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4.3 Crocidura suaveolens gueldenstaedtii The phylogenetic analysis indicated that all Israeli “ C. suaveolens ” sequences are part of clade V – the ‘ gueldenstaedtii ’ clade (Dubey et al. 2007a), and close to the Turkish and Georgian sequences (Figure 5). While the Israeli populations represent the edge of this species complex range, they do not present a mitochondrial genetic difference from the rest of its clade, unlike the situation in C. leucodon. This may be due to their larger population size and wider range in Israel compared to that of C. leucodon (Figure 1B). However, it is also possible that genetic differences exist in the nuclear genome. Based on the high similarity between the Israeli and Balkan sequences we agree with the status of “least concern” listed by the IUCN (Palomo et al. 2016). Our molecular cyt b data unequivocally place Israeli ‘suaveolens’ specimens within the ‘ gueldenstaedtii’ clade. The specific status of the form awaits a thorough taxonomic revision, including an examination of animals from the type localities, preferably alongside specimens from the type localities of other members of the complex (e.g., C. ‘gmelini’, C. suaveolens monacha, C. portali) as well as phylogenetic information from nuclear markers. Until such a study is carried out, we tentatively refer to Israeli specimens as members of C. suaveolens gueldenstaedtii., Published as part of Shpirer, Erez, Haddas-Sasson, Michal, Spivak-Glater, Maya, Feldstein, Tamar, Meiri, Shai & Huchon, Dorothée, 2021, Molecular relationships of the Israeli shrews (Eulipotyphla: Soricidae) based on cytochrome b sequences, pp. 79-89 in Mammalia (Warsaw, Poland) (Warsaw, Poland) 85 (1) on page 86, DOI: 10.1515/mammalia-2019-0143, http://zenodo.org/record/7837749, {"references":["Dubey, S., Cosson, J. F., Magnanou, E., Vohralik, V., Benda, P., Frynta, D., Hutterer, R., Vogel, V., and Vogel, P. (2007 a). Mediterranean populations of the lesser white-toothed shrew (Crocidura suaveolens group): an unexpected puzzle of Pleistocene survivors and prehistoric introductions. Mol. Ecol. 16: 3438 - 3452.","Palomo, L., KryStufek, B., Amori, G., and Hutterer, R. (2016). Crocidura suaveolens. The IUCN Red List of Threatened Species 2016."]}

4.4 Suncus etruscus The Israeli S. etruscus sequences appear to be closely related to the European ones and one Iranian sequence. The differences between the Israeli sequences and those of other western clade members are very small and comparable to the distances observed within the C. suaveolens gueldenstaedtii clade. This suggests that the western clade encompasses individuals from France in the west to Iran in the east and Israel in the south. However, this species has been poorly sampled in molecular studies to date. As a case in point, only a few S. etruscus specimens have been sequenced from Iran where members of both the western and eastern clade are present (Darvish et al. 2017; Ohdachi et al. 2016). More data are needed in order to decipher the population structure of this species., Published as part of Shpirer, Erez, Haddas-Sasson, Michal, Spivak-Glater, Maya, Feldstein, Tamar, Meiri, Shai & Huchon, Dorothée, 2021, Molecular relationships of the Israeli shrews (Eulipotyphla: Soricidae) based on cytochrome b sequences, pp. 79-89 in Mammalia (Warsaw, Poland) (Warsaw, Poland) 85 (1) on page 87, DOI: 10.1515/mammalia-2019-0143, http://zenodo.org/record/7837749, {"references":["Darvish, J., Mahmoudi, A., Pehpuri, A., and Saeidzadeh, S. (2017). New data on distribution and taxonomy of the genus Suncus (Mammalia: Soricidae) in Iran; molecular evidence. Iran. J. Anim. Biosyst. 13: 229 - 235.","Ohdachi, S. D., Kinoshita, G., Oda, S. - i., Motokawa, M., Jogahara, T., Arai, S., Nguyen, S. T., Suzuki, H., Katakura, K., Bawm, S., et al. (2016). Intraspeci fi c phylogeny of the house shrews, Suncus murinus-S. montanus species complex, based on the mitochondrial cytochrome b gene. Mamm. Stud. 41: 229 - 238."]}

