Journal of Helminthology (1977) SI, 229—252

SUPPLEMENTARY REVIEW ARTICLE

A check list of definitive hosts exhibiting evidence of the genus Schistosoma Weinland, 1858 acquired naturally in Africa and the Middle East R. J. PITCHFORD Bilharzia Field Unit, S.A. Medical Research Council, Nelspruit, South Africa

Le Roux (1958a) proposed division of the Genus Schistosoma into six based on the small number of common generic morphological characteristics and because the Genus was overloaded. It seems that no useful purpose is served by this division and this list retains the Genus Schistosoma Weinland 1858. (See also Nelson et al, 1962; Round, 1968). He also proposed, based on intermediate host susceptibility and morphological characteristics of eggs and females from a monkey, that the name Schistosoma capense (Harley, 1864) be reintroduced for the causal parasite of urinary bilharzia in South Africa. In the light of subsequent work on intermediate host susceptibility and egg morphology of African Schistosoma, acceptance might necessitate considerable reclassification and division into innumerable species. Clearly this is not desirable. Amberson and Schwarz (1953) proposed a trinomial nomenclature for the African schistosomes with terminal spined eggs. Acceptance of this proposal would likewise necessitate the creation of numerous subspecies to accommodate, and further confuse, any understanding that might exist of, these schistosomes. Le Roux (1958a) and apparently Round (1968) supported Schistosoma curassoni as a valid species. However Le Roux (1961) stated that it is probable that it is either S. mattheei or S. bovis. The present author agrees and for the purpose of this list S. curassoni is relegated to "species of doubtful validity" requiring further investigation and confirmation of the new generic status proposed by Gretillat (1962). Bhalerao (1932) placed it in synonymy with S. bovis. See also under S. curassoni. The two schistosomes of the hippopotamus, S. hippopotami and S. edwardiense, are retained as valid species. But the substantiating evidence possibly resulting from physiological and anatomical adaptations to the environment of an abnormal host, would seem to be outweighed by the apparent inability of the parasites to continue their life-cycle. See also under these respective Schistosoma sp. There are several reports from North Africa refuting the findings of S. bovis in man. Further south, where S. mattheei is endemic, several reports claim S. bovis infections in man. This is possibly due to the use of the name S. bovis where "bovine" would have been more appropriate, or to justifiable misidentification, or to reports written before S. mattheei was first described in 1929. Certainly some reports of S. bovis from southern Africa are clearly referring to S. mattheei. 229

R. J. PITCHFORD Apparently there is sufficient evidence to place the validity of S. mattheei beyond doubt (Alves, 1949; Dinnik and Dinnik, 1965; Pitchford 1965; Taylor, 1970). If this is accepted, there is evidence that S. bovis and S. mattheei co-exist within the same territory or within the same definitive host (Dinnik and Dinnik, 1965 in Kenya and Tanzania; Graber, 1969, Birgi and Graber, 1969 and Delpy et ah, 1972 in Tchad) but that hydridization between the two is unlikely (Dinnik and Dinnik, 1965; Taylor, 1970). Thus there seems no reason why some observations (van den Berghe 1937b and others in Zaire) should not be classified as mixed infections of these two parasites. (See also under S. mattheei). Reports indicate that S. mattheei and S. bovis overlap in more or less a continuous strip from Nigeria, through Tchad, Zaire, and Ruanda Urundi to southern Kenya, Tanzania and northern Zambia. In southern Tanzania and northern Zambia a third animal schistosome with terminal spined eggs, S. leiperi, overlaps S. bovis and S. mattheei. South of this S. bovis falls away and only S. leiperi and S. mattheei overlap. S. mattheei falls away towards thesouthwest leaving S. leiperi as the only animal schistosomes with terminal spined eggs in East Caprivi, northern Botswana and probably parts of south-western Zambia. North-west of East Caprivi, in Angola, reports are scanty and only S. bovis has been reported. S. leiperi falls away to the south and south-east leaving S. mattheei as the only animal schistosome in Rhodesia, South Africa, Eastern Botswana and Mocambique. Excluding Okavango swamp in northern Botswana and Humansdorp area in the southern Cape there is no evidence of any schistosome south of L 19°00 S and west of L 25c00 E. Towards the south, this area extends east to include the whole of the Orange Free State in South Africa and the Kingdom of Lesotho which is the only country in Africa which can lay claim to no endemic Schistosoma spp. No records could be found of schistosomes in animals from Algeria, Benin, Cameroon, Gabon, Ghana, Ivory Coast, Libya, Liberia, Mauritius, Spanish Sahara, and Togo. The list includes reports of infections acquired in Africa but discovered elsewhere, and also those which might have resulted from ingestion of eggs; it excludes infections acquired by exotic species in artificial environments (zoos) in Africa. Mammalian nomenclature follows the Smithsonian Institution "Preliminary manual for African mammals" as far as possible, in conjunction with Dorst and Dandelot (1972) "A field guide to the larger mammals of Africa" and Roberts (1954) "Mammals of South Africa". 1. SCHISTOSOMA HAEMATOB1UM (Bilharz 1852). Type host Homo sapiens DEFINITIVE HOST AND/OR AS REPORTED

Genus and species

LOCALITY AND AUTHORITY

Vernacular &/or as reported

Primates

Homo sapiens Linnaeus 1758

230

Man

Iran, Iraq, Turkey, Syria, Lebanon, Saudi Arabia, Yemen, South Yemen, Israel. All countries in Africa including Malagassy and Mauritius but excluding Lesotho, Ruanda Urundi and Spanish Sahara. (Wright 1973)

REMARKS

Definitive hosts for Schistosuma species DEFINITIVE HOST AND/OR AS REPORTED

Pan troglodytes Blumenbach 1779 Papio sp. ? P. papio (Desmarest 1822)

LOCALITY A N D AUTHORITY

REMARKS

P. satyrus U.S.A. ex Sierra Leone (de Paoli 1965) Chimpanzee Senegalese baboon Senegal (Taylor et al. 1972)

Passaged with recovery of S. haematobium (presumably) eggs. P. anubis P. doguera Kenya (Nelson 1960; Nelson Mixed with S. mattheei. et Dog faced baboon al. 1962) (J. P. Fisher 1829) P. rhodesiae Rhodesia (Purvis et al. 1965) P. ursinus Chacma baboon (Kerr 1792) Vervet monkey Kenya (Nelson et al. 1962) Cercopithecus (aethiops) pygerethrus C. aethiops (F. Cuvier 1821) Artiodactyla Sus scrofa Linnaeus 1758 Ovis aries Linnaeus 1758 Syncerus caffer (Sparrman 1779)

Large White Boar Tamworth Sow Pig Sheep

Nigeria (Hill & Onabamiro 1960)

African/Cape buffalo

South Africa (Basson et al. One female only. 1970)

Rhodesia (McKenzie 1970)

Mixed with S. mattheei.

2. SCHISTOSOMA BO VIS (Sonsino 1876) Type host Bos taurus Primates Homo sapiens

Man

None passaged experimentally with recovery of S. bovis eggs. Unless otherwise stated all mixed with 5. haematobium.

Perissodactyla Equus caballus Linnaeus 1758

Horse/Equines

E. (Asinus) asinus Linnaeus 1758

Donkey

E. (A.) asinus x E. caballus

Mule

Egypt (Mahfouz 1927; Soliman 1956, 1962) Uganda (Raper 1951)

Cited and doubted by Tadros and Abdelghani (1974) and others. Eggs resembled S. leiperi more than S. bovis (Dinnik & Dinnik 1965). Pure infection.

