fig. 25, and by this level the original lateral internal walls of the
labella have completely disappeared, the hind portion of the apophysis
(<7) now' lying free, imbedded in soft tissues.

J7 4

The axial apophysis (,/ in all the figs.) is a somewhat pear
shaped median structure, the tip of which lies completely free between
the two inner walls (fig. 22) ; its middle portion is, as we have just said
fused on either side to the inner wall. Posteriorly again it expands
into a bilobed bulbous base which instead of being free is surrounded
>y soft tissues, it is, in fact, suspended by fibrous tissue to the under
sur ace of the dorsal gutter (figs. 23-25). It often shews evidence of

its bilateral origin in the existence of a median cleft (fig. 24) potential
or actual v s ^

f t! HE ^?RK ^ 111 aI1 t,le When we have reached the free end

th f L-Xm apophysi-s t,iere now appears on either side of it the tipsof
i a Ut transverse section and attached to the tips, we see also
p! °nS ^ 111 aI1 the fiSs0 which pull on these tips (fig. 22). As we
one -1 H ISd ' We l,ieSe chhinous arms of the fork approaching
(alLr r Ventra,ly (% 25)’ fina% firing, as shewn in fig. 26

sclerites (fig ®V®I'S,on)- • ^ baSC °f tWs f°rk rCStS °n tW0 ventra!
only internal ,'u r m transverse section still nearer the base the
and the vent 1 I?°^S structures are those forming the dorsal canal

fromtL™ To Th" In thiS SeCtion ^ see stretch,.

laterally we see ‘th* * ie ventraI sclerites powerful muscles (r1), and

the incurving and thin" 1™ ^ ^ aCr°SS’ We ma7 note here- 3,501
this point. The vei C aracter the chitin of the lateral wall at
replaced by various o'! SC^e”tes now decrease in size, the muscles are
practically assumed th ^4 ^IOUps °f muscles and the dorsal canal has
course. ‘C S lape !t ,las for a great part of the rest of its

dorsally (fig 28, a). \y epip,laiTnx has appeared, closing in the canal
hi detail. ni<17 now consider some of these parts more

The Epidermis f0

2> 4. 5) presents a fin!]".1'"^ th<? external walls of the labella (figs,
squamae, which struct,! rCtl^ulfted surface due to small’ polygonal
m tegument during the ~ .ltS °f considerable folding of the
,he labella are dosed °f the Iabella (%s- 10-14). When

orsally (fig 4); the* ° eep oblique, bilateral folds are seen
errninatmg at or near the tin commencing near the apex and
behind the lateral group IfT ^ ** 1 the second M commencing
P °f hairs infra). Ventrall y there are no

i75

folds in front of the fork, but there is a deep oblique fold immediately
behind it (fig. 5, c 7). When the labella are everted (figs. 2, 3, 10, II,
12, 14) the folds become very complicated and difficult of interpreta
tion, but there are three deep ones in front of the pulley. Dorsally.
a little posterior to the point where the tip of the fork impinge*
against the inner surface of the external wall (fig. 14, e), a complete
pocket or cul-de-sac is formed (fig. 13). Ventrally there is als -
another deep fold which, when seen in transverse section (fig. 12, c
appears as a projecting and slightly divergent arm.

External Hairs. These occupy three principal areas -.

(1) A ventral series ( d in all the figs.) immediately within the
thin flexible terminal flanges of the labella (figs. 2, 3, 5, 6, 9, &c.). Then-
are from 15 to 16 of these on either side, three of which are much
longer than the rest; all are stout, curved, rest on broad cylindrical
bases, and are equidistant, with the exception of two or three, which
give the otherwise complete line a somewhat irregular appearance
1 hese hairs are easily traced in' all preparations ; and during eversion
they are directed backwards instead of forwards, forming an almost
complete fringe below the teeth. Lying folded back on the proboscis
the tips of the largest reach nearly as far backwards as the fork

(2) A dorsal bilateral row of five or six setae ( [dl in all the figs

1 hese are situated partly within the apical fold of the labella (fig. 4 .
and when the labella are everted they take a more or less transverse
position, with the tips of either row overlapping or meeting along
the median line of the groove of the labella, forming a ladder-like
arrangement (figs. 2, 14). These hairs can be seen only in preparations
that have been thoroughly bleached.

(3) A lateral group of minute hairs (d2 in all the figs.). These
lie immediately behind the tip of the fork when the labella are
t'crted (figs. 2, 14), occupying a sub-dorsal position; but when the
a ella are closed, this group takes a more lateral position (figs. 4, 6)
m t le region opposite the sclerites which form the walls of the dorsal
canal of the labium. The individual hairs are extremely minute (the
ength less than the width of the clear area in which they are placed'
and difficult to see ; each one is placed in the centre of a large, clear
circular or gland-like ” area, which is well marked off from the
slH rounding chitin, and thereby rendered conspicuous.

176

Ventral Flanges or Lips ok the Labella ( c 2 mall the
figs ). These are extremely thin and flexible but are strengthened by
several rows of strongly chitiniscd scales, of which the external row is
much the larger. When the labella are closed these flanges or lips
meet together and completely enclose the .nternal mouth-parts
(fig. iq) ; but when eversion takes place they occupy a dorso-lateral
position, and the edges are then projected upwards and dorsalwanh

(figs. 2. 3, jo, 1 1 ), resting practically on the sub-lying chitin of the
external wall.

The Fork (e in all the figs.). A proboscis mounted in thi
dorso-ventra! position, as in fig. 5, shows at the base of the cleft,
solid clntinous median structure, the axial apophysis (q), and somera
e "n t^1’s a L -shaped piece of chitin, the fork, with its concavit)
pening forward. With this, are seen articulating two lateral sclerite
(figs- 5. 6, el) and posteriorly two ventral sclerites (figs. 5, 6, /). Thf

f • ' ct')<-en the base of the fork and these ventral sclerites i;
I ringed with rows of hairs (fig. 6, fl).

• ^P'Ca^ dl° Pork’ wkick is. as we shall see, an internal structure,
extern ^ ^anges (%• 5). which are continuous with the

sclerite • Lalerall>" the fork articulates with the two lateral

direedv^er S r SeCti°n in ** 25- A These lateral sclerites are

this cormeebor^m/b"0'0 ^ ^ chhinoUS integument, so that by
fork is enabled to n ° f.t,,e d,rect continuity at its tips, the

by the powerful °i U< ° evers,on of the labella when acted upon
the arms of the ^ ° the bu,b- Stretching across between

is first seen when the hind * hT. °f f'bv°US tissue (f,g 25> This
band serves to strenp+1 T ° ^ aX,aI aP°PhXsis disappears. This
to the fibrous tic«s, ^ armS ol t,ie fork- and by its attachments

Peg itself when the T the peg" ,les must exercise traction on the

Position of the hh 11 ^ ta<eS the verticaI position. In the everted
and this organ is S0C dlat we i°se this appearance of a fork.

1 ids change of nositi' ° Stretc^1I'n& across almost as a band (fig. 3-4
dorsally and nosti- ■ °?’ S la^ Sce’ ,s Pr°duced by the tips moving

section the *> that now in a transverse

transversely (figS -,r *S *cn£tk (bg- 12) instead of being cut

be seen (fig. 6s) whi’,~2) n cIeared specimens also the tendons can
* Ch attached “> fork a, its tips (fig. 3. 4

177

INTERNAL ORGANS OF THE LABELLA

These consist of seven different organs and are bilaterally
symmetrical (fig. 8).

(1) The Teeth (j in all the figs.). There are five of these and a
small accessory or rudimentary' one. They are relatively very large,
the length of an individual tooth being equal to about one-fourth the
entire depth of the labellum. They are articulated to an extremely
flexible, narrow plate, which is in turn articulated to the true plate of
the labellum. The first (proximal) and second tooth have each a
strong lateral process (tooth) on the lateral proximal margin,
and the tips of each are serrated on both sides ; the
three succeeding teeth are similar in structure, but the lateral
deep dentitions are now on the opposite sides. In the
everted position the te£th are freely exposed, widely divergent, and
project at right angles and even backwards and dorsalwards. It is
evident from their position and extent of motion that they act as the
chief and, in fact, the only cutting organs of the proboscis.

(2) I HE PETIOLATED Blades (/ in all the figs.). These are
long, curved, blade-like organs covered with minute hairs on the
ventral concave surface. All are attached to long and extremely
slender stalks arising from strongly chitinised projections of the
narrow tooth plate. They arc grouped in the following order : At
the proximal extremity, exterior to the tooth, are two groups ; the first
consists of two or three very small blades attached to a relatively
strong branched stalk ; the second group consists of seven or eight
blades, some of which arc scarcely larger than those of the first set ;
the main stem in also stout, but the lateral stalks carrying the blades
are extremely slender. Between each tooth is a set of four blades
auanged in two pairs, the blades of each pair being widely separated,
so that they resemble, somewhat, the arms of a lyre ; the blades
partly overlap, but the tips of the lower pair extend considerably
beyond the upper pair, and when the labella are closed they reach as
far as the basal row of external squamae fonning the extreme edge of
the labella (fig. 10, in dotted lines). Anterior to the last (fifth) tooth
aie three more sets of blades ; the first set (between the fifth and the
accessory tooth) consists of six or seven blades ; the second of a
similar number, and the third of four or five, of which three resemble

1 78

those carried on the first proximal stalk The stalk to which the
blades are attached, more especially those between the teeth, are, as
we have already stated, extremely slender They are, in fact, scarcely
thicker than the fine marginal hairs of the labella, are filamentous in
character, and in stained preparations present a well-marked granular
appearance. The relative position of these organs in the everted
sections may be seen in figs. 2, 3, j<j, 1 1.

(3) ROD-LIKE Hairs (ni in all the figs.). These are dark in
colour, in fact almost black ; they arc* long, more or less curved
bluntly rounded at the tips, and apparently grooved on one side
1 hey are articulated to strong cylindrical bases similar to those of the
ventral external hairs, are fixed in a semi-erect position, and are
arranged in the following order: One opposite the tip of the firs',
(proximal) tooth, a pair opposite each of the second, third, fourth and
fifth tooth, and one opposite the accessory tooth, making ten in all
S ^1<?se Q1gans are inserted on nucleated cells they are probably
sensory in function. The position of these organs, when the labella
are everted, is shown in figs. 2, 3, 10, 1 1

ihe Operculate Hairs (n in all the figs.). These at
lute cylindrical organs with suddenly truncate extremities, froi
ie edge of which there arises a flat oval-shaped lid or flap, whic

five of T angle °f 45° 0r more froni ‘he margin There a.
hrp-e ip f 1 Se cunous or£ans» one opposite the tip of each of the fiv
int~/ K y f°rm 3 ***** curved line and are fixed to th

of the petiolL^bfedeT3 ThetWeen ^ endS °f teC'h ““ '‘P

labella is shown in fig , , ^ P°S‘tlon of these O'S™* in the evertel

lon&5lwolto,ECiiTEi? HA'RS in a" lhe figs). These are rathe
branched portion^ be’ ^ d'V'ded a little beyond the middle, th'
marked and al extremely slender. They form six well

operculate hairs "earl C°nt,nu®us ;&rouPs« slightly in advance of the
rows (fig. 8). "phe r ' ^‘r >U^ iav,nf? a tendency to arrange itself intc
(figs. 10, 11') i,lcf 1 \ J>c< UP> ,l lateral position in the everted section?

(6) The A 16 Ventml ***** of labella.

These hairs clothTfte FR™GE °F Hairs O’ in a11 the fip)

of the labellum (firr q\ " °f t,,e extreme anterior dorsal margin

1 hls fnnge is broad basally, but gradually

*79

narrows as it approaches the ventral margin. In everted sections
(figs, io, ii) these hairs project dorsally into a pseudo-cavity formed
by the over-arching wall of the epidermis; and when thus in
apposition they shut off the cavity of the labial trough.

(7) Proximal Fringe of Hairs (/>’ in all the figs.). These
are similar to the former, but occupy a relatively small space on the
proximal margin.

LABRUM digs. 14-19)

This commences immediately behind the fork. Dorsally and
laterally for its whole length it is covered with spinose hairs directed
apically. At the apex itself (fig. 16) the hairs are replaced by scales.
Near the tip, the labrum has ventrally a ridge or chitinous band
extending across it (fig. 19), on which are situated four hairs evidently
sensory in function, as nerve endings can be traced into them.
Behind this point the sides widen slightly and along the interior are
here arranged in rows, a series of hairs ( a 2) on tubercles.

I he labrum is at first concavo-convex in section, the convexity
near the tip forming a kind of ridge (fig. 1/). A little further down
it has the appearance of an U fitted into a V (fig. 18). In the cavity
between the two there appear early soft structures which later become
developed into well-marked muscular bands connecting the lateral
walls of the labrum (figs. 29, 30, j4). The sides of the labrum
become soon expanded into lateral flaps, which serve the purpose of
keeping the hypopharynx in its place, the flaps fitting on to the
expanded sides of the latter (figs. 29, 30). Eventually the ventral
wall of the labrum becomes closed ventrally, forming now a
chitinous tube, while the dorsal wall becomes fused and lost in the
substance of the bulb (fig. 31).

THE HYPOPHARYNX (b in all the figs.)

This organ commences at about the same point as the labrum.
ft is a much smaller and weaker tube. At the extreme tip it appears
to be membranous in character and to retain no very definite
constant shape in preserved material. (We would suggest that this
flaccid condition of the extreme tip is to prevent ingress of fluid, e g.,
blood during feeding, while permitting freely of egress of saliva.) It
L

occupies the dorsal gutter of the labium, and is kept in this position
by the pressure of the lateral arms of the labruin which stretch over
on either side for a short distance (figs. 2Q, 30).

In section near the tip it appears as a chi ti nous hoop, formula
circle complete for about seven eighths of its circumference. At firs
the hoop has near the apex on each side a little tubercle, and stretching
across between these tubercles is a thinner chitinous membrane
completing the tube dorsally. As we get nearer the base there is a
distinct separation in the walls of the chitinous ring in the position
where the tubercles first appear, so that a cavity now exists in the
sides (fig. 29). This goes on increasing till eventually we have the
chitinous circular inner tube lying within a sausage-shaped outer
covering, forming a projecting angle on either side (fig. 30, £).

At its base the ventral wall of the hypopharynx becomes fused

with the bottom of the labial gutter. The beginning of this is shown

m 3°' This proceeds until the process is complete on either side

I he fused wall is then absorbed, and thus permits of the inner

(| Oropharyngeal) tube escaping through ventrally. At the same time

! e dorsal wall of the hypopharynx becomes fused with the ventral

-alf of the epipharynx, forming now the pharyngeal tube (fig. 3')

", If *1 !\d°rsal Iialf of llie epipharynx fuses laterally with the sides

3 ,um’ so ^lat ^ie whole of the epipharynx and hypopharynx

• excePti°n of the dorsal wall of the former, have now become

internal structures.

MUSCULAR SYSTEM (s, i„ all the figs.)

two lame c Pro^osc*s it is often possible to separate out

These htto * aratc lllasses °f muscle which terminate in tendons (f.

*»e b?hl“d ;° the t,p °f t,ip ^ ^ - » i

the proboscis H - 1 P ane untl1 we reach the bulbous portion of

they arise, whi 1 ^ ^ tendons &ive Place to the muscles from which
the whole of th COnsist °f two masses filling posteriorly practically
describe as the 1 Cntra^ Portion of the bulb. These muscles we i«a)
the median ***■> we have on either sid>

forming a broad 1 ° j^e ^utter a group of oblique muscles (•''

inserted into tlie '”]•’ 'U'S'n^ from the chitinous integument and

separated from it !'U' limb ^ 29)- Dorsal to this group. nIlri
y the tendon, there is a second oblique group (*J)

converging from the outer wall to the base of the gutter. Both these
groups can be traced as far forwards as about the base of the ventral
sclerites. Anterior to this point we have a well-developed band of
dorso-Iateral muscles stretching from the canal dorsally to the ventral
sclerites ventrally (fig. 28, j1).

Muscles of the Labrum. We have also sub-median muscles
present in the cavity of the labrum stretching from the outer to the
inner walls (figs. 29, 30, r4).

Internal Laminae (fig. 28, s5). In sections in the labella
region, where the dorsal fold is well marked, there appear beneath the
fold of the integument peculiar yellowish or brownish granular bands,
of irregular appearance following the outline of the integument. They
often show little irregular lateral projections. They represent transverse
sections of plates. They are seen in sections as far as the ventral
sclerites. They are, moreover, very well developed also in the cavity
of the labrum in its basal portion. We are unable to suggest what
may be the function of these bodies.

MECHANISM OF THE PROBOSCIS

With regard to the mode of action of these muscles, we consider
that the function of the longitudinal group is to erect the fork and,
by so doing, to produce by the traction on the external chitinous wall,
eversion of the labella and a resulting spread-out condition of the
teeth.

I he oblique muscles' function is, we believe, to produce a semi¬
rotatory movement of the proboscis3 (and labella), so that the teeth
are enabled to exert a cutting action on the skin, the motion being a
to and fro one.

The cutting weapon of the proboscis is undoubtedly the teeth.
These are capable of a variable extent of eversion, and while they arc
m act'on die other structures anterior to them, petiolated blades,
ror -like hairs, etc., are folded back in a plane below the actual tips of
die teeth, and so escape damage.

I he tip of the proboscis then can only cut in the everted position.
" c have in this condition of everted teeth an admirable weapon for
boring a hole into the skin. The arrangement of a carpenter’s augur
"ith its cutting flanges on either side is almost an exact parallel, though
the motion here is a continuous rotatory one, and not a to and fro

A

EXPLANATION OF REFERENCE LETTERS

External Organs.

<i Labrum-epiplmrynx.
ax Ventral ridge of labruni
bearing sensory hairs.

<i* Lateral ventral hairs to
labruni.

aa Apical scales to labruni.

</' Dorsal hairs to labrum.

b Hypopharynx.

c Labium.

c1 Labella.

c* Ventral flange or lip of labella.
t:1 Epidermis of labella.

c‘ hirst dorsal groove in epi¬

dermis of labella.

cb Second dorsal groove in epi¬

dermis of labella.

c° Third dorsal groove in epi¬

dermis of labella (produced
in eversion).

i 7 Ventral groove in epidermis

of labella.

cH Fold forming cul-de-sac.
c® Fold behind fork.

d Ventral external hairs.

dx Dorsal external hairs.

d1 Lateral group of external

hairs.
c Fork.

cx Lateral sclerite to fork.

/ Ventral sclerites.

Jx Hairs at apex of ventral

sclerites.

Palpi.

Apodeme of labrum.

Internal Organs.

i — Tracheal sac.
t1 — Tracheal tube.
j The teeth.
j1 —Accessory tooth.
k -Tooth plate.

/ — Petiolated blades.
m Rod-like hairs.
n Operculate hairs.

o Bifurcated hairs.

p Anterior fringe of hairs.
px Proximal fringe of hairs.

(j The axial apophysis.
r -Dorsal hooked sclerites form
ing upper portion of the
labial gutter.

r1 Sclerite forming base of labial
gutter.

s —Tendon.

s1 Dorso-ventral muscles.
sa —Median oblique muscles.
s8 Ventral oblique muscles.

i4 Transverse muscles in cavity
of labrum.
s6 Internal lamina.
t Chitin cells.
u — Interlocking spines.

S

h

i84

explanation of PLATE XII

E^- 1‘ External mouth parts of Stomoxys calcitrans seen from the
side, x 36, about, a , labrum ; a1, ventral ridge of labrum
bearing sensory hairs; a1, dorsal hairs ; b, hypopharynx;
c, labium; c1, labella ; g, palpi; h, apodeme of labrum;
i, large tracheal tube or sac.

F lg- 2— Labella everted, dorsal aspect. The left half shows the
characteristic scale-like structure of the epidermis. x200-

F,g- 3- Labella everted, ventral aspect. Details of epidermis omitted,
x 200.

big. 4— Labella closed, dorsal aspect, x 200.

Fig. 5.— Labella closed, ventral aspect, x 200.

Plate XII

F'g- 6
Fig- 7

Fig. 8.
Fig- 9-

explanation of PLATE XIII

Plan of labella (closed) as seen from the side, showing the
relative position of the internal and external organs, x 200.

Internal organs of right labelluni and distal portion of labial
trough (r, rx ) as seen in optical section after dissection.
x 250, about.

Fhe same x 600, about. Sclerites forming labial trough

omitted.

Ventral external fringe or lip of the labella (left half) x 5 7a

DORSAL

Plate Xiri

ANTERIOR—

-J

PROXIMAi

-PS

Fig. 6.

DORSAL

VENTRAL

Fig. 9.

West, Newmaji lifch.

Fig. 8.

1 88

EXPLANATION OF PLATE XIV

Fig. io. Section through everted labella in the plane where the tip
of the axial apophysis (q) first appears. The approximate
extent of the petiolated blades is shown in dotted lines.
x 570.

Fig. 1 1.- Section through everted labella in the plane where the base
of the axial apophysis (q) becomes fused to the tooth-pb^
(£) and where the labial gutter (V1) and the dorsal hooked
sclerites (/•) first appear. x 570.

PLATE XIV

Fig. 10

' .* •>>>.

V. •

* : * • T •

«,• .. . .v

ft K j&£l ^

•■/£* r'; •'.• ■- • • • : V •/'

f«^. v

EXPLANATION OF PLATE XV

Fig. 12. — Section through everted labella in the plane of the fork (e)
where the dorsal and ventral folds of the epidermis (d r)
are well marked. In the next proximal section the tips of
the fork are seen to extend into the angle of the third
fold (r6), here indicated by dotted lines. Note that the
tooth plate has entirely disappeared and the labial gutter
0'< r *) is now well formed, though the upper sclerites are
still free. The approximate plane of this section is between
c 6 and e in fig. 14. x 570.

*3- Pocket formed by the infolding of the epidermis in the
plane near d 2 in fig. 14. Here the upper sclerites (r) of
the labial gutter have become fused with the lower (r1).
x 570.

^ '£• !4- -Dorsal aspect of everted labella and distal portion of labium,
from a well cleared preparation. Compare this with fig 2,
which was drawn from a preparation which had not been
sufficiently cleared.

P'g- 15 Distal portion of labrum. x 200.

Fig. 16. -Extreme tip ol same showing the scaly dorsal structure,
x 570.

^ Section through tip of labrum. x 570.

1 ig- 18. Section of labrum through the plane just in front of the
ventral thickening (fig. 15, a1) carrying minute hairs,
x 750.

19 Section of labrum through ventral thickening (fig !5>
showing minute hairs. x 750.

EXPLANATION OF PLATE XVI

Transverse section through the tip of the closed labella
x 600.

Section through the closed labella at the plane where the
teeth (f) have almost disappeared, and where the sclerite
forming the labial gutter (V1) and the tips of the fork (e)
first appear, x 750.

Fig 20

Fig 21

I

, _

194

EXPLANATION OF PLATE XVII

Fig. 22. Section through the closed labella in the plane where the
free end of the axial apophysis (q) first appears. In this
section the teeth have disappeared and the labella is
closed ventrally. x 750.

F'g- 23- Section showing the fusion of the axial apophysis (q) with
the tooth plate (k). x 7150.

Fig. 24.— Another phase of the same. x 750.

I ig. 25. Section through the closed labella in the plane at the base
of the axial apophysis (7). Here the tooth plate has
entirely disappeared ; the arms of the fork are seen con¬
verging, and stretching from tip to tip is a broad
membrane ; here also are seen the lateral sclerities of the
fork (<d) articulating with that organ, x 750.

PLATE XVII

M

M

196

EXPLANATION OF PLATE XVIII

1 ig. 2(>. Section through closed labella at the junction of the fork (i)
with the ventral sclerities (/). x 750.

1 ig. 27. Section through the labium in the plane where the ventral
sclerites ( f ) first appear. The fine hairs at the apex of
these organs are here shown, x 750.

I ig. 28. Section through the labium at the plane where the labruni
(<7) first appears. Here muscular bands stretch across from
the ventral sclerites to the walls of the labial gutter, x 150.

PLATE XVIII

27

28

■ NrwUraJ, ad. Hal. del.

i

I

EXPLANATION OF PLATE XIX

I-ig. 29.— Section approximately through the middle plane of the
proboscis, in which are seen the labrum (a), the
hypopharynx ( b ) completely closed and interlocked with the
labium. Here also the row-lock like appearance of the
labial gutter is well shown and the rod-like sclerites
have completely disappeared, x 750.

F'g- 30.— Section through the bulb of the proboscis. Here the
hypopharynx is seen fusing with the walls of the labial
g-utter , and the large air sacs (i) have taken the place of
the tracheal tubes (fig. 29, i\ etc.), x ;5o.

S 3 • Section through the base of the bulbous portion of the
proboscis, including the apodemes (h) of the labrum. Here

*e wa^ the hypopharynx has become fused with

a rum, forming the pharyngeal tube, x 200.

PLATE XIX

TRYPANOSOME TRANSMISSION
EXPERIMENTS

201

TRYPANOSOME TRANSMISSION
EXPERIMENTS

BY

The late J. EVERETT DUTTON, M.B. (Vict.)

(WALTER MYERS FELLOW, THE UNIVERSITY, LIVERPOOL)

JOHN L. TODD, B.A., M.D., C.M. McGill

(DIRECTOR OK THE RUNCORN RESEARCH LABORATORIES OF THE
LIVERPOOL SCHOOL OF TROPICAL MEDICINE)

AND

J. W. B. HANINGTON, M.D., McGill

(DEMONSTRATOR AND RESEARCH ASSISTANT, LIVERPOOL SCHOOL OF
TROPICAL MEDICINE)

Being the Seventh Progress Report of the Expedition of the Liverpool
School of Tropical Medicine to the Congo, 1903-1905

( Received April 20th, 1907)

I. ATTEMPTS TO TRANSMIT TRYPANOSOMES BY
TSETSE FLIES

In a short series of experiments done in the Gambia in 1902-3, 1
we failed to infect animals with Trypanosoma gambiense or
dimorphon by the bites of captive flies (Glossina palpahs and
Stomoxys (see h page 40). which were either freshly caught or had
been fed previously on an infected animal The members of the
Royal Society’s Commission on Sleeping Sickness in Uganda ha\e
since succeeded, however, in infecting healthy animals wit 1
trypanosoma gambiense by the bites of flies (6 lossina palpalis ) e >
at the most, 48 hours previously on an infected animal (-, page i>
As a result of this work it has again been maintained (13, an
others) that the tsetse fly is merely a mechanical transmitter o
trypanosomes, and is not an alternative host in which the parasite
undergoes definite developmental processes.*

__ *TJ?‘S suggestion first emanated from Bruce, but he J^^m^tTryfartosoina
possibility of there being a developmental process of the trypanosom P

0rucet) >o the tsetse fly ( Glossina morsitans) (14, page 4)-

202

When our expedition readied the Congo, sleeping sickness was
endemic, and the existing conditions were much the same as in
Lganda. We therefore determined to repeat the work done there,
with the primary object of ascertaining the longest period after feeding
on an infected animal at which a tsetse fly, by its bite, is capable of
transmitting trypanosomes to a healthy animal. It must be
remembered that, like Bruce, we worked with adult flies caught in the
bush and that our experimental animals were only partially protected
from the chance bites of insects.

The conditions of our experiments were therefore imperfect. The
flies used should have been raised in the laboratory from the lame
of known parents. Wild flies, caught when adult in an area where
trypanosomiasis is endemic, may have fed when free on infected
animals; it is only in flies bred in the laboratory that the dates of all
infecting feeds can be accurately known. It is possible that the
piogeny of infected tsetse flies are capable, or are alone capable,3 of
transmitting tire trypanosomes ; these points can only be decided by
experiments made with laboratory-bred flies.*

\\ e do not think that the danger of chance infection of laboratory
animals, only/ paitially protected from the bites of insects, is so great
as might be imagined. None of our large stock of rabbits, guinea-
pigs, rats and monkeys were ever found to have been so infected during
33 months spent on the Gambia and on the Congo.

In the Congo two main series of experiments were made with
tsetse fires, one at Leopoldville and the other at Kasongo. A small
nei series of experiments (Experiments 128, 128a, 129 and part

... 'Vas ^onc u*th flies caught along the Congo between Leopold¬
ville and Coquilhatville.f

^ Leopoldville it was very difficult to obtain monkeys and we
1 , °|1Ce, l° USe £u’nea-pigs and rats in some of the experiments,

gi tiey are, as a rule, rather less susceptible to Trypanoswa
gam b tense .4 *

difficult to breed^theiu" °.jfS °! the bionomics of captive tsetse-flies (5), 11 15

flies used in our experiment* ; Yu1 a dozen of the pupae obtained frorn th®
flies. ' p nents ln the Congo developed in the laboratory into adult

distribution of^leeffin^Sifkllit aFea traversed by the Expedition and showing the
Liverpool Sc h ool ^oFt. r op i caHVfed i c i Free State, see Memoir XVIII of the

t .1 .1 wr used small gauze cages and found it difficult to

. i . :/m pal pahs) alive for more than a few days

\i Li >|)oldvillc we attempted to keep them longer by
r ' ai > I i mg their natural habitat in their cages. Cubical

.mi n ug 1 8 inches along each side and containing water
. njj ',\cre placed in a sheltered position out of doors.

■ ' •• ; mental animals were simply turned loose in these

i f , - '.ere found to feed much better on them if they
. t nder these circumstances flies often lived in
" 1 1 ! of months. 1 he method, however, had distinct
•, ,t ' as difficult to feed the flies on large animals and,

1 'i r 1 1 id contents of the cages, it was almost impossible
idler of living flies they contained.5 For this reason.

. •, ii ' done at Kasongo the smaller cages were again

f ;«■, ted ii the experimental animals daily.

r ' animals or men, whose blood contained many

.. . . were chosen for the infecting feeds. In the latter

., .. : . ,lt Kasongo, when it was found that so few of the

enta bad been successful, the flies were fed on
.-.r ... of sleeping sickness, in whose blood there were very

i - ntly no parasites, in order that flies should be
■r wiih V. inch had fed on persons at each stage of the
u, ic, ting hosts have been further described in the

of the successful experiments.

' n . neerning the infectivity of the parasites at various

TX rJ ' • disease can be drawn from an examination of t ie
Ml, Hies used in successful experiments, because
Md uninfected after having been fed upon by the
identical conditions.

, <1|l(l,t,ons permitted, tlie flies were used for several

, example, many of the ^ies use 1

• i infect a healthy animal by Hu es 0

ns previously on an animal tnfected wrth

^FMmu gambienae) had been used before in other e*per—
mjf u then bad therefore fed several times, u i
n pe on infected animals. As far as we ’

•i If ... vs (s used in the experiments made at eop

204

At Kasongo about a score of Glossina fit sea in all were caught. Most
of them were allowed to remain in the rages. Where it is necessary
experiments on which they fed are specified. Rare examples of
Glossina pal l id i pcs or Glass nut want lata may have been present,
undetected in the cages, at both localities.*

At Leopoldville Trypanosoma gantbienst was alone used; at
Kasongo Trypanosoma dimorphvn was also experimented with.

I. FRESHLY-CAUGHT FLIES

To infect susceptible animals by the bites of Glossina caught hi
localities where trypanosomiasis is endemic.

There was a great deal of sleeping sickness in the areas where
the flies were caught at both Leopoldville and Kasongo. Because of
the larger local population, the chances that a newly-caught fly had
previously fed on a person infected with trypanosomes were rather
greater at Leopoldville. The flies fed at Mswata and Coquilhatville
on Experiments 128, 128a, 129, 141 and 21 1 were caught in very
restricted areas where there were many people, the great majority of
whom were infected with Trypanosoma gambiense. At both
Leopoldville and Kasongo trypanosomiasis of animals was far from
rare. Because of the presence of a herd of 280 cattle (about 5 per
cent, infected) and of considerable game at Kasongo, the chances
that a fly caught there had previously fed on an infected animal were
much greater than at Leopoldville.

Leopoldville.

Experiment 45.— Guinea-pig. From December 8, 1903, to April 13, «9°4- 45|
ies were fed. I he animal's blood was examined regularly until October, 1904 ’
trypanosomes were never seen. It died on October j8, 1905, from other causes.
° tr-Vpanosomes or signs of trypanosomiasis were seen at the autopsy. 1

* ^ee (5) page 03.

nr thrift r°utine, the blood in all these experiments was examined, as a rule, tvvo
the hlmiH nf CS a, week' fresh in &-inch square coverslip preparations. In addin'1'
^ 0 eacp animal was occasionally centrifugalised (6).

waslwlh°Ugh U 'S n0t alway-s mentioned, in the following experiments an autopsy
and fresh eVery. CaSe s«e» withi» a few hours or minutes of the animal's death,
By Pr?a.rat,0ns °f body fluids and of organ juices were always examined

enlarge f At °f tryPanos°miasis " are meant in particular, enlarged spleen and
enlarged, often much congested or haemorrhagic, lymphatic glands

205

Experiment 40.— Guinea-pig. From December 8, 1903, to June 9, 1904, 552
flies were fed. The animal's blood was examined regularly until October, 1904 ;
it died on June 3, 1905, of pneumonia. No trypanosomes or signs of trypano¬
somiasis were seen at the autopsy.

River Series.

Experiment 128a. — Monkey ( Cercopitheeus schmidti).* On June 24, 25, 26,
1904, 25 flies caught at Mswata were fed on this animal. Its blood was examined
regularly until September, 19*14, when it died of dysentery ; trypanosomes were not
seen (malaria present |.

Experiment 129. — Monkey ( Cercopitheeus schmidti). I rom June 27 to July ,,

1904, 262 flies caught at Mswata and at the sleeping sickness hospital at Coquilhat-
ville were fed on this animal. Its blood was regularly examined, but trypanosomes
were never seen ; its temperature was always normal. It died (cause .-’) on July 9,
1904; no trypanosomes or signs of trypanosomiasis were seen at an autopsy done
four hours after death, and a rat sub-inoculated with its blood never became
infected.

Experiment 141. — Monkey ( Cercofitheeus schmidti ). 1-rom July 15 to July 1 / ,

1905, 52 flies collected at the Coquilhatville sleeping sickness hospital fed on this
animal. Its blood was regularly’ examined until October 14, 1905, when it was
stolen. Trypanosomes were never seen in its blood, and its temperature was never
abnormal.

Kasongo.

Experiment 182.— Monkey ( Cercopitheeus schmidti). !• rom January 19 to
lebruary 7, 1905, 211 Hies fed on this animal. Its blood was frequently examined
until July 24, 1905, when it was accidentally killed. No trypanosomes were ever
seen (malaria present) during life, and no trypanosomes or signs of trypanosomiasis
were present at the autopsy done three hours after death.

Experiment 198.— Monkey ( Cercofitheeus schmidti). From February S to April
'8, 1905, 2,659 Hies fed on this animal. Trypanosomes, probably Trypanosoma
gambienst , were first seen in its blood on May 13, 1905. It gradually lost weig it
and died of the disease in November, 1905. In March, two G/ossinae fuscae fed
on this animal.

Experiment 203.— Monkey ( Cercopitheeus schmidti). From February 22 to
March 15, 1905, ,i7g9 fljes fecj on this animal. It was accidentally killed on March
'6. Its blood was regularly examined ; no trypanosomes were seen during life 01
at the autopsy.

Experiment 213.— Monkey ( Cercopitheeus schmidti). From March 20 to April
J0- *905. 7*7 flies were fed. The blood was carefully examined up to July 5, when
‘he animal was killed by its cage companions. No trypanosomes were seen during
llfe °r at the autopsy ; nor were there any signs of trypanosomiasis. Two puppies
sub-inoculated at the autopsy with respectively 3 and 5c.cm. of blood never
became infected.

easihKa<h sptcles of monkey used in these experiments has been show n ^
325*5*4 by Trypanosoma gambiense bs the inoculation of blood or cerebro
Plnal fluid i*om sleeping sickness patients.

Successful

Experiment

206

II. INTERRUPTED FEEDING

1 o infect susceptible animals with Trypanosoma gambiense
by the bites of tsetse flies fed immediately previously on an infected animi
or man.

Leopoldville.

Successful Experiment 63. — Guinea-pig. This animal was immobilized and placed

xpenment together with an infected guinea-pig in a rage containing tsetse-flies. From
December 24, 1903, to January 1 5, 1904, 219 Hies fed on one or both animals
I rypanosomes appeared in the blood of this animal on January 15, 1904, and it
died of the disease (and other complications) on March 16, 1904. The morphology
and animal reactions of the parasites agreed with Trypanosome gambiense.

I he flies used in this experiment were infected by feeding on guinea-pigs
inoculated directly from an early case of trypanosomiasis. Trypanosomes were
very numerous in their peripheral blood, and they had been infected for about
two months.

Experiment 76. Monkey ( Colobus sat, mas). Flies in a small cage were

a owed to partially fill themselves with blood from a “ sleeping sickness” patient,
.me were then permitted to finish their feed on this monkey. From January 2 to
prtl r 2 , 1904, when the monkey died of sunstroke, r^7 flies were fed in fe
manner. Trypanosomes were never seen in the monkey’s blood and its temperature
remained normal.

. *'X1 1 ^7-— Guinea-pig. As in Experiment 63, this animal was placed

1 i an in ected one in a cage containing tsetse-flies. From January 23 to March
1 372 "Ies were fed- Its blood was examined regularly until June 7, and

rom tun at intervals until it died of exposure in October, 1905. Trypanosomes
ne\er seen ; there were no signs of trypanosomiasis at the autopsy.

Kasongo.

•xperiment 209.— Monkey [Cercofithecus Schmidt,).

•s rnmnlotolir . j .1 . ...

iX inor o j m nrruuti ). 1'lUlll iUclM-11 j w —j~

having 3 S ,COmIjle,ely filled themselves with blood from this animal aft.
blood wa!mmencei immediately previously to feed on an infected animal, f
on \nril e*annne daily, but trypanosomes were never seen. The animal die
were feause, ? starvation). No trypanosomes or signs of trypanosomias

were seen at the autopsy done immediately after death.

III. THREE TO FIVE HOURS INTERVAL

7o infect a susceptible animal with Trypanosoma gambiense by
hits of tselsi J lies fed from 3 to 5 hours previously on an infected

Very few flies were used in these experiments, since it was
xcee mgly difficult to get them to feed at such a short interval-

Mar^^o^qoe1 J°flirM°nke? \CercoPith“** schmidti). From February 24 <°
result (malaria i re - * were ed- Hlood was examined for trypanosomes without

trypaniomw orKl o t"nt a"imaI dled °f dysentery on Apr,. ,7- *

or signs of trypanosomiasis were seen at the autopsy.

207

Experiment 231.— Monkey ( Cereofithecus schmidti). On April 24, 25, 26, 1905,
22 flies were fed. The blood was regularly examined up to November, 1905.
Trypanosomes were never seen. The animal died of dysentery on l ebruary 21,
1906 ; no trypanosomes or signs of trypanosomiasis were seen at the autopsy.

IV. EIGHT TO TWELVE HOURS INTERVAL

To infect a susceptible animal with trypanosomes by the bites of tsetse
flies fed at least 8 to 12 hours previously on an infected man or animal.

Leopoldville.

Trypanosoma gambiense

Experiment 88. — Monkey ( Cereofithecus schmidti). Prom January 27 to May
28, 1904, 634 flies, previously fed on a sleeping sickness patient, fed on this animal.
Its blood was examined frequently until March 15, 1904, and then at inter\als
until it was killed by rats on March 30, 1905. Its temperature was always normal
and trypanosomes were never seen in its blood.

River Series.

Experiment 128. Monkey ( Cereofithecus schmidti). Prom June 24 to July 22,
1904, 87 flies, previously fed on a monkey infected with 7 ry panosoma gam lense,
were allowed to feed on this animal. Its blood was examined at intervals until
September 28, 1904; trypanosomes were never seen. I he temperature-c art was
not characteristic of trypanosomiasis (malaria present).

Kasongo.

(a) Trypanosoma gambiense

Experiment 187.— Monkey ( Cereofithecus sp. ?)■ P rom January 28 to April 10,
'^S. 939 flies, previously fed on a sleeping sickness patient, fed on this amma .
Its blood was examined almost daily until May 5, 1905, when it was sto en ,
trypanosomes were never seen.

Experiment 197. Monkey (.1 Cereofithecus man a). From February 9 to Apri
2°. >9°5, 943 flies, previously fed on infected guinea-pigs, were fed on this anuna .
Its blood was examined regularly until it died on July 6 (on river steamer) iroi
exposure. Trypanosomes were never seen (malaria present) during life, or at e
autopsy done immediately after death.

(b) Trypanosoma dimorphon

Experiment 216.— Monkey (Cercopitlucus mana). From March 25 to April 25 >
'9°5, 388 flies were fed. Blood was regularly examined until June 13, "

^mal was moribund. It was therefore killed. Trypanosomes were never see
curing life, and no signs of trypanosomiasis were seen at the autopsy. a

df*th ? exposure).*

lb* ^r'nS the journey of the expedition, on foot, from Kasongo
monkeys unavoidably received considerable hard usage.

to Lusambo (5),

2 o8

V. TWELVE TO FIFTEEN HOURS INTERVAL

To infect a susceptible animal with Trypanosoma gambiense by
the bites of tsetse flies fed at least 1 2 to 15 hours previously on an infected
animal or man.

Leopoldville.

Experiment 48.— Guinea-pig. From December 12, 1903, to April 6, 1904, 1,706
flies were fed. The blood was regularly examined until April 12, 1904, when the
animal died, trypanosomes were never seen in its blood nor in any of the body-
Hinds or organ-juices examined at the autopsy, done immediately after death.

1 .xPEKiMENi 6i. Rat. l-roni December 23, 1903, to May 20, 1904,4846165
were fed on this animal. Its blood was regularly examined' until July 3, .904,
v en 1 was accidentally killed. Trypanosomes were not seen.

I. xi iI'.kimi m (>2. Rat. I' rorn December 23, 1903, to February 25, 1904,261
les led on this animal. No trypanosomes were ever seen in its blood. It died
on March 1, 1904, of a lung disease.

Experiment 71.— Rat. From December 18, 1903, until May i5, 1904, 4>7

weit. ec cm this animal. Its blood was regularly examined for trypanosomes
without result until July 7, ,904. when it was accidentally killed.

In the four preceding experiments it was attempted to transm
the trypanosomes from animal to animal; in tiie three following
was attempted to transmit them from man to animal.

this animai ^ V * rom January 16 to April 9, 1904, 999 flies were fed f

result until Ini * ’ "tf re6uIarl3' examined for trypanosomes with negati’
result until July ,, ,904, when it Was accidenta],y

fed on'^his'ammal '' l7s' blood'1”8' Kr0!" ApriI '3 to May 3°. *9°4. 39° fiies wel

at intervals until October 8 wh^ eXa""ned reSuIarl-v until June 1, and afterwarc

aniinarreacdon1o^a,Congo,,cattle'/trypanosonie?X^er'nientR “*

Kasongo.

'905, 542 fl ies,N preyfousk^fecT on ^ SP- ?>- From January 24 to April 5
Its blood was examined f a ■•’leeping sickness patient, fed upon this animal

were never .seen fieqUent,F ,,nt" ”as stolen on May 4 : trypanosomes

April 20, 1905, 46l1iesM°n"k:?' -i erct°^ithecus schmidti). From February 25 to
monkey. ()n April 2c it °''s -v on an infected animal, were fed on this
therefore killed. Nntrvn^!^ ad,-v injured by its cage companions, and was
autopsy, nor were there any’ X'n * durinS Iife <maIaria presenti, or at the

y s,8ns nf trypanosomiasis.

2og

VI. FIFTEEN TO TWENTY HOURS INTERVAL

To infect a susceptible animal with Trypanosoma gambiense by
the bites of tsetse flies fed from 15 to 20 hours previously on an infected
animal.

Leopoldville.

Experiment 97.— Guinea-pig- From February 20 to May 16, 1904, 395 flies
were fed. Its blood was regularly examined for trypanosomes without success
until May 16, when it died from a ruptured stomach. No trypanosomes or signs
of trypanosomiasis were seen at the autopsy.

VII. EIGHTEEN TO TWENTY-FOUR HOURS INTERVAL

To infect a susceptible animal with Trypanosoma dimorphon
by the bites of tsetse flics fed at least IS to 24 hours previously on an
infected animal.

Kasongo.

Experiment 212.- Domestic cat. From March 13 to April 29, I9°5’ 5'0 ^
fed. Blood was examined regularly until July 26, 1905. trypanosomes
seen. When examinations were recommenced in September, 1905, r> P
were present. The animal died in November, 1906, of the disease. 11 ■ ' -

Glossina fusca fed on this animal ; a single Glossina palhdtpes also Fad an
opportunity of feeding. The flies used were infected by fee mg on re „

pigs inoculated about two mouths previously with blood from a sites

infected with Trypanosoma dimorphon ; they frequently ec w icn
were either absent or very scanty in the peripheral blood.

Successful

Experiment

VIII. TWENTY-FOUR HOURS INTERVAL

To infect a susceptible animal with Trypanosoma gambiense
by the bites of tsetse flics fed at least 24 hours previously on an inf
animal or man.

Kasongo.

Experiment 208— Monkey (Cercofilhecus schmidti). I' r0nLd ^'jhis monkey

^S. 590 flies, previously ted OH an infected animal, October 2, 1905,

On June. 16, trypanosomes were first seen in its blood. tie osomes were

from pneumonia. The spleen was considerably enlarge ■ . eding six

seen at the autopsy, and none had been present in the > 00 animal,

"■eeks. in March, 3 fusca had opportunities for feeing oh

A" lhe flies used in this experiment were infected by being e o ^ montbs
"hose blood contained very numerous trypanosomes, >»<,cl,la e' flies had

Previously with blood from a case of sleeping sickness. Many < contained

W three O, more davs previously on sloping sickness cases whose blood
'erv *ew or no trypanosomes.

Successful

Experiment

210

Experiment 218. — Monkey [Cercofithecus sckmidti). From April 3 to April 14,
I9°5» 7 1 previously fed on a sleeping sickness patient, were fed on this

monkey. The blood was examined at intervals until June 5; trypanosomes were
never seen. The animal was moribund on June 5; it was therefore killed and an
autopsy was made immediately. No trypanosomes or signs of trypanosomiasis
were observed.

IX. FOUR DAYS INTERVAL

lo infect a susceptible animal with Trypanosoma gambiense
by the bites of tsetse flies fed at least four days previously on a sleeping
sickness patient.

Leopoldville.

Experiment 74.— Guinea-pig. From December 30, 1903, to May 29, 1904,42
les were e on t is guinea-pig. Its blood was examined regularly until Octobei
904. w en 1 died of coccidiosis. No trypanosomes were ever seen.

ExpERrMi.Ni 122. Monkey' [Cercofithecus sckmidti). From April 24 to June 1
1904, ^o9 flies were fed on this monkey. Its blood was examined regularly unti
j6 W en 11 was accidentally killed. Its temperature had alwavs been normal
and at the autopsy, as during life, trypanosomes were not seen.

It was thought that perhaps the trypanosomes underwent a lengthy
developmental process in the tsetse fly, and that therefore the scanty
success of the preceeding experiments might have been due to the
shortness of the interval between the infecting and transmitting feeds.

n t le two following sets of experiments this period was consequently
greatly increased.

X. TEN DAYS INTERVAL

, lnJec^ a susceptible animal with Trypanosoma dimorphor

animal ^tes 11 ^ ^cas* ^en days previously on an infect a

Kasongo.

fed ; in many instant ^,n*'lca'P*S- 1‘rom March 23 to April 25, 1905. 210 fli?5
transmitting feed.' The hi ' '? * ,ntervenfd between the first infecting feed and the
sonies were ever seen C k'.’° -Wa® examined regularly until August 4 ; no trypano-
i9°5, when the animal i- all,l',atlons were then discontinued until September 18,
autopsy. ammal d‘efl of tuberculosis. No trypanosomes were seen at the

2905. 206 flies were/ fed^°n*<eV ^ erf°l>tt^ecus sckmidti). F'rom April 2 to April 27.
first infecting feed nnri tv,1" nianv jnstances 25 to 30 days intervened between the
UP to July 29 ; no try panosomeT Sm ,ttln^ feed- The blood was examined regularly
September 29, k,,,-’ nf cr. ‘s 'vere ever seen (malaria present). The animal died
were seen at the autnn«« 10eR ’ no ,r>’Panosomes or signs of trvpanosomiasi?
autopsy done immediately after death.

XI. FOURTEEN DAYS INTERVAL OR MORE

To infect a susceptible animal with Trypanosoma gambiense
by the bites of tsetse flies fed at least fourteen days previously on an
infected animal.

Leopoldville.

Experiment 79. — Guinea-pig. From January 11 to May 24, 1904, 156 flies fed
on this animal. Its blood was examined regularly until May 27, and then at
intervals until October, 1904. Trypanosomes were never seen. The animal died
in November, 1905. No trypanosomes were seen at the autopsy.

XII. MISCELLANEOUS

Experiment 21 i. — Guinea-pig. This animal was used for feeding odd batches
of tsetse-flies which for various reasons could not be employed in one of the set
experiments.

From May 16, 1904, to April 30, 1905, 1,278 flies were fed on it ; 200 of these
flies were freshly caught, the remaining had been fed at various antecedent periods
on animals infected with either T ry panosoma gambiense or 'L' ry panosoma dimorphon.
The blood was examined until August, 1905 ; trypanosomes were never seen. The
animal died in September, 1905 (cause of death ?). No trypanosomes or signs of
trypanosomiasis were seen at the autopsy done immediately after death.

In the Gambia it was thought that the excessively dry atmosphere
at the season when our experiments were done might have accounted
for their failure1. This objection cannot be made to the Congo
experiments ; when they were done, the average morning humidity
(6 am.) at Leopoldville was about 45 per cent., at Kasongo about
94 per cent.

Experiments 129, 203, 206, in which the animals were not observed
for a very long period, should probably be disregarded. The
remaining observations should be considered in conjunction with the
published reports of similar experiments done in Uganda 8' 10,

Bruce, Nabarro and Greig (8).— These authors publish three experiments in
''Inch animals were infected with Trypanosoma gambiense , within from two to
(our and a half weeks from the commencement of the experiment, by the bites of
from 200 to 1,000 freshly-caught Glossina palpalis. The rapidity with which these
experiments succeeded is noteworthy. _

In five experiments they succeeded in transmitting Trypanosoma gambiense by

bites of from 250 to 570 flies fed from eight to forty-eight hours previously on
‘nfected animals. The length of time from the commencement of the experiment
t0 ihe recognition of the infection varied from seven to ten weeks.

. r,reig and Gray (10). — In five experiments these authors succeeded in trans¬
iting cattle trypanosomes by the bites of Glossina palpalis. The period elapsing
,.etw1e,en the commencement of the experiment and the detection of the parasites in
^ blood of the experimental animal varied from four and a half to seventeen and

a weeks ; the number of flies fed ranged from 151 to 923.

N

212

It is seen that all the results are in conformity with the
hypothesis that Glossina pal pal is transmits Trypanosoma gambiaut
mechanically, and that it is probably not able to do so when the space
between transmitting and infecting feeds much exceeds 48 horns.
This conclusion is, nevertheless, to our minds a most unsatisfying
one if we are to regard this Glossina as the chief* or only t carrier of
T rypanosoma gambiense.

The experiments of all observers show that it is frequently
necessary to feed hundreds, almost thousands, of flies oil a susceptible
animal before it becomes infected !

It may be objected that in many of these experiments flies may have been
unnecessarily fed on the animals after they were infected but before the infection
was recognised. 'Ihis objection is partially negatived by observations (7, S: see
page 20) showing that the incubation period of a natural infection by Tryfamncni
gambiense may be so short as from two to four weeks.

If the number of successes obtained in such experiments, where
flies are fed on an infected animal whose blood is swarming with
parasites, is so small, it scarcely seems possible that mechanical
transmission by tsetse flies can alone be responsible for the rapi
spread of sleeping sickness of recent years. From these experiment;
it seems that, as a rule, a native must be bitten by a comparatively
laigc number of flies, which have fed not more than 48 hours
previously on a case of human trypanosomiasis§ before he will become
infected ; it must be remembered that trypanosomes are usually rather
rare in the blood of cases of sleeping sickness.

that Varieties of Glossina other than fal falls may <
GloisVnn 7'?J?mbte,,Se must .not be forgotten. It has recently been shown
Greicr an/ (7° T may transmit trypanosomes other than Tryfanosoma gamin
domestic animal ’^ve shown that it will convey the trypanosomes found in vai
in transmitting rlnv,Uganda (IO)' In Experiment 212 of this paper we succe
that ?n Germagn Tptan.°.so.ma din, or f bon by Glossina falfalis. Koch (9) sugf
■pallidi-te anri rv'35* A|.nca cattIe trypanosomes may be transmitted by Glos

K^totefc11 as * which was form

by G l»torn 7^ntS transmitted 7'. brucei, previously only known to be car

^ by the^bite^of1 a^banid'T/l

than m7nP7ZTva^a7bienSuhas as yet been found in nature in no other h
Commission on%iL c® fh,S was wri«en Report VIII of the Royal Socie

have found T. gambLse in^iv^ogs^ reCeived‘ Its authors Mien *

213

Although the chronicity of the disease is well known, it seems
impossible that large percentages of populations, whose vocation does
not keep them constantly on the water, should -become infected in
such a way in places where tsetse flies are far from plentiful, like
Kalombe (17 per cent, infected), Dibwe (18 per cent.), Miambwe
(/percent.), Lokula (6 per cent.) in the Congo Free State.

It seems certain that such a mechanical transmission cannot be
the only way in which Trypanosoma gambiense is transmitted from
man to man. We believe either that something is wrong in the way
in which Glossina palpalis has been used in these experiments, or
that Trypanosoma gambiense can be conveyed by some other means
than by it.*

The status of cattle trypanosomiasis (see page 233) in the Congo Free State
furnishes an additional objection to explaining the propagation of trypanosomiasis
by mechanical transmission alone. A considerable percentage of the cattle are
infected. Trypanosomes are very rarely seen in their blood. Enormous quantities
(to to 5oc.cm. or more) of infected blood must be injected into susceptible animals
before they become infected, and failures to infect are common. It does not seem
possible that the mere mechanical transmission of an infinitesimal quantity of
blood from animal to animal by an insect’s proboscis can adequately explain the
distribution of a strain of trypanosomes possessing so little virulence in direct
inoculations.

II. ATTEMPTS TO TRANSMIT TRYPANOSOMES BY
VARIOUS BLOOD-SUCKING ARTHROPODS

If the tsetse fly is merely a mechanical transmitter of trypanosomes,
there seems no reason why other blood-sucking arthropods should not
also transmit them. To test this possibility the experiments described
below were made with the “ Congo floor maggot ” (larva of
Auchmeromyia luteola ), Anophelines (Pyretophorus coslalis ), Simu-
hum and Ornilhodorus moubaia. None of them were successful. As
before, animals showing large numbers of trypanosomes in their blood
were preferred for the infecting feeds.

1 EXPERIMENTS with larvae of auchmeromyia luteola

I. FRESHLY-CAUGHT LARVAE

to) To ascertain whether larvae caught in native huts in an area
where sleeping sickness is endemic contain trypanosomes in a state capable
°f infecting susceptible animals. _ _

differ/i e9uiterdu’», usually only transmitted by coitus, may be spread in a totally
nt manner by the bites of fleas (25).

214

Experiments 41 and 4 2. — Rats. On November jo, 1903, these two ami;
were inoculated subcutaneously with the fluid obtained by rubbing up 16 lam*
freshly-caught at Wathen, in normal saline. Roth rats were examined freqim
(every three days) until April, 1904; neither ever became infected.

It is noteworthy that no trypanosomes were seen in the blood
from the alimentary canals of six freshly-caught larvae, although
living trypanosomes have been found in the intestines of larvae upio
10 to 13 hours after they had fed on a known infected animal

(b) To ascertain whether larvae, caught in native huh at With:
and Leopoldville, are able to infect a susceptible animal with trypamm
by their bites.

The larvae experimented with were always left in tins containing
dry sand. Animals, on which it was desired that the larvae should
feed, were tied to an arched, wire frame-work and placed in the tin;
so that their cleanly-shaved abdomens rested on the surface of the
sand. Under these conditions, the larvae commenced to feed in about
an hour. They apparently fed daily and for many minutes at a time,
unless the animals were moved, when they immediately burrowed
into the sand.

22a larva p f 1 ~C'uinea-pig. From December 18, 1903, to March 12,19°
intervals nnfa nt 6 .blood Was examined frequently until May, 1904, and then
were never seen °Vember’ I9°s» when the animal died of pneumonia ; trypanosonn

II. INTERRUPTED FEEDING

Lo ascertain whether larvae immediately previously fed o
ima heavily infected with Trypanosoma gambiense were ca
ismittmg the parasite by their bites to a susceptible animal.
xperiments 107a and 108 an infected guinea-pig was pi
mh .L?C °r ftW° llOUrS 111 tJie t,n containing the larvae; it was
nip-hf ^ \\n aCe<^ ^ a healthy animal which was left in the tin c
, ien the infected animals were removed a dozen mag
hirrhh reqi‘einly seen to he feeding on them ; it therefore se
animal ^ ^ that many of these resumed their feed on the sec
which fp>r? 1 1C recorc^s these experiments the number of lai
c uring the night on either or both of the animals is no
Experiment io7ci p..?

both animals. Rlood nea'pig‘ On March n, 1904, 39 larvae fed on on>

October, i9o4) when th - examined regularly until May, and then at intervals 0
somes were never seen * guInea‘P*& was used for another experiment; trvpa

Experiment 108. — Guinea-pig. From March 13 to 27, 1904, 194 larvae were
fed. The blood was examined regularly until the end of May, and then at long
intervals until September, 1905 (death from pneumonia) ; trypanosomes were never
seen.

III. TWELVE HOURS INTERVAL

To infect a susceptible animal with T rypanosoma gambiense by
the bites of larvae fed at least 12 hours* previously on an infected
animal.

Experiment 60. — Guinea-pig. From December 30, 1903, to April 27, 1904, 454
larvae were fed. The blood was frequently examined until May, 1904, and then at
intervals until October, 1905 (death from pneumonia) ; trypanosomes were never
seen.

II. EXPERIMENT WITH ANOPHELINES

The mosquitoes used in this experiment were bred from larvae
caught at Leopoldville; they were practically all Pyretophorus
costalis.

To infect a susceptible animal with Trypanosoma gambiense
by the bites of anopheles fed on an infected animal at least 24 hours
previously.

Experiment 53.— Guinea-pig. From December 15, 1903, to May 8, 1904, about
75 mosquitoes, fed from twelve to 72 hours previously on infected animals,
fed on this guinea-pig. The blood was examined regularly without result until the
animal was accidentally killed on August 8, 1904 ; no trypanosomes or signs of
trypanosomiasis were seen at the autopsy.

HI. EXPERIMENT WITH SIMULlUAl

During the rainy season there were legions of Simulium about
the sleeping sickness hospital at Leopoldville. It was hoped to do a
series of experiments with them, but they were found to be so difficult
to manage that the idea was given up.

Some of these flies, freshly caught at the hospital, were, however, fed on a
guinea-pig; five of them had in addition fed an hour previously on a heavily
infected guinea-pig. The animal was kept under observation for 22 months ; it
never became infected.

IV. EXPERIMENTS WITH ORNJTHODOROS MOUBATA

Because of the following observation, it was thought that this
tick might disseminate sleeping sickness.

At Nyangwe, out of four women presenting themselves as suffering from fever
alter ihe bites of Ornilhodoros moubata, three were infected with trypanosomiasis.
AH four women were wives of an arabised chief and inhabited a house, "
Ornilhodoros moubata swarmed, in a well-kept village situated in open count J ’
tsetse-flies were consequently very rarely seen in the neighbourhood of their
dwelling place. _

.I*te larvae were used several Umts and therefore some

anS‘entS I07a’ 168 and 60 had P°ssibly fed seVCral UmeS

of those employed in
previously on infected

INTERRUPTED FEEDING

1 o infect a susceptible animal with Trypanosoma gambiense
the bites of Ornithodoros moubata which had fed from three to h
minutes previously on a heavily infected animal.

t i E£ET:EN,T I— 1 Guinea-pig. From August 29 to October 25, 1906, 362 ticks
tea. I he blood was examined almost daily without result until January 3, m
when the animal died of pneumonia ; no trypanosomes or signs of trypanosomas

were seen at the autopsy. A rat sub-inoculated on November 18 did not become
infected.

. ^x*’eri'[ent 2. Guinea-pig. from September 27 to November 24, 1906, ;f
6 le 00<^ was examined daily until January 21, 1907 : no trypanosome
were seen . a rat sub-inoculated on December 19 has not become infected.

i'EN 1 IS'oveiuBer a6> 1906, 129 ticks fed. The blood wu

. ,6. a most ai y up to January 21, 1907. No trypanosomes were seen. A

at sub-inoculated on December 19 has not become infected

These animals (Experiments 2 and 3) were inoculated in February,
1907, with Trypanosoma gambiense. Both became infected and their
disease ran a usual course.

Although these experiments are far from conclusive, they seem to

indicate that Trypanosoma gambiense is probably not easily

transmitted by any of the common arthropods experimented with*

The alternative explanation of the comparative insuccess of

attempts to transmit Trypanosoma gambiense by Glossina falfaT

as that the experiments had not been carried out under the best
conditions.

unless thev9were’f<\Ste i trypanosomes could not be transmitted by tsetse flies
He suddoVi pH tK.a tlie at some particular point in their development

trypanosomes m net su^’estlon by the observation that he could transmit cattle

animals whose blood^1 \ ? tlle bites °f tsetse flies previously fed on long-infected
s whose blood contained very few trypanosomes.

th SU^estlon course premises some sort of development of

hap TPanosome in the tsetse fly. From what is known of other
_ oa there seems to be no reason why there should be none.

Gray (i0, page followinS observations should be noted. Greig and

Glossina palpalis bv the hit transmit the cattle trypanosomes conveyed by

Nabarro and Gre.V / , (see page 20t>'

Bruce (12 'J5) a so failed to transmit these trypanosomes by Stomoxyi-

taneous inSKj&(2h T rypanosoma brucei, found no case of spon-
Hies, save Glossina, weregpresentSt°Ck an‘mals *n an area where all sorts of biting

insects they employed**" *th«a™rSe °bservers did not try interrupted feeding.

mechanical transmitters ■'seTbln had. W opportunity of acting »

Gray ( 1 7, succeeded in transmit “.u810? ,of. surra (*<>)• Minchin, Tulloch and
ed interruptedly. lnf> the “ Jinja ” cattle trypanosome by Sfoixox)

- T 7

III. DISSECTIONS OF TSETSE FLIES

The dissections of tsetse flies commenced in the Gambia,1 with the
object of finding such a phase of the trypanosomes, were continued

in the Congo.

Bruce14 found scanty trypanosomes in the proboscis of Glossina
morsitans fed 46 or fewer hours previously on an animal infected
with nagana; it is quite conceivable that the parasites should be
mechanically transmitted by the bites of such insects. It was therefore
important to determine how long after feeding on an infected animal
Trypanosoma gambiense could be found in the proboscides of
Glossina pal pal is. A series of 75 dissections* were done at periods
ranging from one minute to 20 hours after feeding. It was found that
red cells and trypanosomes were almost always present in the labium
up to ten minutes after feeding. Later than this the parasites were
frequently absent, although red cells might still be present. 1 he
longest period after feeding at which trypanosomes were found was
if hours; the longest period for red cells was hours. Red cells
were seen in only six out of 25 dissections done more than two hours

ifter feeding.

Koch9 by pressing the bulb at the base of the tsetse s proboscis
vas able to obtain a clear fluid in which he frequently found
rypanosomes. We found that by irritating a fly' held by the wings
t could frequently be made to spontaneously exude such a fluid ; only
mce was a trypanosome seen in it, and that in a fly which had fed one
md a half hours previously on a heavily-infected animal (red cells
dso present in fluid). It was also found that flies caught after they
Had fed on an infected animal frequently regurgitated in struggling
1 drop of blood, as large as a pin’s head, which was full of parasites,
many of them identical in form with those ingested. This observation
was made up to 28 hours after the last feed on an infected animal.

In tins connection it is well to remember that trypanosomes may live apparently
unaltered in the fly’s stomach for at least 48 hours. Such a regu g ans Qf

Place during the feeding of the fly can be easily conceived as a possib

transmission of the parasite. _ _ _ -

,, *The flies previously fed and kept in test tubes vYere,c^r gt^e 'fly was cut off
th!»,Wlngs wi,h “ little" struggling as possible. The head t JLlirJtation. and the
a quick cut of a fine pair , of scissors, in order to prevent regurgUation, of

proboscis was then detached and dissected for examination in a tin)
normal salt solution.

IV. MORPHOLOGY OF TRYPANOSOMES IN THE
ALIMENTARY CANAL OF CERTAIN
ARTHROPODA*

I. IN G LOSS! A' A /‘ALIA US

The observations here recorded were made on material obtained

from dissections done in the Gambia and in the Congo in 1903 and

the first months of 1904. Work of more immediately practical

importance prevented us from giving as much time to this investigation

as could have been wished, and has deferred the examination of our

stained specimens and publication of our findings until now. The

members of the Royal Society's Sleeping Sickness Commission and

Koch have in the meantime published their work on the same subject

In many points our observations coincide with theirs. For this

reason we do not give detailed descriptions of forms observed, and

this especially since little can be said concerning their significance.

All our work was done with Trypanosoma gambiense and Glossina

paipalis. 1-lies were dissected and examined in fresh and stained

specimens (dry films) at every period from a few minutes up to eleven

days after feeding on an infected animal. Their intestinal contents,

muscles, ovaries, malpighian tubules, blood and salivary glands have

een examined. Only in the alimentary canal have forms been found

VV 1C1 can certainly be connected with the trypanosomes. Active

er^ parasites were seen in the alimentary canal up 1048 hours;

vmg, ut altered, trypanosomes up to 72 hours after feeding. No

gnsa le trypanosomes were ever seen in the faeces of tsetse

In ^ j 'e C*Cai excreted per anum immediately after a fly

has finished its feed.

stor m,HeSt!°n “ ^ a ^ over 24 hours
whole . 'S usua% emptied of blood. In about 50 to 70 hour

faeces S ^ 1S save f°r a small amount of greenish-t

apnarentl 'ng:es[i0n t^e trypanosomes become more active

frequent 7 numerous i longitudinal division forms

hXigested oaV\bCen SCen UP t0 hours after feeding. 3
_ _ Parasites^are distorted, vacuolated and granular,

• ’ g, it must be understood that controls Here constantly examine

are breaking up. They seen) to be degenerating. Groups of five or
more parasites agglomerated by their posterior extremities frequently
occur. These usually degenerate until only shapeless debris and
chromatic granules remain (on one occasion spherical forms were
produced).*

In blood ingested for some days parasites approximating to the male
and female types21 become relatively much more numerous than at
first. The structure of the nucleus varies. The chromatin may be
collected at either pole, or in three or four large masses. It may
be irregularly arranged in transverse rods or distributed in granules
placed on a chromatic reticulum. Sometimes the granules are
placed about the periphery of the nucleus. Chromatophilic granules
occur in the cytoplasm. These have been seen in positions suggesting
that they have been extruded from the nucleus.1' Occasionally one
or perhaps two large faintly pink-staining areas, somewhat diffuse,
but nevertheless possessing a definite outline, have occurred in the
neighbourhood of (usually posterior to) the nucleus. This area
suggests the chlamydoplasm34 observed in Leucocytozoon danielewski.
The blepharoplast often consists of two or more granules. It is
frequently very apparent that it is placed in a “ clear area and that
the thickened edge of the undulating membrane ends not in it but in
a pinkish basal granule or “ diplosome."

Trypanosomes occur which possess a third, deeply-staining
chromatic area (one-third the size of the nucleus) in addition to
nucleus and blepharoplast. Other polynuclear forms resembling in
appearance the polynuclear forms described by Kochf have been seen
(48 hours after feeding). We can say nothing concerning their
significance, but we are not prepared to follow Koch in his interpre¬
tation of them. We believe that the rounded forms, which he thinks
they produce, arise from the englobation of single trypanosomes which
have cast off blepharoplast and undulating membrane and become
spherical. (This process has been observed in blood within three
hours after its ingestion.) The bodies so formed are rounded or oval,
and measure about 40/r to you by 2'on to 3*0/1. I liey consist of a
hght-blue-stained protoplasm with definite contour and at first one,
late^v^ajarge and a small, chromatic masses. We have not seen

the ,t^ccasioi}al confluent collections of degenerated parasites are seen which recall
Pasmodiar’ masses of Plimmer and Bradford (24)- , . „nv

rpiftniKi°Ch gives no measurements. We cannot say whether there is « .

' blance»n size; our forms measure i8-o/t to 25-0/1 by you to 3-75/1.

220

the subsequent formation of a flagellum, and the assumption of
" herpetomonas-like " characters described by Koch (their occurrence
seems most probable).34

The spherical forms are produced in Glossina in the same way as
in the rat flea (see below). The independent movements of flagellum
and cytoplasm are most striking in parasites in the process of exflagel¬
lation. When separated the flagellum and blepharoplast quickly
become motionless ; the cytoplasm is mobile for a little longer. These
rounded forms were not seen to divide, but from the analogy of
Trypanosoma loricatum? 4 and from the difference in size between
the smallest rounded forms (4*5 by 3*75/1) and the average pear-
shaped trypanosome (8‘o/i by 5*0/1) just before ex flagellation is
complete, it seems possible that they may do so at this stage. After
the flagellum and blepharoplast are cast off, the nucleus, at all events
in some cases, fragments into from eight to ten granules. These
then collect together to form a new central mass (some may remain in
the cytoplasm ; are any extruded ?). Before the motor apparatus is
thrown off from the rounded trypanosome it sometimes circumscribes
the parasite so as to simulate a cyst wall no real encystation has been
observed. (Moore and Breinl26 describe the formation of a cyst wall
in similarly rounded parasites observed in trypanosome-infected
animals treated by atoxyl.)

II. IN PED1CUL US

In July and August, 1903, a few pediculi caught on a rat very
heavily infected with 7 rypanosoma gambiense were dissected and
examined fresh at room temperature. Granular, indistinct, obviously
exonerating ParasiIes were of course present. The main change
seen in those apparently normal was the formation of ovoid or
spherical trypanosomes (5-6/1 by 6-4/1 to 8-32/1 by 7-6/1). These had
stained specimens the same appearance as those observed u>
trypanosomes from the gut of tsetse flies. The following short
one observation sufficiently describes what was seen >n

caught lousp . r“t* fiIled with bright red blood, was drawn from a fre

active normal . Was 1 en tease<I out in a minute drop of normal saline.
stumpy forms Wer® Present- All were long and large at first

later the normal na - Some of the parasites were quite granular. One

present which haH w* ^ ® l6SS numerous, and pear-shaped trypanosomes ’

to which the flal n UndulatlnS membrane and flagellum. Similar forms exi
* flagellUm was s*ill attached by the blepharoplast. In these fc

221

flagellum and cytoplasm move quite independently of one another. The movements
of the cytoplasm were both amoeboid and lashing. These forms, as well as some
very stumpy and granular, fairly round trypanosomes, may roll themselves up into
a ball. (If the flagellum be present it is usually external.) Some seemed to be
enclosed in some sort of a membrane, since, although motion was violent, they
never changed their spherical shape. Others straightened out for a moment to
immediately curl up again. In stained specimens no definite cyst wall was ever
seen to enclose a spherical parasite.

One apparently normal trypanosome was first seen at this time inside a large
dear vacuolated cell of the stomach wall.* During the three and a half hours it
was watched it remained active and unable to leave the cell. After four hours
bacteria commenced to multiply in the specimen, and it was necessary soon after
to discard it. No further change was observed in the spherical forms produced by
the rounding of the above described “ pear-shaped ” parasites.

III. IN STOMOXYS

In the Gambia, trypanosomes, identical with those ingested, were
found unchanged in the gut of Stomoxys up to 20 hours after they
had fed heavily on a horse infected with Trypanosoma gambiense.
Longitudinal divisional forms were seen. In one instance phago¬
cytosis (by a cell from the horse’s blood P) of a trypanosome was seen
at this period.

An interesting observation was made in a fly fed 18 hours previously. Two try¬
panosomes were attached to one another ; the separate outlines of each parasite could
be distinguished perfectly. In about an hour and a half, the two parasites had fused
to form a single trypanosome. The newly-formed parasite was thicker than is
normal, had a shorter flagellum and a collection of very minute refractive granules
at either end and no perceptible nucleus in the usual position. After twelve hours the
parasite was motionless ; no further change had occurred. This phenomenon was
twice seen.

IV. IN LARVA OF AUCHMEROMYIA LUTEOLA

The contents of the intestinal canal, the salivary glands and blood of
larvae fed from six to thirty-four hours previously on animals infected
with Trypanosoma gambiense were examined. The changes
observed were very similar to those seen in the other arthropods.
Living trypanosomes were seen in the fresh blood of the diverticulum
up to twelve hours after feeding, and no parasites were seen in the old
black, half-digested blood. At ten hours many parasites were dead ;
numerous agglomerated and degenerating trypanosomes were present ;
some of the stumpier parasites were becoming rounded. The
intestinal contents contained many bacteria ; perhaps for this reason
comparatively few small spherical forms of the trypanosomes were
seen.

of/*** (quoted from 19, page 113) reports having seen intracellular for

Dat hypanosome and of T. Itwisi (?).

222

V. IN ANOPHELINES !

A large series of Anophelines ( Pyretophorus cost alts) was dissected
at periods ranging from twelve hours to eleven days after
feeding on experimental animals heavily infected with Trypanosoma
gambiense. In the interval between feeding and being dissected, the
mosquitoes were fed every two days on uninfected animals. Alimentary
canal, malpighian tubules, ovaries, and salivary glands were carefully
examined, and forms recognisable as connected with the trypanosomes
were seen only in the alimentary canal ; at first some confusion was
caused by the presence of the sporozoites of a coccidium-like
protozoon.14 After 42 hours no trypanosomes were seen. From 12
up to 42 hours after feeding the changes similar to those described in
the gut of other arthropods were observed. Nothing peculiar was
seen.

V. DISCUSSION

When malarial parasites are ingested by a receptive anopheles,
only those parasites survive which are prepared for development in
the environment provided by their new host. The remainder die.
If the blood is heavily infected there is therefore necessarily a large
number of degenerating parasites present in the insect’s alimentary
canal. The same may be true of trypanosomes ingested by tsetse flies,
he mass of degenerating parasites confuses one’s perception and
ia <es it extremely difficult to pick out the forms capable of further
development.

It is by examining in fresh coverslips and, later, stained prepara-
«ons, the parasites in trypanosome-infected blood taken from the gut of
cts, that the changes invariably taking place under such conditions
-■^ki ^ ^eterm*ned- ft seems reasonable to deduce that forms
ana y appearing are those best suited for the new environment,
f ti ^ thCymay P°ssihly concerned with the further development
parasite. Such forms should therefore be most carefully

communication was in the n^int(!i!,’t»!lisJConnection was received while the presen
survives ingestion bv mosnnit,»«c hancJs. Its authors show that Trypanosoma bru:‘
only 14 hours. The^pS * *ours = * remains viruYent to mice to

e si the ingested parasites is not reported on.

223

watched. Up to a certain point observations on trypanosomes in
cultures may be very valuable here.

Bouet (Culture du Trypanosoma de la grenouille, Annales de l’lnst. Pasteur,
Yol. XX, No. 7, page 364) has obtained a very large part of the “ swarm ” cycle (34) of
this parasite in cultures.

Novy and MacNeal, in their studies on mammalian and bird trypanosomes,
illustrate forms which are interesting in this connection. Life History of Trypano¬
soma Icaiisi and T rypanosoma brucei, Journal of Infect. Diseases, Vol. I, No. 4,
Xov. 5, 1904, p. 517. On the trypanosomes of birds, Journ. Infect. Diseases, Vol.
II, No. 2, March 1, 1905, p. 256.

Cerquira, in a thesis published at Rio de Janeiro, describes a most interesting
development in bird trypanosomes in cultures. It bears evident analogies to the
multiplication process described in T. loricatum (34) (reviewed in Journal of Tropical
Medicine, April 1, 1907, p. 118).

A process which has been seen by many observers in trypanosomes
ingested by arthropoda is the formation of spherical bodies without
undulating membrane, flagellum or, at first, differentiated blepharo-
plast. Similar forms have been reported by many different authors
in most of the mammalian trypanosomes when they are placed in
various “ unfavourable circumstances,”19 such as in moribund animals,
in the cerebro-spinal fluid, in organs, kept preparations ol fresh blood,
cultures, gut of lice, fleas and biting flies.

These rounded parasites bear a most striking resemblance to the
rounded form of Trypanosoma loricatum which commences the
interesting cycle of “ swarm ” development first described by
Danielewsky.27 These forms of the mammalian parasites are formed
in much the same way as are the rounded frog trypanosomes. Does
their further development proceed in the same way ? (see footnote to
page 62 of 27). The work of the following observers shows that there
may be some resemblance between their subsequent stages.

Bradford and Plimmer (24) describe the rapid division of spherical “amoeboid
forms” of T. brucei in the blood of a vertebrate host.

Prowazek (22) also reports the division of such bodies in T. brucei ; and
describes the formation of similar spherical bodies in T. lewisi ingested by a louse ;
and he suggests that these later develop a flagellum.

Koch (9) describes the formation of rounded forms, which subsequently develop
a flagellum, in cattle trypanosomes ingested by Glossma. In their deve opmen
these parasites pass through a herpetomonas-like stage.

Lingard (28) describes certain developmental processes in trypanosomes o
ndian cattle where similar rounded forms occur.

Moore and Breinl (26) have observed similar forms in T. gambiense, t special y in
animals treated by atoxyl; their further development has not been followed. A seco
‘vpe of rounded form, produced by the extrusion of the nucleus from an ord
trypanosome, was observed to produce a minute flagellum.

224

Durrant an^ Holmes (30) describe similar rounded forms in T. evanti,
c atter described them as undergoing development. He also describes the second
type of rounded trypanosome and its development as do Breinl and Moore.

It is impossible to say how far this resemblance extends. At all
events, the above observations prove that the spherical forms represent
n stage in a definite cycle of development other than that of
longitudinal division, the only definitely developmental phenomenon
previously established in the life history of trypanosomes. The
observations of Holmes, Moore and Breinl show that there is possibly
a. third line of development in mammalian trypanosomes, in addition
to the multiple segmentation first described in Trypanosoma lewisiT

Some of the rounded forms of mammalian trypanosomes doubtless
degenerate and die, as do similar forms of Trypanosoma loricatum.
But this constitutes no reason for calling all rounded forms degenera¬
tive or involutive, as has been so frequently done. The greatest
caution must be used in applying such terms ; it is chastening to
remember that only ten years ago a flagellating malaria parasite was
w idely believed to be a degenerating form !

Conjugation* has not been seen certainly. Possibly it does not occur
during multiplication in Trypanosoma loricatum by the swarm
cycle; but it is not at all impossible that a union of sexually
differentiated individuals may occur at some other period.!

of a cycle (probably sexual)

It has been stated that the

occurrence

ln niammalian trypanosomes ingested by tsetse flies is inherently
mprobable. The following are the points most usually cited
support this view : _

(0 It is said that there is no need for a sexual cycle since
trypanosomes may be transmitted for an indefinite pe«°d
bites a”ICa^’ ^rom an>nial to animal by simple inoculation and b) )

(2) that the trypanosomes

may be kept for very long periods

wel

diff

sug

225

(3) That another adequate method of reproduction (now probably
at least two) already exists.

(4) That the contents of the intestinal canal of tsetse flies
previously fed on trypanosome-infected animals when inoculated into
susceptible animals does not produce trypanosome infection (nor, in the
cases of bird trypanosomes and mosquitoes, will they produce cultures
of trypanosomes).

None of these arguments is valid. Since those who uphold them
usually oppose Schaudinn’s work-1 on Haemoproteus, etc., we shall
not mention it in answering their arguments, but shall refer only to
the work of other authors. Each objection is answered sepaiatel)
by referring to some occurrence contrary to it in the known life history
of some other protozoon.

(1) Human malaria can be transmitted by simple inoculation,
Trypanosoma lewisi , Piroplasma bovis31* and Haematnoeba relict a
can be propagated indefinitely by inoculation. In all four of these
protozoa a definite sexual process occurs when the mature parasites
come into a favourable environment, the alimentary canal of the

required arthropod.

(2) 7 rypanosoma lewisi has been successfully cultivated.
Prowazek22 describes a sexual conjugation which it undergoes in lice.

Although no other pathogenic protozoa have been cultivated with
'he same success as the trypanosomata, it is well known that certain
non-pathogenic protozoa may be kept for very long periods in pure
cultures without the appearance of any sexual forms. On the
Production of circumstances impelling them, sexual forms may appear
ln these cultures. Granting that the sexual forms are absent from,
not merely unidentified in, the usual cultures of trypanosomes, it y
no means follows that they may not be produced under other
'cuinstances. The conditions causing their appearance are not Pre^

'n the binary culture medium. For the same lack of “ impe mg
C*Uses" the periodic absence of infective sexual forms from the blood
°f Infected animals is easily conceivable. The question natural >

Ses’ w^at are the conditions “impelling” the production ol s-

Professor Nuttall permits us to state that he is of ophuonthat Fir* \as already
imm^TUtted indefinitely from dog to dog by inoculatiom«
the disease in this way for more than twenty ge

226

forms of trypanosomes? The analogy of malaria in man and of
protozoa in cultures suggests an answer — chronic infections.*

(3) Young malarial parasites (ring forms) divide by direct division ;
a cycle of sporulation exists, yet there is also a sexual cycle.

(4) These facts need only signify that the parasites were not, at
the moment of inoculation, in a state favourable to cultivation or to
the production of infection in a vertebrate host.

Coccidium schubergi is parasitic in a centipede. Infection takes place by the
digestive canal. If an infected centipede be eaten by another, infection may be
conveyed by almost every stage of the parasite ingested. The immature gameto-
cytes, however, are unable to infect ; although they are in their normal host their
surroundings are unsuited to their further development. They consequently die.

In cover-slip preparations the malarial gametocyte is fertilized,
development goes but little further ; the surroundings are not
favourable. Why should the sexual cycle of a trypanosome necessarily
occur in the same medium as its asexual one ? Crescents, males at
least, adult malarial parasites, if left in man degenerate ; the young
parasites die in the mosquito. The surroundings suited to one form
are not to the other. Why should the hypothetical sexual form of a

trypanosome develop equally well in either invertebrate or vertebrate
host ?

The cited arguments proffered against the existence of a sexual
cycle of trypanosomes in tsetse flies are contradicted by these known
acts. What are the arguments in favour of it ? They are briefly = —
(0 The analogy of other protozoa.

(2) T he occurrence of parasites possessing sexual characteristics m
ood of infected animalsf (compare Trypanosoma dimorphon )■

(3) Prowazek’s observations on the development of Trypanosoma
tewisi in the rat.22

trvnan KocllS.mcornPlete observations on the development of cattle

trypanosomes in Glossina*

'Pposedly sexual forms, somewhat similar to some of those

made on

somes they ingested onlvltp^f Glossinae only became in
animal m whose blood the n °P-ed lf their feed had been
f The acknowled paras,tes were scanty,
tneasure on a recognition 'of* nature of these forms of course depends ii
een reP°rted in nianv trvnan east a Part of Schaudinn’s work (21). The
• 1 os°mes by many independent observers.

227

described by Koch, are reported in Trypanosoma gambicnse ingested
by Glossina palpal is .* 11

(0) We have observ ed similar forms to (5) and in 011c instance at
least the fusion, we believe, of two living Trypanosoma gambicnse in
blood from the gut of .1 Slomoxys.

The arguments on neither side are conclusive, but we believe those
to be far the stronger which support the existence of a sexual cycle
m the life history of the mammalian trypanosomes, possibly in the
tsetse fly.

It is, we believe, by the most patient and long-continued observation
of living, individual parasites in trypanosome-containing blood
(probably freshly ingested) from the gut of tsetse flies that a definite
answer can best be given to the whole question. The work should be
done under natural conditions in Africa and, naturally, the observations
on living parasites must be controlled and supplemented by the study
of stained specimens. The observer must approach the subject in a
receptive mood and freed from preconceptions.

VI. CONCLUSIONS

from the consideration of all the facts we state the following

propositions :

It is known that,

(0 Mammalian trypanosomes may be mechanically transmitted by
l*le '3*tes °f blood-sucking arthropoda.

^ (2) A cycle of development of the trypanosomes exists in which
°CCUr spherical parasites formed by the throwing off of blepharoplast,
Undulating membrane and flagellum. These forms exist in both the
'ertebrate and the vertebrate hosts; conjugation probably plays no
£!l^he_production of this form. A possibly distinct cycle is

also describe “ ^ trypanosomes "•JOCC)!5rhe^entirelye distinct
,r°m the mam1Chi’ m common with N'ovy (23), they consider to recognisable

as other thamn?u ,an tr.vpanosonies. We have not seen trypai ( instanCe, we
found irvrri n 1 *lf,se 'ingested, in the Hies examined by us. , . but fresh

marn'Oal?an ^SOIVeS in lhe 8"’ "f a freshly-caught Glosano flagellate,
Was also nrob °°d was also present; another parasitic proto a > > which leads
Minchin P e«fem- These Ay trypanosomes (aof have an encysted host>

and so take ,,,l‘ir cysts may be excreted, iftgcsted interesting

,ine; iSStK m a “contaminative cycle.” The hypothesis is a most ^ of
ve,se flie.s PYn sfa,es that the sexual forms of T. gambicnse o not be found.

