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This paper presents a review of host specificity among cestodes collected from Soricidae mammals in Europe and a revision of cestodes preserved in formalin in the collection of the East-Slovakian Museum in Kosice. The morphological re-examination of cestodes showed that cestodes found in non-specific hosts (Soricidae and Talpidae) were identified incorrectly. The majority of the redescribed species are specific to a particular host genus. Only one species of the Staphylocystis Villot, 1877 genus, which could be Staphylocystis brusatae (Vaucher, 1971) according to the number, shape and length of hooks, as originally described from Crocidura suaveolens (Pallas) from Switzerland, was found in a host of the genus Sorex Linnaeus. The host specificity of the majority of Hymenolepididae cestode species is stenoxenous, but this study of the collection and the critical review of cestodes from non-specific Soricidae hosts disclosed that representatives of the genus Staphylocystis are euryxenous. So, these cestodes are better adapted to spreading into new ecosystems. A new species from Neomys fodiens (Pennant) from Slovakia, Western Carpathian, belonging to the genus Coronacanthus Spassky, 1954, Coronacanthus longicirrosusn. sp., is described. The new species is easily distinguishable by the number of small rostellar hooks (12-18, 4-5 mu m), hooks shape, long cirrus (70-86 mu m) and the presence of the vaginal sphincter.

The water shrew species Neomys fodiens and Neomys milleri are morphologically similar and, based on phenotype, it is difficult to distinguish them. Mandibles of both species were studied by means of classic as well as geometric morphometrics. We compared a priori identified mandibles of water shrews captured in two areas where the species ranges partially or entirely overlap, i.e., in Germany and in the south-eastern part of the Balkan Peninsula. Discrimination of the two species by linear measurements was successful in Germany but not in the Balkans. Discriminant analysis revealed clear differences in the shape of the mandible between the species in both regions. Comparison of allopatric and sympatric populations of both species showed that species are smaller and more similar in shape where they occur in sympatry. Our results suggest that geometric morphometrics can be successfully used for species determination, especially in regions where determination by size is not possible.

Soricid (Lipotyphla: Soricidae) inner tooth structures have been described for the first time using high-resolution X-ray computed micro-tomography. The endodontic morphology of shrews is complex and according to our results can be used (i) for the description of additional odontological features, (ii) for phylogenetic investigations on a morphological basis, and (iii) for determining the homology of the tooth elements. The results of the original analysis revealed a gradual process of dentine formation and, correspondingly, decreasing pulp volume in immature and mature specimens of Sorex mirabilis. However, analysis of the age-related changes also revealed an irregularity in the dentine deposition within different parts of a pulp cavity. We named this irregularity “selective pulp overgrowth”. This process was confirmed by the analyses of specimens of Sorex minutissimus, Neomys fodiens and Blarina brevicauda. Attempts to explain selective overgrowth led us to consider the functions of tooth pulp. According to dentistry investigations, one of the several functions of pulp is dentine innervation. We assume that dentine innervation is a significant feature of soricid teeth. A list of twelve characters displays the possible application of pulp endocast morphology to phylogenetic investigations. We suggest that pulp endocast analysis be added to the analysis of the tooth surface to achieve the broadest and most productive use of tooth morphology.

One of the less studied positive interactions among organisms is facilitation. Facilitation may have significant impact on diversity of species especially in low productive environments. We studied the effects of well-known facilitator and ecosystem engineer, the North American beaver (Castor canadensis), on small mammal assemblages in the boreal zone. The small mammals, namely voles, mice, and shrews, were trapped over 2 years in ten beaver-modified habitats and in ten control sites. Contrary to our expectations, we did not observe any differences between species or individual numbers between beaver-modified and control sites. However, there were differences in species composition between sites; grass-eating field voles (Microtus agrestis) and invertebrate-eating shrews (Sorex araneus,Neomys fodiens) tended to be more common in beaver sites while forest-dwelling wood lemmings (Myopus schisticolor) and yellow-necked mice (Apodemus flavicollis) were only captured in control sites. The most common species in both habitats was the bank vole (Myodes glareolus), which is a generalist in its habitat requirements. The bank vole’s population structures were similar between the two habitat types. The actions of beavers in water bodies within boreal forests seem to have no effect on the small mammal diversity and their numbers at the regional scale but may have positive effect on them at the larger landscape level as beavers are increasing the overall habitat diversity in the landscape.

Based on numerous literary sources, the thesis reviews current knowledge of history and range dynamics of the genus Neomys, an indexing element of the west Palearctic mammal fauna. It provides a list of fossil records attributed to the extant genus Neomys and extinct clades composing the European records of tribe Nectogalini, and a survey of hypotheses on their phylogenetic relations. Particular attention is given to the genus Asoriculus which is generally considered as an ancestor of the Quaternary radiation of the clade represented in Europe with genera Nesiotites, Neomys and Macroneomys. The transition from Asoriculus to Neomys is expected to appear during the Q1 stage of the Early Pleistocene, more records identified as fossil species of the latter genus are reported from terminal stage of the Early Pleistocene (Q2). Nevertheless, their relations to extant species were not analysed and past history of the extant European taxa (fodiens, anomalus, milleri) and their range dynamics is despite of molecular data generally unclear. This fact reflects both scarcity of the fossil record and uncertainties regarding patterns of phenotype variation in particular species, their discrimination criteria and orientation of corresponding phylogenetic morphoclines. In these regards, the thesis pays special...