South Africa (Cawston 1921, 1921. Before 5. mattheei de1930; Kisner et al. 1953; de scribed Drawings resemble S. Meillon et al. 1953) mattheei. 1953? Same infection; pure; passaged through Bulinus (Physopsis) sp. with subsequent recovery of 1 male schistosome in mouse. 1930, 1953. Probably mistaken identity. Probably mistaken identity. Zaire (Walravens 1931) Sudan (Abdel-Malek 1969)

Iraq (MacHattie & Chadwick 1932) Sudan (Abdel-Malek 1959a, 1960, 1961, 1969; Soliman 1962) Tchad (Graber 1969; Birgi & Graber 1969) Iraq (MacHattie & Chadwick 1932) Somalia (Sobrero 1960) Sudan (Soliman 1962; AbdelMalek 1961, 1969) Tchad (Graber 1969; Birgi & Graber 1969) Iraq (MacHattie & Chadwick 1932) 231

R. J. PITCHFORD DEFINITIVE HOST AND/OR AS REPORTED

Artiodactyla Bos taurus Linnaeus 1758 (Probably includes B. indicus in part)

Cattle, Cow, Ox

Bos taurus

Bos indicus Linnaeus 1758

Zebu cattle

Bos sp. Ovis aries

Cattle Sheep

232

LOCALITY AND AUTHORITY

REMARKS

Iran (Arfaa et al. 1965) Iraq (MacHattie & Chadwick 1932) Egypt (Sonsino 1876, Soliman 1962 and others) Tunisia (Anderson & Gobert 1936) Morocco (Blanc & Desportes 1936) Mauritania (Marill 1961) Senegal, Niger, Guinea, Mali, Presumably cattle. ? B. indicus ? B. taurus. See also under S. Haute Volta (Gillet 1956b) sp. Senegal. Gillet states "5. bovis has been reported from" these countries but gives no reference. Guinea Bissau (Gillet 1956a) Nigeria (Cowper 1966) Mixed with S. mattheeiinpart. Tchad (Delpy et al. 1972) Sudan (Soliman & le Roux 1955 5. sp. Amended Soliman 1955; Soliman 1962; Abdel- (1962). Malek 1959a, I960, 1961, 1969; Eisa 1963a, b; Dinnik & Dinnik 1965) Ethiopia (Dinnik & Dinnik 1965 not mentioned in text. 1965; Holmes et al. 1975; Lo & Lemma 1975) Somalia (Pellegrini 1953; Sobrero 1960; Dinnik & Dinnik 1965) 1965 not mentioned in text. Congo Republic(Graber 1975) Zaire (van den Berghe 1934, 1937 mixed with S. mattheei. 1937b; Gillet 1950; Fain & Lagrange 1952; Schwetz 1955) Kenya 1965 Mixed with S. mattheei: S. bovis predominant sp. (Dinnik & Dinnik 1965) Uganda (Trap. vet. Bull. 1913; 1913 reported Bilharzia crassa. Dinnik & Dinnik 1965) Ruanda Urundi (Wery 1950; 1950? mixed with 5. mattheei. van den Berghe 1934) Tanzania (Dinnik & Dinnik Mixed with S. leiperi S. bovis 1965) predominant sp. Zambia (Dinnik & Dinnik Mixed with 5. leiperi or S. 1965) mattheei. Angola (Janz & Carvalho 1956) Mocambique (Travassos San- Probably mistaken for S. tos Dias 1954; Azevedo et al. mattheei. See Cruz e Silva 1954) (1971) under S. mattheei. Mocambique. Tchad (Graber 1969; Birgi & Mixed with 5. mattheei in part. Graber 1969) Sudan (Hussein et al. 1975) Kenya (Round 1962) Iran (Arfaa et al. 1965) Iraq (MacHattie & Chadwick 1932) Israel (Olafson 1955, cited by Lengy 1962) Egypt (Sonsino 1877, cited by Soliman 1962; Soliman 1962) Ethiopia (Holmes et al. 1975) Mauritania (Marill 1961) Sudan (Abdel-Malek 1959a, 1960, 1961, 1969; Soliman 1962; Gameel 1974)

Definitive hosts for Schistosoma species DEFINITIVE HOST AND/OR AS REPORTED

LOCALITY A N D AUTHORITY

REMARKS

Tchad (Graber 1969; Birgi & Graber 1969; Delpy et al. 1972) Tanzania (Dinnik & Dinnik 1965)

Mixed with 5. mattheei in part.

Kenya (Round 1962) Zaire (van den Berghe 1934)

Capra hircus Linnaeus 1758

Goat

Camelus dromedarius Camel, Baladi/ Linnaeus 1758 Dromedary camel

S. scrofa

Pig

Phacochoerus aethiopicus (Pallas 1776) Bubalus bubalis Linnaeus 1758

Warthog

Mixed with S. leiperi or S. mattheei. S. bovis predominant sp.

Rhodesian (? Zambian) origin. South Africa (Cawston 1930) Probably mistaken for S. mattheei. Iraq (MacHattie & Chadwick 1932) Somalia (Sobrero 1960) Sudan (Abdel-Malek 1961; Soliman 1962) Mauritania (Marill 1961) Tchad (Graber 1969; Birgi & Graber 1969) Tanzania (Dinnik & Dinnik Mixed with S. leiperi. 1965) Zaire (van den Berghe 1934) Rhodesian (? Zambian) origin. Sudan (Soliman 1955, 1962; Abdel-Malek 1959a, b, 1960, 1961, 1969) Egypt (Soliman 1955, 1962) Somalia (Sobrero 1960) Tchad (Graber 1969; Birgi & Graber 1969) Sudan (Abdel-Malek 1959a, 1960, 1961, 1969) Senegal (Gretillat, 1969) Morphology and eggs reported similar to S. bovis.

Egypt (Nagati 1942, cited by The scarcity of positive findHussein 1973) ings from this animal is interesting. Massoud (1973) failed to infect B. bubalis experimentally; also Arfaa et al. (1965) found no natural infections in Iran. Tragelaphus spekei Limnotragus spekei Ruanda (van den Berghe 1934, 1937a) Sclater 1864 Situtunga, Water kudu Damaliscus korrigum Topi Tchad (Graber 1969) (Ogilby 1833) Kobus (Adenotd) kob Adenota kob Sudan (Myers et al. 1960) (Erxleben 1777) leucotis Kob Tchad (Graber 1969) Defassa waterbuck Tchad (Graber 1969) K. defassa (Ruppell 1826) Redunca redunca Bohor reedbuck Tchad (Graber 1969) (Pallas 1766) Hippolragus equinus Roan antelope Tchad (Graber 1969) (Desmarest 1804) Sylvicapra grimmia S. grimmi Somalia (Sobrero, 1975) Claimed correctly as first Grimm's duiker (Linnaeus, 1758) record of S. bovis in wild herbivore in East Africa, if Ruanda is considered Central Africa, but incorrectly as first record of S. bovis in mammal other than domestic ruminants. (See above). Water buffalo

233

R. J. PITCHFORD DEFINITIVE HOST AND/OR AS REPORTED

Rodentia Praomys {Mastomys) natalensis (Smith 1834) Lophuromys flavopunctatus Thomas 1888

Mastomys natalensis Multimammate mouse Swamp rat

LOCALITY AND AUTHORITY

REMARKS

Kenya (Nelson et al. 1962)

Kenya (Nelson et al. 1962)

Mixed with S. rodhaini.

3. SCHISTOSOMA MATTHEEl Veglia and le Roux 1929 Type host Ovis aries Vernacular &/or Genus and species as reported Primates South Africa (Harley 1864; 1864 depicted as S. capense Man Homo sapiens Cawston 1925, 1930; Porter 1925, 1926, 1930 reported as 1926; Pitchford 1959b, 1965) S. bovis, S. spindalis or S. None passaged or passaged without recovery of typical S. mattheei eggs. spindalis var africana. Unless otherwise stated all mixed Swaziland (Gauldie undated) with S. haematobium, some with South West Africa (Porter Locality doubtful. Reported S. mansoni in addition. 1926) S. spindale var africana. Rhodesia (Blackie 1932; Alves 1958; Goldsmid 1974) Zambia (Hira 1975) Pure or mixed with S. mansoni. Hira implies a scarcity of S. mattheei in Zambia by incorrectly claiming this to be the first report of S. mattheei in Zambia. (See under) Pure cases of S. mattheei in man are rare and even more rare in the gut. He discusses the possibility of these cases being S. intercalatum, but does not mention the prevalence of S. mattheei in domestic or other animals in the vicinity. Botswana (Pitchford 1961) Zaire (van den Berghe 1937b) By implication. See footnote. Nigeria (Ramsay 1934) One pure case. Not stated (le Roux 1947) Baboon (dog faced) Kenya (Nelson 1960; Nelson Mixed with S. haematobium Papio anubis et al. 1962) and pure. P. doguera Tanzania (Nelson 1960) Rhodesia (Blackie 1932) Chacma baboon P. ursinus South Africa (Pitchford et al. P. porcarius 1973, 1974) Baboon Papio sp. Not stated (le Roux 1947) Kenya (Nelson 1960; Nelson Cercopithecus (mitis) Sykes Blue et al. 1962) albogularis monkey (Sykes 1831) C. mitis C. (aethiops) South Africa (Pitchford et al. Vervet monkey pygerethrus 1973) Perissodactyla Rhodesia (presumably) Horse Equus caballus (Lawrence 1976) Zebra, Burchell's South Africa (Pitchford 1961; Equus (Hippotigris) Basson et al. 1971; Pitchford Zebra burchelli Gray 1824 et al. 1973, 1974) Artiodacryla South Africa (Veglia & le Cattle Bos taurus Roux 1929; Pitchford 1959b, 1961; Strydom 1963; Dinnik & Dinnik 1965 and others) Swaziland (Gauldie undated) Botswana (Pitchford 1965) 234

