Talpa, Linnaeus, 1758
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publication ID |
https://doi.org/ 10.1111/zoj.12298 |
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persistent identifier |
https://treatment.plazi.org/id/5223FC41-2D73-FFB8-FEEF-52E4DCC42CB5 |
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treatment provided by |
Felipe |
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scientific name |
Talpa |
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GENUS Talpa View in CoL
T. talyschenses Vereschagin, 1945
Our molecular data provide unequivocal genetic evidence for a separate status of the moles from the Talysh Mts., which are deeply divergent from both the Caucasian mole and the Levant mole, being the sister branch to T. davidiana . According to our time estimates based on the nuclear data set, the age of the Talysh lineage is not less than 2 My. The moles from the Talysh Mts were previously treated as a subspecies of T. levantis ( Sokolov & Tembotov, 1989; Hutterer, 2005), although Zaitsev (1999) after examination of type material expressed doubt whether talyschensis is conspecific with T. levantis .
We conclude that moles occupying an area along the southwestern banks of the Caspian Sea from southernmost Azerbaijan to the central Elburz Mts in Iran, actually belongs to T. talyschensis and not to T. caucasica or T. levantis , as previously proposed (Kryštufek, 2001; Kryštufek & Benda, 2002). This finding adds one more species to the list of the Hyrcanian endemic mammals, which includes Crocidura caspica ( Zaitsev, 1991) , Apodemus hyrcanicus ( Vorontsov et al., 1992) , Microtus shelkovnikovi ( Nadachowski, 2007) , and the Iranian lineage of Glis glis ( Naderi et al., 2013) .
T. caucasica Satunin, 1908 and T. ognevi
Stroganov, 1948
Our results suggest a species status for the two allopatric lineages of the Caucasian mole. The northern lineage is found in the North Caucasus and presumably corresponds to two recognised subspecies: T. c. caucasica Satunin, 1908 from the Central North Caucasus (Kabardino-Balkaria) and T. c. orientalis Ognev, 1926 from the Northwest Caucasus (Adygea). The southern lineage is represented by a sample from northeastern Turkey (Hopa, Cumhuriyet Mahallesi, Artvin), and is evidently very close to T. c. ognevi Stroganov, 1948 (type locality is Bakuriani, Georgia), which is reported for adjacent western Transcaucasia ( Sokolov & Tembotov, 1989). Our additional analysis of cytb ( Fig. S3 View Figure 3 ) showed that sequences of Turkish specimens are very similar to the partial fragments obtained from archived museum specimens of T. c. ognevi originating from Batumi and Borzhomi ( South Georgia); the latter is only ∼ 20 km away from the type locality of ognevi . According to our results the time of the split between T. caucasica and T. ognevi dates back to the second half of the Pliocene (2.5–3 Myr). T. ognevi from Georgia and Turkey is morphologically distinct from true T. caucasica of the North Caucasus being characterized by larger size and more robust dentition ( Sokolov & Tembotov, 1989; Kryštufek & Vohralík, 2001). These differences induced Stroganov (1948) to attrib- ute ognevi to T. romana (as a subspecies T. r. ognevi ). The details of the distributions of T. caucasica and T. ognevi remain to be clarified.
T. levantis group
The most complex genetic structure was found in T. levantis . According to our nuclear data, the two major lineages within the Levant mole may correspond to distinct species that separated more than 2 Myr. The eastern lineage is distributed in the Caucasus, Transcaucasia, and the adjacent parts of northeastern Turkey (Cam Gecidi, Ardahan), while the western lineage occupies the major part of the Turkish range, stretching along the Black Sea shore westwards to the Marmara region. The nomenclatural treatment of these two lineages remains unclear because of the lack of genetic material from the type locality of T. levantis (Alindere, south of Trabzon), which is located between the sampled ranges of the and the eastern clades. If typical T. levantis belongs to the western clade, the senior synonym applicable for the eastern clade is T. transcaucasica Dahl, 1944 (type locality is Voskresenovka, Armenia). However, if the opposite is true, then no name is available for the western clade.
The Asian part of the range is believed to be disjointed (Kryštufek, 2001) with the gap between North– Northwest Turkey (Paphlagonia, i.e., between Zonguldak and Sinop). However, genetic data show no correspondence with this subdivision because the western clade is found both eastwards (Samsung) and westwards (Uludag) from the presumed gap .
