Olduvai's oldest Oldowan.

Previously, Olduvai Bed I excavations revealed Oldowan assemblages <1.85 Ma, mainly in the eastern gorge. New western gorge excavations locate a much older ∼2.0 Ma assemblage between the Coarse Feldspar Crystal Tuff (∼2.015 Ma) and Tuff IA (∼1.98 Ma) of Lower Bed I, predating the oldest eastern gorge DK assemblage below Tuff IB by ∼150 kyr. We characterize this newly discovered fossil and artifact assemblage, adding information on landscape and hominin resource use during the ∼2.3-2.0 Ma period, scarce in Oldowan sites. Assemblage lithics and bones, lithofacies boundaries, and phytolith samples were surveyed and mapped. Sedimentological facies analysis, tephrostratigraphic and sequence stratigraphic principles were applied to reconstruct paleoenvironments and sedimentary processes of sandy claystone (lake), sandstone (fluvial), and sandy diamictite (debris flow) as principal lithofacies. Artifacts, sized, weighed, categorized, were examined for petrography, retouch, and flake scar size. Taxonomic classifications and taphonomic descriptions of faunal remains were made, and phytoliths were categorized based on reference collections. Lithics are dominantly quartzite, mainly debitage and less frequently simple cores, retouched pieces, and percussors. Well-rounded spheroids and retouched flakes are rare. Identifiable taxa, Ceratotherium cf. simum (white rhinoceros) and Equus cf. oldowayensis (extinct zebra), accord with nearby open savanna grasslands, inferred from C3 grass, mixed and/or alternating with C4 grass-dominated phytolith assemblages. Palms, sedges, and dicots were also identified from phytoliths. Diatoms and sponge spicules imply nearby freshwater. The assemblage accumulated at the toe of a Ngorongoro Volcano-sourced fan-delta apron of stacked debris flows, fluvials, and tuffs, preserving fossil tree stumps and wooded grassland phytoliths farther upfan. It formed after the climax of Ngorongoro volcanic activity during a Paleolake Olduvai lowstand and was then buried and preserved by lacustrine clays, marking the first of two lake transgressions, signifying wetter climates. Orbital precessional lake cycles were superposed upon multimillennial (∼4.9 kyr) lake fluctuations.

A new high-resolution 3-D quantitative method for identifying bone surface modifications with implications for the Early Stone Age archaeological record.

Bone surface modifications have become important indicators of hominin behavior and ecology at prehistoric archaeological sites. However, the method by which we identify and interpret these marks remains largely unchanged despite decades of research, relying on qualitative criteria and lacking standardization between analysts. Recently, zooarchaeologists have begun using new technologies capable of capturing 3-D data from bone surface modifications to advance our knowledge of these informative traces. However, an important step in this research has been overlooked and after years of work, we lack both a universal and replicable protocol and an understanding of the precision of these techniques. Here we propose a new standard for identifying bone surface modifications using high-resolution 3-D data and offer a systematic and replicable approach for researchers to follow. Data were collected with a white-light non-contact confocal profilometer and analyzed with Digital Surf's Mountains® software. Our data show that when methods are standardized, results between researchers are statistically indistinguishable. Multivariate analyses using the measured parameters allow discrimination between stone tool cut marks and mammalian carnivore tooth marks with 97.5% accuracy. Application of this method to fossil specimens resulted in 100% correspondence with identifications made by an experienced analyst using macroscopic observations of qualitative features of bone surface modifications. High-resolution 3-D analyses of bone surface modifications have great potential to improve the reliability and accuracy of taphonomic research, but only if our methods are replicable and precise.

OH-65: The earliest evidence for right-handedness in the fossil record.

Labial striations on the anterior teeth have been documented in numerous European pre-Neandertal and Neandertal fossils and serve as evidence for handedness. OH-65, dated at 1.8 mya, shows a concentration of oblique striations on, especially, the left I1 and right I1, I2 and C1, which signal that it was right-handed. From these patterns we contend that OH-65 was habitually using the right hand, over the left, in manipulating objects during some kind of oral processing. In living humans right-handedness is generally correlated with brain lateralization, although the strength of the association is questioned by some. We propose that as more specimens are found, right-handedness, as seen in living Homo, will most probably be typical of these early hominins.

