Beta diversity of gastrointestinal helminths in two closely related South African rodents: species and site contributions.

A fundamental aim of parasite ecology is to understand the mechanisms behind spatial variation in diversity and structure of parasite assemblages. To understand the contribution of individual parasite species and their assemblages to spatial variation in parasite communities, we examined species contributions to beta diversity (SCBD) and local contributions to beta diversity (LCBD) of parasitic gastrointestinal helminths (nematodes and cestodes) in two closely related rodents, Rhabdomys dilectus and Rhabdomys pumilio, from 20 localities across South Africa. Although the two Rhabdomys spp. are morphologically similar, they differ substantially in body size, habitat preference, and sociality. We asked whether the variation in life history traits and infection parameters are associated with SCBD of helminths and whether variation in environmental factors, host population density, and species richness of host communities are associated with LCBD of component assemblages of helminths. We also considered spatial factors to test whether LCBD of helminth assemblages demonstrate geographic structure. We found that the contribution of helminth species parasitic in both hosts to beta diversity significantly increased with characteristic prevalence of these species, whereas mean abundance, type of life cycle, and location in the host's gut had no effect on SCBD. The LCBD of helminth assemblages showed a significant positive correlation with environmental factors in both host species. Our results suggest that predictors of variation in SCBD and LCBD may substantially differ between parasites with different infection parameters and/or parasite communities at different hierarchical scales.

Intra- and interspecific similarity in species composition of helminth communities in two closely-related rodents from South Africa.

To reveal factors responsible for spatial variation in parasite community composition we studied patterns of similarity in helminth species composition in two closely-related rodents (Rhabdomys pumilio and Rhabdomys dilectus) that differ in their social and spatial behaviour and live under different environmental conditions across 20 localities in South Africa. We asked whether the two hosts harbour similar assemblages, whether these are more dissimilar between than within hosts and if host social structure, behaviour or environment affects similarity patterns in helminth infracommunities within and among localities. We also investigated whether similarity in species composition of helminth component communities decreases with an increase of geographic distance between host populations. We found that the pattern of space use by the hosts rather than their social behaviour promotes differences in helminth species composition between host species as well as among host populations from different localities. The rate of distance decay of similarity in species composition of helminth component communities differed between the two hosts due to difference in the degree of environmental variation across their geographic ranges. We conclude that patterns of spatial variation in helminth species composition are driven mainly by host spatial behaviour and, to a lesser extent, by environment-associated factors.

Community structure of helminth parasites in two closely related South African rodents differing in sociality and spatial behaviour.

To understand the effect of social and spatial behaviour of a host on parasite community organization, we studied species co-occurrence and nestedness of assemblages of gastrointestinal helminths in two closely related rodents, solitary and mobile Rhabdomys dilectus and social and territorially conservative Rhabdomys pumilio, and asked whether helminth communities of the two hosts are characterized by a non-random pattern and whether the occurrence or degree of this non-randomness (a) differs between hosts and (b) is associated with abundance, prevalence and diversity of helminths. We found that although the general pattern of helminth co-occurrence was similar in the two hosts, helminth infracommunities of R. dilectus and R. pumilio differed in the relative frequency of positive and negative pairwise species co-occurrences (only positive in the former and both positive and negative in the latter). Nestedness-related patterns in helminth infracommunities were found in R. pumilio (predominantly anti-nested) but not R. dilectus (predominantly non-nested), whereas the opposite was the case for their component communities (non-nested versus nested, respectively). The level of infection was generally associated with the manifestation of non-randomness in helminth assemblages. Different infection parameters affected different structure patterns in the two hosts. We concluded that community structure of helminths in Rhabdomys spp. results from complex interactions between parasite- and host-associated factors.

Helminth parasitism in two closely related South African rodents: abundance, prevalence, species richness and impinging factors.

