Studies on the helminth parasites of voles and shrews from Wales

The incidence and degree of infestation of the Bank vole, Clethrionomys glareolus , the Skomer vole, C. skomerensis , the Common shrew, Sorex araneus and the Pygmy shrew, S. minutus have been studied from a rough grassland and woodland area, Aberystwyth and from Skomer Island in Wales. In addition, small numbers of the Short-tailed vole, Microtus agrestis and the Water shrew, Neomys fodiens were examined for helminth parasites, when they became available. Seven new host records and six new British records are listed. The number of species of helminths in shrews, which consisted largely of digeneans and cestodes, was greater than that in voles. This is undoubtedly linked with differences in the feeding habits of the two hosts.
In Aberystwyth, where the composition of the helminth fauna was found to be more varied than that from Skomer Island, one species of nematode showed evidence of seasonal variation in the degree of infestation of Clethrionomys glareolus and four species, one digenean, two cestodes and one acantocephalan in Sorex araneus. The factors affecting this seasonal fluctuation in parasite numbers are discussed.     

The occurrence of the Pigmy shrew Sorex minutus on moorland, and the implications for its presence in Ireland

Throughout most of its range in western Europe, the Pigmy shrew Sorex minutus is sympatricwith the Common shrew Sorex araneus but greatly outnumbered by it. This makes it difficult to explain the presence of the Pigmy shrew, in the absence of its congener, in Ireland. A limited pitfall trapping exercise in England has shown that in moorland on deep peat, the Pigmy shrew may in fact outnumber the Common shrew. Reference to the literature on the ecology of the two shrews shows that this result is readily explained by existing knowledge of their diet; earthworms are a major prey for Common shrews and are absent from waterlogged, acidic, peats. Further, evidence on the nature of the possible land bridge from Scotland to Ireland suggests that it was at best low-lying, and probably partly flooded. By analogy with moorland, such a land bridge might have been suitable for Pigmy shrews but not for Common shrews.     

Morphometric distances and population structuring in the common shrew <Emphasis Type="Italic">Sorex araneus</Emphasis> L. (Lipotyphla: Soricidae)

The skulls of shrews of genus Sorex from eight samplings from the European part of Russia and two from the vicinity of Novosibirsk were compared. The characteristics were identified using 22 marks on the axial skull. It was found that the centroid size differs significantly in the common and Laxmann’s shrews S. caecutiens and the pygmy shrew S. minutus, while for selected marks, the common and Laxmann’s, as well as pygmy, shrews were significantly different in form, but the differences were very small between Laxmann’s and pygmy shrews. The characteristic features of the biology of the shrews Sorex are discussed, which may contribute to understanding the general laws of the morphological evolution of the genus.     

Density compensation in allopatric populations of the pygmy shrewSorex minutus on Gotland and the outer hebrides: evidence for the effect of interspecific competition

Pygmy shrewsSorex minutus occur allopatrically in Ireland, the Outer Hebrides and Gotland in Sweden, and sympatrically with common shrewsS. minutus on the European mainland and in England. Competition theory redicts higher population density in allopatric populations as a consequence of relaxed competition. Here this prediction is tested by comparing allopatric populations of pygmy shrews in the Outer Hebrides and in Gotland with sympatric populations on the Swedish mainland. Population densities were higher in allopatry than in sympatry. Lower summed densities in allopatry, which is predicted by niche theory, was found on the Outer Hebrides, but not on Gotland. Size distributions of carabids, which are important food items, could not explain the differences in population densities. Since both avian predators and snakes are present in all areas, population density of pygmy shrews during autumn in concluded to be regulated primarly by competition with common shrews in areas of sympatry.     

How did pygmy shrews colonize Ireland? Clues from a phylogenetic analysis of mitochondrial cytochrome b sequences.

