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.     

Phylogenetic relationships of Caucasian shrew Sorex satunini Ogn. (mammalia) in the superspecies Sorex araneus inferred from the data of karyological analysis and the mtDNA cyt b gene sequencing

Using the data of karyological analysis, the phylogenetic relationships of Caucasian shrew Sorex satunini and the cryptic species of superspecies Sorer araneus were examined. In the population of Sorex satunini from the plain of North Ciscaucasia two deeply radiated cytochrome b genes (A and B) were identified. Genetic distance between haplotype A and B groups constituted 0.0675 +/- 0.008, which is higher than any distance in superspecies S. araneus. Possible introgression of type B haplotypes from the populations of the evolutionary lineage S. subaraneus--S. araneus in Pleistocene and the time of the appearance of the chromosomal polymorphism of S. araneus is discussed. Our results show that the use of only one mitochondrial marker can lead to false conclusions on taxonomic diversity     

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.     

Living at the physiological limits: field and maximum metabolic rates of the common shrew (Sorex araneus)

Shrews (genus Sorex, small insectivorous mammals) are well known for their extremely high basal metabolic rates (BMRs) even when corrected for their small body size. We measured energy expenditure of the common shrew (Sorex araneus) under natural conditions (field metabolic rate [FMR]) by doubly labeled water method to test whether FMR is proportional to high BMR in this species. The study was performed in summer in northeastern Poland. In addition to the FMR, we also measured maximum metabolic rates induced by cold exposure and by intense activity (MMRCOLD and MMRRUN, respectively) to evaluate the aerobic reserve (MMR-FMR) in S. araneus. This aerobic reserve was used as an indicator of the potential for metabolic constraints. The FMR averaged 2.31+/-0.32 L CO2 d(-1) (+/-SD) or 58.1+/-8.0 kJ d(-1) in 8.2-g animals. This figure constituted 216%-258% of a value predicted for a "standard" mammal of the same body mass and was the highest mass-specific field metabolic rate in mammals. Because of the high BMR level in S. araneus, the FMR to BMR ratio (2.4) was not far off mammalian standards (median value of 3.1). The rate of water efflux determined in S. araneus (20.2 mL H2O d(-1) or 2.46 mL H2O g(-1) d(-1)) exceeded all figures reported to date in other mammals and was apparently linked to the high FMR level and relatively high water content of shrews' food. Maximal metabolic rates (MMRRUN of 18.1+/-1.6 mL O2 g(-1) h(-1) and MMRCOLD of 23.5+/-1.9 mL O2 g(-1) h(-1)) were not high in proportion to BMR or FMR that resulted in relatively narrow aerobic reserve in S. araneus: 20% when calculated against the MMRRUN and 39% when compared with the MMRCOLD. Our study reveals that S. araneus has high energy costs of living and operates close to its physiological limits.     

Dietary separation in three species of shrew inhabiting water-cress beds

The diets of three species of shrew (Neomys fodiens, Sorex araneus and S. minutus) living sympatrically in an area of water-cress beds were investigated over a two-year period by faecal analysis of live-trapped animals. Thirty-eight different prey types were identified in the diet of N. fodiens , 25 in the diet of S. araneus and 18 in S. minutus. The most important prey items of N. fodiens in all months were freshwater crustaceans and cased trichopteran larvae. Other important items were terrestrial gastropods, coleopterans and dipterans. Neomys fodiens foraged underwater throughout the year and 33–67% of its prey were of aquatic origin. Major prey items of S. araneus throughout the year were lumbricids, terrestrial gastropods and coleopterans. Araneids, isopods and coleopterans were the dominant dietary items of S. minutus. All terrestrial prey items were taken in greater proportions by S. araneus and S. minutus than by N. fodiens , excepting lumbricids which were never eaten by S. minutus. Dietary overlap was greatest between S. araneus and S. minutus at 57%, followed by N. fodiens and S. araneus (44%) and least similar were N. fodiens and S. minutus (36%).     

Structure and seasonal dynamics of an ectoparasite community on the common shrew (Sorex araneus) in the Il'men'-Volkhov lowland

