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The Large-Mammal Fauna from the Kibish Formation Journal of Human Evolution

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The Large-Mammal Fauna from the Kibish Formation Journal of Human Evolution Journal of Human Evolution 55 (2008) 501–512 Contents lists available at ScienceDirect Journal of Human Evolution journal homepage: www.elsevier.com/locate/jhevol The large-mammal fauna from the Kibish Formation Zelalem Assefa a,*, Solomon Yirga b, Kaye E. Reed c a Archaeobiology and Human Origins Programs, Smithsonian Institution, National Museum of Natural History, 4210 Silver Hill Road, Suitland, MD 20746-2863, USA b Biology Department, Addis Ababa University, Science Faculty, P.O. Box 1176, Addis Ababa, Ethiopia c Institute of Human Origins, School of Human Evolution & Social Change, Arizona State University, P.O. Box 874101, Tempe, AZ 85287, USA article info abstract Article history: The Kibish faunal remains are useful for reconstructing the habitat of the earliest documented Homo Received 24 April 2007 sapiens and for understanding the community within which early modern humans existed. A diverse Accepted 15 May 2008 assemblage of large mammals, including many species of bovids, suids, and equids, has been recovered from the Kibish Formation. There are no extinct large mammals represented in the fossil assemblage, and Keywords: the overall taxonomic composition of the fossil fauna is similar to the modern-day wildlife community East Africa living near the Omo River. The fossil faunal assemblage shows a paucity of arboreal primates, and car- Ethiopia nivore species are rare. However, the faunal sample includes possible Cephalophus (duiker) remains and Extant fauna Hylochoerus meinertzhageni (giant forest hog), taxa that are extremely rare in the African fossil record, Hylochoerus Kibish and both indicate more closed habitats. Comparative analyses of the Kibish faunal remains using the Middle Stone Age ecological-diversity approach document close associations with edaphic grassland and woodland Paleoecology vegetation types. These vegetation forms are similar to current habitats surrounding the Omo River. Ó 2008 Elsevier Ltd. All rights reserved. Introduction In this paper, we discuss the vegetation and fauna that exist in the Omo Valley today. This is followed by a description of the fauna Recent expeditions to the Kibish Formation in the Omo River recovered from the members of the Kibish Formation. We then Valley of southern Ethiopia (1999–2003) have yielded a sample of reconstruct the paleoenvironment of the Kibish Formation based mammal remains that comes from all but one of the four members on these faunal remains and discuss the mammalian communities of the formation. Mammalian fauna is useful for reconstructing within which Homo sapiens fossils were recovered. vegetation structure and for understanding the ecological associ- ations of homininsdin this case, early Homo sapiensdwith their environment, including the community in which they lived. The Background mammalian fauna from Kibish represents a poorly known but critical phase in the east African fossil record. According to the Vegetation newly revised dates (McDougall et al., 2005, 2008), the three members of the Kibish Formation represented by mammalian Today the Omo Kibish area supports both wet and dry vegeta- fossils sample the interglacial phases of three different isotopic tion types following the course of the meandering Omo River. stages. Member I accords with isotopic stage 7 (240–200 ka), Closed to open woodlands dominate the inner bends of the river. Member III with isotopic stage 5 (130–75 ka), and Member IV with The floodplains in the extended area away from the riparian the early stages of the Holocene (<10 ka). Specifically, each of the woodland support open, dry habitats such as shrubland, grassland, members results from periodic episodes of high rainfall in the Ethi- and bushland thickets. At a larger scale, the lower Omo regional opian plateau that resulted in northward expansion of Lake Turkana basin has an arid to semiarid tropical climate, characterized by high (Butzer, 1971; Brown and Fuller, 2008; McDougall et al., 2008). The temperatures (Butzer, 1971) and low precipitation, with mean consistencies of the fauna among these three members reflect similar annual rainfall ranging from 320 to 380 mm and a maximum mean habitats. The depositional environments characterize the wet phases temperature of 40 C(Carr, 1976), based on records of nearby sta- of the interglacials, as they are deltaic and littoral deposits. tions such as Ileret, Todenyang, and Lokomarinyang (Butzer, 1971). Although all fossil-bearing localities from the Kibish Formation come from deltaic and littoral beds on both sides of the Omo River, major sedimentary environments of the Omo Kibish area today also * Corresponding author. include crystalline uplands, pediments, and