Inbreeding in the greater white-toothed shrew, Crocidura russula

We combined mark-and-recapture studies with genetic techniques of parentage assignment to evaluate the interactions between mating, dispersal, and inbreeding, in a free-ranging population of Crocidura russula. We found a pattern of limited and female-biased dispersal, followed by random mating within individual neighborhoods. This results in significant inbreeding at the population level: mating among relatives occurs more often than random, and F(IT) analyses reveal significant deficits in heterozygotes. However, related mating partners were not less fecund, and inbred offspring had no lower lifetime reproductive output. Power analyses show these negative results to be quite robust. Absence of phenotypic evidence of inbreeding depression might result from a history of purging: local populations are small and undergo disequilibrium gene dynamics. Dispersal is likely caused by local saturation and (re)colonization of empty breeding sites, rather than inbreeding avoidance.     

Sex-specific selective pressures on body mass in the greater white-toothed shrew, Crocidura russula

The direction, intensity and shape of viability-, sexual- and fecundity selection on body mass were investigated in a natural population of the greater white-toothed shrew (Crocidura russula), combining parentage assignment through molecular techniques and mark-recapture data over several generations. A highly significant stabilizing viability selection was found in both sexes, presumably stemming from the constraints imposed by their insectivorous habits and high metabolic costs. Sexual selection, directional in both sexes, was twice as large in males than in females. Our results suggest that body mass matters in this context by facilitating the acquisition and defense of a breeding territory. No fecundity selection could be detected. The direction of sexual size dimorphism (SSD) was in agreement with the observed pattern of selective pressures: males were heavier than females, because of stronger sexual selection. SSD intensity, however, was low compared with other mammals, because of the low level of polygyny, the active role of females in territory defense and the intensity of stabilizing viability selection.     

Mitochondrial DNA variation along an altitudinal gradient in the greater white-toothed shrew, Crocidura russula

The distribution of mitochondrial control region-sequence polymorphism was investigated in 15 populations of Crocidura russula along an altitudinal gradient in western Switzerland. High-altitude populations are smaller, sparser and appear to undergo frequent bottlenecks. Accordingly, they showed a loss of rare haplotypes, but unexpectedly, were less differentiated than lowland populations. Furthermore, the major haplotypes segregated significantly with altitude. The results were inconsistent with a simple model of drift and dispersal. They suggested instead a role for historical patterns of colonization, or, alternatively, present-day selective forces acting on one of the mitochondrial genes involved in metabolic pathways.     

Disentangling reasons for low Y chromosome variation in the greater white-toothed shrew (Crocidura russula)

Y chromosome variation is determined by several confounding factors including mutation rate, effective population size, demography, and selection. Disentangling these factors is essential to better understand the evolutionary properties of the Y chromosome. We analyzed genetic variation on the Y chromosome, X chromosome, and mtDNA of the greater white-toothed shrew, a species with low variance in male reproductive success and limited sex-biased dispersal, which enables us to control to some extent for life-history effects. We also compared ancestral (Moroccan) to derived (European) populations to investigate the role of demographic history in determining Y variation. Recent colonization of Europe by a small number of founders (combined with low mutation rates) is largely responsible for low diversity observed on the European Y and X chromosomes compared to mtDNA. After accounting for mutation rate, copy number, and demography, the Y chromosome still displays a deficit in variation relative to the X in both populations. This is possibly influenced by directional selection, but the slightly higher variance in male reproductive success is also likely to play a role, even though the difference is small compared to that in highly polygynous species. This study illustrates that demography and life-history effects should be scrutinized before inferring strong selective pressure as a reason for low diversity on the Y chromosome.     

Sexual receptivity and oestrus in the white-toothed shrew, Crocidura russula monacha.

In common white-toothed shrews, the vaginal smear was an inadequate indicator of receptivity. Nucleated epithelial cells usually predominated and most matings took place when the smear was of this type. Cornified cells in the smear in infrequent. Mating tests showed that virgin females 2 to 4 months old, were unreceptive for extended periods. Willingness to mate increased gradually with the age of the female. Females were receptive throughout pregnancy, suggesting a continuous secretion of oestrogen during gestation. There was a post-partum oestrus whether or not the young of the litter were being suckled, and an oestrus at the end of lactation. During mid-lactation, females were in a state of low receptivity, whether or not they were suckling.     

