Microtus (Terricola) subterraneus de Selys-Longchamps 1836

Wilson, Don E. & Reeder, DeeAnn, 2005, Order Rodentia - Family Cricetidae, Mammal Species of the World: a Taxonomic and Geographic Reference (3 rd Edition), Volume 2, Baltimore: The Johns Hopkins University Press, pp. 955-1189 : 1016

Microtus (Terricola) subterraneus de Selys-Longchamps 1836

Microtus (Terricola) subterraneus de Selys-Longchamps 1836 , Essai Monogr. sur les Campagnols des Env. de Liege.: 10.

Type Locality: Belgium, Liege, Waremme.

Vernacular Names: Common Pine Vole.

Synonyms: Microtus (Terricola) atratus (Stein 1931) ; Microtus (Terricola) brauneri Martino 1926 ; Microtus (Terricola) capucinus (Miller 1908) ; Microtus (Terricola) dacius (Miller 1908) ; Microtus (Terricola) dinaricus Kretzoi 1959 ; Microtus (Terricola) ehiki (Wettstein 1927) ; Microtus (Terricola) fingeri (Neuhäuser 1936) ; Microtus (Terricola) fusca (Fatio 1900) ; Microtus (Terricola) grafi Brunet-Lecomte, Nadarowski, and Chaline 1992 ; Microtus (Terricola) hercegovinensis ( Martino 1940) ; Microtus (Terricola) hungaricus (Ehik 1926) ; Microtus (Terricola) incertoides (Wettstein 1927) ; Microtus (Terricola) incertus (Sélys-Longchamps 1841) ; Microtus (Terricola) klozeli (Ehik 1942) ; Microtus (Terricola) kupelwieseri (Wettstein 1925) ; Microtus (Terricola) martinoi (Ehik 1935) ; Microtus (Terricola) matrensis (Ehik 1930) ; Microtus (Terricola) mustersi ( Martino 1937) ; Microtus (Terricola) neglectus Petrov 1992 ; Microtus (Terricola) neuhauseri (Martino and Paspalev 1955) ; Microtus (Terricola) nyirensis (Ehik 1930) ; Microtus (Terricola) rufescente-fuscus (Schinz 1845) ; Microtus (Terricola) rufofuscus (Schinz 1845) ; Microtus (Terricola) subterraneoides Petrov 1992 ; Microtus (Terricola) transsylvanicus (Ehik 1924) ; Microtus (Terricola) transvolgensis (Schaposchnikov and Schanev 1958) ; Microtus (Terricola) ukrainicus (Vinogradov 1922) ; Microtus (Terricola) wettsteini (Ehik 1926) ; Microtus (Terricola) zimmermanni (Matschie 1924) .

Distribution: Atlantic coasts of N and C France through S Netherlands ( Ligtvoet and Straetmans, 1992), across C Europe ( SW Bohemia [Andĕra and Červený, 1994]; Switzerland [ Maurizio, 1994; Hausser, 1995]; Slovakia [ Danko, 1994; Kminiak, 1996; Mošanský, 1994; Stanko, 1995; Stanko and Mošanský, 1994, 2000; Stanko et al., 2000]; Czech Republic [Šmaha, 1996]; Italy [ Amori et al., 1999; Locatelli and Paolucci, 1996 a]) to Ukraine and the Don River ( Gromov and Erbajeva, 1995), south through Romania and Balkan region into N and SE Greece, and Turkey (mountains in N, W and S but not on Anatolian Plateau; Kryštufek and Vohralík, 2001) eastward to south of Trabzon (Macholán et al., 2001 a); isolated populations in Russia (near St Petersburg; Zagorodnyuk, 1989) and Estonia ( Miljutin, 1997, 1998; Timm et al., 1998). Absent from Mediterranean coast and islands.

Conservation: IUCN – Lower Risk (lc).

Discussion: Subgenus Terricola , subterraneus species group (Chaline et al., 1988; Zagorodnyuk, 1990). Reviewed by Niethammer (1982 g). Karyotypic information available in Sablina et al. (1989), Zima (1986), Zima and Kral (1984 a), and Zima et al. (1997 a); biochemical comparisons among M. subterraneus , M. felteni , and other voles implemented by Gill et al. (1987). Synonyms follow Zagorodnyuk (1989), who speculated that the Romanian dacius is a separate species, but Brunet-Lecomte et al. (2001) regarded the minor cytogenetic differences as inconclusive. The taxon grafi was described as a species from late Pleistocene fossils from Bulgaria (Brunet-Lecomte et al., 1992) but is now regarded as a chronological subspecies of M. subterraneus ( Brunet-Lecomte et al., 2001) . Present in the British Isles during early Pleistocene interglacials ( Yalden, 1999).

Regional studies have addressed the discrimination of M. subterraneus . Pelage and vibrissal contrasts with M. tatricus documented by Kratochvíl (1969). Distribution and morphological variation reported by Zagorodnyuk (1989, 1992 a) in taxonomic overview of species in subgenus Terricola ; he clarified distribution of the 2n = 52 and 2n = 54 chromosomal morphs in the Baltic region and morphometrically contrasted M. subterraneus with M. agrestis , M. arvalis , M. levis (formerly rossiaemeridionalis ), and M. oeconomus ( Zagorodnyuk and Mezhzherin, 1992) . Morphometric analysis of m1s vindicated specific separation of M. subterraneus from M. multiplex and M. liechtensteini (Brunet-Lecomte and Kryštufek, 1993), a result also indicated by sequence analysis of the mitochondrial control region ( Haring et al., 2000). Morphometric distinction between M. subterraneus and M. liechtensteini and their distributions in Slovenia discussed by Kryštufek (1991). Vohralík and Sofianidou (1992 a) identified M. subterraneus from SE Greece (Thrace region) but expressed skepticism about the reliability of size distinctions used to segregate it from M. majori . Macholán et al. (2001 a) utilized karyotypic and allozymic analyses to document the distributions of the 2n = 52 and 2n = 54 karyomorphs of M. subterraneus and their chromosomal and genetic distinctions from M. majori (2n = 54); low genetic distance between the two suggests a recent divergence, only 170-350 thousand years ago. Their ranges closely approach one another in NE Turkey but sympatry not yet demonstrated (Macholán et al., 2001 a).

The substantial genetic and morphological variation of M. subterraneus throughout its range, especially the Balkans (Kryštufek et al., 1994), prompted Macholán et al. (2001 a:40) to propose this view: "taking into account close karyotypic and allozymic relationships (and the fact that some intraspecific genetic distances within M. subterraneus are higher than those between M. subterraneus and M. majori ), we cannot rule out a hypothesis that M. subterraneus is paraphyletic, with M. majori (and, by the same token, M. daghestanicus ) being one of its constituents." Further research should explore their reservations .