N'°thincr ..^tned disappeared and after 96 hours parasites rts 0f the flv

eitamined lat eni^]|n8 a trypanosome” was found in the 'arl,' ' ^ tbe copula of
lr-vPanosorn ’ ^ Why should developmental forms or the product ot
oe trypanosome-like in shape?

228

represented by the spherical parasites, . consisting almost entirely of
nucleus, which result from the disintegration of trypanosomes26,

(3) There is reason to suspect that a sexual cycle may also
occur ; it may be in either or in both hosts.

(4) The rapid spread of sleeping sickness cannot be fully explained
by (1) alone; the cycles of development mentioned in (2) and (3)

probably play a very important part in the transmission of the disease.

Runcorn Research Laboratories

April, 1907

LITERATURE

'• Dutton, J. E., and Todd, J. L. Memoir XI, Liverpool School of Tropic.il
Medicine, Sept., 1903.

■i. Bruce, Nabarro and Grf.tg. Reports of the Royal Society's Sleeping Sickness
Commission, No. 4, p. 67-61, Nov., 1903.

3- Kilborne and Smith shewed that the progeny of ticks fed on cattle inf 1,1

with Piroplasma bovis of Texas fever were alone capable of transmitting
the infection.

4- Thomas and Linton. The animal reactions ... of Trypanosoma gambunn

Die Lancet, May 14, 1904.

V Newstead, Dutton and Todd. Insects collected in the Congo t ree State
Annals of Tropical Medicine and Parasitology, Vol. I, No. i, p- ^ l e
>907 (Liverpool School of Tropical Medicine). . .

‘ rTTOX and To»u- Memoir XVI, Liverpool School of Tropical Medicine.
Page 97. August, 1904.

Di ITOK, Todd and Christy. Memoir XIII, Liverpool School of Tr0Pua
Medicine, page 97.

Bruce, Nauarro and Grkig.

'S' Na,!ARRo a"d Greig. U
Commission, July, ^

RKiG. Report V of the Royal Society Sleeping

Sickness

22Q

ii.. Lav era x and Mesxil. Trypanosomes et Trypanosomiases. Masson, Paris,
1894-

i;. Mixchin, Gray and Tclloch. Glosuna palpa/is in its relation to Trypano¬
soma gambiense and other trypanosomes. Proceedings of the Royal Society,
Series 13, Yol. LXXVT3I, Xo. II525, Oct. 12, 1906.
iS, Sergent, Ed. and Et. Etudes sur les trypanosomiases de Rerberie en 1905,
Ann. Inst. Past., t. XX, Aug, 1906, p. 665.

19. I.rHE. Die im blute schmarotzenden Protozoen, in Mense, Ilandbuch der
Tropenkrankheiten, Vol. TIT, part T. This gives an excellent review of what
is known concerning trypanosomes.

Mixchin. Encystation in Trypanosoma gravi, Xovy. Proc. Roy. Soc., Series
13, Yol. LXXIX. Xo. B528, Feb. 22, 1907.
u ScHAuniNN. Generations und Wirtswechsel bei Trypanosoma und Spirochaete.

Arbeiten aus. den. Kaiserlichen Gesundheitsamte, Vol. XX, p. 588, 1904.

■i- Prowazek. Studien fiber Saugetiertrypanosomen Arb. ad. Kaisr. Gesund,
\ol. XXII, second part, page 351, i9°5-

;.l- Now. The trypanosomes of tsetse flies. Journ. of Infect. Diseases, \ol. HI,
No. 5, May, 1906, p. 394.

24 Plisimer and Bradford. Trypanosoma brueti. Quarterly Journal of Micro¬
scopical Science, New Series, Xo. 179 (Vol. XLV, part 3), Feb., 1902, page 449-
25- Kabixovvitsch and Kempnkr. Die trypanosomen in der Menschen— und
Tierpathologie. Centralblatt f tir Bakteriologie. Abt. 1, orig. Hr. XXX ,

30.

P.

P-

13-

34-

the life-history of the parasite of

India.
4.

of

1904, Heft 5.

Moore and Bheinl. A preliminary note on
"Sleeping Sickness.” The Eancet, May 4th, 1907.

Danielewsky. T.a parasitologie comparee du sang, A. Darrc. Kharkoff, 0889.

I.ingaru. Different species of trypanosomata observed in bovines in India.
Hie Journal of Tropical Veterinary Science, Vol. H, No. 1, PaSe
February, 1907.

Hurrant. A trypanosoma found in blood of cattle in India. Journal
Comparative Pathology and Therapeutics, Vol. XVII, part 3, Septem er, 190

Holmes. Evolution of the Trypanosoma evansi. Journal of Comparat
Pathology and Therapeutics, Vol. XVII, part 3, September, i9n4-

Christophers. Preliminary note on the development of Piroplasma
1 1e Indian Medical Gazette, Dec., 1906, p. 467- (Clives a pre 1

account of conjugation). .

T^«ah and Mesnil. Recherches sur le trypanosome des rats. Anna es
Institut Pasteur, 1901, p. 675.

N(,w, Frederick G., MacNeal, Ward J., and Torrey, Harrv
Jrypanosomes of Mosquitoes and other Insects. Journal ot
leases, \ol. IV, No. 2, April 10, 1907. ., ■

D*TTOn> Todd and Tobey. “ Certain Parasitic Protozoa observed m AU ■

- 11 nals of Tropical Medicine and Parasitology. Vol. I, No. 3 (m

N. The

Infectious

CATTLE trypanosomiasis in the
CONGO FREE ST ATE

1

233

CATTLE TRYPANOSOMIASIS IN THE
CONGO FREE ST ATE*

BY

The late J. EVERETT Dl'I ION, M.B. (Vict.)

(WALTER MYERS FELLOW, UNIVERSITY OK LIVERPOOL)

JOHN L. TODD. BA., M.D., C.M. McGill

(DIRECTOR OF THE KISCORN RESEARCH LABORATORIES OF THE
LIVERPOOL SCHOOL OF TROPICAL MEDICINE)

AND

ALLAN KINGHORN. M.B. (Toronto)

(JOHNSTON colonial fellow, I'NiVERsi i y ^ oi ^J^Ychool of
DEMONSTRATOR AND RESEARCH ASSISI ANT , I -
TROPICAL MEDICINE)

Being the Eighth Progress Report of the Expedition of the Liverpool

School of Tropical Medicine to the Congo, WOj-Oj

{Received April 20th, 1907)

I. INTRODUCTION

When Europeans first came to the Congo there were practically no
:uttle in the greater part ol the central area of what is now t
state. Since then advent cattle have constantly been imported from
neighbouring African colonies.t

They have been sent to many posts in different parts of the Congo
with varying success. As a rule, it ,s said that cattle tend to do bes

plain country where there are no buffaloes. _ _ _

. •„ naoer was communicated in 1904

Much of the information contained in this P 1 , reports,

to the Government of the Congo Free State m unp herds Gf cattle.

t The natives in the highlands about Lake nave long, upstanding,

A® a nile the Kivu cattle G ig- 3l are rath*r S th(,m are of the humped zebu
slightly -curved horns 1 B.< aegyfliacus ?). • '”"5 , b the expedition1 generally

'ype. The cattle in the parts of the < ongo V1 sited by ^ be present m the
approximate to the European type. Tsetse ntes cattle taken from Lake

K|vu district. Game' is ‘plentiful. The mortality among c- ^ three have
Kivu to other districts is very great From herds of ^ (.attle died in herds
remained on arriving at Kasongo! It was foun hard. The cause of the

which travelled onlv every third day than m th(> f od but disease. Seventy-

mortality is therefore probably not fatigue and lack ot^ ^ until the forest was
•i'e head of cattle taken North from Luts bi all were dead,

entered. They then commenced to die. and at Mawambi

■

234

II. EXAMINATION OF CATTLE

We examined the cattle of only a few of the more important herds;
those existing at the remaining places visited are only briefly
considered, while herds unvisited are not mentioned.

At ZAMBIE there is a herd of about 190 cattle, all in excellent
condition; in 1904 there were 52 births, and no deaths from disease.
1 hose in charge recognise no disease resembling trypanosomiasis.

At the neighbouring island of MATEBA there is a herd of about
5,000 (?) cattle ; said to be in perfect condition. The original animals
at both Zambie and Mateba were imported from Portuguese West
Africa (San Paul de Loanda, etc.), Walfish Bay, the Canary Islands,
and occasionally from Europe. Both places are on low-lving grass
plains, and tsetse flies, although probably' present, were not seen
(October, 1903), and big game was practically absent. Living animals
are regularly sent from St. Paul de Loanda, from Mateba and from
posts in the Lower Congo to Leopoldville for food ; two said to come
from Mateba (?) were found to be infected with trypanosomes on
their arrival at Leopoldville.

The Leopoldville herd, kept only for food purposes, is heavily
infected 2- 3.

At Yumbie cattle have been present since 1S96, but they have died
as fast as they' reproduced and the herd has not increased. The
principal cause of death is “colic” (?). This disease is probably
trypanosomiasis (see below). When we saw the herd in July, 1904,
the twelve animals present, with one or two exceptions, seemed in
fair condition. In June, 1905, the ten animals then composing the
lerd were seen by Broden.4 They w'ere all in wretched condition
and three of them had trypanosomes in the peripheral blood. In
September, 1904, six heifers arrived from Dolo (near Leopoldville).
From October to June, 1905, there were eight deaths; all probably
from trypanosomiasis.

Yumbie is placed on a high grassy plain with neighbouring forest.
osstna pal pa l is and fuse a (?) are present. Large game is plentiful.

ix young cattle were brought to Basoko from Nouvelle Anvers
114 19 • I he herd now numbers eight (only two deaths from
accidents have occurred); they all seem to be in excellent condition.
ie ier grazes in cultivated ground surrounded by thick forest.

ssina palpal is present, but rare ; big game not plentiful.

2 35

LlSENGI. No deaths recorded ; cattle present for six years.
They graze in uncultivated coffee plantations surrounded by forest.
No buffaloes, many antelopes present, (i loss inn pal -pa l is not seen,
probably present.

At KiRUNDU there were a few miserable-looking trypanosome-
infected cattle, in the hands of the natives.

EALA, August, 1904. Before reaching Eala a report was received
from the Veterinary Surgeon resident there, M. Bertolotti, in which he
described an obscure disease affecting the cattle under his care. I he
symptoms resembled those of trypanosomiasis, and on examination
we succeeded in finding the parasites in one cow (I*ig. 0 in which the
disease was well advanced (no trypanosomes were found by a single
centrifugahsation of the blood of five other animals chosen from the
herd for their lack of condition). Tire extent ol the disease at IE ala is
shown by the following review of the cattle records.

There is at present a herd of 43 cattle at Eala. 1 hese represent
the remnants and product of 51 cattle sent here from the Lower
Congo in May, 1901, one bull and ten cows (of these only two cows
are still alive), and from April to May in 1902, five bulls and thirty-five
cows (of these one bull and 26 cows are still alive).

The great mortality amongst these animals has been almost
exclusively due to trypanosomiasis.*' Calves born at Eala (and at
Eouvelle Anvers) hav e suffered specially from this disease ; of 3b bom
at Eala only 14 are living (for example, of 1 5 bull calves born between
August, 1902, and January, 1904, 12 were dead in August, 1904, of
trypanosomiasis). Some of the young animals born here which are
stdl alive are undersized and obviously ill developed.

Histories of the Infected A nimals

1 OW No. 10. — Brought to Kala from the Lower Congo in 190-2- July *3. =

s » present extremely thin and scarcely able to move (Fig. ■>• Superficial glands
.1 e"ar6ed (M. Rertolotti regards this as one of the most constant symp o^
ne disease- • , . 1 Rlnnd ce

nertoiotti regards this as one oi me * ,

,n -diarrhoea is also a usual symptom). Blood centnfuga isc ,

****** seen. August s, ,904 Animal still thinner ; hardly able to stand
pa ,of weakness; but still has appetite and munches grass. No oe
puj^l haemorrhages observed. Blood centrifugalised and prescapula g
Pjj&ctured ; trypanosomes found by both methods. The animal was killed by
lna air into jugular vein and an autopsy was made at once. _ _

July. ^"'Cal diagnosis; the parasites were not observed in affected anima

236

Autopsy. — Animal is very thin ; muscles pale. Organs all normal, spleen not
enlarged. Superficial lymphatic glands twice as large as normal, waterv, not con¬
gested. Deep glands all much enlarged, about half of them (especially in abdomen
much congested — almost haemorrhagic. Examination for trypanosomes : Trypano¬
somes were found in the blood (jo to a coverslip preparation) only after centrifugalisa-
tion (five coverslip preparations of uncentrifugalised blood were examined without
result). 1 hey were also seen (three to a coverslip) in a preparation of gland juice
obtained from the prescapular gland by a hypodermic syringe before the animal was
dead; none were seen, however, in nine preparations of the juice of glands from various
parts of the body made within one to two hours of the animal's death. Many parasites
were found in the deposit obtained by centrifugalising slightly turbid fluid from
the pleural (6oc.cm. present), the peritoneal (500c. cm.) and the pericardial
(35 c.cm.) cavities. It was noted that the trypanosomes seen in these fluids were
almost invariably attached to a white cell. Forty-five c.cm. of cerebro-spinal fluid
taken immediately after death was centrifugalised and most carefully examined
It contained no trypanosomes and no red or white cells, but it was made slightly
opalescent by a peculiar flocculent material present in small quantities.

C<>\\’ N0. iot at Eala. August 23. 1904

4 noc motions. — Two rats (Fxn

this cow intraperitoneallv on t.,i» ,3~’ ■ CaCh received 4 c.cm. of blood

- - — -Neither had become infected* on At

examined, almost daily, i n 6 1 *a 1 • anima,s mentioned in this paper
nt‘l either the animal became infect rfer "l?*1 scJuare fresh coverslip preparat

ted or the experiment was discontinued.

237

3rd, so they were then both re-inoculated intraperitoneally, one with 5 c. cm. and
the other with 3c. cm. of a mixture of gland juice, from the same cow, and sodium
citrate solution showing one trypanosome to five fields.

A guinea-pig (Kxp. 142) was at the same time inoculated intraperitoneally with
4c.cm. of the same fluid. All three animals were carefully examined ; none ever
became infected.

The herd at Eala grazes on low-lying land surrounded by forest.
Glossina pal pal is 1 exists in small numbers (July, 1904). Big game
is scarce.

Nouvelle Anvers, August 15th, 1904. At Nouvelle Anvers
five of the thinnest of the 34 rattle present were examined ; two had
trypanosomiasis. The first cattle were brought here in 1896. Trom
then to 1900, 16 head (thirteen cows, three bulls) were imported from
the Lower Congo (two from the Canary Islands). Since 1898, 48
calves have been born ; ten have died (one from the results of abortion,
nine from fever and unrecognised diseases), and several have been
sent to other posts. Of the original animals twelve are dead (one
slaughtered, 11 of insufficiently described diseases).*

It is most important to note that seven of these animals came from
Stanley Pool (Kinchassa), and all the animals sent from Kinchassa
to Nouvelle Anvers have died in from one to four years after their
arrival. At present there is only one animal (Bull 18, arrived Match,

H, from Stanley Pool (Leopoldville) and it has trypanosomiasis ;
E Broden has shown trypanosomiasis to be common among cattle
ear Leopoldville.

Though it cannot definitely be said that the animals from Stanley
°°1 did not contract trypanosomiasis at Nouvelle Anvers, it is quite
°ssible that they may have been infected with trypanosomes before
i>eir arrival there. If this is the case the herd has contained infected
ninials since 1899 at least. Its losses since then have certainly been
ieavy, but the percentage of mortality is smaller and the disease seems
ess virulent than at Eala. For instance, one cow born at Nouvelle
Wersin 1896 and another brought there from the Canaries in the
ai»e year are still alive and well

J,TEE* ‘7 — Born here December, .902. In fair condition ; ^PP0^*0^
^lymphatic glands sligh.lv enlarged. t Fairly numerous trypanosomes

gland juice. ' _ _ _

;radu5ien ,he cause death is ill-defined, it is usually because the anima1 ha
- “aHy wasted and died. Such cases are usually ones of trypanosomiasis.
.rypan°"e of the animals in which a most careful examination failed o
in,es a*80 had slightly enlarged glands.

Bull 18. — Age, two (?) years. Imported from Leopoldville, March, 1904. In
good condition ; supposed healthy ; lymphatic glands slightly enlarged. Fairly
numerous trypanosomes seen in blood and prescapular gland juice.

Inoculations. — Rat (Kxp. 145) inoculated Aug. 15 intraperitoneally with ac.cm.
of blood from this animal ; did not become infected.

Monkey ( Cercofithecus schmidti) (Exp. 143) received 8c. cm. of blood intra¬
peritoneally, August 15, 1904. Trypanosomes appeared in its blood September 22.
I hey soon disappeared, and were only occasionally seen during 1904. During 1905
the monkey seemed quite well, and trypanosomes were not seen in its blood, which
was examined at intervals. On January 1, 1906, it was very ill, weak, dull and
apathetic. A single degenerated (?) trypanosome was seen in two coverslip
preparations. Animal died next day ; immediate cause of death was miliary
tuberculosis. #

A rabbit (Exp. 1441 received 3 c.cm. of blood intraperitoneally; never became
infected ; under observation for six months.

I he herd at Nouvelle Anvers grazes for the most part on cultivated
land. Big game is not very abundant, but biting flies of several sorts
(G lossina pit l pal is, Stomoxys and tabanids') are present.

Romee, September 12th, 1904. One or two animals now present
have been here since 189;, but the great majority of the cattle19 have
been brought here since the end of 1902. Most of them are of Lake
Kivu stock and came here by way of Nyangwe. A few head came
from the Nile. Only nine deaths have occurred since March, 1903 ; all
weie cattle which had arrived in poor condition two or three months
previously, it therefore appears that this herd does not lose so many
animals from disease (trypanosomiasis ?) as do those at Eala and
Nouvelle Anvers. For example none of the calves born here (first
m l899) have died- We were shown ten animals that had been kept
apart from the herd of 25 head because of their lack of condition.

■ x A them (anived at Romee, July, 1904) were examined.* Four
, tryPanos°niiasis ; three of these were decidedly thin and had
enlarged lymphatic glands, the fourth had no big glands and was
comparat.vely well-fleshed. All had temperatures of over iori“F.

ur in ected were of Kivu stock ; two had come from Stanley Falls
and the other two were from Nyangwe.

Three rats (Exp. i3J
Cow No. 2 became infer
2i clays.

M9. 151) inoculated intraperitoneally with blood from
»n three, six and eight days and died in 18, 23 and

**n July, 1
the other three

905, one of these animals
had been slaughtered.

was

still living and seemed perfectly well-

239

Two guinea-pigs (Exp. 148, 152I inoculated in the same way, one from one of
the above rats, the other from the cow, showed parasites in twelve and fifteen
days and died in 18 and 23 days of ruptured spleens (much enlarged and friable),
in one case after slight violence.

A rabbit (Exp. 147), weight 51b., became infected in nine days. Parasites
were at first numerous ; they then became scanty and were only seen from time to
time until January, 1905. None were seen after this. Death from pneumonia
October 21, 1905. Spleen not enlarged, firm (5 x *8 X *6 cm.), capsule thickened.
None of glands much enlarged. Animal very fat.

Monkey [Cercofithecus ?> (Exp. 146), weight S«3 lb. Inoculated September 13,
1904, with blood from one of the rats ; showed trypanosomes in its blood three and
six days later and never again. Death from pneumonia December 16, 1905.
Spleen slightly enlarged |5xix i*scm.), firm ; glands not enlarged ; animal fat.

The herd grazes on high, wooded, cultivated grounds. Small
antelopes are not uncommon. G loss inn palpal is is rare.

Lokandu, October 2 1st, 1904. Sixteen head of cattle present.
In January, 1902, there were 23; since then 13 have been sent away
(nine to Romee) and three have died. In only7 two of the deaths wire
symptoms resembling trypanosomiasis present.

We examined eight heads; five had trypanosomiasis. Ihret, all
infected, of those examined were calves born here. All were thin, bad
dry noses, much-enlarged glands (to 3 by 4 by 10 cm.) and slight
temperatures (ior5°F.).

In two of them there were distinct patches of oedema in the loose
skin of the throat. Five of those examined were adult cows imported
from Nyangwe or Kasongo. All were in fair condition ; only one had
enlarged glands. This one, and one other in whom theie was no
glandular enlargement, had trypanosomiasis. The one with enlarge
glands had a temperature of io6‘8°F. Trypanosomes were seen in
small numbers in the blood of only three of these five infected anima s.
They were seen in much larger numbers in the gland juice in eac 1
case.

Inoculations. — A guinea-pig (Exp. .S3) was inoculated intraperitoneally on

October 22, witJ - blLl, from one of the calves contamxng^hree

■ Panosomes to a covcrslip preparation. It was examined careful \ J

'905' but trypanosomes were never seen.

These cattle had scanty pasture in coffee plantations sutroun
h} forest. Antelopes and Glossina pal pal is not uncommon.

Sendwe, November 5th, 1904. Four head of cattle vvere
u re from Kasongo in 1900. The two bulls imported die soon
1 e'r arfival, so the remaining animals were sent away to iV ata p

24 o

In May, 1904, eight head passed through on their way to Romee.

Two died here; only four readied Romee alive (we examined none
of them for trypanosomes). In May, 1904, ten heifers on their way
to Romee arrived here by Kasongo from Lake Kivu. They were not
in good condition ; but this was at the time explained by a two months'
stay at Kibombo poor pasture. A month after arrival they
commenced to die, and in three months five were dead. All had the
same symptoms, “gradual wasting and loss of vigour.” The
remaining animals seemed in fair condition, but all of the four
examined had enlarged glands (see Fig. 2) and temperatures (to
1044 F.). Three of them had trypanosomiasis. Some of them were
later sent to Lokandu.

•1G. 2.— Calk at Sendwe; to show enlarged gland.

it pasture at Sendwe is poor; Glossina pal pal is and big game
are plentiful.

fin 1 November 14th, 1904. A herd, now numbering about

’ haS been here since the end of the Arab war (1894)- is

24I

largely made up of captured cattle, the majority of which probably
came from the neighbourhood of Lake Kivu. 1 he cattle are at
present in splendid condition. Since 1900, 45 calves have been born.
Dunng the same period there have been only five deaths, none of them
from any disease resembling trypanosomiasis.

The country surrounding Nvangwe is high, grassy plain. 1 he
herd never grazes further than one and a half miles from the post
and never goes near a swampy area towards the South. 1 he countiy
about Nyangwe is thickly populated and extensively cultivated. There
is, therefore, little large game in the neighbourhood. Tsetse flies
were not seen anywhere about the post nor on the plains wheie the
cattle graze. They were seen, however, along the river bank a mile
and a half further South.

The blood of 20 cattle chosen because they were less plump than
the remainder of the herd was examined ; gland puncture was done
m 17 in which the glands were sufficiently large. I rypanosomes
were not found.

Kasongo, November 25th, 1904 April 2/th, i9°5- rliere 1S a
herd here of about 260 cattle. Many of them were collected by the
Arabs; additions have been made by the whites. The animals
reproduce well and the herd has been largely increased by local
births ; deaths from diseases are very rare. Its original members
were probably largely from Lake Kivu. Some probably came fiom
the Eastern and Southern shores of Lake 1 anganyika. A ei > ^ev'’
any, were ever brought directly from Katanga, far to the South, or
from Lusambo, in the West.

One hundred of the cattle were examined for trypanosomiasis

during our stay at Kasongo, five were found to be infected. All fi'e
"ere found in the first twenty-two examined. 'I Ins is noteworthy,
Slnce the thinnest animals were taken first. All the animals weie
examined in the same way and each was gone over but
Preparations of gland juice and blood were examined and the
temperature was taken in each case. .

The infected animals were kept under observation for se\
months. The rareness with which parasites were seen in them 1 g
th,s Period (see charts) strongly indicates that many cases remained
^detected by our examinations, and therefore that the proportio

cahle infected is much more than five per cent.

Zilani No. 8 (Nov. 15, 1904). — A young cow, born at Kasongo. She is rather
thin and the natives say she is ill ; but there is no oedema, staring of the coit,
&c. Largest prescapular gland uX5™-, largest femoral gland 9x3cm. Try.
panosomes were found in gland juice, not in blood (see Chart Ii. Inoculations
(see below, page 243).

Moiuma No. 33 (Nov. 25, 1904). — A young cow, born at Kasongo. Apparently
quite healthy. Prescapular gland 12 x 5 cm., soft; femoral 6 x 3 cm. Trypano¬
somes present in blood and gland juice. The course of the disease is given by
Chart II up to l-ebruary 8. From then to April 19 (observations ceased) the
temperature varied, without any sudden changes, between ioi°F. and io4°F.
Blood and gland juice were examined at intervals ; parasites were seen during this
period by gland puncture, only on March 16.

Inoculations. Monkey (Cercofithccus schmidti ) (F.xp. 173), January 11, 1905.
noculated intiaperitoneallv with r8 c.cm. blood showing one trypanosome to cover:
tinder observation until I-ebruary 8 (when inoculated with spirochaetesi : never
infected.

Keowkosiko No. 60 (Nov. 25, 1904). — Cow, born here. It is in fair condition,
per aps slightly thin, l’rcscapular gland measures 10 x5cm., femoral 8 x ;
loth are soft. ( hart III shows the course of the disease up to February 17. From
then to March 9 the temperature continued in the same way and parasite? were
on y seen on March 8, q, 10. I he temperature became irregular and rose constantly
until 105 F. was touched on March 22 (no trypanosomes in blood). The tempera-
ure then became lower and no parasites were seen in blood or gland juice up to
pri 1. , when they were in the blood in small numbers. Observations ceased.

1. 1 boko No. 29 (Nov. 30, 1904).— Fair condition ; glands slightly enlarged
lypanosomes present in gland juice, absent from the blood. As is seen by Chart
, the parasites were never again seen in this animal.

hpp \°L0 ,No- 9 (Dec' *904, Fig. 3). — A steer, adult, born at Nyangwe, has
n nere for three years. Prescapular gland is 15 x Scm.* and soft; femoral is
ii X 2 ( niM so t and mm eable. I rypanosomes were seen in gland juice and not in the

theatn, CO"?e °f !he disease UP to April 14, when the animal was killed for
the autopsy, is indicated on Chart V.

cxaminatinn ThC aninial Was kiIled by Ejecting air into the jugular vein. The
o m l"'3,5 fenced immediately after death. No subcutaneous fat 01
vessels v* l!SC •!, datk' Abdomen contained 400 c.cm. clear fluid; mesenteric
Dleurae rmo ’ °",en,um and retroperitoneal tissues oedematous. Thorax

of nericardi ",d’ Per*cardium 300 c.cm. Between diaphragm and base

aLut base oT i ! 3 .ot oed^atous tissue , cm. in thickness. The fa.

and was vm- 1<? T 3nd a,ong ils 'essels was brownish-canary-yellow in colour
mediastinum °e ematous, as was also the connective tissue of the posterior
weight i'-' ih ° it " * e organs showed any striking macroscopic change (spleen
;; \’fSUbs ancLe Marrow of long bones yellow. Lymfhatic

by a can' . fefor! df,ath the prescapular gland was excised. It was surrounded
All the glands °e( en,atous connective tissue 75 cm. thick; it measured nx6crn

fluid followed tlu^kTife^' l hTl "’T bU‘ dis,inc,1-v “ watery,” so that the
with darker „ w c he arger 6,ands m all parts of the body were rale

destine at the h«s‘ ^ °f the Sma,ler (-5 x . cm.) along the small

( x . e v 1 /m \ ...„. M ,e ,nesentery (2x3 cm.) and about the bronchi and trachea

_ •> e congested or haemorrhagic. Immediately after death lumbar

the capsule ^^This eV-md wh'6" f°r these glands are of course estimates, am
P me. 1 h,s gland when excised on April ,4, ,905, actually measured .

and incl

IX1

243

puncture was done at about the sixth vertebra with a four-inch needle. About
4C.cm. of clear fluid was obtained. Fresh preparations of the following organ
juices and body fluids were examined, juice from the prescapular, mesenteric and
tracheal glands, liver, spleen, marrow from long and short bones, blood, pericardial
and cerebro-spinal fluid. Parasites were only seen in the gland preparations.

Inoculations. — Dog* (Exp. 22 7), April 13, 1905. Inoculated intraperitoneally
with about 150 c.cm. of defibrinated blood from the jugular vein. Died December
12, 1905; no autopsy. No trypanosomes seen.

Dog (Exp. 229), April 13, 1905. Inoculated as above with 92 c.cm. of blood.
No trypanosomes seen until December 1, 1905, when killed because of mange.

Dog (Exp. 229), April 13, 1905. Inoculated as above with 150c. cm. blood.
Killed because of mange, December 1, 1905. Trypanosomes were never seen in the
blood.

1 ,G- 3— Steer, Yolo, at Kasongo. Note emaciation. The type of horns
and tendency to hump indicate a Lake Kivu origin.

Experimental Inoculations at Kasongo
Mosl of the animals inoculated at Kasongo were so frequently
noculated that it has seemed best to treat the majority of them

U-!le,^eparatejteadin? _ _

Tlle d0gs used in all these experiments were native ones bought at Kasongo.

244

Rat (Experiment 132), July 6, 1904. Inoculated with buffalo blood (Buna mi);
never infected. (Exp. 221) April 3, 1905. Inoculated with 5 c.cm. blood from Yth
showing four trypanosomes to cover. Died August 20, 1905 (no autopsy). Never
infected.

Rat (Experiment 133), July 6, 1904. Inoculated with buffalo blood; never infected
November 8, 1904, re-inoculated (8 c.cm.) with buffalo blood. (Exp. 222), April 3,
1905, re-inoculated with 5 c.cm. blood from Yolo showing four trypanosomes to
cover. September 8, cataract in one eye, blood in anterior chamber of the other.
December 8, trypanosomes first seen in blood. December 15, died, blood
swarming with trypanosomes. Glands and spleen enlarged, organs otherwise
healthy.

Rat (Experiment 134), July 6, 1904. Inoculated with buffalo blood; never infected.
(Exp. 177) January 11, 1905, re-inoculated with 12 c.cm. from Moiuma showing one
trypanosome to a cover j never infected. (Exp. 223) April 3, 1905, re-inoculated
with 5 c.cm. blood from Yolo showing four trypanosomes to a cover; accidentally
killed November 4, 1905. No trypanosomes nor signs of trypanosomiasis seen in it
or in a rat sub-inoculated from it.

Rat (Experiment 135), July 6, 1904. Inoculated with buffalo blood; never infected.
(Exp. 224) April 3, 1905, re-inoculated with 5 c.cm. blood from Yolo showing four
trypanosomes to cover ; never infected. Died of skin disease, October 13, 1905.

Rat (Experiment 139), July 27, 1904. Inoculated with 8 c.cm. blood from antelope;
never infected. (Exp. 169) December 27, 1904, re-inoculated with 7-5C.cm. of
mixed blood and citrate solution from Moiuma showing four trypanosomes to
<.over; never infected. March 9, 1905, re-inoculated with <;c.cm. of blood from
Keowkosiko. March 21, 1905, re-inoculated with 5 c.cm. of blood from Zilani.
April 14, died from mechanical effects of inoculation with large amount of blood;
no autopsy ; never infected.

Rat (Experiment 140), July 17, i9o4. Inoculated with 8 c.cm. of blood from ante-
ope; never infected. (Exp. 168) Dec. 27, 1904, re-inoculated with 7-5 c.cm. of blood and
citrate solution from Moiuma showing four trypanosomes to a cover; never infected,
larch 9 1905, re-inoculated with 5 c.cm. blood from Keowkosiko. March 21, I9°5>
re-inoculated with 5 c.cm. blood from Zilani. April 3, 1905, re-inoculated with
5 c.cm. 00 from Y olo. April 13, 1905, re-inoculated with 5 c.cm. blood from
Yolo. Never infected* up to September 19, 1905, when it escaped.

• „ Rat (Experiment 170), December 27, 1904. Inoculated with 2-5 c.cm. blood show-

°Ur ryPan°somes to cover from Moiuma ; five trypanosomes to cover were
N ns nr.ooD next day and not again. March 9, 1905, re-inoculated with
bloid'w 7°v K.eowkosiko- March 21, 1905, re-inoculated with

ttisfp \ ant' pril 13, 1905, re-inoculated with 33 c.cm. blood from Yolo.

DieH1 l™YPANOSOMES TO A FIELD were seen in its blood on April 21, and not again.

ruary r9°6 ; no trypanosomes seen at autopsy; all organs normal save
spleen which was slightly enlarged and fibrous.

F^/UAnr;i(EXPERIMENT 2I9h April >905- Inoculated with 10 c.cm. of blood from
re inomlaf re'lnoculated with 10 c.cm. of blood from Zilani. April 13, i9°5-

at Bom* T I25C-cm. of blood from Yolo. Under observation until stolen

at Roma. Trypanosomes never seen.

iaocJS^L/^^ schmidii) (Experiment 220), very young. April 3, «9°5>
blood from 7 7 ' ? c’“n‘ klood from Yolo. April 5, 1905, re-inoculated with 5 c.cm-
Died of ! Pn I3, I9°5, re-inoculated with 12-5 c.cm. blood from Yolo-

of trypanosorniasL ^autopsy0* ' ParaSitCS neVCr Seen- No fU’panosomes or signs

See charts for number of parasites present in blood inoculated.

1

245

Through the kindness of Commandant Verdick the four infected
cattle which remained when we left Kasongo were kept apart from
the herd at Kasongo. After our departure the animals were left quiet,
and there was a consequent amelioration in their condition. While
we were there they were driven twice daily from their kraal to our
laboratory and back — a distance of about one and a quarter miles.
The fatigue of these journeys and the time last through them from
grazing had certainly an adverse influence. Moiuma and Keowkosiko
both calved. All the animals were in excellent condition in
September, 1905. They were then sent to Lokandu, where there is a
great deal of cattle trypanosomiasis and where Glossiua palpalis is
far from rare, in order to ascertain whether they would resist exposure
to re-infection. (It must be remembered that the pasturage at
Lokandu is not nearly so good as at Kasongo). The steer, three cows
and two calves reached Lokandu on November, 23rd, 1905, in
excellent condition. A calf died in December after twenty days of
manifest diarrhoea. One of the adults died after a “ slow wasting
and great weakness early in 190C. In August, 1906, two of the four
remaining animals were very thin and seemed ill ; the remaining two
are still under observation. The Chef de Secteur at Lokandu, in
conveying this information, reports that the “ health of the herd at
Lokandu is excellent.”

The cattle at Kasongo graze on a wide plain intersected by several
dreams which have occasional clumps of bushes along their banks.
,J/ossina palpalis are very, very rarely found about the grazing
§rcmnd, and large game practically never comes near it.

Tshofa, May 15th, 1905. There was formerly a herd here of
*ome fifty cattle, which was intended to be used for supplying
ullocks for work on the waggon road between Tshofa and Pania
Tutombo. As the animals were not doing well, all, save fourteen
Whlch Were obviously ill, were sent to Lubefu. Of that fourteen only
e'ght remain ; two were killed by a leopard, four died of a chronic
1Sease characterised by general loss of condition and constant
’arrhoea, with straining and small watery stools ; there were no
°,e emas nor lack of appetite. Between June, 190b and I9°4’

aA,°Ut thirteen animals were said to have died here of this disease.
‘ 'nost all the calves bom or brought here have died of it. and the

246

only animal present which was bom here is affected by it (trypano¬
somes seen).

The majority of the cattle brought to Tshofa came from Lusambo.
Cattle have been sent from the Tshofa herd in 1902 to Kisenga and
in 1904 to the Tanganyika, Moero and Lubile districts. Three out of
five animals whose gland juice and blood were examined had
trypanosomiasis.

1 here was good pasture in open country at Tshofa, but Glossm
pal palis and large game were very plentiful.

Cabinda, June 3rd, 1905. Before the advent of Europeans, some
of the important chiefs in this neighbourhood had small herds of ten
to twenty cattle. Cattle did not do well ; many died of diarrhoea, and
there were no large herds. At present a few head are kept here for
vaccine making. They come from Portuguese West Africa by way of
Angola and from as far South as Dilolo. We are told that the Dilolo
natives buy cattle from the Zambesi valley. The blood and gland
juice of six animals were examined once ; trypanosomes were not
seen.

Lusambo, June 21st, 1905. Nearly all the Lusambo cattle come
by way of Malange and Luluabourg from Portuguese West Africa;
formerly herds of a hundred head were brought in by this route to be
exchanged for slaves. It is said that the animals were used for food
and were not kept for breeding. The records of the Lusambo cattle
are very badly kept,* but deaths from disease seem to have been
rare among them. Six of the thinnest animals were chosen and their
blood and gland juice were examined once. Trypanosomes were seen
in one of them. About six cattle were sent from this herd to Kutuon
ake Leopoldville II in 1902-1903 ; all died within a short time. It
is said that the cause of death was trypanosomiasis.

cattle was also inadeonat* USTual one- The system of brands used for marking
animal or to learn was consequently often impossible to iden .

to estimate0 thTincfde^ce of °^ i£ Was often impossible at a

infected animals ft ic' j disease in the past or to determine the ori0
through the institution fV1(*en) that much valuable information would be 0
8 institution of careful systems of recording and identifying cattle.

247

III. OBSERVATIONS ON TRYPANOSOMES IN OTHER DOMESTIC

ANIMALS

Horses, Mules and Donkeys : Comparatively few of these
animals have been brought into the part of the Congo Free State
visited by us. Some were seen in apparently good condition at the
following places: various localities in the Lower Congo, Leopoldville,
Eala (three horses and a foal from Dakar present for fifteen months,
not worked), Coquilhatville (donkeys present four to five years),
Bamamia, four donkeys (one present for eight years), constantly worked.
Nouvelle Anvers, a few horses here were in good condition when we
saw them in 1904 and were still well in June, 1905. Umangi (three
horses, a mare from Teneriffe present for five years), Lisengi (two
horses present for six years). At Nyangwe there is a small drove of
donkeys. None of the above animals were examined. At Kasongo
no trypanosomes were seen in two mules and a donkey ; these three
animals had journeyed considerably in the Eastern part of the Congo
Free State.

In July, 1904, Commandant Sillye bought two stallions near Dakar in Senegal.
They were then in excellent condition, and were six and twelve years old. He
brought them to the Congo and took them on a hard journey from Kasongo almost
to Lake Tanganyika ; from this journey they returned unloaded to Kasongo, where
we saw them on January 14, 1905, after they had had a fortnight’s rest. Both
animals were thin, but this was thought to be due to fatigue and bad pasturage.
There was (Fig. 4), as in the Gambian horses (6), slight abdominal fulness without
definite oedema. Both had rough coats from lack of grooming and were infected
with ticks, fleas, chiggers, trombidium larvae and numerous bots. In addition both
showed the weals of many recent fly-bites. If the animals were left to themselves
they dropped their heads and certainly seemed to lack vigour.

The temperature of both was about ior°F. and their glands were slightly enlarged.
The only sign of oedema ever noticed in either horse was a swelling of the scrotum
in one in October, 1904. The conjunctivae were pale and watery in both.
Trypanosomes were present in good numbers in the blood of both horses. lhe
animals were relieved of the bots, chiggers, ticks and trombidia (lotion of bichloride
of mercury) and were given arsenic in the form of Fowler’s solution. At the en
of January they reached Romee, where they were turned to pasture and the arsenic
was discontinued. For two months both put on flesh. Then the older one ecame
rapidly thinner and very weak ; the appetite remained good. It died pn 17,
1905, after a very profuse perspiration (6). On September 14, 1905, the remaining
borse, though comparatively thin, seemed in fair health and fed well. e scro urn
was often swollen.

Inoculations.— Rat (F.xp. 180). Inoculated January 14, 1905, intraperitoneally
with a mixture of blood, from the older of the two stallions, and citrate solution
showing 15 living trypanosomes to a cover. Parasites appeared in its 00 nex
day and were never absent until it was accidentally killed on March 21.

Autofsy. — Spleen much enlarged (6-25 x i*5 x *75 crn-)» so^> friable , glands all
slightly enlarged.

248

Rat (Exp. 181). Inoculated as in Exp. 180. Parasites appeared January 1:
constantly present till it died of the disease September 24, 1905.

Guinea-pig (Exp. 179). Inoculated as in 180; not infected. March n,
re-inoculated with 1*50. cm. blood from Exp 180 showing 30 trypanosomes to field ,
infected in five days. Parasites constantly present till death from sunstroke. Jrai
18, 1905.

Autopsy (immediately after death). — Dependent part of abdominal wall fin,
water-logged ; vulva oedematous ; spleen soft (5 x 2 5 cm.) j glands not enlarged
Animal well nourished.

Guinea-pig (Exp. 178). Inoculated as in Exp. 180; not infected. Re-inoculated
as in Exp. 179. Trypanosomes in blood in six days, and constantly present rati
death from the disease on September 17, 1905.

F 'G' 4‘ Staluon infected at Kasongo. Note slight fulness of abdomen.

A donkey about seven years old, also belonging to Commando:
Sillye, was examined on January 14th, 1905. This animal probably
came originally from Uganda. Since 1902 it had been constant-,
worked and had just returned from the same journey as the horses
* W.aS yery fat and apparently in the very best of health. A few 5®a-
a po e like trypanosomes were seen in its blood. Although ha c

249

worked it remained in apparently robust health until the end of June,
1905, when it was accidentally killed.

Goats seem to thrive almost everywhere in the Congo.

Sheep are present in few places, but they usually do well.

None of the score of goats and sheep examined by us were
infected with trypanosomes. Broden reports trypanosomiasis in sheep
at Leopoldville.3

IV. TRYPANOSOMES IN BIG GAME

Mswata.

Antelope (adult Tragelaphus scriptus), very scanty stumpy
trypanosomes in blood ; retro-peritoneal glands deeply haemorrhagic,
otherwise apparently healthy.

Rat (Exp. 130), June 26, 1904. Inoculated with 3'SC.cm. blood. Trypano¬
somes seen in its blood only on July 5. Died (cause, exposure ?) Juh 24 , no
trypanosomes or signs of trypanosomiasis at autopsy.

Tshumbiri.

Buffalo (Bos nanus, two adults). Both fat, and save for a caseous
retro-peritoneal gland in one, apparently healthy. Xo trypanosomes
found in blood.

Four rats (Exps. 132, 133, 134, 13S) (see Inoculations at Kasongo, page 243)
were inoculated with from 5 to 4 c. cm. of blood ; none of them became in ectec .

COQUILHATVILLE.

Antelope (young Cephalophus dorsalis ?), no trypanosomes in
blood.

Inoculated two rats (Exp. 139, 140) (see Inoculations at ICasongo); never
became infected.

Stanleyville.

Antelope (adult Cephalophus dorsalis ?). Gland juice and blood
examined ; no trypanosomes seen. Prescapular gland, enlarged f,
4 by 3 by 2 cm., cortical congestion of it and retro-peritoneal glands.

LOKANDU.

Two antelopes (Cephalophus dorsalis ?) gland juice and blood
examined ; no trypanosomes seen. 1 he blood and gland juice of an
adult buffalo were examined and with its blood a rat (Exp. 13 3) "as
re-inoculated ; trypanosomes were not found.

Kasongo.

Three antelopes (adult males, Tragelaphus scriplus).

1. Shot four hours previously, no trypanosomes in gland juice or
blood. Prescapular gland 6 by 3 cm., iliac glands haemorrhagic

2. Shot three hours previously. Trypanosoma theileri in blood.
Inoculated rabbit and two guinea-pigs with blood ; none became
infected with any trypanosome.

3. Shot three hours previously ; no trypanosomes in gland juice,
many in blood. Abdominal glands much congested and enlarged.

Inoculated prom Antelo-pe No. 3. — Guinea-pig (Exp. 188). Inoculated January
28, 1905, with 1-5 c. cm. of blood showing three trypanosomes to field; not infected.
April 19, re-inoculated with 2 c. cm. mixed citrate solution and blood from Exp. 189
showing one trypanosome to field ; June 19 infected ; trypanosomes were constantly
present until animal’s death from pneumonia on August 30, 1905. Spleen
3x i'Sx'75 cm. normal ; glands congested, not much enlarged.

Guinea-pig (Exp. 189). Inoculated with 2-5 c.cm. of blood as in Exp. 188;
infected February 6, parasites constantly present in large numbers until death from
pneumonia, August 30, 1905. Spleen large (6 x 3 x **5 cm.), soft, friable, congested;
glands enlarged and congested. No trypanosomes were seen in blood just before
death or in organ juices ; present in gland juice.

Rat (Exp. 190). Inoculated with 2-5 c.cm. blood as in Exp. 188; not infected.
April 19, re-inoculated with 2-5 c.cm. mixture as in Exp. 188; April 25, trypano¬
somes present, but were not again seen. Animal escaped June 19, 1905.

Rat (Exp. 191). Inoculated and re-inoculated (April 19) as in Exp. 190; infected
May 1. Parasites constantly present until it died of pneumonia September 19, 1905-

Young guinea-pig (Exp. 230). Inoculated April 19, 1905, with 2 c.cm. blood
from Exp. 189 ; infected May 6. Many parasites constantly present until death
from the disease, July 6, 1905.

On the journey from Kasongo to Lusambo the blood of the
following animals was examined by cover-slip preparations : four
Tragelaphus scriplus , one Can is anthus , six Hippotragus equinus,
one Bos nanus (ten antelopes, a jackal and a buffalo). Trypanosomes
were seen in none of them.

V. MORPHOLOGY OF THE TRYPANOSOMES

(a) Trypanosomes of cattle

The trypanosomes seen in stained preparations of blood from the
cattle, horses, antelopes and their sub-inoculations, are so nearly
identical that one description will suffice. Slight variations and
peculiarities will be mentioned separately.

1 hree types of trypanosomes were found corresponding closely to
the description of T rypanosoma dintorphon as given by Dutton and

25J

Todd.6 The first of these is a very short form resembling the
"tadpole” form. It measures between 8 and 14 /j in length (most
commonly 11 to 12-5) by from O’g to 3-4 fx in breadth (average 1-5). In
this type the trypanosome is widest slightly posterior to the nucleus,
and then gradually tapers towards the anterior end. The posterior
extremity presents all gradations in shape from acute to rounded ;
most commonly, however, it is rounded. 1 he anterior extremity is
effilated. The blepharoplast is small, round or oval, and is situated
close to the posterior end and at one edge of the trypanosome. In
most of the parasites the undulating membrane could not be seen ; in
those in which it was visible, it was extremely narrow. 1 he thickened
edge of the undulating membrane is poorly marked and in all cases
ends abruptly with the anterior end of the trypanosome. A tree
flagellum is absent. The nucleus lies in the middle third, and as a
rule occupies the whole width of the body. It is oval, and either stains
quite homogeneously or is composed of a varying number of darkly -
staining chromatin granules (5 to 17) arranged in no definite manner.
These granules were not joined by connecting threads as far as could
be made out. In only a very few cases could a structure suggest ing a
karyosome be seen. The cytoplasm stains a light blue and appears to
be quite homogeneous. It was usually devoid of granules, but in some
cases contained them.

The second type is allied to the “stumpy" form.6 It measures
from 15 to 20/x in length (average 16/x) and from i'5 to 2'5/x in breadth
(average 2 /i). It also lacks a free flagellum or, at most, has an
exceedingly short one. In other respects it does not differ materially
from the description given above.

The third type resembles the “long" form.6 It measures from
20 1° 30jn in length (average 23 to 27/x) by from i’2 to 2/x in bieadth
(average 1-5 tor7/i). The nucleus is rather elongated in the direction
of the long axis of the body and as a rule stains quite intensely and
homogeneously. This form has a fairly long free flagellum which
measures from 2-5 to 6'5/u in length. The posterior extremity is
^her acute and the blepharoplast is situated farther away from it
lhan in the other two types. In other respects it is comparable to the

Uvo °ther varieties.

Eala.

The trypanosomes observed, with one exception, are of the long
type, with a well-defined free flagellum. The exception measured
I4M in length by 9/x in breadth, and conforms in all respects to the
“ tadpole ” type. In these parasites the blepharoplast was of
comparatively large size. Parasites of the “ long ” type in the process of
longitudinal division were seen in preparations of the peritoneal fluid.
In fresh preparations this trypanosome moved so rapidly that it was
extremely difficult to keep it in the field of the microscope. By its
rapid movement this parasite recalls the description of Trypanowu
vivax by Ziemann.20 We are, however, satisfied that it is the “long"
form of I rypanosoma dimorphon.

Nouvelle Anvers.

The parasites conform to the description of the “ tadpole " variety.
One trypanosome of the “ stumpy ” variety, measuring I9'2 by i'9/r,
was also found. In the one sub-inoculation the trypanosomes did not
show any changes.

ROMEE.

The parasites are of the extremely short “ tadpole ” variety. In
the direct sub-inoculations the parasite preserved the same
characteristics with considerable constancy. In two cases (Exps. 148
and 1 51), in addition to the very short forms, trypanosomes
corresponding to the “ stumpy ” forms made their appearance. The
same holds good with regard to the second passage. This fact is of
some importance. Dividing forms of the usual longitudinal type were
observed in all the sub-inoculations.

Lokandu.

1 rypanosomes of the first two types (the “ tadpole ” and “ stumpy”
forms) were found in these cattle. The short “ tadpole ” forms were
very much more common than the “ stumpy ” type.

Sendwe.

The parasites agree in all respects with the “ tadpole ” type. The
examination was rather restricted, however, as the parasites were very
scanty and only a few preparations were available.

Kasongo.

In the cow, Moiuma (No. 33), the trypanosomes are of the third,
the “ long,” type, and possess a well-marked flagellum. In many of

253

these parasites a large vacuole was seen immediately in front of the
blepharoplast. In this parasite the blepharoplast was comparatively
large, and recalls the trypanosome seen at Eala.

In the other cattle, Zilani (No. 8), Yolo (No. 9), and Keowkosiko
(No. 60), the trypanosomes are of the “tadpole” variety. In Yolo,
however, one parasite was seen of the “ long ” type with free flagellum
and comparatively large blepharoplast. I he further history in sub¬
inoculations of the trypanosome derived from Yolo is of great
importance in its bearing on the identity of these cattle trypanosomes.

In the rat (Exp. 222), inoculated from Yolo, all three types were
found. Of these the “ stumpy ” type, measuring from 15 to 1 8/x in length,
was most abundant. In the subsequent passage this differentiation
became more pronounced. The very short forms, measuring up to
14/4 in length disappeared, but the other two types persisted. The
num6er of “ long ” forms increased with the progress of the disease,
as was noted by Dutton and Todd in the case of 7 rypanosoma
dirnorphon (°, page 37).

The remaining two types can still easily be distinguished in
experimental animals. The “ long ” form is more plentiful, is slender,
and has a rather acute posterior extremity. The flagellum is fairly
long and is actively motile, whipping from side to side with great
vigour. The movements of the whole trypanosome are active and
progressive. The trypanosomes of the “ stumpy ” type are rather broad
^d have a blunt posterior extremity. They move slowly, twisting
round on themselves, then straightening out again, and so on. As a
mle they maintain the one position and do not move from place to
place. The undulating membrane moves sluggishly, and there appears
to be no, or, at most, a very short, free flagellum. The body of the
trypanosome is itself contractile. When buried between corpuscles
{be movements of both forms become appreciably slower.

(t>) Trypanosomes of the Kasongo stallions

In the younger horse the parasites were identical with the
tadpole” forms as observed in the cattle. Unfortunately, no su
ln°culations were made from this animal.

In the second horse, “ Toul,” the trypanosomes were of the secon ^
‘stumPy”) and third (“ long ”) types as seen in the cattle. 1 he long

254

forms with free flagellum were in the majority. In both types the
cytoplasm often contained small reddish granules, which were most
frequently observed in the posterior half of the body. In some thirty
sub-inoculations with this strain the trypanosomes of both types have
been constantly present. At the inception of the disease the “stumpy
forms are much more abundant, but as the affection proceeds the
“long" type assumes the preponderance. In the monkey sub-inocidaUi
a few of the “ tadpole” forms were observed.

In fresh preparations the two types can be very easily distinguished.
The long, slender form has a long flagellum and moves very rapidly,
lashing in and out among the corpuscles. It progresses for a short
distance, then stops, and in a few moments moves on again. It
often moves fairly rapidly' across the whole field. Occasionally it
adheres to a corpuscle, and after twisting and lashing about for a short
time breaks free and moves off. While the parasite progresses with
the flagellar end in front as a general rule, it occasionally moves
backwards for a short distance. The second form is broader and
shorter than the first, and is much more sluggish in its movements.
I hese consist chiefly of doubling and twisting of the trypanosome
on itself. 1 1 does not progress far, but remains pretty constantly in
the one spot. The undulating membrane and flagellum are very
poorly developed in this variety. In contradistinction to Dutton and
rodd,b dividing forms, of the usual longitudinal type, were seen in
this “ stumpy ” type.

(c) Antelope trypanosomes

In the antelope ( Tragelaphus scriptus) killed at Mswata
trypanosomes corresponding to the “tadpole” type were seen, but
were very scanty. In the one sub-inoculation the parasites retained

this type. Dividing forms of the usual longitudinal type were
observed.

In the antelope at Kasongo the trypanosomes were of the second
( stumpy ) and third (“ long ’ ) varieties as found in the cattle. In
the sub-inoculated animals these two types persisted; the stumpy
orms being in the majority in the preparations examined. It was
impossible to study this parasite fully, since the animal in which it was
rought to England was accidentally killed and the strain was lost.

255

VI. ANIMAL REACTIONS OF THE TRYPANOSOMES

(a) Trypanosomes of cattle

The slight infectivity for laboratory animals of the trypanosomes
in the majority of the cattle found infected is very striking. It is only
at Roraee that experimental inoculations were at all successful. No
morphological variation in the parasites was noted to accompany this
difference in virulence. All the parasites seen agree with the types
already described. The results of inoculation of various laboratory
animals with the trypanosome obtained from Yolo , Steer 9 at Kasongo
through rat (Exp. 133) are briefly given.

Monkeys (2).

Two monkeys, a Macacus rhesus and a Cercofithecus sp ? were inoculated
intraperitoneal ly with small quantities of infected blood (o'5 and 2 c. cm. respec¬
tively). Trypanosomes were never seen in the Macacus , although it was carefully
eiamined every day. It died 72 days after inoculation, from general tuberculosis.
Parasites were found in the Cercofithecus on the second day after, but were only
present in the peripheral circulation for two days ; they then disappeared and were
never seen up to the time of the monkey’s death, some six weeks after inoculation.
The monkey had a rise in temperature to 1040 F. while trypanosomes were present
in the blood, but afterwards did not have a recurrence of the fever. A rat was
subinoculated from this monkey on the day of its death with 11 c.cm. of almost
pure blood, but has not become infected up to the present, 80 days after inoculation.

Rabbits (3).

The incubation period varies considerably, from four to sixty-nine
flays, the usual time being about three weeks. The disease tends to
he very chronic in these animals, as is strikingly exemplified by the
Mowing experiment : -

A rabbit was inoculated subcutaneously on May 3, 1906, with 4 c.cm. of
™«ted blood and became infected 25 days later. The trypanosomes were present
ln icanty numbers for a week and then disappeared from the peripheral blood.
lnce June 4, 1906, they have never been seen, although the rabbit has been un er
continuous observation. On July 31, two rats were sub-inoculated from the rabbit,
and bo* became infected after an incubation period of between two and three
On November 12, 1906, another rat was sub-inoculated from the rabbit

became infected two weeks later. Finally another rat was sub-inoculated on
'£*** ‘W, and became infected after a prolonged incubation period of 47
J*". 11 wil1 thus be seen that the blood is still infective after the lapse of eleven
D'J?'hs; 11 is said that unfavourable conditions, as lack of food, render amnia s
mW 3rly suscePtible to trypanosomiasis and may cause a declaration of latent
10n In order to test this hypothesis this rabbit was placed on a very restnc : e
. hom December 1, ,9o6, to the middle of February of this year. DunnSjb *

seen IkT* ” Weight from 2’79° 6rm- to 2’°?5 g™- Trypanosomes whe never
Jan0arv hK6 blood> however, and as stated above a rat sub-inoculated at the e
- ,became infected only after a prolonged incubation. The temp

mained normal throughout.

256

Rabbits do not show any symptoms, ’ but remain quite healthy and well-
nourished. The trypanosomes are never very plentiful, and usually show
periodicity. Two rabbits which died had extensive coccidiosis, and their death in
less than four months after inoculation was probably due to this secondary
infection.

Post-mortem. — The spleen is slightly enlarged and rather congested. The
lymphatic glands are usually quite pale, firm, and only very slightly enlarged

Guinea-pigs (3).

The incubation period is prolonged and the disease is also of long duration-
up to 139 days. The animals remain quite healthy and the temperature does not
rise during the disease. The trypanosomes are rather scanty at first, but before
death occurs they increase in number and may finally be fairly plentiful. At the
autopsy of those dead of the disease the spleen is found to be enlarged and rather
soft. The lymphatic glands are increased in sixe to a slight extent and are quite
pale in colour.

Rats (14).

Rats are rather susceptible to the disease. The incubation period is short,
from five to fourteen days. It is interesting to note that in the rat sub-inoculated
directly from the steer ( Yolo No. 9) trypanosomes were not seen until the lapse of
2 49 days. When once established, the disease runs a chronic course as a rule,
lasting for from 40 to 254 days. The usual length is about 80 days. The parasites
are scanty, but usually increase in numbers before the animal's death. Periodicity
is marked ; the parasites may be absent from the blood for two or three weeks
Ihe mode of inoculation has no influence on the incubation period. Large doses
may cause the parasites to appear a little sooner than they otherwise would, but
rats inoculated with such doses do not die any more quickly than those inoculated
with small doses of infected blood. The quick passage of the trypanosome through
a succession of rats (six) did not increase the virulence in any way.

Post-mortem. — The spleen is enlarged, congested and rather firm. The
lymphatic glands are slightly larger than normal, and are quite pale and firm-
The other organs show no changes.

Mice (2).

As in rats, the incubation period is short, about four days, and the disease is
chronic. One mouse is alive 219 days after inoculation and still infected. The
trypanosomes are ahvayrs scanty, and may be absent from the peripheral blood
stream for many days — 78 on one occasion. The post-mortem changes are similar
to those found in rats.

It will thus be evident that most of the common laboratory animals
are susceptible to the infection. Mice and rats are most easily infected,
guinea-pigs least. 1 he disease is chronic and lasts for a comparativel)
long time. It produces few or no symptoms and the trypanosomes
are usually present in but small numbers. The strain is not very
virulent, and resembles in this the strain derived from the horse.

(<£) Horse try pa nos omes

The animal reactions of the strain derived from the second horse,
Toul, are epitomised briefly below.

25 7

Monkeys (2).

Two monkeys were inoculated, a baboon ( Pafio atiubis) and a Macacus rhesus.
The baboon, inoculated intraperitoneally with 5 c.cm. of blood showing one
trypanosome in from one to five fields, never became infected. The Macacus ,
inoculated subcutaneously with 2 c.cm. of citrated infected blood, became infected
after an incubation period of six days. At the same time the temperature rose to
:o5'4°. Unfortunately the further temperature chart is not of much value, as the
monkey developed tuberculosis. The trypanosomes showed periodicity in their
appearance and disappearance in a marked degree. The parasites were scanty at
first, but became more numerous in the course of a couple of days and were
afterwards present in fair numbers. The monkey died 107 days after inoculation,
and at the autopsy the spleen and liver were found to be filled with tubercles. On
the other hand, the lungs appeared quite normal. The lymphatic glands were not
enlarged.

Dogs.

One puppy was inoculated subcutaneously with 1 c.cm. of citrated blood from
a rat, and twelve days later trypanosomes were found in the peripheral circulation.
The parasites have been periodic in their appearance and have never been present
in large numbers. At the time of the first appearance of the parasites there was a
alight rise in the dog’s temperature, but since that there have been no further
increases corresponding to the appearance of trypanosomes in the peripheral
circulation. The dog is quite healthy and presents no symptoms whatever. It is
still living, 76 days after inoculation.

Rabbits (5).

five rabbits have been inoculated with varying doses of infected blood. The
incubation period varies from eleven to twenty-nine days; most usually it is a out
two weeks. The trypanosomes are periodic in their presence in the periphera
blood and are usually rather scanty just before an increase in numbers. One
rabbit only, showed in a marked degree the symptoms usually found in trypano¬
somiasis-emaciation, loss of hair with following ulceration of the skin, oedema of
the external genitals and the base of the ears, increased secretion from the nostrils,
conjunctivitis, haemorrhages into the anterior chambers of the eyes, panophthal¬
mitis and blindness. The animal also became very anaemic and there was a
diminution in both the number of red cells and the percentage of haemoglobin-
AH the others showed no symptoms. The disease is very chronic in rabbits, and
lasted i73 and 178 days in the two which died from the disease. Two other rabbits
led early in the disease from intercurrent affections. The fifth rabbit is s 1
a'lve> 76 days after inoculation. There was no marked rise of temperature when
•be rabbits first became infected.

At the post-mortems of the animals dying of the disease enlargement of the
sPleen and lymphatic glands was the most noticeable feature. The lymphatics
"eie pale in colour and were rather firm in consistence. The spleen was conges e
ind rather soft.

Guinea-pigs (io).

Guinea-pigs appear to be most resistant. The incubation period varied between
'Cu and fifty-two days, six weeks being the average time. The disease, w en o
5t^hed, is 0f iong duration . death ensued in from 46 to 35* days, about 9
J being the average. Two are still alive, 76 days after inoculation. >P
We s,iH present in both. In a few of the guinea-pigs the Para^ . - ,

tinuouidy present from the date of their first appearance in the P P" ^
maiorit 0" Until the animal’s death- They fluctuated in numbers, how ev . I ^
ceil t, ' penodici‘y was noticeable. Occasionally parasites were abs
P eral blood for long periods, e.g. 34 and 67 days.

258

No symptoms of any sort are noticeable in guinea-pigs. The virulence ol the
strain was not increased by successive passages through three guinea-pigs ; the
incubation period and the duration of the disease were not affected. The tempera¬
ture chart does not reveal any rise during the course of the affection. The
trypanosomes are usually rather scanty, but before the death of the animal they
may increase in numbers until they are fairly plentiful.

At the autopsy a more or less marked enlargement of the spleen was seer:
The organ was congested and rather soft. The lymphatic glands, as a rule, were
unchanged, though in a few instances they were slightly enlarged and in one case
haemorrhagic. The other organs appeared normal.

Rats (io).

Rats are rather susceptible to the disease. The incubation period varies
between one and forty-one days ; as a general rule it is about four to six days. The
duration of the disease varied from nine to 253 days, usually about 70. In the
acute cases the parasites are present continuously until death, and are usually
fairly numerous. In the chronic cases periodicity is observed and the trypano¬
somes, when present in the peripheral circulation, are usually in small numbers.
Increase in the virulence of the strain was not obtained by passage through a
succession of four rats. Sub-inoculations made from other experimental animals
became infected in approximately the same time.

Post-mortem. — The spleen is enlarged and is congested and soft in consistence.
Haemorrhagic infarcts were seen in one case. The lymphatic glands may or may
not be enlarged. The enlargement is usually slight when it is present. The
glands are pale and rather firm. The other organs normal. In one case subpleural
petechiae were observed.

Mice (2).

As in rats, the incubation period is short, in our experiments about two days.
I he disease is chronic and lasts for months — as long as five. When first present
the trypanosomes are very scanty. They disappear in a few days and may not be
seen again in the peripheral blood for weeks. In one case they were absent for
fourteen. They reappear before death and increase in number until the animal
succumbs.

Post-mortem . The spleen is very much enlarged, congested, dark purple in
colour, and rather soft in consistence. The lymphatic glands are not enlarged.
The other organs appear normal.

In general, the trypanosome is not very virulent. The incubation
period is prolonged and, while the disease has always terminated
fatally, death only ensues after the lapse of a comparatively long
interval. The low degree of virulence is also manifested by the lack
of symptoms and the absence of heightened temperature in the
experimental animals. Efforts to increase the virulence by successive
passages through any one species of animal (four passages) or through
an alternation of animals, e.g., guinea-pigs, and rats (three of each;
were without result. The parasites are scanty, as a rule, and in the
more chronic cases exhibit periodicity in a very marked degree.

In the following sections the cattle trypanosomes in the Congo
Free State are treated as a whole.

259

VII. DIAGNOSIS

An absolute diagnosis, of course, depends upon the demonstration
of the parasite. The charts of the Kasongo cattle show that every
means may frequently fail to show the parasites in infected animals.
Of all the methods gland puncture was the most successful. In
thirty consecutive examinations, at which parasites were detected
either in the gland juice or in the blood, gland puncture was successful
twenty-nine times, the examination of cover-slip preparations of blood
thirteen times ; on one occasion trypanosomes were present in the
blood and were not seen in the gland juice. T he same precautions
must be taken in puncturing the glands of cattle as in those of men.3
The records of the animal inoculations (particularly at Kasongo,
page 243) show that the animals we employed frequently remained
uninfected after the inoculation of large quantities of blood containing
living parasites. As a diagnostic test the sub-inoculation of laboratory
animals was therefore often less sensitive than the examination of
cover-slip preparations of blood.

If we except the cattle at Romee (seven laboratory animals, all infected), the
bovine trypanosomes (i.e. , first passages) were remarkable for their slight virulence
in direct inoculation. Of twelve rats, four guinea pigs, two rabbits, two monkeys
and four dogs inoculated at various stations, only one rat ever became infected.

It is noteworthy that at present sub-inoculations into ordinary laboratory animals
°f the strain derived from this rat are always successful. The virulence of this
trypanosome in direct inoculations therefore seems to have increased for the
animals employed. The original strains of trypanosomes obtained from the
antelope and from Commandant Sillye’s stallion at Kasongo were much more
•irulent in the first passages.

No idea was formed of the absolute efficiency of gland palpation
as a diagnostic test. In the animals found to be infected the
superficial lymphatic glands were usually considerably enlarged ; some
" ^em, not advanced cases, however, bad no evident glandular
enlargenient. Many of the animals examined once, and not found to be
infected, had glands as large as, or larger than, those seen in the
'nfected animals (this was especially so at Kasongo).

. In the Gambia6 it was noted that a heightened temperature
urnished a valuable indication of possible trypanosome infection in
^■ Charts I, II, HI, IV and V fully confirm this observation;
°Ul a h,gh temperature observed once in an animal just thrown after
a struggle in the sun has but little value. In such circumstances
Stares of from ioic°F. to I03°F„ or more, were noted in
Q

26o

animals carefully examined for parasites without result and believed to
be healthy.

The enormous difficulty of demonstrating the presence of parasites
in animals known to be infected is well illustrated by the observations
at Kasongo. The importance of a well-kept temperature chart is
shown by Liboko (Chart IV), whose temperature was constantly
elevated, although parasites were seen but once.

Animals suspected of trypanosomiasis for any reason should be isolated : their
temperature should be regularly and quietly taken ; their gland juice and blood
should be frequently examined, and laboratory animals should be sub-inoculated
from them with doses of ioc.cm. of blood. An examination of this type extending
over some months is necessary before an animal can even tentatively be said to
be uninfected.

VIII. SYMPTOMATOLOGY

The symptoms are given in the order of their importance, Fever
is practically always present. Listlessness, with progressive weakness,
emaciation and anaemia are usually present. A tendency to diarrhoea,
with small frequent watery stools, and enlargement of superficial
lymphatic glands* are frequently striking features. Oedemas and
running from the eyes and nose were seen in only one or two instances.
Haemorrhages, eye lesions, staring of the coat and loss of hair were not
observed. The appetite remains good to the end. In the autopsies
done the enlargement of the lymphatic glands was by far the most
striking feature ; the glands were either pale and very watery
(particularly superficial glands) or congested and haemorrhagic
(particularly abdominal or mediastinal glands). Oedemas were very
rare, exudates were scanty, the spleen was not enlarged. We saw no
petechial haemorrhages of serous membranes. On the whole, these
findings are identical with those of Broden.3

IX. COURSE OF THE DISEASE

The disease is sometimes exceedingly chronic, and the infected
animal may live for many months. (See Exps. 143, 146, 133. Pa£es
23^. 239, 244, and below).

If marked wasting continues death usually follows in both cattle
and horses within a few weeks or three or four months. Occasional!}
‘"i an“nal at this stage if well cared for will recover flesh, and the

* We are unable to say exactly
apparent. We are inclined to
manifestation.

how long after infection enlarged glands beconif
think that it may be a comparatively *ate

z6i

infection resumes the latent form. As is evident from the observations
in many places, domestic animals may live an apparently normal life
in distncts where trypanosomiasis is rife ; some infected animals
may live for years in apparent health in such districts (see below).

The following observations are interesting in this connection.
The history of Horse VI suggests that animals naturally infected with
Trypanosoma dimorphon may sometimes recover, but are not immune
to re-infection.

A stallion, Horse VI (Kef. 6, page 30), found naturally infected in the Gambia
was brought to England. During two and a half years there were occasional rises
in temperature, trypanosomes were occasionally seen, and its blood was infective to
rats and mice (7). The temperature then became more regular, its blood was no
longer infective, and parasites could not be found. In October, 1905, the super¬
ficial lymphatic glands were just palpable and too small to be punctured. During
the winter of 1905, the animal became thin, &c., but trypanosomes were not seen.
During the whole of 1906 the animal was healthy, its temperature was normal, and
trypanosomes were absent from its blood. The glands remained too small to be
punctured. From time to time inoculations were made into susceptible animals
as follows

July 23, 1906. A white rat, inoculated subcutaneously with 7 c.cm. blood. As
it had not become infected it was re-inoculated on August 14 with 18 c.cm. almost
pure blood. It died on the 5th of September without having become infected,
f ause of death was pneumonia.

October 23, igo6. A guinea-pig received 17 c.cm. of blood subcutaneously and
fiied lebruary 2, 1907. Trypanosomes were never seen, and at the autopsy all the
organs appeared quite normal. On the same day a rat was inoculated with 10 c.cm.
r,f blood. It died on November 5, 1906, from pneumonia ; never infected.

January 15, 1907 . A guinea-pig was inoculated subcutaneously with 20 c.cm.
almost pure blood, and two rats with 20 and 15 c.cm. respectively. They have been
carefully followed to date but have never become infected.

Irom all these points we believe this pony had recovered from a natural
'■'lection by T. dimorfhon.

On the 21st of January, 1907, the pony was inoculated intraperitoneally with
i t em, of infected citrated blood from a mouse infected with T. dimorphon. This
foll°wed between the 6th and 9th of February by a rise in the animal s
emperature, highest 102 5°, and again after a few days intermission to 103-2° on
eibth: With these exceptions the temperature has been since about ioo°.

li *,n lebruary 16, two rats were sub-inoculated from the pony with 1 7 c.cm. of
JJ®each* 0ne °f these became infected on the 6th of March and died six days later
Mar ,nmnerous trypanosomes. The second rat became infected on the 15th o
rr.urti d‘ed 0n the a8th> also heavily infected. In both the spleen was very
en arged. The trypanosomes were of the usual dimorphon type ( s iimpy

“ long ’ forms).

fn addition t0 the rise in temperature the pony lost its appetite and aPPea*®d
careful • °St WClght for several weeks, but is now recovering. The bloo '

*en ■Teixam,ned every day but without result; trypanosomes have never e
enlargemertt ^ands Were PalPated from time to time, but have s ovn

262

The following experiments indicate once more that animals
inoculated in the laboratory with T rypanosoma gambiense may recover
from the consequent infection. The connection between these and the
preceding observations is evident. Their bearing on the question
whether human trypanosomiasis is invariably fatal is most important.

I. A brown stallion was inoculated in the Senegambia (6) from a case of
human trypanosomiasis. For the earlier history consult Memoir X\ I (7) of this
School. In October, 1905, the animal appeared to be quite healthy; no enlarge¬
ment of the glands was present, and no trypanosomes could be found in the blood.
Its temperature was always normal. Sub-inoculations from it were made as follows:

November 1.2, 1906. A guinea-pig was inoculated subcutaneously with ijc.cm.
of almost pure blood. It was accidentally killed on the 31st of December.
Trypanosomes were never seen. On the same date a rat was sub-inoculated with
ioc. cm. Trypanosomes were never seen at any time. The rat died on l ebniary4.
1907, from acute enteritis.

January 16, 1907. A guinea-pig and two rats were inoculated with 20, 17 and
20 c. cm. of pure blood respectively. The guinea-pig and the first rat are still
alive and have never become infected. The second rat died on the nth of
February from an extensive skin disease.

This series of experiments pointed to the fact that the pony had entirely
recovered from the infection by Trypanosoma gambiense. On January 21st it was
re-inoculated subcutaneously with 7 c. cm. of citrated, infected blood, showing one
try'panosome to one to five fields, from a guinea-pig infected with Trypanosoma
gambiense. This, however, had no effect. The temperature has been taken twice
daily for the past year ; there has been no rise, even after the inoculation. The
animal remained in good health ; retained its appetite and did not lose weight. The
pony was most carefully examined day by day, but trypanosomes were never seen.

On March 15 two rats were sub-inoculated, receiving 30 and 250.011.
respectively of almost pure blood. Up to date they have not become infected. On
March 20 the pony was again inoculated intraperitoneally with 25c.cn!. of infected
gambiense blood showing numerous trypanosomes. For a few days afterwards there
was slight auto-agglutination of the cells, but trypanosomes have never been seen.
As in the first instance, no rise of temperature or other morbid symptoms have been
noticed. The experiments are still in progress.

II. A donkey was inoculated in 1904 (7). Since the fall of 1905, the animal
has been in perfect health. No enlargement of the glands, or trvpanosomes, have
e\er been noticed. On January 16, 1907, a guinea-pig (20 c. cm. pure bloodl and a
rat (20c. cm.) were sub-inoculated, but up to date have not become infected. On
January 21 the donkey was inoculated subcutaneously with 7 c.cm. of blood from
a guinea-pig infected with T. gambiense. No rise in temperature or other symptom
followed. For a few days the red cells showed auto-agglutination, but trypano¬
somes were never seen. On March 15 two rats were sub-inoculated from the
donkey with 35 c.cm. of pure blood each. Doth are still alive and have not become
infected. On March 20 the donkey was re-inoculated intraperitoneally with 25 c.cm.
of blood containing T. gambiense (one per field), but without result. No symptom?
of any sort are apparent. The donkey is still under observation.

III. 1 he cow inoculated by Thomas and Dreinl (7) was under observation
until June, 1906, when the animal passed out of our hands. It was in good health
all the time. No trypanosomes were seen and the glands were not enlarged.

263

IV. A monkey ( Cercopiihecus ) was inoculated May 17, 1904, with blood from
i ^eping sickness case. She showed trypanosomes in her blood June 13; they
were seen at intervals up to June 13, 1905, and not again. The animal died
August 5, 1905, from miliary tuberculosis ; no trypanosomes were seen at the
autopsy. (This experiment is reported merely for its interest, as showing a chrome
infection by T. gambiense in a monkey.)

X. CONCLUSIONS DISCUSSED

A. Cattle trypanosomiasis probably exists everywhere in the
Congo Free State. Broden3 is also of this opinion. Since large
game and Glossina are practically omnipresent1 (perhaps with the
exception of high land about Lake Kivu) it seems possible that it
should be so ; at all events, trypanosomes were found in every herd
examined save those at Nyangwe, Cabinda and Lusombo. That the
cursor)' examinations made at these places might easily have failed to
detect infected animals is certain (see charts).

B. The causative trypanosome is probably identical with
Trypanosoma dimorphon. Three ways of identifying trypanosomes
are at present recognised; all of them are liable to error. They are
(1) The morphology of the parasite in infected animals and in
cultures* Cultures have been made of very few of the African
trypanosomes, and were not used in the present instance. (2) Animal
reactions. (3) Cross inoculations.

(1) Morphology. It is well known that the form and dimensions
of all trypanosomes may vary more or less, but when an unknown
trypanosome varies in exactly the same way as does a known
trypanosome (compare the Congo parasites we describe and
trypanosoma dimorphon ) their variations become a strong proof of
identity. And this especially in the present case, since no trypanosome
of domestic animals shows such wide divergencies in form as does
^ dimorphon.

•Ml the forms seen may be approximated to the three types of
irypanosoma dimorphon as described by Dutton and Todd.6 At
^ala’ "’here the cow was suffering from an acute exacerbation of the
d,sease, the - long" type were almost entirely seen, but one “ tadpole
fomnvas observed. At Romee, where the disease was chionic, on y

tary fanM S f^3enat'ons on Trypanosoma brucei and 1 that an

State su§8est a realization of S<*a^* oa Pwould come with

a fuller 1. asSl1ficatlor‘ and differentiation of various haematoz 3 h sts (Koch.
|!SlieknCedg,e °i the,r ,ife cycles in either or ‘1 e^Me der KonigHch

preussivt, lterscheidung der Trvpanosomenarten. Sitzung iqov)

S,Schen Akedemie der Wissenschaften, XLVI Sitzung vom Non. 23, *9 5

264

the “ tadpole ” type was found. In the sub-inoculations, however,
“ stumpy " forms made their appearance. At Lokandu the “tadpole'
and “ stumpy ” forms were seen. At Kasongo “ long " forms were seen
on one cow, and "tadpole " forms in the others. In one of these a
"long’’ form was also observed. The “stumpy and "long "types
was also seen in the antelope at Kasongo. Dutton and Todd
described the “ tadpole ” as being present at the beginning of the
disease, while the other forms of the parasite appeared later.1'1

It seems quite probable that the “ tadpole ” forms are really often
present in chronically infected animals and that they give place to the
“ stumpy ’’ and “ long " forms, as the disease becomes more acute, in
either the original host or in sub-inoculated animals. That this is a
probable explanation is supported by several observations in the
present paper. For example, the history of the parasite as observed
in the case of Yolo (p. 253), where we have the change from the minute
form to the other two types in the sub-inoculations. The similar change
seen by Broden3 in the case of his sheep and its sub- inoculation is of
interest in this connection. Broden states that in the animals sub¬
inoculated from the cattle at Yumbi, the trypanosomes preserved the
characteristics of those seen in the original animals. This observation
seems to conflict with the explanation we have just given. He does
not state definitely that forms with a free flagellum were absent, and
fiom the fact that he compares some of his forms with the hyaline
ones of Bradford and Plimmer which had a free flagellum, it seems
possible that one was also present in some of the trypanosomes he saw.
In this connection it is interesting to note that he observed only the
petite forms in the cattle at Eala, whereas we saw both the "tad¬
pole and the long forms. The occurrence at the same time of
trypanosomes of more than one of the types in several of our animals
certainly also seems to point to the infection being caused by only one
trypanosome rather than by two or more.

(2) Animal Reactions. The virulence of various strains of
trypanosomes (in the laboratory) may vary greatly. The effects of a
parasite on experimental animals can therefore only be taken as a
general indication and cannot be accorded specific value. A parasite
can only be said roughly to be highly, fairly, or slightly virulent.
With Trypanosoma dimorphon the Congo parasites come under the
second class.

265

The variation in virulence observed in the various strains of cattle, horse, and
wild game trypanosomes found in the Congo is certainly not sufficient to suggest that
more than one species of parasite was present.

(3) Cross Inoculation. This method depends upon the belief
that an animal which has acquired immunity* to one trypanosome can
only be re-infected by a trypanosome of a different species. We
believe that this method is open to many fallacies, especially because
of the enormous difficulties of applying it. (a, b, c).

(a) Experiments and observations of many authors show how
extremely difficult it is to be certain that an animal has definitely
recovered from a trypanosome infection. Cases of known infection
often arise in which every means of demonstrating the parasite may

fail.t

(b) For the same reasons an animal may become infected without
the infection being detected.

(0 Our observations on Horse VI show that an animal may be
twice infected by the same parasite.

Laveran (17) infected a rat with T. gambiense which had recovered a year
previously from a previous inoculation. Martin (15) records somew at sum
experiences with a sheep and a goat. lie states that “ une mfime trypanosome peu
revetir des virulences variables et une forme attenuee ne vaccine point con re
forme exaltee,” basing this on the work of Koch and Martini. This complica es
Hie question of the identity of any one trypanosome, and shows that additiona cau ion
must be observed in placing too much reliance on the results of cross inocu a 10ns.
k is altogether probable that the immunity in trypanosome infections is qul e
comparable to that seen in other protozoal diseases such as malaria and spiroc ae e
infections, i.e., only a relatively active and not an absolute immunity is obtained.

Although most authors, in describing a newly-discovered tiypano
some, have found it difficult to say to what species their parasite
belongs, the majority of the trypanosomes, other than 1 rypanosoma
(qi<iperdiim, reported in domestic animals in Alrica, seem to be of the
of either T. brucei or T. evansi. For example, Baleri as described
°y ^azalbou, Jinja cattle disease found by the members of the Sleeping
Sickness Commission in Uganda, Kidei as described by Sanders,
Aino found in Somaliland, El Debab found by Sergent in camels in
Algeria, Mbori affecting animals in French Soudan, Souma desen
’°y Cazalbou, and the trypanosome described by Roger and Greftulhe
'Horses in Algeria. _________

!n our hands the serum reactions dependent upon such immunity

'^unreliable. . u

jJrnk''S P?‘nt cannot be too strongly insisted upon. 11 ; ent^of trypano-

Stred by those engaged in experimental work on the treatment VP

266

The animal trypanosomes we found in the Congo approach
'Trypanosoma dimorphon most nearly in morphology and animal
reactions. On these grounds we identify them with that species.

Broden (3) believes that there are two varieties of trypanosomes, at least, in
the Congo. One of these, corresponding to our “ tadpole ” forms, he calls T. cngh
l ense , while the other he assimilates with T. vivax , described by Ziemann in the
Cameroun. At Leopoldville he found a cow with the two varieties of trypano¬
somes. From the fact that in the sub-inoculations he found only parasites
corresponding to his small forms, he concludes that the animal was suffering from
a double infection. It is possible that the explanation suggested above may apply
in this case. Broden also remarks, in considering the identity of the small form,
that Dutton and Todd were unable to discover their trypanosome in Gambian
cattle. That their examinations were insufficient has been demonstrated by
Captain A. G. Todd, A.V.C. (13), who has since found trypanosomes in these
animals.

In the Soudan, Balfour (14) found a trypanosome in cattle which bears
considerable resemblance to the “tadpole’' type found in the Congo. Five animals—
two monkeys, two rabbits and a dog — were sub-inoculated with small quantities of
blood, but trypanosomes were never seen in any of them. As the author states, the
number of experiments is too small for any dogmatic opinion to be given as to the
non-susceptibility of these animals. It should be again noted here that of many
animals inoculated from cattle by us only one rat became infected. Balfour s
trypanosome was named 7’. nanurn by Laveran. Is it not possible that further
work on this parasite may show that it is merely the tadpole form of T. dimorph^.
In addition to cattle, Balfour found mules infected, but in these the trypanosomes
were frankly of the 7'. dimorphon type (stumpy and long forms). In these the long
slender trypanosomes have a free flagellum.

In French Guinea Martin (15) found that most of the domestic animals suffered
from one of two trypanosomiases. Of these, the most frequent was due to
Trypanosoma dimorphon , and it is interesting to note that in a sheep the trypano¬
somes had a long free flagellum, thus once again bearing out Dutton and Todds
observations. In sub-inoculations, however, the flagellum was lost as in Dutton
and 1 odd’s strain of T. dimorphon. Another interesting point is that in cattle Martin
noted that occasionally' only one type of the parasite was present, most often the
long form.

Quite recently Laveran (16) has described a trypanosome found in sheep
coming from Sdgou on the Upper Niger. This trypanosome presents many of the
characteristics of T. dimorphon , but from the fact that a sheep which had recovered
Irorn the disease, and had afterwards resisted a second inoculation of the same
strain, became infective when inoculated with Trypanosoma dimorphon, Laveran
concludes that it is a separate species, and accordingly names it Trypanosome

7 rypanosoma congolense. Trypanosoma nanum and T rypanosow
vivax, at least, of other African trypanosomes, present marked
affinities with Trypanosoma dimorphon. If they are identical the
distribution of Trypanosoma dimorphon is very wide.

* At this juncture
described by Koch (20,

it is well to consider the variations in form
page 121) to occur in T. brucei.

and virulence

267

c. The transport of cattle from place to place should be
forbidden as far as possible.

Apparently healthy animals may be infected with trypanosomes.
It is easy, therefore, to understand the dangers which lie in moving
cattle. An infected animal might easily introduce a new, and perhaps,
therefore, a more dangerous source of infection into a herd where the
disease does not exist or is present only in a modified form.

Or, again, animals may be introduced into an infected, perhaps
apparently healthy herd, as at Kasongo, only to die of trypanosomiasis.