Paradoxical social effects on response to cold during ontogeny of the common white-toothed shrew,Crocidura russula

Summary Postnatal development of thermoregulatory responses to cold differs between shrews measured singly or in groups as a litter. Single shrews increase their metabolic response to cold within 14 days to maintain normothermia. Shrews measured in groups show little or no metabolic response to cold and do not reach normothermia before the 19th day. Lower body temperatures in groups are explained by torpor, which obviously needs the prerequisite of forming a group. Oxygen consumption of the singly measured shrews depends upon body mass.     

Evolutionary history of the greater white-toothed shrew (Crocidura russula) inferred from analysis of mtDNA, Y, and X chromosome markers

We investigate the evolutionary history of the greater white-toothed shrew across its distribution in northern Africa and mainland Europe using sex-specific (mtDNA and Y chromosome) and biparental (X chromosome) markers. All three loci confirm a large divergence between eastern (Tunisia and Sardinia) and western (Morocco and mainland Europe) lineages, and application of a molecular clock to mtDNA divergence estimates indicates a more ancient separation (2.25 M yr ago) than described by some previous studies, supporting claims for taxonomic revision. Moroccan ancestry for the mainland European population is inconclusive from phylogenetic trees, but is supported by greater nucleotide diversity and a more ancient population expansion in Morocco than in Europe. Signatures of rapid population expansion in mtDNA, combined with low X and Y chromosome diversity, suggest a single colonization of mainland Europe by a small number of Moroccan shrews >38 K yr ago. This study illustrates that multilocus genetic analyses can facilitate the interpretation of species' evolutionary history but that phylogeographic inference using X and Y chromosomes is restricted by low levels of observed polymorphism.     

Geographic variation of the Greater white-toothed shrew (Crocidura russula Hermann, 1780 Mammalia, Soricidae)

A multivariate morphometric study of the Greater white-toothed shrew ( C. russula ) throughout its Palearctic range was carried out to search for patterns of geographic variation within the species boundary. Burnaby's and multiple group principal component analysis allowed the adjustment of raw data with respect to within-sample allometric variation. Multivariate 'size-free'results show a stepped cline with the phenotypical trait reduction and shape change from the eastern to the western Maghreb. Pleistocene fossil mandibles proved to have low phenetic distances with eastern populations (Tunisia, east Algeria) and it is argued that their character set is the primitive condition. The ancestral Mid-Pleistocene shrews lived in a relatively more humid climate. Geo-climatic changes in the north African range during the Quaternary provoked phenetic variation of C. russula and, it can be argued, evolution of the modern western C.r. yebalensis. A historical process can thus be assumed as the main cause of this categorical variation, by segmentation of the species range due to geo-climatic events. Morphometric discontinuity within the C. russula Maghreb range is shown to be congruent with karyological and biochemical studies. Moroccan and Tunisian shrews differ, for example, in NFa chromosomes and electrophoretical traits. A stasipatric process should be invoked to explain categorical variation in the Maghreb range. Colonization and divergence of insular populations results in more or less differentiated geographic races. The populations of Ibiza and Pantelleria are close to the species threshold (Nei's D ≥ 0.1). The process of speciation undergone by the Greater white-toothed shrew results in a complex pattern of geographic variation, including both allopatric and non-allopatric modes.     

