Unlike the Asian and North American Pliocene record, fossil occurrences of Canidae in Europe (and Africa) are uncommon and fragmentary. The revision of canid material from the late Pliocene site of Kvabebi (eastern Georgia) revealed the contemporaneous occurrence of three different taxa: (1)Nyctereutes megamastoides(a derived species of the Eurasian Pliocene raccoon dog-like canids); (2)Vulpescf.V.alopecoides(representing the first occurrence of a member of the vulpine taxonV.alopecoides, a species that was the most widespread fox in the early Pleistocene in western Europe); and (3)Eucyonsp. The latter occurrence at Kvabebi completes our knowledge of the late Pliocene evolutionary history of the latest representatives of the genus in Western Europe and Central Asia. Our revision of Kvabebi canids registers a previously undocumented case of established niche partitioning among early Pliocene sympatric Canidae.

The renowned site of Dmanisi in Georgia, southern Caucasus (ca. 1.8 Ma) yielded the earliest direct evidence of hominin presence out of Africa. In this paper, we report on the first record of a large-sized canid from this site, namely dentognathic remains, referable to a young adult individual that displays hypercarnivorous features (e.g., the reduction of the m1 metaconid and entoconid) that allow us to include these specimens in the hypodigm of the late Early Pleistocene species Canis (Xenocyon) lycaonoides. Much fossil evidence suggests that this species was a cooperative pack-hunter that, unlike other large-sized canids, was capable of social care toward kin and non-kin members of its group. This rather derived hypercarnivorous canid, which has an East Asian origin, shows one of its earliest records at Dmanisi in the Caucasus, at the gates of Europe. Interestingly, its dispersal from Asia to Europe and Africa followed a parallel route to that of hominins, but in the opposite direction. Hominins and hunting dogs, both recorded in Dmanisi at the beginning of their dispersal across the Old World, are the only two Early Pleistocene mammal species with proved altruistic behaviour towards their group members, an issue discussed over more than one century in evolutionary biology.

In the complex scenario of Plio–Pleistocene mammalian faunal turnovers, recent research on canids has revealed an increasingly higher number of species than previously thought. In this framework, Georgia had a key role in the biogeographic dispersion of fauna from/to Asia, Africa, and Europe. Historically attributed to Canis etruscus, the rich Canis material recovered from Dmanisi possesses certain peculiar cranial and dentognathic features, which cannot be regarded only as intraspecific variability. We revealed closer similarities between the Dmanisi wolf and the younger European Canis mosbachensis, rather than with other Early Pleistocene canids as C. etruscus and Canis arnensis. The discovery of a Canis borjgali sp. nov. in Dmanisi, with characteristics close to those of C. mosbachensis, changes radically the idea of Canis lupus evolution as it is conveyed today, invalidating the paradigm C. etruscus–C. mosbachensis–C. lupus lineage. Furthermore, the geographic position of Dmanisi in the Caucasian area offers interesting insights regarding the Asian canids and their dispersion into Europe and Africa, an aspect still poorly investigated. The exquisite state of preservation of the fossil from Dmanisi combined with novel 3D visualization and a digital imaging technique gives us the opportunity to increase the outreach of the research thanks to user-friendly and free tools. Here, for the first time, we employed augmented reality on a few specimens of C. borjgali sp. nov. through a simple web app. The extraordinary chance offered by these technologies has yet to be implemented in scientific research and dissemination, particularly in paleontology.