There is a long-standing debate as to how Ireland attained its present fauna; we help to inform this debate with a molecular study of one species. A 1110 base pair fragment of the mitochondrial cytochrome b gene was sequenced in 74 specimens of the pygmy shrew, Sorex minutus, collected from throughout its western Palaearctic range. Phylogenetic analysis of these sequences revealed several well-supported lineages. Most of the 65 haplotypes belonged to a northern lineage, which ranged from Britain in the west to Lake Baikal in the east. The other lineages were largely limited to Iberia, Italy and the Balkans. One exception, however, was a lineage found in both Ireland and Andorra. This affinity, and the large difference between the mitochondrial sequences of Irish and British individuals, suggest that pygmy shrews did not colonize Ireland via a land connection from Britain, as has been previously supposed, but instead were introduced by boat from southwest continental Europe. All the Irish pygmy shrews analysed were identical or very similar in cytochrome b sequence, suggesting an extreme founding event.     

Species composition, host association and niche differentiation in fleas of small mammals in the Ilmen-Volkhov lowland

Species composition, abundance, annual cycles, and host association of fleas parasitizing small mammals were investigated. The problem of niche differentiation in these insects is considered on the base of the comparative analysis of their annual cycles. The annual cycles of the fleas are revealed to be similar in the case of few number of flea species in parasite community. Thus, two species parasitizing Sorex araneus (Doratopsylla dasycnema and Palaeopsylla soricis), as well as three species associated with Apodemus uralensis (Megabothris turbidus, Ctenophthalmus agyrtes, and Ct. uncinatus) have equal phenology of parasitizing. The fleas community of Clethrionomys glareolus is characterized by a large species number and high diversity of the annual cycles. The differentiation by the seasons of parasitizing is observed most clearly in the dominant flea species, namely Amalaraeus penicilliger, Ct. uncinatus, and Peromyscopsylla bidentata. The periods of imaginal life are overlapped significanly in these species, but they are differed by the season of dominance. Ct. uncinatus predominates in spring and summer, while P. bidentata predominates in autumn, and A. penicilliger predominates in winter and early spring. It may be noted also, that niche partitioning was not observed in the fleas having wide range of hosts. The imaginal life of such fleas usually does not go beyond the warm season.     

Small mammal populations in relation to hedgerow structure in an arable landscape

The population ecology of small mammals in hedgerows in arable farmland in eastern England is described. Features of hedgerows of importance to individual species are examined. Some 97% of the total 3042 mammals captured were wood mouse Apodemus sylvaticus , yellow-necked mouse Apodemus flavicollis , bank vole Clethrionomys glareolus and common shrew Sorex araneus . Small numbers of harvest mice Micromys minutus , field voles Microtus agrestis , pygmy shrews Sorex minutus and water shrews Neomys fodiens were also caught. Wood mouse, the most numerous species, showed a typical pattern of large numbers in autumn and winter, followed by a simultaneous decline over all hedges in early spring. Population changes were less clear in yellow-necked mouse and bank vole but the yellow-necked mouse was more scarce in the second year of study. Common shrews were most numerous in summer and declined rapidly in autumn. Hedgerow coppicing had a marked effect on yellow-necked mouse numbers but not on wood mouse. In an extensive survey of mammal numbers in relation to hedgerow features, ground cover was found to be the single largest factor influencing size of bank vole populations. Hedgerow condition (lack of gaps) was important to yellow-necked mice, which thrived only in well-established hedgerows. Wood mice appeared little influenced by the characteristics of the hedge. Common shrews were more abundant in hedgerows with adjacent permanent water.     

Seasonal dynamics of a flea number (Siphonaptera) on the common shrew (Sorex araneus) in the north part of the Novgorod oblast

In the course of 5-year stationary investigations of the common shrew Sorex araneus in the north part of the Novgorod oblast, 12 flea species have been recorded on this host. Among them, Palaeopsylla soricis and Doratopsilla dasycnema are specific parasites of small insectivores, including the common shrew. Hystrichopsylla talpae is a polyxenous species, parasitizing both rodents and insectivores. Other 9 flea species are not common parasites of S. araneus and apparently have come to this host from other mammal species inhabiting the forest biotopes in the area of investigation. P. soricis and D. dasycnema have similar phenology of parasitism. These fleas appear in spring (April), are present during summer and autumn and disappear in winter. During a year, the abundance of these species shows three pikes, which correspond to three generations: spring (April), summer (June-July) and autumn (September-October) generation. The first species dominates in spring (April-May) and in the autumn-winter period, while the second species predominates in summer. These two species comprise over 90% of total number of fleas collected and determine general dynamics of the flea number on the shrews; during the year the flea number has a sinusoidal increasing from spring to autumn and minimum in winter.     