Additive effects of climate and vegetation structure on the altitudinal distribution of greater white-toothed shrews Crocidura russula in a Mediterranean mountain range

Environmental variation along geographic gradients determines the distribution of animals and plants trough both direct and indirect effects. We analyze the relative contribution of climate and vegetation structure variations along a Mediterranean altitude gradient on the patterns of abundance and occurrence of the greater white-toothed shrew Crocidura russula, a generalist small mammal whose distribution is constrained by cold conditions. Sampling was performed from February 1995 to July 1997 on nine plots covering a wide altitudinal gradient (540–1550 m?a.s.l.). Structural equation models for the direct and indirect effects of climate and vegetation on shrew density and occupancy rates showed a stronger effect of vegetation structure (53 %) than direct climate effects (38 %) on shrew distribution. Shrews were more abundant in the warmer lowland sites, but were able to colonize highland cold areas by selecting habitats with well-developed understory vegetation (low shrubs or bracken). Vegetation effects were additive to climatic restrictions, seemingly providing more favorable microclimatic, and presumably food, conditions under shrub cover. Results indicate that predictions of range changes for shrews under climate change scenarios may underestimate expected range expansions under the current conditions of general land abandonment and shrub encroachment.     

Range expansion of the greater white-toothed shrewCrocidura russula in Switzerland results in local extinction of the bicoloured white-toothed shrewC. leucodon

The distribution limits ofCrocidura russula (Hermann, 1780) andC. leucodon (Hermann, 1780) were investigated during an interval of 25 years in the bottom of the Rhone valley above Lake Geneva, Switzerland (total data set: 105 spatio-temporal occurrences, 1137 shrews). In 1975, the contact zone between the two species was situated in the region of Martigny. In 1999/2000, new sampling revealed three results: (1) The contact zone showed an upward shift of about 25 km. (2) In the expanded range ofC. russula, the resident species has totally disappeared (confirmed by owl pellets analysis). (3) This demonstrates a dominance ofC. russula overC. leucodon. Three hypotheses which may explain the range expansion ofC. russula were evaluated: (1) habitat modification favouring linear dispersal due to the construction of a highway; (2) temporal event favoured by climate fluctuations, or (3) ongoing postglacial colonisation of Europe. Hypothesis 1 was rejected, because the progression of the shrews anticipated the construction. Hypothesis 3 received only weak support because range limits ofC. russula in the region of Nice have been stable for thousands of years. Therefore hypothesis 2, admitting that ongoing climate change has facilitated range expansion, is the most probable.     

Helminth associations in white-toothed shrews Crocidura russula (Insectivora: Soricidae) from the Albufera Natural Park, Spain

The helminths of 218 white-toothed shrews from 29 sites in 2 biotopes in the Albufera Natural Park (Valencia, Spain) were examined from July 1990 to August 1991. An association analysis of helminths occurring at a prevalence of more than 4% was carried out for 4 species of cestodes located in the intestine (Hymenolepis pistillum, H. scalaris, H. tiara, and Pseudhymenolepis redonica) and 3 species of nematodes (Pseudophysaloptera sp. located in the stomach, Stammerinema rhopocephala larvae in the intestine and abdominal cavity, and Porrocaecum sp. in the thoracic and abdominal cavities). Bivariate (species pairs) versus multivariate analyses (associations within the entire set of species) were performed of presence-absence and of quantitative records (influence of intensity on associations). The associations were evaluated with respect to the sex and age of the host and to the sampling date and sites. The host and environment played a limited role, and the major determinant of species assemblage was phylogenetic. Positive associations were found among both the cestodes and the nematodes, whereas negative associations were found between cestodes and nematodes. The type of life cycle was probably the second greatest determinant of species associations. Nematodes using shrews as a paratenic host or as their definitive host were both positively associated.     

Ultrastructural studies on oogenesis in the shrew (Crocidura russula): I. The preantral follicle

Ultrastructural analysis of the ovarian follicle prior to antrum formation in the shrew, Crocidura russula, shows gradual differentiation of mitochondria, endoplasmic reticulum, Golgi complexes, vesicles, and multivesicular bodies correlated with the growth of the oocyte from primordial to tertiary follicle and the development of the follicular wall. The growth rate of the follicle in relation to that of the oocyte was found to be biphasic.     