We investigated patterns of helminth infection in two closely related rodents (social Rhabdomys pumilio occurring mainly in xeric habitats and solitary R. dilectus occurring mainly in mesic habitats) at 20 localities in different biomes of South Africa and asked if between-species differences were mainly caused by difference in sociality or difference in environmental conditions of their respective habitats. Helminths recovered from the gastrointestinal tract totalled 11 nematode and 5 cestode species from R. pumilio and 19 nematode and 7 cestode species from R. dilectus. In both hosts, mean abundance and prevalence of nematodes were higher compared to cestodes. Cestode infection as well as nematode abundance, species richness or prevalence did not differ between the two rodents. However, incidence of nematode infection was significantly higher in R. dilectus than in R. pumilio. Moreover, nematode numbers and species richness in infracommunities of R. pumilio inhabiting the relatively more xeric Karoo biome were significantly lower than in those inhabiting the relatively less xeric Fynbos biome. Although we could not unequivocally distinguish between effects of host sociality and environmental factors on the number of individuals and species of helminths in the two hosts, differences in the incidence of nematode infection between R. pumilio and R. dilectus as well as differences in the number of nematode individuals and species between R. pumilio from the Fynbos and the Karoo suggested the effect of environmental conditions on helminth infection to be more important than that of sociality.

Phylogenetic relationships and systematics of tapeworms of the family Davaineidae (Cestoda, Cyclophyllidea), with emphasis on species in rodents.

The present study aims at clarifying the poorly known phylogenetic relationships and systematics of cestodes of the family Davaineidae Braun, 1900 (Cyclophyllidea), primarily the genus Raillietina Fuhrmann, 1920 and of the subfamily Inermicapsiferinae (Anoplocephalidae) from mammals (mostly rodents, 31 new isolates) and birds (eight new isolates). Phylogenetic analyses are based on sequences of the large subunit ribosomal RNA gene (28S) and mitochondrial NADH dehydrogenase subunit 1 gene (nad1). The main phylogenetic pattern emerging from the present analysis is the presence of three independent lineages within the main clade of the subfamily Davaineinae, one of which is almost entirely confined to species from rodents and the other two show a mixture of species from birds and mammals. It is suggested that the major diversification of the main clade took place in birds, possibly in galliforms. The subsequent diversification included repeated host shifts from birds to mammals and to other birds, and from rodents to other mammals, showing that colonisation of new host lineages has been the main driver in the diversification of davaineine cestodes. It is also shown that all isolates of Inermicapsifer Janicki, 1910, mainly from rodents, form a monophyletic group positioned among Raillietina spp. in the "rodent lineage", indicating that the genus Inermicapsifer is a member of the family Davaineidae. This means that the subfamily Inermicapsiferinae and the family Inermicapsiferidae should be treated as synonyms of the Davaineidae, specifically the subfamily Davaineinae. Three additional genera generally included in the Inermicapsiferinae, i.e. Metacapsifer Spasskii, 1951, Pericapsifer Spasskii, 1951 and Thysanotaenia Beddard, 1911, are also assigned here to the Davaineidae (subfamily Davaineinae). Raillietina spp. were present in all three main lineages and appeared as multiple independent sublineages from bird and mammalian hosts, verifying the non-monophyly of the genus Raillietina and suggesting a presence of multiple new species and genera.

Afrojoyeuxia gen. n. and Hunkeleriella gen. n., two new genera of cestodes (Cyclophyllidea: Anoplocephalidae) from African rodents.

