Rabies in southern Africa

Onde s epoo Jou nal
o
Ve e ina y Resea ch, 60:325-346 (1993)
Rabies
in
sou he n A ica
R.
SWANEPOEL1, B.J.H. BARNARD2, C.D. MEREDITH2, G.C. BISHOP3
G.K. BRUCKNER4, C.M. FOGGIN5
and
O.J.B. HUBSCHLE6
ABSTRACT
SWANEPOEL,
R.
, BARNARD, B.J.H., MEREDITH, C. D
.,
BISHOP, G., BRUCKNER,
GK
, FOGG
IN
, C.M.
& HUBSCHLE, O.J.B. 1993. Rabies
in
sou he n A ica. Onde s epoo Jou nal
o
Ve e ina y Resea ch,
60:325-346
The i s con i med ou b eak o abies
in
A ica, belie ed
o
ha e ollowed he impo a ion o an
in
ec ed
dog om England
in
1892, occu ed
in
he eas e n Cape P o ince o Sou h A ica, and was b ough unde
con ol
in
1894.
An
uncon i med epidemic o abies
in
dogs occu ed
in
wes e n Zambia
in
1901
.
By
he
ollowing yea he disease had appa en ly sp ead along a majo ade ou e, o cause
an
ou b eak
in
Zimbabwe which engul ed mos o he coun y be o e being e adica ed
in
1913. The exis ence o endemic
abies o i e ids (mongooses and gene s) was con i med
in
Sou h A ica
in
1928, and since hen he
i e id disease has con inued o occu widely on he in e io pla eau o he coun y wi h spill-o e o
in ec ion
o
ca le and a a ie y o o he animals. F om abou 1947 onwa ds, an in asi e o m o dog
abies sp ead om sou he n Zambia and/o Angola in o Namibia, ac oss no he n and eas e n Bo swana
in o Zimbabwe and he no he n T ans aal by 1950, en e ed Mozambique
in
1952, and sp ead om he e
o Swaziland
in
1954. Dog abies ex ended om sou he n Mozambique in o Na al
in
1961
o
cause a
majo epidemic which was b ough unde con ol
in
1968. The disease
e
-en e ed no he n Na al om
Mozambique
in
1976 and since hen dog abies has p o ed di icul o con ol
in
he pe i-u ban se lemen s
o Na ai-KwaZulu. The disease sp ead om Na al o Leso ho
in
1982, and in o he T anskei egion o he
eas e n Cape P o ince
in
1987, o each he Ciskei by 1990. The sp ead o he disease
in
dogs was
ollowed by he eme gence o abies o jackals and ca le
in
cen al Namibia, no he n Bo swana, Zimbab-
we and he no he n T ans aal. A unique ou b eak o abies
in
kudu an elope occu ed
in
cen al Namibia
om 1977
o
1985, appa en ly in ol ing o al sp ead o in ec ion be ween indi iduals. A ew cases o abies
in
he ba -ea ed ox we e ecognized each yea
in
Namibia om 1967 onwa ds, and om he 1970s he
occu ence o he disease
in
he ox has eme ged as a dis inc p oblem
in
he no he n Cape P o ince
and sp ead o he wes coas . The abies- ela ed i uses, Lagos ba , Mokola and Du enhage, associa ed
wi h ba s, sh ews and oden s
in
A ica, a e known o ha e caused isola ed cases o disease
in
Sou h A -
ica, and
on
one occasion a small ou b eak in ol ing
si
x ca s and a dog in Bulawayo, Zimbabwe. How-
e e , he esul s o monoclonal an ibody es s on nume ous specimens indica e ha he abies- ela ed
i uses a e no a majo cause o disease
in
sou he n A ica.
INTRODUCTION
1 Na ional Ins i u e o Vi ology, P i a e Bag X4, Sand ingham,
2131
Sou h A ica
2 Onde s epoo Ve e ina y Ins i u e, P i a e Bag X5, Onde s e-
poo , 0110 Sou h A ica
3 Regional Ve e ina y Labo a o y, Alle on, P i a e Bag X2,
Cascades, 3200 Sou h A ica
4 Di ec o a e o Animal Heal h, P i a e Bag X138, P e o ia,
0001
Sou h A ica
5 Ve e ina y Resea ch Labo a o y, Causeway, Ha a e, Zi
m-
babwe
6 Cen al Ve e ina y Labo a o y, Windhoek, Namibia
Rabies ( abidus,
L.
= mad)
is
a highly a al ne ous
disease o humans and all o he wa m-blooded e -
eb a es, caused by a i us which is p esen
in
sali a
la e
in
in ec ion and which is gene ally ansmi ed
by he bi e o diseased animals, mos commonly
dogs and o he ca ni o es. The causa i e agen o
he disease is a membe o he Rhabdo i idae amily
o od o bulle -shaped i uses ( habdos, G . = od),
which ha e a single-s anded, nega i e-sense RNA
genome (complemen a y o mRNA). Wi hin he amily,
325
Rabies
in
so
u he
n
A
i
ca
abies i us is placed in he genus Lyssa i us
(l
yssa,
G .=
age o u y), while
he
amily also includes he
Epheme a i us (bo ine epheme al e e and ela ed
i uses) and Vesiculo i us ( esicula s oma i is and
ela ed i uses) gene a o animal i uses, plus ce -
ain i uses
o ishes,
in e eb a es, and plan s (Cali-
she , Ka aba sos, Zelle , Digou e, Tesh, Shope, T a-
assos Da Rosa & S Geo ge 1989;
Anonymous
1993). The lyssa i uses include abies i us (desig-
na ed lyssa i us se o ype
1)
and he so-called abies-
ela ed i uses, Lagos ba , Mokola and Du enhage
(lyssa i us se o ypes 2, 3 and
4)
which a e associa -
ed wi h ba s, sh ews and oden s in A ica, plus se -
en o he i uses which a e o unknown e e ina y o
medical signi icance (Calishe e
a/
. 1989). Rabies-
ela ed i uses ha e also been isola ed om ba s in
Eu ope, and hese ha e been p o isionally designa -
ed Eu opean ba lyssa i uses 1 (EBL
1)
and 2 (EBL
2)
, bu i is as ye unce ain whe he hese a e o be
ega ded as sub ypes o Du enhage i us o ecog-
nized as sepa a e se o ypes (King 1991
).
I should
be no ed ha despi e ea ly epo s o
he
con a y,
in ec ion o ba s wi h abies i us p ope (lyssa i us
1)
has been con i med only in he Ame icas, whe e
he disease occu s
in
bo h non-haema ophagous and
ampi e ba s (Bae 1975a; 1975b; Be an 1981 ; Smi h
& Bae 1988).
I is o g ea epidemiological signi icance ha ecen
monoclonal an ibody and nucleic acid s udies ha e
shown ha s ains o abies i us (lyssa i us
1)
which
ci cula e in pa icula hos species wi hin gi en geo-
g aphic egions, end o unde go gene ic adap a ion,
esul ing in he de elopmen o so-called bio ypes,
wi h sub le changes in an igenici y and pa hogenici y
(Wik o & Kop owski 1978; Wik o , Flamand & Kop-
owski 1980; Blancou 1988a; Smi h & Bae 1988;
Smi h 1989; Sac amen o, Bou hy & To do 1991;
Smi h, Fishbein, Rupp ech & Cla k 1991 ).
The
bio-
ypes a e uniquely adap ed o ci cula e in speci ic
animals, so ha wi hin an a ec ed a ea he disease
is mani es ed p edominan ly by a ·single hos species,
o less commonly mo e han one, and his
same
hos
appea s o be esponsible o main enance and
sp ead o he i us; disease in o he animals ep e-
sen s spill-o e o in ec ion esul ing om spo adic
con ac wi h he majo hos species (Smi h & Bae
1988; Smi h 1989).
Rabies is widely dis ibu ed in he wo ld, wi h only
ce ain coun ies, mainly islands and peninsulas, be-
ing his o ically ee o he disease o ha ing suc-
ceeded in e adica ing i in ecen imes (Biancou
1988b;
WHO
1989; 1991). I is es ima ed ha he e
a e a leas 25 000 human cases o he disease each
yea , o e
90%
o which esul om ansmission by
dogs (Fe nandes & A ambula 1985; Bagel & Mo sch-
wille 1986). The de eloped na ions o wes e n Eu -
ope and No h Ame ica ha e succeeded
in
d as ically
326
educing he occu ence
o
he disease in
dogs
(u -
ban abies) o e he pas ou decades, bu i onically
hey
ha e expe ienced a ma ked inc ease in
he
dis-
ease o wild e eb a es (syl a ic abies)
o e
he
same pe iod (Smi h & Bae 1988; Blancou
1988b
;
WHO
1991). Dog abies p edomina es in
mos
o
he
de eloping na ions
o
Cen al and Sou h
Ame ica
,
Asia and A ica, and human disease is i e. Syl a ic
abies, on
he
o he hand, is eco ded ela i ely in e-
quen ly in mos o he de eloping na ions,
and
his
is asc ibed pa ly o de icien moni o ing o he disease
in wild e eb a es.
La gely as a esul
o
lack o esou ces, he occu -
ence o sociopoli ical uphea als,
and
he exis ence
o o he na ional p io i ies, abies appea s o be leas
well moni o ed in he con inen
o
A ica (Biancou
1988b;
WHO
1991 ;
Swanepoel1994)
. The disease
has been p esen in No h A ica since an iqui y,
whe e i occu s p incipally as u ban abies (Biancou
1988b).
In
sub-Saha an A ica, whe e
humans
and
o he animals a e mo e widely dis ibu ed han in
no he n A ica, he e has been a g ea e
endency
o epidemics
o
dog abies o sp ead o e la ge a e-
as and o he disease o be obse ed in domes ic
he bi o es and wild e eb a es (Biancou 1988b).
This end is mos no iceable in
he
mo e de eloped
coun ies
o
sou he nmos A ica, whe e
he
high
p opo ions o cases eco ded in wild animals
mus
o
some
ex en e lec mo e in ensi e moni o ing o
he disease. Ne e heless, i is clea om· he his o y
o he disease in he sub-con inen (Swanepoel
1994), p esen ed he e
in
abb e ia ed o m, ha sou h-
e n A ica has a unique blend
o
u ban and syl a ic
abies.
ZAMBIA AND ANGOLA
Rabies
was
appa en ly p esen in
Zambia
du ing he
19 h cen u y, and in 1901 Chie Lewanika o
he
Ba-
o se in he
wes
o
he coun y o de ed
he
des uc-
ion o all dogs in he a ea in an a emp o con ol
a se e e ou b eak o he disease (Edmonds 1922;
Snyman 1940; Shone 1962). The diagnosis o he
disease was i s con i med in 1913 and abies has
con inued o occu h oughou
he
coun y (Zyambo,
Sinyangwe & Bussein 1985; Tuchili 1988; Sinyangwe
1992).
The
disease a ec s
ma
inly dogs, bu
app ec
i-
able numbe s o ca le a e a ec ed in he sou h-cen-
al pa o he coun y, pa icula ly in loca ions
whe e
jackal abies is diagnosed close o na u e ese es
(Zyambo
e a/
. 1985). Rabies
was
i s con i med in
Angola in 1929 and since hen he disease
has
been
diagnosed mainly in dogs, wi h e y
ew
cases being
eco ded in o he dqmes ic o wild animals (Dua e,
Rosliaco , Nsalambi &
Gomes
1985), bu p o ac ed
ci il wa in he coun y has hampe ed moni o ing and
con ol o he disease o e he pas
ew
decades
.
NAMIBIA
An ou b eak o disease i ing he desc ip ion o
a
-
bies and
in
ol ing dogs, ca le and small li es ock
was appa en ly obse ed
in
Namibia
in
1887 (Schnei-
de 1985; Hubschle 1988), and om 1925 onwa ds
he e we e spo adic epo s o ou b eaks o disease
in ol ing dogs, humans and, on one occasion a hy-
ena, in he O ambo, Ka ango and Cap i i S ip dis-
ic s in he no h, bo de ing Angola and Zambia, wi h
an isola ed case being eco ded u he sou h
in
a
child bi en by a dog
in
Swakopmund
in
1928 and
ano he
in
a woman bi en by a wild ca
in
G oo -
on ein
in
1937 (Von Mal i z 1950). A diagnosis o a-
bies was inally con i med
in
a dog om Rundu
in
Ka ango
in
1938, bu a suspec ed ou b eak o he
disease which occu ed on a ms
in
he Gibeon and
Ma ien hal dis ic s
in
he sou h
in
1945 could no
be
con i med (Von Mal i z 1950). The posi ion changed
sha ply wi h he occu ence o he second con i med
case
o
abies
in
194 7, also
in
a dog
in
Rundu,
which was ollowed
in
1948 by he appea ance o
he disease sou h o he E osha Na ional Pa k
in
ca -
le
in
Ou jou dis ic , whence sp ead o disease in-
ol ing black-backed jackals (Canis mesomelas) and
ca le con inued sou hwa ds o each O jiwa ongo
in
1949 and he cen al dis ic s o Gobabis and Wind-
hoek by
1951
(Onde s epoo Ve e ina y Ins i u e
[OVI) 1932-1992, unpublished eco ds; Von Mal i z
1950; Alexande 1952; Schneide 1985}. F om 1967
onwa ds a ew cases o he disease, less han en,
we e eco ded egula ly each yea
in
ba -ea ed oxes
(O ocyon mega/o is) (OVI 1932-1992, unpublished
eco ds). Rabies has emained a p oblem
in
Namib-
ia,
wi h dog and human cases being eco ded mainly
in
he no h whe e he densi y o he u al popula ion
is g ea es , jackal and ca le abies domina ing
in
he
cen al anching a ea, and spo adic disease being
associa ed mainly wi h elids (A ican wild ca s and
ca acals) and i e ids (gene s and mongooses)
in
he sheep- ea ing a eas o he sou h (OVI1932-1992,
unpublished eco ds; Schneide 1985; Depne 1992).