Morphometric geographic variation in the Asian lesser white-toothed shrew Crocidura shantungensis (Mammalia,Insectivora) in East Asia

This study investigated geographic variation in the Asian lesser white-toothed shrew, Crocidura shantungensis Miller, 1901, among a number of localities in East Asia, including the Korean Peninsula, Tsushima Island, Cheju Island, and Taiwan, using univariate and multivariate analyses of cranial morphometric characters. Sexual differences were found in Taiwanese shrews, but were not evident in those from Korea or Tsushima. The samples from Taiwan had a larger overall cranial size than those from Korea and Tsushima, and were well separated from the latter two samples in the discriminant analyses. The overall size of the Cheju shrews was also greater, but lay within the variation of those from Taiwan. Crocidura dsinezumi quelpartis Kuroda, 1934, described from Cheju Island, is actually a junior synonym of C. shantungensis, and is not related to C. dsinezumi.     

Molecular and morphological insights into the origin of the invasive greater white-toothed shrew (<Emphasis Type="Italic">Crocidura russula</Emphasis>) in Ireland

Identifying routes of invasion is a critical management strategy in controlling the spread of invasive species. This is challenging however in the absence of direct evidence. Therefore, indirect methodologies are used to infer possible invasion sources and routes, such as comparisons of genetic and morphological data from populations from invasive ranges and putative source areas. The greater white-toothed shrew (Crocidura russula) was first discovered in Ireland from skeletal remains in the pellets of birds of prey collected in 2007 and is it is now sufficiently established that the species has a detrimental impact on Ireland’s small mammal community. In this study, we address the uncertain origin(s) of the Irish population of C. russula. The cytochrome b gene of mitochondrial DNA was analysed from 143 individuals from throughout its range within a phylogenetic and approximate Bayesian computation framework. These analyses revealed that the Irish population stemmed from Europe as opposed to North Africa. Additionally, mandibles from 523 individuals from Ireland and 28 other European populations were subjected to multivariate and distance-based analyses, which demonstrated an association between the Irish population and those in France, Switzerland and Belgium. When the genetic and morphological analyses were considered together, an origin stemming from France was deemed the most likely scenario for the source of the invasive Irish population. This study has demonstrated the importance of utilising a multidisciplinary approach when attempting to identify the origins and invasion routes of invasive species.     

Mediterranean populations of the lesser white-toothed shrew (Crocidura suaveolens group): an unexpected puzzle of Pleistocene survivors and prehistoric introductions

An earlier study revealed the strong phylogeographical structure of the lesser white-toothed shrew (Crocidura suaveolens group) within the northern Palaearctic. Here, we aim to reconstruct the colonization history of Mediterranean islands and to clarify the biogeography and phylogeographical relationships of the poorly documented Middle East region with the northern Palaearctic. We performed analyses on 998-bp-long haplotypes of the mitochondrial cytochrome b gene of 143 samples collected around the Mediterranean basin, including islands and the Middle East. The analyses suggest that the Cypriot shrew belongs to the rare group of relict insular Pleistocene mammal taxa that have survived to the present day. In contrast, the Cretan, Corsican and Menorcan populations were independently introduced from the Middle East during the Holocene. The phylogeographical structure of this temperate Palaearctic species within the Middle East appears to be complex and rich in diversity, probably reflecting fragmentation of the area by numerous mountain chains. Four deeply divergent clades of the C. suaveolens group occur in the area, meaning that a hypothetical contact zone remains to be located in central western Iran.     

Skewed reproductive success among male white-spotted charr land-locked by an erosion control dam: Implications for effective population size

In 1996 and 1997, the spawning behavior of fluvial white-spotted charr Salvelinus leucomaenis, was observed in the upstream area of an erosion control dam. A small number of males with relatively large body size mated successfully with females as a pair, while almost all satellite males did not sneak successfully, resulting in a non-random mating system. The low sneaking success of subordinate males, in addition to the monopolization of spawning opportunities by a few dominant males, is one of the most important causes of skewed reproductive success among males. The total number of adult fishes in the study area (N: approximately half of the whole tributary above a dam) was estimated as 148 and 102 in 1996 and 1997, respectively. Based on these findings and some further assumptions, the estimated effective population size (N_e) was low in both years. The N_e/N ratio ranged from 0.33 to 0.36 in both years. In addition to reduced population size by construction of an impassable dam, the above-dam population suffered low N_e due to skewed reproductive success among males. The low N_e may be one cause of extinction in above-dam populations of fluvial charr, especially just after the construction of impassable barriers.