The genus Sorex (Mammalia, Soricidae)–distribution and evolutionary aspects of Eurasian species

There are about 60 species of shrews of the genus Sorex distributed over the Nearctic and Palaearctic regions; these are usually divided into the subgenera Sorex, Otisorex and Microsorex . The Old World species are listed and their phylogeny discussed, with special emphasis laid on the karyotypes. It is proposed that the Old World forms should be divided into the subgenera Homalurus, Sorex and Otisorex . It is also suggested that the subgenus Sorex could be divided into at least three taxonomic units.     

Population genetic structure of Sorex unguiculatus and Sorex caecutiens (Soricidae, Mammalia) in Hokkaido, based on microsatellite DNA polymorphism

We investigated the genetic structure of Sorex unguiculatus and Sorex caecutiens populations in Hokkaido, Japan, using hypervariable microsatellite DNA markers. We used five microsatellite loci to type 475 S. unguiculatus individuals from 20 localities on the Hokkaido mainland and four localities from each of four offshore islands (and 11 shrews from one locality in southern Sakhalin for a particular analysis). We used six microsatellite loci to type 240 S. caecutiens individuals from 13 localities on the Hokkaido mainland. Genetic variation was high in mainland populations of both species and low in the island populations of S. unguiculatus. Allelic richness and island size were positively correlated for S. unguiculatus, suggesting that genetic drift occurred on those islands due to small population size. In addition, four insular populations of S. unguiculatus were genetically differentiated from the mainland populations, although clear phylogeographic clustering was not confirmed among populations on the Hokkaido mainland for either S. unguiculatus or S. caecutiens. Heterozygosity excess was observed in more than half of the populations including the mainland populations of the two species, suggesting recent bottleneck events in these populations. Population dynamics of the shrews might be explained by a metapopulation scheme. According to autocorrelation analysis, the extent of non-random spatial genetic structure was approximately 100 km. Isolation by distance was observed in S. unguiculatus, but not in S. caecutiens although there is a positive trend. The lack of correlation for S. caecutiens might have been due to small sample size. Thus, no obvious differences in population genetic structure were found between the two species on the Hokkaido mainland in the present study, while previous investigations using mitochondrial DNA sequences inferred that these two species might have rather different biogeographic histories.     

Mechanisms of chromosomal evolution in three European species of the Sorex araneus-arcticus group (Insectivora: Soricidae)

Presented are the R-banding patterns of the karyotypes of three European species of the Sorex araneus-arcticus group (Insectivora: Soricidae). The eight species of this Holarctic complex are characterized by sharing a male chromosomal set of XY1Y2 elements. Robertsonian and tandem translocations are common in this complex, at the population level as well as at the species level. The rough morphology of the karyotypes looks similar between all the species presently described. An R-banding technique RHG, RBG) has allowed us to make a comparative analysis of the chromosomal similarities in three species, namely S. araneus, S. coronatus and S. granarius. The data provide evidence that Robertsonian and tandem translocations, accompaniei in some cases by centromeric shifts, are the main, if not the only, mechanisms of chromosomal evolution in this grou. It appears that S. granarius presents a karotype which is most similar to the hypotheticar ancestral type from which the chromosomar sets of two other European species might be derived.     

Post-Pleistocene evolution of Bornean shrews Crocidura foetida (Mammalia, Soricidae)

Fossil mandibles of the Bornean shrew Crocidura foetida recovered from excavations at the west mouth of Niah cave, Sarawak, Malaysia, show that the late Pleistocene population at this lowland location was comparable in size with the large subspecies Crocidura foetida doriae , presently occurring at inland, upland locations. Two Holocene specimens fall in the size range of the smaller lowland subspecies C. f. foetida . Comparable post-Pleistocene size-reduction is known among other mammals of Borneo, but this is the first instance of dated examples. The evolutionary trend conforms with Bergmann's 'rule' but, other than climate change, no selective agent is apparent.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 413–419.     

Origin and evolution of the endemic Canary Island shrews (Mammalia: Soricidae)

Until recently the North Atlantic Islands were believed to house only mammals introduced by humans. Recent work has demonstrated that at least the Canary Islands house(d) a native mammal fauna. New data including chromosome numbers, genetic distances and analysis of vocalizations are given for the two extant shrew species, Crocidura canariensis and C. osorio , and their possible sister taxa are evaluated. Evidence is presented for the hypothesis that the two island species originated from two different lineages of the Palaearctic branch of the genus Crocidura. The data support the present status of the Canary Island shrews as local endemics of high conservation priority.     