For example, Kivu cattle brought to the banks of the < ongo ; I.ower ( ongo
cattle taken up the river to Eala. Hroden (3) reports that healthy cattle brought
to Galiema from Dolo, about six miles away, became infected with trypanosomes.
Qlassina falfulis and big game occur about both places ; the possibilities o
infection are equal at both. He suggests that the Dolo cattle, which had no
immunity against the Galiema trypanosome, may have been immune to another or
perhaps merely a less virulent trypanosome occurring at Dolo. \\ e are in entire
agreement with him.

The cattle from Kasongo which we had sent to Lokandu (page ^45) were in
splendid condition. It seemed probable that they were more or less immune to
the Kasongo trypanosome and would live for many months. Out of six amma s
sent, two were dead and two obviously ill within nine months of their arriva a
Lokandu.

If for any reason, e.g., breeding purposes, it is absolutely necessary
to move one or two head of cattle, they should be carefully examined
tor a considerable period (months) before commencing their journey.
While travelling every precaution should be taken to preserve them
from fly-bites. On arrival at their destination they should be isolated
and not drafted into any existing herd until a second examination of

orae months' duration has been completed.

D- Special care must be taken in the choice and main-
eoance of posts for cattle-raising in areas where trypanoso
fliasis is liable to occur.

In the parts of the Congo visited, cattle did best in open plain
-°untry, where tsetse flies (Glossina pal pal is) and big ganu
'.antelopes and buffaloes) were rare and pasturage good.

As Bruce first showed, big game are often infected with try pano
;0mes; ttair wholesale destruction in the neighbourhood of catt
^therefore been frequently advised. To rid a herd of other sources
infection, the infected animals existing in it must also be destroye .

Nothi - - another

’ *,,v '*uvv.i\.u (i 1 1 1 1 1 1 (i 1 n -

hlng is said here concerning treatment. The results presented Qf

- 'Page 275) are, however, quite promising enough to cause
ent ,0 be given a careful trial in trypanosome affections.

268

Since the blood of infected animals remains infective for some time
after their death, and since some flies (S/omoxys and Glossing)
capable of transmitting the disease, will feed on dead animals, great
care should be taken to prevent flies from reaching any part of animals
slaughtered because of trypanosomiasis.8 The G lossinat haunt thick
bush or grass situated near collections of water; cattle should be
kept in clear areas.1 Stonioxys breeds in dung; the kraals should be
kept scrupulously clean.

E. Game, as well as native African cattle and other
domestic animals, may sometimes acquire a tolerance of
certain strains of trypanosomes.

I he African big game is usually said to be immune to trypano¬
somes. I olerant would perhaps be a better word. A buffalo, zebra
and antelopes inoculated with Nagana in Europe have died of the
disease.9 f his " immunity,” therefore, does not extend to every
member of these species, and must be acquired by individuals.
Adult, apparently healthy antelopes (and other wild animals) which
aie infected with trypanosomes are frequently killed. Because of the
percentage* of animals infected and the abundance of biting flies, it is
certain that a young antelope is infected with trypanosomes for the first
time very soon after it is born, and that it will continue to be frequently
re-infected during the whole of its life. The parasites so frequently
found in large adult wild animals, therefore, represent either a new,
recent re-infection or an original infection of long standing. In either
vase the animal has grown up normally, in spite of trypanosome
infections; it is tolerant of the parasites. It seems probable that
the course of trypanosomiasis in African domestic animals may some
times be of a similar type.

artnfiKKarn.g°. trypanosomes were seen in S per cent, of cattle, it
no eC r1S certai,,Iy much larger ; yet this herd is increasing. There «

•splendid condhion01" dUri"g °Ur *** at KaS°ng°’ ^ ^ ^ ^

two JlanOSO,:;iaS" *S c.ornm°n and often fatal among Gambian horses (6). B
s in ie Gambia found to be infected with Trypanosoma dimorfhon

Trale/Z'ihlJVSt 2*'? Cent' infected in South Africa. In the Congo tv

^r*jtre?^teuLch^hren,ige of wi,d aB,nl

269

December, 1902, were still alive, apparently healthy, and were constantly worked
in November, 1906.*

The first part of the history of Horse Yl is another instance of tolerance of
trypanosome infection.

\s will be gathered from the notes on the animals present at various posts
in the Congo, individual cattle, horses, &c., seem to be resistant, since they live
for some years in places where the disease is of a very severe type. C attle are
susceptible to inoculation with Trypanosoma dimorphon, yet at many places in the
Gambia, at Cape St. Mary for example, large herds of cattle in splendid condition
graze over the same ground as the horses. (Captain Todd found trypanosomes
in Gambia cattle, and he states that it is a not infrequent cause of death.)

From all these facts it is not unreasonable to suppose that, like the game,
domesticated animals may thrive in spite of actual trypanosome infection and
exposure to constant re-infection.

It was a “ tolerance ” of this sort which Schilling had in mind when he spoke
of a “ symbiosis ” of trypanosomes with their mammalian hosts (18). Ho describes
cattle which have apparently recovered from infection with 'trypanosoma brucu.
His later publications (19), we believe, simply confirm his former opinion
expressed in (18).

Malaria in African natives forms a striking parallel to this conception.
Children suffer severely from the disease. t Those who survive have parasites
constantly in their blood until they become adult, and even then they are not
infrequently infected, and may have short attacks of fever.

An animal long infected with trypanosomes may have shoit
attacks of fever, with the appearance of numerous parasites in the
Wood, and afterwards completely recover its former tolerant status
(Horse VI). From the fact that adult antelopes are sometimes heavil)
infected with trypanosomes, it seems likely that there may he reciude-
scences of the disease in the same way in them. From the conditions
obtaining in the Gambia, where “ sleeping sickness ’ has been endemic,
for many years, it seems probable that man may acquire a similar
tolerance of Trypanosoma gambiense (b page 23). Almost nothing
!S known of the causes which may make the latent form
ot infection in animal or man virulent, and so produce the
beath of the individual host or an epidemic of trypanosomiasis in
P!ilces where it was formerly sporadic (10, page 319)' ( ’’ Pa£e 2 7)‘
Mc'.i an effect might conceivably be produced by (tf) a diminution m
“Resistance of the host, or by (< b ) an increase in the virulence of the
^ {a) Epidemics of sleeping sickness have seemed to follow

ard Hmes produced by wars and famines,5' 1,1 hi the individual

lhese animals has been treated at intervals with arseJV?Vn the f°rm

5 solution. In March, 1907, he was still in excellent condition.

*e®s Pani^ c.onnecl'on it is noteworthy that the mortality from trypanosom
Particularly great among calves ■ see Eala. Nouvelle Anvers, Tshota,

27 o

animal, hard work, lack of food and concomitant disease sometimes
seem to lead to a great increase in the number of trypanosomes
present and (apparently) in their pathogenic effect.

( 'b ) Laboratory work has also shown that the virulence of trypano¬
somes propagated by direct inoculation from animal to animal may
vary very greatly indeed.

1 homas and Breinl (7, pages 14 and 21) obtained a very virulent strain of
Trypanosoma gambiense. The animal reactions of the common strain of Tryfm »•
soma brucei are no longer the same as those originally described by Bruce (10, page
140). l'he strain of Trypanosoma dimorphon employed by Thomas and Breinl
finally became much more virulent for laboratory animals than it was at first.

1 he strains of trypanosomes obtained from rats in whom Trypanosoma iruai
had recurred after treatment by atoxyl were repeatedly found to be more virulent
than the control strains (page 276).

Whether natural variations in virulence are of the same type as in these
laboratory examples is unknown. An observation bearing on this point is that
there was no difference between the reactions either at the first or succeeding
passages of animals infected with Trypanosoma gambiense in the Gambia, where
sleeping sickness is sporadic, and those infected at Leopoldville or in Uganda,
where that disease is epidemic (n, 12).

XI. CONCLUSIONS

A. C attle trypanosomiasis is very widely distributed in the Congo
Free State.

B. Trypanosoma dimorphon is probably the usual infecting
parasite.

C. All cattle transport should be interrupted or severely controlled.

D. The posts for cattle raising must be carefully chosen and
maintained.

E. Domestic animals probably acquire a relative immunity to some
strains of trypanosomes, and may even recover spontaneously (see
Section IX).

27I

LITERATURE

For a map showing the route followed by the expedition and for observations on
the distribution of biting flies in the Congo, see Newstead, Dutton and Todd,

■ Insects and other Arthropoda collected in the Congo Free State,” Annals
of Tropical Medicine and Parasitology, No. i, Vol. I, 1906.

Rhodes. "Les infections a trypanosomes an Congo.” Bull. soc. beige
detudes coloniales, Feb., 1904, Bruxelles.

Brans. “Second Report of the Laboratory at Leopoldville.” Soc. beige
d'etudes coloniales, 190(1, Bruxelles.

Reference No. 3, page 151.

Doras and Todd. “Gland Palpation in Human Trypanosomiasis,” and
“Distribution and Spread of Sleeping Sickness in the Congo Free State.
Memoir XVIII, Liverpool School of Tropical Medicine, 1906.

Dittos and Todd. “ First Report of the Expedition to Senegambia.”
Memoir XI, Liverpool School of Tropical Medicine, 1903.

Thomas and Bkkinl. “Trypanosomes, Trypanosomiasis and Sleeping
Sickness.” Memoir XVI, Liverpool School of Tropical Medicine, 1905.
Musgrave and Clegg. “Trypanosomes and Trypanosomiasis with special
reference to Surra in the Philippine Islands." Dept, of the Interior,
Biological Laboratory, No. 5, Manila, 1903.

Pummek and Bradford killed a springbok with Trypanosoma brucei.

Kanthack, Di rham and Blandfoku killed hybrids of zebra with horse an
ass by Trypanosoma brucei. A bush buck inoculated died se\en moiit .
later, infected (?)

Martini killed a buffalo and a zebra with the Togoland trypanosome (see 10,
pages 135, 186).

Laveran and Meskil. "Trypanosomes et Trypanosomiases. Masson ei
Cie, Paris, 1904.

Dutton, Todd and Christy. “ Human Trypanosomiasis in the < ongo.
Memoir XIII, Liverpool School of Tropical Medicine, 1904.

Thomas and Linton. “ A Comparison of Animal Reaction of the Trypano¬
somes of Uganda and Congo Free State Sleeping Sickness with those o
Trypanosoma gambiensc." Memoir XIII, Liverpool School o ropi
Medicine, 1904.

Todd, Captain A. G. “ Second Report on the Animal Diseases of the Gambia
to the Colonial Office,” Dec. 18, 1906.

Balfour, A. “Second Report of the Wellcome Research Laboratories at
Khartoum.” 1906.

Mar ns, G. “ Les trypanosomiases de la (iuint-e Iram,aise. A- Mai
Paris, 1906.

L*!**ax, A. “ Nouvelle contribution a lY-tude des trypanosomiases du Haut-
Kiger.” Compt. Rendus de l’Acad. des Sciences, t. CXL1V, p. 243, i9°7-
Laveran. Compt. Rendus de l’Acad. des Sciences, t. CXLI, July 10,
'^s, p. 91.

Sohilun’g. “Ueber die Tsetsekrankheit oder Nagana.’ Arbeiten
aiserlichen Gesundheitsamt, Band XXI, Heft. 3, >9°4- ? 7 t f

hilling. “ Versuche zur Immunisierung gegen Tsetsekrankheit. ei .
Hv° DJ LII, S i49.

Bd- 0.1, o ,49. . ,

^E:in Mensei “ Handbuch dec Tropenkrankheiten,” Barth, L^ip^’ °ces Df
J** 106 and 125. This article gives an excellent review, with refer
important facts in our present knowledge of trypanosomes.

.

CHART III

CHART I

r T ?T

CONCERNING THE TREATMENT OF
EXPERIMENTAL TRYPANOSOMIASIS

CONCERNING THE TREATMENT OF
EXPERIMENTAL TRYPANOSOMIASIS

276

The strains of T. brucei employed killed rats in from five to seven
days after inoculation. In the later experiments a particularly
virulent strain was used which killed rats in from three to four days.
This strain was obtained from a rat in which trypanosomes had
reappeared after treatment by atoxyl. A similar result was obtained
in four out of eight observations ; more work is being done on this
point.

In all cases the rule laid down by Thomas and Breinl1 was followed
that the experimental treatment of animals infected with trypano¬
somiasis must not be undertaken unless the disease is well advanced—
that is, unless the parasites are constantly present and the animal
shows obvious signs of illness (loss of weight, &c.). -The poisonous
dose of each drug, for the animals employed, was always ascertained
as a preliminary step to experiment. The largest possible therapeutic
dose was employed in every instance. The drugs were given
subcutaneously in sterile solutions.

Atoxyl, unless it was otherwise stated, was given in a 5 per cent,
solution in sterile water. It was found necessary to use only fresh
solutions, since atoxyl in solutions deteriorates rapidly.3

W hen experiments were first commenced, the solutions of atoxyl
were kept for some little time.* It was noticed that their toxic effect
on animals infected with T. brucei was evidently great. Out of 121
rats, 40 died within 24 hours ; out of 58 rats at a similar stage of
infection treated with freshly made solutions of atoxyl only 10 died.
This increased toxicity on standing was confirmed by observations on
a solution of 5 per cent, atoxyl in water which had been kept exposed
to light for seven months. Rats inoculated with -5 c.cin. of this solution
died within four hours. It is interesting to note that a similar solution
of atoxyl kept for the same length of time in a dark-coloured bottle
in a cupboard was not nearly so toxic.

T or purposes of comparison it is well to state here that as a general
rule T. brucei reappeared in the circulation of rats, treated by atoxyl
alone, in from 16 to 25 days after the interruption of the treatment.

ut of 1 13 rats treated by atoxyl in the same manner as was employed
in our mercury' experiments (see below) only three are still alive (at
126, 92 and 74 days after the cessation of treatment). In every case

atoxy/wS had8 stSd t0"C eff®Ct of aqUCOUS so,utionS °f

277

in the following experiments, where sub-inoculations were made from
experimental animals to test whether they were infected with trypano¬
somiasis, from i to 2 ccni. of blood were employed.

The routine examination of the blood was made in fresh |-inch
square coverslip preparations. Occasionally the blood of important
animals was centrifugalised.

For the first ten days or a fortnight after the completion of a
treatment the animals were examined daily. As the animals lived
longer the examinations became less and less frequent, until they
were done approximately weekly. The atoxyl-treated controls were
at first examined daily. When it was seen that the trypanosomes
usually reappeared at about 16 to 25 days after the cessation of
treatment examinations were omitted for the first two days after
treatment. The blood of any animal evidently ill was of course
immediately examined and, if necessary, sub-inoculations were made.

II. STRYCHNINE

Van Campenhout advised* that strychnine sulphate should be
used in connection with atoxyl in the treatment of human trypano¬
somiasis. The following experiments were undertaken with the
object of ascertaining whether strychnine sulphate has any action on
trypanosomes when given alone, in conjunction with atoxyl, or when
combined with arsenic or other drugs. Unless it is otherwise stated
T. brucei was used in every case.

(1) Strychnine sulphate alone

(a) Prophylactic. — A rat was given g-ac.cm. of a i per cent, solution of strych¬
nine sulphate in eight doses between October 23rd and November 7t •
inoculated with T. brucci. The infection took the usual course and the animal died

on the seventh day after inoculation. - . . , ,

(*) Curative. — The treatment of four rats commenced on the first day of the
appearance of parasites in the blood. Two rats got altogether 3 c.cm. o a
oent. solution of strychnine sulphate. The remaining two rats receiv e c.c
same solution in two doses. The rat receiving the largest dose showed sig
°f strychnine poisoning but recovered. All the animals died on t e ir a
showing trypanosomes.

in the

'In private communications, reports in the lay pr. ess Utterly^in ^

1 letin de 1 Academie royale de medicine de Belgique ( . strychnine is

in Campenhout makes it clear that he considers that ,th f

obably not 0n the parasite, but that it exercises a beneficial effect on the ner
sue injured by the trypanosomes.

278

Two rats infected with T. gambicnse each received 32c. cm. of a • 1 percent
solution of strychnine sulphate in five doses between the 5th and 16th November
Parasites appeared on the 26th October. Both animals died on the 23rd November.

In none of the above experiments were any signs of degeneration
observed in the trypanosomes during treatment.

(2) Strychnine sulphate followed, by atoxyl

Curative.— Two rats each received -3c.cn!. of a -2 per cent, solution of
strychnine sulphate on the second day of showing trypanosomes in the blood. On
the third day, when the animals were moribund, they each received 7c.cm. of a
•5 per cent, solution of atoxyl. One died. The second recovered and in the
°i "'eek rei e’vecl three -5 c. cm. doses of a -2 per cent, solution of strychnine

su p a e. t s owed trypanosomes on the 17th day after their disappearance as a
result of the atoxyl treatment, and the animal died three days later.

(3) Strychnine sulphate given in conjunction with atoxyl

Curative. 1 he treatment of two rats was commenced on the second day of the
appearance of trypanosomes in the blood. From the Sth to the 17th of October
eac 1 received 3 2 c.cm. of a ‘5 per cent, solution of atoxyl in five doses and 9c.cn!.
o a 2 per cent, solution of strychnine sulphate in nine doses. On the 16th
.November (17 days after stopping the atoxyl treatment) trypanosomes reappeared.

ne annua died four clays later, i'he treatment was repeated in the other (a rat
su inocu ated from this animal at this time died in the usual way from nagana in

a-'s11’ bllt trypanosomes eventually reappeared and the animal died on the
6th December.

, treatnient of two rats was commenced 011 the third day of the appearance

ot l . gambiense. From the 26th of October to the 23rd of November each animal
receive 7 7 c.cm. of atoxyl in eight doses and 18-2 c.cm. of strychnine sulphate in
wen y oses. One of the rats died on the 2nd December, the other on February
26th, 1907; each had many trypanosomes in its blood.

(4) Strychnine sulphate preceded by atoxyl

^uratt The treatment of two rats was commenced on the first day of the
appearance of T. orucei in the blood, 7 c.cm. of a -5 per cent, solution of atoxyl
n^' ln two doses> and then, on the seventh day after the trypanosomes had
animal! ’ reataient b3 strychnine sulphate was commenced. For a fortnight the
on the- J„ecei^e \aiI-v 2c cm- of a -2 per cent, solution. The parasites reappeared

trypanosome’ infection^’ ^ &tOXyI WM St°Pped’ and both animals died °f

auJallnT “ e,aCh received *5 C.cm. of atoxyl on the third and fifth days of
trvnanncmr ° ”’<cei in the blood. Two of them had no further treatment;

each ■ .e> appeared and both died within 22 days. The two remaining rats
davs nf 2 C'UJ}' strFcbn’ne solution in four doses on the sixth and tenth

ec ion. One died of trypanosomiasis in 17 days, the other in 23 days.

(5) Strychnine arseniate

of a -nf, ^ rats inoculated with T. brucei were given from *5 to 1-5c.cn!-

anneannm f ,Ce” S° Utl°n strychnine arseniate on the second day of the
appearance of trypanosomes in the blood.

the third and^f of t,le dlscase was entirely uninfluenced and the animals died on

he ^perimet. 7 ^ of the trypanosomes. In some ot

experiments signs of degeneration of the parasites were noticed.

279

(6) Strychnine arseniate followed, by quinine arseniate

Curative. — Four rats infected with T. brucei each received 2 5 c.cm. of a -06
solution of strychnine arseniate and 2-5c.cn!. of a -i per cent, solution of quinine
arseniate, as in the preceding experiment. The drugs were given separately, at
twelve hour intervals, each in three doses on three successive days. The animals
died on the fourth day, that is two days after the appearance of parasites in the
blood. A few degenerate trypanosomes were noticed on the second day.

From these experiments we conclude that

Strychnine has probably no harmful effect on Trypanosoma
brucei or T. gambiense.

III. ATOXYL, FOLLOWED BY MERCURY

Our work has been directed by the idea5 that perhaps the
recurrences in cases of trypanosome infection, after treatment by
atoxyl, might be due to some resistant stage of the parasite which
survived the first treatment by atoxyl and later gave rise to recur¬
rences of trypanosomes more or less refractory to atoxyl treatment.
It was thought that these hypothetical resistant forms might be
influenced by some other drug. A series of experiments on the
treatment of animals infected by trypanosomes was therefore
commenced in which atoxyl was followed by some other drug. Lp
to the present the best results have been obtained by the use of atoxyl
followed by mercury. We present the results of this treatment on
rats infected with Trypanosoma brucei.

[a) Atoxyl followed by the bichloride of mercury

The same solution of atoxyl was used as in the preceding experi¬
ments. T he mercury was given subcutaneously in a O' 1 per cent,
solution (Liquor Hydrargyri perchloridi, B.P.). 1 he mercury was

never given until parasites could no longer be found by the examina¬
tion of coverslip preparations. That they were probably usually
absent from the blood in an infective form is shown by the fact that
n°ne of four animals inoculated, as controls, with blood taken from
;'uese experimental animals after their treatment by atoxyl have ever
income infected.

Experiment (no).— Four rats received subcutaneously 0-5 c.cm. of a 5 Per
J Solulion of atoxyl (the same strength of solution is used in the same way

«s otherwise stated, in all the following experiments) on the third day of infection,

• e>ghth day after inoculation, with trypanosomes. On the four following days
Cl,Cm of 1116 °'i per cent, solution of perchloride of mercury was given su
"laneousiy (the same solution was employed in the following experiments).

28o

Trypanosomes were never afterwards seen in any of these rats. One rat died 42
days after inoculation with trypanosomes. Another, being moribund, was killed
95 days after inoculation. Trypanosomes were not seen at the autopsy on either
animal, and a rat sub-inoculated from the rat killed has never shown trypanosomes.
1 he remaining two rats are still alive, 181 days after inoculation. Rats and mice
sub-inoculated from them on the 92nd and 156th days have never shown trvpano-
somes and are still alive.

Experiment (134). — Six rats received o-Sc.cm. of atoxyl solution in two doses
on the second and third days after inoculation. The infection was severe ; one rat
died before, two others just after the first dose of atoxyl. Two of the remaining
rats received no further treatment. One died of trypanosomiasis in 27 days, the
other in 22 days, after the cessation of treatment.

In the last rat the course of atoxyl was followed by 1-5c.cn!. of mercury
perchloride given in two doses on the fifth and eleventh days after inoculation,
trypanosomes never reappeared. Thirty-one days after inoculation this rat seemed
ill, it was therefore killed and another rat was sub-inoculated. Trypanosomes
never appeared, and the sub-inoculated rat died of skin disease three months later.

Experiment (149). — Five rats each received 05 c.cm. of atoxyl on the third day
after inoculation. 1 wo rats received no further treatment,- one died (cause of
death ?) in 16 days after inoculation, the other of trypanosomiasis, in 28 days
(parasites reappeared three days before death).

In three rats the atoxyl was followed by 2-7 c.cm. of mercury perchloride given
m four doses on the sixth, seventh, tenth and eleventh days after inoculation.
One died (cause of death ?) in 4o days. The remaining two are still alive, 155
ays after inoculation. Mice sub-inoculated from them 130 days after inoculation
have not become infected.

Experiment (159). — Six rats each received 0*5 c.cm. of atoxyl on the fourth day
after inoculation. Three rats which received no further treatment showed trypano-
somes m from 12 to 24 days and died in from 15 to 27 days after inoculation.

I he other three rats received 2-7 c.cm. of perchloride in four doses on the
se\ent to tenth clays after inoculation. Trypanosomes reappeared in none of
them. One rat died on the eighth day after inoculation (cause ?), another on the
eleventh day (pneumonia). The last rat is alive 133 days after inoculation, and
mice sub-inoculated from it on the 108th day are not infected.

Experiment (161).— Two rats each received 0 5 c.cm. of atoxyl on the fifth day
after moculaUon. One rat had no further treatment and died three days later.

♦v, 6 er rat rece*vec* 1'9c-cm- of perchloride in three doses on the eighth,

en an twelfth days after inoculation. Trypanosomes reappeared on the 15th
ay. Another 0-5 c.cm. of atoxyl was given at once, but the animal died next day.

I reatment was probably commenced too late here ; both animals
were almost moribund when the atoxyl was given. It is possible that
the result might have been better in the mercury-treated rat, had a
larger dose of atoxyl been given. One dose is probably insufficient
for so heavily-infected an animal. It may be questioned whether the
parasites were ever entirely absent from the circulation.

t. f 'XP®®I''EN1 (2^4)- Six rats w-ere inoculated; one died on the third day. v
e tourth day the remaining five rats received 1 c.cm. of atoxyl in two dose;
wo a no urther treatment. Trypanosomes reappeared in them in 20 to 23 day
er the cessation of treatment, and they died a couple of days later, of trypanc
somiasis (30 days after inoculation).

28i

Three rats, after the atoxyl, each received 27 cm of perchloride in four doses
on the fifth to eighth days after inoculation. Trypanosomes were not again seen in
any of them. One rat died (spleen enlarged, cause of death ?) 20 days after
inoculation; the remaining two are alive 123 days after inoculation. Mice
inoculated from them 98 days after inoculation were not infected and are still alive.

Experiment (294). — The treatment of six rats was commenced on the fourth
day after inoculation ; their infection was heavy and two died on the fifth day.
The remaining four rats received 1-40. cm. of atoxyl in three doses on the fourth,
eighth and tenth days after inoculation. Two rats had no further treatment ;
trypanosomes reappeared in them in 34 and 41 days after the cessation of treatment.
They died two to four days later (53 days after inoculation).

In two rats atoxyl was supplemented by 27 c.cm. of perchloride given in four
doses on the fifteenth to nineteenth days after inoculation. Trypanosomes have
reappeared in neither of them. On the thirtieth day after inoculation one of these
rats died (cause of death ?) ; after the second dose of atoxyl this animal had
shown very marked signs of intoxication. The remaining animal is still alive 95
days after inoculation. A rat and mouse sub-inoculated from it 43 and 70 days
after inoculation are alive and uninfected.

Experiment (296). — Four out of 12 rats died during the night of the second
day after inoculation. On the third and fourth days the remaining eight received
ic.cm. of atoxyl in two doses. Four rats received no further treatment ; one of
them died almost immediately. Parasites reappeared in the other three in from
to to 41 days after the cessation of treatment and all died in from 16 to 49 day's
after inoculation.

On the fourth, eighth, tenth and eleventh days after inoculation 27 c.cm. of
perchloride was given in four doses to each of the remaining four rats ; one died
[intoxication ?) after the first dose. All of the three rats were negative until 36 days
after inoculation, when trypanosomes reappeared in one of them. The other two
are still negative at 84 days after inoculation ; mice sub-inoculated from them at
59 days after inoculation have not shown trypanosomes. The rat in which trypano¬
somes reappeared received 1 c.cm. of atoxyl in two doses on the day its relapse
was detected, and the second day after. It then received 17 c.cm. of perchloride
in three doses on the fifth, seventh and ninth days after the relapse. Parasites
again reappeared in 14 days after the relapse. Atoxyl was again immediately given
(ic.cm. in two doses on successive days) ; the trypanosomes disappeared as usual
from the peripheral circulation. The animal died unexpectedly seven days later.
Trypanosomes had not reappeared in its peripheral blood ; spleen and lymphatic
glands were enlarged.

This is the only instance of a recurrence of trypanosomes after a
satisfactory combined treatment by atoxyl and mercury. In this
experiment treatment was certainly commenced very late. There
Was a smaller interval between the administration of the two drugs
than is usual ; but there were probably no infective trypanosomes left
ln Ae peripheral circulation after the atoxyl treatment, since a rat
subinoculated at that time has not shown trypanosomes and still lives.

Experiment (297).— On the third and fourth day after inoculation 10 rats
received ic.cm. of atoxyl in two doses ; two rats died during the night of the third
da-v- Four rats received no further treatment. Trypanosomes reappeared m them

frnra 13 to 24 days after the cessation of treatment and they died two or three
days later.

282

The remaining four rats received 2-7 c.cm. of perchloride in four doses on the
sixth to tenth days after inoculation ; trypanosomes were not again seen in any of
them. One rat died (cause ?) 25 days after inoculation. The remaining three art
still alive, 56 days after inoculation, and mice sub-inoculated from them on the
thirty-first day are alive and have never shown trypanosomes.

When viewed in the severest light the results of these experiments
are distinctly encouraging. The combined treatment by atoxyl and
mercury was given to 25 rats in advanced trypanosomiasis; all would
certainly have died a very few hours later. Of them 13 are still
living (after 181 to 56 days) while all the controls, which received, at
the same time, the same quantity of atoxyl alone, are long since dead
Experiments have been commenced in which the effect of repeated
doses of atoxyl is compared with this combined treatment by atoxyl
and mercury.

The three rats in Experiments 159, 161 (see above) which died
soon after the commencement of treatment, should probably be
disregarded, since in these instances treatment was commenced very
late. The cause of death of many of the treated animals is obscure.
It must be asked whether death was ever wholly due to intoxication
from the drugs administered. There were two recurrences of trypano¬
somes (Experiments 16 1 and 296). Two subinoculations in rats made
from the treated animals dead in Experiments no and 134 are still
alive. It therefore seems doubly doubtful whether death was in these
instances due to trypanosomiasis. If these facts are considered our
results become still more encouraging*

(<£) Atoxyl followed by Donovan's solution f

atnwl^6 m'IE ^3 1 °) ■ Tight rats received 1 c.cm. of a 3 per cent, solution of
rlnriha m W<<! dos^s on ,he fourth and fifth days after inoculation. Two rats died
Donovm’if i>U«rt night- . 0,1 the sixth, eighth and tenth days 1-5 c.cm. of

them are tin° 'i-10” WaS gIven in three doses to the remaining six rats. All of
dav aw „i; 3 ue 44 'fays after inoculation; mice sub-inoculated on the nineteenth

day are alive and have not shown trypanosomes.

This experiment quite confirms those which have preceded it ; it
seems certain that such a combined treatment as we describe should
e given a thorough trial in the treatment of human and animal

trypanosomiasis.

T. 6in S P°ui npa' m ^ 3 f tyPuL at Present under observation, on T. brucei and

complete accord 4,h the SfuiJobtafned^ram!1 d°"leyS h“V<! ” ^ “

t > per cent Iodide of mercury, , per cent. of Iodide of arsenic in water.

2«3

It has been abundantly proved by the work of many previous
observers and by our own preliminary experiments that mercury
perchloride alone will not cure trypanosomiasis. That it has a marked
beneficial effect when preceded by atoxyl is shown by the results
communicated in this paper. How has the mercury acted? A
tempting explanation would be that atoxyl attacks and kills the usual
active well-known phase of the trypanosome found in the peripheral
circulation, but that there exists somewhere in the tissues an inert,-
resting phase, or that such a phase is formed from the active phase
by the action of the atoxyl.12 On this phase the atoxyl has no
poisonous action or an ineffectual one, but on it the mercury salt,
which conversely has no action on the active form, has a powerful
poisonous action. The result is that the inert form exists after atoxyl
treatment in the tissues. On stopping the drug, or on the trypanosome
becoming inured to it,f there is a recurrence of the active fonn and a
fresh outbreak of the disease. The mercury prevents recurrence by
killing the inert form.

This hypothesis suggests the application successively of two or
more remedies, each attacking a different phase of the existence of
protozoan parasites.

Two such remedies attacking two successive phases aie likely to
prove more successful than two remedies attacking the same phase. T
For example, Mesnil and Nicolle and Aubert9 have shown that atoxyl
in alternation with their aniline colours (Ph and Cl) is practically not
more effectual than atoxyl alone.§ Here two agents identical in
action are administered ; the novelty ot a mercury salt aftei atoxy
lies in the fact that the mercury is not active in the same phase as the
atoxyl.

From this point of view it would be of interest to discover whether
the mercury salt would be equally eflectual after the aniline trypano
cides, and this we are proceeding to investigate.

*We owe this suggestion primarily to Dr. lhomas. ,

+ F.hrlich (u) has produced strains of trypanosomes unintluencec y a oxy •

+ Trypanocidal substances known to have a by Ehrlich (8)

[npanosomes are the following: by Ehrlich and -h g ' ’’ - j , lrvDan-blue) and Cl

jCpan-blue; by Mesnil and Nicolle (9), colours V '-\!V: 11 • ; and'llreinl (1), atoxyl.
bWendelstadtand Fulmer (to), malachite green: bv lhomas and Breml < ~

$ Combined treatment along these lines has been attempted by many authors
notably by Laveran (6) and Thomas and Breinl (1).

It is concluded that : —

(1) In the treatment of rats infected by Trypanosoma bructi the
administration of atoxyl followed by mercury perchloride gives better
results than does uncontinucd treatment by atoxyl alone.

(2) The combined treatment should be given a careful trial in
natural trypanosome infections of man and animals.

(3) The treatment must be commenced as early in the infection
as possible ; full therapeutic doses of the drug must be given; fresh
solutions of atoxyl must be employed ; c probably the mercury should

not be given until the parasites have been driven from the peripheral
circulation.

LITERATURE

1.

2.

3-

4-

5-

6.

7-

8.

9-

Thomas and Breinl. Liverpool School of Tropical Medicine. Memoir XVI,

1906.

\ an ( AMi'hNHOUT. Bulletin de l'Academie Koyale de Medicine de Belgique.

1907.

Breinl and Todd. British Medical Journal, January 17th, 1907.

avera.v. Bulletin de l’Academie de Medecine, 26th February, 1907.
Moore, Nierenstein and Todd. A note on the Therapeutics of Trypano-
somiasis, Annals of Tropical Medicine and Parasitology, Vol. 1, No. 1,

C‘ R- de 1Acad- Sciences T. CXL, 17th April, .905, p. 108; also

Ehrlich and Shiga. Berl. Klin. Wochenschrift, 1907, N 13/17.

Meyer”.' Ther&pie der ^egenwart, Vol. 48, p. .29 (Reported by N.

‘?T* anfl ^IC0LLE> anci Albert. Annales de l’Institut Pasteur, 1907,

Deutsche med. Wochenschrift, 17th Nov., 1904'

Wendelstadt and Fulmer.

N- 47. P- 1711.

, Chemotherapeutische Trypanosomenstudien, Berliner Klinische

uresent ’ ,.9°7.’ N°' 9'12’ P- 33- This paper was received after the

present communication was completed and in the printer's hands.

parasite3 of aS^,'VIN'Moo®Ei Preliminary note on the life history of the

parasite of Sleeping-sickness,” Lancet, 3rd April, 1907.

intravenously* (1). "K"’ Wh° " ta*'e the necessary precautions, atoxyl is best givei

CONCERNING CERTAIN PARASITIC
PROTOZOA OBSERVED IN AFRICA

S

28 7

CONCERNING CERTAIN PARASITIC
PROTOZOA OBSERVED IN AFRICA*

Being the Eighth Interim Report of the Expedition of the Liverpool
School of Tropical Medicine to the Congo , 1903-5

BY THE LATE

J. EVERETT DUTTON, M.B. VlCT.

(WALTER MYERS FELLOW, UNIVERSITY OK LIVERPOOL)

JOHN L. TODD, B.A., M.D. McGill

(ASSOCIATE PROFESSOR OK PARASITOLOGY, MCGILL UNIVERSITY, MONTREAL)

AND

E. N. TOBEY, A.B., A.M., M.D. Harvard

(DEMONSTRATOR AND RESEARCH ASSISTANT, LIVERPOOL SCHOOL OF
TROPICAL MEDICINE)

(Received for publication May id, 1907)

Part lit

MAMMALS (continued)

DOGS

leucocytozoon.

In films of fresh blood taken from an apparently healthy native
puppy in the Gambia free gregarine-like forms were seen which
moved slowly in much the same way as do free drepanidia. Intra-
corpuscular forms were not seen ; the stained slides have been

mislaid

From this observation it seems probable that parasites resembling
the haemogregarines described in Indian dogst may also occur in the

Gambia.

'e desire to acknowledge our indebtedness to Dr. ]. W. B- for

ind aid in the preparation of the drawings accompanying Je prewn
unication, and in the examination of some of the slides of Art rop
’art I of this paper appeared in Memoir XXI of the Liverpool School o
Medicine.

ror example, Christophers: Leucocytozoon cants, Scientific Memoirs o e
is of the Medical and Sanitary Department of the Government of Ind .

■ No. 26, 1905.

288

BIRDS

Very many birds of widely different species were examined Only
those in whose blood parasitic protozoa were found are mentioned
here ; several trypanosomes found in Gambian birds are described in
a previous publication, Memoir XI of this School.

Proteosoma was never seen. H alter idium was very common
among the small birds and pigeons examined in the Gambia. In
the Congo Halteridium was seen only once or twice in small birds
and in none of those in which the parasites described below were
found.

ASTURINULA MONOGRAMMICA. (A grey hawk common in the Congo
Free State.)

One of these birds was shot about midday at Lokandu and its
blood was at once examined. In fresh specimens actively moving
trypanosomes were seen ; dried films were therefore made at once.
On studying them parasites of the same type as the LeucocytozoU
of Danielewsky1 (Plates XX-XXIII, figs. 48, 72) were seen to be
present in large numbers.

A second hawk of this species was shot at Lu sambo. Its blood
contained a trypanosome of apparently the same species as that
observed in the first hawk ; the Lcucocytozodn was not seen (one
small slide examined).

Lcucocytozodn zicmanni.

Under the name of Lcucocytozodn zicmanni , Liihe2 gives a good
review of our knowledge concerning this parasite; to his list of
leferences two additional papers3, 8 should be added, in which the
occurrence of similar parasites is reported in America and in Egypt
Although the name Lcucocytozodn has the priority, its use seems
unfortunate, since the parasite to which it here refers is probably
parasitic in other cells besides white cells4,6 and it has no resemblance
to the recently-described, undoubted leucocytozoa of mammals.

Mule (figs. 48-61) and female (figs. 62-67) gametocytes of the
types fully and finally described by Schaudinn4 and previously reported
by Sacharoff, Ziemann,6 Laveran6 and Berestneff7 were frequent. In
addition, many other undescribed forms of the parasite were seen,
rom the nature and scantiness (four slides) of our material, it is

289

impossible to attempt a far-reaching study of the parasite. Never¬
theless, a fairly complete description is given of some forms constantly
observed. This is done with the purpose of insisting on what an
excellent object for study this type of parasite is ; especially since
some of the forms seen seem in some measure to harmonise with
Schaudinn’s description4 of the development of Spirochceta ziemanni
in the blood stream.

Schaudinn gives no particulars but simply states that the development of
Spirochala ziemanni differs in no important point from that of Hamo proteus noctuee.

In our specimens the forms already frequently descnbed as
makrogametocytes (figs. 66, 67, 69, 72, 73). and mikrogametocytes
(figs. 48, 49) are easily recognisable, and in the main their
appearance is in accord with previous descriptions. It is noticeable
that in all forms of the parasite the nucleus of the host-cell is but
little flattened, and practically never takes the dumb-bell-like shape
described by former observers in similar parasites. The position of
the host nucleus in the adult parasite varies according to the position
in which the parasite dries on the slide. Although it is usually-
placed laterally at a point about midway between the two extremities
of the parasite (fig. 49), it may occur in any position. It has frequently
been extruded (fig. 65), and gametocytes without any included host¬
cell nucleus are not uncommon (fig. 66).

The general structure of the parasites of each sex at all stages of
development is the same. Each parasite has periplast, ectoplasm,
endoplasm and nuclear bodies. The “ periplast ” consists of a non-
granular, pink-staining (always ?) sheath which invests the whole of
the parasite. In it run fairly numerous faint, non-staining, longitudina ,
or, if the parasite is twisted, apparently diagonal striations (these can
be seen by careful examination in even the youngest parasites). I he
periplast with its striations is, as a rule, best seen in the more
lightly staining mikrogametocytes (fig. 5°)- tlie aclu^ ^or”ls

the periplast is prolonged at either extremity into long slen
processes. When the matured parasite becomes rounded, preparatory
to conjugation, the periplastic sheath (fig. 7/) 4S thrown off toget er

with the effete host-cell nucleus 1 (fig. 31)-

A similar process probably occurs in the change from the testing

to the motile stages (Schaudinn)4.

Appearances in young parasites (Plate XX), as well as in t e

290

gametocytes, whose effilated extremities have a distinctly granular
structure, indicate that an “ ectoplasm ” exists quite distinct from the
periplast. In the young parasite its tendency is to stain blue; in
the adult it is pink.

The “ endoplasm ” is the most striking part of the whole parasite.
So much so that it, with the nuclear bodies, has been described by
some as the whole of the parasite. In the makrogametocytes the
endoplasm stains a deep blue (fig. 73). It frequently contains large
numbers (20-30) of coarse, or fine, vividly chromatophilic granules
and many small, circular non-staining areas— “ vacuoles.” The

endoplasm of the mikrogametocytes (fig. 49) stains much less deeply,
and rarely contains granules or vacuoles. The endoplasm appears
to consist of a system of wide alveoli filled with a more or less
granular protoplasm. This structure is particularly well seen in the
makrogametocytes. The sexual variation in the endoplasm exists,

though in a minor degree, in some of the smallest parasites (endoplasm
measures 4//, by 3/i).

The form of the nuclear bodies ” varies enormously, according
° the sex and stage of development of the parasite.

I he amount of chromatophilic material-" nuclear material” in

V- C>S11 S<~nse Present 1S much greater in the adult male (fig. 53)
,, n f 6 ema^e parasites (fig. 67). For convenience of description
he various nuclear structures observed at different stages of

“niirl ^m>f ^ , fre, ^ven t^ie following names: “ chlamydoplasm,”*
line ” epharoplast,” with an accompanying “vacuole,” “the

m . fn <;ert'Un “ dots'” whose nature is certainly problematical. It
and th f ear y Understoocl fhat these names are only descriptive
funrtir^ °f r .ma^enal has not enabled us to definitely ascertain the
stain-' "5 Jb°dieS ‘° WWch ** « applied The fixation and

bv nrf mpl°yced are both ^Ity. The parasttes are often d, started

-f f - r-.%

dissected '’ specimens C g fr0m SUch “ fortuitOUS')'

tunatelv a riifF f I!‘ I2, 'S’ 36)- Romanowsky is, unfor-

blepharonh ^ u”' and the details °f dense structures, such as

hopelessly obscured.

' From Gr. Vkauv^ =a cloak 7 -

indicate the diffusely Dink -.r 1 lls name is adopted as a descriptive terra to

preference to other terms indi sur.ro“nt*mg the nucleus. It is employed in
wish to assert nothine rnn ^ *n®- sim*,ar structures since we know nothing, and
fc, concerning its function.

The “ chlamydoplasm ” is a diffuse, light-purple-staining area
whose outlines, though very irregular, are, nevertheless, usually
definite. In the female parasites (figs. 66, 67, 72, 73) and in males
with lines (figs. 33-46) it is comparatively compact. In older
mikrogametocytes its mass becomes greatly increased (figs. 50-5(0)
and its outline is less definite (figs. 58, 61). It may lie compactly
about the nucleus or stretched out (fig. 76) and distributed in any
way through the endoplasm. If it is divided, one part remains with
the nudeus, the other is about, or near, the blepharoplast (Fig. 50).

The “ nucleus ” is employed as an indefinite term to designate
the denser collection of chromatin enclosed in the chlamydoplasm. Its
shape varies considerably ; possibly its nature does so also.

The “ blepharoplast ” is the term applied to a very deeply staining
collection of chromatin usually occurring quite alone in the endoplasm,
and showing a distinctly granular structure (figs. 3» 55’ 4^)-
younger parasites (figs. 3, 5) only two *-° f°ur granules, arranged
in pairs, can be counted. In the older parasites (figs. 68, 71) the
blepharoplast consists of one central granule and from six to eight
other granules joined together by fine lines and situated about the
periphery of the sphere of translucent chromatophilic material in
which they are placed, and outside the chlamydoplasm. As a rule,
no blepharoplast-like body exists separate from the nucleus in
parasites which show the line formation ; rarely one is present
(%• 37). In such parasites the line seems to have split at the end
opposite to that at which the free blepharoplast is placed. This
“blepharoplast” is probably formed by division of the ‘‘nuclear”
mass of chromatin. An oval non-staining or more lightly staining
area, the “ vacuole,” is very constantly associated with the blepharo¬
plast in parasites of all ages. It may vary greatly in size (figs. 53> 73)-
"Dots,” or chromatic granules similar to those illustrated in figs. 15-
!9, 23- 47, 59, were seen in a few instances. They occur in parasites
°f all sexes and sizes in any position. It is impossible to say anythin^
concerning their function from our specimens. In one parasite of
the type illustrated in fig. 71 a chain of five pairs of granules, arranged
'n a column, occurred within the chlamydoplasm and in immediate
connection with the nucleus. In the same way a column of three
Pairs of granules has been seen placed beside the blepharoplast. n
0ne or two cases a pair of granules was joined by a fine thread.

292

Lnfortunately, our material does not permit a full description and
explanation of the developmental phases of these nuclear structures
We can only describe a few of the appearances observed. One
process, however, the formation of a “ line,” seems common to all
sexes and ages of the parasite. 1 he line is widest about the middle,
and tapers to either extremity. It is usually gently curved; rarely
it seems to be slightly wavy. This effect is probably produced by
the over-lying striations of the ectoplasm.

W e have not determined exactly how this line is produced, nor
what is its significance. The morphological changes attending its
appearance in the younger parasites are as follows: — The nuclear
material, in the youngest parasites (fig. 2, 6), consists of a deeply
chromatophilic area, connected with which is a larger pinkish area
often containing a few chromatophilic granules. A line is projected
apparently from the denser nuclear mass (fig. 1). At the distal end
of this line a smaller, densely-staining area appears (fig. 3). The
steps intervening between this stage and that shown in figs. 16, 17,
18, where a commencing line is apparently arising from the lighter,
not the darker, part of the chromatic material, have not been
determined. It is possible that the forms shown in figs. 4, 5, 9. 10
may intervene at this period. In slightly injured parasites, of all
ages (figs. 15, 36), it is seen that the line is composed of several (up
to four counted) filaments. Sometimes (figs. 28, 33, 33a) it may split
longitudinally, apparently normally ; and (?) in such cases the nucleus
sometimes also divides (fig. 39). Multiplication of the parasite has
not been seen. This appearance is the only one observed which in
any w ay suggests division. I he line may also divide transversely in
the following manner. A portion of the line situated in the
chlamydoplasmic area becomes thinned (figs. 26, 32, 40, 41).* An oval
pink area differentiated from the rest of the nucleus surrounds this
constriction. At either side of the constriction a dark granule
ve ops in the line (fig. 32). Connecting these granules is a very fine
me or axial filament.” (The “ nucleus ” in one instance (fig. 26)
was connected to one of these granules by a line.)

n another, slightly squashed specimen a filament connected the
me with a differentiated “nuclear” area situated in the chlamydo-

,hr J hCCa^Tnally the blepharoplast

through a defect in staining, produce

or nucleus may lie over the line and so,
an appearance resembling a constriction.

293

plasm and then with a “ blepharoplast." In some cases the granules
and connecting filament can be discerned in the line even before the
constriction becomes visible ; in two or three specimens a wavy blue
line was associated with the “ line ” (figs. 38, 46, 62). We make no
suggestion concerning its nature. The further development of this
process was seen in only one specimen (fig. 44), here the two halves
of the line are widely separated. The proximal extremity of each
is capped by a dark granule and surrounded by diffuse chromatin
material, while between them runs the faint axial filament, whose
apparent origin has been described.

The process as thus observed seems comparable to the first stages
of the formation of motile apparatus in the trypanosome-like stage of
Haltertdium (Hcenioproteus) noctuce , Schaudinn. The possibility
suggests itself that the formation and transverse division of the
"line” may represent the third division of nuclear material described
bv Schaudinn, and that the axial filament may represent the flagellar
apparatus of a future trypanosome. Unfortunately, our specimens
permit us to go no further than merely to suggest this hypothesis.

Since line formation occurs in parasites of all sizes, it is a process
common to all ages of the parasite. This is also true of the develop¬
ment of the trypanosome-like, motile stages of Hcemoproteus , and is
another point of resemblance between these two processes.

Objections to this interpretation of the line formation are :

(0 None of the smallest trypanosome-like or spirochaete-like stages
representing the motile stage of the youngest intracorpuscular
parasites were seen. This is strange in a bird so heavily infected.
The trypanosome which was present (described below (figs. 29 and
1°))- if it has any connection with the leucocytozoon, probably
represents the motile stage of a mikrogametocyte.

(2) Forms (figs. 45, 46, 63, 64) in which the length of the line has
,ecome much extended do not seem to harmonize with this hypothesis.
Advanced stages of transverse division of the line were seen but
rarely (figs. 26 and 44 are unique).

The line may lie in any position. It may be connected with its
nu(Teus at its centre or by one end (figs. 45> 74)- Although it is
usually almost totally in the endoplasm of the parasite, its extremities
niaV extend into the ectoplasm (figs. 39- 4^> 74)- According to the
ln which the parasite has dried on the slide the line may be at

294

the side of, above, or below the host-cell nucleus. The line is the
most resistent part of the parasite, and is often intact when the rest
of the parasite has been destroyed in the making of the film.

Young forms, smaller than those illustrated (figs. 6, /, 8, 13),
occur in which it is impossible to detect any structure beyond the
presence of a spot of chromatin in a slightly larger area of blue
endoplasm. Every intermediate stage occurs between these tiny
organisms and such larger, line-bearing parasites as figs. 24, 25.
Intermediate stages, without the line, are not seen between parasites
of this size and the adult gametocytes. Intermediate stages with the
line are frequent (fig. 26). The younger parasites are spherical or
have rounded ends (figs. 4, 13, 25); as they become larger their
extremities become pointed and affiliated. Many figures in Plate 1
indicate that the younger parasites are amoeboid. This, and the fact
that one or two very young parasites were seen free in the plasm,
suggests that the parasites may be able to wander from host-cell to
host-cell without the development of a motor apparatus. Such a
process is described in Heemoprotcus noctuee* It is evident from
changes in form of the adult parasites (figs. 59, 63), apparently not
due to artefacts, that they retain up to a late stage something of the
plasticity of the younger parasites. The younger parasites seem to
apply themselves to (figs. 1, 4, 25) or to enter (figs. 13, 16) the host-
cells. The larger parasites engulf them (fig. 26).

It is noteworthy that the parasite’s endoplasm is always in close
connection with the cell nucleus. As far as staining reaction and
appearance goes, it is evident (figs. 6-8, 13, 21, 25) that the host-cells
often are mononuclear white cells ; very rarely granular leucocytes
are attached.

I robably as the infection becomes older, the host -cell, particularly
the nucleus, becomes larger and stains deeply (figs. 24, 26, &c). It
is noteworthy that the nuclei of the host-cells harbouring female
parasites are more often extruded and when present are more
degenerated than is the case in the male parasites (figs. 62, 73)
Since Romanowsky is a stain which does not penetrate, host-cell
nuclei lying beneath the parasite are frequently almost unstained.
In such specimens it is often difficult to make out the relative
arrangement of host-cell and parasite.

295

Mult Female forms.

Adult female parasites, makrogametocytes, are rather less
numerous and larger than adult male parasites. Their general
characteristics, as outlined above, are : deeply-staining endoplasm,
containing more or less numerous chromatophilic granules and small
clear spaces — “ vacuoles ” ; and small amount of nuclear material, as
compared with the male parasites. The average measurements of the
most usual type of makrogametocytes (figs. 65, 67) are: — Total
length, 55'6//; endoplasm, length 20-3^, breadth 9'S^i. Slender forms
igs. 62, 73) in which these measurements were 6yifx, 22 and 8'5^t
respectively, as well as stumpier forms (fig. 69) measuring 47/i, 16^,
and 14 n, were constant types. Much effilated forms (fig. 72) are rare.
They occur most frequently in the thickest parts of our blood smears.

In the most usually seen form (fig. 67) the nuclear material
forms a diffusely-pinkish area (chlamydoplasm ?), usually without
a sharply-defined limit, but possessing a definite oval contour. In it,
or immediately adjacent to it, is the more or less deeply-staining,
often granular, hlepharoplast ( ?). Adjoining the blepharoplast an
oval, more lightly stained area — the vacuole — can usually be
distinguished (fig. 73). Other less conspicuous, dense, chromatic areas
may also occur in the chlamydoplasm (fig. 66). The phenomena
attending line formation in the female (figs. 62, 63, 64) seem to be
analogous to those described in the male and in the young forms ;
because of the darkly staining endoplasm it is, however, very difficult
to follow them.

No changes were observed in the nuclei of the female cells which
seem peculiar to them. For this reason no special description is
§‘ven of them ; the few drawings reproduced indicate the
resemblance between the nuclear changes in the two sexes. I he
Cf|arse alveolar structure of the endoplasm, common to all forms of the
Parasite, is particularly defined in the makrogametocytes. I he curved
outlines of the alveoli, merging into one another, often give the appear-
,lnce °f wavy blue lines running through the parasite. The gianules
vary greatly in number and size. There may be as many as fifty.
W are almost dust-like ; usually they are larger, and may measure
‘Iniost -5 yx in diameter. They frequently occur in pairs and seem to
'K placed superficially in the parasite. They often lie in lines along
''ne ta'nt striations of the ectoplasm. Of the origin and nature of

296

these granules we can say nothing certain. Granules of similar
appearance sometimes occur in much smaller numbers in mikrogame-
tocytes.

A few rounded forms (fig. 31) are present in which the host-cell
nucleus has been extruded and the ectoplasm thrown off. These are
possibly parasites prepared for fertilization.'1 Nothing was ascer¬
tained concerning the nuclear changes at this stage.* Such parasites
measure about 14/z by 9/z.

Adult whet-stone-shaped parasites occur (fig. 70) which possess
general characters intermediate between the male and female adult
types described. They are, therefore, not readily referable to either
sex ; but our material does not permit an assertion as to whether an
indifferent form exists or not.

Adult Male forms.

Besides the forms in which the line occurs or is developing (fig. 3T
there are other parasites where the nuclear material follows a
development of different type (figs. 48, 52) ; it is impossible to say
definitely whether there is any connection between these two
processes.

Apparently an early stage of this second process is illustrated m
figs. 48, 70, 71, 75. There is more or less diffuse and abundant
chlamydoplasm. In the chlamydoplasm occurs a denser, at first circular
or spherical, mass of chromatin — the “ nucleus.” (It may rarely be
placed just outside the chlamydoplasm.) Outside of the chlamydo-
plasm, but occasionally in connection with a detached portion of it,
is a second denser mass of chromatin — the “ blepharoplast." The
blepharoplast is identified by its granular nature and by its darker
staining. 1 he nuclear material becomes arranged in a thick semi¬
circular arc (figs. 49, 76.) At its centre usually occurs a dot, often
connected with the extremities of the arc by fine lines (figs. 53- 55)
The arc of chromatin becomes hemispherical and the dot increases in
size (figs. 50, 52) until the place of the nucleus is taken by two
irregularly oval chromatin masses of approximately equal size (fig- 51)
A stage preliminary to this process is possibly illustrated in fig- 47'
where four brownish granules, of quite a different colour from the
chromatin granules of the blepharoplast, occur in close connection

* l he line in the nucleus of fig. 31 makes it very doubtful to what stage this
parasite really belongs.

297

with the nucleus. This specimen is unique ; somewhat similar
granules once occurred (fig. 59) in connection with the blepharoplast.

It is possible that the division of the nucleus may be by mitosis.
Some half-dozen forms like figs. 54, 56, 59 were seen. Unfortunately,
it was never possible to distinguish the individual spindle fibres or to
count the individual chromosomes. Some spindles, however, had a
distinctly fibrillar appearance, and the chromatin was usually very
distinctly granular at this stage ; in one specimen (fie- 5 6) the granules
could almost be counted — there seemed to be from seven to nine.
The further development of this process was not observed.

The type of parasite in fig. 68 is probably an early stage of this
second process. The nuclear concentration in the chlamydoplasm is
indistinct and circular. The blepharoplast is very well marked ; its
thread-connected granules are distinct, and not infrequently a
chromatophilic granule occurs well outside the blepharoplast, but
still connected to it by a well-defined reddish thread (fig. 68). (In
one such parasite the blepharoplast had divided into two equal parts
connected by a thread.) This apparent extrusion of granules from the
blepharoplast occurs, but less frequently, at other stages (figs. 49. 5°)-

Figs. 51 and 60 are larger parasites apparently undergoing a
similar process. In fig. 57 the blepharoplast seems to be extruding two
granules, while the nucleus is commencing to divide. In fig. 60 the
nucleus, now almost outside the chlamydoplasm, is almost completely
divided, while the granules in the blepharoplast have become much
more distinct. Fig. 58 is probably a stage in this process. It is
suggested that this second process is possibly concerned with the
prostages of mikrogamete formation.

4 trypanosome.

Six trypanosomes which cannot be identified easily with any
Parasite already described were seen in the slides containing the
leucocytozoon. All were of approximately the same type. The usual
Measurements were about I ,* 3*7/4 ; II, 10*5/1 ; III, 1*8/4; IV, 21*4/4;

*The measurements of trypanosomes in this communication were ?nade
Wording to the formula described by us on page 88 of Memoir XXI of this bchool.

I— Posterior extremity of the parasite to centre of the blepharoplast.

II— From the centre of the blepharoplast to the posterior border ot the
nucleus.

HI — From the posterior to the anterior border of the nucleus.

IV— Anterior border of nucleus to posterior extremity of the body 01 me
parasite.

V— Length of free flagellum.

VI— Breadth of body at its widest part.

1‘le total length of the parasite is also given as a measurement.

2g8

V, ii -8/m; VI, 37 fi. Total length, 49 /i. A perfect flagellum was
seen only in one parasite (fig. 29). In fig. 30 the flagellum is much
shorter ; the greater breadth of this parasite is probably due to its
being slightly flattened.

The blepharoplast is a very darkly-staining oblong King longi¬
tudinally in the parasite and placed just posterior to an ill-marked
vacuole. It obviously consists of a collection of four or more granules.
The flagellum, after forming the thickened edge of a wide (1-5/1),
ample, lilac-staining undulating membrane, ends in one instance
(fig- 3°) in a small carmine-coloured, possibly bi-lobed, expansion,
in immediate apposition to the blepharoplast. The granular, palely
staining nucleus measures about 2 by i'3/x, and lies in a sharply-
defined palely-staining area (3 by 2'$n), situated rather posterior to
the middle of the body of the parasite. In one instance a dark
karyosome-like granule, lying in the clear space, is placed do>e
to the nucleus. The nucleus is obscured by the striations of the
body, and it is seen with great difficulty. The finely-aheol r
body cytoplasm is striated through almost its whole length Tin

much-pointed posterior extremity (especially in fig. 29) is very

lightly

the finely-drawn-out, darkly-stained anterior

At the level

stained. Here, as in

extremity of the body, striation can not be detected. - .

of the nucleus about eight, more or less, light striations (my°nenie^
can be distinguished running longitudinally. Ordinarily (hff 'C,! ^

striae are placed at equal distances. In fig. 30 they are so ^

arrange
• disp05

as to make the cytoplasm appear to be arranged in dark stria6 c 1SP ^
in pairs.* By careful examination of fig. 30, it was thought dia
pairs could be distinguished.

The irregular clear areas in the cytoplasm, as illustrated (^f {0

thon£ht

30), occur in the majority of the parasites. They are not --
be artefacts. They seem to be non-staining, refractile granll'eS r
than vacuoles.

The periplast does not stain as pinkly as is usual in tryPan°^j0pe
but it can, nevertheless, be distinguished as a clear refracti'e en
about the body of the parasite.

ox-*C,°mpare With the my°nemes in the motile forms of the mikrog*111^0
■5-pirochcEta ziemanni.

2Q9

“BUSH FOWL”

A few trypanosomes, measuring about 50^ in length, were seen
in a red-legged bush fowl at Tshofa. Stained preparations were not
made. The parasite was remarkable for the extraordinary length
and the flagellum-like fineness of its posterior extremity. About one
quarter of the total body length lay posterior to the blepharoplast.

LITERATURE

1. Danielewsky. Parasitologie comparce du sang, page 23, Kharkoff, 1889.

2. Mense. *• Handbuch der Tropenkrankheiten,” Vol. Ill, page 170.

3. Novy AND MacNeal. On the trypanosomes of birds. Journal of Infectious

Diseases, March 1, 1905, Vol. 2, No. 2, page 256.

4- Schacdinn. Generations-und Wirtswechsel bei Trypanosoma und Spirocliata.

Arb. a. d. Kaiserl. Gesundheitsamt. Bd. XX, Heft 3, page 434.

5- Ziemann. Malaria und andere Blutparasiten, page 128, Jena.

6- Laveran. Contribution a l’etude de Hamamceba Zictnanni. C omptes rendus

de la Societe de Biologie, p. 671, 16th May, 1903.

7. Berestnekf. Ueber das Lcucoc ytozoon Danielewsky Arch, fur Protist, Bd. Ill,
Heft 3, page 376-386.

Neave. Second Report of the Wellcome Research Laboratories, Khartoum,
page 200, 1906. (The budding forms described by this author were not seen
in our specimen.)

BYCANISTES BUCCINATOR (the trumpeter hornbill).

T rypanosomes.*

Two distinct trypanosomes, one small and the other large, were
seen in a bird of this species shot near Coquilhatville in the Congo
Free State. The smaller trypanosomes (Plate XXIV, figs. 1,2,3, 4, 5, 6)
were by far the most numerous. Their appearance in stained
specimens varies very considerably, so that it is possible to descnbe
three distinct types, although forms intermediate between them can
be distinguished. These may be called the “slender" (figs. 1,2),
“ broad ” (figs. 3, 4), and “ stumpy ” (figs. 5, 6) forms, in accordance
with their general appearance. To some extent the structure of these
forms varies with their dimensions. The stouter forms usually stain
more lightly, have the looser cytoplasm, have no vacuole at the
posterior extremity, and their large nuclei do not extend completely
across their bodies.

This association of qualities is not constant, however, and all
gradations were seen between forms possessing them and the, as a
rule, darkly-staining and more compact slender forms. All these
forms are, therefore, considered to be merely variations of one parasite,
which may be described as follows : —

The blepharoplast stains very densely and more darkly than eithe.
nucleus or flagellum. Its position may vary from the extreme posterior
extremity, in particular in the “ broad ” and “ stumpy ” forms, to a spot
i/i or more from the end of the parasite (fig. 1). It is oblong in shape
and is seen to be granular ; in several specimens at least four
granules, sometimes arranged in pairs, can be counted (fig-
Although the arrangement of these granules occasionally suggests
commencing division, none of the ordinary longitudinal division forms
were seen. The blepharoplast may be placed longitudinally,
obliquely, or transversely in the parasite. Just anterior to it there is
often a well-defined vacuole, or, when that is absent, a more lightly
stained area.

The nucleus frequently extends completely across the body of the
parasite and is almost always surrounded by an area stained more
lightly than is the remainder of the body. The relation between
the size of the nucleus and the type of parasite varies considerably,

* The description of this parasite is reproduced by permission from the Journo
of Medical Research. \ ol. XVI, No. 5, March, 1907.

3° i

as a rule, in the “ broad ” and “ stumpy ” forms the nucleus seems
relatively larger, of looser texture, and stains more lightly. Chromatic
granules, number undetermined, occur in the nucleus of each type of
trypanosome (fig. i).

In several preparations (figs. 3, 6) two small darkly-stained and
closely-apposed chromatophilic granules, surrounded by a pinkish
area, occur in close connection with the nucleus, but just outside the
nuclear membrane. In one instance a well-marked chain of oblong,
twin chromatophilic granules runs forward, in an unstained area, from
the anterior edge of the vacuole for about a quarter of the distance
between the blepharoplast and the nucleus. 1 his structure recalls
a similar appearance observed in Trypanosoma karyozcukton .*

Whether staining deeply or lightly, the cytoplasm of the small
trypanosomes is always granular, but the fineness of the granules
varies greatly. Well-marked striations of the protoplasm occur in
each form, although they are best seen in the broader forms (figs. 3>
4, 5). The striations are evidently a superficial structure, and are
usually wound spirally about the parasite. In some specimens a single
striation can be followed for almost two complete turns (figs. 4- 5)-
From their arrangement it is very difficult to estimate the number of
striations, but these seem to be about 7 to 8 (figs. 4, 5)-

The whole parasite is enclosed in a pink -staining periplast, which
Qn be plainly seen at the posterior extremity and in some parasites
all along the edge of the body, where it is unobscured by the
undulating membrane (fig. 3). The ample undulating membrane
seems relatively widest in the “ slender ” forms.

The usual dimensions of each form are given in the following
table : —

Usual Measurements of the Small Trypanosome

Stumpy Broad

Form Form

Measurement

I ...

1*0 ft

i'6/a

II ...

7'° /A

8'o/a

M

Ill ...

2'0 /A

2-4^ •••

n

IV ...

7‘3M

IO'4/A

V ...

7*8 /a

S’O^A

breadth

VI ...

7'o/a

4'8/a

^ °tal length

VII ...

25T/A

... 30-4/*

S RENDER
Form

r6 /a

IO'4/a-
2‘0 /A
7‘2 /A
9-6 /A
2'8 /A
30-8 /A

* See page 2Q7

* Dutton and Todd, 1003. Memoir XI, Liverpool School of Tropical Medicine,
page 53. 9 6

T

302

Only two examples of the large trypanosome (total length about
64 //.), were obtained. Unfortunately, both are so obscured by
surrounding red cells that it is impossible to reproduce them. Their
blepharoplast is placed much nearer to the nucleus than to the
posterior extremity (and in one instance in a vacuole). The nucleus
almost extends across the body. The undulating membrane is ample
and the flagellum seems to be comparatively short. The body of the
parasite is striated longitudinally ; at the level of the nucleus only
seven striations could be counted. In one parasite an appearance
resembling a longitudinal striation was present in the undulating
membrane. The dimensions of this type are : posterior extremity to
centre of blepharoplast. 28 -3/4 (in one parasite only 17/x) ; centre of
blepharoplast to posterior border of nucleus, 3-3/4; anterior border of
nucleus to termination of body, i8'3/4; free flagellum, 8-3/4; width 5%
At Coquilhatville lack of time prevented a careful examination of
the parasite in fresh preparations. Some months later another
hornbill was shot, and its blood was found to contain trypanosomes
resembling the small type described above. Fresh coverslip prepara¬
tions of blood were kept at room temperature (28° C.) and watched
for some hours by Dr. Inge Heiberg. The changes in form indicated
in the following diagram were observed to occur in three hours.

still seen. Occasionally pairs occurred joined by their flagella.
After three hours both normal and very short “stumpy” parasites
were seen, while after four hours, longitudinally dividing, spherical and
irregular, certainly degenerating, parasites were present.

Eighteen hours after the preparation was made dividing form',
similar to those seen at the fourth hour were still present, while a
stouter trypanosome than those seen at the commencement of d-e
observation now appeared for the first time.

303

REPTILES

A considerable number of snakes, chameleons, lizards, monitors
and tortoises of several varieties, as well as three or four crocodiles,
were examined in both the Gambia and in the Congo. Parasites
were found only in the instances described below.

SNAKES

Trypanosomes.

In stained smears of blood taken from a puff-adder caught in the
Gambia, a single trypanosome-like object was seen.

The measurements are as follows :

Measurements:* 1 ... 6'0/a

II ... 6'4/a

III ... 2*4 /A

IV ... 2I'6/a

V ... 8'4/a

VI ... o'6/a

Total length of parasite 45-0 /a

The body is long, narrow and tapers at each end to a pointed
extremity. The body stains a light blue except for two areas between
the blepharoplast and the nucleus, which stain dark blue. The first
dark area, extending from the blepharoplast anteriorly, is 14/^ l°ng >
the second, following a short interval, is 3/a long. I he nucleus is
composed of red chromatin granules and occupies the entire width of
the body. The flagellum is a very slender filament. Only a sug¬
gestion of an undulating membrane can be seen.

Drepanidia. t

Drepanidia of an ordinary type were frequently seen in the same
snake. The nuclei of the cells containing them were displaced, but

t In the present state of o
cation of a Ilaemogregarine
tec ie\v of what is known co
eptiles, see Liihe; in Mense’s “
Leipzig.

304

not injured ; neither was there any destruction of the cell cytoplasm
Two drepanidia were often seen occupying one blood corpuscle,
sometimes on either side of the nucleus, and sometimes at one end of
the corpuscle, which was then double the usual length (Plate XXV
fig- 52)- Two forms of drepanidia were seen, one with a finely-
granular, striated, blue-staining cytoplasm (Plate XXV, figs. 50, 55),
and the other with a lighter-staining, more loosely-woven cytoplasm,
which contained vacuoles, of irregular size and number, at both ends
(Plate XXV, fig. 51). 1 hese latter forms were often much shorter

and wider than the others. The drepanidia measured from 11 to I }n
in length, and from 4*5 to 8/i in width. Free forms of the usual type
were seen ; multiplication forms did not occur. Folded, “two-shanked."
intra-corpuscular parasites were not seen.

A few of the curious bodies illustrated in Plate XXV, figs. 53, 54
and in Plate XXVI II, figs. 56-58, were seen in the blood of this snake
They weie rods from 10 to 19// long and about 1/1 wide. They
stained homogeneously red with Romanowsky; they were, however,
a deeper red at the edges than in the middle. The ends of the rods
were somewhat rounded, but never well-defined. In a free single rod
there was a deep red dot near the centre (Plate XXV, fig. 53). Two
lods were always found in a cell, and they were usually of about the
same length, though occasionally one was longer than the other. The
s weie sometimes placed parallel, sometimes crossed upon one
another, and sometimes came together at one end. They usually
occurred in cells which were lighter-stained and had a rounder, redder
c eus. They' were sometimes, not always, found in cells containing
repanidia. two cases free parasites were seen; in one case two
ro s were found on a degenerated drepanidium ; in the other case, a
single rod was found free.

, • , amk°n> by a study of slides of blood from other snakes in

t lese bodies occur more frequently, has shown that they
P sent t e curled-up remains of the capsules vacated by drepanidia "
An Unidentified Parasite.

fonnrl rOUndec* Parasite of the type seen in frogs (page 335) was

found m the snake (Plate XXV. fig. 55). It Insured W in

w'br* =>1 1 1 conta'nec* short red rods arranged, like the spokes of a

wheel, about a small central red mass.

Private communication.

305

CROCODILES

Trypanosomes.

In stained smears of blood taken from a crocodile (Crocodilus
cat aphr actus ?) shot in the Congo, a trypanosome with a length of
about 35 ft (not including flagellum) was seen. The body was about
2 n wide, the blepharoplast was 3/2 from the posterior extremity and
the nudeus was near the centre of the body and occupied its entire
width. The undulating membrane was well developed.

Drepanidia.

Drepanidia of an ordinary type were seen in the same crocodile
(Plate XXV, fig. 49). They usually measured 12*5/2 in length and
45 p in breadth. The nuclei of the host cells were displaced but not
injured. No destruction of the cytoplasm of the blood corpuscles
was observed, and no division forms were seen. Folded, two-
shanked,” intracorpuscular parasites did not occur.

LITERATURE

Mixchin, Gray and Tulloch. Proc. Roy. Soc., Series 13., Vol. 78, p. 251. Report
the presence of a large trypanosome in a crocodile in l ganda. 1 he\ give
no further particulars.

TORTOISES

Trypanosomes.

In stained smears of blood taken from a tortoise caught in the
Gambia, trypanosomes of one type were seen (Plate XXV, fig. 45)-
The measurements are as follows : —

Measurement I

II

„ HI
IV
V

VI

Total length of parasite
Width of membrane ...

1*2 to 3'5/a
T 5 to i8'5/*
2-2 to 3’5/A
12-5 to 23*0/2
io’o to 15*0/2
2-0 to 3*5/2
35*0 to 58*0/2
2*0/2

The body tapers gradually towards both extremities, which are
sharp-pointed, the anterior extremity being more slender than the

306

posterior. The cytoplasm is finely granular, and in it are several
round unstained spaces I to in diameter. In one specimen there
were seven of these spaces anterior to the nucleus, one at the
posterior edge of the nucleus and four posterior to that. The body
has fine longitudinal striations. The blepharoplast consists of at
least four granules embedded in a matrix, and is about rip long;
a clear area is in connection with it. From it the well-developed
undulating membrane extends in folds to the anterior extremity. The
nucleus is round, sometimes slightly longer than wide, and occupies
two-thirds of the width of the body. No division forms were seen.
Drepanidia*

Drepanidia of an ordinary type were seen in the same tortoise
The nucleus of the host-cell was displaced, but not injured Twc
parasites were sometimes seen in one blood corpuscle. No destruction
of the cytoplasm of host-cells was observed and no multiplication
forms were seen. Some parasites have a cyst wall about them while
t ey are in the coipuscles. This same encysted condition was observed
in extra-coipuscular parasites. Many of the drepanidia have coarse,
colourless, refractile granules (Plate XXV, fig. 4;), either throughout,
r on y in one half of, their bodies. Two forms of parasites were
, in one the nucleus stains a dark purple and has a dense
^ructure, as has its cytoplasm (Plate XXV, fig. 46) ; in the other,
much larger, the nucleus stains a bright carmine, and the

ITT °f thC nudeus and cytoptasm is looser (Plate XXV, fig. 48)
ie drepanidia measure from 10-4 to i7/J. long and from 4 to &

1 e. : lultiphcation forms were not seen. Folded, “ two-shanked ”
mtracorpuscular parasites did not occur.

1 m ri e,r!!^u*)reSental'VeS of c^ass examined were frogs a

•. .lt? . °°d several hundreds of these was searched f
parasites in either the Gamb.a or the Congo.

frogs and toads

I rypanosomes.

r o . 2 \ Congo Tt y patio soma loricatum vel cost a turn2 (Mayer, Jul
^jva^ound m the blood of representatives of the followin
*See footnote to page 303? ~ - ~ - — '

307

species of frogs and toads : — Rana galamensis (D. and B.), Rana
oxyrhynchus (Sund.), Rana mascarensis (B.), Rappia marmorata
Rapp.), and Bnfo regularia (Renn.). A considerable portion of them
was infected. Blood from these infected amphibia was examined
fresh and stained. In the fresh examinations, blood was carefully .
taken aseptically from the heart or, if it was desired to keep the frog
alive, either from a leg vein or a toe. Coverslip preparations were
made and examined periodically. Blood was kept in sealed capillary
pipettes for varying periods and was then used for making films for
staining. Blood was taken from the frogs and, with coverslip
preparations already made, was examined at all hours of the day and
night. Preparations from the organs and bone marrow and of fluid
from the body cavities were examined ; trypanosomes were seen only
in the blood*

Trypanosoma loricatum was present in almost every frog infected
with trypanosomes. It was frequently associated with trypanosomes
of any of the types described below. As a rule, it was noticed that
parasites of the Trypanosoma sanguinis 4 type (Plate XXVI, fig. 27)
were associated with Trypanosoma inopinalum-Wke forms and with
the leaf-like forms3 (Plate XXVI, fig. 28) ; T. mega 4 (Plate XXVII,
figs. 35-39) occurred with forms resembling T. karyozeukton 4
(Plate XXVIII, figs. 40-42) in everything save the absence of the
specific chain of granules. In 99 per cent, of the frogs infected with
trypanosomes Drepanidium was also seen to be present. As a rule,
if there were many trypanosomes there were also many drepanidia.
Striking exceptions to this rule were, however, seen.t

In fresh blood T. loricatum is practically a frilled operculum of
protoplasm, somewhat pointed at the posterior end. It is convex
on one surface and concave on the other. From the median line of

* In a single paper Mayer (2) described two parasites under two specific names
loricatum (or costatum ) and rotatorium. More recent work has s t

probably dealing with two forms of one trypanosome. One of *ese DHed

before disappear. We retain the name loricatum since it was o^ginally appl^
‘0 a parasite resembling that type of trypanosome which seems to be the adult torm
of the haeniatozoon under discussion. A perusal of the pres P P '

^dent that various other forms of T . loricatum have received s^cihc names^

Jhese must also eventually disappear; but until the life history o P

b«en completely worked out it is scarcely worth while discussing this poi .

t For descriptive purposes names already given to various l3Tes tr>
are frequently used in this paper. They are used without question to designate
Parasites resembling those described under these names b} 9

308

the concave surface at a little distance from the postenor extremity
anses the flagellum. The margins of the organism are roughly
serrated. It moves backwards and forwards slowly, now turning over
on itself and now bending antero-posteriorly. As many authors have
observed, its rate of progression is very slow. It is, therefore, easily
watched under the microscope for considerable periods. A side view
is seen in fig. i3, a dorsal view in fig. 2, and a cross section in fig. 3.

1 . loricatum has a peculiar method of division, which we observed
most completely in preparations of fresh blood of Rana galamensis ,
kept aseptically at ;2° F. to 8g° F. for two or three days.

mongst recent authors, Franca and Athias2 and Bouet6 have
parts of the cycle described in this paper. Franca and Athias2
presentatives of the type of T. loricatum , in fresh and stained
P ens, become rounded and, in one case, segment several times.

ainer specimens they saw a trypanosome of their T. rot at or iu m
ype become spherical and divide. They describe most interesting
.1 ‘ binges in which the blepharoplast seems to play

describ^ i° f Centrosome’ and forms suggesting mitotic division are
1. , ° ' n t*lese same specimens small round parasites, possessing

I rop ast and nucleus, which may develop two flagella, occurred.
_ _ tlb panosome of their 7. rotatorium type was also present

F'£s- 20 and 24 are^nla^gea^oocT d iime te^1 iCflly "°UI, actual living sPeciraeD5-

ameius, the remainiier 1000 diameters.

3°9

They describe one or two anomalous forms whose position cannot be
determined at present. Bouet6 made cultures from the blood of frogs
containing trypanosomes of the T. loricatum 2 and the T. rotalorium2
types. The results obtained with either form of parasite were
identical Herpetomonas-like parasites (with an ill-developed undu¬
lating membrane, however) were the most usual forms in cultures.
It is to be noted that every form of parasite between this type and
the trypanosomes originally present in the frog’s blood could be seen
in the cultures.

Some of the parasites, directly after the preparation was made,
were seen to have lost their striations at one or both ends and to have
become granular. This process results in a swelling at one or both
ends (fig. 4). Later the whole parasite becomes granular ; during this

process a well-marked nucleus and blepharoplast are present. The
body now becomes rounded, and the undulating membrane is gradually
peeled off. The flagellum is attached to one end of the organism
5)- The parasite soon becomes completely spherical (fig. 6), and
m this condition its diameter is about 24/i. The flagellum is still

3io

attached, but soon is cast off and is seen lying in the serum, with the
blepharoplast attached (fig. 7). Unfortunately, discarded flagella
were never seen in stained specimens. It is, therefore, impossible to
state whether the whole of the blepharoplast is cast off or not. If
the whole of it is discarded a new body strongly resembling a
blepharoplast is formed very quickly ; because rounded parasites of
all sizes from one equalling the original rounded T. loricatum to those
only 5/z in diameter possess two chromatic areas, one the nucleus
proper, the other resembling a blepharoplast.”

It is suggested by some forms that part of the blepharoplast at
least may remain when the flagellum is thrown off. Chromatophilic
granules not infrequently occur in the cytoplasm of trypanosomes
which probably belong to this period. Concerning their nature we
can say nothing.

The flagellum after being shed loses its lively movements in
three or four seconds, and dies. The parasite is now round and
granular with a highly refractile nucleus (fig. 8).

In one observation the parasite lost its striations, became rounded
and lost its flagellum in eight minutes after the preparation was made
(figs. 9, 10).+

* It would be expected (2) (8) that the blepharoplast would be newly (oW*
from the nucleus.

t 1 he times given for the periods occupied by the various developmental c^3 .■(.
represent actual observations, and may be taken as approximate averages,
same changes may take place more or less rapidly; very frequently the changes ti»*
longer than has been indicated.

It then became amoeboid, and in fifteen minutes had moved
across two fields (Zeiss, 1-30 aperture, achromatic objective; No. 2
eyepiece). The nucleus became distinctly visible (even before the
flagellum was completely cast off), and its structure could easily be
made out (fig. 11).

The parasite now elongates (fig. 1 2), a constriction appears, and
almost before division is complete a similar constriction takes place
in each of the daughter cells to form four cells, all the cells being
almost of the same size as the original mother cell. T he four cells
divide into eight, the eight into sixteen, the sixteen into thirty-two,
and the thirty-two probably into sixty-four*

Preparatory changes were seen in the nucleus before the first
division. These changes were difficult to follow owing to their
rapidity. At one point a rapidly-moving little tongue process appears,
surrounded by a very small differentiated area (fig. 12). In a second

0r two a dull refractile area becomes visible a little distance from it ;
a second later another one appears, but it is somewhat smaller, and
then still another (fig. 12); but almost before this latter has formed,
!‘le four apparently merge into one large, highly-refractile, rounded
area in which the little tongue is seen moving (fig. 1 3). This area is
an obvious feature in many of the cells before division is complete,
t first very bright in appearance, it gradually fades away , thus in
0necase it was formed at 3 21, it was much less visible at 3-24, it
Could not be distinguished at 3-25, and the cells had completely

Danielewsky states that he has seen 120.

divided at 3 30. Another observation showed the area commenting
to be visible at I *5 5» very bright after two minutes with the little
tongue plainly visible ; it was not so bright at 2, and had vanished
at 2’03 ; at this time division of the cytoplasm was well advanced.
Another tongue and area was very bright at 2'oj, and gone at 2*26.
In one case just before its disappearance, the little halo divided,
showing a delicate, thread-like connection between two dots (fig. 14).
These separated completely into two (fig. 15), and then both
disappeared.

A dot was sometimes discernible at one end of the “tongue."
Clear areas, smaller and quite distinct from those described above,
occasionally appeared for a few seconds in the cytoplasm of the
trypanosomes before the first division. One of the products of the
first division not infrequently divides before the other. Often one

or two of the products of the third, fourth or later divisions divides
no further. Indeed, the development of the parasite may be arrested
at any stage. Such individuals become rounded and very granular
and probably usually disintegrate. Some of them may become
encysted resistant forms.6

In dividing, the cells rotate one upon the other in opposite
irections, so that the long axis of one comes to lie at right angles
to that of the other (fig. 16). The nuclei were indistinct. The time
taken to turn around in one case was about thirty minutes.

In one instance the single rounded organism had divided, in fiv'e
our3 a"d forty-two minutes, into sixteen cells, which were all
apparent y inside the outer covering of the original trypanosome
^ S- 17;. In six hours and a half, many of the cells had divided

313

again, and in seven hours there were counted thirty-two cells which
measured 8 /* in diameter. In seven hours and a half, forty-one cells
were counted, though there were probably more.

The next change occurring in these cells is the acquirement of a
flagellum. Each becomes ovoid, then pear-shaped, and from the
more rounded end a flagellum is produced (fig. 18); there are always
one or two parasites in each colony which remain spherical and
develop no further. The colony of cells now takes on a lively
motion. After a while the young trypanosomes become free and

their movements increase in rapidity. 1 hey divide rapidly by
splitting in their longitudinal axis, and thus in the field containing
the original cells, and for many fields around are seen large numbers
(io to 15 to a field) of young actively-moving trypanosomes.

The young trypanosomes when first formed from the cell by the
acquirement of a flagellum, had the shape and size (x 1,000) indicated

3M

in fig. 1 9. A large or small vacuole is generally seen near the pointed
end. One or two highly refractile dots occur in the cytoplasm. The
movements are rapid and take place solely by the lashings of the
flagellum, which acts as a tractellum. The cytoplasm does not take
part m the production of locomotion even when the parasite has
completely separated from its fellows. These small parasites have a
herpetomonas-hke form ; this was especially so after division, when
the body of the young parasite is drawn out and its width is almost
equal throughout the whole length. No evidence of an undulating
membrane was detected in these parasites; nor was the exact

3 2.0.

Fig . xql.

termination of the flagellum in the cytoplasm ascertained in fresh
preparations. In preparations examined thirty hours and forty-five
nutes after they were made, the very active trypanosomes present
of the shape shown in fig. 20. They contained two refractile

n one preparation small free parasites were seen to divide
ecoming rounded. They then divided longitudinally (fig. 21).

u n ftle .same PreParation examined thirty-two and a hi
oms after it was made, agglutination* * of the small trypanoson,

wer^t 1 nCar the edge of the droP of blood.6 The parasit.

h ere attached together by their flagella (fig. 22). Their movemen,

they were absolutely quiescent. Othc

*ThiS Phenomenon was seen uT^Td^T^Ti^'repanition of

3i5

parasites, rounded and attached to one another as in fig. 23, were seen
in this situation. These also became motionless.

Four days later in the same preparation these small parasites
were still very active ; no change having occurred in them, with the
exception of the production of a bulb found at the flagellar end of the
herpetomonas-like forms6 (fig. 24). Five days later many small
parasites were rounded and granular, but were still moving. Six days

Some of the trypanosomes became rounded, but did not develop
further than the first or second division. Their outline became
indistinct, and, instead of fine granulations, clumps of rather coarse
granules appeared, and later showed Brownian movement.

Examples of such parasites watched for five days are as follows:—
Fig. 27. — Shows a coarse granular rounded form.

Fig. 28. — A tri-lobed leaf-like form.

Fig. 29.— A form with fine dancing granules ; on the fifth day the
granules became large and yellow and the organism
disappeared.

