New Oldowan locality Sare-Abururu (ca. 1.7 Ma) provides evidence of diverse hominin behaviors on the Homa Peninsula, Kenya.

The Homa Peninsula, in southwestern Kenya, continues to yield insights into Oldowan hominin landscape behaviors. The Late Pliocene locality of Nyayanga (∼3-2.6 Ma) preserves some of the oldest Oldowan tools. At the Early Pleistocene locality of Kanjera South (∼2 Ma) toolmakers procured a diversity of raw materials from over 10 km away and strategically reduced them in a grassland-dominated ecosystem. Here, we report findings from Sare-Abururu, a younger (∼1.7 Ma) Oldowan locality approximately 12 km southeast of Kanjera South and 18 km east of Nyayanga. Sare-Abururu has yielded 1754 artifacts in relatively undisturbed low-energy silts and sands. Stable isotopic analysis of pedogenic carbonates suggests that hominin activities were carried out in a grassland-dominated setting with similar vegetation structure as documented at Kanjera South. The composition of a nearby paleo-conglomerate indicates that high-quality stone raw materials were locally abundant. Toolmakers at Sare-Abururu produced angular fragments from quartz pebbles, representing a considerable contrast to the strategies used to reduce high quality raw materials at Kanjera South. Although lithic reduction at Sare-Abururu was technologically simple, toolmakers proficiently produced cutting edges, made few mistakes and exhibited a mastery of platform management, demonstrating that expedient technical strategies do not necessarily indicate a lack of skill or suitable raw materials. Lithic procurement and reduction patterns on the Homa Peninsula appear to reflect variation in local resource contexts rather than large-scale evolutionary changes in mobility, energy budget, or toolmaker cognition.

Expanded geographic distribution and dietary strategies of the earliest Oldowan hominins and Paranthropus.

The oldest Oldowan tool sites, from around 2.6 million years ago, have previously been confined to Ethiopia's Afar Triangle. We describe sites at Nyayanga, Kenya, dated to 3.032 to 2.581 million years ago and expand this distribution by over 1300 kilometers. Furthermore, we found two hippopotamid butchery sites associated with mosaic vegetation and a C4 grazer-dominated fauna. Tool flaking proficiency was comparable with that of younger Oldowan assemblages, but pounding activities were more common. Tool use-wear and bone damage indicate plant and animal tissue processing. Paranthropus sp. teeth, the first from southwestern Kenya, possessed carbon isotopic values indicative of a diet rich in C4 foods. We argue that the earliest Oldowan was more widespread than previously known, used to process diverse foods including megafauna, and associated with Paranthropus from its onset.

Ecological perspectives on technological diversity at Kanjera South.

The aspects of hominin behavior responsible for Oldowan stone tool variation are the focus of much debate. There is some consensus that this variation arises from a combination of ecological and cultural factors. The diversity of raw material types and technological strategies present at Kanjera South, Kenya, provide an opportunity to examine the interacting influences of ecology and culture on Oldowan stone tool variation. Here, we combine previous analyses of raw material properties, provenance, and technology with quantitative measures of core reduction intensity and tool utilization to examine the influence of both ecological and technocultural factors on stone tool variation at Kanjera South. The results of this analysis reflect a dynamic relationship between raw material properties, provenance, and hominin mobility. Exotic raw materials are generally more resistant to edge attrition compared with those available locally, which may have incentivized their transport over long distances and more extensive reduction. Cores produced on raw materials from distant sources also exhibit more complex core reduction strategies than locally acquired materials. While this pattern is partially due to the differences in the quality of knappable stone, bifacial centripetal and multifacial core reduction strategies also arise due to the continuous transport and use of exotic raw materials. Moreover, the variation in stone tool reduction is not consistent with neutral models of stone tool transport and discard. These results demonstrate that ecological factors such as raw material provenance and physical properties have strong impacts on reduction intensity and the technological strategies used by hominins.

Bovid mortality patterns from Kanjera South, Homa Peninsula, Kenya and FLK-Zinj, Olduvai Gorge, Tanzania: Evidence for habitat mediated variability in Oldowan hominin hunting and scavenging behavior.

