Journal of Helmintkology (1978) 52, 205-209

Tapeworms of the Chaco Boreal, Paraguay, with two new species GERALD D. SCHMIDT Department of Biology, University of Northern Colorado, Greeley, Colorado 80639, U.S.A. and ROBERT L. MARTIN Department of Biology, University of Maine, Farmington, Maine 04938, U.S.A. ABSTRACT Mathevotaenia paraguayae sp. n. is described from Euphractus sexcinctus. It differs from M. didelphidis (Rud.) in having a transverse cirrus pouch anterior to the vagina. Monoecocestus mackiewiczi sp. n. was found in Phyllotis sp. It differs from M. parcitesticulatus Rego in having a much smaller scolex, a poral ovary, and a cirrus pouch 360 to 440 um long. Other species reported are Taenia macrocystis, Taenia omis'sa, Spirometra mansonoides, Spirometra gracilis, Spirometra longicollis, Diphyllobothrium trinitatis, Atriotaenia parva, and Mathevotaenia tetragonocephala.

This report is based on specimens collected in 1973 and 1974 by the second author in the thorn forest of the Chaco Boreal of Western Paraguay. Living specimens were killed in water and fixed in AFA or formalin. Whole mounts were stained with Semichon's carmine and mounted in balsam. All measurements are in micrometres unless otherwise stated. The results of this survey are summarized in Table 1. Some of these are of uncommon interest and deserve further discussion. Mathevotaenia paraguayae sp. n.

(Figs. 1-3) Twelve worms found in the small intestine of a yellow armadillo, Euphractus sexcinctus, were found to represent a new species, and are the basis of the following description. Description: Scolex (Fig. 1) 215 to 280 long (measured to hind margin of suckers), 375 to 465 greatest width. Suckers oblong, deeply imbedded in scolex, 150 to 200 long, 140 to 190 wide. Opening into sucker is a narrow slit. Neck narrower than scolex, about 400 long. Strobila 20 to 25 mm long, 750 to 800 greatest width near middle of worm. Up to 110 segments per strobila. Proglottids craspedote, anapolytic. Male and female gonads (Fig. 2) mature at same time. Genital ducts dorsal to osmoregulatory canals. Genital atrium 40 to 50 deep, about 10 wide, irregularly alternating. Male genitalia: 16 to 25 testes in continuous field posterior and somewhat lateral to ovary; each 30 to 60 in diameter. Cirrus pouch 100 to 160 long, 30 to 80 wide, transverse in proglottid. Coiled vas deferens engorged with sperm. Internal seminal vesicle absent. Cirrus short, unarmed. Female genitalia: ovary median, not reaching osmoregulatory canals; maximum size 90 to 120 long, 220 to 280 wide. Vitellarium compact, posterior to ovary, 40 to 50 long, 90 to 100 wide. Vagina posterior to cirrus pouch, unarmed. Seminal receptacle small. Uterus develops as a very thin-walled reticulum which quickly fills and disappears, leaving developing eggs scattered throughout medullary parenchyma. Mature oncospheres 22 to 35 diameter, with hooks 16 to 18 long; egg membrane 30 to 40 wide; outer egg capsule 60 to 80 wide (Fig. 3). Type host: yellow armadillo, Euphractus sexcinctus (Desmarest, 1804) Location: small intestine Type locality: Juan de Zalazar, Boqueron, Paraguay Type specimens: USNM Helm. Coll. holotype no. 73085, paratypes no. 73086. 205

