Apodemus sylvaticus (Linnaeus 1758)

[Mus] sylvaticus Linnaeus 1758, Syst. Nat., 10th ed., Vol. 1: 62.

Type Locality: Sweden, Uppsala (neotype designated by Zagorodnyuk [1993]).

Vernacular Names: Long-tailed Field Mouse.

Synonyms: Apodemus albus (Bechstein 1801); Apodemus algirus (Pomel 1856); Apodemus alpinus (Burg 1921); Apodemus bergensis (Krausse 1921); Apodemus butei Hinton 1914; Apodemus callipides (Cabrera 1907); Apodemus candidus (Bechstein 1796); Apodemus celticus (Barrett-Hamilton 1900); Apodemus chamaropsis (Levaillant 1857); Apodemus charkovensis (Migulin 1936); Apodemus clanceyi Harrison 1947; Apodemus creticus Miller 1910; Apodemus cumbrae Hinton 1914; Apodemus dichruroides Miric 1960; Apodemus dichrurus (Rafinesque 1814); Apodemus dichrurus Cabrera 1921; Apodemus eivissensis Alcover 1977; Apodemus fiolagan Hinton 1914; Apodemus flaviventris (Petrov 1943); Apodemus flavobrunneus (Hilzheimer 1911); Apodemus fridariensis (Kinnear 1906); Apodemus frumentariae Sans-Coma and Kahmann 1977; Apodemus ghia Montagu 1923; Apodemus grandiculus Degerbol 1939; Apodemus granti Hinton 1914; Apodemus griseus (Mina Palumbo 1868); Apodemus hamiltoni Hinton 1914; Apodemus hayi (Waterhouse 1838); Apodemus hebridensis (de Winton 1895); Apodemus hermani Felten and Storch 1970; Apodemus hessei Miric 1960; Apodemus hirtensis (Barrett-Hamilton 1899); Apodemus ifranensis Saint Girons and Bree 1962; Apodemus ilvanus Kahmann and Niethammer 1971; Apodemus intermedius (Bellamy 1839); Apodemus isabellinus (Mina Palumbo 1868); Apodemus islandicus (Thienemann 1824); Apodemus krkensis Miric 1968; Apodemus larus Montagu 1923; Apodemus leptodus Kretzoi 1956; Apodemus leucocephalus (Bechstein 1796); Apodemus maclean Hinton 1914; Apodemus maximus (Burg 1925); Apodemus milleri de Beaux 1926; Apodemus nesiticus Warwick 1940; Apodemus niger (Bechstein 1796); Apodemus parvus (Bechstein 1793); Apodemus pecchioli (Pecchioli 1844); Apodemus reboudia (Loche 1867); Apodemus rufescens Saint Girons and Bree 1962; Apodemus spadix Fritsche 1934; Apodemus thuleo Hinton 1919; Apodemus tirae Montagu 1923; Apodemus tural Montagu 1923; Apodemus varius (Bechstein 1796); Apodemus vohlynensis (Migulin 1938) .

Distribution: European and N African: Europe north to Scandinavia; south to NW Turkey (Thrace and NW Anatolia; Filippucci et al., 1996; Pamukoglu and Albayrak, 1996; specimens in USNM); and east to C Belarus, E Ukraine, and closely adjacent W Russia, which is the easternmost limit of the species (Zagorodnyuk, 1993); see maps in Zagorodnyuk et al. (1997:39), Mezhzherin (1997 a:33), Niethammer (1978 c:341), and Mitchell-Jones et al. (1999:275). Range in N Africa extends from Atlas Mtns in Morocco east across Algiers to Tunisia (Aulagnier, 1991; Aulagnier and Thevenot, 1986; Kock and Felton, 1980). Also found on Iceland; Britain, Ireland, and numerous nearby islands; Aegean islands (Kryštufek, 2002 a; Özkan and Kryštufek, 1999); some islands in the Tuscan Arch. (De Marinis et al., 1996); Sardinia, Corsica (Masseti, 1993), and other Mediterranean islands (Alcover and Gosalbez, 1988; Amori, 1993; Amori and Masseti, 1996; Cheylan, 1991).

Conservation: IUCN – Lower Risk (lc).

Discussion: Sylvaemus group. The geographic range as mapped by Corbet (1978 c) east of Belarus and E Ukraine reflects distributions of other species ( A. uralensis, A. witherbyi, A. pallipes, and A. rusiges) once included within A. sylvaticus; " A. sylvaticus is virtually absent from the entire area of the Middle East" (Macholán et al., 2001 b:806). Contrary to published records, A. sylvaticus is not part of the modern Israeli fauna (Filippucci et al., 1989). Tchernov (1979, 1986, 1994, 1996), however, maintains it is and is also represented there by fossils from middle to late Pleistocene cave sediments (late Acheullan to Upper Mousterian), and from 40,000 to 10,000 years B.C. During that Pleistocene interval, A. mystacinus was found along with A. sylvaticus in the same caves, but A. flavicollis was usually absent (Tchernov, 1979). Tchernov’s " sylvaticus " probably represents A. witherbyi (B. Kryštufek, 2002, in litt.; see account of A. witherbyi), which along with A. mystacinus and A. flavicollis are part of the modern Israeli fauna (see accounts of latter two). An Arabian Peninsula record at Qatar is based on Mus (Kock and Nader, 1990) . See Kowalski (2001) for the inclusion of Hungarian leptodus .