Environments and hominin activities across the FLK Peninsula during Zinjanthropus times (1.84 Ma), Olduvai Gorge, Tanzania.

We establish through 13 excavations the landscape context and nature of hominin activities across the Zinjanthropus land surface from which the Leakeys recovered the FLK 22 and FLK NN 1 paleoanthropological assemblages. The land surface was created by fluvial incision of the eastern margin of paleo-Lake Olduvai following a major lake withdrawal. Erosion was uneven, leaving a peninsula bounded by a river channel, the FLK Fault, and a freshwater wetland. This FLK Peninsula supported groves of trees that attracted hominins and carnivores, and that preserved the dense concentrations of carcass remains and stone tools they left behind, including those at FLK 22. Some carcasses appear to have been acquired at the ecotone of the Peninsula and Wetland, where another dense artifact and bone assemblage accumulated. A lesser topographic high at the edge of a Typha marsh in the Wetland was the site of FLK NN 1 and a scatter of large stone tools used possibly for rootstock processing. Our landscape reconstruction delimits the vegetation mosaic indicated by previous work and provides a topographical explanation for the existence of FLK 22 and FLK NN 1. Both are unexpected if the FLK area was the flat, featureless lake margin terrain typical of lake basins similar to paleo-Olduvai. The results show that the Leakeys' sites were not isolated occupation floors but rather parts of a land surface utilized intensively by hominins. Although commonly considered to have been home bases, their likely high predation risk, evidenced by large carnivore feeding traces and the remains of four hominin individuals, suggests visits to them were brief and limited to feeding. Finally, stratigraphic observations confirm that FLK NN 3 accumulated on an older land surface, refuting the hypothesis that the OH 8 foot found there is the same individual as the OH 35 leg from FLK 22.

Crocodylian and mammalian carnivore feeding traces on hominid fossils from FLK 22 and FLK NN 3, Plio-Pleistocene, Olduvai Gorge, Tanzania.

Taphonomic analysis of the Olduvai Hominid (OH) 8 left foot from FLK NN Level 3 and the OH 35 left leg from FLK Level 22 (Zinjanthropus level) in Middle Bed I, Olduvai Gorge, indicates that both were fed upon by crocodiles. Both bear extensive tooth marking, including bisected tooth marks diagnostic of crocodylian feeding. The location of the bisected tooth marks on the distal tibia and the talus indicates disarticulation of the foot by crocodiles. The broken proximal ends of the tibia and fibula are more typical of feeding by a leopard-like carnivore, as is damage to the OH 7 mandible and parietals that are associated with and may derive from the same individual as OH 8. Previous work showing a close articulation of the foot and the leg has been used to suggest that the two specimens belong to the same individual despite deriving from sites separated by 200 m and slightly different stratigraphic levels according to previous work. The location and agent of tooth marking and the nature of gross damage do not refute this hypothesis, but the punctures on the talus and distal tibia differ in size and sharpness. Recent work shows that the stratigraphic discrepancy between OH 8 and OH 35 is greater than previously thought, refuting the single-individual hypothesis. Although seemingly unlikely, this denotes that two hominids represented by rarely found leg and foot elements both lost their left foot to crocodiles at nearby sites within a 6,000 year interval. We cannot determine if the hominids were preyed upon by crocodiles or mammalian carnivores. However, the carnivore damage to them and associated faunal remains suggests that high predation risk constrained hominid activities involving discard of the stone artifacts found at these sites. This finding is inconsistent with the interpretation of the sites as home bases or living floors.

Landscape distribution of Oldowan stone artifact assemblages across the fault compartments of the eastern Olduvai Lake Basin during early lowermost Bed II times.

