Article

Taxonomic status and paleoecology of Rusingoryx atopocranion (Mammalia, Artiodactyla), an extinct Pleistocene bovid from Rusinga Island, Kenya

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Abstract

Rusingoryx atopocranion is a poorly known extinct alcelaphine bovid, documented in Pleistocene deposits associated with Middle Stone Age artifacts on Rusinga Island, Kenya. Following its initial description, Rusingoryx was subsumed into Megalotragus, which includes the extinct giant wildebeests, on the basis of its cranial architecture. Renewed investigations of the Pleistocene deposits on Rusinga Island recovered a large sample of Rusingoryx specimens that provide new taxonomic and paleoecological insight. This study (1) reviews the morphological and phylogenetic evidence concerning the taxonomic status of Rusingoryx and (2) evaluates its paleoecology and dietary habits. The morphology and phylogenetic data indicate that Rusingoryx is distinct from Megalotragus; they likely shared a common ancestor in the late Pliocene. Ecomorphology and mesowear analysis point to a specialized grazing adaptation, and its association with arid-adapted ungulates suggests a preference for arid grasslands. The confirmation of Rusingoryx as a valid taxonomic entity, together with the presence of other extinct taxa (including Megalotragus) on Rusinga Island, suggests an increasingly complex pattern of ungulate biogeography and extinctions in the late Quaternary of East Africa. Rusingoryx appears to have been part of an arid-adapted faunal community that potentially persisted in East Africa until the onset of the Holocene.

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... It is a commonly held view that all very large-bodied alcelaphine antelope in Africa belong to one genus, Megalotragus Van Hoepen, 1932, and that the various fossil species of this genus are closely related (Gentry, 1978(Gentry, , 2010Gentry and Gentry, 1978;Gentry et al., 1995;Vrba, 1979Vrba, , 1997. Prev iously a smaller-bod ied species fro m Rusinga Island, Kenya, Rusingoryx atopocranion Pickford and Thomas, 1984, was referred to the genus Megalotragus (Vrba, 1997), but more recently in a revision of the mo rpho logical and phylogenetic relat ionships of this species it was re-assigned to Rusingoryx as a genus distinct from Megalotragus, although closely related (Pickford and Thomas, 1984;Faith et al., 2011). Previously it was suggested that an East African species, M. kattwinkeli, is ancestral to the southern African M. priscus, wh ich is considered to include two temporal fo rms, M. priscus eucornutus and M. priscus priscus . ...
... However, this hypothesis seems no longer to be suppo rted (Gentry, 2010). There is no fossil evidence yet for the species M. eucornutus and M. priscus outside of southern Africa, although Faith et al. (2011) mentio ns the presence of large wildebeest-like alcelaphine fro m the Wasiriya Beds on Rus inga Island, which they refer to the genus Megalotragus. M. eucornutus is known only fro m Co rnelia-Uit zoek, the type locality of the Cornelian Land Mammal Age (LMA) and from Co rnelia-Mara, a nearby locality of similar age (Brink et al., 2012;Brink, in press). ...
... These features are modifications to allow the mandib le to fit the ext remely elongated skull. A similar, but less extre me widening of the angle between the ramus and the corpus is seen it the mandible of Rusingoryx atopocranion (Faith et al., 2011). Another parallel to this extre me morph ology of the lower jaw can be found in the hartebeest and in the Barbary sheep, Ammotragus lervia. ...
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A reconstruction of the skull of the giant alcelaphine bovid, Megalotragus priscus, is provided based on a brain case and horn cores discovered and excavated at the late Florisian locality of Erfkroon on the Modder River, central Free State Province, South Africa. The sedimentary context of the M. priscus specimen can be correlated with fluvial deposits dated previously by luminescence to the Last Interglacial. Electron Spin Resonance (ESR) analyses of dental specimens from various localities at Erfkroon indicate a terminal Middle Pleistocene and Late Pleistocene age for these deposits. The skull reconstruction of M. priscus is aided by an upper jaw and mandible from the Late Pleistocene locality of Mahemspan. The M. priscus materials from Erfkroon, Mahemspan and other localities allow a re-evaluation of the morphological affinities of the species and it appears to be closer to wildebeest-like alcelaphines (genus Connochaetes) than to hartebeest-like alcelaphines (genera Alcelaphus and Damaliscus). Variability in the fossil horn cores suggests sexual dimorphism and some degree of territorial behaviour. It also suggests geographic variability in the populations of M. priscus in central southern Africa during the later part of the Middle Pleistocene and Late Pleistocene, before its extinction at the end of the Late Pleistocene and early Holocene.
... The fossils discussed here were recovered from Rusinga Island and Karungu in the Lake Victoria Basin (Fig. 2). Rusinga Island is located within Lake Victoria and, although now (Kent 1942;MacInnes 1956;Van Couvering 1972;Leakey 1974;Pickford 1984Pickford , 1986 and are now the subject of renewed investigation focusing on the environmental and ecological context of the Middle Stone Age (MSA) archaeological sites (Tryon et al. 2010Faith et al. 2011). The poorly consolidated Pleistocene deposits on Rusinga Island, known as the Wasiriya Beds, are characterized by weakly-developed paleosols and tuffaceous fluvial sediments recording a complex cut-and-fill system. ...
... The minimum age is provided by a suite of calibrated radiocarbon age estimates on the shells of gastropods that most likely burrowed into the sediments at some point after deposition (Tryon et al. 2010. The fossil mammals from Rusinga Island suggest an expansion of grasslands distinct from the bushland, thicket, and forest found in the region today (Faith et al. 2011Tryon et al. 2012). Open grassland species such as alcelaphine antelopes (wildebeest and allies) are dominant and several extinct specialized grazers are present, including Rusingoryx atopocranion , Damaliscus hypsodon, Megalotragus , and Syncerus antiquus. ...
... The combination of diminished rainfall together with the competitive advantage of C 4 vegetation at lower atmospheric CO 2 concentrations probably accounts for the grassy paleoenvironment implied by the fauna, consistent with paleo-vegetation models for Pleistocene glacial phases (e.g., Cowling et al. 2008;Prentice et al. 2011). In light of lake level fluctuations observed historically (Nicholson 1998) and documented in the late Pleistocene and Holocene geological record (Johnson et al. 1996;Stager et al. 2002Stager et al. , 2011Stager and Johnson 2008), the presence of large gregarious grazers and aridadapted ungulates from Rusinga Island suggests a connection to the mainland (Faith et al. 2011). This is further supported by the presence of a similar fauna from roughly contemporaneous deposits on nearby Mfangano Island, which imply a >25 m decline in lake levels (Tryon et al. in press). ...
Article
This study contributes to the growing complexity of the impala fossil record through a morphological description and analysis of Aepyceros fossils from late Pleistocene deposits in Kenya’s Lake Victoria Basin. We show that the Lake Victoria impala belongs to an extinct species that differs from modern impala and its fossil predecessors by a combination of exceptionally deep mandibles and teeth characterized by greater hypsodonty and occlusal lengths. Whereas modern impala (A. melampus) displays substantial ecological flexibility, these traits in the extinct species suggest a more dedicated adaptation to grazing in open and dry environments. Previous phylogeographic observations indicate that A. melampus was extirpated from East Africa, perhaps during the middle-to-late Pleistocene, and later recolonized from southern Africa. The Lake Victoria impala raises the possibility that the evidence interpreted as extirpation may instead reflect speciation, with A. melampus giving rise to a novel East African species while persisting unchanged in southern Africa. Increased rainfall and rising atmospheric CO2 concentrations at the end of the Pleistocene may have played a role in the disappearance of the extinct form via habitat loss and possibly competition with the more versatile A. melampus.
... The Late Pleistocene geology, fossils and MSA artefacts from Rusinga and Mfangano Islands have been the focus of research since 2009 (Tryon et al., 2010Faith et al., 2011Faith et al., , 2012Faith et al., , 2014Van Plantinga, 2011). The Pleistocene deposits on Rusinga Island, informally designated the 'Wasiriya Beds' by Pickford (1984), unconformably overlie a complex Miocene palaeotopography and are predominately comprised of tuffaceous alluvial and fluvial sediments intercalated with palaeosols that formed during periods of landscape stability, variably reworked tephra and rare tufa deposits (Tryon et al., 2010Van Plantinga, 2011). ...
... Previous research has shown that the Pleistocene deposits surrounding Lake Victoria yield an abundance of well-preserved fossils and MSA ar-tefacts (e.g. Owen, 1937;Kent, 1944;Pickford, 1984;Behrensmeyer et al., 1995;Ditchfield et al., 1999;Plummer et al., 1999;Tryon et al., 2010Tryon et al., , 2012Tryon et al., , 2014Faith et al., 2011Faith et al., , 2012Faith et al., , 2013Faith et al., , 2014Van Plantinga, 2011;Garrett et al., in press). Stone artefacts from Rusinga Island and Karungu include flakes, blades, retouched points and Levallois cores, consistent with a MSA attribution Faith et al., in press b), the industry associated with the earliest modern humans. ...
... The fauna are dominated by alcelaphine bovids (wildebeest and allies) and equids, suggesting environments that were grassier and probably drier than the evergreen bushland, thicket and woodland found in the region today. Oryx (Oryx beisa) and Grevy's zebra (Equus grevyi), which prefer arid to semiarid grasslands and shrublands, and extinct antelopes adapted to grazing in dry grasslands, indicate that the environment was significantly more arid than at present (Tryon et al., 2010Faith et al., 2011Faith et al., , 2012Faith et al., , 2013Faith et al., , 2014. Carbon isotopes of mammalian tooth enamel indicate a diet of predominately C 4 grasses (Garrett et al., in press; Faith et al., in press b). ...
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The effect of changing palaeoclimate and palaeoenvironment on human evolution during the Pleistocene is debated, but hampered by few East African records directly associated with archaeological sites prior to the Last Glacial Maximum. Middle to Late Pleistocene deposits on the shoreline of eastern Lake Victoria preserve abundant vertebrate fossils and Middle Stone Age arte-facts associated with riverine tufas at the base of the deposits, which are ideal for palaeoenvironmental reconstructions. New data from tufas identified on Rusinga Island and on the mainland near Karungu, Kenya are provided from outcrop, thin sections, mineralogical, stable isotopic and U-series dating analyses. Tufa is identified in four sites: Nyamita (94·0 ± 3·3 and 111·4 ± 4·2 ka); Kisaaka, Aringo (455 ± 45 ka); and Obware. The age ranges of these tufa deposits demonstrate that spring-fed rivers were a recurrent, variably preserved feature on the Pleistocene landscape for ca 360 kyr. Poor sorting of clastic facies from all sites indicates flashy, ephemeral discharge, but these facies are commonly associated with barrage tufas, paludal environments with δ13C values of ca 10‰ indicative of C3 plants and fossil Hippopotamus, all of which indicate a perennial water source. Other tufa deposits from Nyamita, Obware and Aringo have a mixed C3/C4 signature consistent with a semi-arid C4 grassland surrounding these spring-fed rivers. The δ18O values of tufa from Nyamita are on average ca 1‰ more negative than calcite precipitated from modern rainfall in the region, suggesting greater contribution of depleted monsoonal input, similar to the Last Glacial Maximum. Microdebitage and surface-collected artefacts indicate that early modern humans were utilizing these spring-fed rivers. The presence of spring−fed rivers would have afforded animals a reliable water source, sustaining a diverse plant and animal community in an otherwise arid environment.
... The sediments at Karungu preserve abundant vertebrate fossils and Middle Stone Age (MSA) artifacts (Owen, 1937;Pickford, 1984;, which are considered the archeological signature of early H. sapiens in East Africa (McBrearty and Brooks, 2000;Tryon and Faith, 2013). The pre-LGM Karungu dataset complements, refines, and expands those from correlative deposits on Rusinga and Mfangano Islands~40 km to the north (Tryon et al., 2010;Faith et al., 2011;Van Plantinga, 2011;Faith et al., 2012;Tryon et al., 2012;Faith et al., 2014;Tryon et al., 2014;Garrett et al., 2015). Previous evidence from MSA archeological and paleontological sites from Rusinga and Mfangano Islands suggests that the contraction of Lake Victoria and expansion of grasslands during the late Pleistocene may have facilitated the dispersal of large-bodied mammals, including humans, across Africa (e.g., Faith Blegen et al. (2015). ...
... Karungu (0.84°S, 34.15°E) is located on the Kenyan margin of Lake Victoria (Fig. 1),~40 km south of Pleistocene localities on Rusinga and Mfangano Islands that have been the focus of research by our team since 2009 (Tryon et al., 2010;Faith et al., 2011;Van Plantinga, 2011;Faith et al., 2012;Tryon et al., 2012;Faith et al., 2014;Tryon et al., 2014;Beverly et al., 2015;Garrett et al., 2015). The Pleistocene deposits at Karungu are exposed at seven sites around the town of Sori. ...
... The presence of gregarious and migratory grazers on Mfangano Island, which is too small to support viable populations of large ungulates, suggests a connection to the mainland. This requires a lake-level decline of at least 25 m (Tryon et al., 2010Faith et al., 2011Faith et al., , 2012Faith et al., , 2014 and comparisons with existing models suggest that such a decline is only possible with a significant rainfall reduction (Broecker et al., 1998;Milly, 1999). Analyses by Faith (2013) indicate peak ungulate diversity in sub-Saharan African game reserves at~800 mm yr −1 and evidence from the paleosols provide quantitative support to explain this high diversity of ungulates. ...
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The effect of changing environment on the evolution of Homo sapiens is heavily debated, but few data are available from equatorial Africa prior to the last glacial maximum. The Karungu deposits on the northeast coast of Lake Victoria are ideal for paleoenvironmental reconstructions and are best studied at the Kisaaka site near Karunga in Kenya (94 to N33 ka) where paleosols, fluvial deposits, tufa, and volcaniclastic deposits (tuffs) are exposed over a ~2 km transect. Three well-exposed and laterally continuous paleosols with intercalated tuffs allow for reconstruction of a succession of paleocatenas. The oldest paleosol is a smectitic paleo-Vertisol with saline and sodic properties. Higher in the section, the paleosols are tuffaceous paleo-Inceptisols with Alfisol-like soil characteristics (illuviated clay). Mean annual precipitation (MAP) proxies indicate little change through time, with an average of 764 ± 108 mm yr −1 for Vertisols (CALMAG) and 813 ± 182 to 963 ± 182 mm yr −1 for all paleosols (CIA-K). Field observations and MAP proxies suggest that Karungu was significantly drier than today, consistent with the associated faunal assemblage, and likely resulted in a significantly smaller Lake Victoria during the late Pleistocene. Rainfall reduction and associated grassland expansion may have facilitated human and faunal dispersals across equatorial East Africa.
... Due to 3 cm of upwardly angled nasal bones, it was initially postulated that Rusingoryx may have had a proboscis or an enlarged, domed nasal region [1]. Due in part to the incompleteness of the type specimen, this reconstruction was dismissed and the taxonomic validity of Rusingoryx was questioned [2] until a more recent reexamination and expanded sample of horn cores and dental material [3]. Here we present six new Rusingoryx cranial specimens excavated en masse from a channel deposit within the type locality [4]. ...
