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.     

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.     

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.     

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.     

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.     

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 electrocardiogram of European shrews

A recording method of electrocardiograms in resting animals is described. The electrocardiograms of Crocidura russula, Crocidura leucodon, Crocidura suaveolens, Neomys fodiens and Sorex araneus are similar to those of other small mammals. T-wave directly follows the QRS-complex. The duration of electrocardiograms depends on heart mass and body temperature. Heart rate shows no influence on the duration of electrocardiograms.     

The feeding ecology of the European Water shrew

This review examines our knowledge of the feeding ecology of the European Water shrew, Neomys fodiens. It describes and discusses the different methods which have been used to study the diets of Water shrews, from the first casual observations to the discovery of food caches and food remains in the wild, and the analysis of stomach contents and faecal pellets of trapped animals. These studies have revealed that Water shrews feed on a variety of invertebrate and vertebrate prey from tiny freshwater dip-teran larvae to sizable fish and frogs. Although aquatic prey are seen to be the major dietary component in most studies, terrestrial prey are also important and may be eaten in an equal proportion. Laboratory studies have increased our knowledge of the foraging behaviour of Water shrews but there is still much to be learnt, especially about underwater prey location.     

Mitochondrial DNA reveals different phylogeographic structures in the water shrews Neomys anomalus and N. fodiens (Insectivora: Soricidae) in Europe

We studied the intraspecific genetic variation of 17 Neomys anomalus (Cabrera, 1907) (eight localities) and 24 Neomys fodiens (Pennant, 1771) (nine localities) in Europe. As molecular marker, we used the sequences of the mtDNA gene for cytochrome b (1140 bp from 24 specimens; 272 bp from 15 specimens). The phylogenetic analysis of N. fodiens shows slight divergence among haplotypes, with only one supported clade including two geographically distant haplotypes from Calabria (southern Italy) and Pyrenees (France); these haplotypes diverge from the others by 2.9% and 2.1%, respectively. In N. anomalus , one haplotype from Spain is very different, diverging from all others by 8–10% of the sequence. This level of divergence is similar to that found between N. anomalus and another species, Neomys teres . There are two other supported clades in N. anomalus , an Italian subclade and an eastern subclade comprising haplotypes from Turkey and Slovenia. Moreover, we found a different genetic structure between the two species with greater divergence in N. anomalus than in N. fodiens. The phylogeographic patterns are discussed in a temporal framework and considering the different ecological characteristics of these two species.     

Synchrony between small mammal population dynamics in marshes and adjacent grassland in a landscape of the Jura plateau, France: a ten year investigation

Small mammal populations were studied during a 10 year multi-annual fluctuation of vole populations in marshes and adjacent grassland of the basin in the Jura mountains, France. Nine species were monitored:Microtus agrestis, Clethrionomys glareolus, Apodemus flavicollis, Sorex araneus Icoronatus, Neomys fodiens, Crocidura leucodon, Mustela nivalis in marshes andArvicola terrestris andMicrotus arvalis in grassland, using live-trapping and index methods during July from 1993 to 2002. Populations of A.terrestris, M. arvalis, M. agrestis, S. araneus I coronatus andM. nivalis exhibited a relatively strong inter-specific temporal synchrony in their multi-annual fluctuations.C. glareolus population fluctuations were not synchronous to theArvicola I Microtus I Sorex group, and were closer to those ofApodemus flavicollis, Neomys andCrocidura. Those results are discussed in the light of earlier studies carried out in the same region and in northern ecosystems.     

Emmonsiosis of wild rodents and insectivores in Czechland

Adiaspores of the fungus Emmonsia crescens were detected microscopically in the lung tissue of 13% of 10.081 small mammals belonging to 24 species examined in 14 areas of the Czech Republic between 1986 and 1997; 441/1.934 (23%) Clethrionomys glareolus, 1/6 (17%) Arvicola terrestris, 357/2.172 (16%) Apodemus flavicollis, 220/1.981 (11%). A sylvaticus, 23/265 (9%) A. microps, 11/81 (14%) Microtus subterraneus, 93/1.275 (7%) M. arvalis, 98/1.439 (7%) M. agrestis, 1/3 (33%) Ondatra zibethicus, 1/1 Cricetus cricetus, 1/20 (5%) Crocidura suaveolens, 2/40 (5%) Neomys fodiens, and 13/529 (2%) Sorex araneus were infected. Emmonsiosis was not recorded among the species of rodents that do not build their nests in the soil (Muscardinus avellanarius, Micromys minutus, Mus musculus, Rattus norvegicus). The overall prevalence of emmonsiosis was significantly higher in adult (19%) than in juvenile (7%) mammals, and in rodents (13%, and 20% in adults) than in insectivores (2%, and 4% in adults). The frequency of infected mammals also varied according to geographic area, altitude, habitat, and season.     

