CESTODES FROM RODENTS IN THE REPUBLIC OF SOUTH AFRICA*

Ondersiepoorl]. vet. Res. 39 (1), 255 1972 CESTODES FROM RODENTS IN THE REPUBLIC OF SOUTH AFRICA* H. MARIA COLLINS, Veterinary Research Institute, Onderstepoort ABSTRACT H. MARIA COLLINS. Cestodes from rodents in the. Onderstepoort]. vet. Res., 39 (1), 255 (1972). The cestodes recovered from various Muridae and Cricetidae in the are recorded. The following species are redescdbed: Paranoplocephala omphalodes (Hermann, 1783) Liihe, 191; lnermicapsifer congolensis Mahon, 1954; (Davaine, 187) Baer, 1956; Catenataenia compacta Ortlepp, 1962; Catenotaenia lobata Baer, 1925; Catenotaenia Iucida Ortlepp, 1962; Raillietina (Raillietina) thr_yonomy.ri Ortlepp, 1938; Raillietina (Raillietina) trapeoides (Janicki, 194) Fuhrmann, 1924; Hymenolepis diminuta (Rudolphi, 1819); Hymenolepi.r straminea (Goee, 1782); Hymenolepis nana (von Siebold, 1852). Three new species are described: Paranoplocephala otomyos from Otomys irroratus; IIymenolepis microcantha and Hymenolepis taterae from Tatera (Tatera) brantsi. Paranoplocephala acanthocirrosa Baer, 1924 is considered a synonym of P. ompha!odes; Catenotaenia capensis Ortlepp, 194 of C. lobata; and Raillietina (Raillietina) namaquensis Mettrick, 1962 of R. (R.) trapeoides. Cestodes recovered from the Myomorpha in the Ethiopian Region are summaried and a hostparasite list is included. INTRODUCTION This investigation is the first systematic survey of the cestodes of some Muridae and Cricetidae in the. Previous records of these parasites in the Ethiopian Region appear in reports on the helminth faunae of various countries. The cestodes that have been recorded from Myomorpha in the Ethiopian Region are summaried in Table 1. Some of the cestodes recovered in this survey have been recorded from many species of rodents in various parts of the world. It is neither possible nor relevant to include all these records. However, definitive hosts listed in the text only include those recorded in this survey. The following authors were consulted to verify the correct names of the host animals: Allen (1939), Cabrera (196), Davis (1968 a, b), De Graaff (1968), Ellerman & MorrisonScott (1951 ), Misonne (1968) and Trough ton (1941 ). In the text the current names of African states are used throughout; in some cases these differ from those used in the literature, as follows: Belgian Congo Rhodesia Southern Rhodesia Somalia Tanania Zambia Italian Somaliland and British Somaliland Tanganyika and Zanibar Northern Rhodesia Union of South Africa In listing the localities in the the various provinces are abbreviated as follows: E.P. Eastern Province (Cape Province) W.P. Western Province (Cape Province) N.W.P. North Western Cape (Cape Province) Ntl. Natal O.F.S. Orange Free State Tvl. Transvaal The following abbreviations are used in the figures: C Cirrus Cm Circular muscles Cp Cirrus pouch Esv Ev Glc Isv Ov Sr T V V d Vit Vs External seminal vesicle Excretory vessels Glandular cells Internal seminal vesicle Ovary Seminal receptacle Testes Vagina Vas deferens Vitellarium Vaginal sphincter PARANOPLOCEPHALA LiiHE, 191 Paranoplocephala omphalodes (Hermann, 1783) Liihe, 191 Synonym: Paranoplocephala acanthocirrosa Baer, 1924 Material (1) Paratype of P. acanthocirrosa (Onderstepoort Helminthological Collection No. T. 11, Onderstepoort) (2) Three adults and fragments of adults from. irroratus, Albany (E.P.) and Pretoria (Tvl.) (3) Fragments of adults from. unisulcatus, Uitenhage (E.P.) and Zoutpansberg (Tvl.) Redescription The total length varies from 61 to 7 4 mm and the maximum width from 5, to 7, mm. The scolex is 76 to 8 fl and the suckers 2 to 24 fl by 176 to 216 fl in diameter. The genital pores alternate irregularly and the genital ducts pass dorsal to the excretory vessels to cross from the medulla into the cortex. There are 9 to 12 testes in two to three layers; they are in a single group aporal to the female genitalia (Fig. 1). The vas deferens, which has thick walls, runs parallel to the vagina; its lumen widens in the vicinity of the seminal receptacle. The muscular cirrus pouch is elongated or pyriform; 44 to 7 fl by 12 to 18 fl wide. The cirrus is densely covered with very small spines. *Thesis submitted in partial fulfilment of the requirements for the degree of Master in Science, in the Department of Zoology, Faculty of Science, Rand Afrikaans University, November, 197 Received on 22 October 1971 Editor 25

Reference Baer (1924; 1926) Baer (1933) Mahon (1954) Mahon (1954) Baer & Fain (1955) Mettrick (1962) Baer (1924; 1926) Joyeux & Baer (193) Joyeux & Baer (193) Baer & Fain (1955) Baylis (1934) Meggitt (1921) Baer (1926) Mahon (1954) Baer (1926) Baylis (1915) Baer (1933) Baer (1926) Ortlepp (1961) Mahon (1954) Baer (1959) Joyeux & Baer (1927) Kofend (1917) Mahon (1954) Baer & Fain (1955) Mahon (1954) Baylis (1949) Joyeux & Baer (193) Ortlepp (1961) Joyeux & Baer (193) J oyeux & Baer (1927) Joyeux & Baer (1927) Baer (1926) n ttl CFl >l tj ttl CFl "rr i'" i'" tj ttl >l CFl ttl i'" ttl "() c to t: n "rr CFl c > TAB.LE 1 Summary of Cestodes recorded from Myomorpha in the Ethiopian Region Cestode Host Locality Paranoplocephala omphalodes (Hermann, 1783). Syn. Paranop!ocepha!a acanthocirrosa Baer, 1924 Otomy>' irroratus (Brants, 1827) Otomys unisulcatus Cuvier, 1829 Rhodesia N \ Paranoplocephala isomydis (Setti, 1892) Oenomys hypoxanthus (Pucheran, 1855) Paranoplocephala otomyos sp. n.. irroratus Inermicapsifer congolensis Mahon, 1954 Cricetomys gambianus Rochebrune, 1885 Dasymus incomtus (Sundevall, 1846) LemniJcomys grise fda (Thomas, 194). irroratus. unisulcatus Praomys (Mastomys) nata!ensis (A. Smith, 1834) Rattus rat/us (Linnaeus, 1758) Rhabdomys pumilio (Sparrman, 1784) lnermicapsifer aberratus Baer, 1924 Thallomys paedulcus (Sundevall, 1846) lnermicapsifer guineensis (Graham, 198) Arvicanthis niloticus (Desmarest, 1822) Syn. Inermicapsifer anibarensis Meggitt, 1921 Cricetomys bunchanani Thomas & Hinton, 1921 C. gambianus Pelomys campanae (Huet, 1888) R. rattus R. pumilio T. paedulcus lnermicapsifer madagascariensis (Davaine, 187) Aethomys (Aethomys) chrysophilus (de Winton, 1897) Syn. Inermicapsifer arvicanthidis (Kofend, 1917) Aethomys kaiseri medica/us (Wroughton, 199) Aethomys (Micaelamys) namaquensis (A. Smith, 1834) Arvicanthis aby.rsinicus (Riippell, 1842) A. niloticus C. gambiam1s Cryptomys mechowi (Peters, 1881) D. incomtus Heliophobius argenteocinereus Noack, 1894 Hybomys univittatus Peters, 1876 L. griselda Lemniscomys stria/us (Linnaeus, 1758) Mus musculus Linnaeus, 1758. hypoxanthus. irroratus Central African Republic Rhodesia Tanania Tanania Ghana Rhodesia Congo Braaville Central African Republic Dahome Sudan Kenya Central African Republic Dahome Dahome

