Rook et al.—Pliocene Canidae from Sighnaghi, Georgia


vertebrate fauna (Vekua, 1972; Hemmer et al., 2004; Agustí et al., 2009; this paper) is as follows:


Reptilia Testudo cernovi transcaucasica Chkhikvadze, 1979 Aves Ioriotis gabuniae Burchak-Abramovich and Vekua, 1971; Struthio transcaucasicus Burchak-Abramovich and Vekua, 1971 Mammalia Rodentia: Hystrix cf. H. primigenia Wagner, 1848 Carnivora: Nyctereutes megamastoides (Pomel, 1842);


Eucyon sp., Vulpes cf. V. alopecoides (Del Campana, 1913); Ursus minimus Devèze and Bouillet, 1827; Lynx issiodorensis (Croizet and Jobert, 1828); Homotherium davitashvili Vekua, 1972; Puma pardoides Owen, 1846; Dinofelis cristata (Falconer and Cautley, 1836); Chasmaportetes lunensis (Del Campana, 1914); Perinium kvabebicus (Bendukize and Vekua, 2012) Arctiodactyla: Propotamochoerus provincialis (Gervais,


1859); Eucladoceros sp.; ?Pseudalces sp.; Procapreolus sp.; Ioribos aceros Vekua, 1972; Protoryx heinrichi Major, 1891; Oryx (Aegoryx) sp.; Parastrepsiceros sokolovi Vekua, 1968; Eosyncerus ivericus Vekua, 1972; Gazella postmitilinii Vekua, 1972


Perissodactyla: Hipparion rocinantis Hernández Pacheco,


1921; Stephanorhinus megarhinus (de Christol, 1834) Hyracoidea: Kvabebihyrax kachethicus Gabunia and


Vekua, 1966 Probiscidea: Anancus arvernensis (Croizet and Jobert, 1828)


Materials and methods


Studied specimens and comparative sample.—The present study is based on comparativemorphological analyses of the Canidae from Kvabebi. The examined and described fossils are housed at the GNM (see abbreviations below). We studied the collections of the IGF as comparative fossil material, and reviewed the relevant literature on Plio-Pleistocene canids (Del Campana, 1913; Thenius, 1954;Qiu andTedford, 1990; Teford andQiu, 1991, 1996;Koufos, 1997; Rook, 2009; Petrucci et al., 2013; Koufos, 2014). Extant specimens from theMZUF,MNHN, and GNMwere also used for morphological and morphometric comparisons. The fossil comparative sample includes specimens of


Nyctereutes donnezani (Depéret, 1890) from La Gloria and Layna; Nyctereutes megamastoides (Pomel, 1842) from the Lower Valdarno Basin, Villaroya, Dafnero 1; Nyctereutes vulpinus Monguillon et al., 2004 from St. Vallier; Nyctereutes sinensis (Schlosser, 1903) and Nyctereutes tingi Tedford and Qiu, 1991 from the Yushe Basin; Eucyon monticenensis (Rook, 1992) from the Cava Monticino and Venta del Moro; Eucyon adoxus (Martin, 1973) from Perpignan; Eucyon marinae Spassov and Rook, 2006 from Muhor-Erig; Eucyon davisi (Merriam, 1911) from North America and the Yushe Basin; Eucyon zhoui from the Yushe Basin; Eucyon odessanus (Odintzov, 1967) from Megalo Emvolon, Alatini and the Odessa Catacombs; Eucyon minor (Teilhard de Chardin and Piveteau, 1930) from Nihewan and Shamar; Vulpes alopecoides (Major, 1875) from the Upper Valdarno Basin, Pirro Nord and Dafnero-1; Vulpes praeglacialis (Kormos, 1932) and Vulpes praecorsac Kormos, 1932 from Spain, Hungary, and Ukraine.


1259 The extant comparative sample includes specimens of


Nyctereutes procyonoides (Gray, 1834), Vulpes vulpes (Linnaeus, 1758), and Vulpes lagopus (Linnaeus, 1758), which are housed in theMZUF, MNHN, and GNM.


Measurements and statistical analyses.—Cranio-dental and postcranial measurements were taken with a digital calliper to the nearest 0.1mm following von den Driesch (1976), with a few modifications.We used linear measurements and the ratio of the width/length of m1 to create scatter plots to test the affinity of the specimens of Vulpes and Eucyon from Kvabebi within the variability of Plio-Pleistocene species of these two genera. For Vulpes, we decided to include the extant species V. vulpes and V. lagopus for the widespread distribution of the former and size affinity of the latter to fossil species. Fossil species taken into consideration are European Plio-Pleistocene ones (e.g., V. praecorsac, V. praeglacialis,and V. alopecoides). We decided to keep convex hulls only for fossil species to identify their variability range compared to extant ones, which are considerably more variable. In the Eucyon scatter plot, we included six species of the Eurasian Pliocene Eucyon (E.marinae,E. adoxus,E.minor, E. davisi, E. zhoui,and E. odessanus) and preferred to leave only the convex hull of E. odessanus for its great variability. In these analyses, we used PAST software ver. 3.08


(Hammer et al., 2001; Hammer, 2016). We further clarified the specific attribution of the material of Nyctereutes by comparing the material from Kvabebi with that of other Eurasian Villafran- chian and extant species of the genus by means of two log-ratio diagrams (Simpson, 1941; Simpson et al., 1960). These graphs calculate the difference between the log-transformed mean value of each single variable for each species and a selected species that is chosen as the comparative baseline. The dental variables selected are P2-P3 L, P4-M2 L, and W for the upper teeth, and p2-p3 L, p4-m2 L, andW.We chose the extant N. procyonoides as a reference baseline and plotted the following fossil species: N. megamastoides from European sites; N. tingi and N. sinensis from China; and Nyctereutes from Kvabebi.


Repositories and institutional abbreviations.—IGF, Museum of Natural History, Geological and Paleontological section, University of Florence (Italy);K, Kvabebi site;MG, S. Janashia Museum of Georgia, part of the Georgian National Museum (Tbilisi, Georgia); MNHN, Museum National d’Histoire Nat- urelle, Paris;MZUF, Museumof Natural History, “La Specola” Zoology section, University of Florence (Italy).


Systematic paleontology


Anatomical abbreviations.—Postcranial: GL, general length; Bd, breadth of the distal epiphysis; Bp, breadth of the proximal epiphysis; BPC, greatest breadth across the coronoid process; DC, depth of the caput femoris; Dd, depth of the distal epiphysis; Dp, depth of the proximal epiphysis; DPA, depth across the processus anconaeus


Order Carnivora Bowdich, 1821 Suborder Caniformia Kretzoi, 1943


Family Canidae Fischer von Waldheim, 1817 Subfamily Caninae Fischer von Waldheim, 1817


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