Based on the study of type material, two new genera of cestodes (Cyclophyllidea: Anoplocephalidae) are proposed for Paranoplocephala Lüihe, 1910 sensu lato species from African rodents. Afrojoyeuxia gen. n., proposed for A. gundii (Joyeux, 1923) comb. n. from Ctenodactylus gundi (Rothmann) (Hystricomorpha: Ctenodactylidae), is characterized by a high length/width ratio of mature proglottids, longitudinally extensive testicular field positioned anterior to the female glands, an ovoid or subspherical cirrus-sac and a thick, conical cirrus. Hunkeleriella gen. n., proposed for H. dasymidis (Hunkeler, 1972) comb. n. from Dasymys incomtus (Sundevall) (Myomorpha: Muridae), differs from related genera mainly by its short (10-20 mm) and wide strobila and neck, unilateral genital pores (exceptionally with a few changes per strobila), the position of the genital pores (slightly anterior to the middle of proglottid margin) and initially tube-like early uterus (later reticulated). Parandrya Gulyaev et Chechulin, 1996, earlier suggested to be a junior synonym of Paranoplocephala, is considered to be a valid, independent genus. Evidence of non-monophyly and need for a taxonomic revision of Paranoplocephala sensu lato, as well as the phylogenetic position of A. gundii and H. dasymidis are discussed.

Andryoides gen. n. and morphological key features in cestodes of the family Anoplocephalidae sensu stricto (Cyclophyllidea) in mammals.

As presently defined, the tapeworm genus Andrya Railliet, 1895 (Cyclophyllidea: Anoplocephalidae sensu stricto) includes the type species A. rhopalocephala (Riehm, 1881) in hares of the genus Lepus Linnaeus (Leporidae) in western Eurasia and four species in cricetid (Neotominae, Sigmodontinae) and octodontid rodents in North and South America. The host range of Andrya is puzzling, because it is the only genus of anoplocephalid (s. s.) cestodes parasitising both rodents and lagomorphs. The present morphological analysis shows that the American species of Andrya share multiple consistent features, in which they differ from those of A. rhopalocephala and the morphologically related Neandrya cuniculi (Blanchard, 1891). The main differences concern the position of the uterus with respect to the longitudinal osmoregulatory canals and testes. Consequently, a new genus Andryoides gen. n. is proposed for the American species, resulting in the following combinations: Andryoides neotomae (Voge, 1946) comb. n. (type species), Andryoides octodonensis (Babero et Cattan, 1975) comb. n., Andryoides vesicula (Haverkost et Gardner, 2010) comb. n. and Andryoides boliviensis (Haverkost et Gardner, 2010) comb. n. However, A. boliviensis is regarded here as a junior synonym of A. vesicula (new synonymy). The present study also defines the morphological key features for all the valid genera of cestodes of the family Anoplocephalidae (s. s.), and discusses the phylogenetic affinities and historical biogeography of Andryoides and other endemic American anoplocephalid cestodes.

Identification of the tapeworm Mosgovoyia pectinata (Anoplocephalidae) in Faroese mountain hares (Lepus timidus).

The mountain hares (Lepus timidus L., 1758) in the Faroe Islands, an archipelago located in the North Atlantic, are known to be commonly infected by tapeworms, the identity of which was unknown. The mountain hare, which now populates 15 of the 18 islands, was introduced from Norway in 1855. In this study, tapeworms collected from four mountain hares from four geographic areas of the Faroe Islands were subjected to molecular identification using the nuclear ribosomal DNA (28S), the mitochondrial cytochrome oxidase subunit 1 (cox1) and the NADH dehydrogenase subunit 1 (nad1) genes. The results indicate unambiguously that the tapeworms were Mosgovoyia pectinata (Goeze, 1782) (Cestoda: Anoplocephalidae sensu stricto). The phylogenetic position and origin of the Faroese M. pectinata are discussed. Given that the parasite is quite common in Norway, from where the mountain hares were introduced, it is conceivable that co-introduction of M. pectinata from Norway to the Faroe Islands took place. The phylogenetic analyses revealed high similarity of the M. pectinata sequences from three regions and the position of the Faroese isolate as the sister lineage of the isolates from Finland and East Siberia.

Taenia arctos n. sp. (Cestoda: Cyclophyllidea: Taeniidae) from its definitive (brown bear Ursus arctos Linnaeus) and intermediate (moose/elk Alces spp.) hosts.