The density and composition of Oldowan stone artifact assemblages deposited during the first ca. 20,000 years of lowermost Bed II times show a recurrent pattern of variation across recognized synsedimentary faults that compartmentalized landscapes of the eastern Olduvai Lake Basin. When active, the faults created minor topographic relief. The upthrown fault footwalls accumulated assemblages with relatively high densities of artifacts, including types retaining potential usefulness, particularly volcanic flaked pieces, manuports, pounded pieces, and split cobbles. Values for these assemblage characteristics decline toward the lower-lying hangingwall of the fault compartments, accompanied by an increase in the proportionate weight of artifact assemblages comprising quartzite, particularly flaking shatter and potentially useful detached pieces. Values reverse once again at faults, either on the downthrown, hangingwall side or on the upthrown side. The patterns are stronger for the volcanic components of the artifact assemblages than for the quartzite components, reflecting the additional influence of distance from the local source on quartzite assemblage characteristics reported previously. The landscape distributions of artifact assemblages are consistent with a landscape-fault model in which minor fault-induced topographic relief at times created a mosaic of vegetation environments repeated within each of the three fault compartments of the lake margin and distal alluvial fan. The fault-compartmentalized landscape model is currently supported only by sediment thickness and facies changes across synsedimentary faults, but it provides predictions for spatial variation in the cover abundance of trees, freshwater reservoirs and associated distributions of resources and hazards associated with stone artifact use and discard that can be tested if sample sizes of key paleoenvironmental indicators are increased.

Dental microwear texture analysis of hominins recovered by the Olduvai Landscape Paleoanthropology Project, 1995-2007.

Dental microwear analysis has proven to be a valuable tool for the reconstruction of aspects of diet in early hominins. That said, sample sizes for some groups are small, decreasing our confidence that results are representative of a given taxon and making it difficult to assess within-species variation. Here we present microwear texture data for several new specimens of Homo habilis and Paranthropus boisei from Olduvai Gorge, bringing sample sizes for these species in line with those published for most other early hominins. These data are added to those published to date, and microwear textures of the enlarged sample of H. habilis (n = 10) and P. boisei (n = 9) are compared with one another and with those of other early hominins. New results confirm that P. boisei does not have microwear patterns expected of a hard-object specialist. Further, the separate texture complexity analyses of early Homo species suggest that Homo erectus ate a broader range of foods, at least in terms of hardness, than did H. habilis, P. boisei, or the "gracile" australopiths studied. Finally, differences in scale of maximum complexity and perhaps textural fill volume between H. habilis and H. erectus are noted, suggesting further possible differences between these species in diet.

Validation of bone surface modification models for inferring fossil hominin and carnivore feeding interactions, with reapplication to FLK 22, Olduvai Gorge, Tanzania.

Resolving the issue of how Early Stone Age hominins acquired large mammal carcasses requires information on their feeding interactions with large carnivores. This ecological information and its behavioral and evolutionary implications are revealed most directly from the tooth, cut, and percussion marks on bone surfaces generated by hominin and carnivore feeding activities. This paper employs a bootstrap method, a form of random resampling with replacement, to refine published neotaphonomic models that use the assemblage-wide proportions of long bones bearing feeding traces to infer the sequences in which Plio-Pleistocene hominins and carnivores accessed flesh, marrow, and/or grease from carcasses. Results validate the sensitivity of the models for inferring hominin feeding ecology, which have been questioned on grounds shown here to be unfounded. The bootstrapped feeding trace models are applied to the late Pliocene larger mammal fossil assemblage from FLK 22 (Zinjanthropus site), Olduvai Gorge, Tanzania. High frequencies of tooth and percussion marking on long bone midshaft fragments from FLK 22 are most consistent with those feeding trace models that simulate hominin scavenging from carcasses defleshed by carnivores, while cut mark data indicate that hominins more often had access to upper forelimb flesh than upper hind limb flesh. Together, the bone surface modification data indicate that hominins typically gained secondary access to partially defleshed carnivore kills, but they also allow for the possibility of some carcasses being processed only by carnivores and only by hominins.

A new horned crocodile from the Plio-Pleistocene hominid sites at Olduvai Gorge, Tanzania.

