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Turner, A.
The evolution of the guild of larger terrestrial carnivores during the Plio-Pleistocene in Africa
1990  Geobios (23): 349-368

The guild of African larger carnivores (_Felidae, Hyaenidae_ and _Canidae_) that existed between c. 4.0 and 1.0 Ma had twice the number of species that exist there today. Lion and leopard appear to have originated in Africa soon after 4.0 Ma, and were confined to that continent until they appeared in Galerian faunas of Europe around 0.9 Ma. Cheetah,; or cheetah-like forms were widely dispersed in both the Old and New World from the mid-Pliocene onwards, and their origins remain unclear. Spotted hyaenas are first known in Africa just after 4.0 Ma and appear in Europe in Galerian faunas, although they may have made an earlier dispersion into Asia. Brown and striped hyaenas appear to have originated in Africa by about 3.0 Ma. The former may be conspecific with the Eurasian Pachycrocuta perrieri, but while the latter appears to have dispersed into Asia during the Pleistocene, records of its presence in Europe require revision. Three other hyaenid species, P. brevirostris, Chasmaporthetes nitidula and C. silberbergi, along with archaic cats of the genera Homotherium, Megantereon and Dinofelis, are known from African Mid Pliocene - Lower Pleistocene deposits and also appear to have been part of an Afro-Eurasian dispersion. The pattern of evolution in the carnivore guild has broad correlations with the patterns of climatic change and the evolution of potential prey species. 

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