The archaeological record has documented Oldowan hominin occupation of habitats ranging from open grasslands to riparian forest by 2.0 Ma. Despite this we have a poor understanding of whether hominin foraging behavior varied in different environmental settings. We compared bovid mortality profiles from the two largest Oldowan zooarchaeological samples, one from a grassland (Excavation 1, Kanjera South, Kenya) and another from a woodland (FLK Zinj, Olduvai Gorge, Tanzania) with bovid mortality samples created by African carnivores in different habitats. Kanjera hominins frequently had early access, likely through hunting, to small (size 1 ≤ 23 kg and size 2 = 24-112 kg) juvenile bovids, creating a mortality pattern similar to that created by grassland dwelling carnivores. Kanjera hominins had more mixed access to large (size 3 = 113-340 kg), often juvenile, bovids and frequently scavenged heads. In contrast, previous work has shown that the few small bovids at FLK-Zinj were predominantly older individuals. Prime adults dominated the FLK-Zinj large bovid sample, leading to a mortality pattern similar to that created by carnivores occupying more closed habitats. Variation in bovid body size and mortality profiles between these archaeological assemblages may reflect the challenges of acquiring fauna in open versus closed habitats with a simple hunting toolkit. The heterogeneous woodland habitat of FLK-Zinj would have provided more opportunities to ambush prey, whereas on grasslands with more limited concealment opportunities Kanjera hominins focused their efforts on vulnerable juvenile prey, some likely acquired after short chases.

Ecomorphological analysis of bovid mandibles from Laetoli Tanzania using 3D geometric morphometrics: Implications for hominin paleoenvironmental reconstruction.

The current study describes a new method of mandibular ecological morphology (ecomorphology). Three-dimensional geometric morphometrics (3D GM) was used to quantify mandibular shape variation between extant bovids with different feeding preferences. Landmark data were subjected to generalized Procrustes analysis (GPA), principal components analysis (PCA), and discriminant function analysis (DFA). The PCA resulted in a continuum from grazers to browsers along PC1 and DFA classified 88% or more of the modern specimens to the correct feeding category. The protocol was reduced to a subset of landmarks on the mandibular corpus in order to make it applicable to incomplete fossils. The reduced landmark set resulted in greater overlap between feeding categories but maintained the same continuum as the complete landmark model. The DFA resubstitution and jackknife analyses resulted in classification success rates of 85% and 80%, respectively. The reduced landmark model was applied to fossil mandibles from the Upper Laetolil Beds (∼4.3-3.5 Ma) and Upper Ndolanya Beds (∼2.7-2.6 Ma) at Laetoli, Tanzania in order to assess antelope diet, and indirectly evaluate paleo-vegetation structure. The majority of the fossils were classified by the DFA as browsers or mixed feeders preferring browse. Our results indicate a continuous presence of wooded habitats and are congruent with recent environmental studies at Laetoli indicating a mosaic woodland-bushland-grassland savanna ecosystem.

Unexpected Convergent Evolution of Nasal Domes between Pleistocene Bovids and Cretaceous Hadrosaur Dinosaurs.

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.

Bovid ecomorphology and hominin paleoenvironments of the Shungura Formation, lower Omo River Valley, Ethiopia.

The Shungura Formation in the lower Omo River Valley, southern Ethiopia, has yielded an important paleontological and archeological record from the Pliocene and Pleistocene of eastern Africa. Fossils are common throughout the sequence and provide evidence of paleoenvironments and environmental change through time. This study developed discriminant function ecomorphology models that linked astragalus morphology to broadly defined habitat categories (open, light cover, heavy cover, forest, and wetlands) using modern bovids of known ecology. These models used seven variables suitable for use on fragmentary fossils and had overall classification success rates of >82%. Four hundred and one fossils were analyzed from Shungura Formation members B through G (3.4-1.9 million years ago). Analysis by member documented the full range of ecomorph categories, demonstrating that a wide range of habitats existed along the axis of the paleo-Omo River. Heavy cover ecomorphs, reflecting habitats such as woodland and heavy bushland, were the most common in the fossil sample. The trend of increasing open cover habitats from Members C through F suggested by other paleoenvironmental proxies was documented by the increase in open habitat ecomorphs during this interval. However, finer grained analysis demonstrated considerable variability in ecomorph frequencies over time, suggesting that substantial short-term variability is masked when grouping samples by member. The hominin genera Australopithecus, Homo, and Paranthropus are associated with a range of ecomorphs, indicating that all three genera were living in temporally variable and heterogeneous landscapes. Australopithecus finds were predominantly associated with lower frequencies of open habitat ecomorphs, and high frequencies of heavy cover ecomorphs, perhaps indicating a more woodland focus for this genus.