Quentin '(1964) Joyeux & Baer (1927) Mahon (1954) Southwell & Lake (1939) Baylis (1939) Joyeux & Baer (193) Ortlepp (1961) Baylis (1934) Ortlepp (1961) Fain (195) Ortlepp (1961) Ortlepp (1961) Baer (1933) Joyeux & Baer (193) Baer (1926) Bay! is (1949) Ortlepp (1961) Fain (195) Baer (1955) Lynsdale (1953) Ortlepp (1962) Hockley (1961) Joyeux & Baer (1927) Pearse(193) [Cited by Dollfus (1953)] T his paper J oyeux & Baer (193) Mettrick (1962) Baylis (1934) Ortlepp (1962) Pearse(193) [Cited by Dollfus (1953)] ] oyeux & Baer (193) Ortlepp (1962) Quentin (1965) Baer (1959) Baer (1959) Joyeux & Baer (193) Joyeux & Baer (193) Bayl is (1939) Meggitt & Subramanian (1927) > ;:d > n t'" t: (Jl P. campanae P. (M.) nata!ensis Rattus paedu!cus (Sundevall, 1846) R. rattus R. pumi!io Saccostomys campestris Peters, 1846 Steatomys pratensis Peters, 1846 Tatera va!ida kempi Wroughton, 196 T. paedu!cus Thamnomys ruti!ans (Peters, 1876) Congo Braaville Guinea Tanania RuandaUrundi Rhodesia Central African Republic Kenya RuandaUrundi Tanania Catenotaenia baeri (Lynsdale, 1953). Syn. Meggittina baeri Lynsdale, 1953 Rat Rhodesia N 1 Catenotaenia compacta Ortlepp, 1962 Catenotaenia cricetomydis (Hockley, 1961) Syn. S krjabinotaenia cricetomydis (Hockley, 1961) C atenotaenia lobata Baer, 1925. Syn. Catenotaenia capensis Ortlepp, 194 A. (A.) chrysophi!us C. gambianuj MaJtomys sp. (with 32 chromosomes) Heliophobius marttngensis Noack, 1887 P. (M.) nata/ensis R. pumilio T. va!ida kempi Central African Republic Dahome Rhodesia Tanania Njgeria Catenotaenia Iucida Ortlepp, 1962 A. (A.) chrysophilus A. (M.) namaquensis Mastomys sp. (with 32 chromosomes) Congo Braaville Catenotaenia pauciproglottis (Quentin, 1965) Stochomys!ongicaudatus (Tullberg, 1855) Central African Republic Raillietina (Raillietina) baeri Meggitt & Subramanian, 1927. Aethomys wa!ambae pedester (Thomas, 1911) Dasymus bentleyae (Thomas, 1892) Ma!ocosmys /ongipes Rochebrune, 1885 Mastomys sp. (with 32 chromosomes) P. (M.) nata!ensis Praomys (Praomys) jack soni (de Winton, 1897) R. rat/us Dahome Central African Republic Central African Republic Congo K inshasa Dahome

Reference Baer (1959) Baer & Fain (1955) Southwell & Lake (1939) O rtlepp (1938) Mettrick (1962) Joyeux & Baer (193) Southwell & Lake (1939) Joyeux & Baer (193) Ortlepp (194) Ortlepp (194) Joyeux & Baer (193) Baer & Fain (1955) Southwell & Lake (1939) Baylis (1934) Joyeux, Baer & Martin (1936) Southwell & Lake (1939) Baer (1933) Baylis (1934) Baylis (1934) Baer (1933) Baer (1933) Southwell & Lake (1939) Southwell & Lake (1939) Baer (1925) Mahon (1954) Baylis (1934) Baer (1933) Joyeux, Baer & Martin (1936) Mahon (1954) Baylis (1939) Baylis (1934) (") tn g (/) I:I1 (/) 'I1 s:::: tl I:I1 >l (/) I:I1 I:I1 '1:1 e H (") 'I1 (/) e TABL E 1 continued Cestode Host Locality Raillietina (Raillietina) mahonae Baer & Fain, 1955 Otomys kempi Dallman, 1915 Thryonomys swinderianus (Temminck, 1827) Raillietina (Raillietina) thryonomysi Ortlepp, 1938. T. swinderianus. irroratus Raillietina (Raillietina) trapeoides (Janicki, 194) Syn. Raillietina (Raillietina) namaquensis Met trick, 1962 A. (M.) namaquensis A. niloticus Rhodesia L. striatus R. pumilio Parotomys brantsi (Smith, 184) Tatera (Tatera) brantsi (A. Smith, 1834) T. va!ida kempi T. SJvinderianus Congo Braaville Anomotaenia beimi Quentin, 1964 Lophuromys sikapusi (Temminck, 1853) Central African Republic N Dilepis do!lfusi Quentin, 1964 Hymenolepis diminuta (Rudolphi, 1819) Mastomys sp. (with 32 chromosomes) Thamnomys surdaster Thomas & Wroughton, 198 Mastomys sp. (with 32 chromosomes) M. musculus Pe!omys fa/lax (Peters, 1852) P. (M.) nata!ensis P. (P.) jacksoni R. rattus S. pratensis Tanania Central African Republic Somalia Rhodesia Tanania Toukoulo Tanania Rhodesia Central African Republic Hymenolepis globirostris Baer, 1925. Rat Hymenolepis microstoma (Dujardin, 1845) A. abyssinicus Dendromys insignis (Thomas, 193) Leggada minutoides (A. Smith, 1834) Mastomys sp. (with 32 chromosomes) M. musculus P. (M.) natalensis Tanania Rhodesia Congo Braaville Somalia T anania

Baylis (1934) Joyeux, Baer & Martin (1936) Joyeux & Baer (193) Joyeux & Baer (193) J oyeux & Baer (193) t:c 8 E (JJ P. (P.) jacksoni R. pumilio Hymenolepis microcantba sp. n. T. (T.) brantsi Hymenolepis nana (von Siebold, 1852) M. musculus Hymenolepis pearsi Joyeux & Baer, 193 H. univittatus Hymenolepis petteri Quentin, 1964 L. sikapusi Hymenolepis taterae sp. n. T. (T.) bran!ji Hymenolepis uncispinosa J oyeux & Baer, 193 H. univittatus P. (M.) natalensis Tanania Somalia Central African Republic

CESTODES FROM RODENTS IN THE REPUBLIC OF SOUTH AFRICA Cp Sr Ov T Vit FrG. 1 Sexually mature proglottid of P. ompha/ode The fanshaped ovary and the lobed vitellarium are situated porally. The vagina is thickwalled; it forms a round or oval seminal receptacle which is poral to the ovary. The uterus is a tranvserse tube with lateral evaginations. In his original description of Paranoplocephala acanthocirrosa Baer (1924) records the length of the cirrus pouch as 8 fl In 1959, Baer described the subspecies Paranoplocephala acanthocirrosa kivuensis from Otomys kempi Dollman, 1915 in the and redescribed the nominate subspecies. In this publication he states that the cirrus pouch of the latter is 366 to 686 fl long and that of P. acanthocirrosa kivuensis 215 to 267 fl The cirrus of the nominate subspecies is covered with spines while that of P. acanthocirrosa kivuensis has fine hairs. The dissimilarity in the lengths of the cirrus pouches and the fact that the cirrus is spiny in one subspecies and hairy in the other are the only differences noted between the subspecies. In this investigation, however, it was found that the cirrus pouch in the paratype of P. acanthocirrosa varies from 378 to 483 fl and in the additional material described here from 4 to 7 fl It is thus clear that the length of the cirrus pouch has a wide range of variation and cannot be considered as a reliable criterion for the erection of a subspecies. Spasskii (1951) questions the validity of P. acanthocirrosa but retains it as a distinct species as the cirrus pouch is stated to be longer (8 f.l) than that of P. omphalodes (2 to 5 f.l) However, as is shown above, this structure may vary from 215 to 8 fl in P. acanthocirrosa and it must be considered a synonym of P. omphalodes. Paranoplocephala otomyos sp. n. Definitive host: Otomys irroratus Type locality: Golden Gate (O.F.S.) Material (1) Holotype from. irroratus, Golden Gate (O.F.S.) (Onderstepoort Helminthological Collection, No. T. 162, Onderstepoort) (2) Fragments of another specimen from the same host species and locality as holotype Description The total length is 73,5 mm and the maximum width 6, mm. The. scolex is 58 fl and the spherical suckers 2 fl in diameter. The excretory system consists of one pair of longitudinal ventral and one pair of longitudinal dorsal vessels; a transverse vessel connects the ventral vessels in the posterior part of each proglottid. The genital pores alternate irregularly and are situated in the middle of the proglottid margin. The genital ducts pass dorsal to the excretory vessels to cross from the medulla into the cortex. There are 75 to 95 testes in two layers situated in a single group aporal to the female genitalia (Fig. 2). em 5p FrG. 2 Sexually mature proglottid of P. otom_yos sp. n. The cirrus pouch is oval or pyriform, 144 to 28 fl long by 12 to 16 fl wide. The cirrus is unarmed, 72 to 8 fl in diameter. Before entering the cirrus pouch, the vas deferens becomes narrow and straight and is surrounded by well developed circular muscles, 12 to 152 fl long and 4 to 8 fl wide. The thickwalled vas deferens has a wide lumen. The fanshaped ovary and the lobed vitellarium are situated porally. The walls of the vagina are thickened. The seminal receptacle is oval or round; 136 to 336 fl long by 64 to 136 fl wide. The uterus is a transverse tube with 2 to 24 saccular lateral evaginations. The ova are 44 to 4 fl and the embryo 14 fl in diameter. This species differs from all the others in this genus in the structure of the cirrus pouch (Fig. 3a, b, and c). FIG. 3a Cirrus pouch of P. ompha/odes 3