The population of A. sylvaticus on Corsica arrived there (presumably by passive human transport from peninsular Italy through the Tyrrhenian islands, Michaux et al., 1996 a) at the beginning of the third millennium B.C. and occurred together with the endemic murine Rhagamys orthodon and arvicoline Microtus henseli, both of which vanished by the end of the first millenium B.C. (Vigne, 1992; Libois et al., 1993; and references cited in those reports). The population on Sicily represents a Holocene anthropogenic introduction (about 750,000 years ago); its origin is not peninsular Italy or W Europe but possibly North Africa or the E Mediterranean basin (Michaux et al., 1998). Allozymic variation among European and North African samples of A. sylvaticus (Filippucci, 1992; Filippucci et al., 2002; Libois et al., 2001) indicated high genetic homogeneity among samples of North African populations and recent derivation from SW Europe most likely through anthropogenic introduction, results corroborated by Michaux et al. (2003) based on phylogenetic analyses of mtDNA cytochrome b sequences. North African populations are no older than Holocene (Kowalski and Rzebik-Kowalska, 1991) or late Pleistocene (Ouahbi et al., 2001). Dobson (1998, 2000) claimed A. sylvaticus is unknown from the Maghreb fossil record prior to the Iron Age (about 3000-2300 years before present) and its presence in North Africa is likely the result of human intervention, but Ouahbi et al. (2001) documented it from late Pleistocene sediments in N Morocco.

Allozymic (Byrne et al., 1990; Fernandes et al., 1991; Gemmeke, 1981 a) and morphometric (Alcantara, 1991; Murbach, 1979) intrapopulational analyses provided results of differentiation within A. sylvaticus and change in its evolutionary history (see Berry, 1973, and references therein); other allozymic and morphometric analyses of European populations presented by Gemmeke (1981 a) and Nauroz (1984); high genetic diversity within samples from W Europe as measured by mtDNA cytochrome b sequences documented by Reutter et al. (2003); and phylogeography based on study of mtDNA cytochrome b sequences, which suggested postglacial colonizations of all of Europe from Iberian and S France refuge regions (Michaux et al., 2002 a, 2003). Several studies have focused on aspects relevant to systematics of A. sylvaticus: morphometric and molecular analyses of samples from Mediterranean region in the context of taxonomy and biogeography (Libois et al., 1993; Michaux et al., 1996 a, b, 1998; Sarà and Casamento, 1995 b), adaptive latitudinal trends in mandible shape (Renaud and Michaux, 2003), insular gigantism in the Mediterranean (Alcover and Gosalbez, 1988; Libois and Fons, 1990, and references cited therein; Michaux et al., 2002 b), B chromosomes in samples from Czech Republic (Zima et al., 1997 d), sex-chromosome heterochromatin variation (Nová et al., 2002), epigenetic characteristics and divergence among Bulgarian populations (Markov and Chassovnikarova, 1999), morphometric analyses of samples from Iberian Peninsula (González-Esteban et al., 1996), variability in allometric relationships between body mass and skull size in Spanish samples (Alcantara et al., 1991), and karyotypes of Macedonian sample (Zima et al., 1997 a). Conspecificity of krkensis with A. sylvaticus documented by Williams et al. (1980) and Dolan and Yates (1981). The name charkovensis, usually associated with A. uralensis (e. g., Mezhzherin, 1997 b; Zagorodnyuk, 1992 b), was identified by Zagorodnyuk (1993) as referring to the easternmost population of A. sylvaticus; he also selected a lectotype. Based only on chromosomal data, Orlov et al. (1996 a, b) regarded vohlynensis as a separate species within the A. sylvaticus superspecies, and one that occurs in Central Europe, the Balkans, and east to the Dnepr basin. Distribution and population density of A. sylvaticus at the easternmost limits of its range (E Ukraine) and comparisons with A. uralensis documented by Naglov (1995). The range expansion of A. sylvaticus into forested tracts that were originally clay semidesert in the Trans-Volga region outlined by Bykov (1990). See account of A. flavicollis for reports documenting morphological, chromosomal, and molecular contrasts between it and the closely related and morphologically similar A. sylvaticus .

European populations reviewed by Niethammer (1978 c) and Mitchell-Jones et al. (1999); N Eurasian segment by Mezhzherin (1997 a, b). Literature covering ecological aspects of A. sylvaticus is voluminous, extending back for decades, and only a few current regional reports focusing on distribution, taxonomy, habitat, and other aspects are listed here for populations in: Pyrenees and NE Spain (Castién and J. Gosálbez, 1992; Torre et al., 1996), Portugal (Santos-Reis and Mathias, 1996), Netherlands (Thissen and Hollander, 1996; Wammes, 1992 a), Belgium (Libois, 1996), N Germany (Dolch et al., 1994), Slovakia (Danko, 1994; Kminiak, 1996; Mošanský, 1994; Stanko and Mosansky, 2000; Stanko et al., 1994), Czech Republic (And�ra and ervený, 1994; Smaha, 1996; Zima and And�ra, 1996); Austria (Bauer and Spitzenberger, 1996), Switzerland (Hausser, 1995; Maurizio, 1994), Italy (Amori et al., 1999, 2002 b; Cagnin et al., 1996; Cantini, 1991; Cresti et al., 1994; Cerone and Aloise, 1994; Locatelli and Paolucci, 1996), E Baltic region (Miljutin, 1997, 1998; Timm et al., 1998), Serbia and Montenegro (Petrov, 1992), Albania (Prigioni, 1969); Bulgaria (Peshev, 1996), Translvanian Romania (Istrate, 1998), Slovenia (Kryštufek, 1991), Thrace (Greece; Vohralík, 1992) and NE Turkey (Kryštufek and Vohralík, 2001). North African populations reviewed by Kock and Felten (1980) and Kowalski and Rzebik-Kowalska (1991); the latter authors also point out that algirus Pomel, 1856 and chamaeropsis Loche, 1867 may refer to species of Mus or Gerbillus rather than Apodemus . Abundance of Algerian populations in different habitats compared with Mus spretus reported by Khidas et al. (2002).