G. D. SCHMIDT and R. L. MARTIN Remarks: In number of testes and size of cirrus pouch Mathevotaenia paraguayae is most similar to M. didelphidis (Rud., 1819) Spassky, 1951. It differs, however, in two easily discernible characters: (1) the cirrus pouch of M. didelphidis curves anteriad at about a 45 degree angle, while it is straight and transverse in M. paraguayae, and (2) the vagina of M. didelphidis opens anterior to the cirrus pouch and has a loop near the distal end, while it terminates posterior to the cirrus pouch and has no loop in the case of M. paraguayae. Monoecocestus mackiewiczi sp. n. (Figs. 4-6) Four worms were found in the intestine of a mouse, Phyllotis sp., and represent a new species. The mouse also appears to be a new species. Description: Scolex (Fig. 4) 175-225 long (measured to hind margin of suckers), 360 to 415 greatest width. Suckers round, deeply imbedded in scolex, 120 to 160 wide. Neck wider than scolex, 280 to 320 long. Stobila 70 to 115 mm long, 3.5 to 4.5 mm greatest width near middle of worm. About 200 segments per strobila. Proglottids craspedote, anapolytic, much wider than long. Male and female gonads (Fig. 5) mature at same time. Genital ducts dorsal to osmoregulatory canals. Genital atrium variable in size and shape depending on extent of withdrawal of cirrus; atrium capable of complete eversion in gravid proglottids. Atrium in mature segments 200 to 270 deep, 140 to 160 wide, irregularly alternating. Male genitalia: 42 to 55 testes in two fields lateral to ovary, occasional few posterior to ovary; usually about two-thirds of them aporal; each 30 to 48 wide. Cirrus pouch in mature segments 360 to 440 long, 140 to 160 wide; transverse in proglottid, exceeding osmoregulatory canals. Internal seminal vesicle present. Cirrus long, densely covered with small spines. Female genitalia: ovary poral, about 160 long, 880 to 1000 wide. Vitellarium compact, posteroventral to ovary, 140 to 175 long, 240 to 320 wide. Vagina opens anterior to cirrus pore, large, persistent. Seminal receptacle small. Uterus an irregular, transverse sac, persistent. Outer egg membrane (Fig. 6) thick, 58 to 60 wide. Oncosphere about 15 wide, pyriform apparatus 20 to 22 long. Type host: Phyllotis sp. (Rodentia: Cricetidae) Location: small intestine Type locality: Juan de Zalazar, Boqueron, Paraguay Type specimens: USNM Helm. Coll. holotype no. 73083, paratypes no. 73084. Remarks: Monoecocestus mackiewiczi is very similar to M. parcitesticulatus Rego, 1960, from guinea pigs in Brazil and Peru. Upon careful comparison it is found to differ in the following ways: (1) the scolex of M. mackiewiczi measures 175 to 200 by 360 to 400, while that of M. parcitesticulatus is 514 by 597; (2) the ovary of M. mackiewiczi is distinctly poral, while it is only slightly poral in M. parcitesticulatus; (3) the cirrus pouch of M. mackiewiczi is 360 to 440 long and 145 to 160 wide, and it far exceeds the ventral osmoregulatory canal, compared with 278 by 87 and barely reaching the canal in M. parcitesticulatus (in mature but not gravid segments in both cases); (4) the pyriform apparatus of M. mackiewiczi is about 20 long, while it is 29 long in M. parcitesticulatus. These differences, especially the first three, indicate that the two are separate species. The new species is named in honour of Dr. John S. Mackiewicz, who has made many important contributions to our knowledge of cestodes. 206

Tapeworms from Paraguay

FIGS. 1-3. Mathevotaenia paraguayae sp. n. from an armadillo. FIG. 1. Scolex and neck. FIG. 2. Mature proglottid. FIG. 3. Egg. FIGS. 4-6. Monoecocestus mackiewiczi sp. n. from a mouse. FIG. 4. Scolex. FIG. 5. Mature proglottid. FIG. 6. Egg.

Notes on other species recovered It was interesting to find Taenia macrocystis cysticerci in Sylvilagus brasiliensis, the type host. To our knowledge it has not been reported in the cottontail rabbit intermediate host since its original description by Diesing (1850), based on material collected by Natterer in Brazil. The adult is known from several species of cat, but Felis geoffroyi is a new host record. Specimens deposited: USNM Helm. Coll. nos. 74614, 74616. We are unaware of any reports of Diphyllobothrium trinitatis since its first description by Cameron (1936) from the crab-eating raccoon in Trinidad. Our records are new for Paraguay and for Nasua nasua, an animal closely related to the raccoon. Specimens deposited: USNM Helm. Coll. no. 73091. Lopez-Neyra and Diaz-Ungria (1957) seem to have been the first to report Spirometra mansonoides from South America (Venezuela). Its presence was confirmed in Brazil and 207

G. D. SCHMIDT and R. L. MARTIN

Ecuador by Mueller et al. (1975), so our finding it in Paraguay is not unexpected. Felis geoffroyi, F. yagouaroundi, Dusicyon gymnocercus, and Cerdocyon thous are all new host records. Specimens deposited: USNM Helm. Coll. no. 73095. Spirometra gracilis has not been reported after the initial description by Baer (1927). The jaguarundi, fox, and Geoffroy's cat are new host records. Specimens deposited: USNM Helm. Coll. no. 73093. Our identification of Spirometra longicollis (Parodi et Widakowich, 1917) comb. n. must be tentative, as the only description of the species, the original one, is incomplete. However, the exceptionally long neck, egg size and other characters are very similar to our specimens; further, the host is the same. The uterus in our specimens is distinctive: there is a single, slender loop to one side of midline or the other, which terminates in a single, simple, median chamber. Specimens deposited: USNM Helm. Coll. no. 73087. Diphyllobothrium decipiens (Diesing, 1850) Gedoelst, 1911 was originally described from several species of cat from Brazil. Although Diesing's description is too meagre to allow recognition of the species, several authors have reported it from Europe, South America, and Asia. Ariola (1900) redescribed the species but apparently with specimens obtained in European domestic cats. We examined the type specimens of Diesing's species and found the uterus, which has never been described, to consist of two thick loops on each side of midline, giving the appearance of a four-chambered sac. This is unlike any species we encountered in this study, and should be a useful character in future recognition of D. decipiens.

