Abstract

We present an inventory of the progress of recent research on the biostratigraphy and palaeoecology of the genus Equus sensu lato in Europe. Our discussion starts with the new hypotheses concerning the dispersal and evolution of non-caballine equids of the Early and the beginning of the Middle Pleistocene, focusing on recent discoveries, description of taxa and revised diagnosis. In particular, we deal with the major debate surrounding the number of lineages, “stenonid horses”, related species and the new subgenus “Sussemionus”. Possible phylogenetic scenarios and relationships with extant species are also taken into consideration. There is consensus that the lineage of true horses emerged in Europe at the beginning of the Middle Pleistocene. Their extensive skeletal plasticity enabled them to survive under a wide range of climatic conditions and environments, from interglacial forests to grass-steppes during glacial episodes. They exhibit homogeneous overall morphology but variability in terms of dental and skeletal proportions. Patterns of general body-size, muzzle shape, macro- and microdonty, proportions of the limbs, robustness of the metapodials and breadth of the third phalanx are influenced by global climate, the characteristics of the vegetation (in particular the degree of openness) and substrate, and are governed by Bergman’s and Allen’s rules or other environmental pressures. In addition to palaeoecological information, these ecomorphological adaptations paradoxically provide real biochronological details in a given geographical province. Throughout the Upper Pleistocene, horses underwent a size diminution initiated at the end of the Middle Pleistocene. After the Last Glacial Maximum, investigation of body-size in different regions of Europe demonstrates the existence of latitudinal and longitudinal clines, and the fragmentation of the horse population. The so-called European wild ass, Equus hydruntinus, has a wide geographical distribution but seems to be less ecologically flexible, or, alternatively, more specialized than true horses, with which it is often found in sympatry. The latest palaeogenetic studies place the species into phylogenetic context within the Asiatic wild asses, however, palaeontological evidence points to Equus hydruntinus as a separate species. Its tooth morphology varies between biogeographical areas.

Highlights

  • It is generally considered that the genus Equus, which most likely emerged some 4.0–4.5 Ma (Orlando et al, 2013), includes all monodactyl equids present in Europe, and the stenonid group was the first to disperse at the Plio-Pleistocene boundary (Equus-Elephant event, Lindsay et al, 1980)

  • Equus livenzovenzis, recorded during the Middle Villafranchian, is for some authors the first species recognized in Eurasia and considered as the phylogenetic root of all the European stenonoid species (Azzaroli, 1992; Alberdi et al, 1998; Palombo and Alberdi, 2017; Bernor et al, 2018; Figure 1)

  • The first cranium of Equus described in Europe is that of Apollonia in Greek Macedonia (=Equus apolloniensis, Koufos et al, 1997; 1.2–0.9 Ma, Koufos and Kostopoulos, 2016) and the latest cranium of stenonids (Allohippus according to Eisenmann) could be that of Ceyssaguet with an age of 1.2 Ma (Equus stenonis, Aouadi and Bonifay, 2008)

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Summary

INTRODUCTION

Over the past 15 years, the discovery of new fossils and the revision of reference collections, added to the increased precision of the dating of certain deposits, have led to a reconsideration of the biochronology of the equids of the Pleistocene (Alberdi et al, 1998; Forsten, 1999a; Eisenmann, 2004a, 2006a, 2010; Alberdi and Palombo, 2013; Palombo and Alberdi, 2017; Van der Made et al, 2017). According to Eisenmann (2006a, 2010, 2017), similarities in the morphology of the teeth suggest, even in the absence of the cranium, a common origin within the branch of ≪ Equus ≫ for certain equids, at least from 1.5 Ma and maybe soon before around 2.5 Ma just above the Gauss-Matuyama limit (Vatera Formation, Greece). These equids (E. granatensis, E. altidens, E. hipparionoides, E. coliemensis, E. suessenbornensis) are grouped together in the subgenus of Equus “Sussemionus” (Eisenmann, 2010). The final part, about the European wild ass Equus hydruntinus, develops different aspects of its evolution

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REMARKS AND CONCLUSION
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