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Mt Kilimanjaro is Africa's highest mountain, and an icon for a country famous for its mammalian fauna. The distribution and abundance of small mammals on the mountain are poorly known. Here we document the distribution of shrews and rodents along an elevational gradient on the southeastern versant of Kilimanjaro. Five sites were sampled with elevational center points of 2000, 2500, 3000, 3500 and 4000 m, using a systematic methodology of standard traps and pitfall lines, to inventory the shrews and rodents of the slope. Sixteen species of mammal were recorded, including 6 shrew and 10 rodent species, and the greatest diversity of both was found at 3000 m, the elevational midpoint of the transect. No species previously unrecorded on Kilimanjaro were observed. Two genera of rodents that occur in nearby mountains (Hylomyscus and Beamys) were not recorded. Myosorex zinki, the only mammal endemic to Mt. Kilimanjaro, which previously was known by only a few specimens collected in the ericaceous or moorland habitat, was found in all but one (the lowest) of the sites sampled, and was one of the most widespread species of small mammal along the gradient. Two shrews (Crocidura allex and Sylvisorex granti) and one rodent (Dendromus insignis) were found throughout the entire transect, with Dendromus being observed at our highest trap point (4240 m). As in similar faunal surveys on other mountains of Tanzania, rainfall influenced the sample success of shrews, but not rodents. Trap success for rodents at 3500 m was notably low. This study contributes further justification for the conservation of the forest habitat of Mt. Kilimanjaro.

The extraction of museum DNA from a unique collection of samples of the “Crocidura pergrisea” species complex, which comprises local endemics of Central and West Asia, allowed us to determine their inter- and intragroup relationships. The first step of this study was the re-evaluation of heavily damaged type specimens of C. armenica via a microcomputed-tomography-based cybertaxonomic approach (CTtax), which enabled a precise description of the species’ morphology; three-dimensional models of the cybertypes were made available through the MorphoBank Repository. We developed the “AProMaDesU” pipeline on the basis of five requirements for micro-CT-based cyber-datasets in relation to mammalian collections. Our second step was a combination of several meticulous approaches to morphological investigation against a background of a cytb-based phylogeny, which helped us to make a taxonomic decision about the status of species of the “pergrisea” group, e.g., C. arispa, C. armenica, and C. serezkyensis, when the morphological results were partly incongruent with the molecular phylogeny. Nevertheless, under two assumptions, our findings preserved a separate species-level status of C. serezkyensis and C. arispa. In addition, we restored the species-level status of C. armenica. This taxonomic decision is based on our morphospace analysis, which revealed unique craniomandibular shape transformations within the rocky shrews that helped them with the transition to a new area of morphospace/trophic niches and consequently separated them from the other analyzed Crocidura groups.

Conventional karyotypes of Crocidura tanakae and C. attenuata collected from mainland China were examined by bone marrow and cell culture methods along with identification to species using molecular data. In contrast to C. tanakae from Taiwan, populations from mainland China were karyotypically polymorphic, varying in diploid number (2n) from 24 to 40 and fundamental number (FN) from 45 to 56. Autosomes are composed of 6–20 metacentric or submetacentric chromosomes, 0–8 subtelocentric chromosomes, and 0–24 telocentric chromosomes. These polymorphic karyotypes indicate the presence of two geographically separated races. The ancestral karyotype of C. tanakae potentially is the same as currently found on Taiwan: 2n = 40 and FN = 56. In contrast to C. tanakae, C. attenuata had a uniform karyotype (2n = 40, FN = 54) in mainland China, suggesting that previously reported karyotypic polymorphism in this species was an error due to the morphological crypsis between C. tanakae and C. attenuata in mainland China. The intricate karyotypic polymorphism of C. tanakae may be indicative of this species undergoing speciation.

Many RNA viruses arise from animal reservoirs, namely bats, rodents and insectivores but mechanisms of virus maintenance and transmission still need to be addressed. The bicolored white-toothed shrew (Crocidura leucodon) has recently been identified as reservoir of the neurotropic Borna disease virus 1 (BoDV-1). Six out of eleven wild living bicoloured white-toothed shrews were trapped and revealed to be naturally infected with BoDV-1. All shrews were monitored in captivity in a long-term study over a time period up to 600 days that differed between the individual shrews. Interestingly, all six animals showed an asymptomatic course of infection despite virus shedding via various routes indicating a highly adapted host-pathogen interaction. Infectious virus and viral RNA were demonstrated in saliva, urine, skin swabs, lacrimal fluid and faeces, both during the first 8 weeks of the investigation period and for long time shedding after more than 250 days in captivity. The various ways of shedding ensure successful virus maintenance in the reservoir population but also transmission to accidental hosts such as horses and sheep. Naturally BoDV-1-infected living shrews serve as excellent tool to unravel host and pathogen factors responsible for persistent viral co-existence in reservoir species while maintaining their physiological integrity despite high viral load in many organ systems.