Within the western and eastern clades, two additional subclades emerge from the mtDNA evidence, with the distance between them being 3–3.5%. Within the last one the substantial difference between morphologically specific lineage from the northwestern Caucasus (Adygea) and Kabardino-Balkarian sample give grounds to treat these subdivisions as subspecies and support the recognition of the Levant moles from northwestern Caucasus as T. l. minima Deparma, 1959.
Genetic differentiation in other species of Talpa
Due to a limited genetic data we could not validate the species status of the major phylogroups within T. caeca and T. stankovici which are characterized by deep phylogeographic breaks in each of these species.
The populations of blind moles from Apennines, Swiss Alps, and Balkans are treated as different subspecies: T. c. caeca Savi, 1822 (Central Italy), T. c. augustana Capolongo and Panasci, 1978 ( Switzerland and North Italy), T. c. steini Grulich, 1971 ( Montenegro), and T. c. hercegovinensis Bolkay, 1925 ( Herzegowina) ( Niethammer, 1990). The mtDNA differentiation observed for the Apennines and the Swiss Alps (western clade) versus the Balkans (eastern clade) is also supported by the divergence at conservative nuclear loci. According to our data the split of these lineages dates back to the Early Pleistocene (approximately 1.1 Myr). Notably, the karyotypes of the Blind mole in the Swiss Alps ( Switzerland, Ticina) and Balkans differ in the chromosome fundamental number ( Niethammer, 1990).
As demonstrated by Tryfonopoulos et al. (2009), the sample of T. stankovici from Greece contains three mitochondrial lineages (central Greece, Pelopónne¯sos, and northwestern Greece). The K2P distances among them vary from 5% to 7%. Based on the available mitochondrial data the tmrca can be tentatively estimated at no less than 1 Myr.
Thus, it is probable that the accumulation of the nuclear data on geographically representative samples of T. caeca and T. stankovici will lead to the revision of the taxonomic status of certain phylogroups.
Finally, we cannot exclude that the Spanish lineage of T. europaea is also a new cryptic species rather than a result of past mtDNA introgression from T. occidentalis ( Feuda et al., 2015) . The available nuclear data on this population is restricted to a single sequence of ApoB, which appeared to be a distinct haplotype. Mitochondrial data placed the split of these lineages at approximately 1 Myr. In the case of T. altaica and T. europaea s.s., the main mitochondrial phylogroups are also recovered with nuclear data. According to the mtDNA data ( Feuda et al., 2015), the split between the Italian and European lineages of T. europaea occurred ∼700– 800 Kya. Thus, it could not be the result of population fragmentation during the last glacial maximum, but most likely reflects a more ancient divergence during some previous interglacial periods. Moreover, Loy & Corti (1996) found a sudden change in mandible morphology over a short geographic range, highlighting a significant morphological difference of both the Spanish and Italian T. europaea versus other populations from western and central Europe, respectively. A morphological gap was also shown between Spain and western France and there is one abrupt change from the Italian populations to geographically adjacent populations in Switzerland, Germany, and Austria ( Loy & Corti, 1996).
Subgeneric taxonomy
As shown above, 12 species of the genus Talpa may be clearly divided into four divergent lineages. One of them ( T. altaica ) was previously attributed to a monotypic subgenus, Asioscalops Stroganov, 1941 , which was established on specific pelvic and dental morphology. The remaining species were included into Talpa s.s. In some cases, Asioscalops was regarded even as a separate genus (e.g., Yudin, 1989). Its isolated position was moderately supported by mitochondrial data ( Colangelo et al., 2010; our data) and cytogenetic evi- dence ( Kratochvíl & Král, 1972; Kawada et al., 2002). However, the nuclear results provide no support for placement of Asioscalops as sister to Talpa s.s. and, hence, for its recognition as a distinct taxon of equivalent rank to the rest of Talpa . Based on our data, we find no justification for splitting extant Talpa into several genera. If the subgeneric rank of the T. altaica lineage is maintained, then one should also reconsid- er the taxonomic status of two other lineages (i.e., the ‘davidiana’ and ‘caucasica’ groups).
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