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Sotnikova, M.V.; Baigusheva, V.S.; Titov, V.V.
Carnivores of the Khapry Faunal Assemblage and Their Stratigraphic Implications 
2002  Stratigraphy and Geological Correlation (10 ): 375-390

Carnivores of the Khapry faunal assemblage that is commonly considered as an analogue of the middle Villafranchian West European fauna (Late Pliocene, Zone MN17) were revised. Members of nine Carnivora genera are described in detail, and five of them are first included in the list of assemblage taxa. Carnivores of the Khapry assemblage are represented by _Nyctereutes megamastoides _(Pomel), _Canis cf. senezensis_ Martin, Lutra sp., _Pannonictis nestii_ (Martelli), _Pliocrocuta perrieri_ (Croizet et Jobert), Pachycrocuta brevirostris (Aumard), Homotherium crenatidens (Fabrini), _Acinonyx pardinensis_ (Croizet et Jobert), and _Lynx issiodorensis_ (Croizet et Jobert). The Khapry carnivore assemblage includes taxa characteristic of the Late Pliocene Eurasian faunas. Its lower stratigraphic limit corresponds to the beginning of the middle Villafranchian, as it follows from presence of Homotherium crenatidens with more advanced dental characteristics than those of homotheres from the lower Villafranchian of Eurasia. The occurrence of Nyctereutes megamastoides that was not encountered above the middle Villafranchian bounds the upper stratigraphic limit of the Khapry Carnivora. The "Canis-event" and "Pachycrocuta-event" are considered to be important records around the middle-upper Villafranchian boundary in Europe. Presence of _Canis senezensis_ is characteristic of the final stage of the middle Villafranchian in Western Europe, and dispersal of Pachycrocuta brevirostris is typical of the lowermost late Villafranchian, whereas the wolflike forms of Canis and Pachycrocuta appeared somewhat earlier in Asia. The Khapry faunal assemblage as a whole is characterized by a certain similarity with Asian analogues of the middle Villafranchian faunas.

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