An unusual de elopmen
in
Namibia in ol ed epi-
demic sp ead o abies
in
kudu an elope (T age-
/aphus s epsice os) om 1977 o 1985
in
he cen-
al anching a ea. A localized ou b eak o abies
in
kudus occu ed
in
Windhoek dis ic
in
1975, bu he
epidemic which ollowed began
in
Okahandja dis ic
in
1977 and o e he nex ew yea s sp ead o Ka i-
bib, Oma u u, Windhoek, O a
i,
O jiwa ongo, Ou jou,
Gobabis, G oo on ein and Tsumeb dis ic s, causing
an es ima ed loss o 30
000-50
000 an elope, o
20%
o he popula ion, by he ime he ou b eak sub-
sided in 1985 (Shaw 1980; Ba na d & Hassel 1981;
Ba na d, Hassel, Geye & De Koke 1982; Hassel
1982; Schneide 1985; Hubschle 1988). I
is
belie ed
ha he kudu popula ion had a ained
an
unp ece-
den ed densi y du ing he 15 yea s p eceding he
R.
SWANEPOEL
e a/
.
epidemic, and his was asc ibed la gely o he con-
se a ion o he an elope because o he inc easing
alue o ophy hun ing and expo o enison, and
he occu ence o a succession o seasons o abo e
a e age ain all. An inc ease
in
jackal abies was
no ed p io o he epidemic and i was su mised ha
abid jackals ini ia ed he in ec ion
in
he kudus, bu
he ea e he numbe o cases eco ded
in
he an e-
lope was disp opo iona ely high
in
compa ison o
ha
in
jackals, sugges ing ha he disease was also
ansmi ed di ec ly be ween kudus. This possibili y
was s eng hened by he ac ha eas wa ds ex en-
sion o he ou b eak was ini ially checked o wo
yea s by a game con ol ence which hinde ed he
passage o an elope bu no small ca ni o es.
I
is belie ed ha ansmission be ween kudus was
a ou ed by hei p opensi y o indulge
in
sel and
mu ual g ooming, and by he ac ha o al ansmis-
sion would ha e been acili a ed
by
he mou h inju-
ies which kudus sus ain when b owsing on he Aca-
cia ho n ees which p edomina e
in
he a ec ed
a ea. Howe e , indi iduals some imes b owse
in
close p oximi y o each o he , so ha ansmission
o in ec ion h ough con amina ion o ege a ion was
possible.
I
was shown ha kudus a e highly sus-
cep ible o in ec ion
by
he o al ou e, and ha in-
ec ed indi iduals exc e e high concen a ions o a-
bies i us
in
sali a (Ba na d
e a/
. 1982).
The s a o he epidemic
in
kudus was ollowed
by
he occu ence o
an
ou b eak o abies
in
he E o-
sha Na ional Pa k
in
he ea ly 1980s, in ol ing ca -
ni o es anging om ba -ea ed oxes and jackals o
lions. The E osha Na ional Pa k lies
in
he pa hway
o he sou hwa ds sp ead o abies
in
1948 and
cases o he disease had been eco ded he e p io
o 1980, bu i
is
belie ed ha he in ec ion was p ob-
ably e-in oduced om Tsumeb dis ic
in
he 1980s
by
jackals which a e able o pene a e he game-
p oo ence o he pa k (Basson 1992).
The o al cases o abies con i med
in
Namibia om
1938-1992
a e summa ized
in
Table 1 (OVI 1932-
1992, unpublished eco ds; Schneide 1985; Depne
1992; Cen al Ve e ina y Labo a o y 1992a, unpub-
lished eco ds).
BOTSWANA
In
Bo swana, he e we e uncon i med ocal ou b eaks
o abies
in
Loba se
in
he sou h-eas
in
1919 and
1922, and
in
Ngamiland dis ic
in
he no h-wes
in
1936, whe e he diagnosis was i s con i med
in
a
dog
in
1938 acco ding o labo a o y eco ds (Ciu e
1927; OVI 1932-1992, unpublished eco ds; Depne
1992; Snyman 1940; Henning 1956; Mans el 1956).
F om unpublished e e ina y co espondence and
eco ds (Foggin 1988; T emle 1993),
i
appea s ha
an ou b eak o dog abies o mo e se ious p opo -
ions was no ed
in
Ngamiland adjacen o he Cap i i
S ip and Ka ango dis ic s o Namibia, a some
327
Rabies
in
sou he n A ica
TABLE 1 To al con i med cases o in ec ion wi h abies and abies- ela ed i uses eco ded
in
ou coun ies o sou he n A ica. Modi ied
and upda ed om Swanepoel (1994). Sou ces o in o ma ion as ci ed
in
ex
Sou h A ica Namibia Bo swana Zimbabwe
Species
1928-1992
1938-1992 1938-1992
1950-1992
Wild
animals
Cynic is penicilla a Yellow mongoose 2 119 -- -
Unspeci ied mongooses 1 3006 16• 158 55b
Gale ella sanguinea Slende mongoose 18 1 -39
Gale el/a pu e ulen a Small g ey mongoose 38 - - -
He pes es ichneumon La ge g ey mongoose ---3
Mungos mungo Banded mongoose 4 1 --
A ilax pa/udinosus Wa e mongoose 13 --1
Pa acynic is selousi Selous' mongoose 1 ---
Helogale pa ula Dwa mongoose 1 ---
/chneumia albicauda Whi e- ailed mongoose 2 --6
Su ica a su ica a Su ica e 118 4 --
Ci e ic is ci e a Ci e 3 - - 29
Gene a gene a Small-spo ed gene 174 23 22 3
Melli o a capensis Honey badge 23 27 16 45
lc onyx s ia us S iped poleca 67c 2 -5
Poecilogale albinucha S iped weasel 1 -- -
Unspeci ied o e species 1 -- -
Pan he a leo Lion -4 - -
Pan he a pa dus Leopa d -2 1 2
Acinonyx juba us Chee ah -4 --
Felis lybica A ican wildca 14 15 -6
Fel
is
ca acal Ca a
cal 14 4 1 -
Felis se al Se al --1 3
Felis nig ipes Small-spo ed ca 3 -1 -
Unspeci ied elids 170d
47d
12d -
Canis mesomelas Black-backed jackal 230° 360 193 1 5951
Canis adus us Side-s iped jackal -- - 166
O ocyon mega/o is Ba -ea ed ox 273 67 6 2
Lycaon pic us Wild dog -1 -3g
Vulpes chama Cape ox 9 8 - -
P o eles c is a us Aa dwol
23
14 -6
Hyaena b unnea B own hyaena 1 2 --
C ocu a c ocu a Spo ed hyaena 1 -4 6
Hys ix a icaeaus alis
Po
c
upine -1 --
Xe us inau is G ound squi el 32 1 --
Pa axe us cepapi T ee squi el 1 - - -
Th yonomys swinde ianus G ea e cane a 2 -- -
Lepus saxa ilis Sc ub ha e 1 ---
O yc e opus a e An bea - - -2
P oca ia capensis Cape hy ax 8 --1
Papio u sinus Chacma baboon 1 - - 2
Ce copi hecus ae hiops Ve e monkey -1 -2
Galago moholi Lesse bushbaby -1 - -
Phacochoe us ae hiopicus Wa hog 1 -- -
Syl icap a g immia Duike 17 9 5 7
Raphice us campes is S eenbok 4 1 --
T age/aphus s epsice os Kudu 3 386 1 2
Tau o agus o yx Eland 1 17 -4
Hippo agus nige Sable - - -2
Alcelaphus buselaphus
Red
ha ebees -1 --
Damaliscus do cas phillipsi Blesbuck 1 - - -
Redunca a undinum Reedbuck 1 - - 1
An ido cus ma supia/is Sp ingbok 3 1 --
Equus bu chelli Bu chell's zeb a 2 -- -
Unspeci ied he bi o es 6 -1 -
Epomopho us wanlbe gi Epaule ed ui ba 14h ---
Nyc e is hebaica Sli - aced ba 1 --1
Uniden i ied chi op e id,
possibly Miniop e us
sch eibe sii
1'
---
Uniden i ied chi op e id Insec i o ous ba 1
Unspeci ied/
un
iden i ied 32i 2 --
To al
wild
animals
4 754 1 023 279 1 999
328
R. SWAN
EP
O
EL
e a/
.
TABLE 1 (con inued)
Coun y Sou h A ica Namibia Bo swana Zimbabwe
Pe iod 1928-
1992
1938-1992 1938-1992
1950-1992
Domes ic
animals
Dogs 3
720
642 640
4
063
Ca s
454
'
70
21
152
Ca le 2
332
1
472
761 1
567
Sheep
127
65
7
81
Goa s 75
76
264
89
Ho ses and donkeys
58
28
42
79
Pigs 25 2 -17
Wa e bu alo -1 --
Guinea pigs 1 ---
To al
domes ic
animals
6
792
2
355
1
735
6 048
To al
11
546
3
378
2
014
8 047
Humans
• Belie ed o be mainly C. penicilla a
Belie ed o be mainly G. sanquinea
c P obably includes some P. albinucha
d
Be
lie ed o be mainly F. lybica
• Possibly includes a ew C. adus us
in
no h-eas e n T ans aal
s age be o e Ma ch 1950, and
by
Sep embe he
disease had c ossed o Se owe
in
he eas and
swep down he eas e n bo de o he sou h o he
coun y, i.e. in ec ion sp ead o all a eas whe e he
human, and he e o e dog, popula ion was mos
dense. Along he way, he in ec ion sp ead in o
sou h-wes e n Zimbabwe and he no he n T ans aal
egion o Sou h A ica. The disease has emained
ac i e
in
all o he a eas o Bo swana ini ially a -
ec ed, bu
in
addi ion o he o iginal p oblem o dog
abies wi h occasional human cases, he e has been
a endency o inc easing numbe s o jackals, ca le
and o he li es ock o
be
in ol ed (Maganu & S au-
ga d 1985; Mosienyane 1988; Masupu 1992; T emle
1993).
F om abou 1980 onwa ds a sepa a e ou b eak o
abies in ol ing domes ic he bi o es and wild ani-
mals de eloped
in
he Ghanzi dis ic on he wes e n
bo de o Bo swana, appa en ly as
an
ex ension o
he kudu epidemic
in
Namibia, and wi hin a ew
yea s had sp ead 1 000
km
sou h-eas wa ds ac oss
he coun y o Kga leng dis ic on he T ans aal
bo de o Sou h A ica (Mosienyane 1988). Sou h-
wa d ex ension occu ed in o Kgalagadi dis ic (Mo-
sienyane 1988), whe e he Gemsbok Na ional Pa k
o Bo swana adjoins he Kalaha i Gemsbok Na ional
Pa k o Sou h A ica, wi h which i is managed as a
uni , and
in
1986 abies was diagnosed
in
he spo -
ed hyaena (C ocu a c ocu a) in he Sou h A ican
pa k (OVI 1932-1992, unpublished eco ds).
The o al cases o abies con i med
in
Bo swana
om 1938-1992 a e summa ised
in
Table 1 (OVI
1932-1992, unpublished eco ds; Maganu & S au-
307
12
1 P obably includes many C. adus us
9 Cap i e animals
32
h Only wo ba s posi i ely iden i ied
as
E.
wah/be g1
' Possible iden i y based on ci cums an ial e iden
ce
I P obably includes domes ic animals
159
ga d 1985; Mosienyane 1988; Masupu 1992; T em-
le 1993).
I
should be no ed ha he e a e disc ep-
ancies wi h he igu es p esen ed
by
Swanepoel
(1994);
in
pa icula , species iden i ies ha e been
published o animals p e iously eco ded as unspe-
ci ied (T emle 1993).
ZIMBABWE
Zimbabwe was appa en ly ee o abies
in
1890
when Eu opean colonis s a i ed
in
he coun y, bu
some o he olde indigenous inhabi an s could ecall
ha he disease had been p esen
in
hei young
days (Edmonds 1922).