Definitive hosts for Schistosoma species DEnNITIVE HOST AND/OR AS REPORTED

B. indicus

Zebu cattle

O. aries

Sheep

C. hircus

Goat

Warthog Phacochoerus aethiopicus Potamochoerus porcus Bushpig (Linnaeus 1758) Giraffa camelopardalis Giraffe (Linnaeus 1758) Tragelaphus Greater kudu strepsiceros (Pallas 1766) T. scriptus Bushbuck (Pallas 1766) T. angasi Nyala Gray 1849 Hippotragus niger Sable Antelope (Harris 1838)

LOCALITY AND AUTHORITY

REMARKS

Rhodesia (Blackie 1932; Condy 1960; Lawrence & Condy 1970; Dinnik & Dinnik 1965 and others) Zaire (van den Berghe 1934, 1936, 1937b) Mocambique (Pitchford & le Roux 1955a; Cruz e Silva 1971)

Egg measurements of 1960 observations very long 201298 nm mean 248 /an. Possibly swollen eggs from bladder. 1937 as S. bovis var mattheei.

Malawi (Dinnik & Dinnik 1965; Pitchford unpublished data) Zambia (le Roux 1931a, b, 1933; Dinnik & Dinnik 1965) Tanzania (Dinnik & Dinnik 1965) Kenya (Dinnik & Dinnik 1965) Ruanda Urundi (van den Berghe 1934) Not stated (le Roux 1947) Tchad (Graber 1969; Birgi & Graber 1969; Delpy et al. 1972) South Africa (Veglia & le Roux 1929; le Roux 1929; Pitchford 1959b, 1961, 1965; Strydom 1963; Hurter & Potgieter 1967 and others) Rhodesia (Blackie 1932; Lawrence & Condy 1970; MacKenzie 1970 and others) Mocambique (Cruz e Silva 1971) Botswana (Blackie 1932) Zambia (le Roux 1931a, b, 1933, 1957; Dinnik & Dinnik 1965) Not stated (le Roux 1947) Tchad (Graber 1969) South Africa (Pitchford 1959b, 1961; Hurter & Potgieter 1967; Pitchford et al. 1973) Mocambique (Cruz e Silva 1971) Not stated (le Roux 1947) South Africa (Pitchford et al. 1974) South Africa (Pitchford et al. 1974) South Africa (Pitchford et al. 1973, 1974) South Africa (Pitchford et al. 1973, 1974)

Cruz e Silva (1971) does not admit the presence of S. bovis in Mocambique. Similarly Dinnik & Dinnik (1965) did not depict S. bovis in Mocambique in their map.

1965 observations mixed with S. leiperi and S. bovis. Mixed with predominant S. bovis. Mixed with predominant S. bovis.

Mixed with S. bovis.

South Africa (Pitchford et al. 1973) South Africa (Pitchford et al 1974) South Africa (Pitchford et al. 1973, 1974) 235

R. J. PITCHFORD DEFINITIVE HOST AND/OR AS REPORTED

H. equinus

Roan Antelope

Kobus ellipsiprymnus Common waterbuck (Ogilby 1833) K. (Adenoid) vardoni Puku (Livingstone 1859) Adenota leche, K. (Adenota) leche Lechwe Gray 1850 Comochaetes (Gorgon) Blue wildebeest taurinus (Burchell 1823) Redunca arundinum Common reedbuck (Boddaert 1785) Aepyceros melampus Impala (Lichtenstein 1812)

Bovidae Syncerus caffer

Unspecified antelopes African/Cape buffalo

LOCALITY AND AUTHORITY

REMARKS

Rhodesia (Pitchford et al. Imported into Kruger National Park, South Africa ex 1974) Wankie National Park, Rhodesia, 3 weeks previously. South Africa (Basson et al. 1971; Pitchford et al. 1973, 1974) Rhodesia (Pitchford 1974) Zambia (le Roux 1938) Zambia (leRoux, 1938,1958b) Mixed with S. leiperi and 5. margrebowiei. South Africa (Ortlepp 1961, Pitchford 1961, Pitchford et al. 1973,1974; Basson et al. 1971) Zambia (le Roux 1938) South Africa (Pitchford & le Roux 1955b, Ortlepp 1961; Pitchford 1961; Basson et al. 1971; Pitchford et al. 1973, 1974) Zambia (le Roux 1933) South Africa (Young & van den Heever 1969; Basson et al. 1970; Pitchford et al. 1973, 1974)

Rodentia South Africa (Pitchford 1959a; Praomys (M.) Multimammate Pitchford & Visser 1962) natalensis mouse South Africa (Pitchford & Vlei rat Ototnys angoniensis Wroughton 1906 Visser 1962) Lemniscomys griselda Single-striped field South Africa (Pitchford & (Thomas 1904) mouse Visser 1962) Tatera leucogaster Schinz's Gerbil South Africa (Pitchford 1959a) (Peters 1852) van den Berghe (1937b) was unable from his observations "to justify S. mattheei as a new species". For present purposes this is immaterial and the facts that emerge from his paper are he (1) found schistosomes in cattle from Katanga (2) was able to differentiate between S. bovis and S. mattheei among these and (3) found evidence in 3-5 % of patients of "bovine type" eggs in urine, identical to those described by Blackie (1932) from Southern Rhodesia. No authenticated reports of 5. bovis in animals or man are known from Rhodesia and it is therefore felt that van den Berghe was dealing with mixed infections of S. bovis and S. mattheei in the cattle and with S. mattheei and/or a hybrid S. haematobium x 5. mattheei in the humans. S. Africa includes Transkei 5. mattheei in man and cattle (Pitchford & Geldenhuys 1960). 4. SCHISTOSOMA INTERCALATUM Fisher 1934. Type host Homo sapiens Vernacular &/or Genus and species as reported Primates Zaire (Fisher 1934; van Wijk Several other reports cited by Man Homo sapiens Fisher (1934) refer to terminal 1975) spined ova in stools published Schwetz (1956) was uncertain of the before his original description. identity of the schistosome he reNotably Chesterman (1923). ported from Hybomys and this record is therefore included under "Schistosoma sp. or author doubtful" Zaire, Gabon and Cameroon Gabon (van Wijk 1975) all transmission foci (van Wijk 1975; Deschiens & Delas Cameroon (Deschiens & Delas 1969). 1969; Wright et al. 1972; van Wijk 1975) Senegal, Upper Volta, Tchad, Isolated cases only (Wright et Central African Republic, al. 1972; van Wijk 1975). Congo, Nigeria (Wright et al. 1972; van Wijk 1975) 236

Definitive hosts for Schistosoma species 5. SCHISTOSOMA LEIPERI le Roux 1955. Type host Tragelaphus spekei DEFINITIVE HOST AND/OR AS REPORTED

Genus and species

LOCALITY AND AUTHORITY

REMARKS

Vernacular &/or as reported

Perissodactyla

E. caballus

Horse

E. (H.) burchelli

E. quagga zambeziensis, Zebra

Zambia (le Roux 1931 b, 1947) 1931 b as 5. spindalis. Claimed as first record of schistosomiasis in horse in Africa (le Roux 1931b). Zambia (le Roux 1955a, 1957) 1955 locality not stated mixed with S. margrebowiei.

Artiodactyla (Zambia and East Caprivi mixed with 5. margrebowiei in Zambia (le Roux 1931a, b, Cattle Bos taunts 1933, 1947; Dinnik & Dinnik 1965) East Caprivi (Pitchford 1976)

Ovis aries

Sheep

Capra hircus

Goat

Tragelaphus spekei

T. spekei selousi, Situtunga, Water kudu

Kobus ellipsiprymnus Common waterbuck K. (A.) leche

A. (A.) vardoni

Onotragus leche, Adenota leche, Onotragus smithemani, Red & Black lechwe, Lechwe

Puku

Damaliscus lunatus Tsessebe (Burchell 1823) Redunca arundinum Common Reedbuck Oribi Ourebia ourebi (Zimmerman 1783) ? Sylvicapra grimmia Duiker (Linnaeus 1758) Tragelaphus scriptus Bushbuck Greater kudu T. strepsiceros Grysbuck Raphiceros sharpei Thomas 1897 Alcelephus lichtensteiniHartebeest (Peters 1849) or possibly A. caama (G. Cuvier 1804)

part). Mixed with 5. mattheei. 1931a, b as S. spindalis.

