ACTA UNIVERSITATIS AGRICULTURAE ET SILVICULTURAE MENDELIANAE BRUNENSIS SBORNÍK MENDELOVY ZEMĚDĚLSKÉ A LESNICKÉ UNIVERZITY V BRNĚ DISCUSSION TO SEVERAL TAPEWORM SPECIES FROM THE FAMILIES HYMENOLEPIDIDAE, ANOPLOCEPHALIDAE AND DAVAINEIDAE pARASITIzING RoDENTS AND MAN

TENORA, F., BARUš, V., PROKEš, M.: Discussion to several tapeworm species from the families Hymenolepididae, Anoplocephalidae and Davaineidae parasitizing rodents and man. Acta univ. agric. et silvic. Mendel. Brun., 2004, LII, No. 1, pp. 23-28 With the more recent knowledge, the hypothesis by Joyeux and Baer (1929) is consulted: “... most of rarer species of tapeworms occurring in man are probably parasites of other mammals, specially of Rodentia .....“. In connection with that, the host specificity in several species from the families Hymenolepididae, Anoplocephalidae and Davaineidae is discussed. So far parasites of rodents are concerned, they are the species rodentolepis straminea, r. fraterna, Hymenolepis diminuta, H. pseudodiminuta, H. hibernia and inermicapsifer arvicanthidis. So far parasites of man are concerned, they are the species rodentolepis nana, Hymenolepis flavopunctata and inermicapsifer madagascariensis. Attention is drawn also to discrepancies in the opinions published on the views of hosts’ specificity or of zoogeographical distribution of several species from the family Davaineidae. Cestoda, Hymenolepididae, Anoplocephalidae, Davaineidae, host specificity, Rodentia, man Ročník LII 3 Číslo 1, 2004 23 In the current medical literature, in university text books and in special literature dealing with the taxonomic appurtenance of cestodes, we often meet with the statement that several tapeworm species from the family Hymenolepididae have as their definitive hosts both man and rodents. This opinion reflects the hypothesis published by Joyeux and Baer (1929) that “.... most of rarer species of tapeworms occurring in man are probably parasites of other mammals, especially of Rodentia ....“. Even when later, these authors leave successively their opinion (see below), a more complex view of the respective facts is missing, which would either support or argue against the above hypothesis. Below, the analysis and the opinion of the complicated situation in the systematic-taxonomical status of the above species is presented. MATERIAL AND METHODS The material was collected from different hosts and localites. Hymenolepididae Ariola, 1899 1. Hymenolepis diminuta (Rudolphi, 1819); host: Mus musculus; locality: Lednice, Czech Republic. 2. H. pseudodiminuta Tenora, Asakawa et Kamiya, 1994; hosts: Apodemus argenteus, A. speciosus; locality: Hokkaido, Japan. 3. Hymenolepis hibernia Montgomery J., Montgomery I. et Dunn, 1987; hosts: Apodemus flavicollis; locality: Lednice, Czech Republic; A. sylvaticus; locality: Hodonín, Czech Republic. 4. rodentolepis nana (von Siebold, 1852); host: man; locality: Nangrahar, Afghanistan. 5. R. fraterna (Stiles, 1906); hosts: Mus musculus; locality: Hodonín, Lednice, Czech Republic; M. musculus var. 24 F. Tenora, V. Baruš, M. Prokeš alb., locality: Nangrahar, Afghanistan (introduction with laboratory animal from Germany); host: Rattus norvegicus; locality: Mutěnice, Czech Republic. 6. R. straminea (Goeze, 1782), hosts: A. flavicollis, A. sylvaticus, A. microps, microtus arvalis; localities: Hodonín, Lednice, Mutěnice, Strážnice, Czech Republic; host: Cricetus cricetus; locality: Hungary. Anoplocephalidae Cholodkovsky, 1902 1. inermicapsifer arvicanthidis (Kofend, 1915); host: Heliophobius argenteocinereus; locality: Malawi, Africa. Davaineidae Braun, 1900 The species from the family Davaineidae presented in this study (raillietina celebensis, r. demerariensis, and R. trinitatae) were studied from the original descriptions and redescriptions presented in the literature cited. Preparation Cestodes, (Hymenolepididae, Anoplocephalidae) were fixed in 4% formaldehyd, stained with acetocarmine, dehydrated in alcohol, cleared in xylene, and mounted in Canada balsam. RESULTS Hymenolepididae The species R. nana was described as a parasite of man under the name Taenia nana, and into the genus Hymenolepis Weinland, 1858, it was transferred by Stiles (1906). The species H. nana was transferred and included in other genera as, e.g., Diplacanthus Weinland, 1858, Vampirolepis Spassky, 1954, Variolepis Spassky et Spasskaya, 1954 (Spassky 1954; Yamaguti 1959; Schmidt 1986 and others). At the present, accepted is commonly the inclusion of the species H. nana in the genus rodentolepis Spassky, 1954 (Vaucher 1994). It should be presented that Spassky (1954), establishing the genus rodentolepis, complicated somewhat the situation with that he included as younger synonyms of the typical species of the genus rodentolepis ( = R. straminea) also H. nana and H. fraterna. This his opinion was traded more frequently in the Russian literature (for more details see Ryzhikov et al. 1978). It was, of course, proved that the species R. straminea differs with a number of characters from R. fraterna and R. nana (Baer and Tenora 1970). We state that tapeworms