Perissodactyla (Rhinocerotidae and Equidae) from Kanapoi

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Abstract

The Kanapoi collection of Rhinocerotidae, first studied by Hooijer and Patterson (1972), now consists of 25 specimens and substantial reinterpretation of their affinities is made here. Kanapoi post-dates the extinction of Brachypotherium and the whole collection belongs to the Dicerotini. It is important because it includes the type-specimen of Diceros praecox, a species that remains poorly known, but looks slightly larger and more primitive than the modern ‘black’ rhino, Diceros bicornis. A second species is probably ancestral to the modern ‘white’ rhino, Ceratotherium simum; it looks identical to the Pleistocene North African Ceratotherium mauritanicum, of which Ceratotherium efficax is probably a synonym. The evolution of the Dicerotini in Africa can be regarded as an increasing divergence in diet and related morphofunctional adaptations in the two lineages. The co-occurrence at Kanapoi of both Diceros and Ceratotherium, with distinct dietary preferences, suggests some habitat heterogeneity, although the low sample size prevents robust paleoecological conclusions.

The Equidae are also rare and consist mostly of isolated teeth. I take the most parsimonious option of tentatively including all of them in a single species, whose identification is left open. Dental features of eastern African Pliocene to Pleistocene hipparions may reflect increasing adaptation to grazing.

Introduction

The Perissodactyla do not make up a high proportion of the Kanapoi large mammals, and no previous publication specifically deals with them, although Hooijer and Patterson (1972) and Hooijer (1975), respectively, studied the Rhinocerotidae and Equidae known at that time, together with fossils from other sites. Since then, renewed research at Kanapoi has significantly increased the faunal sample. The following revision is based upon the study of the material in the National Museums of Kenya (KNM) during several visits between 2002 and 2016, and takes into account all the material of Perissodactyla known to date. It includes no remains of the Chalicotheriidae, although this family was present in Africa from the Early Miocene until the Early Pleistocene with a sparse fossil record (Coombs and Cote, 2010.

Section snippets

Materials and methods

The comparisons of the Equidae are mostly based upon published literature, but the Rhinocerotidae have been extensively compared with the rich collections from other Pliocene and Pleistocene Kenyan sites in the KNM, as well as with the Hadar and Omo material in the National Museum of Ethiopia, Addis Ababa (NME). Late Miocene rhinos were principally examined in the Muséum National d'Histoire Naturelle, Paris (MNHN); Faculté des Sciences, Lyon (FSL); Naturhistorisches Museum, Basel (NHMB);

Systematic paleontology

  • Family Rhinocerotidae Gray, 1821.

  • Description: Hooijer and Patterson (1972) assigned all the Rhinocerotidae material known from Kanapoi at this time to a new species that they called Ceratotherium praecox, assuming that it was ancestral to the modern ‘white’ rhinoceros, Ceratotherium simum. No other study devoted to the Kanapoi rhinos has been published since then, although Harris et al., 2003, Geraads, 2005, Geraads, 2010, Giaourtsakis et al., 2009, and Hernesniemi et al. (2011) discussed them

Paleoecological interpretations

Paleoecological interpretations are hard to draw because of the small size of the collection. For instance, it would not be meaningful to calculate the relative abundance of the browser Diceros vs. the grazer Ceratotherium, but the presence of both genera and the sharp difference in their isotopic values attest to the presence of grassy plains but also of significant arboreal vegetation.

The Equidae are also rare, and the lack of metapodials or phalanges prevents the ecomorphological analyses

Conclusions

In spite of its small size, the Kanapoi rhino sample is important because it corresponds to a poorly documented time period in the evolution of the lineages leading to the modern forms. Still, only the discovery of well preserved, undistorted skulls could settle the disagreements regarding the evolution of the Rhinocerotidae in Africa, because their rather uniform cranial morphology and the scarcity of complete remains too often leads researchers to over emphasize dental features whose

Acknowledgments

I am grateful to C. Ward, F.K. Manthi, and M. Plavcan for having invited me to participate in this special issue about Kanapoi; to all people who allowed me to access modern and fossil collections in their care during the last decades: C. Argot and J. Lesur (MNHN), M. Bertling (GPMM), P. Brewer and A. Currant (NHMUK), G. Daxner-Höck (NHW), B. Engesser (NHMB), G. Senichaw (NME), B. Garevska (NHMMS), E. Heizmann (SMNS), W. Munk (SMNK), M. Muungu (KNM), A. Prieur (FSL); and to V. Eisenmann, L.

References (59)

  • R.L. Bernor et al.

    Hippotherium primigenium (Equidae, Mammalia) from the late Miocene of Höwenegg (Hegau, Germany), Thema: Fossilfundstätte Höwenegg

    Andrias

    (1997)
  • R.L. Bernor et al.

    A contribution to the evolutionary history of Ethiopian hipparionine horses (Mammalia, Equidae): morphometric evidence from the postcranial skeleton

    Geodiversitas

    (2005)
  • R.L. Bernor et al.

    35. Equidae

  • R.L. Bernor et al.

    Eurygnathohippus woldegabrieli sp. nov. (Perissodactyla, Mammalia) from the Middle Pliocene of Aramis, Ethiopia

    Journal of Vertebrate Paleontology

    (2013)
  • T.E. Cerling et al.

    Browsing and grazing in elephants: the isotope record of modern and fossil proboscideans

    Oecologia

    (1999)
  • T.E. Cerling et al.

    Dietary changes of large herbivores in the Turkana Basin, Kenya from 4 to 1 Ma

    Proceedings of the National Academy of Sciences of the United States of America

    (2015)
  • M.C. Coombs et al.

    33. Chalicotheriidae

  • W.O. Dietrich

    Zur Entwicklungsmechanik des Gebisses der afrikanischen Nashörner

    Zentralblatt für Mineralogie

    (1942)
  • V. Eisenmann

    Les Hipparions africains: valeur et signification de quelques caractères des jugales inférieures

    Bulletin du Muséum national d'histoire naturelle 3ème Sér. 438 Sci. Terre

    (1977)
  • V. Eisenmann

    Family Equidae

  • V. Eisenmann

    Les Equidés des gisements de la vallée de l'Omo en Ethiopie (collections françaises)

  • V. Eisenmann et al.

    The hipparion from the late Pliocene of Ahl al Oughlam, Morocco, and a revision of the relationships of Pliocene and Pleistocene African hipparions

    Palaeontologia Africana

    (2007)
  • A. Forsten

    Revision of the Palearctic Hipparion

    Acta Zoologica Fennica

    (1968)
  • D. Geraads

    Sur les relations phylétiques de Dicerorhinus primaevus Arambourg, 1959, Rhinocéros du Vallésien d'Algérie

    Comptes Rendus De l'Academie Des Sciences

    (1986)
  • D. Geraads

    Révision des Rhinocerotidae (Mammalia) du Turolien de Pikermi. Comparaison avec les formes voisines

    Annales de Paléontologie

    (1988)
  • D. Geraads

    Pliocene Rhinocerotidae (Mammalia) from Hadar and Dikika (Lower Awash, Ethiopia), and a revision of the origin of modern African rhinos

    Journal of Vertebrate Paleontology

    (2005)
  • D. Geraads

    34. Rhinocerotidae

  • D. Geraads et al.

    Upper Miocene Rhinocerotidae (Mammalia) from Pentalophos-1, Macedonia, Greece

    Palaeontographica

    (1990)
  • D. Geraads et al.

    Rhinocerotidae from the late Miocene of Bulgaria

    Palaeontographica A

    (2009)
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