Species composition, seasonal dynamics, and a load of ectoparasites per individual and population of the common shrew Sorex araneus have been examined in coniferous and mixed forests of the Ilmen'-Volkhov lowland (a neighbourhood of Oskuy village, Chudovo district, Novgorod Province) during the period 1999-2003. Trapping of mammals was carried monthly, with exception of few accidental gaps. The Gero traps were used for catching micromammalian hosts. The lines of trap were checked 2 times a day, places of lines changed each 3-5 day. Total number of micromammalians collected during the period of study is 3215, including 1115 specimens of the common shrew S. araneus and 246 ones of the pigmy shrew S. minutus. Parasite fauna on the common shrew included 23 ectoparasite species: fleas--12, ixodid ticks--2, gamasid mites--7, and myobiid mites--2 species. Among recorded species, 9 fleas species and some gamasid species are accidental parasites. These accidental ectoparasite species are common to the bank vole Clethrionomys glareolus in the territory investigated. Species composition, occurrence and abundance indices of parasites changed during the year. In total, about 55% shrew specimens are infected with ectoparasites. The infracommunity of ectoparasites on the common shrew usually consists of 6 species or less. Mean number of all ectoparasite individuals per one host specimens varies from 4 to 83. The greatest number of parasites (50 and 83) was recorded on the shrews, which carried 5 and 4 parasites species, respectively. Biodiversity of parasite species in the ectoparasite community on the common shrew and the load of parasites per one host specimen are lower than those in the bank vole. In forest biotopes explored, the most part of temporary ectoparasite species found on the common shrew was also recorded on other small mammals, which could have contacts with this host. It is possible to conclude that among the parasite supracommunity in the explored ecosystem, the temporary ectoparasites represent a "fond" of temporary parasites, which are common for most small mammal species. A role of different shrew and rodent species as main of additional hosts changes depending on a population density of potential mammalian hosts and other environment factors.     

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.     

Small mammals and their ectoparasites from the Scottish islands of Handa (Sutherland), Muck, Pabay, Scalpay and Soay (Inner Hebrides)

Small mammals were trapped on five islands for short periods during the summers of 1964 and 1965, with the following results:
Handa: Rattus norvegicus only, probably no other species present.
Muck: Sorex araneus, S. minutus, Apodemus sylvaticus and Microtus agrestis; R. norvegicus also present.
Pabay: S. minutus and Neomys fodiens ; probably no other species present.
Scalpay: S. araneus, S. minutus and M. agrestis.
Soay: S. araneus and S. minutus ; rodents almost certainly absent.
Ectoparasites (fleas and Acarina) collected from these small mammals are tabulated.     

Karyotype evolution of eulipotyphla (insectivora): the genome homology of seven sorex species revealed by comparative chromosome painting and banding data

The genus Sorex is one of the most successful genera of Eulipotyphla. Species of this genus are characterized by a striking chromosome variability including XY1Y2 sex chromosome systems and exceptional chromosomal polymorphisms within and between populations. To study chromosomal evolution of the genus in detail, we performed cross-species chromosome painting of 7 Sorex species with S. granarius and S. araneus whole-chromosome probes and found that the tundra shrew S. tundrensis has the most rearranged karyotype among these. We reconstructed robust phylogeny of the genus Sorex based on revealed conserved chromosomal segments and syntenic associations. About 16 rearrangements led to formation of 2 major Palearctic groups after their divergence from the common ancestor: the S. araneus group (10 fusions and 1 fission) and the S. minutus group (5 fusions). Further chromosomal evolution of the 12 species inside the groups, including 5 previously investigated species, was accompanied by multiple reshuffling events: 39 fusions, 20 centromere shifts and 10 fissions. The rate of chromosomal exchanges upon formation of the genus was close to the average rate for eutherians, but increased during recent (about 6-3 million years ago) speciation within Sorex. We propose that a plausible ancestral Sorex karyotype consists of 56 elements. It underwent 20 chromosome rearrangements from the boreoeutherian ancestor, with 14 chromosomes retaining the conserved state. The set of genus-specific chromosome signatures was drawn from the human (HSA)-shrew comparative map (HSA3/12/22, 8/19/3/21, 2/13, 3/18, 11/17, 12/15 and 1/12/22). The syntenic association HSA4/20, that was previously proposed as a common trait of all Eulipotyphla species, is shown here to be an apomorphic trait of S. araneus.     

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.     

Helminthofauna of the common shrew Sorex araneus L. (Soricidae) from Samarskaya Luka

Ecological analysis of the helminthofauna of common shrew Sorex araneus L. from Samarskaya Luka (Samara Region) has been carried out. Twenty-three helminth species are found including 3 species of trematodes, 8 cestodes, 10 nematodes, and 2 acanthocephales. Dependence of the structure of helminthofauna on the host age is revealed.     

The helminth fauna of the shrew genus Sorex (Insectivora: Soricidae) in the Kandalksha nature reserve

The helminth fauna of shrews (Sores araneus, S. caecutiens, S. minutus, S. minutissimus) from the Karelian coast and islands of the Kandalaksha nature reserve was studied. 18 helminth species belonging to 3 taxonomic groups, Trematoda (3), Cestoda (8), and Nematoda (7) were found. The main body of the helminth fauna of shrews is represented by the nematodes species Longistriata codrus, L. didas, and the cestode species Ditestolepis diaphana. Inhabiting of shrews in the insular localities leads to the decrease of the species diversity and the abundance of their helminths. The distribution ranges of the shrew parasites have been outlined more accurately.     