Paramyxoviruses, negative-sense single-stranded RNA viruses, pose a critical threat to human public health. Currently, 78 species, 17 genera, and 4 subfamilies of paramyxoviruses are harbored by multiple natural reservoirs, including rodents, bats, birds, reptiles, and fish. Henipaviruses are critical zoonotic pathogens that cause severe acute respiratory distress and neurological diseases in humans. Using reverse transcription-polymerase chain reaction, 115 Crocidura species individuals were examined for the prevalence of paramyxovirus infections. Paramyxovirus RNA was observed in 26 (22.6%) shrews collected at five trapping sites, Republic of Korea. Herein, we report two genetically distinct novel paramyxoviruses (genus: Henipavirus): Gamak virus (GAKV) and Daeryong virus (DARV) isolated from C. lasiura and C. shantungensis, respectively. Two GAKVs and one DARV were nearly completely sequenced using next-generation sequencing. GAKV and DARV contain six genes (3′-N-P-M-F-G-L-5′) with genome sizes of 18,460 nucleotides and 19,471 nucleotides, respectively. The phylogenetic inference demonstrated that GAKV and DARV form independent genetic lineages of Henipavirus in Crocidura species. GAKV-infected human lung epithelial cells elicited the induction of type I/III interferons, interferon-stimulated genes, and proinflammatory cytokines. In conclusion, this study contributes further understandings of the molecular prevalence, genetic characteristics and diversity, and zoonotic potential of novel paramyxoviruses in shrews.

Three species of white-toothed shrews of the order Eulipotyphla are present in central Europe: the bicolored (Crocidura leucodon), greater (Crocidura russula) and lesser (Crocidura suaveolens) white-toothed shrews. Their precise distribution in Germany is ill-defined and little is known about them as reservoirs for zoonotic pathogens (Leptospira spp., Coxiella burnetii, Brucella spp., Anaplasma phagocytophilum, Babesia spp., Neoehrlichia mikurensis and Bartonella spp.). We investigated 372 Crocidura spp. from Germany (n = 341), Austria (n = 18), Luxembourg (n = 2) and Slovakia (n = 11). West European hedgehogs (Erinaceus europaeus) were added to compare the presence of pathogens in co-occurring insectivores. Crocidura russula were distributed mainly in western and C. suaveolens mainly in north-eastern Germany. Crocidura leucodon occurred in overlapping ranges with the other shrews. Leptospira spp. DNA was detected in 28/227 C. russula and 2/78 C. leucodon samples. Further characterization revealed that Leptospira kirschneri had a sequence type (ST) 100. Neoehrlichia mikurensis DNA was detected in spleen tissue from 2/213 C. russula samples. Hedgehogs carried DNA from L. kirschneri (ST 100), L. interrogans (ST 24), A. phagocytophilum and two Bartonella species. This study improves the knowledge of the current distribution of Crocidura shrews and identifies C. russula as carrier of Leptospira kirschneri. However, shrews seem to play little-to-no role in the circulation of the arthropod-borne pathogens investigated.

The Fingui white-toothed shrew Crocidura fingui, categorized as Data Deficient on the IUCN Red List, is endemic to Príncipe Island, Central Africa. It is currently known from < 10 sites and its ecology and any threats are largely unknown. We review its distribution and ecology, and any potential threats, based on camera-trap data, field observations and previously published and unpublished sightings. Crocidura fingui appears to have an island-wide distribution and to use a range of ecosystems, from native forests to urban areas. It is largely nocturnal and exhibits considerable spatiotemporal overlap with introduced mammals such as the African civet Civettictis civetta, domestic cat Felis catus and rats Rattus spp. The species has an area of occupancy < 500 km2 and is known from fewer than five locations, and its population is inferred to be affected by several introduced mammals. We recommend that it is recategorized as Endangered on the IUCN Red List.

Many peninsulas in the temperate zone played an important role as refugia of various flora and fauna, and the southern Korean Peninsula also served as a refugium for many small mammals in East Asia during the Pleistocene. The Asian lesser white-toothed shrew, Crocidura shantungensis, is a widely distributed species in East Asia, and is an appropriate model organism for exploring the role of the Korean Peninsula as a refugium of small mammals. Here, we investigated phylogenetic relationships and genetic diversity based on the entire sequence of the mitochondrial cytochrome b gene (1140 bp). A Bayesian tree for 98 haplotypes detected in 228 C. shantungensis specimens from East Asia revealed the presence of three major groups with at least 5 subgroups. Most haplotypes were distributed according to their geographic proximity. Pairwise FST’s and analysis of molecular variance (AMOVA) revealed a high degree of genetic differentiation and variance among regions as well as among populations within region, implying little gene flow among local populations. Genetic evidence from South Korean islands, Jeju-do Island of South Korea, and Taiwan leads us to reject the hypothesis of recent population expansion. We observed unique island-type genetic characteristics consistent with geographic isolation and resultant genetic drift. Phylogeographic inference, together with estimates of genetic differentiation and diversity, suggest that the southern most part the Korean Peninsula, including offshore islands, played an important role as a refugium for C. shantungensis during the Pleistocene. However, the presence of several refugia on the mainland of northeast Asia is also proposed.