An investigation of the population ecology of syntopic shrews inhabiting water-cress beds

The population ecology of the shrews Neomys fodiens, Sorex araneus and S. minutus living syntopically in an area of commercial water-cress beds was investigated by live-trapping over a two-year period. The shrew population comprised 52% 5. araneus , 31% N. fodiens and 17% S. minutus. Sorex araneus was the most numerous species in all seasons. Peak numbers of shrews occurred in summer with low numbers in the ensuing winter. Input of new individuals was high, especially of N. fodiens which exceeded 60% in most months. Survival in the study area never exceeded 8 months in N. fodiens and 11 months in S. araneus. Both species suffered approximately 50% mortality and emigration in the first two months of life. Apart from a few, more nomadic, shrews, N. fodiens and S. araneus seemed quite sedentary during trapping periods, moving a mean distance of 13.7 m and 17.2 m between successive captures, respectively. The relatively low numbers of recaptures of N. fodiens at successive trapping periods indicated nomadic behaviour. Both species were more active during darkness than during daylight, and were more active in spring and summer than in winter. High turnover of the shrew population was attributed to mortality and emigration. Emigration was possibly encouraged by competition for space, particularly for nesting sites.     

Habitat occurrence and prey distribution of a multi-species community of shrews in the Siberian taiga

The habitat occurrence and invertebrate prey distribution of nine species of shrew in the mid-taiga of central Siberia were investigated. Species richness ranged from 4–9 shrews per habitat. Sorex araneus and S. caecutiens were numerically dominant in all seven habitats (44 and 36% of the total catch, respectively) while Sorex minutus, S. tundrensis, S. isodon , and S. roboratus each constituted 4–6% and Sorex minutissimus, S. daphaenodon , and Neomys fodiens were rare (< 1% each). There was no overall correlation between abundance of shrews and invertebrate prey, but flood-plain habitats supported the greatest abundance and species richness of shrews, and high density and biomass of prey. Oligochaete-eating shrews were twice as numerous here as in other habitats, coincident with high abundance of oligochaetes. The large, earthworm-feeding Sorex roboratus occurred only here. The more acid, typical taiga habitats had lower adundance and species richness of shrews. They had the lowest density and biomass of prey, particularly oligochaetes, and far fewer oligochaete-eating shrews. The relative paucity of shrews in bush-meadow habitats, despite abundant prey, implied that habitat structure influences shrew distribution. Differential numbers of certain species in the presence or absence of larger congeners also suggested that interspecific competitive effects influence habitat selection by shrews. The high species richness of shrews here in the mid-taiga may be accounted for by the heterogeneous nature of the constituent habitats which provide niches for small and large species of shrew with a range of feeding habits.     

The occurrence of Centrorhynchus (Acanthocephala) in shrews (Sorex araneus and Sorex minutus) in the United Kingdom

Encysted acanthocephalans belonging to the genus Centrorhynchus were found in the body cavities of Sorex araneus (common shrew) and Sorex minutus (pygmy shrew) from Boxworth, Cambridgeshire, U.K. Fifty percent of the male S. araneus and 67% of the male S. minutus examined were found to be infected, with the mean intensity (+/-SD) being 54.3 +/- 91.3 and 14.7 +/- 18.4, respectively. The species of Centrorhynchus in the shrews may be Centrorhynchus aluconis, which is distributed widely in tawny owls, Strix aluco, in the United Kingdom. Shrews appear to serve as paratenic hosts for C. aluconis.     

