The taxonomic status of badgers (Mammalia, Mustelidae) from Southwest Asia based on cranial morphometrics, with the redescription of Meles canescens Author Abramov, Alexei V. Author Puzachenko, Andrey Yu. text Zootaxa 2013 3681 1 journal volume 10.11646/zootaxa.3681.1.2 6983f9f1-7d67-45cb-902f-1e7e8cdb614e 1175-5326 222960 035D976E-D497-4708-B001-9F8DC03816EE Meles canescens Blanford, 1875 Meles meles minor Satunin, 1905 . Borzhomi, Georgia . Meles meles arcalus Miller, 1907 . Lassethe Plain, Crete, Greece . Meles meles rhodius Festa, 1914 . Koskino, Rhodes, Greece . Meles meles ponticus Blackler, 1916 . Scalita, 30 miles south of Trebizond, north-east Asia Minor [= Turkey ]. Meles meles severzovi Heptner, 1940 . Arkit, Chodscha-Ata River, Sary-Chilek Lake, Chatkal Ridge, Tien Shan Mts. [=Osh Province, Kirgizia ]. Meles meles canescens natio bokharensis Petrov, 1953 . Eastern Bukhara [= Tajikistan ]. Unavailable name (infrasubspecific rank). Type material and type localities . Lectotype ( Barrett-Hamilton 1899: 383 ): BMNH 74.11.21.1, skull and skin, Abadah, Persia [= Iran ]. Blanford (1875) described this taxon as a full species; later Barrett-Hamilton (1899) reduced Blanford’s M. canescens to subspecific rank. Diagnosis. This species differs from both M. meles and M. leucurus in a combination of cranial and dental characters ( Fig. 3 ). The upper molars have the morphotypes of “ meles - type ” (sensu Baryshnikov et al . 2003 ) with the well-developed external notch between metacone and metaconule. The upper first premolars Pm1 are often absent, whereas the first lower premolars Pm1 are usually present. The second lower premolar is large, usually onerooted or with two fused roots, but sometimes has two roots (as in M. meles ). The upper fourth premolar Pm4 lacks a small cusp on the precingulum at the base of the paracone lingual anterior ridge, and a lingual ridge runs from the paracone apex to the tooth inner projection in front of a well developed protocone. M. canescens is markedly smaller than M. meles , especially from the subspecies M. m. taxus , which is parapatric in SW Asia. From the latter species, it differs in having unflattened auditory bullae, shorter rostrum and mandible, and low crania. From M. leucurus , it differs in having the shape of upper molars “ meles - type ”, presence of first premolars, a large upper canine, narrower zygomatic arches, and wider auditory bullae. The skin can be distinguished from those both of M. leucuru s and of M. anakuma in the type of facial mask, which resembles that of M. meles (Abramov 2003) . Wide black or black-brown longitudinal stripes on either side of the head run from the snout’s tip over eye and ear (both covered from above and below) and a pure white facial stripe is in between the two black bands, covering the head’s back and partly the neck. The snout, cheeks and the ears’ tips white. Overall coloration is paler as comparison to that of M. meles . FIGURE 3. Male skulls of three taxa of the Eurasian badgers: A. Meles meles taxus (ZIN 35056, Leningrad Province, Russia), B. Meles leucurus leucurus (ZIN 1148, Eastern Kazakhstan), C. Meles canescens (ZIN 23800, Azerbaijan). Scale bar 5 cm. Distribution . M. canescens is known from the Caucasus ( Armenia , Georgia , and Azerbaijan ) including the northern slope of the Great Caucasus Mountain Range, Turkey , Iran , Iraq , Syria , Lebanon , Israel , northern Afghanistan , Turkmenistan (Kopetdagh, Balkhany, and Kugitang Mts.), Kirgizia , Uzbekistan and Tajikistan (the foothills of Western Tien Shan Mts. and Pamir-Alai Mts.). It is also found in the Mediterranean islands Crete and Rhodes ( Fig. 4 ). To the east of Caspian Sea, the ranges of M. canescens and M. leucururs are separated by arid desert regions (Kara Kum and Kyzyl Kum deserts). The contact zone between two badger species in Middle Asia is located in the Western Tien Shan Mts. ( Abramov & Puzachenko 2007 ). M. canescens occurs in the foothills of Western Tien Shan (Karzhantau, Ugam, Chatkal, Kuraminsky, and Turkestan ridges). The Asian badger M. leucurus occupies the northern, central and east ridges of Tien Shan Mts. (Talass-Alatau, Kirghiz-Alatau, Kungei-Alatau, Terskei-Alatau, Zailiysky, and Fergana ridges) and plains situated west and north of Western Tien Shan. In the sympatric zone, in the southeast regions of Uzbekistan , two species substantially differ in their