Histological description of seminiferous epithelium and cycle length of spermatogenesis in the water shrew Neomys fodiens (Mammalia: Soricidae)

Recently, we examined the spermatogenesis cycle length in two shrews species, Sorex araneus characterized by a very high metabolic rate and a polyandric mating system (sperm competition) resulting in a short cycle and Crocidura russula characterized by a much lower metabolic rate and a monogamous mating system showing a longer cycle. In this study, we investigated the spermatogenesis cycle in Neomys fodiens showing an intermediate metabolic rate. We described the stages of seminiferous epithelium according to the spermatid morphology method and we calculated the cycle length of spermatogenesis using incorporation of 5-bromodeoxyuridine into DNA of the germ cells. Twelve males were injected intraperitoneally with 5-bromodeoxyuridine, and the testes were collected. For cycle length determination, we applied a recently developed statistical method. The calculated cycle length is 8.69 days and the total duration of spermatogenesis based on 4.5 cycles is approximately 39.1 days, intermediate between the duration of spermatogenesis of S. araneus (37.6 days) and C. russula (54.5 days) and therefore congruent with both the metabolic rate hypothesis and the sperm competition hypothesis. Relative testes size of 1.4% of body mass indicates a promiscuous mating system.     

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.     

Riparian contributions to the diet of terapontid grunters (Pisces: Terapontidae) in wet–dry tropical rivers

The diets of terapontid assemblages in 22 catchments across Australia's wet–dry tropics were investigated in relation to the direct use of terrestrial‐riparian inputs, as well as the role of ontogeny and morphology in mediating consumption of allocthonous material. The diet of several species was restricted almost entirely to instream trophic resources throughout their life history. In contrast, ontogenetic diet shifts towards increasing consumption of terrestrial prey types were a prominent feature of the dietary ecology of some terapontids, with collective allocthonous dietary items making a significant contribution (up to 42%) to diet in larger size classes of several species. For those species consuming terrestrial‐riparian material in their diet, terrestrial invertebrates were the most common prey item; however, terrestrial vegetation, principally riparian fruits, and terrestrial vertebrates were also important dietary inclusions in the larger size classes of particular species. A large mouth gape was the morphological feature most strongly associated with consumption of terrestrial food resources within the Terapontidae. Results indicate that the direct consumption of terrestrially derived food sources in northern Australian aquatic systems may be more important than previously asserted, and that additional research is required to better clarify the role of terrestrial subsidies to these ecosystems.     

Food of introduced mosquitofish: ontogenetic diet shift and prey selection

The mosquitofish Gambusia holbrooki of Lake Banyoles, Catalonia, Spain, were strictly littoral. Their population was dominated by females throughout the year, except in spring when males were more common. Their diet was based on littoral cladocerans, particularly Chydorus sphaericus, Scapholeberis ramneri, Ceriodaphnia reticulata , and Pleuroxus laevis , and nematoceran (basically chironomid) adults. There was a large variety of prey of terrestrial (collembolans, ants) or aquatic neustonic origin ( S. ramneri , emerging nematoceran adults), showing the microhabitat of mosquitofish closely linked to the water surface. In contrast to other studies, terrestrial insects were not especially important in summer. In addition to seasonal and between-site variation, there was an ontogenetic diet shift from microcrustaceans, particularly cladocerans (smallest fish also using diatoms and copepod nauplii) to larger prey, namely nematoceran adults. However, there was an unusual, inverse relationship between fish length and the size of two prey ( S. ramneri and the collembolan Sminthurides sp.), paralleling the overall increase in mean prey size.     

Diet and food selection by Ramnogaster arcuata (Osteichthyes, Clupeidae)

The goals of this study were to describe the diet and estimate the trophic level and food selection of Jenyns's sprat Ramnogaster arcuata, one of the common fishes in the Bahía Blanca estuary in Argentina. The copepods Acartia tonsa [53·7% index of relative importance (I(RI))] and Eurytemora americana (13·4% I(RI)), the mysid Arthromysis magellanica (13·9%I(RI)) and the small shrimp Peisos petrunkevitchi (8·2% I(RI)) were the most important food items for this species, and its diet exhibited monthly variability with respect to the principal prey categories. The zooplankton species in the study area consisted of 32 taxa. Acartia tonsa was the most abundant species [54% individuals (N) m(-3)], followed by Balanus glandula (13·8% N m(-3)) and E. americana (10·1% N m(-3)). The rest of the zooplankton groups represented <3% N m(-3). The most preferred items throughout the year were A. magellanica and Idotea sp. The trophic level value estimated was 3·1.     

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.     

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.     

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.     

Prey selection and foraging constraints in common poorwills (Phalaenoptilus nuttallii: Aves: Caprimulgidae)

Predators that hunt by sight should be constrained by light levels. This problem should be especially acute in aerial predators such as common poorwills ( Phalaenoptilus nuttallii : Caprimulgidae), which capture small insects at night. The purpose of our study was to test the hypothesis that poorwills are constrained to catching larger insect taxa due to limitations of visual orientation under low light. We assessed diet by analysing poorwill faecal samples collected at nest and roost sites in the Cypress Hills of Saskatchewan where prey density is relatively low. Coleopterans (44% and 39% in 1991 and 1992, respectively) and lepidopterans (46% and 55%) dominated the diet, yet these two insect orders were rare in light-suction trap (22% lepidopterans and 1% coleopterans) and sticky trap samples (11% lepidopterans and 2% coleopterans). Based on measurements of femur length, all consumed coleopterans were > 5 mm long, even though 57% of those available were < 5 mm. Our results suggest that poorwills are constrained to taking prey above a certain size, but above this threshold, particular prey types and larger sizes may be selected. Selection for coleopterans may reflect the need to acquire polyunsaturated fat to facilitate the use of torpor.