TABLE 1 Some cestodes of vertebrates from Paraguay (* indicates new host record) Host Geoffroy's cat, Felis geoffroyi paraguae (Pocock)

Parasite

Taenia macrocystis (Diesing, 1850) Liihe, 1910* Spirometra mansonoides (Mueller, 1935)* Spirometra gracilis (Baer, 1927) Wardle, McLeod, et Stewart, 1947* Taenia macrocystis Jaguarundi, Felis yagouaroundi eyra Fischer Spirometra gracilis* Spirometra mansonoides* (?) Spirometra longicollis (Parodi et Widakowich, 1917) comb. n. Taenia omissa Liihe, 1910 Cougar, Felis concolor acrocodia Goldman Spirometra sp. Diphyllobothrium trinitatis Cameron, 1936* Coati-mundi, Nasua nasua aricana Vieira Atriotaneia parva Sandground, 1926 Crab-eating raccoon, Procyon cancrivorus (Cuvier) Diphyllobothrium trinitatis Mathevotaenia paraguayae sp. n. Six-banded armadillo, Euphractus sexcinctus Sparganum sp. flavimanus (Desmarest) Giant anteater, Myrmecophaga tridactyla L. Mathevotaenia tetragonocephala (Bremser in Diesing, 1856) Spassky, 1951 Mathevotaenia sp. Collared anteater, Tamandua tetradactyla (L.) Taenia macrocystis, (cysticerci) Rabbit, Sylvilagus brasiliensis (L.) Spirometra mansonoides* Crab-eating fox, Cerdocyon thous (Bur.) "Fox", Dusicyon gymnocercus (Fisch) Spirometra mansonoides* "Mouse", Phyllotis sp. Monoecocestus mackiewiczi sp. n. Hymenolepis sp. "Mouse", Akodon sp. Brocket deer, Mazama gouazoubira (Fisch) (?) Echinococcus sp., (sterile cyst) Hymenolepis sp. Fishing bat, Noctilio leporinus (L.) Chaco chachalaca, Ortalis canicollis (Wagler) Raillietina sp. 208

Tapeworms from Paraguay ACKNOWLEDGEMENTS We wish to thank the University of Connecticut Research Foundation and National Geographical Society who provided grants to Dr. Ralph M. Wetzel, University of Cormecticut, who led the collecting expedition; Robert J. Eaton of Juan de Zalazar; Hernando Bertoni, Minister of Agriculture and Livestock; Dr. E.. Kritscher, Naturhistorisches Museum Wien, for loan of type specimens of Diphyllobothrium decipiens. Final efforts were sponsored by a grant from the University of Northern Colorado Research Committee.

REFERENCES ARIOLA, V. (1900) Revisione della famiglia Bothriocephalidae s. str. Archives de Parasitologie, 3, 369-484. BAER, J. G. (1927) Die Cestoden der Saugetiere Brasiliens. Abhandlungen der Senckenbergischen Naturforschenden Gesellshaft, 40, 377-386. CAMERON, T. W. M. (1936) Studies on the endoparasitic fauna of Trinidad. III. Some parasites of Trinidad carnivores. Canadian Journal of Research, 14, 25-38. DIESING, C. M. (1850) Systema Helminthum. Vol. I. Vienna, p. 588. LOPEZ-NEYRA, C. R. and DIAZ-UNGRIA, C. (1957) Cestodes de Venezuela. III. Sobre unos cestodes intestinales de reptiles y mamiferas Venezolonos. Memorias del Sociedad de Ciencias Naturales •'La Salle", 17, 28-63. MUELLER, J. F., FROES, O. M. and FERNANDEZ, T. R. (1975) On the occurrence of Spirometra mansonoides in South America. The Journal of Parasitology, 61, 774-775. PARODI, S. F. and WIDAKOWICH, V. (1917) Cestodes del genero Bothryocephalus, parasitos de algunas especies de nuestros felinos salvajes. Revista del Jardin Zoologico de Buenos Aires, II, 13, 222-227. REGO, A. A. (1961) Revisao do genero Monoecocestus Beddard, 1914 (Cestoda, Anoplocephalidae). Memorias do Instituto Oswaldo Cruz, 59, 325-354. Received 1 August, 1977.

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Tapeworms of the Chaco Boreal, Paraguay, with two new species.

Journal of Helmintkology (1978) 52, 205-209 Tapeworms of the Chaco Boreal, Paraguay, with two new species GERALD D. SCHMIDT Department of Biology, Un...
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