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.     

Skeletal morphology of the forefoot in shrews (Mammalia: Soricidae) of the genus Cryptotis, as revealed by digital X-rays

Variation in the forefoot skeleton of small-eared shrews (family Soricidae, genus Cryptotis) has been previously documented, but the paucity of available skeletons for most taxa makes assessment of the degrees of intraspecific and interspecific variation difficult. We used a digital X-ray system to extract images of the forefoot skeleton from 101 dried skins of eight taxa (seven species, including two subspecies of one species) of these shrews. Lengths and widths of each of the four bones of digit III were measured directly from the digital images, and we used these data to quantify variation within and among taxa. Analysis of the images and measurements showed that interspecific variation exceeds intraspecific variation. In fact, most taxa could be distinguished in multivariate and some bivariate plots. Our quantitative data helped us define a number of specific forefoot characters that we subsequently used to hypothesize evolutionary relationships among the taxa using the exhaustive search option in PAUP, a computer program for phylogenetic analysis. The resulting trees generally concur with previously published evolutionary hypotheses for small-eared shrews. Cryptotis meridensis, a taxon not previously examined in recent phylogenies, is rooted at the base of the branch leading to the C. mexicana group of species. The position of this species suggests that the mostly South American C. thomasi group shares an early ancestor with the C. mexicana group.     

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.     

Systematics of white-toothed shrews (Crocidura) (Mammalia: Insectivora: Soricidae) of Taiwan: Karyological and morphological Studies

We define the species boundaries of white-toothed shrews (genus Crocidura ) in Taiwan using karyological and morphological characteristics. Ninety-nine animals were obtained from all over Taiwan at capture rates usually less than 10%. Three species are recognized by distinct cytotypes: Crocidura attenuata tanakae 2n = 40, FN = 56; Crocidura suaveolens hosletti 2n = 40, FN = 50; Crocidura kurodai 2n = 40, FN = 54. A suite of six morphological characters diagnose the three species: shape of skull, position of incisive foramina, shape of fourth upper premolar, shape of pinna, tail vibrissae, and foot pads. A species key and notes on the life history of each species are provided. Finally, we discuss chromosomal evolution and biolgeography of Crocidura in East and South East Asia.     

Molecular taxonomy of Crocidura species (Eulipotyphla: Soricidae) in a key biogeographical region for African shrews,Nigeria

The taxonomy of African shrew species is still unresolved due to their conserved morphology. This also affects knowledge concerning their geographic distribution. In Nigeria, using mitochondrial Cytochrome b gene sequences, we carried out a survey for shrews from the genus Crocidura across various ecological zones to determine taxa that are present and also to assess their phylogeographic structure. Our analyses include 183 specimens collected with Sherman traps from 19 localities around the country. We detected six taxa: Crocidura olivieri lineages II, III and IV, C. hildegardeae, C. jouvenetae, and C. foxi. Among these, C. hildegardeae and C. jouvenetae are reported in Nigeria for the first time. Phylogenetic comparison of our genetic sequences to those generated from other parts of Africa demonstrate that all species in our study, as currently defined, are in need of taxonomic revision. Geographically, Nigeria seems to represent the easternmost boundary for C. olivieri lineage II and C. jouvenetae, and the western distribution limit of C. olivieri lineage IV and C. hildegardeae. The Niger River appears to be the most significant topographical barrier restricting these taxa. This information is vital to preserving the diversity but also managing the epidemiological potential of these small mammals.     