... KNM-RU-52572 is post-depositionally compressed in the sagittal plane. The caudal portion of the cranium matches the holotype, and the dentition follows recent descriptions of fragmentary material [3]. In mature individuals, the frontal, nasal, and premaxillary elements form a prominent, osseous nasal dome. ...
... Nasal vocalizations of Rusingoryx may similarly fall under this evolutionary pretext. Paleoenvironmental evidence from the Wasiriya Beds [3,11,28,29] indicates widespread semi-arid grasslands, and a markedly high degree of hypsodonty in Rusingoryx molars implies a preference for such habitats [3]. Dry environments are ideal for low-frequency vocalizations, as these sound waves can travel over 10 km in such environments [26]. ...
Article
The fossil record provides tangible, historical evidence for the mode and operation of evolution across deep time. Striking patterns of convergence are some of the strongest examples of these operations, whereby, over time, similar environmental and/or behavioral pressures precipitate similarity in form and function between disparately related taxa. Here we present fossil evidence for an unexpected convergence between gregarious plant-eating mammals and dinosaurs. Recent excavations of Late Pleistocene deposits on Rusinga Island, Kenya, have uncovered a catastrophic assemblage of the wildebeest-like bovid Rusingoryx atopocranion. Previously known from fragmentary material, these new specimens reveal large, hollow, osseous nasal crests: a craniofacial novelty for mammals that is remarkably comparable to the nasal crests of lambeosaurine hadrosaur dinosaurs. Using adult and juvenile material from this assemblage, as well as computed tomographic imaging, we investigate this convergence from morphological, developmental, functional, and paleoenvironmental perspectives. Our detailed analyses reveal broad parallels between R. atopocranion and basal Lambeosaurinae, suggesting that osseous nasal crests may require a highly specific combination of ontogeny, evolution, and environmental pressures in order to develop.
... The fossil record provides the requisite empirical evidence for testing this hypothesis. Compared to the record from southern Africa, which has been a focus of modern human origins research for decades, our understanding of the fossil history of East African ungulates over the last *200 kyr is only beginning to come to light (Marean and Gifford-Gonzalez 1991;Marean 1992;Assefa 2006;Assefa et al. 2008;Faith et al. 2011Faith et al. , 2012Faith et al. , 2014. However, the emerging evidence provides compelling examples of climate-driven range shifts that are consistent with hypotheses derived from ungulate biogeography. ...
... Alcelaphine bovids and equids dominate the assemblages, indicating the presence of open grassland vegetation distinct from the historic vegetation (White 1983). Several extinct bovids are present, including Rusingoryx atopocranion, Damaliscus hypsodon, Megalotragus sp., Syncerus antiquus, and an unnamed impala, all of which are characterized by dental or postcranial adaptations to grazing in open habitats (Klein 1980(Klein , 1994Faith et al. 2011Faith et al. , 2012Faith et al. , 2014. The presence of large gregarious grazers on Mfangano Island, some of which are migratory species, suggests a likely connection to the mainland when the deposits accumulated, requiring a ≥25 m reduction in lake level ). ...
... The presence of extinct grazing bovids characterized by massive body mass (Syncerus antiquus and Megalotragus sp.) or extreme hypsodonty (Damaliscus hypsodon, Rusingoryx atopocranion, and the unnamed impala), of which D. hypsodon and R. atopocranion are dominant, also implies the presence of non-analog environments. We have previously interpreted their presence, together with that of Grevy's zebra and oryx, as evidence for aridity (Tryon et al. 2010Faith et al. 2011, although this is at odds with the high diversity (see Faith 2013). A more detailed exploration of this conflict is beyond the scope of this study, but possible explanations could include the presence of a complex Late Pleistocene grazing succession (e.g., Brink and Lee-Thorp 1992) or an extinct migratory system (e.g., Marean 2010;Faith and Thompson 2013). ...
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To better understand the potential role of environmental change in mediating human dispersals across equatorial East Africa, this study examines the biogeographic histories of ungulates, including a summary of current knowledge and fossil evidence stemming from our fieldwork in the Kenyan portion of the Lake Victoria basin. Phylogeographic and paleontological evidence indicates that vegetation changes across Quaternary climate cycles mediated ungulate distributions and dispersals via the opening and closing of biogeographic barriers in equatorial East Africa. Dispersal capabilities would have been enhanced during phases of grassland expansion and diminished during phases of grassland contraction. We propose that the distribution and dispersal of diagnostic technological markers in the archaeological record may be similarly influenced by environmental changes. The Middle Stone Age record from the Lake Victoria region provides intriguing examples of possible environmentally mediated technological dispersals.
... At 66,400 km 2 , Lake Victoria is the largest lake in Africa by surface area (Adams, 1996). The habitats surrounding this lake have undergone substantial climate-driven changes throughout the Quaternary (Bootsma and Hecky, 2003;Nicholson, 1998), likely with profound impacts on human and other animal communities (e.g., Faith et al., 2011;Faith et al., 2015;Tryon et al., 2010;Tryon et al., 2012). However, until recently, an understanding of environmental variation prior to the Last Glacial Maximum has been poorly constrained, and the nature of spatial variation in environments and human behavior obscured. ...
... Stone artifacts from the Wasiriya beds include MSA Levallois cores and Levallois points as well as bifacial and unifacial trimmed points . Fauna are abundant and include both extinct and extant taxa (Faith et al., 2011;Pickford, 1984;Pickford and Thomas, 1984;Tryon et al., 2010). ...
... The eLVB and the Wasiriya beds in general include both extinct and extant taxa . The majority of specimens indicate open, semi-arid grasslands distinct from the evergreen bushlands, woodlands, and forests historically found in the region and otherwise common (Faith et al., 2011;Faith et al., 2012;Faith et al., 2014;Faith et al., 2015;Garrett et al., 2015;Tryon et al., 2010;Tryon et al., 2012). The Nyamita Valley follows the general open-arid pattern seen in the rest of the eLVB . ...
Article
The later Middle through early Late Pleistocene (~100–400 ka) of East Africa is an important time and place for the evolution of our species. This period records the first appearance of Homo sapiens and spans significant technological changes including the decline of large handheld stone tools characteristic of the Acheulean, the development of stone tool technologies collectively known as the Middle Stone Age (MSA). These include diverse Levallois prepared core techniques and the manufacture and use of pointed weapons. It is in association with MSA technologies in sub-Saharan Africa that most of the behaviors characteristic of modern humans first appear. This doctoral dissertation provides new chronological and archaeological data relevant to hominin behavior associated with MSA technology in the Middle and Late Pleistocene of East Africa. Improved chronological resolution is achieved through tephrostratigraphy, the correlation of volcanic ashes, combined with chronometric dating in two regions: the Kapthurin Formation in the Rift Valley, Baringo, Kenya and the eastern Lake Victoria Basin of western Kenya. New data on hominin behavior is provided by archaeological excavations of two sites: 1) The 196-226 ka Sibilo School Road Site in the Kapthurin Formation. 2) The 33–49 ka site of Nyamita Main in the eastern Lake Victoria Basin. The archaeology of the Kapthurin Formation and the eastern Lake Victoria Basin are connected thematically by the presence of MSA technology. These basins are also connected stratigraphically and chronologically, as this study shows, by tephra correlations between them. Results of this work demonstrate: 1) Levallois prepared core techniques, important aspects of MSA technology, are shown to be >380 ka in the Kapthurin Formation, ~100 kyr older than previously estimated in East Africa. 2) Long distance transport (>166 km) of high quality obsidian for stone tool manufacture was a feature of hominin behavior associated with Middle Pleistocene MSA technology ~200 ka ago. 3) MSA technology persisted in East Africa later than 49 ka and perhaps later than 33 ka, after Later Stone Age technologies, often considered categorically superior, are documented in the region. By demonstrating both the early and late presence of various aspects of MSA technology and associated hominin behavior this work shows that tephrostratigraphy and the excavation of new archaeological material in East Africa are productive means of producing new and important data on the MSA and the evolution of human behavior.
... Sparse artifacts and abundant fauna from open-air sites on Rusinga Island, Kenya, highlight the association of MSA humans with a diverse and arid-adapted ungulate community (Tryon et al., 2010(Tryon et al., , 2012Faith et al., 2011Faith et al., , 2013Faith et al., , 2014Faith et al., , 2015Faith et al., , 2016Tryon and Faith, 2013;Blegen et al., 2015;Beverly et al., 2015a, b;Garrett et al., 2015). The Wakondo locality is one of three main Pleistocene collecting areas on Rusinga, and lies on the southeastern slope of the island (UTM: 36M 0630458, 9953261),~20 m above the modern lake level of Lake Victoria ( Fig. 1 A and B). ...
... The Wakondo locality is one of three main Pleistocene collecting areas on Rusinga, and lies on the southeastern slope of the island (UTM: 36M 0630458, 9953261),~20 m above the modern lake level of Lake Victoria ( Fig. 1 A and B). Whereas Wakondo is better known for its early Miocene fossils, including the type specimen of the catarrhine primate Dendropithecus macinnesi (Andrews and Simons, 1977), the locality is also associated with abundant faunal remains and lithic artifacts derived from the Pleistocene Wasiriya Beds (Pickford and Thomas, 1984;Pickford, 1986;Tryon et al., 2010Tryon et al., , 2012Faith et al., 2011). Kent (1942) provides one of the first written observations of Rusinga's Pleistocene deposits, but the earliest formal description was published by Van Couvering (1972), and this work was later extended by Pickford and Thomas (1984) as context for the holotype of the extinct alcelaphin bovid, R. atopocranion. ...
... Our recent work on Rusinga's Pleistocene localities (primarily Wakondo, Nyamita, and Nyamsingula; Fig. 1B) is part of a broader research program reconstructing paleoenvironments, faunal communities, and hominin landscape use around the eastern shores of Lake Victoria (Tryon et al., 2010(Tryon et al., , 2012Faith et al., 2011Faith et al., , 2015Faith et al., , 2016Blegen et al., 2015;Beverly et al., 2015a, b;Garrett et al., 2015). The research in Wakondo's Pleistocene deposits was initiated through limited surface collections and rescue excavations conducted by Miocene paleontological researchers (McNulty et al., 2007;Peppe et al., 2009), who discovered three partial bovid skeletons eroding from sediments at the sub-locality at Wakondo referred to as "Bovid Hill" (Fig. 1D). ...
Article
The foraging behaviors of Middle Stone Age (MSA) early modern humans have largely been based on evidence from well-stratified cave sites in South Africa. Whereas these sites have provided an abundance of data for behavioral reconstruction that are unmatched elsewhere in Africa, they are unlikely to preserve evidence of the diversity of foraging strategies employed by MSA hunters who lived in a variety of ecological and landscape settings across the African continent. Here we describe the results of recent excavations at the open-air site of Bovid Hill at Wakondo, Rusinga Island, Kenya, which yielded 24 in situ MSA artifacts within an assemblage of bones comprised exclusively of the extinct alcelaphin bovid Rusingoryx atopocranion. The excavated faunal assemblage is characterized by a prime-age-dominated mortality profile and includes cut-marked specimens and an associated MSA Levallois blade-based artifact industry recovered from a channel deposit dated to 68 ± 5 ka by optically stimulated lumines-cence. Taphonomic, geologic, and faunal evidence points to mass exploitation of Rusingoryx by humans at Bovid Hill, which likely represents an initial processing site that was altered post-depositionally by fluvial processes. This site highlights the importance of rivers and streams for mass procurement in an open and seasonal landscape, and provides important new insights into MSA behavioral variability with respect to environmental conditions, site function, and tactical foraging strategies in eastern Africa. Bovid Hill thus joins a growing number of MSA and Middle Paleolithic localities that are suggestive of tactical hunting behaviors and mass capture of gregarious ungulate prey.
... However, the impact of these studies is limited because the relevant data archivesdparticularly proxies for temperature, moisture availability, and vegetationdare poorly resolved spatially and temporally (reviewed in Blome et al., 2012). Available faunal evidence indicates that early modern human populations in East Africa were part of extinct non-analog animal communities that were distinct in terms of taxonomic composition compared with regional modern mammal communities (Marean and Gifford-Gonzalez, 1991;Marean, 1992Marean, , 1997Tryon et al., 2010;Faith et al., 2011;Faith et al., 2012Faith et al., , 2013Faith et al., , 2014Tryon et al., 2012), and, as such, the paleoecology of these ancient populations remains poorly understood. Previous paleoenvironmental reconstructions of East African MSA sites using isotopic data have been restricted to the analysis of paleosols from site A5 (dated to~100e80 ka) at Aduma, Ethiopia (Yellen et al., 2005), and no published studies to date have examined the stable isotope composition of fossil mammals from this interval. ...
... The Pleistocene faunas from Rusinga and Mfangano (Table 1) contain the largest number of extinct species of any Pleistocene site in East Africa during the last~400 kyr (cf. Marean, 1992;Assefa et al., 2008;Domínguez-Rodrigo et al., 2008;Faith et al., 2011Faith et al., , 2012Faith et al., , 2013Faith et al., , 2014, with the extinct species Rusingoryx atopocranion (Pickford and Thomas, 1984;Faith et al., 2011) and Damaliscus hypsodon being the most abundant. Relevant to paleoenvironmental reconstruction, the faunas include several taxa indicating locally wet conditions, such as Hippopotamus and two species of reduncine bovids. ...
... The Pleistocene faunas from Rusinga and Mfangano (Table 1) contain the largest number of extinct species of any Pleistocene site in East Africa during the last~400 kyr (cf. Marean, 1992;Assefa et al., 2008;Domínguez-Rodrigo et al., 2008;Faith et al., 2011Faith et al., , 2012Faith et al., , 2013Faith et al., , 2014, with the extinct species Rusingoryx atopocranion (Pickford and Thomas, 1984;Faith et al., 2011) and Damaliscus hypsodon being the most abundant. Relevant to paleoenvironmental reconstruction, the faunas include several taxa indicating locally wet conditions, such as Hippopotamus and two species of reduncine bovids. ...
... The absence of extinct taxa typical of earlier East African sites (780,000 to 500,000 years ago), including the suid Kolpochoerus majus, the giant cercopithecid Theropithecus oswaldi, the elephantid Elephas recki, and the hippopotamid Hippopotamus gorgops (Deino and Potts, 1990;Potts and Deino, 1995;McBrearty et al., 1996;Potts, 1998;Geraads et al., 2004a;Gilbert and Asfaw, 2008), implies substantial faunal turnover during the middle Pleistocene. At the same time, Lainyamok is atypical of subsequent late Pleistocene sites in East Africa, here several extinct bovid species (e.g., long-horn buffalo (Syncerus antiquus), giant wildebeest (Megalotragus sp.), Rusingoryx atopocranion, and a small alcelaphine) are well documented and sometimes very abundant (Marean and Gifford-Gonzalez, 1991;Marean, 1992;Harrison and Baker, 1997;Tryon et al., 2010Tryon et al., , 2012Faith et al., 2011). ...