Taenia arctos n. sp. (Cestoda: Cyclophyllidea: Taeniidae) is described from the brown bear Ursus arctos Linnaeus (definitive host) and moose/elk Alces spp. (intermediate hosts) from Finland (type-locality) and Alaska, USA. The independent status of the new species and the conspecificity of its adults and metacestodes have been recently confirmed by the mtDNA sequence data of Lavikainen et al. (2011; Parasitology International, 60, 289-295). Special reference is given to morphological differences between the new species and T. krabbei Moniez, 1879 (definitive hosts primarily canines for the latter), both of which use the moose/elk (Alces spp.) as intermediate hosts (the latter also uses Rangifer and perhaps other northern ruminants), and between the new species and T. ursina Linstow, 1893, both of which use the brown bear U. arctos as a definitive host. New morphological data are also provided for adults and cysticerci of T. krabbei. The analysis includes potentially useful morphometric features that have not been previously applied to Taenia spp.

Are gastrointestinal parasites associated with the cyclic population dynamics of their arctic lemming hosts?

Many rodents, including most populations of arctic lemmings (genus Dicrostonyx and Lemmus), have cyclic population dynamics. Among the numerous hypotheses which have been proposed and tested to explain this typical characteristic of some terrestrial vertebrate communities, trophic interactions have often been presented as the most likely drivers of these periodic fluctuations. The possible role of parasites has, however, only seldom been assessed. In this study, we genetically measured the prevalence of two endoparasite taxa, eimerians and cestodes, in 372 faecal samples from collared lemmings, over a five year period and across three distant sites in Northeast Greenland. Prevalence of cestodes was low (2.7% over all sites and years) and this taxon was only found at one site (although in 4 out of 5 years) in adult hosts. By contrast, we found high prevalence for eimerians (77.7% over all sites and years), which occurred at all sites, in every year, for both age classes (at the Hochstetter Forland site where both adult and juvenile faeces were collected) and regardless of reproductive and social status inferred from the characteristics of the lemming nests where the samples had been collected. Prevalence of eimerians significantly varied among years (not among sites) and was higher for juvenile than for adult lemmings at the Hochstetter Forland site. However, higher prevalence of eimerians (P t ) was only associated with lower lemming density (N t ) at one of the three sites and we found no delayed density dependence between N t and P t+1 to support the parasite hypothesis. Our results show that there is no clear relation between lemming density and eimerian faecal prevalence in Northeast Greenland and hence no evidence that eimerians could be driving the cyclic population dynamics of collared lemmings in this region.

First Report of Gastrointestinal Parasites from Ansell's Mole-Rat ( Fukomys anselli) in Zambia.

The endoparasite fauna of Ansell's mole-rat, Fukomys anselli (Burda, Zima, Scharff, Macholán and Kawalika) (Rodentia, Bathyergidae), was studied for the first time near Lusaka, Zambia. We recovered 7 endoparasite taxa including 5 nematodes ( Hexametra sp.; Protospirura numidica Seurat, 1914; Protospirura muricola Gedoelst, 1916; Protospirura sp.; and Mammalakis zambiensis Junker, Lutermann and Mutafchiev, 2017 ); and 2 cestodes ( Inermicapsifer sp. and Rodentolepis cf. microstoma Dujardin, 1845) from 272 individuals sampled between February 2009 and February 2010. Species richness (0-2), prevalence (14.7%), and abundance (0.44 ± 0.10) were low, probably as a result of the subterranean lifestyle of the study species. Although effect of host sex on helminth burden was largely absent, helminth burden varied with season and breeding status of the host. This may be a result of seasonal variation in burrow size in response to rainfall, but also the division of labor present in the study species.

Molecular taxonomy and subgeneric classification of tapeworms of the genus Moniezia Blanchard, 1891 (Cestoda, Anoplocephalidae) in northern cervids (Alces and Rangifer).