F'g’- 3°- A coarse granular form without movement. In twenty-four
hours there were large refractile granules, apparently fat
globules. I he flagellum was not so easily seen.

ao.

In one preparation an amoeboid form of trypanosome was see

1/' t contained a few rather coarse granules. On the secon

chjn were somevvhat coarser, and the parasite had slightl;

ge its shape, while the pseudopodia had retracted. On the fiftf

317

day it was obviously degenerated. Some of the herpetomonas-like
forms were seen to divide longitudinally. The further development
of T. loricatum was not observed in fresh preparations.

Stained specimens

From the above-named species of frogs and toads, 220 dried films
of blood and organ juices stained by our usual Romanowsky method
were examined. T. loricatum vel costatum 2 was found in most of
them (Plate XXV, figs. 1, 2, 3 and 4). The measurements of this form
are as follows : —

I— 12 to 42 fJL.

II.— The nucleus and blepharoplast are apparently connected.

III. — 8*2 tO l6 fl.

IV. — 237 to 26/z.

V.— 8’2 to 16/u .

VI. — 22 to 45/4.

Total length of the parasite, 52 to 101^.

It is a curious coincidence that in the twenty parasites measured,
the length of the nucleus is approximately the same as that of the
flagellum.

In an organism of this size and shape, spreading and drying in the
preparation of the film will inevitably produce a certain
amount of distortion (Plate XXV, fig. 0- However, the
shape of the T. loricatum met with in Plate XXV, fig- 3>
corresponds with our description of the parasite seen in fresh
blood (page 30;). The body is ovoid, the posterior extremity being
narrower than the anterior. The organism may be found lying at full
length, or with one end partly turned over, or completely doubled upon
itself. The structure of the cytoplasm is finely granular. The
periplast is pleated into ridges which run longitudinally from the
posterior to the anterior extremity (Plate XXV, fig. 4). This appear¬
ance can occasionally be seen in the most beautiful manner in slightly
disorganised slowly moving parasites, in fresh preparations.
Occasionally the organism is twisted or folded upon itself, when the
ndges consequently seem to run obliquely. The blepharoplast is
usually very small and consists of four or more granules imbedded in
a matrix. The thickened edge of the undulating membrane takes its
origin from it and the nucleus is connected with it (Plate XX >
u

The nucleus in nearly every case is elongated and crescentic, with its
concave side toward the thickened edge of the undulating membrane*,
and its anterior extremity is pointed. It is i'6/« to 27/t wide. In
a very few cases it is round, when its diameter is from 3-3/1 to 4-4/!
(Plate XXV, fig. 3). The undulating membrane is well developed and
runs in folds from the blepharoplast across the middle of the body,
from the edge of which it extends about 27/x before the free flagellum
begins. The structure of the nucleus is more or less complex. Its
edges are sharply defined, and at its middle are many very small red
granules. The anterior portion of it is dense and finely granular and
often contains larger masses of chromatin. In some instances large
masses cf chromatin are found at both ends (Plate XXV, fig. 2). In
the fresh this was found to precede division (see figs. 10 and 11 in
text).

Numerous forms occur which have a round body, round nucleus,
blepharoplast close to the nucleus and a short white line extending
from the blepharoplast, like a short flagellum ; this line, however,
never extends beyond the edges of the parasite. The line resembles
the tongue seen in fresh specimens ; both occur in rounded parasites
of about the same size, but we are unable to state their identity. It
is, however, quite possible that this type of parasite may be a
de\ elopmental form occurring at about this period and that it may be
a product of the first dichotomous division of T. loricatum. Differen¬
tiated areas of obscure nature are sometimes seen in the nucleus, but
in none of our specimens have we observed the interesting nuclear
changes described by Franca and Athias.2

1 he observations by Moore and Breinl8 seem to be of interest in
connection with the peculiarly elongated " nucleus ” of many forms of
J- . loricatum.

In stained smears of kept blood from the same frog, the same cycle

mi tiplication and development can be followed as mentioned
a ove in freshly made preparations of fluid blood. In Plate XXV.
g- c, is shown a parasite rounded and about to cast off its flagellum.

T. loricatum by” Martin to those described under other names that

in a frog ( Hyla 1 is verv evident A T?n hzard, and by Marchoux and Salimbem 15
peculiar in that it had^ nr. < * If Parasite described by the latter authors is

stained with difficulty or mau^1®6 flagellum of 7\ loricatum is often

and its undulating membrane'' 'G °Ut °f Slght beneath the body of the parasite),

in T. loricatlTnliSTbyTr^l *7 3 Td sPur which was ^

y ranca and Athias (2) nor by ourselves.

319

The body measures iSp by 13/z, and the free flagellum was 45/x long
The blepharoplast is of the usual structure with the usual clear space
about it, and from it extends the nucleus as a band rip wide and gp
long.

In Plate XXVI, fig. 14, is shown a round form with the flagellum
cast of. The cytoplasm is coarsely granular and the blepharoplast
which is of the usual structure is situated near the edge of the body.
The nucleus adjoins the blepharoplast on one side. The diameter
of the organism is 22 p. The nucleus is about 3*3/^ in diameter.

A division, probably the first of a round form, is seen in Plate
XXVI. fig. 15. The dividing parasite measures 19 p by 13/x. A group of
16 parasites is shown in Plate XXVI, fig. 18; each of these cells
measures about y$p in diameter. Other small parasites, probably
representing parasites of a third division (Plate XXVI, figs. 16 and 17),
have a diameter from 4*4/* to 6'6p. Plate XXVI, fig. 15, is a division
of forms with two blepharoplasts and two nuclei. Such parasites were
frequently seen. Occasionally round cells are found with multiple
nuclei and blepharoplasts, but with the cytoplasm undivided
(Plate XXVI, fig. 19). Such cells were from 11 to 22// in diameter.

The small round cell now develops a flagellum from the
blepharoplast, and the body elongates in one axis (Plate XXVI,
fig. 20). This parasite measures 6 6/x by 5'5/U., and has a free flagellum
22'8/i long. The blepharoplast is 0‘8 by 0*3 P, and is Vip from the
edge. The nucleus is rip, by r6 p.

The next stage in the development is the formation of herpeto-
monas-like forms (Plate XXVI. fig. 2 1). These have a body length of
from 9/x to 22p, divided as follows Posterior granule (see below)
to posterior extremity o’^p to 4’4/a, posterior granule to nucleus 4 2
to io/i, length of nucleus 2 to 4’4/x, nucleus to blepharoplast (when
blepharoplast is anterior) V2p, length of blepharoplast o 5 to o S/a,
blepharoplast to the anterior extremity 2*2//. to 5*5/*. The flagellum
is from iop to 25 p. The width of the body is i'2p.

The body is narrow, tapering at both ends to a fine point.

There is no undulating membrane (Bouet6 describes one as being
present). There are from 4 to 16 vacuoles lying between the
posterior granule and the blepharoplast. The blepharoplast of the
usual granular type lies either at one side of the nucleus or slightly
anterior to it. From it the flagellum arises and runs through the

320

middle of the anterior portion of the body, or. turning a little to one
side, is closely applied to the margin of the anterior part of the body.
In one specimen a bluish line connected the blepharoplast with the
nucleus. The nucleus is long and narrow, sometimes extending
completely across the body, sometimes occupying half the width.
Just posterior to the nucleus is a vacuole, and often when the nucleus
does not occupy the whole width of the body there is another beside
it. Near the posterior extremity is a red mass, rather more lightly
stained than the blepharoplast, which consists of from one to four
granules imbedded in a matrix ; this we have called the posterior
granule. The size of this mass varies from a small point i/i in
diameter. In one case a blue line was seen to extend from this
mass anteriorly, but only for a short distance. It did not connect the
posterior granule and the nucleus. One of these herpetomonas-like
forms has a large anterior end (Plate XXVI, fig. 23), similar to the
bulbous form seen in the fresh preparations (fig. 24 in text). Masses of
agglutinated parasites of this type were seen (fig. 22). Two anomalous
forms were seen which probably belong to this state. One has a body
length of 10 fi and a width of 4'4//. Numerous filamentous flagella
seem to arise from the blepharoplast at one end, and the organism
seems to be encircled by an undulating membrane. The other is quite
herpetomonas-like except for the presence of four flagella (Plate XXVI,
fig. 22). It measures as follows Posterior extremity to posterior
granule i'2/x ; posterior granule to nucleus 5 ‘4 4 ; length of nucleus
3’3/U ; nucleus to anterior extremity V2/x. The anterior part of the
body in the region of the nucleus and blepharoplast is wider, being
2 2/1 wide. The flagella are 15/i, 4-4//, 4-9/* and 5*5 ft respectively.

Similar forms were seen in the fresh preparations. They bear
some resemblance to the trichomonas described in the intestine of
frogs. The blood was, however, taken and kept with every care, and
there was certainly no contamination. These forms certainly occur
in the blood ; whether they represent a stage in the development of
T. I orica turn or not it is difficult to say.

The next stage in the development is a form resembling
T. inopinatum 3 (Plate XXVI, figs. 25, 26). This stage is found in
fresh blood, in contradistinction to the forms just described, which
were found in kept blood alone. The parasites of the T. inofinatutti
type may be divided into two groups to facilitate measurements. The

321

first group has the blepharoplast close to the nucleus. Its measure¬
ments are : —

Posterior extremity to nucleus
Measurement III

ff

ff

if

IV

V

VI

Total length of the parasite
Width of nucleus

... 5 5 to IO’OyU

... i*6 to 2'7/x

... iro to 20'0/z

... Il'O/i.

... 3 3 to 4*4 /*

... 28’5 to 43‘o/i

I 6 tO 2‘2 fji

The other form of T. inopinatum has the blepharoplast posterior
to the nucleus, and measures as follows : —

Measurement

I ...

II ...

III ...

IV ...
V ...

VI ...

Total length of the parasite
Width of nucleus ...

•8 to I2’0yx

... 2'0 to 4’4/i

2’0 to 2‘2 yU.
... i3’2 to i6'o/i

I2’0 fX,

... r6 to 3’3/x
... 32*5 to 49*0//.

i’6/i

The body of both forms is pointed at both ends, or is sometimes
somewhat blunt at the posterior end. I he blepharoplast* is of the
usual structure and is situated in a plane common to the nucleus, or
posterior to it. The nucleus is round and sometimes occupies the
whole width of the body, and sometimes not. The posterior part of
the body tapers gradually, and at the level of the blepharoplast the
body is widest. The undulating membrane is never very full in these
forms.

Forms resembling T. sanguinis 4 (Plate XXVI, fig. 27) are present.
I hey seem to be a further development of T. inopinatum , since every
gradation exists between these two forms. The undulating membrane
>s the most striking feature. Its thickened edge arises from the
blepharoplast and extends transversely across and beyond the body
to border the wide membrane. The blepharoplast is in the same plane
as the nucleus. The measurements are as follows :

•That is, it consists of 2 to 4 or more granules embedded in a matrix and
surrounded by a clear area.

322

Posterior extremity to nucleus
Measurement III ...

99

99

99

IV ...
V ...
VI ...

Total length of the parasite
Width of undulating membrane

4'4 /*

... 1*1 tO 2*2/1

... 12*0 tO 19‘0/i

... 11*0 to 12*0/1
... 1*1 to 2*7/1

... 30*2 to 39*0/1
27/1

From T. sanguinis there seems to be every gradation to the
“ leaf-like ” trypanosomes of Plate XXVI, fig. 28. These forms occur
as frequently as does T. sanguinis, and are never seen in slides from
which T. sanguinis is absent. The posterior part of this leaf-like form
tapers toward its round and blunt extremity. This portion stains
more deeply and seems to consist of folds tightly folded. At about
the junction of the posterior and middle thirds of the body is the
round nucleus, and in a common plane is the blepharoplast The
folds of the posterior part of the body gradually unfold until about
the middle of the body, where they are completely unfolded. The
body then tapers to a sharp pointed anterior extremity. The
thickened edge of the undulating membrane arises from the

blepharoplast, crosses the body obliquely and continues along the
edge of the body at a distance of 1 n from it to the pointed extremity
where it becomes the free flagellum. The undulating membrane is
not well developed, but the thin edge of the body and anterior portion
of the body seemed, in the fresh, to act as an undulating membrane.
The measurements are : —

Posterior extremity to nucleus
Measurement III ...

„ IV .

„ v .

•* VI, at a level of nucleus

Total length of the'parasite

Division forms of this stage were seen.

I here was another variety of the leaf-form. The posterior part
is sei rated at the extremity and the serrations seem to be the points
of a petal-like arrangement of the posterior end. The general effect
is that of a bud about to open (Plate XXVII, fig. 29).

5-5 to 10*0/1
i*i to 27 n
13*2 to 26 *0/1
9-0 to 1 i*o n
6*6 to 9*0/1
30*0 to 48*0/4

323

The leaf-like form seems to gradually change to the second variety
of T. loricatum described by Franca and Athias.2 The change is
brought about by a shortening of the anterior part of the body and a
complete unfolding of its posterior portion. Forms representing this
transformation are seen in Plate XXVII, figs. 30 and 31. This
completes the cycle from T. loricatum to T. loricatum .*

The first part of this cycle, from T. loricatum to the herpetomonas-
like forms, was actually observed in a single living parasite which was
kept under observation during three days. The second part of the
cyde, from the herpetomonas-like forms back to T. loricatum is
largely based upon the examination of stained preparations of fresh
and “kept " blood. We are fully aware of the necessity for caution
in determining a developmental process from stained specimens. In
the present instance the deductions we have made from our
examination of stained preparations have been frequently confirmed,
and even supplemented by isolated observations on fresh specimens.
Our statement is supported by the observation of Bouet0 who saw
every intermediate stage between rounded forms and adult
trypanosomes of the T . rotatorium type2 in his cultures.

We describe several types of trypanosomes which are constantly
seen. All are simply developmental variations of T . loricatum.

I. — Forms resembling the trypanosome found in Hyla arbor ea.

(a) A short variety (probably identical with T. rotatorium )

(b) A long variety.

(c) A wide variety.

II. — A lanceolate form resembling somewhat a type described by
Laveran and Mesnil.3

HI. — Forms resembling T. mega *

(а) T. mega.

( б ) A coarsely reticulated form.

(r) A form with large red granules in it.

IV. — Forms resembling 1. karyozcukton.

V.— An unplaced trypanosome.

, H'.aule (2) evidently saw a considerable part of this cycle, since he believed
'hat the trypanosomes were produced from the white blood c°rP^cJ®s; flagellum
'hat he hail seen leucocytes each develop an undulating

He also describes trypa'nosomes which cast off their motile apparatus 1 and sc > ag
became leucocytes. ' The way in which such a mistake could arise is e y P e
A clump of rounded parasites at the fourth or fifth division bears a very close
resemblance to a group of white cells.

I. ( a ) A typical short form resembling that found in ftvi
aborea (Plate XXVI, fig. io) is about 2 5/1 long, 9 n wide and has a
flagellum 2.S// long. The blepharoplast is situated 2 3/z from the
posterior extremity, and V2M posterior to the nucleus. The nucleus
is 9 M long. The chromatin was collected in masses at either end of
the nucleus, or was diffusely distributed in fine granules throughout
its extent. The cytoplasm is a rather loose network, and it seems
to be continued into the undulating membrane. The side of the
body carrying the undulating membrane thus appeared to be folded
into a series of foot-like projections produced by the involutions of
the thick and substantial membrane.

This general description applies to the other tw-o forms.

{b) The long form is characterised by its great length, 40 to 63/1
not including flagellum which is from 19-2 to 26 n long (Plate XXV,
figs. 6 and 7), its comparatively narrow body, between greatest
projections 5 5 to 72//, and the appearance of folds or pleats running
longitudinally ; this was especially marked at the posterior end.
Sometimes the body is particularly thick and dark-stained. The
length of the nucleus is 22 to 24/u, and the width is

(0 The wide form (Plate XXVI, fig. 9) looks like the narrow
varieties, with the folds unfolded however. It measures as follows
Measurement I ... 22'0/z

Hlepharoplast is connected with the nucleus.
22.0 /i
19 0/1
14-8/*

63 -0/*

membrane ^ haVe the thickened edge of the undulating

membrane running about I* from the wavy edge of the body. *

round^nuc eus was never found m any of these forms "

, c ' . 16 la”ceoIate variety (Plate XXV, fig. c) 0f T loricate

gradLTly Tapers6 to^ TTu P°Sterior e*^mity, from which the body
has smooth ed ?he pointed anterior extremity. The body

is weTdeveloned , ^ The Undula^ng membrane,

the end of the \ ^ dlreCtIy backwards and then curves aroU>ld

The e iS°2h i0d^ rUn T f°ldS a,°^ the -clian line of the bod/-
- in "

I

II
Ill

1 1 IV

V

1 otal body length

325

The nucleus, a long narrow tube tapering at both ends, especially
:;teriorly, extends from the blepharoplast toward the anterior end.
The length of the nucleus was almost the same as that of the
flagellum*

The measurements of this form are as follows : —

Length of body, 38 to 50 fi ; width, 10 to 16/1 ; blepharoplast to
posterior extremity 2’ 2 to 4-4/11 ; width of nucleus 12 to 2/1 ; length
of nucleus 19 8 to 29/1 ; flagellum, i/-6 to 27/1.

A change in the form of a “ leaf-like ” trypanosome observed in
afresh preparation is illustrated in figs. 32, 33, 34. 1 he parasite was

watched until it had assumed an appearance, always without well-
marked striations, almost identical with T. mega. A single form of
this type, probably identical with fig. 33. was seen in the stained

preparations.

Its measurements are as follows: —

Measurement I

... 9'0/i

II ...

... 2’2/i

„ HI -

... 4'4/i

IV ...

... 20'0/i

v

... 9‘0/i

Total length of parasite

... 45’°/*

We look upon this form as an intermediate stage
leaf-like" forms (see above) and T. mega .4

between the

*It is a curious fact that in f nucleus" w^°t he same as that
bribed (lb and II, see page 3*3) *e length of the nucleus

the flagellum.

326

III. — ( a ) The “ Mega ” forms may have long anterior and posterior
(Plate XXVII, fig. 39) ends, a contracted posterior extremity
(Plate XXVII, figs. 35 and 36), or a short contracted body (Plate
XXVII, fig. 37). In fresh preparations the striations are well seen in
the long forms;* these may coil up exactly as does T. karyozcuklon
(Plate XXVIII, fig. 43) (see below). T. mega has been seen to
gradually become more rounded until it became spherical. It then
lost its flagellum. The forms with contracted posterior extremity and
contracted body (Plate XXVII, figs. 35-37) probably represent stages
in this process. From them development was not observed.

In the type of T . mega with a short contracted body the length is
40 n, the width at the posterior part 20 n, the nucleus 3'3/t, and the
blepharoplast l’l/i posterior to it. The width of the nucleus varies
greatly ; in most cases it extends completely across the body, while
in others it was at one side and only 2'2/i wide.

T. mega (type) measures as follows —

Measurement I

.. 4'0/z

„ II

... 1-3 to

6‘o/i

„ III ...

... ri to

4‘4/i

„ IV ...

... 15-0 to

53 ‘o^

„ V ...

... 8-2 to

ifrQfl

„ VI ...

... 3 ’3 to

iro/t

Total length of parasite

... 52-0 to

io4-o/i

Width of nucleus

2*2 to

The original description of Trypanosoma mega holds good for
these parasites. In a few parasites there was a slight variation in
the structure of the nucleus. The clear area just anterior to the
nucleus was still present, but the nuclear area of irregular chromatin
was replaced by an arrangement of about eight or nine processes
which radiated from a focus placed at the middle of the anterior wall
of the nucleus. 1 hey seemed to be placed in a deep purple stroma,
and some of them contained a few reddish granules. A somewhat
similar appearance is illustrated by Broden.7

(b) 1 here was another form similar to the “ Mega ” type except for
the very coarse reticular structure of the cytoplasm (Plate XXVII,

* It may be stated here that more or less distinct longitudinal striations were
present in every type of trypanosome mentioned in this paper.

327

fig. 38). Other forms were quite different. Only five of these
well-marked forms were seen. The measurements are as follows

Measurement I

„ II ...

„ III ...

„ IV ...

„ v ...

„ VI ...

Total length of parasite
Width of nucleus

8'o to ii'o/a
5'6 to io'o/a
2'2 to 2 7/X
... i5’4 to 21*0/4
... 132/A
... 30 to 8'0/a

... 44’c to 58-0/4

2 '2/A

(0 Another form, seen in both fresh and stained preparations,
which resembles T. mega is a wide trypanosome with large granules
scattered through its body. Some had many granules (Plate XXVII,
%• 34). others had few (Plate XXVII, fig. 33). Usually the granules
stained a bright red, some of them, however, were refractile and
remained unstained. Although the other granules in the same
parasite were stained, we can say nothing concerning the origin of
these possibly chromidial granules.

These forms measure as follows : —

Measurement

I ...

... 9'o to 15'0/A

ft

II ...

... 4'4/a

ft

Ill ...

... 4'4 to 5'0/A

ft

IV ...

... i8'o to 27'0/a

ft

V • ...

... 77/4

ft

VI ...

... 6'6 to io'o/a

Total length of parasite

••• 43'5 to 59‘°y“

I he nucleus in these forms is always pale, and across it can be
seen the striations of the body. The blepharoplast is i'2 // long,
rather narrow, and almost hidden by the large granules, which are
about i'5/i in diameter.

IV- — A trypanosome resembling T. karyozeaktonx is present, but
the characteristic chain between blepharoplast and nucleus can never
be seen. These trypanosomes may be classified, according to the size,
as large, medium, and small, or, better, narrow.

The large form (Plate XXVIII, fig. 42) is sometimes coiled more or
le^s tightly (Plate XXVIII, figs. 43 and 44). This phenomenon was
abo observed in fresh preparations. The further development of these

328

forms was not seen. The edge carrying the undulating membrane
has a wavy outline. The width of the body at the level of the
nucleus is less than just anterior or just posterior to it.

The narrow form (Plate XXVIII, fig. 40) was seen in one case to
have a blue line running from blepharoplast spirally to the posterior
extremity, and the undulating membrane was continued for a short
distance posteriorly as a ridge beyond the blepharoplast. The
medium form is seen in Plate XXVIII, fig. 41.

All the forms have clear spaces around the nucleus and
blepharoplast.

Measurement I

Large

13 2 to 26'O/a

Medium

IO'O/A

„ II

7-1 to

I3'o/a

6'6/a

„ III

3’3 to

5'5P

3'3/i

„ IV

51'0 to

75’o/i

37' 2/*

„ V

17-6 to

27-0/4

r8'7/A

VI

3‘3 to

77/“

3*8/a

Total length of

parasite 102-6 to 134-0/4

767/4

W ldth of nucleus

3‘2 to

57/“

2*2/4

Small

9'0 to 20'0/X
3‘ 8 to 8'5/a
2-2 to 3'5/A
207 to 38'0/a
18*5 to 2yofi
1 '6 to 27 n

59-5 to 87-0/1
1 '6 to 27 /i

V. The remaining trypanosome to be described is a long narrow
form, with a narrow nucleus, and a large blepharoplast (Plate XXVI.
hg. 24). It measures as follows:—

Measurement I
„ II

„ III

,» IV

» v

„ VI

I otal length of parasite

... 33 to 6-6/a
... 13-2 to 19-2/4

... 2-2 tO 3'3/A

6'6/a

... ii'o to i6-o/a

I 'I/A

... 37-0 to 54'0/a

WhThetCytOPlaSm 13 rather C0arsely retic»lar, the body is pointed »
both extremes the large blepharoplast is situated in the centre of

membrane11”’ r°m Wh'Ch ar‘SeS ‘he rather 5canty undulat“’S

a ™?artTe rf embIes T ■ inopinatum in many ways, and may be
a parasite of that form. ^ y 3

329

fhe question arises whether the cycle of multiplication just
•• '"'il is completed in the frog, or whether it normally occurs only
ut side the frog, probably in a second blood-sucking host, as a leech.
IV smallest rounded parasites with flagella and the herpetomonas-
iike forms never occurred in freshly-drawn blood. Bouet6 agrees
with this observation in stating that none of the young parasites were
'ceii in the blood. Franca and Athias,6 however, record that they
small rounded parasites both with and without flagella (probably
terms of T. loricatum ) in blood fixed immediately after withdrawal.
A single small rounded parasite (6// in diameter) was seen in our
er.es of slides of freshly-drawn blood With the exception noted
»vc, every other type of parasite occurring in the developmental
. le described in this paper was observed in fresh-drawn blood.
From these observations it seems that this cycle of “ swarm ” division
may be completed in the amphibian host, but that the smallest forms
arc rarely present in the peripheral blood. It is noteworthy in this
amotion that there is no certain correspondence between the length
f time blood containing 7 . loricatum has been kept and the type of
trypanosomes present in it.

More than once large specimens of T. loricatum were seen in
i -i' substance red blood cells occurred. Marchoux and Salimbeni
Mieve that the cells and the parasite are merely superimposed.
Without wishing to assert that the cell has been ingested, that is,
that its presence in the parasite is due to more than a mechanical
accident, we are confident that we have seen instances where the cell
o definitely within the cytoplasm of the trypanosome. The
■ rvpunosomes may be attacked at any stage of their development by
ytes Frequently they seem to resist successfully and are not
. ted In <'iie instance a leucocyte was seen to ingest the
haemoglobin cont txoma of a disintegrated red cell; it was

' 'ting to note that its protoplasm contained dark brown granules
most unusual occurrence.

LITERATURE

1 l.ar. e du sang. Darre, Kharkoff.

• HU SOT les trypanosomes des Amphibiens.

Archive, dc 1 i de Bacteriologie Camera Pestana. Tome ,

PH** »7 and 335, U*mn« I.ibanjo da Silva. These authors give an
C the work done on frog trypanosomes.

330

3- Lavkran and Mes.vil. Trypanosomes et Trypanosomiases, p. 374, parj5
Masson & Cie.

4. Dutton and Todd. First report of the Expedition to Senegambia, Memoir

XI, Liverpool School of Tropical Medicine.

5. Marchoux and Salimbeni. Un trypanosome nouveau chez une Hyla voisine

de H. latcristriga. Comptes Rendus de la Societe de Biol. T. LXII, p.591.

6. Bouet. Culture du Trypanosome de la grenouille. Annales de L’inst.

Tome XX, No. 7 p. 564, 25 July, 1906.

7. Broden. Rapport sur les Travaux du I.aboratoire medical de Leopoldville,

Vol. II, p. 186, Hayez, Bruxelles.

8. Moore and Breinl. Note on the life cycle of the parasite of Sleeping Sickness.

Lancet, May 4, 1907, p. 1,219.

9. Martin. Sur un trypanosome de Saurien. Comptes Rendus de la Soc. de

Biol. T. LXII, p. 594. April 13, 1907.

Drepanidia .*

I' r csh Preparations.

Diepanidia were present in every species of frog and toad named
on page 307, and in almost every frog examined. In the same frog
{Ran a galamensis) in which the developmental changes in
1 . loncatum were observed, drepanidia were present in almost every
othei led blood corpuscle, though scarcely any young parasites were
seen. The red cells were swollen to nearly twice their normal size,
the colouring matter had disappeared, and the cells in fresh pre¬
parations presented the appearance of a crumpled colourless bag, to
one side of which the parasite lay. The nucleus was pushed to the
periphery of the cell (Plate XXIX, fig. 86).

A few of these diepanidia were found free in fresh preparations
examined immediately after making them, but the number of free
parasites increased enormously after half an hour. The drepanidia,
er eaving the cells and becoming free, were long and had a round

YYVttt rVf' enc* ,n ad vance during progression (Plate

* /)• A little way from this end the body tapered

^ 771°^ pOSterior Which though narrow, was bluntly

rounded off (Plate XXVIII, figs. 68 and 69).

Progression takes place in three ways : —

f ^ie kotty stretched out by a gliding movement.

when n K \ gI?arine"Ilke contracti°ns of the protoplasm. This occurs
When Obstruct^ ,s encountered (Plate XXVIII, figs. 7o and ;.)•

Di-ovir, / ,en Jerkin§: backward of the posterior end which has

Previously curved round to meet the anterior end.

See footnote to page 303. ~ - - - - - —

33i

After about an hour, agglutination of the free parasites was
observed. They joined themselves together by their posterior,
sharper, somewhat granular ends. Two, four, or six parasites came
together in clumps in this way. Contractions occurred from side to
side, causing the parasites to bend. Two days after the fresh
preparations were made, the agglutinated groups of drepanidia were
observed still, the individuals being rather granular and stumpy and
actively motile, but they did not undergo any further change.

Three forms of drepanidia were observed, a small form (Plate
XXVIII, fig. 74) which will be described in stained preparations, a
medium form (Plates XXVII I and XXIX, figs. 82, 68, 69) (the common
form), and a very large form (Plate XXIX, fig. 86). These large
drepanidia often contained large yellowish, highly refractile granules,
which varied in si/.e and number in different parasites. There were
from one to about one hundred of these globules in each parasite
(Plate XXIX, figs. 84, 85, 89, 90. 91. 92). These highly refractile
granules were situated for the most part around the nucleus. In some
of these parasites small dancing granules were seen at the more
slender end.

Stained Specimens.

The following description is based on the examination of dried
and stained films.

The nucleus of the host-cell was displaced (Plate XXVIII, fig. 59)
but not injured ; the cytoplasm, however, suffers severely since the
parasite seems literally to tunnel about within the limits of the cell.
Many corpuscles were found in which the contents seemed to have
been devoured, leaving them looking like wrinkled empty sacks
(Plate XXVIII, fig. 67). Such sacks could be seen with the drepanidia
present, or leaving, or gone. Two and three parasites were sometimes
seen in one blood corpuscle. The drepanidia were not only present in
erythrocytes but were also occasionally found in leucocytes.

As already noted there were three principal forms of drepanidia ;
a large form, a medium form, and a slender form. The medium form
was the most common (Plate XXIX, figs. 81, 82), and was from 15 to
1 8/1 long and from 5 to 6/1 wide. It had a fine granular striated
protoplasm, which sometimes contained coarse red granules, and
sometimes not. Some specimens had a few (four) granules at the
anterior end of the body, and some had a straight blue line running

332

from the nucleus to the anterior extremity. The nucleus was from
3 '3 to 4’4 n in diameter, and was placed about //i from the anterior
extremity and 4// from the posterior extremity.

One and two constrictions in these forms were observed, showing
the method of progression described in fresh specimens (Plate
XXVIII, figs. 70, 71). In a few specimens the nucleus had apparently
divided into two, and the halves were attached by a red line
(Plate XXVIII, figs. 68, 69).

Some of these forms were encysted, both in erythrocytes and free
in the serum. Such encysted forms often had a small amount of
red-staining excretion at both ends (Plate XXVIII, figs. 59, 60).

The “ slender ” forms (Plate XXVIII, figs. 73, 74) were from 13 to
15// long and I'l/z wide. The cytoplasm was light-staining and of a
loose texture. The shorter or posterior end was very faint-staining
and the extremity could be seen with difficulty. The longer or
anterior end was a faint pink, which deepened towards the extremity.
On both sides of the nucleus were areas staining the same as the
cytoplasm of the erythrocytes. There were two, sometimes three,
of these areas which were from o'6 to 17 n long. Near the posterior
extremity there were sometimes from two to four or more red
chromatin granules, and occasionally there were a few in the anterior
part of the body. 1 he nucleus consisted of eight peripherally arranged
chromatic granules, “ chromosomes,”* and one or two darker, central
chromatic granules, “ karyosomes,” all connected by fine lines. In
one instance (Plate XXVIII, fig. 73) a blue spiral line was found to run
from one of the chromatic granules to the anterior extremity of the
body. The nucleus was 2 '2/t long and was placed 9/1 from the
anterior extremity, and 3*3/1 from the posterior extremity. This
slender form may be present in the blood with the larger form,
or it may be present in blood which has none of the larger forms.

These slender ’ drepanidia penetrate the erythrocytes, then lose
both ends, so that only the nucleus and a small amount of cytoplasm
around it is left. The chromatic granules go to the side, and later
they are gathered in a mass at one end of the body, with a few
chromatic granules remaining outside the mass (Plate XXIX, fig. 76).

1 he body of the parasite enlarges and at the extremity opposite the
chromatin mass are several round clear spaces, which may or may

in a nnrl'lT,! 1)6 lln^erstooci that the terms chromosomes and karyosomes are
in a purely descriptive sense. :

333

not contain a granule of chromatin. The mass of chromatin divides
into two, and the cytoplasm may or may not divide synchronously.
In cases where the cytoplasm does not divide synchronously, the
chromatin goes on dividing until there are from io to 16 masses of
chromatin (Plate XXIX, figs. 78-80) arranged almost around the edge
of the cell. These rosettes later show divisions into young parasites
(Plate XXIX. fig. 79). Rosettes were seen leaving the host-cell, and
also found free in the serum (Plate XXVIII, fig. G6). In one slide a
group of eleven young drepanidia that had just escaped from the
rosette condition was seen (Plate XXVIII, fig. 72). These forms were
77 long and ri/x wide. Their cytoplasm was of a faint-staining,
coarse, granular structure. The nucleus was composed of a larger,
central mass of granules and, usually, a chromatin granule on either
side; occasionally both were on one side. In one case the chromatin
seemed to be dividing (Plate XXVIII, fig. 72). These sporulating*
forms always arose from the “slender” drepanidia. In one series of
eight slides from one frog many sporulating and “slender” forms
were seen, but none of the larger drepanidia described in the next
paragraph.

In a few frogs, liana mascariensis , a very large drepanidium was
seen, which was like the common form, but larger. It was lighter
stained and often took a pink colour instead of the normal blue. The
cytoplasm was coarsely granular, and many of the parasites had
non-staining, highly-refractile globules in them. These globules were
from one to one hundred in number, and from cr6/i to 77/1
diameter. They were always found near the nucleus and gave the
appearance of a fatty degeneration of the nucleus. They sometimes
distended the drepanidium until it was nearly round (Plate XXIX,
%• 89), but they usually did not affect the size of the parasite. The
large type of drepanidia presented three forms, a long form, a short or
young fonn, and a folded or “ two-shanked ” form. I he long form
'Plate XXIX, fig. 86) was usually 22-6// long and 7/t wide. The
nucleus was 3-4/* long and 5-6/1 from the posterior extremity and
T^/i from the anterior extremity.

The short, or young form (Plate XXIX, fig. 83) was about 8'8/i long
and 6'6/t wide. The nucleus was 4‘4// 'n diameter and placed in the
centre of the organism.

The folded form was seen in the various stages of unfolding

(pl«e XXIX, figs. 87, 88). _

* *he term is not used in its specific sense.

V

334

Cytamoeba.

Cytamoebae were present in a few of the frogs in considerable
numbers. Amongst the forms seen was the round form (Plate XXIX,
figs. 104, 106, 108, 1 10) previously described in the snake (page 304)
(2'2/z wide). In it were short rods, or sometimes round, red dots
(Plate XX TX, fig. 104). Other forms were oval or finely granular,
sometimes with no apparent internal structure (Plate XXIX, fig. 103).
The round forms were often found near the nucleus, and sometimes
apparently emerging from it, giving the appearance of fragmentation
of the nucleus (Plate XXIX, fig. 1 10). Some of the parasites were
filled with slender rods, which at times projected beyond their edge
for a distance of 4^ (Plate XXIX, figs. 1 12, 1 13). Other forms were
present which looked like masses of rods bound together (Plate XXIX,
figs. 107, 108, 109, 1 1 5); sometimes these bundles of rods were free
in the serum (Plate XXIX, fig. roo). Occasionally short rods were
found free in the erythrocytes, sometimes end to end, sometimes
crossing one another at right angles. The rods found in all the above
forms were of two kinds, a short rod with rounded ends, and a long
filamentous rod. Some of the short rods were seen free in the1
blood plasma in such forms and arrangements that there seems to be
no doubt that some of them at least are bacteria. It is equally
certain that some of the longer slender rods are merely filamentous
cytoplasmic processes of the rounded parasite. In some of the
specimens the body of the cell was stained a light blue, while the
rods within were stained a bright red.

LITERATURE

*• Kruse- Uebef Rlutparasiten. Virchow’s Archives. Jan. 2, 1S90, Yol. uo.
P- 54».

2. Gabritchewsky. Contribution a l’etude de la parasitologie du sang. Annales
Inst. Pasteur, 1890, T. IV, p. 480.

3- I.abbk. Recherches zoologiques et biologiques sur les parasites endoglobulaires

du sang des vertebres. Archives de Zoologie experimentale et generate.
3 Series, T. 2, 1894, p. 5S.

4- Wasielewski. Sporozoenkunde. Jena, 1896, p. 88.

5- Ziemann. Uber Malaria und andere Blutparasiten. Jena, 1798, p. i44-

6- Laveran. Sur le bacille parasite des heinaties de Rana esculenta. Memoir de la

Societe de Biologie, 1899, p. 355. This has a goocl review of the literature.

335

An unidentified parasite

Another parasite, a round, red-staining mass from i'5 to in
diameter, is found in red cells (Plate XXIX, figs. 93, 95, 99). With it
are associated bluish-green, crystal-like bodies (Plate XXIX, figs. 93,
95,97,98). Both the red masses and crystals may be present in
varying numbers (Plate XXIX, figs. 98, 99). The cytoplasm of the
host-cells is usually stained a deeper blue than usual. Many of the
smaller red masses have a well-defined blue area around them, which
suggeststhat this parasite possesses a cytoplasmic body (Plate XXIX,
figs. 94, 96). In stained films of fresh blood the red masses do not
seem to have much structure, but in films of blood kept for two days,
the structure consists of chromatic granules arranged peripherally with
one or two darker chromatic granules in the centre.

This parasite with its crystals was seen in fresh specimens. It
occurred in conjunction with drepanidia and T. loricatum.

FISHES*

Fresh-water fish of many different sorts were examined in the
Gambia and in the Congo. Parasites were found only in the fish
without scales, mentioned below.

Those infected were caught in a very small sluggish stream with
a very dirty bottom ; none of them were over seven inches in length
and none were infected with drepanidia.

Trypanosomes (Plate XXX).

Two “mudfish ” ( C /arias angolensis ) caught at Leopoldville in
the Congo Free State on December 9th and December 30th, 1903,
had actively motile trypanosomes in their blood which may be
divided into three types ; a small, a medium, and a long form. The
small type was identical in both, the medium was present in both, but
with slight differences, and the long form was present only in the fish
°f December 9th. The small form was very frequently seen in the
fish of December 30th, while in the fish of December 9th, only four
were present in a total of ninety-nine parasites. The medium form,
>n the fish of December 30th, was less frequently met with than the
small form, but was often seen in the fish of December 9th-
Of the long forms only thirteen were seen. All the forms were
characterised by the possession of a large four-lobed blepharoplast,

*The description of the parades found in fishc-s is republished by permission from the
Journal of Medical Research.

336

situated at, or very near, the posterior extremity, and by a dear space
about the nucleus. The blepharoplast apparently consisted of four
darkly-staining granules of equal size embedded in a matrix. In a
few specimens there were, perhaps, more than four granules in the
blepharoplast, but in these cases it was never possible to be certain
of the exact number. The three forms of parasites seemed but
different variations of one species, as gradations between all forms
could be seen.

The small type was readily distinguished by its size, its narrow
body, its relatively long nucleus situated more anteriorly than in
the other two types, and with its long diameter parallel to that of the
body — and by its faint-staining reaction. The blepharoplast was
oval, or round, large and distinct (measuring 113 by 075/*). and was
situated at the posterior extremity. The undulating membrane was
well developed, having a width of 075 /*. The long, oval nucleus

was situated at the junction of the middle and anterior thirds of the
body. It was granular in structure and contained a fibrous network
on which were seen eight to sixteen chromatin granules, and one
darker granule — the karyosome. The body protoplasm was
reticular, and contained a few violet-staining granules. Occasionally
parasites were seen in which the body was filled with these coarse
chromatophilic granules. Superficial longitudinal striations to the
number of five could be seen near the nucleus of two slightly injured
parasites. Division forms were very numerous in this type (even
when but four were seen in the fish of December gth, one of them
was dividing). I hey followed the usual method of longitudinal
division. In the division of the blepharoplast, the posterior lobe on
the concave side of the parasite moved behind the other postenor
lobe ; the axis of the posterior lobes thus became longitudinal, while
that of the anterior lobes remained transverse. The two posterior
lobes then separated from the anterior lobes and their axis became
transverse again.

The medium type was characterised by its size, and its darker
staining reaction. The blepharoplast was large, oval or round,
distinct and measured about 1-5 by 075 /x. It was situated at the
posterior extremity, or from 1 to 1*5// anterior to it. The flagellum
was relatively longer than in the short form. The undulating
membrane in parasites from the fish of December 30th was well

337

developed, and wound in numerous folds around the body ; it was
r^fi wide. In trypanosomes from the fish of December gth, the
undulating membrane was poorly developed, had a few folds and
was not more than i /x in width. The nucleus was round to oval, its
long diameter was at right angles to that of the body, and it
occupied the whole width of the parasites. It was granular in
structure and contained a fibrous network on which were seen
sixteen to twenty-four chromatin granules and one or two darker
granules- the karyosomes. T he body was more darkly stained in
the fish of December 30th. Superficial longitudinal striations to the
number of seven were counted near the nucleus. Divisional forms
were few in this type and followed the usual method of longitudinal
division.

The long type was characterised by its size, its tapering
extremities, its comparatively short flagellum and by the possession
of from one to four vacuoles just anterior to the blepharoplast. The
blepharoplast was large, oval or round, and situated 2/x from the
posterior extremity ; its measurements were the same as those of the
medium type. The nucleus had the same characteristics as that of
the middle type, except that it did not quite occupy the width of the
body. The protoplasm was the same as in the medium type, except
for the above mentioned vacuoles, which were always present. Eight
superficial longitudinal striations were present and these were seen to
cross the nucleus. No divisional forms of this type were seen.

The measurements of the three types were as follows :

Small form

Medium form

Long form

Measurement I

... O’O /A

... o-o to 1*5 /X

... 2'0

/*

II

... 127 IX.

I4’2 /*

... 25-0

/*

„ HI

... 3'°

37 /*

... 45-0

/*

IV

... 67 /z

12 ’O [J.

... 19-5

/*

„ V

... 12*0 fX.

I3'5 M

... 10-5

/*

VI

I'O /X

3*o/*

••• 4'5

/*

Total length ...

■ 34' 5 /*

45'° /*

... 6i‘5

/*

Although the trypanosome described by Montel1 closely resembles
our large form, we have seen no account in the literature of any
parasite morphologically identical with those described above.

33«

LITERATURE

Good lists of the publications on the trypanosomata of fishes will be found in
the works of Laveran and Mesnil, “ Trypanosomes et Trypanosomiasis,” and of
Mense, “ Handbuch der Tropenkrankheiten."

The following are important papers not mentioned in these volumes:-

1. Montel. Trypanosoma d’un poisson de Cochinchine. Comptes Rendus de

la Societe de Biologie, T. I, 1905, p. 1016.

2. Lebailly. Recherches sur les Hematozoaires des Teldostdens marins. These

Fac. Med. Paris, 1906, p. 76; and Archiv. de Parasitologie, T. X, 1906.

3. Lebailly, M.C. Trypanosoma bothi iBarbue; n. sp. Comptes Rendus de la

Societe de Biologie, T. IX, 1905, p. 304.

4. Hri mpt. Sur quelques especes nouvelles de Trypanosomes parasites des

Poissons d'eau douce. Comptes Rendus de la Societe Biologie, T. LX, 27th
Jan., 1906, page 160.

5. Brl mpt. Mode de transmission et Evolution des Trypanosomes parasites des

poissons. Comptes Rendus de la Society de Biologie,' 17th Jan., 1906, T. LX.
No. 4, p. 162.

t>. Balfolr. Second Report of the Wellcome Research Laboratories, Khartoum,
page 197.

A Spirochcete (Plate XXXI).

In the blood ol the fish of December 30th a spirochaete-like
organism was found which could be identified with no previously
described spirochaete.

It had a hair-like body; its length was usually 18/1; its breadth
at the widest part was O'6/i. It was widest at one extremity 01
towards the middle; in the latter case, the parasite tapered towards
both ends, but one extremity was always much thicker than the
other. The wider end was rounded, while the other end was drawn
out to a fine and slender point. The parasite usually occurred
singly and lay in a simple curve, forming three-quarters of the
circumference of a circle (Plate XXXI, figs. 1 and 2). Other forros-
coils (fig. 3), compound curves (figs. 4, 9-12) and spirals (figs. 5-8)
veie seen as illustrated. The coils were always formed by the
s ender end of the parasite. The spiral forms were long, measuring
10m _4 2 jn, and in some of these forms there was a suggestion of
membrane (figs. 5 and 6). The parasites were also seen in pairs
in groups of from three to many individuals as shown in figures
!3. 14-16. Ihese organisms seemed to consist of a deeply-staining
core and a lightly-staining periplast. Many of them stained
irregularly by modifications of Romano wsky’s method. The lighter
stained areas (fig. 2) occurred at irregular intervals and were of

339

irregular extent. The periplast could be seen at these lightly stained
areas and it was continued to form the slender extremity from which
the core was also absent. Occasionally granular forms were seen in
which the protoplasm of the core apparently arranged itself into small
granular masses, but no regularity could be detected in the arrange¬
ment of these granules. Undoubted multiplication forms were not

seen. , f

For descriptive purposes we propose for this organism the name o

Sprochceta jonesii.

ARTHROPODA

The protozoa found in this class were encountered by
chance during an investigation of the development of various
haematozoa, while examining various blood-suckers, either as controls,
or for the presence of developmental forms of the blood parasites.
Many tsetse flies and mosquitoes, a few Stomoxys, ticks o vanous
sorts, fleas and lice, and “ Congo floor maggots," were examined.
The only protozoan parasites observed are recorded below.

MOSQUITOES

(1) Myzorhynchus paludis .

A cluster of radiating club-shaped bodies (Colome radiee) was
seen in the thoracic tissues of a female of the above species, examined
for malaria at Lusambo. The parasites became free, deveioped a
flagellum and were actively motile in the salt solution in which
mosquito was dissected.

(2) Pyretophorus costal is. . ,

Some excitement was at first created during the examination o

the alimentary canal of mosquitoes, fed at varying antecedent periods
on animals infected with trypanosomes, by the presence o^vcnmcu
like bodies (Plate XXXII, fig. 14). measuring about iS by /*■
These parasites progressed slowly by active lashing and slower
amoeboid movement ; some similar foims weie motion ess ,

The amoeboid movements were of two sorts. The fast mvcflved
a change in the whole shape of the parasite. T e secon
a simple protrusion of protoplasm from the rounded anterior,
from the effilated posterior extremity.

340

They occurred either within the stomach, in the stomachal tissues,
or in the body cavity of the mosquito. A central differentiated area
could be detected in them by examination in fresh specimens; near
the area were more or less numerous granules which often exhibited
lively Brownian movement. After some time the parasites became
much stumpier (8/i by 16/i), and were later almost spherical. One
parasite, however, remained unchanged and active for twenty-four
hours.

Further search showed that the parasite occurred in freshly-
caught and freshly-hatched mosquitoes, and in the larvae from which
all our experimental mosquitoes were raised, and it was quickly shown
to be a stage in the life cycle of a coccidium. It was most interesting
that practically every larva from one pool was infected with this
coccidium, while none of the larvae and adults (anopheline and
cuhcine) from a pool only a few yards distant were affected. So far
as was observed the parasite did not occasion any excessive mortality
amongst the mosquitoes infected. This seems difficult to understand-
the mosquitoes were probably not watched long enough— since the
tissues of many of them were fairly riddled with coccidium cysts.
Hie water of the heavily-infected pool was centrifugalised and
examined. Coccidium cysts were not recognised, various infusoria
and a clump of herpetomonas-like flagellates were alone seen.

So soon as it was certain that the vermicule-like “sporozoite” hac
no connection with the trypanosomes ingested, the study of this
coccidium was discontinued. It is, therefore, not possible to say
w ether the forms observed represent stages in more than one
^ °r no' ^ seems probable that but one species of coccidium

s present. We describe the forms observed and indicate the
position they seem to occupy in the life cycle of the parasite. The
mobile sporozoites first seen become free through the rupture of a
sporoblast containing eight sporozoites. The number of sporozoites
m each sporoblast is almost invariable. Fig. 15 is unusual in that
nuc eus o two of the sporozoites has divided, and in one instance
sion o t le cytoplasm seems to have commenced. As a rule the
ong .ameters of the sporozoites are parallel. These are, therefore,
gu arly arranged like the segments of an orange (corps en bardlet).
Single sporoblasts surrounded by a definite cyst wall may occur,
y are more usually seen in groups of four, seven, or even as many

341

as eleven sporoblasts packed together in an oocyst with a definite
capsule. The individual sporozoites have no capsule. Their
cytoplasm stains a light blue by Romanowsky's method. It is
alveolar in structure and rarely contains granules of any sort, but
may have occasional granules. The nucleus is placed centrally, is
loose in texture and consists of a varying number of chromatin
granules placed in a more lightly-staining matrix. From the analogy
of other coccidia, we assume that the forms just described represent
the completion of sporogony. The male and female gametes, which
probably commence this cycle, and their conjugation have not been
seen. Neither has the further development of the sporozoites been
observed.

The number of rounded merozoites (figs. 16, 17, 18), present in
every part of the larvae and adults dissected, was often extraordinary.
These organisms measured about 4 to 5/1 in diameter. They were
either free or enclosed in a host cell. On becoming extracellular
they were seen to possess a definite capsule (pink-staining) ; this was
quickly discarded (figs. 1 8, 19) and the empty shells were very
frequently seen. Often merozoites which had just cast their capsules
occurred in groups of four (fig. 18). Four merozoites were, however,
never seen within an unbroken capsule. As a rule whether intra or
extra-corpuscular the merozoites occurred singly. When extra-
corpuscular they were occasionally in pairs. Sometimes large intra-
corpuscular cysts were seen which contained very large numbers of
merozoites. The merozoites (figs. 16, 17, 18) consisted of a pinkish
alveolar cytoplasm in which was situated a nucleus usually surrounded
by an indefinite bluish area. In this bluish area a lighter, rounded
spot was almost invariably present. A few chromatophilic granules
and small clear spaces were frequently seen in the cytoplasm. The
nucleus consisted of irregular masses of chromatin placed in a
matrix. In some forms (fig. 16) the granules were placed at the
Periphery of the nucleus and were joined by threads with a central
darker-staining mass. In fresh specimens they were immobile.

LITERATURE

'• Ross. Parasites of Mosquitoes. Jour, of Hygiene, Vol. VI, No. 2, April
'9°^. The subject is reviewed and discussed.

342

TSETSE FLIES (G/ossina palpalisi

Parasites of the type described below were found in specimens of
Glossina palpalis freshly-caught, and at all periods, up to n days,
after having fed on known animals infected with trypanosomes. All
the parasites were not precisely similar, but one description will suffice
for all. The free rod-like parasites were found only in the
alimentary canal, nearly all of them in its anterior part. They were
most numerous near the oesophagus, in the “ stomach or in the
neighbouring part of the intestine. Cysts containing them were seen
attached to, or actually in, the wall of the intestine. Some of them
were whole, others had burst (or were burst by slight pressure), and
from them were expelled the rod -like bodies. In fresh preparation^
these rods (Plate XXXII, figs, i, 2, 3, 4, 5, 8, 9, 10) were non-motile,
they contained one or two refractile granules and they occurred singly
or, occasionally, in small clumps. The most usual form seen in stained
specimens was a rod about j'Op in length and VOfi in width wi
abruptly rounded ends (figs. 1, 2, 5). It was limited by a definite
outline, but apparently not by a capsule. Its body substance con=
of pale blue protoplasm in which occurred chromatophilic granu 3 ^
rounded clear areas. The chromatophilic granules varied ni^
from extremely fine particles (fig. 6) up to masses measuring * ^
o'3 to 0-5 p in width (fig. 3). They were usually distributed moi ^
less regularly in transverse bands (figs. 1, 2, 5. 10) lying. at
across the parasite. In such parasites the individual granule^ ^
barely be distinguished. Often between each band of granl guCj,
placed one of the rounded, clear areas (figs. 1, 2, 5. U ' sn]all
parasites were probably preparing for transverse division. 1 ^
forms, apparently consisting of one or two rounded se?1110’1^ j the
original rod-like body, were often observed (fig. 7). In xVell aS
parasites showing the band-like distribution of chroniatm- d=bl]ateral
in others in which pairs ol granules were arranged 111 jBjglrt
symmetry (figs. 8, 9), it seemed as though longitudinal dlVl51^s
be about to occur. In many parasites the chromatin gr‘‘n } jarger
distributed absolutely without order (figs. 3, 4, 6). Rarely otli^
and a smaller granule seemed to be in connection with eaC ueptly

freq

i«e'

Rod-shaped parasites of all these three types wcrc ^ ($0$
terminated by a small chromatophilic granule (figs. 4. 5’ 1 ^ wh*c
times placed in a clear area), or by a rounded enlarge111

343

often measured twice the diameter of the rest of the parasite (fig. io).
Rounded parasites measuring from 3 /* to 5// or more in diameter with
irregularly distributed granules and no clear areas occurred (figs. 1 1
and 13). The smaller rounded forms can divide transversely ;
apparently after a concentration of the chromatin (fig. 12).

It seemed possible that this parasite should be placed amongst
the Myxosporidia. In recognition of the collaboration of Dr. Inge
Heiberg in the work of our expedition, and in particular of his
observations on T. loricatum . we suggest for the name of this parasite
Myxosporid turn heibergi. About half of the tsetses dissected at
Leopoldville had these parasites free in their midgut. In the stomach
wall of one freshly caught fly coccidian-like cysts were seen. (“ Corps
en bardlet compare the parasite of Pyretophorus costalis described
above).*

EPILOGUE

Although the facts presented in this paper are admittedly the
results of passing observations, incomplete, disconnected, and made
under unfavourable circumstances, we think that they possess a
certain value as a record alone of protozoan infections encountered by
chance during three years’ work in Africa. Many of the appearances
noted, though incomplete in themselves, are of additional value in
that they confirm the constancy of morphological changes noted in

other allied and better studied parasites.

The following examples are cited. Longitudinal stnations have
been seen in every trypanosome examined ; the myonemes of ot er
authors are, therefore, constant in the trypanosomes we have dealt
with. In several instances the thickened edge of the undulating
membrane of trypanosomes has been observed to terminate, not in
the blepharoplast, but in a lighter-staining pinkish granule, or pair o
granules, closely applied to it. In some of our trypanosomes
structure of the nucleus and the occurrence, m neighbouring
differentiated areas, of paired chromatophilic granules suggests a
connection with the complicated nuclear developmental phenomena
observed in trypanosomes by some observers. The clear, unstame

•This is the last of the parasitic protozoa

tarabia and the Congo. The filariae found in va by5 Professor. H. E.

nd reptiles will be considered in a forthcoming paper o>

tnnett.

344

perinuclear area, the irregular, non-staining almost granular areas, and
the lines in the cytoplasm as well as the chromatophilic granules (see
Plate XXVII, fig. 34) frequently observed in various trypanosomes
described in this paper, are all constant phenomena, which
seem to be of biological significance. In the development of
T. loricatum the work of Danielewsky was confirmed and extended
to prove a most interesting cycle of multiplication in a well-known
trypanosome ; it must be asked whether a similar cycle may not occur
in other trypanosomes. The rough resemblances between the
spirochaetes seen in ulcers and in fish with previously described
parasites of this nature are striking. In the leucocytozoon of birds an
interesting course of development is described. * Unfortunately our
material was too scanty to permit us to fully compare our observations
with those of Schaudinn. Very little has been said concerning the
drepanidia observed. In spite of a very careful search in kept pre¬
parations of blood, we were unable to find any indication of a direct
relation between this parasite and the trypanosomes which were
often associated with it. t Neither were parasites resembling
1 rypanosoma inopinatum seen to become intracorpuscular. X In both
stained and fresh preparations of frogs blood, however, the great
resemblance in size and general appearance between some trypano¬
somes and some drepanidia was very apparent. It was also noticed
that, as a rule, if there were many drepanidia in a given frog, there
were also many trypanosomes.

One of the earliest of the lessons learned from our work on these
protozoa was the entire inadequacy of the methods of preparing
blood-films, ordinarily employed by pathologists, for a morphological
study of protozoa.

The examination of dried films stained by modifications § ot
Romanowsky’s method suffices for the purposes of clinical diagnosis.

I his method will show many of the structures present, it was used in

* The dangers of constructing a part of the life-history of a parasite from
stained specimens alone are apparent. The course of development we described
ma\ be mistaken, but the descriptions of the forms mentioned are accurate and ma)
e relied upon so far as the defects of the method of preparation employed permit

1 Billet, ( ulture d’un trypanosome de la grenouille chez une Hirudinee. C. R-
Acad, des Sci.,1. CXXXIX, Oct. .0, 1904, p 643.

c ^ rypanosoma inopinatum et sa relation avec les Drepanidium. C. R-

boc. de Biol. , July 23, 1904, p. i6x.

Pres! of Th erpoc)1!11 Christophers’ The Practical Study of Malaria, page 10, University

345

the whole of the work described in the present papers, but it fails to
demonstrate finer details properly. Our work must, therefore, be
regarded as incomplete, and certainly the more important parts of it
should be repeated by workers using methods more perfect than those
employed by us.

A second lesson was that too much of the work done on the
pathogenic protozoa, particularly by medical men, has been directed

by conceptions derived from bacteriology.

Days of tedious searching of slides and of observation of parasites,
placed under various conditions, must be spent in studying the
pathogenic protozoa, when minutes would almost suffice in the case of
bacteria. The difficulty of finding the parasites at all is sometimes
extraordinary, and the possible occurrence of a latent infection must
never be forgotten.

The study of the pathogenic protozoa must be approached with
an unbiassed mind and with the remembrance that the known life¬
cycles of several protozoa are exceedingly complicated. We believe
that the continuous observation of living parasites will ultimately
furnish the richest reward. A single positive observation so
obtained is absolute, and outweighs any number of apparent y
antagonistic probabilities obtained by deductions from work done
along apparently parallel lines of research. Of course, the examina¬
tion of fresh preparations should be supplemented by the examina
tion of stained material. Lastly, at the present moment more is
known, in every way, of malaria than of almost any other disease,
observing less studied protozoan infections it will fiequently happen
that our knowledge of what actually does occur in malaria will lea
to the formation of an ultimately successful working hypothesis, or to
the correct interpretation of newly-observed phenomena.

EXPLANATION OF PLATES

All the illustrations accompanying this paper, except where it is
otherwise stated, are drawn to a magnification of 2,000 diameters.
No camera lucida was employed ; the dimensions were obtained by
measurement. A Zeiss 2mm. apochromatic i'40 aperture objective,
with an 8 or 18 eyepiece, was regularly used.

Plate XX

Figs. 1 to 28, except 27. are “Young Forms”

As a rule, ectoplasm, endoplasm and nuclear material can be
distinguished in each of these parasites ; occasionally it is difficult to
do so (fig. 24). The arrangement of the parasites and the host-cell
are well seen in figs. 1, 25, 28. The constant presence of a clearer
area about the chromatic material is to be noted in these parasites.

F'fT- i- Chromatic mass with linear extension possibly antecedent to
nuclear division, as in fig. 3.

Fig. 2 Shows nucleus with vacuole containing indistinct granules.

3- Nuclear material divided into a small and a larger chromatin
mass connected by a line ; possibly stage succeeding fig- 1

Figs. 4, 5 and 9. Chromatin divided into a larger mass and a smaller
granular one, placed in a vacuole. Note the vacuole in
these parasites is frequently pink-staining.

Figs. 6 and 7. — F orms similar to 2 and 5 in very small parasites.

Fig. 8. Very small parasite showing a line.

Fig. 10. Chromatin divided into two equal masses.

Figs. 11 and 12. Are squashed and somewhat degenerated parasites.

They are useful in that they show distinctly the presence
of definite structure in the nuclear material.

347

Fjg j j —Shows a form in a small parasite probably comparable to 9.

The vacuole is well marked. . The endoplasm of the
parasite has pressed upon and considerably distorted the
nucleus of the host-cell.

Fig. 14— Probably not similar to 1, but commencing formation of line.

Figs. 16, 1 7 and 18.— Early formation of line. Note its position in
the chlamydoplasm.

Figs. 19, 20, 21 and 22.— Are unique specimens. They probably
represent important stages of the maturation of the
parasites. We feel it unwise to attempt to discuss their
exact significance. In 22 a wavy bluish-line is disposed
along “ the line.”

Fig. 15. — Is a squashed and degenerated parasite. The immediate
connection of at least one pair of chromatophilic granules
with the line is very evident in 15, 19 and 23.

Figs. 23, 24, 25 and 28. — Are well-grown parasites which retain
their spherical form.

Fig. 26.— Shows commencing transverse division of the line. The
connection of the “ blepharoplast with the blunt
extremity of one half of the line is well shown.

Fig. 27.— Line shows commencing longitudinal division. Note that
there are two distinct areas of chromatophilic material,
beside the chlamydoplasm, lying immediately beneath the
dividing line. (This specimen was unique.)

Figs. 29 and 30. — Trypanosomes.

% 3 1 -—Spherical female. Note dark granule connected with line
running across chlamydoplasm ; this is the only instance in
which such a line was seen. The ectoplasm and host-cel
nucleus are being thrown off.

34»

Plate XXI

With the exception of fig. 38, probably a female, all the parasites

illustrated in this plate are males.

Fig- 33- — Division of the line; nucleus only just commencing to
divide.

Fig- 33a- — Division of line with chlamydoplasm and nucleus. The
ectoplasm has been deleted for convenience of reproduc¬
tion.

Fig- 34- — Formation of line (?). Note the presence of granules of
chromatin.

Fig- 55- — A very common form (see figs. 42, 43, 74). It represents
the line passing through a diffuse chromatic area
(chlamydoplasm) in which lies a denser chromatic area
(nucleus ?), often containing one or more (figs. 42, 41)
deeply chromatic granules (blepharoplast ?). Occasionally
these granules, or similar ones, lie outside the denser
nuclear material (fig. 43).

Fig- 3^. — Degenerated and squashed parasite, showing multiple
nature of line.

Fig- 3 7- — Only three parasites of this type (all practically identical)
were seen. It is remarkable in possessing a blepharoplast-
like mass of chromatin distinct from the main nuclear
structure. 1 here is an indefinite arrangement of fibrillae
and granules about the nucleus and chlamydoplasm.

F'g- 38- A female cell (unique), in which a faint blue line was wound
spirally about one half of the line. Appearances
resembling this were seen in one or two other preparations
(fig. 62).

F’g- 39- Fine dividing longitudinally, nucleus already completely
divided. Parasites of a similar type are not infrequent.

The one illustrated is considerably distorted.

IH1gs. 40 and 41. — Commencing transverse division of the line. In
each parasite the line has become thinner. In fig. 41 the
differentiated area surrounding the constriction is well
seen.

Figs. 42, 43 and 74. — Are ordinary types of adult males with lines.

rigs. 41, 42 and 43— Blepharoplastic (?) granules occur in the two
former within, in the latter without, the nucleus. The
presence in fig. 43 of a bluish area within the chlamydo¬
plasm in immediate connection with the nucleus is
suggestive.

Fig- 44- The line is divided transversely (specimen unique); the
two halves are connected by a filament.

45- (Compare figs. 63 and 64). Probably represents a stage

Fi r antenor 46. (Specimen unique.)

g. 40. l he fine is much involuted. It extends far into the
ectoplasm. A faint blue spiral line (artefact ?) extends,
liom a double granule in the ectoplasm, through the lower
part of the parasite. The arrangement of the nucleus is
not at all definite.

350

Plate XXII

Fig. 47. — (Specimen unique. The colouring of this parasite is much
too vivid.) Probably occurs at almost the same period as
fig. 49. It is remarkable for the four granules placed in
the concavity of the crescentic nucleus. (Similar dots
have been seen in other parasites, but never again in this
position).

Figs. 48, 49. — Are very common types of males. Note the spherical
nucleus with its neighbouring granules in 48 and the
distributed chlamydoplasm in fig. 49. (Compare fig. 75-)

Fig. 50. — Shows a disjointed bit of chlamydoplasm in relation with
the blepharoplast. Note the granules in this position
The alveolar structure of the endoplasm has given the
parasite a lattice-like appearance.

Figs. 5!> 52> 53 and 55. — Are stages in the division of the nucleus. In
fig. 53 the vacuole in association with the blepharoplast
is well seen.

igs- 54> 56 and 59. — The chlamydoplasm is abundant. The nucleus
has formed a spindle-like figure, best seen in fig. 56- ^
fig. 59 the chromatic granules of the figure seem about to
divide, and the vacuole in connection with the blepharo¬
plast is well seen.

Fig- 5 7- Two tiny granules are attached to the blepharoplast by
fine threads. The nuclear chromatin is commencing to
show polar concentration. (See fig. 60.)

Fig. 58. Granules are being extruded (?) from both nucleus and
blepharoplast.

big. 60. Is probably the stage succeeding fig. 5 7. The nucleus ha^
almost divided.

Fig. 61. Shows enormous increase of chlamydoplasm. (Compaq
56.)

352

Plate XXI 1 1
x i>334-

Figs. 62-73, except 68, 70, and 71 are females.

Fig. 62. Is a stage in line formation. Note the wavy blue line
crossing the chlamydoplasm.

Figs. 63 and 64. — Compare fig. 45.

Flg- 65.— Note endoplasm lying over host-cell nucleus. Chlamydo¬
plasm does not stain, so nucleus appears to lie in a vacuole.

Fig. 66. Host-cell nucleus extruded, nucleus well differentiated
Note granules in chlamydoplasm.

P*ff- 67. Host-cell nucleus about to be extruded. Compact
chlamydoplasm, well-defined nucleus (or blepharoplast ?).
This is a very common type.

Fig. 68. (Compare figs. 48, 57, 71, 75.) Chlamydoplasm with
spherical nuclear area. Blepharoplast very granular and
one granule extruded. (Compare fig. 49.)

1 ig. 69. (Compare fig. 67.) Is a common type. The chlamydoplasm
is very diffuse.

big. 70.- -Parasite possessing distinctive characters of neither adult
male nor female.

I lg ‘ z1-— Note deeper-staining area in nucleus. (Compare fig. 68.)

Fig. 72.— Much effilated makrogametocyte ; in one instance two fine
chromatic granules occurred, as illustrated, in the
chlamydoplasm.

S 73 ■ Is a common type of adult female; the vacuole in connec¬
tion with the blepharoplast is well seen.

Fig. 74. — Compare fig. 35.

Pig- 7 5- (Compare fig. 48, &c.) Note the two large ill-defined
granules outside the nucleus in the chlamydopl asm.

S 76. (Compare fig. 49.) Is paJc] y stained ; chlamydoplasm is
much extended.

*> 77- Discarded ectoplasmic sheath and host-cell nucleus of an
adult parasite.

PLATE XXIII.

354

Plate XXIV

Pig- I- — “ Slender ” form.

Fig. 2. — “Slender” form.

Fig. 3. — “ Broad ” form.

Fig. 4.—“ Broad ” form.

Fig. 5. — “ Stumpy ” form.

Fig. 6. Stumpy form ; evidently degenerated but reproduced to
show structure of blepharoplast and nuclear granules.

Dutton, Todd and Tobey.

PLATE XXIV

Trypanosomes.

35^

Plate XXV*

Fig. I. — T. loricatum. This is apparently an ordinary division form
with two blepharoplasts, two nuclei and two flagella. The
nuclei are long, sharply-defined sacs, with many small
chromatin granules in them.

Fig. 2. — T. loricatum This form has a long nucleus with chromatin
massed near both ends. There are longitudinal pleats
and a short free undulating membrane.

Fig. 3. — T. loricatum. This form has a round nucleus. The longi¬
tudinal pleats are present.

Fig. 4. — T. loricatum. This form has deep longitudinal pleats. The
nucleus is not so long as in figs. 1 and 2, but has the
characteristic shape and curves towards the undulating
membrane.

Fig. 5. — T. loricatum. Lanceolate form. The nucleus is a long and
broad band. The undulating membrane runs as a white
band in folds down the middle of the parasite.

Fig. 6. — T. loricatum. Long “ hyla form” (page 324). The body
has longitudinal folds. The nucleus is long, and wide
near the middle, tapering to both extremities.

Fig. 7. — T. loricatum. Long “ hyla form.” The folds are unfolding
No nucleus is visible.

Fig. 8. — T. loricatum. Parasite has become rounded and is about
to lose its flagellum.

Fig- 45- — Trypanosome of the tortoise, showing round unstained
areas.

Fig. 46. — Drepanidia of the tortoise. The nucleus stains purple and
both it and the cytoplasm are denser than in fig. 4^-

Fig- 47- — Drepanidium of the tortoise. The nucleus is red and the
body has many coarse granules in it.

Fig. 48. — Drepanidium of the tortoise. The nucleus is red ar.d
loosely woven, and the cytoplasm is also loosely woven.

Fig- 49- — Drepanidium of the crocodile.

big- 5°- — Drepanidium of the snake — common form.

Fig- 51- — Drepanidium of the snake. The nucleus and cytopL-
are loosely woven, and vacuoles are present at both en -

Fig- 52- — Drepanidium of the snake, showing elongation of the bod}
of the host-cell.

Fig- 53- — A curious appearance in a snake’s blood (page 304); lre'
form, red dot in the middle.

big- 54- The same in a cell with no drepanidium.

big- 55- — The unidentified parasite of the snake — round form.

* Numbers 1-8 are drawn one-half the usual size.

J

.

, *.

358

Plate XXVI

Pig- 9- — T. loricatum. Wide “ hyla form,” with folds unfolding.
Nucleus indicated by a bent band.

1- ig. io. — T. loricatum. Short “ hyla form.” The nucleus is long and
has chromatin massed at both ends. There is an unfold¬
ing of the edge of the body at intervals on one side of the
parasite.

Pig. n. — 7. loricatum. Round form with short curved,. narrow white
band running from blepharoplast towards periphery. The
nucleus has differentiated areas on both sides.

Pig. 12. — 7. loricatum. Round form, dividing form. Two nuclei,
and two blepharoplasts are present.

Pig- T3- — T- ■ loricatum. Round form as in fig. 1 1, but with no
differentiated areas in nucleus.

Fig. 14. — T. loricatum. Rounded form, about to divide, the flagellum
having been lost.

l ig. 15. — T. loricatum. Dividing round form; two nuclei, two
blepharoplasts, but the cytoplasm not quite divided.

Pig. 16. — 7. loricatum. Small round division form.

Pig. 17. 7. loricatum. A still smaller form than the above.

P ig- 18. 7 . loricatum , I he parasite has divided into a group of 16.

Pig. 19. — 7. loricatum. Rounded form with division of nuclei and
blepharoplasts into four, but with no division of the
cytoplasm.

Pig. 20. 7. loricatum. Small divisional form that has just acquired

a flagellum.

Pig. 21. T. loricatum. Herpetomonas-like form. Note the position
of blepharoplast, nucleus, posterior granule, and flagellum.

Fig. 22. T. loricatum. Trichomonas-like form. The anterior
extremity is enlarged and round and from it go four
flagella.

Pig. 23. — T. loricatum. Pferpetomonas-like form with large anterior
end.

P ig. 24. 7. loricatum. This is perhaps a variety of T. inopinatum.

P^g- 2 5- T, loricatum. I his is a common inopinatum- like form.

1 'g- 2b. T loricatum. An inopinatum -\ ike form, wide at the level
of the nucleus.

P 'g- 2 7- T- ■ loricatum. A sanguinis- like form. Note the wide
membrane, the position of the nucleus and the blepharo-
P asP and the shape of the posterior extremity.

P ' g- 28.-7’. loricatum. A “ leaf-like form ” (page 322). Note the
position of nucleus and blepharoplast, and the shape of
posterior and anterior portion of the body.

PLATE XXVI.

P.P. Press. I tu I'.

Plate XXVII

J'ig- 29. — T. loricatum. “Leaf-like form” (page 322), with the
posterior end arranged like a bud about to open. Note
position of nucleus and blepharoplast.

Fig- 3°- — T. loricatum. “ Leaf-like form unfolding. Note position
of nucleus and blepharoplast, and shape of anterior and
posterior portions of body.

Fig- 31- — T. loricatum. The “leaf-like form” has completely
unfolded into an adult type of T. loricatum.
ig- 32- — Intermediate form between the “leaf-like form” of T.
loricatum and T. mega.

Fig- 33- — T. mega. A form with red staining granules in the cyto¬
plasm. I he nucleus is faintly stained.

Fig- 34- — T. mega. Same form as fig. 33, but with more red granules
in the cytoplasm.

1'iiT- 35- — F\ mega. The posterior part of the body is irregularly
contracted. Note the shape of the nucleus.

Fig- 36.- 7'. mega. A somewhat generally contracted form. Note
the shape of the nucleus.

Fig- 37- T. mega. A very much generally contracted form. Note
the shape of the nucleus.

Fig- 3^- T. mega. A variety of T. mega with coarse reticular
structure.

Fig- 39. T. mega. A much elongated form. Note the character of
the nucleus, and the difference between the part of the
body anterior to the nucleus and that posterior to the
nucleus.

plate XXVII.

^7"

362

Plate XXVIII

Fig. 40. — Small form of T. karyozeukton , showing blue spiral line
running from blepharoplast to posterior extremity, and the
thick wide undulating membrane.

Fig. 41. — Medium form of T. karyozeukton , showing granule at
posterior end of thickened edge of undulating membrane.

Fig. 42. — Large form of T. karyozeukton.

Fig. 43- — T. karyozeukton coiled, with the anterior end free.

Fig- 44. — Tighter coil of T. karyozeukton.

Fig. 56. — A curious appearance in a snake’s blood (page 304) in a cell
with a drepanidium.

Fig 57- — J he same lying upon a degenerated drepanidium.

Fig. 58. — The same with short rods of unequal length.

Fig. 59- — Intracellular drepanidium, showing excretion at both
extremities within capsule.

Fig. 60.- Encysted drepanidium, found free in serum, showing
excretion at the extremities.

big. 61. — Division form of drepanidium. The cytoplasm has divided
synchronously with the chromatin.

Fig. 62. — Division form of drepanidium, dividing into two.

Fig 63. — Division form of drepanidium, with the divisions almost
separated.

Fig. 64. Division form of drepanidium, five divisions.

big. 65. — Division form of drepanidium with four divisions.

Fig. 66. Division form of drepanidium on the way out of the
erythrocyte.

big. 67. Common form of drepanidium, almost out of an erythro¬
cyte, showing the eaten-out character of the host cell.

b ig. 68. Common form of drepanidium, showing division of nucleus
into two, connected by a red line.

Fig. 69.— Common form of drepanidium with dividing nucleus.

big. 70. Common form of drepanidium with constriction at one end
tbe body» showing one method of movement

*g‘ 71- Common form of drepanidium with two constrictions.

^2‘ ^ oung parasites of the small form of drepanidium.

73- -Small form of drepanidium, showing nucleus and the blue
spiral line running from a chromatin granule of nucleus
to anterior extremity of body, x 4,000 to show structure
in detail.

^ 74- Small form of drepanidium, showing chromatin granules

at posterior end, also a few anterior to nucleus, x 2,000.

^ ’fa- 75- Free division form of drepanidium.

364

Plate XXIX

i'ig. 76. — Division form with one larger mass of chromatin and
several chromatin dots around it.

^ 77- — Division form with four masses of chromatin.

hig. 78. — Division form with sixteen masses of chromatin.

Fig-- 79 ■ Division form with twelve masses of chromatin and the
division into well-defined " spores.”

I- ig. 80. — Double infection with rosette division forms.

big. 81. Common form of drepanidium with division form beside it
in same host-cell.

big. 82. Common form of drepanidium with division form at an
earlier stage of development.

FjfT- S3.— Young form of large drepanidium.

Fig. 84. Drepanidium showing one globule situated near the
nucleus.

b ig. 85. Drepanidium with about 100 retractile globules in it.

big. 86.— Large form of drepanidium.

Fig. 87. Large form of drepanidium, showing body folded upon
itself.

Fig. 88.
Fig. 89,
Fig. 90.

Fig. 91.
Fig. 92.-
Fig. 93--
Fig- 94-

Fig. 95-
Fig. 96.-
Fig. 97.-
Fig. 98.-
Fig. 99.-
Fig. 100.
Fig. 101.
Fig. 102.-
Fig. 103,
Fig. 104.-
105.-
Fig. 106.
Fig. 107.-
Fig. 108.-
Fig. 109.
Fig. no.
Fig. in,
big. 112.-

I'Z- 1 13 —

Fig. 114.-
Fig. 115.—

Large form of drepanidium, only partly folded upon itself.
—Large form of drepanidium with 34 retractile globules.
Large form of drepanidium with two globules situated
near nucleus.

Drepanidium with 1 1 globules situated near the nucleus.
—Drepanidium with large retractile globules.

An unidentified parasite of frog with crystal.

An unidentified parasite of frog, showing a blue area
sharply defined.

-An unidentified parasite of frog, with crystal in nucleus.

” .» with well-defined blue area.

” .» apparently free in the serum.

•• .1 with three crystals.

' ” •. triple infection.

-Lytomoeba free in serum.

-Cytomoeba in cell.

round form.

round and long forms,
long form.

round forms breaking up nucleus.

round form breaking up nucleus.

long form.

long form.

round form.

long form.

366

Plate XXX

Hg\ i. Small form, showing arrangement of the chromatin granules
of the nucleus and of the granules of the blepharoplast.

h ig. 2. — Small form, showing early stage of division. The chromatin
granules of the nucleus have gathered into two masses,
and the granules of the blepharoplast have changed their
position.

^ iff- 3- Medium form, showing arrangement of the chromatin
granules of the nucleus and of the granules of the
blepharoplast.

^ 4- Medium form, showing striations.

Pig. 5. Long form, showing arrangement of the chromatin granules
and karyosome of the nucleus, the longitudinal striations,
the granules of the blepharoplast and the vacuoles anterior
to it.

5

Duttor.Todd.Tobey

plate XXX

Hemoflagellate.

4

I

368

Plate XXXT

S pi roc ha et a joiiesii from Cl arias angolensis , see page 338.

^'onJodd.Tobey

PLATE XXXI

Hemoflagellate

YAWS

373

YAWS

BY

C. VV. BRANCH, M.B, C.M. (Edin.),

MEDICAL OFFICER, COLONY HOSPITAL, ST. VINCENT, W.I.

(. Received May 13th, 1907)

THE PREVALENCE OF SYPHILIS IN THE
WEST INDIES

The most striking feature of the medical practice in the West
Indies is the prevalence of tertiary syphilis and infantile inherited
syphilis among the labouring class. In some of the islands this
prevalence is quite extraordinary.

On the contrary, in the classification of diseases returned by the
Colonial medical officers, as well as in the reports of hospitals, syphilis
is credited with so few cases that it is incredible that the figures can
be correct. In part this is due to the custom of classing diseases by
the local manifestations, e.g. iritis, necrosis, paralysis, &c. The cases
are thus scattered through the classification, and even if their true
nature has been recognized it is not shown. But it is nevertheless a
fact that great misconception does exist in the minds of many of the
medical profession in the West Indies, and no doubt elsewhere in the
tropics, as to the extent to which syphilis is responsible for the sicknesses
that occur. The chief causes of this failure to recognize the disease
are ignorance of the original nature of syphilis and the tendency to
regard it as a venereal disease.

hailing to attribute a given lesion to syphilis the medical man
naturally turns to tubercle, the effects of which on the natives of the
ropics are in consequence vastly over-rated. Tubercular bone and
joint disease is rare among negroes. Lupus I have never seen,
though I have seen cases classed as such, the scars and subsequent
history of which abundantly proved the syphilitic origin of the lupoid
ulceration. Phthisis is very common among the negroes in some
^est Indian Colonies, but in view of the undoubted rarity of othei

374

tubercular troubles, 1 am inclined to believe that a great deal of the
phthisis is really syphilitic. 1 have several times been agreeably
surprised by the recovery of a phthisis case. Without actually
diagnosing- syphilis, one gets such a belief m iodide, and uses it for
such a wide range of complaints, that some of the lung cases have bad
the good fortune to be accidentally cured in this way. It is a sate
thing to treat all supposed phthisis with iodide if the tubercle bacillus
cannot be found or if there is no opportunity to search for it But
even after discounting for pure syphilis of the lung, there remains a
number of tubercular cases quite out of proportion to the amount of
tubercle of other tissues.

With respect to the West Indies I can personally vouch for the
prevalence of syphilis in the islands St. Croix (Danish West Indies;,
St. Kitts, Nevis, Antigua, St. Lucia and St. Vincent. In Barbados
I think there is not so much of grave syphilis, but the disease is
widely spread nevertheless. Dr. W. J. Branch, after 25 years
practice in St. Kitts, said in reply to a query, " nearly every black or
" coloured person on St. Kitts has, or has had, syphilis in some shape
“ or other, congenital, acquired or both."1 1 had five years
experience in district and hospital work in that island, and can assure
the reader that though this estimate is true, the condition of the
population of St. Vincent is worse. 1 lere I have seen a man in

middle life with tertiary scars dating from childhood, a scar of an old
penile chancre, and a new eruption of secondaries. That is to say, in
St. Vincent people may be “ thrice dipped ” in syphilis. Second
infections are the rule of life if an individual comes to mature years.
I he combination of secondaries and tertiaries is very commonly seen.
In my annual hospital report for 1902-03 I wrote: —

" When to this moral state is added a profound and universal
saturation with syphilis and depletion by ankylostoma, it may easily
be understood that the present labouring population of St. Vincent
1S leased and pauperized as any in the world. The effects of
syp ills depend on the nutrition of the patient ; so that where there
a soil as suited for the exuberant manifestation of the disease a=
° tams here’ 1S not surprising to note the disablement and
increasing pauperism due to syphilitic ulceration and necrosis.

-? _ >P ilitic paralysis, and degenerative neurosis. . • •

'■ Nlcholls’ Report, page 167.

375

•'Medical men from abroad, visiting St. Vincent, are struck by
"the prevalence of disfigured and noseless faces, and the pauper
“asylum is a museum of remnants left by syphilitic disease and the
"surgeon’s knife.”1 This is a lurid picture, but it is paralleled by
the condition of more than one other West Indian colony.

Dr. Blanc, of Tobago, bears witness to the extensive spread of
syphilis in that island. “ In connection with this question, I may say
"that in Charlotteville and Speyside, where yaws was so prevalent,
“a very large proportion of the population suffer from syphilis.”2

Dr. S. Branch, in his hospital report from St. Lucia for 1904-05,
speaks as strongly of the prevalence of syphilis there as I do of
St Vincent.

These, after all, are only statements of opinion, but though
figures are proverbially deceptive, 1 am able to adduce some statistics
in support of my assertions with respect to St. Vincent. We may
take these as practically representative of the state of affairs in the
West Indies generally ; for with few exceptions they all belong to
one type ; the poverty, the race of the masses, the climate, and the
geologic structure are the same.

It was agreed by the Medical Officers of St. Vincent that all cases
of tubercle and syphilis were to be classed as such in the monthly
returns of cases treated, and not scattered under the different organic
systems as local diseases. This has been done since July, 1905, and
I can show some reliable figures for twelve months’ district work in
the Colony. The population of St. Vincent is estimated at 45,000.
The hospital has a daily average of 55 to 60 patients. In the four
years I have been in charge of the hospital there have been 3,269
admissions for all causes. This number is rather swollen by the
cases of ankylostomiasis, many of which are admitted twice or several
times for two days in order to take thymol. In these four years
diere have been 630 admissions for syphilis, of which 42 were for
pnmary. Only about 20 of these cases were suffering from yaws, so
that 1 have not materially swelled the total by including this condition
"'th syphilis. The syphilis cases therefore form about 19 per cent.

the total admissions. When it is remembered that one naturally
avoids filling one’s hospital beds with chronic cases, such as those

'■ ^ospital Report, St. Vincent. Colonial Reprints, No. 20.

2 • Scholls’ Report, p. 166.

3/6

suffering from tertiary ulceration, and that ankylostomiasis has run
the total to an abnormally high figure, it must be admitted that this
percentage shows well the prevalence of syphilis.

F or some time I conducted an out-patient department at the
hospital, as a single-handed effort to deal with some of the tertian
cases of the island. Between July, 1903, and March, 1905, that
is, in twenty-one months I treated 600 cases of syphilis.

In the four years of hospital work referred to, a large number oi
amputations have been necessitated by syphilis. Of 51 amputations
of the leg at the seat of election, 27 were performed for syphilis and
1 1 for elephantiasis. And even of the latter several were brought to
the knife by extensive syphilitic ulceration.

In the reports of the Medical Officers for the district work for
twelve months July, 1903, to June, 1906 -no less than 1,996 cases
of syphilis are returned. This for a population of 45,000 is equal to
4'4 Per cent, per annum of the inhabitants. Of these 1 attended
myself 539 cases, which form about 21 per cent, of all the district
cases treated by me in that time. It must be understood that these
figures referring to syphilis have not been swelled by the inclusion
of the cases of yaws. I hough it is true that only one Medical
Officer in St. Vincent now returns yaws as such, yet this condition is
not often brought to our notice, and the few cases of frambesial
eruption which we have treated would not affect the general truth of
the figures. The one Medical Officer referred to whose district is
that most infected with yaws, only returned 24 cases in the twelve
months.

