Rook et al.—Pliocene Canidae from Sighnaghi, Georgia


considered the ancestor of the extant Vulpes corsac (Linnaeus, 1768) (see Kormos, 1932). Another well-known European species is V. alopecoides described by Major, 1975 from Tasso (Upper Valdarno), and recovered from St. Vallier, Senéze, France (Viret, 1954), and Villaroya and La Puebla de Valverde, Spain (Kurtén and Crusafont Pairó, 1977). Kurtén (1968) suggested the synonymy between V. alopecoides and V. praeglacialis. Never- theless, the literature reveals no unanimous opinion on the status of these two species (Bonifay, 1971; Rabeder, 1976; García and Arsuaga, 1999). The relationship between the two taxa is beyond the scope of this paper; therefore, we have included both in the comparative analyses. MG 29-2013/461 represents the first occurrence of a member the vulpine taxon V. alopecoides,the most widespread fox species in the early Pleistocene of Western Europe.


Eucyon.—The genus Eucyon appears in the late Miocene (late Clarendonian) record of North America with the species Eucyon davisi (Merriam, 1911). Its geographic range remained limited to North America until the latest Miocene, when the genus spread into the Old World, occurring in Central Asia, Africa, and Western Europe (the “Eucyon event”, Sotnikova and Rook, 2010). Eucyon davisi disappeared in North America in the earliest Pliocene (latestHemphillian), and survived into the early Pliocene of easternAsia (China andMongolia). The genus Eucyon reached a relative high diversity in the Pliocene of Eurasia (Rook, 2009; Sotnikova and Rook, 2010), surviving until the late Pliocene in China (E. minor), Mongolia (E. marinae), Tadzhikistan (E. kuruksaensis), Kazakhstan (Eucyon cf. E. odessanus), and southeastern Mediterranean regions (E. Eucyon cf. odessanus, Sarikol Tepe, Turkey). The occurrence of a Eucyon representative within the fauna


of Kvabebi completes our knowledge of the late Pliocene history of the advanced eucyons surviving in Western Europe and central Asia. The Kvabebi finding predates the explosive radiation of the wolf- and coyote-sized Canini that took place in Central Asia (reflected in the appearance of Canis teilhardi, C. longdanensis, C. brevicephalus, and Sinicuon cf. S. dubius), which continued until the end of the Pliocene (Qiu et al., 2004; Sotnikova and Rook, 2010).


Conclusions


The evolutionary history of the Pliocene Canidae in Eurasia is characterized by taxonomic diversification and range expansion. Diversification of Vulpini and Canini showed a maximum in Central Asia at the beginning of the early Pliocene, as evinced by the appearance of several new taxa (Nyctereutes tingi, V. beihaiensis, V. qiuzhudingi, Nurocyon chonokhariensis, Eucyon zhoui) and by the increase in frequency of other species (like the well-known Eucyon davisi). In Western Europe, at the beginning of the Pliocene, the canid


record is limited to the primitive vulpine Nyctereutes donnezani. The peak of canid diversification in Western Europe is recorded later in the early Pliocene, during the late Ruscinian (MN 15), with the differentiation of the advanced raccoon-dog Nyctereutes megamastoides, the appearance of the small fox V. praecorsac,and the emergence of the enigmatic “Canis” michauxi, togetherwith the advanced Eucyon adoxus and Eucyon odessanus.


1269 The revision of the Kvabebi assemblage documents


the earliest occurrence of the typical (later) early Pleistocene species Vulpes alopecoides in the European fossil record. The contemporary early occurrence at Kvabebi of Vulpes with Nyctereutes and Eucyon clearly supports the established niche partitioning among these three sympatric canids.


Acknowledgments


We dedicate this paper to the late Professor A. Vekua for his pioneering work on Kvabebi. The Italian Ministry for Foreign Affairs (DGPCC-V) is acknowledged for financially supporting Italian paleontological research in Georgia. Comparative data were partly derived from Projects FR-TAF-3311, BE-TAF-3607, HU-TAF-6520 awarded to LR and SBL by the SYNTHESYS initiative (http://www.synthesys.info) and financed by theEuropean Community Research Infrastructure Action under the FP7 “Capacities” Program.


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