An articulated skeleton of Eucyon monticinensis (Carnivora: Canidae) from the latest Miocene of Verduno (Italy)

The genus Eucyon includes several Late Miocene–Early Pleistocene canid species, the earliest of which, Eucyon davisi, is regarded as basal in the radiation of the Canini. Eucyon monticinensis is known from Italy (Late Miocene) and Spain (Early Pliocene). Here we report on an articulated skeleton from Verduno (Italy) referred to E. monticinensis based on comprehensive comparisons. The Verduno skeleton stands out as the most complete specimen of Eucyon from Europe, and as one of the most complete in the world. The fossil has allowed us to investigate and reconstruct in detail the morphology of E. monticinensis. Based on morphological similarity, Eucyon debonisi is regarded as a junior synonym of E. monticinensis. Computed tomography-based analysis provides some interesting insights into the comparative anatomy of the brain and frontal sinuses of Eucyon and other canids. We further present a revised phylogeny of the Canidae based on morphological characters. Eucyon davisi is confirmed as the most basal species in a monophyletic clade that also includes E. monticinensis, Eucyon zhoui, and Eucyon adoxus. Eucyon khoikhoi from South Africa and Eucyon ferox from North America show phylogenetic relationships and morphological affinities with the Vulpini and crown-Canini, respectively, which raises questions regarding their systematic status.