Abstract

Rhinoceroses have been considered to have originated from tapiroids in the middle Eocene; however, the transition remains controversial, and the first unequivocal rhinocerotoids appeared about 4 Ma later than the earliest tapiroids of the Early Eocene. Here we describe 5 genera and 6 new species of rhinoceroses recently discovered from the early Eocene to the early middle Eocene deposits of the Erlian Basin of Inner Mongolia, China. These new materials represent the earliest members of rhinocerotoids, forstercooperiids, and/or hyrachyids, and bridge the evolutionary gap between the early Eocene ceratomorphs and middle Eocene rhinocerotoids. The phylogenetic analyses using parsimony and Bayesian inference methods support their affinities with rhinocerotoids, and also illuminate the phylogenetic relationships and biogeography of Ceratomorpha, although some discrepancies are present between the two criteria. The nearly contemporary occurrence of various rhinocerotoids indicates that the divergence of different rhinocerotoid groups occurred no later than the late early Eocene, which is soon after the split between the rhinocerotoids and the tapiroids in the early early Eocene. However, the Bayesian tip-dating estimate suggests that the divergence of different ceratomorph groups occurred in the middle Paleocene.

Highlights

  • Rhinoceroses have been considered to have originated from tapiroids in the middle Eocene; the transition remains controversial, and the first unequivocal rhinocerotoids appeared about 4 Ma later than the earliest tapiroids of the Early Eocene

  • The earliest hyracodontids and amynodontids are represented by Triplopus and Amynodon/Rostriamynodon, respectively, from the early Uintan North American Land Mammal Age (NALMA)[17] and/or Irdin Manhan Asian Land Mammal Age (ALMA)[18] (Fig. 1)

  • The phylogenetic analysis based on both parsimony and Bayesian inference criteria highlights the phylogeny and biogeography of Ceratomorpha, especially for some longstanding controversial groups, such as lophialetids, deperetellids, equivocal early rhinocerotoids, and relationships among rhinocerotoid groups

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Summary

Introduction

Rhinoceroses have been considered to have originated from tapiroids in the middle Eocene; the transition remains controversial, and the first unequivocal rhinocerotoids appeared about 4 Ma later than the earliest tapiroids of the Early Eocene. We describe 5 genera and 6 new species of rhinoceroses recently discovered from the early Eocene to the early middle Eocene deposits of the Erlian Basin of Inner Mongolia, China These new materials represent the earliest members of rhinocerotoids, forstercooperiids, and/or hyrachyids, and bridge the evolutionary gap between the early Eocene ceratomorphs and middle Eocene rhinocerotoids. Both morphological and molecular studies support the idea that Rhinocerotoidea and Tapiroidea form a monophyletic group Ceratomorpha[1,2,3]. Early–middle Eocene in either North America or Asia, some relatively small ceratomorphs have been argued to be rhinocerotoids, such as Fouchia, Dilophodon, Rhodopagus, and Yimengia[21,22,23] (Fig. 1c)

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