4.1 Crocidura ramona Our work, based on sequences from type specimens, corroborates the view that C. ramona is a distinct species and probably endemic to Israel and the West Bank (Dubey et al. 2008; Ivanitskaya et al. 1996). C. ramona has been suggested to be related to the Palearctic “flat-headed rockshrews” (i.e., Crocidura pergrisea, C.arispa, Crocidura armenica, Crocidura serezkyensis and C. zarudnyi) (Burgin et al. 2018b; Kry Š tufek and Vohralík 2001). Although the DNA of most of the latter has not yet been sequenced, our analysis indicates that C. ramona and C. zarudnyi are distantly related (Figure 2). Similarly, we have shown here that C. ramona is not closely related to another silvery-gray shrew – C. nana, as the two formed two distinct and distant clades in our phylogenetic analyses (Supplementary Figure S1). We note that there is some doubt regarding C. nana ’s distribution. It is usually considered to be restricted to Somalia and Ethiopia (Cassola 2019; Hutterer 2005), whereas the samples sequenced in this work originated from Tanzania. We cannot be certain therefore that C. nana is the correct species assignment for the Tanzanian samples that we sequenced. Because no morphological comparisons have been carried out between the skulls of C. katinka and C. ramona, the exact relationship between these two species remains to be determined. Although C. katinka has been suggested to be related to certain African species with cranial similarities (e.g., Crocidura bottegi, C. obscurior, Crocidura bottegoides) (Burgin et al. 2018b; Hutterer and Kock 2002), our analyses have demonstrated that C. ramona is unrelated to C. obscurior. It is also possible that C. ramona is conspecific with C. portali. Thomas (1920) described C. portali as a small shrew (“though not excessively so”), with “pale drab-grey” pelage; and indicated that it has “clearly nothing to do with the C. russula group”. The gross morphology of the holotype (BMNH #19.4.11.9) and its size agree with C. ramona – though a detailed examination is still needed in order to confirm or refute this. Kry Š tufek and Vohralík (2001) suggested that C. portali may be a valid species, related to Crocidura arispa (and other members of the pergrisea group), and a senior synonym of C. ramona. Hutterer and Kock (2002) indicated that C. ramona and C. portali are similar in skull dimensions as well as in pelage, but note that they are also similar to C. gmelini, a species that they note requires “a better definition”. C. gmelini (ranging from Iran to Mongolia) has been synonymized with C. suaveolens, a species distantly related to C. ramona (Figure 2). C. arispa, C. ramona, and C. katinka are currently all considered valid species, while C. portali is not (Burgin et al. 2018b; IUCN 2020). Clearly, a taxonomic revision of these taxa is warranted. Neither the DNA of the C. portali type, nor that of any shrews identified as C. arispa, C. pergrisea, or C. katinka, has been sequenced and, unfortunately, we failed to amplify any cyt b fragment from a tissue of a specimen identified as C. portali (BMNH ZD 1971.817). We thus tentatively ascribe the sequence we obtained to C. ramona, pending a taxonomic revision., Published as part of Shpirer, Erez, Haddas-Sasson, Michal, Spivak-Glater, Maya, Feldstein, Tamar, Meiri, Shai & Huchon, Dorothée, 2021, Molecular relationships of the Israeli shrews (Eulipotyphla: Soricidae) based on cytochrome b sequences, pp. 79-89 in Mammalia (Warsaw, Poland) (Warsaw, Poland) 85 (1) on page 86, DOI: 10.1515/mammalia-2019-0143, http://zenodo.org/record/7837749, {"references":["Dubey, S., Salamin, N., Ruedi, M., Barriere, P., Colyn, M., and Vogel, P. (2008). Biogeographic origin and radiation of the Old World crocidurine shrews (Mammalia: Soricidae) inferred from mitochondrial and nuclear genes. Mol. Phylogenet. Evol. 48: 953 - 963.","Ivanitskaya, E., Shenbrot, G., and Nevo, E. (1996). Crocidura ramona sp. nov. (Insectivora, Soricidae): a new species of shrew from the central Negev desert, Israel. Z. Saugetierkd. 