To determine the habitat characteristics of the Japanese water shrew, Chimarrogale platycephalus, seven stream parameters were studied along five streams. The shrew occurred along three of those streams. To evaluate the stream parameters related to water shrew occurrence as an objective variable, a generalized linear model analysis of the five streams was performed. The results indicated that current velocity was the most important parameter in shrew occurrence at pNeomys fodiens. The number of invertebrates as availability of food resources, however, was not a significant parameter influencing shrew occurrence. Along the banks of the two streams inhabited by C. platycephalus, there are artificially managed leisure fishing sites, and small fish farms have released fish regularly into the streams to stock up fishing resources. Such management of leisure fishing may provide the water shrews with fish as an additional food resource and possibly contribute to the occurrence of C. platycephalus.

Being venomous should help a predator to maximize foraging gain, reduce handling time, and hoard food. Shrews are a good model for study of the relationship between venomousness and optimal foraging as they have to be highly efficient foragers (because of extremely high energy requirements) and comprise a few venomous species. We hypothesized that venom facilitates shrews to overpower large prey and enables hoarding of (mainly large) prey in a comatose state. We compared hunting and hoarding behaviors of the venomous Neomys fodiens and the nonvenomous Sorex araneus, which were allowed to forage on live prey differing in body size. As we predicted, small invertebrates were immediately consumed by both shrew species, whereas larger ones were immobilized and hoarded. Smaller S. araneus hoarded proportionally more food than larger N. fodiens. Although different ways of prey immobilization were expected (venomous paralyzing by N. fodiens and mechanical damage by S. araneus), both species seemed to use mechanical immobilization. However, N. fodiens required significantly less time than S. araneus to subdue prey of proportionally similar sizes and this difference grew with the increase in prey mass. Only N. fodiens was able to overpower and kill frogs (but not larger toads). We concluded that the relatively weak venom of N. fodiens is helpful in overpowering and hoarding mainly medium-sized prey. Thus, venomousness helps N. fodiens to optimize its foraging as it reduces prey handling time, enables it to gain larger energy portions, and facilitates food hoarding, which reduces foraging time.

Background Animal toxins can have medical and therapeutic applications. Principally, toxins produced by insects, arachnids, snakes and frogs have been characterized. Venomous mammals are rare, and their venoms have not been comprehensively investigated. Among shrews, only the venom of Blarina brevicauda has been analysed so far, and blarina toxin has been proven to be its main toxic component. It is assumed that Neomys fodiens employs its venom to hunt larger prey. However, the toxic profile, properties and mode of action of its venom are largely unknown. Therefore, we analysed the cardio-, myo- and neurotropic properties of N. fodiens venom and saliva of non-venomous Sorex araneus (control tests) in vitro in physiological bioassays carried out on two model organisms: beetles and frogs. For the first time, we fractionated N. fodiens venom and S. araneus saliva by performing chromatographic separation. Next, the properties of selected compounds were analysed in cardiotropic bioassays in the Tenebrio molitor heart. Results The venom of N. fodiens caused a high decrease in the conduction velocity of the frog sciatic nerve, as well as a significant decrease in the force of frog calf muscle contraction. We also recorded a significant decrease in the frog heart contractile activity. Most of the selected compounds from N. fodiens venom displayed a positive chronotropic effect on the beetle heart. However, one fraction caused a strong decrease in the T. molitor heart contractile activity coupled with a reversible cardiac arrest. We did not observe any responses of the insect heart and frog organs to the saliva of S. araneus. Preliminary mass spectrometry analysis revealed that calmodulin-like protein, thymosin β-10, hyaluronidase, lysozyme C and phospholipase A2 are present in the venom of N. fodiens, whereas thymosin β4, lysozyme C and β-defensin are present in S. araneus saliva. Conclusion Our results showed that N. fodiens venom has stronger paralytic properties and lower cardioinhibitory activity. Therefore, it is highly probable that N. fodiens might use its venom as a prey immobilizing agent. We also confirmed that S. araneus is not a venomous mammal because its saliva did not exhibit any toxic effects. Electronic supplementary material The online version of this article (10.1186/s12983-017-0230-0) contains supplementary material, which is available to authorized users.

By using the method of geometric morphometry, the article analyses the main directions of shape variability of dentale and lower molars in Palearctic shrews: Sorex araneus, S. caecutiens, S. daphaenodon, S. isodon, S. roboratus, S. tundrensis, S. alpinus, S. asper, S. buchariensis, S. mirabilis, S. raddei, S. volnichini, S. unguiculatus, S. minutes, S. minutissimus, S. gracillimus, S. satunini, Neomys fodiens, Crocidura suaveolens, C. lasiura, C. leucodon, C. sibirica. It is established that intergenus and interspecies differences are poorly expressed in the shape of a dentale. There are three major ways of modifying the shapes of dentale in shrews: (i) massive high proc. coronoidens and low short proc. condiloideus (S. araneus from Altai mountains, S. daphaenodon, S. asper, S. raddei) (ii) low proc. coronoidens and long massive proc. condiloideus (S. roboratus, S. mirabilis, S. alpinus, S. unguiculatus of Sakhalin island, «minutes», N. fodiens (iii) high massive proc. coronoidens and push back proc. condiloideus (Crocidura except leucodon). It is shown that all the variants help to achieve equal morphological and functional effect. Intraspecies variation of the shape of the mandible is reduced to its transformation in according to one of the scenarios mentioned above.