There is no compulsory medical certification of death in
St. Vincent, except for infants under one year, but nearly all the
deaths are certified by medical men. I went through the register for
two years March, 1904, to March, 1906 — to pick out and classify

the causes of death. From the vital statistics I may quote the
following : —

Total deaths . IiiI8

Still Births . ,,1

Syphilis . . . . . . ’ ’ ’ ’ ’ ’ ’ ’ ’ 56

Hereditary Syphilis (infants) . 149

Syphilis, therefore, causes on the average about 14 per cent, of the
deaths in St. Vincent.

377

A high infant mortality (25 per cent, of the total deaths in
St Vincent) and a large number of still-births are observed all over
the West Indies. These alone are evidence of the prevalence of
syphilis, though there are ofher causes — illegitimacy, ignorance and
poverty— which contribute to raise these rates.

Hereditary syphilis in infants is a matter which presses hardly on
Government Medical Officers who are required to attend the
children of labourers free, as they do in many West Indian Islands.
In the twelve months’ period referred to above, I saw no infants
suffering from inherited disease out of a population of about 5,000.
In the whole Colony in the same period there were returned 512
cases of hereditary syphilis. This is in a higher ratio to the total
cases of syphilis than in my practice. I attribute this difference to
my own tendency to class syphilis of children and young persons as
acquired, only counting syphilis in infants as certainly or probably
inherited. Comparing the ratios otherwise, I find a striking agree¬
ment in the figures relating to syphilis returned by myself and the
other Medical Officers.

To recapitulate the proofs of the prevalence of syphilis in
St. Vincent, we note

i- 19 per cent, of the admissions to hospital are for syphilis.

2. 44 per cent, of the inhabitants are attended yearly for syphilis
by the district Medical Officers.

3- In a small hospital in a small colony no less than 27 legs were

removed on account of syphilis in four years.

4- The cases of syphilis form 21 per cent, of the district cases seen

by one Medical Officer.

5- Syphilis is responsible for 14 per cent, of the deaths in the

Colony.

I take St. Vincent as a representative Colony in which the disease
ls well developed, though probably not much more so than in some
other West Indian Colonies.

h now we examine the stages of syphilis seen by the medical
ITlen in St. Vincent, we shall find a preponderance of the tertiary and
uhat seems to be an inadequate proportion of the primary stage.
The figures for the same twelve months’ period are as follows : —

Primary ..
Secondary
Tertiary ..
Hereditary

35

373

1,076

512

37‘s

I may admit that the primaries include several (say four) non
venereal cases returned by myself.

It is true that the primary lesion, even when venereal, is
frequently a very trivial-looking thing to the patient, but many such
sores are aggravated by the dirty habits and incontinence of the
black patients, and must come to hospital for surgical treatment.
In St. Kitts we operated very often on such cases. There are.
therefore, probably not many chancres of the genitals, in St Vincent
at any rate, which are not brought to the surgeon’s notice.

This paucity of venereal chancres in the tropics is referred to by
Numa Rat, whose experience extended to the W est Coast of Afnca
as well as the West Indies. *' And I may add that it is very rarely
" indeed that I have seen the initial lesion of syphilis in a black man
“ in any country. Either the syphilitic chancre in the negro is a much
" less serious affair than it is in a European, or the former considers
" it too trifling, whatever its severity, to trouble the doctor about. 1:
“ is reasonable to assume that in a place like St. Kitts, in which
syphilis prevails so generally, the initial lesion of the affection would
be modified by the presence of an inherited syphilitic taint. 1

It may justly be contended that 31 venereal primaries in 1 -99 1
cases of syphilis is an inadequate proportion. And, again, in four
years only 42 cases of primary have been admitted to hospital,
seems certain that the venereal chancres do not account for the
amount of syphilis in the Colony, and it will be my endeavour to
suggest an explanation in the following pages, which I believe applies
equally to all tropical countries.

TERTIARY SYPHILIS IN EARLY LIFE

While we were speaking of tertiary syphilis, the reader no doubt
has had in his mind the syphilis of adults. But we have now to
consider another phase of the question. A great deal, if not most, d
t ic tertiary in the West Indies is seen or has begun early in the L

of the patients, and much of the juvenile syphilis is of a very severe
character.

. T^e ^arf?e infant mortality indicates what has become of most^
'• Report on Anguilla. Colonial Reprints, No. 19.

379

the inherited syphilis, and it is well known that nowhere is the
occurrence of inherited taint in proportion to the number of infected
parents. If this were not so the human race would have disappeared
at an early period of its existence. I have rarely seen a black or
coloured child over three years of age whose syphilis was undoubtedly
inherited. Eye lesions, sabre shins and cranial malformations occur,
but by no means commonly among the juvenile cases of syphilis.
Hutdiinson’s teeth I have never seen in a dark-skinned person, and

doubt if this mark affects the negro race.

Moreover, the manifestations of disease in young persons are
usually of a severity that one does not associate with inherited taint.
Under the unfavourable conditions of child life among the black
peasants, the bad cases of inherited disease die early. The milder
cases we should not expect to run to such virulent tertiaries as we
commonly see. Those writers who try to prove that some of these
lesions, such as necrosis of the palate, are not caused by syphilis urge
the youth of many of the patients, and thus they indirectly admit the
improbability of their being due to inherited taint. I can therefore
produce these writers in evidence. J. Numa Rat is perhaps the most
careful and accurate observer of yaws. He does not believe that this
condition has any relation to syphilis, but finding a prevalence of
lesions apparently identical with those of tertiary syphilis, he did not
attribute them to inheritance, but to yaws itself.

The very common lupoid ulceration is rarely ascribed to syphilis
in the West Indies for the same reason of the youth of the patients ;
it is called lupus. Yet it invariably gets better under iodide, and often
spontaneously. Just as common in St. Vincent is ulceration and
necrosis in the throat and nose, affecting young people more frequently
than adults. In a recent correspondence in the Journal of Tropical
Medicine, James Leys1 and Numa Rat2 discuss this condition,
which the former found as prevalent at Guam in the Philippines as it
is in the West Indies. It is said to be very common also in Fiji.
Leys suggests the name “ rhinopharyngitis mutilans for the
condition, and both the above writers agree that it cannot be syphilis
because the patients are often young, fourteen to nineteen years. As
a matter of fact, sexual life begins so early with the dark races^ that

i Jour. Trop. Med. February 15th, iqofi.
2- Jour. Trop. Med. May 1st, 1906.

380

a patient of sixteen years could easib have had time to reach this
stage of tertiary from a venereal infection And in my own experience
in St. Vincent most of the few females I have seen with venereal
primary were about or under this age lint I have seen rhinopharyn¬
gitis in patients as young as nine or ten years. Both these writers
ignore the possibility of non-venereal infection in childhood, and
neither even raise the question of inheritance.

The rhinopharyngitis as described by Leys and accepted by Rat
is identical with that we have in St. Vincent. It is such a
characteristic tertiary that it would he diagnosed as such without a
moment’s hesitation if seen in a single individual. But it is very
common in some places, e g. certain parts of the West Indies, Guam,
Fiji, while in other places where there i just as much syphilis it is
only occasionally seen. It is this peculiarity of distribution which
makes the difficulty. I have pointed out that the damp, ho.,
wooded ravines of volcanic islands in the tropics are infested alike by
yaws and rhinopharyngitis. The same climatic conditions determine
the occurrence or favour the development of both. I he point to wli.ch
attention has not been sufficiently directed is that syphilis is modifie
by circumstances of climate, race, personal habits and constitution, o ,
so as to manifest itself in various forms. In St. Vincent, for exampe.
lupoid destruction of the face is the rule on the Windward side o ^
island, and necrosis of the palate and turbinated bones on the Lee ^
It is incorrect to judge of syphilis of dark races in a tropica ^
house by the standard of syphilis in Europe. So certain
rhinopharyngitis mutilans is a tertiary that its presence in ^
he conclusive that the supposed freedom of that group fronl
was a delusion, if such proof was ever seriously required. ^

The condition is this ; there is ulceration of tonsils, fauces,
palate or nose, progressing to destruction and accompanie ^ ^
by necrosis of the hard palate, the turbinated bones and
or all of these parts may he involved. The nose at len?t " 'onf
in, but the ulceration does not usually reach the skin of the ^ ^
may seethe buccal, nasal, and pharyngeal cavities tkr0''n j jjned
space, lying between the tongue and the base of the skull, ^ of
by a vast green-gray ulcer relieved bv blackened re £or the
nea^g^ne. The condition does not appear to

»• Jour. Trop. Med. MaT 15th, 1906.

38i

patients continue slowly losing tissue for years when untreated, and
it may heal even after most extensive destruction has taken place.
There is just as frequent syphilitic ulceration, but of the skin, in other
places where the rhinopharyngitis is not so common.

Hutchinson describes the same lesion as being not infrequent in
the young, and attributes it to inherited taint. It is remarkable,
however, that he does not usually find the other signs of heredity,
notched teeth and keratitis, in such cases. In view of the facts which
1 shall later on present, I have taken the liberty of disagreeing with
the master’s opinion, and believe that this severe tertiary is usually
the result of acquired syphilis.

According to Hutchinson, the tertiaries of inherited syphilis
commonly seen are interstitial keratitis at about the age of pu ) <
deafness ; periosteal nodes between the ages of, say, eight to ten y
sometimes ending in necrosis ; lupoid ulceration and ulceration o
pharynx and palate. I have only seen three instances of deafness m
juvenile syphilitics in St. Vincent. Interstitial keratitis is rare ,
only remember one case. Perhaps the negro race is not pron
though iritis and other eye affections are sometimes seen in acq
syphilis of blacks. Periostitis and synovitis in young persons
children are common enough, but these belong as muc to
acquired disease. Lupoid ulcerations and ulceration of the nOSf,^e
throat are, as we said before, very common in young persons. ese

also occur in acquired syphilis. ,

But it is not, I believe, recognised that cutaneous gumma^ ^
subcutaneous gumma leading to gummatous ulceis occur or a
common in the inherited disease. So frequent is the former
West Indies that it is well known to the natives under t e
“ blind-boil " and “ blue-blister.” It is just as common in chi ren as
in adults.

The terrible and long-lasting ulcers often met with in the young
peasants are sometimes superficial and lupoid or serpigmou , ^
more commonly they are deep subcutaneous gummata. One may ^
the early stage when there is a fluctuating tumour so like y o
mistaken for abscess. I have myself cut into such a gumma
"f eight years, under the impression that it was a cold a etion

found the characteristic soft pale tissue exuding a gummy ^

lll,,n the cut surface. The frequency of these tertiaries

Y

382

conclusive of the occurrence of youthful acquired syphilis, for if even
possibly hereditary, they can only be rarely so.

On the whole, therefore, while there is a great deal of congenital
syphilis of infants, yet I am not satisfied that the amount of youthful
tertiary which we see can be accounted for by inheritance. As before
remarked, most of the congenital disease disappears by the death of
the infants. \\ hat is left is presumably of a milder type, and would
not produce such virulent and lasting tertiaries. When it can be
shown, though this is anticipating my argument, that innocent
syphilis is of frequent occurrence (apart from yaws), I feel I am

justified in saying that the juvenile tertiary as seen is not the result
of inherited taint.

I have for some time been in the habit of jotting down short notes
of syphilitic cases, though this has not been kept up regularly, and
there are long and frequent hiatuses in my record. Yet out of the
notes, such as they are, I can place before my readers enough cases
to illustrate the several points to which I wish to call attention.

I think it will be agreed that the cases following here read much
n°re examples of acquired than hereditary syphilis in spite of the
youthfulness of the patients. I have been limited by selecting cases
in which there was no history of yaws.

l.

T ' years> Jilack. ino note ot earlier history,

lower enri°nf’ «■ 1 1^I1’eilted scars °n both legs. Deep adherent scar over th

At present this • & lbu a‘ lias had an ulcer of the right leg for some yean
of the let. wha-t*-311 en°nnous sloughing ulcer involving half the circumferenc
cicatrized V ! 1San‘UCh enlarSed- The of the circumference of the leg i
years standing US “P?*?00- This is clearly a case of tertiary of man
the primary she ” aCqui/ed dlsease- Yet at the time when she must have ha
primary she was rather too young for venereal infection.

Cask 2.— VV. S.; 13 years; Hindu.

and adherent 1°"? T u°‘h kneeS‘ °n the ri8ht ankle an '"u

volcanic eruption! \ 1 ^ ^ thlS rnore than four years ago (dated by th

This has been of ♦ A arg® f°U‘ tertiary ulcer covers the back of the right fooi
nine years " ° ^ duratio"- The tertiary here dates from the age c

« ask 3.-D. s . Io years; Riack

superficial ukerTon the lefTw*1 °f “ blind'boils ” °n both le&s; S*™

end of left fibula r 1 Ieg‘ Swe,bng of left ankle, and periostitis of lo

H.rd g,a„ds on both g'0i"S' lh°8e ^

circular. Thev arp r™ Sma11 ind°lent ulcers about 6 to 10 mm. wide, a
begin as a pimple whic^hr0^ “ultipIe and symmetrical in arrangement. T1
eaks down and suppurates, leaving a shallow punch

383

ulcer, which is very resistant to treatment. They seem to belong to the “ inter¬
mediate” stage of Hutchinson. This case is one of early tertiary in acquired
disease, at an age which precludes the possibility of venereal infection.

Case 4.— A. M.; 12 years; Black. Denies yaws.

July 2nd, 1903. Left tibia sabre-shaped. Right with a very large node.
Large scar on back of right thigh. Large ulcer inside the upper lip. Columna
nasi has disappeared, and a large ulcer covers the surface of the nasal cavity,
eroding the alae nasi. Several scars of previous skin gummata on the inner
side of nates. Here it may be admitted there is a probability of heredity indicated
by the sabre shin.

Case 5.— W. C.; 18 years; Black. Has been five years in the Pauper Asylum
as a hopelessly disabled person.

October nth, 1902. Left arm has been amputated below the elbow. F.lbow
scarred and ulcerated. Left leg is a scar from hip to toe, with the knee and
ankle contracted in web-like keloids. Right leg is scarred below the knee. Both
feet are masses of ulcers and bone disease.

After more than two years’ treatment with iodide, and amputation of the left
thigh, he was completely healed and left the Asylum. He has remained well to
the present. I think that hereditary syphilis is not known to produce so frightful
a condition.

Case 6. — J. L.; 22 years; Black. Sent to the Pauper Asylum at the age of
>6 years.

October 17th, 1902. Both knees and elbows stiff and contracted; extensive
scarring and lupoid ulceration on both shins. Multiple nodes on forearms and
humeri which give them a distorted appearance. Hard palate perforated. Deaf.

This case had since healed, but lately developed intense headaches and very
marked mental dulness. There is no question of venereal infection in one who
■s already hopelessly disabled at 16 years.

W ^ASE 7'~J- ^*5 22 years; Coloured. Was disfigured at the age of 14 years,
as sent to the Pauper Asylum at 17 years.

u °C.!ober x7tb» !902- Nose is gone, lower lids retracted by scarring. Side of
PPer lips drawn upwards. Face scarred generally. Left leg extensively scarred
h gummatous ulcers on the lower half of the shin. Ulceration of throat. Since
the C- U1K*er P°tassium iodide and sent out. Here again the early age at which

Pa lent was considered incurable precludes the venereal infection.

Case: 8.-W. s., 9 y8ars; Black.

23rt^’ I902- Large keloid on right elbow and a scar on the forearm.
and° v C Uld SCar on le^1 knee. Scar on outer malleolus. Large gummatous ulcer
8ummafaS -vp ankle. The scarring here indicates deep or subcutaneous

a w ich had been appearing during several years.

Case 9 R. §.; j4 years; Hindu.

arm 7tE’ I9°2- Extensive scar all round the left elbow and half of _tbe

^gers ;rnS1Ve scar on ^e back of the hand, and a scar on the wrist. Two
scars at th racted and a few small ulcerated spots on the fingers. ExtensI^e
Large scar 6 baCk °f the riSht elbow> hack of lower forearm, and back of han .
lhe lower “-TV116 front of left knee, and another of the calf. Scarred all over
°ne large a the tibia- Foot a distorted mass of scars and bone disease, «i

tibia. \ aand several small ulcers. Scars on right knee and on lower end ot
Septum naif6 UlCCr °f ankle- Seven superficial scars on cheek, nose and ip-
' 8°ne- Was treated in hospital with a view to amputation of the

3*4

left leg, but when considered fit he declined operation and was discharged. He
has not been heard of since. This is a class of case that is never seen in
Europe. I "have notes of several as severe cases in St. Vincent.

Case io. — L. B.; 9 years; Black.

November 21st, 1902. Perforating ulcer at junction of hard and soft palate.

A very young case of “ rhinopharyngitis."

Case it. — C. A.; 16 years; Black. Says, “never had yaws.”

March 1st, 1906. Scar of veluin palati with a perforation and a cleft of the
palate. Scar on the face, one on the arm, and several on the legs, of tertian-
ulcers. Two of these involved loss of bone from the tibia. Gummatous ulcer
of one foot and the other leg.

Case t2. — A. S-; 12 years; Black.

1-ebruary nth, 1903. Fistula of lachrymal duct from disease of maxillary
bone. Left ala nasi deformed and nostril contracted by a scar. Cartilage of
septum has gone. There is an ulcer of soft palate and fauces. Uvula has
disappeared. Another young case of “ rhinopharyngitis.”

Case 13. — E. W.; 12 years; Black. Had disease of the nose two years ago.

November 22nd, 1902. Ulcer and necrosis of palate. Ulcers of right thigh,
knee and skin. Ulcer and bone disease of first metatarsal. Ulcers of left
thigh, knee and shin. Both knees contracted and legs wasted. Ulcers of right
arm and hand. Extensive scars on left forearm ; large periosteal node on fifth
metacarpal. Some wasting of both arms.

This case is known to have had yaws when two years old.
Cases of tertiary with history of previous yaws may be multiplied
indefinitely.

Among 17 children under the age of 15 years suffering from
tertiary syphilis, yaws was admitted in the case of 12. Of the others
only two had marks of inherited taint. I regret that I cannot quote a
greater number, but one has been so in the habit of expecting the
history of syphilis, that enquiry was not often made on this point, and

I can only find a reply to this question in the notes of seventeen
children.

NON-VENEREAL SYPHILIS

How often do we hear it inferred that a condition cannot be
syphilitic because the patient is young and there is no evidence of
eritance. And how often is there a reluctance to attribute syphilis
to an adult, from a feeling that this is an unjustifiable slander of his

racter. Yet thoughtful writers have warned us against this
attitude.

The physician must forget the local primary lesion, and
„ put aside the idea that the diagnosis of syphilis carries with
11 any stl£ma of impurity.” (Fagge, by Pye Smith.)

385

I cannot state the truth about the West Indian negro better than
by quoting my own words from a report already referred to.

“ This applies with far greater cogency in the West Indies, where
"the presence of a primary lesion on the genitals of an unmarried
“peasant girl implies no immorality. Among the people there is, in
“ a true sense, no sexual immorality. Coitus among children before
“ puberty is as common as kissing among European children. The
"normal sexual life of the adult is one of transient concubinage,
“ which does not restrict casual intercourse any more than does the
“ occasional marriage. The mother of a large family of unknown or
“various paternity finds it very tiresome in the daughter of sixteen
“years to add another infant to her burden. But that is all. She
“ accepts the grandchild with simple faith, and tells you that God
“ sent it.”

Four years ago I was asked by the police to see a case suspected
to be suffering from smallpox (or Trinidad varioloid varicella). I
found the patient, a young woman of decent coloured class, to be a
case I had already been treating for syphilis within the last few
months.

But the interesting feature was that two little girls, aged about
seven and eight years respectively, in the house had now an eruption
°f papular vesicular syphilides. Not long ago I attended the
daughter, aged nine years, of a coloured gentleman. She had been
ailing and getting thin for some months, had had frequent and some¬
times severe rheumatoid pains, and a persistent rash of papules. She
had been under treatment by another physician without improvement.
' offered no diagnosis, but prescribed a hundred powders of grey
powder and saccharated carbonate of iron. The rash rapidly cleared
UP> and the general health became robust many weeks before I
permitted the discontinuance of the powders. These were instances
of non-venereal infection among clean, well-cared for children.

About the same time that the first case above was seen, I was in
consultation on a black peasant girl of sixteen years, suffering from a
Profuse pustular rash, which was thought likely to give rise to a
smallpox scare. My colleagues both diagnosed yaws. I said syp
and from my point of view we agreed. But one of the others, a man
Wlth thlrty years’ experience of the West Indies, objected-- ow
COuld she get syphilis.” I have been treating that girl off and on o

386

the present time for profuse eruptions, iritis, numerous suppurating
glands in the neck, bone aches, periostitis. In fact, she has had
nearly every early syphilitic symptom possible except nerve syphilis
But she has never had either a venereal chancre or a frambesial
eruption.

It is most important that the profession, especially in the tropics,
should realise the importance and frequency of extra -genital syphilis.
We all know that surgeons and nurses get finger syphilis occasionally.
In the island of St. Kitts, with at no time more than eight medical
men in practice, and usually only six, there were in twenty-seven
years five instances of finger syphilis in surgeons. These are well-
authenticated cases.

When we consider how readily syphilis is inoculated, and how the
habits of the poor in the tropics afford opportunities for inoculation,
we shall not wonder that extra-genital primary occurs. In the West
Indies the labourers, especially children, go barefoot, and are,

therefore, frequently receiving bruises and scratches on the feet and
legs.

The negroes never realise the fact of infection or contagion. This

forces itself on our attention by the trouble they give when quarantine

measures are to be carried out. They are too indolent or careless to

make any effort to avoid or prevent infection, even when they

acknowledge that a disease is catching. A child with yaws huddles

ogether with the other children just as naturally as the adults will

ep or co abit with lepers. In I ortola, while I was there, the only

i a 6 ^ ' ° cou^ be called a professional prostitute was an ulcerated

eper. he syphilitic, the yawsy, the leprous and the healthy mix in

pip h aif- .mt“nate contact without a thought. Leprosy, like

no^a^ iaS1f may be the Subject °f reproach, but these diseases are

dislrn aUSe aV°1anCe °f Physical COntact Yaws is hardly even a

of a hen eTP ? C CyeS °f schoolmasters and other coloured folk
a better class than the labourers.

can bendevt CirCUmStances- * » bardly wonderful that yaws

muaculousTr 7 “ * "ative community; and it would be

almost ^universal D 'VCnereal Action did not occur where syphilis is

Jh° 566 the Pe°Ple’ n0t ^

’ my description of West Indian

387

peasant life not unfamiliar, and if a look-out is kept, I am sure others
will verify the existence of non-venereal syphilis for themselves. In
places where yaws, under its various names, is prevalent, it will be
found that frambesial eruptions and non-venereal syphilis of ordinary
type are running together. In the drier and less-wooded localities
the latter will probably be found the more common form.

It must be remembered that in inherited syphilis secondaries do
not occur after the second year, and are usually over in the first year
of life. Tertiaries are the only lesions by which inherited taint can be
manifested in children of, say, three years and upwards. Jonathan
Hutchinson lays down : —

“ If secondary symptoms of the kind described are to occur at all,
“they will show themselves in infancy, and in the vast majority of
“cases within the first three months of life. This is a very important
"fact. If a syphilitic infant survive the first outbreak, in the course
“of from six months to a year the symptoms common to this stage
"(the rash, snuffles, mucous patches, &c.) will wholly disappear, and
" there will follow a period of some years during which no active
‘ symptoms will occur.’’1

When, therefore, undoubted secondary eruptions and swollen
glands are seen in children over the age of two years, they are almost
certainly due to acquired disease, and this certainly rapidly increases
with the age. Such children should be examined for the primary,
which in the bare-footed is usually on the foot, most commonly on the
outer malleolus. The indurated pigmented scar may be seen some
time after the sore has healed, but these sores are often of long
duration, and may be still unhealed months after they were infected
with syphilis. They are often already indolent ulcers before the
infection. I have diagnosed such primaries before the general
symptoms were manifest, and the later eruptions have justified my
diagnosis. More usually, however, one passes the sore by until the
patient is complaining of rheumatoid pains or fever, for sore feet in
children are so common that we cannot treat them all as potentially

syphilitic.

For several years I have been impressed with the frequency of
these extra-genital chancres in children, followed by secondaries of

’■ Syphilis, Ed. 1901, p. 75.

.388

the ordinary type, and some of my colleagues have adopted or
confirmed my opinion. Such cases are seen more commonly in towns
where yaws is never prevalent. I have seen a few such develop
frambesial eruptions after having been diagnosed and treated as
syphilis ; the frambesiae appearing along with the other syphilides.

Nearly everyone familiar with yaws recognises what the natives
all know, that there is a primary in yaws. No doubt this is often a
fungating ulcer rather like a frambesia, but it may be identical with
the foot chancre of ordinary type. Chancres on the penis not
infrequently fungate in the same way, as Hutchinson points out. The
foot chancre, out of which yaws also sometimes originates, is, in my
experience, a small sore about 5 to 10 mm. wide, round, with
indurated base and periphery and raised edges. It may be on one of
the toes, the inner malleolus, the tubercle of the fifth metatarsal or
back of the foot, but the commonest site is the outer malleolus.

In the case of yaws, the natives in some instances do not recognise
the primary, which may be inconspicuous, as is often the case in
syphilis ; and then a large group of frambesiae, especially a horse¬
shoe group, may be called the “ mother-yaw." On the other hand, a
primary may really become the “ master-yaw ” or largest frambesia,

by fungating and having a satellite eruption of papillomata
around it.

It is perhaps unnecessary to insist that there is aways a primary in
yaws. Numa Rat has made this quite clear, and the universal opinion
of the inhabitants of yaws districts themselves leaves no doubt on the
su ject. Notwithstanding, it is urged (e.g. Manson, Tropical Diseases,

F>- 530) as an argument against the syphilis theory that the primary is
wanting in yaws.”

The identity of non-venereal syphilis of the easily-recognised
-r* aS ^ ^rs*- referred to, and the whole group of symptoms
, ed '^WS’ is 80 obvious that the peasants themselves bear witness
, ** *n<dude under this name other skin eruptions, besides

e rambesiae, which they have observed to be associated with the

CSe bave popular names — tubbo, corn-yaws, crab-yaws,
pian-gratelle and dartres. The peasants speak familiarly of “yaws-
pains. hey point to where a patient did “ ketch a yaws,” who has

svnVkT?V 0Pf frambesiae’ but h*d rheumatoid pains and vesicular
syphilides, and is afterwards seen with tertiary

389

It is well to consider some of these other yaws eruptions, which
are so well recognised by the natives and described by writers.

Tubbo is a papilloma, pathologically the same as the frambesia,
though unlike it in appearance, which breaks through the thickened
skin of the sole, and is modified by the pressure of the hard epidermis,
in the same way as the small papules of early psoriasis are modified
when they appear under the skin of the palm.

Corn yaws in English-speaking colonies covers all discrete
papular, vesicular, and even pustulo-vesicular eruptions.

Crab yaws is simply palmar and plantar psoriasis. It is sometimes
severe, and becomes a moist dermatitis. More usually it is seen some
time after the yaws, and is identical with the tertiary plantar
psoriasis. It is chronic, and very resistant to treatment.

Pian gratelle, as described by Numa Rat, Alford Nicholls and
others, is obviously the small papula of syphilis-lichen. It appears at
an early stage in a profuse crop usually, and, as in European syphilis,
it may reappear as a late secondary, and its presence indicates a severe
infection. This tendency to recurrence and this significance of the
papular eruption is insisted on by writers on both syphilis and yaws
(cf. Jamieson, Diseases of the Skin, p. 544, and Alford Nicholls, Report,
p. 285).

Dartre (or as Nicholls calls it, the macula or squama) may be
perhaps in some instances the macular rash, which is the earliest skin
symptom of syphilis. The colour of it cannot be seen on a black
skin, but when severe and on unwashed skin it may possibly result in
an exfoliation of the corneous layer of the epidermis ; but as
described it is the syphilitic scaly psoriasis. This is the earliest
eruption in yaws, and is very common. It is frequently coincident
w'th the papular eruption — lichen — when that is present, and some¬
times lasts to the stage of the frambesial eruption. Its appearance is
absolutely characteristic of scaly psoriasis.

AH these obviously syphilitic conditions are recognised by the
natives to be essentially parts of yaws as a disease. A patient is
said t0 have yaws on the strength of any of these, even by medical
men’ wkhout the frambesial eruption.

Alford Nicholls and Numa Rat, who have made long and careful
°bservations of yaws in its earlier as well as later post-frambesial
' abes, describe these conditions as belonging to yaws. Dartre and

390

pian gratelle especially are noted by them as being characteristic of
yaws. In my own experience, psoriasis and lichen are just as frequent
in occurrence in cases which develop frambesiae as in those which do
not. The rheumatoid pains are almost always a feature of yaws, and
these, as well as the less frequent palmar and plantar psoriasis and
vesicular syphilides, are identical in both diseases.

In short, the vox -pofuli has declared the truth, which the medical
profession has not yet generally accepted, that yaws is non-venereal
(or sometimes venereal) syphilis of the tropics. At any rate, the
people name syphilitic manifestations and associate them with the
frambesial eruption under the one name yaws.

A few recent writers on yaws, under its various names, have seen

and recorded these facts ; proving that yaws is more than the eruption

of frambesiae, as described in text -books (e.g. Scheube). Even those

writers who have not included these symptoms as part of the same

disease have noted the frequency of the occurrence of syphilitic

manifestations together with frambesiae. They regard the smaller

syphilides as evidence of syphilis and reserve the frambesial eruption to

be called yaws, and then speak of the concurrence of the two diseases.

Daniels and Wallbridge note that “ a fair number of such cases are

recorded, and in all the syphilis preceded the yaws.”1 It is doubtful

whether by ‘ syphilis ” they refer to primary syphilis, i.e. the genital

chancre from their point of view, or to small syphilides. Blanc, of

Tobago, observes : — “ I have not seen any yaws patients with syphilis

in the primary or early secondary stages, but have very commonly

<( seen tbem in the early stages of yaws with well-marked papular

“ eruPtlons- which, however, might have been either syphilitic or the

u Precursors of yaws tubercules. These symptoms commonly

‘‘disappear under treatment, but sometimes give rise to a scaly

“ eruptlon very hke psoriasis. . . . Syphilis, as evinced by severe

„ cachexia> or indurated and enlarged glands, enlargements of bones,

„ n°deS’ &c* has been a very frequent complication in the cases of
yaws. 2

There can be no doubt that the other secondary manifestations of
yaws are identical with symptoms elsewhere ascribed to syphilis
ome writers attribute them to yaws, others to a concurrent syphilis

•• J. Hutchinson, Fasciculus, p. i9.

2. Nicholls’ Report, p. 165.

39i

But anyone who cares to look for them will not have far to seek in
almost any case of yaws.

Consistently with the description of the secondaries of yaws, Rat
goes further and describes the tertiaries. In this he is in accord with
Daniels and Corney in Fiji. These symptoms need no description —
periostitic nodes, multiple dactylitis, destructive rhinopharyngitis,
lupoid ulcerations, &c. It is only to be remarked that one does not
realise to what extent tertiary syphilis can go until he has seen the
cases of untreated syphilis in a yaws district.

It is evident that, in the experience of the writers just mentioned,
there is not enough venereal syphilis to account for the extraordinary
prevalence of tertiary, and so they were driven to ascribing the
tertiary, quite correctly, to yaws. They have failed, no doubt, to see
the non-venereal syphilis without frambesiae, which would have
afforded an escape from the difficulty.

Tertiaries identical with those of syphilis are recorded as being
exceedingly prevalent, and are attributed to yaws by Daniels and
others in Fiji, Kynsey in Ceylon, Rat in Dominica. Among ninety
cases of tertiary syphilis of whom the history was ascertained in
St. Vincent, sixty-nine were said to have had yaws and only five
admitted genital syphilis. Some of the remainder had suffered, no
doubt, from extra-genital chancre without frambesiae, and therefore
did not recognise that they had yaws. On the other hand, not all
that claim to have had yaws have had a frambesial eruption, for, as
I pointed out before, the natives, and apparently also some medical
®en> call cases yaws that have had no frambesiae.

From the facts of the polymorphism of the eruptions and the
occurrence of tertiaries, it is evident that the only difficulty in the way
of the general abandonment of the yaws heresy is the frambesial
emption itself. As Jonathan Hutchinson has pointed out, the
Profession did not originally hold yaws to be an independent disease,
*0 that it is quite fair to speak of this belief that yaws is not syphilis as
a heresy. The frambesial eruption is of such importance and,
t0 my mind, such a formidable difficulty that I have reserved its
consideration for another section.

% contention is that syphilis is very common, in fact, almost
universal, among the natives in the tropics ; that in certain districts
:t usually presents an eruption of papillomata, which has given rise to

392

the idea of yaws as a distinct disease ; but that the cases with
papillomata present otherwise all the features of syphilis, just as do
the non-venereal cases. To appreciate the relation of the various
forms of eruptions to each other and to recognise their nature, as well
as to understand many other disease problems, it is necessary to
remember that syphilis is in the tropics not usually a venereal disease.
Notwithstanding the vitality of long-rooted error. I feel sure that
the profession would have more readily accepted the dictum of
Jonathan Hutchinson had we realised the frequency of extra-genital
chancre. It is the teaching of the schools and text-books that syphilis
is a venereal disease, which has blinded us to the fact that in the
tropics it has little to do with sexual intercourse ; that there is, in fact,
far more non-venereal syphilis in the world than syphilis acquired on
the genitals.

Case 14. — D. k..; 6 years; Coloured.

March 27th, 1906. Has had a sore of the toe and paronychia for several
weeks. About two weeks ago began to get “ spots ” on the skin. Has now a
few spots of psoriasis over the body and palms of the hands. Also on the knees
and elbows. Is feverish at nights, has small hard glands in both groins and on
elbows.

Here is a very early case in which, I think, the diagnosis of syphilis
is justified.

Case 15. — J. W.; 13 years; Black.

August 1 st, 1903. Has a small indolent dirty sore on the left outer malleolus,
with raised edges and indurated base and periphery. This has been present for
several weeks. There are tender glands in both groins. That on right larger
and more tender. Small hard glands left side of neck, on both elbows and in both
axillae. 1 wo weeks ago the first crop of eruptions appeared and they have
continued to come out since. Has now a general papular eruption (licheni
tending to vesicular heads. This is very closely set over the whole trunk,
extending to the neck, upper arms, and thighs, where it thins away. A few
papules on the lower part of the cheeks, none on the limbs.

Also an early case, in which the diagnosis seems very evident.

( ask 16. — K. B.j 10 years; Black.

April 6th, 1903. Three months ago had a scratch on right outer malleolus
W 10 ecame a sore. Her mother shows this as what she considered a mother
vaw It is now a pigmented and indurated scar. There are many spots o>
scaiy psoriasis over the body, face and limbs— rheumatoid pains in the right
elbow, both knees and ankles. The mother complains that the child is getting
azy and hard to rouse in the morning. Later she developed evening fever.

September 1st, 1903. Noted that she had attended irregularly. Eruptions
are worse. Has never had frambesiae.

This illustrates the popular identification of syphilis with yaws

393

Case 17— M. Nt.; 14 years; Coloured.

April 22nd, 1903. Has a small excavated ulcer about 6 mm. wide on a raised
indurated base, on the outer side of the right foot. This has been present fox-
several months. Hard enlarged glands in the right groin. Swollen tender glands
in the right side of the neck. A chain of small tender glands in left side of neck.
Fever, malaise.

A patch of superficial flat large papules with crusted scabs, each 10 mm. in
diameter, on the front of the sight thigh. These are like xnild rupia or ecthyma.

Case 18.— L. P.; 21 years; Black.

April 2nd, 1903. Has a deep excavated ulcer on the left outer malleolus.
Does not admit venereal primary. Has no leucorrhoea to indicate cervical
chancre. A coarse papular desquamating eruption of lichen on both hands and
feet. Few papules on the face. Very profuse vesicular eruption on both legs
and thighs. Glands enlarged in both groins and on both elbows.

In spite of the age of the patient, I had no hesitation in attributing
her syphilis to infection on the ankle.

Case 19. — S. X.; 30 years; Black.

March 15th, 1903. Has four small ulcers with indurated bases and raised,
thickened, everted and irregular edges, round the right nipple. These appeared
first three months ago while she was suckling an infant with yaws. They have
all the appearance of primary chancres.

On 6th May, 1903. Complains of pain all over the body, malaise, and
constipation. Looks and says she is very ill. Has nervous sensations and
sleeplessness.

She never developed any eruption. Continued ailing and sickly for at least
two years under treatment for which she attended irregularly, as she felt better or
worse. Treated with small doses of mercury from the first. Never salivated.
At length she complained mostly of gastric troubles. These were not due to
mercury, for she only took doses of 10 minims of liq. hvdr. perch, three times a
day. After two years recovered health.

This was a case of visceral syphilis contracted from yaws. I have
now under treatment a very severe case of gastric syphilis, in which
fte gastritis followed a year after a genital chancre. In this, also,
there was no eruption, but adenitis and iritis.

Iase 20. — I. D.; 9 years; Black.

February 25th, 1904. A wide ulcer on the left outer malleolus, with a dirty-
yellow crust. Profuse patches of psoriasis and areas of lichen over legs, trunk
and arms.

I his would be called an early case of yaws, with the maman-pian,
dartre and pian gratelle. By changing these names it becomes

syphilis.

Case 21. — G. P.; g years; Black.

'9°6. Small punched ulcer about 3 cm. wide on the back of the foot near * e
lues' with raised thickened edges. This has been present for several months.

There are hard glands in both groins, large soft glands in the neck. History
x” in successive crops for several weeks, some months ago.
raes complaining of fever, malaise, and anaemia.

394

In this case the knowledge of the occurrence of non-venereal
syphilis led one to inquire about the eruption. This, together with
the glands, the primary and the cachexia, leave no doubt as to the
diagnosis. Chicken-pox, of course, does not continue in crops for
several weeks.

Case 22. — V. T.; 4 years; Black.

Mother was treated for a copious syphilitic rash eighteen months before. The
patient was treated for an eruption some months ago. Now has a papular rash
(lichen) over neck and shoulders. There are patches of weeping and exfoliating
dermatitis over each side of forehead, on both scapulae, and on back of shoulders.

This is too old for hereditary syphilis. It is not unreasonable to
attribute the rashes to syphilis acquired from the mother.

Case 23. — R. S.; 5 years; Coloured.

I9°5- Multiple ecthymatous sores encrusted with yellow scabs on the legs.
I.arge patch of the same on the left thigh, and a flat condylomatous spot with
dry yellow crust among the ecthymata. Pigmented marks of large pustules about
the body generally. Keratitis of one eye. Treated with mercury.

December 20th, 1905. Eruptions have disappeared, leaving no scars, but
pigmented stains which subsequently faded. Admitted to Hospital for corneal
ulcer and treated till well with mercury internally.

Here, as in the last case, there was no primary to be found, but the
condition was evidently syphilitic, the age too late for secondaries of
inherited taint. The eruption was one like frambesia, and even
presented one spot of condyloma. The crusts, however, did not cover
hypertrophied papillae, but eroded spots.

< ask 24.— M. D.; 3 years; Black. Mother seen on 28th October, 1902, with
pro use eruptions of circular patches of scaly psoriasis over whole body, limbs

January 5th, 1904. T he child has a multiple eruption over both arms and

egs and back, of scaly psoriasis; patches of lichen in spots: and circinate and
exfoliating dermatitis.

The age is too late for the secondaries of inherited syphilis;
besides the mother has acquired syphilis only about fourteen months
before. If the child had inherited syphilis from the father the mother
would have been immune, by Colfcs* law. It is clearly an instance of
acquisition from the mother.

( ASE 25.— N. P.J 8 years; Black.

originated* fmm I9°3’ U*Cer’ 50 nim- wide on the dorsum of the left foot, which
surface S,C'atCh *hree mon,hs “6°. This has a raised touting

large gland in right*6 r.&1S.ef everted edges. A very large gland in the left groin,
neck Has int/ . groln’ g and in left elbow. and chains of shottv glands in the
a desouamatinn cn H V eV"’ a"d with drv hard stool! There is

ie%r °f t"6 and

both

395

This may almost be given as a case of yaws, except that the
eruption was not perfectly frambesial. The patient lived in town ;
had he lived in a country village he would, no doubt, have had yaws.

These few cases, I think, illustrate the various aspects in which
non-venereal syphilis may be presented to the physician : primary
syphilis not followed by frambesia contracted from yaws ; secondaries
in children with and without an evident primary ; secondary in an adult
due to extra-genital infection. Two show the difficulty there may be
in differentiating yaws. I selected five which w'ere seen within five
weeks at a. time when my notes were being regularly kept. This
number indicates how common such cases really are. (Cases 16, 17,
18, 19 and 25.)

ATTITUDE OF THE MEDICAL PROFESSION
TOWARDS YAWS

In consideration of the symptoms of yaws, as referred to above,
there would certainly never have been a doubt as to its identity with
syphilis had it not been for the frambesial eruption. This is so
unusual in European syphilis, and such a distinct and characteristic
manifestation in yaws, that it was natural it should overshadow the
other symptoms. Most physicians, who have not looked carefully
into the matter, have a picture of yaws as consisting only of the
frambesial eruption ; they think this is a harmless, though unpleasant,
disease, which usually runs its course and disappears spontaneously,
without permanent injury to the health. But at first yaws was
included with syphilis, and even since its separation as a disease
Sl{i generis, the doubt that it was correctly separated has existed.

In view of the enormous prevalence of yaws in the tropical world,
lhis doubt gives an importance to the disease which, I think, has not
been appreciated ; for it is a most serious matter if syphilis, the most
destructive and far-reaching in its effects of all diseases, is being
allowed to ravage whole communities untreated and almost unnoticed.

When in recent years the syphilist, who for forty years has been
the authority to the English-speaking profession, pronounces his
unqualified opinion that syphilis is yaws, it is time that the world took
UP the matter seriously. In his preface to Numa Rat’s work on yaws,
Jonathan Hutchinson expressed this view, based, it appears, largely

396

on the evidence offered by Rat himself. Since then, after further
investigation, he has delivered, so to speak, an ex cathedra opinion
to the same effect in Fasciculus XIV of the new Sydenham Sodety’s
Atlas, and presented the evidence for his opinion. Notwithstanding,
the medical profession, with few exceptions, are in the same attitude
of indifference. The new student of tropical medicine will hardly get
from his text-books any indication that there is any serious or
reasonable doubt about the independence of yaws. The 1903 edition
of Manson makes a misleading reference to Hutchinson as holding
that yaws is possibly syphilis modified." The English edition of
Scheube, 1903, only refers to Hutchinson to misunderstand him in a
similar way.

Somewhat in consequence of this, those who believe in the
identity of syphilis and yaws are in a small minority. Not many take
much account of yaws at all. Of those who believe in the
independence of yaws, it is only the few enthusiasts who have studied
the matter that think it a serious disease. It is not to be wondered
at therefore, that in most places yaws is not treated at all. Trinidad,
especially in the island of Tobago, is the only colony in which I have
heard of any rational and systematic attempt to deal with the disease,
but even there, because they do not realise its gravity and true
nature, the measures taken by the medical officers must be incomplete.
In some othei places the yawsy are in effect regarded as social
offenders, and forcibly removed to yaws hospitals for treatment. Such
a course can never obtain the sympathy and support of the people
whom it is meant to benefit.

In the treatment at all yaws asylums and dispensaries, as far as
one can judge from the literature of the subject and the reports, the
am esial eruption is regarded as the disease. When this dries, the

^ 1S. Sa^ t0 cured ; when another crop appears this is

a re apse. All physicians that have given mercury find that the
eiup ion isappears rapidly as a rule, and that it is apt to recur when
e mercury is stopped. Some think that the mercury has a bad
ect, and that the yaws should better be allowed to run its course.
Lertamly the frambesiae disappear under the mercurial treatment,

and as. certa,nly hey sometjmes ^ ^ ^ ^ ^

nor Hn h ° S^>^^ls because a crop of secondaries has faded ;
does he stop mercury on that account ; nor is he surprised if more

397

secondaries appear even while his patient is still taking mercury.
When treating a case of syphilis, one is thinking not of the
secondaries but of the tertiaries. The second stage will usually run
its course and the patient get well in time without mercury. But it
is, as a rule, only the mercury given early that saves the patient from
a life of tertiaries later on.

If the papillomata are all that there is, then yaws is hardly worth
much trouble; but the observations, especially of Numa Rat and
Daniels, have established the fact that there is an aftermath of evils.
They could not help but attribute to the effects of yaws the vast
amount of exaggerated ulceration, periostitis, necrosis, and dactylitis
which they saw around them. Numa Rat boldly calls these the
tertiaries of yaws, though he still believes yaws to be distinct from
syphilis. If the diseases are different, then yaws is the worse of the
two, and it is the most terrible scourge from which the human race
suffers.

If even yaws is not syphilis, then there is still abundant
indication that the treatment is the same, and we should insist on the
early and persevering use of mercury ; for the time for treatment of
tertiary syphilis is in the early secondary stage, and by inference the
same must be true of yaws.

If, however,, we decided to regard yaws as syphilis, its importance
would be at once recognised by every member of the profession ; and
the people, educated in time to pay proper attention to the disease, as
Europeans do to syphilis, would assist in the effort to limit its spread
and cut short its dire results.

A systematic and conscientious treatment of the cases, and a
persistent education of public opinion, would at length effect what all
the yaws asylums, dispensaries and commissions have failed even to
begin.

In view of the theory of Hutchinson that yaws is the original
disease, we may abandon the name syphilis, which bears but an
indifferent repute, and speak of yaws. Recognising’ its nature, we
may teach the prophylaxis of syphilis under its various local names.
The writer has already prepared, with this view, a primer of hygiene
for West Indian negro children, at the request of the Board of
Education of St Vincent.

i

398

I said before, that it is only the frambesial eruption which
constitutes the difficulty in accepting yaws as syphilis.

Manson says: — “I may mention the primary sore, the infection
“ of the foetus, the adenitis, the exanthem, the alopecia, the absence
“ of itching, the iritis, the affection of the permanent teeth, the bone
“ and eye affections, the congenital lesions, the polymorphism of the
“ eruptions, the nerve lesions, and the gummata of syphilis. All these
“ are wanting in yaws."1

To take these in detail : -

(i.) The primary sore. — There is, as we have seen above, no
doubt at all about this. Numa Rat made this quije clear, and the
experiments of Charlouis are convincing. In short, as Hutchinson
says, Alford Nicholls is the only observer who disputes the fact that
there is always a primary lesion in yaws. Careful examination and
enquiry will elicit this quite as often as in cases of syphilis.

(2.) The infection of the foetus. — It would be quite as logical to
deny the inheritance of syphilis. It is so rare for a child to be bom
with syphilitic eruptions that few of us have seen such a case. Many
infants are seen with yaws at the age that they might have the
secondaries of inherited disease. But when such cases occur, we are
in the habit of inferring that the yaws is acquired. Inheritance has
been demonstrated in at least two cases mentioned by Hutchinson.
I have never myself seen an infant known to have hereditary syphilis
develop a frambesial eruption ; but all the infants I have seen thus
have been at once put on mercury, and kept at it as long as I
continued to see them. They get better, or, more usually, I hear
nothing of them for a long time until the parent applies for a death
certificate. It cannot certainly be said that inherited syphilis is not
sometimes frambesial, and it is known that a child may inherit the
yawsy taint and develop the frambesia in due time.2

The adenitis. This is an almost invariable accompaniment of
t e yaws infection. The experiments of Charlouis are conclusive,
besides the experience of actual examination of yaws patients.

(4.) The alopecia. In all my experience of many hundreds of
cases of negro syphilis (non-frambesial) I have only once seen
aopecia, and in that case the diagnosis was at first obscure, but
'• Tropical Diseases, page 530.

2- Hutchinson Fasc., page 18.

399

afterwards cleared up. A physician of far longer experience than
mine of the West Indies tells me he has never seen syphilitic alopecia
in a black patient.

(5.) The absence of itching. — This is admitted to be by no means
an absolute rule in syphilis. Nor is itching at all the rule in yaws.
When we consider the irritation of the small flies around a yawsy
patient, the admission of itching in some cases is not remarkable.

(6.) The exanthem. — It is difficult to understand which particular
eruption is referred to. All or most of the syphilides have been seen
in yaws.

(7.) The iritis. — This is distinctly uncommon in negro syphilis,
and is a comparatively rare symptom anywhere. It is quite feasible
to imagine that, where syphilis is profusely manifested on the skin,
the deeper structures, particularly the nerve system and its
appendages, may escape.

(8.) The affection of the permanent teeth. — This must be
extremely rare, at any rate, among West Indian negroes. I cannot
remember ever having seen notched incisors in a black person.
Finucane and Corney, as mentioned by Hutchinson, report ha\ing
seen such occasionally in Fiji, where syphilis is not admitted to be
present. One has the impression that either the negro race is exempt
from this affection, or that it is caused by mercury given in infancy.

I very frequently look at patients’ teeth to note the tolerance to
mercury, and also to note the irregularities which constitute a stigma
of degeneration.

(9) The bone affections. — These have been described frequently
as tertiaries of yaws. But the author has already denied them to us
by saying that they are due to an independent intercurrent syphilis.

(10.) The congenital lesion. There are none in syphilis, except
the rare pemphigus neonatorum, and the syphilitic causes of still
birth. No one has attempted to prove that miscarriage is any less
frequent among yawsy mothers than among the syphilitic.

(11.) The polymorphism of the eruptions.— This is abundantly
evident in yaws, and is referred to by Manson himself.

('2.) The nerve lesions and the gummata of syphilis. The negro
race is not prone to suffer from nerve syphilis, but all teitiaries and
Parasyphilis are taken from us by the simple assumption of no inter¬
ment syphilis, of which there may be no other evidence.

400

As far, then, as all these differentia are concerned, it is not possible
to separate the diseases. It is impossible to overlook the tertiary
stage of yaws. The indications for treatment are the same as for
syphilis. Even if the "syphilis theory” is not true, it is as if true,
as Paul Carus would say ; and, therefore, it is good to teach and
useful to believe. Any abstruse difference based upon the frambesial
eruption can only have an academic interest.

THE FRAMBESIA OR PAPILLOMATOUS SYPHILIDE

The frambesia has been variously referred to by writers as a
tubercle, a sore, granuloma, or a papule. No doubt, it is not of an
invariable form, but none of these terms are correct. In every case
the frambesia is raised, and consists of a closely-set group of hyper¬
trophied papillae. The dermis is not ulcerated, for properly the
eruption leaves no scar. The frambesia is sometimes naked and
moist, but more usually it is covered by a yellow crust. In some
instances, and especially when the eruption is old, the crust is dark or
dirty brown, but typically it should be yellow. There may be a
yellow secretion under the crust, and in the early and active stages
this is the rule. When old and undergoing resolution the frambesia
is dry, and the crust, if present, is more adherent. If the yaw has
been naked, it becomes in the dry stage covered with a film.

Usually the eruption disappears by slow absorption, each yaw
becoming imperceptibly smaller day by day. No doubt, yaws
occasionally ulcerates by accident or by bad treatment, and leaves a
true scar, but this is by no means its natural tendency.

As a rule, the frambesiae are scattered more or less profusely over
the skin. The corners of the mouth, around the anus and on the
genitals are the favourite sites. Typically each yaw is round or oval,
and measures 8 to 1 5 mm. across. But they vary much in size, and
may be much larger. They are sometimes arranged in a circinate
manner, and by coalescence of contiguous frambesiae a large,
irregular or horse-shoe form may be produced. It is, certainly, a
ery sti iking thing to witness the uniformity of the appearance of a
number of cases. One can go to a country village in a yaws district,
as in most parts of St. Vincent, and collect twenty children in five

401

minutes all showing the eruption in almost identical form. There is
small wonder that the profession, having regard to this eruption
alone, has almost generally fallen into the way of making it constitute
a distinct disease.

Alford Nicholls, who has made a thorough anatomical examination
by sections of the frambesial as well as of the papular and “ macular
eruptions of yaws (fig. XIX, in his report), shows that the papule oi
pian gratelle consists of a few hypertrophied papillae infiltrated with
small round cells, which are also accumulated in the underlying
dermis; the corneous layer of the epidermis is undergoing over¬
growth and exfoliation. This is identical with the structure of a
papule of syphilitic lichen. (The frambesia is beautifully shown in
section in fig. XXII.) It consists entirely of very much elongated
and hypertrophied papillae, with a great infiltration of round cells.
The epidermis is desquamated so that the ends of the papillae are
covered only by the secretion and the crust, which is formed of layers
of dry secretion. The papillae are closely crowded together. This
structure, without the crust, is identical with that of the syphilitic
condyloma. The frambesia is, in fact, one of the large papules of
syphilis. It is not, therefore, a granuloma in the sense that a gumma
is. The small round cells are certainly the elements of granulation
tissue, but the whole mass retains its physiological character as a

group of natural papillae, which do not disappear so as to leave a
scar. They have undergone an evolution and hypertrophy, and will
return to their normal size by involution and absorption of the added
elements. The term granuloma is a misleading one. The red
granular surface of the frambesia, seen after the removal of the crust,
to which it owes its name, from the resemblance to a raspberry, is not
due to granulations such as are on an ulcer.

The proper pathological description of the frambesia is in the
term papilloma ; and it is structurally identical with the papillomata
of syphilis, condylomata and rhagades. In a drying papilloma the
crust fits into the interstices between the papillae, so as to be difficult
to remove. But when it is removed, the individuality and tough
consistence of the papillae can be easily demonstrated.

On moist surfaces, as on the vulva and around the arms,
sometimes at the webs of the fingers and toes, the papilloma is naked.
T^e ends of the papillae are swollen, and squeezed together so

402

give a flat surface. Such frambesiae are indistinguishable from
ordinary condylomata.

We have seen that it is only in the frarnbesial eruption that yaws
differs irom syphilis, but even the frambesia is identical with the
syphilitic papilloma on a moist surface.

There is left, therefore, only the occurrence of condylomata all
over the skin which can be held to be distinctive of yaws, lonathan
Hutchinson points out that a general papillomatous (frambesiform)
eruption does occasionally occur in Europe. It has even been
suggested that “ Sibbens ” was a syphilide of this form.

It is quite feasible to argue that on the thick, highly-developed
and active skin of the dark races this exuberant form of eruption
takes the place of the milder papules known in Europe. The higher
infectiveness and greater prevalence of the frarnbesial syphilis is
readily explained. Syphilis ordinarily (that is, as known among
whites) is contracted from the primary and from moist papillomata-
mucous plaques and condylomata. Yaws, however, has so many of
the moist infective lesions on the surface of the body that contagion
is far more easily effected under any conditions of "life ; and in the
circumstances, and by reason of the habits of the natives, it must be
ost impossible foi each patient not to infect someone else
Infection can, of course, be effected venerea lly. Dr. Hatton, of

D R * have known cases occur from sexual connection.”1

r /’ y ’ . ,f"ada- reP°rts a case, presumably of venereal infection
■ ’ T ' W' ; 1 Cre°le' A‘ 10 ' — perfectly healthy and

“ Ma-t a ° Ct,n^ source contagion, until deflowered by a man in
“ the 1 . ourteen days after this she found some tubercles on

« It th'gh’ f°r Whlch 1 ad™tted *>er to the

and the nT * l**"5 'S suggestion that the man had yaws,

after L?raranCe; * ™ the «* «« thigh so soon

syphil s r:°n r“ e‘l the Case Unrdiabfe- Neither secondary
primary I yaWS “m ' ’erdeS " appear -me while after tire

pnma y sol tho°S 1 *“*«=»« -re all small

1 ‘ ^icholls’ Report, p. 17#

tolls’ Report, p. lg8'

403

It is a fact that yaws is far more common than syphilis in the
villages, and in towns vice versa. When a medical man sees yaws in
an adult, he never thinks of enquiring as to the possibility of its
having been acquired venereally, and when he sees a case of ordinary
syphilis he assumes the venereal infection, in spite, perhaps, of the
asseverations and protests of the patient. In the country districts of
yaws countries the syphilis, however acquired, probably for the most
part shows frambesiae ; while in the towns the non-venereal infection
gives rise to the better-recognised forms of syphilis associated in our
minds with venereal infection. Dr. I ulloch, of 1 obago, remarks
that the “ primary sore .... is especially rare in the outlying
districts where yaws is most common.’

More observations are wanted on two points :

(i.) Syphilis (without frambesiae) contracted from yaws.

(2.) Yaws (with frambesiae) contracted from ordinary syphilis.

As the date of infection will be three weeks or more, in both
cases, before the first manifestation of the primary, and usually two 01
three months before the appearance of the secondaries, it will be
extremely difficult to get histories, especially from ignorant peasants,
to illustrate these two points. I have already described one instance
of syphilis contracted from yaws (Case 19). Some cases aie
available of yaws supervening on venereal chancre, but the history of
these does not include any proof of the nature of the disease from
which the women suffered before they transmitted it as a venereal
chancre.

I shall describe one such case. Similar cases, though not
interpreted as 1 do, arc reported by Numa Rat, W. Boyd and
H. Bennett in the West Indies, and by W allbridge in Fiji.

Rat’s case is as follows:—" A man consulted the doctor about an
" indurated sore of the prepuce, and had at the same time a small
“ulcer on the ankle. During two months he was under mercurial
“ treatment, and had in this time sore throat, roseola, psoriasis and
“enlarged glands. He then noticed a change in the ulcer; the
gradations became pale and large. Three weeks after this the
ulcer healed. A week, again, after this came a wide-spread croP°
“papules. A few of these developed into frambesiae. e

syphilitic roseola co-existed with the yaws papules, and

“ * syphilitic psoriasis with the yaws tubercles.’1 A clearer case of
“ frambesial eruption supervening on venereal infection of syphilis
“cannot be found.”

W. Boyd’s case is given as follows .- — “ A man had a well-marked
“ and indolent Hunterian chancre, and bubo, when he came under my
“ notice, and at the same time a mild eruption of frambesial tubercles,
“ which he stated made their appearance two weeks after the
“ syphilitic sore.”2

H. Bennett’s case: — "The patient was a young woman, who
“ contracted the syphilis first, and then got either a primary or a
“ secondary ulcer inoculated in an unknown manner with the yaws."2
hrorn the confidence with which the syphilis is spoken of, one
presumes that a venereal chancre was known. I have, therefore,
included the case here.

Dr. Wallbridge’s case is quoted by J. Hutchinson3 : — ‘‘The
patient, a white man, had syphilis, and developed frambesiae. The
latter disappeared on his return to England, and later, after he
“ again went to Fiji, he suffered with tertiaries.”

In all these cases it is, of course, impossible to prove that the
persons from whom the disease was contracted had syphilis of
ordinary type.

There still remains one fact which forms the strongest objection
to the identification of yaws with syphilis. This, so far as I know,
has not been referred to by previous writers, and it remains for one,
who for the last twelve years has been holding and teaching the
syphilis theory, to raise what seems to be the strongest argument
against that theory. Although it may be true that the nature of the
dark skin predisposes to the frambesial eruption, it is indisputable
that yaws is seen breeding true on white persons. In a yawsy
village the poor whites get the disease and have the eruption just as
certainly as do the blacks. On a white skin the frambesiae are less
developed. The papilloma is not so prominent, and the crust less
irm. That is, the yaws is flatter, moister, and usually smaller than
a black patient, and it is associated with more of other syphilides,
psoriasis, large moist papules, ecthyma, &c. It is a condyloma

1. Nicholls’ Report, p. 168.

2. Nicholls’ Report, p. 166.

3. Fasciculus, p. 19.

405

of skin, nevertheless, and is of frequent occurrence ; while in Europe
such a lesion is so rare as to be a curiosity.

In considering a probable explanation of this phenomenon, we
must note that even among negroes the frambesial eruption is rarely
seen in towns, and while it is common in some West Indian islands,
it is rare or absent from others.

I have already suggested that damp, hot places in the tropics,
with dense vegetation, show a prevalence of yaws. This, I think,
will be found a rule. Numa Rat states that yaws is absent from
Anguilla,1 which is flat, dry and barren. The typical yaws district is
a deep valley or ravine in a tropical volcanic island, where vegetation
is rank and the rainfall high.

Yaws is, therefore, very prevalent in Dominica, St. Vincent,
St. Lucia and Grenada. It occurs to a less extent in Antigua,
St. Kitts and Nevis. Antigua is flat and not wooded, except at one
part. St. Kitts and Nevis, though lofty and volcanic, have long
slopes sweeping away to the sea. The land here is cleared and
cultivated. There are a few bushy ravines with villages, and in these
yaws may be found.

There is, then, another factor besides the character of the skin
which determines the appearance of frambesiae. We cannot yet
predicate this factor, any more than we can explain many other
examples of the variability of syphilis. Why, for instance, does
hpoid ulceration of the face happen so frequently on the Windward
side of St. Vincent, while destructive rhinopharyngitis is the rule on
the Leeward side? Why do some patients get a formative
periostitis, and others necrosis of bone ? 1 may be allowed to copy

from a recent letter of mine2 : — “ To understand this protean disease,
we must realise that various factors intervene to determine its
manifestations. We can appreciate some of these. T he races and
individuals who tax their nerve system suffer from nerve syphilis,
uhich is comparatively rare among negroes. The negro, with his
highly-specialised and active skin, displays an exuberance of
eruptions rarely seen in Europe. The labourer, exposed to injuries,
develops grave bone lesions. The women in St. Kitts, who

L RePort on Anguilla. Colonial Reprints, No. 20.

Journal of Tropical Medicine, May 15th, 1906.

qo6

“ habitually gratify the desire of sailors for paedicatio mulierum, gel
“ stricture of the rectum. Some of the determining factors we have
“ no knowledge of, others we can now guess at."

The frambesial eruption, therefore, though a striking and
characteristic feature of yaws, we have seen is not unknown in
syphilis. The difference between yaws and syphilis is only in the
frequency of occurrence of this eruption. If all that is called so is
yaws, it must be admitted that the frambesia is not even an essential
of yaws. In face of all the other points of identification, it is
impossible to give this one feature such pre-eminence as to
exclude the others. We must accept the papillomatous eruption only
as a phenomenon of syphilis, of which there are many others still
awaiting explanation. The enquiry into the reason of the frambesial
eruption may well produce a practical result, for if it were possible to
prevent syphilis taking this form, we should thereby materially limit
its infectiveness, and so lessen the amount of syphilis in the world
Here, then, is the field for research into yaws, which may yield a vast
store ol health for the unhappy natives of tropical lands.

Syphilis as yaws is almost general in tropical countries wherever
the conditions of damp, heat and dense vegetation obtain. It is
lecognised that coko in Fiji, puru in the Malay Peninsula, parangi in
Ceylon, galtoo in the West Coast of Africa, tomo in Samoa, are all
identical with yaws in the West Indies. Yaws is well-enough known
in every tropical country to bear a local name, and the synonyms are
as numerous as the languages of the peoples affected. It was endemic
m all parts of the tropical world before the European appeared on
these scenes to confuse the geographical distribution of races and
diseases by his facilities for inter-communication. Syphilis in all
forms is now universal, and was probably so from very ancient times,
ts wide spread proves. I think, that it must have been well

, . a a* an early period before the differentiation and

distribution of human races.

It has been suggested by Hutchinson that yaws was the original
lsease prevalent in the tropics, and that it was introduced into
uiope rom them. Under the influence of a different climate,
and the cleaner habits of the Europeans, it was limited and
modified until it became usually venereal. He thinks that Sibbens
an utton-scurvy were frambesial syphilis. Considering the wide

407

distribution of yaws, it is unlikely that the European countries could
have escaped infection until modern times. The forefathers of the
Caucasic peoples must have been already infected before they left
the race-home on the pleistocene plateau of Sahara.

The experiments of Metchnikoff and Roux have shown that apes
can be inoculated with syphilis. The femur of Pithecanthropus
erectus shows evidences of disease very suggestive of syphilis. I am
aware that these have been attributed to myositis ossificans, but one
would think that the latter is more likely to be a recent disease
resulting from the artificial circumstances of human life. In the
Chaldean epic of Izduhar or Gilgames, it is told how the demi-god,
having incurred the anger of Istar, was afflicted with a plague. An
eruption of sores covered his body, his bones ached, his strength
waned, his hair fell out. At length, under the advice of his beatified
ancestor, Hasisadra, he was restored to health by sea-bathing.
George Smith and other translators of the cuneiform script, following
him, have read this to be leprosy. But every eruption on the skin is
loosely so called by those writing on ancient matters. The leprosy
of the Levitical regulations (Lev., chapters 13 and 14) was certainly
not lepra. It no doubt included more than one disease, but most
likely it referred chiefly to yaws. I here is, perhaps, a reference to
psoriasis in the “ bright spots," and to frambesia in the appearance of
the raw quick flesh. The disease was one of rapid development, for
a suspect was kept under observation for seven days, and then foi
seven days more. Provision is made for restoring the outcast patient
to society when he became clean. 1 his would have been useless in
the case of leprosy. The suggestion that Leviticus refers to yaws is
a very old one, and has been discussed unfavourably by Alford
Nicholls. But there is no common disease except syphilis which
would explain all the regulations. True leprosy, no doubt, came
under the operation of the laws, and was dealt with along with the

syphilis.

It may be claimed that the symptoms of syphilis were sufficiently
"ell known to find a place in the folklore of the early Semitic tribes,
and to be subsequently included by the compilers of the traditional
history and the authors of the great poems; and it was found
Pessary to devise an elaborate legislation to prevent its spread.
The epic of Gilgames cannot be dated earlier than the second

408

millennium B.C. It has already enveloped in myth an invasion of the
Elamites, which occurred about 2300 B.C. But even that is a
respectable antiquity, and if syphilis was known to the poet of this
the oldest epic in existence, there has been ample time and
opportunity for it to spread to Europe before the middle of the second
millennium A.D.

It is now known that endemic non-venereal syphilis may flourish
even in Europe. Metchnikoff1 points out that in rural districts
in Russia the children are the chief sufferers and agents of
the spread of syphilis, just as they are of yaws in the tropics. There
is no reason for postponing the introduction of the disease into
Europe to the 16th century, when it is far more probable that it
has been present and spreading in a non-venereal manner as
long as the human race has been in existence.

I append a few cases of yaws, that is, cases that displayed the
frambesial eruption alone or, as is more usual, accompanied by other
syphilides.

Case 26. — C. W.; 8 years; Black.

January 23rd, 1906. Seen with an ulcer at the web of toes.

February 10th, 1906. Complained of pains, fever, and malaise, given liq.
hydrag. perch.

March 24th, 1906. Onychia of nearly all the toes, whole body spotted with
small round patches of scaly psoriasis. A large frambesia at the arms, which is
said to have appeared a week before. Large glands in both groins. Ulcer on the
plantar surface of web of toes is still in the same state as when first seen. Took
six weekly injections of salalembroth. The frambesia disappeared rapidly, the
skin cleared and the foot healed.

Discharged from treatment with directions to return in a month. Has not
been seen since.

Case 27. — C. B.; 3 years; Black.

March 1st, 1906. Has had yaws lately. One small frambesia still apparent
at the angle of mouth. Had glands on elbows, in back of neck, axillae, below both
angles of jaw, and in groins. I’rofuse eruption of small raw eroded surfaces,
not raised but covered with moist yellow pellicles, in the left axilla. Few similar,
drier and more crusted in the right axilla. Two such on the abdomen. Papules
of similar character, some moist and some crusted, on and behind both auricles.

Case 28. — M. P.; 8 years; Black.

March -th, 1903. Had a sore on right ankle some months before. Large

gland in right groin. Small hard glands on both elbows and many in the neck,

omplains of pain in knees and ankles. Several marks of previous yaws on the
egs. Several frambesiae with crusts 6n the right knee and the back. Papular
eruption (lichen) between shoulder blades and round the axillae.

1. Harben Lectures, 1906.

40Q

Case 29.— M. J.; 12 years; Black.

January 16th, 1903. Numerous small, perfectly-smooth limpets, which when
removed are horny cups covering a granular raised papilloma identical with
frambesia. These are over the face, hands, and body. They are about 6 to 10
mm. in size. A few of the papillomata have yellow crusts as is usual for
frambesiae. On the left ankle is a spreading patch of a mildly lupoid appearance,
the edge being active, and in the centre of this is an ordinary frambesia. This
is probably the mother yaw. Treated with liq. hydrarg. perch.

February 19th, 1903. The limpets and horns are larger.

February 12th, 1904. A small rupia over left eyebrow, small tubboes on both
soles. Circular wall of rupial scab on both hands at the base of the fourth and
fifth fingers. Rhagades between the fingers of the left hand. No marks of the
previous limpet eruption. This exemplifies a dry type of papilloma, and also
shows later secondaries of rupial type.

Case 30. — F. C.; age not noted; Black.

December 30th, 1902. lias a dry crusted frambesia on the wrist. Suppura¬
tive dermatitis of both palms and both soles. Multiple small pigmented marks
as of a vesicular or pustular eruption on both legs. Psoriasis on both elbows and
knees. Took pot. iod. for five weeks and left off attending.

I have seen papular groups, without scaliness, on the knees and elbows in
syphilitic cases so often, that I doubt the opinion that psoriasis in this situation
is never syphilitic.

September 14th, 1903. Tubboes on soles, and plantar .and palmar psoriasis
Icrab yaws), keratitis. V oice raucous. This illustrates the polymorphism of the
eruptions and the symmetry so characteristic of syphilis.

Case 31. — C. B.; 30 years; White.

January 2nd, 1906. Has a child with yaws. Ulcer on heel about 20 mm.
wide, with thickened raised edges, and ichorous discharge. This began as a
pimple two months before. The first eruption was that now on the same foot
and lower leg, which is a moist dermatitis. There are crusted ecthymata and
aciniform eruptions all over the body and sparsely on the limbs and face. Slight
fever every evening. Urine high coloured. Rheumatoid pains in shoulders.
Paronychia of one finger. No enlarged glands. Given liq. hydrag. perch.

February 13th, 1906. Profuse small spots of silvery scaly psoriasis on both
arms and thighs. In the bend of right elbow are three drying frambesiae
Several frambesiae round the trunk, and a profuse eruption of them on the left
foot. Few on the right leg. Groin glands enlarged. No fever pains or
malaise. All frambesiae are small and not much raised. Received a course °‘
seven injections of salalembroth during which the smaller eruptions faded, but
frambesiae continued to appear. Was rested for a month from mercury. e

end of this time the skin was nearly clean and only a few frambesiae left. on-
tinued on liq. hydrag. perch.

Case 32.— M. P.; 8 years; Black. , , ,

>906. Sent to hospital as a case of syphilis by a medical officer who had
ust come out. Frambesiae on two toes. Condyloma of anus. on y
jf prepuce and circinate plaques on the glans penis. F.nlarge §rmr* g a j
ncumcised and treated with six injections of salalembroth. vul e
discharged.

^ASE 33 — A- W.; 9 years; Black. , , k

1906. Two large frambesiae by the side of a scabbec sore °i even

' e !pfl f°ot; and whole body, limbs and face covered with spots o P . j r„e
Palms of the hands. Hard glands in both groins, left being especially large.
Clands on elbow.

4io

Case 34. — Female; 7 years; Black.

1906. Large irregular branched ulcer on a thickened base and with raised
edges, on the back of the left heel. Glands in both groins and shotty glands in
neck.

Several frambesiae on cheek and neck. Very large frambesiae at side and
back of one knee, dry crusted papillomata on forehead and back of neck. These
are essentially the same as yaws, but are dry, and have dark scabs. A
symmetrical lichen covers both buttocks and back of thighs.

Case 35. — L. K.; 19 years; Coloured.

April, 1906. — Had a venereal chancre last year accompanied by fever and
rheumatoid pains. Glands in both groins enlarged. Profuse eruptions of
frambesiform syphilides over both knees, extending to front of thighs and legs.
These are about 12 mm. each. Some are moist with yellow pellicles, under
which is a granular surface of elongated papillae. Some have dry brown crusts,
which when removed expose a frambesial surface.

There are pigmented pits of a recent vesicular eruption on the right arm and
the thighs.

The eruption in this case would have been called yaws by anyone.
It was atypical in two respects. It was quite symmetrical, and the
frambesiae were crowded together so as to be sometimes confluent.
Hut neither of th,ese features are exclusive of yaws. Such cases of
yaws following a venereal infection are, perhaps, more common than
is known. A syphilitic history is rarely enquired into when a patient
displays yaws.

Case 36. — M. W. ; 6 years, Black.

July 16th, 1906. Oval, flat, callous ulcer, with a glazed surface and pigmentec
areola, on the front of the left shin. Enlarged glands in both groins and both
elbows, several large glands in the neck.

A crop of small frambesiae with dirty-brown crusts on the forehead. Several
larger flat condylomata with slight crusts, or naked with moist yellow surface, on
t e ront of the neck and right cheek. Numerous papules varying in size
scattered on the face. On the left cheek a group larger, and almost frambesiform.
■ mall frambesia on the vulva. A profuse eruption of dry papules of all sizes
up to 4 or 5 mm. over both buttocks and back of thighs. On the elbows a warty
erupt,°" of lar8e Papules with hard dry horny tops. Many of the papules else-
" ere have a horny plug or horny apex of epithelium. There is a fine papular
erup ion on the front of the knees. Few scattered papules all over the skin.

is case well exemplifies the polymorphism of the yaws eruptions. There were
at least five syphilides of different appearance present at once

TREATMENT OF SYPHILIS IN NEGROES

If we decide to treat yaws, whether it be syphilis or not. with
mercury, it is necessary to appreciate that the' treatment must be
p severed in for many months. The frequent experience of the
recurrence of frambesial eruptions after being presumablv “ cured " by
mercury, indicates that the mercury must be continued, as in syphilis.

more or less through the first two years off and on. It will be found
a very difficult thing to get a peasant to understand or act up to this,
which in most cases will be only a counsel of perfection. When it
conies to the tertiaries and potassium iodide, the patients readily
leamfrom the relief obtained to value the drug. In St. Kitts and in
Fiji the natives are fully alive to the uses of iodide, and prescribe it
for themselves freely.

1 have noticed doses as high as ninety minims of liq.. hydrag.
perch, mentioned in some of the literature on yaws, and there is very
frequent reference to salivation. I wish to draw attention to the fact
that negroes bear mercury badly. My father, a keen syphilist with
many years’ experience of treating negroes in hospital, where one can
see best what one is doing, warned me of this. Fifteen minims three
times a day is, I believe, about, his limit. My own experience has
amply borne out this. Though one is not afraid of “ touching the
gums of a patient, and, in fact, generally likes to see them just
"touched,” as an indication that mercury is being pushed to its limit,
yet salivation is undesirable. Apart from any physical effect it may
have on the health or strength of the patient, it is apt to shake his
confidence, and perhaps frighten him away altogether. It is
astonishing how quickly some blacks do become salivated. I have
seen a woman very badly ptyalised by five grains of calomel. Blue
ointment applied to a wrist for a few days salivated a man, and a
young woman became salivated before she had finished using half an
ounce of blue ointment by daily inunction of about one dram. In
he last case, there was certainly no idiosyncrasy ; she had taken
mercury, by the mouth in small doses, freely before, and has since
received a course of intramuscular injection of 003 gme of
salalembroth weekly, equivalent to about 'i gr. of corrosi\e
sublimate.

1 have used mercury by inunction of blue ointment, mouth
administration of liq. hydrarg. perch., calomel, and grey powder, and
intramuscular injections of salalembroth. After a fairly large
exPerience, I am confident that mercury is borne best when given by
tne last method Inunction may salivate quickly and suddenly. In
hospital one can watch it, but even so the ptyalism may7 occur
suddenly. Grey powder with carbonate of iron is an excellent
treatment, especially for children with hereditary taint or yaws. It

412

is convenient also for adults, for they can be given a hundred pills at
a time, which last a month. I do not like to exceed o'l gm. thrice a
day. Calomel I do not value for antisyphilitic treatment; but for
general use in dispensary work the liq. hydrarg. perch, in a mixture is
the most useful. I generally colour this lightly with methyl blue,
which gives an impressive and distinctive value to the bottle. One
should begin at the rate of 2 c.cm. of liq. hydrag. perch, per day — say,
10 minims three times a day, and increase to 3 c.cm. It is hardly
necessary to go higher than this. Children bear this and any
mercury well in proportion to the adults. For an infant of three
months with inherited disease I give O' 15 c.cm. three times a day,
equal to about two and a half minims.

Intramuscular injection of soluble salts is, however, far the best
mode of administering mercury in my experience. I cannot speak of
the insoluble preparations, for, being quite satisfied with the salalem-
broth, I have not tried the emulsion of metallic mercury so highly
spoken of by some. In spite of the pain, I have found that out¬
patients attend very well. Out of a number who were to take six
injections each, the average attendance was five times. I have used
this method on infants with hereditary syphilis also, and in yaws.
My experience is not yet very large, but I have had abundant
opportunity of proving that some negroes can carry up to i'5 c.cm. of a
2 per cent, solution of mercury perchloride without any sign of spongy
gums. This is equivalent tp nearly half a grain of the salt, or about
half an ounce of liq. hydrag. perch. Usually, I begin with 07 c.cm. and
work up to I'O, using a solution containing mercury perchloride 2 per
cent., ammonium chloride 1 per cent, and giving one injection weekly.
With care, washing the buttock with an antiseptic and then with
ether, I have not yet seen an abscess result, and only rarely any
induration. For hospital treatment, or for regular yaws dispensaries,
the intramuscular method should be the most certain, the easiest and
most effectual method of giving mercury. The treatment is shorter,
and for that reason more useful in the case of natives. It should be
possible to get most cases to attend for six or eight weeks for a course
o injections, and repeat this twice more within the first year. After
is, one attendance a month for another six months. The symptoms
that may be expected should be explained to the patient, in order that
0 ma^ at °nce rePort himself on the occurrence of any of them ; and

4T3

certainly it is our duty to enlarge to any extent on the terrors of the
tertiaries.

There is no necessity for special yaws asylums, but it is very
desirable to have some provision for hospital treatment of severe and
cachetic cases. Special wards in a general hospital would meet the
requirements.

In the local treatment of yaws, I have obtained best results from
a paint of zinc oxide and calamine, an old-fashioned but ever-useful
preparation. It is readily applied by the patient or his parents ; it
is not easily wiped off, and, most important of all, it keeps the
frambesia dry. I feel confident that a case of yaws is not very likely
to infect other persons from his skin lesions as long as the frambesiae
are kept dry and well coated with this paint. The risk of tonsillar
infection from spoons and cups cannot, of course, be avoided in the
peasant class, and that this is a frequent mode of infection with
syphilis is well recognised.1 But the most important material of
infection in yaws is the secretion of the frambesia, and this can be
limited by local applications, and by' causing the absorption of the
papilloma as rapidly as possible with mercury internally.

*• Kyle, Diseases of the Throat, p. 464.
AA

f

• ' jw:: \ '■

A DESCRIPTION OF SOME GOLD COAST
ENTOMOSTRACA

417

A DESCRIPTION OF SOME GOLD COAST
ENTOMOSTRACA*

BY

W. M. GRAHAM, B.A., M B.

It is a matter of common observation in the Gold Coast Colony,
Ashanti and the Northern Territories that the incidence of Guinea
worm varies locally. Some stations shew a high percentage of cases,
some a low percentage, and some are free from the disease. \\ hy
should such local variations exist ? The probable explanation is the
abundance, the variety or the absence of the Cyclops-host from the
water sources of the locality. Cyclops belongs to a widely-distributed
family, and some species of the family can be found readily in the
water sources of every station. But as Guinea worm disease is not
present at every station, it is evident that the presence of any species
in the water is not sufficient to cause the disease. I he inference
seems probable that a special species of Cyclops is required to act as
efficient host to the worm, and that the absence of such species from
the water sources of a station coincides with the absence of Guinea
worm disease. If this be true, then an exact knowledge of the
species to be found at each station becomes of great practical
importance, but there is at present no classification of Gold Coast
Copepoda. I have, therefore, sought in the present article to furnish
means for the identification of the species found in the water sources
°f the station of Obuasi, and in those of the country within a three-
mile radius round it, as a first instalment. Obuasi is a mining village
on the Gold Coast Government Railway, 124 miles from Sekondi.
The country is hilly and covered with forest. During the dry season
the streams are small, sluggish and full of weed. With the setting in
the heavy rains thev become rapid torrents, and the weed
disappears. During the first period, to the end of April, Cyclops
1 to 5 can be caught in the streams, but after the first week in May

il •r^l'e lhanks of the Editors are herewith given to Dr. Bradv for his kindness in
' entifving and describing this collection of Entomostraca. — Eds.

I have failed to find them there. The country is covered with jj-
holes of varying depth, and along the railway by borrow-pits. 11
water in the shallower pits drys up, and by the end of the dry sc;
the bottoms of the pits have been baked dust dry. After a week
heavy rain these pits begin to retain a permanent pool of wate:
Upon this pool frog spawn soon appears, and shortly afterward-
adult females of No. 6 ( Cyclops leuckarti , Claus) can be found. B\
the middle of May the intermediate nauplius forms have appeared
Then the females become gradually less numerous and males incrc i -
in number, until by the middle of J uly the females have become ran-
than the males.

Meanwhile, Cyclops Nos. 2, 3, 4 and 5 appear in the pools in
relatively small numbers, and are soon accompanied by their naup! in¬
forms.

Lastly, Cyclops No. 7 (C. simillimus , sp. nov., Graham) app<
in the pools (the females first, the males later), and seems to take th>
place vacated by C. leuckarti.

This account brings up the history to the end of July; and 1 w. -
transferred early in August to another station.

The inference previously alluded to, that all species of Copepoda
cannot act as efficient hosts to Guinea worm, is strengthened by th<
following considerations: —

ist.-

- 1 here is a large number of species.

_nd. The habits of the different species vary greatly. Some a
surface feeders, some are found at the bottom. Son
n abit foul, some clean water. Some leave the water i
c imb on stalks of water-weed enveloped in a drop of waU
carried with them, some do not leave the water. Some a:
ound in streams, some are not

3rA”hJhe tdlffeJre”t sPedes are infested by d.fferent parasites
ounc os. 3, 4 and 5 infested only by ecto-parasite
(algae) and No. 6 infested by ento-paras.tes (worms).

4 b.- rhe d.fferent species d.ffer in the date of then appears

or at?0:15' , “e are fOUnd earl>’ ■" May; some aPP«,
of' th' CaS ’ °n y became numerous, in July. The importaK.
It* r,atl°n m date has b«n already explamed b,

me m a former article (B.M.J., 1I/II 05).

A careful examination of the water sources of each station, and a
classification of the Copepoda found in each place, is urgently
required When it has been made, a comparison of the species of
Cyclops found at a station where Guinea worm is common with the
species of Cyclops found at a station where Guinea worm disease is
absent would, I have no doubt, offer an explanation of the observed
local variation in the incidence of the disease. 1 he practical value of
an explanation is evident. It would enable the Medical Officer to
identify those water sources likely to aid in the propagation of the
malady.

Cyclops No. i. ? Cyclops bicolor, G. O. Sars.

Male. — Colour yellowish, with darker coloured first antennae.
First antennae, 11 segments as long as f cephalothorax. lotal
length, 0-46 mm. Cephalothorax and thorax, 0-30 mm. Furca,

0 03 mm.

Female. — Colour as in male. First antennae as in male, but
more slender ; total length, 0-62 mm. Cephalothorax and thoiax,
0-42 mm. Furca, 0'04 mm. Egg sacs, a pinkish colour, carried apait.

In both series the outward tail sita is long. F ound in pool w ith
Calanus No. 1 ; water clear, and used by village as supply.

Cyclops No. 2. Cyclops varicoid.es , sp. nov. (Brady)

Male. — Colour, transparent pale yellow. First antennae, 12
segments, nearly as long as cephalothorax. lotal length, o 59 mm-
Cephalothorax and thorax, 038 mm. F urea, o'04 mm.

Female.— Colour as in male. First antennae as above, but more
slender. Total length, 0-64 mm. Cephalothorax and thorax,
0-39 mm. Furca, 0'04 mm. Egg sacs, a yellow colour, carried close
together.

I-'ound in river water.

Cyclops No. 3. Cyclops longislylis , sp. nov. (Brady)

Male.— Colour, a pale yellowish -green. First antennae, 12
segments as long as cephalothorax and two thoracic segments.
Total length, 070 mm. Cephalothorax and thorax, o-45 mm.
furca, 000 mm.

420

FEMALE. Colour as above. 1- irst antennae as above, but
slender. Total length, 0-84 mm. Cephalothorax and thorax, crjc

mm. Furca, 01 2 mm. Egg sacs, a violet colour, carried slightly
apart.

This species is frequently covered partially or entirely by ecto¬
parasites (algae).

It can be readily distinguished from No. 2 by the great length of
the furca.