Phylogenetic relationships of tapeworms of the genus Moniezia Blanchard, 1891 (Cestoda, Anoplocephalidae) parasitizing the Eurasian elk Alces alces, the moose A. americanus and the reindeer/caribou Rangifer tarandus (Cervidae) were studied using DNA sequences of two mitochondrial genes (cox1 and nad1). Several isolates from domestic ruminants, representing Moniezia expansa (Rudolphi, 1810) sensu lato and M. benedeni (Moniez, 1879) sensu lato, and one unidentified isolate from an African antelope, were also included in the analysis. Both genes identified the same six species of Moniezia, but interspecific phylogenetic relationships were better resolved by the nad1 data. The six species of Moniezia comprised two main clades: clade 1 that originates in bovids, with subsequent colonization of northern cervids in Eurasia, and clade 2 that originates in northern cervids, with subsequent specific divergence within these hosts. Clade 2 has a Holarctic distribution. None of the Moniezia specimens in Alces and Rangifer was conspecific with the species in domestic ruminants, suggesting that the custom of identifying Moniezia spp. in northern cervids either as M. expansa or M. benedeni is incorrect. At least two of the species parasitizing Alces and Rangifer have not been previously recognized. These findings challenge the results of all previous studies concerning the diversity and ecology of Moniezia spp. in northern cervids. The traditional classification into three subgenera (Moniezia Blanchard, 1891, Blanchariezia Skrjabin & Schultz, 1937 and Baeriezia Skrjabin & Schultz, 1937), based on the presence and type of interproglottidal glands, conflicts with the currently observed molecular phylogenetic relationships within the genus Moniezia.

Description and life-cycle of Taenia lynciscapreoli sp. n. (Cestoda, Cyclophyllidea).

A new species of tapeworm, Taenia lynciscapreoli sp. n. (Cestoda, Cyclophyllidea), is described from the Eurasian lynx (Lynx lynx), the main definitive host, and the roe deer (Capreolus capreolus and Capreolus pygargus), the main intermediate hosts, from Finland and Russia (Siberia and the Russian Far East). The new species was found once also in the wolf (Canis lupus) and the Eurasian elk/moose (Alces alces), representing accidental definitive and intermediate hosts, respectively. The conspecificity of adult specimens and metacestodes of Taenia lynciscapreoli sp. n. in various host species and regions, and their distinction from related species of Taenia, was confirmed by partial nucleotide sequences of the mitochondrial cytochrome c oxidase subunit 1 gene. Morphologically, Taenia lynciscapreoli sp. n. can be separated unambiguously from all other species of Taenia by the shape of its large rostellar hooks, particularly the characteristically short, wide and strongly curved blade. If the large rostellar hooks are missing, Taenia lynciscapreoli may be separated from related species by a combination of morphological features of mature proglottids. It is suggested that Taenia lynciscapreoli has been present in published materials concerning the tapeworms of Lynx lynx and Lynx pardinus in Europe, but has been misidentified as Taenia pisiformis (Bloch, 1780). Taenia lynciscapreoli sp. n. has not been found in lynx outside the range of roe deer, suggesting a transmission pathway based on a specific predator-prey relationship. The present study applies a novel, simple approach to compare qualitative interspecific differences in the shape of rostellar hooks.

Cryptic diversity in hymenolepidid tapeworms infecting humans.

An adult hymenolepidid tapeworm was recovered from a 52-year-old Tibetan woman during a routine epidemiological survey for human taeniasis/cysticercosis in Sichuan, China. Phylogenetic analyses based on sequences of nuclear 28S ribosomal DNA and mitochondrial cytochrome c oxidase subunit 1 showed that the human isolate is distinct from Hymenolepis diminuta and Hymenolepis nana, the common parasites causing human hymenolepiasis. Proglottids of the human isolate were unfortunately unsuitable for morphological identification. However, the resultant phylogeny demonstrated the human isolate to be a sister species to Hymenolepis hibernia from Apodemus mice in Eurasia. The present data clearly indicate that hymenolepidid tapeworms causing human infections are not restricted to only H. diminuta and H. nana.