We describe a new dicrocoeliid collected from the Eurasian water shrew Neomys fodiens in Bulgaria based on the combination of morphological characteristics and molecular data. Pojmanskatrema balcanica n. sp., n. gen. is distinguished from all other species of family Dicrocoeliidae Looss, 1900 in having a postovarian vitellarium situated in the posterior half of the body, postacetabular, nearly tandem, lobed testes, and relatively short body. Molecular phylogenetic analysis based on 28S rDNA sequences of the new species and previously sequenced dicrocoeliid taxa has demonstrated close relationship of Pojmanskatrema n. gen. with the genus Eurytrema. Considering the non-monophyletic nature of previously recognized dicrocoeliid sub-families we support the recently proposed dissolution of the sub-families within this family.

When a population shows a marked morphological change, it is important to know whether that population is genetically distinct; if it is not, the novel trait could correspond to an adaptation that might be of great ecological interest. Here, we studied a subspecies of water shrew, Neomys fodiens niethammeri, which is found in a narrow strip of the northern Iberian Peninsula. This subspecies presents an abrupt increase in skull size when compared to the rest of the Eurasian population, which has led to the suggestion that it is actually a different species. Skulls obtained from owl pellets collected over the last 50 years allowed us to perform a morphometric analysis in addition to an extensive multilocus analysis based on short intron fragments successfully amplified from these degraded samples. Interestingly, no genetic divergence was detected using either mitochondrial or nuclear data. Additionally, an allele frequency analysis revealed no significant genetic differentiation. The absence of genetic divergence and differentiation revealed here indicate that the large form of N. fodiens does not correspond to a different species and instead represents an extreme case of size increase, of possible adaptive value, which deserves further investigation.

Remains of 8 species (Soricidae: Sorex minutissimus , S. minutus , S. caecutiens , S. tundrensis , S. isodon , S. araneus , Neomys sp., Crocidura cf. leucodon ) have been found in Late Pleistocene and Holocene layers from 9 localities of the Perm Pre-Ural. Their description, systematic position, and temporal distribution are given. S. tundrensis dominated in the explored territory until the early Holocene. Currently, this species is very rare. Fossil white-toothed shrew ( Crocidura cf. leucodon) has been discovered for the first time in the study area.

Introduced fish represent a major threat for aquatic ecosystems and species and their impact on semiaquatic vertebrates, such as amphibians, has been extensively studied. On the other hand, little is known about their impact on semiaquatic mammals, such as water shrews (Neomys spp., Soricidae), although small mammals have been reported in their diet. The present note reports on a case of predation on an adult Eurasian water shrew Neomys fodiens by a large sized brown trout Salmo trutta belonging to a population introduced in a small pastureland pond in an Italian mountain area. The authors considered that shrews, including relatively large species such as Neomys spp., can be susceptible to predation by introduced fish and they use this finding to discuss about direct and indirect impacts of introduced fish on the populations of shrews inhabiting originally fishless habitats such as headwaters.

The chromosomal relationships between the Japanese water shrew Chimarrogale platycephalus and the Eurasian water shrew Neomys fodiens were investigated by conventional and differential staining. From the side-by-side analysis of matching G-banding patterns from the two species, a high degree of G-band homology was recognized in most of the chromosomes that were compared, although the extent of such homology was unclear in several small chromosomes. In C. platycephalus, large blocks of C-heterochromatin (C-blocks) were found in pair nos. M1, M8, M10 and ST23 and in the Y chromosome as conspicuous C-bands, while N. fodiens had no such C-blocks, having centromeric C-bands on most chromosomes with the exception of several small pairs, which included ST1 and ST23. Silver-stained nucleolus organizers (Ag-NORs) were found only on the secondary constriction of pair no. 1 in C. platycephalus and on the minute short arms of ST23 and ST24 in N. fodiens. A discussion of interspecific differences in C-bandin...

The present study was aimed at analyzing the total mercury in the tissues of mammals adapted to the semi-aquatic or subterranean lifestyle and at analyzing the possible role of their antioxidant system in heavy metal detoxification. The water shrew (Neomys fodiens Pennant, 1771), European mole (Talpa europaea Linnaeus 1758), muskrat (Ondatra zibethicus Linnaeus, 1766), and water vole (Arvicola terrestris Linnaeus, 1758) were the species under study. Our results indicate that the tissue mercury accumulation depends on age, the tissue type and diet of the species. The largest mercury deposition occurred in the water shrew tissues as compared with the other species. Age features of toxicant accumulation in the animals were noted. A correlation between the mercury concentration and the catalase activity in the kidneys of the species investigated was revealed.