Predominantly unisexual male infections. Tanzania (Dinnik & Dinnik Mixed with S. bovis only. 1965) Zambia (le Roux 1931a, b, 1931 as S. spindalis. 1933) Tanzania (Dinnik & Dinnik Mixed with 5. bovis only. 1965) Zambia (le Roux 1931b, 1947, le Roux (1961), amended 1955c, 1957) previous reports of 5. spindale and some Schistosoma sp. to S. leiperi. Not stated (Ortlepp 1961) Also listed as S. spindalis. ? East Caprivi Zambia (le Roux 1957) Botswana (Pitchford 1974) Zambia (le Roux 1931b, 1938, Reported S. spindale(is) 1931b, 1947, 1955a, 1957, 1958b; 1946,1938. See le Roux (1961) Buckley 1946) above. 1946. Not mixed. 1958 also mixed with S. mattheei. 1957 as 2 lechwe species. Botswana (Pitchford 1974; Carmichael et al. 1977) East Caprivi (Pitchford 1976) South Africa and not stated Listed as spindale in S.A. ? (Ortlepp 1961) East Caprivi (lechwe not found in S.A.). Zambia (le Roux 1931b, 1938, Reported 5. spindalis in part. 1957) Botswana (Pitchford 1974) Botswana (Unpublished data)

Zambia (le Roux 1931b, 1938, Reported S. spindalis in part. 1957) Botswana (Pitchford 1974) Not stated (Ortlepp 1961) Listed as 5. spindale. Zambia (le Roux 1957) Zambia (le Roux 1957) Zambia (le Roux 1957) Zambia (le Roux 1957) Zambia (le Roux 1957) Zambia (le Roux 1957)

237

R. J. PITCHFORD DEFINITIVE HOST AND/OR AS REPORTED

Connochaetes taurinus Hippotragus equinus Aepyceros melampus Taurotragus oryx (Pallas 1766) Syncerus coffer

LOCALITY AND AUTHORITY

Blue wildebeest Roan antelope Impala Eland (Cape)

Zambia Zambia Zambia Zambia

African buffalo

Zambia (le Roux 1957) Botswana (Carmichael et al. 1977) Zambia (le Roux 1933,1955c)

Bovidae (Unspecified)

(le Roux (le Roux (le Roux (le Roux

REMARKS

1957) 1957) 1957) 1957)

spp. herbivora unspecified le Roux (1955c) states that S. leiperi parasitized 21 species of herbivore and lists 1 {T. spekei) as type host without naming the other 20. Prior to 1955 he had incriminated E. caballus (1931b, 1947), E. burchelli (1955a), K. leche (1931b, 1938, 1947), K. vardoni, R. arundinum (1931b, 1938), B. taurus (1931a, b, 1933, 1947) and O. aries (1931a, b, 1933) as hosts of S. leiperi (amended in part le Roux 1961). In 1957 he lists 18 species (K. leche as 2) of wild herbivore, including the above 5 as harbouring schistosomes (unspecified) of domestic stock (unspecified but presumably cattle, sheep and horse as above) in Zambia. This list together with cattle, sheep and horse gives the requisite 21 species of herbivore which are therefore included as harbouring 5. leiperi in Zambia. le Roux(1931a, b, 1933,1947,1955a)specified, in part, cattle and sheep as hosts of S. mattheei, cattle, sheep and horse as hosts of S. leiperi, and cattle and horse as hosts of 5. margrebowiei in Zambia. In the present author's opinion it is not clear and is most unlikely, that the 18 species of wild herbivore listed by le Roux (1957) as harbouring schistosomes of domestic stock, all harboured all 3 of these schistosomes as intimated by Round (1968). 6. SCHISTOSOMA EDWARDIENSE Thurston 1964. Type host Hippopotamus amphibius Genus and species Hippopotamus amphibius Linnaeus 1758

Vernacular &/or as reported Hippopotamus

Uganda (Thurston 1963,1964, Not passaged experimentally. 1971) No extra uterine eggs seen. No spine seen on intra uterine eggs: mesenteric vein parasite: No miracidia seen from gut scrapings or faeces (Thurston 1971).

7. SCHISTOSOMA MANSONI Sambon 1907. Type host Homo sapiens Primates

Homo sapiens

Man

Pan troglodytes

Chimpanzee

Papio sp. ? anubis P. anubis

Baboon P. doguera Dog faced baboon

P. ursinus

Chacma baboon

P. papio

West African baboon

238

Saudi Arabia, South Yemen, Records incomplete for many Yemen, Israel; all countries in countries (Wright 1973). Africa including Malagasy but excluding Algeria, Guinea Bissau, Lesotho, Mauritania, Mauritius, Morocco, Niger, Somalia, Spanish Sahara, and Tunisia (Wright 1973) U.S.A. ex Sierra Leone (Hsu & Hsu 1968) Tanzania (Taylor et al. 1972) Tanzania (Fenwick 1969) Uganda (Nelson I960, Fripp 1963 Pers. commun.) Kenya (Miller 1959a, b, 1960, Nelson 1960, Nelson et al. 1962) South Africa (Pitchford et al. 1973) Rhodesia (Goldsmid 1974) U.S.A. ex West Africa (McQuay 1952)

Definitive hosts for Schistosome species DEFINITIVE HOST AND/OR AS REPORTED

Cercopithecus (aethiops) pygerethrus

Insectivora Crocidura luna Dollman 1910 C. oliviera (Lesson 1827) Carnivore Canis familiaris Linnaeus 1758 Artiodactyla Ovis aries Kobus ellipsiprymnus Rodentia Gerbillus pyramidum I. Geoffroy 1825 Praomys (M.) natalensis

Pelomys (P.) fallax (Peters 1852)

Grivet monkey

Lophuromys flavopunctatus

REMARKS

U.S.A. ex Ethiopia (Cheever "It seems likely that our monet al. 1970) keys were infected before U.S.A. ex Tanzania (Cheever capture, although we cannot be sure that they were not et al. 1970) exposed in Africa while being held for shipment." (Cheever et al. 1970). However in view of the generally water-shy habits of C. aethiops, the comparatively high rate of infection and high worm recovery from both groups natural infections might appear a little doubtful.

Katanga red musk Zaire (Stijns 1952) shrew Egyptian musk Egypt (Kuntz 1958) shrew

Not typical S. mansoni.

Dog

Kenya (Nelson et al. 1962)

Sheep Common waterbuck

Rhodesia (McKenzie 1970) South Africa (Pitchford et al. Eggs all dead. 1974)

Egyptian Gerbil

Egypt (Kuntz 1952)

M. coucha, M. natalensis.

Zaire (Schwetz 1953, 1954)

Mastomys sp., Multimammate mouse P. fallax grooved tooth swamp rat

Kenya (Nelson et al. 1962) South Africa (Pitchford 1959a; Pitchford & Visser 1962) Zaire (Schwetz 1953, 1954, Reported S. mansoni var 1956) rodentorum.. Mixed with S. rodhaini or pure. Zaire (Schwetz 1954, 1956) Reported S. mansoni var rodentorum,. Mixed with S. rodhaini. Zaire (Schwetz 1953, 1954, Reported S. mansoni var 1956) rodentorum.. Mixed with S. rodhaini or pure. Zaire (Schwetz 1953, 1954, Reported S. mansoni var 1956) rodentorum.. Mixed with 5". rodhaini or pure. Zaire (Schwetz 1955a)

Oenomys hypoxanthus O. hypoxanthus (Pucheron 1855) Dasymus incomtus (Sundervall 1847)

LOCALITY AND AUTHORITY

D. bentleyae African marsh or water rat L. aquila. Swamp rat

Black rat Rattus rattus Linnaeus 1758 Lemniscomys griselda Single-striped field mouse Otomys angoniensis O. tugelensis, Otomys sp. Vlei rat. Arvicanthis niloticus Nile rat (Desmarest 1822)

Mixed with S. rodhaini.

Reported S. mansoni var rodentorum. Mixed with 5. rodhaini or pure.