R. nana and R. fraterna are species morphologically, anatomically and metrically very similar but biologically distinct. Used other characters than morphological and metrical ones (for a number of features see Kontrimavichus and Smirnova 1991; špakulová 2002), the validity and independence of both species can be confirmed (cf. Skrjabin and Matevosjan 1948; Baer in Baer and Tenora 1970; Joyeux in Hunkeler 1974; Hunkeler 1974; Tenora 2001, 2002; Macnish et al. 2002 and others). However, this cannot exclude that R. fraterna (a parasite of rodents) can be considered, eventually confirmed, to be the ancestor of R. nana (a parasite of man) during the evolutionary events of co-speciation. In contrast to R. nana, the species Taenia diminuta, described from rats, has changed its generic status only once, namely into Hymenolepis Weinland, 1858. Burt (1980) concentrated the highest number of information on the species H. diminuta and states that, in the literature in total, 28 its synonyms and 100 hosts’ species belonging to the orders Primates, Rodentia, Carnivora and Insectivora are reported. It was proved (e.g., Skrjabin and Matevosjan 1948; Tenora and Murai 1972; Tenora and Baruš 1972; Ryzhikov et al. 1978) that is necessary to remove from the synonyms of the species H. diminuta the resurrected species H. megaloon (von Linstow, 1901). It is, of course, wholly evident that the synonym of H. diminuta is H. diminutoides Cholodkowsky, 1913. The complex synonymy and extraordinarily numerous and diverse list of definitive hosts of H. diminuta have led the authors Mas-Coma et al. (1980) to the opinion that, under the name H. diminuta, a complex of more species is reported in the literature. Moreover, this is completed by the information that, in Spain, Apodemus sylvaticus is parasitized by tapeworms morphologically very similar to H. diminuta which will be described as a new species. Later on, erroneous determination of tapeworms as H. diminuta from A. sylvaticus was corrected by Montgomery et al. (1987), who described a new species Hymenolepis hibernia. Tenora et al. (1994) corrected the erroneous determination of the material from Apodemus spp. in Japan with the description of a new species H. pseudodiminuta (its validity was confirmed by Ishih et al. 2003). All above examples document the justification of opinions by Mas-Coma et al. (1980). The analysis of the species complex of the supraspecies H. diminuta means the actual problem. It is highly likely that the species H. diminuta is not, in all probability, regular parasite of man. This is already suggested by ancien efforts by several authors who reported tapeworms from man, similar to H. diminuta, under the names of independent species: Taenia flavopunctata Weinland, 1858, T. flavomaculata Leuckart, 1863, T. varesina Parona, 1884, T. minima Grassi, 1886. Other signal suggesting the possibility and evidently reality for excluding the taxon determined hitherto as H. diminuta parasitizing the man, and for establishing it as an independent taxon, is a low frequency of its findings. So, e.g., Burt (1980) reports 95 findings in man, and of those only two from Europe (determined only on eggs – England, Poland). Contrary, at the same time, the species H. diminuta is a frequent and obligatory parasite in rodents in Europe. The other 93 findings of H. diminuta are from the territory outside Europe (these findings show a mozaic occurrence in the human popuDiscussion to several tapeworm species 25 lation living at a low life level) (Edelman et al. 1965; Burt 1980; Combs and Crompton 1991). The necessity of coming back to the opinion that H. diminuta is a parasite of rodents, and not regularly parasit of man, is also suggested by the fact that the above tapeworm species described as n. sp. from man were declared and arranged as synonyms of H. diminuta as early as in the ancien past, namely by Blanchard (1891) and Grassi and Rovelli (1892). Later on, this opinion was taken over uncritically not only by the medical praxis but also by other specialists. It can be assumed that the species T. flavopunctata is a sibling species of the taxon H. diminuta (cf. Tenora 2001, 2002). This suggests that H. diminuta (a parasite of rodents), or of very similar species, can be the ancestor of H. flavopunctata (a parasite of man). For the exact solution of defined problems, methods of allozyme electroforesis (Beveridge 1998) and modern analyses of biological properties (Brouks 1988, Mongomery et al. 1998, Macnish et al. 2002, Ishih et al. 2003) should be applied. In the case of H. diminuta the authors prefere the host and ecological specificity contrary to the morphology and anatomy of the parasite. The identical problem is in the case of mesocestoides variablilis Mueller, 1927 (parasit of man) and M. lineatus (Goeze, 1782) (parasit of Carnivora), compare Coombs and Crompton, (1991), Fuentes et al. (2002), (2003), Tschertkowa and Kosupko, (1978). The similar problem in the case of Taenia solium (Goeze, 1782) and T. asiatica Eom et Rim, 1993, compare in Hoberg, (2002) and Galan–Puchades and Fuentes, (2000).