biotope preferences. M. canescens occupies mountain biotopes, whereas M. leucurus inhabits plains and semi-deserts. A clear geographic border in the Northern Caucasus between M. canescens and M. meles has not yet been clarified. In some areas of the Northern Caucasus they can occur sympatrically, the possible hybrids with mixed characters were found in north-eastern part of the Northern Caucasus ( Abramov & Puzachenko 2007 ). FIGURE 4. Distribution ranges of the SW Asian badger ( Meles canescens ), Asian badger ( Meles leucurus , a part of its range), and European badger ( Meles meles , a part of its range). Locations of the specimens used in this study are shown (for M. canescens see Appendix). Evolution of the Eurasian badgers . The Meles lineage appears to have evolved in the temperate forest of Asia ( Kurtén 1968 ). Badgers may have originated from the Pliocene genus Melodon Zdansky in China ( Viret 1950 ; Kurtén 1968 ). Earliest known representatives of Meles are the Late Pliocene M. chiai Teilhard de Chardin from China and M. thorali Viret from France ( Kurtén 1968 ). Meles chiai is characterized by the absence Pm1/Pm1 and М1, with the well expressed external notch that is typical for the recent Asian badger M. leucurus . The European M. thorali bears the mixed set of characters (first premolars Pm1/Pm1 not reduced, Pm2 long, with two roots—as in the recent European badger M. meles , but the morphotypes of Pm4 and М1 are typical for M. leucurus ). Meles iberica Arribas et Garrido from Plio-Pleistocene of Spain and M. dimitrius Koufos from the Early Pleistocene of Greece appear to be similar (probably, conspecific) to M. thorali , as Pm1 are present, Pm2 large and М1 with an external notch. Meles hollitzeri Rabeder from the Early Pleistocene deposits of Central Europe ( Austria , Germany ) already had the characters which are typical for M. meles (the presence of Pm1, large Pm2, meles -morphotypes of Pm4 and М1). Presumably, the badger close to M. chiai was an ancestor of the recent forms of Meles . This ancestral form had a wide Palearctic distribution during Late Pliocene. Paleontological evidence has confirmed that Meles reached the Iberian Peninsula before the beginning of the glacial-interglacial cycles in the northern Hemisphere (ca. 2.6 Ma) ( Madurell-Malapeira et al . 2009 ), thus indicating that this genus was widely distributed during the Early Villafranchian, soon after its first appearance in Eastern Asia. At the end of Pliocene—Early Pleistocene, this ancestral form should have split in to two lineages: the western, or European one, and the eastern, or Asian one ( Baryshnikov et al . 2003 ; Abramov & Puzachenko 2005 ). According to the analysis of mitochondrial control region sequences in populations throughout Eurasia, the first split separating the meles-canescens and leucurus-anakuma clusters occurred between 2.87 and 0.55 Mya, most probably at the end of the Pliocene, and just before the beginning of the glacial ages ( Marmi et al . 2006 ). On the basis of paleontological data, it has been suggested that the split between the European and Asian badgers took place in the Middle to Late Villafranchian boundary (ca. 1.8 Ma) or slightly before, through a vicariance process prompted by palaeoclimatic changes ( Madurell-Malapeira et al . 2011b ). The western lineage then evolved through M. thorali (including M. iberica ) to M. hollitzeri and then to recent M. meles and M. canescens , whereas the Eastern lineage has evolved to the recent M. leucurus and M. anakuma . According to a recent taxonomical review of the European Plio-Pleistocene badgers ( Madurell-Malapeira et al . 2011a , b ), all Late Villafranchian European badger remains were assigned to M. meles . According to mtDNA data ( Marmi et al . 2005 , 2006 ), badgers from the Middle East ( M. canescens ) diverged from the European badgers ( M. meles ) between 2.37 and 0.45 Ma and the Japanese ( M. anakuma ) and Asian ( M. leucurus ) badgers diverged between 1.09 and 0.21 Ma. Such a separation could have resulted from mountain glaciations, the extension of the Caspian Sea, and other landscape changes during the glacial epochs, and also from other paleoclimatic factors. The Middle Eastern badgers were apparently isolated from the European ones by the Greater Caucasus Mountain Range, and the Bosporus and Dardanelles straits, which prevented a genetic information exchange.