Journal of Paleontology, 92(2), 2018, p. 305–312 Copyright © 2017, The Paleontological Society. This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/ licenses/by/4.0/), which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited. 0022-3360/18/0088-0906 doi: 10.1017/jpa.2017.68


Gigantic lion, Panthera leo, from the Pleistocene of Natodomeri, eastern Africa


Fredrick K. Manthi,1 Francis H. Brown,2,† Michael J. Plavcan,3 and Lars Werdelin4*


1Department of Earth Sciences, National Museums of Kenya, Museum Hill, P.O. Box 40658-00100, Nairobi, Kenya ⟨fkmanthi@museums.or.ke⟩ 2Department of Geology and Geophysics, University of Utah, Salt Lake City, UT 84112-0102, USA ⟨frank.brown@utah.edu⟩ 3Department of Anthropology, University of Arkansas, Fayetteville, AR 72701, USA ⟨mplavcan@uark.edu⟩ 4Department of Palaeobiology, Swedish Museum of Natural History, Box 50007, S-104 05 Stockholm, Sweden ⟨lars.werdelin@nrm.se⟩


Abstract.—The partial skull of a lion from Natodomeri, northwest Kenya is described. The Natodomeri sites are correlated with Member I of the Kibish Formation, dated to between 195 ka and ca. 205 ka. The skull is remarkable for its very great size, equivalent to the largest cave lions (Panthera spelaea [Goldfuss, 1810]) of Pleistocene Eurasia and much larger than any previously known lion from Africa, living or fossil. We hypothesize that this individual represents a previously unknown population or subspecies of lion present in the late Middle and Late Pleistocene of eastern Africa rather than being an indication of climate-driven size increase in lions of that time. This raises questions regarding the extent of our understanding of the pattern and causes of lion evolution in the Late Pleistocene.


Introduction


The image of the lion is pervasive in human culture from its earliest preserved expression in Paleolithic art (Clottes, 2003) to more ‘modern’ cultural manifestations (e.g., Disney’s ‘The Lion King’). In the scientific world lions are also a focus for attention (Schaller, 1972; Packer, 2015), and although not at present seriously threatened, except in India, lions are nevertheless of some conservation concern due to population declines in recent decades (Bauer et al., 2015). Yamaguchi et al. (2004) reviewed the broad-scale evolution of lions and Barnett et al. (2014) and Bertola et al. (2016) have discussed their recent population history. These publications have shown that lions originated in Africa and, apart fromthe small relict population inwestern India, they are confined to that continent in the present day. In the past, however, lions (whether subspecies of Panthera leo [Linnaeus, 1758] or closely related but distinct species, is amatter of debate) were arguably the mammalian species that, except for humans and our commensals, had the widest geographic distribution, extending from South Africa, across Eurasia and all the way to the eastern seaboard of North America. Ironically, the Late Pleistocene history of lions is far better known from phylogeo- graphy than from fossils, while the fossil record of lions is better known in North America and Eurasia than in Africa. In Africa, the fossil record of lions comprises a few records


from eastern African late Pliocene and early Pleistocene sites and a small number of Middle and Late Pleistocene sites in North and South Africa (Hendey, 1974; Klein, 1986; Werdelin and Peigné, 2010; Geraads, 2012). Practically nothing is known of lion evolution in eastern Africa during the Middle and Late Pleistocene. Herein, we describe and discuss a partial lion skull


† Francis H. Brown is deceased. * Corresponding author.


from Natodomeri in northern Kenya, dated to ca. 196 ka BP. The specimen is interesting not only as the first Kenyan lion fossil found in this time, but also because of its very great size, with measureable skull parameters showing it to be among the largest lion skulls known (including North American and Eurasian fossil lions, which are known to have been on average considerably larger than the extant lion). Carnivore body mass (using lower carnassial length as proxy;


see Van Valkenburgh, 1990) in relation to latitude and climate has been studied by Klein (1986) and Klein and Scott (1989). These authors found that carnivore body mass generally follows ‘Bergmann’sRule,’ with larger individuals occurring at higher latitudes and during colder phases of the glacial cycles. Lionswere included in the latitudinal study and the results showed that the correlation between carnassial length and latitude (proxy for temperature) in lions was low but significant, and would have been higher if the sexes had been separated in the analysis.Alater study on size fluctuation during glacial-interglacial cycleswas carried out on spotted hyenas (Crocuta spp.), and lions may have conformed to the same pattern. With this background, finding the largest lion skull in Africa near the equator is unexpected. Natodomeri is located between settlements at the Kenyan


police posts of Kibish and Lokomarinyang. It lies in the Ilemi Triangle, a region that has been administered by Kenya since at least the 1950s. The name applies to a small ephemeral stream that cuts through a northeast-trending ridge at ~425m in elevation that is underlain by Pleistocene strata, and exposes these strata from ~35.65ºE to 35.80ºE (Fig. 1).


Geologic setting


The Natodomeri area was first noted as being fossiliferous by K.W. Butzer, who visited it during work of the International


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