South Africa (Pitchford & Visser 1962) South Africa (Pitchford 1959a; 1962 mixed with S. mattheei. Pitchford & Visser 1962) Kenya (Nelson et al. 1962) Egypt (Mansour 1973) Mixed with one "terminal spined" male.

8. SCHISTOSOMA RODHAINI Brumpt 1931 Carnivore

Canis familiaris

Dog

Felis (Leptailurus Serval cat serval) Schreber 1776

Ruanda Urundi (Fain et al. Mixed with S. mansoni and 1953;Derameee/a/. 1953) Kenya (Nelson et al. 1962) pure. Zaire (Huygelen 1957) Ruanda Urundi (Deramee et al. 1953; Fain et al. 1953) 239

R. J. PITCHFORD DEFINITIVE HOST AND/OR AS REPORTED

Rodentia Lophwomys flavopunctatus

L. aquila(us) Swamp rat

Praomys (P.) jacksoni P. jacksoni (de Winton 1897) P. {P.) morio Rattus (P.) moris (Trouessart 1881) P. (M.) natalensis M. coucha, R. (M.) coucha multimammate mouse Pelomys (P.)fallax P.frater,P.fallax, P. fallax concolor Groove tooth swamp rat Otomys irroratus Vlei rat Brants 1827 Malacomys longipes Milne Edwards 1877 Thallomys paedulcus (Sundervall 1846) Dasymys incomtus

M. I. centralis T. nigricauda Black tail rat D. bentleyae, African marsh or water rat

Oenomys hypoxanthus

LOCALITY AND AUTHORITY

REMARKS

Zaire (Stijns 1952; Schwetz 1954) Kenya (Nelson et al. 1962) Mixed with S. bovis. Zaire (Stijns 1952) Uganda (Fripp & Goodman 1967) Zaire (Stijns 1952; Schwetz 1952a, b, 1954) Ruanda Urundi (Fain et al. 1953) Zaire (Stijns 1952; Schwetz 1952a, b, 1954, 1956) Ruanda Urundi (Fain et al. 1953) Ruanda Urundi (Fain et al. 1953) Zaire (Gillet 1960) Uganda (Fripp & Goodman 1967) Kenya (Nelson et al. 1962)

Schwetz unmixed or mixed with 5. mansoni var rodentorum. Schwetz unmixed or mixed with S. mansoni var rodentorum.

Zaire (Schwetz 1952a, b, 1954, Schwetz mixed with 5. man1966; Gillet 1960) soni var rodentorum or pure. Zaire (Schwetz 1954) Mixed with S. mansoni var Ruanda Urundi (Fain et al. rodentorum. 1953)

9. SCHISTOSOMA HIPPOPOTAMI Thurston 1963. Type host Hippopotamus amphibius H. amphibius

Hippopotamus

Uganda (Thurston 1963,1971) Not passaged. No extrauterine eggs seen. South Africa (McCulIy et al. "Viable" eggs in sectioned 1967) suprarenals only. Adults mainly in bizarre sites e.g. heart, pulmonary arteries, posterior vanae cavae etc. ? Abnormal host.

10. SCHISTOSOMA MARGREBOWIEIle Roux 1933 Primates Homo sapiens

Man

None passaged experimentally. See also under S. sp. ingested. Perissodactyla Equus cabal/us E. (H.) burchelli

Mali (Lapierre & Hein 1973) Zambia (Cawston 1930)

Recta! biopsy. Mixed with S. mansoni and S. haematobium. Reported S. japonicum in Durban ex Rhodesia.

Horse Zebra, E. quagga zambesiensis

Zambia (le Roux 1947) Zambia (le Roux 1933, 1955a, Locality not stated 1947) Mixed with S. leiperi. Zaire (van den Berghe 1934)

Artiodactyla Tragelaphus spekei T. strepsiceros

T. spekei selousi, Kudu

Hippotragus equinus H. niger

Roan antelope Sable antelope

Kobus (A.) leche

Adenota leche, Kobus leche, Onotragus leche,

Zambia (le Roux 1933) Botswana (Carmichael et al. 1977) Zambia (le Roux 1933) Botswana (Carmichael et al. 1977) Zambia (le Roux 1933, 1938, Locality not stated 1955. 1947, 1955a, 1958b) Mixed with S. leiperi. 1958 mixed with S. mattheei.

240

1955.

Definitive hosts for Schistosoma species DEFINITIVE HOST AND/OR AS REPORTED

Lechwe, Lechwe kobs

A. (A.) vardoni

Puku, Cobus vardoni

K. defossa

Defassa waterbuck Common waterbuck Kob Adenota kob Blue wildebeest Brindled gnu Tsessebe

K. ellipsiprymnus K. (A.) kob Connochaetes (G.) taurinus Damaliscus lunatus Redunca arundinum

Common reedbuck

Aepyceros melampus Impala

REMARKS

LOCALITY AND AUTHORITY

Zaire (van den Berghe 1934, 1937) Botswana (Pitchford 1974) East Caprivi (Pitchford 1974 and unpublished) Not stated (Ortlepp 1961) ? East Caprivi Zambia (le Roux 1933, 1938) Botswana (Pitchford, 1974; Carmichael et al. 1977) Tchad (Graber 1969)

Mixed with S. leiperi. Mixed with S. leiperi.

1974 Mixed with S. leiperi. Mixed with S. bovis.

Botswana (Pitchford 1974)

Mixed with S. leiperi.

Tchad (Graber 1969) Zambia (le Roux 1933)

Mixed with S. bovis.

Botswana (Carmichael et al. 1977; Unpublished data) Zambia (le Roux 1933, 1938) Botswana (Pitchford 1974, Carmichael et al. 1977) Botswana (Carmichael et al. 1977) Botswana (Carmichael et al. 1977) Zambia (le Roux 1933, 1947) East Caprivi (Pitchford 1976 and unpublished)

Mixed with S. leiperi. 1974 Mixed with S. leiperi.

Syncerus caffer

Buffalo

Bos taurus

Cattle

Genus and species

11. SCHISTOSOMA SPP. unspecified or doubtful identity Vernacular &/or as reported

Primates Homo sapiens

Man

Predominantly unisexual male infections. Mixed with S. leiperi. South West Africa (Pitchford Unisexual male infection (imported ?) unpublished) Capra hircus Goat East Caprivi (Unpublished data) le Roux (1933) mentions that S. margrebowiei was invariably found with other Schistosoma spp., S. spindalis (amended le Roux 1961) in the majority of hosts (unspecified) and S. mattheei in a few cases (unspecified).

Cercopithecus Vervet monkey (aethiops) pygerethrus C. aethiops, C. pygere thrus

Zaire (van den Berghe 1934, S. mattheei type: van den Berghe (1937b) apparently 1936) differentiated between S. mattheei and S. bovis but pronounced the schistosome as 5. bovis var mattheei conforming in all respects to S. bovis as modified by MacHattie & Chadwick (1932). As far as can be made out these authors never recognized S. mattheei as a distinct species. See also under 5. mattheei. Identified tentatively as S. Nigeria (Okpala 1961) bovis. Kenya (Nelson 1960; Nelson Single male 5". haematobiutn] et al. 1962) bovis type ? same. South Africa (Porter 1938)

Cercocebus sp. Papio anubis

Egg identity doubtful ? S. mansoni. (Cobbold Bilharzia magna. See under S. sp. of doubtful validity.

U.K. ex Africa Sooty monkey 1859) (Cercopithecus fuliginosus Kenya (Nelson 1960) P. doguera Dog faced baboon

5. haematobium complex. 241

R. J. PITCHFORD DEFINITIVE HOST AND/OR AS REPORTED

LOCALITY AND AUTHORITY

REMARKS

Papio sp.

Baboon

East Africa (Nelson & Pester S. haematobium and S. mat1961) theei. One male schistosome!