Parasites and invasions: changes in gastrointestinal helminth assemblages in invasive and native rodents in Senegal.

Understanding why some exotic species become widespread and abundant in their colonised range is a fundamental issue that still needs to be addressed. Among many hypotheses, newly established host populations may benefit from a parasite loss ("enemy release" hypothesis) through impoverishment of their original parasite communities or reduced infection levels. Moreover, the fitness of competing native hosts may be negatively affected by the acquisition of exotic taxa from invaders ("parasite spillover") and/or by an increased transmission risk of native parasites due to their amplification by invaders ("parasite spillback"). We focused on gastrointestinal helminth communities to determine whether these predictions could explain the ongoing invasion success of the commensal house mouse (Mus musculus domesticus) and black rat (Rattus rattus), as well as the associated decrease in native Mastomys spp., in Senegal. For both invasive species, our results were consistent with the predictions of the enemy release hypothesis. A decrease in overall gastrointestinal helminth prevalence and infracommunity species richness was observed along the invasion gradients as well as lower specific prevalence/abundance (Aspiculuris tetraptera in Mus musculus domesticus, Hymenolepis diminuta in Rattus rattus) on the invasion fronts. Conversely, we did not find strong evidence of GIH spillover or spillback in invasion fronts, where native and invasive rodents co-occurred. Further experimental research is needed to determine whether and how the loss of gastrointestinal helminths and reduced infection levels along invasion routes may result in any advantageous effects on invader fitness and competitive advantage.

Reappraisal of Hydatigera taeniaeformis (Batsch, 1786) (Cestoda: Taeniidae) sensu lato with description of Hydatigera kamiyai n. sp.

The common cat tapeworm Hydatigera taeniaeformis is a complex of three morphologically cryptic entities, which can be differentiated genetically. To clarify the biogeography and the host spectrum of the cryptic lineages, 150 specimens of H. taeniaeformis in various definitive and intermediate hosts from Eurasia, Africa and Australia were identified with DNA barcoding using partial mitochondrial cytochrome c oxidase subunit 1 gene sequences and compared with previously published data. Additional phylogenetic analyses of selected isolates were performed using nuclear DNA and mitochondrial genome sequences. Based on molecular data and morphological analysis, Hydatigera kamiyai n. sp. Iwaki is proposed for a cryptic lineage, which is predominantly northern Eurasian and uses mainly arvicoline rodents (voles) and mice of the genus Apodemus as intermediate hosts. Hydatigera taeniaeformis sensu stricto (s.s.) is restricted to murine rodents (rats and mice) as intermediate hosts. It probably originates from Asia but has spread worldwide. Despite remarkable genetic divergence between H. taeniaeformis s.s. and H. kamiyai, interspecific morphological differences are evident only in dimensions of rostellar hooks. The third cryptic lineage is closely related to H. kamiyai, but its taxonomic status remains unresolved due to limited morphological, molecular, biogeographical and ecological data. This Hydatigera sp. is confined to the Mediterranean and its intermediate hosts are unknown. Further studies are needed to classify Hydatigera sp. either as a distinct species or a variant of H. kamiyai. According to previously published limited data, all three entities occur in the Americas, probably due to human-mediated introductions.

Morphological characterisation of Paranoplocephala bairdi (Schad, 1954) (Cestoda: Anoplocephalidae) in heather voles Phenacomys spp. and tree voles Arborimus spp., and related species in voles and lemmings (Muridae: Arvicolinae).

The taxonomical status of Paranoplocephala bairdi (Schad, 1954)-like cestodes (Anoplocephalidae) in heather voles Phenacomys spp. and tree voles Arborimus spp. (Muridae: Arvicolinae) and their discrimination from five related species of Paranoplocephala is assessed using uni- and multivariate morphometrics. The analyses support the independent status and conspecificity of specimens from Phenacomys spp. and Arborimus spp., and P. bairdi is therefore suggested to be a host-specialist species of heather and tree voles with a wide geographical distribution in North America. A redescription is presented for P. bairdi.

