Anoplocephaline Cestodes Research Articles

Phylogenetic relationships of the anoplocephaline cestodes of Australasian marsupials and resurrection of the genus Wallabicestus Schmidt, 1975

A phylogenetic analysis was carried out on rDNA of 45 species of anoplocephaline cestodes from marsupial hosts. The exclusively Australasian genera Progamotaenia Nybelin, 1917, Triplotaenia Boas, 1902, Paramoniezia Maplestone & Southwell, 1923 and Phascolotaenia Beveridge, 1976 formed a monophyletic clade, and the previously suggested relationship between the Australasian species of the cosmopolitan genus Bertiella Stiles & Hassall, 1902 and species of Progamotaenia was supported. A low degree of phyletic co-evolution was detected within endemic Australasian clades. Colonisation rather than co-speciation appeared to be the principal means of diversification within the Australasian anoplocephaline radiation. The clade of bile duct-inhabiting Progamotaenia species emphasises the role of microhabitat rather than host species as a driver of speciation. Triplotaenia undosa Beveridge, 1976 described from a wide variety of macropodid hosts was found to be polyphyletic and a proposition was made to resurrect Wallabicestus Schmidt, 1975, with W. ewersi Schmidt, 1975 as the type-species and including W. ualabati (Beveridge, 2009) n. comb. [previously Progamotaenia ualabati Beveridge, 2009].

The status of the genus Hokkaidocephala Tenora, Gulyaev & Kamiya, 1999 (Cestoda: Anoplocephalidae), parasites of the endemic Japanese field mice (Apodemus spp.)

The present study reconsiders the status of the genus Hokkaidocephala Tenora, Gulyaev & Kamiya, 1999 by examining available specimens of the type species H. apodemi (Iwaki, Tenora, Abe, Oku & Kamiya, 1994) (syn. Andrya apodemi) and comparing them with related anoplocephaline cestodes. It is shown that Hokkaidocephala apodemi does not differ fundamentally from related genera in the development of the reproductive organs, contrary to the argument of Tenora et al. (1999). In Hokkaidocephala and related genera (Anoplocephaloides Baer, 1923 s. str. and Microcephaloides Haukisalmi, Hardman, Hardman, Rausch & Henttonen, 2008), sperm appears in the seminal receptacle (female organ) simultaneously or slightly earlier than in the seminal vesicles (male organs) and the ovary and vitellarium are absent in postmature proglottides where testes still persist and usually reach their maximum size. However, Hokkaidocephala is considered here a valid genus differentiated from the related genera by its unique uterine structure and development. Presently, Hokkaidocephala includes two host-specific species [H. apodemi and H. baeri (Rausch, 1976) n. comb. (= Anoplocephaloides baeri Rausch, 1976)] parasitizing endemic Japanese field mice (primarily Apodemus argenteus).

Molecular systematics of the Holarctic Anoplocephaloides variabilis (Douthitt, 1915) complex, with the proposal of Microcephaloides n. g. (Cestoda: Anoplocephalidae)

Phylograms based on mitochondrial cytochrome oxidase I gene sequences show that the Anoplocephaloides variabilis (Douthitt, 1915)-like cestodes (Cestoda: Anoplocephalidae) from voles (Microtus spp.) and Paranoplocephala krebsi Haukisalmi, Wickström, Hantula & Henttonen, 2001 from collared lemmings (Dicrostonyx spp.) comprise a monophyletic group within the anoplocephaline cestodes. The patterns of phylogenetic, biological and/or biogeographical distinction suggest six or seven species of A. variabilis-like cestodes, including P. krebsi. However, at this time we decline to describe them as a series of new species as no straightforward morphological differences were found between the A. variabilis-like cestodes. A new genus, Microcephaloides n. g., is proposed for the cestodes earlier assigned to A. variabilis, A. cf. variabilis, A. tenoramuraiae Genov & Georgiev, 1988 and P. krebsi. A redescription is provided for the type-species, M. variabilis n. comb., from pocket gophers (Geomys spp. and Thomomys spp.). In addition to Anoplocephaloides Baer, 1927 (sensu stricto) and Microcephaloides, Paranoplocephaloides Gulyaev, 1996, Flabelloskrjabinia Spasskii, 1951 and Leporidotaenia Genov, Murai, Georgiev & Harris, 1990 are considered valid genera among cestodes previously assigned to Anoplocephaloides (sensu Rausch, 1976). The host spectrum and present phylogenetic data suggest that Microcephaloides has been primarily associated with voles (Microtus spp.) and its basal lineage now occurs in M. guentheri (Danford & Alston) in Turkey. Although the distribution and current host of the basal lineage suggest a western Palaearctic origin, subsequent diversification has probably occurred in eastern Beringia, because most of the more derived lineages occur partly or exclusively in Alaska.