Artiodactyla Bos taurus

Cattle

Bos indicus Bos sp. (presumed)

Zebu cattle Cattle

Swaziland (Pitchford 1958a, 1958 Reported identical to Dir. Med. Ser. I960, 1963, cattle schistosomes in Transvaal. Now considered S. 1967) mattheei. 1960, 1963, 1967 reported S. mattheei or S. bovis. South Africa (Pitchford 1958b) Schistosoma sp. Now considered S. mattheei. Bovine schistosomes. Sudan (Archibald 1933) Ethiopia (Daynes & Graber Schistosoma sp. 1974) Nigeria (Cowper 1966) Reported S. spindale. Zambia (le Roux 1931a, b) Schistosoma sp. Zaire (van den Berghe 1934) See remarks under H. sapiens. Rhodesian (? Zambia) origin. Tanzania (Kinoti 1963; Dinnik 1963 S. bovis complex. 1965 & Dinnik 1965) Intermediate S. bovisj S. leiperi. Ruanda Urundi (van den See above remarks. Berghe 1934) Reported bovine schistosomes. Kenya (Dowdeswell 1938) Malagasy (Horchner & Kar- No extra or intra-uterine ova seen. First report of schistorasch 1972) somes in cattle in Malagasy. Schistosoma sp. Senegal (Vassiliades 1974) Central Africa Republic, Schistosomes of bovis group. Gambia and other countries already listed (WHO Consultants 1956) South Africa (Pitchford 1958b) See remarks above. Zaire (van den Berghe 1934) See remarks above. Schistosoma sp. Zambia (le Roux 1931a) South Africa (Pitchford 1958b) See remarks above. S. bovis/S. Tanzania (Dinnik & Dinnik Intermediate leiperi. 1965) Zaire (van den Berghe 1934) See remarks above. S. bovis type. Kenya (le Roux 1955b) Schistosomes similar to S1. Egypt (Soliman 1962) mattheei. South Africa (Pitchford et al. Eggs resembled large 5. haematobium. ? S. mattheei. 1974) "haematobium—bovis" group. Ruanda (Dinnik et al. 1963) Reported S. bovis type. CalciU.K. ex Tropical Africa fied eggs only. (Porter 1944) South Africa (Basson et al. Typical liver lesions only in 1971) section. Rhodesia (Pitchford 1974) Probably S. leiperi but eggs short (167-232 fim mean 200). Schistosoma sp. Zambia (le Roux 1931b)

Ovis aries Capra hircus

Goat

Camelus dromedarius Camel Hippopotamus amphibius

Hippopotamus

Giraffa camelopardalis Giraffe

Kobus ellipsiprymnus

Waterbuck

K. (A.) leche, Redunca Lechwe, arundinum Reedbuck Tragelaphus spekei Situtunga

Rodentia Arvicanthus niloticus

Nile rat

Otomys angoniensis

Vlei rat

242

Listed as Rhodesia but 18.25 Schistosomes South 23.40 East is East Caprivi (Pike & Condy 1966) Egypt (Mansour 1973)

Reported 5. haematobium. Single male mixed with S. mansoni. South Africa (Pitchford 1959a) Reported S. haematobium. Hybrid S. mattheei x S. haematobium on passage (Pitchford 1961).

Definitive hosts for Schistosoma species DEFINITIVE HOST AND/OR AS REPORTED

Hybomys univittatus (Peters 1876)

H. univittatus

Dasymus incomtus

African marsh or water rat

LOCALITY AND AUTHORITY

REMARKS

Zaire (Schwetz 1956)

Reported S. intercalatum, but "the question of an initial natural infection of wild rats with S. intercalatum is thereforeunsettled" (Schwetz 1956). Kenya (Teesdale & Nelson Single male. Reported S. 1958; Nelson 1960; Nelson et mansoni type: ? all the same. 1962 schistosome sex not al. 1962) stated.

12. SCHISTOSOMA SPP. Probably or possibly ingested Genus and species

Vernacular &/or as reported

Primates Homo sapiens

Man

Nigeria (Cowper & Woodward 1961) Zambia (Buckley 1946) Zaire (d'Haenens & Santele 1955;Walkiers 1928)

£. bovis.

S1. leiperi reported S. spindalis. 1955 S. rodhaini. Egg viability doubted. Fripp (1968); 1928? S1. margrebowiei: reported 5. faradjei. See S. sp. of doubtful validity. East Caprivi (Pitchford 1976 S. margrebowiei. & unpublished) Rhodesia (Goldsmid et al. S. mattheei. 1976) Sudan (Teesdale 1976) S. bovis. Tanzania (MacMahon 1969a) S. bovis. Tchad (Delpy et al. 1972) S. bovis and S. mattheei.

Carnivora Panthera leo Linnaeus 1758 Canis familiaris

Artiodactyla Sus scrofa

South Africa (Pitchford et al. 5. mattheei. 1974) S. mansoni. One dead egg also Tanzania (Mango 1971) reported in liver digest. ? "swept—through biliary system" (Mango 1971). South Africa (Pitchford et al. S. mansoni. 1973)

Lion Dog

Pig

South Africa (Pitchford et al. S. mansoni, S. haematobium, 1973) S. mattheei. Senegal (Lariviere et al. 1960) S. mansoni.

13. SCHISTOSOMA SPP. of doubtful validity or of doubtful occurrence in Africa Schistosoma capense Harley 1864.

Generally considered synonymous with southern strain of S. haematobium. Depicted together with eggs resembling S. mattheei. Reported from man, South Africa.

Schistosoma curassoni Brumpt 1931. Genus and species Vernacular &/or as reported Homo sapiens Man

Senegal (Gretillat 1962b)

Passaged through sheep with recovery of S. curassoni.

B. taurus Domestic cattle Mali (Brumpt 1931) B. taurus, B. indicus, Cattle, Zebu, Mali, Senegal, Mauritania O. aries, C. hircus Sheep, Goats (Gretillat 1962a, 1964) "During 1928 and 1929 M. G. Curasson . . . discovered a schistosome in the mesenteric and portal veins of domestic catt'e from Bamakau (Mali). This species . . . resembles Schistosoma mattheei very closely". "Unfortunately having only 2 females at my disposal and not knowing the morphology of the mature eggs . . . I shall describe the parasite under the provisional name of Schistosoma curassoni, hoping to describe it more fully when more material is at my disposal" (Brumpt 1931). 243

R. J. PITCHFORD Unfortunately no more material apparently became available. However in order to accommodate S. curassoni, Gretillat (1962a) proposed a new genus Proschistosoma, based on developmental differences in the intramolluscan stages. As far as is known this has never been confirmed and the situation in West Africa remains confused regarding the identity of this schistosome. The validity of S. curassoni remains in doubt and "it seems probable that it may be either S. bovis Sonsino 1876 or S. mattheei" (le Roux 1961). Schistosoma faradjei ? Synonymous with 5. margrebowiei or unrecognized entity (le Roux Walkiers 1926 1961; Nelson 1960). Reported from man, Zaire. Abnormal eggs which should no longer be considered as occurring in S. indicum (Montgomery 1906) Africa (le Roux 1961). Reported from zebra and antelopes, Zambia, (le Roux 1931b, 1933). Schistosoma japonicum Misidentification before S. margrebowiei described. Reported from man (Katsurada 1904) ex Rhodesia (Cawston 1930). Schistosoma magna Unrecognized entity (Amberson & Schwarz 1953; le Roux 1961; Nelson Cobbold 1859 1960). Schistosoma mansoni var Synonymous with S. mansoni (Nelson 1960, Nelson et al. 1962; rodentorum Schwetz 1954 Pitchford & Visser 1960). Reported from rodents, Zaire. Schistosoma spindalis Misidentification before S. mattheei Veglia & le Roux 1929 and/or S. leiperi le Roux 1955 described. Reported from man, stock and antelopes, (Montgomery 1906) South Africa, Zambia (Cawston 1925; le Roux 1933; Buckley 1946 and others). Amended le Roux (1961). Schistosoma spindalis var Synonymous with S. mattheei (le Roux 1961). Reported from man, South africana Porter 1926. and South West Africa. ACKNOWLEDGEMENT Thanks are due to the South African Medical Research Council for permission to publish this paper.