In
1902, dog abies appea ed
in
he Bulawayo a ea
in
sou h-wes e n Zimbabwe,
and he e appea s o
be
li le doub ha he disease
was in oduced om wes e n Zambia whe e he dis-
ease was known o be ampan
in
he Ba o seland
a ea
in
1901 ( he e was conside able a ic be ween
hese egions along he majo ade ou e linking he
wo coun ies a he ime) (Edmonds 1922; Shone
1962). Wi hin wo yea s 60 000 dogs we e des oyed
in
an
a emp o con ol he disease
in
Zimbabwe,
and, al hough his mus ha e ep esen ed a con-
side able p opo ion o he popula ion a he ime, he
disease con inued o sp ead h oughou mos o he
coun y. Con ol o he disease was inally achie ed
in
1913, his being asc ibed la gely o he imposi ion
o a dog ax, which p o oked d as ic olun a y
e
-
duc ion o he popula ion
on
he pa o dog owne s
(Edmonds 1922; Shone 1962). The in ec ion appa -
en ly did no become es ablished
in
wild hos s, and
ailu e o he ou b eak o ex end in o Sou h A ica
was asc ibed o p e en i e ac ion
in
he o m o a
329

Rabies
in
sou he n A ica
adical educ ion
o
he dog popula ion wi hin
an
80
km
wide s ip o he T ans aal along he Limpopo
i e whe e i o ms he no he n bo de s o he coun-
y wi h Zimbabwe and Bo swana (Mans el 1956).
A e 1913, Zimbabwe emained ee o abies un il
1938, when wo cases we e diagnosed
in
dogs a
Vic o ia Falls, and, as be o e, he e idence indica ed
ha he in ec ion had been in oduced om Zambia
(Shone 1962). Excep o b idges a ce ain poin s,
he Zambezi i e and he p esen day Lake Ka iba
o m
an
e ec i e na u al ba ie o he sp ead o
abies om Zambia, and a e 1938 Zimbabwe again
emained ee o he disease un il dog abies c oss-
ed
he sou h-wes e n and sou he n bo de s o he
coun y om Bo swana and he T ans aal
in
1950-
i
is belie ed ha he i us was in oduced by dogs
which accompanied people who c ossed he bo de s
illegally o pu chase g ain (Adamson 1954; Shone
1962). The disease sp ead apidly h ough Zimbab-
we, ollowing ou es along he mo e densely popu-
la ed communal a ming a eas, and by 1954 had
eached he no h o he coun y (Adamson 1954;
Shone 1962). The g ow h
in
he human popula ion
since 1913 made i di icul o en o ce o me ly e -
ec i e con ol measu es, such as dog " ie-up" o de s
and he des uc ion o s ays, and om
1951
on-
wa ds mass immuniza ion campaigns we e conduc-
ed wi h Flu y LEP (low egg passage) accine, which
had only ecen ly become a ailable (Adamson 1954;
Shone 1962; Williamson 1976).
By he ea ly 1960s con ol o he disease had been
achie ed o e mos o Zimbabwe, apa om esis-
an oci on he eas e n and wes e n bo de s wi h
Mozambique and Bo swana, and accina ion cam-
paigns we e scaled down (Mackinnon 1963; William-
son 1976; Foggin 1988). F om 1965 onwa ds, how-
e e , poli ical un es culmina ing
in
ci il wa ende ed
i inc easingly di icul o immunize dogs
in
he com-
munal a ming a eas, and he incidence o abies
p og essi ely ose o a eco d le el o
861
con i med
cases
in
1981, a e he cessa ion o he wa
in
1980
(Swanepoel & Foggin 1978; Law ence, Foggin & No -
al 1980; Fogg
in
& Swanepoel 1985; Fogg
in
1988).
Following he o mal ending o he wa , he con ol
o dog abies was complica ed by con inued s i e
in
Ma abeleland
in
he sou h-wes , and by
an
in lux o
e ugees om he ci il wa
in
Mozambique
in
he
eas , while elsewhe e
in
he coun y ackal abies as-
sumed se ious p opo ions (Foggin 1988). Dogs,
jackals and ca le comp ise
91
,
8%
(7
391
/8 04
7)
o
all animals
in
which abies was con i med
in
Zimba-
bwe om 1950 o 1992 inclusi e (Table
1)
(Fogg
in
1988; Bingham 1992; Ve e ina y Resea ch Labo a-
o y 1992b, Zimbabwe, unpublished eco ds) and no
o he coun y has eco ded as many cases o jackal
abies.
A mino po ion o he land
in
Zimbabwe
is
de o ed
o na ional pa ks and u ban de elopmen , while he
330
bulk o he coun y is di ided app oxima ely equally
be ween comme cial and communal a ming. Com-
me cial a ms a e gene ally well wooded, and apa
om la ge wild ca ni o es and he bi o es which
ha e been elimina ed, wildli e, including jackals,
is
gene ally p ese ed o ole a ed, and ew dogs a e
kep (Foggin 1988).
In
con as , communal a ming
a eas a e gene ally o e g azed and de o es ed, wild-
li e
is
sca ce, and dogs a e kep o hun ing. Conse-
quen ly, dog abies has occu ed mainly
in
o close
o communal a ming a eas, and jackal abies has
occu ed almos exclusi ely
on
comme cial a ms
(Fogg
in
1988). Bo h he black-backed jackal and he
side-s iped jackal
(C.
adus us) occu
in
Zimbabwe,
wi h pa ially o e lapping dis ibu ions, and bo h a e
in ol ed
in
ou b eaks o abies.
Jackal abies was i s diagnosed
in
Zimbabwe
in
1952, some 15 mon hs a e he disease had en e ed
he coun y
in
1950, and he i s ou b eaks occu ed
along he eas e n bo de (Cumming 1982; Foggin
1985a; 1988). The ea e , ou b eaks o jackal abies
occu ed
in
widely sepa a ed dis ic s a i egula in-
e als o many yea s (Cumming 1982; Foggin 1988;
Kennedy 1988). Since he ou b eaks always appea -
ed o occu
in
p oximi y o ou b eaks o dog abies
and did no ecu
in
he same a eas o pe iods o
se en yea s o mo e,
i
was a gued ha he i us
was no adap ed o main enance
in
jackals bu had
o be e-in oduced by dogs (Cumming 1982). How-
e e , se e al o he ou b eaks which ha e occu ed
since 1965, and spo adic isola ions o he i us om
jackals, ha e aken place well away om known cen-
es o in ec ion
in
o he species (Foggin 1988). Mo e-
o e ,
in
one pa icula ins ance
i
was clea ha
an
ou b eak o abies
in
dogs
in
ac ollowed sp ead o
he disease
in
jackals (Kennedy 1988).
I
can he e-
o e
be
concluded ha abies appea s o be eadily
ansmi ed by jackals
in
Zimbabwe, and ha he in-
ec ion
is
eely communicable be ween dogs and
jackals, as seems o be he case
in
o he pa s o
sou he n A ica. An al e na i e explana ion gi en o
he ailu e o abies o pe sis
in
ci cumsc ibed a eas
o
comme cial a ming whe e epidemics occu ed
in
jackals, was ha he disease educed he densi y o
jackals
in
hese a eas o below he h eshold e-
qui ed o he sp ead o he in ec ion (Foggin 1988).
As obse ed
in
pa s o Zambia, Namibia, Bo swana
and Sou h A ica whe e jackal abies occu s, ou -
b eaks o he disease
in
jackals
in
Zimbabwe a e in-
a iably accompanied by he occu ence o he dis-
ease
in
ca le, bu he epo ed deg ee o in ol e-
men o ca le a ies:
i
is
a common inding
in
sou h-
e n A ica ha labo a o y con i ma ion o he diag-
nosis
is
sough
in
only a mino p opo ion o sus-
pec ed cases o abies
in
ca le, pa icula ly once he
diagnosis has al eady been es ablished
in
a he d
(Von Mal i z 1950; B uckne , Hu e & Bosho 1978;
Maganu &
S auga ;J
1985; Zyambo
e a/
. 1985; Fogg
in
1985a; 1988; Mosienyane 1988).
In
one ou b eak
in
Zimbabwe i was es ablished by e ospec i e in es i-
ga ion ha 1 200 ca le had died, o which 140 we e
con i med o ha e been abid (Foggin 1988). As wi h
ca le, he occu ence o abies
in
mos o he species
in
Zimbabwe appea s o ep esen spill-o e o in ec-
ion om ei he dogs o jackals, bu he e ha e been
clus e s o cases o he disease
in
he slende mon-
goose (Gale el/a sanguinea)
in
he sou h-wes o he
coun y
on
occasion, which sugges s he exis ence
o
an
independen cycle o ansmission
in
his spe-
cies (Foggin 1988).
SOUTH AFRICA, MOZAMBIQUE AND
SWAZILAND
The e a e his o ical epo s ex ending back o he
eigh een h cen u y o spo adic cases o disease e-
sembling abies
in
dogs and humans ha ing been
obse ed
in
Sou h A ica (Ciu e 1927; Nei z & Ma-
ais 1932; Snyman 1940; Henning 1956). Howe e ,
an
ou b eak o he disease
in
dogs was i s con-
i med
in
he eas e n Cape P o ince
in
1893 by in-
ocula ion o abbi s, and his ep esen s he i s oc-
casion
on
which a diagnosis o abies was con i med
on
he con inen o A ica (Hu cheon 1894). The ou -
b eak was ini ially ecognized
in
Po Elizabe h
in
Ap il1893, bu he esul s o inqui ies sugges ed ha
he i s case had occu ed
in
Sep embe 1892
in
a
dog impo ed om England, which had become abid
a ew weeks a e i s a i al. The ou b eak was be-
lie ed o ha e a ec ed abou 90 dogs, se en ca s
and a ew ca le, bu no wild animals, and had
sp ead o Ui enhage, Jansen ille, Willowmo e and
Albany dis ic s by he ime ha i was b ough unde
con ol
in
Augus 1894, h ough he muzzling and
es ic ion o dogs and he des uc ion o s ays (Hu -
cheon 1894; Edding on 1895; Henning 1956).
A e 1894, abies was no con i med again
in
Sou h
A ica o 34 yea s, bu he e was moun ing anecdo-
al e idence o indica e ha
an
endemic o m o he
disease associa ed wi h i e ids was p esen .
In
pa icula , he e was a gene al belie
in
he eas e n
and no he n Cape P o ince ha bi es om gene s
caused a al, abies-like illness
in
humans, and spe-
ci ic epo s o such inciden s da ed back o 1885
(Fi zsimmons 1919; Clu e 1927; Snyman 1940; Nel-
son 1962). Clu e (1927) documen ed
11
uncon i m-
ed
cases o human abies ( equen ly misquo ed as
en cases) which occu ed
in
he sou he n T ans aal,
O ange F ee S a e and no he n Cape P o ince om
1916 o 1927, ollowing bi es by yellow mongooses
(Cynic is penicilla a), dogs and a small-spo ed gene
(Gene a gene a). The disease was inally con i med
in
1928
in
wo child en bi en by a yellow mongoose
in
Wolma anss ad dis ic
in
he sou h-wes e n T ans-
aal (He zenbe g 1928), and since ha ime abies
has been diagnosed egula ly
in
Sou h A ica.
R.
SWANEPOEL
e a/.
Wi hin a sho pe iod a e he diagnosis o he dis-
ease was con i med
in
1928, abies was ecognized
in
nume ous loca ions
in
Sou h A ica
in
dogs, do-
mes ic ca s, yellow mongooses, su ica es (Su ica-
a
su ica a), gene s and wild elids, and
in
humans
and a m animals which had been bi en by hese
ca ni o es
(Du
Toi 1929; Nei z & Ma ais 1932; Nei z
& Thomas 1933; 1934; Snyman 1937). Ini ial conjec-
u e ha he disease migh ha e a isen by ex ension
om he epidemics o dog abies which had occu ed
in
he eas e n Cape P o ince om 1892 o 1894
and
in
Zimbabwe om 1902 o 1913, ga e way o
he con ic ion ha abies had long been p esen
in
i e ids
in
Sou h A ica, possibly o cen u ies, bu
had simply no been ecognized
(Du
Toi 1929; Nei z
& Ma ais 1932; Nei z & Thomas 1933; 1934;
Du
Toi
1936; Snyman 1937; 1940).
The e e ina y in es iga o s we e well awa e ha he
newly ecognized disease
in
Sou h A ica, which oc-
cu ed p incipally
in
he yellow mongoose, di e ed
undamen ally om wha hey e med classical Eu o-
pean ype dog abies
in
ha he e we e spo adic
cases
in
dogs, bu no eal endency o he in ec ion
o sp ead among hem.
In
ac , Nei z & Ma ais (1932)
p e-emp ed he concep o abies i us bio ypes by
se e al decades when hey s a ed: "The e seems o
be comple e adap a ion [o he i us] o one amily
o Ca ni o a [Vi e idae] and occasionally a domes ic
animal o human being is acciden ally bi en and
con ac s he disease. The e he ou b eak ends
in
spi e o he ac ha condi ions a e being [sic] a-
ou able o he sp ead o he disease
..
. he adap-
a ion o he disease o one amily, is su ely no
some hing which can occu
in
so sho a pe iod as
a decade o wo". La e , b ie e e ence was made
o he ac ha expe imen ally in ec ed mongooses
we e unable o ansmi in ec ion o dogs by bi e, bu
he s ain o i us, species o mongoose and num-
be s o
an
imals
on
expe imen , we e no speci ied
(Alexande 1952).