Eggs Hosts (!') 3 Arvicoia terrestris (Linnaeus, 1758) Microtus arvalis (Pallas, 1779) 46 Microtus agrestis (Linnaeus, 1758) A. terrestris Lagidium pervanum (Meyen, 1953) 47 Various rodents 4247 Cricetus cricetus Linnaeus, 1758. irroratus. irroratus. unisulcatus 5. irroratus 45. kempi 45 M. agrestis M. arvalis Microtus nivalis (Martins, 1842) No. of egg No. of eggs Hosts capsules per capsule 114 C. gambianus D. incomtus 7385 49 C. gambianus C. gambianus 5911 515 Various rodents p:: :>' > (') E {fj TABLE 2 Comparison of P. omphalodes according to various authors Synonym Author Length (mm) Width Scolex Suckers (mm) (!') (!') Cirrus pouch (!') No. of testes Length Width P. omphalodes Liihe (191) 12215 3,45, 1525 35 21 Baer (1927) 1215 5, 152 4 5 24 Spasskii (1951) 428 2,5, 72 374 5 25 Murai (197) 11 2,5 5 226 6285 P. acanthocirrosa ( paratype 1 378483 14721 Additional material 6174 5,7, 768 224 912 47 1218 \.»... P.a. acanthocirrosa Baer (1924; 1926) 456 P.a. kivuensis Baer (1959) 5 P. blanchardi Baer (1927) 24 5, 51 2 5, 585 192 2,54, 7 45 8 2 665 215367 45123 5 5 TABLE 3 Comparison of I. congolensis described by various authors Author Length Width (mm) (mm) Scolex Suckers (!') (!') Cirrus pouch (!') No. of testes Length Width Mahon (1954) 539 5,6,5 66488 2336 5318 144226 8618 Mettrick (1962) 6877 2427 8218 2224 56 112 773 253 7 14 48,593, 2,54,5 4452 162 468 64168 496

CESTODES FROM RODENTS IN THE REPUBLIC OF SOUTH AFRICA (b) (Tv!.); R.pumilio 1 Albany and Uitenhage (E.P.) and Pretoria (Tv!.) Redescription The total length varies from 48,5 to 93, mm and the maximum width from 2,5 to 4,5 mm. The scolex is 44 to 52!Land the suckers 16 to 2!L in diameter. The genital pores are unilateral ; in the specimens from L. griselda the genital pores are on the right side in more than 1 consecutive proglottids and on the left side of the remaining proglottids. The genital ducts pass between the excretory vessels to cross from the medulla into the cortex. There are 4 to 68 testes arranged in two groups, 11 to 25 poral and 27 to 45 aporal (Fig. 4). The area 1JJ Gk FIG. 4 Sexually mature proglottid of I. congolensis FrG. 3b and c Cirrus pouch of P. otomyos sp. n. b. Camera Iucida drawing c. Microphotograph It also differs from Paranoplocephala omphalodes in that the cirrus pouch is smaller and the cirrus unarmed; from Paranoplocephala isomydis in the sie of the strobila, and in the absence of an external seminal vesicle (Baer, 1949; Mahon, 1954), and from Paranoplocephala brevis Kirschenblatt, 1938 in the sie of the strobila (Zarnowski, 1955; Wahl, 1967). INERMICAPSJFER JANICKI, 191 I nermicapsifer congolensis Mahon, 1954 Definitive hosts: Cricetomys gambianus Darymus incomtus Lemniscomys griselda Otomys irroratus. unisulcatus Praomys (Mastomys) natalensis Rattus rattus Rhabdomys pumilio M aterial (1) Ten intact adults from L. griselda 1 Zululand (Ntl.); P. (M.) natalensis 1 Albany and Uitenhage (E.P.) and Pretoria (Tv!.) (2) Fragments of adults from C. gambianus 1 Zoutpansberg (Tv!.); D. incomtus 1 Lower Albany (E.P.);. irroratus 1 Pretoria and Johannesburg (Tv!.);. unisulcatus 1 Uitenhage (E.P.); R. rattus 1 Pretoria anterior to the female genitalia is usually free of testes but two or three may occur here. The cirrus pouch does not reach the excretory vessels; it is oval, 64 to 168!L long by 4 to 96!L wide. The lobed ovary is situated porally. The vagina is thickwalled and in the cortex it is surrounded by glandular cells. There are 59 to 11 egg capsules per proglottid; each contains 5 to 15 eggs. In the original description of this species Mahon (1954) states: "the vagina is seen to stain heavily, and appears to be lined with setae", but she does not mention the glandular cells which surround it. lnermicapsifer madagascariensis (Davaine, 187) Baer, 1956 Definitive hosts : Aethomys (Aethomys) chrysophilus Aethomys (Micaelamys) namaquensis Praomys (Mastomys) natalensis Rattus rattus Rhabdomys pumilio Material (1) Eighteen intact adults from A. (A.) chrysophilus 1 North Eastern Transvaal; A. (M.) namaquensis 1 P. (M.) natalensis 1 Kruger National Park, (Tv!.); R. pumilio and R. rattus 1 Pretoria (Tv!). (2) Fragments of adults from R. pumilio 1 Albany (E.P.) and Zoutpansberg (Tv!.) (3) Fragments of an adult from a child, Pretoria (Tv!.) Redescription The total length varies from 41 to 175 mm and the maximum width from 1, to 2,5 mm. The scolex is 24 to 8!Land the suckers 88 to 28!L by 136 to 28!L in diameter. The genital pores are unilateral; in one 32

H. MARIA COLLINS specimen from R. rattus they are on the right side in the anterior third of the worm and on the left in the remaining twothirds. The genital ducts pass between the excretory vessels to cross from the medulla into the cortex. There are 2 to 64 testes in two groups, 7 to 17 poral and 13 to 47 aporal in position; in some specimens there are a number of testes posterior to the female genitalia which unite the two groups (Fig. Sa, b). 1Jl Frc. 5 Sexually mature proglottid of I. madagascariensis a. Testes in two groups b. Testes confluent posterior to the female genitalia The cirrus pouch extends to the longitudinal excretory vessels and in some instances into the medulla; it is oval, 48 to 176 fl long by 24 to 56 fl wide. The fanshaped ovary is slightly offcentre and the lobed vitellarium somewhat aporal to it. A small seminal receptacle is present. There are 16 to 99 egg capsules per proglottid; they contain 5 to 13 eggs. Fain (195) showed that /. cubensis is a synonym of I. arvicanthidis. Baer (1956) reviewed the literature dealing with this taxon and concludes that I. madagascariensis has priority over other names (Table 4). CATENOT AEN!A JANICKI, 194 The status of the genus Catenotaenia as well as that of its species has been in dispute for many years. At various times it has been placed in the families Taeniidae, Dilepididae, Anoplocephalidae and finally in the Catenotaeniidae. According to Spasskii (1951), Akhumyan (1946) divided it into three genera, Catenotaenia, Mathevotaenia and Skrjabinotaenia, using the distribution of the testes and structure of the uterus as criteria. Spasskii (1951) separates Catenotaenia from Skrjabinotaenia on testes distribution and places Catenotaenia oranensis Joyeux & Foley, 193 and C. lobata in Skrjabinotaenia, which he defines as having the testes in two groups. In the present study, however, it was found that the testes of C. lobata are not always in two dis tinct groups; furthermore the uterus is like that of the other species but unlike that of C. oranensis. It is, therefore, clear that the criteria used for the erection of these genera and subgenera are invalid. Tenora (1959) recognied Catenotaenia and Skrjabinotaenia; in the former the main trunk of the uterus reaches the end of the. proglottid; in the latter the uterus has a short trunk with branches arising only at the base and leading laterally and posteriorly. Tenora further creates three subgenera, i.e. Catenotaenia, Spaskjella and Meggittina, using the distribution of the testes as a criterion. However, Ortlepp (1962) and Wolfgang (1956) are of the opinion that the distribution of the testes is not a valid criterion for the creation of subgenera. The data presented below also show that this character is variable, e.g. in Catenotaenia lobata Baer, 1925 it is dependent on the state of contraction of the worm. Ortlepp (1962) states "a subdivision of the genus into subgenera, if necessary, would be more valid when based on the branched or unbranched nature of the excretory system and not on the spatial arrangement of the testes". If this suggestion is followed the two subgenera would be: (1) Species in which the excretory system consists of one pair of ventral and one pair of dorsal longitudinal vessels. This would include Catenotaenia pusilla (Goee, 1782) Janicki, 194; Catenotaenia ae!!)'ptiaca Wolfgang, 1956; Catenotaenia california Dowell, 1953; Catenotaenia chabaudi Dollfus, 1953; Catenotaenia cricetorum (Kirschenblatt, 1949); Catenotaenia dendritica (Goee, 1782); Catenotaenia geosciuri Ortlepp, 1938; Catenotaenia linsdalei Mcintosh, 1941 ; Catenotaenia reggiae Rausch, 1951; Catenotaenia rhombomydis Schul & Landa, 1935 and C. oranensis. (2) Species in which the excretory system consists of more than one pair of ventral and more than one pair of dorsal longitudinal vessels. These are: Catenotaenia compacta Ortlepp, 1962; Catenotaenia gerbilli (Wertheim, 1954); Catenotaenia Iucida Ortlepp, 1962; C. lobata and Catenotaenia pauciproglottis (Quentin, 1965). The excretory system of Catenotaenia baeri (Lynsdale, 1953), however, is unknown. To avoid further confusion it is considered inadvisable to subdivide the genus until a comparative study of all the species has been made. Catenotaenia compacta Ortlepp, 1962. Definitive hosts: Aethomys (Aethomys) chrysophilus Aethomys (Micaelamys) namaquensis Material (1) Syntypes from A. (A.) chrysophilus, Kaapmuiden (Tv!.) (Onderstepoort Helminthological Collection, No. T. 147, Onderstepoort). (2) Twentytwo adults from A. (M). namaquensis, Kruger National Park, (Tv!.). Redescription The strobila consists of three to six proglottids of which at least one is immature, one mature and one gravid. The total length varies from 2,5 to 11,5 mm and the maximum width from, 7 to 2,5 mm. The scolex is 392 to 92 fl and the suckers 168 to 264 J.L in diameter. The excretory system consists of several longitudinal ventral and dorsal vessels which anastomose. The genital pores alternate regularly. 33