61: 93 - 103.","Burgin, C. J., Colella, J. P., Kahn, P. L., and Upham, N. S. (2018 a). How many species of mammals are there? J. Mammal. 99: 1 - 14.","Cassola, F. (2019). Crocidura nana. The IUCN Red List of Threatened Species 2019. e. T 41341 A 22306927.","Hutterer, R. (2005). Order Soricomorpha. In Wilson DE., Reeder DM.","Hutterer, R., and Kock, D. (2002). Recent and ancient records of shrews from Syria, with notes on Crocidura katinka Bate 1937 (Mammalia: Soricidae). Bonn. Zool. Beitr. 50: 249 - 258.","Thomas, O. (1920). A new shrew and two new foxes from Asia Minor and Palestine. Ann. Mag. Nat. Hist. 5: 199 - 122.","IUCN. 2020. The IUCN red list of threatened species. Gland: Switzerland. Version 2020 - 1. Available at: https: // www. iucnredlist. org. Downloaded on April 2020."]}

4.2 Crocidura leucodon The phylogenetic reconstruction indicates that the Israeli C. leucodon forms a distinct clade related to the eastern clade (Figure 4, Dubey et al. 2007b). The different C. leucodon clades have been suggested to have diverged during the Pleistocene glaciations (Dubey et al. 2007b; Mahmoudi et al. 2019). It is unlikely however that the Israeli clade corresponds to a fourth refugium from the Ice Age. Rather, the observed mitochondrial genetic divergence is possibly the result of the edge position of the Israeli population at the southernmost part of the C. leucodon range. In agreement, Mendelssohn and Yom-Tov (1999) noted that C. leucodon is less abundant than C. s. gueldenstaedtii, basing their conclusion on the number of shrews deposited in museum collections. The current data available at the Steinhardt Museum of Natural History support the view that the range of C. leucodon is more limited than that of C. suaveolens in Israel, since C. leucodon samples are rarer in the collection (79 leucodon vs. 594 suaveolens), and with one exception, restricted to the Mediterranean biome (Figure 1B). Edge populations have been suggested to harbor adaptive traits and genetic variability that may be important when considering future conservation needs (Hampe and Petit 2005; Mátýas et al. 2009). We thus also recommend that further population and genomic studies be carried out on this species, since its population status is currently unknown (Shenbrot et al. 2016), and assessment from museum collections alone may provide a biased representation of the population status., Published as part of Shpirer, Erez, Haddas-Sasson, Michal, Spivak-Glater, Maya, Feldstein, Tamar, Meiri, Shai & Huchon, Dorothée, 2021, Molecular relationships of the Israeli shrews (Eulipotyphla: Soricidae) based on cytochrome b sequences, pp. 79-89 in Mammalia (Warsaw, Poland) (Warsaw, Poland) 85 (1) on page 86, DOI: 10.1515/mammalia-2019-0143, http://zenodo.org/record/7837749, {"references":["Dubey, S., Cosson, J. F., Vohralik, V., KryStufek, B., Diker, E., and Vogel, P. (2007 b). Molecular evidence of Pleistocene bidirectional faunal exchange between Europe and the Near East: the case of the bicoloured shrew (Crocidura leucodon, Soricidae). J. Evol.","Mahmoudi, A., Darvish, J., Siahsarvie, R., Dubey, S., and KryStufek, B. (2019). Mitochondrial sequences retrieve an ancient lineage of bicolored shrew in the Hyrcanian refugium. Mamm. Biol. 95: 160 - 163.","Mendelssohn, H., and Yom-Tov, Y. (1999). Fauna Palaestina: Mammalia of Israel. Jerusalem: The Israel Academy of Sciences and Humanities, pp. 439.","Hampe, A., and Petit, R. J. (2005). Conserving biodiversity under climate change: the rear edge matters. Ecol. Lett. 8: 461 - 467.","Matyas, C., Vendramin, G. G., and Fady, B. (2009). Forests at the limit: evolutionary - genetic consequences of environmental changes at the receding (xeric) edge of distribution. Report from a research workshop. Ann. For. Sci. 66, https: // doi. org / 10.1051 / forest / 2009081.","Shenbrot, G., Hutterer, R., Amori, G., KryStufek, B., Yigit, N., Mitsain, G., and Palomo, L. J. (2016). Crocidura leucodon. The IUCN Red List of Threatened Species 2016. e. T 29651 A 115169304."]}