As he a ea known o be a ec ed by abies expand-
ed, he e was specula ion ha his was due o bo h
ecen sp ead and he ac ha he ue dis ibu ion
o he disease was s ill being elucida ed (Snyman
1940), bu subsequen ly he esul s o delibe a e
in es iga ions e ealed ha he occu ence o en-
demic mongoose abies was con luen o e he
g ea e pa o he in e io pla eau o Sou h A ica
wes o he D akensbe g moun ains (Me edi h 1977;
1982). The only a eas o be excluded we e hose
which ell ou side he dis ibu ion o he yellow mon-
goose- he no he n T ans aal apa om a na ow
cen al s ip passing h ough Sou pansbe g dis ic ,
Na al apa om he no h-eas e n ma gin o he
p o ince, he eas e nmos T anskei po ion o he
Cape P o ince and a na ow coas al egion ex end-
ing om Po Elizabe h owa ds Cape Town (Me edi h
1977; 1982; Smi he s 1983). The mongoose occu s
331
Rabies in sou he n A ica
less abundan ly in Bo swana whe e i is absen
in
he
eas , and i
is
p esen
in
Namibia apa om he
coas al Namib dese (Smi he s 1983).
The yellow mongoose is diu nal and i s ole as a
main enance hos o abies i us is acili a ed by he
ac ha i li es
in
colonies (Snyman 1940; Zump
1968; 1969; 1976; 1982; Smi he s 1983; Wenhold
1990; Raza, Wenhold, Howa d, Ma ais, & Palle
1992). I
is
mos abundan
in
he no h-wes e n O -
ange F ee S a e and
in
he adjacen sou h-wes e n
T ans aal, and mongoose abies
is
mos p e alen
in
his a ea (Snyman 1940; Zump 1976; 1982). O e
much
o
i s dis ibu ion, he yellow mongoose u ilizes
and adap s wa ens pionee ed by he g ound squi el
(Xe us inau is), a oden wi h which he :nongoose
sha es he wa ens
in
appa en ha mony (Snyman
1940; Zump 1976). I ine an g oups o su ica es,
which a e also i e ids, occasionally e ic yellow
mongooses and g ound squi els empo a ily om
wa ens (Snyman 1940; Zump 1982). Compa a i ely
ew cases o abies ha e been eco ded
in
g ound
squi els and su ica es, he dis ibu ions o which
la gely coincide wi h ha o he yellow mongoose,
bu he e appea o ha e been no speci ic a emp s
o de e mine he ela i e popula ion densi ies o he
h ee species.
I
is
possible ha abid g ound squi -
els and su ica es ha e been misiden i ied as yellow
mongooses on occasion, o ca ego ized as uniden i-
ied mongooses.
As ea ly as 1930 e o s we e made o con ol i e -
id abies h ough he e adica ion o he yellow mon-
goose, and om 1939 onwa ds i became ou ine
~o
pump cyanogas (la e phosphine) in o wa ens 1n
loca ions whe e mongoose abies was diagnosed
(Snyman 1940; Zump & De B uyn 1967). Abou
50
000-160
000 hec a es we e ea ed annually un il
excep ionally hea y ains
in
197
4-1976
es ic ed
he access
o
con ol eams o a ec ed si es (Depa -
men o Ag icul u e 1940-1992, unpublished eco ds;
Snyman 1940, Zump &
DeB uyn
1967).
Sine~
hen
mongoose con ol, which had become
p oh1b1 ! ely
expensi e, has been applied much mo e selec i ely
o s a egic loca ions whe e he disease occu s
in
p oximi y o u ban cen es.
I
was ealized om
an
ea ly s age ha ocal e adi-
ca ion
o
he mongoose p o ided only empo a y
con ol
o
abies, and he incidence o he disease
ose p og essi ely
in
each decade om 1950 on-
wa ds
o
each epidemic p opo ions by he ea ly
1970s, despi e he applica ion o he con ol
~eas
u es, and con inued o luc ua e a
h1gh
le els 1n he
1970s and 1980s ollowing he abandonmen o sys-
ema ic mongoose con ol (Table
2)
(OVI1932-1992,
unpublished eco ds; Snyman 1940; Zump 1982).
Rabies o he yellow mongoose emains hype en-
demic in he sou h-wes e n T ans aal and adjoining
no h-wes e n O ange F ee S a e, bu occu s h ough-
332
ou he sou he n T ans aal, and he O ange F ee
S a e whe e he mongoose cons i u es app oxima ely
60%
o all animals
in
which he disease has been
eco ded o e he pas wo decades (OVI
193~-
1992, unpublished eco ds). The e
is
spill~o e
o
In-
ec ion o ca le, dogs, ca s, o he domes 1c an1mals,
he s iped poleca (lc onyx s ia us) and he slende
and wa e (A ilax paludinosus) mongooses. Spo adic
cases
o
abies a e also eco ded
in
su ica es, gen-
e s, ba -ea ed oxes, jackals and wild elids, pa icu-
la ly
in
he d ie wes e n dis ic s (OVI
1932-1992
,
unpublished eco ds). Since sheep ou numbe ca le
in
he a eas whe e mongoose abies occu s, and
bo h species p edispose hemsel es o acial bi es
by displaying cu iosi y owa ds abid mongooses
(Du
Toi 1929·
OVI1932-1992
, unpublished eco ds; Al-
exande 1' 952; Ma e 1962; Mans el 1965), i is cu i-
ous ha he numbe o cases o abies eco ded
in
ca le as ly ou numbe hose
in
sheep (Table 1 ).
The epidemiology o he disease
is
mo e complex
in
he Cape P o ince, whe e he yellow mongoose
cons i u es less han
30%
o abid animals eco ded
o e he pas 20 yea s
(OVI1932-1992,
unpublished
eco ds), and whe e se e al o he ca ni o .es appea
o be in ol ed
in
independen ansm1ss1on o he
i us. Su ica es, s iped poleca s and g ound squi -
els seem o acqui e he in ec ion only whe e he dis-
ease occu s
in
he yellow mongoose, bu he e ha e
been clus e s o cases sugges i e o localized sp ead
in
he wa e mongoose, and
in
he small g ey m.on-
goose (Gale ella pu e ulen a), which occu s mamly
sou h o he O ange Ri e (OVI 1932-1992, unpub-
lished eco ds).
In
he no he n Cape P o ince, a-
bies o gene s and wild elids has been diagnosed
mo e equen ly han elsewhe e
in
he
coun ~y,
bu
i is no clea whe he he e is independen
mcula-
ion o i us
in
hese animals o whe he hey acqui e
in ec ion om mongooses, possibly
in
hei ole as
p eda o s. Howe e , he e has also ?een a
much
highe incidence o abies
in
domes ic ca s 1n he
same a ea han elsewhe e (Ba na d 1979a), and his
could indica e ha he e is sp ead o in ec ion among
elids wi h he A ican wild ca (Felis lybica) se ing
as a link o e al domes ic ca s wi h which i in e -
b eeds.
The e
is
s onge e idence o sugges ha he e
is
independen sp ead o abies
in
he ba -ea ed ox,
a small canid which occu s
in
he d ie wes e n
pa s
o
he sub-con inen , and which subsis s la ge-
ly on e mi es. Spo adic cases o
ab1es
we e e-
co ded
in
he ox om 1955 onwa ds, bu an In-
c ease
in
he incidence o he disease was no ed
in
he no he n and wes e n Cape P o ince du ing he
1970s, sho ly a e he e had been a simila in-
c ease
in
Namibia (see abo e), and om 1980 on-
wa ds he e ha e been up o 24 con i med cases
each yea , wi h p og essi e sp ead o he disease
o he wes coas (almos o he en i ons o Cape
R SWANEPOEL
e
a .
TABLE 2 To al
co
n i
m
ed cases o in ec
i
on wi h abies
an
d abies- ela ed i uses eco ded in an
im
als in Sou h A ica, 1
928
- 1992.
Sou ces o in o
m
a
io
n as o Table 1
Species 28 29 30
31
32 33 34 35
36
37 38
Domes
ic
anima
ls
Dogs -1 1 1 1 1 1 1 2 1 6
Ca
s --- - 2 4 6 1 3 -3
Ca le -1 3 2 2 1 1 1 7 3 16
Sheep - - -1 -2 1 --- -
Goa
s -- - - - - - - - - -
Ho ses and donkeys -- - - - - - 1 --3
Pigs -2 --- - - - - - 2
Guinea pigs - - --- - - - - - -
To al
domes
ic
animals
-4 4 4 5 8 9 4 12 4 30
Wild
an
i
ma
ls
Cynic is
peni
cilla a Yellow mongoose 2 5 1 1 6 15 8 -4 5 6
Unspeci ied mongooses,
p obably C. penicilla a ----------1
Ga/e ella sanguinea
Sl
ende mongoose --- - - - - - - - -
Gale ella pu e u/en a Sma
ll
g ey mongoose - - 1 --------
Mungos mungo Banded mongoose - - - --- - - - - -
A ilax
pal
udinosus
Wa
e mongoose - - ---------
Pa acynic is selousi Selous' mongoose - - - -- - - - ---
Helogale pa ula Dwa mongoose -- - - - - - -- - -
/chneumia albicauda Whi e- ailed mongoose - - -- - - - - - - -
Su ica a su ica a Su ica e - - 2 -1 - - -- - -
Ci e ic is ci e a Ci e --------- - -
Gene a spp.
ma
inly
G.
gene a
Sma
ll
-s
po ed
gene
-2 - - 3 1 - - 2 -1
Unspeci ied i e ids -1 ---------
Melli o a capensis Honey badge - - ---------
lc on
yx
s ia us S iped poleca e ------ - --- -
Poecilogale albinucha S iped weasel -- - - - - - - - - -
Aonyx
ca
pensis o Lu a O e ----- - -----
maculicollis
Felis /ybica A ican wildca -- - - - - - - - --
Felis ca acal Ca acal -- - - - - - - - - -
Felis nig ipes Small-spo ed ca - - - - -------
Unspeci ied elids - - - -
~
1 2 -3 --
Canis spp.
ma
inly
C.
mesome/as Black-backed jackal - - --- - - - - - -
O
ocyon mega/o is Ba -ea ed ox ---------- -
Vulpes chama Cape ox ---- - - - - - - -
P
o
e
les c is a us Aa d
wo
l - - - - - - - - - - -
Hyaena b unnea B own hyaena ---------- -
C ocu a c ocu a Spo ed hyaena -- - - - - - - - - -
Xe us inau
is
G ound squi el - - - - - ---1 --
Pa axe us cepapi T ee squi el --- - - - - - - - -
Th yonomys swinde ianus G ea e cane a ------ - -- - -
Lepus spp
.,
p obably
L.
saxa ilis Sc ub ha e - - -- - - -- - - -
P
oca ia capensis Cape hy ax ------ - -- - -
Papio u sinus Chacma baboon - - - - - - - - - - -
Phacochoe us ae hiopicus Wa hog - - -- - - - - - - -
Sy/ icap a g immia Duike ------- - - - -
Raphice us campes
is
S eenbok --- - - - - - - - -
T agelaphus s epsice os
Ku
du -- - - - - - - - - -
Tau o agus o yx
El
and ------ - --- -
Damaliscus do cas phillipsi Blesbuck ---- - - - - - - -
Redunca a undinum Reedbuck - - - - -------
An ido cus ma supia/is Sp ingbok ------- - - - -
Equus bu chelli Zeb a -- - - - - - - - --
Unspeci ied he bi o es -- - - - - - - - - -
Epomopho us wahlbe gi Wahlbe g's epaule ed
ui ba - - - - - - - - - - -
Unspeci
ie
d p e opodid,
p obably E. wahlbe gi -------- - - -
Nyc e is hebai
ca
Sli - aced ba -- - - - - - - -- -
Uniden i ied chi op e id,
possibly Miniop e us
sch eibe sii -- - - - - - - - - -
Uninden i ied chi op e
id
Insec i o ous ba - - --- - - - - - -
O he unspeci ied o
u
ni
den i ied animals* -- - - - - - 4 -- -
To al
wild
animals
2 8 4 1 10 17 10 4 10 5 8
To al
2 12 8 5 15 25 19 8 22 9 38
* P obably
in
cludes domes ic animals
333
Rabies
in
sou he n A ica
o he u al popula ion
in
he coas al and many o he
midlands dis ic s o Na al a ou ed he sp ead o he
disease
in
dogs, and he epidemic which ollowed
he in oduc ion o he i us in o he p o ince
in
1961
was o an in ensi y unp eceden ed
in
Sou h A ica.
Vigo ous e o s we e made o con ol he disease
(Mans el 1962), and he ou b eak was inally b ough
o
an
end la e in 1968 (OVI
1932-1992
, unpublished
eco ds).
Rabies eappea ed
in
he no he n dis ic s o Na al,
adjacen o he Mapu o dis ic o Mozambique,
in
mid 1976 (OVI 1932-1992, unpublished eco ds) a
a ime when he e was an in lux o e ugees leeing
he unse led condi ions which ollowed he assump-
ion o independence by Mozambique om Po ugal.
