Carnivora from the Kanapoi hominin site, northern Kenya
Highlights
► Kanapoi is the richest early Pliocene carnivore assemblage in sub-Saharan Africa. ► The most abundant species is Parahyaena howelli, which likely denned in the area. ► Kanapoi is the only African locality with two species of the giant otter Enhydriodon. ► Kanapoi marks the first radiation of carnivorans in Africa after the Miocene.
Introduction
The Carnivora of Kanapoi were originally described by Werdelin (2003a) on the basis mainly of material collected by National Museums of Kenya expeditions in the 1990s, directed by Dr. Meave Leakey, with some additional specimens from the Harvard University expeditions in the 1960s directed by Dr. Bryan Patterson. Since the end of the 1990s, collecting at Kanapoi has continued on a smaller scale, led by one of us (FKM). This collecting has yielded additional carnivore material from the site, providing some significant additions to the fauna, including the first instance of two species of Enhydriodon in the same fauna. We have therefore prepared this contribution, to set these new finds in perspective and to take stock of the Kanapoi carnivoran sample as a whole.
The fossiliferous sediments at Kanapoi belong to the Kanapoi Formation, as defined by Feibel (2003). They consist of three sedimentary intervals: a lower fluviatile interval, a lacustrine interval, and an upper fluviatile interval. The stratigraphy and depositional setting of the Kanapoi sequence are discussed in detail by Feibel (2003). The Kanapoi sequence includes three dated tuffs (Leakey et al., 1998): a lower pumiceous tuff within the lower fluviatile interval, dated 4.17 ± 0.03 Ma, an upper pumiceous tuff just below the upper boundary of the lower fluviatile interval, dated 4.12 ± 0.02 Ma, and the Kanapoi Tuff at the top of the lacustrine interval, dated 4.07 ± 0.02 Ma. All of the carnivore specimens studied herein that have secured stratigraphic positions come from either the lower fluviatile interval or the lacustrine interval, with the exception of KNM-KP36600 (Homotherium sp.), which comes from the upper fluviatile interval.
Werdelin (2003a) noted that the majority of published eastern African early Pliocene (taken here as pre-Upper Laetolil beds, i.e., older than ca 3.85 Ma; Deino, 2011) mammal localities have few carnivore fossils associated with them. This is in marked contrast to the several rich Late Miocene localities such as Lothagam (Werdelin, 2003b), Lemudong’o (Howell and García, 2007) and the Lukeino Formation (Morales et al., 2005) in Kenya, and the Adu-Asa Formation in Ethiopia (Haile-Selassie and Howell, 2009). The situation has improved since 2003, however, with, e.g., the publication of carnivores from the informally named Mabaget Fm. overlying the Lukeino Fm. in Kenya (Morales et al., 2005). However, the assemblages from the early Pliocene still remain few and, more importantly, small. Therefore, any additions to these faunas are of great importance to the understanding of the first radiation of carnivores of modern aspect (i.e., post-Miocene) in Africa.
Section snippets
Material and methods
The following measurement abbreviations are used in this paper (upper dentition in upper case letters, lower dentition in lower case). LI1, WI1: length and width I1; LI3, WI3: length and width I3; LP4: buccal length P4; LliP4: lingual length P4; LpP4: length paracone P4; LmP4: length metastyle P4; WaP4 width of P4 including protocone; WblP4: width of metastyle P4; LM1: buccal length M1; LliM1: lingual length M1; WM1: greatest width M1; Lc: length lower canine; Wc: width lower canine; Lp2, Wp2:
Systematic paleontology
Order Carnivora
Family Mustelidae Fischer, 1817
Subfamily Lutrinae Bonaparte, 1838
Tribe Enhydriodontini Pickford, 2007
Genus Enhydriodon Falconer, 1868
Enhydriodon ekecaman Werdelin, 2003
Old material: KNM-KP 10034A-C, right C, P4 (Fig. 1A), M1 (Fig. 2A), and m1.
This material was fully described by Werdelin (2003a) and these descriptions will therefore not be reiterated here. Comparisons with new material of Enhydriodon from Kanapoi are given below.
Enhydriodon cf. Enhydriodon dikikae Geraads et al.,
Discussion
As noted, Kanapoi includes the most diverse carnivore assemblage from the eastern African early Pliocene (Table 1). Nevertheless, the assemblage is quite limited in species richness and any additions to the fauna are highly significant to understanding the carnivore guild of the early Pliocene. This time period (roughly 5.2–3.5 Ma) comes shortly after the late Miocene mass turnover of carnivores, which was a global occurrence, extending between ca 7 and 5 Ma (e.g. Werdelin and Turner, 1996). In
Conclusions
The carnivoran assemblage from Kanapoi now includes at least 12 species, nine of which have been identified to the generic level. This makes it the richest eastern African early Pliocene assemblage thus far described. Comparisons with other early Pliocene sites shows that the majority of taxa at Kanapoi have stratigraphic ranges that extend into the late Pliocene, but only a few back into the Miocene. This places Kanapoi as one of the earliest localities with a significant carnivoran assemblage
Acknowledgements
Permission to conduct field work at Kanapoi was granted by the Government of Kenya, and the National Museums of Kenya (NMK). The NMK and the Turkana Basin Institute (TBI) provided logistical support. The field work at Kanapoi would not have been possible without the involvement of a contingent of Kenyan field assistants, who comprised, R. Moru, J. Edung, J. Mbithi, P. Mulinge, W. Mang’ao, D. Muema, J. Ekeno, J. Nzavi, P. Were, S. Mwololo, J. Lotot, J. Eleman and D. Emuria. We also thank Z.
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