Functional skeletal morphology and its implications for locomotory behavior among three genera of myosoricine shrews (Mammalia: Eulipotyphla: Soricidae)

Myosoricinae is a small clade of shrews (Mammalia, Eulipotyphla, Soricidae) that is currently restricted to the African continent. Individual species have limited distributions that are often associated with higher elevations. Although the majority of species in the subfamily are considered ambulatory in their locomotory behavior, species of the myosoricine genus Surdisorex are known to be semifossorial. To better characterize variation in locomotory behaviors among myosoricines, we calculated 32 morphological indices from skeletal measurements from nine species representing all three genera that comprise the subfamily (i.e., Congosorex, Myosorex, Surdisorex) and compared them to indices calculated for two species with well‐documented locomotory behaviors: the ambulatory talpid Uropsilus soricipes and the semifossorial talpid Neurotrichus gibbsii. We summarized the 22 most complete morphological variables by 1) calculating a mean percentile rank for each species and 2) using the first principal component from principal component analysis of the indices. The two methods yielded similar results and indicate grades of adaptations reflecting a range of potential locomotory behaviors from ambulatory to semifossorial that exceeds the range represented by the two talpids. Morphological variation reflecting grades of increased semifossoriality among myosoricine shrews is similar in many respects to that seen for soricines, but some features are unique to the Myosoricinae. J. Morphol. 276:550–563, 2015. © 2015 Wiley Periodicals, Inc.     

White-toothed shrews (Mammalia, Soricomorpha, Crocidura) of coastal islands of Vietnam

New findings of the white-toothed shrews (Crocidura spp.) from offshore islands of Vietnam are reported. The species identifications have been confirmed by the analysis of complete mitochondrial cytochrome b gene (1140 bp). Crocidura phuquocensis is the only species found in the Phu Quoc Island. Crocidura fuliginosa has been recorded from two islands of the Con Dao Archipelago (Con Son and Bai Canh). The occurrence of Crocidura fuliginosa in Vietnam has been genetically confirmed for the first time. Crocidura attenuata has been collected from the Cat Ba Island for the first time, and this finding corresponds well with the proposal that the species' distribution is confined to the north and east of the Red River only.     

河南信阳发现台湾灰麝鼩北大核心CSCD

于2013年和2018年在河南信阳新县与浉河共采集到3号鼩鼱成年个体标本,基于齿色特征初步鉴定为劳亚食虫目鼩鼱科麝鼩属(Crocidura)物种。对所采3号标本的Cyt b基因进行PCR扩增和测序,测序结果与GenBank数据库进行比对,并通过构建系统发育关系树进一步明确所采3只标本的分类地位。根据标本形态、头骨特征,结合分子生物学证据,确定该物种为台湾灰麝鼩(C.tanakae),为河南哺乳动物分布新记录种。标本现存于郑州大学生物多样性与生态学研究所(编号分别为IBE20131005、IBE2018100501和IBE2018100502)。     

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.     

安徽黄山和宣城发现台湾灰麝鼩

2018年11月在安徽省黄山市太平湖国家湿地公园(30°34′42″N,118°41′47″E)和宣城市旌德县蔡家桥镇(30°21′26″N,118°30′11″E)的针阔混交林采集到3只麝鼩属小型兽类。3只个体体型较小,体重11～14 g;背毛灰褐色,腹部毛色稍淡;尾短而粗壮,上下异色,基部2/3散生稀疏的长刚毛,这些特征与台湾灰麝鼩(Crocidura tanakae)描述一致。基于Cyt b全序列构建的最大似然树显示,这3号标本与从GenBank下载的台湾灰麝鼩(GenBank登录号KX946002～KX946006、AB175080、AB175081)构成单系群(支持率为100%)。这3号标本Cyt b全序列与台湾灰麝鼩地模标本(GenBank登录号AB175080、AB175081)的遗传距离在0.76%～0.85%之间。基于以上结果,确定这3号标本为台湾灰麝鼩,这是该物种在安徽省的首次发现,也是其在华东地区的首个分布记录。     

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.     