BACKGROUND: The fossil record reveals surprising crocodile diversity in the Neogene of Africa, but relationships with their living relatives and the biogeographic origins of the modern African crocodylian fauna are poorly understood. A Plio-Pleistocene crocodile from Olduvai Gorge, Tanzania, represents a new extinct species and shows that high crocodylian diversity in Africa persisted after the Miocene. It had prominent triangular "horns" over the ears and a relatively deep snout, these resemble those of the recently extinct Malagasy crocodile Voay robustus, but the new species lacks features found among osteolaemines and shares derived similarities with living species of Crocodylus. METHODOLOGY/PRINCIPAL FINDINGS: The holotype consists of a partial skull and skeleton and was collected on the surface between two tuffs dated to approximately 1.84 million years (Ma), in the same interval near the type localities for the hominids Homo habilis and Australopithecus boisei. It was compared with previously-collected material from Olduvai Gorge referable to the same species. Phylogenetic analysis places the new form within or adjacent to crown Crocodylus. CONCLUSIONS/SIGNIFICANCE: The new crocodile species was the largest predator encountered by our ancestors at Olduvai Gorge, as indicated by hominid specimens preserving crocodile bite marks from these sites. The new species also reinforces the emerging view of high crocodylian diversity throughout the Neogene, and it represents one of the few extinct species referable to crown genus Crocodylus.

Taphonomic, avian, and small-vertebrate indicators of Ardipithecus ramidus habitat.

Thousands of vertebrate specimens were systematically collected from the stratigraphic interval containing Ardipithecus ramidus. The carcasses of larger mammals were heavily ravaged by carnivores. Nearly 10,000 small-mammal remains appear to be derived primarily from decomposed owl pellets. The rich avifauna includes at least 29 species, mostly nonaquatic forms. Modern analogs of the most abundant birds and of a variety of rodents are associated with mesic woodland environments distant from large water bodies. These findings support inferences from associated geological, isotopic, invertebrate, and large-vertebrate assemblages. The combined results suggest that Ar. ramidus occupied a wooded Pliocene habitat.

A diagnosis of crocodile feeding traces on larger mammal bone, with fossil examples from the Plio-Pleistocene Olduvai Basin, Tanzania.

Neotaphonomic studies have determined the patterns of bone damage created by larger mammalian carnivores when consuming mammalian carcasses. Typically, mammalian carnivores gnaw and break bones to various degrees in order to access marrow, grease, and brain tissue. In contrast, crocodiles attempt to swallow whole parts of mammal carcasses, inflicting in the process tooth marks and other feeding traces on some of the bones they are unable to ingest. Although crocodiles are major predators of larger mammals along the margins of protected tropical rivers and lakes, their feeding traces on bone have received little systematic attention in neotaphonomic research. We present diagnostic characteristics of Crocodylus niloticus damage to uningested mammal bones resulting from a series of controlled observations of captive crocodile feeding. The resulting bone assemblages are composed of primarily complete elements from articulating units, some of which bear an extremely high density of shallow to deep, transversely to obliquely oriented tooth scores over often large areas of the bone, along with shallow to deep pits and punctures. Some of the tooth marks (bisected pits and punctures, hook scores) have a distinctive morphology we have not observed to be produced by mammalian carnivores. The assemblages are also characterized by the retention of both low- and high-density bone portions, an absence of gross gnawing, and minimal fragmentation. Together, the damage characteristics associated with feeding by crocodiles are highly distinctive from those produced by mammalian carnivores. Modern surface bone assemblages along the Grumeti River in Tanzania's Serengeti National Park contain a mixture of specimens bearing damage characteristic of crocodiles and mammalian carnivores. Comparison of Plio-Pleistocene fossil bones from Olduvai Gorge, Tanzania, to bones damaged by captive and free-ranging Nile crocodiles reveals direct evidence of fossil crocodilian feeding from larger mammal bones associated with Oldowan stone artifacts.

Late Pliocene Homo and hominid land use from Western Olduvai Gorge, Tanzania.

Excavation in the previously little-explored western portion of Olduvai Gorge indicates that hominid land use of the eastern paleobasin extended at least episodically to the west. Finds included a dentally complete Homo maxilla (OH 65) with lower face, Oldowan stone artifacts, and butchery-marked bones dated to be between 1.84 and 1.79 million years old. The hominid shows strong affinities to the KNM ER 1470 cranium from Kenya (Homo rudolfensis), a morphotype previously unrecognized at Olduvai. ER 1470 and OH 65 can be accommodated in the H. habilis holotype, casting doubt on H. rudolfensis as a biologically valid taxon.