E ggs 55x49 22x 19 6 34 1216 6 36 11 22x 19 6 2326 Uterine Embryo branches (!') 122 1518 112 1518 1216 912 119 Hosts Various rodents Various rodents T. paedu!cus. irroraltts Various rodents Various rodents H. argentiocinereus P. (M.) natalensis Various rodents Rat Hosts A. (A.) chrysophilus A. (A.) chrysophilus A. (M.) namaquensis (") m [fj >l lj m [fj "rl :;o :;o lj m >l [fj >l :r: m :;o m "tl g t:: (") "rl [fj c >l :r: > "rl TABLE 4 Comparison of I. madagascariensis described by various authors Length Width Scolex Suckers No. of Synonym Author (mm) (mm) (!') (!') testes Cirrus pouch (!') Length IWidth I. madagascariensis Kouri & Rappaport (194) 2742 1,2, 6163 15185 4934 15 75 Baer (1956) 14519 2,6 595 183 4553 91114 257 Ortlcpp (1961) 75 19 34 12 6 No. of No. of capsules eggs per capsule 48175 611 1315 61 12 51 Sie Embryo Hooks of eggs (!') (!') (!') 41175 1,2,5 248 8828 264 48176 2456 141 1,5 3452 7296 4864 I. arvicanthidis Baer (1926) 43 34 Baer (1927) 71 2, 5 5 35 1415 Meggitt & Subramanian (1927) 71 2, 5 35 1415 Baylis (1949) 17925 3, 45 19 4855 112 6 1619 513 591 68 14 812 115 1113 15 1113 8125 1215 Mahon (1954) 2132 1,6,3 376576 14224 287 918 4761 I. cubensis Baer et a/. (1949) 5315 1,52,3 3256 1519 2832 92126 3665 Baylis (1949) 2742 2,1 61 185 3349 15 75 Fain (195) 17 1,8 44 18 4855 92138 558 R. (R.) loechesalavei Dollfus (194) 191 35 148 335 16136 4855 912 12618 41 45175 611 9512 41 758 36 TABLE 5 Comparison of C. compacta by various authors No. of Cirrus pouch (!') Authors prog Length Width Scolex Suckers No. of lottids (mm) (mm) (!') (!') testes Length Width Ortlepp (1962) 613 21 1,8 23 118 273 68 Type specimens 56 911,5 2,2,5 729 22264 118 12336 48 Other material 35 2,56,,71,3 39292 168192 82134 1234 3272 Seminal Vaginal receptacle Vesicle (!') (!') 12 336 1828 136 24 964 16 8296 68144 22184 5414 45 136

H. MARIA COLLINS The genital ducts pass between the excretory vessels when crossing from the medulla into the cortex. T here are 82 to 18 testes in two to three layers which surround the female genitalia completely (Fig. 6). (2) Fragments from P. (M.) natalensis, Albany (E.P.), Brits (Tv!.); R. pumilio, Cradock (E.P.) Redescription The total length is 25, mm and the maximum width 1, to 2, mm. The scolex is 24 to 4 p. and the spherical suckers 14 to 12 p. in diameter. T h e excretory system consists of several longitudinal ventral and dorsal vessels which anastomose. The genital pores alternate irregularly and are situated near the anterior margin of the proglottid. The genital ducts pass between the excretory vessels to cross from the medulla into the cortex. There are 1 to 18 testes which are mainly lateral to the female genitalia but some testes unite the two groups posterior to the ovary (Fig. 7). T hey are 14 to,, 21' FrG. 6 Sexually mature proglottid of C. compacta T he cirrus pouch is oval, 12 to 336 p.long by 32 to 8 p. wide. T he strongly lobed ovary is situated in the centre of the proglottid while the lobed vitellarium is situated porally. T he seminal receptacle is round or oval, 68 to 24 p. by 54 to 16 p. in diameter. The vagina, which is thickwalled, loops dorsoventrally as well as anteroposteriorly and dilates to form a large vesicle before opening in the genital atrium; this vesicle is 22 to 4 p. by 45 to 296 p. in diameter. The uterus consists of a longitudinal stem with 1 to 19 lateral branches which redivide. This species resembles C. lobata, C. Iucida and C. pauciproglottis in that the longitudinal excretory system forms a branching network. C. compacta is the only species in which the genital pores are always regularly alternating ; this may also occur in C. aegyptiaca but is not constant in all the type specimens. Rhab Catenotaenia lobata Baer, 1925 Synonym : Catenotaenia capensis Ortlepp, 194 Definitive hosts: Praomys (Mastomys) natalensis domys pumilio Material (1) Holotype of C. capensis from R. pumilio, Stilbaaistrand (E.P.). (Onderstepoort Helminthological Collection, No. T. 146, Onderstepoort) FrG. 7 2JJ Sexually mature proglottid of C. lobata 128 by 88 to 14 p. in diameter. The vas deferens forms a few coils before entering the cirrus p ouch which is pyriform, 136 to 28 p. long by 4 to 96 p. wide; it extends to the ventral excretory vessels or slightly beyond it to the median wall of the most laterally situated ventral vessel. The ova.ry has long slender lobes and is situated on the midline of the proglottid approximately one third of the length of the proglottid from the anterior margin. The lobed vitellarium is situated p orally. The 35

n ti1 (/J Cl t:l ti1 (JJ '"rl ::::: t:l ti1 '1 (/J ti1 ti1 " c tj:i t: n '"rl (JJ c '1 :I: > '"rl H n > TABLE 6 Comparison of C. lobata described by various authors V>. Synonym Author Length (rnm) C. lobata Baer (1925) 28 Meggitt & Subramanian (1927) 144 Joyeux & Baer (1945) 1214 Mahon (1954) Mettrick (1962) 6 11 Wahl (1967) 158 25, C. capensis Ortlcpp (194) 23 5. lobata Akhumyan (1946) [cited by Spasskii (1951)] 57 Cirrus pouch (ft) Width Scolex Suckers 1\:o. of Uterine Hosts (mm) ([L) ([L) testes Length Width branches,71,5 7 2 2 15 6 Rat 2,5 7 2 15 6 2 Apodemus sylvatim.r (Linnaeus, 1758) H. marunpensis 1,2, 37 122 2 15 17 6 115 Various rodents 2 1442 34 P. (M.) nata!ensis 25 29 9 1 15 17 16 4 11 17 P. (M.) nata!ensis 2,8 35 6 11 2 25 17 2 1<1 astomys sp. r 2,5 432 133 2 16 165 4555 115 Apodemus spp. 1,2, 244 1412 118 136 28 496 142 R. pumilio P. (M.) natalen.ris 2, 41 18 225 15 5 113 R. pumilio 2,5 316 122155 2229 13916 19 14 Various rodents TABLE 7 Comparison of C. Iucida described by various authors No. of Author prog Length lottids (mm) Cirrus pouch ([L) Seminal receptacle ([L) Width Scolex Suckers No. of Uterine Hosts (mm) ([L) ([L) testes Length Width Length Width branches Ortlepp (1962). 18 45 26 17 Type specimens 1316 16,537,5 Other material 37 52 1,3 1214 9 2 27 3 57 1216 1922 A. (A.) chrysophilus 11 33 12 15 175 45 14 15 Ma.rtomys sp. 1,21,5 56 16184 117 144 152 336 568 96168 64 144 15 21 A. (A.) chrysopbilus 152 16 1, 88 12 12 132139 rc 12168 448 64 128 4812 17 A. (A.) chrysophilus A. (M.) namaquensis