... Although Lainyamok no longer includes only extant species, it lacks several extinct taxa commonly found earlier in the middle Pleistocene (Potts, 1998). Fig. 8 illustrates the approximate last appearances of extinct middle-to-late Pleistocene mammals in East Africa (from Marean and Gifford-Gonzalez, 1991;Potts, 1998;Geraads et al., 2004a;O'Regan et al., 2005;Lehman, 2009;Tryon et al., 2010;Faith et al., 2011). Seven of the 12 extinct species are not confidently known from East African fossil sites younger than 400 ka. ...
... However, the possibility of a taxonomically modern Ethiopian fauna at this time should be treated with caution, since the known records include very few specimens firmly identified to species, due to high fragmentation and small samples (e.g., most alcelaphine specimens from Porc Epic Cave and the Kibish Formation are identified only to tribe). Our reclassification of the small alcelaphine material from Lainyamok contributes to a growing literature indicating that a taxonomically modern equatorial East African bovid community did not emerge until the Holocene (Marean and Gifford-Gonzalez, 1991;Marean, 1992;Tryon et al., 2010Tryon et al., , 2012Faith et al., 2011). Although the bovid species found today are well represented in the middle and late Pleistocene records, the numerically dominant taxa are now extinct. ...
Article
The middle Pleistocene fossil mammal assemblage from Lainyamok in the southern Kenya rift has previously been considered the oldest (330–392 ka) African mammal community consisting entirely of extant species, with the dominant bovid tentatively attributed to the southern African blesbok (Damaliscus cf. dorcas). We show that the blesbok-like fossils from Lainyamok belong to an extinct species, described here as Damaliscus hypsodon sp. nov. The D. hypsodon hypodigm includes the previously unnamed small alcelaphine material known from late Pleistocene sites elsewhere in Kenya and Tanzania. Its dental anatomy, together with an ecomorphological analysis of its postcrania, indicates that D. hypsodon grazed in open and arid grassland environments. Although Lainyamok is no longer represented entirely by extant species, the absence of species common earlier in the middle Pleistocene of East Africa suggests substantial faunal turnover between 500 and 400 ka. Damaliscus hypsodon persisted in East Africa until the end of the Pleistocene and its extinction can be attributed to a loss of arid grassland environments at the onset of the Holocene. The fossil evidence from southern Kenya suggests that the development of the taxonomically modern large mammal community was a long-term process characterized by the extinction of grazing specialists, with marked turnover occurring between ~ 500 and 400 ka and near the end of the Pleistocene.
... Increased rainfall would have altered the availability and composition of what were formerly arid grasslands and allowed moist-grass grazers (e.g. wildebeest, plains zebra) to replace arid-adapted grazers (Marean & Gifford-Gonzalez, 1991;Marean, 1992;Faith et al., 2011). Today, Grévy's zebra is unable to compete in the more mesic grasslands favoured by the plains zebra (Bauer et al., 1994;Rubenstein, 2010 Rubenstein (2010) shows that where the two species are found together, increased frequencies of the mesic-adapted plains zebra are associated with diminished forage intake in Grévy's zebra. ...
... The PCoA shows that the fossil distribution of Grévy's zebra parallels that of extinct Pleistocene ungulates, including S. antiquus, Megalotragus, Rusingoryx and the small alcelaphine (Fig. 3). As indicated by their morphological adaptations and palaeoenvironmental associations, these taxa were grazers that preferred open and seasonally arid grasslands (Vrba, 1987;Marean & Gifford-Gonzalez, 1991;Marean, 1992;Klein, 1994;Tryon et al., 2010Tryon et al., , 2012Faith et al., 2011), which seem to have been widespread in East Africa during much of the middle/late Pleistocene (Marean & Gifford-Gonzalez, 1991;Marean, 1992;Potts & Deino, 1995;Tryon et al., 2010Tryon et al., , 2012. These extinct ungulates can be distinguished from their extant relatives by extreme hypsodonty or massive body size, with Megalotragus and S. antiquus among the largest of their respective lineages (Alcelaphini and Bovini). ...
... Although an improved extinction chronology is necessary for some of the East African taxa, it is believed that environmental change at the onset of the Holocene is responsible for the extinctions (Marean & Gifford-Gonzalez, 1991;Marean, 1992;Faith et al., 2011). In particular, the extinctions are attributed to the same factors responsible for the extirpation of Grévy's zebra from southern Kenya, namely increased rainfall and the loss of arid grasslands together with increased competition with mesic-adapted grazers. ...
Article
Within the last several decades, Grévy's zebra (Equus grevyi) has undergone a massive reduction in geographical range and population size, largely as the result of human impacts. To place its recent decline in a deeper prehistoric context, and to understand the factors mediating its range and abundance over geological time frames, this study examines the fossil history of Grévy's zebra in equatorial East Africa. Equatorial East Africa. Presence/absence data for ungulates recovered from fossil sites spanning the last c. 400,000 years in Kenya and Ethiopia were compiled from the literature and from previously unreported palaeontological sites. Associations between Grévy's zebra and other taxa were examined using principal coordinates analysis and non-random species pairs were identified using a Bayesian approach. Changes in rainfall were reconstructed using the average hypsodonty index of ungulate species from fossil assemblages. Grévy's zebra was common during dry phases of the Pleistocene and was found to the south and west of its historical range, coinciding with an expansion of arid grasslands. At the onset of the Holocene, Grévy's zebra was extirpated from southern Kenya and almost completely disappeared from the fossil record. Grévy's zebra was associated with several specialized grazers that became extinct by the end of the Pleistocene. These extinctions and the decline of Grévy's zebra from the Pleistocene to the Holocene are explained by increased precipitation and the consequent loss of arid grasslands at the Pleistocene–Holocene transition. Grévy's zebra is never associated with domestic livestock, unlike the widespread plains zebra. Grévy's zebra thrived in equatorial East Africa during periods of the Pleistocene when environmental conditions favoured an expansion of arid grasslands. Environmental change across the Pleistocene–Holocene transition contributed to decreases in the range size and abundance of Grévy's zebra, setting the stage for the anthropogenic decline observed in recent decades. The spread of pastoralists in the middle Holocene may have additionally contributed to its prehistoric decline. Contemporary climate change warrants further consideration in planning for the long-term survival of Grévy's zebra.
... Pickford ( Pickford and Thomas 1984;Pickford 1986) collected and described Pleistocene fossils from both islands during the course of his wide-ranging surveys and remapping of the islands. We began the first dedicated geological, palaeontological and archaeological study of the Pleistocene deposits on both islands in 2009 ( Tryon et al. 2010;Faith et al. 2011). ...
... The presence of oryx (Oryx gazella) and Grevy's zebra (Equus grevyi), both of which inhabit relatively arid environments today (Kingdon 1982), suggests an arid grassland setting. The arid character of the grasslands is further supported by the presence of extinct bovids that are often associated with arid and open faunal communities ( Klein 1980Klein , 1994Vrba 1987;Marean and Gifford-Gonzalez 1991;Marean 1992), including the giant wildebeest (Megalotragus sp.), the giant buffalo (Syncerus antiquus), a small, unnamed, extremely hypsodont alcelaphine best known also from Lukenya Hill (Marean and Gifford-Gonzalez 1991;Marean 1992) and the medium-sized alcelaphine Rusingoryx atopocranion (Pickford and Thomas 1984;Faith et al. 2011) (Table 1). We follow Gentry (2010) in assigning the extinct giant buffalo to Syncerus rather than Pelorovis (see also Klein 1994, 731). ...
... We follow Gentry (2010) in assigning the extinct giant buffalo to Syncerus rather than Pelorovis (see also Klein 1994, 731). The extinct bovids may have interacted in an arid-adapted grazing succession ( Faith et al. 2011), similar to the grazing succession observed in the modern Serengeti (Bell 1971). Initial isotopic analyses of pedogenic carbonates, soil organic matter and ungulate tooth enamel are consistent with the interpretation that the Wasiriya Beds on Rusinga represent wetter (and perhaps more closed) environments within a larger arid grassland setting ( Garrett et al. 2010). ...
Article
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Surveys and excavations in 2009�2011 recovered fossil and artefact assemblages from late Pleistocene sediments on Rusinga and Mfangano islands (Lake Victoria, Kenya). Radiometric age estimates suggest that the Rusinga material dates to between 100 and 33 kya, whereas that from Mfangano may date to ]35 kya. The preservation of a large and diverse suite of vertebrate fossils is unusual for Pleistocene sites in the Lake Victoria region and the composition of the faunal assemblages from both islands strongly suggest an open, arid, grassland setting very different from that found inwesternKenya today.Middle Stone Age(MSA) artefacts fromRusinga and possible Later Stone Age (LSA) or MSA/LSA assemblages from Mfangano are distinct from Lupemban MSA sites characteristic of the Lake Victoria region and instead share a number of typological and technological features with late Pleistocene sites from open grassland settings in the East African Rift System. This highlights the complex roles that shifting environments, as well as temporal change, may have played in the development of regional variation among EquatorialAfrican artefact assemblages in the Pleistocene. Keywords: Middle Stone Age; Later Stone Age; Quaternary; aridity; Lake Victoria
... The late Pleistocene large mammal communities were composed of numerous extinct taxa, some of which were dominant members of the region's faunas until the onset of the Holocene (MacInnes, 1956;Marean and Gifford-Gonzalez, 1991;Marean, 1992;Faith, 2014;Faith et al., 2015;Lesur et al., 2016;Tryon et al., 2016). This emerging perspective has been reinforced by ongoing research in the Kenyan portions of the Lake Victoria Basin since 2008, which has documented numerous extinct taxa (Rusingoryx atopocranion, Damaliscus hypsodon, Kolpochoerus, and others) in late Pleistocene sediments, including new species or those formerly thought to have disappeared from eastern Africa during the middle Pleistocene (e.g., Tryon et al., 2010Tryon et al., , 2012Tryon et al., , 2016Faith et al., 2011Faith et al., , 2014Faith et al., , 2015Jenkins et al., 2017). These new data show that Homo sapiens in eastern Africa evolved among non-analog faunal communities (e.g., Faith et al., 2016), as has long been recognized for southern Africa (e.g., Klein, 1980). ...
... The Bovid Hill assemblage thus affords a rare opportunity to provide a more holistic understanding of its ecology. In addition to the bonebed accumulation at Bovid Hill, remains of the alcelaphin bovid Rusingoryx have been recovered from other late Pleistocene sediments (∼100-36 ka) around the Kenyan Lake Victoria Basin, including both Rusinga and Mfangano islands and mainland sites Luanda West and Karungu (Faith et al., 2011;O'Brien et al., 2016;Tryon et al., 2016;Blegen et al., 2017;Jenkins et al., 2017). ...
... Past work in the Lake Victoria Basin has documented the expansion of Serengeti-like grasslands across the region in the late Pleistocene (e.g., Tryon et al., 2010Tryon et al., , 2012Tryon et al., , 2016Faith et al., 2015;Garrett et al., 2015), likely in response to increased aridity and desiccation of the lake (e.g., Beverly et al., 2015aBeverly et al., , 2017Beverly et al., , 2020. This interpretation has been heavily influenced by the fossil faunas, including inferences based on the dominance of R. atopocranion, which was assumed to have had an affinity for open grassland habitats similar to extant alcelaphins (e.g., Faith et al., 2011;Faith, 2014). However, the craniodental remains of this species are unusual compared to other bovids and, indeed, are without parallel among other mammals-it has a large, hollow nasal crest otherwise known only from lambeosaurine hadrosaur dinosaurs (O'Brien et al., 2016). ...
Article
Rusingoryx atopocranion is an extinct alcelaphin bovid from the late Pleistocene of Kenya, known for its distinctive hollow nasal crest. A bonebed of R. atopocranion from the Lake Victoria Basin provides a unique opportunity to examine the nearly complete postcranial ecomorphology of an extinct species, and yields data that are important to studying paleoenvironments and human-environment interaction. With a comparative sample of extant African bovids, we used discriminant function analyses to develop statistical ecomorphological models for 18 skeletal elements and element portions. Forelimb and hin-dlimb element models overwhelmingly predict that R. atopocranion was an open-adapted taxon. However, the phalanges of Rusingoryx are remarkably short relative to their breadth, a morphology outside the range of extant African bovids, which we interpret as an extreme open-habitat adaptation. It follows that even recently extinct fossil bovids can differ in important morphological ways relative to their extant counterparts, particularly if they have novel adaptations for past environments. This unusual phalanx morphology (in combination with other skeletal indications), mesowear, and dental enamel stable isotopes, demonstrate that Rusingoryx was a grassland specialist. Together, these data are consistent with independent geological and paleontological evidence for increased aridity and expanded grassland habitats across the Lake Victoria Basin.
... Indeed, the faunal communities of these sites are dominated by arid-adapted taxa, including the extinct alcelaphins Damaliscus hypsodon, Megalotragus, and Rusingoryx atopocranion, and the bovin Syncerus antiquus. In addition, these sites and others in northern Tanzania demonstrate range expansions of arid-adapted taxa such as Oryx beisa and Equus grevyi (Marean and Gifford-Gonzalez, 1991;Marean, 1992a;Faith et al., 2011Faith et al., , 2012Faith et al., , 2013Faith et al., , 2015Tryon et al., 2010Tryon et al., , 2014Tryon et al., , 2015. These ungulate taxa are either rare (D. hypsodon, O. beisa, E. grevyi) or absent (Megalotragus, R. atopocranion, S. antiquus) in the Kibish Formation. ...
... The extinction of arid-adapted grassland ungulates by the Pleistocene-Holocene transition in East Africa has been attributed in part to wetter conditions at the onset of the Holocene (Marean and Gifford-Gonzalez, 1991;Marean, 1992a;Faith et al., 2011Faith et al., , 2012Faith, 2014). The evidence presented here suggesting that the Holocene humid phase is more pronounced than earlier humid phases may explain why arid-adapted grassland ungulates became extinct by the Pleistocene-Holocene transition, but persisted through previous humid phases of the late Quaternary. ...
Article
East Africa has produced the earliest record of Homo sapiens ~200ka and a punctuated record of Middle Stone Age and Later Stone Age behaviors. We lack, however, a detailed late Quaternary paleoenvironmental record for the region, particularly during humid periods. Without a regional record, hypotheses about the evolution and ecology of early Homo sapiens in East Africa remain vague and untestable. The Kibish Formation of southern Ethiopia presents a long, albeit punctuated, record of late Middle Pleistocene to Holocene faunal change in East Africa, which was deposited during humid periods. Here, we present oxygen and carbon stable isotope data of the Kibish ungulates and test whether there are environmental changes within the Kibish Formation. Significant differences in δ18O enamel isotopes are consistent with more humid conditions during the Holocene-age Member IV (~13-4ka) than either Pleistocene-age Member I (~196ka) or Member III (~104ka). Mesowear data document a shift toward more attritional wear among grazers in Member IV and are correlated with more depleted δ18O enamel values, suggesting that the wear pattern shift is linked to the onset of more humid conditions during the Holocene. δ13C enamel values show subtle variations through time, but do not suggest any major changes in diets. We propose that the paleoenvironmental differences evident in Member IV, based on δ18O enamel values, mesowear, and bovid abundances, may be explained by cooler and wetter conditions at the beginning of the Holocene in the lower Omo Valley. The evidence suggesting that the Holocene humid phase is more pronounced than earlier humid phases may explain why arid-adapted grassland ungulates became extinct in East Africa by the Pleistocene-Holocene transition, but persisted through previous humid phases of the late Quaternary.