Anoplocephalid cestodes of wood rats (Neotoma spp.) in the western U.S.A.

AbstractThis study reviews the taxonomy of anoplocephaline cestodes of wood rats, Neotoma cinerea, N. fuscipes and N. mexicana (Sigmodontinae) in the western and south-western U.S.A. The anoplocephaline fauna included five species, only one of which, Andrya neotomae Voge, 1946, was relatively common and occurred in all three host species. Other species were Paranoplocephala freemani Haukisalmi, Henttonen et Hardman, 2006, P. primordialis (Douthitt, 1915), both host-generalist species of North American rodents, and two apparently undescribed species of Paranoplocephala s. str. Aprostatandrya octodonensis Babero et Cattan, 1975 from the indigenous South American rodent Octodon degus is regarded as a junior synonym of A. neotomae. A redescription is provided for A. neotomae.

Morphological Characterisation of Andrya Railliet, 1893, Neandrya n. g. and Paranoplocephala Lühe, 1910 (Cestoda: Anoplocephalidae) in Rodents and Lagomorphs

The taxonomic significance of the main morphological features of the 25 species allocated to Andrya Railliet, 1893 and Paranoplocephala Lühe, 1910 is re-evaluated in the light of the recent molecular phylogenetic hypotheses for anoplocephaline cestodes. The present analysis and the existing phylogenetic data suggest that the structure and complexity of the early uterus are not, as previously assumed, the main phylogenetic or systematic determinants for anoplocephaline cestodes. Instead, the position of the early uterus with respect to other organs, combined with the morphology of the female genitalia, appear to allow a fairly straightforward discrimination of the three genera recognised here, without contradicting current phylogenetic hypotheses. A new genus, Neandrya n. g., is proposed for N. cuniculi (Blanchard, 1891) n. comb. (previously in Andrya), amended diagnoses are provided for Andrya and Paranoplocephala and a diagnostic key to these three genera is presented.

Molecular Phylogeny and Systematics of Anoplocephaline Cestodes in Rodents and Lagomorphs

A molecular phylogenetic hypothesis is presented for the anoplocephaline cestodes of placental mammals based on sequence data from the mitochondrial cytochrome c oxidase I (COI) gene, the nuclear-encoded 28S rRNA gene and the internal transcribed spacer region I of rRNA (ITS1). The material consists of 35 species representing nine genera of cestodes, with emphasis on taxa parasitising rodents and lagomorphs in the Holarctic region. The resulting phylogenies show considerable disagreement with earlier systematic and phylogenetic hypotheses derived from morphology. Specifically, the results contradict the view of uterine morphology being the primary determinant of deeper phylogenetic splits within Anoplocephalinae. Also, the role of genital duplication as a means of generic divergence was not found to follow consistently the pattern suggested by earlier hypotheses. Colonisation of novel host lineages has evidently been the predominant mode of diversification in anoplocephaline cestodes of placental mammals; evidence for phyletic co-evolution was obscure. The phylogenies consistently distinguished a large monophyletic group including all species from arvicoline rodents (voles and lemmings), primarily representing the genera Anoplocephaloides Baer, 1923 and Paranoplocephala Lühe, 1910. Phylogenetic relationships within the "arvicoline clade" of cestodes were generally poorly resolved. Consistent support for nodes above and below the unresolved polytomy indicates a rapid radiation involving a nearly simultaneous diversification of many lineages, a scenario also proposed for the arvicoline hosts.