Old stones' song: use-wear experiments and analysis of the Oldowan quartz and quartzite assemblage from Kanjera South (Kenya).

Evidence of Oldowan tools by ∼2.6 million years ago (Ma) may signal a major adaptive shift in hominin evolution. While tool-dependent butchery of large mammals was important by at least 2.0 Ma, the use of artifacts for tasks other than faunal processing has been difficult to diagnose. Here we report on use-wear analysis of ∼2.0 Ma quartz and quartzite artifacts from Kanjera South, Kenya. A use-wear framework that links processing of specific materials and tool motions to their resultant use-wear patterns was developed. A blind test was then carried out to assess and improve the efficacy of this experimental use-wear framework, which was then applied to the analysis of 62 Oldowan artifacts from Kanjera South. Use-wear on a total of 23 artifact edges was attributed to the processing of specific materials. Use-wear on seven edges (30%) was attributed to animal tissue processing, corroborating zooarchaeological evidence for butchery at the site. Use-wear on 16 edges (70%) was attributed to the processing of plant tissues, including wood, grit-covered plant tissues that we interpret as underground storage organs (USOs), and stems of grass or sedges. These results expand our knowledge of the suite of behaviours carried out in the vicinity of Kanjera South to include the processing of materials that would be 'invisible' using standard archaeological methods. Wood cutting and scraping may represent the production and/or maintenance of wooden tools. Use-wear related to USO processing extends the archaeological evidence for hominin acquisition and consumption of this resource by over 1.5 Ma. Cutting of grasses, sedges or reeds may be related to a subsistence task (e.g., grass seed harvesting, cutting out papyrus culm for consumption) and/or a non-subsistence related task (e.g., production of 'twine,' simple carrying devices, or bedding). These results highlight the adaptive significance of lithic technology for hominins at Kanjera.

Earliest archaeological evidence of persistent hominin carnivory.

The emergence of lithic technology by ≈ 2.6 million years ago (Ma) is often interpreted as a correlate of increasingly recurrent hominin acquisition and consumption of animal remains. Associated faunal evidence, however, is poorly preserved prior to ≈ 1.8 Ma, limiting our understanding of early archaeological (Oldowan) hominin carnivory. Here, we detail three large well-preserved zooarchaeological assemblages from Kanjera South, Kenya. The assemblages date to 2.0 Ma, pre-dating all previously published archaeofaunas of appreciable size. At Kanjera, there is clear evidence that Oldowan hominins acquired and processed numerous, relatively complete, small ungulate carcasses. Moreover, they had at least occasional access to the fleshed remains of larger, wildebeest-sized animals. The overall record of hominin activities is consistent through the stratified sequence - spanning hundreds to thousands of years - and provides the earliest archaeological evidence of sustained hominin involvement with fleshed animal remains (i.e., persistent carnivory), a foraging adaptation central to many models of hominin evolution.

Oldest evidence of tool making hominins in a grassland-dominated ecosystem.

BACKGROUND: Major biological and cultural innovations in late Pliocene hominin evolution are frequently linked to the spread or fluctuating presence of C(4) grass in African ecosystems. Whereas the deep sea record of global climatic change provides indirect evidence for an increase in C(4) vegetation with a shift towards a cooler, drier and more variable global climatic regime beginning approximately 3 million years ago (Ma), evidence for grassland-dominated ecosystems in continental Africa and hominin activities within such ecosystems have been lacking. METHODOLOGY/PRINCIPAL FINDINGS: We report stable isotopic analyses of pedogenic carbonates and ungulate enamel, as well as faunal data from approximately 2.0 Ma archeological occurrences at Kanjera South, Kenya. These document repeated hominin activities within a grassland-dominated ecosystem. CONCLUSIONS/SIGNIFICANCE: These data demonstrate what hitherto had been speculated based on indirect evidence: that grassland-dominated ecosystems did in fact exist during the Plio-Pleistocene, and that early Homo was active in open settings. Comparison with other Oldowan occurrences indicates that by 2.0 Ma hominins, almost certainly of the genus Homo, used a broad spectrum of habitats in East Africa, from open grassland to riparian forest. This strongly contrasts with the habitat usage of Australopithecus, and may signal an important shift in hominin landscape usage.