... The extent to which modern environments provide precise analogues for Pleistocene eastern Africa is uncertain. Fossil fauna from MSA sites include a number of specialized grazers that became extinct by the Holocene, implying important differences in animal communities and grassland composition (Faith et al. 2011Marean 1992Marean , 1997. Five extinct mammals are reported from MSA sites, including an aardvark, Orycteropus crassidens (Lehmann 2009;MacInnes 1956), and four bovids characterized by extreme hypsodonty and/or body mass: a relative of the wildebeest, Rusingoryx atopocranion (Faith et al. 2011;Pickford and Thomas 1984); the giant wildebeest Megalotragus sp. ...
... Fossil fauna from MSA sites include a number of specialized grazers that became extinct by the Holocene, implying important differences in animal communities and grassland composition (Faith et al. 2011Marean 1992Marean , 1997. Five extinct mammals are reported from MSA sites, including an aardvark, Orycteropus crassidens (Lehmann 2009;MacInnes 1956), and four bovids characterized by extreme hypsodonty and/or body mass: a relative of the wildebeest, Rusingoryx atopocranion (Faith et al. 2011;Pickford and Thomas 1984); the giant wildebeest Megalotragus sp. (Kelly 1996;Tryon et al. 2012); an extinct blesbok, Damaliscus hypsodon Marean and Gifford-Gonzalez 1991); and the giant longhorn buffalo Syncerus antiquus (Marean 1992;Tryon et al. 2012). ...
Article
Eastern Africa is an important area to study early populations of Homo sapiens because subsets of those populations likely dispersed to Eurasia and subsequently throughout the globe during the Upper Pleistocene. The Middle Stone Age (MSA) archaeology of this region, particularly aspects of stone-tool technology and typology, is highly variable with only rare cases of geographic and temporal patterning. Although there are differences in timing and perhaps frequency of occurrence, those elements that make up the MSA lithic tool kit are also found at contemporaneous sites elsewhere in Africa and Eurasia, making it difficult to identify a unique archaeological signal for hominin dispersals out of eastern Africa. Rather, regional variation appears to be the outcome of possibly long-term interactions between particular physical and social environments experienced by hominin populations.
... The basal crosssection of what is preserved is more or less circular with weak transverse compression making the horn core distinct from the blesbok- Faith, unpublished). Fossil M. kattwinkeli specimens are from Koobi Fora (Harris, 1991), Rusinga (Faith et al., 2011), and Bouri (Vrba, 1997). like morphology of D. hypsodon; compression grows stronger distally. ...
... It was long argued that the East African faunal community was essentially modern in taxonomic composition by roughly 400,000 years ago (Potts and Deino, 1995;Potts, 1998), although Marean (1992) showed that the Late Pleistocene record at Lukenya Hill was dominated by a small extinct alcelaphin now attributed to Damaliscus hypsodon and also included extinct longhorn buffalo (Syncerus antiquus). Recent work in the Late Pleistocene sediments surrounding Lake Victoria has documented a previously unrecognized array of extinct bovid species (Tryon et al., 2010;Faith et al., 2011Faith et al., , 2012Tryon et al., 2012;Faith et al., 2014;Tryon et al., 2014;Faith et al., in press). The recognition of the extinct alcelaphins Damaliscus hypsodon, Rusingoryx atopocranion, and Megalotragus, in addition to the extinct bovin Syncerus antiquus and a large-bodied hypsodont impala, in southern Kenya less than 100,000 years ago has significantly altered views of the timing of turnover in Pleistocene faunas in East Africa. ...
... Geologic and tephrostratigraphic research around the region has demonstrated that deposits correlative to the Wasiriya Beds exist across several hundred square kilometers on Mfangano Island and along the eastern shore of Lake Victoria Faith et al., 2015;Blegen et al., 2015Blegen et al., , 2017. This research on Rusinga Island and the surrounding region has added many new details to our understanding of the geology, archaeology, and fossil assemblages of the region, which has allowed for studies of the archaeological and paleoecological change during the Late Pleistocene (Tryon et al., 2010Faith et al., 2011Faith et al., , 2014Faith et al., , 2015Garrett et al., 2015;Van Plantinga, 2011;Beverly et al., 2015aBeverly et al., , b, 2017Blegen et al., 2015Blegen et al., , 2017Jenkins et al., 2017). ...
... The fauna from Nyamita is dominated by semiarid grassland taxa (Table 1) including five extinct taxa, Rusingoryx atopocranion (Faith et al., 2011;O'Brien et al., 2016), a herd of which has been recovered 2 km away from the~68 ka bone bed at Bovid Hill, Wakondo (Jenkins et al., 2017), giant wildebeest (Megalotragus sp.), the exceptionally hypsodont, blesbok-like Damaliscus hypsodon , a distinctive, highly hypsodont impala , and the aardvark Orycteropus crassidens (MacInnes, 1956). The presence of hippopotamuses and two species of reedbuck (Redunca redunca and Redunca cf. ...
Article
In 2010, a hominin right humerus fragment (KNM-RU 58330) was surface collected in a small gully at Nyamita North in the Late Pleistocene Wasiriya Beds of Rusinga Island, Kenya. A combination of stratigraphic and geochronological evidence suggests the specimen is likely between ∼49 and 36 ka in age. The associated fauna is diverse and dominated by semiarid grassland taxa. The small sample of associated Middle Stone Age artifacts includes Levallois flakes, cores, and retouched points. The 139 mm humeral fragment preserves the shaft from distal to the lesser tubercle to 14 mm below the distal end of the weakly projecting deltoid tuberosity. Key morphological features include a narrow and weakly marked pectoralis major insertion and a distinctive medial bend in the diaphysis at the deltoid insertion. This bend is unusual among recent human humeri but occurs in a few Late Pleistocene humeri. The dimensions of the distal end of the fragment predict a length of 317.9 ± 16.4 mm based on recent samples of African ancestry. A novel method of predicting humeral length from the distance between the middle of the pectoralis major and the bottom of the deltoid insertion predicts a length of 317.3 mm ± 17.6 mm. Cross-sectional geometry at the midshaft shows a relatively high percentage of cortical bone and a moderate degree of flattening of the shaft. The Nyamita humerus is anatomically modern in its morphology and adds to the small sample of hominins from the Late Pleistocene associated with Middle Stone Age artifacts known from East Africa. It may sample a population closely related to the people of the out-of-Africa migration.
... The basal crosssection of what is preserved is more or less circular with weak transverse compression making the horn core distinct from the blesbok- Faith, unpublished). Fossil M. kattwinkeli specimens are from Koobi Fora (Harris, 1991), Rusinga (Faith et al., 2011), and Bouri (Vrba, 1997). like morphology of D. hypsodon; compression grows stronger distally. ...
... It was long argued that the East African faunal community was essentially modern in taxonomic composition by roughly 400,000 years ago (Potts and Deino, 1995;Potts, 1998), although Marean (1992) showed that the Late Pleistocene record at Lukenya Hill was dominated by a small extinct alcelaphin now attributed to Damaliscus hypsodon and also included extinct longhorn buffalo (Syncerus antiquus). Recent work in the Late Pleistocene sediments surrounding Lake Victoria has documented a previously unrecognized array of extinct bovid species (Tryon et al., 2010;Faith et al., 2011Faith et al., , 2012Tryon et al., 2012;Faith et al., 2014;Tryon et al., 2014;Faith et al., in press). The recognition of the extinct alcelaphins Damaliscus hypsodon, Rusingoryx atopocranion, and Megalotragus, in addition to the extinct bovin Syncerus antiquus and a large-bodied hypsodont impala, in southern Kenya less than 100,000 years ago has significantly altered views of the timing of turnover in Pleistocene faunas in East Africa. ...
Article
The Kibish Formation of southern Ethiopia has yielded the earliest fossils of Homo sapiens, ca. 196 ka, and has thus figured prominently in discussions of the origins of modern humans. Here we describe the fossil Bovidae from the Kibish Formation, a record that spans the late Middle Pleistocene to the early to mid-Holocene, and reconstruct aspects of their dietary ecology using mesowear analyses. All of the Kibish bovids represent extant taxa with the exception of the extinct blesbok-like alcelaphin Damaliscus hypsodon; extinct arid-adapted forms Syncerus antiquus and Megalotragus, common in other Late Quaternary sites, are notably absent. Mesowear of the Kibish bovids suggests that the Late Quaternary specimens were characterized by diets with considerably more abrasion-dominated wear relative to their extant conspecifics. Finally, the Kibish record provides supporting evidence for recent phylogeographic hypotheses by demonstrating significant range expansions of Aepyceros melampus, Connochaetes taurinus, Hippotragus equinus, and, to a lesser extent, Kobus kob in the late Middle Pleis-tocene through the early to mid-Holocene coincident with humid phases that punctuated dry spells of the Late Quaternary.
... It was once thought that East African fauna since the late middle Pleistocene was essentially modern (Potts and Deino, 1995). New work suggests, rather, that truly modern animal communities did not emerge until the Holocene (Marean and Gifford Gonzalez, 1991;Marean, 1992;Tryon et al., 2010;Tryon et al., in review;Faith et al., 2010). This has important paleoecological implications for the paleoenvironmental context of human evolution. ...
... Faunal remains such as reedbuck, waterbuck, and hippo, have been found in association with oryx and Grevy's zebra throughout the Wasiriya Beds. The latter species are arid-adapted and rarely range in regions with >500 mm rainfall/year (Tryon et al., 2010;Faith et al., 2010) and isotopic values from pedogenic carbonates (Garrett et al., 2010) indicate that the Wasiriya Beds likely represented a locally wet setting within a broader open arid grassland context (Tryon et al., in press). Fossil fauna in the Wasiriya Beds include specimens more typical of the modern grasslands east of Lake Victoria, suggesting that the lake was relatively contracted during Wasiriya Beds times (Tryon et al., in press). ...
Article
Wasiriya Beds, for the paleontological work on the fauna of these beds, and for other paleoenvironmental research in these beds. A robust tephrostratigraphic framework was established using diverse statistical methods. Radiocarbon dates confirm a Late Pleistocene age for these deposits. A valley-drainage model was integrated with a facies model to infer their paleoenvironmental history. Sediments suggest a generally sharply alternating wet and dry seasonality. Geological evidence does not suggest that the Lake Victoria region was especially arid just prior to the Last Glacial Maximum. Additional data from the lower Wasiriya Beds could elaborate on their paleoenvironmental significance.
... Among the bovids, alcelaphines and antilopines account for 76% of identified specimens, an abundance paralleled only in contemporary ecosystems characterized by arid to semi-arid grassland environments (e.g., Vrba, 1980;Alemseged, 2003). The dominant bovids, R. atopocranion and D. hypsodon, are characterized by exceptional hypsodonty ( Faith et al., 2011Faith et al., , 2012), a probable adaptation to consuming grasses in dry and gritty environments ( Marean, 1992;Damuth and Janis, 2011). Drier conditions than present are further supported by the recovery of arid-adapted Grevy's zebra and oryx (Oryx beisa; Kingdon, 1982;Faith et al., 2013), although the presence of blue wildebeest suggests at least seasonal availability of moist grasses ( Skinner and Chimimba, 2005). ...
... nov. ( Faith et al., 2011Faith et al., , 2012Faith et al., , 2014Faith, 2014). Of these, only S. antiquus and D. hypsodon are known from other Late Pleistocene faunal assemblages in East Africa ( Marean and Gifford-Gonzalez, 1991;Marean, 1992;Faith et al., 2012;Rowan et al., 2015). ...
Article
The opening and closing of the equatorial East African forest belt during the Quaternary is thought to have influenced the biogeographic histories of early modern humans and fauna, although precise details are scarce due to a lack of archaeological and paleontological records associated with paleoenvironmental data. With this in mind, we provide a description and paleoenvironmental reconstruction of the Late Pleistocene Middle Stone Age (MSA) artifact- and fossil-bearing sediments from Karungu, located along the shores of Lake Victoria in western Kenya. Artifacts recovered from surveys and controlled excavations are typologically MSA and include points, blades, and Levallois flakes and cores, as well as obsidian flakes similar in geochemical composition to documented sources near Lake Naivasha (250 km east). A combination of sedimentological, paleontological, and stable isotopic evidence indicates a semi-arid environment characterized by seasonal precipitation and the dominance of C4 grasslands, likely associated with a substantial reduction in Lake Victoria. The well-preserved fossil assemblage indicates that these conditions are associated with the convergence of historically allopatric ungulates from north and south of the equator, in agreement with predictions from genetic observations. Analysis of the East African MSA record reveals previously unrecognized north-south variation in assemblage composition that is consistent with episodes of population fragmentation during phases of limited dispersal potential. The grassland-associated MSA assemblages from Karungu and nearby Rusinga Island are characterized by a combination of artifact types that is more typical of northern sites. This may reflect the dispersal of behavioral repertoires-and perhaps human populations-during a paleoenvironmental phase dominated by grasslands. Copyright © 2015 Elsevier Ltd. All rights reserved.
... A putative fossil insular bovid from Africa is the al- celaphine Rusingoryx atopocranion Pickford & Thomas 1984, documented in Kenya in Pleistocene deposits from which Mesolithic artifacts are reported (see Faith et al. 2010). Pickford and Thomas (1984) raise the hy- pothesis that this species was an insular taxon restricted to Rusinga Island but geological evidence does not sup- port this scenario (Faith et al. 2010). ...
... A putative fossil insular bovid from Africa is the al- celaphine Rusingoryx atopocranion Pickford & Thomas 1984, documented in Kenya in Pleistocene deposits from which Mesolithic artifacts are reported (see Faith et al. 2010). Pickford and Thomas (1984) raise the hy- pothesis that this species was an insular taxon restricted to Rusinga Island but geological evidence does not sup- port this scenario (Faith et al. 2010). ...
Conference Paper
Insular fossil bovids, ranging in age from the latest Miocene to the Holocene, are widely recorded in Asian and Western Mediterranean islands. Several taxa characterized by different levels of endemism, but no species with an entirely identical adaptation, existed on different islands. Even considering that evolutionary processes, affecting size and morpho-functional features, mainly depend on inter- and intra-specific competition as well as on typology of free available niches, it is difficult to identify a dominant factor in the evolution of insular bovids. Moreover, highly modified insular biota show a mixture of apomorphic and plesiomorphic features that obscure their phyletic relationships, and make their origin and evolutionary patterns problematic to determine. A few common evolutionary traits can however be detected. Body size trends of insular fossil bovids are in agreement with the island rule pattern showing a negative relationship between Si (= mean mass of individuals from an insular population divided by M) and M ( = body mass of individuals of the mainland or ancestral form). The increased hypsodonty of molars (a quite common morphological trend in insular artiodactyls maybe related to an augment in the reproductive lifespan/longevity) is shared by most of insular bovids, including browsers such as the Javanese small boselaphine Duboisia santeng. A recurring pattern in insular bovids (although some exceptions exist ) is the simplification of the horn cores, as shown by both living (e.g., Bubalus depressicornis and Bubalus quarlesi) and highly modified fossil taxa (e.g., Duboisia santeng, Nesogoral spp., Myotragus). Conversely, morphological traits related to the so called ‘low-gear’ locomotion, exhibited by few insular bovids and regarded as typical adaptations acquired by insular artiodactyls, mainly depend on the presence of predators on the island. Assessing causal factors behind these modifications is not an easy task and a number of still unsolved issues need to be scrutinized, firstly to what extent phylogeny and bauplan might affect evolutionary patterns of insular bovids.