Interspecific competition in the shrews Sorex araneus and Sorex minutus (Soricidae, Insectivora): a population study of the Irish pygmy shrew

Sorex araneus and S. minutus , which are likely to show a temporary vertical habitat segregation, are largely sympatric, except in Ireland, where S. minutus occurs alone. Two mechanisms for this segregation may be suggested: a direct interspecific interaction or an evolutionary fixed relationship.
Population densities and surface activity were measured, both in areas where the two species occur together (The Netherlands) and in areas in Ireland, in (order to investigate the possibility of a habitat shift and its influence on population density. No such effect was found in the present study.     

Water and fat contents of British shrews and their role in the seasonal changes in body weight

Water and fat contents of wild and captive Sorex araneus, S. minutus and Neomys fodiens were determined. The influence of sex, locality, body weight and age on water and fat contents of wild shrews are discussed. A seasonal cycle in both water and fat content of wild shrews is demonstrated. An investigation of captive shrews revealed fat contents to be considerably greater than in wild shrews. It is concluded that the seasonal differences in water and fat contents found in this study are of insufficient magnitude to account for the loss in body weight of wild shrews in winter or to be of any great survival value to them.     

The seasonal dynamics of fleas (Siphonaptera) on bank voles (Clethrionomys glareolus) in the north part of Novgorod region

The twelve flea species were revealed on bank voles. Only four of them (Amalaraeus penicillige, Ctenophthalmus uncinatus, Megabothris turbidus, Peromescopsylla bidentata) were abundant in some seasons of the year. Four other species (Ct. agyrtes, Hystrichopsylla talpae, P. silvatica, Rhadinopsylla integella) are the parasites of bank vole too but their numbers were always low in the study area. The four last species (Amphypsylla rossica, M. walkeri, Doratopsylla dasycnema, Palaeopsylla soricis) are not peculiar to bank vole. They occurs on it occasionally from other animals--the voules of Mucrotus and shrews. The most species diversity of fleas on bank vole was observed at the period from August to October, the least one--in late winter, spring and early summer. The total abundance indices of fleas on the voles (mean number of the insects per host) ranged in different months from 0.17 to 5.65. The time of minimum flea numbers was August-September. The peak abundance was reached in springtime (March-April).     

Genetic variation in Irish pygmy shrews Sorex minutus (Soricomorpha: Soricidae): implications for colonization history

The status of the pygmy shrew ( Sorex minutus L.) as a native or an introduced species in Ireland has been subject to much debate. To examine this and other aspects of the colonization history of the Irish pygmy shrew, genetic variation was determined in 247 pygmy shrews collected throughout Ireland, using mitochondrial control region sequences and five polymorphic microsatellite loci. Genetic diversity was low for both types of marker. The median-joining network for control region sequences was star-like, suggesting that the colonization of Ireland involved a small number of founders and rapid population expansion thereafter; this was supported by other statistics. Molecular dating with both mitochondrial DNA and microsatellite data is consistent with a human introduction. This would have been several thousand years ago; a recent colonization within historical times can be ruled out. This is the first detailed population genetic study of the pygmy shrew anywhere in its range.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 918–927.     

Basal rate of metabolism and temperature regulation in Sorex coronatus and S. minutus (Soricidae: Mammalia)

1. The basal rate of metabolism (BMR), body temperature and thermal conductance of Sorex minutus and Sorex coronatus were studied. Special attention was given to measuring BMR in resting and postabsorptive shrews. 2. Both species exhibit a high BMR, respectively 339 and 289% of the value expected on the basis of body mass. 3. The thermal conductance of both species is similar to the predicted value and their body temperature averages 38.6 and 37.6 degrees C respectively. 4. Available data suggest that a very high BMR may be a common feature within the genus Sorex.     

Studies on the distribution, food, breeding biology and relative abundance of the Pygmy and Common shrews (Sorex minutus andS. araneus) in upland areas of northern Engl and.