H. MARIA COLLINS vagina forms one or two coils before it opens into the round or oval seminal receptacle. The uterus has 14 to 2 branches. The type specimens of C. capensis are severely contracted and stain badly but it could be verified that the testes are not confluent along the posterior margin. Ortlepp (194) used the latter character as a criterion for differentiating this species from C. lobata but this is probably caused by the severe contraction of the material. Moreover, the testes distribution in the material described above forms a transition between those described by Baer (1925) and by Ortlepp (194). describes C. lobata with the testes in two groups lateral to the ovary joined by two to five rows of testes behind the female genitalia. According to Spasskii (1951) Akhumyan (1946) considers these two species to be identical but places them in a new genus, Skrjabinotaenia, which is differentiated from Catenataenia on the testes distribution. As this criterion is, however, influenced by the degree of contraction it cannot be used for the erection of a genus. In C. lobata the two groups of testes are united by a few testes along the posterior margin of the proglottid. In the other species the testes are mainly posterior to the female genitalia. Catenotaenia Iucida Ortlepp, 1962 Definitive hosts: Aethomys (Aethomys) chrysophilus Aethomys (Micaelamys) namaquensis Material (1) Syntypes from A. (A.) chrysophilus, Kaapmuiden (Tv!.). (Onderstepoort Helminthological Collection, No. T. 148, Onderstepoort) (2) Fragments from A. (A.) chrysophilus and A. (M.) namaquensis, unknown localities Redescription The strobila consists of 13 to 37 proglottids. The total length is 16,5 to 52, mm; the maximum width 1, to 1,5 mm. The scolex is 5 to 88 fl and the suckers 12 to 184 by 12 to 16 fl in diameter. The excretory system consists of several ventral and dorsal vessels which anastomose. The genital pores alternate irregularly. The genital ducts pass between the excretory vessels to cross from the medulla into the cortex. There are 117 to 144 testes; 66 to 7 fl by 63 to 96 fl in diameter. They are situated in a single layer laterally and posteriorly to the female genitalia (Fig. 8). The cirrus pouch is elongate, 12 to 336 fl long by 4 to 8 fl wide. The ovary consists of long lobes and is situated in the anterior part of the proglottid. The vitellarium is lobed and poral in position. The seminal receptacle is 64 to 168 fl by 48 to 144 fl in diameter. The uterus has 15 to 21lateral branches which may redivide. In C. Iucida the majority of the testes are posterior to the female genitalia with a few lateral; in C. lobata they are mainly lateral with only a few posterior; in C. compacta they surround the female genitalia while in C. pusilla all of them are posterior. FrG. 8 Sexually mature proglottid of C. Iucida 37

Cirrus pouch (!') Length Width 12 66 Capsules Eggs 45 45 45 4 5 12 6 8 59 6 5 8 112 58 14 87 6 12 45 48 67 71 Hosts T. SJvinderianus T. s1vinderianus. irroratus Hosts A. niloticus A. niloticus Mus spp. T. s1vinderianus R. pumilio T. (T.) brantsi L. striatus T. (T.) brantsi P. brantsi A. niloticus A. (M.) namaquensis n tr1 (/).., tl tr1 (/) :: tl tr1.., (/) i tr1 :: tr1 c "' IJj t; n '"r1 (/) c i > '"r1 TABLE 8 Comparison of R. (R.) thryonomysi described by various authors Synonym Author Length (mm) Width (mm) Rostellar hooks Scolex Suckers No. of (!') (!') No. Sie (!') testes R. (R.) thryonomysi Ortlepp (1938) 9,516,,5,6 4248 115 12 1 36 4 56 9,516,,5 1 16 35,8 44,5 332 TABLE 9 Comparison of R. (R.) trapeoides described by various authors (..» Synonym Author Length Width (mm) (mm) Scolex (!') Hooks Cirrus pouch (!') Suckers No. of (!') No. Sie (!') testes Length Width R. (R.) trapeoides Von Janicki (194) 4 1, 182 16 8 1215 Meggitt & Subramanian (1927) 4 1, 67182 16 8 12 15 15132 5275 Southwell & Lake (1939) 5 6 1,2 14 9 15 Ortlepp (194) 2 22 79 1 15 12 162 684 16 6, 7,5 11 14 12 6 28 4 5 17 19 7,59, 1 12 12 17 75 15 15,52, 264 86 x 4 1625 7,811,1 8 16 6416 48 R. (R.) kordofanensis Meggitt & Subramanian (1927) 4 1 1, 6,7 1 12 14 R. (R.) namaquensis Mettrick (1962) 11 1,3 18 24 6 7 19 22 6, 8, 8 11 14 15 7

H. MARIA COLLINS RAILLIETINA FUHRMANN, 192 Raillietina (Raillietina) thryonomysi Ortlepp, 1938 Synonym: Raillietina (S.L.) thryonomysi Ortlepp, 1938 Definitive hosts: Thryonomys swinderianus Otomys irroratus Material (1) Holotype and para types from T. swinderianus, Zululand (Ntl.). (Onderstepoort Helminthological Collection, No. T. 155, Onderstepoort) (2) Fragments of adults from. irroratus, Albany (E.P.) Redescription The total length of the type specimens varies from 9,5 to 16, mm, the maximum width is,5 mm. The rostellum is armed with 1 to 16 hooks 35,8 to 44,5 fl long (Fig. 9). The genital pores are unilateral and situated in the anterior part of the proglottid margin. capsule in the material from. irroratus this species is placed in the nominate subgenus Raillieina. Raillietia (Raillietina) mahonae Baer & Fain, 1955 drffers from this species in having more rostellar hooks (11 to 13 vs. 1 to 16) which are slightly smaller (26 to 35 fl vs. 36 to 45 fl) (Table 8). These two species may be identical but they are kept separate here until more material is available. Baer & Fain (1955) think that the specimens from the which Southwell & Lake (1939) assigned to Raillietina (Raillietina) gracilis (Janicki, 194) are identical with R. (R.) mahonae, which they differentiate from the latter on the shape and sie of the cirrus pouch. Raillietina (Raillietina) trapeoides (Janicki, 194) Fuhrmann, 1924 Definitive hosts: Tatera (Tatera) brantsi Parotomys brantsi Material (1) Fragments of adults from T. (T.) brantsi, J ohannesburg, Roodepoort, Luipaardsvlei, Randfontein (Tv!.) (2) Four adults and fragments from P. brantsi, Port Nolloth (N.W.P.) Redescription The total length varies from 15 to 15 mm and the maximum width from,5 to 2, mm. The scolex is 264 fl in diameter; the rostellum bears 16 to 25 hooks, 7,8 to 11,1 fl long (Fig. 1). The suckers are 86 1y FrG. 9 Rostellar hooks of R. (R). thryonomysi As the specimens are macerated very little of the internal morphology could be determined. There are from 3 to 32 testes, 8 to 11 poral and 21 to 22 aporal to the female genitalia. The vagina is surrounded by a small sphincter. There are several egg capsules per proglottid; these contain four to seven eggs each. The fragments from. irroratus have rostellar hooks of the same sie and shape as those of the holotype. The latter is immature and Ortlepp (1938) could not place it in a subgenus. As there is more than one egg per Frc. 1 Rostellar hooks of R. (R). trapeoides by 4 fl in diameter and are armed with several rows of spines. The genital pores are unilateral; the genital ducts pass between the excretory vessels to cross from the medulla into the cortex. FrG. 11,.._..., 1)J Sexually mature proglottid of R. (R). trapeoides 39

CESTODES FROM RODENTS IN THE REPUBLIC OF SOUTH AFRICA I (b) s\ij! llj FrG. 12 >OOp Sexually mature proglottid of H. diminuta /@ @lis!![ sij (a) [ e j FrG. 13a and b Sexually mature proglottids of H. diminuta showing variations in the number and arrangement of the testes in two individuals 1y FrG. 14 Rostellar hooks of H. nana.cj f FrG. 15 1)' Sexually mature proglottid of H. nana 4