... NTAXA is known to be sensitive to assemblage sample size, with more species expected in larger samples (Grayson, 1984; Connochaetes cf. taurinus BPA (2) Alcelaphus buselaphus modern (20) Connochaetes gnou modern (11) Connochaetes taurinus modern ( (Harris, 1991;Vrba, 1997;Faith et al., 2011). Lyman, 2008). ...
... Understanding the evolutionary history of the herbivore niche within African bovids has traditionally relied on examining anatomical adaptations to diet, particularly those related to digestive strategy (Clauss et al. 2003;Codron et al. 2008a). From a palaeontological point of view, morphological characters discernible in the fossil record, such as anterior dentary shape, mandible shape and size and hypsodonty index, have been used to inform on the dietary preferences of both extant and extinct species (e.g. Gordon and Illius 1988;Solounias and Dawson-Saunders 1988;Solounias et al. , 1995Janis 1990Janis , 1995Janis , 2008Solounias and Moelleken 1993;Gordon 1999, 2001;MacFadden 2000;Williams and Kay 2001;Mendoza et al. 2002;Solounias and Semprebon 2002;Feranec 2007;Mendoza and Palmqvist 2007;Codron et al. 2008b;Faith et al. 2011Faith et al. , 2012Fraser and Theodor 2011). However, the significance of some of these traits in identifying dietary preferences has been re-examined in the light of the potentially confounding effects of phylogenetic affinity (P erez-Barberia and Gordon 1999Gordon , 2001Clauss et al. 2008;Raia et al. 2010). ...
Article
Understanding the evolutionary history of the herbivore niche within African bovids has traditionally relied on examining anatomical adaptations to diet, particularly those related to digestive strategy. More recently, mesowear and stable isotope analyses have been used to great effect to reconstruct dietary preferences. We use these dietary proxies to construct a morphology-free dietary ecospace and examine the topology of the phylogenetic rela-tionships of African bovids mapped onto this ecospace. The reconstructed dietary ecospace provides evidence for four distinct dietary classes: species with C3-or C4-dominated diets that produce low or high occlusal relief, likely related to diets high or low in abrasives, respectively. We detected no evidence for a discrete mixed feeder category; the species often categorized as such represent the end members of groups of species with either C3-or C4-dominated diets. Our analysis reveals high variability within the C4 grazing ecospace, and phylogenetic evidence indicates at least two pathways to grazing, likely related to the abrasive qualities of ingested food, which may be determined by the moisture content or the height of consumed grasses. These different pathways probably contribute to the high diversity of African grazers, both today and in the fossil record. C3 browsers (non-frugivores) also display a high degree of variation, but there are no species associated with highly abrasive diets and there is evidence for only a single evolutionary pathway. We find evidence for only one evolutionary route towards frugivory, which includes species with diets that produce both high and low occlusal reliefs. The cause of abrasive wear in frugivores may be related to grit and/or the hard parts of fruits, but this requires further examination.
... Among Early Stone Age sites in eastern Africa, several long-term archaeological projects have examined variation among archaeological sites and reconstructed paleoenvironments within relatively narrow temporal intervals to identify favored places on the landscape (e.g., Potts et al., 1999;Braun et al., 2008;Blumenschine et al., 2012). Comparable landscape-scale reconstructions for MSA sites in eastern Africa have yet to be done, although there have been efforts in this direction (e.g., Barboni et al., 1999;Ambrose, 2001;Yellen et al., 2005;Assefa et al., 2008;Tryon et al., 2010Tryon et al., , 2012Faith et al., 2011;Brown et al., 2012). Here, we build on these efforts to explore the reasons for open-air site occupation by MSA hominins on the eastern margins of the Lake Victoria basin in Equatorial Africa, drawing on archaeological evidence as well as approaches derived from ecology, geology, historical geography, and ethnobotany. ...
Article
Open-air archaeological sites record only a small fraction of the behavioral traces of mobile forager populations. Whereas caves and rockshelters were often occupied at least in part for protection from the elements, the reasons why human foragers occupied other places on the landscape (however briefly) are varied and not always readily recoverable. We develop a framework for interpreting human use of the landscape and modeling occupation of open-air sites using the archaeological and paleoenvironmental record of Middle Stone Age (MSA) sites from Rusinga and Mfangano Islands, located near the eastern margin of Lake Victoria. Paleoenvironmental reconstructions using fossil faunas suggest an arid grassland setting unlike the present. Paleoecological modeling of the habitats of extant and extinct bovids, combined with GIS-based reconstructions of lake level change, indicate that human occupation of these sites coincided with substantial declines in the level of Lake Victoria. During this time, both Rusinga and Mfangano would have been connected to the mainland and represented local topographic highs within an extensive grassland. Geological, ecological, and ethnobotanical observations suggest that these topographic high points would likely have been important sources of stone raw material, fresh water, and a variety of plant resources for food, fuel, and other purposes. In contrast, the grassy lowland plains were probably exploited primarily as a source of large game, which included numerous species of large gregarious grazers, several of which may have followed now extinct migration routes.
... They could, for example, be factors that have an effect at finer scales than have been measured here, occurred locally (e.g. what allowed both genera to reach such diversity within the Limpopo Basin?), or be factors that have not been accurately mapped, such as soils and geology, fire, pollinator availability or diversity, seed dispersers, or even aspects of long-term faunal dynamics not captured by current patterns of browser richness (Martin 1966;Faith et al. 2011). ...
Article
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1. It has been proposed that, across broad spatial scales, climatic factors are the main drivers of ecological patterns, while biotic factors are mainly important at local spatial scales. However, few tests of the effect of biotic interactions on broad-scale patterns have been conducted; conclusions about the scale-dependence of the importance of biotic interactions thus seem premature. 2. We developed an extensive database of locality records of one of Africa’s most conspicuous groups, the acacias (the genera Senegalia and Vachellia), and used species distribution models (SDMs) to estimate the distribution of all African acacias. 3. African acacias are particularly well adapted against mammalian herbivory; therefore, we hypothesized that browser diversity could be an important driver of acacia richness. Species richness maps for the two genera were created from SDM-generated maps. Ordinary least square (OLS) regressions and, to consider spatial autocorrelation, simultaneous autoregressive (SAR) analyses were used to model richness of the two genera in relation to mammalian browser richness, current environment (including climate), and climate history since the Last Glacial Maximum (LGM). We used variation partitioning to determine what percentage of variation could be explained by these three groups of factors. 4. Both genera showed centres of richness in East Africa and the Limpopo Basin of southern Africa. Browser richness was the best explanatory variable for richness of both genera. Environmental factors explained negligible variation in the richness of Senegalia, but some variation in Vachellia. For both genera, the residuals of the species richness model of one genus also explained much variation in the richness of the other genus, indicating that common factors not considered in the richness analyses here may additionally be driving the richness of both genera. 5. Mechanisms that could generate a correlation between browser and acacia richness are proposed, and differences in the determinants of richness patterns of Senegalia and Vachellia discussed in the light of the two genera’s history of colonization of Africa. 6. Synthesis. This is the first study that demonstrates that consumer diversity can influence richness patterns at continental scales and demonstrates that biotic factors can drive richness even at broad spatial scales.
... The general paleoenvironmental and paleoclimatic conditions, specific habitats and available food resources of early hominins are factors that may inform key questions pertaining to the origin, evolution, and extinction of hominin species, as well as understanding hominin morphological and behavioral adaptations (Dart, 1925; Robinson, 1954; Vrba, 1985; Reed, 1997; Alemseged and Bobe, 2009). Paleoenvironmental reconstruction of early hominin fossil localities in Africa is often based on a variety of independent lines of evidence such as isotopic composition of paleosol carbonate (e.g., Levin et al., 2004; Aronson et al., 2008; Cerling et al., 2011 ), taxonomic uniformitarianism (e.g., Vrba, 1974 Vrba, , 1980 Vrba, , 1985), faunal abundance (e.g., Reed, 1997 Reed, , 2008 Bobe et al., 2007 ), ecomorphology (e.g., Kappelman et al., 1997; DeGusta and Vrba, 2003; Plummer et al., 2008; Faith et al., 2011), and pollen analysis (e.g., Bonnefille et al., 2004). The use of carbon and oxygen stable isotopes of mammalian tooth enamel has also been proven a successful tool for the study of paleodiets and the reconstruction of habitats of both hominin and non-hominin species (Lee-Thorp, 1989; Bocherens et al., 1996; Kohn et al., 1996; Kingston, 1999; Sponheimer et al., 1999; Zazzo et al., 2000; Schoeninger et al., 2003; Cerling et al., 2003a Cerling et al., , 2003c Levin et al., 2008; White et al., 2009; Bedaso et al., 2010). ...
... Together, these lines of evidence provide abundant support for the argument that environmental change across the Pleistocene-Holocene transition played a central role in the CFR extinctions (Klein, 1980;Marean, 1990;Brink, 1999;Marean, 2010;Faith, 2011aFaith, , 2011b. This parallels the emerging picture of late Quaternary megafaunal extinctions in equatorial East Africa, where the losses are restricted to specialized grazers, including some taxa also known from the CFR (e.g., Megalotragus, Syncerus antiquus), and are attributed to changes to the structure of Pleistocene grasslands (Marean, 1990;Marean and Gifford-Gonzalez, 1991;Marean, 1992;Faith et al., 2011). Klein (1980Klein ( , 1984 suggests that environmental change alone is an insufficient explanation for the CFR extinctions. ...
Article
Precipitation, primary productivity, and herbivore biomass are linked to ungulate richness (the number of species) in African ecosystems. This study compares the richness of late-middle to late Pleistocene and Holocene ungulate assemblages in southern Africa's Cape Floral Region (CFR). Regression analysis demonstrates that Pleistocene ungulate assemblages are significantly richer than their Holocene counterparts. Elevated Pleistocene ungulate richness is not explained by differential time-averaging, the presence of extinct taxa in Pleistocene assemblages, or by Middle Stone Age (MSA) to Later Stone Age (LSA) technological change, but instead by a greater number of grazing species in Pleistocene faunal communities. Based on modem African analogs, this implies that the Pleistocene assemblages examined here sample time intervals characterized by elevated primary productivity, particularly of grassland habitats, greater ungulate biomass. and altered rainfall regimes. Declining ungulate richness from the Pleistocene to the Holocene supports the hypothesis that the extinction of specialized grazers at the Pleistocene-Holocene transition in the CFR was driven by declining productivity and availability of grassland habitats. The contrast between rich Pleistocene ungulate communities and impoverished Holocene ungulate communities may also explain important differences in MSA and LSA subsistence behavior. It is proposed that intensified exploitation of fish, shellfish, tortoises, and seabirds by LSA foragers during the Holocene reflects an expansion of diet breadth in response to diminished ungulate biomass on the landscape rather than fundamental behavioral/cognitive advances.
... Population density or size is notoriously difficult to demonstrate from fossil or archaeological data (but see recent efforts for Late Pleistocene western Europe by Mellars and French, 2011). While a number of site-specific approaches have shown that human impacts on various animal taxa can be a good measure of population pressure (Klein, 1980;Stiner et al., 1999;Faith et al., 2011), these sorts of data are not available or reliable for many African sites. Although problematic, our use of site density as a measure of Pleistocene hominin demography for a given time interval remains the best measure available for our pan-African approach. ...
... Stone artifacts from the Wasiriya beds include MSA Levallois cores and Levallois points as well as bifacial and unifacial trimmed points (Tryon et al., 2010). Fauna are abundant and include both extinct and extant taxa (Pickford and Thomas, 1984;Tryon et al., 2010;Faith et al., 2011). Water-dependent taxa are present (e.g., Hippopotamus), but the majority of specimens indicate open, semi-arid grasslands distinct from the evergreen bushlands, woodlands, and forests historically found in the region (Garrett et al., 2015;Tryon et al., 2010Tryon et al., , 2014. ...
Article
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The tephrostratigraphic framework for Pliocene and Early Pleistocene paleoanthropological sites in East Africa has been well established through nearly 50 years of research, but a similarly comprehensive framework is lacking for the Middle and particularly the Late Pleistocene. We provide the first detailed regional record of Late Pleistocene tephra deposits associated with artifacts or fossils from the Lake Victoria basin of western Kenya. Correlations of Late Pleistocene distal tephra deposits from the Wasiriya beds on Rusinga Island, the Waware beds on Mfangano Island and deposits near Karungu, mainland Kenya, are based on field stratigraphy coupled with 916 electron microprobe analyses of eleven major and minor element oxides from 50 samples. At least eight distinct distal tephra deposits are distinguished, four of which are found at multiple localities spanning >60 km over an approximately north to south transect. New optically stimulated luminescence dates help to constrain the Late Pleistocene depositional ages of these deposits. Our correlation and characterization of volcaniclastic deposits expand and refine the current stratigraphy of the eastern Lake Victoria basin. This provides the basis for relating fossil- and artifact-bearing sediments and a framework for ongoing geological, archaeological and paleontological studies of Late Pleistocene East Africa, a crucial time period for human evolution and dispersal within and out of Africa.
... Although sample sizes are small, the relative abundance of alcelaphin specimens among the bovids from the Nyamita excavation is similar to large samples of faunal material from the eLVB (Tryon et al. 2010, and is matched in contemporary arid to semi-arid grassland environments (Alemseged 2003;Vrba 1980). The extinct bovids at Nyamita Main, R. atopocranion and D. hypsodon, are characterized by exceptional hypsodonty (Faith et al. 2011;2012). This adaptation is interpreted as suited to consuming grasses in dry and gritty environments (Damuth and Janis 2011;Faith et al. 2012;Marean 1992 (Tryon et al. 2010;2014) as well as Levallois points (n=2) made in the red and white mottled chert and lava raw materials that dominate the lithic assemblage from the 2013 excavations (see Figure 13). ...
Article
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Here we report tephra correlations, lithic artifacts, obsidian sourcing data, and fauna from nine Late Pleistocene localities of the eastern Lake Victoria basin of western Kenya, as well as new excavations from the 49-36 ka site of Nyamita Main on Rusinga Island. The Late Pleistocene of Africa is an important period for the evolution and dispersals of Homo sapiens. A conspicuous behavioral feature of this period is the replacement of Middle Stone Age (MSA) technologies by Later Stone Age (LSA) technologies. Current research shows this process is complex with the LSA appearing and the MSA disappearing at different times in different places across Africa. Accounting for this pattern requires a precise chronology, detailed evidence of past human behavior and environmental reconstructions of the appropriate scale. Data presented here provide this detail. Tephra correlations improve the regional chronology and expand the lateral area of Late Pleistocene eastern Lake Victoria basin exposures from ~650km 2 to >2500km 2. Lithic artifacts show MSA technology is present younger than 36 ka in western Kenya, 25-35 kyr younger than the first appearance of early LSA technology elsewhere in equatorial East Africa. Obsid-ian sourcing data presented here shows the use of the same raw material sources by MSA and LSA populations through long periods of time from >100 ka through <36 ka. The methods employed here provide the temporal resolution and appropriate geographic scale to address modern human behavioral evolution.