Resumen Se han estudiado los cambios en la distribucion y frecuencia de los micromamiferos semiacuaticos, entre los anos 2000 y 2013-2014, de la red fluvial del parque natural Alt Pirineu, en los Pirineos meridionales. La especie prioritaria del estudio ha sido el desman iberico (Galemys pyrenaicus), pero tambien se han evaluado las frecuencias de los musganos (Neomys fodiens y Neomys milleri) y la rata de agua (Arvicola sapidus), para las que los datos del ano 2000 eran parciales. Es la primera vez que se realiza un estudio diacronico sobre la frecuencia relativa del desman a escala regional y aplicando un metodo identico en los dos muestreos. La metodologia utilizada ha sido la deteccion de indicios de presencia de estas especies en 66 tramos fluviales, prospectados en los dos periodos considerados. Actualmente, los micromamiferos mas comunes son los musganos, presentes en casi todos los tramos (94%), seguidos del desman (45%) y de la rata de agua (34%). Se ha constatado que el desman presenta una regresion intensa en una subcuenca fluvial (Noguera de Cardos) y ha desaparecido en su cabecera. Esta regresion parece atribuible a cambios recientes en la gestion hidroelectrica y se ha iniciado un seguimiento para evaluar esta hipotesis. Palabras clave: Arvicola, Galemys, mamiferos semiacuaticos, Neomys, seguimiento. Abstract We have studied the changes in the distribution and frequency of semiaquatic micromammals in the river network of Alt Pirineu Natural Park (southern Pyrenees, Iberian Peninsula) between 2000 and 2013-2014. The priority of this work has been the Iberian Desman (Galemys pyrenaicus) but we also evaluated the frequency in Water Shrews (Neomys fodiens and Neomys milleri) and Southern Water Vole (Arvicola sapidus), for which 2000 data were incomplete. It is the first time that a diachronic study on the relative frequency of Iberian Desman takes place at regional level and using an identical method in the two surveys. The methodology was the detection of faeces of these species in 66 river sections, prospected in the two periods considered. The most common semiaquatic mammals were Water Shrews, present in almost all river sections (94%), followed by the Desman (45%) and the Water Vole (34%). We found an intense regression of the Desman in a river basin where this species was common in 2000 (Noguera de Cardos) and it is extinct in the upper basin of this river. This local regression is probably attributable to a recent change in the model of hydroelectric management and we started a survey to evaluate this hypotesis. Keywords: Arvicola, Galemys, Neomys, semiaquatic mammals, survey. DOI: 10.7325/Galemys.2015.A4

The trophic mechanisms of differentiation and the relationships between various species of small insectivorous mammals, as well as their interaction with the populations of their prey, were studied. The stomach contents of 3225 animals of seven different species (Sorex araneus, S. caecutiens, S. minutus, S. isodon, S. minutissimus, Neomys fodiens, and Sicista betulina) caught in Karelia were analyzed. These species inhabit the same territories, and their main staple food objects overlap almost completely. Thus, the specific nature of their diets is quantitative rather than qualitative. At the same time, the trophic relation of the animals with soil invertebrates becomes weaker from the even-toothed and common shrews to the northern birch mouse. The ecological differentiation and separation of the forage between close species of insectivores were shown to be primarily provided by the different horizons of their habitat (soil and surface layers, grass and shrub tiers, etc.).

Neomys fodiens Pennant, 1771 —Eurasian Water Shrew Sorex fodiens Pennant, 1771 p.308; Type locality-Berlin, Germany. Neomys fodiens orientis Thomas, 1914 p.564; Type locality- Swamps of Kammanajaretschka River, Semirechyia, Russian central Asia; Won, 1968 p.62; Yoon, 1992 p.22 (Won (1968) and Yoon (1992) used the name orientalis, perhaps a misprint of orientis). N. watasei Kishida, 1930 p.372 (Nomen nudum). N. fodiens watasei: Kuroda, 1941 p.114; Type locality- Toyohara, Sakhalin Island, Russia. N. limchunhunii Won, 1954 p.41; Type locality- Pungseo, Ryanggang Province, Korea. N. fodiens: Won, 1968 p.62; Han, 1994 p.45; Won & Smith, 1999 p.9; Han, 2004a p.28. Range: Eurasian water shrews inhabit the northeastern peninsula from Mt. Baekdu southward to Mt. Seorak National Park along high mountain ranges (Fig. 9). Since the first collection of this species in 1953 near Ryanggang Province, mammalogists have viewed N. fodiens as a species from North Korea (Won 1968). However, since 2007 this shrew has been observed in South Korea (NIBR 2012). Remarks: This species was first described as a new species, Neomys limchunhunii Won, 1955. Korean populations of the Eurasian water shrew are now considered N. fodiens orientis. Literature citations previously referred to this shrew as N. f. orientalis (Won 1968; Yoon 1992; Won & Smith 1999), but Neomys fodiens orientis Thomas, 1914 had priority over Neomys fodiens orientalis Hinton, 1915 as the proper scientific name (Corbet 1978). Conservation status: The North Korean government classified N. fodiens as a ‘Rare’ species (MAB National Committee of DPR Korea 2002)., Published as part of Jo, Yeong-Seok, Baccus, John T. & Koprowski, John L., 2018, Mammals of Korea: a review of their taxonomy, distribution and conservation status, pp. 1-216 in Zootaxa 4522 (1) on pages 20-22, DOI: 10.11646/zootaxa.4522.1.1, http://zenodo.org/record/2610198, {"references":["Pennant, T. (1771) Novae durch verschiedene provinzen des Russischen reichs. Descriptiones animalium, Fol. 1. Gedruckt bey der Kayserlinchen Academie der Wissenschaften, St. Petersburg, 382 pp. [in German]","Thomas, O. (1914) On small mammals from Djarkent, Central Asia. Annals and Magazine of Natural History, Series 8, 13, 563 - 573. https: // doi. org / 10.1080 / 00222931408693525","Won, H. K. (1968) The mammals of Korea. Institute of Science Press, Pyeongyang, 408 pp. [in Korean]","Yoon, M. H. (1992) The wildlife of Korea. Daewonsa Publishing Company, Seoul, 142 pp. [in Korean]","Kishida, K. (1930) Mammal fauna in northern Japan. Zoological Magazine, 42, 372 - 373.","Kuroda, N. (1941) Descriptions of two apparently new forms of mammals from Saghalien and Hokkaido, Japan. Bulletin of the Biogeographical Society of Japan, 11, 113 - 116.","Won, H. K. (1954) Neomys limchunhunii. Bulletin of Institute of Science, 6, 41 - 43.","Han, S. H. (1994) The checklist of Mammals in North Korea. Nature Conservation, 86, 44 - 50. [in Korean]","Won, C. M. & Smith, K. G. (1999) History and current status of mammals of the Korean Peninsula. Mammal Review, 29, 3 - 33. https: // doi. org / 10.1046 / j. 1365 - 2907.1999.00034. x","Han, S. H. (2004 a) Order Insectivora. In: Won, B. O. (Ed.), The mammals of Korea. Dongbang Media Press, Seoul, pp. 17 - 35. [in Korean]","Corbet, G. B. (1978) The mammals of the Palaearctic region: a taxonomic review. British Museum (Natural History), London, 314 pp.","MAB National Committee of DPR Korea (2002) Red Data Book of DPRK (Animal). Democratic People's Republic of Korea, Pyongyang, 316 pp."]}