H. MARIA COLLINS There are from 8 to 16 testes; they are in two groups which are united by 2 to 3 testes posterior to the female genitalia (Fig. 11). The oval cirrus pouch extends to the longitudinal excretory vessels; it is 64 to 16 P long by 4 to 8 fj wide. An internal seminal vesicle is present. The lobed ovary and the vitellarium are situated in the centre of the proglottid. The seminal receptacle is small. The vagina is surrounded by a well developed sphincter muscle 2 fj from its opening into the genital pore; it is 2 fj in diameter, and 3 Plong. There are 14 to 87 egg capsules per proglottid; each contains 6 to 12 eggs. It differs from the other two species paraslt1mg rodents in that its rostellar hooks are smaller (7 to 11 JJ) than those of either R. (R.) gracilis which according to Southwell & Lake (1939) are 4 fk, R. (R.) thryonomysi Ortlepp, 1938 which are 36 to 45 fj long (Table 9). Mettrick (1962) differentiated R. (R.) namaquensis from R. (R.) trapeoides on the basis of a slightly larger cirrus pouch (14 to 15 fj by 7 JJ) and fewer testes (8 to 11). Mettrick's material, however, falls within the range of variation described above and must, therefore, be considered a synonym of R. (R.) trapeoides. HYMENOLEPIS WEINLAND, 1858 The genus Hymenolepis comprises more than 3 species parasitic in birds and mammals (Hughes, 1941 ). Recently Spasskii (1954) and Yamaguti (1959) have each described four new genera and Zarnowski (1955) one to accommodate the species which parasitie rodents. The genera lfymenolepis and Insectivorolepis Zarnowski, 1955 contain those species which have an unarmed rostellum, while the other eight genera are differentiated from one another on the number, sie and shape of the rostellar hooks, the distribution of the testes and the development of the uterus. However, these differences are not always clear and it is sometimes difficult to distinguish between the genera. Hymenolepis and Insectivorolepis, which are both unarmed, must be differentiated on the uterus only: in the former it is saccular and lobed but in the latter an ovoid median sac. While a single difference may be a valid criterion for specific differentiation it is insufficient for the erection of a genus. In the present paper all these genera are, therefore, regarded as synonyms of Hymenolepis. Hymenolepis diminuta (Rudolphi, 1819) Definitive hosts: Mus musculus Praomys (Mastomys) natalensis Rattus rattus Material (1) Four adults from R. rattus, Pretoria (Tvl.) (2) Fragments from M. musculus, Pretoria (Tvl.); P. (M.) natalensis, Johannesburg (Tvl.); R. rattus, Albany (E.P.), Wynberg (W.P.), Durban (Ntl.), Ficksburg (O.F.S.) Redescription The total length varies from 15 to 17 mm and the maximum width from,5 to 4, mm. The scolex is 168 to 336 fj and the suckers 8 to 12 fj in diameter. The rostellum is rudimentary and unarmed. The genital pores are situated in the middle of the proglottid margin; they are usually unilateral. The genital ducts pass dorsal to the longitudinal excretory vessels when crossing from the medulla into the cortex. There are from one to five, usually three, testes [type III of Skrjabin & Matiewossian (1945) cited by Zarnowski (1955)]; they are 88 to 184 fj by 72 to 152 fj in diameter (Fig. 12). The cirrus pouch, which is elongated, extends into the medulla; it is 12 to 44 fj long by 32 to 72 fj wide. A large external seminal vesicle is present. The lobed ovary and the vitellarium are situated in the centre of the proglottid. The seminal receptacle lies anterior to the ovary. The outer membrane of the ovum is 64 to 72 fj by 64 to 66 fj; the inner membrane 3 to 34 fj by 28 to 3 fj; the embryo 26 to 3 fj by 22 to 26 fj with embryonic hooks 16,4 to 19,4 fj long. H. diminuta shows greater variation in the number and the distribution of the testes than the other species of the genus. They vary in number from one to five per proglottid and their spatial arrangement shows marked variation even in a single individual (Fig. 13a, b). The shape of the cirrus pouch is also variable: in the early sexually mature proglottid it is elongated but in some fully mature and gravid proglottids it is pyriform. Hymenolepis nana (von Siebold, 1852) Material Three intact adults from R. pumilio, Pretoria (Tvl.) Redescription The total length varies from 15 to 3 mm; the maximum width is,5 mm. Thescolex is 136 to 176 fj and the suckers 54 to 73 fj in diameter. The rostellum bears a single crown of 24 to 28 hooks, 16,3 to 2,6 fj long (Fig. 14). The genital pores are unilaterally situated in the anterior half of the proglottid margin. The genital ducts pass dorsal to the excretory vessels to cross from the medulla into the cortex. There are two to three testes arranged in a straight line, [type VII of Skrjabin & Matiewossian (1945) cited by Zarnowski (1955)] (Fig. 15). The cirrus pouch is small and pyriform, 52 to 14 fj long by 16 to 32 fj wide. It usually reaches the excretory vessels but in some proglottids these vessels are deflected medially or laterally so that the cirrus pouch may not extend to them or may extend into the medulla. The lobed ovary is in the centre of the proglottid; the seminal receptacle is anterior to it. The ova have polar filaments; the outer membrane is 46 to 62 fj by 36 to 46 fj and the inner membrane 24 to 3 fj by 18 to 28 fj in diameter; the embryo is 18 to 24 fj by 18 to 2 P with embryonic hooks 11,5 to 14,4 JJlong. This worm resembles Hymenolepis straminea (Goee, 1782) in the sie of the rostellar hooks but may be differentiated from it in the shape of these hooks and in the sie of the cirrus pouch. (Table 11 ). 41

Embryonic hooks 14 16 11 11 16,419,4 Hosts R attus nonvegicus (Berkenhout, 1769) other rodents, man Various rodents Rats and mice Cercopithecus schmidti (Linnaeus, 1766) Various rodents Various rodents (Murinae) Rats, mice and man C. schmidti Rodents R. nonvegicus Various rodents n ttl [f) >l tl ttl [f) '"I1 tl >l [f) i ttl ttl '" c tl:l t: n '"I1 [f) c >l ::c > &2 TABLE 1 Comparison of H. diminuta described by various authors Length Width Scolex Suckers Author (mm) (mm) (p.) (p.) Cirrus pouch (p.) Length Width O va (p.) Outer Inner membrane membrane Embryo Liihe (191). 2 6 2,54, 25 5 8 16 6 8 Sluiter et a/. (1921). 16 3,5 Brumpt (1927). 1216 3,5 7 86 36 x 28 6 86 36 x 18 i!3 Meggitt (1927). 29 43 174 26 Meggitt & Subramanian (1927) 12 16 4, 255 174 2 6 Hegner et a/. (1929) 1216 3,5 Joyeux & Foley (193) 12 16 34 174 26 2 28 Tubangui (1931) 1 16 2,54, 26 816 17 3 24 15 18,54, 168336 812 1244 3272 5486 5486 24 4 36 67 86 244 54 86 2 35 6472 3 34 263 64 66 283 22 26

24 2 2 4 4 3 39 4 65 Eggs (f.l) Inner On comembrane sphere 2 34 18 1634 2435 21 27 2425 21 27 18 24 1619 232 243 18 24 18 28 18 26 29 31 28,5 35,3 24 35 22,528,5 21 27 4,5 51 3645 36 43,5 31,54,5 23 5 29 3 Hooks 114 114 9 14 11,5 14,4 1216 1,515 15 15 16 Hosts, rats, mice, rats, mice, various rodents R. nonvegicus, rats, mice, various rodents R. pumilio Various rodents R. rat/us R. norwegicus Various rodents R. nonvegicus R. pumilio E fiomys quercinus (Linnaeus, 1766) ;:r: :;d > 8 r< t: [jj TABLE 11 Comparison of H. nana according to various authors it Rostellar H ooks Length Width Scolex Suckers r Synonym Author (mm) (mm) (f.l) (f.l) No. Sie (f.l) H. nana Brumpt (1927) 1 25,55,7 32 8 243 14 18 Hegner et a!. (1929) 1 45,5,7 25 3 14 18 Hughes (1941) 7 8,27,65 13 4 593 2 26 16 2 Meggitt & Subramanian (1927) 5 4,5,9 21 48 r 3 1418 Sluiter et a!. (1921) 6,9 2535 1 13 2428 15 18 Tubangui (1931) 5 9,2,9 13 48 7 15 2 3 14 18 Wardle & McLeod (1951) 5 6,5 1, 3 4 27 1618 Liihe (191) 4 45,5,9 25 32 79 23 14 18 15 3,5 136 176 54 72 2428 16,3 2,E H.jraterna Baer & Tenora (197) 28 59,6 1, 2633 9 12 2 3 14 18 H. longior Baylis (1922) 45,42,53 21 26 75 93 21 22 19 2 H. nanajraterna Baylis (1922) 2,27,51 16 23 57 22 26 1618 Taenia nana Leuckart (1886) 12 2,5 3 1 2228 18 Taenia murina Dujardin (1845) 25,55 32 8 224 1517 Cirrus pouch (f.l) Outer Length Width membrane 5 6 486 36 48 6 36 44 44 52 6572 18 21 3 6 553 5 7 2 25 3741 36 56 3242 46 62 52 14 16 32 36 46 673 58 49 6 4248 67,5 9 66 87