... Stone artifacts from the Wasiriya beds include MSA Levallois cores and Levallois points as well as bifacial and unifacial trimmed points (Tryon et al., 2010). Fauna are abundant and include both extinct and extant taxa (Pickford and Thomas, 1984;Tryon et al., 2010;Faith et al., 2011). Water-dependent taxa are present (e.g., Hippopotamus), but the majority of specimens indicate open, semi-arid grasslands distinct from the evergreen bushlands, woodlands, and forests historically found in the region (Garrett et al., 2015;Tryon et al., 2010Tryon et al., , 2014. ...
... The presence of arid-adapted species outside of their historic ranges, especially Grevy's zebra (Equus grevyi) and oryx (Oryx cf. beisa), is consistent with a reduction in precipitation (Faith et al., 2013), as is the dominance of extinct species characterized by exceptional hypsodonty (Faith et al., 2011. Drier conditions are further suggested by the geochemical composition of the paleosols at Kisaaka, which provide precipitation estimates of~760e960 mm/yr through the Late Pleistocene sequence (Beverly et al., 2015a). ...
... However, they differ from those of comparative specimens of wildebeest in having simpler infundibula, and most notably so in the second molar. In this regard they more closely resemble the teeth of Alcelaphus and the extinct Rusingoryx but differ from the latter in having a more complex occlusal pattern and pronounced styles and ribs (see Faith et al., 2011). Given the limited number of specimens and difficulties in distinguishing between like-sized alcelaphines, we follow Thomas et al. (1998) (Fig. S4). ...
Article
In recent years, the Arabian Peninsula has emerged as a key region for elucidating hominin and faunal evolution and dispersals between Africa and Eurasia. Central to this research is the middle Pleistocene site of Ti's al Ghadah (TAG) which has yielded a diverse and abundant fossil faunal assemblage and the earliest chronometrically dated evidence for hominins in this part of the world. Here, we present the first detailed taphonomic study of the large Unit 5 fossil assemblage from the site. We aim to assess which actor/s were responsible for the accumulation of the assemblage and evaluate evidence that might be consistent with the accumulation of fauna by hominins. We also describe, for the first time, fossils and lithic artefacts from stratigraphic horizons not previously considered, providing taphonomic insights into their accumulation. The taphonomic work shows that the Unit 5 faunal assemblage was accumulated by ambush predators, likely large felids and hominins, in a lake side environment, and that carcasses were subsequently scavenged by more durophagus carnivores such as hyenas and canids. Less can be reliably said regarding the newly described fossil assemblages given their poor preservation and significant wind abrasion, but large carnivores again appear to have played a role, and hominins probably played a role in the accumulation of at least one of these. This study provides the first detail insights into the interplay between hominins, carnivores, and herbivores in Arabia, and suggests that watering holes have been a focus on the Arabian landscape for resources since the middle Pleistocene.
... Aepyceros includes a South African form, A. helmoedi (Brink et al., 2012), and a hypsodont unnamed species from Kenya (Faith et al., 2014). Several unnamed, potentially new Alcelaphini species have been noted at Middle Pleistocene sites (Klein et al., 2007; de Ruiter et al., 2008; Faith et al., 2011). Extinct species of the genus Antidorcas are known, for example at Redcliff Cave (A. bondi, Cruz-Uribe, 1983) and from South Africa's Western Cape (A. australis. ...
Article
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This paper describes a large collection of Quaternary fossil fauna from the Luangwa Rift Valley, Zambia. Stone Age artefacts have been recovered from stratified fluvial contexts, but no in situ fossil faunas have yet been recovered. We report on 500 fossil specimens collected from the surface of point bars exposed seasonally along the banks of the main Luangwa River channel. We used non-destructive X-ray fluorescence analysis of the fossils' chemical signatures to determine whether they derive from one or many primary contexts, and the relationship between chemical signature and state of preservation. Specimens are identified to taxon (genus) to reconstruct palaeoenvironments and biochronology. A relatively wide range of taxa is identified, including a fossil hominin talus, described here. None of the fossils is positively attributable to extinct species, except a femur of an extinct Theropithecus reported in 2003. Although no additional extinct taxa were identified, some of the remains were attributable to genera that are not currently found in this region. The results suggest that most of the assemblage derives from sediments which are Middle Pleistocene or later, and that past environments in the Luangwa Valley may have differed from the habitat availability found today.
... The Late Pleistocene geology, fossils, and MSA artifacts from Rusinga and Mfangano Islands have been the focus of research since 2009 ( Tryon et al., 2010Tryon et al., , 2012Tryon et al., , 2014Faith et al., 2011Faith et al., , 2012Faith et al., , 2014Van Plantinga, 2011;Jenkins et al., 2017;Blegen et al., in press). More recently, this research has expanded to include the deposits around the region known as Karungu ∼40 km to the south near the town of Sori ( Figure 1; Beverly et al., 2015a,b;Blegen et al., 2015;Faith et al., 2015). ...
Article
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The impact of changing environments on the evolution and dispersal of Homo sapiens is highly debated, but few data are available from equatorial Africa. Lake Victoria is the largest freshwater lake in the tropics and is currently a biogeographic barrier between the eastern and western branches of the East African Rift. The lake has previously desiccated at ~17 ka and again at ~15 ka, but little is known from this region prior to the Last Glacial Maximum. The Pleistocene terrestrial deposits on the northeast coast of Lake Victoria (94–36 ka) are ideal for paleoenvironmental reconstructions where volcaniclastic deposits (tuffs), fluvial deposits, tufa, and paleosols are exposed, which can be used to reconstruct Critical Zones (CZ) of the past (paleo-CZs). The paleo-CZ is a holistic concept that reconstructs the entire landscape using geologic records of the atmosphere, hydrosphere, lithosphere, biosphere, and pedosphere (the focus of this study). New paleosol-based mean annual precipitation (MAP) proxies from Karungu, Rusinga Island, and Mfangano Island indicate an average MAP of 750 ± 108 mm year−1 (CALMAG), 800 ± 182 mm year−1 (CIA-K), and 1,010 ± 228 mm year−1 (PPM1.0) with no statistical difference throughout the 11 m thick sequence. This corresponds to between 54 and 72% of modern precipitation. Tephras bracketing these paleosols have been correlated across seven sites, and sample a regional paleo-CZ across a ~55 km transect along the eastern shoreline of the modern lake. Given the sensitivity of Lake Victoria to precipitation, it is likely that the lake was significantly smaller than modern between 94 and 36 ka. This would have removed a major barrier for the movement of fauna (including early modern humans) and provided a dispersal corridor across the equator and between the rifts. It is also consistent with the associated fossil faunal assemblage indicative of semi-arid grasslands. During the Late Pleistocene, the combined geologic and paleontological evidence suggests a seasonally dry, open grassland environment for the Lake Victoria region that is significantly drier than today, which may have facilitated human and faunal dispersals across equatorial East Africa.
... Ma and the Upper Ndolanya Beds (UNB) at 2.66 Ma (Kaiser et al., 2011). Similarly, Faith and collaborators (Faith et al., 2011) have also used this proxy to investigate the paleoecology of a previously poorly known bovid species from Rusinga Island in Kenya. Most recently, the identification of bovid remains from the Kibish Formation (~196-8Ka), Ethiopia, included the evaluation of mesowear (Rowan et al., 2015). ...
Article
The Pliocene site of Kanapoi is key to our understanding of the environmental context of the earliest species of Australopithecus. Various approaches have been used to reconstruct the environments of this site, and here we contribute new data and analyses using mesowear and hypsodonty. The dental traits of 98 bovids, suids and rhinocerotids from Kanapoi were analyzed using these proxies. Results indicate that most of the animals analyzed had a relatively abrasive diet. Bovids in the assemblage incorporated more grass into their diet than do modern species of the same tribe or genus. Although Pliocene Kanapoi likely had complex environments, our analysis indicates that grassy habitats were a dominant component of the ecosystem, a conclusion that supports the results of previous investigations of the paleoecology of the site.
... Mid-frontal suture well-fused and barely discernible. Teeth are immediately distinguished by their large size (m3 lengths mainly 38e42 mm, which is larger than extant Connochaetes, Fig. A4) and simple occlusal morphology (Gentry and Gentry, 1978a;Faith et al., 2011). The single complete mandible confirms the loss of the second premolar in this species. ...
Article
Eight years of excavation work by the Olduvai Geochronology and Archaeology Project (OGAP) has produced a rich vertebrate fauna from several sites within Bed II, Olduvai Gorge, Tanzania. Study of these as well as recently re-organized collections from Mary Leakey's 1972 HWK EE excavations here provides a synthetic view of the faunal community of Olduvai during Middle Bed II at ∼1.7-1.4 Ma, an interval that captures the local transition from Oldowan to Acheulean technology. We expand the faunal list for this interval, name a new bovid species, clarify the evolution of several mammalian lineages, and record new local first and last appearances. Compositions of the fish and large mammal assemblages support previous indications for the dominance of open and seasonal grassland habitats at the margins of an alkaline lake. Fish diversity is low and dominated by cichlids, which indicates strongly saline conditions. The taphonomy of the fish assemblages supports reconstructions of fluctuating lake levels with mass die-offs in evaporating pools. The mammals are dominated by grazing bovids and equids. Habitats remained consistently dry and open throughout the entire Bed II sequence, with no major turnover or paleoecological changes taking place. Rather, wooded and wet habitats had already given way to drier and more open habitats by the top of Bed I, at 1.85-1.80 Ma. This ecological change is close to the age of the Oldowan-Acheulean transition in Kenya and Ethiopia, but precedes the local transition in Middle Bed II. The Middle Bed II large mammal community is much richer in species and includes a much larger number of large-bodied species (>300 kg) than the modern Serengeti. This reflects the severity of Pleistocene extinctions on African large mammals, with the loss of large species fitting a pattern typical of defaunation or 'downsizing' by human disturbance. However, trophic network (food web) analyses show that the Middle Bed II community was robust, and comparisons with the Serengeti community indicate that the fundamental structure of food webs remained intact despite Pleistocene extinctions. The presence of a generalized meat-eating hominin in the Middle Bed II community would have increased competition among carnivores and vulnerability among herbivores, but the high generality and interconnectedness of the Middle Bed II food web suggests this community was buffered against extinctions caused by trophic interactions.
Article
We report on the Late Pleistocene (36-12 ka) mammals from Kibogo in the Nyanza Rift of western Kenya, providing (1) a systematic description of the mammal remains, (2) an assessment of their paleoenvironmental implications, and (3) an analysis of the biogeographic implications of non-analog species associations. Kibogo has yielded one of the largest paleontological assemblages from the Late Pleistocene of eastern Africa, and it is dominated by grassland ungulates (e.g., equids and alcelaphin antelopes), including an assortment of extralimital (e.g., Equus grevyi, Ceratotherium simum, Redunca arundinum) and extinct species (Syncerus antiquus, Damaliscus hypsodon, Megalotragus sp.). The composition of the fauna, in conjunction with the soils and topography of the region, indicate the local presence of edaphic grassland situated within a broader environment that was substantially grassier and likely drier than at present. In contrast to non-analog faunas from higher latitudes (e.g., North America and western Eurasia), the climatic niches of non-analog species associations strongly overlap, indicating that non-analog climate regimes during the Late Pleistocene of eastern Africa are not necessary to account for the former association of presently allopatric species. The Kibogo faunas add to a growing body of evidence implying that the composition of present-day African herbivore communities is distinct from those of the geologically recent past.
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To understand the current biodiversity crisis, it is crucial to determine how humans have affected biodiversity in the past. However, the extent of human involvement in species extinctions from the Late Pleistocene onward remains contentious. Here, we apply Bayesian models to the fossil record to estimate how mammalian extinction rates have changed over the past 126,000 years, inferring specific times of rate increases. We specifically test the hypothesis of human-caused extinctions by using posterior predictive methods. We find that human population size is able to predict past extinctions with 96% accuracy. Predictors based on past climate, in contrast, perform no better than expected by chance, suggesting that climate had a negligible impact on global mammal extinctions. Based on current trends, we predict for the near future a rate escalation of unprecedented magnitude. Our results provide a comprehensive assessment of the human impact on past and predicted future extinctions of mammals.
Book
Paleozoology and Paleoenvironments outlines the reconstruction of ancient climates, floras, and habitats on the basis of animal fossil remains recovered from archaeological and paleontological sites. In addition to outlining the ecological fundamentals and analytical assumptions attending such analyzes, J. Tyler Faith and R. Lee Lyman describe and critically evaluate many of the varied analytical techniques that have been applied to paleozoological remains for the purpose of paleoenvironmental reconstruction. These techniques range from analyses based on the presence or abundance of species in a fossil assemblage to those based on taxon-free ecological characterizations. All techniques are illustrated using faunal data from archaeological or paleontological contexts. Aimed at students and professionals, this volume will serve as fundamental resource for courses in zooarchaeology, paleontology, and paleoecology.
Article
Endemic bovids are intriguing elements of insular faunas. The living species include the Japanese serow (Capricornis crispus) and the Formosan serow (C. swinhoei), the tamaraw from Mindoro, Philippines, (Bubalus mindorensis) and the anoas (B. depressicornis and B. quarlesi), 2 species of dwarf buffalos endemic to Sulawesi, Indonesia. Fossil endemic bovids are only recorded in some Asian, North American and Western Mediterranean islands. Here we present a comprehensive overview of the changes in body size and evolutionary patterns exhibited by both extant and extinct insular bovids. Our appraisal indicates that each insular representative of Bovidae shows its own peculiar evolutionary model, albeit some parallel trends exist (e.g. reduction in body size, allometric changes in limb bones, alteration of the life history traits). Some changes in morphology (e.g. the simplification of horn cores, the increase in hypsodonty, the acquisition of a 'low-gear' locomotion), for instance, appear as common, albeit not general, patterns triggered by a combination of selective forces. Body size patterns support the 'generality of the island rule' and suggest that biotic interaction had/have a major role in influencing body size evolution in these species, although in different ways on different islands. All things considered, available evidence suggest that a major role in the evolution of insular bovids is played by the structure of the insular community, the nature of available niches and by the dynamics of ecological interactions.
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The use of mesowear to infer diets of extinct species is fast becoming widespread in palaeoecological studies. Nevertheless, traditional mesowear analyses suffer from a specimen number limitation, in that a minimum number of specimens identified to the species level is necessary to make accurate dietary predictions. This is problematic in many fossil African antelope (Mammalia: Bovidae) assemblages, where isolated teeth cannot always be assigned to species. Hereweexplore the possibility of using simple dental metrics to predict diets on the basis of individual teeth as well as gnathic rows using linear discriminant function analyses.We find that browsers are accurately classified at both the individual and species levels, across all models and tooth positions. Mixed feeders and grazers are classified accurately only sometimes, and this is probably a reflection of the more limited sample size of larger bodied species in our study. Body size was a highly significant predictor of the inaccurate classifications obtained in our models, with larger bodied species tending to grazing classifications and smaller bodied species browsing classifications. Nevertheless, the models correctly classify the majority of specimens we examined to their correct trophic group, as determined through stable isotope analyses or as defined through the literature. The methods outlined hold some promise for determining the diets of isolated fossil specimens unassigned to species in a simple manner and, when used in conjunction with other palaeodietary and palaeoecological proxies, may help determine palaeoenvironments more accurately.