The seasonal abundance of oribatid mites (Acari: Cryptostigmata) on an irrigated Kikuyu grass pasture

The seasonal abundance of oribatid mites, the intermediate hosts of anoplocephaline cestodes, was studied on an irrigated Kikuyu grass pasture in Transvaal, South Africa. The following species were consistently present: Galumna nuda. Galumna rasilis, Scheloribates elsi and Tectocepheus sarekensis. The mean total number of mites on the herbage peaked in January (3.7 mites g 1dry herbage) and correlated positively with temperature, relative humidity, rainfall and soil moisture on the day of collection. The mean total number of mites on the herbage including the top 5 cm of soil peaked in April (3.757 mites m 2) and May (3.761 mites m 2), and these numbers correlated positively with radiation on the day of collection. There was a significant decline in the number of mites on the herbage at midday and in the afternoon, but there was no difference in the number on the herbage including the top 5 cm of soil at different times of day. The trends in abundance are also discussed separately for each of the four species.

The genera Paranoplocephala Lühe 1910 and Anoplocephaloides Baer, 1923 (Cestoda: Anoplocephalidae), with particular reference to species in rodents.

The present study redefines the systematic arrangement and taxonomic status of some anoplocephaline cestodes previously allocated among the genera Andrya Railliet, 1893. Paranoplacephala Lühe, 1910, and Aprostatandrya Kirshenblat, 1938. Comparisons indicated that Paranoplocephala omphalodes (Hermann, 1783), type species of Paranoplocephala, differs morphologically from its congeners, all of which conform to the diagnosis of the genus Anoplocephaloides Baer, 1923, hitherto considered a synonym of Paranoplocephala, and that it cannot be distinguished at the generic level from Aprostatandrya macrocephala (Douthitt, 1915), type of the genus Aprostatandrya. The distinction between Andrya and Aprostatandrya based on the presence or absence of a "prostate gland" does not exist, for neither the type species of Andrya, A; rhopalocephala (Riehm, 1881), nor A. cuniculi (Blanchard, 1891) possesses such an organ; rather, the structure of the external seminal vesicle in these cestodes does not differ fundamentally from that in P. omphalodes or in species of Aprostatandrya. Based on these findings, Aprostatandrya Kirshenbalt, 1938 is placed in synonymy with Paranoplocephala Lühe, 1910, and Anoplocephaloides Baer, 1923 is restored to contain all species other than P. omphalodes formerly allocated to the genus. Paranoplocephala. If the pattern of development of the uterus in A. rhopalocephala is found to be like that in P. omphalodes, Paranoplocephala Lühe, 1910 will in turn also become a synonym of Andrya Railliet, 1893. Three additional species of Anoplocephaloides are described, and the zoogeography of Anoplocephaloides spp. and their mammalian hosts is briefly discussed.

On Some Anoplocephaline Cestodes from Pikas, Ochotona spp. (Lagomorpha), in Nepal, with the Description of Ectopocephalium abei gen. et sp. n.

Three species of anoplocephaline cestodes, Anoplocephalinae gen. et sp. indet., Schizorchis cf. altaica Gvozdev, 1951, and Ectopocephalium abei gen. et sp. n., are reported from pikas, Ochotona roylei (Ogilby) and 0. macrotis (Giinther), from central Nepal. The taxonomic status of Schizorchis esarsi Lovekar et al., 1972, and of Schizorchodes dipodomi Bienek and Grundmann, 1972, is discussed, with the conclusion that both species belong in the subfamily Linstowiinae, near the genus Atriotaenia Sandground, 1926. Ectopocephalium gen. n. is distinguished from other genera in the subfamily Anoplocephalinae by the modified anterior portion of the strobila, which is embedded in the tissue of the sacculus rotundus (lower ileum) of the host, and by the arrangement of the genital organs. The attachment of E. abei and the consequent tissue response are described, and the zoogeography of cestodes in pikas is briefly discussed. During March-July 1968, a biological survey of Nepal was undertaken by an expedition from the University of Hokkaido, with support of the Hokkaido University Himalayan Committee. The objectives included an investigation of the small mammals of the region, under the direction of Dr. Hisashi Abe, of Hokkaido University, who kindly made available to us a large series of preserved carcasses for parasitological examination. Information concerning the taxonomy and ecology of these mammals has been published by Abe (1971). Among the preserved materials was a series of pikas, Ochotona spp., from which we obtained anoplocephaline cestodes representing three species. One of these exhibited a combination of characters that was unlike any within the genera recognized in the subfamily Anoplocephalinae. The results of the study of these cestodes are reported in the present

The incidence of anoplocephaline cestodes in a population of rabbits in Surrey, England.