Du ing he eigh yea s since abies had las been di-
agnosed he popula ion o Na al had con inued o
bu geon, and since many u al inhabi an s sough
li elihoods
in
u ban cen es, in o mal se lemen s
lou ished, whe e uncon olled dog popula ions p o-
ided e ile g ound o epidemic sp ead o he dis-
ease. Dog abies sp ead om Na al o he T anskei
a ea o he eas e n Cape P o ince
in
1987, and
by
he ea ly 1990s had eached he Ciskei a ea (OVI
1932-1992, unpublished eco ds; Regional Ve e i-
na y Labo a o y 1987-1992, unpublished eco ds).
Like Na al, he T anskei and Ciskei a eas ha e
dense u al and pe i-u ban popula ions, and he
eme gence o dog abies
in
he eas ep esen s ye
ano he de elopmen
in
he complex pa e n o a-
bies
in
he Cape P o ince.
Since i s e-in oduc ion in o Na ai-KwaZulu
in
1976,
dog abies has p o ed o
be
in ac able. App oxi-
ma ely
75%
o all cases o abies eco ded
in
dogs
in
Sou h A ica since 1976 ha e occu ed
in
Na ai-
KwaZulu, and dogs ep esen abou
88%
o all ani-
mals in which abies has been eco ded
in
he p o -
ince du ing his pe iod (OVI 1932-1992, unpublished
eco ds; Regional Ve e ina y Labo a o y 1987-1992,
unpublished eco ds). Peak accina ion co e age o
59%
o he es ima ed o al dog popula ion o Na ai-
KwaZulu was a ained
in
1980-1981 (Depa men o
Ag icul u e 1940-1992, unpublished eco ds; Swane-
poel1994) bu he immuniza ion o un es ic ed dogs
in
in o mal se lemen s cons i u es a o midable ask
which has been ende ed inc easingly di icul
by
he
poli ical un es which has de eloped
in
he p o ince.
Consequen ly, inadequa e accina ion co e age has
been a ained
in
ecen yea s
in
he s a egically im-
po an loca ions whe e he p oblem is mos se e e.
P io o 1950, mos cases o human abies
in
Sou h
A ica esul ed om bi es by i e ids (and spo adic-
ally o he animals)
in
he sou he n T ans aal, O ange
F ee S a e and Cape P o ince, bu ollowing he in-
cu sion o he canid i us in o he no he n T ans-
aa , and la e Na al, dogs became he mos impo -
340
an sou ce o human in ec ion. Cu en ly, abou 10-
20 cases o human abies a e eco ded each yea
in
Sou h A ica, mainly
in
associa ion wi h dog bi es
in
Na ai-KwaZulu, bu human disease
is
o g owing
impo ance
in
he eas e n Cape P o ince (OVI
1932-
1992, unpublished eco ds; Na ional Ins i u e o Vi ol-
ogy 1953-1992, unpublished eco ds; Ve e ina y Lab-
o a o y, T anskei 1990-1992, unpublished eco ds).
Monoclonal an ibody s udies ha e con i med ha i-
us s ains associa ed wi h endemic i e id abies
in
Bo swana and Sou h A ica can be dis inguished
om he canid s ain which appea s o ha e sp ead
h ough Namibia, Bo swana, Zimbabwe and pa s o
Sou h A ica om abou 1947 onwa ds (Na ional
In
-
s i u e o Vi ology 1953-1992, unpublished eco ds;
Schneide , Ba na d & Schneide 1985; King
1991
).
Vi us associa ed wi h he ba -ea ed ox
in
he no h-
e n and wes e n Cape P o ince con o ms o he ca-
nid ype (Na ional Ins i u e o Vi ology 1953-1992,
unpublished eco ds; King
1991
), and his ends o
con i m he imp ession ha apa om he pu a i e
sp ead o he canid i us ac oss he no he n T ans-
aal and h ough Mozambique in o Na al, he e may
ha e been a u he in oduc ion o he i us in o
Sou h A ica ia he ox om Namibia.
The o al ange o species a ec ed and numbe s o
cases o abies which ha e been con i med since
egula moni o ing was ins i u ed
in
Sou h A ica
in
1928, up o 1992 inclusi e, a e summa ised
in
Ta-
ble
1,
and annual igu es a e p esen ed
in
Table 2.
As a as possible, he in o ma ion was de i ed om
o iginal labo a o y eco ds (OVI 1932-1992, unpub-
lished eco ds; Na ional Ins i u e o Vi ology 1953-
1992, unpublished eco ds; Regional Ve e ina y Lab-
o a o y
1987-1992
, unpublished eco ds; Ve e ina y
Labo a o y, T anskei 1990-1992, unpublished ec-
o ds). O iginal abies eco ds a he Ve e ina y Ins i-
u e, Onde s epoo , emain a ailable only o he
yea s om 1967 onwa ds, bu ansc ip s o he o ig-
inal eco ds o he yea s 1932-1943, and 1952-
1966 inclusi e, had been p epa ed
by
one o he
p esen au ho s
(C
.D.M.
).
Summa ised igu es could
be
de i ed o he yea s 1944-
1951
om exis ing
in
-
o ma ion (OVI 1932-1992, unpublished eco ds; De-
pa men o Ag icul u e 1940-1992, unpublished ec-
o ds), al hough only combined o als could be de e -
mined o he wo yea s 1944 and 1945 (Table
2)
.
Nei z & Ma ais (1932), and Nei z & Thomas (1933;
1934) published de ailed eco ds o he yea s 1928-
1933, bu o he wise he in o ma ion p esen ed
in
Ta-
bles 1 and 2 canno be de i ed accu a ely om pub-
lica ions o depa men al epo s: igu es gi en o
a
-
bies cases eco ded du ing pa icula pe iods a e
ei he impossible o ela e o calenda yea s, o o
econcile wi h each o he wi hou o e laps o gaps,
o else he publica ions and epo s do no s ipula e
any o all species in ol ed, o hey deal wi h only a

pa icula egion, o omi igu es o he sel -go e n-
ing
and independen s a es, o hey p esen igu es
which a e a a a iance wi h labo a o y eco ds
(Du
Toi 1929; 1936; Snyman 1937; 1940; 1953; Alexan-
de 1952; Henning 1956; Mans el
195(?;
Ma e 1962;
Tus in & Smi 1962; Lamb ech s 1964; Nei z 1965;
Zump 1969; B uckne e
a .
1978; 1936; Ba na d
1979a;
b;
Gummow 1985; Gummow & Tu ne 1986;
Keigh ley, S u he s, Johnson & Ba na d 1987; Swa
1989). I should
be
no ed ha Tables 1 and 2 include
cases o in ec ion wi h abies- ela ed i uses eco d-
ed
in
Sou h A ica and Zimbabwe, as discussed be-
low.
LESOTHO
Dog abies sp ead om Na al in o he no h-eas e n
co ne o Leso ho
in
1982, when he disease was e-
co ded he e o he i s ime, and wi hin wo yea s
i
had sp ead h oughou he coun y (Khoma i 1988;
Sco 1988; Khoma i 1992). Rabies has been eco d-
ed
in
dogs, ca s, ca le, sheep, goa s, ho ses and
donkeys
in
Leso ho (Table
1)
, bu moni o ing o he
disease has p o ed o
be
di icul
in
he moun ainous
e ain o he coun y and human disease has been
eco ded wi h disp opo iona e equency. The e a e
ew e al ca ni o es
in
he coun y and he disease
has no been eco ded
in
wild animals. Dog abies
eached he wes e n bo de o Leso ho
in
he mid
1980s, bu did no pene a e deeply in o he O ange
F ee S a e whe e p e en i e accina ion had been
unde aken, and whe e he dog popula ion
is
less
dense.
RABIES-RELATED VIRUSES
La ge numbe s o isola ions o abies i us (lyssa-
i us
1)
we e made om non-haema ophagous ba s
in
he Ame icas du ing he 1950s, and his p omp ed
in es iga ions elsewhe e
in
he wo ld. As a conse-
quence, Lagos ba i us was isola ed om s aw-
colou ed ui ba s (Eidolon hel um)
in
Nige ia
in
1956 (Boulge & Po e ield 1958), bu i was no un-
il1970 ha he i us was iden i ied
as
a habdo i us
an igenically ela ed o, bu dis inc om, abies i us
(Shope, Mu phy, Ha ison, Causey, Kemp, Simpson
& Moo e 1970).
In
1968, a i us was isola ed om
C ocidu a sp. sh ews apped
in
Mokola Fo es nea
lbadan, Nige ia, and om a sh ew ound dead
in
lba-
dan (Kemp, Causey, Moo e, Odeola & Fabiyi 1972;
Kemp, O is, Se ze & Moo e 1974). Iden i ica ion o
he Mokola i us
as
a habdo i us ela ed o abies
i us was epo ed
in
he same publica ion as he
iden i ica ion o Lagos ba i us, and hence he con-
cep o a abies- ela ed subg oup o habdo i uses
was es ablished (Shope e
a/.
1970).
In
1970,
an
adul male li ing
in
he Wa mba hs dis ic abou
1 00
km
no h o P e o ia, Sou h A ica, died o a-
bies-like disease i e weeks a e being bi en by
an
R.
SWANEPOEL
e a/.
insec i o ous ba , possibly Miniop e us sch eibe sii
(Van de Me we 1982), and a i us isola ed om his
b ain was ound o
be
ye ano he abies- ela ed
i us; named Du enhage a e he ic im (Me edi h,
Rossouw & Van P aag Koch
1971
; Tigno , Mu phy,
Cla ke, Shope, Mado e, Bowe , Buckley & Me edi h
1977).
The h ee abies- ela ed i uses ha e been encoun -
e ed
in
only a ew coun ies o wes e n and sou he n
A ica whe e app op ia e in es iga ions ha e been
unde aken (Table
3),
and ecen ly
in
E hiopia (Me-
ba sion, Cox & F os 1992).
In
gene al, isola es ob-
ained om os ensibly heal hy ba s, sh ews o o-
den s
in
su eys we e delibe a ely subjec ed o es s
app op ia e o he iden i ica ion o abies- ela ed i-
uses, whe eas isola es ob ained om specimens
submi ed om humans and lowe animals o he in-
es iga ion o suspec ed abies, we e only ecog-
nized
as
abies- ela ed i uses because he in es i-
ga o s conce ned we e ale o non- speci ic ea u es
which dis inguished he in ec ions om abies: ou-
ine diagnos ic p ocedu es do no allow abies and
abies- ela ed i uses o be di e en ia ed wi h ce -
ain y. Thus, he wo isola ions o Lagos ba i us
made om ca s
in
Sou h A ica and Zimbabwe (Ta-
ble
3),
we e bo h made om animals which had been
accina ed agains abies. The cases occu ed
in
iso-
la ion om known cases o abies, luo escence was
obse ed o
be
weak
in
diagnos ic es s wi h poly-
clonal an i- abies luo escein conjuga e, and he signs
o disease mani es ed by he ca
in
Zimbabwe, le ha -
gy
and pa esis wi hou agg essi eness, we e consid-
e ed o
be
a ypical o abies (OVI 1932-1992, un-
published eco ds; Fogg
in
1988; King & C ick 1988).
Isola ions o Lagos ba i us om ui ba s
in
Sou h
A ica
in
1980 (Table 3), we e made om indi iduals
obse ed o beha e abno mally a a ime when pub-
lic awa eness was heigh ened by he ac ha dog
abies was in
an
epidemic phase
in
Na al
(OVI1932-
1992, unpublished eco ds). Fluo escence wi h poly-
canal an i- abies conjuga e was obse ed
in
he
b ains o 13 o he ba s submi ed om Na al ( en
in
1980 and h ee
in
1981
),
and all 13 cases a e includ-
ed
in
Tables 1 and 2. The a ec ed indi iduals a e
belie ed o ha e been common epaulle ed ui ba s
(Epomopho us wahlbe g ) al hough only one was
posi i ely iden i ied. Only h ee o he b ains om
Pine own and Du ban which luo esced
in
1980 we e
cul u ed and, al hough wo o he isola es ob ained
we e a one s age epo ed o be Mokola i us (Fog-
gin & Swanepoel 1985; Schneide e
a .