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R. J. PITCHFORD FAIN, A. and LAGRANGE, E. (1952) Un foyer be bilharziose bovine a Schistosoma bovis dans PIturi. Atmales des Societes beiges de Medicine tropicale, 32, 49-51. FAIN, A., THIENPOINT, D., HERIN, V. and DERAMEE, O. (1953) Observations sur Schistosoma rodhaini Brumpt an Ruanda Urundi. Annales des Societes beiges de Medicine tropicale, 33,423-443. FENWICK, A. (1969) Baboons as reservoir hosts of Schistosoma mansoni. Transactions of the royal Society of tropical Medicine and Hygiene, 63, 557-567. FISHER, A. C. (1934) A study of schistosomiasis of the Stanleyville district of the Belgian Congo. Transactions of the royal Society of tropical Medicine and Hygiene, 28, 277-306. FRIPP, P. J. and GOODMAN, J. D. (1967) New foci of Schistosoma rodhaini Brumpt in Uganda. Transactions of the royal Society of tropical Medicine and Hygiene, 61, 613-614. FRIPP, P. J. (1968) Some observations on the behaviour of the Kampala strain of Schistosoma rodhaini Brumpt in the laboratory. South African Journal of medical Sciences, 33, 21-30. GAMEEL, A. A. (1974) A clinical pathological study on naturally occurring Schistosoma bovis infection in Sudanese sheep. Ada veterinaria, Jugoslavia, 24, 277-285. Cited in Veterinary Bulletin (1975), 45, 435. GAULDIE, R. D. (undated). Bilharzia in Swaziland. Internal Report Swaziland medical Department, 1-16. GILLET, J. (1950) Contribution a l'etude de la bilharziose urinaire au Congo Beige (2me partie). Annales des Societes beiges de Medicine tropicale, 30, 195-204. GILLET, J. (1956a) Bilharziasis in Portuguese Guinea. World Health Organization/Bilharzia/Conference/44 (mimeographed report). GILLET, J. (1956b) Bilharziasis in French West Africa. World Health Organization/Bilharzia/Conference/41 (mimeographed report). GILLET, J. (1960) Note sur un foyer de bilharziose murine a Schistosoma rodhaini Brumpt au Kivu. Annales de Societes beiges de Medicine tropicale, 40, 639-641. GOLDSMID, J. M. (1974) The intestinal helminthzoonoses of primates in Rhodesia. Annales de Societes beiges de Medicine tropicale. 54, 87-101. GOLDSMID, J. M., ROGERS, S., PARSONS, G. S. and CHAMBERS, P. G. (1976) The intestinal protozoa and helminths infecting Africans in the Gatooma region of Rhodesia. Central African Journal of medicine, 22, 91-95. GRABER, M. (1969) Helminths parasites de certains animeaux domestiques et sauvages du Tchad. Bulletin of Epizootic Diseases in Africa, 17, 403-428. GRABER, M. (1975) Enquete parasitoligique concervant les ranches de la Louila et de la Louboulou—• Niari—Republique populaire du Congo (Brazzaville). Ecole National Veterinaire de Lyon Lyon France. 1-39. Cited in Helminthological Abstracts (1976) 45, 497. GRETILLAT, S. (1962a) Etude du cycle evolutif du schistosome des ruminants domestique di 1'Ouest Africaine et confirmation de l'espice Schistosoma curassoni Brumpt 1931. Annales de Parasitologie humaine et comparee, 37, 556-568. GRETILLAT, S. (1962b) Une nouville zoonose, la bilharziose Ouest Africaine a Schistosoma curassoni Brumpt 1931 commune a l'homme et aux rumenants domestiques. Comptes Rendus des Frances de VAcademic des Sciences Paris, 225, 1805-1807. Cited in Helminthological Abstracts. 1963, 32, 394. GRETILLAT, S. (1969) Le phacochire {Phacochoerus aethiopicus) hote de Schistosoma sp. Bulletin de la Societe de Pathologie exotique, 62, 1090-1095. HARLEY, J. (1864) On the endemic haematuria of the Cape of Good Hope. Medico Chirurgical Transactions, 47, 55-72. HILL, D. H. and ONABAMIRO, S. D. (1960) Vesical schistosomiasis in the domestic pig. British veterinary Journal, 116, 145-150. HIRA, P. R. (1975) Observations on Schistosoma mattheei Veglia & Le Roux 1929, infections in man in Zambia. Annales des Societes beiges de Medicine tropicale, 55, 633-642. HORCHNER, F. and KARRASCH, A. W. (1972) Erstes auftreten von schistosomen bei rindern in Madagascar Berliner und Munchener Tierarztliche wochenschrift, 85, 329-330. HOLMES, P. H., MAMO, E. and LEMMA, A. (1975) Preliminary studies on metabolism in Ethiopian sheep infected with Schistosoma bovis. In Nuclear techniques in helminthological research. Vienna Austria. International atomic Energy Agency, 115-120. HSU, S. Y. and HSU, H. F. (1968) A chimpanzee naturally infected with Schistosoma mansoni. Its resistance against a challenge infection of S. japonicum. Transactions of the royal Society of tropical Medicine and Hygiene, 62, 901-902. HURTER, L. R. and POTGIETER, L. N. D. (1967) Schistosomiasis in small stock in the Potgietersrus veterinary area. Journal of the South African veterinary Association, 38, 444-446. HUSSEIN, M. F. (1973) Animal schistosomiasis in Africa. A review of S. bovis and 5. mattheei. Veterinary Bulletin, 43, 341-347. HUSSEIN, M. F., TARTOUR, G., IMBABI, S. E. and ALI, K. E. (1975) The pathology of naturally occurring bovine schistosomiasis in the Sudan. Annals of tropical Medicine and Parasitology, 69, 217-225. 246

Definitive hosts for Schistosoma species HUYGELEN, C. (1957) Le traitement d'un cas de bilharziose canine a Schistosoma rodhaini par un derive du thioxanthone. Annales des Societes beiges des Medicine tropicale, 37, 993-997. JANZ, G. J. and CARVALHO, A. M. De. (1956) Subsidios para o canhecimento dos bilharzioses em Angola. Anais Institute de Mediciiia tropical (Lisbon), 13, 597-613. KINOTI, G. (1963) S. bovis investigations in Sukumaland. East Africa Institute for medical Research. Annual Report 1962-63, 1-67. KISNER, C. D., STOFFBERG, N. and DE MEILLON, B. (1953) Human infection with Bilharzia bovis. South African medical Journal, 27, 357-358. KUNTZ, R. E. (1952) Natural infections of Egyptian gerbil with S. mansoni. Proceedings of the helminthological Society of Washington, 19, 123-124. KUNTZ, R. E. (1958) Schistosoma sp. in shrews in Lower Egypt. Proceedings of the helminthological Society of Washington, 25, 37-40. LAPIERRE, J. and HIEN, T. V. (1973) Un cas de triple infestation bilharzienne a Schistosoma mansoni, S. haematobium et Rhodobilharzia margrebowieil Annales de Parasitologie Humaine et Comparee, 48, 301-306. LARTVIERE, M., LAPIERRE, J., HOCQUET, P. and CAMERLYNCK, P. (1960) Etude d'un foyer de bilharziose a S. mansoni dans un village du cerde de Thiens (Senegal). Bulletin de la Sociiti Medicate d'Afrique Noire deLangue Francaise, 5,88-94. Cited in Helminthological abstracts (1964) 33,243-244. LAWRENCE, J. and CONDY, J. B. (1970) The developing problem of schistosomiasis in domestic stock in Rhodesia. Central African Journal of Medicine. Supplement, 16, 19-21. LAWRENCE, J. A. (1976) Schistosoma mattheei in the ox. Ph. D. Thesis of University of Rhodesia. 1-308. LENGY, J. (1962) Studies on Schistosoma bovis (Sonsino 1876) in Israel. Bulletin of the research Council in Israel, 10E, 1-36. LE ROUX, P. L. (1929) Remarks on the habits and the pathogenesis of Schistosoma mattheei together with notes on the pathological lesions observed in infected sheep. 15th Annual Report of the Director veterinary Services. Union of South Africa, October. 347-406. LE ROUX, P. L. (1930) Helminthiasis of domestic stock in the Union of South Africa. Journal of the South African veterinary Association, 1, (4), 43-65. LE ROUX, P. L. (1931a) Notes of the more important worms of cattle, sheep, and pigs in Northern Rhodesia. Annual Bulletin. Department animal Health Northern Rhodesia, 9-24. LE ROUX, P. L. (1931b) List of the helminths collected from mammals and birds in the Mazabuka area Northern Rhodesia. Annual Report. Department animal Health. Northern Rhodesia. Appendix B. 31-34. LE ROUX, P. L. (1933) A preliminary note on Bilharzia margrebowiei a new parasite of ruminants and possibly of man in Northern Rhodesia. Journal of Helminthology, 11, 57-62. LE ROUX, P. L. (1938) Annual Report central research station, Mazabuka. Annual Report veterinary Department Northern Rhodesia 1938. Appendix C. 40-60. LE ROUX, P. L. (1947) Twenty four microscope preparations and 615 camera lucida drawings—of species of schistosomes of man and animals. Laboratory meeting. Transactions of the royal Society of tropical Medicine and Hygiene, 41, 428. LE ROUX, P. L. (1955a) Game conservation and severe helminthiasis of the stock in national game reserves. Laboratory meeting. Transactions of the Royal Society of tropical Medicine and Hygiene, 49, 293. LE ROUX, P. L. (1955b) Schistosoma sp. from a goat at Kisumu, Kenya. Laboratory meeting. Transactions of the Royal Society of tropical Medicine and Hygiene, 49, 293. LE ROUX, P. L. (1955c) A new mammalian schistosomes (Schistosoma leiperi) sp. nov. from herbivores in Northern Rhodesia. Transactions of the Royal Society of tropical Medicine and Hygiene, 49,293-294. LE ROUX, P. L. (1957) Report to the Government of the Federation of Rhodesia and Nyasaland. Food and Agricultural Organization, Rome Report 696. LE ROUX, P. L. (1958a) The validity of Schistosoma capense (Harley 1864) amended as a species. Laboratory meeting. Transactions of the Royal Society of tropical Medicine and Hygiene, 52,12-14. LE ROUX, P. L. (1958b) Bone tuberculosis in the red lechwe (Adonota leche) in Northern Rhodesia. Laboratory meeting. Transactions of the Royal Society of tropical Medicine and Hygiene, 52, 15. LE ROUX, P. L. (1961) Some problems in bilharziasis in Africa and the adjoining countries. Journal of Helminthology; R. T. Leiper Supplement, 117-126. LO, C. T. and LEMMA, A. (1975) Studies on Schistosoma bovis in Ethiopia. Annals of Tropical Medicine and Parasitology, 69, 375-382. MacHATTIE, C. and CHADWICK, C. R. (1932) Schistosoma bovis and S. mattheei in Irak with notes on the development of eggs of S. haematobium pattern. Transactions of the royal Society of tropical Medicine and Hygiene, 26, 147-156. MacMAHON, J. E. (1969a) Schistosoma eggs in human faeces. Transactions of the royal Society of tropical Medicine and Hygiene, 63, 545. 247