CESTODES FROM RODENTS IN THE REPUBLIC OF SOUTH AFRICA a 1)1 FrG. 17 Sexually mature proglottid of H. microcantha sp. n. a. Whole mount b. Transverse section 1).1 b' c ( J 1...,.,... I I FrG. 16 Rostellar hooks of H. microcantha sp. n. a. Camera Iucida drawing b. and c. Microphotograph 5 FrG. 18 Rostellar hooks of H. straminea 44

H. MARIA COLLINS As pointed out by Baer & Tenora (197), there is a difference of opinion among workers regarding the specific distinctness of H. nana of man and Hymenolepis fraterna (Stiles, 196) of rodents. In their publication it is stated that Baer considers both species to be valid while Tenora considers them identical. Hymenolepis microcantha sp. n. Definitive host: Tatera (Tatera) brantsi Intermediate host: Unknown Type Locality: Johannesburg (Tvl.) Distribution: Material Holotype from the small intestine of T. (T.) brantsi, Johannesburg (Tvl). (Onderstepoort Helminthological Collection, No. T. 163, Onderstepoort) Description The total length including the scolex is 57, mm and the maximum width is 1, mm. The strobila consists of about 6 proglottids which are all much broader than long; thus the terminal proglottid is 1, mm broad but only 96 fl long. The scolex is 248 fl and the suckers 12 fl in diameter. The rostellum bears a crown of 87 hooks, 8,2 to 11,5 fl long (Fig. 16a, b, c). The ventral excretory vessels, 16 fl in diameter, are united by a transverse vessel in the posterior part of each proglottid; the dorsal excretory vessels are 2 fl in diameter. The genital pores are unilateral and situated in the middle of the proglottid margin. The genital ducts pass dorsal to the excretory vessels when crossing from the medulla into the cortex. The transverse muscles are weakly developed. The three testes are in a straight line, two aporal and one poral [type VII of Skrjabin and Matiewossian (1945) cited by Zarnowski (1955)]; they are oval in shape, 47 to 9 fl by 31 to 67 fl in diameter (Fig. 17a, b). The vas deferens is short and forms a large external seminal vesicle which may partially overlay the cirrus pouch. The latter is long and narrow and extends well into the medulla; in mature proglottids it is 129 to 162 fl long and 24 fl wide; in gravid proglottids it is 16 to 224 fl by 24 fl The ovary is slightly lobed and tranversely elongated. In whole mounts both the ovary and the vitellarium are overlaid by the seminal receptacle and the testes. The pyriform seminal receptacle is situated in the middle of the proglottid anterior to the ovary. On leaving the seminal receptacle the vagina crosses the cirrus pouch and runs anteriorly to it; in the cortex it again crosses the cirrus pouch and opens posteriorly to it in the genital atrium. The lumen of the vagina is wide, about 16 fl in diameter throughout most of its length, but on recrossing the cirrus pouch it narrows gradually to 4 fl at its opening in the genital atrium. There are 24 to 32 oval ova per gravid proglottid; the outer membrane is 67 to 9 fl by 45 to 74 fl and the inner membrane 31 to 38 fl by 29 to 34 fl in diameter; the embryo is 27 to 34 fl by 27 to 31 fl with embryonic hooks 13 to 18 fl long. This species resembles Hymenolepis petteri Quentin, 1964 in the sie of both the rostellar hooks and the cirrus pouch. It differs from the latter, however, in that it has fewer rostellar hooks (87 vs. 128), the cirrus is unarmed, there is no sphincter in the cirrus pouch and the outer membrane of the ovum is not covered in protrusions. H. microcantha is therefore considered distinct from H. petteri. H. microcantha is readily differentiated from the other armed hymenolepids occurring in rodents. According to Hughes (1941), Hymenolepis crassa Janicki, 194 has rostellar hooks 13 fl in length but it has only 24. Mahon (1954) records the length of the rostellar hooks of H. straminea as being 1 to 15 fl, but this species also has only 18 to 3 hooks. The rostellar hooks of H. microcantha are slightly larger (8,2 to 11,5 tl) than those recorded by Baer & Della Santa (196) for Hymenolepis prolifer (Villot, 188) Stammer, 1955, (5,5 to 6,4 tl) However, the latter has 16 to 18 hooks as well as a smaller cirrus pouch and it has been recorded only from Sorex spp. Hymenolepis straminea (Goee, 1782). Definitive hosts: Leggada mimttoides Mus nmsettlus Praom_)'S (Mastomys) natalensis Rhabdomys pumilio Material (1) Three intact adults from the liver of R. pumilio, Pretoria (Tvl.) and five intact adults from the small intestine of P. (M.) natalensis, Brits (Tvl.) (2) Fragments from the small intestine of P. (M.) natalensis and M. musculus, Pretoria (Tvl.); L. mintttoides, Albany (E.P.) The total length varies from 3 to 145 mm and the width from,17 to 2, mm. The scolex is 216 to 336 fl and the suckers 72 to 14 fl in diameter. The rostellum bears a single crown of 22 to 3 hooks, 12,1 to 18,6 fl long (Fig. 18). The genital pores are unilateral, situated in the anterior half of the proglottid margin. The genital ducts pass dorsal to the longitudinal excretory vessels when crossing from the medulla into the cortex. There are two to three testes either in a straight line [type VII of Skrjabin & Matiewossian (1945) cited by Zarnowski (1955)] or in a triangle (type III of Skrjabin & Matiewossian (1945) cited by Zarnowski (1955)]; they are 41 to 12 fl by 22 to 96 fl in diameter (Fig. 19). The cirrus pouch is pyriform, extends to the longitudinal FrG. 19 1)J Sexually mature proglottid of H. straminea 45

E mbryonic hooks 15 19 15 18 18 172 17 2 13,72,2 15,1 19,2 H osts Various rodents C. cricetus M. musculus R. rattus R. nonvegicus M. musculus R. nonvegicus R. rattus Rats and white mice Meriones shawi (Duvernay, 1842) White mice M. musculus M. shmvi P. (M.) natalensis M. musculus M esocricetus aura/us (Waterhouse, 1839) Apodemus spp. R. pumilio M. musculus L. minutoides P. (M.) natalensis n tr1 (fj tr1 (fj >yj : s::: : tl tr1..., (fj..., ::r: tr1 : tr1 "' c IJj 1:"' n >yj (fj c i > >yj &; TABLE 12 Comparison of H. straminea described by various authors Rostellar hooks Cirrus pouch (p.) O va (p.) Synonyms Author Length Width (mm) (mm) Scolex (p.) Suckers (p.) N o. Sie (p.) Length Width Outer Inner mem mem Embryo brane brane H. straminea. Baer & Tenora (1 97) 3515 1,5 2,8 Tenora & Murai (197) 711 1,8 2, 23 15 79 19 28 11 15 12 16 52 54 21 24 12 13 65164 48 82 25 29 5576 2931 4 55 H. microstoma Liihe (191) 162 2,1 45 1 3 11 82 9 77 32 Meggitt & Subramanian 162 2,1 (1927) 45 3 11 829. Joyeux & Koboieff (1928) 8 2, J oyeux & Baer (1936) 8 2, [cited by Mahon (1954)] 2 2 6 27 15 135 5 6 27 15 135 5 9 x 8 4 3 9 x 8 Mahon (1954) 9 2, 187328 5918 1825 1 15 1198 4 65 4872 36 4 D vorak, et a!. (1961) 8 35 2, 2 23 28 15 13 5 8 9 3 Wahl (1967) 14 15 3, 1727 68 2832 11,5 12, 115 135 3545 4755 3 x 33 3841 3145,172, 216 336 72 14 22 3 12,1 18,6 72 2 3264 64 9 28 34 24 32 42 56 24 28 16 24