1985), all
h ee ha e been con i med o
be
Lagos ba i us
in
monoclonal an ibody es s (only hese h ee posi i e-
ly iden i ied isola es a e included in Table
3)
(OVI
1932-1992, unpublished eco ds; Na ional Ins i u e
o Vi ology 1953-1992, unpublished eco ds; Me e-
di h & S anding 1981; C ick, Tigno & Mo eno 1982;
King & C ick 1988). A u he isola ion o Lagos ba
341
Rabies
in
sou he n A ica
TABLE 3 To al isola ions o abies- ela ed i uses
in
A ica
Species Isola ions
Lagos
ba
i us
Eidolon hel um ( ui ba )
Mic op e us pusillus ( ui ba )
Epomopho us wahlbe gi ( ui ba )
Domes ic ca
Nyc e is gambiensis (insec i o ous ba )
Eidolon hel um ( ui ba )
Domes ic ca
Epomopho us wahlbe gi ( ui ba )
Dog
Mok
ola
i
us
C ocidu a sp. (sh ew)
Human
Domes ic ca
Human
C ocidu a sp. (sh ew)
Dog
Domes ic ca
Lophu omys sikapusi (myomo ph oden )
Domes ic ca
Ou enhage
i us
Human
Miniop e us sch eibe sii (insec
i
o ous ba )
Nyc e is hebaica (insec i o ous ba )
a Only one ba posi i ely iden i ied as
E.
wahlbe gi
b Iden i ica ion o species based on ci cums an ial
e idence
c Re e ences o Table 3
a Boulge & Po e ield 1958
b Shope e a/. 1970
1
1
3a
1
1
1
1
1
1
4
1
1
1
1
1
6
1
1
1
1b
1
Yea
1956
1974
1980
1982
1985
1985
1986
1990
1989-1990
1968
1969
1970
1971
1974
1981
1981- 1982
1983
19
89-1990
1970
1981
1986
c Su eau, Ge main, He e, Geo oy, Co ne , Heme & Robin 1977
d Su eau, Tigno & Smi h 1980
e
C icke
a/. 1982
King & C ick 1988
g Me edi h & S anding 1982
h Schneide
e a/.
1985
i Anonymous 1985
j Foggin 1988
k Bishop & Swanepoel 1990
Meba sion e a/. 1992
i us was made om
an
E.
wah/be gi ba ound dead
in
Du ban
in
1990 (Tables
1,
2 and
3)
(Bishop &
Swanepoel 1990), a inding which sugges s ha he
i us is endemic
in
Na al.
A e 1968, Mokola i us was nex isola ed
in
1969
om he ce eb ospinal luid o a young gi l who su -
e ed om non- a al illness wi h e e and seizu es,
and
in
1971
om he b ain o a gi l who died o pa -
aly ic disease, bo h
in
lbadan, Nige ia (Familusi &
Moo e 1972; Familusi, Osunkoya, Moo e, Kemp &
Fabiyi 1972). The possibili y ha he abies- ela ed
i uses we e capable o sp ead among domes ic ca -
ni o es was aised when Mokola i us was isola -
ed om he b ains o six ca s and a accina ed dog
342
Coun y
Lagos, Nige ia
Bozo, Cen al A ican Republic
Pine own/Du ban, Sou h A ica
S ange , Sou h A ica
Kindia, Senegal
Daka , Senegal
Do owa, Zimbabwe
Du ban, Sou h A ica
Addis Ababa, E hiopia
lbadan, Nige ia
lbadan, Nige ia
Umhlanga Rocks, Sou h A ica
lbadan, Nige ia
Yaounde, Came oon
Bulawayo, Zimbabwe
Bulawayo, Zimbabwe
Bo ambi, Cen al A ican Republic
Addis Ababa, E hiopia
Wa mba hs, Sou h A ica
Louis T icha d , Sou h A ica
Esigodini, Zimbabwe
m Kemp e
a/.
1972
n Kemp
e a/.
1974
o Fami
lu
si
& Moo e 1972
p Familu
si
e a/. 1972
Ci cums ances
Su ey
Su ey
Abno mal beha iou
Abno mal beha iou
Su ey
Su ey
Abno mal beha iou
Abno mal beha iou
Abno mal beha iou
Su ey
Non- a al encephali is
Abno mal beha iou
Fa al encepha
li
is
Su ey
Abno mal beha iou
Abno mal beha iou
Su ey
Abno
m
al beha iou
Fa al encephali is
Abno mal beha iou
Su ey
q Le Gonidec, Rickenbach, Robin & Heme 1978
Foggin 1982
s Foggin 1983
Foggin 1985b
u Wik o e a/. 1984
Re e encesc
a,
b
c, d
e-h
i
i
, j
k
I
b,
m,
n
0
h
p
q
j.
- u
j, - u
I
W, X
h
j
Saluzzo, Rollin, Dauga d, Digou e, Geo ges & Su eau 1984
w Me edi h e a/. 1971
x Tigno
e a/.
1977
submi ed om Bulawayo, Zimbabwe,
in
1981 and
1982 o he in es iga ion o suspec ed abies (Ta-
ble
3)
(Foggin 1982; 1983; Wik o , Mac a lan, Foggin
& Kop owski 1984; Fogg
in
1985b; 1988). The ind-
ings
in
Zimbabwe p omp ed e ospec i e in es iga-
ion o a i us which had been isola ed
in
1970 om
a ca
in
Na al, Sou h A ica, and which had been
p ese ed because i eac ed weakly
in
he o iginal
diagnos ic immuno luo escence es wi h an i- abies
conjuga e, and because no abies had been eco d-
ed
in
he a ea o wo yea s (Me edi h 1970). The
isola e was iden i ied as Mokola i us (Table
3)
(Na-
ional Ins i u e o Vi ology 1953-1992, unpublished
eco ds; Schneide
e
a . 1985; King 1991), bu has
also been epo ed as Lagos ba i us (Smi h 1989).
Mokola i us has ne e been isola ed om ba s (Ta-
ble
3}
, and
in
e ospec i is hough ha he ca and
dog in ec ions obse ed
in
Zimbabwe
in
1981 and
1982, we e p obably he esul o spill-o e om an
epidemic
in
small mammals such as sh ews o myo-
mo ph oden s (Fogg
in
1988). Al hough Mokola i us
has no been isola ed
in
sou he n A ica since 1982,
an ibody o he i us was ound
in
oden s in Zimba-
bwe, pa icula ly bush eld ge bils (Ta e a l?ucogas-
e ) (Fogg
in
1988), and i is no able ha i us isola-
ion and cha a e iza ion was no a emp ed
on
he
b ains o wo g ea e cane a s (Th yonomys swinde i-
anus) om he no he n T ans aal which luo esced
in
diagnos ic es s wi h an i- abies conjuga e
in
1985
and 1987 (Tables 1 and
2)
(Swa 1989).
Following i s ini ial isola ion om a human pa ien ,
Du enhage i us was isola ed
in
1981 om a ba ,
possibly
M.
sch eibe sii (Van de Me we 1982),
caugh
in
dayligh by a ca
in
Louis T icha d
in
he
no he n T ans aal (Schneide
e
a/. 1985), and
in
1986 om a common sli - aced ba (Nyc e is heba-
ica) caugh
in
a su ey
in
sou h-eas e n Zimbabwe
(Table
3)
(Foggin 1988).
In
1963, p io o he ecog-
ni ion
o
he abies- ela ed i uses, a i us was iso-
la ed om an
N.
hebaica ba collec ed in a su ey
om a ca e
in
he eas e n T ans aal, bu he isola e
was simply desc ibed as abies i us, p esumably on
he basis
o
his opa hological lesions obse ed
in
mice, and
is
he e o e included
in
Tables 1 and 2,
bu omi ed om Table 3 (Mans el 1965). Fu he -
mo e, immuno luo escence wi h an i- abies conjuga e
was obse ed
in
he b ain
o
an uniden i ied insec i-
o ous ba om he no he n T ans aal
in
1993, bu
no i us could be isola ed o cha ac e iza ion wi h
monoclonal an ibodies, and consequen ly his diag-
nosis is also included
in
Tables 1 and
2,
and omi ed
om Table
3.
Nume ous i uses isola ed
in
sou he n A ica om
he b ains o humans and lowe animals which luo-
esced
in
diagnos ic es s wi h an i- abies conjuga e,
ha e been sc eened wi h monoclonal an ibodies,
and, apa om he cases men ioned abo e (Table
3),
no e idence o in ec ion wi h abies- ela ed i uses
has been ound (Na ional Ins i u e o Vi ology
1953-
1992, unpublished eco ds; King 1991
).
The implica-
ion is ha he abies- ela ed i uses ha e no adap -
ed
o sp ead
in
ca ni o es, bu ha spo adic in ec-
ions may be encoun e ed
in
humans and domes ic
animals, pa icula ly ca s (Table
3)
which a e p eda-
o s o small mammals such as sh ews, myomo ph
oden s and ba s.
The e ha e been nume ous isola ions o ba -associ-
a ed lyssa i uses
in
Eu ope, and a i s i was sus-
pec ed ha Du enhage i us may ha e been in o-
duced wi h ba s om A ica, bu i
is
now clea ha
he e a e wo dis inc sub ypes, Eu opean ba lyssa-
i uses 1 and
2,
associa ed wi h se o ine and myo-
ine ba s espec i ely (King 1991).
R.
SWANEPOEL e a/.
REFERENCES
ADAMSON, J.S. 1954. Ecology o abies
in
Sou he n Rhodesia.
Bulle in o he Wo ld Heal h O ganiza ion, 10:75
3-
759.
ALEXANDER, R.A. 1952. Rabies in Sou h A ica: a e iew o he
p esen posi ion. Jou nal o he Sou h A ican Ve e
in
a y Medical
Associa ion, 23:
135-139
.
ANONYMOUS 1985. Isola ions o Lagos ba i us
in
wes A ica.
In e nal Repo s
o
Cen e Collabo a eu OMS de Re e ence
e Reche che Pou les A bo i us. lns i u Pas eu , Daka , Sene-
gal.
ANONYMOUS 1993. Plena y session o he In e na ional Com-
mi ee o he Taxonomy o Vi uses. /X h In e na ional Cong ess
o
Vi ology, Glasgow, 8-
13
Augus 1993.
BAER, G.
M.
1975a. Rabies
in
nonhema ophagous ba s,
in
The
na u al his o y
o
abies, edi ed by G.M. Bae . New Yo k: Aca-
demic P ess, 2:79-97.
BAER, G.M. 1975b. Bo ine pa aly ic abies and abies
in
he
ampi e ba ,
in
The na u al his o y
o
abies, edi ed by G.
M.
Bae . New Yo k: Academic P ess, 2:
155-175
.
BARNARD, B.J.
H.
1979a. The ole played by wildli e
in
he
epizoo iology o abies
in
Sou h A ica and Sou h Wes A ica.
Onde s epoo Jou nal
o
Ve e ina y Resea ch, 46:155-163.
BARNARD, B.J.H. 1979b. Simp ome an hondsdolheid by huis-
en plaasdie e
in
Suid-A ika en Suidwes-A ika. Jou nal
o
he
Sou h A ican Ve e ina y Associa ion,
50:1
09-111.
BARNARD, B.J.H. & HASSEL, R.H.
1981
. Rabies
in
kudus (T a-
gelaphus s epsice os)
in
Sou h Wes A ica/Namibia. Jou nal
o
he Sou h A ican Ve e ina y Associa ion, 52:309-314.
BARNARD, B.J.H
.,
HASSEL, R.H
.,
GEYER, H.J. & DE KOKER,
W.
C.
1982. Non-bi e ansmission o abies
in
kudu (T agelaphus
s epsice os). Onde s epoo Jou nal
o
Ve e ina y Resea ch, 49:
191
-192.
BERAN, G.W. 1981. Rabies and in ec ions by abies- ela ed
i uses,
in
Vi al zoonoses, edi ed by G.W. Be an, Boca Ra on,
Flo ida: CRC P ess, (CRC Handbook Se ies
in
Zoonoses, Sec-
ion B, ol. 2:57-136).
BINGHAM, J. 1992. Rabies
in
Zimbabwe,
in
P oceedings
o
he
In e na ional Con e ence on Epidemiolog
y,
Con ol
and
P e-
en ion
o
Rabies in Eas e n and Sou he n A ica, Lusaka,
Zambia, 2- 5 June 1992: 29-33.
BLANCOU, J. 1988a. Ecology and _epidemiology o ox abies. Re-
iews
o
In ec ious Diseases, 1 O:S606-S609.
BLANCOU,
J.
1988b. Epizoo iology o abies: Eu asia and A ica,
in
Rabies, edi ed by J.B. Campbell & K.M. Cha l on. Bos on:
Kluwe Academic Publishe s: 243-265.
BOGEL,
K.
& MOTSCHWILLER, E. 1986. Incidence o abies and
pos -exposu e ea men
in
de eloping coun ies. Bulle in
o
he
Wo ld Heal h O ganiza ion, 64:883-887.
BOULGER, L.R. & PORTERFIELD, J.S. 1958. Isola ion
o
a i us
om Nige ian ui ba s. T ansac ions
o
he Royal Socie y
o
T opical Medicine and Hygiene,
52:421-424
.
BRUCKNER, G.K
.,
HURTER, L.R. & BOSHOFF, J.N. 1978. Field
obse a ions
on
he occu ence o abies
in
ca le
in
he magis-
e ial dis ic s o Sou pansbe g and Messina. Jou nal
o
he
Sou h A ican Ve e ina y Associa ion, 49:33-36.
CALISHER, C.G
.,
KARABATSOS, N
.,
ZELLER,
H.
, DIGOUTTE,
J.P
.,
TESH, R.B., SHOPE, R.
E.
, TRAVASSOS DA ROSA,
A.P.A. & ST GEORGE, T.D. 1989. An igenic ela ionships
among habdo i uses om e eb a es and hema ophagous
a h opods. ln e i olog
y,
30:241-257.