Cyclops No. 4. Cyclops virescens, sp. nov. (Brady)

Male. Colour, cephalothorax a pale yellow-green; thorax a
dark green. First antennae, ,0 segments; copper coloured,
earned a sparkling spot below the eye when swimming ; an air bubble

m mouth. Total length, 0-4; mm. Cephalothorax and thorax,
029 mm. Furca, 0-03 mm.

Female.— Colour as above. First antennae as above, but lighter
erm Ur‘ T7T°tal length’ 0'60 mm- Cephalothorax and thorax,
close Together3' ^

Caught in rapid streams, and later on in year in ponds.

Cyclops No. 5. Cyclops pheleratus, Koch

longMa?th7COlT; aubnght C0pper' F,rst antennae' 10 segments, as
taa a t ’ r °nly' T°taI '“*h. o-6o mm, Cephalc
colour ' 0 38 ”m' FurCa' °'°5 Cogs of a light blue

length, 077 m^°loUr as abo''e- Flrst antennae as above. Total
008 mm Emr Cephalothorax and thorax, 0 46 mm. Furca.

This sneeffs r a bnght b‘Ue COl°Ur' Carried close together.

the ! rter read“y and dlmb= - sides of

essel carrying a drop of water with it.

C\ CLOPS No. 6. Cyclops Icuckarli , Claus

segments, as^on^Y^ yeIlow_Sreen- first antennae. 17

0-89 mm.' Cephalothorax and ^ 'T'

the m-ddTeeoPfleiyUl ““ * ^ 1 have «* foundries Wore

421

FEMALE— Colour as above, but not so very pale. F irst antennae
as above. Total length, 1*04 mm. Cephalothorax and thorax,
065 mm. Furca, O' 10 mm. Egg sacs (white) transparent, a long
oval, egg very circular, sacs carried at an angle of more than 450.

Found in stagnant pools in May. Become scarce in July.

Cyclops No. 7. Cyclops simillimus , sp. nov. (Graham)

MALE.— Colour, very pale salmon with orange spots (circular) in
cephalothorax. First antennae, 17 segments, somewhat longer than
cephalothorax and thorax. Total length, 062 mm. Cephalothorax
and thorax, o- 3 5 mm. F'urca, 0^05 mm.

Female. — Colour, pale yellow-green with circular orange spots in
cephalothorax. First antennae as above, a very pale yellow. Length
as above. Total length, o' 80 mm. Cephalothorax and thorax,
049 mm. Furca, 0*05 mm. Egg sacs, a pale yellow, carried apart.

This species also usually carries an air bubble in mouth when
swimming. It differs from No. 6, being smaller and having relatively
shorter tail setae.

Cyclops No. 8. {non del.)

Male. — Not found.

Female. — Colour transparent, with slightly milky spots in cephalo¬
thorax and thorax. First antennae, 17 segments, as long as cephalo¬
thorax and thorax. Total length, O'/g mm. Cephalothorax and
thorax, 0^48 mm. Furca, 0^09 mm. Egg sacs small, milky white,
carried apart.

Caught in well half a mile from the sea. Water in well clear, but
slightly brackish. F rogs in well.

Cal anus No. 1. = Diaptomus innominatus , sp. nov. (Brady)

Male. — Colour, a very pale blue, the antennae being somewhat
darker. First antennae, left 25 segments, right 22 segments, modified
as Jasper. Second antennae, 8 segments. Total length, 0‘95 mm.
Cephalothorax and thorax, 0^67 mm. F urea, 0'05 mm. Legs,
central three pair a violet colour ; fifth pair modified as claspers.

Female. — Colour as above. First antennae, 25 segments.
Second antennae, as above in male. Total length, 1 10 mm.

422

Cephalothorax and thorax, 075 mm. Furca, 0-07 mm. Egg sac,
slightly blue, carried beneath.

When swimming appear transparent, with a dark blue longi¬
tudinal spot in about centre of cephalothorax and thorax.

I have only been able to find it in a single pond under trees, in
which Cyclops No. 1 was also present.

CANTHOCAMPTUS No. i. = Attheyella africana, sp. nov. (Brady)

Female. — Colour, a bright orange. First antennae, eight
segments. Second antennae, four segments, forked. Total
length, 0 35 mm. Cephalothorax and thorax, O’ 17 mm. Egg sac
nearly as long as body.

Very common in certain pools, where they occur in enormous
numbers.

423

NOTES ON DR. GRAHAM’S COLLECTION
OF CYCLOPIDAE FROM THE AFRICAN
GOLD COAST

BV

G. STEWARDSON BRADY, M.D., LL.D., D.Sc., F.R.S.

( Received May 14th, 1907)

The following notes are intended merely as guides to the
identification of species, and do not attempt a complete morphological
account of the various forms. The brief descriptions are based upon
the examination of females only, the males having been left out of the
account.

In addition to the various species of C yclops , the collection
contained specimens of a minute Cyprid, a species of Diaptomus, and
one called by Dr. Graham C anlhocamptus, but which belongs to the
nearly-related genus Attheyella. Both these last-named forms
appear to be new, and I propose for them the names Diaptomus
innominalus and Attheyella africana ; they, however, need complete
figures of structural detail, which at present I am unable to give.

GENUS CYCLOPS

No. i. ? Cyclops bicolor, G. O. Sars.

No. 2. Cyclops varicoides, sp. nov.

Anterior antennae twelve-jointed, reaching to the third cephalo¬
thoracic segment, both branches of the first four pairs of swimming
feet three-jointed, fifth pair rudimentary ; caudal stylets short, about
as long the last caudal segment. Length of body, exclusive of tail
setae, 077 mm.

No. 3. Cyclops longistylis, sp. nov.

Anterior antennae twelve-jointed, reaching to the posterior
border of the first cephalothoracic segment, both branches of the
first four pairs of swimming feet three- jointed, fifth pair consisting o a

424

single small papilliform joint with two terminal setae ; caudal stylets
long and slender, about six times as long as broad, and equal in
length to the three preceding caudal segments. Length of body,
078 mm.

No. 4. Cyclops virescens, sp. nov.

Anterior antennae ten-jointed, reaching to the third cephalo¬
thoracic segment, first four pairs of feet having both branches three-
jointed, fifth pair minute, papilliform, bearing two long apical setae;
caudal stylets short, about equal in length to the last caudal segment.
Length of body, 06 mm.

This species is very similar to C. gracilis . Lilljebon, but differs in
having all the branches of the swimming feet triarticulate ; all the
inner branches in C. gracilis being biarticulate ; there are also other
minor differences.

No. 5. Cyclops phaleratus , Koch

No. 6. Cyclops leuckarti , Claus.

No. 7. Cyclops simillimus , sp. nov.

Anterior antennae seventeen-jointed, reaching nearly to the

postenor extrem.ty of the cephalothorax, all branches of the

swimming feet triarticulate, fifth pair biarticulate, last joint simple,

narrow and bearing two long apical setae ; caudal stylets about twice

as long as broad, and nearly twice as long as the last caudal segment
Length, 077 mm.

.\2(i

EXPLANATION OF PLATE XXXIII

Cyclops bicolor, G. O. Sars
Fig. i. — Female with ovisacs. x 74.

Fig. 2. — Male. x 74.

Cyclops varicoides, n. sp., Brady
Fig- 3- — Female with ovisacs. x 74.

Fig. 4. — Male. x 74.

Cyclops longistylis, n. sp., Brady
Fig-- 5- — Female with ovisac. x 74.

Fig. 6. — Male. x 74.

The figures on this and the subsequent plates are reduced from
1Jr- Oraham s original drawings. _ Eds.

PL A TE XXXIII

to- Graham, ad. not. dtl.

EXPLANATION OF PLATE XXXIV

Cyclops virescens, sp. nov., Brady
Fig. y. — Male. x ;4.

Fig. 8.— Female with ovisacs. x 74.

Cyclops phaleratus, Koch
h ig. 9. — Female with ovisacs. x 74.

Fig. 10. — Immature, x 74.

Fig. 1 1. —Male. x ;4.

Cyclops leuckarti, Claus

* 54-

Fig. 1 2.- -Male.

PLATE XXXIV

H ■ M. Graham, aJ. not. dtl.

- -

430

EXPLANATION OF PLATE XXXV

Cyclops leuckarti, Claus
Fig- 13- — Female with ovisacs. x 74.

Fig. 14. — Immature, x 54.

Cyclops simillimus, sp. nov., Brady
Fig- J5- — Male. x 74.

Fig- 16. — Female with ovisacs. x 74.

Cyclops, ? Sp.

I ig- 17. There were no specimens of this species in Dr. Graham'?

collection when submitted to Dr. Brady for determination

PLATE XXXV

'V/. Graham, ,i J, „«l. J,l.

EXPLANATION OF PLATE XXXVI

Diaptomus INNOMINATUS, sp. nov., Brady
Fig. 1 8. — Female. x 124.

Fig. 19. Male. x 124.

Fig. 20. — Male, fifth pair of legs. x 102.

Pig. 21. — Female, fifth pair of legs. x 102.

Attheyella AFRICANA, sp. nov., Brady
Fig. 22. — Male, x 74.

Fig. 23. -Female, x 74.

PLATE XXXVI

“ -iX Graham, ad. nat. d,l.

ON THE MORPHOLOGY AND LIFE
HISTORY OF SPIROCHAETA DUTTON I

I

435

ON THE MORPHOLOGY AND LIFE
HISTORY OF SPIROCHAETA D UTTONI

ANTON BREINL, M.U.Dr. Prag

DIRECTOR OF TH K RUNCORN RESEARCH LABORATORIES OK THE LIVERPOOL
SCHOOL OF TROPICAL MEDICINE

From the Runcorn Research Laboratories of the Liverpool School
of Tropical Medicine

(Received July 1st , 11)07)

In the course of our experimental study of the spirochaete of
African Relapsing fever, Spirocheeta duttoni} numerous films were
made of the blood, and of the organs, of animals in the diffeient stages
of the disease, with a view of ascertaining the life-history of the
parasite. Considerable uncertainty still exists concerning the
morphology and the life cycle of the whole group of spirochaetes.
While Novy and Knapp2 deny that the parasites undergo any
morphological changes, Prowazek,3 in his work on fowl spirochaetes,
gives them an undulating membrane and a definite nuclear apparatus,
and also describes intra-cellular stages. One terminal flagellum was
observed by Novy and Knapp in Spirocheeta obermeieri , and by
Stephens4 in Spirocheeta duttoni ; Zettnow,5 on the other hand,
describes peritrichal flagella in Spirocheeta duttoni , and Borrel in
fowl spirochaetes.

The stain employed in the present work was Giemsa’s modification
of Romanowsky’s stain, in dry films. Wet films were not found
possess any particular advantage when working with this parasite.

The spirochaete consists of a darkly stained central core, which is
surrounded by a light-stained periplastic sheath. This sheath exten
beyond the central chromatic core, and is drawn out at one en
an elongated filament, thus forming a structure which has been
described by various observers as a terminal flagellum. a emp

436

to demonstrate peritrichal flagella, either in fresh or in stained
specimens, have completely failed. The central core, or chromatic
part, does not always stain uniformly, but in certain parasites lighter
and darker areas are noticeable, either throughout the whole length
or confined to one part of the parasite (figs, i, 2). Very frequently,
especially in spirochaetes which are disappearing from the circulation,
the whole chromatic core seems to be broken up into an irregular
number of granules (fig. 3). In this stage of infection the spirochaetes
often, show one or more swellings, either in the centre or at one end
(figs. 4, 5). A fairly constant appearance, which has been previously
described by various observers in Spirochoeta obermeieri and
Spirochczta duttoni, is a small unstained transverse band situated at
about one-third of the length of the parasite (fig. 6).

A considerable amount of work was done with the object of
demonstrating an undulating membrane. Although in specimens
stained with Giemsa’s solution an appearance was sometimes seen
which resembled an undulating membrane, this was, in my opinion,
due only to the flatten, ng out of the spirals of the paras, te. In wet
trims, even after a prolonged staining by Heidenha.n's method, no
trace of an undulating membrane could be seen.

The division of Spirochata duttoni is, as a rule, transverse,
the paras, tes mcrease in length and become thinner in the middle;

is mner part then elongates more and more until the two
individuals separate (figs. 7, 8). It is very probable that the unstained

future HUe y 56611 T para51te <%■ <9 » ‘he point of the

loTg udtfd °n “ SUbS6qUent f~rse d-sion. Occasionally,
IheTsaooe dlVIS1°nfWaS take place, especially at the time of

and tWs r3nCe ' 3f the Parasites from the peripheral circulation,
(o' increase ° “ the °^ns the paras, te was seen

spirochaete and 't T^t dlV,S,on commencing at one end of the
Judging from tl ^ eXtendmS along its entire length (fig. g)

appear that this' process^ f T P^S'tes at this stafTe, it would
instances, at this stage of the ““P"at,velr . °ne. In rare

engulfed by phagocytes. ’ parasites were seen beinS

occasions,' seen inZT'j m I0' «*■

close to each other t h° W° sPlrochaetes were observed lying
other, touch, ng at certa.n points. The one was stained

437

dark red with Giemsa, the other light blue, with apparently no
chromatic core, but showing an irregular number of dark red granules
situated at the points at which the two spirochaetes were in
apposition. We are inclined to explain this appearance as
conjugation.

Prowazek describes intra-cellular stages of Spirochceia gal l in arum
in the red blood cells. We were able to observe the same phenomenon
in rare instances with SpirocJurta duttoni just before the crisis set in

(%• n).

Although the appearance of the parasites in the peripheral blood
seemed fairly uniform, striking changes were observed in parasites
seen in the organs, notably in the spleen, bone marrow, and liver.

Numerous spirochaetes, especially just before the crisis, when the
blood was still swarming with parasites (principally in the spleen and
bone marrow, rarely in the liver), were seen coiling themselves up
1% I2), a few presenting a swollen appearance (fig. 13), the majority
gradually becoming thinner and rolling themselves up into more and
more complicated skein-like forms (figs. 14, 15) which seemed to
become more irregular as the time of the crisis drew near The
majority of these forms were devoured by the phagocytes of the
spleen, and at the time of the crisis the spleen cells were observed to
be gorged with degenerated spirochaetes. In animals in which the
spleen had been removed an analogous process took place in the liver.

A few similarly shaped parasites underwent a remarkable
change:— The outline remained more regular for a time, and the
parasite surrounded itself with a thin cyst wall, the interior of the
c.vst being filled with a faintly blue stained plasma (fig. 16). These
knns, in scanty numbers, were to be seen even after all the other
krrns had disappeared. They apparently undergo further changes,
as the shape of the parasite becomes more and more indistinct and,
at a still later stage, only the cyst filled with small red granules
Persists.

We were unable to trace the further development of these forms,
as in specimens stained by Giemsa’s method it is impossible to
differentiate them from blood platelets and other constituents. The
'act that the filtrate of spirochaetal blood through a Berkfeld filter is
infective suggests that these small granules may be the forms which
Pass through the filter and give rise to a fresh infection.

438

The life history of the spirochaete might be thus summarised
Just before the crisis the spirochaetes disintegrate, certain of them
coiling up into skeins, the majority of which are phagocytosed by the
spleen. Some of them become encysted and break up into very small
bodies, out of which the new generation of spirochaetes is evolved

LITERATURE

1. Breinl and Kinghorn. An Experimental Study of the Parasite of the African

Tick Fever ( Sfirochala dultoni). Memoir XXI of the Liverpool School of
Tropical Medicine, 1906.

2. Novy and Knapp. Studies in Sfirillum obermeieri and related organisms.
“Journal of Infectious Diseases,” Vol. Ill, No. 3, 1906.

3. Prowazek. Morphologische und entwicklungsgeschichtliche Untersuchungen

iiber Hiihnerspirochaeten.

“ Arbeiten aus dem kaiserlichen Gesundheitsamte,” Vol. XXIII, No. 2, 1906.

4. Stephens. A Note on the Structure of Sfirochteta dultoni. “Lancet,”

August 18th, 1906.

5. Zettnow. Farbung und Theilung bei Spirochaeten.

“ Zeitschrift fur Hygiene und Infektionskrankheiten,” Vol. LII., Heft 3, 1906.
Zettnow. Geisseln bei Jliihner und Recurrensspirochaeten. “Deutsche
Medizinische Wochenschrift,” March 8, 1906, page 276.
f>. Borrel. Cils et division transversale chez le Spirille de la poule.

“ Comptes rendus de la Soci6t<$ de Biologie, Tome 60, p. 238, 1906.

Some forms described in the present paper were observed by this author.

EXPLANATION OF PLATE

1 he accompanying drawings were done with a Zeiss apochromatic
objective 2 mm. aperture r4, ocular 18. Drawn to the scale of 4-5^

g 1 to i_. Fiom the peripheral circulation of infected monkeys
and rats, respectively.

i ig- 13— From the liver of an infected monkey.

HgS’ 14 to l6*~ Forms found in the spleen.

Fig. 4

PLATE XXXVII

THE

CYTOLOGY OF THE TRYPANOSOMES

44i

THE

CYTOLOGY OF THE TRYPANOSOMES

PART I*

BY

J. E. SALVIN-MOORE, A.R.C.S., F.L.S., F.Z.S.

PROFESSOR OF EXPERIMENTAL AND PATHOLOGICAL CYTOLOGY, UNIVERSIIY
OF LIVERPOOL

AND

ANTON BREINL, M.U.Dr. (Prag.)

DIRECTOR OF THE RUNCORN RESEARCH LABORATORIES, LIVERPOOL SCHOOL OF
TROPICAL MEDICINE

( Received July 15th , 1907 )

CONTENTS

PAGE

A. Introductory .

B. The morphology and the life cycle of Trypanosoma

gambiense .

C. The morphology, and the multiplication in the blood of

Trypanosoma brucei .

D. The morphology, and the multiplication in the blood of

Trypanosoma equinum .

E. Consideration of the foregoing observations .

Appendix I .

Appendix II . . .

44 1
448

459

459

460

470

47 1

(A) INTRODUCTORY

The trypanosomes belong to a group of organisms of great
practical importance, since they are related to numeious diseases,
not only affecting many valuable animals, but, in the case of sleeping
sickness, man also. Notwithstanding the facts, the natuie an

* A preliminary account of the observations relating to 7 ■ ^^T^iqoT^Yn a
in the present paper was published in the Lancet, p. ' * j Society , July 20,

subsequent paper by Plimmer and Thomson received by t Trvnanosomes, or

■907, these authors appear to have encountered either the ency - -bed But from
the resistant bodies (latent bodies) which we had previous y • ' it is

the vagueness of their reference, Pro. Roy. boc. «• • described by us

impossible to be certain to which order of structures a >
they do refer.

442

morphology of these organisms are as yet but little understood
Information upon these matters, as well as upon the various lifecycles
they appear to present, is greatly t. 1 be desired Especially is this the
case with regard to their morphology. The numerous descriptions of
their structure and their metamorphoses already in existence have
been drawn in general from the results of methods of research not
calculated to produce any correct appreciation of their true cytological
features. With very few exceptions, the study of the structure of the
trypanosomes has been based, cither upon what can be made out of
the live animals, or else upon observations made upon material after
it has been heated and dried, a method which, to say the least of it,
may be shown, so far .is the finer details of any cell structure are
concerned, to be particularly barbarous.

Trypanosomes, like other unicellular organisms, can, however, be
fixed m a great variety of ways which are commonly used during
cytological research. Hie chief manipulative difficulty 1 1 icv present
is the unreadiness with which they can be made to take any sort
of differential stain. Still this difficulty is not insurmountable; and
ordinary preparations may be produced which stain as completely as
can be desired.

We have used the following fixatives .-—Fleming’s fl«ic
sublimate acetic acid ; osmic acid vapour ; osmic acetic acid vapow
and formalin vapour.

When the animals have been fixed, it is in all cases des, table
use somewhat spec.ai precautions in relation to the stain which t*

Dri'nr i° t e"!ployed' the Pr°cess adopted depending upon tl
are ! °f “PP'yng a mordant, or mordants, before the actual s«ir

FlemtoU fl h •the Wh°Ie’ WC haVe f°Und that ^ fixa,ion "*
of vTe?wh T UnqUest,onab,y rt- best from a morphological f*
the sharpest ^ methods trough which we have obtain*

safrantrin atT,haVC bee”' ^ hand, the
Appendix II "e ™ethyIenc blue atain invented by Breinl (**

iron h" “a to T ° ' 3 S%ht modifi-tio„ of the Heid^”

iron haematoxylm process (see Appendix II)

which5 wTuabr: StTitf.!®1' (and in p-b*<

is necessary in the first il ' d,fferent sPecific {o^\e

.. £ zzzzx?. zsz#

443

This course is unavoidable owing to the confused terminology which
has grown up in the literature, and also on account of the present
necessity of making clear the meaning we attach to different names.
We have further to define our present conception of the nature of
several structures which the trypanosomes possess.

When properly fixed, all the animals we have examined present
an elongated cell form. No anterior or posterior extremity can, except
in the most arbitrary sense, be defined. I he exterior of the protoplasm
is differentiated into a thin outer layer or ectosarc (periplast). Among
the species with which this paper is concerned, the ectosarc is smooth,
and does not present any definite ridges or stripes corresponding to
the structure often described in the larger trypanosomes, such as those
of the frog, and others.

The protoplasmic structure within the ectosarc consists of a \ery
coarse spongioplasm (schaumplasma) containing fine staining granules
embedded in its substance, the meshes of this spongioplasmic netw ork
being filled by a less stainable cytolymph. It is sometimes said that
within the ectosarc, and distinct from the deeper portions of the
spongioplasm, there exists a layer — the endoplasma. We have,
however, not been able to demonstrate the existence of this sub¬
division.

The permanent cell structures contained within the ectosarc
consist of a more or less central area, which, when subjected to Breinl s
stain, assumes a purple colour (see figs. 4, 5, 1 1, 12, 13)- We propose
to call the whole of this area the nucleus. Within the nucleus there
is always to be found a clearly-defined body which stains undei the
same conditions red, and we propose, for reasons which will become
more apparent later, to term this body the inlra-nuclear ccntrosome
(Karyosome, Innenkorper).

It does not appear to be the case, when the animals arc not
dividing, that the nucleus can be correctly said to be bounded by any
definite membrane. In most instances it appears more correct to
say. that there is no definite membrane, but rather that there is a
very sharp division between the spongioplasmic network and the
finer network of the nucleus.

Near the broad end of the animal’s body there is usually to be
found a granule, or small group of granules, which stain like the intr
nuclear centrosome. These, whatever their numbers at any particu a

444

period, we propose to call the extra-nuclear centrosomes (blepharo-
plasts). From one or more of these granules there springs a staining
core, or flagellum, which lies in a thin expansion of the ectosarc,
forming the so-called undulating membrane

For present purposes we have thus the following terminology:—
Ectosarc = (Periplast).

Spongioplasm == (The substance of the network of the proto¬
plasm).

Cytolymph = (The substance between the meshes of the
spongioplasm).

Intra-nuclear centrosomes = (Karyosomes, Innenkorper).
Extra-nuclear centrosomes = (Blepharoplasts, micro-nude;

centrosomes, nucleoli).

Flagellum.

Undulating membranes.

In none of the trypanosomes which we have studied have we
found the slightest indication of the existence of the so-called males,
emales, and indifferent forms. We have found that the often-
asserted existence of these three types in the blood, a suggestion
originating chiefly from Schaudinn* is totally misleading.

So far as is at present known, trypanosomes are parasites
mhabltlngr ^ blood, and body ^ of , grea, ^

Hitherto no non-paras, tic forms have been discovered. As is the
case w,th other parasites of this description, their life histories appear

to anoth h iT Ified l° 5feCUre their ,ransf™ from one □
tissues of “ t WayS' WHen introdu“d into the blood and

the noxious "nfl mU't'PI>' br *** until either

Of some Strain! Tr ! .mfeCti°n deStro^ host, as m the case
runs a different °i ’ gambtense introduced into rats, or the infection

” *> t *

and then falls so thi i ft !” 1C b °od nses to a first maximum,

reappear, and reach firSt ne^ative Phase, the parasites

the infection follows ^n^r^uUrTo ^ T °u In
understood from the diagram given on " “ *

Liverpool -na.fer

see Thomas and BreinI,

445

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Reproduction of the latent bodies and the development of the
Trypanosome therefrom.

It is a very remarkable fact that in some examples of the latter
type of infection the blood during the negative phases, although
apparently containing no trypanosomes at all, and even if it be
properly filtered, is still capable of infecting other animals into which
it may be introduced. These observations suggest the existence of
another stage in the life history of the organism in the same host
during such periods.

According to Schaudinn,* 7 rypanosoma noctiue which infests the
blood of owls in certain parts of Southern Europe is transmitted from
owl to owl by mosquitoes. When withdrawn from the blood of the
owl into the body of the mosquito the trypanosomes pass through a
definite sexual stage, and are again reintroduced through successive
bites, m another stage of their life cycle into the owls once more.

It is very desirable that these observations should be repeated.
F hey have never been properly confirmed ; but, assuming them to be
correct, it by no means necessarily follows that a similar life cycle is

pursued in the case of other trypanosomes with which we are now
acquainted.

There are, indeed, numerous indications which we shall have to
consider in more detail later, that render it probable that a life cycle
such as that described by Schaudinn, involving successive hosts

X? 5eX,UaI ^ aS6XUal Stages altemate’ is th^ exception rather
than the rule. Consider in this connection the following facts-

2VTTOSOmeS/ rm6i‘nSe’ T ■ brucei- T ■ T. efU'Ptr-

bvXle ? °therS’ Can be transferred animal to animal

effe ”d him00 n ° 0b'°0d S™P>e i-oculation of blood can be

of sleeping "l t * app6ars to be 80 effected in the case

hL hXto b ; °Ugh Gl0SS’”a but no sexual stage

Ztt. d;S“Vered “ thiS ^ The observations of Brucrf

with those of Dutto^Tod^and^5 d‘SapPearS after 48 hours, together

of experiments with’ regard to t“lnBton’:|:who made a large number

seem to indicate that the rnfe f 'S matter °n the Upper Congo, all
- e at the infection by these flies is perhaps rather in

# n i ~ - - -

* Schaudinn.

und Sp,ro-

TduS T° dt *9°3 POrtS °f the Ro^ Society. Sleeping Sickness

Tr°P- Med. and Parasit^oTT^o. transrnission experiments. Ann.

447

the nature of an accident than a necessary process involved in the
normal life cycle of the parasites. Some such conclusion is borne out
by other facts in relation to trypanosome infection. Thus in the case
of Dourine,* simple inoculation of blood will transmit the disease, but
it is habitually communicated amongst horses in quite a different
manner, namely by coitus. Consequently, if there is a sexual stage
in the life history of Trypanosoma equiperdum, this sexual stage must
occur normally in the body of the horse. Further strains of trypano¬
somes, such as those of sleeping sickness and Dourine , may be kept for
years in our laboratories through inoculation from animal to animal.
In fact, such strains may be continued in this way for a quite
indefinite period, a process involving an endless number of
generations in the blood, and it consequently follows that if in such
forms the sexual stage occurs only in some other host, this phase can
be dispensed with for an altogether indefinite period.

As a matter of fact, there are yet other observations bearing upon
Schaudinn’s researches, which if they do not necessarily render his
account of the sexual act improbable, seem to clearly indicate that it
may exist in the instance of T rypanosoma nod nee as a very unusual
exception, an exception which may be incapable of throwing any
general light upon the life history of the great group of organisms to
which Trypanosoma nocture belongs. We may refer also particularly
to the author’s account of what he regards as the reduction process.

This, according to Schaudinn.t amounts to a sexual determination,
or differentiation, accomplished through a nuclear division. That is
to say, there occurs in T rypanosoma noctuce a division (heteropolar
mitosis, Schaudinn), which separates the female moiety of an
hermaphrodite nucleus from the male. In other words, Schaudinn
resuscitates (although he does not appear to allude to this fact)
Balfour’s and Minot’s view of the formation of the polar bodies, and
the extrusion of the so-called residual corpuscle during the iormation
of the spermatozoa.

* Rabinowitsch and Kempner. Centralblat fur Bakt. Bd. xxxx lJ7’ mo6

See also Minchin, Gray and Tulloch. Pro. Roy. Soc., London. \oI. 7 » 9 •
See further Laveran and Mesnil. Trypanosomes et trypanosomia,es. a ,
1904.

+ Schaudinn: Neuere Forschungen fiber die Befruehtung bei
erhandlungen der deutschen zoologischen Gesellschaft auf de - • J

sammlung. Leipzig, 1905.

44«

It is necessary to be quite clear about this matter. The
hypothesis respecting- the function of the polar bodies, and so-called
residual corpuscle of the spermatozoa, as the means by which the
opposite “sex-stuff” is got rid of from the ovum, and the
spermatozoon, has for various reasons collapsed some years ago. In
the first place, the polar bodies cannot be homologised in any way
with the residual vesicle. In the second place, it has been clearly
demonstrated that in the vast majority of animals and plants the
sexual reproductive cells, ova, and spermatozoa are precisely
equivalent so far as their nuclei are concerned. * Reduction as now
understood in animals and plants is not sex differentiation, but a
process which results simply in the halving of the number of
chromosomes in those cells which are destined to conjugate.

A great deal of confusion has been produced by Schaudinn’s
inaccurate use of the term reduction, a term which in general biology
has long had a limited and a definite meaning. In dealing with these
matters it must, therefore, be clearly understood that by reduction
chaudmn means sex differentiation, and that the term reduction in
general biology does not mean sex differentiation, and stands for
something quite different.

the morphology and life cycle of trypanosoma
GAMBIENSE

6iZZPanOSOma gamiiense (Dutt°n) is a Parasi* associated with th
Sickness TnngK “ EqUat°nal ^ and known as Sleep,,
the anTmals ““ * transfered simple inoculation into nearly al
may ru“a vfr 7Sed ““"“S' experiments. The infel
inoculation “ ThuS the strai" “-d Liverpool fo,

few days, the rat’s bW,H 5‘mP'y “““* within a Period °f J

dies from the effect 0f the"S, ‘eem'ne 'Vlth Para5i‘es and the animal
as in man the u c invasion- In other cases among the rats,
— bJood rises and falls in a

soPwelldiy observations^ upoi/yrfii V£s.t, literature of this subject. The

it clear tlwTu as to render it unintellipihi” h l*16 Sbove statements are based, being
commnnh (!hlprocess he describes ff ®,why Schaudinn did not himself make
for the Wth the reduction process as <£>V reduction, can have nothing m

the id 't 1° years- It is equally and Studied in animals and Plan'5

among animal^6 \he Process he discussed^nd ,^hy thls author did not point ont
§ mals and plants by Balfour ant^Minof that erroneously supposed to exist

449

450

somewhat irregular series of well-marked periods, the kind of
oscillation produced being indicated in the chart given on page 44).
When an animal has been previously infected, it has been found that
even at a period when no parasites can be detected in the blood, the
blood is nevertheless capable of infecting other animals by sub¬
inoculation.

As is already well known, Trypanosoma gambicnse can be
transmitted from animal to animal by the bites of flies; but the
observations of Bruce* and others have shown that if more than
48 hours elapse after the flies have fed on an infected animal
subsequent bites produce no infection. The observations of Dutton,
1 odd and Haningtonf made on the Congo, show further that it is
often extremely difficult to infect at all with flies, and the authors sum
up the position in respect to this matter in the following paragraph

“ ,Te'eheVe eithCr’ ^ That something is wrong in the way m
«mlCtG/°*S*na *al*alis has ^en used in these experiments' or,

„ W l hat Trypanosoma gambtense can be conveyed by some other
means than by it.” '

o far, then from it being established that Sleeping Sickness is
orma y spread among the African population by the bites of

thfT*' aI°ne' U W°U,d hSem that the most «*ent work on

n he nit "f**" ^ P°SSib'> the Action through fl.es is

Smkness s 6 °J "" T ' “d tha‘ the means ^ which Sleeping

mtermr ha tS;m?eamanner ‘n W’lich " d°“ *PrL in the Afiean
interior, has yet to be discovered.

ryPr0Smna in „u Blood of an InfccUi Ann

vary i “ “l^^T T" " tad

represented in figs. ,B/'. ,, fr°m f°rms smalIer than 1,10
gradually to the f j ’’ we lliay select a series increasii

r™ •*— « - *,»

FWd, ,, does net ^ ,b)° to d« “ ''

trimorphism, corresponding to the s ^ dun0rphlS”’' ‘

b tile so-called male, female, an

* Bruce, loc. cit.

t Dutton, Todd and Hani„g,on, loc. cit

451

indifferent forms described by Schaudinn * Minchin.t and many other
authors. The present examination of the forms appearing in the
blood leads us to believe that there is to be found among these
trypanosomes a series extending from those which are relatively
small, to those which are relatively immense. T he three forms often
described and alluded to as distinct, consequently appear to be
arbitrarily chosen examples in a continuous series of dimensions.

Multiplication tn the Blood

From the time when the parasites appear in the blood of an
infected animal until their numbers reach any particular maximum,
rapid multiplication takes place by longitudinal fission of the
individual trypanosomes, the multiplication being most rapid near the
successive periods of maximum numbers. When the parasites are not
dividing, they present the appearance represented in figs, i and 4.

The nucleus is nearly in the middle of the long axis of the cell,
and consists of an outer stainable mass, enclosing generally a lighter
central area, within which there lies a small, sharply-definable body,
which stains red in contrast to the purple colour of the outer mass
when subject to Breinl’s stain. This central structure forms the
intra-nuclear centrosome (karyosome). At the broad end of the cell
there exists another staining granule which, when the cell is not
dividing, remains single, the granule in question forming the extra
nuclear centrosome (blepharoplast, micro-nucleus, centrosome). It
stains, under the above conditions, like the intra-nuclear centrosome.
Arising directly from the extra-nuclear centrosome, there extends a
delicate thread, which stains more faintly, but in the same manner as
the centrosomes. It is enclosed in a thin expansion of the
ectosarc running along the entire length of the cell. The thread
projects at the narrow end of the cell as a long stained whip-lash.
This thread forms the so-called flagellum, and the ectosarca

expansion the undulating membrane.

The first sign of an approaching fission is generally apparent m
relation to the extra-nuclear centrosome. From this there buds ou
small fragment, figs. 1, 2, and 3, which may become flattened, as m

* Loc. cit.

t I’ro. Roy. Soc., 1907.

452

figs, i and /, and can be seen to be attached to the original extra-
nuclear centrosome by faint unstaining strands, fig. At a late
period there grows out from the new extra-nuclear centrosome a
young flagellum, which gradually extends parallel with the old
flagellum, as in figs. 2, 3, 6.

At a period of the fission roughly corresponding to the above, it
can be observed that the intra-nuclear centrosome has also divided in
the manner represented in figs. 2, 3, and that the staining outer portion
o the nucleus has gradually collected around, and beyond, the
dividing portions of the intra-nuclear centrosome. In this form of
nuclear division characteristic of Trypanosoma gambiense, no chromo¬
somes are formed at all, and the intra-nuclear centrosome behaves
precise y as the so-called karyosome, or centrosome, during the
division of the nucleus in Eug/ena, Eimeria schubergi (Schaudinn),
and other protozoa. At a later stage the division of the intra-nudear
centrosome becomes complete, and the outer staining portion of the
nucleus collects round the two intra-nuclear centrosomes as in

nurlJ’h’ 8’ f0rmmg t0gfether with ^^se two bodies two daughter
nuclei having JUst the same appearance as the parent nucleus. For

is thus n" nr5*6 nUdear divisi°n of trypanosoma gambimt

the presence f ' * “ * °f amitosis somewhat complicated by

the presence of an intra-nuclear centrosome *

the enxttehnesiraaHder the Pro«^ °f drv.sion proceeds b,

represented in %f °,T ° new flagehtim through stages such as

flagella are produced each encL t •” ‘hat ‘W°. aPParentl>' s‘mila'
They form separate A 1 ^ a° exPansi°n of the ectosarc

of the animal's be 1 “dU.atmg membranes and stretch from one end

flagella become eventually' ^ 7 ^ extremities of the

from the narrow , y parate’ and the animal gradually splits

daughter forn^ finally^ separatinp-^16 ^ ^ I5’ * *

represented in figs 1 -> S' PaSS through stages such as those

trypanosomes, each exactly reslih^ ^ ^ eventuall*v tVV°

During the growth and division ofl Parem f°rm' bU‘ Smafc
frequently possible tr, ^ 1 °* * le Parasites in the blood, it is

y Possible to find large numbers of cells in all stages of

deSely^iatedfn6 Cent™esm?rf altat° ”1 be n°ted ,hat in many of the

d to amitosis when this fo-r,/ t extra-nuclear, these bodies may be
0 m of division occurs.

453

division, and rest, wherein there exist, especially towards the narrow
end of the cell, quantities of granules that stain under the action of
Breinl’s stain an intense red. The colour of these granules is quite
distinct from that of either the intra-nuclear centrosome or the extra-
nuclear centrosome, and they cannot be stained with any satisfaction
at all by means of the iron haematoxylin method. 1 hey are quite
irregular in size and number, and also in their appearance, that is to
say, they may appear in all the trypanosomes in all their stages at one
period, and not at another. From these circumstances, we are
inclined to regard these granules as of metabolic origin, and we can
find no evidence that they arise from the nucleus at any time.

These bodies should, however, not be confounded with the minute
granules always seen, more or less throughout the spongioplasmic
network of the entire body. These latter may, and very often do, stain
in the same manner as the intra- and extra-nuclear centrosomes, but
we have been unable to find any indications as to their origin, or that
they any more than the large metabolic granules have any special
relationship with the nucleus.

It is possible, indeed probable, that some of these granules may
correspond to the vegetative and trophic chromidia observed in
Rhizopods by Schaudinn,* and by Hertwigt in Actinospheerium
eichhorni, but owing to the very different methods we have used, we
are not in a position to make any definite statement with respect to
this matter.

At a late stage of division, such as that represented in figs. 14 and
>3. the appearance of the organisms at first sight very much suggests
an act of conjugation, but in all such cases that we have examined we
have found no indication in relation to the nuclei, or the centrosomes

which would suggest a conjugational act.

Along with the regular method of division just described certain
modifications are frequently observed, which, although producing the
most striking appearance (see figs. 8, 9, 10) are nevertheless

apparently always capable of being explained through a disparity in
the stage simultaneously reached by different parts of the cell. Thus
the nucleus may divide completely and then divide again so as to form

P- 3*3-

* Schaudinn. Arbeiten aus dem kaiserlichen Gesundheitsamte, Vol. XIX, i9°3-
tHertwig. Archiv. fur Protistenkunde, Vol. i. ~ TT1

aee also Mesnil. Chroniidies- et questions connexes. Bull. last., lome 11 ,

454

four nuclei, without, however, the extra-nuclear centrosome having
divided more than once. Or the extra -nuclear centrosome may
divide and form three or four flagella without the nucleus having
divided at all. When these unusu.il methods are adopted, the
gigantic trypanosomes figured on Plate 1 are frequently produced.

Changes in the Trypanosomes relative to the Stage of Injection

The appearances described in the preceding paragraphs are those
which are encountered among the parasites during multiplication after
their first appearance in the blood of the infected animal, h
however, other stages of the infection be studied there are to be
found different morphological appearances among the parasites, which
are apparently of the greatest importance. In the case of an amm
which has become infected with Trypanosoma gambiense, and sho*»
a well-marked oscillation in the course ol the disease, it is found t a
as the number of the parasites is rising in the blood — -that *s t0 ^
along the ascending portions of the curve in the diagram

(page 4 i!iai

the phenomena presented are the same as those found in the inu^
stages of the infection which we have just described. If, h°'ve'er^
study preparations made at or near the maxima of the curve^
changes are found to be taking place in the trypanosomes
those of simple division. ^ ^

At the time the curve approaches any maximum, there are ^
found trypanosomes which present the appearance shown m & ^
to 20. From this figure it will be seen that such trypaH0^10^ 3
distinct from those which have been previously described, in
relatively-thick stainable band proceeds from the extra-nuc*ear jr0ll
some. This thick band is found to be most readily stainable
haematoxylin ; it is less readily, but still stainable by the ' tra-
aniline colours which we have employed. It grows from the ci5e
nuclear centrosome not along the surface of the animal as in 1
of the new flagella, but down the interior of the cell to**r ^
nucleus (see figs. 15, i6).v This stainable band, which apP^

* It is probable that the band we here refer to is the same a* tb d- ^
described by Prowazek Studien uber Saugeth.er trypanosomen. .\rb- 3 o0 cegft,

C’ Band XXIIA *905. and also Miss Robinson, Vol- N

blood-inhabiting protozoa. Proceedings of the Royal Society,

455

the periods of the greatest number of parasites in the blood, is fully
twice as thick as the flagellum. As it increases in length it may
reach, or even pass the nucleus ; or it may become coiled upon itself,
as in figs. 16, 17, 18. But whatever form it takes at first, the later
appearance suggests that the band eventually becomes in one way or
another definitely connected up with the nucleus. This suggestion is
strengthened by the subsequent behaviour* of the band, for it is seen
eventually to become gradually less stainable, to break up into a
string of fragments, and finally to disappear. Through all these
later stages of its existence it is most certainly suggested, as in
figs. 19, 20, that it is directly related to the nucleus, that is to say, to
have been, or to be actually connected up with it.

We appear then, at or near the maximum number of the parasites
in the blood, to have direct evidence of some sort of interaction
taking place between the extra-nuclear centrosome and the nucleus.
This phenomenon occurs only among animals in which no evidence of
nuclear division, or cell division, is apparent. In such cases, we do
not find that either the extra-nuclear centrosome or the nucleus is
divided, and throughout the whole development of the stainable band
the nucleus remains in a condition of complete repose.

If now we examine the portions of the curve of infection where the
trypanosomes are decreasing in number, still other phenomena make
their appearance. The numbers of the parasites gradually decrease
in the peripheral blood, and at this time, in such organs as the lungs,
the spleen, and the bone marrow, we find curious changes to be taking
place in vast numbers of the trypanosomes encountered in these
situations. Parasites showing the present changes are rarely found
tn the organs named above during the period when the number of the
Parasites is increasing in the blood, but at the time when the numbers
are beginning to decrease we see in the lungs that numbers of
trypanosomes show alterations in their nuclei. T he protoplasm of
the animal’s body becomes detached from the periphery of the nucleus
" hich lies in a clear space, while the nucleus itself contracts somewhat,
and at the same time a large clear vesicle gradually grows up in
connection with it, in the manner represented in figs. 22, 23. Round
the outside of this vesicle and the nucleus there may be seen a layer
01 protoplasm enclosing in a delicate sheath both nucleus and vesicle
together. When this stage has been reached, the rest of the cell body

456

rapidly disintegrates, the outline of the cell becomes lost, and
flagellum together with the extra-nuclear ccntrosome i. i\
frequently seen detached, and lying loose among the cells :
organs examined (see fig. 26).

These phenomena, as we have said, are encountered in the lun.
but they are also found in the bone marrow and the spleen. Asa:
the above stage has been reached the nuclear bodies and the
disintegrating remains of the parent cells disappear to a large extra:
from the lungs, the nuclear bodies being subsequently found in v<-:.
large numbers both in the spleen and the bone marrow.

At first sight, the appearance we have ju>t described might o
supposed to be simply a phase in the disintegration of the parasites,
and this was, as a matter of fact, the view winch wc were at fiat
inclined to adopt. I here are, however, reasons for thinking that
although the greater part of the protoplasm forming the bodies ot ti.<
parasites undoubtedly does disappear, the peculiar nuclear structures
we have just been considering do not follow the same course. Aft
their detachment from the rest of the disintegrating cell body, the
nuclear corpuscles become impacted in great numbers in the spier:
and bone marrow. Here, instead of disap, in the course of a

few hours, they may remain intact for as | ten days 01

m act, throughout the whole of the negative or latent period of the
infection, when no parasites are present in the peripheral Uood

toterT/1 *Ti°rmati0n’ these pccui: - • Which we propose

although they thus rema.n intact for a relatively Jong peri* it

af exPcriment., Vol. 18 No , T 1 hu»- Rode‘ “d V*IH

inniar to the latent bodies of 4* descr,be bodies mrh;

Bradford^ (C) 'h' ?*** ™ ■» .

Bradford (Quar. Journ. Micro Sri v, 1 ' "'*te form* Phmioer

m the spleen, &c., during Nap ana ini* - ****** also to have found tbu

similar bnm plas™odial production LaJer tl'Scr,be ,hem apparently tb

s.nnlar bod.es as ‘‘formes dmvoluhon ” and ' -scribi^apjOfU

»i£PTnV°*have seen them in the blood 8T Vet Sc. Vol j

he?..?™ .-TtaSS irs

Xo. 1

indicium, and to reg ^ them “ blood °f caiUe in

dHetSethF?ourrnal of

a^e^ISSi0LH-PreCisel>r V : ' '

details of r® b?d,es as true portion ; e-. •'» th.- : . - • - - -• <

The latent bofd,Vlatl0,V0f the lateEt bodies in th'
correspond to m uWe have dealt with n r * pU{ ’ -

espond to the bodies figured by Soiazek Vn^c '^ P«Uly *

457

was still possible that they might eventually simply disappear
altogether, the nuclear constituents of the trypanosome body being
perhaps more resistant than the rest of the protoplasmic structure of
the celL

In order to throw some light upon this matter, we examined a
number of infected animals which had been treated with atoxyl, this
substance having the effect of destroying the parasites in the blood
in the course of a very few hours at the most. When the blood of
such animals was treated by injection of atoxyl and examined during
the time when the parasites were still increasing in number, it was
found that a large percentage of the trypanosomes could be observed
dead among the corpuscles of the blood, but during the rapid
disintegration which follows nothing comparable to the formation of
the latent bodies is encountered. On the contrary, the nuclei in these
instances are among the first of the organs to be affected by a general
disintegration, which rapidly produces masses of debris, wherein it is
only just possible to recognise from their shapes the remains of
trypanosome cells (see fig. 41).

The same appearance is produced through the disintegration and
death of the trypanosomes in the blood of an animal which has been
killed by the disease.

From all this, it would appear that the latent bodies are not
produced during the ordinary course of cell disintegration, and must
be considered from some other point of view.

At this point it is, however, necessary to explain that when animals
are injected with atoxyl at a time when the trypanosomes are
decreasing in numbers in the blood, the disintegration does not
necessarily overtake all the trypanosomes present. It is found, in
fact, that a certain number of trypanosomes under such conditions do
not succumb to the effects of the drug, but round themselves up and
become encysted (see figs. 37, 40). These cysts are, however, very
much larger than the latent bodies. They appear to be true resistant
forms produced directly in response to the drug, and are not in any
way comparable to the latent bodies we have just described.

With regard to the latent bodies which are produced at the
maxima of an oscillating infection (structures which at first sight
might, and probably would, be taken for disintegration products),
these are, as we have seen, eventually collected m the spleen and

45«

the bone marrow, and do not necessarily degenerate there. They
persist in such situations in very large numbers, and each consists of
a flattened nucleus with an intra-nuclear centrosome. There is also
a vesicle attached to the nucleus, and the whole nuclear apparatus is
surmounted by a thin film of protoplasm, figs. 27, 28.

At the periods when there are no trypanosomes to be found in the
blood, these peculiar latent bodies are all the evidence of the existence
of the parasites in an infected animal to be detected microscopically.

At the period of the infection when a few parasites begin to
reappear in the blood, it is possible to still find numbers of latent
bodies in the spleen, and in the bone marrow, wherein the intra¬
nuclear centrosome has divided into two, fig. 28.

Again, at this period it is possible to find forms in which one-half
of the dividing intra-nuclear centrosome has passed out of the nucleus,
fig- 28, forming an extra-nuclear centrosome. Still further, at a
later period, we find forms in which a short flagellum has grown from
e extra nuclear centrosome, and these bodies subsequently appear
gradually transfoim themselves into small trypanosomes in the
manner represented in figs. 28 to 32.

As the latent bodies are gradually transformed into small trypano-
somes, the number of these bodies in the spleen and bone marrow
diminishes, but it appears to be the case that a proportion of what are
apparently latent bodies never really develop into trypanosomes and

thT^hT t0gether’ °r’ in other w°rds. ^at only a proportion of

sumv I" ar£ Under Clrcumstances above described capable of

too com ' rnegatlVe Peri°d' ^ °nCe ”°re f°™ing themselves
into complete trypanosomes.

havJhIthert?bS Wh‘CM WB haVe n°W deSCnbed are aU those which we
of the infection Tn" rats6 ^ “ reIatlon to the succeeding stages

encountered dlring" d^erent ^ Changes might have bee"

although we have had the ■ Pen°ds °f the day ^ night, but
tion continuouslv at nil p.ai^sites ln various animals under observa-
several days L °f ** CUne of Action throughout

discernible. Howevw.1t was 7 'Tu™ nocturnal . aIteration was

number of parasite* 1 ■*’ °Und t^lat t^le rapid diminution of the

So fa T he at Tanably t0°k Place between 2 and 5 ,,

the above observations upon the life cycle of the

459

parasite of sleeping sickness have been carried, they appear thus to
indicate a complete cycle in the blood of a single host, and the stages
of such a life cycle can be semidiagrammatically represented in the
manner given on page 445.

(C) THE MORPHOLOGY AND THE MULTIPLICATION IN THE

BLOOD OF TRYPANOSOMA BRUCEI

The appearance of Trypanosoma brucei in the blood is
represented in figs. 42, 43, 44, 45, 46- 1 he chief morphological

distinctions which the parasite of the disease Nagana presents when
compared with Trypanosoma gambiense are found in relation to the
distribution of the nuclear substance, and the characteis of the
extra-nuclear centrosomes.

The division of this trypanosome in the blood is longitudinal, as in
the case of Trypanosoma gambiense. 1 he nucleus divides
amitotically. The division being first marked by a lengthening of
the extra-nuclear centrosome until this body finally separates, and to
two minute beads. At the same time the nuclear substance also
elongates until we observe forms such as those represented in
%s. 43, 44, 45, 46.

As in Trypanosoma gambiense , the stages of the division o ie
extra-nuclear centrosome and that of the nucleus may not be the
same, at any particular time, and through this circumstance we observe
the same sort of multiple, and abnormal forms as in the case of
Trypanosoma gambiense.

(D) THE MORPHOLOGY AND THE MULTIPLICATION IN THE

BLOOD OF TRYPANOSOMA EQUINUM

In the blood, Trypanosoma equinum possesses much the same
shape as that of either Trypanosoma gambiense 01 Trypanosoma
brucei-, the nucleus is, however, usually placed neater to the b10^
end of the cell. The extra-nuclear centrosome is large, and t e
nuclear division which takes place during the fission of the ce s
possesses points of much interest, since the centrosomes aie moie
conspicuous in Trypanosoma equinum than in many other orm
have examined. The changes which take place in the intra nuc
centrosome during the division of the nucleus can be stu
great clearness.

460

When the nucleus is at rest the intra-nuclear centrosome is
surrounded by a light space, which is in turn enclosed by the stainable
nuclear substance (fig. 47). During division the intra nuclear centro¬
some divides, as in figs. 48 to 52. The nuclear substance becoming
at the same time collected up in the region of the dividing intra¬
nuclear beads. As this process continues, the nuclear substance
eventually forms itself into two cup-shaped masses situated
around, and beyond the intra-nuclear centrosomes. Owing
to this, when Trypanosoma eguinum is dried the edges of
these cups becoming flattened down produce at least two irregular
bands on each side of the intra-nuclear centrosomes, which under
these circumstances may suggest the existence of nuclear chromo¬
somes. This appearance is, however, misleading. The nuclear
division of Trypanosoma eguinum being, as can be seen from figs.
47 to 53, really amitotic, as in the case of Trypanosoma gambiense
and Trypanosoma brucei.

(E) CONSIDERATION OF THE FOREGOING OBSERVATIONS

For the sake of convenience, it is desirable to consider the latent
forms as the starting point in the life cycle of Trypanosoma
gambiense. These bodies, which in an ordinary fluctuating infection
may remain unchanged for long periods in the organs (and, to a less
extent, in the blood), consist at first of a nucleus containing an intra-
ce iular centrosome. The nucleus is related to a vesicle, and the
w o e nuclear apparatus is surrounded by a delicate film of proto-
p m. t a later stage, the intra-nuclear centrosome divides into
two, and one of these bodies passes out of the nucleus into the outer
layer of protoplasm, which gradually increases in extent.

centro,6 abOVe PrOCess results ” the formation of an extra-nuclear

with th°me t hC eXtra"nudear centrosome, and nucleus together

Itinct set f T centrosome, henceforth form two entirely

senes If d StrUCtUreS' which re™,n distinct through a very long
series ot divisions, as represented in j- ,

the letter B. P d in the diagram on page 445, under

461

period at which the black line is formed. At this period the extra-
nuclear centrosome develops a bridge, as it were, and connects itself
for the time with the nucleus. It may be assumed that during this
period some substance from the extra-nuclear centrosome passes into
the nucleus. Anyhow, after this has taken place the remains of the
extra-nuclear centrosome are very shortly cast away, together with the
greater part of the protoplasm forming the rest of the cell, and the
old flagellum.

Thus, if we consider the nuclear apparatus in the latent body as a
whole, this would seem to be divided into two parts during the
development of the trypanosome. After the formation of the cell is
complete, these two structures, the nucleus and the intra-nuclear
centrosome, remain in the same state, and multiply independently
into similarly distinct bodies contained in the cells produced by all
the longitudinal fissions. In other words, there arises from a nucleus
A, two new structures, B and C, both of which differ from A. B and
C multiply independently as the animals divide, but at a subsequent
stage a portion of each B unites again with the C in all the cells, and
the condition of the organism immediately reverts to A once more.

We have thus, after the formation of the latent bodies,
an unequal division of the nuclear apparatus of the latent body,
so as to form two different sets of structures, the nucleus with one
centrosome, and the other centrosome by itself. Each of these then
multiplies indefinitely in number. In individual cells these structures
subsequently unite temporarily, and later the nucleus characteristic of
the latent body is produced once again. In other words, dissimilar
structures are formed from a nucleus by division, both derivatives
multiply by division, and after a time unite in pairs, and the first type
of structure is again produced. There is in this process, when
contemplated from the present standpoint, an obvious similarity to
the two forms of sexual elements in the higher animals and plants , to
two sorts of gametic nuclei, or to a sexual dimorphism. A
dimorphism in the trypanosomes which is in like manner followed by
a reunion, or conjugation between the dissimilar elements, and
succeeded by a reversion to the conditions obtaining before the
dimorphism was produced.

The procedure in the case of the trypanosome nuclear apparatus
differs, however, from that of apparently all other known organisms
DD

462

where the phenomena of sex are discernible, in that the dimorphic
products into which the nuclear apparatus of the latent forms
separate remain contained within the same animal during its
successive fissiparous generation. With the exception of this
difference, however, the phenomena observed are certainly com
parable to the production of sexual gametes and their conjugation
In the forms with which we have been hitherto familiar, the retention
of the nuclear apparatus related to both sexes in one cell, may thus
be nothing more or less than a morphological curiosity, and in no way
necessarily suggests that the process in the trypanosomes we have
been considering is fundamentally different from an ordinary sexual
differentiation.*

Assuming the phenomena with which we have just been concerned
to be of the nature of a sexual process, still another view could
e with regard to them. It may be suggested that the
metamorphoses connected with the appearance of the black line .s an

1 . on P;lrt trypanosomes to become sexually

ifferentiated, but this attempt is not completed, the cells reverting
to their primary condition, in whtch case the process could be
regarded as an example of a special form of parthenogenesis. I»
lation to the above suggestion, it should be noted that, at the time
of he formation of the black line, the whole of the extra-nuclear

lo«Z°7^ DOt reunite With the nucleus' but only 0*
“ of the extra-nuclear centrosome does so. One moiety of Ik

portions yICTtr0050me iS detadled from the °*er. and one of to

forming £ tr^ano ^ ** ^

fronT^1 mTrat Ft ^ Separat'°n of an extra-nuclear centrosome
mtra-nuclear ce t Centr°S°me through *e division of an original

cycle, the nucleus enTeTagam ' ^ ^ B"‘ 3t ™d °f *

centrosome, yet it only doe ,connecOon with the extra-nuclear

On the other hi it may £ T* * 3 ^ ^ ^

- - - - ^ that the extra-nuclear centrosome

nucleu1101^ ^entro°onie Glides” and"?, i*1”*!®™ reduction ir> this process. The
of the'8’ Both nucleus and extra-nuclear ^ & f of this bod-v passes out of the

it re nnt Ceed[ng Missions. Afterwards entrosorne then divide in the production
hat dnr CS Vth the Helens, andThe rest of «*tra-"ucIear centrosome or a part ot

to tii&n ;eJ,li«tion a centrosome is °/t^eKCe11 b°d* disaPPears- b is kn0"T
extent the process of fertilization issiinila0' t^th'0 th<? eIement:

463

divides normally, and that it is the half so produced which enters into
connection with the nucleus during the formation of the black line.

In this case, it may be that the nucleus receives once more a
morphologically complete extra-nuclear centrosome produced in the
ordinary way by division. Our observations are not conclusive with
regard to these matters.

Considering the whole question from another point of view, it
should be remembered that it is only among a percentage of the
trypanosomes present during the periods of the maximum numbers of
the parasites in the blood which can be lounrl at any time to exhibit
the formation of the black line, and we have to assume either that all
the trypanosomes which ultimately form the latent bodies have
passed through this metamorphosis, or that only some of them have
done so.

We have no conclusive evidence upon this question. If, however,
it should eventually be shown that only a fraction of the latent bodies
are produced from trypanosomes which have passed through the
black line metamorphosis, then it will become clear that there are
two series of latent bodies, one class produced after the black line
formation, and the other without this taking place. Should this be
the case, the suggestion that the black line metamorphosis represents
a peculiar form of the sexual act, or conjugation, which we have merely
provisionally considered, will be found probably to be inaccurate.
There will be an actual dimorphism among the latent bodies, and it
will in this case be strongly suggested that the actual sexual act has
yet to be discovered, and has been overlooked. We think it would
be unprofitable to pursue this question further in the present Memoir.
It is obviously a question that can only be properly considered after
the phases of the life cycles of other trypanosomes are available.

In the introductory portion of this paper it was pointed out that
there exists a complete discrepancy between “ reduction as
apparently understood by Schaudinn* and reduction as understood
by biologists in general. Without throwing any reflection whatever
upon the correctness of Schaudinn’s observations in relation to the
phenomena exhibited in the life cycle of Trypanosoma noctua , it is
dear that, whether the process he describes exists or not, this process

Loc. cit.

464

has nothing to do with chromosome reduction as ordinarily under¬
stood. His conception is a resurrection of Balfour and Minot’s idea
regarding the function of the polar bodies in the egg, and supposed
corresponding structures in the male cells, as machinery whereby the
physical representative of the opposite sex is got rid of more or less
completely before fertilization.

In our observations upon Trypanosoma gambiense and other
forms, in which tne life cycle is possibly a complete cycle within
the body of one host (see diagram, page 449), we have encountered
nothing at all resembling the process stated to take place in the life
cycle of Trypanosoma noctuce. Prowazek* dealing with the
morphology and life cycle of Trypanosoma lewisi, maintains that the
sexual act (in the form ol ordinary conjugation) may take place either
in the body of the rat-louse, or in the blood of an infected animal —
the latter more rarely. According to this author, conjugation is
preceded by a reduction process, which he describes in the following
words : —

“ Wie bereits erwahnt wurde, findet auf den mittleren Stadien
“ der Verdauung in Mitteldarm die Reduktion der Flagellaten statt,
“ die aber nicht gleichzeitig die beiden Kerne, den Blepharoplast und
“den centralen Kern, erfasst ; bald ist der letztere schon vollig
“reduziert, wahrend der erstere erst in den Prophasen dieser
" Vorgange steht und umgekehrt.

Im centralen Kern wird vor der Reduktion zunachst das
Karyosom bedeutend deutlicher und intensiver farbbar, das

„ hromatm wird kornig doch vereinigen sich bald diese Korner zu
„ emzelnen Strangen (Taf. II, % 23, 24), die schliesslich nach Art
“ " RClfen daS inzwischen S^eilte Karyosom umgeben. Dieses
„ (Ttadm'n m°chte lch mit den Stadien der Chromosomen Paarung vor
„ fiCr ersten Tei,un§: der Metazoenspermatogenese vergleichen (Taf. II,
“ 5. ^ Pdter findet man in Kernhohlraum wiederum acht mehr

Vie 'T geu mChr°mOSOmen (Taf‘ U’ 26> Ein Stadara der

“Ande fUPPen g iSt n,Cht SGhr deutlich ausgebildet obgle.ch
..doch , ngen m d’eSem S’nne vorhanden waren (Taf. II. fig «):

-TIT r Tn Wegen der Klemheit des Objektes nichts sicheres
- e^ughch aussagen, obzwar in dem abgebildeten Fall doch

* Prowazek, lOC. cit., page 372.

4^5

" 16 Chromosomen, die durch die zwei Teilungen auf vier reduziert
" werden, gezahlt werden konnten. Deutlicher waren die Bilder bei
“ Herpelomonas. Durch die endlich effektiv gewordene Teilung des
“Karyosoms wird der ersle Reduktionskorper gebildet, der selten als
“ein dunkles, korniges Gebilde gegen das spitz.e Ende der Zelle
“abdriickt, sondern meistens dicht am Kern selbst liegen bleibt
“(Taf. II, fig. 30). Bald darauf vollzieht sich noch eine Teilung,
“durch die der zweite Reduktionskorper gebildet wird. Tn fig. 31 der
“Taf. II, bemerkt man terminalwarts rliesen Reduktionskern, der ein
“kleines Karyosom und die vier dicht verbackenen Chromosomen
“enthalt. Demnach muss der reduzierte Kern nur vier Chromosomen
"besitzen. An dem sich reduzierenden Blepharoplast kann man
“ nicht so viel Details erkennen ; zunachst teilt sich der Blepharoplast
“in zwei Teile (Taf. II, fig. 29), von denen der eine Teil durch eine
"heteropole Spindel noch einer Reduktionsteilung unterliegt. Der
'este Reduktionskorper iibernimmt manchmal die undulierende
Saumgeissel und degeneriert erst ziemlich spat. In anderen Fallen
bleibt die Geissel an dem reduzierten Blepharoplast haften (Taf. II,
“ % 28, 29).”

Prowazek describes the nuclear division taking place during the
fission of the parasites in the blood in infections with T. brucei and
T. lewisi as being mitotic, and the nuclei as containing eight chromo¬
somes. Our observations upon T. brucei and the other trypanosomes
with which we have been working, are all similar in regard to this
matter. When these animals have been properly preserved there
appear to be no chromosomes produced, and the type of nuclear
division during the fission of the animals is invariably amitotic, the
extra nuclear centrosome and the nuclear substance dividing like
drops. It is possible, of course, as in the case of the Ciliata, that the
divisions become characteristically mitotic immediately before repro¬
duction, but in none of the trypanosomes which we have examined
(one of which, T. equiperdum , certainly runs its life cycle in a single
dost) have we encountered anything of the kind.

We have had these forms under continuous observation for more
than a year, and for long periods at all hours of the day and night
In the case of Trypanosoma e.quinum , we were at first inclined tc
think that the division was of the mitotic type, but this inference was
soon found to be simply due to the defective manner in which the

466

animals had been preserved and dried, or to other forms of indifferent
fixation. In other words, there are often produced during fission of
the animals under such circumstances appearances in their nuclei due
to coagulation effects which may readily he mistaken for nuclear
chromosomes. As the fixation becomes better, in all the forms with
which we have hitherto dealt, such appearances can, however, be
clearly shown to be illusive, and the division of the nucleus during
fission to be invariably amitotic in character The same inference is
borne out by a study of the living animals.

Among the illustrations given by Minchin [Proceedings of the
Royal Society, Vol. 78, Series B, No. 20, Plate 12], figs. 4, 8, 9. and 1;
have been produced from specimens that have been dried and stained,
and suggest the existence of chromosomes, hut we are inclined to
think that these appearances are simply due to the bad fixation
methods employed, and are really quite misleading.

With regard to the nature of the nuclear division accompanying
fission of the above-mentioned trypanosomes, our results are m
complete accord with those of Laveran. Indeed, with regard to the
nuclear reduction described by Schaudinn,* and finally by Prowazek.
our observations have revealed nothing suggesting anything even
analogous to these descriptions. Prowazek gives a scries of figures
illustrating this process in Trypanosoma lewisi (PI. II, figs. 23, 24,
25). Here nuclei with eight chromosomes are said to be apparent
\\ e cannot detect in the figures themselves the slightest suggestion
of this being the case, and are inclined to think that the irregular
blotches and strands, undoubtedly correctly drawn, have nothing to
do with chromosomes, but are due to the manner in which the
specimens have been preserved. In figs. 27, 28, 29, 30 and 31, the
so-called reduction of the nucleus as well as the extra -nuclear
centrosome (blepharoplast) is represented; but in none of these
figures do we see any indication of either true chromosomes, true
mitotic division, or true reduction as ordinarily understood. In fact, as
ar as the illustrations are concerned, we are unable to find, or to see,
any indication of a reduction process.

It will thus be observed that the results we have obtained.
— *n re^a^on to Trypanosoma gambiense , but also equally

♦Generations und Wirtswechsel, &c. "

t Loc. cit.

4&7

among other forms to be described later, differ not only in degree but
also in kind from those obtained on the one hand by Schaudinn, and
on the other by Prowazek. The descriptions of Schaudinn, in so far
as they bear at all upon the present work, do so, however, through the
investigations of Trypanosoma noctuce , which since it appears to be a
form of trypanosome requiring more than one host for the completion
of its life cycle, may very likely differ in the features of this life cycle
from the more ordinary forms with which we have been concerned.
On the other hand, Prowazek’s results obtained in the case of Nagano.
(that is to say, from one of the trypanosomes considered in the present
paper), so far as the nuclear changes during fission are concerned,
differ entirely from our own ; these latter fall directly into line with
the observations we have made upon other forms, and are quite
incompatible with the description of this process given by Prowazek in
the case of T. lewisi or T. brucei. The question which now confronts
us is upon what cause this difference of results depends. We are
inclined to think that the difference of result is due to the methods
which have been employed. We may as well say here, that from
what we have gathered with respect to the different methods that
have been generally in use, it appears that all the methods involving
the drying of the blood before staining, or, in fact, any method
involving drying at all, is, so far as nuclei are concerned, absolutely
useless from a cytological point of view. Nothing relating to the
delicate mechanism of mitotic division is generally preserved in cells,
whether they belong to unicellular or multicellular organisms, when
dried and stained with Romanowsky, Giemsa, or in any other manner.
Even the resting nucleus itself under such conditions becomes a mere
caricature of the actual structure.

When treated in this way, the irregular or regular blotches and
streaks of stainable matter have nothing in common with, and do not
represent, even in a relative or equivalent sense, the structures actually
present in the cells. Anyone who wishes to verify this fact for
himself will have no trouble in doing so if, for example, he makes a
smear preparation from the testis of a rat, stains after the manner of
Romanowsky, and then compares this with a properly fixed and
stained smear, in the production of which ordinary cytological
precautions have been observed. It is a curious fact that in a rats
testes under these conditions certain cells which really contain

468

sixteen gemim or heterotype chromosomes, when subject to the
action of drying and Romanowsky, very often present (within the ill-
formed area representing the nucleus) six irregular masses of
stainable stuff resembling the so-called chromosomes of the dned
trypanosomes. Such appearances are, however, certainly due to
regularity of coagulation and shrinking during the drying of the cells,
and have nothing in common with the real morphological structures,
(the chromosomes), which either the living cells, or successfully-
preserved cells possess. In view of these circumstances, we are
inclined to regard many accounts of the existence of chromosomes,
spindles, and even the assumed existence of mitotic division among

trypanosomes, as conclusions which appear to be most questionable,
and as requiring in all cases confirmation in a variety of ways which
do not involve the violence dealt to the finer details of cell structure
by drying.

Finally, we see in the case of 7 rypanosoma gambiense that the
life history of this parasite as it lives in rats seems to be complete in
the blood of the rat, and not in any way dependent for its completion
upon the transference of the parasites into the blood of any other
kind of host. In rats the latent forms pass gradually into trypano¬
somes, these m turn divide through many generations, and their
multiplication is followed by a metamorphosis which, whether we
regard it as a special form of sexual process, as a form of patho¬
genesis or as a sexual stage, the fuller details of which have not yet
been elucidated, seems undoubtedly to stand in one of these
relationships to the normal cell multiplications preceding the
formation of latent bod.es. The stage in quest, on results In the
production of the latent bod.es once more, and the cycle is complete,

cvcl c ry, l° thiS COncePtl0" ‘hat, notwithstanding the

a isMhv Pment tt;yPanOSOma SamHe"*C- Stl" there may exist
Pr0bab'hty. of the transference of the trypanosomes

eXu“ZP 7,u°St Wheie a further metamorphosis representing the
be SO the °rgan,5ms ls P“-d through. Tins, of course, may

mstalce of a ^ ““ °f the ^ypanosome of Dour.ne a clear

r.fnt^TT1'" hiSt0ry' "hlch' ™der norma, circun.-
norma^uc:^"^ 'n‘° of host; and, under

whatever sexual stap-e ** >,OSOma e Vul perdu m must pass through
sexual stage ,t may possess, its whole life h.story m fact, in

469

the body of the horse. Dourine can, however, like sleeping sickness,
be inoculated from host to host by simple transmission of blood as well
as by coitus ; in other words, the faculty of being transmitted by
simple inoculation of blood is shared by Trypanosoma equiperdum ,
wherein no other host is usually involved, as well as by Trypanosoma
gambiense. In these circumstances, it is simply natural, assuming
flies to be the agents by which sleeping sickness is transmitted, to
admit that this form of transmission may be merely in the nature of a
mechanical transference, and have no more relation to the sexual act
in the life cycle than has the artificial withdrawal of blood from a
horse infected with Dourine. In other words, it would seem that the
transference by flies in the case of sleeping sickness may have no
more significance with respect to the life history of the parasite than
has the direct inoculation of Dourine from horse to horse by means of
a needle.

As we have already pointed out, the observations of Bruce,
Dutton, Todd and Hanington* and others seem to indicate that the
transference of sleeping sickness, when it is brought about by flies, is
in the nature of a simple inoculation of blood, while it would appear
that Dutton, Todd and Hanington incline further to believe that flies
are not necessarily the normal means by which the propagation of
sleeping sickness takes place.

They sum up the situation in this respect as follows: — “ It seems
“ that all the results are in conformity with the hypothesis that
“ Glossina pal palis transmits Trypanosoma gambiense, and that it
“ is probably not able to do so when the space between the trans-
“ mitting feeds exceeds 48 hours ; this conclusion is, nevertheless, to
" our minds a most unsatisfactory one, if we are to regard these
" Glossinae as the chief or only carriers of Trypanosoma gambiense.

It certainly seems possible that mechanical transmission
" by tsetse flies cannot alone be responsible for the rapid spread of
" keeping sickness of recent years.”

dhese questions, however, open out a wide field of enquiry, which
it is at present unprofitable further to discuss.

Loc. cit.

470

APPENDIX I

METHOD OF PREPARING AND STAINING WET FILMS, USED
DURING THE FOREGOING INVESTIGATIONS

Place a very thin layer of albumen-glycerine on a clean slide
The best method is perhaps to put a drop the size of a large pin’s head
on the slide, and to spread this with a clean duster over the slide.
On top of this layer spread a drop of blood in the usual way, and dip
the slide, while wet, into the fixing solution (Flemming’s strong
solution was usually used). Leave it for about five to ten minutes,
wash immediately in water, and pass the slide through alcohols in
consecutive order, increasing by io per cent, at a time to absolute
alcohol. Then back from absolute into 80 per cent, alcohol in which
is contained iodine and potassium iodide. In order to prepare the
solution of 80 per cent, alcohol containing iodine and potassium iodide,
make up a concentrated solution of potassium iodide in water, and
iodine in alcohol, mix them together, and add them to some So per
cent, alcohol until the mixture becomes a dark brown colour. Leave
the slide in this from five to ten minutes, and then bring it int0
30 per cent, alcohol. Use for staining either aniline safranine faach
isabes), or the following solution :-Prepare a concentrated watery
so ution of safranine (Griibler) and a concentrated alcoholic solution
o sa ranin, mix them in equal parts, and then add pure aniline oil

sixmonfT ‘i?6 l° tlme' and ieaVe tHe S°Iuti0n t0 riP“ thfeC “
Wash nff5' “ tHiS SOluti™ fOT f™> an hour to two hours.

“ 5afranin' and stain afterwards with polych«»”c
(*£££“ Ne,.6™"e methylene blue purissimum

S1 t L,“cT; d water and 5 e— ^boaate'

the stamin, ™ , t0nnpen: the °lder the solution the be#*

differentiate with o °ff ^ methylene blu& !l

comes out Brin? th T tannm> as long as the blue s a

Bnng the Shde up through ^ __ into

the

absolute alcohol Tl r P trough alcohols, as above,

slide into aniline oTl Z ^ ^ a reddlsh Now dip

tinge ; the aniline oil takes 1 1’1fddlS^ C°lour chanSes to a purpl^”
stain left by the on ' °.U &t * le same time the excess of b

Balsam under a coverstp ^ ^ Mount in

- —

47i

APPENDIX II

MODIFICATION OF H El DEN H AIN’S HAEMATOXYLIN METHOD,
USED DURING THE FOREGOING INVESTIGATIONS

Fix and treat the film in the way described in Appendix I. Clear
the slide from the alcohol containing iodine and potassium iodide,
and pass it through successive alcohols (as in Appendix I) into water.
Stain in a 3-t per cent, solution of iron alum for one hour, wash this
off, and stain with the following solution : — -5 gramme haematoxylin
dissolved in 100 c.cm. distilled water, to which, after the haematoxylin
has dissolved, a few drops of concentrated watery solution of lithium
carbonate is added. Stain for half an hour, and then differentiate in
the usual way with iron alum.

472

DESCRIPTION OF FIGURES

In all cases, unless otherwise stated, the figures have been drawn

with a long tube Zeiss, 2 mm. apo. objective and 18 or 27 eyepiece.

PLATE XXXVIII
Trypanosoma gambiensc

Figs. 1, 2, 3 from peripheral circulation stained with iron

haematoxylin.

Figs. 4 and 5 stained with Breinl’s stain.

Fig. 1. — Trypanosome in the resting condition. Nucleus single, not
dividing. Intra-nuclear centrosome single.

Fig. 2. — Trypanosome showing early stages in division of nucleus,
intra-nuclear centrosome divided ; extra-nuclear centro¬
some also divided, new flagellum growing.

Fig- 3- — Trypanosome in the same stage as preceding. Intra-nuclear
centrosome completely divided ; the new flagellum is also
seen during the course of its development.

F*ff- 4- Trypanosome is in the same condition as fig. 1, showing also
the metabolic granules in different parts of the cell.

Fig- 5- -Trypanosome wherein two undulating membranes have
been formed, showing also the metabolic granules.

Fig. 6.— Trypanosome in much the same condition as in fig. 1, but
showing early stages in development of the new flagellum.

F lg' 7 — Later stage in the division, showing the mode of division of
the intra-nuclear centrosome, amitotic fission of the rest
of the nucleus, and the duplication of the extra-nuclear
centrosome.

Fig. 8 Trypanosome showing three flagella and three nuclei in
different stages of division.

Fig. 9— The same showing three flagella and four nuclei.

iff- IO- The same showing two flagella and two nuclei.

iAtut . Trap Med. run! Pfira^itof/igy. PLal&JXXYIU

J-E.S.M. id.nat dt,

Hutn. IitV Xondon

.

474

PLATE XXXIX

Trypanosoma gambiense

Figs, ii to 14 stained with Breinl's stain.

Figs. 15 to 21 stained with the modification of Heidenhain’s

haematoxylin.

Pig. 11. Trypanosome showing division of the intra-nuclear centro-
some, and the nuclear substance. Also multiplication of
the intra-nuclear centrosome so as to form a group.

Pig. 12. Trypanosome showing mode of amitotic separation of the
nuclei. Multiplication of the intra-nuclear centrosome,
and the formation of an independent group of these bodies
In this and the preceding figures metabolic granules are
also seen.

* r3- Later stage in the fission of a trypanosome. The flagella

are being detached from one another at the thin end of the
cell. 1 he cell body is dividing from this end towards the
other. The nuclei are already divided. Metabolic
granules are scattered throughout the spongioplasm.

Fif?- l4- Later stage in the division of a trypanosome, showing the
manner in which the daughter cells separate.

Fig. 15. -Trypanosome, at one of the maximum periods of the
infection, showing a single flagellum and resting nucleus,
and also the origin of the black line from the extra-
nuclear centrosome.

Fig. 16.— A similar stage wherein the black line has reached the
neighbourhood of the nucleus.

Fig- 17.— The same.

lg' 18 ’ TryPanosorne showing the black line coiled upon itse
towards the intra-nuclear centrosome.

'&■ '9— Trypanosome showing early stages in the degeneration 0

the black line, and its later direct association with tb
nucleus.

F'S' 20~ S’m,Iar s‘age. wherein the intra-nuclear centrosome ha:
become divided.

F>g. 2J- So-called involution stage, showing resting nucleus ana
multiplication of the intra-nuclear centrosome.

oAnib. Trap Med. and Parasitology. Plate, XXXIX

JE S.M. ad nut. del.

Suti. Iatt" &»*»■

.

4 76

PLATE XL

Stages in the Metamorphosis of Trypanosoma gambiense in the Organ\

Figs. 22, 23, 24, 26, 27, 28, 29, 30, 31, 32, 34, 35, 36, stained with a
modification of Heidenhain.

Figs. 25 and 33 stained with Breinl.

Fig. 22. — Trypanosome during the decrease of the parasites in the
blood of a rat, showing alteration in the nucleus marked
by the formation of a vacuole.

Fig. 23. — The same.

Fig. 24. — One of the common forms in the lung at this period.

showing the same changes in the nucleus.

Fig. 26. Low power view of trypanosome at this period, showing
the detachment of the latent body from the parent cell
big. 27. Latent bodies. The nucleus is attached to a vacuole and
both are surrounded by a thin film of protoplasm. The
nucleus contains a single intra nuclear centrosome.

Fig. 28. Latent bodies. To the left the intra nuclear centrosome is
shown divided. To the right stages in the division of this

body and the extrusion of one daughter element from the
nucleus.

Fig. 29. Latent bodies showing origin of a new flagellum from th
mtra-nuclear centrosome.

Figs. 30, 31— Later stages in the formation of small trypanosome
from the latent bodies.

F.g- 32— Latent body from the spleen of a rat infected wit
trypanosoma brucci, showing nucleus, vacuole, and th

gin o t e flagellum from the intra nuclear centrosomi
(Compare fig. 29.)

F’g- 33 — Trypanosome showing Mri„ ♦ . .

nucleus ^ early stages in division of thi

34- —Trypanosome drawn to show thn c 1 1

of the protoplasm Schaumplasma struct™

F'gs. 34 to 36. — The

same.

a/lrui;, Trop. Medi. anxl Pa ras ii/j l o </v. Plata XL.

J E s, V.

aJ n»t \ri

HutK 'U0>- loixlo"

*

478

PLATE XLI

Trypanosoma gambiense and Trypanosoma brucei

Figs. 37-41, T. gambiense.

Figs. 42-46, T. brucei.

igs- 37- 38, 4°. 41. 42> 43* 45> 4^, Breinl’s stain.

Fig. 44, Heidenhain.

^i§f- 37- Trypanosome in the blood of a rat after treatment with
atoxyl, showing rounding up of the cell body.

Fig. 38— Further stage in this process, flagellum is still attached. A
slight modification of a membrane is apparent round the
periphery of the cell.

39— The same, flagellum not visible.

Fig. 40. Later stage in the formation of the cyst. The membrane
more apparent.

Fig. 41— Trypanosomes killed by atoxyl in the blood

'S 42' Trypanosoma brucei in resting condition, showing structure
of the nucleus and relation of the schaumplasm.

■g 43. The same, showmg the division of the extra-nuclear
centrosome.

Figs. 44 to 46.— Later stages of division.

JES M -ad ui del.

Jfutb.IdtbT London

c/huiy, Trop. MexL. auruL Parasitology. Plate XU.

—

480

PLATE XL II
Trypanosoma equinum

Figs. 47> 49- 5°. 51* 52. 53. 54. stained with Br.einl.

Tig. 48 stained with modification of Heidenhain.

T 'S'- 47- Resting stage of the trypanosome.

h ig. 48. -Stage showing formation of the new flagellum and division
of the nucleus.

Fig- 49-^ ~ J rypanosome showing details of the division of the intra¬
nuclear centrosome and the nuclear substance,
f 'Ss- 5°. 51- 1 rypanosomes showing later stages of the same process,

f 'Ss- 52* 53- Trypanosomes showing still later stages in the division
of the nucleus and the characters of the intra-nuclear
centrosomes.

'L 54- -Tiypanosome wherein the nucleus has divided into four
constituents, although there is only one flagellum.

del.

Uutk,-LitlW I. cm cl on

k

OBSERVATIONS ON THE SO-CALLED
‘CANARY FEVER’

483

OBSERVATIONS ON THE SO-CALLED
‘CANARY FEVER’

BY

C. E. WALKER

ASSISTANT DIRECTOR OK CANCER RESEARCH AND HONORARY LECTURER ON-
CYTOLOGY TO THE LIVERPOOL SCHOOL OK TROPICAL MEDICINE
IN THE UNIVERSITY OF LIVERPOOL

( Received for publication October \th , 1907)

Hitherto the evidence regarding the disease commonly known as
Canary Fever,’ has been of a most conflicting character. Very
frequently the disorder is of a comparatively trifling nature, and such
cases do not usually come to the knowledge of a medical man.
Private enquiry points to the probability that from 60 to 80 per cent,
of the visitors to the Island suffer from one or more attacks during
the time that they are in the islands. Dr. Taylor, who has practised
among the English visitors in Las Palmas for many years, estimates
that only about 5 per cent, of the cases that occur come under the
observation of a medical man.

The disease is apparently more prevalent during the winter months
than during the summer, but this is probably due to the fact
that larger quantities of food are consumed during the former period
owing to the larger number of visitors, and that as flies are then more
numerous, there is, as we shall see later, a much greater chance of the
food becoming infected.

Clinically the disease is characterised by the suddenness of its
onset, and by its generally attacking a number of individuals in the
same hotel at the same time. Frequently it commences with
vomiting, and as a rule, even when vomiting does not actually occur,
there is a considerable feeling of nausea which lasts for from a few
hours to two or three days. A few hours after the vomiting or nausea,
sometimes synchronously with it, griping pains in the abdomen o a
violent character commence. This is the most marked and constant
symptom, and causes very considerable suffering. The patient now
developes a more or less acute attack of diarrhoea. In the more acute

4«4

cases the discharge frequently becomes mucous in character, and is
sometimes blood-stained. Sometimes the temperature rises, but not,
as a rule, to any considerable extent. The length of the attack varies
in different cases ; a slight one but for two or three days, the more
serious for two or three weeks.

The disorder has been attributed to various causes. The red wine

of the country, the water, an excess of fruit, too much food, and too
little exercise have all been blamed. The most superficial examination
of the available evidence, however, shows that none of these can be
the usual cause of the disease. Individuals who are teetotalers are
as frequently attacked as those who never drink water. Those who
eat a quantity of fruit are no more subject to it than those who
abstain from fruit altogether.

Enquiries made with regard to the milk supply showed that the

dairies in the town of Las Palmas, where the cows are stall-fed, are

generally remarkably clean. 'I hese dairies do not, however, supply

enough milk to meet the demands of visitors, and a large proportion

of it is brought down from the country. The milk supply is generally

the same in the case of the hotels as it is in the case of the private

houses, which fact, as we shall see later, is of special significance.

Also I was able to ascertain that people who drank large quantities

o milk in private houses were not subject to the disease, while those

“els wh° never touched milk were subject to it.

Vith regard to the water supply, there is no evidence incriminating

’ or P^°ple who have stayed in hotels with a certain water supply

su ered from the disease, while people in private houses with a

atf!j SUpp,y derived from the same source have been free from it.

VT S1&n^cant fact which was ascertained at the commence-

cont t j C,enc*uiry was> ^at while visitors at hotels so frequently

1: • • 6 ^^sease> it was practically unknown among people

peonle J Iate k°uses' Dr. Taylor informs me that among English

in his ° 1VC m pr*vate houses the disorder is very rare, and that

coincides confined to visitors at the hotels. This

Dr. Mill”, lrC Wlth the personal evidence I was able to obtain.

practic-illv ' ^ * S° 'n^OIms me that the disease, in his experience, is
practmally confined to the hotels.

classes of t0 comParatively common among the lower

of natives, but I was able to ascertain the occurrence of only

4»5

two cases during the past three years in hotels, and in both these
there was very good evidence that the infection had occurred outside

the hotels in which the men were staying.

These facts suggested the advisability of a comparison between
the sources of food supply, and the methods of keeping and cooking
the food in hotels with those in private houses.

The meat for hotels, in some cases at any rate, is frequently frozen
meat brought to the islands by ship. It often happens that this meat
is more or less exposed on the quay for several hours— sometimes tor
more than twelve when a steamer arrives early in the night.

The larders in many hotels are not fly proof, and the food-stuff is
not adequately protected from flies before and after cooking. In some

cases it is not protected at all. . .

