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Rhinocerotidae Chapter 11 Rhinocerotidae Elina Hernesniemi, Ioannis X. Giaourtsakis, Alistair R. Evans, and Mikael Fortelius Abstract The remains of fossil rhinoceroses from Laetoli Introduction represent at least three taxa: Ceratotherium efficax, Ceratotherium cf. simum, and Diceros sp. The great majority Since the first systematic description by the German paleon- of the material from the Pliocene Laetolil Beds belongs to C. tologist W. O. Dietrich during the Second World War efficax, for which we provide a revised diagnosis. This taxon (Dietrich 1942, 1945), fossil remains of rhinoceroses from has been frequently misidentified and inaccurately referred to Laetoli have been described and discussed under a variety of as C. praecox, C. germanoafricanum, or C. mauritanicum. A names (Arambourg 1947, 1959; Hooijer 1969, 1972; Groves cranium from the Upper Ndolanya Beds shows more derived 1975; Guérin 1979, 1980a, 1987a; Geraads 2005). The dental features, but a precise assignment to C. germanoafri­ Laetoli rhinoceroses are of particular interest for under- canum or C. simum is presently not possible. The occurrence standing the early evolution of the Ceratotherium lineage of true Diceros in the Laetolil Beds is demonstrated by a par- during the Plio-Pleistocene, as well as for interpreting the tial cranium with incomplete dentition, but very few other paleoecological setting of the locality. In particular, the tran- specimens can be potentially attributed to this genus. Analysis sition from a generalized ancestral morphology to the speciali- of occlusal wear patterns suggests that C. efficax was a grazer zed grazing condition of the extant white rhinoceros, or possibly a graze-dominated mixed feeder; in either case it Ceratotherium simum, has been long regarded as a case study probably included a variable component of browse in its diet. for evolutionary research (Osborn 1903; Dietrich 1942; The transition from a Diceros-like ectolophodont dentition Thenius 1969; Stanley 1979). to the full-fledged plagiolophodonty seen in extant Cera­ Our aim is simple: to revise the taxonomy of the rhinoceros totherium simum included a substantial period of stasis, material from Laetoli housed in the collections of the spanning at least the interval represented by the Laetolil Beds. National Museums of Tanzania and Kenya, and in the A shift in the dietary regime towards increased grazing had collections of the Natural History Museum, London. occurred by the Upper Ndolanya time, and this trend continued Furthermore, we describe it with emphasis on functional from the early Pleistocene to the Recent. Based on the available aspects, particularly the dental functional morphology and fossil record, the split of the two lineages leading to the extant wear patterns. We consider the implications of our findings species must have taken place in Africa during the Miocene. for the paleoecology of the Laetoli sequence and the evolu- tionary history of the taxa, including the origins of Keywords Rhinocerotidae • Ceratotherium • Diceros Ceratotherium simum. We also discuss in detail the compli- • Taxonomy • Evolution • Nomenclature • Paleoecology cated taxonomic history of C. efficax (Dietrich, 1942), and • Paleodiet Ceratotherium praecox Hooijer and Patterson, 1972. E. Hernesniemi (*) and M. Fortelius Department of Geosciences and Geography, University of Helsinki, P.O. Box 64 FIN-00014, Helsinki, Finland e-mail: [email protected]; [email protected] Materials and Methods I.X. Giaourtsakis Department of Geo-and Environmental Sciences, Section of Paleontology, Ludwig-Maximilians-University of Munich, Institutional Abbreviations Richard-Wagner-Strasse 10, D-80333 Munich, Germany e-mail: [email protected] AMPG: Athens Museum of Paleontology and Geology, A.R. Evans University of Athens; BMNH: British Museum of Natural School of Biological Sciences, Monash University, Victoria 3800, Australia History (=Natural History Museum), London; BSPG: e-mail: [email protected] Bayerische Staatssammlung für Paläontologie und Geologie, T. Harrison (ed.), Paleontology and Geology of Laetoli: Human Evolution in Context. Volume 2: Fossil Hominins 275 and the Associated Fauna, Vertebrate Paleobiology and Paleoanthropology, DOI 10.1007/978-90-481-9962-4_11, © Springer Science+Business Media B.V. 2011 276 E. Hernesniemi et al. München; FSL: Faculté des Sciences, University of Lyon; Heissig 1975, 1996; Geraads 1994; Fortelius et al. 2003) GSN: Geological Survey of Namibia, Windhoek; IPUW: were studied at BSPG, SMNK, and MNHN; and those from Institut für Paläontologie der Universität, Wien; KMMA: Iran (Maragheh: Osborn 1900; Thenius 1955) were exam- Koninklijk Museum voor Midden-Afrika, Tervuren; KNM: ined at NHMW and MNHN. Casts of specimens from Fort