Chapter
Ancestors of mammals separated from reptiles and birds during the Carboniferous. Early members of the mammalian lineage, called “mammal-like reptiles” as they lack mammalian specializations, flourished during the Permian. Survivors of the Permian-Triassic boundary mass extinction progressively developed mammalian characters. The basic characteristic whereby the Mammalia are defined is the structure of the middle ear. In “reptiles”, including mammal-like reptiles, the lower jaw consists of several bones, one of which, the dentary, contains the teeth, another, the articular, forms a joint with a bone called the quadrate in the cranium, and there is only a single bone, the stapes, in the middle ear. Through the Miocene and Pliocene, mammalian lineages seem to have undergone diversification and extinction at rates that had characterised most of the Cenozoic. It was only the Pleistocene that saw elevated extinction rates, mainly affecting large mammals. The human lineage may have separated from that of chimpanzees some 6–7 million years ago, but about 4 million years ago there was an episode of interbreeding between the two lineages, leaving humans with an X chromosome that is markedly more chimpanzee-like than the rest of the genome. Comparative morphology and DNA analysis agree chimpanzeesare the closest living relatives of humans, followed by gorillas, then orangutans and then gibbons. Most molecular clock calculations indicate that the human and chimpanzee lineages separated some 6 million years ago. Sahelanthropus tchadensis is plausibly promoted as the earliest member of the Hominini.
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The ecological preferences of ruminant artiodactyls are commonly used to reconstruct the paleoenvironment of Neogene fossil localities throughout Africa. However, comparatively little research has focused on the ecology of ruminant artiodactyls from the Miocene of Africa. Here, we contribute new molar mesowear and hypsodonty data for the ruminant artiodactyls from the early and middle Miocene of Kenya and Uganda. Macroscopic dental characteristics of 608 tragulids, stem pecorans, giraffoids, and bovids dated to between 20 and 13.7 Ma were analyzed. Our hypsodonty results reveal that, whereas tragulids remain brachydont throughout the early and middle Miocene, pecoran ruminants experience an increase in hypsodonty due to the appearance of high-crowned bovids and climacoceratids that migrate into eastern Africa in the middle Miocene. Results from dental mesowear analysis suggest that all tragulids and pecorans were likely browsers, with only one taxon showing mesowear values that overlap with both browsers and mixed feeders in both the upper and lower molars (Canthumeryx sirtensis). None of the taxa analyzed had mesowear scores indicative of a grazing diet. Surprisingly, middle Miocene bovids and climacoceratids, despite possessing gross tooth morphologies adapted to abrasive diets, were largely utilizing a browsing diet. Although the early and middle Miocene habitats of eastern Africa were likely very heterogenous, none of the ruminant artiodactyls present in these habitats is interpreted as having incorporated grasses into their diet in significant quantities.
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Much debate has revolved around the question of whether the mode of evolutionary and ecological turnover in the fossil record of African mammals was continuous or pulsed, and the degree to which faunal turnover tracked changes in global climate. Here, we assembled and analyzed large specimen databases of the fossil record of eastern African Bovidae (antelopes) and Turkana Basin large mammals. Our results indicate that speciation and extinction proceeded continuously throughout the Pliocene and Pleistocene, as did increases in the relative abundance of arid-adapted bovids, and in bovid body mass. Species durations were similar among clades with different ecological attributes. Occupancy patterns were unimodal, with long and nearly symmetrical origination and extinction phases. A single origination pulse may be present at 2.0-1.75 Ma, but besides this, there is no evidence that evolutionary or ecological changes in the eastern African record tracked rapid, 100,000-y-scale changes in global climate. Rather, eastern African large mammal evolution tracked global or regional climatic trends at long (million year) time scales, while local, basin-scale changes (e.g., tectonic or hydrographic) and biotic interactions ruled at shorter timescales.
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As described in Part 1 & 2 (Private Game Issue 1 & 2), Earth’s path follows a sequence of natural oscillations effecting global dynamics of constant vegetation and habitat change. Consequently, phyllo-biogeographic existence and evolution (distribution, performance, genetic integrity, survival, and speciation trades) of animal species are affected. Animals move, adapt, or die as a result of changing environmental conditions. The conservation applications of palaeozoological data are numerous (Lyman 2006). One such application concerns the determination whether or not a species is native to a region. Such insight is essential to informing species translocations and to restoring biodiversity in disturbed habitats.
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The only antelopine species found today in southern Africa is the springbok, Antidorcas marsupialis. However, a close relative, A. bondi, was abundant and widespread during the Late Pleistocene. This animal died out at the beginning of the Holocene, ca. 7000 years ago. In a recent study of Florisbad fossil mammals, it was proposed that the extinct springbok was an exclusive grazer on the basis of its specialized dental features, in contrast to the modern springbok, which is a mixed feeder. We provide evidence in support of this hypothesis from stable carbon isotopic analysis of fossil and modern sprinbok teeth. The results are also in accordance with the interpretation that A. bondi coexisted with a wide range of larger-bodied grazing ungulates in a system similar to the grazing succession described for the Serengeti in East Africa.
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Members of the African Bovidae exhibit dietary resource partitioning, which presumably allows coexistence of many species with herbivorous diets. Levels of resource partitioning based on diet include primary food preference, habitat preference, and feeding-height preference. Morphological correlates of these levels of resource partitioning were sought in the skull and vertebral column of 33 bovid species. A quantitative morphometric study of the mandible, skull, and thoracic vertebrae in a large sample of bovids (n > 700) explores these correlates using video-image analysis. Results of this study indicate that many significant morphological differences exist among bovids that have a diet of either grass, dicots, or some combination of these two primary resources. In addition, some variables significantly distinguished bovids with different habitat and feeding-height preferences. These morphological correlates crossed taxonomic boundaries and, in many cases, were related to the structural properties of the herbivorous diet.
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This paper presents an overview of paleoenvironmental changes in East Africa during the late Quaternary based on evidence from pollen, diatoms, microscopic charcoal, and lake level records and associated proxies. The paleoenvironmental records derived from different proxies complement each other to provide a more accurate and complete assessment of the paleoenvironmental changes in East Africa. The records show that the period prior to c. 42,000 14 C yr BP was characterized by warm climatic conditions similar to the present. This was followed by a change to cold dry conditions from 42,000 to 30,000 14 C yr BP , and cold and moist conditions from 30,000 to 21,000 14 C yr BP . Temperatures during the latter period leading to the Last Glacial Maximum (LGM) were probably 2 to 4.1°C lower than the present. Between c. 21,000 and 12,500 14 C yr BP East Africa's environment was generally cool, punctuated by two significant episodes of prolonged desiccation. Warm and moist conditions punctuated by rapid climatic changes prevailed in the region during the deglacial and middle Holocene period. Ice core records document two significant and abrupt drought events in the region, one at ~8300 14 C yr BP and the other at 5200 14 C yr BP . The onset of a longer and more extensive desiccation period commencing ~4000 14 C yr BP was registered in nearly all sites. The climate of East Africa was generally drier than present during the Medieval Warm Period (MWP) while fairly wet conditions prevailed during the Little Ice Age (LIA) interrupted by three episodes of aridity, more severe than those of more recent times. Whereas this review advances our understanding of climate and vegetational changes in East Africa beyond the Last Glacial Maximum, it also highlights limitations of the paradigms that explain the forcing mechanisms behind the changes. However, unequivocal interpretation of the multiproxy data from East Africa with respect to paleoenvironmental changes becomes extremely complex and challenging especially when the anthropogenic input is considered.
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A comprehensive, but simple-to-use software package for executing a range of standard numerical analysis and operations used in quantitative paleontology has been developed. The program, called PAST (PAleontological STatistics), runs on standard Windows computers and is available free of charge. PAST integrates spreadsheettype data entry with univariate and multivariate statistics, curve fitting, time-series analysis, data plotting, and simple phylogenetic analysis. Many of the functions are specific to paleontology and ecology, and these functions are not found in standard, more extensive, statistical packages. PAST also includes fourteen case studies (data files and exercises) illustrating use of the program for paleontological problems, making it a complete educational package for courses in quantitative methods.
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The bovid fossils from Elandsfontein, south-western Cape Province, South Africa, comprise 7257 individually numbered specimens from 18 species. Taxonomic comparisons with Olduvai Gorge and other African sites and the high percentage of extinct forms imply that the bones accumulated in the earlier part of the Middle Quaternary, probably sometime between 700,000 and 400,000 years ago. By extension, this is also the most likely age for the skull cap of archaicHomo sapiens (‘Saldanha Man’) and for the occasional ‘late’ Acheulean stone artifacts that accompany the animal bones. In keeping with geomorphological observations and other aspects of the fauna, the bovids indicate a relatively grassy and moist environment, apparently during an interglaciation that differed significantly from the Holocene. Geomorphological context, the frequent occurrence of partial skeletons, bone damage, and skeletal part representation suggest that carnivore feeding on carcasses scattered across a Mid-Quaternary land surface was probably the main factor shaping the Elandsfontein bone assemblage. Porcupines may also have played a role, but there is little evidence for human activity. The Elandsfontein assemblage thus provides a useful ‘control’ for comparison with bone accumulations where context, associations, and bone damage indicate that people were heavily involved. For example, there are very few young animals in the otherwise attritional profile of ‘giant’ buffalo from Elandsfontein, probably because carnivores often rapidly and completely consumed young carcasses. This suggests that few young carcasses would be available for human scavenging and thus that archaeological attritional profiles in which young individuals are common probably reflect active human hunting, at least of young animals.
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The environmental history of montane southwest Uganda over the last c. 12 300 years is described using data from a 22.84-m long core of sediment from Ahakagyezi Swamp, Rukiga Highlands. These data include plant microfossils and macrofossils, charcoal and nine radiocarbon dates. According to evidence from plant fossils and radiocarbon dates, vegetation in the Ahakagyezi catchment was very different to the expected climax today until c. 10500 BP. The surface of the swamp was dominated by sedges, whilst the adjacent hillsides were largely devoid of trees. Changes in pollen suggest that a form of dry montane scrub was replaced by moist, lower montane forest from c. 11000BP. Subsequent fluctuations in pollen indicate that, once established, the composition of moist, lower montane forest varied during the Holocene and that unreversed forest reduction commenced in the Ahakagyezi catchment at c. 800 BP. On the surface of the swamp itself Syzygium-dominated swamp forest increased in extent from c. 5800 BP, and more rapidly from c. 3100 BP, before beginning a decline c. 1300 BP. The most likely reason for the presence of montane scrub in the catchment, rather than moist lower montane forest, prior to c. 11000 BP (i.e., a period that is believed to precede agricultural activity in the region) is a climate somewhat drier than the present. The spread of moist lower montane forest is believed to indicate the onset of climatic conditions broadly similar to those of today (i.e., the beginning of the Holocene). The causes of Holocene changes in forest composition and extent are most probably due to one or a combination of the following factors: natural succession; regional climate change; and the impact of human activity.
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We provide stable carbon isotope data from 37 species of African bovids to document dietary preferences for C3 browse (or fruits) or C4 grass. These data provide a quantitative measure of the fraction of C4 grass in bovid diets, can be applied on regional to local scales, can be derived from tooth enamel and hair or other tissues, and permit the diets of bovids to be considered in the context of a grazer - browser continuum. We recognize hypergrazers (>95% C4 grass), grazers (70-95% C4 grass), mixed feeders (>30% C4 grass and >30% C3 browse), browsers (70-95% C3 browse), and hyperbrowsers or frugivores (>95% C3 browse or fruit). Our results suggest that, of the extant East African Bovidae, impala (Aepyceros melampus), Thomson's gazelle (Gazella thomsonii), and oribi (Ourebia ourebi) can be construed as mixed feeders. Dietary estimates based on stable isotope analysis are in broad agreement with other measures of diet such as hypsodonty index, mass relationships, and wear scratches on enamel.
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The African long-horned buffalo, Pelorovis antiquus, was once widespread in the savannas and grasslands of southern, eastern, and northern Africa. It apparently disappeared from southern and eastern Africa about 12,000 years ago and from northern Africa about 4000 years ago. Its extinction has been variously attributed to human predation, climatic change, or some combination of the two. Recently, Peters et al. (Late Quaternary extinction of ungulates in Sub-Saharan Africa: a reductionist's approach, Journal of Archaeological Science 21, 17-28, 1994) argued that its demise has been exaggerated and that its postcranial anatomy indicates it was simply a long-horned morph of the extant African buffalo, Syncerus caffer. Both cranial and postcranial similarities to Syncerus can be used to suggest that P. antiquus should be removed from Pelorovis and reassigned to Syncerus, as Syncerus antiquus. However, its status as a distinct (and now extinct) buffalo species is demonstrated by its singular horns, by some dental differences from S. caffer, and above all, by its geographic overlap with S. caffer through much of the middle and late Quaternary, with no evidence for intermediate forms.
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The analysis of fossil ungulate cheek teeth has long been one of the main sources of information about the terrestrial environments of the Cenozoic, but the methods used to extract this information have been either imprecise or prohibitively laborious. Here we present a method based on relative facet development that is quantitative, robust, and rapid. This method, which we term mesowear analysis, is based on the physical properties of ungulate foods as reflected in the relative amounts of attritive and abrasive wear that they cause on the dental enamel of the occlusal surfaces. Mesowear was recorded by examining the buccal apices of molar tooth cusps. Apices were characterized as sharp, rounded, or blunt, and the valleys between them either high or low. The method has been developed only for selenodont and trilophodont molars, but the principle is readily extendable to other crown types. Mesowear analysis is insensitive to wear stage as long as the very early and very late stages are excluded. Cluster analysis of the mesowear variables produces clusters reflecting four main groups from abrasion-dominated to attrition-dominated: grazers, graze-dominated mixed feeders, browse-dominated mixed feeders, and browsers. Most of the relatively few apparent anomalies are explained by more detailed dietary information. Mesowear analysis provides resolution within the main dietary classes and the clustering is virtually identical with and without the index of hypsodonty. Discriminant analysis using all mesowear variables and hypsodonty
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THERE has been considerable debate about the magnitude of the decrease in temperature1,2 and the change in precipitation3 in the African tropics during the last glacial period. With the advent of fossil pollen studies in equatorial regions, it is now generally agreed that the temperature did decrease at this time in tropical regions4, but the magnitude of the temperature fluctuations and discrepancies between the continental5 and marine6 temperature records have yet to be resolved7. Here we present new quantitative estimates of temperature and precipitation using a multivariate analysis8 of pollen time-series data from peat deposits in Burundi for the past 40,000 years9. For the last glacial period, our estimate of a temperature decrease of 4 +/- 2 °C is less than those (ranging from 5 to 8 °C) derived from snow-line and tree-line records5,7. Model simulations7 indicate that the snow-line and tree-line estimates (from high-elevation sites at ~4,000 m above sea level) are incompatible with the marine temperature record. Our lower estimate from a site of intermediate elevation may help resolve the differences between these records. We also estimate that the mean annual rainfall decreased by 30% during the last glacial period, in agreement with the rainfall history inferred from lake level fluctuations10.