H. MARIA COLLINS excretory vessels and may extend into the medulla; it is 72 to 2 fl long by 32 to 64 fl wide. The external seminal vesicle is large in both the mature and early gravid proglottids. The lobed ovary and the vitellarium are situated in the centre of the proglottid. The seminal receptacle is large. The outer membrane of the ovum is 64 to 9 by 42 to 56 fl; the inner membrane 28 to 34 by 24 to 28 fl; the embryo 24 to 32 by 16 to 24 fl with embryonic hooks 15,1 to 19,2 fl long. Baer & Tenora (197) conclude that H. straminea has priority over Hymenolepis microstoma (Dujardin, 1845). The rostellar hooks of this species resemble those of H. nana but they have a well developed guard and short handle while those of H. nana have a smaller guard and a longer handle (Table 12). Hymenolepis taterae sp. n. Definitive host: Tatera (Tatera) brantsi Intermediate host: Unknown Type locality: Witwatersrand (Tv!.) Distribution: Material Syntypes from T. (T.) brantsi, Johannesburg, Roodepoort, Krugersdorp and Randfontein (Tv!.) (Onderstepoort Helminthological Collection, No. T. 164, Onderstepoort) Description The total length varies from 12 to 6 mm; the maximum width is 1, mm. The strobila consists of about 65 proglottids which are much wider than long; thus the terminal proglottids are 1, mm broad but only 14 fl long. The scolex is 152 to 264 fl and the suckers 8 to 144 fl in diameter. The retractable rostellum bears a crown of 24 to 35 hooks 2,9 to 28,1 fl long (Fig. 2). The three oval testes are almost in a straight line, two aporal and one poral [type VII of Skrjabin & Matiewossian (1945) cited by Zarnowski (1955)] (Fig. 21 a, b, c). They are 36 to 9 fl by 25 to 72 fl in (a ) (b) 5y 5}' 1)' FrG. 21 Sexually mature proglottid of H. taterae sp. n. a. Dorsal view b. Ventral view c. Transverse section FrG. 2 Rostellar hooks of H. taterae sp. n. The ventral excretory vessels, 16 fl in diameter, are united by a transverse vessel in the posterior part of each proglottid; the dorsal excretory vessels are 2 fl in diameter. The genital pores are unilateral and situated in the middle of the proglottid margin. The genital ducts pass dorsal to the excretory vessels when crossing from the medulla into the cortex. diameter. The vas deferens is short and forms a large external seminal vesicle which may partially overlay the cirrus pouch. The latter is long and narrow; extends well into the medulla and is 64 to 248 fl long by 16 to 4 fl wide. The lobed ovary and the vitellarium are situated in the centre of the proglottid; in whole mounts they are usually overlaid by the testes and the seminal receptacle. On leaving the seminal receptacle the vagina runs anteriorly to the cirrus pouch but in the region of the excretory vessels it crosses the cirrus pouch to open posteriorly to it in the genital atrium. The lumen of the vagina is 16 fl in diameter throughout most of its length but it gradually narrows to 4 fl before entering 47

CESTODES FROM RODENTS IN THE REPUBLIC OF SOUTH AFRICA the genital atrium. The outer membrane of the ovum is 5 to 74 JL by 38 to 56 JL and the inner membrane 28 to 38 JL by 24 to 34 JL in diameter; the embryo is 24 to 34 JL by 18 to 32 JL with embryonic hooks 14 to 16 JL long. Polar filaments are present. H. taterae resembles Hymenolepis sinensis Oldham, 1929 in the number and sie of the rostellar hooks but those of H. taterae differ in having long, pointed handles which curve downwards. The strobila of H. sinensis has 2, or more proglottids but H. taterae has only 65. In H. sinensis the cirrus pouch is 1 JL long and pyriform while that of H. taterae is 64 to 248 JL in length and long and narrow in shape (Fig. 21). The ovary of H. sinensis is large, occupying a quarter of the width of the proglottid; in H. taterae it occupies about a fifth. In view of these differences, and the fact that H. sinensis originates from China, these species are considered distinct from one another. H. taterae resembles H. straminea and H. nana in the number of rostellar hooks but those of H. taterae are different in shape and somewhat larger than those in the other two species. The shape of the rostellar hook of H. taterae also differentiates it from Hymenolepis australensis Sandars, 1957. According to Baer (1932), Hymenolepis myoxi (Rudolphi, 1819) has spines on the suckers; these are unarmed in H. taterae. Baer (1932) in redescribing Hymenolepis asymmetrica Janicki, 194 records 2 to 22 rostellar hooks which are 19 JL long. In H. asymmetrica the vitellarium is always aporal in relation to the ovary; in H. taterae it is poral in position. H. nana is the only other species in which polar filaments have been recorded. While examining the eggs of this species it was found that the membranes surrounding them become opaque once they are freed from the gravid proglottid and that the polar filaments are visible for a few minutes only. It is, therefore, possible that these structures are also present in other species of the genus, but have not yet been described. HosTPARASITE LIST of MYOMORPHA IN THE ETHIOPIAN REGION Aethomys (Aethomys) chrysophilus Catenotaenia compacta Catenotaenia Iucida Aethomys (Micaelamys) namaquensis Catenotaenia Iucida Raillietina (Raillietina) trapeoides Arvicanthis abyssinicus Hymenolepis microstoma Arvicanthis niloticus Inermicapsifer guineensis Raillietina (Raillietina) trapeoides Cricetomys buchanani Inermicapsifer guineensis Cricetomys gambianus Catenotaenia cricetomydis Inermicapsifer congolensis Inermicapsifer guineensis Cryptomys mechowi Dasymus incomtus Inermicapsifer congolensis Dendromys insignis Hymenolepis microstoma Heliophobius marungensis Catenotaenia lobata Heliophobius argenteocinereus Hybomys univittatus Hymenolepis pearsi Hymenolepis uncispinosa Lemniscomys griselda Inermicapsifer congolensis I nermicapsifer madagascariensis Lemniscomys striatus I nermicapsifer madagascariensis Raillietina (Raillietina) trapeoides Malacomys longipes Raillietina (Raillietina) baeri Leggada minutoides Hymenolepis microstoma Mus musculus Hymenolepis diminuta Hymenolepis microstoma Hymenolepis nana Oenomys I?Jpoxanthus Paranoplocephala omphalodes Otomys irroratus Inermicapsifer congolensis Paranoplocephala omphalodes Paranoplocephala otomyos sp. n. Raillietina (Raillietina) thryonomysi Otomys unisu!catus I nermicapsifer congolensis Paranoplocephala omphalodes Parotomys brantsi Raillietina (Raillietina) trapeoides Praomys (Mastomys) natalensis Catenotaenia lobata Hymenolepis diminuta Hymenolepis microstoma 1fymenolepis uncispinosa I nermicapsifer congolensis Raillietina (Raillietina) baeri Rillietina (Raillietina) madagascariensis Praomys (Praomys) jacksoni Hymenolepis diminuta 1fymenolepis microstoma Pelomys campanae Inermicapsifer guineensis Pelomys fa/lax Hymenolepis diminuta Rattus paedu!cus Rattus rattus Hymenolepis diminuta Inermicapsifer congolensis Inermicapsifer guineensis 48

H. MARIA COLLINS I nermicapsifer madagascariensis Raillietina (Raillietina) baeri Rhabdomys pumilio Catenotaenia lobata Hymenolepis microstoma Inermicapsifer congolensis Inermicapsifer guineensis Raillietina (Raillietina) trapeoides Saccostomys campestris Steatonrys pratensis Hymenolepis diminuta I nermicapsifer madagascariensis Tatera valida kempi Catenotaenia lobata Raillietina (Raillietina) trapeoides Tatera (Tatera) brantsi Hymenolepis microcantha sp. n. Hymenolepis taterae sp. n. Raillietina (Raillietina) trapeoides Thallomys paedulcus Inermicapsifer aberratus I nermicapsifer guineensis Thamnomys surdaster Hymenolepis diminuta Thryonomis S//)inderianus Raillietina (R aillietina) mahonae Raillietina (Raillietina) thryonom_ysi Raillietina (Raillietina) trapeoides AcKNOWLEDGEMENTS I wish to thank Prof. J. Meester, University of Natal and Mr. C. J. Nel, U niversity of Pretoria ; Dr. D. H. S. Davis and staff, Medical Ecology, S.A.I.M.R., Johannesburg; Dr. C. Crocker, Institute of Pathology, University of Pretoria; Dr. G. de Graaff, National Parks Board (formerly University of Pretoria); Dr. A. de Vos, Onderstepoort; Mr. J. Boomker, University of Pretoria, for collecting material. This survey would not have been possible had it not been for their generous assistance. The National Parks Board is thanked for permitting me to collect rodents in the Kruger National Park; Prof. J. Meester very kindly identified the host animals; Prof. R. K. Reinecke, Onderstepoort, read the manuscript ; Miss D. Brooker, Onderstepoort, assisted with the preparation of the drawings; Mrs. Z. A. Lepkowska translated articles from Polish; Mrs. S. E. T. Vermaakt typed the manuscript; Mrs. A. Wreyford typed the various drafts. Finally I wish to thank my promoters Prof. Dr. H. J. Schoonbee and Dr. Anna Verster for encouragement and advice throughout the investigation. REFERENCES ALLEN, G. M., 1939. Checklist of African mammals. Bull. Mus. comp. Zoo/. Harv., 83, 1763. 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