CLUVER, E. 1927. Rabies
in
Sou h A ica. Jou nal
o
he Medical
Associa ion
o
Sou h A ica, 1:247-253.
343
Rabies in sou he n A ica
~~
~~~
--~
------------------------------
--------------------
-- -
CRICK,
J.
, TIGNOR, G.H. & MORENO, K. 1982. A new isola e
o Lagos ba i us om he Republic o Sou h A ica. T ansac-
ions
o
he Royal Socie y
o
T opical Medicine and Hygiene, 76:
211~213.
CUMMING, D.H.M. 1982. A case his o y o he sp ead
o
abies
in
an A ican coun y. Sou h A ican Jou nal
o
Science,
78:443~
447.
DE KOCK,
G.
1949.
In
he se ice
o
he li es ock a ming indus-
y: Di isional Repo o Ve e ina y Se ices, in Annual Repo
o he Sec e a y o Ag icul u e. Fa ming
in
Sou h A ica, 25:521
~
523.
DEPNER,
K.
1992. Rabies
in
Namibia,
in
P oceedings
o
he In-
e na ional Con e ence on Epidemiology, Con ol and P e en ion
o
Rabies
in
Eas e n and Sou he n A ica, Lusaka, Zambia,
2~5
June 1992:
39-44.
DIAS, P.T
.,
NOVOA, A.M. & CLIFF, J.
L.
1985. Rabies
in
Mozam-
bique,
in
Rabies
in
he opics, edi ed by
E.
Kuwe ,
C.
Me ieux,
H.
Kop owski &
K.
Bagel. Be lin: Sp inge -Ve lag:
406~414.
DIAS, P.
T.
1992. Rabies
in
Mozambique,
in
P oceedings
o he
In e na ional Con e ence on Epidemiology, Con ol and P e en-
ion
o
Rabies
in
Eas e n and Sou he n A ica, Lusaka, Zambia,
2~5
June 1992:
24~25.
DUARTE, V.M.S., ROSLIACOV, A.A., NSALAMBI, D , GOMES,
A.F. 1985. Epidemiologie e diagnos ic de
Ia
age en Angola,
in
Rabies
in
he opics, edi ed by
E.
Kuwe ,
C.
Me ieux,
H.
Kop owski &
K.
Bagel. Be lin: Sp inge -Ve lag:
406~414.
DU
TO IT, P.J. 1929.
In
P oceedings
o
he Pan A ican Ag icul-
u al and Ve e ina y Cong ess, P e o ia, Augus 1929:
272
~
284.
DU
TO IT, P.J. 1936. Wild ca ni o a as ca ie s o abies. Qua e ly
Bulle in
o he
Heal h O ganiza ion
o
he League
o
Na ions,
5:
162
~
165
.
EDDINGTON,
A.
1895. Repo o he Di ec o o he Colonial Bac-
e iology Ins i u e o he yea 1894. Depa men o Ag icul u e,
Cape
o
Good Hope.
EDMONDS, C.R. 1922. Diseases
o
Animals in Sou h A ica. Lon-
don: Baillie e, Tindall & Cox:
195~212.
FAMILUSI, J.B. & MOORE, D.L. 1972. Isola ion o a abies- ela ed
i us om he ce eb ospinal luid o a child wi h asep ic me-
ningi is. A ican Jou nal
o
Medical Science,
3:93
~
96.
FAMILUSI, J.B., OSUNKOYA, B.O
.,
MOORE, D.L
.,
KEMP, G.E.
& FABIYI,
A.
1972. A a al human in ec ion wi h Mokola i us.
Ame ican Jou nal
o
T opical Medicine and Hygiene,
21
:
959
~
963.
FERNANDES, M.V. & ARAMBULO Ill, P.V. 1985. Rabies as an
in e na ional p oblem,
in
Wo ld's deb
o
Pas eu , edi ed by H.
Kop owski & S.A. Plo kin. New Yo k: Alan
R.
Liss:
187~218.
FITZSIMONS, F.W. 1919. Na u al his o y
o
Sou h A ica, London:
Longmans G een, 2:7.
FOGGIN, C.M. 1982. A ypical abies i us in ca s and a dog in
Zimbabwe.
The
Ve e ina y Reco d, 110:338.
FOGGIN, C.M. 1983. Mokola i us in ec ion in ca s and a dog
in
Zimbabwe. Ve e ina y Reco d, 113:115.
FOGGIN, C.M. 1985a. The epidemiological signi icance o jackal
abies
in
Zimbabwe,
in
Rabies in he opics, edi ed by
E.
Ku-
we ,
C.
Me ieux,
H.
Kop owski &
K.
Bagel. Be lin: Sp inge -Ve -
lag:
399-405.
FOGGIN, C.M. 1985b. Mokola i us in ec ion
in
domes ic ca s
in
Zimbabwe,
in
Rabies
in
he opics, edi ed by E. Kuwe , C
Me ieux,
H.
Kop owski &
K.
Bagel. Be lin: Sp inge -Ve lag:
65 ~
70.
FOGGIN, C.
M.
& SWANEPOEL,
R.
1985. Rabies
in
A ica wi h
emphasis on abies- ela ed i uses,
in
Wo ld's deb
o
Pas eu ,
edi ed by H. Kop owski, & S.A. Plo kin. New Yo k: Alan R. Liss:
219
~
234
.
344
FOGG
IN
, C.M. 1988. Rabies and abies- ela ed i uses in Zimba-
bwe: his o ical, i ological and ecological aspec s. Ph.D hesis:
Uni e si y
o
Zimbabwe, Ha a e.
GUMMOW,
B.
1985. Rabies on he ise. Depa men
o
Heal h
and Wel a e, P e o ia, Epidemiological Commen s,
12(6):2
~
23.
GUMMOW
, B. & TURNER, 1986. Rabies in Sou h A ica: Epide-
miological ends o he pe iod
1980~1984.
Jou nal
o
he Sou h
A ican Ve e ina y Associa ion,
57:231~237.
HASSEL, R.H. 1982. Incidence o abies in kudu in Sou h Wes
A ica/Namibia. Sou h A ican Jou nal
o
Science,
78:418
~
421.
HENNING, M.W. 1956. Rabies,
in
Animal diseases
in
Sou h A i-
ca
, 2nd ed. Cape Town: Cen al News Agency:
975
~
1022
.
HERZENBERG,
L.
1928. Two cases o hyd ophobia. Jou nal
o
he Medical Associa ion
o
Sou h A ica, 2:659-661 .
HLATSHWAKO,
P.
1992. Rabies
in
Swaziland,
in
P oceedings
o
he In e na ional Con e ence on Epidemiology, Con ol and
P e en ion
o
Rabies
in
Eas e n and Sou he n A ica, Lusaka,
Zambia,
2~5
June 1992:
43~44.
HUBSCHLE, O.J.B. 1988. Rabies
in
he kudu an elope (T agela-
phus s epsice os). Re iews
o
In ec ious Diseases, 1
O:S629
~
S633.
HUTCHEON,
D.
1894. Repo s o he Colonial Ve e ina y Su geon
and Assis an Ve e ina y Su geons o he yea 1893. Depa -
men
o
Ag icul u e, Cape
o
Good Hope.
KEIGHTLEY, A.l. STRUTHERS, J.
K.
, JOHNSON, S & BAR-
NARD, B.J.
H.
1987. Rabies in Sou h A ica:
1980
~
84
.
Sou h
A ican Jou nal
o
Science, 83:466-472.
KEMP, G.E , CAUSEY, O.R., MOORE. D.L , ODEOLA,
A.
& FA-
BIYI,
A.
1972. Mokola i us. Fu he s udies on I bAn 27377, a
new e iologic agen o zoonosis
in
Nige ia. Ame ican Jou nal
o
T opical Medicine and Hygiene,
21
:356~359.
KEMP, G.E
.,
OTTIS, R.C., SETZER, H.W. & MOORE, D.L. 1974.
Isola ion o i uses om wild mammals
in
Wes A ica. Jou nal
o
Wildli e Diseases, 1
0:279
~
293
.
KENNEDY, D.J. 1988. An ou b eak
o
abies
in
no h-wes e n Zim-
babwe
1980~1983.
Ve e ina y Reco d,
122:129~133
.
KHOMARI,
L.
1988. Rabies
in
Leso ho,
in
P oceedings
o
he
In e na ional Con e ence on Epidemiology. Con ol and P e-
en ion
o
Rabies and B ucellosis
in
Eas e n and Sou he n
A ican Coun ies, Gabe one, Bo swana,
23
~
25
No embe 1988:
25~26
.
KHOMARI,
L.
1992. Rabies
in
Leso ho,
in
P oceedings
o
he
In e na ional Con e ence on Epidemiology. Con ol and P e-
en ion
o
Rabies in Eas e n and Sou he n A ica. Lusaka.
Zambia,
2
~
5
June 1992:
45
~
46
.
KING, A. & CRICK,
J.
1988. Rabies- ela ed i uses, in Rabies.
edi ed by J.B. Campbell & K.M. Cha l on. Bos on: Kluwe Aca-
demic Publishe s:
177
~
200
.
LAMBRECHTS, M.C. 1964. A b ie summa y o he sani a y posi-
ion
in
he Republic
o
Sou h A ica. Bulle in
de
/'O ice In e -
na ional des Epizoo ies,
62
:
799
~
805
.
LAWRENCE, J.A., FOGGIN, C.M. & NORVAL, R.A.I. 1980. The
e ec s o wa on he con ol
o
diseases o li es ock
in
Rhodesia
(Zimbabwe). Ve e ina y Reco d, 1
07:82
~
85.
LE GONIDEC, G
..
RICKENBACH. A
..
ROBIN. Y & HEME. G.
1978. lsolemen d'une souche de i us Mokola au Came oun.
Annali di Mic obiologia.
129A:245
~
249
.
LOPES PEREIRA, C.M , PINTO, F.G. & BAULE. C 1988. Rabies
in
Mozambique: upda e,
in
P oceedings
o
he In e na ional Con-
e ence on Epidemiology. Con ol and P e en ion
o
Rabies and
B ucellosis
in
Eas e n and Sou he n A ican Coun ies. Gabe-
one, Bo swana,
23~25
No embe 1988:
37
~
47
.
MACKINNON, J. 1963. Rabies
in
Sou he n Rhodesia 1950-1962.
Bulle in de /'O ice In e na ional des Epizoo ies, 60:87-95.
MAGANU, E.T. & STAUGARD,
F.
1985. Epidemiology o abies
in
Bo swana,
in
Rabies
in
he opics, edi ed by
E.
Kuwe , C. Me-
ieux,
H.
Kop owski &
K.
Bagel. Be lin: Sp inge -Ve lag: 422-
428.
MANSVELT, P.
R.
1956. Rabies
in
he no he n T ans aal (1950)
ou b eak. Jou nal
o
he Sou h A ican Ve e ina y Medical As-
socia ion, 27:167-178.
MANSVEL T, P.R. 1962. Rabies
in
Sou h A ica. Field con ol o
he disease. Jou nal
o
he Sou h A ican Ve e ina y Medical As-
socia ion, 33:313-319.
MANSVEL T, P.R. 1965. The ole o wild li e
in
he epizoo iology
o some in ec ious and pa asi ic animal diseases
in
Sou h A ica.
Bulle in
de
/'O ice In e na ional des Epizoo ies, 64:825-835.
MARE, C.J. 1962. Rabies
in
Sou h A ica. The epizoo iology and
diagnosis o he disease. Jou nal
o
he Sou h A ican Ve e ina y
Medical Associa ion, 33:287-294.
MASUPU, K.V. 1992. Rabies
in
Bo swana,
in
P oceedings
o
he
In e na ional Con e ence on Epidemiology, Con ol and P e en-
ion
o
Rabies
in
Eas e n and Sou he n A ica, Lusaka, Zambia,
2- 5 June 1992: 34-38.
MEBATSION,
T.
, COX, J.H. & FROST, J.W. 1992. Isola ion and
cha ac e iza ion o 115 s ee abies i us isola es om E hiopia
by using monoclonal an ibodies: iden i ica ion o 2 isola es as
Mokola and Lagos ba i uses. Jou nal
o
In ec ious Diseases,
166:972-977.
MEREDITH, C.D. ROSSOUW, A.P. & VAN PRAAG KOCH, H.
1971.
An
unusual case o human abies hough o
be
o chi op-
e an o igin. Sou h A ican Medical Jou nal, 45:767-769.
MEREDITH, C.
D.
1977. Recen s udies
on
abies
in
sou he n A i-
ca,
in
Medicine
in
a opical en i onmen , edi ed by J.H.S Gea .
Cape Town: A.A. Balkema: 387-397.
MEREDITH,
C.
D.
& STANDING, E.
1981
. Lagos ba i us
in
Sou h
A ica. Lance , 1 :832-833.
MEREDITH, C.D. 1982. Wildli e abies: pas and p esen
in
Sou h
A ica. Sou h A ican Jou nal
o
Science, 78:
411
-415.