R. J. PITCHFORD MacMAHON, J. E. (1969b) Eggs resembling those of Schistosoma intercalatum in the faeces of a baboon. Transactions of the royal Society of tropical Medicine and Hygiene, 63, 545. MAHFOUZ, A. H. (1927) A case of Schistosoma bovis in man. Journal of the Egyptian Medical Associaton, 301-306. (Cited by numerous authors). MANGO, A. M. (1971) The role of dogs as reservoirs in the transmission of Schistosoma mansoni. East African Medical Journal, 48, 298-306. MANSOUR, N. S. (1973) Schistosoma mansoni and S. haematobium found as a natural double infection in the Nile Rat, Arvicanthus n. niloticus from human endemic area in Egypt. Journal of Parasitology, 59, 424. MARILL, F. G. (1961) Diffusion de la bilharzioses chez les bovins ovins et caprins en Mauritanie et dans la vallee du Senegal. Bulletin de VAcademie Nationale de Medecine, 145, 147-150. Cited in Tropical Diseases Bulletin, 58, 1258-1259. MASSOUD, J. (1973) Parasitological and pathological observations on Schistosoma bovis Sonsino 1876 on calves, sheep and goats in Iran. Journal of Helminthology, 47, 155-164. McCULLY, R. M., VAN NIEKERK, J. W. and KRUGER, S. P. (1967) Observations on the pathology of bilharziasis and other parasitic infections of Hippopotamus amphibius Linnaeus 1758 from the Kruger National Park. Onderstepoort Journal of Veterinary Research, 34, 563-618. McKENZIE, R. L. (1970) Investigations into Schistosomiasis in sheep in Mashonaland. Central African Journal of Medicine Supplement, 16, 27-28. McQUAY, R. M. (1952) Susceptibility of a Louisiana sp. of Tropicorbis to infection with Schistosoma mansoni. Experimental Parasitology, 1, 184-188. MILLER, J. H. (1959a) Correspondence. East African medical Journal, 36, 56-57. MILLER, J. H. (1959b) The dogfaced baboon {Papio doguerd) a primate reservoir host of Schistosoma mansoni in East Africa. Journal of Parasitology, 45, No. 4, Section 2 (Supplement) 22. MILLER, J. H. (1960) Papio doguera (dog faced baboon), a primary reservoir host of Schistosoma mansoni in East Africa. Transactions of the royal Society of tropical Medicine and Hygiene, 54, 44—46. MYERS, BETTY-JUNE, WOLFGANG, R. W. and KUNTZ, R. E. (1960) Helminth parasites from vertebrates taken in the Sudan (East Africa) Canadian Journal of Zoology, 38, 833-836. NAGATI, H. F. (1942) On some parasites collected in Egypt from food animals. Journal of the Egyptian medical Association, 25, 110-111. (Cited by Hussein, 1973). NELSON, G. S. (1960) Schistosome infections as zoonoses in Africa. Transactions of the royal Society of tropical Medicine and Hygiene, 54, 301-316. NELSON, G. S. and PESTER, F. R. N. (1961) Schistosoma haematobium and S. mattheei, natural infection in a baboon from East Africa. Transactions of the royal Society of tropical Medicine and Hygiene, 55, 305. NELSON, G. S., TEESDALE, C. and HIGHTON, R. B. (1962) The role of animals as reservoirs of bilharziasis in Africa. Bilharziasis. Ciba Foundation Symposium. Churchill, London, 127-149. OKPALA, I. (1961) Studies on Schistosoma haematobium infection in school children in Epe western Nigeria. West African medical Journal, 10, 402-412. OLAFSON, P. (1955) Bovine hyperkeratosis in Israel and some problems in livestock production and animal health. Refuah Veternarith, 12, 443^146. Cited by Lengy (1962). ORTLEPP, R. J. (1961) n Oorsig van Suid-Afrikaanse helminte veral met verwysing na die wat in ons wildherkouers voorkom. Tydskrif vir natuurwetenskappe. (South African academy for Art and Science) 1, 203-212. PELLEGRINI, D. (1953) Alcuve observazione sulla schistosomiasi bovinae in Somalia e sequalazine del Bulinus abyssinicus uella regione del Basso Giura. Rivista di Parassitologia Rome, 14,15-17. Cited in Veterinary Bulletin (1954) 24, 565. PIKE, A. W. and CONDY, J. B. (1966) Fasciola tragelophi sp. nov. from the Sitatunga, Tragelophus spekei Rothschild, with a note on the pharyngeal pouch in the genus Fasciola L. Parasitology, 56,511-520. PITCHFORD, R. J. (1958a) Bilharziasis in Swaziland. Bulletin of the World Health Organization, 18,735-750. PITCHFORD, R. J. (1958b) Animal reservoirs of human bilharziasis in the Eastern Transvaal. Bulletin of the World Health Organization, 18, 1080-1081. PITCHFORD, R. J. (1959a) Natural schistosome infections in South African rodents. Correspondence. Transactions of the royal Society of tropical Medicine and Hygiene, 53, 213. PITCHFORD, R. J. (1959b) Cattle schistosomiasis in man in the Eastern Transvaal. Transactions of the royal Society of tropical Medicine and Hygiene, 53, 285-290. PITCHFORD, R. J. (1961) Observations on a possible hybrid between the two schistosomes S. haematobium and S. mattheei. Transactions of the royal Society of tropical Medicine and Hygiene, 55, 44-51. PITCHFORD, R. J. (1965) Differences in egg morphology and certain biological characteristics of some African and Middle Eastern schistosomes, genus Schistosoma with terminal spined eggs. Bulletin of the World Health Organization, 32, 105-120. 248

Definitive hosts for Schistosome species PITCHFORD, R. J. (1974) Some preliminary observations on schistosomes occurring in antelope in Central Southern Africa. Rhodesian veterinary Journal, 4, 57-61. PITCHFORD, R. J. (1976) Preliminary observations on the distribution, definitive hosts and possible relation with other schistosomes of Schistosoma margrebowiei le Roux, 1933 and Schistosoma leiperi le Roux 1955. Journal of Helminthology, 50, 111-123. PITCHFORD, R. J. and GELDENHUYS, P. J. (1960) Bilharziasis in the Transkei and Eastern Cape Province. South African Journal of laboratory and clinical Medicine, 6, 237-246. PITCHFORD, R. J. and LE ROUX, P. L. (1955a) I. Schistosomes from cattle in the Eastern Transvaal and Portuguese East Africa. Laboratory meeting. Transactions of the royal Society of tropical Medicine and Hygiene, 49, 292. PITCHFORD, R. J. and LE ROUX, P. L. (1955b) Schistosomes from the impala (Aepyceros melampus) in the Eastern Transvaal. Laboratory meeting. 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