Phylogenetic evolution and biogeography of Southeast Asian shrews (genus Crocidura: Soricidae)

Genetic variation of 20 species of shrews from the Malay peninsula, Sumatra, Java, Borneo, the Philippines and Sulawesi was assessed by allozyme electrophoresis at 32 loci. According to Mantel's tests, the genetic differentiation of these species of shrews is not a function of the geographic distance separating them (r = 0.09, NS), but is correlated to the water depth surrounding the islands where they live (r = 0.49, P < 0.01). The results are just the reverse if the correlations are computed for the Sunda Shelf taxa only. In this case, the sampled populations show an isolation-by-distance relationship (r = 0.32; P < 0.01), while no significant correlation with water depth was detected (r = 0.20; P - 0.07). Qualitative predictions based on eustatic sea level variation and water depth were formulated as a model of historic connections between the islands. This palaeogeographic model was tested through Brooks Parsimony Analysis. The assumption of a simple vicariant evolution of the shrews was rejected, but several concordant patterns indicate that the phytogeny of these mammals was indeed shaped by these events. Homoplasies demonstrated that the SE Asian species of Crocidura include composite zoogeographic histories. Sulawesi, for example, supports at least six species, five of which are closely related, while the last one, C. nigripes, is more closely related to a Bornean taxon. This pattern was interpreted as the result of a first wave of colonizers which subsequently radiated, followed by a more recent, second colonization event from Borneo. The overall small genetic distance found within the assemblage of the five old endemics (D_N= 0.151 pL 0.041) suggests that the radiation was not accompanied by extensive differentiation, although from a karyological point of view, they exhibit unusual variations when compared to other Indomalayan Crocidura. By contrast, the four species found on Sumatra are more differentiated (D_N= 0.221 pL 0.063) and never form sister-group relationships in any phylogenetic reconstruction; each one is more closely related to different taxa living outside Sumatra. This suggests that they are probably remnants of an important centre of dispersal for the entire Malay Archipelago. The standard genetic distance averaged among all Southeast Asian species (D_N= 0.235 pL 0.094) is about half that measured within Palearctic or African taxa. Such an overall lower mean level of genetic variability is consistent with the hypothesis of a relatively recent colonization of the Malay Archipelago by shrews of the genus Crocidura.     

Testis size, sperm characteristics and testosterone concentrations in four species of shrews (Mammalia, Soricidae)

The aim of this study was to establish and compare the sperm characteristics in four shrew species in the context of the sperm competition hypothesis. As expected, the large relative testis size in promiscuous species was associated with a high number of cauda epididymal spermatozoa and a high concentration of circulating testosterone. In addition, in Sorex and Neomys, species with high intensity of sperm competition, the spermatozoa stored in cauda epididymis were characterized by high percentage of progressive motility whereas in Crocidura and Suncus, the cauda epididymal spermatozoa were motile but with very low percentage of progressive motility. This capability is achieved only following the passage through the vas gland, a specialized region for sperm storage located along the vas deferens in these shrew species. The hypothesis that sperm competition is positively correlated with spermatozoa length could not be confirmed. In Crocidura and Suncus, the total sperm length is increased by the large sperm head due to a big acrosome. This trait, specific to the subfamily Crocidurinae, may results from a selective pressure independent of the context of sperm competition, related to a specific, but as yet unclear role, for the acrosome during the fertilization.     

Phylogeny and evolution of African shrews (Mammalia: Soricidae) inferred from 16s rRNA sequences

Current phylogenetic hypotheses on the African Crocidurinae (Soricidae) are based upon morpho-anatomical, karyological, and allozyme studies. The present study attempts to resolve the interrelationships among African Crocidurinae and their relationships to Eurasian Crocidurinae and to the subfamily Soricinae, on the basis of partial mitochondrial 16s rRNA sequences (549 bp). This is the first molecular study to include all but one of the nine currently recognized African shrew genera. In agreement with current views, two major lineages emerge. The first lineage includes Myosorex and Congosorex and supports the existence of a myosoricine taxon. The second lineage includes the six remaining genera. The genus Sylvisorex appears to be polyphyletic, whereas species of the controversial genus Crocidura are monophyletic. The genus Suncus presumably originated in Africa. The monospecific genera Ruwenzorisorex and Scutisorex and the two representatives of Paracrocidura cluster with species of other genera. Grouping patterns of species from different continents suggest that there have been multiple exchanges between Africa and Eurasia. The time estimates of these exchanges, inferred from two independent fossil-based calibrations of a molecular clock, coincide with the time estimates for migration events in other mammalian taxa.     