piedmont alluvia. The E-mail address: [email protected] (Z. Assefa). interaction of various soil types, topographic changes, and 0047-2484/$ – see front matter Ó 2008 Elsevier Ltd. All rights reserved. doi:10.1016/j.jhevol.2008.05.015 502 Z. Assefa et al. / Journal of Human Evolution 55 (2008) 501–512 availability of moisture generally provide a mosaic of plant com- variable grazers, browser-grazer intermediates, generalists, and munities in the region, which were assigned by Carr (1976) to the browsers. Thus, in its modern ecological setup, the region supports broad habitat types of plains environments (grasslands), riverine- diverse species preferring both open and closed habitats (Gagnon associated environments (riparian forest and woodland), and and Chew, 2000; Cerling et al., 2003). However, grazers of both anomalous habitats (e.g., sporadic uplands and outcrops, streams, obligate and variable types appear to have higher representation hot springs). Figure 1 shows a schematic profile of vegetation types among the extant bovids of the Omo Kibish area (Yalden et al., across grasslands and riparian woodland. 1984). The majority of grazing species are found mainly within the Both grasslands and riverine-associated vegetation structures different areas of grassland habitats. In the extant Omo region, are dependent on soil types, rainfall, and water-drainage patterns. there are no documented forest frugivorous/browsing bovids (i.e., Higher elevations support higher plant diversity, while in the cephalophines), probably indicating that the modern riverine forest mudflats and floodplains, trees are essentially nonexistent (Carr, is lacking in the type of fruiting plant species required to support 1976, 1998). There are diverse mesic depositional environments, these bovids. such as deltas, silt berms, levees, back swamps, and adjacent flats The Omo Kibish area is known for one of the few sympatric that also support high floral diversity. Vegetation types in the distributions of Burchell’s (Equus burchellii) and Grevy’s zebras modern delta and silt berms share many similarities. The riverine (Equus grevyi). Their joint presence signifies diverse precipitation woodlands occupy the natural levee crest of the river and the and temperature ranges and a wider range of accumulated vege- immediate backslope. Carr (1976, 1998) noted progressive shifts in tation indices, distinct from the predominantly milder or harsher the following sequences along south-to-north meanders of the climates in the nonoverlapping ranges of these two equid species river: grassland and thicket, open woodland, closed woodland, and elsewhere (Bauer et al., 1994). The representation of two species of forest. Such sequences particularly characterize levee vegetation suids, the common warthog (Phacochoerus aethiopicus) and the types along inner bends of the lower 50–60 km of the river. Farther bushpig (Potamochoerus porcus), is also consistent with the mosaic upstream, including the area where the Kibish Formation is environments of Omo Kibish. The Omo Valley also supported the exposed, the forest-cover declines and is replaced by intermediate last known survivors in Ethiopia of the once widespread black vegetation between woodland and forest. The riverine woodlands rhinoceros (Diceros bicornis), which preferentially inhabits include large trees, such as fig (Ficus sycomorus), and two major ecotones between forests and grasslands. Of the diverse species of species of shrubsdgundi (Cordia sinensis) and confetti tree (May- primates documented in the Lower Omo Valley, the forest and tenus senegalensis), which are often found in association with dense woodland support guereza (Colobus guereza), blue monkey (Cer- thickets. The riverine forest, on the other hand, is composed of copithecus mitis), and De Brazza’s monkey (Cercopithecus neglectus), woody plants that grow to a height of 25–30 m, and also maintain whereas other transitional zones, as well as plains and adjacent dense canopy covers. Herbs and shrubs are also found associated flats, support Senegal galagos (Galago senegalensis), vervet mon- with the riverine forest. For a list of plant species, see Carr (1976, keys (Chlorocebus aethiops), and anubis baboons (Papio anubis). The 1998). number of primate species inhabiting the forest today indicates the richness and lateral extent of the forest along the Omo River. Most Fauna riparian forests in semiarid regions only have Chlorocebus aethiops and occasionally Cercopithecus mitis or Colobus guereza. The extant and historically documented faunal composition of the Omo Kibish area is characterized by diverse taxa of large Materials and methods mammals (Table 1). Following Gagnon and Chew’s (2000) dietary- preference assessment of extant African bovids, the species of In the four field seasons conducted between the years 1999 and bovids reported from the
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