Molecular phylogenetics of shrews (Mammalia: Soricidae) reveal timing of transcontinental colonizations

We sequenced 2167 base pairs (bp) of mitochondrial DNA cytochrome b and 16S, and 1390 bp of nuclear genes BRCA1 and ApoB in shrews taxa (Eulipotyphla, family Soricidae). The aim was to study the relationships at higher taxonomic levels within this family, and in particular the position of difficult clades such as Anourosorex and Myosorex. The data confirmed two monophyletic subfamilies, Soricinae and Crocidurinae. In the former, the tribes Anourosoricini, Blarinini, Nectogalini, Notiosoricini, and Soricini were supported. The latter was formed by the tribes Myosoricini and Crocidurini. The genus Suncus appeared to be paraphyletic and included Sylvisorex. We further suggest a biogeographical hypothesis, which shows that North America was colonized by three independent lineages of Soricinae during middle Miocene. Our hypothesis is congruent with the first fossil records for these taxa. Using molecular dating, the first exchanges between Africa and Eurasia occurred during the middle Miocene. The last one took place in the Late Miocene, with the dispersion of the genus Crocidura through the old world.     

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.     

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.     

Mitochondrial simple sequence repeats and 12S-rRNA gene reveal two distinct lineages of Crocidura russula (Mammalia, Soricidae)

A short segment (135 bp) of the control region and a partial sequence (394 bp) of the 12S-rRNA gene in the mitochondrial DNA of Crocidura russula were analyzed in order to test a previous hypothesis regarding the presence of a gene flow disruption in northern Africa. This breakpoint would have separated northeast-African C. russula populations from the European (plus the northwest-African) populations. The analysis was carried out on specimens from Tunisia (C. r. cf agilis), Sardinia (C. r. ichnusae), and Pantelleria (C. r. cossyrensis), and on C. r. russula from Spain and Belgium. Two C. russula lineages were identified; they both shared R2 tandem repeated motifs of the same length (12 bp), but not the same primary structure. These simple sequence repeats were present in 12-23 copies in the right domain of the control region. Within the northeast-African populations, a polymorphism of repeat variants, not yet found in Europe, was recorded. A neighbor-join tree, which was built by sequences of the conserved 12S-rRNA gene, separated the two sister groups; it permitted us to date a divergence time of 0.5 Myr. Our data discriminated two different mitochondrial lineages in accordance with the previous morphological and karyological data. Ecoclimatic barriers formed during the Middle Pleistocene broke the range of ancestral species in the Eastern Algeria (Kabile Mountains), leading to two genetically separate and modern lineages. The northeast-African lineage can today be located in Tunisia, Pantelleria, and Sardinia. The northwest-African lineage (Morocco and West Algeria), reaching Spain by anthropogenic introduction, spread over north Europe in modern times. The Palaearctic C. russula species is monophyletic, but a taxonomical revision (ie, to provide a full species rank for the northeast taxa and to put in synonymy some insular taxa) is required.     

Phylogeny,phylogeography and geographical variation in the Crocidura monax (Soricidae) species complex from the montane islands of Tanzania,with descriptions of three new species

We assess morphological and multilocus genetic variation among 11 isolated montane populations of white‐toothed shrews from Tanzania that have been referred to either Crocidura monax Thomas or C. montis Thomas. The montane sites we sampled represent ‘sky‐islands’ from two geologically distinct archipelagos (Northern Highlands and the Eastern Arc Mountains) and are a significant component of the Eastern Afromontane Biodiversity Hotspot. We used multivariate analyses of morphometric traits and phylogenetic and species‐delimitation analyses of multilocus DNA sequence data to assess species‐level diversity. Our species delimitation analyses included a novel, pairwise validation approach that avoids potential biases associated with specifying a guide tree. These analyses reveal several distinct lineages, which we treat as six allopatric species. Each species is restricted to one, two or four mountains. We use available names to recognize C. monax, C. tansaniana Hutterer and C. usambarae Dippenaar, while naming and describing three new species. Our results demonstrate the effectiveness of combining morphological and genetic data to uncover and describe hidden diversity in a cryptic mammalian system. © 2015 The Linnean Society of London