A new perspective on the evolutionary history of western European Sorex araneus group revealed by paternal and maternal molecular markers

The species of the common shrew (Sorex araneus) group are morphologically very similar, but have undergone a spectacular chromosomal evolution. We investigate here the evolutionary history of the Sorex araneus group distributed in western Europe. In particular, we clarify the position of a difficult species, S. granarius, using sex-specific (mtDNA and Y-chromosome) markers. The karyotype of S. granarius is generally considered similar to the common ancestor of the restricted group considered here. The mtDNA data (1.4 kb) confirms the close relationship between S. granarius and S. araneus sensu stricto (hereafter S. araneus s.s.), but the Y-chromosome (3.4 kb) produces a quite different picture: S. granarius is closely related to another species, S. coronatus. Comparison of mtDNA and Y-chromosome phylogenies suggests that the genetic and chromosomal evolution in this group are disconnected processes. The evolutionary history of the south-western European populations of the S. araneus group can only be understood considering secondary contacts between taxa after their divergence, implying genetic exchanges by means of hybridization and/or introgression.     

Shrews on small islands: epigenetic variation elucidates population stability

Three large (4 to 8 ha) and 14 small islands (0.3 to 2 ha) in a lake in eastern Finland, all situated less than 0.5 km from the mainland, were surveyed for small mammals. Three species of shrew and two species of vole were resident in July 1982: Sorex araneus on 10, S. caecutiens on 2, S. minutus on 5, Microtus agrestis on 12 and Clethrionomys glareolus on 4 islands. Immigrants were trapped from tiny islets, and the data indicate that S. caecutiens and M. agrestis are better dispersers than S. minutus and C. glareolus , respectively. Microtus agrestis, S. araneus and C. glareolus occurred non-randomly, on subsets of the larger islands, while the two small Sorex species occurred more erratically, possibly because of competition with S. araneus . Juvenile sex ratio was male-biased on the mainland but female-biased on large islands, possibly because juvenile males move more and are more likely to emigrate from an island than juvenile females.
Metrical and non-metrical (epigenetic) cranial traits gave similar patterns of population differentiation in S. araneus . Two of the three large-island populations have differentiated from the mainland populations and from each other, suggesting that the populations are relatively stable. Small-island populations, which are often less than 10 individuals in size, showed little differentiation but had more epigenetic traits fixed than large-island and mainland populations (founder effect). This suggests that the small-island populations are unstable, have a high extinction rate.     

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.     

Preferential segregation of metacentric chromosomes in simple Robertsonian heterozygotes of Sorex araneus

One of the hypotheses explaining preferential transmission of metacentrics among simple Robertsonian (Rb) heterozygotes of the common shrew (Sorex araneus L.) invokes the existence of meiotic drive. Thus far, evidence that metacentrics are favoured at meiosis has been obtained indirectly, on the basis of crosses made under controlled conditions. The aim of the present work was to test the hypothesis in a direct study. We analysed products of chromosome segregation among 12 simple heterozygote male subjects from a wild population, with regard to jl, io, nr and mn Rb fusions. We were able to demonstrate significant segregation distortion in favour of all four metacentrics. The level of preferential segregation was independent either of the composition of chromosome arms or the dimensions of metacentrics. We also found that X chromosomes were favoured over Y1Y2 chromosomes during segregation. We discuss the role of meiotic drive in the evolutionary success of metacentric chromosomes in S. araneus, as well as in the emergence of post-hybridization modifications in the zones of contact between races.     

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.     

Intraspecies phylogenetic connections in Sorex araneus L.: a southern Baltic subgroup of chromosome races

Results of chromosome G-banding were used to identify two new karyotypic races of the common shrew (Sorex araneus L.) in the European part of Russia. The chromosomal diagnosis of race St. Petersburg included five metacentrics (hk, ip, jl, mq, and nr) and two acrocentrics (g and o) (2Na = 20); the diagnosis of race West Dvina included six metacentrics (gm, hk, ip, jl, no, and qr) (2Na = 18). The phylogenetic significance of the chromosomal markers gm, hk, and ip is considered and the possibility of reticulate evolution in the species S. araneus is discussed.     

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.     

Genetic and morphological variation in a partially isolated population of Caucasian shrew sorex Satunini (Mammalia)

Morphological and genetic variation at microsatellite loci of Caucasian shrew Sorex satunini Ogn. is examined and compared with that of the common shrew S. araneus L. Genetic distance at microsatellite loci between the common shrew and Caucasian shrew proved to be threefold higher than between chromosome races of the common shrew. The Caucasian shrew manifested low polymorphism in studies of both microsatellites and morphometric mandibular traits. The heterozygote deficit was also typical. These properties may be a consequence of partial isolation of the population and gene drift.