Soup in the hotels is frequently made in a stock-pot which is on y

emptied once or twice during the week.

In private houses the meat supply is generally fiom amnia s
in the islands. Considerable precautions are usually taken to pro e ^
all food from flies. This precaution extends not only to keeping
meat, fish, &c., in fly proof safes, but even dishes of fruit are often
covered with muslin while on the table. Soups are made les i ever)

day, and a stock-pot is not used.

In some of the butchers’ shops (in Las Palmas, for instance
frequently sees meat exposed for sale which is not protecte
flies. I have been unable to trace the destiny of this meat,
case of the private houses where I was able to trace the source ot tne
meat supply, I found that a servant was sent early in the morning
the market. This meat is stated to be freshly killed. It must a *°
be observed that the evidence with regard to the occuirence
disease which is here given applies only to the belter c ass o na n

nd not to the lower classes. . . f .1

Flies are extremely numerous, particularly during t e in
ear when most visitors are staying in the hotels. They a ways ^
0 an extent which can but rarely happen, even in iso a e
nore temperate climates. Again, the habits of the owei c ass ‘
he natives are such as give perhaps the greatest possible number
hances of infection of the food with various nucro-c*ga man*
neans of the flies. For instance, the inhabitants usually defaecaU
n any spot that suits their personal convenience,

4*6

favoured particularly by them in this way is not many yards away
from the unprotected larder window of one of the hotels. This
instance is only quoted as one among a large number.

T emperature seems also to have a very marked effect upon the
frequency of the disorder. In Las Palmas, for instance, the summer
and winter temperatures vary only by about io degrees, and the
temperature does not usually go below about 75 °F., so that any
bacteria carried on to food-stuff by flies would be under very
favourable conditions for rapid multiplication. In the Monte district
oi the same island the temperature is much more variable, and during
the winter is comparatively low. In this district cases of so-called
Canary fever are comparatively rare in the hotels.

These observations indicate that there should be a greater risk of
conveying a large dose of bacteria, and of the toxins they produce, by
means of soup from the stock-pot and of rechauffes, than by any other
kind of food. The soup is rarely if ever brought to boiling point, and
necessarily is allowed to cool every night when the kitchen fire goes
out. Entrees and other forms of rechauffes are made from meat or
is that has been previously cooked, and such material will have been
e t or from 18 to 24 hours before being served without having been
rough t up to the boiling point of water. Micro-organisms will have
een destroyed, and many of their toxic products may have been
1 n^efc>ra^ec^ by the temperature generally required to cook food for
rst time, and with such material the risk of infection or
ft 1. 1U1^ much less than in the case of the soup from the

Suffir' °,r .°[ 3 1 ecbauffe which is rarely if ever brought up to a

sufficiently high temperature. '

hotel 1 - T St l' H1 P'as ^>a^mas. most of the other visitors in the
avoided attacks °f the disorder. My two companions and I
occasion ?°UPS anc* entr^es» and also avoided infection. On one
followed w>reVer’f°nC °f my comPanions tQok an entree. An attack

disease is re e, " a 'en ,n conjunction with the fact that the

in the method oTl- !° ^ !'°tGls’ and keePing in mind the difference

-girded as corroborate evidence^ ***** ^ *° ^

being confined^ hL ^ f,'S°rder’ its epidemic character, and its
hotels, suggest very strongly that it is due to

4*7

bacterial infection of the food. The details which have been given
regarding the differences between the methods pursued in hotels and
in private houses, suggests that the infection is largely brought about
by flies carrying the bacteria on to the food, in which position the
conditions are particularly favourable as regards temperature and
nourishment for their rapid multiplication. 1 his does not necessarily
suggest any specific micro-organism, but is amply sufficient to account
for all the cases of Canary fever that occur. There is, however,
another cause which might explain some cases, which I have proved
by personal experience to be sufficient to produce all the symptoms
of the so-called fever. If two or three men live for many days upon
a small boat during the summer months in England, they will very
likely be suddenly attacked by these symptoms in an acute form unless
all their cooking utensils, plates, forks, &c„ be thoroughly cleaned and
boiled at frequent intervals. The temperature below deck and in the
lockers in a small boat in British waters will frequently be very high
during the summer, and there will be very favourable conditions or
the multiplication of bacteria in any small collections of grease, &c.,
on the cooking and other utensils.

I have been able to find no evidence that any utensils are ever
boiled in the hotels.

The following measures should be adopted in order to avoid, or
at least materially lessen, the chances of food becoming infecte

Meat and fish, particularly, should be protected from flies in
effective a manner as possible before it is brought into the hote s.

When in the hotels all food should be protected from flies ; t ie
larder should be entirely fly-proof ; the entrance should be P*ote<*®
by two doors, between which there is room for a man to stan , °

these doors should close automatically with springs, and it wou
well to have some simple automatic arrangement which would preven
one being opened until the other was closed. It should be easy
catch the few flies that might possibly get into the larder, in spite o

these precautions, by means of fly traps.

Of course the best plan would be to keep the food m a chain er
which was constantly below freezing point. When the oo w
removed, once or perhaps twice during the day, it shou e -ep

fly-proof receptacles. , ,

Meat should be kept hanging up, and not laid upon s e ves.

488

Shelves and tables in the larder, serving rooms and kitchens should
be made of some non-absorptive material, such as marble or slate.
Most of the shelves and tables upon which the food was placed
during the processes of cooking and serving which I saw in the islands
were made of soft wood. No matter how much this wood be scrubbed,
there must always be a certain amount of organic material in a more
or less advanced stage of decomposition in the cracks. In the serving
iooms, kitchens, Sic., and wherever food is exposed for any length of
time to contagion by flies, the food should be covered up as soon as
it is put down. The ordinary wire gauze dish covers are cheap, and
admirably suited to this purpose.

Cooking utensils, plates, dishes, forks, spoons, & c., should be
sterilized shortly before use. 1 his would not involve any very
considerable extra labour, and convenient apparatus would not be
very costly. No pressure of steam would be necessary, only the
utensils should be brought to the temperature of steam. Cleaning
bve steam, such as is done on ships, would be very

opper cooking utensils have the disadvantage that they require

unmg at intervals. "1 here is no means of getting this done in a

ass manner in the islands. 7 he tinning is often irregular, and

•small ^actlcab*e to get such a surface really clean and free from

would cctlons organic material. Something other than copper
would, therefore, be an advantage.

With nUSt jC made ^resb every day, and the stock-pot abolished,
guarded? T * ?*»'**• even if protection from flies is
the material C C firSt and second cooking, it would be well if

some minutes^ Cre 3 br°Ugrht to boiling point and kept so for

Thedreln?tT!!i0nS. mUSt be keP‘

even the whole 1 ° Ut bttle dlfbcu,ty in keeping the kitchen, and

gauze frames tn tv, ^ COniPara,avely free from flies by means of wire
consist of a frame & r- ‘ !ndo.ws anc* double doors : the outside door to
would allow plent f • Wlfe ^aU7e stretched upon it. Such a plan
the majority of the flie^ ^ C°me into tbe rooms, and would exclude
and even by some people inE^T* '** eXteDSively ”

° SUggeSti°n 'S intended that *e kitchens of the hotels are no,

4»9

clean in the ordinary acceptance of the word. For instance, the
kitchens of those I visited would compare very favourably with any
kitchen I have seen in Europe.

What the observations really imply is that precautions which are
sufficient in England to prevent a degree of infection by bacteria
enough to produce symptoms, are wholly inadequate under the
conditions of temperature, &c., in the lower and hotter parts of the
islands.

It is quite possible that there may be one or more specific bacteria
which are specially responsible for the acuteness of the symptoms.
Even if this be the case, however, there seems but little doubt that
the flies are to a large extent responsible for the original infection
of the food. The rapid multiplication of the bacteria and the
consequent production of toxins depends upon the local conditions.
It would seem that the suggested precautions are necessary whether
there be a specific micro-organism or not. It is probable that food is
more frequently infected, even in the best conducted private kitchens
in the towns in the islands, than is the case in Europe, and that
consequently the residents may have acquired a limited degree of
immunity. I met several residents, however, who told me that they
had suffered from attacks after dining at hotels, but not at any
other time.

The term 'Canary Fever’ is obviously' a misnomer. It would
appear probable that a bacterial infection of the food must
frequently happen, and similar epidemics be produced in other
tropical and sub-tropical countries where similar conditions prevail.

I wish to acknowledge the great assistance given to me by
Dr. Taylor and by Dr. Millares. Their local knowledge was of the
greatest value. The enquiry was much facilitated by the ready help
and great frankness with regard to the facts that had come within
his knowledge, shown to me by Mr. Sauerbrei, the manager of the
Hotel Metropole at Las Palmas.

.

CONTRIBUTION A L’ETUDE DE
POROCEPHALUS MONILIFORMIS

4

493

CONTRIBUTION A L’ ETUDE DE
POROCEPHAL US MO NIL IFORMIS

PAR

A. BRODEN

DIRF.CTEUK DU LABOR ATOl RE
ET

J. RODHAIN

M&DECIN DE L’HdPITAl. DKS NOIKS

Laboratoire de Leopoldville (Etat dit C ongo >

( Received November 13 th, 1907)

Nous avons eu recemment 1’occasion d'observer a Leopoldville,
plusieurs cas d’infection par des larves dc Porocephalus moniliformis,
un cas chez l’homme, quatre cas chez le singe.

Les infections par ce parasite doivent etre dans le bassin du
Congo comine ailleurs, extremement rares. Dans les nombreuses
autopsies d’hommes et d’animaux les plus divers que nous avons faites
depuis 1900 a Leopoldville, a Lusambo et dans l’Ubangi, nous
n’avions jamais rencontre de larves de ce parasite. Dutton et Todd
qui ont fait au Congo egalement de nombreuses autopsies, n’ont pas
signale de ces larves.

historique

Pour autant que la litterature nous est accessible ici, peu de cas
d’infection par Porocephalus moniliformis semblent avoir ete
signales. Rencontrees une premiere fois par Piunei en 1846 dans
foie de deux negres au Caire, des larves furent signalees ensuite pa
Bilharz, Fenger, Aitken, Giard, Chalmers. Tons ces cas se rapportent
a des infections chez l’homme. . ,

Nous donnons cette litterature d’apres Braun.1 Depuis 1 99.
de la publication de l’observation de Chalmers, il ne semble pas
ete signale de nouveau cas. _

* During the autopsies of thirty odd natives and some 1”°“ kJvs | and^n each

?A). we saw Porocephalus sb. four times ; twice in man, twee m
instance the parasite was firmly encapsuled in the liver. (J-

ly encapsuled

•Braun. Thierische l’arasiten des Menschen.

3 AuH. 1903-

494

D’apres les observations de ces differents Auteurs, les larves de
Porocephalus s’enkystent dans le foie, dans la muqueuse intestinale,
dans le poumon. Chalmers- en a retrouve a l’autopsie de nombreux
exemplaires libres dans la cavit6 peritoneale et dans l'intestin grele.

Dans certains cas, observations Aitken, Chalmers, le parasite
semble avoir ete la cause directe dc la mort par pneumonie ou
peritonite.

Chez l’animal les larves de Porocephalus moniliformis ont ete
rarement constatees. Pruner en a signale chez la girafe, d’autres
Auteurs chez l’hyene, le Cyrwcephalns mormon et le Cercopithecus
albogularis (Looss3).

Looss se range a l’avis de Neumann4 qui considere les lanes de
Porocephalus signalees jusqua present chez l’homme et quelques
animaux, comme la forme jeune de Porocephalus moniliformis ,
Diesing. Celle-ci se rencontre dans les poumons des grands serpents
africains.

L’evolution du parasite n’est pas connue avec certitude. D’apres
Looss, elle doit etre analogue a relic dc Linguatula rh inaria.

OBSERVATIONS PERSONNELLES

Le 1 1.V.07 est amene a 1’hopital des noirs a Leopoldville, le soldat
Bangala-Mingi, se plaignant de vives douleurs dans le flanc gauche.
L n examen superficiel denotant un etat general grave, le malade est
garde en observation.

De 1 interrogatoire et de l’examen faits par l’un de nous, nous
retenons les faits suivants :

Le soldat est arrive a Leopoldville au commencement d’avril 1907;
avec un detachement il est parti a l’interieur du district, ou il serait
devenu souffrant vers le 15 avril. A ses camarades il se serait plaint
a °rsde \ agues douleurs dans le flanc gauche et de constipation,
bic, CSt Un h°mme largement bati, a bonne musculature, a nutrition
1, , . conservee. L attention est attiree immediatement par
omen dont la moitie gauche est fortement bombee. A la

3InTb' f‘ ,®99> Bd. XXVI, p. 518.

4 PorocTph I1"?* f' Trope,lkrankhe>len, Bd. I, p. ,9S.

CC <raVai1' Sous” apate^r^^S^ici. ^ ** ^ ^

495

palpation l’abdoinen presente une resistance uniforme, assez marquee,
une resistance anormale dans l’hypochondre gauche et en partie dans
l’epigastre. La palpation est tres douloureuse en cette region et ne
permet pas une delimitation exacte. A la percussion, il y a une
sonorite tympanique elevee au niveau de l’hypochondre et du flanc
gauches, sans limites nettes. La limite antero-inferieure de la rate
n’est pas decelable ; le foie n’est pas augmente. Le cceui ne
presente pas d’alterations. Le poumon gauche presente un pen
d'engorgement a la base, le poumon droit normal.

Pas de trypanosomiase. L’examen du sang revele augmentation
notable des leucocytes (myelocytes). La temperature est febrile,
3/'8° C, le pouls tres accelere, filant.

Le malade se plaignanl dc n'avoir pas eu de selles depuis i 5 jours,
prend un purgatif qui provoque 2 selles.

Dans la nuit du n au 12V.07 le malade a des vomissements
sanguinolents dont rexamcn ne denote rien de particulier.

Le malade est tres abattu, refuse toute nourriture. 11 menrt

brusquement dans la nuit du 1 2 au 1 3.

Cette sommaire histoire clinique ne nous donne guere d indications
nettes sur la nature de la maladie. Aussi nous nous etions homes a
supposer l’existence d’une tumeur abdominale de nature indeterminee.

Aulopsie. — Commencee le 13 a 7 heures du matin. Cadavre d un
liomme normalement conforme, dans un etat de nutrition bien
conserve. En rigidite cadaverique ; pas de lesions cutanees.
L'abdomen est fortement ballonn£, la peau y est ties tendue , a
sonorite a la percussion est d’un tympanisme eleve sur toute la paroi
abdominale anterieure.

Apres incision de la paroi. il s’echappe de la cavite abdominale
des gaz fetides en meme temps qu’un liquide epais, sanguino en ,
couleur rouge lie-de-vin. La cavit6 peritoneale renferme environ
200 cc. de ce liquide. A l’examen microscopique on y trouve peu e
leucocytes degeneres, de nombreux bacilles longs, inimobiles, iso
en chainettes plus ou inoins longues ou en filaments.

L'estomac et l’intestin grele sont fortement distendus par cs g
Le long de la grande courbure de 1 estomac, occupan
I’hypochondre gauche, il y a une poche a parois assez epaisses,
presentant a la face externe de nombreuses laches hemorrhagiques
d’un rouge violace. Au palper. cette poche ne presente quu

496

resistance molle, plus ou moins fluctuante. Elle est intimement
adherente d’un cote a la grande courbure de l’estomac et a l’extremite
de lobe gauche du foie, en bas a la partie gauche du colon transverse,
en haut a la face inferieure du diaphragme, du cote externe a la face
interne des cotes, en arrierc a la paroi abdominale posterieure.

En examinant les rapports de la poche, avec le gros intestin, nous
retournons le grand epiploon, et remarquons a la face posterieure de
celui-ci, pres du colon transverse, une larve non enkystee, vivante,
d un blanc laiteux ; 1’extremite cephalique est largement arrondie, la
moiti6 posterieure s’amincit graduellement et se terinine en cone; le
corps presente sur toute son etendue des epaissements annulaires,
rappelant a premiere vue, du moins dans la moitie posterieure, les

tours de spire d une vrille. La larve est legerement fixee a 1’epiploon
par son extremite cephalique ; elle s’en detache facilement et est mise
dans de l’eau physiologique oil elle continue ses mouvements lents de
contraction et d’extension.

A 1 incision de la poche, il s’ecoule en jet un liquide epais,
trouble, sanguinolent, d’un rouge lie-de-vin ; par compression, il en
sort comme c^es caillots de sang nombreux et epais. Incisee
largement, la poche est encore remplie a moitie par une bouillie assez
epaisse, dans laquelle se reconnaissent des tractus fibreux de la rate.
A la paroi de la poche, nous ne trouvons pas d’autre solution de
continuite que celle provoquee par le couteau d’autopsie. L’examen
m lcroscopique du contenu de la poche montre a cote d’elements
cellulaires plus ou moins bien conserves, de nombreux bacilles longs,
i entiques a ceux signals deja dans l’exsudat abdominal.

n est pas hypertrophic, est d’un jaune pale. Sur la face
antero super ieu re, le feuillet peritoneal visceral est particulierement
pai des gaz. Duns la pulpe l’examen denote la presence de
nombreux bacilles identiques a ceux deja signales.

• lnc • hi face externe ne presente pas de lesions macro-
dp npf t rr'1 niuclueuse stomacale presente en differents endroits

1 CS SU US*°ns hemorrhagiques. L’estomac ne presente pas de
communications avec la poche.

Oisophage . pas de lesions macroscopiques.

surfarp fr Srile- uniformement distendu par les gaz, presente a la

pet i tea * ""ri d" n0mbreux endroits, des tidies hemorrhagiques,
petites, irregulieres, d’un rouge vif. Incis^ 5ur toute ga hngeu^ ne

497

presente pas d'ulcerations ni de perforations. Dans la partie
superieure, il renferme de rare AnkyloUomum.

Gros intestine presente egalement a la face externe de petites
suffusions hemorrhagiques, irregulierement disscmiuees ; pas d autres
alterations macroscopiques Dans le contenu du rectum, rares ceufs
i hkylostomum, pas d’autres parasites.

Reins, vessie : pas d’alterat ions macroscopiques.

Cage thoracique: au poumon droit, pas de lesions macroscopiques.
Dans la cavite pleurale droite, quelques c.c. d’exsudat, renfermant des
bacilles identiques a ceux de l’exsudat abdominal.

Le poumon gauche, par toute sa surface, presente des adherences
fortes, fibrineuses, anciennes, avec la face interne de la paroi
thoracique ; la base du poumon adhere fortement au diaphragme, qui
a ce niveau presente des suffusions hemorrhagiques. Le lobe
inferieure presente a la base en arriere, des noyaux fortement
congestionnes.

Ceeur: le pericarde renferme environ 50 c.c. d exsudat sanguino-
lent, tres riche en longs bacilles ; il en est de meme du sang du coeur.
Les valvules sont fortement hyperemiees, mais ne presentent pas
autres lesions.

Les Cultures sur agar et en bouillon faites avec 1 exsudat
abdominal et le sang du coeur, ont donne : celles sur agar, 2 colonies
de microcoques : celles en bouillon, une culture serree de coques. Les
bacilles longs signales a l'examen microscopique, appartenaient done
a une espece anaerobie. Les coques n’ont pas ete identifies. Les
bacilles ne se colorent pas par le gram.

CARACTERES DE LA LARVE

!• Elle se pr^sente sous forme d’un petit vermicule, d’un blanc
laiteux ; le corps est arrondi, compose d’une serie d anneaux, on
f)°rd posterieur est epaissi et surplombe un peu le segment sui
^extremite cephalique vue par la face dorsale est largement arron
A r5 mm. environ du bord anterieur se voit un premier si on peu
Pmfond, suivi bientot d’une serie d’autres tres nets. ,

A la face ventrale, l’etremite cephalique presente a 1
environ du bord anterieur, sur la ligne mediane, un peti on
dfculaire, ou ouverture buccale. Celle-ci ne presente pas d appendices

De chaque cote de la bouche, disposes symetriquemen , y

FF

crochets tres pointus, d’un jaune d’ambre. Ces crochets, que l’animal
sort et rentre alternativement, sont loges dans une depression de
tegument, ou ils sont peu visibles au repos ou a pres la mort.

A un peu plus de i mm. en arriere de I'ouverture buccale, se voit
une premiere indication de sillon

2. Corps : se continue avec 1’extreniite cephalique sans differen
ciation. 11 est compose cJ’une serie d’anneaux, les premiers, pres de
1 extremite cephalique, etant peu distincts. Ces anneaux ou segments
sont au repos, beaucoup plus larges que longs. Vers le bord
posterieur ils s’elargissent un peu et s’epaisissent notablement de
l-K,'on a surplomber la partie anterieure du segment suivant.

La section du corps n’est pas exactement circulaire : la face
dorsale est arrondie, mais la face ventrale est aplatie. Dans la moitie
posterieure meme, la face ventrale presente une surface legerement
< oncave. ( ette concavite est produite par une disposition speciale
du bord posterieur des anneaux a cet endroit. A partir du ioe
segment, le bord posterieur a la face ventrale s’amincit un peu de
chaque cote de la ligne niediane el au niveau de celle-ci presente une
echancrure a concavite clirigee vers la tete. Cette succession

d amincissements et d echancrures produit une espece de sillon pen

profond.

Le tegument du corps ne presente aucun detail de structure; le

rd posterieui des segments ne porte aucune espece d’appendices.

e corps est divise en 18 anneaux plus ou moins nettement

istincts, auxquels il faut ajouter le segment portant la bouche et les

«roc ets ou extremite cephalique et le segment final ou extremite

p sterieure. ( e qui ferait 20 segments, mais nous verrons que le

r anneau piesente encore des indications de subdivisions.

. ^ ' itremitc posterieure : a 1’oeil nu se presente sous forme d’un

on* <'0n'c,ue ^ POInte obtuse. A un grossissement convenable,

1 m Jr rr Stln^llc surt°ut a la face ventrale, deux epaisissements

done ^pleS <UV °rd posterieur des segments du corps, et partageant

distincts ^ Segment terminale en 3 petits segments peu

A la nn' porterait le nombre total des segments a 22.

transversal ° -c SCgment ,crniinal. est un orifice sous forme de fente
an^versale onfice d’excretion (?).

suivant l’etai ^ l"rvc: Durant la vie, les dimensions varient
at de COntraction- ou d ’extension de I’animal. La plus

499

grande longueur que nous ayons mesuree, les trois-quarts environ des
anneaux etant en extension, etait de 23 mm. Apres la mort et
fixation par le sublime, puis alcool a 70%, la longueur est de 16 mm.,
la largeur de 2 a 2*5 mm., la plus grande largeur correspondant a
l'extremite cephalique, immcdiatement en arriere de l’orifice buccale.

La larve extraite a l'autopsie le 1 3. V.07 dans la matinee put ctre
gardee en vie dans de l’eau physiologique et du serum du sang de
bceuf pendant 6 jours : elle fut trouvee morte le matin le 19. V.07.

Cette larve presente des caracteres identiques a ceux decrits par
differents Auteurs pour d’autres exemplaires rencontres chez 1 homme,
et denommes Pentastomum constriction , von Siebold 1852, Poro-
cefhalus constrictus, Stiles 1893. Comme nous l’avons dit plus haut,
Looss et Neumann considerent ces larves comme la forme jeune de
Porocephalus moniliformis , Diesing, 1836. Nous adoptons cette
denomination qui nous parait la plus rationnelle.

Quels ont ete les rapports de cette larve de P orocephalus avec la
maladie et meme la mort de notre sujet?

Differents Auteurs ont signale les ravages causes dans l’organisme
par une ou plusieures de ces larves. Les faits constates jusqua
present tendent a prouver que les embryons parvenus de l’une ou de
l’autre fa^on dans les organes, s’y enkystent et s’y developpent.
Arrivee a un certain stage de developpement, la larve romprait la
poche et chercherait a sortir de l’organisme dans lequel elle est
emprisonnee, pour gagner l’exterieur. Par les crochets dont elle est
armee, par sa taille, elle doit necessairement alors occasionner de
graves desordres.

En examinant les lesions macroscopiques constatees a 1 autopsie
de notre sujet, nous pouvons dire que cet homme a succombe a une
fimtonite suraigue consecutive a une fonte purulent e de la rate.
Quelle a ete la cause des modifications considerables produites dans
la rate? Nous ne croyons pas nous tromper en les attnbuant a a
larve de Porocephalus. L’embryon s’est enkyste, s’est developpe dans
cet organe. Arrive a un certain stage de developpement, la larve a
rompu le kyste et a cherche a sortir de l’organe qui 1 empnsonnai^
Elle a du produire ainsi une irritation considerable suivie ie
dune reaction inflammatoire intense. Ainsi s’expliquent les douleurs
accusees par notre malade dej«a plus d’un mois avant la mort. ^ ur
ees lesions, s’est greffee une infection microbienne, occasionnan

5°°

veritable fonte purulente de la rate avec un d^veloppement
considerable de gaz. Peut-etre, immediateracnt apres la perforation
de la poche par la larve, s’est developpee la peritonite suraigue. Les
caracteres morphologiques et biologiques du bacille, signale dans
1 abces, les divers exsudats et le sang du coeur, nous incitent a croire
que nous avons eu affaire au bacille dc I'ocdeme malin.

LARVES DE POROCEPHALUS CHEZ LE SINGE

I eu de semaines apres le cas du negre que nous venons de relater,
nous avons eu la bonne fortune de retrouver de nombreuses lanes de
Porocephalus chez un singe, Macacus (sp. ?).

Le singe avail ete achete a Leopoldville a un noir descendu du
Ilaut-Congo; il ne paraissait pas malade et fut garde en captivite
pendant plusieurs semaines. 11 fut trouve mort le matin du io.VII.o;.
Bit n qu il n eut pas servi de sujet d ’experiences, nous en fimes
1 autopsie poui determiner la cause de la mort et controler la
parasitologic intest inale. A notre grand etonnement, nous trouvames
de nombreuses larves de Porocephalus , 8 enkystees dans le grand
epiploon, i dans le mesentere, 2 a la face inferieure du diaphragrne.
Nous en donnons plus loin une description sommaire.

Le 3. IX. 07, un autre de nos singes Macacus (sp.?), de meme
espece que le precedent, mourut dans la journee. N’ayant pas servi
aux experiences, l’animal fut neanmoins autopsie.

presentait dans la cavite abdominale de nombreuses larves de
. ° US c. ? enkyst6es ^ans le grand epiploon, 2 dans le

• , e’ ! X^e * ^ace inferieure du diaphragrne, 2 fixees a la
paroi du petit bassin.

I aS de 16sions macroscopiques des organes.
que les Ire T* tr°'Sleme c,e nos s>nges Macacus , de meme espece
TZo ^ P ' rCC°mba- A ’'aUt0PS'e' — trouvames ,3 larves

Tv 6 yS‘“S danS 16 ^and

Le IVnT ma"05C°Pit)Ues d« organes.
fut trouve mort ’ GaramT ^ eSp“C qUe *eS Pn5“dents'

12 enkystees de PoroceptMu T* 'P'Pk>°"

CeS ^ ’®S10ns macrosc°Piques des organes.

ier singe avait paru indispose a certains moments.

50i

Frequemment on l'avait trouve etendu, couche sur le ventre, comme
s’il souffrait de douleurs abdominales.

Chez ces 4 singes, les larves de Porocephalus montraient une
disposition identique.

L'iramense majorite des larves etait fixee au grand epiploon,
quelques unes au mesentere, de tres rares au peritoine parietal : pas
une seule dans les organes, comme le foie ou la rate.

Comme le montre la photographic faite du grand epiploon du
singe No. 3, les larves sont enroulees sur elles-mcmes, entourees d une
mince membrane kystique, a t ravers laquelle les segments du corps
sont visibles. En incisant la membrane pres de l’une des extremites
de la larve, une legere pression suffit pour faire sortir le ver de la
poche.

La membrane qui enferme la larve, est fibrineuse, tres mince, ne
montre pas de vascular isat ion. Le petit kyste qu’elle forme, renferme
outre la larve, une tres petite quantite de liquide un peu trouble, mais
sans elements cellulaires.

Ce kyste est intimement adherent a l’epiploon ou au peritoine
parietal, sur lequel il est fixe : on ne peut Ten detacher sans dechirer
la sereuse.

Immediatement autour du kyste, ni l’epiploon, ni le mesentere, ni
le peritoine parietal, ne presentent d’alterations macroscopiques
permettant de croire a une reaction cellulaire. Le meme dans la
cavite abdominale, il n’y avait pas de lesions macroscopiques des
organes ni d’exsudat.

L’une ou l’autre des larves, enlevee du kyste et raise dans 1 eau
physiologique, ont presente des mouvements analogues a ceux
decrits pour la larve du negre. Elies n’y ont vecu que durant 48
heures au maximum.

Les caracteres morphologiques sont identiques a ceux decrits pour
la larve du negre. Alors que celle-ci apres fixation au sublime, a une
longueur de 16 mm., une largeur de 2 a 2'5 mm., celles des singes n ont

que 10 a 12 mm. de long, et 2 mm. de large.

Mode d' infection : En presence des cas repetes d infection par
des larves de Porocephalus chez des singes tenus en captivite au
meme endroit, nous devons nous demander si ces animaux ne

pas infectes a une source commune. / ,,

Ces singes etaient gardes a la chaine sous la verandah dun

habitation, et pouvaient se promener dans un certain rayon sur le sol
qui entoure la maison. Ils recevaient comine nourriture de la
chikwangue, du pain et du riz cuit, de temps a autre une banane. Tous
avaient ete achetes, 2, 3 ou 4 mois auparavant a des negres descendus
avec ces animaux du Haut-Congo : il ne fut done pas possible de
connaitre plus exactement leur lieu d’origine.

Des quatre singes, seul le qieme avait de temps a autre montredes
signes d’indisposition, en se couchant a plat ventre durant des heures,
refusant toute nourriture.

Actuellement nous possedons encore deux singes qui ont ete en
captivite dans le voisinage immediat des quatre qui ont succombe et
en contact continu avec eux pendant des semaines. Jusqu’a present
m 1 un ni l’autre de ces singes ne presentent des symptoraes de
maladie.

Si Ion admet pour Porocephalus moniliformis , une evolution
analogue a celle de Linguatula , nos singes auraient du etre infectes
p.u les oeufs provenant d’un animal hote de la forme sexuelle, adulte.
Looss, avec Neumann, admet que les grands serpents africains
constituent les hotes des formes adultes. A Leopoldville, les grands
serpents sont relativement rares, et si nos singes ont ete infectes
pendant qu ils etaient en captivite chez nous, comme nous sommes
portes a le croire, nous serions plutot tentes d’admettre que les oeufs
de Porocephalus proviendraient d’un autre animal qu’un serpent.

un autre cote, il ne serait certainement pas impossible que des
““ S ienfermanf un embryon, et tres resistants d’apres Looss, avaient
e \ e mules d une certaine distance jusqu’a notre laboratoire.

. CS c'r^onstances materielles ne nous ont pas permis d’essayer
ecter dauties animaux avec les larves provenant des singes,
et ,, eS qUe I occas'on se presentera, nous examinerons des serpents
1 aut,cs animaux sauvages pour tacher de retrouver la forme adulte

ne / orocephalus moniliformis.

*

EXPLICATION DE LA PLANCHE

Fig- i- Extremite anterieure de larve de Porocephalus moniliformis
du negre, vue par la face dorsale : Gross. 5 diam.

1 ig- 2.- Extremite anterieure, id, vue par la face ventrale.

Fig- 3-

Lane de Porocephalus moniliformis du negre, vue par la
face ventrale : Photographic en grandeur naturelle.

g 4- Larve, id, vue par la face dorsale : Photographic, id.

F'g- 5 — Grand epiploon* du singe No. 3, avec larves enkystees.-
Photographic f grandeur naturelle.

PLATE XU 1 1

Fig. 5

Fig. 3

Fig. 2

Fig. 4

Bredrn, fhM.

ON THE HABITS, LIFE-CYCLE AND
BREEDING PLACES OF THE COMMON
HOUSE-FI ,Y ( MUSCA D OMESTICA, Linn.)

.

ON THE HABIT'S, LIFE-CYCLE AND
BREEDING PLACES OF THE COMMON
HOUSE-FLY (MUSCA DOMESTICA, Linn.)

BY

ROBERT NEWSTEAD, A.L.S., F.E.S., &c.

(LECTURER IX ECONOMIC ENTOMOLOGY \ND PARASITOLOGY)

(Received for publication , November 27 th, 1907)

This paper is reprinted, by kind permission of Dr. E. W. Hope,
Medical Officer of Health, from the preliminary report issued by the
Health Committee of the City of Liverpool, 3rd October, 1907. Some
additional facts regarding the habits of the house-fly have been
obtained recently, but it has been thought desirable to embody these
in the final report, together with the results of further experiments
with insecticides, disinfectants, and other methods of control.

“ This investigation was conducted chiefly with the view of
ascertaining the nature and extent of the breeding places of the
common house-fly {Mu sea domcstica ) in the City of Liverpool; and
also the period of the life cycle of the fly under varying atmospheric
and other conditions; so that some practical measures might be
devised for the destruction of this pest.

“In addition to the common house fly, other species of flies also
occur in dwellings and shops, and several species were bred from
ash-pit refuse, human excreta, &c. One of these, the common blow¬
fly, or blue-bottle ( Calliphora erythroccphala ), may also prove to be
a very important contributory factor in the spread of zymotic diseases,
but it is intended, for the sake of clearness, to deal with these
additional species in the Appendix to this Report.

“ In the popular mind the term ‘ house-fly is applied to almost
all kinds of two-winged flies which are commonly met with m the
dwellings of man. To the zoologist, however, there is but one true
house-fly, and this is the Mu sea domestica described by Linnaeus in
'7 58. This fly is by far the commonest species met with, and quite
9° Per cent, of the flies which infest houses in Liveprool are o 1 11s
kind.

5°‘s

“ Altogether, the refuse from over three hundred ashpits and bins
(chiefly the former) was examined, and 37 middensteads carefully
inspected. Human excreta found in the courts and passages were
also inspected, and breeding-cage experiments with these and the
excreta of domesticated animals were also conducted. The survey
was restricted to five areas in different parts of the City embracing,
in all, visits to 68 streets.*

“ The result of the investigation and survey has proved eminently
satisfactory, both from an economic and scientific standpoint. It
has led to the discovery of the chief breeding places of the fly, and
many new and interesting facts relating to the food of the larval
stages have been brought to light ; so that we are now in possession
of the more important facts relating to the economy of this pest.

“ The chief breeding places of the house-fly may be classified
under the following heads :

“ 1. Middensteads containing horse manure only.

“ 2. Middensteads containing spent hops.

“ 3. Ashpits containing fermenting materials.

“ Leaving for the present the minor breeding places, we may
proceed to consider the chief ones in detail.

1. Stable middens containing horse manure only, were broadly
speaking, found to be the chief breeding places. In the majority of
these the larval stages of the house-fly occurred in countless
thousands, revelling in the heat produced by fermentation. The
adjacent walls often swarmed with newly-hatched flies, and
occasionally one also found enormous masses of their eggs (fig- 2)-
while deep down at the sides, in the cooler portions of the receptacles,
the pupa or chrysalis stage occurred in enormous numbers, looking
like small heaps or collections of reddish berries. Middens containing
a mixture of horse and cow dung were also infected, though to a less
extent than those receptacles containing horse manure only. It is
important to note, however, that in all cases where fowls (not ducks
r geese) were kept and allowed freedom in the yards, relatively few
of the earlier stages of the house-fly were found; and whenever
present were invariably located in places inaccessible to the fowls.

o make ceitain that the fowls were responsible for so remarkable a
jminution of the fly larvae and pupae, a trowel full of these was

treets and also the nature of the receptacles ere here omitted. — R.N’-

509

thrown to some fowls, when they were eaten with as much avidity as
if they had been so many grains of wheat. However much, therefore,
we may deprecate the keeping of fowls in large towns, we must, from
the evidence which has been adduced, consider them as important
contributory factors in the destruction of the earlier stages of the
house-fly. It should be pointed out, however, that fowls are kept in
a very few of the stable yards, so that in the majority of cases the
flies go on breeding uninterruptedly, and, so far as one can gather,
the larvae and pupae have few, if any, other natural enemies but
those already mentioned.

“In one case, where large quantities of a disinfectant (Sharrant s
disinfectant powder) were used in the stable, no larvae or pupae were
found in the manure, though they were swarming in a mass of waste
hops in a separate division of the same midden. Fly larvae were
also absent in another instance where chloride of lime had been used
freely. However, one is not prepared, at the present moment, to
state definitely that the presence of either of these agents had any
deleterious effect on the fly larvae, or that they acted as a deterrent ;
it may have been a simple coincidence, and the matter requires
further investigation.

“All types of middensteads were infected — roofed , vaulted and
open.

" The photographs (figs. 6-9) submitted with this report will afford
some idea of the enormous numbers of the earlier stages of the
house-fly which were found in stable manure.

“2. Only one midden containing warm spent hops was inspected,
and this was found to be as badly infested as any of the stable
middens. The pupae (fig. 1 2) were found collected together in large
masses, and the larvae swarmed in the warmer parts of the material.

"3- A great deal of time was given to the inspection of ashpits,
and it was found that wherever fermentation had taken place, and
artificial heat had been thus produced, such places were infested with
house-fly larvae and pupae, often to the same alarming extent as in
stable manure. Such ashpits as these almost invariably contained
,arffe quantities of old bedding or straw and paper, paper mixed wit
human excreta or old rags, manure from rabbit hutches, &c., or a

mixture of all of these. (See figs. 10, 11.) ,

"About 25 per cent, of the ashpits examined were thus 111 es e

“ House-flies were also found breeding, in smaller numbers, in
ashpits in which no heat had been engendered by fermentation.

“ Both open and closed ashpits were infested, but on the whole
the flies gave preference to the closed receptacles. On opening the
doors of some of the covered ashpits, the flies often came away in
hundreds, appearing like bees round a hive. Ashpits which had been
previously treated with disinfectants were also infested.

" In addition to the foregoing, there were also other collections of
material which afforded temporary breeding places. They are
considered under the following heads: —

1. Collections of fermenting vegetable refuse.

2. Accumulations of manure at the wharves.

“ 3. Bedding in poultry pens.

i- Collections of straw mixed with other vegetable matter and
euthcis, lying in open spaces in poultry yards, were found to contain
enormous numbers of house-fly pupae (fig. 13). The materials had
< \ ic ently fomented, and had also been lying exposed to the weather
or a period of not less than eight weeks.

* be jclrSe accumulation of stable manure lying at the wharf
( arruthers Street) was swarming with fly larvae, and the flies
ec also in laige numbers; so that it is quite evident that a
er o the insects hatch before the manure is placed in the barges
^Tr1™ t0 counttT- Practically all the manure is badly

shn. u 1 W ,en 11 reaches the wharves, so that it is important that it
should be Shipped with as little delay as possible.

jon ^ .. ^ c°mmon practice of leaving 1 bedding ’ material (chiefly

fern Y- m Pou,try pens (containing ducks, geese, &c.) until

such mat 'Y t?kCS Place’ affords breeding places for house-flies, and
such «ater,a] should be removed at much shorter intervals.

refuse- — ^ * breeding places destroyed by speedy removal of

^ a Jectlons °f stable manure removed at short intervals.

“ 7 emptied at intervals of ;-io days.

« j ; Be%'n Pineries.

considered ns tr emptled at ,ntervals of about 14 days may be
removals beino- tocYh'Y breedinS' Peaces, the period between the
under very UnL 1 ^ °l °' <ilC t0 comPiete its life cycle. Only

circumstances, where, for instance, a large amount

511

of fermenting materials are thrown into the receptacle immediately
after it has been emptied, would it be possible for the insect to
complete its metamorphosis in so short a period. In the majority of
cases, therefore, ashpits emptied at such short intervals can only be
considered as temporary breeding places.

“2. Larvae of the house-fly occurred in large numbers in the
receptacle used for holding stable manure in the Corporation Yard,"
Mill Lane. As the manure is removed from this place every week, it
is impossible for the flies to complete their metamorphosis in it.

“ 3. ‘ Bedding ’ in the ‘ piggeries ’ which had been in use for a
few days was found infested with fly larvae. The material used as
‘litter’ was wood chips. This refuse does not lie sufficiently long for
the insect to complete its metamorphosis.

“ The non-breeding places were : -

“ 1. Middens containing excessively moist and non-fermented
cow -dung.

" 2. Patent ash bins.

“ 3. Refuse in ashpits saturated with water.

“4. Human excreta lying in courts and passages.

“ 1. No trace of house-fly larvae or pupae was found in any of the
middens containing cow manure exclusively. 1 he excessive amount
of moisture present in this material was evidently unfavourable to
the development of the larvae. But in all cases where horse manure
was mixed with it, and fermentation had taken place, larvae were
present. The admixture of straw with cow manure would produce
similar results.

“ 2. No house-flies were found breeding in the patent bins, which
is due to the fact that they are emptied daily, or at intervals of a
few days.

“ 3. No house-flies were found breeding in excessively dry refuse,
or refuse thoroughly saturated with water and reeking with inky black
putrid matter.

“ 4- Human faecal matter found lying in the courts and passages
was not infected with fly larvae, and no house-flies were bred from
such matter. But human excreta found in ashpits (several instances
and stable middens (two cases) were found to be infested. _

^ is interesting to note^uTat the bedding material used at this place was

sawdust.

512

“ Reference has already been made to the more important
materials in which the larvae were found breeding, but no details
have been given as to the exact nature of their food. The dietary,
so far as one has been able to see, is almost exclusively that of moist
decaying vegetable matter. Horse manure and spent hops were most
favoured; but they, revelled also in rotten flock beds, straw
mattresses ; they thrived on old cotton garments and rotten sacks,
and waste paper ; and seemed especially partial to the dirty beddings
from rabbits and guinea pigs. T hey fed also on bread, decayed fruits
and vegetables, the excreta of domestic fowls and pigs, and on human
excreta in the ashpits and stable middens. In all cases, however, they
thrived best and occurred in the greatest profusion where fermenta¬
tion had taken place, so that the temperature of the habitat was
raised above that of the outside air.

Like nearly all its congeners, the house-fly undergoes a complete
metamorphosis, in which there are four well-marked stages; these
stages in the life cycle are : —

1. The egg (ovum), commonly' known as ‘fly-blows.’

2. 1 he larva or maggot stage.

* 3- Lhe pupa or chrysalis stage.

4- The imago, or perfect fly. (The final stage.)

1st Stage. The eggs are laid in small irregular clusters, or in
arge collective masses* (figs, i, 2) consisting of many thousands of
1 mdual eggs. They are almost invariably deposited on or in such
stances as will provide food for the larvae or maggots. They are
. , 3 y P ^,Cec^ ‘n narrow crevices near the surface, but, occasionally,
r 3 •stance of from four to six inches below the surface, the
r f . sP0/s cases being fermenting vegetable matter or the
likelv t '^mediately over such materials, or in refuse that is
do not f Crment' * hey are often laid, however, on materials which

» “ - -

due to the delr^ Wh'te’ and Present a highly polished surface,
form of the ' VISC°aS su stance with which they are coated. The
— _ _ _ ggjnay e seen on referring to the micro-photographs

returned8 nS ^ Packed* ,he*r 'BS* >" one small area.

.t, a 1 n to the same snnf r ' ''hen disturbed they flew awav, but

- -poi m a few minutes.— K. X.

5i3

(figs. 3, 4), and the actual size is shown in fig. 1. . The larvae or
maggots hatch from these in periods varying from eight hours to
three to four days ; the average time may be given as twelve hours,
but when laid in fermenting materials the incubation period is
reduced to a minimum of eight to twelve hours.

“ The number of eggs laid by a single fly averages from 1 20 to
140. More than one batch may be laid during the life of the fly, but
this question has not been definitely settled.

“2nd Stage. —The larva or maggot (fig. 9) resembles that of the
common ‘bluebottle-fly,’ or ‘Blow-fly’ ( Calliphora crythrocephala ),
but is much smaller, measuring when fully matured nearly half an
inch in length, and is distinguished also by certain anatomical
characters. It is essentially a vegetable feeder ; animal matter is
eaten only, so far as one has been able to gather, when in the form of
human faeces. It was never found feeding on the carcases of dead
cats and dogs or of birds and fish remains.*

“ They thrive and mature most rapidly, and are always most
abundant in fermenting materials ; but they can also mature in
non-fermenting substances during warm weather, though under such
conditions they do so very slowly. In stable manure they are
generally most numerous a few inches below the surface, and
undoubtedly work their way upwards day by day into the fresh
material, a few hours (five to six) after it has been added to the
previous accumulation. This marked habit is evidently due to the

excessive heat which is engendered in the lower strata of the manure.

“ h'nder the most favourable conditions as to temperature and food
supply they mature in five to eight days ; but, where fermentation
does not take place, this stage, even in hot weather, may be
prolonged to several weeks (six to eight). In middensteads the fully
matured larvae crawl away to the sides or to the top of the wall or
framework of the receptacle ; in ashpits they locate themselves in
various materials as well as ashes, but are evidently partial to old
bedding, paper and rags, usually in or near the centre of fermentation.
After emptying the alimentary tract of organic matter, pupation
takes place rapidly, and the third stage is reached.

" 3rd Stage. The pupa or chrysalis (figs. 7, 10, n) is at first of a
Pale yellowish colour, but rapidly changes to bright red, and finally
t° a dark chestnut colour. It is somewhat barrel-shaped, and varies

. * Taschf iibcrs in l88o, states that he found eggs on the carcases of dead
umals, but does not indicate that the larvae will breed in such substance. R. JN.

CG

514

in length from - rsj of an inch. Small examples are found where
the temperature has been low or excessively hot and somewhat dry.
Large examples invariably occur in fermented materials, more
especially so in stable manure.

“ in stable middens the pupae occur chiefly at the sides or at the
top of the wall or framework of the receptacle, where the temperature
is lowest. In such situations they were often found packed together
in large masses numbering many hundreds (see figs. 6, 7). The flies
emerge from the pupae, under the most favourable conditions, in five
to seven days. In ashpits they occur in the positions already
indicated, and if similar conditions as to heat prevail, the period is
approximately the same ; but in all cases where heat is not produced
by fermentation, the pupal stage may last from 14 to 28 days, or even
considerably more.

4th Stage. I he perfect fly escapes from the pupa by breaking
away the anterior end ; this it accomplishes by inflating the frontal
sac’ wb*ch is situate in the front portion of the head, between the
eyes. By the inflation of this sac, the fly is also enabled to force its
Ua' though the manure or ashes into the open air. When once it
las liberated itself, the wings develop, and when the integument has
SU Ciently hardened the fly takes to wing. Pairing then takes place,
eggs are laid, and another generation is started. The whole cycle
om egg to perfect insect occupies, under the most favourable
, ,ltlCnS’ ^rom ten to fourteen days; but in low temperatures the

e,,C^Ce may ex’tend to several weeks. No growth takes place
after the wings are developed.

j ]r at ^e flies migrate from their breeding places to man's
beino- ^ CJU'te ev*^ent. even to the layman, the primary object

found 1 1PParfnt,y t0 °btain f°°d and sheIter Many flies are als0
they LJT P dmlng Winter and earlv spring, though whether

to ilCIif ZmT:«Iely ” this has not yef been ab,e

remain over tllf» 1 X probable> however, that some pupae* may

the breedino- places" h -'"k ^ ^ f°1Iowin^ spring- NoW that

easy matter t * a'C Cen dlsc°vered, it will be a comparatively
*> matter to ascertain if this is the case or not.

b,lt this u 11 f ° r ‘ u nate^T *d frl not °p todueeU| se^?y ■ was found doting the
not produce a fly in the following year.

winter month?,

“ Little need be said as to the food of the fly itself, as every
layman is conversant with the feeding habits of this little filth-
carrying insect. But the contributory part which many authorities
have claimed that this insect plays in the transmission of zymotic
diseases is due to the almost persistent habit it has of feeding or
alighting upon human excreta.

"In the course of my investigations, more especially on hot days,
numbers of house-flies were seen hovering over or feeding upon such
matter. The faeces were generally those of children, and were lying,
as a rule, a few feet from the doorways in the courts or in the
passages behind the,houses. In one instance no less than five patches
of human excreta were lying in one court, and all of these were
attended by house-flies.

“Temperature, as has already been stated, has a most marked
effect upon the developmental cycle of the fly ; and a sudden check
from heat to cold will materially prolong any one of the stages. Eggs
hatched in eight to twelve hours in a temperature of from 7 5° to
8o° F., at a temperature of 6o° F. in twelve hours, but at 45 F. they
did not hatch until the third day, and then only when placed in a
warmer temperature for the purpose of studying them under the
microscope. The larvae or maggots mature in the shortest period in
fermenting materials at a temperature of between 90° and 98 F.,
but they usually leave the hotter portions of the stable manure when
it reaches a temperature of ioo° to iio° At 540 F. both larval and
pupal stages are considerably prolonged ; larvae kept at this
temperature had not matured at the end of eight weeks, and a
number of pupae kept under similar conditions did not produce flies
until the fourth and fifth weeks.

“ In this Report I have endeavoured to show that :

' I- — The chief breeding places of the house-fly are :

“ (A) Stable middens containing fermenting horse manure or
a mixture of this and cow dung ;

“ (B) Middens containing fermenting spent hops ; and
“(C) Ashpits containing fermenting vegetable matter, or about
25 per cent, of the total number of pits examined.
“II— That covered ashpits and middens were as badly infested as
those which were open.

“ III. — That house-flies breed in all temporary collections of
fermenting matter.

“ IV. — That house-flies breed in relatively small numbers in ashpits
where no fermentation takes place.

“ V. — That they do not breed in ashpits which are emptied at short
internals, or in the patent bins.

" VI. —That the use of disinfectants in ashpits does not prevent the
flies breeding in such receptacles.

“ VII. — That very dry or excessively wet ashes or moist cow dung*
does not harbour them.

" VIII. — That the presence of fowls (not ducks or geese) which had
free access to the stable middens reduced the number of
larvae and pupae to a very marked extent.

“IX. — I hat the life cycle of the fly, in all kinds of fermenting
materials, is reduced to the minimum period of ten to fourteen
days ; and that in the absence of such artificial heat the
cycle may occupy a period of from three to five weeks or
more, according to the temperature of the outside air.

X. 1 hat house-flies do not depend entirely upon excessively warm
weather for breeding purposes, though in hot seasons they
would breed much more rapidly in non-fermenting materials,
and their numbers, under such conditions, would be greatly
increased.

If house-flies are to be reduced to a minimum, I would . . . •
submit the following suggestions

i- That stable manure and spent hops should not be allowed to
accumulate in the middensteads during the months of May to October
inclusive, for a period of more than seven days.

All middensteads should be thoroughly emptied and carefully
swept at the period stated in i.

f he present system of partly emptying such receptacles should
m all cases be discontinued.

r walls of middensteads should also be cemented over, or,

nig t ns, the brickwork should be sound and well pointed.

house fl yGXCe T he6 admix tu re^of 6 W dUng may form a breedi»g Place for the
would also render it suitable for^rS^p^oses^

5i7

3- 1 hat all ashpits should be emptied, during the summer

months, at intervals of not more than ten days.

“4- That the most strenuous efforts should be made to prevent
children defaecating in the courts and passages ; or that the parents
should be compelled to remove such matter immediately ; and
defaecation in stable middens should be strictly forbidden. The
danger lies in the overwhelming attraction which such faecal matter
has for house-flies, which latter may afterwards come into direct
contact with man or his foodstuffs. They may as Vedeer* puts it,
'in a very few minutes . . . load themselves with dejections

‘from a typhoid or dysenteric patient, not as yet sick enough to be
' in hospital or under observation, and carry tne poison so taken up
'into the very midst of the food and water ready for use at the next
‘meal. There is no long roundabout process involved.’

“ 5- Ashpit refuse, which in any way tends to fermentation, such
as bedding, straw, old rags, paper, waste vegetables, dirty bedding
from the ‘ hutches ’ of pet animals, &c., should, if possible, be disposed
of by the tenants, preferably by incineration, or be placed in a
separate receptacle so that no fermentation could take place. If such
precautions were adopted by householders, relatively few house-flies
would breed in the ashpits, and the present system of emptying such
places at longer intervals than, say, four to six weeks, might be
continued.

“6 The application of Paris Greent (poison) at the rate of 2 ozs.
to one gallon of water to either stable manure or ashpit refuse will
destroy gg per cent of the larvae. Possibly a smaller percentage of
Paris Green might be employed with equally good results.

“One per cent, of crude atoxyl in water kills ioo per cent, of fly
larvae.

" The application of either of these substances might, however,
lead to serious complications, and it is very doubtful whether they
could be employed with safety. Paris Green, at the rate of i to 2 ozs.
to 20 gallons of water, is used largely as an insecticide for fruit pests,
h does no harm to vegetation when applied in small quantities ; but
cattle might be tempted to eat the dirty straw in manure which had

MTX^deer, M.B. ‘Flies as sp^d^TTickness in camps.’ Medical

Record, V01. LTV (1898), PP. 429-430.

t This substance is a definite chemical compound of arsenic, copper and acetic acid.

5i«

been treated with this substance, and the results might prove fatal
if large quantities were eaten.

“ j. The use of sun-blinds in all shops containing food which
attracts flies would, in my opinion, largely reduce the number of flies
in such places during hot weather. Small fruiterers' and confectioners’
shops, as a rule, are not shaded by sun-blinds, and in their absence
flies literally swarm on the articles exposed for sale.

“ 8. The screening of middensteads with fine wire gauze would,
undoubtedly, prevent flies from gaining access to manure, &c, but
it is very doubtful if this method would meet with any marked success.
The gauze would rapidly oxidise, the framework supporting it would
probably warp, and numbers of flies would be admitted whenever the
receptacle was opened. Moreover, the erection of such a structure
would prove a great inconvenience and a hindrance to the removal
of the refuse. This, however, does not prejudice the possibility of
erecting a good fly-proof screen in the future.

“ Experiments with crude carbolic acid, cresylic acid, &c, are
being conducted, and the results will be reported later.

“ ha the introductory remarks on the house-fly, reference has been
made to other flies which were found frequenting houses, or were bred
from refuse and excreta during the course of investigation. I he
following is a list of the insects, together with short notes on their
habits and prevalence: —

“ l- Calliphora erythrocephala. The ‘ Blow-fly.’

In some parts of Liverpool this fly is quite as abundant as the
house-fly, and, like the latter, may, from its disgusting habit of
feeding upon faecal matter, also act as a contributory agent in the
spread of zymotic diseases.

1 his fly feeds upon the faeces of man to a greater extent than
the house-fly ; moreover, it is also partial to fruits of various kinds,
especially over-ripe plums and grapes and dried figs and dates, so that
one can readily conceive how it would be possible for the flies to
mechanically transmit the germs of disease from faecal matter to
mt, and tlS a ^aifle percentage is eaten uncooked, the transmission
of infected faecal matter, if present, would be direct.

he female insect lays its eggs in all kinds of raw and cooked
men , t ie carcases of mammals, birds, fish, &c., and wherever such
cmains were found in ashpit refuse, the larvae of this insect swarmed.

5T9

“2. Scalophaga siercoraria. ' Dun ’ or ‘ Yellow Cow Fly.’

“This is a rather large yellow fly which abounds in the country,
but is relatively scarce in the city of Liverpool. It breeds in cow
dung, and was occasionally seen in some of the middens. It rarely
enters houses, and is therefore of no economic importance.

“ 3. Borborus equinus.

“ A minute fly which literally swarms in stable manure all over
the city. As it rarely enters houses or shops, it is of no economic
importance. Fowls eat large numbers. It breeds in the faeces of
the horse.

“4. Stomoxys calcitrans. ' The Stable Fly.'

“ This is a blood-sucking insect, and is one of the recognised
carriers of Trypanosomiasis in tropical countries. In 1906 it was
common in some parts of Liverpool, but this year it has been quite
scarce. It breeds in fermenting horse manure and grass mowings.
It sometimes enters houses and bites both man and his domesticated
animals. Of little economic importance in Liverpool.

“ 5- Homalomyia canicular is.

This species is often common in the dwellings of man, and is for
this reason often mistaken for the house-fly. In Liverpool, however,
it is by no means abundant, and not more than one per cent, of the
flies captured in fly-traps are of this species. It breeds in hoise
manure, and possibly also in the faeces of other animals.

“6. Ant homy ia radicum. ' Root T ly ' or 1 Root Maggot.

“ Closely related to the foregoing species, but not common in
Liverpool. A few examples were captured in fly-traps ; but no
specimens were bred from refuse of any kind.

" 7 • Homalomyia scalaris.

" The larvae of this species were frequently seen in ashpit refuse ,
and a number of these flies were bred from human faeces. It is not
a very abundant species, however, and so far as my investigations
have gone, it has not been found very often in houses or shops. The
larvae feed on all kinds of faecal matter, and are especially partial to
human excreta ; they revel in privies, often congregating togethei on
the shield board in hundreds. Cases of intestinal myiasis in man
have been attributed to the larvae of this fly-

52°

“ 8. Psychoda ( ?) phaleenoides. 1 Owl Midge ’ or ' Moth Fly.’

“ A minute moth-like insect often seen on window-panes in
houses. The larvae of this insect were common in human faeces, and
many examples of the flies were bred from this material. It is also
common in putrid sewage matter. May be looked upon more in the
light of a scavenger, but is of little or no economic importance.

"9. Coleoptera. (Beetles.)

“ Two species were particularly common in ashpits, viz., Crcophilus
maxillosus and PhilonthJis politus. They act as scavengers, feeding
upon all kinds of debris. They rarely enter houses.”

■

PLATE XI, IV

Fig. i. FOUR BATCHES OF EGGS.
NATURAL SIZE

Fig. 2. COLLECTIVE BATCHES OF EGGS IN
STABLE MANURE, NUMBERING ABOUT
il5oo. NATURAL SIZE

I?. Ntwsttad — Photo.

Fig. 3.

EGGS ENLARGED

PLATE X L V

Fig. 4. EGGS GREATLY ENLARGED: ONE SHOWS THE SEGMENTS
OF THE LARVA THROUGH THE CUTICLE

Fig. 5.

I-ARV.E AND PUI’.E IN WASTE PAPER (ASH-PIT REFUSE).
NATURAL SIZE

A’. Nc-jjst tad— Photo.

PLATE XL VI

Fig. 6. MASS OF ABOUT jtj PUI’.E IN' STABLE MANURE.
NATURAL SIZE

Fig.

MASS OF I* UPAS SEPARATED FROM STABLE MANURE SHOW N IN Fig. 6.
NATURAL SIZE

R Nawsteal — Photo.

PLATE XL VII

PLATE XL VIII

Fig. io. LARV/K AND PUP* IN OLD " FLOCK ” BEDDING.
NATURAL SIZE

Fig. LARV* AND PUP.E IN OLD RAGS .ASHPIT REFUSE).

NATURAL SIZE

/

R N twittad— Photo.

PLATE XL IX

I'm. 1 2. MASS OF PUP.E IN SPENT HOPS. NATURAL SIZE

|

I

Fir.. 13.

IM PK IN FEATHERS AND STRAW.

NATURAL SIZE

FROM POULTRY YARDS.

K. Nexvstrad — Photo.

i

523

SOME NOTES ON THE MORPHOLOGY
OF SPIROCHJETA DUTTON I IN THE
ORGANS OF RATS

BY

J. J. van LOG HEM, M.D., Amsterdam

(PRIVAT DOCENT IN BACTERIOLOGY, THE UNIVERSITY OK AMSTERDAM)

From the Runcorn Research Laboratories
( Received for publication , December 3rd, 1907)

The strain of Spirocheeta duttoni, brought from the Congo by
Dutton and Todd, is maintained in the Runcorn Research Labora¬
tories through inoculation of infected blood from rat to rat. By
Levaditi’s1 silver method I have investigated the morphology of the
parasites in the organs of some of these animals killed at different
stages of the infection.

The principle of the silver method may be briefly stated as follows: lhe
organs, after fixation in formol, are impregnated with a solution of silver nitrate,
and then exposed to the reducing action of pyrogallic acid. In thin paraffin
sections the tissues appear of a bright yellow colour, with sufficient differentiation
of the nuclei ; the spirochaetes black, with sharply defined outlines. To bring out
the relation of the parasites to the tissues, Giemsa’s or other stain may be
employed.

The pyridin-silver modification, the ‘second method’ of Levaditi, gave
generally good results, not only with fresh material, but also in organs which had
been kept for some months in formol.

Details of the course of experimental spirochaete-infection in rats
may be found in the extended study of Breinl and Kinghorn;2 here
it is only necessary to repeat that the maximum number of parasites
in the peripheral circulation is reached on about the fifth day after
inoculation, after which the spirochaetes disappear rapidly and
completely, until the first relapse, which occurs in about a week and

extends over one to three days. , ,

Examination of sections (4 to 0 n) of the organs o rats,
at the height of the infection, shows the large and sma 00
and capillaries crowded with spirochaetes, in accordance wi
observations of Levaditi.

The outlines of the parasites are best seen in the capillaries of the
lung and liver, whereas in sections of the heart and large blood-vessels
the spirochaetes are generally found massed in broad strands. In the
spleen at this stage a different condition is observed ; the parasites
being comparatively few in number, and mostly intracellular.

The shape of the spirochaetes is not everywhere the same;
characteristic organisms, showing regular spirals, being generally seen
in lung and heart ; while the liver and the spleen contain atypical
forms, which can be recognised only by their staining reaction, and
by the conditions under which they are found and by the presence of
intermediate stages. These forms are seen as small, circular or oval,
tightly-coiled spirals, about half the size of a red blood-cell.

Similar forms have been described and figured by Breinl and
Kinghorn. These authors found “ occasionally in films made from
the liver and the spleen spirochaetes coiled up into a small compass,
staining a deep red with Giemsa’s stain, and surrounded by a well-
stained membrane.” Levaditi has also observed them as occurring in
the liver of mice, and discusses their probable nature in the light of
similar appearances met with by himself, Manouelian, Cantazucene,
v. Prowazek,3 Schaudinn, and other authors, in the spirillosis of fowls.
Me interprets them as agony-forms, preceding further stages of
degeneration and disintegration. v. Prowazek, on the contrary,
considers these bodies as “ resting stages ” in the life cycle of the
parasite.

With regard to this interesting question, 1 may state that these
forms occur in by far the greatest numbers in the liver, less numerously
in the spleen, scantily in the lung, while in sections of the heart-blood
they could not be demonstrated with certainty.

Moreover, very few, if any, have been observed free in the lumen
of the capillaries, the majority being easily demonstrated m
phagocytes.

1 hese two facts — the excessive disproportion in the number of
thCSe altered forms found in the liver, and their approximately
constant relation to phagocytes — seem to afford a strong argument m
avour of Levaditi’s hypothesis.

Observations were also made at the crisis of the disease. Breinl
an Kinghorn have already pointed out the difficulty of determining

period. In one case described by these authors, the disappearanCf

525

of the parasites from the peripheral circulation occurred between
2 and 5-30 a.m.

In the present case, material at this period was obtained with less difficulty.

A rat (1,540 Bl inoculated on July 20, 1907, showed very numerous parasites in
thick films of tail-blood on the morning of July ^5. It was examined five hours
later, when the parasites were found to have disappeared almost entirely from the
peripheral circulation ; the animal was then killed for the purpose of examining
the organs at the stage of crisis.

Very few spirochaetes were observed in the vessels and capillaries
of lung and kidney. The spleen showed a moderate number of laigc
phagocytes containing partly digested and fragmented spirochaetes.
In the liver, on the contrary, the capillaries were crowded with
parasites, or, to be more exact, were nearly or quite occluded by
swollen mononuclear phagocytes — endothelial cells and cells of
Kupfer— filled with innumerable black granules, many coiled forms

and only a few normal spirochaetes.

This observation seems to lend additional support to t e
hypothesis that the coiled forms are due to the influence o
phagocytosis. At the height of the infection the vessels o t e
different organs were crowded with free, apparently unaltered spiro
chaetes, and the liver showed in addition a great num er 0
intracellular coiled forms ; at the crisis the extracellular parasites ha
disappeared almost entirely, and in the liver were found, besi es t ®
cell-included coiled forms, large numbers of fragmented parasites a
granules, evidently products of intracellular digestion.

The occlusion of many of the capillaries of the liver by swollen
phagocytes seems to account for the hemorrhagic an 1
infarcts observed by Breinl and Kinghorn. Levadit. is als° .
to explain the changes observed by him in the liver o in ec e
by obstruction of the blood-vessels of this organ.

bibliography

Levaditi et Mancunian. Recherches sur Infection *

spirillede la 1 Tick-fever.’ • Annales de 1 InSt„ 1,a>t ’fIC’the Parasite of the
Beeinl and Kinghorn. ' r^xxforr Lverpoo!

African Tick-fever ( Sfirochneta duttoni).

School of Tropical Medicine, September, I9°6- T'ntersuchungen

v. Prowazek. Morphologische und XXIII,

fiber Huhnerspirochaeten Arbeit, a. • a
1906. S. 554.

.

-

MALARIA AND HISTORY

529

MALARIA AND HISTORY

BY

W. H. S. JONES, M.A.

(j*KRSE SCHOOL, CAMBRIDGE)

{Received for publication, December nth, 1907)

It seems likely that disease has exercised considerable influence
upon the history of mankind. In the struggle for existence, man, by
his intelligence, has long since removed wild beasts from the
number of his competitors. In civilised parts of the world, at least,
the struggle is now limited to competition with his fellow-men and
with the parasites of disease. The study of man’s combat with man is
history as at present understood ; the antagonism of man and
parasites may prove to be equally important. Although the
biological study of disease is still in its infancy, the time has come to
collect material, and by careful induction to try to discover any laws
which may appear to have a temporary validity. F urthermore, it is
well so to limit the investigation as to explore more thoroughly a
narrower field before attempting to draw wider inferences or to
formulate more general laws.

The effects of endemic disease are easier to investigate than those
of epidemics, because the influence exerted is continuous, and spiead
over a long period. For a similar reason the question is better
studied from the historical side than by a consideration of the present
state of countries where disease is endemic ; although, of course, the
latter method will furnish valuable information of which use must be
made. Among endemic diseases, malaria, from its wide extension,
the large percentage of a people attacked by it, and its long history,
appears to afford the best starting-point for the enquirer.

Convinced of the truth of the above statements, the writer began
to enquire whether malaria played any part in the history of ancient
Greece and Rome, the decline of which is generally thought to have
commenced during the fourth century B.C. in the case of the former,
and in the second century B.C. in the case of the latter,
suggestion had already been made by Major R. Ross, but there was

HH

5.?o

practically no previous literature on the subject, as the valuable paper
by Professor Aristotle Ivouzis, Tiva irep i eXetoyevmv miperwv,
Athens, 1907, had not yet appeared. On the other hand, the ancient
literature containing references to malaria was of immense size, and
none of it could safely be neglected. The means used to identify
malaria in ancient literature were as follows:

(1) Tertian and quartan fevers are almost certainly malaria.

(2) Quotidian fevers are very likely malaria.

(3) Enlargement of the spleen and early autumnal fevers very

often mean malaria.

(4) Fever in marshy districts is probably malaria.

Of course the ancients knew nothing about microscopes and the
action of quinine.

MALARIA IN GREECE •

^ here is an early reference to 7 ruperos in Homer, but the word
means there, in all probability, “ heat ” not “ fever.” The only other
possible reference in early times is in Theognis, who lived about
54° Bc- at Megara. Tie talks in one passage of lyiriaXos, which
certainly in later Greek often means ague. In a recent volume,
however, of Pauly-Wissowa’s Classical Encyclopaedia, it is suggested
t at this woid was originally connected with the nightmare demon,
n that case Theognis may be referring to the fright-rigors of night¬
mare, and not to malaria.

is impoitant to notice that Hesiod, the Boeotian poet, does not
ntion malaria as one of the farmer’s plagues, though we can be
t iat he would have done so had the disease existed. Boeotia
W malarious, but in very early times just those places in it

da • CcTT ^°r *la,^tat*on wbich are most unhealthy at the present
liisto ' rre> malaria cannot have been in this district from pre-
work r!f t?leS ft 1S tlUe t,iat S“idas quotes eVtaA-r?/? from some lost
Greek it eS'°d'but alth°«gh this word is said to mean ague in later
In th Tta y meant n^tmare at first,
between nen PP°Crat*c XNntmgs, which may be placed approximately
mentioned a 25° BC’. ^ the “ tests ” of malaria given above are
writings cerV*' ^ anc^ ^ must not be forgotten that these
in the Easfor ^ 1InP^> <l long tradition behind them. Accordingly.

parts of Greece, the home of the doctors who wrote

these traditions, malaria was probably known as early as 500 B.C. If
so, it may have come from the East, and malaria is frequently
mentioned in the Sanscrit writings. (See Jolly, Grundriss der Indo-
Arischen Philologic and Altertumskunde , medic in , p. 72.)

But, if the literary evidence can be trusted, the disease did not
appear in Attica before 450 B.C., became more common during the
period 430-400, and finally was so wide-spread as to be designated in
the common speech, though not in the medical writings, by the words
7rupero?, Trupeaato, without further qualification. Every medical
writer after Hippocrates mentions malaria in the clearest possible
language. Even the remittent forms are discussed, although at first
the Greek physicians seem to have been often unable to distinguish
remittent from continuous fevers -a fact not to be wondered at when
we remember that thermometers did not yet exist. Double and mixed
infections were recognised after a while, and Major Ross believes that
in the “semi-tertian" we are to see the double malignant tertian.
Malarial cachexia, with its attendant evils, splenomegaly, anaemia
and dropsy, is described again and again, usually in connection with
marshy places. “ Fever" is often ascribed to over-fatigue, and it is
well known that in a highly malarious country exertion will usually
precipitate an attack. Typhoid, clear descriptions of which are
curiously lacking in the ancient medical writings, generally takes in
them a malarial form, exhibiting tertian periodicity without the
peculiar characteristics of typhoid, diarrhoea and rose-coloured spots,
while pain in the region of the liver is far more common than in
ordinary forms of the disease. Now typhoid often assumes this form
in malarious districts, as the doctors of the non-malarious North
discovered when, in the American Civil War, they treated patients
in the malarious South.

In the non-medical writers malaria is mentioned many times,
though not so often as in Latin literature. I can find clear references
in Sophocles, Aristophanes, Plato, Aristotle, Demosthenes and the
inscriptions.

It is quite impossible, in the absence of direct testimony, to say
when malaria first came into Greece. But the fact that vrvpero 9
in the sense of “ fever ” does not occur before the time of Hippocrates,
combined with the probability that at first older people were
frequently attacked, makes it likely that the disease did not become

532

endemic, at least in Attica, until the close of the fifth century B.c.
But although we cannot date its introduction, it is quite certain that
it became more common after 400 B.c:., until a Greek could talk ot
his touch of fever in much the same way as we talk of influenza, or
an ordinary cold. Nor did malaria change its type. From Hippo¬
crates to Joannes, from the fifth century B.C. to the fourteenth century
A.D., malaria is described in almost precisely the same terms, and
with exactly the same symptoms, and these symptoms arc so clear
that we can recognise all the various forms of the disease as we meet
them at the present time.

MALARIA IN ITALY

W hen and how malaria was introduced into Italy, or when it
became endemic there, is a very complicated question. It is generally
assumed, e.g. by Celli, that malaria was common in pre-historic
times, and that the drainage works at Rome were intended to
diminish the disease, while Cicero is quoted as putting on record
that Romulus founded a city in a healthy spot surrounded by an
unhealthy neighbourhood. It is quite possible that even in early
tunes the disease did exist, but if this be so, it cannot have been
severe, nor can it have been widely spread. For it is certain that
some distiicts, the most highly malarious in the historical period,
were flourishing centres in early times, and could not have been, as
is, indeed, generally acknowledged, very unhealthy. On the other
hand, if the drainage scheme carried out by the kings improved the
Ca ^ C1fy> this improvement did not last, for the Rome of

t C carly Empire was so malarious that it was considered very
c an eroiis to remain in it during the autumn months. It may be
, U ed tka^ Ealy did not sufler much from malaria before 200 B.C,
b . r ter date gradually became more common, as is proved
e"Ces 111 literature, until, during the earlv Empire, many

the”/ 1Str:CtS* and also R°me itself, suffered most severely from

hip-i,]. ? In an cnclemic form. The evidence that Rome was
whelm J T"5 ^ thC begmninS of the Christian era is over-
that the rT 0 1 Study the question, the plainer it becomes

inhabitant.1SeTheeXerftewa marked influence upon the life of every
and there n - r 1006 orace refers to malaria at least six times,

VC re^erences in Martial, including three to the deadly

533

semi-tertian. Horace speaks in the most casual manner of the
foolishness of holding out against the disease until the trembling-fit
causes disaster at the dinner-table. All who could do so left Rome
in the summer, but the poorer people must have suffered severely, as
they appear to have slept very often in booths, or open places, thus
offering themselves as easy victims to the mosquito. Dropsy, a
frequent result of malaria, seems to have been extremely common.
Horace says that a man who will not take exercise will certainly fall
a victim to it, just as we might say that a sedentary occupation must
cause liver-trouble.

Here I should like to meet a possible objection. If malaria was
introduced late in the history of Greece and Rome, why have we
no mention in ancient writers of the time when it first made its
appearance? But it must be remembered that on its first intro¬
duction malaria would certainly be confused with other fevers
(typhoid, for instance) already existing in the country. In fact even
now some kinds of malaria are so like typhoid that the microscope
alone can distinguish between them. It would be only after some
time that malaria could be recognised as a separate disease, and as
late as Galen there is much confusion between the remittent forms
and other pernicious fevers. But in the case of Italy there is veiy
likely a reference in the historian Livy to the time when malaria
first became widely spread over the country. He says that in the
year 208 B.C., an epidemic occurred which did not result in many
deaths, but caused much lingering sickness. This looks like an
epidemic of malaria, and it should be noticed that the date is within
the period of the Hannibalic War, when the land was laid waste and
favourable conditions were given to the mosquito. On other grounds
also, mentioned in this paper, it seems likely that malaria became
common about 200 B.C.

The malarial fevers of Greece and Italy were not confined to the
regular types. Besides the malignant forms included under the
heading “ semi-tertian,” many other dangerous kinds are distinctly
mentioned. In two books of the. Hippocratic corpus, Prorrhetics
and Prognostics , there are frequent references to blackwater fever,
the algide, hyperpyrexial. comatose and other cerebral forms Again
and again mention is made of aphasia, loss of memory, ea ,
convulsions and amblyopia as symptoms of certain kinds of ma aria

534

attacks. It is difficult to discover how far these forms were common
in Rome, owing to the way in which Roman writers on medicine
repeated the remarks of their Greek predecessors, but we are dis¬
tinctly told that the semi-tertian was extremely prevalent.

Both in Greece and in Italy the geographical conditions favour
the development of malaria, as is proved by the prevalence of the
disease in modern times. But, for the present discussion, attention
must be paid to the neighbourhood of Athens and Rome. Near
Athens were two streams, the Cephisus and the Ilissus, which, by
partially drying up in summer, favoured the rapid growth of the
mosquito. Near the Piraeus was a marshy district which must have
proved a continual focus of malaria. Besides these natural breeding-
places, the cisterns in which the Athenians kept their water seem to
h.i\e harboured mosquito larvae. Aristotle tells us that they often
contained the larvae of an insect which was, in all probability,
C hironomus. The streets also of the city were very muddy in wet
weather, so that puddles in out-of-the-way places were certainly
numerous. I may refer the reader to four able articles on malaria
in Greece, by Drs. Savas and Cardamatis, which appear in the viiith
v 190;) volume of Atti della Societa per gh Studi della Malaria.

geographical conditions are shown to be well adapted to the
growth of the mosquito.

T h^fl1*01,116 Simikr COnditions Prevailed. The inundations of the
00 ed a considerable portion of the land near its banks, which
f , notoriously unhealthy. 1 he streets seem to have been as
mu y in wet weather as were those of Athens, while the atrium of
• ; , oman house contained a pool of rain-water which collected

to the impluvium through a hole in the roof, intended no doubt, at

l° ICt °Ut the smoke froni the household hearth.

. , . * C c notlced that just at the time when malaria appears

sLerelirTTuendemiCin Att'Ca and ItalN (4*> and 200 B.C.),
proper control f * C°Untry Waste> and Prevented for many years the
Ihe'spr ^ o^^atln" StrCamS and CaaaIs‘ ^is would favour

535

INFLUENCE OF MALARIA UPON GREEK AND ROMAN HISTORY

In estimating the effects ol malaria upon the history of Greece
and Rome several facts must be borne in mind. In the first place
the ancients had no quinine ; the disease must have run its course
without being mitigated by any efficient remedy. It is therefore
probable that, in Greece at least, some element of the race was
weeded out 1 his would be the Northern strain to which, in all
probability, the Greeks owed their best qualities. Again, the
virulent remittent forms of the disease seem to have been particu¬
larly common. The poet Martial, whose works make but a moderate¬
sized volume, mentions the malignant semi-tertian three times. In
Hippocrates frequent reference is made to those cerebral forms of
malaria which, in the words of Mason, lead to “ permanent psychical
disturbances.” Malarial cachexia, with accompanying derangement
of the digestive system, was very common. Furthermore, the
extent to which malaria occupies the medical treatises is, to say the
least, surprising. By far the greater number of the fever cases in
the Hippocratic writings refer to malaria in its intermittent or
remittent forms ; while in the Latin author Celsus, who flourished
about 50 A.D., other kinds of fever are scarcely mentioned at all, so
that in his book febris is practically equivalent to malaria.

1 he Greeks themselves seem to have noticed that malaria often
produced strange psychological effects. So much is plain from their
use of the term fxe\ayx°^a and its cognates, which, in the common
speech, denoted that a man was crazy, neurotic or even mad, while
they were almost certainly medical terms originally, denoting malarial
cachexia, or, sometimes, the epileptic convulsions which are often
to be observed during a malarial attack. I he problem is made a
little complicated by the fact that Greek medical terms rarely coincide
exactly with any now in use, a source of confusion against which the
historian must be always on his guard. “ Melancholy ’ denoted a good
many kinds of bilious conditions ; but when it is observed that the
Greeks themselves thought that quartans had their origin in black
bile” 0 xXaiva Xo\v), that Galen declares large spleens to be due to
excess of the “ melancholy ” humour, that cases of “ melancholy ’ are
said to be common in autumn, it seems practically certain that the
word was often used to describe malarial states, and that the Greeks

53^

observed how malaria affects the temper, rendering the patient morose
and cross-tempered. It is interesting to note that the word /zeXayxoXia
becomes common in Attic literature just at that time (the last
quarter of the fifth century) when it seems likely that malaria first
became endemic in Attica. Plato in the Timaeus declares that vice
is due to bodily disease, and in particular derives peevishness,
melancholy, rashness, cowardice, forgetfulness and stupidity from
bilious humours finding no outlet from the body.

1 here is a remarkable reference to the influence of malaria upon
character in the Hippocratic treatise Airs, Waters and Places.

I hose who dwell in hollow, hot districts,” says the writer, “where
the winds and water are warm, are neither tall nor straight. If
they drink the water of the place they have diseases of the spleen
and stomach. I hey are stout and fleshy, dark-coloured and bilious.
Hy nature they are neither courageous nor of great powers of
endurance. But there are no ill effects when the water is drained off."

What, then, is the change of character which accompanied the
decline of Greece and Rome? Between 450 and 300 B.c. the Greeks
(at any iate the Athenians, for it is of them that we know most) lost
then manly vigour and intellectual strength. Patriotism was still
considered <1 viitue, but few had the energy and initiative to translate
t ieor) into practice. Love of ease and comfort grew apace,
hilosophy became pessimistic, and there was much brooding over

cat 1. In art there appeared a tendency to sentimentalism. The

of the third century b.c. was unequal to the effort for further
1 h ess, and ne\ er recovered the vital force he once possessed.

n Rome and Italy the change was different. The Roman of the
early Lmpire can scarcely be called weak. But he had changed.

• , ernness had become brutality. He was no longer contented

banen &t Sim^Je llfe’ but *°ved gorgeous display and magnificent
old Rr n urthermore, the population of the city changed. The
lament ,^PP^ent^’ &rew fewer in numbers, for there is a constant
the ininpr'- 1 a-mi 1Cb Were smah> while crowds of foreigners flocked to
armies were °f whom rose to Povver and influence. The

stock. Hist ° en reciuitcc* horn Spaniards and other more virile
the old Rom 1 ans.anc* moralists repeated ad nauseam the truth that
Now SPlnt WaS dead

w ^ would be absurd to maintain that all these changes were

517

produced by malaria. The Greek outgrew his city-state, lost his
faith in religion, and exhausted his strength in a series of suicidal
civil wars. He practised unnatural vice to an extraordinary extent,
and this, with other excesses, produced the natural consequences. But
surely these influences must have been aided in their operation by
the presence of an insidious foe, which weakened the individual from
his birth, and left him an easier victim to the disintegrating forces of
his environment.

The Roman also outgrew his institutions, and no longer found
satisfaction in political life. The farms, which had bred a strong-
race of yeomen, gradually gave place to large grazing estates. Com
could be imported from abroad more cheaply than it could be grown
at home, and the farmers crowded into the already-congested
metropolis. Economic causes, then, as well as political and
psychological forces, were at work both in Greece and in Italy during
the period of decline. But the fact remains that the Greeks became
a race of inefficients, while the Romans of the empire may be roughly
divided into two classes — a few luxurious debauchees and a host of
debased and poverty-stricken retainers.

It is much to be regretted that scientists have paid but little
attention to the effects of malaria upon national prosperity and
national character. The economic effects, indeed, are noticed with
more or less detail by many observers. Celli in his M alaria talks of
the loss of labour and production caused by the disease, and Clemow
describes the appalling incapacitation and economic loss which
accompany its ravages. But its influence upon character has never
been thoroughly investigated. North, in his fascinating work
Roman Fever, does say something on the point, but confines himself
to the general statement that a highly malarious district, if left to
itself, must contain a population that tends to moral degradation.
Professor Nieuwenhuis, of Leyden, who has studied the wild tribes
of Borneo more than any other traveller, writes to tell me t at in
that island malaria actually has the disintegrating effect which
assert it had among the Greeks. The results of his investigations
are to be found in h,s book Qucr durch Borneo. But there certain y
is room for a book containing an adequate study o t is ques ™
both the physical and the psychological standpoints
preliminary to such work seems to be a historical study of the

53*

at which malaria was introduced into various districts, and of the
effects which followed this introduction. Up to the present our
information is very slight, being confined to such instances as the
invasion of Mauritius by malaria in i860.

There is also a pressing need for an investigation into the moral
and intellectual characteristics prevalent in highly malarious districts,
and a comparison of them with those of neighbouring parts, under
similar economic and political conditions, but untroubled by malaria.
This is another unworked field for historians. I take it that the
effects of malaria are threefold :

(1) It may kill, or drive away, the inhabitants. There is
evidence that the population of Greece gradually declined, and
that an inferior race, but one relatively immune to malaria,
supplanted in course of time the older population. I hope to
publish this evidence shortly.

(2) It may cause physical and mental degeneration by
making childhood unhealthy.

(3) It may cause inactivity by punishing over-exertion and
fatigue.

The first thing to do will be to collect material, and for some time
I have been impressing upon anthropologists the necessity of
observing the psychological peculiarities of peoples among whom
malaria is endemic. It is probable enough that the lapses into
barbarism which so often disgrace Europeans living in tropical
countries are at least partially caused by this disease or its sequelae.
I am told that German officials are forced by' the Government to
tan7 quin’ne w*th them when they set out for malarious regions of
Afiica, and the reason assigned is that moral deterioration may follow
malaria unchecked by prophylactic measures. Unfortunately, I have
icon unable to discover from written evidence whether this is correct

CONCLUSION

is certain that both Greece and Italy' were, at least in their
lobl important centres, highly malarious during the period of their
. ^ ht ther Athens and Rome were malarious when they were

f Jn *,<>VVCr anc* greatness it is impossible to state for certain,

ajj ^ 1 sea sc was in all probability rare, even if it was present at

the other hand, the decline was in both cases accompanied

539

by an increase of malaria. The change in the Greek character was
just that which we should expect malaria to produce in a highly
sensitive and cultivated people, while the savage brutality of the later
Romans may be due to the same cause. The peculiar effects of a
disease on national morality will certainly vary with the prominent
national characteristics. The more effeminate Greek grew weak
and inefficient ; the stern Roman became viciously cruel. But it
must always be carefully remembered that other factors, physical and
psychological, contributed to the change in both cases. The growth
of intelligence, resulting in dissatisfaction with existing institutions;
the decay of agriculture or trade ; the slow effects of vice and luxury ;
the exhaustion of the conditions which stimulate a people to aspire to
national greatness all these were doubtless important factors in the
decline of Greece and Rome. But malaria gave rise to physical
conditions which afforded an excellent opportunity for other influences
to produce their full effect. It must also be remembered that these
conditions would be reproduced from generation to generation, for
malaria was continuously present, apparently increasing until nearly
everybody was more or less infected. A temporary disaster, whether
it be war or a virulent epidemic, often stimulates to great deeds and
heroic actions ; but a weakening endemic disease, attacking every
fresh generation as it is born, gives the nation which is its victim
no chance to recuperate.