Checklist of tapeworms (Platyhelminthes, Cestoda) of vertebrates in Finland.

A checklist of tapeworms (Cestoda) of vertebrates (fishes, birds and mammals) in Finland is presented, based on published observations, specimens deposited in the collections of the Finnish Museum of Natural History (Helsinki) and the Zoological Museum of the University of Turku, and additional specimens identified by the present author. The checklist includes 170 tapeworm species from 151 host species, comprising 447 parasite species/host species combinations. Thirty of the tapeworm species and 96 of the parasite/host species combinations have not been previously reported from Finland. The total number of tapeworm species in Finland (170 spp.) is significantly lower than the corresponding figure for the Iberian Peninsula (257 spp.), Slovakia (225 spp.) and Poland (279 spp.). The difference between Finland and the other three regions is particularly pronounced for anseriform, podicipediform, charadriiform and passeriform birds, reflecting inadequate and/or biased sampling of these birds in Finland. It is predicted that there are actually ca. 270 species of tapeworms in Finland, assuming that true number of bird tapeworms in Finland corresponds to that in other European countries with more comprehensive knowledge of the local tapeworm fauna. The other main pattern emerging from the present data is the seemingly unexplained absence in (northern) Fennoscandia of several mammalian tapeworms that otherwise have extensive distributions in the Holarctic region or in Eurasia, including the northern regions. Previously unknown type specimens, that is, the holotype of Bothrimonus nylandicus Schneider, 1902 (a junior synonym of Diplocotyle olrikii Krabbe, 1874) (MZH 127096) and the syntypes of Caryophyllaeides fennica (Schneider, 1902) (MZH 127097) were located in the collections of the Finnish Museum of Natural History.

The impact of climatic factors and host density on the long-term population dynamics of vole helminths.

The seasonal and long-term population dynamics of helminths parasitizing voles suggested that density-dependent factors might be important in the population dynamics of common species, whereas density-independent factors predominate in the regulation of the rare species. To test this, we used single and multiple regression to analyse the effects of climatic factors and host density on populations of six species of vole helminths over 12 years. The data do support the idea of a difference between common and rare species of helminths, but they clearly do not support the above hypothesis. The common helminths Heligmosomum mixtum (Nematoda) and Catenotaenia sp. (Cestoda) responded to changes in temperature sum (>5° C days) and precipitation during summer. The combined effect of climatic factors and host density explained most of the variation in the long-term dynamics of these common species. By contrast, the long-term dynamics of the rare helminths Paranoplocephala kalelai (Cestoda), Mastophorus muris, Capillaria sp. and Syphacia petrusewiczi (Nematoda) were explained less well by weather and host density than those of the common ones. Furthermore, the common and rare helminths differed in some ways in their responses to climatic factors.

Distribution patterns and microhabitat segregation in gastrointestinal helminths of Sorex shrews.

We studied the distribution patterns and microhabitat use in gastrointestinal helminths of the shrews Sorex araneus and S. caecutiens in Finland. The distribution of species prevalences was bimodal, and in S. araneus the abundance (mean intensity) was positively associated with commonness (prevalence), as assumed by the core-satellite species hypothesis (Hanski 1982). However, the positive correlation between prevalence and intensity was observed only when the effects of helminth body size and taxonomic group (cestodes vs nematodes) on intensity were controlled for. The nematodes of the genus Longistriata occurred predictably as core species, whereas the identity of the core cestodes was more variable between host species and regions. Helminth body size and taxonomic group were not related to the degree of aggregation in shrew populations, but helminth body size seemed to explain the differences in the distribution patterns of helminths between shrews and voles. The core species did not show more segregation in microhabitat use than randomly selected species. In fact, the two core nematodes showed largely overlapping intestinal distributions. We conclude that linear intestinal space is not a key resource for shrew nematodes, but it may be for shrew cestodes.