National Museums of Kenya, Nairobi; LGPUT: Laboratory Ternan (Hooijer 1968) and Langebaanweg (Hooijer 1972) of Geology and Palaeontology, University of Thessaloniki; were examined at BMNH and BSPG, respectively. Cranial MNHB: Museum der Naturkunde für Humboldt Universität material of Diceros douariensis from Douaria (Guérin 1966) zu Berlin; MNHN: Muséum National d’Histoire Naturelle, was kindly shown to MF by C. Guérin and digital images of Paris; NHMW: Naturhistorisches Museum, Wien; NME: the specimens were provided by C. Guérin and A. Prieur. National Museum of Ethiopia, Addis Ababa; NMT: National The rhinoceros material from the following localities was Museum of Tanzania, Dar es Salaam; RMNH: Rijkmuseum studied based on the referred publications: Arrisdrift, van Natuurlijke Historie (Naturalis), Leiden; SAM: South Namibia (Guérin 2000, 2003); Ekora Formation, Kenya African Museum, Cape Town; SMF: Forschungsinstitut und (Hooijer and Patterson 1972); Lothagam, Kenya (Hooijer Naturmuseum Senckenberg, Frankfurt am Main; SMNK: and Patterson 1972; Harris and Leakey 2003); Ahl al Staatliches Museum für Naturkunde, Karlsruhe; SMNS: Oughlam and Oulad Hamida, Morocco (Geraads 2005). Staatliches Museum für Naturkunde, Stuttgart; ZMA: Comparative studies with the extant species Ceratotherium Zoological Museum, Amsterdam. simum and Diceros bicornis have been also carried out at the zoological collections of the aforementioned institutions. Material Stratigraphy The material of fossil rhinoceroses from Laetoli is housed at the National Museum of Tanzania in Dar es Salaam, the For temporal resolution of the Laetoli Sequence, we used the National Museums of Kenya in Nairobi, the Museum der stratigraphic position of the localities relative to the marker Naturkunde für Humboldt Universität in Berlin, and the tuffs to create semi-arbitrary Tuff Groups (Harrison and Kweka Natural History Museum of London. Detailed information 2011). The following groups were created: 1-LLB = Lower about the excavation and research history of the locality, as Laetolil Beds, 2-BT3 = below Tuff 3, 3-T3-5 = between Tuffs well as of the geological and stratigraphical setting is pro- 3 and 5, 4-T5-7 = between Tuffs 5 and 7, 5-STRT7 = straddles vided by Harrison (2011), Ditchfield and Harrison (2011), Tuff 7, 6-AT7 = above Tuff 7, 7-UND = Upper Ndolanya Beds. and Harrison and Kweka (2011). The Laetoli material was compared with selected fossils from the Aterir Beds (Hooijer 1973), Chemeron Formation (Hooijer 1969), Kanapoi (Hooijer and Patterson 1972), and Mesowear Koobi Fora and West Turkana (Harris 1976, 1983) at the col- lections of KNM. Additional comparative studies with mate- We applied the original mesowear scoring system and ana- rial from the Plio-Pleistocene localities of Hadar (Guérin lytical techniques introduced by Fortelius and Solounias 1980b; Geraads 2005) and Dikika (Geraads 2005) were car- (2000) for the upper dentition. We also extended the method- ried out at the NME; from the Omo Valley (Arambourg 1947; ology to the cusp sharpness of the buccal enamel band of the Hooijer 1969, 1972, 1973, 1975; Guérin 1985) at NME, lower teeth. We did this by a subjective judgment of equiva- RMNH and MNHN. Pleistocene material from Olduvai lences according to the following guideline: distinct phase I Gorge, Kanjera, Kanam West, and Rawi (Hooijer 1969; facets with sharp boundaries = sharp; distinct phase I facets Groves 1975) was studied at BMNH. The Kohl-Larsen fossil with fuzzy boundaries = rounded; no phase I facets = blunt. collection from East Africa (Dietrich 1942, 1945) was stud- Relief was not scored for lower teeth and, apart from hierar- ied at MNHB. Specimens from Bou Hanifia (Arambourg chic clustering, we have limited our comparisons to cusp 1959; Geraads 1986), Ternifine (Pomel 1895) and several sharpness only. All teeth that were sufficiently well preserved Plio-Pleistocene North African localities (Arambourg 1970) were scored for mesowear, but only the first and second were examined at MNHN. Materials of “Diceros” neumayri molars were included in the final analyses, and only the from Greece (Pikermi, Samos, Axios Valley: Gaudry 1862– uppers in the hierarchic clustering analysis. Results gener- 1867; Geraads 1988; Geraads and Koufos 1990; Giaourtsakis ally remain similar even if premolars and deciduous teeth are et al. 2006; Giaourtsakis 2009) were studied at AMPG, included. We used polysiloxane putty (Provil Novo Putty LGPUT, MNHN, BMNH, NHMW, IPUW, SMNS, BSPG, regular set, Heraeus Kulzer GmbH, Hanau, Germany) to SMF and HLMD; those from Turkey (Various localities: make partial molds of the teeth and synthetic dental stone 11 Rhinocerotidae 277 (Fujirock by GC Europe n.v., Leuwen, Belgium) to make 1942. Serengeticeros efficax
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