MOSIENYANE, M.G. 1988. Rabies si ua ion
in
Bo swana,
in
P o-
ceedings
o
he
In e na ional Con e ence on Epidemiology, Con-
ol and P e en ion
o
Rabies and B ucellosis
in
Eas e n and
Sou he n A ican Coun ies, Gabe one, Bo swana, 23-
25
No-
embe 1988: 9- 13.
NEITZ, W.O. 1965. A check-lis and hos -lis o he zoonoses oc-
cu ing
in
mammals and
bi
ds
in
Sou h and Sou h-Wes A ica.
Onde s epoo Jou nal
o
Ve e ina y Resea ch, 32:189-337.
NEITZ, W.O. & MARAIS, I.P. 1932. Rabies as i occu s in he
Union o Sou h A ica,
in
Eigh een h epo
o
he Di ec o
o
Ve e ina y Se ices and Anima/Indus y, Union
o
Sou h A ica:
71
-89.
NEITZ, W.O. & THOMAS, A.D. 1933. Rabies
in
SoL
' h A ica. Oc-
cu ence and dis ibu ion o cases du ing 1932. Onde s epoo
Jou nal
o
Ve e ina y Science and Animal Indus y, 1 :
51
-56.
NEITZ, W.O. & THOMAS, A.D. 1934. Rabies
in
Sou h A ica. Oc-
cu ence and dis ibu ion o cases du ing 1933. Onde s epoo
Jou nal
o
Ve e ina y Science and Anima/Indus y, 3:335-342.
NELSON, H. 1962. Rabies
in
Sou h A ica. Rabies om he poin
o iew o he medical o ice o heal h o medical p ac i ione .
Jou nal
o
he Sou h A ican Ve e ina y Medical Associa ion, 33:
327-340.
RAZA, O.A.E., WENHOLD, B.A
.,
HOWARD,
P.
MARAIS,
A.
&
PALLED,
J.
1992. Rep oduc ion
in
he yellow mongoose e isi -
ed.
Sou h A ican Jou nal
o
Zoology, 27:192-195.
R.
SWANEPOEL e a/.
SACRAMENTO, D., BOURHY, H. & TOR DO,
N.
1991. PCR ech-
nique as an al e na i e me hod o diagnosis and molecula epi-
demiology o abies i us. Molecula and Cellula P obes, 5:
229-240.
SALUZZO, J.F., ROLLIN, P.E , DAUGARD,
C.
,
DIGOUTIE
, J P ,
GEORGES, A.J. & SUREAU,
P.
1984. P emie isolemen
du
i us Mokola a pa i d'un ongeu (Lophu omys sikapusi). An-
nales de 1'/ns i u Pas eu : Vi ologie, 135E:57--66.
SCHNEIDER, H.P. 1985. Rabies
in
Sou h Wes A ica/Namibia,
in
Rabies in he opics, edi ed by E. Kuwe ,
C.
Me ieux, H.
Kop owski & K. Bagel. Be lin: Sp inge -Ve lag: 520-535.
SCHNEIDER, L.G., BARNARD, B.J.H. & SCHNEIDER, H.
P.
1985.
Applica ion o monoclonal an ibodies o epidemiological in es i-
ga ions and o al accina ion s udies.
I.
A ican i uses,
in
Rabies
in
he opics, edi ed by
E.
Kuwe , C. Me ieux, H. Kop owski &
K. Bagel. Be lin: Sp inge -Ve lag: 47-59.
SCOTI
, C.
D.
1988. Rabies
in
Leso ho. Ve e ina y Reco d, 122:
287.
SHAW, J.J.H. 1980. Hondsdolheid by koedoes. Jou nal
o
he
Sou h A ican Ve e ina y Associa ion,
51
:209.
SHONE, O.K. 1962. Rabies
in
Sou he n Rhodesia 1900-1961 .
Jou nal
o
The Sou h A ican Ve e ina y Medical Associa ion, 33:
567-580.
SHOPE, R.E., MURPHY, F.A., HARRISON, A.K., CAUSEY, O.R.,
KEMP, G.E
.,
SIMPSON, D.I.H. & MOORE, D.L. 1970. Two A i-
can i uses se ologically and mo phologically ela ed o abies
i us. Jou nal
o
Vi ology, 6:690-692.
SINYANGWE, P.
G.
1992.1nP oceedingso heln e na iona!Con-
e ence on Epidemiology, Con ol and P e en ion
o
Rabies
in
Eas e n and Sou he n A ica, Lusaka, Zambia, 2- 5 June 1992:
19-21.
SMITH, J.S. & BAER, G.M. 1988. Epizoo iology o abies: he
Ame icas,
in
Rabies, edi ed by J.B. Campbell & K.M. Ca l on.
Bos on: Kluwe Academic Publishe s: 267-299.
SMITH, J.S. 1989. Rabies i us epi opic a ia ion: use
in
ecologic
s udies. Ad ances
in
Vi us Resea ch, 36:215-253.
SMITH, J.S
.,
FISHBEIN, D. B
.,
RUPPRECHT,
C.
E. & CLARK,
K.
1991. Unexplained abies
in
h ee immig an s
in
he Uni ed
S a es. New England Jou nal
o
Medicine, 324:205-211.
SMITHERS, R.H.N. 1983. Mammals
o
he sou he n A ican sub e-
gion. P e o ia: Uni e si y o P e o ia.
SNYMAN, P.S. 1937. Rabies
in
Sou h A ica. Jou nal
o
he Sou h
A ican Ve e ina y Medical Associa ion, 8:126-133.
SNYMAN, P.S. 1940. The s udy and con ol o he ec o s o a-
bies
in
Sou h A ica. Onde s epoo Jou nal
o
Ve e ina y
Science and Anima/Indus y, 15:9- 140.
SNYMAN, P.
S.
1953. Rabies
in
he Union o Sou h A ica. Bulle in
o
Epizoo ic Diseases
o
A ica, 1 :94- 1 05.
SUREAU, P
.,
GERMAIN, M
.,
HERVE, J.P., GEOFFROY,
B.
,
CORNET, J.P
.,
HEME, G. & ROBIN, Y. 1977. lsolemen du i us
Lagos-ba en Empi e Cen a icaine. Bulle in de
Ia
Socie e de
Pa hologie Exo ique, 70:467-470.
SUREAU, P., TIGNOR, G.H. & SMITH, A.L. 1980. An igenic cha -
ac e iza ion o he Bangui s ain (ANCB-672d) o Lagos ba i us.
Annals
o
Vi ology,
131
:25-32.
SWANEPOEL, R. 1994. Rabies,
in
In ec ious diseases
o
li es ock
wi h special e e ence
o
sou he n
A
ica, edi ed by
JAW
.
Coe ze , G.R. Thomson, R.
C.
Tus in & N.P.
J.
K iek. Cape Town:
Ox o d Uni e si y P ess Sou he n A ica:
in
p ess.
SWANEPOEL, R. & FOGGIN, C.M. 1978. The occu ence, diag-
nosis, ea men and con ol o abies
in
Rhodesia. Cen al
A ican Jou nal
o
Medicine, 24:107-
11
3.
345

Rabies in sou he n A ica
SWART
, 1989. Hondsdolheid
in
Suid-A ika. Ve med, Sou h A i-
can Ve e ina y Medicine,
2:163-166.
TARR,
A.F., O'GRADY, J.M. &
JENKINS,
W.L. 1962. Rabies in
Sou h
A ica
. Rabies and
he
p i a e p ac i ione . Jou nal
o
he
Sou h A ican Ve e ina y Medical Associa ion,
33:323
-325.
TIGNOR,
G.H.
MURPHY,
F.A.,
CLARK
, H.F.,
SHOPE,
R.E.,
MA-
DORE
,
P.
,
BAUER
, S.P
.,
BUCKLEY
, S.M. &
MEREDITH
, C. D.
1977.
Du enhage
i us: mo phological, biochemical, his o-
pa hological and an igenic ela ionships
o
he
abies se og oup.
Jou nal
o
Gene al Vi ology,
37
:
595-611
.
TREMLETT
, 1993. Rabies in Bo swana, a e iew
1977-1992.
Gabo one
: Depa men
o
Animal Heal h and P oduc ion, Minis y
o
Ag icul u e.
TUCHILI,
L.M. 1988.
Epidemiology
and p e en ion
o
abies
in
Zambia
, in P oceedings
o
he In e na ional Con e ence on Epi-
demiology, Con ol and P e en ion
o
Rabies and B ucellosis in
Eas e n and Sou he n A ican Coun ies, Gabe one, Bo swana,
23
-2
5
No embe
1988:
61-65
.
TUSTIN,
R.C. & SMIT, J.D. 1962. Rabies
in
Sou h A ica: an anal-
ysis
o
his ological examina ion. Jou nal
o
he Sou h A ican
Ve e ina y Medical Associa ion,
33:295
-310.
VALADAO
, F.G. 1968.
The
mos
impo an
aspec s
o
he abies
p oblem
in
Mozambique.
Ve e ina ia Mozambicana,
1:65-70
.
VANDER
MERWE,
M. 1982. Ba s
as
ec o s
o
abies. Sou h
A ican Jou nal
o
Science,
78:421-422.
VAN
RENSBURG
, R.D.J. 1965. P elimina y epo on he "hu-
mane
coyo e-ge e " o he con ol
o
he black-backed jackal
Thos mesomelas
in
he T ans aal. Zoologica A icana,
1:193-
198.
VON
MAL
TITZ,
L.
1950. Rabies in he no he n dis ic s
o
Sou h
Wes
A ica. Jou nal
o
he Sou h A ican Ve e ina y Medical As-
socia ion,
21
:
4-12.
WEN
HOLD
, B.A.
1990
.
The
e hology and social s uc u e
o
he
yellow
mongoose
(Cynic is penicilla a). M.Sc. hesis, Uni e si y
o
P e o ia.
WHO
1989. Gene a: Ve e ina y Public Heal h Uni , Di ision
o
Communicable
Diseases, Wo ld Heal h O ganiza ion (Wo ld su -
ey
o
abies; no.
23
[ o he yea s
1986
/1987]).
346
WHO
1991.
Gene a
: Ve e ina y Public Heal h Uni , Di ision
o
Communicable
Diseases, Wo ld Heal h O ganiza ion (Wo ld su -
ey
o
abies; no.
24
[ o he
yea
1988]).
WIKTOR
, T.J. &
KOPROWSKI,
H. 1978.
Monoclonal
an ibodies
agains
abies i us p oduced by
soma ic
cell hyb idiza ion: de-
ec ion
o
an igenic a ian s. Immunology,
75:3938
-
3942.
WIKTOR,
T.J.,
FLAMAND
,
A.
&
KOPROWSKI
,
H.
1980.
Use
o
monoclonoal an ibodies
in
diagnosis
o
abies i us in ec ion and
di e en ia ion
o
abies and abies- ela ed i uses. Jou nal
o
Vi ological Me hods, 1:
33
-46.
WIKTOR,
T.J.,
MACFARLAN
, R.I.,
FOGGIN
, C.M. &
KOPROW-
SKI,
H.
1984
. An igenic analysis
o
abies and
Mokola
i us om
Zimbabwe
using monoclonal an ibodies. De elopmen s
in
Biolo-
gical S anda diza ion,
57:199-211
.
WILLIAMSON
, J.M. 1976.
The
his o y
o
abies
in
Rhodesia
and
con ol measu es. Rhodesia Science News, 10:221-
222.
ZUMPT
,
I.
F.
&
DEBRUYN,
H.W. 1967.
The
me i s
o
"Phos oxin"
in
he e adica ion
o
small Vi e idae. Jou nal
o
he Sou h
A ican Ve e ina y Medical Associa ion,
38:456-457.
ZUMPT
, I.F. 1968.
The
eeding habi s
o
he
yellow
mongoose,
Cynic is penicilla a, he su ica e, Su ica a su ica a and
he
Cape
g ound
squi el, Xe us inau is. 1. Analysis
o
s omach
con en s.
2.
Obse a ions on ee-li ing and cap i e animals. Jou nal
o
he Sou h A ican Ve e ina y Medical Associa ion,
39:89-91
.
ZUMPT
, I.F. 1969. Fac o s in luencing abies ou b eaks: he age
and b eeding cycle
o
he yellow
mongoose
, Cynic is penicilla a
(G. Cu ie ). Jou nal
o
he Sou h A ican Ve e ina y Medical
Associa ion, 40:319-322.
ZUMPT
,
I.
F.
1976.
The
yellow
mongoose
(Cynic is penicilla a) as
a la en
ocus
o
abies
in
Sou h A ica. Jou nal
o
he Sou h
A ican Ve e ina y Medical Associa ion,
47:211-213.
ZUMPT
, I.F. 1982.
The
yellow
mongoose
as a abies
ec o
on
he cen al pla eau
o
Sou h A ica. Sou h A ican Jou nal
o
Science,
78:417
-418.
ZYAMBO
, G.C.N.,
SINYANGWE
, P.G. &
BUSSEIN
, N.A. 1985.
Rabies
in
Zambia
, in Rabies
in
he opics, edi ed by
E.
Kuwe ,
C.
Me ieux, H. Kop owski &
K.
Bagel. Be lin: Sp inge -Ve lag:
415
-421.