179. Mediterranean Water Shrew Neomys anomalus French: Crossope de Miller / German: Sumpfspitzmaus / Spanish: Musgano de Cabrera Other common names: Iberian Water Shrew, Miller's Water Shrew, Southern Water Shrew Taxonomy. Neomys anomalus Cabrera, 1907, “ San Martin de la Vega (province of Madrid, on the Jarama River,” Spain. Six subspecies are recognized. Subspecies and Distribution. N.a.anomalusCabrera,1907—IberianPeninsula. N.a.jostiV.Martino&E.Martino,1940—BalkanPeninsula. N.a.milleriMottaz,1907—Alps. N.a.mokrzeckiiE.Martino&V.Martino,1917—Crimea. N.a.rhenanusLehmann,1976—SGermany. N. a. soricoides Ognev, 1921 — E Poland, S Belarus, and W Russia. Also known from W, SC & E Europe, Turkey, and Iran, but subspecies involved not known. Descriptive notes. Head-body 68-85 mm, tail 46-56 mm, hindfoot 14-8-18-3 mm; weight 9-5-13-5 g. No specific data are available for body weight, but pregnant females can weigh up to 20 g. The Mediterranean Water Shrew is large for the family Soricidae, but it is the smallest species of Neomys. Tail usually exceeds 65% of head-body length. Swimming borders on feet and keel on tail are formed of long stiff hairs but are not as developed as in the other species of Neomys. Pelage is dense and short. Back and sides are dark brown or black; belly is light gray, sometimes with pale yellow or brownish tint. Small light spot behind eye is distinct. Tail is bicolored;its upper part and underside are the same color as back and belly. Dental formula for species of Neomysis 13/2, C 1/0,P2/1,M 3/3 (x2) = 30. Chromosomal complement has 2n = 52 and FN = 98. Xchromosome is large subtelocentric, Y-chromosome is small subtelocentric. There are 44 metacentric and submetacentric autosomes and six acrocentric autosomes. Habitat. Often floodplain habitats and banks of small rivers and creeks flowing through deciduousforests and riverside marshes up to elevations of ¢.2100 m in mountainous areas. The Mediterranean Water Shrew is the most terrestrial species of Neomys and thus capable of living far from water bodies. It can be found in irrigated gardens. In northern regions of the distribution,it is often displaced from favorable near-water habitats by the Eurasian Water Shrew (NN. fodiens), which is better adapted to water activities. In southern regions, many water bodies dry up by mid-summer, and ability to live far from water provides competitive advantage to the Mediterranean Water Shrew. Food and Feeding. The Mediterranean Water Shrew eats terrestrial and water invertebrates, but the latter accounts for a smaller part of the diet. Earthworms, arachnids, adult dipterans, and adult beetles (most commonly carabids) are preferred among terrestrial invertebrates; isopods (Asselus), caddis worms, soldier fly larvae, and horsefly larvae are preferred water invertebrates. Water and terrestrial mollusks were often found in gastric contents. The Mediterranean Water Shrew does not dive to hunt water invertebrates in localities where it lives in sympatry with Eurasian Water Shrews; however, diving is used as a foraging method in Portugal, where other species of Neomys are absent. Prey is immobilized when bitten because saliva enzymes exert a paralytic effect. Breeding. Independent juvenile Mediterranean Water Shrews become detectable in late May, and the latest capture of a lactating female was on 30 October. A female produces up to three litters in a breeding season in Austria. Youth-of-the-year are extensively involved in reproduction. Productivity is generally higher than in most other species of shrews. Pregnant females have 5-13 embryos; highest numbers of embryos were twelve in Poland and Austria and 13 in Italy. Activity patterns. The Mediterranean Water Shrew is semi-aquatic and active almost around the clock; activity at night is somewhat higher than activity during the day. Movements, Home range and Social organization. Mediterranean Water Shrews can live in groups with a high degree of tolerance toward each other. Although aggressive behavior is observed occasionally, it does not prevent several individuals from feeding together. Immigrants can easilyjoin in an established group. Social system is essentially open and non-hierarchical. Status and Conservation. Classified as Least Concern on The IUCN Red List. Distribution of the Mediterranean Water Shrew has many isolated areas, some of which are merely individual capture sites. Many such sites have been identified in the past 20-30 years. So many isolated areas of occurrence over a large distribution suggests that the Mediterranean Water Shrew is a relict species. It is rare or at low densities almost throughout its entire distribution, and it is on the Red List in Ukraine and regional Red Lists of Voronezh and Tambov regions (Russia). Bibliography. Baltiauskas & Balgiauskiené (2012), Bal¢iauskas et al. (2016), Borodin (2013), Churchfield & Rychlik (2006), Esmaeili et al. (2008), Kashtalian (2005), Krushinska & Rychlik (1994), Krystufek & Vohralik (2001), Lay (1967), Rychlik (1998), Sitnikova & Mishta (2008), Spitzenberger (1990b), Tapisso et al. (2013), Zima et al. (2008)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Soricidae, pp. 332-551 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on pages 456-457, DOI: 10.5281/zenodo.6870843