Background Ixodid ticks are important vectors for zoonotic pathogens, with Ixodes ricinus being the most important in Europe. Rodents are hosts of immature life stages of I. ricinus ticks and are considered main reservoirs for tick-borne pathogens, e.g. Borrelia burgdorferi . The aim of this study was to analyse the prevalence as well as genospecies and sequence type (ST) diversity of Borrelia burgdorferi sensu lato in ticks and small mammals from central Germany and to elaborate on the influence of environmental and/or individual host and vector factors on Borrelia prevalence. Methods After species identification, 1167 small mammal skin samples and 1094 ticks from vegetation were screened by B. burgdorferi sensu lato real-time polymerase chain reaction, and positive samples were characterized by multilocus sequence typing. Generalized linear (mixed) models were used to estimate how seasonality, small mammal species/tick life stage and habitat affect individual infection status. Results In total, 10 small mammal species and three tick species, Ixodes ricinus , Ixodes inopinatus (both considered members of the I. ricinus complex) and Dermacentor reticulatus , were investigated. Borrelia DNA was detected in eight host species, i.e. the striped field mouse ( Apodemus agrarius ), the yellow-necked field mouse ( Apodemus flavicollis ), the wood mouse ( Apodemus sylvaticus ), the water vole ( Arvicola amphibius ), the bank vole ( Clethrionomys glareolus ), the field vole ( Microtus agrestis ), the common vole ( Microtus arvalis ), and the common shrew ( Sorex araneus ). Two species were Borrelia negative, the greater white-toothed shrew ( Crocidura russula ) and the pygmy shrew ( Sorex minutus ). The average prevalence was 6.2%, with two genospecies detected, Borrelia afzelii and Borrelia garinii , and at least three STs that had not been previously reported in small mammals. Borrelia prevalence in small mammals did not differ between seasons. Six genospecies of Borrelia —Borrelia afzelii , Borrelia valaisiana , Borrelia garinii , Borrelia lusitaniae , Borrelia spielmanii , and Borrelia burgdorferi sensu stricto—and 25 STs of Borrelia , of which 12 have not been previously described at all and five have not been previously reported in Germany, were detected in 13% of I. ricinus complex ticks. Prevalence was highest in adult females (25.3%) and lowest in nymphs (11.4%). Prevalence was significantly higher in ticks from grassland (16.8%) compared to forests (11.4%). Conclusions The high level of small mammal diversity in this region of Germany seems to be reflected in a wide variety of genospecies and STs of B. burgdorferi . Graphical abstract

Borna disease (BD) is a sporadic neurologic disease of horses and sheep caused by mammalian Borna disease virus (BDV). Its unique epidemiological features include: limited occurrence in certain endemic regions of central Europe, yearly varying disease peaks, and a seasonal pattern with higher disease frequencies in spring and a disease nadir in autumn. It is most probably not directly transmitted between horses and sheep. All these features led to the assumption that an indigenous virus reservoir of BDV other than horses and sheep may exist. The search for such a reservoir had been unsuccessful until a few years ago five BDV-infected shrews were found in a BD-endemic area in Switzerland. So far, these data lacked further confirmation. We therefore initiated a study in shrews in endemic areas of Germany. Within five years 107 shrews of five different species were collected. BDV infections were identified in 14 individuals of the species bicolored white-toothed shrew (Crocidura leucodon, HERMANN 1780), all originating from BD-endemic territories. Immunohistological analysis showed widespread distribution of BDV antigen both in the nervous system and in epithelial and mesenchymal tissues without pathological alterations. Large amounts of virus, demonstrated by presence of viral antigen in epithelial cells of the oral cavity and in keratinocytes of the skin, may be a source of infection for natural and spill-over hosts. Genetic analyses reflected a close relationship of the BDV sequences obtained from the shrews with the regional BDV cluster. At one location a high percentage of BDV-positive shrews was identified in four consecutive years, which points towards a self-sustaining infection cycle in bicolored white-toothed shrews. Analyses of behavioral and population features of this shrew species revealed that the bicolored white-toothed shrew may indeed play an important role as an indigenous host of BDV.