In the preceding pages 1 have indicated very briefly the main
conclusions which it seems just to draw from a study of the classical
literatures. But only the fringe of the subject has been touched,
even within the limited area to which I have confined my own investi¬
gations, and a vast field remains to be worked by those who will take
the trouble to trace the effects of malaria upon other nations. It is
only from a comparison of results obtained in a series of enquiries
that any really valuable knowledge can be expected. The influence
of diseases other than malaria is also most important, and should be
carefully studied. Mr. T. Spencer Jerome, who has for many years
Paid attention to the biological aspect of history, sends at my request
the following note on the transformation of the Roman character.
The remark it contains about pestilence killing off those o t e
greatest nerve force is interesting, as it is just possible that the great
Athenian plague of 430 B.C. weakened the general health by attack-

540

ing and killing those best fitted to be parents of vigorous offspring.
It is a fact that the Athenian population was permanently reduced by
this epidemic, and perhaps malaria, which appears to have become
endemic in Attica soon after 430, found a people already weakened
and less able to resist its ravages.

NOTE

I he influence of diseases on historical development will obviously
be mainly through their effects in weakening individual nervous
systems, and, as a result of this, individual character. Now it seems
\\ell established that all infectious fevers play an important part in
the etiology of the psychoses and neuroses. Typhoid especially is
incontestably a cause of psychasthenia, and often profoundly alters
the whole nervous system. So also influenza. The number of
se\ ere infectious maladies prevalent throughout Roman history is
well known. After the great outbreak of pestilence in the time of

Marcus Aurelius, it kept reappearing from time to time for a century
or more.

Another point deserving consideration is this : — Is it true that the
neurasthenic person is relatively immune to other and more imme-
lately dangeious diseases? Dr. Beard, whose studies of neuras¬
thenia are so well known, asserted this positively- though the
exp a nation he gives seems slightly fanciful* Dr. V. C. Vaughan,
examined the matter of the typhoid outbreaks in the American
army camps at the time of the Spanish war, informed the writer that
e ever seemed to select the soundest men. If this be so, then a
g ontinuance of severe infectious maladies would operate not only
, mUCh nerVOUS debllity- but also by a kind of inverse
Deonlp SC eCtl°n t0 ebminate the relatively sound elements in a

Droblf'inc Jtha\ l*1<? biological method of approaching historical
cease ;rr1rSrbeCnbiegUn: We hope for real progress, and may
tized virtue^ ^ ex? dnatlon of historical developments on hyposta-
step to ;nStitUti°nS and the ^ But. as a prehminary

clarify our kno^'^ biology to Roman history, we need to

human character oTd ° t What WCre the changes in the average

in time. One la Peri°d, and the order of their development

*Sexi7TTv - - - bc scoPtical as to the extent and accuracy

• exual Neurasthenia, by Dr~G jvTDT — ^ - -

y c»r. O. M. Heard, 4th Ed., pp. 63-4, 72-3, 76-8, 116.

54i

of our comprehension of these facts. Tt is probable that Tacitus,
Juvenal and Pliny, as well as most other writers of imperial times,
throw more light upon their own character than upon those depicted
by them, while our knowledge of early Roman character is involved
in an iridescent haze. T. S. JEROME

Capri, Italy

PLUTARCH ON HEALTH

Let us sec whether the theory put forward in the preceding pages
is confirmed by a Greek work which was written about the end of the
first century after Christ.

Among the Moral Treat isos of Plutarch is included a work the
Latin name of which is de lunula sanitate praecepta. It is intended
for the use of those who devote themselves to study or politics (137 c),
and lays down the rules which must be observed by such if they wish
to keep in health. At first sight it appears to be a sensible but
somewhat commonplace series of remarks ; but a more careful reading
proves that it throws a flood of light upon the hygienic conditions of
the period when it was composed. Before proceeding I will give a
short analysis of the contents.

It is necessary to keep the hands warm, as chill in the extremities
invites fever (123 A). It is useful to accustom the body when in health
to the diet which would be necessary in illness, and it should not be
thought insufferable to dine unbathed (123 B — D). The body ought
to be nourished, as a rule, with simple foods, so that, should an
occasion occur when feasting cannot be avoided, no harm results fiom
indulgence. If some high official invite 11s, or other imperative call
come when we are indisposed, it will be less boorish to abstain than
to fall into irXevpiTK or <f>peviTui through false shame (123 L i_4 L) j.

Food and drink are to satisfy hunger and thirst. Dainties should
not be consumed merely because they are costly, or because we wis 1
to boast that we have eaten them. The body must not tyrannise
over the soul, nor yet the soul over the body, so as to cause ovei-
indulgence. A man should take a pride in his power to abstain.
Rich, tempting dishes cause us to eat too much (124 E— 126 B).

Pleasure is impossible without health. Me are wont to neg ec
plain living when we are well, and in sickness to lay the blame upo
climate («ep«9, xwpas'), instead of our own intemperance. en

542

ill we should say to ourselves that drinking cold water, or an untimely
hath, has deprived us of many pleasures. In this way we are made
more careful when in health (126 B 127 B).

Granted that fevers are caused by exertion, heat and chill, too
much food increases the liability (127 B D).

The forewarnings given by fever must not be neglected. Some,
when they feel an attack coming on, betake themselves to baths and

banquets, lest they fall ill before they have satisfied their desires;
others, more refined (/co/iyfrorepoi), are ashamed of showing that they
are unwell, and obey' the call of their companions; most men hope
that the feeling of uneasiness will pass off. But on the morrow they
hu\e to confess to catarrh, fever or colic Then they will beg the
doctor to allow' them wine or cold water. AJJ such should remember
that the unhealthy body feels no pleasure in the indulgences which
caused the trouble (127 D 128 E).

The over-strict diet of one who is always afraid of his health giving
way is certainly to be blamed, as it renders the body liable to fall
sick, while it makes the spirit timid and unenterprising; but it is
c ertainly unwise to wait for those internal pains which are the fore¬
runners of fever before moderating one’s desires and appetites. It
necessary also to be on the watch for bad dreams, crossness of
temper and melancholy (128 E _ 129 C)

W hen \ isiting a sick friend enquiry should be made whether it
P et ora. heat, exertion, lack of sleep or wrong diet that caused
* — *dls answers Wl11 serve as a guide. One should care for
p°^n mode °f life, avoiding all excess (129 D — 130 C).
m . eaChl?- and discussion are excellent physical training. The
'er> o inn-keepers or muleteers can be neglected (130 C 13 1 B).

■17 Crf f,Xe«C*Se’ CO,d batbs are to be avoided. Those who so
diet jl * ’ UnleSS they foIIow in the smallest details that strict
fire 0 3 rii 1SDS; UndeSirabIe- rt is oil body near a

fowl ard'lio-kt fi ^ l°°bed e£2s are not desirable; vegetables,
b"ed& w 6 • arC l° ^ thC Stal)le food Milk as a drink should
after exDosi, lnoderatlon >s good, but not as a “pick-me-up"

be thought JV- Water shouId be drunk several times a day. If it

-^ay be drunV^^ D^^F) ^ ^

lIC eatinfir> a man should exercise his mind with a book or

543

conversation. This will make him less attracted by the pleasures of
the table (133 A- 1 34 A).

Emetics and purges are had. Dieting is the proper remedy for
indigestion. 1 1 something must be done, vomiting is the less evil,
but violent drugs must be avoided. Drinking water or fasting for
a few days may be tried, or even an injection. Most people take
refuge at once in strong purgatives, and suffer for it (134 A — F).

On the other hand, a rigid system of fasting is bad. It is absurd
to keep well by ceasing to perform the functions of living. Nay,
idleness is not healthy (135 A— 136 A).

Toil should not be varied by exhausting pleasures. Love of
honourable pursuits will drown anv desire that is felt for the latter
(136 A-E).

A man should learn all he can about his own constitution, what
suits it and what does not. It is important that care be taken not
to tax it at the change of the seasons (136 E 137 B).

Students must not tax their bodies by too much study, as the
many do by worry and exertion at harvest-time. Otherwise they
will be compelled to lay aside their books, while they are recovering
from a fever (13/ C-E).

It will, I think, be admitted that at the period when the treatise
was composed there was much ill-health. The precepts given by the
writer himself arc strict, and he distinctly states that there were some
who imposed upon themselves such rigid rules of life that health was
obtained at far too high a cost ; for they could not use it without
interfering with those prescribed habits which kept them well/ The
writer does not seem to be referring to infectious sickness, for he
nowhere mentions either contagion or infection. t Indeed, either
ancient Greece was singularly free from infectious maladies (other
than occasional epidemics) or else the Greeks did not think the
danger worth considering. At any rate isolation of the sick, and
similar prophylactic measures, were not generally recognised. £ The
great danger, according to Plutarch, was “ fever.”§ The symptoms of

t When calling1 on* a sick friend the visitor is not supposed to take any

^TSgit'ionbv sulphur was known as early as the Homeric period (Odyssey
481, 4931. but the medical writers appear to know nothing mey

'gwd it as a superstition ? Certain skin and eye _ diseases, with consumption,
egarded as infectious, but not fevers (Aristotle, Prob. vn, °-) n • ,72 E • 137 D.

§ Mentioned several times- 1*3 A; 127 II, E; 128 A, F; 129 lJ , 13- > 37

''ice a more specific name- <f>p€PiTl<;, is given (124 E).

544

fever are not described, but a warning is given not to neglect the
premonitory signs, and among these are crossness of temper and
melancholy * But the risk of falling ill of fever is said to be greatly
increased by certain actions or habits. The “ causes ” of fever
include : —

(1) Violent fatigue (/co7ro<?).t

(2) Extremes of temperature, t- especially chilled extremities and
cold baths at unseasonable times.§

(3) Over-indulgence in food and drink.!!

(4) Insufficiency of rest and sleep.*

In addition to these definite dangers, the general tone of the
treatise implies a strong recommendation to avoid taxing the body
or mind by excess in any form.

Now, with the exception of that prophylaxis which is the direct
result of modern discoveries,** this advice is just that which is now
given to those who dwell in malarious regions. tt If it be urged that
it is unsafe to conclude that malaria is the disease which our writer
had in mind, inasmuch as moderation is a good rule under any
climatic conditions, the following points should be considered.

Only in a highly malarious country can it be said that fatigue will
bring on fever. Of course, fatigue may be the indirect cause of a
feverish chill, but it is in malarious countries that the necessity of
avoiding over-exertion, and the practical certainty of an attack
should this rule be violated, become painfully obvious from repeated
experiences. It is also remarkable that such stress is laid on
keeping the hands warm. In a non-malarious country cold in the
extremities can scarcely be considered a grave danger, but in a
malarious region great care must be taken to avoid chill. It should
also be noticed that the change of the seasons is regarded as an

*129 c.

+ I27 », E; 1 29 D.

g 127 f h**v<rei 9 teal Sih TrepMPeoxt ) ; 129 D.

8,23 Ai 126 F; i27 E; Cf. also ,3, ,

i* 1 * *"4 “ 1 127 C, K ; ,2g A ; ,^9 D ; Cf. also 137 C.

^29 D ( aypvTrviav ) ; 137 D.

++S 8”r USe °f qUinine and niosqui‘o-netting.

Malaria and Manson Tropical Diseases (malaria).

545

especially dangerous period." and malaria is most common just before
the autumnal equinox. Particular attention should be paid to the
difficult passage 137 C,t the general drift of which seems to be that
poor country-folk constantly fall ill during their exertions at harvest¬
time. Indigestion and constipation were evidently common com¬
plaints when the writer lived, as he tells us how the people took refuge
in violent purgatives.* Now although there are many causes of
these stomach complaints, derangement of the digestive organs is
the invariable accompaniment of malarial cachexia. If any doubt
still remains as to the kind of fever which is referred to, the use of
<f>peKm<?§ in 124 B should dispel it at once. 1 his word is certainly
employed by the medical writers to denote a very pernicious kind of
remittent malaria.il

I have tried to prove in the above discussion that the Greeks of
the first century A.D., or at any rate some of them, were liable to
malarial attacks whenever they put the body to unusual exertions ,
in other words that the country, or some part of it, was highly
malarious. What must have been the consequence to the people at
large? What nation can prosper or develop if strain has to be
avoided at all costs? How are war, commerce and successful agri¬
culture to be carried on under such conditions? Surely nothing but
stagnation is possible. Now the literary evidence shows that malaria
was common in Greece during the fifth century, since tertian an
quartan fevers are constantly referred to in the Hippocratic writing
But there are only two allusions before 500 B.C. which can point to
malaria (Trvper'o^ in Iliad xxii, 3 b and fyrlakos in Theogms I74)>
and both of these are doubtful. Hesiod never mentions fever among

1 137 B.

toVa ^ ydp p.ucpoXoyta, Kal dveXevOepia *
pftdvovaiv ot iroWol ire pi re avyicop.L&<is /cap7rwv aa6pd teal
^ovov,, dypvirviac, Kal rr epical*
ov\a tov (Tcoparof, ov/c at-iop eerri oeotev r 1
\o\oyoi Kal 7 toXvtlkoL /

t <t>appaKeia<i, e<f>' a<> o! iroWol <f>epovrai TrpoxeiPU)'i ^

§ For <f)peviTl<; see Hippocrates, Kuhn II, *7» 28, 299, 300, 3 7

" S“ Ul“'S V°L «• . second m Of tie’ fifth

* The word irvpeTOS does not occur again s t0 niean “ heat.” Vi by

'tuw b.c. In Homer it was said by some ancient comm » g to a tradition that malaria

1 they not assume the meaning to be fever . b > F
1 not exist in Homer's time.

54^

the plagues of the Boeotian farmer, but Plutarch, a native of Boeotia,
has a different tale to tell. There can, I think, be no doubt that
malaria was rapidly on the increase^ from the fifth century B.C., and
that it was largely responsible for the lack of energy that the Greeks
began to exhibit during the fourth century.

Plutarch noticed that the feverish attack is often preceded by
melancholy and crossness of temper,* two most noticeable symptoms
of malaria. The physician Hippocrates went much further. After
carefully describing what regions are most malarious, he goes on to
say that the inhabitants of such regions are stunted in growth, and
have neither courage nor powers of endurance.t Plato declared that
ill-health (the symptoms point to malaria as much as anything else)
produces crossness, melancholy, rashness, cowardice, forgetfulness and
stupidity.^ Surely it is reasonable to conclude, not only that
malaria produced disastrous changes in the Greek character, but
‘^so t^at the great thinkers among the Greeks were perfectly con¬
scious that it did. Plato would have introduced remedial measures,!
but, of couise, his advice was not taken, and Greece gradually fell into
the unhealthy state which is manifest in the pages of the treatise
dc sani/atc tuenda.

used in non-medkaf^i become more common, and are regularly

Demosthenes, i i8t 2G r s o mean intermittent (i.e. malarial) fevers. See

*'*9 c.

+ Airs, Waters, Places, Kuhn I, 566 s67
XTimaeus 87 A. 5 5 7'

°ne of' the Smarlf of 'wi!!*1'6 and g°°d institutions (Timaeus 87 B.). This reminds
fibre of malariouks Hoc cit.) that might restore the moral

1 lat° s ,at^ views oS vFce%J &£** drainaSe bri "K back their health. For
6 see Gomperz, Greek Thinkers, Vol. Ill, 225, 226.

TWO NEW HUMAN CESTODES AND A
NEW LINGUATULID

549

TWO NEW HUMAN CESTODES AND A
NEW LINGUATULID

BV

J. W. W. STEPHENS, M.D. Cantab., D.P.H.

(WALTER MYKRS FELLOW IN TROPICAL MEDICINE, THE UNIVERSITY OF LIVERPOOL)

(Received for publication December 20th, 1907J

I. A NEW BOTHRIOCEPHALID IN MAN

( Dibothriocephalus parvus n. sp.)

Three pieces of a tape-worm, none of which had a head, were
received by me from Dr. Elkington, Tasmania, in 1906, with the
following history : ‘ It came from a Syrian, not long arrived from his
native country, and aged 37. It was caught on December 20th, 1898,
at Launceston, with the aid of Filix mas. A quantity had already
come away before he came under treatment. So it is possibly a
Levantine product after all, and not Australian in origin. Lie is
supposed to have come from Beyrout, but this is not certain.

A superficial examination shewed that the tapeworm had all the
appearance of a Dibothriocephalus. I he proglottids had a clearly
defined central uterine rosette, and with a lens the openings of the
cirrus and uterus could be seen, and in several segments the
cirrus was extruded. The surface of many of the segments was
much corrugated with transverse and longitudinal furrows (probably
due partly to the preservative and partly to erosion (digestion) of the
surface.

The strobila consisted of three portions, the lengths of which were
respectively 1025, 940 and 690 mm. In the portion, 1025 mm. long,
there were 420 proglottids, usually broader than long. 1 he smallest
anterior proglottids measure 1*3 mm. broad by o-6 mm. long. The
largest posterior proglottids measure 5‘0 mm. broad by 3 o mm. long.
The terminal segments tend to become quadrate and measure 4 o
by 4-0. Several segments approximately quadrate are, howevei,
interpolated irregularly so that the gradual decrease in breadth and

550

increase in length from one end of the worm to the other is broken
at intervals and is not uniform as in D. latus , &c. As stated above
the dimensions of the largest gravid segments are 5 by 3 mm. The
uterus forms a central rosette with four to five loops on each side of
the median line, occupying about the middle half of the length.

In a proglottid measuring 37 by 2*25 the genital atrium is
situated o^-o^ mm. behind the anterior margin, and the uterine
opening is situated the same distance behind the genital atrium.

The eggs, operculated, average 59^2 by 407//. Calcareous
corpuscles were absent in those segments examined.

Fig. i. Dibothriocephalus parvus

(1) Outline of uterine rosette

(2) Genital atiium, uterine pore and protruded cirrus

1 he question now arises as to the identity of this worm.

0) On comparing it with D. latus we see that the maximum
width of gravid segments is 5 mm., while the minimum width of
t,ravid segments of D. latus is 10-12 mm., and the maximum width
20 I|mi ’ so that the worm is a very much smaller one, as is evident
t e naked eye. I his is also shewn by the size of the quadrate
nents which in D. latus are 6 by 6 mm., while in D. parvus they
4 by 4 mm., and moreover the quadrate segments in D.

551

occur posterior to segments which attain a width of 1 5 mm., while
in D. parvus they occur after segments, the maximum width of which
is only 8 mm., and in the case of the interpolated segments, after
segments which are still narrower.

(/>) The whole strobila appears always to be distinctly thicker
than D. lotus , a point which is especially noticeable along the lateral
margins.

(r) Secondly there is no indication of the serration due to the
projection of the posterior lateral angles, a feature well marked in
D. lotus.

(ft) The eggs of D. parvus (59-2 by 407 A* ) are smaller and
rounder than those of D. I at us (6871 by 44-45^)-

(r) Calcareous bodies are absent in D. parvus , while they aie
present (few) in D. latus.

(a) On comparing with D. cordatus, we find that this has onl>
50 immature segments while in one of the pieces of D. parvus theie
were at least 200 segments before maturity.

(b) The mature segments of D. cordatus are 7-8 mm., t
maximum width of D. parvus is 5 mm.

(c) The quadrate segments of D. cordatus measure 5-b mm.
square. Those of D. parvus 4 by 4 mm.

(d) The uterine loops of D. cordatus are 6-8. Those o
parvus 4-5.

(0 Eggs, D. cordatus 75-80 by 50. D. parvus 59 2 by 4° 7^-

f) Calcareous bodies. D. cordatus , 28-30, numerous. D. parvu ,
absent.

The characters of this bothriocephalid seem to me j

sufficiently distinct to warrant the making of it a new sp -g

propose the name Dibolhriocephalus parvus. The tyPe p
deposited in the museum of the Liverpool School o 1 P

Medicine.

II. A NEW HUMAN CESTODE

( Taenia br earner i , n. sp)

'■ !■'!■ w. ' — """

x :s -

'em, and that they are got through drinking sour m

The most striking feature of the proglottids was their size, being
greater in length, and especially in breadth, than that of any human
Taenia so far described. The measurements were as follows: —
Maximum, 32x9 mm.

Average, 28-6 x 8 5 mm.

Commonest, 29 x 10 mm.

Smallest, 21x6 mm.

bio. 2. Taenia bremneri, n. sp.

Gravid segments of (1) 7\ bremneri, (2) T. sag.nata, (3) T. solium,
(-U L tenne branches enlarged nearly four times

, , to ^ ln^ a sPec*men, the uterine segments were in some segments

forward4 ln,"urnber* not counting the “terminal” one which curves

arrantr*11 as four or five branches on it. The figure shows the

behind the" I:"6 UterUS' The &enital P°re is prominent and lies
behind the middle of the segment.

Lggs : Minimum, 34-2 x 30-4.
Commonest, 38 x 30-4.
Maximum, 45-6 x 41*8.

553

Calcareous bodies are numerous, measuring 1 5*2 /u in diameter.
This species is distinguished from some of the species found in man
by the following points : —

T. africatta. — Segments always broader than long.

T. confusa. — The gravid segments may be as long as 35 mm.,
but the width is only 5 mm., whereas in the present species, with a
length of 32 mm. the width is 9 mm.

T. saginata Gravid segments, 15 to 20, rarely 25 mm. long,
4 to '/ mm. wide.

T. solium. — Gravid segments, 10-12 long by 5 mm. broad.

1 hope soon to be in a position to describe complete specimens of
this worm; for the present I propose the name Taenia bremneri.

The type segments are in the museum of the Liverpool School of
Tropical Medicine.

III. A NEW LINGUATULID

( Porocephalus pattoni, n. sp.)

Habitat: — This linguatulid occurs in the lungs of the Indian rat
snake Zamenis v. Ptyas mucosus or Dhaman.

Diagnosis : — Body greyish white (spirit specimens), showing a
separation into head, neck and body. The head is globular and
flattened ventrally and rounded dorsally, and is separated from the
body by a fairly distinct narrow portion or neck. Of some dozen

Fig. 3. Porocephalus pattoni
(1) Hook, (2) mouth, (3) egg

specimens measured the longest was 1 1*5 centimetres, while the
shortest was 2^5 centimetres. The majority were between 6 an
centimetres. The number of rings varies somewhat, but an a\ erage
size specimen has 36. There are generally two rings on the lea
The body is cylindrical, ends bluntly, and the posterior end exhibits
some tortion. The thickness of the body is about 2’5 mm. n e

.

• •

■ ’■ .. . j . - it .

-J . ■ '' '

.... .

556

EXPLANATION OF PLATE L

x* Dibothnocephalus parvus, n. sp. Portion of strobila, actual
size.

^ ^ oroccphalus pattoni, n. sp. The Linguatulids in situ in the

lung of Zatnenis mucosus.

FlS- 3- Porocephalus pattoni , n. sp. Actual size.

PLATE L

Fig- 2

INDEX

INDEX

PAGE

Index of Authors . iii

General Index . iii

Index of Genera and Species new to Science .. ix

INDEX OF AUTHORS

PAGE

Brady, G. S. . .

. 423

Branch, C. W .

. 37 1

Breinl, A .

. 433. 439

Broden, A .

. 493

Carter, R. Markham .

. >55

Cuffey, E .

Dutton, J. E .

. 1, 199, 231, 285

Graham, W. M .

. 4l5

Hanington, J. W. B .

Jones, W. II. S .

. 529

Kinghom, A .

Looss, A

Moore, Benj .

. 161, 273

Neumann, G .

Newstead, R .

Nierenstein, M .

Rodhain, J .

Ross, E. 11 .

Ross, H. C. . . . .

Salvin-Moore, J. E .

Stephens, J. W. W .

Tobey, E. N .

Todd, J. L . i, 161,

van Loghem, J. J .

Walker, C. E .

PAGE

. »3

i, 169, 507, 554
. 161, 273

493

163

163

>99. 23>.

. 439

169, 549

. 285

273> 2S5

S23

483

GENERAL INDEX

Acarina

PAGE

• 96

Actinospharium cichorni , Chromidia

observed in .

Addenda et Corrigenda .

Aedtomyina .

Aedeomyia squamipetinis .

Amblyomma htbraum sflendidum .

„ variegatum .

,, ? variegatum .

Amphibia, Trypanosomes in .

Anatomy of the proboscis of biting

flies .

Anisocheleomyia quadrimaculata, n. sp.
Anophelinse in Congo Free State ...•••
Anophelines, Attempts to transmit

Trypanosomes by . . .

Anthomyia radicum in Liverpool . • •

Arthropoda, Insects and other, col¬
lected in the Congo

Free State . . .

„ Attempts to transmit I ry-

panosomes by various

blood-sucking . ; .

,, Protozoa found in various

Ascaris lumbricoides .

Asturinula monogrammica .

453

554

27

27

99

99

99

300

169

32

8

213

5*9

1

2*3

339

142

288

PAGE

Atoxyl, Effect of, followed by bi¬
chloride of mercury 16 1
,, followed by Dono¬
van’s solution . 282

„ Result of experiments with ... 275

A ttheyella africana , n. sp . . 422

A ty lotus nigromaculatus . . . . 44

Auchmeromyia luteola, bionomics of ... 50

Trypanosomes
in alimentary
canal of larva
of . 221

bed-bug ( Cimex sp.) .

Beetles in ashpit refuse . . . .

bi-chloride of mercury. Effect ot .

Birds, flat worms in . .

Protozoa observed in .

blood Films, Wet, method of preparing

and staining . •••••••

Modification of ITeiden-

hain’s method .

Borborus equinus breeding in stable

manure . . . . .

Bothriocephalid m man, A new .

Boycia, n. gen. . . .

Boyd a mitnomyiaformts , n. sp .

95

52°

161

>38

287

470

47 1

5*9

459

33

34

iv

PACK

Brady (G. S.) On Dr. Graham’s col¬
lection of Cvclopidae from the African

Gold Coast . 4^3

Branch (C. W.) Yaws . 371

Breinl (A. and others).

On the morphology and life
history of S pi roc hat a

duttoni . 43^

Cytology of Trypanosomes 439
Broden (A.) and Rodhain (J.) Contri¬
bution k I’etude de Porocephalus

moniliformis . 4gi

liufo regularia , Trypanosomes in . 307

Bush Fowl, Trypanosomes in . ^99

Bye a aisles buccinator (Trumpeter horn*

bill), Trypanosomes in .: . joy

Cal anus, sp .

Calliphora eyrthrocephala feeding on

human excreta . 507} qlg

' an ary fever. Observations on the
so-called

48 1

Canthocamptus, sp . 4.1

< nr ter ( R . Markham). On the presence
of Spirochata duttoni in the ova of

(’rni/hodoros moubata .

Oattle Trypanosomasis in Congo Free 3

.State .

C cilia pharoensis . .

Ccratopogon, sp . . g

Cercopithccus mono . .

.. schmidti

Cestodes, Two new human . . r'

ch&i?rr,pti°n. of iarva °f ••••• «

Clnronomus , sp. ... . 3°

C hr y sops dimidi at 'us ’ J ’

........ 4 j, 554

” a correction . er4

» stlacea . Z.

Ctmex lectularius . *54

Cimicidae . 95

aonVrh ngoUnsi% ^ypanosome's' in'
Clonorchts, n.g 0 pi sthorchiidarum .

.. end emuus (= Opisthorchis

sinensis of text books,

.. .

CSS"..in'"=,», dy'dioHim-, 148

Congo Floor 319

mit Trypano¬

somes by .... 21 a

” Structural details

T of larva Of ... 40

i rypanosomes in
alimentary

Congo Free State C?nal of - 221

c;^7P°da c?lle’cted in ...and °ther

Crocodile,' ^Trypano^s as.hPjt refuse.' .520

the^liver and ”

PACE

Cut ex a l bit arsis . 2I

„ annulioris . 2I

,, dissimilis . 2i

»» fatigans . . . 2i

,, laurenti, a. sp . . 24

„ luteolateralis . . . 22

„ met alii cut . 22

par , n. sp . 25

,, thalassius . 21

ti gripes . 22

.. tigripes, var. consimilis . . 23

,, viridis . . . 24

Culiridae . 8

Culicinse . i2

Cyclops , sp. ? . 421

bi color . 423

„ ? bicolor . 419

M leuckarti . 418, 420, 424

,, longistylis , n. sp . 419, 423

,, p heir rot us, . 420, 424

„ simillimus , u. sp . 48, 421, 424

„ varicoides, n. sp . 419, 423

„ virescens , n. sp . 420, 424

Cyclopidae from the Gold Coast,

Description of some . 415, 423

Cytamoebas in frogs . 334

Dejcania , sp., mimicing Glossina . 95

Dermanyssidae in lungs of monkey ... 97
Dermanyssus duttoni, Description of

larva of . 97

Dermatophilus ( sarcofsylla ) penetrans 93
Description of larva of 93

Diaptomus innominatus, n. sp . 423

Dibothriocephalus cordaius . 55'

>. lotus . 55‘

.1 pannes, n. sp. in man 549

Dicrocoelium lanccatum . . 125, 126

.. pancreatum, Syn. ... 128, 132

•> Syn. of Di stoma . '25

ihplo stoma agyptiacum, Syn . '34

Di stoma eras sum . 124

si coelomaticum . . . — >26

11 hepatis cndcmicum . 140, '42

9> 1. innocuum sive pernici-

osum . '4°

,, pancreaticum . , . 124

,, sinense . 141, 142, '43

Distomiasis of the liver and rectum

in man . v . 123, i37

Distomum spathulatum . '4*

Dogs, Protozoa observed in . 287

Drepanidia in Crocodiles . 3°5

91 sp . 3°3

Dutton (J. E. and others): —

,1 Insects and other Arthropoda

collected in the Congo Free

State . 1

.. Trypanosome transmission

experiments . *99

Cattle Trypanosomiasis in

Congo Free State . *3*

11 Concerning certain parasitic

protozoa observed in Africa 2S5

PACK

Dullonia, n. g . 17

„ africana , n. sp . 20

„ tarsalis, n. sp . 18

Eimeria schubtrgi . 45.2

Entomostraca, Description of some

Gold Coast . 415, 423

Eretmapodiics austenii . 14

„ i noma tut, n. sp . 12

Eughna . 452

Eurytrma, nov. gen. of Picrocoe-

liidarum . 127

„ cotlomaticum . 132

„ pancreaticum (Janson)

type of genus Kurytrema 128
Expedition to itie Congo, Seventh Pro¬
gress Report ... 201
„ ,, Eighth Interim

Report of . 287

Fasciola, ref. to genus . 124

Fasciolopsis buski . 123

„ Synonymy of . 124

Films, Methods of preparing ami

staining wet . 470

Fish, Trypanosomes in . 335

Flies as carriers of disease in man ... 485
Flukes in man, mammals and birds.. 123, 134
Fowls, Domestic, feeding on larvae and

pup$ of the house-fly . 509

Frambesia or Papillomatous Syphilide 400
l rogs and Toads, Trypanosomes in ... 306

Gastrodiscus aegypliacus, Syn. of . 134

si poly mast os , Syn . 134

ii second ns, n. sp., in mule 134

11 sonsinoi , Syn . 134

Glossma , Native names of, in Congo... 54
.. Further notes on the distri¬
bution of, in Congo Free

State . 74

is Reference to Koch 's experi¬
ments with . 226

» Trypanosomes in proboscis of 217
>! f 11 sea. Intervals between the

production of larvae

in . 69, 7‘

„ description of larva

and pupa . 71

„ Nocturnal habits of... 72

longi pal pis . : . 7 2

mac ul at a, n. sp . 73

morsitans . 7 1

pallidipes . 7 2

pal pal is . 57, 44b, 45°

description of larva . 57

description of pupa . 5^

distribution of in Congo Free

State . 59

habitats of . 60

habit of following man and
mammals . h2

» in native villages and dwell¬
ings . 64

11 bionomics of in captivity 64, 67-71
ii method of capture and keep¬
ing of in captivity . 68

PAGE

G/ossina, death rate of in captivity ... 69

,, length of life of in captivity 69

,, period of pupation . 69

,, feeding of in captivity . C9

,, used for the transmission of

Trypanosomes . 201

,, results of transmission of

T rypanosomes by inter¬
rupted feeding of . 206

,, results of transmission of

Trypanosomes by intervals
between feeding of ......... 207

Trypanosomes in proboscis

of .

,, Trypanosomes in alimentary

canal of .

,, Ref. to Trypanosomes found

in, by Koch . . .

„ Myxosporidium heibergi, (n.

sp.) in gut of .

G/ossina palpalis, var. wellmani .

Graham (W. A.) A description of some

Gold Coast Entomostraca .

Guinea Worm, Presence of, due to

Cyclops-host . . .

Ilaemamoeba relict a .

Haemaphysalis leachi .

Haemalopota , spp . - .

brunnipennis .

,, dutloni, n. sp .

„ trimaculata, n. sp .

,, native names for in Congo

217

218

227

342

71

4i5

4i7

225

99

43

43

41

42

Free State . 54

FLcemoprotcus . 225

Halteridi um ( lleemoproteus ) noctuez ... 293

Hanington (J- W. B. and others). Try¬
panosome transmission experiment ... 199

Hemiptera . . . 93

llippobosca equina . 9°

llippoboscidse . y . — 9°

Homalcmyia canicularis in stable

manure . 5*9

scalaris, larva of in ash¬
pit refuse, &c . 5*9

Hornbill, Trumpeter, Trypanosomes

observed in . 3°°

Horses infested with larvas of lronr-

bidium .

Hyalomma agyplium . .

llydrachnidae on Mosquitoes . •••;• •

Jlyla arborea, Trypanosomes found in

1 Sct^and^other Arthropoda collected

in the Congo Free State . .

Ixodidm, African species ol ■ —

|erome (T. S.). Transformation of

Tones" (W H^ST). Malaria and history...
L°nghom (A. aid others). Cattle Iry-
pansomiasis in the Congo Free State

Uucccyto«,o« £ . African

hawk .

in Dogs .

96

323

95

98

450

527

231

219

288

287

VI

PACK

Linguatulid, A new . 553

TJpoptena faradoxa , n. sp. . • . 9*

1 ooss (A.). On some parasites in the
'museum of the School of Tropical

Medicine Liverpool . ,-M

Lucilia, spp . 49

„ fuscina . 49

Lyperosia, sp . 53

„ minuta .... . 53

Lypcrosomum, Relation of to Dicro-

coelium . ,J5

Malaria and history . . . • . 5*7

„ influence upon Greek and

Roman History . 535

„ in Greece . . 53°

„ in Italy . ; . 53°

Mammals, Protozoa observed in, in

Africa . • • . . . 2°7

Mansonia uniformis . 2i}

,, ,, africanus . 27

Al argaropus annul at us . 100

tt „ calcaratus . too

M elaniconion rimum . *7

Melipona sp. causing intense annoyance

to man . 95

Mercury, Result of experiment with, on

Trypanosomes . 279

Mimomyia malfeyti, n. sp . 29

,, uniformis . .

Monkeys ( Cercopithecus , spp.) used in
experiments with transmission of

Trypanosomes . 2«7

Moore (Benj., and others). A note on
the therapeutics of Trypanosomiasis 161
Mosquito larvae, Automatic oiler for
destruction and prevention of, in

cesspools, &c . >63

Mosquitoes, Ectoparasites of . gb

„ in Congo Free State . 8

,, Protozoa found in . 339

Mud Fish (C/arias angolensis), Trypano¬
somes in . 335

Mule, Muscid larva of, in . Sq

Musca, spp . 5 3

Musca, A new genus and species of

blood-sucking . 53

Musca domeslica, On the habits, life-

cycle and breeding places of . 507

Muscid Fly, Larvae of in man, rat and

mule . 88, 89

Muscidae . 49

Myiasis, Cutaneous, in man . 88

>» ,, in the rat . SS

j. ,, in the mule . 89

Mymomyia africana, n. sp . 28

M yxosporidium heibergi, in Glossina ... 342

M yzomyia funesta . 9

,, pa/ud is , Protozoa found in 339

M ytorhynchus mauritianus . ' 10

>> paludis . 10

Nierenstein (M. and others):—

A note on the therapeutics of
Trypanosomiasis . 161

Page

Nierenstein (M. and others).—

„ Concerning the treatment of
experimental Trypano¬
somiasis . 273

Meomelaniconion pal pale, n. sp . F

Neumann 10.). Description of two new

species of African rfete . * *3

New stead (R. and others). Insects ana
other Arthropoda col¬
lected in the Congo free

State . 1

„ and Stephens (J. W. \V.).

The anatomy of the pro¬
boscis of biting flies, Pt.

11. Stomoxys . ,<J9

On Chrysops, spp. fiom the
Congo Free State

(Addenda) . . 554

„ On the habits, lile-cyde

and breeding places ol
the common house-fly

(Musca domeslica j . 5°5

Ochromyia anthropophaga in man ...... 5

Oiler, Automatic, for destruction ol

Mosquito larvae . 1 3

Ofisthorchit (Di stoma) tndtmtum as a

distinct sp . >47s

.. f el incus . 13

eeminus . J

„ sinensis . . . ;••• O6- I4'

,, denomination ot

two species of *47
Ornithodoros moubala . 98> '55- J‘3

” ” Sfo\ . >55

,, Attempts to trans¬
mit Trypano¬
somes by . ■ 1 3

Parasites, On some, in the Museum ot
the School of Tropical Medicine, ^

Liverpool . ^

Pediculidae . q4

Pediculus capitis . 04

,, vest i men/ i . . . . .

„ of Rat, Trypansomes in

alimentary canal of . ^

Phlebotornus, sp . ,'2o

Philonthus polities in ashpit refuse .

Pi ropl asm a bovi s . . " ‘ '

Pithecanthropus erect us, lemur of, sno -

ing evidence of ? Syphilis .

Plutarch on health . . . . . '

Pncumonyssus duttoni , description 01 q,

larva of . . . * . .54

Porocephalus annul dtus .

„ moniliformis. Contribu¬

tion a
P etude de 49

„ „ in ^acacU5' ,<?

sp- ; . ?

in various

•1 J>

verte- ,

brates - *

„ pattoni, n. sp . 1

vii

PACK

Proboscis of Biting Hies, Anatomy of... itx,

Protozoa in Arthropoda . . 34 *

„ Concerning parasitic,

observed in Africa . 287

„ Parasitic, in Africa . 285

Psyckoda ? fhalan.idts, breeding ill

human excreta . 520

Psychodidse . jb

Pulicidae . 93

Pycnosoma, sp . 49

„ tiara . i9

„ marginale . 49

„ futorium . 49

Pynlofhorus costalis . 8

„ costalis, Attempts to trans¬

mit Trypanosomes by 213
„ costalis , C'occidium sp. in

tissues of . 339

„ „ Protozoa found

in . 339

„ marshal l i . 9

Rana galamensis, Trypanosomes in . 307

„ mascariensis . . 307, 333

„ „ Trypanosomes in . 307

„ oxyrhynchus, Trypanosomes in ... 307

Rat, Muscid larva of, in . 88

Raffia marmorata , Trypansomes in ... 307

Relapsing fever, African, Parasite of ... 137

Reptiles, Trypanosomes in . . 3°3

Rhificefha/us, Table for the determina¬
tion of species of ... 119

„ bursa . 100

„ ca fen sis . loo

„ duttoni, n. sp . 115

„ longus, n. sp . 117

„ nitens . . io°

„ sanguineus . 100

>, simus . t°°

Rodhain (J.) and llroden (A.). Contri¬
bution a l’dtude de Porocephal us

moniliformis . 49'

Rhizopods, Chromidia observed in . 453

Ross (E. H.) and Boss (II. <\>. An
automatic oiler for the destruction
and prevention of mosquito larva; in
cesspools and other collections of

water . 163

Sahin-Moore (]. K.) and Breinl (A.).

The Cytology of the Trypanosomes . . . 439

Sand Flies . 3b, 37

it description of larva of . 37

i, Life-history of . 37

( Simulium , sp.). Attempts to
transmit Trypanosomes by 213
Sarcofhaga, spp. . . 48

Sarcophagidae . 4^

Sarcoptidae producing symbiotic mange

in rabbit . 97

Sarcofsylla fenelrans . 93

» „ description of

larva of . 93

Scatofhaga stercoraria breeding in cow
dung . 5*9

Simulida; . 36

Simulium, sp . 3b

,, damnosum . 36

,, 0 mat urn . 37

,, structural characters of larva

of . 37

„ habits of larva; of . 37

sp., Attempts to transmit

Trypanosomes by . 213

Snakes, Trypanosomes in . 3°3

S pirochaeta sp. in fish . 338

,, duttoni. Presence of in the

ova of Ornilho-
doros moubata ... 155
,, parasite of African

relapsing fever... 157
„ in the organs of

rats, Some notes
on the mor¬
phology of . 521

,, Morphology and

life-history of ... 4 33

„ gallinarum . 437

M jonesi from Clanas

a ngol crisis . 33^

obermeieri . 435

Stable Flies (Stomoxys, sp.). Anatomy

of proboscis of . 17I

Stegomyia albomarginata, n. sp . 10

argenteopunctata . J4

\\ fascia la . 14

luteocephala, n. sp . .

simpsoni . 14

Stephens (J. W. W.) and Newstead (R.)

The anatomy of the proboscis
of biting flies, Pt. II,

Stomoxys . . 1 9

Two new human Cestodes and

a Linguatulid . 549

Stomoxys, Anatomy of proboscis of...... *7»

Native names for in Congo

Free State .. . . . •••■" ^4

Trypanosomes in alimentary

canal of .

calcitrans . O

„ life history of . 7°

,, in Liverpool . 5*9

omega, n. sp . gj-

sitiens . . . . . F,'

Strychnine sulphate. «£>“£££*

somes . 277

followed by

” ” mercury . 279

Symbiotes communis cum cult . 97

Syphilide, The Frambesia OT 400

^stlndies; The "

- . g

viii

PAGE

Taenia africana . 553

„ bremneri , n. sp . 55 1

,, confusa . 553

,, saginata . . . 553

„ solium . 553

Tatniorhynchus annetti . 2b

„ aurites . 26

,, lenax . 26

Tabauidre . 41

Tabanus albovenlralis , n. sp . 46

,, billingtoni , n. sp . 46

,, canus . 44

,, dorsivitta . 44

fascial us . 14

,, gabonetisis . 45

m g rat us . 45

» far . 45

„ pluto . 45

„ ru fifes . 45

„ tarsalis . 45

, unimaculatus . 40

,, spp. incert . 48

,, Native names of in Congo Free

State . 54

licks. Description of two new species

of African . 113

,, Cist of species from Congo Free

State . 98

Toads and frogs, Trypanosomes in ... 306
Tobey (E. N. and others). Concerning
certain Parasitic Protozoa observed

in Africa . 285

Todd (J. C. and others): —

Insects and other Arthropoda
collected in the Congo Free

State . ,

A note on the Therapeutics of

Trypanosomiasis . 161

Trypanosome transmission

experiments . i99

Cattle Trypanosomiasis in

the Congo Free State . 231

Concerning the treatment of
experimental Trypano¬
somiasis . 2-~

Concerning certain Parasitic
7',,- ; Protozoa observer in Africa 4 ?q

Toxorhynchttes marsh alii i . ..

/ oxorhynchitime . """ . |(

Tortoises, Trypanosomes in . . .

rrtah°d^0n, S<T-r’ in the Muse>'in

Liv^ooi:h^1.°^^o?^..^ed:cine’

Irombidid* on horses at Kasongo 'oS

Trypanosoma brucei . 20,, x6,f 27s, 4ft

” ’• treatment of Rats

infected by . ,61

” Results of an experi-

. mental study on

therapeutics of 27c

” morphology and

A»/,,**multipllcation of 44i
” aimorfhon . 2_

^™"'’Mor.Pho,ogy and
multiplication of .

Trypanosoma equiferdum . 446

„ gambicnse . 201, 217, 441

„ „ in proboscis of

Glossina ... 217

„ „ morphology and

life-cycle ... 441

„ inopinalum 307, 321

,, karyoeeukton . 301

., lewisi 224, 225, 226

,, l orica turn . 223, 307, 306

„ „ description of 307

,, „ vel cost at um ... 306

„ mega . 3°7

,, noclua . 447

,, rotatorium . 323

,, sanguinis . 321 , 307

Trypanosome transmission experiments 199

Trypanosomes in proboscis of Glossina 217
„ Morphology of, in

alimentary canal of

Arthropoda .

,, in alimentary canal of

Glossina palpalis ... 218

,, in alimentary canal of

Pedi cuius, sp . 220

,, in alimentary canal of

Stomoxys . 221

,, in alimentary canal of

Auchtneromyia luteola 222
,, ingested bv insects ... 222

,, iu birds . 29"

,, in bush fowl . 299

,, in trumpeter hornbill— 300

,, in reptilia . 3°3

., in crocodiles . 3°5

., in tortoises . 3°5

in mud fish . 335

Cytology of the . . 439

,, Method of preparing

and staining . 47°

Trypanosomiasis, Note on the thera¬
peutics of . 1'11

,, in Cattle, Congo Free

State . 23'

,, The treatment of

experimental . 273

Tsetse Flies (See also Glossina).

„ „ Native names of in Congo

Free State . 54

,, ,, Field notes and bionomics

of

60

,, „ Observations on captive ••• $7

,, ,, in Congo Free State,

Distribution of, with

map . 5*h '4

,, ,, Protozoa (sp. non del.) in 34-’

Tsetse Fly, A new, from Tshumbiri ... 74

U ranotaenia balfouri .

Walker (C. E.). Observations on the

so-called Canary fever . 4‘>I

Worms, Flat, On some, in the Museum
of the School of Tropical Medicine,

Liverpool . 121

Yaws, The presence of Syphilis in the
West Indies . 37'

IX

INDEX OF GENERA AND SPECIES NEW TO SCIENCE

Protozoa

Myxesforidium heibergi, Dutton, Todd

and Tobey. Congo . 342

Sfirochaeta jonesi, Dutton, Todd and
Tobey. Congo . 338

Trematuda

Clonorchis , Looss . 148

Eurytrema, Looss . 127

Gastrodiscus sccundus , Looss. Assam 134

Cestoidea

Dibothriocephal us parvus, Stephens.

Tasmania . 549

Taenia bremneri, Stephens. Northern
Nigeria . 551

Entomostraca : Cyclopidac

Allheyella africana , Brady. West

Africa . A22

Cyclops longistylis, Brady. West

Africa . 4'9< 423

„ simillitnus, Brady. West

Africa . 421, 424

„ varicoides, Brady. West

Africa . 4>9i 423

1, virescens, Brady. West

Africa . 42°> 424

Diapiomus innominatus , Brady. West

Africa . 423

Ixodidac

Rhipicepkalus duttoni, Neumann. Congo 115

» longus, Neumann. Congo 117

LlNGUATULIDiE

Porocephalus pattoni, Stephens. India 553
Diptera

.4 nisoscheleomyia quadrimaculata ,

Newstead. Congo . 32

Boycia, Newstead . 33

mimomyiaformis, Newstead.

Congo . 34

Culex ti gripes consitnilis, Newstead.

Congo . 23

,, laurenti , Newstead. Congo . 24

,, par, Newstead. Congo . 25

Duttoni a, Newstead .

,, africana, Newstead. Congo 20

,, tarsal is, Newstead. Congo 18

Eretmapodites inornatus, Newstead.

Congo . 12

Glossina maculata , Newstead. Congo... 73
Haematopota duttoni, Newstead. Congo 41
,, trimaculata, Newstead.

Congo . 42

Li po plena paradoxa, Newstead. Congo 91
Mimomyia africana, Newstead. Congo 2b
„ malfeyti, Newstead.

Congo . 29

Neomelaniconion palpale, Newstead.

Congo

3'

Stegomyia albomarginala, Newstead.

Congo . ••• 0

leuteocephala, Newstead.

Congo .

S'tomoxvs omega, Newstead. Congo ... 37

Tabanus albovcntralis, Newstead. Congo 4
billingtoni, Newstead. Congo ^

#

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February i, 1907

Series T.M. Vol, I. No. 1

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Free State. Plates I-VI ... ... *1 .... . \

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J. W. VV. STEPHENS, M.D. (Cantab.), D.P.H.

Lecturer in Economic Entomology and Parasitology
R. N EYVSTEAD, A.L.S., F.E.S.

Professor of Comparative Pathology
H. E. AN NETT, M.D., D.P.H.

i a \ au in /a

VV. CARTER, M.D. , F.R.C.P. |

C. J. MACALISTER, M.D., M.R.C.P. Southern

LLOYD ROBERTS, M.D., F.R.C.S. ) P

Surgeons

W. ALEXANDER, M.D., F.R.C.S. ,

i\ . JONES, F. R . C. S. ( Edi.N. ) Royal Southern

G. P. NEW BOLT, M.B., F.R.C.S. ! Hospital

Director of 7 ropical Research at Runcorn
ANTON BREINL, M.U.Dr.. Prag
Assistant Lecturers

H. WOLFERSTAN THOMAS, M.D., C.M. McGill
AMO N 13 R E I N L, M.U.Dr. Prag

Demonstrators and Research Assistants

V tLL4,Nu1KwNGH0KN’ M-B. Toronto
IL'wN-,10^tV,A B ’ A M* M.D. Harvard
J- W. B. HANING 1 ON, M.D., C.M. McGill

Dean

Iroiessor SirRUBERT BOYCE, M.B., F.R.S.

,nLI, _ _ Laboratory

ON 1 ROPICAL LABORATORY, LIVERPOOL
UNIVERSITY
Research Laboratory
CROFTON LODGE, RUNCORN

Royal SOUTHERN HOSPITAL, LIVERPOOL

* LIVERPOOL

NOTICE

Regarding the Courses of Instruction proposed to be given by the
Liverpool School of Tropical Medicine, and the Examinations for the
Diploma of Tropical Medicine arranged to be held by the University
of Liverpool during 1907 (subject to such alteration as may hereinafter
be decided upon),

Lent Term begins January 15.

Lent Examination, March 25.

Summer Term begins May 1.

Summer Examination, July 15.

Autumn Term begins October 1.

Autumn Examination, December 16.

The full Course of Instruction is open to all qualified medical men,
and the examination to all students who have taken out this full
course.

l ee for the full Course of Instruction Ten guineas.

Fee for the Examination Five guineas.

Fee for the use of a School microscope during one term— len
shillings.

For prospectus and further information, application should be made
to the Dean of the Medical Faculty, University of Liverpool.

Hie following have obtained the Diploma in I ropical Medicine of
die University of Liverpool :

Diploma in L ropical Medicine

Hate of
Diploma

1906 Adie, Joseph Rosamond
'905 Anderson, Catherine Klmslie
190(1 Arnold, Frank Arthur
'9°4 Augustine, Henry Joshua
'906 Kate, John Brabant
1904 Bennett, Arthur King
*907 Bond, Ashton
>905 Brown, Alexander
‘9°4 Bruce, William James
'904 Bvrne, John Scott
'9°5 Caldwell, Thomas Cathcart
'9°ft Chisholm, Janies Alexander
*9°4 Clayton, Thomas Morrison
'9°S Critien, Attilio
'9°4 Dalziel, John McKwen
>9°4 Dee, Peter
1900 Dundas, James

1906 Faichnie, Norman

1907 Fell, Matthew Henry Gregson
*9°4 C.reenidge, Oliver Campbell
'9°4 Hehir, Patrick

■9°5 Hooton, Alfred
’9°5 Hudson, Charles Tilson
■9°5 lllington, Edmund Moritz
'9°ft Jeffreys, Herbert Castelman
,9°7 Kennan, Richard Henry
•9°7 Kenrick, William Hamilton

Dale of
Diploma

ig<u Khan, Saiduzzafor
Laurie, Robert

Le Eanu, George Ernest Hugh
Macfarlane, Robert Maxwell
Mackenzie, Donald l-rancis
Maclurkin, Alfred Robert ^
Maddock, Edward Cecil Gordon
McConnell, Robert Ernest
Moore, James Jackson
Nicholson, James Edward
Nightingale, Samuel Shore
Pailthorpe, Mary Elizabeth
Palmer, Harold Thornbury
Pearse, Albert
Philipson, Nicholas
Radcliffe, Percy Alexander Hurst
Sharman, Eric Harding
Taylor, William Irwin
Thomson, Frank Wyville
Tynan, Edward Joseph
Vallance, Hugh
Walker, George Francis Clegg
Watson, Cecil Francis
Willcocks, Roger Durant
WilTiamson, George Alexandei
Young, John Cameron

1 q«4
1907

■9° 5

1 90b

1904

1 905
1904

' 9°S
1 904

1 9°5

1 906
1906
1906
1904
'9«5
1904
1906
1904

1906

1907
1904
1906
1906
1906
<9°5

EDITORIAL NOTICE

By order of the Committee of the Incorporated Liverpool School
of Tropical Medicine, the series of the Reports of the School, which
have been issued since 1899, will he followed, from January i, 1907,
by the Annals of Tropical Medicine and Parasitology, of which
this is the second number.

The Annals are issued by the Committee of the School, and will
contain all such matter as was formerly printed in the Reports that
is to say, accounts of the various expeditions of the School and of the
scientific work done in its laboratories at the University of Liverpool
and at Runcorn. Altogether twenty-one Memoirs, besides other
works, have been published by the School since 1899, and of these
ten, containing 519 quarto or octavo pages and 95 plates and figures,
were published during the two years 1904 and 1905 ; and there is no
reason to suppose that this rate of production by the workers of the
School alone will diminish in the future. In addition, however, to
School work, original articles from outside on any subject connected
with Tropical Medicine or Hygiene may be published if found
suitable (see notice on back of cover); so that, in all probability, not
less than four numbers of the Annals will be issued annually. Eac 1
number will be brought out when material sufficient for it has een
accumulated.

CONTENTS

PAOf

ROSS, E. H., and ROSS, H. C. . -V '

\n Automatic Oiler for the Destruction and Prevention

of Mosquito Larvae in Cesspools and other sS

collections of water ... ... if}t

STEPHENS, J. W. W., and NEWSTEAD, R.

liie \natomy of the Proboscis of Biting Flies.

Part II, Stomoxys ... ... .

V.vl

'

Dili ON, J. E. ; 'I ODD, J. L. and HANINGTON, J. W. B.

I rypanosome Transmission Experiments . m

DU 1 l0N’ J‘ E ? T0DD» J- D., and KINGHORN, A.

Cat tic Trypanosomiasis in the Congo Free State . . . nt

MOORE, BENJ. j NIERENSTEIN, M., and TODD, j. L.

Concerning the Treatment of Experimental Trypano¬
somiasis

I j r' 1 . 1 HttM M I . 'jj*»

notice

Subscription: Ten Shillings c ' • ^ > Tj

payable m advance to the Secret* SjXPenCe per annum. P°s> free
Medicine, B i0 Exchange ag’ L,verp°<>I School of Tropica!

'■umbers al Six Shillings each ^r L,Verpo0,•

advert.sen.ents should also be addressed

source. Th°y it found su,tabie' fr°m

contrary is stated, will be underin ? tj'pewntten : and unless the
Auuals. Jbey should alway5 be ad? ‘° be olfer'd alone to these

Tropical Medicine and Parasitolnl ?! t0 the Editor- Annals of

1 ejected, the art.de will be returned*!’ University- Liverpool. If
posuo the address given by the authn ?°n 35 possible by registered
nobned to him and the article published f “ * “Z*#® U,e fact «® '■*

.. o ■»

i P

r-M

b *

its

UNIVERSITY OF LIVERPOOL

Kflvcmter 9, 1907

Series T.

.1 1

IVtRPGDLSCHODL
|0F TROPICAL
IrtPPICINE-na

ANNALS

Edited by

DONALD ROSS

,B Elaboration
with

)• K SnpmNS
* NtwntAn
J- L. Tom,
Thomas
Ahtoj, Butra
Su Rt««t Boyce

Annals

of

Tropical Medicine

and

Parasitology

Tttt 1 , ISSUED BY

; “00L SCHOOL OF TROPICAL MEDICIN1-;

““*• "uJKSLr * ««. m.

NORGATE, I4 HENRIETTA

Pnce Se*en Skilli

street, covent garden

lngs and Sixpence

net

Qlntiofaftone

SBRl'M I HKKAI’V

n.- : irom Brand Si rum* arc prepared

1 ! . ».i o! Research Labora?

- - v l- •! ! . «-h:< \\ was .n pioneer in the

i* *' ’i*" : in the liiitish Empire and

• • ’i 1 • 1 w«>nuined a foremost position.

i .ii H« ’A.. ,»• ,'iiw. ir Co. act as distributing
*. ,u 1 ' 1 i' "Jocu* prepared at these

I jhu4ionc%

H

' 1 "• \irmoVK Sf HiM CWelucomk’),
n, icoo (,1 UrlKh Retiring) units,

1 uni: : ) ■ 3000 units, ; 4000
,‘r •. tiV/t j.i -phials containing looo
a* tkfi'mg) in * c.r,, • <) ; 2000 units

• 1 o Ur>it\ in •, c.c., 8 ; 4000 units
1,1 . $c«0< units in y t « , 13 6; 6000

Socc unit in 8 c.c., 23/-;
••*'* 'o <* • . 27 jier phial. Asn-
1 «• (‘W ti t.co.MK.'), from

aMttnriicii oga.rnt . typical members of
” fWft IftcaiJi : of peritonitis

c : in pbiaU containing 35 c.c..,

' ' l1™ } but. ANTI-STREPTOCOC-

1 v iN: (‘WriLcoitf), from
^ ii « rut « iltures of streptococci

* - ’ 1 '■< **urccs, in the following

' r » mc^rikt Fkvkr, Piter

fl " R»m w»nc 1 r.vnt, SEPTICEMIA,

A l I OR At IV R hNnOCAR-
' 1 1 . AAlMn#, 1C .. 2/9; 25 c.c.,

• ' ■ I*' !‘ - • AWTI-OVSBHTERy

1s **-*1 * » '. m pi nuts containing 35 c.c.,

• • venom Serum

* • ' *x • J> j 1 (1 main mg 25 c.c,, 11/.:;

**• -** V k«u. Hosss Sm'M*
Vrui < ntammg 10 c.c, r/-:

. • ;I.IU

- U !<!*■

< »4> <** W.

•(Kalij phiaU

MrvllraJ Uiwy

BtCTMIOLOGt ANh ANALYSIS

m-ii t.nakcn bj Burroughs j
y A Ce c~ u 1 ;:o^ the essentials of
«•'« -,i iLf wnhr. th< fV4Ch of the bus)’

k • ^ uii rotynised. The

4 b caie*, here ilius-

r - b»rdi cornet form the
' • f * *„• . i'4 ucnaK 4 conducting

'*■ 1. »' nnr i-^aivT) and urine analysis

♦» < f TW .mi me and tight weight of
4 *M '-".i (<* bedside .use.

"i\* tin lUcimjoutaiCAL Case

m U--4 • «* - iNnc «iap)«fed bottles, con-
- j»i >4 ^ .H«t, 1 dun., absolute !

— »S A -».* <i »tAW water. t‘i dnn. A
1 *4 bcv.tr Canada Ktlsun, a

*n‘al' 1 WOi •** pair of forcq »s (small

• Tai« o< fr-.l'.tag l«.*r« epa, twelve rJ

microscopic slides, a spirit lamp, one metal case
of needles (straight No. 9), a glass funnel, two
watch-glasses, a packet of filter papers in parch¬
ment envelope, twelve blood collecting pipettes,

(P«H««e4)

X« K« Soidui* S.-»oJ B«.t:jnuuidjicit t»u
UtMortaneau 0 x 3!i x :« .a.

fifty cover slips, a sterile swab, and a glass rod
(for powdering ‘Soloid’ stains, etc.). One tube
each of the following ‘Soloid'Microscopic Stains: —
Methylene Blue, Eosin, Methyl Violet, Fuchsine,
Rotnanowsky Stain, Eosin-Methylene Blue,
Hasmatoxylin ( Delafield), and Toison Blood Fluid.
Nickel-plated metal case, fitted complete and
in doeskin cover, price 25/-.

‘Soloid- BRAND Urine Test Case.— Provides
the practitioner with a compact aseptic pocket-
case, containing the means of applying to urine
qualitative and quantitative tests for albumin,

(axi-.fi’K)

St. -*'A Sotom Br*ua Csis* Ci,‘

lie»fursai«eitl — &V X X in-

• etc. Made of nickel-plated metal, ^
ted. Contains a urinometer m na-
d case, Esbach’s albuminimeter gradu^
ure. pq#e, test-tub®: and stand, . .

, test-papers, analysis charts, Solo'd b<
lets of Fehlings Test, Indigo
Potassium Ferroc)-amde, and Cur

ilete in doeskin cover, price iV*

Leaflets scut giaiis on request

Burroughs Wellcome & Cl

LONDOK (E«C.), N*v VT0«,

Svpmr,

Cape Town

Volume I

November, 1907

No. 3

ANNALS

OF

TROPICAL MEDICINE AND
PARASITOLOGY

I

ISSUED BY

THE LIVERPOOL SCHOOL OF TROPICAL MEDICINE

Editor

Professor RONALD ROSS, Major I.M.S. (Ret.), D.P.H., F.R.C.S.,
D.Sc., LL.D., F.R.S., C.B.

In Collaboration with

J. W. W. STEPHENS, M.D., Cantab., D.P.H.

R. NEVVSTEAD, A.L.S., F.E.S.

J. L. TODD, B.A., M.D., C.M. McGill.

H. WOLFERSTAN THOMAS, M.D., C.M. McGill
ANTON BREINL, M.U.Dr.

Professor S.r ROBERT BOYCE, M.B., F.R.S.

C. Tinting <5- Co., Ltd.

Printers to the University Press of Liverpool
53 Victoria Street

THE INCORPORATED

LIVERPOOL SCHOOL OF TROPICAL MEDICINE

(Affiliated with the University of Liverpool)

Hon. President: Her Royal Highness PRINCESS CHRISTIAN
Ho*. Vice-PnsidetU ; The Duke of Northumberland, K.G.

COMMITTEE

Chairman: Sir ALFRED L. JONES, K.C.M.G.

Vice-Chairman: Mr. WILLIAM ADAMSON. President Royal Southern Hospital

Vice-Chancellor DALE University of Liverpool

Mr. W. B. Bowring Council of University of Liverpool

Dr. CATON J

Professor Sir RUBERT B I , - .

M.B., F.R.S. Senate of University of Liverpool

Professor SHERRINGTON, F.R.S.

Dr. W. Alexander
Professor Carter, M.D.

Mr. J. 0. Strafford
Mr. T. F. Harrison
Mr. Charles Livingston
Mr. A. R. Marshall
Mr. W. Roberts
Mr. Stanley Roger son
Mr. C. Booth (Jun.)

Mr. A. F. Warr
Mr. F. C. Danson

Mr. George BrocklehURST, Hon. Treasurer

' Royal Southern Hospital

I

Chamber of Commerce
1 Steamship Owners Association

1 Shipowners' Association

West African Trade Association

Mr. A. H. Milne,

BA., Hon. Secretary,

B io, Exchange Buildings, Liverpool

Staff

Professor

Major RONALD ROSS, C.B., F.R.S.. F.R.C.S., D.Sc.,
D.P.H. (Ind. Med. Serv. retired).

Lecturer

J. VV. VV. STEPHENS, M.D. (Cantab.), D.P.H.

Lecturer in Economic Entomology and Parasitology
R. NEWSTEAD, A.L.S., F.E.S.

Physicians

W. CARTER, M.D., F.R.C.P.

C. J. MACALISTER, M.D., M.R.C
LLOYD ROBERTS, M.D., F.R.C.

Surgeons

VV. ALEXANDER, M.D., F.R.C.S. i
R. (ONES, F.R.C.S. (EdiN.) [ Royal Southern

G. P. NEW BOLT, M.B., F.R.C.S. j hospital

Honorary Lecturers
H. E. AN NETT, M.D., D.P.H.

Professor of Comparative Pathology
W. T. PROUT, M.B., C.M.G.

C. E. WALKER, F.L.S.

,, _ Assistant Director of Cancer Research

Major C. L. WILLIAMS, M.D., D.P.H.

(Ind. Med. Serv. retired)

Runcorn Research Laboratories
Director : ANTON BREINL, M.U.Dr.. Pkag
Assistant : Vacant

.P

S.

Royal Southern
Hospital

Un Expeditions

wolferstan thomas, m.d., c.m.
ALLAN KINGHORN, M.B
R. E. MONTGOMERY, M R C V S
J- O. W. BARRATT, M.D. n Sc hond\
WARRINGTON YORKE, M.D.

Dean

Professor Sir ROBERT BOYCE, M.B., F.R.S.
Laboratory

J HNSTON TROPICAL LABORATORY, LIVERPOOL
UNIVERSITY

Research Laboratory
CROFTON LODGE, RUNCORN

Hospital

°YAL S°UTHERN- HOSPITAL, LIVERPOOL
• '*■ LIVER POOL

NOTICE

Regarding the Courses ol Instruction proposed to be given by the
Liverpool School of Tropical Medicine, and the Examinations for the
Diploma of Tropical Medicine arranged to be held by the University
of Liverpool during 1907 (subject to such alteration as may hereinafter
be decided upon),

Lent Term begins January 15.

Lent Examination, March 25.

Summer Term begins May 1.

Summer Examination, July 15.

Autumn Term begins October 1.

Autumn Examination, December 16.

The full Course of Instruction is open to all qualified medical men,
and the examination to all students who have taken out this full
course.

Fee for the full Course of Instruction I'en guineas.

Fee for the Examination Five guineas.

Fee for the use of a School microscope during one term — Ten
shillings.

For prospectus and further information, application should be made
to the Dean of the Medical Faculty, University of Liverpool.

The following have obtained the Diploma in Tropical Medicine of
the University of Liverpool :

Diploma in Tropical Medicine

Date of
Difloma

1906 Adie, Joseph Rosamond
>9°7 Allan, Alexander Smith
'907 Allwood, James Aldred
>905 Anderson, Catherine Elmslie
1906 Arnold, Frank Arthur
1904 Augustine, Henry Joshua
1906 Hate, John Rrabant

1904 Bennett, Arthur King

1906 Bennetts, Harold Graves

1907 Bond, Ashton
1907 Branch, Stanley

1905 Brown, Alexander
1904 Bruce, William James

1904 Byrne, John Scott

1905 Caldwell, Thomas Cathcart

1906 Carter, Robert Markham
1906 Chisholm, James Alexander

1904 Clayton, Thomas Morrison

1906 Clements, Robert William

1907 Collinson, Walter Julius

1905 Critien, Attilio

1904 Dalziel, John McEwen

1904 Dee, Peter

1907 Donaldson, Anson Scott

1906 Dundas, James

1906 Faichnie, Norman

1907 Fell, Matthew Henry Gregson
1907 Graham, James Drummond
1904 Greenidge, Oliver Campbell

1904 Hehir, Patrick

1907 iliscock, Robert Carroll

1905 Hooton, Alfred

1905 Hudson, Charles Tilson
190s Hlington, Edmund Moritz
190b Jeffreys, Herbert Castelman

Dale of
Diploma

1907 Kennan, Richard Henry
1907 Kenrick, William Hamilton
1904 Khan, Saiduzzafor

1904 Laurie, Robert

1907 Le Fanu, George Ernest Hugh

1905 Macfarlane, Robert Maxwell

1906 Mackenzie, Donald Francis

1904 Maclurkin, Alfred Robert

1905 Maddock, Edward Cecil Gordon

1907 Maddox, Ralph Henry
1907 McCarthy, John McDonald

1904 McConnell, Robert Ernest

1905 Moore, James Jackson

1904 Nicholson, James Edward

1905 Nightingale, Samuel Shore

1906 Pailthorpe, Mary Elizabeth
1906 Palmer, Harold Thornbury

1906 Pearse, Albert

1904 Philipson, Nicholas

,905 Radcliffe, Percy Alexander Hurst

1907 Ryan, Joseph Charles
1906 Sampey, Alexander William
1904 Sharman, Eric Harding
1906 Smithson, Arthur Ernest
1906 Taylor, Joseph van Someron
1906 Taylor, William Irwin

1904 Thomson, Frank Wyville

1906 Tynan, Edward Joseph

1907 Vallance, Hugh

1904 Walker, George Francis Clegg
1906 Watson, Cecil Francis

1906 Willcocks, Roger Durant
1906 Williamson, George Alexander

1905 Young, John Cameron

EDITORIAL NOTICE

By order of the Committee of the Incorporated Liverpool School
of Tropical Medicine, the series of the Reports of the School, which
have been issued since 1899, will be followed, from January 1, I9°7>
by the Annals of Tropical Medicine and Parasitology, of which
this is the third number.

The Annals are issued by the Committee of the School, and will
contain all such matter as was formerly printed in the Reports— that
is to say, accounts of the various expeditions of the School and of the
scientific work done in its laboratories at the University of Liverp
and at Runcorn. Altogether twenty-one Memoirs, besides other
works, have been published by the School since 1899, and of these
ten, containing 519 quarto or octavo pages and 95 plates and gures,
u,PrP nnKl.'eViMl ftnrma the t wo vears 1004 and 1905 ; and there is no

less than four numbers of th
number will be brought out
accumulated.

.

HOWARDS & SONS

In addition to the Salts of Quinine more commonly used,
manufacture the following Soluble Salts :

Percentage of the Solubility in
Alkaloid in the Salt Cold Water

Quinine Bihydrochlor - 72 o % 1

Rteomminried by medioal men with special experience In the
Tropics as THE BEST QUININE SALT for administration
when febrile symptoms aro presont.

Quinine Bihydrobrom
Quinine Lactate -
Quinine Hydrochlorosulph

N.B.— Compare with Quinine Sulph.

60 0 % 7

78-2 ° „ 10

74'3 % 2

73‘S % 800

HOWARDS' QUININE TABLETS are noted not

Salts contained, but also for their extremely accurate dosaee. This is an
important detail too often neglected, as experiments will prove.

STRATFORD, LONDON, E.

an

TEXT BOOK

The Practical Study
of Malaria and other
Blood Parasites

BY

J. w. W. STEPHENS, M.D. Cantab., D.P.H.

AND

S. R. CHRISTOPHERS, M.B. Vict., I.M-S.

Second Ed.tion, revised and enlarged, col^plare* and

to3 illustrations in the text. 8vo. Price i». 6d. nett.

WILLIAMS & NORGATE, 14 Henrietta Street, London, E.C.

CONTENTS

Kg*?1'- page

DUTTON, J. If., TODD, J. L., and TOBEY, E. N.

Concerning certain Parasitic Protozoa observed in

Africa... ... ... ... ... ... ... 285

BRANCH, C. W.

Yaws . . ... 371

GRAHAM, W. M.

A Description of some Gold Coast Entomostraca ... 415

BRADY, G. S.

Notes on Dr. Graham’s Collection of Cyclopidae from

the African Gold Coast ... ... v. G ... 423

BKEINL, A.

On the Morphology and Life History of Spirochada
duttoni

SALVIN-MOORE, J. E., and BREINL, A.

I he Cytology1 of the Trypanosomes

433

439

NOTICE

Subscription : I en Shillings and Sixpence per annum, post free,
payable in advance to the Secretary, Liverpool School of Tropical
Medicine, B 10 Exchange Buildings, Liverpool. Separate
numbers at least Six Shillings each. Correspondence concerning
advertisements should also be addressed to the Secretary.

Articles for Publication are accepted, if found suitable, from any
1 hey should, if possible, be typewritten; and unless the
contrary is stated, will be understood to be offered alone to these

t™?-5' 1 aV1!?- Sh°uId aIvvays be addressed to the Editor, Annals of
, P‘Cal MediClne and Parasitology, The University, Liverpool. If

dosMoi! tUl aiticle wdl be returned as soon as possible by registered
noLifi,>H ? • rCSb &iven by die auth°r. If it is accepted, the fact will be
° U'!! and the artlcle Published, if possible, in the forthcoming
“ r »The auth°r. is entitled to receive, free of charge, fifty
whpn K ° 1S,W°rk on its publication, provided that he asks for them

^pphefatcostprice^"' Addit‘°na! j§|$ if

UNIVERSITY OF LIVERPOOL

V.

| February 29, 1908

Scries T.M. Vol. I. No. 4

LiVERFGDLSCHODL
OP TROPICAL
/nePICINp-DD

ANNALS

Edited by
RONALD ROSS
in collaboration
with

J. \Y. W. Stephens
R. Newstead
J. L. Todd
H. W. Thomas
Aston Breinl
Sir Robert Boyce

Annals

of

Tropical Medicine

and

Parasitology

ISSUED BY

THE LIVERPOOL SCHOOL OF TROPICAL MEDICINE

I

LIVERPOOL : AT THE UNIVERSITY ^ ^

uTTxrn IFTTA STREET, COVENT GAKUE-in
LONDON: WILLIAMS ft NORGATE, 14 HLNR

Price Seven Shillings and Sixpence net

Tisf ‘ T A B L OI D ’ B«nd

PLEATED COMPRESSED

DRESSINGS

Originated and introduced

by B. W. & Co., they

are made of materials

of the finest quality and
are unique in compact-

• ness and portability.

[oa»hMit *otuai Although occupying a

minimum of space they can be unfolded as easily
as the ordinary bulky Dressings.

STERILISED DRESSINGS

A further important advance originated by
B. W. & Co. is the issue of these Dressings —

Sterilised. Each carefully sterilised and enclosed in
a sterilised impervious covering.

ABRIDGED LIST
Bandages, ‘Tabloid’

ove, flannel, and Triangular (Esmarch’s Pictorial)
Absorbent Cotton Wool, ‘Tabloid’ ...ho

Plain, Boric, Double Cyanide 3%, and Iodoform
Absorbent Cotton Wool between Gauze, ‘Tabloid’ .....

Absorbent Gauze, ‘Tabloid’ ...„0

Plain, Boric, Double Cyanide 3%, Iodoform, and Sal Alembroth 1%
Ltnt, ‘Tabloid’

Plain, Boric, and Carbolised
Tow. ‘Tabloid’ . . . n 0
Carbolised

When ordering, please specify STERILISED, if so dssirsd
See B. W. & Co.’s Price List for prices and details

Burroughs Wellcome & Co., London (Eng.)

rG”Ckti • NEW YOR K AlONTREAL SYDNEY CAPE TQWN

BANDAGES an

opa..o« iaadije • Tabloid' Bandage

F 306

[cormam

Volume I

February, 1908

No. 4

ANNALS

OF

TROPICAL MEDICINE AND
PARASITOLOGY

ISSUED BY

THE LIVERPOOL SCHOOL OF TROPICAL MEDICINE

Editor

Professor RONALD ROSS, Major I.M.S. (Ret.), D.P.H., F.R.C.S
D.Sc., LL.D., F.R.S., C.B.

In Collaboration ■ with

j. W. W. STEPHENS, M.D., Cantab., D.P.H.

R. NEWSTEAD, A.L.S., F.E.S.

J. L. TODD, B.A., M.D., C.M. McGill.

H. WOLFERSTAN THOMAS, M.D., C.M. McGill
ANTON BREINL, M.U.Dr.

Professor S,r RUBERT BOYCE, MB., F.R.S.

&

APR

19

1909

C. Tinting <5r> Co., Ltd.

Printers to the University Press of Liverpool
53 Victoria Street

THE INCORPORATED

LIVERPOOL SCHOOL OF TROPICAL MEDICINE

(Affiliated with the University of Liverpool)

Hon. President : Her Royal Highness PRINCESS CHRISTIAN
Hon. Vice-Presidents : The Duke OF NORTHUMBERLAND, K.G.
Earl Cromer, G.C.B.

COMMITTEE

Chairman : Sir ALFRED L. JONES, K.C.M.G.

Vice-Chairman : Mr. WILLIAM ADAMSON, President Royal Southern Hospital

Hon. Treasurer

Vice-Chancellor Dale
Sir W. B. Bowring, Bart.

Dr. CATON

Professor Sir RUBERT BOYCE,

M.B., F.R.S.

Professor SHERRINGTON, F.R.S.

Dr. W. Alexander

Dr. C. J. Macalister

Mr. J. O. Strafford

Mr. T. F. Harrison

Mr. Charles Livingston

Mr. A. R. Marshall

Mr. W. Roberts

Mr. Stanley Rogerson

Mr. C. Booth (Jun.)

Mr. A. F. Warr
Mr. F. C. Danson
Mr. George Brocklehurst
Mr. J. W. W. Danson

Lord Stanley, K.C.V.O.

University of Liverpool
Council of University of Liverpool

Senate of University of Liverpool

Royal Southern Llosjntal
Chamber of Commerce
j Steamship Owners' Association

Shipowners' Association

]\’cst African Trade Association

)

Mr A. H. Milne, B.A., Secretary,

B io. Exchange Buildings.

Liverpool

Staff

Professor

Major RONALD ROSS, C.B., F.R.S., F.R.C.S., D.Sc.
D.P.H. (Ind. Med. Serv. retired).

Lecturer

J. YV. YV. STEPHENS, M.D. (Cantab.), D.P.H.

Lecturer in Economic Entomology and Parasitology
R. NEWSTEAD, A.L.S., F.E.S.

Physicians

Major RONALD ROSS, C.B., F.R.S., etc. )

C. J. MACALISTER, M.D., M.R.C.P. s

LLOYD ROBERTS, M.D., F.R.C.S. )

Surgeons

YV. ALEXANDER, M.D.', F.R.C.S. |

R. JONES, F.R.C.S. (EdiN.) Royal South*™

G. I>. NEWBOLT, M.B., F.R.C.S. j Hosf,i,u

Honorary Lecturers
H. E. AN NETT, M.D., D.P.H.

... Professor of Comparative Pathology

W. 1. PROUT, M.B., C.M.G
C. E. WALKER, F.L.S.

Assistant Director of Cancer Research

Major C. L. WILLIAMS, M.I)., D.P.H.

(Ind. Med. Serv. retii ud t

Runcorn Research Laboratories

Rrs : ANT0N BREINL. M.U.Dr.. Pkao
Hull Demom^rator: M. NIERENSTEIN. Ph P
kesearch Assistant : ED. HINDLE, A.R.C.S. (Lond)-

On Expeditions

Vi tVamL£?RSTAN ™OMAS, M.D., C..M.
ALLAN KINGHORN, M.B.

L- MONTGOMERY, M.R.C.V.S.
Wa»JX’xt2^RRATT’ AI D-. D.Sc. (Lond.)

YVARRINGTON YORKE, M.D.

Dean

Rofessor SirRUBERT BOYCE, M.B., F.R.S-

JOHNSTON TROPICAl'laboratORY, L1VER^oL
UNIVERSITY

Research Laboratory
CROFTON LODGE, RUNCORN

Hospital

°VAL SOUTHERN HOSPITAL, LIVERPO^1'

,, Secretary's Office

EXCHANGE BUILDINGS. LIVERPOOL

NOTICE

Regarding the Courses oi Instruction proposed to be given by the
Liverpool School of 1 ropical Medicine, and the Examinations for the
Diploma of Tropical Medicine arranged to be held by the University
of Liverpool during 1908 (subject to such alteration as may hereinafter
be decided upon),

Lent Term begins January 14.

Lent Examination, March 23.
Summer Term begins May 1.
Summer Examination, July 13.
Autumn Term begins October 1.
Autumn Examination, December 14

The full Course of Instruction is open to all qualified medical men,
and the examination to all students who have taken out this full
course.

Fee for the full Course of Instruction Ten guineas.

Fee for the Examination Five guineas.

Fee for the use of a School microscope during one term — Ten
shillings.

For prospectus and further information, application should be made
to the Dean of the Medical Faculty, l ni versify of Liverpool.

The following have obtained the Diploma in I ropical Medicine of
the University of Liverpool :

Diploma in Tropical Medicine

Date of
Diploma

1906 Atlie, Joseph Rosamond

1907 Allan, Alexander Smith
1907 Allwood, James Aldred

1905 Anderson, Catherine Klmslie

1906 Arnold, Frank Arthur
1904 Augustine, Henry Joshua
1906 Bate, John Brabant

1904 Bennett, Arthur King

1906 Bennetts, Harold Graves

1907 Bond, Ashton
1907 Branch, Stanley

1905 Brown, Alexander
1904 Bruce, William James

1904 Byrne, John Scott

1905 Caldwell, Thomas Cathcart

IfinA f‘or»or nKort \f arlfhaill

Date of
Diploma

1007 Keane, Joseph Gerald
1907 Kennan, Richard Henry
1907 Kenrick, William Hamilton
1904 Khan, Saiduzzafor

Keane, Joseph Gerald
Kennan, Richard Henry

isennaii, muuuu **„.*._,

Kenrick, William Hamilton
Khan, Saiduzzafor

Mackenzie, Donald Francis

n.)ot) Mackenzie, Donald Francis

Cd I U <V C 1 1 1 I 110111(1'

(*artr.r Marlfh.'llll

1907 Collinson, Walter Julius

1905 Critien, Attilio

1904 Dalziel, John Me K wen
1907 Davey, John Bernard
1904 Dee, Peter
1907 Donaldson, Anson Scott

1906 Dundas, James

1906 Faichnie. Norman

1907 Fell, Matthew Henry Gregson
1907 Gann, Thomas William I rancis
1907 Graham, James Drummond
1904 Greenidce, Oliver Campbell

1904 Hehir, Patrick

1907 Iliscock, Robert Carroll

1905 Hooton, Alfred

1905 Hudson, Charles Tilson

1905 Illington, Edmund Moritz

1906 Jeffreys, Herbert Castelman

Dalziel, John McEwe
Davey, John Bernard

.

EDITORIAL NOTICE

By order of the Committee of the Incorporated Liverpool School
of Tropical Medicine, the series of the Reports of the School, which
have been issued since 1899, will be followed, from January 1, 1907,
by the Annals of Tropical Medicine and Parasitology, of which
this is the fourth number.

The Annals are issued by the Committee of the School, and will
contain all such matter as was formerly printed in the Reports that
is to say, accounts of the various expeditions of the School and of the
scientific work done in its laboratories at the I diversity of Liverpool
and at Runcorn. Altogether twenty-one Memoirs, besides other
works, have been published by the School since 1899, and of these
ten, containing 519 quarto or octavo pages and 95 plates and figures,
were published during the two years 1904 and 1905 ; and there is no
reason to suppose that this rate of production by the workers of the
School alone will diminish in the future. In addition, however, to
School work, original articles from outside on any subject connecte
with Tropical Medicine or Hygiene may be publishe 1
suitable (see notice Oft back of cover); so that, in all pro a 1 ity, n
less than four numbers of the Annals will be issued annua y
number will be brought out when material sufficient for 1 ias

accumulated.

HOWARDS & SONS

In addition to the Salts of Quinine more commonly used,
manufacture the following Soluble Salts :

Percentage of the Solubility in

Alkaloid in the Salt Cold Water

Quinine Bihydrochlor - 72 ’o %

Recommended by medical men with special oxporionco in the
Tropics as THt DMT QUININE 8 ACT for administration
when febrilo symptoms oro prceont.

Quinine Bihydrobrom
Quinine Lactate
Quinine Hydrochlorosulph

N.B. —Compare with Quinine Sulph.

60 0 %
78-2 %
74 '3 %

I

7

10

2

800

HOWARDS’ QUININE TABLETS are noted not only for the Purity of the
Salts contained, but also for their extremely accurate dosage. Tins is an
important detail too often neglected, as experiments will prove.

STRATFORD, LONDON, E.

JUST PUBLISHED

TEXT BOOK

The Practical Study
of Malaria and other
Blood Parasites

BY

J. w. W. STEPHENS, M.D. Cantab., D.P.H.

AND

S. R. CHRISTOPHERS, M.B. Vict., I.M.S.

Third Edition, thoroughly revised, with^ai^ohiured

128 illustrations in the text.

WILLIAMS & NORGATE, 14 Henrietta Street, Lo

CONTENTS

WALKER, C. E.

Observations on the so-called ‘Canary Fever'
BRODE.N, A., et RODHAIN, J.

Contribution a 1'Etude de Porocephalus moniliformis

NEWSTEAD, R.

On the Habits, Life-Cycle and Breeding Places of the
Common House-Fly {Musca domestica , Linn.)

LOGHEM, J. j. van

Some Notes on the Morphology of Spiradurta dultoni
in the Organs of Rats .

JONES, W. H. S.

Malaria and History .

STEPHENS, J. W. W.

fwo New Human Cestodes and a New Linguatulid ...

PACE

4^3

493

507

523

529

549

NOTICE

Subscription: One Guinea per annum, post free, payable in
advance to the Secretary, Liverpool School of Tropical Medicine,
B 10 Exchange Buildings, Liverpool. Separate numbers at least
Shillings each. Correspondence concerning advertisements
•should also be addressed to the Secretary.

Articles for Publication arc accepted, if found suitable, from any
I hey should, if possible, be typewritten ; and unless the
uontrarj ,s stated, will be understood to be offered alone to these
Annals. They should always be addressed to the Editor, Annals of
Tropical Medicine and Parasitology, The University, Liverpool. If
rejected, the article will be returned as soon as possible by registered
post to the address given by the author. If it is accepted, the fact will be
nutjhed to Imn and the article published, if possible, in the forthcoming
number. 1 he author is entitled to receive, free of charge, fifty
icpnnls of his work on its publication, provided that he asks for them

"hen he sends his paper. Additional copies, if required, will be
supplied at cost price.

4

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