Samples of approximately five wild rabbits were obtained each week over the 4-year period 1967–70, and their intestines examined for adult tapeworms. Pour species of anoplocephalinae were found,Cittotaenia pectinata, C. denticulata, C. ctenoidesandAndrya cuniculi. C. pectinatawas found in 29% of the 968 rabbits examined,C. denticulatain 14%,C. ctenoidesin 12% andA. cuniculiin 6%. There were 63 double infections, of which 44 involvedA. cuniculi, and three triple infections. The most frequent number of worms per infection was one forC. pectinataandA. cuniculiand two forC. denticulataandC. clenoides, although the mean numbers for these species were respectively 7·2, 5·3, 2·7 and 1·8.A. cuniculi, and to a lesser extentC. ctenoides, were very restricted in distribution within the study area, occurring only on the higher, chalk downland (250–550 ft A.S.L.). This finding implies that individual rabbits usually spend their lives within relatively small areas. Considering all the material a seasonal trend in host infection rate was apparent only withC. denticulata, which increased in abundance from July to a peak in September and then declined gradually to a low level, lasting from April to June. Infections, including small specimens of any of the four species, were rare from February to June and it is presumed that infected intermediate hosts were correspondingly rare during this period. Except forC. denticulata, which was found more frequently in male rabbits than females, the sex of the host had no influence on its likelihood of infection. However, except forC. pectinata, the age of the host was important. Thus,C. denticulataandA. cuniculiwere significantly more frequent in juveniles than adults whilst the reverse was true forC. ctenoides. Small (‘young’) specimens ofC. denticulataandA. cuniculiwere more frequent in juvenile than adult rabbits, but the difference was not significant for the other two species. It is concluded that onlyC. ctenoidessurvives for more than a year and that this species andC. pectinataare more likely to infect rabbits that are adult than are the other two species.

Distribution and Specificity of Helminths in Microtine Rodents: Evolutionary Implications

Among those genera of microtine rodents having holarctic distribution, the parasitic helminths of North American representatives of Dicrostonyx, LLemmus, Clethrionomys, and Microtus are relatively well known. These helminths demonstrate a highly developed phylogenetic specificity, being restricted essentially to the Microtinae, but few show any evidence of host-specificity at the generic level. Both nematodes and cestodes are prevalent endoparasites of these rodents in boreal North America. For the purposes of the present work, emphasis is placed upon the cestodes, most of which are members of the subfamily Anoplocephalinae. Some of the latter are unusually interesting in that they occur in a variety of host-species over extensive geographic ranges. With the exception of Paranoplocephala omphalodes (Hermann, 1783), it has not been determined whether these cestodes are holarctic. It is evident, however, that some described from microtine rodents in Eurasia correspond morphologically to species in North America. It appears in Alaska that significant relationships between the distribution of certain cestodes and the distribution of their hosts may be recognized. Any attempt to evaluate these apparent relationships is contingent upon determining whether or not these cestodes are holarctic. The study of this problem has now been undertaken in light of work published recently by Eurasian helminthologists and the acquisition of additional comparative material. This paper contains a review of the taxonomic status of anoplocephaline cestodes from microtine rodents and a discussion of some of the evolutionary and zoogeographic implications of distribution and host-occurrence of these and other helminths. I. The genus Andrya. Five species of Andrya parasitic in microtine rodents may be considered valid at the present time. 1. Andrya macrocephala Douthitt, 1915, is the most common cestode observed in species of Microtus in North America. Its unusually wide range of normal morphological variation (Rausch and Schiller, 1949a; Rausch, 1952) has led to the description of some of its variants as distinct species. A. macrocephala has been well characterized from North American material. Two species closely resembling A. mwacrocephala have been described in Eurasia; however, their validity is questionable. One of these, A. bialozuizensis Soltys, 1949, has been reported only from Poland. A. caucasica Kirshenblat, 1938, was described fronm Transcaucasia. It was suggested by Rausch and Schiller (1949a) that A. caucasica might be conspecific with A. macrocephala. This possibility was not considered by Spasskii (1951), in his monograph of anoplocephalid cestodes. Later, Spasskii, Romanova, and Naidenova (1951) made a survey of the helminth parasites of mnuskrats (an introduced species) in Russia, and from this host collected specimens of Andrya in the Kurgansk and Archangel'sk Oblasti. They also had cestodes of this genus from voles in Pribaikal'. These workers noted the lack of significant morphological differences between these specimens and the description of A. macrocephala. It was also observed that the cestodes from the voles of Pribaikal' did not differ in any essential way from A. caucasica and A.