The distribution and relative abundance of the Pygmy and Common shrews were investigated over two years at 42 sites across northern England and at a further seven sites in one year. Most of the sites were in upland areas above 250m in altitude. At almost all sites, the Pygmy shrew was more abundant than the Common shrew, often by a considerable
A correlation exists between the numbers of shrews captured at a site and the abundance of invertebrates revealed by pitfall trapping. I t is recorded that Common shrews fed on earthworms (which are almost completely absent on peatlands) whilst Pygmy shrews did     

Fiber capillarization relative to mitochondrial volume in diaphragm of shrew.

The objective was to examine fiber capillarization in relation to fiber mitochondrial volume in the highly aerobic diaphragm of the shrew, the smallest mammal. The diaphragms of four common shrews [Sorex araneus; body mass, 8.2 +/- 1.3 (SE) g] and four lesser shrews (Sorex minutus, 2.6 +/- 0.1 g) were perfusion fixed in situ, processed for electron microscopy, and analyzed by morphometry. Capillary length per fiber volume was extremely high, at values of 8,008 +/- 1,054 and 12,332 +/- 625 mm(-2) in S. araneus and S. minutus, respectively (P = 0.012), with no difference in capillary geometry between the two species. Fiber mitochondrial volume density was 28.5 +/- 2.3% (S. araneus) and 36.5 +/- 1.4% (S. minutus; P = 0.025), yielding capillary length per milliliter mitochondria values (S. araneus, 27.8 +/- 1.5 km; S. minutus, 33.9 +/- 2.2 km; P = 0.06) as high as in the flight muscle of the hummingbird and small bats. The size of the capillary-fiber interface (i.e., capillary surface per fiber surface ratio) per fiber mitochondrial volume in shrew diaphragm was also as high as in bird and bat flight muscles, and it was about two times greater than in rat hindlimb muscle. Thus, whereas fiber capillary and mitochondrial volume densities decreased with increased body mass in S. araneus compared with S. minutus Soricinae shrews, fiber capillarization per milliliter mitochondria in both species was much higher than previously reported for shrew diaphragm, and it matched that of the intensely aerobic flight muscles of birds and mammals.     

The trophic impact of small mammals in successional grasslands

The habitat use and diets of small mammals inhabiting grassland plots of different successional ages were investigated by live-trapping and faecal analysis over a period of 16 months. The contribution of the major plant life forms and the structural profile of the vegetation of each plot and the availability of insect prey were assessed. The dominant species of small mammal found on each plot were Sorex araneus, S. minutus, Apodemus sylvaticus and Microtus agrestis . Small mammals were most commonly found in the late and mid-successional stages, reflecting the habitat structure and food availability. A wide range of insects and other invertebrates were eaten by all four species and insects formed 50–62% of the animal prey taken. The incidence of certain prey in the diets differed between plots. The daily consumption rate of invertebrates by shrews and mice was estimated at some 6800 prey per ha. The greatest predatory impact came from shews but A. sylvaticus showed increased consumption of invertebrates in spring. The combined predatory impact of the grassland small mammal community on insect populations alone was estimated to average 0.01% per day and is predicted to be greatest in the mid-successional stages.     

Shrews (Sorex spp.) somatometry and reproduction in Slovakia

The morphometrics and reproduction of shrews (Sorex araneus, S. minutus, S. alpinus) was studied in Slovakia. Significant differences in weight, body length and tail length were recorded in adult and subadult shrews. Adult individuals and males had higher mean somatometric values (apart from tail length of subadults). The hind foot length was the least variable characteristic, which can be considered as the most stable taxonomic somatic characteristic of shrews. Weight and body length varied considerably. Values of somatic characteristics in S. araneus and S. minutus increased with increasing altitude, apart from hind foot length in S. araneus, which decreased with increasing altitude. The values of somatic characteristics declined with increasing continentality, but tail length gradually increased in the direction west — east at the highland level. Sorex minutus was characterized by the greatest reproductive activity (length of reproductive cycle April–October, average number of embryos 6.97). The lowest reproductive activity was observed in S. alpinus (average number of embryos 5.72). The mating season of S. araneus began at the end of March and ended in August with an average number of embryos of 6.12.