Habitat, morphology and karyotype of the Saharan shrewCrocidura tarfayaensis (Mammalia: Soricidae)

The Saharan shrewCrocidura tarfayaensis Vesmanis and Vesmanis, 1980, has a limited disribution along the Atlantic coast of Sahara, south of Agadir (Morocco) through Western Sahara into Mauritania and is only known from few captures and some owl pellets. Here we report field data from the successful trapping of five specimens ofC. tarfayaensis in the Guelmim region. The habitat was characterized by sand dunes along a river, with dense shrubberies ofTamarix sp., the huge grassErianthus ravennae (Poaceae) and flat bushes of Atriplex glauca var. ifniensis (Chenopodiaceae). Morphological discrimination withC. whitakeri were examined. The chromosomes ofC. tarfayaensis revealed a karyotype of 2n = 36, similar to that of the Canary shrewC. canariensis and the Sicilian shrewC. sicula. In conclusion,C. tarfayaensis seems to be a descendant of the presumed continental ancestor of the two island species.     

Recombinogenic Telomeres in Diploid Sorex granarius (Soricidae,Eulipotyphla) Fibroblast Cells

The telomere structure in the Iberian shrew Sorex granarius is characterized by unique, striking features, with short arms of acrocentric chromosomes carrying extremely long telomeres (up to 300 kb) with interspersed ribosomal DNA (rDNA) repeat blocks. In this work, we investigated the telomere physiology of S. granarius fibroblast cells and found that telomere repeats are transcribed on both strands and that there is no telomere-dependent senescence mechanism. Although telomerase activity is detectable throughout cell culture and appears to act on both short and long telomeres, we also discovered that signatures of a recombinogenic activity are omnipresent, including telomere-sister chromatid exchanges, formation of alternative lengthening of telomeres (ALT)-associated PML-like bodies, production of telomere circles, and a high frequency of telomeres carrying marks of a DNA damage response. Our results suggest that recombination participates in the maintenance of the very long telomeres in normal S. granarius fibroblasts. We discuss the possible interplay between the interspersed telomere and rDNA repeats in the stabilization of the very long telomeres in this organism.     

A new species of Deinsdorfia (Soricidae, Insectivora, Mammalia) from the Pliocene of Spain

A new species of Deinsdorfia is defined using new material coming from the karstic fissure infilling of Almenara-Casablanca 4 (Castelló, Spain), which shows important differences with the rest of the previously known species of the genus. The definition of a new species is supported by the characters found in extra material coming from the localities of Valdeganga, in the Albacete Province (Spain). The stratigraphical occurrence of this genus of pigmented-toothed shrews reinforces the idea of the Iberian Peninsula as a sink area during the Pliocene, where some Central-European soricids expanded their distribution. Most of those migrations or distributional expansions coincide with moments of climatic changes in the northern hemisphere.     

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.     

Distribution of two karyotypic races of Sorex araneus (Insectivora, Soricidae) in Switzerland and the post-glacial recolonization of the Valais: First results

Two chromosome races of Sorex araneus were found in southern and western Switzerland. It is suggested they were issued by introgression of different sets of metacentrics, coming respectively from north-east and south, into a primitive “acrocentric” population which recolonized the country from the west after the last glaciation.     

First record of Entelodontidae (Mammalia,Artiodactyla) from the late Eocene of Southeast Asia

We describe two entelodontid upper premolars that were recovered from the late Eocene of the Krabi coal mine in southern Thailand. The size and morphology of the material suggest that it can be referred to Entelodon aff. E. gobiensis, a species known from the late Eocene to the early Oligocene of northern Asia and southern China. The Thai material documents for the first time the southernmost occurrence of entelodontids in Asia during the Paleogene and also suggests that Eocene Southeast Asian mammal localities might potentially yield further entelodontid remains mostly associated with selenodont ungulates.     

Soricidae (Mammalia) from the Early Pleistocene of Olduvai Gorge, Tanzania

Soricid remains collected from Bed I of Olduvai Gorge are described. The great majority of the specimens are mandibles. A survey of the mandibles of living African species revealed many differences in characters of the lower teeth and jaw that can be used for identification. On the basis of these characters, nine species are distinguished in the Olduvai collection, of which six are well enough preserved to permit a discussion of their relationships to living species. Three new species and one new subspecies are described. All the Olduvai shrews differ in some respects from their nearest living relatives; three species are close to species from Makapansgat, Swartkrans and Sterkfontein, RSA, though there appear to be slight differences. A change in the soricid fauna takes place within Bed I, interpreted as due to increasing aridity.