181. Eurasian Water Shrew Neomys fodiens French: Crossope aquatique / German: \Wasserspitzmaus / Spanish: Musgano patiblanco Other common names: Northern Water Shrew, Water Shrew Taxonomy. Sorex fodiens Pennant, 1771, Berlin, Germany. Subspecies niethammeri in north-western Spain is morphologically distinct from other populations and may warrant species status. Further taxonomic research is needed. Four subspecies recognized. Subspecies and Distribution. N.f.fodiensPennant,1771—fromAtlanticcoastofcontinentalEuropeEtoObRiverandAltaisteppes(WSiberia). N.f.bicolorG.Shaw,1791—GreatBritain. N.f.niethammeriBuhler,1963—NSpain. N. f. orientis Thomas, 1914 — from Ob River and Altai steppes (W Siberia) E to Pacific coast. Descriptive notes. Head—body 75-103 mm,tail 58-73 mm, hindfoot 16-21 mm; weight 8-5-25 g. The Eurasian Water Shrew is large. Keel occurs along entire length oftail, and swimming borders on feet are well formed with long stiff hairs. Tail is usually longer than 65% of head-body length. Pelage is relatively short. Back and sides are black-brown or black, sometimes with weak grayish tint. Belly is silvery gray, sometimes with brownish tint. White spot behind eye is commonly distinct. Dark spot is occasionally found on chest and chin, varying in shape and size. Tail is bicolored, with dark color of back above contrasting with silvery gray of belly below. Chromosomal complement has 2n = 52 and FN = 98. X-chromosomeis large subtelocentric, Y-chromosome is small subtelocentric. There are 44 metacentric and submetacentric autosomes and SIX acrocentric autosomes. Habitat. Forest zone and foreststeppe and forest-tundra neighboring subzones. The Eurasian Water Shrew avoids extended wooded areas, especially coniferous forests with well-developed moss layer. It inhabits banks of streams and stagnant waters; riverside meadows with quiet pools or other regions of low water velocity are preferred. It is occasionally found far from the water, usually at sites with tall grasses, especially in desolated vegetable gardens or settlements. Food and Feeding. The Eurasian Water Shrew eats terrestrial and water invertebrates, the latter prevailing in the diet. Isopods (Asellus), caddis worms, and larvae of various dipterans are most common dietary components among water invertebrates. Earthworms, mollusks, and various beetles are preferred among terrestrial invertebrates. Vertebrates (fish fry, tadpoles, and young frogs) are regularly found in diets, but their overall contribution is insignificant. The Eurasian Water Shrew dives to catch water invertebrates, occasionally reaching depths of 50-70 cm. Saliva enzymes exert a paralytic effect and serve to immobilize prey. Breeding. Breeding season of the Eurasian Water Shrew lasts throughout the warm period of the year (e.g. April-September in England), but most young are born in summer. Gestation averages 20 days; litters usually have 5-8 young, although captures of pregnant females with 15 embryos are known. Nests are built in hollows between plant roots, driftwood along banks, and rodent burrows. Nests are constructed with plant debris. There is evidence that longevity can occasionally reach 19 months in the wild (i.e. capable of overwintering twice). This feature distinguishes the Eurasian Water Shrew from other shrews, such as those of the genus Sorex. Nevertheless, such cases seem to be extremely rare because long-term observations of many marked Eurasian Water Shrews in southern England showed that they all died after the end of a breeding season and first overwintering. Eurasian Water Shrews can live up to 3-4 years in laboratory conditions. Activity patterns. The Eurasian Water Shrew is semi-aquatic. Activity occurs almost around the clock, most often peaking during dark hours. Maximal activity is observed before sunrise and after sunset. Time spent in nests in winter is generally longer than In summer. Movements, Home range and Social organization. The Eurasian Water Shrew is solitary and promiscuous. Shifting home ranges are characteristic ofits spatial organization; i.e. home range of an individual shifts during its life following a shift in vital resources, such as when the water’s edge moves as a water body dries out. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Eurasian Water Shrew is generally low in abundance, although it is common in favorable habitats and is occasionally codominant in the community of small mammals in Barabinsk Lowland in western Siberia. The Eurasian Water Shrew is on the Red Lists of Latvia, Kyrgyzstan, and 17 regions of the Russian Federation. Bibliography. Aloise et al. (2005), Cantoni (1993), Churchfield (1984a, 1984b, 1990), Dehnel (1950), Kalyakin (1985), Kowalski & Rychlik (2018), Lopez-Fuster et al. (1990), Maksimov (1959), Nesterenko (1999), Okhotina (1984), Reimers & Voronov (1963), Revin (1989), Rychlik (1998), Sheftel (1983), Spitzenberger (1990c), Volpert & Danilov (1999), Won Chang-Man & Smith (1999), Yanushevich et al. (1972), Zima et al. (1998)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Soricidae, pp. 332-551 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 457, DOI: 10.5281/zenodo.6870843