Article
The palaeodiet of seven bovids from Makapansgat Limeworks Cave are analysed using dental mesowear. Results suggest that Tragelaphus pricei had a highly attritional diet and was thus a browser. Tragelaphus sp. aff. T. angasii and Aepyceros sp. were also browsers, having diets similar in texture to the extant mule deer (Odocoileus hemionus). Gazella vanhoepeni had an intermediate attrition-abrasion wear signal and groups most closely with extant mixed feeders. Redunca darti and Makapania broomi are at the abrasion end of the wear continuum and cluster with living grazers, such as the hippotragines and reduncines. Parmularius braini had a highly abrasive diet similar to extreme grazers like the American bison (Bison bison) and topi (Damaliscus lunatus). The bovid mesowear data were compared to previous palaeodietary studies using taxonomic uniformitarianism, ecomorphology (hypsodonty), and stable carbon isotopes on the same Makapansgat taxa. This comparison showed that the mesowear results are most closely in-line with the isotope data, both of which are non-genetic signals that reflect diet during an extended portion of an animal's life.
Article
On a study visit to the Bayerische Staatssammlung fur Palaontologie und historische Geologie, Munich, in the summer of 1992, it was discovered that the holotypes of Megalotragus kattwinkeli and Rhynotragus semiticus and of Taurotragus oryx pachyceros from Olduvai Gorge, had escaped destruction by Allied bombing during WWII, contrary to previous belief. On a further visit in 1994, a discovery was made about the refound holotype of Rhynotragus semiticus which corroborates its synonymy with Megalotragus kattwinkeli. It is now known that the name Rhynotragus semiticus is available from 1925, and not 1935 as thought previously, and it therefore becomes the senior synonym for the commonly used Megalotragus kattwinkeli. The consequences of these discoveries for taxonomy and nomenclature are discussed. -from Authors
Article
Bovidae comprises 17 different genera, including extinct and living representatives. Bovid fossils identified by horn cores first appeared in the early Miocene of Eurasia and Africa, but the family probably originated in the Oligocene. This chapter discusses the bovid fossils from the Daka Member, which include more than 25 well-preserved crania and hundreds of dental and horn core specimens. The assemblage is especially rich in bovines, alcelaphines, and reduncines, hippotragines, tragelaphines, antilopines, caprines, aepycerotines, neotragines, and possibly ovibovines. Bovids contribute substantial paleoecological information as a result of the tendency for niche specificity among bovid tribes, genera, and species. The chapter shows the conservative view of African bovid phylogeny and presents the entire Daka bovid assemblage as a unit. It also provides brief descriptions and evolutionary backgrounds for the bovid taxa present.
Article
This volume, the first in a series devoted to the paleoanthropological resources of the Middle Awash Valley of Ethiopia, studies Homo erectus, a close relative of Homo sapiens. Written by a team of highly regarded scholars, this book provides the first detailed descriptions, photographs, and analysis of the fossil vertebrates-from elephants and hyenas to humans-from the Daka Member of the Bouri Formation of the Afar, a place renowned for an abundant and lengthy record of human ancestors. These fossils contribute to our understanding human evolution, and the associated fauna provide new information about the distribution and variability of Pleistocene mammals in eastern Africa. The contributors are all active researchers who worked on the paleontology and geology of these unique deposits. Here they have combined their disparate efforts into a single volume, making the original research results accessible to both the specialist and the general reader. The volume synthesizes environmental backdrop and anatomical detail to open an unparalleled window on the African Pleistocene and its inhabitants.
Article
Alcelaphine antelopes comprise one of the most species-rich groups among the mammalian assemblages from the Middle Awash, Ethiopia, and in Africa as a whole. I describe a new genus and species Awashia suwai from Matabaietu 3, and other new alcelaphine species, Damaliscus ademassui from Gamedah 1 and Beatragus whitei from Matabaietu 3-5, all dated ca. 2.5 m.y. (millions of years). Other new alcelaphine fossils from Middle Awash include an Early Pliocene species allied to Damalops, Late Pliocene records of Parmularius cf. pandatus and Beatragus antiquus, and Middle Pleistocene records of Megalotragus kattwinkeli, P. angusticornis, Damaliscus niro, Connochaetes taurinus olduvaiensis, Numidocapra crassicornis, and Alcelaphus buselaphus. My comparisons of these fossils with all other known fossil and Recent Alcelaphini includes a cladistic analysis. The results suggest that during or before the Miocene-Pliocene transition two alcelaphine subtribes diverged for which I suggest the names Alcelaphina and Damaliscina. Alcelaphina consists of two ancient subclades: (1) the sister-group of Damalacra neanica and Beatragus known since 5.0-4.5 m.y. ago, and (2) a large clade first recorded 4.4 m.y. ago (genera Damalops, Numidocapra, Alcelaphus, Rabaticeras, Megalotragus, Oreonagor, and Connochaetes) that had a high diversification rate since 3 m.y. ago. The earliest record of Damaliscina is the form that Gentry (1980) named Damalacra acalla, which emerges as the hypothetical direct ancestor of the Early-Middle Pliocene split into Parmularius and the Damaliscus group. The placement of the new genus Awashia remains problematic. A new ovibovine genus and species, Nitidarcus asfawi, and a new caprine genus and species, Bouria anngettyae, both from Bouri 1, are also described. I discuss some evolutionary and biogeographic implications of the new fossils from Middle Awash.
Article
Bovidae contain the cattle, sheep, goats, and antelopes. The word “antelope” is used for bovids outside Europe, mostly in Africa, or not domesticated before Carl Linnaeus' lifetime. It does not correspond with a formal taxonomic category. Most phylogenies postulate bovids being closer to cervids than to giraffids. Unlike the cervoid Moschus in relation to Cervidae, there is no living hornless pecoran thought to be a bovoid (member of a superfamily Bovoidea including Bovidae and any related families, the latter as yet unknown). In Eurasia, tiny bovid-like dental remains are known well back to the early Oligocene of Mongolia, but nothing is known of pre-Miocene ruminants in Africa. Pecorans such as Walangania, Propalaeoryx, and Namibiomeryx do appear in the early Miocene, and the last has been claimed to be a bovid. Subfamilies of Bovidae include Hypsodontinae, Bovinae, Antilopinae, Reduncinae, Oiocerinae, Hippotraginae, and Caprinae. This chapter discusses the overall classification of Bovidae and their evolutionary relationships.
Article
This chapter considers three aspects of the evolution of the Serengeti–Mara ecosystem, focusing on the possible origins of a broadscale migratory system. Firstly, it examines the geoecology of the system, including the evolution of the eastern plain, the wet-season pasture of the wildebeest and zebra today, and the Mara River and Lake Victoria as potential drought refugia. Secondly, it sketches the evolution of large mammals, especially the larger herbivores and carnivores. Thirdly, it assesses the hominin factor, the use of the system by prehistoric hunter-gatherers and pastoralists, and the more dramatic historical human impacts.
Article
This paper describes the fluctuations of Lakes Victoria, Stefanie, Turkana and Naivasha over the last two centuries. A chronology of Lake Victoria back to 700 A.D. is also developed. These chronologies are based mainly on oral traditions of the local peoples, as described in various historical sources, and on reports of European visitors, settlers and explorers. In some cases actual historical levels have been reported. The historical fluctuations are meshed with the modern record to provide a picture of the fluctuations in lake levels until the late twentieth century. The chronologies for Victoria and Stefanie contain much new material, permitting higher temporal resolution and better quantitative assessments, as well as extension of chronologies to the beginning of the 19th century. For Lakes Turkana and Naivasha, chronologies published by other authors are expanded and compared with those for Victoria, Stefanie and other African lakes. A long term chronology for Lake Victoria is developed using the record of the summer Nile flow. These lakes show remarkably similar trends. The most important of these trends are low levels during the first half of the 19th century, very high stands in the last decades of the 19th century, and around the turn of the century a rapid fall to 20th century levels. The lakes returned to relatively high stands in the 1960s, but these generally ended in the 1970s.
Article
A re-examination of the postcranial equid material from the spring deposits of Florisbad and Vlakkraal shows that a form of ass lived in southern Africa during the late Quaternary. On the basis of preliminary comparisons with published reports on ass and half-ass osteology, the Florisbad and Vlakkraal fossil material appears to be more similar to E. (Asinus) asinus than to E. (Asinus) hemionus and is provisionally assigned to E. (Asinus) sp. The presence of a form of ass is significant as it extends the distribution of the subgenus Asinus into the southern African subregion, giving support to the suggested African origin of E. (asinus). Its presence also appears to confirm the hypothesis that during the Last Interglacial senu lato higher levels of grassland productivity allowed a facilitating grazing system which provided niches for specialized grazers that are absent from modern southern grasslands. -from Author
Article
Subsaharan African fossil Bovini belong to two major related lineages, the Simatherium-Pelorovis and Ugandax-Syncerus lineages. Makapansgat Members 3 and 4 contain Simatherium cf. kohllarseni while the bovine from Member 5 is Syncerus cf. acoelotus. The presence of Simatherium in Member 3 and 4 suggests that these assemblages may have accumulated when temperatures in the area, at least during winters, were particularly cold. The interesting taphonomic implications of the high percentage of juveniles (89%) of S. cf. kohllarseni in Member 3 are discussed. A preliminary revision of the Member 3 bovid checklist and a preliminary checklist for Member 5 bovids are given. The bovid evidence suggests a fundamental difference between the Pliocene Member 3 assemblage, with its preponderance of extinct genera and with Asian Siwaliks affinities, and the Pleistocene Member 5 assemblage. - Author Pliocene Siwaliks Pleistocene
Article
Documents the occurrence of the Florisian, or late Quaternary, form of the giant alcelaphine, Megalotragus priscus, from dongas on the Ongers River, near Britstown in the central Karoo. This is significant as it confirms the occurrence of the species in the Karoo and it suggests significantly wetter environments and productive grasslands in the central Karoo in pre-Holocene times. Aridification in recent times is the likely cause of changes in grassland quality and the local dissappearance of these animals, if not their extinction. -Authors
Article
Primary productivity and herbivory were studied in the Serengeti National Park, Tan- zania, and Masai Mara Game Reserve, Kenya, during the annual cycle of 1974-1975, and wet-dry season transitions in 1976-1979. Basic state variables measured were aboveground plant biomass inside permanent and temporary fences, and outside fences. Productivity was calculated as the sum of positive plant biomass increments. Control productivity (cPn) was calculated from biomass dy- namics inside permanent fences. Temporary fences were moved in concert with grazing by the region's abundant ungulates to estimate actual aboveground primary productivity (aPn).Primary productivity was highly stochastic with productive periods poorly synchronized even among nearby sites. Short- term productivities could be extremely high, exceeding 30 g.m-2 .dl. Grazing animals adjusted their densities in relation to grassland productivity. The average proportion of annual aPn that was consumed by herbivores was 0.66, with a minimum of 0.15 and a maximum of 0.94. Green forage was available everywhere late In the wet season in May but was available only at high rainfall sites in the northwest late in the dry season in November. By the end of the dry season, the residual plant biomass outside fences averaged only 8% of cPn. Nomadic grazers moved seasonally in response to grassland pro- ductivity. The growing season ranged from 76 d in low rainfall areas to virtually continuous in high rainfall areas. Annual cPn was linearly related to rainfall and averaged 357 g.m-2.yr-1 over the year and 1.89 g.m-2,d-1 during the growing season. Actual aPn was substantially greater than cPn at most sites, averaging 664 g.m-2.yr-1. Growing season aPn averaged 3.78 g.m-2.d-1. Grazing stimulated net primary productivity at most locations, with the maximum stimulation at intermediate grazing in- tensities. Stimulation was dependent upon soil moisture status at the time of grazing. Rain had a diminishing effect on primary productivity as the wet season progressed and plant biomass accu- mulated. Part of the stimulation of grassland productivity by grazing was due to maintenance of the vegetation in an immature, rapidly growing state similar to that at the beginning of the rainy season. Slnce grazers overrode rainfall-determined productivity patterns, aPn was more closely related to grazing intensity than to ranfall. Grazing was heavier on grasslands that were intrinsically more productive. Rate of energy flow per unit of plant biomass was much higher in grazed vegetation. Grazers ate green leaves almost exclusively during the wet season, but species composition of the diets of different grazers differed markedly. Diets of nomadic grazers were very different in the wet and dry seasons. Vegetation dried out rapidly at the onset of the dry season and dry plant tissues made up a substantial proportion of ungulate dry season diets. However, green forage commonly was more abundant in diets than in the vegetation. Grazing increased both forage quality and its rate of pro- duction. Zebras supplemented a high-bulk diet by eating the seeds of awnless grasses. The foraging patterns of different grazers were differentiated by several vegetation properties, including productivity, structure, and species composition, in a manner suggesting resource partitioning. The relationship between the stability of vegetation functional properties and community species diversity was posltive in five of seven tests. Greater species diversity was associated with greater biomass stability through the seasons, greater resistance to grazing by a single species of ungulate in both the wet and dry seasons, and greater resilience after grazing. Species diversity was not associated with greater resistance to grazing by several ungulate species or to plant species extinction. Specific properties of trophic web members were identified that produced greater functional stability in more diverse communities. Fire does not appear to have important effects upon the functional properties of the grasslands except for a weak stimulation of productivity in the wet season immediately following dry season burning. Fire did have an important effect upon structural properties of the vegetation that would tend to regulate ungulate feeding. The ecology of neither the plants nor the animals in the Serengeti ecosystem can be understood in isolation; many traits of both suggest coevolution among trophic web members. The functional dynamics of the trophic web suggest that the acceleration of energy and nutrient flow rates due to intense herbivory has resulted in the development of an entire consumer food web due to additive fluxes rather than mere quasi-parasitic fluxes from plants to animals.
Article
The dietary regimes of 15 ungulate species from the middle Pleistocene levels of the hominid-bearing locality of Elandsfontein, South Africa, are investigated using the mesowear technique. Previous studies, using taxonomic analogy, classified twelve of the studied species as grazers (Redunca arundinum, Hippotragus gigas, Hippotragus leucophaeus, Antidorcas recki, Homoiceras antiquus, Damaliscus aff. lunatus, Connochaetes gnou laticornutus, Rabaticerus arambourgi, Damaliscus niro, Damaliscus sp. nov., an unnamed “spiral horn” antelope and Equus capensis), one as a mixed feeder (Taurotragus oryx) and two as browsers (Tragelaphus strepsiceros and Raphicerus melanotis). Although results from mesowear analysis sustain previous dietary classifications in the majority of cases, five species were reclassified. Three species previously classified as grazers, were reclassified as mixed feeders (H. gigas, D. aff. lunatus and R. arambourgi), one previously classified as a grazer, was reclassified as a browser (the “spiral horn” antelope), and one previously classified as a mixed feeder, was reclassified as a browser (T. oryx). While current results broadly support previous reconstructions of the Elandsfontein middle Pleistocene environment as one which included a substantial C3 grassy component, the reclassifications suggest that trees, broad-leaved bush and fynbos were probably more prominent than what was previously thought.