Growth in the laboratory of economically important orbited mites.

RECENT quantitative analyses of the microfauna of the soil have shown that oribatid mites are by far the largest representatives. Their economic value is well known; they carry the infective stages of anoplocephaline cestodes, some of which are important parasites of sheep. Attention has recently been directed to their role in the promotion of humus formation from leaf litter on the forest floor, and at the same time it was pointed out that little was known of the biology of these mites1.

Studies on the Life History of the Anoplocephaline Cestodes of Hares and Rabbits

SUMMARY Thousands of free-living mites were collected near Hamburg from areas where wild rabbits were numerous and heavily infected with anoplocephaline cestodes. The mites were dissected and cysticercoids were removed from Scutovertex minutus, Scheloribates laevigatus and Pelops tardus. The larvae were fed to young, domestic rabbits and a cysticercoid froIn S. minutus developed into a sexually mature specimen of Cittotaenia ctenoides. Other mites, collected from areas where they would not be exposed to eggs of anoplocephaline cestodes, were fed eggs of Cittotaenia ctenoides and C. denticulata. Developmental stages of C. ctenoides were recovered from the body cavities of Scutovertex minutus, Galumina obvious, Pelops acromius, Liacarus coracinus, Notaspis coleoptratus, Liebstadia similis, Xenillus tegeocranus, Scheloribates laevigatus, Cepheus cepheiformis, Trichoribates incisellus, and Galumna This content downloaded from 157.55.39.76 on Wed, 20 Apr 2016 06:52:21 UTC All use subject to http://about.jstor.org/terms THE JOURNAL OF PARASITOLOGY nervosus. The larvae attained the cysticercoid stage in the first seven of the species listed and, since development appeared normal in the other mites, it is possible that they might have completed their development if the hosts had lived long enough. Development of C. denticulata was observed in S. minutus, X. tegeocranus, C. cepheiformis, T. incisellus, S. laevigatus, and L. coracinus, although only S. minutus lived long enough in the cultures for cysticercoids to be produced. Infections were obtained only in oribatid mites. None of the rabbits fed cysticercoids from experimentally infected mites gave evidence of infection and it is probable that these larvae were not entirely mature. The developmental stages of C. ctenoides and C. denticulata in the intermediate hosts are described from experimental infections. The development of C. ctenoides, C. denticulata and C. pectinata in the final hosts is described from natural infections. No evidence was found to support the idea that these tapeworms migrate from the intestine to the body cavity of normal, live, rabbits.

The Morphology and Life History of the Cestode, Bertiella Studeri

Summary The problem of specific identity in the genus Bertiella is discussed. Three mature cestodes from a monkey, Macacus rhesus, are identified tentatively as Bertiella studeri. The animal probably acquired the infection in India, but had been in Hamburg for at least six months. The worms and eggs are described and life history experiments are reported. Eggs from gravid proglottids were fed to twenty-four species of free-living mites, collected from different locations in the vicinity of Hamburg. Onchospheres and developing larvae were recovered from the body cavity of Notaspis coleoptratus, Scutovertex minutus, Scheloribates laevigatus and Galumna sp. Cysticercoids were removed from S. laevigatus and Galumna sp. They were spherical, to oval, to pyriform, 0.1 to 0.15 mm. in diameter, and provided with small cercomeres. It is clear therefore that Bertiella, like other anoplocephaline cestodes, Moniezia and Cittotaenia, utilizes free-living mites as intermediate hosts.

The development ofMoniezia expansain the intermediate host

When eggs ofMonieza expansaare fed to galumnid mites, the onchospheres emerge in the intestine and migrate to the body cavity. Here they undergo metamorphosis and develop into cysticercoids. The dissection of mites, at various intervals after exposure, has yielded a successive series of developmental stages. Protocols of the experiments and photographs of living larvae are presented. These results demonstrate that mites serve as intermediate hosts ofMoniezia, and probably also of other anoplocephaline cestodes.