180. Transcaucasian Water Shrew Neomys teres French: Crossope du Caucase / German: Kaukasische Bachspitzmaus / Spanish: Musgano del Caucaso Taxonomy. Neomys teres G. S. Miller, 1908, “ edge of brook in mountains (altitude 7000 feet [= 2134 m]) 25 miles [= 40 km] north of Erzeroum [= Erzurum, Erzurum Province], Turkey.” Neomys teres is ecologically close and very similar in appearance to N. fodiens, although genetically equidistant from N. anomalus and N. fodiens. Subspecific taxonomy needs verification. Four subspecies recognized. Subspecies and Distribution. N. t. teres G. S. Miller, 1908 — Turkey. N. t. balkaricus Ognev, 1926 — Ciscaucasia. N. t. leptodactylus Satunin, 1914 — Transcaucasia. N. t. schelkounikovi Satunin, 1913 — Greater Caucasus. Descriptive notes. Head-body 83-101 mm, tail 58-73 mm, hindfoot 18-2-22-1 mm; weight 11-28 g. The Transcaucasian Water Shrew is very large, larger than any other modern species of Neomys. Keel on tail is well developed and occupies about 50% oftail’s length distally. Swimming borders ofstiff hairs on feet are quite well developed. Tail is usually longer than 65% of head-body length. Pelage is longer and denser compared with other species of Neomys. Eyes are hidden in fur. Upper parts of body are blackbrown, with weak ash or silvery gray tint. Belly is light gray, sometimes with pale yellow or grayish brown tint. Light spot behind eyeis poorly seen. Tail varies from bicolored to light unicolored. Chromosomal complement has 2n = 52 and FN = 98. X-chromosome is medium-sized submetacentric, and Y-chromosome is small metacentric. There are 44 metacentric and submetacentric autosomes and 3 acrocentric autosomes. Habitat. Banks of small rivers and creeks in mixed and coniferous forest belts of mountains or alpine meadows at elevations up to 2500 m. Habitats with rank near-water vegetation are preferred, while habitats near large rivers are avoided, especially when banks are barren of vegetation. In Turkey, the Transcaucasian Water Shrew is occasionally found along irrigation channels and ditches into fields and orchards. Food and Feeding. Gastric contents of Transcaucasian Water Shrews were examined in Armenia, and terrestrial invertebrates were higher in biomass than water invertebrates, accounting for 21% oftotal biomass. More common terrestrial invertebrates included earthworms, adult beetles, adult dipterans, spiders, and harvestmen; caterpillars prevailed in gastric contents of two individuals. Isopods (Asellus), freshwater shrimp, and caddis worms were most prevalent among water invertebrates. Vertebrate tissues were not found in stomachs. The Transcaucasian Water Shrew might be capable of immobilizing its prey, like other species of Neomys, but studies to verify this are lacking. Breeding. Young Transcaucasian Water Shrews are born in May-August. Litters have 2-9 young (average six). Activity patterns. The Transcaucasian Water Shrew is semi-aquatic. Activity periods occur mostly at night or twilight in summer and shift to daytime in winter when individuals are active in the morning and afternoon. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Least Concern on The IUCN Red List. Population sizes are not high, but the Transcaucasian Water Shrew is common. Bibliography. Bannikova & Kramerov (2005), Churchfield et al. (2006), Krystufek & Vohralik (2001), Sokolov & Tembotov (1989), Zaitsev et al. (2014), Zima et al. (1998)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Soricidae, pp. 332-551 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 457, DOI: 10.5281/zenodo.6870843

Objective: The objective of this study was to determine the species composition of mammals and parasites involved in the epizootic process of plague in Armenia and their geographic distribution. Introduction: Plague was first identified in Armenia in 1958 when Y. pestis was isolated and cultured from the flea species Ct. teres collected from the burrows of common voles in the northwestern part of the country. In the process of digitalizing archived data, a statistical and spatial analysis of the species composition of mammals and parasites involved in the epizootic process of plague between 1958 and 2016 was performed. Methods: The plague archives of the NCDCP were exploited. The geographic addresses from which strains of Y. pestis were isolated from mammals and their parasite species were analyzed and grouped into 38 administrative regions (Fig.1). For geostatistical analysis, databases were created using Microsoft Excel and converted into a ESRI Geodatabase (Fig.2). Results: Data from the especially dangerous pathogen laboratories indicate that 9329 Y. pestis strains were isolated in 27 of the 38 regions of the country with 7022 (75%) of the strains found in just four regions: Abovyan 2597 (28%), Sisian 1953 (21%), Martuni 1416 (15%) and Ashotsk 1056 (11%) (Fig.3). During this period, plague bacteria were isolated from 17 mammal species including 15 rodents, Mustela nivalis (weasel), and Neomys fodien s (shrew) (Fig.4). Y. pestis was isolated from 22 species of fleas belonging to 11 genera along with two families of ticks. Of the 9329 bacterial isolates, 6540 (70.2%) came from fleas, 2646 (28.3%) came from mammals and 143 (1.5%) were from ticks (Fig.5). Conclusions: In Armenia, the primary mammalian host for Y. pestis is the common vole Microtus arvalis from which 2600 isolates (27.9%) were taken. Flea species from which large numbers of plague bacteria have been isolated include Ct. teres- 3758 (40.3%), Ct. wladimiri- 1262 (13.5%) and C. caspia- 667 (7.1%).

The present paper is the concluding part of the series describing the Soricidae fauna of the late Early Pleistocene Somssich Hill 2 locality, Southern Hungary. Th e assemblage includes 9 shrew forms: 3 Sorex, 2 Crocidura, 2 Beremendia, 1 Asoriculus, and 1 Neomys species. Th e taxonomic results of the study are summarized, and some new ecological conclusions drawn from the layer-by-layer overview of the specific composition of the fauna are presented here. The forms of cold grassy vegetation dominate in most of the strata, but the indicators of warm grasslands also appear in some layers. The appearance of forest species in most layers correlates with the presence of aquatic shrews. The ratio of cold and warm-indicating species varies in the layers, but dominance of cold indicators decreases in the second half of the succession, while the number of forest and warm indicators increases.

The acoustic signals of shrews (Soricidae) are largely understudied. As shrews are very vocal animals it may be feasible to use acoustic methods in field studies to assess ecological and behavioral data. In this study, we present the first detailed analysis of the twittering calls of six Central European shrew species (Sorex minutus, Sorex araneus, Neomys fodiens, Neomys anomalus, Crocidura russula and Crocidura leucodon). The analysis is based on over 6000 recorded calls from 121 individuals. Our results indicate that there is a large inter-individual variance and a large inter-specific overlap in the acoustic parameters of the calls. Each species uses a large spectral variety of calls without clear species specific call types. A species identification using the Support Vector Machine method on six species shows 66.2% accuracy; however, a pairwise comparison indicates accuracy between 68.5 and 97.3%. We propose to use acoustic monitoring of shrews in comparative studies to estimate the overall shrew activity. Moreover we suggest using the acoustic identification method in areas with few shrew species where the accuracy of the technique can be eligible.

The body size of Palearctic Sorex shrews decreases at higher latitudes, and as such the Bergmann’s rule does not work. However, no analysis has ever been done for water shrew (Neomys fodiens) in the middle of distribution range. Analysis of available literature data showed that some body and skull measurements of N. fodiens are negatively correlated to latitude. Measurements of 158 water shrews from Estonia and Lithuania were also analyzed with respect to the short scale latitudinal pattern. We found that populations are separated (Wilk’s lambda = 0.363, p

Previously, we reported the discovery of a genetically distinct hantavirus, designated Boginia virus (BOGV), in the Eurasian water shrew (Neomys fodiens), as well as the detection of Seewis virus (SWSV) in the Eurasian common shrew (Sorex araneus), in central Poland. In this expanded study of 133 shrews and 69 moles captured during 2010–2013 in central and southeastern Poland, we demonstrate the co-circulation of BOGV in the Eurasian water shrew and SWSV in the Eurasian common shrew, Eurasian pygmy shrew (Sorex minutus) and Mediterranean water shrew (Neomys anomalus). In addition, we found high prevalence of Nova virus (NVAV) infection in the European mole (Talpa europaea), with evidence of NVAV RNA in heart, lung, liver, kidney, spleen and intestine. The nucleotide and amino acid sequence variation of the L segment among the SWSV strains was 0–18.8% and 0–5.4%, respectively. And for the 38 NVAV strains from European moles captured in Huta Dlutowska, the L-segment genetic similarity ranged from 94.1%–100% at the nucleotide level and 96.3%–100% at the amino acid level. Phylogenetic analyses showed geographic-specific lineages of SWSV and NVAV in Poland, not unlike that of rodent-borne hantaviruses, suggesting long-standing host-specific adaptation. The co-circulation and distribution of BOGV, SWSV and NVAV in Poland parallels findings of multiple hantavirus species co-existing in their respective rodent reservoir species elsewhere in Europe. Also, the detection of SWSV in three syntopic shrew species resembles spill over events observed among some rodent-borne hantaviruses.